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ORDER ASAPHIDA last revised 09 June 2006 by S. M. Gon III Diagnostic features shown in this color Introduction: A large (comprising perhaps 20% of trilobite species) and morphologically diverse order of trilobites, most advanced families united by similar ontogeny (the ovoid, effaced "asaphoid" protaspid form with enrolled doublure); most members also have a median ventral suture (only secondarily lost via fusion in two advanced families). Cephalon: often equal/subequal to pygidium (e.g., Asaphoidea), but some not so (e.g., Trinucleioidea); usually with a high degree of cephalic effacement so glabellar furrows are faint or not visible; eyes usually large (some forms secondarily blind); preoccipital glabellar tubercle in advanced forms; cephalic doublure often wide, with terrace ridges; librigena are typically separated by a median ventral suture; dorsal anterior facial sutures often curve adaxially to meet in front of the glabella; sutures opisthoparian; hypostome conterminent or impendent, with only primitive forms (e.g., the Anomocaroidea) natant. Thorax: typically 5 – 12 segments, but 2 - 3 in a few Trinucleioidea, 13+ in some Anomocaroidea, up to 30 in an Alsataspidid (Trinucleioidea). Pygidium: typically large (subisopygous to macropygous), with a wide doublure. Occurrence: Middle-Upper Cambrian boundary to upper Ordovician-lower Silurian. Suborders: None (or nominate Asaphina). Superfamilies: Anomocaroidea, Asaphoidea, Dikelocephaloidea, Remopleuridoidea, Cyclopygoidea, Trinucleioidea. |
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Superfamily Anomocaroidea
Introduction: Primitive Asaphida (possibly ancestral to some
other asaphine groups), including families retaining the natant hypostomal
condition, as well as other ptychoparioid features; protaspides resembling those of Ptychopariida (not
"asaphoid"); not all families included may be monophyletic (i.e.,
Anomocaroidea as described here is likely a paraphyletic group). Anomocarellidae: Anomocarella (=Psilaspis; = Entorachis), Eoanomocarella, Fissanomocarella, Glyphaspis (=Americare), Hanshania, Huayuania, Liocare, Liopeishania, Liopelta, Luia, Lydiaspis, Megalopsis, Neoanomocarella, Orthodorsum, Paranomocarella, Peishania, (=Parapeishania), Peishanoides, Plebiellus. Anomocaridae: Abharella, Afghanocare, Amginia, Anomocare, Anomocarina, Anomocarioides, Anomocariopsis, Callaspis, Chondranomocare, Dilatalimbus, Elandaspis, Eocatuniella, Forchammeria, Formosocephalus, Fuquania, Glyphanellus, Glyphaspellus, Guizhouanomocare, Hanivella, Harataspis, Hunanaspis, Igarkiella, lohomia, Irinia, Jimanomocare, Juraspis, Kokuria, Kolbinella, Kotuia, Leichneyella, Lomsucaspis, Longxumenia, Macrotoxus, Metanomocare, Nadiyella, Palella, Paracoosia (=Manchurocephalus), Parakotuia, Paranomocare, Pjatkovaspellus, Qinlingia, Rectifrontinella, Sachaspis, Schoriecare, Schoriella, Scintilla, Sivovella, Usovinurus, Wutingshania, Yongwolia. Aphelaspididae: Amorphella, Aphelaspidella, Aphelaspis (=Proaulacopleura; =Clevelandella; =Labiostria), Apheloides, Dicanthopyge, Elegantaspis, Erixanium, Eugonocare, Kobayashella, Listroa, Litocephalus, Maduiya, Nganasanella (=Tamaranella), Notoaphelaspis, Olenaspella, Olentella, Paraphelaspis, Pseudaphelaspis, Pseudeugonocare, Taenicephalites, Taenora. Parabolinoididae: Abdulinaspis, Apornodocia, Boestrupia, Croixana, Jasmundia, Kendallina (/Kendallia), Minkella, Orygmaspis, Parabolinoides (=Bernia), Pedinocephalus, Pesaiella, Roksaspis, Stigmacephaloides, Taenicephalops, Taemcephalus (=Bemaspis; =Maustonia), Weishania. Pterocephaliidae: ?Anecocephalus, Beigongia, Bromella, Camaraspis, Cernuolimbus, Cilia, Dikelocephalioides, Dikelocephalites, Dingxiangaspis, Flabellocephalus, Guangxiaspis, Housia (=Housiella), Jubileia, Labiostrella, Labiostrina (=Abia), Longlingaspis, Lunacephalus, Luotuolingia, Morosa, Parahousia, Paramaladioidella, Pauciella, Pedinaspis, Pedinocephalites, Pelicephalus, Petalocephalus, Plesiocilia, Prehousia, Proapatokephaloides, Prodikelocephalites, Pterocephalia (/Pterocephalus; =Hederacauda), Pterocephalina, Pterocephalopsinus (/Pterocephalops LIN & ZHANG in ZHU et al., 1979), Pulchricapitus (=Reaganaspis), Qilianaspis, Sigmocheilus, Stenambon, Strigambitus, Tiantouzhania, Tumicephalus, Uxunella, Xiaoshiella, Yingziaspis, Yokusenia, Yushugouia, Zhenania, Zhuangliella. |
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Superfamily Asaphoidea
Introduction: As in typical Asaphida with the following additional
defining characteristics: Ceratopygidae: ?Ajacicrepida, Asiocephalus, Boschchekulia, Cataplotaspis, Ceratopyge, Cermatops, Charchaqia (=Aplotaspis), Diceratopyge (=Paraceratopyge), Dichelepyge (=Bicornipyge), Dipleuropyge, Guozia, Haniwoides (=Yuepingia), Hedinaspis, Hunanopyge, Hysterolenus (=Ruapyge/ Hectoria), Kaltykelina, Kaufmannella (/Kaufmannia), Kogenium, Lopnorites, Macropyge (=Haniwapyge, =Lichapyge; =Macropygella), Mansuyella, Nannopeltis, Neohedinaspis, Onychopyge (=Prionopyge), Proceratopyge, Promacropyge (=Aksapyge), Pseudohysterolenus, Pseudoyuepingia (=Iwayaspis; =Sayramaspis), Sinoproceratopyge, Tamdaspis (=Psiloyuepingia), Tropidopyge, Wannania, Xiaodaositunia. |
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Trinucleioidea
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Superfamily Trinucleioidea
Cephalon: Opisthoparian or marginal facial sutures, generally
eyeless; glabella typically convex and pyriform,
with 3 or fewer pairs of furrows, preoccipital glabellar tubercle sometimes
present; usually long genal spines. Dionididae: Aethedionide, Digrypos, Dionide (/Dione; /Polytomurus; =Dionidepyga; =Trigrypos), Dionideina, Dionidella, Huangnigangia, Paradionide, Tongxinaspis, Trinucleoides. Liostracinidae: Aplexura, Doremataspis, Liostracina, Lynaspis. Raphiophoridae: Ampyx (/Brachyampyx), Ampyxella, Ampyxina, Ampyxinella, Ampyxoides, Anisonotella, Bulbaspis, Caganaspis, Carinocranium, Cerampyx, Cnemidopyge, Collis, Edmundsonia, Ellsaspis, Endymionia (/Endymion), Globampyx, Jiuxiella (=Miboshania), Kanlingia, Lonchodomas, Malinaspis, Malongullia (=Ampyxinops), Mendolaspis, Metalonchodomas, Miaopopsis, Nambeetella, Nanshanaspis, Parabulbaspis, Parampyx, Pseudampyxina, Pytine, Raphioampyx, Raphiophorus, Raymondella (/Reedaspis), Rhombampyx, Salteria, Sinampyxina, Sinoluia, Taklamakania (=Xinjiangia). Trinucleidae: Anebolithus, Australomyttonia, Bancroftolithus, Bergamia (=Bohemaspis; =Brandysops; =Cochliorrhoe), Bettonolithus, Botrioides, Broeggerolithus (=Ulricholithus), Costonia, Cryptolithoides, Cryptolithus, Deanaspis, Declivolithus, Decordinaspis, Eirelithus, Famatinolithus, Furcalithus, Guandacolithus, Gymnostomix, Hanchungolithus (=Ichangolithus; =Yinjiangolithus), Huenickenolithus, Incaia, Jianxilithus, Lloydolithus, Lordshillia, Marekolithus, Marrolithoides, Marrolithus, ?Microdiscus, Myinda, Myindella, Myttonia, Nankinolithus, Ningkianolithus (=Cerato1ithus; =Hexianolithus), Novaspis, Onnia, Paratretaspis, Paratrinucleus, Parkesolithus, Pragolithus, Protoincaia, Protolloydolithus, Reedolithus, Reuscholithus, Salterolithus (=Smeathenia), Stapeleyelta, Telaeomarrolithus, Tetrapsetlium, Tretaspis, Trinucleus (/Edgellia), Whittardolithus, Xiushuilithus, Yinpanolithus. |
