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Home , Cambrian Stage 4, Crepicephalus, Drumian, Furongian, Guzhangian, Jiangshanian

[Papers in Palaeontology, 2020, pp. 1–22]

LINGULIFORM FROM THE () KARPINSK FORMATION OF NOVAYA ZEMLYA by LARS E. HOLMER1,2 , LEONID E. POPOV3 , MANSOUREH GHOBADI POUR2,3,4 ,INNAA.KLISHEVICH5,YUELIANG1 and ZHIFEI ZHANG1 1State Key Laboratory of Continental Dynamics, Shaanxi Key laboratory of Early Life & Environments, & Department of , Northwest University, Xi’an, 710069, China; [email protected], [email protected], [email protected] 2Department of Earth Sciences, Palaeobiology, Uppsala, SE-752 36, Sweden; [email protected] 3Department of Geology, National Museum of Wales, Cathays Park, Cardiff, CF10 3NP, UK; [email protected] 4Department of Geology, Faculty of Sciences, Golestan University, Gorgan, Iran 5Department of Geology, Institute of Earth Sciences St Petersburg State University, St Petersburg, Russia; [email protected]@yandex.ru

Typescript received 24 September 2019; accepted in revised form 23 January 2020

Abstract: A moderately diverse assemblage of micromor- was completed only at the end of the pelagic and phic linguliform brachiopods, including Tapuritreta gri- shows that they did have direct development, which is bovensis sp. nov., Wahwahlingula? pankovensis sp. nov., characteristic of all recent lingulides. The biogeographical Acrothele sp., Anabolotreta? sp., Orbithele? sp. and Stilp- signature of the new Cambrian fauna from notreta sp., is for the first time described from the Cam- Novaya Zemlya is discussed, and the new fauna gives new brian Karpinsk Formation (, Guzhangian) of information on the poorly known Cambrian margins of the the South Island of Novaya Zemlya. The morphology of Baltica palaeocontinent. the metamorphic shell in Acrothele and Wahwahlingula? suggest that both taxa had a single pair of larval setal sacs, Key words: Brachiopoda, Cambrian, Novaya Zemlya, tax- similar to the recent discinids, but their metamorphosis onomy, ontogeny, biogeography.

THE first evidence of Cambrian deposits on the South Anabolotreta? glabra Streng & Holmer, 2006, Wahwahlin- Island of Novaya Zemlya was obtained by a Norwegian gula? pankovensis sp. nov., Orbithele? sp., Stilpnotreta sp. expedition in 1921 (Holtedahl 1922). Subsequently the cf. Stilpnotreta minuta Holmer et al., 2001, Tapuritreta Cambrian outcrop areas along the west coast of Novaya gribovensis sp. nov. and Treptotreta jucunda Henderson & Zemlya were revisited and sampled occasionally by Rus- MacKinnon, 1981. With the exception of Anabolotreta, sian geologists between 1925 and 1934; they were also the which was previously documented by Popov (1985) from subject of a small-scale geological mapping project the lower part of the Karpinsk Formation at the south between 1948 and 1951 (Romanovich 1970). The first sys- coast of Gribovaya Bay, all other taxa were not known tematic study of the Cambrian geology and palaeontology previously from Novaya Zemlya. of the Novaya Zemlya Archipelago was carried out in The described linguliform brachiopods were extracted 1979–1983, by research teams from the Arctic and Antarc- from black argillaceous dissolved in dilute tic Research Institute, St Petersburg, in connection with (10%), buffered acetic acid. The brachiopod specimens an extensive survey of Cambrian outcrops along the west were coated with gold palladium and then studied and coast of the North and South Island. The main results photographed under a JEOL JCM-5000 Neoscope scan- were published by Solovyev (1988), including a lithostrati- ning electron microscope installed in the Department of graphical subdivision and -based . Natural Sciences, National Museum of Wales, Cardiff. The new assemblage of linguliform brachiopods comes All figured and discussed material is housed in the from the lower part of the Karpinsk Formation, exposed Department of Natural Sciences, National Museum of on the eastern part of the Pan’kov Land peninsula Wales (NMW) under the accession number NMW (Fig. 1). It includes Acrothele sp., Anabolotreta? sp. cf. 2018.4G.

doi: 10.1002/spp2.1314 1 © 2020 The Authors. Papers in Palaeontology published by John Wiley & Sons Ltd on behalf of The Palaeontological Association This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. 2 PAPERS IN PALAEONTOLOGY

FIG. 1. A, simplified map of Novaya Zemlya, showing the location of the investigated area on the South Island. B, simplified geological map of north-western part of Novaya Zemlya South Island showing geographical distribution of Cambrian deposits and position of selected localities. 1, , alluvial deposits; 2, late Palaeozoic, unspecified; 3–5, Cambrian; 3, Karpinsk Formation (upper Guzhangian – ); 4, Snezhyye Gory Formation (upper – Guzhangian); 5, Astafievskaya Formation (unnamed –Wuliuan); 6, fossil localities; 7, marker horizons in late Palaeozoic deposits; 8, faults (prepared by IAK in Corel Draw 15; https://www.corel.com/en/products/coreldraw/). Colour online.

GEOLOGICAL AND GEOGRAPHICAL section of the Gribovaya Bay south coast, the Mendeleev SETTING contains several of Protagraulos, including P. priscus Matthew, 1895 in the lowermost part, suggesting The Cambrian deposits exposed along the west coast of Cambrian Stage 4 (Solovyev et al. 1986). The first bra- Novaya Zemlya are presently subdivided into: (1) the chiopods are known from the overlying unit (Popov 1984; Astafiyev Formation (unnamed Cambrian Stage 4–Wuli- fig. 2, sample 911/26), represented by Acrothyra sp. and uan); (2) the Snezhnyye Gory Formation (upper Wuli- Hadrotreta sp. Their co-occurrence with Acadoparadoxides uan – lower Guzhangian); and (3) the Karpinsk cf. oelandicus (Sjogren,€ 1872) suggests a Wuliuan age. Formation (upper Guzhangian – Jiangshanian; Fig. 2). The Snezhnyye Gory Formation (–Guzhan- On the South Island, the Astafiyev Formation (up to gian), up to 320 m thick, rests conformably on the Asta- 150 m thick in the type section of Gribovaya Bay) com- fiyev Formation. It is composed mainly of oligomict and prises mainly greenish grey–dark grey siltstone and argillite arkosic sandstones, with a minor amount of siltstones intercalations with subsidiary sandstones and lenses of and argillites. The age of the formation is mainly based argillaceous limestones. Its lower contact is not exposed or on occurrences of the , including faulted. On the North Island, the stratigraphical contact paradoxissimus (Wahlenberg, 1821) in the lower part. In with the underlying metamorphosed sediments, assigned to the upper part of the unit at the section on the south the Mendeleev Series (unnamed Cambrian Stage 4–Wuli- coast of Gribovaya Bay, it yielded the brachiopods uan) represents a nonconformity. The stratigraphically Acrothele cf. coriacea Linnarsson, 1876, Prototreta gri- older Lomonosov Series were dated by acritarchs as Edi- bovensis Popov, 1985 and Diraphora simplex Popov, 1985 acaran–basal Cambrian (Korago et al. 1984, 2004). At the (Popov 1985; Solovyev et al. 1986). HOLMER ET AL.: CAMBRIAN BRACHIOPODSFROMNOVAYAZEMLYA 3

FIG. 2. Lithostratigraphical and biostratigraphical subdivision of the Cambrian deposits, and stratigraphical logs of the brachiopod species on Novaya Zemlya South Island (after Popov 1984, 1985; Solovyev et al. 1986 and Solovyev 1988) and its improved correlation with the International Geochronological Scale of the Cambrian System. Stratigraphical position of brachiopod fossil localities is mainly after Popov (1984, 1985) and Solovyev et al. (1986).

