Lower Permian Bryozoa from Ellesmere Island (Canada

eschweizerbartxxx

0031– 0220/07/0081–0017 $ 5.40

© 2007 E. Schweizerbart’sche Verlagsbuchhandlung, D–70176 Stuttgart

Paläontologische Zeitschrift2007, Vol.

81/1

, p. 17–28, 31-03-2007

123456789

10111213141516171819202122232425262728293031323334353637383940414243444546474849505152535455565758

Lower Permian Bryozoa from Ellesmere Island (Canada)

A

NDREJ

E

RNST

, Kiel & H

ANS

A

RNE

N

AKREM

, Blindern

with 6 figures and 5 tables

E

RNST

, A. & N

AKREM

, H.A. 2007. Lower Permian Bryozoa from Ellesmere Island (Canada). – PaläontologischeZeitschrift

81

(1): 17–28, 6 Abb., 5 Tab., Stuttgart, 31. 3. 2007.

Abstract:

A collection of fossils sampled during the 1898–1902 expedition of the

Fram

to the Canadian Arctic Is-lands includes abundant bryozoans from the Lower Permian (Artinskian) Great Bear Cape Formation of EllesmereIsland. From this material a new genus with one new species –

Nansenopora peculiaris

n. gen., n. sp. – as well asthree new species –

Streblotrypella arctica

n. sp.,

Phragmophera patricki

n. sp.

and

Kallodictyon spinatum

n. sp. –are described. Furthermore, the species

Ulrichotrypa ramulosa

B

ASSLER

, 1929 is reported for the first time from theLower Permian of the Arctic region.

Keywords:

Bryozoa • taxonomy • Lower Permian • Canada • palaeobiogeography

Kurzfassung:

Geologische Sammlungen, welche während der Expedition der

Fram

zu den Inseln des arktischen Ka-nada gesammelt wurden, enthalten zahlreiche Bryozoen aus der unterpermischen (Artinskium) Great Bear Cape For-mation von der Insel Ellesmere. Von diesem Material werden eine neue Gattung mit einer neuen Art

–

Nansenoporapeculiaris

n. gen., n. sp. – sowie drei neue Arten –

Streblotrypella arctica

n. sp.,

Phragmophera patricki

n. sp.

und

Kallodictyon spinatum

n. sp. – beschrieben. Außerdem wird die Art


B

ASSLER

, 1929 zum ers-ten Mal im Unterperm aus der arktischen Region festgestellt.

Schlüsselwörter:

Bryozoa • Taxonomie • Unteres Perm • Kanada • Paläobiogeographie

Introduction

The Natural History Museum in Oslo houses extensivegeological collections from the 1898–1902 “Second Ex-pedition in the Fram” to the Canadian Arctic Islands(Fig. 1). The Natural History Museum was formerly Pa-laeontological Museum (prefix PMO), University ofOslo, Norway, and the bryozoans figured herein are keptin the collection of type and illustrated material. Thematerial was collected during the expedition led by Cap-tain Otto Sverdrup (1854–1930), which also includedbotanical, geographical, geological and zoological field-work in the unmapped Arctic area. Per Schei (1875–1905) was the expedition geologist, and he was respon-sible for collecting the samples investigated here. Themuseum collection comprises 5800 catalogued palaeon-tological specimens, including 58 types. Until now, onlyone new bryozoan species has been described from this

material: the Devonian

Fenestella

(s.l.)

scheii

T

OLMA-

CHOFF

, 1926, named after the expedition geologist.T

SCHERNYSCHEW

& S

TEPANOW

(1916) illustrated andbriefly described 17 bryozoan taxa from this unit, whichat the time was believed to be of late Carboniferous age.

The current investigation clarifies questions raisedin M

OROZOVA

& K

RUCHININA

(1986: 25) about their re-ported occurrence of several Artinskian species in theolder Belcher Channel (Sakmarian) and younger Assist-ance (Ufimian) formations. In the revised stratigraphicscheme for the Great Bear Cape area (B

EAUCHAMP

&H

ENDERSON

1994), both these formations are now partof the Great Bear Cape Formation, and material in M

O-

ROZOVA

& K

RUCHININA

(1986) must be considered tobe of Artinskian age. The following species describedby M

OROZOVA

& K

RUCHININA

(1986) from the BelcherChannel Formation should, thus, be considered to derivefrom the Great Bear Cape Formation:

Addresses of the authors: Andrej Ernst, Institut für Geowissenschaften der Christian-Albrechts-Universität zu Kiel, Ludewig-Meyn-Str. 10, D-24118 Kiel, Germany; e-mail <[email protected]>. – Hans Arne Nakrem, Natural History Museum, Universityof Oslo, PO Box 1172 Blindern, NO-0318 Oslo, Norway; e-mail <[email protected]>

eschweizerbartxxx

123456789

10111213141516171819202122232425262728293031323334353637383940414243444546474849505152535455565758

18

A

NDREJ

E

RNST

& H

ANS

A

RNE

N

AKREM

Ramiporidra variolata

(S

HUL

’

GA

-N

ESTERENKO

, 1933)

Tabulipora arcticensis

R

OSS

& R

OSS

, 1962

Rhombotrypella invulgata

T

RIZNA

, 1948

Clausotrypa monticola

(E

ICHWALD

, 1860)

Streblascopora solita

M

OROZOVA

, 1986 in

M

OROZOVA

& K

RUCHININA

1986

Polypora ignota

M

OROZOVA

, 1986 in M

OROZOVA

&K

RUCHININA

1986

Shulgapora pseudokolvae

(S

HUL

’

GA

-N

ESTERENKO

, 1952)

In an initial study of samples from the current collectionE

RNST

& N

AKREM

(2005) identified 37 bryozoan taxawhich are locally abundant (Fig. 2).

