www.sciencemag.org/cgi/content/full/317/5837/499/DC1

Supporting Online Material for

A Cambrian Peak in Morphological Variation Within Trilobite Species

Mark Webster*

*E-mail: mwebster@geosci.uchicago.edu

Published 27 July 2007, Science 317, 499 (2007)

DOI: 10.1126/science.1142964

This PDF file includes:

Materials and Methods SOM Text Figs. S1 to S9 Tables S1 and S2 References

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SUPPORTING ONLINE MATERIAL

MATERIALS AND METHODSPrevious attempts to investigate temporal trends in intraspecific variation have oftenfocused on single components of anatomy (e.g., thoracic segment number, cranidialdimensions; see supporting text) and are therefore limited in morphological coverage.Multivariate morphometric techniques allow a more comprehensive summary of shape,but can suffer from methodological difficulties when applied to disparate morphologies(e.g., problems of landmark homology and of distortion in projecting data from shapespace into tangent space when comparing disparate forms using geometricmorphometrics). Here I explore temporal trends in intraspecific variation as documentedby changes in the frequency and extent of character states coded as polymorphic withintrilobite species in cladistic analyses. This approach minimizes limits on bothmorphological coverage (because continuously varying, meristic, or discretemorphological attributes can be coded as cladistic characters in any analysis) andphylogenetic coverage (since disparate morphologies are often accommodated withincladistic analysis by employing qualitative character states). Furthermore, characters andcharacter state limits utilized in cladistic analyses must have been deemed to bepotentially phylogenetically informative by the original authors: when coding, expressionof alternate character states (between or within species) was implicitly assumed to beunder heritable control and not a result of ecophenotypy. This approach therefore permitsa broad, initial assessment of trends in intraspecific variation in morphological features ofrelevance to evolution. The results of this analysis encourage further investigation intrilobites and other clades, which could employ a more refined methodology (e.g.,geometric morphometrics) at less inclusive phylogenetic scale.

Criteria for Inclusion in the Meta-analysisSixty-eight trilobite cladistic character-taxon matrices were examined for intraspecificpolymorphisms (Table S1). Prior to conducting the meta-analysis, however, stringentcriteria were imposed to reduce the potential for among-study bias in the prevalence ofintraspecific polymorphisms. In order for any given character-taxon matrix to beincluded, the cladistic analysis had to: (A) be at a fine taxonomic level (i.e.,hypothesizing the relationship of species within genera or families); (B) be basedprimarily on mature morphology; and (C) code intraspecific polymorphisms as such (notas missing data, the most common state, etc.; analyses that explicitly excludedpolymorphic characters were not included in the present meta-analysis). Non-independentdata sets (i.e., studies involving a revision of character state coding, or addition of taxa orcharacters, to a previously published matrix) were merged.

Even when a study met these criteria, not all operational taxonomic units (OTUs)coded within the character-taxon matrix were necessarily appropriate for inclusion in themeta-analysis. Additional criteria were applied to further improve the rigor of the dataset. OTUs within a character-taxon matrix were included only if they (D) were genuinespecies (i.e., “Frankenstein” OTUs [constructed from character state data from severalspecies] and hypothetical ancestors were excluded, and subspecies OTUs were lumpedinto species-level OTUs); and (E) were coded from examination of more than onespecimen (so that any intraspecific polymorphism could be potentially detected). Where

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the sample size used for coding a given species was not listed, the systematic literaturefor that species was examined to determine whether more than one specimen was figuredand thus available for coding. However, in several cases this latter criterion had to beassumed. Data for species occurring in multiple studies were merged. Missing,inapplicable, and redundant (when the same species is coded for the same character inmultiple studies) data were excluded.

In rare cases, particular characters had to be excluded from otherwise appropriatecharacter-taxon matrices because the character state definitions mask potentially“significant” intraspecific variation, as when the distinction between the definedcharacter states is smaller than the range of variation permitted within the definedcharacter states (Table S1).

Character state coding for species was not modified herein except for three cases(studies 26a, 32, and 48 in Table S1) in which polymorphisms within some OTUs hadbeen overlooked by the original authors.

Application of these criteria led to the rejection of entire matrices, or to theexclusion of particular characters and/or taxa from some matrices (Table S1). Theremaining data (Table S2), consisting of over 40,000 positively coded character states for982 trilobite species, offers the opportunity to explore temporal change in the frequencyand extent of intraspecific polymorphism. Of course, characters exhibiting “excessive”levels of intraspecific variation are sometimes excluded from cladistic analyses altogetherwithout comment (but see S1), and the full extent of intraspecific variation may not havebeen sampled during character state coding. The frequency of polymorphic cells withinthe culled character-taxon matrices therefore represents a minimum estimate of the extentof intraspecific variation in characters selected for phylogenetic analysis. There is noreason to believe that such an estimate is temporally biased, however, and the patterndetected here appears to be robust.

SUPPORTING TEXTEmpirical support for a decrease in morphological variation in trilobites through time israther limited: intraspecific variation in the number of thoracic segments at maturity,although rare, has been more frequently documented among Cambrian than among post-Cambrian trilobites (S2-S6); and the degree of intraspecific variation in exoskeletalmorphology of the Late Cambrian Dikelocephalus minnesotensis is much greater than inthe Devonian Eldredgeops rana (S7). Several studies find opposite conclusions: nosignificant temporal decline was detected among nine trilobite species ranging from EarlyCambrian to Silurian in age in terms of either intraspecific variation in cranidial shape(S8) or fluctuating asymmetry, a measure of developmental precision (S9); andintraspecific variability in two cranidial dimensions was found to slightly butsignificantly increase among 17 species over a short time span within the Late Cambrian(S10, but see S11).

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SUPPORTING FIGURES

Fig. S1: Diversity (A), origination rate (B), and extinction rate (C) of trilobite generathrough the Paleozoic. Geologic period abbreviations: Cm, Cambrian; O, Ordovician; S,Silurian; D, Devonian; C, Carboniferous; P, Permian. Ma, millions of years beforepresent; myr, million years. Data from ref. S12. Per taxon origination and extinction ratescalculated following ref. S13. Dashed vertical lines correspond to divisions betweentemporal bins employed in the present analysis.

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Fig. S2: Frequency distributions for the proportion of characters coded as polymorphicwithin trilobite species, grouped by time. A, Early Cambrian (median = 2.8%); B, MiddleCambrian (median = 3.6%); C, Late Cambrian (median = 0.0%); D, Early Ordovician(median = 0.0%); E, Middle Ordovician (median = 0.0%); F, Late Ordovician (median =0.0%); G, Early Silurian (median = 0.0%); H, Early Devonian (median = 0.0%); I,Middle Devonian (median = 0.0%); J, Late Devonian (median = 0.0%). Nopolymorphisms are coded among sampled Late Silurian species (n = 28) and post-Devonian species (n = 23). Data from Table S2.

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Fig. S3: Temporal pattern of the frequency (left plots) and relative proportion (right plots)of redlichiid (A, B), ptychopariid (C, D), and agnostid (E, F) trilobite species coded aspolymorphic in at least one character in cladistic analyses. Hatched and solid shading inA, C, and E denotes polymorphic and non-polymorphic species, respectively. Error barsindicate 2-unit profile likelihood confidence intervals. A high proportion of specieswithin each of these phylogenetically early diverging and/or stratigraphically old ordersexhibits some degree of polymorphism. Poor sampling in some time bins renders anytemporal trends within the orders difficult to identify. Nevertheless, a significantly lowerproportion of the sampled Late Cambrian (25%), Early Ordovician (56%), and MiddleOrdovician (44%) ptychopariid species are variable in at least one character relative tothe Middle Cambrian (almost 90%); and the proportion of agnostid species exhibitingpolymorphism significantly declined from the Early (100%) to Middle Cambrian (77%).Data from Table S2. Abbreviations: E, early; M, middle; L, late.

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Fig. S4: Temporal pattern of the frequency (left plots) and relative proportion (right plots)of corynexochid (A, B), lichid (C, D), and asaphid (E, F) trilobite species coded aspolymorphic in at least one character in cladistic analyses. Hatched and solid shading inA, C, and E denotes polymorphic and non-polymorphic species, respectively. Error barsindicate 2-unit profile likelihood confidence intervals. A lower proportion of specieswithin each of these phylogenetically more derived and/or stratigraphically youngerorders (relative to the basal and/or older orders in Fig. S3) exhibits some degree ofpolymorphism. Temporal trends within the orders cannot be identified with confidencedue to poor representation in most time bins. None of the sampled burlingiid species(Middle Cambrian, n = 11; Late Cambrian, n = 3) have been coded with a polymorphism.Data from Table S2. Geologic period abbreviations: Ordov., Ordovician; Sil., Silurian;Dev., Devonian; E, early; M, middle; L, late.

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Fig. S5: Temporal and phylogenetic distribution of trilobite species exhibitingpolymorphism in characters relating to the shape of glabellar lobes (including the depth,orientation, and curvature of glabellar and axial furrows) only. Characters relating toglabellar ornamentation are not included. A, B; Temporal pattern of the frequency (A)and relative proportion (B) of trilobite species coded as polymorphic in at least onecharacter in cladistic analyses. C, D; Differences among trilobite orders in the frequency(C) and relative proportion (D) of species coded as polymorphic in at least one characterin cladistic analyses. Hatched and solid shading in A and C denotes polymorphic andnon-polymorphic species, respectively. Temporal and phylogenetic patterns detectedwhen the meta-analysis is restricted to characters relating to the shape of glabellar lobes(structures deemed homologous across all trilobite clades) are similar to those detectedwhen all characters are considered (compare this figure to Figs. 1, 3A, B). The findingsof the meta-analysis are therefore unlikely to reflect bias in morphological coverageacross clades or studies. Error bars indicate 2-unit profile likelihood confidence intervals.Data from Table S2. Abbreviations as for Fig. S4; Carb., Carboniferous.

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Fig. S6: Temporal and phylogenetic distribution of trilobite species exhibitingpolymorphism in characters relating to the presence or absence of aspects of cephalicornamentation only. Characters with state definitions such as “weak or absent” were notincluded. Multistate characters with one state defined as “absent” and other statesdescribing various conditions relating to “present” were included, but intraspecificpolymorphisms were accepted as such only if the polymorphism related to the “absent”state in addition to one (or more) “present” state. A, B; Temporal pattern of the frequency(A) and relative proportion (B) of trilobite species coded as polymorphic in at least onecharacter in cladistic analyses. C, D; Differences among trilobite orders in the frequency(C) and relative proportion (D) of species coded as polymorphic in at least one characterin cladistic analyses. Hatched and solid shading in A and C denotes polymorphic andnon-polymorphic species, respectively. Presence/absence coding in features of cephalicornamentation offers minimal ambiguity in state definition and recognition, and thereforeminimal potential for among-worker difference in character coding. Althoughintraspecific variation in the presence or absence of cephalic ornamentation is relativelyuncommon, broad temporal and phylogenetic patterns detected when the meta-analysis isrestricted to these characters are similar to those detected when all characters areconsidered (compare this figure to Figs. 1, 3A, B), suggesting that the findings of themeta-analysis are unlikely to reflect bias in character state definition or coding protocolacross clades or studies. Error bars indicate 2-unit profile likelihood confidence intervals.Data from Table S2. Abbreviations as for Fig. S4; Carb., Carboniferous.

