Anales del Jardín Botánico de Madrid 81 (2)
ISSN-L: 0211-1322, eISSN: 1988-3196
https://doi.org/10.3989/ajbm.622

Typification of four Linnaean names in the genus Euphorbia (Euphorbiaceae)

Tipificación de cuatro nombres linneanos del género Euphorbia (Euphorbiaceae)

 

Introduction

 

Euphorbia L. (Euphorbiaceae) is one of the largest genera of flowering plants with about 2,100 species and a worldwide distribution, but is especially diverse in arid and semi-arid regions of the tropics and subtropics (Webster 1994WebsterG.L.1994. Systematics of the Euphorbiaceae. Annals of the Missouri Botanical Gardens81: 1–144.; Govaerts & al. 2000GovaertsR., FrodinD.G. & Radcliffe-SmithA.2000. World checklist and bibliography of Euphorbiaceae (with Pandaceae). Royal Botanic Gardens, Kew.; Frodin 2004FrodinD.G.2004. History and concepts of big plant genera. Taxon53: 753–776.; Bruyns & al. 2006BruynsP.V., MapayaR.J. & HeddersonT.J.2006. A new subgeneric classification for Euphorbia (Euphorbiaceae) in southern Africa based on ITS and psbA-trnH sequence data. Taxon55: 397–420.; Riina & al. 2013; Riina & Berry 2024RiinaR. & BerryP.E. (Coor.) 2024. Euphorbia Planetary Biodiversity Inventory database. Available from: https://euphorbiaceae.org/ [accessed January 2024].; POWO 2024POWO2024. Euphorbia L.In: Plant of the World Online. Royal Botanic Gardens, Kew. https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:327729-2 [accessed January 2024].). Life forms within the genus vary from annual herbs to shrubs and large trees. Analyses of molecular data show that there are four well-defined clades within the genus, which are now treated as subgenera (Bruyns & al. 2006BruynsP.V., MapayaR.J. & HeddersonT.J.2006. A new subgeneric classification for Euphorbia (Euphorbiaceae) in southern Africa based on ITS and psbA-trnH sequence data. Taxon55: 397–420.; Zimmermann & al. 2010ZimmermannN.F.A., RitzC.M. & HellwigF.H.2010. Further support for the phylogenetics relationships within Euphorbia L. (Euphorbiaceae) from nrITS and trnL-trnF IGS sequence data. Plant Systematics and Evolution286: 39–58.): E. subg. Chamaesyce Raf. (Yang & al. 2012YangY., RiinaR., MorawetzJ.J., HaevermansT., AubriotX. & BerryP.E.2012. Molecular phylogenetics and classification of Euphorbia subgenus Chamaesyce (Euphorbiaceae). Taxon61: 764–789.); E. subg. Euphorbia (Dorsey & al. 2013DorseyB.L., HaevermansT., AubriotX.MorawetzJ.J., RiinaR., SteinmannV.W. & BerryP.E.2013. Phylogenetics, morphological evolution, and classification of Euphorbia subgenus Euphorbia. Taxon62: 291–315.); E. subg. Esula Pers. (Riina & al. 2013RiinaR., PeirsonJ.A., GeltmanD.V., MoleroJ., FrajmanB., PahlevaniA., BarresL., MorawetzJ.J., SamakiY., ZarreS., KryukovA., BruynsP.V. & BerryP.E.2013. A worldwide molecular phylogeny and classification of the leafy spurges, Euphorbia subgenus Esula (Euphorbiaceae). Taxon62: 316–342.); E. subg. Athymalus Neck. ex Rchb. (Peirson & al. 2013PeirsonJ.A., BruynsP.V., RiinaR., MorawetzJ.J. & BerryP.E.2013. A molecular phylogeny and classification of the largely succulent and mainly African Euphorbia subg. Athymalus (Euphorbiaceae). Taxon62: 1178–1199.).

The rich morphological variability of Euphorbia (Horn & al. 2012HornJ.W., van EeB.W., MorawetzJ.J., RiinaR., SteinmannV.W., BerryP.E., WurdackK.J.2012. Phylogenetics and the evolution of major structural characters in the giant genus Euphorbia L. (Euphorbiaceae). Molecular Phylogenetics and Evolution63: 305–326.) has attracted human interest around the world since ancient and even prehistoric times (Hargreaves 1981HargreavesB.J.1981. Parallels in plant use in Africa and North America. The Society of Malawi Journal34: 56–71.; Schultes 1987SchultesR.E.1987. Members of Euphorbiaceae in primitive and advanced societies. Botanical Journal of the Linnean Society94: 79–95.; Pauketat & al. 2002PauketatT.R., KellyL.S., FritzG.J., LopinotN.H., EliasS., HargraveE.2002. The residues of feasting and public ritual at early Cahokia. American Antiquity67: 257–279.). Members of Euphorbia are readily distinguishable by their milky latex and specialized inflorescences (cyathia) (Govaerts & al. 2000GovaertsR., FrodinD.G. & Radcliffe-SmithA.2000. World checklist and bibliography of Euphorbiaceae (with Pandaceae). Royal Botanic Gardens, Kew.; Horn & al. 2012HornJ.W., van EeB.W., MorawetzJ.J., RiinaR., SteinmannV.W., BerryP.E., WurdackK.J.2012. Phylogenetics and the evolution of major structural characters in the giant genus Euphorbia L. (Euphorbiaceae). Molecular Phylogenetics and Evolution63: 305–326.; Riina & Berry 2024RiinaR. & BerryP.E. (Coor.) 2024. Euphorbia Planetary Biodiversity Inventory database. Available from: https://euphorbiaceae.org/ [accessed January 2024].). In the genus, several species have a relevant economic importance, either as ornamental and domestic plants, for their appreciated latex, or for their use in folk medicine throughout the centuries (Rizk 1987RizkA.-F.M.1987. The chemical constituents and economic plants of the Euphorbiaceae. Botanical Journal of the Linnean Society94: 293–326.; Schultes 1987SchultesR.E.1987. Members of Euphorbiaceae in primitive and advanced societies. Botanical Journal of the Linnean Society94: 79–95.; Ernst & al. 2015ErnstM., GraceO.M., Saslis-LagoudakisC.H., NilssonN., SimonsenH.T. & RønstedN.2015. Global medicinal uses of Euphorbia L. (Euphorbiaceae). Journal of Ethnopharmacology176: 90–101.; Riina & Berry 2024RiinaR. & BerryP.E. (Coor.) 2024. Euphorbia Planetary Biodiversity Inventory database. Available from: https://euphorbiaceae.org/ [accessed January 2024].).

