A Mid-Ordovician Brachiopod Evolutionary Hotspot in Southern Kazakhstan

Оглавление

Leonid E. Popov. A Mid-Ordovician Brachiopod Evolutionary Hotspot in Southern Kazakhstan

Table of Contents

List of Tables

List of Illustrations

Guide

Pages

A mid-Ordovician brachiopod evolutionary hotspot in southern Kazakhstan

A mid-Ordovician brachiopod evolutionary hotspot in southern Kazakhstan

Introduction

Geological setting and key sections

South Betpak‐Dala

Area 6 km south‐west of Baigara Mountain

Area on the west side of the Karatal River

North Betpak‐Dala

Tastau Formation (New) Derivation of name

Stratotype

Definition of boundaries

Description

Takyrsu Formation (New) Derivation of name

Stratotype

Definition of boundaries

Description

Distribution

Age

The West Balkhash Region

The area south‐west of Lake Alakol

Berkutsyur Formation (New) Derivation of name

Stratotype

Definition of boundaries

Description

Distribution

Age

Remarks

Kopkurgan Formation (New) Derivation of name

Stratotype

Definition of boundaries

Description

Distribution

Age

Brachiopod associations

The Ectenoglossa Association

The Ancistrorhyncha Association

The Baitalorhyncha Association

The Tesikella Association

The Sowerbyella-Altynorthis Association

The Altynorthis‐Lepidomena Association

The Testaprica Association

The Acculina‐Sowerbyella Association

The Eoanastrophia Association

The Chaganella Association

The Bimuria-Grammoplecia Association

Deep water brachiopod associations

The Sowerbyella-Doughlatomena Association

Implications for biodiversity

Localities

North Betpak‐Dala. Locality 154

Locality 162

Locality 163

Locality 166

South Betpak‐Dala. Locality 625/1

Locality 625/3

Locality 765-e

Area 10 km south‐west of Lake Alakol. Locality 735

Area 8 km south‐west of Lake Alakol. Locality 727

Area 7 km south‐west of Lake Alakol. Locality 8120

Locality 8120/4

Locality 8121

Locality 8122

Locality 8124

Locality 8125

Locality 8233

Locality 8235

Locality 8236a

Locality 8236

Area 5.5 km south‐west of Lake Alakol. Locality N-12

Area 4 km south‐west of Lake Alakol. Locality 812

Locality 813

Locality 814

Locality 815

Locality 816

Locality 817

Locality 817a

Locality 8234

Area 6 km south‐west of Baigara Mountain. Locality 1020

Locality 1021

Locality 1022

Locality 1023

West side of the River Karatal, Baigara. Locality 1025

Locality 1026

Locality 1026a

Locality 1026b

Locality 1028

Other areas. Locality 388

Locality 388a

Locality 538

Locality 1501

Locality 8126

Locality 11221

Systematic palaeontology. Remarks

Repositories

Class Lingulata Goryanski & Popov, 1985. Order Lingulida Waagen, 1885. Superfamily Linguloidea Menke, 1828. Family Obolidae King, 1846. Subfamily Obolinae King, 1846. Genus Aploobolus n. gen., n. sp. Derivation of name

Type species and only species

Diagnosis

Remarks

Aploobolus tenuis n. sp

Derivation of name

Holotype

Paratypes

Diagnosis

Description

Subfamily Glossellinae Cooper, 1956. Genus Ectenoglossa Sinclair, 1945. Type species (by original designation)

Ectenoglossa sorbulakensis Popov, 1980b

Holotype

Material

Remarks

Genus Meristopacha Sutton in Sutton et al., 1999. Type species (by original designation)

Remarks

Meristopacha sp

Material

Description

Remarks

Subfamily Elliptoglossinae Popov & Holmer, 1994. Genus Elliptoglossa Cooper, 1956. Type species (by original designation)

Elliptoglossamagna Popov, 1977

Holotype

Material

Remarks

Family Elkaniidae Walcott & Schuchert in Walcott, 1908. Genus Broeggeria Walcott, 1902. Type species (by original designation)

Broeggeria cf. B. putilla (Tenjakova, 1989)

Material

Remarks

Family uncertain. Genus Oxlosia Ulrich & Cooper, 1936. Type species (by original designation)

Oxlosia? sp

Remarks

Superfamily Discinoidea Gray, 1840. Family Trematidae Schuchert, 1893. Genus Trematis Sharpe, 1848. Type species (by subsequent designation of Davidson (1853, p. 130)

