Marine Biodiversity

, Volume 42, Issue 2, pp 161–171

Palolo and un: distinct clades in the genus Palola (Eunicidae, Polychaeta)

Original Paper


The annual swarming of epitokes of the eunicid polychaete Palola viridis has great cultural significance in Samoa and other Indo-Pacific locations. Palola spp. occur worldwide in warm shallow-water environments, but not all populations appear to swarm. Morphological distinctions among Palola species are subtle but, based on phylogenetic analysis of mitochondrial markers, two species groups can be distinguished, each containing numerous subclades. The objectives of this study were to determine the position of P. viridis in the phylogeny of the genus, to assess the species’ genetic distinctness and to describe P. viridis epitokes morphologically. We sequenced epitokes from Samoa (locally known as palolo) and Vanuatu (un) for two mitochondrial markers and performed phylogenetic analyses in combination with previously published sequences. The samples from Samoa and Vanuatu formed a common clade in species group B. We suggest that swarming is restricted to this species group, although probably not to a single species.


Samoa Palola Swarming Epitokes 

Supplementary material

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Table S1(PDF 44.7 KB)
12526_2011_100_MOESM2_ESM.pdf (30 kb)
Table S2(PDF 30 kB)
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Table S3(PDF 30 kB)


  1. Bartlett HK (1947) The Swarming of the polychaete worm Eunice viridis at Misima Is., S.E. Papua. South Austr Nat 24:3–4Google Scholar
  2. Brown D (2009) Spawning behavior of Palola viridis (Polychaeta: Eunicidae) in American Samoa. Coral Reefs 28:535–535CrossRefGoogle Scholar
  3. Brown G (1877) Notes on the Duke of York Group, New Britain, and New Ireland. J R Geogr Soc 47:137–150Google Scholar
  4. Burrows W (1945) Periodic spawning of 'Palolo' worms in Pacific waters. Nature 155:47–48CrossRefGoogle Scholar
  5. Burrows W (1955) Notes on the periodic appearance of the annelid worm Eunice viridis (Gray) in the South-west Pacific Islands. J Polyn Soc 64:137–154Google Scholar
  6. Caspers H (1961) Beobachtungen über Lebensraum und Schwärmperiodizität des Palolowurmes Eunice viridis (Polychaeta, Eunicidae). Int Rev Ges Hydrobiol 46:175–183CrossRefGoogle Scholar
  7. Caspers H (1984) Spawning periodicity and habitat of the palolo worm Eunice viridis (Polychaeta: Eunicidae) in the Samoan Islands. Mar Biol 79:229–236CrossRefGoogle Scholar
  8. Codrington RH (1891) The Melanesians. Studies in their anthropology and folk-lore. Clarendon Press, OxfordGoogle Scholar
  9. Duda TF, Chang D, Lewis BD, Lee TW (2009) Geographic variation in venom allelic composition and diets of the widespread predatory marine gastropod Conus ebraeus. Plos One 4Google Scholar
  10. Excoffier L, Lischer H (2009) Arlequin ver. 3.5. An integrated software package for population genetics data analysis. Bern, SwitzerlandGoogle Scholar
  11. Fauchald K (1992) Review of the types of Palola (Eunicidae: Polychaeta). J Nat Hist 26:1177–1225CrossRefGoogle Scholar
  12. Folmer O, Black M, Hoen W, Lutz R, Vrijenhoek R (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metozoan invertebrates. Mol Mar Biol Biotech 3:294–299Google Scholar
  13. Friedlaender B (1898) Ueber den sogenannten Palolowurm. Biol Centralbl 18:337–357Google Scholar
