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Home , Douvilleiceras, Linguloidea, Naticarius hebraeus, Notocochlis, Pulchelliidae, Rosablanca Formation

PALEOBIOLOGICAL IMPLICATIONS OF THE POST-PALEOZOIC FOSSIL RECORD OF THE LINGULIDE BRACHIOPODS AND ASSOCIATED FAUNAS

ALEXIS ROJAS

A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA

2017

To my wonderful grandparents, who taught me to be resourceful and resilient

ACKNOWLEDGMENTS

I thank the members of my dissertation committee for their time and extreme patience. I am indebted to Liang Mao, whose graduate courses inspired me to adopt a spatially explicit perspective to study the fossil record. I also express my sincere appreciation and gratitude to my advisor, Michał Kowalewski, for his academic guidance and personal support.

I am also grateful to a long list of people affiliated with museums without whom this dissertation would not have been possible: Roger Portell (Florida Museum of

Natural History) for his assistance during multiple surveys of the invertebrate paleontology collections, Sean Roberts (Florida Museum of Natural History) and Mark

Florence (National Museum of Natural History) for their assistance with digital images of specimens, Alfred Dulai (Hungarian Natural History Museum) for sharing information from his ongoing research on lingulides, Eric Simon (Royal Belgian Institute of Natural

Sciences) for providing museum data, and Daniel Levin (National Museum of Natural

History) for his assistance during the visit to the paleobiology collections.

I am very grateful to Dr. Etayo-Serna (Ingeominas), Georgina Guzman

(Universidad Industrial de Santander), Javier Luque (University of Alberta), Jorge

Moreno (Smithsonian Tropical Research Institute), and the family Sandoval- Rueda for their support during field seasons in the town of Zapatoca.

I am indebted to Michael Sandy (University of Dayton) for sharing his expertise on brachiopod taxonomy, providing the serial sections of Sellithyris elizabetha nov sp. and field data on Mexican and European taxa, and Julie Hays for generating biometric information.

I thank the contributors to the PaleoDB who collected Albian ammonoid data, especially Austin Hendy, Matthew Clapham and Loïc Villier, Martin Rosvall (Umeå

University) for his comments on the resolution of the Infomap algorithm, and

Christopher Scotese (http://www.scotese.com) for providing the PALEOMAP and

PointTracker Software.

I am very grateful to my collaborators for their valuable input: Chapter 2 Michael

Sandy; Chapter 3 Gregory P. Dietl and Austin Hendy; Chapter 4, Michał Kowalewski and Roger Portell; Chapter 5 Michał Kowalewski, Pedro Patarroyo and Peter Bengtson.

I also thank reviewers for their comments and constructive criticism: Chapter 4 Patricia

Kelley, Elizabeth M. Harper and David A.T. Harper; Chapter 5 Wolfgang Kiessling,

Dieter Korn, and an anonymous reviewer, and editorJudith Totman.

I am very grateful to my wife, Elizabeth Sandoval, whose constant encouragement was vital in making this dissertation a reality.

This dissertation research was funded in part by the Jon L. and Beverly A.

Thompson Endowment Fund. Additional funding support was provided by a UF-

Department of Geological Sciences Teaching Assistantship. Field Survey was funded in part by the mithsonian Tropical Research Institute. Funding support from a Florida

Museum of Natural History Travel Award and Drs. Emily H. and Harold E. Vokes

Grants-in-Aid for Invertebrate Paleontology Collection-based Research is appreciated.

TABLE OF CONTENTS

page

ACKNOWLEDGMENTS ...... 4

LIST OF TABLES ...... 8

LIST OF FIGURES ...... 9

LIST OF ABBREVIATIONS ...... 11

ABSTRACT ...... 13

CHAPTER

1 INTRODUCTION: THE POST-PALEOZOIC FOSSIL RECORD OF THE LINGULIDE BRACHIOPODS ...... 15

2 LOWER CRETACEOUS BRACHIOPODS FROM COLOMBIA (SOUTH AMERICA): BIOSTRATIGRAPHIC SIGNIFICANCE AND PALEOGEOGRAPHIC IMPLICATIONS ...... 19

Introduction ...... 19 Materials and Methods ...... 20 Sample Collection ...... 20 Data Analysis ...... 21 Systematic Palaeontology ...... 23 Discussion ...... 28

3 A PROXY FOR SITE-SELECTIVE OF DRILLING PREDATORS BASED ON SPATIAL POINT PROCESS MODELING ...... 40

Introduction ...... 40 Materials and Methods ...... 41 Data Collection: Museum Survey ...... 41 Point Pattern Construction ...... 42 Density Analysis of Drillholes ...... 43 Spatial Clustering Analysis of Drillholes ...... 44 Results ...... 45 Discussion ...... 46

4 THE POST-PALEOZOIC FOSSIL RECORD OF DRILLING PREDATION ON LINGULIDE BRACHIOPODS ...... 54

Introduction ...... 54 Materials and Methods ...... 55

Data ...... 55 Methods ...... 57 Results ...... 59 Discussion ...... 62

5 GLOBAL BIOGEOGRAPHY OF ALBIAN AMMONOIDS: A NETWORK-BASED APPROACH ...... 71

Introduction ...... 71 Materials and Methods ...... 72 Data ...... 72 Network Construction and Partitioning ...... 73 Results ...... 74 Discussion ...... 76

6 CONCLUSIONS ...... 87

APPENDIX

A BIOMETRIC DATA ON SELLITHYRIS COMPILED IN THIS STUDY ...... 89

B DRILLING DATA ON CENOZOIC LIROPHORA COMPILED IN THIS STUDY ...... 96

C BODY SIZE DATA OF THE LINGULIDE SPECIMENS SURVEYED AT THE FLORIDA MUSEUM OF NATURAL HISTORY ...... 103

LIST OF REFERENCES ...... 108

BIOGRAPHICAL SKETCH ...... 132

LIST OF TABLES

Table page

2-1 Comparison of Sellithyris elizabetha and other Valanginian forms ...... 38

2-2 Results of the permutational MANOVA ...... 39

3-1 Drillling data on museum samples of Lirophora latirilata ...... 52

3-2 Parameters used to calculate the optimum bandwidth (hopt) ...... 53

4-1 Quantitative data for the localities with drilled specimens...... 69

4-2 Statistical significance of the drilling frequencies...... 70

5-1 Comparison of the Infomap communities in the network GP...... 85

5-2 Comparison of the networks GP and GP-benthos ...... 85

5-3 Centrality scores for the nodes in the network GS...... 86

LIST OF FIGURES

Figure page

2-1 Geographic location of the town of Zapatoca in the Eastern Cordillera of Colombia...... 32

2-2 Stratigraphic section of the Upper Rosablanca Formation in the Laguna del Sapo locality, Zapatoca...... 33

2-3 Bivariate plot of shell length and width of Lingularia sp. from the Upper Rosablanca Formation and other Cretaceous lingulides worldwide...... 34

2-4 Lower Cretaceous brachiopods from Colombia...... 35

2-5 Serial sections through Sellithyris elizabetha nov. sp. from the Lower Cretaceous of Colombia ...... 36

2-6 Morphological and taphonomic data of Sellithyris elizabetha nov. sp. from the Lower Cretaceous of Colombia ...... 37

3-1 Point pattern of naticid-like drillholes on Pliocene and Pleistocene Lirophora latilirata from the Atlantic Coastal Plain...... 49

3-2 Kernel density estimation and hotspot mapping of naticid-like drillholes on Pliocene and Pleistocene Lirophora latilirata from the Atlantic Coastal Plain. .... 50

3-3 K function for naticid-like drillholes on Lirophora latilirata from the Atlantic Coastal Plain...... 51

3-4 Empirical K-function values subtracted by its expected value K(r)−πr2 for predatory drillholes on Lirophora latilirata...... 51

4-1 Schematic diagram of a conjoined lingulide shell in dorsal view indicating the drillhole locations...... 65

4-2 Scatter plot of lingulide size and drillhole size ...... 66

4-3 Drilled specimens of the Pliocene Glottidia inexpectans...... 67

4-4 Quantitative summary of drilling predation on post-Paleozoic lingulide brachiopods...... 67

5-1 Workflow diagram indicating the procedures implemented in the analysis of the global records of Albian ammonoids...... 80

5-2 Bipartite occurrence network of Albian ammonoids G...... 81

5-3 Geographic network GP and Infomap bioprovinces ...... 81

5-4 Comparison of the Infomap results with other estimated grouping structures. ... 82

5-5 Ammonoid occurrence data per grid cell clustered using the unweighted pair- group arithmetic average method ...... 82

5-6 Latitudinal distribution of nodes in the projected network GP grouped by Infomap bioprovince...... 83

5-7 Geographic network GP-benthos and Infomap bioprovinces...... 83

5-8 Taxonomic composition of the Infomap bioprovinces delineated in the projected network GP-benthos...... 84

LIST OF ABBREVIATIONS aA anterior adductor muscle scar aO anterior oblique muscle

AR Austral Realm

ASr Arctic Subrealm bc Body cavity

Bs brachiopod shell

BR Boreal Realm

BPSr Boreal–Pacific Subrealm

Cb crural base

Cpr cardinal process

CSR complete spatial randomness

Crpr crural process

DIG left interval between the pedicle and the maximum width

Dl descending lamella

Dv dorsal valve

EB Edge Betweenness clustering procedure

ECS side-sulci space

ECP folding space

EPS Thickness

FG Fast Greedy clustering procedure

FLMNH Florida Museum of Natural History

HMG height at the maximum width to the left

KDE kernel density estimation

GMI Colombian Geological Service (formerly INGEOMINAS)

LAR Width lc lophophoral cavity

LV Leading Eigenvector clustering procedure

LON total length

LUD length of the dorsal valve

LP Label Propagation clustering procedure pA posteror adductor muscle

PSM depth of the median sulcation

M Myophragm

Ohp outer hinge plate

ML Multi-Level Optimization clustering procedure

MSC Michael Sandy collection

Pf pedicle foramen

S Symphytium

SLG length of the left side sulcation

Sr socket ridge

T Tooth

Tb transverse band

Uc umbonal chamber

TR Tethyan Realm

USNM United States National Museum (Smithsonian Institution)

Vv ventral valve

WT Walktrap clustering procedure

Abstract of Dissertation Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy

PALEOBIOLOGICAL IMPLICATIONS OF THE POST-PALEOZOIC FOSSIL RECORD OF THE LINGULIDE BRACHIOPODS AND ASSOCIATED FAUNAS

Alexis Rojas

August 2017

Chair: Michał Kowalewski Major: Geology

This dissertation represents an assessment of the post-Paleozoic fossil record of the lingulides and associated faunas, with a primary focus on adopting a spatially explicit perspective to study the fossil record. First, I described a fauna of Lower

Cretaceous brachiopods from Colombia, including a new terebratulid species and the first report of Lower Cretaceous lingulides in the region. Second, using a combination of field, museum, and literature data, I evaluated the post-Paleozoic history of drilling predation on the lingulides. Third, I developed a proxy for evaluating site-selective behavior of drilling predators based on spatial point process modeling. Results were consistent with the hypothesis that predation pressures increased through time in marine ecosystems. Fourth, because occurrences of lingulides were insufficient to assess their biogeography, I focused on ammonoid global records in the PaleoDB

(Paleobiology Database) to test the mid-Cretaceous biogeography using a network- based framework. Results supported the validity of previously delineated bioprovinces:

Boreal-Pacific, Arctic, Tethyan, and Austral. The geographic network derived from ammonoids was twice as dense as the one derived from benthic invertebrates and thus more effective in delineating bioprovinces. This dissertation highlights the importance of

the brachiopod fossil record as a source of paleobiological data and the benefits of incorporating spatially explicit approaches to improve our understanding of diverse aspects of the fossil record.

CHAPTER 1 INTRODUCTION: THE POST-PALEOZOIC FOSSIL RECORD OF THE LINGULIDE BRACHIOPODS

Lingulide brachiopods (Family Lingulidae) were globally widespread during most of the Phanerozoic (Holmer and Popov, 2000; Biernat and Emig, 1993), although today they are restricted primarily to low-latitude regions of the world (Campbell et al., 1997;

Emig, 1997). Most Paleozoic taxa are well established and accessible in databases and museum collections (see Paleobiology Database, Peng et al., 2007). In contrast to their

Paleozoic relatives, however, only a few Mesozoic lingulide taxa are described adequately (Biernat and Emig, 1993) and almost nothing is known about their presence and distribution in the Cretaceous rock record of tropical America. Well-described records of Cretaceous lingulides in the region are restricted to the Upper Cretaceous

(Turonian) Lingularia? notialis from Brazil (Holmer and Bengson, 2009). Two lingulide genera, Lingula and Glottidia, are the only Cenozoic representatives of this group.

Glotiddia is restricted to coastal areas from America between 10⁰S and 40⁰N latitude, whereas Lingula has a worldwide distribution, except along the coasts of America

(Campbell et al., 1997). Both genera are supposed to have originated during the early

Cenozoic (Emig, 1997), but their biogeographic origin is still speculative.

Recent studies identified some general trends in lingulide anatomy and body size through the Phanerozoic (Biernat and Emig, 1993; Peng et al., 2007; Zhang et al.,

2009). The overwhelming focus on Paleozoic taxa and a predominant qualitative approach, however, hinder accurate assessments of their post-Paleozoic history. For instance, our current knowledge of the fossil record of drilling predation on these organophosphatic brachiopods remains unexplored, and the hypothesis that lingulides are not an important prey for drilling predators remains largely untested. Chapters

presented below focus on (Chapter 2) describing a lower Cretaceous brachiopod fauna from Colombia, (Chapter 3) developing a proxy for site-selectivity of drilling predators,

(Chapter 4) evaluating the post-Paleozoic history of drilling predation on the lingulides, and (Chapter 5) delineating large-scale Albian (mid-Cretaceous) marine bioprovinces.

Chapter 2, “Lower cretaceous brachiopods from Colombia (South America): biostratigraphic significance and paleogeographic implications” examines the brachiopod fauna of the Rosablanca Formation near the town of Zapatoca (Santander

Province, Colombia). I describe the terebratuloid brachiopod Sellithyris elizabetha nov. sp., and use semi-quantitative taphonomic data to propose a large-scale erosional event that may have affected the Rosablanca Formation during Valanginian times.

Finally, I report the oldest occurrence of Cretaceous lingulides in the region.

Chapter 3, “A proxy for site-selective of drilling predators based on spatial point process modeling” describes a spatially–explicit approach for analyzing drilling traces on marine shelled invertebrates. The overwhelming majority of paleontological studies have concentrated on drilling intensity, ignoring the fact that predatory drillholes in prey skeletons are spatially explicit. Using a two-dimensional morphometric analysis, I constructed a point pattern of naticid-like drillholes that occurred on fossil Lirophora spp. surveyed from the FLMNH. The study comprises four species (i.e., L. glytocyma and L. latilirata) partitioned into four discrete time intervals of varying lengths (Miocene, Early

Pliocene, Late Pliocene and Pleistocene). I fitted kernel density estimators (KDE) to the observed drillholes and used the K-function to investigate clustering under the assumption of complete spatial randomness (CSR). Results indicate that predatory drillholes were significantly clustered. Neverthless, those areas where drillholes tend to

occur at significantly elevated rates relative to other prey shell areas (e.g., umbo, center, edge) differ across the sampled time intervals. This time-variant pattern may indicate temporal changes in site-selective behavior of naticid gastropods and highlights the potential for using this proxy to document the history of the drillhole site-selectivity.

Chapter 4, “The post-Paleozoic fossil record of drilling predation on lingulide brachiopods” provides a tentative evaluation of the post-Palaeozoic history of drilling predation on the lingulides. Research on drilling predation has been concentrated on prey with calcareous skeletons (e.g. molluscs, echinoids, rhynchonelliform brachiopods). This study is based on a compilation of literature sources and surveys of paleontological collections. Results indicate a Mesozoic-to-Cenozoic increase in drilling frequency that is similar to the trends observed in other marine benthic invertebrates and consistent with the hypothesis that predation pressures increased through time in marine ecosystems.

Chapter 5, “Global biogeography of Albian ammonoids: A network-based approach” provides a reproducible quantitative framework for delineating geographic boundaries of marine bioprovinces. Using aggregated species occurrences in the

Paleobiology Database, I generated a geographic network to quantify connectivity of

Albian epicontinental basins and used the flow-based Infomap algorithm to delineate bioprovinces. Those Infomap bioprovinces were largely concordant with the traditional, qualitatively derived biogeographic model. The geographic network derived from ammonoids was twice as dense as the one derived from benthic invertebrates and thus more effective in delineating bioprovinces.

Chapter 2 and Chapter 3 are manuscripts in preparation for submission. Chapter

4 and Chapter 5 were published as research articles in Lethaia (Rojas et al., 2016) and

Geology (Rojas et al., 2017), respectively.

CHAPTER 2 LOWER CRETACEOUS BRACHIOPODS FROM COLOMBIA (SOUTH AMERICA): BIOSTRATIGRAPHIC SIGNIFICANCE AND PALEOGEOGRAPHIC IMPLICATIONS

Introduction

Lower Cretaceous rocks are widely exposed in the Cretaceous basin of

Colombia. Brachiopods, however, are known from only a few localities restricted to the western side of the Eastern Cordillera. A number of studies in this region, stimulated largely by the search for petroleum, provided data for stratigraphic correlations and structural interpretations (e.g., Morales et al., 1958; Etayo, 1979; Villamil, 1998;

Guerrero, 2002a, 2002b). Comprehensive paleontological and biostratigraphic studies, however, are still limited (e.g. Bürgl, 1956; Etayo-Serna, 1964, 1968a, 1968b, 1979;

Villamil, 1998). In recent years, paleontological research has focused largely on the taxonomic description of vertebrates (e.g., Cadena, 2011, 2015; Carrillo-Briceño et al.,

2016; Páramo-Fonseca et al., 2016) and decapods (e.g., Gómez-Cruz et al., 2015;

Luque, 2015). Although brachiopods were globally widespread during the Lower

Cretaceous, only a single taxon has been described recently from Colombia (Schemm-

Gregory et al., 2012) and almost nothing is known about this group in the tropical

America (i.e., Belize, El Salvador and Honduras, to the north in Central America, southwards through Colombia to e.g., Peru, Bolivia, and Paraguay in South America) during that times.

Two brachiopod assemblages were reported in the literature from the Lower

Cretaceous Rosablanca Formation of Colombia. Those fossil assemblages are comprised of a few shells of the species Hadrosia gracilis from Santa Sofía, Boyacá

Province (Schemm-Gregory et al., 2012), and the terebratulide shells described herein as Sellithyris elizabetha nov. sp. from Zapatoca, Santander Province. The latter material

was originally referred to “Terebratula” sella Sowerby (Karsten, 1858; Dietrich, 1938) and Sellithyris sp. (Dietrich, 1938; Sandy, 1991a, b) in paleontological studies based on limited material with uncertain stratigraphic position and geographic location.

This paper documents new material collected during a number of field seasons carried out by the Colombian Geological Service (formerly INGEOMINAS), Florida

Museum of Natural History (FLMNH) and Smithsonian Tropical Research Institute

(STRI) in the town of Zapatoca. A new species, Sellithyris elizabetha nov. sp., is described combining external and internal morphological data, the latter derived from serial sectioning a specimen. In addition, lingulide shells assigned provisionally to the genus Lingularia are reported. It is the first record of lingulide brachiopods from Lower

Cretaceous rocks in northern South America. Finally, the analysis of taphonomic semi- quantitative taphonomic data collected on the external surface of Sellithyris elizabetha nov. sp. indicates an erosional hiatus that may have affected the biochronology of the

Rosablanca Formation. The new material discussed here is reposited at the Colombian

Geological Service.

Materials and Methods

Sample Collection

During a number of field seasons from 2012 to 2014, two stratigraphic sections

(65 and 140 m thick) and a number of nearby sites in the town of Zapatoca were sampled for macrofossils (Figure 2-1). The sampled interval spans the Upper

Rosablanca Formation to the lower beds of an overlying unidentified stratigraphic unit.

The Upper Rosablanca Formation was divided by Guzman (1985) into a number of limestone beds that are herein referred to as horizons (Figure 2-2). Mollusk-dominated fossil assemblages, preserved as internal molds, represent the majority of the

recovered material in those stratigraphic horizons. Less common fossils include echinoderms, brachiopods, arthropods and vertebrate remains. Most terebratuloid shells were recovered on top of the bedding plane of a single horizon (PI). Although these shells were transported over short distances, they are interpreted as being preserved within or near their original stratigraphic horizon. Consequently, these shells are considered here to be in situ elements (Kidwell et al., 1986). Only a few terebratulid brachiopods, including mostly shell fragments, were recovered from the stratigraphically higher non-contiguous horizon O. Bivalve-dominated fossil assemblages, preserved as flattened external molds, represent the majority of the recovered material from the intercalated mudstone beds. A few lingulide shells and their fragments, however, were recovered from two thick, bioturbated and non-contiguous beds situated between horizons UII and T (Figure 2-2).

Data Analysis

The material recovered from the horizons PI, and O is comprised of ~45 % complete shells of adult brachiopods, i.e. conjoined shells ≥15 mm long and complete enough to establish their original dimensions. These terebratulid shells are herein referred to as specimens. For each specimen, the dimensions of 10 morphological characters were measured according to Gaspard and Mullon (1983, Figure 1) to the nearest 0.1 mm using an electronic caliper: total length (LON), length of the dorsal valve

(LUD), width (LAR), thickness (EPS), folding space (ECP), side-sulci space (ECS), depth of the median sulcation (PSM), length of the left side sulcation (SLG), left interval between the pedicle and the maximum width (DIG) and height at the maximum width to the left (HMG) (Appendix A). To assess the variation of the brachiopod assemblage and among materials gathered from horizons PI and O, a principal component analysis

(PCA) was performed on log10-transformed measurements. Statistical analyses were performed with the MVA R-package (Hothorn, and Everitt, 2014). In addition, the taphonomic alteration produced by abrasion and/or dissolution of the shell surface in all specimens was evaluated by looking at dorsal valves under a stereomicroscope (25X).

All specimens were grouped into three categories: good (2 = puncta observed), fair (1 = puncta difficult to discern), and poor (0 = no puncta observed). The difference in the taphonomic alteration between specimens from horizons PI and O was evaluated by a non-parametric permutational MANOVA using the adonis function available in the vegan

R-package. This morphological study was extended to include 161 specimens of

Sellithyris sella from the Isle of Wight, England (MSC collection), and 168 specimens of

S. coahuilensis from Mexico from the National Museum of Natural History, Smithsonian

Institution. Sellithyris coahuilensis was used because of its morphological similarity to S. elizabetha sp. nov. and S. sella to incorporate another species from the genus (the type species).

The Sellithyris sella material used in the morphological study is from the Aptian

(Lower Cretaceous) of Group IV (of Fitton, 1847), Lower Gryphaea Group, Ferruginous

Sands Formation, Lower Greensand, from Atherfield, Isle of Wight, England (Casey,

1961; Simpson, 1985). The specimens were collected from a block of sandstone that contained a single mass of closely packed brachiopod shells. The block of sediment had fallen from the cliff face and was found on the beach. The material of Sellithyris coahuilensis used in the morphological study is from the Valanginian (Neocomian) lower beds of the Barril Viejo Formation, Potrero Oballos in Sierra Hermanos, Coahuila,

Mexico, collection numbers USNM 446382 and USNM 446383 (M. Sandy, personal

communication, April 27, 2017). Because of the small size and fragility of the organophosphate shell, linear measurements of length (L) and width (W) from the lingulides were obtained using ImageJ software (ImageJ, NIH, USA). Abbreviations used herein for collections: IGM = Colombian Geological Service (formerly

INGEOMINAS); USNM = United States National Museum (Smithsonian Institution);

MSC = Michael Sandy collection.

Systematic Palaeontology

Orden Lingulida Waagen, 1885 Superfamily Linguloidea Menke, 1828 Family Lingulidae Menke, 1828 Genus Lingularia Biernat and Emig, 1993 Type species Lingularia similis Biernat and Emig, 1993 Lingularia sp. Figure 2-3U–V Material: Dorsal valve, IGM 880538-1 (Figure 2-3V) (length 2.9 mm, width 1.7 mm); dorsal valve IGM 880538-2 (Figure 2-3U) (length 2.5 mm, width 1.6 mm), and 19 shell fragments IGM 880553.

Locality: Laguna del Sapo, Zapatoca, Santander Province (N 6° 50' 30.90"; W

73° 14' 21.48").

Stratigraphic horizon: Upper Rosablanca Formation, Lingularia bed–1, between horizons TI and T; Lingularia bed–2, between horizons UII and UI.

Description: Shell small (≤3 mm), elongate oval in outline, 1.6−1.7 times as long as width, with maximum length slightly anterior to mid-length; anterior margin rounded, antero-lateral corner broadly rounded, lateral margins gently rounded. Ornamentation consists of strong, concentric, evenly spaced growth lines. Larval shell oval, slightly longer than width (length 0.7 mm, width 0.6 mm in the two examined valves), with shell

rim strongly marked. Dorsal valve with two low plications extending from the umbo to the anterolateral corners, not sufficiently pronounced to define a median fold.

Remarks. Externally, the material is similar in both shape and size to the Berriasian-

Valanginian? Lingularia? sp. from the Barents Sea, but the beak appears to be more prominent in the latter, and the low plications present in the Colombian material appear to be absent in the Arctic species (Sandy in Ärhus et al., 1990, Figure 11a). It is also similar to the Valanginian-Cenomanian Credolingula spp. in having two plications extending from the umbo to the anterolateral corners (Smirnova, 2001, in Smirnova and

Ushatinskaya, 2001). These, however, are not sufficiently pronounced in the Colombian material to define a median fold as in Credolingula. Also, it differs from those in having smaller shells with more rounded anterior margin.

The material available is consistent in shape (i.e. general outline, anterior and lateral margins) and size with the Lower Aptian Lingularia michailovae (Smirnova 2001, pl. 6, figs. 14-17). Maximum width in the Colombian material, however, is located slightly anterior to the mid-length whereas the greatest width is attained in the posterior third of the shell in L. michailovae. The studied material can be distinguished from the

Cenomanian-Turonian Lingularia smirnovae (Biernat and Emig, 1993, Figure 7C-F) in having more rounded lateral and antero-lateral margins. Additionally, the maximum shell width in L. smirnovae is located at the mid-length. The Colombian material is similar in size to Lingula? cretacea Lundgren, 1885 from the Maastrichtian of southern

Scandinavia (Lundgren, 1885; Hansen and Surlyk, 2014) and Germany (Surlyk, 1982), and Campanian-Maastrichtian of England (Johansen and Surlyk, 1990). The low

plications present in the Colombian material are, however, absent in the European species (Hansen and Surlyk, 2014, Figure 4-23).

The fossil record of Cretaceous Lingulidae in the region is restricted to the Upper

Cretaceous (Turonian) Lingularia? notialis from Brazil (Holmer and Bengtson, 2009) and almost nothing is known about their presence and distribution in the Cretaceous rock record of tropical America. Lingularia? notialis differs from the Colombian material in having larger shells (10.2 mm), maximum width located posterior to mid-length and a straight to slightly rounded anterior margin (Holmer and Bengtson, 2009).

The examined material differs from unpublished Cretaceous lingulides (USNM

1665) (mean length 9.6 mm; width to length ratio 0.6; n = 6) recovered near La Plata,

Upper Magdalena Valley of Colombia, in having a smaller shell. It also differs from unpublished Lower Valanginian material from Germany, identified as Lingula truncata

Sowerby (1836) (USNM 190096) (mean length 13.4 mm; width to length ratio 0.6; n =

2), in shell size, shape of the anterior margin and location of the shell’s maximum width.

The anterior margin of the German material is nearly straight and the greatest width is located near the anterior margin of the shell. This species was originally described from

England and later assigned to the genus Credolingula (Smirnova 2001).

Order Terebratulida Waagen, 1883 Suborder Terebratulidina Waagen, 1883 Superfamily Terebratuloidea Gray, 1840 Family Sellithyrididae Muir-Wood, 1965 Subfamily Sellithyridinae Muir-Wood, 1965 Genus Sellithyris Middlemiss, 1959 Type species Terebratula sella J. de C. Sowerby, 1823 Sellithyris elizabetha nov. sp. Figs. 2-3A-P, 2-5, 2-6 1858 Terebratula haueri Klipstein: Karsten, p. 113, pl. VI, Fig. 1. 1938 Terebratula sella Sowerby: Dietrich, p. 106, pl. 22, Fig. 8. v 1985 Terebratulla sella Sow.: Guzman, p. XII-5.

v 1990a Sellithyris sp.: Sandy, p.417, Fig. 5.3. v 2009 Sellithyris sella: Rojas and Patarroyo, p. 130, Figs. 2, 3. Derivation of name: This species is named in honor of Elizabeth Sandoval

Rueda for her continuous support during the time of this research.

