Introduction

During October and November 2015, the author was fortunate to make a second study visit to the Herbario Nacional de Bolivia (LPB), Universidad Mayor de San Andrés, La Paz, Bolivia, to continue with his work on his Annotated Checklist of the Compositae of Bolivia (Hind 2011). On this occasion, amongst other things, I had the opportunity to go through several more recent collections made in the Parque Nacional Madidi, Prov. Tamayo, Depto. La Paz, Bolivia (material collected as part of a long-term collaboration between LPB and Missouri Botanical Garden — MO). Included within the collections were four of an unusual looking species of what appeared to belong to Werneria Kunth (Compositae: Senecioneae: Senecioninae). Other than one evidently alcohol-treated collection, the other three show distinctive purplish abaxial leaf surfaces, purple, sparsely to moderately, villous pedicels, phyllaries with the apex or apical half coloured purple, and white ray limbs markedly pinkish towards the abaxial surface of the apex. A loan of this material from LPB to K was secured, in order to study the specimens further, and prepare a line drawing of the taxon.

Werneria, in the original sense, can be recognised by the following broad diagnostic description: Plants rhizomatous perennials, or dwarf erect or decumbent subshrubs, rarely of solitary rosettiform plants. Leaves simple, alternate or rosulate, rarely distichous, bases often pseudopetiolate and frequently sheathing, or leaves somewhat fleshy, conspicuously 3-partite and narrowed into a pseudopetiolar base. Inflorescences sessile or scapose, single-headed, usually ebracteate, or bracts foliose to scale-like, glabrous or lanate. Capitula usually heterogamous and conspicuously radiate, or sometimes disciform, or rarely homogamous and discoid; involucres campanulate and ecalyculate; phyllaries uniseriate, linear, bases usually fused, apices acute or obtuse; receptacle flat to convex, glabrous and epaleaceous. Ray florets (when present) few to many, female and fertile, ray limbs often appearing entire or scarcely 3-toothed, white or yellow, often discolorous and reddish, pink, or purplish abaxially. Disc florets few to numerous, hermaphrodite, corollas 5-lobed, usually yellow, sometimes off-white or purplish; style arms truncate with sweeping hairs in a crown. Achenes usually glabrous, sometimes setuliferous, setulae of twin-hairs, these long and villous; carpopodium inconspicuous and annular; pappus setae uniseriate, setae persistent and finely barbellate, usually white, whitish or rarely purplish or pinkish.

A comparison of the material mentioned above with the other species listed and accepted by Rockhausen (1939b), Hind (2011), Jørgensen et al. (2005), and Jørgensen (2014), for Bolivia, and those taxa in neighbouring Peru (Beltrán 2017; Beltrán & Leiva 2018), showed no obvious match in the genus Werneria. Analysis of the material in K, and subsequent contemporary studies, have shown this material to be Werneria glandulosa Wedd. (c.f. Calvo et al. 2020a: 33 – 35). Rockhausen (1939b: 334) listed the name under ‘Species incertae et excludendae. A. Species incertae’, with the note that ‘Von folgenden Arten hatte ich keine Material und keine Originale vorliegen. Fernerhin was es nicht möglich, dieselben nach der Beschribung sicher einzureihen.’ (= ‘I had no material and no originals of the following species. Furthermore, it was not possible to classify them with certainty according to the description.’). In adding Werneria glandulosa to the final version of my Annotated Systematic Checklist of Bolivian Compositae (Hind 2011), it became clear that there was a nomenclatural issue with continuing use of the current concepts of Werneria sensu stricto and Xenophyllum V.A.Funk.

Werneria, described in Nova Genera et Species Plantarum (Kunth 1818: 148), contained six species now considered to belong to two genera, Werneria sensu stricto and Xenophyllum (22 spp. — Funk 1997c; Calvo & Moreira-Muñoz 2020). Kunth did not designate a type, nor did he recognise any infrageneric ranks, and the generic description covered the characters of all the species described. The current record for Werneria in Index Nominum Genericorum (Farr & Zijlstra 1996+) indicates that there is no stated or selected type; from my discussion below, this is shown to be incorrect, as suggested by the literature (Rockhausen 1939a; Cabrera 1971; Funk 1997a, Funk 1997c; Freire & Ariza Espinar 2014; Linares-Perea et al. 2014; Calvo et al. 2017; Beltrán & Leiva 2018; Calvo & Meneses 2019; Calvo & Moreira-Muñoz 2019; Calvo et al. 2020a, b).

The issue

Lectotypification of Werneria Kunth

There is a categorical statement in Rockhausen’s revision of Werneria: ‘Gattungstypus ist Werneria rigida.’ (Rockhausen 1939a: 249), and later (p. 293) he specified the type of the name W. rigida as a Humboldt & Bonpland collection. His description of the species is not in conflict with Kunth’s generic description, and there was no sense of doubt over his statement of the generitype. He placed the species in his ‘subgen. Euwerneria (Gay) Rockh. sect. Aciculares Rockh.’; both infrageneric concepts contain the nomenclatural type of the genus (the generitype) and would now be considered as subgen. Werneria and sect. Werneria. This typification appears to have been overlooked, or ignored, by later authors, as many have only cited the second part of Rockhausen’s revision, and continue to use a nomenclaturally incorrect concept of Werneria, unaware of the implications (e.g. Linares-Perea et al. 2014; Calvo et al. 2017; Beltrán & Leiva 2018; Calvo & Meneses 2019; Calvo & Moreira-Muñoz 2019; Calvo et al. 2020a, b).

Three further statements of the ‘lectotype’ of Werneria have been made:

  1. 1)

    Although Cabrera’s early revision of the Argentinian species (Cabrera 1948) mentioned nothing about the type, he later (Cabrera 1971: 275) stated that the lectotype was W. graminifolia Kunth. He repeated this statement (Cabrera 1978: 469), whilst still not stating who designated it; it is assumed that it was Cabrera himself. Calvo et al. (2020a: 6) stated that it was Cabrera in his Flora Patagonica account (Cabrera 1971: 275). Werneria graminifolia, an Ecuadorian endemic, occurs in none of Cabrera’s treatments of the genus.

  2. 2)

    In recognising Misbrookea V.A.Funk, Funk (1997a: 110, 1997b) stated that the ‘type species’ was ‘Werneria nubigena H. B. K., and [the genus] is referred to hereafter as Werneria s.str.’ In both cases she only cited the second part of Rockhausen’s paper on Werneria (Rockhausen 1939b), possibly unaware of the first part where the type citation was made and several new infrageneric taxa described (Rockhausen 1939a). Later, Funk (1997c: 235), in her description and synopsis of Xenophyllum, again mentioned that ‘Werneria s.l.’ contained at least ‘three identifiable groups ... that can be recognized as distinct genera.’ One of these ‘contains the type species, Werneria nubigena Kunth, and can now be referred to as Werneria s. str. ...’; Xenophyllum dactylophyllum (Sch.Bip.) V.A.Funk was designated as the type of Xenophyllum. In none of the accounts is there any indication who designated W. nubigena as the lectotype of Werneria; it can be assumed to have been selected by Funk, unless evidence is eventually found to the contrary.

  3. 3)

    Freire & Ariza Espinar (2014: 220) stated that the lectotype of Werneria was W. graminifolia, as indicated by Cabrera (1971: 275), but also noted the other citations by Rockhausen (but only citing the second part of Rockhausen’s revision, Rockhausen 1939b — where the type citation does not occur) and Funk (1997b). No additional explanation was provided as to why W. graminifolia was accepted as the lectotype over the selection by Rockhausen.

No type was designated or mentioned by Gray (1861), or Blake (1928), when several new species were added in their short synoptic treatments. Tkach et al.’s statement, that Werneria nubigena was the ‘type species of Werneria Kunth’, with reference to the Index Nominum Genericorum website (Tkach et al. 2016: 101), is also without foundation. The website simply records the type as ‘non designatus’, and gives no indication of a lectotype (ING, last accessed 10th October 2020).

