Ecography ECOG-00880Fenberg, P. B., Menge, B. A.,, Raimondi, P. T. and Rivadeneira, M. M. 2014. Biogeographic structure of the northeastern Pacific rocky intertidal: the role of upwelling and dispersal to drive patterns. – Ecography doi: 10.1111/ecog.00880

Supplementary material

  1  

Appendix 1

Meta-analysis. We carried out a survey of studies that examine coastal

biogeographic structure of species inhabiting rocky habitats published from 1992 –

2012 to ask the following questions: (i) which and how many variables have been

mentioned to be important for explaining/predicting biogeographic structure, and (ii)

how many of these variables have been statistically tested? We do this by recording

the number and identity of variables mentioned in each publication and then record

which (if any) of the variables are statistically tested to help predict or describe

observed structure. In total, we surveyed 47 papers covering many coastal regions of

the globe. The main criteria for choosing papers were that they studied marine

biogeographic structure at least at regional scales (using species occurrence and/or

species abundance) and that they primarily studied species living in coastal rocky

environments. Because we also wanted to know what proportion of studies test

variables to predict/explain observed structure, we did not a priori choose studies that

do so. Overall results are presented in tables A1 and A2.

 

  2  

Table A1. Literature survey of the variables cited as potential drivers of the 1  

biogeographic structure of coastal marine organisms. Numbers in parentheses indicate the 2  

number of studies (out of 47 articles, see references) citing its potential importance. 3  

4  Major Processes/Forcing Variables Anthropogenic Forcing (5) Acidification (2)

Fisheries (3)

Invasive Species (2)

Pollution (4)

Population Pressure (2)

Shipping (2)

Unspecified (2)

Climatic (9) Air Temperature (1)

Day Length (1)

Enso Unspecified (2)

Photoperiod (1)

Precipitation (4)

Wind (3)

Unspecified (3)

Dispersal (14) Dispersal Potential (5)

Larval Dispersal (6)

Rafting (2)

Recruitment (1)

Reproduction Strategy (1)

Unspecified (1)

Disturbance (13) Freshwater Input (6)

Ice Scouring (3)

Sediment (8)

Turbidity (2)

Unspecified (1)

Growth Pattern (2) Growth Form (1)

Growth Potential (1)

Growth Rate (2)

Habitat (30) Availability (13)

Geographic Isolation (13)

Heterogeneity (8)

Type (10)

Historic/Evolutionary (18) Glaciation (1)

Macroevolutionary Dynamics (10)

Sea Level Change (1)

  3  

Tectonic Events (7)

Barriers (6)

Evolutionary Unspecified (4)

Oceanographic (43) Coastal Topography (10)

Conductivity (1)

Currents (21)

Depth Unspecified (2)

Enso Effects On Currents (1)

Hydrodynamics (4)

Light Attenuation (1)

Nutrients (15)

Oxygen (2)

pH (4)

Primary Productivity (12)

Salinity (15)

Sea Bottom Slope (2)

Sea Level Change (3)

Water Temperature/SST (38)

Suspension Particulate Matter (1)

Unspecified (2)

Upwelling (14)

Upwelling Seasonality (1)

Wave Exposure (14)

Physiological Tolerance (5) Desiccation Tolerance (2)

Eurybathy (1)

Thermal Stress (1)

Thermal Tolerance (1)

Unspecified (1)

Species Interactions (11) Algal Biomass (2)

Colonization (1)

Competition (3)

Dominance (2)

Facilitation (1)

Food Availability (1)

Functional Group (1)

Predation (2)

Species Richness (1)

Unspecified (3)

 5      6  

  4  

Table A2. Literature survey of studies evaluating the role of environmental variables as potential predictors of marine biogeographic 7  structure. Variables were classified according to 10 major classes of processes. Numbers show the number of variables cited as 8  potential drivers of biogeographic structure. Also shown are the most cited variables across all studies, indicating if the variable is 9  only cited (X) or cited and tested (T) by each particular study (also see Table A1). 10    11   Major Class/Process Top cited variables

