<<

University of Alberta

Select Proetid of southern Morocco

by

Darrin Molinaro

A thesis submitted to the Faculty of Graduate Studies and Research in partial fulfillment of the requirements for the degree of

Master of Science

Department of Earth and Atmospheric Sciences

© Darrin Molinaro Spring 2012 Edmonton, Alberta

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The Tropidocoryphinae and Cornuproetinae represent two subfamilies of proetid trilobites (Class Trilobita, Order ) found throughout the Lower and

Middle Devonian of southern Morocco's Anti-Atlas region. Surveys of the various -bearing horizons within three separate Moroccan basins have yielded a diverse and well preserved collection of genera from both subfamilies. Given the exceptional (complete and articulated) nature of the specimens, systematic description and cladistic analysis, in conjunction with previously described

Moroccan and Central European species of the two subfamilies, was undertaken.

In total, one new (Pontoproetus) and 11 new species (Astycoryphe ditropidia, Tropidocoryphe taharajatensis, T. lahfirensis, T. sculptaridgiata,

Diademaproetus rudimentus, D. issoumourensis, D. corrugatus, D. auxiliairus, D. langus, Pontoproetus truncatus, and P. granulosus) are described and the relationships of Tropidocoryphe-Astycoryphe and Cornuproetus-Diademaproetus-

Pontoproetus are discussed. Acknowledgements

Many special thanks are due to my supervisors Dr. Brian Chatterton and Dr. Lindsey Leighton, for giving me the opportunity to work with them, providing much needed guidance and support, and putting up with my "writing" style. Both of them have provided me with numerous invaluable skills, making me a much better scientist than when I first began this work. Funding for this work was provided by a National Science and Engineering Research Council of Canada Discovery Grant to B. D. E. Chatterton. I would also like to thank the other member of my committee, Dr. Felix Sperling, for his time and comments. Thanks are also due to: Stacey Gibb, for support and guidance beyond measure; Ryan McKellar, for help with PAUP and figure guidelines; Frank Forcino, Emily Stafford, Ben Collins, and Michael Burns for intellectual discussions and support; Allan Lindoe, for the preparation of some specimens, and Kevin Brett, for fieldwork and some original stratigraphic data. I am also in debt to the Department of Earth and Atmospheric Sciences, for their financial support via teaching assistantships and office space. I would also like to thank our Moroccan guide and preparator Hammi Ait H'ssaine, his preparator brother Amur Ait H'ssaine, and their families for their hospitality, support and preparation work on some of the examined specimens; the Berber miners encountered during this work for locating and working the trilobite horizons to produce the bulk of specimens in this study; and the Ministere de l'Energie et des Mines in Rabat, for providing the permits necessary for this work. Lastly, great thanks is also due to my family, in particular my parents for their never ending support and encouragement. Without them I would have never been able to do this work and accomplish what I have. To my grandparents, many thanks for putting up with me not coming home that often and for the many long and always interesting phone conversations. Table of Contents

Chapter 1: Introduction

Introduction 1

Location and geology 2

Examined localities 5

Tindouf Basin 5

Ma'der Basin 6

Tafilalt Basin 8

Purpose and scope 8

Previous research 11

Format 12

Literature cited 18

Chapter 2: Systematic Palaeontology I

Introduction 23

Previous work 25

Materials and methods 27

Stratigraphy 28

Tindouf basin 29

Ma'der basin 30

Terminology 31

Phylogenetic analysis 32

Taxa examined in the analysis 35 Characters and character states 37

Cephalon 42

Glabella 54

Thorax 56

Pygidium 57

Characters not included in the analysis 63

Results 63

Discussion and conclusions 65

Systematic palaeontology 70

Order Proetida Fortey and Owens, 1975 70

Astycoryphe Richter and Richter, 1919 70

Astycoryphe ditropidia new species 72

Tropidocoryphe Novak, 1890 78

Tropidocoryphe amuri Chatterton et al., 2006 79

Tropidocoryphe taharajatensis new species 80

Tropidocoryphe lahfirensis new species 86

Tropidocoryphe sculptaridiata new species 91

Literature Cited 136

Chapter 3: Systematic Palaeontology II

Introduction 142

Previous work 144 Stratigraphy and geology 146

Tindouf basin 146

Ma'der basin 148

Tafilalt basin 151

Terminology 152

Materials and methods 152

Phylogenetic analysis 157

Taxa examined in the analysis 159

Characters and character states 164

Cephalon 165

Thorax 180

Pygidium 181

Characters excluded from the analysis 186

Results 188

Discussion and conclusions 190

Systematic Palaeontology 196

Order Proetida Fortey & Owens, 1975 196

Cornuproetus Richter & Ricter, 1949 196

Diademaproetus Alberti, 1964 199

Diademaproetus praecursor Alberta, 1969 202

Diademaproetus mohamedi Chatterton et al., 2006 204 Diademaproetus rudimentus new species 206

Diademaproetus issoumourensis new species 213

Diademaproetus corrugatus new species 218

Diademaproetus auxiliarus new species 223

Diademaproetus langus new species 229

Pontoproetus new genus 234

Pontoproetus truncatus new species 235

Pontoproetus granulosus new species 240

Literature Cited 304

Chapter 4: Conclusions

Synthesis 309

Future work 312

Literature Cited 314 List of Tables

Chapter 2

Table 2-1. Table outlining included and omitted taxa for the phylogenetic analysis. Images used in coding of specimens are indicated as the source, while material indicates the completeness of specimens being coded. Reasoning provides explanation as to why the taxon was either included or excluded from the analysis. All remaining species not outlined in the table were omitted either based on lack of material (less than 50% of characters could be coded), poor documentation, or not being located in or around the Rheic Ocean of the Devonian. 130

Table 2-2. Results of character state statistical tests on a character by character, state by state basis. Use of Mann-Whittney U, ANOVA, and Hoteling's t-test were dependent on data type. All tests were measured against a 95% confidence interval. When two or more character states are present, a Bonferroni correction adjusted p-value was used to test at the 95% confidence interval. Grey shaded cells represent results which should be used with caution as some of their populations (character states) do not have large enough sample sizes to be statistically accurate. 134

Chapter 3

Table 3-1. Table outlining included and omitted taxa for the phylogenetic analysis. Images used in coding of specimens are indicated as the source, while material indicates the completeness of specimens being coded. Reasoning provides explanation as to why the taxon was either included or excluded from the analysis. All remaining species not outlined in the table were omitted either based on their lack of material (less than 50% of characters could be coded), poor documentation, or not being located in or around the or Devonian Rheic Ocean. 299

Table 3-2. Results of character state statistical tests on a character by character, state by state basis. Use of Mann-Whittney U or Hoteling's t-test were dependent on data type. All tests were measured against a 95% confidence interval. When 2 or more character states are present, a Bonferroni correction adjusted p-value was used to test at the 95% confidence interval. 302 List of Figures

Chapter 1

Figure 1-1. Map of Africa indicating Morocco (in-filled area) (modified from Philip (1991); McKellar & Chatterton (2009); and Gibb & Chatterton (2010)). 13

Figure 1-2. Map of Morocco indicating locations of basins surveyed in this study (images modified from Fetah et al. (1998); Gibb (2005); Chatterton et al. (2006); Gibb & Chatterton (2007,2010); McKellar & Chatterton(2009); Chatterton & Gibb (2010)). 14

Figure 1-3. Map and stratigraphy of select localities of the southern Moroccan Tindouf Basin (image adapted from Fetah et al. (1998); Gibb (2005); Chatterton et al. (2006); Gibb & Chatterton (2007, 2010); McKellar & Chatterton (2009)). 15

Figure 1-4. Map and stratigraphy of select localities of the southern Moroccan Ma'der Basin (image adapted from Fetah et al. (1998); Gibb (2005); Chatterton et al. (2006); Gibb & Chatterton (2007, 2010); McKellar & Chatterton (2009); Chatterton & Gibb (2010)). 16

Figure 1-5. Map and stratigraphy of select localities of the southern Moroccan Tafilalt Basin (image adapted from Fetah et al. (1998); Gibb (2005); Chatterton et al. (2006); McKellar & Chatterton (2009); Gibb & Chatterton (2010)). 17

Chapter 2

Figure 2-1. Locality maps of Africa and Morocco. A: Map of Africa depicting Morocco (shaded area) (Modified from Philip (1991); Gibb (2005); Gibb & Chatterton (2007, 2010); McKellar & Chatterton (2009); Chatterton & Gibb (2010)). B: Map of Morocco indicating locations of basins surveyed in this study (modified from Gibb & Chatterton (2007,2010); McKellar & Chatterton (2009); Chatterton & Gibb (2010)). C and D: Location of individual study basins within Morocco (refer to Figures. 2-3 and 2-4 for more detailed maps of these areas). 97

Figure 2-2. Redrafted version of Feist & Clarkson's (1989) proposed phylogeny of Middle to Late Devonian tropidocoryphid trilobites from the Montagne Noir sequence of France. 98

Figure 2-3. Map of the Tindouf Basin, southern Morocco. The town of Foum Zguid is located to the north, with the Zgulima section located in the Devonian outcrop to the south southeast (indicated by arrow on map). (Diagram adapted from Fetah et al. (1998), Gibb (2005), Chatterton et al. (2006), Gibb & Chatterton (2007, 2010), and McKellar & Chatterton (2009)). 99

Figure 2-4. Map of the Ma'der Basin, southern Morocco. Boxes indicate section localities. (Diagram adapted from Fetah et al. (1998), Gibb (2005), Chatterton et al. (2006), Gibb & Chatterton (2007,2010) McKellar & Chatterton (2009), and Chatterton & Gibb (2010)). 100

Figure 2-5. Stratigraphic columns of Zguilma (adapted from Gibb (2005), Chatterton et al. (2006), Gibb Chatterton (2007,2010) and McKellar & Chatterton (2009)), Assa, and the western slope of Jbel Oufatene (adapted from Gibb (2005), Gibb and Chatterton (2007,2010), and McKellar & Chatterton (2009)). Measurements are in meters and formations are noted to the right of columns. Mined horizons are indicated indicated by arrows and their local names. 101 Figure 2-6. Diagram showing the points of measurement on the cephalon and pygidium, as per Owens (1973). Ratio data were obtained by comparing two measurements, most commonly one measurement in reference to cephalon length (A, sag.). Coordinate data were collected by comparing the position of points (e.g., landmark (3) in relation to the anterior and posterior limits of the cephalon length measurement (A, sag.). 103 Figure 2-7. Cephalic border shape. Arrow indicates area of referenced used to examine this character. A: inflated ('cushion-like') B: flat 104

Figure 2-8. Presence of border rims. A: border rims are not present (absent) B: border rims are present 104

Figure 2-9. Position of cephalic suture angle beta p. Distances Yp and Xp were divided by the length of the cephalon (A, sag. Figure 2-6). Xpand Yp coordinates from all species were then graphed and generic level populations mathematically differentiated using a Hoteling's t-tests with a Bonferroni correction. Three populations were found to be present for the (3 cephalic suture position. 105

Figure 2-10. Position of cephalic suture angle gamma (y). Distances Yy and Xy were divided by the length of the cephalon (A, sag., Figure 2-6). Xy and Yy coordinates from all species were then graphed and generic level populations were mathematically differentiated using a Hoteling's t-test. Two generic level populations were found to be present for the y cephalic suture position. 106

Figure 2-11. Position of cephalic suture angle delta (5). Distances Ys and Xs were divided by the length of the cephalon (A, sag., Figure 2-6). Xs and Y§ coordinates from all species were then graphed and generic level populations mathematically differentiated using a Hoteling's t-test. Two populations were found to be present at generic levels. 107

Figure 2-12. Shape of glabella, as defined by Snadjr (1980,). A: pear-shaped B: sub-cylindrical C: tongue-shaped 108

Figure 2-13. Pygidial shape as defined by Snadjr (1980). A: semi-elliptical B: rounded pentagonal 108

Figure 2-14. Rhachis (axis) shape. A: cone shaped B: cup shaped 109

Figure 2-15. Shape of pygidial pleural ribs. A: bent B: straight 109

Figure 2-16. Cladogram depicting consensus phylogenetic hypothesis based on the 3 most parsimonious trees produced during Heuristic search with no character weighting or ordering. {#} Denotes Bremer support number for clade, all unmarked clades have value of {1>- 110

Figure 2-17. Expanded view of cladogram (Figure 2-16),displaying changes in character states. Cladogram is a re-drafted version of the output from MacClade (Maddison & Maddison 2001) and its data matrix was initially analyzed using PAUP (Swofford 2002). Numbers preceding brackets denote characters, numbers within brackets denote change in character state. All changes shown are unambiguous. (#) = unique, uniform above [#] = homoplasy outside "#" = homoplasy above {#} = homoplasy above and outside 111 Chapter 3

Figure 3-1. Locality maps of Africa and Morocco. A: Map of Africa depicting Morocco (in-filled area) (Modified from Philip (1991) Gibb & Chatterton (2007,2010); McKellar & Chatterton (2009); Chatterton & Gibb (2010)). B: Map of Morocco indicating locations of basins surveyed in this study (modified from Gibb & Chatterton (2007, 2010); McKellar & Chatterton (2009); Chatterton & Gibb (2010)). C, D, E: Location of individual study basins within Morocco (refer to Figures 3-2 to 3-4 for more detailed maps of these areas). 245

Figure 3-2. Map and stratigraphy of select localities of the southern Moroccan Tindouf Basin. Locations of individual taxa indicated on individual stratigraphic columns correspond to appropriate locations on the outcrop map. (Diagrams adapted from Fetah et al. (1998); Gibb (2005); Chatterton et al. (2006); Gibb & Chatterton (2007, 2010); McKellar & Chatterton (2009)). 246

Figure 3-3. Map and stratigraphy of select localities of the southern Moroccan Ma'der Basin. Locations of individual taxa indicated on individual stratigraphic columns correspond to appropriate locations on the outcrop map. (Diagrams adapted from Fetah et al. (1998); Gibb (2005); Chatterton et al. (2006); Gibb & Chatterton (2007,2010); McKellar & Chatterton (2009); Chatterton & Gibb (2010)). 248

Figure 3-4. Map and stratigraphy of select localities of the southern Moroccan Tafilalt Basin. Locations of individual taxa indicated on individual stratigraphic columns with correspond to appropriate locations on the outcrop map. (Diagrams adapted from Fetah et al. (1998); Gibb (2005); Chatterton et al. (2006); Gibb & Chatterton (2007, 2010); McKellar & Chatterton (2009); Chatterton & Gibb (2010)). 251 Figure 3-5. Diagram showing the points of measurement about the cephalon and pygidium (follows Owens 1973). Ratio data were collected by measuring the distance between two points (e.g., length A = length of cephalon) and comparing it to a different measurement. Coordinate data were collected by comparing the position of landmarks (e.g., cephalic suture angle beta (P)) in relation to the length of the glabella (length Ai + A4). 253

Figure 3-6. Position of cephalic suture angle beta (P). Distances Aia- A4b (glabellar length (Figure 3-5, Ai + A4, sag.)), Aia - /?, and A4b - /? were measured and calculated into landmark beta (fi) coordinates Xfi and Yfi using Bookstein shape coordinate analysis. Landmark beta (fi) coordinates from all species were then graphed and generic level populations mathematically differentiated using a Hoteling's t-test. Two populations were found to be present for landmark beta (p) and are referred to as'narrow' and 'wide' morphologies. Overlap between populations exists as a result of variance in landmark position between specimens of the same generic populations. Coding of this character for the phylogenetic analyses was based on the specimens landmark position in relation to the derived equation (which side of the line separating the populations the specimen fell on), not which generic population the specimen was originally part of. 255

Figure 3-7. Position of cephalic suture angle gamma (y). Distances Aia- A4b (glabellar length (Figure 3-5, Ai + A4, sag.)), Aia - y, and A4b - y were measured and calculated into landmark gamma (y) coordinates X y and Yy using Bookstein shape coordinate analysis. Landmark gamma (y) coordinates from all species were then graphed and generic level populations mathematically differentiated using a Hoteling's t-test. Two populations were found to be present for landmark gamma (y) and are referred to as 'narrow' and 'wide' morphologies. Overlap between populations exists as a result of variance in landmark position between specimens of the same generic populations. Coding of this character for the phylogenetic analyses was based on the specimens landmark position in relation to the derived equation (which side of the line separating the populations the specimen fell on), not which generic population the specimen was originally part of.

257

Figure 3-8. Position of cephalic suture angle delta (5). Distances Aia - A4b (glabellar length (Figure 3-5, Ai + A4, sag.), Aia - S, and A4b - $ were measured and calculated into landmark delta (6) coordinates X y and Yy using Bookstein shape coordinate analysis. Landmark delta (5) coordinates from all species were then graphed and generic level populations mathematically differentiated using Hoteling's t-test. Two populations were found to be present for landmark delta (8) and are referred to as 'narrow' and 'wide' morphologies. Overlap between populations exists as a result of variance in landmark position between specimens of the same generic populations. Coding of the character for phylogenetic analyses was based on the derived equation (which side of the line separating the populations) a specimens landmark, not which generic population they were originally part of. 259

Figure 3-9. Cephalic border shape. Arrow indicates area of referenced used to examine this character. A: flat B: inflated ('cushion-like') 261

Figure 3-10. Truncated anterior portion of cephalic border furrow. Arrow indicates area where character was examined. A: truncated (anterior border furrow absent) B: not truncated (anterior border furrow present) 261

Figure 3-11. Shape of glabella as outlined by Snajdr (1980). A: sub-quadrate B: sub-cylindrical C: weakly violin shaped D: strongly violin shaped 262

Figure 3-12. Shape of eye platform. Arrow indicates area of observation for this character. A: none (Absent) B: posterior lobe only C: anterior and posterior lobes present 262

Figure 3-13. Shape of pygidium as outlined by Snajdr (1980). A: segment of circle B: rounded square C: rounded pentagon 263

Figure 3-14. Shape of pygidial pleural ribs as outlined by Snajdr (1980). A: curved B: bent 263

Figure 3-15. Shape of posterior margin of the pygidium. A: rounded (None) B: notched (Anteriorly indented) C: flat (horizontal) 264

Figure 3-16. Cladogram depicting single phylogenetic hypothesis based on the results of exhaustive, branch and bound, and exhaustive analyses. In all analyses no character weighting or ordering was used. {#} Denotes Bremer support number for clade, all unmarked clades have value of {1}. 265

Figure 3-17. Expanded view of cladogram (Figure 3-16), displaying changes in character states. Cladogram is a re-drafted version of the output from MacClade (Maddison & Maddison 2001) and its data matrix was initially analyzed using PAUP (SwofFord 2002). Numbers preceding brackets denote characters, numbers within brackets denote change in character state. All changes shown are unambiguous. 266 List of Plates

Chapter 2

Plate 2-1. Fig. 1-6. Astycoryphe ditropidia n. sp. from the Philonyx/Quadrops horizon, El Otfal Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA13667) (complete specimen). 1, dorsal view of cephalon and thorax; X7.4. 2, dorsolateral view; X6.7. 3, anterodorsal view; X8.2. 4, dorsal view of pygidium; X8.2. 5, lateral view of librigenal spine and macropleural thoracic spine; X10.7. 6, lateral view; X7.6. 112

Plate 2-2. Fig. 1-5. Tropidocoryphe amuri Chatterton et al., 2006 from ZGEE3, Timrhanrhart Formation, Zguilma, Tindouf Basin, southern Morocco (UA13668) (complete specimen). 1, dorsal view of cephalon and thorax; X4.1. 2, lateral view of cephalon; X4.0.3, anterolateral view of cephalon; X4.1. 4, dorsal view of thorax and pygidium; X4.7. 5, anterior view of cephalon; X4.6. Fig. 6. Tropidocoryphe amuri Chatterton et al., 2006 from Assa 2, Khebchia Formation, Assa, Tindouf Basin, southern Morocco (UA13667) (complete specimen). 6, dorsal view; X3.3. 114

Plate 2-3. Fig. 1,3-6. Tropidocoryphe amuri Chatterton et al., 2006 from ZGEE3, Timrhanrhart Formation, Zguilma, Tindouf Basin, southern Morocco (UA13394) (complete specimen). 1, dorsal view of cephalon and thorax; X3.5. 3, anterolateral view of cephalon; X3.8. 4, dorsal view of thorax and pygidium; X3.6. 5, anterior view of cephalon; X3.7. 6. lateral view; X3.0. Fig. 2. Tropidocoryphe amuri Chatterton et al., 2006 from ZGEE1, Timrhanrhart Formation, Zguilma, Tindouf Basin, southern Morocco (UA13670) (partial specimen). 2, dorsal view; X2.8. 116

Plate 2-4. Fig. 1-2,4,6. Tropidocoryphe amuri Chatterton et al., 2006 from ZGEE3, Timrhanrhart Formation, Zguilma, Tindouf Basin, southern Morocco (UA 13669) (complete specimen). 1, dorsal view of cephalon and thorax; X3.1. 2, lateral view; X3.6. 4, dorsal view of pygidium; X3.2. 6, anterolateral view of cephalon; X3.7. Fig. 3, 5, 7. Tropidocoryphe amuri Chatterton et al., 2006 from ZGEE1, Timrhanrhart Formation, Zguilma, Tindouf Basin, southern Morocco (UA13670) (partial specimen). 3, anterolateral view of cephalon; X2.7. 5, anterior view of cephalon; X2.9. 7, dorsal view of pygidium; X4.7. 118

Plate 2-5. Fig. 1-7. Tropidocoryphe taharajatensis n. sp. Tazoulai't Formation, Taharajat, Ma'der Basin, southern Morocco (UA13673) (complete specimen). 1, dorsal view; X2.2. 2, dorsal view of cranidium; X3.6.3, anterolateral view of cephalon; X2.1. 4, dorsal view of cephalon; X2.2. 5, anterior view of cephalon; X2.4. 6, dorsal view of pygidium; X3.1. 7, lateral view; X2.5. 120

Plate 2-6. Fig. 1-2,4-5. Tropidocoryphe taharajatensis n. sp. Tazoulai't Formation, Taharajat, Ma'der Basin, southern Morocco (UA13674) (complete specimen). 1, dorsal view; X3.5. 2, anterior view of cephalon; X3.5. 4, dorsal view of pygidium; X4.3. 5, lateral view; X2.5. Fig. 3 & 6. Tropidocoryphe taharajatensis n. sp. Tazoulai't Formation, Taharajat, Ma'der Basin, southern Morocco (UA13675) (partial specimen). 3, dorsal view; X3.0. 6, anterior view of cephalon; X2.4. 122

Plate 2-7. Fig. 1-3,5,6. Tropidocoryphe lahfirensis n. sp. from the Philonyx/Quadrops horizon, El Otfal Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA13676) (complete specimen). 1, dorsal view; X2.5. 2, anterior view of cephalon; X2.6. 3, anterolateral view of cephalon; X2.3. 5, dorsal view of pygidium; X3.4. 6, lateral view; X2.1. Fig. 4 & 7. Tropidocoryphe lahfirensis n. sp. from the Philonyx/Quadrops horizon, El Otfal Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA13678) (complete specimen). 4, dorsal view of cephalon and thorax; X3.3. 7, dorsal view of pygidium; X3.2. 124

Plate 2-8. Fig. 1,4,5. Tropidocoryphe lahfirensis n. sp. from the Philonyx/Quadrops horizon, El Otfal Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA13679) (complete specimen). 1, dorsal view; X3.6. 4, anterior view of cephalon; X3.5. 5, lateral view; X2.8. Fig. 2,3,6. Tropidocoryphe lahfirensis n. sp. from the Philonyx/Quadrops horizon, El Otfal Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA13677) (complete specimen). 2, dorsal view of cephalon and thorax; X3.2. 3, dorsal view of thorax and pygidium; X2.9. 6, dorsolateral view of cephalon; X3.5. 126

Plate 2-9. Fig. 1-6. Tropidocoryphe sculptaridgiata n. sp. from the Diademaproetus horizon, El Otfal Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA13672) (partial specimen). 1, dorsal view; X4.9. 2, dorsal view of tropidium; X8.3. 3, dorsal view of thorax and pygidium; X5.5. 4, dorsolateral view; X4.1. 5, lateral view; X3.4. 6, posterolateral view; X4.9. 128

Chapter 3

Plate 3-1. Fig. 1,3-6. Diademaproetus praecursor Alberti, 1969 from ZGEE1, Timrhanrhart Formation, Zguilma, Tindouf Basin, southern Morocco (UA13693) (complete specimen). 1, dorsal view of cephalon and thorax; X2.0.3, dorsal view of thorax and pygidium; X2.1. 4, anterodorsal view of cephalon; X2.2. 5, lateral view; XI.6. 6, anterolateral view of cephalon; XI.4. Fig. 2. Diademaproetus praecursor Alberti, 1969 from ZGEE1, Timrhanrhart Formation, Zguilma, Tindouf Basin, southern Morocco (UA13698) (isolated ). 2, ventral view of hypostome; X9.3. 267

Plate 3-2. Fig. 1,3,5-6. Diademaproetus praecursor Alberti, 1969 from Taouz, Tafilalt Basin, southern Morocco (UA13692) (complete specimen). 1, dorsal view of cephalon and thorax; X2.3. 3, dorsal view of thorax and pygidium; X2.4. 5, anterolateral view of cephalon; XI.9. 6, lateral view of cephalon; X2.6. Fig. 2,4,7. Diademaproetus praecursor Alberti, 1969 from the Heliopyge/Kayserops horizon, El Otfal Formation, Jbel Issoumour, Ma'der Basin, southern Morocco (UA13697) (partial specimen). 2, dorsal view of cephalon and thorax; X2.8. 4, dorsal view of thorax and pygidium; X3.1. 7, anterolateral view of cephalon X3.5. 269

Plate 3-3. Fig. 1,3,5,6. Diademaproetus mohamedi Chatterton et al, 2006 from ZGEE1, Timrhanrhart Formation, Zguilma, Tindouf Basin, southern Morocco (UA13383) (partial specimen). 1, dorsal view; X3.7.3, anterior view of cephalon; X3.5. 5, dorsal view thoracic pleurae; X4.7. 6, dorsolateral view; X2.5. Fig. 2 & 4. Diademaproetus praecursor Alberti, 1969 from the Psychopyge horizon, Tazoulai't Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA 13696) (partial specimen). 2, dorsal view; X4.0. 4, dorsolateral view; X2.7. 271

Plate 3-4. Fig. 1-7. Diademaproetus mohamedi Chatterton etal., 2006 from HarpeslThysanopeltis horizon, El Otfal Formation, Ma'der Basin, southern Morocco (UA13699) (complete specimen). 1, dorsal view of cephalon; X2.5. 2, dorsal view of cranidium; X2.6. 3, anterior view of cephalon; X2.8. 4, anterolateral view of cephalon; X1.8. 5, lateral view; X2.3. 6, dorsal view of pygidium; X3.1. 7, dorsal view of thorax; X2.6. 273

Plate 3-5. Fig. 1-7. Diademaproetus rudimentus n. sp.Tazoulai't Formation, Taharajat, Ma'der Basin, southern Morocco. (UA13688) (complete specimen). 1, dorsal view; XI.7. 2, dorsal view of cranidium; X2.5. 3, dorsal view of glabella; X4.5. 4, dorsolateral view of cephalon; X1.7. 5, anterior view of cephalon; X2.2. 6, lateral view; XI.4. 7, dorsal view of pygidium; X3.6. 275

Plate 3-6. Fig. 1-6. Diademaproetus rudimentus n. sp. Tazoulai't Formation, Taharajat, Ma'der Basin, southern Morocco (UA13686) (complete specimen). 1, dorsal view of cephalon and thorax; X2.5. 2, ventral view of hypostome; X5.2.3, dorsolateral view; X2.1. 4, dorsal view of pygidium; X2.6. 5, lateral view; XI.9. 6, anterior view of cephalon; X2.8. 277

Plate 3-7. Fig. 1-6. Diademaproetus corrugatus n. sp. Tazoulai't Formation, Taharajat, Ma'der Basin, southern Morocco (UA13681) (complete specimen). 1, dorsal view of cephalon and thorax; X2.0. 2, dorsal view of librigena; X5.1. 3, dorsal view of thorax and pygidium; X2.7. 4, anterior view of cephalon; X2.3. 5, lateral view of cephalon; X2.2. 6, anterolateral view of cephalon; XI.8. 279

Plate 3-8. Fig. 1-7. Diademaproetus auxiliarus n. sp. from Assa 2, Khebchia Formation, Assa, Tindouf Basin, southern Morocco (UA13689) (complete specimen). 1, dorsal view; X3.1. 2, dorsal view of cranidium; X3.3. 3, dorsal view of glabella; X6.1. 4, dorsolateral view; X2.4. 5, anterior view of cephalon; X3.5. 6, dorsal view of pygidium; X6.0. 7, lateral view; X2.7. 281

Plate 3-9. Fig. 1-3. Diademaproetus auxiliarus n. sp. from Assa 2, Khebchia Formation, Assa, Tindouf Basin, southern Morocco (UA13690) (complete specimen). 1, dorsal view of cephalon and thorax; X3.3. 2, dorsal view of thorax of pygidium; X3.6. 3, dorsolateral view of cephalon; X3.4. Fig. 4-7. Diademaproetus auxiliarus n. sp. from Assa 2, Khebchia Formation, Assa, Tindouf Basin, southern Morocco (UA13691) (partial specimen). 4, lateral view; X6.5. 5, dorsal view of cephalon; X3.5. 6, dorsal view of pygidium; X2.0. 7, ventral view of hypostome; X4.3. 283

Plate 3-10. Fig. 1-7. Diademaproetus langus n. sp. from the Diademaproetus horizon, El Otfal Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA13685) (complete specimen). 1, dorsal view; X2.4. 2, dorsal view of cranidium; X2.7. 3, dorsal view of glabella; X4.8. 4, anterolateral view of cephalon; X2.3. 5, lateral view; X2.3. 6, anterior view of cephalon; X2.5. 7, dorsal view of pygidium; X2.6. 285

Plate 3-11. Fig. 1-5. Diademaproetus langus n. sp. from the Diademaproetus horizon, El Otfal Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA13683) (complete specimen). 1, dorsal view; X2.5. 2, anterolateral view; X2.3. 3, anterior view of cephalon; X2.6. 4, ventral view of hypostome; X4.0. 5, lateral view; XI.8. 287

Plate 3-12. Fig. 1-7. Diademaproetus issoumourensis n. sp. from the Psychopyge horizon, Tazoulait Formation, Jbel Issoumour, Ma'der Basin, southern Morocco (UA13271) (complete specimen). 1, dorsal view; X2.7. 2, dorsal view of cranidium; X3.2.3, dorsal view of anterior lobe of glabella; X6.0. 4, anterolateral view; X2.5; 5, dorsal view of pygidium; X5.4. 6, anterior view of cephalon; X3.1. 7, lateral view; X2.0. 289 Plate 3-13. Fig. 1-5. Diademaproetus issoumourensis n. sp. from the Psychopyge horizon, Tazoulai't Formation, Jbel Issoumour, Ma'der Basin, southern Morocco (UA13682) (complete specimen). 1, dorsal view of cephalon and thorax; X2.4. 2, anterolateral view of cephalon; X2.3. 3, dorsal view of thoracic pleurae; X6.0. 4, dorsal view of thorax and pygidium; X2.9. 5, anterior view of cephalon X2.4. Fig. 6. Diademaproetus langus n. sp. from the Diademaproetus horizon, El Otfal Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA13684) (complete specimen). 6, dorsal view; X2.4. 291

Plate 3-14. Fig. 1-6. Pontoproetus truncatus n. sp. from the Thysanopeltis horizon, Taboumakhlouf Formation, Zireg, Ma'der Basin, southern Morocco (UA13701) (complete specimen). 1, dorsal view; XI.8. 2, anterolateral view; XI.6.3, anterior view of cephalon; X2.0. 4, dorsal view of cephalon; X2.7. 5, lateral view; XI.4. 6, dorsal view of pygidium; X2.7. Fig. 7. Pontoproetus truncatus n. sp. from Timarzite, Ma'der Basin, southern Morocco. (UA13703) (complete specimen). 7, lateral view; X2.3. 293

Plate 3-15. Fig. 1-2. Pontoproetus truncatus n. sp. from Timarzite, Ma'der Basin, southern Morocco (UA13703) (complete specimen). 1, dorsal view; X2.7. 2, dorsal view of glabella; X4.4. Fig. 3-5. Pontoproetus truncatus n. sp. from the Thysanopeltis horizon, Zireg, Ma'der Basin, southern Morocco (UA13702) (complete specimen). 3, dorsal view of glabella; X3.9. 4, dorsal view of cephalon and thorax; X2.1. 5, dorsal view of thorax and pygidium X2.1. 295

Plate 3-16. Fig. 1-6. Pontoproetus grannulosus n. sp. from the Morocconites/Metacanthina equivalent horizon level, Tazoulai't Formation, Jbel El Mrakib, Ma'der Basin, southern Morocco(UA13700) (complete specimen). 1, dorsal view;X2.6.2, anterodorsal view; X2.2. 3, anterior view of cephalon; X2.3. 4, dorsal view of cranidium; X4.1. 5, dorsal view of pygidium; X4.2. 6, lateral view; XI.8. 297 List of Appendices

Appendix 1. Numerical calculations for all examined specimens of the Tropidocoryphinae. 315

Appendix 2. MacClade 4.06 (Maddison and Maddison, 2001) derived data matrix for the examined species of the Tropidocoryphinae. 317

Appendix 3. Numerical calculations for all examined specimens of the Cornuproetinae. 319

Appendix 4. MacClade 4.06 (Maddison and Maddison, 2001) derived data matrix for the examined species of the Cornuproetinae. 322 Chapter 1: Introduction

Introduction

The Devonian period was an extremely important interval of Earth's history for many taxa and communities, with many groups reaching their maximum Paleozoic diversity before experiencing significant losses during the end-Devonian extinctions (Sepkoski et al. 1981). Throughout the Devonian, trilobites were extremely abundant, exhibiting large amounts of both morphological and geographic variation before, like many other taxa, experiencing substantial decline across the period's end extinctions. However, while many trilobite orders perished at the end of the Devonian, only proetids

(Class Trilobita, Order Protida) persisted, surviving another 100 million years until the end mass extinction (Fortey and Owens 1975).

Despite their Late Paleozoic success and survival across the end Devonian extinctions proetid relationships remain poorly understood, especially those of the

Lower and Middle Devonian. Enter Morocco, whose immense palaeontological wealth provides a vast array of Lower and Middle Devonian proetids (Richter &

Richter 1943, 1956; Alberti 1964,1966a, 1966b, 1967a, 1967b, 1969, 1970,1975,

1980,1981a, 1981b, 1981c, 1982a, 1982b, 1988; Pillet 1972; H. Alberti 1975a,

1975b; Feist & Orth 2000). Unfortunately, many previous Moroccan based studies have dealt with only poorly preserved (fragmented and disarticulated) and incomplete specimens, which severely hampers efforts to understand proetid relationships. It has not been until recently (Morzadec 2001; Fortey & Chatterton

1 2003; Schraut & Feist 2004; Chatterton et al. 2006; Gibb & Chatterton 2007,

2010; Lerosey-Aubril et al. 2007; McKellar & Chatterton 2009; Chatterton &

Gibb 2010) that better preserved (fully complete and articulated) Devonian trilobites have been described from Morocco, including proetids.

Using some of these exquisitely preserved Moroccan trilobites, this thesis describes new species and examines the relationships of select genera from two proetid subfamilies, Tropidocoryphinae and Cornuproetinae. Five genera,

Astycoryphe, Tropidocoryphe, Cornuproetus, Diademaproetus, and Pontoproetus new genus are discussed and the following new species are introduced:

Astycoryphe ditropidia, Tropidocoryphe taharajatensis, T. lahfirensis, T. sculptaridgiata, Diademaproetus rudimentus, D. issoumourensis, D. corrugatus,

D. auxiliairus, D. langus, Pontoproetus truncatus, and P. granulosus.

Furthermore, phylogenetic analyses of Tropidocoryphe-Astycoryphe and

Cornuproetus-Diademaproetus-Pontoproetus are undertaken to determine the relationships of the included genera.

Location & geology

Located in the northwestern region of Africa, Morocco is bounded by the

Atlantic ocean to the north west, the Mediterranean Sea to the north, and Algeria and Mauritania to the south and east (Figure 1-lFigure 1-2). Three separate sedimentary basins, within the southern Anti-Atlas region of Morocco, along the

2 Algerian border, are the primary regions of study: Tindouf Basin, Ma'der Basin, and Tafilalt Basin (Figure 1-2 to 1-5).

During the Early and Middle Devonian, present day North Africa is reconstructed as a warm temperate region located between 30° and 60° southern latitude (Scotese 2001). As part of North Africa, Morocco is considered part of the continental shelf of Gondwana and was typically submerged beneath warm epicontinental seas of the Rheic Ocean (Schraut 2000; Gibb & Chatterton 2007;

McKellar & Chatterton 2009). During the Early Devonian closure of the Rheic

Ocean, due to subduction along its northern and southern margins, drew

Euramerica, in the north, and Gondwana, in the south, closer together. Closing of the Rheic Ocean paused briefly during the Middle Devonian as minor sea-floor spreading occurred between Euramerica and Gondwana (Tait et al. 1997).

However, once sea-floor spreading stopped again in the Late Devonian,

Euramerica and Gondwana continued to draw closer together, positioning

Morocco into close association with Amorica (present day western and central

Europe).

In terms of sea-level, the Devonian period is generally considered a time of relatively high sea-level with finer resolution shifts throughout (Vail et al.

1977). Morocco itself witnessed 6 transgressive/regressive cycles in the Early

Devonian and another 6 cycles during the Middle and Late Devonian (Lubeseder et al. 2003; Ouanaimi & Lazreq 2008; McKellar & Chatterton 2009). Becker et al.

(2004) identified many of the regressive cycle packages throughout Morocco's

3 southern Anti-Atlas as a series of thick, resistant, and 'Rich' sandstone units. In contrast, the transgression cycle packages occur as a mixture of shales and shallow water carbonate facies, with the majority of trilobites being found within limestone deposits of the carbonate facies.

In terms of Devonian ocean currents, Morocco was subjected to the southern flow of what Heckle and Witzke (1979) deemed the "Subtropical Gyre," a counter clockwise current bathing Morocco in warm equatorial waters from eastern Euramerica and Amorica (Dopieralska 2009). In conjunction with the proposed Devonian Rheic Ocean currents, numerous similarities between

Moroccan and Bohemian faunas have been proposed (Alberti 1969; 1970; Snadjr

1980; Chlupac 1983; Chlupac et al. 2000; Schraut 2000; Schraut & Feist 2004;

Gibb & Chatterton 2007, 2010; Lerosey-Aubril et al. 2007; McKellar &

Chatterton 2009). Furthermore, classification of Morocco and Amorica within the

'Old World Province" during the Emsian, , and Givetian based upon brachiopod provinces (Johnson & Boucot 1973) indicates a correlation between the two regions. As a result of all the faunal similarities and proposed ocean currents, it is generally accepted that trilobite faunal interchange occurred between southeastern Euramerica, Amorica, and north western Gondwana, primarily Morocco, through much of the Devonian.

4 Examined localities

Tindouf Basin. The most western studied basin, the Tindouf Basin, is subdivided into two main study areas, the Dra Valley (Figure 1-3) and Assa.

The Dra Valley is located south of the town of Foum Zquid and contains within it two main localities. The first and largest locality within the Dra Valley is referred to as 'Zguilma,' and located 53 kilometers southeast of the town Foum

Zguid and part of Jbel Gara El Zguilma. The locality is composed of three trilobite bearing horizons, the upper Emsian horizons ZGEE1 and ZGEE2 (29°

42' 35.2" N and 06° 42' 10.2" W) and the Eifelian horizon ZGEE3 (29° 42' 18.5"

N and 06° 42' 11.5" W). All three trilobite bearing horizons are part of the

Timrhanrhart Formation which consists of nodular to layered limestone interbedded with calcareous shales (Hollard 1978; Bultynck and Hollard, 1980;

Bultynck & Walliser, 2000; El Hassani, 2004; Chatterton et al. 2006; Gibb &

Chatterton 2007, 2010; McKellar & Chatterton 2009).

A second locality within the Dra Valley, known as FZl, was also examined and is found north of Jbel Hamsai at 29° 55' 41.5" N and 07° 03' 15.5" W. FZl displays a similar trilobite fauna as the horizons at Zguilma, and is therefore thought to be of similar late Emsian age and part of the Timrhanrhart Formation

(Gibb & Chatterton 2010).

The third and final examined locality, positioned outside of the Dra Valley but still within the Tindouf Basin, is located northwest of the town of Assa at 28°

30' 06.2" N and 09° 27' 43.4" W. Referred to as 'Assa,' the trilobite bearing

5 horizons at the locality are part of the Khebchia Formation and are late Emsian in age (Hollard 1978; Jansen et al. 2004)

Ma'der Basin. Within the Ma'der Basin attention was paid to the

Devonian outcrops (Figure 1-4), which are located approximately from 31° 08' to

30° 35' N and 05° 03' to 04° 30' W. Localities within this basin include: Jbel

Oufatene, Taharajat, Jbel Issoumour, Timarzite, bou Dib, Jbel El Mrakib, Jbel

Zireg, and El Achana.

Jbel Oufatene is located west northwest of the village Lahfira, in the southwest portion of the Ma'der Basin. The stratigraphically highest trilobite bearing horizon surveyed at Jbel Oufatene is the Diademaproetus horizon (30° 45'

48.6" N and 04° 40' 42.7" W), which is part of the El Otfal Formation and

Eifelian in age (Gibb & Chatterton 2010). Approximately 25 meters below the

Diademaproetus horizon and still within the Eifelian aged El Otfal Formation is the Philonyx/Quadrops horizon (30° 49' 07.0" N and 04° 53' 42.5" W).

Taharajat is located south of Jbel Oufatene, at 30° 47' 48.0" N and 04° 54'

20.3" W, and contains two main trilobite-bearing horizons, the Morocconites horizon and the Hollardops horizon. Because both horizons are close to each other geographically and only separated by three meters stratigraphically, while having extremely similar trilobite faunas, specimens from both horizons are treated as a single collection for the entire locality. Combination of both horizons into a single collection is also helpful in cataloging purchased specimens as the matrices from both horizons are identical and cannot be used to determine which horizon the

6 specimens came from, and Berber miners of this locality seldom note which horizon they collected specimens from. Both horizons are part of the Tazoulai't

Formation, and based on their trilobite and conodont faunas, are late Emsian in age (Gibb & Chatterton 2010).

Jbel Issoumour is located northwest of of the village of bou Dib and north northwest of Jbel Oufatene and Taharajat. Four trilobite bearing horizons along the northern and western face of outcrop were examined. The stratigraphically highest horizons examined are located at the top of Jbel Issoumour (30° 58' 25.2"

N and 04° 01' 49.7" W) are the Lobopyge and Paralejurus horizons of the El Otfal

Formation (Morzadec 2001; Chatterton & Gibb 2010; Gibb & Chatterton 2010).

Below the top of Jbel Issoumour, in the Er Remlia Formation, the Heliopyge/

Kayserops and, slightly stratigraphically lower, the Psychopyge horizon (30° 58'

25.2" N and 04° 01' 52.3" W) were also examined.

Timarzite is located east-northeast of the village of bou Dib, just north of the Number 12 (N12) highway and west-southwest of the town of Mecissi (31°

10' 40.7" N and 04° 58' 55.7" W). The locality is thought to be late Emsian to

Eifelian in age.

Bou Dib is located east of Jbel Issoumour and a few kilometers north of the village of bou Dib, the Harpes/Thysanopeltis horizon was examined. The

Eifelian aged Harpes/Thysanopeltis horizon is located at 31° 05' 16.6" N and 04°

52' 32.2" W, and part of the of the El Otfal Formation (Gibb & Chatterton 2007,

2010).

7 The only examined trilobite bearing horizon examined at Jbel Mrakib consists of the Emsian aged Morocconites/Metacanthina horizon of the Tazoulai't

Formation (Gibb & Chatterton 2010).

Jbel Zireg is located southeast of Jbel Mrakib, in the southeastern portion of the Ma'der Basin. The only horizon examined from Jbel Zireg in this study is the Eifelian aged Thysanopeltis horizon (Gibb & Chatterton 2010), located at 30°

36' 46.1" N and 04° 32' 10.1" W.

El Achana is located at 30° 51' 17.2" N and 04° 58' 35.0" W, with both the

Kolihapeltis horizon and the stratigraphically lower Dicranurus horizon being examined in this study. Both horizons are in age.

Tafilalt Basin. The third and final examined basin, the Tafilalt Basin

(Figure 1-5), is the most eastern of all three examined basins. Only one locality was examined in the Tafilalt basin, Talawarite.

Talawarite is located east-southeast of Hamar Laghdad (Alberti 1969), at

31° 16' 28.5" N and 03° 53' 29.4" W (Gibb & Chatterton 2010). Specimens examined from this locality were collected from both the Gerastos horizion and surrounding surface area. The Gerastos horizon itself is part of the Amerboh

Group and late Emsian in age (Gibb & Chatterton 2010).

Purpose and scope

This thesis can be considered part of the ongoing work of B. D. E.

Chatterton's research group, examining the systematics and relationships of

8 Lower and Middle Devonian Moroccan trilobites. In particular, this thesis examines select genera of the proetid subfamilies Tropidocoryphinae and

Cornuproetinae. In total, five genera, Astycoryphe, Tropidocoryphe,

Cornuproetus, Diademaproetus, and Pontoproetus new genus, are discussed and the following new species introduced: Astycoryphe ditropidia, Tropidocoryphe taharajatensis, T. lahfirensis, T. sculptaridgiata, Diademaproetus rudimentus, D. issoumourensis, D. corrugatus, D. auxiliairus, D. langus, Pontoproetus truncatus, and P. granulosus. Furthermore, phylogenetic analyses of Tropidocoryphe-

Astycoryphe and Cornuproetus-Diademaproetus-Pontoproetus are undertaken to determine the relationships of the included genera.

Motivation for this work is multi-faceted. First, previous description of

Moroccan trilobite material, primarily by Alberti (1969,1970), focused primarily on material from the northern and eastern portions of Morocco. Furthermore, the majority of previous works only described partial (incomplete and disarticulated) remains and made little effort to understand the relationships of the Moroccan taxa with similar taxa of the Devonian Rheic Ocean. Use of such material, at such a limited palaeogeographic scale, has resulted in more than one species name being proposed for different sclerites of the same or species originally found in other countries. This work, while sharing some localities of Alberti's

(1969,1970) work, attempts to rectify many of these problems for the examined taxa by utilizing exquisitely presevered (complete and articulated) specimens and comparing them with similar taxa on a larger paleogeographic scale.

9 Second, another important goal of this thesis is to locate accurately all of the described Moroccan species, both geographically and stratigraphically.

Previous Moroccan trilobite work (e.g. Alberti 1969,1970) mostly neglected both geographic and stratigraphic positioning of samples, often only providing vague locality details. This thesis in large part aims to rectify this by providing more accurate geographic and stratigraphic positions of localities and specimens, as a useful framework for future biostratigraphic and biogeographic correlations.

Third, the phylogenetic analyses of the genera examined in this thesis aim to produce not only a better understanding of the relationships between genera, but also a better understanding of the genera and species themselves. For instance, many of the examined taxa share numerous diagnostic characters with one another, or lack suitable diagnostic characters altogether. This confusion surrounding diagnostic characters has ultimately led to improperly diagnosed taxa and abundant "over-splitting," primarily at the generic level. This thesis statistically examines and discusses both new and previously proposed diagnostic characters of all examined taxa, and in conjunction with the phylogenetic analyses provides proper characters, character states, and diagnoses of taxa. Ultimately, these findings will make the proper identification and naming of new taxa and specimens easier and more useful in the future.

10 Previous research

Although a substantial amount of research has been accomplished regarding the classification of Devonian trilobites worldwide, southern Moroccan proetids have received little attention. In recent years, thanks primarily to B. D. E.

Chatterton's research group, some work has been accomplished on select groups of Lower and Middle Devonian Moroccan proetids (Chatterton et al. 2006; Gibb

& Chatterton 2007, 2010; Chatterton & Gibb 2010). However, there still remains a large number of proetids from southern Morocco in need of work, and this thesis attempts to resolve part of that issue.

Alberti (1969, 1970) originally published on Moroccan trilobites, most significantly describing proetids from the southeastern, Hamar Laghdad, and northwestern regions of the country. Preceding Alberti's Moroccan work, Richter

& Richter (1943,1956) also worked on some Moroccan proetids. Unfortunately, both the works of Alberti (1969, 1970) and Richter & Richter (1943,1956) utilized primarily incomplete (disarticulated and/or fragmental) and poorly preserved specimens. Although some of the specimens examined in this thesis are from similar geographic areas that Alberti (1969,1970) described specimens from, none of the specimens described in this thesis display synapomorphies correlating them with those described by Alberti (1969, 1970).

