Complexity Trends in the Evolutionary History of Dasycladalean Algae Michael Stearns

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Abstract Dasycladalean algae are a group of marine algae that have an excellent fossil record extending back more than 500 million years. In our study, we investigated morphological complexity over the evolutionary history of this group. To determine complexity for each of more than 400 species, we assigned numerical values for six morphological features; the sum for each species forms a quantitative expression of complexity. We found that although maximum complexity increased over time, forms with minimum complexity persisted during most of the evolutionary history of the group.

Degree Type Open Access Senior Honors Thesis

Department Geography and Geology

First Advisor Dr. Steven T. LoDuca

Keywords Algae, Fossil, Dasycladales, Fossil

Subject Categories Geology

This open access senior honors thesis is available at DigitalCommons@EMU: http://commons.emich.edu/honors/29

Complexity Trends in the Evolutionary History of Dasycladalean Algae

Michael Stearns

Advisor: Dr. Steven T. LoDuca

Honors Thesis

ABSTRACT Dasycladalean algae are a group of marine algae that have an excellent fossil record extending back more than 500 million years. In our study, we investigated morphological complexity over the

evolutionary history of this group. To determine complexity for each

of more than 400 species, we assigned numerical values for six

morphological features; the sum for each species forms a quantitative

expression of complexity. We found that although maximum

complexity increased over time, forms with minimum complexity

persisted during most of the evolutionary history of the group.

TABLE OF CONTENTS

INTRODUCTION………………………………………………………….1-2

MORPHOLOGY AND REPRODUCTIVE MODES.…………………...3-7

THE DASYCLAD FOSSIL RECORD……………………………………7-8

METHODS…………………………………………………………………8-10

RESULTS AND CONCLUSIONS……………………………………….10-12

REFERENCES CITED…………………………………………………..13

APPENDIX I: DASYCLAD COMPLEXITY DATABASE…...…………14-24

INTRODUCTION Dasycladalean algae, or dasyclads, are an Order within the Division

Chloraphyta (“green algae”) in the Kingdom Protista (single-celled eukaryotes)

(Van Den Hoek et. al., 1995). The order is based on the radial symmetry and siphonous body plan of the cell (Fig. 1,2). Dasyclads are epifaunal organisms that must live in the photic zone of warm marine water.

Dasycladalean Algae ("Dasyclads") Kingdom Protista Division Chlorophyta “green algae” Class Ulvophyceae flagellar apparatus: cruciate X-2-X-2 roots, with scales mitosis: closed, persistent spindle Order Dasycladales body plan: siphonous thallus architecture: radial symmetry

Figure 1: Dasyclad classification (from Van Den Hoek et. al., 1995)

Dasyclad algae are an important group for many fields of geology. In certain areas where dasyclads were widespread, there are entire packages of carbonate rocks that consist of their skeletons. Dasyclads are used as index fossils for stratigraphic correlation and as environmental facies indicators by

4 sedimentary geologists. Paleontologists use dasyclads to understand evolutionary trends, and biologists use them for genetic research (Berger Kaever

1992).

a b

Figure 2: Dasyclad thalli showing the radial symmetry characteristic of the Order Dasycladales (from Berger and Kaever, 1992)

Figure 3: Fossil dasyclad casements. The central cavity of the large casement contained the main axis; the perforations in the walls contained the laterals (from Berger and Kaever, 1992) MORPHOLOGY AND REPRODUCTIVE MODES

The dasyclad thallus is composed of a central upright main axis and extensions of the main axis called laterals. Many shapes of dasyclad laterals are known, including phliophore, vesicular, acrophore, trichophore, and pirifer.

(Fig. 4) The important thing to note about the different lateral shapes is that only acrophore can be cylindrical.

Figure 4: Lateral shapes, e-g phliophore, h vesicular, l acrophore, m trichophore, n pirifer (from Berger and Kaever, 1992)

Another aspect of thallus morphology is lateral disposition, which refers to the arrangement of the laterals on the main axis. Disposition has two main variations. The first variation, called euspondyl, refers to the arrangement of laterals in whorls around the main axis. The second variation, aspondyl, refers to a random arrangement of laterals on the main axis. (Fig. 5) A structure related to the laterals is called a cortex. This structure is formed of distally inflated and fused laterals that form an outer “shell”. Some dasyclad species have a cortex; others do not (Fig. 6). Branching of the laterals is another aspect of thallus morphology. Laterals may be undivided or have several orders of branching

(Fig. 7).

a b

Figure 5: Lateral disposition: a. aspondyl b. two variations of euspondyl; note the arrangement of laterals into whorls around the main axis (from Berger and Kaever, 1992)

Figure 6: The structure of the cortex. a. External view of the cortex. Note the fusing of the lateral terminations into a solid outer “shell” b. Cutaway view showing the main axis, laterals, and cortex (from Berger and Kaever, 1992)

Figure 7: Lateral branching a. undivided lateral, b-d. Laterals showing several different orders of branching (from Berger and Kaever, 1992) 8

The presence of a heterocladous thallus is another variation in the morphology of the dasyclad thallus. A heterocladous thallus bears alternating whorls of fertile and non-fertile laterals along the length of the main axis (Fig. 8).

Sterile whorls

Fertile whorls

Figure 8: Heterocladous thallus showing the alternation of fertile and sterile whorls (from Berger and Kaever, 1992) There are three different kinds of reproductive modes that dasyclads

exhibit based on the location of cyst production (Berger and Kaever, 1992) (Fig.

9). The vast majority of dasyclads produce reproductive structures called cysts

or gametangia, which contain and protect the gametes. Endospore cyst

production refers to cysts being produced in the main axis. Cladospore cyst

production refers to the cysts being produced in the laterals. Choristospore cyst

production refers to the cysts being produced in specialized structures called

9 gametophores. Gametophores are outgrowths of the laterals, and may extend from any part of the lateral (Fig. 10).

Figure 9: Three modes of cyst production corresponding to different sites of cyst production in the thallus a. Endospore (in the main axis) b. Cladospore (in the lateral) c. Choristospore (in gametophores)

c a b

Figure 10: Three views of cysts and gametophores a. cysts (small spheres) within gametophores (large clear spheres) b. several gametophores extending from the sides of laterals c. view of thallus with many gametophores (from Berger and Kaever, 1992)

THE DASYCLAD FOSSIL RECORD

Dasycladalean algae have a fossil record that extends from the Cambrian

Period, starting 544 million years ago, to the present (Berger and Kaever, 1992).

Most dasycladalean algae produce a calcareous casement made of aragonite

(CaCO3). This casement covers the main axis and all or some of the laterals

(Fig. 11). The majority of the dasyclad fossil record consists of these casements.

In addition, dasyclads that don’t produce this casement have occasionally been preserved as fossils in anoxic lagoon settings (LoDuca, 1997, 2003).

a b

Figure 11: Carbonate casements a. living dasyclad’s casement that covers the entire thallus b. fossil casement showing where the cysts, main axis, and laterals would have been contained c. diagrammatic views of fossil dasyclad casements (from Berger and Kaever, 1992)

METHODS To investigate patterns of complexity in dasyclads over time, we

constructed a large database. In the database, dasyclads were treated at the

species level and at the temporal resolution of geologic Series. The database

spans from the beginning of the Ordovician, 490 million years ago, to the

present. In total, it incorporates over 400 species, including the 35 living species

found in today’s seas. The complete database is provided in Appendix 1. Figure

12 shows some sample entries from the database.

Quantifying relative complexity for dasyclads, as for other organisms, is

challenging. To quantify complexity for the various morphological character

states and for the different reproductive modes in dasyclads, we assigned numerical values based on inferred relative complexity, such that a higher value was assigned to the character state or reproductive mode regarded as more complex (see Table 1). Overall complexity for a given species is the sum of all

the individual complexity values for that species

We used three guidelines to determine the relative complexity for each

character state and reproductive mode: the order of appearance in the fossil

record, geometric complexity, and information from developmental analysis of living species. The character state that appears first in the fossil record was

considered the least complex. The character state with the least amount of

geometric complexity was considered least complex. Finally, character states

that require fewer chemical signals during development, as determined from

study of living species were considered less complex.

All of the morphological and reproductive information in the database was

compiled from peer reviewed journals and monographs, with most coming from

two larger compilations: Bassoullet et. al., 1978 and Deloffre et. al., 1982. Using

the database we produced the graph shown in Figure 13. This graph shows

dasyclad complexity over time, from the Ordovician Period to the present. Total

complexity for each species appears as a data point on the graph.

