Monday, 23 March 2015

Week 16: Woodnymphs to Kingfishers

Day 63: Marvellous spatuletail – Loddigesia mirabilis

What’s in a name?

That which we call a marvellous spatuletail, by any other name, would NOT smell as sweet. Although ‘hummingbird’ itself is an interesting name, with so many hummingbird species of such fantastic diversity, naturalists have given them a variety of interesting and colourful names which encapsulate their beauty. Although more descriptive names might be more useful for hummingbirds, I think the suite of monikers adorning these birds really add to their beauty and mystique.

Some hummingbirds’ names refer to their prominent features including the streamertails (Trochilus), firecrowns (Sephanoides), coronets (Boissoneaua), pufflegs (Haplophaedia), plumeleteers (Chalybura), trainbearers (Lesbia), metaltails (Metallura) and visorbearers (Augastes). Some species appeal to groups of people such as the cowled jacobins (Florisuga), the enticing coquettes (Lophornis) and the native caribs (Eulampis) and incas (Coeligena spp).

Colour is often referenced in hummingbird names from the drab hermits (Phaethorninae) to the brightly-coloured brilliants (Heliodoxa). Many names reflect the jewel-like appearance of hummingbirds such as the topazes (Topaza), emeralds (multiple genera), sapphires (Hylocharis), moutaingems (Lampornis) and sungems (Heliactin). Other groups have a more cosmological feel including the sunbeams (Aglaeactis), hillstars (Oreotrochilus spp.), starfrontlets (Coeligena), comets (multiple genera), starthroats (Heliomaster) and woodstars (multiple genera). Finally, some species appeal to the mythical creatures they resemble such as the woodnymphs (Thalurania), sunangels (Heliangelus), sylphs (Aglaiocercus) and fairies (Heliothryx).

One of the most impressive to look at and glorifyingly named, the marvellous spatuletail is a stunning hummer coated in green and bronze with a dazzling turquoise gorget. The defining feature of this Peruvian endemic is the pair of long, indigo, racquet-like tail feathers which extend from a tail unique for only having four feathers. As with many hummingbirds, it is the males who bear the brightest colours and most extreme ornamentation. He waves his spatulas in a frenetic courtship display in the hope of wooing a mate.


Day 64: Bee hummingbird – Mellisuga helenae

Life in miniature

The bee hummingbird is the smallest bird in the world. If you saw this creature flitting round the gardens of its native Cuba you could be forgiven for thinking it a bee or moth on first inspection. It weighs just 1.6-2 g. A male and female perched together on your hand would weigh as little as a penny! Normally its wings beat around 80 times a second but the courtship flight of a male raises that rate to 200 per second! It is the bird with the fewest feathers and the highest body temperature and its heart beats up to 1,200 times a minute. Meeting its dietary needs means consuming half its weight in food and eight times it weight in water every day. In most hummingbirds, nests are small cups fused to twigs and, unsurprisingly, the bee hummingbird’s is especially tiny, being just 2.5 cm in diameter and made from plant matter, lichen and cobwebs. The smallest bird on the smallest nest must be and endearing sight!


Day 65: Gartered trogon – Trogon caligatus

Toe to toe

The gartered trogon is one of 39 trogon species in the order Trogoniformes. They are colourful, arboreal birds which feed on insects and fruit in generally tropical forests around the world. Trogons are cavity nesters, using their strong bills to gnaw at rotting wood. This has given them the name trogon which means ‘nibbling’ in Greek. Gartered trogons, as well as the related violaceous trogon (T. violaceus), can also nest in wasp, ant or termite nests.

Let’s use this opportunity to talk feet. The archetypal bird foot is anisodactylic with digit 1 pointing backwards and digits 2 to 4 facing forwards. This is more than adequate for the many terrestrial and aquatic orders we have come across so far. However, on the road to passerines it various groups have tried a multitude of digital arrangements.  Trogons are, uniquely, heterodactylic meaning that both digits 1 and 2 face backwards. This arrangement is tailored for an arboreal life, allowing trogons to grip branches firmly. 

This is similar to the zygodactyly of the parrots, cuckoos and woodpeckers however, in their case, it is digits 1 and 4 which face backwards. Owls and ospreys are able to adopt this arrangement when needed, for example when catching prey, but they can also revert back to more typical anisodactyly. In mousbirds and some swifts, both digits 1 and 4 can be rotated to either face forwards or backwards. The swifts use this ability to have all four toes face forward to hook onto rough surfaces. Last but not least, the kingfishers and allies (Coraciiformes) have partially fused second and third digits. Since birds have two limbs devoted to flight, the feet have become increasingly important for interaction with the environment and thus they have been adapted to meet many needs. 


Day 66: Laughing kookaburra – Dacelo novaeguineae

Merry merry king of the bush is he

The laughing kookaburra is one of Australia’s most famous birds, known for its loud, cackling call. Whilst staying in Cairns, a pair of kookaburras (or just kookas) would frequent the garden, sitting together and raising their heads in howls of laughter. They would then plop to the ground where they bounced around in search of food. They eat a lot, mainly taking large insects, reptiles and small mammals and birds. One thing they don’t eat is fish, despite being the largest species of kingfisher.

The laughing kookaburra is one of five kookaburras which also include the demonic blue-winged kookaburra (D. leachii) and the impressive shovel-billed kookaburra (Clytoceyx rex). Kookaburras are part of the Halcyonidae, collectively known as the tree kingfishers. None are fish specialists, preferring to watch and wait from selected vantage points. There are two other families in the kingfisher suborder Alcedines. Our own Eurasian kingfisher (Alcedo atthis) is a member of the river kingfisher family Alcedinidae. These are a group of small, bright kingfishers with short tails and long, thin beaks. By contrast, the water kingfishers of the Cerylidae are larger and chunkier with longer tails and more dour plumage. They are also the only kingfishers found in the New World. Both of the families are expert fishers, plunging head first into the water from well-placed perches.

When allocating Latin names to the kingfishers, naturalists made some interesting decisions. The genus of river kingfishers Alcedo was a reasonable enough starting point. It is the Latinised version of Halcyon which refers to the Greek myth of Alcyone. Alcyone and her husband Ceyx referred to themselves as Hera and Zeus. This sacrilege invited the wrath of Zeus himself who smote Ceyx, aboard his ship, with a lightning bolt. When Alcyone was informed of his demise (via the dream god Morpheus), she threw herself into the sea. Out of pity, the gods transformed the couple into kingfishers. Every year, Alcyone would lay her eggs in a nest on the beach whilst her father, Aeolus, protected her efforts by calming the waters of the sea for a week either side of the winter solstice. This is the probable etymology of the phrase ‘Halcyon days.’

