In one timeline crown birds didn’t make it past the KT event, but ichthyornithids did. Like our birds they quickly radiated into a massive array of species; most of the initial niches were raptorial and insectivorous, but gradually aquatic plant eaters and arboreal frugivores kickstarted an herbivorous revolution, and these would eventually pale with the arrival of blunt-beaked herbivores, the toothed parrots.
The ancestor of the toothed parrots was a semi-aquatic, swamphen like ichthyornithid. It waded in the warm swamps of the Eocene, shifting from a duck or crane-like diet of soft aquatic plants into hardier reeds. To these ends it developed two key adaptations:
The upper jaw completely lost its teeth, becoming a powerful, deep beak. By contrast, the lower jaw essentially lost its rhamphoteca, and became lined with blunt teeth, similar to those of long gone sauropod dinosaurs.
The feet lost their webbing, and became prehensile, allowing the birds to manipulate food items.
The swamphen like forms quickly expanded into larger land herbivores similar to our gastornithids and dromornithids, and their reign wasn’t short either, having remained large browsers until their demise in the Pliocene when grass dominance was too much.
But their bigger success story was when these birds took to the trees, and quickly exploded into a variety of seed eaters and tough vegetation processors, dominating the volant herbivorous flying guilds in the arboreal realm.
Most remained anisodactyl, but some developed zygodactylous and some even heterodactylous feet multiple times. Their teeth remained peg-like in more folivorous and frugivorous niches, but for the more derived granivorous taxa they became increasingly bulkier and encased in bone much like those of the long gone dsungaripterids, making conventional reptilian tooth replacement impossible. In some taxa, some of the teeth moved sideways and lost their role in food processing, instead becoming powerful tusks.
One example of a derived toothed parrot are the elephant-parrots (genus Loxodontornis), birds endemic to African rainforests and dry forests. These gray birds have a hooked upper beak and large lower jaw tusks jutting to the sides, making them look like elephants from the front, helped further by their broad wings. More typical (for the group anyways) dsungaripteroid-like teeth process hard seeds and fruit as well as the occasional bone, while their heterodactylous feet both help manipulate food items as well as provide a stronger grip while climbing. Like all ichthyornithids they lack derived brain anatomy seen in our birds, so they are somewhat less agile flyers and in this particular species rather reluctant at flying.
A clade, the “p-parakeets”, have seen increased miniaturization, specializing on parakeet and finch-like niches. These birds are zygodactylous and have lost the lower jaw teeth, replacing them with a proper hard rhamphotecae; further, they have developed more complex birds, allowing for both increased aerial agility as well as higher intelligence. For now they are small, but these splitting images of our true parrots may yet rule the world.
To honor the new Minotaur Hotel build. Contains spoilers for both the ruthless and normal routes.
He sleeps soundly, arms wrapped around you.
You already don’t really feel the urge to sleep, as his peace and happiness warms your chest.
But you have something important to do, and so you calmly wait until he rolls around, as instincts demand weight to be distributed around different parts of the body as one sleeps.Even if it takes two hours.
You take a deep breath, and become your most ninja self. Limbs are a forest already opened, you just need to avoid the gusts of wind and the cracking of twigs.
You leave the bedroom barefoot. You walk the halls barefoot, and hum a most peculiar request: for light to refract and hide you. This will put a strain on the realm’s power, but the employees only need to get paid next week. You have a pile of diamonds stocked just in case, as recommended by Themba.
Like a sexy ghost you run out of the hotel evilly, reaching the passage to the valley. You are still barefoot, and the pebbles and limestone edges hurt, but you curse the land to feel your pain. It doesn’t actually solve anything, but the land suffers with you.
You carefully dig out at the side of the entrance. It also hurts, but moving the hands slowly over sharp rocks bleeds less than walking enthusiastically over it. Alas, you come across a grim relic: the flayed skin of your beloved, long worn by the Foreman.
Argos is no more, he left as the role was obsolete. You paid no mind to this because you didn’t know the truth about that snake, but an experimental prayer to Wepwawet sent you into a catatonic state where you had holy visions, finally unveiling what the Olympians had hid. You also saw Set with the head of a flamingo, as in the Book of Fayum, bowing in fear before Horus in the form of a crocodile. That was very funny.
You learned of the pelt, and while not spoken outright you put the two and two together when you read the votes of the Olympians. You had immediately requested privacy, crying tears of joy. Asterion was worried when he found you with dried tears but you distracted him away from that, and enjoyed a peaceful day doing pretty much anything that made him happy. Suspicion remained within him, but he had reluctantly accepted your claims of allergies and fungal infections.
You know lying is not healthy for a relationship, but this was a gift too precious for him to learn without surprise.
The path treads on, the land submissive to its master. Occasionally horrific things gnaw and dare come closer, but you are living the power fantasy with well aimed blades cutting even the most harmless of creatures and shredding them into blood butterflies. The jouney is long and your feet are essentially caked in dried blood and screaming at you for not wearing shoes, but it’s fine.
Eventually, you come across a cave with a wine-dark lake. Before it is a statue of Poseidon, which you would normally ignore if not for the most welcoming of circumstances. You utter a silent thanks, not a prayer but a link of gratitude to the figure before you. You don’t much see fit to utter a hymn to the Hellenic gods, but you assume the correct posture for a sacrifice.
