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Department of Evolutionary MorphologyThe History of the DepartmentComparative anatomical studies have been conducted in the Academy of Sciences of Ukraine since 1922. They were initiated at the Chair of Experimental Zoology and at the Omelchenko Biological Institute. Since 1930 these studies have been conducted at the Departments of Experimental and Comparative Morphology of the newly established Institute of Zoology and Biology (renamed as Institute of Zoology of Academy of Sciences of Ukrainian SSR in 1939, now Schmalhausen Institute of Zoology of National Academy of Sciences of Ukraine). The studies were initiated and led by I. I. Schmalhausen (the first Director of the Institute and Head of the Department of Experimental Morphology) until 1941. 
I. I. Schmalhausen
Since Schmalhausen's laboratory was established it was working in three directions: studies of determination and differentiation of body parts (B. I. Balinsky and N. I. Dragomirov); identification of morphogenic factors and role of proportionate and disproportionate growth in determitation of definitive form (I. I. Schmalhausen, N. P. Bordzilowskaja, K. I. Syngajewskaja,V. V. Brunst) and genetics of the growth factors in animals (G. I. Shpet, I. I. Nazarenko, P. O. Sitko, I. M. Krayovyi). 
The Department of Experimental
Morphology (1935)
First two groups of scientists worked in 1930-s at the Department of Experimental Morphology (later the Department of Developmental Mechanics). In 1934, the genetics group was transformed into the separate Department of General Biology. I. I. Agol became the first head of this Department. However, in 1936 he was arrested by false accusation, becoming a victim of Stalinist repression and was executed. Later (in 1937) this subdivision, which was renamed as Department of Genetics, was headed by S. M. Gershenson. Under the guidance of S. M. Gershenson this Department became one of the main centers of genetic studies in the Soviet Union. Studying the development of limb and eye Balinsky and Dragomirov demonstrated that the inductors act in non-specific manner, and showed that the nature of the structures, formed as a result of embryonic induction, is primarily determined by the nature of reactive tissue. Schmalhausen and his colleagues investigated growth processes across a diverse range of organisms. Their research extensively employed experimental approaches alongside mathematical methods and modeling. Schmalhausen interpreted the changing proportions of body parts during growth as a consequence of ontogenetic restructuring, driven by numerous interactions between organs and structures. Furthermore, he recognized the profound evolutionary significance of these processes. 
The Department of Experimental
Morphology (1935)
Based on these findings, throughout the 1930s and 1940s, Schmalhausen developed his theory of organism’s integrity in both ontogeny and phylogeny. He argued that the formation and maintenance of an organism's integrity are ensured by natural selection—specifically through its stabilizing form. Prior to Schmalhausen, stabilizing selection had been viewed as a purely conservative evolutionary force. This theory became a cornerstone of Modern Synthesis in evolutionary biology. During 1930s the Department of Comparative Morphology (headed by N. N. Voskoboinikov) was involved in morphofunctional and evolutionary studies of fish visceral apparatus. In his studies of the development of selachid branchial apparatus, Voskoboinikov demonstrated that branchial arches are formed from mesenchyma of visceral leaf of mesoderm lateral plate. He suggested that phylogenetic complication in branchial apparatus is connected with evolution of alimentary and respiratory functions of the head gut. 