So far so good, we have bird named for a mythical kingfisher. Next, Alcyone’s husband Ceyx gets its own genus, the dwarf kingfishers (Ceyx). I suppose that’s fair as Ceyx was also transformed. We also have the long-tailed paradise kingfishers in the genus Halcyon which is simply the Greek from of Alcedo. This is a touch lazy, although Greek and Latin tautology does exist a lot in binomial names (e.g. Corvus corax means crow-crow in Latin then Greek). Follwing this, naming just got silly. The Dacelo kookaburras we started with are actually just an anagram of Alcedo as is the banded kingfisher genus Lacedo. Finally the belted kingfisher Megaceryle alcyon once again references Alcyone. With a wealth of binomial Latin names in existence, biologists have demonstrated their imagination abundantly but they were having an off day with the kingfishers.


Tuesday, 17 March 2015

Week 15: Nightjars to Emeralds

Day 59: Standard-winged nightjar – Macrodipteryx longipennnis

Flying the flag for nightjars

I struggled to find a decent photo of this splendid nightjar and that fact alone is indicative of nightjar ecology. Like the owls and frogmouths before them, nightjars are crepuscular or nocturnal. However they generally live in dry, open country and nest on the ground so observations are generally made of shapes flying through the twilight or spotlights picking up flashes of white.

Nightjars are able fliers and hawk insects from the air, using their facial bristles to aid in capture and protect the large eyes from flailing insect wings. Measuring nightjar beaks was intriguing as there was little beak protruding from the skull yet the gape was as wide as the head, presumably allowing nightjars to effectively become flying mouths. Their aerial insectivory lends some American species the name 'nighthawk' whilst the name ‘nightjar’ was originally ascribed to the European nightjar (Caprimulgus europaeus) where it referred both to their nocturnal activity and their remarkable ‘churring’ song which reverberates around British heathland. The genus Caprimulgus harks back to an old myth that nightjars suckled from goats (capra=goat, mulgere=to milk).

Although generally cryptic in colouration, male nightjars are often distinguishable from females by patches of white on the throat, wings or tail. These are used in courtship displays; the white reflecting whatever light might be left in the sky. This African species is notably one of a few nightjars that have modified their feathers in unusual ways to aid in courtship. The remarkable ‘standards’ of this species are actually modified primaries which can be erected during courtship. 



Day 60: Edible-nest swiftlet – ­Aerodramus fuciphagus

Enjoy the bird, not the nest

To look at, this swift is a typical member of the swift family Apodidae. Swifts are characterised by their extremely long, scythe-like wings which make them supreme fliers. They are able to reach tremendous speeds with the white-throated needletail holding the record for the fastest bird in flapping flight: an eye-watering 112 km/hr (or more?!). You may have seen our own swifts (Apus apus) hurtling round the spires and towers of our towns and cities where they come to breed in the summer. Like the closely related nightjars, swifts are aerial insectivores however they hunt during the day. They spend the vast majority of their time on the wing, eating, drinking, mating and sleeping in mid-flight! In fact, the genus Apus (from which the names of the family and order are derived) means ‘without feet’, referring to early misconceptions about swifts.

When swifts do come to land it is to breed. For us, in the UK, we are used to seeing the convergently evolved (Passeriformes: Hirundinidae) swallows (Hirundo rustica) and house martins (Delichon urbica) building their nests under eaves and beams where they bind together mud with their saliva. Swifts engage in similar practices but using vegetation over mud. In the Aerodramus swiftlets, saliva actually forms the majority of the nesting material and in two species, the edible-nest swiftlet and the black-nest swiftlet (Aerodramus maximus), saliva is the sole component.

These two species nest in huge caves dotted around the islands of Southeast Asia. There, they form large colonies, covering the walls with glistening nests. Aerodramus swiftlets navigate around the colonies using echolocation, an ability that evolved just twice in the birds (the other incidence being the caprimulgiform oilbird, Steatornis caripensis).

Such large congregations of birds make them easy pickings, not least for humans. In China, the nests of edible-nest and black-nest swiftlets have becoame a delicacy. Served in the infamous bird's-nest soup, swiftlet nests are believed to bring health benefits and an increase in libido. This has resulted in the harvesting of some colonies to near extirpation. In Hong Kong, the centre of swiftlet-nest trade, demand is high and nests can fetch up to $10,000 per kilo. The creation of artificial nesting sites has allowed a regular supply of nests, shifting attention away from the natural sites. As with ivory and tiger claws, it is important that China, and the rest of the world, comes to value our wildlife living rather than dead. 


61: Antillean crested hummingbird - Orthorhyncus cristatus

The first of many

I have a hazy memory of the first time I ever saw a hummingbird. In the year 2000, my sister and I were walking through hotel gardens on the south coast of Grenada in the Lesser Antilles of the Caribbean. There, a tiny bird reared up in front of use before promptly wizzing off in the opposite direction. The bird in question was an Antillean crested hummingbird, one of over 340 hummingbird species in the second largest bird family Trochilidae. There is plenty to say about hummingbirds but luckily I will have a few days to explore them.

Humming birds are generally placed in the same order as the swifts (Apodiformes). Why this might be is an interesting question as hummingbirds are very un-swift-like but both groups share similar wing morphologies. However whereas the swifts use their wings to zoom through the skies, the hummingbirds have their own specially adapted flight. Hummingbirds are expert fliers, able to hover and fly forwards, backwards, up, down and sideways. Using a technique similar to insects, hummingbirds flex their wings in quick beats generating lift on the upward stroke in addition to the downward stroke by inverting the curvature of the wing. With typically between 50 and 80 wing beats per second, the hummingbirds are able to generate sufficient lift to facilitate their astounding aerobatics.

To support their flight, hummingbirds need additional adaptations. They have light, hollow bones with fusion in the vertebrae to remove unnecessary muscles and ligaments. The feet are also tiny to reduce weight and drag. By contrast the sternum and pectoral muscles are enlarged for increased wing power and strong finger bones stabilise the wing. Perhaps the most important requirement for hummingbirds is the highest metabolism of any non-insect animals. Their hearts beat up to 20 times per second and they can breathe 250 times per minute.

Such high energy expenditure requires near-constant feeding and hummingbirds have made use of the most energy-rich and readily-available food supply: nectar. They must visit hundreds of flowers a day in order to drink more than their weight in nectar. Their metabolism ensures sugar makes it to the muscles as soon as possible but this means hummingbirds are never far from dying, so constant is their energy need. Overnight, hummingbirds must shut down, entering torpor where there metabolism decreases and their body temperature halves. This sleep-like state allows them to get through the night until the first drink of the morning. Being a hummingbird sounds knackering! 


