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Sponge spicule assemblages from the Cambrian (Series 2–3) of North Greenland (Laurentia): systematics and biogeography

John S. Peel

To cite this article: John S. Peel (2019) Sponge spicule assemblages from the Cambrian (Series 2–3) of North Greenland (Laurentia): systematics and biogeography, GFF, 141:2, 133-161, DOI: 10.1080/11035897.2019.1578261 To link to this article: https://doi.org/10.1080/11035897.2019.1578261

© 2019 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group.

Published online: 03 Oct 2019.

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Full Terms & Conditions of access and use can be found at https://www.tandfonline.com/action/journalInformation?journalCode=sgff20 GFF 2019, VOL. 141, NO. 2, 133–161 https://doi.org/10.1080/11035897.2019.1578261

ARTICLE Sponge spicule assemblages from the Cambrian (Series 2–3) of North Greenland (Laurentia): systematics and biogeography John S. Peel

Department of Earth Sciences (Palaeobiology), Uppsala University, Uppsala, Sweden

ABSTRACT ARTICLE HISTORY Isolated microscopic spicules from disarticulated scleritomes demonstrate the presence of a diverse sponge Received 23 August 2018 fauna otherwise not evident from the macrofossil record in carbonate successions deposited during the un- Accepted 31 January 2019 named Cambrian Series 2 and Cambrian Series 3 (Miaolingian) in North Greenland. Most of the spicule KEYWORDS morphotypes are not recognised from faunas of articulated sponges known from contemporaneous Cambrian (Series 2–3); siliciclastic strata elsewhere. Assemblages are described in terms of four Cambrian stages and contain Miaolingian; sponge numerous spicule morphotypes not previously recorded from Laurentia. Many of the spicules can be spicules; systematics; correlated worldwide, however, extending current knowledge of the biogeographic distribution of sponge biogeography; North spicule-based taxa in the Cambrian. In particular, similar Cambrian Stage 4 (Series 2) and Miaolingian Greenland; Laurentia assemblages (Wuliuan, Drumian and Guzhangian stages) faunas are recorded from tropical palaeolatitudes in Australia, South China, Siberia and Laurentia, although this may in part reflect a methodology focused on the preparation of the carbonate samples that dominate these successions. New spicule-based taxa: Eiffelia floriformis n. sp., Australispongia? inuak n. sp., Celtispongia dorte n. gen. n. sp., Sanningasoqia borealis n. gen. n. sp., Speciosuspongia inughuitorum n. sp., Sulukispicula gelidae n. gen. n. sp.

Introduction This paper extends the studies by Peel (2017a, 2018)in reviewing sponge spicule assemblages from the lower–middle Cambrian sponges from Laurentia are best known from diverse Cambrian (Cambrian Series 2–3) of North Greenland. Spicule assemblages of articulated specimens in the siliciclastic strata of associations are grouped into four assemblages: Unnamed Stage lagerstätten such as the Burgess Shale and Sirius Passet (Rigby 4 from Cambrian Series 2, and the Wuliuan Stage, Drumian 1978, 1983, 1986; Rigby & Collins 2004; Rigby et al. 2010; Stage and Guzhangian Stage from the Miaolingian Series. In Botting & Peel 2016). Isolated sponge spicules occur widely addition to offering primary documentation of the diversity of but, with few exceptions, they have attracted little attention sponges within carbonate successions from northern Laurentia (Rigby 1975;Pratt2002;Skovsted2006; McMenamin 2008; (present day coordinates), the assemblages permit integration of Harvey 2010; Botting et al. 2015;Peeletal.2016). New records the palaeocontinent into the emerging picture of sponge palaeo- from Cambrian Series 2–3 carbonates of Laurentia (North geography during the Cambrian (Botting & Muir in press). Greenland) were documented by Peel (2017a) in describing Although previously not well known from Laurentia, spicules Silicunculus Bengtson, 1986, Australispongia Dong & Knoll, of disarticulated sponges are widely distributed and often abun- 1996 and Thoracospongia Mehl, 1996, three genera that were dant in other Cambrian palaeocontinents (e.g., Bengtson 1986; originally reported on the basis of sponge spicules from Fedorov in Fedorov & Pereladov 1987;Bengtsonetal.1990; Australia. The Greenland occurrences suggested that spicule- Zhang & Pratt 1994 ;Dong&Knoll1996; Panasenko 1998;Elicki based sponge taxa may have had palaeogeographic and biostra- 2011;Mehl1998; Vasilieva 1998;Beresi2003; Castellani et al. 2012; tigraphic utility in the Cambrian, in addition to demonstrating Kouchinsky et al. 2011, 2015; Sugai et al. 2004;Changetal.2017; a diversity of sponges not otherwise recognised within the Peel 2017a, 2018), showing a morphological diversity that con- relatively well known articulated lagerstätten sponge faunas. tinues to increase into the Ordovician (Webby & Trotter 1993; The notion of widespread distribution of Cambrian sponge Carrera & Maletz 2014). While the focus of systematic studies of spicules gained further support with the description by Peel Cambrian sponges naturally lies with articulated scleritomes, spi- (2018) of spicule assemblages from the Holm Dal Formation cules from acid residues provide a useful source of supplementary (Miaolingian [= Cambrian Series 3], Guzhangian Stage, information since they often represent a size sample, 250–500 µm, Lejopyge laevigata Biozone) of North Greenland which com- not readily discerned in articulated specimens. They also provide pared closely with an assemblage described by Bengtson details of fine surface structures often not preserved (Castellani (1986) from the contemporaneous Mungerebar Limestone et al. 2012)andthenatureoftheaxialfilament of the individual of Queensland, Australia. Although located on separate rays in silicean sponges, a character diagnostic of demosponge or palaeocontinents, both localities were placed near the equator hexactinellid affinity in crown groups (Botting & Muir 2013). In in the Miaolingian (Torsvik & Cocks 2017). present day demosponges, the axial filament is triangular to

CONTACT John S. Peel [email protected] Department of Earth Sciences (Palaeobiology), Uppsala University, Uppsala, Sweden © 2019 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group. This is an Open Access article distributed under the terms of the Creative Commons Attribution-NonCommercial-NoDerivatives License (http://creativecommons.org/licenses/by-nc-nd/4.0/), which permits non-commercial re-use, distribution, and reproduction in any medium, provided the original work is properly cited, and is not altered, transformed, or built upon in any way. Published online 03 Oct 2019 134 J. S. PEEL hexagonal in cross-section but it is square in hexactinellids. Geological background However, the observed canals in most Cambrian specimens are The geological evolution of the Greenland portion of the usually circular (Castellani et al. 2012;Peel2018) although this is Laurentian margin during the lower Palaeozoic was reviewed often a reflection of diagenetic widening of the filament cavity. by Higgins et al. (1991). Cambrian stratigraphy of North Botting & Muir (2013) noted that well-preserved axial filaments Greenland was formally described by Ineson & Peel (1997). are rare in Lower Palaeozoic sponges but increased knowledge of Siliciclastic sediments of the Buen Formation yield the only filament morphology may provide a significant contribution to known articulated sponges (Botting et al. 2015; Botting & Peel resolving the current uncertainty concerning the relationship of 2016; Peel & Willman 2018). Isolated sponge spicules have Cambrian sponges to crown group poriferans (Botting & been recovered from several formations within the overlying Butterfield 2005;Botting&Muir2013, 2018; Botting et al. 2013, carbonate-dominated succession (Figs. 1–3) which forms 2017; Carrera & Botting 2008). a northwards prograding sedimentary complex extending The stauractins, pentactins and hexactins that characterise from Cambrian Series 2 (Stage 3) to the lowest Ordovician. most spicule occurrences in the Cambrian show little mor- This complex is dominated by platform margin carbonate phological variation between localities and are of limited use slope-apron carbonates, deep shelf carbonates and siliciclastic in taxonomy. However, a number of “spicule-based” taxa sediments referred to the Brønlund Fjord and Tavsens Iskappe have been named under zoological nomenclature for mor- groups (Ineson & Peel 1997). Units of dark carbonates and phologically distinct spicules (Mostler 1985, 1996; Bengtson et shales (Fig. 2: Henson Gletscher, Ekspedition Bræ, Holm Dal al. 1990; Mehl 1996; Chen & Dong 2008; Peel 2017a, 2018), formations) are separated by cliff-forming successions of dolo- while Fedorov in Shabanov et al. (1987) and Fedorov in mitised carbonate turbidites and mass flow deposits Fedorov & Peredalov (1987) proposed several lower and (Aftenstjernesø, Sydpasset, Fimbuldal, Perssuaq Gletscher for- middle Cambrian genera on the basis of assemblages of dis- mations) that represent active progradation of the platform articulated spicules. The biological value of spicule-based taxa margin during periods of sea-level highstand (Ineson & Peel may be questioned, but it is unwise to dismiss their usefulness 1997). The dark carbonates and shales were deposited during if recognition increases insight into the diversity and distri- periods of lowstand (Ineson & Peel 1997) and many are richly bution of Cambrian sponges. Thus, the spicule-based taxon- fossiliferous (Peel 1988, 1994; Blaker & Peel 1997; Geyer & Peel omy employed by Peel (2017a, 2018) is maintained herein in 2011; Peel et al. 2016). the description of a number of morphologically distinct spi- Sponge spicules described herein are derived from the starved cules, with the aim of providing a clearer view of the dis- ramp succession of the basal Aftenstjernesø Formation, and the tribution of sponge assemblages in Cambrian carbonate laterally equivalent Kap Troedsson Formation of western sediments.

Figure 1. A–C, locality maps for sponges and sponge spicules from Cambrian Series 2 and Cambrian Series 3 (Miaolingian Series) in North Greenland. Note that locality 4 is indicated both in B and C to provide linkage. 1, Sirius Passet Lagerstätte, Transitional Buen Formation (Cambrian Series 2, Stage 3). 2, northern Freuchen Land, upper Buen Formation (Cambrian Series 2, Stage 4). 3, Navarana Fjord, east side, basal Aftenstjernesø Formation (Cambrian Series 2, Stage 4). 4, southern Freuchen Land, Henson Gletscher Formation (Cambrian Series 2, Stage 4); Ekspedition Bræ Fm. (Cambrian Series 3 [= Miaolingian], Drumian Stage). 5, southern Freuchen Land, Henson Gletscher Formation (Cambrian Series 2, Stage 4). 6, Freuchen Land, western side of glacier feeding Navarana Fjord, Holm Dal Formation (Cambrian Series 3 [= Miaolingian], Guzhangian Stage). 7, West side of Gustav Holm Dal, Fimbuldal Formation (Cambrian Series 3 [= Miaolingian], Drumian Stage). 8, Gustav Holm Dal, Holm Dal Formation (Cambrian Series 3 [= Miaolingian], Guzhangian Satge). 9, Gustav Holm Dal, east side, Aftenstjernesø Formation (Cambrian Series 2, Stage 4); Henson Gletscher Formation (Cambrian Series 3 [= Miaolingian], Wuliuan Stage); Holm Dal Formation (Cambrian Series 3 [= Miaolingian], Guzhangian Satge). 10, Fimbuldal, south side, Henson Gletscher Formation (Cambrian Series 3 [= Miaolingian], Wuliuan Stage). 11, southern Lauge Koch Land, Aftenstjernesø Formation (Cambrian Series 2, Stage 4). 12, west side of Løndal, Henson Gletscher Formation (Cambrian Series 2, uppermost Stage 4). 13, southern Wulff Land, Kap Troedsson Formation (Cambrian Series 2, Stage 4). 14, south-east Wulff Land, Kap Troedsson Formation (Cambrian Series 2, Stage 4). 15, land area south of Nares Land, Kap Troedsson Formation (Cambrian Series 2, Stage 4). Børglum Elv, Buen and Aftenstjernesø formations (Cambrian Series 2, Stage 4). Northern Nyeboe Land, Aftenstjernesø Formation (Cambrian Series 2, Stage 4). GFF 135

Holm Dal, western Peary Land (82°21.5ʹN, 39° 44ʹW; Fig. 1C, locality 8). GGU sample 271471 was collected by J.S. Peel on 25 June 1978, basal member of Aftenstjernesø Formation, south- ern Lauge Koch Land, (82°10ʹN, 40°24ʹW; Fig. 1C, locality 11). GGU sample 315045 was collected by J.S. Peel on 7 July 1984 from the base of the formation on the east side of Navarana Fjord, Lauge Koch Land, on the northern limb of anticline (82°36.5ʹN, 41°15ʹW; Fig. 1C, locality 3). GGU samples 319740 and 319742 were collected by A.K. Higgins on 11 August 1985 from the Aftenstjernesø Formation in northern Nyeboe Land (Fig. 1A), probably at Kap Bryant (approximately 82°19ʹN, 55°27ʹW). GGU sample 319748 was collected by A.K. Higgins on 11 August 1985, south of Kap Bryant, northern Nyeboe Land (82°18.5ʹN, 55°17ʹW), about 8 m above the base of the formation (Higgins et al. 1992, section 25). Kap Troedsson Formation. – All spicules from the Kap Troedsson Formation (Fig. 4) are derived from samples from Cambrian Series 2, Stage 4. GGU samples 270505 and 270506 were collected by J.S. Peel on 23 July 1978 on the west side of the lake in south-east Wulff Land (81°34ʹN, 47°3.6ʹW; Fig. 1B, locality 14). GGU sample 298894 was collected by J.S. Peel and M.R. Blaker on ff – Figure 2. Stratigraphy of Cambrian shelf deposits in the Wul Land Peary Land 3 August 1985 in southern Wulff Land (81°24.5ʹ N, 47°54.5ʹ W; region of North Greenland. Fig. 1B,locality13),about25mabovethebaseoftheformation. GGU sample 315049 was collected by J.S. Peel on 11 July 1984, outcrops, from the lowstand shelf successions of the Henson on the north side of Harder Gletscher, in the land area south of Gletscher and Ekspedition Bræ formations, and from a medial Nares Land (81°10.5ʹN, 44°36.5ʹW; Fig. 1B,locality15). limestone development within the dolomites of the Fimbuldal Henson Gletscher Formation. – Spicules from the Henson Formation (Ineson & Peel 1997). Spicule assemblages from the Gletscher Formation (Cambrian Series 2, Stage 4; Fig. 5) overlying lowstand deposits of the Holm Dal Formation (Series were derived from samples from the lower and upper mem- 3, Guzhangian) were described by Peel (2018). bers of the formation. Spicules from the uppermost part of Methods. – Spicules were recovered during a programme of the upper member (Figs. 6, 7) are from the Miaolingian routine processing of limestone samples in 10% acetic acid. Series, Wuliuan Stage. They were hand picked from sieved residues (250 µm fraction GGU sample 218614 was collected by J.R. Ineson on and larger) prior to study by scanning electron microscopy. 2 July 1979 on the south side of Fimbuldal, westernmost Figures were assembled using Adobe Photoshop CS4. Peary Land (82°17.7ʹN, 39°42ʹW; Fig. 1C, locality 10), from 0.5 m below the top of Henson Gletscher Formation Institutional abbreviations. – GGU: Grønlands Geologiske (Miaolingian Series, Wuliuan Stage). This is locality 4 of Undersøgelse [Geological Survey of Greenland], now a part Geyer & Peel (2011, fig. 1D). of the Geological Survey of Denmark and Greenland (GEUS), GGU samples 225707, 225708, 2235713 and 225714 were Copenhagen, Denmark. PMU: palaeontological type collec- collected by J.S. Peel on 15 July 1979 from the upper Henson tion of the Museum of Evolution, Uppsala University, Gletscher Formation on the west side of Løndal, Peary Land Uppsala, Sweden. (82°17 · 5ʹN, 37°03ʹW; Fig. 1C, locality 12) from strata yield- ing the Ovatoryctocara granulata assemblage (Cambrian Series 2, uppermost Stage 4; Geyer & Peel 2011, fig. 1D; Derivation of samples Peel et al. 2016, figs. 1, 2). GGU sample 271428 was collected by J.S. Peel on The GGU samples that have yielded described and illustrated 20 June 1978 from about 4 m below the top of the formation spicules were collected from localities and horizons identified (thickness 32 m), on the east side of Gustav Holm Dal at its in Figs. 1–3. Spicules also occur in many unlisted collections, junction with Fimbuldal, western Peary Land (82°19.5ʹN, 39° especially within the Henson Gletscher Formation (Geyer & 37ʹW; Fig. 1C, locality 9). Miaolingian Series, Wuliuan Stage, Peel 2011). Ptychagnostus gibbus Biozone (Robison 1984, p. 6). Aftenstjernesø Formation. – Spicules from the Aftenstjernesø GGU samples 298544 and 298545 were collected by J.S. Peel Formation (Fig. 4) are derived from samples from Cambrian on 15 July 1985 from 16 m above the base of the lower member Series 2, Stage 4. (Cambrian Series 2, Stage 4) of the Henson Gletscher Formation GGU sample 218894 was collected by P. Frykman on on a nunatak in southern Freuchen Land (82°09ʹN, 42°25ʹW; 10 July 1979 from the basal member of the formation, Gustav Fig. 1B,C,locality4). 136 J. S. PEEL

