The Science of Nature

, 103:11 | Cite as

Penis morphology in a Burmese amber harvestman

  • Jason A. DunlopEmail author
  • Paul A. Selden
  • Gonzalo Giribet
Short Communication


A unique specimen of the fossil harvestman Halitherses grimaldii Giribet and Dunlop, 2005 (Arachnida: Opiliones) from the Cretaceous (ca. 99 Ma) Burmese amber of Myanmar reveals a fully extended penis. This is the first record of a male copulatory organ of this nature preserved in amber and is of special importance due to the age of the deposit. The penis has a slender, distally flattened truncus, a spatulate heart-shaped glans and a short distal stylus, twisted at the tip. In living harvestmen, the penis yields crucial characters for their systematics. Male genital morphology in H. grimaldii appears to be unique among the wider Dyspnoi clade to which this fossil belongs. The large eyes in the fossil differ markedly from other members of the subfamily Ortholasmatinae to which H. grimaldii was originally referred. Based on recent data, it has been argued that large eyes may be plesiomorphic for Palpatores (i.e. the suborders Eupnoi and Dyspnoi), potentially rendering this character plesiomorphic for the fossil too. Thus, the unique structure of the penis seen here, and the probable lack of diaphanous teeth, present in all other extant non-acropsopilionid Dyspnoi, suggest that H. grimaldii represents a new, extinct family of large-eyed dyspnoid harvestmen, Halithersidae fam. nov.; a higher taxon in amber diagnosed here on both somatic and genital characters.


Arachnida Opiliones Male genitalia Systematics Amber Myanmar 



We thank Jörg Wunderlich (Hirschberg) for making material from his collection available and Jenn Lenihan (Harvard) for producing the micro-CT images. Five anonymous reviewers provided constructive criticism.


