Flamingo teeth

Figure 1. The picture says it all. Like ducks and Pelagornis, pseudo teeth appear in flamingos. Here they are used for filtering. Compare these jaws to those of the right whale, Balaena.

Figure 1. The picture says it all. Like ducks and Pelagornis, pseudo teeth appear in flamingos. Here they are used for filtering. Compare these jaws to those of the right whale, Balaena.

No, they’re not real teeth,
But they act like baleen to filter out tiny brine shrimp and blue-green algae. According to Wikipedia, “Their bills are specially adapted to separate mud and silt from the food they eat, and are uniquely used upside-down. The filtering of food items is assisted by hairy structures called lamellae which line the mandibles, and the large rough-surfaced tongue.”

Figure 2. Phoenicopterus, the flamingo, sometimes enjoys the beach.

Duck teeth
(Fig. 3) are not real teeth either.

Figure 3. Anas, the mallard duck, shares more trait with Aepyornis than with other taxa in the LRT.

Figure 3. Anas, the mallard duck, shares more trait with Aepyornis than with other taxa in the LRT.

Pelagornis teeth
(Fig. 4) are not real teeth either. But, brother they look ral.

Figure 1. Pelagornis skeletal elements.

Figure 4. Pelagornis skeletal elements.

Hesperornis teeth|
(Fig. 5) are real teeth.

Figure 2. Hesperornis skull. Compare this to that of Pelagornis in figure 1.

Figure 5. Hesperornis skull. Compare this to that of Pelagornis in figure 1.

Oscar Reig: a paleoprophet separates archosaurs from lepidosaurs in 1967

But… for the wrong reasons.

Reig 1967 prophetically wrote:
“Archosaurs and lepidosaurs apparently have different origins; the former come from the pelycosaurs, and the latter come from the captorhinomorph cotylosaurs through the Millerettiformes.”

Considered heretical at the time,
Reig’s pronouncement echoes in the large reptile tree (LRT, 1151 taxa).

Here’s the full abstract:
“The characteristics of the first archosaurs, the proterosuchian thecodonts, show that neither of the supposed common ancestors of archosaurs and lepidosaurs could actually be an ancestor of archosaurs. Instead, the evidence seems to indicate that the archosaurian ancestors are probably in the ophiacodont-varanopsid group of the pelycosaurian synapsids. In particular, the Varanopsidae are strongly indicative of proterosuchian relationships, as they have evolved some characters which are elsewhere found only in archosaurs. Archosaurs and lepidosaurs apparently have different origins; the former come from the pelycosaurs, and the latter come from the captorhinomorph cotylosaurs through the Millerettiformes.”

The only thing he got wrong
(as everyone else got wrong until seven years ago) was not splitting the Varanopsidae into the Synapsida and the Prodiapsida, as demonstrated in the LRT. He also thought proterosuchids arose directly from varanopsids like Varanodon (Fig. 1), which converge with proterosuchids in size and skull shapes. There’s even an antorbital fenestra, or elongated naris and a drooping premaxilla in Varanodon. No wonder Reig got excited.

Figure 1. Varanodon the synapsid compared to its analog, Proterosuchus, the archosauriform.

Figure 1. Varanodon the synapsid compared to its analog, Proterosuchus, the archosauriform.

Archosauriforms do arise from
former
varanopsids, like Heleosaurus and Mycterosaurus, but not directly. They have to pass through the diapsid grade, then the basal terrestrial younginiform grade before evolving into proterosuchids.

Lepidosaurs do arise from
captorhinomorphs and millerettids in the LRT, but again, not directly. First they have to pass through the nycteroleterid, owenettid, and basal lepidosauriform grades before evolving into lepidosaurs.

The LRT recovered
two clades of diapsids one closer to lepidosaurs and another closer to archosaurs.

References
Reig OA 1967. Archosaurian reptiles: a new hypothesis on their origins.
Science 157(3788):565-8.

Harrisonavis and the origin of flamingos

Harrisonavis croizeti (Torres et al. 2015, Fig. 1) is very clearly an Oligocene flamingo.  Other than the angle of its beak, it is a close match to the living flamingo, Phoenicopterus (Fig. 1).