the Henson Gletscher Formation (Cambrian Series 2, Stage 4) on a nunatak in southern Freuchen Land (82°09ʹN, 42°25ʹW; Fig. 1B, C locality 4). Ekspedition Bræ Formation. – All spicules from the Ekspedition Bræ Formation (Fig. 8) are from the Miaolingian Series, Drumian Stage. GGU sample 218832 was collected by P. Frykman and J.R. Ineson in 1979 from 68.5 m above the base of the Ekspedition Bræ Formation in its type section (total thickness 82 m; Ineson & Peel 1997, figs. 41, 42) on the western side of Henson Gletscher in southern Lauge Koch Land (82°10ʹN, 40°24ʹW; Fig. 1C, locality 11). GGU sample 271493 was collected by J.S. Peel on 26 June 1978 from about 35 m above the base of the Ekspedition Bræ Formation in its type section in Lauge Koch Land (82°10ʹ N, 40°24ʹ W; Fig. 1C, locality 11). GGU sample 315117 was collected by J.S. Peel on 22 July 1984 from the base of the Ekspedition Bræ Formation on a nunatak in southern Freuchen Land (82°09ʹN, 42°25ʹW; Fig. 1B,C, locality 4). Fimbuldal Formation. – Illustrated spicules from the Fimbuldal Formation (Fig. 9) are derived from the Miaolingian Series, Drumian Stage, Ptychagnostus punctuosus Biozone (Robison 1984). GGU sample 218626 was collected by J.R. Ineson on 1 July 1979 from the south side of Fimbuldal, western Peary Land from the base of the recessive middle unit of the Fimbuldal Formation (82°17.5ʹN, 39°41ʹW; Fig. 1C,locality10). GGU samples 218644 and 218645 were collected by J.R. Ineson on 7 July 1979 from the west side of Gustav Holm Dal, western Peary Land (82°20.3ʹN, 39°45ʹ W; Fig. 1C, locality 7) from just above the middle of a unit of dark, bituminous limestones forming the recessive middle unit of the Fimbuldal Formation in its type section (Ineson & Peel 1997, figs. 44, 45). Holm Dal Formation. – All spicules from the Holm Dal Formation (Fig. 10) are from samples from the Miaolingian Series, Guzhangian Stage, upper Lejopyge laevigata Biozone Figure 3. Distribution of named sponge spicules in Cambrian Series 2 and (Robison 1988; Peel 2018). Miaolingian (Series 3), North Greenland. Spicules from the Kap Troedsson Formation are grouped together with those of the contemporaneous GGU sample 225540 was collected by J.S. Peel on 7 July 1979 Aftenstjernesø Formation. Spicules from Cambrian Series 2, Stage 4, from the from 55 m above the base of the Holm Dal Formation (thickness lower and upper members the Henson Gletscher, are grouped together (1 + 2), 155 m) in its type section (Ineson 1988, Fig. 6; Robison 1988, Fig. while Miaolingian Series spicules from the upper member of the formation are grouped as (3). 2; Peel 2018, Fig. 1) on the east side of Gustav Holm Dal at its junction with Fimbuldal, western Peary Land (82°19.5ʹN, 39° 37ʹW; Fig. 1C,locality9). GGU samples 301338 and 301351 were collected by M.R. GGU samples 225592, 225594, 225595 were collected by J.S. Blaker and J.S. Peel on 15th–17 August 1985 from about 15 m Peel on 13 July 1979 from the Holm Dal Formation in Gustav ʹ ʹ above the base of the lower member of the Henson Gletscher Holm Dal, western Peary Land (82°22 N, 39°41 W; Fig. 1C,local- fi Formation (Cambrian Series 2, Stage 4) on a nunatak in south- ity 8). This is locality 3 of Robison (1988, g.1B,5),seealsoPeel fi ern Freuchen Land (82°09ʹN, 42°25ʹW; Fig. 1B,C,locality4). (2018, g. 1). GGU sample 225592 is from near the middle of an GGU sample 301352 was collected by J.S. Peel on approximately 100 m thick section (and probably about the same 17 August 1985 from the lower member of the Henson stratigraphic level as GGU sample 225540) but GGU samples Gletscher Formation (Cambrian Series 2, Stage 4) at the south- 225594 and 225595 are from the upper part of the section. ern end of a nunatak at the north-east end of the glacier feeding GGU sample 315008 was collected by J.S. Peel on south-west into Jungersen Gletscher (82°10ʹN, 42°4ʹW; Fig. 1C, 28 June 1984 from the Holm Dal Formation on the wes- locality 5). tern side of the glacier feeding into Navarana Fjord (Peel fi ʹ GGU sample 315091 was collected by J.S. Peel on 2018, g. 1), south-east Freuchen Land (82°16.7 N, 41° ʹ 19 July 1984 from 12m above the base of the lower member of 22 W; Fig. 1C,locality6). GFF 137

Figure 4. Sponge spicules from the Aftenstjernesø and Kap Troedsson formations (Cambrian Series 2, Stage 4). A, J, Microcoryne sp., PMU 34329 from GGU sample 270506, Kap Troedsson Formation (locality 14). Arrow in J indicates axial cavity. B, C, H, I, K–M, “Calcihexactina” sp. B, pentactin, PMU 34330 from GGU sample 271471, Aftenstjernesø Formation (locality 11). C, pentactin, PMU 34331 from GGU sample 319740, Aftenstjernesø Formation, northern Nyeboe Land. H, irregular hexactin, PMU 34332 from GGU sample 315045, Aftenstjernesø Formation (locality 3). I, pentactin, PMU 34333 from GGU sample 270505, Kap Troedsson Formation (locality 14). K, pentactin, PMU 34334 from GGU sample 315045, Aftenstjernesø Formation (locality 3). L, pentactin, PMU 34335 from GGU sample 298894, Aftenstjernesø Formation (locality 8). M, pentactin, PMU 34336 from GGU sample 298894, Aftenstjernesø Formation (locality 8). D, G, irregular polyactins. D, PMU 34337 from GGU sample 270506, Kap Troedsson Formation (locality 14). G, PMU 34338 from GGU sample 218894, Aftenstjernesø Formation (locality 8). E, F, Eiffelia sp., worn spicules. E, PMU 34339 from GGU sample 315049, Kap Troedsson Formation (locality 15). F, Eiffelia sp., PMU 34340 from GGU sample 218894, Aftenstjernesø Formation (locality 8). Scale bars 100 µm, except E (500 µm), F (2 mm).

Preservation sponges are demonstrating the importance of Cambrian fossil sponges to the understanding of early sponge evolution While investigation of composition has not been addressed (Pisera 2003, 2006; Botting & Muir 2018). Studies of isolated systematically, most spicules appear to be preserved in silica. sponge spicules may contribute valuable information, but As such, they are resistant to the sample preparation in weak such discussion lies beyond the context of the simple spicule- acetic acid but this resistance in its present state may well be the based taxonomy employed here. Spicules are described with end point of significant diagenetic changes in composition little inference concerning their biological or evolutionary (Botting & Muir 2018) Spicules vary in colouration from trans- relationships, against the background of the phylogenetic parent, through dominant translucent white to matt brown framework reviewed by Botting & Muir (2018, fig. 6). silica; several colours may occur in the same sample and within similar spicules in the same sample, or even within individual spicules. Coatings of microcrystalline silica or pyrite are wide- ?CALCAREA BOWERBANK, 1864 spread. The phosphatised spicules that dominate GGU sample Genus Dodecaactinella Reif, 1968 218832 (Fig. 8J, L–N) obviously have had a complex diagenetic – history on account of their association with the phosphatisation Type species. Dodecaactinella oncera Reif, 1968 from the of the associated hardground. Evidence of wearing due to rede- Ordovician of Estonia. position or corrosion is also common place (Fig. 4E, F) but Discussion. – Polyactinellids were regarded as Calcarea by fl simple breakage may in part re ect preparation techniques. Debrenne & Reitner (2001) but they are unlike other mem- Axial cavities are visible in many spicules but their relatively bers of the group (Botting & Muir 2018); their position is large diameter indicates diagenetic solution or partial recrystal- uncertain. lisation of the spicules (Figs. 5C, 6C) while an outer crust may As a prelude to describing Dodecaactinella cynodontota cover a dissolved interior (Figs. 6E, Z, 9E, J). General degrada- Bengtson & Runnegar in Bengtson et al. 1990,fromthe tion, etching or incomplete recrystallisation of the rays occurs lower Cambrian Ajax Limestone of South Australia, frequently in some samples (Figs. 4H, 5B, 8I) but most are well Bengtson et al. (1990) synonymised Polyactinella preserved. Mostler, 1985 and Sardospongia Mostler, 1985 with Dodecaactinella Reif, 1968, but this synonymy was rejected by Mostler (1996). Mostler (1996) mistakenly Systematic palaeontology identified as the holotype of Dodecaactinella cynodontota, a specimen (Bengtson et al. 1990, fig. 11A) which he PHYLUM PORIFERA GRANT, 1836 subsequently transferred to Dodecaactinella oncera Reif, Discussion. – New insights concerning the mineralogical com- 1968,thetypespeciesofDodecaactinella.Mostler(1996) position, morphology and geological distribution of fossil then designated the second specimen of Dodecaactinella 138 J. S. PEEL

Figure 5. Sponge spicules from the Henson Gletscher Formation (Cambrian Series 2, Stage 4). A, J, K, O, R, Eiffelia floriformis n. sp. A, stauractin, PMU 343412 from GGU sample 298545 (locality 4). J, K, hexactin, holotype, PMU 28801 from GGU sample 225713 (locality 12). O, hexactin, PMU 34342 from GGU sample 225708 (locality 12). R, hexactin, PMU 34343 from GGU sample 225707 (locality 12). B, D, H, I, P, Cjulanciella asimmetrica Fedorov in Fedorov & Peredalov (1987). B, PMU 34344 from GGU sample 301338 (locality 4). D, PMU 34345 from GGU sample 315091 (locality 4). H, PMU 34346 from GGU sample 301351 (locality 4). I, PMU 34347 from GGU sample 301352 (locality 5). P, PMU 34348 from GGU sample 315091 (locality 4). E–G, Dodecaactinella oncera Reif (1968), with arrows indicating accessory rays. E, PMU 34349 from GGU sample 298544 (locality 4). F, PMU 28804 from GGU sample 225713 (locality 12). G, PMU 34350 from GGU sample 298544 (locality 4). C, hexactin, PMU 34351 from GGU sample 315091(locality 4). L,M , Eiffelia sp., from GGU sample 225714 (locality 12). L, PMU 28802. M, PMU 28803. N, stauractin, PMU 28805 from GGU sample 225714 (locality 12). Q, polyactin, PMU 34352 from GGU sample 315091 (locality 4). S, stauractin, PMU 34353 from GGU sample 301352 (locality 4). T, club-like polyactin, PMU 28811 from GGU sample 225708 (locality 12). U, pentactin, PMU 34354 from GGU sample 301352 (locality 4). V, polyactin, PMU 34355 from GGU sample 301352 (locality 4). W, fragment of ray, PMU 34356 from GGU sample 218614 (locality 10). Scale bars 100 µm.

cynodontota illustrated by Bengtson et al. (1990, fig. 11B– junior synonym of cynodontota Bengtson & Runnegar in E) as holotype of the type species of his new genus Bengtson et al. 1990. Bengtsonella. However, the specimen designated by Özdikmen (2009) noted that Bengtsonella Mostler, 1996 Mostler (1996) as the holotype of Bengtsonella australien- is a junior homonym of Bengtsonella Müller & Hinz, 1991 sis n. gen. n. sp. is the true holotype of Dodecaactinella and proposed Mostlerhella nom. nov. as a replacement cynodontota. Since these two nominal species-group taxa name for Mostler’s(1996) genus. The type species is have the same name-bearing type, their names are objec- Mostlerhella australiensis (Mostler 1996), but this is tive synonyms (ICZN article 61–3–4). Thus, the species- ajuniorobjectivesynonymofMostlerhella cynodontota group taxon australiensis Mostler, 1996 is an objective (Bengtson & Runnegar in Bengtson et al. 1990).

Figure 6. Sponge spicules from the Henson Gletscher Formation (Miaolingian Series, Wuliuan Stage), GGU sample 271428 (locality 9). A, polyactin, PMU 34357. B, K, Nabaviella? sp. B, PMU 34358. K, PMU 34359. C, polyactin, PMU 34360. D, E, Cjulanciella asimmetrica Fedorov in Fedorov & Peredalov (1987). D, pentactin, PMU 34361. E, stauractin, PMU 34362. F, G, R, Sanningasoqia borealis n. gen. n. sp. F, stauractin with rays curved within plane of whorls, and axial canal arrowed PMU 34363. G, R, stauractin, holotype, PMU 34364. H–J, pentactins. H, PMU 34365. I, PMU 34366. j, PMU 34367. L, Thoracospongia sp., pentactin, PMU 34368. M, stauractin with reduced ray, PMU 34369. N, stauractin, PMU 34370. O–Q, Celtispongia dorte n. gen. n. sp. O, P, holotype, PMU 34371. Q, PMU 34372. S, hexactin, with axial canal arrowed, PMU 34373. T–V, Z, AA, BB, EE, Thoracospongia lacrimiformis Peel, 2017. T, Y, EE, two conjoined pentactins with minute paratangential rays, holotype, PMU 29990. V, two conjoined pentactins, but paratangential rays broken away, PMU 29992. Z, PMU 34374. AA, PMU 29993. BB, PMU 29991. W, Kuonamia fusiformis (Fedorov in Fedorov & Peredalov 1987), pentactin, PMU 34375. X, Y, CC, DD, Australispongia? inuak n. sp. X, pentactin with one pair of tangential rays reduced, PMU 34376. Y, DD, pentactin, PMU 34377, holotype. CC, pentactin with one pair of tangential rays reduced, PMU 34378. Scale bars 100 µm. GFF 139 140 J. S. PEEL

Figure 7. Sponge spicules from the Henson Gletscher Formation (Miaolingian Series, Wuliuan Stage), GGU sample 271428 (locality 9). A–C, G, I–K, N, R, Speciosuspongia spp., spiderpentactins. A, PMU 34379. B, J, PMU 34380. C, PMU 34381, with axial canals exposed (arrow). E, Q, PMU 34382. F, PMU 34383. G, PMU 34384. I, PMU 34385. K, PMU 34386, stauractin. N, PMU 34387. O, PMU 34388, with axial canal arrowed. R, PMU 34389, with axial canal arrowed. D, L,M, P, Speciosuspongia inughuitorum n. sp., spiderpentactins. D, PMU 34390. L,M , PMU 34391, holotype, with obscure longitudinal ribs on axial ray (arrowed). P, PMU 34392, paratype. H, Abnormisella insperata Fedorov in Fedorov & Peredalov (1987), spiderpentactin, PMU 34393. S, T, W, Z, stauractins. S, stauractin, PMU 34394. T, stauractin with curved rays, PMU 34395. W, stauractin with arrowed axial canal, PMU 34396. Z, irregular stauractin, PMU 34397. V, X, U?, Kuonamia fusiformis (Fedorov in Fedorov & Peredalov 1987). V, hexactin, PMU 34398. X, hexactin, PMU 34399. U, stauractin tentatively assigned, PMU 34400. Y, Cjulanciella asimmetrica Fedorov in Fedorov & Peredalov (1987) PMU 34437, stauractin. AA, Australispongia? inuak n. sp., PMU 34401, stauractin, BB, diactin, PMU 34402. Scale bars 100 µm, except Q (50 µm) and U, W (200 µm). GFF 141

Figure 8. Sponge spicules from the Ekspedition Bræ Formation (Miaolingian Series, Drumian Stage). GGU sample 315117 (locality 4) unless stated. A, pentactin, PMU 34403. B–D, G, H, K, Sulukispicula gelidae n. gen. n. sp., showing blade-like extensions from the four rays (arrows). B, PMU 34404. C, PMU 34405. D, PMU 34406, holotype, oblique view. G, PMU 34407. H, PMU 34408. K, PMU 34409. E, F, I, O, acanthose pentactins. E, PMU 34410. F, PMU 34411. I, O, PMU 34412 from GGU sample 271493 showing concentric structure of ray (locality 11). J, L–N, phosphatised hexactin spicules within re-deposited, phosphate coated, clasts, GGU sample 218832 (locality 11). J, PMU 34413. L, PMU 34414.M, PMU 34415. N, PMU 34416. Scale bars: 100 µm.

Seemingly unaware of Mostler (1996), Mehl & Lehnert (1997) 1985 Polyactinella furcata Mostler, p. 15–16, fig. 3; pl. 1. fig. 5; considered Polyactinella to be a synonym of Dodecaactinella but pl. 2, figs. 1, 2, 7. recognised Sardospongia as a separate genus. They transferred the 1987 Demospongia, forma 2 Fedorov in Shabanov et al., holotype of Dodecaactinella cynodontota (Bengtson et al. 1990, fig. p. 129, pl. 34, figs. 2–5. 11B–E) to Sardospongia but synonymised the original of 1990 Dodecaactinella cynodontota Bengtson & Runnegar in the second illustrated specimen of Dodecaactinella cynodontota Bengtson et al., fig. 11A. (Bengtson et al. 1990, fig.11A)withthetypespecies, 1996 Dodecaactinella cynodontota (oncera), Mostler, pl. 1, Dodecaactinella oncera Reif, 1968. Following Mehl & Lehnert fig. 6. (1997), Dodecaactinella is characterised by three spine-like acces- 1997 Dodecaactinella furcata, Mehl & Lehnert, p. 231, sory rays which are interspersed between the primary triactine fig. 4A–C, fig. 5, D–J. rays whereas Sardospongia lacks these accessory rays. Both 2004 Dodecaactinella cf. D. oncera; Sugai et al., pl. 1, fig. 1, morphologies were illustrated by Fedorov in Shabanov et al. pl. 2, fig. 1. (1987) from the lower Cambrian (Atdabanian Stage) of the 2016 Dodecaactinella sp., Peel, fig. 7F. Anabar uplift, Siberia. According to Mostler (1996), the primary Figured material. – PMU 28804 from GGU sample 225713, triactine rays bifurcate in Sardospongia but divide into three Løndal (Fig. 1C, locality 12), PMU 34349 and 34350 from GGU secondary rays in Mostlerhella (= Bengtsonella). Thus, each mem- sample 298544, southern Freuchen Land (Fig. 1C, locality 4); ber of the two pairs of genera (Dodecaactinella and Polyactinella; Henson Gletscher Formation (Cambrian Series 2, Stage 4). Sardospongia and Mostlerhella) is delimited from its partner by the presence of distal bifurcation or trifurcation of the primary three rays. Following Mostler & Lehnert (1997), this distinction is fi not considered generically signi cant. Polyactinella and Discussion. – Dodecaactinella oncera is common in GGU Mostlerhella are considered to be junior synonyms of sample 298544 from the Henson Gletscher Formation Dodecaactinella and Sardospongia, respectively. (Cambrian Series 2, Stage 4) in southern Freuchen Land (Fig. 1C, locality 4) and seems to be restricted to the upper part of the lower member. Assignment to Dodecaactinella Dodecaactinella oncera Reif, 1968 reflects the presence of the spine-like accessory rays (arrows Fig. 5E–G in Fig. 5E, G). The three main rays are co-planar and each 1968 Dodecaactinella oncera Reif, p. 741, pl. 3, figs, 12, 13. divides into three secondary rays distally (Fig. 5E) which 142 J. S. PEEL

Figure 9. Sponge spicules from the Fimbuldal Formation (Miaoligian Series, Drumian Stage), GGU sample 218644 (locality 7) unless stated. A, B, G, H, Speciosuspongia hunanensis Chen & Dong (2008), spiderpentactins. A, PMU 34417 from GGU sample 218626 (locality 10). B, PMU 34418 from GGU sample 218626 (locality 10). G, PMU 34419. H, PMU 34420 from GGU sample 218626 (locality 10). C, D, I, J, P, Speciosuspongia spp., acanthose spiderpentactins. C, D, PMU 34421. I, PMU 34422 from GGU sample 218626 (locality 10). J, PMU 34423. P, PMU 34424. E, Australispongia sinensis Dong & Knoll (1996), PMU 29994 from GGU sample 218645 (locality 7), pentactine. F, pentactin, PMU 34425. K, acanthose pentactine, PMU 34426 from GGU sample 218626 (locality 10). L, tauactin, PMU 34427. M, stauractin, PMU 34428. N, R–T, Sanningasoqia borealis n. gen. n. sp., stauractins. N, PMU 34429 from GGU sample 21826 (locality 10). R, T, PMU 34430, with detail of axial canal (arrow). S, convex side of spicule, PMU 34431 from GGU sample 218626 (locality 10). O, stauractin, PMU 34432 from GGU sample 218626 (locality 10). Q, X, broken pentactin with detail of axial canal (arrow), PMU 34433 from GGU sample 218626 (locality 10). U, Y, acanthose pentactin with detail of axial canal (arrow), PMU 34434. v, modified stauractin, PMU 34435 from GGU sample 218626. W, Z, Abnormisella insperata Fedorov in Fedorov & Peredalov (1987), spiderpentactin with detail of axial canal (arrow), PMU 34436. Scale bars: 100 µm except X,Y (20 µm) and Z (10 µm). GFF 143