Material: Holotype, conjoined specimen illustrated in this work IGM 880558-2

(Figure 2-3E-H) (length 24.2 mm, width 22.3 mm). Paratypes: Conjoined shells IGM

880558-1 (Figure 2-3A-D); IGM 880558-3 (Figure 2-3I-L); and IGM 880558-4 (Figure 2-

3M-P). In addition, other material for a total of 269 conjoined shells.

Type locality: Laguna del Sapo, Zapatoca, Santander Province (N 6° 50' 30.90";

W 73° 14' 21.48").

Stratigraphic horizon: Upper Rosablanca Formation, horizons PI and O.

Diagnosis: Medium-sized, slightly longer than wide, and strongly bisulcate

Sellithyris; cardinal process low and bilobed; short loop with flat-bladed crural bases, crural process slightly incurved at tip and directed toward the mid-line, and transverse band with flattened crest.

Description: Specimens small to medium in size, outline pentagonal, slightly longer than wide with maximum width located anterior to mid-length. Shells equi- to gently ventri-biconvex in longitudinal section. Anterior margin strongly bisulcate at adult stage, with a short and shallow median sulcus. Inner surface of ventral beak with circular and permesothyridid pedicle foramen, pedicle collar well developed, and symphytium partially obscured. Interior of ventral valve: smooth surface; cyrtomatodont hinge teeth (Figure 2-5H). Interior of dorsal valve: umbonal chamber shallow and slightly wide. Cardinal process thin, wide and bilobed (Figure 52-V). Socket ridges thin, suberect and gently curved (Figure 2-5X). Outer hinge plates concave and extending

towards the crural process. Loop short, 1/3 of length and width of dorsal valve. Crural bases thin, slightly inclined toward the mid-line, and leading into the crural process.

Crural processes thin, high, slightly directed toward the mid-line, gently incurved and thickened at tip, situated anteriorly of mid-loop (Figure 2-5N, O). Descending lamellae short and curved inwards (Figure 2-5Q). Transverse band narrow, high-arched, with steep lateral slopes, and flattened crest (Figure 2-5R). Terminal points short and rounded (Figure 2-5T). Dorsal adductor muscle scars separated by a low and short medium myophragm (Figure 2-5L, M); lateral ridges absent.

Remarks: Sellithyris elizabetha nov. sp. appears distinct from other Valanginian-

Hauterivian forms in the genus Sellithyris (Table 2-1). It differs from the Early

Valanginian Sellithyris coahuilensis from Mexico (Sandy, 1990) by the loop morphology, including the flattened median crest of the transverse band, and the ventromedial- oriented crural processes. These Central and South American species can be distinguished from other Valanginian-Hauterivian forms within the genus (i.e., S. sella,

S. lindensis, S. carteroniana, and S. deningeri) by the lack of lateral ridges dividing the dorsal adductor field. The studied material of S. elizabetha nov. sp. exhibits high variability in outline, ranging from subpentagonal to slightly elongated (Figure 2-3A-P). A few of the elongated forms have elongate ventral umbos similar to those described in some small shells of Sellithyris coahuilensis by Sandy (1990) (Figure 2-3I-L). The anterior commissure of the Colombian material also exhibits high variability ranging from moderately sulciplicate to strongly sulciplicate. The PCA diagram showing the plots of 155 measured shells along the first and second ordination axis suggests that all shells recovered from horizons PI and O of the Rosablanca Formation represent a

single population (Figure 2-6A). Accordingly, the variation of the Colombian material as described above is here attributed to intraspecific variability. A similar approach was used by Gaspard and Mullon (1983) to quantify the morphological variability of

Cenomanian terebratulids in Western Europe. They studied specimens of four species:

Sellithyris tornacensis (d’Archaic); Sellithyris biplicata (Defrance); “Terebratula” phaseolina Lamarck (these all having been considered synonymous by previous authors); and also Platythyris vendeensis Middlemiss. Multivariate analysis combined with internal and microstuctural details resulted in the identification of three species:

Sellithyris tournacensis (d’Archaic); Sellithyris cenomanenis Gaspard; “Terebratula” phaseolina Lamarck, with Platythyris vendeensis Middlemiss considered a subspecies of the latter (M. Sandy, personal communication, April 27, 2017).

Discussion

Available brachiopod material provides information concerning the age of the

Rosablanca Formation in the Santander Province of Colombia. Although brachiopods are reported in only a few stratigraphic levels, the material is sufficient to provide some age constraint on the upper part of this geological unit. The Rosablanca Formation is commonly considered to be Valanginian to Early Hauterivian in age (Guzman, 1985), but the part of the formation that contains Sellithyris elizabetha nov. sp. may be as old as Late Valanginian (cf. Middlemiss, 1984a). The first report of a well-preserved assemblage dominated by “Terebratula sella” from the vicinity of the town of Zapatoca was published by Dietrich (1938), but the exact stratigraphic position of his material was uncertain. Sandy (1991a, p. 420, Figure 5.3a-d) figured a specimen of Sellithyris sp. from Upper Valanginian sediments of Zapatoca (serial sections of this specimen are

presented herein, Figure 2-5), recorded in another work as from the Saynoceras biohorizon (Sandy, 1991b, p. 143).

Here I described Sellithyris elizabetha nov. sp. based on new material recovered from the same geographic area. The genus Sellithyris is currently considered to range from the Berriasian to Turonian (Lee and Smirnova, 2006). Benest et al. (1996) recorded Early Cretaceous Berriasian Sellithyris from Algeria, Africa. The type species

Sellithyris sella (Sowerby, 1823) was a widely-distributed form in Western Europe during the Early Cretaceous, including the Late Valanginian (Middlemiss, 1981, 1984;

Gaspard, 1997, 2005). This species is also known from younger rocks in Spain,

France, England and Hungary (Detre, 1968; Dieni et al., 1973; Middlemiss, 1984;

Gaspard, 1988, 1999). Sellithyris? middlemissi Sandy, 1986 from the Berriasian in south-eastern France (Alpes de Haute-Provence) is distinct from other members of the subfamily Sellithyridinae in lacking biplication of the anterior commissure (Sandy, 1986, p. 184). This form probably represents a juvenile terebratulide of uncertain affinity

(Gaspard, personal communication to M. Sandy).

There is no significant difference between the external morphological features measured on specimens from horizons PI (n = 143) and O (n = 12) (Figure 2-6A), and the variation of the material is here attributed to intraspecific variability. Specimens from horizon O are, however, less common, more fragmented, and exhibit loss of superficial detail (i.e. no puncta or growth lines observed) (Figure 2-6B). The difference in the taphonomic condition between specimens from these two horizons is significant (Table

2-2). Based on both observations and outcomes of the permutational MANOVA, the brachiopods in horizon O were probably excavated from the underlying horizon PI,

which indicates an erosional hiatus that may affect the biochronology of the Rosablanca

Formation. This is compatible with field observations of the molluscan assemblages, which have not yet been quantified.

Brachiopods are common from horizon PI in all studied sections and field stations in the region, making them potentially valuable stratigraphic markers at both local and regional scales. The new data on the stratigraphic distribution of S. elizabetha nov. sp. in the Lower Cretaceous of Colombia not only supports connection with inner-shelf sites in the Mediterranean Tethys, but also with the ancient Gulf of Mexico. Imlay (1937) described both "Terebratula" coahuilensis and "Terebratula" kanei from Valanginian sediments from a number of localities in Coahuila, Mexico. Sandy (1990) later assigned the material of "Terebratula" coahuilensis to the genus Sellithyris. Both forms described by Imlay (1937), however, may belong to a single species.

In terms of Early Cretaceous paleobiogeography, the Colombian, Mexican, and

Spanish localities were probably not very far apart as the Atlantic Ocean was still in early development (García-Ramos, 2009). Owen (1981) also recognized early

Cretaceous faunal links between Tethyan brachiopods on both sides of the opening

Atlantic, e.g., the terebratulide Cyrtothyris from Europe and Mexico and the terebratellidine Colinella from Morocco and Mexico. Sandy (1991a) previously commented on the link that the likely presence of Sellithyris sp. (S. elizabetha nov. sp. herein) provided with European faunas, also commenting on other links between Early

Cretaceous Colombian and European brachiopod faunas: Musculina, Arenaciarcula and

Gemmarcula (Sandy, 1991a).

Organophosphate shells of the brachiopod Lingularia sp. were recovered from slightly bioturbated mudstones named Lingularia bed–1 and Lingularia bed–2. Because species belonging to the genus Lingularia have been recorded from the Carboniferous

(?) to the Upper Cretaceous (Biernat and Emig, 1993, p. 11), the material is of little value in ascertaining the age of the Rosablanca Formation. The beds containing lingulide fragments, however, can be traced laterally for almost 1000 m, allowing local stratigraphic correlations of the unit. Although much of the material is fragmentary, the specimens of Lingularia sp. reported here are the oldest occurrences of Cretaceous lingulides in this region so far. The lingulides have been traditionally considered to be rare elements in Cretaceous near-shore marine communities, however the new material reported here suggests that they may have been common locally.

Figure 2-1. Geographic location of the town of Zapatoca in the Eastern Cordillera of Colombia.

Figure 2-2. Stratigraphic section of the Upper Rosablanca Formation in the Laguna del Sapo locality, Zapatoca, showing basic lithology and brachiopod-yielding horizons. Modified from Guzman (1985).

Figure 2-3. Bivariate plot of shell length (L) and width (W) of Lingularia sp. from the Upper Rosablanca Formation and other Cretaceous lingulides worldwide.

Figure 2-4. Lower Cretaceous brachiopods from Colombia. A–P. Sellithyris elizabetha nov. sp. trom the Rosablanca Formation in Zapatoca, Santander Province. A– D. IGM 880558-1, conjoined shell from horizon O. A. Dorsal view. B. Ventral view. C. Lateral view. D. Anterior view. E–H. IGM 880558-2 holotype, conjoined shell from horizon PI. E. Dorsal view. F. Ventral view. G. Lateral view. H. Anterior view. I–L. IGM 880558-3, conjoined shell from horizon PI. I. Dorsal view. J. Ventral view. K. Lateral view. L. Anterior view. M–P. IGM 880558-4, conjoined shell from horizon PI. M. Dorsal view. N. Ventral view. O. Lateral view. P. Anterior view. Scale 10 mm. Q–T. Hadrosia gracilis Schemm-Gregory et al. 2012, IGM 880558-2, conjoined shell from Rosablanca Formation in Santa Sofia, Boyacá Province. Q. Dorsal view. R. Ventral view. S. Lateral view. T. Anterior view. Scale 10 mm. U–V. Lingularia sp. from Rosablanca Formation in Zapatoca. U. IGM 880538-1, dorsal valve. V. IGM 880538-2 holotype, dorsal valve. Scale 1 mm.

Figure 2-5. Serial sections through Sellithyris elizabetha nov. sp., sectioning perpendicular to commissural plane, sectioning distance indicated in mm from posterior end. Abbreviations: bs, brachiopod shell; cb, crural base; cpr, cardinal process; crpr, crural process; dl, descending lamella; dv, dorsal valve; m, myophragm; ohp, outer hinge plate; pf, pedicle foramen; pf, pedicle foramen; s, symphytium; sr, socket ridge; t, tooth; tb, transverse band; uc, umbonal chamber; vv, ventral valve.

Figure 2-6. Morphological and taphonomic data of Sellithyris elizabetha nov. sp. from the Lower Cretaceous of Colombia. A. Principal components diagram of 155 specimens from horizons PI (n = 143) and O (n = 12) showing their position along the first and second ordination axis based on measured shell dimensions. B. Taphonomic alteration produced by abrasion and/or dissolution on the shell surface in shells from horizons PI (white bars) and O (grey bar).

Table 2-1. Morphological comparison of Sellithyris elizabetha nov. sp., and other Valanginian forms within the genus.

Sellithyris Sellithyris Sellithyris sella Sellithyris lindensis Sellithyris deningeri Sellithyris elizabetha nov. carteroniana coahuilensis (Imlay, (J. de C. Sowerby, 1923) Middlemiss, 1976 Dieni et. al., 1973 sp. (d’Orbigny, 1849) 1940) Size small to medium medium to large small to medium small to medium small to large small to medium rounded, oval or subpentagonal, rhomboidal to pentagonal, alongate, markeldy subpentagonal to pentagonal, slightly longer Outline pentagonal, slightly almost as wide as almost as wide as long longer than wide subtriangular than wide longer than wide long strongly equi- to gently Curvature ventribiconvex biconvex (globose) biconvex biconvex ventribiconvex ventribiconvex sulciplicate to Anterior margin strongly bisulcate strongly sulciplicate strongly bisulcate bisulcate strongly bisulcate bisulcate wide, moderately short, triangular, sligtly obscured Symphytium – very short – wide, triangular wholy visible by dorsal beak

Median myophragm present present present present present present

Lateral ridges present present present present absent absent Cardinal process wide, bilobed wide, bilobated wide, bilobed narrow, bilobed narrow, bilobed wide, bilobed suberect, gently suberect, gently nearly erect, gently Socket ridges erect, gently curved suberect, robust suberect, gently curved curved curved curved fairly wide, end in a point wide, horizontal concave, rarely Outer hinge plates just below (dorsally) of the wide, concave wide, concave wide, concave cuneate resupinate crural process thin, directed thin, directed thin, directed thin, directed Crural bases thin, nearly parallel thin, subparallel venteromedially venteromedially venteromedially venteromedially high, sligtly directed high, sligtly directed base tickened, directed directed base tickened, high, venteromedially, venteromedially, gently Crural processes venteromedially, slightly venteromedially, directed high, subparallel gently incurved and incurved and thickened at incurved at tip gently incurved venteromedially thickened at tip tip short, curved inwards, short and curved short and curved short and curved Descending lamellae short, curved inwards short and curved inwards angle 60° inwards inwards inwards

moderatelly high, broadly narrow, high-arched, narrow, high-arched, narrow, high-arched, Transverse band narrow, low-arched narrow, high-arched arched sligtly trapezoid sligtly trapezoid sligtly trapezoid

Median crest narrowly rounded flattened broadly rounded flattened narrowly rounded flattened

Geographic Great Britain, Central Germany, France, England, Germany Italia Mexico Colombia distribution Europe Switzerland

Stratigraphic Valanginain to Early Late Valanginain to Valanginain to Hauterivian Early Valanginian Early LateValanginian distribution Aptian Early Hauterivian ?Hauterivian

Table 2-2. Results of the permutational MANOVA conducted to evaluate the difference in the taphonomic alteration between specimens from horizons PI and O using the adonis function available in the vegan R-package.

df Sum of squares Mean squares F. model R2 Pr (>F) Taphonomy O 0.0053 0.0026 4.2355 0.0674 0.0010 Horizon PI 0.0020 0.0020 3.1490 0.0251 0.9000 Residuals 114 0.0706 0.0006 0.9075 Total 117 0.0778 1.0000

Comparisons were made using horizons O and PI as groups (strata) within which to constrain 1,000 permutations.

CHAPTER 3 A PROXY FOR SITE-SELECTIVE OF DRILLING PREDATORS BASED ON SPATIAL POINT PROCESS MODELING

Introduction

Predation by drillers on shelled marine invertebrates represents a unique opportunity to quantifying a biological interaction in the fossil record (Kowalewski, 2002).

Predatory drillholes are direct evidence of predation and can be used to assess predator activity (Leighton, 2002; Kowalewski, 2002). These traces provide quantitative data on two aspects of the predator-prey interaction: intensity and selectivity (Kitchell et al., 1981; Kitchell, 1986; Calvet,1992; Dietl and Alexander, 2000). Predation intensity usually refers to the frequency of predatory traces in a given taxon or assemblage, and it is traditionally considered the most powerful tool for quantitatively assessing drilling predation quantitatively in the fossil record (Kowalewski, 2002). Selectivity describes the predator’s preferences for choosing a particular prey species, a prey size class, or a drilling location on the prey skeleton (Kelly, 1988; Kowalewski, 2002).

Experimental studies have shown that drilling predation is a highly-stereotyped behavior (Kelly, 1988; Chiba and Sato, 2012). For instance, naticid gastropods locate drillholes in a stereotyped fashion (Dietl and Alexander, 2000; Dietl et al., 2001), probably related to prey handling during the attack and the prey morphology (Kabat,

1990; Rojas et al., 2015). The overwhelming majority of paleontological studies have focused on drilling intensity, ignoring the fact that drillholes in prey skeletons are spatially explicit. Thus, our knowledge of drillhole-site selectivity by drillling predators remains limited.

Methods for analysis of drillhole-site selectivity were primary developed to test the null hypothesis of equal distribution of drillholes with respect to arbitrary sectors of the prey skeleton. They include goodness-of-fit tests such as Pearson x² and

Kolmogorov-Smirnov approaches (Kowalewski, 1990; Kelley, 1988) and diversity metrics such as the Shannon-Weaver index (Dietl et al., 2001). These sector-based approaches, however, lack spatial information (Baddeley et. Al., 2016). The recognition that drillholes on prey skeletons are spatially explicit and can be mapped enables the development of a proxy for drillhole-site selectivity based on spatial point process theory.

The approach implemented here is a three-step process. First, a point pattern of the location of predatory drillholes on a standardized prey skeleton is created using two- dimensional morphometrics. Second, a data-driven, nonparametric method called

Kernel-Density Estimation (KDE) is used to calculate the density of traces across the prey skeleton and map hotspots. Third, the K-function is used to test the null hypothesis of complete spatial randomness (CSR) of the drillhole locations. This site-selective analisis is illustrated with a case study based on naticid-like drillholes in specimens of the Cenozoic bivalve Lirophora latilirata surveyed from the Florida Museum of Natural

History (FLMNH).

Materials and Methods

Data Collection: Museum Survey

Museum data were obtained by a survey of the Invertebrate Paleontology

Collections at the Florida Museum of Natural History (FLMHN; 56 lots containing 1915 specimens). Cataloged collections of the species Lirophora latilirata (Conrad) from several localities on the Atlantic Coastal Plain were analyzed under a binocular

microscope for presence of naticid-like drillholes (Table 3-1). Predatory drillholes were recognized using the criteria established by Bromley (1981) for identifying the ichnotaxon Oichnus paraboloides, and those representing successful predatory events were recognized following the criteria of Kowalewski (2002). The museum survey was focused on bulk-collected specimen lots that were associated with large collections.

Based on the stratigraphic resolution of the museum lots, drilled speciments were partitioned into three discrete time intervals of varying lengths: Early Pliocene (EP), Late

Pliocene (LP) and Pleistocene (P).

Point Pattern Construction

A total of 278 drilled shells were found and photographed using a standard photogrammetric protocol adapted from Perea et al. (2008). All images were oriented such that the antero-posterior axis was horizontal to facilitate consistency in landmark data collection (Kolbe et al., 2011). The position of the predatory drillholes was quantified using the two-dimensional morphometric approach originally proposed by

Roopnarine and Beussink (1999). This approach was later adapted by Rojas et al.

(2015) to superimpose the drillholes on a standard prey shell using four pseudolandmarks on the external view of the valves: (1) point of maximum curvature of the ventral edge, (2) anterior end of valve, (3) the beak on the outline, and (4) posterior end of the valve. A fifth point corresponds with the predatory drillhole. The pseudolandmarks 2 and 4 are located on the anterior and posterior margins of the bivalve shell and define its maximum length (Kolbe et al., 2011).

The selected pseudolandmarks do not correspond with particular internal homologous traits of the prey, as in Roopnarine and Beussink (1999). Instead, they reflect the general external geometry of the valve, which may be an important

consideration for naticid predators when manipulating their prey. Landmark coordinates obtained for left valves were inverted to compare directly with the right valves following

Kowalewski (2004). The Bookstein baseline registration method for two-dimensional data (Bookstein, 1986) was used to remove shell size, position information and remaining orientation from the analyses using as baseline pseudolandmarks 1 and 3.

The distribution of predatory drillholes in such a standardized prey skeleton (i.e., study area in Bookstein shape units) represents a spatial point pattern (Figure 3-1). Point level attributes are shown in Appendix B. The R functions for landmark-based morphometrics of Claude (2008) were used to undertake this part of the analysis.

Density Analysis of Drillholes

Kernel density estimation was used to visualize the spatial distribution of predatory drillholes on the prey skeleton of L. latilirata. This technique fits a curved surface over each case such that the surface is highest above the case and zero at a specified distance (h) from the case. The Kernel density density is mathematically expressed as (Silverman, 1986):

푛 1 K(푑ᵢ) 푓(푥, 푦) = . ∑ (3˗1) 푛ℎ ℎ 푖=1

where ƒ(x, y) is the density value at drillhole location (x, y), n is the number of cases, h is the bandwidth, di is the distance between drillhole case i and drillhole location (x, y) and K is a Gaussian kernel density function. Drilling frequency (DF), calculated for each museum lot using the following formula: [number of valves drilled] /

[0.5 x total number of valves] (Rojas, et al. 2015), was used to reweight the kernel density estimate and thus it considers variations in sampling effort. The optimum bandwidth hopt was calculated following Fotheringham et al. (2000). Descriptive statistics

and parameters employed to calculate the optimum bandwidth hopt used for the kernel density estimation (KDE) are presented in Table 3-2. Because drillhole configurations are scaled and aligned on the baseline of coordinates (−0.5, 0) and (0.5, 0), derived from Bookstein registration, standard measurements (e.g., mm) can’t be read from these maps. Kernel density estimation was performed using the function density.ppp in the R-package SPATSTAT version 1.23-4. Hotspots for predatory drillholes on the prey skeleton were defined as the upper 10% of estimated density values, following Nelson and Boots (2008).

Spatial Clustering Analysis of Drillholes

The Ripley's K-function is used to analyze the 2-dimensional distribution of predatory drillholes and to quantify deviations from complete spatial randomness (CSR) in a statistically consistent framework. The estimates of K(r) implemented in this study is mathematically expressed as (Ripley, 1988):

푎 퐾(푟) = ∑ ∑ 퐼(푑 ≤ 푟)푒 (3˗2) 푛(푛 − 1) 푖푗 푖푗 푖 푗

where a is the area of the standardized prey skeleton, n is the number of drillholes, and the sum is taken over all ordered pairs of drillholes i and j in the point pattern. dij is the distance between the two drillholes and I(dij ≤ r) is the indicator that equals 1 if the distance is less than or equal to r. The term eij is the edge correction. The expected value of K(r) for a random Poisson distribution is πr2 and deviations from this expectation indicate either clustering or dispersion. Three distinct sceneries are evaluated: (1) null hypothesis [complete spatial randomness]: K(r) = πr2; (2) alternative hypothesis 1 [clustered]: K(r) > πr2; and (3) alternative hypothesis 2 [dispersed]: K(r) <

πr2.

A nonparametric test for complete spatial randomness (CSR) is constructed by simulating s spatial patterns of size n from a homogeneous spatial Poisson process based on the assumption of CSR, and used to compute critical values for the K-function

(Baddeley et al., 2016). The K-function was estimated using the function Kest for a range of values between 0 and 2.5 units (measured in Bookstein shape coordinates), and critical values were computed by Monte Carlo simulation using the function envelope. Functions used for analysis of spatial clustering are available in the R- package SPATSTAT version 1.23-4

Results

Kernel density maps in Figure 3-2D-F show the concentration of drillholes on the prey skeleton per square unit (in Bookstein shape coordinates) at each time interval considered in this study. These kernel estimate surfaces visualize locations of drillhole abundance and scarcity, and the spatial variability in predatory events across the prey shell. The middle shell of the Early Pliocene L. latilirata is the area with major concentration of naticid-like drillholes. High concentrations of those traces are also observed in the umbo and the edge (i.e, edge-drilling) (Figure 3-2D). The composite region comprising the middle shell and the umbo of the Late Pliocene L. latirilata has the major concentration of naticid-like drillholes. Some areas in the ventral edge also exhibs a high concentration of those traces (Figure 3-2E). The umbo, including the edge near to the hinge ligament, and the middle shell areas of the Pleistocene L. latirilata have the major concentration of naticid-like drillholes. A single area in the anterior ventral edge also exhibs a high concentration of traces (Figure 3-2F). Hotspot maps for each time interval are presented in Figure 3-2G-I. Overall, drillholes located on the edge of the shell represents less than the upper 10% of estimated density values and

do not represent hotspots. However, the edge near to the hinge ligament in the

Pleistocene L. latirilata was identified as a hotspot of naticid-like drillholes.

The graphical output of the K-function indicates significant clustering of naticid- like drillholes on the skeleton of L. latilirata across a range of Bookstein shape distances in all time intervals considered in this study. However, those traces were not clustered at very short distances (< 0.06 Bookstein shape distance units) during the Early

Pliocene (Figure 3-3A). The average number of drillholes within a distance r of another drillhole is statistically greater than that expected for a random distribution (Figure 3-3A-

C) and the alternative hypotheses of complete spatial randomness (CSR) and dispersion describing the distribution pattern of naticid-like drillholes on the skeleton of

L. latilirata can be rejected for all time intervals. The K-function results indicate that naticid-like drillholes on the skeleton of L. latilirata are significantly clustered (two-sided

Monte Carlo test with significance level α = 0.05 at the distance r). The results for the

Early and Late Pliocene differs from those from the Pleistocene in the large vertical separation between the empirical and expected K-functions at any interpoint distance r as well as the maximum vertical separation between them (Figure 3-4).

Discussion

Naticid-like drillholes on the skeleton of the Cenozoic Lirophora latilirata were concentrated mostly around the umbo and the middle-shell area. Areas with a high concentration of those traces were also observed in the egde of the prey shell. The number and location of areas with high concentration of naticid-like drillholes vary between time intervals considered in this stufdy. Those areas not always represent hotspots of such predatory traces (Figure 3-3). For instance, cases of edge-drilling, a mode of naticid predation in which a predator drills a hole at a point on the commissure

between the closed valves (Taylor, 1980; Ansell and Morton, 1985; Rojas, et. al. 2016), represented less than the upper 10% of estimated density values and thus are not here considered as hotspots (Figure 3-3G-I). The results indicate that studied populations of

L. latilirata were not subject of significant edge-drilling predation.

Preferences in site-selectivity of unidentified naticid gastropods, as recognized from naticid-like drillholes on modern beach assemblages of a specific prey, may be consistent across space and environments (see Rojas, et. al. 2016). Then, observed differences in the concentration of naticid-like drillholes and the number of hotspots on the skeleton of fossil L. latilirata across time intervals may represent either through-time variations in site-selectivity of the predators or the signal from predators with different preferences. Locality level data were not available to validate these suggestions.

Despite stratigraphic (e.g., long and unequal time intervals) and geographic limitations of the data assembled in this study (e.g., sampling localities relatively far apart one of each other, potential variation in sampled environments), the K-function indicates a significant clustering of the naticid-like drillholes across the skeleton of L. latilirata for all time intervals under consideration. This is the first time that the K-function is used to quantitative test for the site-selectivity of drilling predators. This spatially– explicit approach establishes a reproducible quantitative framework for analyzing drilling traces on marine shelled invertebrates and identifying biotic and abiotic factors that influence this predator–prey interaction. The method described here may be generalized to a wide spectrum of ecologic and taphonomical processes such as encrustation and bioerosion on marine shelled invertebrates. Future studies could define time intervals with similar lengths as well as the consider species with contrasting

morphologies such as smooth versus strongly ornamented taxa to evaluate the potential effect of prey shell morphology on drillhole-site selectivity by drilling predators. The kernel approach implemented here allows the detection of hot spots for predatory drillholes, which may be investigated in terms of underlying variables, such as prey thickness, ornamentation and coloration.

Figure 3-1. Point pattern of naticid-like drillholes on Pliocene and Pleistocene Lirophora latilirata from the Atlantic Coastal Plain.

Figure 3-2. Kernel density estimation and hotspot mapping of naticid-like drillholes on Pliocene and Pleistocene Lirophora latilirata from the Atlantic Coastal Plain. A–C. Point pattern splited per time interval considered in this study. Early Pliocene (28 drillholes), Late Pliocene (63 drillholes) and Pleistocene (73 drillholes). D–F. Weighted spatial kernel density estimation using drilling frequency (DF) as a weight variable. G–I. Hotspot maps indicating the upper 10% of estimated density values. Kernel density maps units are number of drillholes per area; area is measured in square Bookstein Shape units.

Figure 3-3. K function for naticid-like drillholes on Lirophora latilirata from the Atlantic Coastal Plain. A. Early Pliocene (28 drillholes). B. Late Pliocene (63 drillholes). C. Pleistocene (73 drillholes). Empirical (solid lines) and expected (dashed lines) K-functions and upper 95% confidence envelop (grey) estimated from 999 simulations of a Poisson process.

Figure 3-4. Empirical K-function values subtracted by its expected value (K(r)−πr2) for predatory drillholes on Lirophora latilirata. Dashed reference line represents the expected K-function values subtracted by itself (πr2−πr2).

