The Late De vo nian (Frasnian/Famenian) mass ex tinc tion:
a pro posed test of the gla ci ation hy poth e sis
George R. Jr. McGHEE1, *
1 De part ment of Earth and Plan e tary Sci ences, Wright-Rieman Lab o ra to ries, Rutgers Uni ver sity, New Bruns wick, New Jer sey 08903, USA
Mc Ghee, G.R. Jr., 2014. The Late De vo nian (Frasnian/Famenian) mass ex tinc tion: a pro posed test of the gla ci ation hy poth - e sis. Geo log i cal Quar terly, 58 (2): 263–268, doi: 10.7306/gq.1143
In this pa per it is agued that Late Frasnian global cool ing was the first step in the on set of the Late Pa leo zoic Ice Age, and that a Late Frasnian gla ci ation ex isted that was anal o gous to the early Oligocene (Oi-1) gla ci ation that took place in the first step of global cool ing in the on set of the Ce no zoic Ice Age. It is here pro posed that de fin i tive sedimentological ev i dence for the ex - is tence of Late Frasnian gla ci ation, like the Oi-1 gla ci ation, would be the dis cov ery of ice-rafted de bris in ma rine sed i ments of Late Frasnian age sim i lar to the ice-rafted de bris found in Oi-1 ma rine sed i ments (Zachos et al., 1992; Ehrmann and Mackensen, 1992). Rather than metre- to decimetre-sized ice-rafted de bris, a search should be ini ti ated for the pres ence of sand-sized (>250 mm) ice-rafted de bris in Frasnian ma rine strata lo cated off shore from the main Gond wana land mass, and dated to the Late rhenana Cono dont Zone and/or to the linguiformis–Early triangularis cono dont zonal in ter val where the ex - tinc tions and sharp drops in sea-sur face tem per a ture oc curred (Joachimski and Buggisch, 2002).
Key words: mass ex tinc tion, gla ci ation, Late Pa leo zoic Ice Age, Ce no zoic Ice Age.
INTRODUCTION
The cause of the Late De vo nian (Frasnian/Famennian) ex - tinc tion re mains con tro ver sial. Over 36 years of amassed em pir i - cal bi o log i cal data have been used to ar gue for a causal link be - tween global cool ing and the Frasnian/Famennian ex tinc tion (Ta ble 1). How ever, the cause of global cool ing in the Late Frasnian re mains un proved. Cat a strophic causal hy poth e ses have in cluded as ter oid im pacts (McGhee, 2001, 2005) and man - tle-plume vol ca nism (Racki, 1998; Courtillot and Renne, 2003;
Courtillot et al., 2010). Non-cat a strophic, lon ger-term causal hy - poth e ses have in cluded global cool ing pro duced by at mo spheric car bon-di ox ide downdraw by bi o log i cal weath er ing pro cesses (Algeo et al., 2001) or by the chem i cal weath er ing of tec toni - cally-ac tive moun tain ranges (Averbuch et al., 2005).
Global-cool ing pro duced gla ci ation has long been pro posed to have been a trig ger for the Frasnian/Famennian ex tinc tion (for an ex ten sive dis cus sion of pro posed causes of the Late De - vo nian ex tinc tion see McGhee, 1996, 2013). How ever, de - cades of search ing on a world-wide ba sis has failed to un cover hard phys i cal ev i dence for gla ci ation in the Late Frasnian.
Only two ice house in ter vals ex ist in the Phanerozoic in which gla ci ation per sisted for tens of mil lions of years: the Late Pa leo zoic Ice Age and the Ce no zoic Ice Age (Field ing et al., 2008). It is well es tab lished that the de vel op ment of the glaciations of the Ce no zoic Ice Age took place in three steps in geo logic time: the Early Oligocene, Mid dle Mio cene, and Late Plio cene (Lewis et al., 2008). It has re cently been pro posed that the on set of the Late Pa leo zoic Ice Age also took place in three steps: the Late Frasnian, Late Famennian, and Late Visean–Serpukhovian (Barham et al., 2012; McGhee, 2013).
