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Observations on Fur Development in E chidna ( M onotremata, M ammalia) Indicate that Spines Precede Hairs in Ontogeny

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Immuno‐Cross reactivity of transglutaminase and cornification marker proteins in the epidermis of vertebrates suggests common processes of soft cornification across species

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Chicken Corneocyte Cross-Linked Proteome

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Immunolocalization of specific keratin associated beta-proteins (beta-keratins) in the adhesive setae of Gekko gecko

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Keratinization of sheath and calamus cells in developing and regenerating feathers

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Structure of adhesive organ of the mountain‐stream catfish, Pseudocheneis sulcatus (Teleostei: Sisoridae)

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Cornification in developing claws of the common Australian skink (Lampropholis guichenoti) (Squamata, Lacertidae)

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Evolution of tissue-specific keratins as deduced from novel cDNA sequences of the lungfish Protopterus aethiopicus

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Skin lipid structure controls water permeability in snake molts

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Deleterious Mutations of a Claw Keratin in Multiple Taxa of Reptiles

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The Process of Cornification in the Horny Teeth of the Lamprey Involves Proteins in the Keratin Range and Other Keratin-Associated Proteins

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Cornification of the Beak of Rana dalmatina Tadpoles Suggests the Presence of Basic Keratin-Associated Proteins

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Keratinization in crocodilian scales and avian epidermis : evolutionary implications for the origin of avian apteric epidermis

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Morphogenesis of the horny jaws of Rana pipiens larvae

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Claw development and cornification in the passeraceous bird zebrafinch (Taeniatopygia guttata castanotis)

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Perspectives on Hair Evolution Based on Some Comparative Studies on Vertebrate Cornification

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Review: mapping epidermal beta-protein distribution in the lizard Anolis carolinensis shows a specific localization for the formation of scales, pads, and claws

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Cornification in the claw of the amphibianXenopus laevis(Pipidae, Anura) and comparison with claws in amniotes

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Comparative immunolocalization of keratin‐associated beta‐proteins (beta‐keratins) supports a new explanation for the cyclical process of keratinocyte differentiation in lizard epidermis

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Immunocytochemistry suggests that the prevalence of a sub‐type of beta‐proteins determines the hardness in the epidermis of the hard‐shelled turtle

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Cytological Aspects of the Differentiation of Barb Cells During the Formation of the Ramus of Feathers

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Ultrastructural immunolocalization of alpha-keratins and associated beta-proteins (beta-keratins) suggests a new interpretation on the process of hard and soft cornification in turtle epidermis

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The Fossil Record of Feather Evolution in the Mesozoic

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Ultrastructural characteristics of the process of cornification in developing claws of the brushtail possum (Trichosurus vulpecula)

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Presence of a glycine-cysteine-rich beta-protein in the oberhautchen layer of snake epidermis marks the formation of the shedding layer

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Transition from embryonic to adult epidermis in reptiles occurs by the production of corneous beta-proteins

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Th e Integument of Lungfi sh: General Structure and Keratin Composition

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The corneous layer of the claw in the lizard Anolis carolinensis mainly contains the glycine–cysteine-rich beta-protein HgGC3 in addition to hard keratins

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Immunocytochemistry indicates that glycine‐rich beta‐proteins are present in the beta‐layer, while cysteine‐rich beta‐proteins are present in beta‐ and alpha‐layers of snake epidermis

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Molding and Carving Cell Surfaces: The Joke of a Fold and the Origin and Evolution of Feathers

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One More Example of Morphological Convergence: Similarity between the Architectonics of Feather and Hair

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Immunolocalization of transglutaminase and cornification markers proteins in the epidermis of vertebrates suggests common processess of cornification across species.

