Saint Lucia

How the Island Got Its Birds 
Now in the hibernal depths, I cannot help but contemplate distant lands under a tropical sun. Last summer I was lucky enough to go to such a place, the Caribbean island of Saint Lucia. This rocky island thrusts from the sea, a volcanic testament to the grappling tectonic plates below. It is, of course, not alone but one of dozens of volcanic islands that form the Lesser Islands, set at the eastern end of the Caribbean. For me, islands in general are great holiday destinations because they are discrete units and thus ‘doable’ in the allotted time. Add to this tropical rainforests, mountains and coral reefs and you have a potent mix of exploratory fun. Yet as a birder, one could be disappointed. If long bird lists are your game then islands are not the best destinations as their complement of birds tend to be small. Saint Lucia has some 70 regular species compared to the hundreds in the forests of Central and Southern America. But why might this be?

Les Pitons

How natural communities assemble on islands has been debated for decades, ever since Robert MacArthur and E. O. Wilson explored island biogeography in the ‘60s. Early theories of island biogeography were based on the equilibrium balance between immigration and extinction on islands.  These islands were imagined to be blank slates to begin with, new pieces of rock, rising from the ocean and providing new opportunities for life. The inital limitation on island species richness is individuals getting there in the first place. The chance of immigration will depend on the degree of isolation from other land (particularly the mainland). Islands close to shore will accrue species much faster than more distant islands. The second limiting factor is the size of island. It is a general rule in ecology that the greater the area, the more species. Larger islands can have larger areas of habitat allowing larger populations and minimising extinction risk. Larger habitats also allow larger species to exist which require more sizeable territories. Larger islands also are more likely to have a variety of habitats. Complexity of shape, topography and geology can all increase with an island’s size and all of these factors favour habitat heterogeneity. The more complex the habitat, the greater the number of niches, the more species can coexist.

Purple-throated carib

Already we have multiple ideas that could shape the number of species on Saint Lucia and other islands. One thing Classic Island Biogeography does not account for, however, is the creation of new species. There are two possible routes by which islands can create new species. The first involves the immigration of a species which then evolves on the isolated island. This process of anagenesis is responsible for many species that are only found on (endemic to) one or a few islands. Saint Lucia has five endemic bird species: a parrot (Amazona versicolor), a warbler (Leiothlypis luciae), an oriole (Icterus laudabilis), a black finch (Melanospiza richardsoni) and a pewee (Contopus latirostris), all of which I managed to see. Some of these species have sister endemics on other islands and they represent a historical process of immigration and speciation as a lineage has spread throughout the archipelago. Taking Amazona parrots as an example, there is an endemic species on Hispaniola, Puerto Rico, Saint Lucia and Saint Vincent as well as two each on Jamaica and Dominica. There are many lineages like this, related species strung out over island chains and I remember encountering them during my Tring work. Generally these species would have similar niches wherever they were. Thus every island would have its own parakeet, its own fruit dove, its own white-eye or its own flightless rail. These lineages are radiations of geographic opportunity, not specialisation.

Saint Lucia warbler

This brings me to the other form of island speciation: cladogenesis. On islands of sufficient size or habitat differentiation, species can form from the reproductive isolation of different populations. This could happen under the classic model of allopatric speciation where species on different mountains, rivers or habitats speciate. On very large islands such as New Guinea, this form of speciation is not very different from that described about as species could share niches but remain geographically separated. On smaller islands, however, where coexistence is necessary, species must be sufficiently different in order to coexist. Cladogenesis is often unlikely when the immigration of new species is much faster than the evolution of new species in situ. Thus it is only the most isolated islands with the greatest ecological opportunity which produce within-island radiations. Hawai’i comes immediately to mind. The Hawaiian honeycreepers are an unusual lineage descended from South American finches which have diversified in form and function. Some species have short, thick bills and specialise in crushing seeds, others have longer, thinner bills, equally adapt at seizing insects and probing flowers for nectar. The most extreme bills are very long and curved, indicating high levels of specialisation. These birds represent a true adaptive radiation as different species have evolved to exploit different niches in lieu of the lack of competition. This would not happen of the islands were regularly flooded with hummingbirds, finches and warblers which fill the niches on the mainland. Alas, many species of honeycreeper are now extinct.

Mangrove cuckoo

The shadow or extinction is especially dark over the islands of the world. In blissful isolation, unique island fauna have evolved, becoming specialised to their insular ecosystems (often flightless and defenseless). They are not adept at dealing with sudden change. As humans have spread over the world they have caused the extinction of countless organisms. I have spoken in the past of hunters eliminating New Zealand moas yet more destructive has been the suite of introduced species which have had cataclysmic effects on island wildlife. Rats and pigs eat eggs, cats catch birds and mammals, goats strip vegetation and extinction ensues. We have lost many bird species on islands, from the charismatic dodo to numerous elusive and flightless rails. Now the largest threat of all, climate change, threatens what remains as the narrow tolerances of island species offer little leeway when temperatures change and climate events increase in frequency and severity.

Lesser Antillean bullfinch

So given this melee of theories and ideas, what can be made of Saint Lucia’s birds? The island’s habitats are broadly divided into a humid west side and a dry eastern side. Scrub-favouring birds scurry through the eastern brush and cling to the coastline of the west whilst in the montane areas, the humid rainforest produces a more three-dimensional habitat for a higher diversity of species. It is clear that representatives of many species have reached the island and yet there are curious gaps as well, such as the lack of owls, woodpeckers and tanagers, and these could simply reflect historical accident as they are often on adjacent islands. The Lesser Antilles are a chain of relatively close islands (I could see Saint Vincent off the southern edge of Saint Lucia) which means that many species will be able to pass along the archipelago uninhibited and there are species such as seabirds common to many islands of the West Indies and even the mainland.

Grey trembler

The few endemics necessarily represent sedentary species with low dispersal ability, species that got there by accident and didn’t move once they arrived. There is very little opportunity for cladogenesis on such an island where competitors can appear more easily from elsewhere so sister species from different islands tend to differ only in plumage. Finally anthropogenic extinction has probably played a small part in shaping Saint Lucia’s avifauna. The endemic Semper’s warbler (Leucospeza semperi) is likely extinct with the last reliable sighting in 1961. It probably suffered at the paws of the introduced Indian mongoose yet suitable habitat still exists and there remains the possibility of its persistence. Invasive species and loss of rainforest are, and will continue to be, an issue on Saint Lucia but thankfully I am unaware of the loss of any other bird species at present and the once endangered Saint Lucia parrot is on the increase.