Trematis aff. T. parva Cooper, 1956 Plate 2, figure 6

Material

Remarks

Trematis? sp

Material

Remarks

Order Siphonotretida Kuhn, 1949. Superfamily Siphonotretoidea Kutorga, 1848. Family Siphonotretidae Kutorga, 1848. Genus Multispinula Rowell, 1962. Type species (by original designation)

Multispinula sp

Material

Remarks

Class Craniata Williams, Brunton, Carlson, Holmer & Popov, 1996. Order Craniida Waagen, 1885. Superfamily Cranioidea Menke, 1828. Family Craniidae Menke, 1828. Genus Petrocrania Raymond, 1911. Type species (by original designation)

Petrocrania? sp

Material

Remarks

Genus Pseudocrania M'Coy, 1851. Type species (by subsequent designation of Davidson (1853) explanation of pl. 91)

Pseudocraniakaratalensis Popov in Nazarov & Popov, 1980 (Not figured)

Holotype

Material

Remarks

Class Chileata Williams, Brunton, Carlson, Holmer & Popov, 1996. Order Dictyonellida Cooper, 1956. Superfamily Eichwaldioidea Schuchert, 1893. Family Eichwaldiidae Schuchert, 1893. Genus Eichwaldia Billings, 1858. Type species (by original designation)

Eichwaldia sp

Material

Remarks

Class Strophomenata Williams, Brunton, Carlson, Holmer & Popov, 1996 Order Strophomenida Öpik, 1934. Superfamily Strophomenoidea King, 1846. Family Strophomenidae King, 1846. Subfamily Strophomeninae King, 1846. Genus Esilia Nikitin & Popov, 1985. Type species (by original designation)

Esilia cf. E. tchetverikovae Nikitin & Popov, 1985

Material

Remarks

Genus Infurca Percival, 1979. Type species (by original designation)

Infurca? sp

Material

Remarks

Subfamily Furcitellinae Williams, 1965. Genus Katastrophomena Cocks, 1968. Type species (by original designation)

Remarks

Katastrophomenarukavishnikovae (Nikitina, 1985)

Holotype

Material

Description

Remarks

Katastrophomena sp

Remarks

Furcitellinae gen. et sp. indet

Material

Remarks

Family Rafinesquinidae Schuchert, 1893. Subfamily Rafinesquininae Schuchert, 1893. Genus Rhipidomena Cooper, 1956. Type species (by original designation)

Rhipidomena sp

Remarks

Genus Testaprica Percival, 2009. Type species (by original designation)

Testaprica alperovichi n. sp

Derivation of name

Holotype

Material

Diagnosis

Description

Remarks

Genus Colaptomena Cooper, 1956. Type species (by original designation)

Colaptomena insolita (Nikitina, 1985)

Holotype

Material

Description

Remarks

Subfamily Leptaeninae Hall & Clarke, 1894. Genus Leptaena Dalman, 1828. Type species (by subsequent designation of King 1846, p. 28)

Subgenus Leptaena (Ygdrasilomena) Cocks, 2005. Type species (by original designation)

Leptaena (Ygdrasilomena) sp. Figure 17F. Material

Remarks

Genus Doughlatomena n. gen. Type species

Derivation of name

Diagnosis

Discussion

Doughlatomena splendens n. sp

Derivation of name

Holotype

Diagnosis

Material and dimensions

Description

Remarks

Family Glyptomenidae Cooper, 1956. Remarks

Genus Buminomena Popov & Cocks, 2014. Type species (by original designation)

Buminomena sp

Material

Remarks

Glyptomenidae gen. et sp. indet

Remarks

Family Christianiidae Williams, 1953. Genus Christiania Hall & Clarke, 1892. Type species (by original designation)

Remarks

Christiania cf. C. proclivis Popov & Cocks, 2006

Material

Remarks

Christiania cf. C. tortuosa Popov, 1980b

Holotype

Material

Description

Remarks

Superfamily Plectambonitoidea Jones, 1928. Family Plectambonitidae, Jones, 1928. Subfamily Taphrodontinae Cooper, 1956. Genus Bandaleta Nikitin & Popov, 1996. Type species (by original designation)

Bandaleta sp

Material

Remarks

Genus Isophragma Cooper, 1956. Type species (by original designation)

Isophragma princeps Popov, 1980b

Holotype

Material

Description

Remarks

Family Bimuriidae Cooper, 1956. Genus Bimuria Ulrich & Cooper, 1942. Type species (by original designation)

