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Surviving a mass extinction: being small, proliferating rapidly afterward

lpetrich

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 Lilliput effect - after mass extinctions, the surviving species are often much smaller that many of their close relatives before the extinction.

Ancient mass extinction may have shrunk Earth's creatures | Science | AAAS
About 360 million years ago, Earth's seas were filled with myriad fishes, including creatures the size of school buses. Then a mass extinction hit the Age of Fishes. It killed off most of the big guys, according to a new study, and effectively shrunk most vertebrate species to the size of a human forearm or smaller. The findings imply that our planet's next mass extinction—which some believe is already underway—could similarly shrivel any species that remain.

The ancient extinction happened about 359 million years ago, at the end of the Devonian Period. A 100,000-year-long cold spell triggered the growth of glaciers almost down to tropical latitudes, says Lauren Sallan, a paleobiologist at the University of Pennsylvania. Sea level fell substantially, wiping out much of the shallow-water habitat surrounding major landmasses. Because few creatures had yet moved onto land, many ecosystems were devastated. About 96% of the world's vertebrate species disappeared, making it one of Earth's five largest die-offs.
This is the Hangenberg event at the end of the Devonian Period.
Body-size reduction in vertebrates following the end-Devonian mass extinction

After a Mass Extinction, Only the Small Survive - The New York Times
After the mass extinction 359 million years ago, the scientists found, vertebrates were smaller on average than before, and they stayed that way for the next 36 million years.

Peter J. Wagner, a curator of paleobiology at the Smithsonian Institution who was not involved in the study, said one of its strengths was the broad stretch of time it covered.

Previous studies were “more like snapshots of what things were like before and afterwards,” he said. “This study goes much further.” Dr. Wagner, who wrote a commentary in Science accompanying the new study, said the work was also important because it dissected the Lilliput Effect, examining trends in different groups of species.

Some groups simply stayed at the same diminished size during the Mississippian, while others steadily shrank. Sharks, for example, dwindled from over a yard long to about just a few inches. Our own tetrapod relatives shrank from the size of dogs to the size of cats or smaller.
 
The Lilliput effect was also a result of the biggest mass extinction ever, at least in the Phanerozoic: the Permian-Triassic mass extinction.

Changes in size and growth rate of ‘Lilliput’ animals in the earliest Triassic - ScienceDirect
Marine invertebrate taxa that survived the late Permian (latest Changhsingian) mass extinction event are all much smaller than they were prior to the event, an example of the Lilliput effect. New taxa that first appeared in the immediate extinction aftermath are also small compared to their size in younger strata. The Lilliput effect is a temporary phenomenon, with most surviving taxa returning to pre-extinction size in the first two conodont zones of the Triassic.
The Lilliput effect in the aftermath of the end-Permian extinction event - ScienceDirect
Early Triassic animal body fossils and trace fossils are small relative to those in older and younger intervals. Size decreases sharply through the end-Permian extinction event and Permian/Triassic boundary, and the smallest sizes are encountered in the parvus and isarcica Zones of the earliest Induan. Animals appearing within these two zones are also exceedingly small, compared to younger congenerics and conspecifics. Temporary, dramatic size decrease of surviving taxa in the immediate aftermath ...
 
Another one on the end-Devonian mass extinction: The Lilliput Effect: Trends in Body Size Following Ancient Mass Extinctions | EvolutionShorts

Another one on the Permo-Triassic mass extinction: Body Size Reductions in Nonmammalian Eutheriodont Therapsids (Synapsida) during the End-Permian Mass Extinction - these were mammal-like reptiles or protomammals

It also happened in the best-known mass extinction:
Dinosaurs are well-known for being big, and some of the bigger ones made it all the way to the end of the Cretaceous. Here are some of the biggest ones at the end of the Cretaceous:

 Alamosaurus - "Specimens of a juvenile Alamosaurus sanjuanensis have been recovered from only a few meters below the Cretaceous-Paleogene boundary in Texas, making it among the last surviving non-avian dinosaur species." - this sauropod grew as large as other ones before it. Length ~ 30 m, shoulder height ~ 6 m, hip height ~ 5 m, body weight ~ 30 mt.

 Tyrannosaurus - one of the largest theropods. Length ~ 12 m, hip height ~ 4 m, body mass ~ 7 mt.

 Edmontosaurus annectens - one of the largest duckbills. Length ~ 12 m, hip height ~ 4 m, body mass ~ 7 mt.

 Triceratops - largest ceratopsian. Length ~ 8 m, shoulder/hip height ~ 3 m, body mass ~ 9 mt.

 Ankylosaurus - largest ankylosaurid. Length ~ 8 m, shoulder/hip height ~ 2 m, body mass ~ 5 mt.

 Dinosaur size has more. For comparison:

Present-day African bush elephant (Loxodonta africana) - largest present-day land animal. Length ~ 4 m, shoulder height ~ 3 m, body mass ~ 4 mt.

But the only broad-sense dinosaurs to survive are birds, and judging by their descendants, all of the species that survived were capable of flight. That gives the survivors a maximum body mass of about 20 kg, more than 1000 times less than that of the largest dinosaurs at the end of the Cretaceous.
 