The Karpinsk Formation (upper Guzhangian – Jiangsha- c. 10 km north-east of the Gribovaya Bay north coast and c. nian) is presently known only from the South Island, where 8 km east of the abandoned Pan’kov military campus it is exposed between the Bezymyannaya Bay area in the (Figs 1, 2). The age of the unit is based on trilobites docu- south and the Matochkin Shar Strait in the north (Solovyev mented by Walcott & Resser (1924) from Holtedahl’s (1922) et al. 1986). It rests conformably, or possibly with a slight localities 67y and 68b, and it is also based on the trilobites disconformity, on the Snezhnyye Gory Formation. The Kar- reported by Solovyev et al. (1986), as well as the bra- pinsk Formation consists of black argillites and siltstones chiopods reported by Popov (1984, 1985). The trilobite with subsidiary sandstone and beds, and is a total assemblage from the locality 67y, described by Walcott & of 220 m thick at Gribovaya Bay and more than 260 m in Resser (1924), contains mostly endemic species, but includes the east part of the Pan’kov Land peninsula. On the south also Agnostus pisiformis (Wahlenberg, 1818) (for details of coast of Gribovaya Bay, it is disconformably overlain by the the taxon authorship, see Rushton 1978), suggesting a late Blafyel’sk Formation. The linguliform microbra- Guzhangian age. It is possible that the fauna list from the chiopod assemblage described here was recovered from locality 67y given by Walcott & Resser (1924) is composite limestones sampled in 1980 by I. A. Solovyev from Locality and includes both late Guzhangian and taxa. Some 302/3 (approximate coordinates 73°30 N, 53°290 E). The species from the locality 67y assemblage were reported by samples come from the lower part of the Karpinsk Forma- Solovyev et al. (1986) and Solovyev (1988) from three low- tion located at the east side of the Pan’kov Land peninsula ermost informal units in the type section of the Astafiyev 4 PAPERS IN PALAEONTOLOGY

Formation, exposed along the south coast of the Gribovaya Holotype. NMW 2018.4G.3 (Fig. 3F, J; Ld = 2.64 mm, Bay (Solovyev 1988, text-fig. 2). W = 1.80 mm, Il = 0.48 mm, Iw = 1.67 mm, The trilobite assemblage from Holtedahl’s locality 68b, Pl = 0.32 mm, Vl = 1.77 mm), dorsal valve from Snezhnyye documented by Walcott & Resser (1924) contains Irvingella Gory Formation (Miaolingian, Guzhangian), locality 302/3, septentrionalis Walcott & Resser, 1924, which, according to Pan’kov Land peninsula, Novaya Zemlya, South Island. Palmer (1965), is the junior synonym of Irvingella major Ulrich & Resser in Walcott, 1924. The latter occurrence sug- Paratypes. Figured: NMW 2018.4G.2 (Fig. 3G; Lv = 2.84 mm, gests a Jiangshanian age for the uppermost Karpinsk Forma- W = 2.06 mm, Il = 0.54 mm, Iw = 1.32 mm, Pl = 0.34 mm), tion. The brachiopods documented by Walcott (1924) from NMW 2018.4G.7 (Fig. 3D), ventral valves; NMW 2018.4G.1 that locality, including ‘Lingulella’cf.desiderata Walcott, (Figs 3K, 4E), NMW 2018.4G.4 (Fig. 4A, B), NMW 2018.4G.6 1898, Billingsella holtedahli Walcott, 1924, Eoorthis sabus (Fig. 3C), NMW 2018.4G.9 (Figs 3L, 4D, F), NMW 2018.4G.10 Walcott, 1924, Huenellina triplicata Walcott, 1924 and (Fig. 3E, M), NMW 2018.4G.11, (Fig. 4G), NMW 2018.4G.12 (Fig. 3A), NMW 2018.4G.13 (Fig. 3H), dorsal valves. Other Ocnerorthis sp. cf. O. opius (Walcott, 1924) are the youngest specimens: NMW 2018.4G.15, 16, incomplete ventral valves; Cambrian () brachiopod assemblage yet docu- NMW 2018.4G.5, 8, 14, incomplete dorsal valves. Total four mented from the Novaya Zemlya Archipelago. ventral and 11 dorsal valves; all from locality 302/3.

Diagnosis. Shell dorsibiconvex, elongate suboval with SYSTEMATIC PALAEONTOLOGY maximum width at mid-length. Ventral pseudointerarea bisected by deep, narrow, subtriangular groove with steep By Lars E. Holmer, Leonid E. Popov, Mansoureh Ghobadi lateral slopes. Ventral propareas bisected by deep flexure Pour, Yue Liang and Zhifei Zhang lines into two subequal parts. Dorsal visceral area occu- pies about two-thirds of sagittal valve length. Brephic and The morphological terminology used here follows that of adult shell ornamented by densely packed, regular, hemi- Holmer & Popov (2000), while the terminology introduced spherical pits c. 1.8–2.1 lm across. by Zhang et al. (2018) is followed in describing the ontogeny.

Description. Shell dorsibiconvex, elongate, suboval, with maxi- Abbreviations. W, L, T, maximum width, length and height of the mum width at mid-length, c. 135–150% as long as it is wide, shell; Lv, Ld, maximum length of ventral and dorsal valve; Iw, Il, with maximum width at about mid-length. Anterior commissure maximum width and length of pseudointerarea; Pl, Pw, length and evenly rounded, rectimarginate. Ventral valve gently convex, width of median groove or pedicle groove; Cw, Cl, width, length of subacuminate, with high subtriangular almost orthocline pseu- cardinal muscle field; Sl, length of dorsal median septum; Vl, length dointerarea c. 30% as wide as it is long. Pedicle groove deep, of visceral area. narrow subtriangular, with steep lateral slopes, c. 12% of sagittal valve length. Ventral pseudointerareas raised above the valve floor, subdivided into two almost equal parts by the deep flexure Subphylum LINGULIFORMEA Williams et al., 1996 lines (Figs 3G, 4C). Class Gorjansky & Popov, 1985 Dorsal valve evenly convex in sagittal profile with maximum Order Waagen, 1885 height at about mid-length. Dorsal anterior margin evenly rounded. Superfamily LINGULOIDEA Menke, 1828 Dorsal pseudointerarea c. 30%aslongasitiswidemainlyoccupied Family ZHANATELLIDAE Koneva, 1986 by broad median groove lying along the valve floor. Dorsal propar- eas rudimentary, lacking flexure lines. Shell surface smooth with WAHWAHLINGULA Popov et al., 2002 faint growth lines and micro-ornament of faint, closely packed, reg- ular, hemispherical pits c. 1.8–2.1 lm in diameter, which cover all Type species. By original designation antiquissima the shell surface, including the brephic shell (Fig. 3B). Jeremejew, 1856. Ventral valve visceral area, muscle scars and mantle canals are not clearly expressed except for paired umbonal muscle scars located in weakly defined impressions anterolateral of the Wahwahlingula? pankovensis sp. nov. anterior termination of the pedicle groove. Dorsal visceral area weakly impressed, extended anteriorly as a pronounced median Figures 3, 4A–G tongue and terminated at about two-thirds sagittal valve length from the umbo. The latter occupied by a pair of elongate out- side lateral muscle scars (Fig. 3F, ol). A pair of weakly defined LSID. urn:lsid:zoobank.org:act:3E7C8A70-9B35-4DA9-A270- central muscle scars (Fig. 3F, ol) is situated at the anterior end 6A05FA83BC0C of indistinct muscle tracks in the centre of the valve. Posterolat- eral muscle fields (Fig. 3F, plm) representing combined scars of Derivation of name. After the occurrence on the Pan’kov the outside lateral, middle lateral and transmedian muscles Land peninsula. located at the outer margins of the visceral area in front of the HOLMER ET AL.: CAMBRIAN BRACHIOPODSFROMNOVAYAZEMLYA 5

