IBRARIES \llllll ; llllllllll\l\\l\l\l\l\\ This is to certify that the thesis entitled Phylogenetic Analysis of the Rhabdomesine Bryozoans presented by Kurt D. Spearing has been accepted towards fulfillment of the requirements for MEL—degree in Jeological Sciences /W Major professor Date fl‘j’zo/ [271‘ 0—7 639 MS U is an Affirmative Action/Equal Opportunity Institution LIBRARY Michigan State Unlverslty PLACE IN RETURN BOX to remove this checkout from your record. TO AVOID FINES return on or before date due. MTE DUE DATE DUE DATE DUE 1/98 WM“ PHYLOGENETIC ANALYSIS OF THE RHABDOMESINE BRYOZOAN S by Kurt D. Spean'ng A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Department of Geological Sciences 1998 ABSTRACT PHYLOGENETIC ANALYSIS OF THE RHABDOMESINE BRYOZOANS By Kurt D. Spearing The Bryozoan suborder Rhabdomesina is a diverse group of Paleozoic invertebrates that have a well understood morphology, and are well represented in most stratigraphic divisions from the Arenigan to the Dzulfian. There have been several prior attempts to uncover the evolutionary history of the group, but none have been conclusive. This phylogenetic study used groupings of rhabdomesine genera from several sources as a basis for the list of taxa, and the character lists were adapted mostly from the muse on Invertebrate Paleontology volume G (revised). This information was used to create a data matrix which then was entered into the PAUP and MacClade phylogenetic software packages. There were several runs of the data, differing by various removal of characters or taxa in attempts to lessen the effect of missing data. The data was also examined both with and without use of a stratigraphic character. This study found that there are several groups of genera that cluster together consistently, but there are still many areas of uncertainly. There is a general two clade grouping that separates most arthrostylids from most other rhabdomesines. This is consistent with previous studies that divided the group into a rhabdomesine clade and put the arthrostylids into another clade grouped with the fenestrates, and shows a possible polyphyletic origin for the suborder. ACKNOWLEDGMENTS There are several people that should be thanked for their help in the completion of this thesis. Dr. Anstey, for being my advisor. Drs. Taggart and Brandt for being on my committee, and Dr. Gottfried for sitting in when I needed him to. And last but not least, I Should thank Brian and Christina Walter for letting me use their computer over and over for hours (sometimes days) at a time. LIST OF TABLES LIST OF FIGURES INTRODUCTION METHODS RESULTS DISCUSSION FUTURE WORK APPENDIX LIST OF REFERENCES TABLE OF CONTENTS 10 40 45 47 58 LIST OF TABLES TABLE 1: STRATIGRAPHIC RANGES OF THE GENERA IN THIS STUDY TABLE 2: DATA MATRD( TABLE 3: TAXA OR CHARACTERS REMOVED DURING DIFFERENT EXECUTIONS OF THE SOFTWARE TABLE 4: TREE STATISTICS TABLE 5: FAMILIAL GROUPINGS USED BY OTHER AUTHORS 12 41 FIGURE 1 FIGURE 2 FIGURE 3 FIGURE 4 FIGURE 5 FIGURE 6 FIGURE 7 FIGURE 8 FIGURE 9 FIGURE 10 FIGURE 11 FIGURE 12 FIGURE 13 LIST OF FIGURES vi 13 15 I8 20 22 24 27 29 32 34 36 38 55 INTRODUCTION The rhabdomesine suborder of bryozoans is a diverse group of generally slender, cylindrical, dendroid bryozoans that are found in Paleozoic marine sediments worldwide. Their morphology is well understood, and the size of the suborder (approximately 67 genera) made them a workable choice for phylogenetic analysis. They range in age from the Tremadocian (early Ordovician) to the Dzulfian (late Permian). This long time interval makes possible the inclusion of a large number of stratigraphic divisions for data entry in a cladistic study. Stratigraphic data have not been utilized in other cladistic studies specifically directed at rhabdomesines, and only one cladistic study (Blake and Snyder, 1987) has tried to deal with the rhabdomesine suborder on the generic level. Previous attempts to resolve the phylogeny of the rhabdomesines have not been conclusive. Blake and Snyder (1987) could not resolve the evolutionary history of the rhabdomesines using cladistic methods, although they claimed some success with phenetic techniques. They only were able to get satisfactory cladistic resolution when they ran the data in small blocks (14.21 genera). When the whole suborder was included in the analysis they claimed some broad consistencies between the many trees generated, but the overall results were inconclusive, with many equally parsimonious trees generated. Based upon analysis of family level taxa, the suborder Rhabdomesina was found by Anstey and Pachut (1995) to be either (1) a paraphyletic sister group to the fenestrates (their fig. 8.2), or (2) a partly monophyletic (four families) and partly polyphyletic group (5 families belonging to the fenestrates) (their figs. 8.2, 8.8). The Order Cryptostomata, which conventionally includes the rhabdomesines, fenestrates, and ptilodictyines (Blake, 1983b), has been questioned as a clade; earlier results indicate that it might be monophyletic (Anstey and Pachut, 1995, fig. 8.2 and 8.8), polyphyletic (Anstey and Pachut, 1995, fig. 8.3), or paraphyletic (Anstey 1990). Systematists have recognized the affinities of the suborder in diverse ways: some hypotheses have considered the rhabdomesines to be a sister group to the ptilodictyines (Cuffey, 1973; Cuffey and Blake, 1991), whereas Anstey and Pachut (1995) found them to be, in part, a sister group to the fenestrates, and in part a member of that group. There are morphological convergences between some rhabdomesines and some trepostomes (Blake, 1980; Schulga-Nesterenko et al., 1972), particularly with the trepostome Suborder Amplexoporina, and Dzik (1992; 1994) even claimed that the rhabdomesines may be trepostomes. Other cladistic results indicate the possible transfer of the families Arthrostylidae and Hyphasmoporidae from the rhabdomesines to the fenestrates (Anstey and Pachut, 1995). Anstey and Pachut (1995) also showed that when the rhabdomesine families Rhabdomesidae and Bactroporidae were added into an analysis where they had previously been absent, the cryptostome clade was “destroyed” due to the shifting of convergent character states from terminal to basal branches, the net result being the disjunction of the rhabdomesines and the ptilodictyines. The rhabdomesines are a diverse suborder in need of further phylogenetic analysis to address a variety of hypotheses concerning their interrelationships and evolutionary history. Other than paleontologists, few systematists have considered stratigraphic data useful in evaluating phylogeny (Fisher, 1994). The rhabdomesines should benefit from the inclusion of geochronological data in a cladistic analysis, which would be an important ordering tool in comparing alternative phylogenies, and could provide an independent test of the branching order within cladograms (Huelsenbeck, 1994; Fisher, 1994). A cladistic study has the potential to resolve issues concerning the systematics, patterns of descent, rates of evolution, and macroevolutionary patterns displayed by this group. Some of these issues include: the validity of the current familial structure (taxonomy of the group) (Blake, 1983a); questions concerning ancestry and descent, and the effects of mass extinction, radiation, convergence, parallelism, evolutionary reversals, and heterochrony. Answers to many of these questions may present themselves, but the main purpose here will be on a refined tree definition and taxonomic conclusions based on the best supported trees that are generated. This choice was made because once clearer resolution is achieved, the tree structure of the group will be the major key in answering other questions. METHODS To examine the phylogeny of the rhabdomesines two primary computer software packages were used The Phylogenetic Analysis Using Parsimony (PAUP) version 3.1.1 (Swofford, 1993) computer software was the main tool in this study, along with MacCLADE 3.0 (Maddison, and Maddison, 1992), another cladistics program, which is very useful