From the nomenclatural point of view, typification must be the starting point of any taxonomic study, especially in a difficult genus such as Euphorbia. The author who has described more species within this genus has been Linnaeus, with 70 published names. Effective typifications exist for most of these names (see e.g., Jarvis 2007JarvisC.E.2007. Order out of chaos: Linnaean plant names and their types. Linnean Society of London and the Natural History Museum, London.; Geltman 2015GeltmanD.V.2015. Typification of some specific and infraspecific names in Euphorbia (Euphorbiaceae). Novosti Sistematiki Vysshikh Rastenii46: 126–133., 2020GeltmanD.V.2020. A synopsis of Euphorbia (Euphobiaceae) for the Caucasus. Novitates Systematicae Plantarum Vascularium51: 43–78.). However, some Linnaean names, e.g., E. segetalis, and E. exigua var. acuta, are untypified and other (e.g., E. exigua and E. paralias) were not correctly typified and all of them are examined here. This nomenclatural act is necessary for the correct application of these names in this taxonomically difficult group.

Euphorbia exigua is the type of E. sect. Exiguae (Geltman) Riina & Molero (≡ E. subsect. Exiguae Geltman in Novosti Sist. Vyssh. Rast. 32: 101. 2000) (Riina & al. 2013RiinaR., PeirsonJ.A., GeltmanD.V., MoleroJ., FrajmanB., PahlevaniA., BarresL., MorawetzJ.J., SamakiY., ZarreS., KryukovA., BruynsP.V. & BerryP.E.2013. A worldwide molecular phylogeny and classification of the leafy spurges, Euphorbia subgenus Esula (Euphorbiaceae). Taxon62: 316–342.), the name E. exigua var. acuta is treated as heterotypic synonym of E. exigua s. str. Euphorbia segetalis belongs to E. sect. Paralias Dumort., Fl. Belg.: 87. 1827 (≡ Tithymalus subg. Paralias (Dumort.) Raf., Fl. Tellur. 4: 115. 1838 ≡ E. subg. Paralias (Dumort.) Prokh. in Komarov, Fl. URSS 14: 308. 1949) and the type is E. paralias. The purpose of this paper is to contribute to the stability of the nomenclature by the lectotypification of these four names.

Materials and methods

 

This study follows the methods described in detail by Turland & Jarvis (1997TurlandN. & JarvisC.E. (eds.). 1997. Typification of Linnaean specific and varietal names in the Leguminosae (Fabaceae). Taxon 46: 457–485.). The present work is based on the examination of relevant literature and on the study of the specimens conserved in LINN, S-LINN, OXF, and UPS (UPS-BURSER) (herbarium codes according to Thiers 2024ThiersB.2024 [continuously updated]. Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. Available from: https://sweetgum.nybg.org/ih/ [accessed: 30 Jan. 2024]. [continuously updated]). For the selection of types, protologues have been compared with original material, and the most complete, informative specimens or illustrations were selected. The identity of the designated lectotypes was verified with the current use of their respective names.

Results and discussion

 

The case of Euphorbia exigua

 

Linnaeus (1753LinnaeusC.1753. Species plantarum. Impensis Laurentii Salvii, Stockholmiae.: 456) described Euphorbia exigua providing a short diagnosis “EUPHORBIA umbella trifida: dichotoma, involucellis lanceolatis, foliis linearibus” taken from Wiman (1752WimanJ.1752. Specimen academicum quo Euphorbia ejusque historia naturalis et medica exhibetur. Excudit L.M. Höjer, Upsaliae.: pages 19–20 [species no. 29]), followed by the polynomial “Euphorbia inermis, foliis alternis linearibus acutis, umbella universali trifida, partialibus dichotomis diphyllis” cited from Linnaeus (1738LinnaeusC.1738(“1737”). Hortus Cliffortianus. Amstelaedami [Amsterdam].: 199; 1748LinnaeusC.1748. Hortus Upsaliensis. Sumtu & literis Laurentii Salvii, Stockholmiae.: 143), Van Royen (1740Van RoyenA.1740. Florae Leydensis prodromus. Apud Samuelem Luchtmans, Lugduni Batavorum [Leiden].: 197), Dalibard (1749DalibardT.-F.1749. Flora Parisiensis prodromus. Durand, Paris.: 156–157), and associated with the variety acuta, and followed by the synonyms “Tithymalus s.[ive] Esula exigua” cited from Bauhin (1623BauhinC.1623. Pinax theatri botanici. Sumptibus & typis Ludovici Regis, Basileae Helvet [Basel].: 291) and “Esula minima tragi” cited from Dalechamps (1653DalechampsJ.1653. Histoire generale des plantes, tome second. ChezPhilip. Borde, Lur. Arnaud & Cl.Rigaud, Lyon.: 520 [not 1656]).