For more information regarding previous research of the examined subfamilies of this thesis, refer to the "previous research" sections of thesis

11 chapters two and three, where they are dealt with on a subfamily by subfamily basis.

Format

This thesis follows the paper format of thesis submission. Editorial formatting of the individual chapters follows that of the Journal of Systematic

Palaeontology, with each chapter containing specific locality and repository information for the examined specimens. Upon completion, chapters 2 (New tropidocoryphid trilobites (Order Proetida) from Morocco and their influence on our understanding of tropidocoryphid relationships) and 3 (Cornuproetus and

Diademaproetus (Class Trilobita, Order Proetida) from Morocco and their impacts in the cornuproetid, diademaproetid relationship) will both be submitted for publication.

12 Mediterranean M North Atlantic o«« I 1 Ocean

Morocco

South Atlantic Ocean Indian Ocean

0 500 1000 km

Figure 1-1. Map of Africa indicating Morocco (in-filled area) (modified from Philip (1991); McKellar & Chatterton (2009); and Gibb & Chatterton (2010)).

13 -»Taflalt Basin

Hamar l-oum Zguid " ' .RiMam "*®» ==^1" ALufcxiA 3%~ «SL- • i V© '

Hercynian Dolente (Upper Palaeozoic)

Devonian

0 5 10 km

Tindouf Basin -) Ma'der Basin RmeSS

&

Figure 1-2. Map of Morocco indicating locations of basins surveyed in this study (images modified from Fetah et al. 1998; Gibb 2005; Chatterton et al. 2006; Gibb & Chatterton 2007,2010; McKellar & Chatterton 2009; Chatterton & Gibb 2010).

14 Legend

Orfovkian . Devonian Hercynian Dolerite (Upper Palaeozoic)

Figure 1-3. Map and stratigraphy of select localities of the southern Moroccan Tindouf Basin (image adapted from Fetah et al. 1998; Gibb 2005; Chatterton et al. 2006; Gibb & Chatterton 2007, 2010; McKellar & Chatterton 2009; Chatterton & Gibb 2010).

15 Tlmarzite

&

V' s

5 10km I I Legend

1 = bou DTb 4 = Oufat6ne Devonian 2 = El Achana 5 = Mrakib 3 = Issoumour West 6 = Zireg

Figure 1-4. Map and stratigraphy of select localities of the southern Moroccan Ma'der Basin (image adapted from Fetah et al. 1998; Gibb 2005; Chatterton et al. 2006; Gibb & Chatterton 2007,2010; McKellar & Chatterton 2009; Chatterton & Gibb 2010).

16 Hamar Laghdad Talawarite Rissani

Merzouga

Legend • Devonian

Figure 1-5. Map and stratigraphy of select localities of the southern Moroccan Tafilalt Basin (image adapted from Fetah et al 1998; Gibb 2005; Chatterton et al. 2006; McKellar & Chatterton 2009; Gibb & Chatterton 2010).

17 Literature cited

Alberti, G. K. 1964. Neue Trilobiten aus dem Marokkanischen und deutschen Unter- und Mitteldevon. Senckenbergiana lethaea, 45, 115-132, pis. 16-17. Alberti, G. K. 1966a. Note preliminaire sur quelques trilobites (en particulier de Proetides) du Silurien, du Devonien inferieur et du Devonien moyen du Maroc. Notes du Service Geologique du Maroc, 26, 55-69. Alberti, G. K. 1966b. Uber einige neue Trilobiten aus dem Silurium und Devon, besonders von Marokko. Senckenbergiana lethaea, 47, 111-121. Alberti, G. K. 1967a. Neue obersilurische sowie unter- und mittledevonische Trilobiten aus Marokko, Deutschland und einigen anderen europaischen Gebieten. I and II. Senckenbergiana lethaea, 48(5), 463-479, 481-509. Alberti, G. K. 1967b. Neue obersilurische sowie unter- und mitteldevonische Trilobiten aus Marokko, Deutschland und einigen anderen europaischen Gebieten. II. Senckenbergiana lethaea, 48(6), 481-479,481-509. Alberti, G. K. 1969. Trilobiten des jiingeren Siluriums sowie des Unter- und Mitteldevons. I. Mit Beitragen zur Silur-Devon-Stratigraphie einiger Gebiete Marokkos und Oberfrankens. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 510, 1-692. Alberti, G. K. 1970. Trilobiten des jiingeren Siluriums sowie des Unter- und Mittledevons. II. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 252,1-233. Alberti, G. K. 1975. Zur palaozoogrographischen Verbreitung von Warburgekka rugulosa (ALTH 1874) (Trilobitae, Devon-Basis). Mitteilungen aus dem Geologis-Palaeontologischen Intstitut der Universitaet Hamburg, 44, 1-10. Alberti, G. K. 1980. Neue Daten zur Grenze Unter-/Mittel-Devon, vomehmlich aufgrund der Tentaculiten und Trilobiten im Tafilalt (Se-Marokko). Neues Jahrbuch fur Geologie undPalaontologie. Monatshefte, 39, 581-594. Alberti, G. K. 1981a. Beziehungen zwischen "herzynischen" Trilobiten-Faunen aus NW-Marokko und Deutschland (Unter- und Mittle-Devon). Natur und Museum, 111(11), 362-369. Alberti, G. K. 1981b. Scutelluidae (trilobita) aus dem Unter-Devon des Hamar laghdad (tafilalt, SE-Marokko) und das Alter der "mud- mounds" (Ober-Zlichovium bis tiefstes Dalejum). Senckenbergiana Lethaea, 62, 193-204. Alberti, G. K. 1981c. Trilobiten des jiingeren Siluriums sowie des unter-und Mittel-Devons III: mit Beitragen zur Devon-Biostratigraphie (insbesondere nach Nowakiidae) in N-Afrika, Sardinien, Oberfranken und im Harz. Senckenbergiana lethaea, 62, 1-75.

18 Alberti, G. K. 1982a. Der Hamar-Laghdad (Tafilalt, SE-Marokko), eine bedeutende Fundstatte devonischer Trilobite. Natur und Museum, 112(6), 172-182. Alberti, G. K. 1982b. Zur Frage einer Emersion der noerdlichen NW-Sahara am Ende des Lochkoviums (Unter-Devon). Newsletters on Stratigraphy, 11(1), 8-16. Alberti, G. K. 1988. Stratigraphische Untergliederung einiger Lochkovium/Pragium-Grenzprofile (Unter-Devon) in NW-Marokko, Oberfranken und Victoria (SE-Australien) auf Grand von Dacryoconariden und Conodonten. Senckenbergiana lethaea, 68, 479-493. Alberti, H. 1975a. Neue Trilobiten (Waribole) aus dem Ober-Devon IV-V (Nord-Afrika und Mittle-Europa); Beitrag 3. Neues Jahrbuch fur Geologie undPalaeontologie. Abhandlungen, 149(2), 180-210. Alberti, H. 1975b. Neue Trilobiten (Waribole) aus dem Ober-Devon IV-V (Nord-Afrika und Mittle-Europa); Beitrag 4, Nebst Revision einiger Typus-Spezies der Salter 1864, aus dem Ober-Devon II-V. Neues Jahrbuch fiir Geologie und Palaontologie. Abhandlungen, 150(2), 207-226. Becker, T.R., Jansen, U., Plodowski, G., Schindler, E., Aboussalam, S.Z., and Weddige, K. 2004. Devonian litho- and biostratigraphy of the Dra Valley area - an overview. In: El hassani, A. (ed.) Devonian neritic-pelagic correlation and events in the Dra Vallery (western Anti-Atlas, Morocco). Documents de I'Institut Scientifique. 19, 3-18. Bultynck, P. & Hollard, H. 1980. Distribution Comparee de Conodonts et Goniatites Devoniens de Plaines du Dra, de Ma 'der et du Tafilalt (Maroc). Leuven University Press, Belgium, 73 pp. Bultynck, P. & Walliser, O.H. 2000. Devonian boundaries in the Moroccan Anti-Atlas. Courier Forschungsinstitut Senckenburg, 225,211-226. Chatterton, B.D.E., Fortey, R., Brett, K., Gibb, S., and McKellar, R. 2006. Trilobites from the upper Lower to Middle Devonian Timrhanrhart Formation, Jbel Gara el Zguilma, southern Morocco. Palaeontographica Canadiana, 25, 1-177. Chlup&c, 1.1983. Trilobite assemblages in the Devonian of the Barrandian area and their relations to palaeoenvironments. Geologica et Palaeontologica, 17,45-73. Chlupac, I., Feist, R., & Morzadec, P. 2000. Trilobites and standard Devonian stage boundaries. Courier Forschungsinstitut Senckenburg, 220, 87-98. Dopieralska, J. 2009. Reconstructing seawater circulation on the Moroccan shelf of Gondawana during the Late Devonian: evidence from Nd isotope composition of conodonts. Geochemistry, Geophysics, Geosystems, 10(3), 1-13.

19 El Hassani, A. (ed.). 2004. Devonian neritic-pelagic correlation and events in the Dra Valley (western Anti-Atlas, Morocco). Subcommision on on Devonian Stratigraphy. International Meeting on Stratigraphy, Rabat, 1-10 March 2004. Documents de I'Institut Scientifique, 19,1-100. Feist, R. & Orth, B. 2000. Trilobites de la limite Eifelien/Givetien de la region stratotypique (Tafilalt, Maider, Maroc). Pp. 78-91 in A. Tahiri and A. El Hassani (eds.) Proceedings of the Subcommission on Devonian Stratigraphy (SDS) — IGCP 421 Morocco Meeting. Volume 20. Travaux de L 'Institut Scientifique Serie Geologie & Geographie Physique, Rabat, Morocco. Fetah, S. E., Bensaid, M., and Dahmani, M. 1988. Carte Geologique Maroc: Todhra-Ma'der (Anti-Atlas oriental, zones axiale et peripherique Nord du Sud). Editionas de Service Geologique de Maroc, Notes et Memoires No. 243. Royaume du Maroc. Ministere de l'Energie et des Mines, Rabat. Fortey, R. A. and Chatterton, B. D. E. 2003. A Devonian trilobite with an eyeshade. Science, 301, 1689-1689. Fortey, R. A. and Owens, R. M. 1975. Proetida - a new order of trilobites. and Strata, 33, 165 - 222. Gibb, S. 2005. Some proetids (Class Trilobita) from the Lower to Middle Devonian of southern Morocco. M.Sc. thesis, University of Alberta, Edmonton, Alberta, Canada. 211 pg. Gibb, S. & Chatterton, B. D. E. 2007. Timsaloproetus new genus (Proetida: Trilobita) and included species from Lower and Middle Devonian strata of southern Morocco. Journal of , 81(2), 352-367. Gibb, S. & Chatterton, B. D. E. 2010. Gerastos (Order Proetida; Class Trilobita) from the Lower and Middle Devonian of the Southern Moroccan Anti-Atlas region. Palaeontographica Canadiana, 30, 89pg. Heckle, P. H. & Witzke, B. J. 1979. Devonian world palaeogeography determined from distribution of carbonates and related lithic palaeoclimatic indicators; the Devonian System. Special Papers in Palaeontology, 23,99-123. Hollard, H. 1978. Correlations entre niveaux a brachiopodes at a goniatites au voisinage de la limite Devonien inferieur- Devonien moyen dans les plaines du Dra (Maroc presaharien). Newsletters on Stratigraphy, 7(1), 8-25. Jansen, U., Becker, G., Plodowski, G., Schindler, E., Vogel, O. & Weddige, K. 2004. Pragian and Emsian near Aouinet Torkoz (SW Dra Valley, Morocco). Pp. 75-84 in A. El Hassani (ed) Devonian Neritic-Pelagic Correlation and Events in the Dra Valley (Western Anti-Atlas, Morocco). Subcommission on Devonian Stratigraphy. International Meeting on Stratigraphy, Rabat, 1-10 March 2004. Documents de I'Institut Scientifique, 19.

20 Johnson, J. G. & Boucot, A. J. 1973. Devonian Brachiopods. Pp. 89-96 in A. Hallam (ed.) Atlas of Palaeobiogeography. Elsevier Scientific Publishing Company, Amsterdam. Lerosey-Aubril, R., Feist, R., and Chatterton, B. D. E. 2007. The ontogeny and systematics of the otarionine trilobite Otarionella from the Devonian of the Montagne Noir, France and the Maider, Morocco. Geological magazine, 145(1), 55-71. Lubeseder, S., Carr, I. D., & Redfern, J. 2003. A third-order sequence stratigraphic framework for the Devonian of Morocco: its implications for enhanced regional correlation of the Devonian of North Africa. AAPG Hedberg Conference, 1-4. McKellar, R. & Chatterton, B. D. E. 2009. Early and Middle Devonian Phacopidae (Trilobita) of southern Morocco. Palaeontographica Canadiana, 28,1 -109. Morzadec, P. R. 2001. Asteropyginae trilobites from the Devonian of the Anti-Atlas (Morocco). Palaeontographica Abteilung A, 262,53 - 85. Ouanaimi, H. & Lazreq, N. 2008. The 'Rich' group of the Draa Basin (Lower Devonian, Anti-Atlas, Morocco): an integrated sedimentary and tectonic approach. In: Ennih, N. & Liegeois, J. P. (eds.) The boundaries of the West Africa craton. Geological Society, London, Special Publications, 297, 467-482. Philip, G. 1991.Political: Africa, Philip's World Atlas. Reed International Books Ltd., London. Pillet, J. 1972. Les Trilobites du Devonien inferieur et du Devonien moyen du Sud-Est du Massif armoricain. Societe D 'Etudes scientifiques de L 'Anjou. Memoire, 1,1 - 307 & 364. Richter, R. & Richter, E. 1943. Studiem im Palaeozoikum der mittelmeer-laender; 4a, Trilobiten aus dem Devon von Marokko, mit einem Anhand iiber Arten des Rheinlands. Senckenbergiana, 26(1-3), 116-199. Richter, R. & Richter, E. 1956. Annular-Teilung bei Trilobiten am Beispiel besonders von Proetus (Pr.) cuvieri un prox. Senckenbergiana lethaea, 37 (3-4), 343-381. Schraut, G. 2000. Trilobiten aus dem Unter-Devon des siidostlichen Anti-Atlas, Siid-Marokko. Senckenbergiana lethaea,79(2), 361-433. Schraut, G. & Feist, R. 2004. The Devonian styginid trilobite Paralejurus, with new data from Spain and Morocco. Journal of Paleontology, 78(4), 709-722. Scotese, C. R. 2001. Paleogeographic Atlas, PALEOMAP Progress Report 90-0497. Department of Geology, University of Texas at Arlington, Arlington, Texas. 45 p. Sepkoski, J. J., Bambach, R. K., Raup, D. M. & Valentine, J. W. 1981. Phanerozoic marine diversity and the record. Nature, 293,435-437. §najdr, M. 1980. Bohemian Silurian and Devonian Proetidae (Trilobita). Rozpravy Ustredniho ustavu geologickeho, 45, 1-324.

21 Tait, J. A., Bachtadse, V., Franke, W., and Soffel, H. C. 1997. Geodynamic evolution of the European Variscan Fold Belt: palaeomagnetic and geological constraints. Geologische Rundschau, 86(3), 585-598. Vail, P. R., Michum, R. M., Todd, R. G., Widmier, J. M., Thompson, J., Sangree, J. B., Bubb, J. N., and Haleid, W. G. 1977. Seismic stratigraphy and global changes of sea level. In: Payton, C.E. (ed.) Seismic Stratigraphy - Applications to hydrocarbon exploration. AAPG Memoir, 26,49-213.

22 Chapter 2: Systematic Palaeontology

New tropidocoryphid trilobites (Order Proetida) from Morocco and

their influence on our understanding of tropidocoryphid relationships.1

Introduction

Specimens of Tropidocoryphe Novak, 1890 and Astycoryphe Richter &

Richter, 1919 are rare in Lower and Middle Devonian strata. Despite their scarcity, species of these genera and their relatives have been well documented from the Middle and Upper Devonian strata of the Montagne Noir region of

France (Feist 1976, 2003; Feist & Clarkson 1989). Their previous research

proposed that evolution of these genera involved eye reduction and eventual

blindness due to a habitat shift from near-shore, high-photic-level marine

environments to deeper-water, lower-photic-level environments (Feist & Clarkson

1989). Besides intense study in particular regions, such as in France (Feist 1976,

2003; Feist & Clarkson 1989), examination of tropidocoryphid evolution on broader palaeogeographic and taxonomic scales has not been attempted.

Furthermore, quantitative description and statistical testing of synapomorphic character states and the relationships they support have not been examined in conjunction with phylogenetic analysis. This has been the case for both finer

(locality based) and coarser scaled analyses. As a result, the hypothesis that tropidocoryphid evolution is characterized by eye-reduction has never been tested

1 Upon completion of this thesis, this chapter will be submitted it the Journal of Systematic Palaeontology with the co-authors S. Gibb & B. D. E. Chatterton.

23 either quantitatively or on a larger palaeogeographic scale. Without statistically supported characters, character states, and a tested phylogeny, the description of new material and their relationships to known taxa, including assignment to genera are difficult and not always justifiable. Unjustifiable results at even the lowest taxonomic levels (i.e., genus and species) prevent larger scale relationships from being understood. As the old saying goes, "the product is the sum of all its parts."

This study surveys Early and Middle Devonian tropidocoryphid trilobites of the Tindouf and the Ma'der (Figure 2-1) basins of southern Morocco, particularly in relation to previously described European taxa, which were in close proximity to Morocco during the time period. The presence of tropidocoryphid trilobites in Morocco is not surprising, given the palaeogeographic distribution of similarly aged trilobite taxa found in Europe (Chatterton et al. 2006; Gibb &

Chatterton 2007, 2010; McKellar & Chatterton 2009;). Commercial mining of

Moroccan localities by the local Berbers has produced numerous exquisitely preserved (complete and fully articulated) specimens of Tropidocoryhe and

Astycoryphe. The discovery of such well-preserved specimens prompted the present revision of tropidocoryphid trilobites and their relationships from a numerical, statistical and phylogenetic standpoint, an impossible task until now because many previously described tropidocoryphid species were based upon disarticulated and far from complete exoskeletons.

24 The present study performs statistical analysis of both new and previously

discussed characters involved in tropidocoryphid evolution, with the goal of

discerning character and character state validity. Cladistic analysis using

statistically supported characters states is used to examine the relationships and

validity of five previously proposed tropidocoryphid genera: Astycoryphe Richter

& Richter, 1919, Longicoryphe Erben, 1966, Tropidocoryphe Novak, 1890,

Tropicoryphe Snajdr, 1977 and Bojocoryphe Snajdr, 1980. In conjunction with the

cladistic analyses, three new species of Tropidocoryphe (Tropidocoryphe

taharajatensis, T. lahfirensis, and T. sculptaridgiata) along with one new species

of Astycoryphe (Astycoryphe ditropidia) are proposed and described. Results of

these analyses and these new Moroccan specimens will improve our

understanding of the relationships among species of the subfamily

Tropidocoryphinae. It is expected that this work will play a significant role in understanding character selection, description and evolution within the Proetida.

Previous work

Work on tropidocoryphid trilobites commenced in the late 19th century in central European countries, primarily Germany and what is now the Czech

Republic. Many tropidocoryphid species were originally described as members of the genus Proetus Steininger 1831 (Barrande 1846, 1852; Hawle & Corda 1847;

Beyer 1869; Billings 1869; and Whidborne 1889). Following the description of

Tropidocoryphe and its type species Tropidocoryphe filicostata by Novak (1890),

25 morphologically similar species of Proetus were placed within Tropidocoryphe,

and later in the newly described tropidocoryphid genus Astycoryphe (Richter &

Richter 1919).

After a hiatus of work during the Second World War, Pribyl (1946) erected

the subfamily Tropidocoryphinae, which included both Tropidocoryphe and

Astycoryphe. Later, both genera were reported from regions outside their original

European localities, primarily from regions with close palaeogeographical

proximity to the Devonian Rheic Ocean (Alberti 1967, 1969, and 1970; Ormiston

1967; Cisne 1968; Wright & Chatterton 1988). Early work on Moroccan

tropidocoryphids was primarily published by G.K.B. Alberti (1967, 1969, 1970)

who, while providing broad taxonomic surveys of Silurian and Devonian trilobites

from numerous areas throughout the country and parts of Europe, described and

noted examples of both Tropidocoryphe and Astycoryphe. Also of interest were

the new genera Longicoryphe (Erben, 1966), Bojocoryphe (Snajdr, 1976) and

Tropicoryphe (Snajdr, 1977), which are similar in form to both Tropidocoryphe and Astycoryphe.

Work on the Middle and Upper Devonian strata of the Montagne Noir succession of France (Feist 1976) has been important in providing occurrence data for many new and previously known genera into the Upper Devonian. This work paved the way for Feist & Clarkson (1989) to provide the first Middle to Late

Devonian phylogeny of tropidocoryphid trilobites, which included Astycoryphe,

Tropidocoryphe, and Longicoryphe (Figure 2-2). Most importantly, the phylogeny

26 highlighted the enlargement of the preglabellar field, widening of the anterior portions of the cephalic sutures, and reduction of eye size, from stratigraphically

lower to stratigraphically higher taxa. Further work by Feist (2003), based on additional Montagne Noir material, provided more support to this phylogeny and the associated trends in character evolution. To date, however, despite documentation of new Astycoryphe and Tropidocoryphe (Morzadec 1969, 2001;

Pribyl & Vanek 1978; Ellerman 1992; Basse 1997, 2010; Frantisek & Vanek

2000; Chatterton et al. 2006; van Viersen et al. 2009; van Viersen & Prescher

2010) no attempt has been made to test the phylogeny and highlighted characters of Feist & Clarkson (1989) on a larger palaeogeographic scale outside the

Montagne Noir succession.

Material and methods

Specimens used in this study were collected during field work in Morocco by the authors in the spring of 2009 and from previous trips to the same areas by

B.D.E. Chatterton and his research group since 1995. All collected specimens were obtained either directly from trilobite-rich horizons or their associated spoil heaps generated by local fossil mining Berbers. Due primarily to specimen rarity, some specimens were also purchased from Berber miners or their associated fossil shops. Stratigraphic control of purchased specimens was obtained by questioning the miners who originally collected the specimens, and this information was checked by comparing the matrix of the purchased specimen with matrix collected

27 by us from the locality from which the specimen was reported. In all cases, identical matrix matches are present between purchased specimens and their presumed locality, and in most cases conspecific specimens to those purchased were found by us at the appropriate locality and horizon (even if these were not complete specimens).

Following collection, specimens were then shipped directly to the

University of Alberta, where they are now housed. Preparation of specimens at the

University of Alberta was completed using pneumatic chisels and pin-vice by the primary author or by Allan Lindoe, professional fossil preparator. In the case of specimens prepared by our local Moroccan guide and preparator, Hammi Ait

H'ssaine and his family, air propelled calcite abrasion was usually used.

Following preparation, specimens were painted black and coated with an ammonium chloride sublimate to enhance fine detail. A Canon 40d digital camera, with a 60 mm macro Canon EF lens, was then used to photograph the specimens.

Cladistic analyses were run using PAUP* version 4.0b 10 (Altivec)

(Swofford 2002). Results of the analyses were viewed using MacClade 4.08

(Maddison & Maddison 2005). All characters were equally weighted and all character states were unordered.

Stratigraphy

Sites from which specimens in this study were obtained are marked on a series of general and basin diagrams (Figures 2-1,2-3, & 2-4) and the sections

28 within these study sites are illustrated as stratigraphic columns (Figure 2-5). In

conjunction with these figures, individual sites are described in more detail in the

following basin subsections.

Tindouf Basin

Jbel Gara el Zguilma. Specimens of Tropidocoryphe amuri are found in the late Emsian and Eifelian trilobite-bearing horizons at Jbel Gara el Zguilma

(Chatterton et al. 2006), hereafter referred to as 'Zguilma,' of the Dra Valley region. All horizons are part of the Timrhanrhart Formation (Hollard 1978;

Bultynck & Hollard 1980; Bultynck & Walliser 2000; El Hassani 2004) within the

Anti-Atlas region of Morocco. The Late Emsian horizons (ZGEE1 and ZGEE2) are located at 29°42'35.2"N and 06°42'10.2"W, 53 km south southeast of the town

Foum Zguid, and composed of calcareous shale and nodular calcareous mudstone

(Gibb & Chatterton 2010, table 1). The Eifelian horizon (ZGEE3) is located at

29°42'18.5"N and 06°42'11.5"W, and is composed of calcareous mudstone in horizontally laminated shale (Gibb & Chatterton 2010, table 1). Original description of the trilobite faunas and stratigraphy of all three horizons was detailed by Chatterton et al. (2006).

Assa. Tropidocoryphe amuri is found at the trilobite-bearing horizon Assa

2, which is located northwest of the town of Assa. Located at 28° 30' 06.2" N and

09° 27' 43.4" W, the locality of Assa is part of the Khebchia Formation and late

29 Emsian in age (Becker et al. 2004a; Becker et al. 2004b; Hollard 1978; Jansen et al. 2004).

Ma'der Basin

Jbel Oufatene. Specimens of Astycoryphe ditropidia sp. nov.,

Tropidocoryphe sculptaridgiata sp. nov., and Tropidocoryphe lahfirensis sp. nov. come from horizons south of Jbel Issoumour and west northwest of the village

Lahfira. Composed of a thick sequence of Lower and Middle Devonian strata, the locality contains a number of trilobite producing horizons. Both Astycoryphe ditropidia sp. nov. and Tropidocoryphe lahfirensis sp. nov. are found in the

Philonyx/Quadrops horizon, located at 30°49'07.0"N and 04°53'42.5"W.

Tropidocoryphe sculptaridgiata sp. nov. is found in the stratigraphically higher

Diademaproetus horizon, located at 30°50'21.3"N and 04°52'58.7"W. Both beds have a limestone composition, are part of the El Otfal Formation of the Ma'der

Basin (Morzadec 2001), and are Eifelian in age (Gibb & Chatterton 2010, table 1).

Taharajat. The section at Taharajat includes two trilobite-bearing horizons, a stratigraphically lower Morocconites horizon and a slightly stratigraphically higher Hollardops horizon (Metacanthina couche). Due to the small stratigraphic separation (3m), similar limestone lithology, and similar biota of both horizons, specimens are treated as a single collection for the entire locality. This helps in cataloging purchased specimens, as the matrices from both horizons are similar and hamper proper horizon identification. Located at

30 30°47'48.0"N and 04°54'20.3"W, just south of Jbel Oufatene, the locality's trilobite and conodont fauna place it within the late Emsian, Tazoulai't Formation

(McKellar & Chatterton 2009; Gibb & Chatterton 2010). It is the only locality from which Tropidocoryphe taharajatensis sp. nov. has been obtained.

Terminology

Morphological terminology used herein follows that of the revised Treatise on Invertebrate Palaeontology (Whittington & Kelly 1997). Proetid specific terminology is adapted from Owens (1973), Snajdr (1980), and Richter & Richter

(1949). Cephalic and pygidial measurements follow those of Owens (1973), while terminology of overall shape of sclerites, tagma and their features follows Snajdr

(1980). Cephalic suture terminology follows Richter & Richter (1949). Refer to

Figure 2-6 for placement and location of all measurements and positions used in this study. Anatomical directions are used where appropriate, with distal and proximal used in reference to a feature's proximity to the mid-line (sagittal line) or center of the exoskeleton. Abbreviations are also used in accordance to those widely used in trilobite literature (e.g., Occipital lobe = LO). Lastly,

Tropidocoryphe specific characters, such as glabellar furrow shape and tropidial strength are adapted from Chatterton et al. (2006).

New terminology used in this study includes eye length (B) and width of the posttropidial field (As, Figure 2-6). Eye length is a measurement of the maximum exsagittal length of the eye lens field. Lastly, length of the posttropidial

31 field is the sagittal distance from the anteriormost point of the glabella to the

posteriormost point of the tropidium (posteriormost tropidial ridge), measured

along the cephalic midline (sagittal).

Phylogenetic analysis

The goals of this phylogenetic analysis are three-fold. First, the analysis serves as a test to group Moroccan specimens into species. In the case of the new

Moroccan material, numerous complete specimens coded the same. This provided increased confidence in the integrity of these species. For reasons of efficiency, only one similarly coded specimen was included in the analysis to represent the species. Second, the analysis examines the relationships of these species to one another and other well known European and marginal Devonian Rheic Ocean tropidocoryphid species. It was not designed to re-evaluate the taxonomy of previously diagnosed species or the relationships of well documented taxa. Third, following the work of Feist & Clarkson (1989), this analysis re-examines the phylogenetic relationships of Tropidocoryphe, Astycoryphe, and Longicoryphe.

Other genera were included to assess genus level validity and previously hypothesized relationships. Unfortunately within the tropidocoryphids, previous researchers have proposed numerous genera for a group that is neither particularly diverse nor particularly disparate. Generic descriptions have often been based upon incomplete specimens, and genera have often been discriminated

(diagnosed) by few distinctive apomorphies. To date, no attempt at a phylogenetic

32 analysis, or examination of previously proposed synapomorphies, has been made for tropidocoryphids.

Coding of the operational taxonomic units (OTUs) examined in this analysis was conducted using individual specimens, with previously described species being exemplified by either their holotype specimen or a specimen of greater completeness, quality and/or documentation. While use of phylogenetic analysis is not new to trilobite research (Lieberman & Karim 2010), use of individual specimens as OTUs is. Outside of trilobite research, use of specimen based OTUs is not new; Anstey & Pachut (2004) used specimen based OTUs in bryozoans to maximize the morphological disparity between species of the same genera for their analysis. Essentially, use of specimens as OTUs allows for examination of relationships from the lowest, to the highest, taxonomic levels.

Furthermore, use of specimen based OTUs permits new specimens to be diagnosed based on tree topology (where they plot out in relation to other specimens), not preconceived notions.

Examination of previously hypothesized synapomorphies and possible new characters of the examined taxa was accomplished in this study using geometric morphometries and statistics, something never done in tropidocoryphids, let alone trilobites. Such examination of characters, and thus their variability, is crucial towards understanding and delimiting taxon boundaries

(Cronier et al. 2011). Without well delimited boundaries, specimen classification becomes problematic, resulting in relationships becoming unclear and poorly

33 understood. By using morphometries and statistics, this study aims to provide a more current, reproducible, and accurate phylogeny of the examined taxa, given the material currently known and available.

Using the produced phylogeny, new specimens that formed well supported clades, while being distinctly different from other specimens, were considered as either new species or genera depending on their taxonomic level. Specimens which grouped together in the phylogeny as either polytomies or very weakly supported clades were considered similar and thus, part of the same taxon.

Inversely, clades separated by high Bremer support values were generally considered higher taxonomic units than clades separated by lower Bremer support values as they represent more stable groupings of OTUs. However, for the sake of stability, universality, and communication of nomenclature (ICZN 4th edition,

Article 23.2) judgement was used when using Bremer support values to modify previously documented taxonomic classification. Because this analysis is designed/optimized for discovering relationships among species of

Tropidocoryphe rather than related outgroup taxa, we are particularly conservative in proposing nomenclatural and taxonomic changes for outgroup taxa. Only in cases where the diagnoses of taxa proved extremely similar to the diagnoses of other taxa, or based upon indistinguishable characteristics, are taxonomic revisions proposed for examined taxa. In all, use of specimen based OTUs and statistically examined characters and character states in conjunction with

34 phylogenetic analyses allows for a more analytical approach toward naming and

diagnosing new specimens and taxa.

Taxa examined in the analysis

More than a dozen different genera, with over 50 species, have been

assigned to the Tropidocoryphinae (Snajdr 1980) and as described previously,

many of these taxa are problematic. In addition, because of the poor quality of the

majority of the published material, only a limited number of informative

characters can be coded. This low character to taxon ratio decreases the likelihood

of producing a robust phylogeny for all of these taxa, which Wagner (2000)

described as character exhaustion. In order to avoid this problem, the number of

taxa examined at both the genus and species levels needed to be limited (Table

2-1).

One of the primary goals of the present study is to determine the

membership of particular individuals and species within a small group of genera,

inclusion of taxa is based primarily upon the morphological similarity of taxa to

Tropidocoryphe. At the generic level, given the close relationship and historically problematic taxonomy of Tropidocoryphe, Astycoryphe, and Longicoryphe, all

three genera are represented in the analysis. Bojocoryphe and Tropicoryphe are also included, as they are similar and many of their species have at one time or another been assigned to either Tropidocoryphe or Astycoryphe. Furthermore, the diagnoses and descriptions of both Bojocoryphe and Tropicoryphe (Snajdr 1967,

35 1980) are nearly identical to that of Astycoryphe. Lastly, Erbenicoryphe and

Pterocoryphe are included in the analysis to test if the previously proposed

tropidocoryphid relationships of Feist & Clarkson (1989) hold true with additional

new taxa, a larger palaeogeographic scale, and quantified characters. All

remaining tropidocoryphid genera are omitted.

Further omission of taxa, specifically at the species level, was based

primarily on two additional criteria. First, taxa not within close palaeogeographic

proximity to the Devonian Rheic Ocean were omitted as they are all poorly

documented, highly incomplete, and described using poorly preserved material.

Secondly, highly incomplete taxa (less than 50% coded) from within the Devonian

Rheic Ocean were also omitted. Though it has been argued that highly incomplete

taxa are not as detrimental to phylogenetic analyses as are low character numbers

(Wiens 2003), if the distribution of missing data is localized to several taxa the

effects on the analysis can be highly detrimental (Prevosti & Chemisquy 2010).

Furthermore, if a high percentage of missing data is allocated to a single character

or group of characters, character exhaustion can occur. This occurs when a high percentage of incomplete taxa in an analysis prevents characters from being coded for a large portion of the examined taxa. As a result, the characters themselves become highly incomplete, containing many Highly incomplete characters cannot typically be statistically supported and, in the case of this analysis, are also reported as parsimoniously uninformative by the analysis itself, therefore they were omitted in order to produce a more robust phylogeny. In the present study,

36 initial analyses found that omitted taxa typically had issues with respect to both

criteria, and their omission resulted in a more robust phylogeny.

Outgroup taxa were chosen based upon the geologically oldest taxa. As a

result, the Pragian taxa of Bojocoryphe, Tropicoryphe, and Astycoryphe (A.

gracilis) were used as outgroup taxa in this analysis. While it has been argued that

Decoroproetus Pribyl, 1946 is an acceptable outgroup for more derived

tropidocoryphids (Owens 1973, text-fig. 11; Feist 1976, 2003; Snajdr 1980; Feist

& Clarkson 1989), it was omitted from this analysis as, unlike Bojocoryphe and

Tropicoryphe, its diagnosis is notably different from that of Astycoryphe and its

presence in the analysis significantly destabilized the base of the tree while not

altering the overall tree topology. Secondly, because Astycoryphe and the

morphologically similar Bojocoryphe and Tropicoryphe have an earlier

stratigraphic first appearance (Pragian) than Tropidocoryphe (Emsian) and the

remaining examined taxa, only Pragian aged species of Astycoryphe,

Bojocoryphe, and Tropicoryphe were considered as outgroup taxa. Regardless of

the ancestry of tropidocoryphids, the purpose of this analysis is to test the

Astycoryphe-Tropidocoryphe-Longicoryphe relationship, not all the members of

the Tropidocoryphinae. Therefore Decoroproetus was not included as an outgroup for this analysis.

Characters and character states

37 Absence of a well supported character set for tropidocoryphid trilobites, and the Proetida in general, prompted the construction of a tropidocoryphid specific matrix for this study. Many of the characters used herein are proetid characters discussed by Owens (1973) and Snadjr (1980). Others are reworked diagnostic characters for the genera and species studied herein, especially those highlighted by Feist & Clarkson (1989). Where no previously discussed characters were present, but quantitative and statistical examination suggests that there may be, new characters and their states were selected.

Reference taxa and diagrams are included for all characters where coding character states is subjective in terms of morphology; this provides clearer examples of character states such as "nose-like" and "cushion-like" mean in the context of the coded specimens. Of the 31 characters used in the study, 17 are binary and 14 are multistate in nature. Due to the partial and incomplete nature of previously described taxa and greater character richness of the cephalon, there is a significant bias towards cephalic characters (20) compared to pygidial characters

(10) and thoracic characters (1). The complete data matrix for this analysis is included as Appendix 1. Where taxa have been erected using types that are disarticulated and/or skeletons that are incomplete, cranidia are the most frequently described sclerites, followed by pygidial, then free cheeks (librigena), and then other sclerites from the thorax and the underside of the cephalon. A number of taxa have been based purely on cranidia and/or pygidia sclerites.

38 Illustrations of rostral plates, hypostomes and thoracic segments are not available for most named tropidocoryphid species.

Full explanation of all characters and their character states, including justification (statistical where appropriate) for their use, explanations of any modifications made to characters used in other studies, and examples for difficult characters and character states are outlined hereafter. Numerically based characters introduced in this study were established by measuring as many specimens as possible from all examined taxa, while specimen coding was based upon the best individual specimens from each taxon. Graphs were then generated on a character by character basis using measurements from all specimens and taxa examined, with character states distinguished between statistically supported taxon level populations. Thus, multiple specimens were used to examine and generate characters and character states, while the OTUs for the analysis remained individual specimens. For more information regarding character validity, coding and statistics, refer to the remarks section for each character hereafter.

Measurements for the purpose of coding of characters were taken from individual specimens whenever possible using digital calipers. If samples for coding were not available for direct measurement, measurements were taken from strictly dorsal images of specific elements using calipers and a ruler. All measurements, from either specimens or photographs, were taken three times on separate occasions and the average of the three was reported. If no dorsal image was present in the literature or obtainable from institutions where the specimens

39 were housed, measurements were not taken and characters associated with that

measurement were coded as unknown ("?"). Furthermore, unless characters were

clearly identifiable by the primary author (from either photograph or hand sample)

they were coded as unknown; description alone was not considered sufficient for

character state coding.

Statistical tests used to differentiate populations (character states) included

Mann-Whitney U test, ANOVA, Tukey Pairwise test, and Hotelling's t-test (with a

Bonferroni correction where appropriate), and were run using PAST (Hammer

2010). Each test was used depending on the number of populations (character states) that were present when examining all taxa. The Mann-Whitney U test,

ANOVA, and Tukey Pairwise test, were used in conjunction with single variable

(non-coordinate based morphometries) data. The Mann-Whitney U test, which is a non-parametric test that measures the equality of means between two populations, was used when only two populations (character states) were present. This test was

used instead of a standard Students t-test as the distribution and variance between populations was typically unequal, especially when examining gradational characters. Furthermore, the Mann-Whitney U test is considered a more conservative test than either the Students or Welch t-test. When more than two populations of single variable data were present, ANOVA tests were employed in conjunction with a Tukey Pairwise test to compare the means of multiple populations (more than two) at once and then individually. ANOVA tests are not ideal tests alone for multi-population scenarios as they only measure whether the

40 means of any of the populations are distinct. ANOVA tests, by themselves, may report a significant result even if only one of the populations is statistically different, but the others are not. Because of this, characters with three or more character states (populations) were examined using both an ANOVA test

(examining all character states at once) and a Tukey Pairwise test, which runs after the ANOVA test and compares all populations to one another individually, reporting back p-values of whether two individual populations are distinct from one another.

Lastly, a Hotelling's T-squared test was used to examine multi-variable

(coordinate based morphometries) data, and when more than three populations were present, a Bonferroni correction was applied. Hotelling's t-test is a multivariate analog of a Students t-test, measuring the equality of means between two populations in multivariate space. This test was employed when examining bivariate characters and reports both a probability value that the two means are distinct (p-value) and a percent correct classification value (%CC); the latter is a measure of the derived discriminant function to identify specimens to the correct population (Hammer 2010). When more than two populations (character states) were present, a Bonferroni correction was also used to adjust the p-value of individual population vs. population test. Both standard and sequential Bonferroni corrections reduce the individual population vs. population test threshold for rejecting the null hypothesis so that the overall threshold of comparing all populations to one another remains the same (Stafford & Leighton 2011). Because

41 a standard Bonferroni correction is viewed as a more conservative approach than that of a sequential Bonferroni, we only employed standard Bonferroni corrections to be as conservative as possible in determining valid characters and character states.

In all circumstances a confidence interval of 95% was used to determine statistical significance, and thus to separate character states. Furthermore, when multiple statistical tests were performed on a character, statistical support for the differentiation of all character states needed to be found by all tests. If any one of the employed tests could not statistically support differentiation of character states, then all character states and the character were omitted from the analysis.

Refer to Table 2-2 for the statistical breakdown of all tests on a character by character and character state by character state basis.

Cephalon

1. Pretropidial and tropidial area length: length of the pretropidial and

tropidial field, measured from the proximal-most tropidial ridge to the

anterior margin of the cephalon ((A2 + A3) - A5), in relation to (divided by)

the entire length of the cephalon (A, Figure 2-6). If no tropidium is

present, considered as 0.

States: (0) <0.15; (1) 0.16 - 0.25; (2) 0.26 - 0.31; (3) > 0.32

42 Remarks: Alberti (1970), Liitke (1980), and Snajdr (1980) identified preglabellar length as a diagnostic feature, at both the generic and species level, of tropidocoryphid trilobites. Feist & Clarkson (1989) further supported this contention, basing a large part of their proposed evolution of Montage Noir taxa on the lengthening and shortening of the preglabellar area in relation to the length of the entire cephalon. This character was included in the phylogenetic analysis in order to test its validity as a diagnostic character. Interestingly, coding of this character was identical to that of the preglabellar length vs. cephalon length.

Therefore, widening of the preglabellar area is the result of lengthening of only the pretropidial and tropidial areas of the preglabellar area. In order to prevent overweighting of the expansion or contraction of the preglabellar field, only the pretropidial and tropidial area length character was included in the final analysis.

For more information on preglabellar length vs. cephalon length, refer to

"characters not included in analysis".

2. Posttropidial area length: length of the posttropidial area (proximal-most

tropidial ridge to the anterior limit of the glabella (As) in relation to

cephalon length (A, Figure 2-6). If no tropidium present, considered as 0.

States: (0) < 0.05; (1)> 0.05.

43 Remarks: Snadjr (1980) diagnosed Bojocoryphe based on a number of factors, including a short frontal border and preglabellar field as a result of shortening of the posttropidial area, not the pretropidial or tropidial area. As such

the posttropidial field does not correlate with the size of the pretropidial and

tropidial areas, signifying its use as its own character. In order to test if this character is in fact diagnostic of Bojocoryphe, it was included in the analysis.

3. Preglabellar length vs. Glabellar anterior lobe length: length of the

preglabellar area (A2 + A3) divided by the length of the anterior lobe of the

glabella (Ai, Figure 2-6).

States: (0) < 0.84; (2) 0.85 - 1.05; (3) > 1.06.

Remarks: According to Snajdr (1980) glabellar length correlates to exoskeleton thickness in some proetids, with thin shelled species having a shorter glabella than thicker shelled taxa. However, glabellar length is also a diagnostic character of Erbenicoryphe (Feist 1976, 2003), so it was further examined here and included to determine its diagnostic usefulness.

4. Border shape: anterior and lateral borders of the cephalon appears

cushion-like (inflated and vaulted) or flat (no vaulting or inflation) (Figure

2-7).

44 States: (0) flat (non cushion-like); (1) inflated (cushion-like).

Remarks'. Best viewed in lateral or lateral-oblique view of the cephalon and appears as if the frontal border of the cephalon is slightly rolled or upturned in a 'cushion-like' form. Snadjr (1980) used this as a diagnostic character for both

Bojocoryphe and Tropidocoryphe. However, presence of the cushion-like border also occurs prominently in Astycoryphe and in some species of Longicoryphe, making its use as a diagnostic character questionable. Regardless, this character was included for it was a viable diagnostic character for a group of examined specimens.

5. Presence of border rims: presence of one or more fine parallel ridges

running around the anterior border of the cephalon, between the cephalic

margin and cephalic border furrow (Figure 2-8). Counted anterior of the

glabella, in between cephalic suture angles alpha (a).

States: (0) present; (1) absent.

Remarks: As the original description and diagnosis of Astycoryphe by

Richter & Richter (1919) primarily includes features now associated with

Longicoryphe (Erben 1966), Harrington (1959) re-diagnosed Astycoryphe using

45 the presence of border rims as a primary diagnostic feature compared to

Tropidocoryphe. However, since Astycoryphe is not the only tropidocoryphid to display border rims (e.g., Longicoryphe), this character was included to determine its diagnostic capabilities.

6. Position of cephalic suture angle p (Figure 2-9): position of cephalic

suture angle /? in relation to the length of the cephalon (A, Figure 2-6).

Equations used to differentiate character states are the product of the

discriminate analysis when preforming Hotelling T-tests using PAST

(Hammer 2010) and represent the midpoint between the discriminant

scores of generic based populations.

States: (0) narrow morph, where ((72.241*Xp + 20.716*Yp) - 34.559) <

0; (1) moderate morph, where ((72.241*Xp + 20.716*Yp) - 34.559) > 0

and ((-92.823*Xp - 161.63*Yp) + 115.09) > 0; (2) wide morph, where

((-92.823*Xp - 161.63*Yp) + 115.09) < 0.

Remarks'. Diagnoses of Astycoryphe and Tropidocoryphe usually emphasize Astycoryphe having narrowly divergent anterior portions of the cephalic suture, while Tropidocoryphe has highly divergent anterior cephalic suture portions. This character is an attempt to distinguish the degree of divergence at cephalic suture angle P mathematically.

46 Taken as a bivariate measurement, where Xp = P's distance from the mid­ line of the cephalon divided by cephalon length (A) and Yp = P's distance from the most anterior limit of the cephalon divided by cephalon length (A, Figure 2-6).

Graphically plotting the position of angle P using Xp and Yp, as X and Y coordinates respectively, and statistically testing generic level populations was used to determine if the different populations were statistically distinct from one another, and therefore viable character states (Figure 2-9).

7. Position of cephalic suture angle y (Figure 2-10): position of cephalic

suture angle y in relation to length of the cephalon (A, Figure 2-10).

Equations used to differentiate character states are the product of the

discriminate analysis when preforming the Hotelling T-test using PAST

(Hammer 2010) and represent the midpoint between the discriminant

scores of generic based populations.

iStates: (0) anterior morph, where ((-3.582*Xp + 36.856*Yp) - 18.577) > 0 ;

(1) posterior morph, where ((-3.582*Xp + 36.856*Yp) - 18.577) < 0.

Remarks: Similar to character 6, character 7 is an attempt to distinguish the degree of divergence in the anterior portion of the cephalic suture mathematically; examining angle y instead of angle p. Variance in this character however does not support distinction between narrowing or widening of the

47 cephalic suture pattern, but rather an exsagittal (anterior-posterior) shift in the location of the suture angle.

Taken as a bivariate measurement, where X Y = y's distance from the

cephalon mid-line divided by cephalon length (A) and Y Y = y's distance from the most anterior limit of the cephalon divided by cephalon length (A, Figure 2-6).

Graphically plotting the position of angle y using Xy and Yy, as X and Y coordinates respectively, and statistically testing generic level populations was used to determine if the different populations were statistically distinct from one another, and therefore viable character states (Figure 2-10).

8. Position of cephalic suture angle 8 : position of cephalic suture angle d

in relation to sagittal length of the cephalon (A, Figure 2-6). Equations

used to differentiate character states are the product of the discriminate

analysis when preforming the Hotelling T-test using PAST (Hammer 2010)

and represent the midpoint between the discriminant scores of generic

based populations.

States: (0) narrow morph, where ((-7.278*Xp + 82.674*Yp) - 56.178) < 0;

(1) wide morph, where ((-7.278*Xp + 82.674*Yp) - 56.178) > 0.

Remarks: Character 8 is another mathematical approach to distinguish the degree of divergence of the cephalic suture, specifically at angle d. Interestingly,

48 this character demonstrated posterolateral and anteromedial separation of

character states, not just lateral separation as previous studies suggested (Feist &

Clarkson 1989).

Taken as a bivariate measurement, where Xg = 8's distance from the

cephalon mid-line divided by cephalon length (A) and Yg = 8's distance from the

most anterior limit of the cephalon divided by cephalon length (A, Figure 2-6). Xg

and Ys were then both divided by cephalon length (A, Figure 2-6) and plotted as

X and Y coordinates to distinguish distinct populations. Graphically plotting the

position of angle 8 using Xg and Ys, as X and Y coordinates respectively, and statistically testing generic level populations was used to determine if the different populations were statistically distinct from one another, and therefore viable character states (Figure 2-11).

