1. GLOBAL ALPINES ALPINE GARDEN SOCIETY East Anglian Conference 23 April 2005

2. ALPINE PLANTS OF THE AMERICAS John and Hilary Birks

3. The Americas A confession! State of floristic knowledge Environmental features Vegetation zonation Tree-line in the Americas Extent of the alpine-zone in the Americas Richness of alpines General comparisons of alpines in the Americas Alpine species richness patterns Species in common Genera in common Families in common Primarily North American families Primarily South American families Possible floristic history Threats to survival and conservation Final comments

4. A Confession! 'The Americas' is a very misleading title Western North America- 10 visits Patagonia- 2 visits More correct title: 'Alpine Plants of Western North America and Patagonia' Alpine plants difficult to define here because in some areas it is too dry for trees to grow. Steppe zone merges into alpine zone. No clear distinction between an 'alpine' and a 'steppe' plant.

6. State of Floristic Knowledge North America – mainly state floras of variable quality

7. Flora of North America - Started in 1993 8 volumes of a projected 30 volumes ca. 17 000 species, 1 200 genera, 220 families

8. Flora Patagonica1969-19998 volumes ca. 2 400 species, 738 genera, 138 families

9. We will use the only source of consistent nomenclature for North and South American alpines, namely Alpine Garden Society Encyclopaedia of Alpines edited by Kenneth Beckett

10. Left projectorRight projector Klutlan Glacier & St Elias Mountains, Yukon Moraine Lake, Alberta Hellroaring Plateau, MontanaBeartooth Plateau, Montana & Wyoming Big Horns, WyomingGrand Tetons, Wyoming Mount Bierstone, ColoradoPikes Peak, Colorado Borah Peak, IdahoWhite Cloud Peaks, Idaho Cecret Lake & Albion Basin, UtahReid's Peak, Utah Steens Mountain, OregonCrater Lake, Oregon Three Sisters, OregonMt Shasta from Mt Eddy, California Torres del Paine, ChileLago Grey, Chile Cerro Moyano, ArgentinaMount Fitzroy, Argentina Lago Belgrano, ArgentinaLago Nahuel Huapi, Argentina Cerro Chapelco, ArgentinaMoquehue, Argentina Batea Mahuida, ArgentinaVolcan Copahue, Argentina

11. Environmental Features Low temperatures, long-lasting or permanent snow, glaciers Steep rocky slopes. Screes, cliffs, exposed ridges, and summits Temperature decreases, on average, 0.65ºC per 100 m altitude High winds 'Roaring Forties' and 'Furious Fifties' in South America "In few parts of the world is the climate of a region and its life so determined by a single meteorological element, as is the climate of Patagonia by the constancy and strength of the wind." Coronato (1993) Wind chill reduces mean annual temperature perception by 4.2ºC Strong west to east gradients in annual rainfall over small distances (200-300 km) 7 m to 200 mm from western Chile to Puerto Santa Cruz 3 m to 200 mm from coastal Oregon to Steens Mountain

12. snowbeds 1. Central Patagonia – 41ºS – latitude of Bariloche 2500 2000 1500 1000 500 0 3500 2500 15001000 500 steppe open xeric woodland Annual precipitation (mm) Altitude (m)  Temperature WE temperate forest temperate rainforest species- poor steppe deciduous Nothofagus forest evergreen Nothofagus pumilo krummholzAraucaria woodland alpine grasslandalpine semi-desert permanent snow and ice NIVAL SUB-NIVAL ALPINE SUB- ALPINE species-poor temperate rainforest Vegetation Zonation - (Hypothetical!)

13. snowbeds 2. Western North America – 41ºN – latitude of Salt Lake City 4000 3500 3000 2500 2000 1500 1000 500 0 3000 2000 15001000 200 desert prairie open woodland /savannah Annual precipitation (mm) Altitude (m)  Temperature WE temperate forest (Quercus) temperate rainforest Pinus woodland steppe mixed coniferous forest Tsuga-Thuja-Picea- Abies-Sequoia- Sequoiadendron Picea-Abies-Pseudotsuga forest Populus tremuloides Abies lasiocarpa Picea engelmannii Pinus albicaulis Juniperus spp. Pinus longaeva wet meadowsdry grasslandopen fell-field permanent snow and ice NIVAL SUB-NIVAL ALPINE SUB- ALPINE MONTANE FOOTHILLS PLAINS

