The articulated bryozoan genus Cellaria in the southern Zealandian Region: distribution and associated fauna

  • Katerina AchilleosEmail author
  • Abigail M. Smith
  • Dennis P. Gordon
Original Paper


The Zealandian Region centred on New Zealand has the highest diversity of Cellaria species in the world, with three described and 11 undescribed living species. This study assesses their distribution in relation to environmental parameters and characterizes the associated bryozoan fauna in the Southern Zealandian Region based on specimens collected in the years 1911–2018. Cellaria specimens stored in collections in New Zealand were identified to species level, and their metadata were combined with that in the published literature to provide a database for analysis. Distributional data were mapped and assessed in relation to environmental factors: surface chlorophyll-a, sea-surface and seafloor temperature, and substratum type. The bryozoan fauna co-occurring with Cellaria was recorded and characterized according to community assemblage composition. Of the 14 Cellaria species, C. immersa and C. tenuirostris are the most dominant in the southern Zealandian region. Presence/absence records showed that Cellaria species mostly occur at sites with relatively low productivity (0.10–0.99 mg m−3) and relatively high seafloor temperatures (13–14 °C). Sand was identified as the main textural component of the substratum where Cellaria was found. The ability of Cellaria species to colonize soft sediments is of ecological importance in mid-shelf environments where hard substrata are uncommon. The associated bryozoan fauna comprises common elements of bryozoan assemblages and their reef formations around the world and is characterized by species having an erect-rigid colony growth form. The present results can contribute to future ecological analyses using Cellaria species globally as key elements for other bryozoan faunas.


Environmental parameters Chlorophyll-a Surface and seafloor temperature Substratum 



We thank Ms. Sadie Mills (Collection Manager, NIWA) and Ms. Diana Macpherson (Assistant Collection Manager, NIWA) for the donation and loan of specimens. We also thank Dr. Kim Currie (NIWA/University of Otago Research Centre for Oceanography) and the crew of RV Polaris II for support and assistance during the collection of some specimens. Dr. Steve Chiswell (NIWA, Wellington) gave guidance regarding seafloor temperature data. Mr. Aubrey Miller (School of Surveying, University of Otago) gave support and assistance with the use of ArcGIS. We are also grateful for the comments of three anonymous reviewers that improved the manuscript.

Funding information

Research funding to KA is provided by University of Otago Department of Marine Science.

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

Ethical approval

No animal testing was performed during this study.

Sampling and field studies

All necessary permits for sampling and observational field studies have been obtained by the authors from the competent authorities.

Data accessibility statement

All netCDF files for sea surface temperature and chlorophyll-α are available from Nasa Ocean Colour website ( The netCDF file for seafloor temperature is available from CSIRO Atlas of Regional Seas (CARS) website ( Sediment raster data are available from NZODN ( Bathymetry shapefile is available from NIWA website ( All data points extracted from the raster files, coordinates, stations and species details, and associated community are available as supplementary material.

Supplementary material

12526_2019_1009_MOESM1_ESM.xlsx (339 kb)
Supplementary material 1 All sampling locations from 1911 to 2018 including presence and absence records (XLSX 338 kb)
12526_2019_1009_MOESM2_ESM.xlsx (621 kb)
Supplementary material 2 Environmental parameters for all sampling locations, including presence and absence records (XLSX 620 kb)
12526_2019_1009_MOESM3_ESM.xlsx (112 kb)
Supplementary material 3 Associated bryozoan fauna found with Cellaria immersa, C. tenuirostris, C. pilosa and Cellaria sp. 5 (XLSX 111 kb)