Figure 10. Sponge spicules from the Holm Dal Formation (Miaolingian Series, Guzhangian Stage). A, H, Silicunculus australiensis Bengtson (1986). A, PMU 29996 from GGU sample 225540 (locality 9). H, PMU 30003 from GGU sample 225595, detail of acanthose paratangential rays (locality 8). B, C, Silicunculus saaqqutit Peel, 2017. PMU 29999 from GGU sample 225595 (locality 8), holotype, with detail of acanthose paratangential rays (C). D, dichodactin, PMU 31803 from GGU sample 225592 (locality 8). E, diactin with secondary ray arrowed, PMU 31808 from GGU sample 225592 (locality 8). F, hexactin, PMU 31804 from GGU sample 225540 (locality 9). G, Australispongia sinensis Dong & Knoll, 1996, PMU 31818 from GGU sample 225592 (locality 8). I, diactins, PMU 31812 and PMU 31813 from GGU sample 225592 (locality 8). J, Kuonamia fusiformis (Fedorov in Fedorov & Peredalov 1987), stauractin, PMU 31843 from GGU sample 315008 (locality 6). K, Z, Abnormisella insperata Fedorov in Fedorov & Peredalov (1987), spiderpentactins. K, PMU 31819 from GGU sample 225594 (locality 8). Z, PMU 31820 from GGU sample 315008 (locality 6). L, Australispongia sp., PMU 31817 from GGU sample 225594 (locality 8).M, Q, Sanningasoqia borealis n. gen. n. sp., stauractins.M, PMU 31818 from GGU sample 225592 (locality 8). Q, PMU 31814 from GGU sample 315008 (locality 6). N, O, Tallitaniqa petalliformis Peel (2018), stauractins, PMU 31841 from GGU sample 318008 (locality 6). P, Sisamatispongia erecta Peel (2018), paratype, PMU 31846 from GGU sample 315008 (locality 6). R, acanthose pentactin, PMU 31811 from GGU sample 225595 (locality 8). S, pentactin, PMU 31838 from GGU sample 315008 (locality 6). T, partially recrystallised acanthose pentactin with infilled axial cavity (arrow), PMU 31815 from 225 (locality 8). U, V, Seqineqia bottingi Peel (2018), PMU 31821 from GGU sample 225540 (locality 9), holotype. W, acanthose pentactin, PMU 31827 from GGU 225594 (locality 8). X, triactin, PMU 31807 from GGU sample 225540 (locality 9). Y, stauractin, PMU 31832 from GGU sample 315008 (locality 6). Scale bars: 100 µm except U (30 µm). would also confirm this assignment even in the more Dodecaactinella cynodontota (= D. oncera) Bengtson & restricted definition of Mostler (1985, 1996). The secondary Runnegar in Bengtson et al. 1990, from the lower Cambrian rays lie within a plane approximately perpendicular to the Ajax Limestone of South Australia, but not in the holotype of axis of the primary rays. The same pattern is seen in one that species (Bengtson et al. 1990, fig. 11B, C; = Sardospongia illustrated specimen (Bengtson et al. 1990, Fig. 11A)of cynodontota). 144 J. S. PEEL

The irregular, hackly, preservation interpreted by Bengtson Derivation of name. – From the flower-shaped form of the et al. (1990)asareflection of original calcareous composition is spicule. also evident in the specimens from the Henson Gletscher Holotype. – PMU 28801 from GGU sample 225713, Henson Formation (Fig. 5E–G) and other material illustrated by Mostler Gletscher Formation (Cambrian Series 2, Stage 4), Løndal, (1985, 1996), Fedorov in Shabanov et al. (1987) and Dong & Peary Land, North Greenland (Fig. 1C, locality 12). Knoll (1996). Sugai et al. (2004)illustratedDodecaactinella cf. Other figured material. – PMU 28800 from GGU sample D. oncera from the Bagrad and Shashkunar formations 225707, paratype, PMU 34342 from GGU sample 225708, (Cambrian Series 2, Stage 4, Botoman) of Khakasia (south- Løndal (Fig. 1C, locality 12), PMU 34341 from GGU sample ern Siberia) of equivalent age to the North Greenland 298545, southern Freuchen Land (Fig. 1C, locality 4); Henson occurrence. Zhuravlev & Leguta (2005,pl.14,fig. 4) Gletscher Formation (Cambrian Series 2, Stage 4), North described Dodecaactinella sp. from the Sinsk Biota Greenland. (Cambrian Stage 4) of Yakutia, eastern Siberia, but the Diagnosis. – Species of Eiffelia with four to six coplanar rays lack of accessory rays in the illustrated specimen suggests and with about 7–13 small stubby secondary rays radiating assignment to Sardospongia. from the axial area. Skovsted (2006, fig. 7.25–27) illustrated fragments assigned to Dodecaactinella sp. from the Ella Island Formation Description. – Sclerite with four to six coplanar rays. (Cambrian Series 2, Stage 4) of North-East Greenland, but the Individual rays taper uniformly from the axial area, but fragments have parallel rays and more closely resemble become more pointed at their distal termination (Fig. 5A). Phobetractinia Reif, 1968.Elicki(2011) illustrated a similar frag- One central axial surface is smooth, and slightly raised, ment from the Jordan Rift Valley (Cambrian Series 3) and but the opposing surface of hexactin spicules carries about rejected the assignment of the specimens from the Bastion 10–13 short, stubby, radiating secondary rays, seemingly Formation to Dodecaactinella, and another described from the with rounded distal terminations. Rays in the only known Kinzers Formation of Pennsylvania by Skovsted & Peel (2010, stauractin spicule are not perfectly perpendicular to each fig. 3.8), suggesting affiliation to Praephobetractinia Kozur, 1991 other and only seven short secondary rays are present (Kozur 1991;Mostler1996). Wrona (2004, fig. 5A, B) illustrated (Fig. 5A). two specimens from lower Cambrian erratics in Antarctica to Discussion. – Hexactin and stauractin spicules from the Dodecaactinella but the first of these seems to be Phobetractinia. Emyaksin Formation (Cambrian Series 2, Stage 4, Botoman) of northern Siberia referred to Heteractinida indet. by Kouchinsky et al. (2015, fig. 72A, C–E, G, H) are here “Stem Calcarean” assigned to Effelia floriformis n. sp. They are associated with Genus Eiffelia Walcott, 1920 several polyactinous rays which may be part of the same Type species. – Eiffelia globosa Walcott, 1920 from the Burgess skeleton but have not been recognised in the sparse material Shale (Miaolingian) of British Columbia. from North Greenland. Bengtson in Bengtson et al. (1990) illustrated hexactin speci- Discussion.– Botting & Butterfield (2005) considered Eiffelia mens from the early Cambrian of South Australia assigned to to be a stem hexactinellid, intermediate between silicispon- Eiffelia araniformis (Missarzhevsky & Mambetov 1981)insomeof geans and calcareans, but Botting & Muir (2018) interpreted which numerous small tubercles on the axial region were distrib- it as stem group Calcarea + Homoscleromorpha. They noted uted around a short axial ray. A similar array of spicule types from the occurrence of hexactin-based four-rayed spicules in speci- the middle Cambrian of the Georgina Basin (Miaolingian Series, mens of Eiffelia from the Burgess Shale in addition to the Wuliuan Stage) was assigned to Eiffelia sp. by Mehl (1998). Effelia more common hexactins, and both spicule forms are also floriformis n. sp. differs from these in having fewer more promi- recognised within Eiffelia floriformis n. sp. from North nent secondary rays without an axial ray. Greenland. Bengtson in Bengtson et al. (1990) gave an exten- Skovsted (2006, figs. 7.23–24) recorded Eifellia araniformis sive synonymy of Eiffelia. from the Bastion Formation of North-East Greenland Spicules with six relatively robust rays from the (Cambrian Series 2, Stage 4) but the hexactin spicules have Aftenstjernesø and Kap Troedsson formations (Fig. 4E, F) less tapering rays than Eiffelia floriformis and lack secondary and forms from the Henson Gletscher Formation (Fig. rays on the axial area. 5L, M) with more slender, less tapering rays and no secondary rays, are referred to Eiffelia sp. SILICEA Gray, 1867 Eiffelia Floriformis n. sp. Discussion. – Many of the spicules described alphabetically Fig. 5A,J,K,O,R below might be regarded as hexactinellids but hexactin spicules occur in various groups of early sponges (Botting & Butterfield 2016 Heteractinellid sp., Peel, p. 249, fig. 7AC 2005;Harvey2010; Botting & Muir 2013, 2018). Botting & Muir LSID. – urn:lsid:zoobank.org:act:77E98C2F-031B-4A48-8F17- (2018) urged that most hexactinellid-like forms should be 1FC60DDF8D14 regarded as representatives of possible stem groups. GFF 145

Genus Abnormisella Fedorov in Fedorov & Peredalov, that Abnormisella and Speciosuspongia may be closely 1987 related. Hu et al. (2002, pl. 2, figs. 8, 9) illustrated poorly preserved Type species. – Abnormisella insperata Fedorov in Fedorov & spicules from the lower Cambrian Hetang Formation of Peredalov, 1987 from the Kuonamka Formation (Miaolingian Anhui Province, South China, which may represent the oldest Series, Wuliuan Stage, Kuonamkites Biozone) of north- described examples of the “spiderpentactin” morphology. eastern Siberia. Diagnosis. – Spiderpentactin, bilaterally symmetrical about a plane longitudinally bisecting the axial ray and the angle Abnormisella Insperata Fedorov in Fedorov & Peredalov between each pair of paratangential rays. Axial ray long, of (1987) uniform width and circular cross-section. Two pairs of para- Figs. 7H; 9W,Z;10K, Z tangential rays not coplanar, but diverging away from their junction with the axial ray; circular in cross section, with the 1987 Abnormisella insperata Fedorov in Fedorov & Pereladov, distal pair longer than the proximal pair. Proximal pair, when p. 41, pl. 12, figs. 7, 10–12 (figs. 4–6, 8, 9, 14–16 viewed along the plane of symmetry (Fig. 10K), are approxi- unassigned). mately perpendicular to the plane of the axial ray; distal pair 2008 Speciosuspongia wancunensis Chen & Dong, p. 25, form an angle of 50–60 degrees with each other, and lie at fig. 4K, L. 120–130 degrees to the axial ray (emended from Fedorov & 2018 Speciosuspongia cf. wancunensis; Peel, p. 315, fig. 3R, X. Peredalov 1987). Lectotype. – Here designated as the specimen figured by Discussion. – Fedorov in Fedorov & Peredalov (1987) Fedorov in Fedorov & Peredalov (1987, pl. 12, fig. 7; speci- assigned a number of illustrated stauractins and pentactins men 1625/11–5). to an assemblage prioposed as Abnormisella insperata, the type species of Abnormisella. He noted that about 70 % of Figured material. – Henson Gletscher Formation: PMU 34393 the spicules were stauractins, but the illustrated specimens are from GGU sample 271428. Fimbuldal Formation: PMU mainly simple morphologies that are widely distributed in 34436 from GGU sample 218644; Holm Dal Formation: Cambrian sclerite assemblages and therefore unlikely to be PMU 31819 from GGU sample 225594; PMU 31820 from individually diagnostic of Abnormisella. However, several illu- GGU sample 315008. strated specimens of a “spiderpentactin” (Mehl 1998) are Discussion. – The specimen selected here as lectotype is one of distinctive and a specimen with this unusual morphology is several “spiderpentactins” illustrated by Fedorov in Fedorov selected herein as lectotype of the type species, forming the & Peredalov (1987). Speciosuspongia wangcunensis (Chen & basis for emendation of Abnormisella. Dong, 2008) is considered to be a junior subjective synonym Thus, the spicule consist of two dissimilar pairs of short of Abnormisella insperata. This synonymy includes specimens paratangential rays and an oblique axial ray, disposed so that from the Holm Dal Formation (Guzhangian Stage) compared the spicule is bilaterally symmetrical about a plane which long- to Speciosuspongia wangcunensis by Peel (2018; Fig. 10K,Z) itudinally bisects the cylindrical axial ray and the angle between and to additional specimens from the upper Henson each pair of paratangential rays. This distinctive arrangement of Gletscher Formation (Wuliuan Stage; Fig. 7H) and rays is shared with Speciosuspongia Chen & Dong, 2008 from the Fimbuldal Formation (Drumian Stage; Fig. 9W) of Peary Miaolingian Series, Drumian Stage, of western Hunan Province, Land. Abnormisella insperata is also recorded from several China. However, the type species of Speciosuspongia, horizons in the lower Henson Gletrscher Formation Speciosuspongia hunanensis Chen & Dong 2008, is distinguished (Cambrian Stage 4) in southern Freuchen Land (Fig. 1B,C, by its transversely swollen axial ray which has the form of an locality 4). inverted heart, tapering distally. Spicules of this latter type were figured by Mehl (1998,pl.3,figs. 9–11) and are recognised also from North Greenland (Fig. 9B,H). Genus Australispongia (Dong & Knoll, 1996) A second species proposed by Chen & Dong (2008), – Speciosuspongia wangcunensis Chen & Dong, 2008, was Type species. Australispongia sinensis (Dong & Knoll, 1996) characterised by its long, cylindrical axial ray, the same from the Furongian Series (Paibian Stage) of Hunan, China. sclerite morphology recognised in the lectotype of Discussion. – Australispongia is represented in North Abnormisella insperata as illustrated by Fedorov in Fedorov Greenland by the type species, Australispongia sinensis from fi & Peredalov (1987, pl. 12, g. 7). This spicule morphotype is the Fimbuldal Formation (Cambrian Series 3, Drumian, also present in collections from North Greenland (Figs. Ptychagnostus punctuosus Biozone) and Holm Dal 7H, 10K). Formation (Cambrian Series 3, Guzhangian, Lejopyge laevi- Speciosuspongia wangcunensis is considered to be a junior gata Biozone), and Australispongia sp., also from the Holm subjective synonym of Abnormisella insperata.However,at Dal Formation (Peel 2017a, 2018). A possible third species, this time, Speciosuspongia,withtypespeciesSpeciosuspongia Australispongia? inuak n. sp., is described from the Henson hunanensis, is maintained as a separate genus, although the Gletscher Formation (Maiolingian Series, Wuiliuan Stage) of shared spiderpentactin morphology of the spicules suggests Gustav Holm Dal (Fig. 1; locality 9). 146 J. S. PEEL

Australispongia? inuak n. sp. “Calcihexactina” sp. Figs. 6X, Y, CC, DD; 7AA Fig. 4B,C,H,I,K–M

LSID. – urn:lsid:zoobank.org:act:2740A37C-9399-4802-8EED- Figured material. – Aftenstjernesø Formation; PMU 34330 from 4B5EA6C21280. GGU sample 271471 (Fig. 1C, locality 11); PMU 34331 from GGU sample 319740 (Fig. 1A,northernNyeboeLand);PMU Derivation of name. – From inuak (Greenlandic) meaning finger. 34332 and 34334 from GGU sample 315045 (Fig. 1C,locality3); Holotype. – PMU 34377 from GGU sample 271428, Henson PMU 34335 and 34336 from GGU sample 229894 (Fig. 1C, Gletscher Formation, Gustav Holm Dal, western Peary Land, locality 11). Kap Troedsson Formation: PMU 34333 from North Greenland (Fig. 1; locality 9). Mialongian Series, GGU sample 270505 (Fig. 1B,locality13). Wuliuan Stage. Discussion. – Pentactins (and infrequent stauractins and hex- Other figured material. – PMU 34376 and 34378 from the actins) from the Aftenstjernesø and Kap Troedsson forma- same sample and locality as the holotype. tions are characterised by long, slender, parallel-sided rays with a circular cross-section (Fig. 4B,C,H,I,K–M). Diagnosis. – Pentactine with long, slightly curved axial ray Specimens are common but the rays are invariably broken. ornamented by numerous slightly irregular, discontinuous, The four paratangential rays may lie within a single plane narrow longitudinal ridges that are separated by wider con- perpendicular to the axial ray but they are often inclined, or cave interspaces. Paratangential rays subequal in length or slightly curved, towards it (Fig. 4I). A hexactin from with one opposing pair greatly reduced. Navarana Fjord (Fig. 1C, locality 3) has more robust axial Description. – Pentactin with the slightly curved axial ray rays, with the sixth ray inclined (Fig. 4H), but is associated substantially longer and more robust than the four paratan- with the common slender hexactins with rays of equal form. gential rays (Fig. 6Y). Paratangential rays smooth, tapering, of Hyalostelia transitiva Fedorov in Fedorov & Peredalov, 1987 equal length or with one opposing pair greatly reduced (Fig. is a closely similar pentactin from the Kuonamka Formation 6X, CC). Axial ray with slightly irregular, sometimes weakly (Cambrian Series 2, stage 4) of north-eastern Siberia, although anastomosing fine longitudinal ridges separated by concave the genus is based on Carboniferous material (Reid 1968). interspaces (Fig. 6X, Y, CC). Similar pentactins were illustrated by Mao et al. (2013, fig. 2B) from the lower Kaili Formation (Cambrian Series 2, Discussion. – Australispongia? inuak n. sp. is only tentatively Bathyuriscus–Redlichia Biozone) of South China. referred to Australispongia since its axial ray is ornamented Sdzuy (1969)proposedCalcihexactina for spicules from the with narrow longitudinal ridges rather than with the three Wildenstein Formation (Cambrian Series 3) of the Franconian distinct, twisted, flanges seen in the type species, forest,Germany.Henotedthepresence of widened axial canals in Australispongia sinensis (Fig. 10G), known in North the calcareous rays but canals have not observed in the Greenland Greenland from the Holm Dal Formation. However, material. Botting & Muir (2018)consideredthecomposition a single specimen from the Fimbuldal Formation (Fig. 9E) probably to be secondary and the genus to be unrecognisable. which Peel (2017a) also referred to Australispongia sinensis Spicules similar to the Greenland specimens were illu- has five less pronounced flanges. Australispongia sp. from the strated by Brasier (1984, fig. 2R) from the English Midlands Holm Dal Formation (Fig. 10L)differs from Australispongia? (Cambrian Series 2), the Comley Limestone (Cambrian inuak in having blade-like flanges. Series 2) of England (Hinz 1987, pl. 15, figs. 10, 15, 16), the The ornamentation of Australispongia? inuak is also seen in early Cambrian of Hunan, China (Ding & Qian 1988, pl. 4, co-occurring specimens of Thoracospongia lacrimiformis Peel, figs. 1–5), Cambrian glacial erratics in Antarctica (Wrona 2017a (Fig.6T,U,W,AA,BB), suggesting that both spicules 2004, fig. 5D) and the lower Cambrian of North-East may occur within the same scleritome. Spicules of Greenland (Skovsted 2006, fig. 7.21, 7.22). Rays in specimens Australispongia? inuak are markedly less common in GGU sam- from the lower-middle Cambrian of Korea referred to ple 271428 than those of Thoracospongia lacrimiformis.Whilethe Calcihexactina by Lee (2006) are more robust and more four paratangential rays are usually subequal in length (Figs. 6Y, strongly tapering than the Greenland specimens. Specimens DD and 7AA), rare specimens show reduction of one opposing assigned to Calcihexactina by Brock & Cooper (1993) from pair (Fig. 6X, CC). the early Cambrian of South Australia also have more tapered rays than the Greenland spicules, with less sharply defined Genus Calcihexactina Sdzuy, 1969 junctions between the rays. Kouchinsky et al. (2015) illu- strated slender pentactins from the Emyaksin Formation Type species. – Calcihexactina franconica Sdzuy, 1969 from (Cambrian Series 2, stages 2–3) of northern Siberia, but the Wildenstein Formation (Miaolingian) of Germany. these differ from the North Greenland specimens in that the Discussion. – The name Calcihexactina Sdzuy, 1969 has been paratangential rays slope away from the axial ray rather than applied to a variety of widely distributed mainly hexactin shallowly towards it. Cambrian spicules and it is doubtful that its application has any useful meaning (Botting & Muir 2018). Diagnosis is not Genus Celtispongia n. gen. attempted and the name should be restricted to the type suite. In the present context, it is employed solely as a link in the LSID. – urn:lsid:zoobank.org:act:246041F9-65CF-45EA-ADDA- discussion of published records. 80A71773DC7C. GFF 147