14. Tree-Line in the Americas

15. North AmericaSouth America Tree typeConifers (one deciduous tree) Deciduous trees (one conifer) Main species Picea engelmannii, Abies lasiocarpa, Pinus albicaulis, P. aristata, P. longaeva, Juniperus occidentalis, (Populus tremuloides) Nothofagus pumilio, N. antarctica (Araucaria araucana) Altitude at55º1500 m50 m 50º2300 m800 m 45º3000 m1200 m 40º3500 m1800 m Rate of tree-line decrease per 1º latitude 130 m120 m

16. Left projectorRight projector Tree line, Bald Mountain, Utah Picea engelmannii, Wyoming Nothofagus pumilio, ChileNothofagus pumilio, Argentina

17. Extent of the Alpine Zone in the Americas Extent of alpine zone (tree-line to snow-line) o snow-line tree-line Alpine zone NS

18. Richness of Alpines Global10 000 – 15 000 species(6% world's flora) Europe2 500 species(20% Europe's flora) N America1 200 – 1 500 species(9% N American flora) S America? 1 000 species(?% S American flora)

19. General Comparisons of Alpines in the Americas N AmericaS America Low-growing creeping or prostrate shrubs +++ Graminoids (grasses, sedges, etc.) forming tussocks +++ Herbaceous perennials, usually with rosettes +++ Cushion plants +++ Spiny cushions ('vegetable hedgehogs') -+ Giant rosettes -(+) Succulents ++ Geophytes (bulbs, tubers) ++ Annuals (+) ++ = many+ = some(+) = few- = none

20. N AmericaS America Species richness on different mountains HighMedium Generic richnessMediumHigh Endemic richnessLowHigh Range sizeLargeSmall Genetic diversityHighLow

21. Alpine Species Richness Patterns N AmericaS America Mountain areas studied2610 Latitude range68 – 36ºN54 – 27ºS Species richness132-278103-200 Median richness (40-54º latitude) 195144 Difference in richness in 1º areas (40-54º) 35 -73

22. Changes in Alpine Richness with Increasing Latitude and Altitude N America ~ 5 species decrease per 1º latitude increase S America ~ 3 species decrease per 1º latitude increase If we allow for differences in total richness, then percentage decrease in alpine richness with increasing latitude is about the same, 1.7% in North, 1.5% in South America. Number of endemics decreases with increasing latitude in N America but not in S America – ? differences in history. Also a decrease in species richness with increasing altitude N ~30 species decrease per 100 m altitude S ~18 species decrease per 100 m altitude

23. SPECIES IN COMMON (15) Anemone multifida Gentiana prostrata Koenigia islandica Cerastium arvense Phleum alpinum + 4 other grasses 6 species of Carex

24. Left projectorRight Projector Anemone multifida, ChileAnemone multifida, California Gentiana prostrata, ColoradoKoenigia islandica, Bhutan Cerastium arvense, ArgentinaPhleum alpinum, Colorado

25. GENERA IN COMMON (29) EphedraClarkiaCaltha PhaceliaRanunculusEmpetrum SaxifragaDrabaPrimula ArmeriaOenotheraMimulus SileneArenariaPlantago Valeriana ThlaspiRibesTaraxacum RubusFragariaEpilobium Hypochaeris AsterSenecioAstragalus HaplopappusGaliumSisyrinchium

26. Ephedra frustillata, ArgentinaClarkia pulchella, Idaho Caltha leptosepala, ColoradoCaltha sagittata, Argentina Phacelia sericea, ColoradoRanunculus adoneus, Colorado Ranunculus semiverticillatus, ArgentinaRanunculus semiverticillatus, Arg Empetrum nigrum, NorwayEmpetrum rubrum, Argentina Saxifraga oppositifolia, NorwaySaxifraga caespitosa, Norway Saxifraga rhomboidea, ColoradoSaxifraga flagellaris, Colorado Saxifraga magellanica, ArgentinaDraba fladnizensis, Colorado Draba incerta, WyomingDraba gilliesii, Argentina Primula angustifolia, ColoradoPrimula magellanica, Argentina Armeria scabra var. sibirica, ColoradoArmeria maritima ssp. andina, Arg Oenothera caespitosa, IdahoOenothera odorata, Argentina Mimulus tilingii, UtahMimulus cusickii, Oregon Mimulus lewisii, WyomingMimulus glabratus, Argentina Mimulus cupreus, ArgentinaSilene acaulis, Colorado Silene scouleri ssp. hallii, ColoradoSilene magellanica, Argentina Arenaria obtusiloba, ColoradoArenaria serpens, Chile Plantago canescens, AlaskaPlantago sempervivoides, Argentina Valeriana philippiana, ArgentinaValeriana moyanoi, Argentina