Day 62: Violet Sabrewing - Campylopterus hemileucurus

The purple flash

Violet sabrewings are one of a few hummingbird species that I encountered on my trip to Honduras in 2011, my first Noetropical experience since Grenada. The sabrewing is a stunning bird and the name aptly describes both the striking purple colouration and the unusual wing morphology. I had assumed that 'sabrewing' referred to the overall shape of the wing however closer inspection reveals the broad, flattened central rachis of the first two primary feathers which resemble curved swords. We speculated as to the function of these structures and, being only on the males, the most likely explanation seemed to be for noise generation (by rattling) during competitive displays. Sabrewings are polygynous birds, holding leks where they attract and court multiple females.

Another intriguing difference between males and females is the difference in bill morphology. Despite being smaller, the female has a longer and more curved beak. Again, the explanation for this was not obvious but could be a difference in foraging behaviour between males and females. This is highly unusual in the bird world as most members of the same species tend to have similar diets. It could be the case that males and females specialise on different flowers so as not to compete with each other. Alternatively, (and perhaps more likely), if longer, curvier bills are indicative of greater flower specialisation, perhaps females must specialise more in order to access richer nectar sources for their young. As hummingbirds also eat invertebrates, the same arguments apply for insectivory i.e. males hawking for aerial insects whilst females glean them from foliage.

I have already alluded to the close association of hummingbirds and flowers and what a fascinating relationship it is. Hummingbirds, much like bees or butterflies, are in search of the nutritious nectar provided by flowers. Plants are not altruistically offering a free meal but instead aim to be pollinated with pollen transferred from other flowers. This requires that the pollinator regularly visits flowers of the same species which they may not if they are maximising their opportunities by visiting many species. Flowers can increase exclusivity by making nectar trickier to access. The presents an opportunity for some pollinator species to specialise, gaining a monopoly on particular plant species. These evolutionary pressures often lead to arms races with plant and pollinator become increasingly specialised.

The myriad of plant-hummingbird interactions have lead to diverse beak and flower morphologies. Many flowers will have multiple pollinator species whilst many hummingbirds will pollinate multiple plant species. Some hummingbirds have taken specialisation further. Hermits (Phaethornithinae) have very long, curved bills whilst sicklebills (Eutoxeres spp.) have extremely curved bills for flowers of the family Gesneriaceae. The sword-billed hummingbird (Ensifera ensifera) is the most extreme in specialisation. It uniquely has a bill longer than the rest of its body which it uses to drink from just one species: Passiflore mixta. From the perspective of both plants and pollinators, the extent of specialisation can be seen as a choice of strategies with different species (or even sexes, as above) following different evolutionary trajectories. Generalists benefit from accessing abundant yet low value nectar whilst pollinating haphazardly whereas specialists offer a more reliable service in return for a rich reward.With so many possibilities and niches to fill it is perhaps not surprising that up to 25 species can coexist in the same area.

Monday, 9 March 2015

Week 14: Barn Owls to Frogmouths

Day 55: Moluccan masked owl – Tyto sororcula

A rainforest spectre 

In almost every habitat in the world there is an owl who owns the night. Whilst the Accipitriformes and Falconiformes have dominated the diurnal predatory niches, the Strigiformes have taken the equivalent nocturnal positions. The Strigiformes comprise just two families, the Tytonidae (barn owls) and the more speciose Strigidae (true owls). The Molccan masked owl is a member of the former which are known for their heart shaped faces and sometimes ghostly appearance.

There isn’t a huge amount known about this owl species but it is part of a complex of similar species which inhabit the islands of Indonesia and Papua New Guinea. None are well described but one species, the Seram masked owl (Tyto almae), has proven especially elusive. It was first noticed from a photograph taken in 1987. It wasn’t until 2012 that a team from the Natural History Museum of Denmark and the Indonesian Institute of Sciences caught a masked owl in a mist net in Seram (an island in Indonesia) and later described it as a new species. It is still known only from these two records.

The predatory nature of owls might suggest close relatedness to other birds of prey but ornithologists may be quick to point out that owls are distantly related. However, recent research suggests that the predatory hawks, falcons and owls all share a raptorial ancestor but each lineage contains more closely related bird bird groups who have independently evolved almost every other form of dietary ecology. WARNING: below comes a taxonomic blur, not for the faint-hearted.

Traditionally, and matching the order of my work and this blog, the owls are amidst the ‘near-passerines’, the serious of groups that take us closer and closer to the passerines. This started with the soundgrouse followed by the pigeons, parrots, turacos and cuckoos and after the owls come the nightjars, swifts and hummingbirds, mousebirds, trogons, hornbills, kingfishers and woodpeckers.

Recent genomic analysis upsets this tradition remarkably. Soundgrouse and pigeons are shoved to the edge of the Neoaves (i.e. all bird but ratites and fowl) with the mesites, flamingos and grebes in the clade Columbea. The much larger clade Passerea contains, at the edge, the cuckoos, turacos, bustards, nightjars and swifts. Next comes the hoatzin, paired with the shorebirds, followed by the remaining seabirds in the rough order we covered them. The rest of the orders are land birds containing the Afroaves (raptors, owls, mousebirds, trogons, hornbills, kingfishers and woodpeckers). Finally, the nearest birds to the passerines are the two seriemas, the falcons and the parrots. Congratulations if you have followed this, I certainly find it very confusing. For a superior explanation see Jarvis et al (2014).


Day 56: Snowy owl – Bubo scandiacus

Call me Hedwig one more time!

This gorgeous white owl may be famous for being Harry Potter’s companion but reality is far more interesting. As one might predict, the snowy plumage indicates the owl’s circumpolar distribution where it breeds in the northerly reaches of the Arctic tundra. The snowy owl might look cute and cuddly but it is a proficient hunter of a variety of Arctic wildlife. Although the diminutive lemming is its main food source, the snowy owl can hunt large rodents, rabbits and hares as well as ducks, geese, waders and gulls. The females have even been known to take other owls and raptors. Like other owls, snowy owls have very large gapes enabling them to swallow prey whole. All the non-digestible parts of the prey are coughed up as pellets of hair, bone and teeth. Pellets are useful for biologists who wish to characterise the diet of owls. As for the raptors, female owls are the larger of the two sexes. 
As well as being smaller, male snowy owls are more uniformly white and thus a better choice for the Harry Potter films despite playing a female owl!