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Superfamily Dikelocephaloidea
Cephalon: with opisthoparian sutures, glabella typically truncate anteriorly and squat,
1-4 pairs of lateral furrows, 1p may be transglabellar; preglabellar field
variable, sometimes absent, palpebral ridge typically
well-defined, but separate from axial furrow (compare to sister group
Remopleuroidea); genal spines typically present, of various length; median
ventral suture rarely lost to secondary fusion; hypostome conterminant Eurekiidae: Bandalaspis, Bayfieldia, Corbinia, Eurekia, Leocephalus, Lochmanaspis, Magnacephalus, Maladia, Tostonia. Ptychaspididae: Alborsella, Changia (=Coreanocephalus; =Quadraticephalus; =Fengshania), Eoptychaspis, Eowuhuia, Euptychaspis, Idiomesus, Kathleenella, Keithia, Keithiella, Macronoda (=Promesus), Plectrella, Proricephalus, Ptychaspis (=Asioptychaspis), Saukioides (/Pseudosaukia; /Jeholaspis), Sunwaptia, Wilcoxaspis. Saukiidae: Anderssonella (/Anderssonia, /Sunina), Calvinella, Caznaia, Danzhaisaukia, Diemanosaukia, Eosaukia (=Scolosaukia), Galerosaukia, Guangnania, Hamashania, Lichengia, Linguisaukia, Liquania, Lophoholcus, Lophosaukia, Mareda, Metacalvinella, Mictosaukia, Mictosaukioidia, Paracalvinella, Pileaspis, Platysaukia, Prosaukia (=Stenosaukia), Pseudocalvinella, Saukia, Saukiella, Sinosaukia, Stigmaspis, Tellerina, Thailandium, Wedekindiaspis (/Wedekindia). |
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Superfamily Cyclopygoidea
Cephalon: with glabella expanding forward
to anterior margin, effaced in later cyclopygids, may be fused with
occipital ring; fixigenae reduced (except in primitive Taihungshaniidae),
palpebral lobes lack distinct rims, and contact
axial furrows at anterior ends, librigenae fused or separated by
anterior median suture; hypostome relatively transverse,
impendent, often with tripartite posterior margin; eye various sized
(may be hypertrophied and convex (Cyclopyge), typically closely adjoined
to glabella. Nileidae: Aocaspis, Barrandia, Berkutaspis, Borthaspidella, Bumastides, Elongatanileus, Homalopteon, Illaenopsis (=Eurymetopus; =Procephalops; =Rokycania/Pseudobarrandia), Kodymaspis, ?Lakaspis, Neopsilocephalina, Nileus (=Remopleuridioides), Parabarrandia, Parabumastides, Paranileus, ?Peraspis, Petrbokia, Platypeltoides (/Platypeltis), Poronileus, Psilocephalinella (/Psilocephalus/ Psilocephalina/Borthaspis), Shenjiawania, Symphyroxochus, ?Symphysurina (=Symphysurinella; =Symphysuroides), Symphysurus, Troedssonia, Varvia. Taihungshaniidae: Asaphellina, Asaphopsis, Omeipsis, Pacootella, Renhuaia, Taihungshania (=Miquelina), Tungtzuella. |
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Superfamily Remopleuridoidea
Cephalon: with opisthoparian sutures, glabella bulges tranversely anterior of occipital ring,
with up to 3 pairs lateral furrows, eyes medium to very large, with narrow, wire-like socle, palpebral
rims inflated, deep rim furrows, extending into axial furrows anteriorly;