Material and methods

The current investigation is based on material fromGreat Bear Cape, Bjorne Peninsula, SW Ellesmere Is-land, from the Great Bear Cape (GBC) Formation.When T

SCHERNYSCHEW

& S

TEPANOV

(1916) pub-lished their paper on brachiopods and bryozoans, the in-vestigated unit was believed to be of Late Carbonifer-ous age (see also S

CHEI

in S

VERDRUP

1904). Since then,comprehensive work has been carried out on the geolo-gy of the Canadian Arctic Islands the Sverdrup Basinand the modern stratigraphic definitions are presented

in detail in B

EAUCHAMP

& H

ENDERSON

(1994). The up-permost part of the unit is missing at the Bjorne Penin-sula locality (Fig. 3). Rich conodont faunas containspecies of

Adetognathus

,

Sweetognathus

,

Neostrepto-gnathodus

and

Mesogondolella

, placing the unit in thePermian P6b, P7 and P8 (Artinskian) and the top in theP9 (Artinskian – Kungurian) conodont zones of B

EAU-

CHAMP

et al. (1989). The top of the formation is missingdue to Tertiary uplift and subsequent glacial erosionwhere the current material derives from, but the age ofthe investigated bryozoan fauna is most probably Artin-skian.

The museum collection contains both specific sam-ples with visible bryozoans, mainly fenestrates, and alsolarger bioclastic limestone samples which were cut toobtain bryozoans in different internal orientations.About 70 acetate peels and 140 thin sections have beenprepared from selected samples from this material. Allthin sections were prepared from a single block20x20x20 cm, and taxa were described using a binocularmicroscope.

Repository

: Investigated material is housed at the NaturalHistory Museum, University of Oslo, under numbers PMO203.513-203.527, 208.910-208.967.

Fig. 1. Map of Canadian Arctic Islands with localities mentioned in the text.

eschweizerbartxxx

Lower Permian Bryozoa from Ellesmere Island (Canada)

19

123456789

10111213141516171819202122232425262728293031323334353637383940414243444546474849505152535455565758

Systematic palaeontology

Class Stenolaemata B

ORG

, 1926Order Trepostomida U

LRICH

, 1882Suborder Amplexoporina A

STROVA

, 1965Family Anisotrypidae D

UNAEVA

& M

OROZOVA

, 1967

Genus

Ulrichotrypa

B

ASSLER

, 1929

Stenopora

D

YBOWSKI

1877: 65–78.

Ulrichotrypa

B

ASSLER

1929: 55; R

OMANTCHUK

& K

ISELEVA

1968: 56; M

OROZOVA

1970: 86; A

STROVA

1978: 169–170; E

RNST

2001: 141.

Type species

:


B

ASSLER

, 1929; LowerPermian; Timor (Indonesia).

Diagnosis

: Dendroid, massive and encrusting colonies.Autozooecia with circular to polygonal apertures. Auto-zooecial walls in the exozone irregularly thickened. Di-aphragms in the autozooecia lacking. Exilazooecia indifferent numbers. Abundant small stylets in the autozo-oecial walls of the exozone arranged rosette-like. Stel-latopores varying in number from species to species.

Comparison

: The genus

Ulrichotrypa

B

ASSLER

, 1929differs from

Ulrichotrypella

R

OMANTCHUCK

, 1967 bylacking diaphragms in autozooecia, and from

Hinganel-la

R

OMANTCHUCK

, 1967 by absence of diaphragms andpresence of stellatopores.

Stratigraphic

and

geographic range

:

Lower to Upper Perm-ian of Indonesia; Upper Permian, Zechstein of Germany; Low-er Permian of Canada.

Ulrichotrypa ramulosa BASSLER, 1929Figs. 4A–F, Tab. 1

1929 Ulrichotrypa ramulosa BASSLER: 55–56, pl. 10 figs.1–6.

Material: PMO 208.915b, 208.921a, 208.927b, 208.928b,208.932c, 208.933a, 208.941b, 208.955a.

Description: Branching colonies 1.88–2.75 mm in dia-meter with 0.45–0.63 mm wide exozones and 0.63–

Fig. 2. Fragment of the rock thin section (2 × 3 cm) bearing abundant bryozoans, sample no. PMO 208.914. GreatBear Cap Formation (Artinskian), Ellesmere Island, Canada. – P, Polypora sp., R, Rhombotrypella sp., D, Dyscritellavulgata GORJUNOVA, 1975, Ph, Phragmophera patricki n. sp.

eschweizerbartxxx

123456789

10111213141516171819202122232425262728293031323334353637383940414243444546474849505152535455565758

20 ANDREJ ERNST & HANS ARNE NAKREM

1.85 mm wide endozones. Secondary overgrowths com-mon. Autozooecia long, polygonal in cross section inthe endozone, bending sharply and becoming rounded incross section in exozone. Autozooecial apertures round-ed to oval, spaced 0.18–0.32 mm from centre to centre.Diaphragms absent. Stellatopores abundant, restrictedto the exozone, arranged densely in a single row be-tween autozooecial apertures, forming a regular polygo-nal pattern, having irregular stellate shape in cross sec-

tion. Exilazooecia common, rounded to oval in crosssection, restricted to exozone. Autozooecial walls 0.01mm thick, having dark medial lining in endozone; 0.09–0.19 mm thick, displaying reversal U-shaped laminationand monilae-shaped thickenings in exozone. Acantho-styles absent.