Such a sce nario re quires gla ci ation in the Late Frasnian, and the ob ject of the pres ent pa per is to pro pose a de fin i tive test of the hy poth e sis of Late Frasnian gla ci ation.
A COMPARISON OF CENOZOIC AND LATE PALEOZOIC GLOBAL COOLING
The de vel op ment of the glaciations of the Ce no zoic Ice Age took place in three steps in geo logic time: the Early Oligocene, Mid dle Mio cene, and Late Plio cene (Lewis et al., 2008). The first step in the Ce no zoic Ice Age, the Early Oligocene Oi-1 gla - cial pulse (Miller et al., 1991), took place be tween 34 and 33 Ma and lasted for 400,000 years. The on set of the Early Oligocene glaciations co in cided with two sep a rate pulses of ex tinc tion that oc curred in the oceans at about 33 Ma, and a third pulse of ex - tinc tion that oc curred on land at about 32 Ma (Prothero, 1994;
McGhee, 2001; Prothero et al., 2003). The Oi-1 gla cial pulse ini - ti ated the for ma tion of ice sheets in east ern Antarctica, gla ciers
* E-mail: mcghee@rci.rutgers.edu
Received: October 11, 2013; accepted: January 12, 2014; first published online: January 13, 2014
that per sisted for about eight mil lion years (Zachos et al., 2001) be fore re treat ing dur ing a warm ing trend that be gan in the Late Oligocene and ex tended into the Mio cene.
The sec ond step in the Ce no zoic Ice Age was ini ti ated by a sec ond gla cial pulse, the Mi-1, that took place in the Early Mio - cene and lasted for 200,000 years (Zachos et al., 2001). The Mi-1 gla cial pulse ini ti ated a se ries of brief and smaller glaciations in Antarctica and then the Earth be gan to cool rap - idly in the Mid dle Mio cene. The on set of the ma jor ex pan sion of the East Ant arc tic Ice Sheet be tween 14 and 13 Ma (Flower and Ken nett, 1995) trig gered pulses of ex tinc tion in both the ma rine and ter res trial realms dur ing this in ter val of time (Sepkoski, 1996; Lewis et al., 2008). By 12 Ma mas sive ice sheets had formed in east ern Antarctica that per sist to the pres ent day.
The third step in the Ce no zoic Ice Age took place in the Late Plio cene when gla ci ation be came bi po lar, with for ma tion of per - sis tent ice sheets in the north ern hemi sphere of the Earth in ad - di tion to those in the south ern hemi sphere (Zachos et al., 2001). The on set of gla ci ation in the north ern hemi sphere co in - cided with ad di tional ex tinc tion pulses first in the ma rine realm
(Hay ward, 2002) and then later in the ter res trial realm (Barnes et al., 1996).
It has re cently been pro posed that the on set of the Late Pa - leo zoic Ice Age also took place in three steps: the Late Frasnian, Late Famennian, and Late Visean–Serpukhovian (Barham et al., 2012; McGhee, 2013). The pro posed first step in the Late Pa leo zoic Ice Age, Late Frasnian cool ing, took place in two pulses: the first pulse took place at about 376 Ma, is dated to the Late rhenana Cono dont Zone in the Lower Kellwasser strati graphic ho ri zon (Walliser, 1996; Joachimski and Buggisch, 2002). The sec ond cool ing pulse took place at about 375 Ma in the lat est Frasnian, is dated to the linguiformis Cono dont Zone at the Up per Kellwasser strati graphic ho ri zon (Walliser, 1996; Joachimski and Buggisch, 2002). Both of the Kellwasser strati graphic ho ri zons are also as so ci ated with ex - tinc tion pulses: ex tinc tions dated to the Late rhenana Cono dont Zone oc curred in both the ma rine and ter res trial realms (Ray - mond and Metz, 1995; McGhee, 1996), and fur ther ex tinc tions dated to the linguiformis Cono dont Zone oc curred in the ma rine realm (Schindler, 1993; McGhee, 1996).