The position of Oresigonia

Lessing (1832: 393) appears to have been the first author to mention the generic name Oresigonia, albeit referring to two completely different taxa. He first used it under his treatment of Werneria, alongside Werneria disticha Kunth, in referring to Willdenow’s manuscript name, Oresigonia latifolia. It was used lower down on the same page in reference to Culcitium Humb. & Bonpl., albeit as ‘Oresigonia Schlechtd. im Mag. naturf. Fr.’ Lindley (1836: 261) also attributed Willdenow’s mss. name to Werneria and, immediately after, Schlechtendal’s to Culcitium, a position reflected by de Candolle (1838: 323) with the generic name as ‘Oresigonia Willd. herb. ex Less.’ in the synonymic heading to his treatment of Werneria, and on the following page as ‘Oresigonia Schlecht. in Berl. naturf. mag. ex Less.’ in the synonymic heading to his treatment of Culcitium. The explanation of this was effectively provided by Blake (1937: 389 – 390), as simply an error on Lessing’s part, by wrongly referring to Schlechtendal’s genus Lasiocephalus (Schlechtendal 1818: 308) as Oresigonia. Both Lindley and de Candolle had followed Lessing, as did other contemporaneous compendia (e.g. Endlicher 1841: 244), and those from much later (e.g. Baillon 1882: 269 & 261 respectively). I consider this is significant, as Willdenow’s annotations of material seen in B-W are only on sheets of Werneria sensu lato; both species of Lasiocephalus are only annotated as such by Willdenow.

Recognition of the new genus Rockhausenia

Jørgensen (2014: 380) recorded 15 species of Werneria from Bolivia, based in part on the eight previously recorded for the Parque Nacional Madidi (Jørgensen et al. 2005); Misbrookea strigosissima (A.Gray) V.A.Funk (based on W. strigosissima A.Gray), Xenophyllum ciliolatum (A.Gray) V.A.Funk (based on W. ciliolata A.Gray), and X. dactylophyllum (Sch.Bip.) V.A.Funk (based on W. dactylophylla Sch.Bip.), have since been excluded from the original Madidi total of 11. I now recognise 18 taxa from Bolivia belonging to Werneria sensu Funk (1997b), Jørgensen (2014: 380), and Calvo et al. (2020a, b).

Misbrookea, Werneria and Xenophyllum form a well-supported subclade that is sister to a further strongly supported subclade formed by Charadranaetes Janovec & H.Rob., Dorobaea Cass., Garcibarrigoa Cuatrec., Jessea H.Rob. & Cuatrec., Pseudogynoxys (Greenm.) Cabrera and Talamancalia H.Rob. & Cuatrec. (Pelser et al. 2007). Anticona E.Linares, J.Campos & A.Galán (based on Werneria glareophila Cuatrec.) has never been sampled, so its relationship has yet to be elucidated. The generic disposition of the wernerias needs to be reconsidered. Since Rockhausen’s lectotypification is valid and, in recognising Xenophyllum, Funk created a superfluous generic synonym for the wernerias sensu stricto. Providing a solution is a purely nomenclatural issue: the transfer of the remaining wernerias to a new genus, Rockhausenia, which is proposed below.

Synoptic treatment of Rockhausenia

I have refrained from providing a key to species as there is a perfectly adequate key to all species in Calvo et al. (2020a), and a country-specific key is available for Peru by Beltrán (2016) and Argentina (Freire & Ariza Espinar 2014). Adequate keys to taxa originally considered to be in Xenophyllum were provided by Calvo & Moreira-Muñoz (2019) expanded from Funk (1997c).

In the following synoptic treatment of Rockhausenia, under each accepted name synonyms are provided in chronological order. For each name references are provided, and a type citation given for all basionyms. The type citation is a classical one, and is simply a transcription from the basionym protologue. The location of known types is stated (acronyms following Thiers [continuously updated]) and, when available, barcode numbers of type material given. In instances where institutional barcodes unnecessarily start with six or more zeros the barcode number is concatenated to the remaining digits — the suggestion that institutions could ever have that number of specimens when fully digitised is, frankly, ludicrous. Herbarium or manuscript names are also provided where they are relevant to the history of taxa concerned and have appeared in the literature. A full nomenclator of names is also provided.

HAL type material

Whilst Hind & Jeffrey (2001) indicated duplicates of the Humboldt & Bonpland collections exist in both B-Willd and P, there are many duplicates to be found elsewhere. I have come across few duplicates in RB during an earlier study visit but, as many virtual herbaria have now come online over the last few years, a large set is clearly in HAL. Tkach et al. (2016) have stated that the Humboldt & Bonpland specimens in HAL are duplicates of specimens in the Willdenow herbarium (B-W). The collections are unnumbered, having reached the University herbarium via Schlechtendal junior (D. F. L. von Schlechtendal 1794 – 1866); they are all considered syntypes. Handwritten sheet annotations and labelling are considered to be those of Schlechtendal senior (D. F. C. von Schlechtendal 1767 – 1842), whereas the main sheet labels are modern printed labels based on the protologue details and the B-W source of the material.

Rockhausenia D.J.N.Hind, gen. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77300544-1

Type: Werneria nubigena Kunth = Rockhausenia nubigena (Kunth) D.J.N.Hind

Description

Perennial rosulate, sometimes clump-forming, herbs. Rootstock often rhizomatous, lanate or villous amongst leaf bases. Leaves rosettiform, very rarely distichous, sessile, base usually expanded and partially sheathing or amplexicaul, blade linear to linear-lanceolate or spathulate, sometimes fleshy, glabrous or with villous to lanose leaf bases, rarely lamina pubescent, hairs with an eglandular long-flagelliform apical cell on top of a multicellular uniseriate base of thickened-walled cells, margins entire, lobed or pinnatisect, apex rounded, obtuse or acute to acuminate. Inflorescence of solitary, sessile or short- to long- pedicellate to scapiform capitula borne in centre of rosette, pedicel or scape glabrous or pubescent, ebracteolate or bracteolate, bracteoles scale-like. Capitula heterogamous and radiate or, very rarely, disciform or occasionally homogamous and discoid; involucre campanulate, ecalyculate; phyllaries 8 – 21, uniseriate, connate for at least half of length, glabrous or pilose, apices often purplish, penicillate; receptacle flat or convex, glabrous, rarely alveolate, epaleaceous. Ray florets 13 – 21, female, fertile, uniseriate, but sometimes appearing biseriate, ray limb 2 – 3-toothed (sometimes scarcely appearing denticulate), white or cream adaxially, usually discolorous and reddish abaxially, or yellow (and then never discolorous), very rarely purple. Disc florets hermaphrodite, 20 – 200, fertile, corollas actinomorphic, tubular, glabrous, throat narrowly campanulate, 5-lobed, white or yellow, often conspicuously yellow in bud but corollas clearly creamish or whitish when lobes open; anthers yellow, or black or violet, anther collar balusterform, apical anther appendages oblong-lanceolate, obtuse, basal anther appendages sagittate; style base with basal node, style yellow, cream, white or red, style arm apices truncate to obtuse or rounded, papillate, papillae forming a corona. Achenes cylindrical, 8 – 10-ribbed, glabrous or setuliferous, setulae often long and sericeous; carpopodium usually minute and annuliform; pappus setae uniseriate, capillary, barbellate, persistent, bases connate into ring, usually white, sometimes turning purplish with age.

recognition. Species of Rockhausenia are mostly low, perennial mat- or cushion- forming perennials, very rarely aquatics or marginals, although sometimes also found as isolated rosettiform plants/individuals. Species usually have entire (rarely pinnatifid/lobed) concolorous leaves, and characteristically have solitary, sessile or pedicellate, capitula, an involucre of basally fused uniseriate phyllaries, and discolorous ray floret limbs that are usually white (rarely yellow) adaxially and pinkish or purplish abaxially; three species (of the 27 in the genus) possess discoid capitula. In contrast, Werneria spp. usually form dense or loose hummocks or mat, with the rhizomes and upper stems densely leafy for some length, with the exposed leaves green and those within the tussock or mats often brown or blackish (or sometimes whitish). The capitula in Werneria are heterogamous and radiate (very rarely are the rays greatly reduced or absent and the heads effectively disciform), ray limbs are usually white or very rarely violet-purple or yellow. Misbrookea, a rhizomatous rosettiform monotypic genus from Bolivia and Peru, is easily recognised by the long (3 – 5 mm) whitish strigose hairs found on both leaf surfaces and on the involucres (vs glabrous leaves and involucres or villous scape and involucres in few Rockhausenia spp.), and style arm apices with long multicellular hairs.

distribution. The genus Rockhausenia is a New World endemic, essentially restricted to South America, currently with 27 species of the high Andes (of Argentina, Bolivia, Chile, Colombia, Ecuador, Peru and Venezuela), with the exception of the most widespread species, Rockhausenia nubigena (Kunth) D.J.N.Hind, which is found from southern Mexico and Guatemala south to southern Bolivia.

New combinations in Rockhausenia

For convenience, the taxa appear in the order treated by Rockhausen (1939a, b) together with several additional species added where appropriate.