Trace fossils in the aftermath of mass extinction events | Geological Society, London, Special Publications
Here we review published ichnological studies for the Ordovician-Silurian, Permian-Triassic and Cretaceous-Tertiary extinction-recovery intervals. In addition, new information regarding the Triassic-Jurassic ichnological record from England, Austria and the western USA is presented. Trace fossils provide important information on the ecological response of the benthic community at such times. In the immediate post-extinction aftermath, the ichnodiversity, burrow size, depth of bioturbation, and ichnofabric index of the sediments are all much reduced. There is an increase in all these parameters through the post-extinction recovery period.
More Lilliput effect.

I found some interesting papers on birds in this connection. They are descended from theropod dinosaurs, which had a range of sizes from pigeon-sized to elephant-sized like tyrannosaurids. The first "bird" is late-Jurassic pigeon-sized  Archaeopteryx, because of its feathers. But it had more typical non-avian theropod features like teeth, lack of a big breastbone, front-limb claws, and a long tail. Several other small to medium-sized non-avian theropods have recently been found to have feathers, so that's why I put "bird" in quotes.

But by the early Cretaceous, bird ancestors and close relatives of them had emerged that had short tails and big breastbones, much like present-day birds.

The most common of them were "enantiornithines" ("mirror birds"), much like present-day birds, but with teeth and wing-finger claws. Many of them had feet adapted for perching, much like many present-day birds. Grabbing a tree branch with their feet.

In the Late Cretaceous was  Ichthyornis ("fish bird"), much like present-day water birds, but with teeth, and  Hesperornis ("western bird"), with a beak and back teeth. In the last few million years (Maastrichtian) was  Vegavis a bird and no teeth that was much like present-day ducks and geese (Anatidae).
 
Mass extinction of birds at the Cretaceous–Paleogene (K–Pg) boundary | PNAS - "Here, we describe a diverse avifauna from the latest Maastrichtian of western North America, which provides definitive evidence for the persistence of a range of archaic birds to within 300,000 y of the K–Pg boundary" - these toothy birds.

Early Evolution of Modern Birds Structured by Global Forest Collapse at the End-Cretaceous Mass Extinction - ScienceDirect

"Global paleobotanical and palynological data show that the K-Pg Chicxulub impact triggered widespread destruction of forests [8, 9]."

"The fossil record and recent molecular phylogenies support an extraordinary early-Cenozoic radiation of crown birds (Neornithes) after the Cretaceous-Paleogene (K-Pg) mass extinction [1, 2, 3]."
Crown taxa - taxa that include all their present-day members and no extinct members that branched off before the present-day ones' ancestors branched off.

"Here, ancestral state reconstructions of neornithine ecology reveal a strong bias toward taxa exhibiting predominantly non-arboreal lifestyles across the K-Pg, with multiple convergent transitions toward predominantly arboreal ecologies later in the Paleocene and Eocene. By contrast, ecomorphological inferences indicate predominantly arboreal lifestyles among enantiornithines, the most diverse and widespread Mesozoic avialans [5, 6, 7]."

Meaning that present-day birds' early ancestors were ground birds while the mirror birds often lived in trees, much like many present-day birds. After the mirror birds were killed off by the K-Pg disaster, the surviving birds had no competition in the trees, and moved into them.
Palynological data from K-Pg boundary sections worldwide reveal a vegetation response with a fern spike and floral turnover [14], which together indicate forest destruction on a global scale and a protracted (∼1,000 year) onset of the recovery of climax vegetation [15]

...
Global paleobotanical and palynological data show that the K-Pg Chicxulub impact triggered widespread destruction of forests [8, 9]. We suggest that ecological filtering due to the temporary loss of significant plant cover across the K-Pg boundary selected against any flying dinosaurs (Avialae [10]) committed to arboreal ecologies, resulting in a predominantly non-arboreal post-extinction neornithine avifauna composed of total-clade Palaeognathae, Galloanserae, and terrestrial total-clade Neoaves that rapidly diversified into the broad range of avian ecologies familiar today.
Without forests, the mirror birds had nowhere to go.
  • Palaeognathae = ostrich, etc. -- ground birds
  • Galloanserae = Galliformes (chicken, turkey, pheasant, ...), Anseriformes (duck, goose, swan, ...) -- ground, water birds
  • Neoaves = all the rest -- ground, water, tree birds
All three clades diverged from each other late in the Cretaceous.
 
How big was the K-Pg disaster?
The plant fossil record and models of the effects of the Chicxulub impact provide strong evidence for the devastation of forest communities at the K-Pg boundary. Initial disruption came from energy dissipated by the impact blast, leveling trees within a radius of ∼1,500 km, and as intense radiated heat, which may have ignited wildfires on a global scale [29, 30, 31]. This was most likely followed by acid rain resulting from the emission of sulfate-rich vapor [32] and ejection of a large quantity of soot into the atmosphere [33], potentially blocking photosynthetic activity for several years and most likely inducing limited global climate cooling [34, 35, 36, 37]. This phase of suppressed sunlight, notoriously difficult to reconstruct, is supported by the proliferation of saprotrophs thriving on decomposing organic matter [38].