The waters are ink black, but they’re not salty, so you spray them with bath salts. Given the lands restriction on spices and other currencies you were not sure if proper sea salt was something you could summon into existence, so in case you run a conquest in which anyone capable of supplying you with roughly a ton of salt would be granted an elephant-sized sum of platinum (not considered a currency until very recently so not restricted by the Olympians). Asterion assumed you’d be using this salt to improve the wave pool, and you did. But you also stored an amount comparable to your body weight etched within your clothes, then an additional amount in pockets which you attached to your teeth as you left the hotel. You considered also surgically insert an additional kilogram into your thoracic cavity, but it was already filled with love.
You tear your clothes and the pockets and unload them into the water. If not accepted as a sea, then you’d just force the fact that Poseidon started off as a chthonic god associated with rivers anyways. There is no way out (like that Phil Colins song in the Brother Bear movie), you will succeed (unlike anything written by Phil Colins).
Finally, you lay the pelt into the water. It thrashes horribly as if it was alive, and your heart darkens with images of Asterion being flayed. You grimace and wait for the thing to be over with, for the skin to sink into the dark abyss.
Once it is, your reward is clear: something glistens in the water. You remember a book explaining the origins for the whole light thing in the Bible, that to the people of the Ancient Middle East light was used as a metaphor for a god’s presence and darkness for the god’s lack of favor. Well, Poseidon’s will is certainly manifest, and you retrieve the light from the dark waters, like the sun rising from the sea.
You use the remnants of your clothes to shield it, then walk back. Pain nor awareness of nudity can stop you, you run madly with joy cross the desert, until a greater light rises in the horizon.
And, when you reach the passage, a third white form awaits for you. He momentarily sighs in relief when he notices you waving at him, then goes back into panic at seeing your state.
“Have you lost your mind!?” he panics, “[insert your name] you’re bleeding all over! Why would you do this!?”
He edges a bit into the valley side of the entrance, but you don’t need to wait long until he wraps your arms around you and drags you to the safety of the hotel.
“Babe, I got you gift!” you say ecstatically.Asterion shakes his head. You almost visualize him saying something to the effect of “No gift is worth you getting hurt” or something more emotionally powerful, so you waste no time in unwrapping the light and making it touch his hand.
And then he knows.
Words fail to escape him, when he’s in shock, when he hiccups, when tears well in his eyes, when his hand touches his muzzle.
Asterion sniffs, and as he begins to bawl he hugs you dearly, nuzzling you deeply as he weeps. Tears worth thousands of years of torment are let out, and you both sit down. You consider creating some sort of barrier to shield the two of you, but your attempts are met with a violent force. Either you completely abused your power with the light refraction thing or the contracts are no longer operational now that the prisoner is emancipated.
“T-thank you” Asterion says, wiping his eyes, beaming love and joy that make all stars look bleak and lifeless in comparison, “Thank you so much!”
You want to say an righteous “it was the right thing”, a flirty “it’s the least I could do”, perhaps even a sheepsih “you’re welcome”. But silence is the apropriate response. No words can do justice to the weight of this moment. Asterion was already free the moment you came into his life, but the end of so much torment doesn’t come lightly.
So you just remain entwined with each other, two souls basking in a sunrise. Helios was Eleutherios, the liberator, so while entirely accidental on your part you couldn’t help but feel that this dawn was beyond appropriate.
After Asterion’s hiccups die down a little, you dare:
Asterion kisses your forehead, taking in your sweaty, salty scent.
“My mission remains. I don’t know if the contracts uphold, but Hestia’s fire remains as a gift. Now I’m free to go wherever we want, but nothing will really change. Other than the fact that you’ve done yet another miracle, and that I won’t stop loving you for all eternity.”
“Good” you say, the only word out of many in your mind that come out.
To honor the latest release of Minotaur Hotel I decided to do an article on what is known of the Minoan deities.
Known as the “first European city-makers” and a distant precursor to Greece, what is called the Minoan Civilization after King Minos of Crete was a mysterious Bronze Age nation that governed Crete and neighbouring parts of the Aegean. Its age, likely influences over posterior Greek (and by extension western) culture and unique art has long made it a subject of mystique and intrigue. Whereas it’s the severalstillundeciphered scripts and languages or the fact that it seems to have a rare genuinely matriarchal society, it seems the countless research and academia only raises more questions than answers.
One such well documented but ultimately unsuccessful endeavour is identifying the pantheon these people worshipped. It is strongly speculated that Minoan Crete was theocratic (Kristiansen & Larsson, 2005, among several others) and several art either represents cultic activities (such as the famous bull leaping) if not gods themselves, but in the absence of the proper written word this is beyond impossible to ascertain. The implications of understanding Minoan religion are very clear, as beyond offering a snapshot to the lives of these people it also bears the potential implication that many Greek gods and mythological figures ultimately had their origins here.
To completely compile, summarise and synthesize all that has been written on Minoan religion is a task far too vast to implement, so here are some of the most widely agreed upon gods.