Rockhausenia D.J.N.Hind subgen. Anactis (J.Rémy) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300545-1

Werneria Kunth [unranked] § II. Anactis J.Rémy, Fl. Chilena (Gay) 4: 216 (1849). Note: The symbol used by Rémy is a typographical section symbol |§| Elsewhere Rémy used the abbreviation ‘Secc.’, or even in expanso, ‘Seccion’, with the || symbol as a clear lower rank — in Cardamine for example (1: 109), or Nasturtium (1: 117) — where it was certainly used as a section! However, the word ‘Seccion’ has been used as another rank, below tribe, and above genera.

Werneria Kunth subgen. Anactis (J.Rémy) Rockh. (Rockhausen 1939a: 265).Werneria Kunth subgen. Anactis (J.Rémy) Rockh. sect. Pinnatifoliae Rockh., Bot. Jahrb. Syst. 70 (2): 265 (Rockhausen 18 August 1939a). Note: Considered as ‘nom. inval.’ by Calvo et al. (2020a: 6) under Art. 22.2 of the current Code (Turland et al. 2018), as it contained the type of the genus. This is untrue, as the type of the genus was, in their mind, Werneria graminifolia. Since this is clearly not the case, the name is perfectly valid.

Type: Werneria pinnatifida J.Rémy = Rockhausenia pinnatifida (J.Rémy) D.J.N.Hind

1. Rockhausenia pinnatifida (J.Rémy) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300548-1

Werneria pinnatifida J.Rémy, Fl. Chilena (Gay) 4 (2): 216 (1849). Type: [Chile:] ‘Se cria tambien en las cordilleras de los Patos.’ Note: This reference is to the locality provided after Werneria rhizoma — ‘Se halla en las cordilleras de Ovalle, á los Patos.’ Lectotype (selected by Freire & Ariza Espinar 2014: 223): P(02088545 – Gay, s.loc., s.n., the sheet label is marked ‘216’ on the top left hand corner, signifying the page upon which it was published). Note: The image of B†(F0BN15815 — showing it was mounted with a specimen of ‘Werneria incisa’) lacks the original sheet label merely stating ‘Atacama’ on the paper ruler. Calvo et al. (2020a: 63) declared the type collection as ‘C. Gay 703’, presumably based on one of the listed isolectotypes, although not stating which: ‘F(974712), GH[(00936611 — s.n.)], P[(04023489 — numbered ‘450’ on a small paper label, det. ‘ISOLECTOTYPE’ by Calvo, ‘Sept. 2019’)], SGO(000006432)’.

Werneria heteroloba Wedd. (Weddell 1856: 88). Types: ‘Hab. Pérou: Cordillères du département de Cuzco! (Gay). – Bolivie: sur les pelouses marécageuses de la Lancha!, Cordillère de La Paz, avec le W. pygmaea (Wedd.); bord des eaux, dans les montagnes des lagunas de Potosi! (d’Orbigny, no 1415).’ Syntype: Gay s.n., P(06898872, 06898874 — both specimens unmarked as syntypes). Syntype: d’Orbigny 1415, G(00305497), P(02088558, 02088560). Syntype: Weddell, P(00869754 — marked as ‘1’ on the sheet label by Weddell, although only the material that is ‘Werneria pygmaea’ is numbered ‘2’ under each specimen on the sheet itself). Lectotype (selected by Freire & Ariza Espinar 2014: 222, effectively second-step): d’Orbigny 1415, P(02088558); isolectotypes: G(00305497), P(02088560). Note: Rockhausen (1939b) used the word ‘Typus!’ after his citation of the d’Orbigny 1415 collection, although it is unclear from his account if this was merely a citation of a syntype or an effective lectotypification since the other syntypes were not mentioned.

Werneria obtusiloba S.F.Blake (1928: 489). Type: ‘Peru: In sandy soil, with cushion and rosette plants, cordillera east of Carumas, Prov. Moquegua, alt. 4500-4600 m., 7-8 Mar. 1925, A. Weberbauer 7362’. Holotype: F(552591); isotypes: G(00305496), US(44298 = 00037327).

Werneria heteroloba Wedd. var. obtusiloba (S.F.Blake) Rockh. (Rockhausen 1939b: 283).

Werneria heteroloba Wedd. f. microcephala Rockh. (Rockhausen 1939b: 283). Type: ‘Chile: Lago Chungara, in Sümpfen, 4550 m ü.M. (C. Troll Nr. 3222 – März 1927.)’. Holotype: B†. Note: Calvo et al. (2020a: 63) lectotypified the name based on the destroyed type material. There being no extant original material a neotype should have been designated, although this I consider unnecessary.

distribution. Argentina (Catamarca, Jujuy, Salta, San Juan, Tucumán), Bolivia (Cochabamba, La Paz, Oruro, Potosí), Chile (Antofagasta, Arica y Parinacota, Atacama, Coquimbo, Tarapacá), Peru (Ancash, Arequipa, Cusco, Huancavelica, Moquegua, Puno).

note. Beltrán (2017: 48) considered Werneria obtusiloba was separate from W. heteroloba, a position not accepted here.

2. Rockhausenia solivifolia (Sch.Bip.) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300549-1

Werneria solivifolia [as solivaefolia] Sch.Bip., Bonplandia 4 (4): 53 (Schultz Bipontinus 1856). Type: [Peru:] ‘Nun kimmen noch zwei neue Wernerien zur Aburtheilung, welche mir unter [Lechler] Nr. 1710, in uliginosis Cord. de Tuno, Mai 1854 in Cherleria ähnlichen Rasen, in welchen die kleinen Köpfchen versenkt waren, wachsend, mitgetheilt wurden.’ Note: In the subsequent listing (Hohenacker 1856: 55) it appears that Werneria brachypappa was based on Lechler 1710a, and W. solivifolia on Lechler 1710b. Type material: K(000527612 — as Lechler ‘1710’ on the capsule, separated from the block of Distichia muscoides Nees & Meyen (Juncaceae) and placed in the lh capsule). Note: Although Schultz Bipontinus provided a Latin description on P(02088539) there is no material of this species visible on the sheet. Lectotype (selected by Calvo et al. 2020a: 88): K(000527612 — only material in the small capsule on the sheet).

distribution. Bolivia (La Paz, Potosí), Chile (Arica & Parinacota), Ecuador (Pichincha), Peru (Arequipa, Huancavelica, Junín, Lima, Moquegua, Puno, Tacna).

Rockhausenia D.J.N.Hind subgen. Rockhausenia

Werneria Kunth subgen. Werneria [as Euwerneria (Gay) Rockh.] sect. Graminifoliae Rockh., Bot. Jahrb. Syst. 70 (2): 266 (Rockhausen 18 August 1939a).

Rockhausenia D.J.N.Hind subgen. Rockhausenia sect. Integrifoliae (Rockh.) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300550-1

Werneria Kunth subgen. Anactis J.Rémy sect. Integrifolia Rockh., Bot. Jahrb. Syst. 70 (2): 265 (Rockhausen 18 August 1939a). Lectotype (selected by Calvo et al. 2020a: 6): Werneria carnulosa A.Gray = Rockhausenia carnulosa (A.Gray) D.J.N.Hind

3. Rockhausenia carnulosa (A.Gray) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300552-1

Werneria carnulosa A.Gray, Proc. Amer. Acad. Arts 5: 140 (1861). Type: ‘High Andes of Peru. [Wilkes]’. Holotype: GH; isotype: US(00037298).

distribution. Peru (Ancash, Huánaco, Junín, Lima, Pasco).

Rockhausenia subgen. Rockhausenia sect. Rockhausenia

Type: Werneria nubigena Kunth = Rockhausenia nubigena (Kunth) D.J.N.Hind

4. Rockhausenia nubigena (Kunth) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300553-1

Werneria nubigena Kunth in Humb., Bonpl. & Kunth, Nov. Gen. Sp. Pl. 4 (ed. folio, fasc. 16): 151 (1818). Type: [Ecuador:] ‘Crescit in frigidis montis Chimborazi, alt. 1700 hex. (Regno Quitensi.) ... Floret Junio.’ [Humboldt & Bonpland P-Bonpl - ‘3191. Chimborazo’; B-W: ‘3191’] Holotype: P-Bonpl (P00320184); isotypes: B-W(16428-01), HAL(0113455), P(04319323). Note: Tkach et al. (2016: 101) considered the material in HAL a syntype, but also stated that this was the ‘type species of Werneria Kunth’, referring to ‘ING’, this referring to the ING website.

Werneria disticha Kunth (1818: 151). Type: ‘Crescit in summo monte ignivomo Antisanæ, alt. 2100 hex. ... Floret Junio. [Humboldt & Bonpland 2272]’ Holotype: P-Bonpl. (00320185); isotypes: B-W(16429-01), HAL(0113454 — lacking any number or locality on the label), P(02088530, 02088542). Note: Tkach et al. (2016: 101) considered the material in HAL a syntype.