for close examination and study of the trees that PAUP generates. In this study I have used published descriptions and photographs to determine a list of characters displaying derived states in my operational taxonomic units (OTU’s); these states are either nominal (0,1) or ordinal (0,1,2,3, etc.) in nature, based upon qualitative differences or multistate morphoclines. Character states were then polarized [evolutionary direction of a character transformation series from ancestral to derived (Mayr 1991)] by stratigraphic position. Primitive states, polarized by the oldest occurring state, were coded as 0 in the characters that can be clearly polarized and ordered; in cases lacking clear-cut criteria I simply left the characters in an unpolarized and unordered state (see Appendix 1). The main factor in the decisions concerning polarimtion and ordering was a consensus based upon the character states of the genera of the first two stages of the Ordovician (Tremadocian and Arenigian). Characters that did not seem to have clearly defined transitions between the older and younger taxa were treated as unordered characters, which means their states were coded arbitrarily and the software allowed for any state to change to any other state (example 1-2, 2-3, 3-1), as opposed to ordered states where the changes had to occur in a specific order (example 1- 2-3). The geochronological character that I used was listed as an irreversible character, which is not only an ordered character, but one in which changes can only happen in one direction (for example 1-2, 2-3, but not 2-1, 3-2). This was done because that was the only way to introduce geological time in PAUP 3.1.1. The MacClade program has a stratigraphic character, but since most of the processing was done in PAUP, the irreversible method was more practical. To determine the hue of the tree I decided to root it by either using the oldest known genus that is recognized as part of the suborder, namely Arthroclema, or by using all of the taxa that are in the two oldest stratigraphic units (up to seven genera are known from the Tremadocian and Arenigian), the final analyses used the second method, stratigraphic ranges of the genera in this study are listed on Table 1. The OTU’s in my study are the genera of the suborder. To decide which genera to include in the rhabdomesines I used lists of genera from several sources including: the Suborder Rhabdomesina in the Treatise on Invertebrate Paleontology (vol. G revised) (Blake, 1983), the group as defined by Dzik (1992, 1994), and the genera listed on the stratigraphic charts in Ross (1996). Where possible, I used the type species description and photographs to determine character states; however some genera had only partial descriptions so other sources had to be used Some genera also were left out of some of the final studies due to problems in obtaining complete character state data; the most common reason for exclusion was that only silicified remains were known, which usually resulted in the preservation of only external character states (see results). From the various lists I included as many genera as possible from all sources, ending up with as many as 67 OTU’S, which is a much higher number than used by any previous author (between 39 and 58 depending upon taxonomic treatment). Once the characters were coded (see Table 2 and Appendix 1), the PAUP and MacClade programs were executed. Final runs of the PAUP 3.1.1 program included several variations based upon the exclusion of characters and taxa with respect to the missing data (see Table 3). These include: the full matrix (Table 2), the matrix minus the characters with missing data, the matrixminusthetaxawithmissingdata,andthematrixminusthetaxawithmorethan TABLE 1: STRATIGRAPHIC RANGES OF THE GENERA IN THIS STUDY 51.22 <._. Nc-atoporo(9) Neuatoporolla<50> thhrotrupa<51> U1r1chostulus<14> Scoptopora<12) Cunaatopora<3> Holopora<6> Holoclo-a(3) Psoudono-atopora(11) Hoxttcs<7> Huphasnoporo<33) Ogbtnoporatac) Spira(54) Europoracsz) Panirolla<25> P1nogopora<31> Hageholla<38) 8troblocladt¢<40> LInotoxIs<48> H9010toochus<53> Tropidoporo<13) Hongloclona(l7)' Hediapora(17) Rhonbopora<22> Klaucona<23) Fri-oralla(26) Scffordotaxis(27) Holiotrgpa(43) Clausotrupat49) flogaccnthopora<24> Callocladia<39) Cooletubulipora<42> NIktforouellaCZO) PotaloporollaC36) Stroblotrgpclla(32) Reanthoclcna<30) Bhobdoncson<15> flscopora(16) Nickclsopora(19) Stroblotrgpa<35> Stroblcscopora(37) Euthgronbopora(41) Ho.ctaxts<18) Trouatolla<21) OrthoporaCZO) flounrclla<8> BactroporaCZO) Neuatotrupa<45> Enallopora<55) Osburnostglus<10> Glaucononolla(4) Pasaastglus<59> Ooldfussitrgpa<44> 0ttosootaxis<46> Bluunoporo(56) Psoudohornora<57> flrthroclono(2) 15 Figure 2a Arthrostylus Arthroctema Cuneatopora Glauconomella Hebeiema Helopora Hexites Moyerelta Nematopora Osburnostylus Pseudonernatopora Sceptopora Tropidopora Ulrichostytus Rhabdomen Ascopora Mediapora Nematards Nickelsopora Orthopora Trematelia Rhombopora Klaucena Megacanthopora Pamirelia Primoreila Satiordotaxis Bactropora Nikiiorovella Acanthoclema Pinegopora Streblotrypeiia Hyphasmopora Ogbinopora Streblotrypa Petaloporella Strebiascopora Maychella Calloctadia Streblocladla Euthyrombopora Coelotubulipora Heliotrypa Goldtussitrypa Nematotrypa Ottoseetaxis Monglociema Linotaxis Clausotrypa Nematoporolla Arthrotrypa Europora Hyalotoechus Spira Enallopora Alwynopora Pseudohornera Pesnastytus \. I __.__ «- ._.__.__ a. __ .. _____._._ I O 4'___—_.0)________ /--7 ————_.Q.________—.—\. I O / _____ o_____\ Inn 1 /-84 I | I /--83 | | \ \ O ID ,r as \ --------- as I \ \ \'-86 \ ’—--- roa---- /--92 I I | \ ........... I-92 ’--_- roo---- ’---- firthrostulus(1) Neldtopora<9> Ulrichostulus<14> thhroclo-a(2) -(12) Holoe1;-a(5) Cunoatopora(3) Hoycrolla(8) 0sburnostg1us<16> Olaucononclla(4) Pusnastulus<58) Psoudohornora<57) Hoxitos<7> Neantoporolla<56) Hrthrotrupu<51> Stroblocladiat40) Rluunopora(56) Psoudononatopora(11) Tropidopora<13> Ooldfussitrgpa<44> Hodiapora<17) "ouch-110(38) HonglocIa-a(d7) Callocladia(39) Holintrupa<43) Conletubulipora<¢2> Rho-bopora(22) No-atotrgpa(15) Clausotrupa<49) Ennllopora<55) Hogacanthopora<24> Klaucanatza PrincrollaCZS) Saffordotaxis<27) Buctropora<28> Ottosootaxist46) Ntktforovo11a(29) Petaloporollu<36) Pincgopcrat31) Stroblotrgpolla<32) Holopora(6) Spira<54> Palirclla<25> Rhabdo-oson(15> Huph 09binopora<34> Stroblotrupa<35) Stroblascopora<37) Nick-lsopora(19) No-atax13(18> 0rthopora<26> I-BB---- Tronatolla<21> /--77 El U0 /_-_- H9¢lotocchus<53> Reanthoclona<30> \ ----- 83---- Linotaxis<48> Figure 2b 17 There are other polytomies which indicate areas that cannot be resolved. There are also very few nodes that are supported in more than 95% of the trees. Trees three and four were run with the full character set, but eleven genera with missing data were removed (see Table 3). Tree three also has the near basal polytomy one node up from the base; however it is reduced to 7 branches instead of the 11 that were found in figure 1a, and still shows uncertainty about the monophyly of the Arthrostylidae as a basal family. There is again another polytomy of 14 branches at one of the internal nodes; however it is not nearly as extreme as the 27 branches in figure 1a. The tree overall shows much more definition, and familial groupings are much more easily deduced (see conclusions). Tree three’s majority rule data (figure 3b) shows the overall strength of the tree, but also indicates why there are some polytomies in the strict consensus. There are two regions in the tree where the nodes have very low percentages; these correspond with the two large polytomies in the consensus tree. Tree four (figure 4a) has the same problem as all of the trees with no geochronological character, i.e. no structure is visible in the strict consensus tree. In addition very few of the nodes in the majority rule tree (figure 4b) have the high percentages that were present in tree 3, and the basal branches have percentages below 80%. Trees five and six were run with full sets of taxa, but all nineteen characters