CUSHIONS OF SOUTH AMERICA (EXCEPT BRAZIL)

INTRODUCTION

Cushion plants represent a special life form which usually has character combinations such as short-node intervals, compact branches, solitary flowers or few-flowered racemes, and dome- or mat-shaped cushions. They are common among perennial herbs growing on high-altitude mountains and are thought to be associated with dry and cold environments, such as the high Andes and Patagonia, Himalayas, and New Zealand Alps. Aubert et al. (Alpine Botany, 2014) updated the cushion plants catalogue in which they recognized 1,309 species of 63 families and 273 genera.

This post aims to list the cushions from South America in light of new molecular and distributional data. The primary list follows Alpine Cushions (SEE), but with due corrections via POWO (SEE); names assigned to a country in Alpine Cushions but not assigned in the VPA are in salmon color; the reverse situation (which are all accepted here) is marked in blue.

In this compilation we list 280 cushions in 86 genera at 38 families among 20 orders, 18 if them in Eudicots. By country, the diversities are Venezuela (21 spp. in 12 genera at 10 families), Colombia (38 spp. in 20 genera at 13 familes), Ecuador (53 spp. in 28 genera at 18 families), Peru (72 spp. in 35 genera at 22 families), Bolivia (64 spp. in 34 genera at 22 families), Argentina (194 spp. in 67 genera at 31 families), and Chile (158 spp. in 59 genera at 30 families). Gentianaceae, Malvaceae and Saxifragaceae occur only from Ecuador southwards. Fabaceae, Lamiaceae, Verbenaceae and Frankeniaceae occur only from Peru southwards; Oxalidaceae and Bromeliaceae occur only in Bolivia southwards. Restionaceae, Iridaceae, Asteliaceae, Rosaceae, Violaceae, Thymelaeaceae, Plumbaginaceae, Montiaceae, Solanaceae, Rubiaceae occur only in Argentina and Chile; Loasaceae only in Chile.

The Top 5 genera (Azorella, Nototriche, Junellia, Pycnophyllum and Benthamiella) composes more than 1/3 of all cushions in South America (90 spp). None of them occur in Brazil. Taxonomic changes: Phyllactis and Aretioides under Valeriana; Petunia under Fabiana; Mulinum under Azorella; Belloa under Mniodes; Deuterocohnia under Dyckia.

Some claims for cushions in South America are not on this list. Jalcophila M.O. Dillon & Sagást. and Aphanactis Wedd. in Asteraceae (K.Kubitzki, vol. VIII, 2007); Englerocharis Musch. (Brassicaceae) from Peru (Al-Shehbaz et al., Kew Bulletin, 2012); Gamocarpha DC. and Moschopsis Phil. (Calyceraceae) from Chile and Argentina (Pozner & al., Taxon, 2021). Ribes frankei Weigend & Breitkopf (Grossulariaceae), endemic to mountains of Peru, is the unique cushion-like species among this family (Weigend ey al., Novon, 2010).

Although there is no citation in the literature, many species of Poales do Brazil could be considered cushions, within the current limits adopted, namely Paepalanthus glabrifolius Ruhland, P. pseudotortilis Ruhland, Leiothrix plantago (Mart.) Giul. and L. curvifolia Ruhland (Eriocaulaceae), Abolboda killipii Lasser and A. americana (Aubl.) Lanj. (Xyridaceae); all of these are endemic to Brazil except the Abolboda Bonpl., which extend into the adjacent countries of the Guiana Shield. In Mexico there are only 13 cushions in 6 genera of two families: Cactaceae with Echinocereus (2), Epitelantha (1), Ferocactus (1), and Mammilaria (7), and Caryophyllaceae with Arenaria bryoides and Colobanthus quitensis.

TAXONOMY

IRIDACEAE ‣ Tapeinia pumila (G.Forst.) Baill. (AR, CH).

ASTELIACEAE ‣ Astelia pumila (J.R.Forst.) Gaudich. (AR, CH).

RESTIONACEAE ‣ Gaimardia australis Gaudich. (AR, CH).

BROMELIACEAE ‣ Dyckia grisebachii Baker (BL, AR syn. Deuterocohnia brevifolia, inc. Deuterocohnia lorentziana).

