Morphological Shape, Episkeletobiont Analysis, and Life Orientation Study in Pseudoatrypa cf. lineata (Brachiopoda) from the Lower Genshaw Formation of the Middle Devonian Traverse Group, Michigan: A Geometric Morphometric Approach

  • Rituparna Bose
Part of the Springer Theses book series (Springer Theses)


Atrypids examined from the lower Genshaw Formation of the Middle Devonian (early middle Givetian) Traverse Group include a large assemblage of Pseudoatrypa bearing a rich fauna of episkeletobionts. We identified two species of PseudoatrypaPseudoatrypa lineata and Pseudoatrypa sp. A based on ornamentation and shell shape. Qualitative examination suggested that the former had fine-medium size ribbing, narrow hinge line, widened anterior, gentle to steep mid-anterior fold, a more domal shaped dorsal valve, and an inflated ventral valve in contrast to the coarse ribbing, widened hinge line, narrow anterior, gentle mid-anterior fold, arched shape dorsal valve, and flat ventral valve of the latter. Geometric morphometric analysis supported two statistically different shapes (p < 0.01) for the two distinct species. This study further examined these atrypids to investigate the influence of morphology on episkeletobiont settlement on the two Pseudoatrypa species. Among the 343 atrypid hosts examined, nearly 50 % were encrusted by episkeletobionts. Common encrusters included microconchids, bryozoan sheets, and hederellids. Less common encrusters included auloporid corals, cornulitids, tabulate corals, Ascodictyon, craniid brachiopods, and fenestrate bryozoans. Hederellids, auloporid corals, cornulitids, and tabulate corals encrusted a few living Pseudoatrypa hosts, but determination of pre- or post-mortem encrustation by the majority of episkeletobionts is equivocal. In a very few cases, episkeletobionts crossed the commissure indicating the death of the host. Some episkeletobionts, microconchids and the sheet bryozoans, were more common on Pseudoatrypa lineata, which exhibited more dorsal–ventral convexity than Pseudoatrypa sp. A. Perhaps, P. lineata may have provided a larger surface area for episkeletobiont settlement relative to Pseudoatrypa sp. A. In both the host species, encrustation was heaviest on the convex dorsal valve. This suggests that most of the encrustation occurred in a reclining, dorsal-valve-up life orientation of both species, in which the convex dorsal valve was exposed in the water column and the ventral valve remained in contact with the substrate. However, life orientations of these atrypid species could not be confidently predicted simply from the location preferences of episkeletobionts alone, as the life orientation of the host would also have been a hydrodynamically stable orientation of the articulated shell after death. Most episkeletobionts encrusted the posterior region of both dorsal and ventral valves of the two species, which suggests that the inflated areas of these valves, when exposed, favored the settlement of most episkeletobiont larvae.