Thus Saint Lucia may not have the longest bird list in the world but the island’s species tell an interesting tale of colonisation and survival and they include some beautiful and charismatic endemics.

Saint Lucia parrot

Honduras 2015

The Return to Cusuco

After measuring thousands of dead birds I was keen to see some live ones. So it was that I secured a place on the annual Operation Wallacea expedition to Honduras. The last time I had been to Honduras was in 2011 when I joined Operation Wallacea as a volunteer, spending two weeks in the cloud forests of Cusuco National Park and two weeks diving in the Bay Island of Utila. Back then, I did not know where my research interests would take me and I kept my options open, helping with the numerous surveys undertaken by OpWall. Little did I know I would return, four years later, as a member of the ornithology team.

Operation Wallacea is a conservation organisation working in 12 countries around the world. There, they conduct research into the biology and conservation of the natural systems in which they operate. The appeal of Honduras the first time around was the excellent combination of rainforest and reef but this time chance, nostalgia and the drive to go birding took me back. The location was Cusuco National Park, 23,440 hectares of protected are in the Merendon Mountains of northwest Honduras. This comprises 1/3 core protected zone and 2/3 buffer zone with eight camps distributed over an altitudinal range of 1,500 m. Part of the Meso-American biodiversity hotspot, the cloud forest is characterised by its high species richness and endemism making it the 48th most important protected area for biodiversity in the world. My poison is obviously birds, which Cusuco does not lack. With a park species list approaching 300 I was keen to set out and see come for myself.

In Cusuco there is a team for most major groups of organisms, responsible for the standardised, long-term monitoring of said group through their various means. Over time and space we hope to see how the anthropogenic impacts of climate change and habitat degradation affect this precious ecosystem. The large bird team split their time mainly between point counts and ringing. The former allows the detection of a large proportion of the avian diversity over larger areas whilst the latter provides more detailed, individual information on those species that can be caught close to the ground. Not yet being a ringer myself, I was only able to help with ringing for my first week under the close supervision of friend and colleague Sam Jones. The rest of the time I would be entrusted with the point counts.

Slate-throated whitestart

 Learning the Ropes

I was wont to tell people of the difficulty of point counts, mainly to big myself up, but also because it did entail a certain degree of knowledge. To do point counts I was required to be able to ID a good number of species from the off. In the tropics the vast number of bird detections are audial so, as with previous fieldwork in Australia, I needed to spend some time in advance of the trip listening to bird songs and calls. This, coupled with a tome of a bird book and some bedtime reading, meant that on arrival in Base Camp, I was not completely bewildered.

The Base Camp of Cusuco was a familiar site and I was happy to be back. My first morning there took me straight into the forest for my first point count. Point counts are simple in principle. After one minute of acclimatisation, the observer spends ten minutes recording all of the bird species detected. For each individual, the estimated distance and method of detection are recorded. It was reassuring to discover that some of the species had stuck in my mind so I could pick out the ubiquitous slate-coloured solitaires and black-headed nightingale thrushes against the cacophony but there was still a lot to learn.

With a steep learning curve came a steep slope, straight to the camp of Cantiles where I would be spending my first two weeks. I’m not going to lie, that three hour hike nearly killed me and for those of you who’ve done it, I know it wasn’t even that hard but it transpired I was surprisingly (or perhaps unsurprisingly) unfit and those who saw me stumble, red-faced into camp can testify. Thankfully this lack of fitness abated somewhat over my time in Cusuco.

Cantiles was a stunning camp, set on a steep hill among huge, epiphyte-ridden trees. With the highest altitude of any camp, Cantiles also boasts some tricky transects yet these winding routes reach up into the dwarf forest on the very highest peaks of the park. There, amidst stunted, moss-clad trees and mist-drenched bromeliads, I could watch the dawn come, hear the birds sing their chorus and see the wine-throated hummingbirds flit between the flowers (#birdon). However, I still hate mornings and getting up early was a grim necessity especially when it came to reverse transects. Rising at 3 AM to traipse for three hours through the dark, through the bugs, through the grasping, elfin forest was not my idea of fun but balance is important and the sampling of each transect requires surveying to be conducted in both directions, beginning at dawn.

The first week was something of a crash course. Under the esteemed tutelage of Sam, fellow birders Daniel, Adam and I spent that time getting to grips with the protocols and the birds, sometimes literally, where ringing was involved. Despite the knackering transects I was enjoying myself. The first week lent confidence to the second whereupon Adam and I were left in Cantiles to continue the survey effort. We were joined by the first wave of research assistants, an enthusiastic bunch who surprised me in their willingness for early starts. The devilish, mountain-top transects were tempered by long spells of opportunistic birding. These additional surveys are being tested alongside ringing and point counts for their efficacy in discovering the species richness of remote areas. They were also an excuse to sit back in a sunny spot and watch the birds fly by, especially rewarding when conducted in conjunction with a waterfall swim.

Wine-throated hummingbird

Heading West

Thus the second week flew by and it came to leaving time, onwards to the next camp: El Danto. A fellowship of scientists and students departed Cantiles with a daunting ‘8 hour’ trek ahead. I can’t complain much about the 6 ½ hour walk which was much easier than I expected but then I did make my life much simpler by taking the bare minimum to the West Side. The West Side were those few camps reached only by arduous hikes, a long way from the dry pine forests of Base Camp and the East Side. El Danto squatted in a short forest, shrouded in mist, clinging to the banks of a rocky river. Though I did not care for the constant damp or the sheer climb to the long-drops, I liked Danto for its forgiving transects. I was the sole birder at the camp and conducted my first point counts alone. As such, it was in this week that I shifted a gear; now comfortable with the common birds I set out to find more, to bolster my species list. This happened with some success as I found those species that had evaded me for a while (eye-ringed flatbill) and others which represented the first inclination of more disturbed, lower altitudes (masked tityra).

Disturbance was a defining feature of the West Side as the scars of man encroached inexorably on the supposedly protected forest. Whole transects have been lost to deforestation and this became more apparent as I moved on from El Danto. I began a five day journey to my next camp, Capuca. The first stop was Cortecito, a beautiful camp set deep in a valley of huge trees and colourful flowers. It was a shame to only see this camp for a night as it promised interesting birds and gave good views of foraging howler monkeys. A brief liaise with Sam also spurred me on to continue my search for more species.

