Part E, Porifera (Revised), vol. 4 & 5, ch. 16, p. 1043–1106

Full text

PART E, REVISED, VOLUME 4, CHAPTER 19:
SYSTEMATIC DESCRIPTIONS: ARCHAEOCYATHA
F. Debrenne,1 A. Yu. ZhurAvlev,2 and P. D. Kruse3
[113 rue du Long Foin, 91700, Ste Geneviève des Bois, France, francoise.debrenne@gmail.com; 2Área y Museo de Paleontología, Departamento
de Ciencias de la Tierra, Facultad de Ciencias, Universidad de Zaragoza, Spain, ayzhur@mail.ru; 3South Australian Museum, Adelaide, Australia,
archaeo.kruse@gmail.com]
Phylum PORIFERA Grant, 1836
Class ARCHAEOCYATHA
Bornemann, 1884
[nom. correct. vologDin, 1937b, p. 464, pro Archaeocyathinae borne-
mAnn, 1884, p. 706] [=class Archaeocyathinae TAYlor, 1910, p. 105; class
Cyathospongia oKuliTch, 1935, p. 88; class Archaeocyathi R. beDForD
& W. r. beDForD, 1936, p. 9; subphylum Archaeocyatha vologDin,
1937b, p. 464 (Porifera); class Pleospongia oKuliTch, 1943, p. 1; phylum
Archaeocyatha oKuliTch, 1955a, p. 8; phylum Archaeocyathi KrAsnoPe-
evA, 1955, p. 17; subphylum Euarchaeocyatha ZhurAvlevA, 1960b, p. 79,
nom. transl. ZhurAvlev & others in soKolov & ZhurAvlevA, 1983, p. 6,
ex class Euarchaeocyathi ZhurAvlevA, 1960b, p. 79; Salpingidea volog-
Din & YAZmir, 1967, p. 1377; phylum Archaeocyatha hill, 1972, p. 2;
Euarchaeocyatha ZhurAvlevA & mYAgKovA, 1979, p. 521] [equivalent to
superfamily Archaeocyathaceae simon, 1939, p. 5]
Skeleton nonspiculate, calcareous cup
of microgranular microstructure and (with
few exceptions) original magnesium calcite
composition. Cup generally of archaeocya-
than architecture with one or two porous
walls bounding inner or central cavity
respectively; porous to aporose septa, pseu-
dosepta, taeniae, pseudotaeniae, pseudotae-
nial network, dictyonal network, syringes
and/or tabulae may form in intervallum;
a minority are of chaetetid (intervallum
with calicles) or thalamid architecture (cup
consisting of successive chambers). Solitary
or modular. Secondary calcareous skeleton
may be present. [For an explanation of
zonal terms used herein, see Debrenne,
Zhu rAvl ev , & Kru se, 2012, p. 64–65.]
Cambrian (Terreneuvian–Furongian).
Order MONOCYATHIDA
Okulitch, 1935
[nom. correct. oKuliTch, 1955a, p. 9, pro order Monocyathina oKuliTch,
1935, p. 90] [=Archaeolynthida Zhu rAvlevA, 1957, p. 174; =Tectocy-
athida vologDin in vologDin & YAZmir, 1966, p. 948; =suborder Glo-
bosocyathina oKunevA, 1969, p. 74; =suborder Monocyathina Debrenne,
1970a, p. 24; =Tecticyathida vologDin, 1977, p. 93]
Cup one walled, solitary or low modular;
pelta may be present. lower Cambrian
(Tom.1–Bot.3).
Family MONOCYATHIDAE
R. Bedford & W. R. Bedford, 1934
[Monocyathidae R. beDFo rD & W. R. beDForD, 1934, p. 2] [=Rhab-
docnemidae oKuliTch, 1943, p. 45, nom. nov. pro Rhabdocyathidae
vologDin, 1931, p. 52, invalid family-group name based on junior hom-
onym; =Archaeolynthidae ZhurAvlevA, 1949, p. 550; =Monocyathinae
ZhurAvlevA, 1963b, p. 74, nom. transl. ex Monocyathidae R. beDForD &
W. r. beDForD, 1934, p. 2; =Rhabdocyathellidae ZhurAvlevA, 1963b, p.
114; =Capsolynthidae oKunevA, 1969, p. 75; =Crassicyathidae vologDin,
1977, p. 79; =Spinicyathidae vologDin, 1977, p. 103]
Wall with simple pores. lower Cambrian
(Tom.1–Bot.3).
Archaeolynthus TAYl or , 1910, p. 158 [*Monocy-
athus porosus R. beDForD & W. R. beDForD,
1934, p. 2; SD R. beD Fo rD & W. R. beDFor D,
1936, p. 20; lectotype, R. beDForD & W. r.
beDF orD , 1934, fig. 1; Zhur Avle vA, 1963b, fig.
39d; hill, 1965, pl. 2,1; Debrenne , 1969a, pl.
1,3; Debrenne, 1974b, pl. 19,1; SD hill, 1965,
p. 52, NHM S4140, London] [=Ventriculocyathus
vologDin, 1928, p. 31, nom. nud.; =Ventriculo-
cyathus vologDin, 1931, p. 51 (type, V. caulius,
M), for discussion, see hill, 1965, p. 63; =Mono-
cyathus R. beDForD & W. r. beDForD, 1934,
p. 2 (type, M. porosus, SD R. beDForD & W. R.
beD Fo rD , 1936, p. 20; lectotype, hill, 1965,
pl. 2,1; SD hill, 1965, p. 52, S4140, NHM,
London); =Rhabdocnema oKuliTch, 1937a, p.
252, nom. nov. pro Rhabdocyathus v on Toll,
1899, p. 45, non brooK, 1893, cnidarian (type, R.
sibiricus, M), for discussion, see hill, 1965, p. 51;
=Rhabdocyathella vologDin, 1937b, p. 474 (type,
R. lebedevae, M), for discussion, see hill, 1965,
p. 53; =Capsolynthus osADchAYA in ZhurAvlevA &
others, 1967, p. 26 (type, C. helenae, OD); =Corti-
cicyathus vologDin, 1977, p. 46 (type, C. aequi-
porosus, OD); =Crassicyathus vologDin, 1977, p.
79 (type, C. canaliculatus, OD); =Tegminicyathus
vologDin, 1977, p. 98 (type, T. simplex, OD);
=Tytthocyathus vologDin, 1977, p. 98 (type, T.
jenisseicus, OD); =Spinicyathus vologDin, 1977,
p. 103 (type, S. cipis, OD), for discussion, see
Debrenne, ZhurAvlev, & roZ A no v, 1989, p.
94; Debrenne, roZA nov, & Zhu rAvlev , 1990, p.
133]. Wall pores of uniform size. lower Cambrian
(Tom.1–Bot.3): Siberian Platform, Altay Sayan,
Tuva, Mongolia, Transbaikalia, Far East, Australia,
Antarctica, Morocco, Iberia.——Fig. 1,1a–b. *A.
porosus (R. beDForD & W. r. beDForD), Ajax
Limestone, Botoman, Ajax Mine, South Australia,
© 2012, The University of Kansas, Paleontological Institute, ISSN 2153-4012
Debrenne, F., A. Yu. Zhuravlev, & P. D. Kruse. 2012. Part E, Revised, Volume 4, Chapter 19: Systematic
descriptions: Archaeocyatha. Treatise Online XX:1–186, 134 fig.
Ed: add vol.
number

2Treatise Online, number XX
Australia, lectotype, NHM S4140; a, external
longitudinal view of cup, ×5 (Hill, 1965); b,
detail of porosity in external tangential view, ×10
(Debrenne, 1974b).
Kyarocyathus Kruse, 1982, p. 144 [*K. duplus; OD;
holotype, Krus e, 1982, fig. 7H–J, AM FT.8240,
8244, Sydney]. Wall pores of two distinct sizes. lower
Cambrian (Bot.1–Bot.2): Mongolia, Australia.——
Fig . 1,2. *K. duplus, Mount Wright Volcanics,
Botoman, Mt. Wright, New South Wales, Australia,
AM FT.8240, FT.8244, oblique transverse section,
AM FT.8240, ×10 (Kruse, 1982).
Family PALAEOCONULARIIDAE
Chudinova, 1959
[Palaeoconulariidae chuDi novA, 1959, p. 53] [=Debrennecyathidae
vologDin in vologDin & YAZmir, 1966, p. 948, invalid family-group
name based on unavailable genus name; =Debrennecyathidae vologDin,
1977, p. 100]
Wall with attached microporous sheath.
lower Cambrian (Atd.4–Bot.3).
Palaeoconularia chuDinovA, 1959, p. 53 [*P. prima;
OD; holotype, chuDinovA, 1959, fig. 1–2, PIN
1577/1, Moscow] [=Laminaecyathus YAZmir in
vologDin & YAZmir, 1966, p. 948 (type, L. trian-
gulatus, OD); =Debrennecyathus vologDin in
volo gD in & YAZ mi r, 1966, p. 948, nom. nud.,
unavailable genus-group name without associated
nominal species; =Debrennecyathus vologDin,
1977, p. 101 (type, D. pulcher, OD), for discussion,
see Debrenne, Zhur Avlev, & roZAn ov, 1989, p.
122; Debrenne, roZAnov, & ZhurAvlev, 1990, p.
154]. Wall with reticulate carcass pores and attached
microporous sheath. lower Cambrian (Bot.1–Bot.3):
Altay Sayan, Tuva, Mongolia, Transbaikalia.——
Fig. 2,1a–c. *P. prima, Verkhnemonok Formation,
Botoman, Karako l River, western Sayan, Al tay
Sayan, Russia, holotype, PIN 1577/1; a, external
longitudinal view of cup, ×2; b, tangential section
of microporous sheath, ×6; c, tangential section of
carcass pores, ×11 (Chudinova, 1959).
?Butakovicyathus ZhurAvlevA, 1980, p. 175 [*B .
butakovi; OD; holotype, Zhu rAvl evA, 1980, pl.
Archaeolynthus
Kyarocyathus
1a
1b
2
Fig. 1. Monocyathidae (p. 1–2).
Ed: add vol.
number

Systematic Descriptions of the Archaeocyatha 3
Fig. 2. Palaeoconulariidae (p. 2–4).
1a
2
1b
1c
Palaeoconularia
Butakovicyathus

4Treatise Online, number XX
30,1–2, TsSGM 569, Novosibirsk]. Wall carcass
pores of two distinct sizes with attached micro-
porous sheath. [The single available section does
not provide certainty as to wall structure.] lower
Cambrian (Atd.4–Bot.1): Altay Sayan.——Fig.
2,2. *B. butakovi, Krol Formation, Atdabanian,
Mana River, eastern Sayan, Altay Sayan, Russia,
holotype, TsSGM 569, transverse section, ×12
(Zhuravleva, 1980).
Family TUMULIOLYNTHIDAE
Rozanov, 1966
[Tumuliolynthidae roZAnov in roZAnov & missArZhevsKiY, 1966, p. 77]
[=Papulicyathidae vologDin, 1977, p. 62; =Verrucicyathidae vologDin,
1977, p. 63; =Orthocyathidae vologDin, 1977, p. 96]
Wall with simple tumuli. lower Cambrian
(Tom.2–Bot.3).
Tumuliolynthus ZhurAvlevA, 1963b, p. 101 [*Rhab-
docyathus tubexternus vologDin, 1932, p. 64; OD;
holotype, vologDin, 1932, pl. 5,1a, TsNIGRm
209a/2957, St. Petersburg] [=Papulicyathus
vologDin, 1977, p. 62 (type, P. longus, OD);
=Verrucicyathus vologDin, 1977, p. 64 (type,
V. tumefactus, OD), for discussion, see Kru se ,
1982, p. 144; =Isthmocyathus vologDin, 1977,
p. 70 (type, I. articulatus, OD); =Mammaticy-
athus vologDin, 1977, p. 71 (type, M. kyzasicus,
OD ); =Ort h ocya t hus vologDin, 197 7, p. 97
(type, O. bateniensis, OD), non Cyathophyllum
(Orthocyathus) merriAm, 1974, p. 34 (type,
Prismatophyllum flexum sTumm, 1938, p. 483,
OD), cnidarian; for discussion, see Debrenne,
roZAnov, & ZhurAvle v, 1990, p. 165]. Wall with
simple tumuli. lower Cambrian (Tom.2–Bot.3):
Siberian Platform, Altay Sayan, Tuva, Mongolia,
Transbaikalia, Far East, Urals, Australia, Antarc-
tica, Morocco, Iberia.——Fi g. 3,1. *T. tubex-
ternus (vologDin), Verkhneynyrga Formation,
Botoman, Lebed’ River, Altay Mountains, Altay
Sayan, Russia, holotype, TsNIGRm 209a/2957,
transverse section, ×3 (Vologdin, 1932).
Family SAJANOLYNTHIDAE
Rozanov, 1989
[Sajanolynthidae roZAnov in Debrenne, ZhurAvlev, & roZAnov, 1989,
p. 79] [=Sajanolynthidae KAshinA in roZAnov, 1973, p. 85, nom. nud.]
Wall with multiperforate tumuli. lower
Cambrian (Bot.1).
Sajanolynthus vologDin & KAshinA, 1972, p. 152 [*S.
desideratus; OD; holotype, vologDin & KAshinA,
1972, pl. 20,1, KGU 19/1, Krasnoyarsk] [=Pustu-
licyathus vologDin, 1977, p. 94 (type, P. tectus,
OD), for discussion, see Debrenne, Zhur Avlev, &
roZAnov , 1989, p. 129; Debrenne, roZA nov, &
ZhurAvlev, 1990, p. 159]. Wall with multiperfo-
rate tumuli. lower Cambrian (Bot.1): Altay Sayan,
Far East.——Fig. 3,2. *S. desideratus, Torgashino
Formation, Botoman, Uyar River, eastern Sayan,
Altay Sayan, Russia, holotype, KGU 19/1, transverse
section, ×15 (Vologdin & Kashina, 1972).
Family GLOBOSOCYATHIDAE
Okuneva, 1969
[Globosocyathidae oKunevA, 1969, p. 75] [=Propriolynthidae roZAnov,
1973, p. 85, nom. nud.; =Tumuloglobosidae roZAnov, 1973, p. 85, nom.
nud.; =?Tecticyathidae vologDin, 1977, p. 93]
Wall with bracts or scales. lower Cambrian
(Atd.2–Bot.1).
Propriolynthus oKunevA, 1967, p. 133 [*Archaeolyn-
thus vologdini YAKovlev, 1956, p. 855; OD; lecto-
type, YAKovlev, 1956, pl. 1,1, SD oKunevA, 1967,
p. 133, not located] [=Globosocyathus oK une vA,
1969, p. 75 (type, G. bellus, OD); =Tumuloglobosus
oKunevA in oKunevA & rePinA, 1973, p. 93 (type,
T. crassus, OD); =Subiculicyathus vologDin, 1977,
p. 49 (type, Archaeolynthus vologdini YA Kov le v,
1956, p. 855, OD); =?Tecticyathus vologDin,
1977, p. 93 (type, Archaeolynthus peltathus mAslov,
1961, p. 121, OD); =Propricyathus oK une vA in
vologDin, 1977, p. 100 (type, P. maritimus,
OD), for discussion, see Debrenne, ZhurAvlev,
& ro ZAnov , 1989, p. 125; Debrenne, roZAn ov,
& Zh ur Avl ev , 1990, p. 156]. Wall with pores
bearing downwardly projecting, cupped bracts.
lower Cambrian (Atd.2–Bot.1): Siberian Platform,
Altay Sayan, Tuva, Mongolia, Far East.——Fig.
3,3a –b. *P. vo log din i (YA K ov le v ), Dmi trievka
Formation, Botoman, Spassk-Chernigovka area,
Far East, Russia, specimen PGU 30-X2; a, oblique
longitudinal section, ×3; b, tangential section, ×15
(Okuneva, 1967).
Melkanicyathus belYAevA, 1969, p. 88 [*M. limitatus;
OD; holotype, belYAevA, 1969, pl. 38,2–3, DVGU
212/5, Khabarovsk] [=Phymatocyathus vologDin,
1977, p. 72 (type, P. orillatus, OD); =Scyphocy-
athus vologDin, 1977, p. 96 (type, Rhabdocnema
operculatum mAsl ov , 1960, p. 1117, OD), for
discussion, see Debrenne, ZhurAvlev, & roZAnov,
1989, p. 118; Debrenne, roZAnov, & ZhurAvlev,
1990, p. 151]. Wall with pores bearing upwardly
projecting, cupped bracts. lower Cambrian (Bot.1):
Altay Sayan, Far East.——Fi g. 3,4a–b. *M. limi-
tatus, Ust’toka unit, Botoman, Bol’shoy Mel’kan
River, Dzhagdy Range, Far East, Russia, holotype,
DVGU 212/5; a, oblique transverse section, ×6; b,
longitudinal section, ×6 (Belyaeva, 1969).
Family FAVILYNTHIDAE
Debrenne, 1989
[Favilynthidae Debrenne in Debrenne, ZhurAvlev, & roZ Anov, 1989,
p. 79] [=Favilynthidae Debrenne, 1974b, p. 98, nom. nud.]
Wall with canals. lower Cambrian (Atd.1–
Bot.3).
Favilynthus Debrenne in Zhur Avl evA , 1974a, p.
138 [*Monocyathus mellifer R. beDForD & W. R.
beDForD, 1936, p. 12; OD; holotype, R. beDForD &

Systematic Descriptions of the Archaeocyatha 5
Fig. 3. Tumuliolynthidae, Sajanolynthidae, Globosocyathidae, and Favilynthidae (p. 4–6).
Sajanolynthus
2
3a
3b
Melkanicyathus
4a
4b
Favilynthus
5b
5a
Tumuliolynthus
1
6a
6b
Robertiolynthus
Propriolynthus

6Treatise Online, number XX
W. R. beDForD, 1936, fig. 49; ZhurAvlevA, 1963b,
fig. 42; Debrenne, 1974b, pl. 19,5–6, SAM P932-
47, Adelaide]. Wall with horizontal to upwardly
projecting, straight canals. lower Cambrian (Atd.1–
Bot.3): Altay Sayan, Tuva, Mongolia, Far East,
Australia, Antarctica.——Fig. 3,5a–b. *F. mellifer
(R. beDFo rD & W. r. beDFo rD), Ajax Limestone,
Botoman, Ajax Mine, South Australia, Australia; a,
external transverse view, holotype, SAM P932-47,
×6; b, external longitudinal view, topotype, USNM
PU9, ×6 (Debrenne, 1974b).
Robertiolynthus Zhu r Av l e v in voro n ovA and
others, 1987, p. 19 [*R. handfieldi; OD; holotype,
voronovA & others, 1987, pl. 1,1, GSC 90116,
Ottawa] [=Veolynthus boYArinov & KonYAevA
in ZhurAvlevA & others, 1997a, p. 26 (type, V.
jucundus, OD)]. Wall with horizontal to upwardly
projecting, straight canals bearing supplementary
bracts externally. lower Cambrian (Bot.1–Bot.2):
Altay Sayan, Canada.——Fig. 3,6a–b. *R. hand-
fieldi Zhu rAv le v, Se kwi Forma tion, Botoma n,
Mackenzie Mountains, Northwest Territories,
Canada; a, holotype, GSC 90116, longitudinal
section, ×19; b, paratype, GSC 90117, transverse
section, ×19 (Voronova & others, 1987).
Order AJACICYATHIDA
R. Bedford & J. Bedford, 1939
[nom. correct. oKuliTch, 1955a, p. 10, pro order Ajacicyathina R. beDForD
& J. beDForD, 1939, p. 70] [=?order Somphocyathina oKuliTch, 1943,
p. 47, nom. nud.; =Somphocyathida oKuliTch, 1955a, p. 19; =Nocho-
roicyathida ZhurAvlevA in vologDin, 1956, p. 879; =Dokidocyathida
vologDin, 1957a, p. 178; =Bronchocyathida ZhurAvlevA in vologDin,
1957a, p. 180; =Bosceculida KrAsnoPeevA, 1960, p. 41; =Ethmophyllida
vologDin, 1961, p. 178; =Cyclocyathellida vologDin, 1961, p. 179]
Cup two walled, solitary or low modular;
inner wall of centripetal type of devel-
opment; intervallum with septa, with or
without plate tabulae. lower Cambrian
(Tom.1–Toy.3).
Suborder DOKIDOCYATHINA
Vologdin, 1957
[nom. transl. ZhurAvlevA, 1960b, p. 95, ex order Dokidocyathida volog-
Din, 1957a, p. 178]
Intervallum with septa bearing single
longitudinal pore row. lower Cambri an
(Tom.2–Bot.3).
Superfamily
DOKIDOCYATHOIDEA
R. Bedford & W. R. Bedford, 1936
[nom. correct. Debrenne, ZhurAvlev, & roZAnov, 1989, p. 80, pro Doki-
docyathacea Debrenne, 1970a, p. 24, nom. transl. ex Dokidocyathidae R.
beDForD & W. R. beDForD, 1936, p. 12]
Outer wall with simple pores. lowe r
Cambrian (Tom.2–Bot.3).
Family DOKIDOCYATHIDAE
R. Bedford & W. R. Bedford, 1936
[Dokidocyathidae R. beDForD & W. R. beDForD, 1936, p. 12]
Inner wall with simple pores. l o wer
Cambrian (Tom.2–Bot.3).
Dokidocyathus TAYlo r, 1910, p. 146 [*D. simpli-
cissimus; M; lectotype, TAYl o r, 1910, pl. 16,
photos 91–92; Zhu r Av le vA , Ko nY u sh K ov , &
roZA nov , 1964, pl. 4,2–3; ro ZA nov , 1973, pl.
14,1; Debrenne, 1974b, pl. 20,3; SD Debrenne,
1970a, p. 33, SAM T1589A-B, cups F-G, Adelaide]
[=?Velicyathus Debrenne, 1964, p. 125 (type, V.
levillaini, OD); =Dokidolynthus Debrenne, 1974b,
p. 101 (type, Dokidocyathus lenaicus ro ZAn ov in
ZhurAvlevA, KonYushKov, & roZAnov, 1964, p.
83, OD); =?Kamyshovaecyathus YAZmir in Zhu rAv-
le vA, 1974a, p. 183, nom. nud. based on type
species not then available; =?Kamyshovaecyathus
YAZmir in YAZmir, DA lm ATov, & YAZmir, 1975,
p. 38 (type, K. immanis, OD), for discussion, see
Debrenne, ZhurAvlev, & roZA no v, 1989, p.
104; Debrenne, roZAnov, & Zhur Avlev , 1990,
p. 141]. Outer and inner walls with simple pores.
lower Cambrian (Tom.2–Bot.3): Siberian Plat-
form, Altay Sayan, Tuva, Mongolia, Transbaikalia,
Far East, Urals, Australia, Antarctica, Morocco,
Iberia.——Fig. 4,1. *D. simplicissimus, Ajax Lime-
stone, Botoman, Ajax Mine, South Australia,
Australia, lectotype, SAM T1589A-B, cups F-G,
oblique transverse view, ×2.5 (Taylor, 1910).
Family DOKIDOCYATHELLIDAE
Debrenne, 1964
[Dokidocyathellidae Debrenne, 1964, p. 112]
Inner wall with bracts or scales. lower
Cambrian (Atd.2–Bot.1).
Dokidocyathella ZhurAvlevA, 1960b, p. 100 [*D. incog-
nita; OD; holotype, ZhurAvlevA, 1960b, fig. 73, pl.
5,3, TsSGM 205/8, Novosibirsk]. Inner wall with
pores bearing upwardly projecting, S-shaped scales.
lower Cambrian (Atd.2–Bot.1): Siberian Platform,
Altay Sayan, Tuva, Far East.——Fig. 4,2. *D. incog-
nita, Pestrotsvet Formation, Atdabanian, Oy-Muran,
Lena River, Sakha (Yakutia), Russia, holotype, TsSGM
205/8, transverse section, ×10 (Zhuravleva, 1960b).
Incurvocyathus roZ An ov in roZ An ov & missAr-
ZhevsKiY, 1966, p. 50 [*I. voronovae; OD; holotype,
roZAnov & missArZhevsKiY, 1966, pl. 1,4–5, PIN
4597/57, Moscow]. Cup with regular transverse
folds affecting both walls; inner wall with pores
bearing possibly upwardly projecting, S-shaped
scales. lower Cambrian (Atd.2–Bot.1): Altay Sayan,
Tuva. ——Fi g. 4, 3a–b. *I . voronovae , Shangan
Formation, Botoman, eastern Tannu-Ola Range,
Tuva, Russia, holotype, PIN 4597/57; a, longi-
tudinal section (outer wall to left), ×8; b, detail
of longitudinal section (outer wall to left), ×8
(Rozanov & Missarzhevskiy, 1966).
Dokidocy-
athella: I
changed this
occurrence
of Yakutia
to Sakha
(Yakutia),
okay?

Systematic Descriptions of the Archaeocyatha 7
Dokidocyathus Dokidocyathella
Incurvocyathus
Cordobicyathus
12
3a
3b
4a
4b 4c
Fig. 4. Dokidocyathidae, Dokidocyathellidae, and Cordobicyathidae (p. 6–8).

8Treatise Online, number XX
Family CORDOBICYATHIDAE
Perejón, 1975
[Cordobicyathidae Perejón, 1975a, p. 136]
Inner wall with annuli. lower Cambrian
(Atd.2).
Cordobicyathus Perejón, 1975a, p. 136 [*C .
deserti; OD; holotype, Perejón, 1975a,
pl. 3,1–3, CE 3-74-2, Madrid]. Inner wall
with upwardly projecting, S-shaped annuli.
lower Cambrian (Atd.2): Iberia, Germany,
Poland.——Fig. 4,4a–c. *C. deserti, Pedroche
Formation, Atdabanian, Las Ermitas, Cordoba,
Andalusia, Spain, holotype, CE 3-74-2; a,
transverse section, ×6.5 (Perejón, 1975a); b,
oblique section, ×6.5; c, detail of longitudinal
section (outer wall to right), ×15 (Debrenne,
Zhuravlev, & Kruse, 2002).
Superfamily
KIDRJASOCYATHOIDEA
Rozanov, 1964
[nom. transl. Debrenne, ZhurAvlev, & roZAnov, 1989, p. 80, ex Kidrja-
socyathidae roZAnov in ZhurAvlevA, KonYushKov, & roZAnov, 1964, p.
95] [=Kidrjasocyathacea roZAnov, 1973, p. 85, nom. nud.]
Outer wall with independent micropo-
rous sheath. lower Cambrian (Atd.2–Bot.1).
Family KIDRJASOCYATHIDAE
Rozanov, 1964
[Kidrjasocyathidae roZAnov in ZhurAvlevA, KonYushKov, & roZAnov,
1964, p. 95]
Inner wall with simple pores. l o wer
Cambrian (Atd.2–Bot.1).
Kidrjasocyathus roZAnov, 1960b, p. 43 [*K. uralensis;
OD; holotype, roZAnov, 1960b, fig. 1, pl. 1,1a–b;
ZhurAvlevA, KonYushKov, & ro ZA no v, 1964,
pl. 10,3, PIN 4297/9, Moscow]. Inner wall with
simple pores. lower Cambrian (Atd.2–Bot.1): Altay
Sayan, Urals.——Fig. 5a–c. *K. uralensis, Terekla
Formation, Botoman, Kidryassovo, western flank of
Southern Urals, Russia, holotype, PIN 4297/9; a,
transverse section, ×9; b, detail of transverse section
(outer wall to right), ×20 (Debrenne, Zhuravlev, &
Kruse, 2002); c, sketch of transverse section, ×20
(Rozanov, 1960b).
Superfamily
KALTATOCYATHOIDEA
Rozanov, 1964
[nom. transl. Debrenne, ZhurAvlev, & roZAnov, 1989, p. 80, ex Kalta-
tocyathidae roZAnov in ZhurAvlevA, KonYushKov, & roZAnov, 1 964,
p. 92] [=Kaltatocyathacea roZAnov, 1973, p. 85, nom. nud.]
Outer wall with simple tumuli. lower
Cambrian (Atd.1–Bot.1).
Family KALTATOCYATHIDAE
Rozanov, 1964
[Kaltatocyathidae roZAnov in ZhurAvlevA, KonYushKov, & roZAnov,
1964, p. 92]
Inner wall with simple pores. l o wer
Cambrian (Atd.1–Bot.1).
Kaltatocyathus roZ A no v in ZhurA vl evA , KonY-
ushKov, & roZ Anov, 1964, p. 92 [*K. kaschinae;
OD; holotype, Zhu r Av le vA , Kon Y us h Ko v , &
roZAn ov, 1964, pl. 9,7, PIN 4297/47, Moscow]
[=Aroonacyathus grAve sToc K, 1984, p. 46 (type,
A. gregarius; OD), for discussion, see Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 114; Debrenne,
roZAnov, & Zh urAvlev, 1990, p. 147]. Inner wall
with simple pores. lower Cambrian (Atd.1–Bot.1):
Altay Sayan, Transbaikalia, Far East, Australia.——
Fig. 6. *K. kaschinae, Bazaikha Formation, Atdaba-
nian, Bazaikha River, eastern Sayan, Altay Sayan,
Russia, holotype, PIN 4297/47, transverse section,
×19 (Zhuravleva, Konyushkov, & Rozanov, 1964).
Superfamily
PAPILLOCYATHOIDEA
Rozanov, 1989
[Papillocyathoidea roZAn ov i n Debrenne, Zhu rAvlev , & roZAnov,
1989, p. 80] [=Papillocyathacea roZAnov, 1973, p. 85, nom. nud.].
Outer wall with multiperforate tumuli.
lower Cambrian (Atd.4–Bot.1).
Family PAPILLOCYATHIDAE
Rozanov, 1989
[Papillocyathidae roZAnov in Debrenne, ZhurAvlev, & roZAnov, 1989,
p. 80] [=Papillocyathidae roZAnov, 1973, p. 85, nom. nud.]
Inner wall with simple pores. l o wer
Cambrian (Atd.4–Bot.1).
Papillocyathus roZ An ov in ZhurAvlevA, KonY-
ushKov, & ro ZAnov, 1964, p. 94 [*P. vacuus; OD;
holotype, ZhurAvlevA, KonY ushKov, & roZ Anov,
1964, pl. 10,1–2, PIN 4297/48-2, Moscow]. Inner
wall with simple pores. lower Cambrian (Atd.4–
Bot.1): Altay Sayan.——Fig . 7a–b. *P. vacuus,
Balakhtinson Formation, Botoman, Kazyr River,
eastern Sayan, Altay Sayan, Russia, holotype, PIN
4297/48-2; a, transverse section, ×20; b, longitu-
dinal section (outer wall to right), ×20 (Zhurav-
leva, Konyushkov, & Rozanov, 1964).
Superfamily SOANICYATHOIDEA
Rozanov, 1964
[nom. transl. Debrenne, ZhurAvlev, & roZAnov, 1989, p. 80, ex Soani-
cyathidae roZAnov in ZhurAvlevA, KonYushKov, & roZAnov, 1964, p.
97] [=Soanicyathacea roZAnov, 1973, p. 85, nom. nud.]
Outer wall with bracts or scales. lower
Cambrian (Atd.2–Bot.1).

Systematic Descriptions of the Archaeocyatha 9
Family SOANICYATHIDAE
Rozanov, 1964
[Soanicyathidae roZAnov in ZhurAvlevA, KonYushKov, & roZ Anov,
1964, p. 97]
Inner wall with bracts or scales. lower
Cambrian (Atd.2–Bot.1).
Subtilocyathus vologDin, 1960, p. 422 [*Archaeo-
cyathus subtilis vologDin, 1932, p. 41; OD;
lectotype, vologDin, 1932, fig. 32a–b, pl.
7,7, pl. 8,5b, SD Debrenne, Zhu r Av l e v , &
Kru s e, 2002, p. 1557, TsNIGRm 50a/2957,
St . Pe tersb urg] [ = Soanicyathus ro Z A n o v in
ZhurAvlevA, KonYushKov, & ro ZA nov , 1964,
p. 98 (type, S. admirandus, OD), for discussion,
see Debrenne, Zhur Avl ev , & roZ An ov, 1989,
p. 133; Debrenne, roZ A n ov , & ZhurAvlev,
1990, p. 162]. Outer and inner walls with
pores bearing upwardly projecting, cupped
bracts. lower Cambrian (Atd.2–Bot.1): Altay
Sayan, Tuva, Mongolia.——Fig . 8,1a–b. *S.
subtilis (vo lo gD in ), Verkhneynyrga Formation,
Botoman, Lebed’ River, Altay Mountains, Altay
Sayan, Russia, lectotype, TsNIGRm 50a/2957;
a, transverse section, ×4; b, detail of transverse
section, ×20 (Vologdin, 1932).
Batschykicyathus ZhurAvlev i n ZhurAvlev,
ZhurAvlevA, & Fonin, 1983, p. 23 [*B. angu-
losus; OD; holotype, Zh u r A v l e v , Zh u r Av -
le vA, & Fonin, 1983, pl. 3,4, PIN 3848/501,
Moscow]. Outer wall regularly bulging in trans-
verse files; outer and inner walls with pores
bearing upwardly projecting, cupped bracts.
lower Cambrian (Atd.4): Siberian Platform.——
Fig . 8,2. *B. angulosus, Pestrotsvet Formation,
Atdabanian, Bachyk Creek, Lena River, Sakha
(Yakutia), Russia, holotype, PIN 3848/501,
oblique longitudinal section, ×10 (Zhuravlev,
Zhuravleva, & Fonin, 1983).
Family ZHURAVLEVAECYATHIDAE
Rozanov, 1989
[Zhuravlevaecyathidae roZAnov in Debrenne, ZhurAvlev, & roZAnov,
1989, p. 81] [=Zhuravlevaecyathidae roZAnov, 1973, p. 85, nom. nud.]
Inner wall with annuli. lower Cambrian
(Bot.1).
Zhuravlevaecyathus roZAnov in ZhurAvlevA, KonY -
ushKov, & roZAnov, 1964, p. 98 [*Z. pulchellus; OD;
holotype, ZhurAvlevA, KonYushKov, & roZAnov ,
1964, pl. 11,5–6, PIN 4297/54, Moscow]. Outer
wall with pores bearing upwardly projecting, cupped
bracts; inner wall with possibly upwardly projecting,
S-shaped annuli. lower Cambrian (Bot.1): Altay
Sayan.——Fig. 8,3a–b. *Z. pulchellus, Verkhnemonok
Formation, Botoman, Abakan River, western Sayan,
Altay Sayan, Russia; a, holotype, PIN 4297/54, trans-
verse section, ×4; b, paratype, PIN 4297/55, detail of
transverse section, ×3.5 (Zhuravleva, Konyushkov, &
Rozanov, 1964).
Superfamily
KYMBECYATHOIDEA Debrenne,
Rozanov, & Zhuravlev, 1989
[Kymbecyathoidea Debrenne, ro ZAnov , & ZhurAvlev i n Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 81]
Outer wall with canals. lower Cambrian
(Atd.4–Bot.3).
Family KYMBECYATHIDAE
Debrenne, Rozanov, & Zhuravlev, 1989
[Kymbecyathidae Debrenne, r oZAnov, & ZhurAvlev in Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 81]
Inner wall with simple pores. low e r
Cambrian (Atd.4–Bot.3).
Fig. 5. Kidrjasocyathidae (p. 8).
a
b
c
Kidrjasocyathus

10 Treatise Online, number XX
Kymbecyathus Debrenne & Kruse, 1986, p. 241 [*K.
avius; OD; holotype, Debrenne & Kru se, 1986,
fig. 6A–B, VU VC9, Wellington]. Outer wall with
horizontal to upwardly projecting, straight canals;
inner wall with simple pores. lower Cambrian
(Atd.4–Bot.3): Australia, Antarctica.——Fig. 9a–b.
*K. avius, Shackleton Limestone, Botoman, Crack-
ling Cwm, Byrd Glacier, Antarctica, holotype,
VU VC9; a, transverse section, ×3 (Debrenne &
Kruse, 1986); b, oblique longitudinal section, ×3
(Debrenne, Zhuravlev, & Kruse, 2002).
Suborder AJACICYATHINA
R. Bedford & J. Bedford, 1939
[nom. transl. ZhurAvlevA, 1960b, p. 106, ex order Ajacicyathina R.
beDForD & J. beDForD, 1939, p. 70] [=Nochoroicyathina ZhurAvlevA
in vologDin, 1956, p. 879, nom. transl. ZhurAvlevA, 1960b, p. 198, ex
Nochoroicyathida ZhurAvlevA in vologDin, 1956, p. 879; =Schiderty-
cyathina KrAsn oPeevA, 1969, p. 63; =Boscekulcyathina KrA snoPe evA,
1969, p. 63]
Intervallum with septa; pectinate tabulae
or synapticulae may be present. l o wer
Cambrian (Tom.1–Toy.3).
Superfamily
BRONCHOCYATHOIDEA
R. Bedford & J. Bedford, 1936
[nom. transl. ZhurAvlev in voronovA & others, 1987, p. 20, ex Broncho-
cyathidae r. beDForD & j. beDForD, 1936, p. 25] [=Ajacicyathoidea r.
beDForD & J. beDForD, 1939, p. 73, nom. correct. Debrenne & Kruse,
1986, p. 242, pro Ajacicyathacea ZhurAvlevA, 1960b, p. 106, nom. transl. ex
Ajacicyathidae r. beDForD & j. beDForD, 1939, p. 73; =Nochoroicyathacea
ZhurAvlevA in vologDin, 1956, p. 879, nom. transl. ZhurAvlevA, 1960b, p.
198, ex Nochoroicyathidae ZhurAvlevA in vologDin, 1956, p. 879; =Aldano-
cyathacea ZADoroZhnAYA, osADchAYA, & rePinA, 1973, p. 129, nom. transl.
Korshunov, 1983a, p. 96, ex Aldanocyathinae ZADoroZhnAYA, osADchAYA,
& rePinA, 1973, p. 129; =Irinacyathacea ZhurAvlevA in Debrenne, 1972, p.
174, nom. neg.; =Irinaecyathacea ZhurAvlevA in ZhurAvlevA & elKinA, 1974,
p. 44; =Inessocyathacea ZhurAvlevA in ZhurAvlevA & elKinA, 1974, p. 45]
Outer wall with simple pores. lowe r
Cambrian (Tom.1–Toy.2).
Family AJACICYATHIDAE
R. Bedford & J. Bedford, 1939
[Ajacicyathidae R. beDForD & J . beDForD, 1939, p. 73] [=Nochoroicy-
athidae ZhurAvlevA in vologDin, 1956, p. 879; =Kisasacyathidae KonY-
ushKov, 1972, p. 137; =Aldanocyathidae ZADoroZhnAYA, osADchAYA, &
rePinA, 1973, p. 129]
Inner wall with simple pores. l o wer
Cambrian (Tom.1–Toy.2).
Ajacicyathus R. beDFo rD & J. beDFo rD, 1939, p. 73
[*Archaeocyathus ajax TAYlor, 1910, p. 118; OD;
lectotype, TAYlo r, 1910, pl. 1, photo 1a, pl. 7,
photo 39 (below); hill, 1965, pl. 1,6; Debrenne,
1974b, pl. 21,2–4; SD De br e nn e , 1970a, p.
27, SAM T1550A, Adelaide] [=Loculicyathellus
Debrenne, 1969a, p. 310 (type, Archaeocyathus
floreus R. beD Fo rD & W. r. be DFo rD , 1934, p.
2, OD), nom. transl. Debrenne, 1974b, p. 115, ex
Loculicyathus (Loculicyathellus) Debrenne, 1969a, p.
310; =Ajacicyathus (Juricyathus) Debrenne, 1974b,
p. 110 (type, Archaeocyathus aequitriens R. beDForD
& j. beDForD, 1937, p. 35, OD); =Ambistapis
Kruse, 1982, p. 161 (type, A. integer, OD)]. Inner
wall with several rows of simple pores per inter-
sept; stirrup pores may be present; septa aporose
to sparsely porous. lower Cambrian (Atd.1–Toy.2):
Siberian Platform, Altay Sayan, Tuva, Mongolia,
Far East, Australia, Antarctica, Sardinia, France,
Canada.——Fig. 10,1a–b. *A. ajax (TAYlor), Ajax
Limestone, Botoman, Ajax Mine, South Australia,
Fig. 6. Kaltatocyathidae (p. 8).
Fig. 7. Papillocyathidae (p. 8).
Kaltatocyathus
Papillocyathus
a
b

Systematic Descriptions of the Archaeocyatha 11
Australia, lectotype, SAM T1550A; a, oblique
longitudinal view, ×1 (Taylor, 1910); b, detail of
septum and inner wall in longitudinal view (outer
wall to right), ×10 (Debrenne, 1974b).
Davidicyathus Debrenne, roZA nov, & ZhurAvlev,
1990, p. 139 [*?Loculicyathus racemiferus grAve-
sTocK, 1984, p. 48; OD; holotype, grAvesTocK,
1984, fig. 32H, 32L, SAM P21452, Adelaide].
Outer wall with two sizes of pores, the smaller
either isolated or clustered over the larger; inner
wall with several rows of simple pores per inter-
sept; septa completely porous. lower Cambrian
(Atd.4): Australia.——Fig. 10,2a–c. *D. racemiferus
(grAvesTocK); a–b, Wilkawillina Limestone, Atda-
banian, Wilkawillina Gorge, South Australia,
Australia, holotype, SAM P21452, a, transverse
section, ×4; b, longitudinal section, ×9.5 (Grave-
stock, 1984); c, Ajax Limestone, Atdabanian,
Mount Scott Range, South Australia, Australia,
SAM P21455-1, tangential section of outer wall,
×15 (M. Debrenne, new).
Dentatocyathus oKun evA, 1972, p. 57 [*D. mari-
timus; OD; holotype, oK une vA, 1972, pl. 10,7,
PGU 202, Khabarovsk]. Outer wall longitudinally
plicate; inner wall with several rows of simple
pores per intersept; septa completely porous. lower
Cambrian (Bot.1): Altay Sayan, Tuva, Mongolia, Far
East.——Fig. 11,1a–b. *D. maritimus, Dmitrievka
Fig. 8. Soanicyathidae and Zhuravlevaecyathidae (p. 9).
1a
1b
3a
3b
2
Subtilocyathus
Batschykicyathus
Zhuravlevaecyathus

12 Treatise Online, number XX
Formation, Botoman, Knorring Hill, Spassk-
Chernigovka area, Far East, Russia, holotype, PGU
202; a, transverse section, ×5; b, detail of transverse
section (outer wall to right), ×10 (Okuneva, 1972).
Iljinicyathus ZhurAvl evA, 1972b, p. 155 [*I. ulan-
batoriensis; OD; holotype, ZhurAvlevA, 1972b,
pl. 21,1–3, TsSGM 755/1, Novosibirsk]. Cup
in which inner wall shows periodic transverse
folds; inner wall with several rows of simple pores
per intersept; septa completely porous; pectinate
tabulae may be present. lower Cambrian (Atd.2–
Atd.4): Mongolia.——Fig. 11,2a–b. *I. ulanbato-
riensis, formation not known, Atdabanian, northern
Mongolia, holotype, TsSGM 755/1; a, transverse
section, ×4; b, longitudinal section (outer wall to
right), ×5 (Zhuravleva, 1972b).
Kisasacyathus KonYushKov, 1972, p. 137 [*K. micro-
tumulatus; OD; holotype, KonYushKov, 1972, pl.
16,1, PIN 4755/7, Moscow] [=Prethmophyllum
Debrenne, 1974c, p. 174 (type, Archaeocyathus
subacutus R. beDForD & W. R. beDForD, 1934,
p. 2, OD)]. Inner wall with one row of simple
pores per intersept, formed by fluting of inner
edges of septa; septa aporose to sparsely porous.
lower Cambrian (Atd.4–Bot.3): Altay Sayan, Tuva,
Mongolia, Far East, Australia, Antarctica.——Fig.
11,3a–b. *K. microtumulatus, Verkhnemonok Forma-
tion, Botoman, Kizas River, western Sayan, Altay
Sayan, Russia; a, holotype, PIN 4755/7, oblique
longitudinal section, ×6; b, paratype, PIN 4755/8,
transverse section, ×6 (Konyushkov, 1972).
Nochoroicyathus Zhur Av le vA , 1951, p. 78 [*N.
mirabilis; OD; holotype, ZhurAvlevA, 1951, fig.
1a–b, PIN 1168, Moscow, not located] [=Ajacicy-
athellus Debrenne, 1958, p. 64 (type, A. hollardi,
M); =Ascocyathus vologDin, 1960, p. 422 (type,
Archaeocyathus arteintervallum vologDin, 1931,
p. 84, OD); =Howellicyathus vologDin, 1961, p.
180, nom. nud.; =Howellicyathus vologDin, 1962a,
p. 126 (type, Coscinocyathus howelli vologDin,
1940b, p. 88, OD); =Pachecocyathus Perejón,
1971, p. 81 (type, P. cabanasi, OD); =Aldanocy-
athus voronin in Debrenne & voronin, 1971,
p. 30 (type, Ajacicyathus sunnaginicus ZhurAvlevA,
1960b, p. 115, OD), for discussion, see Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 120; Debrenne,
roZ An ov , & Zh ur Avl ev , 1990, p. 153]. Inner
wall with several rows of simple pores per inter-
sept; septa completely porous; pectinate tabulae
may be present. lower Cambrian (Tom.1–Bot.3):
Siberian Platform, Kolyma, Altay Sayan, Tuva,
Mongolia, Transbaikalia, Far East, Urals, Kazakh-
stan, Tarim, Australia, Morocco, Iberia, France,
Sardinia, Germany, Serbia.——Fig. 11,4a–c. *N.
mirabilis; a, Pestrotsvet Formation, Tommotian,
Nokhoroy Creek, Lena River, Sakha (Yakutia),
Russia, holotype, PIN 1168, transverse section,
×6 (Zhuravleva, 1951); b–c, Medvezh’ya Forma-
tion, Tommotian, Kotuy River, Krasnoyarsk region,
Russia; b, TsSGM 205/87, detail of septum in
longitudinal section (outer wall to left), ×6; c,
specimen TsSGM 205/88, detail of transverse
section at inner wall, ×20 (Debrenne, Zhuravlev,
& Kruse, 2002).
Orbiasterocyathus ZhurAvle vA in rePin A & others,
1964, p. 183 [*O. geri; OD; holotype, reP inA &
others, 1964, pl. 11,6, TsSGM 4272/5, Novosi-
birsk]. Cup in which both walls are longitudinally
folded, resulting in stellate transverse section;
inner wall with several rows of simple pores per
intersept; septa completely porous. lower Cambrian
(Atd.3–Atd.4): Altay Sayan.——Fi g. 11,5. *O .
geri, Adiak Formation, Atdabanian, Terensu River,
Gornaya Shoria, Altay Sayan, Russia, holotype,
TsSGM 4272/5, transverse section, ×4 (Repina &
others, 1964).
Fig. 9. Kymbecyathidae (p. 10).
a
b
Kymbecyathus

Systematic Descriptions of the Archaeocyatha 13
Fig. 10. Ajacicyathidae (p. 10–11).
Orbicyathellus osADchAYA in ZADoroZh nAYA, osAD-
chAYA, & rePin A, 1973, p. 133 [*O. bogradi; OD;
holotype, ZADoroZhnAYA, osADchAYA, & rePinA,
1973, pl. 19,1–2, TsSGM 424/1, Novosibirsk].
Cup in which both walls show periodic, synchro-
nous transverse folds; inner wall with stirrup
pores only; septa aporose to sparsely porous. lower
Cambrian (Atd.1–Atd.4): Siberian Platform, Altay
Sayan, Mongolia.——Fig. 12,1a–b. *O. bogradi,
Usa Formation, Atdabanian, Bograd, Batenev
Range, Kuznetsk Alatau, Russia, holotype, TsSGM
424/1; a, oblique longitudinal section, ×4.5; b,
tangential section of inner wall, ×12 (Zadorozh-
naya, Osadchaya, & Repina, 1973).
Orbicyathus vologDin, 1937b, p. 468 [*O. mongo-
licus; M; holotype, vologDin, 1937b, pl. 2,4,
1a
1b Ajacicyathus
2c
2a
Davidicyathus
2b
you asked me
to transpose
the labels
for views 2a
and 2b, but
according to
the mockup,
2b and 2c
needed to
be switched.
Okay now?