Discriminant Function Analysis Anterior Commissure Middle Devonian Ventral Valve Dorsal Valve 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


  1. Ager DV (1961) The epifauna of a Devonian spiriferid. Q J Geol Soc Lond 117:1–10CrossRefGoogle Scholar
  2. Alexander RR (1975) Phenotypic liability of the brachiopod Rafinesquina alternata (Ordovician) and its correlation with the sedimentologic regime. J Paleontol 49:607–618Google Scholar
  3. Alexander RR (1984) Comparative hydrodynamic stability of brachiopod shells on current-scoured arenaceous substrates. Lethaia 17:17–32CrossRefGoogle Scholar
  4. Alexander RR (1990) Mechanical strength of shells of selected extant articulate brachiopods: implications for paleozoic morphologic trends. His Biol Int J Paleobiol 3:169–188CrossRefGoogle Scholar
  5. Alvarez F, Taylor PD (1987) Epizoan ecology and interactions in the Devonian of Spain. Palaeogeogr Palaeoclimatol Palaeoecol 61:17–31CrossRefGoogle Scholar
  6. Anderson WI, Megivern KD (1982) Epizoans from the Cerro Gordo member of the lime creek formation (upper Devonian), Rockford, Iowa. In: Proceedings of the Iowa academy of science, vol 89, pp 71–80Google Scholar
  7. Barringer JE (2008) Analysis of the occurrence of microconchids on middle Devonian brachiopods from the Michigan basin: implications for microconchid and brachiopod autecology. Unpublished M S thesis, Michigan State University, Michigan, pp 1–127Google Scholar
  8. Bartholomew JB (2006) Middle Devonian faunas of the Michigan and Appalachian Basins: comparing patterns of biotic stability and turnover between two paleobiogeographic subprovinces. Unpublished M S thesis, University of Cincinnati, Ohio, pp 1–300Google Scholar
  9. Bassler RS (1939) The Hederelloidea, a suborder of Paleozoic cyclostomatous Bryozoa. In: Proceedings of the United States National Museum, vol 87, pp 25–91Google Scholar
  10. Bookstein FL (1989) Principal warps: thin-plate splines and the decomposition of deformations. IEEE Trans Pattern Anal Mach Intell 11:567–585CrossRefGoogle Scholar
  11. Bookstein FL (1991) Morphometric tools for landmark data: geometry and biology. Cambridge University Press, New YorkGoogle Scholar
  12. Bordeaux YL, Brett CE (1990) Substrate specific associations of epibionts on middle Devonian brachiopods: implications for Paleoecology. Hist Biol 4:203–220Google Scholar
  13. Bose R, Schneider C, Polly PD, Yacobucci MM (2010) Ecological interactions between Rhipidomella (Orthides, Brachiopoda) and its endoskeletobionts and predators from the middle Devonian Dundee Formation of Ohio, United States. Palaios 25:196–210CrossRefGoogle Scholar
  14. Bowen ZP (1966) Intraspecific variation in the Brachial Cardinalia of Atrypa reticularis. J Paleontol 40:1017–1022Google Scholar
  15. Brett CE, Baird GC, Bartholomew AJ, DeSantis MK, Straeten CA (2010) Sequence stratigraphy and a revised sea-level curve for the middle Devonian of eastern North America. Palaeogeogr Palaeoclimatol Palaeoecol 304:21–53CrossRefGoogle Scholar
  16. Brezinski DK (1984) Upper Mississippian epizoans and hosts from southwestern Pennsylvania. In: Proceedings of the Pennsylvania academy of science, vol 58, pp 223–226Google Scholar
  17. Carrera MG (2000) Epizoan-sponge interactions in the early Ordovician of the Argentine Precordillera. Palaios 15:261–272Google Scholar
  18. Copper P (1967) Adaptations and life habits of Devonian atrypid brachiopods. Palaeogeogr Palaeoclimatol Palaeoecol 3:363–379CrossRefGoogle Scholar
  19. Copper P (1973) New Siluro-Devonian atrypoid brachiopods. J Paleontol 47:484–500Google Scholar