Geographic Region Taxa/Habitat studied

Ant

hrop

ogen

ic F

orci

ng

Clim

atic

Dis

pers

al

Dis

turb

ance

Gro

wth

Pat

tern

Hab

itat T

ype

His

toric

Oce

anog

raph

ic

Phys

iolo

gica

l Tol

eran

ce

Spec

ies I

nter

actio

ns

Sea

Tem

pera

ture

Cur

rent

s

Nut

rient

s

Salin

ity

Upw

ellin

g

Wav

e Ex

posu

re

Reference

Global & "colder part of n. hemisphere" Crustose coralline red algae

2 1 3

T X

X

(1)

Southern Polar regions Encrusting faunas (shallow rocky)

1

2

3

X

X

(2)

Global Bivalves

3

T

T

(3)

NE Pacific Rocky intertidal macroinvertebrates and algae

2

1

5

1 T

X

X X

(4)

East coast of S. Africa Intertidal algae

2

4 1

X

X

X

(5)

Chile and Antarctica Decapod crustaceans

1

2 1

X

(6)

  5  

Global Coastal fishes mostly but discussion of other taxa

1 1

2 3 5

X X X

X

(7)

Southern African rocky shores

Rocky intertidal macroinvertebrates and algae

1

4 2

X

X T

(8)

Victoria, Australia Nearshore fish

1

2 1 4

1 X X X X

(9)

Global Rocky shores Rocky intertidal macroinvertebrates and algae 7 3

4

T

(10)

Eastern Pacific Shallow waters (Polar and Temperate)

Isopods

1

2

X X

(11)

Southwest Atlantic Tropical reef fishes

1 2

2 1 4

X X

X

(12)

Brazilian coast Reef fish

1 2

4 1 5

1 T

X

X

(13)

Atlantic Atlantic reef fish

1 2

2 3 4

X X

X X

(14)

Atlantic Opisthobranch gastropods

1

2

2

X X

(15)

SW Atlantic (<80 depth) Hydroids

1

2

X

(16)

Pacific coral region Reef-building coral (41 species) 1 1 3 1 3 2 1 4

3 X X

X

(17)

NE Pacific and NW Atlantic (<200m depth)

Marine algae (NEP) and gastropods (NWA)

1 1 3

X

X X

(18)

  6  

Tropical Eastern Pacific (TEP) Chaenopsid rocky shore reef fish

1

(19)

Chilean coast Sea anemones

1 1

1

2 1

X

(20)

Southeast Australia Rocky intertidal macroinvertebrates

1

1 1 2

1 X X

(21)

Global Nearshore Echinoderms (rocky intertidal and shallow rocky subtidal) 7

2

1

4

1 T

T

X

(22)

Rocky intertidal: Norway, Iceland, NE Canada Rocky intertidal animals

1

(23)

Iberian peninsula (Spain): Intertidal

Peracarid crustaceans (those associated with Corallina elongata)

2

1

5

5 T

(24)

Japan Sea Coastal fishes

1

2

T X

(25)

Chilean coast Rocky intertidal macroinvertebrates

1

3

X X

(26)

Southeast Australia Demersal fishes and macroinvertebrates - shallow rocky reefs

1

8

T X T X

T

(27)

Southwest Atlantic (36-55ºS) Macroalgae

1 1 2

X

X

(28)

Pacific coast of S. America Benthic algae

5

X

X X X

(29)

Japan Rocky intertidal macroinvertebrates and algae

1

1 1 5

X

X

(30)

  7  

Southeast Pacific (Chile mostly) Chilean litoral fishes

1

2

(31)

Portuguese intertidal Epifaunal crustaceans (those associated with macroalgae)

3

1 X X

X

(32)

Australia east coast Rocky intertidal animals (macro fauna)

1

4

T

X

T

(33)

Global Non-halacrid intertidal mites

(34)

Tropical Eastern Pacific (TEP) Shore fishes

1

1 2 3

X

X

(35)

Brazil Reef fish

1 1 1

1 4

X

X

(36)

Eastern Pacific Molluscs (gastropods and bivalves) shallower than 200 m

1

X

(37)

Mediterranean Sea Species associated with Cystoseira crinata assemblages

1 1 5

X X X X X

(38)

Western Indian Ocean - WIO (shallow water) Holothuroids (sea-cucumbers)

1 1

1 3

X X

(39)

