Growth form - Station Alpine Joseph Fourier

Páramos
of Ecuador
III rd Congress of Arctic and Alpine Botanic Gardens
Trento/Viotte – September 2012
Serge Aubert
Station alpine Joseph Fourier
Université de Grenoble

Geography

(mega)Biodiversity
• one of 17 countries in the world which are classified as megadiverse by the group
Conservation International
• more than 1600 bird species, 350 species of reptiles, 400 species of amphibians, 324
species of mammals
• around 15900 species of plants (including ca 4000 orchids), around 4173 of which
are endemic

Alexander von Humboldt (1769-1859) &
Aimé Bonpland (1773-1858)
(par Eduard Ender, ca. 1850)
Humboldt and the biogeography
Mount Chimborazo : 6267 m (DR)

Humboldt and the biogeography
Physical Geography. Humboldt’s Distribution of Plants in
Equinoctial America, According to Elevation Above the Level
of the Sea (1839)
“Outlines of Botanical Geography” and “The Geographical
Distribution of Plants”. From Johnston’s The Physical Atlas
(1849) On this sheet, Johnston incorporated and expanded
data from both Humboldt and Berghaus.

Vegetation

Distribution and climate of páramos
Luteyn 1999 in Sklenar et al 2011 Bot Rev 77:71–108
Hammen & Cleef, 1986 in Sklenar et al 2011 Bot Rev 77:71–108
Luteyn 1999 Páramos: A Checklist of
Plant Diversity, Geographical
Distribution, and Botanical Literature
Venezuelan páramos with Coespeletia
timotensis (photo A. Gröger)
Rundel & al 1994 Tropical Alpine Environments:
Plant Form and Function
Lachemilla orbiculata (Rosaceae, N)
páramo del Angel 3500 m – 6 am

Treeline and páramos

The treeline: Polylepis div sp (Rosaceae)
Polylepis sp (N)
Polylepis incana (N)
Oncidium cucullatum (N) Epidendrum tenuicaule (N) Polylepis racemosa (N)

