Triassic origin of scales, scutes, hair, etc. as biting fly barriers?

During the Middle to Late Triassic

  1. Mammals developed fur/hair.
  2. Aetosaurs developed plates and horns beyond the earlier paired dorsal scutes.
  3. Crocodylomorphs developed large scales beyond the earlier paired dorsal scutes.
  4. Dinosaurs lost those paired scutes and developed placodes and quills. Ultimately these became scales and feathers.
  5. Turtles developed hard scales over a carapace and plastron.
  6. Lepidosaurs developed small scales.
  7. Pterosaurs and their Late Triassic sisters developed pycnofibers

All of these developed on the soft, naked skin
(think of a plucked chicken) that was a universal covering for Carboniferous and Permian tetrapods (Early forms retained large ventral scales inherited from finned ancestors, but these were lost by the Permian).

All of these extradermal structures have one thing in common.
They separated and/or protected the animal’s naked skin from the environment, one way or another. They developed by convergence. Dhouailly 2009 and other workers discussed the chemical similarities of the keratin found in these dermal structures. 

The question is:
What was different about the Triassic environment that was not present in earlier Carboniferous and Permian environments? We can 
eliminate heat, cold, UV rays, rain, aridity, etc. as possible reasons for the development of insulator structures because those factors had always been present. So what was new in the Triassic that affected all terrestrial tetrapods?

Flies and their biting, piercing kin.
“The earliest definitive flies known from the mid-Triassic of France, approximately 230 Ma (Krzemiski and Krzeminska, 2003)” according to Blagoderov, Grimaldi and Fraser 2007. The order Diptera (flies, mosquitos and kin) tend to land on large tetrapods for food, blood, etc. Scales, scutes, hair, feathers, etc. all separate flying insects from the naked skin of Triassic terrestrial tetrapods. Williams et al. 2006 even found mosquito repellents in frog skin. It is notable that, except for armored placodonts and mosasaurus (derived varanid lepidosaurs), aquatic and marine tetrapods also had naked skin with the thalattosaur, Vancleavea, a notable sermi-terrestrial exception. Is that because they had aquatic antecedents in the Triassic that were never affected by flying insects?

It’s not just the insect bite that drives this evolution,
it’s the appearance of new vectors for the rapid spread of disease that drives this evolution.

Interesting coincidence.
If this is not the case, this will take further study.

Figure 1. Lacertulus, a basal squamate from the Late Permian

Figure 1. Lacertulus, a basal squamate from the Late Permian

Carroll and Thompson 1982 report
on the Late Permian lepidosaur, Lacertulus (Fig. 1), “No scales dermal or epidermal are evident in the specimen.”

From the Dhouailly 2009 abstract:
“I suggest that the alpha-keratinized hairs from living synapsids may have evolved from the hypothetical glandular integument of the first amniotes, which may have presented similarities with common day terrestrial amphibians.

Concerning feathers, they may have evolved independently of squamate scales, each originating from the hypothetical roughened beta-keratinized integument of the first sauropsids. The avian overlapping scales, which cover the feet in some bird species, may have developed later in evolution, being secondarily derived from feathers.” Not realized by Dhouailly, the purported clade ‘Sauropsida’ is paraphyletic and a junior synonym for Amniota and Reptilia in the LRT.

Earlier we looked at the first appearances
of hair, quills, pycnofibers and hard scales in a three-part series here, here and here

Exceptionally, humans are terrestrial tetrapods
that have lost most of their hair, more or less returning to the primitive naked state. And yes, flies and mosquitos do bother humans. It is the price we pay for the benefits of naked skin. Clothing helps provide a barrier.

Just because an idea is proposed and a hypothesis is advanced doesn’t make it so. In science ideas have to be confirmed or refuted following their first appearance. If anyone has data concerning scales or other dermal structures in Carboniferous or Permian taxa, please make us aware of those.

Blagoderov V, Grimaldi D and Fraser NC 2007. How Time Flies for Flies: Diverse Diptera from the Triassic of Virginia and Early Radiation of the Order. American Museum Novitates 3572:1-39. DOI: 10.1206/0003-0082(2007)509[1:HTFFFD]2.0.CO;2
Carroll RL and Thompson P 1982.
A bipedal lizardlike reptile from the Karroo. Journal of Palaeontology 56:1-10.
Dhouailly D 2009.
A new scenario for the evolutionary origin of hair, feather, and avian scales Journal of Anatomy 214(4): 587–606. doi: 10.1111/j.1469-7580.2008.01041.x
Krzeminnski, W., and E. Krzeminska. 2003. Triassic Diptera: descriptions, revisions and phylogenetic relations. Acta Zoologica Cracoviensia (suppl.) 46: 153–184.
Maderson PFA and Alibardi L 2000.
The Development of the Sauropsid Integument: A Contribution to the Problem of the Origin and Evolution of Feathers. American Zoologist 40:513–529.
Rohdendorf BB, Oldroyd H and Ball GE 1974. The Historical Development of Diptera. The University of Alberta Press, Edmonton, Canada. ISBN 0-88864-003-X.
Williams CR, Smith BPC, Best SM and Tyler MJ 2006.
Mosquito repellents in frog skin. Biol Lett. 2006 Jun 22; 2(2): 242–245. doi: 10.1098/rsbl.2006.0448