Type species. – Celtispongia dorte n. gen. n. sp. from the Ptychagnostus atavus Biozone (Gapparodus bisulcatus– uppermost part of the upper member of the Henson Westergaardodina brevidens conodont Biozone) of western Gletscher Formation (Miaolingian Series, Wuliuan Stage) of Hunan Province, China is tentatively placed here. North Greenland. Derivation of name. – From the resemblance of the stauractin Genus Cjulanciella Fedorov in Fedorov & Peredalov (1987) spicule, with a central plate, to a Celtic cross. Type species. – Cjulanciella asimmetrica Fedorov in Fedorov Diagnosis. – Stauractin spicules with the point of intersection & Peredalov (1987) from the Kuonamka Formation of the rays lying within a central rectangular plate. Rays leave (Cambrian Series 2, Stage 4) of north-eastern Siberia. the rectangular central plate at its corners. Discussion. – Fedorov in Fedorov & Peredalov (1987, p. 42) Discussion. – Well preserved and abundant spicules from the designated the type species as Cjulanciella asymmetrica but upper Floran–lower Undillan (Miaolingian Series, Drumian elsewhere in the paper he employed the spelling Cjulanciella Stage) of the Georgina Basin were described by Mehl (1998, asimmetrica, including the formal description and some illus- p. 1164) as “platy stauractins” on account of the unique central trations of the type species itself. Cjunlankella asimmetrica plate. Sketches of the spicules were also given by Reitner & Mehl was employed for some figure descriptions (Fedorov in (1995, fig. 1.9, 1.13). The thin blade-like extensions between the Fedorov & Peredalov 1987, pl. 14). All name forms, and proximal parts of the four rays invite comparison with Cjulanciella assymetrica, exist in the sparse literature Tallitaniqa petaliformis Peel, 2018 from the Holm Dal (Fedorov & Peredalov 1987; Rozanov & Zhuravlev 1992; Formation (Fig. 10N, O) and, in particular, Sulukispicula gelidae Buslov et al. 1998, 2001; Zhuravlev & Leguta 2005). n. gen. n. sp. from the Ekspedition Bræ Formation (Fig. 8B,G). fi In Celtispongia dorte, however, the well-developed, rectangular, Mehl (1996)dened and designated the holotype of central plate lies with a single plane, essentially including the four Thoracospongia follispiculata on the basis of a longitudinally ridged follipinule but noted that these were associated with heav- rays, whereas the blade-like extensions in Sulukispicula gelidae fl fi lie oblique to the plane of the rays (Fig. 8D). In both forms, the ily in ated triaxon spicules (often modi ed to stauractins or central disc and blade-like extensions underlie the intersecting pentactins) which she termed pulvinusactins (Mehl 1998). The rays (Figs. 6P,Q,8G,H). follipinules and pulvinusactins were interpreted as forming part ofthesamesponge(Mehl1996, fig. 5, 1998,text-Fig. 2)and a similar reconstruction was presented by Sugai et al. (2004, fig. Celtispongia dorte n. gen. n. sp. 3A), based on specimens from the Bagrad and Sashkunar forma- Fig. 6O–Q tions of south-eastern Siberia. Zhuravlev & Leguta (2005)con- sidered that the pulvinusactins in Mehl (1996, 1998) reconstruction were referable to Cjulanciella and that 1995 (stauractins), Reitner & Mehl, fig. 1.9, 13. Thoracospongia Mehl (1996) should be a junior synonym of 1998 platy stauractins Mehl, p. 1164, pl. 3, figs. 2–5. Cjulanciella Fedorov in Fedorov & Peredalov (1987). The diag- LSID. – urn:lsid:zoobank.org:act:8BFE8B9C-5ABA-49F0-B75C- nostic follipinules of Thoracospongia follispiculata were not 8EF6A311DCF4. recorded by Fedorov in Fedorov & Peredalov (1987)fromthe Derivation of name. – For Dorte Mehl (Dorte Janussen) in Kuonamka Formation (late Stage 4 and Wuliuan Stage) in north- appreciation of her studies of Cambrian sponges. ern Siberia, but Kouchinsky et al. (2011) described both follipi- nules and inflated stauractins from the succeeding Drumian Stage Holotype. – PMU 34371 from GGU sample 271428, upper (Tomagnostus fissus – Paradoxides sacheri Biozone); the latter member of the Henson Gletscher Formation (Miaolingian were not referred to Cjulanciella by name. Series, Wuliuan Stage) of North Greenland (Fig. 1C, local- The synonymy of Thoracospongia with Cjulanciella is not ity 9). followed in the spicule-based taxonomy employed here, although Paratype. – PMU 34372 from GGU sample 271428, same the reconstruction of the sponge wall proposed by Mehl (1996, locality and horizon as holotype. 1998) is considered likely. Furthermore, as discussed by Peel (2017a) and references therein, differences are evident in the Diagnosis. – As for genus. postulated spicule associations of the respective sponges from Description. – Stauractin in which the proximal interspaces which the obese follipinule spicules are derived, especially with between the four rays are joined by a thin rectangular central regard to the presence or absence of pulvinusactins. plate, the angles of which correspond to the four rays (Fig. 6Q). In relative terms, the central plate underlies the rays (Fig. 6O–Q) Cjulanciella asimmetrica Fedorov in Fedorov & Peredalov which are circular in cross-section and rise slightly obliquely (1987) away from the central plate and their common origin. Figs 5B,D,H,I;6D,E;7Y Discussion. – In specimens from North Greenland, the rays are often more strongly differentiated from the central plate 1987 Hexactinellida, forma 9, Fedorov in Shabanov et al., than in Mehl (1998) specimens from the Drumian Stage of pl. 35, figs. 2, 3. the Georgina Basin. A specimen illustrated by Chen & Dong 1987 Cjulanciella asimmetrica Fedorov in Fedorov & (2008, fig. 3c) from the Miaolingian Series, Drumian Stage, Pereladov, p. 43, pl. 13, figs. 1, 2, 10–17, pl. 14, figs. 1–4. 148 J. S. PEEL

1998 Cjulanciella asymmetrica; Buslov et al., figs. 2, 1–4. Yang et al. (2010, pl. 2, Fig. 5–11) are comparable to North 2005 Cjulanciella asimmetrica; Zhuravlev & Leguta, p. 54, Greenland specimens and can be assigned to Cjulanciella. pl. 14, figs. 10, 11. 2015 Pentactines with smooth inflated rays, Kouchinsky et al., p. 502, fig. 70G–L. Genus Kuonamia Doweld, 2016 Figured material. – Henson Gletscher Formation (Cambrian 1987 Disparella Fedorov in Fedorov & Pereladov, p. 41 (non Series 2, Stage 4): PMU 34345 from GGU sample 315091 (Fig. Hessler 1970). 1B, locality 4); PMU 34346 from GGU sample 301351 (Fig. 2016 Kuonamia nom. nov. Doweld (2016, p. 19). fi 1B, locality 4); PMU 34347 from GGU sample 301352 (Fig. 2018 Kuonamia, Peel, p. 312, g. 5G, H, L. 1C, locality 5). Henson Gletscher Formation (Miaolingian Type species. – Disparella fusiformis Fedorov in Fedorov & Series, Wuliuan Stage): PMU 34361, 34362 and 34437 from Peredalov (1987), Cambrian Series 2, Stage 4, Oryctocara GGU sample 271428. Biozone, Kuonamka Formation, Kjuljanka River, Yakutia, North-eastern Siberia. Discussion. – Fedorov in Fedorov & Peredalov (1987) included within Cjulanciella asimmetrica a variety of globose Discussion. – Fedorov and Peredalov (1987, pl. 13) included diactin to hexactin spicules from the Bergeroniellus expansus morphotypes with two to six rays in a spicule assemblage and Oryctocara Biozones (Cambrian Series 2, Stage 4) of the referred to Kuonamia fusiformis; the illustrated isolated speci- Kuonamka Formation of north-eastern Siberia; they are con- mens are considered to be syntypes. sidered to be syntypes. Kouchinsky et al. (2011) included Kuonamia differs from Cjulanciella Fedorov in similar swollen spicules from the same area within Fedorov & Peredalov (1987) in the greater regularity of Thoracospongia sp. cf. follispiculata Mehl (1996), in line the spicule symmetry and the degree of inflation of the with the reconstruction of that taxon presented by Mehl rays. The degree of inflation in Cjulanciella is greater, (1996, 1998). Similar spicules may be common in collections and the angles between adjacent rays in stauractins are from the lower member of the Henson Gletscher Formation generally obtuse or acute. In contrast, rays in Kuonamia in southern Freuchen Land (Fig. 1B, C, locality 4), most often usually meet at about right angles (Fig. 10J). as hexactins; accompanying follipinules of Thoracospongia have not been found. Spicules are often asymmetric (Fig. 5B, D, I) with rays of different lengths (Fig. 5H). The rays Kuonamia fusiformis (Fedorov in Fedorov & Peredalov are typically short and strongly inflated, with their width half 1987) Figs 6W; 7V, X, U; 10J of their length. Their sides are convex proximally but distally they become shallowly concave as they taper to the pointed distal extremity. 1987 Disparella fusiformis Fedorov in Fedorov & Peredalov Specimens from the uppermost Henson Gletscher Formation (1987), p. 42, pl. 13, figs. 3–9. (GGU sample 271428, Fig. 1C, locality 9; Miaolingian Series, 2016 Kuonamia fusiformis; Doweld, p. 19. Wuliuan Stage) include highly inflated pentactins and hexactins 2018 Kuonamia fusiformis; Peel, p. 313, fig. 5G, H, L. in which the rays are almost absorbed into the globose central Figured material. – Henson Gletscher Formation: PMU zone (Figs. 6D and 7Y).Inthisfeaturetheyresemblethepulvi- 34375, 34398, 34399, and tentatively PMU 34400, from nusactins illustrated by Mehl (1998)fromtheWuliuanand GGU sample 271428 (Fig. 1C, locality 9), Miaolingian Drumian Stages of the Georgina Basin. The Greenland specimens Series, Wuliuan Stage, Ptychagnostus gibbus Biozone. Holm are associated with Thoracospongia lacrimiformis (Peel, 2017a, fig. Dal Formation: PMU 31843 from GGU sample 315008 (Fig. 6T,U,V,Z,AA,BB,EE)andwithAustralispongia? inuak n. sp. 1C, locality 6), Miaolingian Series, Guzhangian, Lejopyge lae- (Figs. 6X, Y, CC, DD and 7AA), the latter possibly forming part of vigata Biozone. thesamescleritomeasThoracospongia lacrimiformis. Buslov et al. (1998) illustrated spicules of C. asimmetrica (as Discussion. – Pentactins and hexactins with inflated, tapering C. asymmetrica) from the Chibit Formation of middle–late to fusiform rays are common in GGU sample 271428 from Cambrian age from Gornyi Altai, southern Siberia, including the uppermost Henson Gletscher Formation (Wuliuan Stage; specimens comparable to those figured here from the Henson Figs. 6W, 7V). Associated stauractins (Fig. 7U) have more Gletscher Formation (Stage 4; Fig. 5D,H,P).Cjulanciella asim- slender rays but may belong to the same scleritome. Rare metrica was also described from the Sinsk Biota (Cambrian Stage specimens of a diactin (Fig. 7BB) resemble this stauractin; 4) of Yakutia, eastern Siberia by (Zhuravlev & Leguta 2005). specimens considered to be diactins by Fedorov in Fedorov & Kouchinsky et al. (2015, fig. 70) compared inflated pentactins Peredalov (1987, pl. 13, Figs. 4,5) appear to be monaxons. and hexactins from the Emyaksin Formation of northern Rare stauractins from the Holm Dal Formation placed Siberia (Cambrian Stage 3, Atdabanian Stage, Delgadella anabara here are flattened on one surface but inflated on the opposite Biozone) to Cjulanciella but these, together with spicules from the one (Peel 2018, Fig. 6G,H,L;Fig. 10J). The fusiform rays are Anabar uplift described by Fedorov in Shabanov et al. (1987), are constricted slightly, proximally and narrow rapidly towards older than the North Greenland material. their pointed distal extremity. The rays are slightly more “Stubby hexactins” from the Zhalagou Formation of fusiform than in stauractins from the Kuonamka Formation Guizhou, South China (Cambrian Series 2), illustrated by illustrated by Fedorov in Fedorov & Peredalov (1987, pl. 13, GFF 149

Figs. 6, 9), but are similar to a pentact illustrated by Fedorov et al. 2016; Peel 2018). Sepkoski (2002) classified in Shabanov et al. (1987, pl. 34, fig. 14) from the lower Microcoryne as a sponge. The apparent presence of the axial Cambrian of the Anabar Anticline, Siberia. canal in the Greenland specimen (Fig. 4J, arrow) supports this Buslov et al. (1998) illustrated spicules assigned to interpretation since axial canals are not present in octocor- Kuonamia (as Disparella)cf.fusiformis from the Chibit allian spicules (Goldberg & Benayahu 1987). Formation of Gornyi Altai, southern Siberia, but their rays are tapering when compared to those figured here from the Henson Gletscher Formation (Stage 4; Fig. 7U, Genus Nabaviella Mostler & Mosleh-Yazdi, 1976 V, X). However, the syntype suite illustrated by Fedorov Type species. – Nabaviella gracilis (Mostler & Mosleh-Yazdi, in Fedorov & Peredalov (1987) included both forms. Some 1976) from the Mila Formation (upper Cambrian) of Iran. spicules from the Atdabanian Stage (Cambrian Series 2) assigned to Hexactinellida forma 8 by Fedorov in Shabanov et al. (1987, pl. 34, fig. 14; pl. 35, Figs. 4, 7) Nabaviella? sp. can be assigned to Kuonamia but the greater inflation and Fig. 6B,K slightly irregularity of others promotes referal to Figured material. – PMU 3458 and 34359 from GGU sample Cjulanciella. 271428, Henson Gletscher Formation, Peary Land (Fig. 1C, locality 9). Cambrian, Miaolingian Series, Wuliuan Stage, Ptychagnostus gibbus Biozone. Genus Microcoryne Bengtson in Bengtson et al. (1990) Discussion. – Clavulate anchoring spicules of Nabaviella nor- Type species. – Microcoryne cephalata Bengtson in Bengtson mally display at least five robust paratangential rays (forming et al. (1990) from the lower Cambrian of South Australia. the umbel) which curve back towards a long axial ray or shaft that may develop a distal swelling (Mostler & Mosleh-Yazdi 1976; Bengtson et al. 1990; Dong & Knoll 1996; Mehl 1998). Microcoryne sp. Mostler & Mosleh-Yazdi (1976) assigned a form with just Fig. 4A,J three paratangential rays, in addition to the axial ray, to Figured specimen. – PMU 34329 from GGU sample 270506, Nabaviella? triradiata Mostler & Mosleh-Yazdi, 1976. Mehl Kap Troedsson Formation, Cambrian Series 2 (Stage 4), (1998, pl. 3, Figs. 3, 4) described as the hexactinellid southern Wulff Land (Fig. 1B, locality 14). Nabaviella? similar spicules but with the umbel consisting of two to four paratangential rays from the ?upper Discussion. – The single specimen from North Greenland has Tempeltonian Stage (Wuliuan) of the Georgina Basin. In a preserved length of about 650 µm and shows about 15 thick, these the axial ray is more than five times the length of the radiating rays, with two in one quadrant of substantially greater maximum diameter of the umbel and tapers distally. length than the others (Fig. 4A, J). The shorter rays have Rare spicules from GGU sample 271428 (uppermost rounded terminations, but the longest two are broken distally, Henson Gletscher Formation; Wuliuan Stage) display three one seemingly showing a circular axial canal (arrow in Fig. 4J). rays at 120 degrees to each other which curve slightly The North Greenland specimen closely resembles spicules of towards the axial fourth ray, the length of which is not similar size figured by Elicki (1994, fig. 6.10–16) and Elicki & known (Fig. 6B, K). They appear to represent the broken Geyer (in Heuse et al. 2010) from the Charlottenhof Formation umbels of spicules similar to those illustrated by Mehl (Upper Ludwigsdorf Member; Cambrian Series 2, Stage 4) of the (1998).However,theycanalsobecomparedtentativelyto Görlitz Synclinorium in Germany, and a single specimen from anatriaene spicules figured by Zhuravlev & Leguta (2005,pl. the Burj Formation (Cambrian Series 3) of Jordan (Elicki 2011). 13, Fig. 4) in an articulated specimen of Ivantsovia andreyi There are fewer rays than in Bengtson’sspecimensof Zhuravlev in Zhuravlev and Leguta (2005)fromtheSinsk Microcoryne cephalata described from strata of early Cambrian Lagerstätten (Cambrian Series 2, Stage 4, Botoman Stage) of age (Stages 2 and 3) in South Australia, and the longer rays are Yakutia, eastern Siberia, interpreted as a demosponge. Some more clearly differentiated (Bengtson et al. 1990, fig. 19). The Greenland specimens appear to lack an axial ray and in axial general shape in the Australian material is club-like, with five or view they resemble spicules figured by Fedorov in Shabanov more longer rays fused together into a compact stem. et al. (1987,pl.34,Fig.11)fromthelowerCambrianof Kouchinsky et al. (2015, fig. 72B) interpreted a similar Siberia and by Dong & Knoll (1996, fig. 22) from the spicule from the uppermost Emyaksin Formation (Cambrian Furongian of Hunan, China. Series 2, Stage 4, Botoman Stage) of Bol’shaya Kuonamka, Van Kempen (1990) described a variety of tetraxon spicules northern Siberia, as a heteractinid sponge, pointing out simi- interpreted as derived from demosponges from the Ranken larities with Eiffelia. Limestone (Miaolingian) of the Georgina Basin but most were While not excluding interpretation as a sponge spicule, orthotriaenes, with the three paratangential rays directed away Bengtson (in Bengtson et al. 1990) pointed out similarities (upwards) and forming an obtuse angle with the axial ray. In the between Microcoryne cephalata and octocorallian spicules Greenland spicules, the three paratangential rays are initially figured by Bayer (1956). The latter interpretation was perpendicular to the axial ray before curving back (downwards) repeated by Elicki (1994) and Elicki (2011) and is often towards the axial ray (Fig. 6B). Such anatriaene forms were not noted in published literature (Bengtson 2004, 2005; Han described from Australia but Van Kempen (1990,Fig.3B,C) 150 J. S. PEEL illustrated late Ordovician examples from the Baltic in which the Diagnosis. – As for genus. paratangential rays are much longer and more strongly curved Description. – Stauractins are slightly domed such that the back (downwards) towards the axial ray than in the Greenland spicule has opposing convex and concave surfaces (Fig. specimens. In this respect they resemble Nabaviella? triradiata 6G). The mutually perpendicular rays are straight or and the specimens illustrated by Mehl (1998). Rhebergen & slightly curved along their length; some are slightly curved Botting (2014) commented that some Early Palaeozoic ana- within the plane of the rays producing a swirled effect triaene-like spicules may be of hexactinellid rather than demos- (Fig. 6F). Individual rays are hemispherical in cross- ponge affinity. section with the flattened surface lying within the concave surface of the spicule (Figs. 6G,F,Rand9T),theoppos- ing surface being either uniformly convex or slightly Genus Sanningasoqia n. gen. arched (Fig. 9R–T). Type species. – Sanningasoqia borealis n. gen. n. sp. Discussion. – Stauractins of Sanningasoqia borealis are very LSID. – urn:lsid:zoobank.org:act:E9B994F5-F59E-4AAF-A17E- common in GGU sample 271428 from the uppermost A5D55032D185. Henson Gletscher Formation (Miaolingian Series, Wuliuan Stage) but also range through the Fimbuldal Formation into Derivation of name. – From “sanningasoq” (Greenlandic) the Holm Dal Formation (Guzhangian Stage; Fig. 3). meaning cross. Diagnosis. – Stauractins slightly domed, so that the spicule has a convex and opposing concave surface; straight to Genus Seqineqia Peel, 2018 slightly curved rays are usually perpendicular to each other Type species. – Seqineqia bottingi (Peel, 2018), Holm Dal but may be slightly swirled distally. Rays are hemispherical in Formation (Miaolingian, Guzhangian Stage, Lejopyge laevi- cross-section, with the flat side lying on the concave side of gata Biozone), Peary Land, North Greenland. the spicule. Discussion. – Seqineqia was proposed by Peel (2018) for solari- Discussion. – While the stauractin plan is widespread in form spicules in which the acanthose rays are preserved within sponge spicule assemblages, the general hemispherical cross- a single plane, without any trace of axial rays. The unequal section of the rays is characteristic of Sanningasoqia. One pair depth of the interspaces between rays suggests division from an of rays in the spiderpentactin Speciosuspongia may be hemi- initial stauractin pattern. A simple hexactin with similar slen- spherical in cross-section, particularly in the proximal areas der rays and an 8-rayed polyactin, both with similar coarse (Figs. 7F and 9A, B, H) but in some cases this appears to be spines, were illustrated by Mehl (1998, pl. 3, figs. 13–14) from diagenetic in origin since the axial canals are exposed (Fig. the upper Tempeltonian of the Georgina Basin. 7C, arrow).