Morphology Character State Complexity Value Dispositon Aspondyl 0 Euspondyl 1 Lateral Branching Undivided 0 Branched 1 Cortex No 0 Yes 1 Reproduction Endospore 0 Cladospore 1 Choristospore 2 Lateral Segment Shape Cylindrical 0 Non-cylindrical 1 Heterocladous Thallus No 0 Yes 1

Table 1: System used for quantifying complexity

Figure 12: A portion of the database (For the complete database see appendix 1)

RESULTS AND CONCLUSIONS

Several observations about dasyclad complexity over time emerge for the graph shown in Figure 13. One is that mid-level complexity was established very early in the dasyclad fossil record (Fig. 13). There are two ways to interpret this. 14

First, this could be a real phenomenon, meaning that mid-level complexity

appeared very shortly after the origination of this order. Alternatively, it could be

an indication of a much longer dasyclad history. If prior to the Ordovician Period

dasyclads did not produce casements, then it is likely that they would not have

been preserved as fossils. Therefore, it is possible that dasyclads had a long

evolutionary history before the Ordovician Period that is not preserved in the

fossil record.

A second observation is that overall complexity increased from the

Ordovician Period to present day. Specifically, maximum complexity number

increased from a value of three in the Ordovician Period to seven in the

Cretaceous Period. Minimum complexity also increased. In the Ordovician

Period, minimum complexity was 0; by the Paleogene it was 4. Interestingly,

minimum complexity did not rise as quickly as maximum complexity over

dasyclad evolutionary history (Fig. 13).

A third observation is that during the Triassic and Jurassic Periods,

maximum complexity began to rise dramatically. This corresponds to the arrival

of choristospore reproduction. This observation is especially interesting because

the move from cladospore to choristospore reproduction only increases the

overall complexity value by an increment of one. This suggests that the dramatic

rise in complexity is not only a consequence of the advent of choristospore reproduction.

Finally, the most interesting observation revealed by the dataset is that species with minimum complexity were maintained for over 200 million years

despite the presence of species with higher complexity. In fact, the majority of the dasyclad fossil record from the Ordovician Period to the Neogene Period, is characterized by a minimum complexity of zero or one. Not until late in the

Cretaceous Period did minimum complexity rise above one (Fig. 13). This observation suggests that, in many cases, different morphologies merely corresponded to different ways of attaining equal fitness. In other words, more complex is not necessarily better. Complexity

CO SDPP M mTrJNK P

542 488 443 416 318359 299 251 200 146 65 23 Present

Million years

Figure 13: Graph of dasyclad complexity vs. time. The letters correspond to geologic period names and the numbers correspond to geologic period boundaries

REFERENCES CITED

Bassoullet, J.P., Bernier, P., Conrad, M.A., Deloffre, R., Jaffrezo, M., 1978, Les

Algues Dasycladales du Jurassique et du Cretace: Revision Critique.

Geobios, Lyon, Mem. special 2, 330 p., 2 tab., 40 pl.

Berger, S. and Kaever, M.J. 1992, Dasycladales: An Illustrated Monograph of a

Fascinating Algal Order. Georg Thieme, Stuttgart, 247 p.

Deloffre, R., Genot, P., 1982, Les Algues Dasycladales du Cenozoique: Geobios,

Pau, Mem 4, 247 p., 20pl.

LoDuca, S.T. 2003, The Green Alga Chaetocladus (Dasycladales): Journal of

Paleontology, 71(5), p. 940-949.

LoDuca, S.T. 1997, A New Noncalcified Dasycladalean Alga from the Silurian of

Wisconsin. Journal of Paleontology, 77(6), p. 1152-1158.

Van Den Hoek, C., D.G. Mann, and H.M. Jahns. 1995. Algae: An Introduction to

Phycology. Cambridge University Press, Cambridge, 623 p.

APPENDIX 1: DASYCLAD COMPLEXITY DATABASE w Age Sum Notes d max D max Shape Cortex Lateral d range Species D range Location Arrangement Heterocladus Reproduction Lateral Segment Lateral Branching "Chaetocladus" ottawensis 4 1 1 0 0 1 0 0 2 0.48 "Chaetocladus" ottawensis 5 1 1 0 0 1 0 0 2 0.48 "Chaetocladus" ottawensis 6 1 1 0 0 1 0 0 2 0.48 "Chaetocladus" sardesoni 5 1 1 0 0 1 0 0 2 Acetabularia acetabulum 37 10 1 1 0 2 0 0 4 12 7 55-90 Acetabularia caliculus 37 10 1 1 0 2 0 0 4 6 2 25-35 Acetabularia caliculus 37 5 1 1 0 2 0 0 4 6 2 25-36 Acetabularia caliculus 37 7 1 1 0 2 0 0 4 6 2 25-37 Acetabularia chiavonica 33 4 1 1 0 2 1 0 5 Acetabularia 25- crenulata 37 10 1 1 0 2 0 0 4 15 8 100 Acetabularia 25- crenulata 37 5 1 1 0 2 0 0 4 15 8 101

Acetabularia dentata 37 5 1 1 0 2 0 0 4 4.5 3.5 25-40

Acetabularia dentata 37 7 1 1 0 2 0 0 4 4.5 3.5 25-41

Acetabularia farlowii 37 10 1 1 0 2 0 0 4 4 3 20-30 45- Acetabularia kilneri 37 7 1 1 0 2 0 0 4 14 4 100

Acetabularia major 37 7 1 1 0 2 0 0 4 20 10 60-80 Acetabularia ryukyuensis 37 5 1 1 0 2 0 0 4 10 2 50-60

Acicularia schenckii 37 9 1 1 0 2 0 0 4 7 2 30-50 Acroporella ? assubanipali 29 5 1 1 0 0 0 0 2 0.55 0 1.36 0 12 Acroporella ? Nissorensis 29 4 1 0 0 0 0 0 1 0.22 0.1 0.4 0.04 8

Acroporella anceps 31 4 1 1 0 1 1 0 4 0.51 0.3 0.96 0.46 12-20 Acroporella occidentalis 31 10 1 1 0 1 1 0 4 0.2 1.2 0.73 0.43 8-16

Acroporella radoicici 29 3 1 1 0 0 0 0 2 0.33 0.2 0.7 0.34 8-12 Actinoporella podolica 28 4 1 0 0 1 1 0 3 1.1 0.7 3.2 1.1 13-23 Actinoporella ? Maslovi 29 4 1 0 0 1 1 0 3 0.24 0.1 1.2 0.5 4-9 Actinoporella durandelgai 29 3 1 0 0 1 0 0 2 0.35 0.2 1.45 0.45 7-20 Actinoporella krymensis 28 5 1 0 0 1 1 0 3 Actinoporella podolica 28 4 1 0 0 1 1 0 3 0.85 0.7 3.2 2.5 12-24 Actinoporella podolica 29 4 1 0 0 1 1 0 3 0.85 0.7 3.2 2.5 12-24

Actinoporella sulcata 28 4 1 0 0 1 1 0 3

Actinoporella sulcata 28 4 1 0 0 1 1 0 3 1.1 4 11

Angioporella fouryae 29 3 1 0 1 0 1 0 3 0.75 0.2 4 2.25 30-40 w' 4- Balkhanella hurkai 29 5 1 1 0 1 1 0 4 0.525 0.4 1.18 6? Batophora occidentalis 37 10 1 1 0 2 0 0 4 4 2 Batophora occidentalis var. largoensis 37 10 1 1 0 2 0 0 4 4 2

Batophora oerstedi 37 10 1 1 0 2 0 0 4 13 7 8-16

Belzungia borneti 31 4 0 1 0 1 1 0 3 0.47 0.3 1.5 0.5 8-10

Belzungia silvestril 32 6 1 1 0 1 1 0 4 0.34 0.1 0.78 0.18 7

Belzungia terquemi 31 4 1 1 0 1 1 0 4 0.55 0.2 1.25 0.5 9 62- Calc algal zone Beresella hermineae 17 4 1 0 0 1 0 2 0.222 0.1 0.44 0.12 200 II

Bornetella capitata 37 7 1 1 1 2 1 0 6 10 4 10-14

Bornetella nitida 37 5 1 1 1 2 1 0 6 7.5 4.5 24-30

Bornetella nitida 37 7 1 1 1 2 1 0 6 7.5 4.5 24-30

Bornetella oligospora 37 5 1 1 1 2 1 0 6 6 3 4-12

Bornetella oligospora 37 7 1 1 1 2 1 0 6 6 3 4-12

Bornetella sphaerica 37 7 1 1 1 2 1 0 6 10 4 10-14 20

Broeckella belgica 31 4 1 1 0 1 1 0 4 2.7 1.7 3.01 1.97 8-15

Broeckella belgica 31 5 1 1 0 1 1 0 4 2.7 1.7 3.01 1.97 8-15

Broeckella belgica 31 10 1 1 0 1 1 0 4 2.7 1.7 3.01 1.97 8-15

Broeckella media 32 4 1 1 0 1 1 0 4 0.3 0.66 18 Broeckella ranikotensis 32 5 1 1 0 1 1 0 4 0.83 0.2 2.5 0.5 15-20