Queen of the Gods
By far the most well kown figure attributed to Minoan religion is the Queen or Mother goddess, sometimes known as “Snake Goddess” due to an abundance of figurines depicting women holding snakes. Perhaps surprisingly (or not given the second paragraph), she’s not actually attested anywhere, given that Minoan scripts haven’t yet been deciphered, but her existence can be inferred due to a variety of factors:
Female figurines are by far the most common representation of what could be interpreted as a god in Minoan sites. Chief among these are the aforementioned “snake goddess” figurines. While there is considerable debate on whereas these truly represent deities (plural or singular) or mortal priestesses, they are comparable to apotropaic depictions of surrounding cultures, most notably those of the latter Athena Parthenos which similarly is associated with serpentine iconography as controlling these forces of chaos (Ogden 2013).
The fact that Minoan society was matriarchal in nature, which would lend credence to the supreme being in their cosmology being feminine in nature. While a dominant female deity does not always correlate to a matriarchal society (i.e. Amaterasu, Virgin Mary, et cetera), the opposite, a matriarchal society with a masculine supreme god, is yet to be documented (though see below).
Several Greek mother goddesses such as Demeter and Rheia are thought to have a Cretan origin (Mylonas 1966, Sidwell 1981 among several others), so it’s not terribly hard to see them as “descendents” of this Minoan deity.
The Philistines, contray to biblical assertions on Dagon worship, seem to have favoured a goddess as their primary deity (Schäfer-Lichtenberger 2000, Ben-Shlomo 2019). The Philistines, through genetic legacy and material culture, are now understood to have had an Aegean origin, so again seeing this as a continuation of a Minoan goddess is plausible.
Several names have been speculated for this deity, usually along the lines of the author’s interpretation of Minoan scripts (which should be noted, are not only undeciphered but very likely don’t mention deities at all, since all we have seem to brief texts likely attributed to tax reports). The name “Rhea” doesn’t seem to be of Indo-European origin (Nilsson 1950, Sidwell 1981), making it very likely that this is a theonym with Minoan origins. The same applies to Ariadne (Alexiou 1969) and possibly also Athena (Beekes 2009). Conversely, the Philistine goddess is possibly attested as “Ptgyh” (Ben-Shlomo 2019), a name that is speculated to be related to Greek “Potnia”, “mistress”. In all likelihood, such an important goddess likely was known by a variety of epithets.
Fertility is naturally considered a major function of this mother goddess, but perhaps in ways one might not expect. An emphasis on solar worship has been noted due to temple arrangements and material objects such as “frying pans” with solar iconography (Ridderstad 2009), suggesting that, rather than an earth goddess as one might expect, this was a solar goddess. Solar goddesses are known from a variety of Near Eastern cultures such as Egypt (Sekhmet, Hathor), Anatolia (Arinniti, Istanu, Estan, Wurunsemu) and Canaan (Shapash) so a solar interpretation of the Minoan supreme goddess isn’t unusual. In particular, this might imply a more “chthonic” interpretation of the sun than the Classical “object in the sky”, due to temple angles tressing sunrises and sunsets (Ridderstad 2009), whch is consistent with the Hittite notions of the sun goddess ruling the underworld. Regardless, as noted below in Talos there is also possible evidence for a Minoan male sun.
More unambiguously, this goddess had a civil and possibly domestic function. As noted above snake goddess figurines might be apotropaic in nature, used to ward off evil spirits or more mundane threats like snakes. If Athena is derived from this goddess then a role as the protector of the palace is also implied given Athena’s role in the Mycenaean era, and both Ariadne and Athena are associated with weaving. Conversely, so are solar goddesses in other places, like the Baltic Saule or the Turkic Gun Ana, as the rays of the sun are easily linked to threads, further suggesting this role for the Minoan goddess. Both Rhea and Demeter are also associated with lions, animals that not only are symbolic of the sun but also of a notable sun goddess across the sea, Sekhmet.
The fact that the Minoan ruling goddesses was the possible genesis for several Greek goddesses like Rheia, Demeter, Ariadne and Athena suggests a rather extensive and important function in ancient Cretan religion. Conversely, it might also suggest that what we might attributing to a single goddess was in fact several different deities, but as deities overlap and flow into one another it is possible that these goddesses were either seen as one or acquired independent identities several times throught Minoan history.
The Bull God
The bull is extensively depicted in Minoan art. Most common are bull-leaping frescos depicting youths of both genders leaping or interacting with bulls, suggesting this was a common Minoan sport and perhaps even a religious ritual. But bulls are depicted in many other contexts as well, and as such the existence of an actual Minoan bull god is frequently speculated upon.
In Near-Eastern cultures, bulls are both solar and lunar symbols. On the one hand, the bull’s horn/s resemble/s a lunar crescent, and indeed not only are Middle Eastern male moon gods like Nanna and Suen associated with the bull but even the Greek Selene is described as having a chariot pulled by bulls, suggesting that not even a shift towards a feminine moon deity erased this iconography. On the other hand, a bull is a powerful animal and thus worthy of male solar gods, most notably the Mesopotamian Marduk (literally “calf of the sun”). Sometimes both interpretations show up in the same culture: in Egypt the Apis bull is associated both with Ra and with Osiris as Yah (the moon). Perhaps the same applied to ancient Crete (again, see Talos below), but a lunar bull would certainly be a vivid symbol contrasted against the sun goddess.