The pro posed sec ond step in the Late Pa leo zoic Ice Age, Late Famennian cool ing, is ev i denced by gla cial sed i ments in Gond wana that have been dated to the VCo spore zone on land, and cor re lated to the Late postera Cono dont Zone in the sea (Isaacson et al., 2008). The gla ciers in Gond wana reached their max i mum ex pan sion phase in the lat est Famennian, dated to the LE–LN spore zonal in ter val on land (Streel et al., 2000a;
Caputo et al., 2008), and cor re lated to the Mid dle–Late praesulcata cono dont zonal in ter val in the sea (Isaacson et al., 2008). At the same strati graphic ho ri zon ice-rafted gla cial de - bris, in some cases metre-sized dropstones (Brezinski et al., 2010: fig. 10G), are found in off shore ma rine de pos its of the Cleve land Shale in Ken tucky (Ettensohn et al., 2007). This ice-rafted de bris ap par ently came from con ti nen tal gla ciers in the Ap pa la chian moun tains to the east, a re gion that was as close to the equa tor as 30°S and thus is ev i dence of the in ten - sity of the Late Famennian gla cial cool ing (Brezinski et al., 2009, 2010). Ex tinc tions as so ci ated with the max i mum ex pan - sion phase of the Famennian gla ci ation took place in two pulses: the first pulse oc curred in the ma rine realm at about 360 Ma in the Late Famennian, is dated to the Mid dle praesulcata Cono dont Zone at the Hangenberg Black Shale ho ri zon (Kai ser et al., 2006). The sec ond pulse of ex tinc tion oc curred in the ter - res trial realm and took place at about 359 Ma in the lat est Famennian, is dated to the Late praeculcata Cono dont Zone, and co in cides with the max i mum ex pan sion of the ice sheets on land in Gond wana (Streel et al., 2000a).
The pro posed third step in the Late Pa leo zoic Ice Age took place in the Late Visean–Serpukhovian (Early Car bon if er ous) when gla ci ation be came bi po lar and the ice age en tered its main phase (Stan ley and Powell, 2003; Barham et al., 2012).
This third step in global cool ing co in cided with fur ther biodiversity losses and ma jor eco log i cal dis rup tion in the ma rine realm and, sim i lar to the Late Frasnian, the biodiversity loss was trig gered pri mar ily by speciation sup pres sion (Stan ley and Powell, 2003; McGhee et al., 2012).
Last, the tim ing of the bi o log i cal events that took place dur ing the stepwise on set of the Ce no zoic Ice Age and dur ing the pro - posed stepwise on set of the Late Pa leo zoic Ice Age in the Late De vo nian are very sim i lar. In the on set of the Ce no zoic Ice Age the time in ter val be tween the Oligocene and Mio cene ex tinc tion pulses was 19 mil lion years, and in the pro posed on set of the Late Pa leo zoic Ice Age the time in ter val be tween the Frasnian and Famennian ex tinc tion pulses was 16 mil lion years.
A. MARINE ECOSYSTEMS:
1. Dif fer en tial sur vival of high-lat i tude ma rine fau nas:
– Brachi o pods (Cop per, 1977, 1986, 1998)
– Mi cro bial reef biota (Cop per, 2002)
2. Dif fer en tial sur vival of deep-wa ter ma rine fau nas:
– Glass sponges (McGhee, 1996)
– Rugose cor als (Ol i ver and Ped der, 1994)
– Tornoceratid ammonoids (House, 1988)
3. Mi gra tion of deep-wa ter ma rine fau nas into shal low-wa ters:
– Glass sponges (McGhee, 1996)
– Tornoceratid ammonoids (House, 1988) 4. Blooms in cold-wa ter plank ton:
– Prasinophytes (Streel et al., 2000b)
– Radio lar ians (Racki, 1998, 1999; Cop per, 2002)
– Chitinozoans (Paris et al., 1996; Streel et al., 2000b; Grahn and Paris, 2011)
5. Dif fer en tial sur vival of fresh-wa ter ver sus ma rine spe cies:
– Ac an tho dian fishes (Dennison, 1979)
– Placoderm fishes (Dennison, 1978; Long, 1993) 6. Lat i tu di nal con trac tion of geo graphic range in sur viv ing equa to rial ma rine fau nas:
– Foraminifera (Kalvoda, 1990)
– Stromatoporoid and coral reefs (Stern, 1987; Cop per, 2002)
– Tentaculitoids (Wei et al., 2012)
– Trilobites (Morzadec, 1992) B. TERRESTRIAL ECOSYSTEMS:
1. Dif fer en tial sur vival of high-lat i tude ter res trial biota:
– Land plants (Streel et al., 2000a)
