Names and types relating to the South American genus Lamanonia (Cunoniaceae) and its synonyms, the identity of L. speciosa, and an account of the little-known L. ulei

Summary

Publication details for names that refer to the South American genus Lamanonia and its synonyms, Belangera and Polystemon, are reviewed, including names published by Vellozo, Cambessèdes, David Don, Pampanini and Engler, amongst others. The types of these names, collected by Saint-Hilaire, Glaziou, Sellow and others, are also reviewed and lectotypes designated where appropriate. A specimen of the handwriting of David Don is provided. The types of L. grandistipularis and L. speciosa are clearly conspecific and so L. speciosa is removed from the synonymy of L. ternata and now takes priority over the name L. grandistipularis for a shrub or small tree from campo rupestre in Minas Gerais and Bahia in Brazil; this species has sharply toothed, coriaceous leaves and large stipules that are usually persistent on fertile stems. A description, illustration and distribution map are provided for L. ulei, which has largely been overlooked as an accepted species, but which is distinguished from all other species in the genus by a distinctive layer of dense greyish or fawn indumentum formed of small hairs on the abaxial surface of the leaflets.

Introduction

Lamanonia Vell. (Cunoniaceae) is a small, well defined South American genus from central, eastern and southern Brazil, Paraguay and northern Argentina (Zickel & Leitão 1993; Zuloaga et al. 2008). The species are small to medium-sized trees or occasionally shrubs characterised by opposite and decussate, palmately compound leaves, each usually composed of three or five leaflets that always have toothed margins. Other distinguishing characters of the genus are two pairs of free lateral stipules per node, which can be either persistent or not in mature foliage, axillary racemose inflorescences of white to cream flowers that have a single perianth whorl, usually of six lobes, numerous (c. 25 – 60) stamens in more than one series, and a superior gynoecium composed of two carpels that are fused at the level of the ovary, each with a free style (strictly a stylodium). The fruit is a 2-valved capsule containing numerous small, flattened seeds. The most recent revision, by Zickel & Leitão (1993), accepted five species, and an additional one, L. ulei (Engl.) L. B. Sm., treated as a synonym by these authors, was recently re-instated as distinct (Hopkins et al. 2013). The generic names Belangera Cambess. and Polystemon D. Don are synonyms of Lamanonia.

Lamanonia belongs to the tribe Geissoieae (Bradford & Barnes 2001), along with three genera from the western side of the Pacific Ocean: Geissois Labill. (19 spp., New Caledonia, Vanuatu, Fiji and Vanikoro in the Solomon Islands), Pseudoweinmannia F. Muell. (2 spp., eastern Australia) and Karrabina Rozefelds & H. C. Hopkins (2 spp., eastern Australia) (Hopkins et al. 2013, 2014; Rozefelds & Pellow 2011). Within the Geissoieae, Lamanonia is distinguished by its free lateral stipules, simple axillary inflorescences and stamens in more than one series.

This paper is one of a series that reviews names and their types in the predominantly southern hemisphere family Cunoniaceae; previous publications have dealt with genera from islands in the south-west Pacific (Hopkins 2005, 2006; Hopkins & Bradford 2009). Its principal aim is to establish publication details for all names referable to Lamanonia and its synonyms, and discuss the types of these names, designating lectotypes where appropriate. No attempt is made to revise the taxonomy of the whole genus and comments on synonymy rely largely on the account by Zickel & Leitão (1993), except in a few instances.

Although Zickel & Leitão (1993) provided considerable information on names and types, more is now available, especially concerning the collections of Auguste de Saint-Hilaire (see Pignal et al. 2013). Futhermore, while trying to establish publication details of names in Cunoniaceae for the unpublished families of the World Checklist of Selected Plant Families (available online through Plants of the World Online (POWO, http://plantsoftheworldonline.org), some inconsistencies between different sources became apparent. With databases of plant names, images of many type specimens and some historical taxonomic literature now available online, a review of names and their types in this genus is timely.

Organisation of the paper and methods

The first part of the paper discusses: (A) generic names (numbers I – III) in order of their date of publication, and then in alphabetical order: (B) validly published specific (numbers 1 – 14) and (C) infraspecific names (15 – 17), and finally, (D) invalidly published names (18 – 20). Specific or infraspecific names based on the same type are dealt with together under the combination in Lamononia, if one exists; only a few names in Belangera and Polystemon have not been published as combinations in Lamanonia. The second part of the paper discusses the identity of L. speciosa (E) and presents an account of L. ulei (F). An index to types, including lectoparatypes, is given at the end.

Information on generic names was taken from the International Plant Names Index (IPNI, online) and Index Nominum Genericorum (online). Lists of species names in Lamanonia, Belangera and Polystemon were obtained from IPNI and infraspecific names were found in the taxonomic files of R. D. Hoogland at P. Images of types and other collections were viewed using Jstor Global Plants (online), the Saint-Hilaire Virtual Herbarium (SHVH) (online), the Reflora Virtual Herbarium (online), the Glaziou Virtual Herbarium (online), and the online databases of numerous individual herbaria (including B, BR, E, F, G, L, MICH, MO, NY, P, S, US). Some additional images were supplied directly by herbaria (B, FI, G, MBM, R). Protologues were seen either in the library at K or via the Biodiversity Heritage Library (online). Information on authors and their taxonomic works, including dates of publication, were taken from Taxonomic Literature ed. 2 (TL2, online and see Stafleu et al. 1976, 1979, 1981, 1983, 1985, 1986, 2000) unless otherwise stated. Dates of birth and death are given for many of the botanists and collectors mentioned below, but not the more recent ones.

In accordance with the International Code of Nomenclature (INC) (McNeill et al. 2012: Art. 9), many names are lectotypified, even when based on a single gathering, if the original material consists of more than one element and the author did not specify a holotype in the protologue. When discussing types, “!” after a herbarium abbreviation indicates that I have seen the specimen, and “image!” that I have seen an image of the specimen; however, an online image should not be regarded as part of the original material. The locations of herbarium specimens that I have not seen are cited either because of information in online databases or because the sheet was mentioned by Zickel & Leitão (1993), and for some names, it is likely that further duplicates exist. Specimen barcodes are given, when known, for types, but where a lectotype has been designated from amongst syntypes with different collection numbers, the label details, herbaria and barcodes are not indicated for the remaining syntypes, now lectoparatypes, except in the case of the Saint-Hilaire collections, for which it is necessary to discuss all the syntypes.

Several herbaria, including G and P, sometimes have multiple sheets bearing the same collector’s name and number, which together form part of a single gathering. At P, each herbarium sheet (or occasionally each fragment on one sheet) has been individually barcoded and so no ambiguity is likely. At G, material with a single collection number often comprises several individual specimens that were acquired at different times from different botanists or herbaria, and in the case of Lamanonia, each of these specimens comprises up to three individual sheets. When such a specimen comprises more than one sheet, only one bears a barcode (e.g. G00229802); recognition that the other sheets are part of the same specimen depends on the integrity of the specimen folder at G, although the images are linked under the barcode number in Jstor Global Plants (online). The sheets without barcodes are cited below using the numbers in pencil in the “Herbarium Genavense G” symbol on the sheet (e.g. 7901/12, 7901/13), rather than G00229802_a and G00229802_b, as the latter do not appear in the images seen.

In the account of Lamanonia ulei, both herbarium registration numbers and barcodes are given, when known. The distribution map was produced using SimpleMappr (online).

An overview of the taxonomic history of Lamanonia

Although Lamanonia is now the accepted name for the genus, both this name and Belangera were published almost simultaneously in 1829, and the following year, David Don published the generic name Polystemon (see below). Thus in the early 1830s, a single species had been described in Lamanonia (L. ternata Vell.), four in Belangera (B. cuneata Cambess., B. glabra Cambess., B. speciosa Cambess., B. tomentosa Cambess.) and two in Polystemon (P. pentaphyllus D. Don, P. triphyllus D. Don).

The generic names Lamanonia, Belangera and Polystemon have long been accepted as synonyms (e.g. Engler 1871, 1928; Kuntze 1891; Pampanini 1905; Zickel & Leitão 1993) but until relatively recently, the genus was usually known as Belangera. For instance, in Martius’s Flora brasiliensis, Engler (1871) recognised five species of Belangera (the four described by Cambessèdes plus B. denticulata Moric.); Pampanini (1905) treated seven species (the five recognised by Engler (1871) plus B. chabertii Pamp. and B. paraguaiensis Pamp.) as well as two infraspecific taxa; and Engler (1928) recognised eight (the five from 1871 plus B. chabertii, B. grandistipularis Taub. and B. ulei Engl.). Kuntze (1891) considered that the correct generic name was Lamanonia, rather than Belangera, and he made several new combinations, but it was not until well after Engler’s 1928 account that Lamanonia became widely accepted (e.g. by Biloni 1965; Cuatrecasas & Smith 1971; Hatschbach & Nakamura 1976; Leite 1983; Pirani & de Castro 2011; Smith 1958; Zickel & Leitão 1993; Zuloaga et al. 2008). The name Polystemon has rarely been used except in synonymy.

