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The Late Ordovician Epoch
461 to 444 million years ago

The Late Ordovician sees the high point of the great Ordovician biotic diversification, but also a very severe mass extinction event in its final age, the . It's thus an important time for life, both in terms of new types of organisms appearing and of old ones disappearing for ever.

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We might imagine Ordovician paleogeography as some vast Las Vegas stage, with appreciative crowds (represented by and ) applauding chorus lines of smiling, bespangled terranes as they break off from the background of the north margin, kicking and twirling their way across the southern ocean to the steps of some complex, but orderly, tectonic choreography.

Then again, we might not wish to imagine anything of the sort, since we would then be forced to visualize the first few rows of the audience being violently subducted under the inexorably advancing chorus line, the roar of the crowd being swallowed in the volcanic explosions of the , followed by the howling gale of an Ice Age and mass extinction. This would be ambitious, even for Las Vegas - more than glitz.

The grand collision at the end of the Ordovician occurred in several parts. First was the meeting of Baltica and , a chain of terranes which had rifted off the Gondwanan coast early in the Ordovician. The line of islands moved generally northwest, leaving the expanding behind it and closing the as it neared the Laurentian margin.

By this epoch, Baltica had slowed its own drive to the north, although it was still gradually crossing the equator. Nawrocki & Poprawa (2006). As Avalonia began to close the Iapetus, its northeastern end also met the western edge of Baltica, closing the . This initiated the Caledonian Orogeny in the north of England. Cocks & Torsvik (2005).

Second, the northern arm of the Iapetus, between Baltica and Laurentia, closed at the endof the Late Ordovician. This triggered an extension of the into both Baltica and (to a certain extent) Laurentia. The northern portions of Poland, Denmark, and Germany began to form at this time. Nawrocki & Poprawa (2006).

The Avalonian collision was the second of (about) three such events to affect Laurentia and Baltica over the course of the Middle Paleozoic. Stanley (1998). The collision between western Avalonia and Laurentia, and the addition of more Gondwanan blocks, would prolong the Caledonian events into the .

Some of these non-Avalonian fragments, which comprise a significant part of today's Europe, also began to rift from the coast of Gondwana (particularly ) during the Late Ordovician. The sequence and timing of these events is less clear than it once was. The original story was that Avalonia was succeeded by , a second island arc that left Gondwana and more or less followed Avalonia's path across the waves to Baltica and Laurentia. Armorica was said to include parts of Spain, the Massif Central and Brittany of France, the Saxo-Thuringian Plate of Germany, Bohemia (Perunica), Bosnia, and other indispensible components of the European Union. But, in the last few years, geologists have begun to chip pieces off Armorica (Nysæther et al., 2002; Torsvik & Cocks, 2004), or split it into two separate arcs (Nysæther et al., 2002). In fact, the tendency now is to discard the whole idea of Armorica as unit and refer to it vaguely as the "Armorican Terrane Assemblage" (Cocks & Torsvik, 2005).

Somewhere to the south of Laurentia was the coast of Gondwana. The intervening ocean may have been narrow. Some geologists (e.g. Ian Dalziel) have argued for actual contact between Laurentia and South America in the earlier Ordovician. However, by the Late Ordovician, several pieces of Mexico and Florida probably covered the northern coast of South America (Ortega-Obregón et al., 2008), which was formed by Peru during this epoch.

We will defer further discussion of West Gondwana (South America and Africa) until we can take on the Hirnantian Ice Age in general. In fact, we will slide over the rest of the world very quickly, as we've already spent too much of our own, personal time and space on the events in the Laurussian theater.

As in earlier parts of the Ordovician, West Gondwana sat over the South Pole, with the Pole itself wandering nomadically in what is now the Sahara. The rest of Gondwana extended like a sort of gigantic tail northwards from the West Gondwanan head, terminating in the North China craton, which reached north of the equator. This tail remained more or less on the opposite side of the earth from Baltica and Laurentia. North and west of the East Gondwanan tail, some of the Asian blocks were forming. In particular, Siberia was beginning to pick up and consolidate a number of unaffiliated blocks, as discussed in a recent review by Cocks & Torsvik (2007). ATW080321.