Comparison: The species Ulrichotrypa ramulosaBASSLER, 1929 differs from the most similar species U.zonata BASSLER, 1929 in having larger and more widelyspaced apertures (aperture diameter 0.18–0.32 mm vs.0.15–0.2 mm in U. zonata; 3.0–4.5 apertures per 2 mmvs. 7 apertures per 2 mm in U. zonata).

Stratigraphic and geographic range: Lower Permian; Timor,Indonesia. Great Bear Cape Formation, Lower Permian (Art-inskian); Ellesmere Island, Arctic Canada.

Genus Nansenopora n. gen.

Derivatio nominis: The genus Nansenopora n. gen. is namedafter the famous Norwegian arctic explorer and researcherFRIDTJOF NANSEN.Type species: Nansenopora peculiaris n. gen., n. sp.

Tab. 1. Measurements of Ulrichotrypa ramulosa BASSLER, 1929 [in mm]. – N = number of measurements; X = mean;SD = standard deviation; VC = coefficient of variation; MIN = minimal value; MAX = maximal value.

N X SD CV MIN MAX

branch diameter 7 2.32 0.404 17.429 1.88 2.75

exozone wideness 7 0.53 0.089 16.657 0.45 0.63

endozone wideness 7 1.25 0.518 41.441 0.63 1.85

wall thickness in exozone 10 0.13 0.033 24.727 0.09 0.19

aperture diameter 30 0.25 0.033 13.606 0.18 0.32

aperture spacing from centre to centre 15 0.39 0.052 13.094 0.30 0.46

exilazooecia diameter 30 0.08 0.026 34.137 0.05 0.18

stellatopore diameter 10 0.04 0.009 20.472 0.03 0.06

Fig. 3. Lithological log, stratigraphy and geological de-velopment of the investigated units. Modified from BEAU-CHAMP & HENDERSON (1994).

Fig 4. A–F: Ulrichotrypa ramulosa BASSLER, 1929. A.longitudinal section, PMO 208.932c, scale bar 1 mm; B.cross section, PMO 208.928b, scale bar 1 mm; C. PMO208.932c, longitudinal section of the exozone, scale bar1 mm; D–F. tangential section, D. PMO 203.525, tangen-tial section, scale bar 0.5 mm; E. PMO 208.955a, tan-gential section, scale bar 0.5 mm; F. PMO 208.941b,scale bar 0.2 mm. – G–L: Nansenopora peculiaris n.gen., n. sp. G. cross section, paratype PMO 208.934a,scale bar 1 mm; H–I. holotype PMO 208.960a. H. longi-tudinal section of an encrusting colony, scale bar 1 mm;I. tangential section, scale bar 0.5 mm; J. longitudinalsection, paratype PMO 203.518, scale bar 0.5 mm; K.tangential section displaying stellatopores, PMO208.933b, scale bar 25 μm; L. longitudinal section dis-playing stellatopores, PMO 208.959a, scale bar 0.1 mm.

eschweizerbartxxx

Lower Permian Bryozoa from Ellesmere Island (Canada) 21

123456789

10111213141516171819202122232425262728293031323334353637383940414243444546474849505152535455565758

eschweizerbartxxx

123456789

10111213141516171819202122232425262728293031323334353637383940414243444546474849505152535455565758


Diagnosis: Ramose branching and encrusting colonies.Autozooecia long, polygonal in cross section in endo-zone, becoming rounded in exozone. Hemiphragmsabundant, thick and long. Stellatopores abundant, re-stricted to the exozone, arranged in a single row betweenautozooecial apertures forming a regular pattern, havingwide hyaline cores. Exilazooecia rare.

Comparison: The genus Nansenopora n. gen. is similarto the genus Ulrichotrypella ROMANTCHUCK, 1967 inhaving laminated walls and stellatopores. However, thenew genus differs in having thick and abundant hemi-phragms.

Stratigraphic and geographic range: Great Bear Cape For-mation, Lower Permian (Artinskian); Ellesmere Island, ArcticCanada.

Nansenopora peculiaris n. gen., n. sp.Figs. 4G–L, Tab. 2

Derivatio nominis: The species name (from Latin peculiaris– peculiar) refers to the peculiar morphology of this genus.Type material: Holotype (PMO 208.960a) and two paratypes(PMO 203.518 and PMO 208.934a)Type locality: Ellesmere Island, Arctic Canada.Type horizon: Great Bear Cape Formation, Lower Permian(Artinskian).Material: PMO 208.915a, 208.919a, 208.926a, 208.930b,208.933b, 208.941c, 208.942, 208.943, 208.948, 208.955b,208.956, 208.959a, 208.963a.

Diagnosis: As for genus.

Description: Ramose branching and encrusting colo-nies. Branching colonies 1.26–2.26 mm in diameterwith 0.27–0.78 mm wide exozones and 0.18–0.97 mmwide endozones. Encrusting colonies 0.48–0.63 mm inthickness. Secondary overgrowths common. Autozooe-cia long, polygonal in cross section in the endozone,bending sharply and becoming rounded in cross sectionin exozone. Autozooecial apertures rounded to oval,spaced 0.24–0.44 mm from centre to centre. Basal dia-phragms absent. Hemiphragms abundant, long, curved

distally, up to 7 in each autozooecium. Stellatoporesabundant, restricted to the exozone, arranged densely ina single row between autozooecial apertures, forming aregular polygonal pattern, having stellate shape in crosssection. Exilazooecia rare, small, rounded in cross sec-tion, restricted to exozone, having wide hyaline coresand narrow laminated sheaths. Autozooecial walls 0.02–0.03 mm thick, having dark medial lining in endozone;0.08–0.18 mm thick, displaying reversal U-shaped lam-ination in exozone. Acanthostyles absent.

Comparison: As for genus.