Mammal taxa: origin times

A few days ago, we looked at a revised and expanded cladogram of the Mammalia based on skeletal traits (distinct from and contra to a cladogram based on DNA). Today we add chronology to the cladogram to indicate the first appearance of various mammals and estimate the origin of the various clades (Fig. 1).

Note that derived taxa
that chronologically precede more primitive taxa indicate that primitive taxa had their genesis and radiation earlier than the first appearance of fossil specimens, which always represent rare findings usually during wide radiations that increase the chance the specimen will fossilize in the past and be found in the present day.

Looking at time of mammal taxa origin categories:

Figure 1. Cladogram with time notes for the Mammalia (subset of the LRT).

Figure 1. Cladogram with time notes for the Mammalia (subset of the LRT).

Some notes:

  1. Both prototheres and basal therians were present (and probably widespread) in the Late Triassic.
  2. Derived prototheres appear in the Late Triassic, suggesting an earlier (Middle Triassic?) origin for Mammalia and an earlier (Middle Triassic?) split between Prototheria and Theria.
  3. Both fossorial metatherians and basal arboreal eutherians were present (and probably widespread) in the Late Jurassic. These were small taxa, out of the gaze of ruling dinosaurs.
  4. Large derived eutherians eolved immediately following the K-T boundary in the Paleocene and radiated throughout the Tertiary.
  5. A large fraction of prototherians, metatherians and eutherians are known only from extant taxa, some of which are rare and restricted, not widespread.
  6. Multituberculates and kin are derived placentals close to rodents by homology, not convergence.


Bird, pterosaur, dinosaur simplified chronology

Following the earlier post on non-arboreal post K-T boundary birds…

…this one pretty much speaks for itself.
Here (Fig. 1) is a chronology, very much simplified, of birds, pterosaurs and dinosaurs according to the LRT.

Figure 1. Mesozoic chronology of bird, dinosaur and pterosaur clades.

Figure 1. Mesozoic chronology of bird, dinosaur and pterosaur clades based on taxa in the LRT.

If you’re curious about any of the taxa,
in the chronology, simply use Keywords to locate them.

What is Dyoplax?

Figure 1. Dyoplax arenaceus Fraas 1867 is a mold fossil recently considered to be a sphenosuchian crocodylomorph. Here it nests as a basal metriorhynchid (sea crocodile) in the Late Triassic.

Figure 1. Dyoplax arenaceus Fraas 1867 is a mold fossil recently considered to be a sphenosuchian crocodylomorph. Here it nests as a basal metriorhynchid (sea crocodile) in the Late Triassic.

Dyoplax arenaceus (Fraas 1867, Lucas, Wild and Hunt 1998) is a unique Late Triassic crocodylomorph, one of the three original crocodylomorphs in the 19th century along with two aetosaurs.

No bones are present.
Like Cosesaurus it is a natural cast mold. Fraas thought it had the head of a lizard, but the armor of a gavial. Lucas et al. nested it as the oldest sphenosuchian crocodylomorph. Maish et al. nested it with Erpetosuchus.

Dyoplax nests as a basal metriorhynchid, those Jurassic sea crocodiles with flippers for fore limbs and a curved fish-like tail (Fig. 2), which was probably too early to be present. Unfortunately the tail tip and limb tips are missing from the fossil (Fig. 1)

The fossil appears to have no arch of bones separating the upper and lateral temporal fenestrae, but the intervening squamosal appears to be flipped and displaced near the neck ribs. (Black arrow in fig. 1)

Figure 2. Several Jurassic sea crocs, apparently derived from Late Triassic Dyoplax.

Figure 2. Several Jurassic sea crocs, apparently derived from Late Triassic Dyoplax.

Boy, it’s been awhile
since I posted anything. Feels good to be back… but for how long?