Campbelliella striata 29 4 1 0 0 1 1 0 3 1.22 0.9 2.72 2.07 30-80

Carpenterella jonesi 31 4 1 1 0 2 1 0 5 0.35 0.15 7-11

Carpenterella jonesi 32 4 1 1 0 2 1 0 5 0.35 0.15 7-11 Carpenterella morelleti 31 4 1 1 0 2 1 0 5 0.27 0.1 0.93 0.39 11-15 very little info and both in Catellaria ambia 31 5 0 0 1 0 1 0 2 0.03 0 0.3 Russian Chaetocladus capillatus 9 1 1 0 0 1 0 0 2 3 1 30

Chaetocladus dubius 9 1 1 0 0 1 0 0 2 1.5 40

Chaetocladus hefteri 11 1 1 0 0 1 0 0 2 1.5 0.5 10 Chaetocladus plumula 5 1 1 0 0 1 0 0 2 2 40 Chaetocladus ruedemanni 9 1 1 0 0 1 0 0 2 3.5 30

Chalmasia antillana 37 10 1 1 0 2 0 0 4 6 1 26-40 Chlorocladus australicus 37 7 1 1 0 2 0 0 4 4 1 8-12 21

Clavophysoporella 375- endoi 18 4 1 0 0 0 1 0 2 0.254 0.1 0.88 0.28 217 435- Clavoporella reinae 18 4 1 0 0 0 0 0 1 1 0.5 1.87 0.87 220

Clypeina ? solkani 29 3 1 0 0 1 1 0 3 0.31 0.2 1.2 0.68 8-16

Clypeina ? solkani 29 4 1 0 0 1 1 0 3 0.31 0.2 1.2 0.68 8-16

Clypeina caliciformis 28 3 1 0 0 1 1 0 3 0.96 0.4 3.7 1.48 33-38

Clypeina catinula 28 4 1 0 0 1 1 0 3 0.195 0 2.15 0.17 28-30

Clypeina catinula 29 4 1 0 0 1 1 0 3 0.195 0 2.15 0.17 28-30

Clypeina digitata 32 4 1 1 0 1 0 0 3 0.5 0.3 1.4 0.43 7-20 new locality in east indies Clypeina type locality is in Clypeina digitata 32 7 1 1 0 1 0 0 3 0.5 0.3 1.4 0.43 7-20 Paris Basin

Clypeina elliotti 31 4 1 1 0 1 0 0 3 0.25 0.1 0.8 0.12 9-20 france

Clypeina elliotti 32 4 1 1 0 1 0 0 3 0.25 0.1 0.8 0.12 9-20 france

Clypeina elliotti 33 4 1 1 0 1 0 0 3 0.25 0.1 0.8 0.12 9-20 france

Clypeina elliotti 31 10 1 1 0 1 0 0 3 0.25 0.1 0.8 0.12 9-20 cuba

Clypeina elliotti 32 10 1 1 0 1 0 0 3 0.25 0.1 0.8 0.12 9-20 cuba

Clypeina elliotti 33 10 1 1 0 1 0 0 3 0.25 0.1 0.8 0.12 9-20 cuba Clypeina hanabataensis 28 5 1 0 0 1 1 0 3 1.25 4.5 1.5 27 22

Clypeina helvetica 32 4 1 1 0 1 0 0 3 1.6 0.8 2.25 0.45 18-26 Clypeina infundibuliformis 32 4 1 1 0 1 0 0 3 0.49 0.3 1.18 0.28 12-26

Clypeina inopinata 29 4 1 0 0 1 1 0 3 0.4 0.2 1.7 12-19 may be junior synonym of C. Clypeina johnsoni 32 1 1 1 0 1 0 0 3 0.64 0.3 1.56 0.38 16-30 elliotti

Clypeina jurassica 28 3 1 0 0 1 1 0 3 0.59 0.5 3.44 2.92 5-20

Clypeina jurassica 29 3 1 0 0 1 1 0 3 0.59 0.5 3.44 2.92 5-20

Clypeina jurassica 28 4 1 0 0 1 1 0 3 0.59 0.5 3.44 2.92 5-20

Clypeina jurassica 29 4 1 0 0 1 1 0 3 0.59 0.5 3.44 2.92 5-20

Clypeina jurassica 28 5 1 0 0 1 1 0 3 0.59 0.5 3.44 2.92 5-20

Clypeina jurassica 29 5 1 0 0 1 1 0 3 0.59 0.5 3.44 2.92 5-20

Clypeina jurassica 28 6 1 0 0 1 1 0 3 0.59 0.5 3.44 2.92 5-20

Clypeina jurassica 29 6 1 0 0 1 1 0 3 0.59 0.5 3.44 2.92 5-20 Clypeina marginiporella 31 4 1 1 0 1 0 0 3 0.2 0.76 18-20

Clypeina marteli 29 4 1 1 0 1 1 0 4 0.3 0.2 2 1 7-12

Clypeina marteli 29 5 1 1 0 1 1 0 4 0.3 0.2 2 1 7-12

Clypeina marteli 29 6 1 1 0 1 1 0 4 0.3 0.2 2 1 7-12

Clypeina merienda 31 5 1 1 0 1 0 0 3 1 2.5 50 23

Clypeina parvula 28 4 1 0 0 1 1 0 3 0.12 0.1 0.45 0.36 10-25

Clypeina parvula 29 4 1 0 0 1 1 0 3 0.12 0.1 0.45 0.36 10-25

Clypeina rotella 32 5 1 1 0 1 0 0 3 0.45 0.1 1.19 1.12 22

Clypeina sahnii 31 5 1 1 0 1 0 0 3 0.6 0.4 0.9 0.6 20-26

Clypeina socaensis 31 3 1 1 0 1 0 0 3 0.325 0.2 0.63 0.18 10-20

Clypeina stelliformis 32 4 1 1 0 1 0 0 3 0.5 0.3 1 7-16

Clypeina zumetae 28 4 1 0 0 1 1 0 3 0.4 0.1 1.4 0.1 22-24

Clypeina zumetae 29 4 1 0 0 1 1 0 3 0.4 0.1 1.4 0.1 22-24 Coniporella clavaeformis 27 4 0 0 1 0 1 0 2 0.9 0.1 2.6 8-13 Coniporella clavaeformis 27 4 1 0 1 1 1 0 4 Coniporella micromera 27 4 0 0 1 0 1 0 2 0.6 0.3 24 8-20

Coniporella subtilis 28 4 0 0 1 0 1 0 2 .28?

Coniporella subtilis 28 4 1 0 1 1 1 0 4

Connexia fragilis 18 3 1 1 0 1 1 0 4 0.36 1.87 Conodictyum striatum 28 4 0 0 1 0 1 0 2 7 24-30

Crinella carsica 29 3 1 1 1 0 0 0 3 0.93 0.4 2.4 1.22 24-30 Cylindroporella ? Texana 28 2 1 1 0 1 0 1 4 0.04 0 0.29 0.03 6 or 8

Cylindroporella adducta 29 5 1 0 1 1 1 1 5 0.2 0.6 0.1 Cylindroporella arabica 28 5 1 0 1 1 1 1 5 0.23 0.2 0.57 0.26 6

Cylindroporella arsici 29 3 1 1 1 1 1 1 6 0.5 0.1 2 0.3 10 Cylindroporella barnesii 29 2 1 1 1 1 1 1 6 0.14 0.1 0.55 0.17 6 Cylindroporella 6 benizarensis 29 4 1 0 0 1 1 1 4 0.36 0.3 1 0.53 (12?) Cylindroporella elassonos 30 10 1 1 1 1 1 1 6 0.38 0.4 0.09 12 Cylindroporella elitzae 29 3 1 0 1 1 1 1 5 0.4 0.3 1.98 1.45 8 Cylindroporella elliptica 29 3 1 0 1 1 1 1 5 0.22 0 0.66 0.04 Cylindroporella kochanskyae 30 3 1 1 1 1 1 1 6 0.208 0.1 0.64 0.32 4-6 Cylindroporella sugdeni 29 5 1 1 1 1 1 1 6 0.36 0.1 1.14 0.36

Cymopolia ? velici 29 3 1 1 1 2 1 0 6 1.18 0.3 2.4 0.55

Cymopolia barbata 37 10 1 1 0 2 1 1 6 5 3 10-30

Cymopolia barberae 31 5 1 1 0 2 1 0 5 no data

Cymopolia delicata 34 7 1 1 0 2 0 0 4 0.25 0.1 1 0.4 28-32

Cymopolia dollfusi 32 4 1 1 0 2 0 0 4 0.49 0.1 2.35 1.35 14-16 cortx?