The post-impact recovery of terrestrial plant communities occurred in two phases. The first is marked by the dominance of fern spores in an ∼1-cm-thick interval [39] (Figure 3). Ferns are pioneer re-colonizers of devastated landscapes, and their proliferation represents a classic example of a “disaster flora” composed of taxa capable of rapidly germinating from spores and rhizomes and/or roots. ... The K-Pg fern spike has been identified worldwide and is an indicator of global canopy loss ([14] and references therein). Sedimentation rates based on recent high-resolution radiometric dating of K-Pg bentonites from Montana [40] and the Denver Basin [15] show that establishment of the fern spike occurred within a century of the Chicxulub impact and that the fern spike disaster flora persisted on the order of 1,000 years. This general timescale is corroborated by estimated sedimentation rates from New Zealand (Figure S1). Terrestrial floras were most likely devoid of extensive closed-canopy forests during this phase.

The second phase is marked by the re-establishment of canopy vegetation: the earliest Paleocene marks a change in forest community structure compared to the Cretaceous. Typical earliest Paleocene plant assemblages are characterized by low taxonomic diversity [41, 42, 43] and by a shift of dominance toward new angiosperms and conifers (the disappearance of diverse Cretaceous taxa [K-taxa] and proliferation of Ulmipollenites krempii, Kurtzipites spp., palms [Arecipites spp.], Taxodiaceae, and Pinaceae; Figure 3), long-lived plants that are indicative of modern climax communities [14, 44]. This low-diversity flora persists until the appearance of diversity hotspots ∼1.4 Ma after the K-Pg [45].
 
The ancestors of  Palaeognathae,  Galloanserae, and  Neoaves are all reconstructed as being ground birds.

The phylogeny of Neoaves has been *very* difficult to untangle, because of its rapid divergence after the K-Pg disaster. It has required sequencing the genomes of several species, so one can do whole-genome comparisons. There are seven clades that emerge out of this analysis:
  • Telluraves (landbirds) -- tree
  • Aequornithes (waterbirds) -- water
  • Eurypygimorphae (Eurypygiformes: sunbittern, kagu (ground, tree), Phaethontiformes: tropicbirds (water)) -- 50 Mya, ?
  • Otidimorphae (Musophagiformes: turacos (tree), Otidiformes: bustards (ground), Cuculiformes: cuckoos (tree)) -- 34 Mya, ?
  • Strisores (Caprimulgiformes: nightjars (ground), Nyctibiiformes: potoos (tree), Steatornithiformes: oilbirds (ground), Podargiformes: frogmouths (tree), Apodiformes: swifts, hummingbirds (ground, tree), Aegotheliformes: owlet nightjars (tree)) -- 51 Mya, ?
  • Columbimorphae (Mesitornithiformes mesites, Pterocliformes: sandgrouse, Columbiformes: doves, pigeons) -- 22 Mya, ground
  • Mirandornithes (Phoenicopteriformes flamingos, Podicipediformes grebes) -- 50 Mya, water
with three that are difficult to place:
  • Opisthocomiformes (hoatzin) -- 36 Mya, tree
  • Gruiformes (cranes and rails) -- 60 Mya, water, ground
  • Charadriiformes (shorebirds, gulls and alcids) -- 55 Mya, water

Aequornithes -- 62 Mya, water
  • Gaviiformes: loons
  • Procellariimorphae: (Procellariiformes: albatrosses, petrels, Sphenisciformes: penguins)
  • Pelecanimorphae: (Pelecaniformes: pelicans, Suliformes: boobies, cormorants, Ciconiiformes: storks)
Telluraves -- 62.5 Mya, tree
  • Accipitrimorphae: (Cathartiformes: New World vultures, Accipitriformes: hawks, eagles, Old World vultures) (tree, high ground)
  • Strigiformes: owls (tree)
  • Coraciimorphae: (Coliiformes: mousebirds (tree), Leptosomiformes: cuckoo roller (tree), Trogoniformes: trogons (tree), Bucerotiformes: hornbills, hoopoes (tree), Coraciiformes: kingfisher, roller (tree, ground), Piciformes: woodpeckers, toucans (tree))
  • Australaves: (Cariamiformes: seriemas (ground), Falconiformes: falcons (tree), Psittaciformes: parrots (tree), Passeriformes: passerine birds (tree))
Accipitrimorphae and Strigiformes, and sometimes also Coraciimorphae are sometimes grouped as Afroaves.
 
Big-time insights from a tiny bird fossil | PNAS
noting
Early Paleocene landbird supports rapid phylogenetic and morphological diversification of crown birds after the K–Pg mass extinction | PNAS

An early mousebird, Tsidiiyazhi abini, found in New Mexico. It is close to the size of the present-day  Speckled mousebird The present-day bird is about 35 cm long, with its tail about half that length, and it weighs about 57 grams.

It is from the late Danian in the Paleocene in the Paleogene, about 62.5 million years ago. That's only 3.5 million years after the K-Pg disaster.