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Fig. S7: Temporal and phylogenetic distribution of trilobite species exhibitingpolymorphism in at least one character, using character-taxon matrices in papers authoredby B. D. E. Chatterton. A, B; Temporal pattern of the frequency (A) and relativeproportion (B) of trilobite species coded as polymorphic in at least one character incladistic analyses. C, D; Differences among trilobite orders in the frequency (C) andrelative proportion (D) of species coded as polymorphic in at least one character incladistic analyses. E, F; Temporal pattern of the frequency (E) and relative proportion (F)of proetid trilobite species coded as polymorphic in at least one character in cladisticanalyses. Hatched and solid shading in A, C, and E denotes polymorphic and non-polymorphic species, respectively. The significant post-Early Ordovician drop inproportion of species exhibiting polymorphism, and the high proportion of polymorphicptychopariid species relative to phacopids and lichids, matches the patterns detectedwhen all authors are considered (Figs. 1, 3A, B). The high proportion of proetid speciesexhibiting polymorphism is not seen when all authors are considered, but likely resultsfrom this author’s focus on relatively early members of this long-lived clade (E, F;compare to Figs. 3C, D). Data from studies 2, 3, 5a, 7a, 15, 16, 17, 21, 24, 27, and 36 inTables S1, S2. Error bars indicate 2-unit profile likelihood confidence intervals.Abbreviations as for Fig. S4; Carb., Carboniferous.

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Fig. S8: Temporal and phylogenetic distribution of trilobite species exhibitingpolymorphism in at least one character, using character-taxon matrices in papers authoredby B. S. Lieberman. A, B; Temporal pattern of the frequency (A) and relative proportion(B) of trilobite species coded as polymorphic in at least one character in cladisticanalyses. C, D; Differences among trilobite orders in the frequency (C) and relativeproportion (D) of species coded as polymorphic in at least one character in cladisticanalyses. Hatched and solid shading in A and C denotes polymorphic and non-polymorphic species, respectively. The concentration of polymorphic species in the EarlyCambrian relative to the mid and late Paleozoic, and in the phylogenetically basalredlichiid clade relative to the later and more derived proetids and phacopids, matches thepatterns detected when all authors are considered (Figs. 1, 3A, B). The lack ofpolymorphism within proetid and phacopid species likely results from this author’s focuson relatively late members of these long-lived clades (compare A and B to Figs. 3C-F).Data from studies 6, 8, 10, 11, 12, 13, 14, 20, 26a (not 26b), 34, and 42 in Tables S1, S2.Error bars indicate 2-unit profile likelihood confidence intervals. Abbreviations as forFig. S4; Carb., Carboniferous.

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Fig. S9: Temporal pattern of the frequency (A) and relative proportion (B) of trilobitespecies coded as polymorphic in at least one character in cladistic analyses, restrictinganalysis to characters relating to cranidial outline only. Characters relating to glabellarshape are not included. Hatched and solid shading in A denotes polymorphic and non-polymorphic species, respectively. Temporal patterns detected here are similar to thosedetected when all characters are considered (compare to Fig. 1). The rate of increase indisparity of cranidial shape was highest in the Early Cambrian and slowed from theMiddle Cambrian through Middle Ordovician (fig. 5 in ref. S14), suggesting that rate ofmorphological diversification and frequency of intraspecific variation may be coupled.Error bars indicate 2-unit profile likelihood confidence intervals. Data from Table S2.Abbreviations as for Fig. S4; Carb., Carboniferous.

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SUPPORTING TABLESTable S1: Trilobite cladistic character-taxon matrices examined during the present study.After application of bias-minimizing criteria (see materials and methods), studiesnumbered 1 through 49 were appropriate for inclusion in the present meta-analysis.Justification for the exclusion and/or modification of original matrices is given in theright-hand column. Pending publication, the matrix for study 26b is available from theauthor by request (see also ref. S15).

StudyCode Reference Comments

1N. C. Hughes, A. W. A. Rushton, Palaeontology 33,429 (1990).

2J. M. Adrain, B. D. E. Chatterton, J. Paleontology 64,600 (1990).

Character 17 excluded (variation within states exceedsdifference between states).

3G. D. Edgecombe, B. D. E. Chatterton, Can. J. EarthSci. 27, 820 (1990). Erratencrinurus excluded ("Frankenstein" OTU).

4L. Ramsköld, Trans. R. Soc. Edin. Earth Sci. 82, 143(1991).

Hypothetical ancestor, Lichakephalidae,Ceratocephala, Koneprusia, Miraspis, Selenopeltis,Selenopeltoides (all "Frankenstein" OTUs) andDicranurus (single specimen) excluded. Duplicatecoding with study 21 excluded.

5aL. Ramsköld, B. D. E. Chatterton, Trans. R. Soc.Edin. Earth Sci. 82, 333 (1991).

Subspecies merged. Ketternaspis williamsi A and B("stratigraphically separate morphotypes") treated asdistinct species.

5bM. C. Ebach, Rec. W. Austr. Mus. 20, 353 (2002).[Leonaspis study]

Single species added to matrix of study 5a.

6B. S. Lieberman, G. D. Edgecombe, N. Eldredge, J.Paleontology 65, 824 (1991).

Bouleia, Parabouleia ("Frankenstein" OTUs) andMalvinella australis (single specimen) excluded.Duplicate coding with study 8 excluded.

7aG. D. Edgecombe, B. D. E. Chatterton, J.Paleontology 66, 52 (1992).

Encrinurus ("Frankenstein" OTU) and Anticostispecies (single specimen) excluded.

7b

M. C. Ebach, G. D. Edgecombe, in Fossils,Phylogeny, and Form: An Analytical Approach, J. M.Adrain, G. D. Edgecombe, B. S. Lieberman, Eds.(Topics In Geobiology, vol. 19, KluwerAcademic/Plenum, New York, 2001), pp. 235-289.[Distyrax study]

Three species (but Distyrax quinquecostata = singlespecimen) and four characters added to matrix of study7a.

8 B. S. Lieberman, J. Paleontology 67, 549 (1993). Punillaspis sp. excluded ("Frankenstein" OTU).

9aL. Ramsköld, G. D. Edgecombe, Pal. Zeit. 68, 89(1994).

Hypothetical ancestor excluded.

9b

M. C. Ebach, G. D. Edgecombe, in Fossils,Phylogeny, and Form: An Analytical Approach, J. M.Adrain, G. D. Edgecombe, B. S. Lieberman, Eds.(Topics In Geobiology, vol. 19, KluwerAcademic/Plenum, New York, 2001), pp. 235-289.[Wallacia study]

Two species added to matrix of study 9a (but Wallacian. sp. coded for only 16 characters, so excluded).

10B. S. Lieberman, Bull. Am. Mus. Nat. Hist. 223(1994). [Proetinae study]

Duplicate coding with studies 11, 12, and 13 excluded.

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11B. S. Lieberman, Bull. Am. Mus. Nat. Hist. 223(1994). [Crassiproetus study]

Crassiproetus norwoodensis excluded (singlespecimen).

12B. S. Lieberman, Bull. Am. Mus. Nat. Hist. 223(1994). [Basidechenella study]

Basidechenella witherspooni and B. cartwrightaeexcluded (single specimens).

13B. S. Lieberman, Bull. Am. Mus. Nat. Hist. 223(1994). [Dechenella study]

Pedinodechenella modelli excluded (single specimen).

14B. S. Lieberman, Bull. Am. Mus. Nat. Hist. 223(1994). [Monodechenella study]

"Milesdavis" zlichovianus excluded (single specimen).

15 B. D. E. Chatterton, J. Paleontology 68, 541 (1994). Duplicate coding with study 27 excluded.

16J. M. Adrain, B. D. E. Chatterton, J. Paleontology 68,305 (1994).

Character 12 excluded (variation within states exceedsdifference between states).

17aP. J. Lespérance, S. Desbiens, Pal. Soc. Mem. 42(1995).

17bG. D. Edgecombe, B. D. E. Chatterton, B. G.Waisfeld, N. E. Vaccari, J. Paleontology 72, 678(1998).

Five coding revisions, three additional characters, andtwo species added to matrix of study 17a. Duplicatecoding with study 3 excluded.

18S. R. Westrop, R. Ludwigsen, C. H. Kindle, J.Paleontology 70, 804 (1996).

Duplicate coding with study 47 excluded.

19aG. D. Edgecombe, L. Ramsköld, Geobios 29, 209(1996).

Hypothetical ancestor and Pacificurus ("Frankenstein"OTU) excluded.

19b

M. C. Ebach, G. D. Edgecombe, in Fossils,Phylogeny, and Form: An Analytical Approach, J. M.Adrain, G. D. Edgecombe, B. S. Lieberman, Eds.(Topics In Geobiology, vol. 19, KluwerAcademic/Plenum, New York, 2001), pp. 235-289.[Nucleurus study]

Five coding revisions and one species added to matrixof study 19a. Duplicate coding with study 22 excluded.

20B. S. Lieberman, G. J. Kloc, Bull. Am. Mus. Nat. Hist.232 (1997).

Neometacanthus sp. aff. stellifer excluded (singlespecimen).

21B. D. E. Chatterton, G. D. Edgecombe, N. E. Vaccari,B. G. Waisfeld, J. Paleontology 71, 108 (1997).

Ceratocephala sp. nov. excluded (known from "fewcoarsely silicified fragments").

22J. M. Adrain, G. D. Edgecombe, Pal. Can. 14, 1(1997).

Characters 8 and 35 excluded (variation within statesexceeds difference between states). Struzia obtusa Aand B (stratigraphically and geographically separatemorphotypes) treated as distinct species.

23J. M. Adrain, L. Ramsköld, J. Paleontology 71, 237(1997).

Outgroup excluded. Duplicate coding with study 5aexcluded.

24G. D. Edgecombe, B. D. E. Chatterton, N. E. Vaccari,B. G. Waisfeld, J. Paleontology 71, 419 (1997).

Decoroproetus excluded ("Frankenstein" OTU).Polymorphisms for character 8 given in text (coded asmost common state in matrix).

25F. A. Sundberg, L. B. McCollum, J. Paleontology 71,1065 (1997).

26a B. S. Lieberman, J. Paleontology 72, 59 (1998).

26bWebster, unpublished manuscript (available fromauthor upon request).

Coding revisions and four species added to matrix ofstudy 26a. Duplicate coding with study 35 excluded.

27B. D. E. Chatterton, G. D. Edgecombe, B. G.Waisfeld, N. E. Vaccari, J. Paleontology 72, 273(1998).

State 3 for character 9 by definition involves apolymorphism.

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28 J. M. Adrain, J. Paleontology 72, 698 (1998).Character 17 excluded (state 1 may hidepolymorphisms overlapping with state 0).

29M. C. Ebach, G. D. Edgecombe, J. Paleontology 73,431 (1999).

Mystrocephala excluded ("Frankenstein" OTU).

30 F. A. Sundberg, J. Paleontology 73, 1126 (1999). Duplicate coding with study 45 excluded.

31S. R. Westrop, R. Ludvigsen, J. Paleontology 74,1020 (2000).

32W.-W. Yuan, L.-Z. Li, Z.-Y. Zhou, C.-S. Zhang, ActaPal. Sinica 40, 388 (2001).

Aulacopleura konincki recoded as polymorphic (states2 and 3) for character 11 (see ref. S3).

33 M. C. Ebach, S. T. Ahyong, Cladistics 17, 1 (2001).

34 B. S. Lieberman, J. Paleontology 75, 96 (2001).Olenelloidea ("Frankenstein" OTU) and duplicatecoding with studies 26a, 26b, and 42 excluded.

35M. Webster, H. D. Sheets, N. C. Hughes, in BeyondHeterochrony: The Evolution of Development, M. L.Zelditch, Ed. (Wiley, New York, 2001), pp. 105-144.

Duplicate coding with studies 26a and 26b excluded.

36B. G. Waisfeld, N. E. Vaccari, B. D. E. Chatterton, G.D. Edgecombe, J. Paleontology 75, 827 (2001).

Shumardia crossi, S. granulosa, S. lacrima, S.tarimuensis, S. whittingtoni, Conophrys tauwhena, C.cf. pellizari, Kweichowilla forbesi, K. matchensis, andK.? minaretta excluded (all coded for incorrect numberof characters). State 3 for character 38 by definitioninvolves a polymorphism.

37J. M. Adrain, S. R. Westrop, E. Landing, R. A.Fortey, J. Paleontology 75, 947 (2001).

Dimeropygiella cf. blanda and Pseudohystricurus sp.excluded (single specimens).