9. Number of tropidial ridges: number of tropidial ridges that are

comprised in the tropidium.

States: (0) 0; (1) 1; (2) 2; (3) > 3.

Remarks: The tropidium is a ridge or group of ridges that traverse the preglabellar field and free cheek, parallel to the frontal border (Owens 1973;

Snajdr 1980). Function of this feature remains poorly understood; however it occurs in a great variety of morphologies across numerous, commonly thin

49 shelled, genera. Number and size of tropidial ridges are diagnostic at the species level, and key features for such species as Tropidocoryphe mediterranea and

Bojocoryphe splendens (Feist 1976; Snadjr 1980).

10. Termination of tropidium: position on the librigena (free cheek) where

the tropidium terminates.

States: (0) tropidium terminates anterolateral of angle to of the cephalic suture; (1) tropidium terminates lateral of, or posterolateral of, angle to of cephalic suture.

Remarks: In conjunction with the number of ridges the tropidium is composed of, the robustness of the primary (longest) ridge can vary. More robust tropidia have more prominent ridges that terminate lateral, or posterolateral, of angle co of the cephalic suture. Inversely, taxa with a less robust tropidium have shorter andless prominent tropidial ridges that terminate anterolateral of angle co of the cephalic suture. Robustness of the tropidium was outlined by Chatterton et al. (2006) as a key diagnostic of Tropidocoryphe amuri, and was included to test its diagnostic capabilities amongst other specimens and species.

50 11. Presence of robust genal caeca: robust and prominent ("swelled") genal

caeca are present ahead of the glabella, and bifurcating at the tropidium as

they progress towards the anterior margin of the cephalon.

States: (0) present; (1) absent.

Remarks: Diagnostic feature of Tropidocoryphe outlined by Liitke (1980),

Feist & Clarkson (1989), and Kowalski (1990). Snajdr (1980) debated as to

whether the prominence of genal caeca is a result of either the trilobites increased

sensory system or as a product of a thinner exoskeleton.

12. Palpebral lobe shape: shape of the palpebral lobe when viewed dorsally.

States: (0) bean shaped (e.g., Tropidocoryphe amuri, Plate 2-2, fig. 1); (1)

crescent-shaped (e.g., Astycoryphe ditropidia, Plate 2-1, fig. 1).

Remarks: Shape of the palpebral lobe shows variance in multiple aspects.

Feist & Clarkson (1989) argued that the steepness is of importance; however

taphonomic influences in many specimens, primarily those outside of the

Montage Noir sequence, prevent accurate assessment and coding of such a character. Shape of the palpebral lobe when viewed dorsally appears to be distinct,

with Astycoryphe and many stratigraphically lower taxa having thinner, crescent-

51 shaped lobes and stratigraphically higher taxa, such as Tropidocoryphe portraying

bean shaped lobes.

13. Eye size: length of eye (B) in relation to length of cephalon (A, Figure

2-6). The division point between character states represents the midpoint

between the two statistically distinguishable generic level populations.

States: (0)<0.30; (1) > 0.3.

Remarks: In association with Feist & Clarkson's (1989) proposed evolution of blindness in later Devonian tropidocoryphids, this character attempts to measure eye reduction compared to cephalon length (A, Figure 2-6). Due to taphonomic factors, primarily crushing, eye height was deemed unreliable for measurement and coding. A significant gap is present between the two character states, therefore statistical support for the distinction of populations is not necessary.

14. Border field sculpture: border sculpture.

States: (0) smooth or no sculpture (e.g., Astycoryphe gracilis, Snajdr 1980, pi. XXIX, fig. 18); (1) fine ridges (e.g., Tropidocoryphe amuri, Chatterton et al.

2006, pi. 41.2 and 41.5); (2) granules (e.g., Erbenicoryphe nazairensis, Feist

52 2003, pl.2, fig. 15); (3) tubercles, in the form of larger, more robust, and less dense granules (e.g., Tropidocoryphe lemniscata, Feist 1976, pi. 2, fig. 2).

Remarks: Sculpture elements are used in almost all proetid diagnoses and have been considered important aspects of classification at many taxonomic levels. Specifically for the examined tropidocoryphid taxa, sculpture has been used as a diagnostic feature for many species (Alberti 1969; Feist 1976; Snajdr

1980; Chatterton et al. 2006). Applicable proetid sculpture types of Snajdr (1980) are used as character states here.

15. Presence of occipital lobe spine: presence of a median posterior spine on

the occipital lobe.

States: (0) present; (1) absent.

Remarks: Presence of a posteriorly projecting spine on the posteromedial portion of the occipital lobe is used for diagnosis at the species level (Feist 1976;

Snajdr 1980. An occipital lobe spine can be highly variable in size, ranging from a small point (e.g., Tropidocoryphe pseudofilicostata, Feist 2003, pi. 1, fig. 1), or in more extreme example, a large spine that is longer than the occipital lobe is wide

(e.g., Pterocoryphe larouquettensis, Feist 2003, pi. 3, fig.l).

53 16. Presence of occipital lobe node: presence of a median occipital node.

States: (0) present; (1) absent.

Remarks'. Similar to the presence of the occipital lobe spine, the presence

of an occipital node has been used as a diagnostic character at the species level for

numerous proetid taxa (Snajdr 1980). However, unlike the occipital spine, the

occipital node is not variable in size, only in presence and absence. Most often it

appears as an enlarged tubercle or granule located along the midline of the

cephalon, typically anterior of the occipital spine, if one is present (e.g.,

Tropidocoryphepseudofilicostata, Feist 2003, pi. 1, fig. 1).

Glabella

17. Shape: relative shape (outline) of the glabella (Figure 2-12).

States: (0) pear-shaped; (1) subcylindrical; (2) tongue-shaped.

Remarks: Snajdr (1980) highlighted the generic level diagnostic characteristics of the proetid glabella. Diagnoses of Tropidocoryphe,

Erbenicoryphe, and Pterocoryphe all take into consideration glabellar shape.

54 18. Furrow shape and depth: depth of the glabellar furrows. Primarily the SI

furrow, such that they are tear-drop shaped and deep enough to make the

glabella appear "nose shaped" in dorsal view.

States: (0) deep SI furrow, making glabella appear "nose shaped" (e.g.,

Tropidocoryphe taharajatensis, pi. 2-5 fig. 2); (1) shallow SI furrow (e.g.,

Astycoryphe ditropidia, pi. 2-1 fig. 1).

Remarks: While the number of glabellar furrows, typically three, is quite conservative at lower proetid taxanomic levels, the shape and depth of these furrows can vary at the genus or species level (Snajdr 1980). Chatterton et al.

(2006) in the description of Tropidocoryphe amuri made reference to the glabella appearing "nose shaped". This is a result of the S1 and S2 furrow configuration, as the SI furrow is deep and tear-dropped shaped, pointing towards the constriction point of the anterior lobe of the glabella. Meanwhile the S2 furrow is shallow and located medial of the glabellar constriction point, posterior-medial of cephalic suture angle y, pointing posteromedially. When both the S1 and S2 furrows are shallow the glabella is globose.

19. Length index: length of the anterior lobe of the glabella (Ai) divided by

the length of the cephalon (A, Figure 2-6).

55 States: (0) < 0.4; (1) 0.4 - 0.5; (2) > 0.5.

Remarks: Length of the anterior lobe of the glabella in reference to the

length of the cephalon can be quite variable; however variation can be significant

at higher taxonomic levels (Snajdr 1980). In the case of tropidocoryphids, length

of the anterior lobe of the glabella compared to cephalon length resulted in

statistically significant populations.

20. Sculpture: sculpture of the glabella.

States: (0) smooth or no sculpture; (1) fine ridges; (2) granules; (3)

tubercles, in the form of larger, more robust, and less dense granules.

Remarks: Sculpture of the glabella is typically the most prominent and

easily identifiable of the entire exoskeleton, making it a useful character to assess

lower taxonomic level relationships (Snajdr 1980). Character states are synonymous with previously outlined sculpture states and those of Snajdr (1980).

Thorax

21. Number of segments: number of thoracic segments.

States: (0)9; (1)10.

56 Remarks: The number of thoracic segments is a highly diagnostic

character, specifically at the genus level and higher (Snajdr 1980). Unfortunately,

given the highly incomplete, partial, and only associated nature of much

previously described tropidocoryphid material, this character cannot be coded for

many taxa. As a result, this character was reported as "parsimoniously

uninformative" by the analysis. Despite this labeling the character was maintained

in the analysis because of its high diagnostic importance and to facilitate future work by providing information about this character for genera for whom it is unknown, primarily genera that are more basal on the tree.

Pygidium

22. Length vs. Width index: maximum length of the pygidium (Z) in relation

to the maximum width of the pygidium (W, Figure 2-6).

States: (0) < 0.55; (1) > 0.55.

Remarks: The size of the pygidium, in tropidocoryphids, is typically associated with the size of the cephalon, however variance does promote this character as having some phylogenetic use (Snajdr 1980). Snajdr (1980) diagnosed Tropidocoryphe as having a pygidium whose length/width ratio is

57 between 0.55 and 0.6, while Tropicoryphe has a pygidial length/width ratio of around 0.5. However, statistical examination of this character, for this analysis, could not differentiate genera (primarily Astycoryphe and Tropidocoryphe) from one another. Instead, this character is useful in distinguishing specimens from different localities (collections) and some species from one another.

23. Shape: overall shape (outline) of the pygidium in accordance to Snajdr

(1980) (Figure 2-13).

States: (0) semi-elliptical; (1) rounded pentagonal.

Remarks: Snajdr (1980) stated that the shape (outline) of the pygidium is characteristic of particular genera, with Tropidocoryphe being diagnosed with a semi-elliptical shaped pygidium. However, pygidial shape appears to display no generic level support, as species and genera display a variety of different shapes.

This character was retained for it is diagnostic value at the species level.

24. Rhachis (axis) shape: shape (outline) of the rhachis (Figure 2-14).

States: (0) cup-shaped; (1) cone-shaped.

58 Remarks: Comparison of the outline of the rhachis is used to compare the degree of tapering throughout its length. A cup shaped rhachis displays a low degree of tapering throughout its length. Inversely, a cone shaped rhachis has a higher degree of tapering throughout its length, causing its sides to be more strongly angled and pointed towards the midline of the rhachis.

25. Rhachis (axis) configuration: number of axial segments the rhachis

(axis) is composed of.

States: (0) 6 + 1; (1) 7 + 1; (2) 8 + 1; (3) 9 + 1.

Remarks'. Throughout trilobite ontogeny, rhachis configuration (number of axial segments + terminal piece) is quite flexible during preholaspid growth stages, with additional segments being generated and released into the axial region. At the holaspid growth stage rhachis configuration becomes stable, with thoracic segments no longer being released into the thoracic region from the pygidium during molting (Chatterton & Speyer 1997).

In terms of taxonomy, rhachis configuration of holaspid stage trilobites is most consistent at lower level taxonomic units (e.g., genus and species), fluctuating in only a very narrow range (Snajdr 1980). At higher level taxonomic units (e.g., family and above) rhachis configuration is less consistent and more variable, with different taxa containing different numbers of axial segments. If

59 research material is focused on holaspid stage specimens, rhachis configuration is an informative diagnostic character at lower level taxonomic units.

26. Rhachis (axis) length vs. width: length of the rhachis (Y) divided by the

maximum width (X, Figure 2-6).

States: (0) < 1.00; (1) 1.01 -1.24; (2) > 1.25.

Remarks: Measurement of the overall dimensions of the rhachis have been cited as being diagnostic of Pterocoryphe (Feist 2003), which displays a long and narrow rhachis. Further examination of the character proved diagnostic for more than just Pterocoryphe. The Tukey Pairwise test discriminated character states 1 and 2 (1.506*10" 4) from one another, however this result should be considered with a side of caution as state 2 only contained three samples (all species of

Pterocoryphe). A sample size of three is in no way statistically representative, however given that Pterocoryphe is diagnosed with a long and narrow rhachis and because the three Pterocoryphe appear to be distinct (presence of a wide gap between Pterocoryphe population and the other populations), we considered state

2 a valid character state (population).

27. Rhachis length index: length of the rhachis (Y) in relation to the entire

length of the pygidium (Z, Figure 2-6).

60 States: (0) < 0.65; (1)> 0.65.

Remarks: Snajdr (1980) indicated that the length of the rhachis is a suitable lower taxonomic level character. Furthermore, Snajdr (1980) diagnosed both Tropidocoryphe and Tropicoryphe using this character.

28. Number of pleural ribs: number of pleural ribs on the pygidium.

States: (0) < 4; (1) 5; (2) 6; (3) > 7.

Remarks: Along with the overall shape of the pygidium, its configuration is also diagnostic at the generic and species levels (Snajdr 1980). As adult specimens no longer release pygidial segments into the thoracic region during molting, the number of pygidial ribs becomes stable, and can be measured with certainty.

29. Number of pleural ribs that reach the margin of the pygidium: number

of pleural ribs that reach the margin of the pygidium, intersecting with the

pygidial margin.

States: {0) 1; (1) 2; (2) > 3.

61 Remarks: In association to the number of pleural ribs present on the pygidium, the number of ribs which cross the border furrow and contact the pygidial border is diagnostic at lower taxonomic levels. Examination of only holaspid material is frequired in order for character accuracy.

30. Pleural rib shape: shape of the three most anterior pleural ribs on the

pygidium (Figure 2-15).

States: (0) curved; (1) bent.

Remarks: Shape of the anterior most pairs of pygidial ribs is diagnostic at the generic level. Due to the nature of the pleural ribs behind the first three to take on a similar appearance in both character states (straight and angled posterolaterally away from the pygidial midline) only the first three ribs are suitable for accurate coding. Pleural rib shape can only be accurately coded using holaspids.

31. Pleural region sculpture: sculpture of the pleural region of the pygidium.

States: (0) smooth or no sculpture; (1) fine ridges; (2) granules; (3) tubercles, in the form of larger, more robust, and less dense granules.

62 Remarks: Character states are synonymous with previously outlined sculpture states and those of Snajdr (1980).

Characters not included in analysis

Of all the characters examined in this analysis, the only character omitted which could cause foreseeable doubt is the preglabellar field length ((A2 + A3) -

A5) in relation to cephalon length (A, Figure 2-6). Despite its use as a diagnostic character for both Tropidocoryphe and Astycoryphe, the character and its states are strongly correlated with character 1, pretropidial and tropidial field length (A2

+ A3, Figure 2-6). For both characters the character states and coding are exactly the same. This is the result of the one character being dependent on the other, with an increase in preglabellar length being the result of an increase in the pretropidial and tropidial length. In order to prevent overweighting of a single morphological change by assigning multiple characters to it, the preglabellar field length was culled.

Results

Heuristic parsimony analysis, with all characters unweighted and unordered, yielded three most parsimonious trees with a total length of 143, ensemble consistency index (C.I.) of 0.3636, ensemble retention index (R.I.) of

0.6364, and rescaled consistency index (R.C.) of 0.2359 (see Figure 2-16, for the heuristic consensus phylogenetic hypothesis; see Figure 2-17, for cladogram

63 indicating character changes). Character 21, the number of thoracic segments, was reported as parsimoniously uninformative; however it was not culled from the matrix given its importance for species diagnoses (Alberti 1969). The only differences in the topology of the three resulting trees is the species level relationships of Tropicoryphe, with each tree considering a different Tropicoryphe species the most primitive of the group. Strict and 50% majority rule consensus of the three trees results in the Tropicoryphe clade becoming a polytomy, but the species within the genus still remaining as a distinct clade, separate from other genera. In all circumstances, Tropicoryphe remains sister to Astycoryphe gracilis, and the rest of the tree's topology remains constant.

Branch-and-bound or exhaustive analyses could not be completed for the entire data set because of the large number of taxa and limited number of characters. Only heuristic analyses proved fruitful on the complete data set. In order to be as conservative as possible, and determine if the results of the heuristic analysis were robust enough to be replicated in the more extensive exhaustive,

Longicoryphe, Erbenicoryphe, and Pterocoryphe were separated from

Astycoryphe, Bojocoryphe, Tropicoryphe, and Tropidocoryphe to generate two exhaustive analysis subsets. Both subsets were then subjected to exhaustive analysis under similar conditions as those of the heuristic analysis of the complete data set. Exhaustive analysis results of the Astycoryphe, Bojocoryphe,

Tropicoryphe, and Tropidocoryphe subset produced over 500 trees, with only

Astycoryphe and Tropidocoryphe being supported. Both Bojocoryphe and

64 Tropicoryphe were grouped within the Astycoryphe clade and not well supported as distinct throughout the majority of trees. As for the Longicoryphe,

Erbenicoryphe, and Pterocoryphe subset, exhaustive analyses resulted in the same general conclusions as those of the heuristic analysis of the complete dataset, with

Longicoryphe being the most primitive and Pterocoryphe the most derived. The fundamental topology of the subset analyses was either identical to the heuristic analysis of the complete data set or totally unsupported beyond the generic level, therefore the results of the exhaustive subset analyses were only used to confirm that the results of the original heuristic analysis are conservative and viable.

Discussion and conclusions

Despite only three very similar trees being produced by heuristic analyses, the low value consistency indices and weak Bremer support values for most clades in all three trees prevent any of them, including their consensus, from being considered a strict phylogenetic hypothesis. This is further substantiated by the weakly supported species level relationships present in the exhaustive analyses of the Astycoryphe, Bojocoryphe, Tropicoryphe, and Tropidocoryphe subset. Slight differences in character choice and coding, in both the heuristic and exhaustive analyses, can result in major topological changes taking place throughout the cladogram. As a result, future work will play a major role in understanding tropidocoryphids and their phylogeny better. Until then, only extremely

65 conservative cladistic revisions of the examined tropidocoryphids and their relationships should be attempted using this phylogeny.

In order to be as conservative as possible, stable groupings were identified as clades with a Bremer support value greater than one. This approach aims to accept groups with the highest support values, while suppressing those that are less supported and likely a result of over splitting. As such, both Tropidocotyphe and Pterocoryphe are monophyletic, while Astycoryphe and Longicoryphe are paraphyletic. Astycoryphe is the most primitive tropidocoryphid examined, as it is the stratigraphically lowest taxon and has the most primitive character suit. Given this, Longicoryphe, Erbenicoryphe, Pterocoryphe, and Tropidocoryphe all represent more derived tropidocoryphids. Such a relationship highlights the expansion of the pretropidial area, anterolateral expansion of facial suture angles, and increased pygidial length (Z, Figure 2-6) within the Astycoryphe-

Longicoryphe- Tropidocoryphe and Astycoryphe-Longicoryphe-Pterocoryphe lineages.

The validity of Bojocoryphe and Tropicoryphe as genera is also called into question by this analysis, with both genera not being distinguishable from

Astycoryphe during exhaustive analyses and only weakly supported as distinct from Astycoryphe and one another during the heuristic analysis. Not surprisingly, as the diagnoses of Bojocoryphe and Tropicoryphe are near identical to

Astycoryphe. Furthermore, Tropicoryphe hamalaghdadica and T. marrocanica were originally considered species of Tropidocoryphe and Astycoryphe (Alberti

66 1969), before being designated to Tropicoryphe (Snajdr 1980). Considering these factors and that Tropicoryphe and Bojocoryphe are not easily distinguishable from

Astycoryphe, both Tropicoryphe and Bojocoryphe are considered junior synonyms and their species assigned to Astycoryphe.

Interestingly the Longicoryphe, Erbenicoryphe, and Pterocoryphe relationship appears nearly identical to that proposed by Feist & Clarkson (1989).

The only minor difference in our phylogeny, from that of Feist & Clarkson (1989), is that Longicoryphe is considered ancestral to Tropidocoryphe, instead of vice versa. Given that Longicoryphe is geologically younger than Tropidocoryphe, it is possible that L. brilonensis and L. circumincisa represent reversals, explaining their placement between Astycoryphe and the more derived Longicoryphe and

Tropidocoryphe clade. However, in order to refine such a claim, more work needs to be done, specifically phylogenetic analysis on a more tightly constrained stratigraphic sequence with well documented and complete specimens, allowing reversals to be recognized more easily.

In terms of synapomorphies within the examined taxa, the bulk of characters experience either reversals higher up in the phylogeny or homoplastic occurrences outside of a clade (see Figure 2-16). As a result, very few unique synapomorphies are present in this phylogeny, with only bean shaped palpebral lobes being synapomorphic to Tropidocoryphe and a widely positioned cephalic suture angle P being synapomorphic to Pterocoryphe. Furthermore, numerous autapomorphies are also present, as many species contain unique characteristics

67 not present in other taxa. In an attempt to solve these problems, and strengthen the branch support with additional synapomorphies, future research needs to expand upon the completeness of taxa included in the analysis to reduce the number of unknown character states.

Despite the lack of synapomorphies and the presence of numerous homoplasies in the phylogeny, two evolutionary trends are present. First, the localized Longicoryphe-Erbenicoryphe-Pterocoryphe lineage of the Montagne

Noir sequence demonstrates reduced eye size in comparison to cephalon length, with more derived taxa having smaller eyes compared to more primitive taxa.

This evolutionary trend of eye reduction is identical to the previously documented, early stages, of the tropidocoryphid evolutionary blindness trend proposed by Feist & Clarkson (1989), and supports previous conclusions that

Longicoryphe is ancestral to Erbenicoryphe and Pterocoryphe (Feist 1976, 2003).

Both Tropidocoryphe and Astycoryphe however, do not show any signs of eye reduction when examined across the palaeogeographically larger Devonian Rheic

Ocean region. Therefore, given that eye reduction is localized to taxa of the

Montagne Noir sequence, and that the primitive and derived genera, Astycoryphe and Tropidocoryphe, do not display a similar evolutionary trend, blindness appears to be a localized phenomenon of the Montagne Noir tropidocoryphids and not present in other palaeogeographical regions.

The second, and more palaeogeographically widespread, evolutionary trend observed within the phylogeny of the examined tropidocoryphids is the

68 gradual expansion of the pretropidial and tropidial fields. Primitive taxa, such as

Astycoryphe, display relatively small pretropidial and tropidial fields, with

upturned ('cushion-like') cephalic margins, while more derived taxa,

Pterocoryphe and Tropidocoryphe, have much larger pretropidial and tropidial

fields with flat cephalic margins. In conjunction with expansion of the pretropidial

and tropidial fields, there is also a shift of cephalic suture angle P to a more lateral

position. This shift in best observed in Pterocoryphe, when compared to the more

primitive condition observed in Astycoryphe, P has shifted significantly laterally

(away from the cephalic midline). Although not as significant as Pterocoryphe, the position of landmark P in Tropidocoryphe is also more lateral (away from the cephalic midline) than in the more primitive Astycoryphe.

The lack of phylogenetic support and misinterpretation of the relationships of Tropidocoryphe, Astycoryphe, and Longicoryphe in previous studies are the result of two issues: primarily, many of the described species within these genera are based on partial, fragmented, or incomplete specimens, causing many of the species level relationships to be poorly supported and secondly, the similarity of the diagnoses of some taxa, prevents assignment of species to the proper genus.

Both of these factors are a function of two issues: work on taxa occurring in isolation of other research on similar topics, and a lack of rigorous testing and support of conclusions. Use of rigorously assessed quantitative characters and cladistic analysis reduces these shortcoming, as shown by the proposed phylogeny herein, and should be employed whenever possible. Without a unified approach

69 toward examining these taxa and their relationships, understanding their evolution

and the evolution of higher taxa encompassing them (e.g., Proetida) will not be

easily accomplished and documented accurately.

Systematic palaeontology

Order Proetida Fortey & Owens, 1975

Subfamily Tropidocoryphinae Pribyl, 1946

Genus Astycoryphe Richter & Richter, 1919

Synonymized Genera Tropicoryphe Snajdr, 1977; Bojocoryphe Snajdr, 1976;

Gracilocoryphe Pribyl & Vanek ,1978

T^pe species - Astycoryphe senckenbergiana Richter & Richter, 1919.

Other included species - Astycoryphe gracilis (Barrande, 1846); Astycoryphe memnon (Hawle & Corda, 1847); Astycoryphe Junius (Billings, 1869);

Astycoryphe chctmpernowni (Whidborne, 1889); Astycoryphe westfalica (Richter

& Richter, 1919); Astycoryphe mauretanica Alberti, 1967; Astycoryphe hamlaghdadica (Alberti, 1967); Astycoryphe cimelia Ormiston, 1967;

Astycoryphe arcticus Ormiston, 1967; Astycoryphe marrocanica Alberti, 1969;

Astycoryphe splendens (Snajdr, 1976); Astycoryphe hermon (Snajdr, 1977);

Astycoryphe vermon (Snajdr, 1977); Astycoryphe astyx Snajdr, 1980; Astycoryphe larminiei Wright & Chatterton, 1988; Astycoryphe minuta (Ellerman, 1992);

70 Astycoryphe austriaca Ellerrrtan, 1992; Astycoryphe repens (Frantisek & Vanek,

2000) Astycoryphe planifrons Feist, 2003; Astycoryophe jurosi van Viersen &

Prescher, 2010; Astycoryphe exilis van Viersen & Prescher, 2010; Astycoryphe ditropidia sp. nov.

Diagnosis - Tropidocoryphid with narrow preglabellar field; slightly divergent

anterior portions of cephalic sutures; inflated or "cushion-like" cephalic border

with border rims; tropidium; short pygidium with cup shaped rhachis; and curved

pleural ribs on pygidium.

Remarks - Description and diagnosis of Longicoryphe by Erben (1966), via the designation of A. novaki as its type species, caused the original diagnosis of

Astycoryphe (Richter & Richter 1919) to become problematic. Harrington (1959) listed the diagnostic features of Astycoryphe as being: narrow preglabellar field, slightly divergent anterior portions of the cephalic sutures, presence of border rims about the cephalon, and a short pygidium with pronounced pleural ribs. Feist

& Clarkson (1989) agreed with these diagnostic features, and discussed them in relation to their proposed Late Devonian tropidocoryphid evolutionary sequence.

In light of this study's quantitative and statistical character examination and cladistic analysis, additional diagnostic features of Astycoryphe include: inflated or "cushion-like" cephalic border, tropidium, cup-shaped pygidial rhachis, and curved pleural ribs on the pygidium.

Based on the similarity of diagnoses of Astycoryphe, Tropicoryphe, and

Bojocoryphe, and the unsupported differentiation of the later two genera from

71 Astycoryphe in the cladistic analyses, we consider both Bojocoryphe and

Tropicoryphe junior synonyms of Astycoryphe, This designation increases the

Pragian diversity of Astycoryphe.

Given the current confusing designation of the holotype of Astycoryphe gracilis, as a species of Astycoryphe (Feist 1976, 2003; Snajdr 1980) and as the holotype of Gracilocoryphe (Pribyl & Vanek 1978; Frantisek & Vanek 2000) at the same time, we consider A gracilis a valid member of Astycoryphe. Therefore,

Gracilocoryphe is considered a junior synonym of Astycoryphe and until further study of it's other species, Gracilocoryphe repens Frantisek & Vanek, 2000, can be achieved they should also be included as species of Astycoryphe.

Overall Astycoryphe is a valid genus, differing from Longicoryphe by possessing a narrow preglabellar field, narrow placement of the cephalic suture angle P, shallow glabellar furrows, and a cup-shaped pygidial rhachis.

Astycoryphe ditropidia new species

Plate 2-1, figs. 1-6

Derivation of name - Named in reference to the species' two distinct tropidial ridges. '£)/' being the greek prefix for two and 'tropidia' being the morphological terminology for the ridges encompassing the anterior portion of the genal field.

Diagnosis - Short pretropidial area (less than 0.15x cephalon length (A)); moderately divergent P-P cephalic suture distance; two distinct, well developed,

72 parallel tropidial ridges encompassing part of genal field; cephalic border

sculpture of fine terrace ridges; third most anterior thoracic segment bearing

macropleural spine with border rims on medial flank; long and narrow pygidial

rachis ((Y) greater than 1.25x width (X)); and curved pygidial pleural ribs.

Material - Holotype UA13667 (complete articulated exoskeleton).

Occurrence - Jbel Ofatene Philonyx/Quadrops horizon, west of Lahfira,

southeastern Morocco. Strata are Eifelian (Middle Devonian) in age and part of

the El Otfal Formation of the Ma'dar basin (Morzadec 2001; Gibb & Chatterton

2010).

Description - Cephalon is "D" shaped and moderately convex. Sagittal length (A,

Figure 2-6) 0.4 lx length of entire specimen. Anterior border cushion-like in shape

and adorned by single fine ridge (border rim). Posterior border and medial flank

of genal spine rimmed by numerous fine parallel ridges. Anterior border furrow

narrow and shallow, widening as it approaches genal spine. Tropidium composed

of two distinctive ridges, running parallel to anterior and lateral border furrow.

Tropidial field length (distance between tropidial ridges) approximately 0.04x length of cephalon (A, Figure 2-6). Pretropidial field length (distal most tropidial ridge to anterior limit of cephalon) and posttropidial field length (As) are 0.13x and 0.14x respectively, compared to cephalon length (A, Figure 2-6). Distal tropidial ridge shorter than proximal ridge, terminating lateral of facial suture angle 5 on free cheek. Proximal tropidial ridge larger and more robust than distal ridge, tapering to point lateral of cephalic suture angle ©, where anterior and

73 posterior cephalic border furrows converge. Region proximal of tropidium

displays sparse fine granulose surface texture. Posterior border furrow shallow

and runs laterally from axial furrow, lateral of occipital ring, to junction with

anterior/lateral border furrow of genal spine.

Cranidium. Preglabellar area length (A2 + A3) 0.3 lx cephalon length (A,

Figure 2-6) and convex. Glabellar length (A4 + Ai) approximately 0.70x length of

cephalon (A) (without LO (Ai, Figure 2-6), it is 0.5 lx). Anterior portion of

cephalic suture constricted, with p-(3 distance of 0.79x cephalon length (Figure

2-6, length A). Glabella is subcylindrical in shape, weakly convex and has fine

terrace ridge sculpture throughout. Posteromedian portion of glabella, between

LI, slightly inflated. SI furrows shallow, teardrop-shaped, with broader and

deeper regions near anterior of ovoid LI. S2 furrows much smaller, shallower, to

nearly nonexistent. S2 furrows occur medial to point of glabellar constriction and

angled posteromedially at approximately 40°. Palpebral lobes crescent-shaped and

sloped inwards towards moderately deep axial furrow. Occipital furrow (SO)

moderately deep, shallowing laterally, away from midline. Occipital ring (A4)

0.19x sagittal length of cephalon (A, Figure 2-6) and transversely wider than

widest portion of glabella. Fine granulose sculpture throughout occipital ring.

Lateral occipital ring furrows shallow with median and lateral lobes weakly

convex.

Free Cheek. Eyes are large and strongly convex in both horizontal and lateral planes. Length of eye (B) approximately 0.4lx length of cephalon (A,,

74 Figure 2-6). Long genal spine extends to seventh thoracic axial ring. Medial

aspect of genal spine inflated and outlined with multiple robust parallel ridges.

Lateral border furrow of genal spine is wide and shallow.

Hypostome unknown.

Thorax consists of 9 segments, and represents 0.43x sagittal length of

entire specimen. Widest segment is 0.92x maximum width of specimen. Axial

rings are moderately arched and accentuated by moderately deep axial furrows.

Maximum width of axis is 0.35x maximum width of thorax. Axial rings decrease

in width from anterior to posterior, with most posterior (9th) ring being 0.7 lx

width of most anterior (1st) ring. All axial rings display sculpture of fine terrace ridging throughout. Pleurae incised with moderately shallow pleural furrow running along midlength of segment. Pleural furrows deepest medially, shallowing towards fulcrum. Inner regions of pleurae are transverse, while more distal regions curve posteriorly within approximately 72° of exsagittal line.

Pleurae have fine terrace ridge sculpture which is most prominent distal of fulcrum. Elongate macropleural spines on third thoracic segment. Medial margin of macropleural spines lined with fine parallel border rims, similar to medial aspect of genal spines.

Pygidium 0.16x total length (Z) and 0.60x maximum width (W, Figure

2-6) of the complete specimen. Pygidium rounded-pentagon shaped. Proximal portion of anterior margin of pygidial pleural region is transverse; and distal to fulcrum, margin angles posterolaterally at approximately 46° from transverse.

75 Moderately deep border furrow parallels lateral and posterior margins. Rhachis

(pygidial axis) convex and cone shaped, tapering from anterior to posterior.

Rhachis length (Y) and width (X) are 0.76x and 0.38x, compared to corresponding dimensions of pygidium (Z and W, Figure 2-6). Rhachis comprised of 7 rings and terminal piece. Rhachial rings decrease in width from anterior to posterior, with more anterior rings possessing more pronounced protuberance on posteromedial margin. Anterior ring furrows deeper than more posterior ring furrows. Five pairs of pleural ribs and six interpleural furrows present. Pleural rib structure is imbricate. Interpleural furrows are moderately deep, shallowing distally as they curve posteriorly towards pygidial margin. Ribs decrease in size and convexity from anterior to posterior. Two most anterior rib pairs contact pygidial margin, bisecting pygidial border furrow. More posterior ribs taper out proximal to border furrow. Post axial ridge is weak. Pygidial border furrow shallow, with narrow pygidial border. Rhachis and pleural region surrounding rhachis, lack sculpture. Pleural border furrow and margin with sculpture of fine terrace ridges.

Remarks - Astycoryphe ditropidia is extremely rare at Jbel Ofatene, as only a single specimen is known. Lack of posteromedial occipital ring portion, due to damage, limits identification of median nodes or spines on the specimen.

Surprisingly, Astycoryphe ditropidia is not most similar to other Moroccan representatives of Astycoryphe, but rather is most similar to A.jorusi (van Viersen

& Prescher 2010, pi. 8, fig. 1-10) from Belgium. A. ditropidia is similar to A.

76 jorusi based on narrow positioning of cephalic suture angle 5 and a large anterior lobe of the glabella in relation to cephalon length (A, Figure 2-6). A. ditropida differs qualitatively from A. jorusi by having a tropidium composed of two distinct and robust tropidial ridges (A. jorusi has only one), cephalic border field sculpture of fine terrace ridges, prominent macropleural spine on third thoracic segment with border rims on its medial aspect, and one less pair of pleural ribs that reach the border of the pygidium. Quantitative differences between A ditropidia and A. jorusi are A. ditropidia having a more laterally positioned (away from midline) cephalic suture angle (3, a shorter pretropidial length and tropidial length ((A2 + A3) - As) compared to the length of the cephalon (A), and a longer

(Y) and more narrow (X, Figure 2-6) rhachis.

Astycoryphe ditropidia also bears a slight resemblance to the type species of Astycoryphe, A. senckenbergiana (van Viersen & Prescher 2010, text-fig. 7), by having a strongly upturned ('cushion-like') cephalic border, crescent-shaped palpebral lobes, a subcylindrical shaped glabella, and a rounded-pentagonal shaped pygidium. A. ditropidia can be distinguished from A. senckenbergiana in possessing nine thoracic segments rather then 10 and a rhachis with a configuration of 7+1 instead of 6+1. Both these differences are significant in the fact that they are most likely ontogenetically related (Chatterton & Speyer 1997), with A. senckenbergiana undergoing one additional molt during its meraspid life- cycle stage (adding an additional segment into its thoracic region from its pygidium) before beginning its holaspid (adult) stage. If this is indeed true, A.

77 ditropidia, and the more derived tropidocoryphids, could have evolved through paedomorphosis of A. senckenbergiana.

Lastly, Astycoryphe ditropidia resembles A. gracilis in some features

(Snajdr 1980, pi. XXIX fig. 14-22). Both species have very prominent and distinct border rims, primarily on the medial aspect of the genal spine, a tropidium composed of more than one tropidial ridge, and a short and wide rounded- pentagonal shaped pygidium. However, presence of more than 3 tropidial ridges in the tropidium and the robust genal caeca of A, gracilis distinguishes it from A. ditropidia. Furthermore, A. ditropidia is distinct from A. gracilis by having a shorter pretropidial and tropidial field length compared to cephalon length, cephalic border field sculpture of fine terrace ridges, and a more moderately sized glabella compared to cephalon length .

Genus Tropidocoryphe Novak, 1890

Type species - Tropidocoryphe filicostata Novak, 1890.

Other included species - Tropidocoryphe latens (Barrande, 1846);

Tropidocoryphe ascanius (Barrande, 1852); Tropidocoryphe heteroclyta

(Barrande, 1852); Tropidocoryphe barroisi Mailleiux, 1904; Tropidocoryphe pseudofilicostata Pribyl, 1965; Tropidocoryphe consobrina Alberti, 1967;

Tropidocoryphe richterorum Cisne, 1968; Tropidocoryphe erbeni Morzadec,

1969; Tropidocoryphe mediterranea Feist, 1976; Tropidocoryphe undulans Feist,

78 1976; Tropidocoryphe lemniscata Feist, 1976; Tropidocoryphe etemira Snajdr,

1980; Tropidocoryphe seewartensis Ellerman, 1992; Tropidocoryphe endorfensis

Basse, 1997; Tropidocoryphe amuri Chatterton et al., 2006; Tropidocoryphe

bassei van Viersen et al., 2009; Tropidocoryphe taharajatensis sp. nov.;

Tropidocoryphe lahfirensis sp. nov.; Tropidocoryphe sculptaridgiata sp. nov.

Diagnosis - See Kowalski 1990.

Remarks - Tropidocoryphe represents the most diverse tropidocoryphid genus, and is distinct in comparison to both Astycoryphe and Longicoryphe. It can be differentiated from Astycoryphe by the possession of a long preglabellar field, flat and/or smooth cephalic border, anterolateral placement of the cephalic suture angle (3, presence of robust genal caeca that bifurcates at the tropidium anterior of the glabella, deep SI glabellar furrows, and long (Z, Figure 2-6) pygidium with cone shaped rhachis. Tropidocryphe is also distinguishable from Longicoryphe by the presence of a large preglabellar field in relation to the glabella, cup shaped palpebral lobes, large eyes, and a cup shaped rhachis. The only synapomorphic character for this genus, based on the phylogentic analysis of this study, is bean shaped palpebral lobe (character 12 = 0).

Tropidocoryphe amuri Chatterton et al., 2006

Plate 2-2, figs. 1-6; plate 2-3, figs. 1-6; plate 2-4, figs. 1-7.

Diagnosis - See Chatterton et al. 2006

79 Material - Holotype UA13394 (complete articulated exoskeleton) and paratypes

(UA13667, UA13668, UA13669, and UA13670).

Occurrence - Found at both Zguilma and Assa of the Tindouf basin. At Zguilma

Tropidocoryphe amuri is found in two of the three trilobite producing layers

(ZGEE1 & ZGEE3), which are Late Emsian and Eifelian in age (Chaterton et al.

2006; Gibb & Chatterton 2010). At Assa, T. amuri is found in the late Emsian

aged horizon, Assa 2.

Remarks - Tropidocoryphe amuri was originally described by Chatterton et al.

(2006) from the upper Emsian aged bed ZGEE1 at the base of section in the

Timrhanrhart Formation near Foum Zguid. New material has since been discovered at bed ZGEE3, which is approximately 48 meters higher in section than bed ZGEE1 and Eifelian in age (Chatterton et al. 2006; Gibb & Chatterton

2010). Recovery of T. amuri from both upper Emsian and Eifelian beds suggests that the stratigraphic range of the species is quite broad, longer than any other

Moroccan tropidocoryphid. This new material is identical to that of the holotype of T, amuri (UA13394), with no discernible differences in size, shape or configuration of elements.

Tropidocoryphe taharajatensis new species

Plate 2-5, figs. 1-7 & plate 2-6, figs. 1-6.

80 Derivation of name - Named in reference to the locality of Taharajat, the primary locality in southeastern Morocco where the species is found.

Diagnosis - Pygidium of rounded pentagonal shape; extremely large pretropidial field length (greater than 0.32x total cephalon length (A); short anterior lobe of glabella (less than 0.40x length of entire cephalon (A, Figure 2-6)); pygidial rhachis configuration of 8 segments and 1 terminal piece.

Material - Holotype UA13673 (complete articulated exoskeleton) and paratypes

(UA13674 &UA13675).

Occurrence - From two trilobite producing horizons at Taharajat, Tazoulai't

Formation, Ma'der Basin, south central Morocco, late Emsian based on trilobite and conodont faunas (Gibb & Chatterton 2010).

Description - Cephalon half-moon shaped and weakly convex. Sagittal length (A,

Figure 2-6) is 0.40x length of entire body. Anterior border slightly upturned.

Anterior border furrow broad and concave dorsally, shallowing as it approaches genal spine. Tropidium composed of single distinctive ridge surrounding anterior and lateral regions of genae. Pretropidial field length ((A2 + A3) - As) and posttropidial field length (A5) 0.34x and 0.13x cephalon length (A, Figure 2-6), respectively. Tropidium tapers to point lateral of facial suture angle to, where lateral and posterior border furrows converge. Regions proximal and distal of tropidium display dense fine granulose surface texture. Posterior border furrow shallow, running laterally from occipital ring axial furrow to junction with lateral border furrow of genal spine.

81 Cranidium. Preglabellar area (A2 + A3) 0.47x length of cephalon (A,

Figure 2-6). Anterior portions of cephalic sutures widely spaced, with P-(3 distance of 0.93x cephalon length (A, Figure 2-6). Two robust genal caeca (caecal ridges) project forward from anterolateral corners of glabella to anterior border of cephalon, bifurcating at tropidium. Length of glabella (Ai + A4) approximately

0.54x length of cephalon (A), 0.37x without occipital lobe (LO) (Ai, Figure 2-6).

Glabella pear-shaped, flared posteriorly, with fine granulose sculpture throughout.

Posteromedian portion of glabella (between LI lobes) inflated and convex, looking "nose-like" when viewed dorsally. S1 furrows deep and teardrop shaped.

S2 furrows smaller and shallower, crescent-shaped and angled at approximately

68° to midline. Palpebral lobes bean (kidney) shaped, sloping medially towards axial furrow. Occipital furrow and lateral occipital ring furrows moderately deep, giving occipital ring sigmoidal or w-shaped appearance. Occipital ring length (A4)

0.08x length of cephalon (A, Figure 2-6) and transversely wider than widest portion of anterior glabellar lobe. Fine granulose sculpture throughout occipital ring.

Free Cheek. Eyes small (B), 0.2 lx length of cephalon (A, Figure 2-6), and kidney shaped. Eyes strongly convex in lateral and horizontal planes. Long, angular genal spine extends to eighth thoracic axial ring. Anterior and lateral border furrow broad, shallowing posteriorly. Posterior border furrow narrow and deep, shallowing and widening laterally before intersecting lateral border furrow.

Hypostome unkown.

82 Thorax consists of 9 segments, representing 0.34x sagittal length and

0.80x maximum width of entire specimen. Axial rings moderately arched dorsally and accentuated by fairly deep axial furrows. Spinose median tubercles present on all axial segments, increasing in size posteriorly. Maximum width of axis (tr.) about 0.27x maximum width of thorax. Axial rings increase in width (tr.) posteriorly until 6th thoracic segment. Backwards of 6th thoracic segment axial rings decrease in width (tr.) until pygidium. All axial rings display sculpture of fine granules. Pleurae incised with deep pleural furrows running along segment midlength (tr.). Furrows deepen laterally towards fulcrum, before shallowing and disappearing after fulcrum. Medial portion of pleurae are traverse, while distal portions curving posteriorly at approximately 65° from sagittal line. Sculpture of fine granules present throughout pleurae, decreasing in density away from midline.

Pygidium. Length (Z) and width (W, Figure 2-6) 0.30x and 0.59x total length (sag.) and width (tr.) of entire specimen. Pygidium rounded-pentagonal shape. Medial portion of anterior pleural margin is transverse; distal to fulcrum, margin angles posterolaterally at approximately 54° from transverse. Broad shallow border furrow. Rhachis convex and cone shaped, tapering from anterior to posterior and outlined by moderately deep axial furrows. Rhachis length (Y) and width (X) are 0.53x and 0.25x corresponding dimensions of entire pygidium (Z and W, Figure 2-6). Rhachis configuration of 8 rings and 1 terminal piece.

Rhachial rings marked by prominent axial ring furrows. Anterior rhachial ring

83 furrows distinctly deeper than more posterior furrows. Posterior median tubercles of rhachial rings decrease in size from anterior to posterior, 8th ring lacks node.

Seven pairs of pleural ribs and eight pairs of interpleural furrows. Pleural rib structure imbricate. Most anterior pleural ribs follow shape of thoracic segment's pleural portions. More posterior ribs become increasingly posteriorly directed.

Interpleural furrows moderately deep, shallowing as they approach the pygidial border. Ribs decrease in size and convexity from anterior to posterior. Two most anterior rib pairs contact the pygidial margin, bisecting the pygidial border furrow.

More posterior ribs taper out medial of border furrow. Post-axial ridge weak.

Sculpture of fine granules adorns entire pygidium.

Remarks - Tropidocoryphe taharajatensis is a rare species at Taharajat, with only three specimens known. Chatterton et al. (2006), when describing T. amuri, noted possession of a Taharajat specimen, however considered it T. amuri at the time.

Discovery of two additional specimens from Taharajat, along with examination of gradational characters and phylogenetic analysis of this study, supports the exclusion of these specimens from T. amuri as a new species.

Tropidocoryphe taharajatensis also shows resemblance to the type species of Tropidocoryphe, T. filicostata, which Alberti (1969, pi. 30 fig. 11-13) documented from the northern region of Morocco. Both species portray a tropidium comprised of a single ridge and similarly sized anterior glabellar lobe and glabellar furrows. Furthermore their pygidial shape and pleural rib configuration is identical, along with exoskeletal sculpture. Differences between

84 the species include: divergence of the anterior portion of the cephalic suture, pear- shaped instead of tongue-shaped glabella, and pygidial rhachis configuration of

8+1 instead of 9+1.

Resemblance between Tropidocoryphe taharajatensis, T. pseudofilicostata pseudofilicostata (Snajdr 1980, pi. XVIII fig. 12-17) and T. lemniscata is also quite high. Similarities between the species consist of position of cephalic suture points, shape and configuration of glabella, and the number of pleural ribs on the pygidium. T. taharajatensis differs from T. pseudofilicostata pseudofilicostata in having granulose glabellar sculpture and a pygidial rhachis configuration of 8+1 instead of 9+1. T. taharajatensis differs from T. lemniscata in only having one tropidial ridge, a larger preglabellar field, granulose cephalic border sculpture, and a rounded-pentagonal shaped pygidium.

Similarity between T. taharajatensis and other Tropidocoryphe species is less than that of previously discussed species. T. bassei (van Veirsen et al. 2009, pi. 1 figs. 9-10) lacks granulose sculpture over its entire exoskeleton. T. latens

(Snajdr 1980, pi. XVIII fig. 7-11) having glabellar sculpture of terrace ridges instead of granules, a semi-elliptically shaped pygidium with one less pair of pleural ribs, and a shorter and narrower rhachis with one less rhachial ring.

Compared to T. undulans (Feist 1976, pi. 1 fig. 15-16), T. taharajatensis differs in having a narrower pygidium, with one less pair of pleural ribs, and granulose sculpture throughout the entire exoskeleton.

85 Tropidocoryphe lahfirensis new species

Plate 2-7, figs. 1-7 & plate 2-8, figs. 1-6.

Derivation of name - Named in reference to the village of Lahfira, Morocco.

Located approximately 8 km southeast of the section at Jbel Oufatene, Lahfira is

the closest village to the location from which this species was originally found.

Diagnosis - Tropidocoryphe with short anterior lobe of glabella ((Ai) is less than

0.40x length of entire cephalon (A)); pygidium of rounded pentagonal shape; long

and narrow pygidial rhachis ((Z) is greater than 1.25x width (W, Figure 2-6)); and

pygidial rhachis with configuration of 8 segments and terminal piece.

Material - Holotype UA13676 (complete articulated exoskeleton), and paratypes

(UA 13677, UA13678, UA13679).

Occurrence - From the Philyonix/Quadrops horizon at Jbel Oufatene, El Otfal

Formation, Eifelian, Ma'der basin, southeastern Morocco (Morzadec 2001; Gibb

& Chatterton 2010).

Description - Cephalon. Weakly convex and crescent-shaped. Sagittal length (A,

Figure 2-6) is 0.36x length of entire body. Anterior border slightly upturned.

Cephalic border furrow slightly concave and broad, shallowing towards genal spine. Tropidium composed of single distinctive ridge surrounding anterior and lateral regions of genal fields. Pretropidial field length ((A2 + A3) - A5) and posttropidial field length (As) 0.34x and 0.13x cephalon length (A, Figure 2-6), respectively. Tropidial ridge tapers to point lateral of facial suture angle cd, where

86 lateral and posterior border furrows converge. Dense fine granulose sculpture present throughout regions to both sides of tropidium. Posterior border furrow moderately deep, running transversely from occipital ring axial furrow to junction with lateral border furrow near genal spine.