Figure 1. Click to enlarge. Candidate taxa in the ancestry of flamingos. Most bird experts like long-necked Paleolodus, a tinamou in the LRT. Ignored is Cariama, which shares more traits with flamingos in the LRT.

Figure 1. Click to enlarge. Candidate taxa in the ancestry of flamingos. Most bird experts like long-necked Paleolodus, a tinamou in the LRT. Ignored is Cariama, which shares more traits with flamingos in the LRT.

The trouble comes when you try to delve deeper into flamingo ancestry.
Bird experts, like Mayr 2004, say Palaelodus (Fig. 1 ) is the next outgroup to flamingos. Well, in a way, it is, but the large reptile tree (LRT, 1051 taxa, Fig. 2) nests Palaeodus with Struthio, the ostrich, just outside of the clade that starts with Phoenicopterus, the flamingo, and Cariama, the seriema. So, it’s a close one! Palaelodus was the last of the flying ostriches. All of the basalmost neognaths had long, stilt-like legs.

Figure 1. Subset of the LRT focusing on birds. Here various aspects of birds are shown, including age, teeth, feeding behavior and basic clades.

Figure 2. Subset of the LRT focusing on birds. Here various aspects of birds are shown, including age, teeth, feeding behavior and basic clades.

Figure 1. Nearly proportioned like its giant descendant, Palaeotis was an Eocene ostrich less than 1/3 as tall.

Figure 3. Nearly proportioned like its giant descendant, Palaeotis was an Eocene ostrich less than 1/3 as tall. Compare to Palaelodus in figure 1 and 3.

References
Linnaeus C von 1758. Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata.
Linneaus C 1766. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio duodecima, reformata. Holmiae. (Laurentii Salvii).: 1-532.
Mayr G 2004. Morphological evidence for sister group relationship between flamingos (Aves: Phoenicopteridae) and grebes (Podicipedidae). Zoological Journal of the Linnean Society. 140 (2): 157–169. doi:10.1111/j.1096-3642.2003.00094.x. ISSN 0024-4082.
Molina JI 1782. Saggio sulla Storia Naturale del Chili. Bologna, Stamperia di S. Tommaso d’Aquino. 349 pp.
Olson SL and Feduccia A 1980. Relationships and evolution of flamingos (Aves: Phoenicopteridae). Smithsonian Contributions to Zoology 316: 1–73.
Torres CR, De Pietri VL, Louchart A and van Tuinen M 2015. New cranial material of the earliest filter feeding flamingo Harrisonavis croizeti (Aves, Phoenicopteridae) informs the evolution of the highly specialized filter feeding apparatus. Organisms, Diversity & Evolution. DOI: 10.1007/s13127-015-0209-7

wiki/Seriema
wiki/Flamingo
wiki/Phoenicopteridae

Palaeotis, a tiny Eocene flightless ostrich

Figure 1. Nearly proportioned like its giant descendant, Palaeotis was an Eocene ostrich less than 1/3 as tall.

Figure 1. Nearly proportioned like its giant descendant, Palaeotis was an Eocene ostrich less than 1/3 as tall. Evidently there were larger specimens of the larger winged Palaelodus, but I can only find data for this one at present.

Paelotis weigelti (Lambrecht 1928; middle Eocene; GM 4362) was originally described as an extinct bustard, like Ardeotis. Later (Houde and Haubold 1987) it was referred to the ostrich clade. The wings are small, but the sternum remains large on Paelotis. The pelvis has not attained its larger posterior size yet. Those portions anchor powerful leg muscles on the Struthio, the ostrich. The skull was similar in most respects, but was proportionately larger, like that of a juvenile ostrich. As in Struthio, the scapulocoracoid is fused. The femur was relatively longer. Three toes were present.

Figure 2. Palaeotis untangled, placed into an in vivo pose. Not sure how many pedal digits were present.

Figure 2. Palaeotis untangled, placed into an in vivo pose. Pedal digit 2 is not shown. Note the long humerus and short antebrachium. The bill remains relatively narrow. The sternum remains big.

Figure 3. Palaeotis in situ.

Figure 3. Palaeotis in situ. Note the short right metatarsus. The proximal part must be dislocated beneath the tibia.