The protologue also includes the variety retusa cited through the polynomial “Tithymalus s.[ive] Esula exigua, foliis obtusis” cited from Bauhin (1620BauhinC.1620. Prodromos theatri botanici. Typis Pauli Jacobi, Francofurti ad Moenum [Frankfurt am Main].: 133 [miscited as “132”]; 1623BauhinC.1623. Pinax theatri botanici. Sumptibus & typis Ludovici Regis, Basileae Helvet [Basel].: 291) and indicated with “β”, and followed by the synonym “Tithymalus exiguus saxatilis” cited from Bauhin (1620BauhinC.1620. Prodromos theatri botanici. Typis Pauli Jacobi, Francofurti ad Moenum [Frankfurt am Main].: 133 [miscited as “132”]; 1623BauhinC.1623. Pinax theatri botanici. Sumptibus & typis Ludovici Regis, Basileae Helvet [Basel].: 291) and Magnol (1676MagnolP.1676. Botanicum monspeliense sive plantarum circa Monspeilium nascentium. Francisci Carteron, Lugduni.: 259, t. 258), and indicated with the symbol “γ” and followed by the reference “Burs. XVI: 55”. The β and γ indicates that the protologue includes two varieties, the variety β named retusa, and the unnamed γ. The protologue also includes information about the origin of the two varieties: “Habitat in Lusatia, Gallia, Helvetia, Hispania inter segetes; β. Monspelii in saxosis. γ. Paduae & Massiliae”.

Among the original elements, the references to Magnol (1676MagnolP.1676. Botanicum monspeliense sive plantarum circa Monspeilium nascentium. Francisci Carteron, Lugduni.: t. 258) and Dalechamps (1653DalechampsJ.1653. Histoire generale des plantes, tome second. ChezPhilip. Borde, Lur. Arnaud & Cl.Rigaud, Lyon.: 520) quoted by Linnaeus in the protologue provide illustrations “Tithimalus exiguus saxatilis B prodr” (Fig. 1) and “Esula petite, de Tragus” (Fig. 2), respectively, that can be considered original elements and therefore eligible as lectotype of the name Euphorbia exigua.

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Fig. 1 Original element of Euphorbia exigua L., illustration “Tithimalus exiguus saxatilis B prodr” published by Magnol (1676MagnolP.1676. Botanicum monspeliense sive plantarum circa Monspeilium nascentium. Francisci Carteron, Lugduni.: t. 258). 
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Fig. 2 Original element of Euphorbia exigua L., illustration “Esula petite, de Tragus” published by Dalechamps (1653DalechampsJ.1653. Histoire generale des plantes, tome second. ChezPhilip. Borde, Lur. Arnaud & Cl.Rigaud, Lyon.: 520). 

Sennikov & Geltman (2013SennikovA.N. & GeltmanD.V.2013. (2119) Proposal to conserve the name Euphorbia retusa Forssk. against Euphorbia retusa (L.) Forssk. (Euphorbiaceae). Taxon62: 178–179.: 178) typified the named variety beta E. exigua var. retusa L. from a specimen collected by Loefling in Spain and preserved at LINN (Löfling 372, Herb. Linn. no. 630.29; image available at https://linnean-online.org/6273/#?s=0&cv=0), and proposed the conservation of E. retusa Forssk. (in Fl. Aegypt.-Arab.: 93. 1775) (type: Egypt, Cairo, 1762, Forsskål 1280, C barcode C10002246) against E. retusa (L.) Forssk. (in Fl. Aegypt.-Arab.: 93. 1775), the basionym of which is E. exigua var. retusa. This proposal was recommended (see Committee for Spermatophyta in Taxon 63: 1361. 2014; Wiersema & al. 2018WiersemaJ.H., TurlandN.J., BarrieF.R., GreuterW., HawksworthD.L., HerendeenP.S., KnappS., KusberW.-H., LiD.-Z., MarholdK., MayT.W., McNeillJ., MonroA.M., PradoJ., PriceM.J. & SmithG.F. (eds.). 2018+ [continuously updated]: International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017: Appendices I-VII. Online at <https://naturalhistory2.si.edu/botany/codes-proposals/> [accessed: 20 Jan. 2024].+), however, the name E. exigua var. retusa may still be used at infraspecific rank if desired.

For the name Euphorbia exigua var. acuta, according to Jarvis (2007JarvisC.E.2007. Order out of chaos: Linnaean plant names and their types. Linnean Society of London and the Natural History Museum, London.), as this varietal epithet is associated with the part of the protologue equivalent to E. exigua (rather than the named and unnamed varieties beta and gamma), it is synonymous with var. exigua, and homotypic with it. However, the polynomial mentioned by Linnaeus “Euphorbia inermis, foliis alternis linearibus acutis, umbella universali trifida, partialibus dichotomis diphyllis” is associated with the name acuta and with the reference of Hortus Cliffortianus (Linnaeus 1738LinnaeusC.1738(“1737”). Hortus Cliffortianus. Amstelaedami [Amsterdam].: 199) cited in the protologue from Linnaeus. This reference is also associated with a specimen preserved in the Clifford Herbarium at BM (with barcode BM000628681). The Clifford Herbarium includes plants that were newly cultivated in Europe at the time of collection, as well as specimens from collectors around the world. Linnaeus worked at Clifford’s estate near Haarlem between 1735 and 1737, and described the plants growing there and wrote the Hortus Cliffortianus (1738LinnaeusC.1738(“1737”). Hortus Cliffortianus. Amstelaedami [Amsterdam].), a precursor of his Species Plantarum (1753LinnaeusC.1753. Species plantarum. Impensis Laurentii Salvii, Stockholmiae.). The specimen BM000628681 can be treated as original material used by Linnaeus to describe his Euphorbia exigua var. acuta (see also Radcliffe-Smith in Meikle 1985MeikleR.D.1985. Flora of Cyprus, vol. 2. The Bentham-Moxon Trust, Kew.: 1443). The sheet at BM bears a complete and well-developed plant, and is annotated “Tithymalus. / minor ex procumbens / H.L. foliis pini / exiguus, erectus / Euphorbia / exigua” (Fig. 3). This specimen is designated as the lectotype of the name Euphorbia exigua var. acuta (see below).

