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BORAGINACEAE the Genus Codon Was Formally Established 78 Bothalia 35,1 (2005) Omalycus (1814) predates Calvatia Fr. (1849) by 35 years, DOMINGUEZ DE TOLEDO, L. 1993. Gasteromycetes (Eumycota) del and its adoption to cover species of Calvatia would require centra y oeste de la Argentina. I. Analisis critico de los caracteres taxonomicos, clave de los generos y orden Podaxales. Dar- a considerable number of new combinations, something winiana 32: 195-235. which is highly undesirable. Since Calvatia is already a DURIEU DE MAISONNEUVE, M.C. 1848. Exploration scientifique nomen consen’andum, it would be logical to add Omalycus de I 'Algerie 1,10. Imprimerie royale, Paris. to the list of rejected names against it, which would not pre­ FARR, E.R., LEUSSINK, J.A. & STAFLEU, F.A. (eds). 1979. Index nominum genericorum (Plantarum), vol. II. Regnum vegetabile clude the use of Omalycus for a segregate including C. 101. Bohn, Scheltema & Holkema. Utrecht. cyathiformis. A formal proposal to that effect has been sub­ GREUTER, W„ MCNEILL. J„ BARRIE. F.R., BURDET. H.M., mitted to the journal Taxon. DEMOULIN, V., FILGUEIRAS, T.S., NICOLSON. D.H., SILVA. PC., SKOG, J.E., TREHANE, P.. TURLAND, N.J. & HAWKSWORTH. D.L. 2000. International Code of Botanical AC KNO WLEDGEMENTS Nomenclature (Saint Louis Code) Regnum Vegetabile 138. Koeltz Scientific Books, Konigstein. HAWKSWORTH. D.L.. KIRK. P.M., SUTTON. B.C. & PEGLER. The first author wishes to express his gratitude to Dr D.N. 1995. Ainsworth and Bisby’s dictionary of the Fungi, edn B. Hein (B); Dr M. Pignal and Prof. P. Morat (P); and 8. CAB International, Wallingford. Dr B. Buyck and Prof. A. Coute (PC) for their hospitali­ KIRK, P.M., CANNON, P.F., DAVID. J.C. & STALPERS, J.A. 2001. ty and friendly assistance during visits to the respective Ainsworth and Bisby’s dictionary of the Fungi, edn 9. CABI Publishing, Wallingford. institutions. Dr Vincenzo Migliozzi is thanked for pro­ KREISEL. H. 1967. Taxonomisch-Pflanzengeographische Mono­ viding us with a copy of his paper on Calvatia cyathi­ graphic der Gattung Bovista. Beihefte zur Nova Hedwigia 25: formis. We are also indebted to Dr V. Demoulin, 1-244. Universite de Liege, for his valuable comments on an MIGLIOZZI, V. & COCCIA. M. 1999. Funghi del Lazio. XI. 48-50. earlier draft of this paper. Funding received from the Descrizione di Xerocomus persicolor, Calvatia cyathiformis e Mxriostoma coliforme. Micologia Italiana 3: 46-55. Peninsula Technikon research committee is acknow­ MOYERSOEN. B. & DEMOULIN. V. 1996. Les Gasteromycetes de ledged with gratitude. Corse: taxonomie, ecologie, chorologie. Lejeunia n.s. 152: 1-128. MUSS AT, E. 1901. Synonymia generum, specierum subspecierumque in vol. I-XIV descriptorum. In P.A. Saccardo, Sylloge fungorum REFERENCES omnium hucusque cognitorum 15: 201. Octave Doin Edidit, Paris. RAFINESQUE, C.S. 1814. Precis des decouvertes et travaux somio- BATSCH, A.J.G.C. 1786. Elenchus fungorum. Continuatio prima. Joh. logiques de Mr. C.S. Rafinesque-Schmaltz entre 1800 et 1814; Jac. Gebauer. Halle. ou choix raisonne de ses principales decouvertes en zoologie el CALONGE, F.D. & DEMOULIN, V. 1975. Les Gasteromycetes en hotanique. pour servir d 'introduction a ses ouvrages futurs. d’Espagne. Bulletin trimestriel de la societe mycologique de Royale Typographic Militaire, Palerme [Palermo], France 91: 247-292. ZELLER, S.M. & SMITH, A H. 1964. The genus Calvatia in North DEMOULIN, V. 1971. Le genre Lycoperdon en Europe et en Amerique America. Lloydia 27: 148-186. du Nord. Etude taxonomique et phytogeographique. Doctoral thesis, Universite de Liege. J.C. COETZEE* and A.E. VAN WYK** DESFONTAINES. R.L. 1799. Flora Atlantica. sive Historiaplantarum quae in Atlante, Agro Tunetano et Algeriensi Crescunt. Tomus * Department of Horticulture and Food Technology. Bellville Campus, Secundus. Desgranges. Paris. Cape Peninsula University of Technology. P.O. Box 1906, 7535 Bellville, DE TONI, J.B. 1888. Nidulariaceae, Lycoperdaceae et Hymenogastra- Cape Town. ceae. In P.A. Saccardo, Sylloge fungorum omnium hucusque ** H.G.W.J. Schweickerdt Herbarium. Department of Botany, University cognitorum 7,1: 28-180. Sumptibus auctoris typis seminarii, of Pretoria, 0002 Pretoria. Patavii [Padua]. MS. received: 2004-05-20. BORAGINACEAE CODONOIDEAE, A NEW SUBFAMILY BASED ON CODON The genus Codon was formally established by Carl first ‘logically and philosophically sound basis for a clas­ Linnaeus (1767) in the second volume of the 12th edition sification of plants’ (Stafleu & Cowan 1976). In 1789 of his Systema naturae. He placed the genus in his Class Antoine-Laurent de Jussieu followed with his Genera X: Decandria, Monogynia. The generic name is derived plantarum. He published the description of ‘Borragineae' from the Greek word kodon, a bell (although the flowers as one of 1 (X) orders (i.e. families). Many of his families do not hang down), and alludes to the shape of the flow­ are still maintained in modem classifications. De Jussieu ers of C. royenii L., which are deeply cup-shaped. Codon based ‘Borragineae’ on the genus Borago L. He divided comprises two