Bimuria karatalensis n. sp

Derivation of name

Holotype

Material

Diagnosis

Description

Remarks

Bimuria sp

Material

Remarks

Genus Cooperea Cocks & Rong, 1989. Type species (by original designation)

Cooperea sp

Material

Remarks

Family Leptellinidae Ulrich & Cooper, 1936. Subfamily Leptellininae Ulrich & Cooper, 1936. Genus Acculina Misius in Misius & Ushatinskaya, 1977. Type species (by original designation)

Acculina acculica Misius in Misius & Ushatinskaya, 1977

Type specimens

Material

Remarks

Acculina? sp

Material

Remarks

Genus Dulankarella Rukavishnikova, 1956. Type species (by original designation)

Dulankarella larga Popov, Cocks & Nikitin, 2002

Holotype

Material

Remarks

Genus Kajnaria Nikitin & Popov in Klenina, Nikitin & Popov, 1984. Type species (by original designation)

Kajnaria derupta Nikitin & Popov in Klenina, Nikitin & Popov, 1984

Holotype

Material

Remarks

Genus Leptellina Ulrich & Cooper, 1936. Type species (by original designation)

Leptellina sp

Material

Description

Remarks

Genus Mabella Klenina in Klenina, Nikitin & Popov, 1984. Type species (by original designation)

Mabella sp

Material

Remarks

Subfamily Palaeostrophomeninae Cocks & Rong, 1989. Genus Apatomorpha Cooper, 1956. Type species (by original designation)

Apatomorpha akbakaiensis n. sp

Derivation of name

Holotype

Paratypes

Diagnosis

Description

Remarks

Genus Ishimia Nikitin, 1974. Type species (by original designation)

Ishimia aff. I. ishimensis Nikitin, 1974

Material

Remarks

Genus Lepidomena Laurie, 1991. Type species (by original designation)

Lepidomena betpakdalensis n. sp

Derivation of name

Holotype

Material

Diagnosis

Description

Remarks

Genus Tesikella Popov, Cocks & Nikitin, 2002. Type species (by original designation)

Tesikella necopina (Popov, 1980b)

Holotype

Material

Remarks

Genus Titanambonites Cooper, 1956. Type species (by original designation)

Titanambonites cf. T. magnus Nikitin, 1974

Material

Remarks

Genus Shlyginia Nikitin & Popov, 1983. Type species (by original designation)

Remarks

Shlyginia extraordinaria (Rukavishnikova, 1956)

Holotype

Material

Remarks

Shlyginia sp

Material

Remarks

Family Leptestiidae Öpik, 1933. Genus Leangella (Leangella) Öpik, 1933. Type species (by original designation)

Leangella (Leangella) sp

Material

Remarks

Family Xenambonitidae Cooper, 1956. Subfamily Aegiromeninae Havlíc\̌ek, 1961. Genus Akadyria Nikitina, Neuman, Popov & Bassett in Nikitina et al., 2006. Type species (by original designation)

Akadyria simplex Nikitina, Neuman, Popov & Bassett in Nikitina et al., 2006

Holotype

Material

Remarks

Genus Chonetoidea Jones, 1928. Type species (by original designation)

Chonetoidea? sp

Material

Remarks

Genus Kassinella Borissiak, 1956. Type species (by original designation)

Remarks

Kassinella simorini n. sp

Derivation of name

Holotype

Paratypes

Diagnosis

Description

Remarks

Genus Synambonites Zhan & Rong, 1995. Type species (by original designation)

Synambonites sp

Remarks

Genus Tenuimena Nikitina, Neuman, Popov & Bassett in Nikitina et al. 2006. Type species (by original designation)

Tenuimena aff. T. planissima Nikitina, Neuman, Popov & Bassett in Nikitina et al. 2006

Material

Remarks

Family Hesperomenidae Cooper, 1956. Genus Anoptambonites Williams, 1962. Type species (by original designation)

Remarks

Anoptambonites kovalevskii Popov, Nikitin & Cocks, 2000

Holotype

Material

Remarks

Anoptambonites sp. 1

Remarks

Anoptambonites sp. 2

Material

Remarks

Genus Chaganella Nikitin, 1974. Type species (by original designation)

Chaganella chaganensis Nikitin, 1974

Material

Remarks

Family Sowerbyellidae Öpik, 1930. Subfamily Sowerbyellinae Öpik, 1930. Genus Sowerbyella (Sowerbyella) Jones, 1928. Type species (by original designation)

Remarks

Sowerbyella (Sowerbyella) ampla (Nikitin & Popov, 1996)

