Title Page
Contents
ABSTRACT 16
국문요약 18
GENERAL INTRODUCTION 20
Part Ⅰ. Taxonomic study of Bathynellacea from South Korea and Mongolia 22
Ⅰ-1. Introduction 23
Ⅰ-2. Materials and methods 27
Ⅰ-2-1. Study area and groundwater sampling 27
Ⅰ-2-2. Morphological examination 32
Ⅰ-2-3. General morphology of Bathynellacea 32
Ⅰ-2-4. DNA sequencing and molecular analysis 38
Ⅰ-3. Results 46
Ⅰ-3-1. List of species in this study 46
Ⅰ-3-2. Systematic accounts 48
Ⅰ-4. Discussion 174
Ⅰ-4-1. Proposal of a new genus for Eobathynella gwangjuensis and an additional new putative genus 174
Ⅰ-4-2. DNA barcoding and genetic diversity of Korean parabathynellid species 179
Ⅰ-4-3. Progenetic variation in the genus Allobathynella 180
Ⅰ-4-4. Molecular characterization of bathynellid species from South Korea and Mongolia 181
Ⅰ-5. Conclusion 186
Part Ⅱ. Molecular phylogenetic analysis and estimation of the timing of diversification for parabathynellid genera 187
Ⅱ-1. Introduction 188
Ⅱ-2. Materials and methods 190
Ⅱ-2-1. Taxon selection 190
Ⅱ-2-2. Molecular phylogenetic analysis 190
Ⅱ-2-3. Divergence time estimation 190
Ⅱ-3. Results 194
Ⅱ-3-1. Phylogenetic relationships for 21 parabathynellid genera 194
Ⅱ-3-2. Divergence time estimation 196
Ⅱ-4. Discussion 198
Ⅱ-5. Conclusion 202
SUMMARY 203
REFERENCES 205
APPENDIX 220
Table 1. List of sampling sites with GPS, water system, and collected species in South Korea and Mongolia. An asterisk is added in front of the site numbers of the new species type locality. New species discovered in this study marked in bold. (Abbreviation: R.=River) 31
Table 2. Sequences of primer using in this study. 39
Table 3. Primer set and PCR condition for each gene in this study. 39
Table 4. List of species with corresponding GenBank accession numbers and references used in molecular analysis. Species names obtained as part of the present study are preceded by an asterisk.... 40
Table 5. Voucher and GenBank accession number of specimens registered as part of this study with detail information. Site numbers corresponding with Table 1. (Abbreviations:... 42
Table 6. Genetic p-distance of CO1 sequences (435 bp) among Altainella species. 59
Table 7. Genetic p-distance of CO1 sequences (660 bp) among specimens of Allobathynella species. 95
Table 8. Genetic p-distance of 16S sequences (452 bp) among specimens of Allobathynella species. 95
Table 9. Morphological differences among four species of Arisubathynella, cited and modified from Park & Cho (2015) and Ji & Min (2023a). 110
Table 10. Genetic p-distance of CO1 sequences (726 bp) among specimens of Arisubathynella species. 111
Table 11. Morphological differences among seven species of Eobathynella, cited and modified Ji & Min (2022a). 125
Table 12. Genetic p-distance of CO1 sequences (687 bp) among specimens of Eobathynella species. 126
Table 13. Morphological differences among five species of Hangangbathynella, cited and modified from Ji et al. (2021). 140
Table 14. Genetic p-distance of CO1 sequences (579 bp) among Hangangbathynella species. 141
Table 15. Morphological differences of eight species of Nipponbathynella, cited and modified from Ji & Min (2023c). 162
Table 16. Genetic p-distance of CO1 sequences (789 bp) among Nipponbathynella species. 163
Table 17. Morphological differences between East Asian relatives and Parabathynellidae gen. et sp. nov. 168
Table 18. Genetic p-distance based on 18S rDNA sequences (1,761 bp) of Korean Parabathynellidae. 178
Table 19. List of species with corresponding GenBank accession numbers and references used in molecular phylogenetic analysis based on maximum likelihood and Bayesian inference. 192
Table 20. Inferred divergence time of genera in the family Parabathynellidae. (Abbreviation: HPD=highest posterior probability density). 196
Figure 1. Study area in this study. 1. South Korea, 2. Mongolia. 27