The snowy owl is a more unusual character in the genus Bubo which comprises the horned owls and eagle-owls. The Eurasian eagle-owl is one of the largest, with a wingspan over 6 ft. By comparison, the elf owl (Micrathene whitneyi ) would easily fit in your pocket. The Strigidae are a large family of owls which include most of the owl species you might be familiar with.


Day 57: Short-eared owl – Asio flammeus

Silent but deadly

Two owls down and yet I haven’t discussed some of the fantastic adaptations that unite the owls. Well, what better way to do it than with one of my favourite British birds, the stunning short-eared owl. As we know, owls evolved for the night. As it happens the shorties are observable during the day, as I have seen them over the grassy fields of Portland Bill, however the majority of their hunting occurs at night when a battery of adaptations come into effect.

In the blackness of night, owls must be able to navigate effectively and hunt small prey which may skulk, unseen, in the undergrowth. To accomplish this, owls have fantastic night vision. They have very large eyes with wide pupils which maximise the amount of light passing to their sensitive rod cells. Furthermore, an advanced neural mechanism extracts a superior amount of information from the retina. Owl’s eyes face forward enabling excellent binocular vision so that they can judge distance and depth efficiently (this eye placement gives them their anthropomorphous, ‘wise’ face). Fitting such large eyes into a relatively small head results in their tubular shape. ‘Eyetubes’ cannot be swivelled as eyeballs can, so owls must rotate their heads. They famously do this rather well, being able to rotate the head through up to 270°.

The other sense which owls have mastered is hearing. Don’t be fooled by their ‘short ears’. The ears of the ‘eared owls’ (Asio) are simply tufts of feathers, thought to break up the outline of a roosting bird. The real ears are set on either side of the head, beneath the feathers. Here, we encounter another case of asymmetry as owls’ ears are placed at slightly different heights. This allows owls to pinpoint sounds in three dimensions. We (and many other animals) are good at locating sound horizontally but vertical pinpointing is quite difficult (hence why I sometimes put my head on my side to find singing skylarks). The ears are aided by a ruff of stiff feathers that form a facial disk which collects sound waves and directs them to the ears. Even the beak is designed to minimise sound loss. What was striking when measuring the owls was how small the skeleton was beneath the feathers and how structurally different the facial feathers were to achieve their function.

Owls are excellent at detecting prey but they also need not to be detected. Thus, owls are literally deathly silent. Large, broad feathers allow owls to float around with minimal flapping. Along the leading edge of the wing feathers, serrations disrupt the turbulence that would cause sound in a normal bird whilst velvety down absorbs some of the sound that is still produced. As this silent spectre descends, with talons spread, an unsuspecting vole stands little chance.


Day 58: Papuan frogmouth – Podargus papuensis

Why the wide face?

The frogmouth is a bizarre looking bird - that is, if you were able to spot it as they exhibit impressive cryptic plumage and behaviour. They roost motionless with their necks outstretched which, coupled with their mottled-brown plumage, makes them resemble a branch, complete with a stick-like head tuft. All frogmouths have extremely wide gapes (hence the name) and the Papuan frogmouth has the largest of them all. Like the owls before them, the frogmouths are nocturnal foragers however their foraging mode is slightly more ungainly. They hunt around the ground for insects but will take prey as large as rodents, lizards and frogs. A huge mouth helps the frogmouth to grab prey in the dark without needing the finesse of an owl.

The frogmouths (Podargidae) are part of the nightjar order Caprimulgiformes. The species of this order are characterised by their nocturnal, insectivorous habits and their cryptic plumage. A morphological perspective placed the Caprimulgiformes between the owls and the swifts and it is easy to see why. Since then they have been affiliated with various ‘near-passerine’ groups and genetic evidence has confused matters although there was strong support for a monophyletic clade uniting the nightjars, owlet-nightjars (Aegotheliformes) and the swifts (Apodiformes). This is corroborated in the most recent whole-genome analysis however this clade (the Cypselomorphae or Caprimulgimorphae), is placed nowhere near the owls. Instead, they are located with cuckoos, bustards and turacos on the periphery of the Neoaves. I’m still trying to get my head around this new, counter-intuitive world order of bird.


Monday, 2 March 2015

Week 13: Parakeets to Cuckoos

Day 51: Ring-necked parakeet - Psittacula krameri

Little green men

The large, natural range of this small, long-tailed parrot is a band of sub-Saharan Africa and the Indian Subcontinent yet you probably know it better as the brightly-coloured alien that flocks around the parks and gardens of London and the surrounding counties.

As a popular pet, the ring-necked parakeet found its way into many urban localities where a combination of escapes and deliberate releases has lead to a number of stable feral populations in cities around the world including in South Africa, Japan, Turkey and North Africa. There are even more parakeets in Europe with populations in Belgium, the Netherlands, Germany, Italy and Spain. The biggest feral population is centred on south-west London.

The origin of the 'Kingston parakeets' is a bit of a mystery. Some suggest that they escaped from an aviary after the storm of 1987 or from the set of The African Queen whilst others believe Jimi Hendrix to have released a pair from Carnaby Street in the 1960's. In a city with many green spaces but few predators, the parakeets probably had little problem establishing a population but it would have taken some time for sufficient numbers to build up and become apparent. The population really began to expand during the 1990's, growing exponentially. Currently their numbers are difficult to estimate but there could be as many as 50,000 birds with flocks 6,000 birds strong.

It seems reasonable that such a large population of alien birds would have some impact on the wildlife of London and its environs. One study has found that typical garden birds are avoiding the large, noisy parrots who frequent the bird feeders of the masses. Parakeets also nest in tree holes which necessarily means that other species such as starlings or woodpeckers are loosing prime nesting habitat.

The ring-necked parakeet is just one of many invasive alien species to have reached our shores over the decades and centuries. Successfully invasive 'candidates' must occupy a niche which means either taking a vacant place in the ecosystem or pushing out a native equivalent. Vacant niches are hard to come by so our native flora and fauna tends to take a beating via competition for resources or apparent competition mediated by shared natural enemies such as Squirrel parapoxvirus and crayfish plague. Moreover, invasives are often released from the predation or parasitisation of their native lands allowing unchecked population growth. Grey squirrels, signal crayfish, American mink, harlequin ladybirds and Japanese knotweed are all well-known scourges of British wildlife and the list is growing.    


