First skeletal microfauna from the Cambrian Series 3 of the Jordan Rift Valley (Middle East)

OLAF ELICKI

ELICKI, O., 2011:12:23. First skeletal microfauna from the Cambrian Series 3 of the Jordan Rift Valley (Middle East). Memoirs of the Association of Australasian Palaeontologists 42, 153-173. ISSN 0810-8889.

For the first time, a Cambrian microfauna is reported from the Jordan Rift Valley. The fauna comes from low-latitude carbonates of the Numayri Member (Burj Formation, Jordan) and to a lesser degree the equivalent Nimra Member (Timna Formation, Israel). Co-occuring with trilobite, brachiopod and hyolith macrofossils, the microfauna is represented mostly by disarticulated poriferid (mostly hexactinellids) and echinoderm remains (eocrinoids and edrioasteroids). Among the hexactinellids, Rigbyella sp., many isolated triactins and tetractins, as well as a few pentactins and rare hexactins occur. Additional poriferid spicules come from heteractinids (Eiffelia araniformis [Missarzhevsky, 1981]) and polyactinellids (?Praephobetractinia). Chancelloriids (Archiasterella cf. hirundo Bengtson, 1990, Allonnia sp., Chancelloria sp., ?Ginospina sp.) are a rather rare faunal element. Micromolluscs are represented mainly by an indeterminable helcionellid. The probable octocoral spicule Microcoryne cephalata (Bengtson, 1990), torellellid and hyolithellid hyolithelminths, and a bradoriid arthropod occur as very few or single specimens. The same is the case with a probable siphogonuchitid. The occurrence of a cornulitid related microfossil may extend the stratigraphic range of this fossil group significantly. The rather low-diversity microfauna is overwhelmingly dominated by sessile epibenthic biota. The preferred feeding habit seems to have been suspension feeding and minor deposit feeding. The microfauna from the Jordan Rift Valley is typical for low-latitude carbonate environments of Cambrian Series 3 age that corresponds to the traditional late early to middle Cambrian. Some taxa indicate a closer relation to the equatorial Gondwanan Iran and Australia. Some connection to the European shelf of Perigondwana may also have existed.

O. Elicki ([email protected]), Freiberg University, Geological Institute, Bernhard-von- Cotta Street 2, 09599 Freiberg, Germany. Received 9 August 2011.

Keywords: Small shelly fossils, poriferids, echinoderms, Cambrian, Jordan, Israel, Dead Sea.

CAMBRIAN successions are spectacularly Much of the previous palaeontological work exposed in the Jordan Rift Valley. Outcrops along concentrated on the stratigraphic position and the eastern (Jordanian) shoreline of the Dead Sea correlation of the Dead Sea Cambrian succession. and a little south in the northern Wadi Araba yield As such, there was often focus on macrofossils. fossiliferous deposits of shallow and marginal Micropalaeontological sampling has been lagging marine origin (Burj Formation). In the south- behind in this region. This paper documents the western Wadi Araba (Israel) equivalent Cambrian first skeletal microfauna of Cambrian age from sediments also occur locally (Timna Formation). this region. Cambrian trilobites have been known from the In the course of several joint research projects region for about a century (Dienemann 1915; between the Freiberg working group and King 1923; Richter & Richter 1941). More recent colleagues from Yarmouk University (Irbid) work on trilobites and substantial investigation and the Natural Resources Authority (Amman) on other fossil groups including brachiopods, has resulted in bulk collections of carbonate, hyoliths and trace fossils have been completed siliciclastic and mixed lithological successions (Parnes 1971; Cooper 1976; Bandel 1986; along measured stratigraphic sections from Seilacher 1990a, b; Rushton & Powell 1998). Jordanian Cambrian surface outcrops. These have A summary of the history of palaeontological, undergone palaeontological and sedimentological petrographical, palaeogeographical and facies analyses (Elicki & Shinaq 2000; Elicki et al. investigations in this region is provided by Elicki 2002; Elicki 2007a; Shinaq & Elicki 2007; Geyer (2007a) and Shinaq & Elicki (2007). & Elicki submitted; Hofmann et al. submitted). 154 AAP Memoir 42 (2011)

Powell 1989; Teimeh et al. 1990; Bandel & Shinaq 2003). The oldest Cambrian sedimentary rocks (part of Umm Gaddah Formation and Salib Formation, about 200 m thick; Fig. 2) are continental, fluvial to alluvial siliciclastics (Powell 1989; Amireh et al. 1994; Makhlouf 2003; Amireh et al. 2008). The overlying Burj Formation represents a marine maximum stage transgression (Amireh et al. 1994; Rushton & Powell 1998; Schneider et al. 2007). The start of transgression is indicated by some trace-fossil content in the lower Burj Formation (Tayan Member: Elicki 2007a), pointing to an early Cambrian age of these siliciclastics. The conformably succeeding Numayri Member (limestones, dolostones, minor siliciclastics) includes a variety of marginal- marine environments including open and restricted marine areas, shoals, lagoons, sabkhas (Shinaq & Bandel 1992; Rushton & Powell 1998; Elicki et al. 2002; Shinaq & Elicki 2007). The Numayri Member yields various macrofossils including trilobites, brachiopods and hyolithids, but also trace fossils (Richter & Richter 1941; Parnes 1971; Cooper 1976; Seilacher 1990a; Rushton & Powell 1998). The upper portion of the Burj Formation is represented by the siliciclastic Figure 1. Geographical map of the Jordan Rift Valley. Hanneh Member containing a rich marine Working areas are indicated (for detailed location of ichnofauna (Seilacher 1990a, b; Mángano et al. the sampled sections see Elicki 2007a). 2007; Hofmann et al. 2008; Elicki et al. 2010, Hofmann et al. submitted) and some trilobites The systematic and detailed field work led to the (Elicki & Geyer, submitted). The package is discovery of sections largely unaffected by the conformably overlain by the siliciclastic Umm heavy diagenetic overprint typical of the region, Ishrin Formation which contains trilobites and facilitated detailed micropalaeontological and trace fossils only at its base, but generally investigation of skeletonised body-microfossils. indicates regression and return of continental For the first time, this paper focuses on description conditions (Makhlouf & Abed 1991; Hofmann and documentation of Cambrian microfossils et al. submitted). from this palaeogeographic region. The thickness of the Burj Formation in the The palaeontological specimens figured in this Dead Sea region varies from about 190 m in the paper, are housed at the Geological Institute of north (Wadi Zerqa Ma’in) to approximately 120 Freiberg University, Germany (archive numbers m in the (southern) Safi region (Powell 1989), FG 595 and FG 619). thinning to zero in the basement area of southern Wadi Araba. GEOLOGICAL AND STRATIGRAPHIC Evidence for the stratigraphic position of BACKGROUND the Cambrian succession comes from trilobites. The field area is situated along the eastern These have been found in the marine carbonates shoulder of the Jordan Rift Valley in the Dead Sea (Numayri Member) over the last hundred area, and the northeastern Wadi Araba south of years (Dienemann 1915; King 1923; Richter & the Dead Sea (Jordan). The Timna region (Israel), Richter 1941) and were recently summarised from where some microfauna was kindly provided and reviewed by Rushton & Powell (1998). for investigation by Professor Michael Bassett New discoveries of abundant, well preserved (Cardiff), represents the only locality investigated specimens by the Freiberg working group from from southwestern Wadi Araba (Fig. 1). the carbonate Numayri Member and the Hanneh The Cambrian sediments nonconformably Member siliciciclastics, enabled a critical review overlie Precambrian basement rocks of the of the Cambrian trilobite fauna and confirmed a Arabo-Nubian Shield (Aqaba Complex) or, stratigraphic position within the traditional Early– locally, Neoproterozoic volcanoclastics and Middle Cambrian boundary interval, probably conglomerates (Araba Complex; Bender 1974; equivalent to parts of Series 3, stage 5 (Rushton AAP Memoir 42 (2011) 155

Figure 2. Generalised composite stratigraphic column of the Cambrian in the Dead Sea area (left; Salib Formation and Umm Ishrin Formation are only partially illustrated). Lithostratigraphic range of the Numayri Member in the investigated sections at the right (Numayri Member without fill pattern; Tayan Member and Hanneh Member with cross fill pattern and only partially illustrated). Lithostratigraphic position ofthe described microfaunas are indicated by bars. & Powell 1998; Geyer & Landing 2004; Geyer ml acid) has been used, a method which utilises & Elicki submitted). the absorption of water by crystals of calcium acetate (Nötzold 1965). Sieving procedure was MATERIALS AND METHODS executed using sieves of mesh size 0.3 mm and 0.1 The fossiliferous carbonates were investigated mm. Microscopic work has been achieved using by thin-section analysis and also preparated by a ZEISS SV-11 polarisation light-microscope. laboratory chemical work. Best results have SEM investigation was undertaken at the been attained by careful chemical solution of Geological Institute of Freiberg University with the carbonate rocks using formic acid (about the field emission scanning electron microscope 15%) and in some cases acetic acid (about 15%). JEOL JSM-7001 F. For related compositional In some cases, concentrated acetic acid with identification of the microbiota, an energy added anhydrous copper sulphate (5 g per 100 dispersive X-ray spectroscopy (edx) BRUKER 156 AAP Memoir 42 (2011)