  20. Curry GB (1983) Brachiopod caeca—a respiratory role? Lethaia 16:311–312CrossRefGoogle Scholar
  21. Day J (1998) Distribution of latest Givetian-Frasnian Atrypida (Brachiopoda) in central and western North America. Acta Palaeontologica Polonica 43:205–240Google Scholar
  22. Day J, Copper P (1998) Revision of latest Givetian-Frasnian Atrypida (Brachiopoda) from central North America. Acta Palaeontologica Polonica 43:155–204Google Scholar
  23. Dietl GP, Kelley PH (2001) Mid-Paleozoic latitudinal predation gradient: distribution of brachiopod ornamentation reflects shifting carboniferous climate. Geology 29:111–114CrossRefGoogle Scholar
  24. Ehlers GM, Kesling RV (1970) Devonian strata of Alpena and Presque Isle counties, Michigan. Guidebook for field trips, Michigan Basin Geological Society, pp 1–130Google Scholar
  25. Fagerstrom JA (1996) Paleozoic brachiopod symbioses: testing the limits of modern analogues in paleoecology. Geol Soc Am Bull 108:1393–1403CrossRefGoogle Scholar
  26. Fenton CL, Fenton MA (1932) Orientation and injury in the genus Atrypa. Am Midl Nat 13:63–74CrossRefGoogle Scholar
  27. Fenton CL, Fenton MA (1935) Atrypae described by Clement L. Webster and related forms (Devonian, Iowa). J Paleontol 9:369–384Google Scholar
  28. Gibson MA (1992) Some epibiont-host and epibiont–epibiont relationships from the Birdsong Shale Member of the Lower Devonian Ross Formation (West-Central Tennessee, U.S.A.). Hist Biol 6:113–132Google Scholar
  29. Hammer Ø, Harper D (2005) Paleontological data analysis. Blackwell Publishing, OxfordCrossRefGoogle Scholar
  30. Haney RA, Mitchell CE, Kim K (2001) Geometric morphometric analysis of patterns of shape change in the Ordovician Brachiopod Sowerbyella. Palaios 16:115–125Google Scholar
  31. Hoare RD, Steller DL (1967) A Devonian brachiopod with epifauna. Ohio J Sci 67:291Google Scholar
  32. Hurst JM (1974) Selective epizoan encrustation of some Silurian brachiopods from Gotland. Palaeontology 17:423–429Google Scholar
  33. Kelly AW, Smith GW (1947) Stratigraphy and structure of traverse group in Afton-Onaway area, Michigan. Bull Am Assoc Petrol Geol 31:447–469Google Scholar
  34. Kesling RV, Chilman RB (1975) Strata and megafossils of the middle Devonian silica formation. Pap Paleontol 8:1–408Google Scholar
  35. Kesling RV, Hoare RD, Sparks DK (1980) Epizoans of the Middle Devonian brachiopod Paraspirifer bownockeri: their relationships to one another and to their host. J Paleontol 54:1141–1154Google Scholar
  36. Koch WF (1978) Brachiopod paleoecology, paleobiogeography, and biostratigraphy in the upper middle Devonian of Eastern North America: an ecofacies model for the Appalachian, Michigan, and Illinois basins. Unpublished Ph D thesis, Oregon State University, Oregon, pp 1–311Google Scholar
  37. Lamont A (1934) Brachiopod morphology in relation to environment. Cem Lime Gravel 8:216–219Google Scholar
  38. Leighton LR (1998) Constraining functional hypotheses: controls on the morphology of the concavo-convex brachiopod Rafinesquina. Lethaia 3:293–307Google Scholar
  39. Leighton LR (1999) Possible latitudinal predation gradient in middle Paleozoic oceans. Geology 27:47–50Google Scholar
  40. Leighton LR (2003) Predation on brachiopods. In: Kelley PH, Kowalewski M, Hansen TA (eds) Predator-prey interactions in the fossil record: topics in geobiology, vol 20. Kluwer/Plenum, New York, pp 215–237Google Scholar
  41. Lescinsky HL (1995) The life orientation of concavo-convex brachiopods: overturning the paradigm. Paleobiology 21: 520–551Google Scholar
  42. Macleod N, Forey PL (2002) Morphology, shape, and phylogeny. Taylor and Francis, New YorkGoogle Scholar