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The integument of lungfish: General structure and keratin composition

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Immunogold labeling shows that glycine‐cysteine‐rich beta‐proteins are deposited in the O berhäutchen layer of snake epidermis in preparation to shedding

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Immunolocalization of beta‐proteins and alpha‐keratin in the epidermis of the soft‐shelled turtle explains the lack of formation of hard corneous material

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Gene duplication and recruitment of a specific tropomyosin into striated muscle cells in the jellyfish Podocoryne carnea

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The Early Origin of Feathers

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Update of the keratin gene family: evolution, tissue-specific expression patterns, and relevance to clinical disorders

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Comparison of vertebrate skin structure at class level: A review

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Convergent Evolution of Cysteine-Rich Keratins in Hard Skin Appendages of Terrestrial Vertebrates

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Review: mapping proteins localized in adhesive setae of the tokay gecko and their possible influence on the mechanism of adhesion

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Wanted, dead and alive: Why a multidisciplinary approach is needed to unlock hair treatment potential

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Evolution of Trichocyte Keratins

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Vertebrate keratinization evolved into cornification mainly due to transglutaminase and sulfhydryl oxidase activities on epidermal proteins: An immunohistochemical survey

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Evo Devo of the Vertebrates Integument

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Keratinization and Cornification are not equivalent processes but keratinization in fish and amphibians evolved into cornification in terrestrial vertebrates

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Loss of the ability to regenerate body appendages in vertebrates: from side effects of evolutionary innovations to gene loss

Zaraisky, A. G., Araslanova, K. R., Shitikov, A. D., & Tereshina, M. B. (2024) · Biological reviews/Biological reviews of the Cambridge Philosophical Society · doi:10.1111/brv.13102 · Cited by 14 · see citing works

Analysis and comparison of protein secondary structures in the rachis of avian flight feathers

Lin, P.-Y., Huang, P., Lee, Y.-C., & Ng, C. S. (2022) · PeerJ · doi:10.7717/peerj.12919 · Cited by 13 · see citing works

Structure of Keratins in Adhesive Gecko Setae Determined by Near-Edge X-ray Absorption Fine Structure Spectromicroscopy

Holler, K. R., Rasmussen, M. H., Baio, J. E., Jaye, C., Fischer, D. A., Gorb, S. N., & Weidner, T. (2022) · The Journal of Physical Chemistry Letters · doi:10.1021/acs.jpclett.2c00004 · Cited by 13 · see citing works

Keratinous and corneous-based products towards circular bioeconomy: A research review

Barone, G. D., Tagliaro, I., Oliver‐Simancas, R., Radice, M., Kalossaka, L. M., Mattei, M., Biundo, A., Pisano, I., & Jiménez‐Quero, A. (2024) · Environmental Science and Ecotechnology · doi:10.1016/j.ese.2024.100444 · Cited by 12 · see citing works

General aspects on skin development in vertebrates with emphasis on sauropsids epidermis

Alibardi, L. (2023) · Developmental Biology · doi:10.1016/j.ydbio.2023.05.007 · Cited by 12 · see citing works

Structure and function of skin in the pelagic sea snake, Hydrophis platurus

Lillywhite, H. B., & Menon, G. K. (2019) · Journal of Morphology · doi:10.1002/jmor.20963 · Cited by 12 · see citing works

Skin Appendage Proteins of Tetrapods: Building Blocks of Claws, Feathers, Hair and Other Cornified Epithelial Structures

Holthaus, K. B., Steinbinder, J., Sachslehner, A. P., & Eckhart, L. (2025) · Animals · doi:10.3390/ani15030457 · Cited by 12 · see citing works

An update on thermostable keratinases for protein engineering against feather pollutants

L, B. J. J., & Dhanasingh, I. (2025) · Applied Microbiology and Biotechnology · doi:10.1007/s00253-025-13459-5 · Cited by 11 · see citing works

Development, structure, and protein composition of the corneous beak in turtles

Alibardi, L. (2021) · The Anatomical Record · doi:10.1002/ar.24604 · Cited by 11 · see citing works

Cellular structure of dinosaur scales reveals retention of reptile-type skin during the evolutionary transition to feathers

Yang, Z., Jiang, B., Xu, J., & McNamara, M. E. (2024) · Nature Communications · doi:10.1038/s41467-024-48400-3 · Cited by 10 · see citing works

Early Permian synapsid impressions illuminate the origin of epidermal scales and aggregation behavior

