4 LUNDELLIA DECEMBER, 2017

NOMENCLATURAL NOTES ON THE ANDEAN GENERA PYCNOPHYLLOPSIS AND PYCNOPHYLLUM (CARYOPHYLLACEAE)

Martın´ E. Timana´ Departamento de Humanidades, Seccion´ Geografıa,´ and Centro de Investigacion´ en Geografıa´ Aplicada (CIGA) Pontificia Universidad Catolica´ del Peru,´ Av. Universitaria 1801, San Miguel, Lima 32. Peru´ Email: [email protected] Abstract: The nomenclature of the high Andean genera Pycnophyllopsis Skottsb. and Pycnophyllum J. Remy´ is examined. Eight species of Pycnophyllopsis are recognized; lectotypes or neotypes are selected when required; a new species, Pycnophyllopsis smithii is proposed and two new combinations are made. The genus Plettkea Mattf. is reduced to a synonym of Pycnophyllopsis. Ten species of Pycnophyllum are accepted, including a new species, Pycnophyllum huascaranum and lectotypes or neotypes are selected when needed.

Resumen: Se examina la nomenclatura de los generos´ altoandinos Pycnophyllopsis Skottsb. y Pycnophyllum J. Remy.´ Se reconocen ocho especies de Pycnophyllopsis; se designan lectotipos y neotipos cuando es requerido; se propone una nueva especie, Pycnophyllopsis smithii, y dos nuevas combinaciones. Se aceptan diez especies de Pycnophyllum, incluyendo una nueva especie, Pycnophyllum huascaranum; se designan lectotipos y neotipos cuando es requerido.

Keywords: Caryophyllaceae, Alsinoideae, Pycnophyllopsis, Pycnophyllum, Plettkea, Andes, nomenclature, Peru, Bolivia.

The Caryophyllaceae consists of 100 Andes and the mountain regions of North genera and almost 3000 species (Herna´ndez and Central America. Some genera (includ- et al., 2015). Traditionally, the family has ing several endemics) of the Caryophyllaceae been divided into three subfamilies (Alsi- reach the southern hemisphere, particularly noideae Fenzl, Caryophylloideae Arnott , the high Andes and the south temperate and and Paronychioideae Meisner, but see Har- sub-Antarctic regions. Of the currently baugh et al., 2010) based on various floral recognized genera, 22 are native in South and vegetative features. In that same scheme, America (plus another ten represented by the subfamily Alsinoideae encompasses 28 introduced species), particularly along the genera and nearly 750 species worldwide. Andean region with several of these genera Members of this subfamily are distinguished endemic to the Andes, e.g., Pycnophyllum J. from the Caryophylloideae by their free Remy,´ Pycnophyllopsis Skottsb., Reichella sepals, and from the Paronychioideae by Pax, and Philippiella Speg. their exstipulate leaves and possession of Pycnophyllopsis and Pycnophyllum are true petals (versus petaloid staminodes; among the most poorly known members of McNeill, 1962). the Caryophyllaceae in the New World. Although the family is mainly north Herbarium specimens are, to date, extremely temperate in distribution, with its center of scarce. This under-representation may be diversity in the Mediterranean and Irano- because they are naturally found in the most Turanian region (Bittrich, 1993), several remote parts of the Andes, usually near the genera of Caryophyllaceae are also distinc- snow line. It may also be due to their tive components of alpine floras around the reduced size and, thus, to their inconspicu- world. They are found in the mountain ousness. regions of the Middle East, central Asia, Here I present some long-needed no- alpine New Zealand and Australia, the menclatural notes regarding both Pycnophyl-

LUNDELLIA 20:4–24. 2017 NUMBER 20 TIMANA:´ NOMENCLATURE OF PYCNOPHYLLOPSIS AND PYCNOPHYLLUM 5

lum and Pycnophyllopsis. Because several nally proposed by Skottsberg (1916) to former students of these genera, including include all species of the genus Plettkea Pax, Mushler, and Mattfeld, were based at established by Mattfeld (1934a). the herbarium of the Botanic Garden and When Mattfeld proposed Plettkea Botanical Museum Berlin-Dahlem (B), sev- (Mattfeld, 1934a), he discussed in detail eral, if not most, of the specimens they the position of this taxon (and that of selected as types were destroyed (see Hiepko, Pycnophyllopsis s.s.) in the context of the 1987). Therefore, lectotypification, and in subfamily Alsinoideae. His analysis clearly some cases neotypification, were required. expressed what has been confirmed by later The present work is based upon my doctoral authors (e.g., McNeill, 1962; Bittrich 1993) – dissertation at The University of Texas in namely, that we could establish based on Austin (Timana´, 2005). A revision of each which characters are emphasized, not only genus with detailed descriptions, illustra- multiple classification schemes within the tions, distribution maps and exsiccate is subfamily but also, even generic boundaries. currently in progress. Various genera have been delimited based on single characters; for a detailed discussion MATERIALS AND METHODS on how this has affected large genera such as Arenaria refer to McNeill (1962). Although This study is based upon the examina- Mattfeld (1934b) recognized the close rela- tion of herbarium specimens and extensive tionship of these genera by placing them fieldwork. Specimens borrowed from 67 within a new subtribe (Plettkeinae), in fact, herbaria (see list in Acknowledgments; his separation of Pycnophyllopsis s.s. and abbreviations follow Thiers, continuously Plettkea was primarily based on the weakly- updated) were examined. In addition, nine perigynous ovary of the former versus the major European herbaria (B, BM, CGE, E, strongly perygynous state among Plettkea FI, K, MA, OXF, and P) were visited for the species. Based on the evaluation of multiple study of historically important collections. characters and their variation among indi- The Weberbauer collection deposited at viduals along with a cladistic analysis of MOL was also examined on three occasions. morphological data, Timana´ (2005) demon- Digital specimen images available on the strated that such separation cannot be Internet were also studied. Field observa- maintained and therefore, these two taxa tions and collecting were carried out in Peru, are here merged. Bolivia, Argentina, and Chile between1998 Pycnophyllopsis Skottsb., Kongl. Svenska and 2013. A set of collections is deposited at Vetenskapsakad. Handl. N. S. 56 (5): the Plant Resources Center (TEX-LL), The 216. 1916. University of Texas, Austin. Duplicate sets were left in the country of collection and TYPE: Pycnophyllopsis muscosa Skottsb. additional sets were distributed to various Plettkea Mattf., Schriften Vereins herbaria. ¼ Naturk. Unterweser, N.F. 7: 13. 1934. TYPE SPECIES, designated here: Plettkea cryptantha TAXONOMIC TREATMENT Mattf. Bisexual or dioecious, perennial, densely Pycnophyllopsis Skottsb. caespitose, hemispheric SHRUBS, 5—30 cm in diameter. STEM woody to sub-woody, 1.5—6 Pycnophyllopsis Skottsb. is composed of mm in diameter, branches 1.8—5 mm thick, eight species of cushion-forming plants herbaceous. LEAVES exstipulate, imbricate, occurring near the highest Andean peaks of rarely tetrastichous or spirally arranged, central Peru, Bolivia, central Chile, and adpressed, scarious; sheath glabrous; blade southern Argentina. The genus is here sessile, incurved or flat, lanceolate to nearly circumscribed in a wider sense than origi- triangular, complanate to cymbiform, base 6 LUNDELLIA DECEMBER, 2017