Cranidium. Preglabellar length (A2 + A3) 0.45x length cephalon (A, Figure

2-6). Anterior portions of cephalic sutures transversely expanded, with P-(3 distance of 0.97x cephalon length (A, Figure 2-6). Two robust genal caeca (caecal ridges) project forward from anterolateral corners of glabella to anterior border of cephalon, bifurcating at tropidium. Glabella (Ai + A4) approximately 0.48x length of cephalon (A), and without occipital lobe (LO) (Ai, Figure 2-6) it is 0.36x.

Glabella pear-shaped and flared posteriorly, with sculpture of fine granules throughout. Posteromedial portion of glabella (between LI lobes) inflated and convex, causing glabella to appear "nose-like" viewed dorsally. S1 furrows deep teardrop shaped with anterolateral portion being deepest. S2 furrows small, shallow, crescent-shaped, and angled at approximately 50° to midline. Palpebral lobes bean shaped, with lateral aspect elevated and sloping downwards towards axial furrow. Occipital furrow and lateral occipital ring furrows moderately deep, causing occipital ring (LO) to appear sigmoidal or w-shaped. Occipital ring (LO) length (A4) is 0.12x length of cephalon (A, Figure 2-6) and transversely wider than widest portion of glabella farther forward. Occipital ring with fine granulose sculpture throughout. Posteromedian occipital spine present. Median occipital node absent.

87 Free Cheek. Eyes (B) small, 0.23x length of cephalon (A, Figure 2-6) and

kidney shaped. Eye lens field is strongly convex in both lateral and horizontal

planes. Long, angular genal spine projects posteriorly to eighth thoracic axial ring.

Posterior border furrow narrow and deep, shallowing and widening distally until

intersecting lateral border furrow.

Hypostome unknown.

Thorax consists of 9 segments, representing 0.3 lx sagittal length and

0.92x transverse width of entire specimen. Axial rings moderately arched and

accentuated by moderately deep axial furrows. Spinose median tubercles present

on all axial segments, increasing in size posteriorly. Maximum width (tr.) of axis

approximately 0.26x maximum width of thorax. Axial rings increase in width (tr.)

posteriorly from 1st to 6th segment. Posterior of 6th thoracic segment, axial rings

decrease in width (tr.) backwards towards pygidium. All axial rings display sculpture of fine granules. Pleurae incised with deep pleural furrows running

along midlength (sag.). Furrows deepest at fulcrum, shallowing medially and

laterally. Medial regions of pleurae are transverse, with more distal regions curving posteriorly within approximately 55° of specimen's sagittal midline.

Pleural sculpture consists of fine granules, with granule density decreasing laterally.

Pygidium. Length (Z) and width (W, Figure 2-6) are 0.32x and 0.67x total length (sag.) and width (tr.) of entire specimen. Pygidium rounded-pentagon shape. Medial portion of anterior pygidial pleural margin transverse; distal to

88 fulcrum, margin angles posterolaterally at approximately 62° from transverse

direction. Broad, shallow furrow proximally borders lateral and posterior margins.

Rhachis strongly convex and cone shaped, tapering in width from anterior to

posterior. Rhachis outlined by moderately deep axial furrows on either side.

Rhachis length (Y) and width (X) 0.58x and 0.26x corresponding dimensions of

entire pygidium (Z and W, Figure 2-6). Rhachis configuration of 8 rings and

terminal piece. Rhachial rings divided by prominent axial ring furrows. More

anterior rhachial ring furrows are distinctly deeper than more posterior furrows.

Posterior median tubercles decrease in size from anterior to posterior, with 8th ring

tubercle nonexistent. Seven pairs of pleural ribs and eight pairs of interpleural

furrows. Pleural rib structure is imbricate. Most anterior pleural ribs follow shape

of thoracic pleural regions. More posterior ribs become increasingly posteriorly

directed. Interpleural furrows moderately deep, shallowing as they approach pygidial border. Pleural ribs decrease in size and convexity from anterior to

posterior. Two most anterior rib pairs contact pygidial margin, bisecting pygidial

border furrow. More posterior ribs taper out proximal to border furrow. Post axial

ridge is weak. Sculpture of fine granules throughout pygidium.

Remarks - Like all tropidocoryphids, Tropidocoryphe lahfirensis is rare.

However, it is not as rare as the other species from Jbel Oufatene, T. sculptaridgiata and Astycoryphe ditropidia. Multiple specimens of T. lahfirensis are present in our collection while there is only single example of T. sculptaridgiata and A. ditropidia.

89 Tropidocoryphe lahfirensis most resembles the type species of

Tropidocoryphe, T. filicostata, which Alberti (1969, pi. 30 fig. 11-13) documented

from the northern region of Morocco. Both species have similar sized and

dimensioned preglabellar areas, tropidia, glabellar size and furrow depth, pygidial shape and pleural rib configuration, and sculpture. T. lahfirensis differs from T. filicostata though in having more divergent anterior portions of the cephalic suture, a pear-shaped instead of tongue-shaped glabella, and pygidial rhachis configuration of 8+1 instead of 9+1.

Comparison of Tropidocoryphe lahfirensis to T. pseudofilicostata

pseudofilicostata shows multiple similarities and differences. Similarities consist of the position of cephalic suture points, composition of tropidium, glabellar shape and dimension, and pygidial shape and sculpture. Differences are a narrower preglabellar field, granulose instead of terrace ridge sculpture of the glabella, a longer and narrower rhachis with one less rhachial ring, and one less pair of pygidial pleural ribs in T. lahfirensis.

Compared to other Tropidocoryphe, T. lahfirensis is not very similar. The

German T. bassei (van Veirsen et al. 2009, pi. 1 fig. 9-10) is the only other

Tropidocryphe with which T. lahfirensis could possibly be confused. T. lahfirensis is distinguished by its smaller preglabellar field, a rounded-pentagonal shaped pygidium with longer and narrower rhachis, and granulose sculpture throughout, character stats that T. bassei lacks.

90 In comparison to other Moroccan Tropidocoryphe, T. lahfirensis differs from T. amuri in having a weaker and less robust tropidial ridge, granulose sculpture on the cephalic border, and pygidial pleural regions, anterior-posterior

compression of the posterior section of cephalic suture, and has a longer and

narrower rhachis. Most similar to T. lahfirensis is T. taharajatensis, as both share a number of features, however T. lahfirensis is easily distinguishable by its shorter preglabellar area and one less pair of pygidial pleural ribs (6 instead of 7).

Tropidocoryphe sculptaridgiata new species

Plate 2-9, figs. 1-6.

Derivation of name - Named in reference to the species entire exoskeleton being covered in a sculpture of fine distinct terrace ridging.'Sculpta' referring to the sculpture and 'ridgiata' referring to the terrace ridge sculpture found throughout its exoskeleton.

Diagnosis - Large tropidial field, located between anterior limit of glabella and cephalic border furrow, and composed of multiple short distinct tropidial ridges; prominent sculpture of fine terrace ridges throughout entirety of exoskeleton; long and narrow pygidial rhachis ((Y) greater than 1,25x width (W, Figure 2-6).

Material - Holotype UA13672 (incomplete articulated exoskeleton).

91 Occurrence - Diademaporetus horizon, Jbel Oufatene, El Otfal Formation,

Ma'der basin, southeastern Morocco. The horizon is the uppermost of four

trilobite producing horizons, located approximately 15 meters above the Philony/

Quadrops horizon. Horizon is Eifelian in age (Gibb & Chatterton 2010).

Description - Cephalon. Relatively incomplete. Tropidium composed of multiple

ridges (more than 3) ahead of glabella. Most anterior tropidial ridge robust, with

more posterior ridges becoming less robust towards glabella. Border region with

sculpture of fine terrace ridges. Region behind tropidium displays very sparse and

fine granulose surface texture ahead of glabella. Palpebral lobes with fine terrace

ridge sculpture throughout.

Cranidium. Glabella pear-shaped, flared posteriorly with fine terrace ridge

sculpture throughout. Posteromedian portion of glabella (between LI lobes)

inflated and convex, looking "nose like" viewed dorsally. SI furrows incised and

teardrop shaped, with deepest portion lateral of midline. S2 furrows small,

shallow, crescent-shaped, and angled at approximately 68° from sagittal line.

Palpebral lobes bean shaped, sloping inwards towards axial furrow. Occipital

furrow moderately deep. Occipital ring transversely wider than widest portion of

anterior lobe of glabella. Lateral occipital ring furrows shallow, with median

portion of occipital lobe inflated anteriorly. Sculpture of fine terrace ridging adorns entire occipital ring. Median spine base present at posterior extant of occipital ring.

Free Cheek. Unknown.

92 Hypostome. Unknown.

Thorax. Consists of 9 segments. Axial rings moderately arched and flanked by moderately deep axial furrows. Spinose median tubercle present on axial rings of two most posterior segments. Maximum width of axis (tr.) is approximately 0.36x maximum width of thorax (tr.). Axial rings increase in width

(tr.) posteriorly until 6th segment, posterior of 6th thoracic segment, axial rings decrease in width (tr.). All axial rings display sculpture of fine terrace ridging.

Inner regions of pleurae are transverse, with more distal regions curving posteriorly within 70° of sagittal line, beyond fulcrum. Pleurae incised with moderately deep pleural furrows running along midlength (sag.) of segment.

Pleural furrows deepest at mid-length of inner (tr.) pleural regions, shallowing medially and laterally towards axial rings and fulcrum. Sculpture of fine terrace ridging present throughout pleurae, with ridges decrease in robustness laterally.

Pygidium. Length (Z) and width (W, Figure 2-6) 0.72x and 0.84x maximum length (sag.) and width (tr.) of thoracic region. Pygidium rounded- pentagonal shape. Proximal portion of anterior pleural margin transverse; distal to fulcrum, margin angles posterolaterally at approximately 66° from transverse direction. Broad and shallow furrow borders lateral and posterior margins.

Rhachis convex and cone shaped, with width (tr.) tapering from anterior to posterior. Rhachis outlined by moderately deep axial furrows on either side.

Rhachis length (Y) and width (X) 0.6lx and 0.29x compared to corresponding dimensions of entire pygidium (Z and W, Figure 2-6). Rhachis configuration of 8

93 rings and terminal piece. Rhachial rings separated by axial furrows, with anterior

ring furrows distinctly deeper than more posterior furrows. Posterior median spine of rhachial rings decreases in size from anterior to posterior. Seven pairs of pleural ribs and eight pairs of interpleural furrows. Pleural rib structure is imbricate. Most anterior pleural ribs follow shape of thoracic pleural region. More posterior ribs become increasingly posteriorly directed (parallel to midline). Interpleural furrow moderately deep, shallowing as they approach pygidial border. Ribs decrease in size and convexity from anterior to posterior. Two most anterior rib pairs bisect the pygidial border furrow and contact pygidial margin. More posterior ribs taper out proximal of border furrow. Weak post axial ridge present. Sculpture of fine terrace ridging throughout pygidium.

Remarks - This species is extremely rare at Jbel Oufatene, with only one partial specimen being known. Despite the incomplete cephalon, we consider the presence of multiple tropidial ridges within the tropidium as a diagnostic character of the species, not a developmental pathology of the specimen, as its tropidium's appearance is very similar to that of T. lemniscata and other tropidocoryphids with multiridged tropidia. Furthermore, diagnosis of this specimen as a new species is also supported by its entire exoskeleton exhibiting only terrace ridge sculpture, no granulose sculpture, unlike all other currently described species of

Tropidocoryphe.

According to the phylogenic analysis, Tropidocoryphe sculptaridgiata and

T. mediterranea are most closely related. Similarities between the two include

94 glabellar length (Ai, Figure 2-6), pygidial shape, rhachis configuration, and sculpture. T. sculptaridgiata differs from T. mediterranea in having a tropidium comprised of multiple ridges (not just one), a longer and narrower pygidium, and only having 6 pairs of pleural ribs.

Tropidocoryphe sculptaridgiata also appears similar to T. lemniscata (Feist

1976, pl.2 figs. 1-4), both possessing a tropidium composed of multiple (more than

3) tropidial ridges, an occipital lobe node and spine, a pear-shaped glabella with deep S1 furrows, pygidial rhachis with configuration of 8+1, and bent shaped pygidial ribs. T. sculptaridgiata differs from T. lemniscata by displaying fine terrace ridge sculpture throughout its entire exoskeleton, rounded-pentagonal shaped pygidium, and having only 2 (not 3) pleural ribs that reach the pygidial border.

Compared to the Moroccan material of Tropidocoryphe filicostata (Alberti

1969, pi. 30 fig. 11-13), T. sculptaridgiata is distinguishable by having a tropidium composed of more than 3 ridges, a pear-shaped glabella, one less axial ring (8+1 instead of 9+1), and the entire exoskeleton having fine terrace ridged sculpture. Compared to the slightly different T. pseudofilicostata pseudofilicostata

(Snajdr 1980, pi. XVIII, fig. 12-17), T. sculptaridgiata differs by having a longer and narrower pygidium that is rounded-pentagonal in shape and only 6 pairs of pygidial ribs.

van Veirsen et al. (2009, pi. 1 fig. 9-10) figured and described

Tropidocoryphe bassei from Germany. T. sculptaridgiata differs from T. bassei, in

95 having a larger tropidial field made of multiple tropidial ridges, a rounded- pentagonal shaped pygidium instead of a semi-elliptical shaped pygidium, a longer and narrower pygidial rhachis, and dominated sculpture of fine terrace ridging. Compared to other non Moroccan based Tropidocoryphe (T. undulans, T. latens, and T. lemniscata), T. sculptaridgiata differs by having a multi-ridged tropidium and fine terrace ridged sculpture in many regions.

In relation to the new Moroccan species described herein, Tropidocoryphe sculptaridgiata most closely resembles T, amuri by having a similar shaped glabella, number of thoracic segments, and pygidial shape and dimensions.

However, T. sculptaridgiata differs from T. amuri in having multiple ridges composing its tropidium, a longer and narrower rhachis, and sculpture of fine terrace ridges instead of granules on multiple regions. Compared to new species T. taharajatensis and T. lahfirensis, T. sculptaridgiata differs again by its multi- ridged tropidium and sculpture of fine terrace ridges instead of granules in many regions. Furthermore, T. sculptaridgiata differs in having a longer and narrower rhachis and one less pair of pygidial ribs than T. taharajatensis.

96 AFRICA

SPAIN

• Rz

MOROCCO

Erfoud D - Ma'der basin, Figure 2-4

ALGERIA C - Tindouf Basin, Figure 2-3

Figure 2-1. Locality maps of Africa and Morocco. A: Map of Africa depicting Morocco (shaded area) (Modified from Philip (1991); Gibb (2005); Gibb & Chatterton (2007,2010); McKellar & Chatterton (2009); Chatterton & Gibb (2010)). B: Map of Morocco indicating locations of basins surveyed in this study (modified from Gibb & Chatterton (2007,2010); McKellar & Chatterton (2009); Chatterton & Gibb (2010)). C and D: Location of individual study basins within Morocco (refer to Figures. 2-3 and 2-4 for more detailed maps of these areas)

97 Pteroparia coumiacensis rC Pteroparia ocu/ata Pterocoryphe progrediens Pterocoryphe languedociana Longicoryphe bissousensis Longicoryphe britonensis Longicoryphe circumincisa Erbencoryphe parvula Tropidocoryphe Astycoryphe

Figure 2-2. Redrafted version of Feist & Clarkson's (1989) proposed phylogeny of Middle to Late Devonian tropidocoryphid trilobites from the Montagne Noir sequence of France.

98 Foum Zguid

Mdaouer Srbir

Zguilma

Ordovician Hercynian Dolerite Devonian (Upper Palaeozoic)

Figure 2-3. Map of the Tindouf Basin, southern Morocco. The town of Foum Zguid is located to the north, with the Zgulima section located in the Devonian outcrop to the south southeast (indicated by arrow on map). (Diagram adapted from Fetah et al. (1998), Gibb (2005), Chatterton et al. (2006), Gibb & Chatterton (2007,2010), and McKellar & Chatterton (2009)).

99 Tazoulait

'Lahfira

hwtio«(OOC|

Figure 2-4. Map of the Ma'der Basin, southern Morocco. Boxes indicate section localities. (Diagram adapted from Fetah et al. (1998), Gibb (2005), Chatterton et al. (2006), Gibb & Chatterton (2007,2010) McKellar & Chatterton (2009), and Chatterton & Gibb (2010)).

100 Figure 2-5. Stratigraphic columns of Zguilma (adapted from Gibb (2005), Chatterton et al. (2006), Gibb Chatterton (2007,2010) and McKellar & Chatterton (2009)), Assa, and the western slope of Jbel Oufatene (adapted from Gibb (2005), Gibb & Chatterton (2007, 2010), and McKellar & Chatterton (2009)). Measurements are in meters and formations are noted to the right of columns. Mined horizons are indicated indicated by arrows and their local names.

101 Tindouf Basin Legend Tropdocofypt* Assa U*®l nnauhi e rnnuti PA: Tft&docoryptM 9 icufaisridgufi ItopUocoryph* AWYUNMON. J J JM. C tahfimnsb Attyco/ypt* MirtMddad UnMkm* ^.~~J SttyShtfa ® d*n/x**

100 — Ma'der Basin Oufatdne <— Brschiopod Lag 350_ s

ELOTFAL 3QQ_SS=? FORMATION 0 horizon «— Diadamaproetus horizon Zguilma .— Gerastos/Phacops horizon >— Phitonyx/Quadrops horizon $*e*on«sn*WM 100_ DO TI TAZOULAJT FORMATION M R H , Equivalent to Psychopyge ..'XT1 J A horizon N R H 200 ER REMLIA FORMATION?? A N T

Base of BOUTISKAOUINE "--a Jbel Zguilma FORMATION R - OclontochHe horizon s

50- ffffl -ZGEE3 F © I 0 H R A M F N A 0 D T R A 1 M R O A N TI c-zk-zl O N "i.t.'S.t.'i - Kolihapeltls horizon ZGEE2 -2-— -ZGEE1 e • Dtcranums horizon

102 w Figure 2-6. Diagram showing the points of measurement on the cephalon and pygidium, as per Owens (1973). Ratio data were obtained by comparing two measurements, most commonly one measurement in reference to cephalon length (A, sag.). Coordinate data were collected by comparing the position of points (e.g., landmark p) in relation to the anterior and posterior limits of the cephalon length measurement (A, sag.).

103 Figure 2-7. Cephalic border shape. Arrow indicates area of referenced used to examine this character. A: inflated ('cushion-like') B: flat

Figure 2-8. Presence of border rims. A: border rims are not present (absent) B: border rims are present

104 Position of Cephalic Suture Angle Beta 0

Legend + Population 1 a Population 2 o Population 3

Figure 2-9. Position of cephalic suture angle beta p. Distances Yp and Xp were divided by the length of the cephaion (A, sag., Figure 2-6). Xp and Yp coordinates from all species were then graphed and generic level populations mathematically differentiated using a Hoteling's t-tests with a Bonferroni correction. Three populations were found to be present for the P cephalic suture position.

105 Position of Cephalic Suture Angle Gamma (y) —r ~x'r • • - .* ~.r

Legend a Population 1 +• Population 2

on »a «j» oj ojs Xy

Figure 2-10. Position of cephalic suture angle gamma (y). Distances Yy and Xy were divided by the length of the cephalon (A, sag., Figure 2-6). Xy and Yy coordinates from all species were then graphed and generic level populations were mathematically differentiated using a Hoteling's t-test. Two generic level populations were found to be present for the y cephalic suture position.

106 Position of Cephalic Suture Position Delta (5) .""f .$ *! "'"I . f I ! . i • •!- i r * •.

r«- * i''

Legend d Population 1 + Population 2

om av as e« Xo

Figure 2-11. Position of cephalic suture angle delta (8). Distances Ya and X5 were divided by the length of the cephalon (A, sag., Figure 2-6). Xs and Ys coordinates from all species were then graphed and generic level populations mathematically differentiated using a Hoteling's t-test. Two populations were found to be present at generic levels.

107 Figure 2-12. Shape of glabella, as defined by Snadjr (1980,). A: pear-shaped B: subcylindrical C: tongue-shaped

Figure 2-13. Pygidial shape as defined by Snadjr (1980). A: semi-elliptical B; rounded pentagonal

108 Figure 2-14. Rhachis (axis) shape. A: cone shaped B: cup shaped

Figure 2-15. Shape of pygidial pleural ribs. A: bent B: straight

109 -•Bojocorypha aplendena -•Astycoryphe gracilis -•TFoplcoryphe nramnon -•Troplcoryphe hamlaghdadlca -•Tropleoryphe marrocanica

-•Aatyeoiyphe ditropidla i*.

(!) -•Astycoryphe Jorual •Astycoryphe senckenbarglana Longlcorypha clrcuminclaa Longlcorypha brllonanala Longlcorypha tanulatrlata Longlcorypha lanceolata Longlcorypha antaglabra Erbenlcoryphe nazalrenala

Pterocoryphe languadoclana

Pterocoryphe larouquatlanala Tropidocoryphe undulans •Tropldocorypha latana Tropidocoryphe lemnlscata Tropidocoryphe paeudoflllcoaata

Tropidocoryphe taharajatensls n.«. Tropidocoryphe flllcoatata Tr*« stattaflcc Tropidocoryphe lahflranala n «> L«n0fi"143 C.l.*0.»36 fl-l. »0.S3M Tropidocoryphe amuri R.C. • 0.2399 Tropidocoryphe baaael Tropidocoryphe medlterranea Tropidocoryphe aculptarldglata*

Figure 2-16. Cladogram depicting consensus phylogenetic hypothesis based on the 3 most parsimonious trees produced during Heuristic search with no character weighting or ordering. {#} Denotes Bremer support number for clade, all unmarked clades have value of {1}.

110 Bojocoryphe splendens Astycoryphe gracilis 14(0] 2311] 2IW)2*11 •Tropicoryphe memnon 41015(11 1710120(21 •Tropicoryphe hamlaghdadica 1(2)3(2] 7I0]1«(0] •Tropicoryphe marrocanica •Astycoryphe ditropldia 1(0] 0(1] 9(2] t4f1]2«(2J3

1(2) 7(0} 23(0) 27(11 Longlcoryphe lanceolata 4(0] K1) 14(0] 20(0) 31(01 10)2K» Longlcoryphe anteglabra 23T 29(3] Erbenicoryphe nazalrensls ?111 •(1124(1) 25(0] 30(11 14(2i 15(01 •Pterocoryphe languedoclana 3(2111(01 6(2) 23(0] 2«2 2*3] Pterocoryphe larouquettensls 14(0] 18(0] 20|0] 22(1]29(2)31(01 •Tropldocoryphe undulans 20(3} 29(3] 31|3] Tropldocoryphe latens 4(0) S(1| 11(0}12(0) 23(0)24(1) Tropldocoryphe lemniscata « 14(3 29(11 •Tropldocoryphe pseudoflllcosata 20|1)22(01 25(3) Tropldocoryphe taharajatensls Tropldocoryphe flllcostata 6(0117f2] 25)31 Tropldocoryphe lahflrensls Tropldocoryphe amuri 8(311«0] •Tropldocoryphe basse! 1 PI 23(0) Tropldocoryphe medlterranea 22|0| 2«P| 26(2) Tropldocoryphe sculptaridgiata Figure 2-17. Expanded view of cladogram (Figure 2-16),displaying changes in character states. Cladogram is a re-drafted version of the output from MacCiade (Maddison & Maddison 2001) and its data matrix was initially analyzed using PAUP (Swofford 2002). Numbers preceding brackets denote characters, numbers within brackets denote change in character state. All changes shown are unambiguous. (#) = unique, uniform above [#] = homoplasy outside = homoplasy above {#} = homoplasy above and outside

111 Plate 2-1

Fig. 1-6. Astycoryphe ditropidia n. sp. from the Philonyx/Quadrops horizon, El Otfal Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA13667) (complete specimen). 1, dorsal view of cephalon and thorax; X7.4. 2, dorsolateral view; X6.7. 3, anterodorsal view; X8.2. 4, dorsal view of pygidium; X8.2. 5, lateral view of genal spine and macropleural thoracic spine; XI0.7. 6, lateral view; XI.6.

112 113 Plate 2-2

Fig. 1-5. Tropidocoryphe amuri Chatterton et al., 2006 from ZGEE3, Timrhanrhart Formation, Zguilma, Tindouf Basin, southern Morocco (UA13668) (complete specimen). 1, dorsal view of cephalon and thorax; X4.1. 2, lateral view of cephalon; X4.0. 3, anterolateral view of cephalon; X4.1. 4, dorsal view of thorax and pygidium; X4.7. 5, anterior view of cephalon; X4.6. Fig. 6. Tropidocoryphe amuri Chatterton et al., 2006 from Assa 2, Khebchia Formation, Assa, Tindouf Basin, southern Morocco (UA13667) (complete specimen). 6, dorsal view; X3.3.

114 115 Plate 2-3

Fig. 1,3-6. Tropidocoryphe amuri Chatterton et al., 2006 from ZGEE3, Timrhanrhart Formation, Zguilma, Tindouf Basin, southern Morocco (UA13394) (complete specimen). 1, dorsal view of cephalon and thorax; X3.5.3, anterolateral view of cephalon; X3.8. 4, dorsal view of thorax and pygidium; X3.6. 5, anterior view of cephalon; X3.7. 6. lateral view; X3.0. Fig. 2. Tropidocoryphe amuri Chatterton et al., 2006 from ZGEE1, Timrhanrhart Formation, Zguilma, Tindouf Basin, southern Morocco (UA13670) (partial specimen). 2, dorsal view; X2.8.

116 117 Plate 2-4

Fig. 1-2,4,6. Tropidocoryphe amuri Chatterton et al., 2006 from ZGEE3, Timrhanrhart Formation, Zguilma, Tindouf Basin, southern Morocco (UA13669) (complete specimen). 1, dorsal view of cephalon and thorax; X3.1. 2, lateral view; X3.6. 4, dorsal view of pygidium; X3.2. 6, anterolateral view of cephalon; X3.7. Fig. 3,5,7. Tropidocoryphe amuri Chatterton et al., 2006 from ZGEE1, Timrhanrhart Formation, Zguilma, Tindouf Basin, southern Morocco (UA13670) (partial specimen). 3, anterolateral view of cephalon; X2.7. 5, anterior view of cephalon; X2.9. 7, dorsal view of pygidium; X4.7.

118 119 Plate 2-5

Fig. 1-7. Tropidocoryphe taharajatensis n. sp. Tazoulai't Formation, Taharajat, Ma'der Basin, southern Morocco (UA13673) (complete specimen). 1, dorsal view; X2.2. 2, dorsal view of cranidium; X3.6. 3, anterolateral view of cephalon; X2.1. 4, dorsal view of cephalon; X2.2. 5, anterior view of cephalon; X2.4. 6, dorsal view of pygidium; X3.1. 7, lateral view; X2.5.

120 121 Plate 2-6

Fig. 1-2,4-5. Tropidocoryphe taharajatensis n. sp. Tazoulai't Formation, Taharajat, Ma'der Basin, southern Morocco (UA13674) (complete specimen). 1, dorsal view; X3.5. 2, anterior view of cephalon; X3.5. 4, dorsal view of pygidium; X4.3. 5, lateral view; X2.5. Fig. 3 & 6. Tropidocoryphe taharajatensis n. sp. Tazoulai't Formation, Taharajat, Ma'der Basin, southern Morocco (UA13675) (partial specimen). 3, dorsal view; X3.0. 6, anterior view of cephalon; X2.4.

122 123 Plate 2-7

Fig. 1-3, 5, 6. Tropidocoryphe lahfirensis n. sp. from the Philonyx/Quadrops horizon, El Otfal Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA13676) (complete specimen). 1, dorsal view; X2.5. 2, anterior view of cephalon; X2.6.3, anterolateral view of cephalon; X2.3. 5, dorsal view of pygidium; X3.4. 6, lateral view; X2.1. Fig. 4 & 7. Tropidocoryphe lahfirensis n. sp. from the Philonyx/Quadrops horizon, El Otfal Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA13678) (complete specimen). 4, dorsal view of cephalon and thorax; X3.3. 7, dorsal view of pygidium; X3.2.

124 125 Plate 2-8

Fig. 1,4, 5. Tropidocoryphe lahfirensis n. sp. from the Philonyx/Quadrops horizon, El Otfal Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA13679) (complete specimen). 1, dorsal view; X3.6. 4, anterior view of cephalon; X3.5. 5, lateral view; X2.8. Fig. 2,3,6. Tropidocoryphe lahfirensis n. sp. from the Philonyx/Quadrops horizon, El Otfal Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA13677) (complete specimen). 2, dorsal view of cephalon and thorax; X3.2. 3, dorsal view of thorax and pygidium; X2.9. 6, dorsolateral view of cephalon; X3.5.

126 127 Plate 2-9

Fig. 1-6. Tropidocoryphe sculptaridgiata n. sp. from the Diademaproetus horizon, El Otfal Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA13672) (partial specimen). 1, dorsal view; X4.9. 2, dorsal view of tropidium; X8.3. 3, dorsal view of thorax and pygidium; X5.5. 4, dorsolateral view; X4.1. 5, lateral view; X3.4. 6, posterolateral view; X4.9.

128 129 Table 2-1. Table outlining included and omitted taxa for the phylogenetic analysis. Images used in coding of specimens are indicated as the source, while material indicates the completeness of specimens being coded. Reasoning provides explanation as to why the taxon was either included or excluded from the analysis. All remaining species not outlined in the table were omitted either based on lack of material (less than 50% of characters could be coded), poor documentation, or not being located in or around the Rheic Ocean of the Devonian.

130 Bbjocdzyphe splendens Snajdr 1980, pi. - 1 complete exoskeleton Included - type species of Bojocoryphe Snajdbr 1980 XXVn, figs. (articulated) - diagnosis of Bojocoryphe is nearly 15-19. - 5 cranidia, 6 pygidia identical to that of Astycoryphe (associated) Tropicoryphe memnori memnon (Hawle Snajdr 1980, pi. - 5 partial cranidia and 1 Included - type species of Tropicoryphe &C6iidal847) XIX, figs. 1-4. pygidium (associated) Tropicoryphe marrodanica Alberti 1969, pi. - complete cranidium Included - originally diagnosed as Astycoryphe? (Albert! 1969) 30, figs. 5. (Alberti 1967) however redesignated to Tropicoryphe by Snajdr (1980). Tropicoryphe hamalaghdadica (Alberti Alberti 1969, pi. - 3 cranidia and 2 pygidia Included - originally diagnosed as Tropicoryphe 1967) 30, figs. 6-8. (associated) (Alberti 1967) however redesignated to Tropicoryphe by Snajdr (1980). Astycoryphe senckenbergiana Richter & van Veirsen & - 1 complete exoskeleton Included - type species of Astycoryphe Riehter 1919 Prescher 2010, fig. (articulated) CO 7. Astycoryphe jorusi van Veirsen & - 1 cranidium, 3 librigena, and Included - relatively new species with similar van Vierten and Reseller 2010 Prescher 2010, pi. 1 pygidium (associated) morphology to Astycoryphe ditropidia 8, figs. 1-13. Astycoryphe gracilis Snajdr 1980, pi. - 2 complete cephalia, thoracic Included - only Pragian aged Astycoryphe (Banande 1846) XXIX, figs. 14-22. segments, and 26 pygidia • morphologically similar to Tropicoryphe (associated) Longicoryphe circumincisa Feist 1976, pi. 2, - 5 cranidia and 4 pygidia Included - included to test the proposed figs. 9-17. (associated) tropidocoryphid relationships of Feist & Feist 1976 Clarkson (1989). Longicorypke brilonensis (Richter and Feist 2003, pi. 2, - 1 cranidium and 1 pygidium Included - included to test the proposed Richter 1919) figs. 11-12. (associated) tropidocoryphid relationships of Feist & Clarkson (1989). Longjcoryphe tenuistriata Feist 2003, pi. 2, - 4 cranidia, 1 librigena, and 2 Included - included to test the proposed figs. 5-6. pygidia (associated) tropidocoryphid relationships of Feist & Feist 2003 Clarkson (1989). Feist 2003, pi. 2, -1 cranidium and 2 pygidia Included - included to test the proposed figs. 1-4. (associated) tropidocoryphid relationships of Feist & Feist ^03 Clarkson (1989).

Longwqryphe anteglabra Feist 2003, pi. 2, - 4 cranidia, 1 librigena, and 2 Included - included to test the proposed Feist 200^ . \ figs. 7-10. pygidia (associated) tropidocoryphid relationships of Feist & Clarkson (1989). ErbenicoryphC nctzairensis Feist 2003, pi. 2, - 4 cranidia, 1 librigena, and 2 Included - included to test Longicoryphe and Feist2003 figs. 13-15. pygidia (associated) Pterocoryphe relationship proposed by Feist & Clarkson (1989) Pterocoryphe Umguedociana Feist 1976, pi. 3, - 5 cranidia and 5 pygidia Included - type species of Pterocoryphe YaklW V ' figs. 1-7 (associated) - included to test more derived , .. * . . .\-i\ tropidocoryphid relationships. Pterocoryphe larouquettensis Feist 2003, pi. 3, - 1 cranidium and 2 pygidia Included - included to test Longicoryphe, Feist \9f6- , • ' figs. 1-2. (associated) Erbenicoryphe, Pterocoryphe relationship proposed by Feist & Clarkson (1989) Tropidotdifptie undulans Feist 1976, pi. 1, - 1 cranidium and 1 pygidium Included - included to test the proposed Feist 1976 \ . ' . figs. 15-16. (associated) tropidocoryphid relationships of Feist & Clarkson (1989). Tropidocoryphe latens Snajdr 1980, pi. - 7 partial cranidia and 8 Included - morphologically similar to (Barrandel846) XVin, figs. 7-11. partial pygidial (associated) Tropidocoryphe filicostata Tropidocoryphe lemniscata Feist 1976, pi. 1, - 1 cranidium, 1 librigena, and Included • morphologically similar to Feist 1976 figs. 1-4. 1 pygidium (associated) Tropidocoryphe sculptaridgiata n. sp. - included to test tropidocoryphid Tropidocoryphe pseudqfilicostata Snajdr 1980, pi. - 6 partial cranidia and 18 Included • morphologically similar to pseudofilicostata Pribyl 1965 XVm,figs. 12-17. partial pygidial (associated) Tropidocoryphe filicostata Tropidocoryphe filicostata Alberti 1969, pi. - partial cephalon and partial Included - type species of Tropidocoryphe (Novak 1890) 30, figs. 11-13. pygidium - documented in Morocco (Alberti 1969) Trop(docoryphe cariuri Chatterton et al. - 3 complete exoskeletons Included - complete Moroccan specimen (100% Cbd&itm eta!. 2006 2006, pi. 41, figs. (articulated) coded) with broad stratigraphic range 1-6. - 1 pygidium (associated) Thopidocoryphe bassei van Viersen et al. - 2 complete exoskeletons Included • complete specimen (100% coded) van.Vj^rsen et at. 2009 2009. pi. 1, figs. (articulated) 9-10.

Tropidocoryphe mediterranea Feist 1976, pi. 1, -1 cranidium, 2 librigena, and Included - included to test the proposed Feist 1976 figs. 1,5, 11. 1 pygidium tropidocoryphid relationships of Feist & Clarkson (1989). Decoroproetus decorus Snajdr 1980, pi. - 11 complete articulated Omitted • non-suitable outgroup for the examined PfibyH946 XXI, figs. 1-7. specimens tropidocoryphids based on pygidial shape and formation (Snajdr 1980). Longicoryphe novaki Richter & Richter - line drawing of cephalon and Omitted • only line drawings available, not suitable (Beyert869) 1919, figs. 4-6. pygidium for coding. Erbenicoryphe parvula Feist & Clarkson - 2 partial cranidia and 1 Omitted - not enough suitable material for coding Feist&CTarkson 1989 1989, fig. 3 (A-D) partial librigena (associated) (less than 50% coded) Astycoryphe exilis van Veirsen & -1 partial cranidium and Omitted - not enough suitable material for coding van Viersen and Prescher 2010 Prescher 2010, pi. librigena (less than 50% coded) 8, figs. 1-13. Table 2-2. Results of character state statistical tests on a character by character, state by state basis. Use of Mann-Whittney U, ANOVA, and Hoteling's t-test were dependent on data type. All tests were measured against a 95% confidence interval. When two or more character states are present, a Bonferroni correction adjusted p-value was used to test at the 95% confidence interval. Grey shaded cells represent results which should be used with caution as some of their populations (character states) do not have large enough sample sizes to be statistically accurate.

134 ~ Ot, r ^ s*»W33®sRj£i ,• 'j, S 9^w»& jfeg**/ 1 4 ANOVA State 0 (< 0.15) vs. State 1 (0.16 - 0.25) vs. State 2 (0.26 - 0.31) vs. 2.554*10-'° State 3 (>0.32) Tukey test State 0 (< 0.15) vs. State 1 (0.16 - 0.25) 6.839*10"3 State 1 (0.16 - 0.25) vs. State 2 (0.26 - 0.31) 1.689*10-* State 1 (0.26 - 0.31) vs. State 3 (> 0.32) 2.282* 10"2

2 2 Mann-Whittney U State 0 (< 0.05) vs. State 1 (> 0.05) 0.253* 10"2 3 3 ANOVA State 0 (< 0.84) vs. State 1 (0.85 - 1.05) vs. State 2 (> 0.32) 2.424*10-1C Tukey test State 0 (< 0.84) vs. State 1 (0.85 - 1.05) 1.212*10"* State 1 (0.26 - 0.31) vs. State 2 (> 0.32) 1.703*10-3 6 3 Hotelling's T-test State 0 (Narrow Morph) vs. State 1 (Moderate Morph) 2.506* 10"6 (Bonferroni corrected p-value = (95.65%} 0.0167 for 95%) State 1 (Moderate Morph) vs. State 2 (Wide Morph) 9.778* 10"* (100%) 7 2 Hotelling's T-test State 0 (Anterior Morph) vs. State 1 (Posterior Morph) 6.512* 10"4 (91.3%) 8 2 Hotelling's T-test State 0 (Narrow Morph) vs. State 1 (Wide Morph) 1.337*10"s (91.67%) 19 3 ANOVA State 0 (< 0.4) vs. State 1 (0.4 - 0.5) vs. State 2 (> 0.5)) 5.44*10"14 Tukey test State 0 (< 0.4) vs. State 1 (0.4 - 0.5) 1.212*10^ State 1 (0.4 - 0.5) vs. State 2 (> 0.5) 2.181*10"2 26 3 ANOVA State 0 (< 1.00) vs. State 1 (1.01 -1.24) vs. State 2 (> 1.25) 1.732*10!? Tukey test State 0 (< 1.00) vs. State 1 (1.01 -1.24) 3.321*10"2 State 1 (1.01 -1.24) vs. State 2 (> 1.25) 1,'..'- 27 2 Mann-Whittney U State 0 (< 0.65) vs. State 1 (> 0.65) 2.359* 10"2 Literature cited

Alberti, G. K. 1967. Neue obersilurische sowie unter- und mittledevonische Trilobiten aus Marokko, Deutschland und einigen anderen europaischen Gebieten. I and II. Senckenbergiana lethaea, 48, 463-479, 481-509. Alberti, G. K. 1969. Trilobiten des jiingeren Siluriums sowie des Unter- und Mitteldevons. I. Mit Beitragen zur Silur-Devon-Stratigraphie einiger Gebiete Marokkos und Oberfrankens. Abhandlungen der Senckenbergischen Naturforschenden Gesselschaft, 510, 1-692. Alberti, G. K. 1970. Trilobiten des jiingeren Siluriums sowie des Unter- und Mittledevons. II. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 252, 1-233. Anstey, R. L. & Pachut, J. F. 2004. Cladistic and phenetic recognition of species in the Bryozoan genus Peronopora. Journal of Paleontology, 78(4), 651-674. Barrande, J. 1846. Nouveaux Trilobites. Supplement a la notice preliminaire sur le Systeme Silurien et les trilobites de Boheme, I-IV, 1-40. Barrande, J. 1852. Systeme Silurien du centre de la Boheme. 1 ere Partie. Recherches paleontologiques, IrTrilobites, 935 pp. Basse, B. M. 1997. Trilobiten aus mittlerem Devon des Rhenohercynikums: II. Proetida (2), Ptychopariida, (1). Palaeontographica Abt. A, 246 (3-6), 53-142. Basse, B. M. 2010. Proetoidea Hawle & Corda, 1847 und andere Trilobita aus unterdevonischen Herzynkalken der westlichen Harzgeroder Faltenzone (Zlichovium, Dalejum; sudwestlicher Harz, Rhenoherzynikum): Cornuproetinae Richter, Richter & Struve in Moore, 1959 und Eremiproetinae Alberti, 1967b (1). Freiberger Forschungshefte, 18, 1-67. Becker, T.R., Bockwinkel, J., Ebbighausen, V., Aboussalam, S.Z., El Hassani, A. & Niibel, H. 2004a. Lower and Middle Devonian stratigraphy and faunas at Bou Tserflne near Assa (Dra Valley, SW Morocco). Pp. 90-100 in A. El Hassani (ed) Devonian Neritic-Pelagic Correlation and Events in the Dra Valley (Western Anti-Atlas, Morocco). Subcommission on Devonian Stratigraphy. International Meeting on Stratigraphy, Rabat, 1-10 March 2004. Documents de I'Institut Scientifique, 19. Becker, T.R., Jansen, U., Plodowski, G., Schindler, E., Abzoussalam, S.Z. & Weddige, K. 2004b. Devonian litho- and biostratigraphy of the Dra Valley area — An overview. Pp. 3-18 in A. El Hassani (ed) Devonian Neritic-Pelagic Correlation and Events in the Dra Valley (Western Anti-Atlas, Morocco). Subcommission on Devonian Stratigraphy. International Meeting on Stratigraphy, Rabat, 1-10 March 2004. Documents de I'Institut Scientifique, 19. Beyer, E. 1869. Beitrag zur Kenntis der Fauna des Kalkes von Haina bei Waldgirmes (Wetzlar). Verh. Naturhist. Ver. Rheinland Westfalen.

136 53,56-102. Billings, E. 1869. Description of some new species of fossils with remarks on other already known, from the the Silurian and Devonian rocks of Maine. Proceedings of the Portland Society of Natural History 1,2,104-126. Bultynck, P. & Hollard, H. 1980. Distribution Comparee de Conodonts et Goniatites Devoniens de Plaines du Dra, de Ma 'der et du Tafilalt (Maroc). Leuven University Press, Belgium, 73 pp. Bultynck, P. & Walliser, O.H. 2000. Devonian boundaries in the Moroccan Anti-Atlas. Courier Forschumgsinstitut Senckenburg, 225, 211-226. Cisne, J. L. 1968. Discovery of the subfamily Tropidocoryphinae (Trilobita) in North America. Journal of Paleontology, 42(1), 231-232. Chatterton, B.D.E. & Speyer, S.E. 1997. Ontogeny. Pp. 173-247 In: Whittington, H.B., et al. (Eds.), Arthropoda 1, Trilobita. Part O. (revised) Treatise on invertebrate paleontology. University of Kansas, Lawrence, Geological Society of America, Boulder, Colorado. Chatterton, B. D. E., Fortey, R., Brett, K., Gibb, S., and McKellar, R. 2006. Trilobites from the upper Lower to Middle Devonian Timrhanrhart Formation, Jbel Gara Zguilma, southern Morocco. Palaeontographica Canadiana, 25, 177pg. Cronier, C., Bignon, A., and Francois, A. 2011. Morphological and ontogenetic criteria for defining a trilobite speceis: The example of Siluro-Devonian Phacopidae. Comptes Rendus Palevol, 10, 143-153. Dalman, J. W. 1827. Om palaederna, eler de sa kallade trilobiterna. Kongliga Svenska Vetenskaps-Akademiens Handlingar (for 1826), 113-152, 226-294. Dopieralska, J. 2009. Reconstructing seawater circulation on the Moroccan shelf of Gondwana during the Late Devonian: Evidence from Nd isotope composition of conodonts. Geochemistry, Geophysics, Geosystems, 10(3), 1-13. El Hassani, A. (ed.). 2004. Devonian neritic-pelagic correlation and events in the Dra Valley (western Anti-Atlas, Morocco). Subcommision on on Devonian Stratigraphy. International Metting on Stratigraphy, Rabat, 1-10 March 2004. Documents de I'lnstitut Scientifique, 19, 1-100. Ellerman, 1.1992. Trilobiten aus dem Unterdevon der Karnischen Alpen / Osterreich. Palaeontographica A, 221, 1-62, pis 1-3. Erben, K. 1966. Uber die Tropidocoyphinae (Tril.) - Liefg. 1. Neues Jahrbuch furGeologie undPalaontologie. Abhandlungen, 125, 170-211. Feist, R. 1976. Systematique, phylogenie et biostratigraphie de quelques Tropidocoryphinae (Trilobita) du Devonien Fran9ais. Geobios, 9, 47-80. Feist, R. 2003. Biostatigraphy of Devonian tropidocoryphid trilobites from the Montagne Noir (southern France). Bulletin of Geosciences, 78, 432-446.

137 Feist, R. and Clarkson, E. N. K. 1989. Environmentally controlled phyletic evolution, blindness and extinction in Late Devonian tropidocoryphine trilobites. Lethia, 22, 359-373. Fetah, S. E. M., Bensaid, and Dahmani, M. 1988. Carte Geologique Maroc: Todhra-Ma'der (Anti-Atlas oriental, zones axiale et peripherique Nord du Sud). Editionas de Service Geologique de Maroc, Notes et Memoires No. 243. Royaume du Maroc. Ministere de l'Energie et des Mines, Rabat. Fortey, R. A. & Owens, R. M. 1975. Proetida - a new order of trilobites. Fossils and Strata, 4, 227-239. Frantisek, H. and Vanek, J. 2000. Novf proetidnf trilobiti z prazskeho stupne (spodnf devon) Prazske panve. Palaeont. Bohemiae, 6,16-26. Gibb, S. 2005. Some proetids (Class Trilobita) from the Lower to Middle Devonian of southern Morocco. M.Sc. thesis, University of Alberta, Edmonton Alberta, Canada. 21 lpg. Gibb, S. & Chatterton, B. D. E. 2007. Timsaloproetus new genus (Proetida: Trilobita) and included species from Lower and Middle Devonian strata of southern Morocco. Journal of Paleontology, 81(2), 352-367. Gibb, S. & Chatterton, B. D. E. 2010. Gerastos (Order Proetida; Class Trilobita) from the Lower and Middle Devonian of the Southern Moroccan Anti-Atlas region. Palaeontographica Canadiana, 30, 89pg. Hammer, 0. 2010. PAST - PAleontological STatistics ver.1.99. Harrington, H.J. 1959. Tropidocoryphidae. Pp. 397-398 in Morre, R. C. (ed.) Treatise on invertebrate paleontology, part 0, Arthropoda 1. Geological Society of America, Boulder, and University of Kansas Press, Lawrence. Hawle, I. and Corda, A. J. C. 1847. Prodrom einer Monographic der bohmischen Trilobiten. Abhandlungen der Koniglichen Bohmischen Gesellschaft der Wissenschaften, V. Folgle, 5(5), 1-176. Hollard, H. 1978. Correlations entre niveaux a brachiopodes et a goniatites au voisinage de la limite Devonien inferieur-Devonien moyen dans les plaines du Dra (Maroc presaharien). Newsletters of Stratigraphy, 7(1), 8-25. ICZN. 1999. International code of zoological nomenclature. The International Trust for Zoological Nomenclature. Jansen, U., Becker, G., Plodowski, G., Schindler, E., Vogel, O. & Weddige, K. 2004. Pragian and Emsian near Aouinet Torkoz (SW Dra Valley, Morocco). Pp. 75-84 in A. El Hassani (ed) Devonian Neritic-Pelagic Correlation and Events in the Dra Valley (Western Anti-Atlas, Morocco). Subcommission on Devonian Stratigraphy. International Meeting on Stratigraphy, Rabat, 1-10 March 2004. Documents de Vlnstitut Scientifique, 19. Jell, P. A. and Adrain, J. M. 2002. Available generic names for trilobites. Memoirs of the Queensland Museum, 48(2), 331-553. Kowalski, H. 1990. Trilobiten aus dem Devon der Eifel. Der Aufschluss. Zeitschrift fur die Freunde der Mineralogie und Geologie, 33, 11-63. 138 Lieberman, B. S. and Karim, T. S. 2010. Tracing the trilobite tree from root to the tips: A model marriage of fossils and phylogeny. Structure and Development, 39, 111-123. Liitke, F. 1980. Zur Evolution der altpalaozoischen Proetina (Trilobita). Senckenbergiana lethaea, 61, 73 -144. Maddison, D. R. and Maddison, W. P. 2001. MacClade 4: Analysis of Phylogeny and Character Evolution. Version 4.08. Sinauer Associates, Sunderland, Massachusetts. Maillieux, E. 1904. Quelques mots sur les trilobites du Couvinien des environs de Couvin. Bulletin de la Societe beige de Geologie, de Paleontologie et d'Hydrologie, 17, 579-582. McKellar, R. and Chatterton, B. D. E. 2009. Early and Middle Devonian Phacopidae (Trilobita) of southern Morocco. Paleontographica Canadiana, 28, llOpg. Morzadec, P. 1969. Le Devonien de la Rive Nord de la Riviere du Faou (Finistere). Etude stratigraphique - Etude des Trilobites. Extrait du Bulletin de la Societe Geologique et Mineralogique de Bretagne, Rennes, 58pg. Morzadec, P. 2001. Asteropyginae trilobites from the Devonian of the Anti-Atlas (Morocco). Palaeontographica Abteilung A, 262, 53-85. Novak, O. 1890. Vergleichende Studien an einigen Trilobiten aus dem Hercyn von Bicken, Wildungen, Greirenstein und Bohmen. Paldontologische Abhandlungen, 1, 1-46. Ormiston, A. R. 1967. Lower and Middle Devonian Trilobites of the Canadian Arctic Islands. Geological Survey of Canada Bulletin, 153, 1-148. Owens, R. M. 1973. British Ordovician and Silurian Proetidae (Trilobita). Palaeontographical Society Monographs, 127, 1-98, pis 1-15. Philip, G. 1991. Political: Africa, Philip's World Atlas. Reed International Books Ltd., London. Prevosti, F. J. and Chemisquy, M. A. 2010. The impact of missing data on real morphological phylogenies: influence on the number and distribution of missing data. Cladistics, 26, 326-339. Pribyl, A. 1946. O nekolika novych trilobitovych rodech z ceskeho siluru a devonu. Priroda, Brno, 38, 89-95. Pribyl, A. 1965. Proetidni trilobiti z novych sbiru v eeskem siluru a devonu. II. east. Proetiden aus neueren Auf- sammlungen im bohmischen Silur und Devon (Trilobitae) -1. Easopis Narodniho Muzea, oddil Peirodovidny, 134,91-98. Pribyl, A. and Vanek, J. 1978. Studie zu einigen neuen Trilobiten der Proetidae-Familie. Acta Universitatis Carolinae, Geologica, 1, 163-182. Richter, R. & Richter, E. 1919. Proetiden aus neureren Aufsammlungen im vogtlandischen und sudetischen Oberdevon. Senckenbergiana, 1, 97-130. Ricther, R. & Richter, E. 1919. Der Proetuden Zwieg Astycoryphe - Tropidocoryphe - Pteroparia. Senckenbergiana. 1, 1-17, 25-51.