38

J. M. Adrain, S. R. Westrop, in Fossils, Phylogeny,and Form: An Analytical Approach, J. M. Adrain, G.D. Edgecombe, B. S. Lieberman, Eds. (Topics InGeobiology, Vol. 19, Kluwer Academic/Plenum, NewYork, 2001), pp. 291-322.

39 T. J. Cotton, Palaeontology 44, 167 (2001).Duplicate coding with studies 26a, 26b, and 48excluded.

40

S. T. Turvey, in Palaeobiogeography andBiodiversity Change: The Ordovician and Mesozoic-Cenozoic Radiations, J. A. Crame, A. W. Owen, Eds.(Geological Society of London, Special Publications194, 2002), pp. 53-68.

41J. O. R. Ebbestad, G. E. Budd, Palaeontology 45,1171 (2002).

Burlingia angusta, B. cf. jagoi (single specimens), andduplicate coding with studies 25 and 48 excluded.Russian morphotype of Schmalenseeia amphionuratreated as distinct species.

42 B. S. Lieberman, J. Paleontology 76, 692 (2002).Olenellina ("Frankenstein" OTU) and duplicate codingwith studies 26a, 26b, and 34 excluded.

43 C. Crônier, Acta Pal. Pol. 48, 55 (2003).

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44 D. K. Brezinski, Sp. Pap. Pal. 70, 363 (2003).

45 F. A. Sundberg, J. Paleontology 78, 920 (2004).

Pachyaspis gallagari, Caborcella collaris, C.pseudaulax, Hadrocephalites cecinna, Kochiellaarenosa, K. chares, K. crito, K. gracilis, K. mansfieldi,Kochiellina groomensis, Kochiellina janglensis,Kochina americana, Kochina macrops, Mexicellagrandoculus, and Onchocephalus thia excluded (allsingle specimens).

46 L. Amati, S. R. Westrop, J. Syst. Pal. 2, 207 (2004). Raymondaspis excluded ("Frankenstein" OTU).

47T. J. Cotton, R. A. Fortey, in Crustacea andArthropod Relationships, S. Koenemann, R. A.Jenner, Eds. (Crustacean Issues 16, 2005), pp. 95-136.

Revised character-taxon matrix provided by theauthors.

48J. R. Paterson, G. D. Edgecombe, J. Paleontology 80,496 (2006).

Nephrolenellus geniculatus is recoded as polymorphicfor characters 23 and 29 (see ref. S15). Duplicatecoding with studies 26a, 26b, 34, 35, and 42 excluded.

49W. Yuan, R. A. Fortey, S. T. Turvey, Palaeontology49, 537 (2006).

Character 21 (duplicates character 24) and duplicatecoding with studies 15, 27, and 37 excluded.

-R. A. Fortey, B. D. E. Chatterton, Palaeontology 31,165 (1988).

Excluded (coded from "Frankenstein" OTUs).

-B. D. E. Chatterton, D. J. Siveter, G. D. Edgecombe,A. S. Hunt, J. Paleontology 64, 255 (1990).

Excluded (based on larval characters).

- L. Ramsköld, L. Werdelin, Cladistics 7, 29 (1991). Excluded (polymorphisms coded as uncertain).

-L. Ramsköld, Trans. R. Soc. Edin. Earth Sci. 82, 125(1991).

Excluded (polymorphisms coded as uncertain).

-G. D. Edgecombe, in Extinction and Phylogeny, M. J.Novacek, Q. D. Wheeler, Eds. (Columbia UniversityPress, New York, 1992), pp. 144-177.

Three cladistic analyses, all excluded (coded from"Frankenstein" OTUs).

-R. A. Robison, Grøn. Geol. Unders. Bull. 169, 25(1994).

Excluded (polymorphisms coded as most commonstate).

-L. E. Babcock, Grøn. Geol. Unders. Bull. 169, 79(1994).

Two cladistic analyses, both excluded (coded from"Frankenstein" OTUs).

-M. C. Ebach, Rec. W. Austr. Mus. 20, 353 (2002).[Phacopid study]

Excluded (polymorphisms coded as uncertain).

-C. J. Bentley, J. B. Jago, Mem. Ass. Austr. Pal. 30,179 (2004).

Excluded (coded from "Frankenstein" OTUs).

-J. R. Pollitt, R. A. Fortey, M. A. Wills, J. Syst. Pal. 3,225 (2005).

Excluded (coded from "Frankenstein" OTUs).

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Table S2: Frequency and extent of trilobite intraspecific variation, as documented by thenumber of cladistic characters coded with polymorphic states within each of 982 species.Data are drawn from character-taxon matrices listed in Table S1 (studies numbered 1through 49). Subgeneric assignments are not listed. An asterisk following a species nameindicates that character coding for subspecies-level OTUs were merged for that taxon.Ordinal and geological age assignments are listed for each species; burlingiids (Burlingiaand Schmalenseeia) are of uncertain ordinal assignment. Geological age abbreviations:Camb, Cambrian; Ord, Ordovician; Sil, Silurian; Dev, Devonian; Carb, Carboniferous; E,early; M, Middle; L, Late. Column 5 lists the number of characters for which eachspecies was coded; column 6 lists the number of characters for which each species wascoded as polymorphic.

Study Code Species Order AgeChars.Coded

Polym.Chars.