A. Generic Names

I. Lamanonia Vell. (Vellozo dated 1825, published 7 Sept. – 28 Nov. 1829: 228). Type: L. ternata Vell.

notes. The generic name Lamanonia was established by José Mariano da Conceição Vellozo (1742 – 1811) in Florae fluminensis (Vellozo 1829), which described plants from the area around Rio de Janeiro and which was published after his death. The title page of the text bears the date 1825; however, although the work was printed then, it was not distributed, and thus not validly published in the sense of the ICN (McNeill et al. 2012), until sometime between 7 September and 28 November 1829 (Carauta 1973). The precise date of publication is not known, but in the absence of evidence to show that Lamanonia was not validly published by 28 November 1829, the date on which Belangera was published (see below), we must assume that the former name has priority. In addition to the generic description, Vellozo (1829) gave a brief account of a single species, L. ternata.

etymology. Vellozo named Lamanonia “In memoriam D. Lamanon, globum Comite de Peyrouse circum navigante”. This refers to Jean Honoré Robert de Paul de Lamanon (1752 – 1787), often known as Robert de Lamanon, a French botanist, physicist and meteorologist, who was killed in Samoa on the Comte de Lapérouse’s ill-fated expedition to Oceania (Cartwright 1997).

type of vellozo’s name in lamanonia. According to TL2, little original Vellozo material has ever been traced (Stafleu & Cowan 1986, and see Knapp et al. 2015; Pellegrini et al. 2015). However, illustrations were made to accompany the text of Flora fluminensis (Vellozo dated 1827, published 29 Oct. 1831) and as these are part of the author’s original material, they can serve as lectotypes (see Pellegrini et al. 2015).

II. Belangera Cambess. (Cambessèdes 28 Nov. 1829: 4). Lectotype (selected here): B. speciosa Cambess.

notes. Jacques Cambessèdes (1799 – 1863) first published the name Belangera in what, in the library at K, is a 4-page leaflet with no cover, entitled Portulacearum, Crassulacearum, Ficoidearum, Cunoniacearumque, Brasiliae meridionalis, synopsis (Cambessèdes 1829). When referring to the Cunoniaceae in this leaflet, the title has often been abbreviated in the literature to “Cunon. Syn. Bras. Merid.” although TL2 gives “Portulac. Crassul. Bras. merid. syn.” as the designated abbreviation (Stafleu & Cowan 1976). The leaflet bears the printed date “Novembri 1829” and according to Stafleu & Cowan (1976), the work was published on 28 November 1829. It presents a description of the genus Belangera and diagnoses for four species: B. cuneata, B. glabra, B. speciosa and B. tomentosa, with brief mention of their places of occurrence.

Cambessèdes’s treatment of Cunoniaceae in 1829 was a precursor to his account of the family in volume 2 of Flora Brasiliae meridionalis, which was published soon after (Cambessèdes 1830, in Saint-Hilaire et al. 1829 – 1833). This flora was based on collections made by Auguste de Saint-Hilaire (1779 – 1853) in Brazil and Uruguay between 1816 and 1822 (Pignal et al. 2013), and in it, Cambessèdes re-published word-for-word his generic description of Belangera from 1829 and then gave detailed descriptions of the four species for which he had previously published only diagnoses. The title page of volume 2 bears the printed date 1829 but the account of Cunoniaceae, in part 16 of the entire work in the quarto edition, was published in October to November 1830 (Stafleu & Cowan 1983), with line drawings by Pierre Jean François Turpin (1775 – 1840) for B. glabra, B. speciosa and B. tomentosa appearing on 24 July 1830. In the folio edition, the treatment of Belangera is in vol. 2, pp. 145 – 149, with the illustrations in colour (dates of publication not given by Stafleu & Cowan 1983).

etymology. Cambessèdes (1829) stated that the genus was named in honour of Bélanger (“Dixi in honorem cl. Belanger, horti regii Pondichery rectoris ...”). Charles Paulus Bélanger (1805 – 1881) was a French explorer, naturalist and collector who was sent by the French Colonial Department to India in 1825 to establish a botanic garden in Pondicherry (Matthew 1982; van Steenis-Kruseman 1950).

types of cambessèdes’s names in belangera. According to TL2, Cambessèdes’s own herbarium is at MPU, with further material at LY, P and PC (Stafleu & Cowan 1976); these authors also stated that “Saint Hilaire left his Brazilian herbarium, including the types of the new taxa, published in the books listed here [of which “Fl. Bras. merid.” was one] to P” (Stafleu & Cowan 1983). Although the protologues were published before the account in Flora Brasiliae meridionalis, material at MPU and P with Saint-Hilaire labels thus includes the types of Cambessèdes’s names in Belangera. Some Saint-Hilaire sheets at MPU have small printed labels for Herb. Cambessèdes.

The protologues of Cambessèdes’s names in Belangera did not mention collection numbers and the locality details were brief; Cambessèdes (1830) gave more information on localities but again did not include collection numbers. As we cannot be sure that the illustrations in Flora Brasiliae meridionalis existed when the protologues were published in the earlier pamphlet, they cannot be regarded as part of the original material. Zickel & Leitão (1993) cited these illustrations as the types for three of Cambessèdes’s names and designated a neotype for the name that was not illustrated. However, since original herbarium material exists for all four names, the neotype of B. cuneata is not needed and these names are lectotypified below, following study of the Saint-Hilaire material at P and MPU.

The labels of the Saint-Hilaire specimens of Belangera at P and MPU lack detailed locality data but most have numbers, either on a label in the corner of the sheet, or on a small label attached directly to the plant fragment(s) by a thread, or on both. These correspond to the collection numbers in Saint-Hilaire’s catalogues at P, and to identify a collection precisely, both the catalogue number (e.g. C²) and collection number (e.g. 1204^bis) are needed, and should be cited as “C2 – 1204^bis” (see Pignal et al. 2013). Unfortunately, for a few collections of Belangera, the number attached to the plant fragment does not correspond to that written on the sheet label; in these instances, it is the number attached to the plant fragment that should be cited.

The Saint-Hilaire Virtual Herbarium (online) shows images of his specimens from Brazil that are now at MPU and P, as well as his catalogues, which give a brief description of each collection, sometimes with more information on locality. A few further Brazilian specimens are at K, LE (n.v.), and perhaps elsewhere. Images of some of Saint-Hilaire’s specimens of Lamanonia are also available on Jstor Global Plants (online) and the website of the Muséum national d’Histoire naturelle (MNHN), Paris (online). About half the sheets from P and MPU have a label attached with the name of the plant in spidery writing followed by the symbol “+”. Comparison with examples in Burdet (online) and a Gazette-Drouot catalogue (pdf relating to a sale of letters and manuscripts on 14 April 2015, kindly supplied by Sergio Romaniuc-Neto) show it is Cambessèdes’s writing and it is from amongst these sheets that the lectotypes have been selected. The “+” indicates that the species is not well known, following the convention proposed by de Candolle (1819), which was followed in Saint-Hilaire et al. (1829 – 1833: Preface) (Sergio Romaniuc-Neto pers. comm.).

III. Polystemon D. Don (April – June 1830: 95). Lectotype (selected here): P. pentaphyllus D. Don.

notes. David Don (1799 – 1841) published the name Polystemon in his synopsis of the Cunoniaceae in The Edinburgh New Philosophical Journal (Don 1830). Although the title page of the volume at K bears the printed date “July ... October 1830”, several pages in the first half of the volume, in which Don’s paper appears, have “April ... June 1830” printed at the bottom margin. With the description of his new genus, Don published diagnoses for two species, P. pentaphyllus and P. triphyllus. In a note at the end of his account, Don (1830: 96) equated Polystemon with Cambessèdes’s genus Belangera.

etymology. The generic name refers to the numerous stamens in each flower (Don 1830).

types of don’s names in polystemon. Following the diagnoses of his new species, Don (1830) mentioned collections for both as follows: “Hab. In Brasiliâ. – Sello. Ђ. (V. s. sp. in Herb. Lamb.)”. Friedrich Sello(w) (1789 – 1831) was a German botanist and naturalist who collected in Brazil between 1814 and 1831, sometimes in the company of Auguste de Saint-Hilaire (Herter 1945; Moraes 2008; Stafleu & Cowan 1985). According to Stearn (1973), “Ђ” is the medieval planetary symbol for Saturn and indicates the plants are trees or shrubs. “V. s. sp. in Herb. Lamb.” shows that Don had seen dried material (V. s. = vidi sec = seen dried) in the herbarium of Aylmer Bourke Lambert (1761 – 1842), one of the founding Fellows of the Linnean Society of London. For much of his career, David Don was Keeper of the Lambertian Herbarium and Librarian to the Linnaean Society (Anon 1842). Following Lambert’s death, his herbarium was sold to pay his debts and was thus dispersed, parts being acquired by 18 different institutions in Europe and the U.S.A. (Miller 1970).