From Cortecito, I descended further in the company of herpetologists Justin, Cheyenne and Alex. We ended our West Side experience in Santo Tomas, a tiny yet well furnished ‘camp’ in the buffer zone of the park. Santo Tomas was rife with nostalgia as I skirted the hill where once had stood a camp of real size which served as the main camp of the West Side. In 2011 I had spent my second week here taking in the local bird life but now the camp has all but disappeared. This is in part due to the extent of deforestation in the area rendering the camp impractical to run at full capacity. Now it operates only as an exit camp, releasing staff from the West Side. I knew the birding was good here so I had less than 24 hours to see what I could. My time was well spent and rewarded me with 32 species for my list including the stunning keel-billed toucan and blue-crowned motmot as well as a whole host of lowland ‘trash’ birds (Jones 2015).

Violet sabrewing nest

Back to the East

A long drive and a sumptuous Burger King took us back to Base Camp. One night there gave me time to check out some local birds (white-breasted hawk nest) before heading to Capuca. Capuca, Capuca, Capuca. Rain, mud and illness defined the first half of the week and no amount of tropical skittles or bean-themed card games could lift my cynicism. However, as always, Cusuco provided good company and top chat so that the week was still enjoyable. Moreover it was my first week of school groups which meant ‘inspiring the next generation’. I tried my best to sell birds to the students and succeeded to some small degree but regardless it was good to have them around and see them power through the mud and D & V to emerge enthusiastic and triumphant. The birding that week was a shade less exhilarating than previous weeks but still offered ‘billboard’ views of a resplendent quetzal.

And so I came to my last week. Habitat bloke Rik and I were ushered through Base Camp before a truck took us onwards to Buenos Aires (BA). You do not know luxury until you go to BA! They have plastic, flushing things called toilets, a dense springy layer called a mattress and all the junk food you could want. I loved BA from start to finish. Not only was the living standard higher and the company excellent but the birding was fresh and new too. Very few of the core zone species seep down to the deforested area surrounding the village; instead a suite of more generalist birds took their place. The interesting transects took me through a range of habitats enabling me to increase my list further, despite the protestations of fatigued students. Once again the week passed quickly, a blur of early starts, phallic frozen bananas and various medicines. The end came with one last dawn, one last opportunity to see the sun rise pink above the cloud forest. I joined a group heading down the mountain, comfortable in a seat rather than the bone-breaking bed of a pick-up, whereupon I lounged in the melancholy of leaving and the reverie of what had passed.

I didn't ‘find myself’ or anything but I did confirm that I was doing what I wanted to do: seeing amazing places filled with stunning birds (I saw 108 spp. in the end) in the company of enthusiastic, like-minded and brilliant people. Maybe I will go back...

Resplendent quetzal

Buenos Aires and environs

Week 21: Endangered Species and The End!

This is the last of my instalments based on Tring’s non-passerine birds. These are some of the rarest bird species in the world and there are many commonalities to their endangerment but conservationists are working tirelessly to bring them back from the brink and success is increasing.

Nene – Branta sandvicensis

I am quite familiar with the nene as a small, friendly goose who doesn’t bite your hand like those pesky greylags. I refer to childhood visits to WWT Slimbridge where my sister and I would spend hours feeding grain to the cosmopolitan waterfowl assemblages, inevitably running out of seed half way around the park.

The nēnē, or Hawaiian goose, is common in Slimbridge but on its native Hawaiian islands it became very rare. Up until colonial settlement of the islands, nene were relatively common, the equivalent of Canada geese in North America, but man was responsible for a severe decline via hunting and predation by introduced mongoose, pigs and cats. From Captain James Cook’s arrival in 1778 the population decreased from c. 25,000 to just 30 in 1952. Thankfully, in stepped Sir Peter Scott of WWT fame who spearheaded a captive breeding program to save the nene. Luckily these geese breed well in captivity and so the program and introductions have been successful with the population up to 2,500. Next time I feed a nene I will think of their story and remember the important role we must play in conserving our birds.   

Christmas frigatebird – Fregata andrewsi

The Christmas frigatebird is the rarest of the five frigatebird species in the family Fregatidae and it only breeds on Christmas Island in the Indian Ocean. In the grander scheme of things, frigatebirds are most closely related to cormorants (Phalacrocoracidae) and boobies (Sulidae) and sit in the order Suliformes which has been separated from its pelecaniform cousins.

I always get a sense of awe when I see a frigatebird, not because it’s especially big, impressive or beautiful but because they have a pterosaurian appearance, their angular black wings and tapering tail hanging motionless on the wind. The appearance of ease which they give to flight is not merely an illusion but a reflection of their super-pneumatic bones which comprise just 5% of their weight. This, coupled with the largest wingspan-to-body-weight ratio, means that frigatebirds can spend ages floating on the air, flapping infrequently and expending little energy. Like swifts, they can even over-night on the wing.

Frigatebirds nest in large colonies where males vie for females using huge, inflated gular sacs which are bright red in colour. From above, the females encircle groups of males who point skywards to reveal their pouches, vibrating their bills to produce a drumming sound. Regular readers will not be surprised to see the presence of sexual selection again, especially the colour red which is usually a sign of a healthy diet rich in carotenoids. Furthermore, frigatebirds are unusual amongst seabirds for having blood-borne parasites. Parasites are important in handicap theory whereby only males with the best immune systems can stave off infection and produce their bright colouration.

Parasitised though they may be, frigatebirds are famous for turning parasite themselves. Kleptoparasitism is the act of stealing food from others and frigatebirds are consummate bullies who terrorise the smaller boobies, tropicbirds and gulls, forcing them to give up their catch or the contents of their stomachs. Perhaps piratebird would be a more fitting moniker.

Philippine eagle – Pithecophaga jefferyi

Another critically endangered bird, the Philippine eagle is the largest eagle in terms of length and wing area. It is endemic to the Philippines where deforestation has drastically limited its range and population. It evolved to be the apex predator with little competition on the islands and has such it has grown to a huge size with a fearsome set of talons and a monstrous beak which can make short work of their prey. Their prey is pretty much any terrestrial vertebrate short of humans or larger ungulates. They get the name Pithecophaga for their taste in monkeys (the Philippine long-tailed macaque) but flying foxes, squirrels, giant cloud rats, civets, snakes, lizards, owls, hornbills and even deer also make up this eagles diet. We are used to images of eagles soaring overhead but Philippine eagles are jungle denizens, appearing out of nowhere to seize their prey from branches. For such a large bird they are surprisingly stealthy.