The bull is associated with Dionysus which otherwise is mired in more “exotic” symbols, suggesting that the putative “Minoan Dionysus” might be the bull god. It has long been speculated that the bull god is a male youth and son and consort to the queen of the gods, though women are also depicted bull leaping.
The Greek minotaur has long been speculated to be a remnant of the Minoan bull god, not without reason being so throughly linked with Crete as a concept. In this case, the monstrous depiction is either fully discontinuous from older practises or defamatory, with my personal two cents that it is also a jab against the bull gods of the Phoenicians, accused at the time of human sacrifice by the (infant killing) Greeks. Asterion is said to be the birth name of the minotaur by Pseudo-Apollodorus, but I wouldn’t read much into this since this name (literally “starry one”) is a common Greek name for many figures both historical and mythological, and at any rate a recent Indo-European name at odds with the most likely Pre-Greek Cretan languages.
There is extensive evidence of wine cults in Minoan Crete (Kerényi 1976). This, combined with the Mycenaean depictions of a bull-horned Dionysus (or “di-wo-nu-so” as it is) seems to point to a Minoan origin for this god. “Dionysus” is an Indo-European name connected to Zeus and other sky father figures but the actual character of the god is not easily identified in the PIE world, suggesting a Pre-Greek, local origin. A possible exception is the Lusitanian god Andaeico (Teixeira 2014) which might resemble the putative “flower Dionysus” (see below), but this deity is himself not well understood and might be from an ancient Iberian stratum in Lusitanian culture.
The Mycenaean Dionysus is a figure with stronger ties to death and rebirth than revelry necessarily but the evolution from “eldritch god” to “party dude” might not have been as linear (geddit) a concept as one might expect. Male figurines thought to represent a young god increase in popularity in later stages of Minoan history (Vasilakis 2001) as do male youth figures often identified as “prince of the lilies/flowers” which alongside the wine cults is closer to the Classical Dionysus than the Mycenaean or later Orphic one. However once more in the absence of deciphered scripts it is impossible to say for certainty that these figurines represent deities let alone are Dionysus. Hell, the “flowery figures” have even been interpreted as female at times.
If an actual god, the “Minoan Dionysus” might very well be identified with the bull god, as the bull is a rather odd symbol for the “exotic” attributes the Classical Dionysus is associated with. Ariadne in Greek myth does get hitched with Dionysus; an imbalanced, reversed remnant of the male youth/Minoan queen goddess pairing perhaps?
Perhaps the only Minoan or at least Cretan god we may truly known by name is Talos. In Greek myth Talos is best known as the strange automaton made by Hephaestus, but it was also the Cretan word for “sun”, analogous to “Helios” of mainland Greece according to Hesychius of Alexandria. Zeus was worshipped in Crete as Zeus Talaios, who was associated with the sun, and the Tallaia was a spur of Mt. Ida associated with sunrise rituals (Nilson 1923).
This association of Zeus with Talos is as peculiar as it is extensive. Zeus, a god whose origins are well documented to be Indo-European in nature, is held in Greek myth as born and raised in Crete, and Cretan depictions of Talos differ from those of mainland Greece in having wings. Further, the seduction of Europa by Zeus as a bull links the Classical Zeus to Crete in a very fundamental way. This seems to indicate a rather through syncretism between the Greek/Mycenaean sky god and this indigenous Cretan deity, which in turn implies a rather relevant role to the Minoan Talos.
Conversely, outside of Crete Talos is an enigmatic figure, as noted by Pausanias himself which seems more confused than anything. Certainly, the story of a pre-sci-fi robot is weird, let alone how it relates to an ancient Cretan god, linked to the supreme god of all Greeks down to his very birth.
Talos is truly an anomaly. A solar god which was important enough to warrant syncretism with Zeus, in a matriarchal culture where the sun seems to have been traditionally the supreme goddess herself. Crete was likely never a monolith even at the height of Minoan rule, but all current signs point to Talos being an ancient Cretan deity from before PIE influences in Greece, and he seems so out of place.
My personal two cents is that Minoan cosmology was similar to that of the Hittites and other Anatolian cultures, where the sun is male during the day as it travels through the sky and female at night where it rules the underworld. Talos’ syncretism with Zeus therefore would be derived from representing the male, skyward aspect of the sun, corroborated by worship at the Tallaia. In the original Minoan religion Talos was probably lesser compared to his female aspect (which even as a chthonic deity would easily be accepted as the supreme power; even Mycenaeans favoured the chthonic Poseidon to the celestial Zeus after all), but his roled ensured syncretism with the king of the gods once Crete was conquered.