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Fig. 3 Lectotype of Euphorbia exigua var. acuta L., Herb. Clifford 199, Tithymalus17 (BM, barcode BM000628681). Photography by courtesy of the herbarium BM, reproduced with permission. 

Euphorbia exigua was lectotypified by Jafri & El-Gadi (1982JafriS.M.F. & El-GadiA. 1982. Flora of Libya, vol. 89. Al Faateh University, Department of Botany, Tripoli.: 39) with the specimen Herb. Linnaeus no. 630.27 (LINN), and this typification was subsequently accepted by Jarvis (2007JarvisC.E.2007. Order out of chaos: Linnaean plant names and their types. Linnean Society of London and the Natural History Museum, London.) and Geltman (2020GeltmanD.V.2020. A synopsis of Euphorbia (Euphobiaceae) for the Caucasus. Novitates Systematicae Plantarum Vascularium51: 43–78.). The sheet at LINN bears a complete plant, with leaves and flowers, and is annotated “29 exigua” by Linnaeus at the base of the sheet. However, Linnaeus explicitly cited a specimen in the protologue, i.e., “Burs. XVI: 55”, which is currently preserved in Burser’s herbarium (UPS no. V-174929 at UPS-BURSER) in volume XVI: 55 (see also Juel 1923JuelH.O.1923. Studien in Burser’s Hortus siccus. Nova Acta Regiae Societatis Scientiarum Upsaliensis, ser. 4, 5: 1–144.: 99). This specimen is a syntype and, according to Art. 9.12 of the ICN, syntypes have precedence in lectotype designation over other original material (except isotypes) (see Turland & al. 2018TurlandN.J., WiersemaJ.H., BarrieF.R., GreuterW., HawksworthD.L., HerendeenP.S., KnappS., KusberW.-H., LiD.-Z., MarholdK., MayT.W., McNeillJ., MonroA.M., PradoJ., PriceM.J. &amp;amp; SmithG.F. (eds.). 2018. International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Glashütten, Koeltz Botanical Books.). As it is the only syntype, it must inevitably be chosen as the lectotype for this name.

Therefore, according to Art. 9.12 of the ICN, Jafri & El-Gadi’s type, cited by Jarvis (2007JarvisC.E.2007. Order out of chaos: Linnaean plant names and their types. Linnean Society of London and the Natural History Museum, London.), must be replaced by the specimen in the Burser Herbarium, vol. XVI: 55. The sheet Herb. Burser XVI: 55 (UPS no. V-174929) bears two plants, with leaves and flowers, with a label annotated as “V / Tithymalus exiguus saxatilis Baúh. / Prope Monspelium in monte saxoso / unde Bauhino transmisi. / 55.” (Fig. 4).

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Fig. 4 Lectotype of Euphorbia exigua L., Herb. Burser XVI: 55 (UPS no. V-174929). Photography by courtesy of the herbarium UPS, reproduced with permission. 

Juel (1923JuelH.O.1923. Studien in Burser’s Hortus siccus. Nova Acta Regiae Societatis Scientiarum Upsaliensis, ser. 4, 5: 1–144.) comments on some aspects related to the origin and identity of the specimen preserved in Burser’s herbarium, as follows: “BAUH. Prod., 133, sagt von dieser Art nur: ‘circa Monspelium locis asperis provenit.’ LINNÉ führt in Sp. pl. I, 456, BAUH:S Art als Euphorbia exigua γ. an, und zitiert dann: ‘Burs. XVI. 55.’ Seine Angaben über die Fundorte seiner Var. β. und γ. sind verwechselt: ‘Paduas et Massiliae’ gehört zu β., ‘Monspelii in saxosis’ zu γ., wie es aus einem Vergleich mit BAUH. Prodr. hervorgeht. BURS:S Exemplar besteht aus jungen Individuen einer E. exigua L. mit schmalen und spitzen Blättern”.

In conclusion, because Burser’s material at UPS-BURSER (XVI: 55) (now UPS no. V-174929) has been cited in the protologue by Linnaeus, and both (protologue and specimen) are linked, it is syntype (according to the ICN Art. 9.6) and has precedence in lectotype designation over unmentioned specimens (e.g., Herb. Linnaeus no. 630.27 at LINN) and cited illustrations (e.g., “Tithymalus exiguus saxatilis” in Magnol 1676MagnolP.1676. Botanicum monspeliense sive plantarum circa Monspeilium nascentium. Francisci Carteron, Lugduni. and “Esula minima tragi” in Dalechamps 1653DalechampsJ.1653. Histoire generale des plantes, tome second. ChezPhilip. Borde, Lur. Arnaud & Cl.Rigaud, Lyon.: 520) according to Art. 9.12 of the ICN. It is therefore the obligate lectotype and accordingly designated here as such.

Fortunately, this specimen clearly represents the traditional concept and current application of the name Euphorbia exigua s. str., showing relevant diagnostic characters of the species (e.g., stem leaves linear, ray and raylet leaves narrowly triangular, slightly enlarged at base) (see, e.g., Prokhanov 1949ProkhanovY.I.1949. Euphorbia L.InKomarovV.L. (ed.), Flora SSSR 14: 304–495. Izdatel’stvo Akademii nauk SSSR, Moscow & Leningrad.; Khan 1964KhanM.S.1964. Taxonomic revision of Euphorbia in Turkey. Notes of the Royal Botanical Garden Edinburgh25: 71–161.; Benedí & al. 1997BenedíC., MoleroJ., Simon PalliséJ. & VicensJ.1997. Euphorbia L.InCastroviejoS., AedoC., BenedíC., LaínzM., Muñoz GarmendiaF., Nieto FelinerG. &amp; PaivaJ. (eds.), Flora Iberica 8: 210–285. Real Jardín Botánico, CSIC, Madrid.; Tison & al. 2014TisonJ.-M., JauzeinP. & MichaudH.2014. Flore de la France méditerranéenne continentale. Naturalia publications, Turriers.; Pignatti 1982PignattiS.1982. Flora d’Italia. Vols. 1–3. Edagricole, Bologna.). Moreover, the newly designated lectotype does not change the application of the Linnaean name Euphorbia exigua subsp. exigua.