Order Rhabdomesida ASTROVA & MOROZOVA, 1956Suborder Rhabdomesina ASTROVA & MOROZOVA, 1956

Family Nikiforovellidae GORJUNOVA, 1975

Genus Streblotrypella NIKIFOROVA, 1948

Type species: Streblotrypa major ULRICH, 1889; Lower Car-boniferous, USA.

Tab 2. Measurements of Nansenopora peculiaris n. gen., n. sp. [in mm]. – Abbreviations as in Tab. 1.

N X SD CV MIN MAX

branch diameter 6 1.79 0.401 22.467 1.26 2.26

exozone wideness 6 0.55 0.173 31.797 0.27 0.78

endozone wideness 6 0.70 0.274 39.444 0.18 0.97

aperture diameter 26 0.14 0.035 25.996 0.08 0.19

aperture spacing from centre to centre 20 0.34 0.066 19.223 0.24 0.44

stellatopore diameter 30 0.04 0.007 17.488 0.03 0.05

exilazooecia diameter 10 0.05 0.013 25.891 0.03 0.07

Fig. 5. A–G: Streblotrypella arctica n. sp. A. cross sec-tion of the branch, PMO 208.935b, scale bar 0.2 mm;B. cross section of the branch, PMO 208.924, scale bar0.2 mm; C. tangential section displaying apertures andmetazooecia, paratype PMO 208.916, scale bar 0.2 mm;D–E. PMO 208.930a. D. tangential section displayingapertures and metazooecia, scale bar 0.2 mm; E. tan-gential section displaying apertures and metazooecia,scale bar 0.5 mm; F. tangential section of a dichotomis-ing branch, paratype PMO 208.932a, scale bar 0.5 mm;G. longitudinal section displaying autozooecia, PMO208.964, scale bar 0.5 mm. – H–M: Phragmopherapatricki n. sp. H. tangential section, paratype PMO203.522, scale bar 0.5 mm; I. tangential section, holo-type PMO 208.912, scale bar 0.5 mm; J. longitudinalsection, PMO 203.520, scale bar 0.5 mm, K. cross sec-tion, PMO 208.953, scale bar 0.5 mm; L. cross section ofthe branch, showing exilazooecium, PMO 208.921c,scale bar 0.2 mm; M. cross section displaying arrange-ment of rods, PMO 208.953, scale bar 0.2 mm.

eschweizerbartxxx


123456789

10111213141516171819202122232425262728293031323334353637383940414243444546474849505152535455565758

eschweizerbartxxx

123456789

10111213141516171819202122232425262728293031323334353637383940414243444546474849505152535455565758


Diagnosis: Slender dendroid colonies, with dichotomis-ing branches. Branches circular to semicircular in crosssection. Autozooecia long, budded from a distinct medialaxis. Autozooecial apertures are circular to oval in shape,with proximal peristomes. Hemisepta absent; diaphragmsoccasionally present. Acanthostyles present or absent.Metazooecia few or many. Longitudinal ridges occurring.

Comparison: The genus Streblotrypella NIKIFOROVA,1948 differs from the most similar genus NikiforovellaNEKHOROSHEV, 1948 in having longer autozooecia.

Stratigraphic range: Lower Carboniferous to Lower Perm-ian.

Streblotrypella arctica n. sp.Figs. 5A–G, Tab. 3

Derivatio nominis: The species name refers to the first recordof this species from Arctic.Type material: Holotype (PMO 208.930a) and two paratypes(PMO 208.916 and PMO 208.932a).Type locality: Ellesmere Island, Arctic Canada.Type horizon: Great Bear Cape Formation, Lower Permian(Artinskian).Material: PMO 208.913, 208.914a, 208.921b, 208.924,208.925, 208.931, 208.935b, 208.937, 208.938a, 208.939,208.961a, 208.962a, 208.963b, 208.964, 208.965, 208.966.

Diagnosis: Branching colonies; autozooecia long, bud-ding in spiral order from poorly defined medial axis.Autozooecial apertures oval, forming 6 longitudinalrows at colony surface. Abundant heterozooecia (meta-zooecia) present.

Description: Dendroid colonies with occasionally di-chotomising branches. Branches 0.37–0.60 mm in dia-meter. Autozooecia long, occurring in 6 longitudinalrows, budded from a central axis in spiral order. Auto-zooecial apertures oval, with proximal peristomes, 6–7spaced per 2 mm longitudinally. Diaphragms rare.Metazooecia abundant, up to 8 spaced between adjacentautozooecia, rounded to oval in cross section, restrictedto exozone, often closed by skeleton at colony surface.

Comparison: The species Streblotrypella arctica n. sp.differs from S. ignota (BASSLER, 1929) in having small-er apertures (0.07–0.10 mm vs. 0.25 mm in S. ignota).

Order Cryptostomida VINE, 1884Suborder Intraporina GORJUNOVA, 1993

Family Phragmopheridae GORJUNOVA, 1969

Genus Phragmophera GORJUNOVA, 1969

Phragmophera GORJUNOVA 1969: 129; KARKLINS 1983: 506;GORJUNOVA & LAVRENTJEVA 1993: 108.

Type species: P. eximia GORJUNOVA, 1969, Upper Carbonif-erous; Ural Mountains.