Figure 3. Hexactinellid spicules of Rigbyella sp. from the lower part of the Numayri Member at the unnamed valley immediately south of Wadi At Tayan. All scale bars 100 µm. A, FG 619/13-133; B, FG 619/14-133; B2, detail of basal part of same specimen FG 619/14-133; C, FG 619/15-133; D, FG 619/16-133; E, FG 619/17- 133; F, FG 619/21-133; G, FG 619/33-133; H, FG 619/32-133; I, FG 619/34-133; J, FG 619/22-133; K, FG 619/12-133; L, FG 619/31-133; M, FG 619/29-133; N, FG 619/24-133; O, FG 619/25-133; P, FG 619/27-133; Q, FG 619/23-133; R, FG 619/28-133; S, FG 619/26-133. device with software ESPRIT 1.9 was utilised. Phylum PORIFERA Grant, 1836 Class HEXACTINELLIDA, Schmidt, 1870 SYSTEMATIC PALAEONTOLOGY Order AMPHIDISCOSA Schrammen, 1924 Poriferan spicules Family STIODERMATIDAE Finks, 1960 Sponge spicules were recovered in only a few samples, but normally in very large numbers. Rigbyella Mostler & Mosleh-Yazdi, 1976 Lithotypes from the southern Dead Sea region (Wadi Umm Jafna, Wadi Uhaymir, unnamed Type species. Rigbyella ruttneri Mostler & valley south of Wadi At Tayan) contained the Mosleh-Yazdi, 1976. majority of these microfossils. Dominating types include triactine, tetractine and pentactine Rigbyella sp. (Fig. 3A-S) hexactinellid remains (c. thousands of spicules per kilogram of rock). Heteractinid spicules, in Material. About 450 isolated siliceous spicules. contrast, are of limited quantity. Some spicules could not be assigned systematically (see below). Description. Stiodermatiid (sensu Hooper & Van Nearly all remains are siliceous and come mostly Soest 2002, p. 1214) pinnules with dramatically from the lower portion of the Numayri Member swollen vertical ray perpendicular to a generally (Burj Formation). four-rayed stauractine-like cross built by ‘basal’ spicules lying in same plane (Fig. 3B2). Swelling AAP Memoir 42 (2011) 157 of the vertical ray begins some distance above the spicules from Jordan. Whereas hexactins are the ‘basal’ rays so that there is a bottle-neck- exclusively mentioned in the original diagnosis like transition. In a few cases, the ‘basal’ cross of the genus Rigbyella, the current Treatise (Finks is tilted up to about 45° relative to the vertical et al. 2004) also reports pentactins and includes ray (e.g., Fig. 3C, F). The swollen vertical ray is the spicules described by Rigby (1975) from the always many times the length of the other rays; Wilberns Formation (see above) in this genus. its diameter is also generally larger than the length Siliceous pentactins with swollen, spindle- of the other rays (up to five times). There are shaped vertical ray and tetraradially symmetrical two morphotypes occurring in the sample: the morphology are also described by Bengtson first and most common type is rather elongate (1986) from Furongian carbonates from western and rice-grain shaped (Fig. 3A-M). The second Queensland (Australia). The author mentioned morph is not as tall, commonly about two-thirds also records similar and very abundant spicules the length of the first morphotype, but with with smooth and acanthose surface ornament. distinctly larger diameter and teardrop-shaped He further noted that specimens with excessively (Fig. 3N-S). The distal portion of the spicules is swollen rays similar to those as published by cuspid, and especially in the second morphotype, Rigby (1975, compare explanations above) well developed, (e.g., Fig. 3P, S). are not present in the spicule microfauna from Queensland. Indeed, the spicules from Jordan Discussion. All specimens come from one thin described here, commonly have a distinctly more bioclastic limestone horizon near the base of swollen vertical ray, but, their overall construction the Numayri Member (unnamed adjacent valley is otherwise very close to the spicule morphs south of Wadi At Tayan) immediately on top of figured by Bengtson (1986: Fig. 9C). basal, laminated and stromatolitic dolomites of Bengtson also discussed spicules from the the lowermost Numayri Member. Accompanying early Cambrian of Kazakhstan (referred to fauna consists of small trilobite hash, fragmented Azyrtalia), published by Nazarov (1973, cited brachiopod shells, and isolated eiffeliid and rare by Bengtson 1986) and suggests these may be ‘normal’ hexactinellid spicules. synonymous with Rigbyella. If this assumption is Spicules of similar form are only rarely correct, then Azyrtalia would represent an older reported from Cambrian rocks. Rigby (1975) synomym and would have taxonomic priority. described sponge spicules from Late Cambrian- However, reinvestigation of the cited material is Early Ordovician limestones of Texas with needed to clarify this aspect. pentactins that have a distinctly swollen vertical From the middle Cambrian Georgina Basin in ray. Unfortunately, no photographs are included central northern Australia (Northern Territory), by Rigby, but, simplified drawings indicate Mehl (1995, 1998) described Thoracospongia these specimens are of very similar morphology follispiculata with typical hexactins and pentactins (Rigby, 1975: text-fig. 1, H-N). Rigby (1975) also characterised by having a distinctly swollen reported an association of ‘normal’ and acanthose vertical ray. These forms are quite similar to hexactines and pentactines, and presented the specimens from Jordan. Differences consist a reconstruction of the spicule scleritome, in the occurrence of distinct longitudinally interpreting the swollen spicules as specialised ribs along the swollen ray and in the lack of a morphs that form a dermal pavement surrounding bottle-neck-like transition to the “basal” rays in the interior of the sponge. the material from Australia. Unfortunately, the Mostler & Mosleh-Yazdi (1976) described preservation of the spicules from Jordan is always spicules from the Furongian of Iran (Mila very coarse-grained (diagenetic replacement by Formation), including hexactine spicules with coarse quartz crystals), so that the original surface swollen vertical ray as well as swollen lateral structure can only be suspected at best. In some rays. Sometimes the rays are drop-shaped or cases it cannot be excluded that (faint) primarily cap shaped and pointed. The authors erected longitudinally ribs may have been present in the the taxon Rigbyella ruttneri which includes (1) Jordanian spicules. The Australian spicule fauna swollen hexactins, (2) hexactins with rays of also consists of hexactinellid, demosponge and unequal length, and (3) hexactins with rays with calcarea spicules (e.g. Eiffelia), similar to the split ends. Some of the figured specimens (e.g., association recovered from Jordan. p. 34: plate III figure 6) are very similar to the Considering the morphological variability material from Jordan. The specimens from Jordan of the sclerites from all these regions, the close generally do not have swollen lateral rays (which affinity of the described spicules to Rigbyella are quite common in the Iranian fauna). However, is most reasonable. Minor morphological in cases where only the vertical ray is swollen differences from the only species of this genus in the Iranian material, these are very similar to (R. ruttneri) and the strong diagenetic overprint 158 AAP Memoir 42 (2011)

Figure 4. Unassigned hexactinellid spicules: triactins (A-G), tetractins (H-J’), pentactins (K’-R’), hexactins (S’). Specimens come from the lower portion of the Numayri Member at Wadi Uhaymir (A-D, I-W, Y-Z, A’-B’, E’-G’, K’-O’, Q’-R’) and Wadi Umm Jafna (E-F, X, C’-D’, H’-J’, P’, S’-S’1), as well as from upper portion of the upper Numayri Member at Wadi Zerqa Ma’in (G-H). All scale bars 100 µm. A, FG 619/3-398 B, FG 619/13-398; C, FG 619/18-398; D, FG 619/33-398; E, FG 619/53-398; F, FG 619/65-398; G, FG 619/23-451; H, FG 619/29-451; I, FG 619/2-398; J, FG 619/4-398; K, FG 619/7-398; L, FG 619/8-398; M, FG 619/9- 398; N, FG 619/14-398; O, FG 619/17-398; P, FG 619/15-398; Q, FG 619/16-398; R, FG 619/19-398; S, FG 619/20-398, T, FG 619/23a-398; U, FG 619/25-398; V, FG 619/30-398; W, FG 619/29-398; X, FG 619/63- 398; Y, FG 619/28-398; Z, FG 619/31-398; A’, FG 619/37-398; B’, FG 619/44-398; C’, FG 619/50-398; D’, FG 619/51-398; E’, FG 619/40-398; F’, FG 619/42-398; G’, FG 619/32-398; H’, FG 619/54-398; I’, FG 619/59-398; J’, FG 619/61-398; K’, FG 619/24-398; L’, FG 619/38-398; M’, FG 619/45-398; N’, FG 619/47- 398; O’, FG 619/48-398; P’, FG 619/62-398; Q’, FG 619/5-137; R’, FG 619/28-398; S’1, FG 619/57-398; S’2, detail of same specimen FG 619/57-398. AAP Memoir 42 (2011) 159 in the spicules from Jordan may lead to some 1984 Niphadus xihaopingensis; Duan, p. 168, uncertainty. pl. 4, fig. 17. 1984 Niphadus complanatus; Duan, p. 168, pl. Unassigned hexactinellid spicules. Unassigned 4, fig. 8. isolated triactins, tetractins and pentactins 1986 Lenastella sp.; Laurie, p. 447, fig. 10C. represent the vast majority of the hexactinellid 1990 Eiffelia araniformis (Missarzhevsky); spicule microfauna. Tetractins are often Bengtson et al., p. 27, figs 12, 13. represented by calthrops and triaenes, and less 1994 Eiffelia araniformis (Missarzhevsky); common by stauractins. Hexactins are rare. Elicki, p. 73, figs 14-15. Almost all spicules have a smooth surface 1996 Eiffelia araniformis (Missarzhevsky); (Fig. 4A-Z, A’-R’). Acanthose ornamentation is Culver et al., p. 4, fig. 5.13. extremely rare (Fig. 4S’1, S’2). 2001 Eiffelia cf. araniformis (Missarzhevsky); The majority of spicules occur in huge Sarmiento et al., p. 120, pl. 2, figs 10-11. numbers (mass occurrences within distinct layers) 2004 Eiffelia araniformis (Missarzhevsky); which yield only few other fossils: trilobite Wrona, p. 18, fig. 5H. and hyolith remains. Taxonomic assignment is 2005 Eiffelia araniformis (Missarzhevsky); not useful for these specimens because of the Elicki, p. 163. huge morphological variability of the group. 2006 Eiffelia araniformis (Missarzhevsky); Nevertheless, it can be argued that (1) the huge Elicki, p. 9. number, (2) the nearly exclusive occurrence of the 2006 Eiffelia araniformis (Missarzhevsky); spicules in distinct limestone layers, and (3) the Skovsted et al., p. 1098, figs 7.23, 7.24. content within the taphocoenosis from different 2007b Eiffelia araniformis (Missarzhevsky); localities, suggests the spicules derived from the Elicki, p. 146. same scleritome. Most of the spicules come from carbonate Material. About 10 isolated secondary silicified horizons in the middle to upper portion of the spicules from 2.5 m above the base of the Numayri Numayri Member of Wadi Umm Jafna and Wadi Member at the valley immediately south of Wadi Uhaymir, as well as from a loose boulder at Wadi At Tayan. Uhaymir and from the upper Numayri Member of Wadi Zerqa Ma’in. The variability of spicules Description. The specimens are regularly six- from Jordan is illustrated in Figure 4. rayed. Angle between rays is about 60°. No vertical ray (as sometimes reported for this taxon) Class CALCAREA Bowerbank, 1864 is observed. All rays are arranged in the same Order HETERACTINIDAE Hinde, 1888 plane. The length of the rays varies: in some Family EIFFELIIDAE Rigby, 1986 cases all six rays seem to have the same length (often they are broken), in other cases the length Eiffelia Walcott, 1920 differs. This is also the case regarding the width of the rays. Most specimens have one ray distinctly Type species. Eiffelia globosa Walcott, 1920. elongate (double to triple the length of the others).