ERIOCAULACEAE ‣ Paepalanthus karstenii Ruhland (VZ, CL, EC, PR, BL), P. ferreyrae Moldenke (PR), P. lodiculoides Moldenke (VZ, CL, EC, PR), P. pilosus (Kunth) Kunth (VZ, CL, EC, PR), P. stegolepoides Moldenke (VZ); for pulvinate species in northern Andes, see Hensold (PhytoKeys, 2016).

JUNCACEAE ‣ Distichia acicularis Balslev & Laegaard (EC), D. filamentosa Buchenau (PR, BL, CH), D. muscoides Nees & Meyen (CL, EC, PR, BL, AR, CH), Oxychloe andina Phil. (PR, BL, AR, CH), O. bisexualis Kuntze (AR, CH), O. castellanosii Barros (AR), O. haumaniana (Barros) Barros (AR, CH), O. mendocina Barros (AR), Patosia clandestina (Phil.) Buchenau (BL, AR).

Distichia muscoides Nees & Meyen, D. filamentosa Buchenau, Oxychloe andina Phil.

CYPERACEAE ‣ Carpha alpina R.Br. (CH), Oreobolus cleefii L.E.Moro (CL, EC, PR), O. ecuadorensis T.Koyama (CL, EC, PR), O. goeppingeri Suess. (VZ, CL, EC, PR, BL), O. obtusangulus Gaudich. (VZ, CL, EC, PR, BL, CH), O. venezuelensis Steyerm. (VZ, CL, EC, PR, BL), Rhynchospora oreoboloidea Gómez-Laur. (CL, EC), Phylloscirpus deserticola (Phil.) Dhooge & Goetgh. (EC, PR, BL, AR, CH), Zameioscirpus atacamensis (Phil.) Dhooge & Goetgh. (PR, BL, AR), Z. gaimardioides (É.Desv.) Dhooge & Goetgh. (PR, AR, CH), Z. muticus Dhooge & Goetgh. (PR, BL, AR, CH); for range in northern Andes, see Springer Nature (LINK).

Oreobolus obtusangulus Gaudich., Zameioscirpus atacamensis (Phil.) Dhooge & Goetgh.

XYRIDACEAE ‣ Xyris subulata Ruiz & Pav. (GU, VZ, CL, EC, PR).

POACEAE ‣ Aciachne pulvinata Benth. (VZ, CL, EC, PR, BL), A. acicularis Laegaard (VZ, CL, EC, PR, BL, AR), A. flagellifera Laegaard (CL, EC), Deschampsia venustula Parodi (AR, CH).

Aciachne pulvinata Benth., A. acicularis Laegaard

RANUNCULACEAE ‣ Caltha appendiculata Bercht. & J.Presl (AR, CH), C. sagittata Cav. (EC, PR, BL, CH).

SAXIFRAGACEAE ‣ Saxifraga magellanica Poir. (EC, PR, BL, AR, CH, inc. S. adenodes Poepp. ex Sternb.), Saxifragella bicuspidata (Hook. f.) Engl. (AR, CH).

Saxifraga magellanica Poir.

FABACEAE ‣ Adesmia aconcaguensis Burkart (AR), A. aegiceras Phil. (AR, CH), A. ameghinoi Speg. (AR), A. aueri Burkart (AR), A. caespitosa Phil. (AR, CH), A. digitata Burkart (AR), A. echinus C.Presl (AR, CH), A. erinacea Phil. (BL, AR, CH), A. frigida Phil. (CH), A. glomerula Clos (AR, CH), A. neglecta M.N.Correa (AR), A. horrida Hook. & Arn. (BL, CH), A. occulta (R.E.Fr.) Burkart (PR, BL, CH), A. pentaphylla Phil. (AR, CH), A. salicornioides Speg. (AR, CH), A. serrana M.N.Correa (AR), A. subterranea Clos (AR, CH), Anarthrophyllum burkartii Soraru (AR), A. desideratum (DC.) Benth. (AR, CH), A. gayanum (A.Gray) B.D.Jacks. (AR, CH), A. patagonicum Speg. (AR).