N. N. Voskoboinikov
By the 1930s, Schmalhausen had become the recognized leader of Soviet evolutionary morphologists and embryologists, directly overseeing research in these fields in both Moscow and Kyiv. In 1936, he was appointed director of the Institute of Evolutionary Morphology of the USSR Academy of Sciences in Moscow, and in 1939, he became the chair of the Department of Darwinism at Moscow University. Although he resided permanently in Moscow from the late 1930s, he maintained general leadership of the Institute of Zoology in Kyiv until 1941. During this period, Balinsky carried out much of the essential administrative leadership of the Institute (as deputy director until 1938) and the department. The Institute of Zoology and its morphological divisions suffered irreparable damage due to totalitarian repressions and the World War II. Among the staff of the Department of Developmental Mechanics, Andronikashvili and Syngajewskaja fell victim to the purges. Syngajewskaja spent two years in the Kargopol concentration camp; fortunately, she was able to return to Kyiv and resume her work at the Institute. During the German occupation, the Institute of Zoology was plundered by the Nazis, who looted research collections, libraries, and laboratory equipment. Further losses followed: Voskoboinikov died in evacuation in Kzyl-Orda, and Syngajewskaja passed away in occupied Kyiv. Balinsky emigrated to Western Europe, later continuing his distinguished career in embryological and zoological research at the University of the Witwatersrand in Johannesburg, South Africa. In the postwar years, the Department of Developmental Mechanics was renamed into the Department of Developmental Biology, headed by B. G. Novikov. The research focused on the experimental study of the ontogeny of breed-specific traits in birds and, specifically, on determining the effects of photoperiodism on development. Schmalhausen remained a staff member of the Institute of Zoology, maintaining close ties with his Kyiv colleagues while continuing his research in evolutionary morphology, particularly the further refinement of his theory of stabilizing selection. Additionally, during the early postwar period, D. K. Tretyakov (Director of the Institute from 1944 to 1948). conducted comparative ichthyomorphological studies. The official endorsement of Lysenkoist views as the sole correct doctrine by the Stalinist regime and the subsequent ban on any criticism (1948) led to the near-total cessation of research in genetics and related biological fields in the Soviet Union. This primarily affected the Modern Synthesis studies, since this theory is based on the synthesis of classical Darwinism and Mendelian genetics. Schmalhausen, who stood at the forefront of the anti-Lysenkoist resistance, was branded a leader of 'Morganism' and dismissed from all his posts; research conducted under his supervision or in pursuit of his ideas was banned. At the Institute of Zoology of the Academy of Sciences of the Ukrainian SSR, the Department of Genetics was liquidated, and research topics concerning the integrating role of natural selection were canceled. Schmalhausen was dismissed from the Institute and never returned to work in Kyiv. Tretyakov was removed from his positions as Institute Director and Chair of the Department of Biological Sciences of the AS UkrSSR. Some staff members of the liquidated genetics department were fired, while others were forced to change their research focus. It would take decades to restore genetic and evolutionary research in the post-Stalin era. In 1950, the morphologists of the Institute of Zoology were consolidated into a single unit, initially named the Department of Comparative Morphology (renamed the Department of Evolutionary Morphology in 1960, and the Department of Evolutionary Morphology of Vertebrates from 1984 to 2016). It was headed by V. G. Kasyanenko (the Director of the Institute in 1950-1963).
N. P. Bordzilowskaja, who remained the only staff member to have worked alongside Schmalhausen, served as a vital link between generations and a witness to all the transformations of the department’s research directions and names. Nadia Petrivna was fluent in several foreign languages and, until the end of her life, generously assisted the Institute’s postgraduate students in translating scientific literature. She passed away in 1996. Many scientists across generations remember her with profound gratitude as a highly educated, compassionate, and kind-hearted person. 
N. P. Bordzilowskaja with
the colleagues The direction of research introduced in the Department by Kasyanenko was the comparative anatomical and functional analysis of tetrapod locomotory apparatus. In 1950 – 1963, V. G. Kasyanenko, S. F. Manziy, P. M. Mazhuga, V. I. Tabin, V. M. Samosh, R. G. Radilovska conducted the fundamental comparative studies of joints in extremities of terrestrial vertebrates; P. M. Mazhuga, Yu. P. Antipchuk, O. R. Radzievsky, E. G. Chernyayev and others studied blood vessels in extremities and other organs. At that time, the first degrees of Candidate of Sciences (= PhD) and Doctor of Sciences were defended. The Department raised and became a school for Ukrainian comparative morphologists. In 1962, Kasyanenko started to lose eyesight and shortly had to stop his researches, but remained a scientific consultant of the Department. In 1963, S. F. Manziy became Head of the Department. In addition to comparative studies, the biomechanical studies were initiated, and bionic direction was declared, but soon discontinued because of enormous funding required. However, other new projects were well funded, and new, mainly young collaborators (engineers, post-graduate students) were involved. The staff reached maximum number; it included almost 30 employees at that time.
In 1965, G. B. Agarkov started his researches on nervous system in the Department. His post-graduate students studied innervation of joints (M. M. Ilyenko, 1965), innervation of extremities muscles (M. F. Kovtun, 1966), and cranial nerves (B. G. Khomenko, 1969).
K. P. Melnyk and V. A. Klykova analyzed the skeleton of extremities, S. F. Manziy, O. G. Berezkin, V. I. Klykov and others studied the joints; muscles of extremities and trunk were studied by S. F. Manziy, V. S. Kotok, V. F. Moroz and M. F. Kovtun; the synovial fluid in various mammals was sudied by A. N. Shchegolkov and O. G. Berezkin. As the result, the biological substitute of synovium for medical treatment of osteoarthroses was created. Electromyography, podography and mechanography of joints were used for the first time. The set of instruments used for these studies was designed at the Department by V. P. Tulyupa, V. S. Kotok and V. F. Moroz. Concurrently blood vessels (L. P. Osinsky) and spine (O. Ya. Pylypchuk) were studied. During these years, new scientific units of the Institute were established on the basis of the Department of Evolutionary Morphology. These included the Department of Cytology and Histogenesis, which operated as an independent division of the Institute from 1963 to 2016 (headed by P. M. Mazhuga, Director of the Institute in 1963–1965, and N. V. Rodionova), and the Department of Functional Morphology of Aquatic Animals, which existed as a separate unit from 1969 to 1991 (headed by G. B. Agarkov, and A. P. Manger).