14 Treatise Online, number XX
Fig. 11. Ajacicyathidae (p. 11–16).
4c
1a
Iljinicyathus
2b
2a
1b
Dentatocyathus
4a
4b
3a
3b
Kisasacyathus
Nochoroicyathus
5
Orbiasterocyathus
Orbicyathus
6b
6a

Systematic Descriptions of the Archaeocyatha 15
Fig. 12. Ajacicyathidae (p. 13–16).
Orbicyathellus
Robustocyathellus
2b
2a
1b
1a
3
Rotundocyathus
5
4b
4a
Sibirecyathus
Stapicyathus Urcyathus
6
1a

16 Treatise Online, number XX
not located]. Cup in which both walls show peri-
odic, synchronous transverse folds; inner wall
with several rows of simple pores per intersept;
septa completely porous; pectinate tabulae may
be present. lower Cambrian (Tom.4–Bot.1): Sibe-
rian Platform, Altay Sayan, Tuva, Mongolia,
Transbaikalia, Morocco.——Fig. 11,6a–b. *O.
mongolicus, Burgasutay Formation, Atdabanian,
Seer’ Mountains, Ikh nuuruundyn hotgor, western
Mongolia, holotype; a, longitudinal section, ×3;
b, schematic reconstruction of cup, ×2 (Vologdin,
1937b).
Robustocyathellus KonYushKov, 1972, p. 133 [*R.
spinosus; OD; holotype, Ko nYu shKo v, 1972, pl.
13,1, not located]. Inner wall with one row of
simple pores per intersept; septa aporose to sparsely
porous. lower Cambrian (Atd.1–Bot.3): Siberian
Platform, Altay Sayan, Tuva, Mongolia, Trans-
baikalia, Far East, Tajikistan, southern China,
Canada.——Fig. 12,2a–b. *R. spinosus, Verkh-
nemonok Formation, Botoman, Kizas River,
western Sayan, Altay Sayan, Russia, holotype; a,
oblique longitudinal section, ×6; b, longitudinal
section, ×3 (Konyushkov, 1972).
Rotundocyathus vologDin, 1960, p. 422 [*R.
rotaceus; OD; holotype, vologDin, 1960, fig.
1zh, not located]. Inner wall with one row of
simple pores per intersept; septa completely porous;
pectinate tabulae may be present. lower Cambrian
(At d.2–Bot.1): Siberian Platform, Altay Sayan,
Tuva, Mongolia, Transbaikalia, Far East, Tarim,
Morocco, Iberia, France, Sardinia.——Fig . 12,3.
*R. rotaceus, Verkhneynyrga Formation, Botoman,
Lebed’ River, Altay Mountains, Altay Sayan, Russia,
holotype, transverse section, ×1 (Vologdin, 1960).
Sibirecyathus vologDin, 1937b, p. 468 [*S. nale-
tovi; M; holotype not designated, collection not
located]. Inner wall with one row of simple pores
per intersept; septa completely porous, linked by
synapticulae. lower Cambrian (Tom.3–Bot.3): Sibe-
rian Platform, Altay Sayan, Tuva, Mongolia, Trans-
baikalia, Far East, southern China, Morocco, Iberia,
France, Sardinia, Germany.——Fig. 12,4a–b.
*S. naletovi, Burgasutay Formation, Botoman,
Seer’ Mountains, Ikh nuuruundyn hotgor, western
Mongolia, a, unlocated syntype, transverse section,
×4; b, unlocated syntype, oblique longitudinal
section, ×4 (Vologdin, 1937b).
Stapicyathus Debrenne, 1964, p. 127, nom. transl.
Debrenne, 1970a, p. 43, ex Archaeocyathellus
(Stapicyathus) Debrenne, 1964, p. 127 [*Archaeo-
cyathus stapipora TAYlor, 1910, p. 118; OD; lecto-
type, TAYlor, 1910, pl. 7, photos 37a, 38D, 38G;
Debrenne, 1974b, pl. 24,1; SD Debrenne, 1970a,
p. 43, SAM T1591, Adelaide] [=Sivovicyathus
KonYus hKov, 1972, p. 134 (type, S. abakanensis,
OD); =Nochoroicyathellus osADchAYA in osADchAYA
& others, 1979, p. 154 (type, N. activus, OD), for
discussion, see Debrenne, ZhurAvlev, & roZAnov,
1989, p. 132; Debrenne, roZAnov, & ZhurAvlev,
1990, p. 161]. Inner wall with stirrup pores only;
septa aporose to sparsely porous; pectinate tabulae
may be present. lower Cambrian (Atd.2–Bot.3):
Siberian Platform, Altay Sayan, Tuva, Mongolia,
Transbaikalia, Far East, Australia, Antarctica, South
Africa (allochthonous).——Fig. 12,5. *S. stapipora
(TAYlor), Ajax Limestone, Botoman, Ajax Mine,
South Australia, Australia, lectotype, SAM T1591,
oblique longitudinal section, ×4 (Taylor, 1910).
Urcyathus vologDin, 1940b, p. 64 [*U. asteroides;
OD; holotype, vologDin, 1940b, pl. 14,5, not
located] [=Pectenocyathus KAshinA in rePinA &
others, 1964, p. 211 (type, P. torgaschinicus, OD),
for discussion, see Debrenne, Zhur Av le v , &
roZ An ov , 1989, p. 139; Debrenne, roZ An ov ,
& ZhurAvlev, 1990, p. 166]. Inner wall longitu-
dinally plicate, with several rows of simple pores
per intersept; septa completely porous; pectinate
tabulae may be present. lower Cambrian (Atd.1–
Atd.2): Altay Sayan, Tuva, Mongolia, Iberia,
Germany.——Fig. 12,6. *U. asteroides, Gavr-
ilovskoe Formation, Atdabanian, Gorskino, Salair,
Russia, holotype, oblique transverse section, ×9
(Vologdin, 1940b).
Family DENSOCYATHIDAE
Vologdin, 1937
[Densocyathidae vologDin, 1937b, p. 471] [=Leptosocyathidae volog-
Din, 1961, p. 178; =Tennericyathidae roZAnov in ZhurAvlevA, Korshu-
nov, & roZAnov, 1969, p. 34]
Inner wall with bracts or scales. lower
Cambrian (Atd.1–Bot.3).
Densocyathus vologDin, 1937b, p. 471 [*D.
sanaschticolensis; M; holotype not designated,
collection not located]. Inner wall with several rows
of pores per intersept, bearing upwardly projecting,
S-shaped scales; septa aporose to sparsely porous.
lower Cambrian (Bot.1–Bot.3): Altay Sayan.——
Fig. 13,1. *D. sanaschticolensis, Verkhnemonok
Formation, Botoman, Sanashtykgol Spring,
western Sayan, Altay Sayan, Russia; unlocated
syntype, transverse section of modular skeleton,
×5 (Vologdin, 1937b).
Cadniacyathus R. be DFor D & J. beDF orD, 1937, p.
36 [*C. asperatus; OD; lectotype, R. beDForD & j.
beDForD, 1937, fig. 152; Debrenne, 1974b, pl. 27,2;
SD Debrenne, 1970a, p. 30, USNM PU86616(1),
Washington, D.C.]. Inner wall with several rows of
pores per intersept, bearing upwardly projecting,
planar fused bracts; septa aporose to sparsely porous.
lower Cambrian (Bot.3): Australia, ?Antarctica.——
Fig. 13,2. *C. asperatus, Ajax Limestone, Botoman,
Ajax Mine, South Australia, Australia, lectotype,
USNM PU86616(1), oblique transverse view, ×3
(Debrenne, Zhuravlev, & Kruse, 2002).
Dailycyathus Debrenne, 1970a, p. 32 [*Paranacyathus
margarita R. beDFo rD & J. beD Fo rD , 1937, p.
34; OD; lectotype, R. beDF or D & j. be DF or D,
1937, fig. 138b–c; Debrenne, 1970a, pl. 1,5;
SD Debrenne, 1970a, p. 32, USNM PU87214,

Systematic Descriptions of the Archaeocyatha 17
Fig. 13. Densocyathidae (p. 16–18).
2
1
Densocyathus
Cadniacyathus
3b
3a
Dailycyathus
4b
4a Deceptioncyathus
5
Khirgisocyathus

18 Treatise Online, number XX
specimen 304, Washington, D.C.] [=Joanaecy-
athus grAvesTocK, 1984, p. 53 (type, J. cupu-
losus, OD; =Paranacyathus margarita R. be DFor D
& J. beDF or D, 1937, p. 34), for discussion, see
Debrenne, roZ An ov , & ZhurAvlev, 1990, p.
139]. Inner wall with one row of pores per inter-
sept, bearing upwardly projecting, cupped bracts;
septa aporose to sparsely porous. lower Cambrian
(Atd.4–Bot.3): Altay Sayan, Mongolia, southern
China, Australia.——Fig. 13,3a–b. *D. margarita
(r. be D Fo rD & j. beD Fo rD ), Ajax Limestone,
Atdabanian, Paint Mine, South Australia, Australia;
a, lectotype, USNM PU87214, specimen 304,
transverse view near cup base, ×5; b, paralectotype,
USNM PU87215, tangential view of inner wall,
×5 (Debrenne, 1970a).
Deceptioncyathus grAvesTocK, 1984, p. 53 [*D.
synapticulosus; OD; holotype, gr A v es T o c K ,
1984, fig. 34H–J, SAM P21504-1, Adelaide].
Inner wall with one row of pores per intersept,
bearing upwardly projecting cupped bracts; septa
completely porous, linked by synapticulae. lower
Cambrian (Atd.4): Australia.——Fi g. 13,4a–b.
*D. synapticulosus, Ajax Limestone, Atdabanian,
Mount Scott Range, South Australia, Australia,
holotype, SAM P21504-1; a, transverse section,
×2; b, longitudinal section, ×2 (Gravestock,
1984).
Khirgisocyathus voronin, 1988, p. 5 [*K. primus;
OD; holotype, voronin, 1988, pl. 2,1, PIN
3301/511, Moscow]. Inner wall with several rows
of pores per intersept, bearing upwardly projecting
cupped bracts; septa completely porous. lower
Cambrian (Atd.2): Mongolia.——Fig . 13,5. *K.
primus, Ichituin Formation, Atdabanian, Boro-
Khairkhan-Obo Mountain, Khan-Khukhiy Range,
Mongolia, holotype, PIN 3301/511, oblique trans-
verse section, ×9 (Voronin, 1988).
Leptosocyathellus osADchAYA in osADchAYA & others,
1979, p. 119 [*L. mirandus; OD; holotype, osAD-
chAYA & others, 1979, pl. 5,3, VSEGEI 11594,
St. Petersburg]. Inner wall with stirrup pores only,
bearing upwardly projecting, S-shaped scales;
septa completely porous. lower Cambrian (Atd.2–
Atd.4): Altay Sayan, Iberia.——Fig. 14,1a–b. *L.
mirandus; a, Usa Formation, Atdabanian, Krutoy
Log, Batenev Range, Kuznetsk Alatau, Russia, holo-
type, VSEGEI 11594, transverse section, ×8.5; b,
Usa Formation, Atdabanian, Srednyaya Mountain,
Batenev Range, Kuznetsk Alatau, Russia, VSEGEI
C-69, oblique longitudinal section, ×8 (Osadchaya
& others, 1979).
Leptosocyathus vologDin, 1937b, p. 470 [*L .
curviseptum; OD; holotype, vologDin, 1937b,
fig. 14, not located; =Leptocyathus curvisep-
tatus vologDin, 1940a, p. 146] [=Leptocyathus
vologDin, 1937b, p. 468, nom. null. pro Leptoso-
cyathus vologDin, 1937b, p. 470; non Leptocyathus
milne-eDWArD s & hAime, 1850, a scleractinian;
=Halysicyathus Debrenne, 1965, p. 144 (type, H.
multifurcus, OD)]. Inner wall with one row of
pores per intersept, bearing upwardly projecting,
S-shaped scales; septa aporose to sparsely porous.
lower Cambrian (Atd.1–Bot.3): Siberian Platform,
Altay Sayan, Tuva, Mongolia, Transbaikalia, Far
East, Tajikistan, Australia, Antarctica, Morocco,
Iberia.——Fig. 14,2a–c. *L. curviseptus, Burgasutay
Formation, Botoman, Seer’ Mountains, Ikh nuuru-
undyn hotgor, western Mongolia; a–b, holotype; a,
transverse section, ×1; b, oblique transverse section
of inner wall, ×1 (Vologdin, 1937b); c, topotype,
PIN 3156/3000, transverse section, ×9 (Debrenne,
Zhuravlev, & Kruse, 2002).
Natalijaecyathus KoTe l’n i Kov , 1995, p. 23 [*N.
vadibalaensis; OD; holotype, KoTe l’niKov , 1995,
fig. 1d, pl. 2,7, TsNIGRm 12890/4, St. Petersburg].
Inner wall with stirrup pores only, bearing upwardly
projecting, S-shaped scales; longitudinal bars may
be present, bisecting stirrup pores; septa completely
porous. lower Cambrian (Atd.2): Tuva.——Fig .
14,3a–b . *N. va di balae nsis, Il’chir Formation,
Atdabanian, Vadi-Bala Creek, Tapsa River, Tuva,
Russia, holotype, TsNIGRm 12890/4; a, oblique
transverse section, ×5.5; b, detail of oblique trans-
verse section, ×9 (Kotel’nikov, 1995).
Rectannulus Debrenne, 1977a, p. 106 [*R. wille-
fertae; OD; holotype, Debrenne, 1977a, pl. 6,2–3,
MNHN M80026, specimen IRH4-2b, Paris]. Inner
wall with stirrup pores only, bearing horizontal
to upwardly projecting, S-shaped scales; scales
may be fused into pseudoannuli; septa sparsely to
completely porous. lower Cambrian (Atd.4–Bot.1):
Morocco.——Fig . 15,1a–b. *R. willefertae, Issafen
Formation, Botoman, Jbel Irhoud, Morocco, holo-
type, MNHN M80026, specimen IRH4-2b; a,
detail of transverse section, ×5 (Debrenne, 1977a);
b, oblique longitudinal section, ×5 (M. Debrenne,
new).
Tennericyathus roZ A no v in Zhur Avl ev A, Kor s -
hunov, & ro Z An o v , 1969, p. 35 (roZ A n o v
in ro Z A n o v & others, 1969, p. 182, nom.
nud.) [*T. malycanicus; OD; holotype, Zhu rAv-
le vA, Korshunov, & roZA n ov, 1969, pl. 4,5;
roZA nov , 1973, pl. 9,4, PIN 4297/79, Moscow,
not located] [=Memoriacyathus YAZmir in
ZhurAvlevA, 1974a, p. 215, nom. nud.; =Memo-
riacyathus YAZmir in YAZmir, DAl m A To v , &
YAZmir, 1975, p. 47 (type, M. burjaticus, OD);
=Raropectinus Debrenne & ro Z An o v , 1983,
p. 73 5, nom. nov. pr o Ra roc yathu s osA D ch AY A
in os A D c h AY A & oth ers , 1 979, p. 155 , n o n
volo gD in & jAnK Au sK As , 1968, p. 203, cribri-
cyath (type, R. rarus, OD), for discussion, see
Debrenne, ZhurAvlev, & roZA no v, 1989, p.
135; Debrenne, roZA nov , & ZhurAvlev, 1990,
p. 163]. Inner wall with several rows of pores per
intersept, bearing upwardly projecting, S-shaped
scales; septa completely porous; pectinate tabulae
may be present. lower Cambrian (Atd.1–Bot.1):
Siberian Platform, Altay Sayan, Tuva, Mongolia,
Transbaikalia, Far East.——Fig . 15,2. *T. maly-
canicus, Pestrotsvet Formation, Atdabanian,
Malykan, Lena River, Sakha (Yakutia), Russia,
holotype, PIN 4297/79, detail of oblique trans-
verse section, ×15 (Zhuravleva, Korshunov, &
Rozanov, 1969).

Systematic Descriptions of the Archaeocyatha 19
Family BRONCHOCYATHIDAE
R. Bedford & J. Bedford, 1936
[Bronchocyathidae R. beDForD & J. beDForD, 1936, p. 25] [=Stillicido-
cyathidae Ting, 1937, p. 367; =Thalamocyathidae ZhurAvlevA, 1954, p.
28; =Cyclocyathellidae ZhurAvlevA, 1960c, p. 74; =Trininaecyathidae De-
brenne, 1964, p. 114; =Compositocyathidae ZhurAvlevA in ZhurAv l evA
& others, 1967, p. 52; =Glaessnericyathidae Debrenne, 1970a, p. 35]
Inner wall with annuli. lower Cambrian
(Atd.1–Bot.3).
Thalamocyathus gorDon, 1920, p. 687 [*Archaeo-
cyathus trachealis TAYlor, 1910, p. 125; SD Ting,
1937, p. 368, by elimination; lectotype, TAYlor ,
1910, pl. 8, photo 47(8); hill, 1965, pl. 7,1;
Debrenne, 1973, pl. 1,6; SD Debrenne, 1969b,
p. 262; SAM T1555A, Adelaide] [=Bronchocyathus
R. be DF or D & J. beDFo rD , 1936, p. 25 (type,
Archaeocyathus trachealis TAYlo r, 1910, p. 125,
OD); =Thalamopectinus Debrenne, 1973, p. 8
(type, T. arterialis, OD; =Archaeocyathus trache-
alis TAYl or , 1910, p. 125), for discussion, see
Debrenne & Kruse, 1989, p. 27; =Gordonicyathella
YAZmir in ZhurAvlevA, 1974a, p. 160, nom. nud.;
=Gordonicyathellus YAZmir in YAZmir, DA lmATov,
& YAZmir, 1975, p. 48 (type, G. solidus, OD), for
discussion, see Debrenne, ZhurAvlev, & roZAnov,
1989, p. 136; Debrenne, roZAnov, & ZhurAvlev,
1990, p. 164]. Inner wall with one pore row per
intersept and upright, V-shaped annuli; septa
aporose to sparsely porous; pectinate tabulae may
be present. lower Cambrian (Atd.2–Bot.3): Altay
Sayan, Tuva, Mongolia, Transbaikalia, Australia,
Antarctica, South Africa (allochthonous), ?Falk-
land Islands (allochthonous).——Fi g. 16,1a–b.
*T. trachealis (TAYlor), Ajax Limestone, Botoman,
Ajax Mine, South Australia, Australia, lectotype,
SAM T1555A; a, transverse view, ×3.5 (Debrenne,
1973); b, external longitudinal view of cup, ×8
(Taylor, 1910).
Fig. 14. Densocyathidae (p. 18).
1a
Leptosocyathellus
Leptosocyathus
Natalijaecyathus
2b
2a
1b
2c
3a
3b

20 Treatise Online, number XX
Compositocyathus ZhurAvlevA, 1960b, p. 159
[*Thalamocyathus muchattensis ZhurAvlevA in
ZhurAvlevA & Zelenov, 1955, p. 71; OD;
holotype, ZhurAvlevA & Zelenov, 1955, pl.
2,1–2; Zhur Avle vA, 1960b, pl. 10,3–5, TsSGM
205/47a-b, Novosibirsk]. Inner wall with one pore
row per intersept and planar annuli bearing short
beams that support supplementary microporous
sheath; septa aporose to sparsely porous; pectinate
tabulae may be present. lower Cambrian (Atd.1–
Bot.1): Siberian Platform, Altay Sayan, Tuva,
Transbaikalia.——Fig. 16,2a–b. *C. muchattensis
(ZhurAvlevA), Pestrotsvet Formation, Atdabanian,
Mukhatta River, Lena River, Sakha (Yakutia),
Russia; a, holotype, TsSGM 205/47b, longitudinal
section (outer wall to left), ×7 (Zhuravleva &
Zelenov, 1955); b, specimen TsSGM 323, oblique
transverse section, ×7 (Debrenne, Zhuravlev, &
Kruse, 2002).
Conannulofungia YuAn in Yu An & ZhAng, 1980,
p. 383 [*C. jinshaensis; OD; holotype, Yu A n
& ZhAng, 1980, pl. 2,1a–f, NIGP 51288,
Nanjing]. Inner wall with one pore row per
intersept and upwardly projecting, S-shaped
annuli linked to septa by ribs; septa completely
porous, linked by synapticulae. lower Cambrian
(Bot.1–Bot.2): southern China.——Fi g .
17,1a–c. *C. jinshaensis, Minxinsi (Minghs-
in gs su ) Fo rm a ti on , Bo t om an , Ya n ko ng ,
Guizhou, China; a–b, holotype, NIGP 51288;
a, transverse section, ×4; b, longitudinal section
near inner wall, ×4 (Yuan & Zhang, 1980); c,
specimen MNHN M85006, longitudinal section
near inner wall, ×8 (Debrenne, Zhuravlev, &
Kruse, 2002).
Cyathocricus Debrenne, 1969a, p. 318 [*Archaeo-
cyathus tracheodentatus R. be D Fo r D & W. R.
beDForD, 1934, p. 2; OD; lectotype, R. beDForD &
W. r. beDForD, 1934, fig. 5; Debrenne, 1969a, pl.
5,4–5; SD Debrenne, 1969a, p. 319, NHM S4148,
London; =Ethmophyllum dentatum TAYl or, 1910,
p. 129; lectotype, TAYlor, 1910, pl. 16, photo 89;
Fig. 15. Densocyathidae (p. 18).
2
Tennericyathus
Rectannulus
1b
1a

Systematic Descriptions of the Archaeocyatha 21
Debrenne, 1970a, pl. 1,1; SD Debrenne, 1974b,
p. 132, SAM T1606C-D, Adelaide] [=Cricopectinus
Debrenne, 1970a, p. 32 (type, C. dentulus, OD)].
Inner wall with one pore row per intersept and
commonly horizontally projecting waved annuli
that may mutually coalesce; denticles may be
present on annular rims; septa aporose to sparsely
porous; pectinate tabulae may be present. lower
Cambrian (?Atd.4–Bot.3): Altay Sayan, Tuva, Far
East, Australia, Antarctica, ?Morocco.——Fig.
17,2a–c. *C. dentatus (TAYlo r), Ajax Limestone,
Botoman, Ajax Mine, South Australia, Australia,
lectotype, SAM T1606C-D; a, transverse view,
×6; b, oblique longitudinal view, ×5 (Debrenne,
Zhuravlev, & Kruse, 2002); c, oblique longitudinal
view, ×4 (Taylor, 1910).
Cyclocyathella vologDin in Zhu r Avl e vA, KrAs-
noPe evA, & chernYshevA, 1960, p. 105 [*Cyclo-
cyathus yakovlevi vologDin, 1931, p. 49; OD;
lectotype, vologDin, 1931, pl. 4,7–8; SD
Debrenne, ZhurAvle v, & Krus e, 2002, p. 1569,
TsNIGRm 44a/2956, St. Petersburg] [=Cyclocyathus
vologDin, 1928, p. 30, nom. nud., non milne-
eDWAr Ds & hAime, 1850, p. liv, scleractinian,
Fig. 16. Bronchocyathidae (p. 19–20).
2a
Compositocyathus
2b
1a
1b
Thalamocyathus

22 Treatise Online, number XX
nec DuncAn & ThomPson, 1867, p. 1, rugose
coral; =Cyclocyathus simon, 1939, p. 27 (type, C.
yakovlevi vologDin, 1931, p. 49)]. Inner wall with
one pore row per intersept and inverted V-shaped
annuli; septa completely porous. lower Cambrian
(Atd.2): Altay Sayan, Tuva, Far East.——Fi g.
18,1a–c. *C. yakovlevi (vologDin), Torgashino
Formation, Atdabanian, Kameshki, eastern Sayan,
Altay Sayan, Russia; a, paralectotype, TsNIGRm
45/2956, transverse section, ×6; b, paralectotype,
TsNIGRm 47a/2956, longitudinal section, ×6; c,
schematic sketch of septum in longitudinal section
(outer wall to left), ×8 (Vologdin, 1931).
Denaecyathus ZhurAvlevA in ZhurAvlevA & others,
1967, p. 57 [*D. biporosus; OD; holotype, ZhurAv-
levA & others, 1967, pl. 17,5–6, TsSGM 325/17,
Novosibirsk]. Inner wall with several pore rows
per intersept and upright, V-shaped annuli; septa
aporose to sparsely porous. lower Cambrian (Bot.1):
Altay Sayan, Tuva, Mongolia, Transbaikalia, Far
East.——Fig . 18,2a–c. *D. biporosus, Shangan
Formation, Botoman, Shivelig-Khem River, eastern
Tannu-Ola Range, Tuva, Russia; a, holotype,
TsSGM 325/17, oblique transverse section, ×5;
b, paratype, TsSGM 325/16, oblique longitudinal
section (outer wall at bottom), ×5; c, holotype,
TsSGM 325/17, oblique transverse section (outer
wall to left), ×5 (Zhuravleva & others, 1967).
Gordonicyathus ZhurAvlevA, 1959, p. 426 [*Thala-
mocyathus gerassimovensis KrAsnoPeevA, 1955, p.
95; OD; holotype not designated, collection not
located] [=Sichotecyathus oKu ne vA in oK une vA
& re PinA, 1973, p. 138 (type, S. orientalis, OD),
for discussion, see Debrenne, Zhur Av le v , &
roZ An ov , 1989, p. 109; Debrenne, roZ An ov ,
& ZhurAvlev, 1990, p. 144]. Inner wall with
one pore row per intersept and upright, V-shaped
annuli; septa completely porous; pectinate tabulae
may be present. lower Cambrian (Atd.1–Bot.3):
Siberian Platform, Altay Sayan, Tuva, Mongolia,
Transbaikalia, Far East, Australia.——Fig. 18,3.
*G. gerassimovensis (KrAsno PeevA), Verkhnemonok
Formation, Botoman, Gerasimov Spring, Monok
River, western Sayan, Altay Sayan, Russia, unlo-
cated syntype, oblique transverse section, ×7 (Kras-
nopeeva, 1955).
Gordonifungia roZAnov in rePinA & others, 1964,
p. 193 [*G. batinensis; OD; holotype, rePinA &
others, 1964, pl. 11,1, PIN 4297/24, Moscow].
Inner wall with one pore row per intersept and
upright, V-shaped annuli; septa completely porous,
linked by synapticulae. lower Cambrian (Atd.3–
Bot.1): Altay Sayan, ?Morocco.——Fig. 19,1. *G.
batinensis, Usa Formation, Atdabanian, Verkhnyaya
Erba, Batenev Range, Kuznetsk Alatau, Russia,
holotype, PIN 4297/24, oblique transverse section,
×4 (Repina & others, 1964).
Morenicyathus Perejón, 1975b, p. 169 [*M. arru-
zafai; OD; holotype, Perejón, 1975b, pl. 6,5–6;
Perejón, 1975c, pl. 6,8–9, CE 62-14, Madrid;
=Archaeocyathellus (Protocyathus) cordobae simon,
1939, p. 74; holotype, simon, 1939, pl. 2,11,
SM 26-179e, Frankfurt am Main] [=Kellericyathus
roZAnov, 1973, p. 61, nom. nud.; =Denaecyathellus
osA Dc h AY A in osADc h AY A & others, 1979, p.
122 (type, D. makarichus, OD); =Kellericyathus
roZAnov in Debrenne, ZhurAvlev , & roZAnov,
1989, p. 114 (type, K. altaicus, OD)]. Inner wall
with several pore rows per intersept and upright,
V-shaped annuli; septa completely porous. lower
Cambrian (Atd.2–Atd.3): Kolyma, Altay Sayan,
Iberia.——Fig. 19,2a–b. *M. cordobae (simon)
[=M. arruzafai Perejón], Pedroche Formation,
Atdabanian, Las Ermitas, Cordoba, Andalusia,
Spain, holotype, CE 62-14; a, oblique transverse
section, ×4; b, detail of longitudinal section (outer
wall to left), ×8 (Perejón, 1975c).
Pseudotennericyathellus osADchAYA in osADchAYA &
others, 1979, p. 120 [*Tennericyathus latus osAD-
chAYA in ZADoroZhnAYA, osADchAYA, & rePinA,
1973, p. 134; OD; holotype, ZADoroZhnAYA,
osADchAYA, & reP inA, 1973, pl. 19,3; osA DchAYA
& others, 1979, pl. 5,1–2, TsSGM IGiG424,
Novosibirsk]. Inner wall with several pore rows per
intersept and upwardly projecting, S-shaped annuli;
septa completely porous. lower Cambrian (Atd.2–
Atd.4): Altay Sayan, Mongolia.——Fig. 19,3a–b.
*P. latus (osADchAYA), Usa Formation, Atdabanian,
Bograd, Batenev Range, Kuznetsk Alatau, Russia,
holotype, TsSGM IGIG424; a, transverse section,
×9; b, longitudinal section (outer wall to left), ×9
(Osadchaya & others, 1979).
Sagacyathus Kruse , 1982, p. 178 [*S. stonyx; OD;
holotype, Kruse, 1982, fig. 15D–H, AM F.83576,
Sydney]. Inner wall with one pore row per intersept
and upright, V-shaped annuli bearing denticu-
late rims; septa aporose to sparsely porous; pecti-
nate tabulae may be present. lower Cambrian
(Atd.4–Bot.3): Altay Sayan, Mongolia, Far East,
Australia.——Fig. 19,4a–c. *S. stonyx, Cymbric
Vale Formation, Botoman, Mt. Wright, New South
Wales, Australia, holotype, AM F.83576 ; a, trans-
verse section AM FT.8487, ×6; b, oblique longi-
tudinal section AM FT.8486, ×4; c, longitudinal
section AM FT.8490, ×4 (Kruse, 1982).
Stillicidocyathus Ting, 1937, p. 367 [*Coscinocyathus
aulax TAYlor, 1910, p. 139; OD; lectotype, TAYlor,
1910, pl. 10, photo 57; SD Debrenne, 1969b, p.
263, sAm T1605A-B, Adelaide] [=Glaessnericyathus
Debrenne, 1970a, p. 35 (type, Bronchocyathus
sigmoideus R. beDFor D & J. beDF or D, 1936, p.
25, OD), for discussion, see grAvesTocK, 1984, p.
69; Debrenne, ZhurAvlev , & roZAnov, 1989, p.
133; Debrenne, roZAnov, & Zhur Avlev , 1990,
p. 162]. Inner wall with one pore row per intersept
and upwardly projecting, S-shaped annuli; septa
aporose to sparsely porous; pectinate tabulae may
be present. lower Cambrian (Bot.1–Bot.3): Altay
Sayan, Tuva, Mongolia, Australia, Antarctica,
Iberia, Sardinia.——Fig. 20,1. *C. aulax (TAYlor),

Systematic Descriptions of the Archaeocyatha 23
2b
2a
2c
Cyathocricus
1a
1b Conannulofungia
1c
Fig. 17. Bronchocyathidae (p. 20–21).

24 Treatise Online, number XX
Ajax Limestone, Botoman, Ajax Mine, South
Australia, Australia, lectotype, SAM T1605A-B,
longitudinal view, ×7 (Taylor, 1910).
Svetlanocyathus missArZhevsKiY & roZ Anov, 1962,
p. 43 [*S. primus; OD; holotype, missArZhevsKiY
& roZ Anov , 1962, pl. 3,2a–v; roZA nov , 1973,
pl. 1,5, PIN 4297/19-20, Moscow]. Outer wall
with slitlike, simple pores; inner wall with one
pore row per intersept and inverted V-shaped
annuli; septa aporose to sparsely porous. l ower
Cambrian (Bot.1): Altay Saya n, Tuva.——Fig .
20,2. *S. primus, Shangan Formation, Botoman,
Shivelig-Khem River, eastern Tannu-Ola Range,
Tuva, Russia, holotype, PIN 4297/19-20, oblique
transverse view, ×5 (Debrenne, Zhuravlev, &
Rozanov, 1989).
Taylorcyathus vologDin, 1955, p. 143 [*Cycloc y-
athus subtersiensis vologDin, 1940b, p. 63; OD;
holotype not designated, collection not located]
[=Tersicyathus vologDin, 1955, p. 143 (type,
Cyc locyath us ters ie nsis vologDin, 1931, p. 87,
OD); =Thalamocyathellus osADchAYA in osAD-
chAYA & others, 1979, p. 155 (type, T. inclinatus,
OD), for discussion, see Debrenne, ZhurAvlev,
& ro ZAnov , 1989, p. 134; Debrenne, roZAn ov,
& ZhurAvlev, 1990, p. 163; =Pospelovicyathus
KonYAevA in ZhurAvlevA & others, 1997a, p.
49 (type, P. gravis, OD)]. Inner wall with one
pore row per intersept and upwardly projecting,
S-shaped annuli; septa completely porous; pecti-
nate tabulae may be present. lower Cambrian
(At d.1–Bot.3): Siberian Platform, Altay Sayan,
Tuva, Mongolia, Transbaikalia, Far East, Australia,
southern China, Iberia, France, Sardinia.——Fig.
20,3. *T. subtersiensis (vologDin), Gavrilovskoe
Formation, Atdabanian, Belaya Gorka, Salair,
Russia, topotype, PIN 4754/50, oblique transverse
section, ×6 (Debrenne, Zhuravlev, & Kruse, 2002).
Taylorfungia Perejón, 1989, p. 180 [*Thalamocyathus
synapticulosus Zhu rAv le vA, 1955a, p. 41; OD;
holotype, ZhurAvlevA, 1955a, pl. 5,1, PIN 495,
Moscow, not located]. Inner wall with one pore row
per intersept and upwardly projecting, planar-to-S-
shaped annuli; septa completely porous, linked by
synapticulae. lower Cambrian (Atd.2–Atd.3): Altay
Sayan.——Fi g. 20,4. *T. synapticulosa (ZhurAv-
levA), Usa Formation, Atdabanian, Bol’shaya Erba
(Potekhino), Batenev Range, Kuznetsk Alatau,
Russia, holotype, PIN 495, oblique transverse
section, ×8 (Zhuravleva, 1955a).
Trininaecyathus ZhurAvlevA, 1960b, p. 218 [*T.
macroporus; OD; holotype not located; paratypes,
ZhurAvlevA, 1960b, pl. 18,6–8, fig. 122, TsSGM
205/100, 205/101, Novosibirsk]. Inner wall with
one pore row per intersept and upwardly projecting,
S-shaped annuli bearing denticulate rims; septa
aporose to sparsely porous; pectinate tabulae may
be present. lower Cambrian (Bot.1): Siberian Plat-
form.——Fig . 20,5. *T. macroporus, Perekhod
Formation, Botoman, Botoma River, Sakha
(Yakutia), Russia, paratype, TsSGM 205/100,
oblique transverse section, ×8 (Zhuravleva, 1960b).
Family ETHMOCYATHIDAE
Debrenne, 1969
[Ethmocyathidae Debrenne, 1969a, p. 322] [=Ethmopectinidae De-
brenne, 1970a, p. 25; =Diplocyathidae Debrenne, 1974b, p. 123;
=Zonacyathellidae ZhurAvlevA in ZhurAvlevA & elKinA, 1974, p. 65;
=Baikalocyathinae ZhurAvlevA in ZhurAvlevA & elKinA, 1974, p. 68;
=Inessocyathidae ZhurAvlevA in ZhurAvlevA & elKinA, 1974, p. 106;
=Hyptocyathidae Kruse, 1978, p. 29; =Gnaltacyathidae Kruse, 1982, p.
166; =Baikalopectinidae grAvesTocK, 1984, p. 66]
Inner wall with noncommunicating
canals. lower Cambrian (Atd.1–Bot.3).
Ethmocyathus R. beDForD & W. R. beDF orD, 1934,
p. 2 [*E. lineatus; M; holotype, R. beDForD &
W. R. beDForD, 1934, fig. 8; hill, 1965, pl. 4,2;
Debrenne, 1969a, pl. 5,1–3; Debrenne, 1974b,
pl. 27,1; NHM S4149, M, London] [=Ethmopec-
tinus Debrenne, 1970a, p. 34 (type, E. walteri;
OD), for discussion, see Debrenne, ZhurAvlev, &
roZAnov , 1989, p. 106; Debrenne, roZAn ov, &
ZhurAvlev, 1990, p. 142]. Inner wall with one row
of horizontal to upwardly projecting, straight canals
per intersept, formed by flexure of inner edges of
septa; supplementary screen of planar rings on
central cavity side; septa aporose to sparsely porous;
pectinate tabulae may be present. lower Cambrian
(Bot.3): Australia, Antarctica.——Fi g. 21,1a–b.
*E. lineatus, Ajax Limestone, Botoman, Ajax Mine,
South Australia, Australia, holotype, NHM S4149;
a, longitudinal view of inner wall, ×7; b, oblique
longitudinal view of inner wall (to left) and septa
(to right), ×15 (Debrenne, 1969a).
Afiacyathus voronin, 1962, p. 26 [*A. lativallum;
OD; holotype, voronin, 1962, pl. 4,4–5, PIN
1914/74-80a, Moscow, not located] [=Axiculifungia
F. Debrenne & M. Debrenne in F. Debrenne,
M. Debrenne, & roZ Anov , 1976, p. 102 (type,
Ajacicyathus compositus Debrenne, 1961, p. 9,
OD)]. Inner wall with one row of horizontal to
upwardly projecting, straight canals per intersept;
septa completely porous, linked by synapticulae.
lower Cambrian (Atd.2–Bot.1): Tuva, Morocco,
Iberia, Sardinia, ?Poland.——Fig. 21,2a–b. *A .
lativallum, Shangan Formation, Botoman, Shivelig-
Khem River, eastern Tannu-Ola Range, Tuva,
Russia, holotype, PIN 1914/74-80a; a, oblique
transverse view, ×2; b, oblique longitudinal view,
×2 (Voronin, 1962).
Baikalocyathus YAZmir in ZhurAvlevA, 1974a, p. 55
[*Ethmophyllum rossicum Zhur AvlevA, 1960b, p.
164; OD; holotype, ZhurAvlevA, 1960b, pl. 11,2,
TsSGM 205/51, Novosibirsk] [=Nochoroicyathella
Korshunov, 1983b, p. 110 (type, N. fragilis, OD);
=Baikalopectinus grAvesTo cK, 1984, p. 66 (type,
B. capulus, OD), for discussion, see Debrenne,
ZhurAvlev, & roZAn ov, 1989, p. 95; Debrenne,
roZAnov, & Zh urAvlev, 1990, p. 134]. Inner wall
with one row of downwardly projecting, straight
canals per intersept, bearing supplementary bracts
on central cavity side; septa completely porous;
pectinate tabulae may be present. lower Cambrian
(Atd.1–Bot.1): Siberian Platform, Kolyma, Altay

Systematic Descriptions of the Archaeocyatha 25
Fig. 18. Bronchocyathidae (p. 21–22).
3
2b
2a
2c
Denaecyathus
Gordonicyathus
1a
1b
1c
Cyclocyathella

26 Treatise Online, number XX
Fig. 19. Bronchocyathidae (p. 22).
4a
Pseudotennericyathellus
Sagacyathus
2a
2b
Morenicyathus
1Gordonifungia
3a
4b 4c
3b

Systematic Descriptions of the Archaeocyatha 27
2
3
4
5
Svetlanocyathus
Taylorcyathus
Taylorfungia
Trininaecyathus
1Stillicidocyathus
Fig. 20. Bronchocyathidae (p. 22–24).

28 Treatise Online, number XX
Sayan, Tuva, Mongolia, Transbaikalia, Far East,
Australia, ?Morocco.——Fig. 21,3a–b. *B. rossicus
(ZhurAvl evA); a, Pestrotsvet Formation, Atdaba-
nian, Oy-Muran, Lena River, Sakha (Yakutia),
Russia, holotype, TsSGM 205/51, detail of longi-
tudinal section of septum (outer wall to left),
×10 (Zhuravleva, 1960b); b, Pestrotsvet Forma-
tion, Atdabanian, Mukhatta River, Lena River,
Sakha (Yakutia), Russia, paratype, TsSGM 205/52,
oblique longitudinal section, ×6 (Debrenne,
Zhuravlev, & Kruse, 2002 ).
Carpicyathus osADc hAYA in Zhu rAvl evA & others,
1967, p. 51 [*C. mysticus; OD; holotype, ZhurAv-
levA & others, 1967, pl. 14,3–6, VSEGEI 9594,
St. Petersburg]. Inner wall with several rows of
horizontal to upwardly projecting, straight canals
per intersept, bearing supplementary bracts on
central cavity side; septa completely porous. lower
Cambrian (Atd.2–Bot.2): Altay Sayan, Tuva, Trans-
baikalia, Morocco.——Fig. 21,4a–b. *C. mysticus,
Shangan Formation, Shivelig-Khem River, eastern
Tannu-Ola Range, Tuva, Russia, holotype, VSEGEI
9594; a, transverse section, ×1.5; b, longitudinal
section, ×3.5 (Zhuravleva & others, 1967).
Degeletticyathus ZhurAvlevA in ZhurAvlevA, Kors-
hunov, & roZ Anov, 1969, p. 36 [*Ethmophyllum?
galuschkoi ZhurAvlevA, 1960b, p. 169; OD;
holotype, Zhu rAvl evA, 1960b, pl. 11,7, TsSGM
205/56, Novosibirsk] [=Degeletticyathellus ZhurAv-
levA in ZhurAvlevA & elKinA, 1974, p. 66 (type,
D. lebedevae, OD), for discussion, see Debrenne,
ZhurAvlev, & roZA nov, 1989, p. 103]. Inner wall
with horizontal to upwardly projecting, straight
stirrup canals only; septa aporose to sparsely porous.
lower Cambrian (Atd.2–Bot.1): Siberian Platform,
Altay Sayan, Tuva, Mongolia, Transbaikalia, Tajiki-
stan, Morocco.——Fig. 22,1a–b. *D. galuschkoi
(Zhur AvlevA), Oy-Muran reef massif, Botoman,
Lena River, Sakha (Yakutia), Russia; a, specimen
TsSGM 323/40, transverse section, Oy-Muran,
×5 (Zhuravleva, Korshunov, & Rozanov, 1969);
b, holotype, TsSGM 205/56, longitudinal section
of septum (outer wall to left), Mukhatta River, ×5
(Zhuravleva, 1960b).
Diplocyathellus Debrenne, 1977b, p. 1222, nom. nov.
pro Diplocyathus Debrenne, 1974b, p. 124, non
AllmAn, 1888, p. 16, cnidarian [*Archaeocyathus
retezona TAYlo r, 1910, p. 121; OD; lectotype,
TAYlor, 1910, pl. 6, photo 31; Debrenne, 1974b,
pl. 26,4–5; SD Debrenne , 1974b, p. 124, SAM
T1577A, Adelaide]. Inner wall with horizontal
to upwardly projecting, straight stirrup canals
only, canals branching toward central cavity; septa
aporose to sparsely porous. lower Cambrian (Bot.3):
Australia.——Fig. 22,2. *D. retezona (TAYlo r),
Ajax Limestone, Botoman, Ajax Mine, South
Australia, Australia, paralectotype, SAM T1591,
longitudinal view, ×4 (Debrenne, Zhuravlev, &
Kruse, 2002).
Frinalicyathus Debrenne, roZAnov, & ZhurAvlevA in
ZhurAvlevA & elKinA, 1974, p. 73 [*Leptosocyathus
altaicus roZAnov in rePinA & others, 1964, p. 190;
OD; holotype, rePi nA & others, 1964, pl. 4,2,
PIN 4297/21, Moscow] [=Pseudodegeletticyathellus
osADchAYA in osADc hAYA & others, 1979, p. 123
(type, P. ladae, OD), for discussion, see Debrenne,
roZAnov, & Zh urAvlev, 1990, p. 143]. Inner wall
with downwardly projecting, straight stirrup canals
only, bearing supplementary scales on central cavity
side; septa completely porous. lower Cambrian
(Atd.1–Atd.4): Altay Sayan, Mongolia.——Fig .
22,3. *F. al tai c us (roZA n ov ), Verkhneyn yrga
Formation, Atdabanian, Tyrga River, Altay Moun-
tains, Altay Sayan, Russia, holotype, PIN 4297/21,
transverse section, ×2.5 (Repina & others, 1964).
Gnaltacyathus Kruse, 1982, p. 166 [*G. nodus; OD;
holotype, Krus e, 1982, pl. 3,2–4, AM FT.8453,
8454, 8495b, Sy dney]. Inner wal l wit h hor i-
zontal to upwardly projecting, straight canals,
each canal spanning several intersepts; septa
completely porous. lower Cambrian (Bot.1–Bot.3):
Tuva, ?Mongolia, Australia.——Fig. 22,4a–b. *G.
nodus, Cymbric Vale Formation, Botoman, Mt.
Wright, New South Wales, Australia, holotype,
AM FT.8453, 8454, 8495b; a, transverse section,
AM FT.8495b, ×8; b, longitudinal section, AM
FT.8454, ×8 (Kruse, 1982).
Hyptocyathus Kruse, 1978, p. 30 [*H. licinus; OD;
holotype, Kru se , 1978, fig. 2–3, AM F.83402,
Sydney]. Inner wall with downwardly projecting,
straight stirrup canals only, bearing upwardly
projecting, branching canals on central cavity
side; septa aporose to sparsely porous. lo wer
Cambrian (Bot.3): Australia.——Fi g. 23,1a–c.
*H. licinus, Cymbric Vale Formation, Botoman,
Mt. Wright, New South Wales, Australia, holo-
type, AM F.83402; a, oblique transverse section,
AM FT.14171, ×4; b, longitudinal section, AM
FT.14173, ×4; c, tangential section of inner wall,
AM FT.14174, ×15 (Kruse, 1978).
Inessocyathellus belYAevA in ZhurAvlevA & elKinA,
1974, p. 78 [*I. synapticulosus; OD; holotype,
ZhurAvlevA & elKinA, 1974, pl. 7,3, DVGU,
Khabarovsk]. Inner wall with one row of horizontal
to upwardly projecting, straight canals per intersept;
septa aporose to sparsely porous, linked by synap-
ticulae. lower Cambrian (Bot.3): Far East.——Fig .
23,2a–b. *I. synapticulosus, Ust’toka unit, Botoman,
Verkhneurminsk Spring, Dzhagdy Range, Far East,
Russia, holotype, DVGU, Khabarovsk; a, transverse
section, ×10; b, oblique longitudinal section, ×10
(Zhuravleva & Elkina, 1974).
Inessocyathus Debrenne, 1964, p. 143 [*Archaeocy-
athus spatiosus bornemAnn, 1886, p. 59; OD; lecto-
type, bornemAnn, 1886, pl. 15,1a; SD Debrenne,
1964, p. 143, not located; topotype, Debrenne,
1964, pl. 9,1–2, MNHN M84074, specimen
SPi-13, Paris] [=Voroninicyathus ZhurAvl evA in
ZhurAvle vA & elKinA, 1974, p. 79 (type, Inesso-
cyathus karakolicus voronin, 1969, p. 103, OD);
=Rowanpectinus grAvesTocK, 1984, p. 67 (type,
R. clarus, OD), for discussion, see Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 112; Debrenne,
roZAnov, & Zh urAvlev, 1990, p. 146]. Inner wall
with one row of horizontal to upwardly projecting,
straight canals per intersept; septa completely

Systematic Descriptions of the Archaeocyatha 29
Fig. 21. Ethmocyathidae (p. 24–28).
2b
2a
4b
3b
Aacyathus
Baikalocyathus
1a
1b
Ethmocyathus
4a
Carpicyathus
3a

30 Treatise Online, number XX
Fig. 22. Ethmocyathidae (p. 28).
3
2
4b
Diplocyathellus
Frinalicyathus
Gnaltacyathus
1a
1b
4a
Degeletticyathus
I had to
reduce
view 3
from X3.5
to X2.5 to
fit the two
new views.
Is this
okay? If
not, please
suggest
crop lines
or reduc-
tions for
other views
on the
plate.

Systematic Descriptions of the Archaeocyatha 31
Fig. 23. Ethmocyathidae (p. 28–32).
1b
3a
1c
1a
3b
2b
2a
Hyptocyathus
Inessocyathellus
Inessocyathus

32 Treatise Online, number XX
porous. lower Cambrian (Atd.1–Bot.3): Siberian
Platform, Altay Sayan, Tuva, Mongolia, Far East,
Australia, southern China, Morocco, Iberia, France,
Sardinia, Germany.——Fig. 23,3a–b. *I. spatiosus
(bornemAnn), Matoppa Formation, Botoman,
San Pietro, Sardinia, Italy; a, lectotype, transverse
section, ×2.5 (Bornemann, 1886); b, topotype,
MNHN M84074, specimen SPi-13, longitudinal
section, ×2.5 (Debrenne, Zhuravlev, & Kruse,
2002).
Mackenziecyathus hAnDFielD, 1971, p. 43 [*M .
bukryi; OD; holotype, hAnDFielD, 1971, pl.
5,1a–d, GSC 25334, Ottawa] [=Ussuricyathus
oKu nevA in oK un evA & re Pi nA , 1973, p. 113
(type, U. kropotkini, OD)]. Inner wall with hori-
zontal to upwardly projecting straight stirrup canals
only, bearing supplementary scales on central cavity
side; septa aporose to sparsely porous; pectinate
tabulae may be present. lower Cambrian (Bot.1):
Altay Sayan, Tuva, Mongolia, Far East, Canada,
United States.——Fig . 24,1a–c. *M. bukryi,
unnamed Sekwi Formation equivalent (map unit
5 of hAnDFielD, 1971), Botoman, Coal River,
Yukon Territory, Canada, holotype, GSC 25334;
a, transverse section, ×4; b, longitudinal section
(outer wall to right), ×4; c, tangential section of
inner wall, ×4 (Handfield, 1971).
Rasetticyathus Debrenne, 1971, p. 193 [*R. igle-
siensis; OD; holotype, Debrenne, 1971, fig. 1–2,
not located; =Archaeocyathus acutus bornemAnn,
1886, p. 50; holotype not designated; for discus-
sion, see Debrenne, roZA no v, & ZhurAvlev,
1990, p. 157]. Inner wall with one row of hori-
zontal to upwardly projecting, S-shaped canals
per intersept; septa aporose to sparsely porous;
synapticulae may be present. lower Cambrian
(Bot.1–Bot.2): southern China, Morocco, Iberia,
Sardinia.——Fig. 24,2. *R. iglesiensis, Matoppa
Formation, Botoman, Monte Cuccurinu, Sardinia,
Italy, holotype, transverse section, ×16 (Debrenne,
1972).
Terraecyathus ZhurAvlevA in Zhur AvlevA & elKinA,
1974, p. 104 [*T. lathentis; OD; holotype,
Zhur Avl evA & elKinA, 1974, pl. 23,2, TsSGM
442/37, 38, Novosibirsk] [=Sericyathus voronin,
1988, p. 7 (type, S. tartsinicus, OD)]. Inner
wall with several rows of horizontal to upwardly
projecting, straight canals per intersept; septa
completely porous. lower Cambrian (Atd.1–Bot.1):
Altay Sayan, Tuva, Mongolia, Morocco.——Fig.
24,3a–b. *T. lathentis; a, Adiak Formation, Atda-
banian, Tom’ River, Gornaya Shoria, Altay Sayan,
Russia, holotype, TsSGM 442/37, transverse
section, ×5 (Debrenne, Zhuravlev, & Kruse,
2002); b, Usa Formation, Botoman, Bograd,
Batenev Range, Kuznetsk Alatau, Russia, paratype,
TsSGM 442/38, longitudinal section of septum
(outer wall to left), ×5 (Zhuravleva & Elkina,
1974).
Ussuricyathellus voronin, 1988, p. 6 [*U. bellus;
OD; holotype, voronin, 1988, pl. 1,4, PIN 3175-
920/a-2, Moscow]. Inner wall with several rows of
horizontal to upwardly projecting, straight canals
per intersept; septa aporose to sparsely porous.
lower Cambrian (Bot.1): Mongolia.——Fig. 24,4.
*U. bellus, Burgasutay Formation, Botoman, Seer’
Mountains, Ikh nuuruundyn hotgor, western
Mongolia, holotype, PIN 3175-920/a-2, transverse
section, ×5 (Voronin, 1988).
Zonacyathellus ZhurAvlevA in ZhurAvlevA & elKinA,
1974, p. 66 [*?Zonacyathus monoporosus Zhur Av-
levA in ZhurAvlevA & others, 1967, p. 66; OD;
holotype, Zhu rAvl evA & others, 1967, pl. 23,2;
ZhurAvlevA & elKinA, 1974 , pl. 3,2, TsSGM
325/35, Novosibirsk]. Inner wall with one row of
horizontal to upwardly projecting, straight canals
per intersept; septa aporose to sparsely porous.
lower Cambrian (Bot.1): Tuva.——Fig. 24,5. *Z.
monoporosus (Zhu rAvl evA), Shangan Formation,
Botoman, Shivelig-Khem River, eastern Tannu-Ola
Range, Tuva, Russia, holotype, TsSGM 325/35,
longitudinal section, ×5 (Zhuravleva & others,
1967).
Family SAJANOCYATHIDAE
Vologdin, 1956
[Sajanocyathidae vologDin, 1956, p. 879] [=Formosocyathidae ZhurAvlevA,
1957, p. 175; =Irinacyathidae ZhurAvlevA in Debrenne, 1972, p. 173, nom.
neg.; =Irinaecyathidae ZhurAvlevA in ZhurAvlevA & elKinA, 1974, p. 67]
Inner wall with communicating canals.
lower Cambrian (Atd.1–Toy.2).
Sajanocyathus vologDin, 1940b, p. 81 (vologDin,
1937b, p. 471, nom. nud.) [*S. ussovi; OD; lecto-
type, vologDin, 1940b, pl. 22,8; SD Zhu rAvlev,
2001a, p. 92, PIN 4754/2, Moscow] [=Sayanocy-
athus vologDin, 1937b, p. 479, nom. nud. (type,
Sayanocyathus ussovi vologDin, 1937b, p. 479,
M)]. Inner wall with several rows of anastomosing,
horizontal to upwardly and laterally projecting,
waved canals per intersept; septa aporose to sparsely
porous. lower Cambrian (Bot.1–Toy.2): Siberian
Platform, Altay Sayan, ?Antarctica, ?northeastern
China (Hinggan), ?Sardinia, Canada, United
States.——Fig. 25,1. *S. ussovi, Verkhnemonok
Formation, Botoman, Sanashtykgol River, western
Sayan, Altay Sayan, Russia, lectotype, PIN
4754/2, transverse section of modular skeleton,
×8 (Debrenne, Zhuravlev, & Kruse, 2002).
Chakassicyathus ZhurAvlevA & osADchAYA in ZhurAv-
levA & elKinA, 1974, p. 93 [*Ethmophyllum pseudo-
ratum ZhurAvlevA in ZhurAvlevA & others, 1967,
p. 62; OD; holotype, Zhu rAvlevA & others, 1967,
pl. 21,2, TsSGM 325/28b, Novosibirsk]. Inner wall
with one row of downwardly projecting, straight
porous canals per intersept, bearing supplementary
bracts or annuli on central cavity side; septa aporose
to sparsely porous. lower Cambrian (Bot.1): Altay
Sayan, Tuva.——Fig. 25,2. *C. pseudoratus (ZhurAv-
levA), Shangan Formation, Botoman, Shivelig-Khem
River, eastern Tannu-Ola Range, Tuva, Russia, holo-
type, TsSGM 325/28b, longitudinal section, ×5.5
(Zhuravleva & others, 1967).
Formosocyathus vologDin, 1940b, p. 90 (vologDin,
1937b, p. 471, nom. nud.) [*F. bulynnikovi; OD;
Chakas-
sicyathus:
should this
be "longi-
tudinal
sections"
to describe
views a and
b??