Oresigonia latifolia Willd. ex DC. (de Candolle 1838: 324), nom. nud. pro syn. based upon Willdenow's herbarium name applied to the sheet in B-W(16429-01).

Werneria mocinniana DC. (de Candolle 1838: 324). Type: ‘in Mexico ad cacumen montis cuchumatanes. Orisaba acaulis Moc. Sess. et Cerv. icon fl. mex. ined. ... (v. ic. pict.)’ Note: It is clear that de Candolle only referred to the albeit unpublished plate. However, Funk has labelled a herbarium specimen, MA(606589), as the lectotype of this name; no formal lectotypification of this name has been located.

Oribasia acaulis Moc., Sesse & Cerv. ex DC. (de Candolle 1838: 324), nom. nud. pro syn.

Werneria nubigena Kunth var. α vulgaris Wedd. (Weddell 1856: 80), pro W. nubigena Kunth.

Werneria nubignea Kunth var. β latifolia Wedd. (Weddell 1856: 80), nom. illeg. pro Werneria disticha Kunth

Werneria stuebelii [as stübelii] Hieron. (Hieronymus 1895: 362). Type: ‘Peruvia: crescit prope Challuayacu inter Pacasmayo et Moyobamba, alt. s. m. 3400 m, ubi floret mense Aprili-Junio ([Stübel] coll. peruv. n. 55c).’ Holotype: B†(F0BN015823). Note: Beltrán (2017: 58) stated that P(02088547) was the lectotype of this name; it is not clear who published this lectotypification as Funk’s determinations are unpublished. This is one of two barcoded specimens on a sheet with one collecting label, that of Lechler 2212, and the sheet previously determined as lectotype of W. staticifolia, by Funk — both specimens are conspecific; nowhere on the sheet is there a declaration that this is Stübel 55c.

Oresigonia grandiflora Willd. ex Rockh. (Rockhausen 1939b: 301), nom. nud. pro syn., based on B-W(16428 — Humboldt & Bonpland 3191, an isotype of Werneria disticha Kunth).

[Oresigonia latiflora Willd. ex Rockh. (Rockhausen 1939b: 301) — often cited with this authority having ignored its appearance in de Candolle’s Prodromus, q.v.]

distribution. Bolivia (Cochabamba, La Paz, Tarija), Colombia (?), Ecuador (Azuay, Bolívar, Cañar, Carchi, Chimborazo, Cotopaxi, Imbabura, Loja, Napo, Pichincha, Sucumbíos, Tungurahua), Guatemala (Huehuetenango, Quetzaltenango, San Marcos), Mexico (Chiapas), Peru (Amazonas, Ancash, Apurímac, Arequipa, Ayacucho, Cajamarca, Cusco, Huancavelica, Huánaco Junín, La Libertad, Lambayeque, Lima, Moquegua, Pasco, Piura, Puno, San Martín).

5. Rockhausenia plantaginifolia (Wedd.) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300554-1

Werneria plantaginifolia Wedd. ex Klatt, Ann. K. K. Naturh. Hofmus. Wien 9 (3): 367 (1894). Type: [Bolivia:] ‘Hab.: Prov. Larecaja, vinciniis Sorata, prope las trincheras de Chiliata, in graminosis, Reg. alpina, 3800 m., leg. Mandon, Mart.-Mai 1859, Nr. 89.’ Type material (see note): B†(F0BN015818), G(00305795 — databased as the ‘Neotype’), K(000527610), LL(00374331), P(02690589, 04318258, 04388253), S-R-6526. Lectotype (selected here): P(04318258). Note: Amongst the type material, that selected as the lectotype is the only material of Mandon 89 with the same locality details as provided in the protologue. However, the locality was probably not fully transcribed by Klatt, as it reads ‘Viciniis Sorata: propé las trincheras de Chiliata [sic!] Sa. [sic!] in graminosis. Alt. Reg. alpinae 3800 m. Mart-Mai 1859.’ The remainder of the material, although of Mandon 89, is labelled as ‘Viciniis Sorata; adscensu à Millipaya ad nives, in graminosis. Alt. Reg. alpina: 3800 m. Mai. 1859.’ [trincheras = trenches][‘Chiliata’ is also written ‘Chillata’ on other Mandon collection labels.] [Millipaya (3850 m) is SW of Illampu.].

Werneria plantaginifolia Wedd. ex Klatt var. macrocephala Cuatrec. (Cuatrecasas 1953a: 153). Type: ‘Peru: Dep. Apurimac, prov. Abancay: Punas de Ampay, 3900 m alt., colect. Jun. 1938 Cesar Vargas 1047. (Holotypus, F – no barcode number available [Photo Neg. No. 49324]).’

distribution. Bolivia (La Paz), Peru (Apurímac, Ayacucho, Cusco, Junín).

6. Rockhausenia staticifolia (Sch.Bip.) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300555-1

Werneria staticifolia [as staticaefolia] Sch.Bip., Bonplandia 4 (4): 53 (Schultz Bipontinus 15 Feb. 1856). Type: [Peru:] ‘St. Gavan in summis Cordil. jugis Jul. 1854: Lechler! Nr. 2212.’ Holotype: P(02088546 & 02088547 – the type sheet has two specimens and two barcodes although it is likely that this is simply one collection and specifying one as a lectotype is unnecessary; the sheet is det. Funk, September 2011, as the lectotype; the pencilled lines and arrows on the sheet are not self-explanatory, as apparently only arrowing the labels that do not indicate two different collections); isotypes: B†(F0BN015822 — lacking an image of the label), G(00305495), K(000527608), M(0147066), P(00711471; 02088548 — det. Funk, September 2011, as isolectotype), S-R-6528. Beltrán (2017: 58) only cited ‘Lectótipo: P. P0208854’, failing to provide the last digit and certainly not specifying which of the two sheets this applied to; if it was only to the sheet determined by Funk neither specimen was mentioned.

Werneria nubigena Kunth var. δ caulescens Wedd. (Weddell 1856: 81). Type: not clearly stated. Weddell (1856) noted ‘Pérou: Cordillères des départemnts d’Arequipa et de Cuzco (α, γ, δ et ε) (Gay, Dombey) ...’ as the only mention of material against the varietal name but without indicating to which taxon the material referred as two subvarieties (subvar. leioscapa and erioscapa) were also described, q.v.

Werneria nubigena Kunth var. δ caulescens Wedd. subvar. leioscapa Wedd., (Weddell 1856: 81). Type: not indicated.

Werneria nubigena Kunth var. δ caulescens Wedd. subvar. erioscapa Wedd. (Weddell 1856: 81). Type: not indicated.

Werneria staticifolia Sch.Bip. var. β celmisioides Wedd. (Weddell 1856: 82). Type: [Peru:] ‘sur les sommités de la Cordillère de Tabina (Lechler, no 2111).’ Holotype: P; isotype: K(000527609). Note: Although, at some length, Rockhausen (1939b: 307 – 308) discussed this variety, the complete lack of any pubescence at the base of the involucre suggests it is best placed under R. plantaginifolia, although Schultz Bipontinus clearly noted that the achenes of Lechler 2111 were glabrous, a feature of R. staticifolia! However, Lechler 2111a, labelled as ‘Werneria celmisioides Sch.Bip.’, is in fact a specimen of Hypochaeris, and not a Werneria or Rockhausenia.

Werneria caulescens (Wedd.) Hieron. (Hieronymus 1895: 363), isonym. Note: Hieronymus (1900: 74) oddly stated his combination was based on Weddell’s variety, but his concept was not that of Grisebach’s; Grisebach had based his combination on the same basionym and did not exclude any material. Rockhausen (1939b: 337) indicated this isonym was W. staticifolia.

Werneria caulescens (Wedd.) Rusby (1907: 398), isonym.

distribution. Bolivia (La Paz), Peru (Cusco).

7. Rockhausenia villosa (A.Gray) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300556-1?Werneria nubigena Kunth var. ε dombeyana Wedd. (Weddell 1856: 81). Type: not clearly stated. Note: Weddell (1856: 81) noted ‘Pérou: Cordillères des départemnts d’Arequipa et de Cuzco (α, γ, δ et ε) (Gay, Dombey) ...’ as the only mention of material against the varietal name but without indicating to which taxon the material referred. Rockhausen (1939b: 338) suggested the possible synonymy under W. villosa. Beltrán (2017) indicated it was a synonym of W. nubigena.

Werneria villosa A.Gray, Proc. Amer. Acad. Arts 5: 139 (1861). Type: ‘High Andes of Peru near Alpamarca [Collections of the United States South Pacific Exploring Expedition under Captain Wilkes].’ Holotype: GH; isotypes: K(000527606), P(02088554), US(42760 = 00037309).