FIG. 3. Wahwahlingula? pankovensis sp. nov.; Snezhnyye Gory Formation (Miaolingian, Guzhangian), locality 302/3, Pan’kov Land peninsula, Novaya Zemlya, South Island. A, NMW 2018.4G.12, incomplete dorsal valve exterior. B, I, NMW 2018.4G.8, dorsal valve exterior, dorsal and oblique lateral views. C, NMW 2018.4G.6, dorsal valve umbonal area. D, NMW 2018.4G.7, incomplete ventral valve exterior. E, M, NMW 2018.4G.10, dorsal valve interior, oblique side view of interior. F, J, NMW 2018.4G.3, holotype, dorsal valve interior, oblique side view of interior. G, NMW 2018.4G.2, ventral valve interior. H, NMW 2018.4G.13, incomplete dorsal valve interior. K, NMW 2018.4G.1, incomplete dorsal valve exterior. L, NMW 2018.4G.9, incomplete dorsal valve exterior. Abbreviations: cm, central muscle scars; fl, flexure lines; ol, outside lateral muscle scars; plm, posterolateral muscle fields. Scale bars represent: 100 lm(A–E, H, K–M); 200 lm (F, G, I, J). 6 PAPERS IN PALAEONTOLOGY HOLMER ET AL.: CAMBRIAN BRACHIOPODSFROMNOVAYAZEMLYA 7 propareas. Dorsal median ridge absent. Dorsal mantle canals by a much smaller size, a strongly elongate, suboval, rather than not clearly impressed. The valve floor in posterior half of both a subtriangular shell outline and the lack of a dorsal median valves is often covered with random scattered round-bottomed, ridge. Micro-ornamentation on the brephic shell of Wahwahlin- circular pits. gula antiquissima is yet to be observed.

Remarks on the ontogeny. No protegulum is recognizable at the Superfamily ACROTHELOIDEA Walcott & Schuchert in umbonal area of both valves. The dorsal brephic shell, c. 160 lm Walcott, 1908 wide, is recognized as a slightly swollen area bisected by a shal- Family ACROTHELIDAE Walcott & Schuchert in Walcott, low medial cleft (Figs 3C, 4A, F). A pair of lateral swellings on 1908 both sides of the cleft probably represent impressions of larval Subfamily ACROTHELINAE Walcott & Schuchert in Walcott, setal sacs. The dorsal brephic shell of Wahwahlingula? pankoven- 1908 sis is very similar to the brephic shell of Diencobolus as described by Ghobadi Pour et al. (2011), and both taxa have a pair of lat- Genus ACROTHELE Linnarsson, 1876 eral swellings, which are interpreted as impressions of larval setal sacks. The closest recent analogy can be found in Discinisca, Type species. By original designation, Acrothele coriacea Lin- which also retains a pair of larval setal sacs that disappear narsson, 1876. shortly after hatching (Luter€ 2001; Zhang et al. 2018, fig. 8), but the metamorphosis in Wahwahlingula? pankovensis was com- pleted somewhat later during a free swimming stage. Acrothele? sp. Remarks. Wahwahlingula? pankovensis is the oldest yet recorded Figures 5, 6E–H species of the genus and the only Miaolingian species of the genus yet known. In size, general shell shape and proportions it most 2005 Acrothele sp.; Gonzalez-Gomez, p. 35, fig. 5.1–5.11. resembles Wahwahlingula severensis Popov et al., 2002 from the House Limestone (Barn Canyon and Red Canyon members) and the lower part of Fillmore Formation () of the Ibex Material. NMW 2018.4G.17 (Fig. 5C), 20 (Fig. 5A, G), 21 area, Utah, USA, but it can be distinguished from the latter taxon (Fig. 5D–F, H), 22 (Fig. 5B), ventral valves; NMW 2018.4G.18 by a slightly more elongate shell, ventral propareas distinctly raised (Fig. 6E, F), 19 (Fig. 6G, H) dorsal valves. above the valve floor and subdivided into two almost equal parts by deep flexure lines, shorter dorsal visceral area not exceeding two- Remarks. All specimens are preserved as disarticulated incom- thirds valve length and the invariable absence of the dorsal median plete dorsal and ventral valves. The ventral valve is subconical ridge. In Wahwahlingula? pankovensis faint regular hemispherical with a slightly eccentric apex slightly elongate suboval pedicle pits cover the whole shell surface, while in Wahwahlingula severensis foramen c. 60 lm long and 50 lm wide, which is almost com- the brephic shell is covered by flat-bottomed and cross-cutting pits. pletely enclosed within the metamorphic shell (Fig. 5F, G). The Unlike Wahwahlingula kharbashi Ghobadi Pour et al., 2011, dorsal valve is almost flat with a slightly swollen umbonal region from the Tremadocian (Paltodus deltifer Biozone) of the eastern and rudimentary dorsal pseudointerarea. The ventral interior Alborz Mountains, Iran, and Wahwahlingula? emanuelensis lacks distinctive features, while the dorsal interior is insuffi- Brock & Holmer, 2004, from the Emanuel Formation () ciently known due to poor preservation. of Western Australia, Wahwahlingula? pankovensis has a pitted These shells have an unusual metamorphic shell morphology brephic shell, and an elongate suboval shell with maximum that distinguishes them from other species of Acrothele and the width at mid-length. In addition it completely lacks a dorsal subfamily Acrothelinae. The metamorphic shell is 390–490 lm median ridge, unlike Wahwahlingula kharbashi. wide, ornamented with unusually large hemispherical pits, vary- In comparison with Wahwahlingula antiquissima (Jeremejew, ing in diameter from 10 to 20 lm and separated by wide, flat- 1856), from the lower part of the Tosna Formation ( topped interspaces, and the metamorphic shell is outlined by a lindstromi Biozone), Wahwahlingula? pankovensis is characterized distinct halo (Fig. 5E–H). While a low median tubercle is

FIG. 4. All specimens from Snezhnyye Gory Formation (Miaolingian, Guzhangian), locality 302/3, Pan’kov Land peninsula, Novaya Zemlya, South Island. A–G, Wahwahlingula? pankovensis sp. nov.: A–B, NMW 2018.4G.4, umbonal area of dorsal valve showing brephic shell, enlarged shell surface showing pitted micro-ornament; C, NMW 2018.4G.2, oblique posterior view of ventral pseudoint- erarea; D, F, NMW 2018.4G.9, dorsal valve exterior, enlarged view of umbonal area showing brephic shell; E, NMW 2018.4G.1, enlarged view of umbonal area showing brephic shell; G, NMW 2018.4G.11, incomplete dorsal valve exterior. H–L, Orbithele sp.: H, L, NMW 2018.4G.23, incomplete ventral valve posterior view of pedicle foramen and metamorphic shell, ventral view; I, NMW 2018.4G.24, incomplete ventral valve exterior; J, NMW 2018.4G.26, incomplete ventral valve interior showing internal pedicle tube supported by short septum; K, NMW 2018.4G.25, incomplete ventral valve interior showing internal pedicle tube supported by short septum. Abbreviations: h, halo; ms, metamorphic shell; ss, larval setal sac impression. Scale bars represent: 20 lm (A); 10 lm (B); 100 lm(C–H); 200 lm(I–L). 8 PAPERS IN PALAEONTOLOGY