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For the most part, the Late Ordovician climate was like the climate, but more so. The world continued warm, with two episodes of global warming in the early and middle or late Hirnantian. Cocks & Torsvik (2005). The seas continued to rise. Stanley (1998). All this, as we will see, was good for life. But it ended, quite suddenly (Webby, 2004) with a brief ice age at the end of the Hirnantian.

This particular ice age seems to have been quite different from those of the or . It was perhaps similar to the ice age(s) of the , but considerably shorter. On the evidence of oxygen isotope excursions, the end-Ordovician event lasted only about 500,000 years. Stanley (1998). Even so, it seems to have included several distinct periods of glaciation. It is unclear exactly how many glacial events there were. Depending on location, there may have been only one, or as many as five.

These are minimum figures, based on the number of successive layers of glacial till. It is entirely possible that, in any given location, a late glacial event might destroy all evidence of earlier events. Yet there's bit of a pattern. Ghienne et al. (2007) find two cycles in Algeria. Le Heron et al. (2006) located four in Libya. Connally & Wiltse (1996) describe five in Saudi Arabia. This suggests that the south polar region retained a permanent ice cap, which expanded east or contracted west, depending on the severity of the various glacials and interglacials. Unfortunately, we must also entertain the possibility that the events may, instead, have been localized and asynchronous, since one of the Saudi events has been ascribed to the (earliest ).

Perhaps more interesting is the nature and movement of the ice sheets. In the map from Connally and Wiltse, the main flow is directed into a North African basin, not off the nearest edge of the continental platform. This is confirmed by the studies cited above. In addition the ice flow was highly heterogeneous. That is, it included numerous ice rivers which moved quite quickly while adjacent ice remained almost stationary. The bases of the ice sheets show considerable evidence of subglacial melting and substantial flows of meltwater undercutting the ice.

Two other data are interesting. First, Le Heron et al. (2006) emphasize the local importance of a glacial trampoline effect: subsidence caused by the weight of the ice sheet, followed by post-glacial rebound. This is peculiar for a short-lived ice sheet which ought not to have been particularly thick. Second, Ghienne et al. (2007) argue that, contrary to conventional wisdom (e.g. Benton & Harper, 1997; Stanley, 1998), the Hirnantian ice age occurred during a time of rising sea levels. They assert that the maximum intrusion of marine conditions into the interior of Africa (as judged by the presence of and other marine forms) did not occur until the earliest .

As you have probably guessed, we have not presented these particular facts merely for the general edification of the reader, but rather to introduce yet another futile and unsubstantiated speculation on a subject we are grossly unqualified to address: to wit, the cause of the Hirnantian ice age. But that is the trade-off at the heart of Palaeos. The reader obtains the questionable benefit of our research into the facts, while we are permitted to imagine that someone may imbibe the facile concoctions we brew to make sense of the data.

Our thought is that the Hirnantian ice required only a deep south polar basin and a rising sea. Without question, the oceans gradually transgressed over northern Africa over the course of the Ordovician. At some point, the seas filled the North African Basin. However, contact with the ocean was initially limited and irregular. It may well have been limited to occasional storms, since tempestites and other indicators suggest that the area to the north of the basin was (during the earliest phase of glaciation) "an open shelf that was free of sea ice ... and that was subject to episodic, very high-energy storms." Le Heron et al. (2006).

Since contact with open water was limited, the basin filled with a large volume which was thermally isolated from the oceans and located at or near the South Pole. Consequently, it froze. When it had frozen through, the weight of the ice locally depressed the basin further, accommodating more water. At this point, the coldest regions episodically become cold enough to accumulate "real" ice sheets through precipitation. Given the geography of the area, we would expect to see this ice forming near the Pole and flowing generally eastward (paleo-northeast) from Algeria into Libya and Arabia. This should occur, both because the coldest regions would lie westward and because the west end was in closest contact with the ocean. The west (Algeria) would thus be permanently frozen, while locations to the east would show an increasing number of episodic glacial "events."

