Molecular support for transferring the tropical African species of Gnidia to Lasiosiphon (Thymelaeaceae: Thymelaeoideae) and a worldwide synopsis of the species

Molecular support for transferring the tropical African species of Gnidia to Lasiosiphon (Thymelaeaceae: Thymelaeoideae) and a worldwide synopsis of the species | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Molecular support for transferring the tropical African species of Gnidia to Lasiosiphon (Thymelaeaceae: Thymelaeoideae) and a worldwide synopsis of the species Oluwayemisi Dorcas Olaniyan, J. Stephen Boatwright, Anthony R. Magee, and 2 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-3910184/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 08 Aug, 2024 Read the published version in Plant Systematics and Evolution → Version 1 posted 8 You are reading this latest preprint version Abstract Lasiosiphon Fresen. (Thymelaeaceae: Thymelaeoideae) is now understood to be a morphologically diverse genus distributed across southern and tropical Africa, Madagascar and India. The taxonomic history of Lasiosiphon has been marked by several revisions in its circumscription and status, particularly regarding its relationship with Gnidia L. Our phylogenetic analysis of nuclear (ITS) and plastid ( matK, rbcL, rps16 and trnL-F ) regions of an expanded sampling of the tropical African species of Gnidia provides compelling evidence for including all the tropical African species of Gnidia in Lasiosiphon . The morphological characters of these species are consistent with this conclusion. We accordingly provide 29 new combinations in Lasiosiphon for species of Gnidia that currently lack combinations in that genus. We also provide a comprehensive nomenclature of all currently recognized Lasiosiphon species at the regional level worldwide, as the framework for future taxonomic revisions in the genus. Arthrosolen Africa Asia Gnidia Lasiosiphon Nomenclature Phylogeny Taxonomy Figures Figure 1 Figure 2 1 Introduction The genus Lasiosiphon Fresen. (Thymelaeaceae: Thymelaeoideae) comprises a morphologically diverse group of plants distributed across southern and tropical Africa, Madagascar and India. The genus was established by Fresenius ( 1838 ) for the single species L. glaucus Fresen., based on its capitate inflorescence of pentamerous flowers, with five petaloid scales and ten stamens in two series. The circumscription of the genus subsequently underwent significant changes, primarily due to the increasing morphological and taxonomic complexity of the genus Gnidia . This resulted in uncertainties regarding the relationships and status between Gnidia and allied genera such as Englerodaphne Gilg, Arthrosolen C.A.Mey. and Lasiosiphon in the “ Gnidia group” (sensu Herber 2023). Conflicting views on the importance of floral merosity and the presence or absence of petaloid scales (Fig. 1 ) as generic distinctions led some taxonomists (Gilg 1894, 1921 ; Domke 1934 ; Peterson 1959 , 1978 ; Dyer 1975 ) to include Arthrosolen and Lasiosiphon in an expanded concept of Gnidia whereas others (Endlicher 1847 ; Meisner 1857; Bentham and Hooker 1880 ; Pearson 1910 ; Wright 1915 ; Marloth 1925 ; Burtt-Davy 1926 ; Phillips 1944 , 1951 ; Leandri 1950 ; Hutchinson 1967 ) retained them as separate genera. Notably, Phillips ( 1944 , 1951 ) radically altered the circumscription of Arthrosolen. Previously defined by the absence of petaloid scales, the genus was now defined by a capitate or capitulate inflorescence subtended by colourful, membranous bracts. This revised circumscription of the genus included mainly pentamerous species, mostly lacking petaloid scales (but rarely with petaloid scales). Phillips ( 1944 ) transferred the remaining tetramerous species that had previously been included in Arthrosolen as lacking petaloid scales into Gnidia. In this taxonomy, Lasiosiphon included only pentamerous species with petaloid scales, Arthrosolen comprised mainly pentamerous species without petaloid scales but also one tetramerous species ( Gnidia microcephala Meisn.) with petaloid scales, and Gnidia encompassed tetramerous species with or without petaloid scales. The debate regarding the taxonomy of these genera persisted until Herber ( 2002 , 2003 ) and Peterson ( 2006 ) advocated for a very broad generic circumscription of Gnidia to include both Arthrosolen and Lasiosiphon . They characterized their enlarged Gnidia as having double the number of stamens as the number of sepals, arranged in two whorls. Molecular phylogenetic studies have provided new and valuable insights into the evolutionary relationships among Gnidia and its allied genera (Van der Bank et al. 2002 ; Beaumont et al. 2009 ; Olaniyan 2021 ). These studies showed that Gnidia , in the broad sense of Herber ( 2002 , 2003 ) and Peterson ( 2006 ), is polyphyletic and that Lasiosiphon , along with Arthrosolen sensu Phillips ( 1944 , 1951 ), forms a distinct clade. This clade included all pentamerous species included in Gnidia s.l. plus various tetramerous taxa distinguishable by an involucrate inflorescence. Beaumont et al. ( 2009 ) thus argued for the reinstatement of the genus Lasiosiphon , with an expanded circumscription to include tetramerous species with or without scales. Subsequent authors (Manning and Boatwright 2013 ; Boatwright et al. 2017 ; Magee and Manning 2017 ) implemented the conclusions of Beaumont et al. ( 2009 ) by providing combinations for species of Gnidia from southern Africa and Madagascar that exhibit characteristics consistent with the newly circumscribed Lasiosiphon . These combinations have been widely accepted in the taxonomic community. However, a comparable treatment of the tropical African species remains to be implemented. Following Magee and Manning ( 2017 ), Lasiosiphon can be distinguished “by its pentamerous or sometimes tetramerous flowers in capitate, involucrate inflorescences subtended by foliaceous or papery bracts; a more or less pubescent hypanthium, with the basal part sometimes heavily bearded with long, straight, silky hairs; and waxy, bright yellow to orange or red sepal lobes, usually with petaloid scales in the mouth of the hypanthium tube but these lacking in some species.” Based on the original circumscription of Lasiosiphon and as confirmed by molecular phylogenetic analyses, all the pentamerous taxa in tropical Africa, some seven species, are readily accommodated in Lasiosiphon . As observed by Olaniyan ( 2021 ), however, the tetramerous Gnidia species in tropical Africa, which had not been included in previous phylogenetic analyses, shared the morphological traits consistent with the expanded circumscription of Lasiosiphon . This observation raises the likelihood that these species should be transferred to Lasiosiphon . All the Madagascan species are endemic to the island, and very few species are shared between tropical Africa and either southern Africa or Asia. These four regions thus represent convenient geographical divisions. The tetramerous species of tropical Africa can be segregated into three morphological groups based on the position and structure of their inflorescence: 1. Inflorescences umbellate, terminal and axillary, few-flowered, sessile; bracts foliaceous; pedicel non-hairy ( Gnidia fastigiata Rendle); 2. inflorescences capitate, terminal and/or axillary, few-flowered, usually distinctly long or shortly pedunculate; involucral bracts papyraceous or coloured; pedicel hairy (e.g. G. goetzeana Gilg); and 3. inflorescences composite-like heads, always terminal, many-flowered, distinctly long or shortly pedunculate; involucral bracts papyraceous or coloured; pedicel hairy (e.g. G. mollis C.H.Wright). This paper aims to: 1. expand sampling to include representative samples from each of these morphological groups in a phylogenetic analysis of Thymelaeaceae, allowing us to assess their relationships within the broader context of Gnidia lineages and determine their taxonomic placement in the classification proposed by Beaumont et al. ( 2009 ), Manning and Boatwright ( 2013 ), Magee and Manning ( 2017 ) and Boatwright et al. ( 2017 ); 2. provide any necessary nomenclatural combinations for the Gnidia species in tropical Africa that currently lack combinations in Lasiosiphon ; 3. transfer the three pentamerous Gnidia species from Asia to Lasiosiphon in accordance with the current circumscription of that genus; and 4. compile a comprehensive list of all Lasiosiphon species at regional level as a framework for future taxonomic work in the genus. 2 Material and Methods 2.1 Loci selection and taxon sampling New sequences of tropical African Gnidia were added to a reduced matrix of Olaniyan ( 2021 ) with combined data of five loci (ITS, matK, rbcL, rps16 , and trnL-F ) for phylogenetic analyses. Additionally, taxon sampling included representatives from all major lineages of Gnidia to ensure comprehensive coverage of its diversity. A total of 43 sequences for 13 taxa that were not included in previous studies were added to the pre-existing datasets. These include five newly sampled Gnidia species representing the distinct morphological groups found in tropical Africa. In all, 119 species from 24 genera in the subfamily Thymelaeoideae were analyzed, along with four taxa from the subfamily Octolepidoideae as outgroup taxa. Voucher specimens and GenBank accession numbers are listed in Appendix 1. 2.2 DNA extraction, amplification, and sequencing DNA was extracted from herbarium specimens using the 10X cetyltrimethylammonium bromide (CTAB) method of Doyle and Doyle ( 1987 ) with the addition of 2% polyvinylpyrrolidone (PVP) to prevent the interaction of polyphenolic compounds with the DNA samples. Polymerase Chain Reaction (PCR) amplification and sequencing were performed with parameters and amplification primers optimized for each locus based on previous studies: rbcL and trnL-F regions as in Van der Bank et al. ( 2002 ); ITS as in Beaumont et al. ( 2009 ); matK and rps16 regions as in Motsi et al. ( 2010 ). Geneious v.8.1.9 ( https://www.geneious.com ) was used for the editing and assembly of complementary strands. The sequences were aligned using Multiple Sequence Comparison by Log-Expectation [MUSCLE v3.8.31; (Edgar 2004 )] in the Geneious suite and visually checked for any alignment errors. 2.3 Phylogenetic analyses The nuclear (ITS) dataset, combined plastid ( matK, rbcL, rps16 , and trnL-F ) datasets, and combined (ITS + plastid) datasets were analyzed using the Maximum Parsimony (MP) algorithm of PAUP* version 4.0a168 (Swofford, 2003 ). Tree searches were conducted using a heuristic search with 1,000 random sequence additions and 10 trees held at each step. Tree bisection-reconnection (TBR) branch swapping and MULtrees were in effect. Delayed transformation character optimization (DELTRAN) was used for calculating branch lengths. Branch support was estimated using bootstrap analysis (Felsenstein 1985 ) implemented in PAUP* with 1,000 replicates, employing TBR branch swapping, with ten trees held at each step and random sequence addition. Only values of 50% and above are reported. The following scale was used to evaluate bootstrap support: 50–74%, weak; 75–89%, moderate; 90–100%, strong. The congruence of the individual dataset was assessed by comparing bootstrap percentages. The trees were considered incongruent only when they exhibited ‘hard’ (i.e. with strong bootstrap support) rather than ‘soft’ (i.e., with weak bootstrap support) incongruence (Seelanan et al. 1997 ; Wiens 1998 ). Bayesian inference (BI) analyses were carried out on the nuclear (ITS) dataset, combined plastid datasets, and combined (ITS + plastid) datasets using MrBayes on XSEDE v.3.2.7 (Ronquist et al. 2012 ) as implemented on the CyberInfrastructure for Phylogenetic Research (CIPRES) Science Gateway (Miller et al. 2010 ). Before the Bayesian analysis, the following best-fit model of evolution was determined for each matrix: ITS (TIM1 + I + G), rps16 (TVM + G), matK (TVM + G), rbcL (TVM + I + G), and trnL-F (TPM1uf + G). This was done with jModeltest 2.1.9 v20160115 (Darriba et al. 2012 ) which uses log-likelihood scores to estimate the model of DNA evolution that best suits a specific dataset. Four parallel Markov chain Monte Carlo (MCMC) estimations were run for 10,000,000 generations, with trees sampled every 1000 generations. The stationarity was checked using tracer v 1.7 (Rambaut et al. 2018 ), and all ESS values were found to be above 200 – indicating that the number of generations was sufficient. Suboptimal trees were discarded as the ‘burn-in” phase. The posterior probability values were evaluated using the following scale: below 0.95, weak; 0.95–1.0, strong. Maximum Likelihood (ML) analyses were performed on the nuclear (ITS) dataset, combined plastid datasets, and combined (ITS + plastid) datasets using RAxML-HPC2 on XSEDE (Stamatakis 2014 ) on the CIPRES Science Gateway (Miller et al. 2010 ). A single consensus tree was generated from 1,000 nonparametric bootstrap replicates. The following scale was used to evaluate the node support: 50–74%, weak; 75–89%, moderate; 90–100%, strong. 2.4 Taxonomic treatment For the list of Lasiosiphon species we followed the most recent regional treatments for each geographical region as follows: Southern Africa : Wright in Flora Capensis 5: 1–81 (1915), Manning and Boatwright in Bothalia 43(1): 84–88 (2013); Magee and Manning in Kew Bull. 78(1): 1–6 (2017), and Kolokoto et al. in Bothalia 49(1): 3–5 (2019). Tropical Africa : Pearson in Flora of Tropical Africa 6(1, 2): 212–255 (1910); Staner in Bulletin Jardine Botanique l’Etat. Bruxelles 13(4): 355 (1935); Aymonin in Flore du Gabon 2: 35–95 (1966); Peterson in Flora of Tropical East Africa: 1–37 (1978); Cuccuini in Flora of Somalia 1: 200–202(1993); Peterson in Flora of Ethiopia and Eritrea 2(1): 429–435 (2000); Peterson in Flora Zambesiaca 9(3): 85–117 (2006); Figueiredo and Smith in Strelitzia 22: 162–163 (2008); Madagascar : Rogers in Annals of the Missouri Botanical Garden 96: 324–368 (2009); Boatwright et al. in S. African J. Bot. 112: 336–337 (2017); Asia (Sri Lanka, India, and Arabia) : Gastaldo in Webbia 24(1): 337–389 (1969); Townsend in Revised Handbook of the Flora of Ceylon 2: 501–511 (1981); and Prabhukumar et al. in Phytotaxa 372(2): 180 (2018). 3 Results 3.1 Sequence characteristics A summary of statistics for all datasets is presented in Table S2 . Among the plastid regions, matK had the most variable sites (278; 36%) compared to rps16 (338; 35%), trnL-F (475; 34%) and rbcL (260; 19%). The percentage of parsimony informative characters was higher in matK (163; 21%) and rps16 (201; 21%) than in trnL-F (249; 18%), and rbcL (170; 12%). The ITS region had significantly more variable sites (446; 62%) and a higher number of parsimony informative sites (372; 52%) than any of the plastid regions. It also showed more site changes than the plastid regions (5.3 changes per variable site) reflecting that the ITS tree has a higher phylogenetic signal than the plastid regions. The plastid regions were not analyzed separately due to low sequence divergence but rather in combination. 3.1 Phylogenetic analyses The comparison of the topologies derived from the combined plastid (Fig. S1 ) and ITS dataset (Fig. S2 ) showed significant agreement in the delimitation of the segregate clades of Gnidia . Also, all the genera in the subfamily Thymelaeoideae, including Lasiosiphon , were consistently well supported in both analyses except for Lachnaea L., which did not receive support for its monophyly in both datasets. We combined the two datasets since there were no strongly supported incongruences in them. Additionally, the BI (Fig. S3), ML (Fig. S4) and MP (Fig. S5) analyses of the concatenated dataset produced similar topologies; hence, results and combined support values are displayed on the BI majority rule consensus tree (Fig. 2 ). There was strong support for the monophyly of the subfamily Thymelaeoideae [Bayesian posterior probability (PP) = 1.00, maximum likelihood bootstrap (MLB) = 100%, maximum parsimony bootstrap (MPB) = 98%] with Aquilaria Lam. and Gyrinops Gaertn sister to the rest of the Thymelaeoideae (PP = 100, MLB = 100, MPB = 97). Consistent with the findings of previous studies, Gnidia was polyphyletic in our study and dispersed as ten lineages (Lineages A-I) within Thymelaeoideae. Seven of the lineages (Lineages A-G) were strongly supported as monophyletic (PP = 1.00, MLB = 100, MPB = 100), while the remaining three lineages (Lineages H-J), which were mainly tropical African, were nested in Lasioisphon , rendering Lasiosiphon paraphyletic. There were two strongly supported subclades within Lasiosiphon. One of the lineages of Gnidia (Lineage H) was unresolved in a subclade (PP = 1.00, MLB = 100, MPB = 100) of pentamerous species, while the remaining two lineages (Lineages I-J) were also weakly nested within a subclade (PP = 1.00, MLB = 100, MPB = 100) of mainly tetramerous species. Other genera within the “ Gnidia group” (sensu Herber, 2003 ), viz. Passerina L. Peddiea Harv. ex Hook., Pimelea Banks & Sol. ex Gaertn., Stephanodaphne Baill., Struthiola L., Synaptolepis Oliv. and Wikstroemia were each inferred to be monophyletic with strong supports (Fig. 2 ). The sister groupings Drapetes Banks ex Lam. + Kelleria Endl. and Dais L. + Phaleria Jack was strongly supported in all three analyses (PP = 1.00, MLB = 100, MPB = 100), whereas Dirca L. + Ovidia (Poepp. & Endl.) Meisn. received strong support in the BI and ML (PP = 1.00, MLB = 92) analyses and weak support in the MP analysis (MPB = 76). Lachnaea was found to be paraphyletic with respect to a lineage of Gnidia (Lineage A). Support for the inclusion of this lineage was strong in the BI and ML (PP = 1.00, MLB = 97) analyses but moderate in the MP analysis (MPB = 79). The monophyly of Englerodaphne Gilg received strong support in the BI and ML (PP = 1.00, MLB = 96) analyses but moderate support in the MP analysis (MPB = 76). 4 Discussion This study corroborates and expands upon the conclusions of other research (Van der Bank et al. 2002 ; Beaumont et al. 2009 ; Olaniyan 2021 ) by uncovering the existence of numerous distinct evolutionary lineages within the genus Gnidia . This implies that the existing classification system may not reflect actual evolutionary relationships among Gnidia species, thereby necessitating a reassessment of their taxonomy. Prior researchers (Beaumont et al 2009 ; Foster et al 2016 ; Olaniyan 2021 ) have examined the possibilities of establishing Gnidia as a monophyletic group, using the fundamental principle of monophyly along with certain secondary principles, but have not reached a consensus. A comprehensive analysis of morphological data and the integration of ecological data may be necessary to completely execute the optimal choice. However, while the decision on the reclassification of Gnidia species is still pending, this study is a valuable contribution towards achieving a monophyletic classification for Gnidia . Consistent with the phylogenetic analyses of Olaniyan ( 2021 ), Gnidia fastigiata was resolved in one of the lineages of Gnidia that included mainly southern African species. G. fastigiata differs from other tropical African tetramerous species by its umbellate inflorescence (usually 2–3 flowered), leaf-like bracts, and non-hairy pedicel. This justifies its non-inclusion in the Lasiosiphon clade. Significantly, it is one of two tetramerous tropical African Gnidia species with a geographical range extending to southern Africa, the other being G. microcephala Meisn. [now Lasiosiphon microcephalus (Meisn.) J.C.Manning & Magee (Magee and Manning 2017 )], which is distinct in its capitate involucrate inflorescence and its colored modified bracts. Both nuclear and plastid data strongly support the inclusion of tropical African Gnidia species in Lasiosiphon . Despite less resolution and support among Lasiosiphon species, the analyses reveal two robustly supported subclades within the genus. The first subclade comprises pentamerous species with or without scales from southern Africa and Madagascar. Notably, G. chapmanii B.Peterson (Lineage H), with pentamerous flowers, is nested within this clade. The second subclade encompasses tetramerous species, with four tropical African species (Lineages I & J) represented in the clade. This subdivision hints at potential infrageneric groups within Lasiosiphon , suggesting that floral merosity could serve as an informative character for infrageneric classifications in the genus pending more comprehensive sampling. Lasiosiphon is characterized by its unique floral morphology of an involucrate and pedunculate inflorescence. In a comparative study on bract characteristics within Gnidia s.l., Beaumont et al. (2001) identified three categories: leaf-like bracts, partly modified bracts, and highly modified bracts. The purpose of establishing these categories was to classify the different types of bracts based on their similarity or dissimilarity to cauline leaves. The study revealed that all the Gnidia species except the ones formerly classified in Lasiosiphon are either lacking bracts or bearing unmodified leaf-like bracts. This suggests that the presence or absence of bracts, as well as their level of modification, can be used as a distinguishing characteristic within Gnidia s.l. For example, species of Englerodaphne that were previously put in the genus Gnidia were moved back to that genus because of their unique bract features of ebracteate inflorescence (Olaniyan 2020). In the same vein, a partly or highly modified bract highlights the distinctiveness of Lasiosiphon in the group. Interestingly, Gnidia species in the tropical Africa region (except G. fastigiata ) have an involucre of bracts that exhibit some degree of modification that is absent in other Gnidia lineages, which further supports their inclusion in the Lasiosiphon clade. This floral morphology is a key trait that sets Lasiosiphon apart from other lineages of Gnidia and helps in its identification. 