Seqineqia bottingi Peel, 2018 Sanningasoqia borealis n. gen. n. sp. Fig. 10U,V Figs. 6F, G, R; 9N, R–T; 10M, Q 2018 Seqineqia bottingi Peel, p. 313, fig. 4A–D, F, K. 2018 stauractin, Peel, fig. 3W, Z. Figured material. – PMU 31821 from GGU sample 225540, holotype, Holm Dal Formation (Miaolingian, Guzhangian LSID. – urn:lsid:zoobank.org:act:E9B994F5-F59E-4AAF-A17E- Satge, Lejopyge laevigata Biozone), Peary Land, North A5D55032D185. Greenland (Fig. 1C, locality 9). Derivation of name. – From “borealis” (Latin) meaning north- Discussion. – In addition to the holotype, Seqineqia bottingi ern: the northern cross. (Peel, 2018) is known from five additional specimens from Holotype. – PMU 34364 from GGU sample 271428 (Fig. 1C, the Holm Dal Formation in Gustav Holm Dal (Fig. 1C, locality 9), Henson Gletscher Formation, Miaolingian Series, localities 8 and 9). Peel (2018) noted the abundant acanthose Wuliuan Stage, Ptychagnostus gibbus Biozone. pentactines with widely spaced, similarly shaped spines on all rays that are associated with the solariform spicules (Fig. Paratypes. – Henson Gletscher Formation: PMU 34363 from 10W) and suggested that they may form part of the same the same sample and horizon as the holotype. Fimbuldal spicule skeleton. Carbonaceous sheaths of pentactines with Formation: PMU 34429 and 34431 from GGU sample a coarsely acanthose axial ray were illustrated from the 218626 (Fig. 1C, locality 10), PMU 34430 from GGU sample Forteau Formation (Cambrian Series 2, Stage 4) of western 218644 (Fig. 1C, locality 7); Miaolingian Series, Drumian Newfoundland by Harvey (2010). Stage, Ptychagnostus punctuosus Biozone. Other figured material. – Holm Dal Formation: PMU Genus Silicunculus Bengtson in Bengtson, Conway 31818 from GGU sample 225592 (Fig. 1C,locality8), Morris, Cooper, Jell & Runnegar, 1990 PMU 31814 from GGU sample 315008 (Fig. 1C,locality 6); Miaolingian Series, Guzhangian Stage, upper Lejopyge Type species. – Silicunculus australiensis Bengtson in laevigata Biozone. Bengtson, Conway Morris, Cooper, Jell & Runnegar, 1986 GFF 151 from the Miaolingian Series, Mindyallan Stage of Queensland, Well preserved fragmentary specimens of the cylindrical Australia. sponge Sanshapentella dapingi, measuring about 500 mm in height, were described from the Hetang Formation in southern Anhui, South China, by Xiao et al. (2005).Thespongeischar- Silicunculus australiensis Bengtson in Bengtson, acterised by prominent dermal pentacts in which the four para- Conway Morris, Cooper, Jell & Runnegar, 1990. tangential rays point towards the spongocoel. A short axial ray Fig. 10A,H points in the opposite direction but may be missing in some 1986 Silicunculus australiensis Bengtson, 1986 in Bengtson spicules (Xiao et al. 2005, fig. 8). The overall shape of the sclerite et al., p. 201, figs. 3, 11C. is clearly pointed, V-shaped, away from the spongocoel, with 2017a Silicunculus australiensis; Peel, p. 308, fig. 2A–D, E, F, straight paratangential rays. Spicules of Sisamatispongia differ in H–J. that their rays initially lie within a tangential place before 2018 Silicunculus australiensis; Peel, p. 314, fig. 3A, B, I, becoming strongly recurved (Peel 2018,Fig.5K,N,P;Fig. Y, EE. 10P). Pentactins of Sisamatispongia have not been observed – but it is a rare species in North Greenland. Reitner & Mehl Figured material. PMU 29996 from GGU sample 225540 (1995) gave no information about its abundance or distribution (Fig. 1C, locality 9) and PMU 30003 from GGU sample in the Georgina Basin and it was not described by Mehl (1998). 225595 (Fig. 1C, locality 8), Holm Dal Formation, Spicules described as Hunanospira delicata Qian & Ding in Ding Miaolingian Series, Guzhangian Stage, Lejopyge laevigata & Qian, 1998, from the lower Cambrian Yangjiaping Formation Biozone, Peary Land, North Greenland. of Hunan, China, also have a V-shaped form but are much Discussion. – The pentactins are closely similar to the type smaller than those of Sanshapentella dapingi,althoughXiao specimens of Bengtson in Bengtson et al. (1986) from et al. (2005) considered the two to be related. Bengtson (1986, Australia. Hooked spicules illustrated by Sugai et al. (2004, fig. 9I, J) figured sclerites of a similar V-shaped form to pl. 1, fig. 18) from the Bagrad Formation (Cambrian Series 2, Sanshapentella dapingi, but with more rays, from the Botoman Stage) of Bateny Ridge, south-eastern Siberia, and Guxhangian Stage (lower Mindyallan local stage) of by Dong & Knoll (1996, fig. 7.2, 7.19, 7.23) from the Queensland which he compared to Nabaviella. Furongian of Hunan, China, are tentatively assigned to An unfigured fragment of a stauractin spicule from the upper Silicunculus australiensis. member of the Henson Gletscher Formation (Cambrian Series 2, Stage 4, Ovatoryctocara granulata beds) has a V-shaped form reminiscent of the dominant spicules of Sanshapentella, but the Silicunculus saaqqutit Peel, 2017a four rays are flattened proximally, with a median ridge. Fig. 10B,C 2017a Silicunculus saaqqutit Peel, p. 309, fig., 2D, K, G, L. fi – Sisamatispongia erecta (Peel, 2018) 2018 Silicunculus saaqqutit; Peel, p. 314, g. 3C F. Fig. 10P – Figured material. PMU 29999 from GGU sample 225595, 1995 stauractin; Reitner & Mehl, fig. 1.10. holotype, Holm Dal Formation, Miaolingian Series, 2018 Sisamatispongia erecta Peel; p. 314, fig. 5K, N, P. Guzhangian Stage, Lejopyge laevigata Biozone, of North – Greenland (Fig. 1). Figured material. PMU 31846 from GGU sample 315008, Holm Dal Formation. Lejopyge laevigata Biozone, – ff Discussion. Silicunculus saaqqutit di ers from S. australiensis Miaolingian, Guzhangian Stage, glacier feeding Navarana in lacking a strongly hooked termination of the axial ray. Peel Fjord, southern Freuchen Land (Fig. 1C, locality 6). (2017a, 2018) suggested that Silicunculus saaqqutit may form – part of the same skeleton as S. australiensis since both species Discussion. Peel (2018) recorded just three specimens of this occur together in GGU samples 225594 and 225595. However, delicate, recurved, stauractin from the Holm Dal Formation morphological integration between the two spicule morpholo- at the head of Navarana Fjord (Fig. 1C, locality 6). gies has not been observed. Sisamatispongia erecta is also recognised from the middle Cambrian of the Georgina Basin on the basis of a drawing of a stauractin spicule presented by Reitner & Mehl (1995, Genus Sisamatispongia Peel, 2018 fig. 1.10). Type species. – Sisamatispongia erecta (Peel, 2018), from the Holm Dal Formation (Miaolingian Series, Guzhangian Stage, Genus Speciosuspongia Chen & Dong, 2008 Lejopyge laevigata Biozone), southern Freuchen Land, North Greenland. Type species. – Speciosuspongia hunanensis (Chen & Dong, 2008) from the Miaolingian Series, Drumian Stage, Discussion. – Reitner & Mehl (1995, fig. 1.10) compared Ptychagnostus atavus Biozone (Gapparodus bisulcatus– a figured spicule from the middle Cambrian of the Georgina Westergaardodina brevidens conodont Biozone) of western Basin, Australia, assigned herein to Sisamatispongia erecta, Hunan Province, China. with pentactine spicules of Sanshapentella dapingi (Mehl & Erdtman, 1994) and Hunanospira delicata (Qian & Ding in Diagnosis. – Emended from Chen & Dong (2008): Ding & Qian, 1998) from the lower Cambrian of China. Spiderpentactin with swollen axial ray which may be 152 J. S. PEEL transveresely flattened to globose and rounded in cross- Tempeltonian to lower Undillan (Miaolingian Series, section, tapering distally. Wuliuan–Drumian stages) of the Georgina Basin (Mehl 1998) and from the Fimbuldal Formation of North Discussion. – The name “spiderpentactins” proposed by Mehl Greenland (Fig. 9A, B, G, H). The “basal” surface of the (1998, p. 1162, pl. 3, figs. 6, 9–11) for spicules from the Greenland specimens is concave (Fig. 9B), as are the corre- Gowers Formation (Upper Floran–Lower Undillan; sponding, somewhat flattened, surfaces of the distal pair of Drumian Stage) of the Georgina Basin of Australia aptly paratangential rays, whereas the proximal pair are narrower describes spicules from contemporaneous strata in Hunan, and mainly rounded in cross-section. The axial ray terminates China assigned to Speciosuspongia (Chen & Dong, 2008). distally in a blunt spine or may be extended and slightly Although pentactin in form, the axial ray in Speciosuspongia curved (Fig. 9G); its surface is finely granulated. is inclined at a low angle to the plane delimited by the tips of the four paratangential rays (Figs. 7A and 9G) whereas the axial ray is usually perpendicular to this plane in most other Speciosuspongia inughuitorum n. sp. pentactins (Figs. 4K–M and 6Y). The axial ray in the type Fig. 7D,L,M,P species Speciosuspongia hunanensis Chen & Dong, 2008 from LSID. – urn:lsid:zoobank.org:act:EE011543-3CBE-428D-80D4- China is transversely swollen, shaped in the form of an D146E21C6176. inverted heart with a pointed tip (Fig. 9H), but it may be elongate, tapering, in other material assigned to Derivation of name. – For the Inughuit, the Polar Eskimos, Speciosuspongia (Figs. 7D, L and 9G). The four paratangential the native people of northern Greenland. rays are not equally developed but consist of two pairs of rays. Holotype. – PMU 34391 from GGU sample 271428, The pair closest to the axial ray diverge at an angle of about Fimbuldal Formation. Miaolingian Series, Drumian Stage, 180 degrees and of different length than the distal pair which Ptychagnostus punctuosus Biozone. diverge from each other at an angle of almost 90 degrees. The paratangential rays are usually inclined away from their point Paratypes. – PMU 34390 and PMU 34392 from the same of contact with the axial ray, delimiting a shallow (“basal”) sample and horizon as the holotype. cone. Thus, spicules of Speciosuspongia are not radially sym- Diagnosis. – Species of Speciosuspongia with massive, taper- metrical about the axial ray, as is most often the case in ing, long axial spine and reduced paratangential rays. pentactins, but bilaterally symmetrical about a plane which longitudinally bisects the axial ray and the angle between each Description. – Spiderpentactin in which the long axial ray is of the pairs of paratangential rays. massive and globose in its proximal part, slowly tapering and Abnormisella is similar to Speciosuspongia in being defined curved distally. The paratangential rays are reduced in length, by a spiderpentactin spicule form but is distinguished by the narrow and circular in cross-section, with an angle of only cylindrical form of its elongate axial ray. In contrast, the axial about 50 degrees between the distal pair (Fig. 7L, M, P). ray is distinctly swollen in Speciosuspongia hunanensis. Chen Ornamentation of axial ray with traces of longitudinal ridges & Dong (2008) described a second species of proximally (Fig. 7L, arrow); weakly acanthose distally. Specisosuspongia, Speciosuspongia wancunensis (Chen & Discussion. – Speciosuspongia inughuitorum n. sp. differs from Dong, 2008), also from the Drumian Stage of Hunan. It is Speciosuspongia hunanensis in terms of its massive axial ray characterised by a long cylindrical axial ray and is therefore which is much longer than the inverted heart-shaped ray of transferred to Abnormisella. the type species. Additionally, its paratangential rays are narrower, with the distal pair closer together than in Speciosuspongia hunanensis (Fig. 7P). Speciosuspongia hunanensis Chen & Dong, 2008 Fig. 9A,B,G,H This distinctive spicule was illustrated from the Kuonamka Formation (basal Drumian Stage) of northern Siberia by 1998 platy, spider-like pentactins Mehl, p. 1162, pl. 3, figs. 6, Kouchinsky et al. (2011, fig. 39F, G). Kouchinsky et al. 9–11. (2011) assigned the spicule to Thoracospongia sp. cf. 2008 Speciosuspongia hunanensis Chen & Dong, p. 26, T. follispiculata (Mehl, 1996) but the spiderpentactin form is fig. 4a–j. evident, justifying its transfer to Speciosuspongia. Figured material. – From the Fimbuldal Formation (Miaolingian Series, Drumian Stage, Ptychagnostus punctuo- Speciosuspongia spp. sus Biozone): PMU 34417 from GGU sample 218626, PMU Figs. 7A–G,IC, –K, N, R; 9C, D, I, J, P 34418 from GGU sample 218626, PMU 34419 from GGU – sample 218644, PMU 34420 from GGU sample 218626. Figured material. Upper Henson Gletscher Formation, GGU sample 271428 (Miaolingian Series, Wuliuan Stage, Discussion. – Material illustrated by Chen & Dong (2008, fig. Ptychagnostus gibbus Biozone): PMU 34379–34381, 4A–J) is derived from the Miaolingian Series, Drumian Stage, 34384–34389. Ptychagnostus atavus Biozone (Gapparodus bisulcatus– Fimbuldal Formation (Miaolingian Series, Drumian Stage, Westergaardodina brevidens conodont Biozone) of western Ptychagnostus punctuosus Biozone): PMU 34421 from GGU Hunan Province, China. It has a flattened, inverted heart- sample 218644, PMU 34422 from GGU sample 218626, PMU shaped axial ray also seen in specimens from the ?upper 34423 and 34424 from GGU sample 218644. GFF 153