  1. Bader B (2001) Modern bryomol-sediments in a cool-water, high-energy setting: the inner shelf off northern Brittany. Facies 44:81–103CrossRefGoogle Scholar
  2. Bader B (2002) Bryozoan communities in the Weddell Sea, Antarctica: a first overview. In: Wyse Jackson PN, Butler CJ, Spencer Jones ME (eds) Bryozoan Studies 2001: Proceedings of the 12th international Bryozoology association conference. Dublin, Ireland, Taylor and Francis: CRC Press, pp 1–6Google Scholar
  3. Bader B, Schäfer P (2005) Impact of environmental seasonality on stable isotope composition of skeletons of the temperate bryozoan Cellaria sinuosa. Palaeogeogr Palaeoclimatol Palaeoecol 226:58–71CrossRefGoogle Scholar
  4. Barnes DK, Griffiths HJ (2008) Biodiversity and biogeography of southern temperate and polar bryozoans. Glob Ecol Biogeogr 17:84–99Google Scholar
  5. Bastos AC, Moura RL, Moraes FC, Vieira LS, Braga JC, Ramalho LV, Amado-Filho GM, Magdalena UR, Webster JM (2018) Bryozoans are major modern builders of South Atlantic oddly shaped reefs. Sci Rep 8:9638CrossRefGoogle Scholar
  6. Batson PB, Probert PK (2000) Bryozoan thickets off Otago peninsula. Ministry of Fisheries, Wellington, NZGoogle Scholar
  7. Best BA (1988) Passive suspension feeding in a sea pen: effects of ambient flow on volume flow rate and filtering efficiency. Biol Bull 175:332–342CrossRefGoogle Scholar
  8. Boardman RS, McKinney FK, Taylor PD (1992) Morphology, anatomy, and systematics of the Cinctiporidae, new family (Bryozoa: Stenolaemata). Smithson Contrib Paleobiol 70:1–81CrossRefGoogle Scholar
  9. Bock P (2018) Cellaria Ellis and Solander, 1786. Accessed 10 April 2019
  10. Bock P (2019) World list of recent and fossil Bryozoa. Available online at Accessed 10 April 2019.
  11. Bostock H, Jenkins C, Mackay K, Carter L, Nodder S, Orpin A, Pallentin A, Wysoczanski R (2019a) Distribution of surficial sediments in the ocean around New Zealand/Aotearoa. Part A: continental slope and deep ocean. N Z J Geol Geophys 62:1–23CrossRefGoogle Scholar
  12. Bostock H, Jenkins C, Mackay K, Carter L, Nodder S, Orpin A, Pallentin A, Wysoczanski R (2019b) Distribution of surficial sediments in the ocean around New Zealand/Aotearoa. Part B: continental shelf. N Z J Geol Geophys 62:24–45CrossRefGoogle Scholar
  13. Bradstock M, Gordon DP (1983) Coral-like bryozoan growths in Tasman Bay, and their protection to conserve commercial fish stocks. N Z J Mar Freshw Res 17:159–163CrossRefGoogle Scholar
  14. Branch GM, Attwood CG, Gianakouras D, Branch ML (1993) Patterns in the benthic communities on the shelf of the sub-Antarctic Prince Edward islands. Polar Biol 13:23–34CrossRefGoogle Scholar
  15. Brey T, Gerdes D, Gutt J, Mackensen A, Starmans A (1999) Growth and age of the Antarctic bryozoan Cellaria incula on the Weddell Sea shelf. Antarct Sci 11:408–414CrossRefGoogle Scholar
  16. Brown DA (1952) The tertiary Cheilostomatous Polyzoa of New Zealand. Order of the Trustees of the British Museum, LondonGoogle Scholar
  17. Carter MC (2008). The functional morphology of avicularia in cheilostome bryozoans. Dissertation, Victoria University of WellingtonGoogle Scholar
  18. Di Geronimo I, La Perna R, Rosso A, Sanfilippo R (1998) Notes on two upper-circalittoral assemblages from the Amendolara Bank (northern Ionian Sea). Boll Acc Gioenia Sci Nat Catania 30(353):243–262Google Scholar
  19. Dinno A (2017) Dunn test: Dunn’s test of multiple comparisons using rank sums. R package version 1.3.5.