connate, margin entire, denticulate or cili- (LECTOTYPE, here designated S!); ISOLEC- ate, midvein absent, apex acute, or mucro- TOTYPES: LD!, UPS!). nate, abaxial surface shinny, glabrous, adaxial surface shinny or opaque, glabrous. This is a very peculiar species. It is the INFLORESCENCE terminal, single-flowered; fo- only species that shows a relatively long liose prophylls paired, rarely only one, pedicelate flower — up to 2.5 mm long, subtending the flower; pedicel up to 2.5 compared to all the other species with mm long. FLOWER bisexual or unisexual, pedicels less than 0.5 mm long. Pycnophyl- haplostemonous, tetracyclic, perigynous, lopsis muscosa is also the only species with a sub-periginous or rarely hypogynous, ovate stellate calyx rather than the typical oblong, to obdeltate; receptacular cup present or nearly cylindrical calyx that characterizes all poorly developed, annular, cotyliform or the other species in this genus. infundibuliform; flower disk present or 2. Pycnophyllopsis keraiopetala Mattf., absent. CALYX oblong-cylindrical or stellate. Schriften Vereins Naturk. Unterweser SEPALS 4 or 5, imbricate-decussate, valvate or N.F. 7: 22. 1934. — TYPE: BOLIVIA. La quincuncial, lanceolate to ovate, cymbiform Paz: zwischen Palca und La Paz, an or slightly so, free above the receptacular Felzen, 4800 m, Apr 1908, Pflanz 223 cup, scarious, margin either entire, dentic- (HOLOTYPE: B†). NEOTYPE: BOLIVIA. La ulate, or ciliate, straight to involute, apex Paz: Murillo: Cordillera Real, Chacal- acute, abaxial surface nitid, smooth, or taya, N of Milluni crossing, E of Lake glabrous. PETALS 4 or 5, rarely 3, free, Milluni, 4600 – 4900 m, 168220 S, 688100 alterni-sepalous, translucent, obovate to W, 25 Jan 2000, Timana´ 3804 (NEOTYPE wide obovate, very reduced, membrana- here designated: LPB!; ISONEOTYPES: BM!, ceous, base cuneate or oblong, margin K!, MCSN!, MICH!, LL!, USM!) (Fig. entire, apex deeply bifid to bilobed, gla- 1). brous, staminal disk absent or present, anular. STAMENS (3) 4 or 5, episepalous or Among members of the genus, Pycno- antisepalous, filament base strongly enlarged phyllopsis keraiopetala is the only species or filiform; anthers introrse, dorsifixed; with trimerous petals and stamens, although staminodes (in dioecious species) 5, episep- tetramerous and pentamerous forms are also alous, filament minutely enlarged or fili- found (sometimes all three forms in the form, flattened, sterile anthers dorsifixed. same plant, e.g., Timana´ 3804). In the case PISTIL with ovoid to turbinate, glabrous of trimerous androecia, the two stamens unilocular ovary, gynophore absent; styles missing are always those that would be 2 or 3; stigma terete, slightly clavate; ovules opposite the outermost sepals. This species 3, arranged on the base of a central placental is also characterized by an enlarged filament column. PISTILLODE (in dioecious species) 3- base (described as a nectary by Mattfeld, carpelate, unilocular, reduced ovary ovoid to 1934a), forming a weak staminal disk, a cilindric, glabrous, stigma terete. FRUIT an character shared with P. muscosa. utricule, enclosed in by the persistent sepals. SEED 1, light to dark brown, reniform, 3. Pycnophyllopsis cryptantha (Mattf.) Mo- laterally compressed, rarely shortly crested linari, Polish Bot. J. 61(2): 276 (2016) ” dorsally with surface nitid or opaque, Plettkea cryptantha Mattf., Schriften smooth, minute papilose to asperulate. Vereins Naturk. Unterweser, N. F. 7:15. Fig I, A - K, U. 1934. TYPE: 1. Pycnophyllopsis muscosa Skottsb., Kongl. PERU´ .Morococha,Raimondi 8392 Svenska Vetenskapsakad. Handl. N.S. (HOLOTYPE: B†). NEOTYPE:Lima, 56(5): 216. Fig. 11, Tab. 22(5). 1916. — Huarochirı´: Paso de Anticona, Ticlio, TYPE: ARGENTINA. Patagonia Andi- 118350 S, 768150 W, 4850 m, 16 Dec na, territorio Chubut, Meseta Chalıa,´ c. 1998, Timana´ & Tate 3767 (NEOTYPE 1300 m, 5 Dec 1908, Skottsberg 606 here designated: USM!; ISONEOTYPES: NUMBER 20 TIMANA:´ NOMENCLATURE OF PYCNOPHYLLOPSIS AND PYCNOPHYLLUM 7

FIG. 1. Isoneotype of Pcynophyllopsis keraiopetala. —The University of Texas Herbarium (LL). 8 LUNDELLIA DECEMBER, 2017

BM!, E!, F!, K!, LPB!, MICH!, TEX!, Presl, Reliq. haenk. 2(I): 12. 1831. ” US!). Stellaria laevis (Bartl.) Rohrbach, Lin- naea 37: 275. 1871. TYPE: PERU´ . A very distinctive species, Pycnophyllop- Quebrada de l’Obrachillo, s.d., Haenke sis cryptantha is unique in the genus because s.n. (LECTOTYPE designated here, PR!); of its tetramerous condition and two free ISOLECTOTYPES: HAL 0117903 (photo!), styles. It is also characterized by decussate GOET 000715 (photo!). sepals with the inner pair shorter and non- ciliate as opposed to the outer pair. This In his treatment of the genus Plettkea, Peruvian endemic species is found in Mattfeld (1934a) included the species de- populations near the divide between the scribed by Rohrbach in 1872 as Stellaria Departments of Lima and Junin (central laevis (” Cherleria laevis Bartling), suggest- Peru) in the area known as Ticlio (Anticona ing that this taxon may be congeneric with Pass), around 4800 m and the puna of members of Plettkea but he did not have Cuzco and Puno between 4800 to 5200 m. access to the original specimens collected by T. Haenke in Peru later used by both 4. Pycnophyllopsis macrophylla (Muschl.) Bartling and Rohrbach. The Haenke speci- Molinari, Polish Bot. J. 61(2): 276 men was finally located in the National (2016) ” Pycnophyllum macrophyllum Muschl., Bot. Jahrb. Syst. 45: 458. 1911 Museum of Prague Herbarium (PR), where ” Plettkea macrophylla (Muschl.) Mattf., most collections made by Haenke during the Schriften Vereins Naturk. Unterweser, Malaspina Expedition were sent (Stearn, N.F. 7: 18. Fig II, L- M, V. 1934. TYPE: 1973). After examining this specimen, I PERU´ . Ancash: Cordillera Blanca, 4500 can confirm that this taxon indeed belongs m, 14 May 1903, Weberbauer 2975 in Pycnophyllopsis. (LECTOTYPE designated by Molinari 6. Pycnophyllopsis tetrasticha (Mattf.) Ti- (2016): B†; ISOLECTOTYPE: MOL!). mana´, comb. nov. ” Plettkea tetrasticha Mattf., Schriften Vereins Naturk. Un- Pycnophyllopsis macrophylla differs from terweser, N.F. 7: 21. 1934. ” Pycnophyl- P. cryptantha in the number of floral parts: lum mattfeldii (Mattf.) J.F. Macbr. Publ. P. macrophylla is consistently pentamerous Field Mus. Nat. Hist., Bot. Ser. 13(2): with three free styles; P. cryptantha is a 611. 1937 (non Pycnophyllum tetrasti- tetramerous species with two free styles. In chum J. Remy,´ 1846). — TYPE: PERU´ . addition, the leaves of P. macrophylla are Ancash: Cajatambo, Schwarze Cordil- slightly narrower toward the apex, taking the lere uber¨ Ocros, auf durfig¨ bewachse- geometric form of a narrow triangle, while nem steinigem Boden, 4500 - 4800 m, those of P. cryptantha resemble an equilat- Weberbauer 2804a (B†). NEOTYPE: PE- eral triangle. RU. Ancash: Yungay Province. Huas- The isolectotype was examined and caran National Park, Llanganuco sector, annotated as such during one of my research Quebrada Demanda, west of Chacraraju visits to the MOL herbarium in 2000, as can base camp, 4750 – 4900 m, 98010 S, be attested to in the corresponding JSTOR 778360 W, 13 Apr 1985, Smith & V. Plants photograph (MOL00000509, photo!). Cautivo 10287 (NEOTYPE designated here: Unfortunately, designation of such in Timana´ (2005) does not qualify as effective publica- MO!; ISONEOTYPE F!). tion (McNeil et al., 2012) whereas Molinari’s (2016) designation, based on the study of When Mattfeld (1934a) described Plett- Timana´ (2005), does (see Grayum et al., 2016, kea tetrasticha, he did so by simply stating for further comments on this practice). that this species was in most aspects similar to Pycnophyllopsis weberbaueri (his Plettkea 5. Pycnophyllopsis laevis (Bartl.) Timana´, weberbaueri) but mainly differed in the more comb. nov. ” Cherleria laevis Bartl., robust habit, somewhat larger leaves, and NUMBER 20 TIMANA:´ NOMENCLATURE OF PYCNOPHYLLOPSIS AND PYCNOPHYLLUM 9

FIG. 2. Isotype of Pycnophyllopsis smithii. — The University of Texas Herbarium (LL). 10 LUNDELLIA DECEMBER, 2017

