
Daniel H. Shain
Professor of Zoology
Biology Department
Curriculum Vitae (CV)
Education
- University of New Hampshire, BS/MS (1988)
- Colorado State University, Ph.D. (1994)
- University of California, Berkelely, Post-doc (1995-1998)
Research interests: We are interested in all aspects of worm biology, from the bioenergetic adaptation of glacier ice worms to the secretion of leech cocoons and production of embryonic stem cells. We also have ongoing projects exploring the evolution and adaptation of extremophilic animals (e.g., Rotifera, Tardigrada) that occupy glaciers and hot springs worldwide, including from Iceland, New Zealand, Norway and Tibet.
Selected Publications
(*indicates Rutgers student)
2026
*Hsiao S, Saglam N, Saunders R, Shain DH (2026) Gene Expression Underlying Teloblast Formation in Leech (Annelida: Clitellata). Evolutionary Bioinformatics (in press).
Hill S, Saglam N and Shain D (2026) Reproduction in the Annelida. In: Skinner, Michael K. (ed.) Encyclopedia of Reproduction, 3e, vol. 6, pp. 683–691. US: Elsevier. http://dx.doi.org/10.1016/B978-0-443-21477-6.00114-0
Simoncini A, Cantera I, Giachello S, Valle B, Gobbi M, Fontaneto D, Zawierucha K, Laniecki R, Coulson S, Lencioni V, Cauvy-Fraunié S, Moràn-Ordóñez A, Losapio G, Takeuchi N, Liu Q, Shain DH, Fair H, Janko K, Devetter M, Jiménes-Santos MA, Šabacká M, Falaschi M, Marta S, Ficetola GF (2026) The Global Diversity and Decline of Glacier Animals. PNAS (in revision).
Zawierucha K, Trzebny A, Lukashanets DA, Laniecki R, Kataržyte M, Kalvaitienė G, Valle B, Kowalik J, Stachowiak M, Shain DH, Ambrosini R, Ficetola F, Ono M, Buda J (2026) Microinvertebrates and Associated Algae on Surface Snow (in review).
Avan Mutlu T, *Dunkley, T, Klein EA, Shain DH. Bioengineering the Ice Worm ATP Synthase to Increase Energy Production (in review).
*Sharma R, Shain DH. Reduced CAG Repeats in the Androgen Receptor Gene Can Lead to Polycystic Ovarian Syndrome in Patients with Normal Androgen Levels (in review).
2025
Dunkley T, Shain DH, Klein EA (2025) A histidine-rich extension of F0 subunit ATP6 from the ice worm Mesenchytraeus solifugus increases ATP synthase activity in bacteria. FEBS Lett. doi:10.1002/1873-3468.15100
Saglam N, Saunders R, Shain DH (2025) Species Delimitation Recovers Erpobdellidae (Annelida: Hirundinae) from USA and Türkiye. Zootaxa 5627, 455–479. https://www.mapress.com/zt/
*DiMattia L, Saglam N, Saunders R, Shain DH (2025) An Ice-Inhabiting Bdelloid Rotifer from North America. Extremophiles 29, 30. doi.org/10.1007/s00792-025-01390-6
2024
Shain DH, Irina Rogozhina, Diego Fontaneto, Naim Saglam, Atle Nesje, Jesamine Bartlett, Krzysztof Zawierucha, Einar Arnason, Jørgen Rosvold (2024) Ice-inhabiting species of Bdelloidea Rotifera reveal a pre-Quaternary ancestry in the Arctic cryosphere. Biol. Lett. 20: 20230546. https://doi.org/10.1098/rsbl.2023.0546.
*Hsiao S, Saglam N, *Morrow D, Weisblat DA, Shain DH (2024) Transcriptomic profiling at the maternal-to-zygotic transition in leech, Helobdella austinensis. Genes 15, 283. https://doi.org/ 10.3390/genes15030283
Janko K, Shain DH, Fontaneto D, Kaštánková M, Šabacká M, Rosvold J, Stefaniak J, Hessen DO, Devetter M, Horna P, Drdová EJ, Buda J, Yde JC, Zawierucha K (2024) Islands of Ice: Glacier-Dwelling Metazoans Form Regionally-Distinct Populations Despite Extensive Periods of Deglaciation. Diversity and Distributions 30:e13859. doi.org/10.1111/ddi.13859
Shain DH, Klein EA (2024) Genome assembly of Pseudomonas olympii sp. nov., a psychrotolerant bacterium isolated from Deception Glacier (Washington, USA). Microbiol. Res. Ann. 13, 1-4. doi.org/10.1128/mra.00125-24
2023
Zawierucha K, Stec D, Dearden PK, Shain DH (2023) Two New Tardigrade Genera from New Zealand’s Southern Alp Glaciers Display Morphological Stasis and Parallel Evolution. Molecular Phylogenetic & Evolution 178, 107634.