Holotype

Alakol and North Betpak‐Dala Material

Remarks

Sowerbyella (Sowerbyella) verecunda verecunda Nikitin & Popov, 1983

Holotype

Sowerbyella (Sowerbyella) verecunda baigarensis n. subsp

Derivation of name

Holotype

Paratypes

Other material

Diagnosis

Description

Remarks

Sowerbyella (Sowerbyella) cf. S. (S.) acculica Misius, 1986

Material

Remarks

Genus Gunningblandella Percival, 1979. Type species (by original designation)

Gunningblandella sp. 1

Material

Remarks

Gunningblandella sp. 2

Remarks

Genus Ptychoglyptus Willard, 1928. Type species (by original designation)

Ptychoglyptus sp

Material

Remarks

Order Orthotetida Waagen, 1884. Suborder Orthotetidina Waagen, 1884. Superfamily Chilidiopsoidea Boucot, 1959. Family Chilidiopsidae Boucot, 1959. Subfamily Chilidiopsinae Boucot, 1959. Genus Gacella Williams, 1962. Type species (by original designation)

Gacella sp

Material

Remarks

Suborder Triplesiidina Moore, 1952. Superfamily Triplesioidea Schuchert, 1913. Family Triplesiidae Schuchert, 1913. Genus Grammoplecia Wright & Jaanusson, 1993. Type species (by original designation)

Grammoplecia globosa (Nikitin & Popov, 1985)

Holotype

Material

Remarks

Grammoplecia aff. G. globosa (Nikitin & Popov, 1985)

Material

Remarks

Grammoplecia sp

Material

Remarks

Genus Triplesia Hall, 1859. Type species (by original designation)

Triplesia sp

Remarks

Order Billingsellida Schuchert, 1893. Superfamily Polytoechioidea Opik, 1934. Family Tritoechiidae Ulrich & Cooper, 1936. Genus Eremotoechia Cooper, 1956. Type species (by original designation)

Eremotoechia inchoata Popov, Vinn & Nikitina, 2001

Holotype

Remarks

Eremotoechia spissa Popov, Vinn & Nikitina, 2001

Holotype

Remarks

Order Protorthida Schuchert & Cooper, 1931. Suborder Clitambonitidina Öpik, 1934. Remarks

Superfamily Clitambonitoidea Winchell & Schuchert, 1893. Family Clitambonitidae Winchell & Schuchert, 1893. Genus Atelelasma Cooper, 1956. Type species (by original designation)

Atelelasma sp

Material

Description

Order Orthida Schuchert & Cooper, 1932. Suborder Orthidina Schuchert & Cooper, 1932. Superfamily Orthoidea Woodward, 1852. Family Hesperorthidae Schuchert & Cooper, 1931. Subfamily Dolerorthinae Öpik, 1934. Genus Dolerorthis Schuchert & Cooper, 1931. Type species (by original designation)

Dolerorthis expressa Popov, 1980b

Holotype

Remarks

Genus Sonculina Misius, 1986. Type species (by original designation)

Diagnosis

Remarks

Sonculina cf. S. prima Misius, 1986

Material

Description

Remarks

Sonculina baigarensis n. sp

Derivation of name

Holotype

Paratypes

Diagnosis

Description

Remarks

Hesperorthidae gen. et sp. indet

Material

Remarks

Family Glyptorthidae Schuchert & Cooper, 1931. Genus Eridorthis Foerste, 1909. Type species (by original designation)

Remarks

Eridorthis sp

Material

Description

Remarks

Family Plaesiomyidae Schuchert & Cooper, 1931. Genus Bokotorthis Popov, Nikitin & Cocks, 2000. Type species (by original designation)

Bokotorthis kasachstanica (Rukavishnikova, 1956)

Holotype

Material

Remarks

Superfamily Plectorthoidea Schuchert in Schuchert & LeVene, 1929. Family Plectorthidae Schuchert in Schuchert & LeVene, 1929. Genus Altynorthis n. gen. Derivation of name

Type species

Diagnosis

Species assigned

Species questionably assigned

Remarks

Altynorthis betpakdalensis n. sp

Holotype

Material

Diagnosis

Description

Remarks

Altynorthis tabylgatensis (Misius, 1986)

Holotype

Material

Remarks

Altynorthis vinogradovae n. sp

Derivation of name

Holotype

Paratypes

Diagnosis

Description

Remarks

Genus Lictorthis n. gen. Derivation of the name

Type and only species

Diagnosis

Remarks

Lictorthis cf. L. licta (Popov & Cocks, 2006)