Saint Lucia Parrot – Amazona versicolor

The Amazona parrots are a speciose genus and members in mainland Central and South America can be widespread and common however the few species strung out across the Caribbean tend to be rare. Amazons have low dispersal ability so they are often range restricted and on the islands of the West Indies each species tends to be endemic to just one or two islands. The Caribbean islands have seen a lot of development and habitat loss and this has pushed many endemics closer to extinction and the Saint Lucia parrot is no exception. It declined from around 1,000 individuals to just 150 birds towards the end of the 1970’s. Since then, and with great public support, this parrot has bounced back.

Of these various island-endemic Amazons, I chose to write about the Saint Lucia parrot as I hope to see it in the wild this summer. I am going to Saint Lucia for two weeks and I hope to take in as much of the islands natural history as possible. Saint Lucia has at least four endemic bird species including a warbler (Setophaga delicata), an oriole (Icterus laudabilis) and a black finch (Melanospiza richardsoni) as well as rare reptiles, stunning rainforest and rich coral reef. I cannot wait!

Keel-billed motmot – Electron carinatum

Motmots are stunning birds characterised by their bright colours and long tails which often end in racket-like tips. Unsurprisingly their tails play a role in courtship but one intriguing use seems to be as an anti-pursuit deterrent. According to studies conducted on turquoise-browed motmots (Eumomota superciliosa), motmots wag their tails when they notice a potential predator. The signal says ‘I have seen you and I will fly away so don’t even bother’. This signal benefits both parties as neither wastes its energy in a fruitless pursuit.

During my time in Honduras in 2011 we caught a blue-crowned motmot (Momotus momota) in a mist net and one thing that stood out was the impressive bill which is used to grasp lizards and insects. To catch their prey motmots perch silently then pounce from above. Confined to the New World, the Momotidae occupy a niche similar to the Old World forest kingfishers (Halcyonidae) which share their order. In June I return to Honduras and will hopefully see more of these beautiful birds and maybe even the keel-billed motmot itself.

Guam rail – Gallirallus owstoni

The genus Gallirallus caused some issue when first we passed it in our avian tour as few of the species seemed to be in the collection. Resolution of this issue also lead to multiple explanations for their initial absence. The first was that many of the rail species had only recently joined this genus having traditionally been placed in other genera such as Rallus. Another explanation was the fact that this genus comprises many extremely rare species which had been placed safely in more secure collections.

Ancestors of the Gallirallus rails managed to populate many of the islands across Australasia and the Pacific whereupon they became flightless in the absence of terrestrial predators. Latter introductions of mammals such as cats and rats have put many rail species at risk and even eliminated some. The Tongatapu rail (G. hypoleucus), Dieffenbach’s rail (G. dieffenbachii), Wake Island rail (G. wakensis) and Tahiti rail (G. pacificus) have all gone extinct since the late 18^th century with the New Caledonian rail (G. lafresnayanus) probably extinct. The Okinawa rail (G. okinawae) and Calayan rail (G. calayanus) were only discovered in 1981 and 2004, respectively, demonstrating the cryptic nature of these species.

The Guam rail is a typical member of the genus being a flightless, island endemic. However it is extinct in the wild having been extirpated from Guam in the 1980’s as a result of the introduced brown tree snake. It exists only in captivity where a breeding program hopes to establish a future population. It is a shame that this American territory should lose this species as well as other endemics but the might of American conservation is now behind the Guam rail. 17 zoos are participating in a captive breeding program with around 120 individuals alive in captivity. Eradication of rats and snakes is the key to the success of reintroduction programs such as this and the reintroductions of Guam rails to a 22 hectare enclosure and an isolated atoll have proved successful so there is hope for the future of this species.

Eagle eyed observers might have noticed that the Guam rail is taxonomically out of sync with the rest of the species this week. That is because it was the last species that I measured in Tring. After exhaustively searching for every last species the museum had to offer, chasing up loose ends and find the most elusive specimens, I finished the job after six months measuring. Together, with my friends and colleagues Nico and Sam, we measured 3,046 species from 11,490 skins taking 103,410 measurements. It was a staggering effort, an intellectual journey and thoroughly enjoyable experience. My blog was a vehicle for exploring some of the amazing birds covered in that period (all 90 pictured below!) so do go back and explore my past posts yourself. As for the future, adventure beckons with trips to Honduras and Saint Lucia followed, hopefully, by further avian experiences and I’ll be birding all the way!

Week 20: Large Birds Part II

Marabou stork – Leptoptilos crumenifer

The marabou stork is not the most attractive bird and it would make a terrible dinner guest as carrion forms a large part of its diet. Marabous will follow vultures to carcasses, allowing them to do the difficult work of ripping open the skin, before plunging their own huge bills into the flesh. As with vultures, the lack of facial feathers prevents blood and gore from building up around the head. In addition to scavenging, marabous are generally opportunistic, taking small vertebrates, eggs and invertebrates. This generalism has facilitated an increase in the use of rubbish dumps as sources of food in the burgeoning urban areas of Africa.

The marabou stork is a member of the stork family, Ciconiidae, and order, Ciconiiformes. This is small order of about 19 species which used to include ibises and spoonbills until their move to the Pelecaniformes.

Australian pelican – Pelecanus conspicillatus

Speaking of the Pelecaniformes, the Australian pelican is a handsome example of the eight species of pelican. It is notable for having the longest beak of any living bird! It was also one of the first species I spotted on my trip to Queensland. Unfortunately I did not see it eat. The foraging ecology of pelicans is famous (or perhaps infamous) for the utilisation of the huge gular pouch to catch fish. Whilst small and large fish form the majority of the diet, pelicans will also eat amphibians and crustaceans and some observations from Regent’s Park even show them to eat pigeons! Pelicans can also use the skin of their pouches to regulate their temperature, fluttering the skin to promote cooling in the same way that elephants flap their ears.