At the same time, the ocean would continue to contribute water, which would melt and undermine the basin ice, perhaps causing the complex pattern of ice flow and subglacial melting actually observed. Over the slightly longer run, the rising seas outpaced the growth of the ice sheet and established regular contact with the north African basin. This ended the thermal isolation of the South Polar region. At this point, the ice cap would melt quickly. Isostatic rebound of the crust would briefly displace water, which, together with generally rising sea levels, briefly flooded additional areas post-glaciation, as observed by Ghienne et al. (2007). But the era of rising seas was now over, and the ocean began to retreat from the Sahara early in the Silurian. ATW080322.

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The Late Ordovician was the high point of the mass anti-extinction called the Great Ordovician Biodiversity Event (GOBE). With rare exceptions (, ), all animal groups present in the Late Ordovician attained their highest diversity in that epoch (Webby, 2004a) and global genus-level animal diversity came close to its Paleozoic maximum (Stanley, 1998).

There are any number of plausible reasons why this occurred. Warm epicontinental seas have always promoted the expansion of animal life, and these conditions prevailed in Laurentia, Baltica, Avalonia, Australia, China, Siberia, Mongolia and . At the same time, the cool to cold waters circulating across the long coast of West Gondwana encouraged the evolution of different forms adapted to those conditions. The continents were widely scattered, providing both a larger area of coastal and shelf environments and a large number of partially isolated ecosystems in which species would necessarily diverge through genetic drift.

In addition, the diversity of benthic environments increased. The shift to "calcite" seas meant an increase in the area of carbonate shelf, and the Late Ordovician has thus been described as the "Golden Age of hardground epizoonts." Taylor & Wilson (2003). At the same time, the intensity and depth of softground bioturbation was increasing, greatly expanding the volume of mud & crud environments for mud-sucking worms (or "softground benthic infauna," if you must) and their predators. Stanley (1998). ATW080322

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For the general run of animal life, the GOBE consisted of two diversity peaks separated by a slight dip, followed by a crash in the Hirnantian ice age. The peaks occurred in the "mid Caradoc," which we translate as Late , and the "mid Ashgill," which we suppose to be the mid to late . Webby (2004a). Webby asserts that the Sandbian peak was both lower and more gradual than the abrupt rise in the Katian Age. That is, the Sandbian peak was merely a continuation of the trend line. The slight dip between the peaks may be associated with an event of uncertain nature which resulted in the loss of various groups of Laurentian and in northern Baltica. Id.

Krug & Patzkowsky (2004) raise some serious questions about the methodology supporting this type of analysis. They attempt to correct for sampling bias and find, when this is done, that most of the temporal fine detail disappears. Animal diversity beaks broadly into the first half of the Katian and then remains relatively constant, declining slightly through the rest of the Katian before crashing in the Hirnantian. Yet Cocks & Torsvik (2005) make the qualitative, but weighty, point that the later Katian was marked by an episode of warming "which led to the formation of substantial carbonate mud mounds (bioherms) with very diverse brachiopods, , , and ." Ultimately, we do not yet know enough about the fossil record, and the statistical techniques needed to analyze it. The GOBE certainly happened, but its precise contours are still hazy. ATW080322.

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We have covered the timing and mechanics of the Hirnantian ice age above. The effects of this cold snap on life were variable. Plainly, many species became extinct - 70% of all animal species, according to one estimate. Benton & Harper (1997). Many common families of Ordovician brachiopods, echinoderms, ostracodes, and trilobites disappeared entirely. Many others shifted their ranges. Id. The end of the cooling caused yet further evolutionary angst (Stanley, 1998; Krug & Patzkowsky, 2004), as groups which had just worked out how to adapt to colder conditions became frustrated with an environment that capriciously changed its mind and wanted everything back the way it had been before it installed a state-of-the-art South Polar ice cap. We admit that we have seriously considered extinction ourselves, under similar circumstances.