2. Lat i tu di nal con trac tion of geo graphic range in sur viv ing equa to rial ter res trial biota:
– Land plants (Streel et al., 2000a)
– Tetrapod ver te brates (McGhee, 2013)
T a b l e 1 Em pir i cal bi o log i cal ob ser va tions in ma rine and ter res trial eco sys tems that have been used to ar gue for global cool ing
in the Late Frasnian and Early Famennian
If the Earth cooled at a sim i lar rate in the Ce no zoic and in the Late De vo nian, then the sim i lar tem po ral spac ing and pat - terns of ex tinc tions dur ing these two time in ter vals may not be ac ci den tal, they may re flect the tim ing of the step-wise gla ci - ation of the planet. A sim i lar cool ing rate fur ther sug gests a sim - i lar ity in the forc ing mech a nisms, and hy poth e sized causes of the on set of the Ce no zoic Ice Age in the Oi-1 gla ci ation and the pro posed on set of the Late Pa leo zoic Ice Age in the Late Frasnian are the same: (1) CO2-downdraw due to the weath er - ing of sil i cates in geo graph i cally ex ten sive, tec toni cally-up lifted moun tain ranges and pla teaus, (2) fall ing tem per a tures, trig - gered by the de cline of the green house-gas CO2 be low thresh - old val ues in the at mo sphere, fur ther trig ger ing the sud den growth of con ti nen tal ice sheets, and (3) the sub se quent ex pan - sion and con trac tion of these early ice sheets be ing paced by Milankovitch or bital vari a tions in heat from the sun (Raymo and Ruddiman, 1992; Zachos et al., 2001; Filer, 2002; DeConto and Pol lard, 2003; Pagani et al., 2005; Averbuch et al., 2005;
Berner, 2006; Bowen, 2007; for an ex ten sive dis cus sion see McGhee, 2013).
SEDIMENTARY EVIDENCE FOR GLACIATION IN THE OLIGOCENE AND FRASNIAN?
Ev i dence for the ex is tence of gla ciers in the sec ond and third steps of the Ce no zoic Ice Age, the Mio cene and Late Plio - cene, is un dis puted (Lewis et al., 2008). Ev i dence for gla ci ation in the first step, the Early Oligocene Oi-1 gla ci ation, is much more ten u ous.
The prob lem is that Oi-1 gla cial sed i ments on land in Antarctica have been re moved by the ero sive ac tion of the larger mid dle Mio cene gla ciers, with two pos si ble ex cep tions (Strand et al., 2003; Ivany et al., 2006). The size of the first step Oi-1 gla ciers has been es ti mated to have been in the range of 7.0–11.9 ´ 106 km2 (Zachos et al., 2001; Pusz et al., 2011). In con trast, the size of the sec ond step Mio cene gla ciers has been es ti mated to have been in the range of 14–16 ´ 106 km2 (Westerhold et al., 2005; Wil son and Luyendyk, 2009), to tally cov er ing the ex panse of and eras ing the trace of the ini tial Oligocene gla ciers on the Ant arc tic land mass. Thus the best in - de pend ent sed i men tary ev i dence for the ex is tence of the Oi-1 gla ci ation is gla cially-de rived, ice-rafted de bris in ma rine sed i - ments. Zachos et al. (1992) have doc u mented the pres ence of lay ers of an gu lar quartz sands and heavy min er als at the Oi-1 strati graphic level on the Kerguelen Pla teau in the south ern In - dian Ocean. These lay ers con tain over 200 grains per gram of clastic grains that are larger than 250 mm, which are ar gued to be too large to have been trans ported off shore from Antarctica by wind and thus must have been trans ported by ice (Zachos et al., 1992). In ad di tion, Ehrmann and Mackensen (1992: fig. 8) re ported the pres ence of gravel-sized peb bles at the same strati graphic ho ri zon con tain ing the ice-rafted sand de pos its on the Kerguelen Pla teau. The pres ence of gravel in off shore ma - rine de pos its is un equiv o cal ev i dence of ice raft ing, and the pres ence of ice-rafted de bris as far north as 61°S is ar gued to be ev i dence of ei ther a high fre quency of ice bergs in the area, or of a few large de bris-con tain ing ice bergs, both of which ev i - dence large-scale con ti nen tal Oi-1 gla ci ation rather than small-scale lo cal gla ci ation in Antarctica (see dis cus sion in Ehrmann and Mackensen, 1992).