Systematic Descriptions of the Archaeocyatha 33
1b
3a
1a
2
3b
Mackenziecyathus
Rasetticyathus
Terraecyathus
Ussuricyathellus
5 Zonacyathellus 4
Fig. 24. Ethmocyathidae (p. 32).
1c

34 Treatise Online, number XX
holotype not designated, collection not located].
Inner wall with one row of anastomosing, hori-
zontal to upwardly and laterally projecting, waved
canals per intersept; supplementary spines, annular
structures, and/or microporous sheath may be
present on central cavity side; septa completely
porous; pectinate tabulae may be present. lower
Cambrian (Atd.1–Bot.2): Altay Sayan, Tuva,
Mongolia, Transbaikalia.——Fig. 25,3a–b. *F.
bulynnikovi, Verkhnemonok Formation, Botoman,
Sanashtykgol River, western Sayan, Altay Sayan,
Russia, unlocated syntype; a, transverse section
(outer wall at top), ×6; b, longitudinal section
(outer wall to left), ×6 (Vologdin, 1940b).
Irinaecyathus ZhurAvlevA in ZhurAvlevA & elKinA,
1974, p. 87 [*Ethmophyllum grandiperforatum
vologDin, 1940a, p. 160; OD; lectotype,
vologDin, 1940a, fig. 75, pl. 46,1, SD Zhu rAv-
levA & elKinA, 1974, p. 88, collection not located]
[=Kandatocyathus KAshinA in osADchAYA & others,
Fig. 25. Sajanocyathidae (p. 32–34).
3a
2
1
3b
Sajanocyathus
Chakassicyathus
Formosocyathus

Systematic Descriptions of the Archaeocyatha 35
1979, p. 156 (type, K. kalleganovi; OD), for
discussion, see Debrenne, ZhurAvlev, & roZAnov,
1989, p. 112; Debrenne, roZAnov, & ZhurAvlev,
1990, p. 146]. Inner wall with one row of down-
wardly projecting, straight porous canals per inter-
sept, bearing supplementary bracts or annuli on
central cavity side; septa sparsely to completely
porous; pectinate tabulae may be present. lower
Cambrian (Bot.1–Toy.2): Siberian Platform, Altay
Sayan, Tuva, Mongolia, Transbaikalia, Far East,
southern China.——Fig. 26,1. *I. grandiperforatus
(vologDin), Burgasutay Formation, Botoman,
Seer’ Mountains, Ikh nuuruundyn hotgor, western
Mongolia, topotype, PIN 4327/24-2042/5, trans-
verse section, ×5 (Debrenne, Zhuravlev, & Kruse,
2002).
Kiwicyathus Debrenne & Kru se, 1986, p. 250 [*K.
nix; OD; holotype, Debrenne & Kruse , 1986, fig.
15A–B, VU VC19, Wellington]. Inner wall with
horizontal to upwardly projecting, straight porous
stirrup canals only; septa aporose to sparsely porous.
lower Cambrian (Bot.3): Antarctica.——Fig. 26,2.
*K. nix, Shackleton Limestone, Botoman, Mt.
Egerton, Byrd Glacier, Antarctica, holotype, VU
VC19, transverse section, ×6 (Debrenne & Kruse,
1986).
Palmericyathus hAnDFielD, 1971, p. 44 [*Eth mo-
phyllum lineatum greggs, 1959, p. 66; OD; holo-
type, greggs, 1959, pl. 14,2, GSC 14315, Ottawa;
=Ethmophyllum americanum oKuliTch in co oPer
& others, 1952, p. 30; holotype, cooPer & others,
1952, pl. 7,3–4, USNM 111816, Washington, D.C.
Fig. 26. Sajanocyathidae (p. 34–37).
3a
1
2
3b
Irinaecyathus
Kiwicyathus
Palmericyathus
3c

36 Treatise Online, number XX
Fig. 27. Sajanocyathidae (p. 37).
1a
2c
1b
Zonacyathus
2a
2b
Siderocyathus

Systematic Descriptions of the Archaeocyatha 37
(for discussion, see Debrenne, 1987, p. 270)]. Inner
wall with anastomosing, horizontal to upwardly and
laterally projecting, waved stirrup canals only;
septa aporose to sparsely porous. lower Cambrian
(Bot.1): Canada, United States, Mexico.——Fig.
26,3a–c. *P. americanus (oKuliTch); a–b, Puerto
Blanco Formation, Botoman, Caborca, Sonora,
Mexico, holotype, USNM 111816; a, transverse
section, ×8; b, detail of transverse section (outer
wall at bottom), ×25 (Cooper & others, 1952); c,
[=P. lineatus (greggs)], Laib Formation, Botoman,
Salmo, British Columbia, Canada, holotype, GSC
14315, transverse section (outer wall at top), ×6
(Greggs, 1959).
Siderocyathus Debrenne & gAngloFF in Debrenne,
gAnDin, & gAngloFF, 1990, p. 87 [*S. duncanae;
OD; holotype, Debrenne, gAnDin, & gAngloFF,
1990, pl. 1,7, USNM 443555, specimen IR1-3,
Washington, D.C.]. Inner wall with one row of
short, noncommunicating, horizontal to upwardly
projecting canals per intersept, continuing into
central cavity as communicating waved canals
bearing supplementary bracts on central cavity
side; septa aporose to sparsely porous, linked by
synapticulae. lower Cambrian (Bot.1): Unit ed
States.——Fig. 27,1a–b. *S. duncanae, Valmy
Formation, Botoman, Iron Canyon, Nevada,
United States, holotype, USNM 443555, spec-
imen IR1-3; a, transverse section, ×5; b, longi-
tudinal section, ×5 (Debrenne, Gandin, &
Gangloff, 1990).
Zonacyathus R. beDForD & J. beDForD, 1937, p.
36 [*Archaeocyathus retevallum R. be DF or D &
W. R. beDForD, 1934, p. 2; OD; holotype, R.
beDF orD & W. R. beD For D, 1934, fig. 6; hill,
1965, pl. 4,3; Debrenne, 1969a, pl. 4,4; NHM
S4147, M, London]. Inner wall with one row of
horizontal to upwardly projecting, straight canals
per intersept, canals branching and becoming
porous toward central cavity; septa sparsely to
completely porous. lower Cambrian (?Bot.1,
Bot.2–Bot.3): ?Siberian Platform, Australia.——
Fig . 27,2a–c. *Z. retevallus (R. beDForD & W.
r. beDForD), Ajax Limestone, Botoman, Ajax
Mine, South Australia, Australia; a–b, holotype,
NHM S4147; a, oblique longitudinal view,
×5 (Debrenne, Zhuravlev, & Kruse, 2002); b,
tangential view of inner wall, ×5 (Hill, 1965);
c, paratype, USNM PU86606, longitudinal view
(outer wall to right), ×5 (Debrenne, 1974b).
Family BIPALLICYATHIDAE
Debrenne, Rozanov, & Zhuravlev 1989
[Bipallicyathidae Debrenne, ro ZAn ov, & ZhurAv lev in Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 82]
Inner wall with attached microporous
sheath. lower Cambrian (Atd.2).
Bipallicyathus ZhurAvlev in voronin & others, 1982,
p. 78 [*B. manifestus; OD; holotype, voronin
& others, 1982, pl. 15,6a–b, PIN 3302/3305,
Moscow] [=Kashinaecyathus YAroshevich, 1990,
p. 25 (type, K. salairicus; OD), for discussion, see
Debrenne & Zhur Avle v, 1992b, p. 173]. Inner
wall with one pore row per intersept and attached
microporous sheath; septa completely porous. lower
Cambrian (Atd.2): Altay Sayan, Mongolia.——Fig.
28,1a–c. *B. manifestus, Salaany Gol Formation,
Atdabanian, Khasagt-Khayrkhan Range, Tsagaan
Oloom province, western Mongolia; a, paratype,
PIN 3302/3006, oblique transverse section, ×8;
b–c , holotype, PIN 3302/3305; b, longitudinal
section, ×7; c, detail of inner wall, ×17 (Voronin
& others, 1982).
?Heckericyathus ZhurAvlevA, 1960b, p. 220
[*Ethmophyllum heckeri ZhurAvlevA in ZhurAv-
le vA & Zelenov, 1955, p. 69; OD; holotype,
PIN 1161, Moscow, not located; paratypes,
ZhurAvlevA & Zelenov, 1955, pl. 1,3–4,
TsSGM 205/102, 205/103, Novosibirsk] [=Heck-
erocyathus Zhu r Av l evA in vologDin, 1957a,
p. 180, nom. nud.]. Inner wall with one pore
row per intersept and independent microporous
sheath, each micropore bearing a supplementary
bract; septa completely porous; pectinate tabulae
may be present. [Inner wall bears supplementary
elements atypical of other members of family.]
lower Cambrian (Atd.1–Atd.4): Siberian Plat-
form, Transbaikalia, Far East.——Fig . 28,2a–c.
*H. heckeri (Zhur Av le vA ), Pestrotsvet Forma-
tion, Atdabanian, Oy-Muran, Lena River, Sakha
(Yakutia), Russia; a, paratype, TsSGM 205/102,
transverse section, ×8 (Zhuravleva & Zelenov,
1955); b–c, paratype, TsSGM 205/103; b, detail
of septum in longitudinal section (outer wall to
right), ×16; c, detail of inner wall in oblique
longitudinal section, ×16 (Debrenne, Zhuravlev,
& Kruse, 2002).
Superfamily
PRETIOSOCYATHOIDEA
Rozanov, 1969
[nom. c orrect. Debrenne, ZhurAvlev, & Kruse, herein, pro Pre tiosocy-
athacea roZAnov, 1969, p. 112]
Outer wall with independent micropo-
rous sheath. lower Cambrian (Atd.1–Bot.2).
Family ROBERTOCYATHIDAE
Rozanov, 1969
[Robertocyathidae roZAnov, 1969, p. 112]
Inner wall with simple pores. l o wer
Cambrian (Atd.2–Bot.1).
Robertocyathus roZAnov , 1969, p. 112 [*R. polaris;
OD; holotype, roZA nov, 1969, pl. 42,1–2, PIN
4297/96, Moscow]. Inner wall with several rows
of simple pores per intersept; septa completely
porous. lower Cambrian (Atd.1–Bot.3): Sibe-
rian Platform, Altay Sayan, Australia, Morocco,
Iberia.——Fig. 29,1. *R. polaris, Erkeket Forma-
tion, Botoman, Khorbusuonka River, Olenek Basin,

38 Treatise Online, number XX
Sakha (Yakutia), Russia, holotype, PIN 4297/96,
transverse section, ×7 (Debrenne, Zhuravlev, &
Kruse, 2002).
Mattajacyathus roZAn ov in Debrenne, ZhurAvlev,
& roZAnov , 1989, p. 118 (roZAnov, 1973, p. 61,
nom. nud.; roZ An ov in Debrenne & ro ZA nov ,
1983, p. 735, nom. nud.) [*Robertocyathus arduus
roZAnov, 1969, p. 113; OD; holotype, roZA nov,
1969, pl. 42,3–4; roZA nov , 1973, pl. 5,1, PIN
4297/97, Moscow]. Cup in which both walls show
periodic, synchronous transverse folds; inner wall
with several rows of simple pores per intersept;
septa completely porous. lower Cambrian (Bot.1):
Siberian Platform.——Fig. 29,2a–b. *M. arduus
(roZAn ov), Erkeket Formation, Botoman, Khor-
busuonka River, Olenek Basin, Sakha (Yakutia),
Russia, holotype, PIN 4297/97; a, longitudinal
section, ×7; b, detail of outer wall in tangential
section, ×15 (Debrenne, Zhuravlev, & Kruse,
2002).
Fig. 28. Bipallicyathidae (p. 37).
2c
2b
Heckericyathus
1c
1a
1b
Bipallicyathus
2a

Systematic Descriptions of the Archaeocyatha 39
1
3
2a
4
5
2b
6
Robertocyathus
Mattajacyathus
Urcyathella
Pretiosocyathus
Jangudacyathus
Loculicyathopsis
Fig. 29. Robertocyathidae and Pretiosocyathidae (p. 37–40).

40 Treatise Online, number XX
Urcyathella Zhu rAv l e vA in musA To v & others,
1961, p. 25 [*U. tercyathoides; OD; holotype,
mus AT ov & others, 1961, pl. 3,8–9, TsS GM
264/26, Novosibirsk]. Inner wall longitudi-
nally plicate, with several rows of simple pores
per intersept; septa completely porous. low er
Cambrian (Atd.4–Bot.1): Altay Sayan.——Fig.
29,3. *U. tercyathoides, Balakhtinson Forma-
tion, Atdabanian, Kazyr River, eastern Sayan,
Altay Sayan, Russia, holotype, TsSGM 264/26,
transverse section, ×7 (Debrenne, Zhuravlev, &
Kruse, 2002).
Family PRETIOSOCYATHIDAE
Rozanov, 1969
[Pretiosocyathidae roZAnov, 1969, p. 112]
Inner wall with noncommunicating
canals. lower Cambrian (Atd.1–Bot.1).
Pretiosocyathus roZA nov in roZ Ano v & missAr-
ZhevsKiY, 1966, p. 55 [*P. subtilis; OD ;
holotype, ro ZA n ov & missArZhevsKiY, 1966,
pl. 4,4; roZA nov , 1973, pl. 11,3, PIN 4297/65,
Moscow] [=Cosmocyathus YAZmir in Zhu rAvle vA,
1974a, p. 96, nom. nud.; =Cosmocyathus YAZmir
in YAZmir, DAl mAT ov, & YAZmir, 1975, p. 63
(type, C. perforatus, OD); =Pretiosocyathellus
osA Dc h AY A in osADc h AY A & others, 1979, p.
133 (type, P. toltschiensis, OD); =Grandicyathus
Korshunov, 1983b, p. 109 (type, G. lectus; OD),
for discussion, see Debrenne, Zh ur Av l ev , &
roZA nov, 1989, p. 125; Debrenne, ro ZAn ov, &
ZhurAvlev, 1990, p. 156]. Inner wall with hori-
zontal to upwardly projecting, straight stirrup
canals only; septa completely porous; pecti-
nate tabulae may be present. lower Cambrian
(At d.1–Bot.1): Siberian Platform, Altay Sayan,
Tuva, Mongolia, Transbaikalia, Tajikistan.——
Fig . 29,4. *P. subtilis, Usa Formation, Atdaba-
nian, Bol’shaya Erba, Batenev Range, Kuznetsk
Alatau, Russia, holotype, PIN 4297/65, trans-
verse section, ×7 (Debrenne, Zhuravlev, &
Kruse, 2002).
Jangudacyathus YAZmir in YAZmir, DAl mAT ov, &
YAZmir, 1975, p. 62 (YAZmir in Zhur Av le vA ,
1974a, p. 180, nom. nud.) [*J. simplex; OD;
holotype, YAZmir, DAlmATov, & YAZmir, 1975, pl.
23,5, BGU 0138/21, Ulan-Ude]. Inner wall with
several rows of horizontal to upwardly projecting,
straight canals per intersept; septa aporose to
sparsely porous. lower Cambrian (Bot.1): Trans-
baikalia.——Fig. 29,5. *J. simplex, Uran Forma-
tion, Botoman, Yanguda River, Vitim Highlands,
Transbaikalia, Russia, holotype, BGU 0138/21,
transverse section, ×7 (Debrenne, Zhuravlev, &
Kruse, 2002).
Loculicyathopsis boY A r in o v in Zh u rA v le vA &
others, 1997a, p. 61 [*L. septospinosus; OD;
holotype, Zh u r Av l e vA & others, 1997a, pl.
11,9, ZSGGU 2329/62, Novokuznetsk]. Inner
wall with one row of horizontal to upwardly
projecting, straight canals per intersept; septa
completely porous. lower Cambrian (Atd.2):
Altay Sayan.——Fig. 29,6. *L. septospinosus, Usa
Formation, Atdabanian, Malaya Belokamenka
River, Kiya River, Kuznetsk Alatau, Russia,
holotype, ZSGGU 2329/62, oblique transverse
section, ×5 (Zhuravleva & others, 1997a).
Superfamily
ERBOCYATHOIDEA
Vologdin & Zhuravleva, 1956
[nom. correct. Debrenne & Kruse , 1986, p. 251, p ro Erbocyathacea
ZhurAvlevA, 1960b, p. 187, nom. transl. ex Erbocyathidae vologDin
& ZhurAvlevA in vologDin, 1956, p. 879] [=Bosceculcyathacea KrAs-
noPeevA, 1959, p. 7, nom. transl. hill, 1972, p. 77, ex Bosceculcyathidae
KrAsnoPeevA, 1959, p. 7; =Kordecyathoidea missArZhevsKiY, 1961, p.
21, nom. trans l. missArZhevsKiY in rePinA & others, 1964, p. 218, ex
Kordecyathidae missArZhevsKiY, 1961, p. 21, nom. correct. pro Kordecy-
athacea Debrenne, roZAnov, & ZhurAvlev in Debrenne, ZhurAvlev,
& roZAnov, 1989, p. 82]
Outer wall with attached microporous
sheath. lower Cambrian (Atd.1–Toy.3).
Family ERBOCYATHIDAE
Vologdin & Zhuravleva, 1956
[Erbocyathidae vologDin & ZhurAvlevA in vologDin, 1956, p. 879,
nom. nov. pro Polycyathidae vologDin, 1928, p. 35, invalid name based
on junior homonym] [=Ladaecyathidae Debrenne, 1964, p. 114]
Inner wall with simple pores. l o wer
Cambrian (Atd.1–Toy.3).
Pluralicyathus oKuliTch, 1950c, p. 503, nom. nov. pro
Polycyathus vologDin, 1928, p. 32, non DuncAn,
1876, p. 433, cnidarian] [*Polycyathus heterovallum
vologDin, 1928, p. 36; SD simon, 1939, p. 34;
lectotype, vologDin, 1928, pl. 2,1,2,4,5; SD
Debrenne, ZhurAvle v, & Kruse, 2002, p. 1586,
TsNIGRm 1/a-t/2617, St. Petersburg] [=Erbocyathus
Zhur Avle vA, 1950, p. 857, nom. nud.; =Erbocy-
athus ZhurAvlevA, 1955a, p. 44, nom. nov. pro
Polycyathus vologDin, 1928, p. 32 (type, Polycy-
athus heterovallum, SD simon, 1939, p. 34), non
DuncAn, 1876, p. 433, cnidarian; application by
Debrenne, ZhurAvlev, & Kruse (2003) to suppress
Pluralicyathus and conserve Erbocyathus rejected
by ICZN (2005); =Neocyathus vologDin, 1960,
p. 422 (type, Archaeocyathus laevus vologDin,
1940b, p. 57, OD)]. Inner wall with several rows of
simple pores per intersept; septa aporose to sparsely
porous. lower Cambrian (Bot.1–Toy.3): Siberian
Platform, Altay Sayan, Tuva, Mongolia, Far East,
Uzbekistan.——Fig. 30,1a–b. *P. heterovallum
(vologDin); a, Torgashino Formation, Toyonian,
Uyar River, eastern Sayan, Altay Sayan, Russia, unlo-
cated specimen, transverse section, ×8.5 (Debrenne,
Zhuravlev, & Kruse, 2002); b, Khomustakh Forma-
tion, Toyonian, Amga River, Sakha (Yakutia), Russia,
specimen TsSGM 205/71, section of modular skel-
eton, ×1 (Zhuravleva, 1960b).
Ladaecyathus ZhurAvlevA, 1960a, p. 43 [*Tegerocy-
athus limbatus Zhu rAv le vA, 1955a, p. 46; OD;
holotype, ZhurAvlevA, 1955a, pl. 5,3–4, PIN 494,

Systematic Descriptions of the Archaeocyatha 41
Moscow, not located]. Inner wall with several rows of
simple pores per intersept; septa completely porous;
pectinate tabulae may be present. lower Cambrian
(Atd.4–Bot.3): Siberian Platform, Kolyma, Altay
Sayan, Transbaikalia, Far East, Australia, Antarctica,
Morocco.——Fig. 30,2a–b. *L. limbatus (Zh urAv-
levA), Usa Formation, Botoman, Mt. Martyukhina,
Batenev Range, Kuznetsk Alatau, Russia; a, holotype,
PIN 494, transverse section, ×6.5 (Zhuravleva,
1955a); b, TsSGM 273/4d, longitudinal section of
septum (outer wall to left), ×7 (Debrenne, Zhuravlev,
& Kruse, 2002).
Milaecyathus Debrenne & ZhurAvlev, 2000, p. 49
[*Ladaecyathus melnikovae ZhurAvlev in voronin
& others, 1982, p. 79; OD; holotype, voronin &
others, 1982, pl. 16,2,5, PIN 3302/300v, Moscow].
Inner wall with stirrup pores only; septa completely
porous; pectinate tabulae may be present. lower
Cambrian (Atd.1–Atd.2): Altay Sayan, Mongolia.——
Fig. 30,3a–b. *M. melnikovae (ZhurAvlev), Salaany
Gol Formation, Atdabanian, Salaany-Gol River,
Khasagt-Khairkhan Range, Tsagaan Oloom prov-
ince, western Mongolia, holotype, PIN 3302/300v;
a, transverse section, ×5 (Voronin & others, 1982);
b, detail of outer wall in tangential section, ×25
(Debrenne, Zhuravlev, & Kruse, 2002).
Family PEREGRINICYATHIDAE
Zhuravleva, 1967
[Peregrinicyathidae ZhurAvlevA in ZhurAvlevA & others, 1967, p. 74]
Inner wall with annuli. lower Cambrian
(Bot.1–Bot.2).
Peregrinicyathus ZhurAvlevA in ZhurAvlevA &
others, 1967, p. 75 [*P. dorotheae; OD; holotype,
Zhu rAv le vA & others, 1967, pl. 28,1, TsSGM
325/54, Novosibirsk]. Inner wall with one pore row
per intersept and upright, V-shaped annuli; septa
completely porous. lower Cambrian (Bot.1–Bot.2):
Altay Sayan, Tuva.——Fig. 31,1a–b. *P. dorotheae,
Shangan Formation, Botoman, Shivelig-Khem River,
eastern Tannu-Ola Range, Tuva, Russia; a, holotype,
TsSGM 325/54, transverse section, ×4.5 (Debrenne,
Zhuravlev, & Kruse, 2002); b, paratype, TsSGM
325, specimen 1, thin section 1, sample 314-7,
OR-64, detail of oblique transverse section (outer
wall to left), ×8 (Zhuravleva & others, 1967).
Family VOLOGDINOCYATHIDAE
Yaroshevich, 1957
[Vologdinocyathidae YAroshevich, 1957, p. 1015] [=Bosceculcyathidae
KrAsno PeevA, 1959, p. 7; =Kordecyathidae missArZhevsKiY, 1961, p.
21; =Schidertycyathidae KrAsn oPeevA, 1969, p. 63; =Gumbycyathidae
Debrenne & Kruse, 1986, p. 253]
Inner wall with noncommunicating
canals. lower Cambrian (Bot.1–Toy.2).
Vologdinocyathus YAroshevich, 1957, p. 1015 [*V.
erbiensis; OD; holotype, YAroshevich, 1957, fig.
1a–v, TsSGM 499/1a-b, Novosibirsk] [=Tegero-
cyathella KonYushKov, 1967, p. 109 (type, T.
borovikovi, OD); =Larecyathus KAshinA, 1979, p.
46, nom. nud.; =Larecyathus KAshinA in osAD-
chAYA & others, 1979, p. 145 (type, L. infinitus,
OD), for discussion, see Debrenne, ZhurAvlev, &
roZAnov , 1989, p. 139; Debrenne, roZA nov, &
ZhurAvlev, 1990, p. 167]. Inner wall with one row
of horizontal to upwardly projecting, straight canals
per intersept; septa aporose to sparsely porous.
lower Cambrian (Bot.3–Toy.2): Altay Sayan, Tuva,
Mongolia, Kazakhstan, Uzbekistan, Antarctica,
Greenland.——Fig. 31,2a–b. *V. erbiensis, Usa
Formation, Toyonian, Bol’shaya Erba, Batenev
Range, Kuznetsk Alatau, Altay Sayan, Russia,
holotype, TsSGM 499/1a-b; a, detail of trans-
verse section, ×30; b, detail of septum in longitu-
dinal section (outer wall to left), ×50 (Debrenne,
Zhuravlev, & Kruse, 2002).
Gumbycyathus Kruse, 1982, p. 168 [*G. pythoni; OD;
holotype, Kru se, 1982, pl. 4,1–5, AM F.83930,
Sydney]. Inner wall with several rows of horizontal
to upwardly projecting, straight canals per intersept,
bearing supplementary bracts on central cavity side;
septa completely porous. lower Cambrian (Bot.1–
Bot.3): Mongolia, Australia.——Fig. 32,1a–d. *G.
pythoni, Cymbric Vale Formation, Botoman, Mt.
Wright, New South Wales, Australia, holotype, AM
F.83930; a, transverse section (outer wall to right),
AM FT.8457, ×6; b, detail of longitudinal section
(outer wall to left), AM FT.8455, ×8; c, detail of
outer wall in tangential section, AM FT.8458, ×8;
d, detail of inner wall in tangential section, AM
FT.8456, ×8 (Kruse, 1982).
Inacyathella Debrenne, 1977a, p. 109 [*I. pulchra;
OD; holotype, Debrenne, 1977a, pl. 8,3–4,
MNHN M80037, Paris]. Inner wall with one row
of horizontal to upwardly projecting, S-shaped
canals per intersept; septa completely porous. lower
Cambrian (Bot.1): Morocco.——Fi g. 31,3a–b. *I.
pulchra, Issafen Formation, Botoman, Jbel Irhoud,
Morocco, holotype, MNHN M80037; a, oblique
transverse section, ×3.5 (Debrenne, Zhuravlev,
& Kruse, 2002); b, detail of septum in longitu-
dinal section (outer wall to left), ×10 (Debrenne,
1977a).
Kordecyathus missArZhevsKiY, 1961, p. 21 [*K.
shiveligensis; OD; holotype, missArZhevsKiY,
1961, pl. 1,3, PIN 1914/73M/1, Moscow, not
located]. Inner wall with one row of horizontal to
upwardly projecting, straight canals per intersept,
bearing downwardly projecting cupped bracts
on central cavity side; septa completely porous;
pectinate tabulae may be present. lower Cambrian
(Bot.1–Bot.2): Tuva, Mongolia.——Fig. 31,4a–b.
*K. shiveligensis, Shangan Formation, Botoman,
Shivelig-Khem River, eastern Tannu-Ola Range,
Tuva, Russia, holotype, PIN 1914/73M/1; a,
detail of transverse section (outer wall at top), ×8
(Missarzhevskiy, 1961); b, oblique longitudinal
section, ×3 (Debrenne, Zhuravlev, & Kruse, 2002).
Sanarkophyllum Debrenne & Kr us e , 1986 , p.
254 [*Formosocyathus antarcticus hill, 1964c, p.
616; OD; holotype, hill, 1964c, fig. 1(4a), An
62/1B/p, not located]. Inner wall with one row

42 Treatise Online, number XX
Fig. 30. Erbocyathidae (p. 40–41).
1a
3a
2a
1b
3b
2b
Pluralicyathus
Ladaecyathus
Milaecyathus

Systematic Descriptions of the Archaeocyatha 43
of downwardly projecting, straight canals per
intersept, bearing supplementary bracts on central
cavity side; septa aporose to sparsely porous. lower
Cambrian (Bot.3): Antarctica.——Fig. 32,2a–d.
*S. antarcticum (hill), Shackleton Limestone,
Botoman; a, Plunket Point, Beardmore Glacier,
Antarctica, holotype, An 62/1B/p, oblique trans-
verse section, ×2 (Hill, 1964c); b–d, Holyoake
Range, Nimrod Glacier, Antarctica, specimen GNS
MG509; b, transverse section, ×4; c, oblique longi-
tudinal section, ×3.5; d, detail of repeated longi-
tudinal section (outer wall to left), ×6 (Debrenne,
Zhuravlev, & Kruse, 2002).
Syringocyathus vologDin, 1940b, p. 82 (vologDin,
1937b, p. 471, nom. nud.) [*S. aspectabilis;
OD; lectotype, vologDin, 1940b, pl. 23,3; SD
ZhurAvle v, 2001a, p. 92, PIN 4754/3, Moscow]
[=Schidertycyathus KrAsnoPeevA, 1959, p. 3
(type, S. borucaevi, M); =?Bosceculcyathus KrAs-
noP eevA , 1959, p. 7 (type, B. agyrekensis, OD);
=?Boscekulcyathus KrA sno PeevA, 1959, p. 7, nom.
null.; =Schidertycyathellus KonYushKov, 1967, p.
108 (type, S. borukaevi, OD); =Syringocyathellus
KAsh in A in os AD ch AYA & others, 1979, p. 149
(type, S. kazachstani, OD), for discussion, see
Debrenne, roZA no v, & ZhurAvlev, 1990, p.
162]. Inner wall with several rows of horizontal to
upwardly projecting, straight canals per intersept;
septa completely porous, linked by interseptal
plates. lower Cambrian (Bot.3–Toy.2): Altay Sayan,
Tuva, Kazakhstan, Uzbekistan.——Fig. 32,3. *S.
aspectabilis, Verkhnemonok Formation, Botoman,
Abakan River, western Sayan, Altay Sayan, Russia,
lectotype, PIN 4754/3, oblique transverse section,
×5 (Vologdin, 1940b).
Family TEGEROCYATHIDAE
Krasnopeeva, 1972
[Tegerocyathidae KrAsnoPeevA, 1972, p. 145]
Inner wall with communicating canals.
lower Cambrian (Bot.1–Toy.3).
Tegerocyathus KrAsnoPeevA, 1955, p. 90 (KrAs-
noP ee vA, 1953, p. 52, 56, nom. nud.) [*Ethmo-
phyllum abakanensis vologDin, 1 940b, p.
69; holotype not designated, collection not
located; Zh ur Avl evA , 1960b, p. 192, invalidly
nominated Ethmophyllum edelsteini vologDin,
1931, p. 47, as type species] [=Tegerocoscinus
KrAsnoPeevA, 1972, p. 145 (type, T. tchesno-
kovensis, OD); =Alexandricyathus KAshinA in
osA D ch AY A & others, 1979, p. 142 (type, A .
ultrus, OD), for discussion, see Debrenne,
Zh u r Av l e v , & ro Z A n o v , 1989, p. 134;
Debrenne, roZA no v, & ZhurAvlev, 1990, p.
163]. Inner wall with one row of horizontal
to upwardly projecting, straight porous canals
per intersept; septa sparsely to completely
porous; pectinate tabulae may be present. lowe r
Cambrian (Bot.1–Toy.3): Siberian Platform,
Altay Sayan, Mongolia, Uzbekistan, Antarctica,
Greenland, United States.——Fig . 33,1a–b. *T.
abakanensis (vologDin), Verkhnemonok Forma-
tion, Botoman, Abakan River, western Sayan,
Altay Sayan, Russia, unlocated syntype; a ,
transverse section, ×5; b, longitudinal section,
×5 (Vologdin, 1940b).
Krasnopeevaecyathus ro Z A n ov in reP i n A &
others, 1964, p. 208 [*K. tyrgaensis; OD; holo-
type, rePinA & others, 1964, pl. 21,3–4 ),
PIN 4297/26-27, Moscow] [=Krishnanicyathus
vologDin, 1964b, p. 358 (type, K. elegans,
OD); =Ethmosyringocyathus Ko nYu shK ov, 1972,
p. 138 (type, E. primus, OD)]. Inner wall longi-
tudinally plicate, with several rows of anasto-
mosing, horizontal to upwardly projecting, waved
canals per intersept; septa completely porous.
lower Cambrian (Bot.2): Altay Sayan, United
States.——Fig . 33,2. *K. tyrgaensis, Verkh-
neynyrga Formation, Botoman, Tyrga River,
Altay Mountains, Altay Sayan, Russia, holotype,
PIN 4297/26-27, oblique transverse section,
×3.5 (Repina & others, 1964).
Superfamily
TUMULOCYATHOIDEA
Krasnopeeva, 1953
[nom. correct. ZhurAvlev & roZAnov in voronovA & others, 1987, p.
21, pro Tumulocyathacea Debrenne, 1964, p. 113, nom. transl. ex Tumu-
locyathidae KrAsnoPeevA, 1953, p. 56] [=Geocyathacea Debrenne, 1964,
p. 114, nom. nud., nom. transl. roZAnov, 1973, p. 86 ex Geocyathidae
Debrenne, 1964, p. 114 ]
Outer wall with simple tumuli. lower
Cambrian (Tom.2–Bot.3).
Family TUMULOCYATHIDAE
Krasnopeeva, 1953
[Tumulocyathidae KrAsnoPeevA, 1953, p. 56] [=Kotuyicyathidae roZAnov
in roZAnov & others, 1969, p. 186, nom. nud.]
Inner wall with simple pores. l o wer
Cambrian (Tom.2–Bot.3).
Tumulocyathus vologDin, 1937b, p. 470 [*T. pustulatus;
M; holotype not designated, collection not located]
[=Kotuyicyathus ZhurAvlevA, 1960b, p. 226 (type,
K. kotuyikensis, OD), for discussion, see Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 138; Debrenne,
roZAnov, & ZhurAvlev, 1990, p. 165; =Tumulocy-
athoides boYArinov & KonYAevA in ZhurAvlevA &
others, 1997b, p. 123 (type, T. kiyaensis, OD)]. Inner
wall with one row of simple pores per intersept; septa
completely porous; pectinate tabulae may be present.
lower Cambrian (Tom.2–Bot.1): Siberian Platform,
Altay Sayan, Tuva, Mongolia, Australia.——Fig. 34,1.
*T. pustulatus; Salaany Gol Formation, Atdabanian,
Salaany Gol, Khasagt-Khayrkhan Range, Tsagaan
Oloom province, western Mongolia, specimen PIN

44 Treatise Online, number XX
Fig. 31. Peregrinicyathidae and Vologdinocyathidae (p. 41).
4a
2a
1b
4b
2b
Peregrinicyathus
Vologdinocyathus
1a
3a
3b
Inacyathella
Kordecyathus

Systematic Descriptions of the Archaeocyatha 45
1a
2b
1d
2d
2c
2a
1b
1c
3
Sanarkophyllum
Gumbycyathus
Syringocyathus
Fig. 32. Vologdinocyathidae (p. 41–43).

46 Treatise Online, number XX
3302/710, oblique transverse section, ×7 (Debrenne,
Zhuravlev, & Kruse, 2002).
Isiticyathus Korshunov, 1972, p. 60 [*?Tumulifungia
ultra Korshunov in Zh urAvlevA, Korshunov, &
roZA nov , 1969, p. 38; OD; holotype, Zhur Av-
levA , Korshunov, & roZAn ov, 1969, pl. 10,2;
Korshunov, 1972, pl. 8,6, TsSGM 323/45, Novo-
sibirsk]. Inner wall with one row of simple pores
per intersept; septa completely porous, linked by
synapticulae. lower Cambrian (Atd.4–Bot.1): Sibe-
rian Platform, Transbaikalia.——Fig. 34,2a–b. *I.
ultra (Korshunov), Oy-Muran reef massif, Atda-
banian, Oy-Muran, Lena River, Sakha (Yakutia),
Russia, holotype, TsSGM 323/45; a, transverse
section, ×8 (Zhuravleva, Korshunov, & Rozanov,
1969); b, detail of transverse section (outer wall
at bottom), ×13 (Debrenne, Zhuravlev, & Kruse,
2002).
Kotuyicyathellus osADchAYA in osADchAYA & others,
1979, p. 157 [*K. minus; OD; holotype, osAD-
chAYA & others, 1979, pl. 25,5, VSEGEI 11594,
St. Petersburg] [=Borocyathus voronin, 1988,
p. 8 (type, B. khairkhanicus, OD)]. Inner wall
with several rows of simple pores per intersept;
septa aporose to sparsely porous; pectinate tabulae
may be present. lower Cambrian (Atd.2–Bot.1):
Sib erian Platform, Altay Sayan, Mongolia, Far
East.——Fig. 34,3. *K. minus, Usa Formation,
Atdabanian, Krutoy Log, Batenev Range, Kuznetsk
Alatau, Russia, holotype, VSEGEI 11594, trans-
verse section, ×15 (Osadchaya & others, 1979).
Plicocyathus vologDin, 1960, p. 424 [*P. krassnyi; OD;
holotype, vologDin, 1960, fig. 1m, PIN 4754/45,
Moscow] [=Tumulocyathellus Zhur AvlevA, 1960b,
p. 174, nom. transl. rePinA & others, 1964, p. 194,
ex Tumulocyathus (Tumulocyathellus) ZhurAvle vA,
1960b, p. 174 (type, Tumulocyathus admirabilis
vologDin, 1940b, p. 72, OD); for discussion, see
Debrenne, ZhurAvlev, & roZAnov, 1989, p. 123;
Debrenne, roZAnov, & ZhurAvlev, 1990, p. 155;
=Torosocyathellus osADchAYA in osADchAYA & others,
1979, p. 128 (type, T. torosus, OD)]. Outer wall
longitudinally plicate; inner wall with stirrup pores
only; septa aporose to sparsely porous; pectinate
2
1b
1a Tegerocyathus
Krasnopeevaecyathus
Fig. 33. Tegerocyathidae (p. 43).
I did not
separate
these two
genera into
two separate
figures, as
that would
necessitate
renumbering
from here
to the end,
both the
figures and
the text.

Systematic Descriptions of the Archaeocyatha 47
Fig. 34. Tumulocyathidae (p. 43–47).
2a
3
1
4b
2b
4c
4a
Tumulocyathus
Isiticyathus
Kotuyicyathellus
Plicocyathus
tabulae may be present. lower Cambrian (Atd.1–
Bot.3): Siberian Platform, Kolyma, Altay Sayan, Tuva,
Mongolia, Transbaikalia, Far East, Morocco, Iberia,
Canada, United States, Mexico.——Fig. 34,4a–c. *P.
krassnyi; a, Ust’toka unit, Botoman, Gerbikan River,
Dzhagdy Range, Far East, Russia, holotype, PIN
4754/45, sketch of transverse section, ×5 (Vologdin,
1960); b–c, Ust’toka unit, Botoman, Onnetok River,
Dzhagdy Range, Far East, Russia, specimen DVGU
6M; b, transverse section, ×10; c, oblique transverse
section, ×10 (Belyaeva & others, 1975).
Family SANARKOCYATHIDAE
Hill, 1972
[Sanarkocyathidae hill, 1972, p. 79] [=Sanaricyathidae roZAnov, 1969,
p. 107, name based on invalid generic name Sanaricyathus roZA nov,
1969, p. 108, nom. null. pro Sanarkocyathus ZhurAvlevA, 1963a, p. 118]
Inner wall with bracts or scales. lower
Cambrian (Atd.3–Bot.1).
Sanarkocyathus Zhu rAv le vA , 1963a, p. 118 [*S.
mamaevi; OD; holotype, Zh ur Av le vA, 1963a,

48 Treatise Online, number XX
fig. 2, TsSGM 99/1, Novosibirsk] [=Sanaricyathus
roZAnov, 1969, p. 108, nom. null. pro Sanarkocy-
athus ZhurAvlevA, 1963a, p. 118]. Inner wall with
one row of pores per intersept, bearing possibly
upwardly projecting, S-shaped scales; septa aporose
to sparsely porous. lower Cambrian (Bot.1): Urals,
Altay Sayan.——Fig. 35,1. *S. mamaevi, Sanarka
Formation, Botoman, Sanarka River, eastern flank
of southern Urals, Russia, holotype, TsSGM 99/1,
oblique transverse section, ×6 (Zhuravleva, 1963a).
Neokolbicyathus KonYAevA in ZhurAvlevA & others,
1997b, p.131 [*N. azhuravlevi; OD; holotype,
Zhur Avl evA & others, 1997b, pl. 4,3, ZSGGU
2329/83, Novokuznetsk]. Inner wall with stirrup
pores only, bearing upwardly projecting, S-shaped
scales; septa aporose to sparsely porous. l ower
Cambrian (Atd.4–Bot.1): Altay Sayan, Far East,
Canada.——Fig . 35,2. *N. azhuravlevi, Usa
Formation, Atdabanian, Malaya Belokamenka
River, Kiya River, Kuznetsk Alatau, Russia, holo-
type, ZSGGU 2329/83, oblique transverse section,
×8 (Zhuravleva & others, 1997b).
Ringifungia Korshunov in ZhurAvlevA, Korshunov,
& roZ Anov, 1969, p. 38 [*R. vavilovi; OD; holo-
type, Zh ur Av l evA , Kor sh u n ov , & ro ZA n ov ,
1969, pl. 10,4–5, TsSGM 323/47, Novosibirsk].
Inner wall with one row of pores per intersept,
bearing upwardly projecting, S-shaped scales; septa
completely porous, linked by synapticulae. lower
Cambrian (Atd.3): Siberian Platform.——Fig .
35,3. *R. vavilovi, Perekhod Formation, Atdaba-
nian, Ulakhan-Taryng Creek, Lena River, Sakha
(Yakutia), Russia, holotype, TsSGM 323/47,
oblique transverse section, ×8 (Zhuravleva, Kors-
hunov, & Rozanov, 1969).
Family GEOCYATHIDAE
Debrenne, 1964
[Geocyathidae Debrenne, 1964, p. 114] [=Jakutocyathidae Korshunov,
1972, p. 65; =Eladicyathidae Perejón, 1977, p. 550]
Inner wall with annuli. lower Cambrian
(Atd.1–Bot.1).
Geocyathus ZhurAvlevA, 1960b, p. 234 [*Thalamocy-
athus botomanensis ZhurAvlevA in ZhurAvlevA &
Zelenov, 1955, p. 71; OD; holotype, ZhurAvlevA
& Zelenov, 1955, pl. 2,3–4, TsSGM 205/115a-b,
Novosibirsk; =T. botomaensis Zhu rAvlevA, 1960b,
p. 234, nom. null.] [=Jakutocyathus (Jakutocyathus)
Zhur Avle vA, 1960b, p. 230 (type, J. (J.) latini,
OD); =Eladicyathus Perejón, 1977, p. 550 (type,
E. beticus, OD); for discussion, see Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 108; Debrenne,
roZ An ov , & Zh ur Avl ev , 1990, p. 144]. Inner
wall with one pore row per intersept and upwardly
projecting, S-shaped annuli; septa completely
porous; pectinate tabulae may be present. lower
Cambrian (Atd.1–Bot.1): Siberian Platform, Altay
Sayan, Transbaikalia, Far East, Iberia.——Fig .
35,4a–b. *G. botomanensis (ZhurAvlevA), Perekhod
Formation, Atdabanian, Botoma River, Sakha
(Yakutia), Russia, holotype, TsSGM 205/115a-b;
a, transverse section, ×15; b, longitudinal section,
×15 (Zhuravleva & Zelenov, 1955).
Family KONJUSCHKOVICYATHIDAE
Debrenne & Zhuravlev, 2000
[Konjuschkovicyathidae Debrenne & ZhurAvlev, 2000, p. 49]
Inner wall with noncommunicating
canals. lower Cambrian (Bot.1–Bot.3).
Konjuschkovicyathus Debrenne & ZhurAvlev, 2000,
p. 49 [*Jakutocyathus spinosus KonYushKov, 1972,
p. 140; OD; holotype, Ko nY us hK ov, 1972, pl.
14,6, not located; paratypes, KonYushKov, 1972,
pl. 14,5, PIN 4755/5; Ko n Yu sh Kov , 1972, pl.
16,3, PIN 4755/6, Moscow]. Inner wall with
downwardly projecting, straight stirrup canals only,
bearing supplementary bracts on central cavity side;
septa aporose to sparsely porous; pectinate tabulae
may be present. lower Cambrian (Bot.1–Bot.3):
Altay Sayan, Transbaikalia.——Fig. 35,5a–b. *K.
spinosus (KonYushKov), Verkhnemonok Formation,
Botoman, Malyy Karakol River, western Sayan,
Altay Sayan, Russia; a, holotype, transverse section,
×10; b, paratype, PIN 4755/6, oblique longitu-
dinal section, ×10 (Konyushkov, 1972).
Superfamily LENOCYATHOIDEA
Zhuravleva, 1956
[nom. correct. Debrenne, ZhurAvlev, & roZAno v, 1989, p. 83, pro
Lenocyathacea ZhurAvlevA, 1960b, p. 224, nom. transl. ex Lenocyathidae
ZhurAvlevA in vologDin, 1956, p. 879] [=Rewardocyathacea roZAnov,
1973, p. 86, nom. nud.]
Outer wall with multiperforate tumuli.
lower Cambrian (Atd.1–Bot.1).
Family TOROSOCYATHIDAE
Debrenne, Zhuravlev, & Kruse, 2002
[Torosocyathidae Debrenne, ZhurAvlev, & Kruse, 2002, p. 1594] [=Re-
wardocyathidae roZAnov, 1973, p. 86, nom. nud.; =Rewardocyathidae
roZAnov in Debrenne, ZhurAvlev, & roZAnov, 1989, p. 83, nom. nud.,
based on unavailable genus-group name]
Inner wall with simple pores. l o wer
Cambrian (Atd.1–Bot.1).
Torosocyathus KAshinA in vologDin & KAshinA,
1972, p. 153 [*T. provisus; OD; holotype, vologDin
& KAshinA, 1972, pl. 20a,1, KGU 19/729a, Kras-
noyarsk] [=Rewardocyathus roZAnov, 1973, p. 59,
75, 161, nom. nud.]. Inner wall with stirrup pores
only; septa completely porous. lower Cambrian
(Atd.1–Bot .1): Altay Sayan, Mongolia.——F ig .
36,1. *T. provisus, Balakhtinson Formation, Atda-
banian, Uyar River, eastern Sayan, Altay Sayan,
Russia, holotype, KGU 19/729a, detail of trans-
verse section, ×12 (Vologdin & Kashina, 1972).
Torosocyathella KoTel’ni Kov, 1995, p. 27 [*T. osad-
chajae; OD; holotype, KoTel’ni Kov, 1995, pl. 2,5,
TsNIGRm 12890/9, St. Petersburg]. Inner wall
with several rows of simple pores per intersept;

Systematic Descriptions of the Archaeocyatha 49
Fig. 35. Sanarkocyathidae, Geocyathidae, and Konjuschkovicyathidae (p. 47–48).
2
5a
5b
1
4b
3
4a
Sanarkocyathus
Neokolbicyathus
Ringifungia
Geocyathus Konjuschkovicyathus

50 Treatise Online, number XX
septa completely porous. lower Cambrian (Atd.2):
Tuva.——Fig . 36,2. *T. osadchajae, Il’chir Forma-
tion, Atdabanian, Vadi-Bala, Tapsa River, Tuva,
Russia, holotype, TsNIGRm 12890/9, transverse
section, ×20 (Kotel’nikov, 1995).
Family JAPHANICYATHIDAE
Rozanov, 1989
[Japhanicyathidae roZAnov in Debrenne, ZhurAvlev, & roZAnov, 1989,
p. 83] [=Japhanicyathidae roZAnov, 1973, p. 86, nom. nud.]
Inner wall with annuli. lower Cambrian
(Atd.2–Bot.1).
Japhanicyathus Korshunov i n Zh u r Av l e v A ,
Korshunov, & roZAnov, 1969, p. 45 [*J. genu-
rosus; OD; holotype, ZhurAvlevA, Korshunov, &
roZAnov, 1969, pl. 17,1–2, TsSGM 323/67, Novo-
sibirsk]. Inner wall with one pore row per intersept
and upright, V-shaped annuli; septa completely
porous; pectinate tabulae may be present. lower
Cambrian (Atd.2–Bot.1): Siberian Platform, Far
East.——Fig. 36,3. *J. genurosus, Oy-Muran reef
massif, Atdabanian, Oy-Muran, Lena River, Sakha
(Yakutia), Russia, holotype, TsSGM 323/67, trans-
verse section, ×8 (Zhuravleva, Korshunov, &
Rozanov, 1969).
Family LENOCYATHIDAE
Zhuravleva, 1956
[Lenocyathidae ZhurAvlevA in vologDin, 1956, p. 879]
Inner wall with noncommunicating
canals. lower Cambrian (Atd.2–Bot.1).
Lenocyathus ZhurAvlevA in ZhurAvlevA & Zelenov,
1955, p. 73 (Zh u rAv l evA , 1954, p. 12, nom.
nud.) [*L. lenaicus; OD; holotype, ZhurAvlevA &
Zelenov, 1955, pl. 2,5–6, TsSGM 205/117, Novo-
sibirsk]. Inner wall with one row of horizontal to
upwardly projecting, S-shaped canals per intersept;
septa completely porous; pectinate tabulae may be
present. lower Cambrian (Atd.2–Bot.1): Siberian
Platform, Far East, Morocco.——Fig. 36,4a–b. *L.
lenaicus, Pestrotsvet Formation, Atdabanian, Yudyay,
Botoma River, Sakha (Yakutia), Russia, holotype,
TsSGM, 205/117; a, longitudinal section, ×7; b,
detail of transverse section (outer wall at top), ×20
(Debrenne, Zhuravlev, & Kruse, 2002).
Superfamily
ANNULOCYATHOIDEA
Krasnopeeva, 1953
[nom. correct. Debrenne, ZhurAvlev, & roZAnov, 1989, p. 83, pro Annu-
locyathacea ZhurAvlevA, 1960b, p. 171, nom. transl. ex Annulocyathidae
KrAsnoPeevA, 1953, p. 56]
Outer wall with bracts or scales. lower
Cambrian (Tom.2–Bot.3).
Family TUMULIFUNGIIDAE
Rozanov, 1989
[Tumulifungiidae roZAnov in Debrenne, ZhurAvlev, & roZAnov, 1989,
p. 83] [=Tumulifungiidae roZAnov, 1973, p. 85, nom. nud.]
Inner wall with simple pores. l o wer
Cambrian (Tom.2–Bot.3).
Tumulifungia ZhurAvlevA in DATsenKo & others,
1968, p. 144 (Zh u rAv l ev A in Zhur Av l ev A &
others, 1967, p. 68, nom. nud.) [*T. datzenkoi;
OD; holotype, DATs en Ko & others, 1968, pl.
4,2–3, TsSGM 277/30, Novosibirsk]. Outer wall
with upwardly projecting cu pp ed bracts; inner
wall with one row of simple pores per intersept;
septa completely porous, linked by synapticulae.
lower Cambrian (Atd.1–Bot.3): Siberian Platform,
Altay Sayan, Tuva, Mongolia, Far East, Morocco,
Iberia.——Fig . 37,1. *T. datzenkoi, Shum noy
Formation, Botoman, Sukharikha River, Kras-
noyarsk region, Russia, holotype, TsSGM 277/30,
transverse section, ×11 (Datsenko & others, 1968).
Sclerocyathus vologDin, 1960, p. 424 [*S. scrofu -
losus; OD; holotype, vologDin, 1960, fig. 1z-i,
PIN 4754/1, Moscow]. Outer wall with upwardly
projecting, cupped bracts; inner wall with one row
of simple pores per intersept; septa completely
porous. lower Cambrian (Tom.2–Bot.1): Siberian
Platform, Altay Sayan, Tuva, Mongolia, Far East,
Iberia.——Fig. 37,2a–b. *S. scrofulosus, Bayan-Kol
Formation, Atdabanian, Yenisey River, Shagonar
Mountains, Tuva, Russia, holotype, PIN 4754/1;
a, transverse section, ×4; b, detail of transverse
section, ×8 (Debrenne, Zhuravlev, & Kruse, 2002).
Subtumulocyathellus osADchAYA in osADchAYA &
others, 1979, p. 129 [*S. vulgaris; OD; holotype,
osA Dc hAY A & others, 1979, pl. 11,1, VSEGEI
11594, St. Petersburg] [=Arturocyathus ro ZAnov ,
1973, p. 61, 162, nom. nud. ; =Arturocyathus
roZAnov in Debrenne, ZhurAvle v, & ro ZAnov ,
1989, p. 95 (type, A. borisovi ro ZAno v, 1973, p.
162, OD)]. Outer wall with upwardly projecting,
cupped bracts; inner wall with stirrup pores only;
septa aporose to sparsely porous. lower Cambrian
(At d.1–Bot.1): Siberian Platform, Altay Sayan,
Tuva, Mongolia, Far East.——Fig . 37,3 . *S.
vulgaris, Usa Formation, Atdabanian, Krutoy Log,
Batenev Range, Kuznetsk Alatau, Russia, holotype,
VSEGEI 11594, oblique transverse section, ×10
(Osadchaya & others, 1979).
Tologoicyathus voronin, 1988, p. 9 [*T. ichituin-
icus; OD; holotype, voronin, 1988, pl. 2,3, PIN
3301/516, Moscow]. Outer wall with upwardly
projecting, cupped bracts; inner wall with several
rows of simple pores per intersept; septa completely
porous. lower Cambrian (Tom.4–Bot.1): Mongolia,
Far East. ——F i g. 37,4. *T. ich ituin icu s, Ichi-
tuin Formation, Atdabanian, Boro-Khairkhan-
Obo Mountain, Khan-Khukhiy Range, Mongolia,
paratype, PIN 3301/515, transverse section, ×5
(Voronin, 1988).

