pictorial guide |
ORDER PTYCHOPARIIDA last revised 11 NOV 2019 by S. M. Gon III Introduction: A large, heterogenous order with
primitive origins and problematic classification, with
specialized offshoots that are hard to frame within a
general diagnosis. |
Ellipsocephaloidea .Yunnanocephalidae Ptychoparioidea Menomoniidae Norwoodiidae |
Suborder Ptychopariina
Introduction: Primitive Ptychopariida, a large
and extremely varied group. Superfamily Ellipsocephaloidea
Aldonaiidae: Aldonaia, Granutaspis, Ideria, Perissopyge, Planaspis, Pumilina, Repinaspis, Tuvanella (=Eleganolimba), Tuvanellus, Volonellus. Bigotinidae: Bigotina, Bigotinella, Bigotinops, Bulaiaspis, Hupetina, Neobigotina, Ouijjania, Pruvostina, Serrania. Chengkouiidae: Acanthomicmacca (=Chengkouia; =Jaskovitchella; =Myopsomicmacca), Bidjinella, Changyangia, Micmacca, Turkestanella, Wenganella, Xiuqiella, Zacanthellina, Zhenbaspis (=Yankongia; =Zhenxiongaspis). Ellipsocephalidae: Acadolenus, Alueva, Antatlasia, Argunaspis, Asiatella, Bergeroniaspis, Bergeroniellus, Blayacina, Brevitermierella (=Paratermierella), Cambrunicornia, Catadoxides, Charaulaspis, Chorbusulina, Comluella, Culmenaspis, Ellipsocephalus (=Germaropyge), Ellipsostrenua, Glabrella, Hamatolenus, Hupeolenus, Issafeniella, Kadyella, Kameschkoviella, Kijanella, Kingaspidoides (=Elatius), Kingaspis (=Mesetaia), Krolina, Kymataspis, Latikingaspis, Latouchia, Latuzella, Lermontovia, Limataceps, Limouolenus, Lotzeia, Lusatiops (= Jalonella), Mohicana, Myopsolenus (=Collyrolenus), Myopsostrenua, Nelegeria, Olekmaspis, Ornamentaspis, Orodes, Ourikaia, Paramicmacca, Paraprotolenella, Pauliceps, Planolimbus, Protagraulos, Protaldonaia, Protolenella, Protolenus (/Bergeronia; =Matthewlenus), Pruvostinoides, Pseudoasiatella, Pseudokadyella, Pseudolenus, Pseudoprotolenella, Ptychoparopsis (=Berabichia), Rinconia, Sailycaspis, Sectigena, Strenuaeva (=Hindermeyeria), Strenuella, Tadakoustia, Termieraspis, Termierella, Thoralaspis, Timnaella, Triangulaspis (=Acutaspis; =Angustaeva; =Plenudiscus; =Triangullina), Yeshanaspis. Estaingiidae: Alanisia, Chulanolenus, Coreolenus, Eomalungia, Estaingia (=Hsuaspis, =Pseudichangia; =Zhuxiella; =Sematiscus; =Strenax), Hupeia, Ichangia, Longmenshania, Longxianaspis, Madianaspis, Mundocephalina, Ningxiaspis, Olekinanellus, Paraichangia, Pararaia (=Proichangia; =Tannuolaspis), Protolenoides, Shangsiaspis, Shifangia, Shiqihepsis, Sichuanolenus, Subeia, Szechuanolenus, Yinshanaspis. Palaeolenidae: Alataurus, Bajangoliaspis, Enammocephalus, Ferralsia, Gigoutella, Hybrocephalus, Hoffetella, Latipalaeolenus, Megapalaeolenus, Palaeolenella, Palaeolenides, Palaeolenus, Resimopsis, Schistocephalus, Torgaschina, Ulakhanella, Validaspis. Yunnanocephalidae: Elicicola, Luaspis, Pensacola, Wangzishia, Wenganlenus, Yunnanocephalus (/Pseudoptychoparia). Superfamily Ptychoparioidea
