, Volume 130, Issue 1, pp 1–15 | Cite as

Pattern of occurrence of supraneural coelomopores and intertentacular organs in Gymnolaemata (Bryozoa) and its evolutionary implications

  • Andrew N. Ostrovsky
  • Joanne S. Porter
Review Article


The evolution of bryozoan female gonopores (the supraneural coelomopore (SNP) and the intertentacular organ (ITO)) is considered in the light of two alternative hypotheses. In the first hypothesis it is proposed that the ITO originated from the shortening and fusion of two tentacles possessing terminal pore(s), with further transformation into a simple pore. In the alternative hypothesis it is suggested that the ITO evolved from a coelomopore with a contribution from the basal parts of two disto-medial tentacles in an ancestor. Favouring the second hypothesis, in this paper we present a hypothetical scenario, according to which the earliest gymnolaemate bryozoans with uniserial growth and a broadcasting reproductive pattern possessed the supraneural coelomopore (SNP). This could serve both as a female gonopore and as a conduit for sperm entry. Evolution of large colonies of closely packed zooids led to development of the tube-like intertentacular organ (ITO) that is formed by epithelial proliferation of the basal parts of two dorso-medial tentacles. This prevented egg swallowing in the situation when water exchange was hampered within the large colony. The ITO independently evolved in both ctenostome and cheilostome gymnolaemates when multiserial colonies appeared. Evolution of brooding in species with colonies of closely packed zooids led to reduction of the ITO, except for the cheilostomes Tendra and Thalamoporella that acquired brooding independently. A rudimentary ITO also “survived” in two ctenostomes with the “mixed” type of brooding. An alternative, analogous organ—the ovipositor—has evolved in the cheilostome taxon Schizoporella.


Gonopore Fertilization Spawning Evolution Bryozoa 



Financial support for ANO was provided by the Lise Meitner Foundation, the FWF Grants P19337-B17 and P22696-B17 (Austria) and the RFBR Grants 07-04-00928a and 10-04-00085-a (Russia). Aberystwyth University provided financial support for JSP. Thanks to Dr Michael Winson (Heriot-Watt University), Professors John S. Ryland (Swansea University), Russel Zimmer (University of Southern California), Michael H. Temkin (St Lawrence University) and Dr Alexander Gruhl (Free University of Berlin) for constructive criticism on the early versions of the text and for illustrations. Dr Judith Winston (Virginia Museum of Natural History) and Dr Peter S. Cadman (Swansea University) provided some illustrations. Professor Timothy S. Wood (Wright State University) and Dr M. Carter (Heriot-Watt University) are acknowledged for unpublished data on Hislopia malayensis and Victorella pavida. Hans De Blauwe (Belgium) provided information on the ITO in Arachnidium fibrosum and Alcyonidium parasiticum. We also thank Professor Thomas Bartolomaeus (University of Bonn) and anonymous reviewers whose suggestions substantially helped to improve the manuscript.


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Authors and Affiliations

  1. 1.Department of Invertebrate Zoology, Faculty of Biology and Soil ScienceSt. Petersburg State UniversitySt. PetersburgRussia
  2. 2.Department of Palaeontology, Faculty of Earth Sciences, Geography and Astronomy, GeozentrumUniversity of ViennaViennaAustria
  3. 3.School of Life Sciences, Centre for Marine Biodiversity and BiotechnologyHeriot-Watt UniversityEdinburghUK

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