Britomartis is possibly another deity we might know from a genuinely Minoan or at least Cretan name. Solinus claims it is “sweet virgin” in Cretan and the name doesn’t seem to have Indo-European roots. If true, I’d imagine this theonym is more due to syncretism with the Greek Artemis if anything as I doubt ancient Minoans cared much about virginity as a concept, though Artemis herself may be derived from this deity. Some archaeologists have further suggested that it is an euphemism for the deity’s actual name, since being a goddess of the wilds saying it might have been unwise (Ruck 1994). Another name attributed to her is Diktynna, “hunting nets”, or simply Dicte/Dikte (unsurprisingly, she named said mountain, and was likely its spirit). I’ve never seen the etymology of this name tracked, so I can’t say for sure if it is Greek or Pre-Greek in origin
Britomartis is in Greek myth a mere oread or mountain nymph, said to have invented hunting nets. She is said to have fled Minos’ lust, a tale that even Siculus expressed disbelief at due to her divinity. Thus, although greatly diminuished by Hellenistic times, she was still clearly held to be a deity, and still seems to have been worshipped in Crete during Classical times, frequently appearing in coinage as a winged figured. She is equated to Artemis, a goddess associated with the wilderness and mountains, and it can be assumed she represents a similar “lady of the beasts” archetype. Artemis herself has a name of unclear etymology, and could be of Minoan origin, being perhaps another name for Britomartis.
Some authors tempt to lump Britomartis with the Minoan mother goddess, but to me these seem like clearly distinct figures. Whereas the queen of the gods is a civic, fertility and possibly solar figure, Britomartis is alcearly a goddess of the wild places, perhaps even more specifically the embodiment of Mt. Dicte. Of course, overlap between these two goddesses likely happened at several points in Cretan history.
And that’s it for now.
Other Minoan gods have been positted, including a sea one (naturally), but they aren’t sufficiently supported by everyone in the field at large, so I won’t bother.
Kristiansen, Kristian & Thomas B. Larsson. The Rise of Bronze Age Society: Travels, Transmissions and Transformations. Cambridge: Cambridge University Press, 2005.
Ogden, Daniel (2013). Drakon: Dragon Myth and Serpent Cult in the Greek and Roman Worlds. Oxford University Press. pp. 7–9. ISBN 9780199557325 – via Google Books.
George Mylonas (1966), “Mycenae and the Mycenean world “
Sidwell, R.T. (1981). “Rhea was abroad: Pre-Hellenic Greek myths for post-Hellenic children”. Children’s Literature in Education. 12 (4): 171–176. doi:10.1007/BF01142761. S2CID 161230196.
Christa Schäfer-Lichtenberger, The Goddess of Ekron and the Religious-Cultural Background of the Philistines, Vol. 50, No. 1/2 (2000)
David Ben-Shlomo, Philistine Cult and Religion According to Archaeological Evidence, January 2019Religions 10(2):74, DOI: 10.3390/rel10020074
Nilsson, Martin Persson (1 January 1950). The Minoan-Mycenaean Religion and its Survival in Greek Religion. Biblo & Tannen Publishers. ISBN 9780819602732 – via Google Books.
Beekes, Robert S. P. (2009), Etymological Dictionary of Greek, Leiden and Boston: Brill
Marianna Ridderstad, Evidence of Minoan astronomy and calendrical practices, October 2009
Kerényi, Karl. 1976. Dionysus. Trans. Ralph Manheim, Princeton University Press. ISBN 0691029156, 978-0691029153
Monteiro Teixeira, Sílvia. 2014. Cultos e cultuantes no Sul do território actualmente português em época romana (sécs. I a. C. – III d. C.). Masters’ dissertation on Archaeology.. Lisboa: Faculdade de Letras da Universidade de Lisboa.
Andonis Vasilakis, MINOAN CRETE: FROM MYTH TO HISTORY Paperback – January 1, 2001
Nilsson, “Fire-Festivals in Ancient Greece” The Journal of Hellenic Studies 43.2 1923
Carl A.P. Ruck and Danny Staples, The World of Classical Myth [Carolina Academic Press], 1994
A common trend in pterosaur paleoart in the mid-2000’s was depicting them with two sets of wings. In these depictions the legs, usually free from the brachiopatagium, would be elevated in flight and, thanks to their own uropatagia running alongside them, form a secondary set of wings. Essentially, a membranous version of the tail feathers of birds like swallows or kites.
This trend seems to have been popularised by John Conway and quickly took hold, before disappearing roughly in the early 2010’s. Nowadays, it is rare to see this type of depiction, at least among serious artists.
The appeal of the pterosaur hindwing is pretty basic. At the time, it was increasingly common to try to reinvent pterosaurs away from more bat-like depictions of decades prior and to compare them functionally to birds, including giving them a more “dynamic” look. It was thought that pterosaurs rose their legs in flight like modern gliding lizards and the extent of the brachiopatagium at the time was less publicized (though known since Pterodactylus specimens were first described). Add in the discovery of four winged dinosaurs like Microraptor, and the idea that pterosaurs had a similar bauplan just sort of clicked, you know.
So what happened?
Seeing as this trend died out around the publication of Mark Witton’s book on pterosaurs, more widespread information on pterosaur anatomy can be considered the killer of the pterosaur hindwings. We’ve already known for a while that the hindlegs of pterosaurs were connected to the brachiopatagium and that the cr/uropatagia of pterodactyloids was vestigial at best while that of their earlier cousins formed a blob-like shape not particularly resembling wings, but a few factoids have since shut down this artistic concept:
The idea that pterosaurs needed tails is not well funded. Modern birds can fly without tails and many extinct bird clades like Enantiornithes and lithornithids lacked tail fans and were still efficient flyers. Likewise, many modern bats like flying foxes are in the same boat as pterodactyloid pterosaurs, with vestigial uropatagia.