Diagnosis: Branching bifoliate colonies. Autozooecia inexozones usually at right angles to the mesotheca, ini-tially recumbent, arranged in rhombic pattern; in endo-zones irregularly subcircular in cross section. Autozooe-cial walls extending into peristomes at colony surface.Autozooecial wall boundaries narrowly serrated. Rodsaligned along autozooecial boundaries. Extrazooecialskeleton laminated. Cystiphragms closed; positioned inthe outer endozones and exozones of autozooecia, inregular series. Basal diaphragms thin, slightly curved,regularly spaced in endozones and exozones. Mesozo-oecia abundant, irregularly polygonal to subcircular incross section, variable in size, with few or no dia-phragms, regularly arranged throughout colonies, local-ly filled by extraskeletal material. Mesozooecial wallsextending into peristomes at colony surface. Acantho-styles variable in size or absent.

Comparison: The single genus of the family, Phragmo-phera GORJUNOVA, 1969 differs from other intraporidbryozoans by comprising only branching colonies andby abundant rods as well as abundant cystiphragms.

Stratigraphic and geographic range: Phragmophera eximiaGORJUNOVA, 1969, Upper Carboniferous of the Ural Moun-tains, and P. patricki n. sp. from the Lower Permian of Elles-mere Island (Canada).

Phragmophera patricki n. sp.Figs. 5H–M, 6A–B, Tab. 4

Derivation of the name: The species is named after PatrickWyse Jackson, Irish bryozoologist, who has contributed sig-nificantly to the research of fossil bryozoans.Type material: Holotype (PMO 208.912) and two paratypes(PMO 208.951 and PMO 203.522).Type locality: Ellesmere Island, Arctic Canada.

Tab. 3. Measurements of Streblotrypella arctica n. sp. [in mm]. – Abbreviations as in Tab. 1.

N X SD CV MIN MAX

branch diameter 19 0.47 0.069 14.561 0.37 0.60

aperture width 14 0.09 0.011 11.891 0.07 0.10

metazooecia diameter 8 0.02 0.005 20.574 0.02 0.03

eschweizerbartxxx


123456789

10111213141516171819202122232425262728293031323334353637383940414243444546474849505152535455565758

Type horizon: Great Bear Cape Formation, Lower Permian(Artinskian).Material: PMO 208.911, 208.917, 208.919b, 208.920,208.921c, 208.922, 208.925, 208.930c, 208.934b, 208.936a,208.938b, 208.940a, 208.941d, 208.945a, 208.947, 208.949,208.950, 208.952a, 208.953, 208.954, 208.957, 208.958,208.959b, 208.961b, 208.962b.

Diagnosis: Phragmophera species with small apertures,few and small mesozooecia, and no acanthostyles.

Description: Branching bifoliate colonies. Brancheslens-shaped in cross section with sharp edges, 1.35–3.43mm wide and 0.63–1.29 mm thick. Autozooecia ar-ranged in 7–12 alternating rows on branches, in exo-zones usually at right angles to the mesotheca, irregular-ly subcircular in cross section, spaced 0.40–0.60 mmalong the branch and 0.26–0.42 mm across the branch,measured from centre to centre. Autozooecial walls ex-tending into peristomes at colony surface. Autozooecialwall boundaries narrowly serrated. Rods aligned alongautozooecial boundaries, 0.02–0.05 mm in diameter,spaced densely. Cystiphragms deriving from skeletalcortex, thick-walled, positioned at proximal wall of auto-zooecia in regular rows. Basal diaphragms not observed.Mesozooecia subcircular in cross section, without dia-phragms, narrow, usually filled by skeletal material, ar-ranged 1–2 rows between apertures, locally abundant orabsent. Mesozooecial walls extending into peristomes atcolony surface. Acanthostyles absent. Mesotheca dou-ble-layered, serrated, 0.06–0.09 mm thick. Coloniestend to break alongside the mesotheca.

Comparison: The species Phragmophera patricki n. sp.differs from P. eximia GORJUNOVA, 1969 in the absenceof acanthostyles as well as in having smaller and fewermesozooecia.

Order Fenestellida ASTROVA & MOROZOVA, 1956Suborder Phylloporinina LAVRENTJEVA, 1979

Genus Kallodictyon MOROZOVA, 1981

Kallodictyon MOROZOVA 1981: 47; LAVRENTJEVA 1985: 72.Type species: Kallodictyon perforatum MOROZOVA, 1981;Lower Permian; northern Russia.Diagnosis: Reticulate colonies consisting of anastomo-sing branches. Autozooecia long, polygonal in crosssection in endozone, becoming oval in exozone, origi-nating alongside the reversal wall, arranged in 3–5 rowson branches. Apertures oval, arranged in alternatingrows at obverse colony surface. Heterozooecia (leptozo-oecia after LAVRENTJEVA 1985) representing tubularstructures smaller than autozooecia, restricted to exo-zone, abundant, opening both at obverse and reversecolony sides. Acanthostyles present or absent. Reversalwall thickened.

Comparison: The genus Kallodictyon MOROZOVA,1981 differs from Chainodictyon FOERSTE, 1887 andRhombocladia ROGERS, 1900 in having thickened re-versal wall and abundant heterozooecia.

Stratigraphic and geographic range: Lower Permian, north-ern Russia and Canada.

Kallodictyon spinatum n. sp.Figs. 6C–J, Tab. 5

Derivation of the name: The species refers to the presence ofabundant acanthostyles (from Latin spinae – thorn).Type material: Holotype (PMO 208.941a) and two paratypes(PMO 208.927a and PMO 208.960b).Type locality: Ellesmere Island, Arctic Canada.Type horizon: Great Bear Cape Formation, Lower Permian(Artinskian).Material: PMO 208.910, 208.914b, 208.918, 208.923,208.928a, 208.926b, 208.929, 208.930d, 208.932b, 208.935a,208.936b, 208.940b, 208.944, 208.945b, 208.946, 208.952b,208.961c.

Tab. 4. Measurements of Phragmophera patricki n. sp. [in mm]. – Abbreviations as in Tab. 1.