Marchetti, L., Logghe, A., Buchwitz, M., & Fröbisch, J. (2025) · Current Biology · doi:10.1016/j.cub.2025.04.077 · Cited by 10 · see citing works

Keratin intermediate filament chains in tuatara (Sphenodon punctatus): A comparison of tuatara and human sequences

Parry, D. A. D., & Winter, D. J. (2021) · Journal of Structural Biology · doi:10.1016/j.jsb.2021.107706 · Cited by 10 · see citing works

Immunolocalization of epidermal differentiation complex proteins reveals distinct molecular compositions of cells that control structure and mechanical properties of avian skin appendages

Alibardi, L., & Eckhart, L. (2021) · Journal of Morphology · doi:10.1002/jmor.21357 · Cited by 9 · see citing works

High-Quality Genomes of Pangolins: Insights into the Molecular Basis of Scale Formation and Adaption to Myrmecophagous Diet

Yan, D., Luo, X., Tang, J., Xu, S., Huang, K., Wang, X., Feng, T., Que, T., Jia, M., Guo, X., et al. (2022) · Molecular Biology and Evolution · doi:10.1093/molbev/msac262 · Cited by 9 · see citing works

Creation of a peptide database of corneous beta-proteins of marine turtles for the identification of tortoiseshell: archaeological combs as case study

Solazzo, C., Soulat, J., & Cleland, T. P. (2021) · Royal Society Open Science · doi:10.1098/rsos.201857 · Cited by 7 · see citing works

Epidermal Differentiation Genes of the Common Wall Lizard Encode Proteins with Extremely Biased Amino Acid Contents

Holthaus, K. B., Sachslehner, A. P., Steinbinder, J., & Eckhart, L. (2024) · Genes · doi:10.3390/genes15091136 · Cited by 6 · see citing works

Keratinization and mucogenesis in the epidermis of an angler catfish Chaca chaca (Siluriformes, Chacidae): A Histochemical and fluorescence microscope investigation

Mistri, A., Kumari, U., Mittal, S., & Mittal, S. (2018) · Zoology · doi:10.1016/j.zool.2018.10.001 · Cited by 5 · see citing works

Structure, function and formation of the amniote skin pattern.

Dinh, C. D.-T., & Manceau, M. (2024) · Developmental Biology · doi:10.1016/j.ydbio.2024.09.011 · Cited by 5 · see citing works

Keratin intermediate filament chains in the European common wall lizard (Podarcis muralis) and a potential keratin filament crosslinker

Parry, D. A. D., & Winter, D. J. (2021) · Journal of Structural Biology · doi:10.1016/j.jsb.2021.107793 · Cited by 4 · see citing works

Immunolabeling for filaggrin and acidic keratins in the granular layer of mammalian epidermis indicates that an acidic-basic interaction is involved in cornification

Alibardi, L. (2024) · Tissue and Cell · doi:10.1016/j.tice.2024.102397 · Cited by 3 · see citing works

Keratinization and cornification of avian skin appendages during development. Insights from immunolabeling and electron microscopic studies

Alibardi, L. (2025) · Developmental Biology · doi:10.1016/j.ydbio.2025.02.019 · Cited by 3 · see citing works

Cystatin immunoreactivity in cornifying layers of the epidermis suggests a role in the formation of the epidermal barrier in amniotes

Alibardi, L. (2018) · Zoology · doi:10.1016/j.zool.2018.02.003 · Cited by 3 · see citing works

Immunolabeling indicates that sulfhydryl oxidase is absent in anamniote epidermis but marks the process of cornification in the skin of terrestrial vertebrates

Alibardi, L. (2020) · Journal of Morphology · doi:10.1002/jmor.21299 · Cited by 3 · see citing works

Lipids in the American Alligator stratum corneum provide insights into the evolution of vertebrate skin

Tam, E. A., Robb, F. E., & Champagne, A. M. (2024) · Comparative Biochemistry and Physiology Part A Molecular & Integrative Physiology · doi:10.1016/j.cbpa.2024.111620 · Cited by 2 · see citing works

Efficient DNA extraction and sequencing protocol for keratinised materials of animals