Day 52: Hoatzin - Opisthocomus hoatzin

The crested claw

The hoatzin is a pretty odd bird to look upon. With maroon eyes set in a naked, pink face, a large, orange crest at the end of a long, slender neck and broad, rounded wings and tail the hoatzin resembles the most ancient ancestors of the birds. Furthermore, the chicks possess two clawed fingers on each wing bearing a striking similarity to the feathered arms of the small, Jurassic raptors. The uniqueness of this species meant its placement in its own order: the Opisthocomiformes. However, placement of this order on the bird tree has been very difficult with suggested relatedness to tinamous, gamebirds, rails, bustards, sandgrouse, doves and cuckoos. It took whole-genome sequencing to resolve the issue so that the hoatzin is now paired with the clade comprising Gruiformes (cranes and rails) and Charadriiformes (shore birds).

The hoatzin lives among the dense, luscious vegetation bordering the rivers of the Amazon basin. They are herbivores and their diet requires special fermenting ability akin to cows. Unlike cows (and other ruminants) they do not possess a rumen but have an enlarged crop which they use to digest the vegetation. This decomposition leads to an unpleasant smell, giving the hoatzin the alternative name of stink-bird. The hoatzin is a gregarious nester and does not need to fly far to find food. The chicks are obviously unable to fly immediately after hatching so their claws are used to scramble through the branches. This is especially important for when hawks appear as the chicks drop into the water and swim to safety before climbing out when the coast is clear. The claws of chicks lead early taxonomists to conclude that hoatzins where more closely related to Archeopteryx and its kin but the claws are actually a secondarily derived trait.


Day 53:  Great spotted cuckoo - Clamator glandarius

The expert freeloader

Here we have a handsome representative of the Cuculiformes i.e. the cuckoos. You will all be familiar with our own common cuckoo (Cuculus canorus) with the famous song that gives it its onomatopoeic name. You, hopefully, will also know that the common cuckoo is a brood parasite meaning it lays its eggs in the nests of other bird species. What you may not know is that brood-parasitism is actually present in many cuckoo species in both the Old World (56 species) and the New World (three species). The common cuckoo had already been measured so I could not choose it for discussion however the great spotted cuckoo is a worthy stand-in for a discussion on cuckoos and brood parasitism.

Brood parasitism is a fascinating subject with many interesting processes. To go through these ideas it might be easy to start at the beginning. Rearing offspring is costly but many organisms do it to maximise the survival of their young. However if one could avoid that cost one could have more offspring. Many species engage in extra-pair matings so that others might rear their young. The goldeneye (Bucephala clangula) is a duck that has specialised in laying eggs in the nests of fellow females. Interspecific brood-parasitism is just one step further for a species used to pushing its eggs on others.

One might think that that when the first cuckoos began laying eggs in the nests of other species, those hosts would immediately respond. However for an unwitting host, the oblong objects in her nest have, for millions of years, always been her eggs and so she is not primed to notice if even a conspicuously different egg is added to the brood. Moreover, when the chick hatches along with the others, its loud, begging call and bright gape illicits the same instinctive feeding response as the host's actual offspring.

Clearly it is costly for the host species to expend so much energy feeding a chick that isn't its own so their is an evolutionary pressure on the host to act. The host can avoid parasitism by nesting in less accessible places and earlier in the year or it could learn to recognise the cuckoo and harass it until it flees. If this fails, hosts can learn to asses the number of eggs in the nest or to distinguish the cuckoos egg from her own. The host might respond to parasitism by evicting or piercing the new egg or abandoning the nest entirely. However, once host parents begin to evict parasitic eggs, there is a pressure for the cuckoo's eggs to resemble the host. Thus, an evolutionary arms race ensues as the hosts' powers of discrimination increase hand-in-hand with the mimicry of the cuckoo.

Mimicry requires specialisation on the part of the cuckoo and some species do not bother, preferring to have many, infrequent hosts which never cotton on to the cuckoos' ploys. Other species are able to focus on one, sufficiently abundant host species. However there are a few species, including the common cuckoo, which are able to mimic multiple hosts. This was perplexing for biologists as specialisation has a genetic basis yet members of the same cuckoo species could interbreed without messing up the different specialisations. For this to be possible, egg mimicry should be coded by genes which are only passed down the female line such as mitochondrial genes or genes on the female-determining W chromosome. This would allow females to form specialised gentes (singular gens) whilst males can mate indiscriminately as their genes will not affect the mimicry ability of their offspring.

In addition to egg mimicry, cuckoos have developed many other tactics to maximise the chances of their eggs and chicks. Some cuckoo males resemble hawks and lure aggressive host parents away allowing female cuckoos to stealthily lay an egg (the common cuckoo somewhat resembles the sparrowhawk, Accipiter nisus).Some cuckoo species have short incubation times allowing the chick to hatch before the host's. As soon as the cuckoo hatches, it instinctively rolls the host's eggs, one at a time, into a cleft in its back before thrusting them out of the nest. Whether or not to kill one's step-siblings is an interesting question. Research into brood-parasitic cowbirds suggests that when one's host has large chicks its better to evict them early to avoid later competition whereas, when the host's chicks are small, the cuckoo chick can high-jack the collective begging power of the nestmates before stealing the lions share of food.

Who wins this evolutionary arms race? Well there are many reasons why cuckoos should always be one step ahead. Firstly, cuckoos have much more at stake. Every cuckoo necessarily must outwit a host in order to succeed in reproducing. The hosts, however, are far more numerous and therefore parasitism is unlikely for any given couple, reducing the strength of the pressure. Secondly, as cuckoos' eggs become near exact replicas of the hosts' eggs, the hosts must have incredible powers of discrimination or else suffer the risk of accidentally rejecting their own eggs. Thirdly, it is suggested that some cuckoos have strategies which force hosts to comply. The brilliantly named Mafia hypothesis suggests that if a host rejects a cuckoo egg or chick, the cuckoo parents trash the host nest meaning that the risk of defection is greater then the cost of rearing the cuckoo. Evidence for this startling strategy has been put forward for two species, one of which is the great spotted cuckoo which parasitises magpies and is less likely to predate the nests of compliant hosts. #thuglife


Day 54: Little bronze cuckoo  - Chrysococcyx minutillus

Cuckoo jewel

This gorgeous little cuckoo was a regular and welcome presence during my sampling in the riparian rainforests of Cape Tribulation, Queensland. I liked this species because it had a simple yet pleasant descending call and catching a sight of its metallic green back and red eye was always nice, especially when the bird in question was bashing away at a thick, juicy caterpillar. The little bronze cuckoo was a regular attendant of mixed species flocks. These loose groups of insectivores move through the understory, sallying and rummaging for bugs amongst the foliage. When any given species is locally rare, safety in numbers is provided my members of different species including spectacled monarchs(Symposiachrus trivirgatus), rufous fantails (Rhipidura rufifrons), fairy gerygones (Gerygone palpebrosa) and little shrikethrushes (Colluricincla megarhyncha).