1 Eoasaphus superstes Asaphida L Camb 23 0

1 Proceratopyge conifrons Asaphida L Camb 23 0

1 Sayramaspis yardanshanensis Asaphida L Camb 23 0

1 Pseudoyuepingia whitei Asaphida L Camb 21 0

1 Iwayaspis caelata Asaphida L Camb 22 0

1 Yuepingia niobiformis Asaphida L Camb 22 0

1 Aplotaspis erugata Asaphida L Camb 23 0

1 Haniwoides longus Asaphida L Camb 22 0

1 Tamdaspis tamdensis Asaphida L Camb 20 0

1 Haniwoides varius Asaphida L Camb 23 0

1 Cermatops discoidalis Asaphida L Camb 23 0

1 Cermatops vietus Asaphida L Camb 23 0

1 Guozia crassa Asaphida L Camb 21 0

2 Odontopleura ovata Lichida Sil 21 0

2 Odontopleura brevigena Lichida E Sil 21 0

2 Odontopleura arctica Lichida E Sil 21 0

2 Diacanthaspis serotina Lichida E Sil 19 0

2 Odontopleura greenwoodi Lichida E Sil 21 0

2 Odontopleura maccallai Lichida E Sil 20 0

2 Taemasaspis llandoveryana Lichida E Sil 19 0

2 Odontopleura bombini Lichida E Sil 21 0

2 Odontopleura nehedensis Lichida E Sil 20 0

3 Encrinuroides zhenxiongensis Phacopida L Ord 19 0

3 Curriella tuberculifrons Phacopida Sil 17 0

3 Curriella newlandensis Phacopida E Sil 23 0

3 Curriella clancyi Phacopida E Sil 26 0

3 Encrinurus priscus Phacopida E Sil 27 0

3 "Nucleurus" rotundus Phacopida E Sil 27 0

4 Archaeopleura kazakhensis Lichida L Camb or E Ord 7 0

4 Elbaspis pintadensis Lichida Ord 5 0

4, 21 Ceratocara rarispina Lichida M Ord 45 0

17

4, 21 Ceratocephalina tridens Lichida M Ord 48 0

4 Proceratocephala terribilis Lichida L Ord 21 0

4 Laethoprusia salax Lichida E Sil 19 0

4 Orphanaspis orphana Lichida E Sil 7 0

4 Ceratonurus krejcii Lichida E Dev 21 0

4 Isoprusia mydlakia Lichida M Dev 20 0

5 Acidaspis cincinnatiensis Lichida L Ord 34 0

5 Diacanthaspis cooperi Lichida M Ord 31 0

5 Diacanthaspis scitula Lichida M Ord 29 0

5 Exallaspis bufo Lichida E Sil 34 0

5 Exallaspis acarescola Lichida E Sil 30 0

5 Exallaspis coronata Lichida L Sil 20 0

5 Exallaspis deflexa Lichida E Sil 29 0

5 Exallaspis cf. illinoiensis Lichida E Sil 17 0

5 Exallaspis lothiana Lichida E Sil 28 0

5 Exallaspis muldensis Lichida E Sil 30 0

5 Exallaspis mutica Lichida Sil 32 0

5 Exallaspis quadrimucronata* Lichida E Sil 35 1

5 Exallaspis varbolensis Lichida E Sil 22 0

5 Ketternaspis bispinosa Lichida E Dev 25 0

5 Ketternaspis centrina Lichida E Sil 25 0

5 Ketternaspis chacaltayana Lichida E Dev 18 0

5 Ketternaspis clavata Lichida E Dev 31 0

5 Ketternaspis crenata* Lichida E Sil 35 5

5 Ketternaspis dormitzeri Lichida E Sil 28 0

5 Ketternaspis 'elliptica' Lichida M Dev 32 0

5 Ketternaspis franconica Lichida E Dev 19 0

5 Ketternaspis geinitziana Lichida L Sil 16 0

5 Ketternaspis hirta Lichida Dev 25 0

5 Ketternaspis jaanussoni Lichida E Sil 31 1

5 Ketternaspis lenzi Lichida E Sil 33 1

5 Ketternaspis leridae Lichida E Sil 20 0

5 Ketternaspis leucothea Lichida M Dev 27 0

5 Ketternaspis myops Lichida M Dev 24 0

5 Ketternaspis orphana Lichida E Sil 20 0

5 Ketternaspis parkini Lichida Sil 28 0

5 Ketternaspis pigra* Lichida M Dev 29 2

5 Ketternaspis rattei Lichida Sil 28 0

5 Ketternaspis risbeyi Lichida E Sil 19 0

5 Ketternaspis tuberculata* Lichida E Dev 34 1

5 Ketternaspis wellingtonensis Lichida E Dev 29 0

5 Ketternaspis williamsi A Lichida E Dev 33 2

5 Ketternaspis williamsi B Lichida E Dev 31 0

5 Ketternaspis cf. williamsi Lichida L Sil 31 0

5 Ketternaspis sp. A Lichida E Sil 31 0

5 Ketternaspis n. sp. NSW Lichida E Sil 31 0

5 Leonaspis argentea Lichida E Dev 18 0

18

5 Leonaspis belisarius Lichida E Dev 26 0

5 Leonaspis britannica Lichida E Dev 31 0

5 Leonaspis confluens Lichida E Dev 18 0

5 Leonaspis ezellina Lichida L Sil 18 0

5 Leonaspis glabrata Lichida E Dev 17 0

5 Leonaspis hastata Lichida E Dev 19 0

5 Leonaspis hoernesi Lichida E Dev 28 0

5 Leonaspis jenkinsi Lichida E Dev 28 0

5 Leonaspis lochkovensis Lichida E Dev 31 1

5 Leonaspis leonhardi Lichida L Sil 33 0

5 Leonaspis maura Lichida E Dev 22 0

5 Leonaspis truncata Lichida Dev 20 0

5 Meadowtownella ascita Lichida M Ord 33 0

5 Leonaspis sp. Lichida E Dev 29 2

6, 8 Metacryphaeus giganteus Phacopida E Dev 73 0

6, 8 Metacryphaeus tuberculatus Phacopida M Dev 73 0

6, 8 Plesiomalvinella boulei Phacopida M Dev 73 0

6, 8 Plesiomalvinella pujravii Phacopida M Dev 68 0

6 Malvinella haugi Phacopida E Dev 46 0

6 Malvinella buddeae Phacopida M Dev 51 0

6 Palpebrops goeldi Phacopida M Dev 29 0

6 Palpebrops donegalensis Phacopida E Dev 45 0

6 Vogesina gemellus Phacopida M Dev 18 0

6 Vogesina aspera Phacopida Dev 51 0

6 Vogesina lacunafera Phacopida Dev 51 0

7 Distyrax cooperi Phacopida E Sil 14 0

7 Distyrax n. sp. A Phacopida E Sil 11 0

7 Distyrax n. sp. B Phacopida E Sil 12 0

7 Distyrax n. sp. C Phacopida E Sil 10 0

7 Distyrax peeli Phacopida E Sil 14 0

7 Distyrax pilisterensis Phacopida E Sil 3 0

7 Distyrax n. sp. (Illinois) Phacopida E Sil 9 0

8 Kozlowski superna Phacopida E Dev 37 0

8 Typhlonisus baini Phacopida E Dev 35 0

8 Plesioconvexa praecursor Phacopida E Dev 33 0

8 Eldredgeia venustus Phacopida M Dev 38 0

8 "Clarkeaspis" gouldi Phacopida Dev 31 0

8 Clarkeaspis padillaensis Phacopida Dev 30 0

8 Malvinocooperella pregiganteus Phacopida E Dev 38 0

8 Wolfartaspis cornutus Phacopida M Dev 38 0

8 Metacryphaeus rotundatus Phacopida Dev 38 0

8 Metacryphaeus convexus Phacopida Dev 37 0

8 Metacryphaeus curvigena Phacopida Dev 38 0

8 Metacryphaeus branisai Phacopida Dev 32 0

9 Wallacia hagshawensis Phacopida E Sil 16 0

9 Wallacia pagei Phacopida E Sil 15 0

19

9 Wallacia jaanussoni Phacopida E Sil 17 0

9 Wallacia triangulata Phacopida E Sil 17 0

9 Wallacia expansa Phacopida E Sil 17 0

9 Wallacia masterleei Phacopida E Sil 16 0

9 Wallacia laevis Phacopida E Sil 17 0

9 Wallacia hypolepra Phacopida E Sil 16 0

9 Wallacia elegantula Phacopida E Sil 17 0

9 Wallacia chattertoni Phacopida E Sil 15 0

9 Wallacia hendersoni Phacopida E Sil 16 0

10 Falcoproetus falcatus Proetida E Sil 51 0

10 Proetus latifrons Proetida E Sil 46 0

10 Proetus concinnus Proetida E Sil 51 0

10 Gerastos cuvieri Proetida M Dev 48 0

10 Coniproetus folliceps Proetida M Dev 50 0

10 Coniproetus ryckholti Proetida L Sil 53 0

10 Coniproetus bohemicus Proetida E Dev 53 0

10 Pudoproetus missouriensis Proetida E Carb 53 0

10 Longiproetus tenuimargo Proetida M Dev 51 0

10 "Plesiowensus" confossus Proetida E Sil 50 0

10 Plesiowensus obconicus Proetida L Sil 51 0

10 Arcticormistonia edgecombei Proetida E Dev 52 0

10, 11 Crassiproetus globosus Proetida E Dev 80 0

10, 12 Basidechenella rowi Proetida M Dev 75 0

10, 12 Ormistoniella malaca Proetida E Dev 72 0

10 Aayemenaytcheia paragranulata Proetida E Dev 51 0

10, 13 Lacunoporaspis norrisi Proetida E Dev 92 0

10 Dechenella alpenensis Proetida M Dev 53 0

10, 13 Dechenella perscheii Proetida M Dev 92 0

10 Schizoproetoides richteri Proetida M Dev 50 0

10 Schizoproetus onyx Proetida E Dev 51 0

11 Crassiproetus turgidus Proetida L Sil 11 0

11 Crassiproetus alpenensis Proetida M Dev 32 0

11 Crassiproetus crassimarginatus Proetida M Dev 33 0

11 Crassiproetus traversensis Proetida M Dev 33 0

11 Crassiproetus canadensis Proetida M Dev 33 0

11 Crassiproetus microgranulatus Proetida M Dev 33 0

11 Crassiproetus sibleyensis Proetida M Dev 33 0

11 Crassiproetus calhounensis Proetida M Dev 32 0

11 Crassiproetus stummi Proetida E Dev 11 0

11 Crassiproetus schohariensis Proetida E Dev 10 0

11 Crassiproetus neoturgidus Proetida M Dev 9 0

11 Crassiproetus halliturgidus Proetida M Dev 9 0

12 Basidechenella maura Proetida E Dev 15 0

12 Basidechenella canaliculata Proetida M Dev 20 0

12 Basidechenella clara Proetida E Dev 25 0

12 Basidechenella arkonensis Proetida M Dev 25 0

20

12 Basidechenella nodosa Proetida M Dev 23 0

12 Basidechenella elevata Proetida M Dev 22 0

12 Basidechenella eriensis Proetida M Dev 23 0

12 Basidechenella lucasensis Proetida M Dev 25 0

12 Basidechenella pulchra Proetida M Dev 25 0

12 Basidechenella reimanni Proetida M Dev 24 0

12 Basidechenella hesionea Proetida E Dev 6 0

12 Basidechenella timwhitei Proetida E Carb 17 0

13 Dechenella bathurstensis Proetida M Dev 49 0

13 Dechenella algida Proetida M Dev 49 0

13 Dechenella osborni Proetida M Dev 49 0

13 Dechenella maclareni Proetida M Dev 49 0

13 Dechenella haldemani Proetida M Dev 47 0

13 Dechenella alpenensis Proetida M Dev 49 0

13 Dechenella valentini Proetida M Dev 41 0

13 Dechenella setosa Proetida M Dev 49 0

13 Dechenella rittbergensis Proetida M Dev 49 0

13 Dechenella verneuili Proetida M Dev 45 0

13 Dechenella struvei Proetida M Dev 46 0

13 Dechenella gigouti Proetida M Dev 41 0

13 Dechenella planimarginata Proetida M Dev 43 0

13 Dechenella welleri Proetida M Dev 44 0

13 Dechenella polonica Proetida M Dev 45 0

13 Dechenella carvalhoae Proetida M Dev 43 0

14 Thebanaspis thebana Proetida E Sil 20 0

14 Hedstroemia pachydermata Proetida L Sil 20 0

14 Milesdavis eldredgei Proetida E Sil 10 0

14 Monodechenella boteroi Proetida Dev 10 0

14 Monodechenella halli Proetida M Dev 20 0

14 Monodechenella macrocephala Proetida M Dev 20 0

14 Monodechenella legrandsmithi Proetida M Dev 20 0

14 Monodechenella curvimarginata Proetida M Dev 20 0

14 Monodechenella breviceps Proetida E Carb 19 0

15 Dimeropyge hystrix Proetida M Ord 20 0

15 Dimeropyge labrosa Proetida M Ord 21 0

15 Dimeropyge lucifer Proetida M Ord 19 0

15 Dimeropyge minuta Proetida M Ord 21 0

15 Dimeropyge spinifera Proetida M Ord 21 0

15 Dimeropyge gibbus Proetida M Ord 21 0

15, 27 Dimeropyge speyeri Proetida M Ord 72 0

16 Otarion huddyi Proetida E Sil 14 0

16 Otarion n. sp. A Proetida E Sil 14 0

