Abstract
An Upper Devonian to (?)Lower Mississippian section at Hushoot Shiveetiin gol in the Baruunhuurai Terrane of the Central Asian Orogenic Belt (CAOB) was investigated. A generally well-preserved and diverse ostracod fauna was collected from the Late Devonian (Famennian) Samnuuruul Formation in western Mongolia. It is the first rich assemblage described from this area. The ostracod fauna consists of 19 genera and 25 species. Two of the species are proposed new species, and 13 species are described in open nomenclature. The Mongolian ostracod fauna is similar to coeval faunas in western China and Laurussia, but also contains new, endemic species. Two new taxa are described: Beyrichiopsis hushootensis and Ampuloides beckeri. Ostracod findings are characterized by the Eifelian Mega-Assemblage (I-III) representing a nearshore, variable palaeoenvironment which is in accordance with facies analysis provided by other studies.
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Introduction
Reports on Palaeozoic faunas from Mongolia are sparse. Previous papers have described conodonts, corals and echinoderms (e.g. Bolshakova et al. 2003; Webster and Ariunchimeg 2004; Ariunchimeg et al. 2014; Suttner et al. 2019), but Palaeozoic ostracods from Mongolia are previously unreported. Ostracods can provide critical data on palaeoenvironmental and palaeogeographical reconstructions. Many ostracod species and genera from the Devonian exhibit characteristic assemblages ranging from shallow water to deep marine environments (e.g. Bandel and Becker 1975; Wang 1988; Becker and Bless 1990; Becker and Blumenstengel 1995; Becker 2000; Groos-Uffenorde et al. 2000; Casier 2004; Becker et al. 2004; Crasquin and Horne 2018).
In the framework of the International Geoscience Programme (IGCP) 596 and its successor, the Western Mongolia Working Group, our research group conducted fieldwork in western Mongolia in 2012, 2014 and 2018 (Kido et al. 2013; Ariunchimeg et al. 2014; Suttner et al. 2019). Herein, we describe a diverse, generally well-preserved ostracod fauna from the Hushoot Shiveetiin gol section of the Samnuuruul Formation (Famennian, Late Devonian), Baruunhuurai Terrane and Olonbulag Subterrane (Fig. 1). The fauna provides insight on palaeoenvironmental conditions of the section as well as on taxonomic affinities with other Famennian ostracod communities in the CAOB and elsewhere, particularly with Late Devonian ostracod communities of western Junggar, NW China (Song et al. 2017).
Geological setting
The Central Asian Orogenic Belt (CAOB) is the world’s largest Palaeozoic accretionary belt. Mongolia lies in the center of the CAOB and is composed of up to 44 different terrains, including cratonic, metamorphic, passive margin, island arc, accretionary complex and ophiolitic terranes, ranging in age from Proterozoic through the Phanerozoic (Badarch et al. 2002; Xiao et al. 2010; Metcalfe 2011; Choulet et al. 2012; Yang et al. 2013; Li et al. 2017). Studies by Windley et al. (2007), Donskaya et al. (2013) and Yang et al. (2015) suggest that subduction of the Palaeo-Asian Ocean continued during the Middle and Late Palaeozoic before final closure of this ocean and its Turkestan and Junggar branches during the Late Palaeozoic to the Mesozoic (Safonova et al. 2017). Palaeozoic rocks of western Mongolia described in this paper are part of the complex tectonic framework of the CAOB (for a more detailed discussion on plate tectonics, we refer to Safonova et al. 2017, cum lit.).
Deposits of the Baruunhuurai Terrane have a stratigraphic range from Devonian to Carboniferous and are located in the southwestern part of Mongolia close to the border to China. The investigated section is located in the Olonbulag Subterrane (Fig. 1) and ranges from the Devonian (Famennian) to Carboniferous (?early Mississippian). The exposed section belongs to the Samnuuruul Formation which has a thickness of 250–450 m. A detailed facies and stratigraphic analysis of the Hushoot Shiveetiin gol section is published by Ariuntogos et al. (2020, this issue). The ostracods reported herein were collected from both sections (A and B sections) and have a stratigraphical range based on conodonts from the crepida Biozone to the rugosa trachytera Biozone. The zonation scheme used herein follows Hartenfels (2011) and Spalletta et al. (2017).
