Possible reasons for survival of proetidan trilobites past the Devonian

[DD1991]

It's well-known that extinction events at the end of the Devonian wiped out all trilobite groups except Proetida, which survived until the mass extinction at the end of the Permian. Has there been research in recent years as to why proetidan trilobites outlived the Devonian?

[Kane]

Yes and we will let @piranha provide you with resources. In the interim, understanding the ecological niches, and their transformations, will be of some benefit.

[piranha]

Google Scholar will yield many more papers on the extinction of Cambrian and Ordovician trilobites.

 

In the meantime, these are suggested references that focus on the extinction of Devonian trilobites.

 

A little light reading for the weekend—Enjoy! 

 

 

Bault, V., Balseiro, D., Monnet, C., Crônier, C. 2022

Post-Ordovician Trilobite Diversity and Evolutionary Faunas.

Earth-Science Reviews, 230 (104035)

 

Bault, V., Crônier, C., Allaire, N., Monnet, C. 2021

Trilobite biodiversity trends in the Devonian of North Africa.

Palaeogeography, Palaeoclimatology, Palaeoecology, 565(110208)

 

Becker, R.T., Feist, R., Flajs, G., House, M.R., Klapper, G. 1989
Frasnian-Famennian Extinction Events in the Devonian at Coumiac, Southern France. 
Comptes Rendus de l'Académie des Sciences, Série 2, Stratigraphy, 309(2):259-266

 

Bond, D.P.G., Grasby, S.E. 2017
On the Causes of Mass Extinctions.
Palaeogeography, Palaeoclimatology, Palaeoecology 478:3-29

 

Brauckmann, C., Brauckmann, B. 1986
Famennian Trilobites: an Outline of their Stratigraphical Importance. 
Annales de la Société Géologique de Belgique, 109(1):9-17 

 

Brauckmann, C., Chlupáč, I., Feist, R. 1993
Trilobites at the Devonian-Carboniferous Boundary.
Annales de la Société Géologique de Belgique, 115(2):507-518

 

Chlupáč, I. 1975
The Distribution of Phacopid Trilobites in Space and Time.

Fossils and Strata, 4:399-408

 

Chlupáč, I. 1994
Devonian Trilobites - Evolution and Events.

Geobios, 27(4):487-505

 

Chlupáč, I., Feist, R., Morzadec, P. 2000
Trilobites and Standard Devonian Stage Boundaries.
Courier Forschungsinstitut Senckenberg, 220:87-98

 

Clarkson, E.N.K. 2013
Grzimek's Animal Life Encyclopedia: Extinction. Trilobites.

Gale Research Inc. [pp. 235-244]

 

Copper, P. 1977
Paleolatitudes in the Devonian of Brazil and the Frasnian - Famennian Mass Extinction. 
Palaeogeography, Palaeoclimatology, Palaeoecology, 21(3):165-207

 

Crônier, C., Van Viersen, A.P. 2007
Trilobite Palaeobiodiversity during the Devonian in the Ardennes Massif. 
[Paleobiodiversite des Trilobites en Ardenne Occidentale au Devonien.]
Bulletin de la Société Géologique de France, Série 8, 178(6):473-483

 

Erben, H.K. 1958
Blinding and Extinction of Certain Proetidae (Tril.).

Journal of the Palaeontological Society of India, 3:82-104

 

Feist, R. 1991
The Late Devonian Trilobite Crises.

Historical Biology, 5(2-4):197-214

 

Feist, R. 1995
Effect of Paedomorphosis in Eye Reduction on Patterns of Evolution and Extinction in Trilobites. 
In: Evolutionary Change and Heterochrony. John Wiley, Sons Ltd. Publishing, 298 pp. [pp. 225-244]

 

Feist, R. 1997
Trilobite Extinctions and Recovery at the Late Devonian Kellwasser and Hangenberg Crises.
2nd International Trilobite Conference (Brock University, St. Catharines, Ontario, August 22-24, 1997) Abstracts Volume.

 

Feist, R. 2000
The Frasnian - Famennian Boundary Extinction Event in North Gondwanan Trilobites.  

In: XVI Jornadas de Paleontología & I Congresso Ibérico de Paleontología, Evora (Portugal) p. 214 

 

Feist, R., Clarkson, E.N.K. 1989
Environmentally Controlled Phyletic Evolution, Blindness and Extinction in Late Devonian Tropidocoryphine Trilobites.