139 Richter, R. & Richter, E. 1949. Die Trilobiten der Erdbach-Zone (Kulm) im Rheinischen Schiefergebirge und im Harz. I. Die Gattung Phillibole. Senckenbergiana, 30, 63-94. Salter, J. W. 1864. A Monograph of British Trilobites from the , Silurian, and Devonian Formations. The Palaeontological Society, London, 224 pp. Scotese, C. R., Boucot, A. J., and McKerrow, W. S. 1999. Gondwanan palaeogeography and palaeoclimatology. Journal of African Earth Science, 28(1), 99-114. v Snajdr, M. 1976. New proetid Trilobita from the Silurian and Devonian of the Barrandian (Czechoslovakia). Qas. Mineral. Geol., 21(3), 313-318. Snajdr, M. 1977. New genera of Proetidae (trilobita) from the Barrandian, Bohemia. Vestnik Ustredniho ustavu geologickeho, 52, 293-297. Snajdr, M. 1980. Bohemian Silurian and Devonian Proetidae (Trilobita). Rozpravy Ustredniho ustavu geologickeho, 45, 1-324. Stafford, E. S. and Leighton, L. R. 2011. Vermeij Crushing Analysis: Anew old technique for estimating crushing predation in gastropod assemblages. Palaeogeography, Palaeoclimatology, Palaeoecology, 305, 123-137. Steininger, J. 1831. Bemerkungen iiber die Verseinerungen, welche in dem Ubergangs- Kalkgebirge der Eifel gefunden were. Beilage zum Gymnasial-Programmschrift zu Trier, Blattau, 46p. Swofford, D. 2002. PAUP*4.0bl0 (Altivec): Phylogenetic Analysis Using Parsimony. Sinauer Associates, Inc. Viersen, A. P. van & Prescher, H. 2010. Taxonomy and biostratigraphy of some proetid trilobites in the Middle Devonian of the Ardennes and Eifel (Rhenohercynian Zone). Bulletin de I'lnstitut royal des Sciences naturelies de Belgique, Sciences de la Terre, 80, 5-45. Viersen, A.P. van, Prescher, H. & Savelsbergh, J. 2009. Description of two new trilobites from the Ahrdorf Formation (Middle Devonian) at the "Trilobitenfelder" of Gees, Eifel, Rhenish Mountains. Bulletin de I'lnstitut royal des Sciences naturelles de Belgique, Sciences de la Terre, 79, 43-53. Wagner, P. J. 2000. Exhaustion of morphological character states among fossil taxa. Evolution, 54(2), 365-386. Wiens, J. J. 2003. Incomplete data, missing taxa, and phylogenetic accuracy. Systematic Biology, 52(4), 528-538. Whittington, H. B. and Kelly, S. R. A. 1997. Morphological terms applied to Trilobita. Pp. 313-329 in Moore R. C. and R. L. Kaesler (eds.) Treatise on invertebrate paleontology, part O, Arthropoda 1, Trilobita revised, v. 1: Introduction, Order , Order . Geological Society of America, Boulder, and University of Kansas Press, Lawrence. Whidborne G. F. 1889. A Monograph of the Devonian Fauna of the South of

140 England. Part I. The Fauna of the Limestones of Lummaton, Wolborough, Chircombe Bridge and Chudleigh. Palaeontographical Society, 42, 1-46. Wright, A. J. & Chatterton, B. D. E. 1988. Early Devonian Trilobites from the Jesse Limestone, New South Wales, Australia. Journal of Paleontology, 62, 93 - 103.

141 Chapter 3: Systematic Palaeontology

Cornuproetus and Diademaproetus (Class Trilobita, Order Proetida) from Morocco and their impact on the cornuproetid, diademaproetid relationship.

Introduction

Cornuproetus Richter & Richter, 1919 is a common proetid trilobite genus, found in Morocco and several European countries. In addition to

Cornuproetus, the morphologically similar genus, Diademaproetus Alberti, 1964, is found in many of the same geographic areas and through the same stratigraphic interval. Cornuproetus and Diademaproetus are members of the proetid subfamily

Cornuproetinae Richter & Richter, 1956 and are thought to be closely related

(Alberti 1969). However, despite suggestion of a close relationship between

Cornuproetus and Diademaproetus, quantitative statistical analysis of their morphology, specifically their diagnostic characteristics, has never been undertaken. Furthermore, a phylogenetic analysis involving both genera, to examine how they relate to one another, has also never been performed. Without quantitative statistical analysis of the morphology of both genera and phylogenetic testing of their relationships, valid diagnostic and synapomorphic characters cannot be identified. In turn, if both diagnostic and synapomorphic characters are not identified, discerning their relationships becomes difficult. Ultimately, understanding of both Cornuproetus and Diademaproetus diagnostic characters and their taxonomic (i.e., genus and species) relationships is critical for understanding their larger scale relationships and evolutionary processes.

142 This study surveys newly discovered Early to Middle Devonian aged

Cornuproetus and Diademaproetus trilobites of southern Morocco in conjunction

with previously described species of Morocco and Europe. As a result of the

abundant and very well preserved (complete and articulated) nature of these

Moroccan cornuproetids, and the commercial mining of trilobites by the local

Berber population, this study documents the largest, most complete, and best

preserved collection of Diademaproetus to date. As previous work on these

primarily employed mostly partial and incomplete material, description and

analysis of this study's collection allows for not only the redescription of

previously known species, but also proper diagnosis and description of new

species. Furthermore, increased species level diversity of Diademaproetus allows

better understanding of their diagnostic characteristics and evolution.

The goals of this study are twofold. First, statistical analyses of

Cornuproetus and Diademaproetus characters is used to determine character and

character state validity. Special attention is paid to characters thought to be evolutionarily significant and many of these were previously discussed by Alberti

(1969), Snadjr (1980), Chatterton et al. (2006), and Basse (2010). New characters and character states are included where statistical support suggests they are informative. Second, cladistic analysis, using validated characters and character states, is used to examine the relationships and validity of Cornuproetus,

Diademaproetus, Interproetus Snadjr, 1977, Taciproetus Basse, 2010, and new

Moroccan material. In association with the cladistic analysis, five new species of

143 Diademaproetus: Diademaproetus rudimentus, D. issoumourensis , D. corrugatus,

D. auxiliarus, and D. langus, and one new genus Pontoproetus, with two new

species (Pontoproetus truncatus and P. granulosus), are proposed and described.

Results of this analysis not only increase the known diversity of the

Cornuproetinae, but also improve our understanding of their evolution. This work

will not only increase our understanding of Cornuproetus, Diademaproetus, and

morphologically similar taxa, but also present a suitable methodology in which to

examine these taxa, and other trilobites.

Previous work

Richter & Richter (1919) originally diagnosed Cornuproetus with

Cornuproetus cornutus cornutus (Goldfuss, 1843) as its type species, distinguishing it from Gerastos Goldfuss, 1843. Besides individual species descriptions by Kegel (1926), Erben (1952), and Hass (1968), the majority of

Cornuproetus diversity was not documented until the Morocco and German trilobite surveys of G.K.B. Alberti (1964; 1966; 1967; 1969). Most significantly,

Alberti (1964; 1966; 1967; 1969) described and documented 17 subgenera of

Cornuproetus, with Diademaproetus originally being designated Cornuproetus

(Diademaproetus) (Alberti 1964).

Cornuproetus (Cornuproetus) was the most diverse of these 17 subgenera, with 18 new species being described from Morocco and Germany by Alberti alone from 1964-1969 (Alberti 1964, 1966, 1967, 1969). Within the 18 described

144 species, Alberti (1969) categorized two morphologically distinct subpopulations

of Cornuproetus (Cornuproetus), the first having a violin shaped (well constricted) glabella and wide palpebral lobes, while the second has a subquadrate

(poorly constricted) glabella and narrower palpebral lobes. Cornuproetus

(Diademaproetus), on the other hand, had a much smaller diversity than

Cornuproetus (Cornuproetus), consisting of only 3 species (Alberti 1969). Based on the morphological similarity of all 17 subgenera, their stratigraphic ranges, and the high diversity of Cornuproetus (Cornuproetus) compared to all other subgenera, Alberti (1969, fig. 17) proposed Cornuproetus (Cornuproetus) as the main ancestral lineage that, throughout its stratigraphic range, gave rise to the other cornuproetid subgenera. Given this Cornuproetus (Cornuproetus) ancestry,

Cornuproetus (Diademaproetus) was believed to have originated during the late

Emsian (Alberti 1969).

Following the works of Alberti (1964; 1966; 1967; 1969), descriptions of new species for all of his cornuproetid subgenera eventually led to all 17 subgenera being elevated to genera. Elevation of the 17 subgenera to genera was primarily the result of the recurrence of once "unique" characters as new taxa were discovered, making the characters taxonomically meaningful and no longer

"unique" (Owens 1973). As a result, Cornuproetus (Diademaproetus) was elevated to Diademaproetus. Following this elevation, Cornuproetus diversity continued to grow, with species being described primarily from outside Morocco and within Europe (Owens 1973; Snajdr 1980). Diademaproetus also saw

145 increased diversity, with the description of more European species (Kowalski

1975) and additional Moroccan species (Hammann 1976; Chatterton et al. 2006).

Despite taxonomic elevation and increased diversity in both Cornuproetus and

Diademaproetus, distinguishing these taxa from one another remained difficult, as

diagnostic characters are variably present in the species of both genera.

Ultimately, lack of consensus with regards to the diagnostic characters of both

genera has made proper diagnosis of new material to the appropriate genus

difficult. Furthermore, this problem has been compounded by the diagnosis of

many previously named species being based on largely incomplete and poorly

preserved type specimens.

Stratigraphy and geology

The locations of the three basins in southern Morocco and the localities

where specimens of this study were found are indicated in a series of general,

basin, and stratigraphic diagrams (Figures 3-1 to 3-4). Additional information and

descriptions of the individual sites and horizons, from which specimens were collected, are presented below.

Tindouf Basin

Jbel Gara el Zguilma. Jbel Gara el Zguilma, hereafter referred to as

'Zguilma,' of the Dra Valley region displays three trilobite-bearing horizons

within the Timrhanrhart Formation (Hollard 1978; Bultynck & Hollard, 1980;

146 Bultynck & Walliser, 2000; El Hassani, 2004; Chatterton et al. 2006). The lowest

two of the three horizons (ZGEE1 and ZGEE2) are located at 29° 42' 35.2" N and

06° 42' 10.2" W, 53 km south southeast of the town Foum Zguid, and are late

Emsian in age (Chatterton et al. 2006; Gibb & Chatterton 2010). The stratigraphically highest horizon (ZGEE3) is located at 29° 42' 18.5" N and 06°

42' 11.5" W, and is Eifelian in age (Gibb & Chatterton 2010). Diademaproetus

praecursor Alberti, 1969 is found in all three horizons, while D. mohamedi

Chatterton et al. 2006 is found only in horizon ZGEE3. Both species and the associated trilobite faunas found with them were described in detail by Chatterton et al. (2006).

An additional locality, north of Jbel Hamsai' and closer to Foum Zguid than

Zguilma, was also surveyed. Referred to as FZ1, this locality displays similar trilobite faunas to those that are found in the horizons at Zguilma, and is thought to be of similar late Emsian age and part of the Timrhanrhart Formation (Gibb &

Chatterton 2010). Both Diademaproetus praecursor and D. mohamedi are found at this locality.

Assa. Located at 28° 30' 06.2" N and 09° 27' 43.4" W, the locality of Assa is positioned northwest of the town of Assa, in southern Morocco. Strata at Assa are part of the Khebchia Formation of the Tindouf basin, and are late Emsian in age (Becker et al. 2004a; Becker et al. 2004b; Hollard 1978; Jansen et al. 2004).

Diademaproetus auxiliarus new species is a rare trilobite found at the locality, with three specimens known.

147 Ma'der Basin

Jbel Oufatene. Jbel Oufatene is located in the southwest portion of the

Ma'der Basin, west northwest of the village Lahfira. Within the section surveyed

at Jbel Oufatene, two trilobite-bearing horizons were found to contain

Diademaproetus. The first Diademaproetus bearing horizon is located at 30° 45'

48.6" N and 04° 40' 42.7" W and is part of the El Otfal Formation. Named for its high abundance of Diademaproetus, the horizon contains two diademaproetid species, the more abundant D. langus new species and rarer D. praecursor. Based on the trilobite and conodont faunas, the horizon is Eifelian in age (Gibb &

Chatterton 2010).

The second trilobite-bearing horizon containing Diademaproetus at Jbel

Oufatene is the Psychopyge horizon, which is located approximately 50 meters stratigraphically below the Diademaproetus horizon and is part of the Tazoulai't

Formation. Diademaproetus praecursor is the only diademaproetid found in this late Emsian aged horizon (Gibb & Chatterton 2006).

Taharajat. Located at 30° 47' 48.0" N and 04° 54' 20.3" W, Taharajat is located south of Jbel Oufatene and contains two main trilobite-bearing horizons, the Morocconites horizon and the Hollardops horizon. Both horizons are only separated by three meters stratigraphically and have extremely similar trilobite faunas, specimens from both horizons are treated as a single collection for the entire locality. Combination of both horizons into a single collection is also

148 helpful in cataloging purchased specimens as the matrices from both horizons are identical and cannot be used to determine which horizon the specimens came from, and Berber miners of this locality seldom remember or note which horizon they collected specimens from. Both horizons are part of the Tazoulai't Formation, and based on their trilobite and conodont faunas, are late Emsian in age (Gibb &

Chatterton 2010). Both Diademaproetus rudimentus new species and

Diademaproetus currugatus new species are found at Taharajat, with

Diademaproetus rudimentus being the more abundant of the two species.

Jbel Issoumour. Jbel Issoumour, hereafter referred to as "Issoumour," is a long, curved boomerang-shaped range that is located northwest of the village bou

DIb and north-northwest of Jbel Oufatene and Taharajat. Within the locality diademaproetids are found at four stratigraphic intervals, all of which are located on the western and northern faces of the Issoumour range. Diademaproetus issoumourensis new species and Diademaproetus praecursor are found at and just below the Psychopyge horizon (30° 58' 25.2" N and 04° 01' 52.3" W) of the Er

Remlia Formation (Gibb & Chatterton 2010). Approximately one and a half meters above the Psychopyge horizon, D. praecursor is found in the Heliopyge/

Kayserops horizon. D. praecursor is also found at the top of Jbel Issoumour (30°

58' 25.2" N and 04° 01' 49.7" W) above the cliff-forming "Grande Calcaire" unit, in the stratigraphically higher Lobopyge and Paralejurus horizons of the El Otfal

Formation (Gibb & Chatterton 2010).

149 Timarzite. Located east-northeast of the village of bou Dib, the locality of

Timarzite is located just north of the Number 12 (N12) highway and west-

southwest of the town of Mecissi, at 31° 10' 40.7" N and 04° 58' 55.7" W. Based

on the presence of the conodonts Polygnathus ensensis and Tortodus kockelianus

kockelianus (Bultynck 1987), the locality of Timarzite is believed to be late

Emsian to Eifelian in age (Belka et al. 1997, Figure 3). Pontoproetus truncatus

new species is the only diademaproetid found at the locality.

bou Dib. Located east of Jbel Issoumour and a few kilometers north of the

village of bou Dib, Diademaproetus is found in the Eifelian aged strata of the El

Otfal Formation (Hollard 1974; Morzadec 2001; Gibb & Chatterton 2007, 2010).

Diademaproetus mohamedi is found in the Harpes/Thysanopeltis horizon located

at 31° 05' 16.6" N and 04° 52' 32.2" W, allowing for redescription of this once

incomplete species.

Jbel El Mrakib. Located southeast of the village of Lahfira and northwest of Zireg, Jbel El Mrakib, hereafter referred to as "Mrakib," contains multiple trilobite bearing horizon (Gibb & Chatterton 2010). Pontoproetus granulosus new species is the only diademaproetid found at the locality, stratigraphically equivalent to the Morocconites/Metacanthina horizon of Jbel Oufatene. The

Morocconites!Metacanthina horizon of the Mrakib section is part of the Tazoulai't

Formation and is Emsian in age (Gibb & Chatterton 2010).

Jbel Zireg. Jbel Zireg, hereafter referred to as "Zireg," is located southeast of Mrakib, in the southeastern portion of the Ma'der basin. Pontoproetus

150 truncatus new species is found in the Eifelian Thysanopeltis horizon (30° 36'

46.1" N and 04° 32' 10.1" W), 26 meters above a stratigraphically lower

Ceratarges horizon (Gibb & Chatterton 2010).

El Achana. Located at 30° 51' 17.2" N and 04° 58' 35.0" W, the

Kolihapeltis horizon and the stratigraphically lower Dicranurus horizon of El

Achana both contain Diademaproetus rudimentus. Given the low stratigraphic

position of these horizons and the occurrence of D. rudimentus in stratigraphically lower sections (i.e., late Emsian aged Taharajat), it is not surprising that it is found at El Achana. These levels are Pragian in age.

Tafilalt Basin

Talawarite. Talawarite is located east-southeast of Hamar Laghdad

(Alberti 1969) in the Tafilalt Basin, at 31° 16' 28.5" N and 03° 53' 29.4" W (Gibb

& Chatterton 2010). Diademaproetus praecursor is the only Diademaproetus found at the locality and was collected from both the Gerastos horizon and surface collected from the surrounding area. The Gerastos horizon itself is part of the Amerboh Group and given its trilobite fauna, is late Emsian in age (Gibb &

Chatterton 2010).

Taouz. Located at 31° 04' 01.3" N and 04° 09' 46.1" W, the locality named

Taouz is actually north-northwest of the town of Merzouga, and is closer to

Merzouga than to the town of Taouz. Diademaproetus praecursor is the only diademaproetid species found at this locality.

151 Terminology

Morphological terminology used in this study follows that outlined by

Whittington & Kelly (1997) in the revised Treatise on Invertebrate Paleontology.

This includes the use of abbreviations for some features (e.g., Occipital Lobe =

LO). Specific proetid terminology follows the works of Richter & Richter (1949),

Owens (1973), and Snajdr (1980). Specifically, terminology of the measurement

of tagma and their features follows Owens (1973), terminology of the shape of sclerites and their features follows Snajdr (1980), and terminology of the cephalic sutures follows Richter & Richter (1949). Where appropriate, the anatomical directions, distal and proximal, are used to indicate a feature's proximity to the trilobite's midline (sagittal line). Cornuproetus and Diademaproetus specific terminology is adapted from both Alberti (1969) and Chatterton et al. (2006).

Refer to Figure 3-6 for all measurements and morphometric landmarks examined in this study.

Materials and methods

Collection of the majority of the cornuproetid and diademaproetid material examined in this study was completed by the third author, B.D.E. Chatterton, and the various members of his research group during Moroccan field work in the spring of 2009 and during previous visits to the same areas. Extensive mining of the trilobite-bearing horizons visited during field work, by the local Berber

152 population, not only permitted the collection of specimens directly from the horizons themselves and their directly associated spoil heaps, but also the purchase of rarer specimens from local fossil shops and the miners working the horizons. Without the work of many of these miners, rare species would go undocumented, as the amount of time necessary to find them is beyond the capability of a small research group and the time constraints of field work in the area. In the case of purchased material, stratigraphic control of specimens was ascertained by cross referencing the matrix of purchased specimens with the matrix of the locality from which the specimen was thought or said to have come.

Only when matrices matched the rocks at the supposed locality and conspecific specimens (complete or incomplete) were found at the indicated locality and horizon were purchased specimens used in this study.

Following field work, specimens were either prepared at the University of

Alberta by the authors or Alan Lindoe, a professional fossil preparator, or in

Morocco by our local guide and preparator Hammi Ait H'ssaine and his family.

Some purchased specimens also received some preparation from the miners and/ or fossil shop dealers. However final preparation of such specimens was completed by the authors, Alan Lindoe, or Hammi Ait H'ssaine. Specimens prepared at the University of Alberta were done so using pin-vice and pneumatic chisel, while specimens prepared by Hammi Ait H'ssaine were usually prepared using air propelled calcite abrasion. All specimens are housed in the University of

Alberta, Department of Earth and Atmospheric Sciences type collection.

153 Specimens were painted black and coated with an ammonium chloride sublimate prior to photography in order to enhance fine detail. Either a Canon 40d digital camera, outfitted with a 60 mm macro Canon EF lens and macro flash, or a

Nikon D2X digital camera, outfitted with a 100 mm Zeiss zf macro lens, was used

to photograph the specimens. Line drawings of specimens and their various

features were also constructed by the primary author from the photographs and the

specimens, using Autodesk SketchBook Pro 5.1.

For the phylogenetic analysis, measurements for character and character state statistical testing and coding were primarily taken from specimens. For specimens not available, measurements were taken strictly from dorsal photographs. In all instances, measurements were taken three times on separate occasions, with the average of the three being reported. For specimens where no hand sample or dorsal image were obtainable from the literature or the institution that housed them, measurements were not taken and the characters associated with those measurements were coded as unknown ("?"). Furthermore, description alone was not considered sufficient for character coding; if characters were not directly viewable, using either hand sample or photograph, they are coded as unknown.

Morphometric characters were measured using Bookstein shape coordinate analysis (here after referred to as "Bookstein shape analysis")

(Bookstein et al. 1985, appendix 4). Bookstein shape analysis allows for the comparison of a landmark's position across specimens of varying size. In

154 accordance with the procedures of Bookstein shape analysis ((Bookstein et al.

1985, appendix 4), glabella length (Ai + A4), not cephalon length (A, Figure 3-5), was chosen as the comparative base line because it represents the largest cephalic measurement which is least taxonomically diagnostic among the taxa analyzed

(lacks variability at both the species and generic levels). For example, the length of the preglabellar area (A2 + A3, Figure 3-5) is a diagnostic character within the

Cornuproetinae, and therefore morphologically variable between cornuproetid species. As such, the length of the cephalon (A), which includes the length of the preglabellar area (A2 + A3, Figure 3-5), is not a suitable base line for Bookstein shape analysis as it is partially influenced by taxonomic variation. As a result, clustering of landmark positions between specimens can be the product of base line variation between specimens, rather than variation in examined landmark's position.

Statistical testing of characters and character states in this study employed two different statistical tests, the Mann-Whitney U test and Hotelling's T2 test

(with a Bonferroni correction where appropriate). All statistical testing was accomplished using PAST (Hammer 2010), and a 95% confidence interval was used to determine statistical separation (separate character states). Whenever multiple statistical tests were performed on a single character, statistical support was required from all tests. All potential character states (populations) were initially identified using natural breaks in the data distributions. If any one of the statistical tests could not statistically support the differentiation of character states

155 (populations), then the character was omitted. Only when character states

(populations) were found to be statistically different from one another by all suitable tests were they included in the phylogenetic analyses. Two different statistical tests were run depending on the type of data (univariate or multivariate)

and the number of potential character states (populations) present for each

character. The Mann-Whitney U test is a non-parametric test that measures the equality of two population's means, and was used in conjunction with single

variable (univariate, non-coordinate based morphometric) data. We chose to use

the Mann-Whitney U test instead of the Students t-test or Welch t-test as the

Mann-Whitney U test is considered a more conservative and appropriate test for examining data and populations with unequal variance, which was often the case for the present study's characters.

When examining multivariate (bivariate, coordinate based morphometric) data, a Hotelling's T2 tested was used. Hotelling's T2 test is a multivariate analog of a Students t-test and measures the equality of means between populations in multivariate space. The test itself reports back two values, first the probability that the examined means are distinct (p-value). In conjunction with the Hotelling's T2 test p-value, a percent correct classification value (%CC), which is a measure of the derived discriminant function to identify specimens to the correct populations

(Hammer 2010) was also used to verify that populations where distinct from one another. Whenever three or more potential character states (populations) were present, it was necessary to conduct multiple pairwise Hotelling's T2 tests. In

156 these cases, a Bonferroni correction was applied to the Hotelling's T2 test. By applying a correction to the p-value of individual population vs. population tests when comparing multiple (more than two) populations, a Bonferroni correction reduces the individual population vs. population test threshold for rejecting the null hypothesis so that the overall threshold of comparing all populations to one another remains the same (Stafford and Leighton 2011). Furthermore, only a standard Bonferroni correction was employed, and not a sequential Bonferroni, as the standard Bonferroni correction is deemed a more conservative approach.

PAUP* version 4.0b10 (Altivec) (Swofford 2002) was used to run the phylogenetic analyses. MacClade 4.08 (Maddison & Maddison 2005) was used to view the results of the analyses. All characters are equally weighted and all character states are unordered and reversible. Exhaustive searches using global parsimony were performed. Bremer support values for the best trees were calculated by adding additional steps to produced trees and observing where dichotomies collapsed into polytomies. Bootstrap and Jacknife tests, with 2500 replicates, were also used to examine tree stability.

Phylogenetic analysis

Even though, in the past, numerous cornuproetids have been described and discussed, no phylogenetic testing of proposed hypotheses has been completed.

Furthermore, the lack of agreement surrounding which characters are suitable synapomorphies for diagnosing Cornuproetus and Diademaproetus has caused

157 distinguishing between these genera to be difficult, with many authors listing different diagnostic characters and supporting or refuting those of others. By examining a mixture of the best known species of both Cornuproetus and

Diademaproetus, along with their proposed ancestor Interproetus Snajdr, 1977, this study aims to elucidate the relationships of these genera. Inclusion of new

Moroccan material into this analysis is used to validate their taxonomy and decipher how these taxa fit into cornuproetid relationships. This analysis was not designed to alter previous taxonomy radically. For the sake of stability, universality, and communication of nomenclature (ICZN 4th edition, Article 23.2) judgement is used when modifying previously proposed taxa and classification.

Coding of the operational taxonomic units (OTUs) examined in this analysis was conducted using individual specimens, with previously described species being exemplified by either their holotype specimen or a specimen of greater completeness, quality and/or documentation. Use of individual specimen based OTUs permits both low and high level taxonomic relationships to be viewed in a single analysis. As Anstey & Pachut (2004) have shown, the implementation of specimen based OTUs can maximize the disparity between low level taxonomic units, allowing for even sub-species level relationships to be viewed. As a result, new specimens forming well supported clades with distinct characters and character states are considered valid taxa, while new specimens that form polytomies or very weakly supported clades in our analysis are considered members of the same taxon. In the case of the new Moroccan species,

158 numerous complete specimens coded the same. This provided confidence in the integrity of the species. For reasons of efficiency, only one of several identically coded specimens was included in the analysis to represent each species. Because this analysis is designed/optimized for discovering relationships among species of

Diademaproetus rather than related outgroup taxa, we are particularly conservative in proposing nomenclatural and taxonomic changes for taxa.

Prior to coding and running the analysis, all quantitative characters were examined statistically. Such methods allow for the variance between specimens and their groups to be distinguished and quantified, something which Cronier et al. (2005, 2011) argued is essential for understanding taxon boundaries. Without comprehensive taxon boundaries, the lines between taxa can be subjective and become blurred, as do their relationships with one another. Therefore, to produce the most accurate and conservative phylogeny, characters and character states should be tested statistically and proven distinct from one another. Furthermore, the use of such rigorous methodology promotes better overall taxonomy, as both previously known and new material can be distinguished as either plesiotypes or new taxa. This study's use of morphometries and statistically tested characters, in conjunction with a phylogentic analysis, aims to provide the best and most current phylogeny of cornuproetids.

Taxa examined in the analysis

159 A major hurdle to overcome in this analysis was the selection of taxa to be

included in the phylogenetic analysis. With Cornuproetus being such a diverse

genus, which is not particularly disparate, the number of statistically significant

characters and character states is low. Further adding to this problem is the fact

that many previously described cornuproetid species are known from relatively

incomplete and/or loosely associated remains, causing many of their characters to

be coded as "unknown" (?). Depending on the concentration of these "unknown"

characters, two results are possible. First, if a character itself has a high

"unknown"percentage, it is typically labeled as "parsimonious uninformative" by

the phylogenetic analysis and rendered useless in deciphering the phylogeny.

Second, if there is a high percentage of "unknown" characters concentrated within a single or several taxa, those taxa can act as wild cards, appearing in multiple different positions throughout the phylogeny, severely increasing the number of most parsimonious trees and reducing clade stability of the phylogeny (Prevosti &

Chemisquy 2010). In both cases, the effects of high percentages of "unknown" characteristics within a phylogenetic analysis are highly detrimental and should be avoided.

Additionally, given the confusion over which diagnostic characters discriminate between Cornuproetus and Diademaproetus, many characters thought to distinguish cornuproetid genera from one another may turn out to be insignificant. As a result, the number of statistically insignificant characters,

"unknown" characters, and non-diagnostic characters within cornuproetids leads

160 to what Weins (2003) deemed as "character exhaustion," which is defined as a lack of enough suitable and reliable characters to examine relationships within an entire taxon. Although "character exhaustion" is a problem encountered when attempting to analyze many diverse invertebrate taxa, it must be dealt with in order to produce reliable and suitable phylogenies. For a phylogenetic analysis to be useful (in providing hypotheses of relationships), there must usually be more cladistically informative characters and character states than taxa being examined.

If character exhaustion is a problem, the only way to achieve a sufficiently low enough taxon to character state ratio, to produce a reliable phylogeny, is to eliminate taxa, assuming an exhaustive attempt has been made to determine all cladistically informative characters.

Due to the likelihood of character exhaustion caused by the high number of cornuproetid taxa available for examination, many taxa had to be eliminated, and only a small subset could actually be examined in the analysis. The primary goals of this analysis are to distinguish the relationships and diagnostic characters of Cornuproetus and Diademaproetus, while testing the validity, classification, and relationships of newly described Moroccan material. Culling of taxa from the analysis was based on the following general criteria: First, taxa were limited to those directly within or within close palaeogeographic proximity of the Silurian/

Devonian Rheic Ocean, primarily taxa from Morocco and eastern and central

Europe were selected as they are the best preserved and well documented

Cornuproetids. This limitation was imposed, in large part, to validate and classify

161 the new Moroccan material by examining them with the most closely related taxa.

Second, priority was given to the most complete and well preserved taxa. This

was done to minimize the number of "unknown" character states within the

analysis, thus minimizing character exhaustion and phylogenetic instability. Third,

priority was given to type species as they play a vital role in determining where

the boundaries between genera must be drawn.

Coding of specimens was accomplished in two ways. Previously described

taxa were coded for this study using either the holotype, paratypes, or topotypes

of greater completeness, quality (preservation and preparation), and/or

documentation. New Moroccan material was coded using holotype specimens,

which constituted the most complete and best preserved specimens in our

collection. In all cases, only the most complete and best preserved specimens were

selected as OTUs. Refer to Table 3-1, for a list of all taxa included and the more

significant taxa omitted from this analysis, along with explanations concerning

their inclusion and/or omission and indication of the source material. All taxa not

included within Table 3-1 were omitted from the analysis as they were highly

incomplete (greater than 50% of the taxon was coded as "unknown") or because their morphology was substantially different from that of the included taxa,

therefore they probably are not closely related to the taxa analyzed.

Interproetus was included as the potential outgroup in accordance to

Snajdr (1980) and his arguments in favor of it being ancestral to Cornuproetus.

Furthermore, the stratigraphic range of Interproetus, Middle Silurian to Lower

162 Devonian, favors its putative ancestry to Cornuproetus, which ranges from Lower

Devonian to Upper Devonian. Additional genera omitted from the analysis,

primarily to avoid character exhaustion, include Dalejeproetus Snajdr, 1977 and

Montanproetus Basse, 2010. Both genera are morphologically similar to

Cornuproetus and Diademaproetus, with Dalejeproetus thought to be the evolutionary link (i.e., missing link) between Cornuproetus and Diademaproetus

(Snajdr 1980). Dalejeproetus was included in subsequent phylogenetic analyses to test if the relationships determined in the primary phylogeny remained the same, and were stable. Furthermore, Dalejeproetus was added to the phylogenetic analysis to see if it is a suitable evolutionary link between Cornuproetus and

Diademaproetus. Montanproetus on the other hand, whose diagnostic characters were not statistically supported during character and character state examination, was omitted, as a large proportion of its species are relatively incomplete and a majority of its specimens appear to be immature non-holaspid forms. Use of non- holaspid forms in phylogenetic analyses has been strongly argued against by numerous authors, including most recently Cronier et al. (2011), as ontogenetic

(allometric) change of certain exoskeletal features in relation to one another is not equivalent and trilobite morphology varies greatly across ontogeny (Chatterton &

Speyer 1997). For further discussion regarding both Dalejeproetus and

Montanproetus, and their effects on the proposed phylogeny, refer to the results and discussion sections below.

163 Characters and character states

Despite substantial work on both Cornuproetus and Diademaproetus by

Alberti (1969), little to no discussion and analysis has been done on the characters

he highlighted as evolutionarily significant for these genera. Furthermore, absence

of a well supported character set for the Cornuproetidae, and Proetida, encouraged

the construction of one for this analysis. In order to avoid redefining already

accepted proetid terminology, many of the characters examined herein are ones

discussed by Owens (1973) and Snadjr (1980) when examining proetids in

general. Some characters are also reworked diagnostic characters of either genus,

outlined by either Alberti (1969), Kowalski (1975), Chatterton etal. (2006), or

Basse (2010). Lastly, for features where statistical support of particular characters

and character states are present, but no character or character states had been

previously defined for the feature, new characters and character states are created.

Wherever characters and character states of this analysis were deemed

difficult to code based on verbal description alone, diagrams and reference taxa

are included following the character description to further describe and illustrate

the character and its states. Specifically, this provides clearer examples for characters dealing with the sculpture and shape of specific sclerites and their elements. In total, 31 characters are found to be statistically supported and are used in the phylogenetic analysis. Of those 31 characters, 16 are binary and 15 are multistate. Because of the partial and incomplete nature of many previously described taxa and greater character richness of the cephalon and pygidium, there

164 is significant bias towards cephalic (18) and pygidial characters (10) compared to

thoracic characters (2). Furthermore, due to the high morphological similarity and

poorly documented nature of cornuproetid hypostomes and rostral plates (they are

known for very few taxa), no characters were included pertaining to them. The

complete data matrix for this analysis is included as Appendix 1. Full explanations

of all 31 characters and their character states, including justification (statistical

where appropriate) for their use, and details surrounding any modifications made

to them in the present work, are outlined below. For a statistical breakdown of all

quantitative characters on a character by characters and state by states basis, refer

to Table 3-2.

Cephalon

1. Position of cephalic suture angle P: position of cephalic suture angle /? in

relation to length of the glabella (Ai + A4, Figure 3-5). Equations used to

differentiate character states are the product of the discriminate analysis

when preforming the Hotelling T-test using PAST (Hammer 2010) and

represent the midpoint between the discriminant scores of genus level

populations.

States: (0) narrow morph, where ((-51.258*Xp - 0.265*Yp) + 50.954) < 0;

(1) wide morph, where ((-51.258*Xp - 0.265*Yp) + 50.954) > 0.

165 Remarks: Taken as a bivariate measurement and calculated using

Bookstein shape analysis (Boostein et al. 1985, appendix 4), where Xp = P's

distance from the mid-line of the glabella divided by the length of the glabella

((Ai + A4) and Yp = P's distance from the most posterior limit of the cephalon

divided by the length of the glabella (Ai + A4, Figure 3-5). Plotting the position of

point p using Xp and Yp as X and Y coordinates were used to determine if

different populations (character states) existed (Figure 3-6). Examination of the

position of landmark p produced two generic populations (character states)

(Figure 3-6).

2. Position of cephalic suture Angle y: position of cephalic suture angle y in

relation to length of the glabella (Ai + A4, Figure 3-5). Equations used to

differentiate character states are the product of the discriminate analysis

when preforming the Hotelling T-test using PAST (Hammer 2010) and

represent the midpoint between the discriminant scores of genus level

populations.

States: (0) narrow morph, where ((-63.612*Xy + 17.793*Yy) - 39.198) > 0;

(1) wide morph, where ((-63.612*Xy + 17.793*Yy) - 39.198) < 0.

Remarks'. Taken as a bivariate measurement using Bookstein shape coordinate analysis (Bookstein et al. 1985, appendix 4), this character examines

166 the position of landmark (cephalic suture angle) y in relation to the length of the

glabella (Ai + A4, Figure 3-5). Much like character 1, position of cephalic suture

angle /?, this character uses the same methodology, however substitutes landmark

y for landmark /?. Again, like the position of cephalic suture angle /?, examination

of the position of landmark y produced two generic populations (character states)

(Figure 3-7).

Interestingly, the position of landmark y in cornuproetids appears to track

extremely well with the degree to which the anterior portion of the glabella is

constricted. For example, the closer landmark y is to the cephalic midline, the

more constricted the anterior portion of the glabella is. This tracking is most likely

the result of both characters being morphologically correlated; as one changes so

does the other. In order to remain as conservative as possible, and not overweight

(over code) any one particular morphological change within the examined taxa, only the position of landmark y was included as a character in this analysis. The degree in which the anterior portion of the glabella is constricted was omitted

based on its weaker quantitative ability compared to the position of landmark y.

3. Position of cephalic suture angle 5: position of cephalic suture angle S in

relation to the length of the glabella (Ai + A4, Figure 3-5). Equations used

to differentiate character states are the product of the discriminate analysis

when preforming the Hotelling T-test using PAST (Hammer 2010) and

167 represent the midpoint between the discriminant scores of genus level

populations.

States: (0) narrow morph, where ((-18.679*Xs - 73.526*Ys) + 31.167) >

0 and ((-30.025*Xg - 27.264*Ys) + 29.558) > 0; (1) intermediate morph,

where ((-18.679*Xs - 73.526*Ys) + 31.167) < 0 and ((-57.556*X8 +

36.38*Ys) + 8.319) > 0; (2) wide morph, where ((-30.025*X5 -

27.264*Y§) + 29.558) < 0 and ((-57.556*X8 + 36.38*YS) + 8.319) < 0.

Remarks: Taken as a bivariate measurement using Bookstein shape coordinate analysis (Bookstein et al. 1985, appendix 4), this character examines the position of landmark (cephalic suture angle) S in relation to the length of the glabella and occipital lobe (Ai + A4, Figure 3-5). Much like characters one and two, positions of cephalic suture angles /? and y, this character uses the same methodology, but substitutes landmark 8 for landmarks [i and y. Examination of landmark (cephalic suture angle) 6 produced three populations (character states)

(Figure 3-8).

Interestingly, Alberti (1969) defined two subpopulations (evolutionary groups) of Cornuproetus (Cornuproetus) based primarily on their difference in S -

S distance (Alberti 1969, Fig. 18). This character, position of cephalic suture angle d, can also be used to examine the 6 - S ratio. Due to the bilateral symmetry of the cephalon about its sagittal midline, by multiplying the distance of landmark d

168 from the plane of symmetry (cephalic midline) by two one gets the 3 - 3 distance.

Given that both this character and the 3 - 3 distance used by Alberti (1969) measure the same state, the same subpopulations of Cornuproetus (Cornuproetus) should have been present in our study. However, when examining all

Cornuproetus and Diademaproetus taxa, or only the exact specimens Alberti

(1969) used, two subpopulations of Cornuproetus were not recognized statistically. As a result, the findings of our examination do not support the presence of two subpopulations of Cornuproetus or the use of the 3 - 3 ratio as a species level diagnostic character for Cornuproetus.

Until this study, use of only qualitative characters by Alberti (1969), when examining Cornuproetus, has led to unjustifiable characters being used to diagnose species and describe their evolution. Ultimately, these characters make describing new specimens and their relationships difficult, and may also prevent higher level taxa and their relationships from being understood. Therefore, it must be noted that use of qualitative characters alone when describing specimens and their relationships is often not sufficient, and should be avoided or at least supplemented quantitative characters. As shown by our examination of the 3 - 3 distance, quantitative characters are more conservative and precise, and should be employed whenever possible.

4. Preglabellar area length: length of the preglabellar area (A2 + A3) in

relation to (divided by) the length of the glabella (Ai + A4, Figure 3-5).

169 This character measures both the length of the anterior cephalic projection

and the length of the preglabellar area in relation to each other.

States: (0) < 0.22; (1) > 0.22

Remarks: Alberti (1969) diagnosed Diademaproetus primarily on the presence of a large cephalic border and an anteromedian cephalic projection.

Chatterton et al. (2006), however, stated that the size and shape of the anteromedian cephalic projection is highly variable in both Cornuproetus and

Diademaproetus and should only be used as a species level, not the generic level, diagnostic character within the Cornuproetinae. Furthermore, Chatterton et al.

(2006) described and diagnosed Diademaproetus mohamedi as a species lacking an anteromedian cephalic projection while still possessing a large cephalic border.

In order to determine if the presence of an anteromedian cephalic projection and enlarged cephalic border are in fact valid characters when examining

Cornuproetus and Diademaproetus, both were examined statistically and two separate populations (character states) were found to exist (Table 3-2). Inclusion of this character in the phylogenetic analysis is an attempt to determine if it is valid a diagnostic character (synapomorphy) of Diademaproetus.

5. Border shape: anterior limit (border) of cephalon appears cushion-like

(inflated and vaulted) or is flat, with no vaulting or inflation.

170 States: (0) flat; (1) inflated (cushion-like).

Remarks: Besides a large cephalic border, Chatterton et al. (2006) also diagnosed Diademaproetus with the presence of a flat cephalic border. The opposite of a flat cephalic border is a more inflated, "cushion-like" border (Figure

3-9). Note, in order to be as accurate and conservative as possible, this character should be examined using only the cephalic border of the librigena or at the cephalic suture angle a of the cranidium. Examination of this character using the preglabellar area, between the left and right cephalic suture angles a, should be avoided as the presence of an anteromedian cephalic projection and its associated furrow (depression) can make the cephalic border appear inflated when it is not.

This character was included to test its validity as a diagnostic (synapomorphic) character of Diademaproetus.

6. Number of border rims: number of border rims (fine parallel ridges)

running around the anterior border of the cephalon, between the cephalic

margin and cephalic border furrow. Counted anterior of the glabella,

between the right and left sided cephalic suture angles a of the cranidium.

States: (0) 1 or 2; (1) 3 or 4; (2); 5+.

171 Remarks'. Upon examination of the large number of specimens within our

collection, the number of cephalic border rims present on morphologically similar specimens from the same trilobite-bearing horizons appears to be quite stable,

varying by only a single ridge in rare occurrences. This variance is typically found in specimens with poorer preservation and/or preparation quality, and is most likely the result of the same, not actual absence of the ridge. In order to be as

conservative as possible, coding of this character employed character states with ranges of the number of parallel ridges, to account for minor variance that was not

the result of poor preservation and/or preparation. Note, as additional and more

minor border rims are typically present around the genal spine and lateral cephalic border, their numbers are most stable in the preglabellar area, and therefore should be coded from that region.

7. Truncated anterior portion of cephalic border furrow: cephalic border

furrow is truncated (does not exist) anterior of the glabella, within the

preglabellar field. Also appears as if the cephalic border furrow points

posteriorly once crossing the cephalic sutures towards the cephalic

midline, running into the anterior of the glabella and not forming a

transverse furrow within the preglabellar area (Figure 3-10).

States: (0) truncated; (1) not truncated.

172 Remarks: Unlike all other cornuproetids, truncation (absence) of the

cephalic border furrow in the preglabellar area is only found in species of the new

genus Pontoproetus. Note when coding this character, the presence of the

depression (furrow) associated with an anteromedian cephalic projection may take

the place of the cephalic furrow in the preglabellar area. However, if the

depression of the anteromedian cephalic projection is present, it will not connect

to the cephalic border furrow and still display a truncated cephalic border furrow.

Careful attention needs to be used when coding this character, as these two

furrows can be confused with one another.

8. Presence of epiborder furrow: presence of an additional cephalic border

furrow located between the cephalic border and the genal field. Best

observed anterolateral and lateral of the palpebral lobe, on the distal

portion of the genal field.

States: (0) present; (1) absent.

Remarks: Both Snajdr's (1980) and Chatterton etal. s (2006) diagnoses of

Diademaproetus included the presence of epiborder (additional) border furrows on the cephalon. Primarily, these epiborder furrows appear as additional smaller and less robust furrows between the cephalic border furrow and the genal field on the librigena. The size and number of these epiborder furrows also appear to be

173 quite variable, with some specimens having epiborder furrows that encase the entire genal field or just the area lateral of the eyes on the librigena. We included this character in our analysis in order to determine if it is a valid diagnostic

(synapomorphic) character of Diademaproetus.

9. Length of genal spine: length of the genal spine in relation to the entire

length of the exoskeleton. Relatively measured using the genal spines

point of termination in relation to more posterior sclerites of the

exoskeleton.

States'. (0) terminates anterior of pygidium, adjacent to the thoracic region;

(2) terminates posterior of the thoracic section, adjacent to or behind the pygidium.

Remarks: Alberti (1969), Snajdr (1980), and Chatterton et al. (2006) all considered the length of the genal spine to be a diagnostic character of

Diademaproetus. In particular Alberti (1969) diagnosed Diademaproetus as having a genal spine that terminates within the pygidial region in both fully and flat articulated specimens. Inversely, Cornuproetus has a genal spine that is shorter, terminating adjacent to the thoracic region, instead of the pygidial region, in fully articulated and flat specimens. This character is included in this analysis

174 in order to determine if it is a diagnostic (synapomorphic) character of

Diademaproetus.

10. Shape of glabella: relative shape (outline) of the glabella when viewed

dorsally (Figure 3-11).

States: (0) subquadrate; (1) subcylindrical; (2) weakly violin shaped; (3) strongly violin shaped.

Remarks: Snajdr (1980) outlined the genus and species level diagnostic characteristics of the proetid glabella. Alberti (1969) specifically outlined glabellar shape as an additional character for his division of Cornuproetus into two subpopulations (Alberti 1969, Fig. 18). Despite Alberti's (1969) use of this character in splitting Cornuproetus, we could not find any support for his findings when either all the comuproetids we examined were considered or when just those that Alberti (1969) examined. Both subpopulations of Cornuproetus that Alberti

(1969) defined do not display a uniform glabellar shape, with both populations showing a variety of glabellar shapes. Despite this character's lack of support for diagnosing the supposed subpopulations of Cornuproetus, it did prove to be a conservative character of morphologically similar specimens from within the same trilobite-bearing horizons.

