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Altitudinal gradients
Montane forests in
the tropics
tropical
Temperature at the forest
limit in altitude in
tropical, subtropical and
temperate mountains
subtropical
temperate
Decrease in temperature with altitude: 5-6oC per 1000 m
Mt. Wilhelm, Papua New Guinea
Mean daily T [oC]
30
Mt Wilhelm: 5.4 oC per 1000 m elevation
30
200
25
1200
20
2000
15
2700
10
3700
5
0
m asl.
02-May-10 10-Aug-10 18-Nov-10 26-Feb-11 06-Jun-11 14-Sep-11
Date
25
Mean annual T [oC]
35
20
15
10
y = -0.0054x + 27.776
R2 = 0.987
5
0
0
1,000
2,000
Altitude m
3,000
4,000
Annual range of monthly mean temperature:
an altitudinal gradient
Diurnal
and annual
temperature
range in high
mountains
tropical
-
temperate
Altitude of treeline and snowline at different latitudes
Korner 1998. Oecologia 115:445
Altitudes and tree species at the forest limit in humid Asian mts.
BETULA
PICEA
LARIX
ABIES
Forest altitudinal zonation in Malesia
Altitudinal zonation on some tropical mountains
Malay
Peninsula
Altitudinal zonation in New Guinea
Mt. Wilhem, New Guinea
forest limit at 3,600 m asl
Mt. Kenya
Sinarundinaria alpina, Graminaceae
Mt. Kenya
a) tree and shrub savannah area, up to 2.200 m;
b) mixed forest belt up to 3300m : the blue line represents a Bamboo dominant area, the
yellow line defines a Hagenya-Hypericum zone;
c) alpine zone: composed mainly of Ericaceae and grass thickets.
Tropical alpine
grasslands: Paramo
Tropical alpine
grasslands: Puna
Alpine tropical plants:
meristems isolated
against cold by rosettes,
hairs, dry leaves...
Lobelia telekii (Africa)
Senecio (Africa)
C3 replace C4 grasses along an altitudinal gradient
Kenya
Ecuador [based on 220 species]
Tieszen et al. 1979, Boom et al. 2001
At high temperatures and low CO2, the C3 photosynthetic enzyme rubisco fails to
completely distinguish CO2 and O2. This leads to photorespiration, resulting in
losses of photosynthetic carbon. C4 photosynthesis suppresses photorespiration by
concentrating CO2 internally, but this comes with an energetic cost, which exceeds
the photorespiratory costs of C3 photosynthesis at high CO2 and low temperatures.
Edwards et al. 2010, Science 328, 587
Edwards & Smith 2010, PNAS 107: 2532–2537
C4 grasses dominate in the tropics
C3 grasses
C4 grasses
agriculture
forests
other
Edwards et al. 2010, Science 328, 587
Montane forests
New Guinea, montane forest at 2,200 m asl
Altitudinal trends - trees >20 cm DBH in Africa
basal area m2/ha
no. of species /ha
Altitudinal trends in the main forest characteristics
Pinnate leaves
Leaf size
Frequent
Mesophyll
Rare
Mesophyll
Very rare
Microphyll
Elfin forest
Costa Rica
Elfin x lowland forests
trees
DBH>10cm:
No. trees/ha
Basal area m2/ha
LAI
wood support
efficiency kg
2
wood/m foliage
* m height
Elfin forest
Puerto Rico
4000
44
2.2
1.1
[90 t of wood
to support 2 ha
leaves 4 m
above ground]
Lowland forest
Malaysia
500
30
7.4
0.2
[420 t of wood
to support 7.4 ha
leaves 30 m
above ground]
Biomass and water interception in a mossy elfin forest
Tanzania (2140 m asl)
canopy epiphytes/ litter
leaves
trunk epiphytes
Bryophytes along tropical altitudinal gradient
Two modes of altitudinal species turnover:
with complete nestedness and zero nestedness
Identical altitudinal trends in species richness mean
different trends in mean altitudinal range of species and
beta diversity between adjacent altitudes
1.0
Beta diversity [1-Jaccard] .
10
No. of species
8
6
4
2
0
0.8
0.6
0.4
0.2
0.0
low
high
Altitude
low
high
Altitude
Species turnover along altitudinal gradients:
Rhododendron spp. on Mt. Kinabalu
Rhododendrones: 900 spp. worldwide, 300 spp. in SE Asia, 50 spp. in
Borneo, 25 spp. on Mt. Kinabalu, incl. 5 endemic spp.
Altitudinal distribution of 454 bird species in Papua New Guinea
0 m asl.
4500 m asl.
each row is 100 m elevation belt, each column a bird species
K. Tvardikova, unpubl. data
Checkerboard distribution along an altitudinal gradient
Altitudinal segregation of competing parrots in New Guinea:
2500
Altitude (m)
2000
1500
1000
500
0
Mt. Niba
Sepik
Mts.
Mt.
Karimui
Charmosyna placentalis
Karkar
Is.