Table 3-1. Drilling data on museum samples of Lirophora latilirata compiled in this study. Sample ID # valves # drilled valves DF Geological unit Age UF161845 79 8 0.20 Duplin Early Pliocene UF161844 21 7 0.67 Duplin Early Pliocene UF164230 29 1 0.07 Duplin Early Pliocene UF164234 49 5 0.20 Duplin Early Pliocene UF164014 37 2 0.11 Duplin Early Pliocene UF164012 21 4 0.38 Duplin Early Pliocene UF157413 20 5 0.50 Jackson Bluff Late Pliocene UF157412 25 1 0.08 Jackson Bluff Late Pliocene UF157411 19 2 0.21 Jackson Bluff Late Pliocene UF157410 20 5 0.50 Jackson Bluff Late Pliocene UF157409 27 9 0.67 Jackson Bluff Late Pliocene UF157408 20 2 0.20 Jackson Bluff Late Pliocene UF157622 31 2 0.13 Jackson Bluff Late Pliocene UF157414 20 5 0.50 Jackson Bluff Late Pliocene UF147337 21 8 0.76 Pinecrest Beds Late Pliocene UF137167 13 2 0.31 Pinecrest Beds Late Pliocene UF157627 32 6 0.38 Pinecrest Beds Late Pliocene UF146228 42 2 0.10 Pinecrest Beds Late Pliocene UF157628 26 6 0.46 Pinecrest Beds Late Pliocene UF137878 25 4 0.32 Pinecrest Beds Late Pliocene UF144502 39 1 0.05 Pinecrest Beds Late Pliocene UF147358 35 3 0.17 Pinecrest Beds Late Pliocene UF206594 115 13 0.23 Waccamaw Early Pleistocene UF217095 55 5 0.18 Waccamaw Early Pleistocene UF217094 48 5 0.21 Waccamaw Early Pleistocene UF217097 48 5 0.21 Waccamaw Early Pleistocene UF217099 64 7 0.22 Waccamaw Early Pleistocene UF217098 56 4 0.14 Waccamaw Early Pleistocene UF206587 55 13 0.47 Waccamaw Early Pleistocene UF2107096 53 5 0.19 Waccamaw Early Pleistocene UF137849 23 1 0.09 Bermont Early Pleistocene UF137880 13 1 0.15 Bermont Early Pleistocene UF137172 14 1 0.14 Bermont Early Pleistocene UF131622 23 1 0.09 Bermont Early Pleistocene UF131624 26 2 0.15 Bermont Early Pleistocene UF131623 24 2 0.17 Bermont Early Pleistocene UF127365 20 2 0.20 Bermont Early Pleistocene

Table 3-1. Continued. Sample ID # valves # drilled valves DF Geological unit Age UF131625 30 3 0.20 Bermont Early Pleistocene UF30048 12 3 0.50 Bermont Early Pleistocene UF145969 25 1 0.08 Caloosahatchee Early Pleistocene UF137856 21 2 0.19 Caloosahatchee Early Pleistocene

Table 3-2. Descriptive statistics and parameters used to calculate the optimum bandwidth (hopt). Parameter Value mean center x -0.158 mean center y 0.087 standard distance 0.261 std-x 0.159 std-y 0.208 hopt 0.062 sigma (hopt/2) 0.031

CHAPTER 4 THE POST-PALEOZOIC FOSSIL RECORD OF DRILLING PREDATION ON LINGULIDE BRACHIOPODS1

Introduction

Considerable attention has been devoted to the fossil record of biotic interactions between drilling predators and their marine invertebrate prey, including mollusks (Sohl,

1969; Kitchell, 1986; Kowalewski et al., 1998; Kelley and Hansen, 2003), brachiopods

(Ausich and Gurrola, 1979; Thayer, 1985; Kaplan and Baumiller, 2000; Leighton, 2001,

2003; Hoffmeister et al., 2004; Kowalewski et al., 2005), echinoderms (Baumiller 1993,

1996; Nebelsick and Kowalewski, 1999; Kowalewski and Nebelsick, 2003; Zlotnik and

Ceranka, 2005), and, to a lesser extent, arthropods (Pasini and Garassino 2012;

Klompmaker et al,. 2015), polychaetes (Martinell et al., 2012; Klompmaker, 2012), and a few other higher taxa.

In contrast, our current knowledge of the fossil record of drilling predation on lingulides (Lingulidae: Brachiopoda) remains unexplored. A few studies on durophagous predators (i.e., shell−crushing organism) preying upon lingulides (e.g., Emig and

Vargas, 1990; Mason and Clugston, 1993; Kowalewski et al., 1997; Harris et al., 2005) and incidental reports of drilling attacks (e.g., Paine, 1963; Emig, 1997) suggest that lingulides may be attractive to durophagous, but only occasionally attacked by drillers.

However, in some cases, drilling gastropods may have significant impact on extant lingulide populations (see Paine, 1963). This high drilling frequency may represent a local anomaly rather than a globally widespread phenomenon. Studies of predatory traces (e.g., drillholes, breakage and repair scars) on lingulide shells are restricted to

1 Reprinted with permission from Rojas, A., Portell, R.W., and Kowalewski, M., 2017, The post-Palaeozoic fossil record of drilling predation on lingulide brachiopods: Lethaia, v. 50, p. 296–305.

the Recent Glottidia palmeri from Baja California (Kowalewski and Flessa, 1994;

Kowalewski et al., 1997; Kowalewski and Flessa, 2000) and Glottidia pyramidata from the Gulf of Mexico (Paine, 1963). In addition, naticid gastropods have been pointed out as potential predators of Glottidia audebarti in Central America (Emig, 1983; Emig and

Vargas, 1990). Evidence of durophagous predation has been found in shells of modern

Lingula (C.C.Emig pers. communication 1997 cited in Kowalewski et al., 1997), and studies of stomach contents in a number of fish species have recorded their remains

(Longhurst, 1958; Worcester, 1969; Onyia, 1973). Predatory drillholes have been reported among other inarcticulate brachiopods including modern craniids (Emig, 1997) and fossil acrotretids (Miller and Sundberg, 1984; Chatterton and Whitehead, 1987).

The hypothesis that lingulides are not an important prey of drilling predators remains largely untested. Direct evidence of predator−prey interactions, as documented in the fossil record, can provide data for assessing the paleoecological importance and evolutionary history of this interaction. In this study, we investigate the fossil record of drilling predation on post-Paleozoic lingulides by combining literature sources and surveys of museum collections. In addition, we report new fossil evidence of drilling predation on the Pliocene Glottidia inexpectans from the Atlantic Coastal Plain of

Virginia and Florida.

Materials and Methods

Data

We assembled a dataset of occurrences of post-Paleozoic lingulides, with associated information on stratigraphy and geology (Appendix C). As defined here, a lingulide includes any taxon that can be assigned confidently to the family Lingulidae.

Two main data sources have been combined in the dataset: literature and museum

collection surveys. Museum data were derived via an exhaustive survey of the

Invertebrate Paleontology Collections (both cataloged and uncataloged) at the Florida

Museum of Natural History [FLMNH] (36 lots from 10 localities containing 561 specimens), and a survey of the Paleobiology Collections at the Smithsonian National

Museum of Natural History [NMNH] (35 lots/localities containing 128 specimens). The

Cenozoic Glottidia inexpectans is the only lingulide represented in the FLMNH data

(Table 4-1). It is a taxon known for its occasional occurrences in Pliocene sediments of eastern North America (Campbell et al., 1997). The examined material at the NMNH represents mainly Mesozoic lingulides (83%) and most of them are here provisionally assigned to the genus Lingularia (see Holmer and Bengtson, 2009). Uncatalogued samples of Glottidia dumortieri (7 localities; 37 specimens) from the Neogene of

Belgium and reposited at the Royal Belgian Institute of Natural Sciences [RBINS] are also included in the data.

Our new dataset expands and revises a previous compilation of lingulides constructed by Peng et al. (2007) that was focused on Paleozoic and Early Mesozoic records and did not include any information on predatory interactions. In terms of numbers of specimens and localities, the dataset provides a comparable data coverage for Mesozoic and Cenozoic eras. However, the geographic coverage is temporally discontinuous and spatially incomplete (multi-regional rather than global). The combined literature and museum data represent 8,154 specimens from 307 localities (341 samples). However, only 964 specimens from 139 localities could be examined for evidence of drilling predation, i.e., presence of complete or incomplete drillholes. This limited number of examined specimens reflects the fact that only a small subset of

specimens reported in the literature is figured in publications. Also, not all museum specimens are sufficiently complete to evaluate for presence of drillholes. Examined specimens are represented either by a single valve or a conjoined shell that is sufficiently complete to measure its original width. Those specimens include 721 museum specimens and 243 specimens accessible as figured material in published references. Localities with specimens examined for drilling predation represent 45% of all localities recorded in our dataset.

Methods

Specimens from museum collections and figured material in the published references were examined for complete and incomplete drillholes. In identifying and interpreting those traces of drilling predation we followed standard criteria (Kowalewski,

2002). Drilling frequencies at the locality-level were calculated using the following formula: [number of valves drilled] / [0.5 x total number of valves] (Table 4-1). Shell fragments with a characteristic fracture (half-moon) were identified in some museum specimens, but their biotic origin could not be conclusively demonstrated. These equivocal records of drillholes were not included in quantitative analyses. For all specimens, both drilled and undrilled, the shell width (W) and length (L) were measured using a digital caliper with precision of +/- 0.03 mm. L is measured along the anterior- posterior axis, from the most posterior point of shell (or valve) to the farthest point on anterior margin. W is the maximum dimension measured from right to left and normal to the plane of symmetry (Williams and Brunton, 1997). The size dimension W is used here as a proxy of the lingulide shell size because this dimension is readily obtained for specimens missing the posterior or anterior end.

Whereas museum collections and figured specimens are potentially prone to biases when evaluating for traces of predation (e.g., Kowalewski et al., 2000), it is noteworthy that the museum and figured specimens examined here do not differ significantly in drilling frequency (0.5 % for museum specimens and 1.3 % for figured specimens; p = 0.4; Fisher’s Exact Test). In addition, many of the examined museum lots represent bulk sampled material and are unlikely to have suffered biases associated with selective specimen curation.

The approximate location of each drillhole on its prey valve, relative to the shell outline and muscle scars, was recorded on a schematic diagram of a lingulide shell

(Figure 4-1). The latter is a semi-quantitative approach employed for assessing selectivity in drilling location by predators (Zlotnik, 2001; Baumiller et al., 2006; Tuura et al., 2008), which can be used when quantitative strategies based, for example, on landmark approaches (e.g., Roopnarine and Beussink, 1999; Hoffmeister et al., 2004) are impractical. In addition, we measured the drillhole diameter (D), which is defined here as the maximum diameter of the outer outline of each drillhole (Figure 4-2), and used it as a proxy of the predator size (Kitchell et al., 1981), All measurements on images were made using the ImageJ software (ImageJ, National Institutes of Health,

Maryland, USA). The two-sided Fisher's exact test was used to assess the statistical significance (α = 0.05) of differences in drilling frequency and drilling occurrences between Mesozoic and Cenozoic time intervals. We explored the relationship between drillhole size (D) and prey shell size (W), on drilled specimens that could be measured reliably, using reduced major axis regression. To avoid a potential bias introduced by the selection of specimens photographed in literature sources (i.e., authors consistently

choose a large size class of specimens for illustration) (Krause et al., 2007), the statistical analyses of the differences in size between drilled and undrilled elements were restricted to museum specimens that were bulk-collected (Appendix C). A juvenile assemblage of lingulide shells, recovered from paired valves of Chesapecten jeffersonious at Carters Grove Plantation, Virginia (UF 115656), was not included in prey size analyses because it represents an unusual mass mortality event (Campbell et al., 1997). Because of unbalanced sampling and small sample size of drilled specimens, we used the rank-based Wilcoxon Test. All analyses were performed using R (R

Development Core Team, 2013).

Results

The combined literature and survey data include 8,154 specimens representing

307 localities, including 964 specimens from 138 localities that could be evaluated for drill holes. Each locality in the dataset registers one single species. The dataset comprises a total of six genera, which vary notably in frequency of occurrences. In the total localities of the dataset, the genera Lingularia (39%), Lingula (21%) and Glottidia

(19%) occur commonly, whereas Credolingula, Sinolingularia and Sinoglottidia have been reported from a few only (≤ 4% in all cases). In addition, undetermined lingulide genera are relatively common in the Mesozoic localities (24% of Mesozoic occurrences). The age of the total localities in the dataset ranges from ~250 my (Early

Triassic) to the Recent, covering paleolatitudes from 72° N to 71° S and paleolongitudes from 180° W to 167° E. Mesozoic and Cenozoic represent 59% and 41% of the total localities in the dataset, respectively. However, the data coverage is uneven geographically and temporally, with the total number of localities per period varying from

15 (Paleogene) up to 82 (Triassic).

Drillholes in lingulides are infrequent: a total of 23 drilled specimens (0.3% total specimens; 2.4% inspected specimens) representing eight localities (2.6% total localities; 5.8% examined localities). Despite comparable data coverage for Mesozoic and Cenozoic eras, the two-time intervals differ significantly in drilling frequency (0% inspected specimens and 0% inspected localities for Mesozoic; 3.3% inspected specimens and 15.1% inspected localities for Cenozoic; p < 0.001; Fisher’s Exact Test).

Although multiple genera were sampled at comparable level, only Glottidia yielded drilled specimens (Table 4-1). However, localities that did not include any drilled specimens still provide valuable numerical data about the absence of drilling (i.e., drilling frequency = 0%). Consequently, the data reported here estimate drilling frequency patterns of the family Lingulidae. However, data that pertain to drilling patterns such as site- or size- selectivity are limited to one genus (Glottidia) and may not be valid a generalization for the entire family.

Drilled specimens were found in the Americas (i.e., North and Central) and nearby Antarctic localities (i.e., Seymour and Alexander Islands). The record of drilled shells in Neogene sediments of The Netherlands is the only evidence of drilling predation on lingulides from outside of the previously mentioned regions. The

Netherlands specimens belong to Glottidia dumortieri, the only known occurrence of the genus Glottidia outside the Americas and Antarctica (Chuang, 1964).

The difference in drilling frequency between specimens of the two Cenozoic genera Glottidia and Lingula was not statistically significant (0% inspected specimens for Lingula and 3.6 % for Glottidia; p = 0.4; Fisher’s Exact Test), whereas the difference in the number of inspected localities with drilled specimens was significant (0%

inspected localities for Lingula and 22.9% for Glottidia; p<0.05; Fisher’s Exact Test). For those localities that included drilled specimens, the drilling frequency varies notably, ranging from 0.1 to 20% (Table 4-1). However, some of the locality-level estimates are based on small numbers of specimens (n < 10) and estimates of drilling frequency based on such small samples are suspect. Given that the overall drilling frequency reported here is very low (<3%), very small samples are unlikely to yield any drilled lingulide specimens. Consequently, the very low proportion of localities with drilled specimens reported here is likely to underestimate the proportion of localities with drilled prey specimens. Nevertheless, the pooled data (964 specimens) clearly indicates that drilled lingulide fossils are rare.

Drillholes on lingulide shells cluster around the body cavity and include penetrations of both, dorsal and ventral valves of the prey (Figure 4-1). In addition, a few traces are located in the anterior region of the shell, which roughly corresponds to the lophophoral cavity. We found no significant difference in mean size between drilled

(18.5 mm) and undrilled specimens (17.0 mm) in the museum samples of Glottidia inexpectans (p = 0.3; Wilcoxon Signed-Rank Test) (Table 4-2; Appendix C). There is a strong linear relationship between drillhole diameter and shell size of the drilled specimens (r2 = 0.9, p < 0.001) (Figure 4-2), but these results are highly tentative given that only seven drilled specimens, in which the size measurement W can be obtained confidently, are included in this analysis. Drillholes are typically circular to slightly oval in outline, with a characteristic beveled edge (Kitchell, 1986). Their average maximum outer diameter is 2.3 mm, ranging from 0.7 to 2.8 mm (Figure 4-3).

Discussion

Despite the scarcity of reports documenting drilling predation on extant lingulides and the virtual absence of such reports in the fossil record, the results presented here indicate that lingulide brachiopods have been subject to drilling predation since at least the Eocene. Despite temporal, geographic, and methodological limitations of the data assembled here, the results clearly indicate that drilling predation has been infrequent in the Cenozoic and may have been absent in the Mesozoic (Figure 4-4). High drilling frequencies of naticids feeding on mollusk prey have been reported from the geological units that yielded drilled lingulids in our study (Kelly et al., 2001; Aguilar and Alvarado

2004; Brezina et al. 2016). The much higher frequency of predatory attacks on mollusks suggests that naticids were common in those ecosystems. Consequently, to the relative rarity of drilling attacks on lingulides is unlikely to reflect the scarcity of predators, but rather points to selective preference of mollusk prey by drilling predators. However, there is tentative evidence for somewhat elevated drilling frequencies at the Pliocene locality UF-ZV006 from Virginia and the Eocene Seymour Island (p < 0.1 in both cases;

Table 2). Cenozoic lingulides could have been, occasionally, subject to high predation pressures from drilling organisms, which is consistent with observations on Glottidia pyramidata in present day ecosystems (Paine 1963). This notion is also supported by the tentative evidence for stereotypy suggested by site-selective location of drilling attacks (Figure 4-1) as well as tentative evidence for size-selectivity of drilling predators

(Figure 4-2).

The results reported here are likely to be a conservative estimate of the importance of drilling predation, both in terms of drilling occurrences across localities and drilling frequencies within localities. Because many localities in our data are based

on a limited number of specimens, rarity of localities that yielded drilled specimens may be due to undersampling: when drilling frequency is very low; a small sample is unlikely to capture any drilled specimens. Furthermore, drilling frequencies may be underestimated due to preferential loss of drilled valves. Mollusk-based studies yielded incongruent conclusions regarding the importance of drillholes as a taphonomic agent

(see Roy et al., 1994; Pechenik et al., 2001; Zuschin and Stanton, 2001; Zuschin et al.,

2003; Kelly, 2008), but organophosphatic shells of lingulides tend to be more fragile than most shells of mollusks (Emig, 1990) and thus could be parcticularly prone to this bias. However, all drilled specimens from museum samples are fragmented specimens

(i.e., size dimension L cannot be obtained confidently) that did not fracture at the drillhole (Figure 4-3) indicating that the breakage was not facilitated by perforations (see

Roy et al., 1994). Drilling frequencies at locality-level may be also altered by selective post-mortem transportation (Lever et al., 1961; Kornicker et al., 1963; Chattopadhyay et al., 2014). However, taphonomic studies suggest that lingulids are unlikely to survive substantial transport (Emig, 1986) and should be, therefore, less prone to this bias relative to mollusks. The potential bias due to removal of shells by durophagous predators (Vermeij et al., 1989; Harper et al., 1998; Harper, 2016) is not possible to resolve.

The identification of a drilling predator from its traces is difficult and controversial

(Kowalewski, 2002). However, based on the characteristic beveled shape (Kitchell,

1986) of the observed drillholes, it is likely that drilling-traces on lingulide valves in the museum survey (i.e., Yorktown Formation), and most of those from literature sources

(i.e., La Meseta Formation, Bagaces Formation), were produced by naticid gastropods.

Lingulides are infaunal organisms that preferentially inhabit soft sediments, and they are more likely to be drilled by naticid gastropods, which typically attack infaunal prey

(Gonor,1965; Hughes, 1985). Naticid gastropods are present in those geological units

(e.g., Kelly et al., 2001; Aguilar and Alvarado, 2004; Brezina et al., 2016), which is consistent with our interpretation that drillholes were made by these predators. It is noteworthy that drillholes in extant specimens of G. pyramidata were attributed to naticid predators as well (Paine, 1963).

This study provides a first, tentative insight into the evolutionary history of drilling predation on lingulide brachiopods. Although the absence of drilling in the Mesozoic may reflect sampling limitations of this study, the statistically significant Mesozoic-to-

Cenozoic increase in drilling frequencies is reminiscent of the similar trends observed in other marine benthic invertebrates (e.g., Kowalewski et al., 1998, 2005; Walker and

Brett, 2002; Baumiller et al., 2006; Huntley and Kowalewski, 2007) and consistent with the macroevolutionary hypothesis that predation pressures increased through time

(Vermeij ,1977, 1987). We hope that the results reported here will motivate future quantitative studies on the fossil record of drilling predation on lingulide brachiopods.

Figure 4-1. Schematic diagram of a conjoined lingulide shell in dorsal view indicating the drillhole location for those specimens in which the drilling site can be determined. Grey circles correspond to drillholes on G. inexpectans (this study) and black circles represent estimates gathered from literature sources (see Table 4-1). Asterisks indicate holes drilled through the ventral valve; question mark indicates valves that cannot be identified as either dorsal or ventral. Dorsal interior characters based on Emig (1988). Abbreviations: aA = anterior adductor muscle; aO = anterior oblique muscle; pA = posteror adductor muscle; lc = lophophoral cavity; bc = body cavity.

Figure 4-2. Scatter plot of lingulide size (estimated by the specimens that could be measured reliably in terms of shell width [W]) and drillhole size (estimated by the maximum outer drillhole diameter). Grey circles correspond to drillholes on G. inexpectans (this study) and black circles represent estimates gathered from literature (see Table 4-1).

Figure 4-3. Drilled specimens of the Pliocene Glottidia inexpectans Olsson, 1914. A–B. conjoined shell, arrow indicating a drillhole circular in outline, Tamiami Formation, UF 52890. C. Conjoined shell bearing a circular drillhole with a beveled edge, Peace River Formation, UF 11840. D–E. Ventral valve fragment with a drillhole, Tamiami Formation, UF 92111. D. Internal view showing that the drillhole is located proximal to the pedicle nerve scar. E. External view of the drillhole illustrating the drillhole’s slightly elliptical outline and beveled edge. F–G. Dorsal valve with a drillhole originally figured by Cooper (1988), Yorktown Formation, USNM 551520h, paratype. F. Internal view showing that the drillhole is located proximal to the septa. G. External view illustrating the drillhole’s elliptical outline and beveled edge. All scale bars equal 1 cm.

Figure 4-4. Quantitative summary of drilling predation on post-Paleozoic lingulide brachiopods. A. Specimen level data. B. Locality level data.

Table 4-1. Summary of quantitative data for the Cenozoic localities that yielded drilled specimens of lingulide brachiopods. *Taxonomic names for naticids based on Turgeon et al. (1998). Abbreviations: s = conjoined valves, v = valves, inc = incomplete drillholes, DF = Drilling frequency. # # drilled Age Locality Formation/Member Taxon DF Presumed Predator References elements elements Neverita duplicata Stump Pass Recent not applicable Glottidia pyramidata 97 s 13 0.134 Sinum Paine (1963) Florida, USA perspectivum* San Felipe Kowalewski and Recent not applicable Glottidia palmeri 1 v 1 — Muricidae (?) Baja California, Mexico Flessa (1994) Kingsmill 01 (UF- Olsson (1914); Yorktown Formation Glottidia Pliocene ZV006) Virginia, 10 v 1 0.200 Naticidae Cooper (1988); Sunken Meadow inexpectans USA this work Wilsons Marl Pit 01 Glottidia Pliocene (UF-ZV041) Virginia, Yorktown Formation 18 v 1 inc 0.111 Naticidae (?) this work inexpectans USA Casa de Meadows 02 Glottidia 130 v + 19 Pliocene (UF-CH026) Florida, Tamiami Formation 2 0.024 Naticidae (?) this work inexpectans s USA Ft. Green 13 Dragline Peace River 01 Glottidia Pliocene Formation, 35 v 1 0.057 Naticidae (?) this work (UF-PO002) Florida, inexpectans Bone Valley Member USA Barbudal Creek Aguilar and Pliocene Guanacaste, Costa Bagaces Formation Glottidia sp. unknown 1 — unknown Alvarado (2004) Rica Beugen (64–66 m Miocene below surface) undetermined Glottidia dumortieri 2946 v 2 0.001 Naticidae Dulai (2013) Netherlands Bitner (1996); Seymour Island, 23 s + 22 Eocene La Meseta Formation Glottidia antarctica 2 0.060 unknown Emig and Bitner Antarctica v (2005) *Sympatric naticid gastropods reported by Paine (1963).

Table 4-2. Statistical significance of drilling frequencies for localities in which the recovered elements (i.e., valves and conjoined valves) represent at least five specimens. Estimates of p are based on drilling frequencies expected at a given sample size if frequency of attacks was uniform though time and space and reflected the overall drilling frequency (0.0129) estimated from the pooled data. Estimates of p were obtained using two approaches: Monte Carlo approximations (100,000 iterations) and binomial test. Abbreviation: DF = Drilling frequency.

Number Drilled p-value p-value Age Locality of DF elements simulation binomial test elements Stump Pass Recent 97 13 0.134 0.000 0.000 Florida, USA

Kingsmill 01 (UF-ZV006) Pliocene 5 1 0.200 0.063 0.062 Virginia, USA

Wilsons Marl Pit 01 (UF- Pliocene 9 1 0.111 0.108 0.109 ZV041) Virginia, USA Casa de Meadows 02 Pliocene 84 2 0.024 0.292 0.292 (UF-CH026) Florida, USA Ft. Green 13 Dragline 01 Pliocene 18 1 0.056 0.209 0.207 (UF-PO002) Florida, USA

Beugen (64–66 m below Miocene 1473 2 0.001 0.000 0.000 surface) Netherlands

Eocene Seymour Island, Antarctica 34 2 0.059 0.071 0.070

CHAPTER 5 GLOBAL BIOGEOGRAPHY OF ALBIAN AMMONOIDS: A NETWORK-BASED APPROACH2

Introduction

Biogeographic studies of fossil organisms have contributed to our current understanding of the relationships between plate tectonics and evolution of life

(Lieberman, 2005). However, the impact of those studies extends beyond basic biogeographic questions to include, for example, conservation biology and climate change (Perrin and Kiessling, 2012; García-Molinos et al., 2015). Recently, the network theory has brought significant advances to our understanding of biogeographic patterns in fossil organisms (Brayard et al., 2007; Dera et al., 2011; Sidor et al., 2013; Vilhena et al., 2013; Dunhill et al., 2016). One of the most relevant features of network analysis is its ability to detect community structure — natural partitioning of network nodes into densely connected subgroups (Newman and Girvan, 2004). The network-based approach has been demonstrated to be a powerful tool in modern biogeography

(Vilhena and Antonelli, 2015; Edler at al., 2016). Here, we employ a network approach to examine the biogeography of Albian (mid-Cretaceous) ammonoids using data from the Paleobiology Database (PaleoDB), a major geoinformatics initiative aimed at providing fossil occurrence data across all taxa retrievable from the geological record

(Peters and McClennen, 2016).

Large scale biogeographic regions for the Cretaceous have been delimited by ammonoid experts using qualitative assessments of taxonomic inventories (e.g.,

2 Reprinted with permission from Rojas, A., Patarroyo, P., Mao, L., Bengtson, P., and Kowalewski, M., 2017, Global biogeography of Albian ammonoids: A network-based approach: Geology, v. 45, p. 659–662.

Kennedy and Cobban, 1976; Jagt-Yazykova, 2011). This approach in ammonoid research has deep historical roots and remains unverified by quantitative assessments

(Ifrim et al., 2015). Because of the high dispersal potential of ammonoids, their biogeographic partitioning is likely to reflect large-scale physical, climatic and/or biotic environmental changes (Bengtson and Kakabadze, 1999). We focused on the Albian because this stage is represented by the largest number of ammonoid records in

PaleoDB, compared to the other Cretaceous stages. We explicitly test the qualitatively- derived biogeographic model for the Albian ammonoids as defined in the biogeographic synthesis of Lehmann et al. (2015): Boreal Realm (subdivided into Boreal–Atlantic,

Boreal–Pacific, and Arctic subrealms), Tethyan Realm, and Austral Realm. These bioprovinces have been consistently recognized in studies on the distribution of ammonite taxa (e.g., Owen, 1973; Kennedy and Cobban, 1976; Page, 1996).

Materials and Methods

Data

Occurrence data of Albian ammonoids were downloaded from the PaleoDB on

20 January 2016. The search was restricted to occurrences with species-level resolution, and those with uncertain or provisional taxonomic identification (i.e., qualified by aff., cf., ex.gr., sensu lato, or quotation marks) were excluded. The following data fields were downloaded: collection number, genus name, species name, geologic formation, and present-day latitude and longitude. The reduced data set contained a total of 1795 occurrences that met the filtering criteria. Paleogeographic coordinates were determined using the PointTracker Software and rotated occurrences plotted on the plate tectonic configuration from the PALEOMAP (Scotese, 2013). The stage-level resolution used in this analysis reflects data limitations: the number of species

occurrences currently available is insufficient to conduct a meaningful network analysis at finer stratigraphic resolution.

Network Construction and Partitioning

The network analysis implemented here is a four-step process (Figure 5-1). First, we aggregate the rotated occurrence data into a geographical grid of 5 × 5 degree cells.