The Samnuuruul Formation is dominated by siliciclastic rocks with numerous volcaniclastic deposits. Limestones are limited, occur as thin-bedded layers, and are rich in fossils, such as corals, conodonts, crinoids, trilobites, bryozoans and ostracods. The section is divided into an eastern (A) and a western (B) part which was correlated by a diagnostic thick-bedded pyroclastic marker horizon (Fig. 2). Correlation is also proven by conodont data (Ariuntogos et al. 2020, this issue). Sedimentary analysis indicates variable depositional facies settings ranging from shallow-intertidal to open marine (for detailed sedimentology and facies analysis see Ariuntogos et al. 2020, this issue).
Materials and methods
In this study, 12 samples contain ostracod specimens as single valves and carapaces. The limestone and marl samples between 1 and 4 kg were collected from the eastern (A) and western (B) Hushoot Shiveetiin gol section (see Suttner et al. 2019, Ariuntogos et al. 2020, this issue; Fig. 2). These samples were dissolved in formic and/or acetic acids and sieved into 63 μm, 125 μm, 250 μm and 500 μm fractions. The fractions were dried in an oven at about 40 °C and the most promising conodont-sized fractions were separated by liquid sodium polytungstate. After conodont separation, the light fractions were picked for ostracods and other microfossils. The ostracods were mounted on aluminium stubs, sputtered with gold using a Quorum (Q150R ES) and imaged using a Quanta 650 Field Emission scanning electron microscope in the Central Research Laboratory, Çukurova University, Turkey. The studied ostracods are deposited in the collections of the Mongolian University of Science and Technology, under the collection numbers MUST-RCSP-Ost.00001-98.
Ostracod data
Ostracods were identified in samples from the eastern (A) and western (B) Hushoot Shiveetiin gol section ranging from the crepida into the rugosa trachytera biozones in the Famennian (Ariuntogos et al. 2020, this issue). A systematic list of identified ostracod taxa (19 genera and 25 species) were given Appendix 1 and their stratigraphical distributions in the studied area is provided in Fig. 3. The ostracod fauna of the Hushoot Shiveetiin gol section includes species belonging to the orders Palaeocopida, Platycopida and Podocopida (Figs. 3, 4, 5, 6, 7, and 8). In this study the Palaeozoic ostracod classification by Becker (2002) and Liebau (2005) was used. Ten species out of a total number of 25 species belong to previously known taxa. Two new species are proposed, and because of the materials are insufficient for formal description and naming of a new taxon, thirteen species are left in open nomenclature.
Systematic Palaeontology
The new species and open nomenclature are given in this part.
Class Ostracoda Latreille, 1802
Order Palaeocopida Henningsmoen, 1953
Superfamily Aechminacea Bouček, 1936.
Family Aechminidae Bouček, 1936.
Aechmina sp. (Fig. 4/i)
Material: Two carapaces, partly damaged
Occurrence. It was found in gracilis gracilis Biozone of Hushoot Shiveetiin gol section (Unit IV).
Superfamily Hollinoidea (=Hollinacea) Swartz 1936
Family Ctenoloculinidae Jaanusson and Martinsson, 1956
Parabolbina sp. (Fig. 4/j)
Material: Three valves, partly damaged.
Remarks: Because of the limited number of carapaces and valves, Parabolbina sp. is described in open nomenclature.
Occurrence: It was found in gracilis gracilis Biozone of Hushoot Shiveetiin gol section (Unit IV).
Superfamily Primitiopsoidea (=Primitiopsacea) Swartz 1936
Family Pribylitidae Pokorný, 1958
Coryellina sp. (Fig. 4/o)
Material: One valve.
Remarks: Coryellina sp. is poorly preserved and does not allow a precise identification.
Superfamily Kirkbyoidea (=Kirkbyacea Ulrich and Bassler 1906)
Family Amphissitidae Knight, 1928
Polytylites sp. (Fig. 5/f)
Material: Seven valves, partly damage.
