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Electronic Supplementary Material (ESM) for:
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The high fidelity of the cetacean stranding record: insights into measuring diversity by integrating
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taphonomy and macroecology
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Nicholas D Pyenson1,2*
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Department of Paleobiology, National Museum of Natural History, Smithsonian Institution,
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Washington, DC 20013, USA
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Departments of Mammalogy and Paleontology, Burke Museum of Natural History and Culture, Box
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353010, University of Washington, Seattle, WA 98195-3010, USA.
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*Author for correspondence (pyensonn@si.edu)
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Table of Contents:
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1. Supplementary Materials and Methods
p. 2
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2. Supplementary Table and Figure Captions
p. 8
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3. Supplementary Table S1
p. 10
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4. Supplementary Table S2
p. 11
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5. Supplementary Table S3
p. 12
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6. Supplementary Figure S1
p. 13
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7. Supplementary Figure S2
p. 14
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8. Supplementary Figure S3
p. 15
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9. References for Supplementary Material
p. 16
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1. Supplementary Materials and Methods
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(a) Strandings and sightings datasets
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I collected strandings and sightings data from publicly archived or available datasets from multiple
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coastlines around the world. Data were collected in their raw format, which was usually represented in a
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spreadsheet format. In such cases, datasets were arranged in rows by finely resolved taxa (e.g., species)
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and organized into columns over time intervals (i.e., consecutive years). For some raw datasets, I
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removed categories that were too broad or taxonomically imprecise. When such categories represented
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large numerical abundances, their deletion from the pooled datasets was justified because it did not
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affect the presence or absence of individual taxa. Taxonomic conventions follow those presented in each
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dataset, despite some determinations (e.g., certain delphinid species) that were outdated or paraphyletic.
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Raw data from both dead (strandings) and live (sightings) datasets were considered as
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occurrence data (i.e., presence/absence data). Absence from these compilations should not be considered
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as true, ecological absences, merely as samples of diversity within specific spatiotemporal settings.
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Following Pyenson [S1], raw dead and live data were stratified into three sets that approximately reflect
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three taxonomic hierarchies: species-, genus-, and family-level data. Beginning with the raw data, I
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pooled all permutations of different species groupings into species level groups. I deleted any category
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qualified with ‘‘unidentified.’’ Then, I pooled all species, including unidentified species from known
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genera into their respective genera. Similarly, I deleted any genus level category qualified by
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‘‘unidentified.’’ Lastly, I pooled all genera, including unidentified genera belonging to known families,
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into their respective families. Suprafamilial- to subordinal-level data are essentially uninformative for
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the purposes of this study, and they were disregarded.
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Below, I elaborate on the data sources for each country’s coastline, in alphabetical order.
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(i) Australia. Both dead and live data for Australia were collected from the National Whale and Dolphin
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Sightings and Strandings Database (available online at http://data.aad.gov.au/aadc/whales/) of the
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Australian Government’s Department of Environment and Water Resources. These data are organized
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on a species account basis, ranging over different ranges of years (sometimes non-consecutively).
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Occurrence data from “unknown years” were vetted and entered if they were “collected, observed” or
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provided any specificity solely relating to the year in which the occurrence happened.
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(ii) Galapagos Islands (Ecuador). Death assemblage data for the Galapagos Islands derives from data
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published in Palacios et al. 2004 [S2]. I adjusted the dataset to exclude all pre-1971 strandings data, as a
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measure to match the time interval covered by the live survey data from Palacios 2003 [S3: table A4.1].
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This necessary step meant deleting all pre-1971 records from table 2 in Palacios et al. 2004 [S2], and
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verified against figure 1 in the same publication. Ultimately this modification deleted mostly records of
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delphinid strandings, but it also removed the occurrence of Mesoplodon gingkodens, a rare ziphiid that
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only occurred in the stranding data. This pre-1971 deletion procedure depressed the total richness of the
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death record slightly, and voided any presence data for Pseudorca crassidens and all Mesoplodon spp. in
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the Galapagos Islands. Only cumulative strandings data were available over the time interval matching
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all sightings data, and thus these data could not be included in the rarefaction analyses.
