Monsters in the dark: systematics and biogeography of the stygobitic genus Godzillius (Crustacea: Remipedia) from the Lucayan Archipelago

  • Lauren Ballou Department of Marine Biology, Texas A&M University at Galveston, United States
  • Thomas M. Iliffe Department of Marine Biology, Texas A&M University at Galveston, United States
  • Brian Kakuk Bahamas Caves Research Foundation, Marsh Harbor, Abaco Island, the Bahamas
  • Brett C. Gonzalez Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington DC, United States
  • Karen J. Osborn Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington DC, United States
  • Katrine Worsaae Department of Biology, University of Copenhagen, Denmark
  • Kenneth Meland Department of Biological Sciences, University of Bergen, Norway
  • Kenneth Broad Department of Environmental Science and Policy, Rosenstiel School of Marine and Atmospheric Science & Abess Center, University of Miami, United States
  • Heather Bracken-Grissom Institute of Environment and Department of Biology, Florida International University - Biscayne Bay Campus, United States
  • Jørgen Olesen Natural History Museum of Denmark, University of Copenhagen, Denmark
Keywords: anchialine, cave, new species, phylogeny


Remipedia is a stygobitic group commonly associated with coastal anchialine caves. This class consists of 12 genera, ten of which are found within the Lucayan Archipelago. Herein, we describe a new species within the genus Godzillius from Conch Sound Blue Hole, North Andros Island, Bahamas. Godzillius louriei sp. nov. is the third known remipede observed from a subseafloor marine cave, and the first from the Godzilliidae. Remipedes dwell within notoriously difficult to access cave habitats and thus integrative and comprehensive systematic studies at family or genus level are often absent in the literature. In this study, all species of Godzillius are compared using morphological and molecular approaches. Specifically, the feeding appendages of G. louriei sp. nov., G. fuchsi Gonzalez, Singpiel & Schlagner, 2013 and G. robustus Schram, Yager & Emerson, 1986 were examined using scanning electron microscopy (SEM). Species of Godzillius are identified based on the spines of maxilla 1 segment 4 and by the denticles on the lacinia mobilis of the left mandible. A molecular phylogeny using the mitochondrial 16S rRNA and nuclear histone 3 genes recovered G. louriei sp. nov. within the Godzillius clade and 16S genetic distances revealed a 13–15% difference between species of Godzillius.


Altschul S.F., Gish W., Miller W., Myers E.W. & Lipman D.J. 1990. Basic local alignment search tool. Journal of Molecular Biology 215: 403–410.

Benson D.A., Boguski M.S., Lipman D.J., Ostell J. & Ouellette B.F.F. 1998. GenBank. Nucleic Acids Research 26: 1–7.

Bishop R.E., Humphreys W.F., Cukrov N., Žic V., Boxshall G.A., Cukrov M., Iliffe T.M., Kršinić F., Moore W.S., Pohlman J.W. & Sket B. 2015. ‘Anchialine’ redefined as a subterranean estuary in a crevicular or cavernous geological setting. Journal of Crustacean Biology 35: 511–514.

Brankovits D., Pohlman J.W., Niemann H., Leigh M.B., Leewis M.C., Becker K.W., Iliffe T.M., Alvarez F., Lehmann M.F. & Phillips B. 2017. Methane- and dissolved organic carbon-fueled microbial loop supports a tropical subterranean estuary ecosystem. Nature Communications 8: 1–12.

Cánovas F., Jurado-Rivera J.A., Cerro-Gálvez E., Juan C., Jaume D. & Pons J. 2016. DNA barcodes, cryptic diversity and phylogeography of a W Mediterranean assemblage of thermosbaenacean crustaceans. Zoologica Scripta 45: 659–670.

Colgan D.J., McLauchlan A., Wilson G.D.F., Livingston S.P., Edgecombe G.D., Macaranas J., Cassis G. & Gray M.R. 1998. Histone H3 and U2 snRNA DNA sequences and arthropod molecular evolution. Australian Journal of Zoology 46: 419–437.

Daenekas J., Iliffe T.M., Yager J. & Koenemann S. 2009. Speleonectes kakuki, a new species of Remipedia (Crustacea) from anchialine and sub-seafloor caves on Andros and Cat Island, Bahamas. Zootaxa 2016 (1): 51–66.

Darriba D., Taboada G.L., Doallo R. & Posada D. 2012. jModelTest 2: more models, new heuristics and parallel computing. Nature Methods 9: e772.