Remarks

Family Giraldiellidae Williams & Harper, 2000. Genus Phaceloorthis Percival, 1991. Type species (by original designation)

Phaceloorthis? corrugata n. sp

Derivation of name

Holotype

Topotypes

Diagnosis

Description

Remarks

Genus Scaphorthis Cooper, 1956. Type species (by original designation)

Scaphorthis recurva Nikitina, 1985

Holotype

Material

Remarks

Suborder Dalmanellidina Moore, 1952. Superfamily Dalmanelloidea Schuchert, 1913. Family Dalmanellidae Schuchert, 1913. Subfamily Dalmanellinae Schuchert, 1913. Genus Onniella Bancroft, 1928. Type species, (by original designation)

Onniella? sp

Material

Remarks

Superfamily Enteletoidea Waagen, 1884. Family Draboviidae Havlíc\̌ek, 1950. Subfamily Draboviinae Havlíc\̌ek, 1950. Genus Pionodema Foerste, 1912. Type species (by original designation)

Pionodema opima Popov, Cocks & Nikitin, 2002

Holotype

Material

Remarks

Order Pentamerida Schuchert & Cooper, 1931. Suborder Syntrophiidina Ulrich & Cooper, 1936. Superfamily Camerelloidea Hall & Clarke, 1894. Family Camerellidae Hall & Clarke, 1894. Genus Ilistrophina Popov, Cocks & Nikitin, 2002. Type species (by original designation)

Ilistrophina tesikensis Popov, Cocks & Nikitin, 2002

Holotype

Material

Remarks

Genus Plectocamara Cooper, 1956. Type species (by original designation)

Plectocamara sp

Material

Description

Remarks

Family Parastrophinidae Schuchert, 1929. Genus Eoanastrophia Nikiforova & Sapelnikov, 1973

Type species (by original designation)

Remarks

Eoanastrophia kurdaica Sapelnikov & Rukavishnikova, 1975

Holotype

Material

Remarks

Genus Parastrophina Nikiforova & Sapelnikov, 1973. Type species (by original designation)

Parastrophina? sp

Remarks

Order Rhynchonellida Kuhn, 1949. Superfamily Ancistrorhynchoidea Cooper, 1956. Family Ancistrorhynchidae Cooper, 1956. Genus Ancistrorhyncha Ulrich & Cooper, 1936. Type species (by original designation)

Ancistrorhyncha modesta Popov in Nikiforova & Popov, 1981

Holotype

Material

Remarks

Genus Lydirhyncha n. gen

Derivation of name

Type species

Diagnosis

Species assigned

Species questionably assigned

Discussion

Lydirhyncha tarimensis (Sproat & Zhan, 2018)

Holotype

Material

Description of the Kazakh material

Remarks

Family Sphenotretidae Savage, 1996. Genus Baitalorhyncha n. gen. Type species

Derivation of name

Diagnosis

Species questionably assigned

Remarks

Baitalorhyncha rectimarginata n. sp

Derivation of name

Holotype

Paratypes

Diagnosis

Description

Remarks

Family Oligorhynchidae Cooper, 1956. Genus Paraoligorhyncha Popov in Nikiforova & Popov, 1981. Type species (by original designation)

Paraoligorhyncha? sp

Material

Remarks

Order Atrypida Rzhonsnitskaya, 1960. Suborder Atrypidina Moore, 1952. Superfamily Atrypoidea Gill, 1871. Family Atrypidae Gill, 1871. Genus Qilianotryma Xu, 1979. Type species (by original designation)

Qilianotryma cf. Q. suspectum (Popov in Nikiforova et al., 1982)

Remarks

Suborder Lissatrypidina Copper, 1996. Superfamily Protozygoidea Copper, 1986. Family Kellerellidae n. fam. Diagnosis

Genera assigned

Remarks

Genus Costistriispira n. gen. Derivation of name

Type and only species

Diagnosis

Remarks

Costistriispira proavia n. sp

Derivation of name

Holotype

Material

Diagnosis

Description

Remarks

Costistriispira? sp

Remarks

Family Cyclospiridae Schuchert, 1913. Genus Rozmanospira Popov, Nikitin & Sokiran, 1999. Type (by original designation) and only species

Remarks

Rozmanospira mica (Nikitin & Popov in Klenina, Nikitin & Popov 1984)

Holotype

Material

Remarks

Acknowledgements

References

PLATES 1 – 21

Appendix

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