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The Nomenclature and Application of the Names Euphorbia Candelabrum Welw

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The Nomenclature and Application of the Names Euphorbia Candelabrum Welw Bruyns & Berry • Euphorbia candelabrum Welw. and E. ingens TAXON 68 (4) • August 2019: 828–838 NOMENCLATURE The nomenclature and application of the names Euphorbia candelabrum Welw. and Euphorbia ingens in tropical Africa Peter V. Bruyns1 & Paul E. Berry2 1 Department of Biological Sciences, University of Cape Town, 7701, Rondebosch, South Africa 2 University of Michigan Herbarium, Department of Ecology and Evolutionary Biology, 3600 Varsity Drive, Ann Arbor, Michigan, 48108, U.S.A Address for correspondence: Peter V. Bruyns, [email protected] DOI https://doi.org/10.1002/tax.12091 Abstract During the last 40 years, one of the most widespread and conspicuous succulent trees in East and north-east Africa has been referred to as Euphorbia candelabrum Kotschy or as E. candelabrum Trémaux ex Kotschy. This name is a later homonym of E. candelabrum Welw., and consequently it is illegitimate. The species to which the name E. candelabrum Kotschy has been widely applied is shown to be conspecific with E. ingens, which occurs from southern Ethiopia to subtropical South Africa. Keywords Euphorbiaceae; illegitimate name; synonymy; Welwitsch; widespread species ■ INTRODUCTION E. ampliphylla Pax (Bruyns & al., 2011) in E. sect. Euphorbia (Bruyns & al., 2006). These three species form large trees 8 to With the publication of accounts of Euphorbia in Flora of 30 m tall (as in Figs. 1D and 2A,D), with a broad, candelabrum- Tropical East Africa (Carter, 1988b), Flora of Somalia (Holmes, shaped crown arising from a thick, ± cylindrical trunk covered 1993), Flora of Ethiopia and Eritrea (Gilbert, 1995) and Flora by corky, brown bark. As in E. ammak Schweinf. (which is not Zambesiaca (Carter & Leach, 2001), much of the major diver- closely related, Bruyns & al., 2011), their branches are perennial sity of Euphorbia in Africa has been investigated taxonomi- and are not gradually shed as the tree grows taller (Dyer, 1957), cally, though these works are floristic and do not constitute as happens in all other arborescent African species of Euphor- thorough revisions of the species in the respective areas. A bia. Young plants of these three species are particularly distinc- complete revision of the African species of Euphorbia has tive before they begin to branch (which usually happens at 1 to also not been attempted so far. Areas where Euphorbia is 3 m tall), as they bear oblanceolate leaves at least 5 cm long diverse and where accounts are still lacking include Angola towards their tips (with strong pairs of thorns behind each leaf) and southern Africa (here taken to consist of Namibia, Leso- that are mostly (except in E. ampliphylla) much larger than the tho, South Africa and Swaziland). Although White & tiny scale-like leaf-rudiments on the branches. These species al. (1941) gave a detailed account of the succulent species in share also the unusual feature of a “lobed calyx” beneath the southern Africa, this is now considerably out of date. ovary (Brown, 1912; Leach, 1992), which is also recorded for In southern Africa ± 160 species of Euphorbia occur natu- other species (not all closely related) including E. calycina rally and, of these, around 74% are endemic. Of those that are N.E.Br. (Brown, 1912) and is shown here in Fig. 3D. not endemic, some occur near the borders of southern Africa North of Zimbabwe, very similar-looking plants to and extend beyond them, often only for a short distance. Exam- E. ingens have tended to be known as E. candelabrum Kotschy ples are E. rowlandii R.A.Dyer on the NE border of South Africa (or E. candelabrum Trémaux ex Kotschy) since Carter (1987). It (extending into neighbouring parts of Moçambique and Zimba- is difficult to see what separates these more northerly plants from bwe) and E. eduardoi L.C.Leach near the NW border of E. ingens. We investigate this further and show that these are Namibia (extending to just north of Lobito in Angola). Others, actually the same species. We also show that the name such as E. ingens E.Mey. ex Boiss. and E. grandicornis A.Blanc, E. candelabrum Kotschy is illegitimate, being a later hom- appear to have substantially larger distributions outside southern onym of E. candelabrum Welw., a different species from Angola Africa than in it. In preparation for an account of the southern (which is also arborescent but closely related to E. con finalis R.A. African species of Euphorbia, the question arises as to how large Dyer and not to E. ingens: P.V. Bruyns, unpub. data). We further the distribution is for these more widespread species. In the case show that it is impossible to apply the name E. candelabrum of E. ingens, which is considered here, there is also the problem Kotschy