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(iii) Greece and the Greek Island Archipelago. Data from the Greek stranding record (including the
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Ionian, Aegean, Cretan and northwest Levantine seas and the northern Cretan Passage) were tabulated
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from [S4], with some modifications and updates to the dataset indicated by the corresponding author, A.
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Frantzis, following personal correspondence via email in 2010. I followed the extrapolation in [S4: table
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1] for the three most abundant delphinid species, but restricted the temporal coverage of these data from
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1991 to 2001. Ref. [S4] also provided the live data as sightings in table 1 of that document.
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(iv) Ireland. All of the dead and live data from Ireland are available online through the public database
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maintained by the Irish Whale and Dolphin Group (IWDG), a non-profit private company and registered
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charity. All records are validated (and available online at: http://www.iwdg.ie). Because many of the
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entries on the database used common names that did not necessarily identify taxonomic occurrences to
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the species level, I re-grouped the dataset according to taxonomic categories that were different than
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those available on the IWDG database. Also, I restricted the database searches to the following
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geographic regions: Porcupine Bank, Celtic Sea, Irish Sea, North Channel, North Coast, Northwest and
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Southwest regions, St. George's Channel, and West Coast. Only cumulative strandings data were
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available over the time interval matching all sightings data, and thus Ireland’s data could not be included
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in the rarefaction analyses.
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(v) The Netherlands. For The Netherlands, both dead and live data were collected from the Dutch
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Seabird Group (NZG) Marine Mammal Database database originally created by C. J. (Kees)
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Camphuysen (available online at: http://home.planet.nl/~camphuys/Cetacea.html). The (NZG) Marine
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Mammal Database collected all strandings and sightings of marine mammals in The Netherlands and the
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Southern North Sea starting in 1971. Personal correspondence with Kees Camphuysen in 2010 indicated
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that Camphuysen no longer maintains the dataset, and the dataset continues to be maintained by the
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Dutch stranding program of Naturalis in Leiden (see http://www.walvisstrandingen.nl/). Some of the
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dead and live data were compiled in van der Meij & Camphuysen 2006 [S5: table 2]. Total sightings
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data were collected from the NZG Marine Mammal Database posted on Camphuysen’s website, and
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strandings data were collected from a spreadsheet formatted from data on the same site, sent via
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personal correspondence with Camphuysen in 2010.
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(vi) New Zealand. Death assemblage data from New Zealand derived from the Department of
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Conservation (DOC)’s marine mammal stranding database, which is administered by A. van Helden at
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the Museum of New Zealand, Te Papa Tongarewa. Only strandings data from New Zealand were
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available; no sightings data exist because of patchy, local surveys that account for singular species and
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non-existent platform coverage across the entirety of the North and South islands. Data were collected
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via personal email request to DOC and A. Van Helden in 2009.
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(vii) United States. Dead and live data for the US Pacific coast were collected and organized by Pyenson
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[S1]. See references and details therein for the assembly of that particular dataset, including the
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compilation of live survey data from multiple sources.
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Death assemblage data for the northeastern US Atlantic coast were collected from the Northeast
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Regional Office of NOAA’s National Marine Fisheries Service Marine Mammal Stranding Network.
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Live survey data from the same region were collected by Palka 2006 [S6: table 7]. These data were
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collected using both aerial and ship-based survey platforms in US Navy’s Northeast (NE) operating area
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(OPAREA), which is a geographic area broadly consistent with the Northeast Stranding Region (see
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[S6: figure 1]). Live occurrence data were reported as summary of abundance estimates for all regions,
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years (1998-2004) and species, reported in table 7 of that document.