Remarks: Some morphological features on valves (a rounded lobe occurs in the median part, surface reticulate) are observed. But, they are not detailed enough for formal description and naming of a new species.
Occurrence: Polytylites sp. was found in the rhomboidea Biozone of Hushoot Shiveetiin gol section, sample HS-C-1b (Unit IV).
Family Nezamysliidae Żbikowska 1983
Nezamyslia sp. (Fig. 5/g-i)
Material: Three valves, all partly damage.
Remarks: The material is poorly-preserved, thus this species cannot be determined exactly and attributed to a known or new species.
Occurrence: It is observed in the rhomboidea Biozone.
Family Kirkbyellidae Sohn, 196
Berdanella sp. (Fig. 5/j-k)
Material: 4 valves
Description: Outline subquadrate in lateral view, dorsal border straight. Median sulcus (S2) distinct. A lateral ridge with a small posteromedian lobe is on ventral half of the valve. Surface fine reticulate.
Remarks: In outline and surface fine reticulate, the specimens are close to Kirkbyella (Berdanella) unicornis (Coryell and Malkin), 1936, but differ by a lateral ridge with a small posteromedian lobe.
Dimension: L = 1.023–1.433 mm, H = 0.600–0.783 mm.
Occurrence: Berdanella sp. was found in rugosa trachytera Biozone of the Hushoot Shiveetiin gol section, sample HS-C-W1 (Unit VII).
Palaeocopina? indet. 1 (Fig. 5/l-n)
Material: Thirteen valves.
Dimensions: L = 0.600–0.690 mm, H = 0.400–0.480 mm.
Description: Carapace small, nonsulcate, amplete, straigth and long dorsal margin, cardinal angles well defined, obtuse, adventral structure developed and running almost parallel to the free margins, adductorial pit is not clear. The rows of reticulations are observed. The perpendicular ridges are dominated and subparallel to the free margin.
Remarks: The form described herein correlates with the lateral outline and ornamentation of carapace to Reticestus sp. Olempska, 1979 and Reticestus? n.sp. Zagora 1968, except anterodorsal and posterodorsal part and general shape.
Occurrence: The rugosa trachytera Biozone of the Hushoot Shiveetiin gol section, HS-C-W1 (Unit VII).
Palaeocopina? indet. 2 (Fig. 5/o, Fig. 6/a-f).
Material: 33 corroded carapaces.
Dimensions: L = 0.538–0.728 mm, H = 0.400–0.480 mm.
Description: The carapaces and valves are poorly preserved. There is a denticulate dorsal border. Indistinct shallow sulcus (S2). Sexual dimorphism presumably present. The carapace outline of male and female is different. Anterior and posterior margins of females are rounded. Male carapace is preplete (Fig. 5/o, 6/a,b,f). Female carapace is amplete (Fig. 6/c–e).
Occurrence: It occurs from the rhomboidea to the gracilis gracilis zones of the Hushoot Shiveetiin gol section (Unit IV).
Order Platycopida Sars 1866
Superfamily Kloedenelloidea (=Kloedenellacea Ulrich and Bassler 1908)
Kloedenellidae indet. (Fig. 6/g-h)
Material: Two valve
Dimension: (one specimen measured). L = 1.025 mm, H = 0.575 mm.
Occurrence: HS-C-2 includes undetermined kloedenellid ostracod in the rhomboidea Biozone of the eastern section (A) Hushoot Shiveetiin gol section.
Family Beyrichiopsidae Henningsmoen, 1953
Genus Beyrichiopsis Jones and Kirkby 1886
Type-species: Beyrichiopsis fimbriata Jones and Kirkby 1886
Beyrichiopsis hushootensis sp. nov. (Fig. 6/j-o, Fig. 7/a)
Holotype: Right valve, (Fig. 6/o, MUST-RCSP-Ost.00041)
Paratype: Right valve of paratype A (Fig. 6/m, MUST-RCSP-Ost.00038), left valve of paratype B (Fig. 7/a, MUST-RCSP-Ost.00040).