175 11. Palpebral lobe shape: relative shape (outline) of the palpebral lobes,

when viewed dorsally.

States: (0) bean-shaped; (1) crescent-shaped.

Remarks: Alberti (1969, Fig. 18) depicted a difference in palpebral shape, when viewing the cranidium dorsally, for his two hypothesized subpopulations of

Cornuproetus. However, similar to the 3-3 distance and the shape of the glabella, two subpopulations of Cornuproetus are not supported by the shape of the palpebral lobes. This is the case for both all the examined Cornuproetus specimens and not just those examined by Alberti (1969). During our examination of this character, however, it was found to be conservative within morphologically similar specimens from the same trilobite-bearing horizons.

12. Palpebral rim structure: shape and sculpture of the lateral edge of the

palpebral lobe, surrounding the upper portion of the eye's lens field.

States'. (0) normal, no sculpture; (1) fine ridges; (2) granulose sculpture.

Remarks: Examination of morphologically similar taxa from the same trilobite-bearing horizons demonstrated that the palpebral rim sculpture, around the upper lens field of the eye, is quite conservative. Three typical sculpture types

176 (character states) were identified. Note, this is not the sculpture of the palpebral

lobe, but its lateral border (margin). The sculpture of the palpebral lobe was found

to be different, and more conservative, than this character.

13. Eye pad shape: relative shape (outline) and configuration of eye platform

when viewed dorsally (Figure 3-12).

States'. (0) absent; (1) posterior lobe only; (2) anterior and posterior lobes

present.

Remarks'. Examination of both Cornuproetus and Diademaproetus revealed that eye platform shape is variable between trilobite-bearing horizons and species. Morphologically similar specimens within trilobite-bearing horizons display similar eye platform shapes, with little variance.

14. Definition of occipital lobe furrows: size and depth of the occipital lobe

furrows when viewed dorsally.

States: (0) deep occipital lobe furrows; (1) shallow occipital lobe furrows.

177 Remarks: Size and depth of the occipital lobe furrows was found to be quite conservative within morphologically similar specimens from the individual trilobite-bearing horizons.

15. Occipital lobe median node size: presence and size of the median

occipital lobe node.

States: (0) absent; (1) small; (2) large.

Remarks: The presence and size of a median occipital lobe node is a typical proetid species level diagnostic character (Snajdrl980). Many species of both Cornuproetus and Diademaproetus are described as possessing a median occipital lobe node, therefore this character was used to determine if it has any weight on the relationship between the genera. Note, coding of this character does not include the height of the node, just its basal area. Small nodes have a basal area similar to or smaller than that of granules and tubercles, while large nodes have a basal area significantly larger than that of the granules or tubercles.

16. Border field sculpture: border sculpture.

States: (0) smooth or no sculpture; (1) granules; (2) fine ridges.

178 Remarks: Sculpture is used in many proetid diagnoses, and types of sculpture are considered important towards classification at multiple taxonomic levels. In regards to both Cornuproetus and Diademaproetus, sculpture has been used as a diagnostic feature for many species (Alberti 1969; Snajdrl980).

Applicable sculpture types used as character states for this character follow those outlined for proetids by Snajdr (1980). Care must be taken in determining details of sculpture, since specimens prepared using air abrasive techniques often have destroyed fine details of surface sculpture.

17. Genal field sculpture: sculpture of the genal field, the area proximal to

the cephalic border furrow and distal to the glabella and palpebral lobes.

States: (0) fine ridges; (1) granules; (2) isolated granules; (3) fine ridges and granules.

Remarks: Similar to the cephalic border field sculpture, the genal field sculpture of specimens appears to be quite conservative among morphologically similar specimens from individual trilobite-bearing horizons. The sculpture in this region is so similar that in some instances the same number, position, and pattern of individual granules is identical between different specimens.

18. Glabella sculpture: sculpture of the glabella.

179 States: (0) granules; (1) fine ridges; (2) fine ridges and granules.

Remarks: Glabellar sculpture is typically the most easily recognizable and coded of all the sculpture characters, commonly used as a species level diagnostic character for cornuproetids (Alberti 1969; Owens 1973; Snajdrl980). Similar to the genal field sculpture, this character typically displays very conservative sculpture patterns between morphologically similar specimens.

Thorax

19. Thoracic segments pleural region shape: shape of the thoracic segments

pleural region distal of the fulcrum (distal morphology of thoracic

segments).

States: (0) pointed; (1) rounded or blunt.

Remarks'. The general shape of thoracic segments pleural region, distal of the fiilcrum, was found to be quite conservative among morphologically similar specimens from the same trilobite-bearing horizons. Note, in order to be as conservative as possible, coding of this character was done using only the three most posterior thoracic segments. Segments closer to the cephalon, were avoided when coding this as they are not easily visible given the large genal spines. Also,

180 the tips of the most anterior thoracic segments are often narrower (exsag.) and

more pointed than those of more posterior thoracic segments.

20. Axis sculpture: sculpture of the thoracic segments axial region.

States: (0) granules; (1) fine ridges; (2) fine ridges and granules.

Remarks: Similar to the previously discussed characters involving sculpture, the placement and pattern of the axis sculpture was found to be highly conservative among morphologically similar specimens from the same trilobite- bearing horizons. The sculpture of the thoracic axial region was typically identical to that of the occipital lobe.

21. Pleural sculpture: sculpture of the thoracic segments' pleural region.

States'. (0) granules; (1) fine ridges; (2) fine ridges and granules.

Remarks'. Similar to the sculpture of the thoracic segments axial region, but on the pleural region.

Pygidium

181 22. Pygidial length vs. width: maximum length of the pygidium (Z) in

relation to the maximum width of the pygidium (W,, Figure 3-5).

States: (0) > 0.37; (1) < 0.37.

Remarks: Alberti (1969) and Chatterton et al. (2006) both diagnosed

Diademaproetus as having a short and broad pygidium. In neither publication,

however, is theer an explanation of what the qualitative terms short and broad refer to or are comparable to. As such, our quantitative examination of the character is an attempt to make this character more easily understandable and easily coded.

23. Pygidial shape: overall shape (outline) of the pygidium as defined by

Snajdr (1980) (Figure 3-13).

States: (0) segment of a circle; (1) rounded square; (2) rounded pentagon.

Remarks: Alberti (1969) diagnosed Diademaproetus on the presence of a rounded-square pygidium. Furthermore, Snajdr (1980) stated that the shape of the pygidium is an important characteristic of many proetid genera.

24. Rhachis length vs. width: length of the rhachis (Y) divided by its maximum

width (X, Figure 3-5).

182 States: (0) > 0.85; (1) <0.85.

Remarks: Snajdr (1980) discussed the size, shape, and configuration of the pygidial rhachis as all significant proetid characteristics at differing taxonomic levels. Examination of the rhachis (size) length vs. width for cornuproetids revealed that two populations (character states) are present for the examined taxa.

25. Pleural rib shape: shape of the two most anterior pleural ribs of the

pygidium (Figure 3-14).

States: (0) curved; (1) bent.

Remarks: Examination of morphologically similar specimens from individual trilobite-bearing horizons demonstrates that the shape and size of the most anterior pleural ribs of the pygidium are quite conservative, but typically vary between taxa.

26. Most anterior pleural rib reaches pygidial margin: this character

involves whether or not the most anterior (1st) pleural rib of the pygidium

reaches the pygidial border.

183 States: (0) 1st rib reaches pygidial border; (1) 1st rib does not reach the pygidial border, and does not cross the pygidial pleural furrow.

Remarks: In all collections of similar specimens from a particular locality, the specimens shared either state 0 or state 1 of this character and do not show the other state in the same collection

27. Presence of border notch: presence and shape of a posteromedian

pygidial notch or indent, at the most posterior portion of the pygidium

(Figure 3-15).

States: (0) none (rounded); (1) notched, indented; (2) flat (transverse).

Remarks: Alberti (1969) and Snajdr (1980) both refered to a pygidial notch

(indent) in the pygidial border, posterior of the rhachis. Further examination of morphologically similar specimens from individual trilobite-bearing horizons during this study found this characteristic to be quite frequent and stable among morphologically similar specimens. Given it's previous discussion as a species level diagnostic character, and its presence in many of the examined taxa, it was included in the phylogenetic analysis.

184 28. Presence of post rhachial bump: presence or absence of a post rhachial

bump, posterior of the rhachis.

States: (0) present; (1) absent.

Remarks-. Presence of a post rhachial bump, behind the rhachis on the pygidium, is typical of many proetids, varying in shape and size at the species level (Snajdr 1980).

29. Presence of border rims: presence or absence of a border rim around the

pygidium, distal of the pygidial border furrow.

States: (0) present; (1) absent.

Remarks: Much like the cephalon, the presence of a border rim around the pygidium occurs conservatively within morphologically similar specimens from the same trilobite-bearing horizons. However, unlike the presence of multiple border rims on the cephalon, only one border rim is present on the pygidial border.

30. Pleural region sculpture: sculpture on the pleural region of the pygidium.

States: (0) granules; (1) fine ridges; (2) fine ridges and granules.

185 Remarks'. Sculpture types synonymous with those previously discussed for

other sclerites and regions of the exoskeleton. Determination of this character

requires well preserved and well prepared specimens.

31. Rhachis sculpture: sculpture of the rhachis on the pygidium.

States: (0) granules; (1) fine ridges; (2) fine ridges and granules; (3) smooth (no sculpture).

Remarks: Sculpture types synonymous with those previously discussed for other sclerites and regions of the exoskeleton. Again, as for character 30, determination of this character requires well preserved and well prepared specimens.

Characters excluded from the analysis

Although 31 characters were found to be useful for determining the relationship between Cornuproetus and Diademaproetus, numerous other characters had to be excluded as they either lacked variance or played no significant role in understanding the relationships of the examined taxa (were not cladistically informative). Following this, is a brief discussion on a character by character basis of the characters that were omitted from the analysis as they

186 proved unfruitful towards understanding the relationships among the examined

taxa.

Rhachis configuration and number of thoracic segments. Snajdr (1980)

discussed and promoted the number of axial segments within the pygidial rhachis

as being a very important evolutionary and diagnostic character for many proetids.

As additional segments are released into the thorax from the pygidium during a

trilobites' meraspid (juvenile, non-mature) period, their number should stabilize

when the trilobite reaches its holaspid (adult, mature) period (Chatterton & Speyer

1997). Therefore, when examining holaspid specimens, reduced or additional segments within a specimen's thorax or pygidium typically signifies a significant

change in the ontogeny. Therefore the specimen most likely belongs to a different

taxon compared to the specimens who all have the same number of segments in

their pygidium and thorax. It should also be noted that while many Cambrian trilobites do show plasticity in the number of segments, proetids typically do not, and are considered more conservative in regards to their segment numbers

(Owens 1973; Chatterton & Speyer 1997; Hughes et al. 2006).

Within the examined cornuproetids, both the number of thoracic segments

(10) and the configuration of the pygidial rhachis (four segments and one terminal piece) are quite conservative. All examined cornuproetids of this study possessed ten thoracic segments, preventing the character from being informative at a low taxonomic level of study. Variance is found however, within the rhachis configuration of Cornuproetusperaticus Owens, 1973, which has a rhachial

187 configuration of five and one instead of the much more common four and one.

Originally, following Owens (1973) description of C. peraticus, Snajdr (1980) thought the species to be a member of lnterproetus. However, this is likely not the case, for the majority of lnterproetus species, including the holotype, have a rhachis configuration of four and one. Unfortunately, given that C. peraticus is the only species to have a different rhachis configuration compared to other cornuproetids, the character is labelled as "parsimonious uninformative" by the analysis and rendered useless (an autapomorphy for the species). Despite the conclusions of the phylogenetic analysis, the rhachial configuration is still useful for diagnosing species from one another. Indicating that at least one Cornuproetus species changed the number of thoracic segments that has been assigned to it.

Number and shape of glabellar furrows and depth of interpleural furrows. Alberti (1969) and Snajdr (1980) discussed the number and shape of glabellar furrows and the depth of thoracic and pygidial interpleural furrows as being diagnostic at higher taxonomic levels, and therefore being evolutionarily significant characters. However, given the extremely low taxonomic level of this study, no variance within either of these characters was found. As a result the characters were culled from this analysis. It should be noted, that if testing higher level phylogenetic relationships these characters should be considered in the phylogenetic analysis if they show variance between taxonomic units.

Results

188 Heuristic parsimony, branch and bound, and exhaustive analyses, all with equally weighted and unordered characters, all yielded the same single tree with a total length of 102 steps, ensemble consistency index (C.I.) of 0.4902, ensemble retention index (R.I.) of 0.5098, and rescaled consistency index (R.C.) of 0.6667

(see Figure 3-16, for the cladogram, with Bremer support values; see Figure 3-17, for the same cladogram indicating character state changes). Results of the exhaustive search returned a skewed single sided tree distribution (landscape), with one additional step (103) producing 164 trees and two additional steps (104) producing 1464 trees. Bootstrap and Jackknife support analyses, both using 2500 replicates, resulted in similar consensus trees, with all clades having a Bremer support of greater than one, and the Pontoproetus clades, being present. All other areas with a Bremer support of one collapsed into polytomies in these resampling analyses.

Following the original phylogenetic analysis, insertion of Dalejeproetus dalejensis (Pribyl, 1971), the type species of Dalejeproetus into the analysis resulted in the the same overall phylogeny being produced, albeit with weaker

Bremer support values and indices and the tree having 107 steps instead of 102.

Dalejeproetus dalejensis plotted out between Cornuproetus cornutus cornutus and

Diademaproetus rudimentus new species in all instances.

Similarly to Dalejeproetus, Montanproetus capuvena Basse, 2010, the type species of Montanproetus, and Montanproetus gordios Basse, 2010, were both added to a third round of analyses with all the originally examined taxa, not

189 including Dalejeproetus. Inclusion of these two species resulted in an increased

number of most parsimonious trees, re-shaping of the tree distribution from a

skewed to proper bell curve, and loss of all Bremer support values above one, compared to the results of the primary analysis. Within the multiple most parsimonious trees produced by the addition of Montanproetus, only the

Montanproetus species themselves shifted throughout the trees. The original taxa from the primary analysis continued to hold the same positions in relation to one another in all most parsimonious trees.

Discussion and conclusions

Given the problem of diagnosing Cornuproetus and Diademaproetus, so as to distinguish them from one another, and the effects of character exhaustion, the production of only a single most parsimonious tree by all three (heuristic parsimony, branch and bound, and subset exhaustive) analyses, a Bremer support value above one at the Cornuproetus-Diademaproetus boundary, and a skewed single sided tree distribution (landscape) during the subset exhaustive analysis, suggests that the produced phylogeny (Figure 3-16 & 3-17) is a useful hypothesis of relationships among the examined taxa. Furthermore, production of identical, albeit not as well supported, consensus trees by character resampling (Bootstrap and Jackknife) analyses further strengthens the results of the exhaustive analysis and the characters used. As a result, this robust phylogeny allows for a better understanding of Cornuproetus and Diademaproetus, and their relationship.

190 Being conservative, generic level taxonomic units were identified using clades with high Bremer support values. This approach was employed as the best method to replicate taxonomic ranking, as genera should be morphologically distinct, therefore well supported and identifiable from one another in a phylogeny than species. As a result of this methodology, the Cornuproetus-Diademaproetus boundary was drawn between Cornuproetus cornutus cornutus and the new species Diademaproetus rudimentus, where a Bremer support value of two exists and two synapomorphic character states (presence of a wide (W) and short (Z) pygidium and shorter (Y) and wider (X, Figure 3-5) pygidial rhachis) are present.

As for additional synapomorphies within the examined taxa, the bulk of characters represent either reversals higher up, or homoplastic occurrences lower down, in the phylogeny (Figure 3-17), therefore synapomorphies are rare. The most notable synapomorphy present, besides those at the Cornuproetus-

Diademaproetus boundary, is the truncated anterior portion of the cephalic border furrow (character 7, state 0) which is exclusive to the most derived species and the new genus Pontoproetus. Other synapomorphic characters include the flat cephalic border (character 5, state 0) of the more derived diademaproetids and

Pontoproetus, and the fine ridge sculpture of the thoracic axis between the new species Diademaproetus langus and the pontoproetids. Additionally, numerous autapomorphies also exist within the phylogeny, as many species contain unique characteristics not found in others.

191 Besides synapomorphic characters, the size and shape of the anteromedian cephalic projection appears to be less plastic than Chatterton et al. (2006) suggested and distinctly increases in size and decreases in width from primitive to more derived diademaproetids that exhibit it. For example, the new species

Diademaproetus issoumourensis displays a very weak anteromedian cephalic projection, which appears as the anterior portion of the cephalic border coming to a rounded point, compared to Diademaproetus preacursor which has a short and wide projection, and finally followed by the new species Diademaproetus langus which has a very long and narrow projection.

In order to test the validity of the division between Cornuproetus and

Diademaproetus, Dalejeproetus dalejensis, a hypothesized evolutionary intermediate between the two examined genera (Snajdrl980), was coded and inserted into the analysis. While addition of the species did increase the number of steps within the tree (107 instead of 102) and reduce the tree's Bremer support, the tree topology remained identical and D. dalejensis was positioned exactly as hypothesized as an evolutionary intermediate, between C. cornutus cornutus and

Diademaproetus rudimentus. This placement of D. dalejensis further strengthens our previously defined Cornuproetus-Diademaproetus boundary, as a wide (W) and narrow (Z, Figure 3-5) pygidium remained a synapomorphic character of

Diademaproetus. Closer examination of the Dalejeproetus-Diademaproetus relationship, which is outside of the scope of this study, needs to be accomplished before a differential diagnosis between the two genera and the phylogenetic

192 importance of the length and width of the pygidial rhachis in this situation can be determined.

Given the extremely similar diagnoses of Montanproetus and Taciproetus

(Basse 2010) to Cornuproetus (Alberti 1969), inclusion of some of their species into the analysis was used test their validity as genera. Taciproetus adeps Basse,

2010 was included throughout all phylogenetic analyses as it was relatively complete (greater than 50% coded) and did not severely degrade tree support.

Basse's (2010) diagnosis of Taciproetus as being a cornuproetid was based on it having a strongly convex glabella; subquadratic to weakly violin-shaped glabella; ledge like anterior border of the cephalon, with border furrow pushed against the anterior limit of glabella; pygidial flanks with weak relief and furrows flexing backwards; and a pygidial border that is marginally bent downwards. Within our analysis, T. adeps plots out within a subclade of Cornuproetus, between

Cornuproetus peraticus Owens, 1973 and Cornuproetus chlupaci Alberti, 1967.

Furthermore, T. adeps is diagnosed in our analysis as being a Cornuproetus with an intermediately positioned cephalic suture angle delta, a single cephalic border rim, normal palpebral lobe structure, and small occipital node. None of the diagnostic features of Taciproetus, as outlined by Basse (2010), were found to be either statistically significant when compared with Cornuproetus, reported as parsimoniously uninformative characters, or useful in diagnosing Taciproetus apart from Cornuproetus. Given these short comings of the diagnostic features of

Taciproetus, the fact that many of its assigned species were originally considered

i

193 members of Cornuproetus by Alberti (1969), and the very similar morphology of

Taciproetus to Cornuproetus, Taciproetus is considered herein a junior synonym of Cornuproetus.

As for Montanproetus, inclusion of its species into the analysis was problematic. First, many Montanproetus species are highly incomplete (less than

50% coded), being diagnosed using only cranidia or partial cephala and associated pygidia. Second, inclusion of its species into the analysis severely reduced Bremer support values, causing the analysis to produce many, instead of just one, most parsimonious trees. Within the produced trees, the only difference in tree topology amongst themselves was the placement of the Montanproetus species between the

Interproetus and Diademaproetus clades. Its species were essentially not phylogenetically different from Cornuproetus. Furthermore, the tree topology of both the Interproetus and Diademaproetus clades, when Montanproetus was included, was identical to when it was omitted. This problematic position of

Montanproetus is likely a result of two issues, many of its species being relatively incomplete and the fact that it is not morphologically distinct from Cornuproetus.

The latter of these issues is evident given the diagnosis of the genus by Basse

(2010), which includes: convex anterior border of cranidium, presence of two furrows anterior of the glabella in at least one postlarval phase, violin-shaped glabella, large palpebral lobes, wide (W) and short (Z, Figure 3-5) pygidium, and deep interpleural pygidial furrows. First, presence of features in single post-larval phases is not a useful diagnostic character as it is unclear which post-larval phase

194 of ontogeny is being refered to, and if only phases not depicting the proposed character are available, how can one accurately diagnose the species. Characters such as these, with ontogenetic variability, are not suitable diagnostic characters for taxa and should be avoided (Cronier et al. 2011). Secondly, none of the proposed diagnostic characters by Basse (2010) for Montanproetus are either statistically significant or diagnostic for the genus's separation from

Cornuproetus . Therefore, given the problematic nature of Montanproetus, the characters used to diagnose it, and the fact that many of its species were once classified as Cornuproetus (Alberti 1969), herein Montanproetus is considered a junior synonym of, and its species assigned to, Cornuproetus.

In large part, the results of character exhaustion and morphologically similar species on a phylogenetic analysis can be overcome when statistically tested characters and relatively complete OTUs are examined. In the case of

Cornuproetus and Diademaproetus, use of these methods in conjunction with specimen based OTUs produce a well supported phylogeny that not only clarifies the diagnoses of the two similar genera, but also withstands the inclusion of additional taxa thought to be of significance. This result supports the hypothesis that the use of properly assessed quantitative characters and cladistic analysis can produce reliable results and useful phylogenies for even morphologically similar taxa. Furthermore, such quantitative methodology and phylogenetic testing of new species needs to become common practice when examining morphologically similar taxa. Without quantification of characters, subjective terminology leads to

195 improper characters coding and possibly false relationships. Ultimately, these things can lead to over-splitting of taxa, which as a result, hampers the assignment of new species and material to the proper taxon. Without a unified approach towards examining these taxa and their relationships, understanding of their evolution and that of trilobites as a whole will not be easily obtained.

Systematic palaeontology

Order Proetida Fortey & Owens, 1975

Subfamily Cornuproetinae Richter & Richter 1949

Genus Cornuproetus Richter & Richter, 1919

Junior Synonyms Taciproetus Basse, 2010 & Montanproetus Basse, 2010

Type species - Cornuproetus cornutus cornutus (Goldfuss, 1843)

Other included species - Cornuproetus complanatus (Barrande, 1846);

Cornuproetus curtus (Barrande, 1852); Cornuproetus pictus (Giebel, 1858);

Cornuproetus adeps Kegel, 1926; Cornuproetus pernix pernix Kegel, 1926;

Cornuproetus taciturnus Kegel, 1926; Cornuproetus midas midas (Erben, 1952);

Cornuproetus exanthemoides exanthemoides (Erben, 1952) Cornuproetus chouberti chouberti Alberti, 1964; Cornuproetus fauremuretae Alberti, 1964;

Cornuproetus chlupaci Alberti, 1967; Cornuproetus hercules Alberti, 1967;

Cornuproetus mauretanicus Alberti, 1967; Cornuproetus rudrichteri Alberti 1967;

196 Cornuproetus steinbergicus Alberti, 1967; Cornuproetus walliseri Alberti, 1967;

Cornuproetus chouberti acrodactylium Hass, 1968; Cornuproetus exanthemoides subsolanus Hass, 1968; Cornuproetus comutus marrakechensis Alberti, 1969;

Cornuproetus pernix boutsharafinensis Alberti, 1969; Cornuproetus midas amlanensis Alberti, 1969; Cornuproetus tindoufanus tindoufanus Alberti, 1969;

Cornuproetus infans Alberti, 1969; Cornuproetus maidericus Alberti, 1969;

Cornuproetus mendax Alberti, 1969; Cornuproetus pribyli Alberti, 1969;

Cornuproetusperaticus Owens, 1973; Cornuproetus tindoufanus fornitox Snajdr,

1980; Cornuproetus augur Snajdr, 1980; Cornuproetus trambus Snajdr, 1980;

Cornuproetus comutus archaeocornutus Snajdr, 1980; Cornuproetus capuvena

Basse, 2010; Cornuproetus gordios Basse, 2010; Cornuproetus montanus Basse,

2010.

Diagnosis - Cornuproetid with short preglabellar field; inflated or "cushion-like" cephalic border; narrow positioning of cephalic suture angles gamma (y) and delta

(8); sculpture of fine granules throughout genal field; comparatively short (for the subfamily) genal spine which terminates alongside thoracic region; long (Z) and narrow (Z) pygidium (length (Z) is greater than 0.37x the maximum width (W); long (Y) and narrow (X) pygidial rhachis (length (Y) is greater than 0.85x the maximum width (X, Figure 3-5); and pygidial rhachis and pleural region both display sculpture of granules, primarily along the posterior limit of rhachial segments and pleural ribs.

197 Remarks - Cornuproetus is the most diverse and stratigraphically abundant of all cornuproetid genera, with over 30 species being known from the Late Silurian to the Middle Devonian. With such a large diversity and long stratigraphic range we still considered it likely, as did Alberti (1969) that Cornuproetus is a true evolutionary rootstock for many of the other, and much less diverse, cornuproetid genera. However, exactly how and when all of these other genera evolved from

Cornuproetus is still unclear and in need of further elucidation.

Based on the similarity of diagnosis between Cornuproetus, Taciproetus, and Montanproetus, and the unsupported statistical and phylogenetic differentiation of the latter two genera from Cornuproetus, we consider both

Taciproetus and Montanproetus as junior synonyms of Cornuproetus. Failure to asses statistically and phylogenetically both Taciproetus and Montanproetus, had led to over-splitting of Cornuproetus and ultimately the proposal of problematic genera that made designation of new species to the proper genus extremely difficult. Since many species of both Taciproetus and Montanproetus were originally described as Cornuproetus, and they all statistically and phylogenetically plotted out within Cornuproetus, both genera are synonymized and considered junior synonyms of Cornuproetus.

Within Cornuproetus itself, Alberti (1969) diagnosed two subpopulations based upon supposed differences in their 5-5 distance, glabellar shape, and palpeberal lobe shape. Upon quantitative examination of the specimens examined by Alberti (1969) and then an entire sample of all current Cornuproetus species,

198 no statistically significant or intuitive subpopulations of Cornuproetus were

determined. We therefore regard these subpopulations as misleading, and

therefore his division of Cornuproetus into two separate evolutionary lineages as

difficult to justify. Most likely Cornuproetus was a diverse genus, some of whose

many different species eventually gave rise to different comuproetid genera. This

is supported by Cornuproetus being represented as paraphyletic, rather than

monophyletic, in our proposed phylogeny. In essence, a central group of

Cornuproetus species existed throughout the stratigraphic range of the genus, and

as new morphological variance progressed in some species new genera were

formed. For example, Cornuproetus cornutus cornutus is distinctly more similar

to Diademaproetus than other examined Cornuproetus species (Figures 3-16 &

3-17), but not similar enough to be diagnosed as Diademaproetus, and therefore is

a possible Cornuproetus ancestor of Diademaproetus.

Genus Diademaproetus Alberti, 1964

T^pe species - Diademaproetus holzapfeli (Novak, 1890)

Other included species - Diademaproetus praecursor Alberti, 1969;

Diademaproetus antatlasius Alberti, 1969; Diademaproetus menzeni Kowalski,

1975; Diademaproetus morenaica Hammann, 1976; Diademaproetus mohamedi

Chatterton et al. 2006; Diademaproetus rudimentus sp. nov.; Diademaproetus

199 issoumourensis sp. nov.; Diademaproetus corrugatus sp. nov.; Diademaproetus

auxiliarus sp. nov.; Diademaproetus langus sp. nov..

Diagnosis - Long preglabellar area (greater than or equal to 0.28x length of

glabella and occipital lobe (A2 + A3)); cephalic border sculpture of fine ridges;

bean-shaped palpebral lobes; long genal spines that extend past thorax and

terminate parallel to pygidium; wider (X) and shorter (Y) pygidial rhachis (length

(Y) is less than or equal to 0.85x the maximum width (X, Figure 3-5)).

Remarks - Although widely agreed that Diademaproetus was a descendant of

Cornuproetus, the lack of agreement on the diagnoses of these genera made

designation of new material to the proper genus difficult. Alberti (1969) originally

diagnosed Diademaproetus as a cornuproetid genus with an anteromedian

cephalic projection, anteromedian cephalic projection depression, anterior

cephalic border furrow contacting the anterior limit of glabella, glabella

subquadratic in shape, large eyes, extremely long genal spines that extend past

the thorax and terminate alongside the pygidium, and a short (Z) and wide (W,

Figure 3-5) pygidium. Chatterton et al. (2006) however, considered that the

presence of an anteromedian cephalic projection was not necessarily diagnostic of

the genus based on the fact that occurrence was variable among species of this genus. Chatterton et al. (2006) also suggested that the presence of a flat and broad cephalic border, with epiborder furrows, is distinctive of the genus.

In an attempt to solve the diagnostic confusion surrounding

Diademaproetus, all previously used diagnostic characters of the genus were

200 coded and added to the phylogenetic analysis involving both Cornuproetus and

Diademaproetus. From the resulting phylogeny (Figures 3-16 & 3-17) the

synapomorphic quality of a long preglabellar field (character 4, state 1) and a

short (Y) and wide (X, Figure 3-5) pygidial rhachis (character 24, state 1) in the

more derived cornuproetids indicated that these characters are most diagnostic for

Diademaproetus. The presence of a long genal spine is also likely a diagnostic

character of Diademaproetus, however with some specimens' genal spine length

not being known (taxa are represented by incomplete and unarticulated

specimens), it cannot be confirmed whether this is true for all Diademaproetus

species. As for the remaining 'diagnostic' characters stated by Alberti (1969) and

Chatterton et al. (2006), many of them are not consistent for all species of

Diademaproetus and therefore not crucial diagnostic characters.

In addition to producing an acceptable diagnosis for Diademaproetus, this study also greatly increases the diversity the genus. Originally, Alberti (1969) only described three species of Diademaproetus, with three more additional species

being described later (Kowalski 1975; Hammann 1976; Chatterton et al 2006).

Such a large increase in Moroccan diversity of Diademaproetus possibly signifies it's Devonian area of origin. In total, all but two species of Diademaproetus are found in Morocco. Furthermore, in accordance with the phylogenetic analysis, the most primitive diademaproetid (D. rudimentus) and closely related cornuproetids

{Cornuproetus cornutus cornutus) are all found in similar aged, Middle Devonian,

Moroccan strata. Given the Devonian palaeogeographic proximity of Morocco to

201 the other European areas where Diademaproetus is found (all within the Devonian

Rheic Ocean), the high density of Moroccan Diademaproetus species, and the

similar stratigraphic age of primitive Moroccan diademaproetids and derived

cornuproetids, it is possible that Emsian aged Morocco was the palaeogeographic

epicenter of Diademaproetus evolution from Cornuproetus. Then, following the

evolution of Diademaproetus in Morocco, diademaproetid species spread to other

areas of the Middle Devonian Rheic Ocean, such as Eastern and Central Europe.

Such an evolutionary theory of Diademaproetus, from Cornuproetus, is in

agreement with the works of Alberti (1969, Fig. 17), therefore our study has been

able to identify the Emsian aged diademaproetids missing from the equation.

Diademaproetus praecursor Alberti, 1969

Plate 3-1, figs. 1-6; plate 3-2, figs. 1-7; plate 3-3, figs. 2 & 4.

Diagnosis - See Chatterton et al. (2006)

Material - Articulated specimens ( UA13692, UA13693, UA13694, UA13695,

UA13696, UA13697) and isolated hypostome (UA13698).

Occurrence - Found in multiple horizons, localities and basins in southern

Morocco. The following is a basin by basin, locality by locality, breakdown of the known occurrences of D. praecursor.

Tindouf Basin. Zguilma. All three trilobite-bearing horizons of the

Timrhanrhart Formation, ZGEEl, ZGEE2, and ZGEE3. Horizons ZGEEl and

202 ZGEE2 are late Emsian in age, while horizon ZGEE3 is stratigraphically higher than horizons ZGEE1 and ZGEE2 and Eifelian in age (Chatterton et al. 2006;

Gibb & Chatterton 2007, 2010),

Ma'der Basin. Jbel Oufatene. Found in the 'Diademaproetus' and

'Psychopyge' horizons west of the village of Lahfira. The 'Diademaproetus horizon' is part of the El Otfal Formation and Eifelian (Middle Devonian) in age

(Morzadec 2001; Gibb & Chatterton 2010), while the 'Psychopyge horizon' is part of the Tazoulai't Formation and Late Emsian in age (Gibb & Chatterton 2010).

Jbel Issoumour. Found in the 'Psychopyge,''Kayserops,''Lobopyge,' and

'Paralejurus' horizons on the western face of Jbel Issoumour. Both the

'Psychopyge' and ''Kayserops'' horizons are part of the Er Remlia Formation

(Gibb & Chatterton 2010), while the 'Lobopyge' and 'Paralejurus' horizons are stratigraphically higher in section and part of the El Otfal Formation (Gibb &

Chatterton 2010).

Tafilalt Basin. Talawarite. Found in the 'Gerastos horizon,' which is part of the Amerboh Group and late Emsian in age (Gibb & Chatterton 2010). Also found in surface collections of the area surrounding the locality.

Taouz. Found in surface collections surrounding the locality whose strata are of Emsian age (Gibb & Chatterton 2010).

Description - See Chatterton et al (2006)

Remarks - Diademaproetus praecursor is the most stratigraphically and palaeogeographically wide ranging species of Diademaproetus. In southern

203 Morocco alone, D. praecursor is found within all three examined basins (Tindouf,

Ma'der, and Tafilalt) and in both Emsian (late) and Eifelian (early) aged strata.

This wide stratigraphic and palaeogeographic range of the species, which no other

Moroccan diademaproetid species exhibits, signifies D. precursor as the most stable morphotype, and thus possibly root stock species, of Diademaproetus. This

is further supported by our phylogenetic analysis, in which D. praecursor is

located exactly between the more primitive (Emsian aged) and more derived

(Eifelian aged) Moroccan diademaproetids (Figure 3-16). Essentially, D.

praecursor bridges the gap between the Emsian and Eifelian species of

Diademaproetus.

Diademaproetus praecursor differs from the type species of

Diademaproetus, D. holzapfeli by having a slightly smaller (less protuberant) anteromedian cephalic projection with a smaller and shallower associated furrow; a more narrow positioned cephalic suture angle beta (P); glabellar and genal field sculpture of granules instead of fine ridges and granules; glabellar sculpture of granules also extends farther forward on the glabella; a flat, instead of slightly notched/indented, posterior border of the pygidium; and pygidial rhachis sculpture of fine ridges and granules instead of simply granules.

Diademaproetus mohamedi Chatterton et ah, 2006

Plate 3-3, figs. 1, 3, 5-6 & plate 3-4, figs. 1-7.

Diagnosis - See Chatterton et al. (2006)

204 Material - Holotype UA13383 (complete articulated specimen except for damaged pygidial pleural region). Paratype UA13699 (complete articulated specimen).

Occurrence - Tindouf Basin. From the trilobite-bearing horizon 'ZGEE3' at

Zguilma. Strata are Eifelian in age and part of the Timrhanrhart Formation

(Chatterton et al. 2006; Gibb & Chatterton 2007,2010). Tafialt Basin. From the

'Harpes/Thysanopeltis horizon' at bou Dib. Strata are part of the El Otfal

Formation, which is Eifelian in age (Gibb & Chatterton 2007, 2010).

Description - See Chatterton et al. (2006).

Remarks - Diademaproetus mohamedi is an extremely rare Diademaproetus species, with only two specimens being known, coming from the ZGEE3 trilobite- bearing horizon of Zguilma, and one specimen from bou Dib.

Interestingly, Diademaproetus mohamedi from Zguilma is one of only two

Diademaproetus species to lack an anteromedian cephalic projection completely, the anterior cephalic border being perfectly rounded and without any form of tapering into a point about the cephalic midline. Morphological similarity of this species with D. praecursor promoted the diagnosis of the species as

Diademaproetus rather than Cornuproetus, ultimately leading Chatterton et al.

(2006) to conclude that the presence of an anteromedian cephalic projection was not a diagnostic requirement for Diademaproetus. Our proposed phylogeny of

Cornuproetus and Diademaproetus supports the conclusions of Chatterton et al.

(2006), placing D. mohamedi besides D. praecursor topologically within the

205 Diademproetus group and not promoting the presence of an anteromedian

cephalic projection as a necessary diagnostic character for Diademaproetus.

Furthermore, the presence of a very small (pointed) anteromedian cephalic

projection in the bou D!b specimens of D. mohamedi suggests that the presence of

anteromedian projection can be variable for this species.

Diademaproetus mohamedi differs from D. holzapfeli in lacking an

anteromedian cephalic projection and its pygidial rhachis having sculpture of fine

ridges and granules, concentrated along the posterior limit rhachial segments,

instead of just granules. D. mohamedi differs from D. praecursor in lacking or

having a very reduced anteromedian cephalic projection; having a more widely positioned cephalic suture angle beta (P); and glabellar and genal field sculpture of fine ridges and granules instead of just granules.

Diademaproetus rudimentus new species

Plate 3-5, figs. 1-7 & plate 3-6, figs. 1-6.

Derivation of name - Named in reference to the species' primitive character suite. Since the species is thought to be one of the earliest species of

Diademaproetus, we considered its features rudimentary compared to those of more derived species the genus.

Diagnosis - Diademaproetus with sculpture of large isolated granules on genal field, near junction of lateral and posterior border furrows, at base of genal spine;

206 glabellar sculpture of fine ridges and granules; and posterior margin of thoracic pleurae with sculpture of granules.

Material - Holotype UA13686 (complete and fully articulated exoskeleton) and paratypes UA13687, UA13688, UA13704, and UA13705.

Occurrence - Ma'der Basin. From Taharajat, south of Jbel Oufatene, and El

Achana. Strata at Taharajat are late Emsian in age (Gibb & Chatterton 2010), while at El Achana they are believed to be Pragian in age.

Description - Cephalon. "D" shaped and strongly convex. Sagittal length (A,

Figure 3-5) 0.38x length of entire specimen. Cephalic border inflated, appearing convex or cushion-like when viewed laterally. Anterior and lateral portions of cephalic border rimmed by three fine parallel ridges (border rims), between cephalic border and cephalic border furrow. Cephalic border furrow broad and moderately deep anterior of glabella, narrowing as it approaches librigena and genal angle. Cephalic border furrow deepest at base of genal spine. Posterior cephalic border furrow moderately deep, running laterally from axial furrow opposite occipital ring to junction with lateral cephalic border furrow. Posterior border furrow longest at its midwidth, anterior of cephalic suture angle omega

(ft).

Cranidium. Preglabellar area (A2 + A3) slightly inflated, 0.22x cephalon length (A, Figure 3-5). Sagittal length of glabella (Ai + A4) 0.78x length of cephalon (A, Figure 3-5). Length of anterior portion of glabella (Ai) is 0.56x cephalon length (A, Figure 3-5). P - P distance 0.67x length of cephalon (A,

207 Figure 3-5). y - ydistance 0.58x length of cephalon (A, Figure 3-5). 5-8 distance

0.87x length of cephalon (A, Figure 3-5). Anterior portion of glabella weakly- violin shaped, moderately convex and with a sculpture of sparse fine ridges and granules, granules evenly positioned throughout, while fine ridges are rare and most prominent on anterior portion of glabella. When viewed laterally, anterior portion of glabella weakly convex and steeply angled downwards (ventrally) towards cephalic border. Posterior portion of glabella less steeply angled than anterior portion, with highest dorsal point of glabella being its posterior margin along cephalic midline. In dorsal view, anterior portion of glabella slightly constricted posteromedially of cephalic suture angle gamma (y). Glabella contains

4 pairs of glabellar furrows. Furrows SI and S3 are largest and located medial to cephalic suture angles delta (5) and gamma (y) respectively, appearing linear in shape and curving posteromedially towards cephalic midline. Furrows S2 and S4 are smaller and semi-circular in shape, located anteromedially of SI and S3 respectively. Palpebral lobes are bean-shaped and slightly sloped downwards medially, towards glabella. Palpebral lobe lateral border is rimmed with sculpture of fine granules. Shallow and narrow palpebral furrow is present. Occipital furrows (SO) shallow. Occipital lobe (A4) 0.22x length of cephalon (A, Figure

3-5) and transversely wider than widest portion of glabella. Occipital ring with sculpture of granules.

Librigena. Eyes large and strongly convex in both sagital and vertical planes. Both anterior and posterior portions of eye platform present. Anterior

208 portion of eye platform larger than posterior portion and rounder in shape. Genal field with sculpture of fine ridges and granules positioned along lateral genal field border. Long, medially curved genal spines longer than rest of cephalon (A,

Figure 3-5) and terminate lateral of pygidium. Lateral border furrow and posterior border furrows both merge at genal angle. Medial aspect of genal spine slightly more inflated than lateral aspect. Genal spine furrow widest and deepest at genal angle, narrowing and shallowing posteriorly.

Hypostome. Length (HL) 1.04x width (HWi, Figure 3-5). Central hypostomal body slightly longer than wide, with anteromedial convex bulge

(rhynchos) and rounded posterior. Anterior border raised (ventrally in life position) above lateral and posterior borders. Central hypostomal body separated from anterior border by shallow anterior border furrow. Lateral borders narrow, with moderately deep and short lateral border furrows anterior of slightly swollen maculae. Posterior border semi-circular in shape posteriorly, with short pair of posterolateral marginal spines. Both lateral and posterior borders slightly convex.

Anterior wings are steeply downsloped (dorsally) laterally and bluntly trapizoidal.

Entire hypostome covered by sculpture of dense anastomosing terrace ridges.

Terrace ridges on lateral and posterior borders run parallel to hypostomal border.

Thorax consists of 10 segments, representing 0.49x sagittal length (when articulated) and 0.78x maximum width (tr.) of entire specimen. Axial rings moderately arched and bordered laterally by moderately deep axial furrows.

Maximum width of axis (tr.) approximately 0.47x maximum width (tr.) of thorax.

209 Axial rings increase in width from most anterior segment to third segment. Axial rings posterior of third segment decrease in width from anterior to posterior, with most posterior (10th) axial ring being 0.55x width (tr.) of third axial ring. Axial ring sculpture consists of both fine terrace ridges and granules, with granules clustered on posterior portion of axial rings. Higher amounts of fine ridge sculpture are present on more posterior axial segments, closer to axial furrows.

Most anterior axial segments have largest concentration of granules, with little or no fine ridge scultpure. Pleurae incised with moderately deep pleural furrows, running laterally from axial furrow. Medial anterior and posterior margins of pleurae transverse, with more lateral regions curved posterolaterally to within approximately 75° of sagittal midline. Lateral portion of pleural terminate in distinct point. Sculpture of pleurae consists of fine granules, denser along posterior border of pleural regions.

Pygidium. 0.13x total length (sag.) and 0.37x maximum width (tr.) of complete specimen. Pygidium shaped as segment of circle. Proximal portion of pleural anterior margin transverse. Lateral of fulcrum, pleural margin angled posterolaterally at roughly 40° from transverse direction. Shallow and broad border furrow encases lateral and posterior pygidial border. Posterior border, rounded in shape. Rhachis moderately convex and cone-shaped, tapering towards sagittal pygidial midline from anterior to posterior. Rhachis length (Y) and width

(X) 0.68x and 0.36x pygidial length (Z) and width (W, Figure 3-5) respectively.

Rhachis composed of 4 rhachial rings and terminal piece. Rhachial ring furrows

210 shallow posteriorly, with more anterior rhachial ring furrows distinctly deeper.

Small post rhachial bump present. Three pairs of curved pleural ribs and two pairs of moderately deep pleural furrows. Pleural ribs curved in shape, decreasing in size from anterior to posterior. First (most anterior) pleural rib longest, extending to pygidial margin. Pleural furrows are moderately deep, shallowing distally to pygidial border. Both rhachis and pleural region with sculpture of fine granules located on posterior borders of rhachial rings and pleural ribs.

Remarks - Diademaproetus rudimentus is common at Taharajat, with multiple

(more the 10) specimens in our collection. Specimens of D. rudimentus are rare at

El Achana, with only two poorly preserved and prepared (chipped) specimens being known. Quantitatively, the Taharajat and El Achana specimens are very similar, however, due to the poor quality of preservation and preparation of the El

Achana specimens, some qualitative characters (e.g. sculpture) cannot be used to compare them. Therefore, given that the specimens are proportionally quite similar, and other characters are unknown, we treat them as the same species herein.

In conjunction with the Taharajat specimens, multiple hypostomes, a typically rare sclerite, of Diademaproetus rudimentus are also known. Specimen

(UA13686) possesses its hypostome in life postion. Phylogenetically,

Diademaproetus rudimentus is the most primitive Diademaproetus known from

Morocco, topologically situated between Cornuproetus cornutus cornutus and the remaining diademaproetids.

211 Diademaproetus rudimentus differs from Cornuproetus cornutus cornutus in having a longer preglabellar field (A2 + A3); cephalic border with sculpture of fine ridges instead of fine granules; genal field sculpture of granules clustered around the lateral genal field border instead of being evenly positioned throughout the entire genal field; subcylindrical shaped glabella with additional fine ridge sculpture instead of a strongly violin shaped glabella with only granulose sculpture; shallow rather than deep occipital lobe furrows; lacking a distinct occipital median node; slightly wider pygidial rhachis (X, Figure 3-5); posterior pygidial margin that is rounded instead of flat; and lacking a pygidial border rim.

Compared to Diademaproetus praecursor, D. rudimentus differs by having a wider positioned cephalic suture angle beta (P); narrower positions of cephalic suture angles gamma (y) and delta (8); an inflated ("cushion-like") cephalic border instead of a flat one; three cephalic border rims instead of two; no cephalic epiborder furrow; weakly violin shaped glabella with sculpture of fine ridges and granules instead of a strongly violin shaped glabella with sculpture of only granules; thoracic pleural sculpture lacking fine ridges; longer pygidium (Z,

Figure 3-5); rounded posterior pygidial margin instead of a flat one; a post rhachial bump; and pygidial rhachis and pleural region with sculpture of granules only.

Diademaproetus rudimentus differs from D. holzapfeli in having narrower positions of cephalic suture angles gamma (y) and delta (8); inflated ("cushion­ like") cephalic border instead of a flat; three cephalic border rims instead of two;

212 no cephalic epiborder furrow; genal field sculpture without fine ridges; weakly violin shaped glabella with sculpture of fine ridges and granules instead of a strongly violin shaped glabella with sculpture of only granules; thoracic pleural sculpture of granules; longer pygidium (Z, Figure 3-5); rounded pygidial margin instead of a slightly notched one; and pygidial sculpture of granules throughout.

Diademaproetus issoumourensis new species

Plate 3-12, figs. 1-7 & plate 3-13, figs. 1-5.

Derivation of name - Named in reference to the locality from which the species is found, on the western face of Jbel Issoumour of the Ma'der Basin, southern

Morocco.

Diagnosis - Diademaproetus with flat cephalic border; wide positioned cephalic suture angle gamma (y); well defined (deep) occipital lobe furrows; thoracic axis with sculpture of granules; short (Z) and wide (W) pygidium (length (Z) is less than 0.37x the maximum width (W, Figure 3-5)); and pygidial rhachis with sculpture of fine ridges and granules, primarily on posterior portion of rhachial segments.

Material - Holotype UA13271, complete and fully articulated exoskeleton and paratype UA 13682.

213 Occurrence - From the 'Psychopyge horizon' of the western side of Jbel

Issoumour. The horizon is part of the Er Remlia Formation and late Emsian (Gibb

& Chatterton 2010).

Description -Cephalon. "D" shaped and moderately convex. Sagittal length (A,

Figure 3-5) 0.4 lx length of entire specimen. Cephalic border flat and with sculpture of fine ridges. Anterior and lateral portions of cephalic border rimmed by three fine parallel ridges (border rims), between cephalic border and cephalic

border furrow. Anterior of glabella, cephalic border furrow broad and shallow, narrowing and deepening to librigena and genal angle. Cephalic border furrow deepest at base of genal spine. Posterior cephalic border furrow moderately deep, running laterally from axial furrow of occipital ring to junction with lateral cephalic border furrow. Posterior border furrow widest and shallowest at its midlength, anterior of cephalic suture angle omega (Q).