flowers notably longer. No duplicates of the staminodes 0; pistillode 0.8—1.0 mm long, original Berlin type material have been ovoid, ovary 0.5—0.7 3 0.05 mm. PISTILLATE located; therefore, determining the identity FLOWER perigynous, ovate, 2.2—2.5 3 1.5— of this species has been based on the various 1.7 mm; receptacle cup crateriform, 0.5 mm measurements made and the geography of long; calyx ovoid; sepals 5, imbricate, the original specimen and current collec- straight, stramineous, lanceolate, cymbi- tions. The specimen selected perfectly falls form, 1.5—1.8 3 0.7—0.8 mm, margins within the size ranges given by Mattfeld for entire, slightly involute and apex acute, flower, receptacular cup, sepals, and pistil. surface smooth; petals 5, light cream colored to pale stramineous, translucent, obovate, 7. Pycnophyllopsis smithii Timana´, sp. nov. 0.2—0.3 3 0.2 mm, apices deeply bifid, ´ — TYPE: PERU. Lima: Huarochirı,´ divided 1/2—3/4 the petal length; flower Paso de Anticona, Ticlio, 4850 m, 16 disk absent; staminodes 5, 0.25—0.30 mm Dec 1998, Timana´ & Tate 3768 (HOLO- long, filament base terete; pistil 1.5—2.0 mm TYPE: USM!; ISOTYPES: BM!, K!, F!, LPB!, long its ovary obpyriform to turbinate, 0.6— MICH!, LL!) (Fig. 2). 0.8 3 0.6 - 0.8 mm; styles 3; stigma terete. FRUITS 1.5 3 1 mm. SEED 1, brown, reniform, Perennial, caespitose SHRUB, (5) 8—20 laterally flattened, 0.8 3 0.8 mm, shiny, cm in diameter, dioecious. ROOTS woody, minutely papillose. robust. STEM woody, 3—5 mm in diameter, Phenology — Collected in flower from primary branches 2 mm in diameter, December to May and in fruit from August yellowish green to stramineous. LEAVES to January. imbricate, (2.0) 2.5—3.5 (3.8) mm long, Distribution — Primarily found in the sheath stramineous, 0.8—1.0 mm long, divide between Lima and Junin, central smooth; blade incurved, stramineous to Peru, in the area locally known as the slightly yellowish-green, lanceolate to nearly Anticona Pass, from 4300 to almost 4900 triangular, cymbiform, (1.7) 2.0—2.5 3 m, where it is sympatric with Pycnophyllopsis 0.8—1.2 mm wide, margin minute ciliate cryptantha and members of the genus to denticulate, cilia sparsely distributed Pycnophyllum. Pycnophyllopsis smithii has along the blade margin, single-celled, 0.05 also been reported in the Department of mm long, venation a weak midvein on Ancash, within the boundaries of Huascaran abaxial side, blade apex mucronate, abaxial National Park, at nearly 4800 m. surface nitid, smooth, adaxial surface shiny. Eponymy: It is a great honour to name INFLORESCENCES terminal, included; prophylls this new species after the American botanist paired, 1.2—1.5 mm long; pedicel 0.1—0.3 David N. Smith (1945–1991). David spent mm long. PLANTS dioecious. STAMINATE nearly four years botanizing the highest FLOWER perigynous, ovate, 2—3.0 3 1.0— peaks of central Peru, particularly the 1.3 mm with a receptacle cup present, Cordillera Blanca (Ancash) as part of his crateriform, 0.7—0.9 mm long; calyx ovoid; doctoral dissertation on the flora and sepals 5, imbricate, slightly incurved apically, vegetation of Peru’s Huascaran National stramineous, lanceolate to narrowly ovate, Park (Smith, 1988). His numerous collec- slightly cymbiform, 1.2—1.5 3 0.5 (-0.7) tions of Pycnophyllopsis and Pycnophyllum mm, margin entire, slightly involute distally have greatly enhanced our understanding of with apex acute, surface smooth, with a fine these two genera, having an impact parallel midvein visible adaxially; petals 5, light to that of A. Weberbauer a century earlier. cream-colored to pale stramineous, translu- Additional Specimens examined — cent, obovate, 0.3—0.4 3 0.2—0.3 mm, its PERU´ . Ancash: Recuay, Huascaran National apex bifid, divided 1/2 —2/3 the petal Park, quebrada Quenua Ragra, 4700 - 4850 length; floral disc absent; stamens 5, epi- m, 9858 S, 77813 W, 11 Mar 1985, Smith et sepalous, 1.0—1.2 mm long, filament base al. 11772 (MO). Junin: Ticlio, 4740 m, Dec mostly filiform, rarely slightly enlarged; 1986, Rivas et al. s.n. (USM); Capillacocha, NUMBER 20 TIMANA:´ NOMENCLATURE OF PYCNOPHYLLOPSIS AND PYCNOPHYLLUM 11

20 km E Carhuamayo, 4300 – 4350 m, 25 of a tightly arranged, very compact cushion Nov 1951, Tovar 381 (TEX). Lima: where the most visible part is the light green- Huarochirı,´ Anticona, 4843 m, 8 Aug 1987, colored leaf tips. In P. weberbaueri the leaves Chanco et al. 1219 (USM); Anticona Pass, are also imbricate, yet they do not overlap to approx. 140 km E of Lima on road to La the degree previously described, giving the Oroya, ca 4890 m, 16 Dec 1978, Dillon & overall impression of a more loose cushion, Turner 1301 (MO, TEX); Ticlio, 4800 m, 6 with various brown-colored branches clearly May 2001, Leon´ s.n. (TEX, 2 sheets); Ticlio, visible. Finally, one needs to consider the 4850 m, 14 Feb 1954, Rauh P86 (F); Canta, geographic distribution. The two taxa, both Huascoy, 4600 m, 27 Sep 1974, Waechter dioecious, and with inconspicuous flowers 217PW (GOET). Pasco:Huayllay,Lago and probably very limited dispersal mecha- Huaron,´ 4750 m, 14 Jan 1971, Ellenberg nisms, are geographically separated by a 4061 (US). distance of nearly 1000 km by two of the Pycnophyllopsis smithii is perhaps the highest mountain ranges in this part of the most difficult species to distinguish using Andes. Placing them within the same species just a single set of characters. Morphologi- would suggest the possibility of potential cally, it most resembles the disjunct Pycno- gene flow between the two populations. phyllopsis weberbaueri with which it Given the evidence at hand, this seems to be maintains differences that, although gradual, very unlikely. With these considerations in are consistent. The two differ in leaf margin: mind I have therefore decided to circum- P. weberbaueri clearly has noticeable cilia scribe this new species. (up to 0.2 mm) whereas those of P. smithii 8. Pycnophyllopsis weberbaueri (Muschl.) can be better characterized as denticulate, Timana´, Monogr. Syst. Bot. Missouri with projections less than 0.05 mm that can Bot. Gard. 127: 1267. 2014. ” Pycno- be observed at 40X magnification. In phyllum weberbaueri Muschl., Bot. addition, leaves of P. smithii are normally Jahrb. Syst. 45: 455. 1911. ” Plettkea shortly mucronulate, while in P. weberbaueri weberbaueri (Muschl.) Mattf., Schriften they are consistently acute and without any Vereins Naturk. Unterweser, N.F. 7:19. further projection. Although flowers of both Fig III, O-Q. 1934. TYPE: PERU.´ species are approximately the same length, Arequipa:Caylloma,Vincocaya, the pedicel of P. smithii is relatively shorter 4300—4400 m, 24 Aug 1902, Weberba- (0.1 – 0.3 mm) than those of P. weberbaueri uer 1373 (HOLOTYPE: B†; LECTOTYPE (0.3 – 0.5 mm). Petals of P. smithii are also (designated here): MOL!). shorter (0.2 – 0.4 mm) and more deeply cleft apically (over ½the total petal length) Although it is morphologically rather compared to those of P. weberbaueri (0.4 – uniform, this dioecious species has flowers 0.8 mm, petals cleft ½—¼the petal that are consistently pentamerous with three length). The stamens of P. smithii are up to free styles. It shares several similarities with 1.2 mm in length, while those of P. Pynophyllopsis smithii, from central Peru [see weberbaueri can be as long as 2 mm and the discussion under P. smithii for various the staminodes of the former species are also distinguishing features that separate these smaller. The style of P. smithii is somewhat two species]. Pycnophyllopsis weberbaueri is longer relative to the total length of the pistil found sympatrically with P. keraiopetala,a (mean 60% of the total pistil length) ¼ bisexual species with a weakly perigynous compared to the corresponding character in flower and a reduced number of stamens. P. weberbaueri (mean 48%). There are also differences in the overall¼ aspect of these Pycnophyllum J. Remy´ species: although they both have imbricate leaves, each opposite pair in P. smithii is Pycnophyllum, a genus of ten species, is greatly overtopped by the corresponding one of the few known genera in the lower (inferior) pair, giving the appearance Caryophyllaceae restricted exclusively to 12 LUNDELLIA DECEMBER, 2017

the high central Andes. The genus is here CALYX ovoid, cylindrical to obovoid. SEPALS circumscribed as in Mattfeld’s (1922) orig- 5, very rarely 6, imbricate quincuncial, wide inal concept, namely excluding members of to narrowly ovate, cymbiform to slightly so, Pycnophyllopsis Skottsb. and Plettkea Mattf. free, scarious, with a membranous margin or which were previously merged into Pycno- not, central callous present or absent, phyllum by MacBride (1937). margin entire, usually involute, apex acute to obtuse, abaxial side nitid, smooth, Pycnophyllum J. Remy´ in Ann. Sci. Nat. Bot., glabrous, colored or not. PETALS 0 to 5, free, Ser.´ 3, 6: 355 (Dec. 1846), non Pycno- alternisepalous, translucent or cream-col- phyllum J. Tuzson, 1911 (” Pychnophyl- ored to light brown to violaceous, oblong lum A. T. Brongniart in Orbigny, Dict. to obovate, membranous, base cuneate, Universel Hist. Nat. 13: 114. Jul-Dec margin entire, apex dentate to divide, or 1849). truncate, glabrous. STAMENS 3 to 5, anti- sepalous, heterandrous, rarely homandrous, TYPE SPECIES, designated here: Pycnophyl- filament base filiform, terete or complanate, lum molle J. Remy´ sometimes weakly subulate; anthers introrse, Stichophyllum Phil. in Fl. atacam. 19, dorsifixed; staminal disk present, flattened ¼ Tab. 1d (1860). TYPE SPECIES: Stichophyllum to prominent; staminodes (in pistillate bryoides Phil. (” Pycnophylum bryoides flowers) 2 to 5, antisepalous, filaments terete (Phil.) Rohrb.) or complanate, sterile anthers dorsifixed. Dioecious, perennial, densely caespitose, PISTIL 1.5—4.5 mm long; ovary 3-carpellate, hemispheric to irregularly shaped SHRUBS, unilocular, ovoid to ellipsoid, frequently 20—ca 200 cm in diameter. STEM woody, trigonous, glabrous, sometimes minute stip- 5—10 mm in diameter; branches herba- itate; style single, stigma terete, shortly ceous, crowded or flabelliform, proximal capitate at the apex with 3–6 ovules arranged branches sometimes enclosed by shriveled at the base of a central placental column; leaves. LEAVES exstipulate, imbricate, tetra- pistillode (in staminate flowers) 3-carpellate, stichous or spirally arranged, adpressed or unilocular, 1—2 mm long, reduced ovary spreading from the branch axis, scarious to ovoid to cylindrical, sometimes apically membranous, sheath 0.5—2.5 mm long, compressed, 0.4—1.5 mm long, glabrous, scarious to membranous, smooth, glabrous; stigma terete. FRUIT an utricle or septicidal blade sessile, incurved, straight, or recurvate, capsule opening by 3 valves, or an irregular lanceolate to wide ovate, complanate to circumscissile capsule opening near the base cymbiform, scarious, base connate, margin of the ovary; ovoid or ellipsoid, usually entire, sometimes expanded into a translu- trigonous, 1.0—3.5 mm long, 1—2 mm cent membrane, then the blade composed of wide, enclosed by persistent sepals. SEEDS 1 a scarious callous and a peripheral mem- to 6, light to dark brown, reniform, laterally brane, apex obtuse, acute or long aristate, compressed, surface nitid or opaque, midvein obscure or weakly developed on the smooth or minute papilose. adaxial side, abaxial side nitid or opaque, Despite its overall morphological simi- smooth, glabrous. INFLORESCENCE terminal, larities with members of Pycnophyllopsis and solitary, weakly exserted above the leaves, Arenaria L., species of Pycnophyllum form a single flowered; bracts 4—10 (12), opposite, single, coherent group united by a combi- decussate, complanate to slightly cymbi- nation of various morphological characters form, ovate to spathulate, membranous, not found among other members of Car- base obtuse, margin entire, apex obtuse to yophyllaceae. In addition to their cushion- rounded, rarely acute, medial callous present like life form, species of Pycnophyllum are or absent, smooth, glabrous; pedicel 0—0.4 characterized by dioecy, simple, opposite mm long. FLOWERS unisexual, haplostemo- leaves generally fused at the based forming a nous, tetracyclic, hypogynous, ovate, ob- distinctive sheath, a complete lack of any ovate, or oblong; receptacular cup absent. form of pubescence or marginal cilia, NUMBER 20 TIMANA:´ NOMENCLATURE OF PYCNOPHYLLOPSIS AND PYCNOPHYLLUM 13