The habitat of pelicans is generally, and unsurprisingly, based particularly around wetlands (although I have seen brown pelicans [Pelecanus occidentalis] off the beaches of Miami and the Caribbean). There they join a wider assemblage of pelecaniform species. The Pelecaniformes have evolved various ways of plying the waterways for food with much emphasis on beak morphology. The statuesque herons come in a range of sizes but most are expert anglers. The ibises (or, more correctly, ibides) take a more relaxed view, probing the substrate for invertebrates whilst the spoonbills sweep the water with their huge utensils. Perhaps the oddest members of the order are the hamerkop (Scopus umbretta) and shoebill (Balaeniceps rex), two monotypic species related to pelicans. The former is a small, hammer-headed wading bird which build huge nests whilst the latter is a large, stork-like bird with an even larger, whale shaped bill (hence the Latin 'king whale-head'). The shoebill also has a dastardly stare which could strike fear into anyone.

Andean condor – Vultur gryphus

The Andean condor means playing the superlative game again. I had thought that this condor had the second largest wingspan of any living bird but apparently the two largest pelicans are also bigger. Moreover, the splitting of the wandering albatross species complex into many species throws a few more, longer winged species into the mix. The Andean condor does, however, have the largest wing area of any living bird species which is not surprising as the barn doors of the condor are far wider than the tapered wings of the albatrosses.

As the name suggests, the Andean condor hails from the Andes mountain range which forms the spine of South America. Without a sternum, the condor is not designed for flapping, instead it relies heavily on thermals to keep it aloft, keeping to high places and using the topography of the land to soar about. Seeing these birds in their element must surely be an amazing sight and its unsurprising that the ancient Native Americans worshipped them as sun deities. Their legendary appearance lives on in the coats of arms of no fewer than five South American countries.

Soaring around, looking for carrion is what traditionally makes the Andean condor a vulture, one of the seven species in the New World vulture family Cathartidae. In the second half of the 20^th century, some ornithologists began to realise that the similarity between the Old and New World vultures might only be skin deep. Morphological and karyotypic (number and shape of chromosomes) data were used to support the grouping of New World vultures with the storks with the apparent similarity with Old World vultures resulting from convergent evolution. Whilst this convergence in morphology and ecology still stands, the evolutionary relationships have been better resolved with the placement of New World vultures as a sister clade to the rest of the birds of prey (excluding falcons), either sharing the order Accipitriformes or being elevated to their own order Cathartiformes.   

Lammergeier – Gypaetus barbatus

The lammergeier (or bearded vulture) is a true vulture in the sense of being a member of the Aegypiinae Old World vultures however it is atypical in its appearance and ecology. It has orangey-yellow head plumage to match its torso and the face is completed with a stiff, black beard (barbatus = ‘bearded’). Although lammergeiers are scavengers, their diet is unique for being based largely on bone marrow. Obviously bone marrow can be pretty difficult to access. For smaller bones, lammergeiers can crack the bone open but for bigger bones, they have a rather clever technique. Lammergeiers have learnt to carry bones up to 150 m into the air, dropping them from a height so that they fall and crack open on the ground. Adult birds can carry bones weighing nearly as much as they do! Sometimes this method is even extended to crack open tortoises. Bone cracking is a classic example of a learned technique and immature birds can spend years honing their skills. Once ingested the bones need industrial strength acid dissolve. As such, lammergeiers have stomach acid at pH 1 which dissolves the calcium rich bone in a day.

Lammergeiers cling to high altitude regions across Southern Europe, Central Asia and Africa. The mountainous habitat is a harsh environment but living off bones means that food can often be found. Furthermore, when carcasses are all but stripped of flesh, the marrow can live on, desiccated within the bones.

Great bustard – Otis tarda

The great bustard is generally cited as the heaviest flying bird, weighing up to 18 kg. It is also, arguably, the most sexually dimorphic bird species in terms of size as males can weigh up to four times the weight of females. Great bustards have a patchy distribution across the grassland and steppes of Iberia, Central Europe and Asia. They were once part of the British countryside too but they were hunted to extinction by the 1840s. Now the Great Bustard Project is working to establish a new breeding population in the rolling fields of Wessex. Since 2004, birds reared from Russian and then Spanish eggs have been introduced annually. Bustards bread for the first time in nearly 200 years in 2009 and the population is slowly growing.

In the UK, great bustards have much in common with the common crane as both were almost absent from our countryside and now programmes are tirelessly working to bring them back with the Great Crane Project experiencing much success. Evidence of these two species in our heritage is subtle but present. Most noticeably, bustards, cranes and other species are found in the heraldic emblems of the past. Bustards are found on the coats of arms of Wiltshire and Cambridgeshire. Peregrine falcons are found on the arms of Surrey, Bagshot and the University of Lincoln whilst Lincolnshire’s is capped by a lapwing. Northamptonshire even has a white falcon that looks a bit like a gyrfalcon (unlikely that gyrs have ever visited Northants). Finally, in Cornwall, red-billed choughs used to be a common sight, flying on broad wings along the cliffs, and as such they grace the coat of arms of Cornwall. Another word for chough is beckit and the 12^th Century Bishop Thomas Beckit had three choughs on his coat of arms, as did the seat of his power, Canterbury. During the reign of the Tudors, Cardinal Thomas Wolsey took the choughs of his namesake for his own crest and bestowed that crest on his new college of Christchurch in Oxford.

Red-crowned crane – Grus japonensis

Cranes are members of the Gruidae family which sits in the Gruiformes order with the rails, crakes and gallinules. This relationship might seem surprising but when measuring both the rails and cranes I could see the similarity in bill morphology. Cranes are essentially rails on steroids. 

The red-crowned crane is one of the largest and (in my opinion) most beautiful of the 15 crane species. Cranes are characterised by the long legs and necks which are stretched out in flight. They have a generally omnivorous diet, although a degree of niche segregation can occur when multiple crane species live in sympatry. China is the hotspot of crane diversity, containing, at various points, eight species. Many are migratory so may either only winter or breed in China. The red-crowned crane exhibits different migration strategies between the two disjunct populations. In Japan, the cranes barely move at all, hopping just 150 km whilst the mainland race breeds in Northern China and Siberia then splits up and migrates around 1,500 km to Korea and Jiangsu, China.

The red-crowned crane is one of the most endangered crane species. It is possible that the risk to cranes is somehow linked to diet as it has been noted that the most common species are largely herbivorous within the scale of omnivory whilst the red-crowned and (rarer) whooping cranes feature more animals in their diet including invertebrates, fish, amphibians, reptiles and birds. A carnivorous diet results in a degree of dispersal whilst foraging as opposed to the gregarious nature of more herbivorous cranes. Red-crowned cranes do, however, breed in groups, taking advantage of safety in numbers. Cranes are typically monogamous, keeping the same partner for many years. Couples maintain their bonds via the rhythmic, synchronised dancing as well as duetting with loud, trumpeting calls.

