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Why Swans are bizarre and tragic

January 31, 2012

Cygnus buccinator, the largest species of waterfowl (after dromornithids, several hawaiian waterfowl and Cnemiornis became foie gras).

Appearently, through dubious research, the animal that most represents me is the swan, both as a totem and as a daemon (read His Dark Materials, scum). I must say I’ve always felt drawn towards large, white waterbirds, but it is still something of a shock.

Regardless, swans, I think, are very interesting for mutiple reasons. They seem like nature’s experiment at creating something that should not exist: from an ancient small, bipedal theropod came a being that is fautlessly angelic, with a serpentiforme neck, a soft, platypus like beak (Anseriformes are notable amidst modern birds for barely having a true rhamphoteca, restricted to the very tip of their jaws as that iconic “tooth”; the rest of the beak is covered by a very thin and soft keratin covering, almost like colourful leather), and angelic wings capable of breaking human bones. They are what I like to call an “angelic abomination”*; by themselves, they’re beautiful, but their anatomy seems like an abominable distortion when compared to even other birds.

*Yes, I do realise how redundant that is. If you don’t know why, then you don’t know how judeo-christian angels look like (hint: they’re not pretty people).

On top of that, they’re specialised aquatic herbivores living on inland waterways on cold climates. Surprised, huh?

How the “ugly” duck[ling]s ascended

Coscoroba coscoroba. While it's relationships with true swans are not well figured out, it most defenitely resembles the prototype.

Swans are almost consistently classified as within Anserinae, the clade that also includes most anatids called geese. I’ve seen arguments that anserines are polyphyletic and thus not a true clade, but for the most part they still tend to be classified as a single clade. True swans almost certainly diverged from other geese early on, during the early/middle Miocene; the modern south american Coscoroba Swan (Coscoroba coscoroba) is sometimes considered the closest living relative of Cygnus, thus rendering it a “true swan”, while some studies place it at the base of Anserinae while Cygnus diverged from the main lineage imediately after. Regardless, Coscoroba is perhaps a living image of what the swan ancestor probably looked like: a red billed, medium sized bird not too different from more “generic” dabbling ducks. Unlike other anserines, which are specialised terrestrial grazers, swans are specialised aquatic herbivores, feeding on vegetation growing on water, both at the surface and on the bottom, so their ancestor was certainly a rather generalistic anatid. Unlike other duck lineages, this lifestyle appearently did not warrant specialisations to diving, just the much cheaper development of a long, sauropod like neck in order to reach the bottom; indeed, given that swans are huge birds that take part in long migrations, becoming specialised divers would perhaps be a handicap.

Swans are thought to have evolved in western Eurasia, more or less in the area that today is eastern Europe; at the time these birds evolved, the Paratethys Sea still covered large extensions of  Europe and western Asia. Fossil sites from this epoch in eastern Europe unveil rich wetland and coastoal fauna; crocodyllians, choristoderes, sirenians, several dolphin-like basal odontocetes (some possibly related to the modern Ganges/Indus river dolphins), pseudodontorns and giant salamanders are among the many species present in those estuarine ecosystems that are obviously absent from modern Europe. The early swans were the product of these ancient subtropical coastoal wetland ecosystems, and might have even fed on seagrasses alongside the now gone european manatees and dugongs.

Subsequently, Cygnus became widespread across the northern hemisphere, but their diversity center seems to have remained in Europe, even after the Paratethys and it’s coastoal wetlands began to dry out; only by the Pliocene did the european species began to decline in number, presumably as the climate became colder on the onset of the ice ages. Currently, the only species occuring naturally in Europe are the sometimes sedentary, sometimes migratory Mute Swan (Cygnus olor), the very migratory Whooper Swan (Cygnus cygnus) and some wintering populations of Bewick’s Swan (Cygnus [columbianus?] bewickii), but the ancient european swan diversity blessed the continent with yet another species in the Pleistocene:

Cygnus falconeri, before utterly destroying a poor dwarf elephant, by Talk about overkill.

Cygnus falconeri was bigger than the modern largest swans (a tough competion between the mute and trumpeter species) by one third, the largest individuals being as tall as 2,10, being slightly smaller than an ostrich and certainly capable of attacking the scalp of most people. This bird was likely the largest animal in it’s insular ecosystem of Pleistocene Malta; the native dwarf elephants were considerablyheavier, but dwarfed (eh) by it’s enormous frame, likely augmented by it’s enormous wingspan. Given it’s size (modern swans already have efford taking off) and it’s insular habitat, it was most likely flightless, but it retained it’s adaptations for flight, namely the enormous wings (it is possible that, much like the Nene-nui, Cygnus falconeri was “semi-flightless”, with some individuals capable of flight, others theoretically capable of flight but not displaying the behaviour, and others completly flightless; interbreeding with mute swans could have muddied things further, just like the hybridisation between the Nene and the Nene-nui).