FIG. 3. Lectotype (P 00156958) of Pycnophyllum molle. — Reproduced by kind permission, Ó MNHN collection-Paris. 14 LUNDELLIA DECEMBER, 2017

hypogynous flowers, 5 free sepals, 3 to 5 Given its wide geographic distribution antisepalous stamens (or staminodes) form- across various degrees of latitude, Pycno- ing a well-defined staminal disk, a pistil phyllum molle shows a considerable degree terminating in a single style with 1—3 of variation. Although the main source of stigmatic lobes, and a one to three-seeded variation is in leaf size, shape, and color, the fruit. Fruits can be indehiscent (P. sect. floral structure remains rather constant. Gymnopycnophyllum), or dehiscent (3- Much of the apparent variation among P. valved capsule) as in P. sect. Pycnophyllum. molle specimens is caused by the different branching patterns detected within the same Pycnophyllum sect. Pycnophyllum cushion – a condition produced by either intrinsic causes, such as branch age, or ” Pycnophyllum sect. Kirrhanthemum extrinsic ones, such as micro-topography. Mattf. in Feddes Repert. Spec. Nov. Regni Consequently, one often finds a wide range Veg. 18: 174 (1922); nom. superfl. of variation within a single individual. The same effect has been noticed in P. tetra- 1. Pycnophyllum molle J. Remy,´ Ann. Sci. stichum. Nat. Bot., Ser.´ 3, 6: 355, Tab. 20, fig. 2-8 (Dec. 1846) —TYPE: BOLIVIA. Potosi: 2. Pycnophyllum huascaranum Timana´, sp. Lagunas de Potosi, March, D’Orbigny nov. — TYPE: PERU´ . Ancash: Carhuaz, 442.(LECTOTYPE,designatedhere:P! Huascaran National Park, lateral valley (P00156958); ISOLECTOTYPES:P of Quebrada Ishinca, trail to Lago (P00156959, photo!), F! (fragment; Ishinca, 4730—4930 m, 98230 S, 778250 871384), G! (G00226948) (Fig. 3). W, 12 Feb 1985, Smith, Valencia & Gonzales 9453;(HOLOTYPE:MO! Pycnophyllum aculeatum Muschl., Bot. 3311207); ISOTYPES:AAU!,F!MO! Jahrb.¼ Syst. 45: 456, Fig. 1. (1911). TYPE: (3311209) (Fig. 4). PERU. Arequipa: Vincocaya, en el ferroca- rril de Arequipa a Puno, 4300—4400 m, s.d. Pycnophyllum huascaranum differs from (24 Aug 1902, per Mattfeld), Weberbauer Pycnophyllum molle in the dark orange to 1374 (HOLOTYPE: B†; LECTOTYPE, designated orange-yellow blade callous, the strongly here: MOL!) involute blade margin, 6(8) bracts, and the Pycnophyllum markgrafianum Mattf., 3-valved capsule. ¼ Feddes Repert. Spec. Nov. Regni Veg. 18: Hemispherical, convex SHRUB, up to 60 175 (1922). TYPE: PERU. Ancash: Cajatam- cm in diameter and 25 cm high; stem bo, 4400 m, 6 Apr 1903, Weberbauer 2783 woody, 6—9 mm in diameter; branches (HOLOTYPE: B†; LECTOTYPE, designated here: compact, crowded, primary branches terete K!; ISOLECTOTYPES: G!, MOL!) to clavate, 2.5—3.5 mm wide, light brown to The French traveller, palaeontologist yellowish green. LEAVES imbricate, appressed and explorer Alcide d’Orbigny (1802-1857) to the branch axis, 2.3—3.1 mm long; sheath visited Bolivia from 1830 to 1833; his plant translucent, 0.5—0.8 mm long; blade in- collections were studied by Jules Ezechiel curved, light stramineous, ovate to obovate, Remy´ (1826-1893) and published in the slightly cymbiform, 1.3—2.1 3 1.0—1.4 Annales des Sciences Naturalles in two mm, margin membranous, translucent, articles, Analecta Boliviana I and Analecta strongly involute, blade venation obscure, Boliviana II (Remy,´ 1846, 1847; Stafleu and apex obtuse to rounded, apiculate, less than Cowan 1981, 1983). There are two d’Or- 0.1 mm long, blade surface shiny, smooth, bigny’s specimens at the Paris herbarium blade callous present, obspathulate, 1/2 — 2/ belonging to this species, but only the one 3 the total blade length, dark orange to selected as lectotype carries the label which orange yellow. INFLORESCENCE single, termi- coincides with the locality described by nal, bracts 6(8), obovate to obspathulate, Remy´ in the original description. 3.0—3.5 3 1.3—1.6 mm, bract apex obtuse NUMBER 20 TIMANA:´ NOMENCLATURE OF PYCNOPHYLLOPSIS AND PYCNOPHYLLUM 15

FIG. 4. Holotype (MO 3311207) of Pycnophyllum huascaranum. — Reproduced by kind permission,Ó Mo. Bot. Garden. (MO). 16 LUNDELLIA DECEMBER, 2017