  20. Eckman JE, Okamura B (1998) A model of particle capture by bryozoans in turbulent flow: significance of colony form. Am Nat 152:861–880CrossRefGoogle Scholar
  21. Gordon DP (1984) The marine fauna of New Zealand: Bryozoa, Gymnolaemata from the Kermadec ridge. N Z Oceanogr Inst Mem 91:1–198Google Scholar
  22. Gordon DP (1986) The marine fauna of New Zealand: Bryozoa: Gymnolaemata (Ctenostomata and Cheilostomata Anasca) from the Western South Island continental shelf and slope. N Z Oceanogr Inst Mem 95:1–121Google Scholar
  23. Gordon DP (1989) Intertidal bryozoans from coral reef-flat rubble Sa 'aga, Western Samoa. N Z J Zool 16:447–463CrossRefGoogle Scholar
  24. Gordon DP, Mawatari SF (1992) Atlas of marine-fouling Bryozoa of new-Zealand ports and harbours. Misc Publ N Z Oceanogr Inst 107:1–52Google Scholar
  25. Gordon DP, Taylor PD (1999) Latest Paleocene to earliest Eocene bryozoans from Chatham Island, New Zealand. Bull Br Mus Nat Hist 55:1–45Google Scholar
  26. Gordon DP, Taylor PD (2001) New Zealand recent Densiporidae and Lichenoporidae (Bryozoa: Cyclostomata). Species Divers 6:243–290CrossRefGoogle Scholar
  27. Gordon DP, Taylor PD (2015) Bryozoa of the early Eocene Tumaio limestone, Chatham Island, New Zealand. J Syst Palaeontol 13:983–1070CrossRefGoogle Scholar
  28. Gordon DP, Clark AG, Harper JF (1987) Bryozoa. In: Pandian TJ, Vernberg FJ (eds) Animal energetics. Academic Press, New York, pp 173–199Google Scholar
  29. Gordon DP, Mawatari SF, Kajihara H (2002) New taxa of Japanese and New Zealand Eurystomellidae (phylum Bryozoa) and their phylogenetic relationships. Zool J Linnean Soc 136:199–216CrossRefGoogle Scholar
  30. Gordon DP, Taylor PD, Bigey FP (2009) Phylum Bryozoa: moss animals, sea mats, lace corals. In: Gordon DP (ed) New Zealand Inventory of Biodiversity: 1. Kingdom Animalia: Radiata, Lophotrochozoa, Deuterostomia. Canterbury University Press, Christchurch, pp 271–297Google Scholar
  31. Hageman SJ, Bayer MM, Todd CD (1999) Partitioning phenotypic variation: genotypic, environmental and residual components from bryozoan skeletal morphology. J Nat Hist 33:1713–1735CrossRefGoogle Scholar
  32. Harmelin JG (1968) Bryozoaires récoltés au cours de la campagne du Jean Charcot en Méditerranée orientale (Août-Septembre 1967). I. Dragages. Bull Mus Natl Hist Nat 40:1179–1208Google Scholar
  33. Harmelin JG (1976) Le sous-ordre Tubuliporina (Bryozoaires Cyclostomes) en Mediterranée. Mém Inst Océanogr Monaco 10:1–326Google Scholar
  34. Harmelin JG, d’Hondt JL (1992) Bryozoaires des parages de Gibraltar (campagne océanographique BALGIM, 1984). 1–Chéilostomes. Bull Mus Natl Hist Nat 14:23–67Google Scholar
  35. Harmelin JG, d'Hondt JL (1993) Transfers of bryozoan species between the Atlantic Ocean and the Mediterranean Sea via the strait of Gibraltar. Oceanol Acta 16:63–72Google Scholar
  36. Hayward PJ, Thorpe JP (1988) Species of Chaperiopsis (Bryozoa, Cheilostomata) collected by discovery investigations. J Nat Hist 22:45–69CrossRefGoogle Scholar
  37. Hayward PJ, Thorpe JP (1989) Membraniporoidea, Microporoidea and Cellarioidea (Bryozoa, Cheilostomata) collected by discovery investigations. J Nat Hist 23:913–959CrossRefGoogle Scholar
  38. Henrich R, Freiwald A, Betzler C, Bader B, Schäfer P, Samtleben C, Brachert TC, Wehrmann A, Zankl H, Kühlmann DH (1995) Controls on modern carbonate sedimentation on warm-temperate to arctic coasts, shelves and seamounts in the northern hemisphere: implications for fossil counterparts. Facies 32:71–108CrossRefGoogle Scholar