N X SD CV MIN MAX

aperture width 30 0.15 0.028 19.001 0.10 0.20

aperture length 30 0.30 0.041 13.903 0.23 0.40

aperture spacing along branch 30 0.52 0.057 10.951 0.40 0.60

aperture spacing across branch 30 0.36 0.041 11.519 0.26 0.42

mesozooecial diameter 22 0.04 0.015 43.283 0.02 0.07

branch width 10 2.13 0.571 26.855 1.35 3.43

branch thickness 10 0.96 0.167 17.430 0.63 1.29

eschweizerbartxxx

123456789

10111213141516171819202122232425262728293031323334353637383940414243444546474849505152535455565758


Tab. 5. Measurements of Kallodictyon spinatum n. sp. [in mm]. – Abbreviations as in Tab. 1.

N X SD CV MIN MAX

branch width 7 0.84 0.199 23.850 0.57 1.11

branch thickness 7 0.90 0.150 16.635 0.66 1.10

aperture width 13 0.18 0.023 12.852 0.14 0.22

heterozooecia diameter 20 0.06 0.015 26.035 0.03 0.09

acanthostyle diameter 10 0.03 0.006 22.588 0.02 0.04

Fig. 6. A–B: Phragmophera patricki n. sp. A. cross section of the branch displaying mesotheca, PMO 208.947, scalebar 0.1 mm; B. longitudinal section displaying mesotheca, PMO 208.952a, scale bar 0.1 mm. – C–J: Kallodictyon spina-tum n. sp. C. mid tangential section displaying autozooecial shape and leptozooecia, PMO 208.926b, scale bar 0.5 mm;D–E. oblique section of the branch, displaying autozooecia and leptozooecia, paratype PMO 208.960b, scale bar 1 mmfor D and 0.5 mm for E; F. mid tangential section of fusing branches, PMO 208.936b, scale bar 2 mm; G. tangentialsection displaying apertures, leptozooecia and acanthostyles, PMO 208.928a, scale bar 0.5 mm; H. cross section of thebranch, PMO 208.946, scale bar 0.5 mm; I. cross section of the branch, paratype PMO 208.927a, scale bar 0.5 mm; J.tangential section displaying apertures, leptozooecia and acanthostyles, holotype PMO 208.941a, scale bar 0.2 mm.

eschweizerbartxxx


123456789

10111213141516171819202122232425262728293031323334353637383940414243444546474849505152535455565758

Diagnosis: Reticulate colonies with irregularly anasto-mosing branches. Leptozooecia abundant both on reverseand obverse surface. Abundant acanthostyles present.

Description: Reticulate colonies consisting of irregular-ly anastomosing branches. Autozooecia long, originat-ing alongside the reversal wall, overlapping two follow-ing autozooecial generations, polygonal in cross sectionin endozone, rhombic in mid tangential section, becom-ing oval in exozone, arranged in 3–5 rows on branches.Apertures oval, arranged in alternating rows at obversecolony surface. Leptozooecia abundant, irregularlyspaced between autozooecial apertures on the obversecolony side; abundant and arranged in longitudinal rowsat the reverse colony side. Acanthostyles small, abun-dant both at obverse and reverse colony surface, havingdistinct cores and dark laminated sheaths. Endozonalwalls granular, 0.02 mm thick. Exozonal wall laminat-ed, 0.08–0.14 mm thick. Longitudinal ridges separatingleptozooecia at the reverse colony surface developed.

Comparison: The species Kallodictyon spinatum n. sp.differs from K. perforatum MOROZOVA, 1981 in havingnarrower branches (0.57–1.10 mm vs. 1.50–1.70 mm inK. perforatum) and in presence of acanthostyles.

Discussion

The majority of the bryozoans from the Great Bear CapeFormation discussed herein have an Early Permian(Sakmarian – Artinskian) distribution in the Urals andthe Timan-Pechora region of Russia, but a stratigraphic-al Artinskian – Kungurian range in Svalbard, ArcticNorway. The genus Phragmophera GORJUNOVA, 1969was previously only known from the Upper Carbonifer-ous of the Urals. The new species from the Great BearCape Formation is the second known species of this ge-nus. The genus Kallodictyon MOROZOVA, 1981 is previ-ously known only from the Lower Permian of northernRussia. The genus Streblotrypella NIKIFOROVA, 1948 iscommon in the Carboniferous, but few species were re-ported from the Lower Permian of Timor, Mongolia andMiddle Asia. The species Ulrichotrypa ramulosaBASSLER, 1929 is previously known only from the Low-er Permian of Timor.

The overall species composition of the Great BearCape Formation is most similar to the Kungurian faunasof Spitsbergen (Svalbard) and Timan-Pechora (westernSiberia), as well as with the Artinskian of Bjørnøya(Svalbard). Faunal migrations through time from theSverdrup Basin to the Svalbard and Timan-Pechora Ba-sins may explain this discrepancy in stratigraphic posi-tion. The Great Bear Cape Formation fauna is quitedifferent from the Sakmarian and Artinskian faunas ofTiman-Pechora.

Acknowledgements

BENOIT BEAUCHAMP and CHARLES M. HENDERSON (Calgary)are thanked for help on the geology of the investigated materi-al, as well as suggestions for language improvements. Wethank CAROLINE BUTTLER (Cardiff) for helpful taxonomiccomments. The Deutsche Akademische Austauschdienst(DAAD) supported ANDREJ ERNST’S study visit to the NaturalHistory Museum, University of Oslo, with a one-year fellow-ship (grant D/02/00949).