References
Houde P and Haubold H 1987. Paleotis weigelti  restudied: a small middle Eocene ostrich (Aves: Struthioniformes) Palaeovertebrata, Montpellier 17(2):27–42.
Lambrecht K 1928. Palaeotis weigelti n. g. sp., eine fossil trappe aus der mitteleozanen Braunkohle des Geiseltales. Jahrbuch hallesch. Verband., Halle, n.s., 7:11.

Back to that elusive completely resolved LRT

You know
I’m constantly adding taxa to the large reptile tree (LRT, 1147 taxa) and that every addition affects and is affected by every other taxa as the software seeks to find the most parsimonious combination of scores that ultimately should reflect evolutionary events in nature. I’m also constantly making corrections as I see them (sort of like Fig. 1) a YouTube video you can click to view. (This was considered the entertainment in the 1960s, shortly after the wide adoption of color TV).

Recently I fell behind
and did not realize the tree had lost resolution. That happens because I only test relevant parts (like birds) when I add in a new taxon. The problem is, when you take the relevant subsets down to examine the issues, you might get, as I did, your resolution back. The taxa are bing affected from afar.

It came to my attention
that the LRT, as a whole unit, had some issues. Only a few bad scores in a cladogram of this size can rapidly and algorithmically multiply the number of MPTs (most parsimonious trees). They’re easy to find. I only need to know to look for them.

Most of the issues were centered around
basal mammals and their pre-mammal analogs.

Now everything is back in order.
All the plates are spinning again. The tree is fully resolved, as I just tested it in whole, not in part.

Bottom line:
Admit your errors. Fix them. Keep moving forward. Watch out for more errors creeping in.

My what big eyes you have, Lyriocephalus!

Looking like some sort of medieval fever dream,
meet Lyriocephalus, the hump-nosed lizard (Fig. 1), a cousin to Draco, the gliding lizard. Distinct from Draco, the body of this insectivore is laterally compressed, not laterally extended.

Figure 1. Lyriocephalus in vivo.

Figure 1. Lyriocephalus in vivo.

Probably the largest eyes
relative to the skull of any tetrapod. Lyriocephalus, is an arboreal jungle lizard with an anterodorsal naris and a small antorbital fenestra. Note the arching postorbital contacting the prefrontal.

Figure 2. Lyriocephalus skull in several views. Note the arching of the postorbital to contact the prefrontal.

Figure 2. Lyriocephalus skull in several views. Note the arching of the postorbital to contact the prefrontal. And did I mention that antorbital fenestra?

Lyriocephalus scutatus (Merrem 1820) is represented by a skeleton at Morphospace.org where you can rotate the skeleton on your monitor. Note the brevity of the tail of this agamid iguanid, There are more in vivo pix here. And a video here.

Figure 3. Lyriocephalus skeleton from Morphobank.org, where you can rotate digitized skeletons.

Figure 3. Lyriocephalus skeleton from Morphobank.org, where you can rotate digitized skeletons.

References
Merrem B 1820. Versuch cines Systems Amphihien Tentamen Systcmatis Amphibiorum. Marburg, Krieger.

wiki/Draco
wiki/Lyriocephalus

Taxa in the heron-cuckoo clade

As earlier, here are a selection of extant taxa in a subset of the LRT in their evolutionary order. This is the heron-cuckoo clade (Figs. 1, 2).

Figure 1. Taxa in the parallel heron and cuckoo clades.

Figure 1. Taxa in the parallel heron and cuckoo clades, all within a single clade.

Eudromius – long-leg terrestrial omnivore (paleognath clade)

Sagittarius – stilt-leg, hook-beak terrestrial predator (hawk/owl clade)

Heron clade

Ardeotis – stilt-leg, straight-beak terrestrial predator

Ciconia – stilt-leg, straight-beak terrestrial predator

Butroides – stilt-leg, straight-beak terrestrial predator

Ardea – stilt-leg, straight-beak terrestrial predator

Cuckoo clade

Psophia – stilt-leg, curved-beak terrestrial omnivore

Menura – stilt-leg, curved-beak terrestrial insectivore

Geococcyx – stilt-leg, straight-beak terrestrial predator/insectivore

Monias – aerial, curved-beak aerial/terrestrial omnivore

Coccyzus – aerial, curved-beak aerial

Figure 1. Subset of the LRT focusing on birds. Here various aspects of birds are shown, including age, teeth, feeding behavior and basic clades.