FIG. 5. Acrothele sp.; Snezhnyye Gory Formation (Miaolingian, Guzhangian), locality 302/3, Pan’kov Land peninsula, Novaya Zem- lya, South Island. A, G, NMW 2018.4G.20, incomplete ventral valve exterior, enlarged view of metamorphic shell. B, NMW 2018.4G.22, incomplete ventral valve interior. C, NMW 2018.4G.17, incomplete ventral valve interior. D–F, H, NMW 2018.4G.21, incomplete ventral valve exterior, enlarged view of metamorphic shell, surface ornament of postmetamorphic shell, pitted ornament of metamorphic shell. Scale bars represent: 200 lm(A–D); 50 lm (E, H); 100 lm (F, G).

present in front of the pedicle foramen, there is no sign of et al. 2018, fig. 8); however, unlike recent lingulides, the pres- tubercles or spines in the metamorphic shell (Fig. 5F, G). ence of a large metamorphic shell suggests that metamorphosis Instead a pair of inflated lobes diverges from the outer margin in the Acrothelidae was completed at the end of the prolonged of the pedicle foramen towards the anterior margin of the ven- pelagic stage; this shows that indirect development was still char- tral metamorphic shell. Only a single pair of spines is present in acteristic of the life cycle of these lingulides. front of the posterior margin of the dorsal metamorphic shell In contrast, other species of Acrothele are characterized by (Fig. 6E–H). The spine bases continue anteriorly as a pair of well-ordered, flat-based imprints varying in diameter from 3.2 divergent, anteriorly inflated lobes terminated close to the ante- to 9.5 lm, forming relatively open hexagonal arrays and sepa- rior margin of the metamorphic shell, but no distinct anterior rated by narrow, rounded walls (Williams 2003). Other charac- pair of tubercles is present (Fig. 6F, G). These dorsal lobes are teristic features of Acrothele are the presence of four short considered here as impressions of a single pair of larval setal spines on the dorsal metamorphic shell and a pair of promi- sacks, by analogy with recent discinids (e.g. Luter€ 2001; Zhang nent tubercles or short spines anterolateral of the pedicle HOLMER ET AL.: CAMBRIAN BRACHIOPODSFROMNOVAYAZEMLYA 9

FIG. 6. All specimens from Snezhnyye Gory Formation (Miaolingian, Guzhangian), locality 302/3, Pan’kov Land peninsula, Novaya Zemlya, South Island. A–D, Orbithele? sp.: A–B, D, NMW 2018.4G.29, enlarged micro-ornament of postmetamorphic shell, pitted micro-ornament of metamorphic shell, incomplete dorsal valve exterior; C, NMW 2018.4G.28, incomplete dorsal valve interior. E–H, Acrothele sp.: E–F, NMW 2018.4G.18, incomplete dorsal valve exterior, enlarged metamorphic shell; G–H, NMW 2018.4G.19, enlarged metamorphic shell, incomplete dorsal valve exterior. Abbreviations: h, halo; ms, metamorphic shell; pms, postmetamorphic shell; ss, larval setal sac impression. Scale bars represent: 100 lm (A, F, G); 20 lm (B); 500 lm (C, D); 200 lm (E, H). foramen in the ventral valve. This type of metamorphic shell designation is postponed due to insufficient material presently morphology is characteristic, in particular, of Acrothele coriacea available; however, very similar and most probably conspecific from the Miaolingian of Baltoscandia (Holmer & Popov 2000; shells were described as unnamed species of Acrothele by Gon- fig, 46.1), the type species of the genus; and of Acrothele vertex zalez-Gomez (2005), from the Val d ‘Homs Formation (Guz- Reed, 1910 from the Parahio Formation (Wuliuan, Orycto- hangian) of Montagne Noire at southern France. These shells cephalus salteri Biozone) of the Indian Himalayas (Popov et al. have a distinct similarity to the specimens from Novaya Zem- 2015; fig. 15D, I, L–P), which is probably the earliest formally lya in having a small pedicle foramen mainly within the meta- designated species of the genus. morphic shell, a lobate ventral metamorphic shell lacking While these acrothelid shells from Novaya Zemlya definitely spines, and a dorsal metamorphic shell with a single pair of represent a new, as yet unnamed taxon, a formal taxonomic spines, while the pitted micro-ornament looks almost identical. 10 PAPERS IN PALAEONTOLOGY

Genus Orbithele Sdzuy, 1955 Anabolotreta? sp. cf. Anabolotreta? glabra Streng & Holmer, 2006 Type species. By original designation Discina contraria Figure 7 Barrande, 1868. 1985 Anabolotreta sp.; Popov, p. 25; pl. 2, fig. 15.

Orbithele? sp. Figures 4H–L, 6A–D Material. NMW 2018.4G.35, 36, ventral valves, and NMW 2018.4G.37 dorsal valve. Material. Figured: NMW 2018.4G.23 (Fig. 4H, L), NMW 2018.4G.24 (Fig. 4I), NMW 2018.4G.25 (Fig. 4K), NMW Description. Shell slightly ventribiconvex, slightly transverse, sub- 2018.4G.26 (Fig. 4J), ventral valves; NMW 2018.4G.28 (Fig. 6C); oval to almost subcircular. Anterior commissure broadly NMW 2018.4G.29 (Fig. 6A, B, D) dorsal valves. Other speci- rounded, rectimarginate. Posterior commissure almost straight, mens: NMW 2018.4G.27, 30–34, ventral valves. Total 10 ventral c. 55% as wide as maximum shell width. Ventral valve low, and two dorsal valves; all from locality 302/3. moderately convex, with evenly convex sagittal profile anterior to the apex. Ventral pseudointerarea procline, bisected by a Remarks. All available specimens represent incomplete ventral weakly defined, shallow intertrough and indistinctly separated and dorsal valves. Nevertheless, they show a number of distinc- from the lateral sides of the valve. Pedicle foramen posteroven- tive morphological features, including a low conical ventral trally directed, small, subcircular, within the metamorphic shell. valve with subcentral apex, a strongly elongate pedicle foramen Dorsal valve sagittal profile gently and evenly convex. Dorsal crossing the outer boundary of the metamorphic shell bearing pseudointerarea low, occupied mainly by the broad, gently con- a pair of low umbonal spines, and finely pustulose ornament cave median groove, propareas rudimentary. Shell surface orna- on the postmetamorphic shell. These features can also be mented with faint, crowded growth lines and up to three strong observed in a number of late Miaolingian acrothelides provi- growth lamellae in the anterior half of the valves. sionally assigned to Orbithele (e.g. Zell & Rowell 1988; Percival Ventral interior with a strong, boss-like, posteriorly grooved & Kruse 2014), but there is no evidence of marginal spines apical process with undercut anterior slope and steep, subparallel characteristic of the Early Ordovician species of the genus lateral slopes, situated anterior to the internal foramen. Two including the type species. The ventral interior has a pedicle small circular umbonal muscle scars (apical pits) located antero- tube on the posterior slope of the valve, supported anteriorly laterally to the internal foramen and surrounded by faint ridges. by a short, blade-like median septum with a short rod in front Cardinal muscle fields slightly thickened, clearly defined, pos- of the internal foramen, which are among the most distinctive terolaterally located. Mantle canals not clearly impressed. Dorsal features of Orbithele (Popov & Holmer 1994). The dorsal valve interior with short, rounded, subtriangular, posterolaterally is characterized by having a marginal umbo and a low dorsal located cardinal muscle fields, and vestigial faint median ridge median ridge bisecting the visceral area (Fig. 6C). The meta- recognizable only in central part of the valve. No median but- morphic shell of Orbithele? sp. shows typical features first tress. described by Henderson (1974). It has a median tubercle in front of the pedicle foramen and a pair of short spines on the Remarks. In spite of the limited material available for study, the ventral valve (Fig. 4I, L). The dorsal metamorphic shell (pre- affinity of these shells to Anabolotreta? sp. cf. A. glabra is evident, served on a single specimen), is poorly preserved, but the pres- because they are characterized by a pedicle foramen enclosed ence of two pairs of short spines is evident. The pitted within the metamorphic shell, evenly convex anterior slope of the ornament of the metamorphic shell is identical to that ventral valve, concentric ornament with a few growth lamellae in described by Williams (2003) and is characterized by densely the anterior part of the shell, a prominent, medially grooved apical packed, irregular, flat-based imprints strongly variable in size process, and a broad and short dorsal median groove. However, (3.2–11 lm), separated by narrow, convex walls bearing occa- unlike the types from the lowermost Dunderberg Formation sional small hemispherical pits less than 1 lm across. Due to (Guzhangian, –Crepicephalus ) of Nevada the poor preservation of the shells from Novaya Zemlya the and the specimens from the Huaqiao Formation (Drumian–Guz- material is kept under open nomenclature. hangian, atavus–Linguagnostus reconditus trilobite zones) of section, in Province, South China, the shells from Novaya Zemlya have a vestigial dorsal median ridge Order ACROTRETIDA Kuhn, 1949 clearly expressed only in the central part of the valve and a pair of Superfamily ACROTRETOIDEA Schuchert, 1893 well-defined umbonal muscle scars (apical pits) anterolateral to Family ACROTRETIDAE Schuchert, 1893 the internal foramen. Therefore, their species attribution is con- Anabolotreta Rowell & Henderson, 1978 sidered here as provisional. In poor development of concentric growth lamellae, the specimens from Novaya Zemlya are similar Type species. By original designation, Anabolotreta tegula Rowell to Anabolotreta groenlandica Zell & Rowell, 1988, from the Guz- & Henderson, 1978. hangian part of the Holm Dal Formation of central North HOLMER ET AL.: CAMBRIAN BRACHIOPODS FROM NOVAYA ZEMLYA 11