5. Conclusion The polyphyletic nature of the genus Gnidia has been a topic of recent discussion. Previous research has shown the existence of multiple lineages within the genus, and this study supports that conclusion. Olaniyan ( 2021 ) sampled the full morphological diversity of Gnidia except the tropical African species, resulting in the discovery of additional Gnidia lineages. Her findings indicated that no additional lineages are expected, except for the tropical African species which were not adequately represented in her study. She proposed that these should be classified under the genus Lasiosiphon based on a review of critical literature that shed lights on their morphological characteristics. The sampling of species from tropical Africa confirmed that they do indeed belong to the Lasiosiphon clade, providing compelling evidence for the transfer to that genus of the tropical African species that remain in Gnidia . We, therefore, provide new generic combinations for the tropical African species of Gnidia that lack combinations in Lasiosiphon . 6 Taxonomic Treatment Lasiosiphon Fresen. in Flora 21(2): 602. 1838; Meisner in Candolle, Prodr. [A. P. de Candolle] 14(2): 593. 1857; Wright in Fl. Cap. (Harvey) 5(2, 1): 69. 1915; Pearson in Fl. Trop. Afr. 6(1, 2): 227. 1910; Domke in Biblioth. Bot. 27(111): 135. 1934; Phillips in Gen. S. Afr. Fl. Pl. (ed.2): 528. 1951. ≡ Gnidia subg. Lasiosiphon (Fresen) Engler, Veg. Erde [Engler] 9(3, 2): 634. 1921. ≡ Gnidia sect. Lasiosiphon (Fresen.) Staner in Bull. Jard. Bot. État Bruxelles 13(4): 355. 1935. Type— Lasiosiphon glaucus Fresen. Arthrosolen [unranked] Cephalodaphne C.A.Mey. in Bull. Cl. Phys.-Math. Acad. Imp. Sci. Saint-Pétersbourg 1: 359. 1843, syn. nov . ≡ Arthrosolen sect. Cephalodaphne (C.A.Mey) Tiegh in Bull. Soc. Bot. France 40: 75. 1893. —TYPE: Not designated [Species included: A. calocephalus (Meisn.) C.A.Mey., A. polycephalus (E.Mey. ex Meisn.) C.A.Mey., A. anthylloides (L.f.) C.A.Mey.]. Gnidia sect. Involucratae Gilg in Engler & Prantl., Nat. Pflanzenfam. 3(6a): 227. 1894, syn. nov. —LECTOTYPE ( designated here ): Lasiosiphon involucratus (Steud ex A.Rich) Olaniyan (≡ Gnidia involucrata Steud. ex A.Rich). Atemnosiphon Leandri in Not. Syst. 13: 44. 1947, syn. nov. —LECTOTYPE ( designated here ): Lasiosiphon coriaceus Leandri (≡ Atemnosiphon coriaceus (Leandri) Leandri). Arthrosolen sensu Phillips in J. S. African Bot. 10: 64. 1944, non Arthrosolen C.A.Mey. in Bull. Cl. Phys.-Math. Acad. Imp. Sci. Saint-Pétersbourg 1: 359. 1843. —TYPE: Arthrosolen polycephalus (E.Mey. ex Meisn.) C.A.Mey. [≡ Gnidia polycephala (E.Mey. ex Meisn.) Gilg]. Nomenclatural note : Meyer (1843: 359) designated the species Arthrosolen spicatus C.A.Mey. as the type of his new genus Arthrosolen but the typesetting on this page is slightly confusing and Phillips ( 1951 : 528) overlooked this, and in consequence lectotypified the genus against A. polycephalus (E.Mey. ex Meisn.) C.A.Mey. This lectotypification is nomenclaturally superfluous but has been generally accepted until now. Arthrosolen polycephalus falls within the circumscription of Lasiosiphon and as a result, the generic name Arthrosolen has been accepted as a synonym of Lasiosiphon . The designated type of the genus, A. spicatus , does not, however, fall within the morphological circumscription of Lasiosiphon and it has been retrieved as nested within one of the other lineages of Gnidia . The generic name Arthrosolen is therefore to be treated as applying to the lineage containing G. spicata and is not to be treated as a synonym of Lasiosiphon . Species from southern Africa : Lasiosiphon albosericeus (M.Moss ex B.Peterson) Olaniyan, comb. nov. ≡ Gnidia albosericea M.Moss ex B.Peterson in Bothalia 14(1): 81. 1982. ≡ Lasiosiphon ornatus Burtt Davy, Man. Pl. Transvaal 1: 206. 1926 as nom. nov. [non G, ornata (Meisn.) Gilg]. —TYPE: Swaziland, Hlatikulu, Dec 1910, M. Stewart 14 (Holotype: K Barcode K000322563 [web!]; Isotype: SAM Barcode SAM0002522-0 [web!]). Lasiosiphon anthylloides (L.f.) Meisn. in Prodr. [A.P. de Candolle] 14(2): 595. 1857. ≡ Passerina anthylloides L.f., Suppl. Pl.: 225. 1782. ≡ Arthrosolen anthylloides (L.f.) C.A.Mey. in Bull. Cl. Phys.-Math. Acad. Imp. Sci. Saint-Pétersbourg 1: 359. 1843. ≡ Gnidia anthylloides (L.f.) Gilg in Engler & Prantl., Nat. Pflanzenfam. 3(6a): 227. 1894. —TYPE: South Africa, Western cape, "Cap. bonai. spei.” [Cape of good hope], without date, Thunberg s.n. (Lectotype: UPS-THUNB 9566, designated by Rogers ( 2006 :486); isolectotypes: S barcode S-G-4573 [web!], SBT barcode SBT12559 [web!]). Lasiosiphon anthylloides var. vulgaris (Meisn.) Meisn. in Prodr. [A. P. de Candolle] 14(2): 596. 1857. ≡ Passerina anthylloides var. vulgaris Meisn. in Linnaea 14(5): 393. 1841. —TYPE: South Africa, “Hue spectat specim. Burchell (cat. No. 2203) in Herb. DC”, without date, Burchell 2203 (Holotype: G-DC [n.v.]). Other original material: South Africa, “Vanstadesberg, inter Zandplaat et Komga, usque ad alt. 3,000 ped., 27 Dec 1829, Drège 7347 (HBG barcode HBG512551 [web!], P barcode P00541076 [web!]); without date, Drège 4659 (NY barcode NY0386302 [web!]). Lasiosiphon anthylloides var. glabrescens (Meisn.) Meisn. in Prodr. [A.P. de Candolle] 14(2): 596. 1857. ≡ Passerina anthylloides var. glabrescens Meisn. in Linnaea 14(5): 393. 1841. —TYPE: South Africa, Kwazulu Natal, “Inter gramina ad sylvae marginem prope Port Natal, infra 200 ped. alt. (V,c, 34)", 8 Apr 1832, Drège s.n. (Syntypes: HAL barcode HAL0076215 [web!], P barcode P00648792 [web!]). Lasiosiphon anthylloides var. macrophyllus (Meisn.) Meisn. in Prodr. [A.P. de Candolle] 14(2): 596 (1857) [as macrophylla]. ≡ Passerina anthylloides var. macrophylla Meisn. in Linnaea 14(5): 393 (1841). —TYPE: South Africa, "Zwischen Omsamwubo und Omsamcaba, auf Grasfeldern, unter 1000 Fuss, (V,b,64)” 21 Feb 1832, Drège 4658a (Syntypes: HAL barcode HAL0076214 [web!], P barcode P00541075 [web!], NY barcode NY01288019 [web!]); “Zwischen Omtendo und Omsamculo, auf Sandhügeln unter 500 Fuss, (V,c,8)” Feb without year, Drège 4658b (Syntype: NY barcode NY00386301 [web!]). Lasiosiphon burchellii Meisn. in Prodr. [A.P. de Candolle] 14(2): 594 (1857). ≡ Gnidia burchellii (Meisn.) Gilg in Engler & Prantl., Nat. Pflanzenfam. 3(6): 227 (1894). ≡ Gnidia burchellii var. villosus Meisn. in Prodr. [A.P. de Candolle] 14(2): 594 (1857), nom. illeg. Superfl. —TYPE: South Africa “between Matlowing River and Takun”, 13 Jul 1812, Burchell 2203 (Syntypes: G barcode G00015757 [web!], G-DC barcode G00131382 [web!], GH barcode GH00443740 [web!], GOET barcode GOET002835 [web!], GRA barcode GRA0027039-0 [web!], K barcodes K000324212 [web!] and K000324211 [web!], M barcode M0106537 [web!], NY barcodes NY01287757 [web!] and NY01287758 [web!], P barcode P00541085 [web!], PRE barcodes PRE0587576-0 [web!] and PRE0307854-0 [web!], S barcode S11-24304 [web!]); South Africa, “Ad pedem montium Stormbergen”, Sept without year, Ecklon & Zeyher s.n. (Syntypes: W barcodes W1889-0280784 [web!] and W1889-0280828 [web!]). Lasiosiphon caffra Meisn. in Prodr. [A. P. de Candolle] 14(2): 593 (1857) [as Lasiosiphon caffer ]. ≡ Gnidia caffra (Meisn.) Gilg in Veg. Erde [Engler] 9(3, 2): 634 (1921). —TYPE: South Africa, Gauteng, “In Caffrariae monte Magalisberg”, without date, Zeyher 1488 (Holotype: NY01288012 [web!], isotypes: K barcodes K000324205 [web!] and K000324206 [web!]). Lasiosiphon calocephalus (Meisn.) Domke in Biblioth. Bot. 27(111): 95. 1934. ≡ Passerina calocephala Meisn. in Linnaea 14(5): 393. 1841. ≡ Arthrosolen calocephalus (Meisn.) C.A.Mey. in Bull. Cl. Phys.-Math. Acad. Imp. Sci. Saint-Pétersbourg 1: 359. 1843. ≡ Gnidia calocephala (Meisn.) Gilg in Engler & Prantl., Nat. Pflanzenfam. 3(6a): 228. 1894. —TYPE: South Africa, “inter Omsamwubo et magnam cataractam alt. 1000–1500 ped. (V. b.)”, 24 Feb 1832, Drège 4664a (Syntypes: NY barcode NY00386305 [web!], P barcode P00648740 [web!]); “in collibus inter Omcomas et Port Natal, infra 200 ped. alt. (V. c.)", 24 Apr 1832, Drège 4664b (Syntypes: HAL barcode HAL0111338 [web!], HBG barcode HBG512552 [web!], NY barcode NY00386304 [web!], P barcode P00648739 [web!], S barcode S11-21273 [web!]). Lasiosiphon canoargenteus C.H.Wright in Fl. Cap. (Harvey) 5(2, 1): 70 (1915) [as Lasiosiphon canoargentea ]. ≡ Gnidia canoargentea (C.H.Wright) Gilg in Veg. Erde [Engler] 9(3, 2): 634 1921. —TYPE: Transvaal, Lydenburg District, on the sides of mountains near Lydenburg, Jun 1874, McLea sub H. Bolus 3020 (Lectotype: K barcode K000324203 [web!], designated by Burt Davy (1926: 207); SAM barcode SAM0001519-0 [web!]). Lasiosiphon capitatus (L.f.) Burtt Davy in Man. Fl. Pl. Transvaal [Burtt Davy] 1: 207. 1926. ≡ Gnidia capitata L.f., Suppl. Pl.: 224. 1782. —TYPE: South Africa, “Habitat in Cap. bonae spei” without date, Smith 688.1 [Lectotype: LINN-SM, designated by Rogers ( 2006 :486)]. Lasiosiphon deserticola (Gilg) C.H.Wright in Fl. Cap. 5(2, 1): 71. 1915. ≡ Gnidia deserticola Gilg in Bot. Jahrb. Syst. 19(2–3): 263. 1894. ≡ Arthrosolen deserticola (Gilg) Compton in Trans. Roy. Soc. South Africa 19(3): 300. 193. —TYPE: South Africa, Carroogebiet, Hantam-Gebirge, without date, Meyer s.n. (Syntype: B †); Grootsriviers, Carroo" Mar 1819, Mundt & Maire s.n (Syntypes: B†, A barcode A00135116 [web!]). Lasiosiphon dregeanus (Meisn.) Endl., Gen. Pl. [Endlicher] Suppl. 4(2): 67. 1848. ≡ Gnidia dregeana Meisn. in Linnaea 14(5): 426. 1841. —TYPE: South Africa, Prope Port Natal (V. c.), Apr 1832, Drège 4661 (Holotype: NY barcode NY01287771 [web!]; isotype: P barcode P00541064 [web!). Lasiosiphon esterhuyseniae Magee & J.C.Manning in Bothalia 72(1): 2 (2017). —TYPE: South Africa, Northern Cape, Hay Div., Floradale, Apr 1940, E. Esterhuysen 2315 (Holotype: NBG!) Lasiosiphon kraussianus (Meisn.) Burtt Davy, Man. Fl. Pl. Transvaal [Burtt Davy] 1: 207 (1926). ≡ Gnidia kraussiana Meisn. in London J. Bot. 2: 552 (err. typ. 452) (1843). ≡ Lasiosiphon kraussii Meisn. in Candolle, Prodr. [A. P. de Candolle] 14(2): 596 (1857), nom. illeg. superfl. pro Gnidia kraussiana Meisn. (1843). —Type: South Africa, KwaZulu-Natal, "Ad latera montium tabularium prope Port Natal (V.c.)”, Sept 1839, Krauss 455 (Holotype: NY barcode NY01287756 [web!], isotypes: FI barcode FI000346 [web!], K barcodes K000324197 [web!] and K000324198 [web!], P barcode P00541092 [web!]). Nomenclatural note : Gnidia kraussiana Meisn. (1843) was described based on a specimen collected by the German scientist and traveler C.F. Krauss. Meisner (1857) subsequently changed the epithet from the adjectival form kraussiana to the genitive form kraussii when he transferred the species to Lasiosiphon , citing Gnidia kraussiana Meisn. in synonymy. This is not a correctable error (Turland et al., 2018: ICN Art. 60.1), and the name Lasiosiphon kraussii must therefore be regarded as an illegitimate superfluous name for Gnidia kraussiana (Turland et al., 2018: ICN Art. 52.1). The valid transfer of the name Gnidia kraussiana to Lasiosiphon must date from the first publication of the combination Lasiosiphon kraussianus by Burtt-Davy ( 1926 ). Lasiosiphon kuntzei (Gilg) R.Kolokoto & Magee in Bothalia 49(1): 3. 2019. ≡ Gnidia kuntzei Gilg in Revis. Gen. Pl. 3(3): 280. 1898. —TYPE: South Africa, Eastern Cape, “Middelburg Road, Capland”, 17 Feb 1894, Kuntze s.n . (Holotype: NY barcode NY01287798 [web!]; isotypes: GH barcode GH00443746 [web!], US barcode US1269408 [web!], Z barcode Z28526 [web!]). Lasiosiphon macropetalus (Meisn.) Meisn. in Prodr. [A. P. de Candolle] 14(2): 594. 1857. ≡ Gnidia macropetala Meisn. in London J. Bot. 2: 553 (err. typ. 453). 1843. —TYPE: South Africa, “in graminosis ad sylvarum margines ubique circa Port Natal (V.c.)”, 1 Jun 1839, Krauss 237 (Holotype: NY01287796 [web!]; isotypes: G barcode G00015749 [web!], FI barcode FI000345 [web!], K barcode K000324187 [web!], M barcode M0106560 [web!], P barcode P00648778 [web!], PRE barcode PRE0307855-0 [web!], TUB barcodes TUB002871 [web!] and TUB002872 [web!]). Lasiosiphon meisnerianus Endl., Gen. Pl. [Endlicher] Suppl. 4(2): 67. 1848. ≡ Gnidia cuneata Meisn. in Linnaea 14(5): 427. 1841, as nom. nov. [non Lasiosiphon cuneatus Decne (1844)]. —TYPE: South Africa, “Cap de Bonne Espérence” [Cape of good hope], 1828, Zehyer 86 (Holotype: MPU [n.v.], isotypes: G barcode G00190999 [web!], M barcode M0145888 [web!], P barcode P00541094 [web!]). Lasiosiphon meisnerianus var. angustifolius (Meisn.) Meisn. in Prodr. [A.P. de Candolle] 14(2): 594. 1857. ≡ Gnidia cuneata var. angustifolia Meisn. in Linnaea 14(5): 427. 184. —TYPE: South Africa, "In collibus lapidosis ad Klein Vischrivier, alt. 2000 ped. (II.a.)", 13 Oct 1829, Drège 2336 , (Syntypes: HAL barcode HAL0111685 [web!], P barcode P00541077 [web!]). Lasiosiphon meisnerianus var. spathulatus (Meisn.) Meisn. in Prodr. [A.P. de Candolle] 14(2): 594. 1857. ≡ Gnidia cuneata var. spathulate Meisn. in Linnaea 14(5): 427. 1841. —TYPE: South Africa, "In collibus ad Vischrivier prope Trompetersdrift, alt. 600–800 ped. (V. a.)", without date, Drège 4657 (Syntype: NY barcode NY01287755 [web!]); Drège 7351 (Syntype: P [n.v.]). Lasiosiphon microcephalus (Meisn.) J.C.Manning & Magee in Kew Bulletin 78(1): 12. 2017. ≡ Gnidia microcephala Meisn. in Prodr. [A.P. de Candolle] 14(2): 589. 1857. ≡ Arthrosolen microcephalus (Meisn.) E.Phillips in J. S. African Bot. 10: 63. 1944, nom. illeg. , non A. microcephalus S.Moore in J. Bot. 57: 116. 1919. —TYPE: South Africa, Gauteng, “monte Magalisberg”, without date, Zeyher 1492 (Syntypes: G barcodes G00165832 [web!] and G00190996 [web!], G-DC barcodes GDC00162612/1 [web!] and GDC00162612/2 [web!], GRA barcode GRA0027036 [web!], K barcode K000322515 [web!], P barcodes P00541080 [web!] and P00541081 [web!], P00541079 [web!], S barcode S11-24462 [web!], SAM [web!]). Lasiosiphon microphyllus (Meisn.) Meisn. in Prodr. [A.P. de Candolle] 14: 593 (1857). ≡ Gnidia microphylla Meisn. in Linnaea 14(5): 432. 184. —TYPE: South Africa, Northern Cape, “in collibus siccis prope Kuigunjels ad fl. Garip, alt. 200 ped.” [dry hills near Kuigunjels, near the mouth of the Orange River, 200 feet], without date, Drège 2976a (Holotype: NY barcode NY01288010 [web!]). Lasiosiphon pedunculatus (Beyers) J.C.Manning & Boatwr. in Bothalia 43(1): 88. 2013. ≡ Gnidia pedunculata Beyers in Bothalia 32: 79. 2002. —TYPE: South Africa, Western Cape, Knersvlakte, Olifants River Settlement 316, near eastern border, 6 Aug 1993, Le Roux & Hilton-Taylor 27 (Holotype: NBG barcode NBG0199291-0 [web!]; isotypes: K barcode K000322601 [web!], PRE barcode PRE0569222-0 [web!]). Lasiosiphon polyanthus (Gilg) C.H.Wright in Fl. Cap. (Harvey) 5(2, 1): 74. 1915. ≡ Gnidia polyantha Gilg in Bot. Jahrb. Syst. 19(2): 265. 1894. —TYPE: South Africa, Eastern Cape, “Saddle between Iggakancu & Bazeia Mountains”, Jun without year, Baur 646 (Holotype: K✝; isotypes: K barcodes K000324183 [web!] and K000324184 [web!]). Lasiosiphon polyanthus var. microcalyx Burtt Davy, Man. Pl. Transvaal [Burtt Davy] 1: 206. 1926. —TYPE: South Africa, "Pietersberg Distr., Marovuni, infrequens", without date, Junod 145 (Holotype: K [n.v.]). Lasiosiphon polycephalus (E.Mey. ex Meisn.) H.Pearson in Fl. Trop. Afr. 6(1, 2): 228. 1910. ≡ Passerina polycephala E.Mey. ex Meisn. in Linnaea 14(5): 390. 1841. ≡ Arthrosolen polycephalus (E.Mey. ex Meisn.) C.A.Mey. in Bull. Cl. Phys.-Math. Acad. Imp. Sci. Saint-Pétersbourg 1: 359. 1843. ≡ Gnidia polycephala (E.Mey. ex Meisn.) Gilg in Engler & Prantl., Nat. Pflanzenfam. 3(6a): 227. 1894. —TYPE: South Africa, “Locis depressis argillosis inter Poortje et Nieuwkerkshoogte, alt. 4000–5000 ped. (I.c.)” 10 Sept 1829, Drège 724 (Syntypes: HAL barcode HAL0111341 [web!], HBG barcode HBG512593 [web!], HEID barcode HEID702658 [web!], JE barcode JE00019704 [web!], NY barcode NY01104569 [web!], P barcode TUB barcode TUB002886 [web!], W barcode W 1889-0282097 [web!]); “Nieuweveld, alt. 3000–4000 ped. (l.d.)”, 24 Oct 1826, Drège s.n. (Syntypes: HAL barcode HAL0111340 [web!], HBG barcode HBG512653 [web!], MO barcode MO2302117 [web!], P00648737 [web!]); “Circa Limoenfontein in Winterveld, alt. 3000–3500 ped. (I.d.)”, without date, Burchell 2222 (Syntypes: G-DC [n.v.], NY barcode NY01104568 [web!], W barcode W1889-0282096 [web!]). Lasiosiphon pulchellus (Meisn.) Decne. in Voy. Inde [Jacquemont] 4: 149. 1844. ≡ Gnidia pulchella Meisn. in Linnaea 14(5): 425. 1841. ≡ Gnidia pulchella var. glabrata (Meisn.) Meisn. in Linnaea 14(5): 425. 1841, nom. illeg. Superfl . —Type: South Africa, "Ad rivulum inter Omtata et Omgaziana, alt. 800 ped. (V.b.), without date, Drège s.n. (Syntype: NY [n.v.]); "Katberg, alt. 4000–4500 ped. (I.a.)", without date, Drège 7354 (Syntype: NY barcode NY01287790 [web!]). Lasiosiphon pulchellus var. dasyphyllus (Meisn.) Meisn. in Prodr. [A.P. de Candolle] 14(2): 595. 1857. ≡ Gnidia pulchella var. dasyphylla Meisn. in Linnaea 14(5): 425. 184. —TYPE: South Africa, "in collibus siccis prope Zwartbulletje, 2500 ped.", without date, Drège 2976b (Syntype: NY barcode NY01288029 [web!]); “ad Zondagrivier prope Blaauwekrans, alt. 1700 ped. (II. c.)” without date, Drège s.n. (Syntype: NY [n.v.]). Lasiosiphon rigidus J.C.Manning & Boatwr. in Bothalia 43(1): 85. 2013. —TYPE: South Africa, Northern Cape, Tankwa [Tanqua] Karoo National Park, SW foot of Leeuberg, along drainage lines, 20 Jun 2012, Manning 3363 (Holotype: NBG barcode NBG0279875-0 [web!]; isotypes: K [n.v.], MO [n.v.], PRE!). Lasiosiphon robustus (B.Peterson) Olaniyan comb. nov. ≡ Gnidia robusta B.Peterson in Bothalia 14: 82. 1982. ≡ Lasiosiphon nanus Burtt Davy, Man. Pl. Transvaal 1: 207. 1926 as nom. nov. [non G. nana (L. f.) Wikstr.] —TYPE: Swaziland, Hlatikulu, without date., M. Stewart s.n . (Holotype: K barcode K000322535 [web!]). Lasiosiphon rubescens (B.Peterson) J.C.Manning & Magee in Kew Bull. 78(1): 12. 2017. ≡ Gnidia rubescens B.Peterson in Bot. Not. 119(2): 345. 1966. —TYPE: South Africa, Limpopo, “Transvaal - Near Mara, about 18 miles from Louis Trichardt on Mara-Vivo road”, 3 Apr 1957, Meeuse 10196 (Holotype: LD barcode LD1001442 [web!]; isotypes: BOL barcode BOL136956 [web!], BR barcode BR0000006249892 [web!], EA barcode EA000002189 [web!], G barcode G00165800 [web!], GB barcodes GB0048776 [web!] and GB0048777 [web!], LMA barcode LMA0031290-0 [web!], M barcode M0106561 [web!], MO barcode MO313015 [web!], PRE barcodes PRE0587577-0 [web!] and PRE0297723-0 [web!], S barcode S05-9781 [web!]). Lasiosiphon sericocephalus (Meisn.) J.C.Manning & Boatwr. in Bothalia 43(1): 88. 2013. ≡ Arthrosolen sericocephalus Meisn. in Prodr. [A.P. de Candolle]14(2): 561. 1857. ≡ Gnidia sericocephala (Meisn.) Gilg ex Engl., Veg. Erde [Engler] 9(3, 2): 634. 192. —TYPE: South Africa, “In mont. Macalisberg Caffrariae”, Zeyher 1494 (Holotype: G-DC; Isotype: K barcode K000324226 [web!]). Lasiosiphon splendens (Meisn.) Endl., Gen. Pl. [Endlicher] Suppl. 4(2): 67. 1848. ≡ Gnidia splendens Meisn. in Linnaea 14(5): 428. 1841. —TYPE: South Africa, Kwazulu Natal, "In herbosis prope Port Natal, infra alt. 100 ped. (V. c. 33)", without date, Drège 4656 (Holotype: NY barcode NY01287785 [web!]; isotypes: HAL barcode HAL0111548 [web!], HBG barcode HBG512608 [web!], K barcode K000324192 [web!], P barcode P00541044 [web!]). Lasiosiphon suavissimus (Dinter) Domke in Biblioth. Bot. 27(111): 91. 1934. ≡ Gnidia suavissima Dinter in Repert. Spec. Nov. Regni Veg. 16: 340. 1920.—TYPE: Namibia, "Namaland: Zwischen Angras Juntas und Buntfeldschuh", Jul 1913, Schäfer 504 (Lectotype: GLM barcode GLM101451 [web!], designated by Otte et al. (2011:258); isolectotype: B barcode B 10 0349157 [web!]). Lasiosiphon triplinervis (Meisn.) Decne. in Voy. Inde [Jacquemont] 4: 149. 1844. ≡ Gnidia triplinervis Meisn. in Linnaea 14(5): 429. 184. —TYPE: "Inter frutices in saxosa ripa fl. Omtendo infra alt. 100 ped. (V. c.)”, without date, Drège 4662 (Holotype: NY barcode NY01287803 [web!]; isotype: HAL barcode HAL0111551 [web!], TUB barcodes TUB002883 [web!] and TUB002883 [web!]). Species from Madagascar : Lasiosiphon ambondrombensis Boiteau in Bull. Acad. Malgache, n.s., 24: 83. 1941 [as ambondrombense]. ≡ Gnidia ambondrombensis (Boiteau) Z.S.Rogers in Ann. Miss. Bot. Gard. 96(1): 339. 2009. —TYPE: Madagascar, Fianarantsoa, Mt. Ambondrombe, rocky summit, 1900 m, 11 Apr 1941, P. Boiteau (Hb. Jard. Bot. Tananarive) 4643 (Holotype: P barcode P00370331 [web!], isotypes: MO barcode MO1181707 [web!], TAN). Lasiosiphon bojerianus Decne., Voy. Inde [Jacquemont] 4: 149. 1844. ≡ Gnidia bojeriana (Decne.) Gilg in Engler & Prantl., Nat. Pflanzenfam. 3(6a): 228. 1894. —TYPE: Madagascar, Antananarivo, Emirnae mtns., without date, W. Bojer s.n. (Lectotype: P barcode P00370315 [web!], designated by Rogers ( 2009 :342); isolectotypes: BM [n.v.], G barcode G00131578 [web!], P barcodes P00370314 [web!] and P00370316 [web!]). Lasiosiphon danguyanus (Leandri) Boatwr. & J.C.Manning in S. African J. Bot. 112: 337. 2017. ≡ Gnidia danguyana Leandri, Bull. Soc. Bot. France 77: 35 (1930). —TYPE: Madagascar, Toamasina, Tampina forest, Dec 1923, M. Louvel 118 (Lectotype: P barcode P00380376 [web!], designated by Rogers ( 2009 :342). Lasiosiphon daphnifolius (L.f.) Boatwr. & J.C.Manning in S. African J. Bot. 112: 337. 2017. ≡ Gnidia daphnifolia L.f., Suppl. Pl. 225 (1782) [as daphnaefolia]. ≡ Gnidia daphnifolia var. glabra L.f., Suppl. Pl. 225 (1782). ≡ Dessenia daphnifolia (L.f.) Raf. in Fl. Tellur. 4: 106. 1838 [as daphnefolia]. —TYPE: Madagascar, Herb. Smith 688.5 (Lectotype: LINN-SM, designated by Rogers ( 2006 :486). Lasiosiphon gilbertae (Drake) Boatwr. & J.C.Manning in S. African J. Bot. 112: 337. 2017. ≡ Gnidia gilbertae Drake in Bull. Mens. Soc. Linn. Paris 2: 1218. 1896. —TYPE: Madagascar, Mahajanga, ‘Madounga et Antsalahanki’, 1876, A. Grandidier s.n . (Holotype: P barcode P00380410 [web!]). Lasiosiphon gnidioides (Baker) Boatwr. & J.C.Manning in S. African J. Bot. 112: 337. 2017. ≡ Dais gnidioides Baker in J. Linn. Soc. Bot 20: 244 (1883). ≡ Arthrosolen gnidioides (Baker) Leandri in Bull. Soc. Bot. France 76: 1043 (1929). ≡ Gnidia gnidioides (Baker) Domke in Biblioth. Bot 27(111): 46 (1934). —TYPE: Madagascar, Antananarivo, ‘grassy hills of the province of Imerina’, without date, R. Baron 2061 (Lectotype: K barcode K000324220 [web!], designated by Rogers ( 2009 :351); isolectotype: P00380432 [web!]). Lasiosiphon hibbertioides S.Moore in J. Bot. 58: 189 (1920). ≡ Gnidia hibbertioides (S.Moore) Z.S.Rogers in Ann. Miss. Bot. Gard. 96(1): 349. 2009. —TYPE: Madagascar, without date, J. Thompson & J. Forbes s.n . (Holotype: BM barcode BM000647723 [web!]). Lasiosiphon humbertii Leandri in Bull. Soc. Bot. France 76: 1039. 