Saglam N, *Melissaratos DA, Shain DH (2023) Biocontrol of Snail-Borne Parasites with the Glossiphoniid Leech, Helobdella austinensis. Biol. Lett. 19, 20220484. https://doi.org/10.1098/rsbl.2022.0484
2022
Shain DH, Iakovenko N, Novis P, Cridge A, Plášek V, Dearden PK (2022) Microinvertebrate colonization of New Zealand’s thermally extreme environments. Evol. Biol. 49, 414-423
*Osorio T, Shain DH†, *Scoma ER†, *Melissaratos DS, Riggs LM, Hambardikar V, Solesio ME (2022) The Glacier Ice Worm, Mesenchytraeus solifugus, Elevates Mitochondrial Inorganic Polyphosphate (PolyP) Levels in Response to Stress. Biology 11, 1771.
2021
Shain DH, Novis P, Cridge A, Zawierucha K, Dearden PK (2021) Five animal phyla in glacier ice reveal unprecedented biodiversity in New Zealand’s Southern Alps. Scientific Reports 11, 3898 https://doi.org/10.1038/s41598-021-83256-3
Shmakova L, Malavin S, Iakovenko N, Vishnivetskaya T, Shain D, Plewka M, Rivkina E (2021) A living bdelloid rotifer from 24,000-year-old Arctic permafrost. Current Biology, 31, R712-13. https://doi.org/10.1016/j.cub.2021.04.077
Zawierucha K, Uetakeb J and Shain DH (2021) A Compendium of Tropical Ice Masses. The Encyclopedia of Conservation, Reference Module in Earth Systems and Environmental Sciences. Elsevier https://doi.org/10.1016/B978-0-12-821139-7.00033-7
2020
*Lang SA, McIlroy PJ, Shain DH (2020) Structural evolution of the glacier ice worm F1Fo ATP Synthase complex. The Protein Journal 39, 152-159. doi: 10.1007/s10930-020-09889-x
Gagliardi LJ, Shain DH (2020) Electrostatic contributions to carcinogenesis. Open Journal of Biophysics 10, 27-45.
Elser JJ, Wu C, González AL, Shain DH, Smith HJ, Sommaruga J, Williamson CE, Brahney J, Hotaling S, Vanderwall J, Yu J, Aizen V, Aizen E, Battin TJ, Camassa R, Feng X, Jiang H, Lu L, Qu JJ, Ren Z, Wen J, Wen L, Woods HA, Xiong X, Xu J, Yu G, Saros JE (2020) Rules of life and the fading cryosphere: impacts in alpine lakes and streams. Global Change Biology 26, 6644-6656.
Zawierucha K, Porazinska DL, Ficetola GF, Ambrosini R, Baccolo G, Buda J, Ceballos JL, Devetter M, Dial R, Franzetti A, Fuglewicz U, Gielly L, Łokas E, Janko K, Jaromerska, Kościński A, Kozłowska A, Ono M, Parnikoza I, Pittino F, Poniecka E, Sommers P, Schmidt PK, Shain DH, Sikorska S, Uetake J, Takeuchi N (2020) A hole in the nematosphere: tardigrades and rotifers dominate the cryoconite hole environment, whereas nematodes are missing. Journal of Zoology 313, 18-36.
Saglam N, *Saunders R, Shain DH, Saidel WM (2020) Detailed ultrastructure of the Hirudo salivary gland. Micron 136. doi.org/10.1016/j.micron.2020.102887.
Gagliardi LJ, Shain DH (2020) Biophysical mechanism for zinc as an anticancer agent. Medical Hypotheses 144, 110273.
2019
Zawierucha K and Shain DH (2019) Disappearing Kilimanjaro snow – are we the last generation to explore equatorial glacier biodiversity? Ecology and Evolution 9, 8911–8918. DOI:10.1002/ece3.5327
Kutschera U and Shain DH (2019) Hirudinea Lamarck 1818: Evolutionary origin and taxonomy of the six medicinal leeches (genus Hirudo) known today. Biomedical Research and Reviews 3, 1-4.
