Hostname: page-component-848d4c4894-75dct Total loading time: 0 Render date: 2024-05-06T08:32:10.730Z Has data issue: false hasContentIssue false

Electron backscatter diffraction (EBSD) study of elongatoolithid eggs from China with microstructural and parataxonomic implications

Published online by Cambridge University Press:  25 April 2024

Xufeng Zhu
Affiliation:
Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, 100044 Beijing, China. National Natural History Museum of China, 100050 Beijing, China. College of Earth and Planetary Sciences, University of Chinese Academy of Sciences, 100049 Beijing, China.
Qiang Wang*
Affiliation:
Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, 100044 Beijing, China.
Xiaolin Wang
Affiliation:
Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, 100044 Beijing, China. College of Earth and Planetary Sciences, University of Chinese Academy of Sciences, 100049 Beijing, China. Centre for Research and Education on Biological Evolution and Environment, Nanjing University, 210023 Nanjing, China.
*
Corresponding author: Qiang Wang; Email: wangqiang@ivpp.ac.cn

Abstract

Electron backscatter diffraction (EBSD) has been widely used in recent studies of eggshells for its convenience in collecting in situ crystallographic information. China has a wide variety of dinosaur eggshells, although nearly none have been studied with this technique. Elongatoolithid eggs include many oogenera, although the microstructural differences of some were not highly appreciated, leading to several parataxonomic problems. In this paper, we surveyed seven elongatoolithid oogenera in China using EBSD in order to acquire more information about their microstructural variation. It is shown in this paper that in some elongatoolithid eggshells, scaly calcite grains that form the squamatic ultrastructure are not the only form of calcite in the continuous layer. Large columnar grains separated by high-angled grain boundaries and slender subgrains separated by radially arranged low-angled grain boundaries could exist in certain areas of the eggshells such as Macroolithus and Macroelongatoolithus. This paper discusses the criteria for identifying squamatic ultrastructure and proposes type I (rich in rugged high-angled grain boundaries) and type II (rich in both rugged high- and low-angled grain boundaries) squamatic ultrastructures. A pathological layer is found in Undulatoolithus pengi. An external zone is identified in the eggshell of Heishanoolithus changii, which does not support its position within the oofamily Elongatoolithidae. We argue that Paraelongatoolithus no longer belongs to Elongatoolithidae based on a combination of reticulated ornamentation, columnar continuous layer, and acicular mammillae. The high structural variation in elongatoolithid eggshells also implies that it may be inappropriate to relate all previous elongatoolithid eggshells to oviraptorosaurs and assume they are non-monophyletic.

Type
Article
Copyright
Copyright © The Author(s), 2024. Published by Cambridge University Press on behalf of Paleontological Society

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Literature Cited

Agnolin, F. L., Powell, J. E., Novas, F. E., and Kundrát, M.. 2012. New alvarezsaurid (Dinosauria, Theropoda) from uppermost Cretaceous of north-western Patagonia with associated eggs. Cretaceous Research 35:3356.CrossRefGoogle Scholar
Bailleul, A. M., O'Connor, J., Zhang, S., Li, Z., Wang, Q., Lamanna, M. C., Zhu, X., and Zhou, Z.. 2019. An Early Cretaceous enantiornithine (Aves) preserving an unlaid egg and probable medullary bone. Nature Communications 10:1275.CrossRefGoogle ScholarPubMed