Derivation of name: The name is derived from Hushoot Shiveetiin gol section, Baruunkhuurai Terrane, Mongolia.
Type locality and horizon: The rugosa trachytera Biozone of the Hushoot Shiveetiin gol section, sample HS-C-W1 (Unit VII).
Material: Fifty-six valves.
Diagnosis: Subquadrate and subovate, comma-shaped crests, S2 short.
Dimensions: Holotype: L = 0.980 mm, H = 0.600 mm; Paratypes: L = 0.940–1.020 mm, H = 0.560–0.600; other specimens: L = 0.785–1.080 mm, H = 0.446–0.600 mm.
Description: Outline subquadrate-elongate in lateral view, right valve larger than left valve; hinge line straight; cardinal angle obtuse; median node (L2) prominent; adductorial sulcus (S2) prominent as narrow and short; conspicuous marginal frill; comma-shaped crest on valve surface; marginal ridge present; surface smooth.
Discussion: Beyrichiopsis hushootensis sp. nov. may be compared with Beyrichiopsis glyptopleuroides Green 1963 in Becker et al. 1974. The new species differs in having a comma-shaped lateral crest and smooth ornamentation.
Order Podocopida Sars 1866
Superfamily Bairdioidea (=Bairdiacea Sars 1887)
Family Bairdiidae Sars, 1888
Bohlenatia sp. (Fig. 7/c)
Material: One valve
Dimension: L = 0.600 mm, H = 0.188 mm
Occurrence: It was found in the gracilis gracilis Biozone of the section Hushoot Shiveetiin gol section.
Pustulobairdia sp. (Fig. 7/d)
Material: One valve
Dimension: L = 0.600 mm, H = 0.300 mm.
Occurrence: It was observed in the crepida Biozone of Hushoot Shiveetiin gol section.
Superfamily Bairdiocypridoidea (=Bairdiocypridacea Shaver 1961)
Family Bairdiocyprididae Shaver 1961
Baschkirina sp. (Fig. 7/e)
Material: One carapace
Dimension: L = 0.616, H = 0.350 mm
Occurrence: It was found in the rugosa trachytera Biozone of the Hushoot Shiveetiin gol section.
Bairdiocypris sp. (Fig. 7/f-g)
Material: Five carapaces
Dimension: L = 0.925–0.616 mm, H = 0.550–0.350 mm
Occurrence: It occurs from the crepida to gracilis gracilis biozones of Hushoot Shiveetiin gol section.
Superfamily Healdioidea (=Healdiacea Harlton 1933)
Family Pachydomellidae Berdan and Sohn, 1961
Genus Ampuloides Polenova 1952
Type-species: Ampuloides verrucosa Polenova 1952
Ampuloides beckeri sp. nov. (Fig. 7/k-o)
Holotype: Left valve (Fig. 7/l, MUST-RCSP-Ost.00031)
Paratype: Left valve of Paratype A (Fig./k, MUST-RCSP-Ost.00069), left valve lateral Paratype B (Fig. 7/m, MUST-RCSP-Ost.00035), left valve of Paratype C (Fig. 7/n, MUST-RCSP-Ost.00036)
Derivation of name: Dedicated to German ostracodologist Prof. Dr. Gerhard Becker.
Type locality and horizon: The rhomboidea Biozone of the Hushoot Shiveetiin gol section, sample HS-C-2 (Unit IV).
Material: Twenty-one valves.
Diagnosis: Carapace very swollen, inflated in dorsal view, a narrow groove, cardinal extremity, carapace surface warty (small tubercles).
Dimensions: Holotype: L = 0.690 mm, H = 0.430 mm; Paratypes: L = 0.527–0.700 mm, H = 0.310–0.377.
Description: The lateral outline of the carapace is preplete, valves strongly asymmetrical and inflated, a narrow groove runs parallel to the front edge in the lateral view of valve, the most swollen part of the carapace located in the posterior half, left valve overlaps the right one, cardinal extremity in anterodorsal part of left valve, depressing straight hinge margin, the bow-shaped projection occurs in the ventral margin of the left valve, the surface is covered with small rounded tubercles (warty). The tubercles are only visible in well-preserved specimens.