Its main competition is an early penguin, Waimanu, from about 60.5 million years ago.


What motivated me to start this thread:
Genomic Signature of an Avian Lilliput Effect across the K-Pg Extinction | Systematic Biology | Oxford Academic

The earliest divergences of birds have a big problem when one tries to date them using molecular-clock techniques. Their age is much older than what one finds in the fossil record, from <75 Mya to >160 Mya -- the proto-bird Archaeopteryx lithographica is about 150 Mya.
Reconciling molecular divergence time estimates with the known crown bird fossil record thus suggests a hidden acceleration of the avian molecular clock at some point in avian evolutionary history (e.g., Alroy 1999; Benton 1999; Bromham 2003). However, a plausible mechanism for such an acceleration has yet to be articulated, casting doubt on this interpretation (Easteal 1999).

Mass extinction events have been characterized by marked reductions in body size among surviving lineages relative to their pre-extinction antecedents (Twitchett 2007). This phenomenon, known as the “Lilliput Effect” (Urbanek 1993) is difficult to observe directly in many clades (including birds) because it requires an exceptionally well-sampled fossil record immediately before and after an extinction event. The challenge is exaggerated when surviving lineages are predicted to be very small-bodied, and therefore subject to taphonomic bias against their preservation and discovery (Brown et al. 2013). Body size is correlated to a constellation of traits related to life history and demography, including generation length, population size, longevity, and metabolic rate (Simpson 1944; Western and Ssemakula 1982; Brown 1995; Roff 2002). As a result, pronounced changes in body size may be correlated with changes in rates of nucleotide substitution, an expectation that stems from the nearly neutral theory of molecular evolution (Kimura 1968; Ohta 1973; Nabholz et al. 2013; Figuet et al. 2016), as well as the metabolic theory of ecology (e.g., Brown et al. 2004; see Supplementary material available on Dryad for a detailed discussion). Invoking the Lilliput Effect as a hypothesis to explain a hidden period of increased substitution rates in a clade implies two general predictions: (i) that small body sizes are associated with faster substitution rates and (ii) that survivors of a mass extinction are characterized by reduced size relative to their pre-extinction relatives.
The authors then did some simulations of the effects of small-size speedups on what one calculates for divergence time. They conclude that that would likely resolve the discrepancies.
 
Bird neurocranial and body mass evolution across the end-Cretaceous mass extinction: The avian brain shape left other dinosaurs behind

The K-Pg mass extinction included a lot of taxa that we don't ordinarily think suffered very much.

Severe extinction and rapid recovery of mammals across the Cretaceous–Palaeogene boundary, and the effects of rarity on patterns of extinction and recovery - Longrich - 2016 - Journal of Evolutionary Biology - Wiley Online Library
To better understand terrestrial extinction and recovery and how sampling influences these patterns, we collected data on the occurrence and abundance of fossil mammals to examine mammalian diversity across the K-Pg boundary in North America. Our data show that the extinction was more severe and the recovery more rapid than previously thought. Extinction rates are markedly higher than previously estimated: of 59 species, four survived (93% species extinction, 86% of genera). Survival is correlated with geographic range size and abundance, with widespread, common species tending to survive. This creates a sampling artefact in which rare species are both more vulnerable to extinction and less likely to be recovered, such that the fossil record is inherently biased towards the survivors. The recovery was remarkably rapid. Within 300 000 years, local diversity recovered and regional diversity rose to twice Cretaceous levels, driven by increased endemicity; morphological disparity increased above levels observed in the Cretaceous.
Here also, the survivors could diversify because they had much less competition.

Mass extinction of lizards and snakes at the Cretaceous–Paleogene boundary | PNAS
A revision of fossil squamates from the Maastrichtian and Paleocene of North America shows that lizards and snakes suffered a devastating mass extinction coinciding with the Chicxulub asteroid impact. Species-level extinction was 83%, and the K-Pg event resulted in the elimination of many lizard groups and a dramatic decrease in morphological disparity. Survival was associated with small body size and perhaps large geographic range. The recovery was prolonged; diversity did not approach Cretaceous levels until 10 My after the extinction, and resulted in a dramatic change in faunal composition.
The Lilliput effect again.
 
More about mammals and K-Pg.
The existence of relatively large mammals in the basal Palaeocene is particularly striking in the light of the extinction of small dinosaurs. Basal Pu1 communities include a number of species that ranged from 500 g to several kilograms, approaching or exceeding the size of small Late Cretaceous dinosaurs such as alvarezsaurids (Longrich & Currie, 2009a) and microraptorines (Longrich & Currie, 2009b). Mammals may have been able to increase in size very rapidly, even within a few thousand years, but the fact that Cretaceous Protungulatum is as large (in fact, larger than) its basal Palaeocene counterpart implies that this lineage crossed the K-Pg boundary at relatively large body size, and other large-bodied species are likely to have done so as well. Other, smaller species found in the Bug Creek and Mantua Lentil tend to be comparable in size to the same species or closely related species found in the Maastrichtian, which implies that we are not seeing rapid increases in body size between the K-Pg boundary and the appearance of these faunas, although larger species do begin to appear very soon after.