Fraas O 1867. Dyoplax arenaceus, ein neuer Stuttgarter Keuper-Saurier. Jh. Verein vaterländ. Naturk. Württemberg 23:108-112; Stuttgart.
Lucas SG, Wild R, Hunt AP 1998. Dyoplax O. Fraas, a Triassic sphenosuchian from Germany. Stuttgarter Beiträge zur Naturkunde, B. 263: 1–13.
Maisch MW;  Matzke AT; Rathgeber T 2013. Re-evaluation of the enigmatic archosaur Dyoplax arenaceus O. Fraas, 1867 from the Schilfsandstein (Stuttgart Formation, lower Carnian, Upper Triassic) of Stuttgart, Germany. Neues Jahrbuch für Geologie und Paläontologie – Abhandlungen. 267 (3): 353–362.


What is the enigmatic Otter Sandstone (Middle Triassic) diapsid?

Coram, Radley and Benton 2017
presented a “small diapsid reptile [BRSUG 29950-12], possibly, pending systematic study, a basal lepidosaur or a protorosaurian.” According to Coram et al. “The Middle Triassic (Anisian) Otter Sandstone was laid down mostly by braided rivers in a desert environment.”

Figure 1. The Middle Triassic Otter Sandstone diapsid BRSUG 29950-12 under DGS nested with basalmost lepidosaurs like Megachirella.

Figure 1. The Middle Triassic Otter Sandstone diapsid BRSUG 29950-12 under DGS nested with basalmost lepidosaurs like Megachirella. Skeleton is exposed in ventral (palatal) view.

The LRT is here to nest and identify published enigmas
The large reptile tree (LRT 1041 taxa) nests BRSUG 29950-12 with the basalmost lepidosaur Megachirella. They are a close match and preserve nearly identical portions of their skeletons (Fig. 2). Megachirella was originally considered a sister to Marmoretta, another basal sphenodontian from the much later Middle/Late Jurassic.

FIgure 2. Megachirella (Renesto and Posenato 2003) is a sister to the BSRUG diapsid.

FIgure 2. Megachirella (Renesto and Posenato 2003), also from Middle Triassic desposits, is a sister to the BSRUG diapsid and provides a good guide for its eventual reconstruction.

At the base of the Lepidosauria
in the LRT nests Megachirella, derived from a sister to Sophineta (Early Triassic) and Saurosternon + Palaegama (Latest Permian) and kin. Sisters to Megachirella within the Lepidosauria include the tritosaurs Tijubina + Huehuecuetzpalli (Early Cretaceous), Macrocnemus (Middle Triassic) and the prosquamate Lacertulus (Late Permian). Also similar and related to Palaegama is Jesairosaurus (Middle Triassic). So the genesis of the Lepidosauria is Late Permian. The initial radiation produced taxa that continued into the Early Cretaceous. The radiation of derived taxa continued with three major clades, only one of which, the Tritosauria, is now completely extinct.

It is important to remember that lepdiosaurs and protorosaurs are not closely related, but arrived at similar bauplans by convergence, according to the LRT. The former is a member of the new Lepidosauromorpha. The latter is a member of the new Archosauromorpha. Last common ancestor: Gephyrostegus and kin.

Nesting at the base of the Lepidosauria
in the Sphenodontia clade makes the BSRUG specimen an important taxon. Let’s see if and when this taxon is nested by academic workers that they include all of the pertinent taxa and confirm or re-discover the Tritosauria. The LRT provides a good list of nearly all of the pertinent taxa that should be included in that future study, many of which are listed above. Based on that list, the BSRUG specimen is a late-survivor of a perhaps Middle Permian radiation of basal lepidosaurs.

Coram RA, Radley JD and Benton MJ 2017. The Middle Triassic (Anisian) Otter Sandstone biota (Devon, UK): review, recent discoveries and ways ahead. Proceedings of the Geologists’ Association in press.

Arcticodactylus a tiny Greenland Triassic pterosaur

Arcticodactylus cromptonellus (Kellner 2015, originally Eudimorphodon cromptonellus Jenkins et al. 1999, 1999; MGUH VP 3393) Late Triassic ~210mya ~8 cm snout to vent length was a tiny pterosaur derived from a sister to Eudimorphodon ranzii and phylogenetically preceded Campylognathoides and BSp 1994 specimen attributed to Eudimorphodon. Whether it was a juvenile or a tiny adult cannot be determined because juveniles and even embryos are identical to adults in pterosaurs. Note that that rostrum was not shorter and the orbit was not larger than in sister taxa. This specimen is one of the smallest known pterosaurs., but not THE smallest (Fig. 1) contra the Wikipedia article. That honor goes to B St 1967 I 276.

Figure 1. Articodactylus is evidently NOT the smallest pterosaur. That honor still goes to an unnamed specimen (not a Pterodactylus kochi juvenile) B St 1967 I 276.