The red-crowned crane has important symbolism for both the Chinese and the Japanese. In China, and particularly Taoism, the crane symbolises longevity and immortality. In Japan, red-crowned cranes were said to live for 1,000 years whilst they are also recognised for their loyalty in monogamy and their strength in migration. I think they are impressive birds and should rightly be respected and cherished.

Week 19: Large Birds Part I

Now for some of the species that were too large for the normal collection and they are large in every sense of the word. Beware there will be a lot of superlatives!

Ostrich – Struthio camelus

Behold, the largest species of bird in the world! Well that was obviously no surprise but even though ostriches are well known they are still very impressive beasts. Males can stand up to 2.8 m high and weigh in at 145 kg plus they also hold the land-speed record for birds, running at speeds of 70 km/h. Ostriches also lay the largest eggs of any birds, 20 times the weight of a chicken's, yet the eggs are the smallest in relation to body size of any bird.

The two ostriches (S. camelus and the Somali ostrich S. molybdophanes) are ratites, a term generally reserved for flightless birds of the order Struthioniformes including emus, rheas, cassowaries and kiwis. With the flux of modern taxonomy, the validity of this clade is in question as the volant tinamous should be nested within the flightless birds suggesting independent routes to flightlessness. As well as the extant ratites, there were larger and more bizarre members of the group. On New Zealand the moas reigned supreme until the relatively recent immigration of Mouris whilst on Madagascar, the huge elephant birds stalked the forest.

Together the ratites have a Gondwanan distribution. This refers to a geographical distribution of organisms contingent on the past arrangement of continents. During the evolution of birds in the Cretaceous, ancestors of the ratites were distributed over the southern continent of Gondwana (South America, Africa, India, Australasia and Antarctica). Once Gondwana began to break up during this period, each new continent took its own ratites. Emus, cassowaries, kiwis and moas evolved in Australasia, South America evolved rheas and Africa took the ostriches. Well at least that was the traditional view; more recent genetic work places the separation of these lineages as more recent than the continental splits. This implies that volant ancestors colonised the different continents and independently became flightless, probably in lieu of the lack of large terrestrial vertebrates following the extinction of the dinosaurs.

Southern Cassowary – Casuarius casuarius

Speaking of dinosaurs, the southern cassowary could be lifted straight from the Cretaceous! Of the three Australasian cassowaries, the southern cassowary is the largest and its bright blue neck with pink wattles stands out against the black, ‘furry’ body. It is the species of most relevance to me as I was lucky enough to see several individuals of this vulnerable, enigmatic species in the wild whilst undertaking fieldwork in the rainforests of Far North Queensland. One encounter stands out in particular.

My friend, Emily, and I were stepping along one of our paths into the rainforest when Emily froze and back-tracked suddenly, cursing as she went! I then spotted the issue when I noticed a cassowary standing about 15 m ahead. My first thought was ‘photo opportunity’, so I whipped out my phone and took terrible snaps. But then it turned to face us. We did nothing at first until it started walking towards us. Suddenly, through my mind flashed the stories of people being head-butted with their huge casques or lacerated with their elongated claws. We started walking back down the track but the cassowary began to pick up the pace. The more we sped up, the more it did, until soon we were running as fast as we could which was difficult due to the suddenly claustrophobic foliage and lianas. Finally (after what was probably just seconds) we emerged into the sun light and the cassowary did not follow. Our hearts thumping, we regained our composure but the fear of that encounter remained for the duration of our work!

Green peafowl – Pavo muticus

I have to admit I did not know what this species looked like beforehand but being a congener of the familiar peacock (Indian peafowl, Pavo cristatus) I surmised correctly that it would be impressive. Moreover, the unfamiliarity gives it an exotic feel where we might take peacocks for granted. This denizen of Southeast Asian forests shares many of the traits that make the peacock famous including the huge ‘tail’ of many eyes which is actually formed from upper tail coverts rather than the rectrices. The tail of the peacock is famous in biology as the pin-up for sexual selection. That such an elaborate ornament should evolve under natural selection troubled Charles Darwin greatly who said that ‘the sight of a feather in a peacock’s tail, whenever I gaze at it, makes me sick!’ In time, Darwin, and the generations of biologists to follow, have expounded theories of sexual selection which explain the ornaments of peacocks and other animals. Whether choosing ornamented males for their sexy sons are their good genes, it seems that beauty is the way to go.

Emperor penguin – Aptenodytes forsteri

Of the 19 penguin species, the emperor is the largest, as the name might suggest. It is also the only bird species to breed in the Antarctic winter. The emperor penguins are famous for their annual 100 km trek into the Antarctic desert whereupon they breed, the female laying a single egg. The first duty of the male is incubation so that the female might return to the sea to breed. The egg is passed carefully between the feet of the male to the female, avoiding the lethally cold ground at all costs. With the female departed the male must while away the winter months, burning precious fat reserves until the female returns. During this time, huddling is imperative to conserve heat as are the suite of hypothermic adaptations including blubber, down and the most densely packed feathers of any bird which can be erected to trap additional air.

Soon after the chick hatches it is fed its first protein rich regurgitation.Then, the female returns with her own supply, locating the male through vocal recognition and relieving him of his duties so that he can travel back to the sea. Thereafter, rearing the chick is taken in turn. The free parent must run a gauntlet of leopard seals and orcas to reach open water whereupon they may fly through the water, diving up to 500 m deep, feasting upon Antarctic fish, squid and krill. 

A month and a half into life and the chicks are big enough to be left alone whilst the parent both feed at sea. This is not really ‘alone’, however, as the chicks form large crèches, avoiding the bitter wind and the predatory Southern giant petrels. As summer approaches, the chicks begin to moult into juvenile plumage and make to join their parents in the booming summer seas which, by this point, are considerably closer owing to the melting sea ice.

 Wandering albatross – Diomedia exulans

The wandering albatross has the largest wingspan of any living bird at up to 3.5 m (11 ft 6 in) which is slightly absurd if you think about it. Most of their life is spent at sea with monogamous adults only returning to land to dance and breed on rocky islands in the Southern Ocean. They are at the slowest end of the life-history continuum. The life-history continuum represents the trade-off that all organisms have to face: whether to live longer or reproduce more. Large organisms tend to be at the slow end and wandering albatrosses can live beyond 50 and reach sexual maturity in their teens, thereafter only rearing one chick every two years. 