The implication is that whereas large body size alone may explain the disappearance of most dinosaurs, other factors must explain the disappearance of small nonavian dinosaurs, some of which appear to have been below the 5 kg size threshold necessary for survival. Some other aspect of mammalian biology, for example adaptations for nocturnal foraging, must have predisposed mammals to survive where these dinosaurs did not.
Weird. The small dinosaurs were likely warm-blooded, as were the mammals, so warm-bloodedness may not have made much difference.

Our data also suggest that the recovery was more rapid than previously thought. C29r faunas span a range of time, some deposited within ~50 000 years of the boundary, others perhaps 200–300 000 years after.
Much of that repopulation was from immigrant species from elsewhere, with some of them coming from Asia. That continent had a land connection to North America back then, and was much farther from Chicxulub, and thus less affected.
The initial wave of colonization probably took place very rapidly. Both Bug Creek (Lofgren, 1995) and Mantua Lentil (Jepsen, 1940) occur in thick, coarse channel sands that cut down into the underlying Maastrichtian. These channels are likely to be part of the pulse of continental erosion (Misra & Froelich, 2012) caused by widespread vegetation die-off and deforestation at the K-Pg boundary (Nichols & Johnson, 2008; Schulte et al., 2010). If so, initial colonizations would appear to precede complete revegetation of the landscape, suggesting that they occurred within a few thousand years of the extinction.
Once they arrived, they soon split up into lots of different species.
Within 300 000 years, many faunas had experienced recovery of local diversity, and regional diversity had dramatically increased, driven by increased endemicity. Invasive lineages are highly diverse: multiple species of Protungulatum, Baioconodon, Mimatuta, Oxyacodon, Catopsalis and Procerberus are present. These species complexes may represent the radiation of a single colonist species, analogous to insular radiations such as Darwin's finches (Grant & Grant, 2011). If so, multiple speciation events occurred over a period of 200–300 000 years, implying rates of speciation comparable to or exceeding those seen in extant dispersal-driven radiations, such as Darwin's finches (Grant & Grant, 2011), Hawaiian honeycreepers (Lerner et al., 2011) and African cichlids (Elmer et al., 2009).
Honeycreeper: a kind of passerine bird. Like Darwin's finches, more passerine birds, their ancestors most likely flew there. Cichlids in African lakes have been abundantly researched.

Something remarkable:
Yet given the high rates of extinction seen here, the success of mammals in the Palaeocene seems to stem less from high rates of survival than from their ability to adapt to the aftermath. Squamates, crocodyliforms and turtles, which fared as well or better than mammals during the extinction, did not manage to radiate to occupy and dominate terrestrial herbivore and carnivore niches to the same extent as mammals, and instead show relatively limited niche expansion in the Palaeocene (Hutchison, 1982; Longrich et al., 2012). Adaptability, not survivability, was the mammals’ key to success. But why mammals were so successful is still not clear.
 
Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary
Pterosaurs were the first vertebrates to evolve powered flight and the largest animals to ever take wing. The pterosaurs persisted for over 150 million years before disappearing at the end of the Cretaceous, but the patterns of and processes driving their extinction remain unclear. Only a single family, Azhdarchidae, is definitively known from the late Maastrichtian, suggesting a gradual decline in diversity in the Late Cretaceous, with the Cretaceous–Paleogene (K-Pg) extinction eliminating a few late-surviving species. However, this apparent pattern may simply reflect poor sampling of fossils. Here, we describe a diverse pterosaur assemblage from the late Maastrichtian of Morocco that includes not only Azhdarchidae but the youngest known Pteranodontidae and Nyctosauridae. With 3 families and at least 7 species present, the assemblage represents the most diverse known Late Cretaceous pterosaur assemblage and dramatically increases the diversity of Maastrichtian pterosaurs. At least 3 families—Pteranodontidae, Nyctosauridae, and Azhdarchidae—persisted into the late Maastrichtian. Late Maastrichtian pterosaurs show increased niche occupation relative to earlier, Santonian-Campanian faunas and successfully outcompeted birds at large sizes. These patterns suggest an abrupt mass extinction of pterosaurs at the K-Pg boundary.
So the pterosaurs also were done in by the K-Pg disaster.

In the Late Cretaceous, birds and pterosaurs avoided competing by doing niche partitioning: birds were small and pterosaurs were large. No known birds were larger than 1 m wingspan or 5 kg body weight, and most pterosaurs had wingspans of 1 to 10 m.

Likewise, on land, (non-avian) dinosaurs were mostly large and mammals were mostly small.

The Lilliput effect, in both cases. It was small ones that survived the K-Pg disaster.

After that disaster, birds and mammals soon started becoming large. The largest wingspans I can find for birds is 3.5 - 4 meters (present-day albatrosses and recently-extinct teratorns), well in the pterosaur range, and for mammals, the largest land ones, like elephants and the extinct rhinoceros Indricotherium are and were impressively large, but not quite as large as several dinosaurs. Predators like Tyrannosaurus were much larger than even the largest mammalian land predators, like the polar bear.
 