with missing data were removed (see Table 3). In tree five (figure 5a) the basal polytomy that has occurred in all of the geochronological trees is present once again, but it now includes 20 branches. In this tree the internal polytomy of 33 branches is the dominant l8 Figure 3a Arthrostylus Cuneatopora Helocierna Pseudonematopora Hexites Hyphasrnopora Ogbinopora Spire Tropidopora Trematella Rhabdomeson . Aseopora Medlapora Rhombopora Klaucena Primorella Saiiordotaxis Clausotrypa Megacanthopora Nemataxis Nickeisopora Pamirela Nikitorovella Petaloporeila Acanthoclerna Pinegopora Streblotrypelia Streblotrypa Streblascopora Unotaxis Hyalotoechus Orthopora Moyerella HeIOpora Bectropora Ottoseetaxis Glauconomella Pesnastylus Nematoporella Arthrotrypa Osburnostylus Nematopora Sceptopora Ulrichostylus Alwynopora Pseudohomera Arthroclema \ I I I I I I I I I I I I 8__-_____-_____\ 8—————————_—\ p 8_-_————_H an. ’-———-————_—-- ’--—-——— 19 roe I-- ran \ ------- O ’_--—-—-‘-_—_——__— q——-——---—-__——_- 0 I ’___fl___\ ’——————-——— Figure 3b thhrostglus<1> Nonatopora(9) Ulrichostglus<14> Cunoatopora(3) Holopora(6) Holoelola(5) Psoudonouatopora<11> Hoxttos<7> H9phasuopora<33> Ogbtnopora(34) Spira<44> Strcblotrupa(35) Stroblascoporot37) Pantrolla<25> PInogopora<3I> L1notaxis<39) Hyolotoochus<43) Hodtapora<17) MonboporaQZ) . K10ucona<23> Princrolla<26> SaffordotaxtstZ?) Clausotrgpatdc) flogaeanthoporatZI) NIKIforovollo<29> Pataloporella<36> flcanthoclc-atao) Streblotrgpolla<32> Tropidoporo(13) TronatolloCZI) Rhabdonoson<15) Ascopora(16) Nickolsopora<19> Neuataxis<18> 0rthoporo<20> Hagar-110(8) Buctropora<28> Ottosootaxts(38) Glauconooolla(4) Posnastulus<47) Neuatoporolla<41> thhrotrgpa<42> Osburnostulus(10) 8coptoporo<12> Rluunopora(45) Psoudohornoratlb) flrthroclo-a(2) 20 Figure 4a Ill Arthrostylus Arthrociema Cuneatopora Glaucommella HeIocIema Helopora Hexites Moyerella Nematopora Osburnostyius Pseudonematopora Sceptopora Tropidopora Ulrichostylus Rhabdomcson Ascopora Mediapora Nemataxis Nickelsopora Orthopora Tremateila Rhombopora Klaucena Megacanthopora Pamirela Primoreila Sationdotaxis Bactropora Nikiiorovella Acanthoclema Pinegopora Streblotrypella Hyphasmopora Ogbinopora Streblotrypa Petaloporella Streblascopora Ottoseotaxis Linotaxls Clam Nematoporella Arthrctrypa Hyalotoechus Spire Alwynopora Pseudohomera Pesnastylus 21 [--- firthrostulus(1) [69--- Neoctopora<9> [ ------------------------- 78 Ulrichostglus<14> I [ ----------- firthroclona<2> I I [--- Cunaatoporo<3> I I I -------------------------------- 100--- Hoytrclla<8> I l I [--- Holopora(6) 'I I I [93'-- Spira<44> I I I [-52 ----- Panirolla<25> I I I [-62 -------- Hodiapora<17) [51 I I [---50 ----------- Nonataxis(18) I I I I I I ----------- Ascopora(16) I I I I I I /--- H9phasnopora<33> I I I I /-52 His 100--- cobrnoporuao I I I I I I I I [-98 ----- Strablotrgpa<35> I I I I 1-66 \94 \-97 -------- Strgbrascopomca'n I I I I I I \ Nickolsopora<19> I I I I [67 \ r -— Rhabdouoson<15> l \-86 I I I [--- 0rthopora<20> I l l-94 l-56 \ 100--- Tronatolla(21) ' I I I I I \ Rcanthocloua(36> I I I I [-66 [--- Linotaxis<39> I I I I I \ ----- -95--- Hualotoochus<43) | I I I l [--- RhonboporaCZZ) I I I | I- [99--- Clousotrgpa<40) I I I I I [-99 ----- Hogacanthopora<24) I I I I [94 ['99 -------- K10ueona<23> I I [94 I l | [-99 ----------- Fri-oralla(26) + I I I I I I [99 Saffordotaxts<27> | I I I I I I [-99 [--- BactroporatZB) I I l I I l-87 I I \ ---------- reo--- cttosutaxrsaa) I I I I I I I \ ------- 86 /"' HIKIIOPOVIIIOCZO) | \-89 I \-94 I \ —69--- PataloporollaC36) I I I I \‘ PInogopora<31) I I I \-- StroblotrgpcllaC32) I I \ """""" = Oshnrnostulus<16> I I [--- Glaucononolla(4) I \ ------------- 100--- Posnastulustd?) I [--- Holoclona(5) + --------------------------- 93--- Seoptoporo<12> I I ----- Hoxitos<7> I [100 [--- Neuatoporollo<41> I I 100--- thhrotrupa(42) I l-BO [--- Psoudono-atoporctlt) | {-73 \---69--- Aluunopora(45) \ ---------------------------------- -63 \ ----------- Psoudohornnra<46) \ Tropidopora(13) Figure 4b 22 Figure 5a Anhrostylus Nematopora Ulrichostylus Arthrostyloecaa' Cuneatopora Helopora Glauconomefla Pesn Ius Taeniaosdtytctya Nematoporella Arthrotrypa Kielanopora HelocIema Hexites Pseudonematopora Tropidopora Rhabdomeson Ascopora Mediapora Nemataxls Nlckelso ta Tnamatal Rhombopora Megacanthopora Klaucena Pamirella PrimoreIIa Saffordotaxis Spira NikiIomveIla Acanthoclema PineQOpora StrebIotrypeIIa Hyphasmopora OgbinOpora Streblotlypa Petaloporella Streblascopora Maychella StrablocIadIa Callodadia Euthyrombopora CoelotubuIIpora Heliotrypa Mongloclema Linotaxis Clausotrypa Europora Hyantoechus Mo erella Ort opora Heminematopora Hemiuldchootylus Osbumostylus Scaptopora Bactropora Goldfussitrypa Nematotrypa Ottoseetaxis Kieleepora Ralflnella Oils ta Ena Iopora Parachasmatopora AIwynopora Pseudohornera Arthroclema 23 I--- fi-Urutylwfl) [100“- "autumn-«12) [100------ Ulrichostglw<17) ,’ 3" Seopt [--- Cmtopora“) ,' :% "output-(K?) [--- "clock-«6) [-96--- Strnblmopor <40) [-97 [--- Hun-:00) I \100--- Pmtoporau4> I79 I ------ Pimoporu(34) I I I I--- W66) | \-95-97--- 0¢inoporc<37> I \------ Stnblou-wa‘fl) I ------ 46 ----------- Puiullatze) I I l--- SaffordotuisGO) I +-----190--- Spit-068) I \---- ----- -- was) I [------ ”twat-«(20) I [- ------- -99 /--- BholboporaQS) l l \IOO--- momma?) I I [‘"n- Klan-lewd”) l-fi-BB I ----- -90 l--- PrisonllaQO) /-5: I I I I \-oo--- Heliotnpcfl'l) I I I I [--- mum“) I I Hu noon- Strqblocladtau3) /-94 | I I 1-99- ----- Cinemas) MK I I [03 --- Calloclautaaz) I I I \-82 \----97--- Coolotuwuporafls> ----------- (>44 I I \ ------------- ------- Nucleic-chum?) I91 \ ------------ - ------ ---- Reaper-«(19) l +- -------------- -- --------- out“) I I I ------ lelfmllataz) I \ ------------------ 90 [--- Potalopmllaam I \-96--- Honglocl-ufil ) l-Ba \--- Ltmtuistsz) l-9E l , I In ropidoporaflb) I I I [-97--- Ntdmlsoporfi I [100 * manual) I I I ‘. 1 <33) I I I I--- Nuclease” I +100 ‘\ 93 StrdalotrwcllaGS) \95 I I l--- flog-mlmun no: I I g c: Orthopmaa) | I [--- Enema!) | .‘ :3 Ottomtaxiflso) | \. Human) \ Mtyluua) / ----------------- RrU'rutuloociuQ) | [--- Gianna-110(5) I I ----- 190-" Pmtylufls?) I I ----- Tamflodictw(4§> | [100 [190 l--- N-atopmlhK“) | I \97 “09"- flr‘h-otrwaGS) ---+ | I \ --------- Rhiannon“ “2 /-67 +- ------------- Hutulrlchostuluw) I +44 l--- I“. Hie-amt” I I I l-68--- Emllopora(63) I | +------65 ------ Productopm (M) A 55 I ‘, Mlflmllatu) I | ~-----—- --------- Ruin-atopma) | | +. --------------- 011m“! ) | | \ ----------------- Elmer-«(65) | \ -------------------- PW(M) \ Boldfussunpaue) \‘ Whack-«2) Figure 5b 24 Figure 6a Arthrostylus Arthroclema Anhrostyloecia Cuneatopora GIauconomeIIa HeIocIema Helopora Heminematopom Hemiulrichostylus Hexites Moyerella Nematopom Osbumostylus Pseudonematopora Sceptopora Tropidopora Ulrichostylus Rhabdomeson Ascopora Mediapora Nemataxis NIckeIsopora Orthopora Trematalla Rhombopora Klaucena Megacanthopora PamireIIa Pn'rnoreIIa Saffordotaxns’ Bactropora NikiforoveIIa Acanthoclama Pinegopora Streblotrypella Hyphasmopora Ogbinopora Streblotrypa Petaloporella Streblascopora Maychella CaIIocIadIa StreblocIadia Euthyrombopora CoeIqubuIIpora Taeniodictya Heliotrypa Goldfussitrypa Nematotrypa Ottoseetaxis Mongloclema Linotaxis Clausotrypa Nematoporella Arthrotrypa Europora Hyalotoochus ira Kaalcepora RaIfIneIIa Ojlepora Kielanopora EnalIopora Parachasmatopom Alwynopora Psaudohomera Pu 25 l---- [ 54---- ' I \-....