16 Otarion brauni Proetida E Sil 14 0

16 Otarion coppinensis Proetida E Sil 14 0

16 Otarion beukeboomi Proetida L Sil 14 0

16 Otarion hollandi Proetida Sil 14 0

21

16 Otarion diffractum Proetida L Sil 14 0

17 Encrinuroides autochthon Phacopida M Ord 19 0

17 Encrinuroides capitonis Phacopida L Ord 20 0

17 Physemataspis coopi Phacopida L Ord 20 0

17 Walencrinuroides gelaisi Phacopida L Ord 18 0

17 Encrinurus gibber Phacopida L Ord 15 0

17 Encrinuroides hornei Phacopida M Ord 16 0

17 Encrinuroides insularis Phacopida M Ord 19 0

3, 17 Encrinuroides lapworthi Phacopida L Ord 37 0

3, 17 Encrinuroides neuter Phacopida L Ord 41 0

17 Encrinuroides obesus Phacopida L Ord 19 0

17 Encrinuroides periops Phacopida L Ord 19 0

17 Encrinuroides polypleura Phacopida M Ord 18 0

17 Ceraurus rarus Phacopida L Ord 20 0

17 Cybele sexcostatus Phacopida L Ord 18 0

17 Encrinurus stincharensis Phacopida M Ord 19 0

17 Encrinuroides tholus Phacopida L Ord 16 0

3, 17 Encrinuroides torulatus Phacopida L Ord 41 0

3, 17 Encrinuroides uncatus Phacopida L Ord 41 0

17 Frencrinuroides edelsi Phacopida L Ord 19 0

17 Erratencrinurus spicatus Phacopida L Ord 19 2

18 Agnostus pisiformis Agnostida L Camb 28 0

18 Peronopsis brighamensis Agnostida M Camb 26 0

18 Agnostus punctuosus Agnostida M Camb 28 0

18 Agnostus affinis Agnostida M Camb 28 2

18 Zeteagnostus incautus Agnostida M Camb 27 0

18 Zeteagnostus scarifatus Agnostida M Camb 28 0

18 Acidusus atavus Agnostida M Camb 27 4

18 Acidusus acidusus Agnostida M Camb 28 1

18 Acidusus germanus Agnostida M Camb 28 2

18 Acidusus occultatus Agnostida M Camb 28 0

18 Ptychagnostus michaeli Agnostida M Camb 28 1

18 Ptychagnostus akanthodes Agnostida M Camb 27 1

18 Ptychagnostus cassis Agnostida M Camb 26 1

18 Agnostus aculeatus Agnostida M Camb 25 1

18 Agnostus hybridus Agnostida M Camb 28 4

18 Agnostus seminula Agnostida M Camb 28 1

18 Onymagnostus semiermis Agnostida M Camb 28 1

18 Agnostus barrandei Agnostida M Camb 27 2

18 Agnostus praecurrens Agnostida M Camb 28 3

18, 47 Agnostus gibbus Agnostida M Camb 119 7

18 Triplagnostus scopus Agnostida M Camb 28 1

18 Triplagnostus ultimus Agnostida M Camb 27 0

18 Aotagnostus culminosus Agnostida M Camb 27 1

18 Aotagnostus aotus Agnostida M Camb 27 2

18 Aotagnostus magniceps Agnostida M Camb 27 1

22

18 Myrmecomimus tribulis Agnostida M Camb 28 1

18 Myrmecomimus saltus Agnostida M Camb 26 0

18 Agnostus nathorsti Agnostida M Camb 28 1

18 Goniagnostus scarabaeus Agnostida M Camb 28 3

18 Agnostus spiniger Agnostida M Camb 24 2

18 Goniagnostus oxytaurus Agnostida M Camb 27 1

18 Goniagnostus temniscatus Agnostida M Camb 27 1

18 Agnostus fissus Agnostida M Camb 28 2

18 Agnostus perrugatus Agnostida M Camb 26 0

18 Lejopyge acantha Agnostida M Camb 22 0

18 Agnostus armatus Agnostida M Camb 24 1

18 Lejopyge calva Agnostida M Camb 21 1

18 Battus laevigatus Agnostida M Camb 23 0

18 Lejopyge multifora Agnostida M Camb 22 0

18 Phalacroma? dubium Agnostida M Camb 26 0

18 Agnostus lundgreni Agnostida M Camb 27 1

18 Lejopyge rigbyi Agnostida M Camb 27 0

18 Tomagnostus bothrus Agnostida M Camb 27 2

18 Agnostus elegans Agnostida M Camb 27 0

19 Brianurus jeffersoni Phacopida E Sil 20 0

19 "Encrinurus" nodai Phacopida E Sil 20 0

19 Nucleurus abyssalis Phacopida E Sil 22 0

19 Nucleurus anticostiensis Phacopida E Sil 21 0

19 Nucleurus diabolus Phacopida E Sil 23 0

19 Nucleurus inexpectans Phacopida E Sil 18 0

19 "Encrinurus" mareki Phacopida E Sil 20 0

19 Nucleurus mullochensis Phacopida E Sil 24 0

19 Nucleurus meifodensis Phacopida E Sil 21 0

19 sp. 2 Phacopida E Sil 19 0

19 Elsarella rotunda Phacopida E Sil 24 0

19 Elsarella rumbaensis Phacopida E Sil 20 0

19 "Encrinurus" kiltsiensis Phacopida E Sil 15 0

19 "Encrinurus" selistensis Phacopida E Sil 18 0

19 sp. 3 Phacopida E Sil 15 0

19, 22 Billevittia adraini Phacopida E Sil 55 0

19, 22 Balizoma variolaris Phacopida E Sil 55 0

19, 22 Stuszia dimitrovi Phacopida E Sil 54 0

20 Pelitlina smeenki Phacopida E Dev 62 0

20 Treveropyge prorotundifrons Phacopida E Dev 63 0

20 Pseudocryphaeus michelini Phacopida E Dev 64 0

20 Pseudocryphaeus munieri Phacopida E Dev 64 0

20 Tolkeinia granulispina Phacopida E Dev 51 0

20 Braunops obsoletus Phacopida E Dev 65 0

20 Braunops prostellans Phacopida E Dev 63 0

20 Hallandclarkeops cantamoricus Phacopida M Dev 65 0

20 Neocalmonia quadricosta Phacopida L Dev 64 0

23

20 Bellacartwrightia phyllocaudata Phacopida M Dev 66 0

20 Bellacartwrightia pleione Phacopida M Dev 28 0

20 Bellacartwrightia jennyae Phacopida M Dev 66 0

20 Bellacartwrightia whiteleyi Phacopida M Dev 66 0

20 Bellacartwrightia calderonae Phacopida M Dev 66 0

20 Coltraneia celtica Phacopida E Dev 64 0

20 Bradocryphaeus afghanica Phacopida M Dev 63 0

20 Heliopyge turolensis Phacopida M Dev 63 0

20 Paracryphaeus hercyniae Phacopida E Dev 61 0

20 Pilletina aequisulcata Phacopida E Dev 62 0

20 Breizhops lanceolatus Phacopida E Dev 65 0

20 Breizhops chaconae Phacopida E Dev 64 0

20 Breizhops bigranulosus Phacopida E Dev 64 0

20 Philipsmithiana burtandmimiae Phacopida M Dev 66 0

20 Philipsmithiana hyfinkeli Phacopida M Dev 66 0

20 Modellops struvei Phacopida E Dev 64 0

20 Asteropyge punctata Phacopida Dev 66 0

20 Rhenops babini Phacopida E Dev 62 0

20 Rhenops n. sp. Phacopida L Dev 60 0

20 Greenops chilmanae Phacopida M Dev 65 0

20 Greenops grabaui Phacopida M Dev 66 0

20 Greenops widderensis Phacopida M Dev 66 0

20 Greenops barberi Phacopida M Dev 66 0

20 Deloops hassi Phacopida M Dev 65 0

20 Stummiana osismorum Phacopida M Dev 63 0

20 Stummiana arkonensis Phacopida M Dev 59 0

20 Armorigreenops leoi Phacopida M Dev 61 0

20 Kennacryphaeus harrisae Phacopida M Dev 63 0

20 Kayserops inflatus Phacopida E Dev 65 0

20 Kayserops traversensis Phacopida M Dev 61 0

21 Ceratocara argentina Lichida L Ord 29 0

21 Ceratocara sexpares Lichida L Ord 23 0

21 Ceratocara shawi Lichida M Ord 29 0

21 Ceratocephala triacantheis Lichida M Ord 29 0

21 Ceratocephala laciniata Lichida M Ord 29 0

21 Ceratocephala relativa Lichida L Ord 28 0

21 Ceratocephala graffhami Lichida M Ord 22 0

22 Frammia arctica Phacopida L Sil 38 0

22 Frammia bachae Phacopida E Sil 35 0

22 Mackenziurus ceejayi Phacopida E Sil 36 0

22 Mackenziurus deedeei Phacopida E Sil 37 0

22 Struszia epsteini Phacopida E Sil 34 0

22 Avalanchurus garfunkeli Phacopida E Sil 31 0

22 Fragiscutum glebale Phacopida L Sil 37 0

22 Struszia harrisoni Phacopida E Sil 37 0

22 Frammia hyperborea Phacopida L Sil 38 0

24

22 Struszia indianensis Phacopida L Sil 33 0

22 Aegrotocatellus jaggeri Phacopida L Sil 38 0

22 Mackenziurus joeyi Phacopida E Sil 31 0

22 Mackenziurus johnnyi Phacopida L Sil 34 0

22 Mackenziurus lauriae Phacopida E Sil 25 0

22 Avalanchurus lennoni Phacopida E Sil 37 0

22 Struszia martini Phacopida E Sil 37 0

22 "Struszia" mccartneyi Phacopida E Sil 37 0

22 Aegrotocatellus nankerphelgeorum Phacopida L Sil 29 0

22 Struszia obtusa A Phacopida L Sil 38 0

22 Struszia obtusa B Phacopida E Sil 32 0

22 Struszia onoae Phacopida E Sil 35 0

22 Struszia petebesti Phacopida E Sil 37 0

22 Struszia ramskoeldi Phacopida L Sil 27 0

22 Mackenziurus reimeri Phacopida E Sil 37 0

22 Fragiscutum rhytium Phacopida L Sil 38 0

22 Struszia rosensteinae Phacopida L Sil 37 0

22 Frammia rossica Phacopida L Sil 22 0

22 Avalanchurus simoni Phacopida E Sil 36 0

22 Avalanchurus starri Phacopida L Sil 34 0

5, 23 Edgecombeaspis beni Lichida E Sil 37 2

5, 23 Edgecombeaspis belli Lichida E Sil 41 1

5, 23 Edgecombeaspis besti Lichida E Sil 37 1

5, 23 Edgecombeaspis boltoni Lichida E Sil 40 1

23 Edgecombeaspis jahansi Lichida E Sil 17 0

23 Edgecombeaspis johansonae Lichida E Sil 17 0

5, 23 Edgecombeaspis longstaffei Lichida E Sil 46 1

23 Edgecombeaspis soehni Lichida E Sil 15 0

23 Edgecombeaspis ulrichi Lichida L Ord 17 1

24 Eremiproetus agellus Proetida L Ord 16 0

24 Stenoblepharum astinii Proetida L Ord 18 1

24 Stenoblepharum dactylum Proetida L Ord 18 1

24 Stenoblepharum kullsbergense Proetida L Ord 17 0

24 Stenoblepharum norvegicum Proetida L Ord 18 0

24 Stenoblepharum planum Proetida L Ord 18 2

24 Stenoblepharum strasburgense Proetida L Ord 18 1

24 Stenoblepharum warburgae Proetida L Ord 18 0

24 Stenoblepharum zalesskyi Proetida L Ord 15 0

25 Arthricocephalus chauveaui Corynexochida E Camb 32 0

25 Changaspis placenta Corynexochida E Camb 36 0

25 Opsioryctocephalus ophis Corynexochida M Camb 33 0

25 Oryctocephalites burgessensis Corynexochida M Camb 36 0

25 Oryctocephalites opiki Corynexochida M Camb 36 0

25 Oryctocephalites palmeri Corynexochida E Camb 34 1

25 Oryctocephalites rasettii Corynexochida M Camb 34 1

25 Oryctocephalites resseri Corynexochida M Camb 31 0

25

25 Oryctocephalites reynoldsi Corynexochida M Camb 36 0

25 Oryctocephalites reynoldsiformis Corynexochida M Camb 36 0

25 Oryctocephalites runcinatus Corynexochida M Camb 33 0

25 Oryctocephalites salteri Corynexochida M Camb 34 0

25 Oryctocephalites typicalis Corynexochida M Camb 31 0

25 Oryctocephalites walcotti Corynexochida M Camb 34 0

25 Oryctocephalops frischenfeldi Corynexochida M Camb 34 0

25 Oryctocephalus indicus Corynexochida M Camb 36 1

25 Oryctocephalus lancastrioides Corynexochida M Camb 36 0

25 Oryctocephalus nyensis Corynexochida M Camb 36 1

25 Oryctocephalus orientalis Corynexochida M Camb 35 0

25 Oryctocephalus primus Corynexochida M Camb 31 0

25 Protoryctocephalus wuxunensis Corynexochida E Camb 34 0

26 Olenellus transitans Redlichiida E Camb 77 0

26 Mesonacis vermontanus Redlichiida E Camb 76 1