The need to functionally compare pterosaurs to birds has reached an all time low. Most notable is the notion that pterosaurs used their forelimbs to launch; this would make the seperation of the legs from the brachiopatagium less urgent.
It is not entirely clear if pterosaurs could raise their hindlimbs like gliding lizards do. A 2018 study suggests pterosaurs had such erect hindlimbs that they were just as incapable as birds of raising their hindlegs, but this study has since been criticised for ignoring soft tissues. It seems most likely pterosaurs could still raise their hindlegs, but this probably only helped reduce drag, rather than form a secondary wing set.
The pterosaur hindwing is an interesting hypothesis, and while it doesn’t seem to have survived the test of time it was an interesting flame while it lasted.
By the Early Cretaceous, both birds and pterosaurs had achieved a cosmopolitan distribution, thanks to their abilty to fly. A lot has been written on the diversity of both groups up to the end of the Mesozoic, but it is widely agreed that even in the Maastrichtian pterosaurs were found in all continents including Oceania (Witton 2008) and Antarctica (Kellner 2019). And the same applied for birds, with both northern hemisphere formations like Hell Creek and southern hemisphere ones like the Lecho Formation offering insights to the diverse avifauna of this epoch.
In spite of this, however, there are two places in the world in which either group is completely and conspicuously absent. Both bird and pterosaur fossils are highly susceptible to preservation bias and rare in non-Lagerstätte sites (Silverstone 2016 among several others), but these particular sites are noted for their high degree of preservation and presence of microfauna from small amphibians to insects.
The first is the Kem Kem Group, from the Cenomanian of Morocco. This ecosystem, representing several estuarine stages from more inland freshwater habitats to coastoal waters, preserves a very rich vertebrate and invertebrate fauna, with dinosaurs like Spinosaurus being the best known residents. Among the finds is a very rich pterosaurian fauna dominated by azhdarchoids and pteranodontians: this included the tapejarid Afrotapejara, an undescribed azhdarchid, the chaoyangopterid Apatorhamphus, the incertae sedis azhdarchoids Leptostomia, Xericeps and Alanqa (the latter two have variously been interpreted as either azhdarchids or thalassodromines; see Longrich 2018 and Campos 2021 for instance) and severl ornithocheirids/anhanguerids such as Anhanguera, Coloborhynchus, Siroccopteryx and Nicorhynchus. Yet there is not a single avian fossil in sight, in spite of the fact that almost all contemporary formations have rich avifaunas, including contemporary African formations such as the Zebbag Formation (Contessi 2012). While an attempt in the paper is used to explain the absence of mammals (namely, competition with notosuchians), it only notes the strangeness of the lack of avian fossils. In general pterosaurs seem to occupy a very wide range of trophic niches in the Kem Kem Group, from the small probing Leptostomia to the frugivorous Afrotapejara to the durophagous Alanqa to the piscivorous ornithocheirids/anhanguerids to the raptorial Apatorhamphus and Xericeps, so one might be tempted to explain the absence of birds as particularly aggressive competitive exclusion on the part of pterosaurs. Still, this pterosaurian diversity is not too far off from those seen in other formations with birds, so the Kem Kem environment must have truly been strange to favor pterosaurs alone.
The other are the various Maastrichtian formations of India and Madagascar, which then were either still together as an island or just recently seperated. A diverse avifauna is known from the Maevarano Formation that includes Enantiornithes (both pengornithids such as Falcatakely as well as Vorona, now interpreted as an enantiornithean rather than as an ornithurine as previously believed; see Hartman 2019 and Pei 2020), flying dromaeosaurs such as Rahonavis and even possible relatives of Sapeornis (O’Connor 2010). Indian formations meanwhile preserve neither bird nor pterosaur fossils, but they don’t approach the sheer level of preservation detail as the Maevarano, which also includes postcranial remains of allotherian mammals, frogs and other fragile specimens. The absence of pterosaurs from Indo-Madagascar is just as puzzling as the absence of birds from the Kem Kem Group; pterosaurs in the Maastrichtian essentially surrounded this landmass, with South America, Antarctica, Australia and New Zealand each having specimens and Africa in particular preserving a rich pterosaurian fauna in the Ouled Abdoun Basin (Longrich 2018). Furthermore, unlike the Kem Kem Group there doesn’t seem to be anything close to a competitive displacement: none of the Maevarano birds are larger than average for Mesozoic birds, and with the exception of possibly Vorona and Rahonavis nearly all seem to occupy primarily arboreal niches. True, the toucan billed Falcatakely, the flying dromaeosaur Rahonavis and omnivoropterygids are all unusual elements for a Maastrichtian avifauna, but none seem to push the boundaries of the Late Cretaceous avian morphospace.
Ultimately, both cases are mysteries, but perhaps this is indicative that even cosmopolitan groups can have sufficiently large gaps in their range.