Bi, S., Amiot, R., Peyre de Fabrègues, C., Pittman, M., Lamanna, M. C., Yu, Y., Yu, C., et al. 2021. An oviraptorid preserved atop an embryo-bearing egg clutch sheds light on the reproductive biology of non-avialan theropod dinosaurs. Science Bulletin 66:947954.CrossRefGoogle ScholarPubMed
Bouwman, H., Yohannes, Y. B., Nakayama, S. M. M., Motohira, K., Ishizuka, M., Humphries, M. S., van der Schyff, V., du Preez, M., Dinkelmann, A., and Ikenaka, Y.. 2019. Evidence of impacts from DDT in pelican, cormorant, stork, and egret eggs from KwaZulu-Natal, South Africa. Chemosphere 225:647658.CrossRefGoogle ScholarPubMed
Canudo, J. I., Gasca, J. M., Aurell, M., Badiola, A., Blain, H.-A., Cruzado-Caballero, P., Gomez-Fernández, D., et al. 2010. La Cantalera: an exceptional window onto the vertebrate biodiversity of the Hauterivian–Barremian transition in the Iberian Peninsula. Journal of Iberian Geology 36:205224.CrossRefGoogle Scholar
Cheng, Y., Ji, Q., Wu, X., and Shan, H.. 2008. Oviraptorosaurian eggs (Dinosauria) with embryonic skeletons discovered for the first time in China. Acta Geologica Sinica—English Edition 82:10891094.CrossRefGoogle Scholar
Choi, S. 2020. Paleontological, neontological, and taphonomic studies for amniote eggshells with analytical approaches. Seoul National University, Seoul.Google Scholar
Choi, S., and Lee, Y.-N.. 2019. Possible Late Cretaceous dromaeosaurid eggshells from South Korea: a new insight into dromaeosaurid oology. Cretaceous Research 103:104167.CrossRefGoogle Scholar
Choi, S., Han, S., Kim, N.-H., and Lee, Y.-N.. 2018. A comparative study of eggshells of Gekkota with morphological, chemical compositional and crystallographic approaches and its evolutionary implications. PLoS ONE 13:e0199496.CrossRefGoogle ScholarPubMed
Choi, S., Han, S., and Lee, Y.-N.. 2019. Electron backscatter diffraction (EBSD) analysis of maniraptoran eggshells with important implications for microstructural and taphonomic interpretations. Palaeontology 62:777803.CrossRefGoogle Scholar
Choi, S., Barta, D. E., Moreno-Azanza, M., Kim, N., Shaw, C. A., and Varricchio, D. J.. 2022a. Microstructural description of the maniraptoran egg Protoceratopsidovum. Papers in Palaeontology 8:e1430.CrossRefGoogle Scholar
Choi, S., Kim, H., Paik, I., Park, Y., Jung, H., and Xu, X.. 2022b. Turtle eggs from the Lower Cretaceous Hasandong Formation (South Korea) with relict aragonite under significant thermal maturity. Journal of Vertebrate Paleontology 42:e2183866.CrossRefGoogle Scholar
Choi, S., Kim, N.-H., Kim, H.-I., Kweon, J. J., Lee, S. K., Zhang, S., and Varricchio, D. J.. 2022c. Preservation of aragonite in Late Cretaceous (Campanian) turtle eggshell. Palaeogeography, Palaeoclimatology, Palaeoecology 585:110741.CrossRefGoogle Scholar
Choi, S., Hauber, M. E., Legendre, L. J., Kim, N.-H., Lee, Y.-N., and Varricchio, D. J.. 2023. Microstructural and crystallographic evolution of palaeognath (Aves) eggshells. eLife 12:e81092.CrossRefGoogle ScholarPubMed
Clark, J. M., Norell, I. M. A., and Chiappe, L. M.. 1999. An oviraptorid skeleton from the Late Cretaceous of Ukhaa Tolgod, Mongolia, preserved in an avianlike brooding position over an oviraptorid nest. American Museum Novitates 3265:136.Google Scholar
Cusack, M. 2016. Biomineral electron backscatter diffraction for palaeontology. Palaeontology 59:171179.CrossRefGoogle Scholar
Cusack, M., Dauphin, Y., Chung, P., Pérez-Huerta, A., and Cuif, J.-P.. 2008a. Multiscale structure of calcite fibres of the shell of the brachiopod Terebratulina retusa. Journal of Structural Biology 164:96100.CrossRefGoogle ScholarPubMed