Alokistocaridae: Alokistocare (=Pseudoalokistocare), Alokistocarella, Alokistocaropsis, Altiocculus, Amecephalina, Amecephaloides, Amecephalus (=Strotocephalus), Annamitia, Arcadiaspis, Arellanella, Atopiaspis, Beldirella, Binella, Bythicheilus, Chancia, Chanciaopsis, Danzhaiaspis, Diaoyaspis, Ehmania, Ehmaniella (=Anomalocephalus; =Clappaspis), Elrathia, Elrathiella (=Coelaspis; =Glossocoryphus), Eokaotaia, Erdoradites, Furia, Ganovexopyge (/Scottia), Huochengella, Inglefieldia, Jenkinsonia, Kailiella, Kaotaia, Katunicare, Kistocare, Langqia, Lenacare, Nelgakia, Parapachyaspis, Parehmania (=Mcnairia; =Rowia; =Thompsonaspis), Pedinocephalina, Peregrinaspis, Plesiamecephalus, Proehmaniella, Proveedoria, Pseudomexicella, Schopfaspis, Trachycheilus, Tympanuella, Utaspis. Antagmidae: Antagmella, Antagmus, Bagradia, Bicella, Bilimbataia, Cambrophatictor, Crassifimbra, Cyphambon, Erzishania (=Oreisator), Houmengia, Katunia, Lennontoviella, Longshania, Luaspides, Mantoushania, Onchocephalina, Onchocephalus (=Litocodia), Paraantagmus, Periomma, Plesioperiomma, Shilengshuia, Sombrerella, Wanbeiaspis, Xiangqianaspis, Xiaofangshangia, Xiaomajiella, Yuehsienszella. Asaphiscidae: Anomocarellius, Asaphiscus, Blainia, Blainiopsis, Blountiella, Blountina, Canotiana (=Williamsina), Cinnella, ?Conoides, Dunderburgella, Edithiella, Eoasaphiscus HAJRULLINA, Eokaninia (/Kaniniella SIVOV), Eoproetus, Erbenia, Eteraspis, Iniotoma, Kaninia (/Kaniniella SIVOV; =Dolgaia), Kaniniella KOBAYASHI, Lioparia (/Lorentzia, /Pseudoliostracina; =Emmrichella; =Liaoyangaspis), Luyanhaoaspis (/Luaspis PENG et al., 1995), Mindycrusta, Paraorlovia, Vega, Verkholenella. Atopidae: Atopina, Atops (=Ivshiniellus), ?Avalonia, Pseudatops. Bolaspididae: Acrocephalops, Bolaspidaspis, Bolaspis, Eldoradia, Rawlinsella. Cedariidae: Bonneterrina (=Holstonia; =Piedmontia), Carinamala, Cedaria, Cedarina, Henadoparia, Jimachongia, Vernaculina. Changshaniidae: Benxiella, Changshania (=Metachangshania; =Prochangshania), Changshanocephalus, Kazelia (=Kazellina), Mecophyrs, Narinosa, Parachangshania, Paramenomonia, Paraqingshuiheella (=Qingshuiheella), Pseudowentsuia, Suribongia, Wentsuia. Conocoryphidae: Bailiaspis, Bailiella (=Liaotungia; =Liocephalus; =Tangshihella), Cainatops (=Comucoryphe), Conocoryphe (/Conocephalites; =Conocephalus; =Couloumania), Ctenocephalus, Elyx (/Eryx), Hartella, Parabailiella, Tchaiaspis. Conokephalinidae: Buitella, Catuniella, Conokephalina (=Lobocephalina; =Ruzickaia/Lobocephalus), Gorskia, Maspakites, Meisteraspis, Meisterella, Miranda, Oirotella, Suludella, Westergaardella. Crepicephalidae: Bagongshania, Beikuangaspis, Cayupania, Coosella (=Wilsonella), Coosia, Coosina, Coosinoides, Crepicephalina (=Mesocrepicephalus), Crepicephalus, Hsuchuangia, Idioura, Kasatchaspis, Neimonggolaspis, Perforina, Pseudocrepicephalus, Sinocoosella, Sinocrepicephalus, Sneedvillia, Temnoura (=Asteromajia), Tetraceroura, Uncaspis, Zaozhuangaspis. Diceratocephalidae: Anopocodia, Aulacodigma, Cyclolorenzella, Diceratocephalus, Fenghuangella (=Cyclolorenzellina), Hwangjuella, Jiangnania, Tangshihlingia, Tholifrons (=Paraphoreotropsis), Torifera, Xiangia. Elviniidae: Chariocephalus, Dartonaspis, Drumaspis, Dunderbergia, Dytremacephalus, Elburgia, Elvinaspis, Elvinia (=Moosia), Elviniella, Elvinioides, Elyaspis, Enshia, Irvingella (=Irvingellina; =Parairvingella; =Komaspis), Jessievillia, Kujandina, Maladioides, Maladiopsis, Megadundabergia, Metisaspina, Onchopeltis, Paraenshia, Parakomaspis, Pesaia, Protemnites (=Prismenaspis), Pseudomaladioides, Pseudosaukia, Qingshuihella, Schmidtaspis, Yunlingia. Eulomidae: Acrocephalaspina, Altaiaspis, ?Amzasskiella (=Triplacephalus), Archaeuloma, Baikadamaspis, Bilacunaspis, Butyrinia, Crucicephalus, Dolgeuloma (/Rosovaspis/ Pseudoacrocephalites ROSOVA), Duplora, Euduplora, Euloma (=Calymenopsis), Guizhoucephalina, Iveria, Karataspis, Ketyna (=Kujandaspis), Lateuloma, Limpeina, Loparella, Lopeuloma, Luyanhaoia, Miaeuloma, ?Natmus, Pareulorna (=Gansucephalina), Pesaiina, Plecteuloma, Probilacunaspis, Proteuloma (=Mioeuloma), Pseudoacrocephalites MAKSIMOVA, Sanduspis, Spineuloma, Stigmatoa. Ignotogregatidae: Ignotogregatus. Inouyiidae: Catinouyia, Eoinouyia, Huainania, Inouyia, Parahuainania, Parainouyia, Parajialaopsis, Parawuania, Plesiowuania, Proinouyia, Pseudinouyia. Isocolidae: Cyphoniscus, Effnaspis, Hanzhongaspis, Holdenia (/Tiresias), Isocolus (/Astyages), Kielanella, Liangshanocephalus, Paratiresias, Pradesia, Pseudopetigurus, Taimyraspis, Thoralocolus, Triarthroides. Kingstoniidae: Acheilus CLARK, Ankoura, Blountia (=Homodictya; =Protillaenus; =Stenocombus), Brachyaspidion (/Brachyaspis MILLER, 1936), Bynumia, Bynumina, Calvipelta, Clelandia (/Harrisia; =Bynumiella), Ithycephalus, Kingstonella, Kingstonia (=Ucebia), Kingstonioides, Komaspidella (=Buttsina; =Ataktaspis), Larifugula, Maryvillia, Pugionicauda, Saonella, Shuizuia, Wanwanaspis, Wanwanoglobus, Yanzhuangia. Lisaniidae: Dazhuia, Eoshcngia (=Baojingia), Extrania, Klimaxocephalus, Lisania (=Aojia), Megalisania, Metalisania, Paralisaniella, Paraojia, Parashengia, Platylisania, Quandraspis, Redlichaspis (=Lisaniella), Rinella, Shengia, Xichuania. Llanoaspididae: Amquia, Arcuolimbus, Deiracephalus (/Asteraspis), Genevievella (=Placoscma; = Nixonella; =Torridella), Llanoaspidella, Llanoaspis, Metisaspis, Nahannicephalus, Paracedaria (/Pilgrimia), Rogersvillia, Sacha, Stenelymus, Tagenarella. Lonchocephalidae: Amiaspis, Bolaspidellus, Calymenidius, Caulaspina, Caulaspis, Durinia, Glaphyraspis (=Raaschella), Graciella, Hawkinsaspis (/Hawkinsia), Interalia, Kuraspis, Kuraspoides, Lazarenkiura, Letniites, Lonchocephalus (=Bucksella), Monosulcatina, Neoglaphyraspis, Nordia, Olegaspis, Prolonchocephalus, Pseudotalbotina, Quebecaspis, Raaschellina, Talbotina, Terranovella, Trymataspis, Weeksina. Lorenzellidae: Damiaoaspis, Eujinnania, Inouyops, Inoyellaspis, Jiangjunshania, Lonchinouyia, Lorenzella, Paralorenzangella (/Paralorenzella Q.Z. ZHANG), Paralorenzella LUO, Paraporilorenzella, Porilorenzella (=Jinnania), Pseudolorenzella, Ptyctolorenzella, Zhongweia. Mapaniidae: Angsiduoa, Hualongia, Mapania, Mapanopsis, Metanomocarella, Paramapania, Pseudomapania, Quitacetra, Quitalia. Marjumiidae: Anemocephalops, Crepichilella, Glyphopeltis, Holmdalia, Ithyektyphus, Lecanopleura, Loulania, Marjumia, Modocia (=Armonia; =Metisia; =Perioura; =Semnocephalus), Nasocephalus, Nericella, Nericia, Pearylandia, Petruninaspis, Schyilaspis, Shickshockia, Syspacheilus. Menomoniidae: Balderia, Biaverta, Bolaspidella (= Deissella; =Howellaspis), Bridgeia, Coenaspis, Coenaspoides, Deltophthalmus, Dresbachia, Hysteropleura (=Apedopyanus), Josina, Knechtelia, Menomonia (=Densonella/Millardia), Tavsenia, Verditerrina. Nepeidae: Ascionepea, Ferenepea, Folliceps, Loxonepea, Nepea, Penarosa (=Trinepea). Norwoodiidae: Cedaraspis, Garbiella, Hardyoides (=Norwoodina), Holcacephalus, Levisaspis, Norwoodella, Norwoodia (=Whitfieldina), Paranorwoodia, Xenocheilos. Papyriaspididae: Badainjaranaspis, Metapianaspis, Papyriaspis, Pianaspis, Prohedinaspis, Prohedinia (=Tosotychia), Sanduhedinaspis, Wandelella, Wangcunia, Wudangia. Phylacteridae: Aphelotoxon (=Ponumia), Cliffia, Drabia, Phylacterus (=Liostracinoides), Westonaspis. Proasaphiscidae: Acanthocephalus Alomataspis, Chelidonocephalus, Dananzhuangaspis, Daopingia, Deltocephalus, Derikaspis, Dignaceps, Eoasaphiscellus (/Eoasaphiscus LU & YUAN), Eymekops (=Kolpura), Farsia, Gangdeeria, Gloria, Grandioculus (/Megalophthalmus; =Honania), Guankouia, Hadraspis, Hatangia, Heukkyoella, Holanshania, Honanaspis, Hsiaoshia, Huaibeia, Hundwarella (=Anomocaraspis), Hunjiangaspis, Iranochresterius, Iranoleesia (/Irania; =Michaspis; =Heyelingella), Itcheriella, Jiangsuaspis, Jiangsucephalus, Jiangsuia, Jiubaspis, Jixianella (/Jixiania), Koptura (=Parakoptura), Kopungiella, Kutsingocephalus, Leichuangia, Lioparella (=Zhuozishania), Luliangshanaspis, Luonanocephalus, Manchuriella, Maotunia, Memmatella, Miaobanpoia, Paofeniellus, Paragangdeeria, Parazhongtiaoshanaspis, Plectrocrania, Plesigangdeeria, Proasaphiscina, Proasaphiscus, Pseudanomocarina, Pseudonericella, Raduginella, Severina, Shanxiella, Shuangshania, Stella, Sudanomocarina, Szeaspis (=Spitiaspis), Tankhella, Tengfengia, Teratokoptura, Tylotaspis, Ulania, Urjungaspis, Wutaishanaspis, Xiangshania, Xinglongia, Yanshania, Yujinia, Zhaishania, Zhongtiaoshanaspis. Ptychopariidae: Achlysopsis, Altikolia, Altitudella, Amecephalites, Asthenopsis, Austinvillia, Balangcunaspis, Bashania, Bathyholcus, Bathyocos, Billingsaspis, Binodaspis (=Xilingxia), Blairella, Bolaspidina, Brunswickia, Bulkuraspis, Caborcella, Callidaspina, Callidaspis, Cathayanella, Champlainia, Chengshanaspis, Chinghisicus, Chunghwaella, Conopolus, Dananzhuangia, Deltina, Douposiella (=Tongshania), Elrathina, Entsyna, Eodouposiella, Eokochaspis, Eosoptychoparia, Eospencia, Eurostina, Finecrestia, Gaotanaspis, Gaphuraspis, Gedongaspis, Gunnia (=Ellotia; =Yiliangaspis), Hadrocephalites, Hadrokraspedon, Hamptonella, Hejinaspis, Hemicricometopus, Hewenia, Holasaphus, Horbusonia, Illtydaspis, Jangudaspis, Jialaopsis, Jianchangia, Jimaoshania, Jiumenia, Kermanella, Kochaspis (/Palaeocrepicephalus), Kochiella (=Eiffelaspis), Kochiellina, Kochina, Kounamkites, Kunmingaspis (=Benxiaspis), Laminurus, Laoyingshania, Lianglangshania, Loriella, Luguoia, Luxella, Lyriaspis, Majiangia, Manailina, Meitania, Metisella, Mexicella, Monanocephalus, Mopanshania, Mrassina, Mufushania, Nangaocephalus, Nangaoia, Nangaops, Nanoqia, Nassovia, Nelsonia, Neokochina, Nyella, Olenekina, Onchocephalites, Ontoella, Orienturus, Orlovia, Orloviella, Pachyaspidella, Pachyaspis, Palmeraspis, Panacus, Paraeosoptychoparia, Paragunnia, Paramecephalus (=Parahiolites), Paraperiomma, Paraplagiura, Parapoulsenia, Parashuiyuella, Paraziboaspis, Perimetopus, Periommella, Piazella, Pingluaspis, Piochaspis, Plagiura (=Ruichengella; =Plagiurella), Pokrovskayaspis, Poulsenella, Poulsenia, Poulseniella, Probowmania, Probowmaniella (=Proshantungaspis), Probowmanops, Proliostracus, Promeitania, ?Protohedinia, Pseudoliostracus, Psilostracus, Ptychoparella (=Eoptychoparia; =Syspacephalus; =Elrathma), Ptychoparia (=Agraulopsis; =Ptychoparioides), Qiaotouaspis, Qingshuiheia, Reedus, Regina, Regius, Runnania, Salankanaspis, Sanhuangshania, Sanwania, Schistometopus, Semisphaerocephalus, Seriaspis, Shanganella, Shantungaspis, Shuiyuella, Sinoptychoparia, Spencella, Spencia (=Stauroholcus), Stoecklinia, Sujaraspis, Suluktella, Taijiangia, Taniaspidella, ?Townleyella, Trachyostracus, Trigonyangaspis (/Trigonaspis), Tukalandaspis, ?Ulrichaspis, Variopelta, Vermontella, Vica, Volocephalina, Wanhuaia, Weijiaspis, Wuhaina, Xiangshanaspis, Xingrenaspis (=Spitella; = Danzhaina; =Wuxunaspis), Yaoyiayuella, Yohoaspis, Yuknessaspis, Ziboaspidella, Ziboaspis. Shumardiidae: Acanthopleurella, Changchowilla, Conophrys, Elaphraella, Eoshumardia, Gaoloupingia, Koldinioidia (=Akoldinioidia), Kuruktagaspis, Kweichowilla, Leioshumardia, Leoforteyia, Liriamnica, Parashumardia, Puanocephalus, Shumardia, Shumardoella (/Shumardella), Shumardops, Stigmametopus, Tarimella (=Yinganaspis), Thomondia, Trianguraspis. Solenopleuridae: Abakanopleura, Acanthometopus, Acrocephalaspis, Aiaiaspis, Aidarella, Aikhaliella, Albansia, Aldanaspis, Badulesia, Bigranulella, Bijelina, Braintreella, Canotaspis, Catasolenopleura, Changqingia (=Austrosinia), Colliceps, Conicephalus, Crusoia, Daldynaspis, Datongites, Denaspis, Eilura, Ejinaspis, Erratojincella, Foveatella, Gonzaloia, Gushanaspis, Herse (/Solenopleurina; =Parasolenopleura WESTERGARD), Huzhuia, Hyperoparia, Jiagouia, Jincella, Kabuqiia, Kaipingella, Karagandoides, Keguqinia, Lashushania, Levisia, Lingyuanaspis, Liosolenopleura, Maiaspis, Manublesia, Markhaspis, Mataninella, Menocephalites (=Parataitzuia), Minupeltis, Mukrania, Munija, Neoacrocephalites, Neosolenopleurella, Nilegna, Ninaspis, Notocoryphe, Paracrocephalites (/Arctaspis), Paramenocephalites (=Solenoparina), Parasolenoparia, Parasolopleurena (/Parasolenopleura POLETAEVA), Parayabeia, Pardailhania, Perneraspis (/Perneria), Pingluia, Plesisolenoparia, Proavus, Protrachoparia, Pseudosolenoparia, Reillopleura, Rimouskia, Rina, Sao (=Acanthocnemis; =Acanthogramma, =Crithias; =Endogramma; =Enneacnemis; =Goniacanthus; =Micropyge; =Monadina; =Monadella; =Selenosema; =Staurogmus; =Tetracnemis), Shanghaiaspis, Sohopleura, Solenoparia, Solenoparops, Solenopleura, Solenopleurella, Solenopleuropsis, Squarrosoella, Suyougouia, Tabalqueia, Tangwangzhaia, Tjungiella, Trachoparia, Usumunaspis, Velieuxia, Wafangdiania, Xianfengia, Yabeia. Tricrepicephalidae: Erediaspis, Meteoraspis (=Greylockia; = Coleopachys), Prometeoraspis, Tricrepicephalus (/Paracrepicephalus). Utiidae: Amginouyia, Antagmopleura (=Poljakovia), Chondragrautina, Chondagraulos, Okunevaella, Utia. Wuaniidae: Eotaitzuia, Houmaia, Inouyella, Jixianaspis, Latilorenzella (=Wuania), Lophodesella, Megagraulos, Pseudosolenopleura, Rencunia, Ruichengaspis, Wuanoides, Wuhushania, Yunmengshania. |
. . Triarthrus |
Suborder Olenina
Cephalon: with narrow border, opisthoparian
sutures (exceptionally proparian), genal angle rounded
or spined, glabella tapering forward with simple,
sigmoid, or bifurcate furrows (furrows absent in some
forms); librigenae yoked (fused by a narrow strip of
cephalic doublure), or rarely, separated by anterior
median suture; eyes small to medium, usually with