Werneria caulescens (Wedd.) Griseb. (Grisebach 1879a: 208, 1879b: 208). Note: Both the proposed combinations of Hieronymus (1895) and Rusby (1907) refer to material that Rockhausen (1939b: 337) considered belonged to W. staticifolia (= R. staticifolia), q.v.

?Werneria dombeyana (Wedd.) Hieron. (Hieronymus 1895: 363).

Werneria acerosifolia Hieron. (Hieronymus 1895: 363). Type: ‘Peruvia: crescit prope Fraileyacu inter Ventilla y Bayazan, collecta fuit in itinere inter Pacasmayo et Moyobamba, floret mense Maio ([Stübel] coll. peruv. n. 25a).’ Holotype: B†(F0BN015800). Note: Beltrán (2017: 56) considered this a synonym of W. pumila, along with W. canaliculata. Calvo et al. (2020a: 95 – 96) proposed the current synonymy, but refrained from neotypification (because of the poor state of the, now destroyed, type material illustrated in the Field Museum Berlin Type negative collection).

Werneria caulescens (Wedd.) Rusby (Rusby 1907: 398), isonym.

distribution. Argentina (Jujuy, Salta, Tucumán), Bolivia (Chuquisaca, Cochabamba, La Paz, Oruro, Potosí, Tarija), Peru (Ancash, Apurímac, Ayacucho, Cajamarca, Cusco, Huancavelica, Huánuco, Junín, La Libertad, Lambayeque, Lima, Pasco, Piura, Puno, San Martín).

8. Rockhausenia pumila (Kunth) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300557-1

Werneria pumila Kunth in Humb., Bonpl. & Kunth, Nov. Gen. Sp. Pl. 4 (ed. folio, fasc. 16): 148 (1818). Type: ‘Crescit cum praecedentibus.’ [Werneria rigida & humilis: Crescit in summis Andibus Quitensium, alt. 1800 – 2000 hex. [Humboldt & Bonpland]] Holotype: P(00320182); isotype: B-W(16430-01). Note: Nordenstam’s citation of the holotype in P (Nordenstam 1999: 314), Calvo et al. (2020a: 71) considered to be effective lectotypification.

Werneria densa Benth. (Bentham 1845: 211). [Colombia:] ‘[Hartweg] 1168. ... In sabulosis montis Antisana, altit. 15,000 ped.’ Holotype: K(specimen unbarcoded and still on loan to US — ex Herb. Benthamianum); isotypes: E(00413267), K(000527603 — ex Herb. Hookerianum), LD(1211958), P(02088562, 02088563), US(00037304). Note: Regrettably, the specimen chosen by Calvo et al. (2020a: 71) as the lectotype of the name was not the specimen used by Bentham in writing his account of the Hartweg collections — the material ex Herb. Benthamianum (Bentham 1839 – 57).

Werneria calyculata Turcz. (Turczaninow 1851: 204). Type: ‘In summis alpibus Quitensibus. Jameson coll. n. 618 – No. 809 ejusdem collectionis videtur ad W. frigidam HB. Kunth spectare.’ Type material: BM, G(00305796, 00305797), K(000374197 — mounted with Holton 399, Purdie s.n. and Pearce s.n. — ex ‘Bogata, 9-10000 ft March 1864’), KW(001001520 — ‘618/ From the summit of the Andes’ in Jameson’s hand, and additionally labelled ‘In summis alpibus Quitensibus/ Jameson coll: n. 618’ from the protologue). Lectotype (selected by Calvo et al. 2020a: 71): KW(001001520).

Werneria pumila Kunth var. β subspathulata Wedd. (Weddell 1856: 82), based on ‘W. densa Benth., Pl. Hartw., 214; Wlprs. Repert., VI, 254.’

Oresigonia angustifolia Willd. ex Rockh. (Rockhausen 1939b: 310), nom. nud. pro syn., based on B-W(16430-01 — an isotype of Werneria pumila Kunth)

distribution. Colombia (Auca, Caldas, Risaralda, Tolima, Valle del Cauca), Ecuador (Azuay, Bolívar, Chimborazo, Cotopaxi, Imbabura, Loja, Morana-Santiago, Napo, Pichincha, Tungurahua, Zamora-Chinchipe), Peru (Piura).

9. Rockhausenia graminifolia (Kunth) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300558-1

Werneria graminifolia Kunth in Humb., Bonpl. & Kunth, Nov. Gen. Sp. Pl. 4 (ed. folio, fasc. 16): 148 (1818). Type: ‘Crescit in alta planitie Antisanae, qua montem Chusolongi spectat, alt. 2100 hex. ... Floret Julio.’ Holotype: Humboldt & Bonpland 2243, P-Bonpl., P(00320183 — determined by Rockhausen 28 Aug. 1937); isotype: B-W(16431-01 — lacking original label and det. Willdenow as Oresigonia parviflora).

Werneria nubigena Kunth var. γ graminifolia (Kunth) Wedd. (Weddell 1856: 81).

Oresigonia parviflora Willd. ex Rockh. (Rockhausen 1939b: 312), nom. nud. pro syn., based on B-W(16431-01 — lacking original label, but det. by Willdenow).

distribution. Ecuador (Azuay, Bolívar, Cañar, Chimborazo, Cotopaxi, Imbabura, Napo, Pichincha, Tungurahua).

10. Rockhausenia apiculata (Sch.Bip.) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300559-1

Werneria apiculata Sch.Bip., Bonplandia 4 (4): 52 (Schultz Bipontinus 1856). Type: ‘Meine Werneria apiculata kommt unter 2 Nummern von sterilen Andenweiden vor.’ [Note: In a later list of determinations it is clear that these were Lechler 1737 and 1964 from Peru. Inference from Schultz Bipontinus (1856: 50 – 54) is that these were collected between May (cf. Lechler 1710[a] — Werneria brachypappa) and June 1854 (cf. Lechler 1963 — Achyropappus cryptocephalus).] Syntype: Lechler 1737, B†(F0BN15801), G(00305499), K(000527601), P(02088532, 02088533, 02088534), S-R-6521. Syntype: Lechler 1964, B†. Lectotype (selected by Freire & Ariza Espinar 2014: 224): Lechler 1737, P(02088534); isolectotypes as cited above. Note: Freire & Ariza Espinar (2014: 224) maintained this species as a variety of Werneria pygmaea.

Werneria pygmaea Gillies ex Hook. & Arn. var. ε apiculata (Sch.Bip.) Wedd. (Weddell 1856: 84).

distribution. Argentina (Catamarca, Jujuy, La Rioja, Salta), Bolivia (Chuquisaca, Cochabamba, La Paz, Oruro, Potosí, Tarija), Peru (Ancash, Apurímac, Arequipa, Ayacucho, Cajamarca, Cusco, Huancavelica, Huánuco, Junín, Lima, Moquegua, Pasco, Puno, Tacna).

note. Funk (1997b) included Rockhausenia apiculata as a synonym of R. pygmaea. However, there are very clear ecological preferences between the two species, R. apiculata growing in rather drier parts of the Puna where no standing water occurs, and it does not occur in peat bogs — bofedales. Rockhausenia pygmaea always occurs in peat bogs in the wetter area. Morphological differences can also clearly separate the two taxa (see Rockhausen’s key, 1939b: 280).

11. Rockhausenia lanatifolia (J.Calvo & R.I.Meneses) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300560-1

Werneria lanatifolia J.Calvo & R.I.Meneses, Phytotaxa 422 (2) 202 (2019). Type: ‘PERU. Cusco: cordillera de Vilcanota, cuenca de la laguna Sibinacocha, cerro Pumachunta, 13°50'S, 71°4'W, 4950 m, 4 April 2012, A. Palabral & al. 774 (holotype: LPB s.n.; isotype: US s.n.).’

distribution. Bolivia (La Paz), Peru (Cusco).

12. Rockhausenia cornea (S.F.Blake) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300561-1

Werneria cornea S.F.Blake, J. Wash. Acad. Sci. 18 (18): 497 (1928). Type: ‘Peru: Dry gravelly slopes, Punco, Dept. Huánaco, about 34 km. west of Huallanca, altitude about 4115 m., 1 Oct. 1922, Macbride & Featherstone 2477 (type no. 518903, Field Mus.; dupl. no. 1,121,767, U. S. Nat. Herb.).’ Holotype: F(518903 = [?material not yet barcoded]); isotype: US(1121767 = 00037301).

distribution. Peru (Ancash, Huancavelica, Junín, La Libertad, Pasco).

note. Rockhausen (1939b: 316) placed this in the synonymy of Werneria pygmaea. However, the colour of the ray limbs (at least in herbarium specimens), and other leaf characters, would preclude that. Like Beltrán (2017), I prefer to keep the two taxa separate.