Adesmia erinacea Phil., A. occulta (R.E.Fr.) Burkart

OXALIDACEAE ‣ Oxalis compacta Gillies ex Hook. & Arn. (AR), O. erythrorrhiza Gillies ex Hook. & Arn. (AR), O. holosericea Phil. (CH), O. hypsophila Phil. (BL, CH, AR), O. loricata Dusén (AR), O. muscoides Phil. (AR).

Oxalis hypsophila Phil.

ROSACEAE ‣ Acaena alpina Poepp. ex Walp. (AR, CH), A. caespitosa Gillies ex Hook. & Arn. (AR, CH), A. confertissima Bitter (AR), A. lucida Vahl (AR, CH).

VIOLACEAE ‣ Viola auricolor Skottsb. (AR, CH), V. dasyphylla W.Becker (AR).

THYMELAEACEAE ‣ Drapetes muscosus Hook.f. (AR, CH).

BRASSICACEAE ‣ Brayopsis colombiana Al-Shehbaz (CL, EC), B. calycina (Desv.) Gilg & Muschl. (PR, BL, AR), B. diapensioides (Wedd.) Gilg & Muschl. (PR, BL), Dactylocardamum imbricatifolium Al-Shehbaz (PR), Delpinophytum patagonicum (Speg.) Speg. (AR), Draba aretioides Humb. & Bonpl. ex DC. (EC), D. depressa Hook.f. (CL, EC, PR), D. gilliesii Hook. & Arn. (AR, CH), D. obovata Benth. (EC), Eudema nubigena Humb. & Bonpl. (EC), E. hauthalii Gilg & Muschl. (AR, CH), E. werdermannii O.E.Schulz (AR, CH), Lithodraba mendocinensis (Hauman) Boelcke (AR), Onuris hatcheriana (Gilg ex Macloskie) Gilg & Muschl. (AR, CH), Physaria okanensis Al-Shehbaz & Prina (AR), Xerodraba colobanthoides Skottsb. (AR), X. glebaria (Speg.) Skottsb. (AR), X. lycopodioides (Speg.) Skottsb. (AR), X. monantha (Gilg ex Kuntze) Skottsb. (AR), X. patagonica (Speg.) Skottsb. (AR, CH), X. pectinata Skottsb. (AR, CH), X. pycnophylloides (Speg.) Skottsb. (AR).

Draba depressa Hook.f., Dactylocardamum imbricatifolium Al-Shehbaz

Brayopsis colombiana Al-Shehbaz, Eudema nubigena Humb. & Bonpl.

GERANIACEAE ‣ Geranium crassipes Hook. ex A.Gray (CL, PR), G. sericeum Willd. ex Spreng. (EC).

Geranium sericeum Willd. ex Spreng.

MALVACEAE ‣ Nototriche azorella Hill (PR, CH), N. auricoma A.W. Hill (CH), N. compacta A.W. Hill (PR).

, CH, AR, CH), N. ecuadoriensis Fryxell (EC), N. flabellata (Wedd.) A.W. Hill (PR, BL), N. holoserica A.W. Hill (AR, CH), N. jamesonii A.W.Hill (EC, syn. N. hartwegii A.W. Hill), N. obcuneata (Baker f.) A.W. Hill (PR, BL, CH), N. orbignyana (Wedd.) A.W. Hill (PR, BL, CH), N. parviflora A.W. Hill (AR, CH), N. pediculariifolia (Meyen) A.W. Hill (PR, BL, CH), N. philippii A.W. Hill (AR, CH), N. phyllanthos (Cav.) A.W. Hill (EC, PR, inc. N. pichinchensis (Bonpl.) A.W. Hill), N. polygama Krapov. (BL), N. pulvilla A.W. Hill (AR), N. sajamensis (Hieron.) A.W. Hill (PR, BL), N. stipularis (Phil.) A.E.Martic. (BL, CH), N. turritella A.W. Hill (PR, BL, CH).

Nototriche compacta A.W. Hill, Nototriche obcuneata (Baker f.) A.W. Hill, Nototriche phyllanthos (Cav.) A.W. Hill

FRANKENIACEAE ‣ Frankenia triandra Remy (PR, BL, AR, CH; inc. Pycnophyllum pilgerianum Muschl).

Frankenia triandra Remy

PLUMBAGINACEAE ‣ Armeria curvifolia Bertero (AR, CH), A. maritima Willd. (AR, CH).