Whilst the generic name Diomedia references the Ancient Greek hero of the Trojan War. Diomedes, who’s companions transformed into birds, the specific epithet exulans means ‘exile’ or ‘wanderer’ noting the vast areas this albatross covers. One impressive individual was observed to travel 6,000 km in just 12 days.

The albatross is famous for its role in the Rime of the Ancient Mariner, the epic poem by Samuel Taylor Coleridge, where the ‘bird of good omen’ was a welcome companion to the sailors and where the killing of one is a terrible crime resulting in the burden of an albatross around the neck. Today, it is humans that are the harbingers of doom for the albatrosses as plastic pollution threatens colonies around the world.

Week 18: Woodpeckers

Day 71: White-browed piculet – Sasia ochracea

Bamboo bambino

This pretty little bird is a close relative of the woodpeckers, being in the same family Picidae but the piculet subfamily Picuminae. As, essentially, miniature woodpeckers, the piculets (literally ‘small woodpecker’) have many of the adaptations of their larger cousins which I will discuss later. One thing all woodpeckers do is nest in holes which they drill into trees. In the case of the white-browed piculet, it is not trees that house the nests but bamboo stems. They prove that pandas are not the only bamboo specialists although this piculet is not found in Southwest Asia rather than China. 

Day 72: White-naped woodpecker – Chrysocolaptes festivus

Pack a hard hat

The white-naped woodpecker is a member of the vibrant flamebacks (Chrysocolaptes spp.) As well as the gold and orange colouration that characterises this genus, the male white-naped woodpecker also sports a splendid, crimson crest. The red crest is actually common to many woodpeckers species including the pileated woodpecker which probably inspired the cartoon Woody Woodpecker. In fact the red in the male’s crest is something I want to pick up on as, across the woodpeckers, I found that the amount of the red on the head was diagnostic of the sex of the individuals. I accept that there are exceptions but, by-and-large, all that was required to sex a specimen was to check the extent of red on the head with males generally having more. Although the dimorphism of pheasants, ducks etc was obvious, it varied greatly from species to species yet with woodpeckers I do not think I have encountered such consistent dichromatism.

The white-naped woodpecker is a very typical woodpecker and thus exhibits the adaptations that define the clade. Woodpeckers obviously peck wood. This is generally an activity undertaken to procure insects hidden beneath the bark of trees but woodpeckers also use their chisel-like bills to excavate holes and to drum in spring courtship. Drumming is a physically challenging act with woodpeckers experiencing deceleration rates of 1,000 g! To cope with this, they have relatively small brains orientated within their reinforced skulls in such a way as to maximise contact (minimising pressure) and minimise contact time. The strenuous activity of pecking up to 12,000 times a day is supported by strong neck muscles, gripping, zygodactylic feet and a thick, rigid tail which braces the whole bird. Additionally woodpeckers protect themselves from flying debris with feathered, closable nostrils and translucent nictitating membranes which flick across the eyes.

Once suitable excavations are made into bark, woodpeckers insert their extremely long, barbed tongues into crevices to wheedle out ants and larvae. Extending up to three times the length of the bill, these incredible tongues are retracted back into the head, coiling up behind the brain where they aid the cushioning tissue encasing the brain. Woodpeckers are biological machines.

Day 73: Lesser spotted woodpecker – Dendrocopos minor

The wish list

Lesser spotted woodpeckers are one of four woodpecker species found in the UK, the others being the enigmatic wryneck (Jynx torquilla), the ubiquitous great spotted woodpecker (Dendocopos major) and the laughing, ant-eating, weasel-sporting green woodpecker (Picus viridis). The lessser spotted woodpecker is of significance for its placement on two special lists. Firstly, it is the last species that I shall discuss from the main collection at Tring. After five months in the Natural History Musuem’s bird collection we have reached the bottom of our list of non-passerines. However, this is not quite the end as we have yet to measure the very large bird species (ratites, storks, cranes etc) housed in the Large Collection as well as the endangered and extinct species and species known only from type specimens. These remaining species will be discussed next week but for now, the majority of the job is done!

The second list upon which the lesser spotted woodpecker finds itself is my wish list. As a birder (and part-time twitcher) I keep a list of all the bird species I have observed in the UK (224 and growing). Whilst recent additions have been rare birds twitched in the West Country (black-winged stilt, penduline tits and Bonaparte’s gull) I am inclined to spend some time finding the less rare but less twitchable birds of this country. The sparrow-sized lesser spotted woodpecker is far less common than its larger counterpart. Pairs keep large territories and signal to each other with high-pitched, staccato drumming a noise which, coupled with calling, can give them away. Otherwise, their unobtrusive habits and sparse distribution make them difficult to see and the closest I have come is hearing a call or two in the New Forest. The season for drumming is dwindling so I may have to wait until next year to see one but hope to discover for myself this species, and others, in the years to come.

And thus, whilst one ornithological adventure draws to an end, the lesser spotted woodpecker is one of many I hope to come.

Week 17: Bee-eaters to Honeyguides

Day 67: Rainbow bee-eater – Merops ornatus

No sting in the tail

Bee-eaters are a charismatic group of birds in the family Meropidae. They are conspicuous birds, known for their bright and varied palette of plumage colouration. They are also conspicuous in their behaviour, frequenting prominent perches such as telephone wires from which they can spy prey up to 50 m away. They are capable of catching all manner of insects on the wing but, as the names suggests, bees can form a large part of their diet. Large and/or venomous prey is taken back to the perch where it is neutralised with bashing and scraping. Aerial insects abound so foraging opportunities are plentiful. Bee-eaters are limited more by nesting opportunities, requiring dry substrates in which to build their burrows. They have a largely tropical and subtropical distribution across the Old World with many species migrating annually.

The rainbow bee-eater is a typical bee-eater species. It is, however, the only species I have seen (despite multiple trips to the Med where European bee-eaters can be found). Rainbow bee-eaters were a common sight for me in the open areas of the Daintree Rainforest in Queensland. They are by no means confined to this habitat, perching anywhere where flying insects are in abundance including parks and gardens. I was observing them during the winter/dry months after they have spent the summer breeding in Southern Australia. They are a fun bird to watch and their bubbly calls are a delight.

