Ecological Archives E087-073-A1

Len N. Gillman and Shane D. Wright. 2006. The influence of productivity on the species richness of plants: a critical assessment. Ecology 87:1234–1243.

Appendix A. Summary assessments of productivity-plant species richness relationships included in Mittelbach et al. (2001).

Taxa

Habitat

Productivity measure

Extent§

Grain||

Comments

Relationship reported by Mittelbach et al. (2001) ¶, ††.

Included

Relationship reported here

Biomass

R2

Reference‡‡

T

Forest, East Asia

Production

CG

C

 

+ve

Y

+ve

-

0.84

(Adams and Woodward 1989)

T

Forest, Europe

Production

CG

C

 

+ve

Y

+ve

-

0.84

(Adams and Woodward 1989)

T

Forest, North America

Production

CG

C

 

+ve

Y

+ve

-

0.91

(Adams and Woodward 1989)

T

North America

AET

CG

C

Reanalyzed and reported by Whittaker and Heegaard (2003) as a +ve relationship.

OLS, U-S;

GLM UM.

 

+ve

 

0.67

(Currie and Paquin 1987)

VP

Global

Production

CG

C

Mosaic diversity

+ve

Y

+ve

   

(Scheiner and Rey-Benayas 1994)

H, S, T

Islands

AET

CG

C

 

+ve

Y

+ve

 

0.91

(Wright 1983)

T

Global

AET

CG

C

Refer to text

UM

N

     

(Latham and Ricklefs 1993)

T

Tropical Forest, America

Rainfall

CG

C

Reanalyzed below using tree turnover, which is a preferable index for productivity.

GLM +ve; OLS NS.

N

     

(Phillips et al. 1994)

T

Forest, Asia

Rainfall

CG

C

N<10

NS

N

     

(Phillips et al. 1994)

T

Forest, Global

Rainfall

CG

C

High rainfall confounded with high altitude and latitude.

UM

N

     

(Phillips et al. 1994)

VP

Global

Production

CG

F

Smaller plot sizes towards the poles. However, plot sizes were not correlated with productivity and number of stems per plot is negatively correlated with productivity.

+ve

Y

+ve

 

0.43

(Scheiner and Rey-Benayas 1994)

H, S

Desert, North America

Warm season rainfall

CG

F/C

Quadrat numbers varied among sites (32–48); area of quadrats varied among sites (58–231 m2).

NS

N

     

(Pianka 1967)

H, S

Desert, Kalahari

Rainfall

CG

F/C

Quadrat numbers varied among sites (16–41); area of quadrats varied among sites (58–232 m2)

NS

N

     

(Pianka 1971)

H, S, T

Forest, Neotropics

Rainfall

CG

F

No control for latitude (high rainfall sites are up to 20° further from the equator than the other sites); Very high rainfall (> 4000 mm/yr) is unlikely to represent high productivity (Kay et al. 1997); 0.1-ha plots.

UM

N

     

(Gentry 1988)

H, S, T

Forest, Africa

Rainfall

CG

F

N<10; Sites with high rainfall include an active volcano and very poor soils.

NS

N

     

(Gentry 1988)

T

Forest, Tropical South America

Rainfall

CG

F

Rainfall cannot be used as a surrogate for productivity in this case because productivity declines with increasing rainfall beyond 2500 mm/yr. Tree richness data from Clinebell et al (1995) (see below). 0.1-ha plots.

UM

N

     

(Kay et al. 1997)

H, S

Desert

Rainfall

R

F

High rainfall sites were also at high altitude. However, an over estimation of productivity in the upper productivity range will have had a conservative influence on the reported relationship; OLS Plinear = 0.028; 25-m line intercept samples.

GLM, NS; OLS +ve.

Y

+ve

 

0.27

(Brown 1973)

H

Bogs, North America

Rainfall, MAT, freezing days

R

C

GLM Plinear= 0; OLS Plinear< 0.001

GLM +ve; OLS +ve.

Y

+ve

 

0.60

(Glaser 1992)

H, S, T

Wet forest, Australia

Nutirent

R

C

GLIM: Pquad > 0.05;
GLIM: Plinear < 0.001;
OLS Plinear = 0.013

U-S

Y

+ve

 

0.56

(Beadle 1966)

S, T

South Africa

Rainfall

R

C

Rainfall confounded with low temperatures at high altitude. However, an over-estimation of productivity in the high range could not have contributed to the reported relationship.