Mark Witton and Darren Naish, A Reappraisal of Azhdarchid Pterosaur Functional Morphology and Paleoecology, PLoS One. 2008; 3(5): e2271. Published online 2008 May 28. doi: 10.1371/journal.pone.0002271
Alexander Kellner, Taissa Rodrigues, Fabiana R. Costa, Luiz C. Weinschütz, Rodrigo G. Figueiredo, Geovana A. de Souza, Arthur S. Brum, Lúcia H.S. Eleutério, Carsten W. Mueller, Juliana M. Sayão, Pterodactyloid pterosaur bones from Cretaceous deposits of the Antarctic Peninsula, Earth Sciences • An. Acad. Bras. Ciênc. 91 (suppl 2) • 2019 • https://doi.org/10.1590/0001-3765201920191300
O’Connor and Forster, 2010. A Late Cretaceous (Maastrichtian) avifauna from the Maevarano Formation, Madagascar. Journal of Vertebrate Paleontology. 30(4), 1178-1201.
Nizar Ibrahim, Paul C. Sereno, David J. Varricchio, David M. Martill, Didier B. Dutheil, David M. Unwin, Lahssen Baidder, Hans C. E. Larsson, Samir Zouhri, Abdelhadi Kaoukaya, Geology and paleontology of the Upper Cretaceous Kem Kem Group of eastern Morocco, https://doi.org/10.3897/zookeys.928.47517 (21 April 2020)
Martin-Silverstone, Elizabeth; Witton, Mark P.; Arbour, Victoria M.; Currie, Philip J. (2016). “A small azhdarchoid pterosaur from the latest Cretaceous, the age of flying giants”. Royal Society Open Science. 3 (8): 160333. Bibcode:2016RSOS….360333M. doi:10.1098/rsos.160333. PMC 5108964. PMID 27853614.
Longrich, Nicholas R; Martill, David M; Andres, Brian (2018). “Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary”. PLOS Biology. 16 (3): e2001663. doi:10.1371/journal.pbio.2001663. PMC 5849296. PMID 29534059.
Campos, H. B. N. (July 31, 2021). “A new azhdarchoid pterosaur from the Late Cretaceous Javelina Formation of Texas”. Biologia. doi:10.1007/s11756-021-00841-7.
Hartman, Scott; Mortimer, Mickey; Wahl, William R.; Lomax, Dean R.; Lippincott, Jessica; Lovelace, David M. (10 July 2019). “A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight”. PeerJ. 7. doi:10.7717/peerj.7247. PMC 6626525. PMID 31333906.
Pei, Rui; Pittman, Michael; Goloboff, Pablo A.; Dececchi, T. Alexander; Habib, Michael B.; Kaye, Thomas G.; Larsson, Hans C.E.; Norell, Mark A.; Brusatte, Stephen L.; Xu, Xing (August 2020). “Potential for Powered Flight Neared by Most Close Avialan Relatives, but Few Crossed Its Thresholds”. Current Biology. doi:10.1016/j.cub.2020.06.105.
(Once again thanks to Ron Blakey, NAU Geology, for the free use in desecrating the maps)
A basic map of ratite “urheimats” during the earliest Cenozoic: red = ostriches and kin, indigo = “lithornithids”, green = rheas, yellow = casuariforms, dark red/crimson = moas and tinamous and brown = kiwis and elephant birds.
A Laurasian origin for Paleognathae has been proposed in several recent studies, in contrast to the old idea that ratites are gondwannan vicariants. The two most basal clades, “lithornithids” and ostriches, certainly are dominant during the Paleogene of Laurasia and predate gondwannan ratite fossils. Lithornithids are thought to have had a circumpolar distribution and are found in European and North American sites as recently as the mid-Eocene. Ostriches are also found in all of Laurasia but have diversified into several clades as they lost the ability to fly early on: Geranoididae in North America, Palaeotiidae in Europe and Eogruidae in Asia. Of these, it seems eogruids might be the most likely ancestors of modern ostriches, which would later invade Africa and India in the Miocene.
Rheas have a poor fossil reccord, but putative members occur in the Paleocene and Eocene of South America and Antarctica. They don’t seem to have expanded beyond these landmasses.
Tinamous are known from the fossil reccord prior to the Pliocene, but moas have been in New Zealand most likely since the Paleocene, as the known Miocene forms are already flightless and specialised. They might have crossed Antarctica or flat out flown across the Pacific.
Casuariforms are best known from Australia but the earliest representative, Diogenornis, comes from South America. As Diogenornis still has relatively large wings its unclear if it and emus/cassowaries lost the ability to fly independently or if they crossed Antarctica into Australia (or vice versa).
Kiwis and elephant birds have the most drastic geographic distance. Given hw specialised elephant birds are I’m assuming they might have been on Madagascar since the Paleocene, so the still volant kiwi ancestors might have crossed the Indian Ocean with India as a stepping island, then Australia before arriving to New Zealand in the Miocene. Or they could have both come from either Asia and migrated southwards or Antarctica and inversely migrated northwards. The absence of Australian fossils is of particular interest.