Hotaling S, Shain DH, *Lang SA, Bagley RK, Tronstad LM, Weisrock DW and Kelley JL (2019) Long-distance dispersal, ice sheet dynamics, and mountaintop isolation underlie the genetic structure of glacier ice worms. Proc. Roy. Soc. B 286, 20190983. http://dx.doi.org/10.1098/rspb.2019.0983
Gagliardi LJ, Shain DH (2019) Electrostatic mechanism for depolymerization-based poleward force generation at kinetochores. Open Journal of Biophysics 9, 198-203.
2018
Saidel WM, Saglam N, Salas-de la Cruz D, *Saunders R, Shain DH (2018) Elaborate Ultrastructure of the Hirudo (Annelida: Hirudinae) Cocoon Surface. J Morphology 279, 545-553.
Saglam N, *Saunders R, *Lang SA, Shain DH (2018) Phylogeny and cocoon production in the parasitic leech Myzobdella lugubris Leidy, 1851 (Hirudinidae, Piscicolidae). Acta Parasitologica 63, 15-26.
Gagliardi LJ, Shain DH (2018) Emergent mitotic chromosome motions from a changing intracellular pH. Open Journal of Biophysics 8, 1-13.
Saglam N, Kutschera U, *Saunders R, Saidel WM, *Balombini KLW, Shain DH (2018) Morphologic and phylogenetic resolution of the Helobdella stagnalis complex (Annelida: Hirudinidae). Zootaxa 4403(1) 061-086. https://doi.org/10.11646/zootaxa.4403.1.3
- Saglam N, Kutschera U, Saunders R, Saidel WM,Balombini KLW and Shain DH (2019) Correct spelling of a recently described species of Helobdella (Annelida:Clitellata: Hirudinea). Zootaxa 4571, 277.
- Saglam N andShain DH(2025) Correction of the specific epithet in Helobdella serendipitous Saglam et al. 2018 (Annelida: Hirudinea) as Helobdella serendipitosa, in accordance with Latin grammar. Zootaxa
*Lang SA, Shain DH (2018) Atypical Evolution of the F1F0 ATP Synthase Regulatory ATP6subunit in Glacier Ice Worms (Annelida: Clitellata: Mesenchytraeus). Evolutionary Bioinformatics 14, 1-4. doi: 10.1177/1176934318788076
Hill SD, Saglam N and Shain DH (2018) Reproduction in the Annelida. In M. K. Skinner (Ed.), Encyclopedia of Reproduction. Vol. 6, pp. 526–532. Academic Press: Elsevier. http://dx.doi.org/10.1016/B978-0-12-809633-8.20648-1. ISBN: 9780128118993
2017
*Lang SA, Saglam N, *Kawash J, Shain DH (2017) Punctuated invasion of water, ice, snow and terrestrial ecozones by segmented worms (Oligochaeta: Enchytraeidae: Mesenchytraeus). Proc. Roy. Soc. B 284, 20171081. http://dx.doi.org/10.1098/rspb.2017.1081
2016
Kotchoni SO, Gachomo EW, *Slobodenko K, Shain DH (2016) AMP DEAMINASE suppression increases biomass, cold tolerance and oil content in green algae. Algal Research 16, 473-480.
Saltmarsh DM, Bowser ML, Morton JM, *Lang SA, Shain DH, Dial RJ (2016) Distribution and abundance of exotic earthworms within a boreal forest system in southcentral Alaska. NeoBiota 28, 67-86.
Dial RJ, Becker M, Hope AG, Dial CR, Thomas J, *Slobodenko K, Golden TS, Shain DH (2016) The role of temperature in the distribution of the glacier ice worm, Mesenchytraeus solifugus (Annelida: Oligochaeta: Enchytraeidae). Arctic, Antarctic, and Alpine Res. 48, 199-211.
Shain DH, Halldórsdóttir K, Pálsson F, Aðalgeirsdóttir G, Gunnarsson A, Jónsson T, *Lang SA, Pálsson HS, Steinþórssson S, Arnason E (2016) Colonization of Maritime Glacier Ice by Bdelloid Rotifera. Mol. Phylog. Evol. 98, 280–287.
Saglam N, *Saunders R, *Lang S, Shain DH (2016) A new species of Hirudo (Annelida: Hirudinidae): biogeographical history of Eurasian medicinal leeches. BMC Zoology 1:5, DOI 10.1186/s40850-016-0002
Gagliardi LJ, Shain DH (2016) Electrostatic forces drive poleward chromosome motions at kinetochore. Cell Division 11, 1-8.
*Rossi A, Saidel WM, *Gravante C, Sayers CW, Shain DH (2016) Mechanics of cocoon secretion in a segmented worm (Annelida: Hirudinae). Micron 86, 30-35.