- I / ........... l +. .......... I h- --------- I l ,---- I I IIIO---- I I I \---- I +--96 I---- ' 1-91 \-90---- | I I I---- I I l l-9o---- I I +--oo I---- l I--oo I \-9o---- I I I +- ---------- I I I x ------- -- | I--oo I I---- l I I \ ----- "190-“- | I-eo \ I I as \ l I \ __ I I / I I t— I I | [---- | | I /-----oo---- | | I 1-73 I ------- ""+ | I I \--09 /---- I | I I--7a \-95---- I l I I \----------—--- I I | t- l I I-sa I /--- -------- I I I I I I I---- I I I +--71-----es I-oo---- l I I I I +--o¢ ------- I I I I I x ----------- I I I I I l---- l l I I +------------oo---- | | I I \ | I [--55 \ I I | ¢= I I l #- I [~63 I I , ....... I I I I t- 4:7 /---- I I I [-53 I \1.g---- I I I I I #- | I I I I \ I I I I :— l I I I--51 :- l I I l I \ I I I I f I I I I \ I I I I I I I I /--7a I I .............. \-62 I I I | :— | | l I +‘-n ’-”-------- I I I I I I +-- -------- I I I \ 73 \-83 I ------- I 4-70 I I I I---- I I l I \--78-96---- I I I I I \---- I I l I \ ------- I I l \ I l \ l \ \ Ifigun36b firthrostulus<1) NOIatoporc<12> Ulrichostulus<17> firthroclcla(2) firthrostgloociata) Honing-atopor¢(8) Honinlrichostylu<9> chitcs(10) Honotoporolla<34> firthrotrupa<55) Psoudoncnctoporo<14> Tacntodictuo<46) Strubloclodia<43> Parcehcsuatoporc<64> Ktulanoporo(62) fllugnopora<65J Bulfinclla<60> OjlnporctGl) OIOUOOBOIOIIOC5) Pashastglus(o7) PsuudohornoruCBOJ K1ulccpora<59) Osburnostglus Cunoatoporc<‘) Holoporo<7) Tropidoporu<16> Nickulsopora<22) Euthuro-bopora((¢) Clausotrgp¢<53) Ennlloporacoa) Tronatclla<24> Princrclla<29) Holiotrgp¢<47) "09°ccnthoporc(27) Colloclodia<42) Coclotubultporc<43) "clatotrgpa(49) Spirc<59) flcnataxtsCZI) Pauirnlla<20> Saffordotaxis<39) BuctroporoCSl) Ottosootaxis(30) N1k1forouclla<32> flccnthocla-OC33) floucrolla(11) Pin-gopora(34> Strcblotrupcllat33) <36) 09b1nopora<37) PctaloporullaC39) "ouch-110(41) Ooldfussitrupa<4l> flonqloclono(51) L1notaxts(52> Strtblotrgpc(3l) 8trcbloscoporc<49> Holoclc|a<6) Sc-ptoporo<15> 26 feature, causing very little structure to be seen in this consensus, and only a few of the branching patterns that are common in the other trees are present The majority rule tree for this data (figure 5b) also shows the effect of losing information by eliminating characters. Even in this tree there are ten polytomies. To have multiple polytomies in a tree of this type indicates extreme uncertainty in the tree structure. Tree six (figure 6a) has the same dominant polytomy that occurs in all of the consensus trees that lack the geochronological character. The majority rule tree (figure 6b) for this run of the data shows even more polytomies (16) than the tree with the temporal character (figure 5b), and also shows some very low percentages on several nodes. Trees seven and eight were run using the full data matrix as shown in Table 2 (with the exceptions noted in the appendix). I made all of the previous runs because I had achieved Iess than satisfactory results with the complete data set, and I had wanted to see if the exclusion of characters or taxa with missing data would improve the definition of the suborder. The same near basal polytomy exists in tree seven (figure 7a), which leaves the Arthrostylidae in this area of indecision at the base of the tree, along with the Bactroporidae, which has brought two new genera in to a small grouping with it. The internal polytomy in this tree is also very similar to the ones seen in other runs of the data, and is somewhat severe (28 branches). This polytomy includes most of the members of the other families mentioned in Blake 1983b (see also Table 5). The majority rule tree for data set seven is fairly strong, with high percentages at most nodes, 27 Figure 7a ArthrostyIus Arthrostyloecta Cuneatopora I-IeIocIerna Pseudonamatopora Hexites Hyphasmopora Ogbinopora Spira Tropidopora Rhabdomeeon Ascopora Medlapora Nemataxis Nickelsopora Tmmatella Rhombopom Klaucena Megacantnopora PamiraIIa Primorella Saflordotaxls Nikitorovella PetanporeIIa Acanthoclema Pinegopora Streblotrypalla Streblotrypa Streblascopora Maychella Strablocladia CaIIocIadIa Euthyrombopora CoeIotubuIIpora HeIIotrypa Mongloclema LinotaxIs H alotoechus C ausotrypa Europora Orthopora MoyareIIa Helopora Osbumostylus Nematotrypa Glauconomalla Pesnastylus Heminemat HemiuIrIchostyIus Nematopora Sceptopora Ulrichostylus Bactropora Ottoseetaxis Goldfussitrypa TaeniodIctya' NematoporoIIa Arthrotrypa KieIcepora RaIfinoIIa O Iepora KIeIanopora Enallopora Parachasmatopora AIwyrIopora Pseudohomem Arthroclema 28 8_____-__________\ p ’——————-————_——-— 3-“ 4’ -—-———*—-——\ I!” I “'- I!“ \'-!OO--- \ ........... 199 -99 \ \ I----- \------------99 /--- roo--- ’-.- ioo--- ,-_§::§:::r_. ’—— /_u-_ /_¢__? fl (bar-”1.91m“ ) Motown-1K 12) Ulrichostuluu?) Sccptopord 15> Brthrostulocciam) Gimme-0110(5) Pmtglmm?) Tomiodictuauo) "Camper-0110(8) flrtlrotnpaCSS) Humor-“62) Prochmtoporfl“) Hniulridtostglum) Kicle-poro(59 ) Emlloporc<63> scum-11am.) himtm") 011mm! ) Bactroporatal ) Dunc-mix“) Goldfmi Irma.) manor-a“) ‘ Hamper-0(7) MtotI-QpaGO) Oshtrnostglusc 13) Nancie-0(6) PMtopmt M) Hod test 10) (36) Ofiinoponfla'l) spa-«50> Swot-«56) Pair-allot”) legopor-«a‘n "cud-Idle“! ) Strobloclodlcua) LimtaxtsGZ) Mlotuchuflfl) Mime.) W005) KIWCZO) Primllflfl) Saffa-dotuxtsai) Haiku-wad?) Blanch-mafia) "mutiny-C(27) Cal loclodhKQ) CulottbuuPM-«OSJ Mk! rm11a Potatome “(3') hemmed-“33> smut-ml “(33) Tropidopor“ 16) Michele-1K3! ) Tract“ 1604) Nickolsmta) sauna-wean) sulucopm(fl) EU (44) /—————————————————8——————————————————\ ” l—s—p manna: ) (trumpet-«23) Hagar-.1 lot 1 1 ) Ema-“2) /__8__\ /__-————-———-_—————+-———————_-—-—-—-— Figure 7b 29 Figure 8a ArthrostyIus AnhrocIema Arthrostyloocla Cuneatopora GlauconomoIIa Heloclema Helopora Heminematopora Hemiulrichoctylus HexItas Moyerella Nematopora OsburnostyIus Pseudonematopora Sceptopora Tropidopora UIIIchostyIus Rhabdomoson Ascopora Medlapora Nemataxrs‘ Nickelsopom Orthopora Tramatella Rhombopora Klaucana Megacanthopora PaminaIIa Primorella Saffordotaxis Bactropora NikItoroveIIa Acanthoclema Pinegopora Strablotrypotla Hyphasmopora Ogbinopora Streblotrypa PetaloporoIIa Streblascopora Maychella CaIIocIadIa StreblocIadIa Euthyrombopora Coalotubulipora Taeniodictya Heliotrypa Goldfussitrypa Nematotrypa OttoseetaxIs Mongloclama LinotaxIs Clause NematoporoIIa Arthrotrypa Europora Hyantoechus Spira KIeIcepora Ralfinella OIIopora KIeIanopora EnaIIopora Parachas Alwynopom Psoudohomora Pasnutylus ‘II An I NI 30 firmwlufl 1) "camper“ 12) Ulriehostulufl 17) firthrocluntz ) 109m“ ) flour-l I“ 1 1 ) Pmtulusto 67 H-lulr-tchostglum > Hal thou-«33> Niki for-oval 10(32) ' (36> Debt-mama?) LtmtmdsGZ) Pctclopml “(39) Huguenot“) ---------------------- —oo \ {I /——————O———————\ "anoint-us: > Monm< (>25 Klan-rat“) Primlla<29> Saffordotcxls‘30) Bahama-«<31 > Mama-«27> Clmtrwo<53) Emlloporflfl) ,o____‘|_____\ ,'___§ ___\ ,r._._u_._.\ L- a'—3—3 50) Stroblocladtaua) ,r— P taper-cc“) KIIIWwZJ ”I "-'I “I ‘— Figurc 8b leatbs Hutu-alumna“) Hart 9..“ 10 ) "camper-cl 10¢“) Rimmed!) Khlcmtfl) PMtM) Ml final “(60 ) 011mm! ) 31 the exceptions being at some of the internal areas where the groupings are consistent, but the internal arrangement of the given group is in question. Tree eight (figure 8a) is another polytomy, the same problem inherent to the other atemporal strict consensus trees. The majority rule tree (figure 8b) has a basal polytomy, and has the added problem of low percentages on many basal branches. Trees nine and ten both use the entire data matrix but characters 2 (number of metapores) and 11 (development of the zooecial bend) were changed from an unordered state to an ordered state. This was done because progressions from none to few to many, and strong to weak to negligible seemed to have» a logical basis, even though some of the earlier runs did not support a clear order in these characters (see Appendix 1 for more details). Tree nine (figure 93) has a large polytomy one step up from the tree base, just as many of the other trees did, showing no clear relationship among the arthrostylids and other early genera. The upper part of the tree has a moderately well developed structure and has only two small polytomies. The majority rule tree (figure 9b) for this data set is strong with percentages dropping below 90% only in the arrangement of the arthrostylids. Tree ten (figure 10a), like all of the other trees that lack the geochronological character, consists only of a universal polytomy. The majority rule tree (figure 10b), however, shows a very strikingly different structure than the other trees in this study, but the supporting percentages on the basal branches are very low. Trees eleven and twelve used the full data matrix, but the coding had been altered so that the oldest genus Arthroclema is a true ancestral genus. What this means is that all of the characters were recoded so that the Arthroclema had all zeroes for its character 32 Figure 9a Arthrostylus Arthro Cuneatopora HeIocIema Pseudonematopora Hexites I-Iyphasmopora Ogbinopora Spira Streblotrypa StrebIascopora Europora Pamiralla Pinegopora Linotaxls Hyalotoechus Tropidopora Rhabdomeson Ascopora Nickelsopora Euthyrombopom Mediapora Rhombopora Klaucena PrimoreIIa Satfordotaxie Clauso Megacanthopora Maychella Streblocladla CoeIotubuIIpora CaIIocIadia Heliotrypa Mongloclema TrarnateIIa Nemataxrs’ NikIforoveIIa PetaloporeIIa Acanthocloma Streblotrypolla Orthopora Moyarella Helopora Bactropora Nematotrypa GIauconomeIIa Pesnastylus Heminomatopora Homiulrichoetylus Nematopora Osbumostylus Sceptopora Ulrichostylus Taeniodictya NematoporeIIa Arthrotrypa GoIdIussItrypa Ottoseetaxis Kielcepora Halflnella O'Iepora K eIanopora Enallopora Parachasmatopora AIwynopora Pseudohornera Arthroelema w /————_—_———_—_——_—+_—--———-——-———-—— s____-_____I______________\ ’ -—-—-—————_-—-—————- 33 “ ” _fl‘_——_‘-——“—— M /———8———\ I I /--- I I--reo--- I 1-04 I ----- I I \reo I--- I -oe \97--- I \ I10. 