  39. Hermansen P, Larsen PS, Riisgård HU (2001) Colony growth rate of encrusting marine bryozoans (Electra pilosa and Celleporella hyalina). J Exp Mar Biol Ecol 263:1–23CrossRefGoogle Scholar
  40. Hirose M, Mawatari SF, Scholz J (2013) Distribution and diversity of erect bryozoan assemblages along the Pacific coast of Japan. In: Ernst A, Schäfer P, Scholz J (eds) Bryozoan studies 2010. Springer, Berlin, pp 121–136CrossRefGoogle Scholar
  41. Humphries S (2007) Body size and suspension feeding. In: Hildrew AG, Raffaelli DG, Edmonds-Brown R (eds) Body size and the structure and function of aquatic ecosystems. Cambridge University Press, Cambridge, pp 16–32CrossRefGoogle Scholar
  42. Jebram D (1977) Experimental techniques and culture methods. In: Woollacott RM, Zimmer RL (eds) Biology of bryozoans. Academic Press, New York, pp 273–306CrossRefGoogle Scholar
  43. John F, Sanford W (2011) An {R} companion to applied regression, 2nd edn. Sage, Thousand Oaks, CA Google Scholar
  44. Key MM, Jackson PNW, Falvey LW, Roth BJ (2014) Use of fossil bryozoans in sourcing lithic artefacts. Geoarchaeology 29:397–409CrossRefGoogle Scholar
  45. Kirchenpauer GH (1869) Neue Bryozoen. Museum Godeffroy Catalog IV(xxv-xxxiv):118–119Google Scholar
  46. Kobluk DR, Cuffey RJ, Fonda SS, Lysenko MA (1988) Cryptic Bryozoa, leeward fringing reef of Bonaire, Netherlands Antilles, and their paleoecological application. J Paleontol 62:427–439CrossRefGoogle Scholar
  47. López de la Cuadra CM, García-Gómez JC (1996) The species of Cellaria (Bryozoa: Cheilostomatida) with large avicularia from West Africa. J Nat Hist 30:153–161CrossRefGoogle Scholar
  48. Lukasik JJ, James NP, McGowran B, Bone Y (2000) An epeiric ramp: low-energy, cool-water carbonate facies in a tertiary inland sea, Murray Basin, South Australia. Sedimentology 47:851–881CrossRefGoogle Scholar
  49. McKinney FK, Jackson JB (1989) Life on and in sediments: problems of substratum stability. In: McKinney FK, Jackson JB (eds) Bryozoan evolution. University of Chicago Press, Chicago, pp 190–208Google Scholar
  50. McKinney FK, Jaklin A (2000) Spatial niche partitioning in the Cellaria meadow epibiont association, northern Adriatic Sea. Cah Biol Mar 41:1–17Google Scholar
  51. McKinney FK, Jaklin A (2001) Sediment accumulation in a shallow-water meadow carpeted by a small erect bryozoan. Sediment Geol 145:397–410CrossRefGoogle Scholar
  52. Mortimer N, Campbell H (2014) Zealandia: our continent revealed. Penguin group (NZ) for GNS science, AucklandGoogle Scholar
  53. Moyano HI (1973) Briozoos marinos Chilenos I. Briozoos de la Islas de Pascua I. Gayana Zoologia, Concepcion 25:1–22Google Scholar
  54. NASA Goddard Space Flight Center, Ocean Ecology Laboratory, Ocean Biology Processing Group (2018a) Moderate-resolution Imaging Spectroradiometer (MODIS) Terra Chlorophyll Data; Reprocessing. NASA OB.DAAC, Greenbelt, MD, USA. Accessed on 11/29/2018Google Scholar
  55. NASA Goddard Space Flight Center, Ocean Ecology Laboratory, Ocean Biology Processing Group (2018b) Moderate-resolution Imaging Spectroradiometer (MODIS) Terra Sea Surface Temperature Data; Reprocessing. NASA OB.DAAC, Greenbelt, MD, USA. Accessed on 11/29/2018Google Scholar
  56. Nelson CS, Keane SL, Head PS (1988) Non-tropical carbonate deposits on the modern New Zealand shelf. Sediment Geol 60:71–94CrossRefGoogle Scholar