1929. ≡ Gnidia humbertii (Leandri) Z.S.Rogers in Ann. Miss. Bot. Gard. 96(1): 351. 2009. —TYPE: Madagascar, Fianarantsoa, ‘Isalo, mouth of Sakamarekely & Sambalinieto rivers, 500–1000 m, 19 Oct 1924, H. Humbert 2844 (Lectotype: P barcode P00373493 [web!], designated by Rogers ( 2009 :351); isolectotype: G barcode G00010149 [web!]). Lasiosiphon leandrianus Boatwr. & J.C.Manning in S. African J. Bot. 112: 337. 2017. ≡ Gnidia decaryana Leandri in Bull. Mus. Natl. Hist. Nat. sér. 2(1): 436 (1929) as nom. nov. [non Lasiosiphon decaryi Leandri]. —TYPE: Madagascar, Toliara, ‘Fort-Dauphin’, 3 Jul 1926, R. Decary 4332 (Holotype: P barcode P00373426 [web!], isotypes: P barcode P00380392, [web!] TAN [n.v.]). Lasiosiphon linearis Leandri in Bull. Soc. Bot. France 76: 1040. 1930. ≡ Lasiosiphon decaryi var. linearis (Leandri) Leandri in Bull. Mus. Natl. Hist. Nat. sér. 2(3): 154. 1931. ≡ Gnidia linearis (Leandri) Z.S.Rogers in Ann. Miss. Bot. Gard. 96(1): 353. 2009. —TYPE: Madagascar, [Toliara/Fianarantsoa], savanna betw. Bemketa [Bereketa] & Malio, 15 Jun 1923, H. Poisson 692 (Holotype: P barcode P00373448 [web!]). Lasiosiphon occidentalis Leandri in Notul. Syst. (Paris) 13: 47. 1947. ≡ Gnidia occidentalis (Leandri) Z.S.Rogers in Ann. Miss. Bot. Gard. 96(1): 358. 2009. —TYPE: Madagascar, Mahajanga, Kamakama forest, Ankara plateau, 14 Jul 1901, H. Perrier de la Bâthie 1276 (Lectotype: P barcode P00373473 [web!], designated by Rogers ( 2009 :358). Lasiosiphon perrieri Leandri in Notul. Syst. (Paris) 13: 49. 1947. ≡ Gnidia perrieri (Leandri) Z.S.Rogers in Ann. Miss. Bot. Gard. 96(1): 360. 2009. —TYPE: Madagascar, Fianarantsoa, Andringitra Massif (Iratsy), valley of Riambava & Antsifotra, 2000–2500 m, 27 Nov 1924, H. Humbert 3827 (Lectotype: P barcode P00380352 [web!], designated by Rogers ( 2009 :360); isolectotypes: BM barcode BM000812676 [web!], K barcode K000634424 [web!], G barcode G00010144 [web!], K barcode K000634424 [web!], MO barcode MO1181820, TAN barcode TAN000732 [web!], US barcode US00956206 [web!]). Lasiosiphon razakamalalanus (Z.S.Rogers) Boatwr. & J.C.Manning in S. African J. Bot. 112: 337. 2017. ≡ Gnidia razakamalalana Z.S.Rogers in Adansonia sér. 3(28): 156. 2006. —TYPE: Madagascar, Toliara, Fivondronona Fort-Dauphin, Ivohibe Forest, 112 m, 29 Nov 2005, R. Razakamalala, E. Ramisy and B. Mara 2670 (Holotype: MO barcode MO1287062 [web!], isotypes: G barcode G00424957 [web!], K barcode K001089638 [web!], P barcode P02141588 [web!], TAN). Species from Asia (Arabia, India and Sri Lanka) : Lasiosiphon glaucus Fresen. in Flora 21(2): 603. 1838. ≡ Gnidia glauca (Fresen) Gilg in Engler. & Prantl., Nat. Pflanzenfam. 3(6a): 227. 1894. —TYPE: Ethiopia, “Abyssinia”, Rueppel s.n . (Holotype: FR barcode FR0030066 [web!]). Lasiosiphon sisparensis (Gardner) Meisn. in DC. Prodr. [A. P. de Candolle] 14(2): 598. 1857. ≡ Lasiosiphon eriocephalus var. sisparensis (Gardner) Gamble in Fl. Madras 2(7): 1244. 1925. ≡ Gnidia sisparensis Gardner in J. Nat. Hist. 7: 457. 1847. ≡ Gnidia glauca var. sisparensis (Gardner) Kumari in Fl. Tamil Nadu 1(2): 214. 1987. — TYPE: India, Tamil Nadu: Neilgherry (Nilgiri) mountains, Sispara, without date, Gardner s.n. (Lectotype: K barcode K000357981 [web!], designated by Prabhukumar et al. ( 2018 :180); isolectotype: K barcode K000357982 [web!]). Lasiosiphon socotranus Balf.f. in Proc. Roy. Soc. Edinburgh 12(113): 92. 1884. ≡ Gnidia socotrana (Balf.f.) Gilg in Engler & Prantl., Nat. Pflanzenfam. 3(6a): 228. 1894. —TYPE: Yemen, Socotra, Feb-Mar 1880, Balfour 518 (Lectotype: K barcode K000322641 [web!], designated by Gastaldo ( 1969 :369); isolectotypes: BM barcode BM000910837 [web!], EA barcode E00239397 [web!], E barcodes E00239400 [web!] and E00239402 [web!]). Lasiosiphon somalensis (Franch) H.Pearson in Fl. Trop. Afr. 6(1, 2): 229. 1910. ≡ Arthrosolen somalensis Franch. in Sert. Somal.: 62 (t. 6). 1882. ≡ Gnidia somalensis (Franch) Gilg in Ann. 1st. Bot. Roma, 6: 98. 1897. —TYPE: Somalia, “Ouanentab, plateau de Yaffer”, without date, Révoil 123 (Holotype: P barcode P00541095 [web!]). Species from tropical Africa Lasiosiphon apiculatus (Oliv.) Olaniyan, comb. nov. ≡ Gnidia involucrata var. apiculata Oliv. in Trans. Linn. Soc. London 29(3): 143, pl. 91. 1875. ≡ Gnidia apiculata (Oliv.) Gilg in Bot. Jahrb. Syst. 19(2–3): 263. 1894. —TYPE: Sudan, “Bare heights of Madi”, Dec 1862, Speke & Grant s.n . (Holotype: K barcode K000322646 [web!]). Lasiosiphon apiculatus fo. pyramidalis (Aymonin) Olaniyan, comb. nov. ≡ Gnidia apiculata fo. pyramidalis Aymonin in Fl. Gabon 11: 95, pl. 14, f. 6. 1966. —TYPE: Central African Republic, Oubangui, 60 Km North of Bambari, 5 Oct 1921, Tisserant 307 (Holotype: BM barcode BM000910854 [web!], isotype: P barcode P00541078 [web!]) Lasiosiphon bambutanus (Gilg & Ledermann ex Engl.) Olaniyan, comb. nov. ≡ Gnidia bambutana Gilg & Ledermann ex Engl. in Veg., Erde [Engler] 9(3, 2): 631. 1921. —TYPE: Cameroon “In den Babutubergen an der Grenze der sudansichen Parksteppenprovinz”, without date, Lederman s.n (Holotype: B†); Cameroon, Bangou [Bangan], Dec 1938, H. Jacques-Félix 2855 (Neotype: P00541086 [web!], designated by Aymonin ( 1965 :324). Lasiosiphon baumianus (Gilg) Olaniyan, comb.nov. ≡ Gnidia baumiana Gilg in Kunene-Sambesi-Exped.: 311. 1903. —TYPE: Angola, “Chitanda bei Kasinga”, Oct 1899, Baum 211 (Holotype: B†, isotypes: G barcode G00165820 [web!], HBG barcode HBG512557 [web!], M barcode M0106538 [web!], S barcodes S10-25321 [web!] and S10-25326 [web!], W barcode W19010009368 [web!]) Lasiosiphon capitatus (L.f.) Burtt Davy (for protologue see under southern African taxa) Lasiosiphon chapmanii (B.Peterson) Olaniyan, comb. nov. ≡ Gnidia chapmanii B.Peterson in Kew Bull. 31(1): 177. 1976. —TYPE: Malawi, Mlanje Mountain, 7 Jun 1970, Brummit 11346 (Holotype: K barcode K000049217 [web!], isotypes: EA barcode EA000002370 [web!], GB barcode GB0048775 [web!], LI barcode LISC011492 [web!], MAL, PRE barcode PRE0308076-0 [web!], SRGH [n.v.], UPS [n.v.]). Lasiosiphon chrysanthus (Solms ex Schweinf.) Olaniyan, comb. nov. ≡ Arthrosolen chrysanthus Solms ex Schweinf. in Beitr. Fl. Aethiop.: 165 .1867. ≡ Gnidia chrysantha (Solms ex Schweinf.) Gilg in Bot. Jahrb. Syst. 19(2–3): 258 (1894). —TYPE: Ethiopia “Bei Duli im Fesoghlu”, 21 Apr 1848, Cienkowski s.n (Holotype: B†); Ethiopia, “near Nadda”, 13 Oct 1954, Mooney 6257 (Neotype: K barcode K000322639 [web!], designated by Gastaldo ( 1969 :351). Lasiosiphon deserticola (Gilg) C.H. Wright (for protologue see under southern African taxa) Lasiosiphon dumicola (S.Moore) Olaniyan, comb. nov. ≡ Gnidia dumicola S.Moore in J. Bot. 57: 114. 1919. —TYPE: Angola, “between Forte Princeza Amelia and Limbala Monelilo”, 26 Sept, 1905, Gossweiler 2023 (Holotype: BM barcode BM000910844 [web!], isotypes: K barcode K000322604 [web!], BR barcode BR0000006299842 [web!]). Lasiosiphon elkerensis (Friis & Sebsebe) Olaniyan, comb. nov. ≡ Gnidia elkerensis Friis & Sebsebe in Webbia 73(2): 214. 2018. —TYPE: Ethiopia, Somalia Region “Afder Zone, El Kere Woreda”, 30 Dec 2016, Melaku Wondafrash, Shewangziwe Lemma, Wege Abebe, Shambel Alemu & Elias Tadesse 4182 (Holotype: ETH [n.v.], isotypes: C barcode C10022126 [web!], K [n.v.]). Lasiosiphon eminii (Engl. & Gilg) H.Pearson in Fl. Trop. Afr. 6(1, 2): 229. 1910 [as “ emini ”]. ≡ Gnidia eminii Engl. & Gilg in Bot. Jahrb. Syst. 19(2–3): 265. 1894 [as “ emini ”]. —TYPE: Tanzania, “Massaisteppe, Wald westlich Jrangi”, Jun 1892, Stuhlmann 4224 (Holotype: B†). Lasiosiphon foliosus (H.Pearson) Olaniyan, comb. nov. ≡ Arthrosolen foliosus H.Pearson in Fl. Trop. Afr. 6(1, 2): 237. 1910 [as foliosa ]. ≡ Gnidia foliosa (H.Pearson) Engl., Veg. Erde [Engler] 9(3, 2): 633. 1921. —TYPE: Sudan, “Upper Guinea. Soudan: between Bammako [Bamako] and Guignula”, 22 Jan 1899, Chevalier 256 (Holotype: K barcode K000322627 [web!], isotypes: BR barcode BR0000006299132 [web!], P barcodes P00516687 [web!] and P00516688 [web!]). Lasiosiphon fruticulosus (Gilg) Olaniyan, comb. nov. ≡ Gnidia fruticulosa Gilg in Bot. Jahrb. Syst. 23(1–2): 207. 1896. —TYPE: Angola, “Huilla” Antunes sub De Kindt 116 (Syntype: not located), 121 (Syntype: not located), and 316 (Syntypes: K barcode K000322562 [web!], LISC barcode LISC011490 [web!], P barcodes P00648774 [web!] and P00648775 [web!], W barcode W20020014719 [web!], WU barcode WU0026027 [web!]). Lasiosiphon fulgens (Welw. ex Hiern) Olaniyan, comb. nov. ≡ Gnidia fulgens Welw. ex Hiern, Cat. Afr. Pl. 1 (4): 923. 1900 [ Gnidia fulgens Welw., Apont. 548: 1859, nom nud.]. ≡ Arthrosolen fulgens (Welw. ex Hiern) H.Pearson in Fl. Trop. Afr. 6(1, 2): 237. 1910. —TYPE: Angola, Pungo Andongo, at Mutollo and in marshy meadows near Sobato Nbilla, Jan-Mar 1857, Welwitsch 6483 (Syntypes: BM barcode BM000910843 [web!], LISU barcodes LISU221402 [web!], LISU221403 [web!] and LISU221404 [web!], M barcode M0106551 [web!]). Lasiosiphon glaucus Fresen. (for protologue see under Asian taxa). Lasiosiphon goetzeanus (Gilg) Olaniyan, comb.nov. ≡ Gnidia goetzeana Gilg in Bot. Jahrb. Syst. 30(3–4): 363. 190. —TYPE: Tanzania, Njombe district,“Langenburg [Lumbira]”, May 1899, Goetze 895 (Lectotype: BR barcode BR0000006299101, designated by Robyns ( 1975 :43). Lasiosiphon gossweileri (S.Moore) Olaniyan, comb. nov. ≡ Arthrosolen gossweileri S.Moore in J. Bot. 57: 116. 1919. ≡ Gnidia gossweileri (S.Moore) B.Peterson in Fl. Zambes. 9(3): 98. 2006. —TYPE: Angola, “Munonque”, 20 Apr 1906, Gossweiler 3090 (Holotype: BM barcode BM000910842 [web!], isotypes: K barcode K000322605 [web!], BR barcode BR0000006249830 [web!]). Lasiosiphon gossweileri subsp. petersonii (Verdc.) Olaniyan, comb. nov. ≡ Gnidia gossweileri subsp. petersonii Verdc. in Fl. Zambes. 9(3): 98. 2006. —TYPE: Zambia, Mwinilunga District, 18 km East of Kalene Hill, 16 Dec 1963, Robinson 6111 (Holotype: K barcode K000322606 [web!]). Lasiosiphon hockii (De Wild) Olaniyan, comb. nov. ≡ Gnidia hockii De Wild in Repert. Spec. Nov. Regni Veg. 11: 536. 1913. —TYPE: Democratic Republic of the Congo, “Ober-Katanga: Elisabethville”, Sept 1911 Hock s.n . (Holotype: BR barcode BR0000008964878 [web!]). Lasiosiphon involucratus (Steud ex A.Rich) Olaniyan, comb. nov. ≡ Gnidia involucrata Steud. ex A.Rich. in Tent. Fl. Abyss. 2: 234. 1850. —TYPE: Ethiopia, Tigre, Sana to Terrfera, 5 Oct 1838, Schimper 770 (Lectotype: P barcode P00541070, designated by Gastaldo ( 1969 :347); isolectotypes: BR barcodes BR0000006249809 [web!] and BR0000006249854 [web!], BR0000006249847 [web!], G barcodes G00015753 [web!], G00015754 [web!] and G00015755 [web!], G-DC barcode GDC00131478 [web!], HAL barcode HAL0111681 [web!], K barcode K000322632 [web!], LG barcode LG0000090028878 [web!], M barcodes M0106555 [web!] and M0106556 [web!], MPU barcode MPU007261 [web!], P barcode P00648788 [web!], REG barcode REG000882 [web!], S barcode S05-9784 [web!], TUB barcodes TUB002865 [web!] and TUB002866 [web!]) Lasiosiphon kasaiensis (S.Moore) Olaniyan, comb. nov. ≡ Gnidia kasaiensis S.Moore in J. Bot. 57: 113. 1919. —TYPE: Democratic Republic of Congo, Kasai District, Sankuru River, Kässner 3322 (Holotype: BM barcode BM000910841 [web!], isotype: BR barcode BR0000008965530 [web!]). Lasiosiphon kraussianus (Meisn.) Burtt Davy (for protologue see under Asian taxa). Lasiosiphon kraussianus var. mollissimus (E.A. Bruce) Olaniyan, comb, nov. Gnidia kraussiana var. mollissima (E.A.Bruce) A. Robyns in Fl. Afr. Centr., Spermatophytes: 53. 1975. Lasiosiphon mollissimus E.A.Bruce in Bull. Misc. Inform. Kew 1940(2): 50. 1940. —TYPE: Zambia, "Northern Rhodesia. Abercorn District, Near Malombe, on the road to Kambole, 2 Jun 1936, Burtt 6115 (Holotype: K barcode K000322622 [web!], isotypes: BR barcode BR0000006249878 [web!], EA, K barcode K000322623 [web!]). Lasiosiphon kundelungensis (S.Moore) Olaniyan, comb. nov. ≡ Gnidia kundelungensis S.Moore in Journal of Botany, British and Foreign 57: 114. 1919. —TYPE: Democratic Republic of Congo, “West Kundelungu”, 17 May 1908, Kassner 2793 (Holotype: BM barcode BM000910840 [web!], isotypes: HBG barcode HBG512730 [web!], K barcode K000322631 [web!], P barcodes P00541090 [web!] and P00541091 [web!]). Lasiosiphon latifolius (Oliv.) Brenan in Kew Bull. 4(1): 93. 1949. ≡ Arthrosolen latifolius Oliv. in Trans. Linn. 2(15): 348. 1887. ≡ Gnidia latifolia (Oliv.) Gilg in Pflanzenw. Ost-Afrikas 2(C): 283 1895. —TYPE: Tanzania, Kilimanjaro, without date, Johnston s.n. (Lectotype: K barcode K000322589 [web!], designated by Gastaldo ( 1969 :365). Lasiosiphon lampranthus (Gilg) H. Pearson in Fl. Trop. Afr.6(1, 2): 233. 1910. ≡ Gnidia lamprantha Gilg in Bot. Jahrb. Syst. 19(2–3): 264. 1894. —TYPE: Tanzania, Bukoba District, Karagwe, 1892, Stuhlmann 3204 (Lectotype: K barcode K000322594 [web!], designated by Gastaldo ( 1969 :362). Lasiosiphon microcephalus (Meisn.) J.C.Manning & Magee (for protologue see under southern Africa taxa). Lasiosiphon mollis (C.H.Wright) Olaniyan, comb. nov. ≡ Gnidia mollis C.H.Wright in Bull. Misc. Inform. Kew 1906(1): 23. 1906. —TYPE: Tanzania, “lower plateau north of Lake Nyasa [Lake Malawi]”, without date, Thomson s.n. (Syntypes: K barcodes K000322661 [web!] and K000378910 [web!]); Mozambique, “between Unangu and Lake Shiré”, Nov. 1899, Johnson 12 (Syntype: K barcode K00322619 [web!]). Lasiosiphon newtonii (Gilg) Olaniyan, comb. nov. ≡ Gnidia newtonii Gilg in Bot. Jahrb. Syst. 23(1–2): 205 (1896). ≡ Arthrosolen newtonii (Gilg) H.Pearson in Fl. Trop. Afr. 6(1, 2): 236. 1910. —TYPE: Angola, “Huilla [Huila], Chella, Monpulla, Humpata, Biballa, in montibus divulgata”, without date, Newton 10 (Holotype:B†, isotype: K barcode K000378890 [web!]). Lasiosiphon oliverianus (Vatke ex Engl. & Gilg) Olaniyan, comb. nov. ≡ Gnidia oliveriana Vatke ex Engl. & Gilg in Bot. Jahrb. Syst. 19(2–3): 262. 1894. —TYPE: Angola, “Malandsche” [Malanje], Nov-Dec 1879, Mechow 325 (Holotype: B†; Lectotype: K barcode K000322559 [web!], designated by Robyns ( 1975 :56); isolectotypes: BR barcode BR0000006249991 [web!], G barcodes G00015475 [web!], G00015717 [web!], JE barcodes JE00003506 [web!], JE00003507 [web!], M barcode M0106558 [web!]). Lasiosiphon pleurocephalus (Gilg) Olaniyan, comb. nov. ≡ Gnidia pleurocephala Gilg in Kunene-Sambesi-Exped.: 310. 1903. ≡ Arthrosolen pleurocephalus (Gilg) H.Pearson in Fl. Trop. Afr. 6(1, 2): 236. 1910[as pleurocephala ]. —TYPE: Angola, “Quiriri”, 18 Apr 1900, Baum 828 (Holotype: B†, isotypes: G barcode G000190891 [web!], W barcode W19010006459 [web!]). Lasiosiphon poggei (Gilg) Olaniyan, comb. nov. ≡ Gnidia poggei Gilg in Bot. Jahrb. Syst. 19(2–3): 259. 1894. ≡ Arthrosolen poggei (Gilg) H.Pearson in Fl. Trop. Afr. 6(1, 2): 235. 1910. —TYPE: Angola, by the River Kuango, Sept 1876, Pogge 208 (Holotype: B†). Lasiosiphon quarrei (A. Robyns) Olaniyan, comb. nov. ≡ Gnidia quarrei A. Robyns in Bull. Jard. Bot. Natl. Belg. 45: 223. 1975. —TYPE: Democratic Republic of Congo: “Haut-Katanga, monts Marungu, Kakera, terr. Moba, en savane herbeuse”, Dec 1945, Quarré 7599 (Holotype: BR barcode BR0000008964885 [web!]). Lasiosiphon polyanthus (Gilg) C. H. Wright (for protologue see under southern African taxa). Lasiosiphon polycephalus (E.Mey. ex Meisn.) H.Pearson (for protologue see under southern African taxa). Lasiosiphon rendlei (Hiern) Olaniyan, comb. nov. ≡ Gnidia rendlei Hiern in Cat. Afr. Pl. (Hiern) 1: 924. 1900. —TYPE: Angola, “Huilla, between Mumpulla and Nene”, Oct 1859, Welwitsch 6478 (LISU-221410, syn.); Angola, “between Lopollo and Catumba”, Nov 1859, Welwitsch 6477 (Syntypes: BM barcode BM000910848 [web!], LISU barcode LISU221408 [web!], M barcode M010652 [web!]). Lasiosiphon rubrocinctus (Gilg) Olaniyan, comb. nov. ≡ Gnidia rubrocincta Gilg in Bot. Jahrb. Syst. 19(2): 259. 1894. —TYPE: Angola, “Oberes Kongogebiet, am Quango”, Sept 1876, Pogge 209 (Holotype: B†). Lasiosiphon sericocephalus (Meisn.) J.C.Manning & Boatwr. (for protologue see under southern African taxa) Lasiosiphon somalensis (Franch) H.Pearson (for protologue see under Asian taxa) Lasiosiphon stenophyllus (Gilg) Olaniyan, comb. nov. ≡ Gnidia stenophylla Gilg in Bot. Jahrb. Syst. 19(2): 259. 1894. —TYPE: Tanzania, Lushoto district, Usambara Mts., Kwa Mshuza, Aug 1893, Holst 8963 (Holotype: B†, Lectotype: K barcode K000322650 [web!]; designated by Peterson ( 1978 :24); isolectotypes: COI barcode COI00005795 [web!], G barcodes G00015473 [web!], G00015474 [web!], HBG barcode HBG512609 [web!], JE barcode JE00003502 [web!], M barcode M0106576 [web!], P barcodes P00541073 [web!], P00541074 [web!], S barcode S05-9780 [web!]). Lasiosiphon usafuae Gilg Olaniyan, comb. nov. ≡ Gnidia usafuae in Gilg in Bot. Jahrb. Syst. 30(3–4): 363. 190. —TYPE: Tanzania, Mbeya/Chunya Districts, slopes of Poroto Mts. Usafua, Jun 1899, Goetze 1042 (Holotype: B†; isotypes: BR barcodes BR0000006249861 [web!], BR0000006249984 [web!], E, G barcode G00022850 [web!], K barcodes K000322659 [web!], K000322660 [web!], P). Lasiosiphon welwitschii (Hiern) Olaniyan, comb. nov. ≡ Gnidia welwitschii Hiern, Cat. Afr. Pl. 1(4): 923. 1900. —TYPE: Angola, Huila, Apr 1860, Welwitsch 6482 (Syntypes: BM barcode BM000910855 [web!], LD barcode LD1501359 [web!], LISU barcodes LISU221405 [web!], LISU221406 [web!] and LISU221407 [web!], M barcodes M0106566 [web!], M0106567 [web!]). Declarations Acknowledgements This article is part of the PhD Thesis of O. D. Olaniyan. We acknowledge the funding provided by the University of Johannesburg and the African Centre for DNA Barcoding. We thank the staff of SANBI (Pretoria National Herbarium and Compton Herbarium) for providing herbarium specimens and facilities for the herbarium work. Author contributions: ODO: Laboratory performance of the research, data analyses, and writing of the first draft of the manuscript. JSB: Conceptualization of the research, interpretation of the result, taxonomic and nomenclatural treatment, reviewing and editing of the manuscript. ARM: Conceptualization of the research, interpretation of the result, taxonomic and nomenclatural treatment, herbarium collections, reviewing and editing of the manuscript. JCM: Conceptualization of the research herbarium collections, taxonomic and nomenclatural treatment, reviewing and editing of the manuscript. MVDB: Funding of the project, supervision of the laboratory work, data analyses, reviewing and editing of the manuscript. Conflict of Interest: All the authors have no conflict of interest regarding this research. References Aymonin GG (1965) Sur un Gnidia (Thyméléacées) à inflorescence complexe du Cameroun. Bull. Soc. Bot. 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Motsi MC, Moteetee AN, Beaumont AJ, Rye BL, Powell MP, Savolainen V, Van der Bank M (2010) A phylogenetic study of Pimelea and Thecanthes (Thymelaeaceae): Evidence from plastid and nuclear ribosomal DNA sequence data. Austral Syst Bot 23(4):270–284 Oguri E, Tagane S, Chhang P, Toyama H, Murakami N, Yahara T (2017). Flora of Bokor National Park, Cambodia VI: A new species of Wikstroemia (Thymelaeaceae), W. bokorensis . Phytotaxa 317(4):280–285. https://doi.org/10.11646/Phytotaxa 317.4.2 Olaniyan OD, Van der Bank M, Boatwright JS, Magee AR, Manning JC (2020) A taxonomic revision of Englerodaphne Gilg (Thymelaeaceae: Thymelaeoideae). S. African J. Bot. 130: 348–355. https://doi.org/10.1016/j.sajb.2020.01.015 Olaniyan OD (2021) Systematics of the subfamily Thymelaeoideae (Thymelaeaceae) in southern Africa. PhD Thesis, University of Johannesburg, Johannesburg. ProQuest Dissertations and Theses Global (Publication No. 29356354). Pearson HHW (1910). Order CXVIII: Thymelaeaceae. In: Thiselton-Dyer WT (ed) Flora of Tropical Africa. Reeve, L. & Co., England, pp 212–255 Peterson B (1959) Some interesting species of Gnidia . Bot Not 112(4):465–480 Peterson B (1978) Flora of Tropical East Africa. Thymelaeaceae. Crown Agents for Overseas Governments & Administrations, London Peterson B (2000) Thymelaeaceae. In: Edwards S, Mesfin T, Sebsebe D, Hedberg I (eds) Flora of Ethiopia and Eritrea: Magnoliaceae to Flacourtiaceae vol 2. National Herbarium, Addis Ababa University, Addis Ababa, pp 429–434 Peterson B (2006) Thymelaeaceae. In: Pope GV, Polhill RM, Martins ES (eds). Flora Zambesiaca. Royal Botanical Gardens, London, pp 85–117 Phillips EP (1944) Notes on some genera of the Thymelaeaceae. J S African Bot 10:61–67. Phillips EP (1951) The genera of South African flowering plants (No. 25). Cape Times Ltd., Govt. Printers, Capetown Prabhukumar KM, Rogers ZS, Hareesh VS, & Balachandran I (2018) Reinstatement and lectotypification of Gnidia sisparensis (Thymelaeaceae), a species endemic to India. Phytotaxa 372(2):179–182. https://doi.org/10.11646/phytotaxa.372.2.5 Rambaut A, Drummond AJ, Xie D, Baele G, Suchard MA (2018) Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Syst biol 67(5):901-9044. Rautenbach M (2006) Gnidia L. (Thymelaeaceae) is not monophyletic: Taxonomic implications for Gnidia and its relatives in Thymelaeoideae [Unpublished Master's thesis]. University of Johannesburg, Johannesburg Robinson C (2004) Molecular phylogenetics of Lachnaea (Thymelaeaceae): evidence from plastid and nuclear sequence data [Unpublished Master's thesis]. University of Johannesburg, Johannesburg Robyns A (1975) Thymelaeaceae. In: Bamps P (ed.) Flore d’Afrique Centrale . Jardin Botanique Nationale de Belgium, pp. 1–68 Rogers ZS, Spencer, MA (2006) Typification of plant names in Thymelaeaceae published by Linnaeus and Linnaeus filius. Taxon 55(2):483–488. https://doi.org/10.2307/25065596 Rogers ZS (2009) A Revision of Malagasy Gnidia (Thymelaeaceae, Thymelaeoideae). Ann Missouri Bot Gard 96(2):324–369 Ronquist F, Teslenko M, Van Der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Syst biol 61(3):539-542 Seelanan T. Schnabel A, Wendel JF (1997) Congruence and consensus in the cotton tribe (Malvaceae). Syst Biol 22(2):259–290 Stamatakis A (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 30(9):1312–1313 Staner P (1935) Les Thyméléacées de la Flore du Congo Belge. Bulletin Du Jardin Botanique de l’Etat Bruxelles 13:321–372 Swofford DL (2003) PAUP*: Phylogenetic Analysis Using Parsimony, version 4.0. Sunderland, MA: Sinauer Associates Townsend CC (1981) Thymelaeaceae. In: Dassanayake MD, Fosberg FR (eds) A revised Handbook to the Flora of Ceylon Vol 2. Amerind Publishing Co., Delhi, pp 501-511 Van der Bank M, Fay MF, Chase MW (2002) Molecular