Cranidium. Preglabellar area (A2 + A3) 0.29x length of cephalon (A,

Figure 3-5). Sagittal length of glabella (Ai + A4) 0.7 lx length of cephalon (A,

Figure 3-5). Length of anterior portion of glabella (Ai) is 0.60x cephalon length

(A, Figure 3-5). P- P distance 0.58x length ofcephalon (A, Figure 3-5). y-y distance 0.5 lx length of cephalon (A, Figure 3-5). 5-5 distance 0.79x length of cephalon (A, Figure 3-5). Anterior margin of genal field slightly inset posteriorly, such that anterior margin of genal field, between cranidium and librigena, does not run parallel to cephalic border furrow. Glabella weakly-violin shaped, moderately convex. Glabellar sculpture of moderately large granules and fine

214 ridges, slightly clustered around anterior area and towards sagittal midline. When viewed laterally, anterior portion of glabella is slightly less convex than posterior

portion, but is steeply angled ventrally towards cephalic border. Posterior portion

of glabella less steeply angled than anterior portion, with highest point (dorsally) of glabella being its posterior margin along cephalic midline. In dorsal view, anterior portion of glabella slightly constricted posteromedially of cephalic suture angle gamma (y). Glabella possesses 4 pairs of glabellar furrows. Furrows SI and

S3 are largest and located medial to cephalic suture angles delta (5) and gamma

(y) respectively, appearing linear in shape and curving posteromedially towards cephalic midline. Furrows S2 and S4 are smaller and semi-circular in shape, located anteromedially of S1 and S3 respectively. Palpebral lobes are bean-shaped and slightly sloped ventromedially, towards glabella. Palpebral lobe lateral border rimmed with sculpture of fine granules. Occipital furrows (SO) moderately deep.

Occipital lobe (A4) 0.14x length of cephalon (A, Figure 3-5). Occipital ring with sculpture of moderately large granules clustered towards posterior border.

Librigena. Eyes, large and strongly convex in both horizontal and lateral planes. Both anterior and posterior portions of eye platform present. Anterior portion of eye platform larger than posterior portion and rounder in shape. Genal field with sculpture of sparse granules primarily positioned next to lateral genal field border. Long, medially curved genal spines that terminate lateral of pygidium. Lateral border furrow and posterior border furrows both merge at genal angle. Medial aspect of genal spine slightly more inflated than lateral aspect,

215 which is flat. Genal spine furrow widest and deepest at genal angle, narrowing and shallowing posteriorly.

Hypostome. Unknown.

Thorax consists of 10 segments, representing 0.48x sagittal length (when articulated) and 0.82x maximum width (tr.) of entire specimen. Axial rings are moderately arched and bordered laterally by moderately deep axial furrow.

Maximum width of axis (tr.) is approximately 0.48x maximum width (tr.) of thorax. Axial rings decrease in width (tr.) from anterior to posterior, with most posterior (10th) axial ring being 0.59x width (tr.) of most anterior (1st) axial ring.

Axial ring sculpture consists of both fine terrance ridges and granules, with granules being clustered around posteromedian portion of axial rings. Higher amounts of fine ridge sculpture are present on more posterior axial segments, with most anterior segments adorned with largest concentration of granules. Pleurae incised with moderately deep pleural furrows, shalloing laterally from axial furrow. Medial anterior and posterior margins of pleurae transverse, with more lateral regions curved posterolaterally within approximately 65° of sagittal midline. Lateral portion of pleurae terminate in distinct point. Sculpture of pleurae consists of fine granules, forming line of granules along posterior margin of pleural regions.

Pygidium. 0.12x total length (Z) and 0.5 lx maximum width (W, Figure

3-5) of complete specimen. Pygidium shaped as segment of circle. Proximal portion of pleural anterior margin transverse. Lateral of fulcrum, pleural margin

216 curved posterolateral^ at roughly 47° from transverse. Shallow and broad border

furrow encases lateral and posterior pygidial border. Posterior border, rounded in

shape. Rhachis moderately convex and cone shaped, tapering towards pygidial

midline from anterior to posterior. Rhachis length (Y) and width (X) 0.72x and

0.35x pygidial length (Z) and width (W, Figure 3-5) respectively. Rhachis

composed of 4 rhachial rings and terminal piece. Rhachial ring furrows shallow

posteriorly, with more anterior rhachial ring furrows deeper than posterior ones.

Small post rhachial bump present. Three pairs of curved pleural ribs and three

pairs, two deep and one shallow, of pleural furrows. Pleural ribs curved in shape,

decreasing in size from anterior to posterior. First (most anterior) pleural rib

longest, extending all way to pygidial margin. Pleural furrows are moderately

deep, shallowing as they approach pygidial border. Rhachis with sculpture of

moderately large granules. Pleural region with sculpture of fine granules and fine

ridges on posterior borders of pleural ribs.

Remarks - Diademaproetus issoumourensis is a rarer cornuproetid, with only 3 specimens known from Jbel Issoumour. Phylogenetically, D. issoumourensis is a

relatively primitive diademaproetid, topologically located between D. rudimentus

and the more derived diademaproetids. Significantly, D. issoumourensis possesses a primitive anteromedian cephalic projection, forming a slight triangular shape instead of the typical round shape found in more primitive cornuproetids.

Diademaproetus issoumourensis differs from D, rudimentus in having

217 more widely positioned cephalic suture angles gamma (y) and delta (8); a flat

instead of inflated ("cushion-like") cephalic border; three, not two, cephalic

border rims; a genal field sculpture of evenly spaced granules throughout; deep,

instead of shallow, occipital lobe furrows; a small occipital median node; a

thoracic axis sculpture of only granules; thoracic pleural sculpture of fine ridges; a

short pygidium (Z, Figure 3-5); and a pygidial axis sculpture of fine ridges.

Compared to Diademaproetus praecursor, D. issoumourensis differs by

having a wider positioned cephalic suture angle beta (P); three cephalic border

rims instead of two; no cephalic epiborder furrow; weakly violin shaped glabella

instead of strongly violin shaped glabella; deep, not shallow, occipital lobe furrows; a small occipital median node; a thoracic axis sculpture lacking fine ridges; a rounded posterior pygidial margin instead of a flat one; and a post rhachial bump.

Compared to Diademaproetus holzapfeli, D. issoumourensis differs by having three cephalic border rims instead of two; no cephalic epiborder furrow; a genal field sculpture lacking fine ridges; a weakly violin shaped glabella, instead of strongly violin shaped glabella; thoracic axial sculpture lacking fine ridges; thoracic pleural sculpture with ridges; a rounded posterior pygidial margin instead of an anteriorly notch one; and a pygidial pleural sculpture of granules instead of fine ridges.

Diademaproetus corrugatus new species

Plate 3-7, figs. 1-7.

218 Derivation of name - Named in reference to the species' distinct sculpture of fine ridges throughout the cephalic genal field. This sculpture when viewed dorsally makes the genal field appear "corrugated" or "composed of alternating minor ridges and furrows.

Diagnosis - Diademaproetus with flat cephalic border; wide positioned cephalic suture angle gamma (y); genal field sculpture of fine ridges; presence of single epiborder furrow, located anterior of genal spine; flat posterior pygidial border; and pygidial rhachis with sculpture of fine ridges and granules.

Material - Holotype UA13681, complete and fully articulated exoskeleton.

Occurrence - From Taharajat, south of Jbel Oufatene in the Ma'der basin. Strata are late Emsian in age (Gibb & Chatterton 2010).

Description - Cephalon. "D" shaped and moderately convex. Sagittal length (A,

Figure 3-5) 0.37x length of entire specimen. Cephalic border flat and with sculpture of fine ridges. Anterior and lateral portions of cephalic border rimmed by single fine ridge (border rim), running parallel to cephalic border, between cephalic margin and cephalic border. Anterior of glabella, cephalic border furrow broad and shallow, narrowing and deepening towards librigena and genal angle.

Single small and shallow epiborder cephalic furrow present between lateral cephalic border furrow and genal field. Border furrow deepest at base of genal spine, shallowing anteromedially as it progresses around genal field. Posterior cephalic border furrow moderately deep, running laterally from axial furrow of

219 occipital ring to junction with lateral cephalic border furrow and epiborder furrow.

Posterior border furrow widest and shallowest at its midlength, anterior of cephalic suture angle omega (iQ).

Cranidium. Preglabellar area (A2 + A3) slightly inflated, and 0.28x length of cephalon (A, Figure 3-5). Sagittal length of glabella (Ai + A4) is 0.72x length of cephalon (A, Figure 3-5). Length of the glabella (Ai) is 0.5 lx cephalon length (A,

Figure 3-5). P - P distance 0.7 lx length of cephalon (A, Figure 3-5). y - y distance

0.66x length of cephalon (A, Figure 3-5). 5-5 distance 0.94x length of cephalon

(A, Figure 3-5). Glabella weakly-violin shaped, moderately convex. Glabellar sculpture of moderately large granules. Viewed laterally, anterior portion of glabella is slightly less convex then posterior portion, but is steeply angled ventrally close to margins. Posterior portion of glabella is less steeply angled than anterior, with dorsally highest point of glabella being its posterior margin along cephalic midline. Dorsal view, anterior portion of glabella is slightly constricted posteromedially of cephalic suture angle gamma (y). Glabella has 4 pairs of glabellar furrows. Furrows SI and S3 are largest and located medial to cephalic suture angles delta (6) and gamma (y) respectively, appearing linear in shape and curving posteromedially towards cephalic midline. Furrows S2 and S4 are smaller and semi-circular in shape, located anteromedially of SI and S3, respectively.

Palpebral lobes are bean-shaped and sloped slightly ventrally, towards glabella.

Palpebral lobe lateral border is rimmed with sculpture of fine granules. Occipital furrows (SO) are moderately deep. Occipital lobe (A4) is 0.2 lx length of cephalon (A sloped) and transversely wider than widest portion of glabella. Occipital ring with sculpture of moderately large granules, clustered along posterior border, and fine ridges.

Librigena. Eyes, are large and strongly convex in horizontal and lateral planes. Anterior and posterior lobes of eye platforms are present. Anterior lobe of eye platform is larger and rounder than posterior lobe. Genal field with sculpture of fine ridges throughout. Long, medially curved genal spines are longer than cephalon (A, Figure 3-5) and terminate lateral of pygidium. Lateral cephalic border furrow, epiborder furrow, and posterior cephalic border furrow all merge at genal angle. Medial aspect of genal spine slightly more inflated than flat lateral aspect. Genal spine furrow is widest and deepest at genal angle, narrowing and shallowing posteriorly.

Hypostome. Unknown.

Thorax consists of 10 segments, representing 0.46x sagittal length of entire specimen. Axial rings are moderately arched and bordered laterally by moderately deep axial furrows. Axial rings decrease in width (tr.) from anterior to posterior, with most posterior (10th) axial ring being 0.58x width of most anterior (1st) axial ring. Axial ring sculpture consists of both fine terrace ridges and granules. Pleurae are incised with moderately deep pleural furrows. Pleural furrows are deepest slightly proximal to fulcrum, shallowing laterally. Anteromedial and posteromedial margins of pleurae are transverse, with more lateral regions curved

221 posterolateral^ within approximately 70° of sagittal midline. Pleurae with

sculpture of fine ridges.

Pygidium. 0.14x (Z, Figure 3-5) total length of complete specimen.

Shallow and broad border furrow encases lateral and posterior pygidial border.

Posterior border is flat. Rhachis is moderately convex and cone shaped, tapering

towards sagittal pygidial midline from anterior to posterior. Rhachis length (Y)

and width (X) 0.87x and 0.29x pygidial length (Z) and width (W, Figure 3-5)

respectively. Rhachis is composed of 4 rhachial rings and terminal piece. Rhachial

ring furrows become shallower posteriorly. Anterior rhachial ring furrows deeper

than posterior ones. Small post rhachial bump present. Four pairs of curved

pleural ribs and three pairs of moderately deep pleural furrows. Pleural ribs

curved in shape, decreasing in size from anterior to posterior. Pleural furrows are

moderately deep, shallowing distally.

Remarks - Diademaproetus corrugatus is much rarer than D. rudimentus at

Taharajat, with only one specimen known. Unfortunately, the pygidium of the

holotype was damaged during preparation so not all pygidial characters could be

coded, primarily those involving sculpture. Phylogenetically, D. corrugatus is

located between D. issoumourensis and more derived diademaproetids.

Diademaproetus corrugatus differs from D. issoumourensis in having two

cephalic border rims, not three; a cephalic epiborder furrow; genal field sculpture of fine ridges instead of granules; shallower occipital furrows; thoracic pleural

222 sculpture of fine ridges, no granules; a longer pygidium (Z, Figure 3-5); and a flat posterior pygidial margin instead of a rounded one.

Compared to Diademaproetus praecursor, D. corrugatus differs by having a wider positioned cephalic suture angle beta ((3); narrower position of cephalic suture angle delta (8); a genal field sculpture of fine ridges instead of granules; a weakly violin shaped glabella instead of a strongly violin shaped one; a small occipital median node; thoracic pleural sculpture without granules; a comparatively longer pygidium (Z, Figure 3-5); and a small post rhachial bump on the pygidium. Diademaproetus corrugatus differs from D. holzapfeli by having a more narrowly positioned cephalic suture angle delta (5); genal field sculpture lacking granules; a weakly violin shaped glabella, not a strongly violin shaped glabella; and a flat pygidial posterior margin instead of one that is anteriorly notched.

Diademaproetus auxiliarus new species

Plate 3-8, figs. 1-7 & plate 3-9, figs. 1-7.

Derivation of name - Named because of its close resemblance to

Diademaproetus praecursor and information it provided in deciphering the gradational increase in anteromedian cephalic projection size from primitive to more derived diademaproetids. "Auxiliary" meaning "providing supplementary or additional help or support."

223 Diagnosis - Diademaproetus with flat cephalic border; narrow positioned cephalic suture angle gamma (y); intermediate position of cephalic suture angle delta (8); presence of a single epiborder furrow, located anterior of genal spine; shallow glabellar furrows; flat posterior border of pygidium; and pygidial rhachis with sculpture of granules.

Material - Holotype UA13689 (complete and fully articulated exoskeleton), paratypes UA13690 and UA13691.

Occurrence - From the Assa locality, northwest of the town of Assa. Strata are part of Khebchia Formation of the Tindouf Basin and late Emsian in age (Becker et al, 2004a; Becker et al. 2004b; Hollard 1978; Jansen et al. 2004).

Description - Cephalon "D" shaped and moderately convex. Sagittal length (A,

Figure 3-5) 0.40x length of entire articulated specimen. Anterior border flat, broad, and rimmed by two fine parallel ridges (border rims) between cephalic border and cephalic margin. Short and wide anteromedian cephalic projection protrudes anterior of preglabellar field. Weak anteromedian cephalic projection furrow (depression) present between cephalic border furrow and anterior margin of anteromedian cephalic projection. Anterior cephalic border furrow narrow and shallow (almost inconspicuous) anterior of glabella, widening and deepening distinctly towards genal angle. Cephalic epiborder furrow parallels lateral cephalic border furrow. Cephalic border furrow shallow and narrow, deepening towards genal angle. Posterior cephalic border furrow moderately deep, running laterally from axial furrow to junction with lateral cephalic border and base of genal spine.

224 Cranidium. Preglabellar area length (A2 + A3) 0.30x cephalon length (A,

Figure 3-5). Cephalic border broad and flat, with sculpture of fine ridges. Area

anterior of cephalic border furrow (A3) and length from anterior limit of glabella

to the cephalic border furrow (A2) are 0.23x and 0.07x cephalon length (A, Figure

3-5) respectively. Sagittal length of glabella, including occipital lobe (LO), (Ai +

A4) 0.70x length of cephalon (A, Figure 3-5). Length of anterior portion of glabella (Ai) 0.53x length of cephalon (A, Figure 3-5). (3 - P distance 0.56x length of cephalon (A, Figure 3-5). y - y distance 0.5 lx length of cephalon (A, Figure

3-5). 8-8 distance 0.80x length of cephalon (A, Figure 3-5). Glabella subcylindrical in shape, moderately convex and with sculpture of fine ridges and granules. Anterior portion of glabella slightly constricted posterior-medially of cephalic suture angle gamma (y). Glabella possesses four shallow furrows, with most posterior S1 furrows longest and deepest, located medial of cephalic suture angle delta (8), roughly linear in shape with their medial portions deflected posterior-medially towards cephalic midline approximately half way along their length. S2 furrows ovoid, located anterior-medially of SI furrows. S3 furrows similar shaped as S1 furrows, however smaller and located medial of cephalic suture angle gamma (y). S4 furrows small and roughly ovoid, located anteromedially of S3 furrows. Palpebral lobes are bean-shaped and slightly sloped ventrally towards lateral portion of glabella. Occipital furrows (SO) are shallow.

Occipital lobe (A4) 0.17x length of cephalon (A, Figure 3-5), and adorned with granules..

225 Librigena. Eyes, large and strongly convex in horizontal and lateral planes.

Anterior and posterior lobes of eye platform present. Anterior lobe of eye platform larger and rounder than posterior lobe. Genal field with sculpture of granules.

Long, medially curved genal spines are longer than cephalon (A, Figure 3-5) and terminate lateral of pygidium. Lateral border furrow and posterior border furrow all merge into genal spine furrow at genal angle. Medial aspect of genal spine inflated, lateral aspect flat like cephalic border. Genal spine furrow is widest and deepest at genal angle, narrowing and shallowing posteriorly.

Hypostome. Central hypostomal body with anteromedial convex bulge

(rhynchos) and rounded posterior. Anterior border raised (ventrally in life position) above lateral and posterior borders. Central hypostomal body seperated from anterior border by anterior border furrow. Lateral borders narrow, with moderately deep and short lateral border furrows. Posterior margin semi-circular in shape. Both lateral and posterior borders slightly convex. Anterior wings down- sloped laterally, and bluntly triangular. Entire hypostome covered by sculpture of dense anastomosing terrace ridges. Terrace ridges on lateral and posterior borders run parallel to hypostomal border.

Thorax consists of 10 segments, 0.44x sagittal length (when articulated) and 0.78x width (tr.) of entire specimen. Axial rings moderately arched and bordered laterally by moderately deep axial furrow. Maximum width of axis (tr.) is approximately 0.40x maximum width (tr.) of thorax. Axial rings increase in width from anterior to posterior in first five segments, fifth axial segment being

226 widest (tr.). Posterior of fifth segment, axial rings decrease in width. Most posterior (10th) axial ring is narrowest and 0.59x as wide (tr.) as widest (5th) axial ring. Axial rings sculpture consists of fine ridges and granules. Pleurae are incised with moderately deep pleural furrow running laterally from near axial furrow beyond pleural midwidth. Pleural furrows deepest near pleural fulcrum. Medial regions of pleurae are transverse, lateral regions curved posterolaterally within approximately 67° of sagittal midline.

Pygidium is 0.16x total length (Z) and 0.59x maximum width (W, Figure

3-5) of complete specimen. Pygidium is segment of circle shaped. Anterior margin of medial pleural region is transverse. Lateral of fulcrum, pleural margin angles slightly posterolaterally at roughly 40° from transverse. Shallow and broad border furrow encases lateral and posterior pygidial border. Posterior margin is flat. Rhachis is strongly convex and cone shaped, tapering from anterior to posterior. Rhachis length (Y) and width (X) 0.7lx and 0.24x pygidial length (Z) and width (W, Figure 3-5) respectively. Rhachis composed of 4 rhachial rings and terminal piece. Anterior rhachial ring furrows deeper than more posterior rhachial ring furrows. Rhachis with sculpture of granules. Four pairs of curved pleural ribs and three pairs of moderately deep interpleural furrows. Interpleural furrows shallow towards pygidial border. Pleural rib structure is imbricate. Pleural furrows deep, shallowing laterally towards pygidial border. Pleural pygidial field sculpture of fine ridges and granules.

227 Remarks - Diademaproetus auxiliarus is found at Assa. Of specific interest is the slightly more pronounced anteromedian cephalic projection of D. auxiliarus compared to D. issoumouensis, which forms a noticeable projection of the anterior margin of the cephalon. Furthermore, the anterior cephalic projection in D. auxiliarus is less prominent and wider than that of the slightly more derived D. praecursor. Phylogenetically, D. auxiliarus is located between D. corrugatus and

D. praecursor.

Compared to Diademaproetus corrugatus, D. auxiliarus differs in having a more narrowly positioned cephalic suture angle gamma (y); widely positioned cephalic suture angle delta (5); genal field sculpture of granules, not fine ridges; glabellar and thoracic pleural sculpture of fine ridges and granules; and no occipital median node. Diademaproetus auxiliarus differs from D. praecursor by having a more widely positioned cephalic suture angle beta ((3); narrowly positioned cephalic suture angle gamma (y); weakly violin shaped glabella, with sculpture of fine ridges and granules; and longer pygidium (Z, Figure 3-5); and most anterior pleural rib that terminates prior to reaching the lateral pygidial margin.

Diademaproetus auxiliarus differs from D. holzapfeli in having a narrowly positioned cephalic suture angle gamma (y); genal field sculpture lacking fine ridges; weakly violin shaped, not strongly violin shaped glabella; long pygidium

(Z, Figure 3-5); most anterior pleural rib that terminates prior to the lateral pygidial margin; and a flat, not anteriorly notched, posterior pygidial margin.

228 Diademaproetus langus new species

Plate 3-10, figs. 1-7; plate 3- 11, figs. 1-5; plate 3-13, fig. 6.

Derivation of name - Named in reference to the genus' distinct long and narrow

anteromedian cephalic projection, which appears tongue like due to its large and

relatively deep associated furrow. Langue meaning "tongue" in French.

Diagnosis - Diademaproetus with five or more cephalic border rims

encompassing anterior cephalic projection; wide position of cephalic suture angle

delta (8); prominent glabellar furrows; genal field sculpture of fine granules;

glabella weakly violin shaped, with sculpture of fine ridges and granules;

palpebral lobes and thoracic axial regions with sculpture of fine ridges; thoracic

pleural region lacking sculpture (smooth); rounded pentagon shaped pygidium;

and pygidial rhachis and pleural region lacking sculpture (smooth).

Material - Holotype UA13685 (complete and fully articulated exoskeleton) and

paratypes UA13683 and UA13684.

Occurrence - From the 'Diademaproetus horizon' at Jbel Oufatene west

northwest of the village of Lahfira in the Ma'der Basin, southern Morocco. Strata

are part of the El Otfal Formation and Eifelian (Gibb & Chatterton 2010).

Description - Cephalon is "D" shaped and moderately convex. Sagittal length (A,

Figure 3-5) is 0.44x length of entire specimen. Cephalic border is flat, with sculpture of fine ridges throughout. Anterior portion of cephalic border rimmed is

by five fine ridges (border rim), running parallel to cephalic border, between 229 cephalic margin and border furrow. Anteromedian cephalic projection is long and narrow and back by deep anteromedian cephalic projection furrow (depression) that is merged with anterior portion of cephalic border furrow. Cephalic border furrow is broad and moderately deep, narrowing as it approaches librigena.

Posterior cephalic border furrow is moderately deep, running laterally from axial furrow of occipital ring to junction with lateral cephalic border furrow. Posterior border furrow is widest and shallowest next to occipital lobe (LO), narrowing and deepening as it approaches genal angle.

Cranidium. Preglabellar area (A2 + A3) is slightly inflated, 0.33x length of cephalon (A, Figure 3-5). Sagittal length of glabella (Ai + A4) is 0.67x length of cephalon (A, Figure 3-5). Length of anterior portion of glabella (Ai) is 0.47x cephalon length (A, Figure 3-5). p - p distance is 0.44x length of cephalon (A,

Figure 3-5). y - y distance is 0.49x length of cephalon (A, Figure 3-5). 8-8 distance is 0.73x length of cephalon (A, Figure 3-5). Glabella is weakly-violin shaped and moderately convex. Glabellar sculpture consists of granules, concentrated away from center and towards margins of frontal lobe and occipital ring, with fine ridges. Viewed laterally, anterior portion of glabella is less convex then posterior, and steeply angled ventrally adjacent to preglabellar furrow.

Posterior portion of glabella is less steeply angled than anterior portion, with highest (dorsally) point of glabella at posterior margin along cephalic midline.

Dorsal view, anterior portion of glabella is slightly constricted posteromedially of cephalic suture angle gamma (y). Glabella contains 4 pairs of glabellar furrows.

230 SI furrows are shallow, large, located medial to cephalic suture angle delta (5), and triangular in shape. S2 furrows are small, shallow, semi-circular in shape, and located anteromedially of S1 furrows. S3 furrows have triangular shape, not as deep and large as S1 furrows, and are positioned medially of cephalic suture angle gamma (y). S4 furrows are shallow, linear, and posteromedially directed towards cephalic midline anterior of S3 furrow. Palpebral lobes are bean-shaped and slightly sloped ventrally, towards glabella. Palpebral lobes lateral border is rimmed with fine ridge sculpture. Occipital furrows (SO) are deep and small.

Occipital median node is present. Occipital lobe (A4) is 0.20x length of cephalon

(A, Figure 3-5), and wider (tr.) than widest portion of frontal lobe of glabella.

Occipital ring has sculpture of granules and fine ridges clustered around posterior border.

Librigena. Eyes are large and strongly convex in horizontal and lateral planes. Anterior and posterior lobes of eye platform are present. Anterior lobe of eye platform is larger and rounder than posterior lobe. Genal field has sculpture of very fine ridges. Long, medially curved genal spines are longer than cephalon (A,

Figure 3-5) and terminate lateral of pygidium. Lateral cephalic border furrow and posterior cephalic border furrow merge at genal angle. Medial aspect of genal spine is slightly more inflated than flattened lateral aspect. Genal spine furrow is widest and deepest at genal angle, narrowing and shallowing posteriorly.

Hypostome. Length (HL) is 1.06x width (HWi, Figure 3-5). Central hypostomal body slightly longer than wide, with anterior-medial convex bulge

231 (rhynchos) and rounded posterior. Anterior border raised (ventrally in life position) above lateral and posterior borders. Central hypostomal body seperated from anterior border by anterior border furrow. Lateral borders narrow with moderately deep and short lateral border furrows. Posterior border semi-circular in shape posteriorly, with short pair of posterior-lateral marginal spines. Both lateral and posterior borders slightly convex. Anterior wings are steeply down- sloped laterally and bluntly triangular. Entire hypostome covered by sculpture of dense anastomosing terrace ridges. Terrace ridges on lateral and posterior borders run parallel to hypostomal margin.

Thorax. Consists of 10 segments, representing 0.40x sagittal length (when articulated) of entire specimen. Axial rings are moderately arched and bordered laterally by moderately deep axial furrows. Axial rings decrease in width from anterior to posterior, with most posterior (10th) axial ring being 0.67x width (tr.) of most anterior (1st) axial ring. Axial ring sculpture consists of fine terrace ridges.

Pleurae are incised with moderately deep pleural furrows, running laterally from near axial furrow to posterior of fulcrum. Pleural furrows are deepest proximally, shallowing distally. Medial anterior and posterior margins of pleurae are transverse, with more lateral regions curved posterolateral^ within approximately

67° of sagittal midline. Pleural region lacks sculpture (smooth).

Pygidium is 0.16x total length (Z, Figure 3-5) of complete specimen, and has rounded square shape. Shallow and broad border furrow encases lateral and posterior pygidial border. Posterior border is distinctly notched anteriorly. Rhachis

232 is moderately convex and cone shaped, tapering backwards. Rhachis length (Y)

and width (X) are 0.76x and 0.3lx pygidial length (Z) and width (W, Figure 3-5)

respectively. Rhachis is composed of 4 rhachial rings and terminal piece. Rhachial

ring furrows become shallower posteriorly, with most anterior rhachial ring

furrow deepest. Small post rhachial bump is present. Three pairs of curved pleural

ribs and three pairs of moderately deep pleural furrows are present. Interpleural furrows are shallow. Pleural ribs are curved, decreasing in size from more anterior

to more posterior ribs, with first rib reaching pygidial margin. Pleural furrows are

moderately deep, shallowing distally to pygidial border. Rhachis lacks sculpture

(smooth), and pleural region has sculpture of fine granules.

Remarks - Diademaproetus langus is the most common diademaproetid found in

the 'Diademaproetus horizon' of Jbel Oufatene. Phylogenetically, D. langus is the

most derived Diademaproetus, being situated next to D. holzapfeli and the new, even more derived, genus Pontoproetus. D. langus also possesses the longest and

narrowest anteromedian cephalic projection of all diademaproetids.

Diademaproetus langus differs from D. holzapfeli in displaying a more

narrowly positioned cephalic suture angle gamma (y); a wider positioned cephalic suture angle delta (5); a five, instead of two, cephalic border rims; no cephalic epiborder furrows; a weakly violin shaped glabella; a smooth genal field; thoracic axial sculpture of only fine ridges; no (smooth) thoracic pleural sculpture; a rounded pentagon shaped pygidium; and no (smooth) sculpture on the pygidial rhachis.

233 Genus Pontoproetus gen. no v.

Type species - Pontoproetus truncatus sp. nov.

Other included species - Pontoproetus granulosus sp. nov.

Derivation of name - Named in reference to the absence (truncation) of the anterior cephalic border furrow within the preglabellar field. Upon initial examination by the primary author the truncated portion of the cephalic border furrow appeared to be "bridged" by the exoskeleton. Hence the term "Pont" which is french for "bridge," and proetus in reference to the order of trilobites that this genus belongs to.

Diagnisis - Corauproetid genus with flat cephalic border; long (sag.) preglabellar area (greater than or equal to 0.25x length of cephalon (A); truncated anterior portion of cephalic border furrow (Figure 3-10, image A); wide position of cephalic suture angle beta ((3); strongly violin shaped glabella with sculpture of granules; prominent (deep) occipital lobe furrows; moderately long genal spines, that terminate anterior of pygidium; thoracic axis with sculpture of fine ridges; short (sag.) and wide (tr.) pygidium ((Z) is less than 0.37x maximum width (W,

Figure 3-5)); and posterior pygidial margin slightly notched.

Remarks - Pontoproetus is a useful taxon to encompass a distinct group of

Moroccan cornuproetids with a truncated, or absent, anterior cephalic border furrow in front of glabella. This truncation of the anterior cephalic border furrow causes the cephalic border furrow to curve posteriorly, after it crosses onto the

234 cranidium from the librigena. Interestingly, both Diademaproetus corrugatus and

D. auxiliarus appear to have a primitive state of this trait, with their anterior

cephalic border furrows slightly indented backwards on the cranidium. This

indent is not strong enough to cause the cephalic border furrow to contact the

anterior portion of the glabella, however it does prevent the cephalic border furrow from having a continuous arch or crescent shape. Both species of

Pontoproetus also possess a short anteromedian cephalic projection that is round

in shape, rather than the more triangular shape found in some species of

Diademaproetus, and a triangular shaped pattern of granulose sculpture on the glabella. Furthermore, the overall convexity of the Pontoproetus cephalon is less, and the rhachial ring furrows of the pygidium are more sigmoidal than crescent shaped.

Pontoproetus truncatus new species

Plate 3-14, figs. 1-7 & plate 3-15, figs. 1-5.

Derivation of name - Named in reference to the species' distinct truncated cephalic border furrow within the preglabellar field.

Diagnosis - Pontoproetus with wide positioned cephalic suture angle gamma (y); narrow positioned cephalic suture angle delta (5); normal (not furrowed or sculpted) palpebral rim structure; anterior and posterior lobes of eye platform present; and glabellar sculpture of fine granules restricted to anterior portion and

235 midline of glabella, forming triangular shaped pattern; and presence of single

pygidial border rim.

Material - Holotype UA13701 (complete articulated specimen) and paratypes

UA13702 & UA13073..

Occurrence - Found at Timarzite, east northeast of the town bou Dib, and in the

'Thysanopeltis horizon,' Taboumakhlouf Formation, Jbel Zireg. Based on conodont faunas, Timerazite is Eifelian (Bultynck 1987; Belka et al. 1997) while

the strata are Eifelian (Gibb & Chatterton 2010).

Description - Cephalon is "D" shaped and weakly convex. Sagittal length (A,

Figure 3-5) is 0.37x length (sag.) of entire specimen. Cephalic border is flat, with sculpture of fine ridges. Anterior and lateral portions of cephalic border are rimmed by single fine ridge (border rim), running parallel to cephalic border, between cephalic margin and cephalic border furrow. Short and rounded anteromedian cephalic projection protrudes anterior of preglabellar field.

Anteromedian cephalic projection is accompanied by broad and shallow anteromedian cephalic projection furrow (depression) posterior of anteromedian cephalic projection. Anterior of glabella, cephalic border furrow is absent, with deep cephalic border furrow arching posteriorly and intersecting preglabellar furrow after crossing onto cranidium from librigena. Narrow posterior cephalic border furrow of moderate depth runs laterally from axial furrow towards front of occipital ring to junction with lateral cephalic border furrow and genal spine.

236 Cranidium. Preglabellar area (A2 + A3) is slightly inflated, 0.32x length of cephalon (A, Figure 3-5). Sagittal length of glabella (Ai + A4) is 0.68x length of cephalon (A, Figure 3-5). Length of anterior of glabella (Ai) is 0.5 lx cephalon length (A, Figure 3-5). p - p distance is 0.52x length of cephalon (A, Figure 3-5). y - y is distance 0.43x length of cephalon (A, Figure 3-5). 8 - 5 is distance 0.67x length of cephalon (A, Figure 3-5). Glabella is strongly-violin shaped and moderately convex. Glabellar sculpture consists of small granules, which are clustered into a triangular configuration in anterior portion of glabella, and fine ridges. Dorsal view, anterior portion of glabella is highly constricted posteromedially of cephalic suture angle gamma (y). Glabella contains 4 pairs of glabellar furrows. SI furrows are very shallow, large, located medial to cephalic suture angle delta (8), and triangular in shape. S2 furrows are small, shallow, semi-circular in shape, and located anteromedially of SI furrows. S3 furrows display similar triangular shape as S1 furrows, however are not as deep and long as SI furrows and positioned medially of cephalic suture angle gamma (y). S4 furrows are shallow, linear, posterior-medially directed towards cephalic midline, and anterior of S3 furrows. Palpebral lobes are bean-shaped and slightly sloped ventrally, towards glabella. Marginal rim of palpebral lobes lack sculpture.

Occipital furrow (SO) is deep. Small occipital median node is present. Occipital lobe (A4) is 0.17x length of cephalon (A, Figure 3-5) and transversely wider than widest portion of glabella. Occipital ring has sculpture of fine ridges.

237 Librigena. Eyes are large and strongly convex in horizontal and lateral planes. Anterior and posterior lobes of eye platform are present. Anterior lobe of eye platform is only slightly larger and more rounded than posterior lobe. Distinct eye socle is present, forming continuous ridge underneath entire eye. Genal field has sculpture of fine ridges. Long, medially curved genal spines are longer than cephalon (A) and terminate anterior of pygidium. Lateral cephalic border furrow and posterior cephalic border furrow merge at genal angle. Medial aspect of genal spine slightly more inflated than flat lateral aspect. Genal spine furrow is widest and deepest at genal angle, narrowing and shallowing posteriorly.

Hypostome. Unknown.

Thorax. Consists of 10 segments, representing 0.44x sagittal length of entire specimen. Axial rings are moderately arched and bordered laterally by axial furrows of moderate depth. Axial rings increase in width from most anterior (1st) segment to fourth (4th) segment, before decreasing in width posteriorly from fourth (4th) segment to most posterior (10th) segment. Width (tr.) of narrowest

(10th) axial ring is 0.68x width (tr.) of widest (4th) axial ring. Axial ring sculpture consists of fine terrace ridges. Pleurae are incised with moderately deep pleural furrows. Distal ends of thoracic segments are rounded. Pleural furrows are deepest near fulcrum. Medial anterior and posterior margins of pleurae are transverse, with more lateral regions curved posterolaterally approximately 74° of sagittal midline. Pleurae have sculpture of fine ridges.

238 Pygidium is 0.19x total length (Z, Figure 3-5) of complete specimen, and

segment of circle shape. Shallow and broad border furrow encases lateral and

posterior pygidial border with single fine ridge around margin. Posterior border is

weakly notched anteriorly. Rhachis is moderately convex and cone shaped,

tapering backwards. Rhachis length (Y) and width (X) are 0.78x and 0.29x

pygidial length (Z) and width (W, Figure 3-5) respectively. Rhachis is composed

of 4 curved rhachial rings and terminal piece. Rhachial ring furrows become

shallower posteriorly, with most anterior rhachial ring furrow being deepest, and

containing preannular lobes laterally in first two furrows. Small post rhachial

bump present. There are three pairs of curved pleural ribs and two pairs of

shallow pleural furrows. Pleural ribs are curved, decreasing in size from anterior

to posterior. Pleural furrows are moderately shallow, shallowing distally. Both

rhachis and pleural region have sculpture of fine ridges.

Remarks - Pontoproetus truncatus is quite common in the ' Thysanopeltis horizon' at Jbel Zireg and from Timarzite.

Pontoproetus truncatus differs from Diademaproetus holzapfeli by having a weakly convex cephalon; a round, rather than triangular shaped, anteromedian cephalic projection; a narrower positioned cephalic suture angle delta (5); a truncated (absent) anterior cephalic border furrow; no cephalic epiborder furrow; a glabella with granulose sculpture concentrated strictly into posteriorly directed triangle on anterior portion of the frontal lobe of the glabella; a genal field and thoracic axial sculpture lacking granules; and a fine pygidial border rim.

239 Compared to Diademaproetus langus, Pontoproetus truncatus differs by

having a weakly convex forward cephalon; round, rather than triangular shaped,

anteromedian cephalic projection; wider positioned cephalic suture angle gamma

(y); a narrower positioned cephalic suture angle delta (8); only one cephalic

border rim; a truncated (absent) anterior cephalic border furrow; a strongly violin

shaped glabella with granules forming a triangular pattern pointing posteriorly

from the anterior portion of the glabella; a palpebral lobe rim with no sculpture;

thoracic pleural field sculpture of fine ridges; a segment of circle pygidium; and

an entire pygidium (rhachis and pleurae) with sculpture of fine ridges.

Pontoproetus granulosus new species

Plate 3-16, figs. 1-6.

Derivation of name - Named in reference to the species distinct and less constrained granulose sculpture throughout the glabella and around the rim of the palpebral lobes.

Diagnosis - Pontoproetus with narrow positioned cephalic suture angle gamma

(y); wide positioned cephalic suture position delta (8); palpebral lobes rimmed with granules; only posterior lobe of eye platform present; and glabellar sculpture of granules less constrained and more widely spaced.

Material - Holotype UA13700, partial articulated exoskeleton missing right librigena.

240 Occurrence - Found in the 'Morocconites/Metacanthina horizon' at Jbel El

Mrakib. Strata are part of the Tazoula'it Formation and Emsian in age (Gibb &

Chatterton 2010)

Description - Cephalon is "D" shaped and weakly convex. Sagittal length (A,

Figure 3-5) is 0.32x length (sag.) of entire specimen. Cephalic border is flat, with

sculpture of fine ridges throughout. Anterior and lateral portions of cephalic border are rimmed by single fine ridge. Short and rounded anteromedian cephalic projection protrudes anterior of preglabellar field. Anteromedian cephalic projection accompanied by narrow and shallow anteromedian cephalic projection furrow (depression) posterior of anteromedian cephalic projection. Anterior of glabella, cephalic border furrow is absent, with distinct cephalic border backwards and medially, intersecting preglabellar furrow. Border furrow narrows and deepens towards genal angle. Narrow posterior cephalic border furrow runs laterally from axial furrow, opposite occipital ring, to junction with lateral cephalic border furrow and genal spine.

Cranidium. Preglabellar area (A2 + A3) is slightly inflated, 0.3 lx length of cephalon (A, Figure 3-5). Sagittal length of glabella (Ai + A4) is 0.69x length of cephalon (A, Figure 3-5). Length of anterior portion of glabella (Ai) is 0.53x cephalon length (A, Figure 3-5). P - P is distance 0.52x length of cephalon (A,

Figure 3-5). y-y distance is 0.50x length of cephalon (A, Figure 3-5). 8-8 distance is 0.73x length of cephalon (A, Figure 3-5). Glabella is strongly-violin shaped and moderately convex. Glabellar sculpture consists of moderate sized

241 granules, which are concentrated along glabellar mid-line, in frontal lobe and back of occipital ring, and fine ridges. Dorsal view, anterior portion of glabella is distinctly constricted posteromedially near cephalic suture angle gamma (y).

Glabella has 4 pairs of glabellar furrows. SI furrows are shallow, large, located medial to cephalic suture angle delta (5), and triangular in shape. S2 furrows small, shallow, semi-circular in shape, and located anteromedially of S1 furrows.

S3 farrows display similar triangular shape to SI furrows, not as deep and large as

SI furrows, and positioned medial of cephalic suture angle gamma (y). S4 furrows are shallow, linear, and posteromedially directed towards cephalic midline anterior of S3 furrow. Palpebral lobes are bean-shaped and slightly sloped ventrally, towards glabella. Palpebral lobe lateral border has sculpture of fine granules.

Occipital furrow (SO) is deep. Small occipital median node is present. Occipital lobe (A4) is 0.16x length of cephalon (A, Figure 3-5), and transversely wider than widest portion of anterior lobe of glabella. Occipital ring has sculpture of fine ridges, with fine granules along posterior margin.

Librigena. Eyes are large and strongly convex in horizontal and lateral planes. Only posterior circular lobes of eye platform are present. Genal field has sculpture of fine ridges. Long, medially curved genal spines are longer than cephalon (A, Figure 3-5) and terminate in front of pygidium. Lateral cephalic border furrow and posterior cephalic border furrow merge at genal angle. Medial aspect of genal spine is slightly more inflated than flat lateral aspect.. Genal spine furrow is widest and deepest at genal angle, and narrows and shallows posteriorly.

242 Hypostome. Unknown.

Thorax. Consists of 10 segments, representing 0.42x sagittal length of entire specimen. Axial rings are moderately arched and bordered laterally by moderately impressed axial furrows. Axial rings increase in width from most anterior (1st) segment to fourth (4th) segment, before decreasing in width posteriorly from fourth (4th) segment to most posterior (10th) segment. Width (tr.) of narrowest (10th) axial ring is 0.68x width (tr.) of widest (4th) axial ring. Axial ring sculpture consists of fine terrace ridges. Pleurae are incised with moderately deep pleural furrows. Thoracic segments terminate in rounded point. Pleural furrows are deepest near fulcrum, shallowing proximally and laterally.

Anteromedial and posteromedial margins of pleurae are transverse, with more lateral regions curved posterolaterally to approximately 76° of sagittal midline.

Pleurae have sculpture of fine ridges.

Pygidium is 0.26x total length (Z, Figure 3-5) of complete specimen, and forms segment of circle shape. Shallow and broad border furrow encases lateral and posterior pygidial border. Posterior border is weakly notched anteriorly.

Rhachis is moderately convex and cone shaped, tapering backwards. Rhachis length (Y) and width (X) are 0.76x and 0.33x pygidial length (Z) and width (W,

Figure 3-5) respectively. Rhachis is composed of 4 curved (arched) rhachial rings and terminal piece. Rhachial ring furrows become shallower posteriorly, with most anterior rhachial ring furrow being deepest. Small post rhachial bump is present. Three pairs of curved pleural ribs and three pairs of shallow pleural

243 furrows are present. Pleural ribs are curved, decreasing in size from anterior to posterior. Interpleural furrows are shallow. Pleural furrows are shallow, shallowing distally. Both rhachis and pleural region have sculpture of fine ridges.

Remarks - Pontoproetus granulosus is rare, and has been found in the

'Morocconites/Metacanthina horizon' of Jbel Mrakib. Only a single partially articulated specimen is known.

Like Pontoproetus truncatus, P. granulosus possesses a truncated anterior cephalic border furrow and rounded anteromedian cephalic projection. However,

P. granulosus differs from P. truncatus by having a narrower anteromedian cephalic projection depression; more narrowly positioned cephalic suture angle gamma (y); more widely positioned cephalic suture angle delta (8); glabellar sculpture of granules that is less constricted to a triangular shape on the frontal glabellar lobe and more spread out about the glabellar midline and back of the occipital lobe; a palpebral lobe rimmed with shallow granules; and no pygidial border rim.

Pontoproetus granulosus differs from Diademaproetus langus by possessing a weakly convex cephalon; a round, rather than triangular shaped, anteromedian cephalic projection; only one cephalic border rim, not five; a truncated (absent) anterior cephalic border furrow; palpebral lobe rimmed with granules; a thoracic pleural region and the entire pygidium sculpted with fine ridges; and a segment of circle shaped pygidium.

244 AFRICA

SPAIN

• Fes E -Tafialt Basin, MOROCCO Figure 3-4

Erfoud ilattKl D - Ma'der basin, Ouomnle a Figure 3-3 Foum Zguid ™.«l%l ALGERIA C - Tindouf Basin, Figure 3-2

Figure 3-1. Locality maps of Africa and Morocco. A: Map of Africa depicting Morocco (in-filled area) (Modified from Philip (1991) Gibb & Chatterton (2007, 2010); McKellar & Chatterton (2009); Chatterton & Gibb (2010)). B: Map of Morocco indicating locations of basins surveyed in this study (modified from Gibb & Chatterton (2007,2010); McKellar & Chatterton (2009); Chatterton & Gibb (2010)). C, D, E: Location of individual study basins within Morocco (refer to Figures 3-2 to 3-4 for more detailed maps of these areas).

245 Figure 3-2. Map and stratigraphy of select localities of the southern Moroccan Tindouf Basin. Locations of individual taxa indicated on individual stratigraphic columns correspond to appropriate locations on the outcrop map. (Diagrams adapted from Fetah et al. (1998); Gibb (2005); Chatterton et al. (2006); Gibb & Chatterton (2007, 2010); McKellar & Chatterton (2009)).

246 Tindouf Basin

Outcrop Map Hffcynian Dotorite (Uppf P>Nitozoic)

Devonian

Ordovidan

w j£- 10 km Stratigraphy Zouama

100_ ""mtr*"!- y- TI Zguilma M R ^0 Diademaprvetus auxiliarus AH Assa RN € Diademaptwtuspraecursor Note The locality of Ana » pMl of (he Tindouf Basin, H but notmriodcrf on the aboveastcrop map M His A laoMed fiatber sontkwat« 28*3©"06J*N and N 09^743.4-W. : _ .T - r _ T Diademaprottus mohamedi • i - ^ J Base of Limestone ' Jbei Zguilma

Shaly Limestone 50„ - ZGEE3 Interbedded Limestone » and calcareous shale R 100 F Shale with calcareous O nodules I R MA Calcaraous Shale lit T -r-:€ O -S--3- N Sandstone Iftfl € •3-. -ZGEE2 SIRy Shale ZGEE1 €

247 Figure 3-3. Map and stratigraphy of select localities of the southern Moroccan Ma'der Basin. Locations of individual taxa indicated on individual stratigraphic columns correspond to appropriate locations on the outcrop map. (Diagrams adapted from Fetah et al. (1998); Gibb (2005); Chatterton et al. (2006); Gibb & Chatterton (2007,2010); McKellar & Chatterton (2009); Chatterton & Gibb (2010)).

248 Ma'der Timerzk#* Basin Ciprc 4 Outcrop Map

j j Devonian

Location of Ma'der Basin sections: 1 - bou Ofb

3 * Issoumour West 4 = Oufatdne 9 — MrSKIO 6-Zireg L_LJ!r ^

Stratigraphy Oufat&ne E Pontopmetus truncates • Brachiopod Lag B Pontoproetus granulosus (5 Diademaproetus /angus ELOTFAL Diademaproetus issoumourensis FORMATION 9 CO Diademaproetus mohamedi *— Diademaproetus horizon © «— Garastos/Phacops horizon r— Coltranafa horizon iwmMni € Diademaproetuspmecursor _ QwKftgpjhorizon <2*4 m«M) TAZOULATT FORMATION Limestone €> , Equivalent to Psychopyga horizon

Nodular Limestone igggg ER REMLIA FORMATION 77 ME Shaly Limestone E BOUTISKAOUINE S iSS© FORMATION - OdontochHa horizon Interbedded Limestone and calcareous shale

m-. Shale with calcareous <3W3 nodules

Calcareous Shale

Shale

«— Kolihapeftis horizon Silty Shale •— Dicranurus horizon

249 DfOtapt rrxrprtynarncu* hortton

Harpes/Thysanopeftis horizon

7?iy®«rtOp«rtj» honzori

Psychopyge horizon

Issoumour Mrakib 100- TABOUMAKHLOUF FORMATION — Thysanopeltis horizon

Gerastos horizon Coral Cemlarges horizon 185 Gralnstone ^.Parwkyurvshomoa€ «^.£a6cwy* horizon horizon . QuadropsfTraveropyge horizon F E F L 0 B 0 R 0 R U M O M A A T D T F T1 1 1 A O B O L N N 50- 150-

T F - 2oc horizon A 0 Z R 0 M u A 1 T A 1 Drotops r O - megekmarticus T N horizon

horizon KonepnisSa horizon a

Equivalent Psychopyge/Comum Mraklblna - horizon 100 - ~ horizon

250 Figure 3-4. Map and stratigraphy of select localities of the southern Moroccan Tafilalt Basin. Locations of individual taxa indicated on individual stratigraphic columns with correspond to appropriate locations on the outcrop map. (Diagrams adapted from Fetah et al. (1998); Gibb (2005); Chatterton et al. (2006); Gibb & Chatterton (2007, 2010); McKellar & Chatterton (2009); Chatterton & Gibb (2010)).