Discussion. – The great morphological variation shown by Holm Dal Formation, but in the latter they are smaller and spiderpentactins from the Miaolingian of North Greenland more regularly disposed on stauractins in which the rays are might promote the notion that they should all be placed perpendicular to each other (Fig. 10N, O). Mao et al. (2013, within a single taxon. Priority dictates that this should be fig. 2A1) illustrated a spicule from the lower Kaili Formation assigned to Abnormisella rather than Speciosuspongia. This (Series 2, Stage 4) of South China in which the rays are stated step is not followed, however, partly on account of the type to carry blade-like extensions, but the rays are perpendicular materials being insufficiently described, but mainly due to to each other in contrast to Sulukispicula gelidae; it might be a lack of understanding of the significance of the spiderpen- referred tentatively to Sulukispicula. tactin spicule in poriferan taxonomy. It is clear from their wide distribution (Fedorov & Peredalov 1987; Heredia et al. 1987; Mehl 1998; Kouchinsky et al. 2011; Peel 2018) that the Sulukispicula gelidae n. gen. n. sp. – development of spiderpentactins represents an important Fig. 8B D, G, H, K event in middle Cambrian sponge evolution. Thus, LSID. – urn:lsid:zoobank.org:act:12A8262E-49A8-45C1-927B- Abnormisella and Speciosuspongia are maintained for spicules 99DAE78DE056. which are morphologically close to their respective type spe- – “ ” cies. While one additional new species, Speciosuspongia Derivation of name. From gelidus (Latin) meaning icy. inughuitorum n. sp., is proposed, most North Greenland Holotype. – PMU 34406 from GGU sample 315117, basal specimens are grouped together as Speciosuspongia spp. Ekspedition Bræ Formation (Miaolingian Series, Drumian fi until their relationships are clari ed. Stage, Ptychagnostus atavus Biozone) southern Freuchen The axial ray in most North Greenland specimens is Land (Fig. 1B, C locality 4). more uniformly rounded in cross-section than the flattened – – form seen in Speciosuspongia hunanensis. In general pro- Paratypes. PMU 34404, 34405 and 34407 34409, from the portions,however,mostspecimens from the upper Henson same sample as the holotype. Gletscher Formation (Wuliuan Stage) are otherwise closely Description. – Stauractin in which the four almost coplanar similar to Speciosuspongia hunanensis (Fig. 7A,B,F,G,J). rays are usually not regularly perpendicular to each other but Comparable specimens from the Fimbuldal Formation are meet at between about 80–120 degrees. Rays are distally often more acanthose (Fig. 9C,D,J).Thereisalsocon- circular in cross-section but blade-like extensions (arrows in siderable variation in the length and degree of tapering of Fig. 8B, D, G, H) on the presumed “basal” surface (Fig. 8B) the axial ray (Figs. 7E,I,Oand9P);itmaybebent(Fig. form narrow wings on each side of the rays in their proximal 7A, E) and in rare specimens an axial ray may be lacking part, delimiting a concave area between. Distally, the blade- (Fig. 7K). The paratangential rays are equally variable in like extensions may terminate abruptly (Fig. 8D) or taper into form and length, the latter is also clearly evident in the the main part of the ray (Fig. 8K). Chinese material of Speciosuspongia hunanensis (Chen & – Dong 2008, fig. 4A–J).InGreenlandmaterial,raysforming Discussion. Available spicules are usually broken distally, the distal pair are often shallowly concave on the “basal” sometimes corroded so as to display traces of the axial cavity, surface whereas the proximal pair are usually circular in but demonstrate the variation from perpendicular to each cross-section (Figs. 7F and 9B). The proximal paratangen- other (Fig. 8K) to meeting irregularly at angle of about – tialraysmayalsovaryinlengthfromshortspines(Fig. 9C, 80 120 degrees (Fig. 8B, H). D) to long rays (Fig. 9A). Genus Tallitaniqa Peel, 2018 Genus Sulukispicula n. gen. Type species. – Tallitaniqa petalliformis (Peel, 2018), from the LSID. – urn:lsid:zoobank.org:act:F784D7D6-FF9D-4699-9BB2- Holm Dal Formation. Lejopyge laevigata Biozone, Cambrian A00464A9DA5C. Series 3, Guzhangian Stage, Holm Dal, Peary Land (Fig. 1, locality 9). Type species. – Sulukispicula gelidae n. gen. n. sp. from the Ekspedition Bræ Formation (Miaolingian Series, Drumian Stage, Ptychagnostus atavus Biozone) of North Greenland. Tallitaniqa petalliformis Peel, 2018 Fig. 10N,O Derivation of name. – From “suluk” (Greenlandic) meaning wing, an allusion to the blade-like winged margins of the 2018 Tallitaniqa petalliformis Peel, p. 315, fig. 5D–E, O, Q. spicule rays. Holotype. – PMU 31841 (315008-A1) from GGU sample Diagnosis. – Stauractins in which the rays are usually copla- 315008, Holm Dal Formation (Miaolingian Series, nar, but not perpendicular to each other. Blade-like exten- Guzhangian Stage, Lejopyge laevigata Biozone) from the wes- sions on the presumed “basal” surface (Fig. 8B) form narrow tern side of the glacier feeding into Navarana Fjord, south- wings on each side of the rays in their proximal part, delimit- east Freuchen Land (Fig. 1C, locality 6). ing a concave channel between. Discussion. – Peel (2018) noted about 20 spicules of Discussion. – The blade-like extensions on the ray are remi- Tallitaniqa petalliformis from samples collected at the same niscent of Tallitaniqa petalliformis (Peel, 2018) from the locality and horizon as the holotype. The raised edges of the 154 J. S. PEEL petal-like excavations on the rays are reminiscent of the Discussion. – This strongly inflated, globose, sclerite is rare in blade-like wings on rays of Sulukispicula gelidae from the GGU sample 271428. It lacks the distally pointed axial ray Ekspedition Bræ Formation (Fig. 8B–D, G, H) but are more and longitudinal ribs characteristic of Thoracospongia lacri- regularly formed. In contrast to Tallitaniqa petalliformis, miformis, with which it occurs, but compares closely to most rays in the stauractins of Sulukispicula gelidae do not smooth sclerites described by Castellani et al. (2012) from intersect at right angles. the Furongian Series of southern Sweden. The degree of inflation of the axial ray is similar to that in the type species, Thoracospongia follispiculata, from the Upper Floran–Lower Genus Thoracospongia Mehl, 1996 Undillan of the Georgina Basin (Mehl 1998), but the latter is Type species. Thoracospongia follispiculata Mehl, 1996 longitudinally ribbed. Thoracospongia lacrimiformis Peel, 2017a Fig. 6T– V, Z, AA–BB, EE Sponge spicule assemblages Buen Formation 2004 pinular hexactine, Sugai et al., pl. 1, fig. 21. 2017a Thoracospongia lacrimiformis Peel, p. 311, fig. 3A–H. Siliciclastic sediments of the Buen Formation (Cambrian Series 2, Stages 3–4) crop out extensively in a broad belt Figured material. – PMU 29990, holotype, PMU 29991–29993, extending eastwards from Aftenstjernesø and in a second paratypes, PMU 34374 from GGU sample 271428, uppermost zone along the north coast from the outer reaches of J.P Henson Gletscher Formation (Maiolingian Series, Wuliuan Koch Fjord, through Freuchen Land to northern Wulff Stage), Gustav Holm Dal, western Peary Land, North Land and beyond (Ineson & Peel 1997, 2011; Fig. 1). The Greenland (Fig. 1; locality 9). outer shelf mudstones yield the only articulated sponges Discussion. – More than a hundred spicules are known from known from the Cambrian of North Greenland, with the same sample and locality as the holotype. The type spe- a diverse assemblage dominated by protomonaxonids cies, Thoracospongia follispiculata (Mehl, 1996) from the described from the Sirius Passet Lagerstätte (Cambrian Stage Miaolingian Series (Templetonian–Undillan; Wuliuan and 3; Fig. 1C, locality 1) by Botting & Peel (2016). Botting et al. Drumian stages) of Australia (Mehl 1996, 1998)differs in (2015) also described crown group demosponge microscleres. having a more inflated, blunt, axial ray with more prominent The lagerstätte occurs in the lowest fossiliferous beds of the paratangential rays. The paratangential rays of specimens formation in a facies transitional to deep trough clastics described by Kouchinsky et al. (2011) from the Kuonamka which crop out along the northernmost coast of Greenland Formation of north-eastern Siberia (Miaolingian Series, (Higgins et al. 1991; Ineson & Peel 1997, 2011; Peel & Ineson Drumian Stage) are much larger and more inflated than in 2011; Peel & Willman 2018). T. lacrimiformis, which also differs in terms of its finer, more Rare, poorly preserved articlulated sponges occur in south- clearly delimited, longitudinal ornamentation on the axial ray. ern outcrops of the formation in strata near the middle of the Sugai et al. (2004, pl. 1, fig. 21) illustrated a spicule from the formation in the Børglum Elv region (Fig. 1A; Peel & Bagrad Formation (Cambrian Series 2, Botoman Stage) of the Willman 2018) in mudstones deposited during the early Bateny Ridge, south-eastern Siberia, which can be assigned to part of Stage 4. Fragmentary large specimens and isolated T. lacrimiformis. spicules of Lenica cf. unica Goryanskii, 1977 from the upper Peel (2017a) discussed the spicular composition of scleri- Buen Formation in northern Freuchen Land (Botting & Peel tomes containing follipinules of Thoracospongia (or similar spi- 2016; Peel & Willman 2018; Fig. 1C, locality 2) are associated ff cules) and other grossly inflated spicules (pulvinusactins of Mehl with Ratcli espongia freuchenensis Botting & Peel, 2016. 1998) described by Rigby (1975), Mostler and Mosleh-Yazdi (1976), Pittau et al. (2003), Elicki (2011), Kouchinsky et al. Aftenstjernesø and Kap Troedsson formations (2011) and Castellani et al. (2012). He noted that Sugai et al. (2004, Fig. 3a) suggested that several pentactine spicules from The Aftenstjernesø Formation crops out across North the Shashkunar Formation (Botoman Stage) of Gorny Altai Greenland from northern Nyeboe Land to Børglum Elv formed part of the scleritome of Thoracospongia, but the diag- (Fig. 1A). At the type section in southern Lauge Koch Land nostic strongly modified pentactin (follipinule of Mehl 1998) (Fig. 1C, locality 11) the formation is 62 m thick and domi- was not included in their figured reconstruction. However, its nated by wavy bedded and graded dolomites with prominent occurrence in the contemporaneous Bagrad Formation was breccias (Ineson & Peel 1997). The basal member (about 2 m) noted (Sugai et al. 2004, pl. 1, fig. 21; Peel 2017a). of fossiliferous glauconitic, phosphoritic and pyritic dolomite represents a starved outer shelf ramp succession that can be traced throughout the outcrop area. In areas southwest of Thoracospongia sp. Nordenskiöld Fjord, across southern Wulff Land (Fig. 1B), Fig. 6L this basal member is equivalent to proximal ramp limestones Figured material. – PMU 34368 from GGU sample 271428, and shales of the Kap Troedsson Formation (Ineson & Peel uppermost Henson Gletscher Formation, Gustav Holm Dal, 1997). western Peary Land, North Greenland (Fig. 1; locality 9). The most characteristic spicules in samples from the basal Miaolingian Series, Wuliuan Stage. Aftenstjernesø and Kap Troedsson Formations are pentactins GFF 155 tentatively assigned to Calcihexactina (Sdzuy, 1969), origin- The lower and upper members of the Henson Gletscher ally described from the Wildenstein Formation (Miaolingian Formation are richly fossiliferous (Peel 1994; Geyer & Peel Series) of the Franconian forest, Germany (Figs. 3 and 4B,C, 2011;Peeletal.2016).TheboundarybetweenCambrian H, I, K–M). Specimens are common but the long, slender, Series 2 and Cambrian Series 3 (= Miaolingian Series) is parallel-sided rays are always broken. Sdzuy (1969) noted the located within the upper member in the Lauge Koch Land to presence of a wide axial canal in the rays but canals have not Løndal area (Fig. 1C, localities 10–12), where Wuliuan observed in the Greenland specimens. Similar spicules from (Cambrian Stage 5) deposits overlie the Ovatoryctocara Cambrian Series 2 were illustrated from North-East granulata assemblage (Cambrian Series 2, upper Stage 4; Greenland (Skovsted 2006), from central England (Brasier Geyer & Peel 2011;Peeletal.2016), but Miaolingian faunas 1984; Hinz 1987), northern Siberia (Fedorov & Peredalov have not been recognised in southwest Freuchen Land (Fig. 1987; Kouchinsky et al. 2015), Hunan, China (Ding & Qian 1C, localities 4 and 5). In northern Nyeboe Land, 1988) and Antarctica (Wrona 2004). Other specimens Miaolingian faunas of Wuliuan and Drumian age were assigned to Calcihexactina in the literature are more robust described by Peel (1994). (Korea, Lee 2006; South Australia, Brock & Cooper 1993). Spicules from the lower member of the Henson However, pentactins and associated stauractins and hexactins Gletscher (Fig. 3) are most common in collections from are widespread in the Cambrian fossil record and no doubt southern Freuchen Land (Fig. 1C, localities 4 and 5; Geyer occured in many species of sponges; they are not usually & Peel 2011, fig. 4). Pentactins and hexactins with long named (e.g. Dong & Knoll 1996; Elicki 2011; Mao et al. 2013). narrow rays are common (Fig. 5S,U),asisCjulanciella Spicules with six rays assigned to Eiffelia Walcott, 1920 asimmetrica (Fig. 5B,D,H,I,P).Apartfromthetype are widely distributed but often difficult to distinguish from material from the Kuonamka Formation of northern accompanying 6–0 spicules of chancelloriids due to poor Siberia (Fedorov in Fedorov & Peredalov 1987), preservation. Syndepositional erosion, redeposition and Cjulanciella has been illustrated from southern Siberia pervasive phosphatisation often removes all trace of the (Buslov et al. 1998; Sugai et al. 2004), eastern Siberia hollow, chambered rays and basal pores characteristic of (Zhuravlev & Leguta 2005), Australia (Mehl 1996, 1998) the chancelloriid sclerites and the identity of many 6–0 and Guizhou Province, China (Yang et al. 2010). sclerites is only evident from comparison to other co- Dodecaactinella oncera is common at one horizon (GGU occurring morphotypes which find no equivalence in sample 298544) about 16 m above the base of the formation Eiffelia.SpiculesofEiffelia from the Aftenstjernesø and (Fig. 5E–G). Abnormisella insperata occurs sporadically. Kap Troedsson formations usually have rays shortened by Sponge spicules from the Ovatoryctocara granulata assem- erosion (Fig. 4E and F) and may be flat, with coplanar rays, blage (Cambrian Series 2, upper Stage 4) were described from or slightly domed. Similar spicules were described by Elicki the basal part of the upper member of the Henson Gletscher (2011) from the Miaolingian Series of Jordan and Formation in Løndal by Peel et al. (2016). Hexactins of Kouchinsky et al. (2015, fig. 72D, H) from the Emyaksin Eiffelia floriformis n. sp. are characteristic but uncommon Formation (Cambrian Series 2, Stage 4, Botoman) of (Fig. 5J, K, R); a single stauractin is also known from the Siberia. uppermost beds of the lower member of the Henson Rare polyactins from the Kap Troedsson Formation include Gletscher in southern Freuchen Land (Fig. 5A). Eiffelia flor- Microcoryne Bengtson in Bengtson et al., 1990 (Fig. 4A,J), iformis is also recognised in the uppermost beds of the originally described from South Australia, but possibly also Emyaksin Formation (Cambrian Series 2, Stage 4, Botoman, present in the uppermost Emyaksin Formation of Siberia Calodiscus–Erbiella Biozone) of north-east Siberia in material (Kouchinsky et al. 2015). Robust irregular 6 and 7-rayed forms described by Kouchinsky et al. (2015). A club-shaped poly- (Fig. 4D, G) are reminiscent of an 8-rayed spicule figured by actin (Fig. 5T), also present in the Bagrad Formation Bengtson et al. (1990, fig. 21E) from the lower Cambrian (Cambrian Series 2, Botoman Stage) of south central Siberia Kulpara Limestone of South Australia and Acanthosphaera echi- (Khakassia Republic; Sugai et al. 2004), is similar to monax- nus (Duan, 1986) from the lower Cambrian of Hubei Province, ons figured by Yang et al. (2010, pl. 1, Fig. 1–7) from the China. lower Cambrian of Guizhou Province, China. They occur together with hexactins of Cjulanciella and slender pentactins and hexactins similar in form to those present in the lower member of the Henson Gletscher Formation (Fig. 5D, P and Henson Gletscher Formation Fig. 5S, U, respectively). The Henson Gletscher Formation is 62 m thick at its type With the exception of rare fragments of coarsely ribbed rays section in southern Lauge Koch Land (Fig. 1C,locality11) (Fig. 5W) from GGU sample 218614 (Fig. 1C,locality10),sponge but thickens to 112 m in southwest Freuchen Land (Fig. 1, spicules of Wuliuan age are known only from GGU sample locality 4). It can be recognised as far west as northern Nyeboe 271428 from 4 m below the top of the Henson Gletscher Land (Fig. 1A) and east to the Løndal area (Fig. 1C). Lower Formation in Gustav Holm Dal (Fig. 1C,locality9).Theprocessed and upper members of dark, thin-bedded limestones and sample yielded several thousand spicules, numerically dominated dolomites are separated by a middle member of fine grained by stauractins, many of which are similar to spicules referred to cream sandstone in outcrops between Nordenskiöld Ford and Hyalostelia transitiva Fedorov in Fedorov & Peredalov, (1987) Løndal but the sandstones are inconspicuous in northern out- from the Kuonamka Formation (Cambrian Series 2, Stage 4, crops (Ineson & Peel 1997). Oryctocara Biozone and Miaolingian Series, Wuliun Stage) of 156 J. S. PEEL northern Siberia (Fedorov & Peredalov 1987;Kouchinskyetal. wackestones that are interbedded with grey calcareous mud- 2011). Stauractins with an oval or circular cross section are most stones (Ineson & Peel 1997). The formation attains common and may attain 1 cm in diameter. They are planar to a thickness of 82m at the type locality in eastern Lauge slightly domed, with straight or slightly curved rays (Fig. 6N, 7S Koch Land (Fig. 2, locality 11) but thins to the west across and T), although stauractin spicules with more rapidly tapering southern Freuchen Land, becoming dominantly argillaceous. rays (Fig. 7U) or a pronounced bend in one or more rays occur R.A. Robison (written communication 1987, cited by Peel & infrequently (Fig. 7T); pentactins are rare. Stauractins of Streng 2015) dated the current samples to the Ptychagnostus Sanningasoqia borealis are also very common, characterised by atavus Biozone, the basal zone of the Drumian Stage rays with a hemispherical cross section (Fig. 6F,G,R). (Miaolingian Series; Fig. 2), equivalent to the lower part of Pentactins of Thoracospongia lacrimiformis,withthe Bolaspidella Biozone, but see discussion by Geyer & Peel a pointed, strongly inflated, axial ray and greatly reduced (2017). paratangential rays (Fig. 6T,U–W, Z, AA, BB, EE), are Limestone at the base of the Ekspedition Bræ Formation on associated with rare pentactins with a similarly ornamen- a nunatak in southern Freuchen Land (GGU sample 315117; ted but elongate narrow axial ray. The latter are assigned Fig. 1B, C locality 4) yields stauractins of Sulukispicula gelidae, to Australispongia? inuak sp.nov.(Fig. 6X,Y,CC,DD) known only from this locality, associated with rare and mainly but may form part of the same scleritome. Thoracospongia poorly preserved pentactins with an elongate axial ray (Fig. 8E, lacrimiformis is recognised also in the Bagrad Formation F). A similar etched pentactin from the lower part of the (Cambrian Series 2, Botoman Stage) of the Bateny Ridge, formation in its type section (Fig. 2C, locality 11) displays south-eastern Siberia (Sugai et al. 2004,pl.1,fig. 21). The concentric banding in the rays (Fig. 8I, O) similar to that type species, Thoracospongia follispiculata,fromthe illustrated by Castellani et al. (2012) in a pentactine from the Miaolingian Series (Templetonian–Undillan; Wuliuan Furongian of Sweden, but it is not known if the feature is and Drumian stages) of Australia (Mehl 1996, 1998), has original, as illustrated by Müller et al. 2007, fig. 6) in recent a more globose axial ray, although Reitner & Mehl (1995, material, or a result of diagenesis. Fig. 1.4) and Mehl (1996, pl. 2, Fig. 2) also illustrated A phosphatised hardground (Peel 2017b) located in the a pointed spicule from the Georgina Basin. The axial ray upper part of the formation in its type section (Fig. 2C, in pentactin and hexactin follipinule spicules from the locality 11) is associated with small scours containing rede- Kuonamka Formation of north-east Siberia (Miaolingian posited sponge spicules. The spicules are phosphatised, often Series, Drumian Stage) assigned by Kouchinsky et al. partially coated in pyrite, and usually eroded with rounded (2011)toT. follispiculata varies from globular to elongate, ends to the rays. Many occur within rounded, phosphate but the paratangential rays are much larger than in coated clasts (Fig. 8J, M, N). Stauractins and pentactins with T. lacriformis. perpendicular or oblique limbs are most common, with less Rare pentactins with a globular axial ray, but without frequent hexactins (Fig. 7J, L). The paratangential rays may longitudinal ridges (Fig. 6L), compare well with obese pen- lie within a single plane or have one or more non-planar tactins described from the Furongian (Stage 10) of Sweden by rays. Castellani et al. (2012). The latter are associated with pentac- tins and hexactins similar in shape to the abundant staurac- Fimbuldal Formation tins of GGU sample 271428 (Fig. 7W), but stauractins are rare in the Swedish material while pentactins and hexactins are The Fimbuldal Formation is a mainly cliff-forming unit com- rare in the Greenland sample. posed of pale carbonates and thin darker interbeds (Ineson & Hexactins with inflated, rapidly tapering, pointed rays (Fig. Peel 1997). Dolostone breccia beds are conspicuous, particu- 6W), associated with similar pentactins (Fig. 5X), were illu- larly in the upper part of the formation. At its type section in strated from the Undillan of Queensland by Mehl (1998 , pl. 1, Gustav Holm Dal (Fig. 1C, locality 7) the formation attains Fig. 1) and are here referred to Kuonamia. Plate-like staur- a thickness of about 180 m, but it thins to less than half this at actins proposed as Celtispongia dorte n. gen. n. sp. (Fig. 6P– Henson Gletscher. Faunas indicative of the Ptychagnostus pun- Q) were reported to be abundant in the Undillan of the tuosus Biozone (Miaolingian Series, Drumian Stage, Robison Georgina Basin by Mehl (1998, p. 1164). 1984) occur in a recessive limestone interval near the middle of Rare spicules assigned to Nabaviella?sp.(Fig. 6B,K) the formation, from which all spicules (Fig. 3, 9) are derived. were also illustrated by Mehl (1998)fromtheGeorgina Stauractins of Sanningasoqia borealis, with their characteris- Basin (Wuliuan Stage) of Australia. Similar spicules, but tic hemispherical cross-section (Fig. 9N,R,S),arecommon,as without an axial ray, were figured by Fedorov in are slender stauractins and rarer pentactins (Fig. 9M). Pentactins Shabanov et al. (1987) from the lower Cambrian of of Speciosuspongia show variation in the relative lengths of the Siberia and by Dong & Knoll (1996)fromthe rays (Fig. 9A–D), and the axial ray varies from flattened in the Furongian of Hunan, China. type species, Speciosuspongia hunanensis (Fig. 9A, B), to swollen, sometimes elongate, and acanthose (Fig. 9C, D, J). The distal pair of paratangential rays may be hemispherical in cross- section (Fig. 9B, H), as in Sanningasoqia borealis. Ekspedition Bræ Formation In addition to North Greenland, Speciosuspongia hunanensis The Ekspedition Bræ Formation consists mainly of pale- has been described from the Miaolingian Series, Drumian Stage weathering, thin-bedded, grey lime mudstones and skeletal of western Hunan Province, China (Chen & Dong 2008)and GFF 157 from the ?upper Tempeltonian to lower Undillan (Miaolingian pentactins assigned to Abnormisella insperata (= Speciosuspongia Series, Wuliuan and Drumian Stages) of the Georgina Basin, cf. wancunensis of Peel 2018), the axial ray is strongly oblique to Australia (Mehl 1998). Speciosuspongia inughuitorum is also the paired paratangential rays (Fig. 10K, Z). Pentactins of both known from the Kuonamka Formation (basal Drumian Stage) morphotypes maybe be smooth or finely acanthose. Stauractins of northern Siberia (Kouchinsky et al. (2011). with spindle-shaped, fusiform, rays are common, while rare staur- Australispongia sinensis (Fig. 9E) and Tallitaniqa petalli- actins show strongly tapering rays or oblique rays traversed by formis are each represented by single specimens. Other rare broad depressions near the origin (Fig. 10Y). Stauractins of spicules (Fig. 9K, L) resemble a pinular pentactin and tauactin Sanningasoqia borealis in which the rays have a hemispherical figured by Mehl (1998, pl. 2, figs. 10, 13) from the cross-section (Fig. 10M, Q) occur rarely at Navarana Fjord, and Templetonian (Miaolingian Series) of the Georgina Basin. are also known from Gustav Holm Dal. Rare, triactins with needle-like rays (Fig. 10X) resemble some spicules of Inflexiostella incognita FedorovinShabanovetal.(1987)from Holm Dal Formation Cambrian Series 2 (Atdabanian Stage) in the Anabar Anticline, Sponge spicule assemblages from the Holm Dal Formation although the latter tend to be slightly irregularly curved. (Fig. 3, 10) were described by Peel (2018) from Gustav Holm Tallitaniqa petalliformis (Fig. 10N, O) occurs in several fl Dal (Fig. 1C, localities 8 and 9) and Navarana Fjord (Fig. 1C, samples from Navarana Fjord while in ated stauractins of locality 6) who concluded that compositional differences Kuonamia fusiformis (Fig. 10J) and erect stauractins of between the two assemblages probably reflected the more Sisamatispongia erecta (Fig. 10P) are rare. argillaceous character of the formation at Gustav Holm Dal. Peel (2018) compared spicule assemblages from the Holm Robust acanthose pentactins in which the five rays are of Dal Formation with spicules from the contemporaneous similar size (Fig. 10W) dominate samples from Gustav Holm Mungerebar Limestone of Queensland (Miaolingian Series, Dal, although the axial ray may be significantly longer. Peel basal Mindyallan Stage, Acmarhachis quasivespa Biozone; (2018) suggested that the presence of similar widely spaced Guzhangian Stage) described by Bengtson (1986). thorn-like spines on the solariform spicules of Seqineqia bot- Silicunculus australiensis (Fig. 10A), Australispongia sinensis tingi (Fig. 10U, R) could indicate that the two morphotypes (Fig. 10G), coarsely acanthose pentactins (Fig. 10R, T), formed part of the same sponge scleritome, although the a dichodiactin (Fig. 10D) and similar slender pentactins smaller solariform spicules are rare. Other common spicules (Fig. 10S) occur in both assemblages. Reitner & Mehl (1995, fi from Gustav Holm Dal include smooth, slender stauractins g. 1.10) illustrated a spicule that is here assigned to and pentactins. Australispongia sinensis (Fig. 10G), Sisamatispongia erecta from the middle Cambrian Silicunculus australiensis (Fig. 10A, H) and Silicunculus saaq- (Miaolingian) of the Georgina Basin, Australia, but its strati- fi qutit (Fig. 10B, C) are widely distributed but uncommon. graphic derivation was not more closely speci ed. Large, smooth stauractins with straight to curved rays inter- Dong & Knoll (1996) described spicule assemblages from the secting at or near right angles dominate samples from Navarana Chefu and lower Bitiao formations (Cambrian Series 4 [= Fjord. In accompanying pentactins the axial ray varies from short Furongian Series], Paibian Stage) of Hunan, South China, (Fig. 10S)tolong;itmaybestraightorslightlycurved,and which include possible fragments of Sisamatispongia erecta perpendicular or slightly inclined to the paratangential rays and Silicunculus. Abnormisella insperata (Fig. 10K,Z)and which may themselves be subequal in size. In contrast, in a diactin with a secondary ray (Fig. 10E)alsoappeartobe