I made the qualification "land", because some mammals became partially or completely aquatic, like sirenians and cetaceans. The largest predators are sperm whales and the largest filter feeders blue whales.

Of dups and dinos: evolution at the K/Pg boundary - ScienceDirect
  • Although many plants are extant polyploids, the number of ancient polyploidy events that survived on the long term is limited.
  • Many whole genome duplications seem to be clustered in time around the K/Pg boundary.
  • Both adaptive and neutral processes might have contributed to promote the establishment of polyploids.

Global fires after asteroid impact probably caused mass extinction - Balcerak - 2013 - Eos, Transactions American Geophysical Union - Wiley Online Library
then
Revisiting wildfires at the K‐Pg boundary - Morgan - 2013 - Journal of Geophysical Research: Biogeosciences - Wiley Online Library
then
K‐Pg extinction: Reevaluation of the heat‐fire hypothesis - Robertson - 2013 - Journal of Geophysical Research: Biogeosciences - Wiley Online Library
The global debris layer created by the end-Cretaceous impact at Chicxulub contained enough soot to indicate that the entire terrestrial biosphere had burned. Preliminary modeling showed that the reentry of ejecta would have caused a global infrared (IR) pulse sufficient to ignite global fires within a few hours of the Chicxulub impact. This heat pulse and subsequent fires explain the terrestrial survival patterns in the earliest Paleocene, because all the surviving species were plausibly able to take shelter from heat and fire underground or in water.
They then consider the difficulty of a seeming lack of charcoal from all the burned trees.
We show that the apparent charcoal depletion in the Cretaceous-Paleogene layer has been misinterpreted due to the failure to correct properly for sediment deposition rates. We also show that the mass of soot potentially released from the impact site is far too low to supply the observed soot. However, global firestorms are consistent with both data and physical modeling.
Seems very horrible -- that asteroid strike set much of our planet's vegetation on fire.

Fossil evidence reveals how plants responded to cooling during the Cretaceous-Paleogene transition | BMC Plant Biology | Full Text
It was somewhat cooler after the K-Pg disaster's dust settled.

Unexpected resilience of species with temperature-dependent sex determination at the Cretaceous–Palaeogene boundary | Biology Letters

Impact of K-Pg Mass Extinction Event on Crocodylomorpha Inferred from Phylogeny of Extinct and Extant Taxa | bioRxiv

The authors could recognize the K-Pg even in their family trees of extinct crocodilian relatives. They also note that three crocodilians survived the K-Pg disaster: the ancestors of the alligators, crocodiles, and gharials. Their ancestors had diverged some 90 - 118 Mya in the Campanian in the early Cretaceous.
 
Mammals across the K/Pg boundary in northeastern Montana, U.S.A.: dental morphology and body-size patterns reveal extinction selectivity and immigrant-fueled ecospace filling | Paleobiology | Cambridge Core
My results reveal several key findings: (1) latest Cretaceous mammals, particularly metatherians and multituberculates, had a greater ecomorphological diversity than is generally appreciated, occupying regions of the morphospace that are interpreted as strict carnivory, plant-dominated omnivory, and herbivory; (2) the decline in dental-shape disparity and body-size disparity across the K/Pg boundary shows a pattern of constructive extinction selectivity against larger-bodied dietary specialists, particularly strict carnivores and taxa with plant-based diets, that suggests the kill mechanism was related to depressed primary productivity rather than a globally instantaneous event; (3) the ecomorphological recovery in the earliest Paleocene was fueled by immigrants, namely three multituberculate families (taeniolabidids, microcosmodontids, eucosmodontids) and to a lesser extent archaic ungulates; and (4) despite immediate increases in the taxonomic richness of eutherians, their much-celebrated post-K/Pg ecomorphological expansion had a slower start than is generally perceived and most likely only began 400,000 to 1 million years after the extinction event.
The Lilliput effect here also.


Frontiers | Phoenix from the Ashes: Fire, Torpor, and the Evolution of Mammalian Endothermy | Physiology
 
There were some other disasters at the time of the Chicxulub impact.  Cretaceous–Paleogene extinction event

Stable climate in India during Deccan volcanism suggests limited influence on K–Pg extinction - ScienceDirect
Large igneous provinces (LIPs) have been temporally correlated to mass extinctions throughout the Phanerozoic, including the emplacement of the Deccan Volcanic Province (DVP; 66.3–65.6 Ma) in western and central India, which has been invoked as either a cause or exacerbating factor in the Cretaceous-Paleogene (K-Pg) extinction. However, relatively little is known about local paleoclimatic and paleoenvironmental responses to volcanism. To investigate the DVP's role as a driver of local environmental change and to provide climatic background for known ecological shifts, new inter-basaltic paleosol profiles at the eastern edge of the DVP are used in conjunction with profiles from the literature to reconstruct paleoclimate and terrestrial environments before and after the K–Pg. These profiles provide a novel opportunity to study the sediments within basalt flows before, during, and after a mass extinction event and in the midst of a LIP emplacement event. Paleoclimate proxies and the Floral Humidity Province proxy reflect little long-term change in either climate or environment across the K–Pg, with stable precipitation values and temperatures accompanied by a constant forest signal. These interpretations are corroborated by macrofloral records and sedimentology from India, which suggest some environmental turnover but generally support a forested, fluvio-lacustrine environment throughout the duration of volcanism. Our results support the possibility of rapid recovery times for terrestrial ecosystems during volcanism and suggest that while DVP eruptions may have exacerbated long-term environmental perturbation, the emplacement of the DVP is not likely to have caused the terrestrial mass extinction at the K–Pg boundary.
So the Deccan Traps were formed a bit after the Chicxulub impact. They were likely formed by the  Réunion hotspot - Réunion Island is about 200 km SW of Mauritius and 900 km east of Madagascar.