Eiffelia araniformis (Missarzhevsky, 1981) Discussion. A full discussion of this species was (Fig. 5A-D) provided by Bengtson et al. (1990). Here, the opinion of Bengtson et al. (1990) is followed 1981 Lenastella araniformis; Missarzhevsky in concerning the assignment of the genus Lenastella Missarzhevsky & Mambetov, p. 76, pl. 12, Missarzhevsky, 1981, to the synonymy of Eiffelia figs 1, 10. Walcott, 1920, and in interpreting the various 1981 Lenastella aculeata; Missarzhevsky in Lenastella species erected by Missarzhevsky Missarzhevsky & Mambetov, p. 76, pl. 12, (1981) as belonging to the morphological range figs 2-4. of Eiffelia araniformis. 1981 Lenastella mucronata; Missarzhevsky in The specimens from Jordan come from Missarzhevsky & Mambetov, p. 77, pl. 12, bioclastic, oncolitic limestone with accompanying figs 8, 9, 11. small trilobite and brachiopod hash, Rigbyella 1981 Lenastella umbonata; Missarzhevsky in (see above), and rare hyolithid, chancelloriid and Missarzhevsky & Mambetov, p. 77, pl. 12, echinoderm remains. In most cases, the bioclasts fig. 12. are oncolitic. 1984 Actinoites universalis; Duan, p. 167, pl. 4, Eiffelia araniformis is known from the early figs 11, 13-14. and middle Cambrian interval almost worldwide. 1984 Actinoites simplex; Duan, p. 167, pl. 4, fig. 6. 160 AAP Memoir 42 (2011)

Figure 5. Heteractinids (A-D), ?polyactinellids (E-O), chancelloriids (P-Z), and other spicules (A’) from the Numayri Member (A-Z, A’) and the equivalent Nimra Member (Israel, W-Z). Scale bars D-V = 100 µm, A-C, W-X = 1 mm. A-D, Eiffelia araniformis (Missarzhevsky, 1981); A, FG 619/1-133; B, FG 619/2-133; C, FG 619/3-133; D, FG 619/4-133; E, polyactinellid praephobetractinid spicule, FG 619/11-133; F-O, ?polyactinellid spicules; F, FG 619/6-398; G, FG 619/12-451; H, FG 619/21-398; I, FG 619/34-398; J, FG 619/35-398; K, FG 619/39-398; L, FG 619/43-398; M, FG 619/46-398; N, FG 619/56-398; O, FG 619/60-398; P-Q, Archiasterella cf. hirundo Bengtson, 1990; P, FG 619/10-133; Q, FG 619/6-133; R-T, Allonnia sp.; R, FG 619/5-133; S, FG 619/9-133; T, FG 619/52-133; U, ?Ginospina sp., FG 619/8-133; U1, lateral view; U2, plan view; V-Z, Chancelloria sp.; V, FG 619/7-133; W, FG 619/19-200; X, FG 619/20-200; Y, FG 619/21-200; Z, FG 619/23- 200; A’, Microcoryne cephalata (Bengtson, 1990), FG 619/40-133. Unassigned sponge spicules. One specimen of the early Cambrian to early Ordovician (Mostler a polyactinellid, triradiate spicule (Fig. 5E) was 1985, 1996; Kozur 1991). Some similarity exists observed along with Rigbyella near the base of with Dodecaactinella Reif, 1968. However, a the Numayri Member at the valley immediately distinct difference to Dodecaactinella is in the south of Wadi At Tayan (see above). The spicule presence of the unbranched and clearly elongate has Mg-calcareous mineralogy. Two of the rays third main ray, and in the angles between rays (lower rays according to Finks et al. 2004) are in the specimen from Jordan. These characters dichotomously branched into two secondary rays; can be observed despite the broken preservation branching begins close to the point of divergence and are consistent with the diagnostic features of the main rays. The third main ray (upper of Praephobetractinia. These characters are also first-order ray) is distinctly longer, straight, and present in specimens from Greenland (Skovsted not branched, but clearly more tilted. Between 2006) and Pennsylvania (Skovsted & Peel 2010), the lower rays (originated by branching), the erroneously reported as Dodecaactinella. A original presence of a now broken additional ray specimen from Sardinia figured by Mostler (1985, is indicated by a round breaking suture. This ray pl. 3, fig. 3 = “phobetractine spicule”) and Finks arrangement and branching clearly point to the et al. (2004, p. 760. fig. 7a) is most similar to the systematic affiliation close to the calcisponge specimen described here. Praephobetractinia Kozur, 1991, known from A few spicules were recovered from limestone AAP Memoir 42 (2011) 161 samples of the lower portion in the Numayri of rays (4+0), ray configuration and proportions Member at Wadi Umm Jafna and Wadi Uhaymir. suggest the affiliation with A. hirundo from South Most specimens come from a loose boulder Australia (Bengtson in Bengtson et al. 1990). The sampled in the latter region (Fig. 5F-O). original spicule surface pattern is not visible due The spicules illustrated here are characterised to coarse recrystallisation. Because of this poor by conspicuous irregularity in the pattern of preservation of the material, the spicules from branching, ray arrangement and ray symmetry. Jordan are referred to Archiasterella cf. hirundo. The composition of the spicules appears to Archiasterella hirundo is known from the late be siliceous, but it is difficult to be sure this early Cambrian of South Australia, Germany, reflects the original composition because of Spain and probably from southern China the strong diagenetic overprint of silification (Bengtson et al. 1990; Elicki 1994; Fernández- in these sections. An axial canal is not visible Remolar 2001; Sarmiento et al. 2001; Moore et and this may be the result of diagensis as well. al. 2010). The polyactinellid or dodecaactinellid nature (Calcarea, Heteractinida) of some spicules Allonnia Doré & Reid, 1965 is possible (Reitner personal communication 2011). One of the spicules (Fig. 5M) has a ray Type species. Allonnia tripodophora Doré & configuration similar to phobetractins, but with Reid, 1965. an additional ray in a third plane. Mehl (1998 p. 1167, figs 7, 8, 10, and personal communication Allonnia sp. (Fig. 5R-T) 2011) figured demospongid specimens from the middle Cambrian of the Georgina Basin Material. Only few spicules, probably large (Australia) which resembles some of the material number of related single rays; from lower from Jordan (e.g. Fig. 5H). Numayri Member of the valley south of Wadi At The coeval fauna consists of abundant Tayan and from the higher Numayri Member of hexactinellid spicules and a few trilobite and Wadi Umm Jafna. hyolith remains. Discussion. The arrangement and number Phylum unassigned of rays (4+0) in these specimens are in the Class COELOSCLERITOPHORA Bengtson & range of the genus Allonnia. Jiang (in Luo et Missarzhevsky, 1981 al. 1982) erected Onychia tetrathallis from Order CHANCELLORIIDA Missarzhevsky, 1989 southern China for four-rayed spicules, which Family CHANCELLORIIDAE Walcott, 1920 were assigned later to the genus Allonnia by Qian & Bengtson (1989). The specimens from Archiasterella Sdzuy, 1969 Jordan are poorly preserved, so that species determination is difficult. Nevertheless, the Type species. Archiasterella pentactina Szduy, number, arrangement and orientation of the four 1969. rays point to a systematic affiliation with the species A. tetrathallis. Archiasterella cf. hirundo Bengtson, 1990 (Fig. Allonnia is known from Cambrian strata 5P-Q) worldwide and A. tetrathallis also has a worldwide distribution in the early and middle Cambrian. 2001 Archiasterella cf. hirundo Bengtson; Fernández-Remolar, p. 59, figs 3e-f, 8c-d, f. Chancelloria Walcott, 1920 2001 Archiasterella cf. hirundo Bengtson; Sarmiento et al., p. 120, pl. 2, fig. 5. Type species. Chancelloria eros Walcott, 1920. 2010 ?Archiasterella hirundo Bengtson; Moore et al., p. 1053, figs 11.1-11.3, 11.6, 11.7. Chancelloria sp. (Fig. 5V-Z)