Systematic Descriptions of the Archaeocyatha 51
Fig. 36. Torosocyathidae, Japhanicyathidae, and Lenocyathidae (p. 48–50).
2
1
4b
3
4a
Torosocyathus
Torosocyathella
Japhanicyathus
Lenocyathus

52 Treatise Online, number XX
Family ANNULOCYATHIDAE
Krasnopeeva, 1953
[Annulocyathidae KrAsnoPeevA, 1953, p. 56]
Inner wall with annuli. lower Cambrian
(Atd.2–Bot.3).
Annulocyathus vologDin, 1937b, p. 468 [*A. pulcher;
M; lectotype, Debr en ne, Zhu rAvl ev, & Kru se,
2002, fig. 32E; SD Debrenne, ZhurAvlev, &
Krus e, 2002, p. 1597, PIN 4754/5, Moscow].
Oute r wa ll with upwardly proje cti ng, cup ped
bracts; inner wall with one pore row per intersept
and upright V-shaped annuli; septa completely
porous. lower Cambrian (Bot.1): Altay Sayan, Far
East.——Fig. 38,1a–b. *A. pulcher, Verkhnemonok
Formation, Botoman, Sanashtykgol River, western
Sayan, Altay Sayan, Russia, lectotype, PIN 4754/5;
Fig. 37. Tumulifungiidae (p. 50).
Sclerocyathus
2a
4
1
2b
3
Tumulifungia
Subtumulocyathellus
Tologoicyathus

Systematic Descriptions of the Archaeocyatha 53
a, transverse section, ×11 (Debrenne, Zhuravlev,
& Kruse, 2002); b, sketch of longitudinal section
(outer wall to left), ×5.5 (Vologdin, 1937b).
Annulocyathella vologDin, 1962a, p. 123 [*Annulo-
cyathus lavrenovae KrAsnoPeevA, 1955, p. 99; OD;
holotype, KrAsnoPeevA, 1955, pl. 3,2; vologDin,
1962a, fig. 86, not designated; =Anulocyathus lavre-
novi KrAsnoPeevA, 1937, p. 33; holotype, KrAsnoP-
eevA, 1937, pl. 4,38–39,41,43–44; pl. 16,109; pl.
19,118, not designated]. Outer wall with upwardly
projecting, cupped bracts; inner wall with one
pore row per intersept and upwardly projecting,
S-shaped annuli; septa aporose to sparsely porous.
lower Cambrian (Atd.4–Bot.3): Altay Sayan.——
Fig. 38,2a–b. *A. lavrenovae (KrA sno PeevA), Usa
Formation, Botoman, Bol’shaya Erba, Batenev
Range, Kuznetsk Alatau, Russia; a, unlocated
syntype, oblique transverse section, ×8; b, unlo-
cated syntype, sketch of longitudinal section (outer
wall to left), ×8 (Krasnopeeva, 1955).
Annulofungia KrAsnoPeevA, 1955, p. 99 (KrAs-
noPe evA, 1953, p. 56, nom. nud.) [*Anulocyathus
taylori KrA snoPe evA, 1937, p. 34; OD; holotype,
KrAsnoPe evA, 1937, pl. 4,46–47; pl. 18,115–116;
pl. 22,130; pl. 24,137, not designated, collection
not located] [=Kiyafungia boYArinov in ZhurAv-
levA & others, 1997b, p. 130 (type, K. concinna,
OD)]. Outer wall with upwardly projecting,
cupped bracts; inner wall with one pore row per
intersept and upright, V-shaped annuli; septa
completely porous, linked by synapticulae. lower
Cambrian (Atd.4–Bot.1): Altay Sayan.——Fig.
38,3a–b. *A. taylori (KrA sn oPee vA), Usa Forma-
tion, Botoman, Mt. Aydachikha, Batenev Range,
Kuznetsk Alatau, Russia, unlocated specimen; a,
longitudinal section, ×6; b, transverse section, ×6
(Debrenne, Zhuravlev, & Kruse, 2002).
Hemithalamocyathus Ting, 1937, p. 367 [*Archaeo-
cyathus sibiricus Toll, 1899, p. 40; M; lectotype,
Toll, 1899, pl. 6,5; SD Debrenne, Zh urAvlev, &
Kruse, 2002, p. 1598, TsNIGRm 24a/11533, St.
Petersburg]. Outer wall with upwardly projecting,
cupped bracts; inner wall with several pore rows
per intersept and upright, V-shaped annuli; septa
completely porous. lower Cambrian (Atd.4–Bot.1):
Altay Sayan.——Fig . 38,4. *H. sibiricus (Toll),
Torgashino Formation, Torgashino, Krasnoyarsk
region, eastern Sayan, Altay Sayan, Russia, unnum-
bered paralectotype, oblique longitudinal section
(outer wall to left), ×10 (Toll, 1899).
Family JAKUTOCARINIDAE
Debrenne, Rozanov, & Zhuravlev, 1989
[Jakutocarinidae Debrenne, roZAnov , & ZhurAvl ev in Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 83]
Inner wall with noncommunicating
canals. lower Cambrian (Atd.1–Bot.3).
Jakutocarinus ZhurAvlevA, 1960b, p. 232 [*Jakuto-
cyathus (Jakutocarinus) jakutensis; OD; holotype,
Zhur Avle vA, 1960b, pl. 20,2, TsSGM 205/113,
Novosibirsk]. Outer wall with upwardly projecting,
cupped bracts; inner wall with several rows of
downwardly projecting, straight canals per inter-
sept, bearing supplementary bracts on central cavity
side; septa completely porous; pectinate tabulae may
be present. lower Cambrian (Atd.1–Bot.1): Siberian
Platform, Altay Sayan, Tuva, Mongolia, Trans-
baikalia.——Fig. 39,1. *J. jakutensis, Pestrotsvet
Formation, Atdabanian, Mukhatta River, Sakha
(Yakutia), Russia, holotype, TsSGM 205/113,
detail of oblique transverse section (outer wall at
top), ×15 (Debrenne, Zhuravlev, & Kruse, 2002).
Kosticyathus Debrenne & ZhurAvlev, 2000, p. 49
[*Porocyathus sheglovi KonYush Kov, 1972, p. 138;
OD; holotype, KonYushKov, 1972, pl. 16,4–5,
PIN 4755/9, Moscow]. Outer wall with upwardly
projecting, cupped bracts; inner wall with one
row of downwardly projecting, straight canals
per intersept, bearing supplementary bracts on
central cavity side; septa completely porous. lower
Cambrian (Bot.1–Bot.3): Altay Sayan.——Fig .
39,2. *K. sheglovi (KonYushKov), Verkhnemonok
Formation, Botoman, Malyy Karakol River, western
Sayan, Altay Sayan, Russia, holotype, PIN 4755/9,
longitudinal section, ×6 (Konyushkov, 1972).
Kruseicyathus boYAr ino v & Kon YAevA in Zhu rAv-
levA & others, 1997b, p. 134 [*K. notabilis; OD;
holotype, ZhurAvlevA & others, 1997b, pl. 4,5,
ZSGGU 2329/86, Novokuznetsk]. Outer wall
with upwardly projecting, cupped bracts; inner wall
with horizontal to upwardly projecting, S-shaped
canals, each canal spanning several intersepts;
septa completely porous. lower Cambrian (Bot.1):
Altay Sayan.——Fig. 39,3a–b. *K. notabilis, Usa
Formation, Botoman, Malaya Belokamenka River,
Kiya River, Kuznetsk Alatau, Russia; a, paratype,
ZSGGU 2329/85, transverse section, ×10; b,
holotype, ZSGGU 2329/86, oblique longitudinal
section, ×10 (Zhuravleva & others, 1997b).
Rossocyathella Zhu rAv l evA , 1960b, p. 178 [*R.
ninaekosti; OD; holotype, ZhurAvlevA, 1960b, pl.
12,5, PIN 1038, Moscow, not located]. Outer wall
with upwardly projecting, cupped bracts; inner wall
with one row of downwardly projecting, straight
canals per intersept, bearing supplementary bracts
on central cavity side; septa aporose to sparsely
porous. lower Cambrian (Bot.1): Siberian Platform,
Altay Sayan, Tuva.——Fig. 39,4a–b. *R. ninaekosti,
Perekhod Formation, Botoman, Botoma River,
Sakha (Yakutia), Russia, holotype, PIN 1038; a,
sketch of longitudinal section (outer wall to left),
×40; b, detail of transverse section (outer wall at
bottom), ×15 (Zhuravleva, 1960b).
?Russocyathus ZhurAvlevA, 1955b, p. 628 [*R. basa-
ichensis; OD; holotype, Zhu rAvle vA, 1955b, fig.
1E, 2v; rePinA & others, 1964, pl. 19,2, PIN 1039,
Moscow, not located]. Outer wall with probable
upwardly projecting, cupped bracts; inner wall
with one row of probable horizontal to upwardly
projecting, S-shaped canals per intersept; septa
aporose to sparsely porous. [Limited type material
does not provide certainty as to orientation of cup
and hence as to presence or absence of canals and/

54 Treatise Online, number XX
Fig. 38. Annulocyathidae (p. 52–53).
3a
3b
2b
2a
Annulocyathella
Annulofungia 4Hemithalamocyathus
1b
1a
Annulocyathus

Systematic Descriptions of the Archaeocyatha 55
Fig. 39. Jakutocarinidae and Gagarinicyathidae (p. 53–56).
2
5
3b
4a
4b
6
1
3a
Jakutocarinus
Kosticyathus
Kruseicyathus
Rossocyathella
Russocyathus Gagarinicyathus

56 Treatise Online, number XX
or bracts in walls.] lower Cambrian (Atd.3–Bot.1):
Altay Sayan, Tuva.——Fig. 39,5. *R. basaichensis,
Torgashino Formation, Atdabanian, Torgashino,
Krasnoyarsk region, eastern Sayan, Altay Sayan,
Russia, holotype, PIN 1039, sketch of oblique
longitudinal section, ×20 (Zhuravleva, 1955b).
Family GAGARINICYATHIDAE
Debrenne, Rozanov, & Zhuravlev, 1989
[Gagarinicyathidae Debrenne, ro ZAnov, & ZhurAvlev, in Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 84]
Inner wall with communicating canals.
lower Cambrian (Atd.3–Bot.1).
Gagarinicyathus ZhurAvlevA in DATsen Ko & others,
1968, p. 146 [*G. ethmophylloides; OD; holotype,
DATsenKo & others, 1968, pl. 5,1, TsSGM 277/36,
Novosibirsk]. Outer wall with upwardly projecting,
cupped bracts; inner wall with one row of horizontal
to upwardly projecting, straight porous canals per
intersept; septa completely porous. lower Cambrian
(Atd.3–Bot.1): Siberian Platform.——Fig. 39,6. *G.
ethmophylloides, Shumnoy Formation, Botoman,
Sukharikha River, Krasnoyarsk region, Russia, holo-
type, TsSGM 277/36, oblique longitudinal section,
×4.5 (Datsenko & others, 1968).
Superfamily
ETHMOPHYLLOIDEA
Okulitch, 1937
[nom. transl. ZhurAvlev in voronovA & others, 1987, p. 23, ex Ethmo-
phyllidae oKuliTch, 1937b, p. 358] [=Carinacyathoidea KrAsnoPeevA,
1953, p. 52, nom. transl. ZhurAvlev in voronovA & others, 1987, p.
23, ex Carinacyathidae ZhurAvlevA, 1960b, p. 240, nom. correct. pro Ca-
rinocyathidae KrAsnoPeevA, 1953, p. 52; =Fansycyathacea Korshunov &
roZAnov in ZhurAvlevA, Korshunov, & roZAnov, 1969, p. 46; =Hupe-
cyathelloidea roZAnov, 1969, p. 111, nom. correct. Debrenne, ZhurAvlev,
& roZAnov, 1989, p. 84, pro Hupecyathellacea roZAnov, 1969, p. 111]
Outer wall with canals. lower Cambrian
(Atd.1–Toy.1).
Family FALLOCYATHIDAE
Rozanov, 1969
[Fallocyathidae roZAnov in ZhurAvlevA, Korshunov, & roZAnov, 1969,
p. 47] [=Sekwicyathidae roZAnov, 1973, p. 85, nom. nud.]
Inner wall with simple pores. l o wer
Cambrian (Bot.1–Bot.2).
Fallocyathus roZAnov in ZhurAvlevA, Korshunov, &
roZAnov, 1969, p. 47 [*F. dubius; OD; holotype,
Zhu rAv l evA , Korshunov, & roZA no v, 1969,
pl. 18,5–6; pl. 19,2, PIN 4297/84, Moscow].
Outer wall with horizontal to upwardly projecting,
straight canals, bearing supplementary bracts exter-
nally (imparting overall inverted V-shaped appear-
ance to outer wall); inner wall with several rows of
simple pores per intersept; septa completely porous;
pectinate tabulae may be present. lower Cambrian
(Bot.1): Siberian Platform, Iberia.——Fig. 40,1.
*F. dubius, Oy-Muran reef massif, Atdabanian,
Oy-Muran, Lena River, Sakha (Yakutia), Russia,
holotype, PIN 4297/84, oblique transverse section,
×12 (Zhuravleva, Korshunov, & Rozanov, 1969).
Sekwicyathus hAnDFielD, 1971, p. 34 [*S. nahan-
niensis; OD; holotype, hAnDFielD, 1971, p. 34,
pl. 2,5, GSC 25317, Ottawa; ?=Archaeocyathus
nevadensis oKuliTch, 1935, p. 101]. Outer wall
with subspherical chambered canals each with base
commencing in intervallum, canals subdivided
by stipules (imparting overall inverted V-shaped
appearance to outer wall); inner wall with several
rows of simple pores per intersept; septa aporose
to sparsely porous. lower Cambrian (Bot.1–Bot.2):
Altay Sayan, Iberia, Canada, United States.——
Fig. 40,2a–c. *S. nahanniensis; a, Sekwi Forma-
tion, Botoman, Mackenzie Mountains, North-
west Territories, Canada, holotype, GSC 25137,
oblique longitudinal section, ×15 (Handfield,
1971); b, Atan Group, Botoman, Gataga River,
British Columbia, Canada, specimen GSC 69260,
transverse section, ×10 (Debrenne, Zhuravlev, &
Kruse, 2002); c, Sekwi Formation, Botoman, Mack-
enzie Mountains, Northwest Territories, Canada,
specimen GSC 90132, longitudinal section, ×10
(Voronova & others, 1987).
Yukonocyathus hAnDFielD, 1971, p. 51 [*Y. francesi;
OD; holotype, hAnDFielD, 1971, pl. 8,1a–c, GSC
25351, Ottawa]. Outer wall with horizontal to
upwardly projecting, S-shaped canals, bearing
supplementary bracts externally (imparting overall
inverted V-shaped appearance to outer wall); inner
wall with one row of simple pores per intersept,
formed by fluting of inner edges of septa; septa
aporose to sparsely porous. lower Cambrian (Bot.1):
Canada, United States.——Fig. 40,3a–b. *Y. fran-
cesi, Sekwi Formation, Botoman, Frances Lake,
Yukon Territory, Canada, holotype, GSC 25351;
a, transverse section, ×6; b, longitudinal section
(outer wall to left), ×6 (Handfield, 1971).
Family GLORIOSOCYATHIDAE
Rozanov, 1969
[Gloriosocyathidae roZAnov, 1969, p. 108]
Inner wall with bracts or scales. lower
Cambrian (Atd.1–Bot.1).
Gloriosocyathus roZ A n o v , 1969, p. 108 [*G .
permultus; OD; holotype, ro ZA no v, 1969, pl.
40,3, PIN 4297/95, Moscow]. Outer wall with
horizontal to upwardly projecting, S-shaped
canals; inner wall with one row of pores per inter-
sept, bearing upwardly projecting, S-shaped scales;
septa completely porous. lower Cambrian (Bot.1):
Siberian Platform, Iberia.——Fi g . 41,1. *G.
permultus, Erkeket Formation, Botoman, Khor-
busuonka River, Olenek Basin, Sakha (Yakutia),
Russia, holotype, PIN 4297/95, oblique transverse
section, ×10 (Debrenne, Zhuravlev, & Kruse,
2002).
Gandinocyathus F. Debrenne & M. Debrenne in
gAnDin, F. Debrenne, & M. Debrenne, 2007, p.
41 [*G. gravestocki, OD; holotype, F. Debrenne,

Systematic Descriptions of the Archaeocyatha 57
Yukonocyathus
Fig. 40. Fallocyathidae (p. 56).
2a
3b
2c
2b
1
3a
Fallocyathus
Sekwicyathus

58 Treatise Online, number XX
gAnDin, & m. Debrenne, 1993, pl. 3,1, MNHN
M84234, Paris]. Outer wall with horizontal to
upwardly projecting, straight canals, bearing
supplementary bracts externally (imparting overall
inverted V-shaped appearance to outer wall);
inner wall with one row of pores per intersept,
bearing upwardly projecting, cupped bracts;
septa completely porous. lower Cambrian (Bot.1):
Sardinia.——Fig. 41,2. *G. gravestocki, Matoppa
Formation, Botoman, Matoppa Valley, Sardinia,
Italy, holotype, MNHN M84234, oblique longi-
tudinal section, ×10 (F. Debrenne, Gandin, & M.
Debrenne, 1993).
Nalivkinicyathus boYArinov & osADchAYA in osAD-
chAYA & gAn Ac hK ovA , 1986, p. 170 [*Porocy-
athellus cyroflexus boY Ar i nov & osA Dc hA YA in
osADchAYA & others, 1979, p. 132; OD; holotype,
osADchAYA & others, 1979, pl. 8,1–2; osAD chAYA
& gAnAchKovA, 1986, pl. 18,1–2, VSEGEI 11594,
St. Petersburg] [=Nalivkinicyathus osADchAYA in
Debrenne & roZAnov, 1983, p. 735, nom. nud.,
nom. nov. pro Porocyathellus boYArin ov & osAD-
chAYA in osADc hAYA & others, 1979, p. 131, non
Debrenne, 1977a, p. 107, archaeocyath]. Outer
wall with downwardly projecting, straight canals,
bearing supplementary bracts externally (imparting
overall upright V-shaped appearance to outer wall);
inner wall with one row of pores per intersept,
bearing upright, V-shaped scales; septa completely
porous. lower Cambrian (Atd.1–Bot.1): Altay Sayan,
Iberia.——Fig. 41,3a–b. *N. cyroflexus (boYA -
rinov & osADchAYA), Usa Formation, Atdabanian,
Krutoy Log, Batenev Range, Kuznetsk Alatau,
Russia, holotype, VSEGEI 11594; a, transverse
section, ×4; b, detail of septum in longitudinal
section (outer wall to right), ×12 (Osadchaya &
others, 1979).
Family KIJACYATHIDAE
Zhuravleva, 1964
[Kijacyathidae ZhurAvl evA in rePinA & others, 1964, p. 195] [=Fan-
sycyathidae Korshunov & roZAnov in ZhurAvle vA, Korshunov, &
roZAnov, 1969, p. 47]
Inner wall with annuli. lower Cambrian
(Atd.2–Bot.3).
Kijacyathus ZhurAvl evA, 1959, p. 424 [*K. chomen-
tovskii; OD; holotype, Zh ur Avl evA , 1959, fig.
2b–g, PIN 1431, Moscow, not located]. Outer
wall with horizontal to upwardly projecting,
S-shaped canals; inner wall with on e pore row
per intersept and upright, V-shaped annuli; septa
completely porous. lower Cambrian (Atd.3–
Bot.1): Altay Sayan, Mongolia, Far East.— —
Fig. 42,1a–b. *K. chomentovskii, Usa Formation,
Atdabanian, Kiya River, Kuznetsk Alatau, Russia,
holotype, PIN 1431; a, transverse section, ×8
(Debrenne, Zhuravlev, & Kruse, 2002); b, detail
of septum in longitudinal section (outer wall to
left), ×13 (Zhuravleva, 1959).
Aporosocyathus Krus e , 1978, p. 32 [*A. mucro-
porus; OD; holotype, Kru s e, 1978, fig. 4A–B,
AM FT.15203, 15204, Sydney]. Outer wall with
horizontal to upwardly projecting, S-shaped canals,
bearing supplementary bracts externally (imparting
overall inverted V-shaped appearance to outer wall);
inner wall with one pore row per intersept and
upright, V-shaped annuli; septa aporose to sparsely
porous; pectinate tabulae may be present. lower
Cambrian (Bot.2–Bot.3): ?Mongolia, Australia,
Antarctica, ?Canada.——Fig. 42,2a–b. *A. mucro-
porus, Cymbric Vale Formation, Botoman, Mt.
Wright, New South Wales, Australia, holotype, AM
FT.15203, 15204; a, oblique transverse section,
AM FT.15204, ×8 (Kruse, 1978); b, detail of longi-
tudinal section (outer wall to right) AM FT.15203,
×6 (Kruse, 1982).
Fansycyathus Korshunov & roZ An ov in Zh urAv-
le vA , Korshunov, & ro ZA n o v, 1969, p. 48
[*F. lermontovae; OD; holotype, Zh u rAv l evA ,
Korshunov, & roZ A n o v , 1969, pl. 17,7 ;
roZAnov, 1973, pl. 20,2, PIN 4297/83, Moscow].
Outer wall with horizontal to upwardly projecting,
S-shaped canals, bearing supplementary bracts
externally (imparting overall inverted V-shaped
appearance to outer wall); inner wall with one
pore row per intersept and upright, V-shaped
annuli; septa completely porous; pectinate tabulae
may be present. lower Cambrian (Atd.2–Bot.1):
Siberian Platform.——Fig. 42,3. *F. lermontovae,
Oy-Muran reef massif, Atdabanian, Oy-Muran,
Lena River, Sakha (Yakutia), Russia, holotype,
PIN 4297/83, longitudinal section, ×12 (Zhurav-
leva, Korshunov, & Rozanov, 1969).
Flexanulus Debrenne, 1975, p. 335 [*F. oosthui-
zeni; OD; holotype, Debrenne, 1975, fig. 3a–b,
SAM(C) K4495 B-12a, Cape Town]. Outer wall
with horizontal to upwardly projecting, S-shaped
canals, each with base commencing in inter-
vallum, bearing supplementary bracts externally
(imparting overall inverted V-shaped appearance
to outer wall); inner wall with one pore row per
intersept and upwardly projecting, S-shaped
annuli; septa aporose to sparsely porous. l ower
Cambrian (Bot.3): Antarctica, South Africa
(allochthonous).——Fig . 43,1a–b. *F. oosthui-
zeni, Dwyka Subgroup, Botoman (allochthonous
in Permian), Zwartskraal, South Africa, holotype,
SAM(C) K4495 B-12a; a, transverse section,
×10; b, longitudinal section, ×10 (Debrenne,
1975).
Protocyathus ForD, 1878, p. 124 [*P. rarus; M; holo-
type, ForD, 1878, fig. 1a–b, NYSM 52, Albany].
Outer wall with horizontal to upwardly projecting,
straight stirrup canals only; inner wall with stirrup
pores only, bearing horizontal planar to waved
annuli, one per several horizontal pore files; septa
aporose to sparsely porous. lower Cambrian (Bot.1–
Bot.2): Canada, United States.——Fig. 43,2a–b.
*P. rarus, Brown’s Pond Formation, Botoman, Troy,
New York, United States, holotype, NYSM 52; a,
detail of septum and inner wall in longitudinal
view (outer wall to left), ×15; b, longitudinal
view of outer wall, ×15 (Debrenne, Zhuravlev, &
Kruse, 2002).

Systematic Descriptions of the Archaeocyatha 59
Yudjaicyathus ZhurAvlev in ZhurAvlev, ZhurAvlevA,
& Fonin, 1983, p. 25 [*Y. astashkini; OD; holo-
type, Zhu rAv lev , Zh ur Avle vA, & Fonin, 1983,
pl. 4,1, PIN 3848/505, Moscow]. Outer wall
with horizontal to upwardly projecting, straight
canals, bearing supplementary bracts externally
(imparting overall inverted V-shaped appearance
to outer wall); inner wall with one pore row per
intersept and upwardly projecting, S-shaped annuli;
septa completely porous; pectinate tabulae may be
present. lower Cambrian (Atd.3–Atd.4): Siberian
Platform.——Fig. 43,3. *Y. astashkini, Pestrotsvet
Formation, Atdabanian, Bachyk Creek, Lena River,
Sakha (Yakutia), Russia, holotype, PIN 3848/505,
oblique longitudinal section, ×14 (Zhuravlev,
Zhuravleva, & Fonin, 1983).
2
3b
1
3a
Gloriosocyathus
Gandinocyathus
Nalivkinicyathus
Fig. 41. Gloriosocyathidae (p. 56–58).

60 Treatise Online, number XX
Fig. 42. Kijacyathidae (p. 58).
1a
3
2b
1b
Kijacyathus
Aporosocyathus
Fansycyathus
I reduced view
2a to X8 and
changed the text
too. Is that what
you requested?
2a

Systematic Descriptions of the Archaeocyatha 61
1a
2a
3
1b
Flexanulus
Protocyathus
Yudjaicyathus
2b
Fig. 43. Kijacyathidae (p. 58–59).

62 Treatise Online, number XX
Family CARINACYATHIDAE
Krasnopeeva, 1953
[nom. correct. ZhurAvlevA, 1960b, p. 240, pro Carinocyathidae KrAsnoPe-
evA, 1953, p. 56] [=Porocyathidae ZhurAvlevA in vologDin, 1957a, p.
179; Hupecyathellidae roZAnov, 1969, p. 111]
Inner wall with noncommunicating
canals. lower Cambrian (Atd.1–Bot.3).
Carinacyathus vologDin, 1932, p. 37 [*C. loculatus;
M; holotype, vologDin, 1932, fig. 28, pl. 10,5;
M; vologDin, 1940b, fig. 77, pl. 27,7, TsNIGRm
45a/2957, St. Petersburg] [=Carinocyathus
vologDin, 1937b, p. 471, nom. null.; =Porocyathus
ZhurAvlevA in vologDin, 1957a, p. 179, nom. nud.;
=Porocyathus ZhurAvlevA, 1960b, p. 180 (type, P.
pinus, OD); =Fossilicyathus Korshunov, 1983b,
p. 111 (type, F. evidens, OD), for discussion, see
Debrenne, Zh urAvlev, & roZAnov , 1989, p. 98;
Debrenne, roZAnov, & ZhurAvlev, 1990, p. 136].
Outer wall with downwardly projecting, straight
canals, bearing supplementary bracts externally
(imparting overall upright V-shaped appearance to
outer wall); inner wall with one row of downwardly
projecting, straight canals per intersept, bearing
supplementary bracts on central cavity side; septa
completely porous; pectinate tabulae may be present.
lower Cambrian (Atd.1–Bot.3): Siberian Platform,
Kolyma, Altay Sayan, Tuva, Mongolia, Transbai-
kalia, Far East.——Fi g. 44,1a–b. *C. loculatus,
Verkhneynyrga Formation, Botoman, Lebed’ River,
Altay Mountains, Altay Sayan, Russia, holotype,
TsNIGRm 45a/2957; a, transverse section (outer
wall at top), ×10; b, longitudinal section (outer wall
to right), ×10 (Vologdin, 1932).
Hupecyathellus roZA no v in DATs en Ko & others,
1968, p. 149 [*H. schuberti; OD; holotype,
DATsenKo & others, 1968, pl. 14,1–3; roZA nov,
1973, pl. 7,1, PIN 4297/75, Moscow]. Outer wall
with downwardly projecting, S-shaped canals,
bearing supplementary independent microporous
sheath externally; inner wall with several rows
of horizontal to upwardly projecting, S-shaped
canals per intersept; septa completely porous.
lower Cambrian (Bot.1): Siberian Platform.——
Fig. 44,2a–c. *H. schuberti, Shumnoy Formation,
Botoman, Sukharikha River, Krasnoyarsk region,
Russia, holotype, PIN 4297/75; a, oblique longi-
tudinal section, ×5; b, detail of septum in longi-
tudinal section (outer wall to right), ×7; c, detail
of outer wall in tangential section, ×17 (Datsenko
& others, 1968).
Porocyathellus Debrenne, 1977a, p. 107 [*P. bouddi;
OD; holotype, Debrenne, 1977a, pl. 6,1, MNHN
M80025, IRH 2 1aL, Paris]. Outer wall with
horizontal to upwardly projecting, S-shaped canals;
inner wall with several rows of horizontal to
upwardly projecting, S-shaped canals per intersept;
septa completely porous. lower Cambrian (Bot.1):
Morocco.——Fig. 44,3. *P. bouddi, Issafen Forma-
tion, Botoman, Jbel Irhoud, Morocco, holotype,
MNHN M80025, IRH 2 1aL, oblique longitudinal
section, ×10 (Debrenne, 1977a).
Vologdinocyathellus KonY ushKov , 1972, p. 136 [*V.
schischlovi; OD; holotype, KonYushKov , 1972, pl.
15,2, not located]. Outer wall with horizontal to
upwardly projecting, S-shaped canals; inner wall
with one row of horizontal to upwardly projecting,
straight canals per intersept; septa completely
porous. lower Cambrian (Bot.1): Altay Sayan.——
Fig . 44,4a–b. *V. schischlovi, Verkhnemonok
Formation, Botoman, Bol’shoy Karakol River,
western Sayan, Altay Sayan, Russia; a, holotype,
transverse section, ×4; b, unnumbered paratype,
oblique longitudinal section, ×4 (Konyushkov,
1972).
Family ETHMOPHYLLIDAE
Okulitch, 1937
[Ethmophyllidae oKuliTch, 1937b, p. 358] [=Archaeocyathellidae simon,
1939, p. 73; =Dupliporocyathidae YAZmir in YAZmir, D AlmATov, &
YAZmir, 1975, p. 58; =Kolbicyathidae Debrenne, roZAnov, & ZhurAvlev
in Debrenne, ZhurAvlev, & roZAnov, 1988, p. 97]
Inner wall with communicating canals.
lower Cambrian (Atd.4–Toy.1).
Ethmophyllum meeK, 1868a, p. 64 [*E. whitneyi;
OD; lectotype, oKuliTch, 1943, pl. 3,15; hill,
1965, pl. 4,1; SD oKuliTch, 1943, p. 66, USNM
15307 1,1b, thin sections A, Washington, D.C.].
Outer wall with subspherical, chambered canals,
each with base commencing in intervallum,
canals subdivided by stipules (imparting overall
inverted V-shaped appearance to outer wall);
inner wall with one row of anastomosing, hori-
zontal to upwardly and laterally projecting, waved
canals per intersept, arising from fluted inner
edges of septa; septa aporose to sparsely porous.
lower Cambrian (Atd.4–Bot.2): Canada, United
States.——Fig . 45,1a– c. *E. whitneyi, Poleta
Formation, Botoman, Silver Peak, Nevada, United
States; a–b, lectotype, USNM 15307 1,1b, thin
sections A; a, longitudinal section, ×5; b, trans-
verse section, ×5; c, topotype, MCZ 9314, detail
of septum in longitudinal section (outer wall to
left), ×9 (Debrenne, Zhuravlev, & Kruse, 2002).
Angaricyathus ZhurAvle vA, 1965, p. 7 [*A. cyrenovi;
OD; holotype, ZhurAvlevA, 1965, pl. 2,1, TsSGM
215, specimen 2, thin section 1, Novosibirsk].
Outer wall with horizontal to upwardly projecting,
straight canals, bearing supplementary bracts
externally (imparting overall inverted V-shaped
appearance to outer wall); inner wall with one
row of anastomosing, horizontal to upwardly
and laterally projecting, straight to waved canals
per intersept; septa completely porous; pectinate
tabulae may be present. lower Cambrian (Toy.1):
Transbaikalia, ?Sardinia.——Fig . 45,2a–b. *A.
cyrenovi, Kacha Formation, Toyonian, Kookta
River, Transbaikalia, Russia; a, holotype, TsSGM
215, specimen 2, oblique longitudinal section,
×6; b, paratype, TsSGM 215, specimen 4, trans-
verse section, ×6 (Zhuravleva, 1965).
Aulocricus Debrenne, 1987, p. 270 [*A. arellani;
OD; holotype, Debrenne, 1987, pl. 1,7, USNM

Systematic Descriptions of the Archaeocyatha 63
Fig. 44. Carinacyathidae (p. 62).
4a
Carinacyathus
Porocyathellus
Vologdinocyathellus
1b
3
1a
2b
2c
Hupecyathellus
4b
2a

64 Treatise Online, number XX
Fig. 45. Ethmophyllidae (p. 62–65).
2b
2a
Angaricyathus
Ethmophyllum
1a
1c
3a
3b
Dupliporocyathus
1b

Systematic Descriptions of the Archaeocyatha 65
111823, Washington, D.C.]. Outer wall with
horizontal to upwardly projecting, straight canals,
bearing supplementary bracts externally (imparting
overall inverted V-shaped appearance to outer
wall) and subdivided by stipules; inner wall with
anastomosing, horizontal to upwardly and laterally
projecting, straight stirrup canals only, bearing
supplementary planar annuli on central cavity side;
septa aporose to sparsely porous. lower Cambrian
(Bot.1): Canada, United States, Mexico.——Fig.
46,1a–b. *A. arellani, Puerto Blanco Formation,
Botoman, Caborca, Sonora, Mexico; a, holotype,
USNM 111823, transverse section, ×10; b, para-
type, USNM 414812, longitudinal section, ×10
(Debrenne, 1987).
Cordilleracyathus hAnDFielD, 1971, p. 49 [*C. blus-
soni; OD; holotype, hAnDFielD, 1971, pl. 7,2,
GSC 25345, Ottawa]. Outer wall with horizontal
to upwardly projecting, S-shaped canals, subdi-
vided by stipules and bearing supplementary bracts
externally (imparting overall inverted V-shaped
appearance to outer wall); inner wall with one
row of anastomosing, horizontally to upwardly
and laterally projecting, S-shaped canals per inter-
sept, formed by fluting of inner edges of septa,
bearing supplementary scales on central cavity
side; septa aporose to sparsely porous; pectinate
tabulae may be present. lower Cambrian (Atd.4–
Bot.2): Koryakia, Canada, United States, Mexico,
?Greenland.——Fig. 46,2a–c. *C. blussoni, Sekwi
Formation, Botoman, Caribou Pass, Northwest
Territories, Canada; a, holotype, GSC 25345,
detail of tangential section (outer wall at bottom,
inner wall at top), ×12; b, paratype, GSC 25348,
transverse section, ×10; c, paratype, GSC 25347,
detail of longitudinal section (outer wall to right),
×12 (Handfield, 1971).
?Dupliporocyathus YAZmir in YAZmir, DAlm ATov ,
& YAZmir, 1975, p. 59 (YAZmir in Zhur AvlevA ,
1974a, p. 119, nom. nud.) [*D. tumulosus; OD;
holotype, YAZmir, DAlmATo v, & YAZmir, 1975,
pl. 21,5–7, BGU 0138/17, Ulan-Ude]. Outer wall
longitudinally plicate, canal shape and orientation
uncertain; inner wall with one row of downwardly
projecting, straight canals per intersept, bearing
supplementary bracts on central cavity side; septa
aporose to sparsely porous. [Limited type material
does not provide certainty as to inner wall struc-
ture: canals with supplementary bracts, or V-shaped
scales.] lower Cambrian (Atd.4): Transbaikalia.——
Fig. 45,3a–b. *D. tumulosus, Oldyndy Formation,
Atdabanian, Ul’dzuytuy Creek, Vitim Highlands,
Transbaikalia, Russia, holotype, BGU 0138/17;
a, transverse section, ×10; b, oblique longitudinal
section, ×10 (Yazmir, Dalmatov, & Yazmir, 1975).
Kolbicyathus ZhurAvl ev in Debrenne, Zhur Avlev,
& roZ An o v , 1988, p. 97 [*Vologdinocyathus
kolbiensis Zhur AvlevA, 1959, p. 425; OD; holo-
type, Zhu rAvle vA, 1959, fig. 2d; TsSGM 282/2,
Novosibirsk]. Outer wall with horizontal to
upwardly projecting, S-shaped canals, bearing
supplementary bracts externally (imparting overall
inverted V-shaped appearance to outer wall);
inner wall with horizontal to upwardly projecting,
straight stirrup canals only, bearing supplementary
bracts on central cavity side; septa completely
porous; pectinate tabulae may be present. lower
Cambrian (Bot.1): Altay Sayan, Mongolia.——
Fig. 47,1a–b. *K. kolbiensis (Zh ur Avl evA), Usa
Format ion , Boto man , Pe trov ka, Kiya River,
Kuznetsk Alatau, Russia, holotype, TsSGM 282/2;
a, transverse section, ×5; b, detail of same, ×15
(Debrenne, Zhuravlev, & Kruse, 2002).
Parethmophyllum Debrenne, 1987, p. 270 [*Ethmo-
phyllum cooperi oKuliTch in cooPer & others,
1952, p. 29; OD; holotype, co oPe r & others,
1952, pl. 7,1–2; Debrenne, 1987, pl. 1,2,4, USNM
111814, Washington, D.C.]. Outer wall with hori-
zontal to upwardly projecting, straight canals; inner
wall with one row of anastomosing, horizontal
to upwardly and laterally projecting, straight to
waved canals per intersept, formed by fluting of
inner edges of septa, bearing supplementary bracts
on central cavity side; septa aporose to sparsely
porous. lower Cambrian (Bot.1): United States,
Mexico.——Fig. 47,2a–c. *P. cooperi (oKuliTch),
Puerto Blanco Formation, Botoman, Caborca,
Sonora, Mexico; a, paratype, USNM 1118 13,
oblique longitudinal section, ×6 (Debrenne,
Zhuravlev, & Kruse, 2002); b–c, holotype, USNM
111814; b, transverse section, ×6; c, longitudinal
section, ×6 (Debrenne, 1987).
Squamosocyathus ZhurAvlevA, 1960b, p. 183 [*S.
taumatus; OD; holotype, Zhur AvlevA, 1960b, pl.
13,5a–b, TsSGM 205/66a-b, Novosibirsk]. Outer
wall with horizontal to upwardly projecting straight
canals, bearing supplementary bracts externally
(imparting overall inverted V-shaped appearance to
outer wall); inner wall with one row of horizontal
to upwardly projecting, straight porous canals per
intersept; septa completely porous; pectinate tabulae
may be present. lower Cambrian (Atd.3–Bot.1):
Siberian Platform, ?Altay Sayan.——Fig. 46,3a–c.
*S. taumatus, Perekhod Formation, Atdabanian;
a, Botoma River, Sakha (Yakutia), Russia, para-
type, TsSGM 205/67, longitudinal section, ×6
(Zhuravleva, 1960b); b–c, Yudyay, Lena River, Sakha
(Yakutia), Russia; b, holotype, TsSGM 205/66a-b,
detail of septum in longitudinal section (outer wall
to left), ×12 (Debrenne, Zhuravlev, & Kruse, 2002);
c, paratype, TsSGM 205/68, transverse section, ×6
(Zhuravleva, 1960b).
Stephenicyathus ZhurAvlev in vo ronovA & others,
1987, p. 26 [*S. rowlandi; OD; holotype, voronovA
& others, 1987, pl. 6,1, GSC 90145, Ottawa]. Outer
wall with horizontal to upwardly projecting, straight
canals, bearing supplementary bracts externally
(imparting overall inverted V-shaped appearance
to outer wall) and subdivided by stipules; inner
wall with one row of anastomosing, horizontal to
upwardly projecting, waved canals per intersept,
formed by fluting of inner edges of septa; septa
aporose to sparsely porous; pectinate tabulae may
be present. lower Cambrian (Bot.1): Canada, United
States.——Fig. 47,3a–b. *S. rowlandi, Sekwi Forma-
tion, Botoman, Mackenzie Mountains, Northwest

66 Treatise Online, number XX
Fig. 46. Ethmophyllidae (p. 62–65).
2a 2b
2c
Cordilleracyathus
1a
1b Aulocricus
3a
3c
Squamosocyathus
3b

Systematic Descriptions of the Archaeocyatha 67
Fig. 47. Ethmophyllidae (p. 65–68).
3a 3b
Stephenicyathus
2a
2b 2c
Parethmophyllum
1a
1b Kolbicyathus

68 Treatise Online, number XX
Territories, Canada; a, paratype, GSC 90146, longitu-
dinal section, ×12; b, holotype, GSC 90145, oblique
longitudinal section, ×5 (Voronova & others, 1987).
Superfamily TERCYATHOIDEA
Vologdin, 1939
[nom. correct. Debrenne & Kruse, 1986, p. 256, pro Tercyathacea ZhurAv-
levA, 1960b, p. 184, nom. transl. ex Tercyathidae vologDin in simon,
1939, p. 11] [=Piamaecyathacea ZhurAvlevA, 1960a, p. 44, nom. transl.
ZhurAvlevA, 1960b, p. 50, ex Piamaecyathidae ZhurAvlevA, 1960a, p. 44]
Outer wall clathrate. lower Cambrian
(Atd.4–Toy.1).
Family PIAMAECYATHELLIDAE
Rozanov, 1974
[Piamaecyathellidae roZAnov in boroDinA, 1974, p. 157] [=Piamaecy-
athellidae roZAnov, 1973, p. 86, nom. nud.]
Inner wall with simple pores. l o wer
Cambrian (Bot.2).
Piamaecyathellus roZAnov in rePinA & others, 1964,
p. 217 [*P. simplex; OD; holotype, rePinA & others,
1964, pl. 23,5, PIN 4297/28, Moscow]. Inner wall
with several rows of simple pores per intersept; septa
completely porous; pectinate tabulae may be present.
lower Cambrian (Bot.2): Altay Sayan.——Fig. 48,1.
*P. simplex, Verkhneynyrga Formation, Botoman,
Kyzyl-Tash, Bol’shaya Isha River, Altay Mountains,
Altay Sayan, Russia, holotype, PIN 4297/28, trans-
verse section, ×5 (Repina & others, 1964).
Family BOTOMOCYATHIDAE
Zhuravleva, 1955
[Botomocyathidae ZhurAvlevA, 1955b, p. 628] [=Botomacyathidae
ZhurAvlevA in vologDin, 1956, p. 879, nom. null.]
Inner wall with annuli. lower Cambrian
(Atd.4–Bot.3).
Botomocyathus ZhurAvlevA, 1955b, p. 629 (ZhurAv-
levA, 1954, p. 12, nom. nud.) [*B. zelenovi; OD;
holotype, Zh ur Avl evA , 1955b, fig. 2e, TsSGM
205/69, Novosibirsk] [=Botomacyathus ZhurAvlevA
in vologDin, 1956, p. 879, nom. null.]. Inner wall
with one pore row per intersept and upright, V-shaped
annuli; septa completely porous; pectinate tabulae
may be present. lower Cambrian (Atd.4–Bot.1):
Siberian Platform, Altay Sayan, Far East.——Fi g.
48,2a–c. *B. zelenovi, Perekhod Formation, Botoman,
Botoma River, Sakha (Yakutia), Russia; a–b, holotype,
TsSGM 205/69; a, oblique longitudinal section,
×6; b, detail of outer wall in tangential section,
×16; c, paratype TsSGM 205/70, detail of longitu-
dinal section (outer wall to right), ×10 (Debrenne,
Zhuravlev, & Kruse, 2002).
Clathrithalamus Debrenne & Kr use, 1986, p. 256
[*C. mawsoni; OD; holotype, Debrenne & Kruse,
1986, fig. 21A–C, GNS MG511, Lower Hutt]. Inner
wall with one pore row per intersept and upright,
V-shaped annuli; septa aporose to sparsely porous;
pectinate tabulae may be present. lower Cambrian
(Bot.3): Antarctica.——Fig. 48,3a–c. *C. mawsoni,
Shackleton Limestone, Botoman, Holyoake Range,
Nimrod Glacier, Antarctica, holotype, GNS MG511;
a, transverse section, ×5 (Debrenne & Kruse, 1986);
b, detail of outer wall in tangential section, ×16
(Debrenne, Zhuravlev, & Kruse, 2002); c, detail
of oblique longitudinal section, ×7 (Debrenne &
Kruse, 1986).
Family OLGAECYATHIDAE
Borodina, 1974
[Olgaecyathidae boroDinA, 1974, p. 158]
Inner wall with noncommunicating
canals. lower Cambrian (Bot.2).
Olgaecyathus boroDinA, 1974, p. 158 [*O. fistu-
losus; OD; holotype, bor oDi nA, 1974, pl. 16,8,
TsSGM 429/2, Novosibirsk]. Inner wall with
one row of horizontal to upwardly projecting,
straight canals per intersept; septa completely
porous; pectinate tabulae may be present. lower
Cambrian (Bot.2): Altay Sayan.——Fig. 49. *O.
fistulosus, Verkhnemonok Formation, Botoman,
Kazly River, western Sayan, Altay Sayan, Russia,
holotype, TsSGM 429/2, oblique longitudinal
section, ×8 (Debrenne, Zhuravlev, & Kruse,
2002).
Family TERCYATHIDAE
Vologdin, 1939
[Tercyathidae vologDin in simon, 1939, p. 11] [=Tercyathidae volog-
Din, 1937b, p. 459, nom. nud., invalid family-group name based on
unavailable genus name; =Piamaecyathidae ZhurAvlevA, 1960a, p. 44]
Inner wall with communicating canals.
lower Cambrian (Bot.1–Toy.1).
Tercyathus vol o g D i n in si m o n , 1939, p. 40
(vologDin, 1932, p. 55, nom. nud., without desig-
nated type species) [*T. duplex vologDin, 1932, p.
56; OD; lectotype, vologDin, 1932, pl. 13,2; SD
boroDin A, 1974, p. 154, TsNIGRm 71a-g/2957,
St. Petersburg] [=Piamaecyathus ZhurAvlevA,
1960a, p. 45 (type, P. sajanicus, OD), for discus-
sion, see Debrenne, ZhurAvlev, & roZA no v,
1989, p. 135; Debrenne, roZAnov, & ZhurAvlev,
1990, p. 164; =Rugocyathus vologDin, 1962d,
p. 13 (type, R. venustus, M), nom. nud.]. Inner
wall with one row of anastomosing, horizontal
to upwardly and laterally projecting, straight to
waved canals per intersept; supplementary spines,
annular structures and/or microporous sheath may
be present on central cavity side; septa sparsely
to completely porous; pectinate tabulae may be
present. lower Cambr ian (B o t.1 –Toy.1 ): Altay
Sayan.——Fig. 50,1a–c. *T. duplex, Verkhneynyrga
Formation, Botoman, Lebed’ River, Altay Moun-
tains, Altay Sayan, Russia; a, lectotype, TsNIGRm
71a-g/2957, longitudinal section of inner wall
(central cavity to right), ×6 (Vologdin, 1932); b,
unlocated specimen, longitudinal section, ×6; c,

Systematic Descriptions of the Archaeocyatha 69
Fig. 48. Piamaecyathellidae and Botomocyathidae (p. 68).
1
3a
3b
3c
2a
2b
2c
Piamaecyathellus
Botomocyathus
Clathrithalamus

70 Treatise Online, number XX
unlocated specimen, transverse section, ×6 (Repina
& others, 1964).
Clathricyathellus boroDinA, 1974, p. 150 [*Clath-
ricyathus robustus vologDin, 1932, p. 53; OD;
lectotype, vologDin, 1932, pl. 12,6; SD boroDinA,
1974, p. 151, TsNIGRm 68a/2957, St. Petersburg]
[=Lebedicyathus boroDinA, 1974, p. 164 (type,
L. duplicatus, OD)]. Inner wall with one row of
downwardly projecting, straight porous canals per
intersept; supplementary spines, annular struc-
tures, and/or microporous sheath may be present
on central cavity side; septa completely porous;
pectinate tabulae may be present. lower Cambrian
(Bot.3): Altay Sayan.——Fig. 50,2a–b. *C. robustus
(vologDin), Verkhneynyrga Formation, Botoman,
Lebed’ River, Altay Mountains, Altay Sayan, Russia;
a, lectotype, TsNIGRm 68a/2957, oblique transverse
section, ×4; b, paralectotype, TsNIGRm 69a/2957,
longitudinal section, ×4 (Vologdin, 1932).
Clathricyathus vologDin in simon, 1939, p. 25
(VologDin, 1932, p. 50, nom. nud., without
designated type species) [*C. firmus vologDin,
1932, p. 50; OD; lectotype, vologDin, 1932, pl.
10,12; SD boroDi nA, 1974, p. 150, TsNIGRm
65, 65a-v/2957, St. Petersburg] [=Clathrocyathus
vologDin, 1937b, p. 469, nom. null.]. Inner
wall with one row of amalgamating, downwardly
projecting, straight canals per intersept, continuing
into larger, horizontal to upwardly projecting,
confluent canal system on central cavity side;
septa completely porous; pectinate tabulae may be
present. lower Cambrian (Bot.3): Altay Sayan.——
Fig. 51,1a–b. *C. firmus, Verkhneynyrga Forma-
tion, Botoman, Lebed’ River, Altay Mountains,
Altay Sayan, Russia, lectotype, TsNIGRm 65,
65a-v/2957; a, transverse section, ×7; b, oblique
longitudinal section, ×7 (Vologdin, 1932).
Tercyathellus boroDinA, 1974, p. 155 [*T. capisterium;
OD; holotype, boroDinA, 1974, fig. 13, pl. 10,3,
TsSGM 429/1, Novosibirsk] [=Kazlycyathus boro-
DinA, 1974, p. 163 (type, K. flexuosus, OD), for
discussion, see Debrenne, ZhurAvlev, & roZAnov,
1989, p. 135; Debrenne, roZAnov, & ZhurAvlev,
1990, p. 164]. Inner wall with several rows of anas-
tomosing, horizontal to upwardly and laterally
projecting, straight to waved canals per intersept,
continuing into larger, horizontal to upward ly
projecting, confluent canal system on central cavity
side; septa sparsely to completely porous; pectinate
tabulae may be present. lower Cambrian (Bot.2):
Altay Sayan.——Fig. 51,2. *T. capisterium, Verkh-
nemonok Formation, Botoman, Kazly River, western
Sayan, Altay Sayan, Russia, holotype, TsSGM 429/1,
oblique transverse section, ×8 (Borodina, 1974).
Superfamily
SIGMOCYATHOIDEA
Krasnopeeva, 1953
[nom. correct. Debrenne & Kruse, 1986, p. 255, pro Sigmocyathacea
Debrenne, 1970a, p. 25, nom. transl. ex Sigmocyathidae KrAsnoPeevA,
1953, p. 56, as Sygmocyathidae nom. null.]
Outer wall with annuli. lower Cambrian
(Bot.3).
Family SIGMOCYATHIDAE
Krasnopeeva, 1953
[nom. correct. ZhurAvlevA, 1960b, p. 49, pro Sygmocyathidae KrAsnoPe-
evA, 1953, p. 56, nom. null., based on erroneous spelling of generic name]
Inner wall with annuli. lower Cambrian
(Bot.3).
Sigmocyathus R. beDForD & J. beDForD, 1936,
p. 23 [*Coscinocyathus didymoteichus TAY lo r ,
1910, p. 140; OD; lectotype, TAYlo r, 1910, pl.
10, photo 58; Debrenne & roZAno v, 1972, pl.
43,1,3; roZ Ano v, 1973, pl. 4,3; SD Debrenne,
1970a, p. 42, SAM T1606B-D, Adelaide]
[=Hemistillicidocyathus Ting, 1937, p. 368 (type,
Coscinocyathus didymoteichus TAY lo r, 1910, p.
140, OD)]. Outer wall with upwardly projecting,
S-shaped annuli; inner wall with one pore row
Fig. 49. Olgaecyathidae (p. 68).
Olgaecyathus

Systematic Descriptions of the Archaeocyatha 71
1a
1b
1c
Tercyathus
2b
2a Clathricyathellus
Fig. 50. Tercyathidae (p. 68–70).

72 Treatise Online, number XX
per intersept and upwardly projecting, S-shaped
annuli; septa aporose to sparsely porous. l ower
Cambrian (Bot.3): Australia, ?Antarctica.——Fig.
52a–c. *S. didymoteichus (TAYlo r), Ajax Lime-
stone, Botoman, Ajax Mine, South Australia,
Australia, lectotype, SAM T1606B-D; a, longi-
tudinal view of septum (outer wall to left), ×8;
b, external view of outer wall, ×8 (Debrenne &
Rozanov, 1972); c, transverse view, ×7 (Debrenne,
Zhuravlev, & Kruse, 2002).
Didymocyathus Debrenne & ro ZA no v, 1972, p.
236 [*D. hillae; OD; holotype, Debrenne &
roZ Ano v, 1972, pl. 42,2–3; pl. 43,2; pl. 44,1,
USNM PU299, Washington, D.C.]. Outer wall
with upwardly projecting, S-shaped annuli;
inner wall with one pore row per intersept and
upwardly projecting, S-shaped annuli; septa
completely porous. lower Cambrian (Bot.3):
Australia, ?Antarctica.——Fig. 53a–c. *D. hillae,
Ajax Limestone, Botoman, Ajax Mine, South
Australia, Australia; a, holotype, USNM PU299,
longitudinal view of septum (outer wall to left),
×5 (Debrenne, Zhuravlev, & Kruse, 2002); b–c,
topotype, USNM PU296; b, longitudinal view of
outer wall, ×5; c, longitudinal view of inner wall,
×5 (Debrenne & Rozanov, 1972).
Family WRIGHTICYATHIDAE
Kruse, 1978
[Wrighticyathidae Kruse, 1978, p. 34]
Inner wall with communicating canals.
lower Cambrian (Bot.3).
Wrighticyathus Kruse, 1978, p. 34 [*W. nexus; OD;
holotype, Krus e, 1978, fig. 7A–E, AM F.83298,
Sydney]. Outer wall with upwardly projecting,
S-shaped annuli; inner wall with one row of hori-
zontal to upwardly projecting, straight canals
per intersept; septa completely porous. low er
Cambrian (Bot.3): Australia.——Fig . 54a–c. *W.
nexus, Cymbric Vale Formation, Botoman, Mt.
Wright, New South Wales, Australia, holotype,
AM F.83298; a, transverse section, AM FT.8268,
Fig. 51. Tercyathidae (p. 70).
2
1a
1b
Clathricyathus
Tercyathellus

Systematic Descriptions of the Archaeocyatha 73
×4 (Kruse, 1978); b, longitudinal section, AM
FT.8265, ×4 (Debrenne, Zhuravlev, & Kruse,
2002); c, detail of outer wall in longitudinal
section, AM FT.8264, ×10 (Kruse, 1978).
Suborder ERISMACOSCININA
Debrenne, Rozanov, & Zhuravlev,
1989
[Erismacoscinina Debrenne, r oZAnov, & ZhurAvlev in Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 85] [?=Erismacoscinina ZhurAvlev ,
1988, p. 105, nom. nud.]
Intervallum with septa and plate tabulae;
synapticulae may be present. lower Cambrian
(Tom.2–Bot.3).
Superfamily
SALAIROCYATHOIDEA
Zhuravleva, 1956
[nom. transl. ZhurAvlev, 1988, p. 105, ex Salairocyathidae ZhurAvlevA in
vologDin, 1956, p. 879] [=Erismacoscinacea Debrenne, 1964, p. 166,
nom. transl. Debrenne, 1970a, p. 25, ex Erismacoscinidae Debrenne,
1964, p. 166]
Outer wall with simple pores. lowe r
Cambrian (Tom.2–Bot.3).
Family ASTEROCYATHIDAE
Vologdin, 1956
[Asterocyathidae vologDin, 1956, p. 879] [=Erismacoscinidae Debrenne,
1964, p. 166; =Syringocoscinidae vologDin & YAZmir, 1967, p. 1375]
Inner wall with simple pores. l o wer
Cambrian (Tom.2–Bot.3).
Asterocyathus vologDin, 1940b, p. 92 [*A. salairicus;
OD; holotype not designated, collection not
located]. Inner wall longitudinally plicate, with
several rows of simple pores per intersept; septa
completely porous; tabulae with normal pores.
lower Cambrian (Atd.1–Atd.2): Altay Sayan,
Tuva, Mongolia.——Fig . 55,1. *A. salairicus,
Gavrilovskoe Formation, Atdabanian, Belaya
Gorka, Gorskino, Salair, Russia, unlocated syntype,
oblique transverse section, ×5 (Vologdin, 1940b).
Antoniocoscinus ZhurAvlev in Debrenne, ZhurAvlev,
& roZAnov, 1988, p. 98 [*Coscinocyathus vsevolodi
Korshunov in Zhu r Av l e vA , Korshunov, &
roZA nov , 1969, p. 51; OD; holotype, Zhur Av-
levA, Korshunov, & ro ZAnov , 1969, pl. 20,2,4;
Korshunov, 1972, pl. 15,5; pl. 16,5), TsSGM
323/84, Novosibirsk]. Inner wall with one row
of simple pores per intersept; septa completely
porous; tabulae with normal pores. lower Cambrian
(Atd.4–Bot.3): Siberian Platform, Tuva, Transbai-
kalia, Morocco, Iberia, Sardinia.——Fig. 55,2a–b.
Fig. 52. Sigmocyathidae (p. 70–72).
b
a
c
Sigmocyathus

74 Treatise Online, number XX
*A. vsevolodi (Korshunov), Oy-Muran reef massif,
Botoman, Oy-Muran, Lena River, Sakha (Yakutia),
Russia, holotype, TsSGM 323/84; a, transverse
section, ×4 (M. Debrenne, new); b, detail of
longitudinal section (outer wall to left), ×10 (Kors-
hunov, 1972).
Erismacoscinus Debrenne, 1958, p. 65 [*E. marocanus;
M; holotype, Debrenne, 1958, pl. 3,12,14–16;
Debrenne, 1964, pl. 20,1–2, MNHN M80139,
specimen H2, Paris] [=Pluralicoscinus Debrenne,
1963b, p. 135 (type, P. alanisensis, OD); =Syrin-
gocoscinus YAZmir in vologDin & YAZmir, 1967,
p. 1376 (type, S. angulatus, OD), for discussion,
see Debrenne, Zhur Avlev, & roZAn ov, 1989, p.
105; Debrenne, roZAnov, & Zhur Avlev , 1990,
p. 141; =Churanocyathus sun DuKov , 1984, p. 14
(type, C. aculeatus, OD)]. Inner wall with several
rows of simple pores per intersept; septa completely
b
a
c
Didymocyathus
Fig. 53. Sigmocyathidae (p. 72).

