Age of Crinoids”: a Mississippian Biodiversity Spike Coincident with Widespread Carbonate Ramps

THE “AGE OF CRINOIDS”: A MISSISSIPPIAN BIODIVERSITY SPIKE COINCIDENT WITH WIDESPREAD CARBONATE RAMPS

Thomas W. Kammer Department of Geology and Geography West Virginia University www.geo.wvu.edu Palaios, June 2006

Kammer, T.W. and Ausich, W.I. 2006. The “Age of Crinoids”: A Mississippian Biodiversity Spike Coincident with Widespread Carbonate Ramps. Palaios, v. 21, p. 238-248. Key Points of this Talk

• Key Point 1 – Crinoids reached their maximum generic richness during the Mississippian. • Key Point 2 – During the Mississippian the transition between the Middle Paleozoic and Late Paleozoic macroevolutionary faunas occurred. • Key Point 3 – The evolutionary replacement of the camerates by the advanced cladids. One pinnulate clade replaces another. • Key Point 4 – Crinoids were abundant as evidenced by widespread global encrinites. • Key Point 5 – The Late Devonian (F-F) extinction event wiped out reef ecosystems. • Key Point 6 – Carbonate ramps replaced carbonate platforms, increasing habitat space for stenohaline crinoids. Echinoderms: 21 classes (16 extinct) including:

• Crinoids •Blastoids • Cystoids • Edrioasteroids •Asteroids • Ophiuroids • Echinoids • Holothurians Crinoid Blastoid Cystoid Edrioasteroid Asteroid Ophiuroid Echinoids: sand dollar (left) sea biscuit (below) Holothurian: sea cucumber Diversity of crinoid genera over geologic time. High modern diversity is a taphonomic artifact (Broadhead and Waters, 1980). Classification of Crinoids

• Paleozoic Crinoids – Subclass Camerata: pinnulate arms – Subclass Disparida: nonpinnulate arms – Subclass Cladida: nonpinnulate and pinnulate – Subclass Flexibilia: nonpinnulate arms • Mesozoic and Cenozoic Crinoids – Subclass Articulata: pinnulate arms • Isocrinids - stalked • Comatulids - free living Encrinus, the survivor of the Permian extinction event, founder of the Articulata Endoxocrinus at a depth of 692 m, Bahamas

Photo by Chuck Messing Endoxocrinus, 430 m, Bahamas

Photo by Chuck Messing Neocrinus, 424 m, Bahamas

Photo by Chuck Messing Neocrinus, 424 m, Bahamas. Current = 20cm/s

Photo by Chuck Messing Tube feet on the pinnules of crinoid arms

Photo by Dave Meyer Modern comatulid crinoids, Bahamas

Photo by Dave Meyer Modern comatulid crinoids, Bahamas. They can tolerate salinities between 24-36‰.

Photo by Dave Meyer Classification of Crinoids

• Paleozoic Crinoids – Subclass Camerata: pinnulate arms – Subclass Disparida: nonpinnulate arms – Subclass Cladida: nonpinnulate and pinnulate, or primitive and advanced cladids – Subclass Flexibilia: nonpinnulate arms • Mesozoic and Cenozoic Crinoids – Subclass Articulata: pinnulate arms • Isocrinids - stalked • Comatulids - free living Examples of Mississippian Crinoid Clades

A. Camerate crinoid with pinnulate arms. Abatocrinus grandis

B. Disparid crinoid with nonpinnulate arms. Synbathocrinus swallovi

C. Advanced cladid crinoid with pinnulate arms. Decadocrinus tumidulus

D. Primitive cladid crinoid with non-pinnulate arms. Cyathocrinites barydactylus

E. Flexible crinoid with nonpinnulate arms. Onychocrinus ulrichi Actinocrinites: a camerate; note the pinnulate arms Cyathocrinites, a primitive cladid with nonpinnulate arms Actinocrinites and the primitive cladid Barycrinus

Sepkoski, J.J., Jr. 2002. A Compendium of Fossil Marine Animal Genera. Bulletins of American Paleontology, No. 363, 560 pages.

Paleozoic Generic Diversity by Clade from Sepkoski, 2002

Loch Prag Emsi Eife Give Fras Fame Tour Vise Serp Bash Mosc Step Advanced Cladids 0 0 4 3 8 11 8 34 76 53 50 100 89 Camerates 22 19 46 42 33 14 9 49 52 14 9 13 6 Disparids 13 15 22 21 14 7 6 9 9 5 5 6 5 Primitive Cladids 16 20 32 27 25 14 9 16 22 8 11 11 11 Flexibles 9 6 5 9 11 9 8 13 16 5 5 6 6

Totals 60 60 109 102 91 55 40 121 175 85 80 136 117 Devonian Mississippian Pennsylvanian Example of Database Entries First Occurrence Last Occurrence ADVANCED CLADIDS Aaglacrinus C (Mosc-u) C (Step-u) Aatocrinus C (Mosc-u) P (Wc) Abrotocrinus C (Tour-l) C (Vise-l) Acylocrinus C (Tour-u) Adacrinus C (Mosc-u) C (Step-u) ADIAKRITOCRINUS C TOUR Adinocrinus C VISE Aenigmocrinus C (Vise-u) Aesiocrinus C (Bash) P WC Affinocrinus C (Bash-u) C (Mosc-l) Agassizocrinus C (Vise-u) C (Serp-l) Aglaocrinus C (Bash) C STEP Alcimocrinus C (Serp-l) C (Mosc-l) Allosocrinus C (Mosc-u) P WC AMABILICRINUS D FAME C TOUR AMADEUSICRINUS D FAME Ampelocrinus C (Vise-u) C (Serp-l) AMPULLACRINUS C SERP Anartiocrinus C (Vise-u) C (Serp-l) Anchicrinus C (Bash-l) C (Mosc-u) Update of Sepkoski’s database based on the literature, 1996-2004. Comparison of total generic richness by stage.