References

ASTROVA, G.G. 1965. Morphologiâ, istoriâ razvitiâ i sistema ordo-vikskih i siluriiskih mšanok [Morphology, history of develop-ment, and system of the Ordovician and Silurian Bryozoa]. –Trudy Instituta Akademii Nauk SSSR 106:1–432.

ASTROVA, G.G. 1978. Istoriâ razvitiâ, sistema i filogeniâ mšanok.Otrâd Trepostomata [The history of development, system, andphylogeny of the Bryozoa. Order Trepostomata]. – Trudy Paleon-

Instituta Akademii Nauk SSSR 169: 1–240.ASTROVA, G.G. & MOROZOVA, I.P. 1956. K sistematike mšanok otrâda

Cryptostomata [On systematics of the order Cryptostomata]. –Doklady Akademii Nauk SSSR [Reports of the Academy of Sci-ences of USSR] 110 (4): 661–664.

BASSLER, R.S. 1929. The Permian Bryozoa of Timor. – Paläontologievon Timor. Lieferung 16, Abhandlung 28: 37–90.

BEAUCHAMP, B.; HARRISON, J.C. & HENDERSON, C.M. 1989. UpperPaleozoic stratigraphy and basin analysis of the Sverdrup Basin,Canadian Arctic Archipelago: Part 1, time frame and tectonic ev-olution. – Current Research, Part G, Geological Survey of Cana-da, Paper 89-16: 105–113.

BEAUCHAMP, B. & HENDERSON, C.M. 1994. The Lower PermianRaanes, Great Bear Cape and Trappers Cove formations, Sver-drup Basin, Canadian Arctic: stratigraphy and conodont zonation.– Bulletin of Canadian Petroleum Geology 42 (4): 562–597.

BORG, F. 1926. Studies on recent cyclostomatous Bryozoa. – Zoologis-ka Bidrag fran Uppsala 10: 181–507.

DUNAEVA, N.N. & MOROZOVA, I.P. 1967. Osobennosti razvitiâ i sis-tematieskoe nekotoryh paleozojskih trepostomat[Peculiarities of development and systematic position of somePaleozoic Trepostomata]. – 4: 86–94.

DYBOWSKI, W. 1877. Ueber die Gattung Stenopora LONSDALE, mitbesonderer Berücksichtigung der Stenopora columnarisSCHLOTHEIM sp. – Verhandlungen der Russisch-KaiserlichenMineralogischen Gesellschaft zu St. Petersburg 12 (2): 65–78.

EHRENBERG, C.G. 1831. Animalia invertebrata exclusis insects. Sym-bolae Physicae, seu Icones et descriptiones Corporum Natural-ium novorum aut minus cognitorum, ... – Pars Zoologica 4.

EICHWALD, E. 1860. Lethaea Rossica, ou Paleontologie de la Russie. I.Ancienne Periode. 355–419, 434–435, 450–452, 476–494, Stutt-gart (E. Schweizerbart).

ERNST, A. 2001. Bryozoa of the Upper Permian Zechstein Formation ofGermany. – Senckenbergiana lethaea 81 (1): 135–181.

ERNST, A. & NAKREM, H.A. 2005. Bryozoans from the Artinskian(Lower Permian) Great Bear Cape Formation, Ellesmere Island(Canadian Arctic). – In: MOYANO, G.I.; CANCINO, J.M. & WYSE

JACKSON, P.N., eds., Bryozoan Studies 2004 – Proceedings of the13th International Bryozoology Association Conference, Con-cepcion, Chile, January 2004: 63–68, Balkema Publishing House.

FOERSTE, A.F. 1887. The Clinton Group of Ohio (Bryozoa). – Bulletinof Scientific Laboratory, Denison University 2 (1/2): 71–88,149–176.

GORJUNOVA, R.V. 1969. O novom rode mšanok iz verhnego karbonaUrala [On a new bryozoan genus from the Upper Carboniferousof Urals]. – 2: 129–131.

Paleontologiceskogo

tologiceskogo

pololozenie

Paleontologiceskij zurnal


https://www.researchgate.net/publication/279578821_The_lower_Permian_Raanes_Great_Bear_Cape_and_Trappers_Cove_Formations_Sverdrup_Basin_Canadian_Arctic_stratigraphy_and_conodont_zonation?el=1_x_8&enrichId=rgreq-52c6c94d053b390da122c9d9e5b456be-XXX&enrichSource=Y292ZXJQYWdlOzIyNzA3MjY0NTtBUzoxMjQxODk5MTA1MDc1MjBAMTQwNjYyMDU5MDU5OA==




eschweizerbartxxx

123456789

10111213141516171819202122232425262728293031323334353637383940414243444546474849505152535455565758


GORJUNOVA R.V. 1975. Permskie mšanki Pamira [Permian Bryozoansof the Pamir]. – Trudy Instituta AkademiiNauk SSSR [Transactions of the Paleontological Institute ofAcademy of Sciences of the USSR] 148: 1–125.

GORJUNOVA, R.V. 1993. – In: GORJUNOVA, R.V. & LAVRENTJEVA,V.D. Morfologiâ i sistema kriptostomidnyh mšanok [Morpholo-gy and system of the cryptostome bryozoans]. – Trudy Paleontol-

Instituta Rossiiskoi Akademii Nauk [Transactions ofthe Paleontological Institute of the Russian Academy of Scienc-es] 257: 1–152.

KARKLINS, O.L. 1983. Systematic descriptions for the Suborder Ptilo-dictyina. – In: ROBISON, R.A.. ed., Treatise on Invertebrate Pale-ontology, Part G (1): Bryozoa (revised): 489–529, Boulder (Ge-ological Society of America and University of Kansas Press).