GLM, UM; OLS +ve

Y

+ve

 

0.59

(O'Brien 1998)

H

Desert-dry forest, Namibia

Rainfall

R

C

Altitude confounded with rainfall. However, this could not have contributed to the reported relationship.

+ve

Y

+ve

 

0.77

(Schulze et al. 1996)

T

Madagascar

Rainfall

R

F

High altitude confounded with high rainfall at one site. However, this could not have contributed to the reported relationship; 0.1-ha plots; OLS +ve P< 0.05

OLS +ve

Y

+ve

 

0.44

(Ganzhorn et al. 1997)

S, T

Tropical savanna, Australia

Rainfall

R

F

Reanalyzed and reported by Whittaker and Heegaard (2003) as a +ve relationship; 0.04-ha plots.

OLS U-S

Y

+ve

 

0.30

(Williams et al. 1996)

H, S, T

Monsoon forest, Australia

Rainfall

R

F

GLM Plinear = 0;
OLS Plinear < 0.001;
0.84-ha plots.

GLM +ve; OLS +ve

Y

+ve

 

0.55

(Russel-Smith 1991)

H

Savannah, Venezuela

Rainfall

R

C

OLS Plinear = 0.04;
GLM Plinear = 0.088;
100-m2 plots.

GLM +ve; OLS +ve

Y

+ve

 

0.62

(Sarmiento 1983)

H, S

Wetland, Canada

SCB

R

F

Standing crop not necessarily correlated with productivity; 0.25-m2 plots.

GLM -ve; OLS -ve.

Y

 

-ve

0.27

(Moore and Keddy 1989)

H, S, T

Lakeshore, Canada

SCLB

R

F

0.25-m2 plots

GLM UM; OLS -ve

Y

 

-ve

0.16

(Wisheu and Keddy 1989)

H

Grassland, Kenya

SCB

R

F

Canopy intercept of five pins/m × four replicates.

NS

Y

 

NS

-

(McNaughton 1983)

Moss

Forest, Australia

Rainfall

R

-

N<10

 

N

     

(Fensham and Streimann 1997)

H, S

Arid zone, Chile

Rainfall

R

-

N<10

 

N

     

(Meserve and Glanz 1978)

H, S, T

Forest, USA

Moisture

R

-

N<10

 

N

     

(Monk 1965)

H, S

Grassland

Rainfall

R

-

N<10

 

N

     

(Nicholson and Hulett 1969)

S, T

Forest, USA

Rainfall

R

-

N<10

 

N

     

(Ohmann and Spies 1998)

H

Petland, Canada

Nitrogen

R

-

N<10

NS

N

     

(Wells 1996)

T

Forest, Australia

Rainfall

R

C

A decline in species richness of a drought-adapted taxonomic group as rainfall increases is not a decline in richness due to productivity. Instead it reflects the replacement of drought-adapted plants by rainforest taxa.

UM

N

     

(Hughes et al. 1996)

Cacti

Argentina

Summer Rainfall

R

C

A decline in species richness of a desert-adapted taxonomic group (cacti) as rainfall increases is not a decline in richness due to productivity. Instead it reflects the replacement of desert-adapted plants by drought intolerant taxa.

UM

N

     

(Mourelle and Ezcurra 1996)

H, S, T

Dry/heath forest, Australia

Nutrient

R

C

N<10; OLS P > 0.1

 

N

     

(Beadle 1966)

H, S, T

California

Rainfall

R

C

No control for altitude or latitude.

UM

N

     

(Richerson and Lum 1980)

H

Desert, Egypt

Rainfall

R

C

Diversity count limited to grass species. A decline in grass species with increasing rainfall is likely to be due to the replacement of grasses with woody vegetation.

UM

N

     

(Vogel et al. 1986)

H, S, T

Forest, Canada

Moisture index

R

C

Moisture was confounded with altitude; GLM and OLS P > 0.05; 2.4-ha plots.

UM

N

     

(La Roi and Hnatiuk 1980)

H

Grassland, Kenya

Moisture index

R

F

Moisture confounded with altitude; 1m2 plots.

UM

N

     

(Tieszen et al. 1979)

H, S

Desert, USA

Rainfall

R

C

Rainfall confounded with altitude; No control for latitude;
GLIM: Pquad > 0.05;
OLS: Plinear > 0.05; 0.15-ha plots.