200

180

160

140

120 a r Updated, 2005 100 ne Sepkoski, 2002 Ge 80

h g i e e s r e p p c a f v a u s r sh sc me a e Pr Ei Gi Fr To Vi Se St Lo Ems Fa B Mo Key Point 1 – Crinoids reached their maximum generic richness during the Mississippian. This diversity peak is bracketed between the Late Devonian (F-F) extinction event and the Late Mississippian (Serpukhovian) extinction event (Stanley and Powell, 2003). There are five major clades of Paleozoic crinoids that make up this diversity peak. Middle Paleozoic Crinoid Fauna Late Paleozoic Crinoid Fauna Miss. 180 160 140 120 Advanced Cladids

a Camerates r 100 e Primitive Cladids n 80 Disparids Ge 60 Flexibles 40 20 0

h g e e s e p h c p si f se s s oc a m iv ra m our i er e Ei G F T V St L Pr E Fa S Ba Mo Key Point 2 – During the Mississippian the transition between the Middle Paleozoic and Late Paleozoic macroevolutionary faunas occurred as the advanced cladids replaced the camerates as the dominant clade (Ausich et al., 1994; Baumiller, 1994). The overlap of these two macroevolutionary faunas is a primary factor in the high crinoid generic richness. Middle Paleozoic Crinoid Fauna Late Paleozoic Crinoid Fauna

100% Miss.

80% Advanced Cladids

60% Camerates a r

e Primitive Cladids n Disparids Ge 40% Flexibles

20%

h g i e e s e r e p c s f v u s r sh s oc ra ra m i a ep Ei Gi F To V o St L P Em Fa Se B M Key Point 3 - The evolutionary replacement of the camerates by the advanced cladids. One pinnulate clade replaces another pinnulate clade. The muscular articulations of the advanced cladid arms were apparently superior to the ligamentary articulations of camerate arms. Miss. Miss. 100% 140

90% 120 80%

70% 100

60% Advanced Cladids 80 a a r r Camerates e

50% ne n Ge

Ge 60 40%

30% 40

20% 20 10%

0% 0

h g e e s e e h p h g e e s e r e h p c si f a ur s rp s sc c si v u s rp s sc e ra m iv m te ra Eif ra e P Ei G Fr To Vi Se S Lo P Gi F am To Vi S St Lo E Fa Ba Mo Em F Ba Mo Muscular articulations in modern crinoid arms

Macurda et al., 1978 Van Sant, 1964 Arm articulations in Mississippian crinoids

Camerate: Platycrinites Advanced Cladid: Abrotocrinites

Van Sant, 1964 Van Sant, 1964 Ligaments only (mutable collagen?) Muscles and Ligaments Key Point 4 Crinoids were not only diverse during the Mississippian, they were also abundant as shown by widespread global encrinites, defined as >50% crinoidal packstones and grainstones. For example, the encrinital Burlington and Keokuk limestones of North America extend over 74,000 km2 (Ausich, 1997). Encrinital limestones are found throughout the Mississippian on multiple continents Age of Crinoids

Restored The Mississippian was also a time Present record of unusually high carbonate production and accumulation.

Walker et al., 2002, J. Geol., 110:75. Tournaisian and Visean carbonate ramp. Limestone bluffs along the Mississippi River in Illinois. Tournaisian and Visean carbonate ramp. Burlington and Keokuk limestones, Iowa. Visean crinoidal bioherm, Fort Payne Fm., Kentucky Visean crinoidal bioherm, Fort Payne Fm., KY Crinoid calyces in the Fort Payne Fm. Visean advanced cladid crinoids on a carbonate ramp, Main Ls., Richmond, England Tournaisian encrinite, Clitheroe, England Tournaisian encrinite, Clitheroe, England Crinoid calyces, Clitheroe, England We had to set a limit on how many crinoids we could collect. Tournaisian bioherm and ramp, Pembroke, Wales Tournaisian encrinite, Pembroke, Wales Castlemartin Tank Firing Range, British Army, Pembroke, Wales - 2003

Tournaisian ramp, Hook Head, Ireland Tournaisian encrinite, Hook Head, Ireland Tournaisian encrinite, Hook Head, Ireland Key Point 5 The Late Devonian (F-F) extinction event wiped out reef ecosystems (Copper, 1994, 2002). This resulted in widespread carbonate ramps during the Mississippian (Ahr, 1989; Wright and Faulkner, 1990). Key Point 6 The open circulation of ramps, versus the more restricted circulation on platforms, would have maintained stenohaline conditions over much of the ramp. Modern comatulid crinoids are strictly stenohaline and are absent from carbonate platforms that experience summertime hypersalinities (Clark, 1921; Fell, 1966). Key Point 6 (cont’d) Modern crinoids can tolerate salinities between 24-36%o (Clark, 1921). Summertime salinities on the west side of Andros Island reach 45%o (Bathurst, 1976).

Smith, 1940 Reefs Circulation on a carbonate platform versus carbonate ramp. (A) Platform-edge reef system restricts circulation on the inner platform; evaporation increases salinity. (B) Carbonate ramp lacks a platform-edge reef system resulting in more open circulation and stenohaline conditions. FWB: fair- weather wave base. 10’s - 100’s km

~ 40 m

~ 40 m Summary The “Age of Crinoids” resulted from a biodiversity spike associated with the overlap of two macroevolutionary faunas as the camerates and advanced cladids were co- dominant, and from the simultaneous occurrence of widespread carbonate ramps that provided an extensive habitat for crinoids. London 2004