This leaves Africa as the only continent without indigenous ratites prior to the Miocene. Eremopezus and “aepyornithid-like” forms might be the original African ratites, but for now their relations to the rest of Aves remain unknown.
In an alternate timeline two lineages of placoderms convergent with syngnathid fish evolved:
– Longmaiformes were a clade of arthrodires of uncertain affinities as they more or less appear in the mid-Devonian fossil reccord as they are, with the general consensus leaning towards being part of Coccosteina. They developed long, tubular jaws similar to those of the Cretaceous turtle Ocepechelon, the upper jaw dental plates flanking it to keep the suction tube’s shape and the lower jaw’s dental plate reduced to a small hook keep the jaw closed, with only a small jaw motion required for a gap to be formed and prey sucked in. Like seahorses and trumpetfish a bent neck was developed, allowing a greater degree of motion and thus more efficient prey capture. In derived taxa the pectoral fins became the dominant propulsion mechanism while the tail lost its fin and became prehensile. Like in most placoderms they were viviparous, the females giving birth to a few large pups.
– Jurakaniformes were a clade related to Entelognathus and Qilinyu, and as just converged more deeply with syngnathids, their small jaws at the end of a long tubular snout much their actinopterygian mimics. Derived taxa took a more serpentine apparence, relying on their dorsal and anal fins to swim, and like both Longmaiformes and syngnathids they developed a bent neck. Little is known about their reproductive habits, but the presence of claspers suggests internal fertilization at least.
Both clades survived the Devonian mass extinction, presumably due to more generalistic habits as well as the spread of coastoal swamps and other well vegetated aquatic biomes that favoured their unique lifestyle. Longmaiformes in particular were well successful in marine habitats, some reaching large sizes and becoming some of the largest marine vertebrates of the Paleozoic. Jurakaniformes, however, were the longer survivors: longmaiform diversity plummeted in the early Permian and eventually the group became extinct in the Capitanian Extinction event, while seahorse and pipefish like jurakaniforms made it into the earliest Triassic, before dying out, presumably due to predation from marine temnospondyls and reptiles.
Placodonts were some of the most resilient groups of Triassic marine reptiles, consistently surviving minor extinction events with minimal diversity loss until the mass extinction.
In one timeline, this did not happen. Instead, their diversity would continue, in some ways replacing the turtles from our timeline.
The Mesozoic reccord of placodonts is relatively stable. Their diversity remains mostly consistent in spite of several other extinction events affecting other marine reptiles (such as thalattosaurs, phytosaurs and pleurosaurs), there being two major spikes of diversity: in the early Jurassic and late Cretaceous.
Soon after the TJ extinction event, Placochelyidae increases significantly in species diversity. They dominate marine habitats from the near shore to the open sea, a few species even occuring in brackish and freshwater habitats like in the Phu Kradung Formation of Thayland. Mesozoic taxa have developed flippers and exploited open sea habitats long before our turtles, the closest analogues in our timeline being some thalassochelydids in the Late Jurassic. Some taxa became cephalopod and jellyfish specialists and lost their teeth, developing long toothless jaws to snatch their prey. Most however remained in the coastoal regions eating hard shelled invertebrates, albeit most genera having a cosmopolitan distribution.
Placochelyids undergo the Jurassic/Cretaceous boundary virtually unaffected unlike other marine reptile groups, but begin to lose steam in the Cenomanian turnovers. They remain relatively important components of marine faunal assemblages, but they are upstaged by other clade of placodonts, the cyamodontids.
Cyamodontids spent most of the Mesozoic up to that point as relatively unspecialised coastoal and freshwater durophagists, like weird reptilian skates. Everything changed in the post-Cenomanian Cretaceous, where a decline in placochelyid diversity allowed them to explode ecologically and become the dominant placodont linage. Most similarly remained coastoal/freshwater taxa, but several took to life in the open seas, developing flippers and either reducing their carapace like leatherback sea turtles (as well as a few placochelyids) or by inversely expanding it into broad lift-generators in order to remain suspended effortlessly, relying on small but powerful flippers for propulsion and steering. Like in placochelyids some became toothless, but instead developed suction feeding like our turtle Ocepechelon.
While cyamodontids became the dominant placodonts, placochelyids still retain a relatively high degree of diversity, and both taxa passed through the KT extinction event with few casualties, aside from the more pelagic taxa. They continued to dominant early Cenozoic oceans until the Eocene, when global cooling events caused their extinction.
This freed the way for the third and last lineage of placodonts, the henodontids. Through most of the Mesozoic these reptiles remained in freshwater and brackish water biomes, but with the onset of the Cenozoic they began to explore marine niches, just before the extinction of their relatives. Surviving the climatic turmoils much better (barring a massive species loss in the Pliocene/Pleistocene cooling), these remaining placodonts are truly the turtles of this world, having lost their teeth in favour of a strong beak. They occur in freshwater and marine habitats, using their broad carapaces to generate lift while the flippers do the work. They feed on all manner of items from crustaceans to algae to fallen fruit to vertebrates they suction feed. They haven’t managed to return to land, the niche of tortoises instead occupied by notosuchians and sphenodonts, but at over 300 species in terms of diversity do they really need to?