GLOBAL ALPINES

ALPINE GARDEN SOCIETY

East Anglian Conference

23 April 2005

ALPINE PLANTS OF THE AMERICAS

John and Hilary Birks

The Americas

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

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.

State of Floristic Knowledge

North America – mainly state floras of variable quality

Flora of North America - Started in 1993

8 volumes of a projected 30 volumes

ca. 17 000 species, 1 200 genera, 220 families

Flora Patagonica 1969-1999 8 volumes

ca. 2 400 species, 738 genera, 138 families

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

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Klutlan Glacier & St Elias Mountains, Yukon

Moraine Lake, Alberta

Hellroaring Plateau, Montana Beartooth Plateau, Montana & Wyoming

Big Horns, Wyoming Grand Tetons, Wyoming

Mount Bierstone, Colorado Pikes Peak, Colorado

Borah Peak, Idaho White Cloud Peaks, Idaho

Cecret Lake & Albion Basin, Utah Reid's Peak, Utah

Steens Mountain, Oregon Crater Lake, Oregon

Three Sisters, Oregon Mt Shasta from Mt Eddy, California

Torres del Paine, Chile Lago Grey, Chile

Cerro Moyano, Argentina Mount Fitzroy, Argentina

Lago Belgrano, Argentina Lago Nahuel Huapi, Argentina

Cerro Chapelco, Argentina Moquehue, Argentina

Batea Mahuida, Argentina Volcan Copahue, Argentina

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

snowbeds

1. Central Patagonia – 41ºS – latitude of Bariloche

2500

2000

1500

1000

500

0 3500 2500 1500 1000 500

steppeopen xeric woodland

Annual precipitation (mm)

Alt

itud

e (

m)

Tem

pera

ture

W E

temperate foresttemperate

rainforest

species-poor

steppe

deciduous Nothofagus forest

evergreen Nothofagus pumilo krummholz

Araucaria woodland

alpine grassland alpine semi-desert

permanent snow and ice NIVAL

SUB-NIVAL

ALPINESUB-

ALPINE

species-poor

temperate rainforest

Vegetation Zonation - (Hypothetical!)

snowbeds

2. Western North America – 41ºN – latitude of Salt Lake City

4000

3500

3000

2500

2000

1500

1000

500

03000 2000 1500 1000 200

desert

prairieopen

woodland /savannah

Annual precipitation (mm)

Alt

itud

e (

m)

Tem

pera

ture

W E

temperate forest

(Quercus)

temperate rainforest

Pinus woodland steppe

mixed coniferous forest Tsuga-Thuja-

Picea-Abies-Sequoia-Sequoiadendron

Picea-Abies-Pseudotsuga forest

Populus tremuloide

s

Abies lasiocarp

a

Picea engelmann

ii

Pinus albicauli

s

Juniperus spp.

Pinus longaev

a

wet meadows dry grassland open fell-field

permanent snow and ice NIVAL

SUB-NIVAL

ALPINE

SUB-ALPIN

E

MONTANE

FOOTHILLS

PLAINS

Tree-Line in the Americas

North America South America

Tree type Conifers (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 at

55º

1500 m 50 m

50º

2300 m 800 m

45º

3000 m 1200 m

40º

3500 m 1800 m

Rate of tree-line decrease per 1º latitude

130 m 120 m

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Tree line, Bald Mountain, Utah

Picea engelmannii, Wyoming

Nothofagus pumilio, Chile Nothofagus pumilio, Argentina

Extent of the Alpine Zone in the Americas

Extent of alpine zone (tree-line to snow-line)

o snow-line

• tree-line

Alpine zone

N S

Richness of Alpines

Global 10 000 – 15 000 species (6% world's flora)

Europe 2 500 species (20% Europe's flora)

N America 1 200 – 1 500 species (9% N American flora)

S America ? 1 000 species (?% S American flora)

General Comparisons of Alpines in the Americas

N America

S 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

N America

S America

Species richness on different mountains

High Medium

Generic richness Medium High

Endemic richness Low High

Range size Large Small

Genetic diversity High Low

Alpine Species Richness Patterns

N America S America

Mountain areas studied

26 10

Latitude range 68 – 36ºN 54 – 27ºS

Species richness 132-278 103-200

Median richness (40-54º latitude)

195 144

Difference in richness in 1º areas (40-54º)

35 - 73

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

SPECIES IN COMMON (15)