1"- I on. /——————8——————\ n /___8___\ p —____8____\ p ,__8___\ mum-- \ ----- m I--- \--IOI-- /——-—8———\ /_-_-__8______\ ,________-_s___________\ Figure 9b flrthrostulus<1> Nanotoporo(12) Ulrichostulus arthrostulo-cto<3) OlauconOIOIIOCS) Peshastqlus(67) Tooniodictgatdo) Nanotoporollc<54) arthrotrupo(53> Kielanopora<62) Honinlrichostolu<9> Kiolcoporo<59J Enallopora<63) Parachusnotoporctb‘) Bulfinclla<60) Huntncnatoporo<8> III-Mommas) 011cporc<61) Psoudohornnru(66) Boldfussttrgp¢(4fl) 0ttos..tnu13(5l) Buctropora<31> Honototrgpa<49> Holoelu-cCG) Psoudon-Iotoporocxa) chitcs<10) HUPhOSIoporo(36) 09bin0por¢(37) Spiro(58) Stroblotrupc<38> Strnblascoporo Linotaxis<52> H9clotoochus<57> Ntktferovollo‘32) Potatoporollc<39> flcanthoclo-a<33) Stroblotrupcllccas) Nanotaxis(21) Traptdoporarre> Hickclsopora(22> EuthUPOIboporc<44) Hodtuporc(20) Rho-boporc(23) KlauccnOCZB) Princrolla<29> Saffordotaxtst30) Clausotrgp¢(53) "coaccn thopor¢( "OUcholla(41) Strtblocladla(43) CoclotubultporoCGS) Callocladto<42) Holtetrgp¢(47) fl°fl910clooo(51) TrI-atolla(24) 0rth°p°ro<23> floqcrclla<11) °'°“'"°8tulus<13> arthroclo-nCZ) 27) 34 Figure 10a Arthrostylus Arthroclema Arthno Cuneatopora GIauconomeIIa HeIocIema Helopora Heminematopora HemiuIrIchoetyIus Hexites MoyereIla Nematopora Osbumostylus Pseudonernatopora Sceptopora Tropidopora UI ‘ us Rhabdomeson Ascopora Mediapora Nemataxis Nickelsopora Orthopora TramateIIa Rhombopora Klaucena Megacanthopora Pamirella Primorella Saftordotaxia Bactropora NIkItoroveIIa AcanthocIema Pinegopora StreblotrypeIIa Hyphasmopora Ogbinopora Streblotrypa Petaloporella Streblascopora MaychoIIa CaIIocIadIa Streblocladia Euthyrombopora Coelotubulipora Taeniodictya HeIiotrypa GoIdIussItrypa Nematotrypa Ottoseetaxts Mongloclema LInotaxis Clausotrypa Nematoporella Arthrotrypa Europora Hyantoechus Spire KIeIoepora RaIfIneIIa OIIepora Kielanopora EnaIIopora p Alwynopora Pseudohomera Pas ’-§:a-. 3-x I ’—3——\ ”'3-‘h ’v_.ro..\. 35 I ----- I ------- -05 I--- \94--- / a? ‘l l”— / " -———-————————— + —-—————-——-——-— ,_é__\ ,_-é__ 3___\ I-— ,_-_é____\ /——-———g—————\ I I“ /:—\ ’--- \ ,_$__ :se--- /----- I ----- -04 I--- I tee-~- I91 I--------- I I I l--- I \-07 tee--- I I-0e----- \00-------- -0- ’-------- I I l--- \-----90 I90--- \-07----- ’--- I99--- I-oo I--- I \95--- on I--- \--07--- ’---- =00 I--- I..--- ’--- :9e--- ’--- I99--- 02 I--- reen- Figure 10b armour-u ) 11.09.0000“ 12) Int-1011001310“ 17) hthroclma) 610500100090“) Hoyt-01 10( 1 1 ) mun-<13) 1101001000“) 800ptopor0( 15) hmwlmma) 01000000001106) Pmtyluflfi?) Hutulridnostuluw) H.10000t000r-00) 0] 10por0<61 ) K1010Q090<59) PWGO) B01 f1rr0110<60) Hod 100(10) “moor-0110(8) htI'I-otrwdfl) SN10010010GS) PMW Elma-065) - K101mpm<62) PWtopm< 14) 100010010th5) loch-0900061 ) Ottosutcxiflfl) Troptdoporoub) 001de! ”0(0) H01101np0(47) Print-0110(29) 80ff00-0010xtfla) Klmafi) 1101000000) 8010060) M< 10) "101(0180900002) Euthuro-bopora<44> 33009001“ 19) Wane) ”mood-067) smut-worse) SNIWGO) Mt0x10(21 ) Orthopmaa) Tr-010110(20) H9010“00st(57) W66) Pair-0110a.) "(Ufa-00011002) 901010000011000) P1w<34> 3100010u-Ip01 10(35) flannel-0(3) L1mtu10<52) 6001mm.) mud“ ) 11009100100061 ) C011001d10(42) €0010tthu1 tom-0(a) W005) "WW0(27) c1mtnp0(53) Emulator-0(a) mamas) 36 Figure 11a ArthrostyIus Arthro ' Cuneatopora GIauconomoIIa Pesnastylus I-IeIocIama Pseudonematopora HexItes Hyphasmopora Ogbinopora Spira Tropidopora Rhabdomeson Ascopora Mediapora Nmtaads Nickels ra Tmmatel Rhombopora Klaucena Megacanthopora PamiraIIa PrimoraIla Saffondotaxis NIkiIoroveIIa PetaloporeIIa Acanthoclema Pinegopora Streblotrypolla StrebIorrypa Streblascopora Maychella Streblocladia CaIIocIadIa Euthyrombopora CoelotubuIIpora Heliotrypa Mongloclema Linotaxis H alotoechus C usotrypa Europora Orthopora MoyereIIa Helopora Heminamatopora Hemiulrichostylus Nematopora Osburnostylus Sceptopora UIrichostyIus Bactropora TaenIodIctya Nematoporella Arthrotrypa Goldtussitrypa Nematotrypa Ottoseetaxis Kielcepora Ralflnella O Iepora Kelanopora Enallopora Parachasmatopora Alwynopora Pseudohomera ArthrocIema 37 8-———————————————-——-——-—\ §____-______________\ ’ ———-———--———_-——_—_—— ’—-_-_"—-————————“— ,_-_______é__________ /——_—d_____\ Figure 11b P-hflnwwmfln ['99-- “tum-0‘12) l9I---- Ulrichostglus<17> I 75 - 800pt00000<15> I I--'-- flrthr0019100010(3) I I l-- H00in000topor0(8) I l ------- “-5941" tumor-0(a) I I \---- Ojlnporo<01> I I I- 0100000000110(5) I I I-1OI-- P0snastulus(67) I I I I-- T00010010t90(46) 48 P“ I45hflWflbmnmunGm I I I I93 199----- H000t00000110<54) I I I I-ee \ nnnnnn H00101r10h00t910<9> I II l\~«-~n~flnuunmqmmmn I I99 I I I-- K1010000r0<59) I II W6 I4?~Euflqmwwm I I I I ’73--- 001(100110(69) \-92 I \ ----- -61---- K1010009000<62) :\ I gmrwm-mmu '“ fmfluunuv \ :96-- 0tt0000t0x10(5|) I'- Cun00t00000<0> I”~HquwO) h-mum-mm) I199-- Ps0udon000t0poru<14> 1”---'- M1W I90 \199-- 8tr0bloc10010<03) I I I-- Linotauis<32> I\ ammmpuu-nswv I I Hour-poroc20> PWI I mnummua> I I I I I'--- K10000n0(25) I I I I I9? I- Fri-0r0110(29) I \90 I96 I-oe \-90-- earrordotauistao) I I I I [99 \------ H011010gp0<47> I l I \-95 \—------ Cl0usotrgp0<53) I V” V”“””““ Hanna” I I I“ Calloclod1o(‘2) N7 \ .Jwanummuumma> I I I“""'-"'- WHO) I I I I------ fl00000r0(19) II Pfll fiwwn-MQMMMMunD I I I \9! I I- 81r0blotrup0(39) flBI I \Hfllfltn9hfikmmmmun III N1 “Emu-h (fl) I I I I I I'- H1k1f00000110<32> [~96 I \ ----- -98 \ ------ --199-- P010109000110<39) I I I \ flcnnth001000(33) I I \ SNIoWUaCfl) 199 \ H000t0u10<21> I I I- 70001000000(16) [199 I I-99-- H0091001000(51) I I \ fl Tmulllflzi) 100 \ G'thopora(23) ~95 \ H090r0110(11) \ B0ctr00000(31) Ihnuflunflfl 0000rnostulus<13) —~m~u-mm> 38 Figure 123 Arthrostylus Arthrocloma Arthrostyloacla Cuneatopora Glauconomella HeIocIema Helopora Heminematopora HemiuIrichostyIus Hexites MoyeroIIa Nematopora Osbumostylus Pseudonamatopora Sceptopora Tmpidopora UIn'chostyIus Rhabdomeson Ascopora MedIapora Nemataros' Nickelsopora Orthopora TremateIIa Rhombopora Klaucana Megacanthopora PamireIIa PrimoreIIa Saflordotaxis Bactropora NIkIforovelIa Acanthoclema Pinegopora StrebIotrypeIIa Hyphasmopora Ogbinopora StrebIotIypa PetaloporeIIa Streblascopora MaycheIIa CaIIocIadia StreblocIadIa Euthyrombopora Coelotubullpora Taeniodictya Heliotrypa GoldtussItrypa Nematotrypa Ottoseetaxis Mongloclema Linotaxis Clausotrypa Namatoporella Arthrotrypa Eumpora Hyantoechus Ira EIaeIcepora RaIIIneIIa OIIepora KIeIanopora EnaIIopora Paradrasmbpora AIwynopora Psoudohomera Pesnaetylua /————————————+—————————————3 39 ’--- I------99--- I \--- I-oo I-------- I I I--- \-05 I97--- \-90----- l--- I--- roe-~- 100 IJ8-—‘0 ,_I__, 0 ’0——.