Cymopolia edwardsi 32 4 1 1 0 2 0 0 4 0.95 0.5 2.35 1.23 >20 cortx?

Cymopolia elongata 32 4 1 1 1 2 1 0 6 1.89 1.7 2.8 2.3 18-32

Cymopolia incomplete entongensis 31 5 1 1 1 2 0 0 5 0.24 0 0.58 0.01 fossil Cymopolia incomplete entongensis 32 5 1 1 1 2 0 0 5 0.24 0 0.58 0.01 fossil Cymopolia incomplete entongensis 33 5 1 1 1 2 0 0 5 0.24 0 0.58 0.01 fossil Cymopolia eochoristoporica 30 5 1 1 0 2 1 0 5 0.86 0.1 2.2 0.2 12-14

Cymopolia fragilis 32 5 1 1 0 2 0 0 4 0.28 2.8 1.88 12-14

Cymopolia frugifera 31 4 1 1 0 2 1 0 5 0.5 0.1 1.16 0.38

Cymopolia heraki 30 3 1 1 0 2 1 0 5 0.94 0.7 4.5 3.07 20-35 Cymopolia inflataramosa 31 4 1 1 1 2 1 0 6 1.55 1 2 1 20

Cymopolia longistila 29 3 1 1 0 2 1 0 5 0.82 0.3 2.37 0.81

Cymopolia mayaense 32 10 1 1 1 2 1 0 6 0.94 0.7 4.5 3.7 12-35

Cymopolia miocenia 32 3 1 1 1 2 1 0 6 0.6 0.4 1.4 1 Cymopolia morelletorum 30 10 1 1 0 2 1 0 5 0.55 0.1 1.25 0.2 22-24 Cymopolia morelletorum 30 10 1 1 1 2 1 0 6 0.55 0.1 1.25 1.2 22-24

Cymopolia pacifica 32 7 1 1 1 2 1 0 6 0.9 0.6 2.2 0.8 16-40

Cymopolia paktia 32 5 1 1 0 2 1 0 5 1.45 0.9 0.28 0.04 10-15

Cymopolia paronai 31 4 1 1 1 2 1 0 6 0.581 0.2 1.25 0.25 16-18

Cymopolia perkinsi 29 2 1 1 0 2 0 0 4 1.062 0.2 2.42 0.37

Cymopolia rarifistulosa 32 3 1 1 1 2 1 0 6 1 2 Cymopolia rarifistulosa 32 4 1 1 1 2 1 0 6 1 2

Cymopolia tibetica 30 5 1 1 0 2 1 0 5 0.75 0.6 1.5 0.4 10-12

Cymopolia tibetica 30 5 1 1 1 2 1 0 6 0.75 0.2 1.15 0.05 9-12

Cymopolia tingriensis 31 5 1 1 0 2 1 0 5 0.62 0.1 1.24 0.17 26

Cymopolia tingriensis 32 5 1 1 0 2 1 0 5 0.62 0.1 1.24 0.17 26 Cymopolia van bosseae 37 5 1 1 1 2 1 1 7 1 0.5 17-26 Cymopolia van bosseae 37 10 1 1 1 2 1 1 7 1 0.5 17-27

Cymopolia zitteli 32 4 1 1 1 2 1 0 6 0.6 0.3 1.5 0.7 22-34 Dactylopora cylindrica 32 4 1 1 1 2 1 0 6 4 0.5 7 4.9 28 no w or D Dactylopora deloffri 31 3 1 1 1 2 1 0 6 0.515 0.2 1.19 range

Dactylopora gusici 31 3 1 1 1 2 1 0 6 0.75 3 no w Dactylopora "alternating praturloni 32 4 1 1 1 2 1 1 7 2.58 0.1 5.51 0.73 43-50 whorls"

Dasycladus densus 37 7 1 1 0 2 0 0 4 4 2 10-12

Dasycladus ramosus 37 7 1 1 0 2 1 0 5 8 4 13-18 Dasycladus vermicularis 37 10 1 1 0 2 0 0 4 6 3 8-12

Dasyporella sp. 10 0 0 0 0 0 0

Dasyporella sp. 9 0 0 0 0 0 0

Dasyporella sp. 8 0 0 0 0 0 0

Dasyporella sp. 7 0 0 0 0 0 0 0 Digitella branching? No dactyloporoides 32 4 1 0 1 2 1 0 5 0.8 0.5 3 1.4 w

Digitella jacquei 31 4 1 0 1 2 1 0 5 1.1 2.7

Digitella radoicicae 31 3 1 0 1 2 1 0 5 1.5 0.4 2.72 0.45

Digitella rotunda 32 4 1 0 1 2 1 0 5 0.56 0.1 1.5 needs reproduction Dinarella kochi 26 3 1 1 0 1 1 0 4 0.74 0.2 1.85 0.18 12-16 information very little info, just calcareous Diplopora ? sp. 26 4 1 1 0 1 1 0 4 2.8 0.6 3.8 0.6 sleeve

Diplopora annulata 24 4 1 0 0 1 1 0 3 Diplopora annulatissima 23 4 1 0 0 1 1 0 3 1.4 3.1 0.9 Diplopora clavaeformis 23 4 1 0 0 1 1 0 3

Diplopora helvetica 23 4 1 0 0 1 1 0 3 0.5 0 3.1 2 10

Diplopora hexaster 23 4 1 0 0 1 1 0 3 0.9 0.3 2.3 1.4 13

Diplopora johnsoni 29 4 1 1 0 1 1 0 4 1.4 4 0.4 no w

Diplopora nodosa 24 4 1 1 0 0 1 0 3 3.6

Diplopora phanerospora 23 4 1 0 0 1 1 0 3 2.2 1.7 2.9 1.8 6-12

Diplopora philosophi 23 4 1 0 0 1 1 0 3 1.5 1.1 2.2 1.5

Diplopora praecursor 23 4 1 0 0 1 1 0 3 1.7 3.2

Diplopora uniseialis 23 4 1 0 0 1 1 0 3 1.2 0.9 2 1.3 very little info Dissocladella cretica 25 3 1 0 1 1 1 0 4 just skeleton very little info Dissocladella cretica 26 3 1 0 1 1 1 0 4 just skeleton

Dissocladella deserta 31 5 1 1 1 1 1 0 5 0.29 0.1 0.47 0.08 12

Dissocladella deserta 32 5 1 1 1 1 1 0 5 0.29 0.1 0.47 0.08 12

Dissocladella lunata 31 4 1 1 1 1 1 0 5 0.182 0.1 0.42 0.13 4-6 Dissocladella savitriae 30 8 1 1 1 1 1 0 5 1.6 0.5 2.05 0.64 4-6 Dissocladella savitriae 30 8 1 1 1 1 1 0 5 1.6 0.1 2 0.5 40-44 Dissocladella savitriae 31 8 1 1 1 1 1 0 5 1.6 0.1 2 0.5 40-44 Dissocladella undulata 30 6 1 1 1 1 1 0 5 0.1 0 0.32 0.08 8 Dobunniella coriniensis 27 4 1 1 0 1 1 0 4 0.697 0.3 1.7 0.84 6-8

Donezella lunaensis 17 4 1 0 0 0 1 0 2 0.12 0 0.23 0.07

Donezella lutugini 17 4 1 0 0 0 1 0 2 0.045 0 0.6 0.45

Donezella lutugini 18 4 1 0 0 0 1 0 2 0.045 0 0.6 0.45

calc algal zone Dvinella comata 18 4 1 1 0 0 0 2 0.198 0.1 0.32 0.07 50-75 III Eodasycladus ogilviae 26 3 1 1 1 1 1 0 5 2 1.4 3.5 1.8 Eodasycladus ogilviae 26 4 1 1 1 1 1 0 5 2 1.4 3.5 1.8 Eugonophyllum mulderi 18 4 0 0 1 0 1 0 2 Euspondyloporella duplicata 29 3 1 1 0 1 1 0 4 1.8 0.7 6.1 1.7 34-40 Euteutloporella triasina 24 4 1 0 0 0 1 0 2 4.7

Fanesella dolomitica 26 4 1 1 0 1 1 0 4 1 0.6 3.3 0.7

Ferganella asiatica 31 8 1 1 0 2 0 0 4 0.16 0.4 pakistan Furcoporella diplopora 32 4 1 1 1 1 0 4 0.325 0.2 0.6 0.34 12 sporangia? Furcoporella diplopora 32 5 1 1 1 1 0 4 0.325 0.2 0.6 0.34 12 sporangia?