to rounded, callous bract absent, pedicel 0— or crystalline rocks, some glacial debris, 0.2 mm long. STAMINATE FLOWER obovate, some large outcrops; on soil as a yellow obconical to ovoid; sepals slightly incurved, mound, level site, 4250 m, 22 Jul 1975, from light cream to pale yellow, peripherally Bunin s.n. (COLO); Huarapasca, estacion 40 membranous, scarious medially, obovate, miles east of Huaraz, 5500 m, 3 Oct 1922, carinate to slightly cymbiform, 3.8—4.1 3 MacBride & Featherstone 2488 (BM, F, G, 1.5—2.5 mm, margin slightly involute, apex MA, NY, S, US); Cordillera Huayhuasch, obtuse to rounded, abaxial surface smooth, 4600 m, 19 Jul 1954, Rauh & Hirsch P1894 veins obscure, callous present, 2/3—3/4 the (F); Cordillera Raura, 4900 m, 21 Jul 1954, sepal length, light yellow; petals 5, translu- Rauh & Hirsch P1902 (F); south of Huara´z, cent, obovate, 1.1—1.4 3 0.1—0.5 mm, apex 32 km from Pachacoto towards La Union, obtuse to truncate; stamens 5, heterandrous, Punta Huarapasca, 4750 m, 6 Apr 1988, 1.4—2.3 mm long, filament base filiform, Renvoize & Laegaard 5132 (K); Huascaran floral disk poorly developed; pistillode 1.3— National Park, between pass and Carpa, rio 1.7 mm long, pistillode ovary ovoid, 0.6— Pachacoto drainaje, 4600–4350 m, 98530 S, 0 0.7 3 0.4—0.7 mm. PISTILLATE FLOWER 77814 W, 31 Mar 1985, Smith & Escalon obovate, obconical to ovoid; sepals slightly 10216 (F, MO); Huascaran National Park, incurved, from light cream to pale yellow, Llanganuco sector, quebrada Demanda, peripherally membranous, scarious medial- between Lago 69 and glacier coming from ly, obovate, carinate to slightly cymbiform, Nevado Pisco, 4800 m, 98000 S, 778360 W, 12 3.4—3.8 3 2—2.3 mm, margin slightly Apr 1985, Smith & Cautivo 10273 (AAU); involute, apex obtuse to rounded, abaxial Huascaran National Park, quebrada Quenua surface smooth, veins obscure, callous pre- Ragra, 4700-4600 m, 98580 S, 778130 W, 10 sent, 2/3—3/4 the sepal length, light yellow; May 1985, Smith et al. 10654 (MO); petals 5, translucent, obovate, 0.8—1.3 x Huascaran National Park, 3-4 km below nearly 0.2 mm, apex obtuse to truncate; Cahuish Tunnel, 4500-4550 m, 98410 S, staminodes 5, homandrous, less than 0.2 778140 W, 10 Jul 1985, Smith & Buddensiek mm long, filament flattened, floral disk 11115 (F, MO); Huascaran National Park, minute; pistil 1.8—2.5 mm long, ovary Paron´ Valley, E of Lake, 4600-4900 m, 88580 ovoid, 1.2—1.5 3 0.9—1.2 mm, ovules 3, S, 778380 W, 28 Sep 1985, Smith 11506 (F, stigma trifid. FRUIT a 3-valved capsule, 1.5— MO); Huascaran National Park, lateral 1.8 3 1.2—1.5 mm, ovoid. SEEDS 2—3, light valley of quebrada Ishinca, 4730-4930 m, brown, asymmetrically reniform, ca 1 x ca 98230 S, 778250 W, 12 Feb 1985, Smith et al., 0.7 mm, minute papillose. 9453 (AAU, F, MO); Huascaran National Phenology — Flowering from October to Park, quebrada Alpamayo, at foot of Alpa- April and fruiting from February to May. mayo and Quitaraju, 4600-4750 m, 88530 S, Distribution — Endemic to the Depart- 778410 W, 8 Mar 1985, Smith et al. 9738 ment of Ancash, Peru, in the vicinity of (AAU, F, MO); Huascaran National Park, Huascaran National Park, from 4200 to 5000 pass between quebrada Los Cedros and m. Hatuncocha, 4600-4850 m, 88510 S, 778450 Eponymy — This new species is named W, 12 Mar 1985, Smith & Valencia 9949 after the type locality, Huascaran National (AAU, MO); Pisco Creek, Llanganuco Val- Park, a magnificent Peruvian National Park ley, S. exposure, ca 3200 m, Aug 1959, where a number of interesting caryophylla- Tothill 147 (CORD, F, UC). ceous species are found. This taxon is characterized by its robust, Additional Specimens examined — PE- densely packed primary branches, its mark- RU. Ancash: Llanganuco valley, slopes of edly involute leaf margin, particularly to- Huandoy, Cordillera Blanca, 18 Jul 1968, ward the apex, the conspicuously bright Balthgate 10 (E, F); near lago Safuna, orange to light brown, shiny, protruding northern Cordillera Blanca, northeast of callous and dehiscent fruit. The callous can Alpamayo, 4250 m, rocky sites with granitic sometimes be so conspicuous that, to an NUMBER 20 TIMANA:´ NOMENCLATURE OF PYCNOPHYLLOPSIS AND PYCNOPHYLLUM 17

untrained eye it may resemble the sporangia Pycnophyllum have been described as inde- of some Sellaginella species (some Pycno- hiscent or irregularly rupturing (Mattfeld, phyllum specimens have been determined as 1922; Bittrich, 1993) P. macropetalum clearly such in the past). The fruits are also peculiar produces a 3-valved capsule. because they can unambiguously be consid- ered capsules, opening from the bottom of 5. Pycnophyllum holleanum Mattf., Feddes the ovary into three valves. Repert. Spec. Nov. Regni Veg. 18: 177. 1922. TYPE: PERU. Junin: Huancayo, 3. Pycnophyllum spathulatum Mattf., am Gletscher Chuspicocha uber¨ der Feddes Repert. Spec. Nov. Regni Veg. Hacienda Acopalca, nordostlich¨ von 18: 175 (1922). TYPE: BOLIVIA. Zwi- Huancayo, geogr. Br. ca. 12850 S, 4900- schen Tomarape und Sajama, 4500 m, 5000 m, 27 Mar 1913, Weberbauer 6522 Oct 1876, Stubel¨ 3 (HOLOTYPE, B†). (HOLOTYPE: B†; LECTOTYPE here designat- NEOTYPE: BOLIVIA. al este del pueblo ed: K!; ISOLECTOTYPES: B!, BM!, CAS!, de Sajama. Bosque abierto de Polylepis DS!, F! (2 sheets), MA!, MO!, MOL!, tarapacana, sobre las laderas del cerro NY!, PH!, S!, UC!, US! Sajama, 4750 m, 1 Jun 1991, Beck 19899 (NEOTYPE designated here: LPB!; ISONEO- Pycnophyllum holleanum is one of the TYPE: TEX!). most distinctive and handsome species in this genus. It is characterized by squarrose As with many others of Stubel’s¨ speci- leaves, a sharp awn, and the light brown mens deposited in Berlin, the type of color of the foliar callous. Morphologically it Pycnophyllum spathulatum is no longer is most similar to P. aristatum from which it extant. The only fertile specimen (with differs in the shorter awn and the blade pistilate flowers) gathered in the same shape (ovate in the latter). Mattfeld de- locality as the type is that of S.G. Beck, scribed this species based on a subsequently which is therefore selected here as the destroyed Berlin specimen. Of the 15 neotype. Morphologically Pycnophyllum spa- isotypes studied, only two (K and MOL) thulatum is most similar to P. macropetalum, have the same complete collection informa- from which it can be easily distinguished by tion as that of the protologue. The Kew its upright, uniformly terete, brownish- specimen label reads ‘‘Ex Museo Botanico yellow branches, whereas P. macropetalum Berolinensi’’ and shows the accession date as has lax, light green to yellow, club-shaped ‘‘24 Apr. 1923’’ (one year after Mattfeld’s branches. monograph). The Berlin isolectotype cited above was acquired from the Stockholm 4. Pycnophyllum macropetalum Mattf., herbarium in June 1961, thus it presumably Feddes Repert. Spec. Nov. Regni Veg. cannot be considered part of the original ´ 18: 176 (1922) — TYPE: PERU. [Tacna] material studied by Mattfeld. Cerro Tacora, 4500 m, 7 Oct 1876, Stubel¨ 110. (HOLOTYPE: B†; LECTOTYPE, 6. Pycnophyllum aristatum Mattf., Feddes designated here: K!). Repert. Spec. Nov. Regni Veg. 18: 178 (1922). TYPE: PERU. Ancash: Pallasca, Pycnophyllum macropetalum exhibits 4600—4800 m, Aug 1918, Weberbauer numerous unique characteristics that make 7250 (HOLOTYPE: B†; LECTOTYPE here it easily recognizable: it not only has the designated: K!; ISOLECTOYPE: F!). largest petals in the genus but also it is the only species in which petals are longer than Pycnophyllum aristatum is another dis- the sepals; in addition, P. macropetalum is tinctive species in this genus. It is easily the only species with coloured sepals – from recognized by the long, filamentous, spiraled light brown to a rather intense violaceous awns, the flabelliform disposition of branch- color, particularly toward the apical region es, and the light brown coloration of the of the sepals. Although fruits in the genus young branches. 18 LUNDELLIA DECEMBER, 2017