Ehrmann and Mackensen (1992) and Rob ert et al. (2002) also ar gued that changes in clay min er al ogy in ma rine sed i -
ments that take place at the Oi-1 strati graphic ho ri zon, such as a shift from smectite-dom i nated clays to illite- and chlorite-dom i - nated clays, ev i dence a shift to weath er ing in cooler cli mates in Antarctica and hence can be taken as fur ther ev i dence of gla ci - ation. Ev i dence of cli ma tic cool ing is not un equiv o cal ev i dence of gla cier for ma tion, how ever, thus the best phys i cal sedimentological ev i dence for Oi-1 gla ci ation re mains the ice-rafted de bris (Ehrmann and Mackensen, 1992; Zachos et al., 1992).
Like wise, ev i dence for the ex is tence of gla ciers in the pro - posed sec ond and third steps of the Late Pa leo zoic Ice Age, the Late Famennian and Late Visean–Serpukhovian, is un dis puted (Field ing et al., 2008; Barham et al., 2012). Sedimentological ev i dence for gla ci ation in the first pro posed step, a Late Frasnian gla ci ation, does not ex ist at pres ent.
Anal o gous to the Oi-1 gla ci ation at the on set of the Ce no - zoic Ice Age, it is here ar gued that gla cial sed i ments of Late Frasnian age at the on set of the Late Pa leo zoic Ice Age prob a - bly never will be dis cov ered on land as they will have been re - moved by the ero sive ac tion of the sub se quent much larger ice sheet that formed in Gond wana in the Late Famennian. The min i mum size of the sec ond step Late Famennian gla ciers has been mea sured to have been 16 ´ 106 km2 in west ern Gond - wana (Isaacson et al., 2008). I here pro pose that gla ciers ap - prox i mately 50 to 71% the size of the Famennian ice sheet, or 8–11 ´ 106 km2, were pres ent in west ern Gond wana in the Late Frasnian. The scal ing used to ob tain that es ti mate is based upon the scal ing of the size range of the first step Oi-1 gla ciers to the size range of the sec ond step Mio cene gla ciers in the on - set of the Ce no zoic Ice Age, and the as sump tion that that scal - ing was sim i lar in the size ranges of the first step Late Frasnian gla ciers to the sec ond step Famennian gla ciers in the pro posed on set of the Late Pa leo zoic Ice Age. As in the case of the Oi-1 gla ci ation in Antarctica, the much larger Late Famennian gla - ciers would have to tally cov ered the ex panse of and erased the trace of the ini tial Late Frasnian gla ciers on Gond wana, thus sedimentological ev i dence for Late Frasnian gla ci ation must be sought for in ma rine sed i ments off shore from the Gond wana land mass.
The large Late Famennian gla ciers pro duced metre-sized ice-rafted dropstones (Brezinski et al., 2010: fig. 10G). The pro - posed smaller Late Frasnian gla ciers, like the smaller Oi-1 gla - ciers, po ten tially pro duced sand-sized (> 250 mm) ice-rafted de - bris sim i lar to the ice-rafted de bris found in Oi-1 ma rine sed i - ments (Zachos et al., 1992; Ehrmann and Mackensen, 1992).
To test the hy poth e sis that gla ciers formed in the Late Frasnian a world-wide search should be ini ti ated for the pres ence of ice-rafted de bris in ma rine strata dated to the Late rhenana Cono dont Zone and/or to the linguiformis–Early triangularis cono dont zonal in ter val where the ex tinc tions and sharp drops in sea-sur face tem per a ture oc curred (Joachimski and Buggisch, 2002).