Euphorbia exigua L., Sp. Pl.: 456. 1753 subsp. exigua. Type (lectotype designated here): Herb. Burser XVI: 55 (UPS no. V-174929 [digital image!]). Fig. 4.

Euphorbia exigua var. acuta L., Sp. Pl.: 456. 1753. Type (lectotype designated here): Herb. Clifford 199, Tithymalus 17 (BM, barcode BM000628681 [digital image!]). Fig. 3.

Euphorbia exigua var. retusa L., Sp. Pl.: 456. 1753. Lectotype (designated by Sennikov & Geltman in Taxon 62: 178. 2013SennikovA.N. & GeltmanD.V.2013. (2119) Proposal to conserve the name Euphorbia retusa Forssk. against Euphorbia retusa (L.) Forssk. (Euphorbiaceae). Taxon62: 178–179.): Spain, s.d., Löfling 372, Herb. Linn. no. 630.29 (LINN [digital image!]. Image of the lectotype available at https://linnean-online.org/6273/#?s=0&cv=0

The case of Euphorbia paralias

 

Jafri & El-Gadi (1982JafriS.M.F. & El-GadiA. 1982. Flora of Libya, vol. 89. Al Faateh University, Department of Botany, Tripoli.: 44) treat two specimens at LINN (Herb. Linn. nos. 630.44 and 630.45) as the type of E. paralias, but none of these specimens in this herbarium are Linnaean original material for this name (Jarvis 2007JarvisC.E.2007. Order out of chaos: Linnaean plant names and their types. Linnean Society of London and the Natural History Museum, London.). Similarly, the specimen Herb. Linn. no. 199.15 (S-LINN) (image available at https://linnaeus.nrm.se/botany/fbo/e/eupho/euphpar2.html.en) indicated by Geltman (2015GeltmanD.V.2015. Typification of some specific and infraspecific names in Euphorbia (Euphorbiaceae). Novosti Sistematiki Vysshikh Rastenii46: 126–133.: 130; 2020GeltmanD.V.2020. A synopsis of Euphorbia (Euphobiaceae) for the Caucasus. Novitates Systematicae Plantarum Vascularium51: 43–78.: 65) as the “lectotype” is not original material (see Jarvis 2007JarvisC.E.2007. Order out of chaos: Linnaean plant names and their types. Linnean Society of London and the Natural History Museum, London.). This herbarium sheet bears a poorly preserved specimen that is annotated at the base of the sheet as “37. Paralias”, and on the verso as “It. M. Káhl [= Iter Mårten Káhler] / a Linné P. / Paralias”. According to Jarvis (2007JarvisC.E.2007. Order out of chaos: Linnaean plant names and their types. Linnean Society of London and the Natural History Museum, London.: 214-215, 600), this material was sent by Kähler and did not reach Linnaeus until after 1753 (see Jarvis 2007JarvisC.E.2007. Order out of chaos: Linnaean plant names and their types. Linnean Society of London and the Natural History Museum, London.: 214–215, 600). There are more than 50 herbarium sheets of mostly Mediterranean species in the Linnaean Herbarium that Linnaeus received from Mårten Kähler, many of which can be correlated with the manuscript list “Italica Koehleri” in LINN, thought to date from 1757. Among Kähler’s listed names can be found the name E. paralias (image available at https://linnean-online.org/162314/#?#/1&s=0&cv=1&z=-0.0932%2C0%2C1.1865%2C1.6044). Therefore, the “lectotype” designated by Geltman (2015GeltmanD.V.2015. Typification of some specific and infraspecific names in Euphorbia (Euphorbiaceae). Novosti Sistematiki Vysshikh Rastenii46: 126–133.: 130) (see also Geltman 2020GeltmanD.V.2020. A synopsis of Euphorbia (Euphobiaceae) for the Caucasus. Novitates Systematicae Plantarum Vascularium51: 43–78.: 65) is ineffective according to ICN Art. 9.3.

Linnaeus (1753LinnaeusC.1753. Species plantarum. Impensis Laurentii Salvii, Stockholmiae.: 458) described E. paralias providing the diagnosis “EUPHORBIA umbella subquinquefida: bifida, involucellis cordato-reniformibus, foliis sursum imbricatis”, cited from Wiman (1752WimanJ.1752. Specimen academicum quo Euphorbia ejusque historia naturalis et medica exhibetur. Excudit L.M. Höjer, Upsaliae.: page 31 [species no. 53]), followed by three synonyms: (1) “Euphorbia inermis, foliis setaceo-linearibus confertis, umbella universali multifida, partialibus ramose bifidis” cited from Linnaeus (1737LinnaeusC.1738(“1737”). Hortus Cliffortianus. Amstelaedami [Amsterdam].: 200 [no. 22]), Van Royen (1740Van RoyenA.1740. Florae Leydensis prodromus. Apud Samuelem Luchtmans, Lugduni Batavorum [Leiden].: 193); (2) “Tithymalus maritimus” from Bauhin (1623BauhinC.1623. Pinax theatri botanici. Sumptibus &amp; typis Ludovici Regis, Basileae Helvet [Basel].: 291) and Dodoëns (1616DodoënsR.1616. Stirpium historiae pemptades sex sive libri XXX, varie ab auctore, pauilo ante mortem, aucti et emendati. Ex Officina Plantiniana, apud Balthasarem et Ioannem Moretos frates, Antuerpiae.: 370, f. I. 2); and (3) “Tithymalus paralios” from Mattioli (1586MattioliP.A.1586. De plantis epitome utilissima, Petri Andreae Matthioli senesis; novis iconibus et descriptionibus pluribus nunc primum diligenter aucta, a d. Ioachimo Camerario; accessit catalogus plantarum, quae in hoc compendio continentur, exactiss. Frankfurt.: 962). The protologue includes as locality “Habitat in Europae arena maritima”. Mattioli (1586MattioliP.A.1586. De plantis epitome utilissima, Petri Andreae Matthioli senesis; novis iconibus et descriptionibus pluribus nunc primum diligenter aucta, a d. Ioachimo Camerario; accessit catalogus plantarum, quae in hoc compendio continentur, exactiss. Frankfurt.: 962) and Dodoëns (1616DodoënsR.1616. Stirpium historiae pemptades sex sive libri XXX, varie ab auctore, pauilo ante mortem, aucti et emendati. Ex Officina Plantiniana, apud Balthasarem et Ioannem Moretos frates, Antuerpiae.: 369, 370) provided illustrations that can be considered original material used by Linnaeus to describe E. paralias and are identifiable with the species.