Zhou, Y., Gao, P., Cui, L., Liu, B., Su, C., Zhang, Q., Ma, Y., Lan, T., Yang, S., & Xu, Y. (2026) · Forensic Science International Genetics · doi:10.1016/j.fsigen.2026.103437 · Cited by 1 · see citing works

Ultrastructural Characteristics of Keratinization and Secretion in the Epidermis of Some Antarctic Fishes in Relation to Their Cold Adaptation

Alibardi, L., & Motta, C. M. (2026) · Acta Zoologica · doi:10.1111/azo.70028 · Cited by 1 · see citing works

Development and structure of the skin in the Australian lungfish (Neoceratodus forsteri) in relation to epidermal adaptation of tetrapods

Alibardi, L. (2023) · Acta Zoologica · doi:10.1111/azo.12462 · Cited by 1 · see citing works

Introduction to the Development of Skin in Vertebrates

Alibardi, L. (2023) · Journal of Developmental Biology · doi:10.3390/jdb11010007 · Cited by 1 · see citing works

Peer Review #3 of "Analysis and comparison of protein secondary structures in the rachis of avian flight feathers (v0.1)"

(2022) · PeerJ · doi:10.7287/peerj.12919v0.1/reviews/3

Peer Review #1 of "Analysis and comparison of protein secondary structures in the rachis of avian flight feathers (v0.1)"

(2022) · PeerJ · doi:10.7287/peerj.12919v0.1/reviews/1

Peer Review #2 of "Analysis and comparison of protein secondary structures in the rachis of avian flight feathers (v0.2)"

(2022) · PeerJ · doi:10.7287/peerj.12919v0.2/reviews/2

Peer Review #2 of "Analysis and comparison of protein secondary structures in the rachis of avian flight feathers (v0.1)"

(2022) · PeerJ · doi:10.7287/peerj.12919v0.1/reviews/2

Peer Review #1 of "Analysis and comparison of protein secondary structures in the rachis of avian flight feathers (v0.2)"

(2022) · PeerJ · doi:10.7287/peerj.12919v0.2/reviews/1

Spatio-temporal expression patterns of glycine-rich beta proteins and cysteine-rich beta proteins in setae development of Gekko japonicus

Xia, L., Li, C., Chen, S., Lyu, L., Xie, W., Yan, J., Zhou, K., & Li, P. (2024) · BMC Genomics · doi:10.1186/s12864-024-10426-8

Papillary and Callous Scales in the Integument of Agamid Lizards (Agamidae, Sauria) as a Phenomenon of Extraordinary Development of the Corneous Layers

Dujsebayeva, T. N., Ananjeva, N. B., Rastegar‐Pouyani, N., Al-Johany, A. M. H., & Melnikov, D. A. (2025) · Animals · doi:10.3390/ani15050743

Additional file 7 of Spatio-temporal expression patterns of glycine-rich beta proteins and cysteine-rich beta proteins in setae development of Gekko japonicus

Xia, L., Li, C., Chen, S., Lyu, L., Xie, W., Yan, J., Zhou, K., & Li, P. (2024) · Figshare · doi:10.6084/m9.figshare.26727943

Additional file 9 of Spatio-temporal expression patterns of glycine-rich beta proteins and cysteine-rich beta proteins in setae development of Gekko japonicus

Xia, L., Li, C., Chen, S., Lyu, L., Xie, W., Yan, J., Zhou, K., & Li, P. (2024) · Figshare · doi:10.6084/m9.figshare.26727949

Additional file 2 of Spatio-temporal expression patterns of glycine-rich beta proteins and cysteine-rich beta proteins in setae development of Gekko japonicus

Xia, L., Li, C., Chen, S., Lyu, L., Xie, W., Yan, J., Zhou, K., & Li, P. (2024) · Figshare · doi:10.6084/m9.figshare.25940277

Additional file 3 of Spatio-temporal expression patterns of glycine-rich beta proteins and cysteine-rich beta proteins in setae development of Gekko japonicus

Xia, L., Li, C., Chen, S., Lyu, L., Xie, W., Yan, J., Zhou, K., & Li, P. (2024) · Figshare · doi:10.6084/m9.figshare.26727928