Figure 1. Subset of the LRT focusing on birds. Here various aspects of birds are shown, including age, teeth, feeding behavior and basic clades.

 

Taxa in the chicken-sparrow-parrot clade

Sometimes it helps to see photos
(Fig. 1) rather than taxonomic names in a cladogram (Fig. 2). See for yourself the evolutionary changes that attend modern representatives of pre- or post-K/T boundary radiation of birds. Seems like when phylogenetic miniaturization happened (with Passer) then the arboreal realm, perching on small twigs, becomes possible. And long legs are primitive, at least until the giants appear.

Figure 1. Taxa in the chicken, sparrow, parrot clade, subset of the LRT.

Figure 1. Taxa in the chicken, sparrow, parrot clade, subset of the LRT.

Eudromius – long-leg terrestrial omnivore (paleognath clade)

Sagittarius – stilt-leg, hook-beak terrestrial predator (hawk/owl clade)

Ardeotis – stilt-leg, straight-beak terrestrial predator (heron/cuckoo clade)

Chauna – long-leg, terrestrial herbivore

Crex – short-leg, terrestrial herbivore

Gallus – short-leg, terrestrial herbivore

Passer – short-leg, miniaturized aerial/arboreal herbivore

Opisthocomus – short-leg, aerial/arboreal herbivore

Ara – short-leg, aerial/arboreal herbivore

Gastornis (Diatryma) and Dinornis – [extinct] giant. long-leg, terrestrial herbivore

Figure 1. Subset of the LRT focusing on birds. Here various aspects of birds are shown, including age, teeth, feeding behavior and basic clades.

Figure 2. Subset of the LRT focusing on birds. Here various aspects of birds are shown, including age, teeth, feeding behavior and basic clades.

 

What are woodpeckers?

Wikipedia reports:
“The Picidae [including woodpeckers] are just one of eight living families in the order Piciformes, the others being barbetstoucans, and honeyguides in the clade Pici, and the jacamars and puffbirds in the clade GalbuliDNA sequencing has confirmed the sister relationships of these two groups.”

Figure 2. Melanerpes, a woodpecker in the LRT, nests with dippers, swifts and wrens.

Figure 2. Melanerpes, a woodpecker in the LRT, nests with dippers, swifts and wrens, all derived from grackles, crows and jays. Nuthatches are sisters to woodpeckers.

Here
the large reptile tree (LRT, 1146 taxa) nests woodpeckers with another vertical bark clinging clade: the wren-sized nuthatches. Outgroups include grackles, crows and jays, several nodes away from deep-billed barbets and toucans.

That’s one more example
demonstrating that DNA fails to duplicate morphology, which literally has to trump DNA because phenomics is the only method that permits the identification of a gradual accumulation of derived traits… and includes fossil taxa. DNA works within genera (like humans), but fails at larger phylogenetic distances. And DNA works in the realm of trust. You hope the results are correct, because there is no way to check them…except by comparing with morphological results.

Figure 2. Sitta the nuthatch skull and in vivo. Pecking insects from tree bark is the first stage in drilling tree bark for insects.

Figure 2. Sitta the nuthatch skull and in vivo. Pecking insects from tree bark is the first stage in drilling tree bark for insects.

Woodpecker and nuthatch skulls are nearly identical
Slender laminations of the maxilla to the tip of the beak help strengthen it’s nail-like tip. Nuthatches do to rotate pedal digit 4 to the back (zygodactyly) like woodpeckers do, so nuthatches are the more primitive of the two clades.

Zygodactyly is convergent several times in neornithine birds.

Figure 3. Skull of the woodpecker, Melanerpes.

Figure 3. Skull of the woodpecker, Melanerpes.

And what about that hyper-hyoid?
The woodpecker’s tongue is anchored by a super-long Y-shaped hyoid. Here (Fig. 4) is the best illustration I have seen about how that works and how it develops.

Figure 4. The woodpecker tongue and hyoid and how it works to extend the tongue into wood to extract burrowing insects.

Figure 4. The woodpecker tongue and hyoid and how it works to extend the tongue into wood to extract burrowing insects.