According to Jarvis (2007JarvisC.E.2007. Order out of chaos: Linnaean plant names and their types. Linnean Society of London and the Natural History Museum, London.: 513), there are three herbarium sheets with specimens that are part of the original material: a sheet is preserved in the Burser Herbarium at UPS (linked to the Bauhin synonyms) (see Savage 1937SavageS.1937. Caroli Linnaei determinationes in Hortum Siccum Joachimi Burseri: The text of the manuscript in the Linnaean Collections. The Linnean Society, London.), another sheet is preserved at Clifford Herbarium at BM (linked to the synonyms taken from Linnaeus’s Hortus Cliffortianus published in 1737 on page 200), and finally there is a sheet preserved in the Linnaean herbarium at S-LINN.

The sheet Herb. Burser XVI(2): 39 (UPS no. V-174912) bears a specimen well-preserved, and a label annotated as “Tithymalus maritimus / Monspelii et Liborni”. The sheet Herb. Clifford: 200, Euphorbia 22 (now barcode BM000628687) bears a complete and well-preserved specimen annotated as “Tithÿmalus / cÿparissias. / paralias / Euphorbia / Cyparissias / 23”, “25”, and “p. 200. Euphorbia 22” (see below).

Finally, the sheet S-LINN 199.13 bears a complete and well-developed plant and is annotated “Euphorbia / 37. Paralias / Sp. pl. 458”, and on the verso “Herb. Alstroemerii / Euphorbia paralias Linn / Dahl a Linné P.” (image available at http://linnaeus.nrm.se/botany/fbo/e/eupho/euphpar1.html.en). Anders Dahl received specimens from both Linnaeus and his son, and also made his own collections in the Uppsala Botanic Garden. Many of these are now to be found in the Linnaean collection at S, annotated with a characteristic form on the verso of each sheet, and often with the words “Dahl a Linne P.”, added by him (Jarvis 2007JarvisC.E.2007. Order out of chaos: Linnaean plant names and their types. Linnean Society of London and the Natural History Museum, London.: 200). Accordingly, S-LINN 199.13 can be considered an original material used by Linnaeus to describe his species, and therefore eligible as lectotype.

Although these three specimens (at BM, UPS and S-LINN 199.13) are undoubtedly original material of E. paralias, the specimen Herb. Clifford: 200, Euphorbia 22 (BM000628687) can be identified as belonging to E. aleppica L. A lectotypification of this name on the plant preserved of this sheet at BM would be nomenclaturally disruptive because this plant is inconsistent with the current usage of the name Euphorbia paralias.

In conclusion, among all the originals elements mentioned, specimens at UPS-BURSER, BM and S-LINN, and illustrations of Mattioli (1586MattioliP.A.1586. De plantis epitome utilissima, Petri Andreae Matthioli senesis; novis iconibus et descriptionibus pluribus nunc primum diligenter aucta, a d. Ioachimo Camerario; accessit catalogus plantarum, quae in hoc compendio continentur, exactiss. Frankfurt.: 962) and Dodoëns (1616DodoënsR.1616. Stirpium historiae pemptades sex sive libri XXX, varie ab auctore, pauilo ante mortem, aucti et emendati. Ex Officina Plantiniana, apud Balthasarem et Ioannem Moretos frates, Antuerpiae.: 369, 370), I designate the specimen at S-LINN as the lectotype of E. paralias. This specimen is a complete and informative element, and matches with the traditional concept and current use of the name, showing some diagnostic features (e.g., glabrous, glaucous, somewhat fleshy, leaves 3–30 × 2–15 mm, lowest ones obovate-oblong, middle ones elliptic-oblong and upper ones ovate; all entire, adaxially concave, imbricate; ray-leaves like the upper cauline, raylet-leaves suborbicular-rhombic to reniform, strongly adaxially concave, rays 3-6, glands emarginated) (e.g., Radcliffe-Smith & Tutin 1968Radcliffe-SmithA. & TutinT.G.1968. Euphorbia L.InTutinT.G. & al. (eds), Flora Europaea 2: 213–226. Cambridge University Press, Cambridge.; Radcliffe-Smith 1982Radcliffe-SmithA.1982: Euphorbia LInDavisP.H. (ed.), Flora of Turkey and the East Aegean Islands 7: 571–630. Edinburgh University Press, Edinburgh.; Benedí & al. 1997BenedíC., MoleroJ., Simon PalliséJ. & VicensJ.1997. Euphorbia L.InCastroviejoS., AedoC., BenedíC., LaínzM., Muñoz GarmendiaF., Nieto FelinerG. &amp; PaivaJ. (eds.), Flora Iberica 8: 210–285. Real Jardín Botánico, CSIC, Madrid.; Tison & al. 2014TisonJ.-M., JauzeinP. & MichaudH.2014. Flore de la France méditerranéenne continentale. Naturalia publications, Turriers.).