‘———\ ’--- I--reo--- I I--- I-90 I93--- I \-00----- I-00 /--- I I99--- \----02 I--- \02--- 3---- I-oo----- I0 I —-—”——— I---— —9e--- 3“? d’--:'--‘u l-- [~95 _fl‘ "I ’--- \--199--- \----------- I /_-_£___ 4’-—J&-—-‘u ’1’. —-————— II. ————————_————__—-—_—————\ d'---—It-—--—-—-—---——-——- kin-00191000 ) 11.01.0009“ 12) Ulrtehostulust17) firth-001.0(2) 1101001000“) Oahu-0001910“ 13) 800910900“ 15) Pmupord 14) Tamiodtctwua) ”tn-«(91000106 ) 0100000000110(5) .Pmtulw‘fi?) Hawk-1015001910“) H-1n-0t000r-00) 011000I-0(61 ) manor-0(4) flood-01 10( 11 ) 00000000003) L100t0x10<52) heathen-063) um {00090110(32) P0t01000r~0110<39) P100909000(30) . 8N10W0110<35) 11901010000005?) PCU‘OI 10(20) (19) N10k018090r0<22) ”tumor-0(1)?) 8.01.0:10Q1 ) 1401000000) San-0(a) Nimrod.) M110<41 ) ”unlock-“51 ) C011001d10(02) “0101:0001 100F003) (25) Klan-0“”) "mm-ecu) ClmW(53) amnesia-0(a) mamas) 700001.0110(24) Traptdoporocre> III-1000011009) “flu-detain”) intone-061 ) Ottmtodua) 11011001.pdf!) (“1de1 W001.) smug-40010013) PWtopou-q(“) flleS) K101at000I-0(62) K1010qa0r0(59) Panoramic-0(a) 801f1m110(69) M11000.) "-01.000001 10(54) NMWG5) states in the data matrix. This was done because it is the oldest taxon in the study which, if the fossil record were complete, should be the ancestor; however it has more in common with some of the other younger taxa and varies on some points from the trends in the other older taxa. These are the reasons that it was not used as the sole basis for character polarization. Tree eleven (figure 11a) has a structure with a polytomy one step up from the base of the tree, as well as a large internal polytomy similar to trees 5 and 7. Figure 11b (the majority rule data) shows two large groupings as well as the ancestral genus, but the reasons for the polytomies in 11a are clear because the majority rule data shows mid to low range percentages in many locations. Tree twelve (figure 123) has the same universal polytomy that is present in all of the other trees that lack the geochronological character. The majority rule information (figure 12b) for the tree shows a fairly good structure, but the percentages around the base are ofien well below 90%. DISCUSSION As mentioned above, in this study the geochronological character was invaluable in the production of a definable tree structure in the Rhabdomesina, as shown strongly in the strict consensus trees. The 50% majority rule trees also showed structure, but the percentage of consistencies among the trees were ofien quite low in the non- geochronological trees, which shows their apparent weakness in comparison to the geochronological trees. For an analysis of the familial structure, unless otherwise stated, 41 TABLE 5: FAMILIAL GROUPINGS USED BY OTHER AUTHORS .6 3:03:05 .0 vocsfioE .6 cocoacoE 42 TABLE 5: CONTINUED .5 30:08:25. 50 VOCQEOE 43 I shall be referring to the families as defined by Blake (1983a), but I have included a table listing the groupings defined by other authors as well (see Table 5). I will use the majority rule trees from data sets seven, nine, and eleven; even though other trees were better in some statistical ways, I will use the trees based upon the whole data set for this discussion. Though there are many shifts in overall tree structure, several similarities in generic position occur in all three trees. First, the family Arthrostylidae has split up. There are two well-defined groups, and a number of genera that have scattered around the tree. An interesting development is that all of the unilaminate arthrostylids are grouped together with all of the unilaminates that were not included in Blake 1983b (see Table 3) in one clade, with just the unilaminate genera Arthrostylus and Streblocladia appearing elsewhere on the tree, however the internal arrangement of genera is variable. Another aspect of this group is that three radial genera appear consistently, Arthrotrypa, Nematoporella, and Taeniodictya. The other main stable arthrostylid group is much smaller, and unlike the unilaminate group its internal arrangement seems stable. This group consists of the genera Arthrostylus, Nematopora, Ulrichostylus, and Sceptropora. The radial arthrostylid Arthroclema, the oldest genus in the suborder, always appears alone at the base of the tree (as I have mentioned above). Eight other arthrostylids are in other places around the tree. The family Rhabdomesidae has been scattered, but a small grouping has developed around the type genus. This small cluster consists of Rhabdomeson, Ascopora, Nickelsopora, and E uthyrombopora, and in two of the three trees Streblotrypa and Streblascopora are also present The Rhomboporidae stayed together fairly well, and also incorporated several of the uncertain genera, in a large consistent grouping. This group includes Rhombopora Klaucena, Primorella, Safl'ordotaxis, Helionypa, Clausomrpa, Megacanthopora, Callocladia, Coelotublipora, and Mediapora. The Bactroporidae jumped around the tree, sometimes pairing with Cuneatopora and Nematotrypa, other times matching up with Ottoseetaxis and Goldfizssitrypa, but it did not have a stable position among the three trees. The Nikiforovelidae was dismantled as a family; genera placed in this family were often within the same larger clade, but not together in a cohesive unit In the Hyphasmoporidae the genera were split up with only two groups of two genera consistently together. Streblotrypa and Streblascopora, as well as Hyphasmopora and 0gbinopora, but the placement of these pairings was unstable. There were also several pairings that were consistent in the three trees but their locations were variable. These were: Cuneatopora and Helopora, Ottoseetaxis and Goldfilssinypa, Linotaxis, and Hyalotoechus, and Nikg'forovella and Petaloporella. Only one large consistent grouping was found that did not precisely match up with the current familial structure, including: H eloclema, Pseudonematopora, Hexites, Hyphasmopora, 0gbinopora, and Spira, and in two of the three trees E uropora and Pamirella as well. It does not seem that any large scale reworkings of rhabdomesine familial structure can be made from this study, due to a lot of ambiguity in the high numbers of 45 trees; however it does seem that the unilaminate arthrostylids should be placed in their own family. One other finding of note is that all of the trees that showed structure divided the suborder into two general groups, one that included most of the arthrostylids, and one that included the majority of the remaining families. Both clades had somewhat variable members depending upon which tree was observed, but several members of each clade remained the same. This information correlates with some of the findings of Anstey and Pachut (1995). In their study of all Paleozoic bryozoan families, the Arthrostylidae were in a group with the fenestrates, and the Rhabdomesine families were in another group. This two clade structure is possibly the most important result of this study, as it shows a possible polyphyletic origin for the suborder, and when compared to prior studies it shows that the Arthrostylidae may have stronger affinities to the fenestrates than the rhabdomesines. FUTURE WORK Future work based upon this study should go in two directions: 1) Use this study as a starting point for a more thorough study of the original type specimens, 2) There are less than 450 genera of Paleozoic bryozoans known, and an analysis including all of them would provide a means of looking at the relationships of all of the genera without starting with predesigned orders to trap studies of the systematics. I believe that this more comprehensive generic level study of Paleozoic bryozoans as a whole could very well clarify the phylogeny of this group, and answer some of the questions that this study could not. APPENDIX A APPENDIX A List of chiacters and their states along with their types and a brief explanation of each. L Apertural arrangement: 0 linear rows, 1 rhombic, 2 irregular [unordered] This character describes the arrangement of the apertural openings in the colonial skeleton. The most primitive state is that where the openings are in a linear arrangement that runs down the length of the colony, usually two to four rows are encountered. The rhombic arrangement is most easily described as a situation where the openings are in diagonal rows, where the apertures (when