Figure 11. Biogeography of Cambrian sponge spicule assemblages. The four assemblages of spicules described from North Greenland (Cambrian Series 2, Stage 4; Wuliuan Stage; Drumian Stage; Guzhangian Stage) can be recognised in tropical successions from Siberia, South China and Australia. 158 J. S. PEEL represented in the assemblages described by Dong & Knoll Disclosure statement (1996). However, Konyrium Nazarov & Popov 1976, No potential conflict of interest was reported by the author. a distinctive element in the assemblages described by Bengtson (1986) and Dong & Knoll (1996), has not been recognised in North Greenland, although it was described from the References Rabbitkettle Formation (Cambrian, Furongian Series) of the MacKenzie Mountains of northwest Canada by Pratt (2002). Bayer, F.M., 1956: Octocorallia. In R.C. Moore (ed.): Treatise on inverte- brate paleontology, part F, Coelenterata, F166–F231. Geological Society of America and University of Kansas Press, Lawrence. Biogeography Bengtson, S., 1986: Siliceous microfossils from the Upper Cambrian of Queensland. Alcheringa 10, 195–216. doi:10.1080/03115518608619155 When considered together with the earlier studies by Peel Bengtson, S., 2004: Early skeletal fossils. In J.H. Lipps & B.M. Waggoner (2017a, 2018), the present account documents a substantial (eds.): Neoproterozoic-Cambrian biological revolutions,67–77. increase in the known diversity of Cambrian sponge spicules Paleontological Society Papers. Bengtson, S., 2005: Mineralized skeletons and early animal evolution. In D. from Laurentia, albeit just from the Greenland sector. From E.G. Briggs (ed.): Evolving form and function: fossils and development, a biogeographic point of view, however, the Laurentian assem- 101–124. Peabody Museum of Natural History, New Haven, Connecticut. blages confirm the wide distribution of many of the spicule Bengtson, S., Conway Morris, S., Cooper, B.J., Jell, P.A., & Runnegar, B. morphotypes between Cambrian palaeocontinents (Fig. 11). N., 1990: Early Cambrian fossils from South Australia. Memoirs of the – The similarity with Drumian and Guzhangian assemblages Association of Australasian Palaeontologists 9, 1 364. Beresi, M.S., 2003: Cambrian sponge spicules and chancelloriid sclerites described from the Georgina Basin by Bengtson (1986)and from the Argentine Precordillera: a review. Geologica Acta 1, 73–84. Mehl (1998) is particularly striking and similar assemblages Beresi, M.S., Aceñolaza, G., & Nieva, S., 2006: Cambro-Ordovician sponges can be traced from Australia, through South China and and spicule assemblages from Northwest Argetina: new data from the Siberia, into Laurentia. Collectively, these occurrences are tro- siliciclastic platforms of western Gondwana. Neues Jahrbuch für Geologie – pical, but this distribution is probably strongly influenced by und Paläontologie Monatshefte, 2006(7)403 420. Blaker, M.R. & Peel, J.S., 1997: Lower Cambrian trilobites from North a methodology focused on carbonate rich sequences. It should Greenland. Meddelelser om Grønland, Geoscience 35, 1–145. be noted, however, that assemblages from Siberia described by Botting, J.P. & Butterfield, N.J. 2005: Reconstructing early sponge rela- Panasenko (1998) and Sugai et al. (2004) are derived from cherts tionships by using the Burgess Shale fossil Eiffelia globosa, Walcott. in siliciclastic sediments. Furthermore, while sponge spicules are Proceedings of the National Academy of Sciences 102, 1554–1559. very widely distributed, the scarcity of published descriptions of Botting, J.P., Cárdenas, P., & Peel, J.S., 2015: A crown group demos- ponge from the early Cambrian Sirius Passet Biota, North Greenland. assemblages promotes caution. Published Cambrian spicule Palaeontology 58, 35–43. doi:10.1111/pala.12133 records from high latitudes may show lower diversity but pre- Botting, J.P. & Muir, L.A., 2013: Spicule structure and affinities of the servational and lithological factors are evident (Beresi 2003; Late Ordovician hexactinellid-like sponge Cyathophycus loydelli from Beresi et al. 2006;Castellanietal.2012;Muiretal.2013). the Llanfawr Mudstones Lagerstätte, Wales. Lethaia 46, 454–469. As regards articulated sponges, Carrera & Botting (2008), Botting, J.P. & Muir, L.A., 2018: Early sponge evolution: A review and phylogenetic framework. Palaeoworld 27, 1–29. doi:10.1016/j. Muir et al. (2013) and Botting & Muir (2018, in press)noted palwor.2017.07.001 that the Cambrian record is intermittent, with that from the late Botting, J.P. & Muir, L.A., in press: Dispersal and endemic diversifica- Cambrian (Furongian Series) being especially poor. Furongian tion: differences in non-lithistid spiculate sponge faunas between the spicules are not known from Greenland but have been described Cambrian Explosion and the GOBE. Palaeoworld. doi:10.1016/j. from elsewhere in Laurentia by Rigby (1975)andPratt(2002). palwor.2018.03.002 Articulated sponges occur mainly in offshore siliciclastic envir- Botting, J.P., Muir, L.A., & Lin, J.P., 2013: Relationships of the Cambrian – onments and are well represented in lower and middle Protomonaxonida (Porifera). Palaeontologia Electronica 16, 9A, 1 23. Botting, J.P. & Peel, J.S., 2016: Early Cambrian sponges of the Sirius Cambrian lagerstätten, notably from Laurentia and South Passet biota, North Greenland. Papers in Palaeontology 2, 463–487. China. However, faunas are often distributed between lower doi:10.1002/spp2.1048 and higher latitudes without particular palaeogeographic restric- Botting, J.P., Zhang, Y., & Muir, L.A., 2017: Discovery of missing link tion, but calcareans and some lithistids may show a tropical between demosponges and hexactinellids confirms palaeontological fi pattern (Muir et al. 2013; Botting & Muir 2018, in press). model of sponge evolution. Scienti c Reports 7, 5286. doi:10.1038/ s41598-017-05604-6 Bowerbank, J.S., 1864: A monograph of the British Spongidae, Volume 1. Acknowledgments The Ray Society, London. 290. Brasier, M., 1984: Microfossils and small shelly fossils from the Fieldwork was undertaken as part of the North Greenland Project (1978- Lower Cambrian Hyolithes Limedstone at Nuneaton, English – 80, 1984-85) organised by the Geological Survey of Greenland (GGU). Midlands. Geological Magazine 121, 229 253. doi:10.1017/ Scanning electron microscope facilities were provided by the Microscopy S0016756800028296 and Imaging unit, Evolutionary Biology Centre, Uppsala University. Brock, G.A. & Cooper, B.J., 1993: Shelly fossils from the early Cambrian Artem Kouchinsky and Joseph P. Botting reviewed the manuscript and (Toyonian) Wirrealpa, Aroona Creek, and Ramsay limestones of – I am particularly grateful to the latter for many valuable comments. South Australia. Journal of Paleontology 67, 758 787. doi:10.1017/ S0022336000037045 Buslov, M.M., Saphonova, I.Y., Watanabe, T., Obut, O.T., Fujiwara, Y., Data archiving statement Iwata, K., Semakov, N.N., Sugai, Y., Smirnova, L.V., & Kazansky, A. Y., 2001: Evolution of the Paleo-Asian Ocean (Altai-Sayan Region, This published work and the nomenclatural acts it contains have been central Asia) and collision of possible Gondwana-derived terranes registered in ZooBank: with the southern marginal part of the Siberian continent. http://zoobank.org./References/16B0FD52-721E-4572-86F8-5AAFFC5426B8. Geosciences Journal 5, 203–222. doi:10.1007/BF02910304 GFF 159