phylogenetics of Thymelaeaceae with particular reference to African and Australian genera. Taxon 51(2): 329–339 Van Niekerk A (2005) Phylogenetic relationships and speciation in the genus Passerina L. (Thymelaeaceae) inferred from chloroplast and nuclear sequence data [Unpublished Master's thesis]. University of Johannesburg, Johannesburg Wiens JJ (1998) Combining Data Sets with Different Phylogenetic Histories. Syst Biol 47(4):568–581 Wright CH (1915) Order CXVIII: Thymelaeaceae. In: Thiselton-Dyer WT (ed) Flora capensis. L. Reeve & Co., London, pp 1–81 Additional Declarations No competing interests reported. Supplementary Files Appendix1andTableS1.docx TableS2Typeofdataincluded.docx FigureS1CombinedPlastiddataset.tiff Fig. S1. Bayesian Inference (BI) majority-rule consensus tree from the combined plastid dataset ( rbcL , matK , rps16 and trnL-F ) of Thymelaeaceae. Bayesian posterior probability (PP) and Maximum-likelihood bootstrap (MLB) values are indicated above the branches, and Maximum Parsimony bootstrap (MPB) support values are indicated below the branches. FigureS2ITSdataset.tiff Fig. S2. Bayesian Inference (BI) majority-rule consensus tree from the ITS dataset of Thymelaeaceae. Bayesian posterior probability (PP) and Maximum-likelihood bootstrap (MLB) values are indicated above the branches, and Maximum Parsimony bootstrap (MPB) support values are indicated below the branches. FigureS3CombinedBayestree.tiff Fig. S3. Bayesian Inference (BI) majority-rule consensus tree from the combined dataset (nuclear + plastid) of Thymelaeaceae. Bayesian posterior probability (PP) values are indicated above the branches. FigureS4CombinedRaxMLtree.tiff Fig. S4. Phylogram of 116 taxa within Thymelaeaceae, as inferred through maximum-likelihood analysis of the combined (nuclear + plastid) dataset using RAxML. Maximum-likelihood bootstrap (MLB) values are indicated above the branches. FigureS5CombinedParsimonytree.tiff Fig. S5. One of the most parsimonious trees obtained from the maximum parsimony analyses. Maximum Parsimony bootstrap (MPB) support values are indicated above the branches. DataMatrices.zip Cite Share Download PDF Status: Published Journal Publication published 08 Aug, 2024 Read the published version in Plant Systematics and Evolution → Version 1 posted Editorial decision: Revision requested 04 May, 2024 Reviews received at journal 26 Mar, 2024 Reviewers agreed at journal 15 Feb, 2024 Reviewers agreed at journal 13 Feb, 2024 Reviewers invited by journal 12 Feb, 2024 Editor assigned by journal 02 Feb, 2024 Submission checks completed at journal 31 Jan, 2024 First submitted to journal 30 Jan, 2024 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-3910184","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":270370202,"identity":"74f17fad-a6bf-4f8f-aec5-df331c90a56f","order_by":0,"name":"Oluwayemisi Dorcas Olaniyan","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA2klEQVRIiWNgGAWjYFAC5gYGBgMgzd4ApogBjFAtPAdI0gICEglEauBnb2z8XFBwWN5c8o1BAUONHYN8+wH8WiR7DjZLzzA4bLhzdo6BAcOxZAbGHgLWGdxIbJDmMTjMuOE2SAvbAQZmBsJamn8DtdhvuHkGqOXfAQY2/gcEtbSBbEnccIPHwICx7QADD6FwAPqlzZrHID15w5m0AoPEvmQeCQkCtvCzNx++zfPH2nbD8cPbDD58s5OT7ydgCxQ0gwg2A6BiHqLUA0EdiGAm4KJRMApGwSgYqQAAs8s/1NNUf5YAAAAASUVORK5CYII=","orcid":"","institution":"University of Johannesburg","correspondingAuthor":true,"prefix":"","firstName":"Oluwayemisi","middleName":"Dorcas","lastName":"Olaniyan","suffix":""},{"id":270370203,"identity":"b77a1258-b030-49dc-a4a8-3426de73dcb5","order_by":1,"name":"J. Stephen Boatwright","email":"","orcid":"","institution":"University of the Western Cape","correspondingAuthor":false,"prefix":"","firstName":"J.","middleName":"Stephen","lastName":"Boatwright","suffix":""},{"id":270370204,"identity":"6beb7e0e-3c27-4dbd-be3e-47126527bf09","order_by":2,"name":"Anthony R. Magee","email":"","orcid":"","institution":"Compton Herbarium, South African National Biodiversity Institute","correspondingAuthor":false,"prefix":"","firstName":"Anthony","middleName":"R.","lastName":"Magee","suffix":""},{"id":270370205,"identity":"878567d4-363a-434c-bd21-4a89d4dec122","order_by":3,"name":"John C. Manning","email":"","orcid":"","institution":"Compton Herbarium, South African National Biodiversity Institute","correspondingAuthor":false,"prefix":"","firstName":"John","middleName":"C.","lastName":"Manning","suffix":""},{"id":270370206,"identity":"012b6608-2677-4656-871a-a5bf6ab37e50","order_by":4,"name":"Michelle Bank","email":"","orcid":"","institution":"University of Johannesburg","correspondingAuthor":false,"prefix":"","firstName":"Michelle","middleName":"","lastName":"Bank","suffix":""}],"badges":[],"createdAt":"2024-01-30 09:44:21","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-3910184/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-3910184/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s00606-024-01912-3","type":"published","date":"2024-08-08T15:57:12+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":50551560,"identity":"569f4b79-ddbd-4b07-8a12-8b3a87143f8c","added_by":"auto","created_at":"2024-02-02 11:13:20","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":1022812,"visible":true,"origin":"","legend":"\u003cp\u003eDiversity of petaloid scales and flower merosity in \u003cem\u003eGnidia\u003c/em\u003e group.\u003cem\u003e \u003c/em\u003e\u003cstrong\u003eA\u003c/strong\u003e, Pentamerous flower with five membranous petaloid scales in \u003cem\u003eLasiosiphon glaucus\u003c/em\u003e Fresen.; \u003cstrong\u003eB\u003c/strong\u003e, Tetramerous flower with four densely hairy petaloid scales in \u003cem\u003eGnidia pinifolia \u003c/em\u003eL; \u003cstrong\u003eC\u003c/strong\u003e, Tetramerous flower with eight fleshy petaloid scales in \u003cem\u003eGnidia clavata\u003c/em\u003e Schinz;\u003cem\u003e \u003c/em\u003e\u003cstrong\u003eD\u003c/strong\u003e, Tetramerous flower with four fleshy petaloid scales in \u003cem\u003eGnidia tenella \u003c/em\u003eMeisn;\u003cem\u003e \u003c/em\u003e\u003cstrong\u003eE\u003c/strong\u003e,\u003cem\u003e \u003c/em\u003ePentamerous flower without petaloid scales in\u003cem\u003e Lasiosiphon rubescens \u003c/em\u003e(B.Peterson) J.C.Manning \u0026amp; \u003ca href=\"https://www.ipni.org/a/20013214-1\"\u003eMagee\u003c/a\u003e;\u003cem\u003e \u003c/em\u003e\u003cstrong\u003eF\u003c/strong\u003e, Tetramerous flower without petaloid scales in \u003cem\u003eGnidia chrysantha \u003c/em\u003e(Solms) Gilg. Photos: Dinesh Valke (A); Carina Lochner (B); Annelise Vlok (C); Nicola van Berkel (D); Wynand Uys (E); Markus de Klerk (F).\u003c/p\u003e","description":"","filename":"Figure1Diversityofpetaloidscales.png","url":"https://assets-eu.researchsquare.com/files/rs-3910184/v1/6f898348c010febfd00a5e1e.png"},{"id":50551562,"identity":"4a0cb507-1b84-4449-bdc2-ca021c8253bf","added_by":"auto","created_at":"2024-02-02 11:13:20","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":578740,"visible":true,"origin":"","legend":"\u003cp\u003eBayesian Inference (BI) majority-rule consensus tree from the combined dataset (\u003cem\u003erbcL\u003c/em\u003e, \u003cem\u003ematK\u003c/em\u003e, \u003cem\u003erps16\u003c/em\u003e, \u003cem\u003etrnL-F, \u003c/em\u003eand\u003cem\u003e \u003c/em\u003eITS)\u003cem\u003e \u003c/em\u003eof Thymelaeaceae. Bayesian posterior probability (PP) and Maximum-likelihood bootstrap (MLB) values are indicated above the branches, and Maximum Parsimony bootstrap (MPB) support values are indicated below the branches.\u003c/p\u003e","description":"","filename":"Figure2Combinedtree.png","url":"https://assets-eu.researchsquare.com/files/rs-3910184/v1/87778c6bfe9f2899675ae127.png"},{"id":62298280,"identity":"9d4e03b9-eb89-4826-83f3-4c8bc8a610d6","added_by":"auto","created_at":"2024-08-12 16:11:35","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":3822206,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-3910184/v1/453e23b2-3014-4a44-b225-258c102c1a54.pdf"},{"id":50551569,"identity":"650f60d5-7d5f-4c25-a35c-a94160a9990b","added_by":"auto","created_at":"2024-02-02 11:13:25","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":84366,"visible":true,"origin":"","legend":"","description":"","filename":"Appendix1andTableS1.docx","url":"https://assets-eu.researchsquare.com/files/rs-3910184/v1/9d0a1154976b84094b1c4bce.docx"},{"id":50551561,"identity":"c3a4321b-32fe-4ee1-98bf-598e5a7728fd","added_by":"auto","created_at":"2024-02-02 11:13:20","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":19785,"visible":true,"origin":"","legend":"","description":"","filename":"TableS2Typeofdataincluded.docx","url":"https://assets-eu.researchsquare.com/files/rs-3910184/v1/9e02b0ee1314b0614d4eb063.docx"},{"id":50551564,"identity":"259b3ed7-f698-49a5-938c-eac2d20d31ff","added_by":"auto","created_at":"2024-02-02 11:13:21","extension":"tiff","order_by":3,"title":"","display":"","copyAsset":false,"role":"supplement","size":1726262,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eFig. S1.\u003c/strong\u003e Bayesian Inference (BI) majority-rule consensus tree from the combined plastid dataset (\u003cem\u003erbcL\u003c/em\u003e, \u003cem\u003ematK\u003c/em\u003e, \u003cem\u003erps16\u003c/em\u003e and \u003cem\u003etrnL-F\u003c/em\u003e)\u003cem\u003e \u003c/em\u003eof Thymelaeaceae. Bayesian posterior probability (PP) and Maximum-likelihood bootstrap (MLB) values are indicated above the branches, and Maximum Parsimony bootstrap (MPB) support values are indicated below the branches.\u003c/p\u003e","description":"","filename":"FigureS1CombinedPlastiddataset.tiff","url":"https://assets-eu.researchsquare.com/files/rs-3910184/v1/6e5750f0b204337e9e37927a.tiff"},{"id":50551570,"identity":"2bca7163-652f-4196-806c-ef3d410c4f57","added_by":"auto","created_at":"2024-02-02 11:13:25","extension":"tiff","order_by":4,"title":"","display":"","copyAsset":false,"role":"supplement","size":604230,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eFig. S2.\u003c/strong\u003e Bayesian Inference (BI) majority-rule consensus tree from the ITS dataset\u003cem\u003e \u003c/em\u003eof Thymelaeaceae. Bayesian posterior probability (PP) and Maximum-likelihood bootstrap (MLB) values are indicated above the branches, and Maximum Parsimony bootstrap (MPB) support values are indicated below the branches.\u003c/p\u003e","description":"","filename":"FigureS2ITSdataset.tiff","url":"https://assets-eu.researchsquare.com/files/rs-3910184/v1/ec300497b6c2c6fdceb93225.tiff"},{"id":50551566,"identity":"ee0a8752-70be-4a7a-99fa-f6663478d068","added_by":"auto","created_at":"2024-02-02 11:13:21","extension":"tiff","order_by":5,"title":"","display":"","copyAsset":false,"role":"supplement","size":539610,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eFig. S3. \u003c/strong\u003eBayesian Inference (BI) majority-rule consensus tree from the combined dataset (nuclear + plastid) of Thymelaeaceae. Bayesian posterior probability (PP) values are indicated above the branches.\u003c/p\u003e","description":"","filename":"FigureS3CombinedBayestree.tiff","url":"https://assets-eu.researchsquare.com/files/rs-3910184/v1/5927ebcb4e64ead733070011.tiff"},{"id":50551825,"identity":"34c3cd65-b7c8-488c-bb53-ad4368105067","added_by":"auto","created_at":"2024-02-02 11:21:21","extension":"tiff","order_by":6,"title":"","display":"","copyAsset":false,"role":"supplement","size":530688,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eFig. S4.\u003c/strong\u003e Phylogram of 116 taxa within Thymelaeaceae, as inferred through maximum-likelihood analysis of the combined (nuclear + plastid) dataset using RAxML. Maximum-likelihood bootstrap (MLB) values are indicated above the branches.\u003c/p\u003e","description":"","filename":"FigureS4CombinedRaxMLtree.tiff","url":"https://assets-eu.researchsquare.com/files/rs-3910184/v1/ce3189c82b702560ac452f25.tiff"},{"id":50551826,"identity":"e078585f-3749-41fc-9557-de72db8781f7","added_by":"auto","created_at":"2024-02-02 11:21:21","extension":"tiff","order_by":7,"title":"","display":"","copyAsset":false,"role":"supplement","size":517380,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eFig. S5. \u003c/strong\u003eOne of the most parsimonious trees obtained from the maximum parsimony analyses. Maximum Parsimony bootstrap (MPB) support values are indicated above the branches.\u003c/p\u003e","description":"","filename":"FigureS5CombinedParsimonytree.tiff","url":"https://assets-eu.researchsquare.com/files/rs-3910184/v1/edc721e9c42057706526cbed.tiff"},{"id":50551563,"identity":"1f43885f-6614-4b7b-87c8-e48a84f28856","added_by":"auto","created_at":"2024-02-02 11:13:20","extension":"zip","order_by":8,"title":"","display":"","copyAsset":false,"role":"supplement","size":71276,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cbr\u003e\u003c/p\u003e","description":"","filename":"DataMatrices.zip","url":"https://assets-eu.researchsquare.com/files/rs-3910184/v1/dc8098e5245bf6b4eeaa8313.zip"}],"financialInterests":"No competing interests reported.","formattedTitle":"Molecular support for transferring the tropical African species of Gnidia to Lasiosiphon (Thymelaeaceae: Thymelaeoideae) and a worldwide synopsis of the species","fulltext":[{"header":"1 Introduction","content":"\u003cp\u003eThe genus \u003cem\u003eLasiosiphon\u003c/em\u003e Fresen. (Thymelaeaceae: Thymelaeoideae) comprises a morphologically diverse group of plants distributed across southern and tropical Africa, Madagascar and India. The genus was established by Fresenius (\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e1838\u003c/span\u003e) for the single species \u003cem\u003eL. glaucus\u003c/em\u003e Fresen., based on its capitate inflorescence of pentamerous flowers, with five petaloid scales and ten stamens in two series. The circumscription of the genus subsequently underwent significant changes, primarily due to the increasing morphological and taxonomic complexity of the genus \u003cem\u003eGnidia\u003c/em\u003e. This resulted in uncertainties regarding the relationships and status between \u003cem\u003eGnidia\u003c/em\u003e and allied genera such as \u003cem\u003eEnglerodaphne\u003c/em\u003e Gilg, \u003cem\u003eArthrosolen\u003c/em\u003e C.A.Mey. and \u003cem\u003eLasiosiphon\u003c/em\u003e in the \u0026ldquo;\u003cem\u003eGnidia\u003c/em\u003e group\u0026rdquo; (sensu Herber 2023). Conflicting views on the importance of floral merosity and the presence or absence of petaloid scales (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) as generic distinctions led some taxonomists (Gilg 1894, \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e1921\u003c/span\u003e; Domke \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1934\u003c/span\u003e; Peterson \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e1959\u003c/span\u003e, \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e1978\u003c/span\u003e; Dyer \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1975\u003c/span\u003e) to include \u003cem\u003eArthrosolen\u003c/em\u003e and \u003cem\u003eLasiosiphon\u003c/em\u003e in an expanded concept of \u003cem\u003eGnidia\u003c/em\u003e whereas others (Endlicher \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1847\u003c/span\u003e; Meisner 1857; Bentham and Hooker \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1880\u003c/span\u003e; Pearson \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e1910\u003c/span\u003e; Wright \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e1915\u003c/span\u003e; Marloth \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e1925\u003c/span\u003e; Burtt-Davy \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e1926\u003c/span\u003e; Phillips \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e1944\u003c/span\u003e, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e1951\u003c/span\u003e; Leandri \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e1950\u003c/span\u003e; Hutchinson \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e1967\u003c/span\u003e) retained them as separate genera. Notably, Phillips (\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e1944\u003c/span\u003e, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e1951\u003c/span\u003e) radically altered the circumscription of \u003cem\u003eArthrosolen.\u003c/em\u003e Previously defined by the absence of petaloid scales, the genus was now defined by a capitate or capitulate inflorescence subtended by colourful, membranous bracts. This revised circumscription of the genus included mainly pentamerous species, mostly lacking petaloid scales (but rarely with petaloid scales). Phillips (\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e1944\u003c/span\u003e) transferred the remaining tetramerous species that had previously been included in \u003cem\u003eArthrosolen\u003c/em\u003e as lacking petaloid scales into \u003cem\u003eGnidia.\u003c/em\u003e In this taxonomy, \u003cem\u003eLasiosiphon\u003c/em\u003e included only pentamerous species with petaloid scales, \u003cem\u003eArthrosolen\u003c/em\u003e comprised mainly pentamerous species without petaloid scales but also one tetramerous species (\u003cem\u003eGnidia microcephala\u003c/em\u003e Meisn.) with petaloid scales, and \u003cem\u003eGnidia\u003c/em\u003e encompassed tetramerous species with or without petaloid scales.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe debate regarding the taxonomy of these genera persisted until Herber (\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2002\u003c/span\u003e, \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2003\u003c/span\u003e) and Peterson (\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2006\u003c/span\u003e) advocated for a very broad generic circumscription of \u003cem\u003eGnidia\u003c/em\u003e to include both \u003cem\u003eArthrosolen\u003c/em\u003e and \u003cem\u003eLasiosiphon\u003c/em\u003e. They characterized their enlarged \u003cem\u003eGnidia\u003c/em\u003e as having double the number of stamens as the number of sepals, arranged in two whorls.\u003c/p\u003e \u003cp\u003eMolecular phylogenetic studies have provided new and valuable insights into the evolutionary relationships among \u003cem\u003eGnidia\u003c/em\u003e and its allied genera (Van der Bank et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Beaumont et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Olaniyan \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). These studies showed that \u003cem\u003eGnidia\u003c/em\u003e, in the broad sense of Herber (\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2002\u003c/span\u003e, \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2003\u003c/span\u003e) and Peterson (\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2006\u003c/span\u003e), is polyphyletic and that \u003cem\u003eLasiosiphon\u003c/em\u003e, along with \u003cem\u003eArthrosolen\u003c/em\u003e sensu Phillips (\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e1944\u003c/span\u003e, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e1951\u003c/span\u003e), forms a distinct clade. This clade included all pentamerous species included in \u003cem\u003eGnidia\u003c/em\u003e s.l. plus various tetramerous taxa distinguishable by an involucrate inflorescence. Beaumont et al. (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) thus argued for the reinstatement of the genus \u003cem\u003eLasiosiphon\u003c/em\u003e, with an expanded circumscription to include tetramerous species with or without scales.\u003c/p\u003e \u003cp\u003eSubsequent authors (Manning and Boatwright \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Boatwright et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Magee and Manning \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) implemented the conclusions of Beaumont et al. (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) by providing combinations for species of \u003cem\u003eGnidia\u003c/em\u003e from southern Africa and Madagascar that exhibit characteristics consistent with the newly circumscribed \u003cem\u003eLasiosiphon\u003c/em\u003e. These combinations have been widely accepted in the taxonomic community. However, a comparable treatment of the tropical African species remains to be implemented.\u003c/p\u003e \u003cp\u003eFollowing Magee and Manning (\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), \u003cem\u003eLasiosiphon\u003c/em\u003e can be distinguished \u0026ldquo;by its pentamerous or sometimes tetramerous flowers in capitate, involucrate inflorescences subtended by foliaceous or papery bracts; a more or less pubescent hypanthium, with the basal part sometimes heavily bearded with long, straight, silky hairs; and waxy, bright yellow to orange or red sepal lobes, usually with petaloid scales in the mouth of the hypanthium tube but these lacking in some species.\u0026rdquo; Based on the original circumscription of \u003cem\u003eLasiosiphon\u003c/em\u003e and as confirmed by molecular phylogenetic analyses, all the pentamerous taxa in tropical Africa, some seven species, are readily accommodated in \u003cem\u003eLasiosiphon\u003c/em\u003e. As observed by Olaniyan (\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), however, the tetramerous \u003cem\u003eGnidia\u003c/em\u003e species in tropical Africa, which had not been included in previous phylogenetic analyses, shared the morphological traits consistent with the expanded circumscription of \u003cem\u003eLasiosiphon\u003c/em\u003e. This observation raises the likelihood that these species should be transferred to \u003cem\u003eLasiosiphon\u003c/em\u003e. All the Madagascan species are endemic to the island, and very few species are shared between tropical Africa and either southern Africa or Asia. These four regions thus represent convenient geographical divisions.\u003c/p\u003e \u003cp\u003eThe tetramerous species of tropical Africa can be segregated into three morphological groups based on the position and structure of their inflorescence: 1. Inflorescences umbellate, terminal and axillary, few-flowered, sessile; bracts foliaceous; pedicel non-hairy (\u003cem\u003eGnidia fastigiata\u003c/em\u003e Rendle); 2. inflorescences capitate, terminal and/or axillary, few-flowered, usually distinctly long or shortly pedunculate; involucral bracts papyraceous or coloured; pedicel hairy (e.g. \u003cem\u003eG. goetzeana\u003c/em\u003e Gilg); and 3. inflorescences composite-like heads, always terminal, many-flowered, distinctly long or shortly pedunculate; involucral bracts papyraceous or coloured; pedicel hairy (e.g. \u003cem\u003eG. mollis\u003c/em\u003e C.H.Wright).