LAVRENTJEVA, V.D. 1979. Phylloporinina – novyj podotrâd paleozo-jskih mšanok [Phylloporinina – a new suborder of PalaeozoicBryozoa]. – 1: 59–68.

LAVRENTJEVA, V.D. 1985. Mšanki podotrâda Phylloporinina [Bryo-zoans of the suborder Phylloporinina]. – Trudy go Instituta Akademii Nauk SSSR [Transactions of the Paleonto-logical Institute of Academy of Sciences of the USSR] 214: 1–100.

MOROZOVA, I.P. 1970. Mšanki pozdnej permi [Late Permian Bryozoa].– Trudy Instituta Akademii Nauk SSSR[Transactions of the Paleontological Institute of Academy of Sci-ences of the USSR] 122: 1–347.

MOROZOVA, I.P. 1981. Pozdnepermskie mšanki severo-vostoka SSSR[Late Paleozoic bryozoans of the Northwest of the USSR]. –Trudy Instituta Akademii Nauk SSSR[Transactions of the Paleontological Institute of Academy of Sci-ences of the USSR] 188: 1–119.

MOROZOVA, I.P. & KRUCHININA, O.N. 1986. Permskie mšanki Arktiki(zapadnyj sektor) [The Permian bryozoans of the Arctic Region(Western Sector)]. – 144 p., Moscow (Nauka).

NEKHOROSHEV, V.P. 1948. Devonskie mšanki Altaâ [Devonian Bryo-zoa of the Altai]. – Paleontologiâ SSSR [Paleontology of theUSSR] 3 (2/1): 1–172.

NIKIFOROVA, A.I. 1948. mšanki Karatau[Lower Carboniferous Bryozoa of Karatau]. – 53 p., Alma-Ata(Akademiâ Nauk Kazahskoi SSSR).

ROGERS, A.F. 1900. A new genus and species of Bryozoa from the CoalMeasures of Kansas and Missouri. – The Kansas UniversityQuarterly A (9): 1–12.

ROMANTCHUK, T.V. & KISELEVA, A.V. 1968. Novye pozdnepermskiemšanki Dalnego Vostoka [New Late Permian bryozoans of theFar East]. – 4: 55–60.

ROMANTCHUK, T.V. 1967. Novye mšanki otrâda Trepostomata iz verh-nej permi Habarovskogo kraâ [New bryozoans of the OrderTrepostomata from the Upper Permian of the Khabarovsk Re-gion]. – 2: 69–63.

ROSS, C.A. & ROSS, J.R.P. 1962. Faunas and correlation of the latePaleozoic rocks of north-east Greenland, pt 4: Bryozoa. – Med-delelser om Grønland 167: 1–65.

SVERDRUP, O. 1904. New Land: Four Years in Arctic Regions. – Vol.I: 1–496, vol. II: 1–504, London (Longmans and Green).

SHUL’GA-NESTERENKO M.I. 1933. Mšankovaâ fauna uglenosnyh i po-duglenosnyh otlozhenij Pecorskogo kraya [Bryozoan fauna fromthe coal-bearing and subjacent series of the Pechora Land]. –Trudy vseso oznogo geologo-razvedo nogo ob’edineni [Trans-actions of the United geological and prospecting Service ofUSSR]: 1:157, Moscow (Akademia Nauk SSSR).

SHUL’GA-NESTERENKO M. I. 1952. Novye ni nepermskie mšanki Pri-ural’â [New Lower Permian bryozoans of Ural Mountains]. –Trudy Paleontologieskogo Instituta Akademii Nauk SSSR[Transactions of the Paleontological Institute of Academy of Sci-ences of the USSR] 37: 1–79.

TOLMACHOFF, I.P. 1926. On the Fossil Faunas from Per Schei's SeriesD from Ellesmere Land. – Report of the Second Norwegian Arc-tic Expedition in the “Fram” 1898–1902, 38: 1–106; Oslo (DetNorske Videnskaps-Akademi i Oslo).

TRIZNA, V.B. 1948. Permskie mšanki s r. Sylva [Permian bryozoansfrom the river Sylva]. – In: „Mikrofauna neftânyh mestoro denyjSSSR“ [Microfauna of the oilfields of the SSSR], Paper 1, TrudyVNIGRI, New Series 31: 137–188, Leningrad (VNIGRI).

TSCHERNYSCHEW, T. & STEPANOW, P. 1916. Obercarbonfauna vonKönig Oscars und Heibergs Land. – Report of the Second Norwe-gian Arctic Expedition in the “Fram” 1898–1902, 34: 1–67, Oslo(Det Norske Videnskaps-Akademi i Kristiania [Oslo]).

ULRICH, E.O. 1882. American Paleozoic Bryozoa. – Journal of theCincinnati Society of Natural History 6: 245–279.

ULRICH, E.O. 1889. – In: MILLER, S.A. North American geology andpaleontology for the use of amateurs, students and scientists. –664 p., Cincinnati (Western Methodist Book Concern).

VINE, G.R. 1884. Fourth report of the Committee consisting of Dr. H.R.Sorby and Mr. G.R. Vine, appointed for the purpose of reportingon fossil Polyzoa. – Reports of the 53rd Meeting of the BritishAssociation for Advances in Sciences: 161–209.

Manuskripteingang / manuscript received 25. 1. 2006;Manuskriptannahme / manuscript accepted 15. 9. 2006.

Paleontologiceskogo

ogiceskogo


Paleontologicesko-

Paleontologiceskogo

Paleontologiceskogo

Niznekamennougol’nye



u c a

z

z

Lower Permian Bryozoa from Ellesmere Island (Canada

Documents

Transcript of Lower Permian Bryozoa from Ellesmere Island (Canada