Systematic Descriptions of the Archaeocyatha 75
porous; tabulae with normal pores. lower Cambrian
(Tom.2–Bot.3): Siberian Platform, Kolyma, Altay
Sayan, Tuva, Mongolia, Transbaikalia, Far East,
Urals, Tajikistan, Australia, Antarctica, Tarim,
southern China, Morocco, Iberia, France, Sardinia,
Germany.——Fig. 55,3a–b. *E. marocanus, Amou-
slek Formation, Atdabanian, Jbel Taïssa, Morocco,
holotype, MNHN M80139, specimen H2; a,
longitudinal section (outer wall to left), ×4; b,
transverse section, ×4 (Debrenne, 1958).
Ichnusocyathus Debrenne, 1977a, p. 103 [*Archaeo-
cyathus ichnusae meneghini, 1881, p. 201; OD;
lectotype, bornemAnn, 1886, pl. 13,3–4; SD
Debrenne, 1964, p. 129, not located]. Inner
wall with several rows of simple pores per inter-
sept; septa aporose to sparsely porous; tabulae
with normal pores. lower Cambrian (Bot.1):
Morocco, Sardinia.——Fig. 56a–b. *I. ichnusae
(meneghini), Matoppa Formation, Botoman,
Monte Gloria, Canal Grande, Sardinia, Italy;
a, lectotype, oblique section, ×4 (Bornemann,
1886); b, topotype, MNHN M84258, specimen
RFB 14/1, transverse section, ×4 (Debrenne,
Zhuravlev, & Kruse, 2002).
Retecoscinus ZhurAvlevA, 1960b, p. 247 [*Coscino-
cyathus retetabulae vologDin, 1931, p. 75; OD;
lectotype, vologDin, 1931, pl. 22,1e; SD Debrenne,
ZhurAvl ev, & Krus e, 2002, p. 1617, TsNIGRm
94a/2956, St. Petersburg]. Inner wall with several
rows of simple pores per intersept; septa completely
porous; tabulae with slitlike pores. lower Cambrian
(Tom.3–Bot.1): Siberian Platform, Altay Sayan, Far
East, Morocco, Iberia, France, Germany.——Fig.
57,1a–b. *R. retetabulae (vologDin), Usa Formation,
Atdabanian, Nizhnyaya Ters’ River, Kuznetsk Alatau,
Russia, lectotype, TsNIGRm 94a/2956; a, transverse
section, ×2; b, unlocated specimen TsNIGRm, sketch
of oblique transverse section, ×2.5 (Vologdin, 1931).
Rozanovicoscinus Debrenne, 1970a, p. 41 [*R. fonini;
OD; holotype, Debrenne, 1970a, pl. 2,1, USNM
PU86614, Washington, D.C.]. Outer wall longi-
tudinally plicate; inner wall with several rows of
simple pores per intersept; septa aporose to sparsely
porous; tabulae with normal pores. lower Cambrian
(Atd.4–Bot.3): Australia.——Fig. 57,2. *R. fonini,
Ajax Limestone, Botoman, Ajax Mine, South
Australia, Australia, holotype, USNM PU86614,
transverse view, ×5 (Debrenne, 1970a).
Fig. 54. Wrighticyathidae (p. 72–73).
a
b
c
Wrighticyathus

76 Treatise Online, number XX
Fig. 55. Asterocyathidae (p. 73–75).
2a
1
3a 3b
2b
Asterocyathus
Antoniocoscinus
Erismacoscinus

Systematic Descriptions of the Archaeocyatha 77
Family RUDANULIDAE
Debrenne, Rozanov, & Zhuravlev, 1989
[Rudanulidae Debrenne, roZA no v, & ZhurAv le v in Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 85]
Inner wall with bracts or scales. lower
Cambrian (Bot.1–Bot.3).
Rudanulus Debrenne in ZhurAvlevA, 1974b, p.
79 [*Coscinocyathus petersi R. beDForD & W. R.
beDForD, 1934, p. 3; OD; holotype, r. beDForD
& W. r. beDForD, 1934, fig. 13; Debrenne, 1969a,
pl. 10,4–5; M, NHM S4158, London]. Outer wall
longitudinally plicate; inner wall with several rows
of pores per intersept, bearing upwardly projecting,
S-shaped scales; septa completely porous; tabulae
with normal pores. lower Cambrian (Bot.1–Bot.3):
Australia, southern China.——Fi g. 58,1a–c. *R.
petersi (R. beD F or D & W. r. beD Fo r D), Ajax
Limestone, Botoman, Ajax Mine, South Australia,
Australia, holotype, NHM S4158; a, transverse view,
×8 (Debrenne, 1969a); b, longitudinal view, ×8; c,
detail of transverse view, ×16 (M. Debrenne, new).
Pilodicoscinus Debrenne & jiAng, 1989, p. 826
[*P. yuani; OD; holotype, Debrenne & j iAng,
1989, pl. 2,6, MNHN M85002, specimen 2-13,
Paris]. Cup in which outer wall shows periodic
transverse folds; inner wall with several rows of
pores per intersept, bearing upwardly projecting,
cupped bracts; septa aporose to sparsely porous;
Fig. 56. Asterocyathidae (p. 75).
a
b
Ichnusocyathus

78 Treatise Online, number XX
tabulae with normal pores. lower Cambrian (Bot.3):
southern China.——Fig . 58,2. *P. yuani, Tsang-
langpu Formation, Botoman, Yangchang, Yunnan,
China, holotype, MNHN M85002, specimen
2-13, longitudinal section, ×7.5 (Debrenne &
Jiang, 1989).
Yhecyathus belY Ae vA & Yu An , 1995, p. 140 [*Y.
futchinensis; OD; holotype, be lYA evA & YuA n,
1995, fig. 1d, NIGP NF6H1, Nanjing]. Cup with
regular transverse folds affecting both walls; inner
wall with several rows of pores per intersept,
bearing upwardly projecting, cupped bracts; septa
completely porous; tabulae with normal pores.
lower Cambrian (Bot.1): southern China.——Fig .
58,3. *Y. futchinensis, Xiannudong Formation,
Botoman, Fuchin, Shaanxi, China, holotype, NIGP
NF6H1, oblique longitudinal section, ×2 (Belyaeva
& Yuan, 1995).
Family SALAIROCYATHIDAE
Zhuravleva, 1956
[Salairocyathidae ZhurAvlevA in vologDin, 1956, p. 879]
Inner wall with annuli. lower Cambrian
(Atd.2–Bot.1).
Salairocyathus vologDin, 1940b, p. 89 [*S .
zenkovae; OD; holotype not designated, collec-
tion not located] [=Rimotabulocyathus YA ro -
shevich, 1990, p. 26 (type, R. bulynnikovi,
OD)]. Inner wall with one pore row per
intersept and upright V-shaped annuli; septa
completely porous; tabulae with slitlike pores.
lower Cambrian (Atd.2): Altay Sayan.——Fi g.
59,1. *S. zenkovae, Gavrilovskoe Formation,
Atdabanian, Belaya Gorka, Salair, Russia, unlo-
cated syntype, oblique longitudinal section, ×6
(Vologdin, 1940b).
Kotuyicoscinus sunDuKov, 1983, p. 16 [*K. minaevae;
OD; holotype, sunDuKov, 1983, pl. 1,7, SNIIG-
GiMS 1580/2, Novosibirsk]. Inner wall with several
pore rows per intersept and upwardly projecting,
S-shaped annuli; septa completely porous; tabulae
with normal pores. lower Cambrian (Atd.2–Atd.3):
Siberian Platform.——Fig. 59,2a–b. *K. minaevae,
Kyndyn Formation, Chomp-Yurekh Creek, Kotuy
River, Krasnoyarsk region, Russia; a, holotype,
SNIIGGiMS 1580/2, oblique longitudinal section,
×9; b, paratype, SNIIGGiMS 1580/1, longitudinal
section, ×16 (Sundukov, 1983).
Polystillicidocyathus Debrenne, 1959a, p. 14 [*P.
erbosimilis; OD; holotype, Debrenne, 1959a,
fig. 1; Debrenne, 1964, pl. 17,1–2, MNHN
M80166, specimen Ki140, Paris]. Inner wall with
one pore row per intersept and upright, V-shaped
annuli; septa completely porous; tabulae with
normal pores. lower Cambrian (Bot.1): Altay Sayan,
Morocco.——Fig . 59,3a–b. *P. erbosimilis, Issafen
Formation, Botoman, Tizi Oumeslema, Morocco;
holotype, MNHN M80166, specimen Ki140,
modular skeleton; a, oblique transverse view, ×4;
b, longitudinal view, ×3 (Debrenne, 1964).
2Rozanovicoscinus
1a
1b
Retecoscinus
Fig. 57. Asterocyathidae (p. 75).

Systematic Descriptions of the Archaeocyatha 79
Fig. 58. Rudanulidae (p. 77–78).
2
3
Pilodicoscinus
Yhecyathus
1a
1b
1c
Rudanulus

80 Treatise Online, number XX
Fig. 59. Salairocyathidae (p. 78).
3a 3b
Polystillicidocyathus
2a
2b
Kotuyicoscinus
1
Salairocyathus

Systematic Descriptions of the Archaeocyatha 81
3
Crucicyathus
Dentatocoscinus
2a
2b
1Crassicoscinus
Fig. 60. Crassicoscinidae (p. 82).

82 Treatise Online, number XX
Family CRASSICOSCINIDAE
Debrenne, Rozanov, & Zhuravlev, 1988
[Crassicoscinidae Debrenne, r oZAnov, & Zhu rAvlev in Debrenne,
ZhurAvlev, & roZAnov, 1988, p. 98]
Inner wall with noncommunicating
canals. lower Cambrian (Atd.4–Bot.1).
Crassicoscinus ro Z A n o v & Zh u r A v l e v in
Debrenne, Zhu r A v l e v , & roZ A n o v , 1988,
p. 98 [*Coscinocyathellus vulgaris ro Z An ov in
reP i n A & othe rs, 1964, p. 227 ; O D; holo -
ty pe, rePi n A & others, 1964, pl. 24,2, PIN
4297/29, Moscow]. Inner wall with several rows
of horizontal to upwardly projecting, straight
canals per intersept; septa completely porous;
tabulae with normal pores. lower Cambrian
(Atd.4–Bot.1): Altay Sayan.——Fi g. 60,1. *C.
vulgaris (ro Z An o v), Uba Formation, Atdaba-
nian, Verkhnyaya Tyrga River, Altay Mountains,
Altay Sayan, Russia, holotype, PIN 4297/29,
transverse section, ×8 (Debrenne, Zhuravlev,
& Kruse, 2002).
Crucicyathus gr Av e s T o c K , 1984, p. 74 [*C .
repandus; OD; holotype, gr Ave s To c K , 1984,
fig. 42A–B, D, SAM P21585, Adelaide].
Outer wall longitudinally plicate; inner wall
with several rows of horizontal to upwardly
projecting, S-shaped canals per intersept; septa
completely porous; tabulae with normal pores.
lower Cambrian (Atd.4): Australia.——Fig .
60,2a–b. *C. repandus, Ajax Limestone, Atda-
banian, Mount Scott Range, South Australia,
Australia, holotype, SAM P21585; a, longitu-
dinal section, ×3.5; b, transverse section, ×3
(Gravestock, 1984).
Dentatocoscinus ZhurAvlev i n Debrenne,
Zhur Avl ev , & roZAn ov, 1988, p. 98 [*Astero-
tumulus sektensis Korshunov & ZhurAvlevA,
1967, p. 10; OD; holotype, Korshunov &
ZhurAvlevA, 1967, pl. 2,5, TsSGM 247/11,
Novosibirsk]. Outer wall longitudinally plicate;
inner wall with several rows of horizontal to
upwardly projecting, straight canals per inter-
sept, bearing supplementary bracts on central
cavity side; septa completely porous; tabulae
with normal pores. lower Cambrian (Bot.1):
Siberian Platform.——Fi g. 60,3. *D. sektensis
(Korshunov & ZhurAvlevA), Sekten Forma-
tion, Botoman, Tuora-Sis Range, Lena River,
Sakha (Yakutia), Russia, holotype, TsSGM
247/11, detail of oblique transverse section, ×9
(Korshunov & Zhuravleva, 1967).
Superfamily KASYRICYATHOIDEA
Zhuravleva, 1961
[nom. transl. Debrenne, ZhurAvlev, & Kruse, 2002, p. 1620, ex Kasyri-
cyathidae ZhurAvlevA in musATov & others, 1961, p. 29]
Outer wall with independent micropo-
rous sheath. lower Cambrian (Atd.1–Bot.3).
Family AGYREKOCYATHIDAE
Konyushkov, 1967
[Agyrekocyathidae KonYushKov, 1967, p. 110]
Inner wall with simple pores. l o wer
Cambrian (Atd.1–Bot.1).
Agyrekocyathus Kon Yu sh Ko v, 1967, p. 110 [*A.
malovi; OD; holotype, KonYushKov, 1967,
pl. 1,1 0, TsNIGRm 8722/6, St. Petersburg]
[= Mennericyathus Debrenne & roZ An o v in
ZhurAvlevA, 1974a, p. 216 (type, Tomocyathus
kundatus ro Z A n ov in ro Z A n o v & missAr-
ZhevsKiY, 1966, p. 63, OD)]. Inner wall with
several rows of simple pores per intersept; septa
completely porous; tabulae with normal pores.
lower Cambrian (Atd.1–Bot.1): Siberian Platform,
Altay Sayan, Mongolia, Far East, Kazakhstan,
Tajikistan, Australia, Antarctica, Morocco, Iberia,
Sardinia.——Fig. 61,1. *A. malovi, Boshchekul’
Formation, Atdabanian, Agyrek Mountains,
northern Kazakhstan, holotype, TsNIGRm
8722/6, transverse section, ×5 (Debrenne,
Zhuravlev, & Kruse, 2002).
Family XESTECYATHIDAE
Debrenne, Rozanov, & Zhuravlev, 1989
[Xestecyathidae Debrenne, roZAn ov, & Zh urAv lev i n Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 86]
Inner wall with bracts or scales. lower
Cambrian (Bot.3).
Xestecyathus Kru s e, 1982, p. 193 [*X. zigzag;
OD; holotype, Kru se , 1982, pl. 14,1–7, AM
F.83405, Sydney]. Inner wall with several rows of
pores per intersept, bearing upwardly projecting,
S-shaped scales; septa completely porous; tabulae
with normal pores. lower Cambrian (Bot.3):
Australia.——Fig. 62a–c. *X. zigzag, Cymbric
Vale Format ion , Bo tom an, Mt. Wrig ht, New
South Wales, Australia, holotype, AM F.83405; a,
tangential section of inner wall, AM FT.8526, ×5;
b, transverse section, AM FT.8527, ×5; c, longitu-
dinal section, AM FT.12793, ×5 (Kruse, 1982).
Family KASYRICYATHIDAE
Zhuravleva, 1961
[Kasyricyathidae ZhurAvlevA in musATov & others, 1961, p. 29]
Inner wall with communicating canals.
lower Cambrian (Bot.1).
Kasyricyathus Zhu rAvl evA in mus ATov & others,
1961, p. 30 [*K. schirokovae; OD; holotype,
mus AT ov & others, 1961, pl. 6,3–4, TsS GM
264/36, Novosibirsk]. Inner wall with one row of
horizontal to upwardly projecting, straight canals
per intersept; septa completely porous; tabulae
with normal pores. lower Cambrian (Bot.1): Altay
Sayan.——Fig. 61,2. *K. schirokovae, Balakhtinson
Formation, Botoman, Kazyr River, eastern Sayan,
Please make
a short
note at the
order level
explaining
that super-
families and
families are
not in alpha
order but
(roughly) in
order of wall
type.

Systematic Descriptions of the Archaeocyatha 83
2
Membranacyathus
Kasyricyathus
3a
3b
Agyrekocyathus
1
Fig. 61. Agyrekocyathidae, Kasyricyathidae, and Membranacyathidae (p. 82–83).
Altay Sayan, Russia, holotype, TsSGM 264/36,
transverse section, ×8 (Musatov & others, 1961).
Family MEMBRANACYATHIDAE
Debrenne, Zhuravlev, & Kruse, 2002
[Membranacyathidae Debrenne, ZhurAvlev, & Kruse, 2002, p. 1622]
Inner wall with microporous sheath. lower
Cambrian (Atd.1–Atd.2).
Membranacyathus roZA no v, 1960a, p. 664 [*M.
repinae; OD; holotype, roZ A no v , 1960a, fig.
1zh-z; roZ An ov, 1973, pl. 13,2, PIN 4297/15,
Moscow]. Inner wall with several rows of pores per
intersept and continuous microporous sheath; septa
completely porous; tabulae with normal pores.
lower Cambrian (Atd.1–Atd.2): Altay Sayan.——
Fig . 61,3a–b. *M. repinae, Adiak Formation,
Atdabanian, Mrassu River, Gornaya Shoria, Altay
Sayan, Russia, holotype, PIN 4297/15; a, oblique
transverse section, ×4.5 (Rozanov, 1960a); b, detail
of longitudinal section (outer wall to left), ×8
(Debrenne, Zhuravlev, & Kruse, 2002).
Superfamily
POLYCOSCINOIDEA
Debrenne, 1964
[nom. transl. Debrenne, ZhurAvlev, & Kruse, 2002, p. 1623, ex Polycoscinidae
Debrenne, 1964, p. 194] [=Anaptyctocyathoidea Debrenne, 1970a, p. 25, nom.
correct. Debrenne & Kruse, 1986, p. 260, pro Anaptyctocyathacea Debrenne,
1970a, p. 25; Lunulacyathacea Debrenne, 1973, p. 18, nom. nud.; Lunula-
cyathoidea Debrenne in Debrenne, ZhurAvlev, & roZAnov, 1989, p. 87]
Outer wall with attached microporous
sheath. lower Cambrian (Atd.3–Bot.3).

84 Treatise Online, number XX
a
c
b
Xestecyathus
Fig. 62. Xestecyathidae (p. 82).

Systematic Descriptions of the Archaeocyatha 85
Family ANAPTYCTOCYATHIDAE
Debrenne, 1970
[Anaptyctocyathidae Debrenne, 1970a, p. 25]
Inner wall with simple pores. l o wer
Cambrian (Atd.4–Bot.3).
Anaptyctocyathus Debrenne, 1969a, p. 340, nom.
transl. Debrenne, 1970a, p. 28, ex Alataucyathus
(Anaptyctocyathus) Debrenne, 1969a, p. 340
[*Coscinocyathus cribripora R. beDForD & W. R.
beDForD, 1934, p. 3; OD; lectotype, R. beDForD
& W. r. beD F or D , 1934, fig. 15; Debrenne,
1969a, pl. 11,1,3; Debrenne, 1973, pl. 2,6 ;
SD Debrenne, 1969a, p. 340, NHM S4160,
London]. Inner wall with several rows of simple
pores per intersept; septa completely porous;
tabulae with normal pores. lower Cambrian
(Atd.4–Bot.3): Australia, Antarctica.——Fi g .
63a–c. *A. cribripora (R. be D F o rD & W. r.
beDFo rD), Ajax Limestone, Botoman, Ajax Mine,
South Australia, Australia, lectotype, NHM
S4160; a, longitudinal view, ×5; b, transverse
view, ×5; c, detail of outer wall, ×15 (Debrenne,
Zhuravlev, & Kruse, 2002).
Family POLYCOSCINIDAE
Debrenne, 1964
[Polycoscinidae Debrenne, 1964, p. 194] [=Lunulacyathidae Debrenne,
1973, p. 18, nom. nud.; =Lunulacyathidae Debrenne in Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 87]
Inner wall with bracts or scales. lower
Cambrian (Atd.3–Bot.3).
Polycoscinus R. beDForD & J. beDForD, 1937, p.
37 [*P. contortus; OD; holotype, R. beDForD &
J. beD ForD , 1937, fig. 157; Debrenne, 1973, pl.
3,7; M; USNM PU87217, specimen 222, Wash-
ington, D.C.] [=Erugatocyathus Debrenne, 1969a,
p. 334 (type, Coscinocyathus papillatus R. beDFor D
& W. R. beDForD, 1934, p. 3; OD), nom. transl.
Debrenne, 1970a, p. 33, ex Tomocyathus (Erugatocy-
athus) Debrenne, 1969a, p. 334)]. Inner wall with
several rows of pores per intersept, bearing down-
wardly projecting, cupped bracts; septa sparsely
to completely porous; tabulae with normal pores.
lower Cambrian (Atd.3–Bot.3): Australia, Antarc-
tica.——Fig. 64a–c. *P. contortus, Ajax Limestone,
Atdabanian, Paint Mine, South Australia, Australia,
holotype, USNM PU87217, specimen 222; a,
transverse view of modular skeleton, ×3; b, detail
Anaptyctocyathus
c
a
b
Fig. 63. Anaptyctocyathidae (p. 85).

86 Treatise Online, number XX
of inner wall, tangential view, ×9; c, detail of outer
wall, tangential view, ×15 (Debrenne, Zhuravlev,
& Kruse, 2002).
Lunulacyathus Debrenne, 1973, p. 17 [*Coscinocy-
athus minimiporus R. beDForD & J. beDForD, 1937,
p. 37; OD; lectotype, R. beDForD & J. beDForD,
1937, fig. 155; Debrenne, 1973, pl. 4,6; SD
Debrenne, 1973, p. 17, USNM PU86705, Wash-
ington, D.C.]. Outer wall with attached micro-
porous sheath and supplementary cupped bracts;
inner wall with several rows of pores per intersept,
bearing downwardly projecting, cupped bracts;
septa completely porous; tabulae with normal pores.
lower Cambrian (Bot.3): Australia.——Fig. 65a–c.
*L. minimiporus (R. beDForD & j. be DForD), Ajax
Limestone, Botoman, Ajax Mine, South Australia,
Australia, lectotype, USNM PU86705; a, transverse
view, ×8; b, detail of outer wall in longitudinal
view, ×30 (Debrenne, Zhuravlev, & Kruse, 2002);
c, longitudinal view (outer wall to right), ×8 (M.
Debrenne, new).
Family VERONICACYATHIDAE
Debrenne, Zhuravlev, & Kruse, 2002
[Veronicacyathidae Debrenne, ZhurAvlev, & Kruse, 2002, p. 1625]
Inner wall with noncommunicating
canals. lower Cambrian (Atd.4–Bot.3).
Veronicacyathus Debrenne, 1973, p. 19 [*V. fron -
deus Debrenne, 1973, p. 20; OD; holotype,
Debrenne, 1973, pl. 2,4–5; pl. 3,1 ; pl. 4,8,
USNM PU86731, specimen 200, Washington,
D.C.; =Coscinocyathus tatei eTheriDge, 1890,
p. 18; lectotype, eTheriDge, 1890, pl. 3,2–4;
Debrenne, ZhurAv l ev , & grAve sT oc K, 1993,
fig. 3–4; SD Debrenne, Zhu rAvl ev , & gr Ave-
sTo cK, 1993, p. 182, choice following elimina-
tion of all other specimens by TAT e , 1892, p.
188, SAM T1245, Adelaide]. Inner wall with
several rows of horizontal to upwardly projecting,
straight canals per intersept, bearing spines
projecting radially across orifice to form screen;
c
a
bPolycoscinus
Fig. 64. Polycoscinidae (p. 85–86).

Systematic Descriptions of the Archaeocyatha 87
septa aporose to sparsely porous; tabulae with
normal pores. lower Cambrian (Atd.4–Bot.3):
Australia, Antarctica.——Fig. 66,1a–b. *V. tatei
(eTheriDge), Parara Limestone, Botoman, Pavy
Gully, Ardrossan, South Australia, Australia,
lectotype, SAM T1245; a, transverse section,
×3; b, longitudinal section (outer wall to left),
×3 (Debrenne, Zhuravlev, & Gravestock, 1993).
Bractocyathus Kru se , 1978, p. 41 [*B. labiosus;
OD; holotype, Kru se , 1978, fig. 11A–E, AM
F.83335, Sydney]. Inner wall with several rows
of horizontal to upwardly projecting, straight
canals per intersept; upwardly projecting, planar
bract arises within each canal, supporting
spines projecting radially to form screen; septa
completely porous; tabulae with normal pores.
lower Cambrian (Atd.4–Bot.3): Australia, Antarc-
tica.——Fig. 66,2a–d. *B. labiosus, Cymbric Vale
Formation, Botoman, Mt. Wright, New South
Wales, Australia; a– b, holotype, AM F.83335
; a, transverse section, AM FT.8302, ×4; b,
longitudinal section, AM FT.8303, ×4 (Kruse,
1978); c, paratype AM F.83278, detail of outer
wall, tangential section, AM FT.8317, ×10; d,
topotype, detail of inner wall, tangential section,
AM FT.10077, ×6 (Kruse, 1982).
Family ZONACOSCINIDAE
Debrenne, 1971
[Zonacoscinidae Debrenne, 1971, p. 194]
Inner wall with communicating canals.
lower Cambrian (Bot.1).
Zonacoscinus Debrenne, 1971, p. 194 [*Z. tumu-
losus; OD; holotype, Debrenne, 1971, fig. 3;
Debrenne, 1972, pl. 4,5–6, MNHN M84037,
specimen Ci 15U 21-2, Paris]. Inner wall with
several rows of horizontal to upwardly projecting,
b
Lunulacyathus
a
c
Fig. 65. Polycoscinidae (p. 86).

88 Treatise Online, number XX
Fig. 66. Veronicacyathidae (p. 86–87).
1a
2d
2c
2b
1b
Veronicacyathus
2a Bractocyathus

Systematic Descriptions of the Archaeocyatha 89
straight canals per intersept, canals branching
toward central cavity; septa completely porous;
tabulae with normal pores. lower Cambrian (Bot.1):
Sardinia.——Fig. 67,1. *Z. tumulosus, Matoppa
Formation, Botoman, Monte Cuccurinu, Sardinia,
Italy, holotype, MNHN M84037, specimen Ci
15U 21-2, transverse section (outer wall at top),
×10 (Debrenne, 1972).
Orienticyathus be lY A e v A , 1969, p. 95 [*O .
mamontovi; OD; holotype, be lY A e vA , 1969,
pl. 36,1–2, DVGU 6M, Khabarovsk]. Inner
wall with several rows of upright, V-shaped
canals per intersept; septa completely porous;
tabulae with normal pores; synapticulae may be
present. lower Cambrian (Bot.1): Far East.——
Fig . 67,2a– b . *O. mam onto vi, Ust’t oka uni t,
Botoman, Gerbikan River, Dzhagdy Range, Far
East, Russia; a, holotype, DVGU 6M/K8/3-3,
detail of transverse section (outer wall to right),
×13; b, paratype, DVGU 6M/K8/3-2, detail of
longitudinal section (outer wall to right), ×13
(Debrenne, Zhuravlev, & Kruse, 2002).
Superfamily
ETHMOCOSCINOIDEA
Zhuravleva, 1957
[nom. transl. Debrenne, roZAnov, & ZhurAvlev in Debrenne, ZhurAv-
lev , & roZAn ov, 1989, p. 87, ex Ethmocoscinidae Zh urAv levA i n
vologDin, 1957a, p. 181] [=Tumulocoscinacea ZhurAvlevA, 1960b, p.
265, nom. nud., nom. transl. roZAnov, 1973, p. 86, ex Tumulocoscininae
ZhurAvlevA, 1960b, p. 265; =Tumulocoscinoidea ZhurAvlevA, 1960b, p.
265, nom. transl. roZAnov in Debrenne, ZhurAvlev, & roZAnov, 1989,
p. 86, ex Tumulocoscininae ZhurAvlevA, 1960b, p. 265]
Outer wall with simple tumuli. lower
Cambrian (Atd.2–Bot.3).
Fig. 67. Zonacoscinidae (p. 87–89).
1Zonacoscinus
Orienticyathus
2a
2b

90 Treatise Online, number XX
Fig. 68. Tumulocoscinidae (p. 90–91).
23
4a
1a
1b
4b
Tumulocoscinus
Orbicoscinus
Retetumulus
Asterotumulus
Family TUMULOCOSCINIDAE
Zhuravleva, 1960
[nom. transl. Debrenne, 1970a, p. 25, ex Tumulocoscininae ZhurAvlevA,
1960b, p. 265]
Inner wall with simple pores. l o wer
Cambrian (Atd.2–Bot.1).
Tumulocoscinus Zhu rAv levA , 1960b, p. 265 [*T.
atdabanensis; OD; holotype, Zhur AvlevA, 1960b,
pl. 3,3b; pl. 23,10, PIN 1161, Moscow, not
located]. Inner wall with several rows of simple
pores per intersept; septa aporose to sparsely
porous; tabulae with normal pores. l ower
Cambrian (Atd.2–Bot.1): Siberian Platform,
Altay Sayan.——Fig . 68,1a–b. *T. atdabanensis,

Systematic Descriptions of the Archaeocyatha 91
Perekhod Formation, Atdabanian; a , Yudyay,
Lena River, Sakha (Yakutia), Russia, holotype,
PIN 1161, transverse section, ×12 (Zhuravleva,
1960b); b, Achagyy-Taryng Creek, Lena River,
Sakha (Yakutia), Russia, specimen TsSGM 323/91,
oblique longitudinal section, ×15 (Zhuravleva,
Korshunov, & Rozanov, 1969).
?Asterotumulus KAshinA in rePinA & others, 1964, p.
229 [*A. receptori; OD; holotype, rePinA & others,
1964, pl. 15,3, TsSGM KGU1313/61, Novosi-
birsk]. Outer wall with probable tumuli; inner
wall longitudinally plicate, with several rows of
simple pores per intersept; septa completely porous;
tabulae with normal pores. [Limited type material
does not provide certainty as to which wall is outer
and which inner, thereby creating uncertainty as to
whether accepted outer wall bears tumuli or bracts.]
lower Cambrian (Atd.3–Bot.1): Altay Sayan.——
Fig. 68,2. *A. receptori, Bazaikha Formation, Atda-
banian, Bazaikha River, eastern Sayan, Altay Sayan,
Russia, holotype, TsSGM KGU1313/61, transverse
section, ×7 (Repina & others, 1964).
Orbicoscinus Debrenne, 1977a, p. 111 [*O. schaerti;
OD; holotype, Debrenne, 1977a, pl. 10,4, MNHN
M80045, specimen IRH24-1c, Paris]. Cup in
which both walls show periodic, synchronous
transverse folds; inner wall with one row of simple
pores per intersept; septa completely porous;
tabulae with normal pores. lower Cambrian (Bot.1):
Moro cco .——Fig . 68,3. *O. schaerti, Issafen
Formation, Botoman, Jbel Irhoud, Morocco, holo-
type, MNHN M80045, specimen IRH24-1c,
longitudinal section, ×5 (Debrenne, 1977a).
Retetumulus Debrenne, 1977a, p. 112 [*R. dutuiti;
OD; holotype, Debrenne, 1977a, pl. 10,1, MNHN
M80042, specimen IRH34-3f, Paris]. Inner wall
with several rows of simple pores per intersept;
septa completely porous; tabulae with slitlike
pores. lower Cambrian (Bot.1): Morocco.——Fi g.
68,4a–b. *R. dutuiti, Issafen Formation, Botoman,
Jbel Irhoud, Morocco, holotype, MNHN M80042,
specimen IRH34-3f; a, detail of outer wall (at top)
in transverse section, ×20 (Debrenne, 1977a);
b, oblique transverse section, ×6 (Debrenne,
Zhuravlev, & Kruse, 2002).
Family ETHMOCOSCINIDAE
Zhuravleva, 1957
[Ethmocoscinidae ZhurAvlevA in vologDin, 1957a, p. 181]
Inner wall with noncommunicating
canals. lower Cambrian (Bot.3).
Ethmocoscinus simon, 1939, p. 28 [*Coscinocy-
athus papillipora R. beD For D & W. R. beDForD,
1934, p. 4; OD; holotype, R. beD Fo r D & W.
R. beD Fo rD, 1934, fig. 18; hill, 1965, pl. 8,7;
Debrenne, 1969a, pl. 3,3, NHM S4164, M,
London]. Inner wall with one row of horizontal
to upwardly projecting, S-shaped canals per inter-
sept; septa aporose to sparsely porous; tabulae
with normal pores. lower Cambrian (Bot.3):
Australia.——Fig. 69,1a–b. *E. papillipora (R.
beD Fo rD & W. r. beDF or D ), Ajax Limestone,
Botoman, Ajax Mine, South Australia, Australia,
holotype, NHM S4164; a, detail of outer wall
in longitudinal view, ×7; b, transverse view, ×4
(Hill, 1965).
Superfamily
COSCINOPTYCTOIDEA
Debrenne, Rozanov, &
Zhuravlev, 1989
[Coscinoptyctoidea Debrenne, roZ Anov, & ZhurAvlev in Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 86]
Outer wall with multiperforate tumuli.
lower Cambrian (Atd.1–Bot.3).
Family GEYERICOSCINIDAE
Debrenne & Zhuravlev, 2000
[Geyericoscinidae Debrenne & ZhurAvlev, 2000, p. 50]
Inner wall with simple pores. l o wer
Cambrian (Atd.1–Bot.1).
Geyericoscinus Debrenne & Zhu rAvl ev, 2000, p.
50 [*Coscinocyathus equiporus Debrenne, 1959b,
p. 8; OD; lectotype, De br enn e, 1959b, pl. 1,4;
Debrenne, 1964, pl. 28,4; SD Debrenne, 1963a,
p. 23, MNHN M80081, specimen TAI 1-5-4T,
Paris]. Inner wall with several rows of simple pores
per intersept; septa completely porous; tabulae
with normal pores. lower Cambrian (Atd.1–
Bot.1): Morocco.——Fig. 69,2a–b. *G. equiporus
(Debrenne), Amouslek Formation, Atdabanian,
Jbel Taïssa, Morocco, lectotype, MNHN M80081,
specimen TAI 1-5-4T; a, transverse section, ×5
(Debrenne, 1959b); b, detail of transverse section
(outer wall at bottom), ×15 (Debrenne, 1964).
Family COSCINOPTYCTIDAE
Debrenne, Rozanov, & Zhuravlev, 1989
[Coscinoptyctidae Debrenne, ro ZAnov , & ZhurAvlev in Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 86]
Inner wall with bracts or scales. lower
Cambrian (Bot.3).
Coscinoptycta broili, 1915, p. 121 [*Coscinoptycha
convoluta TAY lo r , 1910, p. 141; SD simon,
1939, p. 26; lectotype, TAYl or , 1910, fig. 7–8,
pl. 11, photo 60; SD Debrenne, Zh ur Avlev , &
Kruse , 2002, p. 1630, SAM T1594-6, Adelaide]
[=Coscinoptycha TAY lo r , 1910, p. 141 (type,
C. convoluta, SD simon, 1939, p. 26), non
meYricK, 1881, p. 700, insect]. Cup in which
both walls show synchronous transverse folds;
inner wall with several rows of pores per inter-
sept, bearing probably downwardly projecting,
cupped bracts; septa completely porous; tabulae
with normal pores. lower Cambrian (Bot.3):

92 Treatise Online, number XX
Australia, Antarctica.——Fi g . 70a–c. *C .
convoluta ( TAYl or ), Ajax Limestone, Botoman,
Ajax Mine, South Australia, Australia, lectotype,
SAM T1594-6; a, detail of outer wall in tangen-
tial section, ×7; b, transverse section (outer wall
at bottom), ×4; c, detail of transverse section
(outer wall at top), ×9 (Debrenne, Zhuravlev,
& Kruse, 2002).
Family JEBILETICOSCINIDAE
Debrenne, Rozanov, & Zhuravlev, 1989
[Jebileticoscinidae Debrenne, r oZAnov, & ZhurAvlev in Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 86]
Inner wall with noncommunicating
canals. lower Cambrian (Bot.1).
Fig. 69. Ethmocoscinidae and Geyericoscinidae (p. 91).
2a
2b
Geyericoscinus
1a Ethmocoscinus
1b

Systematic Descriptions of the Archaeocyatha 93
Jebileticoscinus Debrenne, 1977 a, p. 114 [*J.
huvelini; OD; holotype, Debrenne, 1977a, pl.
11,2, MNHN M80048, specimen IRH4-1d, Paris]
[=Pachycoscinus Debrenne, 1977a, p. 117 (type, P.
hollardi, OD), for discussion, see Debrenne,
Zhu r Av le v , & Kru s e , 2002, p. 1630]. Inner
wall with several rows of horizontal to upwardly
projecting, straight canals per intersept; septa
completely porous; tabulae with normal pores.
lower Cambrian (Bot.1): Morocco.——Fig. 71,1.
*J. huvelini, Issafen Formation, Botoman, Jbel
Irhoud, Morocco, holotype, MNHN M80048,
specimen IRH4-1d, longitudinal section, ×4
(Debrenne, 1977a).
Irhoudicoscinus Debrenne, 1977a, p. 117 [*I. destom-
besi; OD; holotype, Debrenne, 1977a, pl. 12,3–4,
MNHN M80052, specimen IRH2-1a, Paris].
Inner wall with one row of horizontal to upwardly
projecting, straight canals per intersept; septa
completely porous; tabulae with normal pores.
lower Cambrian (Bot.1): Morocco.——Fig. 71,2.
*I. destombesi, Issafen Formation, Botoman, Jbel
Irhoud, Morocco, holotype, MNHN M80052,
specimen IRH2-1a, oblique transverse section, ×4
(Debrenne, Zhuravlev, & Kruse, 2002).
Superfamily
SIGMOCOSCINOIDEA
R. Bedford & J. Bedford, 1939
[nom. correct. Debrenne & Kruse, 1986, p. 264, pro Sigmocoscinacea
Debrenne, 1970a, p. 25, nom. transl. ex Sigmocoscinidae R. beDForD &
J. beDForD, 1939, p. 76]
Outer wall with bracts or scales. lower
Cambrian (Bot.1–Bot.3).
Family SYLVIACOSCINIDAE
Debrenne, Rozanov, & Zhuravlev, 1989
[Sylviacoscinidae Debrenne, r oZAnov, & ZhurAvlev in Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 87]
Inner wall with simple pores. l o wer
Cambrian (Bot.1–Bot.3).
Sylviacoscinus Debrenne in Zhur Avl evA , 1974b,
p. 119 [*Coscinocyathus sylvia R. beDF or D & J.
beDForD, 1937, p. 37; OD; holotype, R. beD ForD
& J. beDForD, 1937, fig. 156, USNM PU86706,
specimen 221, M, Washington, D.C.]. Outer wall
with upwardly projecting, denticulate, curved
Fig. 70. Coscinoptyctidae (p. 91–92).
b
a
c
Coscinoptycta

94 Treatise Online, number XX
scales; inner wall with one row of simple pores
per intersept; septa aporose to sparsely porous;
tabulae with normal pores. lower Cambrian (Bot.3):
Australia.——Fig. 72a–c. *S. sylvia (R. beDFo rD
& j. beDFor D), Ajax Limestone, Botoman, Ajax
Mine, South Australia, Australia, holotype, USNM
PU86706, specimen 221; a, detail of outer wall,
longitudinal view, ×15; b, transverse view, ×7; c,
detail of inner wall, internal longitudinal view, ×9
(Debrenne, Zhuravlev, & Kruse, 2002).
Family SIGMOCOSCINIDAE
R. Bedford & J. Bedford, 1939
[Sigmocoscinidae R. beDForD & J. beDForD, 1939, p. 76]
Inner wall with annuli. lower Cambrian
(Bot.3).
Sigmocoscinus R. be DFor D & J. beDF orD, 1936, p.
24 [*S. sigma; OD; lectotype, R. beDF or D & J.
beDFo rD, 1936, fig. 98; SD hill, 1965, p. 111,
USNM PU86686, specimen 235, Washington,
D.C.]. Outer wall with upwardly projecting,
S-shaped scales; inner wall with one pore row per
intersept and upwardly projecting, S-shaped annuli;
septa completely porous; tabulae with normal
pores. lower Cambrian (Bot.3): Australia, Antarc-
tica.——Fig. 73,1a–d. *S. sigma, Ajax Limestone,
Botoman, Ajax Mine, South Australia, Australia,
lectotype, USNM PU86686, specimen 235; a,
transverse view, ×8; b, detail of inner wall, internal
longitudinal view, ×11; c, detail of outer wall,
longitudinal view, ×11; d, longitudinal view, ×8
(Debrenne, Zhuravlev, & Kruse, 2002).
?Statanulocyathus Debrenne, 1975, p. 342 [*S.
oosthuizeni; OD; holotype, Debrenne, 1975, fig.
7a–b, SAM(C) K44945, Cape Town]. Outer wall
with upwardly projecting, cupped bracts; inner
wall with one row of pores per intersept, bearing
upwardly projecting, cupped bracts; upwardly
projecting, arcuate annuli at each tabula; septa
completely porous; tabulae with normal pores.
[Inner wall structure comprises bracts together
with unusual, hypertrophied annuli, the appro-
priate taxonomic treatment of which is uncer-
tain.] lower Cambrian (Bot.3): South Africa
(allochthonous).——Fig . 73,2. *S. oosthuizeni,
Dwyka Subgroup, Botoman (allochthonous in
Permian), Zwartskraal, South Africa, holotype,
SAM(C) K44945, oblique longitudinal section,
×10 (Debrenne, 1975).
Fig. 71. Jebileticoscinidae (p. 93).
2
Jebileticoscinus
1
Irhoudicoscinus

Systematic Descriptions of the Archaeocyatha 95
Superfamily
POROCOSCINOIDEA
Debrenne, 1964
[nom. transl. Debrenne, Zh urAvlev, & Kruse, 2002, p. 1633, ex Poro-
coscinidae Debrenne, 1964, p. 190] [=Rozanovicyathacea Korshunov in
ZhurAvlevA, Korshunov, & roZAnov, 1969, p. 54; =Schumnyicyathoidea
Debrenne, ZhurAvlev, & roZAnov, 1989, p. 87, nom. transl. Debrenne,
ZhurAvlev, & Kruse, 2002, p. 1635, ex Schumnyicyathidae Debrenne, ro-
ZAnov, & ZhurAvlev in Debrenne, ZhurAvlev, & roZAnov, 1989, p. 87]
Outer wall with canals. lower Cambrian
(Atd.1–Bot.3).
Family ROZANOVICYATHIDAE
Korshunov, 1969
[Rozanovicyathidae Korshunov in ZhurAvlevA, Korshunov, & ro ZA-
nov, 1969, p. 54]
Inner wall with simple pores. l o wer
Cambrian (Bot.1).
Rozanovicyathus Korshunov in Zhu r Av l e vA ,
Korshunov, & roZAnov, 1969, p. 54 [*R. alexi; OD;
holotype, Zhur AvlevA, Korshunov, & roZ Anov,
1969, pl. 23,1,3, TsSGM 323/93, Novosibirsk].
Outer wall with horizontal to upwardly projecting,
S-shaped canals, bearing supplementary bracts exter-
nally (imparting overall inverted V-shaped appearance
to outer wall); inner wall with one row of simple pores
per intersept; septa completely porous; tabulae with
slitlike pores. lower Cambrian (Bot.1): Siberian Plat-
form.——Fig. 74,1. *R. alexi, Mukhatta Formation,
Botoman, Mukhatta River, Sakha (Yakutia), Russia,
holotype, TsSGM 323/93, transverse section, ×5
(Zhuravleva, Korshunov, & Rozanov, 1969).
Family TATIJANAECYATHIDAE
Korshunov, 1976
[Tatijanaecyathidae Korshunov, 1976, p. 149] [=Schumnyicyathidae
Debrenne, roZ Anov, & ZhurAvlev i n Debrenne, ZhurAvlev, & ro-
ZAnov, 1989, p. 87]
Inner wall with annuli. lower Cambrian
(Bot.1).
Muchattocyathus roZ An ov in F. Debrenne, M.
Debrenne, & roZ Anov, 1976, p. 103 (roZ Anov,
1973, p. 61, nom. nud.) [*M. sibiricus; OD; holo-
type, F. Debrenne, M. Debrenne, & ro ZA nov ,
1976, pl. 1,5, PIN 4597/142, Moscow] [=Tatijan-
aecyathus Korshunov, 1976, p. 149 (type, T. laetus,
OD), for discussion, see Debrenne, Zhur Avlev, &
roZAnov , 1989, p. 120; Debrenne, roZAn ov, &
ZhurAvlev, 1990, p. 152]. Outer wall with down-
wardly projecting, straight canals, bearing supple-
mentary bracts externally (imparting overall upright
V-shaped appearance to outer wall); inner wall with
one pore row per intersept and upwardly projecting,
S-shaped annuli; septa completely porous, linked
by synapticulae; tabulae with normal pores. lower
Cambrian (Bot.1): Siberian Platform.——Fig. 74,2.
*M. sibiricus, Oy-Muran reef massif, Botoman,
Oy-Muran, Lena River, Sakha (Yakutia), Russia,
holotype, PIN 4597/142, oblique transverse section,
×7 (F. Debrenne, M. Debrenne, & Rozanov, 1976).
Schumnyicyathus ZhurAvlevA in DATsenKo & others,
1968, p. 164 [*S. validus; OD; holotype, DATsenKo &
Fig. 72. Sylviacoscinidae (p. 93–94).
Sylviacoscinus
b
a
c

96 Treatise Online, number XX
Fig. 73. Sigmocoscinidae (p. 94).
Statanulocyathus
1d
1c
1a
1b
Sigmocoscinus
2

Systematic Descriptions of the Archaeocyatha 97
others, 1968, pl. 9,3, TsSGM 278/72, Novosibirsk].
Outer wall with horizontal to upwardly projecting,
S-shaped canals and supplementary attached micro-
porous sheath; inner wall with several pore rows per
intersept and upwardly projecting, S-shaped annuli;
septa completely porous; tabulae with normal pores.
lower Cambrian (Bot.1): Siberian Platform.——Fig.
74,3 a–b. *S. validus, Shumnoy Formation, Botoman,
Sukharikha River, Krasnoyarsk region, Russia; a,
holotype, TsSGM 278/72, transverse section, ×12;
b, paratype, TsSGM 278/73, oblique longitudinal
section, ×12 (Datsenko & others, 1968).
Family POROCOSCINIDAE
Debrenne, 1964
[Porocoscinidae Debrenne, 1964, p. 190]
Inner wall with noncommunicating
canals. lower Cambrian (Atd.3–Bot.3).
Porocoscinus Debrenne, 1964, p. 190 [*P. flexi-
bilis; OD; holotype, Debrenne, 1964, pl. 28,1–3,
MNHN M84108, specimen S Sc 5-4b, Paris]
[=Coscinoteichus Debrenne, 1964, p. 180 (type,
C. minimiporus, OD); =Chengkoucyathus YuA n,
1974, p. 81 (type, C. shabaensis, OD); =Flexicy-
athus Kruse, 1978, p. 40 (type, F. rudens, OD), for
discussion, see Debrenne, ZhurAvlev, & roZAnov,
1989, p. 124; Debrenne, roZAnov, & ZhurAvlev,
1990, p. 155; Debrenne, ZhurAvle v, & Kru se,
2002, p. 1635]. Outer wall with horizontal to
upwardly projecting, S-shaped canals, bearing
supplementary bracts externally (imparting overall
inverted V-shaped appearance to outer wall); inner
wall with several rows of inverted V-shaped canals
per intersept; septa completely porous; tabulae
with normal pores. lower Cambrian (Bot.1–Bot.3):
Australia, Antarctica, southern China, Iberia,
Sardinia.——Fig. 75,1a–b. *P. flexibilis, Matoppa
Formation, Botoman, Serra Scoris, Sardinia,
Italy, holotype, MNHN M84108, specimen S Sc
5-4b; a, transverse section (outer wall at top), ×8
(Debrenne, Zhuravlev, & Kruse, 2002); b, longi-
tudinal section (outer wall to left), ×8 (Debrenne,
1964).
Geniculicyathus Debrenne, 1960, p. 118 [*G. varius;
M; holotype, Debrenne, 1960, fig. A, MNHN
M80154, specimen HD40, Paris]. Outer wall with
horizontal to upwardly projecting, straight canals,
bearing supplementary bracts externally (imparting
overall inverted V-shaped appearance to outer wall);
inner wall with several rows of horizontal to upwardly
projecting, S-shaped canals per intersept; septa
completely porous; tabulae with normal pores. lower
Cambrian (Atd.3): Morocco.——Fig. 75,2a–b. *G.
varius, Amouslek Formation, Atdabanian, Jbel Taïssa,
Morocco, holotype, MNHN M80154, specimen
HD40; a, transverse section, ×4; b, longitudinal
section, ×4 (Debrenne, Zhuravlev, & Kruse, 2002).
Tubicoscinus Debrenne in Debrenne, ZhurAvlev, &
roZAn ov, 1989, p. 137 (Tubicoscinus Debrenne,
1970b, p. 207, nom. nud., proposed conditionally,
ICZN Art. 15) [*Coscinocyathus tuba bornemAnn,
1884, p. 704; OD; holotype, bornemAnn, 1886,
pl. 15,2a; Debrenne, 1964, pl. 18,1–2, GML 930,
Halle]. Outer wall with horizontal to upwardly
projecting, straight canals, bearing supplemen-
tary bracts externally (imparting overall inverted
V-shaped appearance to outer wall); inner wall
with one row of horizontal to upwardly projecting,
slightly S-shaped canals per intersept; septa
completely porous; tabulae with normal pores.
lower Cambrian (Bot.1): Iberia, Sardinia.——Fig.
75,3a–b. *T. tuba (bornemAnn), Matoppa Forma-
tion, Botoman, San Pietro, Sardinia, Italy, holotype,
GML 930; a, longitudinal section, ×7 (Borne-
mann, 1886); b, detail of same, ×14 (Debrenne,
1964).
Superfamily
MOOTWINGEECYATHOIDEA
Kruse, 1982
[nom. transl. Debrenne, roZAnov, & ZhurAvlev in Debrenne, ZhurAv-
lev, & roZAnov, 1989, p. 87, ex Mootwingeecyathidae Krus e, 1982,
p. 194]
Outer wall clathrate. lower Cambrian
(Bot.3).
Family MOOTWINGEECYATHIDAE
Kruse, 1982
[Mootwingeecyathidae Kruse, 1982, p. 194]
Inner wall with bracts or scales. lower
Cambrian (Bot.3).
Mootwingeecyathus Kru se , 1982, p. 195 [*M .
mootwingeensis; OD; holotype, Kr us e , 1982,
fig. 20–21, pl. 15,4–11, AM F.83344, Sydney].
Inner wall with several rows of pores per inter-
sept, bearing upwardly projecting, S-shaped
scales; septa completely porous; tabulae
with normal pores. lower Cambrian (Bot.3):
Australia.——Fig. 76a–d. *M. mootwingeensis,
Cymbric Vale Formation, Botoman, Mt. Wright,
New South Wales, Australia, holotype, AM
F.83344; a, oblique longitudinal section, AM
FT.14162, ×8; b, detail of outer wall in tangen-
tial section, AM FT.8175, ×30; c, transverse
section, AM FT.14163, ×7; d, detail of inner
wall, transverse section, AM FT.14163, ×3 0
(Kruse, 1982).
Order PUTAPACYATHIDA
Vologdin, 1961
[Putapacyathida vologDin, 1961, p. 177]
Intervallum with plate tabulae;
redimiculi may be present on intervallum
side of either or both walls. lower Cambrian
(Bot.1–Bot.3).