Lethaia, 22(4):359-373

 

Feist, R., McNamara, K.J. 2007
Biodiversity, Distribution and Patterns of Extinction of the Last Odontopleuroid Trilobites during the Devonian (Givetian, Frasnian).

Geological Magazine, 144(5):777-796

 

Feist, R., McNamara, K.J. 2013

Patterns of Evolution and Extinction in Proetid Trilobites during the Late Devonian Mass Extinction Event, Canning Basin, Western Australia.

Palaeontology, 56(2):229-259

 

Feist, R., McNamara, K.J., Crônier, C., Lerosey-Aubril, 2009

Patterns of Extinction and Recovery of Phacopid Trilobites during the Frasnian-Famennian (Late Devonian) Mass Extinction Rvent, Canning Basin, Western Australia.

Geological Magazine, 146(1):12-33

 

Feist, R., Petersen, M.S. 1995
Origin and Spread of Pudoproetus, A Survivor of the Late Devonian Trilobite Crisis.
Journal of Paleontology, 69(1):99-109

 

Feist, R., Schindler, E. 1994
Trilobites during the Frasnian Kellwasser Crisis in European Late Devonian Cephalopod Limestones.
Courier Forschungsinstitut Senckenberg, 169:195-223 

 

Hahn, G., Hahn, R., Brauckmann, C. 1994
Trilobiten mit "Drevermannia-Habitus" im Unter-Karbon.
[Trilobites of "Drevermannia Habit" from the Lower Carboniferous.]
Courier Forschungsinstitut Senckenberg, 169:155-193

 

House, M.R. 2002.

Strength, Timing, Setting and Cause of Mid-Palaeozoic Extinctions.

Palaeogeography, Palaeoclimatology, Palaeoecology, 181(1-3), pp.5-25.

 

Kaiser, S.I., Aretz, M., Becker, R.T. 2016
The Global Hangenberg Crisis (Devonian–Carboniferous Transition): Review of a First-Order Mass Extinction.
Geological Society of London, Special Publications, 423:387-437

 

Kaiser, S.I., Becker, R.T., Steuber, T., Aboussalam, S.Z. 2011
Climate-Controlled Mass Extinctions, Facies, and Sea-Level Changes around the Devonian-Carboniferous Boundary in the Eastern Anti-Atlas (SE Morocco).

Palaeogeography, Palaeoclimatology, Palaeoecology 310(3-4):340-364

 

Lerosey-Aubril, R., Feist, R. 2012

Quantitative Approach to Diversity and Decline in Late Palaeozoic Trilobites. [pp. 535-555]

In: Earth and Life: Global Biodiversity, Extinction Intervals and Biogeographic Perturbations through Time. Springer Publishing

 

Lieberman, B.S. 1994
Evolution of the Trilobite Subfamily Proetinae Salter, 1864, and the Origin, Diversification, 

Evolutionary Affinity, and Extinction of the Middle Devonian Proetid Fauna of Eastern North America. 

American Museum of Natural History, Bulletin, 223:1-176 

 

McGhee, G.R. 1996

The Late Devonian Mass Extinction: The Frasnian / Famennian Crisis.

New York: Columbia University Press, 378 pp.

 

McGhee, G.R. 2013
Grzimek's Animal Life Encyclopedia: Extinction. The Late Devonian and End-Devonian Extinctions.
Gale Research Inc. [pp. 449-457]

 

McNamara, K.J., Feist, R. 2008
Patterns of Trilobite Evolution and Extinction During the Frasnian/Famennian Mass Extinction, Canning Basin, Western Australia.

Cuadernos del Museo Geominero, 9:269-274

 

McNamara K.J., Feist, R. 2016

The Effect of Environmental Changes on the Evolution and Extinction of Late Devonian Trilobites from the Northern Canning Basin, Western Australia.

Geological Society, London, Special Publications, 423:251-271

 

McNamara, K.J., Feist, R., Ebach, M.C. 2009

Patterns of Evolution and Extinction in the Last Harpetid Trilobites During the Late Devonian (Frasnian).

Palaeontology, 52(1):11-33

 

Owens, R.M. 1994
Pseudoextinctions in Late Devonian Proetide Trilobites.

Historical Biology, 9(3):207-221

 

Talent, J.A. 2012
Earth and Life: Global Biodiversity, Extinction Intervals and Biogeographic Perturbations Through Time.
Springer Scientific Publishing, 1100 pp.