Goniolina geometrica 28 4 1 1 1 1 1 0 5 26.6 11.1

Goniolina geometrica 28 4 1 0 1 1 1 0 4

Goniolina janeti 28 4 1 0 1 1 1 0 4

Goniolina thurmanni 28 4 1 0 1 1 1 0 4

Griphoporella curvata 25 4 0 0 1 0 1 0 2 3.2 2.4 3.6 2.5 38 Griphoporella gumbeli 24 4 0 0 1 0 1 0 2 2.2 0.9 2.9 0.9 28

Griphoporella piae 28 3 0 1 0 0 0 0 1 0.75 0.3 1.5 0.3

Griphoporella piae 29 3 0 1 0 0 0 0 1 0.75 0.3 1.5 0.3 Griphoporella undulata 28 4 0 1 1 0 1 0 3 Gyroporella aff. Vesiculifera 26 4 0 0 0 0 1 0 1 25-35 limited data Gyroporella ampleforata 23 4 0 0 0 1 0 0 1 Gyroporella cf. 332- Nipponica 18 4 1 1 0 0 1 0 3 1.025 0.5 1.75 0.75 250 Gyroporella constricta 18 3 0 0 0 1 1 0 2 0.32 0.85 Gyroporella intraseptata 18 3 0 0 0 1 1 0 2 1.08 1.64

Gyroporella likana 18 3 0 0 0 1 1 0 2 1.02 1.36

Gyroporella maxima 23 4 0 0 0 1 0 0 1 5.4 3.2 6.8 3.3

Gyroporella prisca 18 3 0 0 0 1 1 0 2 2.32 3.1 Gyroporella vesiculifera 25 4 0 0 0 1 0 0 1 2.3 5.1 3.9 30

Halicoryne morellti 34 3 1 1 0 2 1 1 6 0.4 0.2 1.3

Halicoryne spicata 37 5 1 1 0 2 0 0 4

Halicoryne spicata 37 7 1 1 0 2 0 0 4

Halicoryne whrightii 37 5 1 1 0 2 0 0 4 8-12 Hamulusella tribe sedalanensis 31 5 1 1 0 2 1 0 5 0.126 0 1.02 0.81 12 Neomerinae Harlanjohnsonella annulata 30 3 0 0 0 0 1 0 1 1.68 1.2 2.25 0.25 48-50

Helioporella cylindrica 29 3 1 0 0 1 1 1 4 1.85 0.5 2.83 0.61 no w

Herakella paradoxa 18 3 1 1 0 1 1 0 4 0.85 2.3 Heterocladus waukeshaensis 7 1 1 0 0 1 0 1 3 0.5 10 Heteroporella ? poorly Paucicalcarea 29 4 1 0 0 1 1 1 4 0.57 0.3 0.15 0.65 9-18 preserved

Heteroporella anici 28 3 1 0 0 1 1 1 4 0.13 0.1 0.4 0.11 12-16

Heteroporella crosi 26 4 1 1 0 1 1 1 5 0.5 0.1 2.1 0.4 6-7 Heteroporella ellenbergeri 26 4 1 1 0 1 1 1 5 0.84 0.2 1.73 0.33 8-10 Heteroporella Occurs in israelense 29 5 1 1 0 1 1 1 5 1.2 0.5 1.65 0.71 24-25 Israel Heteroporella lemmensis 28 4 1 0 0 1 1 1 4 0.1 0.1 0.68 0.35 7-9 Heteroporella lemoinei 29 3 1 1 0 1 1 1 5 0.31 0.1 0.85 0.15 10-12

Heteroporella lepina 30 4 1 1 0 1 1 1 5 0.4 0.2 0.8 0.4 10-15 Heteroporella lusitanica 28 4 1 1 0 1 1 1 5 0.5 0.3 0.1 0.02 8-12

Indopolia feddeni 31 8 1 1 0 2 0 0 4 0.55 0.55 20-24 pakistan

Indopolia parvula 31 8 1 1 0 2 0 0 4 0.12 0.27 #'s are fertile max infertile Indopolia satyavanti 32 5 1 1 0 2 0 1 5 0.64 0.4 1.16 0.61 28 min #'s are fertile max infertile Indopolia satyavanti 32 8 1 1 0 2 0 1 5 0.64 0.4 1.16 0.61 28 min

Ivdelipora gloriosa 11 5 1 1 0 0 1 0 3

Ivdelipora gloriosa 12 5 1 1 0 0 1 0 3

Ivdelipora gloriosa 13 5 1 1 0 0 1 0 3 Jodotella sloveniaensis 31 3 1 1 0 2 1 0 5 0.275 0.1 0.65 0.25 18

Jodotella thilense 31 4 1 1 0 2 1 0 5 0.171 0.1 0.71 0.22

Jodotella veslensis 31 4 1 1 0 2 1 0 5 0.43 0.2 1.29 0.41 18-24

Jodotella volpensis 31 4 1 1 0 2 1 0 5 0.415 0.2 1.1 0.35 16

Kopetdagaria bifaria 29 3 0 1 1

Likanella ? danilovae 29 3 1 1 0 1 1 1 5 no dimensions

Likanella ? danilovae 29 4 1 1 0 1 1 1 5 no dimensions

Likanella bartheli 28 4 1 0 0 1 1 1 4 0.07 0 0.46 0.26 6 Likanella campanensis 29 4 1 1 0 1 1 1 5 0.5 0.3 1.4 0.5 12 double whorl

Likanella hummudai 30 6 1 0 0 1 1 1 4 1.5 0.94 30-55

Likanella pejovicae 29 3 1 1 0 1 1 1 5 0.72 0.4 3.04 1.84 30-52

Likanella pejovicae 29 4 1 1 0 1 1 1 5 0.72 0.4 3.04 1.84 30-52

Linoporella ? buseri 26 3 1 1 0 1 0 0 3 0.8 0.4 2.4 0.9 12-18

Linoporella ? lucasi 28 4 1 1 1 1 1 0 5 2.1 0.6 3.1 0.7

Linoporella capriotica 30 2 1 1 1 1 1 0 5 1.2 0.4 4 2.5 4-5 mexico

Linoporella capriotica 28 4 1 1 0 1 0 0 3

Linoporella elliotti 29 3 1 1 0 1 1 0 4 0.5 0.1 1.3 0.5 15-20 Linoporella kapelensis 28 3 1 1 1 1 1 0 5 1.4 0.4 2.59 0.37 35-40

Litostroma europaea 18 3 1 1 0 1 1 0 4

Lulipora schattrovii 11 5 1 0 0 0 0 0 1

Lulipora schattrovii 12 5 1 0 0 0 0 0 1

Lulipora schattrovii 13 5 1 0 0 0 0 0 1

Macroporella ? retica 28 4 1 0 1 0 1 0 3 3.1 2 3.9 2.2 60 unnamed Macroporella ? sp. 26 4 1 0 1 0 1 0 3 0.7 species

Macroporella alpina 23 4 0 0 0 0 0 0 0 0.8 Macroporella bellerophontis 23 4 0 0 0 0 0 0 0 picture has no cortex but Berger and Kaever says Macroporella Macroporella beneckei 24 4 0 0 0 0 0 0 0 0.8 2.4 1.1 35 has cortex book says cylindrical but we decided that species with cortex Macroporella dinarica 23 4 0 0 0 0 0 0 0 cannot have 0 34

for lateral shape Macroporella occurs in the embergeri 29 1 0 1 0 1 0 3 0.375 0 0.98 0.08 9-10 Tethyan belt Macroporella espichelensis 28 4 1 0 1 0 1 0 3 0.31 0.1 1.9 0.73 24 295- Macroporella ginkeli 18 4 1 0 0 0 1 0 2 1.16 0.6 1.75 0.38 355 Macroporella helvetica 23 4 0 0 0 0 0 0 0 synonym with Salpingoporella Macroporella incerta 29 3 1 0 1 0 1 0 3 verticillata genus defined Macroporella nisi 29 5 1 0 1 0 1 0 3 0.192 0 1.2 35 by endospore? Macroporella perforatissima 23 4 0 0 0 0 0 0 0 1.5 2.4 0.6 22 Macroporella praturloni 28 3 1 0 1 0 1 0 3 1.6 1 4 2.8 20-40 Macroporella praturloni 29 3 1 0 1 0 1 0 3 1.6 1 4 2.8 20-40 Medusagraptus graminiformis 10 1 0 0 0 0 0 0 0 Medusagraptus miribalas 8 1 0 0 0 0 0 0 0 Medusagraptus miribalas 9 1 0 0 0 0 0 0 0

Medusagraptus sp. 6 5 0 0 0 0 0 0 0 Mellporella 250- anthracoporellaformis 17 4 1 1 0 0 1 0 3 1.98 1.1 2.43 1.3 130 Mellporella 150- beundermani 18 4 1 1 0 0 1 0 3 1.074 0.2 1.35 0.3 90 Microsporangiella langi 31 4 1 1 0 1 1 1 5 0.216 0.1 0.43 0.23 8-10 35