Cusack, M., England, J., Dalbeck, P., Tudhope, A. W., Fallick, A. E., and Allison, N.. 2008b. Electron backscatter diffraction (EBSD) as a tool for detection of coral diagenesis. Coral Reefs 27:905911.CrossRefGoogle Scholar
Dalbeck, P., and Cusack, M.. 2006. Crystallography (electron backscatter diffraction) and chemistry (electron probe microanalysis) of the avian eggshell. Crystal Growth & Design 6:25582562.CrossRefGoogle Scholar
Dauphin, Y., Luquet, G., Perez-Huerta, A., and Salomé, M.. 2018. Biomineralization in modern avian calcified eggshells: similarity versus diversity. Connective Tissue Research 59:6773.CrossRefGoogle ScholarPubMed
Dong, Z.-M., and Currie, P. J.. 1996. On the discovery of an oviraptorid skeleton on a nest of eggs at Bayan Mandahu, Inner Mongolia, People's Republic of China. Canadian Journal of Earth Sciences 33:631636.CrossRefGoogle Scholar
Eagle, R. A., Enriquez, M., Grellet-Tinner, G., Pérez-Huerta, A., Hu, D., Tütken, T., Montanari, S., et al. 2015. Isotopic ordering in eggshells reflects body temperatures and suggests differing thermophysiology in two Cretaceous dinosaurs. Nature Communications 6:8296.CrossRefGoogle ScholarPubMed
Ewert, M. A., Firth, S. J., and Nelson, C. E.. 1984. Normal and multiple eggshells in batagurine turtles and their implications for dinosaurs and other reptiles. Canadian Journal of Zoology 62:18341841.CrossRefGoogle Scholar
Fanti, F., Currie, P. J., and Badamgarav, D.. 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE 7:e31330.CrossRefGoogle ScholarPubMed
Grellet-Tinner, G., and Dyke, G. J.. 2005. The eggshell of the Eocene bird Lithornis. Acta Palaeontologica Polonica 50:831835.Google Scholar
Grellet-Tinner, G., and Makovicky, P.. 2006. A possible egg of the dromaeosaur Deinonychus antirrhopus: phylogenetic and biological implications. Canadian Journal of Earth Sciences 43:705719.CrossRefGoogle Scholar
Grellet-Tinner, G., Corsetti, F., and Buscalioni, A. D.. 2010. The importance of microscopic examinations of eggshells: Discrimination of bioalteration and diagenetic overprints from biological features. Journal of Iberian Geology 36:181192.CrossRefGoogle Scholar
Grellet-Tinner, G., Sim, C. M., Kim, D. H., Trimby, P., Higa, A., An, S. L., Oh, H. S., Kim, T., and Kardjilov, N.. 2011. Description of the first lithostrotian titanosaur embryo in ovo with neutron characterization and implications for lithostrotian Aptian migration and dispersion. Gondwana Research 20:621629.CrossRefGoogle Scholar
Grellet-Tinner, G., Murelaga, X., Larrasoaña, J. C., Silveira, L. F., Olivares, M., Ortega, L. A., Trimby, P. W., and Pascual, A.. 2012. The first occurrence in the fossil record of an aquatic avian twig-nest with Phoenicopteriformes eggs: evolutionary implications. PLoS ONE 7:e46972.CrossRefGoogle ScholarPubMed
Grellet-Tinner, G., Spooner, N. A., and Worthy, T. H.. 2016. Is the “Genyornis” egg of a mihirung or another extinct bird from the Australian dreamtime? Quaternary Science Reviews 133:147164.CrossRefGoogle Scholar
Grellet-Tinner, G., Lindsay, S., and Thompson, M.. 2017. The biomechanical, chemical, and physiological adaptations of the eggs of two Australian megapodes to their nesting strategies and their implications for extinct titanosaur dinosaurs. Journal of Microscopy 267:237246.CrossRefGoogle ScholarPubMed
Han, F., Yu, Y., Zhang, S., Zeng, R., Wang, X., Cai, H., Wu, T., et al. 2023. Exceptional early Jurassic fossils with leathery eggs shed light on dinosaur reproductive biology. National Science Review 11:nwad258.CrossRefGoogle Scholar