Material. 3 spicules, numerous probably related Material. About 20 spicules, but many probably single rays; from lower Numayri Member of the related single rays; from lower Numayri Member valley south of Wadi At Tayan. of the valley south of Wadi At Tayan and from Timna area (upper Nimra Member of Timna Discussion. The typical rays, the characteristic Formation, which is more or less equivalent to arrangement and curvature, and canal openings the Numayri Member in Jordan). (foramina) on the lower side, identifies these spicules as chancelloriid and archiasterellid Description. Rosette-like sclerites with 5-6 rays (Szduy 1969; Bengtson et al. 1990). The number of very similar length and shape. Openings on the 162 AAP Memoir 42 (2011) lower side are round to slightly oval. Rays only Geyer, p. 107, fig. 2.13. slightly bent upwards; no vertical ray present. Material. One spicule from a loose boulder of the Discussion. The preservation of the sclerites is middle Numayri Member sampled at the mouth poor, so that detailed systematic affiliation cannot of the Wadi Qunai. be determined. Many accompanied unassigned specimens with chancelloriid characteristics Discussion. Sclerites with this characteristic possibly belong this genus. For some specimens, mace-shaped morphology were described by the foramen on the lower side of the sclerite Bengtson (in Bengtson et al. 1990) who grouped can faintly be seen. The shape and organisation them into the new genus Microcoryne (type of the spicule suggests affinity with the genus species: M. cephalata Bengtson, 1990). The Chancelloria, a genus which is extremely morphological variability of the spicules from common in Cambrian microfaunas worldwide. Australia is quite large. The phosphate mineralogy is assumed to be a secondary replacement. The Family SISSOSPINIDAE Missarzhevsky, 1989 Jordanian material coincides fully with spicules of M. cephalata described from the late early Ginospina Missarzhevsky, 1989 Cambrian of Germany (Elicki 1994; Elicki & Geyer 2010). The only difference is in the type Type species. Ginospina araniformis Missar- of preservation which is distinctly coarser in the zhevsky, 1989. Jordanian material due to different diagenetic overprint. ?Ginospina sp. (Fig. 5U1-U2) As discussed by Bengtson (1990), Microcoryne seems to be closest to the sclerites of octocorals. Material. One specimen from the lower Numayri In Jordan, Microcoryne is a very rare constituent Member of the valley south of Wadi At Tayan. in bioclastic carbonates which also yield trilobite hash. Besides the specimen from Jordan described Discussion. This secondarily silicified, poorly here, M. cephalata is hitherto reported from the preserved chancelloriid sclerite has 7 upright higher early Cambrian of South Australia and rays of variable length. Rudimentary foramina Germany. are visible on the lower side. A conspicuous character is the remarkable, somewhat elliptical Echinoderm remains arrangement of the rays. This kind of ray- Echinoderm remains are represented by isolated arrangement in chancelloriids is similar to the plates and ossicles of diagenetically altered, genus Ginospina from the early Cambrian secondarily phosphatised or silified composition. (Tommotian) of Siberia (Missarzhevsky 1989). They occur in bioclastic limestones in various levels within the whole Numayri Member. The Phylum, class, order & family unassigned majority of the ossicles described here come from the Wadi Zerqa Ma’in section which represents Microcoryne Bengtson, 1990 the uppermost part of the Numayri Member (Shinaq & Elicki 2007). The limestones of Type species. Microcoryne cephalata Bengtson, this section are interpreted as deposited within 1990. a shallow, open-marine, occasionally storm- influenced environment (Shinaq & Bandel 1992; Microcoryne cephalata (Bengtson, 1990) (Fig. Shinaq & Elicki 2007). 5A’) Phylum ECHINODERMATA Klein, 1734 1990 Microcoryne cephalata; Bengtson in Class, order & family unassigned Bengtson et al., p. 35, fig. 19. 1994 Microcoryne cephalata Bengtson; Elicki, p. gen. et sp. indet. (Fig. 6A-Z, A’-E’) 74, pl. 2, figs 6.10-6.16. 2010 Microcoryne cephalata Bengtson; Elicki & Material. Hundreds of ossicles from Wadi Zerqa

Figure 6. Echinoderm ossicles. All scale bars 100 µm. A, FG 619/7-115; B, FG 619/13-559; C, FG 619/2-115; D, FG 619/8-559; E, FG 619/3-559; F, FG 619/18-115; G, FG 619/1-115; H, FG 619/7-137; I, FG 619/2-559; J, FG 619/8-362; K, FG 619/13-362; L, FG 619/17-362; M, FG 619/1-559; N, FG 619/1-477; O, FG 619/9- 559; P, FG 619/11-137; Q, FG 619/11-559; R, FG 619/12-559; S, FG 619/12b-362, T, FG 619/16-362; U, FG 619/3-115; V, FG 619/4-115; W, FG 619/11-362; X, FG 619/8-115; Y, FG 619/20-115; Z, FG 619/15-137; A’, FG 619/15-559; B’, FG 619/19-115; C’, FG 619/5-115; D’, FG 619/8-115; E’, FG 619/6-115. AAP Memoir 42 (2011) 163 164 AAP Memoir 42 (2011) AAP Memoir 42 (2011) 165