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(b) Taxonomic accumulation curves
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As a form of sample-based rarefaction, EstimateS computes taxonomic accumulation curves based on
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iterative resampling of separate collections of richness data (i.e., stranding occurrences). The actual
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accumulation curves are the result of Monte Carlo resampling from the stranding dataset, which
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randomly accumulates reiterations of the original stranding occurrences. Sample-based rarefaction is
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more appropriate for the time-series stranding data in this study than standard individual rarefaction
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analyses (e.g., analytical rarefaction) because the latter samples from a single pool of data without
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replacement [S7-S8]. By resampling with replacement, EstimateS smoothes collection curves and
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provides confidence intervals for comparison with other collection curves. The input data file for
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EstimateS contained cetacean stranding occurrences for a given stranding network region that were
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grouped by year and sorted by taxonomic level. In EstimateS, I selected nominal parameters, including
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50 randomization runs, a strong hash encryption for the random number generator, and randomization
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without replacement.
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(c) Coastline lengths
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Measuring coastline length has long posed a significant mathematical problem [S9] because their vector
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paths are essentially fractal. Thus, different methods exist to measure the total coastline of a given
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country. Maps of larger regions will tend to simplify coastlines for economic use of map space, whereas
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maps of island archipelagos tend to show features in more detail out of navigational necessity. This
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differential thus affects comparisons of coastline lengths for countries with maps at different scales.
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Published accounts of coastline lengths (e.g., [S10]) rarely identify whether measurements were derived
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from single sources with constant scale, which is a necessary control for comparing coastline lengths. I
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collected coastline length data from Coastal and Marine Ecosystems — Marine Jurisdictions section of
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the World Resources Institute’s EarthTrends database [S11], except for Australia and its states, which
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relied on data from the GEODATA Coast 100K 2004 database. These coastline lengths are published by
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Geoscience Australia [S12], and they are nationally uniform because they derived from the 1:100 000
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scale National Topographic Map Series. Lastly, the Galapagos Islands coastline length derives from an
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estimation provided by the Galapagos Conservancy [S13].
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2. Supplementary Table and Figure Captions.
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Supplementary Table S1. Total compilations for living and dead species in each dataset for coastlines in
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this study. Summary includes totals both with and without New Zealand’s death record.
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Supplementary Table S2. Coastline lengths for countries (including both US coasts) used in this
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analysis. Coastline lengths in kilometers (km). See Supplementary References for data sources.
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Supplementary Table S3. Spearman rank order correlation tests on ranked relative abundance values
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from living and death assemblages from Queensland, Western Australia and all of Australia.
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Results are grouped by taxonomic rank, and by coastline; coefficient represents values for
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Spearman's r; and p values are one-tailed. Bold indicates values p > 0.01, and represent
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correlations that are not significant.
Supplementary Figure S1. Compositional fidelity metrics for live-dead records across the globe, at the
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(a) species, (b) genera and (c) family levels. Part (b) is a reproduction of Figure 1 from the main
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text. Light gray indicates live-dead (LD) values; medium gray indicates dead-live (DL) values;
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and dark gray indicates abundance-corrected dead-live values (DLa). Countries are ranked, left
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to right, in increasing coastline size; see main text for further details.
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Supplementary Figure S2. Taxonomic accumulation curves generated from sample-based rarefaction
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curves of stranding data, at the (a) species, (b) genera and (c) family levels. Stranding data from
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Ireland and the Galapagos Islands could not be used in this analysis, although New Zealand’s
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stranding data was included.
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Supplementary Figure S3. Rank-order family level abundances of cetaceans for both living and death
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assemblages from Australia. Live-dead pairs are ranked, top to bottom, in increasing coastline
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size. Abundance values are percentages of total number of sighting occurrences (live) or total
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number of stranding occurrences (dead), in both cases reflecting relative abundance; bold
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indicates zero values. From top to bottom as follows: (a) Queensland; (b) Western Australia; (c)
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all of Australia (data from Figure 2g).
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3. Supplementary Table S1.
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Coastline
Australia
Greece
Galapagos Is.