Mizzia velebitana 14 4 1 0 0 1 1 0 3 2.2 1.3 2.6 0.7

Mizzia yabei 14 4 1 0 0 1 1 0 3 0.5 0.2 1 0.5 11

Mizzia yabei 20 5 1 0 0 1 1 0 3

Mizzia yabei 21 5 1 0 0 1 1 0 3

Mizzia yabei 22 5 1 0 0 1 1 0 3 Montenegrella fustiformis 29 3 1 1 1 1 1 0 5 1.3 0.4 2.96 1.01 24-38 Montenegrella tuberifera 29 3 1 1 1 1 1 0 5 1.37 0.4 2.78 0.93 24-30

Montenegrella verae 29 3 1 1 1 1 1 0 5 1.18 0.4 2.59 0.56 24-30

Montiella macropora 31 4 1 1 0 2 0 0 4 0.8 1.3 Berserella's reproduction is Montiella munieri 28 4 1 1 0 2 0 0 4 0.8 1.3 unknown

Morelletina ucrainica 31 3 1 1 0 2 1 0 5 0.45 1.05 not accepted Morelletopora as nammalensis 31 8 1 0 0 2 0 0 3 0.7 2 14-24 Dasycladales

Munieria baconica 29 4 1 0 0 0 0 0 1 1.1 0.5 16 Myrmekioporella mosana 28 1 1 1 1 1 0 5 2.028 0.6 2.6 0.8 28 Neogyroporella elegans 28 5 1 1 1 1 1 0 5 no data Neomeris (Larvaria) auversiensis 32 4 1 1 1 2 1 0 6 0.38 0.2 0.92 0.41

Neomeris (Larvaria) craniphora 31 4 1 1 1 2 1 0 6 0.8 0.4 1.5 0.76 Neomeris (Larvaria) davisi 32 4 1 1 1 2 1 0 6 0.2 0.6 0.3 Neomeris (Larvaria) defrancei 31 4 1 1 1 2 1 0 6 0.5 0.2 0.8 0.26 Neomeris (Larvaria) encrinula 32 4 1 1 1 2 1 0 6 0.15 0.9 0.46 Neomeris (Larvaria) filiformis 32 4 1 1 1 2 1 0 6 0.39 0.3 0.78 0.41 Neomeris (Larvaria) limbata 32 4 1 1 1 2 1 0 6 0.245 0.2 0.78 0.38 Neomeris (Larvaria) montensis 31 4 1 1 1 2 1 0 6 0.3 0.1 1 0.3 Neomeris (Larvaria) reticulata 32 4 1 1 1 2 1 0 6 0.6 0.3 1.05 0.51

Neomeris aciculata 31 3 1 1 1 2 1 0 6 0.15 0 0.4 0.05

Neomeris alternans 33 4 1 1 1 2 1 0 6 0.8 0.4 1.25 0.5

Neomeris ambigua 34 3 1 1 1 2 1 0 6 1 0.5 1.2 0.5

Neomeris annulata 32 4 1 1 1 2 1 0 6 0.7 1

Neomeris annulata 37 10 1 1 1 2 1 0 6 2 0.2 20-60

Neomeris annulata 37 7 1 1 1 2 1 0 6 2 0.2 20-60

Neomeris arenularia 32 5 1 1 1 2 1 0 6 0.78 0.4 1.47 0.81 14-21

Neomeris arenularia 32 4 1 1 1 2 1 0 6 0.78 0.4 1.47 0.81 14-21 Neomeris avellanensis 31 4 1 1 1 2 1 0 6 0.33 0.8 0.2 18-20 37

Neomeris bilimbata 37 7 1 1 1 2 1 0 6 2 1 30-40 Neomeris bowdenensis 35 10 1 1 1 2 1 0 6 1.6 1.5 2.7 0.7

Neomeris budaense 29 2 1 1 0 2 1 0 5 0.45 0.1 0.8 0.01

Neomeris bureaui 32 4 1 1 1 2 1 0 6 0.5 0.2 1.5 0.5 12-18

Neomeris cokeri 37 10 1 1 1 2 1 0 6 3.5 2 12-56

Neomeris courtyi 34 4 1 1 1 2 1 0 6 0.8 1.25

Neomeris cretacea 29 2 1 1 1 2 1 0 6 1.5 0.3 2 0 mexico

Neomeris cretacea 29 3 1 1 1 2 1 0 6 1.5 0.3 2 0 mexico

Neomeris cretacea 29 4 1 1 1 2 1 0 6 1.5 0.3 2 0 mexico

Neomeris cretacea 29 5 1 1 1 2 1 0 6 1.5 0.3 2 0 mexico

Neomeris cretacea 29 6 1 1 1 2 1 0 6 1.5 0.3 2 0 mexico

Neomeris dumetosa 37 10 1 1 1 2 1 0 6 2.6 0.6 24-40

Neomeris dumetosa 37 7 1 1 1 2 1 0 6 2.6 0.6 24-40 Neomeris fercourtensis 32 4 1 1 1 2 1 0 6 0.53 0.2 1.03 0.49

Neomeris fragilis 32 4 1 1 1 2 1 0 6 1.42 0.7 1.96 0.47

Neomeris geometrica 32 4 1 1 1 2 1 0 6 0.2 1 Neomeris herouvalensis 32 4 1 1 1 2 1 0 6 1.592 1 2.16 1.25

Neomeris ignota 34 3 1 1 1 2 1 0 6 3.6 4 0

Neomeris larvariodes 32 4 1 1 1 2 1 0 6 0.51 0.2 1.54 0.51 10-15 Neomeris lusanovkaensis 31 3 1 1 1 2 1 0 6 0.6 1.15

Neomeris mucosa 37 10 1 1 1 2 1 0 6 3 1.5 28-48

Neomeris mucosa 37 7 1 1 1 2 1 0 6 3 1.5 28-48 Neomeris occidentalis 30 10 1 1 0 2 1 1 6 0.425 0.3 0.76 0.36 16-36

Neomeris oligospora 32 4 1 1 1 2 1 0 6 0.25 0.65

Neomeris pfenderae 29 2 1 1 1 2 1 0 6 0.2 0.03

Neomeris plagnensis 31 4 1 1 1 2 1 0 6 0.8 0.5 1.18 0.55 Neomeris pseudo- eruca 32 4 1 1 1 2 1 0 6 1.1 0.4 1.3 0.4

Neomeris pustulosa 32 4 1 1 1 2 1 0 6 1.8

Neomeris radiata 32 4 1 1 1 2 1 0 6 0.5 0.2 1.01 0.33 Neomeris scrobiculata 32 4 1 1 1 2 1 0 6 0.8 0.4 1.37 0.44 11-17

Neomeris stipitata 37 5 1 1 1 2 1 0 6 3 1 16-32 Neomeris van bosseae 37 7 1 1 1 2 1 0 6 4 2 32-44 Neomeris venezuelensis 35 9 1 1 1 2 1 0 6 0.77 0.3 1.36 0.16 cortex with perforated Neomizzia elongata 26 4 1 1 1 1 1 0 5 0.15 0.3 12-16 plates 39

alternating whorls but not Neoteutloporella between fertile gallaeformis 28 3 1 0 0 1 1 0 3 0.85 0.5 2.5 18-23 and infertile Neoteutloporella 80- missing socialis 28 4 1 0 0 1 1 0 3 0.4 0.1 2.3 0.8 100 planches 21

Oligoporella duplicata 23 4 1 0 0 1 0 0 2 1.7 1.3 2.5 1.4 12-25

Oligoporella pilosa 23 4 1 0 0 1 0 0 2

Oligoporella prisca 23 4 1 0 0 1 0 0 2

Oligoporella serripora 23 4 1 0 0 1 0 0 2 Family Orioporella bonieri 31 4 1 1 0 2 1 0 5 Acetabularia

Orioporella briardi 31 4 1 1 0 2 1 0 5

Orioporella malaviae 30 8 1 1 0 2 1 0 5 100

Orioporella villattae 31 4 1 1 0 2 1 0 5

Ortonella myrae 17 4 1 1 0 0 0 0 2 0.05 0 Pagodaporella Family wetzeli 32 5 1 1 0 2 1 0 5 0.22 0 0.34 8 Dasycladacea? Palaeodasycladus barrabei 26 4 1 1 0 1 1 0 4 0.6 2.3 0.7 20-25 Palaeodasycladus elongatulus 26 4 1 1 0 1 1 0 4 2.1 0.7 2.1 0.2 Palaeodasycladus mediterraneus 26 5 1 1 0 1 1 0 4 1 0.8 2.8 2.24 Palaeodasycladus mediterraneus 26 6 1 1 0 1 1 0 4 1 0.8 2.8 2.24