\u003c/p\u003e \u003cp\u003eThis paper aims to: 1. expand sampling to include representative samples from each of these morphological groups in a phylogenetic analysis of Thymelaeaceae, allowing us to assess their relationships within the broader context of \u003cem\u003eGnidia\u003c/em\u003e lineages and determine their taxonomic placement in the classification proposed by Beaumont et al. (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2009\u003c/span\u003e), Manning and Boatwright (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), Magee and Manning (\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) and Boatwright et al. (\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2017\u003c/span\u003e); 2. provide any necessary nomenclatural combinations for the \u003cem\u003eGnidia\u003c/em\u003e species in tropical Africa that currently lack combinations in \u003cem\u003eLasiosiphon\u003c/em\u003e; 3. transfer the three pentamerous \u003cem\u003eGnidia\u003c/em\u003e species from Asia to \u003cem\u003eLasiosiphon\u003c/em\u003e in accordance with the current circumscription of that genus; and 4. compile a comprehensive list of all \u003cem\u003eLasiosiphon\u003c/em\u003e species at regional level as a framework for future taxonomic work in the genus.\u003c/p\u003e"},{"header":"2 Material and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1 Loci selection and taxon sampling\u003c/h2\u003e \u003cp\u003eNew sequences of tropical African \u003cem\u003eGnidia\u003c/em\u003e were added to a reduced matrix of Olaniyan (\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) with combined data of five loci (ITS, \u003cem\u003ematK, rbcL, rps16\u003c/em\u003e, and \u003cem\u003etrnL-F\u003c/em\u003e) for phylogenetic analyses. Additionally, taxon sampling included representatives from all major lineages of \u003cem\u003eGnidia\u003c/em\u003e to ensure comprehensive coverage of its diversity. A total of 43 sequences for 13 taxa that were not included in previous studies were added to the pre-existing datasets. These include five newly sampled \u003cem\u003eGnidia\u003c/em\u003e species representing the distinct morphological groups found in tropical Africa. In all, 119 species from 24 genera in the subfamily Thymelaeoideae were analyzed, along with four taxa from the subfamily Octolepidoideae as outgroup taxa. Voucher specimens and GenBank accession numbers are listed in Appendix 1.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e2.2 DNA extraction, amplification, and sequencing\u003c/h2\u003e \u003cp\u003eDNA was extracted from herbarium specimens using the 10X cetyltrimethylammonium bromide (CTAB) method of Doyle and Doyle (\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1987\u003c/span\u003e) with the addition of 2% polyvinylpyrrolidone (PVP) to prevent the interaction of polyphenolic compounds with the DNA samples.\u003c/p\u003e \u003cp\u003ePolymerase Chain Reaction (PCR) amplification and sequencing were performed with parameters and amplification primers optimized for each locus based on previous studies: \u003cem\u003erbcL\u003c/em\u003e and \u003cem\u003etrnL-F\u003c/em\u003e regions as in Van der Bank et al. (\u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2002\u003c/span\u003e); ITS as in Beaumont et al. (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2009\u003c/span\u003e); \u003cem\u003ematK\u003c/em\u003e and \u003cem\u003erps16\u003c/em\u003e regions as in Motsi et al. (\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Geneious v.8.1.9 (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.geneious.com\u003c/span\u003e\u003cspan address=\"https://www.geneious.com\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e)\u003c/span\u003e was used for the editing and assembly of complementary strands. The sequences were aligned using Multiple Sequence Comparison by Log-Expectation [MUSCLE v3.8.31; (Edgar \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2004\u003c/span\u003e)] in the Geneious suite and visually checked for any alignment errors.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e2.3 Phylogenetic analyses\u003c/h2\u003e \u003cp\u003eThe nuclear (ITS) dataset, combined plastid (\u003cem\u003ematK, rbcL, rps16\u003c/em\u003e, and \u003cem\u003etrnL-F\u003c/em\u003e) datasets, and combined (ITS\u0026thinsp;+\u0026thinsp;plastid) datasets were analyzed using the Maximum Parsimony (MP) algorithm of PAUP* version 4.0a168 (Swofford, \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). Tree searches were conducted using a heuristic search with 1,000 random sequence additions and 10 trees held at each step. Tree bisection-reconnection (TBR) branch swapping and MULtrees were in effect. Delayed transformation character optimization (DELTRAN) was used for calculating branch lengths. Branch support was estimated using bootstrap analysis (Felsenstein \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1985\u003c/span\u003e) implemented in PAUP* with 1,000 replicates, employing TBR branch swapping, with ten trees held at each step and random sequence addition. Only values of 50% and above are reported. The following scale was used to evaluate bootstrap support: 50\u0026ndash;74%, weak; 75\u0026ndash;89%, moderate; 90\u0026ndash;100%, strong. The congruence of the individual dataset was assessed by comparing bootstrap percentages. The trees were considered incongruent only when they exhibited \u0026lsquo;hard\u0026rsquo; (i.e. with strong bootstrap support) rather than \u0026lsquo;soft\u0026rsquo; (i.e., with weak bootstrap support) incongruence (Seelanan et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Wiens \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e1998\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBayesian inference (BI) analyses were carried out on the nuclear (ITS) dataset, combined plastid datasets, and combined (ITS\u0026thinsp;+\u0026thinsp;plastid) datasets using MrBayes on XSEDE v.3.2.7 (Ronquist et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) as implemented on the CyberInfrastructure for Phylogenetic Research (CIPRES) Science Gateway (Miller et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Before the Bayesian analysis, the following best-fit model of evolution was determined for each matrix: ITS (TIM1\u0026thinsp;+\u0026thinsp;I\u0026thinsp;+\u0026thinsp;G), \u003cem\u003erps16\u003c/em\u003e (TVM\u0026thinsp;+\u0026thinsp;G), \u003cem\u003ematK\u003c/em\u003e (TVM\u0026thinsp;+\u0026thinsp;G), \u003cem\u003erbcL\u003c/em\u003e (TVM\u0026thinsp;+\u0026thinsp;I\u0026thinsp;+\u0026thinsp;G), and \u003cem\u003etrnL-F\u003c/em\u003e (TPM1uf\u0026thinsp;+\u0026thinsp;G). This was done with jModeltest 2.1.9 v20160115 (Darriba et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) which uses log-likelihood scores to estimate the model of DNA evolution that best suits a specific dataset. Four parallel Markov chain Monte Carlo (MCMC) estimations were run for 10,000,000 generations, with trees sampled every 1000 generations. The stationarity was checked using tracer v 1.7 (Rambaut et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), and all ESS values were found to be above 200 \u003cb\u003e\u0026ndash;\u003c/b\u003e indicating that the number of generations was sufficient. Suboptimal trees were discarded as the \u0026lsquo;burn-in\u0026rdquo; phase. The posterior probability values were evaluated using the following scale: below 0.95, weak; 0.95\u0026ndash;1.0, strong.\u003c/p\u003e \u003cp\u003eMaximum Likelihood (ML) analyses were performed on the nuclear (ITS) dataset, combined plastid datasets, and combined (ITS\u0026thinsp;+\u0026thinsp;plastid) datasets using RAxML-HPC2 on XSEDE (Stamatakis \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) on the CIPRES Science Gateway (Miller et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). A single consensus tree was generated from 1,000 nonparametric bootstrap replicates. The following scale was used to evaluate the node support: 50\u0026ndash;74%, weak; 75\u0026ndash;89%, moderate; 90\u0026ndash;100%, strong.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003e2.4 Taxonomic treatment\u003c/h2\u003e \u003cp\u003eFor the list of \u003cem\u003eLasiosiphon\u003c/em\u003e species we followed the most recent regional treatments for each geographical region as follows:\u003c/p\u003e \u003cp\u003e \u003cb\u003eSouthern Africa\u003c/b\u003e: Wright in Flora Capensis 5: 1\u0026ndash;81 (1915), Manning and Boatwright in Bothalia 43(1): 84\u0026ndash;88 (2013); Magee and Manning in Kew Bull. 78(1): 1\u0026ndash;6 (2017), and Kolokoto et al. in Bothalia 49(1): 3\u0026ndash;5 (2019).\u003c/p\u003e \u003cp\u003e \u003cb\u003eTropical Africa\u003c/b\u003e: Pearson in Flora of Tropical Africa 6(1, 2): 212\u0026ndash;255 (1910); Staner in Bulletin Jardine Botanique l\u0026rsquo;Etat. Bruxelles 13(4): 355 (1935); Aymonin in Flore du Gabon 2: 35\u0026ndash;95 (1966); Peterson in Flora of Tropical East Africa: 1\u0026ndash;37 (1978); Cuccuini in Flora of Somalia 1: 200\u0026ndash;202(1993); Peterson in Flora of Ethiopia and Eritrea 2(1): 429\u0026ndash;435 (2000); Peterson in Flora Zambesiaca 9(3): 85\u0026ndash;117 (2006); Figueiredo and Smith in Strelitzia 22: 162\u0026ndash;163 (2008); \u003cb\u003eMadagascar\u003c/b\u003e: Rogers in Annals of the Missouri Botanical Garden 96: 324\u0026ndash;368 (2009); Boatwright et al. in S. African J. Bot. 112: 336\u0026ndash;337 (2017);\u003c/p\u003e \u003cp\u003e \u003cb\u003eAsia (Sri Lanka, India, and Arabia)\u003c/b\u003e: Gastaldo in Webbia 24(1): 337\u0026ndash;389 (1969); Townsend in Revised Handbook of the Flora of Ceylon 2: 501\u0026ndash;511 (1981); and Prabhukumar et al. in Phytotaxa 372(2): 180 (2018).\u003c/p\u003e \u003c/div\u003e"},{"header":"3 Results","content":"\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003e3.1 Sequence characteristics\u003c/h2\u003e \u003cp\u003eA summary of statistics for all datasets is presented in Table \u003cspan refid=\"MOESM2\" class=\"InternalRef\"\u003eS2\u003c/span\u003e. Among the plastid regions, \u003cem\u003ematK\u003c/em\u003e had the most variable sites (278; 36%) compared to \u003cem\u003erps16\u003c/em\u003e (338; 35%), \u003cem\u003etrnL-F\u003c/em\u003e (475; 34%) and \u003cem\u003erbcL\u003c/em\u003e (260; 19%). The percentage of parsimony informative characters was higher in \u003cem\u003ematK\u003c/em\u003e (163; 21%) and \u003cem\u003erps16\u003c/em\u003e (201; 21%) than in \u003cem\u003etrnL-F\u003c/em\u003e (249; 18%), and \u003cem\u003erbcL\u003c/em\u003e (170; 12%). The ITS region had significantly more variable sites (446; 62%) and a higher number of parsimony informative sites (372; 52%) than any of the plastid regions. It also showed more site changes than the plastid regions (5.3 changes per variable site) reflecting that the ITS tree has a higher phylogenetic signal than the plastid regions. The plastid regions were not analyzed separately due to low sequence divergence but rather in combination.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003e3.1 Phylogenetic analyses\u003c/h2\u003e \u003cp\u003eThe comparison of the topologies derived from the combined plastid (Fig. \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e) and ITS dataset (Fig. \u003cspan refid=\"MOESM2\" class=\"InternalRef\"\u003eS2\u003c/span\u003e) showed significant agreement in the delimitation of the segregate clades of \u003cem\u003eGnidia\u003c/em\u003e. Also, all the genera in the subfamily Thymelaeoideae, including \u003cem\u003eLasiosiphon\u003c/em\u003e, were consistently well supported in both analyses except for \u003cem\u003eLachnaea\u003c/em\u003e L., which did not receive support for its monophyly in both datasets. We combined the two datasets since there were no strongly supported incongruences in them. Additionally, the BI (Fig. S3), ML (Fig. S4) and MP (Fig. S5) analyses of the concatenated dataset produced similar topologies; hence, results and combined support values are displayed on the BI majority rule consensus tree (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThere was strong support for the monophyly of the subfamily Thymelaeoideae [Bayesian posterior probability (PP)\u0026thinsp;=\u0026thinsp;1.00, maximum likelihood bootstrap (MLB)\u0026thinsp;=\u0026thinsp;100%, maximum parsimony bootstrap (MPB)\u0026thinsp;=\u0026thinsp;98%] with \u003cem\u003eAquilaria\u003c/em\u003e Lam. and \u003cem\u003eGyrinops\u003c/em\u003e Gaertn sister to the rest of the Thymelaeoideae (PP\u0026thinsp;=\u0026thinsp;100, MLB\u0026thinsp;=\u0026thinsp;100, MPB\u0026thinsp;=\u0026thinsp;97). Consistent with the findings of previous studies, \u003cem\u003eGnidia\u003c/em\u003e was polyphyletic in our study and dispersed as ten lineages (Lineages A-I) within Thymelaeoideae. Seven of the lineages (Lineages A-G) were strongly supported as monophyletic (PP\u0026thinsp;=\u0026thinsp;1.00, MLB\u0026thinsp;=\u0026thinsp;100, MPB\u0026thinsp;=\u0026thinsp;100), while the remaining three lineages (Lineages H-J), which were mainly tropical African, were nested in \u003cem\u003eLasioisphon\u003c/em\u003e, rendering \u003cem\u003eLasiosiphon\u003c/em\u003e paraphyletic. There were two strongly supported subclades within \u003cem\u003eLasiosiphon.\u003c/em\u003e One of the lineages of \u003cem\u003eGnidia\u003c/em\u003e (Lineage H) was unresolved in a subclade (PP\u0026thinsp;=\u0026thinsp;1.00, MLB\u0026thinsp;=\u0026thinsp;100, MPB\u0026thinsp;=\u0026thinsp;100) of pentamerous species, while the remaining two lineages (Lineages I-J) were also weakly nested within a subclade (PP\u0026thinsp;=\u0026thinsp;1.00, MLB\u0026thinsp;=\u0026thinsp;100, MPB\u0026thinsp;=\u0026thinsp;100) of mainly tetramerous species. Other genera within the \u0026ldquo;\u003cem\u003eGnidia\u003c/em\u003e group\u0026rdquo; (sensu Herber, \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2003\u003c/span\u003e), viz. \u003cem\u003ePasserina\u003c/em\u003e L. \u003cem\u003ePeddiea\u003c/em\u003e Harv. ex Hook., \u003cem\u003ePimelea\u003c/em\u003e Banks \u0026amp; Sol. ex Gaertn., \u003cem\u003eStephanodaphne\u003c/em\u003e Baill., \u003cem\u003eStruthiola\u003c/em\u003e L., \u003cem\u003eSynaptolepis\u003c/em\u003e Oliv. and \u003cem\u003eWikstroemia\u003c/em\u003e were each inferred to be monophyletic with strong supports (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The sister groupings \u003cem\u003eDrapetes\u003c/em\u003e Banks ex Lam. + \u003cem\u003eKelleria\u003c/em\u003e Endl. and \u003cem\u003eDais\u003c/em\u003e L. + \u003cem\u003ePhaleria\u003c/em\u003e Jack was strongly supported in all three analyses (PP\u0026thinsp;=\u0026thinsp;1.00, MLB\u0026thinsp;=\u0026thinsp;100, MPB\u0026thinsp;=\u0026thinsp;100), whereas \u003cem\u003eDirca\u003c/em\u003e L. + \u003cem\u003eOvidia\u003c/em\u003e (Poepp. \u0026amp; Endl.) Meisn. received strong support in the BI and ML (PP\u0026thinsp;=\u0026thinsp;1.00, MLB\u0026thinsp;=\u0026thinsp;92) analyses and weak support in the MP analysis (MPB\u0026thinsp;=\u0026thinsp;76). \u003cem\u003eLachnaea\u003c/em\u003e was found to be paraphyletic with respect to a lineage of \u003cem\u003eGnidia\u003c/em\u003e (Lineage A). Support for the inclusion of this lineage was strong in the BI and ML (PP\u0026thinsp;=\u0026thinsp;1.00, MLB\u0026thinsp;=\u0026thinsp;97) analyses but moderate in the MP analysis (MPB\u0026thinsp;=\u0026thinsp;79). The monophyly of \u003cem\u003eEnglerodaphne\u003c/em\u003e Gilg received strong support in the BI and ML (PP\u0026thinsp;=\u0026thinsp;1.00, MLB\u0026thinsp;=\u0026thinsp;96) analyses but moderate support in the MP analysis (MPB\u0026thinsp;=\u0026thinsp;76).\u003c/p\u003e \u003c/div\u003e"},{"header":"4 Discussion","content":"\u003cp\u003eThis study corroborates and expands upon the conclusions of other research (Van der Bank et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Beaumont et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Olaniyan \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) by uncovering the existence of numerous distinct evolutionary lineages within the genus \u003cem\u003eGnidia\u003c/em\u003e. This implies that the existing classification system may not reflect actual evolutionary relationships among \u003cem\u003eGnidia\u003c/em\u003e species, thereby necessitating a reassessment of their taxonomy. Prior researchers (Beaumont et al \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Foster et al \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Olaniyan \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) have examined the possibilities of establishing \u003cem\u003eGnidia\u003c/em\u003e as a monophyletic group, using the fundamental principle of monophyly along with certain secondary principles, but have not reached a consensus. A comprehensive analysis of morphological data and the integration of ecological data may be necessary to completely execute the optimal choice. However, while the decision on the reclassification of \u003cem\u003eGnidia\u003c/em\u003e species is still pending, this study is a valuable contribution towards achieving a monophyletic classification for \u003cem\u003eGnidia\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eConsistent with the phylogenetic analyses of Olaniyan (\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), \u003cem\u003eGnidia fastigiata\u003c/em\u003e was resolved in one of the lineages of \u003cem\u003eGnidia\u003c/em\u003e that included mainly southern African species. \u003cem\u003eG. fastigiata\u003c/em\u003e differs from other tropical African tetramerous species by its umbellate inflorescence (usually 2\u0026ndash;3 flowered), leaf-like bracts, and non-hairy pedicel. This justifies its non-inclusion in the \u003cem\u003eLasiosiphon\u003c/em\u003e clade. Significantly, it is one of two tetramerous tropical African \u003cem\u003eGnidia\u003c/em\u003e species with a geographical range extending to southern Africa, the other being \u003cem\u003eG. microcephala\u003c/em\u003e Meisn. [now \u003cem\u003eLasiosiphon microcephalus\u003c/em\u003e (Meisn.) J.C.Manning \u0026amp; Magee (Magee and Manning \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2017\u003c/span\u003e)], which is distinct in its capitate involucrate inflorescence and its colored modified bracts.\u003c/p\u003e \u003cp\u003eBoth nuclear and plastid data strongly support the inclusion of tropical African \u003cem\u003eGnidia\u003c/em\u003e species in \u003cem\u003eLasiosiphon\u003c/em\u003e. Despite less resolution and support among \u003cem\u003eLasiosiphon\u003c/em\u003e species, the analyses reveal two robustly supported subclades within the genus. The first subclade comprises pentamerous species with or without scales from southern Africa and Madagascar. Notably, \u003cem\u003eG. chapmanii\u003c/em\u003e B.Peterson (Lineage H), with pentamerous flowers, is nested within this clade. The second subclade encompasses tetramerous species, with four tropical African species (Lineages I \u0026amp; J) represented in the clade. This subdivision hints at potential infrageneric groups within \u003cem\u003eLasiosiphon\u003c/em\u003e, suggesting that floral merosity could serve as an informative character for infrageneric classifications in the genus pending more comprehensive sampling.\u003c/p\u003e \u003cp\u003e \u003cem\u003eLasiosiphon\u003c/em\u003e is characterized by its unique floral morphology of an involucrate and pedunculate inflorescence. In a comparative study on bract characteristics within \u003cem\u003eGnidia\u003c/em\u003e s.l., Beaumont et al. (2001) identified three categories: leaf-like bracts, partly modified bracts, and highly modified bracts. The purpose of establishing these categories was to classify the different types of bracts based on their similarity or dissimilarity to cauline leaves. The study revealed that all the \u003cem\u003eGnidia\u003c/em\u003e species except the ones formerly classified in \u003cem\u003eLasiosiphon\u003c/em\u003e are either lacking bracts or bearing unmodified leaf-like bracts. This suggests that the presence or absence of bracts, as well as their level of modification, can be used as a distinguishing characteristic within \u003cem\u003eGnidia\u003c/em\u003e s.l. For example, species of \u003cem\u003eEnglerodaphne\u003c/em\u003e that were previously put in the genus \u003cem\u003eGnidia\u003c/em\u003e were moved back to that genus because of their unique bract features of ebracteate inflorescence (Olaniyan 2020). In the same vein, a partly or highly modified bract highlights the distinctiveness of \u003cem\u003eLasiosiphon\u003c/em\u003e in the group. Interestingly, \u003cem\u003eGnidia\u003c/em\u003e species in the tropical Africa region (except \u003cem\u003eG. fastigiata\u003c/em\u003e) have an involucre of bracts that exhibit some degree of modification that is absent in other \u003cem\u003eGnidia\u003c/em\u003e lineages, which further supports their inclusion in the \u003cem\u003eLasiosiphon\u003c/em\u003e clade. This floral morphology is a key trait that sets \u003cem\u003eLasiosiphon\u003c/em\u003e apart from other lineages of \u003cem\u003eGnidia\u003c/em\u003e and helps in its identification.\u003c/p\u003e"},{"header":"5. Conclusion","content":"\u003cp\u003eThe polyphyletic nature of the genus \u003cem\u003eGnidia\u003c/em\u003e has been a topic of recent discussion. Previous research has shown the existence of multiple lineages within the genus, and this study supports that conclusion. Olaniyan (\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) sampled the full morphological diversity of \u003cem\u003eGnidia\u003c/em\u003e except the tropical African species, resulting in the discovery of additional \u003cem\u003eGnidia\u003c/em\u003e lineages. Her findings indicated that no additional lineages are expected, except for the tropical African species which were not adequately represented in her study. She proposed that these should be classified under the genus \u003cem\u003eLasiosiphon\u003c/em\u003e based on a review of critical literature that shed lights on their morphological characteristics. The sampling of species from tropical Africa confirmed that they do indeed belong to the \u003cem\u003eLasiosiphon\u003c/em\u003e clade, providing compelling evidence for the transfer to that genus of the tropical African species that remain in \u003cem\u003eGnidia\u003c/em\u003e. We, therefore, provide new generic combinations for the tropical African species of \u003cem\u003eGnidia\u003c/em\u003e that lack combinations in \u003cem\u003eLasiosiphon\u003c/em\u003e.