98 Treatise Online, number XX
Superfamily
PUTAPACYATHOIDEA
R. Bedford & J. Bedford, 1936
[nom. correct. Debrenne, ZhurAvlev, & Kruse, 2002, p. 1638, pro Puta-
pacyathacea Debrenne, 1970a, p. 24, nom. transl. ex Putapacyathidae R.
beDForD & J. beDForD, 1936, p. 24] [=Alphacyathoidea R. beDForD &
J. beDForD, 1939, p. 72, nom. transl. Debrenne, roZAnov, & ZhurAvlev
in Debrenne, ZhurAvlev, & ro ZAnov, 1989, p. 88, ex Alphacyathidae
R. beDForD & J. beDForD, 1939, p. 72; =Aptocyathacea KonYushKov
in ZhurAvlevA, KonYushKov, & roZ Anov , 1964, p. 102, nom. nud .,
nom. transl. roZ Anov, 1973, p. 85, ex Aptocyathidae KonYushKov in
ZhurAvlevA, KonYushKov, & roZAnov, 1964, p. 102]
Outer wall with simple pores. lowe r
Cambrian (Bot.1–Bot.3).
Family ALPHACYATHIDAE
R. Bedford & J. Bedford, 1939
[Alphacyathidae R. beDForD & J. beDForD, 1939, p. 72] [=Aptocyathidae
KonYushKov in ZhurAvlevA, KonYushKov, & roZAnov, 1964, p. 102]
Inner wall with simple pores. l o wer
Cambrian (Bot.1–Bot.3).
Alphacyathus R. beDF or D & J. beDF or D, 1939,
p. 72 [*Dictyocyathus annularis R. beD Fo r D &
W. R. beDF or D , 1936, p. 13; OD; lectotype,
R. be DF or D & W. R. beD F or D, 1936, fig. 55;
ZhurAvlevA, KonYushKov, & ro ZA nov , 1964,
fig. 53; SD Debrenne, 1969a, p. 305, SAM
P942, Adelaide; =Dictyocyathus simplex TAYlo r,
Fig. 74. Rozanovicyathidae and Tatijanaecyathidae (p. 95–97).
Schumnyicyathus
3a
3b
2Muchattocyathus
Rozanovicyathus 1

Systematic Descriptions of the Archaeocyatha 99
3a
3b
Geniculicyathus
2a
2b
Tubicoscinus
1a
1b
Porocoscinus
Fig. 75. Porocoscinidae (p. 97).

100 Treatise Online, number XX
1910, p. 144; lectotype, TAYl or , 1910, fig. 34;
SD Debrenne, Zhu rAv le v, & Kru se , 2002, p.
1638, SAM T1598A, B, Adelaide]. Inner wall
with stirrup pores at each tabula and one file of
simple, intertabular pores; tabulae with normal
pores; longitudinal lintels form septumlike plates
in some intertabulae. lower Cambrian (Bot.3):
Australia.——Fig. 77,1a–c. *A. simplex (TAYlor),
Ajax Limestone, Botoman, Ajax Mine, South
Australia, Australia; a, lectotype, SAM T1598,
longitudinal view, ×12; b, paralectotype, USNM
PU86714, specimen 225, transverse view, ×12
(Debrenne, Zhuravlev, & Kruse, 2002); c, para-
lectotype, NHM S4822, longitudinal view, ×12
(M. Debrenne, new).
Aptocyathus vologDin, 1937b, p. 471 [*A. gordoni;
M; lectotype, SD ZhurAvlevA, KonYushKov, &
roZ An ov , 1964, p. 106, collection IK Bazhe-
nova, spec ime n 28-40 48, thi n sec tion 3/10 ,
not located] [=Aptocyathella Kon Y u sh K ov in
ZhurAvlevA, KonYushKov, & ro ZA nov , 1964,
p. 111 (type, A. prima, OD); =Galinaecyathus
KonYushKov in Zhur Avl evA , KonYushKov, &
roZ A n ov , 1964, p. 102 (type, G. lebedensis,
OD); =Arthrocyathus vo lo g Di n , 1977, p. 61
(type, A. articulatus, OD)]. Inner wall with
several files of simple pores per intertabulum;
tabulae with normal pores. lower Cambrian
(Bot.1–Bot.2): Altay Sayan, Urals, Australia,
Iberia, Sardinia.——Fi g . 77,2a–b. *A .
gordoni, Verkhnemonok Formation, Botoman,
Sanashtykgol River, western Sayan, Altay Sayan,
Russia; a, topotype, possibly of type series, PIN
4754/42, transverse section of modular skeleton,
×10 (Vologdin, 1940b); b, unlocated syntype,
sketch of transverse and longitudinal sections of
modular skeleton, ×10 (Vologdin, 1937b).
Family PUTAPACYATHIDAE
R. Bedford & J. Bedford, 1936
[Putapacyathidae R. beDForD & J. beDForD, 1936, p. 24]
Inner wall with bracts or scales. lower
Cambrian (Bot.3).
Putapacyathus R. beDForD & J. beDForD, 1936, p.
24 [*P. regularis; OD; holotype, R. beDForD & J.
beDForD, 1936, fig. 97; ZhurAvlevA, KonYushKov,
& roZ Ano v, 1964, fig. 68; hill, 1965, pl. 9,1,
USNM PU86699-115, M, Washington, D.C.].
Inner wall with several files of pores per intertab-
ulum, bearing upwardly projecting cupped bracts;
tabulae with normal pores; sporadic septa may be
present. lower Cambrian (Bot.3): Australia.——Fig.
78a–c. *P. regularis, Ajax Limestone, Botoman, Ajax
Mine, South Australia, Australia, holotype, USNM
Fig. 76. Mootwingeecyathidae (p. 97).
Mootwingeecyathus
d
c
a
b

Systematic Descriptions of the Archaeocyatha 101
PU86699-115; a, transverse view, ×6; b, oblique
longitudinal view, ×6 (Debrenne, Zhuravlev, &
Kruse, 2002); c, internal longitudinal view of inner
wall, ×4.5 (Hill, 1965).
Superfamily HUPECYATHOIDEA
Debrenne, Rozanov, & Zhuravlev,
1990
[Hupecyathoidea Debrenne, roZAnov, & ZhurAvlev, 1990, p. 121]
Outer wall with canals. lower Cambrian
(Atd.4).
Family HUPECYATHIDAE
Debrenne, Rozanov, & Zhuravlev, 1990
[Hupecyathidae Debrenne, roZAnov, & ZhurAvlev, 1990, p. 121]
Inner wall with bracts or scales. lower
Cambrian (Atd.4).
Hupecyathus Debrenne, 1964, p. 198 [*H. sphinc-
toides; OD; holotype, Debrenne, 1964, pl. 15,7,
MNHN M80258, specimen Ki135, Paris]. Outer
wall with horizontal to upwardly projecting,
straight canals, bearing supplementary bracts
externally (imparting overall inverted V-shaped
appearance to outer wall); inner wall with several
files of pores per intertabulum, bearing upwardly
projecting, cupped bracts; tabulae with normal
pores, linked by pillars. lower Cambrian (Atd.4):
Morocco.——Fig . 79a–b. *H. sphinctoides, Amou-
slek Formation, Atdabanian, Ouijane, Morocco,
holotype, MNHN M80258, specimen Ki135; a,
transverse section, ×6; b, detail of longitudinal
section (outer wall to right), ×13 (Debrenne ,
1964).
Superfamily
CHABAKOVICYATHOIDEA
Rozanov, 2002
[Chabakovicyathoidea roZAn ov i n Debrenne, ZhurAvlev, & Kruse,
2002, p. 1639] [=Chabakovicyathacea roZAnov, 1973, p. 85, nom. nud.]
Outer wall pustular. lower Cambrian
(Bot.1).
Alphacyathus
2a
2b
1c
Aptocyathus
1a
1b
Fig. 77. Alphacyathidae (p. 98–100).

102 Treatise Online, number XX
Putapacyathus
a
b
c
Fig. 78. Putapacyathidae (p. 100–101).

Systematic Descriptions of the Archaeocyatha 103
Family CHABAKOVICYATHIDAE
Rozanov, 2002
[Chabakovicyathidae roZA nov in Debrenne, ZhurAvlev , & Kruse,
2002, p. 1639] [=Chabakovicyathidae roZAnov, 1973, p. 85, nom. nud.]
Inner wall with simple pores. l o wer
Cambrian (Bot.1).
Chabakovicyathus Kon Yu s hK ov in Zh ur Av le vA ,
Kon Yu s hK ov , & roZA n ov , 1964, p. 114 [*C.
tumulatus; OD; holotype, ZhurAvlevA, KonY-
ushKov, & roZAnov, 1964, pl. 14,6, not located].
Inner wall with several files of simple pores per
intertabulum; tabulae with normal pores. lower
Cambrian (Bot.1): Urals.——Fig . 80a–b. *C.
tumulatus, Terekla Formation, Botoman, Terekla
River, western flank of Southern Urals, Russia; a,
longitudinal section, specimen PIN 4327/80, ×16
(Debrenne, Zhuravlev, & Kruse, 2002); b, holo-
type, oblique transverse section, ×28 (Zhuravleva,
Konyushkov, & Rozanov, 1964).
Order CAPSULOCYATHIDA
Zhuravleva, 1964
[nom. transl. ZhurAvle v & roZ Anov in voronovA & others, 1987, p.
29, ex Capsulocyathina Zhu rAvlevA in ZhurAvl evA, Ko nYus hKov, &
roZAnov , 1964, p. 59] [=Coscinocyathida Zhu rAvlevA, 1955a, p. 10;
=Clavicyathida vologDin, 1977, p. 110]
Thalamid cup, single- or multichambered;
inner wall of invaginal type of development;
septa and/or plate tabulae may be present in
intervallum of multichambered cups. lower
Cambrian (Tom.1–Bot.3).
Suborder CAPSULOCYATHINA
Zhuravleva, 1964
[Capsulocyathina ZhurAvlevA in ZhurAvlevA, KonYushKov, & roZAnov,
1964, p. 59] [?=Uralocyathina Debrenne, 1964, p. 113]
Cup single chambered and subspherical,
or multichambered without septa. lower
Cambrian (Tom.1–Bot.3).
Family CRYPTOPOROCYATHIDAE
Zhuravleva, 1960
[Cryptoporocyathidae ZhurAvlevA, 1960b, p. 92] [=Cryptaporocyathidae
ZhurAvlevA, 1963b, p. 117, nom. null.; =Capsulocyathidae ZhurAvlevA in
ZhurAvlevA, KonYushKov, & roZAnov, 1964, p. 60; =Gerbicanicyathidae
belYAevA, 1969, p. 90; =Vasicyathidae vologDin, 1977, p. 104; =Clavicyathidae
vologDin, 1977, p. 110; ?=Complicatocyathidae YAroshevich, 1990, p. 23]
Outer wall with simple pores. lowe r
Cambrian (Tom.1–Bot.3).
Cryptoporocyathus Zhur AvlevA, 1960b, p. 92 [*C.
junicanensis; OD; holotype, Zhu rAvlevA, 1960b,
pl. 4,9; ZhurAvlevA, 1963b, pl. 12,3), not located;
paratype, TsSGM 205/6, Novosibirsk] [=Crypta-
porocyathus Zhu rAv l evA , 1963b, p. 117, nom.
null.]. Cup single chambered; outer wall pores
of two distinct sizes; inner wall simple. l ow er
Cambrian (Tom.1–Tom.4): Siberian Platform.——
Fig . 81,1a–b. *C. junicanensis; a, Medvezh’ya
Formation, Tommotian, Moyero River, Krasnoyarsk
region, Russia, holotype, oblique section, ×20; b,
Pestrotsvet Formation, Tommotian, Aldan River,
Sakha (Yakutia), Russia, paratype, TsSGM 205/6,
section of outer wall (inner cavity at bottom), ×16
(Debrenne, Zhuravlev, & Kruse, 2002).
Hupecyathus
a
b
Fig. 79. Hupecyathidae (p. 101).

104 Treatise Online, number XX
Capsulocyathus vologDin in Zh u r A v l e vA ,
KonYushKov, & roZAnov, 1964, p. 61 (vologDin,
1962c, p. 75 (type, C. capsulifer, OD), nom. nud.,
not described, figured or separately diagnosed
until vologDin, 1977, p. 76, fig. 44, wherein a
holotype was invalidly nominated from material
other than type or topotype material) [*C. subcal-
losus ZhurAvlevA in ZhurAvlevA, KonYushKov, &
roZAnov, 1964, p. 62; OD; holotype, ZhurAvlevA,
KonYushKov, & roZAnov, 1964, pl. 1,8(3), TsSGM
283/5, Novosibirsk] [=Capsulicyathus vologDin,
1977, p. 75 (type, C. capsulifer, OD), nom. van.;
=Mesocyathus vologDin, 1977, p. 95 (type, M.
plasticus, OD); =Vasicyathus vologDin, 1977, p. 104
(type, V. urniformis, OD); =Clavicyathus vologDin,
1977, p. 110 (type, C. clavellatus, OD), for discus-
sion, see Debrenne, ZhurAvlev, & roZAnov, 1989,
p. 97; Debrenne, roZAnov, & ZhurAvlev, 1990, p.
135]. Cup single chambered; outer and inner walls
with simple pores. lower Cambrian (Tom.3–Bot.3):
Altay Sayan, Tuva, Mongolia, Transbaikalia, Far
East, Urals, Morocco, Iberia, Sardinia.——Fig. 81,2.
*C. subcallosus, Bazaikha Formation, Atdabanian,
Bazaikha River, eastern Sayan, Altay Sayan, Russia,
holotype, TsSGM 283/5, longitudinal section, ×6
(Zhuravleva, Konyushkov, & Rozanov, 1964).
?Complicatocyathus YAroshevich, 1990, p. 23 [*C.
rozanovi; OD; holotype, YAroshevich, 1990,
pl. 12,1, TsSGM 901/5a, Novosibirsk]. Cup
with regular transverse folds affecting both walls,
forming empty multichambered cups; outer and
inner walls with simple pores. [Limited type mate-
rial does not provide certainty as to whether cup is
multichambered.] lower Cambrian (Atd.2): Altay
Sayan.——Fig. 81,3. *C. rozanovi, Gavrilovskoe
Formation, Atdabanian, Gavrilovskoe, Salair,
Russia, holotype, TsSGM 901/5a, longitudinal
section, ×4 (Yaroshevich, 1990).
Gerbicanicyathus belYAevA, 1969, p. 90 [*G. emili;
OD; holotype, belYAevA, 1969, pl. 37,1, DVGU
55/68, Khabarovsk]. Cup multichambered; outer
and inner walls with simple pores. lower Cambrian
(Bot.1–Bot.2): Far East.——Fig . 82,1. *G. emili,
Ust’toka unit, Botoman, Gerbikan River, Dzhagdy
Range, Far East, Russia, holotype, DVGU 55/68,
oblique longitudinal section, ×8 (Belyaeva, 1969).
Mirandocyathus belYAevA, 1974, p. 121 [*M. artus;
OD; holotype, belYA evA, 1974, pl. 3,8, DVGU
13M/572/3, Khabarovsk]. Cup multichambered;
outer wall with simple pores; inner wall with pores
bearing downwardly projecting, cupped bracts.
lower Cambrian (Bot.1): Far East.——Fig . 82,2.
*M. artus, Ust’toka unit, Botoman, Gerbikan River,
Dzhagdy Range, Far East, Russia, holotype, DVGU
13M/572/3, longitudinal section, ×7 (Belyaeva,
1974).
Polythalamia Debrenne & Woo D , 1990, p. 436
[*P. americana; OD; holotype, Debrenne &
WooD , 1990, fig. 1B, USNM 434924, specimen
GA5.18F, Washington, D.C.]. Cup multicham-
bered, globose chambers propagating linearly or
glomerately; outer wall with few or no pores;
inner wall with simple pores. lower Cambrian
(Bot.1–Bot.2): Altay Sayan, Tuva, Mongolia,
United States.——Fig . 82,3. *P. americana,
Valmy Formation, Botoman, Galena Canyon,
Nevada, United States, holotype, USNM 434924,
specimen GA5.18F, longitudinal section, ×20
(Debrenne & Wood, 1990; copyright Cambridge
University Press).
Fig. 80. Chabakovicyathidae (p. 103).
Chabakovicyathus
a
b

Systematic Descriptions of the Archaeocyatha 105
Family URALOCYATHELLIDAE
Zhuravleva, 1964
[Uralocyathellidae ZhurAvlevA in ZhurAvlevA, KonYushKov, & roZAnov,
1964, p. 72]
Outer wall with independent micropo-
rous sheath. lower Cambrian (Bot.1).
Rhabdolynthus Zhu r Av le vA , 1960b, p. 91 [*R.
conicus; OD; holotype, ZhurAvlevA, 1960b, pl.
4,8; ZhurAvlevA, 1963b, pl. 9,11–12, TsSGM
205/5, Novosibirsk] [=Uralocyathella ZhurAvlevA
in Zhu rAv levA , KrAsn oP eevA , & ch er nY sh evA,
1960, p. 99 (type, U. repinae, OD); =Miricyathus
vologDin, 1977, p. 88 (type, M. aseptatus, OD),
for discussion, see Debrenne, Zhu r Av le v , &
roZAn ov, 1989, p. 127; Debrenne, ro ZAnov , &
ZhurAvlev, 1990, p. 157]. Cup single chambered;
outer wall with independent microporous sheath;
inner wall with simple pores. lower Cambrian
(Bot.1): Siberian Platform, Altay Sayan.— —
Fig. 82,4a–b. *R. conicus, Perekhod Formation,
Botoman, Atdaban, Lena River, Sakha (Yakutia),
Russia, holotype, TsSGM 205/5; a, longitudinal
section, ×4; b, detail of wall in longitudinal
section (inner cavity to right), ×7.5 (Zhuravleva,
1963b).
Family TYLOCYATHIDAE
Zhuravlev, 1988
[Tylocyathidae ZhurAvlev, 1988, p. 106]
Outer wall with attached microporous
sheath. lower Cambrian (Tom.4–Bot.3).
Tylocyathus vologDin, 1977, p. 102 (vologDin in
vologDin & YAZmir, 1966, p. 948, nom. nud.)
[*T. inaequilateralis vologDin, 1977, p. 102; OD;
holotype, vologDin, 1977, pl. 1,7, PIN 1924-41,
Moscow; =Uralocyathella bullata Zh urAv levA in
Fig. 81. Cryptoporocyathidae (p. 103–104).
Capsulocyathus
3
Cryptoporocyathus
Complicatocyathus
1a
1b
2

106 Treatise Online, number XX
musATov & others, 1961, p. 19; OD; holotype,
musATov & others, 1961, pl. 1,7, TsSGM 264/7,
Novosibirsk]. Cup single chambered; outer wall
with attached microporous sheath, each micro-
pore bearing a supplementary bract; inner wall
with simple pores. lower Cambrian (Bot.1–Bot.3):
Altay Sayan, Far East.——Fi g. 83,1. *T. bullatus
(ZhurAvlevA), Balakhtinson Formation, Botoman,
Kazyr River, eastern Sayan, Altay Sayan, Russia,
holotype, TsSGM 264/7, oblique longitudinal
section, ×5.5 (Musatov & others, 1961).
Korshunovicyathus ZhurAvlev in Debrenne,
Zhur Avle v, & roZAno v, 1988, p. 99 [*Crypta-
porocyathus melnikovi Korshunov & Zh urAvlevA,
1967, p. 5; OD; holotype, Korshunov & ZhurAv-
levA, 1967, pl. 1,1, TsSGM 247/1, Novosibirsk].
Cup single chambered; outer wall with attached
microporous sheath; inner wall with simple pores.
lower Cambrian (Tom.4–Atd.1): Siberian Plat-
form.——Fig. 83,2a–b. *K. melnikovi (Korshunov
& ZhurAvlevA); a, Tyuser Formation, Atdabanian,
Ulakhan-Ald’arkhay Creek, Lena River, Tuora-Sis
Range, Sakha (Yakutia), Russia, holotype, TsSGM
247/1, detail of transverse section, ×20 (Korshunov
& Zhuravleva, 1967); b, Pestrotsvet Formation,
Atdabanian, Isit’, Lena River, Sakha (Yakutia),
Fig. 82. Cryptoporocyathidae and Uralocyathellidae (p. 104–105).
Rhabdolynthus
2
3
4a
4b
1
Polythalamia
Mirandocyathus
Gerbicanicyathus

Systematic Descriptions of the Archaeocyatha 107
3b
Fig. 83. Tylocyathidae and Fransuasaecyathidae (p. 105–108).
Korshunovicyathus
4c
3a
4a
4b
2b
1
Yukonensis
Fransuasaecyathus
Tylocyathus
2a

108 Treatise Online, number XX
Russia, specimen PIN 4220/117, transverse section,
×8 (Debrenne, Zhuravlev, & Rozanov, 1988).
Family FRANSUASAECYATHIDAE
Debrenne, 1964
[Fransuasaecyathidae Debrenne, 1964, p. 113] [=Acanthopyrgidae
hAnDFielD, 1971, p. 31]
Outer wall with simple tumuli. lower
Cambrian (Atd.1–Bot.3).
Fransuasaecyathus Zhu rA vl e vA , 1960b, p. 103
[*F. subtumulatus; OD; holotype, Zh ur Avl evA ,
1960b, pl. 5,5, TsSGM 205/10, Novosibirsk]
[=Bullicyathus vologDin, 1977, p. 105 (type,
B. pyxidatus, OD); =Marginicyathus vologDin,
1977, p. 107 (type, M. cardiosimilis, OD)]. Cup
single chambered; outer wall with simple tumuli;
inner wall with simple pores. lower Cambrian
(Atd.1–Bot.3): Siberian Platform, Mongolia,
Transbaikalia, Far East.——Fi g. 83,3a–b. *F.
subtumulatus, Perekhod Formation, Atdabanian,
Yudyay, Lena River, Sakha (Yakutia), Russia,
holotype, TsSGM 205/10; a, detail of wall in
oblique longitudinal section, ×20; b, oblique
longitudinal section, ×8(Debrenne, Zhuravlev,
& Kruse, 2002).
Yukonensis ÖZDiKmen, 2009, p. 216, n om. nov.
pr o Acanthopyrgus hAnDFielD, 1967, p. 209,
non DescAmPs & WinTreberT, 1966, p. 28
(type, Geloius finoti bolivAr, 1905, p. 285,
OD), insect [*Acanthopyrgus yukonensis hAnD-
FielD, 1967, p. 209; OD; holotype, hAnDFielD,
1967, pl. 23,1,4, GSC 21059, Ottawa]. Cup
multichambered with a tabula of rods linked by
synapticulae and an external thorny corolla at
each chamber junction; outer wall with simple
tumuli; inner wall with simple pores. l owe r
Cambrian (Bot.1): Canada, United States.——
Fig. 83,4a–c. *Y. yukonensis (hAnDFielD); a –b,
Adams Argillite, Botoman, Tatonduk River,
Alaska, United States; a , transverse section,
specimen USGS 5156-CO (A1), ×6 (Debrenne,
Zhuravlev, & Kruse, 2002); b , transverse
section, specimen USGS 5156-CO (A4), ×10
(Nitecki & Debrenne, 1979); c, Sekwi Forma-
tion, Botoman, Mackenzie Mountains, Yukon
Territory, Canada, holotype, GSC 21059, longi-
tudinal section, ×4.5 (M. Debrenne, new).
Family TUBERICYATHIDAE
Debrenne, Rozanov, & Zhuravlev, 1989
[Tubericyathidae Debrenne, roZ Anov , & ZhurAvlev in Debrenne,
ZhurAvl ev, & roZAnov , 1989, p. 89] [=Tubericyathidae vologDin,
1977, p. 31, nom. nud.]
Outer wall with bracts or scales. lower
Cambrian (Bot.1).
Tubericyathus vologDin, 1977, p. 90 [*T. clath-
ratus; OD; holotype, vologDin, 1977, fig. 57,
pl. 13,9, PIN 1924/38, Moscow] [=Arminocyathus
vologDin, 1977, p. 113 (type, A. fungiformis,
OD)]. Cup single chambered; outer wall with pores
bearing upwardly projecting, cupped bracts; inner
wall with simple pores. lower Cambrian (Bot.1):
Altay Sayan, Mongolia.——Fig . 84a–b. *T. clath-
ratus, Usa Formation, Botoman, Sukhie Solontsy
Valley, Batenev Range, Kuznetsk Alatau, Russia;
a, holotype, PIN 1924/38, oblique longitudinal
section, ×10; b, paratype, PIN 1924/39, oblique
transverse section, ×10 (Vologdin, 1977).
Suborder COSCINOCYATHINA
Zhuravleva, 1955
[nom. transl. ZhurAvlevA, 1 960b, p. 245 , ex order Coscin ocyath ida
ZhurAvlevA, 1955a, p. 25]
Cup multichambered; intervallum with
septa. lower Cambrian (Atd.1–Bot.3).
Fig. 84. Tubericyathidae (p. 108).
b
aTubericyathus

Systematic Descriptions of the Archaeocyatha 109
Fig. 85. Coscinocyathidae, Mawsonicoscinidae, and Coscinocyathellidae (p. 110).
Coscinocyathus
32a
2b
Coscinocyathellus
1a
1b
Mawsonicoscinus
I had to crop
view b on the
top (now the
rigtht side)
to make it fit.
Okay? Also,
do you have a
higher resolu-
tion file? This
one is only
100 dpi!!

110 Treatise Online, number XX
Superfamily
COSCINOCYATHOIDEA
Taylor, 1910
[nom. correct. Debrenne, ZhurAvlev, & roZAnov, 1989, p. 89, pro Coscino-
cyathacea ZhurAvlevA, 1960b, p. 245, nom. transl. ex Coscinocyathidae TAY-
lor, 1910, p. 137] [=Mawsonicoscinoidea Debrenne & Kruse, 1986, p. 258]
Outer wall tabular with simple pores.
lower Cambrian (Atd.2–Bot.3).
Family COSCINOCYATHIDAE
Taylor, 1910
[Coscinocyathidae TAYlor, 1910, p. 137] [=Poletaevacyathidae vologDin,
1962a, p. 125].
Inner wall with simple pores. l o wer
Cambrian (Atd.2–Bot.3).
Coscinocyathus bor n em A nn , 1884, p. 704 [*C.
dianthus; SD by exercise of ICZN plenary powers
by melville, 1974, p. 155, following application
by Debr enn e, 1970b, p. 207, negating SD of C.
tuba bornemAnn, 1884, p. 704 by Ting, 1937,
p. 360 (now type of Tubicoscinus Debrenne in
Debrenne, ZhurAvlev, & roZA no v, 1989, p.
137); lectotype, bornemAnn, 1886, pl. 17,2–7;
Debrenne, 1964, pl. 21,1–2; SD Debrenne,
1964, p. 169, GML An597, Halle] [=Poletaevacy-
athus vologDin, 1959b, p. 88 (type, P. obrutchevi,
M)]. Inner wall with several rows of simple pores
per intersept; septa completely porous; tabulae
with normal pores. lower Cambrian (Atd.2–Bot.3):
Siberian Platform, Altay Sayan, Tuva, Mongolia,
Transbaikalia, Far East, Tajikistan, Morocco,
Iberia, Sardinia.——Fig. 85,1a–b. *C. dianthus,
Matoppa Formation, Botoman, Canal Grande,
Sardinia, Italy, lectotype, GML An597; a, trans-
verse section, ×4; b, longitudinal section, ×4
(Bornemann, 1886).
Family MAWSONICOSCINIDAE
Debrenne & Kruse, 1986
[Mawsonicoscinidae Debrenne & Kruse, 1986, p. 258]
Inner wall with noncommunicating
canals. lower Cambrian (Bot.3).
Mawsonicoscinus Debrenne & Krus e, 1986, p.
259 [*M. sigmoides; OD; holotype, Debrenne
& Kruse , 1986, fig. 22, GNS MG513, Lower
Hutt]. Inner wall with one row of horizontal
to upwardly projecting, S-shaped canals per
intersept; septa completely porous; tabulae with
normal pores. lower Cambrian (Bot.3): Antarc-
tica.——Fig . 85,2a –b. *M. sigmoides, Sh ack -
leton Limestone, Holyoake Range, Nimrod
Glacier, Antarctica, holotype, GNS MG513;
a, longitudinal section (outer wall to right),
×3.5; b, transverse section, ×3.5 (Debrenne &
Kruse, 1986).
Family COSCINOCYATHELLIDAE
Zhuravleva, 1956
[Coscinocyathellidae ZhurAvlevA in vologDin, 1956, p. 879]
Inner wall with communicating canals.
lower Cambrian (Bot.1–Bot.3).
Coscinocyathellus vologDin, 1940b, p. 91 (vologDin,
1937b, p. 471, nom. nud.) [*C. parvus; OD; lectotype,
vologDin, 1940b, pl. 29,1; SD ZhurAvlev, 2001a,
p. 92, PIN 4754/4, Moscow]. Inner wall with several
rows of horizontal to upwardly projecting, straight to
waved canals per intersept; septa completely porous;
tabulae with normal pores. lower Cambrian (Bot.1–
Bot.3): Altay Sayan.——Fig. 85,3. *C. parvus, Verkh-
nemonok Formation, Botoman, Sanashtykgol River,
western Sayan, Altay Sayan, Russia, lectotype, PIN
4754/4, oblique longitudinal section, ×8 (Vologdin,
1940b).
Superfamily
CALYPTOCOSCINOIDEA
Debrenne, 1964
[nom. correct. Debrenne, roZA nov , & Zh ur Av lev in Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 89, pro Calyptocoscinacea Debrenne,
1964, p. 115]
Outer wall tabular with independent micro-
porous sheath. lower Cambrian (Atd.1–Bot.1).
Family TOMOCYATHIDAE
Debrenne, Rozanov, & Zhuravlev, 1989
[Tomocyathidae Debrenne, roZA nov , & ZhurAvl ev in Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 89]
Inner wall with simple pores. l o wer
Cambrian (Atd.1–Atd.3).
Coscinocyathella vologDin, 1959b, p. 87–88
(vologDin, 1957d, p. 699, nom. nud.) [*C. nikitini;
M; holotype, vologDin, 1957d, fig. 1(11), PIN
1800/1,1a, M, Moscow] [=Tomocyathus roZAnov,
1960a, p. 664 (type, T. operosus, OD), for discussion,
see Debrenne, Zhu rAvlev, & roZA nov, 1989, p.
100; Debrenne, roZAnov, & ZhurAvlev, 1990, p.
138]. Inner wall with several rows of simple pores
per intersept; septa completely porous; tabulae with
normal pores. lower Cambrian (Atd.1–Atd.3): Altay
Sayan, Tuva, Mongolia.——Fig. 86a–b. *C. nikitini,
Usa Formation, Atdabanian, Kiya River, Kuznetsk
Alatau, Russia, holotype, PIN 1800/1,1a; a, transverse
section, ×4; b, longitudinal section, ×4.5 (Debrenne,
Zhuravlev, & Kruse, 2002).
Family CALYPTOCOSCINIDAE
Debrenne, 1964
[Calyptocoscinidae Debrenne, 1964, p. 115]
Inner wall with independent microporous
sheath. lower Cambrian (Bot.1).

Systematic Descriptions of the Archaeocyatha 111
Coscinocyathella
a
b
Fig. 86. Tomocyathidae (p. 110).
Calyptocoscinus Debrenne, 1964, p. 196 [*Coscino-
cyathus cornucopiae bornemAnn, 1884, p. 704;
OD; lectotype, bornemAnn, 1886, pl. 16,1; SD
Debrenne, 1964, p. 196, GML block B, Halle,
requires restudy]. Inner wall with several rows
of pores per intersept and independent micro-
porous sheath; septa completely porous; tabulae
wi th nor mal pores . low e r C amb ria n (Bo t .1):
Sardinia.——Fig. 87a–e. *C. cornucopiae (borne-
mAnn), Matoppa Formation, Botoman, Monte
Gloria, Canal Grande, Sardinia, Italy, topotype,
MNHN M84106; a, oblique transverse section,
×6; b, transverse section, ×7; c, detail of inner
wall, longitudinal section (central cavity to right),
×30; d, detail of transverse section (outer wall
at bottom), ×10; e, longitudinal section, ×5
(Debrenne, 1964).
Superfamily
ALATAUCYATHOIDEA
Zhuravleva, 1955
[nom. correct. Debrenne, ZhurAvlev, & ro ZAn ov, 1989, p. 89 pr o
Alataucyathacea ZhurAvlevA, 1960b, p. 264, nom. transl. ex Alataucyathi-
dae ZhurAvlevA, 1955b, p. 626] [=Mrassocyathoidea vologDin, 1960,
p. 130, nom. correct. Debrenne, ZhurAvlev, & roZAnov, 1989, p. 85,
pro Mrassucyathacea ZhurAvlevA & roZAnov in rePinA & others, 1964,
p. 230, nom. transl. ex Mrassocyathidae vologDin in ZhurAvlevA, KrAs-
noPeevA, & chernYshevA, 1960, p. 130, as Mrassucyathidae nom. null.]
Outer wall tabular with multiperforate
tumuli. lower Cambrian (Atd.1–Atd.2).
Family ALATAUCYATHIDAE
Zhuravleva, 1955
[Alataucyathidae ZhurAvlevA, 1955b, p. 626] [=Mrassocyathidae volog-
Din, 1956, p. 879, nom. nud.; =Alataucyathinae Zhur AvlevA, 1955b,
p. 626, nom. transl. Zhur AvlevA , 1960b, p. 264, ex Alataucyathidae
ZhurAvlevA, 1955b, p. 626; =Mrassocyathidae vologDin in ZhurAvlevA,
KrAsnoPeevA, & chernYshevA, 1960, p. 130, nom. correct. Debrenne,
roZAnov, & ZhurAvlev in Debrenne, ZhurAvlev, & roZAnov, 1989,
p. 86 pro Mrassucyathidae vologDin in ZhurAvle vA, KrAs noPeevA, &
chernYshevA, 1960, p. 130]
Inner wall with simple pores. l o wer
Cambrian (Atd.1–Atd.2).
Alataucyathus ZhurAvlevA, 1955b, p. 626 [*A. jaros-
chevitschi; OD; holotype, Zhur AvlevA, 1955b, fig.
1a, 2g–d, PIN 1040, Moscow, not located] [=Mras-
socyathus KrAsnoPeevA in vologDin, 1956, p. 879,
nom. nud.; =Mrassocyathus Kr Asn oP ee vA, 1960,
p. 43 (type, M. micropora, OD); =Mrassucyathus
KrAsnoPeevA in ZhurAvlevA, KrAsnoPeevA, & cher-
nYshevA, 1960, p. 130 (type, M. schoriensis, OD),
nom. null.]. Inner wall with several rows of simple
pores per intersept; septa completely porous; tabulae
with normal pores. lower Cambrian (Atd.1–Atd.2):
Altay Sayan, Tuva, Mongolia.——Fig. 88a–b. *A.
jaroschevitschi; a, Usa Formation, Atdabanian, Mt.
Martyukhina, Kuznetsk Alatau, Russia, holotype,
PIN 1040, oblique longitudinal section, ×4; b, Usa
Formation, Atdabanian, Sukhie Solontsy Valley,
Batenev Range, Kuznetsk Alatau, Russia, unlocated
specimen, neither the holding institution nor the
collection number are known, transverse section, ×4
(Debrenne, Zhuravlev, & Kruse, 2002).
Superfamily
CLATHRICOSCINOIDEA
Rozanov, 1964
[nom. correct. ZhurAvlev, 1988, p. 105, pro Clathricoscinacea Debrenne,
1964, p. 115, nom. transl. ex Clathricoscinidae roZAnov in rePinA &
others, 1964, p. 223]
Outer wall tabular and pseudoclathrate.
lower Cambrian (Bot.1–Toy.1).
Family CLATHRICOSCINIDAE
Rozanov, 1964
[Clathricoscinidae roZAnov in rePinA & others, 1964, p. 223]
Inner wall with simple pores. l o wer
Cambrian (Bot.1–Toy.1).

112 Treatise Online, number XX
Calyptocoscinus
a
c
b
d
e
Fig. 87. Calyptocoscinidae (p. 111).
Clathricoscinus Zh u r Av l e vA , 1955b, p. 627
[*Coscinocyathus infirmus vologDin in ZhurAv-
le vA, 1955b, p. 627; OD; holotype, ZhurAv-
levA, 1955b, fig. 2a, PIN 1040, Moscow; collec-
tion not located] [=Asterocyathellus vologDin,
1962a, p. 126 (type, A. compositus, OD), for
discussion, see Debrenne, ZhurAvlev, &
roZ A no v, 1989, p. 99; Debrenne, roZ An o v,
& ZhurAvlev, 1990, p. 136]. Inner wall with
several rows of simple pores per intersept;

Systematic Descriptions of the Archaeocyatha 113
septa completely porous; tabulae with normal
pores. lower Cambrian (Bot.1–Toy.1): Kolyma,
Altay Sayan, Tuva, Mongolia, Transbaikalia,
Far East, southern China.——Fig . 89,1a–b.
*C. infirmus (vologDin), Usa Formation,
Botoman, Bol’shaya Erba, Batenev Range,
Kuznetsk Alatau, Russia, syntype, PIN 1040; a,
transverse section, ×10; b, tangential section of
outer wall (at top), ×10 (Debrenne, Zhuravlev,
& Kruse, 2002).
Family LANICYATHIDAE
Debrenne, Rozanov, & Zhuravlev, 1989
[Lanicyathidae Debrenne, roZA nov , & ZhurAvlev in Debrenne,
ZhurAvlev, & roZAnov, 1989, p. 90]
Inner wall with noncommunicating
canals. lower Cambrian (Bot.1).
Lanicyathus belYAevA in belYAevA & others, 1975, p.
87 [*L. albus; OD; holotype, belYA evA & others,
1975, pl. 19,3; pl. 37,2–3, PIN DVIMS5157/6,
Moscow]. Inner wall with several rows of horizontal
to upwardly projecting, straight to waved canals
per intersept; septa completely porous; tabulae
with normal pores. lower Cambrian (Bot.1): Far
East.——Fig. 89,2a–b. *L. albus, Ust’toka unit,
Botoman, Lan River, Dzhagdy Range, Far East,
Russia, holotype, PIN DVIMS5157/6; a, longi-
tudinal section, ×7; b, transverse section, ×7
(Debrenne, Zhuravlev, & Kruse, 2002).
Order ARCHAEOCYATHIDA
Okulitch, 1935
[nom. correct. Zhu rAvle vA, 1955a, p. 11, pro order Archaeocyathina
oKuliTch, 1935, p. 90] [=Anthomorphida oKuliTch, 1935, p. 90,
nom. correct. oKuliTch, 1955a, p. 18, pro order Anthomorphina
oKuliTch, 1935, p. 90; =Syringocnemidida oKuliTch, 1935, p. 90,
nom. correct. Debrenne, 1964, p. 117, pro order Syringocnemina
oKuliTch, 1935, p. 90; =Spirocyathida R. beDForD & W. R. beD-
ForD , 1936, p. 13, nom. correct. hill, 1972, p. 103, pro ord er Spiro -
cyathina R. beDForD & W. R. be DFor D, 1936, p. 13; =Metacyathida
R. beDForD & W. R. beDForD, 1936, p. 16, nom. correct. oKuliTch,
1955a, p. 14, pro order Metacyathina R. beDForD & W. R. beDForD,
1936, p. 16; =order Dictyocyanthina (sic) R. beDForD & J. beDForD,
1937, p. 37, nom. nud., proposed conditionally; =Archaeosyconida
ZhurAv levA , 1955a, p. 12; =superorder Loculicyathina ZhurAvle vA,
1955a, p. 9, nom. transl. vologDin, 1962a, p. 118, ex order Locu-
licyathida Zhur Avle vA, 1955a, p. 9, nom. correct. vologDin, 1961,
p. 178, pro Loculocyathida ZhurAvlevA, 1955a, p. 9, invalid name
based on nom. null.; =Rhizacyathida ZhurAvlevA, 1955b, p. 629, for
discussion, see hill, 1972, p. 103, 133; =Bicyathida vologDin 1956,
p. 878; =Syringocnematida ZhurAvl evA in Zhu rAvlevA, K rAsnoPeevA,
& chernYshevA, 1960, p. 139; =Thalassocyathida vologDin, 1961,
p. 177; =Tersiida vologDin, 1961, p. 181; = superorder Bicyathina
vologDin, 1962a, p. 117; =Archaeopharetrida Debrenne, 1970a,
p. 25; =Metaldetida Debrenne, 1970a, p. 25; =Paranacyathida
Debrenne, 1970a, p. 25; =Paracoscinida Debrenne, 1970a, p. 25,
for discussion, see Debrenne & ZhurAvle v, 1992b, p. 111; =Meta-
coscinida Debrenne, 1974a, p. 187]
Cup two walled, of solitary or low- to
high-modular organization, with septal
type of development; inner wall of centrip-
etal type; intervallum with taeniae, pseu-
dosepta, pseudotaenial network, dictyonal
network, calicles, or syringes; segmented
or independent (membrane and plate)
tabulae may be present. lower Cambrian
(Tom.1–Toy.3), middle Cambrian, upper
Cambrian (Furongian).
Alataucyathus a
b
Fig. 88. Alataucyathidae (p. 111).

114 Treatise Online, number XX
Lanicyathus
2a
2b
Clathricoscinus
1a
1b
Fig. 89. Clathricoscinidae and Lanicyathidae (p. 112–113).
Suborder LOCULICYATHINA
Zhuravleva, 1955
[nom. transl. Debrenne, 1991, p. 219, ex superorder Loculicyathina
ZhurAvlevA, 1955a, p. 9, nom. transl. vologDin, 1962a, p. 118, ex order
Loculicyathida vologDin, 1961, p. 178, nom. correct. pro Loculocyathida
ZhurAvlevA, 1955a, p. 9, invalid name based on nom. null.]
Cup modular (pseudocolonies formed by
interparietal budding) or rarely solitary; inter-
vallum with pseudosepta; synapticulae and
plate tabulae may be present. lower Cambrian
(Tom.1–Bot.3), upper Cambrian (Furongian).
Superfamily
LOCULICYATHOIDEA
Zhuravleva, 1954
[nom. transl. et correct. Debrenne & Zhur Avle v, 1992b, p.112, e x
Loculocyathidae Zhu rAvl evA, 1954, p. 27, invalid name based on
nom. null.]
Outer wall simple, with pores
of cambroid type. lower Cambrian
(Tom.1–Bot.3); upper Cambrian
(Furongian).

Systematic Descriptions of the Archaeocyatha 115
Family LOCULICYATHIDAE
Zhuravleva, 1954
[nom. correct. ZhurAvlevA, 1960b, p.107, pro Loculocyathidae ZhurAvl-
evA, 1954, p. 27, invalid name based on nom. null.] [=Robustocyathidae
Debrenne, 1964, p. 113; =Paranacyathidae Debrenne, 1970a, p. 38, nom.
nud.; =Ardrossacyathidae grAvesTocK, 1984, p. 109]
Inner wall with simple pores. l o wer
Cambrian (Tom.1–Bot.3), upper Cambrian
(Furongian).
Loculicyathus vologDin, 1931, p. 54 (vologDin,
1928, p. 30, nom. nud.) [*L. tolli; M; lecto-
type, vologDin, 1931, pl. 19,1; SD Debrenne,
ZhurAvl ev, & Krus e, 2002, p. 1651, TsNIGRm
58a/2956, St. Petersburg] [=Loculocyathus
vologDin, 1937b, p. 468, nom. null.]. Inner wall
with one row of simple pores per intersept; pseudo-
septa finely porous. lower Cambrian: (Atd.2–Bot.3):
Siberian Platform, Altay Sayan, Tuva, Mongolia,
Transbaikalia, Far East, Urals, Australia, Iberia,
Sardinia, United States, Mexico.——Fig. 90,1a–b.
*L. tolli, Torgashino Formation, Atdabanian,
Kameshki, eastern Sayan, Altay Sayan, Russia; a,
lectotype, TsNIGRm 58a/2956, transverse section,
×8.5; b, paralectotype, TsNIGRm 57a/2956, longi-
tudinal section, ×6 (Vologdin, 1931).
?Antarcticocyathus Debrenne, roZ A n o v , &
Webers, 1984, p. 298 [*A. webersi; OD; holo-
type, Debrenne, roZ Anov, & Webers, 1984, fig.
5.1–5.2, 6.3, USNM 333901, specimen Ant-1,
Washington, D.C.]. Outer wall pores in irregular,
undulating quasitransverse (or less commonly
quasilongitudinal) rows over entire wall plate; inner
wall with one row of simple pores per intersept;
pseudosepta coarsely porous. [Genus is otherwise
typical of suborder, but bears a continuous outer
wall of distinctive porosity, the appropriate taxo-
nomic treatment of which is uncertain.] upper
Cambrian (Furongian: Paibian): Antarctica.——
Fig . 90,2a–b. *A. webersi, Minaret Formation,
Furongian: Paibian, Springer Peak, Heritage Range,
Ellsworth Mountains, Antarctica; a, holotype,
USNM 333901, specimen Ant-1, longitudinal
section, ×4.5 (Debrenne, Rozanov, & Webers,
1984); b, paratype, USNM 333906, specimen
Ant-2, oblique transverse section, ×4 (Debrenne,
Zhuravlev, & Kruse, 2002).
Ardrossacyathus R. beD ForD & J. beDForD , 1937, p.
31 [*A. endotheca; OD; holotype, R. beDFo rD &
J. beDFor D, 1937, fig. 125, M, USNM PU86766,
specimen 354, Washington, D.C.] [=Met a l -
detimorpha R. beDForD & J. beDForD, 1937,
p. 31 (type, M. yorkei, OD), for discussion, see
ZhurAvlev & grAvesTocK, 1994, p. 31; =Dzhag-
dycyathus belYAevA in belYAevA & others, 1975, p.
102 (type, D. crinitus, OD); =Egiinocyathus Fonin,
1983, p. 12 (type, E. ornatus, OD), for discus-
sion, see Debrenne & ZhurAvlev, 1992b, p. 121;
Zhur Avle v & gr Ave sTo cK, 1994, p. 31]. Inner
wall with several rows of simple pores per intersept;
pseudosepta finely porous. lower Cambrian (Bot.1–
Bot.3): Mongolia, Far East, Australia.——Fig .
90,3. *A. endotheca, Botoman, Parara Limestone,
Ardrossan, South Australia, Australia, topotype,
SAM P32041, tangential section of outer wall, ×7
(Zhuravlev & Gravestock, 1994).
Cambrocyathellus ZhurAvlevA, 1960b, p. 284 [*C.
tschuranicus; OD; holotype, ZhurAvlevA, 1960b,
pl. 28,3, PIN 1161, Moscow, not located] [=Robus-
tocyathus ZhurAvlevA, 1960b, p. 133 (type, Archaeo-
cyathus robustus vologDin, 1937a, p. 25, OD);
=Ramuscyathus (Ramuscyathus) Fonin in voronin &
others, 1982, p. 101 (type, Loculocyathus tuberculatus
vologDin, 1940a, p. 87, OD; =R. (R.) artus Fonin
in voronin & others, 1982, p. 102; =Ramuscyathus
(Parvuscyathus) Fonin in voronin & others, 1982,
p. 103 (type, R. (P.) pannonicus, OD), for discus-
sion, see Debrenne & ZhurAvlev, 1992b, p. 122].
Outer wall pores restricted to interseptal areas;
inner wall with one row of simple pores per inter-
sept; pseudosepta coarsely porous. lower Cambrian
(Tom.1–Atd.4): Siberian Platform, Altay Sayan, Tuva,
Mongolia, Far East, Kazakhstan, Australia.——
Fig. 91,1a–b. *C. tschuranicus, Pestrotsvet Forma-
tion, Tommotian; a, Churan, Lena River, Sakha
(Yakutia), Russia, holotype, PIN 1161, transverse
section, ×5.5; b, Krestyakh, Lena River, Sakha
(Yakutia), Russia, specimen PIN 1161, longitudinal
section of modular skeleton (outer wall to right), ×4
(Debrenne, Zhuravlev, & Kruse, 2002).
Mikhnocyathus mAslov, 1957, p. 307 [*M. zolaensis;
OD; lectotype, mAslo v, 1957, fig. 2; Debrenne
& Zhu rAvle v, 1992b, pl. 5,6; SD Debrenne &
Zhu rAv l ev , 1992b, pl. 5, fig. 6 caption, PIN
2038(1), Moscow] [=Zolacyathus vologDin,
1962d, p. 10 (type, Z. loculosus, M)]. Inner wall
with several rows of simple pores per intersept;
pseudosepta coarsely porous; rare plate tabulae.
lower Cambrian (Atd.2–Atd.3): Altay Sayan, Tuva,
Mongolia, Transbaikalia, ?Sardinia.——Fi g .
91,2a–b. *M. zolaensis, Bystraya Formation, Atda-
banian, Zola Valley, Transbaikalia, Russia, lecto-
type, PIN 2038(1); a, transverse section, ×3.5; b,
longitudinal section, ×3.5 (Maslov, 1957).
Neoloculicyathus voronin, 1974, p. 134 [*N. primus;
OD; holotype, voronin, 1974, pl. 6,4; voronin,
1979, pl. 12,5; Debrenne & ZhurAvlev, 1992b, pl.
1,5, PIN 2742/4, Moscow]. Inner wall with several
rows of simple pores per intersept; pseudosepta
coarsely porous. lower Cambrian (Atd.1–Bot.1):
Siberian Platform, Altay Sayan, Tuva, Mongolia,
Transbaikalia, Far East, Urals, Australia, Morocco,
Iberia, Germany.——Fig. 91,3a–b. *N. primus,
Bazaikha Formation, Atdabanian, Bazaikha River,
eastern Sayan, Altay Sayan, Russia; a, paratype, PIN
2742/3, longitudinal section, ×6; b, holotype, PIN
2742/4, longitudinal section, ×4 (Voronin, 1974).
Okulitchicyathus ZhurAvlevA, 1960b, p. 281 [*Ajaci-
cyathus discoformis ZhurAvl evA in Zhur AvlevA &
Zelenov, 1955, p. 68; OD; holotype, Zh urAvlevA
& Zelenov, 1955, pl. 1,1, PIN 100(1), Moscow, not
located] [=Lermontovaecyathus Korshunov, 1972,
p. 59 (type, L. isiti, OD; =Ajacicyathus discoformis
ZhurAvlevA in ZhurAvlevA & Zelenov, 1955, p. 68,
for discussion, see Debrenne & ZhurAvlev, 1992b,

116 Treatise Online, number XX
Fig. 90. Loculicyathidae (p. 115).
3
2a
2b
Antarcticocyathus
Loculicyathus
1a
Ardrossacyathus
1b

Systematic Descriptions of the Archaeocyatha 117
2a
2b
Cambrocyathellus
Neoloculicyathus
1a Mikhnocyathus
1b
Fig. 91. Loculicyathidae (p. 115).
3a 3b

118 Treatise Online, number XX
p. 128); =Alconeracyathus Perejón, 1973, p. 185
(type, A. melendezi, OD; =Archaeocyathellus (Archaeo-
fungia) andalusicus simon, 1939, p. 76); =Anda-
lusicyathus Perejón in Debrenne, 1975, p. 352,
nom. nud.; =Andalusicyathus Perejón, 1976, p. 17
(type, Archaeocyathellus (Archaeofungia) andalusicus
simon, 1939, p. 76, OD); =Urdacyathus Perejón &
moreno, 1978, p. 201 (type, U. pradoanus, OD),
for discussion, see Debrenne & ZhurAvlev, 1992b,
p. 119]. Inner wall with several rows of simple
pores per intersept; pseudosepta coarsely porous;
plate tabulae and synapticulae may be present.
lower Cambrian (Tom.1–Atd.4): Siberian Platform,
Mongolia, Kazakhstan, ?Australia, Iberia.——Fig .
92,1a–b. *O. discoformis (ZhurAvlevA), Pestrotsvet
Formation, Tommotian; a, Aldan River, Sakha
(Yakutia), Russia, specimen PIN 1162, external
view of inner wall, ×6 (Zhuravleva, 1960b); b,
Churan-Zhurinskiy Mys area, Lena River, Sakha
(Yakutia), Russia, specimen MNHN M810058,
transverse section (outer wall at top), ×6 (Debrenne,
Zhuravlev, & Kruse, 2002).
Paranacyathus R. beDForD & J. beDForD, 1937, p.
34, nom. nov. pro Paracyathus R. beDForD & W.
R. beDForD, 1936, p. 17, non milne- eD WAr Ds
& hAime, 1848, p. 318 (type, P. procumbens, SD
milne-eDWArDs & hAime, 1850, p. xv), cnidarian
[*Paracyathus parvus R. beDForD & W. R. beDForD,
Fig. 92. Loculicyathidae (p. 115–119).
2a
2b
Okulitchicyathus
Paranacyathus
1a
1b

Systematic Descriptions of the Archaeocyatha 119
1936, p. 17; OD; holotype, R. be DF or D & W.
r. beDForD, 1936, fig. 76; Debrenne, 1974c, pl.
19,1–4; Debrenne & ZhurAvlev, 1992b, pl. 1,6),
SAM P992-134, -135, M, Adelaide]. Inner wall with
one, rarely two rows of simple pores per intersept;
pseudosepta finely porous. lower Cambrian (Bot.1–
Bot.3): ?Altay Sayan, ?Canada, Australia, Antarctica,
Morocco.——Fig. 92,2a–b. *P. parvus (R. beDForD
& W. r. beDForD), Ajax Limestone, Botoman, Ajax
Mine, South Australia, Australia, holotype, SAM
P992; a, detail of intervallum, longitudinal view
(outer wall to left), ×10 (Debrenne, Zhuravlev, &
Kruse, 2002); b, external longitudinal view of outer
wall, ×6 (Debrenne, 1974c).
Family EREMITACYATHIDAE
Debrenne, 1992
[Eremitacyathidae Debrenne in Debrenne & ZhurAvlev, 1992b, p. 112]
[=Eremitacyathidae ZAmArreño & Debrenne, 1977, p. 55, nom. nud.]
Inner wall with canals. lower Cambrian
(Atd.2).
Eremitacyathus ZAmArreño & Debrenne, 1977,
p. 55 [*E. fissus; OD; holotype, ZAmArreño
& Debrenne, 1977, pl. 5a–b; Debrenne &
Zhu rAv le v, 1992b, pl. 3,4, MNHN M84016,
specimen Spe 10-1a, Paris]. Inner wall with
one canal-like opening per intersept, longitu-
dinall y cont inuous along entire cup, bounded
by longitudinal plates bearing denticulate rims
and lacking transverse partitions; pseudosepta
coarsely porous; synapticulae may be present.
lower Cambrian (Atd.2): Iberia.——Fig . 93a–b.
*E. fissus, Pedroche Formation, Atdabanian, Las
Ermitas, Cordoba, Andalusia, Spain, holotype,
MNHN M84016, specimen Spe 10-1a; a, trans-
verse section, ×2.5; b, longitudinal section, ×2.5
(Zamarreño & Debrenne, 1977).
Superfamily
SAKHACYATHOIDEA
Debrenne & Zhuravlev, 1990
[nom. transl. Debrenne & ZhurAvlev, 1992b, p. 112, ex Sakhacyathidae
Debrenne & ZhurAvlev, 1990, p. 302]
Outer wall pustular. lower Cambrian
(Tom.2–Atd.2).
Eremitacyathus
a
b
Fig. 93. Eremitacyathidae (p. 119).