Bosque siempreverde montano alto

TROPICAL Endemic
(E: 17)
Plant diversity and distribution of genera
Neotropical
(N: 242)
Wide
Tropical
(WT: 58)
TEMPERATE Austral-
Antarctic
(AA: 34)
Holarctic (H:
41)
Wide
temperate
(WTe: 76)
COSMOPOLITAN Worldwide, or
nearly so,
Distribution
(C: 41)
Confined (almost)
exclusively to
páramo
from lowlands to the
alpine zone, also
outside páramo
Widely distributed in
tropical America,
occurring also in the
Palaeotropics
South temperate
distribution
North temperate and
mediterranean
distribution
Temperate and cool
regions of
both hemispheres
TOTAL = 509 genera / 127 families / 3500 species
Aragoa, Ascidiogyne, Blakiella, Bucquetia, Castratella, Cotopaxia, Espeletia, Floscaldasia,
Hinterhubera,Laestadia, Myrrhidendron, Nephropteris, Neurolepis, Perissocoeleum, Plutarchia,
Raouliopsis, Westoniella
Arcytophyllum, Baccharis, Cavendishia, Ceratostema, Chuquiraga, Chusquea,
Diplostephium, Disterigma, Distichia, Dorobea, Epidendrum, Gynoxys, Halenia, Lachemilla,
Lasiocephalus, Loricaria,Jamesonia, Monnina, Nototriche, Odontoglossum, Oncidium, Opuntia,
Oreopanax, Pentacalia, Perezia, Polylepis, Puya, Werneria, Xenophyllum, etc.
Achyrocline, Begonia, Bothriochloa, Buddleja, Bulbostylis, Cheilanthes, Clethra, Conyza, Cyperus,
Dicksonia, Dioscorea, Elaphoglossum, Eragrostis, Eriocaulon, Grammitis, Hedyosmum,
Hymenophyllum, Hypoxis, Ilex, Maytenus, Myrsine, Ocotea, Paspalum, Passiflora, Peperomia,
Persea, Pilea, Piper, Phytolacca, Pteris, Schefflera, Sporobolus, Tournefortia, Xyris, etc.
Acaena, Azorella, Calandrinia, Calceolaria, Colobanthus, Cortaderia, Cotula, Desfontainia,
Drimys, Dysopsis, Escallonia, Fuchsia, Gaultheria, Gunnera, Libertia, Lilaea, Lilaeopsis,
Lomatia, Muehlenbeckia, Myriactis, Myrteola, Nertera, Oreobolus, Oreocallis, Oreomyrrhis,
Orthrosanthus,Ourisia, Pernettya, Prumnopitys,Rostkovia, Tristerix, Ugni, Uncinia, Weinmannia
Antennaria, Astragalus, Bartsia, Berberis, Castilleja, Cerastium, Cirsium, Draba, Erigeron,
Erysimum, Geum, Helianthemum, Hypochaeris, Lathyrus, Lithospermum, Lupinus, Muhlenbergia,
Oenothera, Parietaria, Pedicularis, Phacelia, Potentilla, Ribes, Rhamnus, Salvia, Saxifraga,
Sibthorpia, Silene, Stachys, Thalictrum, Trichophorum, Vaccinium, Verbena, Viburnum, Vicia
Agrostis, Alopecurus, Anemone, Arenaria, Brachypodium, Bromus, Calamagrostis, Callitriche,
Caltha, Cardamine, Cardionema, Carex, Crassula, Coriaria, Cynoglossum, Cystopteris, Danthonia,
Daucus, Deschampsia, Descurainia, Dryopteris, Elatine, Elodea, Elymus, Ephedra, Epilobium,
Festuca, Galium, Gentiana, Gentianella, Geranium, Glandularia, Glyceria, Gnaphalium, Gratiola,
Hieracium, Hierochloe, Hordeum, Hypericum, Isoëtes, Isolepis, Juncus, Lepidium, Limosella,
Luzula, Lysimachia, Melica, Mimulus, Montia, Myosotis, Paronychia, Pilularia, Pingjuicula,
Plagiobothrys, Plantago, Poa, Polypogon, Polystichum, Potamogeton, Puccinellia, Ranunculus,
Rubus, Rumex, Senecio, Sedum, Sisyrinchium, Spergularia, Stellaria, Stipa, Thelypteris, Trisetum,
Urtica, Valeriana, Veronica, Viola
Adianthum, Anagallis, Asplenium, Athyrium, Azolla, Bidens, Blechnum, Botrychium, Eleocharis,
Chamaesyce, Culcita, Cuscuta, Cynanchum, Drosera, Equisetum, Eryngium, Euphorbia,
Habenaria, Huperzia, Hydrocotyle, Lobelia, Lemna, Lycopodiella, Lycopodium, Malaxis, Morella,
Myriophyllum, Nicotiana, Ophioglossum, Oxalis, Pellaea, Polypodium, Pteridium, Rhynchospora,
Scirpus, Selaginella, Solanum, Stuckenia, Utricularia, Wahlenbergia, Woodsia

Ramsay & Oxley (1997) Plant Ecology 131: 173–192
Diversity of growth forms
Ten growth forms of vascular
plants in the Ecuadorian paramos
• stem rosette (e.g., Espeletia
pycnophylla)
• (b) basal rosette (e.g. Puya hamata)
• (c) tussock (e.g., Calamagrostis
coarctata )
• (d) acaulescent rosette (e.g.,
Hypochaeris sessiliflora)
• (e) cushion (e.g., Plantago rigida)
• (f) upright shrub (e.g., Valeriana
microphylla)
• (g) prostrate shrub (e.g., Pernettya
prostrata)
• (h) erect herb (e.g., Bartsia laticrenata)
• (i) prostrate herb (e.g., Geranium
multipartitum)
• (j) trailing herb (e.g., Vicia setifolia).

Diversity of Asteraceae
Gynoxys cuicochensis (N) Dorobea pimpenilifolia (N) Culcitium canescens (N)
Pentacalia vaccinioides (N) Werneria nubigena (N) Perezia pungens (N)
Hypochaeris sessiliflora (H)
Senecio formosoides (WT)
Diplostephium ericoides (N) Chuquiraga jussieui (N) Lasiocephalus ovatus (N) Hieracium frigidum (WTe)

Diversity of Asteraceae: Loricaria
Loricaria thuyoides (N)
Loricaria ferruginea (N)

Diversity of Asteraceae: Baccharis
Baccharis odorata (N) Baccharis genistelloides (N)
Baccharis caespitosa = B. alpina (N)
« Kriechpolster » (Rauh 1939)
Ruthsatz 1978
Darwiniana, 21, 494–539

Frailejones : Espeletia pycnophylla
Páramo de Frailejones

Growth form: arborescent caulescent rosette
Schultze, Beck, Müller-Hohenstein 2005
Plant Ecology, Springer

Growth form: basal rosette
Puya clava-herculis (Bromeliaceae, N)