Figure 2. Map showing 17 sampling sites in South Korea. Type locality of new species discovered in this study is indicated by a red solid square. The numbers shown in the map... 28
Figure 3. Map showing six sampling sites in Mongolia. Type locality of new species discovered in this study is indicated by a red solid square. The numbers shown in the map correspond to the... 29
Figure 4. Six sampling sites in Mongolia shown in Fig. 3. A, site 18; B, site 19; C, site 20; D, site 22; E, site 21; F, site 23. 30
Figure 5. Habitus of two families of Bathynellacea covered in this study: Bathynellidae (above), Parabathynellidae (below). Scale bar: 0.5 mm. 34
Figure 6. General morphology of Bathynellacea with characters labeled. 35
Figure 7. Appendages of Bathynellacea: antennae and mouthparts. A, AI; B, AII; C, Lbr; D, Md; E, MxI; F, MxII. 36
Figure 8. Appendages of Bathynellacea: thoracopods, uropod and furca. A, Th; B, ThVIII of male; C; Upd; D, Pleotelson and furcal rami. 37
Figure 9. Map showing of collecting sites of Altainella species. A, Distribution of all previously recorded Altainella species (black solid square) and type locality of Altainella sp. nov. (red solid... 52
Figure 10. Altainella sp. nov., male. Scale bar: 0.5 mm. 53
Figure 11. Altainella sp. nov., holotype male. A, AI; B, AII; C, Lbr; D, Pg. Scale bars: 0.05 mm. 54
Figure 12. Altainella sp. nov., holotype male. A, Md; B, MxI; C, MxII; D, ThI; E, ThII; F, ThIII. Scale bars: 0.05 mm. 55
Figure 13. Altainella sp. nov., holotype male. A, ThIV; B, ThV; C, ThVI; D, ThVII. Scale bars: 0.05 mm. 56
Figure 14. Altainella sp. nov., holotype male. A, ThIV; B, ThV; C, ThVI; D, ThVII. Scale bars: 0.05 mm. 57
Figure 15. Altainella sp. nov., allotype female. A, ThIV; B, ThV; C, ThVI; D, ThVII. Scale bars: 0.05 mm. 58
Figure 16. Distribution map of all previously reported Morimotobathynella species (black solid circle) and type locality of new species of Morimotobathynella. The number in the map... 64
Figure 17. Morimotobathynella sp. nov., female. Scale bar: 0.5 mm. 65
Figure 18. Morimotobathynella sp. nov., holotype female. A, AI; B, AII; C, Lbr; D, Pg. Scale bars: 0.05 mm. 66
Figure 19. Morimotobathynella sp. nov., holotype female. A, Md; B, MxI; C, MxII; D, ThI; E, ThII. Scale bars: 0.05 mm. 67
Figure 20. Morimotobathynella sp. nov., holotype female. A, ThIII; B, ThIV; C, ThV; D, ThVI. Scale bar: 0.05 mm. 68
Figure 21. Morimotobathynella sp. nov., (A, B, D) holotype female; (C) allotype male. A, ThVII; B, ThVIII; C, ThVIII; D, Pld. Scale bars: 0.05 mm. 69
Figure 22. Morimotobathynella sp. nov., holotype female. A, Upd; B, Furcal rami. Scale bars: 0.05 mm. 70
Figure 23. Map showing the distribution of all previously recorded Allobathynella species. 1. A. imjinensis, 2. A. hongcheonensis, 3. A. bangokensis, 4. A. gangneungensis, 5. A. wonjuensis, 6.... 81
Figure 24. Map showing the collecting sites of Allobathynella species in this study. The type locality of new species is marked with red square. The numbers in the map correspond with the... 82
Figure 25. Habitat photograph of A. boseongensis sp. nov. 83
Figure 26. Habitat photograph of A. danyangensis sp. nov. 83
Figure 27. Allobathynella cheongdoensis, female. Scale bar: 0.5 cm. 84
Figure 28. Allobathynella yecheonensis, female. Scale bar: 0.5 cm. 84
Figure 29. Allobathynella danyangensis sp. nov., female. Scale bar: 0.5 mm. 85
Figure 30. Allobathynella boseongensis sp. nov., holotype female. A, habitus; B, AI; C, AII; D, Lbr; E, Md; F, MxI. Scale bars: 0.5 mm. 86
Figure 31. Allobathynella boseongensis sp. nov., holotype female. A, MxII; B, ThI; C, ThII; D, ThIII; E, ThIV. Scale bars: 0.5 mm. 87
Figure 32. Allobathynella boseongensis sp. nov., holotype female. A, ThV; B, ThVI; C, ThVII. Scale bar: 0.5 mm. 88
Figure 33. Allobathynella boseongensis sp. nov., (A-D) holotype female; (E) allotype male. Scale bar: 0.5 mm. 89
Figure 34. Allobathynella danyangensis sp. nov., holotype female. A, AI; B, AII; C, Lbr; D, Md (right); E, Md (left). Scale bars: 0.5 mm. 90