In early 1970s, M. F. Kovtun initiated ecomorphological studies. Morphological characters were compared with biological features of the studied animals, particularly with their locomotion. Primarily the bats were studied. The hypotheses of bat origin and evolution were proposed. A lack of homology between the bat patagium and the patagium of gliding mammals was found; monophyly of bats was supported.
In 1984, M. F. Kovtun became Head of the Department. Functional and ecomorphological studies continued, and embryological studies were started. In 1984, the Department organized the All-Union Conference on Biomechanics, and in 1985 — the All-Union Conference on Bat Studies. 
The experimental station of the Institute
of Zoology in the Feofaniia tract (southern outskirts of Kyiv):
The researchers of the Department studied functional morphology and ecomorphology of bats (M. F. Kovtun, I. M. Kovalyova, R. Y. Lykhotop, N. F. Zhukova, S. Y. Ledenev, Ya. A. Omelkovets, I. I. Dzeverin), gallinaceous birds (V. F. Sych, I. A. Bogdanovich) and flycatchers (O. M. Peklo). At the same time, functional-morphological and biomechanical studies of ungulates and some other animals were continued (O. G. Berezkin, K. P. Melnik, V. I. Klykov, V. A. Klykova, V. F. Moroz, V. S. Kotok, L. P. Osinsky). The concept of the origin of bats was being developed (M. F. Kovtun, I. M. Kovalyova, R. Y. Lykhotop, N. F. Zhukova, S. Y. Ledenev). Some new concepts concerning the origin of land locomotion apparatus of birds (I. A. Bogdanovich) and the origin of birds (I. A. Bogdanovich in coauthorship with E. N. Kurochkin from Paleontological Institute, Russian Academy of Sciences). Multivariate statistical techniques were applied in the studies of bat variation and evolution (I. I. Dzeverin). The researches of the Department studied the embryogeny of skull and extremities skeleton in bats (M. F. Kovtun, R. Y. Lykhotop, S. Y. Ledenev), and later in corvid birds (M. F. Kovtun, Y. V. Shatkovsky, O. V. Shatkovska). 
Researchers from the Department
of Evolutionary Morphology of Vertebrates and their colleagues
(December 1993)
Gradually, comparative anatomical, ecomorphological and comparative embryological studies covered most vertebrate taxa and systems of organs, including digestive (N. F. Zhukova), respiratory (I. M. Kovalyova), locomotor (I. A. Bogdanovich, I. M. Kovalyova, S. Y. Ledenev), cardiovascular (L. P. Osinsky), reproductive and excretory (O. V. Nechayeva, S. O. Ghilevych) systems, skin (O. P. Koval), termoregulatory organs (I. P. Zakrevskaya). The researches of the Department studied the skull embryogenesis in some reptiles (O. M. Yaryhin, H. V. Sheverdyukova). In the framework of cooperation between the Institute of Zoology and Lesya Ukrainka Volyn National University, a study on the embryology of Osteichthyes was started (Y. V. Stepanyuk and M. F. Kovtun). 