New Zealand Rocky intertidal Macroalgae and invertebrates 1 2

2

5

1 X

X

X

(40)

N. Indian Ocean Macroalgae

4

T

X

X

(41)

NE Pacific Rocky intertidal Macroalgae and invertebrates

3 1 2

3

9 1 1 T X X T T T

(42)

  8  

New Zealand rocky intertidal

Rocky intertidal macroinvertebrates and algae

(43)

South Africa Macroalgae and invertebrates (220 species) 1

1

6

X

X

X X

(44)

Chilean coast Peracarid crustaceans (those associated with macroalgae)

1

3

X X

(45)

South Africa Coastal fishes

1 1 6

X X X

X X

(46)

S. Australia Macroalgae (1487 species from herbarium specimens)

1 4

1 X X X

X

(47)

 12   13  

  9  

1. Adey, W. H. and Steneck, R. S. 2001. Thermogeography over time creates 14  

biogeographic regions: A temperature/space/time-integrated model and an 15  

abundance-weighted test for benthic marine algae. — J. Phycol. 37: 677-698. 16  

2. Barnes, D. K. A. and De Grave, S. 2001. Ecological biogeography of southern polar 17  

encrusting faunas. — J. Biogeogr. 28: 359-365. 18  

3. Belanger, C. L. et al. 2012. Global environmental predictors of benthic marine 19  

biogeographic structure. — Proc. Natl. Acad. Sci. U. S. A. 109: 14046-14051. 20  

4. Blanchette, C. A. et al. 2008. Biogeographical patterns of rocky intertidal communities 21  

along the Pacific coast of North America. — J. Biogeogr. 35: 1593-1607. 22  

5. Bolton, J. J. et al. 2004. Where is the western limit of the tropical Indian Ocean 23  

seaweed flora? An analysis of intertidal seaweed biogeography on the east coast 24  

of South Africa. — Mar. Biol. 144: 51-59. 25  

6. Boschi, E. E. and Gavio, M. A. 2005. On the distribution of decapod crustaceans from 26  

the Magellan biogeographic province and the Antarctic region. — Sci. Mar. 69: 27  

195-200. 28  

7. Briggs, J. C. and Bowen, B. W. 2012. A realignment of marine biogeographic 29  

provinces with particular reference to fish distributions. — J. Biogeogr. 39: 12-30. 30  

8. Bustamante, R. H. and Branch, G. M. 1996. Large scale patterns and trophic structure 31  

of southern African rocky shores: The roles of geographic variation and wave 32  

exposure. — J. Biogeogr. 23: 339-351. 33  

9. Colton, M. A. and Swearer, S. E. 2012. Locating faunal breaks in the nearshore fish 34  

assemblage of Victoria, Australia. — Mar. Freshw. Res. 63: 218-231. 35  

  10  

10. Cruz-Motta, J. J. et al. 2010. Patterns of spatial variation of assemblages associated 36  

with intertidal rocky shores: A global perspective. — Plos One 5: e14354. 37  

11. Espinosa-Perez, M. D. and Hendrickx, M. E. 2006. A comparative analysis of 38  

biodiversity and distribution of shallow-water marine isopods (Crustacea : 39  

Isopoda) from polar and temperate waters in the East Pacific. — Belg. J. Zool. 40  