Ma’in (uppermost Numayri Member); usual main arguments for a non-echinoderm relation components in bioclastic limestones of various are the occurrence of hollow tubes instead of stratigraphic levels of the Numayri Member. non-hollow trabeculae, in two apatitic layers, and in non-phosphatisation of echinoderm Description. A wide range of morphology is ossicles within the same sample. However, these represented by disarticulated plates with micro- arguments are rather weak regarding the complex porous stereom ultrastructure. But, there are more diagenetic alteration of such remains. The hollow or less three characteristic morphological types: tubes (which can be observed also on ‘true’ (1) plates with very distinct epispires (Fig. 6A-L), echinoderm remains, e.g. with epispires) may (2) pentamerous and irregular to slightly rounded very well represent coating processes which can plates (Fig. 6M-W), and (3) irregular brick-like be observed not only in echinoderms, but also on segments (Fig. 6X-Z, A’-E’). many Cambrian shelly microfossils from Jordan and elsewhere (e.g. Shinaq & Elicki 2007, p. 266 Discussion. The value of echinoderm residues fig. 7.14; Zamora et al. in press, fig. 5B-C; from preserved as isolated ossicles is generally limited. Turkey by personal observation of the author). Only in some cases it is possible to suggest It is quite common that microfossils from the systematic affinity. Nevertheless, such remains same horizon and sample show very different have been mentioned from carbonates of many preservation: shell-pseudomorphs, phosphatic early to middle Cambrian successions worldwide coatings and steinkerns. The preservation reported such that echinoderms have obviously represented by Clausen & Álvaro (2006) is also known from an important constituent of shallow marine many other localities, and is observed in the carbonate habitats at this time. material from Jordan. It is interpreted here as a The morphologies mentioned above are typical diagenetic phenomenon. for edrioasteroids and eocrinoids, but some of them do also occur in cincta. Ossicles of type 3 Phylum MOLLUSCA Cuvier, 1797 morphology (Fig. 6X-Z, A’-E’) may represent Class HELCIONELLOIDA Peel, 1991 brachiole or ambulacral parts of an eocrinoid or Order HELCIONELLIDA Geyer, 1994 ring elements surrounding the central platform Family HELCIONELLIDAE Wenz, 1938 of an edrioasteriod. Plates of types 1 and 2 (Fig. 6A-W) are probably edrioasteroid interambulacral gen. et sp. indet. (Fig. 7A-O) or cover plates, or eocrinoid thecal elements (compare Smith 1985, p. 718, text-fig. 2, p. 722, Material. Dozens of silicified specimens text-fig. 5). Interestingly, no columnal (barrel- exclusively from Timna area (higher Nimra shaped) stem elements, no typical cinctan plates, Member of Timna Formation). The specimens and also no stylocone segments have been have kindly been provided by Professor Mike recovered from the huge number of remains; Bassett (Cardiff University). different from other Gondwanan occurrences of comparative age (e.g., Elicki et al. 2003; Clausen Description. Cap-shaped, cyrtoconic, planspiral 2004; Elicki 1994, 2006; Shinaq & Elicki 2007; univalves with wide aperture. Aperture is Elicki & Gürsu 2009; Clausen & Smith 2008; somewhat elliptical in outline due to slightly Zamora et al. 2009; Zamora 2010). compressed shape of cap. The apex is distinctly The echinoderm remains from the Jordanian coiled (but less than a complete whorl), not only successions suggest eocrinoid and edrioasteroid tilted or bent. The apex is partly isolated by a weak affinity. Internal ultrastructure of some of the constriction. The margin of the shell is planar remains seems to be identical to those reported and slightly curved posteriorly. Ornamentation by Kruse et al. (2004) from the Ord Basin consists of numerous fine radiating ridges; faint (Australia) and by Clausen & Álvaro (2006) from grow lines are occasionally preserved. the Cantabrian Mountains and interpreted by the latter authors as non-echinoderm problematicum Discussion. The preservation of the specimens (?lobopodian Cantabria labyrinthica Clausen & is poor. The coarsely silicified material does not Álvaro, 2006). As stated by these authors, the allow a detailed determination. In only few cases Figure 7. Helcionellid gen. et sp. indet. from the Nimra Member of the Timna Formation (Israel). All scale bars 100 µm. A1, FG 619/7-200; A2, oblique lateral view of same specimen FG 619/7-200; A3, half-lateral view of apical area of same specimen FG 619/7-200; B, FG 619/1-200; C, FG 619/5-ISR3B; D, FG 619/2-ISR3; E, FG 619/3-ISR3; F, FG 619/1-ISR3; G, FG 619/13-200; H, FG 619/6-ISR3B; I, FG 619/6-200; J, FG 619/12-200; K1, FG 619/4-200; K2, half-lateral view of apical area of same specimen FG 619/4-200; L, FG 619/8-200; M1; FG619/10-200; M2, detail of same specimen FG 619/10-200; N1, FG 619/17-200; N2, FG 619/17-200; O, FG 619/18-200. 166 AAP Memoir 42 (2011) very faint radiating rugae and growth lines can be early Cambrian skeletal fauna and mostly grouped observed. The general shape and kind of coiling within the class Coeloscleritomorpha Bengtson & indicate a helcionellid affinity. Missarzhevsky (1981). They are known mainly The remarkable radiating ridges are also from Asia (China, Mongolia, Siberia), but have reported from other Cambrian univalved been reported from France, Australia, Iran and molluscs. Runnegar & Jell (1976) erected a Greenland (Luo et al. 1984; Missarzhevsky species of Latouchella with such characteristics 1989; Qian 1989; Kerber 1988; Qian & Bengtson (L. penecyrano), but Latouchella has a stronger 1989; Bengtson et al. 1990; Hamdi 1995; coiling and a more elongate ventral margin Eskova & Zhegallo 1996; Steiner et al. 2004; than the specimens from Israel. Regarding the Skovsted 2006). Whilst most occurrences are general shape and ornamentation, there is also from the lower portion of the early Cambrian, some similarity to the genera Kutanjia (middle some siphogonuchitids (especially from the Cambrian of the Northern Territory, Australia; occurrences outside Asia, see citations above) Kruse 1991), Kalbyella (middle Cambrian of are reported from younger strata, up to the Denmark and Australia; figured by Berg-Madsen traditional Early-Middle Cambrian boundary & Peel 1978, and by Runnegar in Bengtson et al. interval. However, this stratigraphic phenomenon 1990) and to Yochelcionella (as in Atkins & Peel could rather reflect the occurrence of special 2004). Nevertheless, distinct further differences in palaeoecological conditions and lithofacies than shell morphology do not allow suggestion of any a true palaeogeographic migration pattern. closer affinity of the specimens described here. The systematic position of monoplacophorans Hyolithelminths (Fig. 8F-J) and helcionellids is under debate and they are affiliated to various higher taxonomic groups Material. Six incomplete specimens. All come by different authors. For major discussion of the from bioclastic limestones of the higher portion systematics and on the palaeobiological aspects of the Wadi Zerqa Ma’in section (uppermost see Bengtson et al. (1990), Peel (1991), Runnegar Numayri Member). (1996), Gubanov & Peel (2000) and Parkhaev (2002). Discussion. The palaeobiology, taxonomy and systematic affiliation of hyolithelminths ?Siphogonuchitids (Fig. 8A-E) (interpreted as annelids or cnidarians) is not clear and is still controversial (compare e.g. Material. From limestone of the Wadi Zerqa Landing 1988, Brasier in Cowie & Brasier 1989, Ma’in section (uppermost Numayri Member) Bengtson in Bengtson et al. 1990, Paterson et come about ten fragments grouped together here al. 2007; Topper et al. 2009; Skovsted & Peel as probable siphogonuchitids. 2011). The original diagnostic definition of the only two families (torellellids and hyolithellids) Description. All specimens are fragmentary. is rather imprecise and highly controversial too. The remains are asymmetrical and of phosphatic Specimens mentioned here, are formally related in mineralogy (EDX analysis data). Their cross- the ‘classical’ sense into those with circular cross- section is angled to irregularly subtriangular sections (hyolithellid) and those with flattened with rounded corners. No distinct keel is visible. cross-sections (torellellid). Preservation is in A characteristic feature is the ornamentation of phosphatic mineralogy, but mostly in steinkerns. transverse, densely spaced lineations (of about Hyolithellid hyolithelminths (Fig. 8H-J) have 5-10 µm). A basal opening (rounded and slightly been recognised by three fragmentary preserved elongate) can be seen in a few cases (Fig. 8C-D). steinkerns of regular, nearly straight (extremely The apex is unknown due to poor preservation. low angle of widening) tubes with circular cross-section. No tapering is visible. The most Discussion. Skovsted (2006) described very conspicuous character is the prominent growth similar microfossils from the upper lower annulations (regularly perpendicular to the tube Cambrian Bastion Formation of eastern Greenland direction) which occur in distances of commonly as “siphogonochitids?”. Skeletal elements from 40–50 µm, but sometimes this distance is reduced southern China and northern Iran determined to 20 µm (Fig. 8H-I). In one specimen (Fig. 8J), as Siphogonuchites and Lopochites (Qian & very fine longitudinal corrugation can be observed Bengtson 1989; Hamdi 1995; Steiner et al. 2004) between the growth lines. However, it has to be are also similar to the remains from Jordan, but stressed that this specimen represents a steinkern the preservation of the latter is much too poor for and that these features reflect the inner surface more detailed determination. of the tube. Siphogonuchitids are typical elements of the Torellellid hyolithelminths (Fig. 8F-G) have AAP Memoir 42 (2011) 167