 

Twitchett, R.J. 2005
The Palaeoclimatology, Palaeoecology and Palaeoenvironmental Analysis of Mass Extinction Events.
Palaeogeography, Palaeoclimatology, Palaeoecology, 232(2-4):190-213

 

Wagnera, P.J., Estabrook, G.F. 2014
Trait-Based Diversification Shifts Reflect Differential Extinction Among Fossil Taxa.
Proceedings of the National Academy of Sciences, USA. 111(46):16419-16424

 

Yuan, J., Xiang, L. 1998
Trilobite Fauna at the Devonian-Carboniferous Boundary in South China (S-Guizhou and N-Guangxi).
National Museum of Natural Science, Special Publication, 8:1-281

[Coco]

Hi,

 

Woh ! All this ! Enjoy reading !

 

Coco

[piranha]

 

McGhee, G.R. 2013
The Late Devonian and End-Devonian Extinctions.
Grzimek's Animal Life Encyclopedia: Extinction. [pp. 449-457]

[piranha]

"The terminal Devonian Hangenberg Extinction Event had an extremely severe effect on trilobite evolution, as demonstrated by extermination of the
order Phacopida and the high level of generic extinctions, comparable only to those observed during the Middle Devonian events (Fig. 6). The reduced-eyed and blind taxa among the Proetida became extinct without exception, most likely trapped by the sudden rise in sea level and the spread of oceanic anoxia. The survivors probably originated from conservative, normally sighted communities in shallow shoal refugia that were beyond the devastating effects of the oxygen depleted conditions at the onset of the Hangenberg black shale. The Hangenberg Event recalls the terminal Frasnian mass extinction in terms of both survivorship and post-event recovery. However, unlike the situation after the Upper Kellwasser Event, postevent recovery was immediate and worldwide after the Hangenberg biocrisis, even in North America where Pudoproetus originated in the Late praesulcata Zone before the defined D/C boundary (Feist and Petersen 1995). The total worldwide absence of reduced-eyed trilobites in this terminal Devonian period may indicate a considerable eustatic shallowing. The first phillipsiids, representatives of the subfamilies Archegoninae and Weaniinae, developed contemporaneously at that time and crossed the D/C boundary without any loss before their main radiations in the Early Carboniferous (Brauckmann et al. 1993). A major biotic turnover in trilobite diversity was associated with the Hangenberg Event, i.e. shortly before the internationally defined D/C boundary. The subsequent D/C lineages have yet to be defined within the continuously radiating trilobite lineages of ‘Carboniferous aspect’ (Chlupáč et al. 2000)."

 

text from:

 

Lerosey-Aubril, R., Feist, R. 2012

Quantitative Approach to Diversity and Decline in Late Palaeozoic Trilobites. [pp. 535-555]

In: Earth and Life: Global Biodiversity, Extinction Intervals and Biogeographic Perturbations through Time. Springer Publishing.  PDF LINK

 

 

 

"Near the Devonian/Carboniferous boundary, another important biotic turnover linked to anoxia/dysoxia and sea-level fall occurred (Walliser, 1996; House, 2002; Kaiser et al., 2016; Zhang et al., 2020). Trilobites were strongly affected by this event (Fig. 2), with the disappearance of the order Phacopida as well as blind and reduced-eye Proetida trilobites (Lerosey-Aubril and Feist, 2012). The Hangenberg event was lethal for both shallow and deep-water trilobites (Chlupáč, 1994), but this event had a stronger impact on deeper water trilobites and consequently, survivors mostly lived in shallower areas (Feist and Petersen, 1995). Important faunal changes coincided with this event (Brauckmann et al., 1993). The Kellwasser Fauna almost disappeared with the Hangenberg event at the end of the Devonian, resulting in the extinction of phacopids (Lerosey-Aubril and Feist, 2012) and the decline of cyrtosymbolines, so that this subfamily was considered extinct for a long time (Feist, 1991). Climate cooling and glaciation at the Devonian-Carboniferous boundary (Isaacson et al., 2008; Kaiser et al., 2016) affected more severely a fauna accustomed to warmer temperatures."