On the question of DNA vs. Morphology:
A recent reviewer of Feduccia A (2017) wrote: “Conclusions from cladistic methodology
– easily misled by close evolutionary convergence (Ridley 1986) – have sometimes been overturned by its more prestigious younger sister, DNA phylogenetics, which is less
vulnerable to closely convergent evolution. I vividly remember the shocked disbelief of many biologists when DNA phylogenetics revealed that the tenrecs and shrew-moles of
Madagascar were more closely related to elephants than to Eurasian shrews or hedgehogs (Stanhope et al. 1998).”

Do you see how DNA was given more ‘prestige’?
How morphology was ‘easily misled’? How it was ‘vulnerable’ to closely convergent evolution? The author sets you for the big unbelievable (and in the LRT, untenable, unverifiable, unreasonable) relationship of tenrecs to elephants. Do not believe such rubbish. If you wonder about such things, test them yourself, like I did. You’ll find that even with objective scoring, close convergence and all the other evils that befall phenomics (morphology), the unbiased software will carry to past all that to the land of verifiable, repeatable results.

Bird cladogram 2018 and the origin of penguins

Bird interrelationships
are coming together (Fig. 1). Here, wrens, dippers and woodpeckers now nest with crows, jays and grackles. Little else has changed. A few more taxa were also added.

Figure 1. Subset of the LRT focusing on birds. Here various aspects of birds are shown, including age, teeth, feeding behavior and basic clades.

Figure 1. Subset of the LRT focusing on birds. Here various aspects of birds are shown, including age, teeth, feeding behavior and basic clades. Four images change every 5 seconds.

The interesting thing I find in this cladogram
(Fig. 1) from the large reptile tree (LRT, 1146 taxa) is the highly derived placement of the penguin Aptenodytes. That’s really to be expected. However, a basal penguin, Waimanu (Slack et al. 2006; Fig. 2) nests from the Middle Paleocene (61.6 mya), soon after the K-T extinction event. That’s not really expected if one follows traditional cladograms that have birds radiating quickly and widely from the K-T boundary on out. Or maybe they did!

Figure 2. Waimanu is the penguin ancestor from the Middle Paleocene.

Figure 2. Waimanu is the penguin ancestor from the Middle Paleocene.

Wikipedia reports,
“Both DNA sequence analyses and anatomy argue for a close relationship between penguins and loons, with penguins being specialized for wing-propelled diving, and loons for foot-propelled diving.” Slack et al. 2006 nest penguins with storks, loons an petrels in order of increasing distance. I don’t see kingfishers in that list. And I wonder which stork was tested, because the Jabiru (see below) is traditionally considered a stork.

Figure 1. The extant murre, Uria, and the extant penguin, Aptenodytes to scale.

Figure 3. The extant murre, Uria, and the extant penguin, Aptenodytes to scale.

The LRT
(subset Fig. 1) nests penguins and murres with kingfishers (volant diving birds, Fig. 4) and the giant stork-like kingfisher, Jabiru, (which, of course, does not dive). These are close to loons + grebes (poorly flying diving birds) and terns (oceanic soaring diving birds, Fig. 5).

Figure 1. Megaceryle, the belted kingfisher may be a neotonous jabiru (genus: Jabiru).

Figure 4. Megaceryle, the belted kingfisher may be a neotonous jabiru (genus: Jabiru).

With Neornithes originating in the Early Cretaceous,
evolution had plenty of time to create penguins throughout the Cretaceous. But we don’t find their fossils then. Did Neornithes remain in small, out of the way enclaves, barely evolving, waiting, like mammals, to emerge after the Cretaceous?

Figure 3. Skeleton of Thalasseus, the crested tern.

Figure 5. Skeleton of Thalasseus, the crested tern. Scale bar = 10 cm.

Or did neornithine birds evolve throughout the Cretaceous
in areas that don’t often preserve fossils? Good question. 

Here’s a video worth seeing on Kingfishers.
Think of them as pre-penguins.

References
Slack KE. et al. 2006. Early Penguin Fossils, plus Mitochondrial Genomes, Calibrate Avian Evolution. Molecular Biology and Evolution, 23(6): 1144-1155.

online here.

http://www.otago.ac.nz/geology/research/paleontology/waimanu.html