Hickey, J. J., and Anderson, D. W.. 1968. Chlorinated hydrocarbons and eggshell changes in raptorial and fish-eating birds. Science 162:271273.CrossRefGoogle ScholarPubMed
Holm, L., Blomqvist, A., Brandt, I., Brunström, B., Ridderstråle, Y., and Berg, C.. 2006. Embryonic exposure to o,p′-DDT causes eggshell thinning and altered shell gland carbonic anhydrase expression in the domestic hen. Environmental Toxicology and Chemistry 25:27872793.CrossRefGoogle Scholar
Jackson, F. D., and Schmitt, J. G.. 2008. Recognition of vertebrate egg abnormalities in the Upper Cretaceous fossil record. Cretaceous Research 29:2739.CrossRefGoogle Scholar
Jackson, F. D., and Varricchio, D. J.. 2003. Abnormal, multilayered eggshell in birds: implications for dinosaur reproductive anatomy. Journal of Vertebrate Paleontology 23:699702.CrossRefGoogle Scholar
Jackson, F. D., and Varricchio, D. J.. 2010. Fossil eggs and eggshell from the lowermost Two Medicine Formation of western Montana, Sevenmile Hill locality. Journal of Vertebrate Paleontology 30:11421156.CrossRefGoogle Scholar
Jackson, F. D., Garrido, A., Schmitt, J. G., Chiappe, L. M., Dingus, L., and Loope, D. B.. 2004. Abnormal, multilayered titanosaur (Dinosauria: Sauropoda) eggs from in situ clutches at the Auca Mahuevo locality, Neuquén Province, Argentina. Journal of Vertebrate Paleontology 24:913922.CrossRefGoogle Scholar
Jain, S., Bajpai, S., Kumar, G., and Pruthi, V.. 2016. Microstructure, crystallography and diagenetic alteration in fossil ostrich eggshells from Upper Palaeolithic sites of Indian peninsular region. Micron 84:7278.CrossRefGoogle ScholarPubMed
Jin, X., Varricchio, D. J., Poust, A. W., and He, T.. 2020. An oviraptorosaur adult–egg association from the Cretaceous of Jiangxi Province, China. Journal of Vertebrate Paleontology 39:e1739060.CrossRefGoogle Scholar
Jin, X. S., Azuma, Y., Jackson, F. D., and Varricchio, D. J.. 2007. Giant dinosaur eggs from the Tiantai basin, Zhejiang Province, China. Canadian Journal of Earth Sciences 44:8188.CrossRefGoogle Scholar
Keymer, I. F. 1980. Disorders of the avian female reproductive system. Avian Pathology 9:405419.CrossRefGoogle ScholarPubMed
Kim, N.-H., Choi, S., Kim, S., and Lee, Y.-N.. 2019. A new faveoloolithid oogenus from the Wido Volcanics (Upper Cretaceous), South Korea and a new insight into the oofamily Faveoloolithidae. Cretaceous Research 100:145163.CrossRefGoogle Scholar
Lincer, J. L. 1975. DDE-induced eggshell-thinning in the American kestrel: a comparison of the field situation and laboratory results. Journal of Applied Ecology 12:781.CrossRefGoogle Scholar
Lundholm, C. E. 1997. DDE-induced eggshell thinning in birds: Effects of p,p′-DDE on the calcium and prostaglandin metabolism of the eggshell gland. Comparative Biochemistry and Physiology Part C: Pharmacology, Toxicology and Endocrinology 118:113128.Google ScholarPubMed
Mikhailov, K. E. 1991. Classification of fossil eggshells of amniotic vertebrates. Acta Palaeontologica Polonica 36:193238.Google Scholar
Mikhailov, K. E. 1994. Theropod and protoceratopsian dinosaur eggs from the Cretaceous of Mongolia and Kazakhstan. Paleontological Journal 28:101120.Google Scholar
Mikhailov, K. E. 1997a. Avian eggshells: an atlas of scanning electron micrographs. British Ornithologists’ Club, Tring.Google Scholar
Mikhailov, K. E. 1997b. Fossil and recent eggshell in amniotic vertebrates: fine structure, comparative morphology and classification. Special Papers in Palaeontology 56:180.Google Scholar
Mikhailov, K. E. 2014. Eggshell structure, parataxonomy and phylogenetic analysis: some notes on articles published from 2002 to 2011. Historical Biology 26:144154.CrossRefGoogle Scholar