27. FAMILIES IN COMMON (16) RanunculaceaeAquilegia, Pulsatilla, Clematis Hamadryas RosaceaeDryas, Potentilla, Kelsia, Petrophyton Acaena Cruciferae (Brassicaceae) Physaria, Smelowskia, Lesquerella Xerodraba, Stenodraba, Onuris, Eudema, Menonvillea Leguminosae (Fabaceae) Lupinus, Trifolium, Oxytropis, Astragalus Astragalus, Adesmia, Anarthrophyllum PortulacaceaeLewisia, Claytonia, Calyptridium Calandrinia

28. Aquilegia caerulea, ColoradoAquilegia jonesii, Wyoming Pulsatilla occidentalis, OregonPulsatilla patens, Wyoming Clematis columbiana var. tenuiloba, WyomingClematis hirsutissima, Wyoming Hamadryas kingii (male), ArgentinaHamadryas delfinii (male), Argentina Hamadryas sempervivoides, ArgentinaHamadryas sempervivoides (male), Argentina Dryas octopetala, SloveniaDryas drummondii, Yukon Potentilla fruticosa, ColoradoKelseya uniflora, Idaho Petrophyton caespitosum, IdahoAcaena magellanica, Chile Physaria alpina, Colorado Smelowskia calycina, ColoradoLesquerella alpina, Wyoming Xerodraba patagonica, ArgentinaStenodraba pusilla, Argentina Onuris graminifolia, ArgentinaEudema monantha, Argentina Menonvillea nordenskjoeldii, ArgentinaMenonvillea rigida, Argentina Lupinus breweri, OregonTrifolium nanum, Colorado Oxytropis nana, WyomingOxytropis lambertii, Wyoming Astragalus kentrophytus, MontanaAstragalus utahensis, Idaho Astragalus whitneyi var. siskyouensis, CaliforniaAstragalus cruickshankii, Argentina Adesmia corymbosa, ArgentinaAdesmia longipes, Argentina Adesmia salicornioides, ArgentinaAdesmia parviflora, Argentina Adesmia villosa, ArgentinaAdesmia boronoides, Argentina Anarthrophyllum desideratum, Chile Lewisia rediviva, Idaho Claytonia megarhiza, ColoradoCalyptridium umbellatum, Oregon Calandrinia affinis agg., Argentina

29. RubiaceaeGalium Oreopolus CampanulaceaeCampanula Pratia CactaceaeMamillaria, Opuntia Maihuenia, Maihueniopsis, Austrocactus, Gymnocalcium Umbelliferae (Apiaceae) Oreoxis, Angelica Mulinum, Azorella, Bolax, Pozoa EricaceaeCassiope, Phyllodoce, Rhododendron, Ledum Pernettya

30. Left projectorRight projector Galium boreale, AlaskaOreopolus glacialis, Argentina Campanula rotundifolia, ColoradoCampanula lasiocarpa, Yukon Campanula shetleri, CaliforniaPratia repens, Argentina Mamillaria vivipara, N DakotaOpuntia polycantha, Idaho Maihuenia patagonica, ArgentinaMaihueniopsis platycantha, Arg Austrocactus patagonicus, ArgGymnocalcium gibbosum, Argentina Oreoxis humilis, ColoradoAngelica grayi, Colorado Mulinum spinosum, Argentina Azorella monantha, ArgentinaAzorella patagonica, Argentina Bolax gummifer, ArgentinaPozoa vulcanica, Argentina Cassiope mertensiana, AlaskaPhyllodoce empetriformis, Oregon Rhododendron occidentale, CalLedum groenlandicum, Yukon Pernytta mucronata, ArgentinaPernytta pumila, Argentina