  43. Morris RW, Felton SH (1993) Symbiotic association of Crinoids, Platyceratid Gastropods, and Cornulites in the Upper Ordovician (Cincinnatian) of the Cincinnati, Ohio region. Palaios 8:465–476Google Scholar
  44. Morris RW, Felton SH (2003) Paleoecologic associations and secondary tiering of Cornulites on crinoids and bivalves in the Upper Ordovician (Cincinnatian) of southwestern Ohio, southeastern Indiana, and northern Kentucky. Palaios 18:546–558Google Scholar
  45. Pitrat CW, Rogers FS (1978) Spinocyrtia and its epizoans in the traverse group (Devonian) of Michigan. J Paleontol 52:1315–1324Google Scholar
  46. Richards RP (1969) Biology and ecology of Rafinesquina alternata (Emmons). Geol Soc Am Abstr Programs 6:41–42 (North Central Section)Google Scholar
  47. Richards RP, Shabica CW (1969) Cylindrical living burrows in Ordovician dalmanellid brachiopod beds. J Paleontol 43:838–841Google Scholar
  48. Richards RP (1972) Autecology of Richmondian brachiopods (Late Ordovician of Indiana and Ohio). J Paleontol 46:386–405Google Scholar
  49. Richards RP (1974) Ecology of the Cornulitidae. J Paleontol 48:514–523Google Scholar
  50. Rodland DL, Kowalewski M, Carroll M, Simoes MG (2004) Colonization of a ‘Lost World’: encrustation patterns in modern subtropical brachiopod assemblages. Palaios 19:381–395Google Scholar
  51. Rodrigues SC, Simoes MG, Kowalewski M, Petti MAV, Nonato EF, Martinez S, Rio CDJ (2008) Biotic interaction between spionid polychaetes and bouchardiid brachiopods: paleoecological, taphonomic and evolutionary implications. Acta Palaeontologica Polonica 53:657–668Google Scholar
  52. Rohlf FJ (1990) Fitting curves to outlines. In: Proceedings of the Michigan Morphometrics Workshop, pp 167–177Google Scholar
  53. Rohlf FJ, Slice DE (1990) Extensions of the Procrustes method for the optimal superimposition of landmarks. Syst Zool 39:40–59Google Scholar
  54. Rohlf FJ (1999) Shape statistics: procrustes superimpositions and tangent spaces. J Classif 16:197–223Google Scholar
  55. Rohlf FJ, Marcus LF (1993) A revolution in morphometrics. Trends Ecol Evol 8:129–132Google Scholar
  56. Rudwick MJS (1962) Filter-feeding mechanisms in some brachiopods from New Zealand. Zool J Linn Soc (Lond) 44:592–615Google Scholar
  57. Sandy MR (1996) Oldest record of peduncular attachment of brachiopods to crinoid stems, Upper Ordovician, Ohio, USA (Brachiopods; Atrypida: Echinodermata; Crinoidea). J Paleontol 70:532–534Google Scholar
  58. Schneider CL (2003) Hitchhiking on Pennsylvanian echinoids: epibionts on archaeocidaris. Palaios 18:435–444Google Scholar
  59. Schneider CL (2009a) Epibionts on Late Carboniferous through Early Permian echinoid spines from Texas, USA, Echinoderms 2006. In: Proceedings of the 12th international echinoderm conference, Durham, 7–11 August 2006, New Hampshire, U.S.A. CRC Press, Boca Raton, pp 1–679Google Scholar
  60. Schneider CL (2009b) Substrate preferences of Middle and Late Devonian Hederella from the midcontinent USA. In: Hageman SJ, Key MM Jr, Winston JE (eds) Bryozoan Studies 2007, Proceedings of the 14th IBA Conference, Boone, North Carolina, July 1–8, 2007, vol 15. Virginia Museum of Natural History Special Publication, Martinsville, pp 295–300Google Scholar
  61. Schneider CL, Leighton LR (2007) The influence of spiriferide microornament on Devonian epizoans. Geol Soc Am Abstr Programs 39:531Google Scholar
  62. Schneider CL, Webb A (2004) Where have all the encrusters gone? Encrusting organisms on Devonian versus Mississippian brachiopods. Geol Soc Am Abstr Programs 36:111Google Scholar
  63. Shou-Hsin C (1959) Note on the palaeoecological relation between Aulopora and Mucrospirifer. Acta Palaeontologica Sinica 7:502–504Google Scholar