This spatial resolution is widely used in studies of mid-Cretaceous climate and paleogeography (e.g., Fluteau et al., 2007). Second, aggregated data are used to generate a bipartite network (G) between species (S) and grid cells (P) denoted G = (V,

E), where V is the node set of two disjoint subsets (S, P) (Figure 5-2). E  P × S is the edge set which links P and S subsets. The incidence matrix (B) representing the connections between species (S) and grid cells (P) has the elements Bij such that

1, if taxon j belongs to grid cell i 퐵 = { (5˗1) 푖푗 0, otherwise

Since no edges are possible within node subsets P and S, the proportion of all possible edges that are actually present (i.e., density) in the bipartite network G was calculated as |V| / (|P| × |S|). Third, we performed a weighted projection from the bipartite network G onto the node subset P (Alzahrani and Horadam, 2016). The projection procedure generates a geographic network GP = (P, EP) in which two grid cells k and l  P are linked together if they have at least one common taxon in S (Figure

5-3). The adjacency matrix A representing the connections between grid cells in the projected network GP has the elements Akl such that

1, if grid cells 푙 and 푘 have taxa in common 퐴 = { (5˗2) 푘푙 0, otherwise

For a grid cell k  P, let (k) denote the set of neighbors of k. Then, the connection strength (CS) between distinct grid cells k and l in P are given by

훤(푘) ∩ 훤(푙) 퐶푆 = , 푘 ≠ 푙 (5˗3) 푘푙 퐶(푘) + 퐶(푙)

where C(k) and C(l) are the total number of collections recorded at grid cells k and l. This common neighbor index, standardized using the number of collections, takes into account variations in sampling effort. We also performed a projection from the bipartite network G onto the node subset S and generates a network GS = (S, Es), in which two species are linked if they occur together at least in one grid cell. Fourth, we applied the Infomap clustering algorithm (Rosvall et al., 2009) to partition the grid cells within the projected network GP into bioprovinces. Infomap finds the modular structure of the network with respect to flow by using random walks. The algorithm calculates the theoretical limit of how concisely we can describe the trajectory of a random walker on the network and selects the partition that gives the shortest description length. The analyses were performed using the R-package igraph 0.6 (Csardi and Nepusz, 2006).

Results

The bipartite occurrence network of Albian ammonoids (G) comprises a total of

540 nodes partitioned into 78 grid cells (P) and 462 taxa (S). This network is fairly sparse, comprising 859 edges that represent only 2% of all possible connections

(density 0.02). The number of edges per node of the subset P is highly variable (mean degree 11.0 ± 12.5 SD) and correlated with the number of sampled formations per grid cell in the dataset (r = 0.7, n = 78, p < 0.01). Ammonoid species (subset node S) are connected with ~2 grid cells on average (1.9 ± 1.6 SD) (Figure 5-2). The emergent projection GP is a spatially explicit arrangement of 78 nodes connected by 433 weighted

edges. The GP projection covers most of the Albian epicontinental basins and quantifies the strength of the connectivity across most marine regions (Figure 5-3A). This geographic network is fairly sparse (density 0.14), and the number of edges per node approximates a power law distribution. The maximum node-to-node distance (diameter) of the network GP, as measured by number of edges, equals 5. The numbers of mutual connections between GP network nodes range from 1 to 43, with ~12 mutual connections, on average. A small fraction of nodes (~6%) are disconnected from all remaining nodes.

The Infomap algorithm applied to the geographic network GP partitioned data into four non-overlapping bioprovinces (Figure 5-3B). The number of nodes per Infomap bioprovince ranged from 9 to 40 (Table 5-1). Disconnected nodes were disregarded because they did not contain meaningful information about the overall network structure. The Infomap bioprovinces match closely the traditional qualitatively established biogeographic units: Boreal Pacific Subrealm, Arctic Subrealm, Tethyan

Realm and Austral Realm. The modularity score of this division of the network GP – the fraction of edges within the given Infomap bioprovince minus the fraction expected if edges were distributed at random – is relatively low (Q = 0.24), but indicates the presence of a significant community structure (Newman, 2006a).

To find out how stable this division of the network GP is, we compared the results obtained by Infomap to six different partitioning procedures using the Normalized Mutual

Information similarity score (NMI) (Figure 5-4). According to the NMI, the Label

Propagation procedure (Raghavan et al., 2007) produces grouping most closely aligned with the Infomap output. The increasingly less concordant (although still generally

consistent) groupings were derived by Walktrap (Pons and Latapy, 2006), Multilevel

(Blondel et al., 2008), Fastgreedy (Clauset et al., 2004), and Leading Eigenvector

(Newman, 2006b) partitioning procedures. The Edge Betweenness partitioning algorithm (Girvan and Newman, 2002) produced numerous small-sized communities.

Because two partitioning algorithms can reach a similar level of performance (NMI) but produce different grouping structures (Orman et al., 2011), we implemented a qualitative comparison of the different partitions using the largest node overlap across procedures. Despite some variations, the grouping structure of the network GP was relatively consistent regardless of the algorithm and most nodes fell within the corresponding Infomap community. However, some of those algorithms failed to distinguish Boreal subrealms, whereas some algorithms subdivided further the Tethyan

Realm (Figure 5-4).

The Infomap bioprovinces differ from those obtained by an agglomerative hierarchical clustering method (UPGMA) in which a few of the nodes clustered out of their geographic context (Figure 5-5). The latitudinal distribution of the Infomap bioprovinces is presented in Figure 5-6. The network analysis was also implemented for

Albian benthic invertebrates in the PaleoDB. The construction procedure resulted in a geographic network with a similar size but less dense that the one derived from ammonoids (Figure 5-7). The clustering procedure resulted in a network partition with a higher modularity for the benthic invertebrates compared to the ammonoid (Table 5-2), and failed to reproduce the ammonoid Infomap bioprovinces.

Discussion

The partitioning of the geographic network GP into four groups using the Infomap algorithm resulted in an analytical outcome that is largely concordant with the traditional

biogeographic model of Albian ammonoids. The most obvious difference between the qualitative and network-based biogeographic model is that Infomap failed to delineate the Boreal–Atlantic Subrealm, either reflecting limitations of our data or inaccuracies of the traditional biogeographic model. This Boreal–Atlantic Realm has been, however, recognized in qualitative studies based on non-ammonoid taxa (e.g., Iba et al., 2011).

Our outcome may also reflect the resolution limits of the Infomap algorithm implemented in igraph, which uses standard teleportation; a procedure used in random walk-based methods that allows walkers to randomly teleport across the network on any node

(Lambiotte and Rosvall, 2012). Because the primary goal of this study was the global- scale test of major biogeographic bioprovinces, this limitation is not critical. The results presented here offer tentative evidence that the Boreal Realm was partitioned into Arctic and Pacific subrealms. The hierarchical cluster UPGMA failed to replicate the Infomap bioprovinces regardless of the level of similarity at which grid cells are joined (Figure 5-

5), pointing to the advantage of Infomap for detection of biogeographic structure.

The location of the Austral-Tethyan boundary in our biogeographic model differs substantially from the one suggested in Lehmann et al. (2015). In our model, the Austral

Realm is restricted to the Australian Epicontinental Sea. However, a node representing the Northern Territory of Australia clustered with the Tethyan Realm (Figure 5-3B). This configuration is driven by the occurrence of widespread forms in Australian borderlands and endemism of the Great Artesian Basin (Wright, 1963; Henderson, 1990). The

Tethyan Realm in our biogeographic model extends from the open waters of the

Tethyan Ocean, the Central Atlantic and the Paleo-Caribbean to ~60° S paleolatitude, and includes the northernmost Antarctic Peninsula region. This asymmetric pattern

may be controlled by the paleogeographic position of both South America and Africa, which were still relatively close and largely located in the Southern Hemisphere. Such a geographic configuration appears to have provided a number of relatively well- connected epicontinental basins suitable for ammonoids. In addition, southern latitudes were relatively warm and the northernmost Antarctic Peninsula region maintained relatively strong marine connections with South America and Africa (Bice and Norris,

2002; Martin and Hartnady, 1986).

Overall, the spatial distribution of the Infomap bioprovinces is reminiscent of the known large-scale paleogeographic and oceanographic features of the Albian earth: restricted Arctic Ocean, relatively open Tethyan region, and partly isolated South

Atlantic Ocean (Sewall et al. 2007). The latitudinal distribution of the Infomap bioprovinces (Figure 5-6) appears to support the hypothesis that latitude-related factors also played an important role in shaping the biogeographic partitioning of the mid-

Cretaceous ammonoids by reducing dispersal and constraining bioprovinces in space

(Reboulet, 2001; Ifrim et al., 2015). However, these interpretations may partly depend on geographic and stratigraphic resolutions of analyzed data, and may thus be inapplicable for analyses carried out at finer observational scales.

Our results shed new light onto the controversial affinities of the Albian ammonoid faunas from the Netherlands Antilles (see Owen and Mutterlose, 2006). For multiple network partitioning procedures, the grid cell containing this tropical region clustered with the Arctic Subrealm (Figure 5-3B). The spatial relationship among grid cells of the Infomap bioprovince representing the Boreal Realm in our model suggests that a Boreal affinity for areas proximal to the paleo-equator is unlikely. This outcome is

consistent with the Tethyan affinity suggested for this region previously (Owen and

Mutterlose, 2006). Centrality measurements on the projected network GS (Table 5-3) allows us to identify important or highly connected species.

Despite having a similar size, the geographic network derived from ammonoid data is twice as dense as the network derived for Albian benthic marine invertebrates.

The overall comparison indicates that benthic invertebrates do not replicate the biogeographic patterns observed in the ammonoids, and thus confirm the premise that ammonoid biogeographic partitioning is more likely to reflect large-scale environmental changes (Bengtson and Kakabadze, 1999).

The approach utilized in this study establishes an alternative, objective standard framework for studying the spatiotemporal dynamics of the marine paleobioprovinces over evolutionary timescales (e.g., origination, extinctions, expansions, contractions, and migrations). It is also a powerful methodological framework for comparative biogeographic studies within and across taxa. This approach complements qualitative studies by providing a spatially unambiguous and reproducible strategy for quantitative delineation of bioprovinces.

Figure 5-1. Workflow diagram indicating the procedures implemented in the analysis of the global records of Albian ammonoids.

Figure 5-2. Bipartite occurrence network of Albian ammonoids G.

Figure 5-3. Geographic network GP and Infomap bioprovinces. A. Projected one-mode network GP. Links are colored indicating their connection strength (CS). B. Community structure in the projected network GP. Nodes are colored indicating the Infomap bioprovinces. Abbreviations: BPSr—Boreal–Pacific Subrealm; ASr—Arctic Subrealm; TR—Tethyan Realm; AR—Austral Realm. Black unfilled squares are disconnected nodes.

Figure 5-4. Comparison of the Infomap results with other estimated grouping structures. LP—Label Propagation; WT—Walktrap; ML—Multi-Level Optimization; FG— Fast Greedy; LV—Leading Eigenvector; EB—Edge Betweenness. Disconnected nodes are represented as white unfilled areas.

Figure 5-5. Ammonoid occurrence data per grid cell (i.e., incidence matrix B) clustered using the unweighted pair-group arithmetic average method (UPGMA). Distances between grid cells were calculated using Bray–Curtis dissimilarity. For easier comparison of the cluster topology with the Infomap bioprovinces, the height of the tree nodes was adjusted so that the tree will have a distance of one unit between each parent/child nodes. The height range was also adjusted to 1. This analysis was performed using the vegan package for R software (version 2.3-4, Oksanen et al., 2016).

Figure 5-6. Latitudinal distribution of nodes in the projected network GP grouped by Infomap bioprovince.

Figure 5-7. Geographic network GP-benthos and Infomap bioprovinces. A. Projected network GP-benthos derived from Albian benthic marine invertebrate records in the PaleoDB. Links are colored indicating their connection strength (CS). B. Infomap bioprovinces. Black unfilled squares are isolated nodes.

Infomap Bioprovince IB1 IB2 IB3 IB4 IB5 IB6 IB7 Anthozoa 1 36 91 — — — — Asteroidea 1 14 — — 100 — — Bivalvia 24 26 1 100 — — 100 Calcarea — — — — — — — Crinoidea 7 — — — — — — Demospongea — — — — — — — Echinoidea 9 2 — — — — — Gastropoda — — — — — — — Hexactinellida — — — — — — — Holothuroidea — — 0 — — — — Lingulata 1 6 — — — — — Malacostraca 7 7 2 — — 75 — Ophiuroidea — 0 0 — — 25 — Rhynchonellata 51 8 5 — — — —

Figure 5-8. Taxonomic composition of the Infomap bioprovinces delineated in the projected network GP-benthos.

Table 5-1. Comparison of the estimated community structures in the projected network derived from Albian ammonoid records in the PaleoDB (GP). Number of Algorithm Comunities size Q† NMI§ comunities* InfoMap (IM) 4 40, 15, 9, 9 0.24 N.A.#

Label Propagation (LP) 4 40, 12,12, 9 0.24 0.94 Walktrap (WT) 3 40, 24, 9 0.26 0.92 Multilevel (ML) 4 25, 22, 17, 9 0.31 0.78 Fastgreedy (FG) 4 25, 23, 16, 9 0.30 0.78 Leading Eigenvector 5 26, 19, 14, 9, 2 0.28 0.70 (LE) Edge Betweenness 23, 9, 7, 7, 5, 3, 2, 2, 9 0.18 0.65 (EB) 2 *Isolate nodes have not been taken into account since they provide no significant information on the overall network structure. †Q = Modularity. §NMI = Normalized Mutual Information score. #N.A. = not applicable.

Table 5-2. Comparison of the projected networks derived from Albian ammonoid (GP) and benthic marine invertebrate records (GP-benthos) in the PaleoDB

Network Ammonoids (GP) Benthos (GP-benthos) Number of nodes 78 77 Number of links 433 176 Density 0.14 0.06 Modularity (Q) 0.23 0.37 Infomap comunities (size ≥ 2 nodes) 4 7 Disconnected nodes 5 15

Table 5-3. Centrality scores for the nodes in the projected network GS (Figure 5-3B). Degree Centrality (DEC) ranks higher a node that has a high number of connections to other nodes. This metric identifies highly connected species (i.e., important nodes in the network) and is related to the extent of their geographic distribution as well as the number of different taxa recorded per grid cell; Betweenness Centrality (BTC) ranks higher a node that connects along shortest paths with many other nodes. This metric is used to classified species as peripheral, intermediate or central in the network (Ma, et. al., 2016); Eigenvector Centrality (EVC) ranks higher a node if it connects to highly connected nodes. Those nodes not necessarily have a high number of connections but are connected to important nodes in the network. Note that only ranked higher nodes are included here. Species DEC EVC BTC Desmoceras (Desmoceras) latidorsatum 215 0.93 0.62 Phylloceras velledae 193 1.00 0.21 Hysteroceras orbignyi 175 0.93 0.17 Hysteroceras subbinum 164 0.65 0.17 Anagaudryceras sacya 162 0.41 0.83 Douvilleiceras mammillatum 159 0.50 0.72 Anisoceras perarmatum 159 0.77 0.26 Anisoceras armatum 156 0.79 0.18 Dipoloceras (Dipoloceras) cristatum 152 0.70 0.35 Beudanticeras beudanti 146 0.61 0.52 Lechites (Lechites) gaudini 136 0.76 0.15 Douvilleiceras orbignyi 133 0.55 0.18 Puzosia quenstedti 133 0.56 0.18 Phylloceras (Hypophylloceras) seresitense 130 0.69 0.06 Hamites venetzianus 125 0.46 0.15 Tetragonites rectangularis 119 0.43 0.39 Hysteroceras carinatum 119 0.52 0.06 Hysteroceras binum 117 0.52 0.07 Protanisoceras blancheti 115 0.44 0.29 Hamites virgulatus 115 0.31 0.15 Tetragonites timotheanus 114 0.49 0.25 Stoliczkaia (Stoliczkaia) notha 114 0.69 0.04 Hysteroceras varicosum 113 0.55 0.03 Dipoloceras (Dipoloceras) bouchardianum 112 0.54 0.03 Goodhallites goodhalli 111 0.48 1.00 Neophlycticeras (Neophlycticeras) brottianum 110 0.58 0.05 Hysteroceras choffati 109 0.39 0.09 Tetragonites subtimotheanus 104 0.32 0.31 Puzosia (Puzosia) mayoriana 103 0.57 0.05 Salaziceras (Salaziceras) salazacense 100 0.37 0.04

CHAPTER 6 CONCLUSIONS

The assessment of the post-Paleozoic fossil record of the lingulides and associated faunas yielded the following findings: The terebratuloid brachiopods of the

Upper Rosablanca Formation represent a new form, Sellithyris elizabetha nov. sp., and suggest affinity with the Valanginian species S. coahuilensis from the ancient Gulf of

Mexico. Taphonomic semi-quantitative data suggest a large-scale erosional event that may have affected the Upper Rosablanca Formation during Valanginian times. The specimens of Lingularia sp. reported here are the oldest occurrences of Cretaceous lingulides in the region so far. This new material suggests that lingulide brachiopods may have been common locally.

The chapters dedicated to study of drilling predation resulted in the description of a spatially–explicit procedure for analyzing drilling traces on marine shelled invertebrates that was illustrated using naticid-like drillholes on fossil Lirophora. The estimated K-function indicated that predatory drillholes on the prey skeleton were significantly clustered. Those areas where drillholes tend to occur at a significant higher rate than in the rest of the prey skeleton (i.e., umbo, center or both areas) vary across time intervals. This case study illustrates the potential of the point pattern analysis to assess a wide spectrum of ecologic and taphonomical processes on marine shelled invertebrates such as drilling predation, encrustation and bioerosion.

The evaluation of the post-Palaeozoic history of drilling predation on the lingulide brachiopods indicates that they have been subject to drilling predation since at least the

Eocene. Variation in drilling frequencies at the locality level suggests that lingulides may occasionally experience somewhat elevated predation pressures from drilling

organisms. The observed Mesozoic-to-Cenozoic increase in drilling frequencies on lingulides is similar to the trends observed in other marine benthic invertebrates and consistent with the hypothesis that predation pressures increased through time in marine ecosystem.

Although the occurrences of lingulides were insufficient to assess their biogeography, global records of ammonoids in the PaleoDB we used to test the mid-

Cretaceous biogeography using a network-based framework. Overall, the results supported the Boreal-Pacific, Arctic, Tethyan, and Austral bioprovinces. The geographic network derived from ammonoids was twice as dense as the one derived from benthic invertebrates and thus more effective in delineating bioprovinces. The network-based approach described in this study establishes a reproducible quantitative framework for delineating geographic boundaries of marine bioprovinces, and tracking biogeographic changes over evolutionary time scales.

APPENDIX A BIOMETRIC DATA ON SELLITHYRIS COMPILED IN THIS STUDY

Table A-1. Biometric data compiled in this study. Morphological characters measured according to Gaspard and Mullon (1983, Figure 1). Localities for Sellithyris elizabetha nov. sp.: El Sapo, La Virgen, Pico de la Vieja and Cementerio, Zapatoca, Colombia (this study); Sellithyris sella: Isle of Wight, England (MS collection and data); Sellithyris coahuilensis: Coahuila, Mexico (USNM collections and MS data).

ID Section Level LON LUD LAR DIG HMG EPS ECP PSM ECS SLG 1 La Virgen P 24.47 20.52 18.89 17.08 13.89 15.11 6.11 4.88 15.75 11.38 2 La Virgen P 26.03 21.69 19.61 18.84 14.6 17.72 7.81 4.81 15.86 9.43 3 La Virgen P 25.47 20.84 21.32 20.48 15.93 14.85 7.39 4.37 16.41 9.77 4 La Virgen P 23.12 19.97 18.33 15.81 14.69 13.61 5.26 2.85 13.84 7.04 5 La Virgen P 24.89 21.34 20.48 19.2 16.66 13.9 7.72 4.49 15.41 8.69 6 La Virgen P 18.9 16.35 14.95 14.05 11.08 10.47 5.53 1.74 11.85 4.09 7 La Virgen P 26.12 21.03 20.48 18.56 15.96 13.17 8.25 2.8 16.54 7.17 8 La Virgen P 25.8 21.65 20.86 19.36 16.58 14.62 5.53 3.52 15.84 9.19 9 El Sapo P 18.71 16.34 16.31 15.42 12.77 10.42 4.99 1.55 14.12 7.11 10 El Sapo P 20.37 17.34 19.42 15.45 12.35 12.15 6.24 2.77 14.85 8.73 11 El Sapo P 20.38 17.83 20.46 17.67 14.25 11.98 5.91 2.57 17.09 9.19 12 El Sapo P 23.93 19.96 19.96 17.83 14.75 15.37 5.74 5.36 16.81 10.72 13 Pico la vieja P 18.62 16.07 15.06 14.13 12.74 9.85 5.19 2.78 10.2 5.41 14 Pico la vieja P 19.26 16.37 17.36 14.51 11.72 11.07 6.12 2.35 14.51 6.58 15 Pico la vieja P 21.27 17.26 17.35 17.85 15.75 13.64 5.45 5.12 13.37 6.56 16 Pico la vieja P 19.91 16.65 18.44 15.47 12.11 11.71 5.47 2.12 14.05 6.49 17 Pico la vieja P 21.53 17.88 18.88 17.97 15.49 12.8 5.01 3.03 14.51 7.52 18 La Virgen P 26.05 21.66 20.69 19.09 16.75 16.41 6.78 5.39 15.68 8.71 19 La Virgen P 27.22 22.52 23.69 20.53 18.32 16.48 9.58 4 18.19 9.08 20 La Virgen P 28.99 26.1 26.69 21.68 18.52 18.38 8.65 3.76 21.24 12.01 21 La Virgen P 27.02 22.07 24.34 20.93 18.38 16.62 6.38 5.03 15.87 8.85 22 La Virgen P 21.68 18.3 18.64 16.07 14.3 12.73 6.18 2.09 14.7 6.53 23 El Sapo P 17.75 14.62 16 13.39 9.72 9.25 4.58 1.92 11.39 6 24 El Sapo P 18.08 14.81 15.61 14.1 12.6 10.37 4.89 2.02 12.7 5.86 25 El Sapo P 17.64 15.41 14.61 14.61 12.41 9.17 4.9 2.69 11.5 5.27 26 El Sapo P 19.6 16.47 15.1 14.23 12.94 10.97 5.54 2.17 12.11 6.64 27 El Sapo P 18.58 16.03 17.55 14.31 13.39 10.63 4.5 2.8 13.99 7.94 28 El Sapo P 19.64 16.85 18.19 15.31 14.08 11.37 5.69 2.94 15.08 8.71 29 El Sapo P 20.8 17.66 19.23 16.36 12.8 12.15 5.93 3.1 15.75 8.66 30 El Sapo P 20.56 17.76 16.85 15.39 14.8 10.37 4.83 1.8 13.52 8.21 31 La Virgen II P 16.7 14.94 15.49 11.96 11.58 10.08 5.26 1.66 12.63 6.67

Table A-1. Continued.

ID Section Level LON LUD LAR DIG HMG EPS ECP PSM ECS SLG 32 La Virgen II P 19.85 16.27 18.22 15.73 11.82 12.08 5.29 2.55 14.17 6.31 33 La Virgen II P 22.13 19.37 19.13 17.15 14.16 12.15 5.23 2.46 14.79 7.88 34 La Virgen II P 20.18 16.37 16.98 14.78 14.76 12.65 6.02 3.46 13.72 7.56 35 La Virgen II P 23.2 18.8 18.73 18.23 15.93 13.29 5.81 2.77 14.78 7.11 36 — P 22.05 17.93 15.45 17.07 15.9 13.93 5 2.11 12.17 6.62 37 — P 24.53 19.79 19.2 18.11 16.88 14.87 5.41 2.28 15.88 9.72 38 — P 25.38 16.89 19.65 20.41 17.12 14.64 6.5 3.49 14.54 6.41 39 — P 20.52 17.13 18.28 15.7 14.95 11.92 6.21 1.61 13.91 5.78 40 — P 21.47 17.88 19.08 16.41 15.07 13.01 5.93 2.09 14.65 8.57 41 El Sapo P 22.59 19.09 22.74 16.75 13.92 13.81 6.86 3.65 18.74 9.4 42 El Sapo P 22.92 19.53 22.05 17.54 15.79 12.73 5.57 2.39 17.05 9.32 43 El Sapo P 21.33 17.94 18.76 16.22 15.54 11.7 3.88 3.05 13.7 8.53 44 El Sapo P 22.17 19.93 21.51 17.34 16.49 11.32 10.59 3.33 18.97 8.28 45 El Sapo P 19.81 17.56 16.5 15.83 14.75 12.01 5.95 2.93 13.07 6.48 46 El Sapo P 15.76 13.18 14.41 12.14 11.18 8.8 5.29 1.47 11.81 5.42 47 El Sapo P 17.02 14.92 17.27 13.46 12.41 9.62 5.86 1.41 11.62 6 48 El Sapo P 18.53 15.71 16.44 14.68 14.38 10.33 5.09 1.91 13.89 7.01 49 El Sapo P 22.81 18.76 17.44 18.74 17.47 12.54 6.77 3.45 14.61 8.29 50 El Sapo P 19.93 17.13 17.13 15.21 14.84 10.23 4.67 2.25 13.53 7.33 51 El Sapo P 19.45 16.56 17.72 15.77 14.94 10.66 4.95 2.36 14.04 8.26 52 El Sapo P 21.09 18.32 17.23 16.23 15.82 11.54 4.65 2.34 13.9 7.85 53 El Sapo P 21.25 17.19 16.73 13.69 15.27 11.18 6.12 3.55 13.92 7.56 54 El Sapo P 21.43 18.59 19.49 17.52 17.32 13.51 5.05 2.59 15.3 6.89 55 Pico la vieja P 17.89 15.41 17.03 13.6 12.4 9.58 5.72 2.08 11.69 5.63 56 Pico la vieja P 19.52 16.11 16.46 15.87 14.79 10.63 4.59 2.57 10.98 5.95 57 El Sapo P 17.13 14.95 16.44 11.68 12.21 10 6.16 1.63 14.07 4.73 58 El Sapo P 26.27 22.44 21.13 18.56 16.71 13.59 6.19 2.34 16.73 8.01 59 La Virgen P 16.16 14.57 15.38 11.65 11.06 9.61 5.52 1.82 11.19 5.24 60 La Virgen P 22.64 19.1 20.46 18.04 15.86 12.85 5.37 2.74 15.86 9.09 61 La Virgen P 19.05 16.46 16.35 13.9 13.25 10.45 6.84 2.65 12.54 6.53 62 La Virgen P 19.81 16.61 17.69 15.61 13.9 12.18 5.11 4.13 13.51 6.98 63 La Virgen P 21.41 18.48 18.46 16.16 14.01 11.34 5.55 2.41 13.64 5.07 64 La Virgen P 20.78 17.78 18.61 15.8 14.36 12.34 4.47 3.66 12.63 7.03 65 La Virgen P 19.04 15.41 15.82 14.26 12.19 10.57 4.14 2.53 13.35 6.23 66 La Virgen P 21.08 18.22 16.61 15.17 14.06 12.77 6.08 2.95 14.3 7.5 67 La Virgen P 20.61 17.87 19.57 17.14 14.08 11.12 6.24 2.37 15.61 7.8 68 La Virgen P 21.33 17.73 18.55 15.54 13.47 11.92 4.38 3.01 15.01 8.46 69 La Virgen P 21.1 18.22 16.77 15.72 14.41 11.42 5.66 3 13.91 7.19

Table A-1. Continued.