Discussion. Ampuloides beckeri sp. nov. differs from Ampuloides pumilus Olempska, 1979 (Famennian of the Holy Cross Mountains, Poland) by the absence of a nodular process occuring in the anterodorsal part and shape/outline of the carapace. Also, Ampuloides beckeri sp. nov. differs from Ampuloides kellerwaldensis Casier and Lethiers, 1999 (Schmidt Quarry, Frasnian/Famennian section in the Kellerwald, Germany) by the presence of an anterior furrow and by the smaller development of cardinal extremities.
Palaeoenvironmental analysis
Ostracods can provide critical data for reconstructing Palaeozoic environments, especially in the Devonian. Ostracods can be used to distinguish ecologically three mega-assemblages (ecotype), such as the Eifelian (thick-shelled and sculptured ostracod assemblages), Thuringian (thin-shelled species with smooth and spinose valves) and Myodocopid (=Entomozoacean) mega-assemblages (ecotype) (finger-print ostracods) (e.g. Bandel and Becker 1975; Groos-Uffenorde et al. 2000; Becker 2000; Becker et al. 2004; Casier 2004, 2017; Crasquin and Horne 2018). Also, in-depth overall information and references are given and discussed on palaeoecology of ostracods by Casier (2017). Three mega-assemblages and six assemblages of ostracods numbered 0 to V indicative for lagoonal-, semi-restricted- and marine environments above and below fair weather wave-base or storm wave-base in the Devonian are described by Casier (2017). Wang (1988) proposed five ostracod associations (leperditiid, palaeocopid, smooth-podocopid, spinose-podocopid and entomozoacean) ranging from nearshore to deep basins in Palaeozoic strata of South China.
The Devonian ostracod genera and species from the Hushoot Shiveetiin gol section (Baruunhuurai Terrane) are characterized by palaecopids, platycopids and podocopids (Fig. 9). Calcareous levels (Unit II, IV, V and VII) from the section yielded many ostracod specimens even if they lived in a high physical stress environment characterized by frequent and often voluminous pyroclastic eruptions (Ariuntogos et al. 2020, this issue). However, some genera and species are particularly useful for palaeoenvironmental interpretation. Critical assemblages in this study are characterized by the smooth-shelled and ornamented-shell (reticulate, spinous and node-bearing) ostracods.
Hollinoideans lived in marine environments and lived worldwide in nektobentonic, epineritic and pelagic facies realms. They occur together with kloedenellids and eridostracans (Becker 1980, 1982, 2002) and are indicative of more restricted environments. Hypotetragona occurs in marine near-shore environments (Bless 1983). Also, weakly ornamented ostracods such as the Knoxiella-Beyrichiopsis assemblage represent a supratidal environment in the late Famennian in the Dinantian Basin of Belgium (Becker et al. 1974). Primitiopsoidean occur in epineritic as well as in pelagic facies realms (Becker 2002; Adamczak 2005). Kirkbyoidean were marine ostracods ranging from near-shore (e.g. found in shallow-water sections in the Eifel area, Germany) to deep-sea (such as in Thuringia, Germany; see Bandel and Becker 1975; Olempska 1999). The studies on fossil and modern bairdiids indicate that they occur from near-shore to bathyal environments of normal marine salinity (Melnyk and Maddocks 1988). Thin-shelled, spinose bairdiids indicating Thuringian mega-assemblage are absent in the samples from the Hushoot Shiveetiin gol section.