Are there any anom a lous oc cur rences of gravel or clastic grains larger than 250 mm in any of the off shore-ma rine black-shale strata of the Lower or Up per Kellwasser ho ri zons?
Ab sence of ev i dence is not ev i dence of ab sence, as even in the Oi-1 gla ci ation ice-rafted de bris is not uni ver sally found in the strati graphic re cord: for ex am ple, ice-rafted sand and gravel is pres ent on the Kerguelen Pla teau in the In dian Ocean but ab - sent on the Maud Rise in the At lan tic Ocean (Ehrmann and Mackensen, 1992). Yet even the dis cov ery of one site with ma - rine strata con tain ing ice-rafted de bris at the same ho ri zon as one of the Lower or Up per Kellwasser ho ri zons would con firm the ex is tence of gla ci ation in the Late Frasnian.
SUMMARY AND CONCLUSIONS
The old est gla ci ation usu ally as so ci ated with the on set of the Late Pa leo zoic Ice Age is the for ma tion of the Late Famennian (Late De vo nian) ice sheet in Gond wana (Isaacson et al., 2008; Caputo et al., 2008). How ever, it has been ar gued that an older gla cial phase ex isted in the Late Frasnian (Late De vo nian), and that it is from this gla cial phase that the on set of the Late Pa leo zoic Ice Age should be mea sured (McGhee, 2013: 203–212). If the Late Frasnian is taken as the first step in the on set of the Late Pa leo zoic Ice Age, and the Late Famennian gla ci ation is taken as the sec ond step, then the tim - ing of the pro posed on set of the Late Pa leo zoic Ice Age be - comes strik ingly sim i lar to the first two steps in the on set of the Ce no zoic Ice Age. Last, the hy poth e sized causes of the on set of the Ce no zoic Ice Age in the Oi-1 gla ci ation and the pro posed on set of the Late Pa leo zoic Ice Age in the Late Frasnian are the same: (1) CO2-downdraw due to the weath er ing of sil i cates in geo graph i cally ex ten sive, tec toni cally-up lifted moun tain ranges and pla teaus, (2) fall ing tem per a tures, trig gered by the de cline of the green house-gas CO2 be low thresh old val ues in the at mo - sphere, fur ther trig ger ing the sud den growth of con ti nen tal ice sheets, and (3) the sub se quent ex pan sion and con trac tion of these early ice sheets be ing paced by Milankovitch or bital vari - a tions in heat from the sun (Raymo and Ruddiman, 1992;
Zachos et al., 2001; Filer, 2002; DeConto and Pol lard, 2003;
Averbuch et al., 2005; Pagani et al., 2005; Berner, 2006;
Bowen, 2007).
Ac tual Oi-1 gla cial sed i ments on land in Antarctica have been re moved by the ero sive ac tion of the larger mid dle Mio - cene gla ciers and sub se quent gla ciers, and thus the best sed i - men tary ev i dence for the ex is tence of the Oi-1 gla ci ation is gla - cially-de rived, ice-rafted de bris in ma rine sed i ments (Ehrmann and Mackensen, 1992; Zachos et al., 1992). Anal o gous to the Oi-1 gla ci ation, it is ar gued that gla cial sed i ments of Late Frasnian age prob a bly never will be dis cov ered on land as they will have been re moved by the ero sive ac tion of the sub se quent much larger ice sheet that formed in Gond wana in the Late Famennian.
In con clu sion, to test the hy poth e sis that gla ciers formed in the Late Frasnian a search should be ini ti ated for the pres ence of sand-sized (>250 mm) ice-rafted de bris pres ent in Frasnian ma rine strata lo cated off shore from the main Gond wana land - mass, and dated to the Late rhenana Cono dont Zone and/or to the linguiformis–Early triangularis cono dont zonal in ter val where the ex tinc tions and sharp drops in sea-sur face tem per a - ture oc curred (Joachimski and Buggisch, 2002).
Ac knowl edge ments. I thank K.G. Miller for dis cus sions con cern ing the on set of the Ce no zoic Ice Age, and P.B. Wignall, M.M. Joachimski, and P.E. Isaacson for con struc - tive com ments on im prov ing the manu script.
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