According to Miller (1970), the Sellow specimens from Lambert’s herbarium are mostly at G with some at BM and K. Before his death, Lambert had sent some Sellow specimens to William J. Hooker in Glasgow and these presumably included a fragment of Lamanonia now at K (K000761379) on a sheet that bears Hooker's stamp. (Another sheet at K (K000761377), Sello 453, was received from B.) The specimens at G labelled “Herb. Delessert” were bought by Rich, as Delessert’s agent, at the sale of Lambert’s herbarium, and those now at BM were bought by Brown (Miller 1970). Index Herbariorum (Thiers online) and TL2 (Stafleu & Cowan 1985) list further herbaria where Sellow material can be found, but although some of these institutions may have duplicates of the specimens from Herb. Lambert, they are unlikely to have the material seen by Don.

The Sellow material of Lamanonia at K (two sheets) and BM (one sheet) has no original label data, no indication of the name Polystemon and no handwriting that might be that of David Don, and thus none of these specimens is likely to be the material studied by him.

At G, four sheets of Lamanonia collected in Brazil by Sellow are from Herb. Delessert; two have “Polystemon triphyllus D. Don” and two have “Polystemon pentaphyllus D. Don” written on pinned labels that also bear the stamp for “Hb. Delessert”. The names may be in David Don’s hand, based on a comparison with samples provided by the Linnean Society of London (Fig. 1), although this requires verification. Because the type material at G lacks collection numbers, it is difficult to determine whether or not Sellow sheets in other herbaria are duplicates of these collections.

Fig. 1

Specimen of the handwriting of David Don. Reproduced by permission of the Linnaean Society of London.

B. Validly published species names

1. Belangera lamanonia Steud. (Steudel 1840: 195), nom. illegit., nom. superfl.Basionym: Lamanonia ternata Vell. Type: as for Lamanonia ternata.

notes. From the layout of his list of species in the genus Belangera, Steudel (1840) clearly had the intention of making a new combination to replace the name Lamanonia ternata Vell., which was cited as a synonym. This makes B. lamanonia a superfluous name.

2. Belangera paraguaiensis Pamp. (Pampanini 1905: 53). Lectotype (selected here): Balansa 3183 [cited in protologue as 3188], Paraguay, Cordillèra de Peribebui, 1879, G – image! [G00229802] (fl.); isolectotypes: BM! [BM000548803] (buds, fl., fr.); F – image! [V0063194F]; G ×7 – images! [G00229803, G00229804, G00229805, G00229806, 7901/10 (no barcode), 7901/12 (no barcode), 7901/13 (no barcode)]; K! [K000486104] (fl. & fr.); L ×3 – images! [L0019930, L0019931, L0019932]; P ×8! [P00697283, P00697284, P01817785 (fl.), P01817786 (fr.), P01817787, P01817788, P05616946, P05616951]; NY [NY533108] (no image); U – image! [U0001475] (buds & fl.).

notes. In the protologue, Pampanini (1905) cited two collections: “1. – « Cordillère de Peribebui, – 1879 – [B. Balansa, – Pl. du Paraguay, (1878–1884), n. 3188] » (D, D C). 2. – « Vallée de l'Y-acan-guazu, près de Valenzuela, sur le versant humide des montagnes – 15 mars 1884 – [B. Balansa, – Pl. du Paraguay, (1878–1884), n. 4752] » (D).”. “D C” signifies the De Candolle herbarium at G and “D” the Delessert herbarium, now part of the general herbarium at G. Sheets of Balansa 4752 are now lectoparatypes.

According to Zickel & Leitão (1993), Belangera paraguaiensis is a synonym of Lamanonia cuneata. No combination in Lamanonia is given in IPNI (online) for the epithet paraguaiensis.

3. Lamanonia brasiliensis Zickel & Leitão (1993: 80). Type: Ratter et al. 3524, Brazil, Federal Distr., Fazenda Agua Limpa (University of Brasília field station), near Vargem Bonita, c. 18 km SSW of Brasília TV tower, 1 Sept. 1976. Holotype: UEC (fl. & fr.); isotypes: E – image! [E00319711]; K! [K000761380]; MO [MO-1551635] (no image seen); NY – image! [NY00922466]; additional isotype (fide Zickel & Leitão 1993): UB.

notes. Lamanonia brasiliensis was an accepted species in the revision of Zickel & Leitão (1993).

4. Lamanonia chabertii (Pamp.) L. B. Sm. (Smith 1958: 283).Basionym: Belangera chabertii Pamp. (Pampanini 1904: 328). Lectotype (selected here): Glaziou 8247, Brazil, (either “Rio de Janeiro, 1876” or “São Paulo, Campos da Bocaina, 8 Jan. 1876”), G – image! [G00357597] (fl.); isolectotypes: B – image! [B109009669]; BR – image! [BR0000013505233]; G ×2 – images! [G00357597, 6864-2 (no barcode)]; K! [K00761375]; P ×5! [P00697272, 00697273, P00697274, P02441814 + 1 sheet without barcode]; R – image! [R000008836]; S ×3 – images! [S 08-7441, S 08-7442, S 08-7443 (no barcodes)]; additional isolectotype cited by Zickel & Leitão (1993) (as an isotype): LE.

notes. In the protologue of Belangera chabertii, Pampanini (1904) cited the following specimen: “Hab. « Prope Rio de Janeiro (Glaziou n. 8247). » – Typ. in Herb. Delessert”, and these details were repeated in Pampanini (1905). Two sheets of Glaziou 8247 at G are from Herb. Delessert. Zickel & Leitão (1993) stated that one was the holotype and the other an isotype, but did not distinguish between them, either in print or by labelling the sheets. The sheet that gives locality data is now the lectotype.

All the type material cited above as Glaziou 8247 appears to be from a single gathering. The lectotype bears the locality “Prope Rio de Janeiro”, with the date 1876. However, among the sheets of Glaziou 8247 at P, three (P00697272, P00697273, P02441814) state “Brésil, S. Paulo, Campos de Bocaina, près de la fazenda, 8 janv 1876”. The sheet at B has “Rio de Janeiro” on the label for Herb. A. W. Eichler, but Engler’s det. slip repeats the information from the three Paris sheets and this locality is cited in Engler (1928). In his own account of his collections from Brazil, Glaziou (1905 – 1913: 199) gave the locality “Campos da Bocaina, São Paulo” for this collection, which was listed under Belangera denticulata. Several places named Bocaina occur in the state of São Paulo, and the present-day Parque Nacional de Boacina is on the border between the states of São Paulo and Rio de Janeiro. Some comments about the labelling of Glaziou material are made under Lamanonia grandistipularis.

Lamanonia chabertii was an accepted species in the revision of Zickel & Leitão (1993), known at the time only from the type, and said to be distinguished by its rather small leaflets and hairs on the filaments.

5. Lamanonia cuneata (Cambess.) Kuntze (1891: 227).Basionym: Belangera cuneata Cambess. (Cambessèdes 28 Nov. 1829: 4). Lectotype (selected here): A. de Saint-Hilaire C1 – 133, Brazil, Minas Gerais, P! [P00594819] (fr.); isolectotypes: P! [P00594818] (133, s.loc.); MPU – image! [MPU011798] (133, s.loc.).

notes. In the protologue of Belangera cuneata, Cambessèdes (1829) mentioned cuneate leaflets with puberulous lower surfaces and the fruits (capsules hirsute-tomentose, the hairs turning reddish) and he gave the locality as “Minas Geraës” (sic). The collection cited here as the type is the only Saint-Hilaire material seen with this name (Table 1). All three sheets are labelled “133”, and the lectotype also mentions a catalogue number and the province of “Minas Geraes” (sic). All the sheets have dehisced fruits and appear to be from the same gathering.