Common short spiny podocopid (Pustulobairdia, Ampuloides) association with kirkbyoid (Amphissites) forms are found in Unit II (sample HS-C-11a, Figs. 2 and 3). The thick-shelled and sculptured palaeocopids (Hollinella, Pribylites, Nesamyslia and Amphissites) and smooth to weak ornamented podocopids (Bairdia, Ampuloides and Cribroconcha) are very common in the upper part of Unit IV (sample number HS-C-2, Figs. 2, 3, 9). Paleocopids are dominant in the other parts of Unit IV. Podocopids are also observed in the upper part of Unit IV (sample number HS-C-W15). Relatively increases in the number of ostracod genera and species point to fully marine conditions which correspond with limestone or slightly silicified limestone. But, the ostracods are represented by a few species and genera in the upper parts of Unit IV (sample number HS-C-W14) and Unit V (sample number HS-C-W12). A very diverse assemblage containing palaecopid + platycopid + podocopid ostracods were observed at the end of Unit VII (sample HS-C-W1, Figs. 2, 3, 9). This assemblage was found in short-lived pure limestone and might be related to a sea-level rise. However, the Hushoot Shiveetin gol section is characterized by a very variable facies setting ranging from lagoonal, shallow-intertidal to open marine. Hydrodynamic conditions are also very variable ranging from high-energy environment to semi-restricted conditions and a number of remarkable sea-level changes (Ariuntogos et al. 2020, this issue).
Overall, the ostracod assemblage of the Hushoot Shiveetiin gol section is considered to be ecologically equivalent to the Eifelian mega-assemblage which include semi-restricted (Ass. I), marine above (Ass. II) or below (Ass. III) fair-weather wave-base or storm wave-base environments. These assemblages correspond to foreshore Ostracod Assemblage-1 (OA1), nearshore Ostracod Assemblage-2 (OA2) and offshore Ostracod Assemblage-23 (OA3) environments of the Hongguleleng Formation (western Junggar, NW China) in an open oceanic island arc (Song et al. 2019). Also, these results can be correlated with the Eifelian mega-assemblage indicative of a nearshore-offshore setting (palaeocopid and smooth-podocopid associations) in the Gelaohe Formation from the Devonian-Carboniferous (D/C) transition, South China (Song and Gong 2019).
The Eifelian mega-assemblage is characterized by variable environments which are confirmed by sedimentological criteria and facies analysis (Ariuntogos et al. 2020, this issue).
Palaeobiogeographical relationships
The Hushoot Shiveetiin gol section (Baruunhuurai Terrane) belongs to an active island arc complex in the CAOB. Ostracod assemblages of this section consist of cosmopolitan, regional and endemic, new species. Some of the identified ostracods are similar to coeval faunas reported from studies in Russia, China, Poland, Belgium, Germany, Spain, Czech and Turkey and references are given below (Fig. 10).
Hollinella (Keslingella) aff. praecursor occurs in the rhomboidea Biozone of the eastern and western parts of the Hushoot Shiveetiin gol section. This species is known from the Late Devonian in Spain (Iberian Chains, Gozalo 1994), from the Middle Devonian from Red Coral Limestone in the Czech Republic (Late Givetian, Pokorny 1950), the Middle Devonian from the Eifel area, Germany (Ahrdorf Beds, Eifelian, Bless and Jordan 1971), from Late Devonian in Belgium (Frasnian, Dinant Basin, Casier and Olempska 2008a), and from the Late Devonian from Turkey (Famennian in Taurids, E. Şeker Zor, personal communication, October 14, 2019).
Hollinella (Keslingella) lionica occurs in the gracilis gracilis and rugosa trachytera biozones of the western part. It is known from the Late Devonian in France (Frasnian of Ardennes and Boulonnais, Lethiers 1984), the Dinant Basin in Belgium (Becker 1971; Casier and Olempska 2008a, b), and the Late Devonian in Turkey (Famennian in the Taurids; E. Şeker Zor, personal communication, October 14, 2019).
Amphissites irinae occurs in both, the eastern and western sections within crepida and rhomboidea biozones. This species is known from the Late Devonian (Frasnian) of Russia (Egorov 1953; Zaspelova 1959; Evdokimova 2015; Zhuravlev et al. 2006), Poland (Zbikowska 1983), Germany (Harz Mountains, Blumenstengel 1969), and Spain (Famennian of Cantabrian Mountains, Becker 1977).
Amphissites centrocostatus occurs from the rhomboidea to rugosa trachytera biozones of the western section. This species is known from the Late Devonian in Germany (Harz Mountains, Frasnian, Blumenstengel 1969), Spain (Famennian, Central Pyrenees; Sanchez de Posada et al. 2008), and from the Pontids in Turkey (Nazik et al. 2012).