Ireland
Netherlands
New Zealand
US Atlantic
US Pacific
Total species (with NZ)
Total species (without NZ)
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Live
1522
681
2879
10635
18250
not applicable
631405
618616
1283988
1283988
Dead
1117
354
86
1336
3329
2108
2434
2083
12847
10739
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4. Supplementary Table S2.
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Country
Australia
New Zealand
Greece
Ireland
US Pacific Coast
Netherlands
Galapagos Is.
US Atlantic Coast
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Coastline length (km)
59736
17209
15147
6437
2081
1912
1609
1605
Source
Ref. [S12]
Ref. [S11]
Ref. [S11]
Ref. [S11]
Ref. [S1]
Ref. [S11]
Ref. [S13]
Ref. [S14]
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5. Supplementary Table S3.
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Queensland
Western Australia
Coefficient
p value
n
Coefficient
p value
n
Species
0.568
<0.001
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0.471
0.0013
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Genera
0.317
0.0659
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0.477
0.0093
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Families
0.571
0.0901
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0.857
0.0069
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Australia (all states)
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Coefficient
p value
n
Species
0.411
0.0034
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Genera
0.422
0.0159
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Families
0.595
0.0598
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6. Supplementary Figure S1.
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7. Supplementary Figure S2.
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8. Supplementary Figure S3.
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9. Supplementary References
S1
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stranding record in eastern North Pacific Ocean. Paleobiol. 36, 453– 480.
S2
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Pyenson, N. D. 2010 Carcasses on the coast: measuring the ecological fidelity of the cetacean
Palacios, D. M., Salazar, S. K. & Day, D. 2004 Cetacean remains and strandings in the
Galápagos Islands, 1923-2003. Latin Amer. J. Aq. Mamm. 3, 127-150.
S3
Palacios, D. M. 2003 Oceanographic conditions around the Galápagos Archipelago and their
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influence on cetacean community structure. Unpublished Ph.D. Thesis, Oregon State University,
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Corvallis, Oregon. Available online http://www.pfeg.noaa.gov/~dpalacio/pubs.html
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S4
Frantzis, A., Alexiadou, P., Paximadis, G., Politi, E., Gannier, A. & Corsini-Foka, M. 2003
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Current knowledge of the cetacean fauna of the Greek Seas. J. Cetacean Res. Manag. 5, 219-
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232.
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S5
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dolphins (Cetacea) in the southern North Sea: 1970–2005. Lutra 49, 3–28.
S6
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S7
S8
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Gotelli, N. & Colwell, R. K. 2001 Quantifying biodiversity: procedures and pitfalls in the
measurement and comparison of species richness. Ecol. Lett. 4, 379–391.
S9
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Colwell, R. K., & Coddington, J. A. 1994 Estimating terrestrial biodiversity through
extrapolation. Phil. Trans. Roy. Soc. London B 345, 101–118.
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Palka, D. L. 2006 Summer abundance estimates of cetaceans in US North Atlantic Navy
Operating Areas. US Dep. Commer., Northeast Fish. Sci. Cent. Ref. Doc. 06-03, 1-41.
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18
van der Meij, S. E. T. & Camphuysen, C. J. 2006 Distribution and diversity of whales and
Mandelbrot, B. 1967 How long is the coast of Britain? Statistical self-similarity and fractional
dimension. Science 156, 636-638.
S10
US CIA World Factbook. Available online https://www.cia.gov/library/publications/the-worldfactbook/
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S11
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World Resources Institute. 2010 EarthTrends Database. Persistent URL
http://earthtrends.wri.org
S12
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Geoscience Australia. Coastline lengths. Available online
http://www.ga.gov.au/education/geoscience-basics/dimensions/coastline-lengths.html
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S13
Galapagos Conservancy. 2010 Galapagos Map (PDF file). Downloaded 16 June 2010.
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S14
NOAA (National Oceanic and Atmospheric Administration). 1975 The coastline of the United
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States. Washington: U.S. Government Printing Office.