Mikhailov, K. E. 2019. Conservative nature of biomineral structures as a challenge for the cladistic method of phylogeny reconstruction (illustrated by two groups of dinosaur eggs). Paleontological Journal 53:551565.CrossRefGoogle Scholar
Moreno-Azanza, M., Mariani, E., Bauluz, B., and Canudo, J. I.. 2013. Growth mechanisms in dinosaur eggshells: an insight from electron backscatter diffraction. Journal of Vertebrate Paleontology 33:121130.CrossRefGoogle Scholar
Moreno-Azanza, M., Canudo, J. I., and Gasca, J. M.. 2014. Unusual theropod eggshells from the Early Cretaceous Blesa Formation of the Iberian Range, Spain. Acta Palaeontologica Polonica 59:843854.Google Scholar
Moreno-Azanza, M., Bauluz, B., Canudo, J. I., Gasca, J. M., and Torcida Fernández-Baldor, F.. 2016. Combined use of electron and light microscopy techniques reveals false secondary shell units in Megaloolithidae eggshells. PLoS ONE 11:e0153026.CrossRefGoogle ScholarPubMed
Moreno-Azanza, M., Bauluz, B., Canudo, J. I., and Mateus, O.. 2017. The conservative structure of the ornithopod eggshell: electron backscatter diffraction characterization of Guegoolithus turolensis from the Early Cretaceous of Spain. Journal of Iberian Geology 43:235243.CrossRefGoogle Scholar
Moreno-Azanza, M., Díaz-Berenguer, E., Silva-Casal, R., Pérez-García, A., Badiola, A., and Canudo, J. I.. 2021. Recognizing a lost nesting ground: First unambiguous Testudines eggshells from the Eocene, associated with the pleurodiran Eocenochelus (Huesca, Northern Spain). Palaeogeography, Palaeoclimatology, Palaeoecology 576:110526.CrossRefGoogle Scholar
Norell, M. A., Clark, J. M., Demberelyin, D., Rhinchen, B., Chiappe, L. M., Davidson, A. R., McKenna, M. C., Altangerel, P., and Novacek, M. J.. 1994. A theropod dinosaur embryo and the affinities of the flaming cliffs dinosaur eggs. Science 266:779782.CrossRefGoogle ScholarPubMed
Norell, I. M. A., Clark, J. M., Chiappe, L. M., and Dashzeveg, D.. 1995. A nesting dinosaur. Nature 378:774776.CrossRefGoogle Scholar
Oestreicher, M. I., Shuman, D. H., and Wurster, C. F.. 1971. DDE reduces medullary bone formation in birds. Nature 229:571571.CrossRefGoogle ScholarPubMed
Oser, S. E., Chin, K., Sertich, J. J. W., Varricchio, D. J., Choi, S., and Rifkin, J.. 2021. Tiny, ornamented eggs and eggshell from the Upper Cretaceous of Utah represent a new ootaxon with theropod affinities. Scientific Reports 11:10021.CrossRefGoogle ScholarPubMed
Pérez-Huerta, A., and Cusack, M.. 2009. Optimizing electron backscatter diffraction of carbonate biominerals—resin type and carbon coating. Microscopy and Microanalysis 15:197203.CrossRefGoogle ScholarPubMed
Pérez-Huerta, A., and Dauphin, Y.. 2016. Comparison of the structure, crystallography and composition of eggshells of the guinea fowl and graylag goose. Zoology 119:5263.CrossRefGoogle ScholarPubMed
Pérez-Huerta, A., Dauphin, Y., Cuif, J. P., and Cusack, M.. 2011. High resolution electron backscatter diffraction (EBSD) data from calcite biominerals in recent gastropod shells. Micron 42:246251.CrossRefGoogle ScholarPubMed
Prior, D. J., Mariani, E., and Wheeler, J.. 2009. EBSD in the earth sciences: applications, common practice, and challenges. Pp. 345360 in Schwartz, A. J., Kumar, M., Adams, B. L., and Field, D. P., eds. Electron backscatter diffraction in materials science. Springer US, Boston.CrossRefGoogle Scholar
Pruett-Jones, S. G., White, C. M., and Emison, W. B.. 1981. Eggshell thinning and organochlorine residues in eggs and prey of peregrine falcons from Victoria, Australia. Emu—Austral Ornithology 80:281287.CrossRefGoogle Scholar