FIG. 7. Anabolotreta? cf. glabra Streng & Holmer, 2006. All specimens from Snezhnyye Gory Formation (Miaolingian, Guzhangian), locality 302/3, Pan’kov Land peninsula, Novaya Zemlya, South Island. A, D, NMW 2018.4G.35 ventral valve exterior, oblique side view. B, E, NMW 2018.4G.36, ventral valve interior, oblique posterior view of exterior. C, F, NMW 2018.4G.37, dorsal valve interior, oblique posterolateral view of interior. Scale bars represent 100 lm.

Greenland, but can be readily distinguished in having a pedicle Formation (Miaolingian, Guzhangian), locality 302/3, Pan’kov foramen enclosed within the metamorphic shell, an evenly convex Land peninsula, Novaya Zemlya, South Island. (not concave) anterior slope of the ventral valve, a prominent, boss-like apical process with steep, subparallel lateral margins, Paratypes. Figured: NMW 2018.4G.41 (Fig. 8K, L), NMW small, as well as well-defined apical pits surrounded by ridges, and 2018.4G.42 (Fig. 8I, J), NMW 2018.4G.44 (L = 1.53, T = 0.80; a dorsal vestigial median ridge. A single dorsal valve briefly Fig. 8B), NMW 2018.4G.45 (Fig. 8M), NMW 2018.4G.46 described and illustrated by Popov (1985) from the Karpinsk For- (Fig. 8H), ventral valves and NMW 2018.4G.47 (Fig. 9H, K), mation of the Gribovaya Bay southern coast is closely similar and NMW 2018.4G.50 (Ld = 1.47 mm, W = 1.80; Fig. 9B, F, I), considered here as conspecific to the shells from the Pan’kov Land NMW 2018.4G.52 (Ld = 2.02 mm, W = 1.81 mm, peninsula. Iw = 1.00 mm, Pw = 0.47 mm, Sl = 1.23 mm; Cl = 0.53 mm, Cw = 82 mm; Fig. 9G, J); NMW 2018.4G.53 (Ld = 2.05 mm, W = 2.10 mm, Iw = 1.07 mm, Pw = 0.53 mm, Sl = 1.33 mm; Genus TAPURITRETA Popov et al., 2009 Cl = 0.61 mm, Cw = 1.02 mm; Fig. 9A, C, D). Other material: NMW 2018.4G.54.1–4, ventral valves; NMW 2018.4G.48, 49, 51, – Type species. By original designation Tapuritreta angusta Popov 54.5 15 dorsal valves. Total 10 ventral and 17 dorsal valves from et al., 2009. locality 302/3.

Diagnosis. Ventral valve between one half and two-thirds as high as long, with procline deltoid pseudointerarea, almost straight in Tapuritreta gribovensis sp. nov. sagittal profile and bisected by vestigial intertrough. Dorsal pseu- Figures 8, 9 dointerarea slightly exceeding half of maximum shell width. Dorsal cardinal muscle scars gently impressed less than one-third as long as the valve. LSID. urn:lsid:zoobank.org:act:6227614F-1EC5-4726-BBF3- F19A5E6AA148 Description. Shell strongly dorsibiconvex, subcircular, almost as long as wide, with maximum width at mid-length. Ventral valve Derivation of name. After Gribovaya Bay south of the type locality. conical, about two-thirds as high as long with a strongly flattened, distinctly procline deltoid interarea almost straight in sagittal profile Holotype. NMW 2018.4G.43 (Lv = 1.30 mm, W = 1.47 mm, (Fig. 8F). Intertrough very weakly defined mainly by slight deflec- T = 0.64 mm; Fig. 8E–G), ventral valve from Snezhnyye Gory tions of growth lines (Fig. 8G, I). Pedicle foramen small, circular, 12 PAPERS IN PALAEONTOLOGY

FIG. 8. Tapuritreta gribovensis sp. nov.; Snezhnyye Gory Formation (Miaolingian, Guzhangian), locality 302/3, Pan’kov Land penin- sula, Novaya Zemlya, South Island. A, L, NMW 2018.4G.45, ventral valve: lateral view of exterior, oblique posterior view of interior showing apical process and internal pedicle tube attached to the inner side of the posterior slope of the valve. B, NMW 2018.4G.44, oblique posterior view of interior showing internal foramen and apical process. C, G, NMW 2018.4G.46, oblique posterior views of ventral valve exterior, enlarged metamorphic shell. H, I, NMW 2018.4G.42, oblique posterior view of ventral valve exterior, enlarged metamorphic shell. D–F, NMW 2018.4G.43, holotype, ventral valve exterior: ventral, oblique lateral and posterior views. J–K, NMW 2018.4G.41, incomplete ventral valve interior, oblique lateral view of interior showing high, septiform apical process. Abbreviations: ap, apical process; ipt, internal pedicle tube; um, umbonal muscle scar. Scale bars represent: 200 lm (A, G, H, L); 500 lm (B, D–F, J, K); 50 lm (C, I). HOLMER ET AL.: CAMBRIAN BRACHIOPODS FROM NOVAYA ZEMLYA 13