Pycnophyllum sect. Gymnopycnophyllum Perez,´ 26-28 Jan 1879, Hieronymus & Pax, Bot. Jahrb. 18: 34 (1893). Niederlein 365 (HOLOTYPE: B†; LECTOTYPE designated here: G!; ISOLECTOTYPES: S!, E!, TYPE SPECIES: Pycnophyllum tetrastichum UC!). ´ Pycnophyllum mucronulatum Mattf., J. Remy. ¼ See discussion under Pycnophyllum tet- Feddes Repert. Spec. Nov. Regni Veg. 18: rastichum 173 (1922) TYPE: ARGENTINA. Salta: Pycnophyllum sect. Diphtheranthe- Nevado del Castillo, 19-23 Mar 1873, mum¼ Mattf., Feddes Repert. Spec. Nov. Hieronymys & Lorentz 31 (HOLOTYPE: B†; Regni Veg. 18: 170 (1922). TYPE SPECIES: LECTOTYPE designated here: GOET!; ISOLECTO- Pycnophyllum convexum Griseb. TYPES: F! [fragment]: G-DC!) Pycnophyllum sect. Drudea (Griseb.) In the protologue of P. argentinum, Pax Pax,¼ Bot. Jahrb. 18: 34 (1893). TYPE (1893) included a second specimen, Stubel¨ 4, SPECIES: Pycnophyllum lechlerianum Rohrb. then at the Berlin herbarium (B) but now destroyed. Mattfeld (1922) however pointed 7. Pycnophyllum bryoides (Phil.) Rohrb., out it belonged to P. tetrastichum. Pycno- Linnaea 36: 662 (Dec 1870) ” Sticho- phyllum convexum as circumscribed here phyllum bryoides Phil., Fl. atacam. 19. includes two other described Argentinean Tab. Id. (1860) — TYPE: CHILE. taxa, namely, P. argentinum Pax and P. Atacama: Alto de Puquios in deserto mucronulatum Mattf. Although the study of Atacama, Feb. 1854, R. Philippi s.n. type material of these taxa shows some (HOLOTYPE: SGO 48876! p.p., marked as differences between them, the analysis of No. 2). multiple populations across their geographic range demonstrates that these three taxa Pycnophyllum bryoides is one of the cannot be kept separate. As discussed under most distinctive and well-defined species in P. tetrastichum, the levels of phenotypic P. sect. Gymnopycnophyllum. It can easily be variation even within individual plants show recognized by its uniformly spirally arranged that the species limits set by Mattfeld are not leaves, and by the presence of five small, meaningful. apically divided petals, a unique condition among members of this section. While 9. Pycnophyllum tetrastichum J. Remy,´ various specimens collected by Philippi from Ann. Sci. Nat. Bot., Ser. 3, 6: 356, Tab. the Atacama Desert region have been 20, fig. 1 (Dec. 1846). TYPE: BOLIVIA. located, only the SGO holotype carries the Potosi: 4000 m, D’Orbigny s.n. (LECTO- exact information provided in the proto- TYPE selected here: P!, P00156960; ISO- logue. Except for the HAL specimen that was LECTOTYPE: P!, P04925579) (Fig. 5). collected in 1854, all of the others were Pycnophyllum lechlerianum Rohrb., collected later or lack enough information to ¼ decide their nomenclatural status. Linnaea 36: 664 (Dec 1870); Drudea lycopo- dioides Griseb., Symb. Fl. Argent. 25 (1879), 8. Pycnophyllum convexum Griseb., Pl. pro parte. TYPE: PERU, Azangaro, ‘‘in lorentz. 28 (1874); also in: Abh. Konigl.¨ montibus asperis lapidosis,‘‘ Jun, Lechler Ges. Wiss. Gottingen¨ 19: 76 (1874) — 1742 (HOLOTYPE: B†; LECTOTYPE designated TYPE: ARGENTINA. Catamarca: Vayas here: G!; ISOLECTOTYPES:BR!,CGE!,F!, altas pr. Belen alt. 9-11000 ft, Jan 1872, GOET!, K!, L!, M!, P!, SGO!, S!, TCD!). Lorentz 638 (HOLOTYPE: GOET!; ISOTYPE Pycnophyllum filiforme Mattf., Feddes CORD!) Repert.¼ Spec. Nov. Regni Veg. 18: 172 (1922). TYPE: PERU. Arequipa: Caylloma, Pycnophyllum argentinum Pax, Bot. 158200-158300 S, am See Villafro uber¨ Jahrb.¼ Syst. 18: 33 (1893). TYPE: ARGEN- Caylloma, 4700—4800 m, 29 Mar 1914, TINA. La Rioja: Sierra Famatina, Cueva de Weberbauer 6885 (HOLOTYPE: B†; LECTOTYPE NUMBER 20 TIMANA:´ NOMENCLATURE OF PYCNOPHYLLOPSIS AND PYCNOPHYLLUM 19

FIG. 5. Lectotype (P 00156960) of Pycnophyllum tetrastichum. — Reproduced by kind permission, Ó MNHN collection-Paris. designated here: MOL!; ISOLECTOTYPES: G!, designated here: NY!; ISOLECTOTYPES: GH!, US!). US!). Pycnophyllum glomeratum Mattf., When Remy´ proposed the genus Pycno- Feddes¼ Repert. Spec. Nov. Regni Veg. 18: phyllum in 1846, he included two species: P. 171 (1922). TYPE: PERU. Puno: Sandia, molle and P. tetrastichum. The description of zwischen Poto und Ananca, offene durftig¨ the latter makes reference to a single bewachsene Polsterpflanzenmatte, 4600- specimen that included both species (‘‘mixta 4700, 7 May 1902, Weberbauer 952 (HOLO- in herbario parisiensi cum praecendete’’). TYPE: B†; LECTOTYPE designated here: K!; However, currently, the D’Orbigny 442 ISOLECTOTYPE MOL!). specimen at P (P00156958, Fig. 3) only Pycnophyllum leptothamnum Mattf., consists of material corresponding to P. Notizbl.¼ Bot. Gart. Berlin-Dahlem. 10: 1051 molle. A second sheet at P (P00156960, Fig. (1930). TYPE: PERU. Cuzco: Quispicanchis, 5) containing P. tetrastichum, is attributed to in der Puna-Matte nahe den Gletschern des D’Orbigny but without collection number, Auzangate, 4500-4600 m, 11 Feb 1929, A. and includes the annotation ‘‘etais avec la Weberbauer 7772 (HOLOTYPE B†; LECTOTYPE congen´ ere` ’’ – which can be translated as ‘‘it 20 LUNDELLIA DECEMBER, 2017

was with its congener’’, namely Pycnophyl- see any set of correlated characters (and lum molle.Thusthisspecimenishere particularly discrete characters) to recognize designated as the lectotype. and support more than one species. While In his 1922 revision of the genus some regional variation may be suggested, Pycnophyllum and subsequent 1930 publi- additional detailed field studies and sam- cation, Mattfeld described several new pling is needed to finally justify species species in P.sect.Gymnopycnophyllum, recognition. some of which are here reduced to August H. R. Grisebach (1879) placed synonyms of Pycnophyllum tetrastichum. Rohrbach’s Pycnophyllum lechlerianum and Such a merger requires justification. First, his own Colobanthus lycopodioides into an an analysis of Mattfeld’s 1922 key to species unusual new genus, Drudea,combining shows that, for the most part, the differ- these two species as Drudea lycopodioides. ences supporting his species are primarily This arrangement was based on two speci- based on the shape (outline) and the size of mens: Lechler 1078, the type of Colobanthus flowers and/or leaves. This leads to a rather lycopodioides Griseb. (1854) and Lechler artificial separation since these two charac- 1742, the type of Pycnophyllum lechlerianum ters are each part of a continuum. Matt- Rohrb. (1870). Pax (1893) later reduced feld’s new species were based only upon one Grisebach’s anomalous species to a synonym or two specimens (some of them fragmen- of P. lechlerianum Rohrb. (keeping C. tary), and while the differences he suggested lycopodioides as a synonym as well) and may well apply to such a small sample, a proposed Pycnophyllum sect. Drudea to thorough examination of additional speci- include it. Finally, Skottsberg (1916) emend- mens revealed less support for such divi- ed Pax’s section by removing Colobanthus sion. Second, plants of this taxon show a lycopodioides Griseb. from synonymy and tremendous amount of phenotypic plastic- reducing P. lechlerianum to a synonym of C. ity, even within the same individual cushion lycopodioides. plant. While carrying out field studies in Regarding the enigmatic Pycnophyllum Peru and Bolivia, I found that, depending lechlerianum Rohrb. that puzzled previous on factors such as branch age, position workers such as Grisebach, Pax, and Matt- within the cushion (central vs. peripheral), feld with its alleged 3-styled flowers, al- and micro-topography, characters used by though ten of the 12 type specimens (Lechler Mattfeld such as phylotaxis, leaf, and 1742) are sterile (as reported by all previous internode size, and even branching organi- workers), two (G, TCD) do bear flowers zation may vary tremendously. Other diag- which show a single style fused lengthwise nostic features such as shape, size, and with three stigmatic regions, as in all other position of the blade apex may dramatically members of Pycnophyllum. Consequently, change along a single primary branch the separation of this taxon into a separate depending on the leaf distance from the section, as P. sect. Drudea (Griseb.) Pax, is branch tip. Lastly, after comparing Matt- no longer supported. feld’s descriptions with extant type material, I have found that several, if not all, of his 10. Pycnophyllum kobalanthum Mattf., linear measurements, have been clearly Feddes Repert. Spec. Nov. Regni Veg. over-estimated. This fact is most burden- 36: 274, Tab. 179 (pp) (1934) — TYPE: some when, as stated earlier, his species BOLIVIA. Cordillera Real, Chacaltaya, definitions are based mainly on organ size. 4600 m, 21 Apr 1928, Troll 1906 Consequently, after long consideration and (HOLOTYPE: B!; ISOTYPE: M!). in light of these observations, I have decided to combine these previously recog- In my 2005 study, I considered P. nized species under the oldest binomial kobalanthum a synonym of P. tetrastichum. available, namely P. tetrastichum. While this New material from LPB made available to solution may be less than perfect, I fail to me in 2015 convinced me to maintain its NUMBER 20 TIMANA:´ NOMENCLATURE OF PYCNOPHYLLOPSIS AND PYCNOPHYLLUM 21