\u003c/p\u003e"},{"header":"6 Taxonomic Treatment","content":"\u003cp\u003e \u003cb\u003eLasiosiphon\u003c/b\u003e Fresen. in Flora 21(2): 602. 1838; Meisner in Candolle, Prodr. [A. P. de Candolle] 14(2): 593. 1857; Wright in Fl. Cap. (Harvey) 5(2, 1): 69. 1915; Pearson in Fl. Trop. Afr. 6(1, 2): 227. 1910; Domke in Biblioth. Bot. 27(111): 135. 1934; Phillips in Gen. S. Afr. Fl. Pl. (ed.2): 528. 1951. \u0026equiv;\u003cem\u003eGnidia\u003c/em\u003e subg. \u003cem\u003eLasiosiphon\u003c/em\u003e (Fresen) Engler, Veg. Erde [Engler] 9(3, 2): 634. 1921. \u0026equiv;\u003cem\u003eGnidia\u003c/em\u003e sect. \u003cem\u003eLasiosiphon\u003c/em\u003e (Fresen.) Staner in Bull. Jard. Bot. \u0026Eacute;tat Bruxelles 13(4): 355. 1935. Type\u0026mdash;\u003cem\u003eLasiosiphon glaucus\u003c/em\u003e Fresen.\u003c/p\u003e \u003cp\u003e \u003cem\u003eArthrosolen\u003c/em\u003e [unranked] \u003cem\u003eCephalodaphne\u003c/em\u003e C.A.Mey. in Bull. Cl. Phys.-Math. Acad. Imp. Sci. Saint-P\u0026eacute;tersbourg 1: 359. 1843, \u003cb\u003esyn. nov\u003c/b\u003e. \u0026equiv;\u003cem\u003eArthrosolen\u003c/em\u003e sect. \u003cem\u003eCephalodaphne\u003c/em\u003e (C.A.Mey) Tiegh in Bull. Soc. Bot. France 40: 75. 1893. \u0026mdash;TYPE: Not designated [Species included: \u003cem\u003eA. calocephalus\u003c/em\u003e (Meisn.) C.A.Mey., \u003cem\u003eA. polycephalus\u003c/em\u003e (E.Mey. ex Meisn.) C.A.Mey., \u003cem\u003eA. anthylloides\u003c/em\u003e (L.f.) C.A.Mey.].\u003c/p\u003e \u003cp\u003e \u003cem\u003eGnidia\u003c/em\u003e sect. \u003cem\u003eInvolucratae\u003c/em\u003e Gilg in Engler \u0026amp; Prantl., Nat. Pflanzenfam. 3(6a): 227. 1894, \u003cb\u003esyn. nov.\u003c/b\u003e \u0026mdash;LECTOTYPE (\u003cb\u003edesignated here\u003c/b\u003e): \u003cem\u003eLasiosiphon involucratus\u003c/em\u003e (Steud ex A.Rich) Olaniyan (\u0026equiv;\u0026thinsp;\u003cem\u003eGnidia involucrata\u003c/em\u003e Steud. ex A.Rich).\u003c/p\u003e \u003cp\u003e \u003cem\u003eAtemnosiphon\u003c/em\u003e Leandri in Not. Syst. 13: 44. 1947, \u003cb\u003esyn. nov.\u003c/b\u003e \u0026mdash;LECTOTYPE (\u003cb\u003edesignated here\u003c/b\u003e): \u003cem\u003eLasiosiphon coriaceus\u003c/em\u003e Leandri (\u0026equiv;\u0026thinsp;\u003cem\u003eAtemnosiphon coriaceus\u003c/em\u003e (Leandri) Leandri).\u003c/p\u003e \u003cp\u003e \u003cem\u003eArthrosolen\u003c/em\u003e sensu Phillips in J. S. African Bot. 10: 64. 1944, non \u003cem\u003eArthrosolen\u003c/em\u003e C.A.Mey. in Bull. Cl. Phys.-Math. Acad. Imp. Sci. Saint-P\u0026eacute;tersbourg 1: 359. 1843. \u0026mdash;TYPE: \u003cem\u003eArthrosolen polycephalus\u003c/em\u003e (E.Mey. ex Meisn.) C.A.Mey. [\u0026equiv;\u0026thinsp;\u003cem\u003eGnidia polycephala\u003c/em\u003e (E.Mey. ex Meisn.) Gilg].\u003c/p\u003e \u003cp\u003e \u003cem\u003eNomenclatural note\u003c/em\u003e: Meyer (1843: 359) designated the species \u003cem\u003eArthrosolen spicatus\u003c/em\u003e C.A.Mey. as the type of his new genus \u003cem\u003eArthrosolen\u003c/em\u003e but the typesetting on this page is slightly confusing and Phillips (\u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e1951\u003c/span\u003e: 528) overlooked this, and in consequence lectotypified the genus against \u003cem\u003eA. polycephalus\u003c/em\u003e (E.Mey. ex Meisn.) C.A.Mey. This lectotypification is nomenclaturally superfluous but has been generally accepted until now. \u003cem\u003eArthrosolen polycephalus\u003c/em\u003e falls within the circumscription of \u003cem\u003eLasiosiphon\u003c/em\u003e and as a result, the generic name \u003cem\u003eArthrosolen\u003c/em\u003e has been accepted as a synonym of \u003cem\u003eLasiosiphon\u003c/em\u003e. The designated type of the genus, \u003cem\u003eA. spicatus\u003c/em\u003e, does not, however, fall within the morphological circumscription of \u003cem\u003eLasiosiphon\u003c/em\u003e and it has been retrieved as nested within one of the other lineages of \u003cem\u003eGnidia\u003c/em\u003e. The generic name \u003cem\u003eArthrosolen\u003c/em\u003e is therefore to be treated as applying to the lineage containing \u003cem\u003eG. spicata\u003c/em\u003e and is not to be treated as a synonym of \u003cem\u003eLasiosiphon\u003c/em\u003e.\u003c/p\u003e \u003cp\u003e \u003cb\u003eSpecies from southern Africa\u003c/b\u003e:\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon albosericeus\u003c/b\u003e (M.Moss ex B.Peterson) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia albosericea\u003c/em\u003e M.Moss ex B.Peterson in Bothalia 14(1): 81. 1982. \u0026equiv;\u003cem\u003eLasiosiphon ornatus\u003c/em\u003e Burtt Davy, Man. Pl. Transvaal 1: 206. 1926 as nom. nov. [non \u003cem\u003eG, ornata\u003c/em\u003e (Meisn.) Gilg]. \u0026mdash;TYPE: Swaziland, Hlatikulu, Dec 1910, \u003cem\u003eM. Stewart 14\u003c/em\u003e (Holotype: K Barcode K000322563 [web!]; Isotype: SAM Barcode SAM0002522-0 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon anthylloides\u003c/b\u003e (L.f.) Meisn. in Prodr. [A.P. de Candolle] 14(2): 595. 1857. \u0026equiv;\u003cem\u003ePasserina anthylloides\u003c/em\u003e L.f., Suppl. Pl.: 225. 1782. \u0026equiv;\u003cem\u003eArthrosolen anthylloides\u003c/em\u003e (L.f.) C.A.Mey. in Bull. Cl. Phys.-Math. Acad. Imp. Sci. Saint-P\u0026eacute;tersbourg 1: 359. 1843. \u0026equiv;\u003cem\u003eGnidia anthylloides\u003c/em\u003e (L.f.) Gilg in Engler \u0026amp; Prantl., Nat. Pflanzenfam. 3(6a): 227. 1894. \u0026mdash;TYPE: South Africa, Western cape, \"Cap. bonai. spei.\u0026rdquo; [Cape of good hope], without date, \u003cem\u003eThunberg s.n.\u003c/em\u003e (Lectotype: UPS-THUNB 9566, designated by Rogers (\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2006\u003c/span\u003e:486); isolectotypes: S barcode S-G-4573 [web!], SBT barcode SBT12559 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon anthylloides\u003c/b\u003e var. \u003cb\u003evulgaris\u003c/b\u003e (Meisn.) Meisn. in Prodr. [A. P. de Candolle] 14(2): 596. 1857. \u0026equiv;\u003cem\u003ePasserina anthylloides\u003c/em\u003e var. \u003cem\u003evulgaris\u003c/em\u003e Meisn. in Linnaea 14(5): 393. 1841. \u0026mdash;TYPE: South Africa, \u0026ldquo;Hue spectat specim. Burchell (cat. No. 2203) in Herb. DC\u0026rdquo;, without date, \u003cem\u003eBurchell 2203\u003c/em\u003e (Holotype: G-DC [n.v.]). Other original material: South Africa, \u0026ldquo;Vanstadesberg, inter Zandplaat et Komga, usque ad alt. 3,000 ped., 27 Dec 1829, \u003cem\u003eDr\u0026egrave;ge 7347\u003c/em\u003e (HBG barcode HBG512551 [web!], P barcode P00541076 [web!]); without date, \u003cem\u003eDr\u0026egrave;ge 4659\u003c/em\u003e (NY barcode NY0386302 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon anthylloides\u003c/b\u003e var. \u003cb\u003eglabrescens\u003c/b\u003e (Meisn.) Meisn. in Prodr. [A.P. de Candolle] 14(2): 596. 1857. \u0026equiv;\u003cem\u003ePasserina anthylloides\u003c/em\u003e var. \u003cem\u003eglabrescens\u003c/em\u003e Meisn. in Linnaea 14(5): 393. 1841. \u0026mdash;TYPE: South Africa, Kwazulu Natal, \u0026ldquo;Inter gramina ad sylvae marginem prope Port Natal, infra 200 ped. alt. (V,c, 34)\", 8 Apr 1832, \u003cem\u003eDr\u0026egrave;ge s.n.\u003c/em\u003e (Syntypes: HAL barcode HAL0076215 [web!], P barcode P00648792 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon anthylloides\u003c/b\u003e var. \u003cb\u003emacrophyllus\u003c/b\u003e (Meisn.) Meisn. in Prodr. [A.P. de Candolle] 14(2): 596 (1857) [as macrophylla]. \u0026equiv;\u003cem\u003ePasserina anthylloides\u003c/em\u003e var. \u003cem\u003emacrophylla\u003c/em\u003e Meisn. in Linnaea 14(5): 393 (1841). \u0026mdash;TYPE: South Africa, \"Zwischen Omsamwubo und Omsamcaba, auf Grasfeldern, unter 1000 Fuss, (V,b,64)\u0026rdquo; 21 Feb 1832, \u003cem\u003eDr\u0026egrave;ge 4658a\u003c/em\u003e (Syntypes: HAL barcode HAL0076214 [web!], P barcode P00541075 [web!], NY barcode NY01288019 [web!]); \u0026ldquo;Zwischen Omtendo und Omsamculo, auf Sandh\u0026uuml;geln unter 500 Fuss, (V,c,8)\u0026rdquo; Feb without year, \u003cem\u003eDr\u0026egrave;ge 4658b\u003c/em\u003e (Syntype: NY barcode NY00386301 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon burchellii\u003c/b\u003e Meisn. in Prodr. [A.P. de Candolle] 14(2): 594 (1857). \u0026equiv;\u003cem\u003eGnidia burchellii\u003c/em\u003e (Meisn.) Gilg in Engler \u0026amp; Prantl., Nat. Pflanzenfam. 3(6): 227 (1894). \u0026equiv;\u003cem\u003eGnidia burchellii\u003c/em\u003e var. \u003cem\u003evillosus\u003c/em\u003e Meisn. in Prodr. [A.P. de Candolle] 14(2): 594 (1857), \u003cb\u003enom. illeg. Superfl.\u003c/b\u003e \u0026mdash;TYPE: South Africa \u0026ldquo;between Matlowing River and Takun\u0026rdquo;, 13 Jul 1812, \u003cem\u003eBurchell 2203\u003c/em\u003e (Syntypes: G barcode G00015757 [web!], G-DC barcode G00131382 [web!], GH barcode GH00443740 [web!], GOET barcode GOET002835 [web!], GRA barcode GRA0027039-0 [web!], K barcodes K000324212 [web!] and K000324211 [web!], M barcode M0106537 [web!], NY barcodes NY01287757 [web!] and NY01287758 [web!], P barcode P00541085 [web!], PRE barcodes PRE0587576-0 [web!] and PRE0307854-0 [web!], S barcode S11-24304 [web!]); South Africa, \u0026ldquo;Ad pedem montium Stormbergen\u0026rdquo;, Sept without year, \u003cem\u003eEcklon \u0026amp; Zeyher s.n.\u003c/em\u003e (Syntypes: W barcodes W1889-0280784 [web!] and W1889-0280828 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon caffra\u003c/b\u003e Meisn. in Prodr. [A. P. de Candolle] 14(2): 593 (1857) [as \u003cem\u003eLasiosiphon caffer\u003c/em\u003e]. \u0026equiv;\u003cem\u003eGnidia caffra\u003c/em\u003e (Meisn.) Gilg in Veg. Erde [Engler] 9(3, 2): 634 (1921). \u0026mdash;TYPE: South Africa, Gauteng, \u0026ldquo;In Caffrariae monte Magalisberg\u0026rdquo;, without date, \u003cem\u003eZeyher 1488\u003c/em\u003e (Holotype: NY01288012 [web!], isotypes: K barcodes K000324205 [web!] and K000324206 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon calocephalus\u003c/b\u003e (Meisn.) Domke in Biblioth. Bot. 27(111): 95. 1934. \u0026equiv;\u003cem\u003ePasserina calocephala\u003c/em\u003e Meisn. in Linnaea 14(5): 393. 1841. \u0026equiv;\u003cem\u003eArthrosolen calocephalus\u003c/em\u003e (Meisn.) C.A.Mey. in Bull. Cl. Phys.-Math. Acad. Imp. Sci. Saint-P\u0026eacute;tersbourg 1: 359. 1843. \u0026equiv;\u003cem\u003eGnidia calocephala\u003c/em\u003e (Meisn.) Gilg in Engler \u0026amp; Prantl., Nat. Pflanzenfam. 3(6a): 228. 1894. \u0026mdash;TYPE: South Africa, \u0026ldquo;inter Omsamwubo et magnam cataractam alt. 1000\u0026ndash;1500 ped. (V. b.)\u0026rdquo;, 24 Feb 1832, \u003cem\u003eDr\u0026egrave;ge 4664a\u003c/em\u003e (Syntypes: NY barcode NY00386305 [web!], P barcode P00648740 [web!]); \u0026ldquo;in collibus inter Omcomas et Port Natal, infra 200 ped. alt. (V. c.)\", 24 Apr 1832, \u003cem\u003eDr\u0026egrave;ge 4664b\u003c/em\u003e (Syntypes: HAL barcode HAL0111338 [web!], HBG barcode HBG512552 [web!], NY barcode NY00386304 [web!], P barcode P00648739 [web!], S barcode S11-21273 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon canoargenteus\u003c/b\u003e C.H.Wright in Fl. Cap. (Harvey) 5(2, 1): 70 (1915) [as \u003cem\u003eLasiosiphon canoargentea\u003c/em\u003e]. \u0026equiv;\u003cem\u003eGnidia canoargentea\u003c/em\u003e (C.H.Wright) Gilg in Veg. Erde [Engler] 9(3, 2): 634 1921. \u0026mdash;TYPE: Transvaal, Lydenburg District, on the sides of mountains near Lydenburg, Jun 1874, \u003cem\u003eMcLea sub H. Bolus 3020\u003c/em\u003e (Lectotype: K barcode K000324203 [web!], designated by Burt Davy (1926: 207); SAM barcode SAM0001519-0 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon capitatus\u003c/b\u003e (L.f.) Burtt Davy in Man. Fl. Pl. Transvaal [Burtt Davy] 1: 207. 1926. \u0026equiv;\u003cem\u003eGnidia capitata\u003c/em\u003e L.f., Suppl. Pl.: 224. 1782. \u0026mdash;TYPE: South Africa, \u0026ldquo;Habitat in Cap. bonae spei\u0026rdquo; without date, \u003cem\u003eSmith 688.1\u003c/em\u003e [Lectotype: LINN-SM, designated by Rogers (\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2006\u003c/span\u003e:486)].\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon deserticola\u003c/b\u003e (Gilg) C.H.Wright in Fl. Cap. 5(2, 1): 71. 1915. \u0026equiv;\u003cem\u003eGnidia deserticola\u003c/em\u003e Gilg in Bot. Jahrb. Syst. 19(2\u0026ndash;3): 263. 1894. \u0026equiv;\u003cem\u003eArthrosolen deserticola\u003c/em\u003e (Gilg) Compton in Trans. Roy. Soc. South Africa 19(3): 300. 193. \u0026mdash;TYPE: South Africa, Carroogebiet, Hantam-Gebirge, without date, \u003cem\u003eMeyer s.n.\u003c/em\u003e (Syntype: B \u0026dagger;); Grootsriviers, Carroo\" Mar 1819, \u003cem\u003eMundt \u0026amp; Maire s.n\u003c/em\u003e (Syntypes: B\u0026dagger;, A barcode A00135116 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon dregeanus\u003c/b\u003e (Meisn.) Endl., Gen. Pl. [Endlicher] Suppl. 4(2): 67. 1848. \u0026equiv;\u003cem\u003eGnidia dregeana\u003c/em\u003e Meisn. in Linnaea 14(5): 426. 1841. \u0026mdash;TYPE: South Africa, Prope Port Natal (V. c.), Apr 1832, \u003cem\u003eDr\u0026egrave;ge 4661\u003c/em\u003e (Holotype: NY barcode NY01287771 [web!]; isotype: P barcode P00541064 [web!).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon esterhuyseniae\u003c/b\u003e Magee \u0026amp; J.C.Manning in Bothalia 72(1): 2 (2017). \u0026mdash;TYPE: South Africa, Northern Cape, Hay Div., Floradale, Apr 1940, \u003cem\u003eE. Esterhuysen 2315\u003c/em\u003e (Holotype: NBG!)\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon kraussianus\u003c/b\u003e (Meisn.) Burtt Davy, Man. Fl. Pl. Transvaal [Burtt Davy] 1: 207 (1926). \u0026equiv;\u003cem\u003eGnidia kraussiana\u003c/em\u003e Meisn. in London J. Bot. 2: 552 (err. typ. 452) (1843). \u0026equiv;\u003cem\u003eLasiosiphon kraussii\u003c/em\u003e Meisn. in Candolle, Prodr. [A. P. de Candolle] 14(2): 596 (1857), \u003cb\u003enom. illeg. superfl. pro\u003c/b\u003e \u003cem\u003eGnidia kraussiana\u003c/em\u003e Meisn. (1843). \u0026mdash;Type: South Africa, KwaZulu-Natal, \"Ad latera montium tabularium prope Port Natal (V.c.)\u0026rdquo;, Sept 1839, \u003cem\u003eKrauss 455\u003c/em\u003e (Holotype: NY barcode NY01287756 [web!], isotypes: FI barcode FI000346 [web!], K barcodes K000324197 [web!] and K000324198 [web!], P barcode P00541092 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eNomenclatural note\u003c/b\u003e: \u003cem\u003eGnidia kraussiana\u003c/em\u003e Meisn. (1843) was described based on a specimen collected by the German scientist and traveler C.F. Krauss. Meisner (1857) subsequently changed the epithet from the adjectival form \u003cem\u003ekraussiana\u003c/em\u003e to the genitive form \u003cem\u003ekraussii\u003c/em\u003e when he transferred the species to \u003cem\u003eLasiosiphon\u003c/em\u003e, citing \u003cem\u003eGnidia kraussiana\u003c/em\u003e Meisn. in synonymy. This is not a correctable error (Turland et al., 2018: ICN Art. 60.1), and the name \u003cem\u003eLasiosiphon kraussii\u003c/em\u003e must therefore be regarded as an illegitimate superfluous name for \u003cem\u003eGnidia kraussiana\u003c/em\u003e (Turland et al., 2018: ICN Art. 52.1). The valid transfer of the name \u003cem\u003eGnidia kraussiana\u003c/em\u003e to \u003cem\u003eLasiosiphon\u003c/em\u003e must date from the first publication of the combination \u003cem\u003eLasiosiphon kraussianus\u003c/em\u003e by Burtt-Davy (\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e1926\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon kuntzei\u003c/b\u003e (Gilg) R.Kolokoto \u0026amp; Magee in Bothalia 49(1): 3. 2019. \u0026equiv;\u003cem\u003eGnidia kuntzei\u003c/em\u003e Gilg in Revis. Gen. Pl. 3(3): 280. 1898. \u0026mdash;TYPE: South Africa, Eastern Cape, \u0026ldquo;Middelburg Road, Capland\u0026rdquo;, 17 Feb 1894, \u003cem\u003eKuntze s.n\u003c/em\u003e. (Holotype: NY barcode NY01287798 [web!]; isotypes: GH barcode GH00443746 [web!], US barcode US1269408 [web!], Z barcode Z28526 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon macropetalus\u003c/b\u003e (Meisn.) Meisn. in Prodr. [A. P. de Candolle] 14(2): 594. 1857. \u0026equiv;\u003cem\u003eGnidia macropetala\u003c/em\u003e Meisn. in London J. Bot. 2: 553 (err. typ. 453). 1843. \u0026mdash;TYPE: South Africa, \u0026ldquo;in graminosis ad sylvarum margines ubique circa Port Natal (V.c.)\u0026rdquo;, 1 Jun 1839, \u003cem\u003eKrauss 237\u003c/em\u003e (Holotype: NY01287796 [web!]; isotypes: G barcode G00015749 [web!], FI barcode FI000345 [web!], K barcode K000324187 [web!], M barcode M0106560 [web!], P barcode P00648778 [web!], PRE barcode PRE0307855-0 [web!], TUB barcodes TUB002871 [web!] and TUB002872 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon meisnerianus\u003c/b\u003e Endl., Gen. Pl. [Endlicher] Suppl. 4(2): 67. 1848. \u0026equiv;\u003cem\u003eGnidia cuneata\u003c/em\u003e Meisn. in Linnaea 14(5): 427. 1841, as nom. nov. [non \u003cem\u003eLasiosiphon cuneatus\u003c/em\u003e Decne (1844)]. \u0026mdash;TYPE: South Africa, \u0026ldquo;Cap de Bonne Esp\u0026eacute;rence\u0026rdquo; [Cape of good hope], 1828, \u003cem\u003eZehyer 86\u003c/em\u003e (Holotype: MPU [n.v.], isotypes: G barcode G00190999 [web!], M barcode M0145888 [web!], P barcode P00541094 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon meisnerianus\u003c/b\u003e var. \u003cb\u003eangustifolius\u003c/b\u003e (Meisn.) Meisn. in Prodr. [A.P. de Candolle] 14(2): 594. 1857. \u0026equiv;\u003cem\u003eGnidia cuneata\u003c/em\u003e var. \u003cem\u003eangustifolia\u003c/em\u003e Meisn. in Linnaea 14(5): 427. 184. \u0026mdash;TYPE: South Africa, \"In collibus lapidosis ad Klein Vischrivier, alt. 2000 ped. (II.a.)\", 13 Oct 1829, \u003cem\u003eDr\u0026egrave;ge 2336\u003c/em\u003e, (Syntypes: HAL barcode HAL0111685 [web!], P barcode P00541077 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon meisnerianus\u003c/b\u003e var. \u003cb\u003espathulatus\u003c/b\u003e (Meisn.) Meisn. in Prodr. [A.P. de Candolle] 14(2): 594. 1857. \u0026equiv;\u003cem\u003eGnidia cuneata\u003c/em\u003e var. \u003cem\u003espathulate\u003c/em\u003e Meisn. in Linnaea 14(5): 427. 1841. \u0026mdash;TYPE: South Africa, \"In collibus ad Vischrivier prope Trompetersdrift, alt. 600\u0026ndash;800 ped. (V. a.)\", without date, \u003cem\u003eDr\u0026egrave;ge 4657\u003c/em\u003e (Syntype: NY barcode NY01287755 [web!]); \u003cem\u003eDr\u0026egrave;ge 7351\u003c/em\u003e (Syntype: P [n.v.]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon microcephalus\u003c/b\u003e (Meisn.) J.C.Manning \u0026amp; Magee in Kew Bulletin 78(1): 12. 2017. \u0026equiv;\u003cem\u003eGnidia microcephala\u003c/em\u003e Meisn. in Prodr. [A.P. de Candolle] 14(2): 589. 1857. \u0026equiv;\u003cem\u003eArthrosolen microcephalus\u003c/em\u003e (Meisn.) E.Phillips in J. S. African Bot. 10: 63. 1944, \u003cb\u003enom. illeg.\u003c/b\u003e, non \u003cem\u003eA. microcephalus\u003c/em\u003e S.Moore in J. Bot. 57: 116. 1919. \u0026mdash;TYPE: South Africa, Gauteng, \u0026ldquo;monte Magalisberg\u0026rdquo;, without date, \u003cem\u003eZeyher 1492\u003c/em\u003e (Syntypes: G barcodes G00165832 [web!] and G00190996 [web!], G-DC barcodes GDC00162612/1 [web!] and GDC00162612/2 [web!], GRA barcode GRA0027036 [web!], K barcode K000322515 [web!], P barcodes P00541080 [web!] and P00541081 [web!], P00541079 [web!], S barcode S11-24462 [web!], SAM [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon microphyllus\u003c/b\u003e (Meisn.) Meisn. in Prodr. [A.P. de Candolle] 14: 593 (1857). \u0026equiv;\u003cem\u003eGnidia microphylla\u003c/em\u003e Meisn. in Linnaea 14(5): 432. 184. \u0026mdash;TYPE: South Africa, Northern Cape, \u0026ldquo;in collibus siccis prope Kuigunjels ad fl. Garip, alt. 200 ped.\u0026rdquo; [dry hills near Kuigunjels, near the mouth of the Orange River, 200 feet], without date, \u003cem\u003eDr\u0026egrave;ge 2976a\u003c/em\u003e (Holotype: NY barcode NY01288010 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon pedunculatus\u003c/b\u003e (Beyers) J.C.Manning \u0026amp; Boatwr. in Bothalia 43(1): 88. 2013. \u0026equiv;\u003cem\u003eGnidia pedunculata\u003c/em\u003e Beyers in Bothalia 32: 79. 2002. \u0026mdash;TYPE: South Africa, Western Cape, Knersvlakte, Olifants River Settlement 316, near eastern border, 6 Aug 1993, \u003cem\u003eLe Roux \u0026amp; Hilton-Taylor 27\u003c/em\u003e (Holotype: NBG barcode NBG0199291-0 [web!]; isotypes: K barcode K000322601 [web!], PRE barcode PRE0569222-0 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon polyanthus\u003c/b\u003e (Gilg) C.H.Wright in Fl. Cap. (Harvey) 5(2, 1): 74. 1915. \u0026equiv;\u003cem\u003eGnidia polyantha\u003c/em\u003e Gilg in Bot. Jahrb. Syst. 19(2): 265. 1894. \u0026mdash;TYPE: South Africa, Eastern Cape, \u0026ldquo;Saddle between Iggakancu \u0026amp; Bazeia Mountains\u0026rdquo;, Jun without year, \u003cem\u003eBaur 646\u003c/em\u003e (Holotype: K✝; isotypes: K barcodes K000324183 [web!] and K000324184 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon polyanthus\u003c/b\u003e var. \u003cb\u003emicrocalyx\u003c/b\u003e Burtt Davy, Man. Pl. Transvaal [Burtt Davy] 1: 206. 1926. \u0026mdash;TYPE: South Africa, \"Pietersberg Distr., Marovuni, infrequens\", without date, \u003cem\u003eJunod 145\u003c/em\u003e (Holotype: K [n.v.]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon polycephalus\u003c/b\u003e (E.Mey. ex Meisn.) H.Pearson in Fl. Trop. Afr. 6(1, 2): 228. 1910. \u0026equiv;\u003cem\u003ePasserina polycephala\u003c/em\u003e E.Mey. ex Meisn. in Linnaea 14(5): 390. 1841. \u0026equiv;\u003cem\u003eArthrosolen polycephalus\u003c/em\u003e (E.Mey. ex Meisn.) C.A.Mey. in Bull. Cl. Phys.-Math. Acad. Imp. Sci. Saint-P\u0026eacute;tersbourg 1: 359. 1843. \u0026equiv;\u003cem\u003eGnidia polycephala\u003c/em\u003e (E.Mey. ex Meisn.) Gilg in Engler \u0026amp; Prantl., Nat. Pflanzenfam. 