Table 1. Saint-Hilaire specimens referrable to Lamanonia (syn. Belangera), arranged by name and barcode number. Sheets at P and K have all been seen and for material at MPU, online images were studied (see Jstor Global Plants, the website of the Muséum national d’Histoire naturelle (MNHN) Paris and the Saint-Hilaire Virtual Herbarium (SHVH)). In column 4, small labels attached to fragments often have the catalogue letter indicated as a squiggle below the collection number. Only sheet P00594817 has two fragments with different collection numbers, although on some other sheets, only one of the fragments has a numbered label attached and so it is not clear whether all belong to the same gathering. LHS = left hand side; RHS = right hand side; “C2 – 1240” means: Catalogue C2, collection number 1240 (see text). In column 6, the locality data use the spellings given on the sheet label; some are printed, others are hand-written in ink. In column 8, types are said to refer to a name in Lamanonia, but note that all these names were originally published in Belangera.

As mentioned above, Zickel & Leitão (1993) designated a neotype (Hatschbach 16173, Brazil, Paraná, Porto Amazonas, Rio Iguaçu, 16 Nov. 1967, fr., MBM; isoneotypes MO, US) for this name because no illustration was provided in Cambessèdes (1830), but this neotype is not required because original material exists. Lamanonia cuneata was an accepted species in Zickel & Leitão’s revision.

6. Lamanonia denticulata (Moric.) Kuntze (1891: 227).Basionym: Belangera denticulata Moric. (Moricand Jan. – June 1847: 153, tab. 90). Lectotype (selected here): Blanchet 3253, Brazil, Bahia, G – image! [G00357595] (old fl.); isolectotypes: BM ×2!; BR ×3 – images! [BR0000013327309, BR0000013327316, BR0000013327323]; F ×2 – images! [V0063192F, V0063193F]; G ×4 – images! [G00357594, G00357596, 6864-11 (no barcode), 6864-14 (no barcode)]; K ×2! [K000486105, K000486106]; NY – image! [NY0356071]; P ×7! [P00697275, P00697276 & P00697279 (mounted on same sheet), P00697277 & P00697278 (mounted on same sheet), P05616937, P05519309 (Blanchet s.n.)]; additional isolectotype, cited by Zickel & Leitão (1993) (as an isotype): LE.

notes. Moïse-Étienne Moricand (1779 – 1854), known as Stefano, worked in the later part of his life with A. P. de Candolle in Geneva (Stafleu & Cowan 1981), and a label on the lectotype of Belangera denticulata at G indicates the sheet was previously in Moricand’s own herbarium. Moricand (1847) cited the type as “Hab. in sylvis, prov. Bahiensis. Blanchet Exsic. N° 3253”. Some collections provide further locality details, including “Jacobina” (on a sheet at F), “Solidado” (sheets at BR, F) and “circa Igregia Vilha” (sheets at BM, P).

Lamanonia denticulata is a synonym of L. ternata (fide Zickel & Leitão 1993).

7. Lamanonia glabra (Cambess.) Kuntze (1891: 227).Basionym: Belangera glabra Cambess. (Cambessèdes 28 Nov. 1829: 4). Lectotype (selected here): A. de Saint-Hilaire C – 66, Brazil, Minas Gerais, MPU – image! [MPU 011041] (fl. & fr.); isolectotypes: P ×3! [P00594820, P00594822, P00594649]; probable isolectotype: P! [P0059824 p.p., fragm. on right hand side)]. (Catalogue number given as C1 in SHVH.)

notes. In the protologue, Cambessèdes (1829) mentioned oblong leaflets narrowing at the base with the undersides glabrous and the fruits (capsules almost glabrous). He gave the collection locality as “Minas Geraës”, which is also on the printed label of one of the isolectotypes (P00594822).

The Saint-Hilaire material of Belangera glabra at P and MPU appears to represent two gatherings (Table 1). The lectotype and isolectotypes (C1 – 66) have flowers and sometimes young or mature fruits (the latter in the lectotype and P00594824). The lectoparatypes (D – 122) (P00594821, P00594823) have flowers. Sheet P00594824 is listed in SHVH as No. 1289, but the number is written on the sheet label in pencil and appears to have been added subsequent to the original labelling; the fragment on the left-hand side of this sheet matches D – 122. Based on the shape of the leaflets, collection D – 122, and probably sheet P00594821, was the basis of tab. 115 in Cambessèdes (1830).

Zickel & Leitão (1993) included this name in the synonymy of the widespread and variable Lamanonia ternata.

8. Lamanonia grandistipularis (Taub.) Taub. (Taubert 1892: 16).Basionym: Belangera grandistipularis Taub. (Taubert 1890: 17). Lectotype (selected here): Glaziou 17623, Brazil, Minas Gerais, Pico de Itabira de Campo, [21 Dec. 1888], B – image! [B109009678] (fl.); isolectotypes: C – image! [C10009982]; K! [K000486111]; P ×2! [P00697280, P00697281]; additional isolectotype listed by Zickel & Leitão (1993) (as an isotype): R.

notes. In the protologue of Belangera grandistipularis, Taubert (1890) cited the type material as “Habitat in Brasilia loco non indicato : Glaziou n. 17623”. However, two sheets at P, as well as Glaziou (1905 – 1913), give the locality for this collection as “Pico d'Itabira do Campo, Minas”. Two peaks with the name Pico do Itabira (at 19°35'S 43°15'W and 20°14'S 43°52'W) are near the town of Itabira (19°37'S 43°13'S) in the State of Minas Gerais but I have been unable to find a locality of Itabira de Campo.

The French botanist and traveller Auguste François Marie Glaziou (1828 – 1906) collected in Brazil between 1861 and the mid 1890s (Glaziou Virtual Herbarium online; Stafleu & Cowan 1976). In addition to the collections he made himself, it is known that he acquired material from other collectors and distributed it with his own labels and numbers, and that sometimes he also altered locality data (see Prance 1971; Wurdack 1970).

Among the specimens listed under Lamanonia grandistipularis by Zickel & Leitão (1993) is “Minas Gerais, Pico D’Itabira do Campo, 20 – XII – 1988 [sic], fl., Schwacke, (R 63041)”, although the sheet was determined by Zickel in 1989 as L. ternata, and the date of collection is likely to be 1888. The locality is the same as that of the type of L. grandistipularis, and Prance (1971) stated that “much Schwacke material was pirated by Glaziou”. While it is possible that the type of L. grandistipularis might have been collected by Schwacke and relabelled by Glaziou, Schwacke R 63041 represents a separate collection that does not closely resemble Glaziou 17623. Zickel & Leitão (1993) also cited Saint-Hilaire 1249 (LE) under the name L. grandistipularis, which was said to be from Rio de Janeiro. This number does not correspond to that on any of the Saint-Hilaire sheets at P or MPU and I have not seen the specimen.

Lamanonia grandistipularis was an accepted species in Zickel & Leitão’s treatment of 1993, but it is now a synonym of L. speciosa (see below under E).

9. Lamanonia speciosa (Cambess.) L. B. Sm. (Smith 1958: 283).Basionym: Belangera speciosa Cambess. (Cambessèdes 28 Nov. 1829: 4). Lectotype (selected here): A. de Saint-Hilaire s.n., Brazil, Minas Gerais, [Minas Novas], P! [P00594825] (fl.); isolectotypes: MPU – image! [MPU011042] (“Penha”, buds); P! [P00594826] (s.loc., buds & fl.).

notes. In the protologue of Belangera speciosa, Cambessèdes (1829) mentioned ovate-ellipic, acute leaflets with glabrous undersides and gave the locality as “Minas Novas”. In Cambessèdes (1830), this was expanded to “In pascuis prope pagum Nossa Sñra da Penha in parte provinciae Minas Geraës dictâ Minas Novas à fidelissimo famulo Laruotte lecta”. (In pasture near the district of Nossa Sñra da Penha, in part of the province of Minas Gerais called Minas Novas ...). “Penha” is written on a small label attached to one of the fragments on sheet MPU011042. Minas Novas (17°15'S 42°36'W) is just south of the Rio Jequitinhonha and according to Herter & Rambo (1953), Saint-Hilaire collected there in 1817. The three sheets cited here as type material represent all the Saint-Hilaire material that I have seen with the name B. speciosa and all appear to be from the same gathering (Table 1). The lectotype (P00594825) is most probably the basis for tab. 117 in Cambessèdes (1830).

Zickel & Leitão (1993) treated Lamanonia speciosa as a synonym of L. ternata; however, the type of L. speciosa is conspecific with that of L. grandistipularis and has priority over the latter name (see below under E).