Unlike modern swans, Cygnus falconeri probably foraged more on land, something encouraged by the dry mediterranean climate and by the absence of competitors and predators (Gyps melitensis was appearently the largest maltese predator through most of the Pleistocene). In particular, it’s long neck could had allowed the exploitation of browsing niches, much like an avian giraffe (or, if you preffer, an anatid moa/vorompatra), although given the prefference of modern anserines for grazing, I suppose that it was probably a closer equivalent to the long gone diplodocid sauropods, which were long necked grazers as well. This bird governed Malta through most of the Pleistocene, but became extinct before human beings arrived to Malta. The general consensus seems to be that it disappeared due to the temporary landbridge that formed between Malta and Italy, allowing predatory mammals like wolves, hyenas and bears to feast on the native fauna, which evolved with vultures as their only predators; however, I’ve seen discussions that declare a much latter date for the extinction of Cygnus falconeri.

Meanwhile, North America and Australia appearently became the centers of swan diversity, with several species known from fossil sites  in both landmasses; most of these species became extinct when humans arrived, just like the contemporary giant mammals that co-existed with these birds. Swans never really became established in Africa, and so far only one species is known from South America.

Swan phylogeny has often been the subject of debate, but there seems to be a consensus that two groups are formed: one consisting of the Black-necked Swan (Cygnus melancoryphus) and the Black Swan (Cygnus atratus), the two southern hemisphere species, and another group composed of the Arctic species, the Whooper Swan, the Tundra Swan (Cygnus columbianus), the Trumpeter Swan (Cygnus buccinator) and the Bewick’s Swan. The first appear to be clearly left-overs of an older lineage common in the northern hemisphere, reduced to southern hemisphere relics (the extinct australian forms, as well as a few of the extinct north american ones, were almost certainly part of this lineage as well), while the latter evolved much more recently, from a single population that was fragmented by the expansion of the glaciers, and indeed nearly all Arctic swans look the same (the trumpeter probably diverged first, followed by the whooper).

Things are more problematic, however, when it comes to the placement of the Mute Swan. Different studies have either placed this species as part of the southern hemisphere swan clade, sometimes as even the sister taxa to the Black Swan, or as having diverged after them and thus being more closely related to the Arctic swans. In terms of behaviour, mute swans more closely resemble their southern relatives, being mostly sedentary and only occasionally migratory or nomadic, although the migratory behaviour of the Arctic swans is most certainly a more recent adaptation, having evolved in order to cope with the ice ages. In terms of colouration, both the mute swan and the southern hemisphere species have bright red/orange beaks, while the Arctic swans either have completly black or black and yellow beaks, although again it is possible that the Arctic species evolved from species with red beaks that lost that colour in order to cope with the harsher climates. Curiously, both southern hemisphere species are dark in colour while their northern cousins are pure white; this in part supports the idea that they are closely related and that may indicate that the ancestral swans were dark coloured, but if the Mute Swan is part of this clade, then it lost the dark feathers for some undiscernible reason.

Swans are Crane mimics

The Siberian Crane co-exists with two swan species; it competes with both for food. Curiously, it's wintering spots tend often to further away than the swans', further evidencing competition between these birds.

Swans, as previously discussed, are specialised herbivores, feeding almost exclusively on aquatic vegetation, and on submerged vegetation in particular. For this end, they became long necked birds, well adapted to probe deep waters, and they thrive on cold freshwater ecosystems such as the lakes that dot the tundra. They’re also white birds that migrate for miles each year, and are monogamous even though their closest relatives are infamous for their deviant sexual habits. Their trachea’s form loops on their sternum, even diving on it’s inside, an adaptation with the means of aiding with their voacalisations; due to this loop, and their long neck, a vast section of “dead air” means that swans rely a lot on anaerobic muscle power.

If this sounds familiar, it’s because you’re at least somewhat aware of the existence of Gruidae, a lineage of stork-like specialised Gruiformes known vernacularly as “cranes”. And within this clade, it’s the genus Grus in particular that I am talking about; the other cranes spend most of their time foraging on land like anorectic bustards, but Grus cranes specialised in feeding in wetlands like their rail relatives, using their long beaks and necks to gain access to submerged aquatic vegetation to feed on. Unlike swans, they aren’t very competent swimmers, but when you have long legs to wade around, swimming becomes irrelevant. Specially since most wetlands are not even very deep anyway.

Indeed, both swans and most Grus cranes function similarly in their freshwater ecosystems; both birds are almost exclusively vegetarian, feeding on the same plant species, both are primarily Arctic migrators with a few species living permanently on warmer climates (Australia in particular was until very recently a hotspot of both crane and swan diversity), both nest on the ground/shallow waters and both are vicious enough that the only predators that consistently pose a threat are eagles. Even the native mythologies about cranes and swans tend to offer similar connatations to both birds.

Indeed, I dare say that competition between both clades seems to have shape the evolution of both Cygnus and Grus; in North America, the native cranes tend to be more carnivorous, while the european and australian species are generally more terrestrial, all coinciding with the existence of hotspots of swan diversity; there’s also no cranes in South America, and the presence of a single swan species suggests that the latter won the evolutionary battle for the continent’s appearently few resources that matter to these birds. Likewise, swans as a whole are less common on the hotspot of crane diversity, eastern Asia, and their wintering spots tend also to be different, as an efford to avoid competition.