U-S

N

     

(Barbour and Diaz 1973)

S, T

Beech forest, China

Rainfall

R

F

Rainfall confounded by latitude, altitude and mean temperature; Highest diversity at medium rainfall, but also at greatest temp and lowest latitude; Number and size of plots varied

GLM UM; OLS NS

N

     

(Cao and Peters 1997)

T

Forest, Costa Rica

SCB

R

F

No control for altitude or latitude; Standing crop is not necessarily a good estimate of productivity for trees; 0.37-ha plots.

NS

N

     

(Huston 1980)

H, S, T

Forest, USA

Production

R

F

N<10; 100-m2 plots.

NS

N

     

(Westman and Whittaker 1975)

H

Steppes - forest, Syria

Rainfall

R

C

No control for altitude; Sites modified by grazing stock; Limited taxonomic group used (legumes); 207-m2 plots.

UM

N

     

(Ehrman and Cocks 1990)

H, S

Alpine, Australia

Nitrate

R

F

No control for climate. Authors stated that climate variables were more important than soil; 25-m2 plots.

UM

N

     

(Kirkpatrick and Bridle 1998)

S, T

Chaparral, USA

Rainfall

R

F

No control for altitude or latitude; 8-m2 plots.

NS

N

     

(Keeley 1992)

H, S, T

Forest, North America

Rainfall

R

F

N<10; No control for altitude (1350 m alt variation), or latitude; 0.1-ha plots.

NS

N

     

(Gentry 1988)

H, S

Grassland

Rainfall

R

C

N<10; Stands 8–200 ha.

 

N

     

(Diamond and Smeins 1988)

H, S, T

Arid -rainforest, Australia

Nutrient

R

C

Only rainforest genera were counted and therefore the study does not provide a valid estimates of diversity in dry zones; OLS: P > 0.1

GLM U-S; OLS NS

N

-

   

(Beadle 1966)

H

Fen, UK

Production

L

F

Change in St. Crop measured over time; Past management may have influenced diversity; 4-m2 plots.

U-S

Y

U-S

 

0.41

(Wheeler and Shaw 1991)

T

Forest, Canada

Nutrient index

L

F

GLM Plinear> 0.05;

400-m2 Plots.

GLM -ve; OLS U-S

Y

U-S

 

0.58

(Nantel and Neumann 1992)

H

Fen, UK

Production

L

F

Change in St. Crop measured over time. 0.25-m2 plots.

U-S

Y

U-S

 

0.35

(Wheeler and Giller 1982)

H, S, T

Forest, USA

Above ground production

L

F

Productivity was estimated from core increments, volume estimates and wood density; OLS P>0.05; 0.1–0.5-ha plots.

GLM +ve;

OLS NS.

N

NS

   

(Whittaker 1966)

H, S, T

Shrub - Forest, USA

Rainfall

L

F

All sites with low rainfall (< 380 mm/yr); 1-ha plots.

GLM +ve; OLS +ve

Y

+ve

 

0.62

(Harner and Harper 1976)

H

Grassland, Netherlands

Nitrogen

L

F

1-m2 plots.

+ve

Y

+ve

 

0.23

(Werger et al. 1983)

H, S, T

Desert, woodland, forest, USA

Biomass production

L

F

OLS P > 0.1; 0.1-ha plots.

GLM U-S;

OLS NS.

 

NS

-

 

(Whittaker and Niering 1975)

H

Desert, Israel

SCB

L

F

All plants were annuals, therefore, biomass can be used as an estimate for productivity;
0.1-m2 plots. GLM -ve Plinear > 0.05;
OLS Plinear > 0.1

GLM -ve; OLS NS.

 

NS

   

(Danin 1976)

H

Desert, USA

SCB

L

F

Site modified by grazing stock prior to 1977; Biomass dies back each year, therefore biomass can be used as an index for productivity; 0.25-m2 plots.

 

Y

UM

 

0.15

(Guo and Berry 1998)

H, S

Sand dune, Israel

SCB

L

F

Biomass justified as an index for productivity; OLS MOS P < 0.001; GLM MOS P < 0.001; 0.09-m2 plots.

UM

Y

UM

 

0.31

(Kutiel and Danin 1987)

H, S, T

Deciduous forest, Canada

Nutrient index

L

F

Highest richness also highest nutrient; Residuals normal RJ = 0.968,
Pnon-normal > 0.100;
400-m2 plots.

UM

Y

UM

 

0.93

(Loucks 1962)

H, S

Inselbergs, Ivory Coast

Rainfall

L

F

Inselbergs are less abundant in high rainfall areas and this implies that the –ve relationship may be an area effect; 3-m2 plots.