  1. Dunlop JA, Mammitzsch L (2010) A new genus and species of harvestman from Baltic amber. Palaeodiversity 3:23–32Google Scholar
  2. Dunlop JA, Bartel C, Mitov PG (2012) An enigmatic spiny harvestman from Baltic amber. Foss Rec 15:91–101CrossRefGoogle Scholar
  3. Dunlop JA, Anderson LI, Kerp H, Hass H (2003) Preserved organs of Devonian harvestmen. Nature 425:916CrossRefPubMedGoogle Scholar
  4. Dunlop JA, Anderson LI, Kerp H, Hass H (2004) A harvestman (Arachnida: Opiliones) from the Early Devonian Rhynie cherts, Aberdeenshire, Scotland. Trans R Soc Edinburgh, Earth Sci 94:341–354Google Scholar
  5. Garwood RJ, Dunlop JA, Giribet G, Sutton MD (2011) Anatomically modern Carboniferous harvestmen demonstrate early cladogenesis and stasis in Opiliones. Nat Comm 2:444CrossRefGoogle Scholar
  6. Garwood RJ, Sharma PP, Dunlop JA, Giribet G (2014) A new stem-group Palaeozoic harvestman revealed through integration of phylogenetics and development. Curr Biol 24:1–7CrossRefGoogle Scholar
  7. Giribet G, Dunlop JA (2005) First identifiable Mesozoic harvestman (Opiliones: Dyspnoi) from Cretaceous Burmese amber. Proc R Soc B 272:1007–1013PubMedCentralCrossRefPubMedGoogle Scholar
  8. Giribet G, Tourinho AL, Shih C, Ren D (2012) An exquisitely preserved harvestman (Arthropoda, Arachnida, Opiliones) from the Middle Jurassic of China. Org Div Evol 12:51–56CrossRefGoogle Scholar
  9. Groh S, Giribet G (2015) Polyphyly of Caddoidea, reinstatement of the family Acropsopilionidae in Dyspnoi, and a revised classification system of Palpatores (Arachnida, Opiliones). Cladistics 31:277–290CrossRefGoogle Scholar
  10. Gruber J (2007) Taxonomy: Dyspnoi. In: Pinto da Rocha R, Machado G, Giribet G (eds) Harvestmen. The biology of Opiliones. Harvard University Press, Cambridge, pp 131–159Google Scholar
  11. Karaman I (2005) Evidence of spermatophores in Cyphophthalmi (Arachnida, Opiliones). Rev suisse Zool 112:3–11CrossRefGoogle Scholar
  12. Kury AB (2013) Order Opiliones Sundevall, 1833. Zootaxa 3703:27–33CrossRefGoogle Scholar
  13. Macías-Ordóñez R, Machado G, Pérez-González A, Shultz JW (2010) Genitalic evolution in Opiliones. In: Leonard J, Córdoba-Aguilar A (eds) The evolution of primary sexual characters in animals. Oxford University Press, Oxford, pp 285–306Google Scholar
  14. Martens J (1976) Genitalmorphologie, System und Phylogenie der Weberknechte (Arachnida, Opiliones). Entomol German 3:51–68Google Scholar
  15. Martens J (1983) Europäische Arten der Gattung Sabacon Simon 1879 (Arachnida: Opiliones: Sabaconidae). Senckenbergiana Biol 63:265–296Google Scholar
  16. Martens J (1986) Die Grossgliederung der Opiliones und die Evolution der Ordnung (Arachnida). Actas X Congr Int Arachnol, Jaca 1:289–310Google Scholar
  17. Perreau M, Tafforeau P (2011) Virtual dissection using phase-contrast X-ray synchrotron microtomography: reducing the gap between fossils and extant species. Syst Ent 36:573–580CrossRefGoogle Scholar
  18. Ross A, Mellish C, York P, Crighton B (2010) Burmese amber. In: Penney D (ed) Biodiversity of fossil in amber from the major world deposits. Siri Scientific Press, Manchester, pp 208–235Google Scholar
  19. Schönhofer AL (2013) A taxonomic catalogue of the Dyspnoi Hansen and Sørensen, 1904 (Arachnida: Opiliones). Zootaxa 3679:1–68CrossRefPubMedGoogle Scholar
  20. Schönhofer AL, Martens J (2012) The enigmatic Alpine opilionid Saccarella schilleri gen. n., sp. n. (Arachnida: Nemastomatidae)—isolated systematic placement inferred from comparative genital morphology. Org Div Evol 12:409–419CrossRefGoogle Scholar
  21. Schwendinger PJ, Gruber J (1992) A new Dendrolasma (Opiliones, Nemastomatidae) from Thailand. Bull Br arachnol Soc 9:57–60Google Scholar
  22. Sharma PP, Giribet G (2014) A revised dated phylogeny of the arachnid order Opiliones. Front Genet 5:255PubMedCentralPubMedGoogle Scholar
  23. Shear WA (2010) New species and records of ortholasmatine harvestmen from México, Honduras, and the western United States (Opiliones, Nemastomatidae, Ortholasmatinae). ZooKeys 52:9–45CrossRefPubMedGoogle Scholar
  24. Shi G, Grimaldi DA, Harlow GE, Wang J, Wang J, Yang M, Lei W, Li Q, Li X (2012) Age constraint on Burmese amber based on U–Pb dating of zircons. Cret Res 37:155–163CrossRefGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2016

Authors and Affiliations

  • Jason A. Dunlop
    • 1
    Email author
  • Paul A. Selden
    • 2
    • 3
  • Gonzalo Giribet
    • 4
  1. 1.Museum für Naturkunde, Leibniz Institute for Evolution and Biodiversity ScienceBerlinGermany
  2. 2.Department of Geology and Paleontological InstituteUniversity of KansasLawrenceUSA
  3. 3.Department of Earth Sciences, Natural History MuseumLondonUK
  4. 4.Museum of Comparative Zoology, Department of Organismic and Evolutionary BiologyHarvard UniversityCambridgeUSA

Personalised recommendations