136: 219-247. 41  

12. Floeter, S. R. and Gasparini, J. L. 2000. The southwestern Atlantic reef fish fauna: 42  

composition and zoogeographic patterns. — J. Fish Biol. 56: 1099-1114. 43  

13. Floeter, S. R. et al. 2001. Geographic variation in reef-fish assemblages along the 44  

Brazilian coast. — Glob. Ecol. Biogeogr. 10: 423-431. 45  

14. Floeter, S. R. et al. 2008. Atlantic reef fish biogeography and evolution. — J. 46  

Biogeogr. 35: 22-47. 47  

15. Garcia, F. J. and Bertsch, H. 2009. Diversity and distribution of the Gastropoda 48  

Opisthobranchia from the Atlantic Ocean: A global biogeographic approach. — 49  

Sci. Mar. 73: 153-160. 50  

16. Genzano, G. N. et al. 2009. Hydroid assemblages from the Southwestern Atlantic 51  

Ocean (34-42 degrees S). — Mar. Ecol. 30: 33-46. 52  

17. Glynn, P. W. and Ault, J. S. 2000. A biogeographic analysis and review of the far 53  

eastern Pacific coral reef region. — Coral Reefs 19: 1-23. 54  

18. Harley, C. D. G. et al. 2003. Environmental variability and biogeography: the 55  

relationship between bathymetric distribution and geographical range size in 56  

marine algae and gastropods. — Glob. Ecol. Biogeogr. 12: 499-506. 57  

  11  

19. Hastings, P. A. 2000. Biogeography of the Tropical Eastern Pacific: distribution and 58  

phylogeny of chaenopsid fishes. — Zool. J. Linn. Soc. 128: 319-335. 59  

20. Haussermann, V. 2006. Biodiveristy of Chilean sea anemones (Cnidaria: Anthozoa): 60  

distribution patterns and zoogeographic implications, including new records for 61  

the fjoord region. — Investigaciones Marinas 34: 23-35. 62  

21. Hidas, E. Z. et al. 2007. Is the species composition of rocky intertidal invertebrates 63  

across a biogeographic barrier in south-eastern Australia related to their potential 64  

for dispersal? — Mar. Freshw. Res. 58: 835-842. 65  

22. Iken, K. et al. 2010. Large-scale spatial distribution patterns of echinoderms in 66  

nearshore rocky habitats. — Plos One 5: e13845. 67  

23. Ingolfsson, A. 1992. The origin of the rocky shore fauna of Iceland and the Canadian 68  

maritimes. — J. Biogeogr. 19: 705-712. 69  

24. Izquierdo, D. and Guerra-Garcia, J. M. 2011. Distribution patterns of the peracarid 70  

crustaceans associated with the alga Corallina elongata along the intertidal rocky 71  

shores of the Iberian Peninsula. — Helgol. Mar. Res. 65: 233-243. 72  

25. Kafanov, A. I. et al. 2000. Ichthyofaunistic biogeography of the Japan (East) Sea. — J. 73  

Biogeogr. 27: 915-933. 74  

26. Lancellotti, D. A. and Vasquez, J. A. 1999. Biogeographical patterns of benthic 75  

macroinvertebrates in the southeastern Pacific littoral. — J. Biogeogr. 26: 1001-76  

1006. 77  

27. Leaper, R. et al. 2012. Comparing large-scale bioregions and fine-scale community-78  

level biodiversity predictions from subtidal rocky reefs across south-eastern 79  

Australia. — J. Appl. Ecol. 49: 851-860. 80  

  12  

28. Liuzzi, M. G. et al. 2011. Latitudinal gradients in macroalgal biodiversity in the 81  

southwest Atlantic between 36 and 55 degrees S. — Hydrobiologia 673: 205-214. 82  

29. Meneses, I. and Santelices, B. 2000. Patterns and breaking points in the distribution 83  

of benthic algae along the temperate Pacific coast of South America. — Rev. Chil. 84  

Hist. Nat. 73: 615-623. 85  

30. Nakaoka, M. et al. 2006. Similarity of rocky intertidal assemblages along the Pacific 86  

coast of Japan: effects of spatial scales and geographic distance. — Ecol. Res. 21: 87  

425-435. 88  

31. Ojeda, F. P. et al. 2000. Biogeographic patterns of Chilean littoral fishes. — Rev. 89  

Chil. Hist. Nat. 73: 625-641. 90  

32. Pereira, S. G. et al. 2006. Biogeographic patterns of intertidal macroinvertebrates and 91  

their association with macroalgae distribution along the Portuguese coast. — 92  

Hydrobiologia 555: 185-192. 93  

33. Poloczanska, E. S. et al. 2011. Little change in the distribution of rocky shore faunal 94  

communities on the Australian east coast after 50 years of rapid warming. — J. 95  

Exp. Mar. Biol. Ecol. 400: 145-154. 96  

34. Proches, S. and Marshall, D. J. 2001. Global distribution patterns of non-halacarid 97  

marine intertidal mites: implications for their origins in marine habitats. — J. 98  

Biogeogr. 28: 47-58. 99  

35. Robertson, D. R. and Cramer, K. L. 2009. Shore fishes and biogeographic 100  

subdivisions of the Tropical Eastern Pacific. — Mar. Ecol. Prog. Ser. 380: 1-17. 101  