 

"The Kellwasser Fauna had the shortest duration, being restricted to the Late Devonian and Tournaisian. This is the only evolutionary fauna associated with a low total diversity, because Middle Devonian environmental changes decimated the previous evolutionary faunas. Dominated by Phacopidae and Cyrtosymbolinae, this evolutionary fauna invaded both shallow and deep environments. The taxa of this fauna are characterized by important morphological changes such as the eye-reduction. At that time, trilobites inhabited warm, carbonate environments. The gradual cooling that began in the Famennian and the Hangenberg event at the Devonian–Carboniferous boundary led to the disappearance of this evolutionary fauna."    

 

"It is complicated to know whether predation, resource competition or asymmetrical effect of Devonian events had an influence, but faunal changes occurring in the Devonian–Permian had an evident consequence on trilobite macroevolution."

 

text from:

 

Bault, V., Balseiro, D., Monnet, C., Crônier, C. 2022

Post-Ordovician Trilobite Diversity and Evolutionary Faunas.

Earth-Science Reviews, 230 (104035):1-20

[piranha]

"Trilobite extinction and survival patterns associated with the Hangenberg Crisis have been reviewed by Brauckmann & Brauckmann (1986), Hahn (1990), Brauckmann et al. (1993), Hahn et al. (1994) and Chlupáč et al. (2000). No species is known to have passed through the complete crisis interval, and there are only a few surviving genera within the Proetida, represented by the Brachymetopidae (a neritic family), Proetidae (Drevermanniinae) and Phillipsiidae (with two surviving subfamilies, the Archegoninae = Cyrtosymbolinae, but alternatively placed in the Proetidae, and Weaniinae). Therefore, the re-radiation must have come from survivors within those groups, which evolved within still unknown refugia areas. A re-dispersal of descendent species began in the upper (kockeli Zone) and/or post-crisis interval. There is an open debate whether some forms, such as Pudoproetus, represent Elvis Taxa homoeomorphic with Middle Givetian genera (in this case of the Proetinae), or whether they indicate a survival in refugia areas for more than 15 Ma (see Feist & Petersen 1995). The second interpretation (e.g. Yuan & Xiang 1998; Hahn et al. 2012) implies an enigmatic prevention of recovery and spread in practically all of the Upper Devonian (UD) and an even more enigmatic palaeoecological factor that enabled a sudden spread to North America, North Africa, Europe, Central Asia and South China at the end of the crisis interval. A similar discussion is required to clarify whether there was true survival of proetid genera or whether Carboniferous taxa represent iterative homoemorphs, e.g. in the case of the Famennian Drevermannia (Drevermannia) and the Carboniferous Dr. (Pseudodrevermannia) and Dr. (Paradrevermannia) (see Gandl et al. 2015).

 

The main trilobite extinction was equally severe in western (Europe–North Africa) and eastern (South China) Prototethys regions (see Yuan & Xiang 1998). In the pelagic realm the last Phacopida, represented by the blind Dianops, the small-eyed Weyerites, and the genus Rabienops (‘Phacops’ granulatus Group) with median-sized eyes, died out at the onset of the HBS. The shallow-water genus Omegops, however, obviously survived in Xinjiang briefly into the lower/middle crisis interval, based on co-occurrences with miospores of the LN Zone and the youngest Cymaclymenia faunas (Zong et al. 2012, 2014). Based on geochemistry, Carmichael et al. (2015) placed the HBS level in the Bulongguor type section of the Junggar Basin much below the phacopid extinction. This questions the assumption that Omegops specimens from the kockeli Zone of the Ardennes (Conil et al. 1986) are reworked. The globally youngest phacopid is a probably reworked specimen from the sulcata Zone of the DCB stratotype (Flajs & Feist 1988). It seems likely that the final demise of the order was a case of ‘survivor extinction’, as in the case of the contemporaneous clymeniids. So far there is no explanation as to why the widespread phacopids died out in the neritic realm whilst the associated Brachymetopidae managed to survive in widely separate regions of the Rhenish Massif (e.g. Michels 1986) and South China (Yuan & Xiang 1998). The post-crisis recovery was very fast in both magnafacies and led to the appearance of many new proetid subfamilies and genera (e.g. Brauckmann et al. 1993)."

 

text from:

 

Kaiser, S.I., Aretz, M., Becker, R.T. 2016
The Global Hangenberg Crisis (Devonian–Carboniferous Transition): Review of a First-Order Mass Extinction.
Geological Society of London, Special Publications, 423:387-437  PDF LINK