31. Compositae (Asteraceae) Aster, Erigeron, Cirsium, Senecio, Ligularia, Hymenoxis, Raillardella, Hulsea, Chaenactis, Crepis, Saussurea, Townsendia, Wyethia Nassauvia, Leuchera, Haplopappus, Perezia, Nardophyllum, Brachyclados, Senecio ScrophulariaceaePenstemon, Castilleja, Besseya, Pedicularis Calceolaria, Ourisia, Euphrasia Labiatae (Lamiaceae) Monardella Satureja

32. Aster coloradoensis, ColoradoErigeron compositus, Colorado Cirsium scopulorum, ColoradoSenecio canus, California Ligularia holmii, ColoradoHymenoxis acaulis var. caespitosa, Colorado Hymenoxis grandiflora, ColoradoRaillardella argentea, Oregon Hulsea nana, CaliforniaChaenactis alpina, Colorado Crepis nana, YukonSaussurea weberi, Colorado Townsendia leptotes, WyomingWyethia helianthoides, Idaho Nassauvia glomerulosa, ArgentinaNassauvia juniperina, Argentina Nassauvia lagascae, ArgentinaNassauvia lagascae var. globosa, Argentina Nassauvia pygmaea, ArgentinaNassauvia magellanica, Argentina Nassauvia axillaris, ArgentinaLeuchera purpurea, Chile Haplopappus prunelloides, ArgentinaPerezia recurvata, Argentina Nardophyllum bryoides, ArgentinaBrachyclados caespitosus, Argentina Senecio miser, ArgentinaSenecio tricuspidatus, Argentina Penstemon rupicola, OregonPenstemon davidsonii, Oregon Penstemon hallii, ColoradoPenstemon eriantherus, Wyoming Penstemon uintahuensis, UtahCastilleja miniata, Oregon Castilleja rhexifolia, ColoradoCastilleja puberula, Colorado Besseya alpina, ColoradoBesseya ritteriana, Colorado Pedicularis kanei, AlaskaPedicularis groenlandica, Colorado Calceolaria uniflora, ArgentinaCalceolaria polyrhiza, Chile Calceolaria biflora, ArgentinaCalceolaria crenatifolia, Argentina Ourisia poeppigii, ChileOurisia poeppigii, Argentina Ourisia fragrans, ArgentinaEuphrasia meiantha, Argentina Monardella odoratissima, UtahSatureja darwinii, Argentina

33. LiliaceaeCalochortus, Fritillaria, Allium, Erythronium, Veratrum, Xerophyllum Tristagma IridiceaeIris Olsynium, Sisyrinchium OrchidaceaeSpiranthes, Habenaria, Cypripedium Gavilea, Chloraea

34. Left projectorRight projector Calochortus gunnisonii, ColoradoFritillaria pudica, Wyoming Allium acuminatum, OregonErythronium grandiflorum, Wyoming Veratrum californicum, UtahXerophyllum tenax, Alberta Tristagma nivale, ArgentinaTristagma sessile, Argentina Tristagma patagonicum, ArgentinaIris missourensis, Wyoming Olsynium junceum, Argentina Olsynium biflorum, ArgentinaOlsynium biflorum agg., Argentina Olsynium frigidum, ArgentinaSisyrinchium macrocarpum, Argentina Spiranthes romanzoffiana, AlbertaHabenaria dilatata, Alberta Cypripedium californicum, CaliforniaGavilea lutea, Chile Chloraea alpina, ArgentinaChloraea magellanica, Argentina

35. PRIMARILY NORTH AMERICAN FAMILIES (10) Salicaceae-Salix Polemoniaceae-Linanthus, Polemonium, Phlox, Gilia, Ipomopsis, Collomia Gentianaceae-Gentiana, Swertia, Frasera Polygonaceae-Eriogonum, Polygonum Crassulaceae-Sedum, Clementsia = Rhodiola Boraginaceae-Eritrichium Primulaceae-Douglasia, Dodecatheon, Androsace Papaveraceae-Papaver Linaceae-Linum Pinaceae-Pinus longaeva, P. aristata