Figure 1. Articodactylus is evidently NOT the smallest pterosaur. That honor still goes to an unnamed specimen (not a Pterodactylus kochi juvenile) B St 1967 I 276.

Distinct from E. ranzii,
the skull of Arctiodactylus had a rounder, less triangular orbit. The jugal was not as deep. The sternal complex did not have small lateral processes. The humerus was not as robust. The fingers were longer an more gracile. The prepubis was distinctly shaped.

Distinct from
Bergamodactylus the femur and tibia were smaller but the metatarsals were longer, compact and nearly subequal in length with IV smaller than III.

Jenkins FA Jr, Shubin NH, Gatesy SM and Padian K 1999. A primitive pterosaur of Late Triassic age from Greenland. Journal of the Society of Vertebrate Paleontology 19(3): 56A.
Jenkins FA Jr, Shubin NH, Gatesy SM and Padian K 1999. A diminutive pterosaur (Pterosauria: Eudimorphodontidae) from the Greenlandic Triassic. Bulletin of the Museum of Comparative Zoology, Harvard University 155(9): 487-506.
Kellner AWA 2015. Comments on Triassic pterosaurs with discussion about ontogeny and description of new taxa. Anais da Academia Brasileira de Ciências 87(2): 669–689


Rough chronology of basal tetrapods and basal reptiles

Today we’ll look at WHEN
we find fossils of basal tetrapods and basal reptiles. According to the large reptile tree (959 taxa, LRT, subset shown in Fig. 1), oftentimes we find late survivors of earlier radiations in higher strata. The origin of Reptilia (amphibian-like amniotes) extends back to the Devonian and Early Carboniferous now, not the Late Carboniferous as Wikipedia reports and as the Tree of Life project reports.

Figure 1. Color coded chronology of basal tetrapods and reptiles.We're lucky to know these few taxa out of a time span of several tens of millions of years.

Figure 1. Color coded chronology of basal tetrapods and reptiles.We’re lucky to know these few taxa out of a time span of several tens of millions of years. Click to enlarge.

The Late Devonian 390–360 mya
Here we find late survivors of an earlier radiation: Cheirolepis, a basal member of the Actinopterygii (ray-fin fish) together with Eusthenopteron and other members of the Sarcopterygii (lobe-fin fish). Coeval are basal tetrapods, like Acanthostega and basal reptiles, like Tulerpeton. These last two launch the radiations we find in the next period. The presence of Tulerpeton in the Late Devonian tells us that basal Seymouriamorpha and Reptilomorpha are waiting to be found in Devonian strata. We’ve already found basal Whatcheeriidae in the Late Devonian taxa Ichthyostega and Ventastega.

Early Carboniferous 360–322 mya
Here we find the first radiations of basal reptilomorphs, basal reptiles, basal temnospondyls,  basal lepospondyls and microsaurs, lacking only basal seymouriamorphs unless Eucritta is counted among them. It nests outside that clade in the LRT.

Late Carboniferous 322–300 mya
Here we find more temnospondyls, lepospondyls and phylogenetically miniaturized archosauromorphs, likely avoiding the larger predators and/or finding new niches. Note the first prodiapsids, like Erpetonyx and Archaeovenator, appear in this period, indicating that predecessor taxa like Protorothyris and Vaughnictis had an older, Late Carboniferous, origin. Not shown are the large basal lepidosauromorphs, Limnoscelis and Eocasea and the small archosauromorphs, Petrolacosaurus and Spinoaequalis.

Early Permian 300–280 mya
Here we find the first fossil Seymouriamorpha and the last of the lepospondyls other than those that give rise to extant amphibians, like Rana, the frog. Here are further radiations of basal Lepidosauromorpha, basal Archosauromorpha (including small prodiapsids), along with the first radiations of large synapsids.

Late Permian 280–252 mya
Here we find the next radiation of large and small synapsids, the last seymouriamorphs, and derived taxa not shown in the present LRT subset.

Early/Mid Triassic 252 mya–235 mya
Among the remaining basal taxa few have their origins here other than therapsids close to mammals. Afterwards, the last few basal taxa  listed here, principally among the Synapsida, occur later in the Late Triassic, the Jurassic and into the Recent. Other taxa are listed at the LRT.

What you should glean from this graphic
Taxa are found in only the few strata where fossilization occurred. So fossils are incredibly rare and somewhat randomly discovered. The origin of a taxa must often be inferred from phylogenetic bracketing. And that’s okay. This chart acts like a BINGO card, nesting known taxa while leaving spaces for taxa we all hope will someday fill out our card.