Hearing little bronze cuckoos and the other species of the rainforest was not only a pleasantry but a necessity for the purposes of fieldwork. From day one of sampling I needed to be able to identify the species I heard to get an accurate representation on of the presence and absence of the jungle denizens. This meant learning the calls of up to 100 species in advance with only a week or so in the field to actually hear and see them in real life. Thus, much time was devoted to sitting in front of an Australian bird app, playing calls until my partner and I could ID successfully. By the end of sampling, some species were so familiar I could ID them in my sleep but rarer individuals such as the congeneric shining bronze cuckoo (Chrysococcyx lucidus), still required the playback of our digital recordings.

This process of learning bird song in advance gave a real appreciation for its necessity in fieldwork and in birding in general. In the rainforest, most identification is based on song as small, skulking birds at some distance through the dense foliage can be difficult to observe. I'm pretty good at identifying British birds by song or call but that is from years of passive (and sometimes active) learning, one species at a time. However I had never attempted the songster crash course before on other international trips (which would have been especially useful in Honduras). From now on I will make an effort to get to know the birds of a new country in advance, by sight and by sound.

The little bronze cuckoo is another brood-parasitic cuckoo but there are many members of the Cuciliformes which are neither brood-parastic nor especially cuckoo-like including the charismatic roadrunners (Geococcyx spp.).


Wednesday, 25 February 2015

Week 12: Keas to King Parrots

Day 47: Kakapo - Strigops habroptilus

The lovable owl-parrot

If you don't know what a kakapo is yet, you need to get it in your life (see kakapo and Stephen Fry). This is a parrot with an interesting ecology, an endangered past and an adorable personality.

Kakopo is the Maori for night parrot and Strigops means owl-eyes; together these names sum up much of their ecology. The kakapo is endemic to New Zealand and, like the kiwi before it, has lost the power of flight in the absence of terrestrial enemies. Despite this, the kakapo's wings are not redundant but are used to parachute down through the branches up which it ably climbs with sturdy legs. Predators were not, however, absent altogether in the evolution of the kakapo. The existence of visually proficient aerial predators (such as the Haast's eagle or yore) pressurised the kakopos to stay low and hidden, employing cryptic green plumage and a freezing response to danger. At night, they could skulk in relative safety, using their huge eyes, whisker-like vibrissae and keen sense of smell to navigate and forage for plants, fruits and seeds.

As one might imagine, this suite of adaptations offers little protection against a relatively recent threat. Arriving in New Zealand around 700 years ago, the Maoris (formerly Polynesian) and their dogs hunted kakapos for food and feathers and had no problem picking the petrified parrots from the forest floor. Fast-forward 600 years and European settlers began to arrive, bringing with them their own mammalian entourage of cats, rats, stoats and ferrets which made a quick meal of the kakapos and their eggs. The ruthless efficiency of naturalists also played its part, each scientist or collector desperate for a kakapo in their cabinet of curios. Soon it was extinct on North Island and fast declining on South Island.

Into the 20th century, naturalists began to find and relocate kakapos to relative safety which tended to be islands or rugged glacial valleys. However it became apparent during the 1970's that only males were being identified until a small population was found on Stewart Island. A kakapo recovery programme set about relocating all kakapos to islands where predators could be removed and vegetation managed. Strangely, the sex ratio of relocated kakapos was skewed towards males (2:1) which was a concern for conservationists.

A typical conservation strategy may provide supplementary feeding for individuals to increase health and reproductive success. When applied to kakapos, however, the sex ratio became even more skewed in favour of males! In steps the Trivers-Willard hypothesis which states that a mother should manipulate the sex ratio of her offspring based on her condition. The idea is that her condition is correlated with her offspring's condition and condition matters more to males. Females are likely to breed regardless of their size and health however in males (i.e. the competitive sex) a big male may have many offspring whilst a small male has few to none. Thus a well fed female should have male offspring. This insight from evolutionary biology identified both the problem and the solution. The solution was to feed females up to their threshold for reproduction but not so far as to reach their male-biased threshold. The population is now (slowly) recovering and evolutionary theory saves the day!


Day 48: Major Mitchell's cockatoo - Lephochroa leadbeateri

A big fan

The cockatoos (superfamily Cacatuoidea) are one of the three superfamilies in the parrot order Psittaciformes, the others being Strigopoidea (kea, kaka and kakapo) and Psittacoidea (true parrots). All parrots possess a huge, sharp beak which is generally used for crushing fruit and nuts. They also have unusual, zygodactyl feet which means two toes face forward and two face backward. This adaptation makes parrots both impressive climbers and manipulators of food and even tools.

Major Mitchell's cockatoo is one of many cockatoo species in the Cacatuidae family which are confined to Australasia. The cockatoos are noted for their splendid crests: long, often colourful feathers which they fan out when aroused. The word cockatoo has Indonesian origins where it refers to cockatoos' vice like bills. These are kept sharp by the slow grinding of the mandibles. Cockatoos are able to tackle very tough nuts which no animal can deal wiith, pinning them against the roof of their beak whilst the lower mandible dismantles them.

Cockatoos are prevalent in the pet trade. Like many parrots they are colourful and intelligent and they can live for many years with a Major Mitchell's cockatoo called Cookie celebrating his 82nd birthday this year! This cockatoo is named after Major Thomas Mitchell, a 19th century explorer who remarked that 'few birds more enliven the monotonous hues of the Australian forest than this beautiful species whose pink-coloured wings and flowing crest might have embellished the air of a more voluptuous region'.


Day 49: Papuan lorikeet - Charmosyna papou

Red lorikeet, yellow lorikeet

The Papuan lorikeet is a stunning examples of the tribe of Australasian parrots known as lories and lorikeets (Loriini). The dichotomy between lories and lorikeets is not taxonomic but simply a grouping based on tail length with the latter having much longer, tapering tails (this distinction applies to parrots and parakeets too). Like most parrots, lories and lorikeets are also very colourful. The most well known is probably the rainbow lorikeet (Trichoglossus moluccanus), a raucous and conspicuous denizen if Australian suburbs. In my humble opinion, there is such a thing as an overly colourful bird so I chose the slightly more refined Papuan lorikeet.

Unusually amongst the parrots, the Loriini are specialised nectarivores. The genus Trichoglossus refers to their hairy ('tricho') tongue ('glossus') which is covered in tiny papillae that hold on to the nectar as the tongue is withdrawn into the mouth. The lorikeets are joined by the pint-sized fig-parrots and the budgerigar of pet fame in the parrot subfamily Loriinae.


Day 50: Eclectus parror - Eclectus roratus

Who wears the bright trousers?