Growth form: tussocks
Calamagrostis sp (Poaceae, WTe) - Páramo herbáceo
Chusquea sp (Poaceae, N)
Stipa ichu (Poaceae, WT) - pajonal Rhynchospora sp (Cyperaceae, C), Cortaderia nitida (Poaceae, AA)

Growth form: cushions
Xenophyllum humile (Asteraceae, N)

Growth form: cushions
Azorella pedunculata (Apiaceae, AA) - Páramo herbáceo y de almohadillas
« Vollkugelpolster »
(Rauh 1939)

Growth form: cushion bogs
Plantago rigida (Plantaginaceae, WT)

Growth form: cushion bogs
Distichia muscoides (Juncaceae, N)
« Rasenpolster » (Rauh 1939)
Ruthsatz 1978
Darwiniana, 21, 494–539

Growth form: cushion bogs
Biogeography of Oreobolus inferred from
molecular phylogeny suggests a longdistance
dispersal from Australasia to southern South
America and subsequent northward migration to
páramo of northern Andes and Central America;
páramo branches in bold (modified from
Chacón et al., 2006

Growth forms: Viola (WTe)
Viola pygmea
Viola sp Viola sp Viola cotyledon (Chile, Argentina)

Growth forms: cushions & facilitation
Sklenar 2009 Flora 204: 270–277
Huperzia crassa (Lycopodiaceae, C), Gentiana
sedifolia (WTe), Disterigma empetrifolia
(Ericaceae, N), etc.

Growth forms: cushions & competition
Azorella pedunculata > Plantago rigida
Azorella pedunculata - Plantago rigida
Azorella pedunculata < Plantago rigida
Azorella pedunculata - Distichia muscoides

Growth forms: cushions
Valeriana rigida (Valerianaceae, WT)
« Rosettenpolster » (Rauh, 1939)

Growth forms, biogeography and evolution
Valeriana microphylla (Wte)
Valeriana plantaginea
Asian origin of the genus (and family) and
subsequent migration via Europe to North America.
From North America there may have been two
colonization events of the South American continent
(páramo branches in bold) (From Bell & Donoghue,
2005 in Sklenar & al 2011)

Growth forms, biogeography and evolution
Gentiana sedifolia (WTe)
Gentianella hirculus (WTe)
G. rapunculoides Gentianella rapunculoides
Gentianella hyssopifolia
Gentianella originated in
Asia, then dispersed to
other north temperate
regions, and colonization of
South America likely
occurred via North America
(in Sklenar & al 2011)

Gentianaceae: genus Halenia (N)
Asia : 2 species
N America: 1 species
C America: 15 species
S America: 50 species including 39 in paramos
Richness diversity suggested neotropical element while
Reconstructed phylogeny suggests that Halenia
originated in eastern Asia and then immigrated via North
and Central America to South America with three
independent immigrations from Central America to the
northern Andes (von Hagen & Kadereit, 2003 in Sklenar
& al 2011)

Growth forms, biogeography and evolution
Draba steyermarki (Brassicaceaee, H)

Growth forms, biogeography and evolution
Cavendishia tarapotana (N)
Ceratostema alatum (N)
Bosque siempreverde montano alto
Ericaceae
Disterigma empetrifolia (N)
Vaccinium floribundum (H)
Pernettya prostrata (AA)

Elaphoglossum sp
(Lomariopsidaceae, WT)
Ferns
Jamesonia imbricata (Pteridaceae, N)
Cladogram of the Jamesonia-Eriosorus
complex suggests two colonizations
and three independent radiations in the
Andes. Repeated evolution of the
“jamesonia” growth form is inferred
which renders both
Jamesonia and Eriosorus paraphyletic
(modified from Sánchez-B., 2004, Fig.
6)

Espeletia pycnophylla
(Asteraceae)
Puya clava-herculis
(Bromeliaceae)
El Angel, 4000 m
Comparison with other tropical mountains
Sagina abyssinica (Caryophyllaceae)
Helichrysum manii (Asteraceae)
Mount Cameroun (4100 m)
Senecio adnivalis (Asteraceae)
Lobelia wollastonii (Lobeliaceae)
Ruwenzori (3500 m - Photo C. Ruiz)
Eriocaulon pulvinatum
(Eriocaulacceae)
Oreobolus pumilio
(Cyperaceae)
Potentilla foersteriana
(Rosaceae)
Trikora, N-G, 4000 m
Photos: J-M Mangen

Muy ricos la carne… y el vino
Muchas gracias por su atención

Available on the web
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Slides of the conference