MONTIACEAE ‣ Calandrinia caespitosa Gillies ex Arn. (AR), C. colchaguensis Barnéoud (AR), C. ferruginea Barnéoud (CH), C. potentilloides Barnéoud (AR), C. sericea Hook. & Arn. (AR).

CACTACEAE ‣ Airampoa soehrensii (Britton & Rose) Lodé (PR, BL, CH, inc. Tunilla soehrensii (Britton & Rose) D.R. Hunt & Iliff), Austrocylindropuntia floccosa (Salm-Dyck ex Winterfeld) F.Ritter (PR, BL), Copiapoa ahremephiana N.P.Taylor & G.J.Charles (CH), C. cinerascens (Salm-Dyck) Britton & Rose (CH), C. dealbata F.Ritter (CH), C. longistaminea F.Ritter (CH), C. solaris (F.Ritter) F.Ritter (CH), Cumulopuntia boliviana (Salm-Dyck) F.Ritter (PR, BL, AR, CH; inc. Maihueniopsis bolivianum (Salm-Dyck) R. Kiesling and M. pentlandii (Salm-Dyck) R. Kiesling), Gymnocalycium monvillei (Lem.) Pfeiff. ex Britton & Rose (AR), M. glomerata (Haw.) R. Kiesling (BL, AR, CH), Opuntia atacamensis Phil. (CH).

Austrocylindropuntia floccosa (Salm-Dyck ex Winterfeld) F.Ritter, Cumulopuntia boliviana (Salm-Dyck) F.Ritter, Tunilla soehrensii (Britton & Rose) D.R. Hunt & Iliff

CARYOPHYLLACEAE ‣ Arenaria bisulca (Bartl.) Fenzl ex Rohrb. (PR, AR), A. denticulata Phil. (CH), A. dicranoides Kunth (VZ, EC, PR, BL), A. musciformis Triana & Planch. (VZ, CL, EC), A. venezuelana Briq. (VZ, CL), Colobanthus quitensis (Kunth) Bartl. (CL, EC, PR, BL, CH), C. lycopodoides Griseb. (AR, CH), C. subulatus (d'Urv.) Hook.f. (AR, CH), Paronychia sessiliflora Nutt. (BL), Pycnophyllum argentinum Pax (BL, AR), P. bryoides (Phil.) Rohrbach (PR, BL, AR, CH), P. glomeratum Mattf. (PR, AR, CH), P. macropetalum Mattf. (PR, BL, AR, CH), P. mattfeldii J.F. Macbr. (PR, BL), P. molle Remy (CL, PR, BL, AR, CH), P. spathulatum Mattf. (BL, CH), P. stuebelii Mattf. (BL), P. tetrastichum Remy (PR, BL, AR, CH, inc. P. kobalanthum Mattf.), P. muscosa Skottsb. (AR), P. convexum Griseb. (AR), P. mucronulatum Mattf. (AR), Philippiella patagonica Speg. (AR, CH), Reicheella andicola (Phil.) Pax (CH), Silene haumanii Bocquet (AR).

Arenaria bisulca (Bartl.) Fenzl ex Rohrb., A. dicranoides Kunth, A. musciformis Triana & Planch.

Pycnophyllum bryoides (Phil.) Rohrbach, P. molle Remy

Colobanthus quitensis (Kunth) Bartl.

LOASACEAE ‣ Loasa caespitosa Phil. (CH).

ERICACEAE ‣ Disterigma empetrifolium (Kunth) Nied. (VZ, CL, EC, PR, BL).

Disterigma empetrifolium (Kunth) Nied.

GENTIANACEAE ‣ Gentianella armerioides (Griseb. ex Gilg) J.S.Pringle (PR, BL), Halenia minima C.K. Allen (EC), Halenia pusilla Gilg (PR, BL).

Halenia minima C.K. Allen

RUBIACEAE ‣ Oreobolus obtusangulus Gaudich. (AR), O. glacialis (Poepp.) Ricardi (AR, CH).