Tolokiwa
C. rubronotata
New
Britain
C. rubrigularis
Global elevation
patterns of bird diversity
decreasing diversity
low-elevation plateaus
low-elevation plateaus
with mid-peaks
unimodal midelevational
peaks
McCain, Global Ecology and
Biogeography, (2009) 18, 346–360
Species richness along altitudinal gradients:
• monotonous decrease from the lowland maximum
OR
• peak at mid-elevations?
Factors causing decreasing species richness with altitude:
- harsh environmental conditions (temperature)
- diminishing habitat area with altitude
Mid-elevation peak:
- possibly as a result of geometric constraints:
both lowland and high-montane species can overlap to the midelevation
Geometric constraints:
Madagascar study of mycalesine butterflies
Mycalesine butterfly
species richness
Henotesia fraterna
Lees et al. 1999. Biol. J. Linn. Soc. 67:529
Himalayas
Birds
Mammals
Lepidoptera, Mt Wilhelm, New Guinea
Ants: decreasing abundance and
species richness with altitude,
practically absent above 1,800 m
Mammals
Altitudinal gradients in New Guinea
No. of species
100
total species: 133
max per 100m: 78 [59%]
80
60
40
20
3400
3100
2800
2500
2200
1900
1600
1300
1000
700
400
100
0
Altitude [m]
Birds
500
Altitude [m]
Altitude [m]
3400
3100
2800
2500
2200
1900
3400
3100
2800
2500
2200
1900
1600
1300
1000
0
700
0
400
100
100
50
1600
100
200
1300
150
300
1000
200
400
700
No. of species
250
total species: 700
max. per 100m: 468 [67%]
400
total species: 459
max. per 100m: 300 [65%]
300
No. of species
Butterflies
100
350
1
Jaccard similarity .
Jaccard similarity .
1.0
0.9
0.8
0.7
0.8
0.6
0.4
0.2
0
0.6
0
500 1000 1500 2000 2500 3000 3500
0
500 1000 1500 2000 2500 3000 3500
Altitudinal difference [m]
Altitude m
Mammals
Linear (Butterflies)
Birds
Linear (Birds)
Butterflies
Linear (Mammals)
Mammals
Birds
Butterflies
Species turnover:
- between neighbouring 100m
altitudinal zones
- with increasing altitudinal
distance
Butterflies along altitudinal
transect at Mt Wilhelm:
200 m asl
700 m asl
1200 m asl
1700 m asl
2200 m asl
are lowland species just
disappearing with altitude,
or are only new montane
species appearing, or both
at the same time?
Legi 2011
Rapoport’s rule:
do montane species have wider altitudinal ranges?
Mean altitudinal range [m] .
2500
2000
1500
1000
500
0
0
max
Altitude [m]
Birds
min
range width
midpoint
500 1000 1500 2000 2500 3000 3500
Butterflies
Mammals
Land area decreases enormously
with altitude:
we need to consider species-area
effect in all altitudinal patterns
New Guinea: Surface Area km2
New Guinea: Surface Area km2
1,000,000
400,000
350,000
100,000
300,000
250,000
10,000
200,000
1,000
150,000
100,000
100
50,000
0
Altitude
Altitude
4550
4250
3950
3650
3350
3050
2750
2450
2150
1850
1550
1250
950
650
350
50
50
35
0
65
0
95
0
12
50
15
50
18
50
21
50
24
50
27
50
30
50
33
50
36
50
39
50
42
50
45
50
10
Species-area curves [power and logarithmic]
fitted in altitudinal richness gradient:
area alone predicts a strong diversity trend
lowland
100
y = 63.993Ln(x) - 419.79
R2 = 0.8243
10
1000
10000
100000
1000000
Land area per 100m alt. [km2]
Species-area curve S=b*Az with z=0.25
and b estimated from lowland data used to
predict altitudinal trends in species
richness and density in birds
350
S=b*A
300
0.25
250
200
150
100
50
0
0
500 1000 1500 2000 2500 3000 3500
No. of bird species per 1000 km 2
y = 3.3162x0.4137
R2 = 0.7881
alpine
No. of bird species
No. of bird species
1000
50
S = b * A0.25
45
40
35
30
25
20
15
10
5
0
0
500 1000 1500 2000 2500 3000 3500
Altitude [m]
S observed
Altitude [m]
S predicted
S observed
S predicted
350
Spp. per 1000 km2
mammals, birds
300
40
250
200
30
150
20
100
10
50
Spp. per 1000 km2 butterflies
50
0
0
1000000 100000 10000
1000
100
Land area [km2]
0 m asl.
3,500 m asl.
Species per 1000 km2
50
25
0
0
500
1000 1500 2000 2500 3000 3500
Altitude [m]
Mammals
Birds
Butterflies
Species density:
typically [but not always]
increases with altitude, as
expected from a power
species area relationship
Climate change: species climbing up montane slopes
Mt. Kinabalu, Lepidoptera: average increase in altitude 67 m from 1965 to 2007
Chen et al., PNAS 106:1479, 2009
Altitudinal gradient: the mother of all environmental gradients
W. Barthlott, G. Kier, H. Kreft, W. Küper, D. Rafiqpoor & J. Mutke 2005
Altitudinal gradient: the mother of
all environmental
Beta
diversity of gradients
vertebrates
Birds
Amphibians
Mammals
decrease in species overlap over 500 km