120 Treatise Online, number XX
Family SAKHACYATHIDAE
Debrenne & Zhuravlev, 1990
[Sakhacyathidae Debrenne & ZhurAvlev, 1990, p. 302]
Inner wall with simple pores. l o wer
Cambrian (Tom.2–Atd.2).
Sakhacyathus Debrenne & ZhurAvlev, 1990,
p. 302 [*Paranacyathus subartus Zhu r Avl e vA ,
19 60b, p. 291; OD; holoty pe, ZhurAvlevA,
1960b, pl. 28,6, TsSGM 205/149, Novosibirsk]
[= Orbiparanocyathus be lY Ae vA , 1996, p. 109
(type, O. zolaensis, OD)]. Inner wall with one,
rarely two rows of simple pores per intersept;
pseudosepta finely porous. lower Cambrian
(Tom.2–Atd.2): Siberian Platform, Altay Sayan,
Tuva, Mongolia, Transbaikalia.——Fi g. 94a–b.
*S. subartus (Zhur Avl ev A), Pestrotsvet Forma-
tion, Atdabanian, Mukhatta River, Sakha
(Yakutia), Russia; a, holotype, TsSGM 205/149,
longitudinal section (outer wall to left), ×7
(Debrenne, Zhuravlev, & Kruse, 2002); b ,
transverse section, specimen PIN 4451/9, ×15
(Debrenne & Zhuravlev, 1992b; copyright
Publications Scientifiques du Muséum national
d’Histoire naturelle, Paris).
Superfamily
CHANKACYATHOIDEA
Yakovlev, 1959
[nom. transl. Debrenne & ZhurAvlev, 1992b, p. 112, ex Chankacyathidae
YAKovlev, 1959, p. 93]
Outer wall with canals. lower Cambrian
(Atd.4–Bot.3).
Family CHANKACYATHIDAE
Yakovlev, 1959
[Chankacyathidae YAKovlev, 1959, p. 93]
Inner wall with simple pores. l o wer
Cambrian (Bot.1–Bot.3).
Chankacyathus YAKov lev , 1959, p. 91, fig. 1 [*C.
strachovi; OD; nom. correct. oKu ne vA, 1969, p.
82, pro C. strachovii; holotype not designated,
collection not located]. Outer wall with horizontal
to upwardly projecting, straight canals, bearing
supplementary bracts externally (imparting overall
inverted V-shaped appearance to outer wall); inner
wall with one row of simple pores per intersept;
pseudosepta finely porous. lower Cambrian (Bot.1–
Bot.3): Far East, ?Australia.——Fig . 95,1a–b.
*C. strachovi, Dmitrievka Formation, Botoman,
Kar’ernaya, Far East, Russia, specimen PGU 202
133/52; a, transverse section of modular skeleton,
×7; b, longitudinal section, ×6.5 (Okuneva, 1969).
Family TCHOJACYATHIDAE
Debrenne & Zhuravlev, 1992
[Tchojacyathidae Debrenne & ZhurAvlev, 1992b, p. 113]
Inner wall with canals. lower Cambrian
(Atd.4).
Tchojacyathus roZAn ov, 1960b, p. 46 [*T. validus;
OD; holotype, roZAnov , 1960b, pl. 1,3 (non fig.
2); ZhurAvlevA, Ko nYushKov, & roZ Anov, 1964,
pl. 11,1, PIN 4297/11, Moscow]. Outer wall with
horizontal to upwardly projecting, S-shaped canals;
inner wall with one row of horizontal to upwardly
projecting, S-shaped canals per intersept; pseu-
dosepta coarsely porous. lower Cambrian (Atd.4):
Sakhacyathus
a
b
Fig. 94. Sakhacyathidae (p. 120).

Systematic Descriptions of the Archaeocyatha 121
Altay Sayan.——Fig . 95,2a–b. *T. validus, Uba
Formation, Atdabanian, Tyrga River, Altay Moun-
tains, Altay Sayan, Russia, holotype, PIN 4297/11;
a, longitudinal section (outer wall to left), ×6; b,
transverse section, ×6 (Rozanov, 1960b).
Suborder ANTHOMORPHINA
Okulitch, 1935
[nom. transl. Debrenne, 1991, p. 219, ex Anthomorphida oKuliTch,
1955a, p. 18, nom. correct. pro order Anthomorphina oKuliTch, 1935,
p. 90] [=subclass Anthocyatha oKuliTch, 1943, p. 46; =Araneocyathida
vologDin, 1961, p. 182]
Cup solitary or modular (pseudocolonies
formed by external budding); intervallum
with pseudosepta and membrane tabulae.
lower Cambrian (Bot.1).
Superfamily
ANTHOMORPHOIDEA
Okulitch, 1935
[nom. transl. Debrenne & ZhurAvlev, 1992b, p. 113, ex Anthomorphidae
oKuliTch, 1935, p. 97]
Outer wall simple, with pores of anthoid
type; microporous membranes of similar
structure to tabulae may be present. lower
Cambrian (Bot.1).
Fig. 95. Chankacyathidae and Tchojacyathidae (p. 120–121).
Tchojacyathus
2a
2b
Chankacyathus
1a
1b

122 Treatise Online, number XX
Family ANTHOMORPHIDAE
Okulitch, 1935
[Anthomorphidae oKuliTch, 1935, p. 97] [=Anthomorphinae oK u-
liTch, 1 935, nom. transl. Fonin, 1 985, p . 121, ex Anthomorphidae
oKuliTch, 1935, p. 97; =Araneocyathidae vologDin, 1956, p. 878;
=Serligocyathidae vologDin, 1959a, p. 670; =Rudicyathinae Fonin
in Zhur Avlev, ZhurAv levA, & Fonin, 1983, p. 26; =Vertocyathinae
Fonin, 1985, p. 110]
Inner wall with simple pores. l o wer
Cambrian (Bot.1).
Anthomorpha bor n e m A n n , 1884, p. 705 [*A .
margarita; M; lectotype, bornemAnn, 1886,
pl. 28,1a,4–6; Debrenne, 1964, pl. 45,1; SD
Debrenne, 1964, p. 233, GML 897a, Halle]. Inner
wall with one row of simple pores per intersept;
pseudosepta aporose even in early ontogenetic
stages; membrane tabulae may be present. lowe r
Cambrian (Bot.1): Tuva, ?Far East, Morocco, Iberia,
France, Sardinia.——Fig. 96,1a–b. *A. margarita,
Matoppa Formation, Botoman; a, Cuccuru Contu,
Sardinia, Italy, lectotype, GML 897a, transverse
section, ×4 (Debrenne, Zhuravlev, & Kruse, 2002);
b, Gonnesa, Sardinia, Italy, paralectotype, MNHN
M84133, specimen C GON 3-7, longitudinal
section, ×3 (Debrenne, 1964).
Tollicyathus chernYshevA, 1960, p. 77 [*T. ishensis;
OD; holotype, chernYshevA, 196 0, pl. 4,1,
ZSGGU 503/1, Novokuznetsk] [=Nellicyathus
Fonin in rePinA & others, 1964, p. 247 (type, N.
nelliae, OD); =Rudicyathus Fonin in Zhur Avlev,
ZhurAvlevA, & Fonin, 1983, p. 26 (type, R. tersus,
OD); =Vertocyathus Fonin, 1985, p. 110 (type, V.
reduncus, OD), for discussion, see Debrenne &
ZhurAvlev, 1992b, p. 67]. Inner wall with one
row of simple pores per intersept; pseudosepta
with pores restricted to outer wall area, but coarsely
porous in early ontogenetic stages; membrane
tabulae may be present. lower Cambrian (Bot.1):
Altay Sayan, Tuva, Mongolia.——Fig. 96,2a–b.
*T. ishensis, Verkhneynyrga Formation, Botoman,
Bol’shaya Isha River, Altay Mountains, Altay Sayan,
Russia, holotype, ZSGGU 503/1; a, longitudinal
section, ×5; b, transverse section, ×5 (Cherny-
sheva, 1960).
Family SHIVELIGOCYATHIDAE
Fonin, 1983
[nom. transl. Debrenne & ZhurAvlev, 1992b, p. 113, ex Shiveligocy-
athinae Fonin, 1983, p. 12]
Inner wall with canals. lower Cambrian
(Bot.1).
Shiveligocyathus missArZhevsKiY, 1961, p. 19 [*S.
vesiculoides; OD; holotype, missArZhevsKiY, 1961,
pl. 1,1, PIN 1914/75M/44, Moscow, not located]
[=Voznesenskicyathus roDionovA in Zhu rAvl evA
& others, 1967, p. 99 (type, V. florens, OD), for
discussion, see Debrenne & ZhurAvlev, 1992b, p.
130]. Inner wall with several rows of horizontal to
upwardly projecting, straight communicating canals
per intersept; pseudosepta finely porous; membrane
tabulae may be present. lower Cambrian (Bot.1):
Altay Sayan, Tuva, Mongolia.——Fig. 97 a–b.
*S. vesiculoides, Shangan Formation, Botoman,
Shivelig-Khem River, eastern Tannu-Ola Range,
Tuva, Russia, paratype, PIN 1914/75M/00; a,
longitudinal section, ×2 (Debrenne, Zhuravlev,
& Kruse, 2002); b, oblique transverse section, ×2
(Missarzhevskiy, 1961).
Suborder ARCHAEOCYATHINA
Okulitch, 1935
[nom. transl. ZhurAvle vA, 1960b, p. 271, ex Archaeocyathida Zhu rAv-
levA, 1955a, p. 17, nom. correct. pro order Archaeocyathina oKuliTch,
1935, p. 90] [=Archaeosyconina ZhurAvlevA, 1955a, p. 12, nom. transl.
ZhurAv levA, 1960b, p. 303, ex order Archaeosyconida Zhu rAvle vA,
1955a, p. 12, nom. correct. Debrenne, 1964, p. 117, pro Archaeosyconiina
ZhurAvlevA, 1960b, p. 303; =Dictyocyathina vologDin, 1956, p. 878,
nom. transl. Fonin in voronin & others, 1982, p. 83, ex Dictyocyathida
vologDin, 1956, p. 878; =Chouberticyathina Debrenne, 1970a, p. 25,
nom. transl. Fonin in voronin & others, 1982, p. 83, ex Chouberticy-
athida Debrenne, 1970a, p. 25]
Cup solitary or modular (latter by external
budding and/or longitudinal subdivision;
encrusting forms develop by addition of
new central cavities); intervallum with
taeniae, pseudosepta, or pseudotaenial or
dictyonal network; segmented tabulae may
be present. lower Cambrian (Tom.2–Toy.3),
middle Cambrian.
Superfamily
DICTYOCYATHOIDEA
Taylor, 1910
[nom. transl. WooD, evAns, & Zhu rAvlev, 1992, p. 492, ex Dictyocy-
athidae TAYlor, 1910, p. 111]
Outer wall simple, either rudimentary (of
marginal intervallar elements only) or basic
(of marginal intervallar elements with addi-
tional linking lintels); segmented tabulae may
be present. lower Cambrian (Tom.2–Toy.1),
middle Cambrian.
Family DICTYOCYATHIDAE
Taylor, 1910
[Dictyocyathidae TAYlor, 1910, p. 111] [=subfamily Dictyocyathinea
hernánDeZ-sAmPelAYo, 1933, p. 159; =Prismocyathidae Fonin, 1960, p.
725; =Paracoscinidae Debrenne, 1970a, p. 38, nom. nud.; =Paracoscinidae
Debrenne, 1974a, p. 252; =Chouberticyathidae Debrenne, 1974a, p. 192;
=Graphoscyphiidae Debrenne, 1974a, p. 204, nom. correct. Kruse, 1982,
p. 196, pro Graphoscyphidae Debrenne, 1974a, p. 204]
Inner wall with simple pores. l o wer
Cambrian (Tom.2–Toy.1), middle Cambrian.
Dictyocyathus bor ne m An n , 1891a, p. 500 [*D.
tenerrimus; M; lectotype, bornemAnn, 1891a, pl.
42,5; pl. 43,4–6; SD Debrenne, 1964, p. 200, not

Systematic Descriptions of the Archaeocyatha 123
located; =Coscinocyathus verticillus bornemAnn,
1886, p. 65; lectotype, bornemAnn, 1886, pl.
15,3g; Debrenne, 1964, pl. 34,5; SD Debrenne,
1964, p. 205, GML 899c, Halle] [=Prismocyathus
Fonin, 1960, p. 725 (type, P. praesignis, OD);
=Spongiosicyathus ZhurAvlevA in DAT se nK o &
others, 1968, p. 174 (type, Dictyocyathus translu-
cidus ZhurAvlevA, 1960b, p. 275, OD); =Prismo-
cyathellus Fonin, 1990, p. 152 (type, Prismocyathus
verisimilis Fonin, 1960, p. 726, OD; =Prismocy-
athus praesignis Fonin, 1960, p. 725)]. Outer wall
basic; inner wall with one row of simple pores
per intersept; dictyonal network. lower Cambrian
(Tom.2–Bot.1), middle Cambrian (Guzhangian):
Siberian Platform, Kolyma, Altay Sayan, Tuva,
Mongolia, Far East, Kazakhstan, Morocco, Iberia,
Sardinia, Germany, Tom.2–Bot.1; Antarctica,
Guzhangian.——Fig . 98,1a–b. *D. verticillus
(bornemAnn), Matoppa Formation, Botoman,
Cuccuru Contu, Sardinia, Italy; a, topotype,
Tollicyathus
Anthomorpha
2a
2b
1a
1b
Fig. 96. Anthomorphidae (p. 122).

124 Treatise Online, number XX
MNHN M84248, specimen CCC 9-1a, oblique
longitudinal section, ×6 (Debrenne, Zhuravlev, &
Kruse, 2002); b, lectotype, GML 899c, transverse
section, ×6 (Debrenne, 1964).
Cellicyathus Debrenne & ZhurAvlev, 1990, p. 300
[*Maturocyathus ornatus Fonin, 1985, p. 118; OD;
holotype, Fon in, 1985, pl. 22,2, PIN 1915/280,
Moscow]. Outer wall basic, tabular; inner wall
tabular, with one row of simple pores per intersept;
taeniae coarsely porous; synapticulae and simply
porous segmented tabulae may be present. lower
Cambrian (Bot.2–Toy.1): Siberian Platform, Altay
Sayan, Tuva.——Fig. 98,2a–b. *C. ornatus (Fonin),
Shangan Formation, Botoman, Ulug-Shangan River,
eastern Tannu-Ola Range, Tuva, Russia; a, holotype,
PIN 1915/280, transverse section, ×5; b, paratype,
PIN 1915/300, longitudinal section (outer wall to
right), ×3.5 (Fonin, 1985).
Chouberticyathus Debrenne, 1964, p. 208 [*C.
clatratus; OD; holotype, Debrenne, 1964, pl.
32,1–3, MNHN M80272, specimen Ki 140 P-6,
Paris]. Outer wall imperforate (possibly rudi-
mentary); inner wall with one row of simple
pores per intersept; taeniae coarsely porous. lower
Cambrian (Bot.1): southern China, Morocco,
Iberia, Sardinia.——Fig. 98,3a–b. *C. clatratus,
Issafen Formation, Botoman, Tizi Oumeslema,
Morocco, holotype, MNHN M80272, specimen
Ki 140 P-6; a, transverse view, ×6; b, longitu-
dinal view, ×6 (Debrenne & Zhuravlev, 1992b;
copyright Publications Scientifiques du Muséum
national d’Histoire naturelle, Paris).
Graphoscyphia Debrenne in ZhurAvlevA, 1974a, p.
164 [*Protopharetra graphica R. beDF orD & W. R.
beDForD, 1934, p. 4; OD; lectotype, R. beDForD
& W. r. beDForD, 1934, fig. 22; Debrenne, 1969a,
pl. 12,5; SD Debrenne, 1969a, p. 346, NHM
S4170, London]. Outer wall basic; inner wall with
one row of simple pores per intersept; pseudosepta
coarsely porous, linked by synapticulae. l ower
Cambrian (Atd.4–Bot.3): Altay Sayan, Australia,
Antarctica, Mexico.——Fig. 98,4a–b. *G. graphica
(R. beDFo rD & W. r. beDFor D), Ajax Limestone,
Botoman, Ajax Mine, South Australia, Australia,
lectotype, NHM S4170; a, internal longitudinal
view of inner wall, ×6; b, transverse view, ×6
(Debrenne, Zhuravlev, & Kruse, 2002).
Molybdocyathus Debrenne & gAngloFF i n
Debrenne, gAnDin, & gAngloFF, 1990, p. 92
[*M. juvenilis; OD; holotype, Debrenne, gAnDin,
& gAngloFF, 1990, pl. 2,13, USNM 443573,
specimen IR 23.7a’, Washington, D.C.]. Outer
wall rudimentary; inner wall with one row of
simple po re s per inters ept; dictyona l network.
lower Cambrian (Bot.1–Bot.2): Altay Sayan, Tuva,
Mo ngolia , Unit ed States. ——F i g . 99,1a–b.
*M. juvenilis, Valmy Formation, Botoman, Iron
Canyon, Nevada, United States; a, holotype,
USNM 443573, specimen IR 23.7a’, transverse and
and longitudinal sections of modular skeleton, ×8;
b, paratype, USNM 443568, specimen IR 14.2,
and longitudinal section of modular skeleton, ×8
(Debrenne, Gandin, & Gangloff, 1990).
Paracoscinus R. beDForD & W. R. beDForD, 1936,
p. 18 [*P. mirabile; OD; holotype, R. beDForD &
W. r. beDForD, 1936, fig. 85; Debrenne, 1974a,
fig. 37a–b, SAM P988-169, -170, -171, Adelaide].
Outer wall basic, tabular; inner wall with one row
of simple pores per intersept, each pore subdivided
by median longitudinal rod; pseudosepta finely
porous; segmented tabulae. lower Cambrian (Bot.3–
Toy.1): Altay Sayan, Australia.——Fig. 99,2a–d. *P.
mirabile, Ajax Limestone, Botoman, Ajax Mine,
South Australia, Australia; a, holotype, SAM P988-
169, -170, -171, transverse view, ×6 (Debrenne &
Zhuravlev, 1992b; copyright Publications Scienti-
fiques du Muséum national d’Histoire naturelle,
Paris); b, paratype, USNM PU86680, specimen
241A, external view of outer wall, ×6 (Debrenne,
Zhuravlev, & Kruse, 2002); c–d, holotype, SAM
P988-169, -170, -171; c, longitudinal view, ×4; d,
internal view of inner wall, ×7 (Debrenne, 1974a).
?Retilamina Debrenne & jAmes, 1981, p. 370
[*R. amourensis; OD; holotype, Debrenne &
jAmes, 1981, pl. 54,4, GSC 62128, specimen
169-5acT1, Ottawa]. Encrusting, domelike cup;
Fig. 97. Shiveligocyathidae (p. 122).
Shiveligocyathus
a
b

Systematic Descriptions of the Archaeocyatha 125
Fig. 98. Dictyocyathidae (p. 122–124).
Chouberticyathus
Graphoscyphia
Cellicyathus
Dictyocyathus
3a
3b
4a 4b
2a
2b
1a
1b

126 Treatise Online, number XX
Fig. 99. Dictyocyathidae (p. 124).
Molybdocyathus
Paracoscinus 2c
2a
2b
1a
1b
2d

Systematic Descriptions of the Archaeocyatha 127
upper wall (interpreted as outer) with pores
regularly arranged but not at each intertaenia;
pores commonly produced as chimneys; lower
(possibly inner) wall rudimentary; dictyonal or
more probably pseudotaenial network. [Atypical
cup shape does not provide certainty as to which
wall is outer and which inner, and nature of inter-
vallar elements and accepted inner wall remain
uncertain.] lower Cambrian (Bot.2): Canada,
United States, Mexico.——Fig. 100. *R. amou-
rensis, Forteau Formation, Botoman, Mount St.
Margaret, Newfoundland, Canada, holotype, GSC
62128, specimen 169-5acT1, oblique section, ×6
(Debrenne, Zhuravlev, & Kruse, 2002).
Family CLARUSCOSCINIDAE
Debrenne & Zhuravlev, 1992
[Claruscoscinidae Debrenne & ZhurAvlev, 1992b, p. 114] [=Clarus-
coscinidae Debrenne i n Debrenne, gAnDin, & roWlA nD, 1989, p.
167, nom. nud.]
Inner wall with bracts, fused bracts, or
pore tubes. lower Cambrian (Bot.1–Toy.1).
Claruscoscinus hAnDFielD, 1971, p. 74 [*Eucyathus
billingsi vologDin, 1940b, p. 48; OD; holotype
not designated, collection not located] [=Monstri-
cyathus vologDin, 1977, p. 60 (type, M. tubi-
formis, OD); =Arisacyathus KAshinA in osA DchAYA
& others, 1979, p. 166 (type, A. diligens, OD;
=Eucyathus billingsi vologDin, 1940b, p. 48);
=Maturocyathus Fonin, 1985, p. 114 (type, M.
makarovi, OD; =Eucyathus billingsi vologDin,
1940b, p. 48); =Costocyathus Fonin, 1985, p.
119 (type, C. mactus, OD), for discussion, see
Debrenne & Zh urAvl ev, 1992b, p. 123]. Outer
wall basic, tabular; inner wall with one row of pores
per intersept, bearing upwardly projecting, straight
to S-shaped pore tubes; pseudosepta finely porous;
segmented tabulae. lower Cambrian (Bot.1–Toy.1):
Altay Sayan, Tuva, Mongolia, Transbaikalia, Far
East, Canada, United States.——Fig. 101,1a–c. *C.
billingsi (volog Din ), Verkhnemonok Formation,
Botoman, Berezovaya River, Abakan River, western
Sayan, Altay Sayan, Russia; a, unlocated syntype,
longitudinal section, ×4 (Vologdin, 1940b); b,
transverse section, syntype, specimen PIN 4754/6,
×5 (Debrenne, Zhuravlev, & Kruse, 2002); c, unlo-
cated syntype, longitudinal section, ×4 (Vologdin,
1940b).
Fenestrocyathus hAnDFielD, 1971, p. 72 [*F.
complexus; OD; holotype, hAnDFielD, 1971, pl.
14,5; pl. 15,1, GSC 25388, Ottawa]. Outer wall
basic; inner wall with one row of pores per inter-
sept, bearing upwardly projecting, S-shaped bracts
or fused bracts; dictyonal network. lower Cambrian
(Bot.1–Bot.2): Altay Sayan, Mongolia, Canada,
United States.——Fig. 101,2. *F. complexus, Sekwi
Formation, Botoman, Mackenzie Mountains,
Northwest Territories, Canada, holotype, GSC
25388, transverse section of modular skeleton, ×5
(Debrenne, Zhuravlev, & Kruse, 2002).
Landercyathus Debrenne & gAngloFF in Debrenne,
gAnDin, & gAngloFF, 1990, p. 91 [*L. lewandowskii;
OD; holotype, Debrenne, gAnDin, & gAngloFF,
1990, pl. 1,13, USNM 443571, specimen IR 23a,
Washington, D.C.]. Outer wall simple; inner wall
with one row of horizontal to upwardly projecting,
straight to waved canals per intersept; canals may
penetrate intervallum forming astrorhizae; dictyonal
network. lower Cambrian (Bot.2): United States.——
Fig. 101,3. *L. lewandowskii, Valmy Formation,
Botoman, Iron Canyon, Nevada, United States,
holotype, USNM 443571, specimen IR 23a, oblique
longitudinal section, ×4 (Debrenne, Gandin, &
Gangloff, 1990).
Stevocyathus Debrenne in Debrenne, gAnDin, &
roWlAnD, 1989, p. 166 [*S. elictus; OD; holotype,
Retilamina
Fig. 100. Dictyocyathidae (p. 124–127).

128 Treatise Online, number XX
Landercyathus
Stevocyathus Fenestrocyathus
4a
4b
2
3
Claruscoscinus
1c
1a
1b
Fig. 101. Claruscoscinidae (p. 127–129).

Systematic Descriptions of the Archaeocyatha 129
Debrenne, gAnDin, & roWlAnD , 1989, pl. 12,1,
MNHN M83100, specimen CR2-8, Paris]. Outer
wall basic; inner wall with one row of pores per
intersept, bearing upwardly projecting, S-shaped
bracts or fused bracts; taeniae coarsely porous,
linked by synapticulae; simple segmented tabulae
may be present. lower Cambrian (Bot.2): United
States, Mexico.——Fig. 101,4a–b. *S. elictus, Puerto
Blanco Formation, Botoman, Caborca, Sonora,
Mexico; a, paratype, MNHN M83107, specimen
CR2*1-8, transverse section, ×6 (Debrenne,
Zhuravlev, & Kruse, 2002); b, holotype, MNHN
M83100, specimen CR2-8, oblique longitudinal
section, ×6 (Debrenne, Gandin, & Rowland, 1989).
Family PYCNOIDOCOSCINIDAE
Debrenne, 1974
[Pycnoidocoscinidae Debrenne, 1974a, p. 256] [=Pycnoidocoscinidae
Debrenne, 1970a, p. 40, nom. nud.]
Inner wall compound. lower Cambrian
(Bot.3).
Pycnoidocoscinus R. beDForD & W. R. beDForD, 1936,
p. 19 [*P. pycnoideum; OD; lectotype, R. beDForD &
W. r. beDForD, 1936, fig. 87; SD Debrenne, 1970a,
p. 40, SAM P990-175, -176, -177, Adelaide]. Outer
wall basic; inner wall compound consisting of wall
carcass and additional microporous sheath formed by
tabulae; pseudosepta finely porous; segmented tabulae.
lower Cambrian (Bot.3): Australia, ?Canada.——Fig.
102a–b. *P. pycnoideum, Ajax Limestone, Botoman,
Ajax Mine, South Australia, Australia, lectotype,
SAM P990-175, -176, -177; a, transverse view (outer
wall at top), ×4; b, tangential view of inner wall, ×6
(Debrenne, Zhuravlev, & Kruse, 2002).
Superfamily
ARCHAEOCYATHOIDEA
Hinde, 1889
[nom. correct. ZhurAvle v in vor ono vA & oth ers, 1987, p. 32, p ro
Archaeocyathacea simon, 1939, p. 6, nom. transl. ex Archaeocyathidae
TAYlor, 1910, p. 105, nom. correct. pro family Archaeocyathinae hinDe,
1889, p. 141] [=Flindersicyathoidea R. beDForD & J. beDForD, 1939, p.
78, nom. correct. Debrenne & Kruse, 1986, p. 268, pro Flindersicyathacea
grAvesTocK, 1984, p. 115, nom. transl. ex Flindersicyathidae R. beDForD
& J. beDFor D, 1939, p. 78; =Vadimocyathacea KAshinA in os ADchAYA
& others, 1979, p. 160]
Outer wall concentrically porous. lower
Cambrian (Atd.1–Toy.3).
Family ARCHAEOPHARETRIDAE
R. Bedford & W. R. Bedford, 1936
[Archaeopharetridae R. beDForD & W. R. beDForD, 1936, p. 17] [=Dic-
tyocoscinidae R. beDForD & W. R. beDForD, 1936, p. 14, for discussion,
see ZhurAvlev & grAvesTocK, 1994, p. 34; =Protopharetridae vologDin,
1957a, p. 182; =Flindersicoscinidae Debrenne, 1974a, p. 246; =Salanycy-
athidae Fonin in voronin & others, 1982, p. 95; =Hawkercyathidae
grAvesTocK, 1984, p. 115]
Inner wall with simple pores. l o wer
Cambrian (Atd.1–Bot.3).
Archaeopharetra R. beDF or D & W. R. beD F or D,
1936, p. 17 [*A . typica; OD; holotype, R.
beD F o r D & W. R. beD F or D , 1936, fig. 75;
Zhur Avle vA, 1963b, fig. 67a; Debrenne, 1974a,
fig. 3b; SD hill, 1965, p. 115, SAM P969,
Adelaide; =Dictyocyathus irregularis TAY l o r,
1910, p. 145; lectotype, TAY l o r , 1910, pl.
12, photo 66; SD Debrenne, ZhurAvlev, &
Krus e, 2002, p. 1665, SAM T1590, Adelaide]
[=Dictyocoscinus R. beDForD & W. R. beDForD,
19 36, p. 14 (ty pe, D. bel tana , OD; =Di cty o -
cyathus irregularis TAYl o r, 1910, p. 145, for
discussion, see ZhurAvlev & grAvesTocK,
1994, p. 34); =Tubocyathus vologDin, 1937b,
p. 473 (type, T. smolianinovae, M); =Tubicy-
athus vologDin, 1940a, p. 114, nom. null.;
=Tubulocyathus vologDin, 1956, p. 880, nom.
null.; =Flindersicoscinus Debrenne, 1970a, p.
34 (type, Flindersicyathus tabulatus R. beDF or D
& J. beD F or D , 1937, p. 29, OD); =Salanycy-
athus Fonin in voronin & others, 1982, p.
95 (type, S. marginatus, OD); =Hawkercy-
athus gr Av e s T o c K , 1984, p. 115 (type, H.
insculptus, OD), for discussion, see Debrenne
& ZhurAvlev, 1992b, p. 120]. Inner wall
with one row of simple pores per intersept;
pseudotaeniae coarsely porous; concentrically
porous segmented tabulae may be present.
lower Cambrian (Atd.1–Bot.3): Altay Sayan,
Tuva, Mongolia, Far East, Australia, Antarc-
tica, South Africa (allochthonous), southern
China, ?Iberia.——Fi g. 103,1a–b. *A. irregu-
laris (TAYl or ), Ajax Limestone, Botoman, Ajax
Mine, South Australia, Australia; a, lectotype,
Pycnoidocoscinus
a
b
Fig. 102. Pycnoidocoscinidae (p. 129).

130 Treatise Online, number XX
Markocyathus
2a
2b
Archaeopharetra
1a
1b
Fig. 103. Archaeopharetridae (p. 129–133).

Systematic Descriptions of the Archaeocyatha 131
Spirocyathella
2c
2a
2b
Dictyosycon
1
Fig. 104. Archaeopharetridae (p. 132–133).

132 Treatise Online, number XX
Protopharetra
c
a
b
Fig. 105. Archaeopharetridae (p. 133).
SAM T1590, oblique longitudinal view, ×5
(Debrenne, Zhuravlev, & Kruse, 2002); b, (=A.
typica R. beDForD & W. R. beDForD), holotype,
SAM P969, longitudinal view, ×8 (Debrenne,
1974a).
Dictyosycon Zhu r Av le vA , 1960b, p. 307, nom.
transl. Debrenne & Zh ur Avl ev, 1992b, p. 124,
ex Sphinctocyathus (Dictyosycon) ZhurAvlevA,
1960b, p. 307 [*Sphinctocyathus (Dictyosycon)
gravis; OD; holotype, Zhu rAv l evA , 1960b, pl.
31,7, TsSGM 205/169, Novosibirsk]. Inner
wall with one row of simple pores per intersept;
dictyonal network; simple or concentrically
porous segmented tabulae may be present. lower
Cambrian (Atd.1–Atd.4): Siberian Platform,
Altay Sayan, Tuva, Iberia.——Fig. 104,1. *D.
gravis, Pestrotsvet Formation, Atdabanian,
Oy-Muran, Lena River, Sakha (Yakutia), Russia,
holotype, TsSGM 205/169, oblique longitudinal
section, ×4 (Zhuravleva, 1960b).
Markocyathus Debrenne in Debrenne, gAnDin, &
roWl An D, 1989, p. 165 [*M. clementensis; OD;
holotype, Debrenne, gAnDin, & roW l A n D ,
1989, pl. 11,1–2, MNHN M83096, specimen

Systematic Descriptions of the Archaeocyatha 133
CL-1e, Paris]. Inner wall with several rows of
simple pores per intersept; taeniae coarsely
porous; concentrically porous segmented
tabulae. lower Cambrian (Bot.2): Canada,
Mexico.——Fig. 103,2a–b. *M. clementensis,
Puerto Blanco Formation, Botoman, Caborca,
Sonora, Mexico, holotype, MNHN M83096,
specimen CL-1e; a, detail of inner wall in
tangential section, ×12 (Debrenne, Zhuravlev,
& Kruse, 2002); b, transverse and longitudinal
sections of modular skeleton, ×3 (Debrenne,
Gandin, & Rowland, 1989).
Protopharetra bornemAnn, 1884, p. 705 (borne-
mAnn, 1883, p. 274, nom. nud.) [*P. polymorpha
bornemAnn, 1886, p. 46; SD simon, 1939,
p. 34; lectotype, bornemAnn, 1886, pl. 5,
fig. 4 bottom; SD simon, 1939, p. 35, not
located; topotypes, MNHN M84120, speci-
mens CGR3/3, GLA3.3, GLC10.II.1b, Paris]
[=Volvacyathus Debrenne, 1960, p. 118 (type,
V. proteus, OD), for discussion, see Debrenne
& Zhu rAv lev , 1992b, p. 128]. Inner wall with
one row of simple pores per intersept; taeniae
coarsely porous, linked by rare synapticulae.
lower Cambrian (Atd.1–Bot.3): Altay Sayan,
Tuva, Far East, Tajikistan, Canada, United
States, southern China, Morocco, Iberia, France,
Sardinia, Germany.——Fig. 105a–c. *P. poly-
morpha, Matoppa Formation, Botoman, Canal
Grande, Sardinia, Italy; a, topotype, MNHN
M84120, transverse section of modular skeleton,
×3 (Debrenne, Zhuravlev, & Kruse, 2002); b,
lectotype, transverse section, ×3 (Bornemann,
1886); c, topotype, MNHN M84120, detail of
outer wall in tangential section, ×7 (Debrenne,
Zhuravlev, & Kruse, 2002).
Spirocyathella vologDin, 1939, p. 227 [*S. kyzlar-
tauensis; OD; holotype not designated, collec-
tion not located] [=Aruntacyathus Kr u se in
WAlTer, 1980, chart, nom. nud.; =Amaded-
cyathus Kru s e in WAlTer, 1980, chart, nom.
nud.; =Aruntacyathus Kru se in Krus e & Wes T,
1980, p. 172 (type, A. toddi, OD); =Spi roc y-
athellus Fonin in voronin & others, 1982, p.
98, lapsus calami pro Spirocyathella vologDin,
1939, p. 227, for discussion, see Debrenne &
ZhurAvlev, 1992b, p. 131]. Inner wall with
several rows of simple pores per intersept; pseu-
dotaenial network coarsely porous; concentri-
cally porous segmented tabulae. lower Cambrian
(Atd.4–Bot.2): Altay Sayan, Urals, Canada,
United States, Mexico, Australia, Antarctica,
South Africa (allochthonous), France.——Fig .
104,2a–c. *S. kyzlartauensis, Terekla Forma-
tion, Botoman, Mt. Kizlar-Tau, western flank
of southern Urals, Russia; a, topotype, PIN
4451/26, longitudinal section, ×8 (Debrenne
& Zhuravlev, 1992b; copyright Publications
Scientifiques du Muséum national d’Histoire
naturelle, Paris); b–c, unlocated syntype, 4-M,
thin section 4; b, transverse section, ×6; c,
longitudinal section, ×6 (Vologdin, 1939).
Family ARCHAEOCYATHIDAE
Hinde, 1889
[nom. correct. TAYlor, 1910, p. 105, pro family Archaeocyathinae hinDe,
1889, p. 141] [=Spirocyathidae TAYlor, 1910, p. 112; =Archaeocyathinae
hernánDeZ-sAmPelAYo, 1933, p. 158, nom. correct. Fonin, 1985, p. 69,
pro Archaeocyathinea hernánDeZ-sAmPelAYo, 1933, p. 158; =Sigmofungi-
idae R. beDForD & W. R. beDForD, 1936, p. 16, nom. correct. Debrenne,
1970a, p. 42, pro Sigmofungidae R. beDForD & W. R. beDForD, 1936, p.
16; =Flindersicyathidae R. beDForD & J. beDForD, 1939, p. 78; =Flinder-
sicyathinae R. beDForD & J . beDForD, 1939, p. 78, nom. transl. Fonin,
1985, p . 93, ex Fli ndersicyathidae R. beDForD & J. beDForD, 1939, p.
78; =Pycnoidocyathidae oKuliTch, 1950b, p. 394; ?=Protocyclocyathidae
vologDin, 1956, p. 878; ?=Protocyclocyathellidae vologDin, 1956, p.
878, lapsus calami Debrenne & jAmes, 1981, p. 366, pro Protocyclocy-
athidae vologDin, 1956, p. 878; =Syringsellidae KrAsnoPeevA, 1961, p.
248; =Archaeofungiidae vologDin, 1962c, p. 90, nom. correct. hill, 1965,
p. 58, pro Archaeofungidae vologDin, 1962c, p. 90; =Vadimocyathidae
KAshinA in osADchAYA & others, 1979, p. 161; =Claruscyathinae Fonin
in ZhurAvlevA & Fonin, 1983, p. 49]
Inner wall with bracts, fused bracts, or
pore tubes. lower Cambrian (?Atd.4, Bot.2–
Toy.3).
Archaeocyathus billings, 1861, p. 3 [*A. atlanticus
billings, 1861, p. 5; SD WA lcoTT , 1886, p. 75;
holotype, billings, 1861, fig. 5; hinDe, 1889,
pl. 5,8–10; oKuliTch, 1943, pl. 5,1–2, GSC
369, Ottawa] [Original spelling was Archeocy-
athus billings; subsequent authors have used the
diphthong] [=Spirocyathus hinDe, 1889, p. 136
(type, Archeocyathus atlanticus billings, 1861,
p. 5, M); =Retecyathus vologDin, 1932, p. 20,
nom. nud.; =Claruscyathus vologD in, 1932, p. 25
(type, C. cumfundus, M); =Eucyathus vologDin,
1937b, p. 466, nom. nud.; =Flindersicyathus R.
beDForD & J. beDForD, 1937, p. 28, nom. nud.;
=Flindersicyathus R. beDF o rD & J. beDF o rD ,
1939, p. 78 (type, F. decipiens, OD);=Eucyathus
vologDin in simon, 1939, p. 29 (type, Clarus-
cyathus cumfundus vologDin, 1932, p. 25, OD);
=Retecyathus vologDin i n simon, 1939, p. 36
(type, R. laqueus vologDin, 1932, p. 20, OD,
=Claruscyathus cumfundus vologDin, 1932, p.
25); =Syringsella KrAsnoPee vA, 1961, p. 248 (type,
S. ynyrgensis, OD); =Batenevia KrAsnoPeevA,
1961, p. 249 (type, B. pellisi, OD); =Sanxiacyathus
YuAn & ZhAng, 1977, p. 8 (type, S. hubeiensis,
OD); =Bijacyathus Kr As n oP e evA , 1978, p. 81
(type, Archaeocyathus regularis KrAsnoPeevA in
ZhurAvlevA, KrAsnoPeevA, & chernYshevA,
1960, p. 135, M, =Retecyathus kusmini vologDin,
1932, p. 21); =Retecyathus (Pararetecyathus) YuAn
& ZhAng, 1978, p. 139 (type, R. (P.) curvatus,
OD); =Vadimocyathus KAsh inA in osA Dch AYA &
others, 1979, p. 161 (type, V. chikinevae, OD), for
discussion, see Debrenne & ZhurAvlev, 1992b, p.
120]. Inner wall with one row of pores per inter-
sept, bearing upwardly projecting, straight pore
tubes; pseudotaenial network coarsely porous;
concentrically porous segmented tabulae. l ower
Cambrian (?Atd.4, Bot.2–Toy.3): Siberian Plat-
form, Altay Sayan, Tuva, Mongolia, Transbaikalia,
Far East, Uzbekistan, Canada, United States,
Mexico, Australia, Antarctica, southern China,

134 Treatise Online, number XX
Iberia, Sardinia.——Fig . 106,1a–b. *A. atlan-
ticus, Forteau Formation, Botoman, Anse au
Loup, Labrador, Canada, holotype, GSC 369; a,
longitudinal section, ×2.5 (Okulitch, 1943); b,
transverse section, ×3 (Debrenne, Zhuravlev, &
Kruse, 2002).
Arrythmocricus Debrenne & jAmes, 1981, p. 366 [*A.
kobluki; OD; holotype, Debrenne & jAmes, 1981,
pl. 53,3–4, GSC 62123, Ottawa]. Inner wall with
one row of pores per intersept, bearing upwardly
projecting, S-shaped bracts or fused bracts; pseu-
dotaenial network coarsely porous. lower Cambrian
(Bot.1–Bot.3): Canada, United States, Mexico.——
Fig. 106,2a–b. *A. kobluki, Forteau Formation,
Botoman, Fox Cove, Labrador, Canada, holotype,
GSC 62123; a, longitudinal section of modular
skeleton, ×5; b, longitudinal section (inner wall at
top), ×5 (Debrenne & James, 1981).
Archaeocyathus
Arrythmocricus
2a
2b
1a
1b
Fig. 106. Archaeocyathidae (p. 133–134).

Systematic Descriptions of the Archaeocyatha 135
Pycnoidocyathus
c
a
b
Fig. 107. Archaeocyathidae (p. 135–136).
Pycnoidocyathus TAYlor , 1910, p. 131 [*P. synap-
ticulosus; SD R. beDF or D & J. beDFor D, 1939,
p. 78; lectotype, TAYlo r, 1910, pl. 12, photo 69;
Debrenne, 1974a, fig. 13a–b; SD Debrenne,
1970a, p. 40, SAM T1587A,B,C, Adelaide]
[=Archaeofungia TAYlor , 1910, p. 131 (type, A.
ajax, M); =Batenevicyathus YAroshevich, 1962,
p. 117, 122 (type, B. zhuravlevae, OD), for
discussion, see Debrenne & ZhurAvlev, 1992b,
p. 129]. Inner wall with one row of pores per

136 Treatise Online, number XX
intersept, bearing upwardly projecting, straight
pore tubes; taeniae coarsely porous, linked at base
by synapticulae; during ontogeny, taeniae become
progressively less porous, more planar and
without synapticulae. lower Cambrian (Bot.2–
Toy.3): Altay Sayan, Tuva, Mongolia, Far East,
Australia, Antarctica, southern China, Iberia,
Sardinia, Greenland, Canada, United States,
Mexico.——Fig . 107a–c. *P. synapticulosus,
Ajax Limestone, Botoman, Ajax Mine, South
Australia, Australia, lectotype, SAM T1587A–C;
a, transverse view, ×1; b, longitudinal view, ×1;
c, detail of outer wall in tangential view, ×4
(Debrenne, Zhuravlev, & Kruse, 2002).
Sigmofungia R. beDForD & W. R. beDForD, 1936,
p. 16 [*S. flindersi; M; lectotype, R. beDForD
& W. r. beDForD, 1936, fig. 82; hill, 1965,
pl. 6,1–2; Debrenne, 1974a, fig. 30a–b; SD
hill, 1965, p. 89, SAM P963-115, -116,
Adelaide] [=Palmericyathellus Debrenne, 1970a,
p. 37 (type, Sigmofungia tabularis R. beD Fo rD
& j. be DF or D, 1937, p. 29, OD, =Sigmofungia
flindersi R. beDFo rD & W. r. beDF or D, 1936, p.
16), for discussion, see Debrenne & Zhu rAvle v,
1992b, p. 130; =Palmericyathus Debrenne in
ZhurAvle vA, 1974b, p. 15, nom. null., non hAnD-
FielD, 1971, p. 44 (type, Ethmophyllum lineatus
greggs, 1959, p. 66, OD), archaeocyath]. Inner
wall with one row of pores per intersept, bearing
upwardly projecting, S-shaped pore tubes; taeniae
finely porous, linked by synapticulae; concentri-
cally porous, segmented tabulae. lower Cambrian
(Bot.2–Bot.3): Australia, Antarctica, Mexico.——
Fig . 108a–c. * S. flindersi, Ajax Limestone,
Botoman, Ajax Mine, South Australia, Australia,
lectotype, SAM P963-115, -116; a, transverse
and longitudinal views, ×3 (Hill, 1965); b, detail
of inner wall in oblique view, ×11 (Debrenne,
1974a); c, detail of outer wall in tangential view,
×10 (Debrenne, Zhuravlev, & Kruse, 2002).
Family ARCHAEOSYCONIDAE
Zhuravleva, 1954
[Archaeosyconidae ZhurAvlevA, 1954, p. 30]
Inner wall compound. lower Cambrian
(Bot.1–Bot.3).
Archaeosycon TAYlor, 1910, p. 111 [*Archaeocyathus
billingsi WAl co TT , 1886, p. 74; M; holotype,
WAl co TT , 1886, pl. 3,3a–c; oKuliTch, 1943,
pl. 14,2–3, USNM 15302, Washington, D.C.]
[=Pustulacyathellus Debrenne & gAngloFF in
voronovA & others, 1987, p. 42 (type, P. copulatus,
OD), for discussion, see Debrenne & Zhur Avlev,
1992b, p. 121]. Outer wall tabular; inner wall
compound, comprising wall carcass and tabulae;
taeniae coarsely porous; concentrically porous
segmented tabulae. lower Cambrian (Bot.1–Bot.3):
Canada, United States.——Fig. 109a–b. *A. bill-
ingsi (WAl coT T), Forteau Formation, Botoman,
Anse au Loup, Labrador, Canada, holotype, USNM
15302; a, longitudinal section, ×3; b, transverse
section, ×3 (Okulitch, 1943).
Superfamily METACYATHOIDEA
R. Bedford & W. R. Bedford, 1934
[nom. correct. Debrenne & Kruse, 1986, p. 266, pro Metacyathacea Fonin,
1983, p. 11, nom. transl. ex Metacyathidae R. beDForD & W. R. beDForD,
1934, p. 5] [=Spirillicyathacea grAvesTocK, 1984, p. 111]
Outer wall compound. lower Cambrian
(Tom.2–Bot.3).
Family COPLEICYATHIDAE
R. Bedford & J. Bedford, 1937
[Copleicyathidae R. beDFor D & J. beDForD, 1937, p. 29] [=Tabulacy-
athellidae Fonin in voronin & others, 1982, p. 86; =Spirillicyathidae
grAvesTocK, 1984, p. 111].
Inner wall with simple pores. l o wer
Cambrian (Tom.2–Bot.3).
Copleicyathus R. beDForD & J. beDForD, 1937, p.
29 [*C. confertus; OD; holotype, R. beDForD &
J. beDFo rD, 1937, fig. 116; hill, 1965, pl. 10,4;
Debrenne, 1974a, fig. 27, USNM PU86741-783,
Washington, D.C.]. Outer wall compound with
completely subdivided pores; inner wall with several
rows of simple pores per intersept; pseudotaenial
network coarsely porous. lower Cambrian (Atd.3–
Atd.4): Australia.——Fig. 110a–b. *C. confertus,
Ajax Limestone, Atdabanian, Paint Mine, South
Australia, Australia, holotype, USNM PU86741-
783; a, transverse view, ×5; b, longitudinal view,
×5 (Hill, 1965).
Agastrocyathus Debrenne, 1964, p. 209 [*Proto-
pharetra gregaria Debrenne, 1961, p. 21; OD;
holotype, Debrenne, 1961, pl. 2,5–6, MNHN
M80138, HD71, Paris]. Outer wall compound
with incipient subdivision of intervallar cells; inner
wall with one row of simple pores per intersept;
taeniae coarsely porous, linked by synapticulae.
lower Cambrian (Atd.2–Atd.4): Morocco,
Iberia.——Fig. 111a–c. *A. gregarius (Debrenne),
Amouslek Formation, Atdabanian, Jbel Taïssa,
Morocco, holotype, MNHN M80138, HD71; a,
longitudinal section of modular skeleton, ×3; b,
transverse section of modular skeleton, ×4; c, detail
of outer wall in tangential section, ×10 (Debrenne,
Zhuravlev, & Kruse, 2002).
Gabrielsocyathus Debrenne, 1964, p. 248 [*Meta-
coscinus gabrielsensis oKuliTch, 1955b, p. 61;
OD; holotype, oKuliTch, 1955b, pl. 1,1,2,5,
GSC 12357, Ottawa]. Outer wall compound with
completely subdivided pores; inner wall with several
rows of simple pores per intersept; taeniae finely
porous, linked by synapticulae; simple segmented
tabulae. lower Cambrian (Bot.2): Canada, United
States.——Fig . 112,1a–c. *G. gabrielsensis
(oKuliTch), Atan Group, Botoman, McDame
Lake, British Columbia, Canada, holotype, GSC
12357; a, transverse section, ×2.5; b, longitudinal
section, ×2.5; c, transverse section (outer wall at

Systematic Descriptions of the Archaeocyatha 137
Sigmofungia
c
a
b
Fig. 108. Archaeocyathidae (p. 136).