Farr M. 2017. The Darkness Beckons: The History and Development of World Cave Diving. Verte-brate Publishing.

Farr M. & Palmer R. 1984. The blue holes: description and structure. In: Ford T.D. (ed.) Bahamas Blue Holes 1981–1982. Cave Science 11 (1): 9–22.

Gonzalez B.C., Singpiel A. & Schlagner P. 2013. Godzillius fuchsi, a new species of Remipedia (Godzilliidae) from Abaco Island, Bahamas. Journal of Crustacean Biology 33: 275–285.

Gonzalez B.C., Martínez A., Borda E., Iliffe T.M., Fontaneto D. & Worsaae K. 2017. Genetic spatial structure of an anchialine cave annelid indicates connectivity within – but not between – islands of the Great Bahama Bank. Molecular Phylogenetics and Evolution 109: 259–270.

Guindon S. & Gascuel O. 2003. A simple, fast, and accurate method to estimate large phylogenies by maximum likelihood. Systematic Biology 52: 696–704.

Hart C.W. Jr, Manning R.B. & Iliffe T.M. 1985. The fauna of Atlantic marine caves: evidence of dispersal by sea floor spreading while maintaining ties to deep waters. Proceedings of the Biological Society of Washington 98: 288–292.

Hoang D.T., Chernomor O., von Haeseler A., Minh B.Q. & Vinh L.S. 2018. UFBoot2: improving the ultrafast bootstrap approximation. Molecular Biology and Evolution 35: 518–522.

Hoenemann M., Neiber M.T., Humphreys W.F., Iliffe T.M., Li D., Schram F.R. & Koenemann S. 2013. Phylogenetic analysis and systematic revision of Remipedia (Nectiopoda) from Bayesian analysis of molecular data. Journal of Crustacean Biology 33: 603–619.

Hunter R.L., Webb M.S., Iliffe T.M. & Alvarado Bremer J.R. 2008. Phylogeny and historical biogeography of the cave-adapted shrimp genus Typhlatya (Atyidae) in the Caribbean Sea and western Atlantic. Journal of Biogeography 35: 65–75.

Juan C., Guzik M.T., Jaume D. & Cooper S.J.B. 2010. Evolution in caves: Darwin’s ‘wrecks of ancient life’ in the molecular era. Molecular Ecology 19: 3865–3880.

Kalyaanamoorthy S., Minh B.Q., Wong T.K.F., von Haeseler A. & Jermiin L.S. 2017. ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods 14: 587–589.

Katoh K., Rozewicki J. & Yamada K.D. 2019. MAFFT online service: multiple sequence alignment, interactive sequence choice and visualization. Briefings in Bioinformatics 20: 1160–1166.

Kearse M., Moir R., Wilson A., Stones-Havas S., Cheung M., Sturrock S., Buxton S., Cooper A., Markowitz S., Duran C., Thierer T., Ashton B., Meintjes P. & Drummond A. 2012. Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28: 1647–1649.

Koenemann S. & Iliffe T.M. 2014. Class Remipedia Yager, 1981. In: Klein V.K. (ed.) Treatise on Zoology-Anatomy, Taxonomy, Biology. The Crustacea. Volume 4, part A: 125–177. Brill, Leiden.

Koenemann S., Iliffe T.M. & Yager J. 2004. Kaloketos pilosus, a new genus and species of Remipedia (Crustacea) from the Turks and Caicos Islands. Zootaxa 618 (1): 1–12.

Koenemann S., Schram F.R., Hönemann M. & Iliffe T.M. 2007. Phylogenetic analysis of Remipedia (Crustacea). Organisms Diversity & Evolution 7: 33–51.

Kumar S., Stecher G. & Tamura K. 2016. MEGA7: Molecular evolutionary genetics analysis version 7.0 for bigger datasets. Molecular Biology and Evolution 33: 1870–1874.

Lefébure T., Douady C.J., Gouy M. & Gibert J. 2006. Relationship between morphological taxonomy and molecular divergence within Crustacea: proposal of a molecular threshold to help species delimitation. Molecular Phylogenetics and Evolution 40: 435–447.

Miller M.A., Pfeiffer W. & Schwartz T. 2010. “Creating the CIPRES Science Gateway for inference of large phylogenetic trees.” In: Proceedings of the Gateway Computing Environments Workshop (GCE): 1–8. IEEE Xplore, New Orleans.