SOLANACEAE ‣ Benthamiella azorella (Skottsb.) C.Soriano (AR), B. azorelloides Speg. (AR, CH), B. chubutensis C.Soriano (AR), B. graminifolia Skottsb. (AR), B. lanata C.Soriano (AR), B. longifolia Speg. (AR), B. nordenskjoldii Dusén (AR), B. patagonica Speg. (AR, CH), B. pycnophylloides Speg. (AR), B. skottsbergii C.Soriano (AR), B. sorianoi S.Arroyo (AR), B. spegazziniana C.Soriano (AR, CH), Combera minima Sandwith (CH), Fabiana australis Alaria (AR), F. foliosa (Speg.) S.C.Arroyo (AR), F. nana (Speg.) S.C.Arroyo (AR), Pantacantha ameghinoi Speg. (AR).

CALCEOLARIACEAE ‣ Calceolaria mendocina Phil. (AR, CH), C. pennellii Descole & Borsini (AR), C. pinifolia Cav. (AR, CH), C. prichardii (Rendle) Kraenzl.

LAMIACEAE ‣ Hedeoma mandoniana Wedd. (PR, BL, AR).

Hedeoma mandoniana Wedd.

PLANTAGINACEAE ‣ Plantago barbata G.Forst. (AR, CH), P. rigida Kunth (VZ, CL, EC, PR, BL), P. sempervivoides Dusén (AR, CH), P. tubulosa Decne (VZ, EC, PR, BL), P. uniglumis Wallr. ex Walp. (AR, CH).

Plantago rigida Kunth, P. tubulosa Decne 

VERBENACEAE ‣ Junellia aretioides (R.E.Fr.) Moldenke (BL, AR, CH), J. azorelloides (Speg.) Moldenke (AR), J. caespitosa (Gillies & Hook., CH) Moldenke (AR), J. congesta (Tronc.) Moldenke (AR), J. digitata (Phil.) Moldenke (BL, CH, AR, CH), J. erinacea (Gillies & Hook.) Moldenke (AR), J. micrantha (Phil.) Moldenke (CH), J. minima (Meyen) Moldenke (PR, BL, AR), J. mulinoides (Speg.) Moldenke (AR), J. odonellii Moldenke (AR, CH), J. pappigera (Phil.) N. O'Leary & P. Peralta (AR, CH), J. patagonica (Speg.) Moldenke (AR, CH), J. silvestrii (Speg.) Moldenke (AR, CH), J. toninii (Kuntze) Moldenke (AR, CH), J. uniflora (Phil.) Moldenke (AR, CH).

Junellia minima (Meyen) Moldenke 

ASTERACEAE ‣ Abrotanella diemii Cabrera (AR), A. emarginata (Cass. ex Gaudich.) Cass. (AR, CH), A. linearifolia A.Gray (CH), A. purpurea Swenson (AR, CH), A. submarginata A.Gray (CH), A. trichoachaenia Cabrera (AR, CH), A. trilobata Swenson (AR, CH), Baccharis aretioides Turcz. (EC), Brachyclados caespitosus (Phil.) Speg. (AR), Belloa chilensis Remy (BL, AR, CH), Burkartia lanigera (Hook. & Arn.) Crisci (AR), Chuquiraga aurea Skottsb. (AR), Erigeron leptopetalus Phil. (AR, CH), E. rosulatus Wedd. (PR, BL, AR, CH), Gamochaeta nivalis Cabrera (AR,CH), Lucilia kunthiana (DC.) Zardini (VZ, CL, EC, PR, BL), Mniodes aretioides Cuatrec. (PR, BL), M. caespititia (Wedd.) Quip. & M.O.Dillon (BL), M. longifolia (Cuatrec. & Aristeg.) Sagást. & M.O.Dillon (VZ, CL, EC, PR, BL), M. radians (Benth.) Sagást. & M.O.Dillon (VZ, CL, EC), M. schultzii (Wedd.) S.E.Freire, Chemisquy, Anderb. & Urtubey (EC, PR, BL, AR, CH inc. M. virescens (Wedd.) Cabrera), Monticalia microdon (Wedd.) B.Nord. (EC), Nardophyllum bryoides (Lam.) Cabrera (AR, CH), Nassauvia ameghinoi Speg. (AR), N. pentacaenoides Speg. (AR), Raouliopsis pachymorpha (Wedd.) S.F.Blake (CL), R. seifrizii S.F.Blake (CL), Senecio culcitenellus Cuatrec. (AR, CH), S. depressus Hook. & Arn. (AR, CH), S. digitatus Phil. (AR, CH), S. skottsbergii Cabrera (AR, CH), S. subdiscoideus Sch.Bip. ex Wedd. (PR, BL, AR, CH), Werneria aretioides Wedd. (PR, BL, AR, CH), W. humilis Kunth (CL, EC, PR, BL), W. poposa Phil. (PR, BL, AR), W. pumila Kunth (CL, EC, PR, AR), W. pygmaea Gillies ex Hook. & Arn. (VZ, CL, EC, PR, BL, CH), W. rosea Hieron. (EC), Xenophyllum incisum (Phil.) V.A.Funk (AR, CH), X. pseudodigitatum (Rockh.) V.A.Funk (AR, CH).