Hypotetragona tremula is known from the Late Devonian (Frasnian) of Dinant Basin, Belgium (Becker 1971; Becker and Bless 1974 in Becker et al. 1974; Casier and Olempska 2008a, b).
Bairdia hypsela occurs from the crepida to the gracilis gracilis biozones of both sections. It is known from Upper Devonian-Carboniferous strata from the Holy Cross Mountains, Poland, from the Dinant Basin in France and Belgium, as well as from Germany (Rhenish Massif and Thuringia), from Turkey (Taurids and Pontids), and from Morocco and northern Algeria (Olempska 1979; Becker et al. 1993; Casier et al. 2005; Nazik et al. 2012).
Three new species (Microcheilinella bulongourensis, Pribylites junggarensis and Cribroconcha honggulelengensis) which were identified in the Late Devonian of western Junggar, NW China by Song et al. (2017) were also found in Hushoot Shiveetiin gol section. Cribroconcha primaris occurs in the rhomboidea Biozone of the eastern section. It is known from Tournaisian from the Southern Urals (Kotschetkova 1980), the Devonian/Carboniferous boundary beds of Kozhim, Polar Urals (Nemirovskaya et al. 1992) and of South Urals (Kulagina 2013).
Furthermore, the new species (Beyrichiopsis hushootensis, Ampuloides beckeri) and species described in open nomenclature (Palaeocopina? indet. 1 and 2, Nezamyslia sp., Polytylites sp., Parabolbina sp., Coryellina sp., Berdanella sp., Pustulobairdia sp., Aechmina sp., Bairdiocypris sp., Baschkirina sp., Bohlenatia sp. and Kloedenellidae indet.) are considered as endemic species. These data may support the hypothesis that the CAOB was a diversity hotspot during the Famennian also for ostracods beside other organisms (see Waters et al. 2020, this issue). As a result, the studied ostracod fauna consist of Euramerican/Laurusian assemblages (Hollinella (Keslingella) lionica, Hollinella (Keslingella) aff. praecursor, Amphissites irinae, Amphissites centrocostatus, Hypotetragona tremula and Bairdia hypsela), China (Microcheilinella bulongourensis, Pribylites junggarensis and Cribroconcha honggulelengensis) and endemics.
Conclusions
In total, 25 ostracod species belonging to 19 genera from the Late Devonian (Famennian) Hushoot Shiveetiin gol section (Mongolia) are documented and figured. They were found in sediments ranging from the crepida to rugosa trachytera biozones. The Mongolian ostracod fauna is similar to coeval faunas known from Laurussia but also from China. Two new taxa are described. Ostracod assemblages are ecologically equivalent to the Eifelian mega-assemblage (I-III) and generally represent nearshore, variable palaeoenvironment. The diverse ostracod Eifelian mega-assemblage expanding from shallow-water refugia supports the idea of the CAOB as a biodiversity hotspot in the aftermath of the F/F extinction.
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Acknowledgments
The authors are grateful to Jana Anger (Senckenberg Institute, Frankfurt) for conodont sample preparations and selections of ostracod specimens under the binocular microscope. Also, A. Nazik would like to thank Peter Königshof (Senckenberg Institute, Frankfurt) inviting me for studying ostracod findings from Mongolia. We appreciate the Çukurova Üniversity, Central Laboratory Management and Expert Derya Akalan taking of scanning electron microscope photographs. The PhD student (A.M.) and her supervisor (P.K.) acknowledge funding from both, German Academic Exchange Service (DAAD, Research Grant–Doctoral Programme in Germany, September 01, 2018–September 01, 2021; 57381412) and German Science Foundation (DFG, KO 1622-19/1). This paper is a contribution to IGCP Project 652 (Reading geologic time in Palaeozoic rocks: the need for an integrated stratigraphy [2017–2021]). We are grateful to Junjun Song (China) and Marie-Béatrice Forel (France) for their critical comments and suggestions on the manuscript.