10. Lamanonia ternata Vell. (Vellozo dated 1825, published 7 Sept. – 28 Nov. 1829: 228). Lectotype (selected here): Brazil, [Rio de Janeiro], habitat silvis Regii Praedii Sanctae Crucis. Original parchment plate of Flora Fluminensis in the Manuscript Section of the Biblioteca Nacional, Rio de Janeiro [cat. no. mss1198654_107] [image!] and subsequently published in Vellozo, Fl. Flumin. Icones 5: tab. 104 (dated 1827, published 29 Oct. 1831).

notes. The protologue did not mention a collection number but gave the habitat and locality: “Habitat silvis Regii Praedii Sanctae Crucis” (Grows in woods of the region of the farm of the Sacred Cross). According to Lima (1995), this is now in the neighbourhood of Santa Cruz in the city of Rio de Janeiro.

As no herbarium specimen of Lamanonia ternata collected by Vellozo is known to exist (see above), the illustration prepared to accompany the text is designated as the lectotype. Several recent publications, including Knapp et al. (2015), Moraes (2005), Pastore (2013) and Pellegrini et al. (2015), have lectotypified Vellozo names in other families. With the exception of Pastore (2013), they have all stated that the lectotypes are the original parchment plates held in the Biblioteca Nacional, Rio de Janeiro (https:/www.bn.br/) and in some cases they overturned previous lectotypifications based on the published versions of these plates. My lectotypification of L. ternata follows the format they have established.

Zickel & Leitão (1993) treated Lamanonia ternata as a widespread species that has numerous synonyms and varies in the number of leaflets (three or five) and the amount of indumentum on the foliage (glabrous to tomentose). Because the illustration that is now the lectotype is highly stylised, it would be useful to designate an epitype to fix the usage of this name; however, this should be done as part of a wider study of infraspecific variation within L. ternata, which is beyond the scope of this paper.

11. Lamanonia tomentosa (Cambess.) Kuntze (1891: 227).Basionym: Belangera tomentosa Cambess. (Cambessèdes 28 Nov. 1829: 4). Lectotype (selected here): A. de Saint-Hilaire B – 2073, Brazil, MPU – image! [MPU011043] (fl. & fr.); isolectotypes: K! [K000486108] (B – 2073, fl. buds); P! [P00594814] (B – 2073, fl. & fl. buds). (Catalogue number given as B1 in SHVH.)

notes. In the protologue of Belangera tomentosa, Cambessèdes (1829) mentioned oblong, acuminate leaflets with the lower surface pubsecent-tomentose, and white, tomentose fruits. The collection locality was given as “Prov. S. Pauli, Minas Geraës”.

Saint-Hilaire’s material of Belangera tomentosa comprises several collections and some of the label data are contradictory (Table 1). Ideally, besides having tomentose indumentum on the undersurface of the leaflets, the lectotype should have fruits, indicate the locality of Saint Paul and have a label in Cambessèdes’s handwriting, but none of the sheets fulfils all of these criteria.

Labels on three sheets listed in Table 1 mention the locality Saint Paul: P00594816 has mature fruits but the numbers on the sheet and fragment labels do not correspond, and neither P05616957 nor P05616858 (C2 – 1204 ^bis and 1204 respectively) was labelled Belangera tomentosa by Cambessèdes. The upper fragment on sheet P00594817 has the same collection number (1204^bis) as one of the preceding sheets, suggesting it also came from Saint Paul, whereas the lower fragment on this sheet (C – 342) has dehisced fruits and leaves that are a good match with those of P00594813, in which again, the number on the fragment (C – 342^bis) does not correspond to that on the sheet label (B1 – 2073). Of the four remaining sheets, MPU011043 (2073) (lectotype) appears to be the basis for tab. 116 in Cambessèdes (1830), and K000486108 (B – 2073) and P00594814 (2073) almost certainly belong to the same gathering although they lack fruits. P00594815 (B1 – 107) is the only sheet with immature fruits.

Zickel & Leitão (1993) placed Lamanonia tomentosa in the synonymy of L. ternata.

12. Lamanonia ulei (Engl.) L. B. Sm. (Smith 1958: 283).Basionym: Belangera ulei Engl. (Engler dated 1930, published 1928: 236). Lectotype (selected here): E. Ule Herb. Brasil. 4551 [cited in protologue as 4581], Brazil, Prov. Rio de Janeiro, Pedra do Cônico bei Nova Friburgo, 1300 m, 17 Jan. 1898, B – image! [B109009682] (fl.); isolectotype: HBG – image! [HBG506927].

notes. Although the name Belangera ulei was said by Zickel & Leitão (1993) to be a nomen nudum, the synoptic key in Engler (1928) provided diagnostic information for this species, allowing it to be distinguished from other members of the genus, and so the name was validly published. The protologue gave the collection locality as stated above but did not cite a particular sheet of Ule 4551 as the holotype, and so a lectotype is designated here.

The date of publication of Engler's account of the Cunoniaceae in Die Naturlichen Pflanzenfamilien was given by Stafleu & Cowan (1976) as 1930, which is the date that appears on the title page of the volume, and Stafleu & Mennega (2000) provided no further information. However, evidence from the volume itself (Engler 1928: 229), from the annual report of Berlin Botanical Garden and Museum (Anon 1929) and from Diels (1931: lvi) suggests that the treatment of Cunoniaceae was issued as a preprint in 1928. No copies of this were located in the libraries of B, G, K, L or P in 1999 (N. Kilian pers. comm.; H. M. Burdet pers. comm.; pers. obs.) and thus we have no firm evidence that this preprint was distributed in 1928 in a manner to constitute valid publication. However, the date 1928 is accepted here in the absence of proof establishing some other date of publication (following ICN, McNeill et al. 2012: Art. 31.1). The acceptance of 1928 in preference to 1930 has no nomenclatural implications in Lamanonia.

Lamanonia ulei was included in the synonymy of L. ternata by Zickel & Leitão (1993) but was re-instated as an accepted taxon in Hopkins et al. (2013) because of the distinctive greyish or fawn indumentum on the abaxial surface of the leaflets. Because no detailed account of this species has ever been published, a description and illustration are given below (see F).

13. Polystemon pentaphyllus D. Don (1830: 95). Lectotype (selected here): Sello(w) s.n., Brazil (buds): G – image! [G0015573 (6864/44)]; isolectotypes: BM!; G – image! [G00105572 (6864/48)] (buds). The following are probably also isolectotypes: BR ×2 – images! [BR0000013548490, BR0000013521912]; P! [P05519326].

notes. The two sheets at G labelled Polystemon pentaphyllus (G00105573, G00105572) both have a mixture of 3- and 5-foliolate, apparently glabrous leaves and inflorescences in bud, and are sufficiently similar to belong to a single gathering. Both bear a label indicating the name Belangera glabra in the same writing as P. pentaphyllus and the lectotype also gives the name “B. speciosa Camb.”. None of the labels on either sheet appears to mention growing in a vineyard, as mentioned by Don (1830: 95, “Arbor habitu Viticis”), although parts of the principal label for Herb. Delessert on the lectotype are now illegible. The sheet cited by Zickel & Leitão (1993) as an isotype (with number 6864/44) is now designated as the lectotype. The two sheets at BR have similar characters and are therefore likely to be isolectotypes; both are labelled B. speciosa.

The name Polystemon pentaphyllus was regarded as a synonym of Belangera speciosa by Don (1830), Engler (1871) and Pampanini (1905) (but see below under E), and as a synonym of Lamanonia ternata by Kuntze (1891), Leite (1983) and Zickel & Leitão (1993).

14. Polystemon triphyllus D. Don (1830: 95). Lectotype (selected here): Sello(w) s.n., Brazil: G – image! [G00105570 (6864/67)] (fr.); isolectotype: Sello(w) 1411, G – image! [G00105571 (6864/68)] (fl. & y. fr.).

notes. The two sheets at G labelled Polystemon triphyllus have 3-foliolate leaves; G00105570 has almost mature fruits and G00105571 has old flowers and young fruits. The leaves are sufficiently similar for them to be part of the same gathering. The sheet cited by Zickel & Leitão (1993) as an isotype (with number 6864/67) is now the lectotype. One of the labels for Herb. Delessert on sheet G00105571 has the name “Belangera tomentosa Cambess.” in the same hand as the name Polystemon triphyllus, and a small label with the number 1411 has been stuck onto this label.

Polystemon triphyllus was equated with Belangera tomentosa by Don (1830), Engler (1871) and Pampanini (1905), and with Lamanonia ternata by Zickel & Leitão (1993).