  64. Slice DE (2001) Landmark coordinates aligned by procrustes analysis do not lie in Kendall’s shape space. Syst Biol 50:141–149Google Scholar
  65. Sparks DK, Hoare RD, Kesling RV (1980) Epizoans on the brachiopod Paraspirifer bowneckeri (Stewart) from the Middle Devonian of Ohio, University of Michigan Museum of Paleontology, Papers on Paleontology, vol 23, pp 1–50Google Scholar
  66. Spjeldnaes N (1984) Epifauna as a tool in autecological analysis of Silurian Brachiopods. The Spec Pap Palaeontol 32:225–235Google Scholar
  67. Stumm EC (1951) Check list of fossil invertebrates described from the middle Devonian traverse group of Michigan: Contributions from the Museum of Paleontology, vol 9. University of Michigan, Ann Arbor, pp 1–44Google Scholar
  68. Sumrall CD (2000) The biological implications of an edrioasteroid attached to a pleurocystitid rhombiferan. J Paleontol 74:67–71Google Scholar
  69. Taylor PD, Wilson MA (2002) A new terminology for marine organisms inhabiting hard substrates. Palaios 17:522–525Google Scholar
  70. Taylor PD, Wilson MA (2003) Palaeoecology and evolution of marine hard substrate communities. Earth Sci Rev 62:1–103Google Scholar
  71. Taylor PD, Wilson MA (2008) Morphology and affinities of hederelloid “bryozoans”. In: Hageman SJ, Key MM Jr, Winston JE (eds) Bryozoan Studies 2007: Proceedings of the 14th international bryozoology conference, Boone, North Carolina, July 1–8, 2007, vol 15. Virginia Museum of Natural History Special Publication, Martinsville, pp 301–309Google Scholar
  72. Thayer CW (1974) Substrate specificity of Devonian epizoan. J Paleontol 48:881–894Google Scholar
  73. Vermeij GJ (1977) The Mesozoic marine revolution: evidence from molluscs, predation, and grazing. Paleobiology 3:245–258Google Scholar
  74. Voros A (2010) Escalation reflected in ornamentation and diversity history of brachiopod clades during the Mesozoic marine revolution. Palaeogeogr Palaeoclimatol Palaeoecol 291:474–480Google Scholar
  75. Warthin AS, Cooper GA (1935) New formation names in the Michigan Devonian. J Wash Acad Sci 25:524–26Google Scholar
  76. Warthin AS, Cooper GA (1943) Traverse rocks of Thunder bay region, Michigan. Bull Am Assoc Petrol Geol 27:571–595Google Scholar
  77. Watkins R (1981) Epizoan ecology of the type Ludlow Series (Upper Silurian), England. J paleontol 55:29–32Google Scholar
  78. Webster CL (1921) Notes on the genus Atrypa, with description of new species. Am Midl Nat 7:13–26Google Scholar
  79. Wiedman LA (1985) Community paleoecological study of the silica shale equivalent of Northeastern Indiana. J Paleontol 59:160–182Google Scholar
  80. Wilson MA, Taylor PD (2001) “Pseudobryozoans” and the problem of encruster diversity in the Paleozoic. PaleoBios 21:134–135Google Scholar
  81. Wylie AS, Huntoon JE (2003) Log curve amplitude slicing—visualization of log data from the Devonian Traverse Group in the Michigan Basin, U.S. Bull Am Assoc Petrol Geol 87:581–608Google Scholar
  82. Zapalski MK (2005) Paleoecology of Auloporida: an example from the Devonian of the Holy Cross Mts., Poland. Geobios 38:677–683Google Scholar
  83. Zelditch ML, Swiderski DL, Sheets HD, Fink WL (2004) Geometric morphometrics for biologists, a primer. Elsevier Academic Press, New YorkGoogle Scholar
  84. Zhan R, Vinn O (2007) Cornulitid epibionts on brachiopod shells from the Late Ordovician (middle Ashgill) of East China. Estonian J Earth Sci 56:101–108Google Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2013

Authors and Affiliations

  1. 1.City College of New YorkNew YorkUSA

Personalised recommendations