ID Section Level LON LUD LAR DIG HMG EPS ECP PSM ECS SLG 70 La Virgen P 21.35 16.9 17.8 15.63 15.02 10.38 5.12 2.41 13.35 5.67 71 La Virgen P 17.34 14.69 14.64 13.63 13 10.25 5.67 2.61 12.36 5.47 72 La Virgen P 17.7 14.99 15.38 13.81 12.04 9.65 3.56 1.36 11.14 6 73 La Virgen P 18.41 15.78 16.31 14.25 12.97 9.85 4.2 2.05 11.15 6.06 74 La Virgen P 21.72 18.77 19 16.07 15.6 11.55 6.67 2.95 15.61 8.1 75 La Virgen P 22.94 19.54 21.22 18.05 16.1 11.95 4.95 2.58 15.62 9.21 76 La Virgen P 21.4 17.77 19.5 15.98 13.92 11.54 6.91 1.89 17.28 8.41 77 La Virgen P 19.81 16.57 18.08 15.48 13.81 10.94 6.52 2.76 14.43 6 78 La Virgen P 22.14 19.64 20.36 17.27 15.87 12.13 7.3 3.16 14.69 7.23 79 Pico la vieja P 22.16 17.84 19.05 16.99 17.03 11.71 6.79 2.58 17.26 9.16 80 Pico la vieja P 22.23 17.68 19.07 16.9 14.95 11.58 5.9 4.4 15.93 9.23 81 Pico la vieja P 24.09 21.02 19.86 17.75 16.51 12.93 7.7 3.27 15.36 8.55 82 Cementerio P 15.78 13.4 14.55 12.58 11.07 7.86 4.75 1.47 11.28 5.67 83 Cementerio P 18.05 15.45 14.52 14.15 12.08 9.69 4.61 3.53 12.1 4.71 84 Cementerio P 17.75 14.74 15.93 14.42 13.44 10.21 5.03 3.38 13.39 6.81 85 Cementerio P 18.92 16.06 15.78 14.81 12.44 10.38 6.26 3.18 12.67 5.74 86 Cementerio P 21.38 18.12 18.54 17 16.21 11.49 7.31 4.33 15.6 6.18 87 El Sapo P 16.81 14.26 14.84 13.14 11.09 8.26 4.96 1.9 11.04 5.26 88 El Sapo P 18.38 15.53 17.75 15.01 12.82 10.48 4.88 4.07 13.2 7.47 89 El Sapo P 20.44 16.91 17.57 15.69 14.11 10.17 4.92 2.49 13.91 8.43 90 El Sapo P 20.84 17.3 17.23 16.58 16.23 12.38 6.17 4.24 15.53 7.02 91 El Sapo P 22 18.6 19.11 16.1 13.32 11.34 4.5 2.09 15.73 7.72 92 cementerio P 21.32 18.02 19.06 16.96 15.97 12.14 6.34 2.78 15.37 7.76 93 cementerio P 18.87 15.1 17.03 14.61 14.32 10.17 5.69 2.8 13.55 6.29 94 cementerio P 16.08 13.93 15.08 12.86 11.02 8.75 3.31 2.04 10.33 5.82 95 cementerio P 16.38 14.32 14.33 11.89 10.74 8.36 6.66 2.09 11.94 4.71 96 cementerio P 17.2 14.64 15.13 13.89 13.26 9.04 4.54 1.84 11.15 5.43 97 El Sapo P 21.45 18.55 19.01 14.92 13.64 11.74 5.06 1.9 15.48 9.71 98 El Sapo P 21.04 18.15 18.48 15.77 14.23 12.48 5.33 2.74 14.16 8.77 99 El Sapo P 20.87 18.16 18.31 16.43 14.09 11.09 4.62 2.94 13.78 7.23 100 El Sapo P 20.6 18.36 18.63 15.89 13.81 11.36 4.77 3.1 15.81 8.06 101 El Sapo P 18.24 15.92 17.2 13.4 11.29 11.7 4.58 1.38 12.82 6.21 102 La Virgen P 19.01 15.99 16.37 15.01 12.63 11.13 6.2 3.35 15.26 7.75 103 La Virgen P 18.74 16.27 17.16 14.13 12.61 9.97 5.77 2.55 15.4 7.75 104 La Virgen P 17.9 16.32 17.38 13.87 11.89 9.14 5.91 1.96 13.66 5.83 105 La Virgen P 20.89 18.4 17.92 14.77 13.06 11.09 6.62 2.62 14.14 8.68 106 La Virgen P 18.41 16.21 15.57 12.81 11.14 10.02 4.61 1.76 13.84 6.08 107 La Virgen P 19.11 16.01 18.83 13.86 12.49 11.17 4.43 2.88 13.81 7.79

Table A-1. Continued.

ID Section Level LON LUD LAR DIG HMG EPS ECP PSM ECS SLG 108 La Virgen P 18.8 16.2 17.23 14.17 12.21 9.74 5.98 1.6 13.66 5.89 109 La Virgen P 17.3 14.91 16.28 13.27 11.25 10.22 5 1.39 14.22 5.36 110 La Virgen P 26.75 21.51 20.08 19.27 19.23 15.37 6.02 3.52 16.69 11.67 111 La Virgen P 22.68 20.15 20.41 16.32 14.9 12.47 8.1 3.3 17.61 7.57 112 La Virgen P 21.4 18.63 20.17 15.33 13.9 11.42 6.64 1.45 15.16 5.81 113 El Sapo P 23.9 20.09 22.43 17.83 15.31 13.16 7.33 3.42 18.55 10.25 114 El Sapo P 19.94 17.29 18.73 15.67 13.23 11.14 6.32 1.83 15.79 8.04 115 El Sapo P 19.36 16.27 17.67 15.52 13.24 11.59 5.97 2.09 15.06 7.06 116 El Sapo P 17.9 14.86 15.21 13.09 12.42 9.5 4.91 2.74 13.22 7.56 117 El Sapo P 19.77 16.7 16.13 15.07 12.67 10.6 5.81 2.05 13.46 6.57 118 El Sapo P 19.3 17.08 16.03 14.18 12.21 10.8 5.3 2.02 13.6 7.35 119 El Sapo P 18.34 15.83 17.31 14.11 12.58 9.9 4.5 1.74 13.1 5.75 120 El Sapo P 20.73 17.58 18.94 15.36 12.29 11.67 5.5 2.81 14.62 7.78 121 El Sapo P 24.1 20.57 20.44 18.29 14.92 14.05 5.21 3.42 18.18 11.57 122 La Virgen P 17.38 15.63 16.54 12.68 11.83 10.01 6.32 2.16 14.37 7.39 123 La Virgen P 17.7 14.82 15.59 13.07 12.22 9.7 5.77 2.24 13.48 5.82 124 La Virgen P 18.89 16.37 17.15 14.73 13.21 10.12 4.9 2.02 12.73 5.7 125 La Virgen P 18.95 16.15 16.56 14.29 13.79 11.38 5.35 2.01 13.99 7.08 126 La Virgen P 21.36 17.94 18.91 16.82 13.32 12.57 6.47 3.4 16.94 9.1 127 La Virgen P 20.54 17.27 19.47 16.98 14.48 11.89 7.11 2.56 18.43 8.58 128 La Virgen P 23.33 18.48 20.91 18.07 15.59 14.97 6.51 3.94 16.53 10.63 129 La Virgen P 21.13 18.17 16.43 15.08 14.95 12.75 5.07 2.99 13 7.22 130 La Virgen P 26.14 22.68 22.37 18.15 18 15.41 6.9 3.87 17.45 10.11 131 El Sapo P 24.25 20.55 22.29 18.77 16.15 14.48 8.59 2.9 20.13 9.76 132 El Sapo P 19.41 16.85 17.68 14.72 13.58 9.69 5.17 2.92 14.08 6.46 133 El Sapo P 16.3 14.77 15.26 12.75 11.6 9.26 4.17 0.78 11.82 5.31 134 El Sapo P 21.01 18.66 19.98 17.12 14.07 12.39 5.95 3.45 14.25 7.16 135 El Sapo P 19.02 16.9 18.52 14.47 13.62 10.34 5.22 2.94 15.53 8.15 136 El Sapo P 20.31 17.61 17.41 14.83 13.22 11.81 5.26 2.3 14.25 7.78 137 La Virgen P 19.77 17.34 17.97 14.64 13.67 11.37 5.1 1.95 14.65 7.4 138 Pico la vieja P 24.04 22.8 22.23 17.17 15.58 12.76 7.09 3.25 17.11 9.33 139 Pico la vieja P 15.88 14.23 16.25 13.7 11.65 9.27 5.47 2.17 12.38 4.82 140 Pico la vieja P 17.97 14.94 16.82 13.16 11.1 9.78 5.94 2.89 14.6 6.79 141 Pico la vieja P 18.05 15.32 14.25 13.58 12.23 9 5.41 1.43 12.62 7.19 142 La Virgen P 27.2 19.35 22.3 18.24 15.27 13.5 6.39 2.23 17.44 8.95 143 El Sapo P 26.59 22.72 20.34 20 18.96 13.72 6.32 3.47 17.32 9.14 144 El Sapo O 24.16 21.25 19.28 18.92 16.22 14.02 6.49 2.7 16.07 7.96 145 El Sapo O 26.37 23.5 19.3 19.42 17.9 15.69 5.83 2.88 15.61 8.82

Table A-1. Continued.

ID Section Level LON LUD LAR DIG HMG EPS ECP PSM ECS SLG 146 La Virgen O 23.46 20.21 20.55 18.59 16.2 13.9 5.55 3.7 14.97 7.92 147 El Sapo O 25.12 20.65 19.24 20.91 19.64 14.17 6.23 2.93 13.6 6.8 148 El Sapo O 20.7 18.14 17.64 14.9 14.23 10.17 4.86 1.61 12.7 5.41 149 Cementerio O 27.19 23.52 22.55 19.19 16.62 16.48 8.11 3.94 17.8 9.14 150 Cementerio O 24.71 22.08 20.12 18.36 15.61 14.42 8.54 3.36 16.6 6.64 151 Cementerio O 24.16 21.02 20.75 19.02 17.06 13.31 7.78 3.43 17.89 7.77 152 Cementerio O 18.62 16.53 16.61 13.79 12.02 10.39 6.76 3.34 12.63 4.08 153 El Sapo O 26.89 24.13 22.27 20.56 19.2 15.24 7.24 4.24 18.22 10.74 154 El Sapo O 20.56 16.91 18.98 16.01 14.48 11.24 5.43 2.95 15.26 8.66 155 El Sapo O 17.32 14.76 15.93 13.06 11.91 10.44 5.13 1.62 12.58 5.96 156 Coahuila − 23.65 19.73 21.09 16.98 13.55 12.73 7.19 1.22 16.86 8.33 157 MRSC − 21.01 18.23 16.62 15.79 13.98 8.11 5.26 1.21 12.15 4.99 158 MRSC − 20.18 17.84 17.04 15.37 13.56 8.73 5.2 0.51 13.21 4.85 159 MRSC − 22.72 19.82 19.41 17.73 15.08 9.94 6.04 1.27 14.05 5.42 160 MRSC − 21.15 17.91 19.01 16.75 14.28 8.48 3.71 0.56 14.66 6.17 161 MRSC − 21.67 18.11 19.82 13.87 11.01 10.79 5 0.63 13.98 4.33 162 MRSC − 22.22 18.98 17.91 16.68 14.65 11.64 4.4 0.46 14.96 7.78 163 MRSC − 22.76 19.46 20.69 16.24 11.84 9.81 6.75 0.93 16.4 6.87 164 MRSC − 22.13 19.36 21.03 16.82 12.68 10.92 5.64 1.38 16.54 7.72 165 MRSC − 23.46 20.14 20.16 16.94 14.3 10.62 6.55 0.79 16.09 4.44 166 MRSC − 24.33 21.59 22.64 17.5 13.96 11.33 6.91 0.77 15.87 5.43 167 MRSC − 23.71 20.98 22.24 17.79 13.93 12.41 7.66 1.76 18.52 6.6 168 MRSC − 22.65 20.19 19.37 16.27 13.31 10.56 7.54 0.79 13.62 3.65 169 MRSC − 25.24 22.78 23.65 19.54 15.72 11.51 5.95 0.86 17.77 8.71 170 MRSC − 19.46 16.54 15.66 14.26 11.69 8.38 4.11 0.63 10.57 5.4 171 MRSC − 26.47 23.04 22.03 19.56 15.86 14.37 7.77 4.45 20.01 12.33 172 MRSC − 24.01 20.31 21.16 16.39 12.06 11.84 5.86 0.68 17.14 7.37 173 MRSC − 24.89 21.72 20.57 17.19 14.27 11.61 5.86 0.44 16.11 5.69 174 MRSC − 25.25 21.62 20.95 19.21 16.32 12.47 5.31 0.99 15.82 7.99 175 MRSC − 25.66 21.63 23.34 19.66 15.28 13.07 6.83 1.43 20.03 9.24 176 MRSC − 26.36 22.22 21.92 18.92 14.95 12.37 5.72 1.45 16.93 8.58 177 MRSC − 25.92 21.65 24.83 21.07 15.72 12.41 6.42 1.21 18.97 8.83 178 MRSC − 27.53 23.47 23.3 20.54 15.65 12.99 6.84 1.07 19.02 10.37 179 MRSC − 25.62 22.29 23.82 20.22 15.79 12.66 6.82 2.33 19.72 11.33 180 MRSC − 27.13 23.43 22.58 18.66 16.36 12.95 7.86 2.61 18.89 8.74 181 MRSC − 24.95 21.31 24.84 20.91 17.58 12.42 7.35 0.99 18.97 6.18 182 MRSC − 26.97 22.88 24.02 21.32 17.66 13.86 7.95 1.68 20.67 8.47

Table A-1. Continued.

ID Section Level LON LUD LAR DIG HMG EPS ECP PSM ECS SLG 183 MRSC − 26.02 22.09 23.94 20.2 15.82 14.14 9.31 3.11 20.48 8.03 184 MRSC − 25.95 22.92 23.62 19.25 15.11 12.04 7.64 1.18 18.23 7.95 185 MRSC − 24.63 21.65 22.86 19.21 15.49 13.35 7.86 1.2 17.91 5.81 186 MRSC − 27.84 24.41 23.33 21.57 17.99 13.05 7.17 0.92 18.89 7.38 187 MRSC − 27.13 23.66 25.53 21.44 17.53 13.01 8.27 2.72 20.85 9.32 188 MRSC − 26.51 22.4 26.38 21.2 18.2 14.88 7.51 1.1 20.42 10 189 MRSC − 27.62 23.99 24.41 19.5 15.59 14.81 6.33 1.7 19.68 11.16 190 MRSC − 28.48 24.55 24.61 20.76 17.19 14.77 7.31 1.73 20.61 12.28 191 MRSC − 28.25 25.18 26.93 18.86 13.85 13.53 9.36 3.67 22.21 10.91 192 MRSC − 29.17 23.81 26.57 22.77 18.8 14.47 9.85 2.11 22.54 10.65 193 MRSC − 29.51 25.99 24.13 21.43 16.97 13.72 7.66 1.8 20.64 10.12 194 USNM − 24.74 21.33 25.19 21.04 14.71 15 7.87 3.32 19.37 9.85 195 USNM − 19.08 15.87 16.63 13.7 10.64 10.64 6.26 2.37 14.63 9.53 196 USNM − 13.85 11.58 10.59 10.38 8.47 8.18 3.98 0.86 9.2 4.79 197 USNM − 14.42 12.52 12.87 11.25 9.36 7.63 4.11 0.64 10.48 4.71 198 USNM − 16.2 13.51 13.51 12.34 10.92 8.23 3.72 1.32 10.46 6.99 199 USNM − 13.89 12.55 11.92 11.31 8.97 7.16 3.12 0.97 9.35 5.3 200 USNM − 16.81 14.21 15.82 13.42 11.59 9.27 5.96 1.46 13.12 5.39 201 USNM − 15.26 12.68 13.26 10.47 8.44 7.59 4.16 0.82 10.22 6.38 202 USNM − 12.21 10.62 9.37 9.6 9.02 6.42 2.59 0.61 8.76 4.84 203 USNM − 17.28 15.01 14.87 12.58 10.82 8.81 4.37 1.03 11.42 6.84 204 USNM − 18.13 16.13 16.86 13.61 11.03 10.72 5.86 1.08 14.56 6.65 205 USNM − 17.28 13.97 15.34 13.24 10.7 9.68 4.85 1.28 12.86 7.24 206 USNM − 12.92 11.09 11.91 9.75 7.42 6.95 4.39 0.85 9.67 4.46 207 USNM − 14.21 12.4 12.56 11.02 9.75 7.85 3.48 1.31 10.95 5.57 208 USNM − 14.15 11.59 12.34 11.53 9.43 8.27 3.56 0.99 9.53 6.71 209 USNM − 14.37 8.29 13.67 11.29 9.56 8.07 3.34 1.14 10.45 4.85 210 USNM − 12.89 10.99 10.81 8.85 7.71 6.9 3.31 0.71 8.68 4.65 211 USNM − 14.14 12.12 12.83 11.26 9.98 7.78 3.89 1.06 10.87 5.53 212 USNM − 12.07 10.51 11.25 10.23 8.87 6.33 3.53 0.73 9.63 4.73 213 USNM − 14.91 12.43 13.87 11.54 9.71 8.85 4.66 1.21 11.36 6.09 214 USNM − 16.73 14.51 13.28 12.66 10.71 8.15 3.81 1.15 11.35 6.36 215 USNM − 12.61 10.57 10.91 8.56 7.53 7.28 3.61 1.04 9.01 4.95 216 USNM − 11.78 9.91 10.11 9.42 8.11 7.03 3.55 1.15 8.47 5.25 217 USNM − 13.84 11.61 12.82 10.89 9.36 8.05 4.15 1.57 11.27 6.09 218 USNM − 13.17 11.56 11.4 9.52 7.99 7.72 3.84 1.18 10.34 6.62 219 USNM − 12.83 11.09 11.48 9.47 8.43 6.65 4.04 1.11 10.21 5.01

Table A-1. Continued.

ID Section Level LON LUD LAR DIG HMG EPS ECP PSM ECS SLG 220 USNM − 12.03 10.06 10.41 9.17 7.97 7.41 3.09 1.1 8.85 4.82 221 USNM − 10.14 8.96 9.06 8.16 6.62 5.81 1.77 0.73 6.52 3.45 222 USNM − 13.51 12.01 11.99 10.51 8.11 7.68 4.22 1.01 10.53 5.13 223 USNM − 15.07 13.04 12.89 10.18 7.32 9.14 4.23 1.81 10.81 7.23 224 El Sapo − 18.34 16.02 17.05 14.01 11.49 10.09 4.3 2.5 11.9 7.39 225 El Sapo − 16.43 13.52 14.75 13.5 11.75 8.26 4.14 2.21 12.08 5.78

APPENDIX B DRILLING DATA ON CENOZOIC LIROPHORA COMPILED IN THIS STUDY

Table A-2. Shell dimensions of FLMNH specimens used in this study. Abbreviations: n = undrilled specimen, y = drilled specimen, D = drillhole diameter.

Lot # Prey FLMNH-ID Unit Age Valve Bx By size drilled L. glytocyma UF190402 37 5 Oak Grove Sand EM 1 0.36 0.2 L. glytocyma UF190402 37 5 Oak Grove Sand EM 0 0.34 0.22 L. glytocyma UF190402 37 5 Oak Grove Sand EM 1 -0.49 0.1 L. glytocyma UF190402 37 5 Oak Grove Sand EM 0 0.07 0.23 L. glytocyma UF190402 37 5 Oak Grove Sand EM 1 -0.24 -0.34 L. glytocyma UF190401 51 12 Oak Grove Sand EM 1 0.25 0.08 L. glytocyma UF190401 51 12 Oak Grove Sand EM 1 0.34 0.18 L. glytocyma UF190401 51 12 Oak Grove Sand EM 0 0.22 0.3 L. glytocyma UF190401 51 12 Oak Grove Sand EM 1 -0.49 0 L. glytocyma UF190401 51 12 Oak Grove Sand EM 0 0.28 0.2 L. glytocyma UF190401 51 12 Oak Grove Sand EM 0 -0.36 -0.07 L. glytocyma UF190401 51 12 Oak Grove Sand EM 1 0.37 0.22 L. glytocyma UF190401 51 12 Oak Grove Sand EM 0 0.27 0.3 L. glytocyma UF190401 51 12 Oak Grove Sand EM 0 0.21 0.31 L. glytocyma UF190401 51 12 Oak Grove Sand EM 0 0.15 0.28 L. glytocyma UF190401 51 12 Oak Grove Sand EM 0 -0.49 0.03 L. glytocyma UF190401 51 12 Oak Grove Sand EM 1 -0.49 0.03 L. glytocyma UF190392 56 19 Oak Grove Sand EM 1 -0.49 0.3 L. glytocyma UF190392 56 19 Oak Grove Sand EM 0 0.38 0.26 L. glytocyma UF190392 56 19 Oak Grove Sand EM 1 0.34 0.24 L. glytocyma UF190392 56 19 Oak Grove Sand EM 0 0.32 0.22 L. glytocyma UF190392 56 19 Oak Grove Sand EM 1 0.23 0.09 L. glytocyma UF190392 56 19 Oak Grove Sand EM 0 0.43 0.07 L. glytocyma UF190392 56 19 Oak Grove Sand EM 0 0.23 0.27 L. glytocyma UF190392 56 19 Oak Grove Sand EM 0 0.43 0.09 L. glytocyma UF190392 56 19 Oak Grove Sand EM 1 -0.38 -0.2 L. glytocyma UF190392 56 19 Oak Grove Sand EM 0 0.41 0.16 L. glytocyma UF190392 56 19 Oak Grove Sand EM 1 -0.41 -0.14 L. glytocyma UF190392 56 19 Oak Grove Sand EM 1 0.08 0.24 L. glytocyma UF190392 56 19 Oak Grove Sand EM 0 0.3 0.29 L. glytocyma UF190392 56 19 Oak Grove Sand EM 1 0.39 0.12 L. glytocyma UF190392 56 19 Oak Grove Sand EM 1 -0.48 0.12 L. glytocyma UF190392 56 19 Oak Grove Sand EM 0 -0.45 -0.1 L. glytocyma UF190392 56 19 Oak Grove Sand EM 1 -0.5 0.12 L. glytocyma UF190392 56 19 Oak Grove Sand EM 1 0.4 0.13

Table A-2. Continued.

Lot # Prey FLMNH-ID Unit Age Valve Bx By size drilled L. glytocyma UF190392 56 19 Oak Grove Sand EM 0 0.45 0.06 L. glytocyma UF190395 56 13 Oak Grove Sand EM 0 0.41 0.12 L. glytocyma UF190395 56 13 Oak Grove Sand EM 0 0.29 0.25 L. glytocyma UF190395 56 13 Oak Grove Sand EM 0 0.39 0.16 L. glytocyma UF190395 56 13 Oak Grove Sand EM 0 0.3 0.23 L. glytocyma UF190395 56 13 Oak Grove Sand EM 1 0.18 0.15 L. glytocyma UF190395 56 13 Oak Grove Sand EM 1 0.24 0.23 L. glytocyma UF190395 56 13 Oak Grove Sand EM 0 0.16 0.09 L. glytocyma UF190395 56 13 Oak Grove Sand EM 0 0.36 0.2 L. glytocyma UF190395 56 13 Oak Grove Sand EM 0 0.4 0.11 L. glytocyma UF190395 56 13 Oak Grove Sand EM 0 0.17 0.21 L. glytocyma UF190395 56 13 Oak Grove Sand EM 1 0.24 0.19 L. glytocyma UF190395 56 13 Oak Grove Sand EM 0 0.04 0.03 L. glytocyma UF190395 56 13 Oak Grove Sand EM 0 0.05 0.09 L. glytocyma UF190386 29 8 Oak Grove Sand EM 1 0.33 0.18 L. glytocyma UF190386 29 8 Oak Grove Sand EM 0 0.21 0.05 L. glytocyma UF190386 29 8 Oak Grove Sand EM 0 0.02 0.18 L. glytocyma UF190386 29 8 Oak Grove Sand EM 1 -0.21 -0.35 L. glytocyma UF190386 29 8 Oak Grove Sand EM 1 -0.4 -0.18 L. glytocyma UF190386 29 8 Oak Grove Sand EM 1 -0.48 0.01 L. glytocyma UF190386 29 8 Oak Grove Sand EM 1 0.19 0.25 L. glytocyma UF190386 58 8 Oak Grove Sand EM 0 0.4 0.14 L. glytocyma UF190391 58 8 Oak Grove Sand EM 1 0.35 0.18 L. glytocyma UF190391 58 8 Oak Grove Sand EM 0 0.35 0.21 L. glytocyma UF190391 58 8 Oak Grove Sand EM 0 0.29 0.08 L. glytocyma UF190391 58 8 Oak Grove Sand EM 1 0.08 -0.01 L. glytocyma UF190391 58 8 Oak Grove Sand EM 0 0.09 0.31 L. glytocyma UF190391 58 8 Oak Grove Sand EM 0 0.32 0.13 L. glytocyma UF190391 58 8 Oak Grove Sand EM 1 0.06 0.04 L. glytocyma UF190391 58 8 Oak Grove Sand EM 0 0.43 0.11 L. glytocyma UF190390 41 8 Oak Grove Sand EM 0 0.26 0.31 L. glytocyma UF190390 41 8 Oak Grove Sand EM 0 0.26 0.19 L. glytocyma UF190390 41 8 Oak Grove Sand EM 1 0.4 0.18 L. glytocyma UF190390 41 8 Oak Grove Sand EM 1 -0.27 -0.31 L. glytocyma UF190390 41 8 Oak Grove Sand EM 1 -0.46 -0.08 L. glytocyma UF190390 41 8 Oak Grove Sand EM 0 -0.29 0.12 L. glytocyma UF190390 41 8 Oak Grove Sand EM 1 -0.22 -0.36 L. glytocyma UF190393 62 18 Oak Grove Sand EM 1 0.32 0.19 L. glytocyma UF190393 62 18 Oak Grove Sand EM 1 -0.48 0.03 L. glytocyma UF190393 62 18 Oak Grove Sand EM 1 0.2 0.2

Table A-2. Continued.

Lot # Prey FLMNH-ID Unit Age Valve Bx By size drilled L. glytocyma UF190393 62 18 Oak Grove Sand EM 1 0.16 0.39 L. glytocyma UF190393 62 18 Oak Grove Sand EM 0 0.41 0.17 L. glytocyma UF190393 62 18 Oak Grove Sand EM 1 0.37 0.18 L. glytocyma UF190393 62 18 Oak Grove Sand EM 0 0.24 0.14 L. glytocyma UF190393 62 18 Oak Grove Sand EM 1 0.26 0.31 L. glytocyma UF190393 62 18 Oak Grove Sand EM 0 0.14 0.21 L. glytocyma UF190393 62 18 Oak Grove Sand EM 0 0.3 0.25 L. glytocyma UF190393 62 18 Oak Grove Sand EM 1 0.37 0.22 L. glytocyma UF190393 62 18 Oak Grove Sand EM 1 0.1 0.15 L. glytocyma UF190393 62 18 Oak Grove Sand EM 1 0.39 0.24 L. glytocyma UF190393 62 18 Oak Grove Sand EM 1 0.42 0.16 L. glytocyma UF190393 62 18 Oak Grove Sand EM 1 0.35 0.17 L. glytocyma UF190393 62 18 Oak Grove Sand EM 1 0.08 0.12 L. glytocyma UF190393 62 18 Oak Grove Sand EM 1 -0.19 0.05 L. glytocyma UF190393 62 18 Oak Grove Sand EM 1 0.42 0.14 L. latilirata UF30048 12 3 Bermont P 1 -0.04 0.41 L. latilirata UF30048 12 3 Bermont P 0 -0.06 -0.01 L. latilirata UF30048 12 3 Bermont P 1 -0.04 -0.13 L. latilirata UF206594 115 13 Waccamaw P 1 0.12 0.21 L. latilirata UF206594 115 13 Waccamaw P 0 -0.21 -0.03 L. latilirata UF206594 115 13 Waccamaw P 0 -0.22 0.34 L. latilirata UF206594 115 13 Waccamaw P 1 0.13 0 L. latilirata UF206594 115 13 Waccamaw P 0 0.35 0.33 L. latilirata UF206594 115 13 Waccamaw P 0 -0.12 0.19 L. latilirata UF206594 115 13 Waccamaw P 1 -0.09 0.16 L. latilirata UF206594 115 13 Waccamaw P 0 -0.24 0.35 L. latilirata UF206594 115 13 Waccamaw P 0 -0.3 0.23 L. latilirata UF206594 115 13 Waccamaw P 0 -0.11 0.05 L. latilirata UF206594 115 13 Waccamaw P 0 -0.07 0.24 L. latilirata UF206594 115 13 Waccamaw P 1 0.32 0.25 L. latilirata UF217095 55 5 Waccamaw P 1 0.23 0.34 L. latilirata UF217095 55 5 Waccamaw P 1 -0.07 0.13 L. latilirata UF217095 55 5 Waccamaw P 0 0.11 0.12 L. latilirata UF217095 55 5 Waccamaw P 1 0.09 0.22 L. latilirata UF217094 48 5 Waccamaw P 0 0.43 0.26 L. latilirata UF217094 48 5 Waccamaw P 0 -0.11 0.23 L. latilirata UF217094 48 5 Waccamaw P 1 0.09 0.34 L. latilirata UF217094 48 5 Waccamaw P 0 0.34 0.43 L. latilirata UF217094 48 5 Waccamaw P 1 0.26 -0.13 L. latilirata UF217097 48 5 Waccamaw P 0 0.37 0.34

Table A-2. Continued.