with certainty to any of the species found today in as to what it ought to be called over the wide swathe of Africa north-east Africa and that it may be a different species from where it occurs, as Dyer (1957) already mentioned. E. ingens. Consequently we recommend that the populations from Euphorbia ingens belongs to a group of three closely East and north-east Africa that have been named E. candelabrum related species which includes E. abyssinica J.F.Gmel. and Kotschy by several authors, should be included in E. ingens. Article history: Received: 4 Dec 2018 | returned for (first) revision: 20 Feb 2019 | (last) revision received: 18 Mar 2019 | accepted: 9 Apr 2019 Associate Editor: Erin Tripp | © 2019 International Association for Plant Taxonomy 828 Version of Record TAXON 68 (4) • August 2019: 828–838 Bruyns & Berry • Euphorbia candelabrum Welw. and E. ingens ■ WAS EUPHORBIA CANDELABRUM WELW. after being read at a meeting of the Society in June 1854. VALIDLY PUBLISHED? The relevant extracts (Welwitsch, 1855) are: “I have already become acquainted with and plundered Euphorbia candelabrum Welw. (Fig. 1A–C) was pub- upwards of forty miles of coast, from the Guizembo River lished in an English translation of a letter in German written (three miles N. of Ambriz) to near the mouth of the mighty by Friedrich Welwitsch to Richard Kippist and published in Cuanza (about 9 30′ S. Lat.), and possess the materials for a the Proceedings of the Linnean Society of London in 1855, Flora of Loanda, of five to six * miles [he referred here to Fig. 1. Euphorbia candelabrum Welw. and E. abyssinica J.F.Gmel. A, E. candelabrum, two trees south of Sumbe, Angola; B, E. candelabrum, flowering and fruiting branch, east of Sumbe, Angola; C, E. candelabrum, young tree ±1.5 m tall, east of Egito Praia, Lobito, Angola; D, E. abyssinica, tree near Adishihu, Ethiopia; E, E. abyssinica, fruiting branches on tree near Adwa, Ethiopia; F, E. abyssinica, young plant ± 0.5 m tall, with 8-angled stem, near Adishihu, Ethiopia. — Photos: P.V. Bruyns. Version of Record 829 Bruyns & Berry • Euphorbia candelabrum Welw. and E. ingens TAXON 68 (4) • August 2019: 828–838 German miles = ± 4.5 British miles] in circumference Although this letter did not give much detail of his […].” (p. 328) “Euphorbia candelabra”, there is no other species in the area “Of Euphorbia I have already found near Loanda a gigan- (i.e., within a radius of ± 20 miles of Luanda, Angola) to tic species, with a stem 2½ feet in diameter and upwards of which it could refer (Leach, 1974). The manner in which it 30 feet high, forming woods, as the Pinus sylvestris does with dominated the countryside was well-known, as shown clearly us! This species, which is readily discernable even from ship- in the sketch by Rose Monteiro that forms Plate 1 in Monteiro board, is not noticed in the ‘Flora Nigritiana.’” (p. 328) (1875) and is reproduced here as Fig. 4. Since ICN Art. 38.1 “In the woods of Euphorbia (Candelabra n.sp.) is found a does not specify how detailed a description or diagnosis ought wonderfully beautiful terrestrial Orchidea […].” (p. 329) to be, the fact that one was provided, that it characterizes the Fig. 2. Euphorbia ingens E.Mey ex Boiss. A, Tree near Isiolo, Kenya; B, Fruiting branch, near Penge, South Africa; C, Part of flowering branch, north of Narok, Kenya; D, Tree near Zion City, South Africa (with Aloe marlothii); E, Young tree ± 1 m tall, with 5-angled stem, near Zion City, South Africa; F, Part of flowering branch, Pretoria, South Africa. — Photos: P.V. Bruyns. 830 Version of Record TAXON 68 (4) • August 2019: 828–838 Bruyns & Berry • Euphorbia candelabrum Welw. and E. ingens species in the area where it was found and that this diagnosis Welwitsch’s intentions in 1854 for his letter, the first comment was made available by publication makes E. candelabrum is without foundation. As pointed out above and by Leach Welw. in Proc. Linn. Soc. 2: 329 (1855, as ‘candelabra’) both (1974), whether the context was “friendly” or not, the phrases validly and effectively published (ICN Art. 32.1, 38.1). Hiern given characterize the species in the area that Welwitsch (1900: 946) considered, erroneously, that Welwitsch pub- referred to (namely a radius of ± 20 miles around Luanda, lished the “name only” here and so he did not treat this as val- Angola) and therefore do constitute a diagnosis. The fact that idly published. other similar trees were known elsewhere is irrelevant: in the Welwitsch published the name again in Annaes do Con- area Welwitsch referred to there is no other such species. selho Ultramarino Parte Não Official in 1856 (p. 251): Therefore, these arguments do not show that E. candelabrum “N.o 5 – Euphorbia spec. (Euphorbia candelabrum Welw Welw. 1855 was not validly published. mspt) | Arvore de 30 até 45 pés de altura em fórma de cande- To contest the validity of E.
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