distinct eye ridges. [Members of Olenina form the
foundation for the Order Olenida (Adrain 2011) and will
be extracted from Ptychopariida at this website in the
near future.] |
Cattilicephalidae
Pemphigaspis
|
Uncertain Suborder or Polyphyletic Families: Avoninidae: Avonina, Wongia. Catillicephalidae: Acheilops, Agelagma (?=Paradistazeris), Buttsia, Buttsiella, Catillicephala (/Cephalocoelia), Catillicephalites, Coephalocoeliaspis, Cryptoderaspis, Distazeris, Galeaspis, Lajishanaspis, Madarocephalus, Matania, Onchonotellus (=Onchonotina; =Guotangia; =Seletella), Onchonotopsis, Onchonotus, Pemphigaspis (=Hallaspis), Peracheilus (=Acheilus RAYMOND), Qilianshania, Stenochilina, Theodenisia (/Denisia; =Calculites; =Mannschreekia), Triarthropsis, Tumidulaspis, Tuojiangella, Urbanaspis, Waergangia, Welleraspis (=Avonaspis), Yukonaspis. Ityophoridae: Frognaspis, Ityophorus. Plethopeltidae: Arapahoia (=Hesperaspis), Exigua (=Brassicicephalus), Koldinia, Koldiniella, Kuljumbina, Lampropeltis, Leiocoryphe, Meniscocoryphe, Notaiella, Plethopeltides, Plethopeltis (=Plethometopus; =Enontioura), Plethopeltoides (=Kulyumbopeltis), Poriplethopeltis, Pseudokoldinella, Semicyclocephalus, Stenopilus, Strotocephala, Tolstotchichaspis. ADDITIONAL CLASSIFICATION NOTES FOR PTYCHOPARIIDA: This order is certainly most problematic. Defined largely by combinations of primitive character states (such as tapering glabella, preglabellar field, natant hypostome, small, furrowed pygidium, and subdued exoskeletal sculturing), the fate of the Ptychopariida may be to slowly dwindle as groups such as the Proetida, Asaphida, and Harpida are defined via their shared derived characters and extracted from the Ptychopariida clade. The origins of the Ptychopariida and other Early Cambrian trilobite lineages have been explored by Jell (2003) who noted that at least two major lineages arose independently in different parts of the Early Cambrian world out of Fallotaspididae (one of the earliest trilobite families, a member of Order Redlichiida) to produce taxa that are now referred to as members of Ellipsocephaloidea (generally recognized as the earliest of the Ptychopariida). More recently (e.g, Hollingsworth 2008), members of the ellipsocephaloid family Bigotinidae (Serrania and Hupetina) actually appear in strata lower and earlier than fallotaspidoids in Morocco and Spain! In addition, Jell 2003 suggests that another lineage gives rise to Anomocaridae (primitive members of Order Asaphida) and Proasaphiscidae (primitive Ptychoparioidea). This would make the Ptychopariida polyphyletic, and throws a complicaton into the origins of the Asaphida. Jell also questions the primacy of the conterminant hypostomal state, suggesting that the earliest trilobites were natant, and that both facial sutures and the conterminant condition arose early and independently in several lines. The "Librostome Orders" (Ptychopariida, Asaphida, Proetida, Harpida, and possibly the Phacopida), have sister taxa within the Ptychopariida, making it a paraphyletic taxon. The problem is exacerbated by the inadequate diagnoses of the constituent families; even the well-accepted clades within Ptychopariida, such as the family Olenidae, were united by few shared derived characters (but see Adrain 