The Boltysh impact structure: An early Danian impact event during recovery from the K-Pg mass extinction
oth the Chicxulub and Boltysh impact events are associated with the K-Pg boundary. While Chicxulub is firmly linked to the end-Cretaceous mass extinction, the temporal relationship of the ~24-km-diameter Boltysh impact to these events is uncertain, although it is thought to have occurred 2 to 5 ka before the mass extinction. Here, we conduct the first direct geochronological comparison of Boltysh to the K-Pg boundary. Our 40Ar/39Ar age of 65.39 ± 0.14/0.16 Ma shows that the impact occurred ~0.65 Ma after the mass extinction. At that time, the climate was recovering from the effects of the Chicxulub impact and Deccan trap flood volcanism. This age shows that Boltysh has a close temporal association with the Lower C29n hyperthermal recorded by global sediment archives and in the Boltysh crater lake sediments. The temporal coincidence raises the possibility that even a small impact event could disrupt recovery of the Earth system from catastrophic events.
 
Looking back further, to the  Permian–Triassic extinction event -

Abruptness of the end-Permian mass extinction as determined from biostratigraphic and cyclostratigraphic analyses of European western Tethyan sections
The Permian-Triassic (P-Tr) boundary is marked by the most severe mass extinction in the geologic record, but the time interval over which the extinctions occurred has been a subject of much debate. Published biostratigraphic data from P-Tr boundary sections in the southern Alps, Italy, were used to test the effects of sampling and species abundance on the record of the latest Permian extinction. The results for foraminiferal taxa match simulations for abrupt extinctions; the extinction level is close to the base of the Tesero horizon of the Werfen Formation. Using biostratigraphic estimates of sedimentation rates, we constrain the extinction interval to
<10 k.y.

High-resolution cyclostratigraphy on a 10^4 yr scale across the P-Tr boundary in a core from the Carnic Alps (Austria) shows significant cycles in the ratio ~40:10:4.7:2.3 m, identified with Milankovitch cycles of ~412:100:40:20 k.y. (eccentricity 1 and 2, obliquity, and precession). Cycle analysis indicates continuity of deposition across the P-Tr boundary, and an average accumulation rate of ~10 cm/k.y. The results define the dramatic faunal shift across the boundary within an interval of <8 k.y., and the accompanying sharp negative global carbon isotope shift within <30 k.y., suggesting a catastrophic cause.
Different triggers for the two pulses of mass extinction across the Permian and Triassic boundary | Scientific Reports
Widespread ocean anoxia has been proposed to cause biotic mass extinction across the Permian–Triassic (P–Tr) boundary. However, its temporal dynamics during this crisis period are unclear. The Liangfengya section in the South China Block contains continuous marine sedimentary and fossil records. Two pulses of biotic extinction and two mass extinction horizons (MEH 1 & 2) near the P–Tr boundary were identified and defined based on lithology and fossils from the section. The data showed that the two pulses of extinction have different environmental triggers. The first pulse occurred during the latest Permian, characterized by disappearance of algae, large foraminifers, and fusulinids. Approaching the MEH 1, multiple layers of volcanic clay and yellowish micritic limestone occurred, suggesting intense volcanic eruptions and terrigenous influx. The second pulse occurred in the earliest Triassic, characterized by opportunist-dominated communities of low diversity and high abundance, and resulted in a structural marine ecosystem change. The oxygen deficiency inferred by pyrite framboid data is associated with biotic declines above the MEH 2, suggesting that the anoxia plays an important role.


Lethally Hot Temperatures During the Early Triassic Greenhouse
Too-Hot Times
Climate warming has been invoked as a factor contributing to widespread extinction events, acting as a trigger or amplifier for more proximal causes, such as marine anoxia. Sun et al. (p. 366; see the Perspective by Bottjer) present evidence that exceptionally high temperatures themselves may have caused some extinctions during the end-Permian. A rapid temperature rise coincided with a general absence of ichthyofauna in equatorial regions, as well as an absence of many species of marine mammals and calcareous algae, consistent with thermal influences on the marine low latitudes. Sea surface temperatures approached 40°C, which suggests that land temperatures likely fluctuated to even higher values that suppressed terrestrial equatorial plant and animal abundance during most of the Early Triassic.