species status. P. kobalanthum is an endemic Excluded names Bolivian species found between 4000 and 4600 m; it can be recognized by its Pycnophyllum aschersonianum Muschl., characteristic gibbous leaves with the apex Bot. in Jahrb. Syst. 45: 455. 1911 — TYPE: suddenly arched into a small mucron. PERU. Ancash: Cordillere Janashallath, inter Additional Representative specimens: BO- Huaraz et Chavin de Huantar, in formatione LIVIA. Chuquisaca:CordilleradeLos plantis caespitosis ac pulvinaribus composi- Sombreros, near summit along road from ta, 4500- 4600 m, 3 Jul 1903, Weberbauer 0 0 Icla to Tarvita, 19834 50’’ S, 64837 50’’ O, 3293 (HOLOTYPE: B†). 3929 m, 1 Dec 2005, Wood et al. 22156 (LBP, Because of the lack of original material TEX). La Paz: Area´ Natural de Manejo and the contradictory nature of Muschler’s Integrado Apolobamba, cerro Socondori description (he described bisexual flowers Grande, aprox. 8 km NNE de Ulla Ulla, for a dioecious species), I cannot assign this 15800042’’ S, 69813002’’ O, 4600 m, 27 Feb taxon to any known species. The closest 2008, Meneses 4506 (LBP, TEX); Umanata, taxon with which this description fits is that Jaquewi Lampampa, 15826058’’ S, 69804056’’ of Pycnophyllum huascaranum, although the O, 4149 m, 25 Nov 2003, Valdivia et al. 88. two greatly differ in petal size. Muschler (LBP, TEX); camino a Alto Chacaltaya, 200 described his specimen as having petal and m entrando hacia el camino de la comp. stamens of equal size, which is not the case Minera Kellguani, 4320 m, 21 Dec 1986, in the newly described species where the Valenzuela 917 (LBP, TEX); Patapampa, 2.5 petals are always much shorter than the km del desvıo,´ 4440 m, 21 Dec 1986, stamens. Mattfeld (1922) pointed out the Valenzuela 937 (LBP, TEX). Oruro: entre overall leaf similarities with P. molle and P. Challapata y Potosı,´ 4050 m, 27 Feb 1991, spathulatum but was unable to assign a Navarro 234 (LBP, TEX); bosque de Poly- species status. lepis, arriba del pueblo Sajama, 4600 m, 1 Jun 1991, Hensen 2623 (LBP, TEX); Parque Pycnophyllum carinatum Muschl., Bot. Nacional Sajama, cerro Huincurata, Jahrb. Syst. 45: 457 (1911) ” Arenaria 18807005’’ S, 68857044’’ O, 4555 m, 25 Mar carinata (Muschl.) Molinari, Polish Bot. J. 2012, Pozo & Zeballos 759 (LBP, TEX). 61(2): 275 (2016). SYNTYPES: PERU. Potosı´: al norte de Pulacayo, aprox. 13.5 km, Junin: between Tarma and La Oroya, 4000 cerca a la cabezera Calzada, 20815014.66’’ S, m, 14 Feb 1903, Weberbauer 2529 (B†); 6684106.25’’ W, 4470 m, 10 Mar 2010, Puno, Sandia, between Poto and Ananca, Zenteno & Mamani 9774 (LBP, TEX); along 4600-4700 m, 7 May 1902, A. Weberbauer road between Villa Alota and Viscacacha, 951 (B†). 21823038’’ S, 67843011’’ O, 4100 m, 7 Mar Mushler described this species based on 2009, Kool et al. 1221 (LBP, TEX); al oeste- two gatherings by August Weberbauer. sur-oeste cruce a Nazanerito v Kucho, Weberbauer 2529 is most probably destroyed 21826011’’ S, 65853002’’ O, 4398 m, 10 Mar – it has not been located at either B, MOL, 2012, Zenteno et al. 11985 (LBP, TEX); al SO or any other herbarium. The second speci- de Kucho en lınea´ recta hacia Moiinete men, Weberbauer 951, has not been located aprox 18.93 km, 21837041’’ S, 66808048’’ W, in either of the two herbaria. A specimen 4453 m, 12 Mar 2012, Zenteno et al. 12108 with the same collection number is listed as (LBP, TEX); along road between Uyuni and the holotype of Bulbophyllum incarum Laguna Colorada, after passing Laguna Kraenzl. (Orchidaceae, Kraenzlin, 1905) Cachi, 21849057’’ S, 69812016’’ W, 4600 m, and cited as such by Schweinfurth in his 7 Mar 2009, Kool et al. 1223 (LBP, TEX); on ‘‘Orchids of Peru’’ (Schweinfurth, 1960), pass above Belen Pampa, along road from however, the image of the F photograph Potosı´ to Challapata, 19813011’’ S, 66803016’’ presented in JSTOR Global Plants carries the O, 4265 m, 14 Mar 2009, Kool et al. 1241 number 591 (F0BN018331, photo!) instead (LBP, TEX). of 951. In 1994, Christenson (1994) de- 22 LUNDELLIA DECEMBER, 2017

scribed the orchid collection at the Herbar- the Cordillera Illapel in the Province of ium Hamburgense (HGB); he reported (p. Coquimbo, Chile (Philippi, 1892). Philippi 348) specimen Weberbauer 951 as the missed the small, inconspicuous flowers holotype of B. incarum (HBG501372, pho- concealed by the dense indumentum that to!). The Weberbauer specimen that Musch- characterizes Pycnophyllum lanatum and ler examined for the description of P. therefore proposed the new species with carinatum remains a mystery. He described some reservation, adding a question mark a ciliated leaf margin, an unknown charac- after the generic name, thus as Pycnophyllum teristic among species of Pycnophyllum; ? lanatum. Mattfeld (1922), the last mono- Mattfeld (1922) suggested this could be a grapher of Pycnophyllum, was unable to member of Arenaria sect. Dicranilla; finally, study any part of the original material and Macbride (1937) speculated this taxon could listed this taxon as species dubia, placing be conspecific with Arenaria boliviana Wil- some doubts on its generic position due to liams. Thus, while Molinari’s combination its heavy white indumentum. The examina- might be correct from a nomenclatural tion of additional specimens has demon- point of view, it is also purely speculative strated that this taxon does not belong in since the true identity of this taxon cannot Pycnophyllum. Further details will be given be confirmed. in a publication in preparation (Timana´, in prep.) Pycnophyllum dicranoides (Kunth) Muschl, Bot. Jahrb. Syst. 45: 454 (1911) ” Arenaria Pycnophyllum peruvianum Muschl., Bot. dicranoides Kunth, Nov. Gen. et Spec. VI: 34 Jahrb. Syst. 45: 457 (1911) ” Arenaria (Apr. 1823 -folio). — TYPE: ECUADOR. peruviana (Muschl.) Molinari, Polish Bot. ‘‘Cresit in monte Antisana, alt. 2100 hex. J. 61(2): 276 (2016). TYPE: PERU. Junin: La floret Junio.’’ A. Humboldt & A. Bonpland, Oroya, 4300 m, Feb 1903, Weberbauer 2597 s.n (HOLOTYPE: P! P00274230). (LECTOTYPE selected by Molinari, 2016: This is a synonym of Arenaria dicra- MOL!). nioides Kunth. Muschler (1911) cited We- As proposed earlier by Mattfeld (1922), berbauer 5120 (B) when proposing this this is a member or Arenaria sect. Dicranilla. combination but no duplicates have been Mushler described ciliated leaf margins and located. bisexual flowers, both features known to be absent in members of Pycnophyllum. The Pycnophyllum horizontale Muschl., Bot. Weberbauer specimen cited by Molinari and Jahrb. Syst. 45: 454 (1911) ” Arenaria deposited at MOL emerges under unusual horizontalis (Muschl.) Molinari, Polish Bot. circumstances. As part of my research I J. 61(2): 275 (2016). TYPE: PERU. Lima: visited MOL in 1999, 2000 and 2016; during ‘‘Andes altissimae supra Lima’’, 4500m, Aug my first two visits Weberbauer 2597 was not 1905, Weberbauer s.n. (HOLOTYPE: B†). in the collection. In fact, such a specimen is Muschler (1911) described ciliated not mentioned in Velarde’s authoritative leaves and bisexual flowers in this specimen, catalogue of Weberbauer’s isotypes kept at none of which occur in Pycnophyllum. the MOL herbarium (Velarde 1969). The Mattfeld (1922) suggested this could be a specimen location is not listed in neither the member of Arenaria sect. Dicranilla. No Cano and Sa´nchez report on endemic duplicates have been located. Caryophyllaceae of Peru (Cano & Sa´nchez, 2006), nor is included, to July 2017, among Pycnophyllum lanatum Phil. in Anal. Univ. MOL type specimens submitted to JSTOR Santiago 81: 774 (1893) — TYPE: CHILE. Plants in 2013. The specimen itself lacks all Las Mollucas, Cordillera de Illapel, Jan.1888, the characteristic elements of the Weber- R. Philippi s.n. (HOLOTYPE: SGO!) bauer specimens stored at MOL: no original This name was published by R.A. label (with Weberbauer’s distinctive hand- Philippi based on specimens collected near writing), barcode, or official herbarium seal NUMBER 20 TIMANA:´ NOMENCLATURE OF PYCNOPHYLLOPSIS AND PYCNOPHYLLUM 23