36. Rocha, L. A. 2003. Patterns of distribution and processes of speciation in Brazilian 102  

reef fishes. — J. Biogeogr. 30: 1161-1171. 103  

  13  

37. Roy, K. et al. 1994. Eastern Pacific Molluscan provinces and latitudinal diversity 104  

gradient - no evidence for "Rapoport's rule". — Proc. Natl. Acad. Sci. U. S. A. 91: 105  

8871-8874. 106  

38. Sales, M. et al. 2012. Biogeographical patterns of algal communities in the 107  

Mediterranean Sea: Cystoseira crinita-dominated assemblages as a case study. — 108  

J. Biogeogr. 39: 140-152. 109  

39. Samyn, Y. and Tallon, I. 2005. Zoogeography of the shallow-water holothuroids of 110  

the western Indian Ocean. — J. Biogeogr. 32: 1523-1538. 111  

40. Schiel, D. R. 2011. Biogeographic patterns and long-term changes on New Zealand 112  

coastal reefs: Non-trophic cascades from diffuse and local impacts. — J. Exp. 113  

Mar. Biol. Ecol. 400: 33-51. 114  

41. Schils, T. and Wilson, S. C. 2006. Temperature threshold as a biogeographic barrier 115  

in northern Indian Ocean macroalgae. — J. Phycol. 42: 749-756. 116  

42. Schoch, G. C. et al. 2006. Fifteen degrees of separation: Latitudinal gradients of 117  

rocky intertidal biota along the California Current. — Limnol. Oceanogr. 51: 118  

2564-2585. 119  

43. Shears, N. T. et al. 2008. Evaluation of biogeographic classification schemes for 120  

conservation planning: Application to New Zealand's coastal marine environment. 121  

— Conserv. Biol. 22: 467-481. 122  

44. Sink, K. J. et al. 2005. Biogeographic patterns in rocky intertidal communities in 123  

KwaZulu-Natal, South Africa. — Afr. J. Mar. Sci. 27: 81-96. 124  

45. Thiel, M. 2002. The zoogeography of algae-associated peracarids along the Pacific 125  

coast of Chile. — J. Biogeogr. 29: 999-1008. 126  

  14  

46. Turpie, J. K. et al. 2000. Biogeography and the selection of priority areas for 127  

conservation of South African coastal fishes. — Biol. Conserv. 92: 59-72. 128  

47. Waters, J. M. et al. 2010. Australia's marine biogeography revisited: Back to the 129  

future? — Austral. Ecol. 35: 988-992. 130  

  15  

Table A3. Summary of the modified mCART to account for geographic differences in 131  sampling effort, for all species and separately by each classification scheme. We were 132  not able to find larval development type for 27 species of animalia. 133   134  

Scheme (number of species) # Significant

breaks

Pseudo-R2

median (95% CI)

All Species (406) 5 0.77 (0.66-0.83) Animalia (235) 5 0.75 (0.65-0.84) Algae (171) 4 0.76 (0.65-0.83) Direct develop (43) 5 0.76 (0.62-0.87) Planktonic non-feeding larvae (85) 5 0.73 (0.62-0.83) Planktotrophic larvae (80) 4 0.69 (0.51-0.85)

135  

  16  

136  Table A4. Adjusted Rand Index between biogeographic classification schemes (see main text). Jackknifed 95%CI are indicated in 137  parentheses. 138   139  

All species

Algae

Animalia

Direct develop

Planktonic feeding larvae

Algae 0.971

(0.928-1.000)

Animalia 0.996 0.967

(0.989-1.000) (0.924-1.000)

Direct develop 0.703 0.716 0.705

(0.593-0.815) (0.609-0.825) (0.594-0.818)

Planktotrophic larvae 0.445 0.420 0.448 0.193

(0.358-0.537) (0.335-0.509) (0.359-0.542) (0.085-0.305)

Planktonic non-feeding larvae 0.889 0.871 0.891 0.734 0.372

(0.802-0.977) (0.779-0.965) (0.804-0.980) (0.632-0.839) (0.271-0.475)