13. Rockhausenia canaliculata (Sch.Bip.) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300562-1

Werneria canaliculata Sch.Bip., Bonplandia 4 (4): 52 (Schultz Bipontinus 15 Feb. 1856), p.p. Type: ‘Meine Werneria canaliculata vom Tobina hat ein sehr langes, … Dahin gehört [Lechler] Nr. 2048 und auch, in einer spätern Entwicklungsstufe sich befindend, [Lechler] Nr. 2111, welches mich wegen der verlängerten, …’. Syntype: Lechler 2048, G(00305798; 00305799 — the capsule material corresponds to R. pygmaea), K(000374200 — determined as ‘isolectotype’ by Calvo in 2019, mounted with a Jameson s.n. collection), P(02088555, 02088556). Syntype: Lechler 2111, K(000527609), P. Lectotype (selected by Calvo et al. 2020a: 21): Lechler 2048, P(02088555). Note: Lechler 2111 is referrable to Werneria staticifolia Sch.Bip. var. celmisioides Wedd., q.v. = Rockhausenia staticifolia (Sch.Bip.) D.J.N. Hind.

Werneria pumila Kunth var. γ pinifolia Wedd. (Weddell 1856: 82). Type: ‘W. canaliculata Schultz Bip. in Bonplandia, ann. 1856, p. 52, pro parte.’ Type material: Lechler 2048, G(00305798; 00305799 — the capsule material corresponds to R. pygmaea), K(000374200 — mounted with a Jameson s.n. collection), P(02088555, 02088556).

distribution. Bolivia (La Paz), Peru (Cusco, Huancavelica, Junín, Puno).

14. Rockhausenia huascarana (J.Calvo, H.Beltrán & Trinidad) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300563-1

Werneria huascarana J.Calvo, H.Beltrán & Trinidad, Willdenowia 50 (1): 9 (Calvo et al. 2020b). Type: ‘Holotype: Peru, Ancash, Carhuaz, Huascarán National Park, quebrada Ulta, near Ulta pass, 09°07'S, 77°30'W, 4870 m a.s.l., 28 Jul 1985, D. N. Smith 11308 (USM accession no. 69998; isotype: MO n.v.).’ Holotype: USM; isotype: MO.

distribution. Peru (Ancash).

15. Rockhausenia caespitosa (Wedd.) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300564-1

Werneria caespitosa Wedd., Chloris Andina 1 (3): 83 (Weddell 30 June 1856). Type: ‘Hab. Pérou!: Cordillères du département de Lima (Dombey); ...’ Holotype: Dombey 964, P(02088584 — also given the ined. name of ‘Werneria caricoides Wedd.’ on the larger Herb. Mus. Paris. label). Note: An additional label on the sheet reads ‘uñacusma. para bubas en decoction. rapace.’ probably referring to a use of the plant.

Werneria caespitosa Wedd. var. β haenkei Wedd. (Weddell 1856: 83). Syntypes: [Peru:] ‘Cordillères de Cuzco! (Wedd.[ell]). Bolivie? (Haenke).’ Syntype: Haenke, P(00869742 — annotated as ‘Werneria Haenkei’ by Schultz Bipontinus, as indicated by Weddell). Syntype: ‘in locis paludosis Perou, prov. Carabaya.’ Weddell 4753, P(02088565 — ‘Werneria minima var. caespitosa Wedd. Chloris/ in locis paludosis. Perou. prov. Carabaya. /Flores alba. Weddell n. 4753.’). Note: Although the variety was listed by Foster (1958: 222), Rockhausen (1939b: 316) suggested that this is only a Peruvian species, although it is likely that the Bolivian record was based on Troll 2130 or Mandon 103 [P(04319316)](from collections mentioned within Rockhausen’s paper); Rockhausen only recognised the species. Calvo et al. (2020a: 103) left the name as unverified, even though Funk had seen the material in P, q.v.

Werneria haenkei Sch.Bip. ex Wedd. (Weddell 1856: 83), nom. nud. pro syn.

Werneria acicularis A.Gray ex Rockh. (Rockhausen 1939b: 315), nom. nud.

distribution. Bolivia (Cochabamba, La Paz), Peru (Ancash, Cusco, Huancavelica, Huánuco, Junín, Lima, Pasco, Puno).

16. Rockhausenia pygmaea (Gillies ex Hook. & Arn.) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300565-1

Werneria pygmaea Gillies ex Hook. & Arn., J. Bot. (Hooker) 3: 348 (Hooker & Arnott 1841). Type: ‘Valle de los Ciegos, Andes of Mendoza; Dr. Gillies.’ Types: E[ex GL](00322678 — label marked as ‘Werneria pusilla n sp/ 164 Valle de los Ciegos/Andes of Mendoza/50 Don J. Gillies.’; this is mounted with E00322677 — label marked as ‘Werneria pygmea/95 n sp. Gillies/on moist banks/Alto de los Morros/Andes of Mendoza/J Gillies’, and E00322679 — is unnumbered but is labelled ‘Werneria pygmaea n sp./Gillies/on moist banks/at the foot & descent from El Alto/ de los Morros. Andes, Mendoza/J Gillies’), K(000374203 — Werneria pusilla n. sp. 164 [with the addition of ‘pygmaea HA’ between ‘164’ and ‘n. sp.’]/ Valle de los Ciegos/ Andes of Mendoza/ J. Gillies/ 50 Don’ — this is mounted with a Reed s.n. collection from ‘Baños del Toro’). Note: Freire & Ariza Espinar (2014: 220) stated that the holotype was in E, E(00322678), certainly unaware of the duplicate in K (then on loan since 1986).

Werneria minima Meyen & Walp. (Meyen & Walpers 1843: 277). Type: ‘Peruvia: in planitie circa Tacoram, alt. 14–17,000 ped. (v.s.)’. Note: Although it is highly likely that the type collection was Meyen’s and originally in B, it is not at all clear if the citation included material of the variety (q.v. var. β pygmaea Walp.).

Werneria rhizoma J.Rémy (1849: 215). Type: [Chile:] ‘Se halla en las cordilleras de Ovalle, á los Patos. [Gay s.n.]’ Holotype: P; isotype: ?K(000527751 — s.loc.).

Werneria brachypappa Sch.Bip. (Schultz Bipontinus 1856: 53), non Phil. (Philippi 1873: 501)[= Rockhausenia cochlearis (Griseb.) D.J.N.Hind]. Type: ‘In uliginosus. Cord. de Tuno. Mai 1854. Lechler 1710a’. Type material: P(02088539 — determined as holotype by Funk, Sept. 2011, a sheet with Schultz Bipontinus’s Latin notes and an annotated Lechler collection label, together with a capsule which Schultz Bipontinus clearly wrote ‘a’ after the ‘1710’; 02088531; 02088540 — of the 4 ‘specimens’ on the sheet the upper rh is material of Distichia muscoides, barcoded as P02088541).

distribution. Argentina (Catamarca, Jujuy, La Rioja, Mendoza, Nequén, Salta, San Juan, Santa Cruz, Tucumán), Bolivia (Chuquisaca, Cochabamba, La Paz, Oruro, Potosí, Tarija), Chile (Antofagasta, Arica y Parinacota, Atacama, Aysén, Coquimbo, Metropolitan District, Tarapacá, Valparaíso), Colombia (Arauca, Boyacá, Caldas, Cesar, Cundinamarca, La Guajira, Magdalena, Nariño, Norte de Santander, Riseralda, Santander, Tolima, Valle del Cauca), Ecuador (Azuay, Bolívar, Cañar, Carchi, Chimborazo, Cotopaxi, Imbabura, Napo, Pichincha, Tungurahua), Peru (Amazonas, Ancash, Arequipa, Ayacucho, Cusco, Huancavelica, Huánuco, Junín, La Libertad, Lima, Moquegua, Pasco, Puno, San Martín, Tacna), Venezuela (Mérida).

note. Funk (1997b) clearly included Rockhausenia apiculata as a synonym of R. pygmaea. However, there are very clear ecological preferences between the two species highlighed above, and they can be separated morphologically.

17. Rockhausenia microphylla (H.Beltrán & S.Leiva) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300566-1

Werneria microphylla H.Beltrán & S.Leiva, Phytotaxa 372 (4): 297 (2018: 297). Type: ‘PERU. Lima: prov. Huarochiri, distr. Chicla, Abra Anticona, 113534S, 761138W, 4877 m, 29 April, H. Beltrán, S. Castillo & M. Arakaki 7970 (holotype: USM 253391; isotypes: HAO, HUS, HUT).’ Holotype: USM(253391); isotypes: HAO, HUSA, HUT.

distribution. Bolivia (Cochabamba, La Paz), Peru (Ancash, Huancavelica, Lima, Puno).