Pu, H., Zelenitsky, D. K., , J., Currie, P. J., Carpenter, K., Xu, L., Koppelhus, E. B., et al. 2017. Perinate and eggs of a giant caenagnathid dinosaur from the Late Cretaceous of central China. Nature Communications 8:14952.CrossRefGoogle ScholarPubMed
Romanoff, A. L., and Romanoff, A. J.. 1949. The avian egg. Wiley, New York.Google Scholar
Sabath, K. 1991. Upper Cretaceous amniotic eggs from Gobi Desert. Acta Palaeontologica Polonica 36:151192.Google Scholar
Sato, T., Cheng, Y.-N., Wu, X., Zelenitsky, D. K., and Hsiao, Y.. 2005. A pair of shelled eggs inside a female dinosaur. Science 308:375375.CrossRefGoogle ScholarPubMed
Schwarzer, R. A., Field, D. P., Adams, B. L., Kumar, M., and Schwartz, A. J.. 2009. Present state of electron backscatter diffraction and prospective developments. Pp. 120 in Schwartz, A. J., Kumar, M., Adams, B. L., and Field, D. P., eds. Electron backscatter diffraction in materials science. Springer US, Boston.Google Scholar
Sellés, A. G., Vila, B., and Galobart, À.. 2017. Evidence of reproductive stress in titanosaurian sauropods triggered by an increase in ecological competition. Scientific Reports 7:13827.CrossRefGoogle ScholarPubMed
Shao, Z., Fan, S., Jia, S., Tanaka, K., and , J.. 2014. Intact theropod dinosaur eggs with embryonic remains from the Late Cretaceous of southern China. Geological Bulletin of China 33:941948.Google Scholar
Sochava, A. V. 1969. Dinosaur eggs from the Upper Cretaceous of the Gobi Desert. Paleontological Journal 4:517527.Google Scholar
Tanaka, K., Lv, J., Kobayashi, Y., Zelenitsky, D. K., Xu, L., Jia, S., Qing, S., and Tang, M.. 2011. Description and phylogenetic position of dinosaur eggshells from the Luanchuan area of western Henan Province, China. Acta Geologica Sinica—English Edition 85:6674.CrossRefGoogle Scholar
Tanaka, K., Zelenitsky, D. K., Saegusa, H., Ikeda, T., DeBuhr, C. L., and Therrien, F.. 2016. Dinosaur eggshell assemblage from Japan reveals unknown diversity of small theropods. Cretaceous Research 57:350363.CrossRefGoogle Scholar
Trimby, P., and Grellet-Tinner, G.. 2011. Using electron backscatter diffraction to aid identification of fossilized dinosaur eggshells. Microscopy and Microanalysis 17:574575.CrossRefGoogle Scholar
Tyler, C. 1965. A study of the egg shells of the Sphenisciformes. Journal of Zoology 147:119.Google Scholar
Varricchio, D. J., and Jackson, F. D.. 2004. A phylogenetic assessment of prismatic dinosaur eggs from the Cretaceous Two Medicine Formation of Montana. Journal of Vertebrate Paleontology 24:931937.CrossRefGoogle Scholar
Vila, B., Sellés, A. G., and Beetschen, J.-C.. 2017. The controversial Les Labadous eggshells: a new and peculiar dromaeosaurid (Dinosauria: Theropoda) ootype from the Upper Cretaceous of Europe. Cretaceous Research 72:117123.CrossRefGoogle Scholar
Wang, Q., Wang, X.-L., Zhao, Z.-K., and Jiang, Y.-G.. 2010a. A new oogenus of Elongatoolithidae from the Upper Cretaceous Chichengshan Formation of Tiantai Basin, Zhejiang Province. Vertebrata PalAsiatica 48:111118.Google Scholar
Wang, Q., Zhao, Z., Wang, X., Jiang, Y., and Zhang, S.. 2010b. A new oogenus of macroelongatoolithid eggs from the Upper Cretaceous Chichengshan Formation of the Tiantai Basin, Zhejiang Province and a revision of the macroelongatoolithids. Acta Palaeontologica Sinica 49:7386.Google Scholar
Wang, Q., Zhao, Z., Wang, X., Li, N., and Zou, S.. 2013. A new form of Elongatoolithidae, Undulatoolithus pengi oogen. et oosp. nov. from Pingxiang, Jiangxi, China. Zootaxa 3746:194200.CrossRefGoogle ScholarPubMed