36. Salix reticulata ssp. nivalis, Colorado Linanthus (= Linanthastrum) nuttalii, Oregon Polemonium viscosum, WyomingPolemonium confertum, Montana Phlox diffusa, OregonPhlox multiflora, Idaho Gilia aggregata, OregonIpomopsis globularis, Colorado Collomia debilis, UtahCollomia larsenii, Oregon Gentiana detonsa, WyomingGentiana algida, Colorado Swertia perennis, ColoradoFrasera speciosa, Wyoming Eriogonum ovalifolium, OregonEriogonum cespitosum, California Eriogonum umbellatum var. porteri, Colorado Polygonum bistortoides, Colorado Sedum divergens, OregonClementsia rhodantha, Colorado Eritrichium aretioides, WyomingEritrichium howardii, Wyoming Douglasia montana, WyomingDodecatheon pulchellum, Wyoming Androsace chaemasjne ssp. lehmanniana, Alberta Papaver kluanense, Colorado Linum kingii, UtahPinus longaeva, California

37. PRIMARILY SOUTH AMERICAN FAMILIES (12) Solanaceae-Benthamiella, Petunia, Fabiana, Jaborosa Verbenaceae-Junellia, Glandularia, Acantholippia Oxalidaceae-Oxalis Tropaeolaceae-Tropaeolum Bignoniaceae-Argylia Alstroemeriaceae-Alstroemeria Amaryllidaceae-Rhodophiala Calyceraceae-Moschopsis, Nastanthus Proteaceae-Embothrium Santalaceae-Quinchamalium Violaceae-Rosulate Viola Araucariaceae-Araucaria araucana

38. Benthamiella longifolia, ArgentinaBenthamiella nordenskjoldii, Argentina Benthamiella patagonica, ArgentinaBenthamiella azorella, Argentina Petunia patagonica, Argentina Fabiana nana, ArgentinaJaborosa volkmanii, Argentina Junellia azorelloides, Argentina Junellia mulinoides, ArgentinaJunellia minutifolia, Argentina Junellia micrantha, ArgentinaJunellia patagonica, Argentina Glandularia micrantha, ArgentinaAcantholippia seriphioides, Argentina Oxalis enneaphylla, Argentina Oxalis erythrorhiza, ArgentinaOxalis nahuelhuapiensis, Argentina Oxalis laciniata, Argentina Oxalis adenophylla, Argentina Oxalis loricata, Argentina Tropaeolum incisum, ArgentinaArgylia bustillosi, Argentina Rhodophiala andicola, ArgentinaRhodophiala mendocina, Argentina Moschopsis rosulata, ArgentinaMoschopsis subandina, Argentina Nastanthus spathulatus, ArgentinaAlstroemeria patagonica, Argentina Embothrium coccineum, ChileEmbothrium coccineum, Argentina Quinchamalium chilense, ArgentinaViola reichii, Chile Viola sacculus, Argentina Viola cotyledon, Argentina Viola columnaris, ArgentinaViola ? petraea, Argentina Viola dasyphylla, ArgentinaViola coronifera, Argentina Viola auricolor, Argentina Viola vulcanica, ArgentinaViola sacculus, Argentina Araucaria araucanaTorres del Paine, Chile

39. POSSIBLE FLORISTIC HISTORY 1.Little direct fossil evidence – very few studies 2.Higher frequency of endemics in South America 3.Species ranges and genetic diversity generally smaller in S America Circum- arctic plants Beringia plants Basin plants Ice-free mountain areas Andean plants Steppe plants Circum-antarctic plants Ice-sheets in North Ice-sheets in West and South

40. THREATS TO SURVIVAL AND CONSERVATION Live in a period of rapid environmental change – climate, land-use, atmospheric nitrogen, plant introductions, mountain developments. Provide potential threats to the survival of alpines. N AmericaS America Introduced species+++ Land-use changes-+ Hydroelectric development+- Ski development++ Atmospheric nitrogen deposition++- Global warming+++ ++ = high threat+ = some threat- = no likely threat

41. FINAL COMMENTS 1.Both continents have wonderful, attractive, interesting, and very different floras. 2.Striking differences between the two continents in terms of tree-line and extent of alpine zone, probably due to the major differences in wind severity. 3.Many areas remain to be explored and documented. Great amount to be discovered in both North and South America. 4.Much remains to be brought in cultivation. 5.American alpines, like alpines elsewhere, are threatened by impacts of 'global change', especially global warming, atmospheric nitrogen deposition, and loss of habitat.

42. Acknowledgements Peter ErskinePhil & Gwen Phillips Marcela FerreyraLoren Russell Martin & Anna SheaderLoraine Yeatts Cathy Jenks