Palaeodasycladus mediterraneus 26 4 1 1 0 1 1 0 4 1.3 0.3 wall thickness Palaepimastoporella of 200-300 espielensis 15 4 0 0 0 1 1 0 2 micrometers Palaepimastoporella espielensis 15 4 0 0 0 0 0 0 0 0.3 0.1

Parkerella binodosa 32 4 1 1 0 2 1 0 5 0.5 0.1 1.1 0.32 Parkerella binodosiformis 31 3 1 1 0 2 1 1 6 0.32 0.1 0.95 0.05

Parkerella montensis 31 4 1 1 0 2 1 0 5 0.5 0.2 1.4 0.58

Petrascula ? Globosa 28 4 1 1 0 1 1 0 4 1.6 0.3 7.5 1.5 40-50

Petrascula ? heraki 26 3 1 1 0 1 1 0 4 0.59 0.2 1.59 0.22 14-16 Petrascula bursiformis 28 4 1 1 0 1 1 0 4 no data Petrascula bursiformis 28 4 1 1 0 1 0 0 3 8.4 7.5 12.8 10 28-29

Petrascula piai 26 3 1 1 0 1 1 0 4 0.55 0.2 3.33 1.48 18-20

Petrascula? Globosa 28 4 1 1 0 1 0 0 3 2.1 0.8 7.5 1.5

Physoporella dissita 23 4 1 0 0 1 0 0 2 Physoporella minutula 24 4 1 0 0 0 0 0 1 2.3 Physoporella pauciforata 23 4 1 0 0 1 0 0 2 Physoporella pauciforata var.? Lotharingica 23 4 1 0 0 1 0 0 2 1 0 2 0.3

Physoporella praealpina 23 4 1 0 0 1 0 0 2 2.4 1.2 3.6 1.1 38

Piania niniyurensis 31 8 1 1 0 2 1 0 5 0.5 1.2 28-30

Polyphysa clavata 37 6 1 1 0 2 0 0 4 2 1 5-13

Polyphysa clavata 37 5 1 1 0 2 0 0 4 2 1 5-14

Polyphysa clavata 37 7 1 1 0 2 0 0 4 2 1 5-15

Polyphysa exigua 37 5 1 1 0 2 0 0 4 3 1 6-14

Polyphysa exigua 37 6 1 1 0 2 0 0 4 3 1 6-15

Polyphysa exigua 37 7 1 1 0 2 0 0 4 3 1 6-16 Polyphysa myriospora 37 9 1 1 0 2 0 0 4 4 1.5 7-9

Polyphysa parvula 37 10 1 1 0 2 0 0 4 3 1 12-19

Polyphysa parvula 37 5 1 1 0 2 0 0 4 3 1 12-20

Polyphysa parvula 37 6 1 1 0 2 0 0 4 3 1 12-21

Polyphysa parvula 37 7 1 1 0 2 0 0 4 3 1 12-22

Polyphysa peniculus 37 7 1 1 0 2 0 0 4 11 7 8-15 Polyphysa polyphysoides 37 10 1 1 0 2 0 0 4 4 2 12-27

Polyphysa pusilla 37 10 1 1 0 2 0 0 4 2.5 1.5 6-17 Praturlonella salernitana 31 4 1 1 0 1 1 0 4 1 0.9 2.2 1.8 8-70

Primicorallina trentonensis 4 4 1 1 0 0 0 0 2 Pseudoactinoporella fragilis 29 4 1 1 1 1 1 0 5 0.59 0.4 2.7 9-22

Pseudoclypeina cirici 28 3 1 1 0 2 1 0 5 1.7 0.1 9 1 Pseudoclypeina crnogorica 29 3 1 1 0 2 1 0 5 4 1.2 20-22 Pseudoclypeina farinacciae 28 3 1 1 0 2 0 0 4 1.24 12-18 Pseudocymoporella anadyomenea 28 5 1 1 0 2 1 0 5 2.73 2 3.9 2.4 45-48 Iraq Pseudocymoporella anadyomenea 29 5 1 1 0 2 1 0 5 2.73 2 3.9 2.4 45-48 Iraq Pseudocymoporella anadyomenea 30 5 1 1 0 2 1 0 5 2.73 2 3.9 2.4 45-48 Iraq Pseudocymoporella jurassica 28 3 1 1 0 2 1 0 5 1.5 0.9 3.6 2.3 48-52 Pseudocymoporella pluricellata 29 3 1 1 0 2 1 0 5 Pseudoepimastopora 190- ? Impera 18 4 1 0 1 0 1 0 3 1.72 0.3 2.11 0.33 200 poorly known Pseudoepimastopora may not be jurassica 28 5 1 0 0 1 0 2 dasyclad

Radoiciciella subtilis 28 3 1 1 1 2 1 0 6 1 0.3 1.4 0.2 8-14

Radoiciciella subtilis 29 3 1 1 1 2 1 0 6 1 0.3 1.4 0.2 8-14 Rostroporella aviformis 32 4 1 1 0 2 0 0 4 1.5 0.7 3.5 2.2 35-40 Salopekiella inopinata 30 3 1 0 0 0 0 0 1 0.75 0.4 1.27 0.64 22-30 Salpingoporella ? atlantica 29 1 1 0 0 1 1 0 3 0.375 0 0.71 0 43

Salpingoporella ? atlantica 29 5 1 0 0 1 1 0 3 0.375 0 0.71 0 Salpingoporella adriatica 29 3 1 0 1 1 1 0 4 0.5 0.2 2.62 1.26 20-22 Salpingoporella annulata 29 4 1 0 0 1 1 0 3 0.25 0.2 0.64 0.34 8-12 Salpingoporella from the middle arabica 29 5 1 0 0 1 1 0 3 0.48 0.3 0.73 0.42 8-10 east

Salpingoporella cemi 29 3 1 0 0 1 1 0 3 0.52 0.8 1.6 0.5 10-15

Salpingoporella cemi 29 4 1 0 0 1 1 0 3 0.52 0.8 1.6 0.5 10-15 Salpingoporella dinarica 29 3 1 0 0 1 1 0 3 0.4 0.3 0.56 0.32 8-10 Salpingoporella genevensis 29 4 1 0 0 1 1 0 3 0.51 0.2 1 0.4 6-9 Salpingoporella muhlbergii 29 4 1 0 0 1 0 0 2 0.2 0.1 0.5 0.2 8

Sariosiella feremollis 31 4 1 1 1 1 1 0 5 0.15 0.1 1 0.66 7-10

Sariosiella feremollis 32 4 1 1 1 1 1 0 5 0.15 0.1 1 0.66 7-10

Sestrophaera liasina 26 4 1 0 1 1 1 0 4 Teutloporella aequalis 24 4 1 0 0 1 0 0 2 Teutloporella gigantea 23 4 1 0 0 1 0 0 2 Teutloporella herculea 24 4 1 0 0 1 0 0 2 6.9

Teutloporella hirsuta 11 5 1 0 0 0 0 0 1

Teutloporella hirsuta 12 5 1 0 0 0 0 0 1

Teutloporella hirsuta 13 5 1 0 0 0 0 0 1

Teutloporella nodosa 24 4 1 0 0 1 0 0 2 1.4 4.6 2 30-40

Teutloporella sp. 23 4 1 0 0 1 0 0 2 1.4

Teutloporella tenuis 23 4 1 0 0 1 0 0 2

Teutloporella triasina 23 4 1 0 0 1 0 0 2 Teutloporella vicentina 24 4 1 0 0 1 0 0 2 3 Thrysoporella cancellata 32 4 1 1 1 1 1 0 5 0.32 0 0.51 0.06 7

Trinocladus atacis 31 4 1 1 0 1 1 0 4 0.23 0 1.6 0.97 10-16

Trinocladus bellus 31 5 1 1 0 1 1 0 4 0.24 0.1 0.55 0.12 12-14

Trinocladus bellus 32 5 1 1 0 1 1 0 4 0.24 0.1 0.55 0.12 12-14 Trinocladus megacladus 31 5 1 1 0 1 1 0 4 0.38 0.2 0.72 0.14 13-14 Trinocladus megacladus 32 5 1 1 0 1 1 0 4 0.38 0.2 0.72 0.14 13-14 Trinocladus perplexus 31 5 1 1 0 1 1 0 4 0.32 0.2 0.62 0.38 6-10 Trinocladus perplexus 32 5 1 1 0 1 1 0 4 0.32 0.2 0.62 0.38 6-10 Trinocladus pinarensis 30 10 1 1 0 1 1 0 4 0.35 2 0.5 12-13

Triploporella remesi 28 4 1 1 0 1 0 0 3 Uglasiella aurigerensis 31 4 1 1 0 1 1 0 4 0.714 0.4 1.26 0.51 12-25