Among the Sellow collections of Lamanonia that I have seen are: Sello 453 (fl.) (K! [K000761377], P! [P05537945]), originally labelled Belangera tomentosa; Sellow 5087 (buds) (P! [P05519305]), B. cuneata; Sellow s.n. (fr.) (P! [P05519339]), B. glabra; Sello s.n. (fl.) (P! [P05519340]), B. glabra; Sello(w) s.n. (buds) (K! K000761379), with no original determination. None of these appears to belong to the same gathering as either of collections given above as the types of Polystemon pentaphyllus or P. triphyllus.

C. Validly published infraspecific names

15. Belangera glabra Cambess. var. intermedia Pamp. (Pampanini 1905: 54). Lectotype (selected here): Martius Herb. Flora Brasiliensis 93, Brazil, [Matto Grosso, Porto do Corumbá]: G – image! [G00229817] (fl.); isolectotypes: BR – image! [BR0000013327293] (Martius 93, also labelled No. 375); E – image! [E00319710]; K ×2! [K000486107, K000761378]; L – image! [L0019923]; NY – image! [NY00918700]; P! [P00697282]; additional isolectotype cited by Zickel & Leitão (1993) (as an isotype): LE.

notes. In the protologue, Pampanini (1905) cited two collections: “1. – « Brésil, St. Paul [Lund] » (D C). 2. – sine loco – « ex Martii Herb. Florae Brasiliensis, n. 93 » (D C).” He took the epithet intermedia from Martius (1837), whose name Belangera intermedia was invalidly published (see below). The locality details for the lectotype are taken from Martius (1837).

Zickel & Leitão (1993) included this name in the synonymy of Lamanonia ternata.

16. Belangera speciosa Cambess. forma pubescens Pamp. (Pampanini 1905: 56). Type: Glaziou 2495, Brazil, Prov. Rio de Janeiro. Holotype: FI – image! [FI012926] (fl. buds); isotypes: BR ×3 – images! [BR0000013505240, BR0000013505257, BR0000013505264]; K! [K000761376] (fl.); NY – image! [NY00533064] (fl. buds); P ×4 – images! [P05537925, P05537929, P05616959, P05616961]; additional isotypes (fide Zickel & Leitão 1993): MO, R.

notes. In the protologue, Pampanini (1905) cited a single collection as follows: “1. – Province of Rio de Janeiro [A. Glaziou, n. 2495] » – ex Mus. Kew. – sub B. tomentosa – (C.-I) ”. “C.-I” indicates the Centrale Italiano Herbarium at Florence. Because only one sheet was cited, it is the holotype, assuming that no further sheets of this collection exist at F. The locality indicated on the specimen label is “Rio de Janeiro” and the material consists of leaves with racemes of flowers in bud.

Numerous other sheets of Glaziou 2495 have been located, at BR, K, NY and P, and all have either flowers in bud, or at anthesis, or both, but it is possible that they represent more than one collection. Those that resemble the holotype most closely have been designated as isotypes, including the sheet at K, which I previously indicated erroneously (on the sheet) as the holotype.

The localities stated on the labels of sheets of Glaziou 2495 are either “Brésil”, “Rio de Janeiro” or “Corcovado, et Serra dos Orgãnos (Rio Jan.)” (P05616959 only), and the latter locality was given for this number and two others in Glaziou (1905 – 1913). One sheet (P05616961) that may be part of the same material as the holotype has the locality “Chemin des Macacos, Province de R. J.”, as do two others (P05537926, P05537927), although the latter possibly do not belong to the same gathering as the sheet at FI and so they have been omitted from the list of isotypes.

Engler determined the three sheets at BR as belonging to Belangera glabra. Zickel & Leitão (1993) put B. speciosa forma pubescens into the synonymy of Lamanonia ternata.

17. Belangera tomentosa Cambess. var. calvata Chodat & Hassl. (Chodat & Hassler 1903: 540). Lectotype (selected here): Hassler 6583a, Paraguay, Upper course of the river Y-acá, 1900, G – image! [G00229812] (fl.); isolectotypes: B – image! [B109009675]; BM! [BM000953539]; K! [K000486109]; G ×9 – images! [G00229810, G00229811, G00229813, 7901/19, 7901/20, 7901/21, 7901/26, 7901/27, 7901/29]; MICH – image! [MICH1192175]; MO – image! [MO-247934]; MPU – image! [MPU011797]; P ×3! [P03322143, P03322152, P05537934]; S – image! [S08-7445 (no barcode)]; UC [UC940752]; US – image! [US00036514].

notes. The protologue referred to two collections: “ad marginem fluminis Y-aca in campo rupestre pr. Chololo, Dec., n. 6583a florifera et 6583b. Jan. fructifera”. The sheet at K of Hassler 6583a gives a variant of this locality (“In regione cursus superioris flumis Y-acá”). Numerous duplicates of both syntypes exist and according to Stafleu & Cowan (1979), Émile Hassler’s (1864 – 1937) own herbarium from Paraguay is at G, with other sets at K, P, S and W (plus B†). Sheets of Hassler 6583b are now lectoparatypes.

Belangera tomentosa var. calvata was cited as a synonym of Lamanonia cuneata by Zickel & Leitão (1993). Stearn (1973) gave the meaning of “calvata” as hairless, glabrous or bald.

D. Invalid names

18. Belangera hirta Glaz. (Glaziou July 1906: 199), nom. subnud. Specimen cited: Glaziou 21119a, Brazil, Goyaz, Corrego do Brejo, 9 May 1895, G – image! [G00229818] (fl.); P ×2! [P05537930, P05537935]; R – image! [R000008838]).

notes. After the name “B. hirta Glaz.”, Glaziou (1906) stated “sp. n.? [...] ined.” and for a description, wrote only “Grand arbre, fl. blanchâtres”. As this is inadequate as a description or diagnosis, the name is invalid. He referred to a single collection: “in herb. Berol. [...] Corrego do Brejo, Goyaz, n° 21119 a”. At B, no material of this number has been found (Robert Vogt, pers. comm. 11 May 2016), although a sheet of Glaziou 21119 is specimen of Weinmannia. The sheet at G (G00229818) was determined as Belangera tomentosa by Pampanini in 1904 and as Lamanonia ternata by Zickel in 1989. The two sheets at P are both labelled “Belangera hirta Glaz. sp. nov.”.

Glaziou’s account of his collections (Glaziou 1905 – 1913) was published in vol. 3 of Mém. Soc. Bot. France, in several parts, labelled a, b and c, but all are part of a single work. The fascicle in which Belangera hirta appeared has the date “juillet 1906” printed on the last page (p. 200), although the front cover bears the date “1907”, with “22 Février 1908” in small print by the bottom margin. Stafleu & Cowan (1976) gave the date of publication of this fascicle as July 1906.

19. Belangera intermedia Mart. (Martius 1837: 95), nom. subnud. Specimen cited: Martius 93, Porto do Corumbá, prov. Mato Grosso, fl. Aug. Oreas. (See Belangera glabra var. intermedia for further details).

notes. The collection cited by Martius (1837) is now the type of Lamanonia glabra var. intermedia Pamp. “Oreas” refers to the plant growing in the mountains (Stearn 1973).

20. Belangera riedelina Casar. ex Engl. (Engler 1870: 591), nom. nud., pro syn. sub Belangera speciosa. Specimen cited: none.

notes. As Giovanni Casaretto’s name was simply cited by Engler (1870, 1871) as a synonym of Belangera speciosa, the name is invalid.

E. The identity of Lamanonia speciosa

As mentioned above, in the most recent revision of this genus, Zickel & Leitão (1993) placed the name Lamanonia speciosa (and its basionym Belangera speciosa) in the synonymy of L. ternata, and treated L. grandistipularis as a separate taxon. However, type specimens of L. speciosa (Saint-Hilaire s.n.) and L. grandistipularis (Glaziou 17623) compared at P are strikingly similar and clearly conspecific. The type material of both names has large stipules that are persistent at several nodes on fertile stems (in the remaining taxa the stipules are usually smaller and generally caducous on fertile stems), as well as 5-foliolate leaves that have glabrous, rather coriaceous leaflets with sharply toothed margins. The leaflets in both types are of similar size and shape and most are slightly bullate, with the intervenium raised on the upper surface between the impressed secondary veins (in other species the intervenium is ± flat). Types of both names have axillary racemes that exceed the leaves, although the inflorescences of Glaziou 17623 are longer than those in the Saint-Hilaire material. The similarity between the types is illustrated in Fig. 2, which shows the drawing of L. speciosa from Cambessèdes (1830, as B. speciosa), based on the specimen that is now the lectotype of that name, next to a sheet of Glaziou 17623. (Note also the similarity between Fig. 2A and the drawing of L. grandistipularis in Pirani & de Castro 2011: 43, Fig. 1A).

