More fun with ammonoids

A few years ago I posted on the joyful eccentricities of the heteromorph ammonites, who refused to be bound by conventional notions of what ammonites were supposed to look like. Ammonoids also experimented with some interesting morphological choices in their early days. Here are a couple of examples of inspired ammonoids you might have come across scuba diving in the Late Devonian: 

Below is Parawocklumeria, looking like a Paleozoic premonition of the globigerine foraminifera (although somewhat larger).

Parawocklumeria paradoxa, Plate 19 from Wedekind (1918) (described as Wocklumeria paradoxa)

Despite the inflated appearance, Parawocklumeria is still doing the typical coiling. Another ammonoid, Solicylmenia, seems to have decided that coiling is all well and good, but it's even better when done triangularly. What exactly this taxon got out of this, I'm not sure; it seems to have worked well enough for Soliclymenia but not well enough for anyone else.

Solicylmenia paradoxa, Plate 16 from Münster (1839) (described as Cylmenia paradoxa). I realize the quality isn't great, but on the other hand it's fun to track down the original illustrations. (And yes, both chosen species here happen to be paradoxa; I wasn't being lazy with cut and paste.)

References

Münster, G. Graf zu. 1839. Nachtrag zu den Clymenien des Fichtelgebirges. Beiträge zur Petrefactenkunde 1:35-43.

Wedekind, R. 1918. Die Genera der Palaeoammonoidea (Goniatiten). Palaeontographica 62:85–184, pl. 14–22.

Fossil Felids of the National Park Service

For this year's annual focus on a fossil group in National Park System units, I am going to pay my Internet cat tax and present a quick look at the fossil felids of the parks. If you're not familiar with the history of Felidae, it may come as a surprise that cats are a relatively recent innovation. They have only a few tens of millions of years under their collective belts, and didn't arrive in North America until Pseudaelurus in the early Miocene. (Note: barbourofelids and nimravids may give off saber-toothed cat vibes, but they aren't Felidae.)

It turns out that there are 24 NPS units with cat body or trace fossils. All of the records, unsurprisingly, are early Miocene in age or younger. The great majority are Quaternary (concerning the question of when to cut off the paleontological record, caves don't discriminate if you happen to be 11,000 years old rather than 12,000 years old, and neither do we): only seven of the 24 park units have pre-Q cats. These are Big Bend National Park, Death Valley NP, Hagerman Fossil Beds National Monument, John Day Fossil Beds NM, Lake Mead National Recreation Area, Mojave National Preserve, and Niobrara National Scenic River.

These maps are so much easier to make now that I have one file with all of the parks as points, and I can just turn them on and off. 1. John Day Fossil Beds National Monument; 2. Oregon Caves NM; 3. Lava Beds NM; 4. Hagerman Fossil Beds NM; 5. Yellowstone National Park; 6. Great Basin NP; 7. Death Valley NP; 8. Tule Springs Fossil Beds NM; 9. Lake Mead National Recreation Area; 10. Mojave National Preserve; 11. Joshua Tree NP; 12. Grand Canyon NP; 13. Chaco Culture National Historical Park; 14. White Sands NP; 15. Carlsbad Caverns NP; 16. Guadalupe Mountains NP; 17. Big Bend NP; 18. Amistad NRA; 19. Waco Mammoth NM; 20. Niobrara National Scenic River; 21. Ozark National Scenic Riverways; 22. Chickamauga & Chattanooga National Military Park; 23. Potomac Heritage National Scenic Trail; 24. Valley Forge NHP.

Geographically the sites are concentrated in the southwestern US; in fact, 15 of the sites are in Arizona, California, Nevada, New Mexico, or Texas. This seems to say something about cat biogeography, given these are mostly Quaternary sites and the NPS has a pretty good Quaternary record in terms of geographic spread. I wouldn't base a thesis on it or anything, though.

It's not stated on the map, but the Quaternary record is heavy on caves; a dozen of the records are from caves or rock shelters. These included significant records at Carlsbad Caverns NP, Grand Canyon NP, Guadalupe Mountains NP, Potomac Heritage NST (Cumberland Bone Cave), and Valley Forge NHP (Port Kennedy Bone Cave).

Most of the records are body fossils, but at least three have cat trace fossils: tracks at Death Valley NP and White Sands NP, and tracks and cave scratches at Chickamauga & Chattanooga NMP. One of these sites has yielded a track type specimen, Felipeda scrivneri Sarjeant et al. (2002) from Death Valley NP. There are also eight fossil felid species named from body fossils discovered within or potentially within NPS areas (all named before the units were established):

• Felis lacustris Gazin (1933) from Hagerman (now Puma lacustris)
• Machairodus? hesperus Gazin (1933) from Hagerman (now Megantereon hesperus)
• Felis augustus Leidy (1872) possibly from Niobrara (now Panthera onca [augusta], and Pleistocene instead of Miocene in age)
• Felis (Pseudaelurus) intrepidus Leidy (1858) possibly from Niobrara (now Pseudaelurus intrepidus)
• Crocuta inexpectata Cope (1895) from Valley Forge (now Miracinonyx inexpectatus) 
• Lynx calcaratus Cope (1899) from Valley Forge (now considered a synonym of Lynx rufus)
• Smilodon gracilis Cope (1880) from Valley Forge
• Uncia mercerii Cope (1895) from Valley Forge (now considered a synonym of Smilodon gracilis)