138 Treatise Online, number XX
bottom), ×2.5 (Debrenne, Zhuravlev, & Kruse,
2002).
Metacyathellus Debrenne & Zhur Avle v, 1990, p.
302 [*Metaldetes? caribouensis hAnDFielD, 1971,
p. 64; OD; holotype, hAnDFielD, 1971, pl. 11,2,
GSC 25367, Ottawa]. Outer wall compound with
completely subdivided pores; inner wall with one
to two rows of simple pores per intersept; taeniae
coarsely porous; compound segmented tabulae.
lower Cambrian (Atd.4–Bot.3); Australia, Antarctica,
Canada, United States.——Fig. 112,2a–c. *M. cari-
bouensis (hAnDFielD), Sekwi Formation, Botoman; a,
Caribou Pass, Northwest Territories, Canada, holo-
type, GSC 25367, transverse section, ×4 (Hand-
field, 1971; reproduced with the permission of the
Minister of Public Works and Government Services
Canada, 2006 and courtesy of Natural Resources
Canada, Geological Survey of Canada); b, Mackenzie
Mountains, Northwest Territories, Canada, spec-
imen GSC 90187, detail of outer wall in tangential
section, ×10 (Voronova & others, 1987); c, Caribou
Pass, Northwest Territories, Canada, holotype, GSC
25367, longitudinal section, ×6 (Handfield, 1971;
reproduced with the permission of the Minister of
Public Works and Government Services Canada,
2006 and courtesy of Natural Resources Canada,
Geological Survey of Canada).
Spinosocyathus ZhurAvlevA, 1960b, p. 276 [*S. maslen-
nikovae; OD; holotype, Zhu rAvl evA, 1960b, pl.
25,1b, TsSGM 205/134, Novosibirsk]. Outer wall
compound with incipient pore subdivision; inner
wall with one row of simple pores per intersept;
pseudotaenial network coarsely porous; compound
segmented tabulae. lower Cambrian (Tom.2–Atd.2):
Siberian Platform, Mongolia, Iberia.——Fig .
113,1a–b. *S. maslennikovae, Pestrotsvet Formation,
Tommotian, Churan, Lena River, Sakha (Yakutia),
Russia; a, holotype, TsSGM 205/134, transverse
section (outer wall at bottom), ×8 (Zh ura v-
leva, 1960b); b, oblique longitudinal section of
modular skeleton, specimen TsSGM 144-32/4, ×3
(Debrenne, Zhuravlev, & Kruse, 2002).
Spirillicyathus R. beDFo rD & J. beD Fo rD , 1937,
p. 30 [*S. tenuis; OD; holotype, R. beDForD &
J. beDForD, 1937, fig. 118; Debrenne, 1974a,
fig. 10, USNM PU493967, specimen 358, Wash-
ington, D.C.] [=Spiralicyathus R. beDForD & J.
beDForD, 1937, fig. 118 caption, nom. null.]. Outer
wall compound with completely subdivided pores;
inner wall with one to two rows of simple pores per
intersept; pseudotaenial network coarsely porous.
lower Cambrian (Atd.4–Bot.1): Australia, southern
China.——Fig. 113,2a–c. *S. tenuis, Ajax Lime-
stone, Atdabanian, Paint Mine, South Australia,
Australia; a–b, holotype, USNM PU493967, spec-
imen 358; a, transverse view, ×9; b, longitudinal
view, ×9 (Debrenne, 1974a); c, Wilkawillina Lime-
stone, Atdabanian, Wilkawillina Gorge, South
Australia, Australia, specimen SAM P21741, tangen-
tial section of outer wall, ×10 (Gravestock, 1984).
Tabulacyathellus missArZhevsKiY in rePinA & others,
1964, p. 249 [*T. bidzhaensis; OD; holotype,
rePinA & others, 1964, pl. 7,4–6, PIN 4297/22,
Moscow]. Outer wall compound with completely
subdivided pores; inner wall tabular with several
rows of simple pores per intersept; pseudotaenial
network coarsely porous; compound segmented
tabulae. lower Cambrian (Atd.2): Altay Sayan, Tuva,
Mongolia.——Fig. 114a–c. *T. bidzhaensis, Usa
Formation, Atdabanian, Sukhie Solontsy Valley,
Batenev Range, Kuznetsk Alatau, Russia, holotype,
PIN 4297/22; a, tangential section of outer wall,
×5; b, longitudinal section (outer wall to left), ×5;
c, transverse section, ×5 (Repina & others, 1964).
Family JUGALICYATHIDAE
Gravestock, 1984
[Jugalicyathidae grAvesTocK, 1984, p. 114]
Inner wall with bracts, fused bracts
or pore tubes. lower Cambrian (Atd.4–
Bot.2).
Fig. 109. Archaeosyconidae (p. 136).
Archaeosycon
a
b

Systematic Descriptions of the Archaeocyatha 139
Jugalicyathus grAvesTocK, 1984, p. 114 [*J. tardus;
OD; holotype, grAvesTocK, 1984, fig. 56H–I, SAM
P21747, Adelaide]. Outer wall compound with
incipient subdivision of intervallar cells; inner wall
with one row of pores per intersept, bearing upwardly
projecting, straight pore tubes; pseudosepta finely
porous. lower Cambrian (Atd.4): Australia.——Fig.
115,1a–b. *J. tardus; a, Wilkawillina Limestone,
Atdabanian, Wilkawillina Gorge, South Australia,
Australia, paratype, SAM P21749, oblique transverse
section, ×8 (Debrenne, Zhuravlev, & Kruse, 2002);
b, Ajax Limestone, Atdabanian, Mount Scott Range,
South Australia, Australia, holotype, SAM P21747,
longitudinal section, ×1 (Gravestock, 1984).
Alaskacoscinus Debrenne, gAngloFF, & Zhu rAvlev
in Debrenne & ZhurAvlev, 1990, p. 300 [*A.
tatondukensis; OD; holotype, De b r e n n e &
ZhurAvlev, 1990, pl. 1,5, UAM UA2534, 2535,
Fairbanks]. Outer wall tabular, compound with
completely subdivided pores; inner wall tabular
with one row of pores per intersept, bearing
upwardly projecting, S-shaped pore tubes; pseu-
dosepta finely porous; segmented tabulae with
subdivided pores. lower Cambrian (Bot.2): United
States.——Fig. 115,2a–b. *A. tatondukensis, Adams
Argillite, Botoman, Tatonduk River, Alaska, United
States; a, holotype, UAM UA2534, longitudinal
section (outer wall to right), ×4; b, paratype, UAM
UA2536, longitudinal section, ×5 (Debrenne &
Zhuravlev, 1990).
Family METACYATHIDAE
R. Bedford & W. R. Bedford, 1934
[Metacyathidae R. beDForD & W. R. beDForD, 1934, p. 5] [=Metacos-
cinidae R. beDForD & W. R. beDForD, 1936, p. 18; =Cambrocyathidae
oKuliTch, 1937a, p. 251; =Cambrocyathinae oKuliTch, 1937a, p. 251,
nom. transl. Debrenne, 1964, p. 218, ex Cambrocyathidae oKuliTch,
1937a, p. 251; =Metaldetinae Debrenne, 1964, p. 218; =Metafungiidae
Debrenne, 1974a, p. 216]
Inner wall compound. lower Cambrian
(Atd.4–Bot.3).
Metaldetes TAYlor , 1910, p. 151 [*M. cylindricus;
M; holotype, TAYlor, 1910, pl. 15, photo 86–88,
fig. 11, 37, 38; Debrenne, 1974a, fig. 21a–b,
M, SAM T1592A, Adelaide] [=Metafungia R.
beDForD & W. R. beDForD, 1934, p. 5 (type,
M. reticulata, M); =Metacyathus R. beDF or D &
W. R. be DFor D, 1934, p. 5 (type, M. taylori, M,
=Archaeocyathus dissepimentalis TAY lo r, 1910,
Copleicyathus
a
b
Fig. 110. Copleicyathidae (p. 136).

140 Treatise Online, number XX
p. 128); =Metacoscinus R. beDForD & W. R.
beDForD, 1934, p. 6 (type, M. reteseptatum, M,
=Archaeocyathus retesepta TAYlor, 1910, p. 120);
=Cambrocyathus oKuliTch, 1937a, p. 251 (type,
Archaeocyathus profundus billings, 1861, p. 4,
OD); =Metethmophyllum oKuliTch, 1943, p. 78
(type, Ethmophyllum meeki WA lc oT T, 1889, p.
34, OD); =Bedfordcyathus vologDin, 1957a, p.
182 (type, Metacyathus irregularis R. beDForD &
W. R. be DFo rD, 1934, p. 6, M, =Archaeocyathus
dissepimentalis TAYlor, 1910, p. 128); =Praefungia
Debrenne in ZhurAvlevA, 1974b, p. 42, nom.
correct. Debrenne, 1974a, p. 227, pro Pruefungia,
lapsus calami (type, Metaldetes superbus R. beDForD
& W. R. beDForD, 1936, p. 18, OD), for discus-
sion, see Debrenne & ZhurAvlev, 1992b, p. 127].
Outer wall compound with completely subdivided
pores; inner wall compound with several rows of
completely subdivided pores per intersept; taeniae
coarsely porous, linked by synapticulae in early
ontogenetic stages but rarely so in mature cups;
compound segmented tabulae. lower Cambrian
(Atd.4–Bot.3): Far East, Australia, Antarctica,
Canada, United States, Mexico.——Fig. 116a–e.
*M. cylindricus, Wilkawillina Limestone, Botoman,
Wilson, South Australia, Australia, holotype, SAM
T1592A; a, transverse section, ×4; b, detail of inner
wall in transverse section, ×8; c, detail of outer
wall in transverse section, ×8; d, detail of taenia
in longitudinal section, ×8; e, longitudinal section
Agastrocyathus
c
a
b
Fig. 111. Copleicyathidae (p. 136).

Systematic Descriptions of the Archaeocyatha 141
Fig. 112. Copleicyathidae (p. 136–138).
Gabrielsocyathus
Metacyathellus
2c
2b
1c
1a
1b
2a
I had to
rotate view
c to make
the figures
fit--if this
is not okay,
please mark
crop lines
on either
views c or a
so I can fit
them side
by side.
Thanks!

142 Treatise Online, number XX
of modular skeleton, ×4 (Debrenne, Zhuravlev, &
Kruse, 2002).
Changicyathus Debrenne & Zhu rAvl ev , 1990, p.
301 [*Cambrocyathellus tenuicaulus ZhAng &
YuA n, 1985, p. 523; OD; holotype, ZhAng &
YuAn , 1985, pl. 2,6; Debrenne & ZhurAvlev,
1990, pl. 1,7, NIGP 82277, specimen 17f(10-
14), Nanjing]. Outer wall compound with
completely subdivided pores; inner wall
compound with incipient pore subdivision;
taeniae coarsely porous; compound segmented
tabulae. lower Cambrian (Bot.1): ?Tajikistan,
southern China.——Fi g. 117. *C. tenuicaulus
(ZhAng & Yu A n ), Xiannudong Formation,
Spinosocyathus
Spirillicyathus 2c
2a
2b
1a
1b
Fig. 113. Copleicyathidae (p. 138).

Systematic Descriptions of the Archaeocyatha 143
Botoman, Nanzhen, Sichuan, China, holotype,
NIGP 82277, specimen 17f(10-14), oblique
longitudinal section of modular skeleton, ×6
(Debrenne & Zhuravlev, 1990).
Superfamily
NAIMARKCYATHOIDEA
Wrona & Zhuravlev, 1996
[Naimarkcyathoidea WronA & ZhurAvlev, 1996, p. 28]
Outer wall pustular. lower Cambrian (Bot.3).
Family NAIMARKCYATHIDAE
Wrona & Zhuravlev, 1996
[Naimarkcyathidae WronA & ZhurAvlev, 1996, p. 29]
Inner wall with bracts, fused bracts or
pore tubes. lower Cambrian (Bot.3).
Naimarkcyathus WronA & Zh urAvle v, 1996, p. 29
[*N. elenae; OD; holotype, WronA & ZhurAvl ev,
19 96, pl. 7,2, ZPAL Ac. I/M1 0DI, Wars aw].
Inner wall with one row of pores per intersept,
bearing upwardly projecting, straight pore tubes;
pseudotaenial network coarsely porous. l owe r
Cambrian (Bot.3): Antarctica.——Fig. 118a–b.
*N. elenae, Polonez Cove Formation (allochtho-
nous), Botoman, Mazurek Point, King George
Island, southern Shetland Islands, Antarctica; a,
holotype, ZPAL Ac.I/M10DI, transverse section,
×5; b, paratype, ZPAL Ac.I/M10CI, longitudinal
section, ×5 (Wrona & Zhuravlev, 1996).
Superfamily
WARRIOOTACYATHOIDEA
Debrenne & Zhuravlev, 1992
[Warriootacyathoidea Debrenne & ZhurAvlev, 1992b, p. 115]
Outer wall with canals. lower Cambrian
(Atd.3–Atd.4).
Family WARRIOOTACYATHIDAE
Debrenne & Zhuravlev, 1992
[Warriootacyathidae Debrenne & ZhurAvlev, 1992b, p. 115]
Inner wall with bracts, fused bracts or
pore tubes. lower Cambrian (Atd.3–Atd.4).
Warriootacyathus grAvesTocK, 1984, p. 126
[*W. wilkawillinensis; OD; holotype, gr Av e-
sT o cK , 1984, fig. 62A,D–F, SAM P21806-1,
Adelaide]. Outer wall with horizontal to
upwardly projecting, straight canals; inner wall
with one row of pores per intersept, bearing
upwardly projecting, straight to waved pore
tubes; pseudosepta coarsely porous. l o w e r
Cambrian (Atd.3–Atd.4): Australia.——Fig .
119a–c. *W. wilkawillinensis, Wilkawillina
Limestone, Atdabanian, Wilkawillina Gorge,
South Australia, Australia, holotype, SAM
P21806-1; a, tangential section of outer wall,
×7; b, tangential section of inner wall, ×3; c,
longitudinal section of septum (outer wall to
right), ×3 (Gravestock, 1984).
Fig. 114. Copleicyathidae (p. 138).
Tabulacyathellus
c
a
b

144 Treatise Online, number XX
Alaskacoscinus
2a
2b
Jugalicyathus
1a 1b
Fig. 115. Jugalicyathidae (p. 139).

Systematic Descriptions of the Archaeocyatha 145
Fig. 116. Metacyathidae (p. 139–142).
Metaldetes
c
d
e
a
b

146 Treatise Online, number XX
Superfamily
BELTANACYATHOIDEA
Debrenne, 1974
[nom. correct. Debrenne & ZhurAvlev, 1992b, p. 115, pro Beltanacyatha-
cea grAvesTocK, 1984, p. 123, nom. transl. ex Beltanacyathidae Debrenne,
1974a, p. 243] [=Beltanacyathidae Debrenne, 1970a, p. 30, nom. nud.]
Outer wall with subdivided canals. lower
Cambrian (Atd.3–Bot.3).
Family MAIANDROCYATHIDAE
Debrenne, 1974
[Maiandrocyathidae Debrenne, 1974a, p. 235]
Inner wall with simple pores. l o wer
Cambrian (Bot.3).
Maiandrocyathus Debrenne in Zhur AvlevA, 1974a,
p. 209 [*Metacoscinus insigne R. beDFo rD & W. R.
beDForD, 1936, p. 18; OD; holotype, R. beD ForD
& W. R. beDForD, 1936, fig. 84; Debrenne, 1974a,
fig. 28, M, SAM P986-167, -168, Adelaide]. Inner
wall with one to two rows of simple pores per
intersept; taeniae coarsely porous. lower Cambrian
(Bot.3): Australia.——Fig. 120,1a–b. *M. insigne
(R. beDFo rD & W. r. beDFor D), Ajax Limestone,
Botoman, Ajax Mine, South Australia, Australia,
holotype, SAM P986-167, -168; a, tangential view
of outer wall, ×3; b, longitudinal view of septum
and exocyathoid buttress (outer wall to right), ×3
(Debrenne, Zhuravlev, & Kruse, 2002).
Ataxiocyathus Debrenne in Zhu rAv l evA , 1974a,
p. 5 2 [*Paranacyathus grandis R . beDForD
& J. beDForD, 1937, p. 34; OD; holotype,
R. be D F o r D & j. beD F o r D , 1937, fig. 140;
Debrenne, 19 7 4 c, pl . 20 , 3 – 4 , M, US NM
PU86821, specimen 311, Washington, D.C.].
Inner wall with one row of simple pores per
intersept; pseudosepta finely porous. l owe r
Cambrian (Bot.3): Australia.——Fig. 120,2a–c.
*A. grandis (R. beDForD & j. beDForD) ,
Ajax Limestone, Botoman, Ajax Mine,
South Australia, Australia, holotype, USNM
PU86821, specimen 311; a, transverse view,
×5 (Debrenne, Zhuravlev, & Kruse, 2002); b,
longitudinal view of septum (outer wall to left),
×6 (M. Debrenne, new); c , tangential view of
outer wall, ×6 (Debrenne, 1974c).
Family BELTANACYATHIDAE
Debrenne, 1974
[Beltanacyathidae Debrenne, 1974a, p. 243] [=Beltanacyathidae De-
brenne, 1970a, p. 30, nom. nud.]
Inner wall with bracts, fused bracts,
or pore tubes. lower Cambrian (Atd.3–
Atd.4).
Beltanacyathus R. beDForD & J. beDForD, 1936,
p. 23 [*B. ionicus; OD; lectotype, R. beD Fo rD
& j. beD ForD , 1936, fig. 95–96; hill, 1965, pl.
6,3; SD hill, 1965, p. 89, USNM PU86716-
271, Washington, D.C.; =Archaeocyathus wirri-
alpensis TAYlor, 1910, p. 124; holotype, TAY lor,
1910, pl. 8, photo 43–44; Debrenne, 1974a, fig.
33b; M, SAM T1581A-E, Adelaide] [=Fridaycy-
athus grAve sT oc K, 1984, p. 125 (type, F. bise-
rialis, OD); =Bayleicyathus grAvesTocK, 1984,
p. 131 (type, B. bowmani, OD), for discussion,
see Debrenne & Zh u rAv l ev , 1992b, p. 122].
Inner wall with one row of pores per intersept,
bearing upwardly projecting, straight pore tubes;
pseudosepta finely porous; segmented tabulae.
Changicyathus
Fig. 117. Metacyathidae (p. 142–143).

Systematic Descriptions of the Archaeocyatha 147
lower Cambrian (Atd.3–Atd.4): Australia.——Fig.
121a–d. *B. wirrialpensis (TAYlor); a–b, holotype,
Wilkawillina Limestone, Atdabanian, Wirrealpa
Mine, South Australia, Australia, SAM T1581A-E;
a, transverse section (outer wall at top), ×2; b,
longitudinal section (outer wall to left), ×2.5
(Taylor, 1910); c–d, [=B. ionicus], lectotype,
Ajax Limestone, Atdabanian, Paint Mine, South
Australia, Australia, USNM PU86716-271; c ,
transverse view, ×2; d, longitudinal view, ×2
(Hill, 1965).
Superfamily
TABELLAECYATHOIDEA
Fonin, 1963
[nom. transl. Debrenne & ZhurAvlev, 1992b, p. 116, ex Tabellaecyathidae
Fonin, 1963, p. 15] [=Taeniaecyathellacea KonYushKov, 1972, p. 141]
Outer wall tabellar. lower Cambrian
(Bot.2–Bot.3).
Family TABELLAECYATHIDAE
Fonin, 1963
[Tabellaecyathidae Fonin, 1963, p. 15] [=Taeniaecyathellidae
Kon Yu s hK ov , 1972, p. 142; =Karakolocyathidae Kon Y us hK ov ,
1972, p. 142]
Inner wall with bracts, fused bracts or
pore tubes. lower Cambrian (Bot.2–Bot.3).
Taeniaecyathellus ZhurAvlevA, 1960a, p. 45 [*T.
semenovi; OD; holotype, Zh ur Av le vA , 1960a,
fig. 1i–k, TsSGM 273/7, Novosibirsk] [=Tabel -
laecyathus Fonin, 1963, p. 15 (type, T. totus, OD);
=Cambronanus Fonin, 1963, p. 19 (type, C. multi-
cavitatus, OD); =Karakolocyathus KonY us hKov ,
1972, p. 142 (type, K. loculatus, OD; =Tabellae-
cyathus totus Fonin, 1963, p. 16), for discussion,
see Debrenne & Zh u rAv l ev , 1992b, p. 131].
Inner wall with several rows of pores per intersept,
bearing upwardly projecting, straight pore tubes;
dictyonal network. lower Cambrian (Bot.2–Bot.3):
Altay Sayan.——Fig. 122a–b. *T. semenovi, Verkh-
nemonok Formation, Botoman, Malyy Karakol
River, western Sayan, Altay Sayan, Russia, holotype,
TsSGM 273/7; a, oblique longitudinal section, ×5;
b, detail of outer wall in tangential section, ×21
(Debrenne, Zhuravlev, & Kruse, 2002).
Suborder DICTYOFAVINA
Debrenne, 1991
[Dictyofavina Debrenne, 1991, p. 219]
Skeleton solitary or modular, latter
as branching or massive pseudocolonies
(both by intercalicular budding); inter-
vallum with calicles. lower Cambrian
(Atd.1–Bot.2).
Naimarkcyathus
a
b
Fig. 118. Naimarkcyathidae (p. 143).
Superfamily
USLONCYATHOIDEA
Fonin, 1966
[nom. transl. Debrenne, ZhurAvlev, & Kruse, 2002, p. 1679, ex Uslon-
cyathidae Fonin in vologDin & Fonin, 1966, p. 187] [=Dictyofavoidea
Debrenne & ZhurAvlev, 1992a, p. 596]
Outer wall simple. lower Cambrian
(Atd.1–Bot.2).

148 Treatise Online, number XX
Family USLONCYATHIDAE
Fonin, 1966
[Usloncyathidae Fonin in vologDin & Fonin, 1966, p. 187] [=Dictyo-
favidae Debrenne & ZhurAvlev, 1992a, p. 596]
Inner wall with simple pores. l o wer
Cambrian (Atd.1–Bot.2).
Usloncyathus Fonin in vologDin & Fonin, 1966,
p. 188 [*U. miculus; OD; holotype, vologDin
& Fonin, 1966, fig. 1a, PIN 2486/143, Moscow]
[=Falsocyathus Fonin in vologDin & Fonin, 1966,
p. 189 (type, F. vastulus, OD; =U. miculus Fonin in
vologDin & Fonin, 1966, p. 188); =Nostrocyathus
Fonin in vologDin & Fonin, 1966, p. 189 (type,
N. aculeatus, OD; =U. miculus Fonin in vologDin
& Fonin, 1966, p. 188); =Cavocyathus Fonin in
vologDin & Fonin, 1966, p. 189 (type, C. pusilus;
OD; =U. miculus Fonin in vologDin & Fonin,
1966, p. 188), for discussion, see Debrenne &
Zhur Avlev , 1992b, p. 132; =Dictyofavus gr Ave-
sTocK, 1984, p. 98 (type, D. obtusus, OD)]. Outer
and inner walls rudimentary; calicles hexagonal
in cross section with several pore rows per facet.
lower Cambrian (Atd.1–Atd.4): Altay Sayan, Tuva,
Mongolia, Transbaikalia, Far East, Australia.——
Fig . 123,1. *U. miculus, Bystraya Formation,
Atdabanian, Uslon Valley, Transbaikalia, Russia,
holotype, PIN 2486/143, longitudinal section, ×8
(Vologdin & Fonin, 1966).
Kechikacyathus Debrenne & ZhurAvlev, 1992a, p.
598 [*K. natlaensis; OD; holotype, Debrenne &
ZhurAvlev, 1992a, pl. 1,3, GSC 90166, Ottawa].
Outer wall basic; inner wall rudimentary; calicles
hexagonal in cross section with one pore row per
facet. lower Cambrian (Bot.1–Bot.2): Canada.——
Fig. 123,2a–b. *K. natlaensis, Sekwi Formation,
Botoman; a, Kechika River, British Columbia,
Canada, paratype, GSC 103939, GAM-78-G,
detail of outer wall in tangential section, ×10; b,
Natla, Mackenzie Mountains, Northwest Territo-
ries, Canada, holotype, GSC 90166, longitudinal
section, ×5 (Debrenne & A. Zhuravlev, 1992a).
Zunyicyathus Debrenne, Kru se , & ZhAng, 1991,
p. 286 [*Agastrocyathus grandis YuA n & ZhAng,
1980, p. 387; OD; nom. correct. Debrenne, Krus e
& ZhAng, 1991, p. 286, pro Agastrocyathus grandus
YuAn & ZhAng, 1980, p. 387; holotype, YuAn &
ZhAng, 1980, pl. 1,3, NIGP 56292, Nanjing].
Outer and inner walls rudimentary; calicles tetrag-
onal in cross section with one pore row per facet.
lower Cambrian (Bot.1–Bot.2): Tajikistan, southern
China, United States.——Fig . 124. *Z. grandis
(YuAn & ZhAng), Jindingshan (Chintingshan)
Formation, Botoman, Jindingshan, Guizhou,
China, specimen MNHN 85103, longitudinal
section of modular skeleton, ×5 (Debrenne, Kruse,
& Zhang, 1991).
Superfamily
KERIOCYATHOIDEA
Debrenne & Gangloff, 1992
[Keriocyathoidea Debrenne & gAngloFF in Debrenne & ZhurAvlev,
1992a, p. 598]
Outer wall concentrically porous. lower
Cambrian (Bot.1–Bot.2).
Family KERIOCYATHIDAE
Debrenne & Gangloff, 1992
[Keriocyathidae Debrenne & gAngloFF in Debrenne & ZhurAvlev ,
1992a, p. 598] [=Keriocyathidae Debrenne & gAngloFF in Debrenne,
gAnDin, & gAngloFF, 1990, p. 93, nom. nud.]
Inner wall with simple pores. l o wer
Cambrian (Bot.1–Bot.2).
Keriocyathus Debrenne & gAngloFF in Debrenne,
gAnDin, & gAngloFF, 1990, p. 93 [*K. arachnaius;
OD; holotype, Debrenne, gAnDin, & gAngloFF,
1990, pl. 1,9, USNM 443557, specimen IR24.10,
Warriootacyathus
c
a
b
Fig. 119. Warriootacyathidae (p. 143).

Systematic Descriptions of the Archaeocyatha 149
Washington, D.C.]. Inner wall basic; calicles tetrag-
onal in cross section with one pore row per facet.
lower Cambrian (Bot.1–Bot.2): Altay Sayan, Far
East, United States.——Fig . 125a–b. *K. arach-
naius, Valmy Formation, Botoman, Iron Canyon,
Nevada, United States; a, holotype, USNM 443557,
specimen IR24.10, transverse section, ×7.5; b, para-
type, USNM 443572, longitudinal section, ×7.5
(Debrenne, Gandin, & Gangloff, 1990).
Superfamily
GATAGACYATHOIDEA
Debrenne & Zhuravlev, 1992
[Gatagacyathoidea Debrenne & ZhurAvlev, 1992a, p. 598]
Outer wall compound. lower Cambrian
(Bot.2).
Fig. 120. Maiandrocyathidae (p. 146).
Maiandrocyathus
2c Ataxiocyathus
2a
2b
1a
1b

150 Treatise Online, number XX
Beltanacyathus
c
d
a
b
Fig. 121. Beltanacyathidae (p. 146–147).

Systematic Descriptions of the Archaeocyatha 151
Taeniaecyathellus
a
b
Fig. 122. Tabellaecyathidae (p. 147).
Family GATAGACYATHIDAE
Debrenne & Zhuravlev, 1992
[Gatagacyathidae Debrenne & ZhurAvlev, 1992a, p. 598]
Inner wall with simple pores. l o wer
Cambrian (Bot.2).
Gatagacyathus Debrenne & Zhur Avlev , 1992a, p.
598 [*G. mansyi; OD; holotype, Debrenne &
ZhurAvlev, 1992a, pl. 1,4, GSC 103942, specimen
GAM76.8G.XI.3L, Ottawa]. Outer wall compound
with incipient pore subdivision; inner wall rudi-
mentary; calicles hexagonal in cross section with
one pore row per facet. lower Cambrian (Bot.2):
Canada, United States.——Fig. 126. *G. mansyi,
Rosella Formation, Botoman, Kechika River,
British Columbia, Canada, holotype, GSC 103942,
specimen GAM76.8G.XI.3L, longitudinal section,
×3.5 (Debrenne, Zhuravlev, & Kruse, 2002).
Suborder SYRINGOCNEMINA
Okulitch, 1935
[nom. correct. Debrenne, ZhurAvlev, & Kruse, herein, pro Syringicnemi-
dina KrAsnoP eevA, 1980, p. 159, nom. transl. ex order Syringocnemina
oKuliTch, 1935, p. 90] [=Syringocyathina Debrenne, 1991, p. 219]
Skeleton solitary or modular, latter as
branching pseudocolonies (by longitudinal
fission); intervallum with syringes. lower
Cambrian (Atd.4–Bot.3).
Superfamily
AULISCOCYATHOIDEA
Debrenne & Zhuravlev, 1992
[Auliscocyathoidea Debrenne & ZhurAvlev, 1992b, p. 117]
Outer wall simple. lower Cambrian
(Atd.4–Bot.3).
Family AULISCOCYATHIDAE
Debrenne & Zhuravlev, 1992
[Auliscocyathidae Debrenne & ZhurAvlev, 1992b, p. 117]
Inner wall with simple pores. l o wer
Cambrian (Atd.4–Bot.3).
Auliscocyathus Debrenne in ZhurAvlevA, 1974a, p.
53 [*Spirocyathus multifidus R. beDForD & W. R.
beDForD, 1936, p. 14; OD; lectotype, R. beDForD
& W. R. beDForD, 1936, fig. 65; Debrenne, 1974a,
fig. 8a; SD Debrenne, 1974a, p. 199, SAM P950-
81, Adelaide]. Outer and inner walls rudimentary;
syringes tetragonal in cross section with one pore
row per facet. lower Cambrian (Atd.4–Bot.3): Tuva,
Australia, Antarctica.——Fig. 127a–c. *A. multifidus
(R. beDFo rD & W. r. beDFo rD), Ajax Limestone,
Botoman, Ajax Mine, South Australia, Australia,
lectotype, SAM P950-81; a, longitudinal view, ×5
(Debrenne, 1974a); b, oblique longitudinal view,
×5; c, detail of syringes in longitudinal intervallar
view, ×8 (Debrenne, Zhuravlev, & Kruse, 2002).
Superfamily
SYRINGOCNEMOIDEA
Taylor, 1910
[nom. correct. Debrenne, ZhurAvlev, & Kruse , herein, pro Syrin -
gocnemidoidea Debrenne & ZhurAvlev, 1992b, p. 117, nom. transl. e x
Syringocnemidae TAYlor, 1910, p. 113]
Outer wall concentrically porous. lower
Cambrian (Bot.1–Bot.3).
Family TUVACNEMIDAE
Debrenne & Zhuravlev, 1990
[nom. correct. Debrenne, ZhurAvlev, & Kruse, herein, pro Tuvacnemidi -
dae Debrenne & ZhurAvlev, 1990, p. 300]
Inner wall with simple pores. l o wer
Cambrian (Bot.1–Bot.3).

152 Treatise Online, number XX
Tuvacnema Debrenne & Zhur Av l e v , 1990, p.
301 [*Syringocnema tannuolensis roDionovA in
ZhurAvlevA & others, 1967, p. 106; OD; holotype,
ZhurAvlevA & others, 1967, pl. 58,4, VSEGEI
9594, St. Petersburg, not located]. Inner wall with
several rows of pores per syrinx; syringes hexagonal
in cross section with several pore rows per facet.
lower Cambrian (Bot.1–Bot.3): Tuva.——Fig. 128.
*T. tannuolensis (roDionovA), Shangan Formation,
Botoman, Shivelig-Khem River, eastern Tannu-Ola
Range, Tuva, Russia, holotype, VSEGEI 9594,
transverse section, ×7 (Zhuravleva & others, 1967).
Family SYRINGOCNEMIDAE
Taylor, 1910
[Syringocnemidae TAYlor, 1910, p. 113] [=Syringocnematidae vologDin,
1928, p. 31; =Syringocnemitidae Ting, 19 37, p. 370 ; =Syringo cnemi-
didae Debrenn e, 1964, p. 117; =Pseudosyringocnemididae De brenne,
1975, p. 355]
Inner wall with bracts, fused bracts, or
pore tubes. lower Cambrian (Bot.1–Bot.3).
Syringocnema TAYlor, 1910, p. 153 [*S. favus; M;
holotype, TAYlo r, 1910, pl. 14, photos 78–79,
M, SAM T1597A,B,E, Adelaide]. Inner wall with
one row of pores per syrinx, bearing upwardly
projecting, S-shaped pore tubes; syringes hexagonal
in cross section with several pore rows per facet.
lower Cambrian (Bot.3): Australia, Antarctica.——
Fig. 129a–d. *S. favus, Ajax Limestone, Botoman,
Ajax Mine, South Australia, Australia; a–c, holo-
type, SAM T1597A,B,E; a, transverse view, ×3;
b, oblique longitudinal view, ×3.5; c, detail of
syringes in longitudinal view (outer wall to left),
×5; d, paratype, SAM T1558, detail of syringes in
oblique transverse view, ×5 (Debrenne, Zhuravlev,
& Kruse, 2002).
Pseudosyringocnema hAnDF ie lD, 1971, p. 76 [*P.
uniporum; OD; holotype, hAnDFielD, 1971,
pl. 15,3, GSC 25392, Ottawa]. Inner wall with
one row of pores per syrinx, bearing upwardly
projecting, S-shaped pore tubes; syringes hexag-
onal in cross section with one pore row per
transverse facet and several pore rows per lateral
Fig. 123. Usloncyathidae (p. 148).
Kechikacyathus
Usloncyathus
2b
1
2a

Systematic Descriptions of the Archaeocyatha 153
facet. lower Cambrian (Bot.2–Bot.3): Altay
Sayan, Antarctica, Canada, United States.——
Fig . 130,1a–b. *P. uniporum, unnamed Sekwi
Formation equivalent (map unit 5 of hAnDFielD,
1971), Botoman, Coal River, Yukon Territory,
Northwest Territories, Canada, holotype, GSC
25392; a, longitudinal section, ×4 (Debrenne,
Zhuravlev, & Kruse, 2002); b, oblique longitu-
dinal section, ×4 (Handfield, 1971; reproduced
with the permission of the Minister of Public
Works and Government Services Canada, 2006
an d courte sy of Natural Resourc es Ca nada,
Geological Survey of Canada).
Syringothalamus Debrenne, gAngloFF, & ZhurAvlev
in Debrenne & Zhu rAv le v, 1990, p. 301 [*S.
crispus; OD; holotype, Debrenne & Zhu rAvle v,
1990, pl. 1,1, UCMP D6610, Berkeley]. Inner wall
with one row of pores per syrinx, bearing upwardly
projecting, S-shaped fused bracts; syringes hexag-
onal in cross section with one pore row per facet.
lower Cambrian (Bot.1): United States.——Fig.
130,2a–c. *S. crispus, Poleta Formation, Botoman,
Lida, Palmetto Mountains, Nevada, United States;
a, holotype, UCMP D6610, detail of outer wall in
tangential section, ×11 (Debrenne, Zhuravlev, &
Kruse, 2002); b, paratype, UCMP D6620, trans-
Zunyicyathus
Fig. 124. Usloncyathidae (p. 148).

154 Treatise Online, number XX
(Bot.1–Bot.2): Canada, United States.——Fig .
131a–c. *W. colvillensis (greggs); a, Sekwi Forma-
tion, Botoman, Mackenzie Mountains, Northwest
Territories, Canada, specimen GSC 90169, trans-
verse section, ×7 (Voronova & others, 1987); b,
Maitlen Formation, Botoman, Colville, Washington,
United States, holotype, GSC 14317, transverse
section, ×7.5 (Greggs, 1959); c, Sekwi Formation,
Botoman, Mackenzie Mountains, Northwest Territo-
ries, Canada, GSC 90170, oblique transverse section,
×7.5 (Voronova & others, 1987).
Superfamily
KRUSEICNEMOIDEA
Debrenne & Zhuravlev, 1990
[nom. correct. Debrenne, Zh urAvl ev, & Kruse , herein, pro Kruseic-
nemidoidea Debrenne & ZhurAvlev , 1992b, p. 117, nom. transl. ex
Kruseicnemididae Debrenne & ZhurAvlev, 1990, p. 301]
Outer wall pustular. lower Cambrian (Bot.3).
Family KRUSEICNEMIDAE
Debrenne & Zhuravlev, 1990
[nom. correct. Debrenne, ZhurAvlev, & Kruse, herein, pro Kruseicnemi-
didae Debrenne & ZhurAvlev, 1990, p. 301]
Inner wall with bracts, fused bracts or
pore tubes. lower Cambrian (Bot.3).
Kruseicnema Debrenne, grAvesTocK, & ZhurAvlev
in Debrenne & ZhurAvlev , 1990, p. 301 [*Syrin-
gocnema gracilis gorDon, 1920, p. 699; OD;
holotype, gorDon, 1920, pl. 4,43,46, NHM
S10412, London]. Outer wall pustules bearing
supplementary multiperforate tumuli; inner wall
with one row of pores per syrinx, bearing upwardly
projecting, S-shaped pore tubes; syringes hexagonal
in cross section with several pore rows per facet.
lower Cambrian (Bot.3): Australia, Antarctica,
South Africa (allochthonous), Falkland Islands
(allochthonous).——Fig. 132,1a– b. *K. gracilis
(gorDon), allochthonous, Botoman, Weddell Sea,
Antarctica, holotype, NHM S10412; a, oblique
Keriocyathus
b
Fig. 125. Keriocyathidae (p. 148–149).
Fig. 126. Gatagacyathidae (p. 151).
Gatagacyathus
a
verse section, ×5; c, holotype, UCMP D6610,
oblique longitudinal section, ×5 (Debrenne &
Zhuravlev, 1990).
Williamicyathus ZhurAvlev in voronovA & others,
1987, p. 34 [*Syringocnema colvillensis greggs,
1959, p. 72; OD; holotype, greggs, 1959, pl. 13,6,
GSC 14317, Ottawa]. Inner wall with one row
of pores per syrinx, bearing upwardly projecting,
planar, fused bracts; syringes hexagonal in cross
section with one pore row per transverse facet and
several pore rows per lateral facet. lower Cambrian

Systematic Descriptions of the Archaeocyatha 155
longitudinal section, ×9; b, transverse section, ×9
(Gordon, 1920).
Superfamily
FRAGILICYATHOIDEA
Belyaeva, 1975
[nom. transl. Debrenne & ZhurAvlev, 1992b, p. 117, ex Fragilicyathidae
belYAevA in belYAevA & others, 1975, p. 117]
Outer wall with canals. lower Cambrian
(Bot.1).
Family FRAGILICYATHIDAE
Belyaeva, 1975
[Fragilicyathidae belYAevA in belYAevA & others, 1975, p. 117]
Inner wall with bracts, fused bracts, or
pore tubes. lower Cambrian (Bot.1).
Fragilicyathus belYAe vA, 1969, p. 98 [*F. zhurav-
levae; OD; holotype, be lYAe vA, 1969, pl. 37,7,
DVGU 6M/212/15-3, Khabarovsk]. Outer wall
with horizontal to upwardly projecting, straight
canals; inner wall with one row of pores per syrinx,
bearing upwardly projecting, S-shaped pore tubes;
syringes hexagonal in cross section with several
pore rows per facet. lower Cambrian (Bot.1): Far
East.——Fig . 132,2. *F. zhuravlevae, Ust’to ka
unit, Botoman, Gerbikan River, Dzhagdy Range,
Far East, Russia, holotype, DVGU 6M/212/15-3,
oblique longitudinal section, ×5.5 (Debrenne,
Zhuravlev, & Kruse, 2002).
Order
KAZACHSTANICYATHIDA
Konyushkov, 1967
[Kazachstanicyathida KonYushKov, 1967, p. 105]
Cup multichambered, solitary or
modular, with massive modular types by
individualization of modules around new
central cavities; development of thalamid
type, with stromatoporoid growth pattern;
chambers of subspherical to laterally
c
a
b
Auliscocyathus
Fig. 127. Auliscocyathidae (p. 151).

156 Treatise Online, number XX
elongate shape, with pillars. lower Cambrian
(Bot.1–Bot.3).
Suborder
KAZACHSTANICYATHINA
Konyushkov, 1967
[nom. transl. Debrenne & ZhurAvlev, 1992b, p. 118, ex Kazachstanicy-
athida KonYushKov, 1967, p. 105] [=Kazakhstanicyathida hill, 1972, p.
130, nom. null.; =Korovinellina Debrenne, 1991, p. 219]
Initial chambers hollow and elongate;
pillars developed in subsequent chambers;
inner wall invaginal. lower Cambrian (Bot.1–
Bot.3).
Family KOROVINELLIDAE
Khalfina, 1960
[Korovinellidae KhAlFinA, 1960, p. 80] [=Kazachstanicyathidae KonYush-
Kov, 1967, p. 106; =Kazakhstanicyathidae hill, 1972, p. 130, nom.null.]
Outer and inner walls with simple pores.
lower Cambrian (Bot.1–Bot.3).
Korovinella rADugin i n KhAlFinA, 1960, p. 80
[*Clathrodictyon sajanicum YAvorsKY, 1932, p.
614; OD; holotype, YAvorsKY, 1932, fig. 4–5, M,
TsNIGRm 4a,b/4070, St. Petersburg] [=Kazach-
stanicyathus KonYu shKov, 1967, p. 106 (type, K.
fistulatus, OD); =Kazakhstanicyathus hill, 1972,
p. 130, nom. null.]. Outer and inner walls tabular;
chambers of simple segmented tabulae and pillars.
lower Cambrian (Bot.3): Altay Sayan, Kazakh-
stan.——Fig. 133,1a–b. *K. sajanica (YAvorsKY),
Verkhnemonok Formation, Botoman, Sanashtykgol
Spring, western Sayan, Altay Sayan, Russia; a,
oblique transverse section, topotype, PIN 4754/10,
×10 (Debrenne, Zhuravlev, & Kruse, 2002); b,
holotype, TsNIGRm 4a,b/4070, transverse section
of modular skeleton, ×10 (Yavorsky, 1932).
Bicoscinus Debrenne, 1977a, p. 127 [*B. sdzuyi; OD;
holotype, Debrenne, 1977a, pl. 14,2, MNHN
M80058, specimen IRH13-1d, Paris]. Outer wall
aporose (possibly rudimentary); inner wall simple;
tabulae. lower Cambrian (Bot.1): Morocco.——Fig.
133,2. *B. sdzuyi, Issafen Formation, Botoman, Jbel
Irhoud, Morocco, holotype, MNHN M80058,
specimen IRH13-1d, oblique longitudinal section,
×5 (Debrenne, 1977a).
Suborder ALTAICYATHINA
Debrenne, 1991
[Altaicyathina Debrenne, 1991, p. 219]
Initial chambers subspherical; pillars
present in initial and subsequent chambers.
lower Cambrian (Bot.1–Bot.2).
Family ALTAICYATHIDAE
Debrenne & Zhuravlev, 1992
[Altaicyathidae Debrenne & ZhurAvlev, 1992b, p. 118]
Outer and inner walls with simple pores.
lower Cambrian (Bot.1–Bot.2).
Altaicyathus vologDin, 1932, p. 26 [*A. notabilis; M;
lectotype, vologDin, 1932, pl. 1,5; SD Debrenne
& ZhurAvlev, 1992b, p. 48, TsNIGRm 290/2957,
St. Petersburg] [=Praeactinostroma KhAlFinA, 1960,
p. 81 (type, Actinostroma vologdini YAv or s KY ,
1932, p. 613, OD); =Cambrostroma vlAs ov, 1961,
p. 29 (type, C. rossicum, OD); =Abakanicyathus
KonYushKov in Zhur Avl evA , KonYushKov, &
roZAnov, 1964, p. 127 (type, A. karakolensis, OD),
for discussion, see Debrenne & ZhurAvlev, 1992b,
p. 119; =Altaicyathus notabilis vologDin, 1932,
p. 26)]. Outer and inner walls tabular; chambers
of simple segmented tabulae and pillars; exaules
and astrorhizae may be present. lower Cambrian
(Bot.1–Bot.2): Altay Sayan, Mongolia, Far East,
United States.——Fig. 134. *A. notabilis, Verkh-
neynyrga Formation, Botoman, Lebed’ River,
Altay Mountains, Altay Sayan, Russia, lectotype,
TsNIGRm 290/2957, longitudinal section, ×9
(Vologdin, 1932).
NOMINA DUBIA
Adaecyathus Fonin in Zhu rAvl ev, Zh urAv levA, &
Fonin, 1983, p. 28 (Fonin in KrAsnoPe evA, 1978,
p. 81, nom. nud.) [*A. gravis; OD].
Tuvacnema
Fig. 128. Tuvacnemidae (p. 152).
Korovinella: I
deleted comma
from simple
segmented
here too, okay?

Systematic Descriptions of the Archaeocyatha 157
Syringocnema
c
a
b
d
Fig. 129. Syringocnemidae (p. 152).

158 Treatise Online, number XX
Syringothalamus
2c
2a
Pseudosyringocnema
1a
1b
Fig. 130. Syringocnemidae (p. 152–154).
2b

Systematic Descriptions of the Archaeocyatha 159
Araneocyathus vologDin in simon, 1941, p. 5
(vologDin, 1937b, p. 466, nom. nud.) [*A. curvus
vologDin, 1940a, p. 64; SD simon, 1941, p. 5].
Archaeocyathellus ForD, 1873b, p. 135 [*Archaeocy-
athus? rensselaericus ForD, 1873a, p. 211; M].
ARCHAEOCYATHOSPONGIA vo logDi n, 1940a,
p. 27 (class).
Archaeofungiella ZhurAvlevA in ZhAuTiKov & others,
1976, p. 137 [*A. chingisiensis; OD].
ARCHAEOPHYLLIDA oKuliTch, 1943, p. 46, nom.
correct. oKuliTch, 1955a, p. 10, pro order Archaeo-
phyllina oKuliTch, 1943, p. 46]
ARCHAEOPHYLLIDAE vologDin, 1940b, p. 97
[=Archaeophyllidae vologDin, 1931, p. 60, nom.
nud.].
Archaeophyllum vologDin in simon, 1939, p.
21 (vologDin, 1931, p. 61, nom. nud.) [*A.
edelsteini vologDin, 1931, p. 62; SD simon,
1939, p. 21].
BACATOCYATHIDAE ZhurAvlevA, 1960b, p. 268,
nom. correct. hill, 1965, p. 116, pro Batchatocy-
athidae ZhurAvlevA, 1960b, p. 268].
Bacatocyathus vologDin, 1940b, p. 95, nom. correct.
hill, 1965, p. 116, pro Bačatocyathus vologDin,
1940b, p. 95 [*B. kazakevici; OD] [=Batschatocy-
athus vologDin, 1956, p. 878, nom. null.; =Batcha-
tocyathus ZhurAvlevA, 1960b, p. 268, nom. null.].
Beticocyathus simon, 1939, p. 73 [*B. beticus; OD].
BICYATHIDAE vologDin, 1937b, p. 472.
Bicyathus vologDin, 1939, p. 235 (vologDin,
1937b, p. 472, nom. nud.) [*B. angustus; OD].
Butovia vologDin, 1931, p. 63 [*B. serrata; M].
CROMMYOCYATHINA R. beDForD & J. beDForD,
1939, p. 79 (order).
Dendrocyathus oKuliTch & rooTs, 1947, p. 44 [*D.
unexpectans; M].
Echinocyathus H. Termier & G. Termier, 1950, p.
47 [*E. goundafensis; OD] [=Dictyocyathus (Echino-
cyathus) H. Termier & G. Termier, 1950, p. 47,
nom. transl. Debrenne, 1964, p. 207, ex Echino-
cyathus H. Termier & G. Termier, 1950, p. 47].
Echinocyathus vologDin, 1960, p. 424, non H.
Termier & G. Termier, 1950, p. 47 (type, E.
goundafensis; OD) [*E. bilateralis; OD].
ETHMOLYNTHIDAE ZhurAvlevA, 1963b, p. 112,
nom. transl. hill, 1972, p. 51, ex Ethmolynthinae
ZhurAvlevA, 1963b, p. 112.
Ethmolynthus ZhurAvlevA, 1963b, p. 112 [*E. rosa-
novi; OD].
EXOCYATHA oKuliTch, 1943, p. 42 (subclass).
EXOCYATHIDAE R. beDFor D & J. beDForD, 1939,
p. 82.
Exocyathus R. beDForD & J. beDForD, 1937, p. 32
[*E. australis; OD].
Gorskinocyathus vologDin, 1960, p. 422 [*Archaeo-
cyathus gorskinensis vologDin, 1940b, p. 60; OD].
Kameschkovia vologDin, 1957a, p. 183 (vologDin,
1956, p. 880, nom. nud.) [*Labyrinthomorpha perfo-
rata vologDin, 1940b, p. 40; M].
LABYRINTHOCYATHIDAE YAroshevich, 1962,
p. 117.
Labyrinthocyathus YAroshevich, 1962, p. 117 [*L.
grandiporosus; M].
Labyrinthomorpha vologDin, 1931, p. 35 [*L. tolli;
M] [=Labirinthomorpha vologDin, 1928, p. 32,
nom. nud.].
LABYRINTHOMORPHIDA vologDin, 1961, p.
180 (order).
Fig. 131. Syringocnemidae (p. 154).
Williamicyathus
c
a
b

160 Treatise Online, number XX
LABYRINTHOMORPHIDAE vologDin, 1962a, p.
125 [=Labirinthomorphidae vologDin, 1928, p.
32, nom. nud.].
LABYRINTHOMORPHINA vologDin, 1961, p.
180 (superorder), nom. transl. vologDin, 1962a,
p. 125, ex order Labyrinthomorphida vologDin,
1961, p. 180].
LEECYATHIDAE vologDin, 1957c, p. 495 [=Leecy-
athidae vologDin, 1956, p. 879, nom. nud.].
Leecyathus vologDin, 1957c, p. 495 [*Archaeocyathus
yavorskii vologDin, 1931, p. 86; OD] [=Zeecyathus
vologDin, 1956, p. 879, nom. nud., lapsus calami
pro Leecyathus].
Leiocyathus vologDin, 1959a, p. 671 [*L. inaequi-
taenialis; OD].
Nevadacyathus oKuliTch, 1943, p. 59 [*Archaeocy-
athus septaporus oKuliTch, 1935, p. 101; M].
Pinacocyathus R. beDFor D & W. R. beDF orD, 1934,
p. 4 [*P. spicularis; M].
Potekhinocyathus vologDin, 1957d, p. 699 [*P.
bateniensis; M].
Protocyclocyathus vologDin, 1955, p. 142 [*Cyclo-
cyathus irregularis vologDin, 1940b, p. 62; M].
RHIZACYATHIDAE R. beD Fo rD & J. beD Fo rD ,
1939, p. 69.
Rhizacyathus R. beD ForD & J. beDF orD, 1939, p. 69
[*Protopharetra radix R. beDF orD & J. beD For D,
1937, p. 28; OD].
Salopicyathus vologDin, 1962c, p. 86 [*S. compla-
natoporosus; OD].
Fragilicyathus
Kruseicnema
1a
1b
2
Fig. 132. Kruseicnemidae and Fragilicyathidae (p. 154–155).

Systematic Descriptions of the Archaeocyatha 161
Bicoscinus
Korovinella
1a
1b 2
Septocyathus vologDin, 1937b, p. 468 [*S .
pedaschenkoi; M].
Serligocyathus vologDin, 1959a, p. 671 [*S. luka-
shevi; OD].
SOMPHOCYATHIDAE oKuliTch, 1935, p. 98.
Somphocyathus TAYl or , 1910, p. 134 [*S. coral-
loides; M].
Sphinctocyathus (Sphinctocyathus) Zhu r Avl evA ,
1960b, p. 304 [*S. (S.) oimuranicus; OD].
Squamella vologDin, 1977, p. 75,non borY De sAinT-
vincenT, 1826, p. 90 (type, S. limulina; M), rotifer
[*S. prima; OD] [=Squamellicyathus vologDin,
1977, p. 22, nom. nud.].
TAB ULACY ATHI DA vologDin, 1956, p. 878
(order), nom. correct. hill, 1972, p. 121 pro Tabu-
locyathida vologDin, 1956, p. 878, lapsus calami.
TABULACYATHIDAE vologDin, 1956, p. 878,
nom. correct. hill, 1972, p. 123, pro Tabulath-
yathidae vologDin in rePin A & others, 1964, p.
249, nom. correct. pro Tabulocyathidae vologDin,
1956, p. 878, lapsus calami.
Tabulacyathus vologDin, 1932, p. 30 [*T. taylori;
M] [=Tabulocyathus vologDin, 1937b, p. 471,
nom. null.]
TABULOIDEA vologDin, 1957a, p. 183 (class).
TANNUOLACYATHIDAE Debrenne, 1964, p. 188.
Tannuolacyathus vologDin, 1957c, p. 496 [*T.
multiplex; OD].
TEREKTIGOCYATHIDAE vologDin, 1962b, p.
419.
Terektigocyathus vologDin, 1962b, p. 420 [*T.
primus; OD].
Tersia vologDin, 1931, p. 70 [*T. filiforma; M].
Tersiella vologDin, 1962a, p. 129 [*Tersia nodosa
vologDin, 1940a, p. 34; OD].
THALASSOCYATHIDAE vologDin, 1962a, p. 116.
Fig. 133. Korovinellidae (p. 156).
Thalassocyathus vologDin, 1957d, p. 699 [*T.
acutatus; M].
Torgaschinocyathus vologDin, 1957d, p. 699 [*T.
spinosus; M].
Turgidocyathus vologDi n, 1960, p. 422 [*T. ippoli-
tovensis; OD].
Altaicyathus
Fig. 134. Altaicyathidae (p. 156).

162 Treatise Online, number XX
Tuvacyathus volog Din , 1940a, p. 112 (volog Din ,
1937b, p. 471, nom. nud.) [*T. mollimurus; M].
URALOCYATHIDAE vologDin & ZhurAvlevA
in vologDin, 1956, p. 878 [=Vacuocyathidae
vologDin, 1962c, p. 77].
Vacuocyathus oKuliTch, 1950a, p. 392, nom . nov.
pro Coelocyathus vologDin, 1939, p. 237, non
sArs, 1857, p. 126, cnidarian, nec schlüTer,
1886, p. 899, cnidarian [*Coelocyathus kidrjass-
ovensis vologDin, 1939, p. 237, OD; =Coelocyathus
kidrjassovensis vologDin, 1937b, p. 478, nom.
nud.] [=Uralocyathus ZhurAvl evA, 1960b, p. 102
(type, Coelocyathus kidrjassovensis vologDin, 1939,
p. 237, OD), nom. nov. pro Coelocyathus vologDin,
1939, p. 237, archaeocyath].
VESICULOIDA vologDin, 1956, p. 878 (order).
VESICULOIDAE vologDin, 1931, p. 34, invalid
family-group name based on unavailable genus
name.
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Systematic Descriptions of the Archaeocyatha 167
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Systematic Descriptions of the Archaeocyatha 185
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