The Meropidae, along with the three kingfisher families, are members of the order Coraciiformes. Coraciiforms lack a unifying name despite being generally colourful, small-to-medium-sized predators. The order includes the acrobatic, Old World rollers (Coraciidae), the stockier, Malagasy ground-rollers (Brachypteraciidae), the Neotropical, long-tailed motmots (Momotidae) and the tiny Caribbean todies (Todidae). Formerly a member of this order, the cuckoo roller (Leptosomus discolour) is an obscure bird from Madagascar and the Comoro islands now placed in its own order: Leptosomiformes.

Day 68: Southern ground hornbill – Bucorvus leadbeateri

Luscious lashes

This looker is the largest species of hornbill. Hornbills are notable for their large, curved bills which resemble cows’ horns (buceros in Greek). Their bills, supported by their uniquely-fused first two vertebrae, serve a variety of purposes including nest construction, fighting and preening but their most obvious use if for crushing food such as fruit and small animals. Many species have a hollow casque atop the bill which can be used to resonate calls or to joust with competitors (in the helmted hornbill, Rhinoplax vigil). Whilst their plumage is usually dark, the bills and naked facial skin of hornbills is often brightly coloured with yellows and reds that reflect the carotenoids of their diet. Once again, this is classically honest signalling of health for mate choice. 

Hornbills are largely arboreal, found in habitats from savannah to rainforest across Africa, southern Asia and Melanesia. The Southern hornbill and its sister species (the Abyssinian ground hornbill, B. abyssinicus) are a bit of an exception amongst the hornbills, being far more terrestrial than their relatives. In fact they might forsake the trees altogether were in not for the tree cavities in which they need to nest. The two ground hornbills can be found striding across the plains of sub-Saharan Africa in search of small vertebrates and large invertebrates. To cope with the dusty habitats in which they forage, the ground hornbills have very long lashes and bare skin around the face and throat. In the Southern ground hornbill, this skin is bright red.

Ground hornbills are some of the most (if not the most) long-lived birds with life expectancies around 30 years and with individuals living up to 70 years in captivity. They are also obligate cooperative breeders. Individuals must spend half-a-dozen years assisting at the nests of older birds (parents) before becoming breeders themselves. Cooperative breeding always necessitates explanation. Usually in birds it is a bet-hedging strategy; if it is too risky to start your own family (i.e. low chance of success) better to stay at home and help rear your younger siblings. Perhaps the risk in starting a new family for hornbills is either not gaining a suitable territory (or nest site) or not being able to bear the costs of rearing large chicks without help.

Ground hornbills share the family Bucorvidae with the Bycanistes genus of hornbills. The rest of the hornbills, from the small, African Tockus species to the large, casqued Buceros species, are found in the family Bucerotidae. Hornbills traditionally had the order Bucerotiformes to themselves but recently they have been joined by hoopoes (Upupidae) and wood-hoopoes (Phoeniculidae). Hoopoes (Upupa epops) are a fantastic bird with broad, barred wings and a salmon pink head and crest. Their name is an onomatopoeic rendering of their call (an oop-oop-oop) as is their generic name Upupa whilst the specific epithet epops was the name of a hoopoe in the Ancient Greek play The Birds by Aristophanes.

Day 69: Emerald toucanet – Aulacorhynchus prasinus

Big-billed birds

Whilst bearing a passing resemblance to hornbills (although not the species pictured above), the toucans, toucanets and aracaris are quite unrelated with their own family, Ramphastidae, in the order Piciformes.  Their resemblance (especially the large, flashy bills) reflects the convergent evolution of toucans into a similar niche of arboreal faunivory but in the forests of the Neotropics. Although sometimes making up more than half the length of the bird, the toucan’s beak is surprisingly light, being filled with spongy keratin. Serrations on their billss allow toucans to crush fruit whilst the length allows them to lazily reach many fruit without having to move. As with hornbills, toucan beak are huge advertising boards of colour.  

The emerald toucanet is (as the name suggests) one of the smaller toucan species. Though their bills are not as brightly coloured as some of the larger toucans and aracaris, the plumage of the green toucanets (Aulacorhynchus spp.) is vivid, the bright green sometimes embellished with red or blue. The emerald toucanet was my first wild toucan, seen feeding in the cloud forest canopy of Honduras. This was closely followed by the ornate collared aracari (Pteroglossus torquatus) and the spectacular keel-billed toucan (Ramphastos sulfuratus).

Day 70: Greater honeyguide – Indicator indicator

Show me the honey!

As mentioned above, toucans are members of the Piciformes. This is the last order of birds on our list so the end is very much in sight!  I do not have space to detail the bee-eater-like jacamars (Galbulidae) and the related puffbirds (Bucconidae) nor the four, chunky barbet families. However I will, of course, discuss at length the largest piciform family: the woodpeckers (Picidae). But first, honeyguides.

Honeyguides (Indicatoridae) look a bit out of place in the Piciformes, an order of well-built birds with large heads and beaks. In fact the honeyguides look like passerines, the humongous order of perching birds which would come next in the sequence however you should note the woodpecker-like zygodactylic feet. Although they are relatively dull birds, honeyguides are unusual in their regular consumption of wax which is taken mainly from bee hives. The greater honeyguide, feeds on the eggs, larvae, pupae and wax of bees nests and this has lead to a surprising relationship with man. As suggested by both the common name and the Latin Indicator, honeyguides lead humans (African bushmen) to honey.

Honey-guiding is a classic of example of a mutualism where individuals help each other for mutual benefit (symbiosis is a word often used instead of mutualism but symbiosis actually refers to organisms that inextricably depend on others where this relationship is not necessarily mutualistic. Conversely mutualists need not be obligate mutualists, but merely cooperative from time to time). It is relatively easy to explain this mutualism as a form of ‘partner choice’ where both parties must cooperate in order to receive the reward. Honeyguides lead hunters to bees’ nests with loud, distinct calls. After the humans have smashed open the nests and taken the honey, the birds can eat their fill. If the humans were to cheat the system, leaving nothing for the honeyguides, the selection pressure for the birds to guide would disappear followed by the behaviour itself. Likewise, the birds save a lot of time guiding the humans rather than waiting for them to find the bees themselves. Thus cooperation is needed on both sides for a successful relationship.  

Honey-guiding has only been demonstrated in this Indicator species despite the whole family sharing the name. One thing all members do share is brood parasitism, as seen in the cuckoos.