Euphorbia paralias L., Sp. Pl.: 458. 1753. Type (lectotype designated here): Herb. Linn. no. 199.13 (S-LINN; code S 09-28595) (image of the lectotype available at http://linnaeus.nrm.se/botany/fbo/e/eupho/euphpar1.html.en)

The case of Euphorbia pinea

 

In the protologue of E. pinea, Linnaeus (1767LinnaeusC.1767. Caroli a Linné ... Systema Naturæ per Regna Tria Naturæ, secundum Classes, Ordines, Genera, Species, cum characteribus & differentiis. Tomus II. Editio Duodecima, Reformata. Impensis Laurentii Salvii, Stockholmiae.: 524) provided, with the number “64”, the phrase name “E. [Euphorbia] umbella quinquefida: dichotoma, involucel. [involucellis] cordatis, fol. [foliis] linearibus acuminates confertis, caps. laeviusculis”. This polynomial is quasi a verbatim copy of either Linnaeus polynomial of E. segetalis (1753LinnaeusC.1753. Species plantarum. Impensis Laurentii Salvii, Stockholmiae.: 458) cited from Wiman (1752WimanJ.1752. Specimen academicum quo Euphorbia ejusque historia naturalis et medica exhibetur. Excudit L.M. Höjer, Upsaliae.: page 22 [species no. 34]) “EUPHORBIA umbella quinquefida: dichotoma, involucellis cordatis acutis, foliis lineari-lanceolatis, ramis floriferis”.

The name has been treated as E. pinea or E. segetalis var. pinea (L.) Lange (in Willk. & Lange, Prodr. Fl. Hispan. 3: 499. 1877) (see Riina & al. 2013). However, there is no consensus on the taxonomic value of this taxon, and the name E. pinea is currently treated as a synonym of E. segetalis (Benedí & al. 1997BenedíC., MoleroJ., Simon PalliséJ. & VicensJ.1997. Euphorbia L.InCastroviejoS., AedoC., BenedíC., LaínzM., Muñoz GarmendiaF., Nieto FelinerG. &amp; PaivaJ. (eds.), Flora Iberica 8: 210–285. Real Jardín Botánico, CSIC, Madrid.). I was unable to locate any original material in the Linnaean or Linnaean-linked herbaria, and there appear to be no extant original elements (Jarvis 2007JarvisC.E.2007. Order out of chaos: Linnaean plant names and their types. Linnean Society of London and the Natural History Museum, London.). Given the absence of original material its application is clearly uncertain (see Benedí 1997BenedíC., MoleroJ., Simon PalliséJ. & VicensJ.1997. Euphorbia L.InCastroviejoS., AedoC., BenedíC., LaínzM., Muñoz GarmendiaF., Nieto FelinerG. &amp; PaivaJ. (eds.), Flora Iberica 8: 210–285. Real Jardín Botánico, CSIC, Madrid.). It is recommended to be rejected as a nomen ambiguum.

Euphorbia pinea L., Syst. Nat., ed. 12, 2: 333. 1767, nomen ambiguum.

The case of Euphorbia segetalis

 

Linnaeus (1753LinnaeusC.1753. Species plantarum. Impensis Laurentii Salvii, Stockholmiae.: 458) described E. segetalis providing the short diagnosis “EUPHORBIA umbella quinquefida: dichotoma, involucellis cordatis acutis, foliis lineari-lanceolatis, ramis floriferis”, cited from Wiman (1752WimanJ.1752. Specimen academicum quo Euphorbia ejusque historia naturalis et medica exhibetur. Excudit L.M. Höjer, Upsaliae.: page 22 [species no. 34]), followed by five synonyms: (1) “Euphorbia inermis, foliis alternis linearibus acutis, partialibus umbellae ovato-rhombeis, petalis bicornibus” cited from Linnaeus (1748LinnaeusC.1748. Hortus Upsaliensis. Sumtu & literis Laurentii Salvii, Stockholmiae.: 142), as “Hort. ups. 142.* [that * is important and something Linnaeus used to indicate that this publication has a detailed description for the species]; (2) “Euphorbia inermis, foliis linearibus acutis ad umbellam quinis isocelibus, ad umbellulas ter dichotomas ovato-trigonis” cited from Sauvages (1751SauvagesF.B. de la Croix. 1751. Methodus foliorum; seu Plantae florae Monspeliensis. A La Haye [The Hague].: 46); (3) “Tithymalus annuus, lunato flore, linariae folio longiore” from Morison (1699MorisonR.1699. Plantarum historiae universalis Oxoniensis, vol. 3. Theatro Sheldoniano, Oxonii.: 339, sect. 10, t. 2, fig. 3); (4) “Tithymalus maritime affinis, linariae folio” from Bauhin (1623BauhinC.1623. Pinax theatri botanici. Sumptibus &amp; typis Ludovici Regis, Basileae Helvet [Basel].: 291); and finally the synonym (5) “Tithymalus segetum longifolius” from Ray (1724RayJ.1724. Synopsis Methodica Stirpium Britannicarum, 3th ed. Impensis Gulielmi & Joannis Innys Regiae Societatis Typographorum, Londini.: 312). The protologue includes as locality “Habitat in Mauritania”. Morison (1699MorisonR.1699. Plantarum historiae universalis Oxoniensis, vol. 3. Theatro Sheldoniano, Oxonii.: sect. 10, t. 2, fig. 3) provided an illustration (Fig. 5) that can be considered original material used by Linnaeus to describe E. segetalis and is identifiable with the species.

media/e152_005.jpeg
Fig. 5 Lectotype of Euphorbia segetalis L., illustration “Tithÿmalus Linariae folio longiore” published by Morison (1699MorisonR.1699. Plantarum historiae universalis Oxoniensis, vol. 3. Theatro Sheldoniano, Oxonii.: sect. 10, t. 2, fig. 3). 