140   141  

  17  

Table A5. Relative importance (pseudo-R2) of the 29 environmental variables used to predict NEP biogeographic structure according 142  

to random forest modeling for different classification schemes. Significant predictors (p <0.05) highlighted in bold. Also indicated is 143  

the variance inflation factor (VIF) for each variable (values >5 indicate strong collinearity of the predictor). UDSSI stands for 144  

upwelling/downwelling seasonal switch index (see main text). 145  

Predictor

VIF All Species

Animals

Algae

Direct develop

Planktonic non-feeding

Planktotrophic

Silicate 143.8 0.13 0.14 0.13 0.13 0.11 0.09 Nitrate 150.3 0.12 0.12 0.12 0.14 0.10 0.02 Sea Surface Temperature 37.6 0.12 0.12 0.12 0.13 0.12 0.04 Phosphate 84.8 0.11 0.11 0.12 0.13 0.10 0.01 UDSSI 9.2 0.07 0.07 0.07 0.09 0.08 0.01 Air Temperature 54.6 0.05 0.06 0.07 0.05 0.08 0.06 Salinity 5.0 0.05 0.05 0.04 0.01 0.06 0.12 Precipitation Seasonality 35.6 0.05 0.05 0.05 0.02 0.04 0.13 Mean Tide Range 51.6 0.04 0.05 0.03 0.00 0.04 0.10 Upwelling Index 39.4 0.04 0.05 0.05 0.03 0.02 0.10 Photosynthetically Available Radiation 34.4 0.04 0.04 0.03 0.03 0.03 0.09 Oxygen 29.7 0.03 0.03 0.03 0.04 0.02 0.03 Cloud Cover 18.1 0.02 0.02 0.03 0.01 0.01 0.04 Annual Precipitation 57.0 0.02 0.02 0.02 0.01 0.05 0.03 Primary Productivity 3.2 0.01 0.01 0.01 0.04 0.02 0.02 pH 31.6 0.01 0.01 0.01 0.04 0.02 0.03 LGM Ice Extent - 0.01 0.01 0.00 0.00 0.00 0.02 Temperature Seasonality 21.8 0.01 0.01 0.01 0.01 0.01 0.02

  18  

Human Population 2.4 0.01 0.01 0.00 0.00 0.00 0.01 Air Temperature Annual Range 13.1 0.00 0.00 0.00 0.01 0.01 0.01 Wave Run-up 45.1 0.00 0.00 0.00 0.00 0.00 0.00 Shelf Area 2.2 0.00 0.00 0.00 0.00 0.00 0.00 Calcite 3.5 0.00 0.00 0.00 0.00 0.00 0.01 Significant Wave Height 27.1 0.00 0.00 0.00 0.00 0.00 0.00 Intertidal Slope 15.8 0.00 0.00 0.00 0.00 0.00 0.00 Upper Biologic Limit 4.4 0.00 0.00 0.00 0.00 0.00 0.00 Diffuse Attenuation 26.6 0.00 0.00 0.00 0.00 0.00 0.00 Chlorophyll a 29.6 0.00 0.00 0.00 0.00 0.00 0.00 Transect length 5.3 0.00 0.00 0.00 0.00 0.00 0.00

  19  

Supplementary Material Figure Legends 146   147  Figure A1. Accounting for bias in the assessment of NEP biogeographic 148  

structure. A) Sampling frequency across the NEP. Each bar represents the sum 149  

of the number of sites and times sampled per 1º bin. Each site (n=102) was 150  

sampled once (n=37), twice (n=51) or three times (n=14). Blue dotted lines 151  

show the position of 8 breaks identified by the bias-uncorrected mCART 152  

shown in B). C) The bias-corrected procedure showing the breaks identified 153  

by 10,000 runs of the mCART. D) A single realization of the corrected 154  

mCART. 155  

156  

157  

Figure A2. Bias corrected assessment of the biogeographic structure of NEP 158  

species according to different groupings. A) All species combined, B) 159  

Animalia, C) Algae, D) Direct develop, E) Planktonic non-feeding larvae, F) 160  

Planktonic feeding (Planktotrophic) larvae. The locations of significant 161  

biogeographic breaks are depicted as peaks (ºN in red).162  

  20  

Figure A1.

  21  

Figure A2.