FIG. 9. Tapuritreta gribovensis sp. nov.; Snezhnyye Gory Formation (Miaolingian, Guzhangian), locality 302/3, Pan’kov Land penin- sula, Novaya Zemlya, South Island. A, C, D, NMW 2018.4G.53, enlarged view of dorsal metamorphic shell, dorsal valve exterior, dorsal valve interior. B, F, I, NMW 2018.4G.50, enlarged view of dorsal metamorphic shell, dorsal valve exterior, oblique posterior view of exterior. E, NMW 2018.4G.52, dorsal valve interior. G, J, NMW 2018.4G.49, dorsal valve interior, oblique lateral view of interior. H, K, NMW 2018.4G.47, dorsal valve interior, oblique lateral view of interior of juvenile individual. Scale bars represent: 50 lm (A, B); 500 lm(C–G, I, J); 200 lm (H, K). 14 PAPERS IN PALAEONTOLOGY c. 50 lm in diameter, facing posteroventrally, enclosed within the Guzhangian) in their small size and in having a subcircular, metamorphic shell (Fig. 8J). Anterior slope of the ventral valve very slightly ventribiconvex shell with a vestigial deltoid pseudointer- gently convex to almost straight in sagittal profile. Dorsal valve area, low apical process in front of the internal opening of the sagittal profile very gently convex with maximum height between pedicle foramen, extending to the mid-valve, as well as in the the umbo and mid-length. Dorsal interarea orthocline, c. 50–55% as absence of distinct denticulation of a ventral valve posterior wide as the valve, with a broad shallow median groove almost half margin and a median buttress. However, the new material differs as wide as the pseudointerarea. Dorsal sulcus very shallow, originat- in the complete absence of a dorsal median ridge. All listed ing at the umbo, strongly widening towards the anterior margin. characters, except the shape of the apical process, clearly dis- Postmetamorphic shell smooth, with faint growth lines. Metamor- criminate the Novaya Zemlya specimens from Stilpnotreta phic shell, c. 180–200 lm wide, ornamented with faint, densely magna. Nevertheless, their species affiliation remains provisional packed, slightly irregular, hemispherical pits, c. 1–3 lm in diameter because of insufficient material available for study. (Fig. 9A, B). Ventral interior with weakly impressed cardinal muscle fields and high, septiform apical process undercut anteriorly (Fig. 8B) Treptotreta Henderson & MacKinnon, 1981 and enclosing the internal pedicle tube situated on the posterior slope of the valve (Fig. 8L). Ventral umbonal muscle scars gently impressed as a pair of small hemispherical pits, posterolateral of Type species. Treptotreta jucunda Henderson & MacKinnon, the internal foramen (Fig. 8B). Ventral mantle canals not 1981; middle Cambrian (Goniagnostus nathorsti Biozone), Mail- impressed. Dorsal interior with weakly impressed, suboval cardi- change Limestone, western Queensland, Australia. nal muscle fields divergent anterolaterally, terminated anteriorly at c. 27–29% valve length from the umbo. Dorsal medium sep- tum moderately high, blade-like, triangular, terminated at c. 63– Treptotreta jucunda Henderson & MacKinnon, 1981 81% anterior from the umbo, and with the highest point at Figures 10E–M, 11 mid-length. Median buttress narrow, poorly defined, widening posteriorly towards dorsal pseudointerarea. 1981 Treptotreta jucunda; Henderson & MacKinnon, p. 296, fig. 5A–I, text-figs 3–4. Remarks. Tapuritreta gribovensis differs from the type species 2014 Treptotreta jucunda Henderson & MacKinnon; Per- Tapuritreta angusta in having a distinctly procline deltoid pseu- cival & Kruse, p. 389, fig. 22 (full synonymy). dointerarea bisected by a vestigial intertrough and shallow dorsal 1985 Angulotreta postapicalis Palmer; Popov, p. 21; pl. 2, sulcus. figs 5–14; text-fig. 3. Tapuritreta gribovensis differs from Tapuritreta reclinata Streng et al., 2011 from the Yudachica Member (Cambrian, Furongian) of the Oaxaquia Terrane, Mexico, in having a less Holotype. JCF 10714 (housed in James Cook University, Towns- convex ventral valve with an almost straight (not convex) lateral ville), ventral valve from Mailchange Limestone, Drumian, Wes- profile of ventral pseudointerarea, a distinct dorsal median sul- tern Queensland. cus, less prominent median buttress and dorsal median septum, and smaller and weakly impressed cardinal muscle scars not Material. NMW 2018.4G.55–58, 63, 78 ventral valves; 59–62, exceeding one-third valve length. 65–66, dorsal valves plus NMW 2018.4G.67.1–3 ventral valves; NMW 2018.4G.77.4–10 dorsal valves.

Stilpnotreta Henderson & MacKinnon, 1981 Remarks. The specimens from the Guzhangian of Novaya Zem- lya are assigned to Treptotreta jucunda because they exhibit a Type species. By original designation, Stilpnotreta magna Hen- number of diagnostic characters of the taxon as revised by Perci- derson & MacKinnon, 1981. val & Kruse (2014); they are similar in having the following characters: a moderately high, conical ventral valve with a pro- cline deltoid pseudointerarea divided by an indistinct inter- trough defined by deflection of growth lines (Fig. 9J, K); small, Stilpnotreta sp. cf. Stilpnotreta minuta Holmer et al., 2001 circular pedicle foramen, completely enclosed into metamorphic – Figure 10A D shell (Fig. 9M); a strong ridge-like, apical process widening ante- riorly and perforated by the internal foramen at the anterior ter- mination (Fig. 9C, L); a narrow dorsal pseudointerarea Material. NMW 2018.4G.38 (Fig. 10C), dorsal valve; NMW occupying less than half maximum valve width; and a high, 2018.4G.39 (Lv = 0.51 mm, W = 0.51 mm; Fig. 10A), NMW blade-like, triangular dorsal median septum with a rod-like pro- 2018.4G.40 (Lv = 0.56 mm, W = 0.62 mm; Fig. 10B, D), ventral jection at the tip (Fig. 11D–F). valves; all from locality 302/3. Compared with the Baltoscandian material of the species, described by Topper et al. (2013a, b) from The Alum Shale For- Remarks. These shells are most similar to Stilpnotreta minuta mation (Furongian, Jiangshanian, Parabolina brevispina Zone) of Holmer et al., 2001 from the Sarykumy Formation (Miaolingian, V€astergotland,€ Sweden, the shells from Novaya Zemlya differ by HOLMER ET AL.: CAMBRIAN BRACHIOPODS FROM NOVAYA ZEMLYA 15