Given the sexual dimorphism and dichromatism we have encountered so far on this journey it may not surprise you that male eclectus parrots look different from females. What is surprising, however, is both the extent of the dichromatism and the fact that it is the female that stands out most, in scarlet and blue, compared to the predominantly green males. This disparity has confused biologists for centuries who even originally recognised males and females as separate species.

The appearance of eclectus parrots reflects a unique and complex interplay of ecology, reproduction and evolution. The main constraint for eclectus parrots in the rainforests of Australasia is nesting availability. They require hollow tree cavities to nest which are hard to come by. Females invest much of their time competing for nest holes leading to 'social selection'. Social selection is a term used by some biologist to encompass the evolutionary change resulting from the pressures of social interactions including, but not limited to, classical sexual selection. It is well understood in evolutionary theory that males that compete with each other for females can avoid damage by honestly signalling their health and ability with colours and ornaments. Well the reasoning goes that there will be situations where conspecifics compete for commodities other than mates which result in the same pressure to appear impressive. Thus, competing female eclectus parrots evolved their colouration in competition for nest holes.

In this situation, the sex roles have not been reversed (as they were for jacanas and phalaropes); males still compete for females yet male eclectus parrots look so...green. Well that green is perfect camouflage for the male as it forages amongst the foliage, dodging the keen eyes of falcons and owls.  However, in the arena around the nest hole, the green plumage suddenly stands out and the males are able to compete in the traditional fashion. Similarly the females plumage is effective camouflage when she dives into the safety of her cavity. Moreover, unbeknownst to the keen-eyed predators, these parrots are even more eclectic in ultraviolet light, part of the spectrum which parrots but not peregrines can see in.

Limited nest holes have further implications for the eclectus parrot. When females can only breed in cavities the limiting factor becomes, not females, but holes and sometimes there aren't enough to go around. As a result, females cannot afford to leave their hard-won homes so they desperately need food provisioning. She will, therefore, mate with multiple males (polyandry) and the uncertainty of paternity causes the potential fathers to all help out. This isn't the best situation for a male when holes, and thus mating opportunities, are rare and the offspring they're feeding might not even be their own! To maximise their reproductive success, males occupy huge territories which include multiple nesting holes. This complicated and unique (amongst birds) mating system is known as polygynandry: the only way to succeed is to cheat!


Sunday, 22 February 2015

Week 11: Sandgrouse to Fruit-doves

Day 43: Pin-tailed sandgrouse – Pterocles alchata

Pterocles Queen of the Desert

This is one of the most beautiful of the 16 species of sandgrouse which form their own order, the Pteroclidiformes. The name sandgrouse refers both to the generally dry habitat (and matching colour) of these birds as well as their superficial resemblance to grouse. Sandgrouse are roughly grouse like in shape and sexes are dimorphic with males bearing more ornate plumage. In the case of the pin-tailed sandgrouse the male is handsome in ochre and burnished gold with a blue eye-ring and a finely tapered tail.

One difference between sandgrouse and grouse, however, becomes obvious in flight. Sandgrouse have long, pointed wings which allow them to cover large distances at speed. This is necessary for life in the plains when water can be patchily distributed. This puts some constraint on parental provisioning, however sandgrouse have specially adapted belly feathers which enable parents to absorb water and carry it back to their chicks. Drinking at watering-holes must be brief as predators frequent such sites so sandgrouse maximise their chances by congregating in large numbers, usually only visiting at dawn or dusk.

The resemblance to grouse meant that sandgrouse were formerly considered members of the Galliformes. Now they are placed next to the pigeons although they might be more closely related to shorebirds.


Day 44: Crested pigeon – Ocyphaps lophotes

Respect the pigeon

Here we enter one of the largest non-passerine bird orders: the Columbiformes. This is the order of the pigeons and doves, two terms which mean little biologically but, generally, pigeons tend to be larger species whilst doves are smaller. Some people may have a derogatory opinion of pigeons but this is probably in part due to the most familiar species, the feral pigeon (Columba livia domestica) which derives from the cliff-living rock dove via domesticated populations. With over 300 species there is , however, more to the pigeons than the humble ‘flying-rat’!

The pigeons vary in size from the goose-sized crowned pigeons (Goura spp.) to the sparrow-sized New World ground-doves (Columbina). Although there is only one pigeon family (Columbidae) the pigeons and doves are split into around ten subfamilies including the typical pigeons (Columbinae) and the arboreal doves (Treroninae). Columbids are mostly granivorous however the latter subfamily specialise in fruit.

The crested pigeon is a small Australian pigeon of open habitats. It is notable for its erect crest which is surprisingly rare in the Columbidae given the importance of visual cues in courtship.


Day 45: Nicobar pigeon – Caloenas nicobaricai

You should meet his cousin

The Nicobar pigeon is a fantastic beast to behold. This large pigeon is covered in the most exquisite pointed, metallic feathers from bronze to blue. As well as the Nicobar islands the pigeon is found on small forested islands as far east as Palau. It is generally gregarious, roaming the forest floor in search of seeds and fruits. As well as their distinctive plumage the Nicobar pigeon has a notable vulturine look which gives an air of the prehistoric.

This brings me to its most important taxonomic relevance. The Nicobar pigeon is the closest living relative of the dodo. Everyone knows about the dodo, that charismatic species that met its demise at the hands of humans. The dodo was a large flightless bird that evolved in isolation from predators on the island of Mauritius. Like many flightless birds we have already come across, the dodo was at considerable risk when humans came to the island at the end of the 16th Century. Although hunting would have placed pressure on the birds the greater issue came from habitat destruction and nest predation by introduced pigs and macaques. With its loss, as well as the extinction of giant tortoises (Cylindraspis spp.), Mauritius lost both charismatic species and important ecosystem engineers, responsible for shaping the vegetation of the island. On nearby Rodrigues the swan-sized, flightless Rodrigues solitaire was another giant pigeon to become extinct because of man.


Day 46: Mariana fruit-dove – Ptilinopus roseicapilla

Jewel of the pigeons

The Mariana fruit-dove is a member of the largest genus of columbids comprising around 50 species distributed around the islands of Southeast Asia and Oceana. Fruit-doves generally occupy the same niche wherever they are, being arboreal frugivores. This raises an interesting question about what causes some groups to be so large. In some instances a clade will hit upon an adaptation which allows the group to specialise into many different niches facilitated by the new adaptation. Hummingbirds, for example, have capitalised on their hovering flight and long bills to diversify and this is generally seen as an adaptive radiation.