Lot # Prey FLMNH-ID Unit Age Valve Bx By size drilled L. latilirata UF217097 48 5 Waccamaw P 0 -0.11 -0.07 L. latilirata UF217097 48 5 Waccamaw P 0 0.36 -0.01 L. latilirata UF217097 48 5 Waccamaw P 1 -0.12 -0.07 L. latilirata UF217097 48 5 Waccamaw P 0 0.34 0.43 L. latilirata UF217099 64 7 Waccamaw P 1 -0.21 0.17 L. latilirata UF217099 64 7 Waccamaw P 1 -0.06 -0.06 L. latilirata UF217099 64 7 Waccamaw P 0 0.12 0.01 L. latilirata UF217099 64 7 Waccamaw P 0 0.4 0.38 L. latilirata UF217099 64 7 Waccamaw P 1 0.03 0.17 L. latilirata UF217099 64 7 Waccamaw P 0 -0.03 0.34 L. latilirata UF217099 64 7 Waccamaw P 0 0.34 0.38 L. latilirata UF217098 56 4 Waccamaw P 0 0.09 0.17 L. latilirata UF217098 56 4 Waccamaw P 0 0.31 -0.08 L. latilirata UF217098 56 4 Waccamaw P 0 0.24 -0.14 L. latilirata UF217098 56 4 Waccamaw P 1 -0.24 0.24 L. latilirata UF206587 55 13 Waccamaw P 0 0.4 0.26 L. latilirata UF206587 55 13 Waccamaw P 0 0.38 0.34 L. latilirata UF206587 55 13 Waccamaw P 0 0.35 -0.07 L. latilirata UF206587 55 13 Waccamaw P 0 0.03 0.24 L. latilirata UF206587 55 13 Waccamaw P 1 0.05 0.22 L. latilirata UF206587 55 13 Waccamaw P 1 0.02 0.2 L. latilirata UF206587 55 13 Waccamaw P 0 -0.07 -0.1 L. latilirata UF206587 55 13 Waccamaw P 0 -0.11 0.24 L. latilirata UF206587 55 13 Waccamaw P 0 -0.15 0.12 L. latilirata UF206587 55 13 Waccamaw P 0 -0.02 0.24 L. latilirata UF206587 55 13 Waccamaw P 0 0.01 -0.07 L. latilirata UF206587 55 13 Waccamaw P 0 -0.01 -0.02 L. latilirata UF206587 55 13 Waccamaw P 0 0.32 0.17 L. latilirata UF2107096 53 5 Waccamaw P 0 -0.05 0.28 L. latilirata UF2107096 53 5 Waccamaw P 0 -0.19 0.62 L. latilirata UF2107096 53 5 Waccamaw P 1 0.19 -0.07 L. latilirata UF2107096 53 5 Waccamaw P 1 0.18 0.39 L. latilirata UF2107096 53 5 Waccamaw P 1 -0.13 0.28 L. latilirata UF137849 23 1 Bermont P 0 0.29 0.29 L. latilirata UF137880 13 1 Bermont P 1 -0.04 0 L. latilirata UF137172 14 1 Bermont P 1 0.1 0.27 L. latilirata UF131622 23 1 Bermont P 1 0.09 0.2 L. latilirata UF131624 26 2 Bermont P 0 -0.03 0.18 L. latilirata UF131624 26 2 Bermont P 0 0.11 0.41 L. latilirata UF131623 24 2 Bermont P 0 -0.13 0.18

Table A-2. Continued.

Lot # Prey FLMNH-ID Unit Age Valve Bx By size drilled L. latilirata UF131623 24 2 Bermont P 0 -0.05 0.08 L. latilirata UF127365 20 2 Bermont P 1 -0.03 -0.09 L. latilirata UF131625 30 3 Bermont P 0 0.15 -0.01 L. latilirata UF131625 30 3 Bermont P 0 0.17 0.26 L. latilirata UF131625 30 3 Bermont P 0 0.26 0.17 L. latilirata UF161845 79 8 Duplin EP 1 -0.13 0.21 L. latilirata UF161845 79 8 Duplin EP 0 0.23 -0.03 L. latilirata UF161845 79 8 Duplin EP 1 -0.48 0.01 L. latilirata UF161845 79 8 Duplin EP 1 0.38 0.16 L. latilirata UF161845 79 8 Duplin EP 1 0.06 0.06 L. latilirata UF161845 79 8 Duplin EP 0 0.05 0.26 L. latilirata UF161845 79 8 Duplin EP 1 -0.24 0.21 L. latilirata UF161845 79 8 Duplin EP 1 -0.02 0.19 L. latilirata UF161844 21 7 Duplin EP 1 0.01 0.34 L. latilirata UF161844 21 7 Duplin EP 1 -0.17 0.26 L. latilirata UF161844 21 7 Duplin EP 0 0.02 0.19 L. latilirata UF161844 21 7 Duplin EP 1 -0.35 -0.22 L. latilirata UF161844 21 7 Duplin EP 0 0.02 0.13 L. latilirata UF161844 21 7 Duplin EP 0 0.04 0.35 L. latilirata UF161844 21 7 Duplin EP 1 0.27 0.06 L. latilirata UF161844 21 7 Duplin EP 0 0.35 0.25 L. latilirata UF164230 29 1 Duplin EP 0 0.03 0.2 L. latilirata UF164234 49 5 Duplin EP 1 -0.15 0.11 L. latilirata UF164234 49 5 Duplin EP 0 0.31 0.15 L. latilirata UF164234 49 5 Duplin EP 0 0.02 -0.08 L. latilirata UF164234 49 5 Duplin EP 0 -0.07 0.21 L. latilirata UF164234 49 5 Duplin EP 1 -0.13 0.2 L. latilirata UF164014 37 2 Duplin EP 0 -0.26 0.26 L. latilirata UF164014 37 2 Duplin EP 0 -0.05 0.15 L. latilirata UF164012 21 4 Duplin EP 0 0.16 0.2 L. latilirata UF164012 21 4 Duplin EP 1 -0.02 0.16 L. latilirata UF164012 21 4 Duplin EP 1 0.07 0.27 L. latilirata UF164012 21 4 Duplin EP 0 0.22 0.35 L. latilirata UF157413 20 5 Jackson Bluff LP 1 0.12 0.14 L. latilirata UF157413 20 5 Jackson Bluff LP 1 0.17 0.23 L. latilirata UF157413 20 5 Jackson Bluff LP 1 0.28 0.02 L. latilirata UF157412 25 1 Jackson Bluff LP 0 -0.16 0.09 L. latilirata UF157411 19 2 Jackson Bluff LP 0 0.23 0.21 L. latilirata UF157411 19 2 Jackson Bluff LP 0 0.21 0.23 L. latilirata UF157410 20 5 Jackson Bluff LP 1 0.29 0.3

100

Table A-2. Continued.

Lot # Prey FLMNH-ID Unit Age Valve Bx By size drilled L. latilirata UF157410 20 5 Jackson Bluff LP 1 0.31 0.19 L. latilirata UF157410 20 5 Jackson Bluff LP 0 0.21 0.03 L. latilirata UF157410 20 5 Jackson Bluff LP 1 0.11 0.07 L. latilirata UF157410 20 5 Jackson Bluff LP 1 0.2 0.18 L. latilirata UF157409 27 9 Bluff LP 0 0.13 0.27 L. latilirata UF157409 27 9 Bluff LP 1 -0.21 0.39 L. latilirata UF157409 27 9 Bluff LP 0 -0.11 0.32 L. latilirata UF157409 27 9 Bluff LP 0 -0.02 0.07 L. latilirata UF157409 27 9 Bluff LP 0 -0.08 -0.19 L. latilirata UF157409 27 9 Bluff LP 0 -0.31 -0.28 L. latilirata UF157409 27 9 Bluff LP 0 0.41 0.32 L. latilirata UF157409 27 9 Bluff LP 1 0.12 0.18 L. latilirata UF157408 20 2 Bluff LP 0 0.25 0.13 L. latilirata UF157408 20 2 Bluff LP 1 0.02 0.26 L. latilirata UF157622 31 2 Bluff LP 0 -0.38 0.51 L. latilirata UF157622 31 2 Bluff LP 0 0.21 0.22 L. latilirata UF157414 20 5 Jackson Bluff LP 1 0.1 0.2 L. latilirata UF157414 20 5 Jackson Bluff LP 1 -0.09 0.14 L. latilirata UF157414 20 5 Jackson Bluff LP 0 0.26 0.26 L. latilirata UF157414 20 5 Jackson Bluff LP 1 0.26 0.22 L. latilirata UF157414 20 5 Jackson Bluff LP 1 0.22 0.13 L. ballista UF132582 9 1 Arcadia M 0 -0.46 -0.07 L. ballista UF189358 9 1 Arcadia M 0 0.27 0.06 L. ballista UF189357 9 1 Arcadia M 0 0.37 0.11 Lirophora sp. UF32169 22 4 Arcadia M 1 0.19 0.23 Lirophora sp. UF32169 22 4 Arcadia M 0 0.2 0.22 Lirophora sp. UF32169 22 4 Arcadia M 0 -0.08 -0.32 Lirophora sp. UF32169 22 4 Arcadia M 0 0.33 0.14 Lirophora sp. UF32181 22 1 Arcadia M 1 -0.19 0.22 L. latilirata UF157413 20 5 Macasphalt Shell LP 1 -0.25 0.14 L. latilirata UF157413 20 5 Macasphalt Shell LP 0 -0.01 0.31 L. latilirata UF145969 25 1 Caloosahatchee P 0 0.16 0.17 L. latilirata UF137856 21 2 Caloosahatchee P 0 -0.15 0.19 L. latilirata UF137856 21 2 Caloosahatchee P 1 0.06 0.01 L. latilirata UF147337 21 8 Pinecrest Beds LP 1 -0.01 0.15 L. latilirata UF147337 21 8 Pinecrest Beds LP 1 0.38 0.21 L. latilirata UF147337 21 8 Pinecrest Beds LP 0 0.12 0.37 L. latilirata UF147337 21 8 Pinecrest Beds LP 0 -0.09 -0.37 L. latilirata UF147337 21 8 Pinecrest Beds LP 1 -0.09 -0.33 L. latilirata UF147337 21 8 Pinecrest Beds LP 1 -0.49 0.01

101

Table A-2. Continued.

Lot # Prey FLMNH-ID Unit Age Valve Bx By size drilled L. latilirata UF147337 21 8 Pinecrest Beds LP 1 0 0.29 L. latilirata UF147337 21 8 Pinecrest Beds LP 0 0.05 0.18 L. latilirata UF137167 13 2 Pinecrest Beds LP 0 0.03 0.08 L. latilirata UF137167 13 2 Pinecrest Beds LP 0 0.02 0.06 L. latilirata UF137167 13 2 Pinecrest Beds LP 1 -0.17 0.37 L. latilirata UF157627 32 6 Pinecrest Beds LP 1 0.46 0.06 L. latilirata UF157627 32 6 Pinecrest Beds LP 0 0.2 0.17 L. latilirata UF157627 32 6 Pinecrest Beds LP 1 0.38 0.14 L. latilirata UF157627 32 6 Pinecrest Beds LP 1 -0.05 0.3 L. latilirata UF157627 32 6 Pinecrest Beds LP 0 0.38 0.15 L. latilirata UF157627 32 6 Pinecrest Beds LP 1 0 0.28 L. latilirata UF146228 42 2 Pinecrest Beds LP 1 0.06 0.32 L. latilirata UF146228 42 2 Pinecrest Beds LP 1 0.26 0.18 L. latilirata UF157628 26 6 Pinecrest Beds LP 0 0.17 0.18 L. latilirata UF157628 26 6 Pinecrest Beds LP 1 -0.2 -0.22 L. latilirata UF157628 26 6 Pinecrest Beds LP 0 0.14 0.33 L. latilirata UF157628 26 6 Pinecrest Beds LP 0 0.34 0.2 L. latilirata UF157628 26 6 Pinecrest Beds LP 1 0.4 0.09 L. latilirata UF157628 26 6 Pinecrest Beds LP 1 0.05 0.23 L. latilirata UF137878 25 4 Pinecrest Beds LP 0 0.03 0.43 L. latilirata UF137878 25 4 Pinecrest Beds LP 0 0.38 0.19 L. latilirata UF137878 25 4 Pinecrest Beds LP 1 0.36 0.22 L. latilirata UF137878 25 4 Pinecrest Beds LP 1 0.45 0.2 L. latilirata UF144502 39 1 Pinecrest Beds LP 1 0.07 0.18 L. latilirata UF147358 35 3 Pinecrest Beds LP 1 0.28 0.2 L. latilirata UF147358 35 3 Pinecrest Beds LP 1 0.34 0.19 L. latilirata UF147358 35 3 Pinecrest Beds LP 1 -0.17 0.42

102

APPENDIX C BODY SIZE DATA OF THE LINGULIDE SPECIMENS SURVEYED AT THE FLORIDA MUSEUM OF NATURAL HISTORY

Table A-3. Shell dimensions of FLMNH specimens used in this study. Abbreviations: n = undrilled specimen, y = drilled specimen, D = drillhole diameter.

sample ID FLMNH L W drilled D (this study) catalog number (mm) (mm) (y/n) S00128 UF-52889 — — n — S00129 UF-40885 — — n — S00130 UF-56278 — — n — S00131 UF-63819-1 — — n — S00131 UF-63819-2 — — n — S00132 UF-39731-1 — 14.5 n — S00132 UF-39731-2 — 21.0 n — S00133 UF-47128 — 18.7 n — S00134 UF-48798-1 — 16.1 n — S00135 UF-52890 40.8 19.0 y 2.9 S00136 UF-39533-1 — — n — S00136 UF-39533-2 — — n — S00137 UF-92111-1 — — n — S00137 UF-92111-2 — — n — S00137 UF-92111-3 — — n — S00137 UF-92111-4 — 15.6 y 2.3 S00138 UF-112933-1 37.2 — n — S00138 UF-112933-2 30.4 — n — S00139 UF-32269-1 — — n — S00139 UF-32269-2 — 11.5 n — S00140 UF-112915-1 — 20.3 n — S00140 UF-112915-2 — 35.0 n — S00141 UF-112860-1 — — n — S00141 UF-112860-2 — — n — S00142 UF-112882 — — n — S00143 UF-112815 — — n — S00144 UF-32272 — — n — S00145 UF-32240 36.1 — n — S00146 UF-245228 — 17.3 n — S00147 UF-155933-1 — 15.1 n — S00147 UF-155933-2 — 17.7 n — S00147 UF-155933-3 — 16.9 n —

103

Table A-3. Continued.

sample ID FLMNH L W drilled D (this study) catalog number (mm) (mm) (y/n) S00147 UF-155933-4 37.4 — n — S00147 UF-155933-5 — — n — S00147 UF-155933-6 — — n — S00147 UF-155933-7 — — n — S00147 UF-155933-8 — 15.1 n — S00147 UF-155933-9 — — n — S00147 UF-155933-10 — — n — S00148 UF-116730 — — n — S00149 UF-128720 — — n — S00150 UF-129544 — — n — S00151 UF-245227 30.6 13.5 n — S00152 UF-245229 22.2 11.2 n — S00153 UF-245230 — — n — S00154 UF-245232 — — n — S00155 UF-245231 — 13.8 n — S00156 UF-177300 28.0 11.6 n — S00157 UF-177321 — — n — S00158 UF-115658-1 — 18.2 n — S00158 UF-115658-2 — 16.5 n — S00158 UF-115658-3 — — n — S00158 UF-115658-4 — — n — S00158 UF-115658-5 — 14.1 n — S00158 UF-115658-6 — 13.1 n — S00158 UF-115658-7 — — n — S00158 UF-115658-8 — 19.0 n — S00158 UF-115658-9 — 20.4 n — S00158 UF-115658-10 — — n — S00159 UF-115657 — 19.0 n — S00160 UF-245259-1 — — n — S00160 UF-245259-2 — — n — S00160 UF-245259-3 — — n — S00160 UF-245259-4 — — n — S00160 UF-245259-5 — — n — S00161 UF-11840 — — y 3.4 S00162 UF-115656-1 13.8 5.5 n — S00162 UF-115656-2 12.3 5.1 n — S00162 UF-115656-3 14.8 5.7 n — S00162 UF-115656-4 12.7 5.1 n — S00162 UF-115656-5 13.1 5.3 n — S00162 UF-115656-6 12.0 4.8 n —

104

Table A-3. Continued.

sample ID FLMNH L W drilled D (this study) catalog number (mm) (mm) (y/n) S00162 UF-115656-7 13.4 5.1 n — S00162 UF-115656-8 13.2 5.2 n — S00162 UF-115656-9 14.1 5.2 n — S00162 UF-115656-10 — 5.1 n — S00162 UF-115656-11 — 5.5 n — S00162 UF-115656-12 — 5.4 n — S00162 UF-115656-13 — 5.3 n — S00162 UF-115656-14 — — n — S00162 UF-115656-15 — — n — S00162 UF-115656-16 — — n — S00162 UF-115656-17 12.6 5.0 n — S00162 UF-115656-18 12.0 4.7 n — S00162 UF-115656-19 14.0 5.5 n — S00162 UF-115656-20 13.5 5.4 n — S00162 UF-115656-21 14.9 5.5 n — S00162 UF-115656-22 12.7 5.0 n — S00162 UF-115656-23 13.2 4.9 n — S00162 UF-115656-24 13.0 5.0 n — S00162 UF-115656-25 11.5 4.8 n — S00162 UF-115656-26 — — n — S00162 UF-115656-27 — — n — S00162 UF-115656-28 — — n — S00162 UF-115656-29 — — n — S00162 UF-115656-30 — — n — S00162 UF-115656-31 — — n — S00162 UF-115656-32 — — n — S00162 UF-115656-33 — — n — S00162 UF-115656-34 — — n — S00162 UF-115656-35 — — n — S00162 UF-115656-36 — — n — S00162 UF-115656-37 — — n — S00162 UF-115656-38 — — n — S00162 UF-115656-39 — — n — S00162 UF-115656-40 — — n — S00162 UF-115656-41 — — n — S00162 UF-115656-42 — — n — S00162 UF-115656-43 — — n — S00162 UF-115656-44 — — n — S00162 UF-115656-45 — — n — S00162 UF-115656-46 — — n —

105

Table A-3. Continued.

sample ID FLMNH L W drilled D (this study) catalog number (mm) (mm) (y/n) S00162 UF-115656-47 — — n — S00162 UF-115656-48 — — n — S00162 UF-115656-49 — — n — S00162 UF-115656-50 — — n — S00162 UF-115656-51 — — n — S00162 UF-115656-52 — — n — S00162 UF-115656-53 — — n — S00162 UF-115656-54 — — n — S00162 UF-115656-55 — — n — S00162 UF-115656-56 — — n — S00162 UF-115656-57 — — n — S00162 UF-115656-58 — — n — S00162 UF-115656-59 — — n — S00162 UF-115656-60 — — n — S00162 UF-115656-61 — — n — S00162 UF-115656-62 — — n — S00162 UF-115656-63 — — n — S00162 UF-115656-64 — — n — S00162 UF-115656-65 — — n — S00162 UF-115656-66 — — n — S00162 UF-115656-67 — — n — S00162 UF-115656-68 — — n — S00162 UF-115656-69 — — n — S00162 UF-115656-70 — — n — S00162 UF-115656-71 — — n — S00162 UF-115656-72 — — n — S00162 UF-115656-73 — — n — S00162 UF-115656-74 — — n — S00162 UF-115656-75 — — n — S00162 UF-115656-76 — — n — S00162 UF-115656-77 — — n — S00162 UF-115656-78 — — n — S00162 UF-115656-79 — — n — S00162 UF-115656-80 — — n — S00162 UF-115656-81 — — n — S00162 UF-115656-82 — — n — S00162 UF-115656-83 — — n — S00162 UF-115656-84 — — n — S00162 UF-115656-85 — — n — S00162 UF-115656-86 — — n —

106

Table A-3. Continued.

sample ID FLMNH L W drilled D (this study) catalog number (mm) (mm) (y/n) S00162 UF-115656-87 — — n — S00162 UF-115656-88 — — n — S00162 UF-115656-89 — — n — S00162 UF-115656-90 — — n — S00162 UF-115656-91 — — n — S00162 UF-115656-92 — — n — S00162 UF-115656-93 — — n — S00162 UF-115656-94 — — n — S00162 UF-115656-95 — — n — S00162 UF-115656-96 — — n — S00162 UF-115656-97 — — n — S00162 UF-115656-98 — — n — S00162 UF-115656-99 — — n — S00162 UF-115656-100 — — n — S00163 UF-30936-1 — — n — S00163 UF-30936-2 — — n — S00163 UF-30936-3 — — n —

107

LIST OF REFERENCES

Abrahao, B., Soundarajan, S., Hopcroft, J., and Kleinberg, R., 2014, A separability framework for analyzing community structure: ACM Transactions on Knowledge Discovery from Data, v. 8, p. 1–29, doi: 10.1145/2527231.

Abrahao, B., Soundarajan, S., Hopcroft, J., and Kleinberg, R., 2012, On the separability of structural classes of communities, in ACM Press, p. 624, doi: 10.1145/2339530.2339631.

Aguilar, T., and Alvarado, G.E., 2004, Tafonomía y sedimentología de la Paleobiota Estuarina en los Cerros Barbudal (Costa Rica) sepultada por vulcanismo neógeno: Revista Geológica de América Central, v. 30, p. 111–, doi: 10.15517/rgac.v0i30.7276.

Aguirre-Urreta, M.B., Price, G.D., Ruffell, A.H., Lazo, D.G., Kalin, R.M., Ogle, N., and Rawson, P.F., 2008, Southern Hemisphere Early Cretaceous (Valanginian-Early Barremian) carbon and oxygen isotope curves from the Neuquén Basin, Argentina: Cretaceous Research, v. 29, p. 87–99, doi: 10.1016/j.cretres.2007.04.002.

Alzahrani, T., and Horadam, K.J., 2016, Community Detection in Bipartite Networks: Algorithms and Case studies, in Lü, J., Yu, X., Chen, G., and Yu, W. eds., Complex Systems and Networks, Berlin, Heidelberg, Springer Berlin Heidelberg, p. 25–50.

Amédro, F., and Robaszynski, F., 2005, Corrélation des successions de l’Albien par les ammonites entre la Province Nord-Pacifique et les Provinces européenne et arctique du Domaine boréal: zonation, eustatisme et paléobiogéographie: Geobios, v. 38, p. 585–607, doi: 10.1016/j.geobios.2004.04.004.

Ansell, A., and Morton, B., 1985, Aspects of naticid predation in Hong Kong with special reference to the defensive adaptations of Bassina (Callanaitis) calophylla (Bivalvia). (B. Morton & D. Dudgeon, Eds.): Hong Kong, Hong Kong University Press, Proceedings of the Second International Workshop on the Malacofauna of Hong Kong and Southern China, 635-660 p.

Aragó, P., Juan, P., Díaz-Avalos, C., and Salvador, P., 2016, Spatial point process modeling applied to the assessment of risk factors associated with forest wildfires incidence in Castellón, Spain: European Journal of Forest Research, v. 135, p. 451–464, doi: 10.1007/s10342-016-0945-z.

Århus, N., Kelly, S.R.A., Collins, J.S.H., and Sandy, M.R., 1990, Systematic palaeontology and biostratigraphy of two Early Cretaceous condensed sections from the Barents Sea: Polar Research, v. 8, p. 165–194, doi: 10.3402/polar.v8i2.6811.

108

Asratian, A.S., Denley, T.M.J., and Häggkvist, R., 1998, Bipartite graphs and their applications: Cambridge, Cambridge University Press, Cambridge tracts in mathematics 131, 259 p.

Ausich, W.I., and Gurrola, R.A. Two boring organisms in a Lower Mississippian Community of Southern Indiana: Journal of Paleontology, v. 53, p. 335–344.

Bachnou, A., and Boughdiri, M., 2005, Morphométrie des brachiopodes par modélisation mathématique combinée des tracés ouvert et fermé. Exemple dʼapplication parmi les Terebratulidae du Jurassique moyen: Revue de Paléobiologie, v. 24, p. 789–802.

Baddeley, A., Rubak, E., and Turner, R., 2016, Spatial point patterns: methodology and applications with R: New York, CRC Press, Taylor & Francis Group, Champan & Hall/CRC Interdisciplinary Statistics Series, 810 p.

Baddeley, A., and Turner, R., 2005, Spatstat: An R package for analyzing spatial point patterns: Journal of Statistical Software, v. 12, p. 1–42.

Ballesteros, I., and Nivia, A., 1985, La Formación Ritoque: registro sedimentario de una albufera de comienzos del Cretácico, in Etayo-Serna, F. and Laverde-Montano, F. eds., Proyecto Cretácico, Publicaciones Geológicas Especiales del Ingeominas, v. 16, p. XIV 1-17.

Baumiller, T.K., 1993, Boreholes in Devonian blastoids and their implications for boring by platyceratids: Lethaia, v. 26, p. 41–47, doi: 10.1111/j.1502- 3931.1993.tb01508.x.

Baumiller, T.K., 1996, Boreholes in the Middle Devonian blastoid Heteroschisma and their implications for gastropod drilling: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 123, p. 343–351, doi: 10.1016/0031-0182(95)00124-7.

Baumiller, T., Aleksandra Bitner, M., and Emig, C., 2006, High frequency of drill holes in brachiopods from the Pliocene of Algeria and its ecological implications: Lethaia, v. 39, p. 313–320, doi: 10.1080/00241160600847546.

Benest, M., Gaspard, D., and Ghali, M., 1996, Les brachiopodes de l’avant-pays ouest- algérien lors de la phase transgressive maximale du Berriasien supérieur; systématique, environnement, paléogéographie: Geobios, v. 29, p. 13–34, doi: 10.1016/S0016-6995(96)80068-4.

Bengtson, P., and Kakabadze, M.V., 1999, Biogeography of Cretaceous ammonites - a review of procedures and problems: Neues Jahrbuch fur Geologie und Palaontologie - Abhandlungen, v. 212, p. 221–239.

Bice, K.L., and Norris, R.D., 2002, Possible atmospheric CO2 extremes of the Middle Cretaceous (late Albian-Turonian): Paleoceanography, v. 17, p. 22-1-22–17, doi: 10.1029/2002PA000778.

109

Biernat, G., and Emig, C.C., 1993, Anatomical distinctions of the Mesozoic lingulide brachiopods: Acta Paleontologica Polonica, v. 38, p. 1–20.

Bitner, M.A., 1996, Brachiopods from the Eocene La Meseta Formation of Seymour Island, Antarctic Peninsula: Palaeontologia polonica, v. 55, p. 65–100.

Bivand, R., and Gómez-Rubio, V., 2008, Applied spatial data analysis with R: New York, Springer, Use R, 378 p.

Blondel, V.D., Guillaume, J.-L., Lambiotte, R., and Lefebvre, E., 2008, Fast unfolding of communities in large networks: Journal of Statistical Mechanics: Theory and Experiment, v. 2008, p. P10008, doi: 10.1088/1742-5468/2008/10/P10008.

Bogdanova, T.N., and Hoedemaeker, P.J., 2004, Barremian – early Albian Deshayesitidae, Oppeliidae, Desmoceratidae and Silesitidae of Colombia: Scripta Geologica, v. 128, p. 183–313.

Bolstad, P., 2008, GIS fundamentals: A first text on geographic information systems: Minnesota, Eider Press, 620 p.

Bookstein, F.L., 1986, Size and shape spaces for landmark data in two dimensions: Statistical Science, v. 1–2, p. 181–222.

Borgatti, S.P., and Everett, M.G., 1997, Network analysis of 2-mode data: Social Networks, v. 19, p. 243–269.

Brayard, A., Escarguel, G., and Bucher, H., 2007, The biogeography of Early Triassic ammonoid faunas: Clusters, gradients, and networks: Geobios, v. 40, p. 749– 765, doi: 10.1016/j.geobios.2007.06.002.

Brayard, A., Escarguel, G., Monnet, C., Jenks, J.F., and Bucher, H., 2015, Biogeography of Triassic Ammonoids, in Klug, C., Korn, D., De Baets, K., Kruta, I., and Mapes, R.H. eds., Ammonoid Paleobiology: From macroevolution to paleogeography, Dordrecht, Springer Netherlands, v. 44, p. 163–187, http://link.springer.com/10.1007/978-94-017-9633-0_7

Brezina, S.S., Cech, N., Martín Serralta, D., and Casadío, S., 2016, Cannibalism in Naticidae from the La Meseta Formation (Eocene, Antarctica): Antarctic Science, v. 28, p. 205–215, doi: 10.1017/S0954102015000656.

Briggs, J.C., 1988, Biogeography and Plate Tectonics.: Burlington, Elsevier, 203 p.

Bromley, R.G., 1981, Concepts in ichnotaxonomy illustrated by small round holes in shells: Acta Geológica Hispánica, v. 16, p. 55–64.

Bürgl, H., 1956, Catálogo de los amonitas de Colombia. Parte I. Pulchelliidae: Boletín de Geología, v. IV, p. 1–119.

110

Cadena, E.A., 2011a, First Record of Eucryptodiran Turtles from the Early Cretaceous (Valanginian), at the Northernmost Part of South America: South American Journal of Herpetology, v. 6, p. 49–53, doi: 10.2994/057.006.0107.

Cadena, E.A., 2011b, First Record of Eucryptodiran Turtles from the Early Cretaceous (Valanginian), at the Northernmost Part of South America: South American Journal of Herpetology, v. 6, p. 49–53, doi: 10.2994/057.006.0107.

Cadena, E., 2015, The first South American sandownid turtle from the Lower Cretaceous of Colombia: PeerJ, v. 3, p. e1431, doi: 10.7717/peerj.1431.

Calvet, C., 1992, Borehole site-selection in Naticarius hebraeus. (Chemnitz in Karsten, 1769) (Naticidae: Gastropoda): Orsis, v. 7, p. 57–64.

Calzada-Badia, S., 1975, Braquiópodos infracretácicos del Levante español: Trabajos del museo geológico del seminario C. de Barcelona, v. 14, p. 1–71.

Campbell, M.R., Campbell, D.C., Campbell, L.D., and Portell, R.W., 1997, Distribution and paleoecology of Glottidia inexpectans Olsson, 1914 (Brachiopoda: Lingulidae): Tulane Studies in Geology and Paleontology, v. 30, p. 159–170.