Anemone multifida

Gentiana prostrata

Koenigia islandica

Cerastium arvense

Phleum alpinum + 4 other grasses

6 species of Carex

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Anemone multifida, Chile Anemone multifida, California

Gentiana prostrata, Colorado Koenigia islandica, Bhutan

Cerastium arvense, Argentina Phleum alpinum, Colorado

GENERA IN COMMON (29)Ephedra Clarkia Caltha

Phacelia Ranunculus Empetrum

Saxifraga Draba Primula

Armeria Oenothera Mimulus

Silene Arenaria Plantago

Valeriana

Thlaspi Ribes Taraxacum

Rubus Fragaria Epilobium

Hypochaeris

Aster Senecio Astragalus

Haplopappus Galium Sisyrinchium

Ephedra frustillata, Argentina Clarkia pulchella, Idaho

Caltha leptosepala, Colorado Caltha sagittata, Argentina

Phacelia sericea, Colorado Ranunculus adoneus, Colorado

Ranunculus semiverticillatus, Argentina Ranunculus semiverticillatus, Arg

Empetrum nigrum, Norway Empetrum rubrum, Argentina

Saxifraga oppositifolia, Norway Saxifraga caespitosa, Norway

Saxifraga rhomboidea, Colorado Saxifraga flagellaris, Colorado

Saxifraga magellanica, Argentina Draba fladnizensis, Colorado

Draba incerta, Wyoming Draba gilliesii, Argentina

Primula angustifolia, Colorado Primula magellanica, Argentina

Armeria scabra var. sibirica, Colorado Armeria maritima ssp. andina, Arg

Oenothera caespitosa, Idaho Oenothera odorata, Argentina

Mimulus tilingii, Utah Mimulus cusickii, Oregon

Mimulus lewisii, Wyoming Mimulus glabratus, Argentina

Mimulus cupreus, Argentina Silene acaulis, Colorado

Silene scouleri ssp. hallii, Colorado Silene magellanica, Argentina

Arenaria obtusiloba, Colorado Arenaria serpens, Chile

Plantago canescens, Alaska Plantago sempervivoides, Argentina

Valeriana philippiana, Argentina Valeriana moyanoi, Argentina

FAMILIES IN COMMON (16)

Ranunculaceae

Aquilegia, Pulsatilla, ClematisHamadryas

Rosaceae Dryas, Potentilla, Kelsia, PetrophytonAcaena

Cruciferae(Brassicaceae)

Physaria, Smelowskia, LesquerellaXerodraba, Stenodraba, Onuris, Eudema, Menonvillea

Leguminosae(Fabaceae)

Lupinus, Trifolium, Oxytropis, AstragalusAstragalus, Adesmia, Anarthrophyllum

Portulacaceae Lewisia, Claytonia, CalyptridiumCalandrinia

Aquilegia caerulea, Colorado Aquilegia jonesii, Wyoming

Pulsatilla occidentalis, Oregon Pulsatilla patens, Wyoming

Clematis columbiana var. tenuiloba, Wyoming Clematis hirsutissima, Wyoming

Hamadryas kingii (male), Argentina Hamadryas delfinii (male), Argentina

Hamadryas sempervivoides, Argentina Hamadryas sempervivoides (male), Argentina

Dryas octopetala, Slovenia Dryas drummondii, Yukon

Potentilla fruticosa, Colorado Kelseya uniflora, Idaho

Petrophyton caespitosum, Idaho Acaena magellanica, Chile

Physaria alpina, Colorado Physaria alpina, Colorado

Smelowskia calycina, Colorado Lesquerella alpina, Wyoming

Xerodraba patagonica, Argentina Stenodraba pusilla, Argentina

Onuris graminifolia, Argentina Eudema monantha, Argentina

Menonvillea nordenskjoeldii, Argentina Menonvillea rigida, Argentina

Lupinus breweri, Oregon Trifolium nanum, Colorado

Oxytropis nana, Wyoming Oxytropis lambertii, Wyoming

Astragalus kentrophytus, Montana Astragalus utahensis, Idaho

Astragalus whitneyi var. siskyouensis, California Astragalus cruickshankii, Argentina