3(6a): 227. 1894. \u0026mdash;TYPE: South Africa, \u0026ldquo;Locis depressis argillosis inter Poortje et Nieuwkerkshoogte, alt. 4000\u0026ndash;5000 ped. (I.c.)\u0026rdquo; 10 Sept 1829, \u003cem\u003eDr\u0026egrave;ge 724\u003c/em\u003e (Syntypes: HAL barcode HAL0111341 [web!], HBG barcode HBG512593 [web!], HEID barcode HEID702658 [web!], JE barcode JE00019704 [web!], NY barcode NY01104569 [web!], P barcode TUB barcode TUB002886 [web!], W barcode W 1889-0282097 [web!]); \u0026ldquo;Nieuweveld, alt. 3000\u0026ndash;4000 ped. (l.d.)\u0026rdquo;, 24 Oct 1826, \u003cem\u003eDr\u0026egrave;ge s.n.\u003c/em\u003e (Syntypes: HAL barcode HAL0111340 [web!], HBG barcode HBG512653 [web!], MO barcode MO2302117 [web!], P00648737 [web!]); \u0026ldquo;Circa Limoenfontein in Winterveld, alt. 3000\u0026ndash;3500 ped. (I.d.)\u0026rdquo;, without date, \u003cem\u003eBurchell 2222\u003c/em\u003e (Syntypes: G-DC [n.v.], NY barcode NY01104568 [web!], W barcode W1889-0282096 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon pulchellus\u003c/b\u003e (Meisn.) Decne. in Voy. Inde [Jacquemont] 4: 149. 1844. \u0026equiv;\u003cem\u003eGnidia pulchella\u003c/em\u003e Meisn. in Linnaea 14(5): 425. 1841. \u0026equiv;\u003cem\u003eGnidia pulchella\u003c/em\u003e var. \u003cem\u003eglabrata\u003c/em\u003e (Meisn.) Meisn. in Linnaea 14(5): 425. 1841, \u003cb\u003enom. illeg. Superfl\u003c/b\u003e. \u0026mdash;Type: South Africa, \"Ad rivulum inter Omtata et Omgaziana, alt. 800 ped. (V.b.), without date, \u003cem\u003eDr\u0026egrave;ge s.n.\u003c/em\u003e (Syntype: NY [n.v.]); \"Katberg, alt. 4000\u0026ndash;4500 ped. (I.a.)\", without date, \u003cem\u003eDr\u0026egrave;ge 7354\u003c/em\u003e (Syntype: NY barcode NY01287790 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon pulchellus\u003c/b\u003e var. \u003cb\u003edasyphyllus\u003c/b\u003e (Meisn.) Meisn. in Prodr. [A.P. de Candolle] 14(2): 595. 1857. \u0026equiv;\u003cem\u003eGnidia pulchella\u003c/em\u003e var. \u003cem\u003edasyphylla\u003c/em\u003e Meisn. in Linnaea 14(5): 425. 184. \u0026mdash;TYPE: South Africa, \"in collibus siccis prope Zwartbulletje, 2500 ped.\", without date, \u003cem\u003eDr\u0026egrave;ge 2976b\u003c/em\u003e (Syntype: NY barcode NY01288029 [web!]); \u0026ldquo;ad Zondagrivier prope Blaauwekrans, alt. 1700 ped. (II. c.)\u0026rdquo; without date, \u003cem\u003eDr\u0026egrave;ge s.n.\u003c/em\u003e (Syntype: NY [n.v.]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon rigidus\u003c/b\u003e J.C.Manning \u0026amp; Boatwr. in Bothalia 43(1): 85. 2013. \u0026mdash;TYPE: South Africa, Northern Cape, Tankwa [Tanqua] Karoo National Park, SW foot of Leeuberg, along drainage lines, 20 Jun 2012, \u003cem\u003eManning 3363\u003c/em\u003e (Holotype: NBG barcode NBG0279875-0 [web!]; isotypes: K [n.v.], MO [n.v.], PRE!).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon robustus\u003c/b\u003e (B.Peterson) Olaniyan \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia robusta\u003c/em\u003e B.Peterson in Bothalia 14: 82. 1982. \u0026equiv;\u003cem\u003eLasiosiphon nanus\u003c/em\u003e Burtt Davy, Man. Pl. Transvaal 1: 207. 1926 as nom. nov. [non \u003cem\u003eG. nana\u003c/em\u003e (L. f.) Wikstr.] \u0026mdash;TYPE: Swaziland, Hlatikulu, without date., \u003cem\u003eM. Stewart s.n\u003c/em\u003e. (Holotype: K barcode K000322535 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon rubescens\u003c/b\u003e (B.Peterson) J.C.Manning \u0026amp; Magee in Kew Bull. 78(1): 12. 2017. \u0026equiv;\u003cem\u003eGnidia rubescens\u003c/em\u003e B.Peterson in Bot. Not. 119(2): 345. 1966. \u0026mdash;TYPE: South Africa, Limpopo, \u0026ldquo;Transvaal - Near Mara, about 18 miles from Louis Trichardt on Mara-Vivo road\u0026rdquo;, 3 Apr 1957, \u003cem\u003eMeeuse 10196\u003c/em\u003e (Holotype: LD barcode LD1001442 [web!]; isotypes: BOL barcode BOL136956 [web!], BR barcode BR0000006249892 [web!], EA barcode EA000002189 [web!], G barcode G00165800 [web!], GB barcodes GB0048776 [web!] and GB0048777 [web!], LMA barcode LMA0031290-0 [web!], M barcode M0106561 [web!], MO barcode MO313015 [web!], PRE barcodes PRE0587577-0 [web!] and PRE0297723-0 [web!], S barcode S05-9781 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon sericocephalus\u003c/b\u003e (Meisn.) J.C.Manning \u0026amp; Boatwr. in Bothalia 43(1): 88. 2013. \u0026equiv;\u003cem\u003eArthrosolen sericocephalus\u003c/em\u003e Meisn. in Prodr. [A.P. de Candolle]14(2): 561. 1857. \u0026equiv;\u003cem\u003eGnidia sericocephala\u003c/em\u003e (Meisn.) Gilg ex Engl., Veg. Erde [Engler] 9(3, 2): 634. 192. \u0026mdash;TYPE: South Africa, \u0026ldquo;In mont. Macalisberg Caffrariae\u0026rdquo;, \u003cem\u003eZeyher 1494\u003c/em\u003e (Holotype: G-DC; Isotype: K barcode K000324226 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon splendens\u003c/b\u003e (Meisn.) Endl., Gen. Pl. [Endlicher] Suppl. 4(2): 67. 1848. \u0026equiv;\u003cem\u003eGnidia splendens\u003c/em\u003e Meisn. in Linnaea 14(5): 428. 1841. \u0026mdash;TYPE: South Africa, Kwazulu Natal, \"In herbosis prope Port Natal, infra alt. 100 ped. (V. c. 33)\", without date, \u003cem\u003eDr\u0026egrave;ge 4656\u003c/em\u003e (Holotype: NY barcode NY01287785 [web!]; isotypes: HAL barcode HAL0111548 [web!], HBG barcode HBG512608 [web!], K barcode K000324192 [web!], P barcode P00541044 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon suavissimus\u003c/b\u003e (Dinter) Domke in Biblioth. Bot. 27(111): 91. 1934. \u0026equiv;\u003cem\u003eGnidia suavissima\u003c/em\u003e Dinter in Repert. Spec. Nov. Regni Veg. 16: 340. 1920.\u0026mdash;TYPE: Namibia, \"Namaland: Zwischen Angras Juntas und Buntfeldschuh\", Jul 1913, \u003cem\u003eSch\u0026auml;fer 504\u003c/em\u003e (Lectotype: GLM barcode GLM101451 [web!], designated by Otte et al. (2011:258); isolectotype: B barcode B 10 0349157 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon triplinervis\u003c/b\u003e (Meisn.) Decne. in Voy. Inde [Jacquemont] 4: 149. 1844. \u0026equiv;\u003cem\u003eGnidia triplinervis\u003c/em\u003e Meisn. in Linnaea 14(5): 429. 184. \u0026mdash;TYPE: \"Inter frutices in saxosa ripa fl. Omtendo infra alt. 100 ped. (V. c.)\u0026rdquo;, without date, \u003cem\u003eDr\u0026egrave;ge 4662\u003c/em\u003e (Holotype: NY barcode NY01287803 [web!]; isotype: HAL barcode HAL0111551 [web!], TUB barcodes TUB002883 [web!] and TUB002883 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eSpecies from Madagascar\u003c/b\u003e:\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon ambondrombensis\u003c/b\u003e Boiteau in Bull. Acad. Malgache, n.s., 24: 83. 1941 [as ambondrombense]. \u0026equiv;\u003cem\u003eGnidia ambondrombensis\u003c/em\u003e (Boiteau) Z.S.Rogers in Ann. Miss. Bot. Gard. 96(1): 339. 2009. \u0026mdash;TYPE: Madagascar, Fianarantsoa, Mt. Ambondrombe, rocky summit, 1900 m, 11 Apr 1941, \u003cem\u003eP. Boiteau\u003c/em\u003e (Hb. Jard. Bot. Tananarive) \u003cem\u003e4643\u003c/em\u003e (Holotype: P barcode P00370331 [web!], isotypes: MO barcode MO1181707 [web!], TAN).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon bojerianus\u003c/b\u003e Decne., Voy. Inde [Jacquemont] 4: 149. 1844. \u0026equiv;\u003cem\u003eGnidia bojeriana\u003c/em\u003e (Decne.) Gilg in Engler \u0026amp; Prantl., Nat. Pflanzenfam. 3(6a): 228. 1894. \u0026mdash;TYPE: Madagascar, Antananarivo, Emirnae mtns., without date, \u003cem\u003eW. Bojer s.n.\u003c/em\u003e (Lectotype: P barcode P00370315 [web!], designated by Rogers (\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2009\u003c/span\u003e:342); isolectotypes: BM [n.v.], G barcode G00131578 [web!], P barcodes P00370314 [web!] and P00370316 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon danguyanus\u003c/b\u003e (Leandri) Boatwr. \u0026amp; J.C.Manning in S. African J. Bot. 112: 337. 2017. \u0026equiv;\u003cem\u003eGnidia danguyana\u003c/em\u003e Leandri, Bull. Soc. Bot. France 77: 35 (1930). \u0026mdash;TYPE: Madagascar, Toamasina, Tampina forest, Dec 1923, \u003cem\u003eM. Louvel 118\u003c/em\u003e (Lectotype: P barcode P00380376 [web!], designated by Rogers (\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2009\u003c/span\u003e:342).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon daphnifolius\u003c/b\u003e (L.f.) Boatwr. \u0026amp; J.C.Manning in S. African J. Bot. 112: 337. 2017. \u0026equiv;\u003cem\u003eGnidia daphnifolia\u003c/em\u003e L.f., Suppl. Pl. 225 (1782) [as daphnaefolia]. \u0026equiv;\u003cem\u003eGnidia daphnifolia\u003c/em\u003e var. \u003cem\u003eglabra\u003c/em\u003e L.f., Suppl. Pl. 225 (1782). \u0026equiv;\u003cem\u003eDessenia daphnifolia\u003c/em\u003e (L.f.) Raf. in Fl. Tellur. 4: 106. 1838 [as daphnefolia]. \u0026mdash;TYPE: Madagascar, \u003cem\u003eHerb. Smith 688.5\u003c/em\u003e (Lectotype: LINN-SM, designated by Rogers (\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2006\u003c/span\u003e:486).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon gilbertae\u003c/b\u003e (Drake) Boatwr. \u0026amp; J.C.Manning in S. African J. Bot. 112: 337. 2017. \u0026equiv;\u003cem\u003eGnidia gilbertae\u003c/em\u003e Drake in Bull. Mens. Soc. Linn. Paris 2: 1218. 1896. \u0026mdash;TYPE: Madagascar, Mahajanga, \u0026lsquo;Madounga et Antsalahanki\u0026rsquo;, 1876, \u003cem\u003eA. Grandidier s.n\u003c/em\u003e. (Holotype: P barcode P00380410 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon gnidioides\u003c/b\u003e (Baker) Boatwr. \u0026amp; J.C.Manning in S. African J. Bot. 112: 337. 2017. \u0026equiv;\u003cem\u003eDais gnidioides\u003c/em\u003e Baker in J. Linn. Soc. Bot 20: 244 (1883). \u0026equiv;\u003cem\u003eArthrosolen gnidioides\u003c/em\u003e (Baker) Leandri in Bull. Soc. Bot. France 76: 1043 (1929). \u0026equiv;\u003cem\u003eGnidia gnidioides\u003c/em\u003e (Baker) Domke in Biblioth. Bot 27(111): 46 (1934). \u0026mdash;TYPE: Madagascar, Antananarivo, \u0026lsquo;grassy hills of the province of Imerina\u0026rsquo;, without date, \u003cem\u003eR. Baron 2061\u003c/em\u003e (Lectotype: K barcode K000324220 [web!], designated by Rogers (\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2009\u003c/span\u003e:351); isolectotype: P00380432 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon hibbertioides\u003c/b\u003e S.Moore in J. Bot. 58: 189 (1920). \u0026equiv;\u003cem\u003eGnidia hibbertioides\u003c/em\u003e (S.Moore) Z.S.Rogers in Ann. Miss. Bot. Gard. 96(1): 349. 2009. \u0026mdash;TYPE: Madagascar, without date, J. \u003cem\u003eThompson \u0026amp; J. Forbes s.n\u003c/em\u003e. (Holotype: BM barcode BM000647723 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon humbertii\u003c/b\u003e Leandri in Bull. Soc. Bot. France 76: 1039. 1929. \u0026equiv;\u003cem\u003eGnidia humbertii\u003c/em\u003e (Leandri) Z.S.Rogers in Ann. Miss. Bot. Gard. 96(1): 351. 2009. \u0026mdash;TYPE: Madagascar, Fianarantsoa, \u0026lsquo;Isalo, mouth of Sakamarekely \u0026amp; Sambalinieto rivers, 500\u0026ndash;1000 m, 19 Oct 1924, \u003cem\u003eH. Humbert 2844\u003c/em\u003e (Lectotype: P barcode P00373493 [web!], designated by Rogers (\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2009\u003c/span\u003e:351); isolectotype: G barcode G00010149 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon leandrianus\u003c/b\u003e Boatwr. \u0026amp; J.C.Manning in S. African J. Bot. 112: 337. 2017. \u0026equiv;\u003cem\u003eGnidia decaryana\u003c/em\u003e Leandri in Bull. Mus. Natl. Hist. Nat. s\u0026eacute;r. 2(1): 436 (1929) as nom. nov. [non \u003cem\u003eLasiosiphon decaryi\u003c/em\u003e Leandri]. \u0026mdash;TYPE: Madagascar, Toliara, \u0026lsquo;Fort-Dauphin\u0026rsquo;, 3 Jul 1926, \u003cem\u003eR. Decary 4332\u003c/em\u003e (Holotype: P barcode P00373426 [web!], isotypes: P barcode P00380392, [web!] TAN [n.v.]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon linearis\u003c/b\u003e Leandri in Bull. Soc. Bot. France 76: 1040. 1930. \u0026equiv;\u003cem\u003eLasiosiphon decaryi\u003c/em\u003e var. \u003cem\u003elinearis\u003c/em\u003e (Leandri) Leandri in Bull. Mus. Natl. Hist. Nat. s\u0026eacute;r. 2(3): 154. 1931. \u0026equiv;\u003cem\u003eGnidia linearis\u003c/em\u003e (Leandri) Z.S.Rogers in Ann. Miss. Bot. Gard. 96(1): 353. 2009. \u0026mdash;TYPE: Madagascar, [Toliara/Fianarantsoa], savanna betw. Bemketa [Bereketa] \u0026amp; Malio, 15 Jun 1923, \u003cem\u003eH. Poisson 692\u003c/em\u003e (Holotype: P barcode P00373448 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon occidentalis\u003c/b\u003e Leandri in Notul. Syst. (Paris) 13: 47. 1947. \u0026equiv;\u003cem\u003eGnidia occidentalis\u003c/em\u003e (Leandri) Z.S.Rogers in Ann. Miss. Bot. Gard. 96(1): 358. 2009. \u0026mdash;TYPE: Madagascar, Mahajanga, Kamakama forest, Ankara plateau, 14 Jul 1901, H. Perrier de la \u003cem\u003eB\u0026acirc;thie 1276\u003c/em\u003e (Lectotype: P barcode P00373473 [web!], designated by Rogers (\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2009\u003c/span\u003e:358).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon perrieri\u003c/b\u003e Leandri in Notul. Syst. (Paris) 13: 49. 1947. \u0026equiv;\u003cem\u003eGnidia perrieri\u003c/em\u003e (Leandri) Z.S.Rogers in Ann. Miss. Bot. Gard. 96(1): 360. 2009. \u0026mdash;TYPE: Madagascar, Fianarantsoa, Andringitra Massif (Iratsy), valley of Riambava \u0026amp; Antsifotra, 2000\u0026ndash;2500 m, 27 Nov 1924, \u003cem\u003eH. Humbert 3827\u003c/em\u003e (Lectotype: P barcode P00380352 [web!], designated by Rogers (\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2009\u003c/span\u003e:360); isolectotypes: BM barcode BM000812676 [web!], K barcode K000634424 [web!], G barcode G00010144 [web!], K barcode K000634424 [web!], MO barcode MO1181820, TAN barcode TAN000732 [web!], US barcode US00956206 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon razakamalalanus\u003c/b\u003e (Z.S.Rogers) Boatwr. \u0026amp; J.C.Manning in S. African J. Bot. 112: 337. 2017. \u0026equiv;\u003cem\u003eGnidia razakamalalana\u003c/em\u003e Z.S.Rogers in Adansonia s\u0026eacute;r. 3(28): 156. 2006. \u0026mdash;TYPE: Madagascar, Toliara, Fivondronona Fort-Dauphin, Ivohibe Forest, 112 m, 29 Nov 2005, \u003cem\u003eR. Razakamalala, E. Ramisy and B. Mara 2670\u003c/em\u003e (Holotype: MO barcode MO1287062 [web!], isotypes: G barcode G00424957 [web!], K barcode K001089638 [web!], P barcode P02141588 [web!], TAN).\u003c/p\u003e \u003cp\u003e \u003cb\u003eSpecies from Asia (Arabia, India and Sri Lanka)\u003c/b\u003e:\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon glaucus\u003c/b\u003e Fresen. in Flora 21(2): 603. 1838. \u0026equiv;\u003cem\u003eGnidia glauca\u003c/em\u003e (Fresen) Gilg in Engler. \u0026amp; Prantl., Nat. Pflanzenfam. 3(6a): 227. 1894. \u0026mdash;TYPE: Ethiopia, \u0026ldquo;Abyssinia\u0026rdquo;, \u003cem\u003eRueppel s.n\u003c/em\u003e. (Holotype: FR barcode FR0030066 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon sisparensis\u003c/b\u003e (Gardner) Meisn. in DC. Prodr. [A. P. de Candolle] 14(2): 598. 1857. \u0026equiv;\u003cem\u003eLasiosiphon eriocephalus\u003c/em\u003e var. \u003cem\u003esisparensis\u003c/em\u003e (Gardner) Gamble in Fl. Madras 2(7): 1244. 1925. \u0026equiv;\u003cem\u003eGnidia sisparensis\u003c/em\u003e Gardner in J. Nat. Hist. 7: 457. 1847. \u0026equiv;\u003cem\u003eGnidia glauca\u003c/em\u003e var. \u003cem\u003esisparensis\u003c/em\u003e (Gardner) Kumari in Fl. Tamil Nadu 1(2): 214. 1987. \u0026mdash; TYPE: India, Tamil Nadu: Neilgherry (Nilgiri) mountains, Sispara, without date, \u003cem\u003eGardner s.n.\u003c/em\u003e (Lectotype: K barcode K000357981 [web!], designated by Prabhukumar et al. (\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2018\u003c/span\u003e:180); isolectotype: K barcode K000357982 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon socotranus\u003c/b\u003e Balf.f. in Proc. Roy. Soc. Edinburgh 12(113): 92. 1884. \u0026equiv;\u003cem\u003eGnidia socotrana\u003c/em\u003e (Balf.f.) Gilg in Engler \u0026amp; Prantl., Nat. Pflanzenfam. 3(6a): 228. 1894. \u0026mdash;TYPE: Yemen, Socotra, Feb-Mar 1880, \u003cem\u003eBalfour 518\u003c/em\u003e (Lectotype: K barcode K000322641 [web!], designated by Gastaldo (\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1969\u003c/span\u003e:369); isolectotypes: BM barcode BM000910837 [web!], EA barcode E00239397 [web!], E barcodes E00239400 [web!] and E00239402 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon somalensis\u003c/b\u003e (Franch) H.Pearson in Fl. Trop. Afr. 6(1, 2): 229. 1910. \u0026equiv;\u003cem\u003eArthrosolen somalensis\u003c/em\u003e Franch. in Sert. Somal.: 62 (t. 6). 1882. \u0026equiv;\u003cem\u003eGnidia somalensis\u003c/em\u003e (Franch) Gilg in Ann. 1st. Bot. Roma, 6: 98. 1897. \u0026mdash;TYPE: Somalia, \u0026ldquo;Ouanentab, plateau de Yaffer\u0026rdquo;, without date, \u003cem\u003eR\u0026eacute;voil 123\u003c/em\u003e (Holotype: P barcode P00541095 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eSpecies from tropical Africa\u003c/b\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon apiculatus\u003c/b\u003e (Oliv.) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia involucrata\u003c/em\u003e var. \u003cem\u003eapiculata\u003c/em\u003e Oliv. in Trans. Linn. Soc. London 29(3): 143, pl. 91. 1875. \u0026equiv;\u003cem\u003eGnidia apiculata\u003c/em\u003e (Oliv.) Gilg in Bot. Jahrb. Syst. 19(2\u0026ndash;3): 263. 1894. \u0026mdash;TYPE: Sudan, \u0026ldquo;Bare heights of Madi\u0026rdquo;, Dec 1862, \u003cem\u003eSpeke \u0026amp; Grant s.n\u003c/em\u003e. (Holotype: K barcode K000322646 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon apiculatus\u003c/b\u003e fo. \u003cb\u003epyramidalis\u003c/b\u003e (Aymonin) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia apiculata\u003c/em\u003e fo. \u003cem\u003epyramidalis\u003c/em\u003e Aymonin in Fl. Gabon 11: 95, pl. 14, f. 6. 1966. \u0026mdash;TYPE: Central African Republic, Oubangui, 60 Km North of Bambari, 5 Oct 1921, \u003cem\u003eTisserant 307\u003c/em\u003e (Holotype: BM barcode BM000910854 [web!], isotype: P barcode P00541078 [web!])\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon bambutanus\u003c/b\u003e (Gilg \u0026amp; Ledermann ex Engl.) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia bambutana\u003c/em\u003e Gilg \u0026amp; Ledermann ex Engl. in Veg., Erde [Engler] 9(3, 2): 631. 1921. \u0026mdash;TYPE: Cameroon \u0026ldquo;In den Babutubergen an der Grenze der sudansichen Parksteppenprovinz\u0026rdquo;, without date, \u003cem\u003eLederman s.n\u003c/em\u003e (Holotype: B\u0026dagger;); Cameroon, Bangou [Bangan], Dec 1938, \u003cem\u003eH. Jacques-F\u0026eacute;lix 2855\u003c/em\u003e (Neotype: P00541086 [web!], designated by Aymonin (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1965\u003c/span\u003e:324).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon baumianus\u003c/b\u003e (Gilg) Olaniyan, \u003cem\u003ecomb.nov.\u003c/em\u003e \u0026equiv;\u003cem\u003eGnidia baumiana\u003c/em\u003e Gilg in Kunene-Sambesi-Exped.: 311. 1903. \u0026mdash;TYPE: Angola, \u0026ldquo;Chitanda bei Kasinga\u0026rdquo;, Oct 1899, \u003cem\u003eBaum 211\u003c/em\u003e (Holotype: B\u0026dagger;, isotypes: G barcode G00165820 [web!], HBG barcode HBG512557 [web!], M barcode M0106538 [web!], S barcodes S10-25321 [web!] and S10-25326 [web!], W barcode W19010009368 [web!])\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon capitatus\u003c/b\u003e (L.f.) Burtt Davy (for protologue see under southern African taxa)\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon chapmanii\u003c/b\u003e (B.Peterson) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia chapmanii\u003c/em\u003e B.Peterson in Kew Bull. 31(1): 177. 1976. \u0026mdash;TYPE: Malawi, Mlanje Mountain, 7 Jun 1970, \u003cem\u003eBrummit 11346\u003c/em\u003e (Holotype: K barcode K000049217 [web!], isotypes: EA barcode EA000002370 [web!], GB barcode GB0048775 [web!], LI barcode LISC011492 [web!], MAL, PRE barcode PRE0308076-0 [web!], SRGH [n.v.], UPS [n.v.]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon chrysanthus\u003c/b\u003e (Solms ex Schweinf.) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eArthrosolen chrysanthus\u003c/em\u003e Solms ex Schweinf. in Beitr. Fl. Aethiop.: 165 .1867. \u0026equiv;\u003cem\u003eGnidia chrysantha\u003c/em\u003e (Solms ex Schweinf.) Gilg in Bot. Jahrb. Syst. 19(2\u0026ndash;3): 258 (1894). \u0026mdash;TYPE: Ethiopia \u0026ldquo;Bei Duli im Fesoghlu\u0026rdquo;, 21 Apr 1848, \u003cem\u003eCienkowski s.n\u003c/em\u003e (Holotype: B\u0026dagger;); Ethiopia, \u0026ldquo;near Nadda\u0026rdquo;, 13 Oct 1954, \u003cem\u003eMooney 6257\u003c/em\u003e (Neotype: K barcode K000322639 [web!], designated by Gastaldo (\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1969\u003c/span\u003e:351).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon deserticola\u003c/b\u003e (Gilg) C.H. Wright (for protologue see under southern African taxa)\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon dumicola\u003c/b\u003e (S.Moore) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia dumicola\u003c/em\u003e S.Moore in J. Bot. 57: 114. 1919. \u0026mdash;TYPE: Angola, \u0026ldquo;between Forte Princeza Amelia and Limbala Monelilo\u0026rdquo;, 26 Sept, 1905, \u003cem\u003eGossweiler 2023\u003c/em\u003e (Holotype: BM barcode BM000910844 [web!], isotypes: K barcode K000322604 [web!], BR barcode BR0000006299842 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon elkerensis\u003c/b\u003e (Friis \u0026amp; Sebsebe) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia elkerensis\u003c/em\u003e Friis \u0026amp; Sebsebe in Webbia 73(2): 214. 2018. \u0026mdash;TYPE: Ethiopia, Somalia Region \u0026ldquo;Afder Zone, El Kere Woreda\u0026rdquo;, 30 Dec 2016, \u003cem\u003eMelaku Wondafrash, Shewangziwe Lemma, Wege Abebe, Shambel Alemu \u0026amp; Elias Tadesse 4182\u003c/em\u003e (Holotype: ETH [n.v.], isotypes: C barcode C10022126 [web!], K [n.v.]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon eminii\u003c/b\u003e (Engl. \u0026amp; Gilg) H.Pearson in Fl. Trop. Afr. 6(1, 2): 229. 1910 [as \u0026ldquo;\u003cem\u003eemini\u003c/em\u003e\u0026rdquo;]. \u0026equiv;\u003cem\u003eGnidia eminii\u003c/em\u003e Engl. \u0026amp; Gilg in Bot. Jahrb. Syst. 19(2\u0026ndash;3): 265. 