Mylroie J.E. & Carew J.L. 1990. The flank margin model for dissolution cave development in carbonate platforms. Earth Surface Processes and Landforms 15: 413–424.

Mylroie J.E. & Mylroie J.R. 2011. Void development on carbonate coasts: creation of anchialine habitats. Hydrobiologia 677: 15–32.

Natural Earth 2020. Natural Earth. Free vector and raster map data at 1 : 10m, 1 : 50m, and 1 : 100m scales. Available from [accessed 18 May 2021].

Neiber M.T., Hansen F.C., Iliffe T.M., Gonzalez B.C. & Koenemann S. 2012. Molecular taxonomy of Speleonectes fuchscockburni, a new pseudocryptic species of Remipedia (Crustacea) from an anchialine cave system on the Yucatán Peninsula, Quintana Roo, Mexico. Zootaxa 3190 (1): 31–46.

Nguyen L.T., Schmidt H.A., von Haeseler A. & Minh B.Q. 2014. IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution 32: 268–274.

Olesen J., Meland K., Glenner H., van Hengstum P.J. & Iliffe T.M. 2017. Xibalbanus cozumelensis, a new species of Remipedia (Crustacea) from Cozumel, Mexico, and a molecular phylogeny of Xibalbanus on the Yucatán Peninsula. European Journal of Taxonomy 316: 1–27.

QGIS Development Team 2020. QGIS Geographic Information System, ver. 3.12. Open Source Geospatial Foundation. Available from [accessed 19 Mar. 2021].

Pais F.S.M., Ruy P.C., Oliveira G. & Coimbra R.S. 2014. Assessing the efficiency of multiple sequence alignment programs. Algorithms for Molecular Biology 9: 1–8.

Palmer R. 1997. Deep into Blue Holes. Media Publishing, Nassau, Bahamas.

Palumbi S., Martin A., Romano S., McMillan W.O., Stice L. & Grabowski G. 2002. The Simple Fool’s Guide to PCR. Ver. 2. Department of Zoology and Kewalo Marine Laboratory, Honolulu.

Rambaut A., Drummond A.J., Xie D., Baele G. & Suchard M.A. 2018. Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systematic Biology 67: 901–904.

Reid W.V. 1998. Biodiversity hotspots. Trends in Ecology & Evolution 13: 275–280.

Ronquist F., Teslenko M., van der Mark P., Ayres D.L., Darling A., Höhna S., Larget B., Liu L., Suchard M.A. & Huelsenbeck J.P. 2012. MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539–542.

Schram F.R., Yager J. & Emerson M.J. 1986. Remipedia Part 1. Systematics. San Diego Society of Natural History, Memoir 15: 1–60.

Song H., Buhay J.E., Whiting M.F. & Crandall K.A. 2008. Many species in one: DNA barcoding overestimates the number of species when nuclear mitochondrial pseudogenes are coamplified. Proceedings of the National Academy of Sciences 105: 13486–13491.

Surić M., Juračić M., Horvatinčić N. & Bronić I.K. 2005. Late Pleistocene–Holocene sea-level rise and the pattern of coastal karst inundation: records from submerged speleothems along the Eastern Adriatic Coast (Croatia). Marine Geology 214: 163–175.

van Hengstum P.J., Cresswell J.N., Milne G.A. & Iliffe T.M. 2019. Development of anchialine cave habitats and karst subterranean estuaries since the last ice age. Scientific Reports 9: 1–10.

Yager J. 2013. Speleonectes cokei, new species of Remipedia (Crustacea: Speleonectidae) from a submerged ocean cave near Caye Chapel, Belize. Zootaxa 3710 (4): 354–362.

Zhou X., Shen X.X., Hittinger C.T. & Rokas A. 2017. Evaluating fast Maximum Likelihood-based phylogenetic programs using empirical phylogenomic data sets. Molecular Biology and Evolution 35: 486–503.

How to Cite
Ballou, L., Iliffe, T. M., Kakuk, B., Gonzalez, B. C., Osborn, K. J., Worsaae, K., Meland, K., Broad, K., Bracken-Grissom, H., & Olesen, J. (2021). Monsters in the dark: systematics and biogeography of the stygobitic genus Godzillius (Crustacea: Remipedia) from the Lucayan Archipelago. European Journal of Taxonomy, 751(1), 115-139.
Research article