FIG. 10. Specimens from Snezhnyye Gory Formation (Miaolingian, Guzhangian), locality 302/3, Pan’kov Land peninsula, Novaya Zemlya, South Island. A–D, Stilpnotreta sp.: A, NMW 2018.4G.40, ventral valve exterior; B, D, NMW 2018.4G.39, ventral valve int- erior, oblique lateral view of exterior; C, NMW 2018.4G.38, dorsal valve interior. E–M, Treptotreta jucunda Henderson & MacKinnon, 1981: E–F, H–I, NMW 2018.4G.58, ventral valve exterior, ventral, oblique lateral and oblique posterior views, enlarged pitted micro- ornament of metamorphic shell; G, NMW 2018.4G.57, ventral valve interior; J, M, NMW 2018.4G.78, ventral valve posterior view, enlarge posterior view of the umbonal area showing pedicle foramen; K, NMW 2018.4G.73, ventral valve posterior view; L, NMW 2018.4G.55, oblique anterior view of ventral valve interior. Abbreviations: ap, apical process; cmf, cardinal muscle field; um, umbonal muscle scar. Scale bars represent: 200 lm(A–D, J–L); 500 lm(E–H); 5 lm (I); 20 lm (M). 16 PAPERS IN PALAEONTOLOGY HOLMER ET AL.: CAMBRIAN BRACHIOPODS FROM NOVAYA ZEMLYA 17 having a posteriorly inclined umbonal area of the ventral valve, Alvaro et al. (2013, fig. 19.5), shows that the Cambrian a straight to gently convex (not concave) ventral valve sagittal (Miaolingian) trilobite faunas of Novaya Zemlya appear profile, and a less prominent rod on the tip of the dorsal median within the distinct cluster, uniting the Miaolingian faunas septum. from Avalonia, Bohemia, Holy Cross Mountains, Mor- The shells from the Karpinsk Formation on the southern occo, Sweden/Norway, Turkey and the Armorican cluster coast of Gribovaya Bay, described and illustrated by Popov of terranes. (1985) as Angulotreta postapicalis Palmer, 1954, are morphologi- The published information on Cambrian (Wuliuan– cally very close to the specimens from the Pan’kov Land penin- sula and are here reassigned to Treptotreta jucunda. Jiangshanian) brachiopod faunas from Novaya Zemlya has mainly been published in Russian and is not accessi- ble (Walcott 1924; Popov 1984). The previously described SIGNIFICANCE OF THE FAUNA faunas are characterized by a low richness, which makes it difficult to use them for a detailed biogeographical Cambrian (Miaolingian–Furongian) deposits of the epei- assessment. Nevertheless, as discussed below, five succeed- ric Baltoscandian Basin preserve an extensive record of ing brachiopod assemblages can be recognized in the successive brachiopod faunas (Walcott 1912; Martinsson Cambrian sedimentary succession of Novaya Zemlya. 1968; Popov et al. 1989; Holmer & Popov 1990; Puura & The earliest brachiopod assemblage yet documented Holmer 1993; Topper et al. 2013a, b, etc.) However, the from Novaya Zemlya, is recovered from the lower part of existing information on the brachiopod faunas that Astafiyev Formation and is Wuliuan in age (Fig. 2). This inhabited the Cambrian margins of the Baltica palaeocon- is a low richness micromorphic linguliform brachiopod tinent is very poor, because the Palaeozoic continental association including unnamed species assigned to shelves of Baltica were mainly destroyed by the subse- Acrothyra and Hadrotreta, and early representatives of quent continental collisions later in the Palaeozoic (Cocks Acrothele (Popov 1985), which was a cosmopolitan genus. & Torsvik 2005; Torsvik & Cocks 2017). The only possi- Acrothyra is otherwise confined to Laurentia (Rowell ble preserved information from the marginal faunas 1980; Skovsted et al. 2017), while the poorly known type derives from the Holy Cross Mountains, Poland and the species Acrothyra proavia (Matthew, 1899), probably orig- Novaya Zemlya Archipelago. The published documenta- inated from the West Avalonia location at Cape Breton, tion on the Cambrian brachiopods of the Holy Cross suggesting that it was dispersed across the Iapetus Ocean Mountains is inadequate, while its position at the conti- during Miaolingian time. This is supported also by its nental margin of Baltica during the Cambrian times is Baltica (Novaya Zemlya) occurrence. The data presented somewhat controversial (Cocks & Torsvik 2005; Zyli_ nska by Skovsted et al. (2017) show a close association of 2013; Torsvik & Cocks 2017). The Novaya Zemlya Archi- Acrothyra and Hadrotreta with outer shelf settings, which pelago is widely considered as the outer part of the Tima- is consistent with the distribution of these taxa in the nid Orogen, which represents a terrane assemblage that Wuliuan deposits of Novaya Zemlya, where they occur in was accreted along the active Baltica continental margin limestone nodules within laminated, grey shale as well as sometime in the Infra-Cambrian (Torsvik & Cocks 2017). co-occurring with the trilobite (Solovyev During the Cambrian – Middle Ordovician the whole et al. 1986). Remarkably, there is no record of these Baltica continent underwent a large scale rotation of c. acrotretides from the Baltoscandian Basin. Hadrotreta is a 120°, which probably occurred mainly during the Furon- widespread taxon that achieved an almost cosmopolitan gian – Early Ordovician (Torsvik & Rehnstrom€ 2001; distribution at low latitudes during Miaolingian time Cocks & Torsvik 2005). Thus, during Guzhangian time, (Popov et al. 2015), but it is yet unknown from temper- the Novaya Zemlya sector was located at the easternmost ate to high-latitude Gondwana including North Africa, corner of Baltica, being separated from the North African Central Iran and Mediterranean peri-Gondwana, as well sector of Gondwana by a narrow oceanic space (Torsvik as from the Baltoscandian Basin. The geographically clos- & Cocks 2017; fig. 5.1). It is not surprising that the most est Hadrotreta occurrence to Novaya Zemlya was proba- recent analysis of Cambrian trilobite biogeography by bly documented from the Parahio Formation

FIG. 11. Treptotreta jucunda Henderson & MacKinnon, 1981; Snezhnyye Gory Formation (Miaolingian, Guzhangian), locality 302/3, Pan’kov Land peninsula, Novaya Zemlya, South Island. A–B, NMW 2018.4G.60, dorsal valve interior, oblique lateral view of interior. C–D, NMW 2018.4G.62, dorsal valve interior, oblique lateral view of interior showing two small septal rods. E, NMW 2018.4G.59, dorsal valve interior, oblique lateral view of dorsal valve interior showing prominent septal rod. F, NMW 2018.4G.61, dorsal valve inte- rior. G, J, K, NMW 2018.4G.65, dorsal valve exterior, enlarged view of metamorphic shell pitted micro-ornament, oblique lateral view of metamorphic shell. H, NMW 2018.4G.64, dorsal valve exterior. I, NMW 2018.4G.66, dorsal valve exterior. Scale bars represent: 500 lm(A–G, I); 200 lm (H); 10 lm (J); 50 lm (K). 18 PAPERS IN PALAEONTOLOGY

( salteri trilobite Zone) of the Indian biogeographical distribution suggests that the dispersal of Himalayas (Popov et al. 2015), but otherwise, the bra- Diraphora originated from Gondwana locations, where it chiopod faunas from these regions are very different, as probably first appeared. follows from the biogeographical analysis published in the The late Guzhangian linguliform microbrachiopod cited paper. fauna is derived from the lower part of the Karpinsk For- The next brachiopod association occurs in the upper mation (Koldiniella mittella – Modocia arctica local trilo- part of the Snezhnyye Gory Formation and is dated as bite Zone), which by occasional occurrences of the early Guzhangian (Baltoscandian Paradoxides forchammeri eponymous species can be correlated with the Agnostus trilobite Superzone). The assemblage includes only two pisiformis Zone of Baltoscandia (Solovyev 1986). The full taxa, the acrotretide Prototreta gribovensis and the orthide brachiopod assemblage (Fig. 2) includes species of Diraphora simplex, which form a characteristic bra- Acrothele, Anabolotreta?, Orbithele?, Stilpnotreta, Tapu- chiopod shell bed. Both taxa belong to the local endemic ritreta, Treptotreta and Wahwahlingula, described here, species, but to geographically widespread genera. No sim- plus Dictyonina, which was briefly discussed and illus- ilar taxa have yet been recorded from Baltoscandia. trated by Popov (1985). Among them Diraphora simplex requires special attention The acrotheloids Acrothele and Orbithele? and the because its occurrence at Novaya Zemlya probably repre- paterinide Dictyonina are relatively common and cos- sents one of the earliest signs of rhynchonelliform bra- mopolitan components in Drumian–Guzhangian linguli- chiopod proliferation on the shelves of Baltica together form brachiopod faunas, while the presence of with Oligomys exporrectus (Linnarsson, 1876), which Tapuritreta and Wahwahlingula is somewhat unusual, appears about that time in Scandinavia. The Gondwanan because these are their only documented Miaolingian occurrences of Diraphora in Central Iran (Wolfart 1974) occurrences to date. The Furongian occurrences of Tapu- and Australia (Kruse 1990, 1991) are thought to be of ritreta are confined to Alborz (Popov et al. 2009) and early Wuliuan age and therefore the earliest. In Laurentia, Oaxaquia (Streng et al. 2011) terranes, where both are Diraphora is confined to the late Wuliuan – Drumian associated with temperate latitude Peri-Gondwana. The (Bell 1941), while in the peri-Siberian Altai Region proliferation of Wahwahlingula occurred only near the (Aksarina & Pelman 1978) and Novaya Zemlya it is of Cambrian–Ordovician boundary, when species of the early Guzhangian age (Fig. 12). The observed pattern of genus dispersed across Baltica (Popov et al. 1989),