Cardozo, E.P., and Ramirez, C.Q., 1985, Ambientes de depósito de la Formación Rosablanca: area de Villa de Leiva, in Etayo-Serna, F. and Laverde-Montano, F. eds., Proyecto Cretácico, Publicaciones Geológicas Especiales del Ingeominas, v. 16, p. XIII 1-13.

Carrillo-Briceño, J.D., Cadena, E.A., Dececchi, A.T., Larson, H.C.E., and Du, T.Y., 2016, First record of a hybodont shark (Chondrichthyes: Hybodontiformes) from the Lower Cretaceous of Colombia: Neotropical Biodiversity, v. 2, p. 81–86, doi: 10.1080/23766808.2016.1191749.

Casey, R., 1961, The stratigraphical palaeontology of the Lower Greensand: Palaeontology, v. 3, p. 487–621.

Catzigras, F., 1948, Sur une espèce hauterivienne de Térébratulideés suisse et méditerranéenne: Bulletin de la Société Géologique de France, v. 18 Série 5, p. 391–394.

Chatterton, B.D.E., and Whitehead, H.L., 1987, Predatory borings in the inarticulate brachiopod Artiotreta from the Silurian of Oklahoma: Lethaia, v. 20, p. 67–74, doi: 10.1111/j.1502-3931.1987.tb00762.x.

Chattopadhyay, D., Rathie, A., Miller, D.J., and Baumiller, T.K., 2014, Hydrodynamic effects of drill holes on postmortem transportation of bivalve shells and its taphonomic implications: Palaios, v. 28, p. 875–884, doi: 10.2110/palo.2013.029.

Chiba, T., and Sato, S., 2012, Size-selective predation and drillhole-site selectivity in Euspira fortunei (Gastropoda: Naticidae): implications for ecological and

111

palaeoecological studies: Journal of Molluscan Studies, v. 78, p. 205–212, doi: 10.1093/mollus/eys002.

Chuang, S.H., 1964, The affinity of Lingula dumortieri Nyst with Glottidia: Journal of Paleontology, v. 38, p. 155–157.

Claude, J., 2008, Morphometrics with R: New York, Springer.

Clauset, A., Newman, M.E.J., and Moore, C., 2004, Finding community structure in very large networks: Physical Review E, v. 70, doi: 10.1103/PhysRevE.70.066111.

Cooper, A., 1988, Some Tertiary Brachiopods of the East Coast of the United States: Smithsonian Contributions to Paleobiology, v. 64, p. 1–45.

Cooper, G.A., 1983, Terebratulacea (Brachiopoda), Triassic to Recent: A Study of the Brachidia (Loops): Smithsonian Contributions to Paleobiology, p. 1–445, doi: 10.5479/si.00810266.50.1.

Crame, J.A., Lomas, S.A., Pirrie, D., and Luther, A., 1996, Late Cretaceous extinction patterns in Antarctica: Journal of the Geological Society, v. 153, p. 503–506, doi: 10.1144/gsjgs.153.4.0503.

Crame, J.A., Pirrie, D., and Riding, J.B., 2015, Mid-Cretaceous stratigraphy of the James Ross Basin, Antarctica: Geological Society, London, Special Publications, v. 258, p. 7–19, doi: 10.1144/GSL.SP.2006.258.01.02.

Csardi, G., and Nepusz, T., 2006, The igraph software package for complex network research: InterJournal Complex Systems, v. 1695.

Danon, L., Díaz-Guilera, A., Duch, J., and Arenas, A., 2005, Comparing community structure identification: Journal of Statistical Mechanics: Theory and Experiment, v. 2005, p. P09008–P09008, doi: 10.1088/1742-5468/2005/09/P09008.

Darlington, P.J., 1957, Zoogeography: The Geographical Distribution of Animals: New York, John Wiley & Sons, 675 p.

Dera, G., Neige, P., Dommergues, J.-L., and Brayard, A., 2011, Ammonite paleobiogeography during the Pliensbachian–Toarcian crisis (Early Jurassic) reflecting paleoclimate, eustasy, and extinctions: Global and Planetary Change, v. 78, p. 92–105, doi: 10.1016/j.gloplacha.2011.05.009.

Dêtre, C.S., 1968, The Development of the commissure of the Lower Cretaceous Terebratellid Sellithyris sella (J de C. Sowerby) in the course of ontogeny: Evi Jelentése Magyar Fold- tani Intézet, p. 353–378.

Dieni, I., Middlemiss, F.A., and Owen, E.F., 1973, The lower Cretaceous brachiopods of east-central Sardinia: Bollettino della Società Paleontologica Italiana, v. 12, p. 166–216.

112

Dietl, G.P., and Alexander, R.R., 2000, Post-Miocene Shift in Stereotypic Naticid Predation on Confamilial Prey from the Mid-Atlantic Shelf: Coevolution with Dangerous Prey: Palaios, v. 15, p. 414–429.

Dietl, G.P., Alexander, R.R., Kelley, P., and Hansen, T., 2001, Stereotypy of naticid predation on bivalves since the cretaceous: trends, controlling factors, and implications for escalation: PaleoBios, v. 21, p. 78.

Dietrich, W., 1938, Lamelibranquios cretácicos de la Cordillera Oriental, in Estudios Geológicos y Paleontológicos sobre la Cordillera Oriental de Colombia, Bogota, Ministerio de Industria y Petrologia, v. 2, p. 81–108.

Ding, Z., Zhang, X., Sun, D., and Luo, B., 2016, Overlapping Community Detection based on Network Decomposition: Scientific Reports, v. 6, p. 24115, doi: 10.1038/srep24115.

Ditchfield, P.W., Marshall, J.D., and Pirrie, D., 1994, High latitude palaeotemperature variation: New data from the Thithonian to Eocene of James Ross Island, Antarctica: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 107, p. 79– 101, doi: 10.1016/0031-0182(94)90166-X.

Dos Santos, D., Fernandez, H., Cuezzo, M.G., and Dominguez, E., 2008, Sympatry Inference and Network Analysis in Biogeography: Systematic Biology, v. 57, p. 432–448, doi: 10.1080/10635150802172192.

Dos Santos, D.A., Gabriela Cuezzo, M., Celina Reynaga, M., and Dominguez, E., 2012, Towards a Dynamic Analysis of Weighted Networks in Biogeography: Systematic Biology, v. 61, p. 240–252, doi: 10.1093/sysbio/syr098.

Doyle, P., and Pirrie, D., 1999, Belemnite Distribution Patterns, in Olóriz, F. and Rodríguez-Tovar, F.J. eds., Advancing Research on Living and Fossil Cephalopods, Boston, MA, Springer US, p. 419–436.

Dulai, A., 2013, Sporadic Miocene brachiopods in the Naturalis Biodiversity Center (Leiden, the Netherlands): Records from the Mediterranean, the North Sea, and the Atlantic Ocean: Fragmenta Palaeontologica Hungarica, v. 30, p. 15–51.

Dunhill, A.M., Bestwick, J., Narey, H., and Sciberras, J., 2016c, Dinosaur biogeographical structure and Mesozoic continental fragmentation: a network- based approach: Journal of Biogeography, v. 43, p. 1691–1704, doi: 10.1111/jbi.12766.

Emig, C.C., 1986, Conditions de fossilisation du genre Lingula (Brachiopoda) et implications paléoécologiques: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 53, p. 245–253, doi: 10.1016/0031-0182(86)90061-1.

Emig, C., 1997, Ecology of inarticulated brachiopods and biogeography of the inarticulated brachiopods, in Kaesler, R.L. ed., Treatise on invertebrate

113

paleontology. Part H, Brachiopoda revised, Lawrence, Geological Society of America and University of Kansas Press, v. 1, p. 471–526.

Emig, C.C., 1990, Examples of post-mortality alteration in Recent brachiopods shells and (paleo) ecological consequences: Marine Biology, v. 104, p. 233–238.

Emig, C.C., 1983, Taxonomie du genre Glottidia (Bachiopodes, Inarticulés): Bulletin du Muséum national d’histoire naturelle, v. 4, p. 469–489.

Emig, C.C., and Bitner, M.A., 2005, Glottidia (Brachiopoda: Lingulidae) from the Eocene la Meseta Formation, Seymour Island, Antarctica: Palaeontology, v. 48, p. 423– 431, doi: 10.1111/j.1475-4983.2005.00454.x.

Emig, C.C., and Bitner, M.A. The brachiopod Lingula in the Middle Miocene of the Central Paratethys: Acta Palaeontologica Polonica, v. 50, p. 181–184.

Emig, C.C., and Vargas, J., 1990, Glottidia audebarti (Broderip) (Brachiopoda, Lingulidae) from the Gulf of Nicoya, Costa Rica: Revista de Biologia Tropical, v. 38, p. 251–258.

Etayo-Serna, F., 1968a, Apuntaciones acerca de algunos ammonites interesantes en el Hauteriviano y del Barremiano, de la region Villa de Leiva (Boyacá, Colombia S.A.): Boletín de Geologia, p. 51–70.

Etayo-Serna, F., 1968b, El sistema cretáceo de la región de Villa de Leiva y zonas próximas: Revista de Geología Colombiana, p. 5–74.

Etayo-Serna, F., 1964, Posición de las faunas en los depósitos cretácicos colombianos y su valor en la subdivisión cronológica de las mismas: Boletín de Geologia, p. 5–142.

Etayo-Serna, F., 1979, Zonation of the Cretaceous of central Colombia by ammonites: Publicación Especial Ingeominas, v. 2, p. 1–186.

Etayo-Serna, F., Renzoni, G., and Barrero, D., 1976, Contornos sucesivos del mar Cretáceo en Colombia, in Etayo-Serna, F. and Cáceres-Giron, C. eds., Memoria del Primer Congreso Colombiano de Geología, Agosto 4-8 de 1969, p. 217–252.

Everett, M.G., and Borgatti, S.P., 2013, The dual-projection approach for two-mode networks: Social Networks, v. 35, p. 204–210, doi: 10.1016/j.socnet.2012.05.004.

Everitt, B., 2011, An introduction to applied multivariate analysis with R: New York, Springer, 273 p.

Fitton, W.H., 1847, A stratigraphical account of the section from Atherfield to Rocken- end, on the south-west coast of the Isle of Wight: Quarterly Journal of the Geological Society of London, p. 289–327.

114

Fluteau, F., Ramstein, G., Besse, J., Guiraud, R., and Masse, J.., 2007, Impacts of palaeogeography and sea level changes on Mid-Cretaceous climate: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 247, p. 357–381, doi: 10.1016/j.palaeo.2006.11.016.

Fortunato, S., 2010, Community detection in graphs: Physics Reports, v. 486, p. 75– 174, doi: 10.1016/j.physrep.2009.11.002.

Fotheringham, A.S., Brunsdon, C., and Charlton, M., 2000, Quantitative geography: perspectives on spatial data analysis: London, Sage Publications, 288 p.

García Molinos, J., Halpern, B.S., Schoeman, D.S., Brown, C.J., Kiessling, W., Moore, P.J., Pandolfi, J.M., Poloczanska, E.S., Richardson, A.J., and Burrows, M.T., 2015, Climate velocity and the future global redistribution of marine biodiversity: Nature Climate Change, v. 6, p. 83–88, doi: 10.1038/nclimate2769.

García-Ramos, D., 2009, Relevancia en Sistemática del estudio mediante secciones seriadas del cardinalio de los Terebratuloidea y Loboidothyridoidea (Brachiopoda) cretácicos: Asociación Cultural Paleontológica, p. 34–72.

Gaspard, D., 1997, Distribution and recognition of phases in Aptian-Turonian [Cretaceous] Brachiopod development in NW Europe: v. 48, p. 145–161.

Gaspard, D., 1999, Distribution of Lower Cretaceous brachiopods in Europe (Berriasian to Aptian): Bulletin de la Societe Geologique de France, v. 170, p. 311–326.

Gaspard, D., 1988, Sellithyridinae, Terebratulidae du Crétacé d’Europe Occidentale – Dynamique des populations – Systématique et Evolution: Paris, CNRS, Cahiers de Paléontologie, 243 p.

Gaspard, D., and Mullon, C., 1983, Méthodes de discrimination ap- pliquées à des populations de Sellithyridinae (Brachiopodes) du Cénomanien d’Europe Occidentale: Bulletin de la Société Géologique de France, v. 7, p. 679–688.

Girvan, M., and Newman, M.E.J., 2002, Community structure in social and biological networks: Proceedings of the National Academy of Sciences, v. 99, p. 7821– 7826, doi: 10.1073/pnas.122653799.

Gómez-Cruz, A., Bermúdez, H.D., and Vega, F., 2015, A new species of Diaulax Bell, 1863 (Brachyura: Dialucidae) in the Early Cretaceous of the Rosablanca Formation, Colombia: Boletín de la Sociedad Geológica Mexicana, v. 67, p. 103– 112.

Gordillo, S., Bayer, M.S., Boretto, G., and Charó, M., 2014, Taphonomy, in Mollusk shells as bio-geo-archives, Cham, Springer International Publishing, p. 7–14, doi: 10.1007/978-3-319-03476-8_2.

Guerrero, J., 2002a, A proposal on the classification of systems tracts: application to the

115

allostratigraphy and sequence stratigraphy of the Cretaceous Colombian Basin. Part 1: Barremian to Maastrichtian: Geología Colombiana, p. 27–49.

Guerrero, J., 2002b, A proposal on the classification of systems tracts: application to the allostratigraphy and sequence stratigraphy of the Cretaceous Colombian Basin. Part 1: Berriasian to Hauterivian: Geología Colombiana, p. 3–25.

Guillaume, J.-L., and Latapy, M., 2004, Bipartite structure of all complex networks: Information Processing Letters, v. 90, p. 215–221, doi: 10.1016/j.ipl.2004.03.007.

Guzman, G., 1985, Los grifeidos infracretácicos Aetostreon couloni y Ceratostreon boussingaulti de la Formación Rosablanca, como indicadores de oscilaciones marinas, in Etayo-Serna, F. and Laverde, F. eds., Proyecto Cretácico, Bogota, Publicaciones Geológicas Especiales del Ingeominas 16, p. XII 1-16.

Hallam, A., 1983, Early and Mid-Jurassic molluscan biogeography and the establishment of the Central Atlantic Seaway: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 43, p. 181–193, doi: 10.1016/0031- 0182(83)90010-X.

Hansen, T., and Surlyk, F., 2014, Marine macrofossil communities in the uppermost Maastrichtian chalk of Stevns Klint, Denmark: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 399, p. 323–344, doi: 10.1016/j.palaeo.2014.01.025.

Harper, E.M., 2016, Uncovering the holes and cracks: from anecdote to testable hypotheses in predation studies (A. Smith, Ed.): Palaeontology, v. 59, p. 597– 609, doi: 10.1111/pala.12255.

Harper, E.M., Forsythe, G.T.W., and Palmer, T., 1998, Taphonomy and the Mesozoic Marine Revolution: Preservation State Masks the Importance of Boring Predators: Palaios, v. 13, p. 352, doi: 10.2307/3515323.

Harris, J.E., Parkyn, D.C., and Murie, D.J., 2005, Distribution of Gulf of Mexico Sturgeon in Relation to Benthic Invertebrate Prey Resources and Environmental Parameters in the Suwannee River Estuary, Florida: Transactions of the American Fisheries Society, v. 134, p. 975–990, doi: 10.1577/T04-100.1.

Hartigan, J.A., 1975, Clustering algorithms: New York, Wiley, Wiley series in probability and mathematical statistics, 351 p.

Henderson, R.A., 1990, Late Albian ammonites from the Northern Territory, Australia: Alcheringa: An Australasian Journal of Palaeontology, v. 14, p. 109–148, doi: 10.1080/03115519008527815.

Henderson, R.A., and McKenzie, E.D., 2002, Idanoceras, a New Heteromorph Ammonite Genus from the Late Albian of Eastern Australia: Journal of Paleontology, v. 76, p. 906, doi: 10.1666/0022-

116

3360(2002)076<0906:IANHAG>2.0.CO;2.

Hoffmeister, A.P., Kowalewski, M., Baumiller, T.K., and Bambach, R.K., 2004, Drilling predation on Permian brachiopods and bivalves from the Glass Mountains, west Texas: Acta Palaeontologica Polonica, v. 49, p. 443–454.

Holmer, L.E., and Bengtson, P., 2009, The first occurrence of a lingulid brachiopod from the Cretaceous of Sergipe, Brazil, with a restudy of “Lingula” bagualensis Wilckens, 1905 from southern Patagonia: Paläontologische Zeitschrift, v. 83, p. 255–266, doi: 10.1007/s12542-009-0024-6.

Holmer, L.E., and Popov, L.E., 2000, Lingulida, in Kaesler, R.L. ed., Revised Brachiopoda. -Treatise on Invertebrate Paleontology Part H, Lawrence, Geological Society of America and University of Kansas, v. 2, p. 32–95.

Hothorn, T., and Everitt, B., 2014, A handbook of statistical analyses using R: Boca Raton, CRC Press, Taylor & Francis Group, 421 p.

Hughes, R., 1985, Predatory behavior of Natica unifasciata feeding intertidally on gastropods: Journal of Molluscan Studies, v. 51, p. 331–335.

Huntley, J.W., and Kowalewski, M., 2007, Strong coupling of predation intensity and diversity in the Phanerozoic fossil record: Proceedings of the National Academy of Sciences, v. 104, p. 15006–15010, doi: 10.1073/pnas.0704960104.

Iba, Y., Mutterlose, J., Tanabe, K., Sano, S. -i., Misaki, A., and Terabe, K., 2011, Belemnite extinction and the origin of modern cephalopods 35 m.y. prior to the Cretaceous-Paleogene event: Geology, v. 39, p. 483–486, doi: 10.1130/G31724.1.

Ifrim, C., Lehmann, J., and Ward, P., 2015, Paleobiogeography of Late Cretaceous Ammonoids, in Klug, C., Korn, D., De Baets, K., Kruta, I., and Mapes, R.H. eds., Ammonoid Paleobiology: From macroevolution to paleogeography, Dordrecht, Springer Netherlands, v. 44, p. 259–274.

Imlay, R.W., 1937, Lower Neocomian fossils from the Miquihuana region, Mexico: Journal of Paleontology, v. 11, p. 552–574.

Jagt-Yazykova, E.., 2011, Palaeobiogeographical and palaeobiological aspects of mid- and Late Cretaceous ammonite evolution and bio-events in the Russian Pacific: Scripta Geologica, p. 15–121.

Johansen, M.B., and Surlyk, F. Brachiopods and the Stratigraphy of the Upper Campanian and Lower Maastrichtian Chalk of Norfolk, England: Palaeontology, v. 33, p. 823–872.

Johnson, J.G., 1986a, Revision of Lower Devonian (Emsian) Brachiopod Biostratigraphy and Biogeography, Central Nevada: Journal of Paleontology, v.

117

60, p. 825–844.

Johnson, J.G., 1986b, Revision of Lower Devonian (Emsian) brachiopod biostratigraphy and biogeography, central Nevada: Journal of Paleontology, v. 60, p. 825–844, doi: 10.1017/S0022336000042979.

Kaplan, P., and Baumiller, T.K., 2000, Taphonomic Inferences on Boring Habit in the Richmondian Onniella meeki Epibole: Palaios, v. 15, p. 499–510, doi: 10.1669/0883-1351(2000)015<0499:TIOBHI>2.0.CO;2.

Karsten, H., 1886, Géologie de l’ancienne Colombie bolivarienne, Venezuela. Nouvelle- Grenade et Ecuador: Berlin, R. Friedländer & sohn, 62 p.

Kéfi, S., Berlow, E.L., Wieters, E.A., Joppa, L.N., Wood, S.A., Brose, U., and Navarrete, S.A., 2015, Network structure beyond food webs: mapping non-trophic and trophic interactions on Chilean rocky shores: Ecology, v. 96, p. 291–303, doi: 10.1890/13-1424.1.

Kelley, P.H., 1988, Predation by Miocene Gastropods of the Chesapeake Group: Stereotyped and Predictable: Palaios, v. 3, p. 436, doi: 10.2307/3514789.

Kelley, P.H., 2008, Role of bioerosion in taphonomy: effect of predatory drillholes on preservation of mollusc shells, in Wisshak, M. and Tapanila, L. eds., Current Developments in Bioerosion, Berlin, Heidelberg, Springer Berlin Heidelberg, p. 451–470.

Kennedy, W.J., and Cobban, W.A., 1976, Aspects of ammonite biology, biogeography, and biostratigraphy: Special Papers in Palaeontology, p. 1–94.

Kidwell, S.M., Fursich, F.T., and Aigner, T., 1986, Conceptual Framework for the Analysis and Classification of Fossil Concentrations: Palaios, v. 1, p. 228, doi: 10.2307/3514687.

Kitchell, J., 1986, The evolution of predator-prey behavior: naticid gastropods and their molluscan prey, in Nitecki, M. and Kitchell, J. eds., Evolution of Animal Behavior: Paleontological and Field Approaches, Oxford University Press, p. 184.

Kitchell, J., Boggs, C., Kitchell, J., and Rice, J., 1981, Prey Selection by Naticid Gastropods: Experimental Tests and Application to Application to the Fossil Record: Paleobiology, v. 7, p. 533–552.

Klompmaker, A.A., 2012, Drill hole predation on fossil serpulid polychaetes, with new data from the Pliocene of the Netherlands: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 321–322, p. 113–120, doi: 10.1016/j.palaeo.2012.01.024.

Klompmaker, A.A., Portell, R.W., Lad, S.E., and Kowalewski, M., 2015, The fossil record of drilling predation on barnacles: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 426, p. 95–111, doi: 10.1016/j.palaeo.2015.02.035.

118

Klug, C., Korn, D., De Baets, K., Kruta, I., and Mapes, R.H. (Eds.), 2015, Ammonoid Paleobiology: From macroevolution to paleogeography: Dordrecht, Springer Netherlands, Topics in Geobiology, v. 44.

Kolbe, S.E., Lockwood, R., and Hunt, G., 2011, Does morphological variation buffer against extinction? A test using veneroid bivalves from the Plio-Pleistocene of Florida: Paleobiology, v. 37, p. 355–368, doi: 10.1666/09073.1.

Kornicker, L.S., Wise, C.D., and Wise, J.M. Factors affecting the distribution of opposing mollusc values: Journal of Sedimentary Research, v. 33, p. 703–712.

Kowalewski, M., 1990, A hermeneutic analysis of the shell-drilling gastropod predation on mollusks in the Korytnica Clays (Middle Miocene; Holy Cross Mountains, Central Poland): Acta Geologica Polonica, v. 40, p. 183–1213.

Kowalewski, M., 2004, Drill holes produced by the predatory gastropod Nucella lamellosa (muricidae): palaeobiological and ecological implications: Journal Molluscan Studies, v. 70, p. 359–370, doi: 10.1093/mollus/70.4.359.

Kowalewski, M., 2005, Secondary Evolutionary Escalation Between Brachiopods and Enemies of Other Prey: Science, v. 308, p. 1774–1777, doi: 10.1126/science.1113408.

Kowalewski, M., 1996, Taphonomy of a Living Fossil: The Lingulide Brachiopod Glottidia palmeri Dall from Baja California, Mexico: Palaios, v. 11, p. 244–265.

Kowalewski, M., 2002, The fossil record of predation: an overview of analytical methods, in Kowalewski, M. and Kelley, P. eds., The fossil record of predation: New Haven, CT, Paleontological Society Special Papers, v. 8, p. 3–42.

Kowalewski, M., Dulai, A., and Fürsich, F.T., 1998, A fossil record full of holes: The Phanerozoic history of drilling predation: Geology, v. 26, p. 1091, doi: 10.1130/0091-7613(1998)026<1091:AFRFOH>2.3.CO;2.

Kowalewski, M., Dyreson, E., Marcot, J., Vargas, J., Flessa, K., and Hallman, D., 1997, Phenetic discrimination of biometric simpletons; paleobiological implications of morphospecies in the lingulide brachiopod Glottidia: Paleobiology, p. 444–469.

Kowalewski, M., and Flessa, K., 1994, A predatory drillhole in Glottidia palmeri Dall (Brachiopoda; Lingulidae) from Recent tidal flats of northeastern Baja California, Mexico: Journal of Paleontology, v. 68, p. 1403–1405.

Kowalewski, M., and Flessa, K.W., 1996, Improving with age: The fossil record of lingulide brachiopods and the nature of taphonomic megabiases: Geology, v. 24, p. 977, doi: 10.1130/0091-7613(1996)024<0977:IWATFR>2.3.CO;2.

Kowalewski, M., and Flessa, K.W., 2000, Seasonal predation by migratory shorebirds recorded in shells of lingulid brachiopods from Baja California: Mexico. Bulletin of

119

Marine Science, v. 66, p. 405–416.

Kowalewski, M., Flessa, K.W., and Marcot, J., 1997, Predatory scars in the shells of a Recent lingulid brachiopod: paleontological and ecological implications: Acta Palaeontologica Polonica, v. 42, p. 497–532.

Kowalewski, M., and Nebelsick, J.H., 2003, Predation on Recent and Fossil Echinoids, in Kelley, P.H., Kowalewski, M., and Hansen, T.A. eds., Predator—Prey Interactions in the Fossil Record, Boston, MA, Springer US, p. 279–302.

Kowalewski, M., Simões, M.G., Torello, F.F., Mello, L.H.C., and Ghilardi, R.P., 2000, Drill holes in shells of Permian benthic invertebrates: Journal of Paleontology, v. 74, p. 532–543, doi: 10.1017/S0022336000031796.

Krause, R.A., Stempien, J.A., Kowalewski, M., and Miller, A.I., 2007, Body size estimates from the literature: utility and potential for macroevolutionary studies: Palaios, v. 22, p. 60–73, doi: 10.2110/palo.2005.p05-122r.

Lambiotte, R., and Rosvall, M., 2012, Ranking and clustering of nodes in networks with smart teleportation: Physical Review E, v. 85, doi: 10.1103/PhysRevE.85.056107.

Lancichinetti, A., Fortunato, S., and Radicchi, F., 2008, Benchmark graphs for testing community detection algorithms: Physical Review E, v. 78, doi: 10.1103/PhysRevE.78.046110.

Landré, M., 2012, Geoprocessing Journey-to-Work Data: Delineating Commuting Regions in Dalarna, Sweden: ISPRS International Journal of Geo-Information, v. 1, p. 294–314, doi: 10.3390/ijgi1030294.

Lee, D.L., and Smirnova, T.N., 2006, Terebratuloidea, in Treatise on Invertebrate Paleontology, Part H (Brachiopoda Revised), Lawrence, The Geological Society of America and The University of Kansas Press, v. 5, p. 2054–2080.

Lehmann, J., Ifrim, C., Bulot, L., and Frau, C., 2015, Paleobiogeography of Early Cretaceous Ammonoids, in Klug, C., Korn, D., De Baets, K., Kruta, I., and Mapes, R.H. eds., Ammonoid Paleobiology: From macroevolution to paleogeography, Dordrecht, Springer Netherlands, v. 44, p. 229–257.

Leighton, L.R., 2002, Inferring predation intensity in the marine fossil record: Paleobiology, v. 28, p. 328–342.

Leighton, L.R., 2001, New example of Devonian predatory boreholes and the influence of brachiopod spines on predator success: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 165, p. 53–69, doi: 10.1016/S0031-0182(00)00153-X.

Leighton, L.R., 2003, Predation on brachiopods, in Kelley, P.H., Kowalewski, M., and Hansen, T.A. eds., Predator−Prey Interaction in the Fossil Record, New York,

120

Plenum Press, p. 215–237.

Lever, J., Kessler, A., Van Over Beeke, P., and Thijssen, R., 1961, Quantitative beach research II, The “‘hole effect’”: A second mode of sorting of lamellibranch valves on sandy beaches: Netherlands Journal of Sea Research, v. 1, p. 339–358.

Lieberman, B.S., 2005, Geobiology and paleobiogeography: Tracking the coevolution of the Earth and its biota: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 219, p. 23–33, doi: 10.1016/j.palaeo.2004.10.012.

Lieberman, B.S., 2003, Paleobiogeography: The Relevance of Fossils to Biogeography: Annual Review of Ecology, Evolution, and Systematics, v. 34, p. 51–69, doi: 10.1146/annurev.ecolsys.34.121101.153549.

Lieberman, B.S., 2000, The Relevance of Hierarchy Theory to Biogeography and Paleobiogeography, in Paleobiogeography, Boston, MA, Springer US, v. 16, p. 5–27.

Longhurst, A.R. An ecological survey of the West African marine benthos: Colonial Office Fishery Publications, v. 11, p. 1–102.

Lundgren, B., 1883, Undersökningar öfver Brachiopoderna i Sveriges kritsystem: Lunds Universitets. Årsskrift, p. 1–72.

Luque, J., 2015, The oldest higher true crabs (Crustacea: Decapoda: Brachyura): insights from the Early Cretaceous of the Americas (J. Jagt, Ed.): Palaeontology, v. 58, p. 251–263, doi: 10.1111/pala.12135.