  14. Galloway TW, Welch PS (1911) Studies on a phosphorescent Bermudan annelid, Odontosyllis Enopla Verrill. Trans Am Microsc Soc 30:13–39CrossRefGoogle Scholar
  15. Gambi MC, Cigliano M (2006) Observations on reproductive features of three species of Eunicidae (Polychaeta) associated with Posidonia oceanica seagrass meadows in the Mediterranean Sea. Sci Mar 70S3:301–308Google Scholar
  16. Giangrande A (1997) Polychaete reproductive patterns, life cycles and life histories: an overview. Oceanogr Mar Biol Annu Rev 35:323–368Google Scholar
  17. Gray JE (1847) An account of Palolo, a sea worm eaten in the Navigator Islands. Proc Zool Soc 15:17–18Google Scholar
  18. Hall TA (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic acid symposium series 41:95–98Google Scholar
  19. Hauenschild C, Fischer A, Hoffmann DK (1968) Untersuchungen am pazifischen Palolawurm Eunice viridis (Polychaeta) in Samoa. Helgo Wiss Meeresunters 18:254–295CrossRefGoogle Scholar
  20. Hofmann DK (1972) Nachweis der Epitokie als Fortpflanzungsmodus des Polychaeten Eunice siciliensis (Polychaeta: Annelida). Mar Biol 14:341–344CrossRefGoogle Scholar
  21. Hofmann DK (1974) Maturation, epitoky and regeneration in the polychaete Eunice siciliensis under field and laboratory conditions. Mar Biol 25:149–161CrossRefGoogle Scholar
  22. Hofmann DK (1975) Reproductive forms in eunicid polychaetes inhabiting the "fonds coralligènes in the region of Banyuls-s.-M. (Mediterranean Sea) with particular reference to Eunice siciliensis. Pubbl St Zool Napoli 39:242–253Google Scholar
  23. Kohn AJ (1959) The ecology of Conus in Hawaii. Ecol Monogr 29:47–90CrossRefGoogle Scholar
  24. Kohn AJ (1968) Microhabitats, abundance and food of Conus on atoll reefs in the Maldive and Chagos Islands. Ecology 49:1046–1062CrossRefGoogle Scholar
  25. Kohn AJ (1980) Abundance, diversity and resource use in an assemblage of Conus species in Enewetak Lagoon. Pac Sci 34:359–369Google Scholar
  26. Kohn AJ (2001) Maximal species richness in Conus: diversity, diet and habitat on reefs of northeast Papua New Guinea. Coral Reefs 20:25–38CrossRefGoogle Scholar
  27. Kohn AJ, Nybakken JW (1975) Ecology of Conus on Eastern Indian Ocean fringing reefs: diversity of species and resource utilization. Mar Biol 29:211–234CrossRefGoogle Scholar
  28. Korringa P (1947) Relations between the moon and periodicity in the breeding of marine animals. Ecol Monogr 17:347–381CrossRefGoogle Scholar
  29. Kumar V, Dey A, Singh A (2009) MEGA: A Bio Computational Software for Sequence and Phylogenetic Analysis. In: Ao SI, Gelman L, Hukins DWL, Hunter A, Korsunsky AM (eds) World Congress on Engineering 2009, vol I, and Ii. Int Assoc Engineers-Iaeng, Hong Kong, pp 1863–1865Google Scholar
  30. Leviten PJ (1978) Resource partitioning by predatory gastropods of the genus Conus on subtidal Indo-Pacific coral reefs: the significance of prey size. Ecology 59:614–631CrossRefGoogle Scholar
  31. Martens JM, Heuer U, Hartmann-Schroeder G (1995) Swarming of Palola viridis Gray and other Polychaeta such as Lysidice oele Horst and Lumbrineris natans sp. nov. on Ambon (Moluccas, Indonesia). Mitt Hamburg Zool Mus Inst 92:7–33Google Scholar
  32. Mondragón C (2004) Of winds, worms and mana: the traditional calendar of the Torres Islands, Vanuatu. Oceania 74:289–308Google Scholar