Buslov, M.M., Sennikov, N.V., Iwata, K., Zybin, V.A., Obut, O.T., Gray, J.E. 1867: Notes on the arrangement of sponges, with the descriptions Gusev, N.I., & Shokal’skii, S.P., 1998: New data on the structures of some new genera. Proceedings Zoological Society London, Lit. IV, and age of olistostromal and sand-silt rock masses of the Gornyi 492–558. Altai Series in the southeast of Gorny Altai Anui-Chuya zone. Han, J., Zhang, X., & Komiya, T., 2016: Integrated evolution of cnidarians Geologiya i Geofyziki 39, 789–798. [in Russian]. and oceanic geochemistry before and during the Cambrian explosion. In Carrera, M.O. & Botting, J.P., 2008: Evolutionary history of Cambrian S. Goffredo & Z. Dubinsky (eds.):The Cnidaria, past, present and future: spiculate sponges: implications for the Cambrian evolutionary fauna. the world of Medusa and her sisters,15–29. Springer, Switzerland. Palaios 23, 124–138. doi:10.2110/palo.2006.p06-089r Harvey, T.H.P., 2010: Carbonaceous preservation of Cambrian hex- Carrera, M.O. & Maletz, J., 2014: Ordovician sponge spicules from actinellid sponges. Biology Letters 6, 834–837. doi:10.1098/ Spitsbergen, Nevada and Newfoundland: new evidence for hex- rsbl.2010.0377 actinellid and demosponge early diversification. Journal of Hessler, R.R., 1970: The Desmosomatidae (Isopoda: asellota) of the Gay Systematic Palaeontology 12, 961–981. doi:10.1080/ Head – Bermuda Transect. Bulletin of the Scripps Institution of 14772019.2013.846946 Oceanography 15, 1–185. Castellani, C., Maas, A., Haug, C., Haug, J.T., & Waloszek, D., 2012: Heuse, T., Blumenstagel, H., Elicki, O., Geyer, G., Hansch, W., Maletz, J., Isolated sponge spicules from the late Cambrian Alum Shale Sarmiento, G.N., & Weyer, D., 2010: Biostratigraphy – the faunal Formation (‘Orsten’ nodules) of Sweden. Bulletin of Geosciences 87, province of the southern margin of the Rheic Ocean. In 443–460. doi:10.3140/bull.geosci.1328 U. Linneman & R.L. Romer (eds.): Pre-Mesozoic geology of Saxo- Chang, S., Feng, Q., Clausen, S., & Zhang, L., 2017: Sponge spicules from Thuringia – from the Cadomian active margin to the Variscan the lower Cambrian in the Yanjiahe Formation, south China: the Orogen,99–170. Schweitzerbart, Stuttgart. [Elicki, O. & Geyer, G., earliest biomineralizing sponge record. Palaeogeography, Cambrian biostratigraphy, 103–114]. Palaeoclimatology, Palaeoecology 474, 36–44. doi:10.1016/j. Higgins, A.K., Ineson, J.R., Peel, J.S., Surlyk, F., & Sønderholm, M., 1991: palaeo.2016.06.032 Lower Palaeozoic Franklinian Basin of North Greenland. Bulletin Chen, F. & Dong, X., 2008: Middle Cambrian highly modified sponge Grønlands Geologiske Undersøgelse 160, 71–139. spicules from western Hunan, South China. Acta Scientiarum Higgins, A.K., Soper, N.J., & Davis, N.C. 1992: Lower Palaeozoic outer Naturalium Universitatis Pekinensis 44, 883–889. shelf and slope lithostratigraphy, Franklinian basin, North Greenland. Debrenne, F. & Reitner, J., 2001: Sponges, cnidarians, and ctenophores. Grønlands Geologiske Undersøgelse Open File Report 92/8,1–57. In A.Y. Zhuravlev & R. Riding (eds.): The Ecology of the Cambrian Hinz, I., 1987: The Lower Cambrian microfauna of Comley and Radiation, 301–325. Columbia University Press, New York. Rushton, Shropshire/England. Palaeontographica A 198, 41–100. Ding, W. & Qian, Y., 1988: Late Sinian to early Cambrian small shelly Hu, J., Chen, Z., Xue, Y., Wang, J., Wang, J., & Yuan, X., 2002 : Sponge fossils from Yangjiaping, Shimen, Hunan. Acta Micropalaeontologia spicules in Early Cambrian Hetang Formation, Xiuning, southern Sinica 5, 39–55. Anhui. Acta Micropalaeontologica Sinica 19, 53–62. Dong, X. & Knoll, A.H., 1996: Middle and Late Cambrian sponge Ineson, J.R., 1988: Lithostratigraphy and depositional setting of the spicules from Hunan, China. Journal of Paleontology 70, 173–184. Holm Dal Formation (Middle Cambrian), central North Greenland. doi:10.1017/S0022336000023271 Meddelelser om Grønland, Geoscience 20, 9–21. Doweld, A.B., 2016: Kuonamia, a new generic replacement name for Ineson, J.R. & Peel, J.S., 1997: Cambrian shelf stratigraphy of North Disparella Fedorov and Pereladov 1987 (Porifera: hexactinellida) non Greenland. Geology of Greenland Survey Bulletin 173, 1–120. Hessler 1970 (Crustacea: isopoda). Research & Reviews: Journal of Ineson, J.R. & Peel, J.S., 2011: Geology and depositional setting of the Sirius Zoological Sciences 4, 19. Passet Lagerstätte (early Cambrian) of North Greenland. Canadian Duan, C., 1986: Small shelly fossils from the lower Cambrian Xihaoping Journal of Earth Sciences 48, 1259–1281. doi:10.1139/e11-018 Formation in the Shennongjia District, Hubei Province – hyoliths and Kouchinsky, A., Bengtson, S., Clausen, S., Gubanov, A., Malinky, J.A., & fossils skeletons of unknown affinities. Bulletin Tianjin Institute of Peel, J.S., 2011: A Middle Cambrian fauna of skeletal fossils from the Geology and Mineral Resources 7, 143–188. Kuonamka Formation, northern Siberia. Alcheringa 35, 123–189. Elicki, O., 1994: Lower Cambrian carbonates from eastern Germany: doi:10.1080/03115518.2010.496529 palaeontology, stratigraphy and palaeogeography. Neues Jahrbuch Kouchinsky, A., Bengtson, S., Clausen, S., & Vendrasco, M.J., 2015:Anearly für Geologie und Paläontologie Abhandlungen 191, 69–93. Cambrian fauna of skeletal fossils from the Emyaksin Formation, northern Elicki, O., 2011: First skeletal microfauna from Cambrian Series 3 of the Siberia. Acta Palaeontologica Polonica 60, 421–512. Jordan Rift Valley (Middle East). Memoirs of the Association of Kozur, H., 1991: Erster Nachweis von Polyactinellidae Mostler Australasian Palaeontologists 42, 153–173. (Calcispongea, Porifera) im Perm und Revision der Gattung Fedorov, A.B. & Peredalov, V.S., 1987: Spicules of siliceous sponges from Phobetractinia Reif. Zeitschrift für Geologische Wissenschaften 19, the Kuonamka Formation of north-eastern Siberian Platform [in 585–591. Russian]. In S.P. Bulynnikova & I.G. Klimova (eds.): Novye vidy Lee, B., 2006: Skeletal microfossils from the lower and middle Cambrian drevnih rastenij i bespozvonocnych fanerozoa Sibiri,36–46. of southwestern Mungyeong, Korea. Journal of the Paleontological SNIGGIMS, Novosibirsk. Society of Korea 22, 293–303. Geyer, G. & Peel, J.S., 2011: The Henson Gletscher Formation, North Mao, Y., Li, X., Lin, J., Muir, L.A., & Botting, J.P., 2013: A fossil Greenland, and its bearing on the global Cambrian Series 2–Series 3 assemblage of sponge spicules from the lower Kaili formation boundary. Bulletin of Geosciences 86, 465–534. doi:10.3140/bull. (Cambrian Series 2), South China. Acta Geologica Sinica 87, 883–884. geosci.1252 McMenamin, M.A.S., 2008: Early Cambrian sponge spicules from the Cerro Geyer, G. & Peel, J.S., 2017: Middle Cambrian fossils from the Clemente and Cerro Rajón, Sonora, México. Geologica Acta 6, 363–367. Ekspedition Bræ Formation of North Greenland, with a reappraisal Mehl, D., 1996: Phylogenie und Evolutionsökologie der Hexactinellida of the genus Elrathina (Trilobita, Ptychopariida). Journal of (Porifera) im Paläozoikum. Geologische Paläontologische Mitteilungen Paleontology 91, 265–293. doi:10.1017/jpa.2016.152 der Universität Innsbruck 4, 1–55. Goldberg, W.M. & Benayahu, Y., 1987: Spicule formation in the Mehl, D., 1998: Porifera and Chancelloriidae from the Middle Cambrian Gorgonian coral Pseudoplexaura flagellosa. 1: demonstrationof intra- of the Georgina Basin, Australia. Palaeontology 41, 1153–1182. cellular and extracellular growth and the effect of ruthenium red during Mehl, D. & Erdtman, B.-D., 1994: Sanshapentella dapingi n. gen. n. sp. – decalcification. Bulletin of Marine Science 40, 287–303. a new hexactinellid sponge from the Early Cambrian (Tommotian) of Goryanskii, V. Yu., 1977: First discovery of sponge remains from the China.). Berliner Geowissenschaften Abhandlungen (E) 13, 315–319. Lower Cambrian in eastern Siberia. Ezhdegodnik Vsesoyuznogo Mehl, D. & Lehnert, O., 1997: Cambro-Ordovician sponge spicule Paleontologicheskogo Obschestva 20, 274–278. [In Russian]. assemblages in the Ordovician of the Argentine Precordillera and Grant, R.E., 1836: Animal kingdom. In R.B. Todd (ed.): The cyclopedia of paleoenvironmental ties. Neues Jahrbuch für Geologie und anatomy and physiology, 107–118. Gilbert and Piper, London. Paläontologie Abhandlungen 204, 221-246. 160 J. S. PEEL

Missarzhevsky, V.V. & Mambetov, A.M., 1981: Stratigraphy and fauna of Mountains, northwestern Canada. Journal of Paleontology 76, the Precambrian–Cambrian boundary beds in Malyi Karatau. AN 565–569. doi:10.1017/S0022336000037380 SSSR Trudy Geologicheskogo Instituta 326, 1–92. [in Russian]. Reid, R.E.H., 1968: Hyalostelia smithii (Young & Young) and the sponge Mostler, H., 1985: Neue heteractinide Spongien (Calcispongia) aus dem genus Hyalostelia Zittel (Class Hexactinellida). Journal of Paleontology Unter- und Mitterlkambrium Südwestsardinien. Berichte des natur- 42, 1243–1248. wissenschaftlichen-medizinischen Verein Innsbruck 72, 7–32. Reif, W.E., 1968: Schwammreste aus dem oberen Ordovizium von Mostler, H., 1996: Polyactinellide Schwämme, eine auf das Paläzoikum Estland und Schweden. Neues Jahrbuch für Geologie und beschränkte Calcispongien-Gruppe. Geologische Paläontologische Paläontologie, Monatshefte 12, 733–744. Mitteilungen der Universität Innsbruck 21, 223–243. Reitner, J. & Mehl, D., 1995: Early Palaeozoic diversification of sponges: Mostler, H. & Mosleh-Yazdi, A., 1976: Neue Poriferen aus oberkambrischen new data and evidences. Geologische Paläontologische Mitteilungen der Gesteinen der Milaformationen im Elburzgebirge (Iran). Geologische Universität Innsbruck 20, 335–347. Paläontologische Mitteilungen der Universität Innsbruck 5, 1–36. Rhebergen, F. & Botting, J.P., 2014: A new Silurian (Llandovery, Muir, L.A., Botting, J.P., Carrera, M.G., & Beresi, M., 2013: Cambrian, Telychian) sponge assemblage from Gotland, Sweden. Fossils and Ordovician and Silurian non-stromatoporoid Porifera. In D.A. Strata 60, 1–88. T. Harper & T. Servais (eds.): Early Palaeozoic biogeography and Rigby, J.K., 1975: Some unusual hexactinellid sponge spicules from the palaeogeography,81–95. Geological Society, London. Memoirs 38 Cambrian Wilberns Formation of Texas. Journal of Paleontology 49, Müller, K.J. & Hinz, I., 1991: Upper Cambrian conodonts from Sweden. 412–415. Fossils and Strata 28, 1–153. Rigby, J.K., 1978: Porifera of the Middle Cambrian Wheeler Shale, from Müller,W.E.G.,Li,J.,Schröder,H.C.,Qiao,L.,&Wang,X.,2007:Theunique the Wheeler amphitheater, House Range, in western Utah. Journal of skeleton of siliceous sponges (Porifera; Hexactinellida and Demospongiae) Paleontology 52, 1325–1345. that evolved from the Urmetazoa during the Proterozoic: a review. Rigby, J.K., 1983: Sponges of the Middle Cambrian Marjum Limestone Biogeosciences 4, 219–232. doi:10.5194/bg-4-219-2007 of the House Range and Drum Mountains of western Millard County, Nazarov, B.B. & Popov, L.Y., 1976: Radiolaryii, bezzamkovye brakhiopody Utah. Journal of Paleontology 57, 240–270. i organizmy neyasnogo sistematicheskogo polozheniya iz srednego ordo- Rigby, J.K., 1986: Sponges of the Burgess Shale (Middle Cambrian), vika vostochnogo Kazakhstana. Paleontologicheskii Zhurnal. 1976 (4), British Columbia. Palaeontolographica Canadiana 2, 1–105. 33–42. Rigby, J.K., Church, S.B., & Anderson, N.J., 2010: Middle Cambrian Özdikmen, H., 2009: Substitute names for eight sponge genus group sponges from the Drum Mountains and House Range in western names (Porifera). Munis Entomology & Zoology 4, 212–218. Utah. Journal of Paleontology 84, 66–78. doi:10.1666/08-046.1 Panasenko, E.S. 1998: New unusual microfossils from pelagic silicon Rigby, J.K. & Collins, D., 2004: Sponges of the Middle Cambrian Burgess deposits of the Cambria. Dalnevostochnij geologiceskij institut, Shale and Stephen Formations, British Columbia. Royal Ontario Rossiya Akademia Nauk, Vladivostok.[FEGI]. http://www.fegi.ru/pri Museum Contributions in Science 1, 1–155. morye/geology/panas.htm [in Russian]. Robison, R.A. 1984: Cambrian Agnostida of North America and Peel, J.S. (ed.), 1988: Stratigraphy and palaeontology of the Holm Dal Greenland; part 1, Ptychagnostidae. Paleontological Contributions of Formation (late Middle Cambrian), central North Greenland. In the University of Kansas, Paper 109,1–59. Meddelelser om Grønland Geoscience Vol. 20, 1–168. Robison, R.A., 1988: Trilobites of the Holm Dal Formation (late Middle Peel, J.S., (ed.), 1994: Palaeontology, stratigraphy and environmental Cambrian), central North Greenland. Meddelelser om Grønland setting of Middle Cambrian outer shelf deposits, North Greenland. Geoscience 20, 23–103. Grønlands Geologiske Undersøgelse Bulletin 169, 1–155. Rozanov, A.Y. & Zhuravlev, A.Y., 1992: The Lower Cambrian fossil record Peel, J.S., 2017a: First records from Laurentia of some middle Cambrian of the Soviet Union. In J.H. Lipps & P.W. Signor (eds.): Origin and (Series 3) sponge spicules. Alcheringa 41, 306–314. doi:10.1080/ evolution of the Metazoa, 205–282. Plenum Press, New York. 03115518.2017.1282983 Sdzuy, K., 1969: Unter- und mittelkambrische Porifera (Chancellorida Peel, J.S., 2017b: The oldest pelmatozoan encrusted hardground and und Hexactinellida). Paläontologische Zeitung 43, 115–147. holdfasts from Laurentia (Cambrian Series 2–3). GFF 139, 195–204. doi:10.1007/BF02987647 doi:10.1080/11035897.2017.1347196 Sepkoski, J., 2002: A compendium of fossil marine animal genera. Peel, J.S., 2018: Sponge spicules from the Holm Dal Formation Bulletins of American Paleontology 363, 1–560. (Cambrian Series 3, Guzhangian) of North Greenland (Laurentia). Shabanov, Y.Y., Astashkin, V.A., Pegel, T.V., Egorova, L.I., Zhuravleva, I.T., GFF 140, 306–317. doi:10.1080/11035897.2018.1479444 Pelman, Y.L., Sundukov, V.M., Stepanova, M.V., Sukhov, S.S., Peel, J.S. & Ineson, J.R., 2011: The Sirius Passet Lagerstätte (early Cambrian) Fedorov, A.B., Shiskin, B.B., Vaganova, N.V., Ermak, V.I., of North Greenland. Palaeontographica Canadiana 31, 109–118. Ryabukha, K.V., Yadrenkina, A.G., Abaimova, G.P., Lopushinskaya, T. Peel, J.S. & Streng, M., 2015: A new middle Cambrian bradoriid arthro- V., Sychev, O.V., & Moskalenko, T.A., 1987: Nizhnii paleozoi yugo- pod from Greenland and western Canada. Journal of Paleontology 89, zapadnogo sklona Anabarskoi anteklizy (pro materialum bureniya). 96–102. doi:10.1017/jpa.2014.8 Nauka, Novosibirsk. 206. [in Russian]. Peel, J.S., Streng, M., Geyer, G., Kouchinsky, A., & Skovsted, C.B., 2016: Skovsted, C.B., 2006: Small shelly fauna from the upper lower Cambrian Ovatoryctocara granulata assemblage (Cambrian Series 2–Series 3 Bastion and Ella Island formations, North-East Greenland. Journal of boundary) of Løndal, North Greenland. Australasian Palaeontological Paleontology 80, 1087–1112. doi:10.1666/0022-3360(2006)80[1087: Memoirs 49, 241–282. SSFFTU]2.0.CO;2 Peel,J.S.&Willman,S.,2018: The Buen Formation (Cambrian Series 2) Biota Skovsted, C.B. & Peel, J.S., 2010: Small shelly fossils from the basal of North Greenland. Papers in Palaeontology 4, 381–432. doi:10.1002/ Kinzers Formation (Cambrian series 2, stage 4) of Pennsylvania. spp2.1112 Journal of Paleontology 84, 754–762. doi:10.1017/S00223 Pisera, A., 2003: What can we learn about siliceous sponges from 36000058467 palaeontology. Bollettino dei Musei e degli Instituti Biologici della Sugai,Y.,Iwata,K.,Sennikov,N.V.,Obut,O.T.,&Khlebnikova,T.V.,2004: (R.) Universita di Genova 68, 55–69. Sponge spicules from the lower Cambrian of Bateny Ridge and Gorny Altai Pisera, A., 2006: Palaeontology of sponges — a review. Canadian Journal (Bagrad and Shashkunar formations). Novosti paleontologii i stratigrafii. of Zoology 84, 242–261. doi:10.1139/z05-169 Pridozhenyie k Zhurnaly Geologiya i Geofyzika 6, 59–73. Pittau, P., Cotza, F., & Del Rio, M., 2003: Early Silurian siliceous sponge Torsvik, T.H. & Cocks, L.R.M., 2017: Earth history and palaeogeography. spicule assemblages from the Sardinia Hercynian Chain (Italy). Cambridge U.P. Cambridge, 332. Bollettino della Società Paleontologica Italiana 42, 225–239. Van Kempen, T.M.G., 1990: On the oldest tetraxon megascleres. In Pratt, B.R., 2002: Occurrence of the siliceous sponge spicule Konyrium K. Rützler (ed.): New perspectives in sponge biology,9–16. (Hexactinellida) in the Upper Cambrian of the MacKenzie Smithsonian Institution Press, Washington D.C. GFF 161

Vasilieva, N.I., 1998: Small shelly fossils and Lower Cambrian biostrati- of sponges. Palaeogeography, Palaeoclimatology, Palaeoecology 220, graphy of the Siberian Platform. Trudy VNIGRI, Skt Petersburg. 89–117. doi:10.1016/j.palaeo.2002.02.001 1–139. [in Russian]. Yang, X., Zhao, Y., Zhu, M., Wu, W., & Yang, K., 2010:Sponge Walcott, C.D., 1920: Cambrian geology and paleontology IV: 6 – Middle spicules from the Jiumenchong and Zhalagou Formations Cambrian Spongiae. Smithsonian Miscellaneous Collections 67, 261–364. (Cambrian) of Guizhou. Acta Micropalaeontologica Sinica 27, Webby, B.D. & Trotter, J., 1993: Ordovician sponge spicules from New 243–252. South Wales, Australia. Journal of Paleontology 67, 28–41. Zhang, X. & Pratt, B.R., 1994: New and extraordinary sponge spicule doi:10.1017/S0022336000021144 assemblage from China. Geology 22, 43–46. doi:10.1130/0091-7613- Wrona, R., 2004: Cambrian microfossils from glacial erratics of King (1994)022<0043:NAEECS>2.3.CO;2 George Island, Antarctica. Acta Palaeontologica Polonica 49, 13–56. Zhuravlev, A.Y. & Leguta, A.V., 2005:Sponges.In A.G. Ponomarenko Xiao, S., Hu, J., Yuan, X., Parsley, R.L., & Cao, R., 2005: Articulated (ed.): Unikal’niye Sinskiye mestonakhozhdeniya rannekembriyskikh sponges from the lower Cambrian Hetang formation in southern organizmov (Sibirskaya platforma),42–54.Nauka,Moscow.Trudy Anhui, South China: their age and implications for the early evolution Paleontologicheskogo Instituta 284, 1‒144 [in Russian].