26 Mesolenellus hyperborea Redlichiida E Camb 75 3

26 Elliptocephala logani Redlichiida E Camb 56 0

26 Kjerulfia lata Redlichiida E Camb 56 0

26 Esmeraldina rowei Redlichiida E Camb 75 0

26 Holmiella preancora Redlichiida E Camb 57 1

26 Fritzolenellus truemani Redlichiida E Camb 77 1

26 Mummaspis occidens Redlichiida E Camb 73 4

26 Laudonia bispinata Redlichiida E Camb 76 8

26, 35 Bristolia bristolensis Redlichiida E Camb 82 9

26 Fremontella halli Redlichiida E Camb 54 5

26 Lochmanolenellus mexicana Redlichiida E Camb 56 1

26, 35 Nephrolenellus multinodus Redlichiida E Camb 83 9

26 Bolbolenellus euryparia Redlichiida E Camb 56 8

26 Olenelloides armatus Redlichiida E Camb 68 7

26 Peachella iddingsi Redlichiida E Camb 73 10

26 Biceratops nevadensis Redlichiida E Camb 74 5

27 Chomatopyge canasi Proetida M Ord 55 1

27 Chomatopyge falcata Proetida M Ord 54 2

27 Chomatopyge marginifera Proetida M Ord 39 0

27 Hystricurus ellipticus Proetida E Ord 41 0

27 Hystricurus flectimembrus Proetida E Ord 53 1

27 Hystricurus globosus Proetida E Ord 53 0

27 Hystricurus hillyardensis Proetida E Ord 52 2

27 Hystricurus millardensis Proetida E Ord 51 2

27 Hystricurus paragenalatus Proetida E Ord 54 2

27 Hystricurus paucituberculata Proetida E Ord 53 0

27 Hystricurus penchiensis Proetida E Ord 54 3

27 Ischyrophyma tuberculata Proetida M Ord 38 1

27 Lasarchopyge benedettoi Proetida L Ord 55 0

27 Lasarchopyge correae Proetida M Ord 54 0

27 Mesotaphraspis circumflexa Proetida L Ord 40 2

26

27 Mesotaphraspis inornata Proetida M Ord 55 4

27 Mesotaphraspis parva Proetida M Ord 55 3

27 Mesotaphraspis? dalfaskensis Proetida L Ord 54 1

27 Paratoernquistia chinchensis Proetida M Ord 22 3

27 Paratoernquistia reedi Proetida M or L Ord 40 1

27 Paratoernquistia sanchezae Proetida L Ord 55 2

27 Paratoernquistia translata Proetida L Ord 36 1

27 Psalakilus typicus Proetida E Ord 52 2

27 Toernquistia anxiensis Proetida M Ord 17 0

27 Toernquistia minuta Proetida M Ord 17 2

27 Toernquistia nicholsoni Proetida L Ord 33 1

28 Hammannopyge unica Phacopida L Ord 33 0

28 Holia anacantha Phacopida L Ord 30 0

28 Youngia boucoti Phacopida E Sil 37 0

28 Parayoungia brennardi Phacopida E Sil 34 0

28 Acanthoparypha chiropyga Phacopida M Ord 38 0

28 Holia cimelia Phacopida L Ord 38 0

28 Hyrokybe lightfooti Phacopida E Sil 38 0

28 Hyrokybe copelandi Phacopida E Sil 38 0

28 Acanthoparypha echinoderma Phacopida M Ord 38 0

28 Parayoungia eleyae Phacopida E Sil 38 0

28 Acanthoparypha evitti Phacopida M Ord 38 0

28 Parayoungia folinsbeei Phacopida E Sil 32 0

28 Acanthoparypha goniopyga Phacopida L Ord 32 0

28 Parayoungia mclaughlini Phacopida E Sil 33 0

28 Hyrokybe hadnagyi Phacopida E Sil 37 0

28 Youngia johnsoni Phacopida E Sil 37 0

28 Hyrokybe julli Phacopida E Sil 38 0

28 Youngia kathyae Phacopida E Sil 38 0

28 Hyrokybe lenzi Phacopida E Sil 38 0

28 Acanthoparypha perforata Phacopida L Ord 38 0

28 Pandaspinapyga salsa Phacopida L Ord 33 0

28 Holia secristi Phacopida L Ord 38 0

28 Youngia steineri Phacopida E Sil 38 0

28 Parayoungia tuberculata Phacopida E Sil 33 0

29 Radnoria syrphetodes Proetida E Sil 10 0

29 Cordania becraftensis Proetida E Dev 10 0

29 Cordania buicki Proetida E Dev 11 0

29 Cordania cyclurus Proetida E Dev 10 0

29 Cordania falcata Proetida E Dev 11 1

29 Cordania gaspeiou Proetida E Dev 8 0

29 Cordania macrobius Proetida E Dev 11 0

29 Cordania wessmani Proetida E Dev 11 0

30 Ptychoparella cordillerae Ptychopariida M Camb 47 1

30, 45 Eoptychoparia piochensis Ptychopariida M Camb 94 8

30 Pianaspis sors Ptychopariida M Camb 49 1

27

30 Papyriaspis lanceola Ptychopariida M Camb 50 0

30 Amecephalus idahoensis Ptychopariida M Camb 50 3

30, 45 Amecephalus piochensis Ptychopariida M Camb 95 8

30, 45 Amecephalus cleora Ptychopariida M Camb 78 8

30 Amecephalus laticaudum Ptychopariida M Camb 50 2

30 Altiocculus americanus Ptychopariida M Camb 43 0

30 Altiocculus harrisi Ptychopariida M Camb 50 1

30 Altiocculus drumensis Ptychopariida M Camb 43 1

30 Altiocculus concavus Ptychopariida M Camb 44 1

30 Proehmaniella basilica Ptychopariida M Camb 50 1

30 Ehmaniella waptaensis Ptychopariida M Camb 50 1

30 Ehmaniella fronsplanata Ptychopariida M Camb 50 6

30 Elrathiella euthyopsis Ptychopariida M Camb 44 0

30 Alokistocare paranotatum Ptychopariida M Camb 43 0

30 Alokistocare swasii Ptychopariida M Camb 31 1

30 Alokistocare subcoronatum Ptychopariida M Camb 47 2

31 Hysteropleura macgerriglei Ptychopariida L Camb 14 0

31 Tavsenia ditrema Ptychopariida M Camb 14 0

31 Hysteropleura lacinia Ptychopariida M Camb 13 0

31 Hysteropleura adraini Ptychopariida L Camb 14 0

31 Hysteropleura edgecombei Ptychopariida L Camb 14 0

31 Hysteropleura ramskoldi Ptychopariida L Camb 14 0

31 Knechtelia festiva Ptychopariida M Camb 13 0

31 Bolaspidella housensis Ptychopariida M Camb 14 0

32 Aulacopleura andersoni Proetida E Sil 7 0

32 Aulacopleura hercynica Proetida L Sil 4 0

32 Aulacopleura konincki Proetida E Sil 11 1

32 Aulacopleura krizi Proetida E Sil 9 0

32 Aulacopleura nitida Proetida E Sil 2 0

32 Aulacopleura reedi Proetida L Ord 5 0

32 Aulacopleura? ranfordi Proetida E Sil 8 0

32 Aulacopleura soror Proetida E Sil 5 0

32 Aulacopleura wulongensis Proetida E Sil 11 0

32 Aulacopleura beyrichi Proetida M Dev 11 0

32 Aulacopleura bohemica Proetida E Dev 11 0

32 Aulacopleura dawanensis Proetida E Ord 4 0

32 Aulacopleura ignorata Proetida M Dev 11 0

32 Aulacopleura longecornuta Proetida M Dev 10 0

32 Aulacopleura roquemaillerensis Proetida E Sil 11 0

32 Aulacopleura szechuanica Proetida E Ord 11 0

32 Songkania hanjiadianensis Proetida E Sil 11 0

32 Songkania pijiazhaiensis Proetida E Sil 5 0

32 Songkania smithi Proetida E Sil 8 0

32 Songkania socialis Proetida E Sil 11 1

33 Hemiarges wesenbergensis Lichida L Ord 11 0

33 Acanthopyge australiformis Lichida E Dev 15 0

28

33 Acanthopyge australis Lichida E Dev 12 0

33 Acanthopyge balliviani Lichida Dev 15 0

33 Acanthopyge brankiensis Lichida E Dev 15 0

33 Acanthopyge brevis Lichida E Dev 8 0

33 Acanthopyge campbelli Lichida E Dev 15 0

33 Acanthopyge consanguinea Lichida E Dev 15 0

33 Acanthopyge decheni Lichida E Dev 8 0

33 Acanthopyge docekali Lichida E Dev 15 0

33 Acanthopyge edgecombei Lichida E Dev 15 0

33 Acanthopyge erinacea Lichida E Dev 14 0

33 Acanthopyge hirsuta Lichida E Sil 15 0

33 Acanthopyge limbata Lichida Dev 13 0

33 Acanthopyge orientalis Lichida L Sil 13 0

33 Acanthopyge parapleura Lichida E Sil 12 0

33 Acanthopyge parva Lichida E Dev 13 0

33 Acanthopyge pragensis Lichida L Sil 13 0

33 Acanthopyge pustulosa Lichida E Dev 13 0

33 Acanthopyge richteri Lichida L Sil 15 0

33 Acanthopyge rohri Lichida E Sil 13 0

33 Acanthopyge sinuata Lichida E Dev 15 0

33 Acanthopyge trinodis Lichida E Dev 12 0

33 Acanthopyge uralensis Lichida Dev 11 0

33 Acanthopyge sp. Victoria Lichida E Dev 14 0

34 Pseudojudonia egregia Redlichiida E Camb 34 3

34 Plesionevadia burri Redlichiida E Camb 39 1

34 Nevadella cartlandi Redlichiida E Camb 38 0

34 Nevadella eucharis Redlichiida E Camb 54 0

34 Nevadella mountjoyi Redlichiida E Camb 56 3

34 Nevadella perfecta Redlichiida E Camb 56 5

34 Cirquella espinata Redlichiida E Camb 41 1

34 Cirquella nummularia Redlichiida E Camb 41 0

34 Cirquella nelsoni Redlichiida E Camb 32 1

34 Bradyfallotaspis fusa Redlichiida E Camb 39 0

34 Paranevadella subgroenlandicus Redlichiida E Camb 35 0

34 Cambroinyoella wallacei Redlichiida E Camb 51 0

34 Geraldinella corneiliana Redlichiida E Camb 41 0

26, 34 Cambropallas telesto Redlichiida E Camb 89 2

26, 34 Andalusiana cornuta Redlichiida E Camb 79 5

26, 34 Gabriellus sp. Redlichiida E Camb 77 2

26, 34 Callavia broeggeri Redlichiida E Camb 80 1

34 Callavalonia callavei Redlichiida E Camb 47 0

34 Sdzuyomia lotzei Redlichiida E Camb 45 0

34 Bondonella typica Redlichiida E Camb 56 2

26, 34 Neltneria jacqueti Redlichiida E Camb 91 0

35 Bristolia harringtoni Redlichiida E Camb 12 1

35 Nephrolenellus? n. sp. Redlichiida E Camb 12 1

29

36 Akoldinioidia dydimacantha Ptychopariida L Camb 38 2

36 Shumardia aculeata Ptychopariida L Ord 47 2

36 Shumardia analoga Ptychopariida M Ord 40 1

36 Shumardia angulata Ptychopariida E Ord 45 0

36 Shumardia gadwanensis Ptychopariida E Ord 47 1

36 Conophrys acutifrons Ptychopariida E Ord 46 0

36 Conophrys alata Ptychopariida E Ord 45 1

36 Conophrys changshanensis Ptychopariida E Ord 50 2

36 Conophrys fabiani Ptychopariida E Ord 51 1

36 Conophrys gaoluoensis Ptychopariida E Ord 50 2

36 Conophrys gonioloba Ptychopariida E Ord 41 0

36 Conophrys minutula Ptychopariida E Ord 46 0

36 Conophrys pentagonalis Ptychopariida Ord 34 0

36 Conophrys pusilla Ptychopariida E Ord 47 1

36 Conophrys rushtoni Ptychopariida E Ord 50 2

36 Conophrys salopiensis Ptychopariida E Ord 51 1

36 Conophrys sulcatus Ptychopariida E Ord 50 0

36 Conophrys wrighti Ptychopariida E Ord 46 2

36 Leoforteyia exopthalma Ptychopariida M Ord 46 2

36 Leoforteyia hintzei Ptychopariida E Ord 52 0

36 Leoforteyia ludvigseni Ptychopariida E Ord 48 0

36 Changchowilla? carrerai Ptychopariida M Ord 50 0

36 Changchowilla gracielae Ptychopariida M Ord 46 0

36 Changchowilla riojana Ptychopariida M Ord 51 1

36 Changchowilla sanjuanina Ptychopariida M Ord 49 2

36 Kweichowilla piojensis Ptychopariida L Ord 48 2

36 Kweichowilla salasae Ptychopariida M Ord 52 0

36 Kweichowilla sterrenae Ptychopariida L Ord 47 1

36 Kweichowilla? acuticaudata Ptychopariida E Ord 46 0

36 Kweichowilla? lacrimosa Ptychopariida M Ord 42 0