Figure 35. Allobathynella danyangensis sp. nov., holotype female. A, MxI; B, MxII; C, ThI; D, ThII; E, ThIII. Scale bars: 0.05 mm. 91
Figure 36. Allobathynella danyangensis sp. nov., holotype female. A, ThIV; B, ThV; C, ThVI; D, ThVII. Scale bar: 0.05 mm. 92
Figure 37. Allobathynella danyangensis sp. nov., holotype female. A, ThVIII; B, Pld; C, Upd; D, telson and furcal rami. Scale bars: 0.05 mm. 93
Figure 38. Allobathynella danyangensis sp. nov., (A-C) paratype males; (D) paratype female. A, ThVIII, ventral view; B, ThVIII, lateral view; C, ThVIII, ventral view; D, Upd. Scale bars: A... 94
Figure 39. Distribution map of all previously reported Arisubathynella species (black solid circle) and type locality of new species of Arisubathynella (red solid square). The numbers in the... 102
Figure 40. Habitat photograph of A. pocheonensis sp. nov. 103
Figure 41. Arisubathynella cheongmiensis, male. Scale bar: 0.5 mm. 104
Figure 42. Arisubathynella hongcheonensis, female. Scale bar: 0.5 mm. 104
Figure 43. Arisubathynella pocheonensis sp. nov., female. Scale bar: 0.05 mm. 104
Figure 44. Arisubathynella pocheonensis sp. nov., holotype male. A, AI; B, AII; C, Lbr; D, Md; E, MxI; F, MxII. Scale bars: 0.05 mm. 105
Figure 45. Arisubathynella pocheonensis sp. nov., holotype male. A, ThI; B, ThII; C, ThIII. Scale bar: 0.05 mm. 106
Figure 46. Arisubathynella pocheonensis sp. nov., holotype male. A, protopod, epipod and endopod of ThIV; B, exopod of ThIV; C, ThV; D, ThVI. Scale bar: 0.05 mm. 107
Figure 47. Arisubathynella pocheonensis sp. nov., (A-C) holotype male, (D) allotype female. A, ThVII; B, ThVIII; C, Pld; D, ThVIII and Pld; Scale bars: 0.05 mm. 108
Figure 48. Arisubathynella pocheonensis sp. nov., holotype male. A, Upd; B, pleotelson and furcal rami. Scale bars: 0.05 mm. 109
Figure 49. Map showing the distribution of all previously reported Eobathynella species. 116
Figure 50. Map showing the collecting sites of Eobathynella species in this study. The type locality of new species is marked with red square. The numbers in the map correspond with the... 117
Figure 51. Habitat photographs of E. gwangjuensis sp. nov. 118
Figure 52. Eobathynella cf. yeojuensis from Yeongju-si, South Korea (female). Scale bar: 0.5 mm. 119
Figure 53. Eobathynella cf. yeojuensis from Uiseong-gun, South Korea (female), Scale bar: 0.5 mm. 119
Figure 54. Eobathynella gwangjuensis sp. nov., (A) paratype female, (B-E) holotype female. A, habitus; B, AI; C, AII; D, Lbr; E, Md. Scale bars: A = 0.5 mm; B-E = 0.05 mm. 120
Figure 55. Eobathynella gwangjuensis sp. nov., holotype female. A, MxI; B, MxII; C, ThI; D, ThII; E, ThIII. Scale bars: 0.05 mm. 121
Figure 56. Eobathynella gwangjuensis sp. nov., holotype female. A, ThIV; B, ThV; C, ThVI; D, ThVII. Scale bar: 0.05 mm. 122
Figure 57. Eobathynella gwangjuensis sp. nov., holotype female. A, pleotelson and furcal rami; B, Upd. Scale bar: 0.05 mm. 123
Figure 58. Eobathynella gwangjuensis sp. nov., yellow arrows pointing numerous spinules on basipod of male ThVIII. (A, B) paratype males, (C) paratype female. A, ThVIII; B, Th VIII... 124
Figure 59. Distribution map of all previously reported Hangangbathynella species (black solid circle) and type locality of new species of Hangangbathynella (red solid square). The numbers in... 132
Figure 60. Habitat photograph of H. mihoensis sp. nov. 133
Figure 61. Hangangbathynella taechooni, female. Scale bar: 0.5 mm. 133
Figure 62. Photograph of Male ThVIII of H. taechooni using scanning electron microscope (SEM). A, ThVIII; B, Exopod of ThVIII. Scale bars: A = 0.5 mm, B = 0.02 mm. 134
Figure 63. Hangangbathynella mihoensis sp. nov., holotype male. A, habitus; B, AI; C, AII; D, Lbr; E, Md; F, MxI; G, MxII. Scale bars: A = 0.5 mm; B-G = 0.05 mm. 135
Figure 64. Hangangbathynella mihoensis sp. nov., holotype male. A-E, ThI-V. Scale bar: 0.05 mm. 136