Researchers from the Department
of Evolutionary Morphology of Vertebrates (June 2005)
In 2013, I. I. Dzeverin took over the leadership of the department. The primary research areas now encompass morphometrics and the integration of statistical methods into comparative morphological and ecological data (I. I. Dzeverin, P. E. Gol’din, M. A. Ghazali, A. V. Romaniuk, S. V. Davydenko, O. O. Vertsimakha). Other key directions include multifaceted studies of marine mammal biology (P. E. Gol’din, K. O. Vishnyakova, O. V. Gladilina, O. V. Savenko, S. V. Davydenko, J. F. Ivanchikova, V. S. Telizhenko, P. A. Otriazhyi), as well as research in paleontology (L. V. Popova, P. E. Gol’din, S. V. Davydenko, P. A. Otriazhyi), taxonomy and phylogenetics (P. E. Gol’din, L. V. Popova, M. A. Ghazali, I. I. Dzeverin, S. V. Davydenko, V. S. Telizhenko, P. A. Otriazhyi). Furthermore, the department focuses on ecology and the scientific basis for biodiversity conservation (P. E. Gol’din, J. F. Ivanchikova, K. O. Vishnyakova, O. V. Gladilina, M. A. Ghazali). At the same time, the department has maintained its traditional focus on comparative and functional morphology (I. O. Bogdanovich, P. E. Gol’din, O. V. Shatkovska, I. M. Kovalyova, S. V. Davydenko, P. A. Otriazhyi) and embryology (M. F. Kovtun, O. M. Yaryhin, H. V. Sheverdyukova). In 2016, the former Department of Cytology and Histogenesis was merged into the Department of Evolutionary Morphology. Histological research previously conducted there (N. M. Akulenko, N. V. Zolotova-Haidamaka) has since continued within the Department of Evolutionary Morphology. Collaboration in the field of evolutionary embryology also continues with specialists from the Lesya Ukrainka Volyn National University (Y. V. Stepanyuk, O. V. Kostielova). The department’s researchers have investigated the evolutionary pathways and patterns of cetaceans (P. E. Gol’din, S. V. Davydenko, V. S. Telizhenko, P. A. Otriazhyi), bats (I. I. Dzeverin, M. A. Ghazali), rodents (L. V. Popova, I. I. Dzeverin), carnivorans (P. A. Otriazhyi, P. E. Gol’din, A. V. Romaniuk), and various avian groups (I. O. Bogdanovich, O. V. Shatkovska, M. A. Ghazali). Their theoretical work focuses on the mechanisms of evolution (I. I. Dzeverin, M. A. Ghazali, V. S. Telizhenko, P. E. Gol’din, S. V. Davydenko), organismal modularity and integration (I. I. Dzeverin, M. A. Ghazali, P. E. Gol’din, V. S. Telizhenko, S. V. Davydenko, O. O. Vertsimakha), and the role of heterochrony in evolution (P. E. Gol’din, O. M. Yaryhin). The staff has been instrumental in introducing cutting-edge scientific tools to Ukraine, including geometric morphometrics (P. E. Gol’din, M. A. Ghazali, L. V. Popova, A. V. Romaniuk, S. V. Davydenko), molecular phylogenetics (M. A. Ghazali, V. S. Telizhenko), phylogenetic comparative methods (M. A. Ghazali, I. I. Dzeverin), evolutionary quantitative genetics (I. I. Dzeverin, M. A. Ghazali), paleohistology (S. V. Davydenko), bioinformatics (V. S. Telizhenko), bioacoustics (J. F. Ivanchikova), and mathematical methods for analyzing the structure of complex systems (O. O. Vertsimakha). The department’s researchers have carried out numerous scientific projects, including two major grants supported by the National Research Foundation of Ukraine: 'Integration of mammalian organism as a proxy of stability at aquatic and aerial life (as illustrated by skeleton traits)' (2020–2023, team leader: P. E. Gol’din) and Pathways of evolution of mammalian and avian body size: trajectories of adaptation and breaking constraints' (launched in 2026, team leader: P. E. Gol’din). Additionally, the department hosts the project Establishing the bank of cetacean tissue samples in Ukraine,' commissioned by the Secretariat of the ACCOBAMS Agreement (Monaco) (team leader: P. E. Gol’din). The department’s staff have taken an active role in implementing the Institute of Zoology's educational and scientific program. From 2016 to 2020, the Department functioned as a Graduating Unit; later, this responsibility was transferred to the Research and Education Training Unit, which continues to involve department staff (the group is headed by I. I. Dzeverin, who also serves as the Educational and scientific program Coordinator). Researchers from the department have developed and currently teach specialized courses for postgraduate students (I. I. Dzeverin, M. A. Ghazali, P. E. Gol’din, L. V. Popova, S. V. Davydenko) and provide organizational and methodological support for the teaching and thesis defense processes (I. I. Dzeverin, L. V. Popova, N. V. Zolotova-Haidamaka, M. A. Ghazali). The Department also hosts the preparation of doctoral dissertations, as well as master’s and bachelor’s theses. In its activities, the Department of Evolutionary Morphology strives to bridge classical approaches with modern methodologies, maintaining a cutting-edge level of research. While upholding the traditions of evolutionary and ontogenetic research established by Schmalhausen, the department's staff fully integrate the theoretical and methodological innovations of recent decades and adhere to contemporary global scientific standards. Sustaining this level of scientific excellence has taken on particular significance for the department's staff, as it has for all Ukrainian scientists, amidst the challenges of the Russian-Ukrainian war. |
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I. I. Schmalhausen Institute of Zoology, 2004-2026 |