Erigeron rosulatus Wedd. and Werneria aretioides Wedd.

Mniodes schultzii (Wedd.) Cabrera, Senecio subdiscoideus Sch.Bip. ex Wedd., Xenophyllum humile (Kunth) V.A.Funk

STYLIDIACEAE ‣ Donatia fascicularis J.R.Forst. & G.Forst. (AR, CH), Phyllachne uliginosa J.R. Forst. & G. Forst. (AR, CH).

CAMPANULACEAE ‣ Lysipomia aretioides Kunth (EC, PR), L. muscoides Hook.f. (CL, EC, PR).

Lysipomia muscoides Hook.f.

CAPRIFOLIACEAE ‣ Valeriana alypifolia Kunth (EC), V. aretioides Kunth (EC, PR, BL), V. corynodes Borsini (AR), V. densa (Wedd.) Höck (BL), V. macrorhiza Poepp. ex DC. (BL, AR, CH), V. rigida Ruiz & Pav. (CL, EC, PR, BL), V. sedifolia d'Urv. (AR, CH, inc. Aretiastrum magellanicum (Decne.) Skottsb.).

Valeriana rigida Ruiz & Pav.

APIACEAE ‣ Azorella albovaginata (Gillies & Hook.) G.M.Plunkett & A.N.Nicolas (AR, CH), A. ameghinoi Speg. (AR, CH), A. aretioides (Kunth) Willd. ex DC. (CL, EC, PR, AR), A. biloba (Schltdl.) Wedd. (CL, EC, PR, BL, AR, CH), A. corymbosa (Ruiz & Pav.) Pers. (CL, EC, PR), A. compacta Phil. (PR, BL, AR, CH), A. crassipes Phil. (AR, CH), A. crenata (Ruiz & Pav.) Pers. (VZ, CL, EC, PR), A. diapensioides A.Gray (PR, BL, AR, CH), A. diversifolia Clos (AR, CH), A. filamentosa Lam. (AR, CH), A. fuegiana Speg. (AR, CH), A. hallei (Skottsb.) G.M.Plunkett & A.N.Nicolas (AR), A. julianii Mathias & Constance (VZ, CL), A. lycopodioides Gaudich. (AR, CH), A. madreporica Clos (AR, CH), A. microphylla (Cav.) G.M.Plunkett & A.N.Nicolas (AR), A. monantha Clos (AR, CH), A. multifida (Ruiz & Pav.) Pers. (VZ, CL, EC, PR, BL), A. nivalis Phil. (AR, CH), A. patagonica Speg. (AR, CH), A. pectinata Phil. (CH), A. pedunculata (Spreng.) Mathias & Constance (CL, EC), A. pulvinata Wedd. (PR, BL), A. prolifera (Cav.) G.M.Plunkett & A.N.Nicolas (AR, CH), A. ruizii G.M.Plunkett & A.N.Nicolas (AR, CH), A. selago Hook.f. (AR, CH), A. spinosa (Ruiz & Pav.) Pers. (AR, CH), A. trifoliolata Clos (AR, CH), A. trifurcata (Gaertn.) Pers. (AR, CH), A. trisecta (H.Wolff) Mart.Fernández & C.I.Calviño (BL, AR, CH; syn. Mulinum crassifolium), A. valentini (Speg.) Mart.Fernández & C.I.Calviño (AR, CH), Bolax caespitosa Hombr. & Jacquinot ex Decne (AR, CH), B. gummifera (Lam.) Spreng. (AR, CH).

Azorella compacta Phil., A. diapensioides A.Gray