Funding
This study was supported by the German Academic Exchange Service (DAAD, Research Grant–Doctoral Programme in Germany, September 01, 2018–September 01, 2021; 57381412) and German Science Foundation (DFG, KO 1622-19/1).
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Appendix
Appendix
Taxonomic list of identified ostracod taxa from Hushoot Shiveetiin gol section, Mongolia
Class Ostracoda Latreille, 1802
Order Palaeocopida Henningsmoen, 1953
Superfamily Hollinoidea (=Hollinacea) Swartz 1936
Family Hollinellidae Bless and Jordán 1972
Hollinella (Keslingella) lionica Becker and Bless, 1971 (Fig. 4/a-d)
Hollinella (Keslingella) aff. praecursor Pokorny, 1950 (Fig. 4/e-g)
Superfamily Aechminoidea Bouček, 1936
Family Aechminidae Bouček, 1936
Aechmina sp. (Fig. 4/i)
Family Ctenoloculinidae Jaanusson and Martinsson, 1956
Parabolbina sp. (Fig. 4/j)
Superfamily Primitiopsoidea (=Primitiopsacea) Swartz 1936
Family Pribylitidae Pokorný, 1958
Pribylites junggarensis Song and Crasquin, 2017 in Song et al. 2017 (Fig. 4/k-n)
Coryellina sp. (Fig. 4/o)
Superfamily Kirkbyoidea (=Kirkbyacea Ulrich and Bassler 1906)
Family Amphissitidae Knight, 1928
Amphissites centrocostatus Blumenstengel, 1969 (Fig. 5/a)
Amphissites irinae Glebovskaja and Zaspelova in Egorov, 1953 (Fig. 5/b-e)
Polytylites sp. (Fig. 5/f)
Family Nezamysliidae Żbikowska, 1983
Nezamyslia sp. (Fig. 5/g–i)
Family Kirkbyellidae Sohn, 1961
Berdanella sp. (Fig. 5/j–k)
Palaeocopina? indet. 1 (Fig. 5/l–n)
Palaeocopina? indet. 2 (Fig. 5/o, Fig. 6/a–f)
Order Platycopida Sars 1866
Superfamily Kloedenelloidea (=Kloedenellacea Ulrich and Bassler 1908)
Family Kloedenellidae
Kloedenellidae indet. (Fig. 6/g–h)
Family Geisinidae Sohn, 1961
Hypotetragona tremula Becker, 1971 (Fig. 6/i)
Family Beyrichiopsidae Henningsmoen, 1953
Beyrichiopsis hushootensis sp. nov. (Fig. 6/j–o, Fig. 7/a)
Order Podocopida Sars 1866
Superfamily Bairdioidea (=Bairdiacea Sars 1887)
Family Bairdiidae Sars, 1888
Bairdia hypsela Rome 1971 (Fig. 7/b)
Bohlenatia sp. (Fig. 7/c)
Pustulobairdia sp. (Fig. 7/d)
Superfamily Bairdiocypridoidea (=Bairdiocypridacea Shaver, 1961)
Family Bairdiocyprididae Shaver, 1961
Baschkirina sp. (Fig. 7/e)
Bairdiocypris sp. (Fig. 7/f–g)
Family Pachydomellidae Berdan and Sohn 1961
Microcheilinella bulongourensis Song and Crasquin, 2017 (Fig. 7/h–j)
Ampuloides beckeri sp. nov. (Fig. 7/k–o)
Superfamily Healdioidea (=Healdiacea Harlton 1933)
Family Healdiidae
Cribroconcha primaris Kotschetkova, 1980 (Fig. 8/a–c)
Cribroconcha honggulelengensis Song and Crasquin, 2017 (Fig. 8/d–f)
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Nazik, A., Königshof, P., Ariuntogos, M. et al. Late Devonian ostracods (Crustacea) from the Hushoot Shiveetiin gol section (Baruunhuurai Terrane, Mongolia) and their palaeoenvironmental implication and palaeobiogeographic relationship. Palaeobio Palaeoenv 101, 689–706 (2021). https://doi.org/10.1007/s12549-020-00446-z
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DOI: https://doi.org/10.1007/s12549-020-00446-z