Attaining consensus on cat taxonomy and nomenclature can be like, well, herding cats. It doesn't help that it can be difficult to tell cats apart; see a record of "Panthera onca" at Carlsbad Caverns NP becoming Panthera atrox (Kottkamp et al. 2022), and "Puma concolor" fossils at Grand Canyon NP becoming Miracinonyx trumani (Hodnett et al. 2022; take the skull for a spin here). Nevertheless, the Quaternary sample can be divided among seven species or species groups. Three are extinct:

• American cheetah (Miracinonyx inexpectatus and M. trumani): found at Carlsbad, Grand Canyon, Potomac Heritage, and Valley Forge
• American lion (Panthera atrox): found at Carlsbad, Potomac Heritage, and Tule Springs
• Saber-toothed cats (Smilodon spp.): found at Potomac Heritage, Tule Springs, Valley Forge, and Waco Mammoth

Two are still around but not this far north:

• Jaguar (Panthera onca): found at Lava Beds, Oregon Caves, Ozark, Potomac Heritage, and Valley Forge (these are all pretty far north for something we associate with tropical jungles!)
  
• Jaguarundi (Herpailurus yagouaroundi): found at Valley Forge

Finally, two are still found in the United States:

• Bobcat (Lynx rufus, with allowance for Lynx sp.): Amistad, Carlsbad Caverns, Chaco Culture*, Great Basin*, Grand Canyon, Guadalupe Mountains, Joshua Tree*, Lava Beds, Tule Springs, Valley Forge, and Yellowstone* (*=Holocene only)
• Cougar/mountain lion/puma (Puma concolor): Carlsbad Caverns, Chaco Culture*, Guadalupe Mountains, and Tule Springs

The most species-rich sites are:

• Carlsbad Caverns NP (American cheetah, American lion, bobcat, cougar)
• Hagerman Fossil Beds NM (American cheetah, Homotherium sp. [a saber-toothed cat], Lynx rexroadensis, Megantereon hesperus [or cultridens; another saber-toothed cat], Puma lacustris)
• Tule Springs Fossil Beds NM (American lion, bobcat, cougar, saber-toothed cat)
• Valley Forge NHP (American cheetah, bobcat, jaguar, jaguarundi, saber-toothed cat)

Of these four, the Hagerman assemblage is Pliocene, the Valley Forge assemblage is middle Pleistocene, and the other two are late Pleistocene. In the Pleistocene, at least, a robust cat assemblage may include a big lion-type cat, a saber-toothed cat, a smaller big cat (but apparently not cougars and jaguars at the same place), and a bobcat-sized cat.

Felids are relatively uncommon compared to the other big terrestrial carnivorans. Fossil dogs and bears are more widely distributed in NPS units than cats. Furthermore, all but one NPS assemblage that has cats also has dogs, bears, or both (the exception being the Pliocene tracks of Death Valley NP). These groups, though, are for another time...

References

Cope, E. D. 1880. On the extinct cats of America. American Naturalist 14(12):833–858.

Cope, E. D. 1895. The fossil Vertebrata from the fissure at Port Kennedy. Proceedings of the Academy of Natural Sciences of Philadelphia 47:446–450.

Cope, E. D. 1899. Vertebrate remains from Port Kennedy bone deposit. Journal of the Academy of Natural Sciences of Philadelphia, 2nd series, 11(3):193–286.

Gazin, C. L. 1933. New felids from the upper Pliocene of Idaho. Journal of Mammalogy 14:251–356.

Hodnett, J. P., R. White, M. Carpenter, J. Mead, and V. L. Santucci. 2022. Miracinonyx trumani (Carnivora; Felidae) from the Rancholabrean of the Grand Canyon, Arizona and its implications on the ecology of the “American cheetah.” New Mexico Museum of Natural History Bulletin 88:157–186.

Kottkamp, S., V. L. Santucci, J. S. Tweet, R. D. Horrocks, and G. S. Morgan. 2022. Pleistocene vertebrates from Carlsbad Caverns National Park, New Mexico. New Mexico Museum of Natural History and Science Bulletin 88:267–290.

Leidy, J. 1858. Notice of remains of extinct Vertebrata, from the valley of the Niobrara River, collected during the exploring expedition of 1857, in Nebraska, under the command of Lieut. G. K. Warren, U.S. Top. Eng., by Dr. F. V. Hayden, Geologist to the expedition. Proceedings of the Academy of Natural Sciences of Philadelphia 10:20–29.

Leidy, J. 1872. Remarks on some extinct vertebrates. Proceedings of the Academy of Natural Sciences of Philadelphia 24:38–40.

Sarjeant, W. A. S., R. E. Reynolds, and M. M. Kissell-Jones. 2002. Fossil creodont and carnivore footprints from California, Nevada, and Wyoming. Pages 37–50 in R. E. Reynolds, editor. Between the Basins: Exploring the western Mojave and southern Basin and Range Province. California State University, Fullerton, California.