Wang, S., Zhang, S., Sullivan, C., and Xu, X.. 2016. Elongatoolithid eggs containing oviraptorid (Theropoda, Oviraptorosauria) embryos from the Upper Cretaceous of Southern China. BMC Evolutionary Biology 16:67.CrossRefGoogle ScholarPubMed
Weishampel, D. B., Fastovsky, D. E., Watabe, M., Varricchio, D., Jackson, F., Tsogtbaatar, K., and Barsbold, R.. 2008. New oviraptorid embryos from Bugin-Tsav, Nemegt Formation (Upper Cretaceous), Mongolia, with insights into their habitat and growth. Journal of Vertebrate Paleontology 28:11101119.CrossRefGoogle Scholar
Wu, H.-J., Tseng, Y.-C., Tsao, S.-H., Chiang, P.-L., Tai, W.-Y., Hsieh, H.-I., Yu, H.-T., and Juang, J.-Y.. 2023. A comparative study on the microstructures, mineral content, and mechanical properties of non-avian reptilian eggshells. Biology 12:688.CrossRefGoogle ScholarPubMed
Xing, L., Niu, K., Ma, W., Zelenitsky, D. K., Yang, T.-R., and Brusatte, S. L.. 2022. An exquisitely preserved in-ovo theropod dinosaur embryo sheds light on avian-like prehatching postures. iScience 25:103516.CrossRefGoogle ScholarPubMed
Xu, L., Xie, J., Zhang, S., Choi, S., Kim, N.-H., Gao, D., Jin, X., Jia, S., and Gao, Y.. 2022. Fossil turtle eggs from the Upper Cretaceous Gaogou Formation, Xiaguan-Gaoqiu Basin, Neixiang County, Henan Province, China: Interpretation of the transformation from aragonite to calcite in fossil turtle eggshell. Cretaceous Research 134:105166.CrossRefGoogle Scholar
Yang, T.-R., Wiemann, J., Xu, L., Cheng, Y.-N., Wu, X., and Sander, M.. 2019. Reconstruction of oviraptorid clutches illuminates their unique nesting biology. Acta Palaeontologica Polonica 64:581596.CrossRefGoogle Scholar
Young, C.-C. 1954. Fossil reptilian eggs from Laiyang, Shantung, China. Scientia Sinica 3:505522.Google Scholar
Young, C.-C. 1965. Fossil eggs from Nanhsiung, Kwangtung and Kanchou, Kiangsi. Vertebrata PalAsiatica 9:159170.Google Scholar
Zelenitsky, D. K., and Sloboda, W. J.. 2005. Eggshells. Pp. 398404 in Currie, P. J. and Koppelhus, E. B., eds. Dinosaur provincial park: a spectacular ancient ecosystem revealed. Indiana University Press, Bloomington.Google Scholar
Zelenitsky, D. K., and Therrien, F.. 2008. Unique maniraptoran egg clutch from the Upper Cretaceous Two Medicine Formation of Montana reveals theropod nesting behaviour. Palaeontology 51:12531259.CrossRefGoogle Scholar
Zelenitsky, D. K., Hills, L. V., and Currie, P. J.. 1996. Parataxonomic classification of ornithoid eggshell fragments from the Oldman Formation (Judith River Group; Upper Cretaceous), southern Alberta. Canadian Journal of Earth Sciences 33:16551667.CrossRefGoogle Scholar
Zelenitsky, D. K., Carpenter, K., and Currie, P. J.. 2000. First record of elongatoolithid theropod eggshell from North America: the Asian oogenus Macroelongatoolithus from the Lower Cretaceous of Utah. Journal of Vertebrate Paleontology 20:130138.CrossRefGoogle Scholar
Zhao, H., and Zhao, Z.-K.. 1999. A new form of elongatoolithid dinosaur eggs from the Lower Cretaceous Shahai Formation of Heishan, Liaoning Province. Vertebrata PalAsiatica 37:278284.Google Scholar
Zhao, Z. 1975. The microstructure of fossil dinosaur eggs from Nanxiong County, Guangdong Province: concurrent with a discussion on the problem of the classification of dinosaur eggs. Vertebrata PalAsiatica 13:105117.Google Scholar
Zhao, Z., Wang, Q., and Zhang, S.. 2015. Dinosaur eggs. Pp. 1163 in Palaeovertbrata Sinica, Vol. 2. Science Press, Beijing.Google Scholar
Zhu, X., Fang, K., Wang, Q., Deng, L., Liu, Y., Wen, J., Wang, X., and Wang, X.. 2021. The first Similifaveoloolithidae (Wormoolithus luxiensis oogen. et oosp. nov.) from the Upper Cretaceous of Jiangxi Province, China. Historical Biology 33:689698.CrossRefGoogle Scholar