1894 [as \u0026ldquo;\u003cem\u003eemini\u003c/em\u003e\u0026rdquo;]. \u0026mdash;TYPE: Tanzania, \u0026ldquo;Massaisteppe, Wald westlich Jrangi\u0026rdquo;, Jun 1892, \u003cem\u003eStuhlmann 4224\u003c/em\u003e (Holotype: B\u0026dagger;).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon foliosus\u003c/b\u003e (H.Pearson) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eArthrosolen foliosus\u003c/em\u003e H.Pearson in Fl. Trop. Afr. 6(1, 2): 237. 1910 [as \u003cem\u003efoliosa\u003c/em\u003e]. \u0026equiv;\u003cem\u003eGnidia foliosa\u003c/em\u003e (H.Pearson) Engl., Veg. Erde [Engler] 9(3, 2): 633. 1921. \u0026mdash;TYPE: Sudan, \u0026ldquo;Upper Guinea. Soudan: between Bammako [Bamako] and Guignula\u0026rdquo;, 22 Jan 1899, \u003cem\u003eChevalier 256\u003c/em\u003e (Holotype: K barcode K000322627 [web!], isotypes: BR barcode BR0000006299132 [web!], P barcodes P00516687 [web!] and P00516688 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon fruticulosus\u003c/b\u003e (Gilg) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia fruticulosa\u003c/em\u003e Gilg in Bot. Jahrb. Syst. 23(1\u0026ndash;2): 207. 1896. \u0026mdash;TYPE: Angola, \u0026ldquo;Huilla\u0026rdquo; \u003cem\u003eAntunes sub De Kindt 116\u003c/em\u003e (Syntype: not located), \u003cem\u003e121\u003c/em\u003e (Syntype: not located), and \u003cem\u003e316\u003c/em\u003e (Syntypes: K barcode K000322562 [web!], LISC barcode LISC011490 [web!], P barcodes P00648774 [web!] and P00648775 [web!], W barcode W20020014719 [web!], WU barcode WU0026027 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon fulgens\u003c/b\u003e (Welw. ex Hiern) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia fulgens\u003c/em\u003e Welw. ex Hiern, Cat. Afr. Pl. 1 (4): 923. 1900 [\u003cem\u003eGnidia fulgens\u003c/em\u003e Welw., Apont. 548: 1859, nom nud.]. \u0026equiv;\u003cem\u003eArthrosolen fulgens\u003c/em\u003e (Welw. ex Hiern) H.Pearson in Fl. Trop. Afr. 6(1, 2): 237. 1910. \u0026mdash;TYPE: Angola, Pungo Andongo, at Mutollo and in marshy meadows near Sobato Nbilla, Jan-Mar 1857, \u003cem\u003eWelwitsch 6483\u003c/em\u003e (Syntypes: BM barcode BM000910843 [web!], LISU barcodes LISU221402 [web!], LISU221403 [web!] and LISU221404 [web!], M barcode M0106551 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon glaucus\u003c/b\u003e Fresen. (for protologue see under Asian taxa).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon goetzeanus\u003c/b\u003e (Gilg) Olaniyan, \u003cem\u003ecomb.nov.\u003c/em\u003e \u0026equiv;\u003cem\u003eGnidia goetzeana\u003c/em\u003e Gilg in Bot. Jahrb. Syst. 30(3\u0026ndash;4): 363. 190. \u0026mdash;TYPE: Tanzania, Njombe district,\u0026ldquo;Langenburg [Lumbira]\u0026rdquo;, May 1899, \u003cem\u003eGoetze 895\u003c/em\u003e (Lectotype: BR barcode BR0000006299101, designated by Robyns (\u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e1975\u003c/span\u003e:43).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon gossweileri\u003c/b\u003e (S.Moore) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eArthrosolen gossweileri\u003c/em\u003e S.Moore in J. Bot. 57: 116. 1919. \u0026equiv;\u003cem\u003eGnidia gossweileri\u003c/em\u003e (S.Moore) B.Peterson in Fl. Zambes. 9(3): 98. 2006. \u0026mdash;TYPE: Angola, \u0026ldquo;Munonque\u0026rdquo;, 20 Apr 1906, \u003cem\u003eGossweiler 3090\u003c/em\u003e (Holotype: BM barcode BM000910842 [web!], isotypes: K barcode K000322605 [web!], BR barcode BR0000006249830 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon gossweileri\u003c/b\u003e subsp. \u003cb\u003epetersonii\u003c/b\u003e (Verdc.) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia gossweileri\u003c/em\u003e subsp. \u003cem\u003epetersonii\u003c/em\u003e Verdc. in Fl. Zambes. 9(3): 98. 2006. \u0026mdash;TYPE: Zambia, Mwinilunga District, 18 km East of Kalene Hill, 16 Dec 1963, \u003cem\u003eRobinson 6111\u003c/em\u003e (Holotype: K barcode K000322606 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon hockii\u003c/b\u003e (De Wild) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia hockii\u003c/em\u003e De Wild in Repert. Spec. Nov. Regni Veg. 11: 536. 1913. \u0026mdash;TYPE: Democratic Republic of the Congo, \u0026ldquo;Ober-Katanga: Elisabethville\u0026rdquo;, Sept 1911 \u003cem\u003eHock s.n\u003c/em\u003e. (Holotype: BR barcode BR0000008964878 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon involucratus\u003c/b\u003e (Steud ex A.Rich) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia involucrata\u003c/em\u003e Steud. ex A.Rich. in Tent. Fl. Abyss. 2: 234. 1850. \u0026mdash;TYPE: Ethiopia, Tigre, Sana to Terrfera, 5 Oct 1838, \u003cem\u003eSchimper 770\u003c/em\u003e (Lectotype: P barcode P00541070, designated by Gastaldo (\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1969\u003c/span\u003e:347); isolectotypes: BR barcodes BR0000006249809 [web!] and BR0000006249854 [web!], BR0000006249847 [web!], G barcodes G00015753 [web!], G00015754 [web!] and G00015755 [web!], G-DC barcode GDC00131478 [web!], HAL barcode HAL0111681 [web!], K barcode K000322632 [web!], LG barcode LG0000090028878 [web!], M barcodes M0106555 [web!] and M0106556 [web!], MPU barcode MPU007261 [web!], P barcode P00648788 [web!], REG barcode REG000882 [web!], S barcode S05-9784 [web!], TUB barcodes TUB002865 [web!] and TUB002866 [web!])\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon kasaiensis\u003c/b\u003e (S.Moore) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia kasaiensis\u003c/em\u003e S.Moore in J. Bot. 57: 113. 1919. \u0026mdash;TYPE: Democratic Republic of Congo, Kasai District, Sankuru River, \u003cem\u003eK\u0026auml;ssner 3322\u003c/em\u003e (Holotype: BM barcode BM000910841 [web!], isotype: BR barcode BR0000008965530 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon kraussianus\u003c/b\u003e (Meisn.) Burtt Davy (for protologue see under Asian taxa).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon kraussianus\u003c/b\u003e var. \u003cb\u003emollissimus\u003c/b\u003e (E.A. Bruce) Olaniyan, \u003cem\u003ecomb, nov. Gnidia kraussiana\u003c/em\u003e var. \u003cem\u003emollissima\u003c/em\u003e (E.A.Bruce) A. Robyns in Fl. Afr. Centr., Spermatophytes: 53. 1975. \u003cem\u003eLasiosiphon mollissimus\u003c/em\u003e E.A.Bruce in Bull. Misc. Inform. Kew 1940(2): 50. 1940. \u0026mdash;TYPE: Zambia, \"Northern Rhodesia. Abercorn District, Near Malombe, on the road to Kambole, 2 Jun 1936, \u003cem\u003eBurtt 6115\u003c/em\u003e (Holotype: K barcode K000322622 [web!], isotypes: BR barcode BR0000006249878 [web!], EA, K barcode K000322623 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon kundelungensis\u003c/b\u003e (S.Moore) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia kundelungensis\u003c/em\u003e S.Moore in Journal of Botany, British and Foreign 57: 114. 1919. \u0026mdash;TYPE: Democratic Republic of Congo, \u0026ldquo;West Kundelungu\u0026rdquo;, 17 May 1908, \u003cem\u003eKassner 2793\u003c/em\u003e (Holotype: BM barcode BM000910840 [web!], isotypes: HBG barcode HBG512730 [web!], K barcode K000322631 [web!], P barcodes P00541090 [web!] and P00541091 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon latifolius\u003c/b\u003e (Oliv.) Brenan in Kew Bull. 4(1): 93. 1949. \u0026equiv;\u003cem\u003eArthrosolen latifolius\u003c/em\u003e Oliv. in Trans. Linn. 2(15): 348. 1887. \u0026equiv;\u003cem\u003eGnidia latifolia\u003c/em\u003e (Oliv.) Gilg in Pflanzenw. Ost-Afrikas 2(C): 283 1895. \u0026mdash;TYPE: Tanzania, Kilimanjaro, without date, \u003cem\u003eJohnston s.n.\u003c/em\u003e (Lectotype: K barcode K000322589 [web!], designated by Gastaldo (\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1969\u003c/span\u003e:365).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon lampranthus\u003c/b\u003e (Gilg) H. Pearson in Fl. Trop. Afr.6(1, 2): 233. 1910. \u0026equiv;\u003cem\u003eGnidia lamprantha\u003c/em\u003e Gilg in Bot. Jahrb. Syst. 19(2\u0026ndash;3): 264. 1894. \u0026mdash;TYPE: Tanzania, Bukoba District, Karagwe, 1892, \u003cem\u003eStuhlmann 3204\u003c/em\u003e (Lectotype: K barcode K000322594 [web!], designated by Gastaldo (\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1969\u003c/span\u003e:362).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon microcephalus\u003c/b\u003e (Meisn.) J.C.Manning \u0026amp; Magee (for protologue see under southern Africa taxa).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon mollis\u003c/b\u003e (C.H.Wright) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia mollis\u003c/em\u003e C.H.Wright in Bull. Misc. Inform. Kew 1906(1): 23. 1906. \u0026mdash;TYPE: Tanzania, \u0026ldquo;lower plateau north of Lake Nyasa [Lake Malawi]\u0026rdquo;, without date, \u003cem\u003eThomson s.n.\u003c/em\u003e (Syntypes: K barcodes K000322661 [web!] and K000378910 [web!]); Mozambique, \u0026ldquo;between Unangu and Lake Shir\u0026eacute;\u0026rdquo;, Nov. 1899, \u003cem\u003eJohnson 12\u003c/em\u003e (Syntype: K barcode K00322619 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon newtonii\u003c/b\u003e (Gilg) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia newtonii\u003c/em\u003e Gilg in Bot. Jahrb. Syst. 23(1\u0026ndash;2): 205 (1896). \u0026equiv;\u003cem\u003eArthrosolen newtonii\u003c/em\u003e (Gilg) H.Pearson in Fl. Trop. Afr. 6(1, 2): 236. 1910. \u0026mdash;TYPE: Angola, \u0026ldquo;Huilla [Huila], Chella, Monpulla, Humpata, Biballa, in montibus divulgata\u0026rdquo;, without date, \u003cem\u003eNewton 10\u003c/em\u003e (Holotype:B\u0026dagger;, isotype: K barcode K000378890 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon oliverianus\u003c/b\u003e (Vatke ex Engl. \u0026amp; Gilg) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia oliveriana\u003c/em\u003e Vatke ex Engl. \u0026amp; Gilg in Bot. Jahrb. Syst. 19(2\u0026ndash;3): 262. 1894. \u0026mdash;TYPE: Angola, \u0026ldquo;Malandsche\u0026rdquo; [Malanje], Nov-Dec 1879, \u003cem\u003eMechow 325\u003c/em\u003e (Holotype: B\u0026dagger;; Lectotype: K barcode K000322559 [web!], designated by Robyns (\u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e1975\u003c/span\u003e:56); isolectotypes: BR barcode BR0000006249991 [web!], G barcodes G00015475 [web!], G00015717 [web!], JE barcodes JE00003506 [web!], JE00003507 [web!], M barcode M0106558 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon pleurocephalus\u003c/b\u003e (Gilg) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia pleurocephala\u003c/em\u003e Gilg in Kunene-Sambesi-Exped.: 310. 1903. \u0026equiv;\u003cem\u003eArthrosolen pleurocephalus\u003c/em\u003e (Gilg) H.Pearson in Fl. Trop. Afr. 6(1, 2): 236. 1910[as \u003cem\u003epleurocephala\u003c/em\u003e]. \u0026mdash;TYPE: Angola, \u0026ldquo;Quiriri\u0026rdquo;, 18 Apr 1900, \u003cem\u003eBaum 828\u003c/em\u003e (Holotype: B\u0026dagger;, isotypes: G barcode G000190891 [web!], W barcode W19010006459 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon poggei\u003c/b\u003e (Gilg) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia poggei\u003c/em\u003e Gilg in Bot. Jahrb. Syst. 19(2\u0026ndash;3): 259. 1894. \u0026equiv;\u003cem\u003eArthrosolen poggei\u003c/em\u003e (Gilg) H.Pearson in Fl. Trop. Afr. 6(1, 2): 235. 1910. \u0026mdash;TYPE: Angola, by the River Kuango, Sept 1876, \u003cem\u003ePogge 208\u003c/em\u003e (Holotype: B\u0026dagger;).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon quarrei\u003c/b\u003e (A. Robyns) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia quarrei\u003c/em\u003e A. Robyns in Bull. Jard. Bot. Natl. Belg. 45: 223. 1975. \u0026mdash;TYPE: Democratic Republic of Congo: \u0026ldquo;Haut-Katanga, monts Marungu, Kakera, terr. Moba, en savane herbeuse\u0026rdquo;, Dec 1945, \u003cem\u003eQuarr\u0026eacute; 7599\u003c/em\u003e (Holotype: BR barcode BR0000008964885 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon polyanthus\u003c/b\u003e (Gilg) C. H. Wright (for protologue see under southern African taxa).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon polycephalus\u003c/b\u003e (E.Mey. ex Meisn.) H.Pearson (for protologue see under southern African taxa).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon rendlei\u003c/b\u003e (Hiern) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia rendlei\u003c/em\u003e Hiern in Cat. Afr. Pl. (Hiern) 1: 924. 1900. \u0026mdash;TYPE: Angola, \u0026ldquo;Huilla, between Mumpulla and Nene\u0026rdquo;, Oct 1859, \u003cem\u003eWelwitsch 6478\u003c/em\u003e (LISU-221410, syn.); Angola, \u0026ldquo;between Lopollo and Catumba\u0026rdquo;, Nov 1859, \u003cem\u003eWelwitsch 6477\u003c/em\u003e (Syntypes: BM barcode BM000910848 [web!], LISU barcode LISU221408 [web!], M barcode M010652 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon rubrocinctus\u003c/b\u003e (Gilg) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia rubrocincta\u003c/em\u003e Gilg in Bot. Jahrb. Syst. 19(2): 259. 1894. \u0026mdash;TYPE: Angola, \u0026ldquo;Oberes Kongogebiet, am Quango\u0026rdquo;, Sept 1876, \u003cem\u003ePogge 209\u003c/em\u003e (Holotype: B\u0026dagger;).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon sericocephalus\u003c/b\u003e (Meisn.) J.C.Manning \u0026amp; Boatwr. (for protologue see under southern African taxa)\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon somalensis\u003c/b\u003e (Franch) H.Pearson (for protologue see under Asian taxa)\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon stenophyllus\u003c/b\u003e (Gilg) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia stenophylla\u003c/em\u003e Gilg in Bot. Jahrb. Syst. 19(2): 259. 1894. \u0026mdash;TYPE: Tanzania, Lushoto district, Usambara Mts., Kwa Mshuza, Aug 1893, \u003cem\u003eHolst 8963\u003c/em\u003e (Holotype: B\u0026dagger;, Lectotype: K barcode K000322650 [web!]; designated by Peterson (\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e1978\u003c/span\u003e:24); isolectotypes: COI barcode COI00005795 [web!], G barcodes G00015473 [web!], G00015474 [web!], HBG barcode HBG512609 [web!], JE barcode JE00003502 [web!], M barcode M0106576 [web!], P barcodes P00541073 [web!], P00541074 [web!], S barcode S05-9780 [web!]).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon usafuae\u003c/b\u003e Gilg Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia usafuae\u003c/em\u003e in Gilg in Bot. Jahrb. Syst. 30(3\u0026ndash;4): 363. 190. \u0026mdash;TYPE: Tanzania, Mbeya/Chunya Districts, slopes of Poroto Mts. Usafua, Jun 1899, \u003cem\u003eGoetze 1042\u003c/em\u003e (Holotype: B\u0026dagger;; isotypes: BR barcodes BR0000006249861 [web!], BR0000006249984 [web!], E, G barcode G00022850 [web!], K barcodes K000322659 [web!], K000322660 [web!], P).\u003c/p\u003e \u003cp\u003e \u003cb\u003eLasiosiphon welwitschii\u003c/b\u003e (Hiern) Olaniyan, \u003cb\u003ecomb. nov.\u003c/b\u003e \u0026equiv;\u003cem\u003eGnidia welwitschii\u003c/em\u003e Hiern, Cat. Afr. Pl. 1(4): 923. 1900. \u0026mdash;TYPE: Angola, Huila, Apr 1860, \u003cem\u003eWelwitsch 6482\u003c/em\u003e (Syntypes: BM barcode BM000910855 [web!], LD barcode LD1501359 [web!], LISU barcodes LISU221405 [web!], LISU221406 [web!] and LISU221407 [web!], M barcodes M0106566 [web!], M0106567 [web!]).\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis article is part of the PhD Thesis of O. D. Olaniyan. We acknowledge the funding provided by the University of Johannesburg and the African Centre for DNA Barcoding. We thank the staff of SANBI (Pretoria National Herbarium and Compton Herbarium) for providing herbarium specimens and facilities for the herbarium work.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003cstrong\u003eAuthor contributions:\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eODO: Laboratory performance of the research, data analyses, and writing of the first draft of the manuscript. JSB: Conceptualization of the research, interpretation of the result, taxonomic and nomenclatural treatment, reviewing and editing of the manuscript. ARM: Conceptualization of the research, interpretation of the result, taxonomic and nomenclatural treatment, herbarium collections, reviewing and editing of the manuscript. JCM: Conceptualization of the research herbarium collections, taxonomic and nomenclatural treatment, reviewing and editing of the manuscript. MVDB: Funding of the project, supervision of the laboratory work, data analyses, reviewing and editing of the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflict of Interest:\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAll the authors have no conflict of interest regarding this research.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAymonin GG (1965) Sur un \u003cem\u003eGnidia\u003c/em\u003e (Thym\u0026eacute;l\u0026eacute;ac\u0026eacute;es) \u0026agrave; inflorescence complexe du Cameroun. Bull. Soc. Bot. France 112(5\u0026ndash;6): 321\u0026ndash;325. https://doi.org/10.1080/00378941.1965.10838246\u003c/li\u003e\n\u003cli\u003eAymonin GG (1966) Thym\u0026eacute;l\u0026eacute;ac\u0026eacute;es. In: Aubr\u0026eacute;ville A (ed) Flore du Gabon, vol 2. 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Sunderland, MA: Sinauer Associates\u003c/li\u003e\n\u003cli\u003eTownsend CC (1981) Thymelaeaceae. In: Dassanayake MD, Fosberg FR (eds) A revised Handbook to the Flora of Ceylon Vol 2. Amerind Publishing Co., Delhi, pp 501-511\u003c/li\u003e\n\u003cli\u003eVan der Bank M, Fay MF, Chase MW (2002) Molecular phylogenetics of Thymelaeaceae with particular reference to African and Australian genera. Taxon 51(2): 329\u0026ndash;339\u003c/li\u003e\n\u003cli\u003eVan Niekerk A (2005) Phylogenetic relationships and speciation in the genus \u003cem\u003ePasserina\u003c/em\u003e L.\u003c/li\u003e\n\u003cli\u003e(Thymelaeaceae) inferred from chloroplast and nuclear sequence data [Unpublished \u003c/li\u003e\n\u003cli\u003eMaster\u0026apos;s thesis]. University of Johannesburg, Johannesburg\u003c/li\u003e\n\u003cli\u003eWiens JJ (1998) Combining Data Sets with Different Phylogenetic Histories. Syst Biol 47(4):568\u0026ndash;581\u003c/li\u003e\n\u003cli\u003eWright CH (1915) Order CXVIII: Thymelaeaceae. In: Thiselton-Dyer WT (ed) Flora capensis. L. Reeve \u0026amp; Co., London, pp 1\u0026ndash;81\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"plant-systematics-and-evolution","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"plsy","sideBox":"Learn more about [Plant Systematics and Evolution](http://link.springer.com/journal/606)","snPcode":"606","submissionUrl":"https://submission.nature.com/new-submission/606/3","title":"Plant Systematics and Evolution","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Arthrosolen, Africa, Asia, Gnidia, Lasiosiphon, Nomenclature, Phylogeny, Taxonomy","lastPublishedDoi":"10.21203/rs.3.rs-3910184/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-3910184/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e \u003cem\u003eLasiosiphon\u003c/em\u003e Fresen. (Thymelaeaceae: Thymelaeoideae) is now understood to be a morphologically diverse genus distributed across southern and tropical Africa, Madagascar and India. The taxonomic history of \u003cem\u003eLasiosiphon\u003c/em\u003e has been marked by several revisions in its circumscription and status, particularly regarding its relationship with \u003cem\u003eGnidia\u003c/em\u003e L. Our phylogenetic analysis of nuclear (ITS) and plastid (\u003cem\u003ematK, rbcL, rps16\u003c/em\u003e and \u003cem\u003etrnL-F\u003c/em\u003e) regions of an expanded sampling of the tropical African species of \u003cem\u003eGnidia\u003c/em\u003e provides compelling evidence for including all the tropical African species of \u003cem\u003eGnidia\u003c/em\u003e in \u003cem\u003eLasiosiphon\u003c/em\u003e. The morphological characters of these species are consistent with this conclusion. We accordingly provide 29 new combinations in \u003cem\u003eLasiosiphon\u003c/em\u003e for species of \u003cem\u003eGnidia\u003c/em\u003e that currently lack combinations in that genus. We also provide a comprehensive nomenclature of all currently recognized \u003cem\u003eLasiosiphon\u003c/em\u003e species at the regional level worldwide, as the framework for future taxonomic revisions in the genus.\u003c/p\u003e","manuscriptTitle":"Molecular support for transferring the tropical African species of Gnidia to Lasiosiphon (Thymelaeaceae: Thymelaeoideae) and a worldwide synopsis of the species","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-02-02 11:13:16","doi":"10.21203/rs.3.rs-3910184/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2024-05-05T02:04:52+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2024-03-26T16:34:48+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"402e4208-4585-4ad0-b8d2-bff76c8e7233","date":"2024-02-16T01:36:26+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"ebdf19b8-3920-41ce-b53e-097619c8c0df","date":"2024-02-13T08:14:57+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2024-02-12T12:19:50+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-02-02T08:54:37+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2024-02-01T02:48:23+00:00","index":"","fulltext":""},{"type":"submitted","content":"Plant Systematics and Evolution","date":"2024-01-30T09:37:03+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"plant-systematics-and-evolution","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"plsy","sideBox":"Learn more about [Plant Systematics and Evolution](http://link.springer.com/journal/606)","snPcode":"606","submissionUrl":"https://submission.nature.com/new-submission/606/3","title":"Plant Systematics and Evolution","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"8283fbda-5fba-43d4-8c0d-33777b547e61","owner":[],"postedDate":"February 2nd, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2024-08-12T16:00:39+00:00","versionOfRecord":{"articleIdentity":"rs-3910184","link":"https://doi.org/10.1007/s00606-024-01912-3","journal":{"identity":"plant-systematics-and-evolution","isVorOnly":false,"title":"Plant Systematics and Evolution"},"publishedOn":"2024-08-08 15:57:12","publishedOnDateReadable":"August 8th, 2024"},"versionCreatedAt":"2024-02-02 11:13:16","video":"","vorDoi":"10.1007/s00606-024-01912-3","vorDoiUrl":"https://doi.org/10.1007/s00606-024-01912-3","workflowStages":[]},"version":"v1","identity":"rs-3910184","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-3910184","identity":"rs-3910184","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}