described species, C. royenii and C. 28 genera into three different groups using fruit mor­ schenckii Schinz, both endemic to Namibia and South phology as a distinguishing character: 1, berry-like fruits; Africa. A possible undescribed third species is found in 2, one- or two-locular capsules; and 3, four separate nut­ the southern part of Namibia and is currently under lets. He regarded Codon as a genus of uncertain position. investigation. Of the five genera of Hydrophyllaceae known to him. It was in France that a move towards more ‘natural’ De Jussieu (1789) assigned Hydrophyllum L., Phacelia groupings of plants was first made. It is clear from his Juss. and Ellisia L. to ‘Borragineae’ and Nama L. and writings that Linnaeus recognized natural affinities, but Hydrolea L. to ‘Convolvuli’. R. Brown separated the for­ that ease of classification and identification were his mer trio of genera as the natural order Hydrophylleae in main objectives (Gunn & Codd 1981). Michel Adanson’s 1810, and the latter two as the natural order ‘Hydroleae’ Families des plantes (1763-64) can be regarded as the in 1818. Choisy (1833) treated the Hydroleae in a mono­ Bothalia 35,1 (2005) 79 graph, recognizing the genera Hydrolea, Nama, Wigan- but they would most probably also require placement in dia Kunth and Romanzoffia Cham.; to these he added one or more additional subfamilies. Eriodictyon Benth. in 1846, and at the same time vigor­ ously defended the distinctness of the Hydroleae. De Phytogeographically the restriction of Codon to the arid Candolle (1846: 589) was the first to place Codon in the southwestern comer of Africa is of special interest. Its nearest family Hydrophyllaceae. Gray (1875) united all the gen­ relatives appear to be those members of Boraginaceae s. I. pre­ era mentioned in the family Hydrophyllaceae, which he viously placed in Hydrophyllaceae s. str., found mainly in divided into four tribes. Baillon (1890) merged Hydro­ North America, but with members of both Nama and phyllaceae under Boraginaceae, but his view was not fol­ Phacelia also occurring in South America (Deginani 1999). lowed at the time. Hydrophyllaceae was restored by Brand Among plants, African-New World distributions are rather (1913) in a monograph of the family. unusual, but for southern Africa, involve as many as seven families and many more genera (Goldblatt 1978). It is intrigu­ Until recently, most authors accepted the Hydrophylla­ ing that several other plant groups with a distribution pattern ceae as a separate family. A comparison between Hydro­ comparable to that of Codon, namely with a disjunct presence phyllaceae and Boraginaceae in southern Africa based on in southwestern and in northeastern Africa, show links with pollen and macromorphological characters, however, taxa in the New World. Among these are the boraginaceous shows a strong overlap of features. The surface structure of genus Wellstedia as well as members of Calliandra, Cae- pyrenes of Ehretia P.Browne shows similarity with the salpinia, Haematoxylon. Hoffinannseggia. Parkinsonia, Xero- seeds of Nama (compare Retief & Van Wyk 2001: 15 and cladia (all Fabaceae), Nicotiana (Solanaceae), Thartmosma Chance & Bacon 1984: 832), although this may not be (Rutaceae) and Tumera (Tumeraceae) (Van Wyk & Smith meaningful, as the outer surfaces in these two structures are 2001). These African-New World disjuncts may have been obviously not homologous. Of more significance is the established in different ways, but one possible explanation is likeness between pollen grains of Wellstedia Balf.f. and based on the proximity of Africa and South America during those of Eriodictyon, Nama and Phacelia (compare Gondwana times and for quite some time after the break-up Constance & Chuang 1982 and Retief & Van Wyk 2005); of the supercontinent. tapetal orbicules or Ubisch bodies—sporopollenin particles Members of the new subfamily Codonoideae, estab­ usually lining the inner tangential tapetal cell walls of secre­ lished here, show the strongest affinity with the subfamilies tory tapetums—of Wellstedia and Codon show similarity in Wellstedioideae and Ehretioideae in Boraginaceae s.l.—in morphology (Retief et al. 2001). The broad family concept similar pollen, inflorescence and trichome morphology. of Baillon (1890) is followed here, and we agree with the However, the Codonoideae differ from Wellstedioideae in Angiosperm Phylogenetic Group (APG) (i 998, 2003) and the 10 or 12 (not 4) corolla lobes and in the seed which is Langstrom & Chase (2002) who regard Hydrophyllaceae subglobose and glabrous (not truncate and pubescent). They and Lennoaceae as synonyms of Boraginaceae s.I. differ from Ehretioideae in habit in that they are annual or short-lived perennial herbs (not shrubs or trees) and in the Modem views on the delimitation of Boraginaceae fruit which is a capsule (not a drupe). differ, for example, 1, segregating a separate family, Heliotropiaceae (Diane et al. 2002) from Boraginaceae Codonoideae Retief & A.E. van Wyk, subfam. nov. s./.; or 2, recognizing several segregate families: Bora­ Type: Codon royenii L.
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