In 1752 was published “Specimen academicum quo Euphorbia ejusque historia naturalis et medica praeside ... Carolo Linnaeo ... sistit Johannes Wiman”, a Dissertation of Johannes Wiman (published by Höjer in Upsala) that was cited extensively by Linnaeus under the Euphorbia species named in Species Plantarum (1753LinnaeusC.1753. Species plantarum. Impensis Laurentii Salvii, Stockholmiae.) (see Wiman 1752WimanJ.1752. Specimen academicum quo Euphorbia ejusque historia naturalis et medica exhibetur. Excudit L.M. Höjer, Upsaliae.); the reference there uses the species number in the dissertation (e.g., in this case “Diss. euph. 34”) rather than the page number (Jarvis 2007JarvisC.E.2007. Order out of chaos: Linnaean plant names and their types. Linnean Society of London and the Natural History Museum, London.: 90).

I have not been able to locate any original material in any Linnaean or Linnaean-linked herbaria (see Jarvis 2007JarvisC.E.2007. Order out of chaos: Linnaean plant names and their types. Linnean Society of London and the Natural History Museum, London.). Therefore, the only extant original material eligible for typification of the name E. segetalis is the illustration of Morison (1699MorisonR.1699. Plantarum historiae universalis Oxoniensis, vol. 3. Theatro Sheldoniano, Oxonii.: sect. 10, t. 2, fig. 3). Linnaeus’s diagnosis matches with the Morison illustration except for the inflorescences (“umbella quinquefida”) that is not highlighted in the figure. However, this general character is not considered diagnostic of the species as currently recognized. In conclusion, the illustration “Tithÿmalus Linaria folio longiore” published by Morison is designated as the lectotype for the name Euphorbia segetalis (see Fig. 5).

Fortunately, Morison’s illustration matches with the traditional concept and current use of the name, showing some diagnostic features (e.g., leaves linear to linear-lanceolate, entire, ray-leaves elliptic-oblong, raylet-leaves deltate-rhombic, obtuse; base cuneate to subcordate) (see e.g., Prokhanov 1949ProkhanovY.I.1949. Euphorbia L.InKomarovV.L. (ed.), Flora SSSR 14: 304–495. Izdatel’stvo Akademii nauk SSSR, Moscow & Leningrad.; Khan 1964KhanM.S.1964. Taxonomic revision of Euphorbia in Turkey. Notes of the Royal Botanical Garden Edinburgh25: 71–161.; Radcliffe-Smith & Tutin 1968Radcliffe-SmithA. & TutinT.G.1968. Euphorbia L.InTutinT.G. & al. (eds), Flora Europaea 2: 213–226. Cambridge University Press, Cambridge.; Radcliffe-Smith 1982Radcliffe-SmithA.1982: Euphorbia LInDavisP.H. (ed.), Flora of Turkey and the East Aegean Islands 7: 571–630. Edinburgh University Press, Edinburgh.; Benedí & al. 1997BenedíC., MoleroJ., Simon PalliséJ. & VicensJ.1997. Euphorbia L.InCastroviejoS., AedoC., BenedíC., LaínzM., Muñoz GarmendiaF., Nieto FelinerG. &amp; PaivaJ. (eds.), Flora Iberica 8: 210–285. Real Jardín Botánico, CSIC, Madrid.; Tison & al. 2014TisonJ.-M., JauzeinP. & MichaudH.2014. Flore de la France méditerranéenne continentale. Naturalia publications, Turriers.).

After searching Robert Morison’s own material of E. segetalis in his herbarium (Morisonian Herbarium, in OXF), a relevant sheet has been found: Morison III. 339 no. 3. However, the Morison Herbarium was not used by Linnaeus, and so Morison’s specimens do not form original material (see Jarvis 2007JarvisC.E.2007. Order out of chaos: Linnaean plant names and their types. Linnean Society of London and the Natural History Museum, London.), but are of interest because they served as voucher specimens for Morison’s illustrations. This sheet contains a specimen with leaves and inflorescences. On the sheet, the label on the bottom left is in the hand of Jacob Bobart the Younger (1641–1719), who was the editor of and primary contributor to volume 3 of Morison’s work. The label is annotated as “Tithymalus annuus lunato flore / Linariaefolio longiore. 3 H. Oxfr. 339. / Tithymalus maritime affinis Linariae / folio. CBP. 291” (Fig. 6). The annotations on the bottom in red and black ink are in the hand of George Claridge Druce (1850–1932), whilst ‘M.P.H. II, 339. no 3 HEFG’ is in the hand of Henry Edward Fowler Garnsey (1826–1903) (Stephen Harris, OXF, pers. comm.). The specimen is clearly identifiable as Linnaeus’s E. segetalis and matches with the current application of the name.

media/e152_006.jpeg
Fig. 6 Specimen of Euphorbia segetalis preserved at Morisonian Herbarium, in OXF (Morison III. 339 no. 3), a voucher specimen for Morison’s illustration “Tithÿmalus Linariae folio longiore” published by Morison (1699MorisonR.1699. Plantarum historiae universalis Oxoniensis, vol. 3. Theatro Sheldoniano, Oxonii.: sect. 10, t. 2, fig. 3). 

Euphorbia segetalis L., Sp. Pl.: 458. 1753. Type (lectotype designated here): [illustration] “Tithÿmalus Linariae folio longiore” in Morison, Pl. Hist. Univ.: sect. 10, t. 2, fig. 3. 1699. Fig. 5.

Acknowledgments

 

I would like to express my great appreciation to Stephen Harris (OXF) and Mats Hjertson (UPS) for their help in the study of herbarium sheets.

Authorship contribution statement

 

P. Pablo FERRER-GALLEGO: Conceptualization, Data curation, Investigation, Writing–original draft, Writing–review & editing.

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