A 70% species-level extinction wouldn't involve much family level extinction at all, if the extinct species were randomly distributed. Further, to use an industrial analogy, this extinction did not represent the kind of wholesale elimination of jobs groups which occurs when the local factory closes, or when technology replaces assembly lines with assembly language. The end-Ordovician is more closely analogous with the case in which two local firms merge and become a regional producer. Some jobs become redundant, some become more specialized, and some new ones are created. Many individual lives and careers are drastically altered, for better or worse, but the overall economic dislocation is minor and temporary.

In one sense, this analogy is exact. The physical merger of Avalonia, Baltica, and Laurentia wouldn't leave room for three different species of brachiopod that did exactly the same thing. These species would specialize or become extinct. To that extent, an end-Ordovician extinction would have occurred with or without an ice-age. Similarly, any rapid series of environmental changes will favor generalists over specialists. By definition, generalists will overlap in ecospace more than specialists, and diversity will necessarily suffer. As human beings, we are in a particularly poor position to complain about that result. Finally, the ice-age almost completed the Ordovician trend toward replacing the old Cambrian fauna, e.g. trilobites and "inarticulate" brachiopods, with the characteristic animals of the Paleozoic, e.g. , , , and brachiopods which, if no more eloquent than their ancestors, were none the less articulate enough for the needs of the Silurian. ATW080323.

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  • Benton MJ & DAT Harper (1997), Basic Paleontology. Addison Wesley Longman, 342 pp.
  • Cocks LRM & TH Torsvik (2005), Baltica from the late Precambrian to mid-Palaeozoic times: The gain and loss of a terrane’s identity. Earth-Sci. Rev. 72: 39-66.
  • Cocks LRM & TH Torsvik (2007), Siberia, the wandering northern terrane, and its changing geography through the Palaeozoic. Earth-Sci. Rev. 82: 29-74.
  • Connally T & E Wiltse (1996), Saudi Sandstone Correlations. Middle East Well Eval. Rev. 16: 26-41.
  • Ghienne J-F, K Boumendjel, F Paris, B Videt, P Racheboeuf & HA Salem (2007), The Cambrian-Ordovician succession in the Ougarta Range (western Algeria, North Africa) and interference of the Late Ordovician glaciation on the development of the Lower Palaeozoic transgression on northern Gondwana. Bull. Geosci. 82: 183–214.
  • Krug AZ & ME Patzkowsky (2004), Rapid recovery from the Late Ordovician mass extinction. Proc. Nat. Acad. Sci. (USA) 101: 17605-17610.
  • Le Heron DP, J Craig, OE Sutcliffe & R Whittington (2006), Late Ordovician glaciogenic reservoir heterogeneity: an example from the Murzuq Basin, Libya. Marine Petrol. Geol. 23: 655-677.
  • Nawrocki J & P Poprawa (2006), Development of Trans-European Suture Zone in Poland: from Ediacaran rifting to Early Palaeozoic accretion. Geol Quart. 50: 59-76.
  • Nysæther E, TH Torsvik, R Feist, HJ Walderhaug & EA Eide (2002), Ordovician palaeogeography with new palaeomagnetic data from the Montagne Noire (Southern France). Earth Planet. Sci. Let. 203: 329-341.
  • Ortega-Obregón C, LA Solari, JD Keppie, F Ortega-Gutiérrez, J Solé & S Morán-Ical (2008), Middle-Late Ordovician magmatism and Late Cretaceous collision in the southern Maya block, Rabinal-Salamá area, central Guatemala: Implications for North America–Caribbean plate tectonics. GSA Bull. (in press).
  • Stanley, SM (1998), Earth System History. WH Freeman & Co., 615 pp.
  • Torsvik, TH & LRM Cocks (2004), Earth geography from 400 to 250 Ma: a palaeomagnetic, faunal and facies review. J. Geol. Soc. 161: 555-572.
  • Webby BD (2004a), Introduction in BD Webby, F Paris, ML Droser & IG Percival (eds.), The Great Ordovician Biodiversification Event. Columbia Univ. Press, pp. 1-37.
  • Webby BD (2004), Stromatoporoids in BD Webby, F Paris, ML Droser & IG Percival (eds.), The Great Ordovician Biodiversification Event. Columbia Univ. Press, pp. 112-118.


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