Fig. 2

Comparison of Lamanonia speciosa and L. grandistipularis. A drawing of Belangera speciosa Cambess., the basionym of L. speciosa (Cambess.) L. B. Sm., from Cambessèdes (1830: tab. 117), based on Saint-Hilaire s.n. [P00594825]; B image of Glaziou 17623, isolectotype of L. grandistipularis (Taub.) Taub. [http://specimens.kew.org/herbarium/K000486111] reproduced from The Herbarium Catalogue, with permission of the Trustees of the Royal Botanic Gardens, Kew.

The types of Lamanonia speciosa and L. grandistipularis belong to a species that is distinct from the remainder of the material that Zickel & Leitão (1993) called L. ternata (which includes the types of L. glabra and L. tomentosa, amongst others). According to these authors, although L. ternata and L. grandistipularis both have leaves with either three or five leaflets, in L. grandistipularis they are coriaceous, shiny and glabrous above and puberulous beneath, and in L. ternata they are chartaceous, opaque, glabrous to puberulous above, and glabrous, puberulous or tomentose beneath. In their key, Zickel & Leitão distinguished L. grandistipularis principally by its stipules, which were said to be large (c. 1.5 × 1 cm), persistent and semi-cordate, where as in L. ternata (and others) they are smaller (c. 0.4 × 0.3 cm), caducous or persistent, falcate and chartaceous. (However, their description of L. ternata gave the size range of the stipules as 0.8 – 1.4 × 0.2 – 0.7 cm, and they thus ± accommodated the illustration of B. speciosa from Cambessèdes 1830). L. grandistipularis is a low shrub or small tree 2 – 6 m high (the remaining taxa in the genus are generally but not always larger), and it is known principally from the Espinhaço Range in Minas Gerais and Bahia, where it grows in campo rupestre (savanna in rocky, mountainous areas) and associated cerrado (savanna) (Pirani & de Castro 2011; Zickel & Leitão 1993). L. ternata was said to be more widely distributed in southern, central and eastern Brazil, where its distribution overlaps with that of L. grandistipularis, and it occurs in a variety of wooded habitats. As noted by Zickel & Leitão (1993), juvenile material of L. ternata has more sharply toothed leaflets and larger, more persistent stipules than the adult foliage, and thus bears a closer resemblance to L. grandistipularis.

Unfortunately, the information available regarding the habitat and elevation for the type collections of Lamanonia grandistipularis and L. speciosa is inadequate to show whether either was collected in campo rupestre, and even the locality of the type of L. grandistipularis is not entirely certain because of the labelling problems associated with Glaziou. As far as we know, Glaziou 17623 came from near Itabira in central Minas Gerais but no habitat is given on the label. The habitat for the type of L. speciosa, from Minas Novas to the north-east of Itabira, also in Minas Gerais, is stated to be pasture, which might indicate cattle pasture or cerrado/campo rupestre.

The name Lamanonia (or Belangera) speciosa has been much mis-interpreted, almost since first publication. Although “Belangera glabra” is written on the lectotype of Polystemon pentaphyllus, in print, Don (1830) incorrectly equated P. pentaphyllus with B. speciosa, perhaps because the type material of both names has 5-foliolate, glabrous or almost glabrous leaves. Don made no mention of the stipules of P. pentaphyllus; Cambessèdes (1829) had not mentioned these structures in the protologue of B. speciosa, describing them only in 1830, when he said that they were oblong, falcate and 8 – 10 lines long (in contrast to the stipules of his other three species that were all between 3 and 4 lines long).

Following on from Don, some material of Lamanonia with glabrous leaves has been identified as L. (or Belangera) speciosa, irrespective of the size and persistence of the stipules and the thickness of the leaflets. Engler (1871), Kuntze (1891) and Pampanini (1905) all (wrongly) treated Lamanonia ternata as a synonym of B. speciosa, and conversely, Leite (1983) included B. speciosa in the synonymy of L. ternata. In addition, L. (or B.) speciosa has often been used on specimen labels and in flora accounts (including Biloni 1965; Cuatrecasas & Smith 1971; Hatschbach & Nakamura 1976) for material that was later placed in L. ternata by Zickel & Leitão (1993). One dissenting voice was Smith (1958), who treated L. speciosa and L. ternata as distinct species; however, he differentiated between them only by the number of leaflets, which alone, is insufficient to distinguish two taxa. Much of the material in herbaria that has at some time been labelled as L. (or B.) speciosa is, in fact, closer to the type of B. glabra.

In contrast to the situation regarding Lamanonia (or Belangera) speciosa, the name L. (or B.) grandistipularis has generally been regarded as referring to a distinct species (e.g. Engler 1928), although Leite (1983) put it into synonymy under L. ternata before Zickel & Leitão (1993) reinstated it as a separate taxon.

In conclusion, the names Lamanonia speciosa and Belangera speciosa must be removed from the synonymy of L. ternata and the epithet speciosa has priority over grandistipularis for the shrub or small tree from campo rupestre that has a combination of rather coriaceous, sharply toothed leaflets and comparatively large stipules that are generally persistent on fertile stems.

Lamanonia speciosa (Cambess.) L. B. Sm. (Smith 1958: 283).Basionym: Belangera speciosa Cambess. (Cambessèdes 1829: 4). Type: see above.Synonyms: Belangera grandistipularis Taub. (Taubert 1890: 17), synon. nov. Lamanonia grandistipularis (Taub.) Taub. (Taubert 1892: 16), synon. nov. Type: see above.

F. Lamanonia ulei

Lamanonia ulei (Engl.) L. B. Sm. (Smith 1958: 283).Basionym: Belangera ulei Engl. (Engler 1928: 236). Type: see above.

Treelets or small trees, 4 – 10 m tall. Indumentum on leaf-bearing twigs, petioles and inflorescence axes fawn, composed of two layers, the upper layer hirsute to tomentose (hairs c. 1 mm long), over a lower dense layer of minute hairs. Apical bud of shoot circular in outline and swollen, 3 – 4 mm diam., densely hirsute, hairs fawn. Stipules 4 per node, free, rhombic-elliptic, 6 × 3 mm, with the adaxial surface bearing long and short hairs, caducous, leaving scars 1 mm long. Leaves opposite and decusssate, each 3-foliolate, the median leaflet larger than the laterals; in fertile material: petioles (1.6 –) 2 – 3 cm long; median leaflets with a short petiolule 3 – 6 mm long, the blade ovate, (4.6 –) 5 – 9 × (2 –) 2.3 – 4 cm, cuneate at the base, usually acute or obtuse at the apex, tending to be conduplicate distally; lateral leaflets ± sessile and symmetrical at the base, the blades ovate, (3.3 –) 4 – 7 × (1.1 –) 1.7 – 3.6 cm; margins serrate; upper surface of leaflets glabrous except along the midrib towards the base of the blade, with midrib and secondary veins impressed; lower surface of leaflets with midrib and secondary veins prominent, clearly visible and appearing darker than the intervenium, bearing fawn hairs to c. 0.5 mm long, glabrescent; intervenium covered by a dense, almost felty indumentum of minute, dull white, fawn or yellowish grey hairs; tertiary veins less clearly visible than the secondary veins to almost obscure, covered by minute hairs; secondary veins pinnate, 12 – 16 on either side of the midrib in median leaflets, each one ending at the sinus of a tooth; tertiary veins somewhat ladder-like. Leaves in sterile material larger, median leaflets to 14 cm long. Inflorescences axillary, racemose; axis 6.5 – 12.5 cm long (including a basal peduncle of 1.5 – 4 cm), projecting beyond the subtending leaves. Flowers well spaced along the axis, white or cream at anthesis; pedicels 2 – 3 mm long (– 5 mm in fruit), buds ovoid, 4 – 5 mm long, pedicels and buds covered by dense, fawn indumentum; calyx lobes (4) 6, triangular, 4.5 × 1.5 mm, coriaceous, outer surface with faint longitudinal ridges, inner surface hairy; corolla absent; stamens numerous, in > 1 series; filaments c. 4 mm long but not all of equal length, narrowly cylindrical-subulate, hairy in bud especially towards the base; anthers 0.5 × 0.5 mm, versatile, not apiculate; gynoecium ovoid, 4 × 3 mm, densely golden-hairy except around the base where nectar is probably secreted, comprising 2 locules, each containing numerous ovules in two rows; styles 2 per ovary, c. 1.5 mm long, glabrous. Capsules numerous per infructescence, dry, woody, brown, narrowly ovoid to ± cylindrical, 10 – 13 mm long (including 2 apical beaks 1 mm long) × 4 – 5 mm diam., with a constriction at the base (former nectar-secreting region), sitting above a wider annulus bearing the calyx scars, narrowing towards the pointed apex, dehiscing almost to the base into 2 valves, the valves scarcely separating; except for the glabrous constricted region, outer surface of valves bearing fawn indumentum in two layers, the upper one pubescent (hairs to 0.5 mm long), in the lower layer hairs minute, vermiform; inner surface of each half-capsule glabrous and bony, deeply incised longitudinally down the mid-line dividing each locule into 2 sub-locules. Seeds numerous (c. 9 per half-locule, i.e. 36 per capsule), densely packed and imbricate, 3.5 – 5 × 1.8 – 2 mm, glabrous, orange-red (when dry), not all identical in size and shape, thin and either flat or sometimes with a raised lateral auricle, each with a very narrow, marginal wing. Fig. 3.