genal spines present. Idahoiidae: Aguilarella, Arrhenaspis, Brabbia, Comanchia, Duibianaspis, Elviraspis, Langyashania, Lauzonella, Leviscila, Loganellus (=Highgatea), Maladioidella (=Kuruktagella; =Cedarellus), Noelaspis, Patronaspis, Psalaspis, Pyttstrigis, Saratogia (=Idahoia, =Meeria), Shitaia, Valtoressia, Wafangia, Wilbernia, Zhuitunia. Remopleurididae: Aktugaiella, Amphitryon (/Caphyra; =Brachypleura), Aotiaspis, Apatokephalina, Apatokephaloides, Apatokephalops (=Aristokainella; =Wanliangtingia), Apatokephalus, Apiflabellum, Arator, Artokephalus, Atratebia, Auricula, Binervus, Blosyropsis, Cavia, Deanokephalus, Diplapatokephalus, Dislobosaspis, Eoapatokephalus, Eorobergia, Haniwa, Hastiremopleurides, Hexacopyge, Hualongella, Hukasawaia, Hypodicranotus, Ivshinaspis, Jiia, Jingheella, Jinshaella, Kainella, Kainellina, Kainelloides, Lacorsalina, Lingukainella, Lohanpopsis, Loshanella, Lulongia, Makbelaspis, Mendosina, Menoparia, Naustia, Oculeus, Poletaevia, Portentosus, Praepatokephalus, Proapatokephalops, Pseudokainella (=Elkanaspis; =Parakainella; =Fatocephalus), Pugilator, Remopleurella, Remopleurides, Remopleuridiella, Richardsonaspis, Richardsonella (=Lakella; =Protapatokephalus), Robergia, Robergiella, Scinocephalus, Sculptaspis, Sculptella, Sigmakainella, Spinacephalus, Taishania, Teratorhynchus, Tibikephalus, Tramoria, Yosimuraspis (=Eoyosimuraspis; =Metayosimuraspis). |
| ADDITIONAL CLASSIFICATION NOTES FOR ASAPHIDA:
Fortey & Chatterton (1988) provided an extensive summary of evidence uniting the clades within the suborder Asaphina, and later, arguments were presented by Fortey (1990) for the recognition of an emmended Order Asaphida, as described here. The assumption is that the ventral median suture evolved only one time, thereby uniting all members bearing that feature as a monophyletic group. As a stem group composed of primitive asaphid families (such as Anomocaridae), the Anomocarioidea is paraphyletic, giving rise to the derived superfamilies, and presumably including the most primitive transitional taxa for each (such as some Auritamiidae). The ontogeny of the advanced members of the Asaphida (e.g., Asaphoidea, Trinucleioidea, Cyclopygoidea) are consistent, with an effaced, globular protaspis. For more details on ontogeny see Rudy Lerosey's page: "Trilobite larvae & orders: Asaphida" Whittington (2003) argues that at least some of the Nileidae (e.g., Lakaspis, Peraspis, and Symphysurina; see Cyclopygoidea above) should not be placed in that family, and that most nileids should be considered Illaenina (Corynexochida). If so, then the argument of the ventral median suture as a central synapomorphy for the Asaphida might require reconsideration. Expanded and complete genera listings for the families above are from Jell & Adrain (2003). . To view a chart of relationships within the Asaphida, click here (warning: the image is large!) Fortey, R. A. 1990. Ontogeny, hypostome attachment, and trilobite classification. J. of Paleontology. 33:529-76. Fortey, R. A. 2001. Trilobite systematics: the last 75 years. J. of Paleontology. 75(6):1141-51. Fortey, R. A. and B. D. E. Chatterton. 1988. Classification of the Trilobite Suborder Asaphina. Palaeontology 31(1):165-222 Jell, P.A. & J.M. Adrain. 2003. Available generic names for trilobites. Memoirs of the Queensland Museum 48(2):331-553. Whittington, H.B. 2003. The trilobite family Nileidae: Morphology and Classification. Palaeontology 46(4):635-46. |