Figure 8. Unassigned small shelly fossils from Wadi Zerqa Ma’in section (uppermost Numayri Member). All scale bars 100 µm. A-E, ?Siphogonuchitids; A, FG 619/32133-451; B, FG 619/32150-451; C, FG 619/19-362; D, FG 619/20-362; E, FG 619/22-115; F-G, torellellid hyolithelminths; F, FG 619/31-451; G1, FG 619/21- 451; G2, lateral view of same specimen FG 619/21-451; H-J, hyolithellid hyolithelminths; H, FG 619/7-362; I, FG 619/20-451; J, FG 619/6-362; K-L, ?cornulitids; K1, FG 619/32092-362; K2, detail of same specimen FG 619/32092-362; L, FG 619/32148-451; M-O, opercula of hyoliths; M, FG 619/45-133; N, FG 619/46-133; O, FG 619/50-133; P, bradoriid, FG 619/7-115; P1, lateral view; P2, posterolateral view. been identified by two fragments of phosphatic conical, thick-walled tubes have flattened cross- tubular steinkerns. The features are mostly sections, but in early growth stages it is more identical to hyolithellids except in cross-section circular. The only larger fragment shows distinct and angle of divergence. Here, the tube is flattened and irregular sinuosity. The angle of tube widening so the cross-section is elliptical. The ellipticity is very low and not constant. Outer surface is is variable when comparing both specimens, but ornamented by pronounced transverse annulations also within the same specimen, and illustrates (rings), running characteristically irregular so that the large morphological variability of these they often come closer until fusion, producing hyolithelminths. a somewhat cellular wall structure (Fig. 8K-L). Longitudinally, distinct and densely arranged ?Cornulitids (Fig. 8K-L) striae between the rings are remarkable (Fig. 8K2). These characters coincides in some extant Material. Two incomplete specimens from the to torellellid hyolithelminths (mineralogy, cross- uppermost Numayri Member of the Wadi Zerqa section, general shape). The transition from Ma’in section. Shinaq & Elicki (2007, p. 266, circular to elliptical cross-section is already fig 7.23-7.24) referred them to Torellella and known from such specimens (e.g. Rozanov stressed the problematic taxonomy because of 1982; Missarzhevsky 1989). In contrast, the very some anomalous characters in the specimens special structure of the characteristic annulations from Jordan. which led to a kind of cellular appearance of the outer wall, and the sturdy striation between Discussion. The two tube fragments are of these rings is not known from hyolithelminths in phosphatic mineralogy (EDX analysis data). The general. Additionally, although some sinuosity 168 AAP Memoir 42 (2011) is already described for Torellella (e.g. Rozanov Discussion. The specimen is of phosphate 1982; Missarzhevksy 1989; Elicki 1994), such composition (EDX analysis data) and has a a high degree as in the material from Jordan bivalved preplete carapace of asymmetrical, has not previously been recorded. However, sub-semicircular outline. Valves gape anteriorly these characteristics are typical for cornulitids, a a little. The dorsal margin is straight and has problematic group described from the Ordovician rounded, but angular corners anteriorly and to Carboniferous and controversial interpreted as posteriorly (Fig. 8P1). The anterodorsal area of the annelids, tentaculitids, microconchids, cnidarians, carapace is slightly flattened close to the margin. molluscs, bryozoans or phoronids (Fisher 1962; Outer surface is smooth to somewhat corrugate; Morris & Mollins 1971; Morris & Felton 2003; no lobes, ridges or other ornamentation is visible. Herringshaw et al. 2007; Zhan & Vinn 2007). The The large depression of the carapace (Fig. 8P1) is only rather problematic feature in the specimens a secondary feature due to compaction, which is from Jordan is their comparatively small size. interpreted here as an indication of some organic Until more material is available, the specimens constituent in a non-calcitic carapace reacting described here are classified as problematica flexibly. Due to the limited morphological with probable cornulitid affinity. If the assumed features, a detailed determination is problematic. affinity is confirmed, the material from Jordan would represent the oldest occurrence of this CONCLUSIONS fossil group. For the first time, the Cambrian carbonates of the Jordan Rift Valley have been Hyolith opercula (Fig. 8M-O) micropalaeontologically investigated. During the last few years, several research projects Material. Five, partly corroded specimens from focusing on the Cambrian successions of Jordan the Numayri Member of the Wadi Umm Jafna (Burj Formation) have been initiated (Elicki et section. al. 2002; Elicki 2007a; Shinaq & Elicki 2007; Elicki submitted; Geyer & Elicki submitted; Discussion. Three different morphotypes of Hofmann et al. submitted). This paper presents hyolith opercula were found. The specimens are the micropalaeontological outcomes of this preserved as very fragile phosphatic moulds of research and the first microfauna published from different morphology. this region. The first type (Fig. 8O) has a rather circular The carbonates have been deposited in outline with relatively narrow but long and rather a structured complex of shallow marine straight to slightly outwardly curved clavicles environments under tropical/subtropical (indicated by the ridge-like elevations of the conditions (Shinaq & Elicki 2007; Elicki et al. phosphate envelope). The clavicles enclose an 2002; Elicki submitted). The development of the angle of about 50°. The second type (Fig. 8N) carbonate platform (Numayri Member) in the has a subrounded to slightly triangular outline. early Series 3 of the Cambrian (near the traditional Clavicles are well developed and are slightly Early-Middle Cambrian boundary interval) begins curved comarginally. They are closer to the margin and ends with stromatolitic fabrics. Carbonate of the operculum as in the first morphotype and facies types in between are rather diversified seemingly broader and shorter. The angle between and yield a lot of shelly macro- and microbiota. the clavicles is about 80°. The third type (Fig. 8M) Whereas the former are represented by trilobites, has a quite different, distinctly triangular outline. brachiopods and hyoliths, the microfauna contains The ventral side is rather flat and the two flanks poriferids, echinoderms, helcionellid molluscs, enclose an angle of about 120° meeting each other chancelloriids, hyolithelminths, bradoriids by a pointed peak. Clavicles are rather short and and some problematica, as well as very rare run comarginally. (?)octocorals, (?)siphogonochitids and (?) The general outline and clavicle configuration cornulitids. Because of the diagenetic overprint of type 1 correspond to Hyolithes kingi, the and related large mineral replacement, as well as very common and only described hyolith of the due to the lack of knowledge on the morphological Numayri Member. diversity of some skeletal taxa (e.g., poriferids, echinoderms), a detailed taxonomic determination Bradoriids (Fig. 8P1-P2) is not possible in several cases. Nevertheless, two genera and one species of poriferids, four Material. One specimen from limestone of the genera and one species of chancellorrids, and one Wadi Zerqa Ma’in section (uppermost Numayri probable octocoral species have been identified. Member). By far the most of the microfossils belong to echinoderms and poriferids. Isolated AAP Memoir 42 (2011) 169 echinoderm ossicles could be identified as (Uppsala, Sweden) provided important comments eocrinoid and edrioasteroid elements. Among and suggestions which improved the paper. Rafie poriferids, hexactinellid spicules (Rigbyella Shinaq (Irbid, Jordan), Thomas Wotte (Münster, sp. and triactins, tetractins, pentactins, and Germany) and Thomas Biener (Ludwigsfelde, hexactins) are predominant. Heteractinids Germany) helped significantly during field work (Eiffelia araniformis [Missarzhevsky, 1981]) in Jordan. Jan Fischer (Freiberg, Germany) is and polyactinellids (?Praephobetractinia) are thanked for technical assistance. Field work rather rare. Chancelloriids are not very common would not have been possible without the and are represented by Archiasterella cf. hirundo generous assistance provided by the Natural Bengtson, 1990, Allonnia sp., Chancelloria sp., Resources Authority of Jordan (Amman, Jordan). and ?Ginospina sp. Molluscs are known from Their long-lasting and fruitful cooperation is the microfossil content of Jordan by only few greatly appreciated. Investigations have received opercula of hyoliths. From the Nimra Member of significant support from the German Research the Timna area in Israel (which coincides to the Foundation (DFG, Germany). Numayri Member of Jordan), a poorly preserved helcionellid mollusc has been identified in large REFERENCES numbers. Microcoryne cephalata (Bengtson, Amirieh, B.S., Schneider, W. & Abed, A.M., 1994. 1990) – a probable octocoral sclerite – is Evolving fluvial-transitional-marine deposition represented by one specimen. Few specimens of through the Cambrian sequence of Jordan. torellellid and hyolithellid hyolithelminths were Sedimentary Geology 89, 65-90. able to be determined. Bradoriid arthropods are Amirieh, B.S., Ameireh, M.N. & Abed, A.M., 2008. represented by one specimen. Some rare and Tectono sedimentary evolution of the Umm Ghaddah broken microfossils point to a siphogonuchitid Formation (late Ediacaran–early Cambrian) in affinity. The occurrence of a probably cornulitid Jordan. Journal of Asian Earth Sciences 33, 194- related problematicum is reported; if correct, this 218. would represent the oldest occurrence of this Atkins, C.J. & Peel, J.S., 2004. New species of fossil type so far. Yochelcionella (Mollusca ; Helcionelloida) from the The diversity of the microfossil assemblage is Lower Cambrian of North Greenland. Bulletin of the not very large. The fauna is clearly dominated by Geological Society of Denmark 51, 1-9. sessile (and minor mobile) epibenthic forms. Few Bandel, K., 1986. The reconstructionof “Hyolithes representatives may have realised a necto-benthic kingi” as annelid worm from the Cambrian of Jordan. life-style. Most organisms seem to have been Mitteilungen des Geologisch-Paläontologischen suspension feeders with fewer deposit feeders. Instituts der Universität Hamburg 61, 35-101. Such type of microfauna is well known Bandel, K. & Shinaq, R., 2003. Sediments of the from many late early to middle (Series 3) Precambrian Wadi Abu Barqa Formation influenced Cambrian low-latitude carbonate environments. by life and their relation to the Cambrian sandstones Nevertheless, some taxa from the Jordan Rift in southern Jordan. Paläontologie, Stratigraphie, Valley may support a somewhat closer relation to Fazies 11 (Freiberger Forschungshefte C 499), Iran and Australia (e.g., Rigbyella, Archiasterella 78-91. cf. hirundo Bengtson, 1990) which were part of Bender, F., 1974. Geology of Jordan. Bornträger, equatorial Gondwana. Some connection may have Stuttgart, 196 p. existed to the European shelf of Perigondwana, Bengtson, S., 1986. Siliceous microfossils from the too, as suggested by the shared occurrence of Upper Cambrian of Queensland. Alcheringa 10, Microcoryne cephalata (Bengtson, 1990) and 195-216. again of Archiasterella cf. hirundo Bengtson, Bengtson, S. & Missarzhevsky, V.V., 1981. 1990. Coeloscleritophora – a major group of enigmatic Cambrian metazoans. US Geological Survey Open- ACKNOWLEDGEMENTS file Report 81-743, 19-21. I thank Michael Bassett (Cardiff, UK) and Bengtson, S., Conway Morris, S., Cooper, B.J., Jell, Leonid Popov (Cardiff, UK) for making available P.A. & Runnegar, B.N., 1990. Early Cambrian microfossils from their own field work in Israel. fossils from South Australia. Memoirs of the Important suggestions in poriferan investigation Association of Australasian Palaeontologists 9, came from Joachim Reitner (Göttingen, Germany) 364 p. and Dorte Mehl (Frankfurt/Main, Germany). Berg-Madsen, V. & Peel, J.S., 1978. Middle Cambrian Pierre Kruse (Adelaide, Australia) kindly helped monoplacophorans from Bornholm and Australia, in discussing helcionellids. Glenn A. Brock and the systematic position of the bellerophontiform (Sydney, Australia) is deeply thanked for help molluscs. Lethaia 11, 113-125. with English. He and Christian B. Skovsted Clausen, S., 2004. New Early Cambrian eocrinoids 170 AAP Memoir 42 (2011)