2012, below), and, as in the Asaphida, there are primitive representatives lacking some of the derived states, and advanced representatives that have secondarily lost them. The process of clarifying the Ptychopariida is hampered by lack of complete specimens, lack of sufficient ontogenetic data, lack of hypostome and other ventral characters, etc. It will take much effort and some fortuitous breakthroughs in both data and analysis to deal with the Ptychopariida. There are a number of families and genera that are for the present considered part of the Ptychopariida, but for which additional detail is difficult to ascertain. This by definition makes the Ptychopariida a "garbage heap" taxon, an inelegant status. Adrain (2012) supported splitting of Order Olenida out of Ptychopariida based on the following: "Secondarily silicified trilobites yield new data on ventral structures and developmental morphologies, and have revealed the surprising presence of a complex articulation of the librigenal anterior projection and the cranidium in several groups of trilobites, many of which have not been considered related to one another. We propose the term "stylidion" for this structure. It involves a series of tiny pillars running dorsally from the inner edge of the librigenal doublure and articulating in a series of small pits on the inner (ventral) surface of the cranidium. These pits are often expressed on the dorsal (external) surface of the cranidium as tiny tubercles. The pillars are matched on the outer (ventral) surface of the librigenal doublure by another series of small pits. This morphology has not previously been documented. Most trilobites have a simple inner doublural margin and no cranidial modifications in this region. We hypothesize that the stylidion is an unreversed synapomorphy of the recently proposed Order Olenida. While available data are scant, all firmly associated larvae and early growth stages of olenide species have a characteristic morphology involving transversely complete glabellar furrows (most trilobites have the furrows expressed only abaxially), anteriorly placed eye ridges, and tuberculate fixigenae. These will likely prove to be additional synapomorphies of the order, supplementing the stylidion, which is developed later in ontogeny." A page for the newly established Olenida is forthcoming. The genera listed here are from Jell & Adrain 2003.
Adrain, J.M. 2012. Toward a phylogenetic classification of the Trilobita: Synapomorphies and structure of the Order Olenida Adrain 2011. Presentation at Geological Society of America 2012. GSA Abstracts Vol 44 (7): 373. Fortey, R. A. Trilobite systematics: the last 75 years. J. of Paleontology. 75(6):1141-51. Hollingsworth, J.S., 2008. The first trilobites in Laurentia and elsewhere. In: I.Rábano, R. Gozalo and D. García-Bellido (Eds.), Advances in trilobite research. Cuadernos del Museo Geominero, nº 9. Instituto Geológico y Jell, P.A. 2003. Phylogeny of Early Cambrina trilobites. Special Papers in Palaeontology 70:45-57. Jell, P.A. & J.M. Adrain. 2003 Available generic names for trilobites. Memoirs of the Queensland Museum 48(2):331-553 |
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Ptychopariida Gallery
A small gallery of images from the web