36 Acanthopleurella grinrodi Ptychopariida E Ord 47 1

36 Koldinioidia typicalis Ptychopariida L Camb 47 2

36 Tarimella bituberculata Ptychopariida M Ord 50 0

37 Ischyrophyma? borealis Proetida M Ord 16 0

15, 27, 37 Dimeropygiella caudanodosa Proetida E Ord 94 1

37 Ischyrotoma juabensis Proetida M Ord 24 0

37 Dimeropygiella mccormicki Proetida E Ord 25 0

37 Pseudohystricurus orbus Proetida E Ord 17 0

15, 27, 37 Dimeropygiella ovata Proetida E Ord 96 0

37 Ischyrotoma parallela Proetida E Ord 15 0

27, 37 Ischyrotoma twenhofeli Proetida M Ord 77 1

37 Ischyrotoma sp. Proetida M Ord 22 0

37 Parahystricurus smithiae Proetida E Ord 25 0

38 Euptychaspis kirki Asaphida L Camb 23 0

38 Euptychaspis typicalis Asaphida L Camb 25 0

38 Euptychaspis jugalis Asaphida L Camb 19 0

30

38 Kathleenella subula Asaphida L Camb 25 0

38 Kathleenella maritima Asaphida L Camb 20 0

38 Kathleenella hamula Asaphida L Camb 22 0

38 Larifugula triangula Asaphida L Camb 25 0

38 Leiobienvillia leonensis Asaphida E Ord 25 0

38 Sunwaptia carinata Asaphida L Camb 22 0

38 Wilcoxaspis bulbosa Asaphida L Camb 22 0

38 Macronoda prima Asaphida L Camb 22 0

38 Macronoda punctata Asaphida L Camb 23 1

38 Keithiella depressa Asaphida L Camb 25 0

39 Atopina antiqua Ptychopariida E Camb 48 2

39 Atops rupertensis Ptychopariida E Camb 84 1

39 Atops trilineatus Ptychopariida E Camb 74 2

39 Bailiaspis bobrovi Ptychopariida M Camb 60 3

39 Bailiaspis dalmani Ptychopariida M Camb 65 1

39 Bailiaspis glabrata Ptychopariida M Camb 62 1

39 Bailiaspis venusta Ptychopariida M Camb 65 6

39 Bailiella aequalis Ptychopariida M Camb 63 1

39 Bailiella baileyi Ptychopariida M Camb 75 5

39 Bailiella emarginata Ptychopariida M Camb 66 3

39 Bailiella lantenoisi Ptychopariida M Camb 82 1

39 Bailiella levyi Ptychopariida M Camb 72 4

39 Conocoryphe caecigena Ptychopariida M Camb 58 2

39 Conocoryphe sulzeri Ptychopariida M Camb 89 1

39 Cornucoryphe schirmi Ptychopariida M Camb 65 1

39 Couloumania heberti Ptychopariida M Camb 80 2

39 Ctenocephalus bergeroni Ptychopariida M Camb 65 1

39 Ctenocephalus coronatus Ptychopariida M Camb 80 1

39 Ctenocephalus antiquus Ptychopariida M Camb 55 2

39 Ctenocephalus exsulans Ptychopariida M Camb 64 1

39 Ctenocephalus matthewi Ptychopariida M Camb 63 0

39 Ctenocephalus terranovicus Ptychopariida M Camb 53 2

39 Dasometopus breviceps Ptychopariida M Camb 72 1

39 Dasometopus granulatus Ptychopariida M Camb 60 4

39 Dasometopus maensis Ptychopariida M Camb 81 3

39 Elyx laticeps Ptychopariida M Camb 72 1

39 Elyx matthewi Ptychopariida M Camb 64 3

39 Hartshillia clivosa Ptychopariida M Camb 41 0

39 Hartshillia inflata Ptychopariida M Camb 63 0

39 Hartshillina spinata Ptychopariida M Camb 59 1

39 Holocephalina leve Ptychopariida M Camb 59 1

39 Holocephalina primordialis Ptychopariida M Camb 72 3

39 Holocephalites incertus Ptychopariida M Camb 60 2

39 Meneviella venulosa Ptychopariida M Camb 88 1

39 Meneviella viatrix Ptychopariida M Camb 85 3

39 Parabailiella languedocensis Ptychopariida M Camb 72 5

39 Pseudatops reticulatus Ptychopariida E Camb 61 0

31

39 Sdzuyella stremina Ptychopariida M Camb 46 1

39 Tchaiaspis sdzuyi Ptychopariida M Camb 57 3

39 Tchaiaspis sp. nov. Ptychopariida M Camb 54 3

39 Acontheus sp. nov. Ptychopariida M Camb 67 1

39 Agraulos ceticephalus Ptychopariida M Camb 77 1

39 Clavigellus annulus Ptychopariida M Camb 73 2

39 Elrathia kingii Ptychopariida M Camb 87 1

39, 48 Eoredlichia intermedia Redlichiida E Camb 111 0

26, 39 Olenellus thompsoni Redlichiida E Camb 140 1

39 Olenoides serratus Corynexochida M Camb 82 1

39 Ptychoparia striata Ptychopariida M Camb 88 1

40 Bavarilla hofensis Phacopida E Ord 25 0

40 Calymenella preboisseli Phacopida E Ord 25 1

40 Calymenesun granulosa Phacopida M Ord 25 0

40 Neseuretinus turcicus Phacopida L Ord 25 0

40 Neseuretus arenosus Phacopida E Ord 23 0

40 Neseuretus attenuatus Phacopida M Ord 16 0

40 Neseuretus avus Phacopida M Ord 25 2

40 Neseuretus chaschuilensis Phacopida M Ord 19 0

40 Neseuretus caerhunensis Phacopida E Ord 23 0

40 Neseuretus concavus Phacopida M Ord 25 1

40 Neseuretus elegans Phacopida M Ord 25 0

40 Neseuretus henkei Phacopida M Ord 25 0

40 Neseuretus intermedius Phacopida M Ord 23 1

40 Neseuretus leonensis Phacopida E Ord 19 0

40 Neseuretus lipanensis Phacopida M Ord 25 0

40 Neseuretus monensis Phacopida E Ord 24 0

40 Neseuretus murchisoni Phacopida E Ord 24 2

40 Neseuretus parvifrons Phacopida E Ord 25 2

40 Neseuretus planus Phacopida M Ord 23 0

40 Neseuretus ramsayensis Phacopida E Ord 25 1

40 Neseuretus sanlucasensis Phacopida M Ord 21 0

40 Neseuretus sexangulus Phacopida E Ord 16 0

40 Neseuretus shensiensis Phacopida E Ord 25 0

40 Neseuretus tristani Phacopida M Ord 25 0

40 Neseuretus turveyi Phacopida M Ord 25 0

40 Neseuretus vaningeni Phacopida M Ord 25 0

40 Pharostomina oepiki Phacopida E Ord 24 0

40 Pradoella pradoi Phacopida E Ord 25 1

40 Protocalymene mcallisteri Phacopida M Ord 25 0

40 Reedocalymene expansa Phacopida M Ord 25 0

40 Sarrabesia teichmuelleri Phacopida M Ord 25 0

40 Vietnamia douvillei Phacopida L Ord 23 0

41 Burlingia hectori Uncertain M Camb 22 0

41 Burlingia jagoi Uncertain M Camb 22 0

41 Burlingia obscura Uncertain M Camb 19 0

32

41 Burlingia ovata Uncertain M Camb 19 0

41 Schmalenseeia acutangulata Uncertain M Camb 22 0

41 Schmalenseeia amphionura Uncertain L Camb 22 0

41 Schmalenseeia amphionura (Russian) Uncertain L Camb 19 0

41 Schmalenseeia anthrotryphe Uncertain M Camb 21 0

41 Schmalenseeia gostinensis Uncertain M Camb 19 0

41 Schmalenseeia longa Uncertain M Camb 13 0

41 Schmalenseeia sinensis Uncertain M Camb 19 0

41 Schmalenseeia spinulosa Uncertain M Camb 19 0

41 Schmalenseeia cf. spinulosa Uncertain L Camb 16 0

41 Schmalenseeia transversa Uncertain M Camb 8 0

41, 48 Dolerolenus zoppii Redlichiida E Camb 55 0

25, 41 Lancastria roddyi Corynexochida E Camb 55 0

41 Olenus truncatus Ptychopariida L Camb 22 0

41 Peltura scarabaeoides Ptychopariida L Camb 22 0

42 Profallotaspis jakutensis Redlichiida E Camb 24 0

42 Pelmanaspis jurii Redlichiida E Camb 24 0

42 Eofallotaspis tioutensis Redlichiida E Camb 23 0

42 Choubertella spinosa Redlichiida E Camb 29 0

42 Fallotaspis typica Redlichiida E Camb 25 0

34, 42 Parafallotaspis grata Redlichiida E Camb 54 0

42 Archaeaspis hupei Redlichiida E Camb 24 0

42 Archaeaspis nelsoni Redlichiida E Camb 24 0

42 Fallotaspidella musatovi Redlichiida E Camb 24 0

42 Lemdadella linaresiae Redlichiida E Camb 29 1

43 Acuticryphops acuticeps Phacopida L Dev 23 0

43 Trimerocephalus caecus Phacopida L Dev 23 2

43 Trimerocephalus lelievrei Phacopida L Dev 23 1

43 Trimerocephalus shotoriensis Phacopida L Dev 23 2

43 Trimerocephalus dianopsoides Phacopida L Dev 23 1

43 Trimerocephalus tardispinosus Phacopida L Dev 23 0

43 Trimerocephalus procurvus Phacopida L Dev 23 0

43 Trimerocephalus interruptus Phacopida L Dev 23 0

43 Trimerocephalus sponsor Phacopida L Dev 23 0

43 Trimerocephalus polonicus Phacopida L Dev 18 0

43 Trimerocephalus mastophthalmus Phacopida L Dev 23 2

43 Trimerocephalus nigritus Phacopida L Dev 23 1

43 Trimerocephalus trifolius Phacopida L Dev 19 0

43 Trimerocephalus steinachensis Phacopida L Dev 18 0

44 Thigriffithides roundyi Proetida E Carb 25 0

44 Paladin chesterensis Proetida E Carb 25 0

44 Paladin wilsoni Proetida E Carb 25 0

44 Paladin genevievensis Proetida E Carb 25 0

44 Paladin rosei Proetida E Carb 25 0

44 Paladin longispina Proetida E Carb 23 0

44 Paladin osagensis Proetida E Carb 22 0

33

44 Paladin morrowensis Proetida E Carb 25 0

44 Paladin girtyianus Proetida E Carb 25 0

44 Paladin helmsensis Proetida E Carb 25 0

44 Paladin rarus Proetida E Carb 25 0

44 Paladin moorei Proetida E Carb 25 0

44 Paladin czarnieckii Proetida E Carb 25 0

44 Paladin eichwaldi Proetida E Carb 25 0

44 Paladin mucronatus Proetida E Carb 25 0

44 Paladin cuspidatus Proetida E Carb 25 0

44 Paladin maillieuxi Proetida E Carb 25 0

44 Paladin lutugini Proetida L Carb 24 0

44 Paladin cervilatus Proetida L Carb 22 0

44 Paladin trigonopyge Proetida E Permian 24 0

45 Elrathina antiqua Ptychopariida M Camb 66 6

45 Pachyaspis deborra Ptychopariida M Camb 47 9

45 Ptychoparella sp. A Ptychopariida E Camb 67 12

45 Syspacephalus gregarius Ptychopariida M Camb 64 9

45 Amecephalites sundbergi Ptychopariida M Camb 58 8

45 Amecephalus arrojosensis Ptychopariida M Camb 69 7

45 Amecephalus agnesensis Ptychopariida M Camb 67 6

45 Amecephalus sp. A Ptychopariida M Camb 47 7

45 Caborcella arrojosensis Ptychopariida M Camb 46 3

45 Caborcella reducta Ptychopariida M Camb 47 2

45 Eokochaspis delamarensis Ptychopariida M Camb 68 9

45 Eokochaspis longspina Ptychopariida M Camb 65 10

45 Eokochaspis metalaspis Ptychopariida E Camb 69 7

45 Eokochaspis nodosa Ptychopariida M Camb 68 7

45 Eokochaspis? cabinensis Ptychopariida M Camb 65 9

45 Hadrocephalites convexus Ptychopariida M Camb 60 8

45 Hadrocephalites lyndonensis Ptychopariida M Camb 64 6

45 Hadrocephalites rhytidodes Ptychopariida M Camb 64 7

45 Inglefieldia porosa Ptychopariida E Camb 45 9

45 Kochaspis liliana Ptychopariida M Camb 64 15

45 Kochiella augusta Ptychopariida M Camb 66 8

45 Kochiella brevaspis Ptychopariida M Camb 64 7

45 Kochiella eiffelensis Ptychopariida M Camb 59 5

45 Kochiella gibbosa Ptychopariida M Camb 47 7

45 Kochiella mackenziensis Ptychopariida M Camb 50 4

45 Kochiella maxeyi Ptychopariida M Camb 59 5

45 Kochiella rasettii Ptychopariida M Camb 68 10

45 Kochiella tuberculata Ptychopariida E Camb 49 6