from the Iberian Chains (NE Spain) and their role in at the northern edge of the Arabian-Nubian Shield: nonreefal benthic communities. Eclogae geologicae sedimentary facies architecture and palaeoecological Helvetiae 97, 371-379. context. Journal of Asian Earth Sciences. Clausen, S. & Álvaro, J.J., 2006. Skeletonized Elicki, O., Altumi, M.M., Leuschke, A., Farouk, S. microfossils from the Lower-Middle Cambrian & Hofmann, R., 2010. Cambrian trace fossils from transition of the Cantabrian Mountains, northern the north-eastern Africa - Middle East segment of Spain. Acta Palaeontologica Polonica 51(2), 223- Perigondwana. Zitteliana, series B 29, 31-32. 238. Elicki, O. & Geyer, G., 2010. Cambrian biostratigraphy. Clausen, S. & Smith, A.B., 2008. Stem structure and 103-114 in Linnemann, U. & Romer, R.L. (eds.), evolution in the earliest pelmatozoan echinoderms. Pre-Mesozoic Geology of Saxo-Thuringia – From Journal of Paleontology 82, 737-748. the Cadomian Active Margin to the Variscan Cooper, G.A., 1976. Lower Cambrian brachiopods Orogen. Schweizerbart, Stuttgart. from the Rift Valley (Israel and Jordan). Journal of Elicki, O. & Gürsu, S., 2009. First record of Pojetaia Paleontology 50, 269-289. runnegari Jell, 1980 and Fordilla Barrande, Cowie, J.W. & Brasier, M.D., 1989. The Precambrian- 1881 from the Middle East (Taurus Mountains, Cambrian Boundary. Clarendon Press, Oxford, Turkey) and critical review of Cambrian bivalves. 213 p. Paläontologische Zeitschrift 83(2), 267-291. Culver, S.J., Repetski, J.E., Pojeta, J. & Hunt, D., Elicki, O., Hamann, Y. & Münzberger, P., 2003. 1996. Early and Middle(?) Cambrian metazoan Biofazies, Paläoökologie und Ablagerungsmodell and protistan fossils from West Africa. Journal of des Finalstadiums einer kambrischen Paleontology 70, 1-6. Karbonatplattform-Entwicklung: die Fauna Dienemann, W., 1915. Älteres Paläozoikum von der Campo Pisano Formation SW-Sardiniens. Südsyrien und Westarabien. Centralblatt für Paläontologie, Stratigraphie, Fazies 11 (Freiberger Mineralogie, Geologie und Paläontologie 16, Forschungshefte C 499), 1-33. 23-26. Elicki, O., Schneider, J.W. & Shinaq, R., 2002. Dore, F. & Reid, R.E., 1965. Allonia tripodophora Prominent facies from the Lower/Middle nov. gen, nov. sp., nouvelle éponge du Cambrien Cambrian of the Dead Sea area (Jordan) and their inférieur de Carteret (Manche). Comptes Rendus palaeodepositional significance. Bulletin de la Sommaires des Séances de la Société géologique Société géologique de France 173(6), 147-152. de France, 20-21. Elicki, O. & Shinaq, R., 2000. Kambrische Lagunen- Duan, C., 1984. Small shelly fossils from the Lower Karbonate vom südlichen Toten Meer (Wadi Tayan, Cambrian Xihaoping Formation in the Shennongjia Jordanien). Paläontologie, Stratigraphie, Fazies 8 District, Hubei Province – hyoliths and fossil (Freiberger Forschungshefte C 490), 51-66. skeletons of unknown affinities. Bulletin of the Esakova, N.V. & Zhegallo, E.A., 1996. Biostratigrafiya Tianjin Institute of Geology and Mineral Resources, i fauna nizhnego kembriya Mongolii. Trudy, 7, 143-188. Sovmestnaya Rossiysko-Mongolskaya Elicki, O., 1994. Lower Cambrian carbonates from Paleontolicheskaya Ekspeditsiya 46, 214 p. eastern Germany: palaeontology, stratigraphy and Fernández-Remolar, D.C., 2001. Chancelloriidae del palaeogeography. Neues Jahrbuch für Geologie und Ovetiense inferior de la Sierra de Córdoba, España. Paläontologie, Abhandlungen 191/1, 69-93. Revista Española de Paleontología 16, 39-61. Elicki, O., 2005. The utility of late Early to Middle Fisher, D.W., 1962. Small conoidal shells of uncertain Cambrian small shelly fossils from the western affinities. 98-14 in Moore, R.C. (ed.), Treatise of Mediterranean. Geosciences Journal 9(2), 161-171. Invertebrate Paleontology, Part W – Miscellanea. Elicki, O., 2006. Microbiofacies analysis of Cambrian Geological Society of America, New York and offshore carbonates from Sardinia (Italy): University of Kansas Press, Lawrence. environment reconstruction and development of a Finks, R.M., Reid, R.E.H. & Rigby, J.K., 2004. drowning carbonate platform. Carnets de Géologie Treatise on invertebrate paleontology, part E: 2006/01, 1-22. porifera – revised, vol. 3: Porifera (Demospongea, Elicki, O., 2007a. Facies development during late Hexactinellida, Heteractinida, Calcarea). Early-Middle Cambrian (Tayan Member, Burj Geological Society of America, Boulder and Formation) transgression in the Dead Sea Rift University of Kansas Press, Lawrence, 872 p. Valley, Jordan. Carnets de Géologie 2007/07, 1-20. Geyer, G. & Elicki, O., submitted. Cambrian trilobites Elicki, O., 2007b. Paleontological data from the Early of Jordan: taxonomy, stratigraphic affinity, and Cambrian of Germany and paleobiogeographical functional morphology revised. Acta Geologica implications for the configuration of central Polonica. Perigondwana. Geological Society of America, Geyer, G. & Landing, E., 2004. A unified Lower – Special Paper 423, 143-152 Middle Cambrian chronostratigraphy for West Elicki, O., submitted. A Cambrian carbonate platform Gondwana. Acta Geologica Polonica 54, 179-218. AAP Memoir 42 (2011) 171

Gubanov, A.P. & Peel, J., 2000. Cambrian 154 p. monoplacophoran molluscs (Class Helcionelloida). Makhlouf, I.M., 2003. Braided river model and American Malacostracal Bulletin 15(2), 139-145. associated facies of lower Cambrian age in South Hamdi, B., 1995. Precambrian-Cambrian deposits in Jordan. Africa Geoscience Review 10(3), 289-300. Iran. Treatise on the Geology of Iran 20, 353 p. Makhlouf, I.M. & Abed, A.M., 1991. Depositional Herringshaw, L.G., Thomas, A.T. & Smith, M.P., 2007. facies and environments in the Umm Ishrin Systematics, shell structure and affinities of the Sandstone Formation, Dead Sea area, Jordan. Palaeozoic Problematicum Cornulites. Zoological Sedimentary Geology 71, 177-187. Journal of the Linnéan Society 150 , 681-699. Mángano, M.G. , Hofmann, R., Elicki, O. & Shinaq, Hinz-Schallreuter, I., 1993. Cambrian ostracodes R., 2007. Paleoecologic and paleoenvironmental mainly from Baltoscandia and Morocco. Archiv für controls of trace fossils from a Lower to Middle Geschiebekunde I/7, 385-448. Cambrian tide-dominated delta, Hanneh Member, Hofmann, R., Mángano, M.G. & Elicki, O., 2008. Burj Formation, Southern Death Sea, Jordan. 101 Multiple behaviours in under-utilised ecospace: in MacEachern, J.A., Gingras, M.K., Bann, K.L. & insights from a Cambrian marine system. Erlanger Pemberton, S.G. (eds.), Ichnological applications Geologische Abhandlungen, Sonderband 6, 91. to sedimentological and sequence stratigraphic Hofmann, R., Mangano, M.G., Elicki, O. & Shinaq, problems. SEPM Research Conference, Price, Utah, R., submitted. Paleoecologic and biostratigraphic USA, May 20-26, abstract volume. significance of trace fossils from Middle Cambrian Mehl, D., 1995. Phylogenie und Evolutionsökologie shallow- to marginal-marine environments, Hanneh der Hexactinellida (Porifera) im Paläozoikum. Member, Burj Formation, Southern Dead Sea, Geologische und Paläontologische Mitteilungen Jordan. Journal of Paleontology. der Universität Innsbruck, Sonderband 4, 1-55. Hooper, J.N.A. & Van Soest, R.W.M., 2002. Systema Mehl, D., 1998. Porifera and chancelloriidae from the Porifera – A guide to the classification of sponges. Middle Cambrian of the Georgina basin, Australia. Kluwer Academic/Plenum Publishers, 1708 p. Palaeontology 41, 1153-1182. Kerber, M., 1988. Mikrofossilien aus unterkambrischen Missarzhevsky, V.V., 1981. Early Cambrian hyoliths Gesteinen der Montagne Noire, Frankreich. and gastropods of Mongolia. Paleontologicheskiy Palaeontographica, Abt. A 202, 127-203. Zhurnal 1, 21-28. King, W.B.R., 1923. Cambrian Fossils from the Dead Missarzhevsky, V.V., 1989. Oldest skeletal fossils and Sea. Geological Magazine 60, 507-514. stratigraphy of Precambrian and Cambrian boundary Kozur, H., 1991. Erster Nachweis von Polyactinellidae beds. Trudy Gologicheskiy Instituta 443, 236 p. Mostler (Calcispongea, Porifera) im Perm und Missarzhevsky, V.V. & Mambetov, A.M., 1981. Revision der Gattung Phobetractinia Reif. Stratigraphy and fauna of the Precambrian- Zeitschrift für Geologische Wissenschaften 19(5), Cambrian boundary beds of Malyj Karatau. Trudy 585-591. Geologicheskiy Instituta 326, 1-92. Kruse, P.D., 1991. Cambrian fauna of the Top Springs Moore, J.L., Porter, S.M., Steiner, M. & Li, G., Limestone, Georgina Basin. The Beagle, Records of 2010. Cambrothyra ampulliformis, an unusual the Northern Territory Museum of Arts and Sciences coeloscleritophoran from the Lower Cambrian of 8(1), 169-188. Shaanxi Province, China. Journal of Paleontology Kruse, P.D., Laurie, J.R. & Webby, B.D., 2004. 84, 1040-1060. Cambrian Geology and palaeontology of the Ord Morris, R.W. & Mollins, H.B., 1971. The distribution Basin. Memoirs of the Association of Australasian and palaeoecological interpretation of Cornulites Palaeontologists 30, 1-58. in the Waynesville Formation (Upper Ordovician) Landing, E., 1988. Lower Cambrian of Eastern of southwestern Ohio. Ohio Journal of Science Massachusetts: stratigraphy and small shelly fossils. 71(3), 159-170. Journal of Paleontology 62, 661-695. Morris, R.W. & Felton, S.H., 2003. Paleoecologic Laurie, J.R., 1986. Phosphatic fauna of the early Associations and Secondary Tiering of Cornulites Cambrian Todd River Dolomite, Amadeus Basin, on Crinoids and Bivalves in the Upper Ordovician central Australia. Alcheringa 10, 431-454. (Cincinnatian) of Southwestern Ohio, Southeastern Luo Huilin, Jiang Zhiwen, Wu Xiche, Song Xueliang & Indiana, and Northern Kentucky. Palaios 18, 546- Ouyang Lin, 1982. The Sinian-Cambrian boundary 558 in eastern Yunnan. People’s Publishing House, M o s t l e r , H ., 1985. Neue heteractinide Yunnan, 265 p. Spongien (Calcispongea) aus dem Unter- und Luo Huilin, Jiang Zhiwen, Wu Xiche, Song Xueliang, Mittelkambrium Südwestsardiniens. Berichte Ouyang Lin, Xing Yusheng, Liu Guizhi, Zhang des naturwissenschaftlich-medizinischen Vereins Shishan, Z. & Tao Yonghe, 1984. Sinian-Cambrian Innsbruck 72, 7-32. boundary stratotype section at Meishucun, Jinning, Mostler, H., 1996. Polyactinellide Schwämme, eine Yunnan, China. People’s Publishing House, Yunnan, auf das Paläozoikum beschränkte Calcispongien- 172 AAP Memoir 42 (2011)