Fig. 3

Lamanonia ulei. A leafy shoot with inflorescences in bud, * indicating that part of one of the most distal pair of leaves has been removed for clarity; the apical bud of the shoot is at the base of this petiole; B terminal node of a shoot showing an apical bud, a pair of axillary buds with stipular scars at their bases and one stipule remaining, and the bases of two opposite petioles; C part of the lower surface of a leaflet; D detail of C, showing the midrib and secondary vein with longish hairs, and the intervenium and tertiary veins covered by a dense, felty coat of minute hairs; E flower, lateral view, after the removal of part of the perianth and some stamens; F fruit, lateral view; G adaxial surface of one half of a fruit; H detail of the fruit wall, showing long hairs overlying a minute, vermiform indumentum on the surface; J both surfaces of a seed. (A – E from Plowman & de Lima 12870; F – J from Moraes 151). drawn by andrew p. brown.

distribution. South-eastern Brazil (Map 1).

Map 1

The distribution of Lamanonia ulei in south-eastern Brazil.

specimens examined. brazil. Minas Gerais: Caparaó, Parque Nacional do Caparaó, próximo a Cachoeira Bonita, 1700 – 1800 m, 14 June 1991 (fl.), G. & M. Hatschbach 55459 & J. M. Silva (MBM – image! [MBM0151324]); Fazenda Silva, Alto da pedra Caratinga, 17 Sept. 1929 (fr.), I. Kuhlmann 7 (RB – image! [RB111079, barcode: RB00073232]). Rio de Janeiro: Petrópolis, A.P.A. Petrópolis, Vale das Videiras, Serra da Maria Comprida, Morro do Cuca, 1500 m, s.dat. (fr.), M. A. Moraes 151 & E. Fernades (K!, RB – image! [RB462639, barcode: RB00593594]); Nova Friburgo (fl.), Luis Edmundo Pâes 111 (RB – image! [RB56549, barcode: RB00073197]); Município de Santa Maria Madalena, Santa Maria Madalena, Pedra Dubois, 21°58'S 42°01'W, 900 – 1195 m, 22 Feb. 1983 (buds), T. C. Plowman & H. C. de Lima 12870 (K!; NY [NY533109]); Pedra do Cônico bei Nova Friburgo, 1300 m, Jan. 1898 (buds & fl.), E. Ule Herb. Bras. 4551 (B – image! [barcode: B109009682], holotype; HBG – image! [barcode: HBG506927], isotype). São Paulo: Serra da Bocaina, 1800 m, 18 May 1957 (st.), A. C. Brade s.n. (RB – image! [RB73894, barcode: RB00073182]). S.loc.: (st.), A. de Saint-Hilaire 735 (P! [barcode: P05616955]).

habitat. Forest, including secondary forest, at c. 1000 – 1750 m elevation.

conservation status. The extent of occurrence (EOO) and area of occupancy (AOO) (see IUCN 2001, 2016), calculated in GeoCAT (online), suggest a threat status of either NT (Near Threatened) or EN (Endangered) respectively. Without more information about the distribution of this plant, its occurrence within protected areas and possible threats to its habitat, a provisional threat status of NT seems appropriate.

notes. When publishing the basionym of Lamanonia ulei, Engler (1928) distinguished his new species by the dense, grey, felty indumentum on the lower surface of the leaflets (“Blattchen unterseits dicht graufilzig”). This layer of small hairs is quite distinct from the pubescent to tomentose indumentum seen in some specimens of L. ternata, and does indeed differentiate L. ulei from all the remaining species in the genus. The dense arrangement of the hairs means that while the secondary veins can be seen on the lower leaf surface in L. ulei, the tertiary and quaternary venation are not visible.

Lamanonia ulei is not a common tree and I have seen material or images of only eight collections. The description given here is based primarily on Plowman & de Lima 12870 and Moraes 151, the specimens available at K, so it is possible that a greater range in characters, such as leaf size, will be found when other collections are included. The number of stamens per flower needs to be determined from better material than is currently available to me.

The earliest of the collections listed here was made by Saint-Hilaire between 1816 and 1822. This sheet (P05616955) bears a label in the bottom left-hand corner stating “Rameau d’une Cunoniacée, sans fleurs et sans fruits, non décrite dans la flore” in Cambessèdes’s hand. No locality is indicated on the sheet other than Brazil, nor is a catalogue number given. The material appears to be from a sapling as it has much larger leaflets, longer internodes and more persistent stipules than normally seen in collections from mature plants with flowers and/or fruits.

Index to the types of names in Lamanonia, Belangera and Polystemon

Balansa 3183 — lectotype (G) & isolectotypes of B. paraguaiensis Pamp.

Balansa 4752 — syntypes, now lectoparatypes of B. paraguaiensis Pamp.

Blanchet 3253 — lectotype (G) & isolectotypes of B. denticulata Moric. & L. denticulata (Moric.) Kuntze

Glaziou 2495 — holotype (FI) & isotypes of B. speciosa Cambess. forma pubescens Pamp.

Glaziou 8247 — lectotype (G) & isolectotypes of B. chabertii Pamp. & L. chabertii (Pamp.) L. B. Sm.

Glaziou 17623 — lectotype (B) & isolectotypes of B. grandistipularis Taub. & L. grandistipularis (Taub.) Taub.

Glaziou 21119a — specimen cited under B. hirta, nom. inval.

Hassler 6583a — lectotype (G) & isolectotypes of B. tomentosa Cambess. var. calvata Chodat & Hassl.

Hassler 6583b — syntypes, now lectoparatypes of B. tomentosa Cambess. var. calvata Chodat & Hassl.

Martius Herb. Flora Brasiliensis 93 — lectotype (G) & isolectotypes of B. glabra Cambess. var. intermedia Pamp.

Ratter et al. 3524 — holotype (UEC) & isotypes of L. brasiliensis Zickel & Leitão

Saint-Hilaire (B1 –) 107 — syntype, now lectoparatype of B. tomentosa Cambess.

Saint-Hilaire (B –) 2073 — lectotype (MPU) & isolectotypes of B. tomentosa Cambess. & L. tomentosa (Cambess.) Kuntze

Saint-Hilaire C (or C1) – 66 — lectotype (MPU) & isolectotypes of B. glabra Cambess. & L. speciosa (Cambess.) Kuntze

Saint-Hilaire (C1 –) 133 — lectotype (P) & isolectotypes of B. cuneata Cambess. & L. cuneata (Cambess.) Kuntze

Saint-Hilaire C – 342 and C – 342bis — syntypes, now lectoparatypes of B. tomentosa Cambess.

Saint-Hilaire C – 343 — syntype, now lectoparatype of B. tomentosa Cambess.

Saint-Hilaire C (or C2) – 1204bis — possibly lectoparatypes of B. tomentosa Cambess.

Saint-Hilaire D – 122 — syntype, now lectoparatype of B. glabra Cambess. & L. glabra (Cambess.) Kuntze

Saint-Hilaire s.n. “Penha” — lectotype (P) & isolectotypes of B. speciosa Cambess. & L. glabra (Cambess.) L. B. Sm.

Sello(w) s.n. — lectotype (G) & isolectotypes of P. pentaphyllus D. Don

Sello(w) s.n. — lectotype (G) & isolectotypes of P. triphyllus D. Don

Ule Herb. Bras. 4551 — lectotype (B) & isolectotype of B. ulei Engl. & L. ulei (Engl.) L. B. Sm.