  57. O’Dea A, Jackson JB (2002) Bryozoan growth mirrors contrasting seasonal regimes across the isthmus of Panama. Palaeogeogr Palaeoclimatol Palaeoecol 185:77–94CrossRefGoogle Scholar
  58. Ogle DH, Wheeler P, Dinno A (2018) FSA: fisheries stock analysis. R package version 0.8.22.
  59. O'Hara TD (2001) Consistency of faunal and floral assemblages within temperate subtidal rocky reef habitats. Mar Freshw Res 52:853–863CrossRefGoogle Scholar
  60. Peake BM, Walls DJ, Gibbs MT (2001) Spatial variations in the levels of nutrients, chlorophyll a, and dissolved oxygen in summer and winter in Doubtful Sound, New Zealand. N Z J Mar Freshw Res 35:681–694Google Scholar
  61. Probert PK, Batham EJ, Wilson JB (1979) Epibenthic macrofauna off southeastern New Zealand and mid-shelf bryozoan dominance. N Z J Mar Freshw Res 13:379–392CrossRefGoogle Scholar
  62. Ridgway KR, Dunn JR, Wilkin JL (2002) Ocean interpolation by four-dimensional least squares - application to the waters around Australia. J Atmos Ocean Technol 19:1357–1375CrossRefGoogle Scholar
  63. Riisgård HU, Larsen PS (2010) Particle capture mechanisms in suspension-feeding invertebrates. Mar Ecol Prog Ser 418:255–293CrossRefGoogle Scholar
  64. Riisgård HU, Manríquez P (1997) Filter-feeding in fifteen marine ectoprocts (Bryozoa): particle capture and water pumping. Mar Ecol Prog Ser 154:223–239CrossRefGoogle Scholar
  65. Rosso A (1996a) Popolamenti e tanatocenosi a Briozoi di fondi mobili circalitorali del Golfo di Noto (Sicilia SE). Nat Sicil 20(Ser. 4):189–225Google Scholar
  66. Rosso A (1996b) Valutazione della biodiversità in Mediterraneo: l’esempio dei popolamenti a briozoi della Biocenosi del Detritico Costiero. Biol Mar Meditt 3:58–65Google Scholar
  67. Rosso A, Sanfilippo R, Taddei Ruggiero E, Di Martino E (2013) Faunas and ecological groups of Serpuloidea, Bryozoa and Brachiopoda from submarine caves in Sicily (Mediterranean Sea). Boll Soc Paleontol Ital 52:167–176Google Scholar
  68. Rowden AA, Warwick RM, Gordon DP (2004) Bryozoan biodiversity in the New Zealand region and implications for marine conservation. Biodivers Conserv 13:2695–2721CrossRefGoogle Scholar
  69. Sebens KP, Johnson AS (1991) Effects of water movement on prey capture and distribution of reef corals. Hydrobiologia 226:91–101CrossRefGoogle Scholar
  70. Sebens KP, Witting J, Helmuth B (1997) Effects of water flow and branch spacing on particle capture by the reef coral Madracis mirabilis (Duchassaing and Michelotti). J Exp Mar Biol Ecol 211:1–28CrossRefGoogle Scholar
  71. Smith AM (1995) Palaeoenvironmental interpretation using bryozoans: a review. Geol Soc Lond Spec Publ 83:231–243CrossRefGoogle Scholar
  72. Smith AM (2007) Age, growth and carbonate production by erect rigid bryozoans in Antarctica. Palaeogeogr Palaeoclimatol Palaeoecol 256:86–98CrossRefGoogle Scholar
  73. Smith AM (2014) Growth and calcification of marine bryozoans in a changing ocean. Biol Bull 226:203–210CrossRefGoogle Scholar
  74. Smith AM, Gordon DP (2011) Bryozoans of southern New Zealand: a field identification guide. Ministry of Fisheries, Wellington, NZGoogle Scholar
  75. Smith AM, Key MM (2004) Controls, variation, and a record of climate change in detailed stable isotope record in a single bryozoan skeleton. Quat Res 61:123–133CrossRefGoogle Scholar