18. Rockhausenia glaberrima (Phil.) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300567-1

Werneria glaberrima Phil., Anales Mus. Nac. Chile, Segunda Secc., Bot. 8: 40 (Philippi 1891). Type: [Chile:] ‘Ad Linsor lecta.’ Type material: Pizarro (1960: 165) cited two collections against this name SGO44588, and SGO60590. Note: several other collections may well represent type material, although none (except the LP sheet) have the specific locality: B†(F0BN015809 — as ‘Tarapacá’), K(000527750 — as ‘Tarapacá’ — two duplicates), LP(002606 — as ‘Linsor 19.II.1885’), US(00037305 — as ‘Tarapacá’; 00622820 — a capsule containing one capitulum from K).

distribution. Bolivia (La Paz, Oruro, Potosí), Chile (Antofagasta, Arica y Parinacota, Tarapacá).

Rockhausenia D.J.N.Hind subgen. Rockhausenia sect. Spathulifoliae (Rockh.) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300569-1

Werneria Kunth subgen. Werneria [as Euwerneria (Gay) Rockh.] sect. Spathulifoliae Rockh., Bot. Jahrb. Syst. 70 (2): 266 (Rockhausen 18 August 1939a). Lectotype (selected here): Werneria spathulata Wedd. = Rockhausenia spathulata (Wedd.) D.J.N Hind

19. Rockhausenia weberbaueriana (Rockh.) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300570-1

Werneria weberbaueriana Rockh., Bot. Jahrb. Syst. 70 (3): 323 (Rockhausen 6 Oct. 1939b). Type: ‘Peru: Cordillera Blanca bei Huarez [sic!], auf dürftig bewaschsenem Steinschutt, 4300—4500 m ü. M. (A. Weberbauer Nr. 2984 – Mai 1903.). Holotype: B†. Neotype (selected by Calvo et al. 2020b: 6): ‘Peru, Ancash, Huari [Asunción], Huascarán N.P., just crossing the Ulta pass, 4870 m a.s.l., 0907S, 7730W, 28 Jul 1985, D. N. Smith 11303 (US barcode US-00622845; isoneotypes: F accession no. 1960115, MO accession no. 3316165, USM accession no. 69993).’

distribution. Peru (Ancash). Note: Beltrán (2017: 48) had previously suggested Lima, although that is considered to be through misidentification of material.

20. Rockhausenia rockhauseniana (H.Beltrán, Trinidad & J.Calvo) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300571-1

Werneria rockhauseniana H.Beltrán, Trinidad & J.Calvo, Willdenowia 50 (1): 7 (in Calvo et al. 2020b). Type: ‘Holotype: Peru, Ancash, Huaylas, Huascarán N.P., pass between quebrada Los Cedros and Hatuncocha, 08°51'S, 77°45'W, 4600–4800 m a.s.l., 12 Mar 1985, D. N. Smith & R. Valencia 9950 (USM accession no. 68139; isotypes: F accession no. 1962951, LPB s.n., MO accession no. 3316189, QCNE accession no. 58168, US barcode US-00622663).’ Holotype: USM(68139); isotypes: F(1962951), LPB, MO(3316189), QCNE(58168), US(00622663).

distribution. Peru (Ancash, Lima).

21. Rockhausenia aretioides (Wedd.) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300572-1

Werneria aretioides Wedd., Chloris Andina 1 (3): 86 (Weddell 30 June 1856). Type: ‘Hab. Bolivie: sur les montagnes des lagunas de Potosi! (d’Orbigny, no 1400).’ Holotype: P(02088552); isotype: P(02088553). Note: Freire & Ariza Espinar (2014: 221) lectotypified this name based on the same material.

Werneria minima Meyen & Walp. var. β pygmaea Walp. (Walpers in Meyen & Walpers 1843: 277). Type: ‘Peruvia: in planitie circa Tacoram, alt. 14–17,000 ped. (v.s.)’. Note: Although it is highly likely that the type collection was Meyen’s and originally in B, it is not at all clear if the citation, of a single collection, was just that of the species or included material of the variety mixed with it.

distribution. Argentina (Jujuy, Salta), Bolivia (La Paz, Oruro, Potosí), Chile (Antofagasta, Arica y Parinacota, Tarapacá), Peru (Moquegua, Tacna).

22. Rockhausenia pectinata (Lingelsh.) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300573-1

Werneria ciliata Wedd. ex Sch.Bip. (Schultz Bipontinus 1866: 530), nom. nud. pro syn. (based on Mandon 99).

Werneria pectinata Lingelsh., Repert. Nov. Spec. Regni Veg. 8 (157/159): 6 (Lingelsheim 1910). Type: ‘Bolivien: Am Chacaltaya, 4800 m (Buchtien, no. 1596!).’ Original material: ?B or ?WRSL; isotype: US(1098581 = 00037343). Note: In the first part, solely authored by Pax, of a series of papers on new Bolivian taxa (see C. de Candolle 1911, 1914; Cogniaux et al. 1910; Lingelsheim et al. 1909a, 1909b; Pax 1908) Pax described many new taxa based on collections of ‘Dr. Otto Buchtien’, Miguel Bang and H. Rusby that had been made available by Dr Baenitz of the ‘botanischen Museum zu Breslau’, now known as Wroclaw University (WRSL ≡ BRSL) in Poland. The location of the original material of these taxa will depend upon where these collections were finally deposited — either in WRSL or possibly in B (now destroyed — for several families), with several widely distributed isotypes of many taxa; a large set of duplicates is in US, although label collection dates are somewhat at odds with the protologues. Lingelsheim, Pax and Winkler were all staff members in the ‘Botanischen Museum zu Breslau’ which is certainly where they described the material and would most certainly be the most appropriate herbarium to cite if lectotypes were to be selected if type material still exists there; investigations still need to be undertaken.

Werneria knocheae Perkins (1913: 230). Types: ‘Bolivien: Aguila, 17° südl. Breite, 67° westl. Länge, an der Cordillera Real, 5200 m ü. M. (Edith Knoche n. 2, 14, 21. – Im April 1909 blühend).’ Syntype: Knoche 21, B†(F0BN015813 — lacking a sheet label in the photograph). Note: There are no images of the other Knoche syntypes in F.

distribution. Bolivia (Cochabamba, La Paz, Oruro), Peru (Ancash, Arequipa, Ayacucho, Cusco, Huancavelica, Junín, Lima, Moquegua, Puno).

23. Rockhausenia castroviejoi (J.Calvo & H.Beltrán) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300574-1

Werneria castroviejoi J.Calvo & H.Beltrán, Phytotaxa 408 (2) 139 (Calvo & Beltrán 2019). Type: ‘PERU. Lima: [Huarochií], límite con dpto. Junín, entre Casapalca y Ticlio, [11°35'S 76°11'W], 4840, 1 December 1977, S. Castroviejo, M. Costa & E. Valdés-Bermejo 1112 (holotype: MA 867835).’

distribution. Peru (Ancash, Junín, La Libertad, Lima).

24. Rockhausenia cochlearis (Griseb.) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300575-1

Werneria brachypappa [as brachypappus] Phil., Anales Univ. Chile 43 (1): 501 (Philippi 1873), nom. illeg., non Sch.Bip. (Schultz Bipontinus 1856: 53)[= Rockhausenia pygmaea (Gillies ex Hook. & Arn.) D.J.N.Hind]. Type: [Chile: Antofagasta.] ‘Hallada por el señor don Guillermo Doell cerca de las salitreras de Antofagasta, en el desierto de Atacama.’ Cabrera (1948: 58) stated: ‘Salitreras de Antofagasta, leg. Doll. 1873 (Tipo: SAGO.).’ Calvo & Moreira-Muñoz (2019: 174) considered this first-step lectotypification and provided second-step lectotypification based on SGO(000006428); isolectotypes: LP(010363), SGO(000006429).

Werneria cochlearis Griseb., Symbol. Fl. Argent.: 208 (Grisebach 1879a); Abh. Königl. Ges. Wis. Göttingen 24 (1): 208 (Grisebach 1879b). Type: [Argentina:] ‘S[alta].: Nevado del Castillo.’ Types: B†(F0BN015805 — ‘L & H [= Lorentz & Hieronymus] 114’), CORD(00006520), GOET(6285 — databased as the ‘holotype’), US(00037300). Lectotype (selected by Freire & Ariza Espinar 2014: 221): ‘19/23 Mar 1873, P. G. Lorentz & G. Hieronymus 114’, CORD(000006520); isolectotype: US(280924 = 00037300). Calvo & Moreira-Muñoz (2019: 174) were of the opinion that the lectotype was selected by Cabrera (1948: 60). However, Cabrera only stated that the isotype was in CORD, and made no comment on any possible material in GOET, contrary to the suggestion by Calvo & Moriera-Muñoz (2019: 175). The material in K(000527749), whilst clearly a Lorentz collection is only localised to ‘Salta.’, and is unnumbered.