  33. Palumbi SR (1996) Nucleic acids II: the polymerase chain reaction. In:Hillis DM, Moritz C, Mable BK (eds) Molecular systematics.Sinauer, Sunderland, pp 205–247Google Scholar
  34. Pamungkas J (2011) Delicious! Marine worms from Ambon Island, Indonesia. Habitat Mar:35–37Google Scholar
  35. Pleijel F (2001) Eunicidae. In: Rouse G, Pleijel F (eds) Polychaetes. Oxford University Press, New York, pp 155–159Google Scholar
  36. Powell T (1883) Remarks on the structure and habits of the coral-reef annelid, Palolo viridis. J Linn Soc 16:193–396Google Scholar
  37. Ronquist F, Huelsenbeck JP (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19:1572–1574PubMedCrossRefGoogle Scholar
  38. Rouse GW (2000) The epitome of hand waving? Larval feeding and hypotheses of metazoan phylogeny. Evol Dev 2:222–233PubMedCrossRefGoogle Scholar
  39. Saito M, Okamoto A, Kohno T, Takakura S, Shinozaki H, Isonishi S, Yasuhara T,Yoshimura T, Ohtake Y, Ochiai K, Yokota J, Tanaka T (2000) Allelic imbalance and mutations of the PTEN gene in ovarian cancer. Int J Cancer 85:160–165Google Scholar
  40. Schröder O (1905) Beiträge zur Kenntnis der Bauchsinnesorgane (Bauchaugen) von Eunice viridis Gray sp. (Palolo). Zeitschr Wiss Zool 79: 132–149, Taf. 137–138Google Scholar
  41. Schulze A (2006) Phylogeny and genetic diversity of palolo worms (Palola, Eunicidae, Polychaeta) from the tropical North Pacific and the Caribbean. Biol Bull 210:25–37PubMedCrossRefGoogle Scholar
  42. Seeman B (1862) Viti. An account of a government mission to the Vitian or Fijian Islands in the years 1860–1861. Macmillan, CambridgeCrossRefGoogle Scholar
  43. Stair JB (1897) Palolo, a sea-worm eaten by the Samoans. J Polyn Soc 6:141–144Google Scholar
  44. Tamura K, Dudley J, Nei M, Kumar S (2007) MEGA4: Molecular evolutionary genetics analysis (MEGA) software version 4.0. Mol Biol Evol 24:1596–1599PubMedCrossRefGoogle Scholar
  45. Tamura K, Nei M, Kumar S (2004) Prospects for inferring very large phylogenies by using the neighbor-joining method. Proc Nat Acad Sci USA 101:11030–11035PubMedCrossRefGoogle Scholar
  46. Taylor JD (1986) Diets of sand-living predatory gastropods at Piti Bay, Guam. Asian Mar Biol 3:47–58Google Scholar
  47. Thompson JD, Higgins DG, Gibson TJ (1994) Clustal W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position specific gap penalties and weight matrix choice. Nucleic Acids Res 22:4673–4680PubMedCrossRefGoogle Scholar
  48. Treadwell AL (1922) Leodicidae from Fiji and Samoa. Papers from the Department of Marine Biology of the Carnegie Institution of Washington 18:127–170Google Scholar
  49. Woodworth WM (1903) Preliminary report on the "Palolo" worm of Samoa, Eunice viridis (Gray). Am Nat 37:875–818, 871 figCrossRefGoogle Scholar
  50. Xiong B, Kocher TD (1991) Comparison of mtDNA sequences from seven morphospecies of black flies. Genome 34:651–701Google Scholar
  51. Zanol J, Halanych KM, Struck TH, Fauchald K (2010) Phylogeny of the bristle worm family Eunicidae (Eunicida, Annelida) and the phylogenetic utility of noncongruent 16S, COI and 18S in combined analyses. Mol Phylogenet Evol 55:660–676PubMedCrossRefGoogle Scholar

Copyright information

© Senckenberg, Gesellschaft für Naturforschung and Springer 2011

Authors and Affiliations

  1. 1.Department of Marine BiologyTexas A&M University at GalvestonGalvestonUSA

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