taken in groups of four) form rough rhombuses. The irregular arrangement, as can be deduced by its name, are the colonies that do not seem to have any consistent pattern to their apertures. Originally I thought that there seemed to be a noticeable trend from linear to rhombic to irregular when the genera were looked at through time, however I later decided that, all I could be sure of was that the linear state was primitive, and the other two states were derived, but not necessarily from one to the other. In the final analyses this character was dropped in favor of characters 15, 16, and 17. These characters looked more specifically at the three aspects of character 1, whether or not the Openings are rhombic, linear or organized. L Metapores: 0 none, 1 few or not in all species, 2 many [unordered] Metapores are variably sized cavities in the surface of the zoarial wall, most likely used to space the zooecia in the colony. They are present in many genera, but not always in all species of the genus. Also there are varying numbers occurring on different genera, 47 48 some have only a small number scattered around the zoaria, while others are covered almost solidly with them. The absence of this structure is clearly synapomorphic, but there never seemed to be a clear trend between state 0 and the other two states. I tried nmning this character as both an ordered and an unordered character, with few differences being shown, but the consistency index for this character was higher in the unordered state, so I left it that way for the most of the final analyses. The exceptions to this were trees nine and ten, which were run to see if their ordering had a significant impact. 1 Paurostyles: 0 no, 1 yes [ordered] Characters 3-6 all deal with the presence or absence of various types of spine like protrusions from the external zoarial wall, approximately perpendicular to the zoarial surface and parallel to the zooecium. There are four different types of these stylets that exist within the suborder, which are all pictured and described in Blake (1983a pp. 537- 541). The first that I have listed is the type known as paurostyles. These stylets are the smallest and most diffith to see, they have somewhat clear cores and appear as small deflections in the lamellae of the wall. I usually had to rely on written descriptions to account for this character, as these structures are very difficult to see in photographs. Their presence was apomorphic due to their inconsistent presence in the early genera of the suborder. _4_._ Acanthostyles: 0 no, 1 yes [ordered] The second and most prominent type of stylets were the acanthostyles. These stylets are the largest type of stylet, which can be seen as large hollow cores that run 49 perpendicular to the surface of the zoaria; these were seen in many genera. Their presence was also apomorphic due to inconsistent presence in the early genera (See also character 3) ; Aktinotostyles: 0 no, 1 yes [ordered] The third type of style is the aktinotostyles; these are also quite large structures which can be differentiated from the acanthostyles by their dark cores. Their presence was apomorphic due to their rare occurrence, being in only a few genera (See also character 3). Q Heterostyles: 0 no, 1 yes [ordered] The final type of stylet is the heterostyle; these small rare structures only occur in three genera, and for that reason their occurrence was considered an apomorphy. These structures were difficult to see and I had to rely on written descriptions to use them in the data matrix. (See also character 3). 1, Vertical axial zooecia: 0 no, 1 yes [ordered] This character is somewhat rare and exists only in a few genera. It looks for the presence of a cluster of zooecia in the central region of the zoaria that are running parallel to the colony branch; its rare presence is an apomorphic state. L Branch cross-section: 0 subcircular, circular, round, 1 polygonal [ordered] Character eight was looking at the overall shape of the zoarial branch in cross section. State zero was used for any degree of roundness that was observed in the taxa, while the apomorphic state was reserved for the younger occurrence of a rigid polygonal state. 50 2, Longitudinal ridges: 0 yes, 1 no [ordered] This character was looking for ridge-like structures that ran the length of the zoarial branches. Since they were common in the older taxa I decide to make their presence the plesiomorphic state. _1_9_. Zoarial jointing: 0 yes, 1 no [ordered] This character dealt with the presence of segmented joints in the zoarial branches. As the character was common only in the older taxa I decided to make its absence the apomorphic state. fl Zooecial bend: 0 strong, 1 weak, 2 negligible [unordered] The zooecial bend is the curvature of the zooecial chamber as it runs from the zoarial surface to the center of the zoaria. As in many of my unordered characters, I could easily determine the plesiomorphic state, but had difficulty deciding an order for the other states. I experimented with it both ways and from the trees that were produced I decided that I could not justify a specific order for this character, however I did use this as an ordered character in the runs that created trees nine and ten, to see ifthere was an impact on the final analyses. 1;, Zooecia on both sides: 0 no, 1 yes [ordered] This character describes the unilaminate or radial form of the zoarial branches. The simplest description is the one I originally used in the character name, a state where zooecia exist on only one side. Almost all of the unilaminate genera are early in the 51 history of the suborder, so that is why having zooecia on both sides is an apomorphy. (See character 23) 1_3_. Time: 0 Ordovician, 1 Silurian, 2 Devonian, 3 Carboniferous, 4 Permian (not used in final study in favor of character 14) [irreversible up] This coarse grained geochronological character is a carry over from when I did not have enough information to complete character 14. l_4_. Time: 0 Tremadocian, 1 Arenigian, 2 Llanvirnian, 3 Caradocian, 4 Ashgillian, 5 L Silurian, 6 U Silurian, 7 L Devonian, 8 M Devonian, 9 U Devonian, 10 L Carboniferous, 11 M Carboniferous, 12 U Carboniferous, 13 L Permian, 14 U Permian [irreversrble up] This geochronological character is one of the major focuses of my study. As I have previously mentioned not everyone sees the value in the use of time as a cladistic character, but in this study this character was the only saving one that was able to put conclusive order in the strict consensus trees. The irreversrble up setting allows for the character to change as far as it needs to in the higher states, but cannot change backwards (as time cannot go backwards). I; Ordered zooecial arrangement: 0 yes, 1 no [ordered] This character looks for organization in the arrangement of the apertural openings, by looking at the older genera in the suborder 1 determined that order is plesiomorphic, and that the genera showing strong disorder were apomorphic. ( See character 1 for further discussion). 52 Q Zooecia in rows: 0 yes, 1 no [ordered] This character regards the linear arrangement of the apertures, usually in two to six rows along the zooecial branches. By looking at the oldest genera in the suborder I decided that linear arrangement was synapomorphic and non-linear arrangements were apomorphic. (see character 1 for further discussion). 11 Zooecia rhombicly arranged: 0 no, 1 yes [ordered] Much like the two previous characters this one deals with the arrangement of the apertural openings. As the oldest genera were all arranged in a linear fashion, I used the absence of rhombic arrangement as the plesiomorphic state, and the presence of this arrangement as the apomorphic state. (See character 1 for further discussion). 