Werneria denticulata S.F.Blake, Contr. U.S. Natl. Herb. 22 (8): 651 (1924), as nom. nov. pro W. brachypappa Phil.

distribution. Argentina (Catamarca, Jujuy, La Rioja, Salta, Tucumán).

25. Rockhausenia orbignyana (Wedd.) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300576-1

Werneria orbignyana Wedd., Chloris Andina 1 (3): 85 (Weddell 30 June 1856). Type: ‘Hab. Bolivie: au sommet de la Cordillère de La Paz! (d’Orbigny, no 338 [sic! = '328' on sheet]).’ Holotype: P (00869837 - designated as 'LECTOTYPE' by Calvo in Sept. 2019, although cited as 's.n.' in Calvo et al. 2020a: 57, the material not having been returned from loan at that point).

Werneria orbignyana Wedd. var. breviradiata A.Gray (1861: 139). Type: [Peru:] ‘High Andes of Peru, near Casa Cancha. [Wilkes s.n.]’ Holotype: GH; isotypes: K(000527755), US(00037328).

Werneria nuda A.Gray, Proc. Amer. Acad. Arts 5: 139 (1861), nom. nud. (based on the type of W. orbignyana var. breviradiata).

Werneria mandoniana Sch.Bip., Bull. Soc. Bot. France 12: 80 (Schultz Bipontinus 1865), nom nud. (based on Mandon 102).

Werneria pygmaea Gillies ex Hook. & Arn. var. rhodopappa Phil., Anales Mus. Nac. Chile, Segunda Secc., Bot. 8: 41 (Philippi 1891). Type: [Chile: Antofagasta] ‘De Machuca 3200 m. s. m. [17 Feb. 1885], pappus pulcherrime roseus.’ Lectotype (selected by Calvo & Moreira-Muñoz 2019: 174): SGO(000006435); isolectotype: SGO(000006442).

Werneria mandoniana Wedd. ex Klatt (1894: 367). Type: [Bolivia:] ‘Hab.: Prov. Larecaja, viciniis Sorata, valle inter las trincheras de Chiliata et montem Hlampie, in scopulosis, Alt. Reg. alpina, 3800 m., leg. Mandon, Sept. 1858, Nr. 102.’ Type material: B†(F0BN015816), BR(5531547), G(00305793, 00305794), K(000527754), P(00711470, 04318256), S-R-6524, US(1706004 = 00037324, 1803309 = 00037325). Note: The locality referred to by Klatt is written by Mandon as ‘Valle inter las Trincheras de Chillata et montem Illampu. in scopulosis ...’. Illampu (6368 m) is the fourth highest peak in Bolivia. Whilst there is a Laguna Chillata (4235 m), NE of Illampu, there is no modern reference to La Trincheras de Chillata (or Chiliata).

Werneria orbignyana Wedd. var. longifolia Rockh., Bot. Jahrb. Syst. 70 (3): 331 (Rockhausen 1939b). Type: ‘Peru: Departm. Lima, Antaicocha, Cerro Colorado, östlich Canta, offene sandige Hänge, 4000–4100 m ü. M. (Fr. W. Pennell Nr. 14690 – Juni 1925.) Typus!’. Note: Rockhausen (1939b: 331) did not state where the type was located, nor whether he had seen any of the other duplicates. Type material: F(558651), GH(00936610), NY(04241910), PH(00348278), US(00622823). Lectotype (selected by Calvo et al. 2020a: 57): PH(00348278).

distribution. Bolivia (Cochabamba, La Paz, Oruro), Chile (Antofagasta, Arica y Parinacota, Tarapacá), Peru (Ancash, Arequipa, Ayacucho, Cajamarca, Cusco, Huancavelica, Junín, Lima, Moquegua, Puno, Tacna).

26. Rockhausenia spathulata (Wedd.) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300577-1

Werneria spathulata Wedd., Chloris Andina 1 (3): 85 (Weddell 30 June 1856). Type: ‘Hab. Bolivie: partie supérieure du ravin de Chuquiaguillo, dans les pelouses un peu tourbeuses de la Lancha!, Cordillère de La Paz (Wedd.).’ Holotype: P(02088537); isotype: P(02088538). Type material: P(02088537; 02088538 — a capsule containing 2 separate portions of flowering stem and a separate leaf). Lectotype (selected by Freire & Ariza Espinar 2014: 224): P(02088537 — det. by Funk 2011, as the holotype).

distribution. Argentina (Jujuy, Salta, Tucumán), Bolivia (La Paz, Oruro, Potosí), Chile (Antofagasta, Arica y Parinacota, Tarapacá), Ecuador (Napo), Peru (Arequipa, Huacavelica, Lima, Moquegua, Puno, Tacna).

27. Rockhausenia glandulosa (Wedd.) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300578-1

Werneria glandulosa Wedd., Chloria Andina 1 (3): 85 (Weddell 30 June 1856). Type: [Hab. Pérou: fentes des rochers, dans les Cordillères de Carabaya! (Wedd[ell].)’. Holotype: P(02088561 — the pre-printed label reads ‘AMERIQ. méridle. Pérou, prov. de Carabaya./ Juin–Juillet 1847./ M. H. Alg. Weddell, 1848./ Cat. prop. No.’, and in Weddell’s hand ‘4738?’, a single plant mounted on the sheet with 2 small leaf fragments in a capsule). Lectotype (selected by Calvo et al. 2020a: 33): P(02088561). Epitype (selected by Calvo et al. 2020a: 33): ‘Peru. Cusco Paucartambo, Parque Nacional del Manu, Acjanaco, cerro Inanbari [Inambari], 3,850 m, 21 Mar 1992, A. Cano 5241 (USM-107649!).’

distribution. Bolivia (La Paz), Peru (Cusco, Puno).

note. As this species is still relatively poorly illustrated, bar the colour images provided by Alfredo Fuentes (LPB) to Calvo et al. (2020a), which he also provided to me after my earlier study visit in 2015, I have provided a black and white line illustration to allow some of the critical details to be seen, and comparison made with similar species — see Fig. 1.

Fig. 1
figure 1

Rockhausenia glandulosa. A habit of flowering plant; B abaxial view of leaf; C detail of leaf hair; D capitulum; E l.s. capitulum, showing fistulose scape apex beneath receptacle; F phyllaries; G detail of apex of scape bract; H ray floret; J detail of style arms of ray floret; K disc floret corolla opened out showing attachment point of filaments; L side view of disc floret corolla lobe; M detail of style arms of disc floret; N stamen; P achene and base of pappus setae; Q detail of apex of pappus seta. All from Fuentes et al. 12607 (LPB). drawn by hazel wilks.

Full size image

Two new combinations in Werneria

In maintaining all xenophyllums as wernerias, new combinations are required for the following two species:

Werneria funkiana (J.Calvo) D.J.N. Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300579-1

Xenophyllum funkianum J.Calvo, PhytoKeys 139: 30 (in Calvo & Funk 2020). Type: ‘Ecuador. Chimborazo: Mt. Chimborazo area, at the end of Polylepis road and beginning of hike to Polylepis forest, 1°31'50"S, 78°52'55"W, 4233 m, 20 Apr 2018, V. A. Funk & J. M. Bonifacino 14059 (holotype: US!: isotypes: MO!, QCA!).’ Holotype: US; isotypes: MO, QCA.

distribution. Ecuador (Bolívar, Chimborazo).

Werneria lorochaqui (J.Calvo & V.A.Funk) D.J.N.Hind, comb. nov.http://www.ipni.org/urn:lsid:ipni.org:names:77300580-1

Xenophyllum lorochaqui J.Calvo & V.A.Funk, Phytokeys 139: 34 (2020). Type: ‘Argentina. Catamarca: El Cajón, Negroara, [26°24'S, 66°22'W], 15 Jan 1914, L. Catillón 3365 (holotype: LIL-26677!; isotypes: BM s.n.!, BR s.n.!, US barcode 00622893!, W-334!).’ Holotype: LIL(26677); isotypes: BM, BR, US(00622893), W. Note: The number provided by Calvo & Funk (2020: 34) for the isotype in W is too small to be either an accession or barcode number, with or without any missing zeros.

distribution. Argentina (Catamarca, Jujuy, Salta, Tucumán).