251 u . Tafilalt Ha mar Laghdad Talawarite Basin Rissam

c

Outcrop Map 0 5 10 km • 1 I I

Stratigraphy Talawarite 40

Diademaprvetus praecursor

Shale

IntertMddad Limestone uMitoi hoftzoo and cateawx* shate ©

Stale wflh calcareous

Ptmoop* toftoyl found In tltu Deitomtm** horizon

252 Figure 3-5. Diagram showing the points of measurement about the cephalon and pygidium (follows Owens 1973). Ratio data were collected by measuring the distance between two points (e.g., length A = length of cephalon) and comparing it to a different measurement. Coordinate data were collected by comparing the position of landmarks (e.g., cephalic suture angle beta (P)) in relation to the length of the glabella (length Ai + A4).

253 254 Figure 3-6. Position of cephalic suture angle beta (P). Distances Aia- A4b (glabellar length (Figure 3-5, Ai + A4, sag.)), Aia - P, and A4b - ft were measured and calculated into landmark beta (fi) coordinates and Yp using Bookstein shape coordinate analysis. Landmark beta (fi) coordinates from all species were then graphed and generic level populations mathematically differentiated using a Hoteling's t-test. Two populations were found to be present for landmark beta (P) and are referred to as'narrow' and 'wide' morphologies. Overlap between populations exists as a result of variance in landmark position between specimens of the same generic populations. Coding of this character for the phylogenetic analyses was based on the specimens landmark position in relation to the derived equation (which side of the line separating the populations the specimen fell on), not which generic population the specimen was originally part of.

255 Position of Cephalic Suture Angle Beta

Population 1 Population 2

Narrow Morph

256 Figure 3-7. Position of cephalic suture angle gamma (y). Distances Au - A4b

(glabellar length (Figure 3-5, Ai + A4, sag.)), Aia - y, and A4b - y were measured and calculated into landmark gamma (7) coordinates X y and Yy using Bookstein shape coordinate analysis. Landmark gamma (y) coordinates from all species were then graphed and generic level populations mathematically differentiated using a Hoteling's t-test. Two populations were found to be present for landmark gamma (y) and are referred to as 'narrow' and 'wide' morphologies. Overlap between populations exists as a result of variance in landmark position between specimens of the same generic populations. Coding of this character for the phylogenetic analyses was based on the specimens landmark position in relation to the derived equation (which side of the line separating the populations the specimen fell on), not which generic population the specimen was originally part of.

257 Position of Cephali Suture Angle Gamma

Population 1 ° Population 2

• Q - ° oo *n

Narrow Morph

258 Figure 3-8. Position of cephalic suture angle delta (5). Distances Aia - A4b

(glabellar length (Figure 3-5, Ai + A4, sag.), Aia - <5, and A4b - S were measured and calculated into landmark delta (5) coordinates X y and Yy using Bookstein shape coordinate analysis. Landmark delta (5) coordinates from all species were then graphed and generic level populations mathematically differentiated using Hoteling's t-test. Two populations were found to be present for landmark delta (8) and are referred to as 'narrow' and 'wide' morphologies. Overlap between populations exists as a result of variance in landmark position between specimens of the same generic populations. Coding of the character for phylogenetic analyses was based on the derived equation (which side of the line separating the populations) a specimens landmark, not which generic population they were originally part of.

259 Position of Cephalic Suture Angle Delta

= Population 1 • Population 2 +• Population 3 rSE?

Narrow Morph f -t j <9 o ^3 • Sara*

260 Figure 3-9. Cephalic border shape. Arrow indicates area of referenced used to examine this character. A: flat B: inflated ('cushion-like')

Figure 3-10. Truncated anterior portion of cephalic border furrow. Arrow indicates area where character was examined. A: truncated (anterior border furrow absent) B: not truncated (anterior border furrow present)

261 A B C D

Figure 3-11. Shape of glabella as outlined by Snajdr (1980). A: sub-quadrate B: sub-cylindrical C: weakly violin shaped D: strongly violin shaped

Figure 3-12. Shape of eye platform. Arrow indicates area of observation for this character. A: none (Absent) B: posterior lobe only C: anterior and posterior lobes present

262 ABC

Figure 3-13. Shape of pygidium as outlined by Snajdr (1980). A: segment of circle B: rounded square C: rounded pentagon

A B

Figure 3-14. Shape of pygidial pleural ribs as outlined by Snajdr (1980). A: curved B: bent

263 / A B Figure 3-15. Shape of posterior margin of the pygidium. A: rounded (None) B: notched (Anteriorly indented) C: flat (horizontal)

264 Inteiproetus irrtermedius intermedius Inteiproetus pratux Inteiproetus venustus tE Inteiproetus xenon "pSS Comuproetus aoeps Comuproetus chluoad Comuproetus curtus Comuproetus comutus comutus Diademaproetus rudimentus- v Diademaproetus i880umouren8is».w. Diademaproetus cornigatus^w Diademaproetus auxiBarus... Diademaproetus praecursor Diademaproetus mohamedi Diademaproetus holzapfeli Tree statistics Diademaproetus lanausn •• Length: 102 steps Pontoproetus CX: 0.4902 tiuncatus- H> RX: 0.5098 Pontoproetus granulosus». R.C.: 0.6667

Figure 3-16. Cladogram depicting single phylogenetic hypothesis based on the results of exhaustive, branch and bound, and exhaustive analyses. In all analyses no character weighting or ordering was used. {#} Denotes Bremer support number for clade, all unmarked clades have value of {1}.

265 Inteiproetus intermedius intermedius 6(2) IK 1)25(1} Interproetus praliix Inteiproetus ,°l" venustus 16(2] 36(1) Interproetus xenon Comuproetus peraticus Comuproetus miimiam15(2) Cormiproetus chlupaci 1U0 27B 2*1 Comuproetus curtus iq31«p)J2J|2|2«tO! Comuproetus comutus cornutus IS(0jl?(2|2l<0) Diademaproetus i-riijoi rudimentusn.p 14J0J 20(O)22(1j)O{2} Diademaproetus issoumourensis-. »P. l7J0jl«t®]2lOl Diademaproetus jfitftomw corrugatusnsp. • Diademaproetus 6iO}fV27f2| *P»]2«0! auxiliarusn.sP. Diademaproetus praecursor I«0|22|S1 Diademaproetus mohamedi Diademaproetus hoizapfeli «2| Mft 12(11210) Diademaproetus 2M2|3IO) langus n. «P. 3-TMI Pontoproetus i -W0120( I) W\ 2«fO)»^0J truncatus n. »P. <#> = changed above Pontoproetus (#) = unique, uniform above granulosus n.sP. [#j = homoplasy outside "#" = homoplasy above {#} = homoplasy above & outside

Figure 3-17. Expanded view of cladogram (Figure 3-16), displaying changes in character states. Cladogram is a re-drafted version of the output from MacClade (Maddison & Maddison 2001) and its data matrix was initially analyzed using PAUP (Swofford 2002). Numbers preceding brackets denote characters, numbers within brackets denote change in character state. All changes shown are unambiguous.

266 Plate 3-1

Fig. 1,3-6. Diademaproetus praecursor Alberti, 1969 from ZGEE1, Timrhanrhart Formation, Zguilma, Tindouf Basin, southern Morocco (UA13693) (complete specimen). 1, dorsal view of cephalon and thorax; X2.0.3, dorsal view of thorax and pygidium; X2.1.4, anterodorsal view of cephalon; X2.2. 5, lateral view; XI.6. 6, anterolateral view of cephalon; X1.4. Fig. 2. Diademaproetus praecursor Alberti, 1969 from ZGEE1, Timrhanrhart Formation, Zguilma, Tindouf Basin, southern Morocco (UA13698) (isolated hypostome). 2, ventral view of hypostome; X9.3.

267 268 Plate 3-2

Fig. 1,3,5-6. Diademaproetus praecursor Alberti, 1969 from Taouz, Tafilalt Basin, southern Morocco (UA13692) (complete specimen). 1, dorsal view of cephalon and thorax; X2.3.3, dorsal view of thorax and pygidium; X2.4. 5, anterolateral view of cephalon; XI.9. 6, lateral view of cephalon; X2.6. Fig. 2,4, 7. Diademaproetus praecursor Alberti, 1969 from the Heliopyge/Kayserops horizon, El Otfal Formation, Jbel Issoumour, Ma'der Basin, southern Morocco (UA13697) (partial specimen). 2, dorsal view of cephalon and thorax; X2.8. 4, dorsal view of thorax and pygidium; X3.1. 7, anterolateral view of cephalon X3.5.

269 270 Plate 3-3

Fig. 1,3, 5,6. Diademaproetus mohamedi Chatterton et al., 2006 from ZGEE1, Timrhanrhart Formation, Zguilma, Tindouf Basin, southern Morocco (UA13383) (partial specimen). 1, dorsal view; X3.7. 3, anterior view of cephalon; X3.5. 5, dorsal view thoracic pleurae; X4.7. 6, dorsolateral view; X2.5. Fig. 2 & 4. Diademaproetus praecursor Alberti, 1969 from the Psychopyge horizon, Tazoulait Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA 13696) (partial specimen). 2, dorsal view; X4.0. 4, dorsolateral view; X2.7.

271 272 Plate 3-4

Fig. 1-7. Diademaproetus mohamedi Chatterton et al, 2006 from Harpes! Thysanopeltis horizon, El Otfal Formation, Ma'der Basin, southern Morocco (UA13699) (complete specimen). 1, dorsal view of cephalon; X2.5. 2, dorsal view of cranidium; X2.6.3, anterior view of cephalon; X2.8.4, anterolateral view of cephalon; XI.8. 5, lateral view; X2.3. 6, dorsal view of pygidium; X3.1. 7, dorsal view of thorax; X2.6.

273 274 Plate 3-5

Fig. 1-7. Diademaproetus rudimentus n. sp.Tazoula'it Formation, Taharajat, Ma'der Basin, southern Morocco. (UA13688) (complete specimen). 1, dorsal view; X1.7. 2, dorsal view of cranidium; X2.5. 3, dorsal view of glabella; X4.5. 4, dorsolateral view of cephalon; XI .7. 5, anterior view of cephalon; X2.2. 6, lateral view; XI.4. 7, dorsal view of pygidium; X3.6.

275 276 Plate 3-6

Fig. 1-6. Diademaproetus rudimentus n. sp. Tazoulai't Formation, Taharajat, Ma'der Basin, southern Morocco (UA13686) (complete specimen). 1, dorsal view of cephalon and thorax; X2.5. 2, ventral view of hypostome; X5.2. 3, dorsolateral view; X2.1. 4, dorsal view of pygidium; X2.6. 5, lateral view; XI.9. 6, anterior view of cephalon; X2.8.

277 278 Plate 3-7

Fig. 1-6. Diademaproetus corrugatus n. sp. Tazoula'it Formation, Taharajat, Ma'der Basin, southern Morocco (UA13681) (complete specimen). 1, dorsal view of cephalon and thorax; X2.0. 2, dorsal view of librigena; X5.1. 3, dorsal view of thorax and pygidium; X2.7. 4, anterior view of cephalon; X2.3. 5, lateral view of cephalon; X2.2. 6, anterolateral view of cephalon; X1.8.

279 280 Plate 3-8

Fig. 1-7. Diademaproetus auxiliarus n. sp. from Assa 2, Khebchia Formation, Assa, Tindouf Basin, southern Morocco (UA13689) (complete specimen). 1, dorsal view; X3.1. 2, dorsal view of cranidium; X3.3. 3, dorsal view of glabella; X6.1. 4, dorsolateral view; X2.4. 5, anterior view of cephalon; X3.5. 6, dorsal view of pygidium; X6.0. 7, lateral view; X2.7.

281 282 Plate 3-9

Fig. 1-3. Diademaproetus auxiliarus n. sp. from Assa 2, Khebchia Formation, Assa, Tindouf Basin, southern Morocco (UA13690) (complete specimen). 1, dorsal view of cephalon and thorax; X3.3. 2, dorsal view of thorax of pygidium; X3.6. 3, dorsolateral view of cephalon; X3.4. Fig. 4-7. Diademaproetus auxiliarus n. sp. from Assa 2, Khebchia Formation, Assa, Tindouf Basin, southern Morocco (UA13691) (partial specimen). 4, lateral view; X6.5. 5, dorsal view of cephalon; X3.5. 6, dorsal view of pygidium; X2.0. 7, ventral view of hypostome; X4.3.

283 284 Plate 3-10

Fig. 1-7. Diademaproetus langus n. sp. from the Diademaproetus horizon, El Otfal Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA13685) (complete specimen). 1, dorsal view; X2.4. 2, dorsal view of cranidium; X2.7. 3, dorsal view of glabella; X4.8. 4, anterolateral view of cephalon; X2.3. 5, lateral view; X2.3. 6, anterior view of cephalon; X2.5. 7, dorsal view of pygidium; X2.6.

285 286 Plate 3-11

Fig. 1-5. Diademaproetus langus n. sp. from the Diademaproetus horizon, El Otfal Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA13683) (complete specimen). 1, dorsal view; X2.5. 2, anterolateral view; X2.3. 3, anterior view of cephalon; X2.6.4, ventral view of hypostome; X4.0. 5, lateral view; XI.8.

287 288 Plate 3-12

Fig. 1-7. Diademaproetus issoumourensis n. sp. from the Psychopyge horizon, Tazoulait Formation, Jbel Issoumour, Ma'der Basin, southern Morocco (UA13271) (complete specimen). 1, dorsal view; X2.7. 2, dorsal view of cranidium; X3.2. 3, dorsal view of anterior lobe of glabella; X6.0. 4, anterolateral view; X2.5; 5, dorsal view ofpygidium; X5.4. 6, anterior view of cephalon; X3.1. 7, lateral view; X2.0.

289 290 Plate 3-13

Fig. 1-5. Diademaproetus issoumourensis n. sp. from the Psychopyge horizon, Tazoulai't Formation, Jbel Issoumour, Ma'der Basin, southern Morocco (UA13682) (complete specimen). 1, dorsal view of cephalon and thorax; X2.4. 2, anterolateral view of cephalon; X2.3.3, dorsal view of thoracic pleurae; X6.0. 4, dorsal view of thorax and pygidium; X2.9. 5, anterior view of cephalon X2.4. Fig. 6. Diademaproetus langus n. sp. from the Diademaproetus horizon, El Otfal Formation, Jbel Oufatene, Ma'der Basin, southern Morocco (UA13684) (complete specimen). 6, dorsal view; X2.4.

291 292 Plate 3-14

Fig. 1-6. Pontoproetus truncatus n. sp. from the Thysanopeltis horizon, Taboumakhlouf Formation, Zireg, Ma'der Basin, southern Morocco (UA13701) (complete specimen). 1, dorsal view; X1.8. 2, anterolateral view; X1.6. 3, anterior view of cephalon; X2.0.4, dorsal view of cephalon; X2.7. 5, lateral view; X1.4. 6, dorsal view of pygidium; X2.7. Fig. 7. Pontoproetus truncatus n. sp. from Timarzite, Ma'der Basin, southern Morocco. (UA13703) (complete specimen). 7, lateral view; X2.3.

293 294 Plate 3-15

Fig. 1-2. Pontoproetus truncatus n. sp. from Timarzite, Ma'der Basin, southern Morocco (UA13703) (complete specimen). 1, dorsal view; X2.7. 2, dorsal view of glabella; X4.4. Fig. 3-5. Pontoproetus truncatus n. sp. from the Thysanopeltis horizon, Zireg, Ma'der Basin, southern Morocco (UA13702) (complete specimen). 3, dorsal view of glabella; X3.9. 4, dorsal view of cephalon and thorax; X2.1. 5, dorsal view of thorax and pygidium X2.1.

295 296 Plate 3-16

Fig. 1-6. Pontoproetus grannulosus n. sp. from the Morocconites!Metacanthina equivalent horizon level, Tazoulai't Formation, Jbel El Mrakib, Ma'der Basin, southern Morocco(UA 13700) (complete specimen). 1, dorsal view;X2.6. 2, anterodorsal view; X2.2.3, anterior view of cephalon; X2.3. 4, dorsal view of cranidium; X4.1. 5, dorsal view of pygidium; X4.2. 6, lateral view; XI.8.

297 298 Table 3-1. Table outlining included and omitted taxa for the phylogenetic analysis. Images used in coding of specimens are indicated as the source, while material indicates the completeness of specimens being coded. Reasoning provides explanation as to why the taxon was either included or excluded from the analysis. All remaining species not outlined in the table were omitted either based on their lack of material (less than 50% of characters could be coded), poor documentation, or not being located in or around the Silurian or Devonian Rheic Ocean.

299 Snajdr 1980, pi. - 4 complete exoskeletons Included - type species of Interproetus XLV, fig. 19 and (articulated) • classified as ancestral genus to pi. XLVI, fig. 15. - 42 cranidia, 16 pygidia Cornuproetus and many morphologically r \' ~-A 4. '/--a* ,.' . V5. *• •" 'r--v -i" ; . (associated) similar genera by Snajdr (1980) Snajdr 1980, pi. - 43 cranidia and 16 pygidia Included • well preserved and documented XLVn, figs. 8 and (associated) Interproetus species 10. • one of the stratigraphically oldest species • *, -% ••-•.<, J »t;!j , j» of Interproetus, found in both Wenlock and *• , n * ;* • r\, * r's ^ V j' • "*'*' '• '• *•*»>••»"•' *" * •''* • Ludlovian (Silurian) strata brterproetus veriustus Snajdr 1980, pi. - complete cranidium Included - originally diagnosed as Astycoryphe? XLWI, figs. 6,7, (Alberti 1967) however re-designated to (B^Pl and 12. Tropicoryphe by Snajdr 1980 Snajdr 1980, pi. - 50+ cranidia, 20 librigena, and Included - extremely well preserved and XLVI, figs. 17, 19, 30 pygidia (associated) documented Interproetus species u Sa^lSpo o and 21. o Alberti 1969, pi. 7, -1 complete cephalon Included - type species of Cornuproetus (Gbld fig. 10. (articulated) and one thoracic ^Sa)v " » r i >'Jh *' . t * segment Cornuproetusperaticw Owens 1973, pi. 7, - 1 complete articulated Included - odd species of Cornuproetus which has a OWens, 1973 figs. 3-6. specimen 5+1 rhachial arrangement rather than the usual 4+1 - possibly a species of Interproetus (Snajdr 1980) Cornuproetus chlupaci Alberti 1969, pi. 8, - 12 cranidia and 1 pygidium Included - Early Devonian (Pragian) Cornuproetus Alberti, 1967 fig. 1-12. (associated) species found in Morocco - relatively complete and well preserved Cornuproetus curtus Snajdr 1980, pi. -10 cranidia and 19 pygidia Included - relatively complete and well documented (Barrande 1852) XLI, fig. 8-12. (associated) Cornuproetus species. Taciproetus adeps Basse 2010, pi. 2, - cranidia and pygidia Included - best preserved and documented newly (Lutke, 1977) fig. 11-12. (associated) proposed genus by Basse (2010) with very similar diagnosis as Cornuproetus Alberti 1969, pi. - complete cephalon with 5 Included - type species of Diademaproetus 16, fig. 5-6. articulated thoracic segments f »_ j? . •* *.i£k. *»«.,, 5 .->*«**: "* ? and 1 pygidium (associated) Chatterton el al. - numerous complete articulated Included - Moroccan Diademaproetus originally 2006, pi. 2, fig. specimens described by Alberti (1969), and re- 4 * v i , »« 1-4. described by Chatterton et al. (2006) - found throughout many of our localities. Diademaproetus mohar^tedi Chatterton el at. - 1 complete articulated Included - Moroccan Diademaproetus found at Chatterton et ai.,2006 2006, pi. 4, fig. specimen with partial pygidium Zguilma. 1-7. Duut&iitiproetus antatltisius ' Alberti 1969, pi. -1 cephalon and partial thorax Omitted - Moroccan Diademaproetus originally AJberti, 1969 16, fig. 7. and new described by Alberti (1969) material - Not complete enough, less than 50% coded. CO o C6rmqhq$h&4ftmutus Alberti 1969, pi. 7, - 2 incomplete cranidia Omitted - Incomplete, less than 50% coded marrakedftensis ,' ' ' figs. 11-12. Alberti. 1969 CornuprpetuS pernixpernix Alberti 1969, pi. - 2 incomplete cranidia Omitted - Incomplete, less than 50% coded fKeeel.1926) 10, fig. 5-6. Taciproetus taditurnus Alberti 1969, pi. 9, -1 partial cranidium Omitted - Incomplete, less than 50% coded (Kegel, 1926) fig. 12. Montanpraetus capuverta Basse 2010, pi. 2, - numerous partial cranidia and Omitted - Incomplete, less than 50% coded Basse, 2010 fig. 22-33. pygidia Montanprvetus rudrichteri Alberti 1969, pi. 9, - 2 partial cranidia Omitted - Incomplete, less than 50% coded fig. 10-11. fAlberti, 1967) Table 3-2. Results of character state statistical tests on a character by character, state by state basis. Use of Mann-Whittney U or Hoteling's t-test were dependent on data type. All tests were measured against a 95% confidence interval. When 2 or more character states are present, a Bonferroni correction adjusted p-value was used to test at the 95% confidence interval.

302 Character # #of States Test Tests P-value (%CC) l 2 Hotelling's T-test State 0 (Narrow Morph) vs. State 1 (Wide Morph) 4.381* 1012 (90.24%) 2 2 Hotelling's T-test State 0 (Narrow Morph) vs. State 1 (Wide Morph) 3.54*10"12 (81.61%) 3 3 Hotelling's T-test State 0 (Narrow Morph) vs. State 1 (Intermediate Morph) 2.47* lO"7 (Bonferroni corrected p-value = (73.13%) 0.0167 for 95%) State 0 (Intermediate Morph) vs. State 1 (Wide Morph) 5.98*10-7 (88%) State 0 (Narrow Morph) vs. State 2 (Wide Morph) 1.57*10"8 (90.7%) 4 2 Mann-Whittney U State 0 (< 0.28) vs. State 1 (> 0.28) 8.83*10-" 22 2 Mann-Whittney U State 0 (> 0.37) vs. State 1 (< 0.37) 4.79*10-5 24 2 Mann-Whittney U State 0 (> 1.10) vs. State 1 (< 1.10) 5.39* 10-4 Literature cited

Alberti, G. K. 1964. Neue Trilobiten aus dem Marokkanischen und deutschen Unter- und Mitteldevon. Senckenbergiana lethaea, 45, 115-132, pis. 16-17. Alberti, G. K. 1966. Note preliminaire sur quelques trilobites (en particulier de Proetides) du Silurien, du Devonien inferieur et du Devonien moyen du Maroc. Notes du Service Geologique du Maroc, 26, 55-69. Alberti, G. K. 1967. Neue obersilurische sowie unter- und mittle devonische Trilobiten aus Marokko, Deutschland und einigen anderen europaischen Gebieten. I and II. Senckenbergiana lethaea, 48, 463-479, 481-509. Alberti, G. K. 1969. Trilobiten des jiingeren Siluriums sowie des Unter- und Mitteldevons. I. Mit Beitragen zur Silur-Devon-Stratigraphie einiger Gebiete Marokkos und Oberfrankens. Abhandlungen der Senckenbergischen Naturforschenden Gesselschaft, 510, 1-692. Anstey, R. L. & Pachut, J.F. 2004. Cladistic and phenetic recognition of species in the Ordovician bryozoan genus Peronopora. Journal of Paleontology, 78(4), 651-674. Barrande, J. 1846. Notice preliminaire sur le systeme Silurien at les Trilobites de Boheme. Hirschfield, Liepzig, 97 pp. Barrande, J. 1852. Systeme Silurien du centre de la Boheme. 1 ere Partie. Recherchespaleontologiques, IrTrilobites, 935 pp. Basse, B. M. 2010. Proetoidea Hawle & Corda, 1847 und andere Trilobita aus unterdevonischen Herzynkalken der westlichen Harzgeroder Faltenzone (Zlichovium, Dalejum; sudwestlicher Harz, Rhenoherzynikum): Cornuproetinae Richter, Richter & Struve in Moore, 1959 und Eremiproetinae Alberti, 1967b (1). Freiberger Forschungshefte, 18,1-67. Becker, T.R., Bockwinkel, J., Ebbighausen, V., Aboussalam, S.Z., El Hassani, A. & Niibel, H. 2004a. Lower and Middle Devonian stratigraphy and faunas at Bou Tserfine near Assa (Dra Valley, SW Morocco). Pp. 90-100 in A. El Hassani (ed) Devonian Neritic-Pelagic Correlation and Events in the Dra Valley (Western Anti-Atlas, Morocco). Subcommission on Devonian Stratigraphy. International Meeting on Stratigraphy, Rabat, 1-10 March 2004. Documents de I'Institut Scientifique, 19. Becker, T.R., Jansen, U., Plodowski, G., Schindler, E., Abzoussalam, S.Z. & Weddige, K. 2004b. Devonian litho- and biostratigraphy of the Dra Valley area - An overview. Pp. 3-18 in A. El Hassani (ed) Devonian Neritic-Pelagic Correlation and Events in the Dra Valley (Western Anti-Atlas, Morocco). Subcommission on Devonian Stratigraphy. International Meeting on Stratigraphy, Rabat, 1-10 March 2004. Documents de I'Institut Scientifique, 19. Belka, Z., Kaufmann, B., & Bultynck, P. 1997. Conodont-based quantitative biostratigraphy for the Eifelian of the eastern Anti-Atlas, Morocco.

304 Geological Society of America Bulletin, 109(6), 643-651. Bookstein, F. L., Chernoff, B. C., Elder, R. L., Humphries, J. M., Smith, G. R., and Strauss, R. E. 1985. Morphometries in evolutionary biology. Academy of Natural Sciences of Philadelphia Special Publication, 15, 227pp. Bultynck, P. 1987. Conodonts from a potential Eifelian-Givetian Global Boundary Stratotype at Jbel Ou Driss, southern Ma'der, Morocco. Bulletin de I 'institut Royal des Sciences Naturelles de Belgique, Sciences de la Terre, 57, 149-181. Bultynck, P. & Hollard, H. 1980. Distribution Comparee de Conodonts et Goniatites Devoniens de Plaines du Dra, de Ma 'der et du Tafilalt (Maroc). Leuven University Press, Belgium, 73 pp. Bultynck, P. & Walliser, O.H. 2000. Devonian boundaries in the Moroccan Anti-Atlas. Courier Forschumgsinstitut Senckenburg, 225, 211-226. Chatterton, B.D.E. & Speyer, S.E. 1997. Ontogeny. Pp. 173-247 In: Whittington, H.B., et al. (Eds.), Arthropoda 1, Trilobita. Part O. (revised) Treatise on invertebrate paleontology. University of Kansas, Lawrence, Geological Society of America, Boulder, Colorado. Chatterton, B. D. E., Fortey, R., Brett, K., Gibb, S., and McKellar, R. 2006. Trilobites from the upper Lower to Middle Devonian Timrhanrhart Formation, Jbel Gara Zguilma, southern Morocco. Palaeontographica Canadiana, 25, 177 pp. Cronier, C., Auffray, J.C., & Courville, P. 2005. A quantitative comparison of the ontogeny of two closely related phacopid species of the Upper Devonian. Lethaia, 38, 1-14. Cr6nier C., Bignon, A., & Francois, A. 2011. Morphological and ontogenetic criteria for defining a trilobite species: The example of Siluro-Devonian Phacopidae. Comptes Rendus Palevol, 10, 143-153. El Hassani, A. (ed.). 2004. Devonian neritic-pelagic correlation and events in the Dra Valley (western Anti-Atlas, Morocco). Subcommision on on Devonian Stratigraphy. International Metting on Stratigraphy, Rabat, 1-10 March 2004. Documents de I'Institut Scientifique, 19, 1-100. Erben, H. K. 1952. Proetus (Cornuproetus) corrugatus Erben 1952 als objektives Homonym von Proetus corrugatus van Ingen 1901. Neues Jahrbuch fur Geologie und Palaontologie, Monatshefte, 10, 449-450. Fetah, S. E., Bensa'id, M., and Dahmani, M. 1988. Carte Geologique Maroc: Todhra-Ma'der (Anti-Atlas oriental, zones axiale et peripherique Nord du Sud). Editions de Service Geologique de Maroc, Notes et Memoires No. 243. Royaume du Maroc. Ministere de l'Energie et des Mines, Rabat. Fortey, R. A. & Owens, R. M. 1975. Proetida - a new order of trilobites. Fossils and Strata, 4, 227-239.

305 Gibb, S. 2005. Some proetids (Class Trilobita) from the Lower to Middle Devonian of southern Morocco. M.Sc. thesis, University of Alberta, Edmonton Alberta, Canada. 21 lpg. Gibb, S. & Chatterton, B. D. E. 2007. Timsaloproetus new genus (Proetida: Trilobita) and included species from Lower and Middle Devonian strata of southern Morocco. Journal of Paleontology, 81(2), 352-367. Gibb, S. & Chatterton, B. D. E. 2010. Gerastos (Order Proetida; Class Trilobita) from the Lower and Middle Devonian of the Southern Moroccan Anti-Atlas region. Palaeontographica Canadiana, 30, 89pg. Giebel, C. 1858. Die silurische Fauna des Unterharzes. Abhandlungen des Naturwissenschaftlichen Vereines der Provinzen Sachsen und Thuringen, 1,261-332, Taf. 1-7. Goldfuss, A. 1843. Systematische Ubersicht der Trilobiten und Beschreibung einiger neue Arten derselben. Neues Jahrbuch Jur Mineralogie, Geognosie, Geologie, und Petrefakten-Kunde, Stuttgart, 1843, 537-567. Hammann, W. 1976. Trilobiten aus dem oberen Caradoc der ostlichen Sierra Morena (Spanien). Senckenbergiana lethaea, 57, 35-85. Hammer, 0.2010. PAST - PAleontological STatistics ver.1.99. Hass, W. 1968. Trilobiten aus dem Silur und Devon von Bithynien (NW-Tiirkei). Palaeontographica Abteilung A, 130, 60-207. Hollard, H. 1974. Recherches sur la stratigraphie des Formations du Devonien Moyen, de l'Emsian superieur au Frasnien, dans le Sud du Tafilalt et dans le Ma'der (Anti-Atlas oriental). Notes du Service Geologique de Maroc, 36(264), 7-68. Hollard, H. 1978. Correlations entre niveaux a brachiopodes at a goniatites au voisinage de la limite Devonien inferieur- Devonien moyen dans les plaines du Dra (Maroc presaharien). Newsletters on Stratigraphy, 7(1), 8-25. Hughes, N.C., Minelli, A., and Fusco, G. 2006. The ontogeny of trilobite segmentation: a comparative approach. , 32(4), 602-627. ICZN. 1999. International code of zoological nomenclature. The International Trust for Zoological Nomenclature. Jansen, U., Becker, G., Plodowski, G., Schindler, E., Vogel, O. & Weddige, K. 2004. Pragian and Emsian near Aouinet Torkoz (SW Dra Valley, Morocco). Pp. 75-84 in A. El Hassani (ed) Devonian Neritic-Pelagic Correlation and Events in the Dra Valley (Western Anti-Atlas, Morocco). Subcommission on Devonian Stratigraphy. International Meeting on Stratigraphy, Rabat, 1-10 March 2004. Documents de I'Institut Scientifique, 19. Kegel, W. 1926. Unterdevon von bohmischer Facies (Steinberger Kalk) in der Lindener Mark bei GieGen. Abhandlungen der Preussischen Geologischen Landesanstalt, N.F., 100: 1-77.

306 Kowalski, H. 1975 Cornuproetus (Diademaproetus) aus dem Eifelium (Mittle devon) der Eifel. Der Aufschluss. Zeitschrift fur die Freunde der Mineralogie und Geologie, 26, 97-101. Maddison, D. R. and Maddison, W. P. 2005. MacClade 4: Analysis of Phylogeny and Character Evolution. Version 4.08. Sinauer Associates, Sunderland, Massachusetts. McKellar, R. and Chatterton, B. D. E. 2009. Early and Middle Devonian Phacopidae (Trilobita) of southern Morocco. Paleontographica Canadiana, 28, llOpg. Morzadec, P. 2001. Asteropyginae trilobites from the Devonian of the Anti-Atlas (Morocco). Palaeontographica Abteilung A, 262, 53-85. Owens, R. M. 1973. British Ordovician and Silurian Proetidae (Trilobita). Palaeontographical Society Monographs, 127, 1-98, pis 1-15. Philip, G. 1991. Political: Africa, Philip's World Atlas. Reed International Books Ltd., London. Prevosti, F.J., & Chemisquy, M.A. 2010. The impact of missing data on real morphological phylogenies: influence of the number and distribution of missing entries. Cladistics, 26, 326-339. Pribyl, A. 1971. Proetidni trilobiti z novych sberu v ceskem siluru a devonu. cast III. Casopis Narodniho Muzea, oddil Pfirodovedny, 140, 81-89. Richter, R. & Richter, E. 1919. Proetiden aus neueren Aufsammlungen im vogtlandischen und sudetischen Oberdevon. Senckenbergiana, 1, 97-130. Richter, R. & Richter, E. 1949. Die Trilobiten der Erdbach-Zone (Kulm) im Rheinischen Schiefergebirge und im Harz. I. Die Gattung Phillibole. Senckenbergiana, 30, 63-94. Richter, R., Richter, E. 1959. Grundlagen fur die Beurteilung und Einteilung der Scutelluildae (Tril.). Senckenbergiana lethaea, 37, 79-124. Snajdr, M. 1977. New genera of Proetidae (Trilobita) from the Barrandian, Bohemia. Vestnik Ustredniho ustavu geologickeho, 52, 293-297. Snajdr, M. 1980. Bohemian Silurian and Devonian Proetidae (Trilobita). Rozpravy Ustredniho ustavu geologickeho, 45, 1-324. Stafford, E. S. and Leighton, L. R. 2011. Vermeij Crushing Analysis: Anew old technique for estimating crushing predation in gastropod assemblages. Palaeogeography, Palaeoclimatology, Palaeoecology, 305, 123-137. Swofford, D. 2002. PAUP*4.0bl0 (Altivec): Phylogenetic Analysis Using Parsimony. Sinauer Associates, Inc. Wagner, P. J. 2000. Exhaustion of morphological character states among fossil taxa. Evolution, 54(2), 365-386. Wiens, J. J. 2003. Incomplete data, missing taxa, and phylogenetic accuracy. Systematic Biology, 52(4), 528-538. Whittington, H. B. and Kelly, S. R. A. 1997. Morphological terms applied to Trilobita. Pp. 313-329 in Moore R. C. and R. L. Kaesler (eds.) Treatise on invertebrate paleontology, part O, Arthropoda 1, Trilobita revised, v. 1:

307 Introduction, Order Agnostida, Order Redlichiida. Geological Society of America, Boulder, and University of Kansas Press, Lawrence.

308 Chapter 4: Conclusion

Synthesis

" If we knew what we were doing, it wouldn't be called Research" (A. Einstein).

Throughout this thesis numerous approaches were explored and methods tested, many of which did not make it into the final product. Examination of select tropidocoryphid and comuproetid genera from the Lower and Middle Devonian strata of southern Morocco's Anti-Atlas region has resulted in the description of one new genus and eleven new species. Comparison of the new species with similar species from larger palaeogeographic scales (e.g., the Devonian Rheic

Ocean), by means of phylogenetic analyses, has provided both a better understanding of species and genus level diagnostic characters and the relationships between the examined taxa. Furthermore, the relatively well supported and intuitive phylogenetic results suggest that the use of statistically supported characters and character states in phylogenetic analyses is beneficial in determining the relationships of closely related and morphologically similar taxa.

Members of the subfamily Tropidocoryphinae (e.g. Astycoryphe and

Tropidocoryphe) were found in the Emsian and Eifelian strata of the Tindouf and

Ma'der basins. Only one specimens of Astycorphe (Astycoryphe ditropidia) was found, in the Philonyx/Quadrops horizon at Jbel Oufatene. Tropidocoryphe was found in: Assa (Tropidocoryphe amuri); Zguilma, in the trilobite bearing horizons

ZGEE1 and ZGEE3 (Tropidocoryphe amuri); Jbel Oufatene, in the Philonyx/

309 Quadrops horizon (Tropidocoryphe lahfirensis) and the Diademaproetus horizon

(Tropidocoryphe sculptaridgiata)\ and Taharajat (Tropidocoryphe taharajatensis).

Results from the phylogenetic analysis of Astycoryphe, Tropicoryphe,

Bojocoryphe, Longicoryphe, Tropidocoryphe, Erbenicoryphe, and Pterocoryphe suggest that Astycoryphe is the most primitive genus, with Longicoryphe,

Tropidocoryphe, Erbenicoryphe, and Pterocoryphe all being more derived. Lack of morphological distinction between Tropicoryphe, Bojocoryphe, and

Astycoryphe suggests that both Tropicoryphe and Bojocoryphe are junior synonyms of Astycoryphe. Furthermore, results of the phylogenetic analysis support the hypothesized Longicoryphe-Erbenicoryphe-Pterocoryphe relationship proposed by Feist & Clarkson (1989).

Members of the subfamily Cornuproetinae (primarily Diademaproetus) were found throughout the Tindouf, Ma'der, and Tafilalt basins. In the Tindouf

Basin, Diademaproetus praecursor was found in all three trilobite-bearing horizons (ZGEE1 - ZGEE3) at Zguilma and the locality FZ1, while

Diademaproetus mohamedi was found found only in ZGEE3 at Zguilma. In the

Ma'der Basin, Diademaproetus and Pontoproetus were found at Jbel Oufatene, in the Diademaproetus horizon (.Diademaproetus langus and D. praecursor);

Taharajat (Diademaproetus rudimentus and D. currugatus); Jbel Issoumour, in and just below the Psychopyge horizon (Diademaproetus issoumourensis and D. praecursor), in the Heliopyge/Kayserops horizon (Diademaproetus praecursor), and in the Lobopyge and Paralejurus horizons (.Diademaproetus praecursor)-,

310 Timarzite (Pontopwetus truncatus)', bou DIb, in the Harpes/Thysanopeltis horizon

(Diademaproetus mohamedi); Jbel El Mrakib, in the stratigraphically equivalent to the Morocconites/Metacanthina horizons (Pontopwetus granulosus); Jbel

Zireg, in the Thysanopeltis horizon {Pontopwetus truncatus); and at El Achana, in the Kolihapeltis and Dicranurus horizons (Diademaproetus rudimentus). In the

Tafilalt Basin, Diademaproetus praecursor was found at Talawarite, in the

Gerastos horizon, and Taouz. Results of phylogenetic analyses involving

Cornuproetus, Diademaproetus, and Pontopwetus suggest that Cornuproetus is the most primitive genus, with Pontopwetus being more derived than

Diademaproetus. Lack of morphological distinction between Taciproetus,

Montanproetus, and Cornuproetus, and the phylogenetic instability that

Taciproetus and Montanproetus bring to the Cornuproetus-Diademaproetus relationship, promotes both Taciproetus and Montanproetus as junior synonyms of

Cornuproetus.

The phylogenetic analyses completed throughout this thesis were by no means 'easy,' and often very time consuming. Lack of suitable characters and character states, for both the Tropidocoryphinae and Cornuproetinae, promoted the construction and statistical testing of both new and previously discussed proetid characters and character states. Furthermore, character exhaustion and a high abundance of incomplete taxa (taxa which were less than 50% coded) resulted in phylogenetic instability. However, as the results of the final phylogenetic analyses demonstrate, the effects of character exhaustion on a

311 phylogenetic analysis can be overcome when statistically tested characters and

relatively complete (taxa which are more than 50% coded) taxa are examined.

Furthermore, these results encourage the use of properly assessed quantitative

characters and cladistic analysis when examining and describing morphologically similar taxa. Failure to follow such methods creates confusing diagnoses and

over-splitting of taxa, hampering the assignment of new specimens to the proper genus and the understanding of the taxon relationships. Without a unified approach towards examining trilobite taxa and their relationships, proper

understanding of their evolution cannot be achieved.

Future work

As is the case with most scientific research, answering one question typically leads to many more questions being asked. As Gibb (2005) stated, much work still needs to be completed on Moroccan proetids. Although this thesis was derived to accomplish that goal, there still remain numerous Moroccan proetid taxa to be examined and described. Of particular importance are the remaining genera of the Cornuproetinae, primarily those which were once considered subgenera of Cornuproetus (Alberti 1969). While Moroccan examples of these genera do exist, much better preserved and more complete specimens are undoubtedly available and would likely add important information to the evolutionary relationship of Cornuproetus and Diademaproetus. Furthermore, expansion of the phylogenetic methodology and hypotheses presented in this

312 thesis to include more taxa from a larger palaeogeographic area would be useful in examining higher level taxon relationships and Devonian trilobite evolution.

313 Literature cited

Alberti, G. K. 1969. Trilobiten des jiingeren Siluriums sowie des Unter- und Mitteldevons. I. Mit Beitragen zur Silur-Devon-Stratigraphie einiger Gebiete Marokkos und Oberfrankens. Abhandlungen der Senckenbergischen Naturforschenden Gesselschaft, 510, 1-692. Feist, R. and Clarkson, E. N. K. 1989. Environmentally controlled phyletic evolution, blindness and extinction in Late Devonian tropidocoryphine trilobites. Lethia, 22, 359-373. Gibb, S. 2005. Some proetids (Class Trilobita) from the Lower to Middle Devonian of southern Morocco. M.Sc. thesis, University of Alberta, Edmonton Alberta, Canada. 21lpg.

314 Appendix 1

Numerical calculations for all examined specimens of the Tropidocoryphinae.

315 <(A2 + A3)-A5) A! AS: Aj(A2 Beta(§) Population; Xy Gamma (y) Population: XS j YS Delta (5) Population Z : W 0.44 0.33 0.32 0.68 0.48 0.27 0.33:0.78 0.57 1.07 0.40-0.26 0.28; 0.73 0.61 1.59 n/a n/a n/a ! n/a 0.64 .4 0.42! 0.34 p.33lpJ9 0.64 ; 1.29 0.4310.28 0.29:0.76 0.32^0.74 0.66 1.16 0.31i0.21 0.32 i 0.66 0.69 1.06 • 0.3 i 0.15 0.32:0.65 ; 0.35 0.24 0.34 0.74 0.67 1.03 , 0.34 0.66 0.35 i 0.22 0.35 10.67 0.68 1.03 0.37* 0.21 0.70 1.17 0.70 1.18 CO 0.72 1.22 o> . n/a 0.55 1.14 ^4110.26 0.48 1.22 0.46! 0.32 0.2910.64 0.72 1.01 0.38j0.34 0.73 0.96 0.39 i 0.19 0.301 0.66 0.70 0.89 0.33 i 0.25 0.32 0.67 0.70 : 1.15 0.68 > 1.24

0.3510.32 0.3010.71 0.73 ; 1.16 0.65:0.56 0.29! 0.76 0.72 . 1.29 0.62! 0.44 0.35 0.72 0.66 1.39 0.39; 0.24 0.38•0.65 0.69 1.25 0.45 0.31 0.31 0.79 0.43I 0.32 0.53 0.32 0.34 ? 0.72 0.58 : 1.25 n/a n/a 0.42 i 0.28 0.63 Ml a/ a 0.43 0.31 . . . . n/a 0.44 0.3 0.62 . 1.22 0.62 ' 1.25 Appendix 2

MacClade 4.06 (Maddison and Maddison, 2001) derived data matrix for the examined species of the Tropidocoryphinae.

317 [ CM CM CM CO CM T- CM CM CM T- C" T- - O CM CM x- CO CM CM O O" r-

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I oo o o o o o o e» o- T- O CM O O O t- T- T- o o O T-

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r* T" T- CO T"» T- COCMOCMOT-T-OOOCMCM CM CO

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318 Appendix 3

Numerical calculations for all examined specimens of the Comuproetinae.

319 |j I18i & 1 !• 8 I I l~ 1 8j Ij i. i 8 I i-1 I & 11:111 V i Sj ii 5 5= 5* i 5 5- 5 51 5; i 5 i s i 5 i i i sj 5! i* si s s

»: a a 3S. *X JS S.S2; *; *> | * 4S b; s' 6 "sT i a B s j| 3: 5 5: Ii I e, t : 9j ' f ' t * ! I* •i SI I«;« 5 «» « S ? 5 S ! 3! !! t ca aSi 8,4 «

«: « f a «. ta « «! «' «! •• I* B! 8' 8j ! 8 ft' • * B 8 8. • 8! 8: 8- Si S< 3 8 C Si «! ft nai 8 runs, x' Sii a: it! r- a; s k fi ft ill.! i rtilliii 11 I I I I !; «| il || J|» 5 ; E j [; 11.1};| Ij 1111,11 1 am a a io; iHEEfiiiic e{ a «} •; « o o « « *1 8

Es liIi i.R1 1;8i 3 Ii ;I iS i

1 : 5' 2 E S S- 2 "' 5s = 5'! 5 s * = S 8 3 S. *: 2: 5? §- s. I 3 r; i.I Rgi 15 § I 5I *? §2 ii ?! ? ?:!' §" 5i| Ij §; 11 *! 8* Si 81 =1 8 ; s 5 R: ;i 8 a 8 ?• 8 5; 9, 8 8 B; 9) 8! 8 *• 8; 8 8 C

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ft Mil\ 11iuhni11 M i 1S! i i ' I «I

J II181i11.1 i I ij l^iji 11«' i 11 SI I' SI sj I 8 l! 51 ij i| 5; 3; i; i, i, i i, g i| i ii ij i j; is i i ii 1 ij ii i i

320 use

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I I ------| I t {* I i - i - - - - - it I - I - - i - - !-

• • » • .• 1 s •* 1 >• * • .p 'a I ' IV'II IINIjvillilili i-fljl 11 S11 ;8 0 • '» » (O •» 1 » ® 0 '» • P FT IP • ; • » "• >0

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i i I i 3 - - I ,} - - - " - - » I" j- - - I - i- !- I ;-

s :B s :a 2 * * B 8 » !B IS ;* jg 11111 ll:i 1 ! » ii* |S

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Is 11 i •* • .« i I li t .* * * I * ? li-iH i»•»; i»i i« |i j I"!!'! IliliS

IIIHHi i ;l'«I.1 -l i 11« 11 i 1 n J Kill •».«.»:»|i p 1 »Ji 1 Appendix 4

MacClade 4.06 (Maddison and Maddison, 2001) derived data matrix for the examined species of the Cornuproetinae.

322 I o o CM CM CMCMOCMO-OJCOT-T- I o o CM O 1-T-OCMO-Ot-T-T- I ° T- 1— 1— 1— T— T- T- O T— IC" o o O O o o C^- T- O O O T-'O o o I eg CM CM o o O O O T- C^- CM IO O CM CM T- T- T- lc" o T~ O o o OOOOC-r-Or-T— 1° o o 0-o ooooooooo 1° o O o O O -i- 1° o O o O o o o o o;o> O CM O O I o o O o o o f- O T- o O T- 1°" CV. Cv. CM c^- O* 1- T- CM O CM I- CM CO I CM c^- CM o- C^- CM CM ; CM CM O CM CM r-

\o- o» ; c^- cw. o- O O o o o- 0- i- O o o CM CM CM CM O CM CM O CM CM CM CM O CO CO I— CM T- O 1- O O O IT- T- O O o CM CM CM CM CM CM CM CM CM CM CM O 1- T- CM T- T- T- r- O- O O O i- r- O T- T— 1T" 1° T- C" i- O i- r~ O o o I CM 0» O CM CM CM CM CM CM CM CM CM r- lc- o O C^- CM CM CM CM CM CM F O CM O T- c^' O O ' O O O O O

ICO CM CM CO CO CO CM CM CM CM CM I C^- FS.. ' -r-

1- o o o i— O O T- -r- r- R~ T"" T" I" T- T- O O

11- CM CM CM O 1- o o o T- O O CM O O o o © o o o o o o I O o O *- i— t— , f- r» r* r" r~ 1° o O T- O C- O 1- CM O CM I o o O 1- O t- r- O O i- O o o o o i- i- O'

323