Abstract
Global warming is widely regarded to have played a contributing role in numerous past biotic crises. Here, we show that the end-Permian mass extinction coincided with a rapid temperature rise to exceptionally high values in the Early Triassic that were inimical to life in equatorial latitudes and suppressed ecosystem recovery. This was manifested in the loss of calcareous algae, the near-absence of fish in equatorial Tethys, and the dominance of small taxa of invertebrates during the thermal maxima. High temperatures drove most Early Triassic plants and animals out of equatorial terrestrial ecosystems and probably were a major cause of the end-Smithian crisis.
So the overheating of the P-Tr mass extinction persisted in lesser degree. During the event itself, temperatures rose some 6 C near the equator and more than that away from it, an average of 8 C.
 
Modern snakes evolved from a few survivors of dino-killing asteroid -- ScienceDaily
noting
Evolution and dispersal of snakes across the Cretaceous-Paleogene mass extinction | Nature Communications
Abstract:
Mass extinctions have repeatedly shaped global biodiversity. The Cretaceous-Paleogene (K-Pg) mass extinction caused the demise of numerous vertebrate groups, and its aftermath saw the rapid diversification of surviving mammals, birds, frogs, and teleost fishes. However, the effects of the K-Pg extinction on the evolution of snakes—a major clade of predators comprising over 3,700 living species—remains poorly understood. Here, we combine an extensive molecular dataset with phylogenetically and stratigraphically constrained fossil calibrations to infer an evolutionary timescale for Serpentes. We reveal a potential diversification among crown snakes associated with the K-Pg mass extinction, led by the successful colonisation of Asia by the major extant clade Afrophidia. Vertebral morphometrics suggest increasing morphological specialisation among marine snakes through the Paleogene. The dispersal patterns of snakes following the K-Pg underscore the importance of this mass extinction event in shaping Earth’s extant vertebrate faunas.
The oldest divergences in the family tree of snakes is about 90 - 110 million years ago, in the early Cretaceous. That gave four lineages, and there are several divergences just before or just after the K-Pg mass extinction. So there were as few as four surviving snake species. Needless to say, the survivors proliferated and diversified over the Cenozoic.

Three of the four lineages are of scolecophidians ("worm snakes", blind snakes, or thread snakes), burrowing snakes. The ancestral snake was likely a scolecophidian.

The remaining one of them was an alethinophidian ("true snakes"), and they go back some 100 million years to the mid-Cretaceous.

Its first division was into amerophidians, a small group of snakes including pipesnakes, and afrophidians, including just about all of the better-known snakes. These ones are known for "macrostomy", the ability to stretch their mouths to swallow large prey whole. Snakes in general do that, though non-afrophidians like blind snakes tend to prefer easier-to-swallow prey.

Afrophidians quickly split into henophidians, including boas, pythons, and the like, known for constricting their prey, and caenophidians, the large majority of present-day snake species, including rattlesnakes, cobras, garter snakes, and the like.

Snakes continued to diversify over the Cenozoic.
 
Mass extinction of lizards and snakes at the Cretaceous–Paleogene boundary | PNAS
The Cretaceous–Paleogene (K-Pg) boundary is marked by a major mass extinction, yet this event is thought to have had little effect on the diversity of lizards and snakes (Squamata). A revision of fossil squamates from the Maastrichtian and Paleocene of North America shows that lizards and snakes suffered a devastating mass extinction coinciding with the Chicxulub asteroid impact. Species-level extinction was 83%, and the K-Pg event resulted in the elimination of many lizard groups and a dramatic decrease in morphological disparity. Survival was associated with small body size and perhaps large geographic range. The recovery was prolonged; diversity did not approach Cretaceous levels until 10 My after the extinction, and resulted in a dramatic change in faunal composition. The squamate fossil record shows that the end-Cretaceous mass extinction was far more severe than previously believed, and underscores the role played by mass extinctions in driving diversification.
Squamata = lizards + snakes

Postnatal ontogeny and the evolution of macrostomy in snakes | Royal Society Open Science - "Macrostomy is the anatomical feature present in macrostomatan snakes that permits the ingestion of entire prey with high cross-sectional area." It requires some adaptations in mouth and skull and jawbone anatomy, as one might expect.

"Snakes are gape-limited predators that swallow their prey whole, that is, without mechanical reduction through an intraoral treatment prior to ingestion." - that is, they don't tear apart or bite into pieces or chew their prey. They swallow it whole. One can find some videos online of pet snakes swallowing rats and rabbits whole, and stretching their mouths around their meals.

"Among the wide diversity of extant snakes, basal forms such as scolecophidians (worm-like snakes) and basal alethinophidians (pipe snakes, shieldtail snakes) occupy underground macrohabitats and feed on prey of small size and/or small cross-sectional area, such as insects, earthworms and elongated vertebrates. By contrast, alethinophidian snakes included in the clade Macrostomata (pythons, boas, dwarf boas and colubroids) have developed in extreme this feeding strategy ingesting large prey with large cross-sectional area in relation to their head dimensions."

Some macrostomate snakes have reverted to eating thin prey like earthworms, and their mouth/skull/jawbone features have also reverted.
 
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