East of the Sun and West of the Moon, mating with both in exchange for eggs

The greek god of light Apollo, riding a swan like a pro.

As to be expected, swans were always seen as icons of angelic grace. Their white feathers were honoured in India as the symbol of ascension, representing enlightment as they were always on water but never wet, as saints were always on this world but never attached. The Celts honoured the swan as the herald of Lugh, god of the Sun, love and arts, while the Greeks likewise connected the swan to their own god of light and rationality, Apollo[n]; saint/goddess/whatever Brigid was also associated with swans, as was Artemis and Selene, the latter sometimes depicted with swan wings. Swans feature in norse mythology as drinking from the Well of Urd, painting their feathers white and obtain countless wisdom; the greeks called the black swan “avis rara”, mocking the notion of dark coloured swans as impossible (the swans, of course, got the last laugh), while all over Europe and North America the swans were seen as holy harbingers of Spring, their white wings signalling the rise of the glorious Spring Sun while the ice melted.

However, somestimes swan symbolism gets plain weird. Zeus is iconic for turning into a swan to rape Leda, a legend thought to have been brought upon by the nasty raping habits that Anatidae as a whole praises (amusingly enough, swans don’t seem to engage in rape behaviour, but contrary to popular perceptions of monogamy, they are rather promiscuous, cheating on each other all the time; monogamy is just social among birds, rarely extending to perfect fidelity). In Australia, swans often represent ancient tribes that became prideful or evil in some way, being turned into swans in order to warn future generations. In Finnish Mythology, a swan lives in Tuonela, the world of the dead, swimming in it’s river in solemn creepiness.

Even the seemingly obvious connatations with Apollo seem rather non-inoccent upon further inspection. Apollo, like his sacred birds, is a god of light and sometimes of the Sun (when mixed with Helios) or the Moon (in his aspect as a silver plaguedealer), and his association with arts has been linked with the mute swan’s own lack of vocalisations, in the famous Swan Song that heraldic swans sing as they’re about to die. Apollo is also a god associated with homosexuality (or, more accurately, bisexuality, but I digress); his most iconic lovers were Hyacinth, Cyparissus, Hymenaios and Branchus (the latter also being his son).

In reality, swans are among the animals with the most cases of documented homosexual behaviour. Over one quarter of all black swan pairings are thought to be between males of the same sex, and the same is thought to be the case of other swan species. This is notable because most homosexual pairings among birds are thought to be lesbian in nature, with seabirds in particular often having entire colonies of female/female pairings. Swans, penguins and flamingos, on the other hand, seem to exhibit primarily male/male couples; this dichotomy between male homosexuality and female homosexuality among birds is not well understood, but it is thought to be derived from their breeding strategies.

Contrary to what homophobes think, being gay =/= impairs reproductive advantage. In many birds colonies where homosexuality is prevalent, the opposite sex still functions as donors. Male seagulls provide sperm, female swans provide eggs. As often noted, this is not indicative of bisexuality, as the birds in same sex pairings may only mate once or twice in their entire lifetime with a bird of the opposite sex, thus indicating that it’s only opportunism brought about by parental instincts. In most birds, said donors tend to be the males, because they are sadly the most expendable gender when it comes to reproduction, but in swans the male is stronger than the female as usually amidst anatids, so females in areas where homosexual behaviour is predominant are set free from their reproductive duties, left to enjoy an hedonistic lifestyle while the males receive fertilized eggs.

Both males in the couple then incubate the eggs and raise the chicks, much like hetero pairs. Studies show that, because males are stronger than females, chicks raised this way have higher chances of survival in harsher conditions, with predators consistently kept at bay. However, this strategy seems to work less efficiently when conditions are less harsh, presumably due to some advantage straight parents have in normal conditions. This doesn’t seem to be the case with lesbian couples amidst other bird species, whose efficiency doesn’t faulter when conditions are harsh or not.

Directing swan homosexuality with deities associated with homosexuality seems at large fitting, even if in retrospect. The concept of the Swan Song is nearly always accompanied with the concept that the Mute Swan lives in agony, tormented by existence and crying a final hymn when life reaches it’s finale, dying in joy to ascend to the Sun (in Pythagorean Theology, Helios acts as a psychopomp, because in reality there are five suns; the physical Sun we see, Helios, and the Suns in higher realities, the next one in the line being Apollo, which receives the ones who found enlightment).

A life tormented, to find joy in death, is easily the most chilling connatation the swan has received, not only as a metaphor for repressed homosexuality or people with suicidal wishes in general, but because swans are white, and white, not black, has always been the colour of death in nearly all cultures, with black only representing death in modern christian thought. To see the swan as harbinger of death is not delusional (swans can still break your arm, after all), but rather an expression of purity gone wrong, of a life so pure that it doesn’t belong to this reality, that belongs decaying in a swamp or ascending to the Sun like a phoenix. Both Greek and Finnish mythology intersect in this aspect, given the previous mention of the swan in Tuonela.

In life or death, the swan is something that should not be, an angel from a reptile, a crane in a goose, a soul in constant agony in the most cheery of all bodies.

One Comment leave one →
  1. April 23, 2013 7:18 am

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