Figure 65. Hangangbathynella mihoensis sp. nov., (A, B) holotype male, (C, D) paratype male. A, ThVI; B, ThVII; C, ThVIII. Scale bars: A-C = 0.5 mm; D = 0.05 mm. 137
Figure 66. Hangangbathynella mihoensis sp. nov., holotype male. A, pleotelson and furcal rami; B, Upd. Scale bars: 0.05 mm. 138
Figure 67. Hangangbathynella mihoensis sp. nov., (A-D, F) paratype female; (E) allotype female; (G, H) paratype male. A, AII; B, Lbr; C, Md; D, MxI; E, ThVIII; F, pleotelson and... 139
Figure 68. Map showing the distribution of all previously reported Nipponbathynella species. 148
Figure 69. Map showing the type localities of two new Nipponbathynella species. The numbers in the map correspond with the site number indicated in Table 1. 149
Figure 70. Habitat photograph of N. yeongchunensis sp. nov. 150
Figure 71. Nipponbathynella gapcheonensis sp. nov., female. Scale bar: 0.5 mm. 151
Figure 72. Nipponbathynella yeongchunensis sp. nov., female. Scale bar: 0.5 mm. 151
Figure 73. Nipponbathynella gapcheonensis sp. nov., holotype female. A, AI; B, AII; C, Lbr; D, Md; E, MxI. Scale bars: 0.05 mm. 152
Figure 74. Nipponbathynella gapcheonensis sp. nov., holotype female. A, MxII; B, ThI; C, ThII. Scale bars: 0.05 mm. 153
Figure 75. Nipponbathynella gapcheonensis sp. nov., holotype female. A, ThIII; B, ThIV; C, ThV. Scale bars: 0.05 mm. 154
Figure 76. Nipponbathynella gapcheonensis sp. nov., holotype female. A, ThVI; B, Th VII; C, ThVIII. Scale bars: 0.05 mm. 155
Figure 77. Nipponbathynella gapcheonensis sp. nov., (A, B) holotype female; (C) paratype male. A, Upd; B, Pleotelson and furcal rami; C, ThVIII. Scale bars: 0.05 mm. 156
Figure 78. Nipponbathynella yeongchunensis sp. nov., holotype female. A, AI; B, AII; C, Lbr; D, Md; E, MxI. Scale bars: 0.05 mm. 157
Figure 79. Nipponbathynella yeongchunensis sp. nov., holotype female. A, MxII; B, ThI, C, ThII; D, ThIII. Scale bars: 0.05 mm. 158
Figure 80. Nipponbathynella yeongchunensis sp. nov., holotype female. A, ThIV; B, ThV; C, ThVI. Scale bars: 0.05 mm. 159
Figure 81. Nipponbathynella yeongchunensis sp. nov., (A, B) holotype female; (C) Allotype male. A, ThVII; B, ThVIII; C, ThVIII. Scale bars: 0.05 mm. 160
Figure 82. Nipponbathynella yeongchunensis sp. nov., holotype female. A, Urd; B, Pleotelson and furcal rami. Scale bars: 0.05 mm. 161
Figure 83. Map showing the type locality of Parabathynellidae gen. et sp. nov. The number in the map correspond with the site number indicated in Table 1. 169
Figure 84. Parabathynellidae gen. et sp. nov., holotype male. A, AI; B, AII; C, Lbr; D, Md; E, MxI. Scale bars: 0.05 mm. 170
Figure 85. Parabathynellidae gen. et sp. nov., holotype male. A, MxII; B, ThI; C, ThII; D, ThIII. Scale bars: 0.05 mm. 171
Figure 86. Parabathynellidae gen. et sp. nov., holotype male. A, ThIV; B, ThV; C, ThVI; D, ThVII. Scale bars: 0.05 mm. 172
Figure 87. Parabathynellidae gen. et sp. nov., holotype male. A, ThVIII; B, Upd; C, pleotelson and furcal rami. Scale bars: 0.05 mm. 173
Figure 88. 18S-based (1,761 bp) ML tree based on TIM2e+R2 model for seven Korean genera, including two new genera proposed in the present study. 177
Figure 89. 18S-based ML tree and map showing the sites correspond to the colors and shapes in the ML tree. A, 18S-based (2,016 bp) ML tree based on TNe+R3 model for family Bathynellidae including newly determined sequences of Korea (blue box) and Mongolia specimens (yellow... 185
Figure 90. ML and BI analyses based on the 18S rDNA sequences and map showing the representing genera of region. A, 18S-based (1,800 bp) ML and BI phylogenetic reconstruction based on TNe+R3 and GTR+I+G model for respective analysis (majority consensus tree of ML). The... 195
Figure 91. An estimated divergence generated from relaxed molecular clock analysis using 18S rDNA sequences. Estimated mean divergence time (Mya) and posterior probabilities (PP) are annotated at nodes. The 95 % highest posterior density (HPD) interval is marked by horizonal... 197