-ve

Y

-ve

 

0.4

(Porembski et al. 1995)

H, S, T

Fynbros - shrubland, South Africa

Soil fertility index

L

F

GLM UM Pquad > 0.05; OLS P > 0.1

GLM UM; OLS NS.

Y

 

NS

 

(Cowling 1990)

H, S

Wetland, USA

SCLB

L

F

Biomass was found by the study to be less important than salinity and elevation; Plants included perennials; OLS Pquad > 0.05; 1-m2 plots.

GLM UM

OLS NS

Y

 

-ve

 

(Gough et al. 1994)

H

Coastal wetlands, Louisiana

Biomass

L

F

1-m2 plots.

NS

Y

 

NS

 

(Grace and Pugesek 1997)

H, S

Coastal dunes, Netherlands

SCLB

L

F

Woody species excluded; 0.02-m2 plots.

UM

Y

 

UM

 

(Klinkhamer and deJong 1985)

H, S, T

Grassland-forest, Estonia

SCB

L

F

Species/500 ramets + species/m2; Plants included perennials; OLS P>0.05.

OLS NS;

GLM UM

Y

 

NS

 

(Zobel and Liira 1997)

H, S

Shoreline, Canada

SCB

L

F

A mixture of perennials and annuals; 0.25-m2 plots.

UM

Y

 

UM

0.44

(Shipley et al. 1991)

H, S, T

Lakeshore, Canada

SCB

L

F

Only vegetation up to 50 cm high was included; 0.25-m2 plots.

UM

Y

 

UM

0.13

(Wilson and Keddy 1988)

H, S, T

Lakeshore, Canada

SCLB

L

F

Only two sites used, one dominated by Typha; Plot sizes may have been too small for vegetation type; 0.25-m2 plots.

UM

Y

 

UM

0.16

(Wisheu and Keddy 1989)

H

Salt marsh, Spain

SCB

L

F

Biomass confounded with salinity and water logging; 0.25-m2 plots.

UM

N

     

(Garcia et al. 1993)

T

Forest, USA

SCB

L

C

1.6-ha plots.

+ve

Y

 

+ve

0.19

(Risser and Rice 1971)

H, S

Alpine, PNG

SCB

L

F

4-m2 plots; OLS P>0.1

GLM +ve;

OLS NS

Y

 

NS

 

(Walker 1968)

H

Desert, Greenland

Nitrogen

L

 

N<10

 

N

     

(Bay 1997)

H

Grassland, Argentina

Rainfall

L

 

N<10

 

N

     

(Cabido et al. 1997)

H

Shrub/grassland, Argentina

Annual Rainfall

L

C

N<10

 

N

     

(Cavagnaro 1988)

H, S, T

Shrubland - forest, New Mexico

SCB

L

F

N<10; 12.5-m2 plots.

 

N

     

(Cully and Cully Jr 1989)

H

Forest, Mexico

Biomass

L

 

N<10

 

N

     

(Hietz and Hietz-Seifert 1995)

H

Sand plain, USA

Nitrogen

L

F

Highly modified environment with exotic species; Relationship reflects successional processes, including increases in native species over time; 0.5-m2 plots.

-ve

N

     

(Inouye et al. 1987)

H, S

Dunes, Netherlands

Nitrogen

L

F

Nitrogen is not an adequate surrogate for productivity in dunes where water availability is a more important limiting factor for productivity.

NS

N

     

(Laan 1979)

H, S, T

Alpine, Sweden

Growing Days

L

F

Plots too small for 5-m high vegetation; 0.5-m2 plots.

UM

N

     

(Nilsson and Wilson 1991)

H, S, T

Lakeshore, Sweden

Growing Days

L

F

Plots too small for 5-m high vegetation; 0.5-m2 plots.

UM

N

     

(Nilsson and Wilson 1991)

H

Alpine tundra, Wyoming

SCB

L

F

Not all species were reported, only the 14 most common; Sites modified by sheep grazing; 0.15-m2 plots.

UM

N

     

(Scott and Billings 1964)

H, S, T

Lakeshore, Canada

SCLB

L

F

N<10; 0.25-m2 plots.

 

N

     

(Day et al. 1988)

H, S, T

Forest, USA

Production

L

F

N<10;
0.1-ha plots.

NS

N

     

(Whittaker and Woodwell 1969)

H, S

Shrubland - desert, Israel

Annual Rainfall

L

C

N<10; 0.2-ha plots.