Uglasiella aurigerensis 32 4 1 1 0 1 1 0 4 0.714 0.4 1.26 0.51 12-25

Uncatoella verticillata 11 5 1 1 0 2 0 0 4 Uragiella suprajurassica 28 4 1 0 0 1 1 0 3 Uraloporella sieswerdai 17 4 1 1 0 0 0 0 2 0.49 0.2 0.65 0.22 80-35

Uteria brocchii 31 4 1 1 0 1 1 1 5 1 0.3 2.5 1

Uteria encrinella 32 4 1 1 0 1 1 1 5 0.8 0.5 2.8 1.9 Vermiporella 152- hispanica 18 4 1 0 0 0 0 0 1 0.735 0.2 1.13 0.25 190 Vermiporella velebitana 14 4 0 0 0 0 0 0 0 Zapoporella cantariensis 18 4 1 1 0 0 0 2

Zittelina dumasi 32 4 1 1 1 2 1 0 6 1.25 2.5

Zittelina elegans 32 4 1 1 1 2 1 0 6 5

Zittelina parisiensis 32 4 1 1 1 2 1 0 6

Zittelina simplex 32 4 1 1 1 2 1 0 6 Medusagraptus hispidus 5 5 0 0 0 0 0 0 0 7-9 Medusagraptus setaceus 5 5 0 0 0 0 0 0 0 Medusagraptus pavonicus 5 5 0 0 0 0 0 0 0 Medusagraptus floralis 5 5 0 0 0 0 0 0 0

Medusagraptus divaricuatus 5 5 0 0 0 0 0 0 0 Medusagraptus plumosus 5 5 0 0 0 0 0 0 0

Exigraptus clavulus 4 5 0 0 0 0 0 0 0 Chaetocladus capillatus? 9 5 1 1 0 1 0 0 3 Callithamnopsis delicatula 7 1 1 1 0 0 0 0 2 Callithamnopsis fruticosa 7 1 1 1 0 0 0 0 2 Chaetomorpha? prima 7 1 1 0 0 0 0 0 1 Chaetocladus sardesoni 6 1 1 0 0 0 1 0 2

Articella rotela 4 1 1 0 0 0 0 0 1 0.4 0.7

Articella rotela 5 1 1 0 0 0 0 0 1 0.4 0.7

Articella rotela 6 1 1 0 0 0 0 0 1 0.4 0.7

Novantiella silurica 7 1 1 0 0 0 0 0 1

Novantiella silurica 8 1 1 0 0 0 0 0 1

Novantiella silurica 9 1 1 0 0 0 0 0 1

Intermurella scotica 4 1 1 0 0 0 0 0 1

Intermurella scotica 5 1 1 0 0 0 0 0 1

Intermurella scotica 6 1 1 0 0 0 0 0 1

Inopinatella lawsoni 9 4 1 1 0 0 0 0 2

Inopinatella lawsoni 9 4 1 1 0 0 0 0 2 Medusagraptus mirabilis 9 1 0 0 0 0 0 0 0 Dasyporella norvegica 6 4 0 0 0 0 0 0 0 Novantiella ordiviciana 6 4 1 0 0 0 0 0 1 Archaeobatophora typa 6 1 1 1 0 0 0 0 2

Imperiella iranica 22 5 1 1 0 2 1 0 5 0.686 3.84

Dasyporella silurica 6 5 1 0 0 0 0 0 1 Moniliporella multipora 6 5 1 0 0 0 0 0 1

Plexa varia 6 5 1 0 0 0 0 0 1

Acicularia antiqua 30 5 1 1 0 2 0 0 4

Acicularia elongata 28 4 1 1 0 2 0 0 4 Acicularia (Briardina) sp. 31 5 1 1 0 2 0 0 4 Anthracoporella spectabilis 22 5 0 1 0 0 0 0 1 Anthracoporella spectabilis 21 5 0 1 0 0 0 0 1 Anthracoporella spectabilis 20 5 0 1 0 0 0 0 1 Anthracoporella mercurii 22 5 0 1 0 0 0 0 1

Anthracoporella mercurii 21 5 0 1 0 0 0 0 1 Anthracoporella mercurii 20 5 0 1 0 0 0 0 1 Atractyliopsis darariensis 22 5 0 0 0 1 0 0 1

Clypeina lucasi 29 5 1 0 0 2 1 0 4 Cymopolia anadyomenea 30 5 1 1 0 2 1 0 5 Cymopolia kurdistanensis 31 5 1 1 0 2 1 0 5 Griphoporella cf. perforatissima 29 5 0 0 1 1 1 0 3 Griphoporella cf. perforatissima 28 5 0 0 1 1 1 0 3 "Griphoporella arabica" 32 5 0 0 1 1 1 0 3 "Griphoporella arabica" 31 5 0 0 1 1 1 0 3 Permoperplexella attenuata 22 5 0 0 0 0 0 0 0 Permoperplexella attenuata 21 5 0 0 0 0 0 0 0 Permoperplexella attenuata 20 5 0 0 0 0 0 0 0 Pseudoepimastopora ampullacea 22 5 1 1 0 1 1 0 4 Pseudoepimastopora ampullacea 21 5 1 1 0 1 1 0 4 Pseudoepimastopora ampullacea 20 5 1 1 0 1 1 0 4 Pseudoepimastopora cf. likana 22 5 1 1 0 1 1 0 4

Pseudoepimastopora cf. likana 21 5 1 1 0 1 1 0 4 Pseudoepimastopora cf. likana 20 5 1 1 0 1 1 0 4 Salpingoporella apenninica 28 5 1 0 0 0 0 0 1 Trinocladus tripolanus 30 5 1 1 0 1 1 0 4 Trinocladus radoicicae 30 5 1 1 0 1 1 0 4

Apidium sp. 6 4 1 1 1 0 1 0 4 Atractyliopsis cumberlandensis 14 4 0 0 0 1 0 0 1

Atractyliopsis minima 15 4 0 0 0 1 0 0 1

Dutroella scotti 15 4 1 0 1 1 1 0 4

Cyclocrinitides sp. 4 1 0 1 0 1 0 3

Cyclocrinitides sp. 5 1 0 1 0 1 0 3

Cyclocrinitides sp. 6 1 0 1 0 1 0 3

Cyclocrinitides sp. 7 1 0 1 0 1 0 3

Cyclocrinitides sp. 8 1 0 1 0 1 0 3

Cyclocrinitides sp. 9 1 0 1 0 1 0 3

Cyclocrinitides sp. 10 1 0 1 0 1 0 3

Eovelebitella robertsi 13 4 1 0 0 0 1 0 2

Eovelebitella robertsi 14 4 1 0 0 0 1 0 2

Eovelebitella robertsi 15 4 1 0 0 0 1 0 2 Pekiskopora macqueeni 14 2 1 0 0 0 0 0 1 Pekiskopora macqueeni 15 2 1 0 0 0 0 0 1 Pekiskopora macqueeni 16 2 1 0 0 0 0 0 1 Pekiskopora macqueeni 17 2 1 0 0 0 0 0 1 Pekiskopora macqueeni 18 2 1 0 0 0 0 0 1 Pekiskopora macqueeni 19 2 1 0 0 0 0 0 1 Columbiapora johnsoni 14 2 1 0 0 0 0 0 1 Columbiapora johnsoni 15 2 1 0 0 0 0 0 1 Columbiapora johnsoni 16 2 1 0 0 0 0 0 1 Columbiapora johnsoni 17 2 1 0 0 0 0 0 1 Columbiapora johnsoni 18 2 1 0 0 0 0 0 1 Columbiapora johnsoni 19 2 1 0 0 0 0 0 1 Planches II is Nanopora anglica 15 4 1 0 0 1 1 0 3 missing Pseudonanopora stockmansi 15 4 1 0 0 1 1 0 3 Anatolipora carbonica 15 4 1 0 0 1 1 0 3

Cabrieropora pokornyi 15 4 1 0 0 0 1 0 2 Eovelebitella occitanica 15 4 1 0 0 0 1 0 2 Nanopora fragilissima 14 1 1 0 0 1 1 0 3 Nanopora fragilissima 15 1 1 0 0 1 1 0 3 Nanopora fragilissima 16 1 1 0 0 1 1 0 3

Windsopora radiata 14 1 1 0 0 0 0 0 1

Windsopora radiata 15 1 1 0 0 0 0 0 1

Windsopora radiata 16 1 1 0 0 0 0 0 1 Sphinctoporella lisburnensis 14 1 0 0 0 0 0 0 0 Sphinctoporella lisburnensis 15 1 0 0 0 0 0 0 0 Sphinctoporella lisburnensis 16 1 0 0 0 0 0 0 0