  76. Smith AM, Nelson CS, (2003) Effects of early sea-floor processes on the taphonomy of temperate shelf skeletal carbonate deposits. Earth-Science Reviews 63 (1–2):1–31Google Scholar
  77. Smith AM, Nelson CS (1994) Calcification rates of rapidly colonising bryozoans in Hauraki gulf, northern New Zealand. N Z J Mar Freshw Res 28:227–234CrossRefGoogle Scholar
  78. Smith AM, Taylor PD, Spencer HG (2008) Resolution of taxonomic issues in the Horneridae (Bryozoa: Cyclostomata). Ann Bryozool 2:359–411Google Scholar
  79. de Soares MO, da Lotufo TMC, Vieira LM, Salani S, Hajdu E, Matthews–Cascon H, Leão ZM, de Kikuchi RKP (2017) Brazilian marine animal forests: a new world to discover in the southwestern Atlantic. In: Rossi S, Bramanti L, Gori A, Orejas C (eds) Marine animal forests: the ecology of benthic biodiversity hotspots. Springer International Publishing, Cham, pp 73–110Google Scholar
  80. Sonar MA, Badve RM, Gaikwad SG (2010) Usefulness of Bryozoa to deduce palaeoenvironment: a case study from the Holocene of west coast of Maharashtra and Goa. Gondwana Geol Mag 25:69–80Google Scholar
  81. Steger KK, Smith AM (2005) Carbonate mineralogy of free-living bryozoans (Bryozoa: Otionellidae), Otago shelf, southern New Zealand. Palaeogeogr Palaeoclimatol Palaeoecol 218:195–203CrossRefGoogle Scholar
  82. Strathmann RR (1982) Cinefilms of particle capture by an induced local change of beat of lateral cilia of a bryozoan. J Exp Mar Biol Ecol 62:225–236CrossRefGoogle Scholar
  83. Strathmann RR, McEdward LR (1986) Cyphonautes' ciliary sieve breaks a biological rule of inference. Biol Bull 171:694–700CrossRefGoogle Scholar
  84. Taylor PD, Gordon DP, Batson PB (2004) Bathymetric distributions of modern populations of some common Cenozoic Bryozoa from New Zealand, and paleodepth estimation. N Z J Geol Geophys 47:57–69CrossRefGoogle Scholar
  85. Tenison-Woods JE (1880) Palaeontology of New Zealand. Part 4. Corals and Bryozoa of the Neozoic period in New Zealand. Government printer, WellingtonGoogle Scholar
  86. Wass RE, Yoo JJ (1975) Bryozoa from Site 282 West of Tasmania. In: Kennett JR, Houtz RE, Andrews PB, Edwards VP, Gostin VA, Hajos M, Hampton MA, Jenkins DG, Margolis SV, Ovenshire AT, Perch–Nielsen K (eds) Initial reports on the Deep Sea Drilling Project, 29. U.S. Government Printing Office, Washington, pp 809–831.Google Scholar
  87. Waters AW (1887) On tertiary chilostomatous Bryozoa from New Zealand. Q J Geol Soc 43:40–72CrossRefGoogle Scholar
  88. Wickham H (2009) ggplot2: elegant graphics for data analysis. Springer-Verlag, New YorkCrossRefGoogle Scholar
  89. Winston JE (1978) Polypide morphology and feeding behavior in marine ectoprocts. Bull Mar Sci 28:1–31Google Scholar
  90. Wood AC, Probert PK, Rowden AA, Smith AM (2012) Complex habitat generated by marine bryozoans: a review of its distribution, structure, diversity, threats and conservation. Aquat Conserv Mar Freshwat Ecosyst 22:547–563CrossRefGoogle Scholar
  91. Wood AC, Rowden AA, Compton TJ, Gordon DP, Probert PK (2013) Habitat-forming bryozoans in New Zealand: their known and predicted distribution in relation to broad-scale environmental variables and fishing effort. PLoS One 8:e75160CrossRefGoogle Scholar

Copyright information

© Senckenberg Gesellschaft für Naturforschung 2019

Authors and Affiliations

  1. 1.Department of Marine ScienceUniversity of OtagoDunedinNew Zealand
  2. 2.National Institute of Water and Atmospheric ResearchWellingtonNew Zealand

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