 

N

     

(Kutiel et al. 1995)

H, S

Grassland, Tanzania

Annual Rainfall

L

C

N<10; 0.48-ha plots.

 

N

     

(Schmidt 1975)

H

Alpine, Europe

Production

L

F

N<10; 1.25-m2 plots.

 

N

     

(Onipchenko et al. 1998)

H, S

Grassland, USA

Productivity

L

F

N<10; 1.5-m2 plots.

 

N

     

(McNaughton 1968)

H, S

Fen, Poland

Production

L

F

N<10; 10-m2 plots.

+ve

N

     

(Wassen et al. 1990)

H, S

Desert, Argentina

Rainfall

L

C

OLS: Plinear > 0.1; 1536-m2 plots.

+ve

N

     

(Barbour and Diaz 1973)

H

Grassland, USA

Production

L

C

Sites modified by mowing and stock grazing; Fewer samples taken in low diversity areas; Patch sizes of low diversity stands were very small; 15-m2 plots.

-ve

N

     

(Redmann 1975)

H

Bog, UK

Biomass production

L

F

N<10; 3.1-m2 plots.

 

N

     

(Forrest and Smith 1975)

H, S, T

Wetland forest

SCB

L

F

N<10; 3-m2 plots.

-

N

     

(Kirkman et al. 1998)

H, S, T

Wetland, Alaska

Productivity Index

L

F

Above ground herb productivity measured, but not tree productivity; 55-m2 plots.

OLS NS; GLM, UM

N

     

(Pollock et al. 1998)

H

Grassland - marsh, USA

Drainage

L

C

Highly modified sites from burning, mowing; Drainage is not a good index for productivity as both excessive drainage and water-logging are likely to depress productivity; 75-m2 plots.

UM

N

     

(Dix and Smeins 1967)

H, S, T

Forest, USA

Elevation

L

F

N<10; 0.1-ha plots.

 

N

     

(Glenn-Lewin 1975)

H, S

Subalpine, USA

Biomass

L

C

N<10; 364-m2 plots.

 

-

     

(Kuramoto and Bliss 1970)

T

Rainforest, Borneo

Est. St. Crop

L

 

N<10

 

N

-

-

 

(Aiba and Kitayama 1999)

H, S, T

Desert, Chile

Rainfall

L

C

N<10

 

N

-

   

(Gutierrez et al. 1998)

S, T

Deciduous forest, USA

Annual Rainfall

L

F/C

Sample area varied from 2–28 ha; Not all species reported in results.

NS

N

-

   

(Keever 1973)

H

Desert, USA

Rainfall

L

 

N<10; Highest rainfall sites dominated by exotic weeds.

-ve

N

-

   

(Beatley 1969)

H, S, T

Forest, Australia

P

L

F

N<10; 0.1-ha plots.

 

N

-

   

(Blanche and Westoby 1995)

H

Grassland/woodland, UK

St. Crop + Litter Biomass

L

F

Canopy cover varied among sites; Highly modified sites; OLS Pquad > 0.1;

GLM Pquad > 0.5;

0.25-m2 plots.

GLM UM; OLS NS

N

-

-

 

(Al-Mufti et al. 1977)

H, S

Fynbos, South Africa

Foliage Area Profile

L

F

The plots were too small for large shrubs; A justification for using Foliage Area Profile as a surrogate for productivity was not provided; 1-m2 plots.

UM

N

-

   

(Bond 1983)

† Taxa: LP = lower plants (lichen and bryophytes); H =herbs, grasses and sedges; S = shrubs; T = trees, A = Angiosperms; VP = all vascular plants; P = all plants.

‡ Extent: L = local-landscape extent (<200 km between sites); R = regional extent (200–3000 km between sites); CG= continental to global extent (>3000 km between sites).

§ Grain: F= fine grain (point diversity and alpha diversity); C = coarse grain (gamma diversity).

|| Productivity measure: AET = Actual evapotranspiration; PET = Potential evapotranspiration; NPP = Net primary production; T = Temperature; MAT = Mean annual temperature; SCB = Standing crop biomass; SCLB = Standing crop plus litter biomass.

¶ Form of relationships: +ve = positive monotonic relationship; -ve = negative monotonic relationship; UM = unimodal relationship; NS =nonsignificant relationship; US = U-shaped.

†† Regression type used: OLS = Ordinary Least Squares; GLM = General Linear Model.

‡‡ See Appendix C for full citations.  



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