Examining the fruits-doves, however, reveals little niche diversification within the genus. One adaptation that precluded their diversity was the large gape which facilitates frugivory yet there is little further specialisation. Instead the clue is in thier short, rounded wings which are not evolved for strong, regular flight. When this is coupled with the distribution of fruit-doves over the islands of the Indo-Pacific it becomes apparent why there are so many species. After the origin of the genus (probably in Southeast Asia), sporadic events caused birds to hop islands and start populations in isolation. The lack of regular dispersal meant that gene flow was soon cut short and the populations diverged and speciated in allopatry. When this process is iterated across the Indo-Pacific, the species richness of Ptilinopuus becomes the result of a series of vicariant events.

It should be noted that fruit-doves are not identical despite ecological similarity. In fact they are the most colourful and aesthetically diverse group of pigeons, mostly green with patches of bright orange, purple or magenta. The greater similarity in appearance of some species to others tracks the evolutionary history of their divergence as they jumped from island to island. The Mariana fruit-dove is a typical Ptilinopus species endemic to the Northern Mariana Islands and Guam (both US territories). I chose this species because it looks cool. 


Monday, 9 February 2015

Week 10: Skimmers to Auks

Day 39: Black skimmer - Rynchops niger

Just skimming the surface

The black skimmer is one of three skimmer species in their own family: the Rynchopidae. Skimmers are yet another fantastic example of the avian beak as a highly adapted tool. In this instance, the lower mandible extends much further than the upper. The skimmer flies low over water and drags the lower mandible along the surface before hooking up an unsuspecting fish. Of the skimmers, the black skimmer is unique in being the only bird with slitted pupils. The cat-like vertical slits are thought to minimise the amount of light reaching the eye from staring so constantly at the water's surface at close proximity.

The skimmers are one of the firm member families of the charadriiform suborder Lari. I say firm as ,whilst taxonomists can agree on the inclusion of five families, others, such as the coursers (Glareolidae) and Egyptian plover (Pluvianidae), have debated inclusion. The remaining families in this order will be represented over the coming days but for the skuas: large, brown sea birds known for their kleptoparasitic bullying of other birds.


Day 40: Lesser black-backed gull - Larus fuscus

One ring to rule them all

As gulls go, the lesser black-backed gull does not stand out ecologically or aethetically (all though I think the smartness of gulls is under-appreciated). Larus is a genus of large, white-headed gulls in the gull family Laridae (other gull genera tend to have dark heads). The reason that the lesser black-backed gull is so interesting is because of its evolutionary relationship with the familiar seaside herring gull (Larus argentatus)

To us in Britain these two gulls are easy to tell apart with the lesser black-backed gull being darker with the bright yellow legs and the herring gull being pale-backed and pink-legged (the latter has also probably stolen your lunch). However they are part of a ring complex of species. As one moves east over Europe to Siberia the lesser black-backed gulls grow slightly paler becoming the Heuglin's gull (Larus heuglini). Continue through to far-east Siberia and Heuglin's gull is replaced by the grey-backed, pink-legged Vega gull (Larus vegae). Cross into North America and, paler still, we have the American herring gull (Larus smithsonianus). Finally, across the Pond and back in Britain the European herring gull is the dominant Larus species. 

Thus, when moving east, the differences between populations are slight and all species can hybridise with neighbours but by the time the circuit is complete the sympatric herring and lesser black-backed gulls are so different as to be obviously separate species. Ring species complexes like this are a nightmare for taxonomists. However they are a brilliant, if rare, demonstration of the fuzziness of evolution and how gradual changes can accrue almost imperceptibly over space and/or time.


Day 41: Inca tern - Larosterna inca

Movember

The Inca tern is a beautiful tern from the coasts of Peru and Chile. The bright-white moustache of the Inca tern gives it a unique look among the terns but otherwise its ecology is representative of the tern family Sternidae. The terns are able fliers with very long, tapered wings and tails. Flight is very important to the terns for two main reasons. Firstly, terns spend a great deal of the time on the wing, using their keen eyes to spot fish before plunging from a height to seize their prey in their sharp beaks. Secondly, many terns are migratory with the Arctic tern (Sterna paradisaea) travelling up to 91,000 km each year. Though I have not seen Inca terns in the wild they do form part of the unnatural bird community found in Living Coasts Zoo and Aquarium in Torquay, a fantastic attraction which gives close looks of many overlooked but fantastic coastal animals.


Day 42: Atlantic puffin - Fratercula arctica

Friar of the cliffs

Puffins! If you do not know what a puffin is then you live in a burrow (much like a puffin) and if you don't love puffins then you have a heart of stone. One of our most charismatic birds, the puffin is famous for its bright bill and pompous gait as it struts around the grassy knolls of rocky coastlines. The colourful beak that we know and love only looks like this during the breeding season when pairs return to land to breed. The bright colours are used to signal dominance whilst couples court by rattling their beaks together in 'billing' courtship displays. In winter the beak shrinks and becomes grey in colour as individuals live a largely solitary existence on the open ocean.

The word puffin derives from the Middle English 'poffin' which actually referred to the cured carcasses of Manx shearwaters (hence Puffinus puffinus) which were originally known as Manx puffins. The word later came to describe the puffins we know, perhaps because they nest in burrows as Manxies do. Otherwise they are very different, especially when it comes to locomotion. Whilst shearwaters can traverse huge distances with relative ease, puffins have very short wings which are well adapted for swimming but terrible for flight. Their swimming ability meant that puffins were thought to be part fish, part bird, a loop-hole which allowed Catholics to eat them on Fridays and during Lent!

The Atlantic puffin shares the genus with two other puffin species whose name Fratercula refers to their resemblance to little friars. The puffins are part of the auk family (Alcidae), a relatively small group of piscivorous sea birds. Cliffs in the UK can be populated with thousands of auks and other seabirds which form classist tower-blocks. Guillemots (Uria aalge) occupy the majority of the cliff face with the chunkier razorbills (Alca torda) cramming in above. The kittiwakes (Rissa tridactyla), small, gentile gulls, take the steepest part of the cliffs whilst shags (Phalacrocorax aristotilis) live nearest the waves. Above them all, the puffins share the cliff tops with tube-nosed fulmars (Fulmaris glacialis) and nocturnal Manxies. These seabird stacks show vertical niche stratification both on the cliffs and under the water where their diving abilities determine what prey different species can access. Of all the seabirds, auks have forsaken their flying ability in favour of increased diving ability with Uria guillemots reaching down to 100 m. Of these so-called 'northern penguins', one auk went so far as to lose flight altogether but this, as with many flightless birds, would be its downfall. The great auk (Pinguinus impennis) was hunted to extinction in the mid 19th century.