and number (compared with, for example, friendship with me since then. Jorge Chia- the type specimen of Pycnophyllum filiforme pella (INBIV, Cordoba)´ and Carlos Martel available in JSTOR Plants (Weberbauer 6885, (U. of Ulm) provided much-needed support MOL-00000494, photo!). The fragmentary and advice when needed. Sabine von Mering material is sterile, thus allowing limited (B) assisted me with arranging a visit to the comparison with Mushler’s description and Berlin herbarium and issues related to therefore it is impossible to assert that it is Weberbauer’s type specimens. I thank indeed a P. peruvianum specimen. Christine Niezgoda (F), M. Jeanson (P), Carmen Ulloa (MO) and Lauren Peters Pycnophyllum pilgerianum Muschl., Bot. (MO) for facilitating obtaining digital im- Jahrb. Syst. 45: 454 (1911) – TYPE: BOLI- ages of type specimens. I would like to thank VIA. Puna de Patanca, 3700 m, 8 Jan 1904, curator Tom Wendt (TEX) for acquiring all Fiebrig 2617 (HOLOTYPE: B†); LECTOTYPE, herbarium loans required and supporting designated here: US!; ISOLECTOTYPES: BM!, my project. The following herbaria are G! [photo], LD!, Z! (2 sets) ” Frankenia acknowledged for providing specimens on triandra J. Remy.´ See Whalen, 1987. loan for morphological study: AAS, AAU, AK, ASU, B, BA, BAB, BM, BR, C, CAS, Pycnophyllum stuebelii Mattf., Feddes Re- CGE, CHR, COL, COLO, CORD, CTES, E, pert. Spec. Nov. Regni Veg. 18: 171 (1922). F, FI, G, GB, GH, GOET, H, HAL, HO, K, L, TYPE: BOLIVIA. Puna de Sicasica, ca. 5000 LD, LP, LPB, LY, M, MA, MCNS, MEL, m, 2 Nov 1876, Stubel¨ 68 (HOLOTYPE: B†). MICH, MO, MOL, MSC, NA, NHA, NSM, Pycnophyllum stuebelii is a most enig- NSW, NY, OS, OXF, P, PH, PR, PRE, RNG, matic taxon. Mattfeld commented that this S, SGO, SI, TCD, TEX, TUB, UC, UPS, US, species presented similarities with Pycno- USM, W, WIS, WU, Z. RBG Kew’s Latin phyllum lechlerianum (here P. tetrastichum). America Research Fellowship Programme Based on the leaf and bract sizes it would fit (KLARF) allowed me to visit several major near Mattfeld’s P. glomeratum but the lack of European herbaria to study historical col- original material precludes confirming this. lections. The Department of Humanities and the Applied Geography Research Center Pycnophyllum sulcatum Griseb., Pl. lorentz. (CIGA) at PUCP provided me with logistic 28, tab. 1, fig. 1 (Dec. 1874); also in Abh. support and funding to visit B and TEX in Kon.¨ Ges. Wiss. Gottingen¨ 19: 76, tab. 1, fig. 2015. This paper is dedicated to the memory 1 (1874). TYPE: ARGENTINA. Catamarca: of Dr. William ‘‘Bill’’ Anderson, former in salsi convallium alpinarum inter Laguna Director of the MICH herbarium, and dear blanca et Nacimiento, Jan 1872, Lorentz 458 friend and teacher. (LECTOTYPE (selected by Whalen, 1987): GOET!); ISOLECTOTYPE: CORD (photo!). ” Frankenia triandra J. Remy.´ See Whalen, LITERATURE CITED 1987. Bittrich V. 1993. Caryophyllaceae. In: K. Kubitzki, J. Rohwer, V. Bittrich, eds. The Families and Genera of ACKNOWLEDGMENTS Vascular Plants 2. Magnoliid, Hamamelid, and Caryophyllid Families. Berlin: Springer-Verlag, pp. This work represents a portion of a 206–236. Cano, A. & I. Sa´nchez. 2006. Caryophyllaceae Ph.D. Dissertation submitted to the Gradu- endemica´ del Peru.´ Rev. Peru. Biol. 13: 242s–252s. ate School of The University of Texas, Christenson, E.A. 1994. Significant collections of Austin. I would like to thank my academic Orchidaceae conserved in herbarium Hamburgense advisor, Dr. Beryl B. Simpson who guided (HBG). Brittonia 46: 344–354. me throughout this research. Dr. Richard Grayum, M. H, B. Hammel, and N. Zamora. 2016. The Cutting Edge 32(2): April. URL: http://www. (Rich) Rabeler (MICH) introduced me to mobot.org/MOBOT/research/Edge/welcome.shtml. the world of the Caryophyllaceae 20 years Grisebach, A. H. R. 1854. Systematische Bemerkungen. ago and has shared his expertise and Gottingen,¨ Dieterich. 24 LUNDELLIA DECEMBER, 2017

———. 1879. Symbolae ad Floram Argentinam. Pax, F. 1893. Uber¨ die Verbreitung der Gottingen,¨ Dieterich. sudamerikanischen¨ Caryophyllaeceae und die Arten Harbaugh, D. T., M. Nepokroeff, R. K. Rabeler, J. der Republica Argentina. Bot. Jahrb. Syst. 18: 1–35. McNeill, E. A. Zimmer, and W. L. Wagner. 2010. Philippi, R. A. 1892. Plantas nuevas chilenas. Anales A new lineage-based classification of the family Univ. Chile 81: 761–775. Caryophyllaceae. Int. J. Pl. Sci. 171(2): 185–198. Remy,´ J. 1846. Anacleta Boliviana. Ann. Sci. Nat., Bot. Herna´ndez-Ledesma P., W. G. Berendsohn, Th. ser.´ 3, 6: 345–357 Borsch, S. von Mering, H. Akhani, S. Arias, I. ———. 1847. Anacleta Boliviana. Ann. Sci. Nat., Bot. Castaneda-Noa,˜ U. Eggli, R. Eriksson, H. Flores- ser.´ 3, 8: 224–239. Olvera, S. Fuentes-Baza´n, G. Kadereit, C. Klak, N. Rohrbach, P. 1870. Beitrage¨ zur Systematik der Korotkova, R. Nyffeler, G. Ocampo, H. Ochoter- Caryophyllinen. I. Uber die Gattung Pycnophyllum. ena, B. Oxelman, R. K. Rabeler, A. Sanchez, B. O. Linnaea 36: 651–664. Schlumpberger, and P. Uotila. 2015. A taxonomic ———. 1872. Beitrage¨ zur Systematik der Caryophyl- backbone for the global synthesis of species linen. III. Linnaea 37: 183–312. diversity in the angiosperm order Caryophyllales. Schweinfurth, C. 1960: Orchids of Peru. Fieldiana Bot. Willdenowia 45: 281–383. 30: 533–786. Hiepko, P. 1987. The collections of the Botanical Skottsberg, C. 1916. Die Vegetationsverhaltnisse¨ langs¨ Museum Berlin-Dahlem (B) and their history. der Cordillera de los Andes S. von 418 S. Br. Kungl. Englera 7: 219–252 Svenska Vetenskapsakad. Handl. 56: 1–366 Kraenzlin, F.W.L. 1905. Orchidaceae novae Austro- Smith, D. N. 1988. Flora and vegetation of the americanae plerumque Peruanae. Repert. Spec. Huascaran National Park, Ancash, Peru, with Nov. Regni Veg. 1: 85–92 preliminary taxonomic studies for a manual of the MacBride, J. F. 1937. Caryophyllaceae. Flora of Peru. flora. Ph.D Dissertation. Iowa State University. Publ. Field Mus. Bot. 13(2): 578–639. Ames, Iowa. Stafleu, F.A. and R. S. Cowan. 1981. Taxonomic Mattfeld, J. 1922. Revision der Gattung Pycnophyllum Literature. A Selective Guide to Botanical Publica- Remy. Feddes Repert. 17: 167–179. tions and Collections with Dates, Commentaries, and ———. 1930. Pycnophyllum leptothamnum. Notizbl. Types, ed 2, vol. 3. Dr. W. Junk, The Hague. Bot. Gart. Berlin–Dahlem 10: 1051–1052. ———. 1983: Taxonomic Literature. A Selective Guide ———. 1934a. Plettkea eine neue Gattung der to Botanical Publications and Collections with Dates, Alsinoideae aus den Hochanden Perus. Schriften Commentaries, and Types, ed 2, vol. 4. Dr. W. Junk, Vereins Naturk. Unterweser. N.F. VII: 1–22. Publishers, The Hague. ———. 1934b. Nachtrag zu den Caryophyllaceae. In: Stearn, W. T. 1973. An introduction to K. B. Presl’s A. Engler and K. Prantl (eds.), Die naturlichen¨ Reliquiae Haenkeanae (1825- 1835). In: K. B. Presl’s Pflanzenfamilien, ed. 2, 16c: 365 – 367. Leipzig: Reliquiae Haenkeanae seu descriptiones et icones Engelmann. plantarum, quas in America meridionali et boreali, McNeill, J. 1962. Taxonomic studies in the Alsinoi- in insulis Philippinis et Marianis collegit Thaddaeus deae: I. Generic and infra-generic groups. Notes Haenke. Reprint Edition. A. Asher. Amsterdam. Roy. Bot. Gard. Edinburgh 24 (2): 79–155. Thiers, B. [continuously updated]. Index Herbario- McNeill, J.; Barrie, F.R.; Buck, W.R.; Demoulin, V.; rum: A global directory of public herbaria and Greuter, W.; Hawksworth, D.L.; Herendeen, P.S.; associated staff. New York Botanical Garden’s Knapp, S.; Marhold, K.; Prado, J.; Prud’homme Virtual Herbarium. http://sweetgum.nybg.org/ih/ Van Reine, W.F.; Smith, G.F.; Wiersema, J.H.; Timana´, M. E. 2005. Systematic Studies in Pycnophyl- Turland, N.J. 2012. International Code of Nomen- lum and Pycnophyllopsis (Caryophyllaceae) of the clature for algae, fungi, and plants (Melbourne Code) High Andes.Ph.D.Dissertation.Austin:The adopted by the Eighteenth International Botanical University of Texas, School of Biological Sciences. Congress Melbourne, Australia, July 2011. Regnum Velarde, O. 1969. Cata´logo de isotipos de la coleccion´ Vegetabile 154. A.R.G. Gantner Verlag KG. de plantas peruanas de A. Weberbauer que se Molinari, E. 2016. Further nomenclatural changes conserva en el herbario de la Universidad Nacional regarding Peruvian endemics. Polish Bot. J. 61: Agraria del Peru.´ Raymondiana 2: 115–147. 275–277. Whalen, M. A. 1987. Systematics of Frankenia Muschler, R. 1911. Caryophyllaceae andinae. Bot. (Frankeniaceae) in North and South America. Syst. Jahrb. Syst. 45: 441–461. Bot. Monogr. 17: 1–93.