Adesmia corymbosa, Argentina Adesmia longipes, Argentina

Adesmia salicornioides, Argentina Adesmia parviflora, Argentina

Adesmia villosa, Argentina Adesmia boronoides, Argentina

Anarthrophyllum desideratum, Chile Anarthrophyllum desideratum, Chile

Lewisia rediviva, Idaho Lewisia rediviva, Idaho

Claytonia megarhiza, Colorado Calyptridium umbellatum, Oregon

Calandrinia affinis agg., Argentina Calandrinia affinis agg., Argentina

Rubiaceae GaliumOreopolus

Campanulaceae

CampanulaPratia

Cactaceae Mamillaria, OpuntiaMaihuenia, Maihueniopsis, Austrocactus, Gymnocalcium

Umbelliferae(Apiaceae)

Oreoxis, AngelicaMulinum, Azorella, Bolax, Pozoa

Ericaceae Cassiope, Phyllodoce, Rhododendron, LedumPernettya

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Galium boreale, Alaska Oreopolus glacialis, Argentina

Campanula rotundifolia, Colorado Campanula lasiocarpa, Yukon

Campanula shetleri, California Pratia repens, Argentina

Mamillaria vivipara, N Dakota Opuntia polycantha, Idaho

Maihuenia patagonica, Argentina Maihueniopsis platycantha, Arg

Austrocactus patagonicus, Arg Gymnocalcium gibbosum, Argentina

Oreoxis humilis, Colorado Angelica grayi, Colorado

Mulinum spinosum, Argentina Mulinum spinosum, Argentina

Azorella monantha, Argentina Azorella patagonica, Argentina

Bolax gummifer, Argentina Pozoa vulcanica, Argentina

Cassiope mertensiana, Alaska Phyllodoce empetriformis, Oregon

Rhododendron occidentale, Cal Ledum groenlandicum, Yukon

Pernytta mucronata, Argentina Pernytta pumila, Argentina

Compositae (Asteraceae)

Aster, Erigeron, Cirsium, Senecio, Ligularia, Hymenoxis, Raillardella, Hulsea, Chaenactis, Crepis, Saussurea, Townsendia, Wyethia

Nassauvia, Leuchera, Haplopappus, Perezia, Nardophyllum, Brachyclados, Senecio

Scrophulariaceae

Penstemon, Castilleja, Besseya, Pedicularis

Calceolaria, Ourisia, Euphrasia

Labiatae(Lamiaceae)

Monardella

Satureja

Aster coloradoensis, Colorado Erigeron compositus, Colorado

Cirsium scopulorum, Colorado Senecio canus, California

Ligularia holmii, Colorado Hymenoxis acaulis var. caespitosa, Colorado

Hymenoxis grandiflora, Colorado Raillardella argentea, Oregon

Hulsea nana, California Chaenactis alpina, Colorado

Crepis nana, Yukon Saussurea weberi, Colorado

Townsendia leptotes, Wyoming Wyethia helianthoides, Idaho

Nassauvia glomerulosa, Argentina Nassauvia juniperina, Argentina

Nassauvia lagascae, Argentina Nassauvia lagascae var. globosa, Argentina

Nassauvia pygmaea, Argentina Nassauvia magellanica, Argentina

Nassauvia axillaris, Argentina Leuchera purpurea, Chile

Haplopappus prunelloides, Argentina Perezia recurvata, Argentina

Nardophyllum bryoides, Argentina Brachyclados caespitosus, Argentina

Senecio miser, Argentina Senecio tricuspidatus, Argentina

Penstemon rupicola, Oregon Penstemon davidsonii, Oregon

Penstemon hallii, Colorado Penstemon eriantherus, Wyoming

Penstemon uintahuensis, Utah Castilleja miniata, Oregon

Castilleja rhexifolia, Colorado Castilleja puberula, Colorado

Besseya alpina, Colorado Besseya ritteriana, Colorado

Pedicularis kanei, Alaska Pedicularis groenlandica, Colorado

Calceolaria uniflora, Argentina Calceolaria polyrhiza, Chile

Calceolaria biflora, Argentina Calceolaria crenatifolia, Argentina

Ourisia poeppigii, Chile Ourisia poeppigii, Argentina

Ourisia fragrans, Argentina Euphrasia meiantha, Argentina

Monardella odoratissima, Utah Satureja darwinii, Argentina

Liliaceae Calochortus, Fritillaria, Allium, Erythronium, Veratrum, Xerophyllum

Tristagma

Iridiceae Iris

Olsynium, Sisyrinchium

Orchidaceae Spiranthes, Habenaria, Cypripedium

Gavilea, Chloraea

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Calochortus gunnisonii, Colorado Fritillaria pudica, Wyoming