45 Mexicella antelopea Ptychopariida M Camb 64 7

45 Mexicella granulata Ptychopariida M Camb 47 7

45 Mexicella mexicana Ptychopariida M Camb 66 14

45 Mexicella obscurus Ptychopariida M Camb 62 8

45 Mexicella robusta Ptychopariida M Camb 65 7

45 Mexicella stator Ptychopariida M Camb 66 9

34

45 Nyella granosa Ptychopariida M Camb 63 6

45 Nyella rara Ptychopariida M Camb 46 5

45 Nyella? plana Ptychopariida M Camb 64 8

45 Nyella clinolimbata Ptychopariida M Camb 51 4

45 Nyella columbiana Ptychopariida M Camb 42 4

45 Nyella immoderata Ptychopariida M Camb 59 7

45 Nyella skapta Ptychopariida M Camb 47 2

45 Onchocephalus depressus Ptychopariida M Camb 46 4

45 Onchocephalus fieldensis Ptychopariida M Camb 47 6

45 Onchocephalus maior Ptychopariida M Camb 46 4

45 Parapoulsenia lata Ptychopariida M Camb 47 6

45 Schistometopus? minor Ptychopariida M Camb 47 6

45 Tonopahella goldfieldensis Ptychopariida M Camb 67 5

45 Tonopahella walcotti Ptychopariida M Camb 67 9

46 Illaenus crassicauda Corynexochida E Ord 20 0

46 Illaenus sarsi Corynexochida E Ord 20 0

46 Illaenus aduncus Corynexochida M Ord 20 0

46 Illaenus planifrons Corynexochida E Ord 17 0

46 Illaenus oscitatus Corynexochida M Ord 18 0

46 Illaenus praecurrens Corynexochida Ord 18 0

46 Illaenus fallax Corynexochida L Ord? 19 0

46 Illaenus roemeri Corynexochida M Ord 19 0

46 Illaenus lacertus Corynexochida L Ord 17 0

46 Illaenus viator Corynexochida L Ord 19 0

46 Illaenus utahensis Corynexochida M Ord 19 0

46 Illaenus marginalis Corynexochida M Ord 20 0

46 Illaenus conradi Corynexochida L Ord 20 0

46 Illaenus angusticollis Corynexochida L Ord 19 0

46 Illaenus depressicapitatus Corynexochida L Ord 17 0

46 Illaenus latiaxiatus Corynexochida L Ord 17 0

46 Illaenus fieldi Corynexochida L Ord 15 0

46 Illaenus conifrons Corynexochida M Ord 15 0

46 Nanillaenus? raymondi Corynexochida M Ord 19 0

46 Nanillaenus aduncus Corynexochida L Ord 17 0

46 Nanillaenus mackenziensis Corynexochida L Ord 18 0

46 Thaleops ovata Corynexochida L Ord 20 0

46 Thaleops longispina Corynexochida L Ord 19 0

46 Thaleops anusacermissima Corynexochida L Ord 18 0

46 Thaleops laurentiana Corynexochida M Ord 20 0

46 Thaleops jaanussoni Corynexochida L Ord 20 0

46 Thaleops mobydicki Corynexochida L Ord 20 0

47 Condylopyge amitina Agnostida E Camb 94 2

47 Peronopsis roddyi Agnostida E Camb 92 3

47 Abakolia minutus Agnostida M Camb 98 3

47 Abakolia pauca Agnostida M Camb 90 7

47 Acidiscus birdi Agnostida E Camb 94 2

35

47 Acidiscus theristes Agnostida E Camb 101 4

47 Acimetopus bilobatus Agnostida E Camb 91 1

47 Alaskadiscus spinosus Agnostida E Camb 104 3

47 Analox bipunctata Agnostida E Camb 83 3

47 Bathydiscus dolichometopus Agnostida E Camb 86 3

47 Bolboparia superba Agnostida E Camb 93 1

47 Calodiscus lobatus Agnostida E Camb 94 8

47 Cephalopyge notabilis Agnostida E Camb 84 3

47 Chelediscus acifer Agnostida E Camb 97 2

47 Chelediscus chathamensis Agnostida E Camb 84 3

47 Cobboldites comleyensis Agnostida E Camb 84 2

47 Cobboldites itsariensis Agnostida E Camb 82 6

47 Dawsonia bohemicus Agnostida M Camb 92 2

47 Dawsonia dawsoni Agnostida M Camb 95 2

47 Delgadella caudatus Agnostida E Camb 91 4

47 Delgadella lenaicus Agnostida E Camb 97 1

47 Delgadella amouslekensis Agnostida E Camb 84 2

47 Dicerodiscus tsunyiensis Agnostida E Camb 84 2

47 Egyngolia willochra Agnostida E Camb 100 4

47 Egyngolia obtusa Agnostida E Camb 102 6

47 Egyngolia zaicevi Agnostida E Camb 57 2

47 Ekwipagetia marginata Agnostida E Camb 100 6

47 Ekwipagetia plicofimbria Agnostida E Camb 100 5

47 Eodiscus borealis Agnostida M Camb 97 4

47 Eodiscus scanicus Agnostida M Camb 99 9

47 Hebediscina sardoa Agnostida E Camb 99 5

47 Hebediscina blagonravovi Agnostida E Camb 90 6

47 Hebediscina yuqingensis Agnostida E Camb 99 5

47 Hebediscus attleborensis Agnostida E Camb 84 2

47 Helepagetis bitruncula Agnostida M Camb 101 6

47 Jinghediscus nummularius Agnostida M Camb 89 4

47 Kiskinella cristata Agnostida M Camb 98 1

47 Korobovia ocellata Agnostida E Camb 94 3

47 Lenadiscus unicus Agnostida E Camb 91 4

47 Leptochilodiscus punculatus Agnostida E Camb 85 4

47 Leptochilodiscus succinctus Agnostida E Camb 95 5

47 Litometopus longispinus Agnostida E Camb 86 1

47 Luvsanodiscus gammatus Agnostida E Camb 101 3

47 Macannaia maladensis Agnostida M Camb 110 5

47 Mallagnostus desideratus Agnostida E Camb 52 1

47 Mallagnostus limbatus Agnostida E Camb 84 3

47 Mallagnostus llarenai Agnostida E Camb 91 5

47 Meniscuchus menetus Agnostida E Camb 88 4

47 Meniscuchus nanus Agnostida E Camb 84 1

47 Natalina incita Agnostida E Camb 90 6

47 Natalina dilata Agnostida E Camb 86 3

47 Neocobboldia dentata Agnostida E Camb 97 2

36

47 Neopagetia rjonsnitzkii Agnostida E Camb 101 3

47 Ninadiscus strobulatus Agnostida E Camb 89 5

47 Oodiscus subgranulatus Agnostida E Camb 87 1

47 Opsidiscus bilobatus Agnostida M Camb 96 7

47 Opsidiscus microspinus Agnostida M Camb 108 5

47 Opsidiscus longispinus Agnostida M Camb 70 2

47 Pagetia bootes Agnostida M Camb 110 2

47 Pagetia prolata Agnostida M Camb 111 3

47 Pagetides elegans Agnostida E Camb 109 6

47 Pagetides fragrum Agnostida E Camb 101 5

47 Pseudocobboldia pulchra Agnostida M Camb 93 4

47 Runcinodiscus index Agnostida E Camb 91 2

47 Semadiscus sollennis Agnostida E Camb 58 1

47 Serrodiscus speciosus Agnostida E Camb 98 5

47 Serrodiscus gravestocki Agnostida E Camb 91 4

47 Serrodiscus daedalus Agnostida E Camb 91 4

47 Serrodiscus ctenoa Agnostida E Camb 98 3

47 Sinodiscus shipaiensis Agnostida E Camb 101 2

47 Sinodiscus subquadratus Agnostida E Camb 94 4

47 Sinopagetia jinnanensis Agnostida M Camb 97 1

47 Stigmadiscus stenometopus Agnostida E Camb 84 2

47 Tannudiscus altus Agnostida E Camb 83 6

47 Tannudiscus balanus Agnostida E Camb 94 3

47 Tchernyshevioides ninae Agnostida M Camb 96 3

47 Tsunyidiscus niutitangensis Agnostida E Camb 109 3

47 Tsunyidiscus kaiyangensis Agnostida E Camb 106 6

47 Tsunyidiscus aclis Agnostida E Camb 103 4

47 Tsunyidiscus orientalis Agnostida E Camb 104 3

47 Tsunyidiscus longquanensis Agnostida E Camb 104 3

47 Weymouthia nobilis Agnostida E Camb 72 2

47 Yukonia intermedia Agnostida E Camb 105 6

47 Yukonides lacrinus Agnostida E Camb 98 6

48 Olenellus fowleri Redlichiida E Camb 41 0

35, 48 Bristolia anteros Redlichiida E Camb 36 0

35, 48 Nephrolenellus geniculatus Redlichiida E Camb 45 3

26, 48 Wanneria walcottana Redlichiida E Camb 100 5

26, 48 Elliptocephala asaphoides Redlichiida E Camb 109 5

26, 48 Holmia kjerulfi Redlichiida E Camb 102 1

42, 48 Fallotaspis bondoni Redlichiida E Camb 61 3

26, 34, 42, 48 Daguinaspis ambroggii Redlichiida E Camb 129 1

42, 48 Archaeaspis macropleuron Redlichiida E Camb 48 0

42, 48 Repinaella sibirica Redlichiida E Camb 42 1

26, 34, 48 Judomia tera Redlichiida E Camb 113 0

26, 34, 48 Nevadia weeksi Redlichiida E Camb 115 0

48 Emuella polymera Redlichiida E Camb 42 0

48 Balcoracania dailyi Redlichiida E Camb 42 0

37

48 Redlichia takooensis Redlichiida E Camb 40 0

42, 48 Lemdadella antarcticae Redlichiida E Camb 55 1

48 Yinites wanshougongensis Redlichiida E Camb 38 0

48 Abadiella huoi Redlichiida E Camb 36 0

48 Eccaparadoxides pusillus Redlichiida M Camb 42 1

48 Anabaraspis splendens Redlichiida E Camb 36 2

48 Xystridura templetonensis Redlichiida M Camb 43 3

42, 48 Bigotina bivallata Redlichiida E Camb 57 3

48 Estaingia bilobata Redlichiida E Camb 43 0

48 Ichangia ichangensis Redlichiida E Camb 33 0

48 Ellipsocephalus gripi Redlichiida M Camb 41 1

15, 27, 49 Dimeropyge clintonensis Proetida M Ord 93 1

15, 27, 49 Dimeropyge virginiensis Proetida M Ord 93 0

15, 27, 49 Dimeropyge dorothyae Proetida L Ord 91 0

27, 37, 49 Ischyrotoma anataphra Proetida E Ord 90 2

27, 37, 49 Ischyrotoma stubblefieldi Proetida M Ord 93 1

37, 49 Ischyrotoma wahwahensis Proetida E Ord 40 0

37, 49 Dimeropygiella fillmorensis Proetida E Ord 40 0

15, 27, 37, 49 Dimeropygiella blanda Proetida E Ord 108 0

37, 49 Pseudohystricurus obesus Proetida E Ord 32 0

49 Pseudopetigurus deprati Proetida E Ord 24 0

49 Pseudopetigurus ovatus Proetida L Ord 19 0

49 Glaphurella harknessi Proetida L Ord 17 0

49 Celmus granulatus Proetida M Ord 24 0

49 Celmus michuelmus Proetida M Ord 21 0

15, 27, 49 Genalaticurus genalatus Proetida E Ord 93 1

27, 49 Hystricurus scrofulosus Proetida E Ord 74 0

TOTALS 982 species 40957 1080

SUPPORTING REFERENCESS1. J. J. Wiens, in Phylogenetic Analysis of Morphological Data, J. J. Wiens, Ed.

(Smithsonian Institution Press, Washington, 2000), pp. 115-145.S2. K. J. McNamara, Biol. Rev. 61, 121 (1986).S3. N. C. Hughes, R. E. Chapman, Lethaia 28, 333 (1995).S4. N. C. Hughes, R. E. Chapman, J. M. Adrain, Evol. Dev. 1, 24 (1999).S5. N. C. Hughes, Int. Comp. Biol. 43, 185 (2003).S6. N. C. Hughes, in Evolving Form and Function: Fossils and Development, D. E. G.

Briggs, Ed. (Peabody Museum of Natural History, Yale Univ., New Haven,2005), pp. 139-158.

S7. N. C. Hughes, Geology 19, 913 (1991).S8. L. H. Smith, Paleobiology 24, 17 (1998).S9. L. H. Smith, Lethaia 31, 99 (1998).S10. J. H. Ashton, A. J. Rowell, Paleobiology 1, 161 (1975).S11. N. C. Hughes, Smithsonian Contributions to Paleobiology 79, (1994).

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S12. J. J. Sepkoski Jr., Bull. American Paleontology 363, (2002).S13. M. Foote, in Deep Time: Paleobiology’s Perspective, D. H. Erwin, S. L. Wing, Eds.

(Paleontological Society, 2000), pp. 74-102.S14. M. Foote, Paleobiology 19, 185 (1993).S15. M. Webster, J. Paleontology, in press.