Gruppe. Geologisch-Paläontologische Mitteilungen Rozanov, A.Yu., 1982. The Precambrian/Cambrian Innsbruck 21, 223-243. boundary in the geosynclinals areas (the reference Mostler, H. & Mosleh-Yazdi, A., 1976. Neue section of Salany-Gol, MPR). Nauka, Moscow, Poriferen aus oberkambrischen Gesteinen der 150 p. Milaformation im Elbruzgebirge (Iran). Geologisch- Runnegar, B., 1996. Early evolution of the mollusca: Paläontologische Mitteilungen Innsbruck 5(1), the fossil record. 77-87 in Taylor, J (ed.), Origin 1-36. and evolutionary radiation of the mollusca. Oxford Müller, K., 1964. Ostracoda (Bradorina) mit University Press. phosphatischen Gehäusen aus dem Oberkambrium Runnegar, B. & Jell, P.A., 1976. Australian Middle von Schweden. Neues Jahrbuch für Geologie und Cambrian molluscs and their bearing on early Paläontologie, Abhandlungen 121, 1-46. molluscan evolution. Alcheringa 1, 109-138. Nötzold, T., 1965. Die Präparation von Gyrogoniten Rushton, A.W.A. & Powell, J.H., 1998. A review of the und kalkigen Charophyten-Oogonien aus festen stratigraphy and trilobite faunas from the Cambrian Kalksteinen. Monatsberichte der deutschen Burj Formation in Jordan. Bulletin of the Natural Akademie der Wissenschaften 7, 216-221. History Museum London (Geology) 54(2), 131-146. Parkhaev, P.Yu., 2002. Phylogenesis and the system of Sarmiento, G.N., Fernández-Remolar, D. & Cambrian univalved molluscs. Paleontologicheskiy Göncüoglu, M.C., 2001. Cambrian small shelly Zhurnal 36, 25-39. fossils from the Çal Tepe Formation, Taurus Parnes, A., 1971. Late Lower Cambrian trilobites Mountains, Turkey. Coloquios de Paleontología from the Timna area and Har’Amram (southern 52, 117-134. Negev, Israel). Israel Journal of Earth Sciences Schneider, W., Amireh, B.S. & Abed, A.M., 2007. 20, 179-205. Sequence analysis of the early Paleozoic Paterson, J.R., Skovsted, C.B., Brock, G.A. & sedimentary systems of Jordan. Zeitschrift der Jago, J.B., 2007. An early Cambrian faunule Deutschen Gesellschaft für Geowissenschaften from the Koolywurtie Limestone Member (Parara 158(2), 225-247. Limestone) Yorke Peninsula, South Australia and Sdzuy, K., 1969. Unter- und mittelkambrische its biostratigraphic significance. Memoirs of the Porifera (Chancelloriida und Hexactinellida). Association of Australasian Palaeontologists 34, Paläontologische Zeitschrift 43, 115-147. 131-146. Seilacher, A., 1990a. Paleozoic trace fossils. 649- Peel, J.S., 1991. The classes Tergomya and 670 in Said, R. (ed.), The Geology of Egypt. A.A. Helcionelloida, and early molluscan evolution. Balkema Rotterdam. Bulletin Grønlands Geologiske Undersøgelse 161, Seilacher, A., 1990b. Cruziana stratigraphy of „non- 11-65. fosiliferous“ Palaeozoic sandstones. Geological Powell, J.H., 1989. Stratigraphie and sedimentation of Journal 3, 447-476. the phanerozoic rocks in Central and South Jordan, Shinaq, R. & Bandel, K., 1992. Microfacies of Part A: Ram and Khreim Groups. Natural Resources Cambrian limestones in Jordan. Facies 27, 52-57. Authority, Geology Directorate, Geological Shinaq, R. & Elicki, O., 2007. The Cambrian Mapping Division Bulletin 11, 1-72. sedimentary succession from the Wadi Zerqa Ma’in Qian, Y., 1989. Early Cambrian small shelly (northeastern Dead Sea area, Jordan): lithology and fossils of China with special reference to the fossil content. Neues Jahrbuch für Geologie und Precambrian-Cambrian boundary. Stratigraphy and Paläontologie, Abhandlungen 243(3), 255-271. Palaeontology of Systemic Boundaries in China, Siveter, D.J. & Williams, M., 1997. Cambrian Precambrian-Cambrian Boundary 2, 341 pp, 99 pl. bradoriid and phosphatocopid arthropods of North Qian Y. & Bengtson, S., 1989. Palaeontology and America. Special Papers in Palaeontology 57, 1-69. biostratigraphy of the Early Cambrian Meishucunian Skovsted, C.B., 2006. Small shelly fauna from Stage in Yunnan Province, South Chna. Fossils and upper Lower Cambrian Bastion and Ella Island Strata 24, 1-156. formations, north-east Greenland. Journal of Reif, W.-E., 1968. Schwammreste aus dem oberen Paleontology 80, 1087-1112. Ordovizium von Estland und Schweden. Neues Skovsted, C.B. & Peel, J.S., 2010. Early Cambrian Jahrbuch für Geologie und Paläontologie, brachiopods and other shelly fossils from the basal Monatshefte 9, 522-529. Kinzers Formation of Pennsylvania. Journal of Richter, R. & Richter, E., 1941. Das Kambrium am Paleontology 84, 754-762. Toten Meer und die älteste Tethys. Abhandlungen Skovsted, C.B. & Peel, J.S., 2011. Hyolithellus in Senckenbergiana Naturforschende Gesellschaft life position from the Lower Cambrian of North 460, 1-51. Greenland. Journal of Paleontology 85, 37-47. Rigby, J.K., 1975. Some unusual hexactinellid sponge Smith, A.B., 1985. Cambrian eleutherozoan spicules from the Cambrian Wilberns Formation of echinoderms and the early diversification of Texas. Journal of Paleontology 49, 412-415. edrioasteroids. Palaeontology 28, 715-756. AAP Memoir 42 (2011) 173

Steiner, M., Li, G., Qian, Y. & Zhu, M., 2004. north Spain show echinoderms diversified earlier in Lower Cambrian Small Shelly Fossils of northern Gondwana. Geology 38, 507-510. Sichuan and southern Shaanxi (China), and their Zamora, S., Gozalo, R. & Liñan, E., 2009. Middle biostratigraphic importance. Geobios 37, 259-275. Cambrian gogiid echinoderms from Northeast Spain: Teimeh, M., Taani, Y., Abu Lihie, O. & Abu Saad, Taxonomy, palaeoecology, and palaeogeographic L., 1990. A study of the Palaeozoic formations of implications. Acta Palaeontologica Polonica 54, Jordan at outcrop and in the subsurface including 253-265. measured sections and regional isopach maps. Zamora, S., Lefebvre, B., Álvaro, J.J., Clausen, S., Natural Resources Authority, Geology Directorate, Elicki, O., Fatka, O., Jell, P., Kouchinsky, A., Geological Mapping Division Subsurface Geology Lin, J.-P., Nardin, E., Parsley, R., Rozhnov, S., Bulletin 1, 1-72. Sprinkle, J., Sumrall, C.D., Vizcaïno, D. & Smith, Topper,T.P., Brock, G.A., Skovsted, C.B. & Paterson, A.D., in press. Cambrian echinoderm diversity and J.R., 2009. Shelly fossils from the lower Cambrian palaeobiogeography. In Harper, D. & Servais, T. Pararaia bunyerooensis Zone, Flinders Ranges, (eds), Early Palaeozoic Palaeobiogeography and South Australia. Memoirs of the Association of Palaeogeography. Geological Society Special Australasian Palaeontologists 37, 199-246. Publication, London. Wrona, R., 2004. Cambrian microfossils from glacial Zhan, R. & Vinn, O., 2007. Cornulitid epibionts on erratics of King George Island, Antarctica. Acta brachiopod shells from the Late Ordovician (middle Palaeontologica Polonica 49, 13-56. Ashgill) of East China. Estonian Journal of Earth Zamora, S., 2010. Middle Cambrian echinoderms from Sciences 56(2), 101-108.