Allium acuminatum, Oregon Erythronium grandiflorum, Wyoming

Veratrum californicum, Utah Xerophyllum tenax, Alberta

Tristagma nivale, Argentina Tristagma sessile, Argentina

Tristagma patagonicum, Argentina Iris missourensis, Wyoming

Olsynium junceum, Argentina Olsynium junceum, Argentina

Olsynium biflorum, Argentina Olsynium biflorum agg., Argentina

Olsynium frigidum, Argentina Sisyrinchium macrocarpum, Argentina

Spiranthes romanzoffiana, Alberta Habenaria dilatata, Alberta

Cypripedium californicum, California Gavilea lutea, Chile

Chloraea alpina, Argentina Chloraea magellanica, Argentina

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

Salix reticulata ssp. nivalis, Colorado

Linanthus (= Linanthastrum) nuttalii, Oregon

Polemonium viscosum, Wyoming Polemonium confertum, Montana

Phlox diffusa, Oregon Phlox multiflora, Idaho

Gilia aggregata, Oregon Ipomopsis globularis, Colorado

Collomia debilis, Utah Collomia larsenii, Oregon

Gentiana detonsa, Wyoming Gentiana algida, Colorado

Swertia perennis, Colorado Frasera speciosa, Wyoming

Eriogonum ovalifolium, Oregon Eriogonum cespitosum, California

Eriogonum umbellatum var. porteri, Colorado

Polygonum bistortoides, Colorado

Sedum divergens, Oregon Clementsia rhodantha, Colorado

Eritrichium aretioides, Wyoming Eritrichium howardii, Wyoming

Douglasia montana, Wyoming Dodecatheon pulchellum, Wyoming

Androsace chaemasjne ssp. lehmanniana, Alberta

Papaver kluanense, Colorado

Linum kingii, Utah Pinus longaeva, California

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

Benthamiella longifolia, Argentina Benthamiella nordenskjoldii, Argentina

Benthamiella patagonica, Argentina Benthamiella azorella, Argentina

Petunia patagonica, Argentina Petunia patagonica, Argentina

Fabiana nana, Argentina Jaborosa volkmanii, Argentina

Junellia azorelloides, Argentina Junellia azorelloides, Argentina

Junellia mulinoides, Argentina Junellia minutifolia, Argentina

Junellia micrantha, Argentina Junellia patagonica, Argentina

Glandularia micrantha, Argentina Acantholippia seriphioides, Argentina

Oxalis enneaphylla, Argentina Oxalis enneaphylla, Argentina

Oxalis erythrorhiza, Argentina Oxalis nahuelhuapiensis, Argentina

Oxalis laciniata, Argentina Oxalis laciniata, Argentina

Oxalis adenophylla, Argentina Oxalis adenophylla, Argentina

Oxalis adenophylla, Argentina Oxalis loricata, Argentina

Tropaeolum incisum, Argentina Argylia bustillosi, Argentina

Rhodophiala andicola, Argentina Rhodophiala mendocina, Argentina

Moschopsis rosulata, Argentina Moschopsis subandina, Argentina

Nastanthus spathulatus, Argentina Alstroemeria patagonica, Argentina

Embothrium coccineum, Chile Embothrium coccineum, Argentina

Quinchamalium chilense, Argentina Viola reichii, Chile

Viola sacculus, Argentina Viola sacculus, Argentina

Viola cotyledon, Argentina Viola cotyledon, Argentina

Viola columnaris, Argentina Viola ? petraea, Argentina

Viola dasyphylla, Argentina Viola coronifera, Argentina

Viola auricolor, Argentina Viola auricolor, Argentina

Viola vulcanica, Argentina Viola sacculus, Argentina

Araucaria araucana Torres del Paine, Chile

POSSIBLE FLORISTIC HISTORY1. 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

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 America

S America

Introduced species + ++

Land-use changes - +

Hydroelectric development + -

Ski development + +

Atmospheric nitrogen deposition

++ -

Global warming ++ +++ = high threat + = some threat - =

no likely threat

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.

Acknowledgements

Peter Erskine Phil & Gwen Phillips

Marcela Ferreyra Loren Russell

Martin & Anna Sheader Loraine Yeatts

Cathy Jenks