Ma, B., Wang, H., Dsouza, M., Lou, J., He, Y., Dai, Z., Brookes, P.C., Xu, J., and Gilbert, J.A., 2016, Geographic patterns of co-occurrence network topological features for soil microbiota at continental scale in eastern China: The ISME Journal, v. 10, p. 1891–1901, doi: 10.1038/ismej.2015.261.

Manceñido, M.O., 2002, Paleobiogeography of Mesozoic brachiopod faunas from Andean–Patagonian areas in a global context: Geobios, v. 35, p. 176–192, doi: 10.1016/S0016-6995(02)00058-X.

Manceñido, M.O., and Owen, E.F., 2007, Post-Paleozoic Rhynchonellida, in Kaesler, R.L. and Sun, D.L. eds., Treatise on invertebrate paleontology. Part H, Brachiopoda revised, Lawrence, Geological Society of America and University of Kansas Press, v. 6, p. 2727–2741.

Marchand, D., Thierry, J., and Tintant, H., 1985, Influence des seuils et des haut-fonds sur la morphologie et l’évolution des Ammonoidés: Bulletin de la Section des Sciences, p. 191–202.

Marcinowski, R., and Wiedmann, J., 1990, The Albian ammonites of Poland: Palaeontologia polonica, p. 3–94.

121

Martin, A.K., and Hartnady, C.J.H., 1986, Plate tectonic development of the south west Indian Ocean: A revised reconstruction of East Antarctica and Africa: Journal of Geophysical Research, v. 91, p. 4767, doi: 10.1029/JB091iB05p04767.

Martinell, J., Kowalewski, M., and Domènech, R., 2012, Drilling Predation on Serpulid Polychaetes (Ditrupa arietina) from the Pliocene of the Cope Basin, Murcia Region, Southeastern Spain (C. Lalueza-Fox, Ed.): PLoS ONE, v. 7, p. e34576, doi: 10.1371/journal.pone.0034576.

Mason, W.T., and Clugston, J.P., 1993, Foods of the Gulf Sturgeon in the Suwannee River, Florida: Transactions of the American Fisheries Society, p. 378–385.

McKerrow, W.S., and Scotese, C.R., 1990, Paleozoic Paleogeography and Biogeography: London, The Geological Society, Geological Society Memoir 12, 453 p.

Melián, C.J., and Bascompte, J., 2004, Food web cohesion: Ecology, v. 85, p. 352–358, doi: 10.1890/02-0638.

Middlemiss, F.A., 1962, Brachiopod ecology and Lower Greensand palaeoecology. Palaeontology, 5, 253-267.: Paleontology, v. 5, p. 253–267.

Middlemiss, F.A., 1981, Brachiopod events in the European Middle Cretaceous (Aptian- Cenomanian): Cretaceous Research, v. 2, p. 377–382, doi: 10.1016/0195- 6671(81)90025-2.

Middlemiss, F.A., 1968a, Brachiopodes du crétacé inférieur des Corbière orientales (Aude): Annales de paléontologie (Invertébrès), v. 54, p. 173–197.

Middlemiss, F.A., 1984a, Cretaceous Terebratulid Events in Western and Southern Europe and their relation to the stage boundaries: Cretaceous Research, v. 5, p. 345–348, doi: 10.1016/S0195-6671(84)80029-4.

Middlemiss, F.A., 1984b, Distribution of Lower Cretaceous brachiopods and its relation to climate, in Brenchley, P. ed., Fossils and Climate, Chichester, John Wiley and Sons Ltd., p. 165–170.

Middlemiss, F.A., 1959, English Aptian Terebratulidae: Palaeontology, v. 2, p. 94–142.

Middlemiss, F.A., 1980, Lower Cretaceous Terebratulidae from South-Western Morocco and their Biogeography. Palaeontology: Palaeontology, v. 23, p. 515–556.

Middlemiss, F.A., 1968b, Observations on the ontogeny of the brachiopod Sellithyris sella: Bulletin Indian Geologists Association, v. 1, p. 1–17.

Middlemiss, F.A., and Brenchley, P. (Eds.), 1984, Distribution of Lower Cretaceous brachiopods and its relation to climate, in Fossils and Climate, Chichester, John Wiley and Sons Ltd., p. 165–170.

122

Miller, R.H., and Sundberg, F.A., 1984, Boring Late Cambrian organisms: Lethaia, v. 17, p. 185–190, doi: 10.1111/j.1502-3931.1984.tb01615.x.

Moalic, Y., Desbruyeres, D., Duarte, C.M., Rozenfeld, A.F., Bachraty, C., and Arnaud- Haond, S., 2012, Biogeography Revisited with Network Theory: Retracing the History of Hydrothermal Vent Communities: Systematic Biology, v. 61, p. 127– 137, doi: 10.1093/sysbio/syr088.

Mora, A., Parra, M., Strecker, M.R., Kammer, A., Dimaté, C., and Rodríguez, F., 2006, Cenozoic contractional reactivation of Mesozoic extensional structures in the Eastern Cordillera of Colombia: Tectonics, v. 25, p. n/a-n/a, doi: 10.1029/2005TC001854.

Morales, L.G., 1958, General Geology and Oil Occurrences of Middle Magdalena Valley, Colombia: South America, in Habitat of Oil, Tulsa, Okla., U.S.A, American Association of Petroleum Geologists, AAPG symposium, p. 641–695.

Mutterlose, J., 1992, Migration and evolution patterns of floras and faunas in marine Early Cretaceous sediments of NW Europe: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 94, p. 261–282, doi: 10.1016/0031- 0182(92)90123-M.

Nebelsick, J.H., and Kowalewski, M., 1999, Drilling Predation on Recent Clypeasteroid Echinoids from the Red Sea: Palaios, v. 14, p. 127, doi: 10.2307/3515369.

Nelson, T.A., and Boots, B., 2008, Detecting spatial hot spots in landscape ecology: Ecography, v. 31, p. 556–566, doi: 10.1111/j.0906-7590.2008.05548.x.

Newman, M.E.J., 2004a, Detecting community structure in networks: The European Physical Journal B - Condensed Matter, v. 38, p. 321–330, doi: 10.1140/epjb/e2004-00124-y.

Newman, M.E.J., 2004b, Fast algorithm for detecting community structure in networks: Physical Review E, v. 69, doi: 10.1103/PhysRevE.69.066133.

Newman, M.E.J., 2006a, Finding community structure in networks using the eigenvectors of matrices: Physical Review E, v. 74, doi: 10.1103/PhysRevE.74.036104.

Newman, M.E.J., 2006b, Modularity and community structure in networks: Proceedings of the National Academy of Sciences, v. 103, p. 8577–8582, doi: 10.1073/pnas.0601602103.

Newman, M.E.J., 2010, Networks: an introduction: New York, Oxford University Press, 772 p.

Newman, M.E.J., and Girvan, M., 2004, Finding and evaluating community structure in networks: Physical Review E, v. 69, doi: 10.1103/PhysRevE.69.026113.

123

Nilsen, E.B., Pedersen, S., and Linnell, J.D.C., 2008, Can minimum convex polygon home ranges be used to draw biologically meaningful conclusions? Ecological Research, v. 23, p. 635–639, doi: 10.1007/s11284-007-0421-9.

Ogg, J.G., Agterberg, F.P., and Gradstein, F.M., 2004, The Cretaceous Period, in Gradstein, F.M., Ogg, J.G., and Smith, A.G. eds., A Geologic Time Scale 2004, New York, Cambridge University Press, p. 344–383.

Olsson, A., 1914, New and interesting Neocene fossils from the Atlantic Coastal Plain: Bulletins of American paleontology, v. 5, p. 1–35.

Onyia, A.D., 1973, A contribution to the food and feeding habit of the threadfin Galeoides decadactylus: Marine Biology, v. 22, p. 371–378, doi: 10.1007/BF00391397. d’Orbigny, A., 1851, Paleontologie Française. Description des Mollusques et Rayonnés Fossiles, Terrains Crétacés. 4, Brachiopodes: 390 p.

Orman, G.K., Labatut, V., and Cherifi, H., 2011, Qualitative Comparison of Community Detection Algorithms, in Cherifi, H., Zain, J.M., and El-Qawasmeh, E. eds., Digital Information and Communication Technology and Its Applications, Berlin, Heidelberg, Springer Berlin Heidelberg, v. 167, p. 265–279.

Owen, H.G., 1973, Ammonite faunal provinces in the Middle and Upper Albian and their palaeogeographical significance, in Casey, R. and Rawson, P.F. eds., The Boreal Lower Cretaceous, Liverpool, Seel House Press, Geological Journal Special Issue 5.

Owen, H.G., 1999, Correlation of Albian European and Tethyan ammonite zonations and the boundaries of the Albian Stage and substages: some comments: v. 3, p. 129–149.

Owen, E.F., 1981, Distribution of some Mesozoic brachiopods in North America, in Gray, J., Boucot, A.J., and Berry, W.B.N. eds., Communities of the Past, Stroudsburg, Hutchinson and Ross, p. 297–309.

Owen, H.G., 1984, The Albian Stage: European province chronology and Ammonite Zonation: Cretaceous Research, v. 5, p. 329–344, doi: 10.1016/S0195- 6671(84)80028-2.

Owen, H.G., and Mutterlose, J., 2006, Late Albian ammonites from offshore Suriname: implications for biostratigraphy and palaeobiogeography: Cretaceous Research, v. 27, p. 717–727, doi: 10.1016/j.cretres.2005.12.001.

Page, K.N., 1996, Mesozoic Ammonoids in Space and Time, in Landman, N.H., Tanabe, K., and Davis, R.A. eds., Ammonoid Paleobiology, Boston, MA, Springer US, v. 13, p. 755–794.

124

Paine, R., 1963, Ecology of the brachiopod Glottidia pyramidata: Ecological Monographs, v. 33, p. 187–213.

Páramo-Fonseca, M.E., Gómez-Pérez, M., Noé, L.F., and Etayo-Serna, F., 2016, Stenorhynchosaurus munozi, gen. et sp. nov. a new pliosaurid from the Upper Barremian (Lower Cretaceous) of Villa de Leiva, Colombia, South America: Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales, v. 40, p. 84, doi: 10.18257/raccefyn.239.

Pasini, G., and Garassino, A., 2012, Naticid gastropod and octopodid cephalopod predatory traces: evidence of drill holes on the leucosid crab Ristoria pliocaenica (Ristori, 1891), from the Pliocene of the “La Serra“ quarry (Tuscany, Italy): Atti della Società italiana di scienze naturali e del Museo civico di storia naturale di Milano, v. 153, p. 257–266.

Patarroyo, P., 2000a, Amonitas del Barremiano en Villa de Leyva-Boyacá (Colombia- Sur America): Datos bioestratigráficos preliminares: Zentralblatt der Geologie und Paläontologie, p. 789–798.

Patarroyo, P., 1997a, Barremiano Inferior en la base de la Formación Paja, Barichara, Santander- Colombia: Geología Colombiana, p. 135–138.

Patarroyo, P., 2000b, Distribución de ammonitas del Barremiano de la Formación Paja en el sector de Villa de Leyva (Boyacá, Colombia). Bioestratigrafía.: Geología Colombiana, p. 149–162.

Patarroyo, P., 2004, Entwicklung der Ammoniten der Familie Pulchelliidae aus dem Barreme von Zentral-Kolumbien Suedamerika: Revue de Paléobiologie, v. 23, p. 1–65.

Patarroyo, P., 1997b, New aspects of the Early Cretaceous Paja Formation in Central Colombia, South America: Zentralblat für Geologie und Paläontologie, p. 547– 558.

Patzkowsky, M.E., 1995, Gradient Analysis of Middle Ordovician Brachiopod Biofacies: Biostratigraphic, Biogeographic, and Macroevolutionary Implications: Palaios, v. 10, p. 154–179, doi: 10.2307/3515180.

Pechenik, J.A., Hsieh, J., Owara, S., Wong, P., Marshall, D., Untersee, S., and Li, W., 2001, Factors selecting for avoidance of drilled shells by the hermit crab Pagurus longicarpus: Journal of Experimental Marine Biology and Ecology, v. 262, p. 75– 89, doi: 10.1016/S0022-0981(01)00284-2.

Peng, Y., Shi, G.R., Gao, Y., He, W., and Shen, S., 2007, How and why did the Lingulidae (Brachiopoda) not only survive the end-Permian mass extinction but also thrive in its aftermath? Palaeogeography, Palaeoclimatology, Palaeoecology, v. 252, p. 118–131, doi: 10.1016/j.palaeo.2006.11.039.

125

Perea, J., García, A., Acero, R., Valerio, D., and Goméz, G., 2008, A photogrammetric methodology for size measurements: Application to the study of weight–shell diameter relationship in juvenile Cantareus aspersus snails: Journal of Molluscan Studies, v. 74, p. 1–5.

Perrin, C., and Kiessling, W., 2012, Latitudinal Trends in Cenozoic Reef Patterns and their Relationship to Climate, in Mutti, M., Piller, W., and Betzler, C. eds., Carbonate Systems during the Oligocene-Miocene Climatic Transition, Oxford, UK, Wiley-Blackwell, p. 17–33, doi: 10.1002/9781118398364.ch2.

Peters, S.E., and McClennen, M., 2016, The Paleobiology Database application programming interface: Paleobiology, v. 42, p. 1–7, doi: 10.1017/pab.2015.39.

Pons, P., and Latapy, M., 2006, Computing Communities in Large Networks Using Random Walks: Journal of Graph Algorithms and Applications, v. 10, p. 191–218.

Poulsen, C.J., Barron, E.J., Arthur, M.A., and Peterson, W.H., 2001, Response of the Mid-Cretaceous global oceanic circulation to tectonic and CO2 forcings: Paleoceanography, v. 16, p. 576–592, doi: 10.1029/2000PA000579.

Poulsen, C.J., Gendaszek, A.S., and Jacob, R.L., 2003, Did the rifting of the Atlantic Ocean cause the Cretaceous thermal maximum? Geology, v. 31, p. 115, doi: 10.1130/0091-7613(2003)031<0115:DTROTA>2.0.CO;2.

Raghavan, U.N., Albert, R., and Kumara, S., 2007a, Near linear time algorithm to detect community structures in large-scale networks: Physical Review E, v. 76, doi: 10.1103/PhysRevE.76.036106.

Reboulet, S., 2001, Limiting factors on shell growth, mode of life and segregation of valanginian ammonoid populations: Evidence from adult-size variations: Geobios, v. 34, p. 423–435, doi: 10.1016/S0016-6995(01)80006-1.

Rhus, N., Kelly, S.R.A., Collins, J.S.H., and Sandy, M.R., 1990, Systematic palaeontology and biostratigraphy of two Early Cretaceous condensed sections from the Barents Sea: Polar Research, v. 8, p. 165–194, doi: 10.1111/j.1751- 8369.1990.tb00383.x.

Riding, J.B., and Crame, J.A., 2002, Aptian to Coniacian (Early–Late Cretaceous) palynostratigraphy of the Gustav Group, James Ross Basin, Antarctica: Cretaceous Research, v. 23, p. 739–760, doi: 10.1006/cres.2002.1024.

Ripley, B.D., 1988, Statistical Inference for Spatial Processes: Cambridge, Cambridge University Press, doi: 10.1017/CBO9780511624131.

Rojas, A., Gracia, A., and Patarroyo, P., 2015, Brachiopods from off the San Bernardo Archipelago (Colombian Caribbean), with comments on specific synonymies in Tichosina, Cooper, 1977: Zootaxa, v. 3914, p. 55, doi: 10.11646/zootaxa.3914.1.3.

126

Rojas, A., Hendy, A., and Dietl, G.P., 2015, Edge-drilling behavior in the predatory gastropod Notocochlis unifasciata (Lamarck, 1822) (Caenogastropoda, Naticidae) from the Pacific coast of Panama: taxonomic and biogeographical implications: Vita Malacologica, v. 13, p. 63–72.

Rojas, A., and Patarroyo, P., 2009, Occurrence of Sellithyris sella (Brachiopoda) in the Rosablanca Formation, Boyacá-Colombia: Boletín de Geología, v. 31, p. 129– 132.

Rojas, A., Patarroyo, P., Mao, L., Bengtson, P., and Kowalewski, M., 2017, Global biogeography of Albian ammonoids: A network-based approach: Geology, v. 45, p. 659–662.

Rojas, A., Portell, R.W., and Kowalewski, M., 2017, The post-Palaeozoic fossil record of drilling predation on lingulide brachiopods: Lethaia, v. 50, p. 296–305, doi: 10.1111/let.12198.

Roopnarine, P., and Beussink, A., 1999, Extinction and naticid predation of the bivalve Chione von mühlfeld in the late Neogene of Florida: Paleontologia Electronica, v. 2, p. 1–14.

Rosvall, M., Axelsson, D., and Bergstrom, C.T., 2009, The map equation: The European Physical Journal Special Topics, v. 178, p. 13–23, doi: 10.1140/epjst/e2010- 01179-1.

Rosvall, M., and Bergstrom, C.T., 2008, Maps of random walks on complex networks reveal community structure: Proceedings of the National Academy of Sciences, v. 105, p. 1118–1123, doi: 10.1073/pnas.0706851105.

Sandy, M.R., 1991a, Aspects of Middle-Late Jurassic-Cretaceous Tethyan brachiopod biogeography in relation to tectonic and paleoceanographic developments: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 87, p. 137–154, doi: 10.1016/0031-0182(91)90133-C.

Sandy, M.R., 1991b, Biogeographic affinities of some Jurassic-Cretaceous brachiopod fauna from the Americas and their relation to tectonic and paleoceanographic event, in MacKinnon, D.I., Lee, D.E., and Campbell, J.D. eds., Brachiopods through time, Rotterdam, Balkema, p. 415–422.

Sandy, M.R., 1990, Early Cretaceous brachiopods from Mexico and their paleobiogeographic significance: Journal of Paleontology, v. 64, p. 942–956.

Sandy, M.R., 1986, Lower Cretaceous brachiopods from Provence, France, and their biostratigraphical distribution: Bulletin of the British Museum (Natural History), v. 40, p. 177–196.

Sandy, M.R., 1997, Mesozoic brachiopods of Mexico – A review and some comments on their paleobiogeographic affinities and paleoecology: Revista Mexicana de

127

Ciencias Geológicas, v. 14, p. 219–224.

Sandy, M.R., Murphy, M.., and Rodda, P.U., 1995, Dzirulina (Brachiopoda; Terebratellidina) from California, U.S.A.–additional record of an Early Cretaceous transatlantic brachiopod genus.: Journal of Paleontology, v. 69, p. 838–845.

Schemm-Gregory, M., Rojas-Briceño, A., Patarroyo, P., and Jaramillo, C., 2012, First report of Hadrosia Cooper, 1983 in South America and its biostratigraphical and palaeobiogeographical implications: Cretaceous Research, v. 34, p. 257–267, doi: 10.1016/j.cretres.2011.11.005.

Scotese, C.R., 2000, PointTracker, Version 7h. Arlington, Tex: Arlington, Texas, http://www.scotese.com.

Scotese, C.R., 2011, The PALEOMAP Project PaleoAtlas vol. 2, Cretaceous paleogeographic and plate tectonic reconstructions: Arlington, Texas, http://www.scotese.com.

Sewall, J.O., van de Wal, R.S.W., van der Zwan, K., van Oosterhout, C., Dijkstra, H.A., and Scotese, C.R., 2007, Climate model boundary conditions for four Cretaceous time slices: Climate of the Past, v. 3, p. 647–657, doi: 10.5194/cp-3-647-2007.

Sidor, C.A., Vilhena, D.A., Angielczyk, K.D., Huttenlocker, A.K., Nesbitt, S.J., Peecook, B.R., Steyer, J.S., Smith, R.M.H., and Tsuji, L.A., 2013, Provincialization of terrestrial faunas following the end-Permian mass extinction: Proceedings of the National Academy of Sciences, v. 110, p. 8129–8133, doi: 10.1073/pnas.1302323110.

Silverman, B.W., 1986, Density estimation: London, Chapman and Hall, Monographs on statistics and applied probability, 175 p.

Simpson, G.G., 1940, Mammals and land bridges: Journal of the Washington Academy of Sciences, v. 30, p. 137–163.

Simpson, M.I., 1985, The stratigraphy of the Atherfield Clay Formation (Lower Aptian, Lower Cretaceous) at the type and other localities in southern England: Proceedings of the Geologists’ Association, v. 96, p. 23–45, doi: 10.1016/S0016- 7878(85)80012-2.

Smirnova, T.N., and Baraboshkin, E.Y., 2004, Valanginian-Lower Hauterivian brachiopod assemblages from the Kacha-Bodrak interfluve, southwestern Crimea: Stratigrafiya, Geologicheskaya korrelaziy, v. 12, p. 152–166.

Smirnova, T., and Ushatinskaya, G., 2001, New lingulids (Brachiopoda) from the Lower Cretaceous of European Russia, with notes in [sic] the microstructure of their shells: Paleontological Journal, p. 387–395.

Smith, A.G., Barry, T., Bown, P., Cope, J., Gale, A., Gibbard, P., Gregory, J., Hounslow,

128

M., Kemp, D., Knox, R., Marshall, J., Oates, M., Rawson, P., Powell, J., et al., 2015, GSSPs, global stratigraphy and correlation: Geological Society, London, Special Publications, v. 404, p. 37–67, doi: 10.1144/SP404.8.

Sobolevsky, S., Szell, M., Campari, R., Couronné, T., Smoreda, Z., and Ratti, C., 2013, Delineating Geographical Regions with Networks of Human Interactions in an Extensive Set of Countries (Y. Moreno, Ed.): PLoS ONE, v. 8, p. e81707, doi: 10.1371/journal.pone.0081707.

Sowerby, J., 1823, The Mineral Conchology of Great Britain. p. 115–160.

Stigall, A.L., and Lieberman, B.S., 2006, Quantitative palaeobiogeography: GIS, phylogenetic biogeographical analysis, and conservation insights: Journal of Biogeography, v. 33, p. 2051–2060, doi: 10.1111/j.1365-2699.2006.01585.x.

Surlyk, F., 1982, Brachiopods of the Campanian-Maastrichtian boundary sequence, Kronsmoor (NW Germany): Geologisches Jahrbuch, v. 61, p. 259–277.

Taylor, J.D., 1980, Diets and habits of shallow water predatory gastropods around Tolo Channel, Hong Kong, in Morton, B. ed., The Malacofauna of Hong Kong and Southern China, Hong Kong, Hong Kong University Press, Proceedings of the First International Workshop on the Malacofauna of Hong Kong and Southern China, p. 163–180.

Vega, F., Nyborg, T., Kovalchuk, G., Etayo-Serna, F., Luque, J., Rojas, A., Patarroyo, P., Porras-Muzquiz, H., Armstrong, A., Bermúdez, H., and Garibay, L., 2010, On some Panamerican Cretaceous crabs (Decapoda: Raninoida): Revista Mexicana de Ciencias Geológicas, v. 62, p. 263–279.

Vermeij, G., 1987, Evolution and Escalation: An Ecological History of Life: New Jersey, Princeton University Press, 527 p.

Vermeij, G.J., 1989, Successful and unsuccessful drilling predation in recent pelecypods: Veliger, v. 32, p. 266–273.

Vermeij, G.J., 1977, The Mesozoic Marine Revolution: Evidence from Snails, Predators and Grazers Paleobiology: Paleobiology, v. 3, p. 245–258.

Vilhena, D.A., and Antonelli, A., 2015, A network approach for identifying and delimiting biogeographical regions: Nature Communications, v. 6, p. 6848, doi: 10.1038/ncomms7848.

Vilhena, D.A., Harris, E.B., Bergstrom, C.T., Maliska, M.E., Ward, P.D., Sidor, C.A., Strömberg, C.A.E., and Wilson, G.P., 2013, Bivalve network reveals latitudinal selectivity gradient at the end-Cretaceous mass extinction: Scientific Reports, v. 3, doi: 10.1038/srep01790.

Villamil, T., 1998, Chronology, relative sea-level history and a new sequence

129

stratigraphic model for basinal Cretaceous facies of Colombia, in Paleogeographic Evolution and Non-Glacial Eustacy, Northern South America, SEPM (Society for Sedimentary Geology), v. 58, p. 161–216, http://sp.sepmonline.org/content/sepspecpub/sepsppag/1.toc (accessed January 2015).

Viviers, M.C., Koutsoukos, E.A.M., da Silva-Telles, A.C., and Bengtson, P., 2000, Stratigraphy and biogeographic affinities of the late Aptian–Campanian ostracods of the Potiguar and Sergipe basins in northeastern Brazil: Cretaceous Research, v. 21, p. 407–455, doi: 10.1006/cres.2000.0205.

Walker, S.E., and Brett, C.E., 2002, Post-Paleozoic patterns in marine predation: was there a Mesozoic and Cenozoic marine predatory revolution, in Kowalewski, M. and Kelley, P.H. eds., The Fossil Record of Predation: Methods, Patterns, and Processes., New Haven, CT, Paleontological Society Special Papers, v. 8, p. 119–194.

Wallace, A.R., 1876, The Geographical Distribution of Animals: New York, Harper & Brothers, 503 p.

Wiedmann, J., 1978, Ammonites from the Curacao Lava Formation, Curacao, Caribbean: Geologie en Mijnbouw, v. 57, p. 361–364.

Williams, A., and Brunton, C.H., 1997, Morphological and anatomical terms applied to brachiopods, in Kaesler, R.L. ed., Treatise on invertebrate paleontology. Part H, Brachiopoda revised, Lawrence, Geological Society of America and University of Kansas Press, v. 1, p. 423–440.

Williams, A., Kaesler, R.L., Moore, R.C., Geological Society of America, and Paleontological Society (Eds.), 2006, Rhynchonelliformea (part): Boulder, The Geological Society of America, Treatise on invertebrate paleontology Brachiopoda, v. 5, 1689 p.

Worcester, W., 1969, On Lingula reevei [Unpublished Master Science Thesis]: University of Hawaii, 49 p.

Wright, C.W., 1963, Cretaceous ammonites from Bathurst Island, Northern Australia: Palaeontology, v. 6, p. 597–614.

Xie, J., Kelley, S., and Szymanski, B.K., 2013, Overlapping community detection in networks: The state-of-the-art and comparative study: ACM Computing Surveys, v. 45, p. 1–35, doi: 10.1145/2501654.2501657.

Xu, B., Liang, Z., Jia, Y., and Zhou, B., 2013, Community Detection Based on Robust Label Propagation Algorithm, in Yuan, Y., Wu, X., and Lu, Y. eds., Trustworthy Computing and Services, Berlin, Heidelberg, Springer Berlin Heidelberg, v. 320, p. 303–310.

130

Zell, P., Beckmann, S., Stinnesbeck, W., and Flores-Ventura, J., 2013, First record of Duvalia ex. gr. lata (Cephalopoda, Coleoidea) from Mexico: Boletin de la Sociedad Geologica Mexicana, v. 65, p. 527–531.

Zhang, Z., Shu, D., Han, J., and Liu, J., 2005, Morpho-anatomical differences of the Early Cambrian Chengjiang and Recent lingulids and their implications: Differences between Early Cambrian and Recent lingulids: Acta Zoologica, v. 86, p. 277–288, doi: 10.1111/j.1463-6395.2005.00211.x.

Zhou, J., Poulsen, C.J., Pollard, D., and White, T.S., 2008, Simulation of modern and middle Cretaceous marine δ 18 O with an ocean-atmosphere general circulation model: Paleoceanography, v. 23, pA3223, doi: 10.1029/2008PA001596.

Złotnik, M., 2001, Size-related changes in predatory behaviour of naticid gastropods from the Middle Miocene Korytnica Clays, Poland: Acta Palaeontologica Polonica, v. 46, p. 87–97.

Złotnik, M., and Ceranka, T., 2005, Patterns of drilling predation of cassid gastropods preying on echinoids from the Middle Miocene of Poland: Acta Palaeontologica Polonica, v. 50, p. 409–428.

Zuschin, M., Stachowitsch, M., and Stanton, R.J., 2003, Patterns and processes of shell fragmentation in modern and ancient marine environments: Earth-Science Reviews, v. 63, p. 33–82, doi: 10.1016/S0012-8252(03)00014-X.

Zuschin, M., and Stanton, R.J., 2001, Experimental Measurement of Shell Strength and Its Taphonomic Interpretation: Palaios, v. 16, p. 161, doi: 10.2307/3515527.

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BIOGRAPHICAL SKETCH

Alexis Rojas received a Bachelor of Geology in the Department of Geosciences at the National University, Bogotá, Colombia in 2009. After graduation, he joined the

Center for Tropical Paleoecology and Archaeology at Smithsonian Tropical Research

Institute (STRI) were he assisted in a number of geological and paleontological projects until 2011. He enrolled in the Department of Geological Sciences at the University of

Florida in 2012 and worked as a teaching assistant in Geological Sciences and research assistant in the Florida Museum of Natural History under the supervision of

Michał Kowalewski. His research interests encompass the post-Paleozoic fossil record of the Neotropical brachiopods and the drilling predation. His current research mainly focuses on adopting a spatially explicit perspective to study the fossil record. He received his Ph.D. from the University of Florida in the summer of 2017.

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