Additional file 4 of Spatio-temporal expression patterns of glycine-rich beta proteins and cysteine-rich beta proteins in setae development of Gekko japonicus

Xia, L., Li, C., Chen, S., Lyu, L., Xie, W., Yan, J., Zhou, K., & Li, P. (2024) · Figshare · doi:10.6084/m9.figshare.26727934

Additional file 6 of Spatio-temporal expression patterns of glycine-rich beta proteins and cysteine-rich beta proteins in setae development of Gekko japonicus

Xia, L., Li, C., Chen, S., Lyu, L., Xie, W., Yan, J., Zhou, K., & Li, P. (2024) · Figshare · doi:10.6084/m9.figshare.26727940

Additional file 5 of Spatio-temporal expression patterns of glycine-rich beta proteins and cysteine-rich beta proteins in setae development of Gekko japonicus

Xia, L., Li, C., Chen, S., Lyu, L., Xie, W., Yan, J., Zhou, K., & Li, P. (2024) · Figshare · doi:10.6084/m9.figshare.26727937

Additional file 1 of Spatio-temporal expression patterns of glycine-rich beta proteins and cysteine-rich beta proteins in setae development of Gekko japonicus

Xia, L., Li, C., Chen, S., Lyu, L., Xie, W., Yan, J., Zhou, K., & Li, P. (2024) · Figshare · doi:10.6084/m9.figshare.25940098

Additional file 8 of Spatio-temporal expression patterns of glycine-rich beta proteins and cysteine-rich beta proteins in setae development of Gekko japonicus

Xia, L., Li, C., Chen, S., Lyu, L., Xie, W., Yan, J., Zhou, K., & Li, P. (2024) · Figshare · doi:10.6084/m9.figshare.26727946

The Epithelial Egg Tooth of the Chicken Shares Protein Markers with the Embryonic Subperiderm and Feathers

Sachslehner, A. P., Steinbinder, J., Hess, C., Mlitz, V., & Eckhart, L. (2025) · Journal of Developmental Biology · doi:10.3390/jdb14010001

The developing alligator tongue undergoes a soft form of cornification associated with intermediate filament keratins

Alibardi, L. (2026) · Journal of Anatomy · doi:10.1111/joa.70117

Review. Cell Biology and Immunohistochemical Evidence of the Transition Between an Aquatic to a Terrestrial‐Waterproof Epidermis in the Archosaurian Alligator

Alibardi, L. (2026) · Journal of Experimental Zoology Part B Molecular and Developmental Evolution · doi:10.1002/jez.b.70017

Sauropsids Cornification is Based on Corneous Beta‐Proteins, a Special Type of Keratin‐Associated Corneous Proteins of the Epidermis

Alibardi, L. (2016) · Journal of Experimental Zoology Part B Molecular and Developmental Evolution · doi:10.1002/jez.b.22689 · Cited by 30 · see citing works

Keratinization and Cornification are not equivalent processes but keratinization in fish and amphibians evolved into cornification in terrestrial vertebrates

Alibardi, L. (2022) · Experimental Dermatology · doi:10.1111/exd.14525 · Cited by 16 · see citing works

Evolutionary origin and diversification of epidermal barrier proteins in amniotes

Strasser, B., Mlitz, V., Hermann, M., Rice, R. H., Eigenheer, R. A., Alibardi, L., Tschachler, E., & Eckhart, L. (2014) · Molecular Biology and Evolution · doi:10.1093/molbev/msu251 · Cited by 135 · see citing works

General aspects on skin development in vertebrates with emphasis on sauropsids epidermis

Alibardi, L. (2023) · Developmental Biology · doi:10.1016/j.ydbio.2023.05.007 · Cited by 12 · see citing works

Cornification in reptilian epidermis occurs through the deposition of keratin‐associated beta‐proteins (beta‐keratins) onto a scaffold of intermediate filament keratins

Alibardi, L. (2012) · Journal of Morphology · doi:10.1002/jmor.20086 · Cited by 37 · see citing works