1_8. Branched zoaria: 0 no, 1 yes [ordered] This character is used to describe zooarial forms that have a branching pattern of some form, but not a pinnate one (see character 19). An extreme majority of the zoarial shapes displayed by this group are branched in some way, however a few of the earliest genera, including the oldest one, are single unbranched forms. Character 18 was used to show this difference, branching was considered an apomorphic state, while non branching forms were considered plesiomorphic. fl Pinnate zoaria: 0 no, 1 yes [ordered] This character was originally included when I thought that a clear connection to the fenestrate bryozoans (which often have pinnate or grid-like forms) was going to be more noticeable. However when I began to code characters I realized that almost none 53 displayed this unusual branching form, and in the end only two taxa were apomorphic for this state, Glauconomella and Pesnasozlus. 29, Hemisepta: 0 none, 1 upper wall, 2 lower wall, 3 both walls [unordered] This character deals with protrusions from the walls of the zooecial chamber, and which walls they protrude from. Again, the plesiomorphic state was easily determined but the order in the apomorphic states was not clear. I tried running this character in an ordered state, but even then no clear patterns emerged, so I left the character in an unordered state. __l_. Peristomal ridges: 0 yes, 1 no [ordered] Peristomes are small ridges that form around the zooecial opening in the colony branches. These ridges are common in the older taxa of the suborder, and that is why I decided that their presence should be plesiomorphic and their absence apomorphic. 22, Stylets: 0 no, 1 yes [ordered] This character was to reinforce situations when I knew that stylets were present but could not be completely certain of the type. In following with the other stylet characters I made their presence an apomorphy. Q. Zooecial pattern in cross-section: O radial, l bilaminate, 2 strongly radial, 3 close packed, 4 hollow central chamber, 5 unilaminate, 6 hexagonal, 7 NOT USED, 8 medial rows (not used in final study in favor of characters 12, 26-32) [unordered] This character is a description of the arrangement of the zooecia and the other internal structures as viewed from a cross section of the zoarial branches. I tried for 54 some time to find a way for this character to be put in an ordered state, but to no avail. The blank spot in state 7 is due to a state that I had thought was to be included, but only after the information had been entered into the computer did I realize that it was not needed See the aforementioned characters for details of the states; all characters have the presence of the state as an apomorphy. 2_4_. Large central style: 0 yes, 1 no [ordered] This rare trait was seen in only a few genera, and since only one occurred outside of the Ordovician I decided to make the trait a plesiomorphy, and the lack of it an apomorphy. 2_5, Zooecial shape (as defined in Blake, 1983a, pg. 535.) 0 type 7, 1 type 1, 2 type 2, 3 type 3, 4 type 4, 5 type 5, 6 type 6 (not used in final study in favor of characters 33-39) [unordered] The trait can only be best described by looking at the figure in Blake 1983a (reproduced as Figure 13). Each type is a different kind of zooecial chamber construction The types range from a nearly perpendicular chamber in type 6, to a straight diagonal chamber like in type 1. Originally I tried to polarize this character, but had too many difficulties in deciding the order after the first, so I converted the character into seven smaller characters where each type was apomorphic for the presence of a given type. (See characters 33-39). 26, Hollow central x-section: 0 no, 1 yes This describes the hollowness of the center of some zoarial branches, in some cases possibly due to a form of encrusting behavior. (See character 23). 55 Figure 13 Taken From Blake 1983b, pp. 535 56 _2_]_._ Radial central x-section: 0 no, 1 yes A radial cross section appears to have the supporting walls of the zoaria radiating out from the center of the branch. (See character 23). _2_8, Bilaminate central x-section: 0 no, 1 yes The zooecial chambers have a bilaminate pattern in this type of cross section. (See character 23). _2_9_. Trepostome like central x-section: 0 no, 1 yes This character has a cross section that is convergent with the kind that is seen in the order Trepostomata. All of the zooecial chambers are closely packed together in a tight bundle. (See character 23) 39, Strongly radial central x-section: 0 no, 1 yes This type of cross section is similar to the radial type, but the radiating wall structures have a strong spoke like look to them. (See character 23). 31, Medial rows in central x-section: 0 no, 1 yes The type of cross section that has medial rows will have one or two rows of zooecia that will seem to be parallel to the branch and will be the center for the lateral zooecia. (See character 23) 32, Hexagonal central x-section: 0 no, 1 yes A hexagonal cross section has a radial appearance, but the spokes are so strong that the entire branch takes on a hexagonal shape. (See character 23) _3, Type 1: Ono, 1 yes (See character 25) 3_4_. Type 2: Ono, 1 yes (See character 25) 3; Type 3: Ono, 1 yes (See character 25) 3_6._ Type 4: Ono, 1 yes (See character 25) 3_7, Type 5: Ono, 1 yes (See character 25) 38, Type 6: Ono, 1 yes (See character 25) 3_9_. Type 7: Ono, 1 yes (See character 25) 57 LIST OF REFERENCES LIST OF REFERENCES ANSTEY, R. 1990. Bryozoans. p. 232-252. In K. MCNAMARA (ed), Evolutionary trends. Belhaven Press, London. ANSTEY, R. and PACHUT, J. 1995. Phylogeny, diversity history, and speciation in Paleozoic bryozoans. p. 239-284. In D. ERWIN, and R ANSTEY (eds), New approaches to speciation in the fossil record. Columbia University Press, New York. BLAKE, D. 1980. Homeomorphy in Paleozoic bryozoans: a search for explanations. Paleobiology 6: 451-465. - BLAKE, D. 1983a. Introduction to the Suborder Rhabdomesina. p. 530-592. In R. ROBINSON (ed), Treatise on invertebrate paleontology part G, Bryozoa (revised) Geological Society of America, Inc. and The University of Kansas, Boulder and Lawrence. BLAKE, D. 1983b. The Order Cryptostomata, p. 440-452. In R ROBINSON (ed), Treatise on invertebrate paleontology part G, Bryozoa (revised). Geological Society of America, Inc. and The University of Kansas, Boulder and Lawrence. BLAKE, D. and SNYDER, E. 1987. Phenetic and cladistic analyses of the Rhabdomesina (Bryozoa) and similar taxa: a preliminary study. p. 33-40. In J. ROSS. (ed. ), Bryozoa: past and present. Western Washington University, Bellingham. CUFFEY, R. 1973. An improved classification, based upon numerical-taxonomic analyses, for the higher taxa of Entoproct and Ectoproct bryozoans. p. 549-564. In G. LARWOOD (ed), Living and fossil Bryozoa. Academic Press, New York. CUFFEY, R and BLAKE, D. 1991. Cladistic analysis of the Phylum Bryozoa. p. 97-108. In P. BIGEY (ed): Bryozoa living and fossil. Bull. Soc. Sci. Nat. Ouest Fr., Mem. HS 1. 58 S9 DZIK, J. 1992. Early astogeny and relationships of the Ordovician rhabdomesine bryozom. Acta Palaeontologica Polonica 37: 37-54. DZIK, J. 1994. Bryozoa of the Mojcza Limestone. Palaeontologia Polonica 53: 253-282. FISHER, D. 1994. Stratocladistics: morphological and temporal patterns and their relation to phylogenetic process. p. 133-171. In L. GRANDE, and O. RIEPPEL (eds.), Interpreting the hierarchy of nature from systematic patterns to evolutionary theories. Academic Press, San Diego. HUELSENBECK, J. 1994. Comparing the stratigraphic record to estimates of phylogeny. Paleobiology, 20: 470-483. MADDISON, W. and MADDISON, D. 1992. MacClade 3.05, Sinauer Associates, Inc. Sunderland, MA. MAYR, E. and ASHLOCK, P. 1991. Principles of systematic zoology. McGraw- Hill, inc. NY. ROSS, J .R.P. eta]. 1996. Bryozoan evolution and dispersal and Paleozoic sea- level fluctuations pp. 243-258 In: GORDON, D.P. et. al. Bryozoans in space and time. National Institute of water and Atmospheric Research Ltd. Wellington, New Zealand 442 p. SCHULGA-NESTERENKO, M, et.a1. 1972. Order Cryptostomata p. 111-148. In T. SARYCHEVA. Fundamentals of paleontology Vol. 7, Bryozoa. Indian National Documentation Centre, New Delhi. SWOFFORD, 1993. Phylogenetic Analysis Using Parsimony. Smithsonian Institution, Washington, DC. "‘liliflllllllllllll“