FIG. 12. Palaeogeographical reconstruction for Cambrian, Guzhangian age showing the position of Novaya Zemlya on the eastern margin of the Baltica continent, and the geographical distribution of selected brachiopod taxa documented from the Cambrian of Novaya Zemlya. Relative position of major early Palaeozoic continents (e.g. Laurentia, Baltica, Gondwana and Siberia) mainly after Torsvik & Cocks (2017) with significant emendations for tropical peri-Gondwana, related to the position of the North China Palaeo- continent and terranes associated with Kazakh Archipelago (for further discussion see Popov & Cocks 2017). Colour online. HOLMER ET AL.: CAMBRIAN BRACHIOPODS FROM NOVAYA ZEMLYA 19

Laurentia (Popov et al. 2002; Holmer et al. 2005) and requires further study; the recorded rhynchonelliform Alborz (Ghobadi Pour et al. 2011), and has little rele- brachiopod species are local endemics, and two of them, vance to the brachiopod biogeography in Guzhangian Eoorthis sabus and Huenellina triplicata, first appear in time. Anabolotreta? glabra is a distinct species, which Jiangshanian. The genus Eoorthis is a cosmopolitan taxon, occurs in Laurentia (Streng & Holmer 2006) and South which appears in Laurentia at about the same time as in China (Engelbretsen & Peng 2007). Novaya Zemlya, while Huenellina is unknown outside Treptotreta jucunda has a long stratigraphical range Novaya Zemlya. As for the family Huenellidae, it exhibits (from Drumian to Jiangshanian) and a wide geographical pantropical distribution throughout the Furongian distribution, including the Australian sector of Gondwana (Carlson 2002), being documented from Laurentia, (Percival & Kruse 2014), New Zealand (Henderson & Siberia and Kazakh terranes. MacKinnon 1981), South China (Engelbretsen & Peng In general, the Miaolingian–Furongian brachiopod fau- 2007), and probably the Atasu–Zhamshi Terrane of Kaza- nas of Novaya Zemlya are characterized by low richness khstan (Holmer et al. 2001), but it is remarkably absent and show little overlap with contemporaneous faunas of from Laurentia (Fig. 12). Its appearance in Baltoscandia the Baltoscandian Basin in their generic composition. The (Topper et al. 2013a, b) was delayed until the Paibian. Guzhangian brachiopod assemblage shows mixed biogeo- Stilpnotreta has a pantropical distribution (Fig. 12), but graphical signals suggesting faunal exchange with temper- its reported occurrence in the Furongian of Baltoscandia ate latitude (Tapuritreta, Diraphora) and tropical (Puura & Holmer 1993) is questionable. (Stilpnotreta, Treptotreta) peri-Gondwana. This was prob- The Furongian brachiopod faunas at Novaya Zemlya ably facilitated by the geographical position of the region are characterized by a proliferation of rhynchonelliform on the east margin of Baltica, as well as favourable ocea- brachiopods, which is not found in Baltoscandia, and nic currents (Fig. 12). The Furongian (Paibian–Jiangsha- they are represented by different taxa in these areas. The nian) is associated with a proliferation of Paibian assemblage includes Billingsella holtedahli and rhynchonelliform brachiopods, which is unknown from Ocnerorthis opius (see Walcott 1924). Both belong to gen- Baltoscandia, while the appearance of Billingsella and era that are known from Laurentia (Freeman & Stitt Ocnerorthis at Novaya Zemlya pre-dates their appearance 1996), where they make a later appearance in the Jiang- in Laurentia. shanian (Elvinia Zone). The only other Billingsella species reported from Baltica is Billingsella? lindstromi (Lin- Acknowledgements. LEH is supported by the Swedish Research narsson, 1876) from the Guzhangian (Paradoxides for- Council (VR Project no. 2018-03390) and by a Zhongjian Yang chammeri Zone) of Sweden. This species remains very Scholarship from the Department of Geology, Northwest poorly known. It was originally considered as an orthide University, Xi’an. LEP is supported by the National Museum of (Linnarsson 1876), and subsequently re-assigned to Bill- Wales. MGP acknowledges support from the Golestan Univer- sity, and logistical support from the National Museum of Wales, ingsella by Walcott (1912), but this record is still uncer- Cardiff and the Uppsala University. IAK acknowledges support tain. Rhynchonelliform brachiopods are common in the from the St Petersburg State University. ZFZ sincerely acknowl- peri-Gondwanan terranes of Iran, and, in particular, from edges the National Natural Science Foundation of China Alborz (Popov et al. 2013), where billingsellids and syn- (41425008, 41621003, 41720104002 and 41890844) and the 111 trophioids form extensive shell beds in the Mila Forma- Centre (D17013) for the continuous financial support for the tion Member 3 (Paibian–Jiangshanian). The record of palaeobiology group in Xi’an. The Changjiang Scholars rhynchonelliform brachiopods from North Africa and (T2016155 to ZFZ) and Wanrenjihua programs (W03020685 to Mediterranean Gondwana is relatively poor. It includes ZFZ) are sincerely acknowledged. Michal Mergle and Uwe only reports on the occasional occurrence of Billingsella Balthasar commented on an earlier draft of this manuscript. and Saccogonum species in Morocco and Spain (Havlıcek & Josopait 1972; Mergl et al. 1990; Alvaro et al. 2007). Author contributions. LEH, LEP, MGP, YL and ZFZ are respon- Mergl (1983) described Billingsella destombesi from the sible for newly designated brachiopod taxa, while IAK and LEP are responsible for the geological information presented in the ‘middle’ Cambrian of Anti-Atlas, Morocco and if the paper. Guzhangian age of this taxon is confirmed, it could be the earliest species of the genus yet recorded. The youngest Cambrian (Furongian) brachiopod asso- DATA ARCHIVING STATEMENT ciation from the uppermost Karpinsk Formation has a Jiangshanian age. It includes ‘Lingulella’ cf. desiderata, This published work and the nomenclatural acts it contains have Billingsella holtedahli, Eoorthis sabus, Huenellina triplicata been registered in ZooBank: http://zoobank.org/references/ and Ocnerorthis sp. cf. O. opius documented by Walcott E2FC1C9C-7113-4779-A4A3-ACB485CF1C90 (1924) from Holtedahl’s locality 68b. This fauna has unfortunately never been re-studied and the linguliform Editor. Javier Alvaro 20 PAPERS IN PALAEONTOLOGY

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