Work out what Head of Biscay meant and what Bermeo lost when it stopped being it in 1602

Go into the Torre Ercilla for the fishermen's museum, which is the town's actual history

Find the Puerta de San Juan, the last surviving gate of the medieval walls

Watch the fleet work the old port, where nearly the whole town still lives

Walk out to the Atalaya and look along the coast towards Matxitxako and Gaztelugatxe

Bermeo was founded in 1236 and held the title of Head of Biscay from 1476 to 1602 — the principal town of the whole lordship, with privileges to match and the leading voice in the Juntas Generales. It lost the title to Bilbao and never got it back. The old port is still one of the most important fishing harbours on the Cantabrian coast, ninety-five per cent of the population lives round it, and Gaztelugatxe is just along the coast. The Torre Ercilla holds the fishermen's museum.

Bermeo: The Head of Biscay

Get to a hide on the marshes on a rising tide, which is when the waders come in off the flats

Work out the three bands — mountain, marsh and cliff coast — and how far apart they actually are

Go to the Urdaibai Bird Center at Gautegiz Arteaga for the ringing and the counts

Work out the scale: 22,452 hectares, about a tenth of Biscay, protected since 1984

Walk out to the cliffs and islands at the mouth, which are the seaward end of the same reserve

Urdaibai is the estuary of the Oka behind Gernika — 22,452 hectares, about 10% of the whole of Biscay, made a UNESCO Biosphere Reserve in 1984. It is the most important wetland in the Basque Country by size and condition and a major resting and wintering ground for migrating birds, which is why it was declared a Special Protection Area for birds in 1994 and added to the Ramsar list. It works as three bands running into each other: the mountain, the flats and marshes, and the coast with its cliffs, ports and beaches, and between them they hold the greatest diversity of landscape in the Basque Country.

Urdaibai: The Estuary Behind Gernika

Climb all 241 steps of the Urizarreta path down to the hermitage

Find the bell tower and read why its ringing rope was removed in 2021 to protect nesting seabirds

Stop at the upper viewpoint on the way down — the only point on the path showing both the east and west coastline together

Photograph the islet and its stepped causeway from the water's edge before climbing back up

A hermitage on a rock islet off the Biscay coast, reached only by a stepped stone causeway. Access has required a timed online reservation since 2021 to manage crowds drawn partly by its appearance as Dragonstone in Game of Thrones.

San Juan de Gaztelugatxe: 241 Steps to the Islet

Go to the Casa de Juntas and find the oak under which the fueros were sworn

Go through the Peace Museum, which reconstructs what 26 April 1937 was actually like from inside the town

Work out what was claimed afterwards — that the Basques burned their own town — and how long that lie survived

Find the reproduction of Picasso's Guernica in the town, and work out why the original is not here

Work out why this town and not another: it was the symbol of Basque self-government, and it was market day

Gernika was the symbolic seat of Basque law — the lords of Biscay swore to respect the fueros under an oak tree here. On Monday 26 April 1937 the German Condor Legion and the Italian Aviazione Legionaria bombed the town on market day. It held about five thousand people plus refugees, perhaps ten thousand in all. The rebel side then claimed for decades that the Basques had burned their own town, a lie that outlasted the war and the dictatorship. Picasso painted Guernica that year, and the town has been a byword for the bombing of civilians ever since.

Gernika: The Oak Tree and the Bombing

Check the tide and walk the causeway across to the island of San Nicolás

Go into the basilica of Santa María and look at the altarpiece, among the largest in Spain

Watch the boats come in and work out what a working Basque fishing port still looks like

Find out what the Antzar Eguna in September is and how it is done now

Walk the coast path along the cliffs either side of the town

Lekeitio has two beaches with the island of San Nicolás between them, and at low tide a stone causeway comes out of the water so you can walk across to it. The town is a working fishing port with a late Gothic basilica whose altarpiece is one of the largest in Spain. Its September festival includes the Antzar Eguna, in which crews in boats grab at a goose suspended over the harbour — now done with a dead bird, having been done otherwise for a very long time, which is the sort of thing Basque coastal towns do not apologise for.

Lekeitio: A Fishing Town and an Island

Find Jeff Koons' 12m-tall floral sculpture "Puppy" — replanted twice a year with 60,000 flowering plants — guarding the Guggenheim's entrance

Inside, find Richard Serra's "The Matter of Time" in Gallery 104 — eight weathering-steel sculptures filling a room 130m long

Walk from the Guggenheim to the Bilbao Fine Arts Museum through Doña Casilda Iturrizar Park, and compare the two museums' opposite approaches to art

Cross the Zubizuri footbridge and find Calatrava's glass-brick deck, now covered with a non-slip mat after years of complaints it was dangerous when wet

Bilbao was a dying port city with a poisoned river in 1991 when it gambled public money on a Frank Gehry building nobody could picture. It opened in 1997, drew a million visitors in its first year, and gave planners everywhere a phrase — the Bilbao effect — for the belief that one spectacular building can turn a city round. Most attempts since have failed. What is often missed is that Bilbao also cleaned the Nervión, built a metro and moved the port, and the Fine Arts Museum five minutes away had a better collection all along.

The Guggenheim Bilbao and the Bilbao Effect

Ride the hanging gondola across the Nervión between Portugalete and Getxo

Take the panoramic lift to the 45m-high walkway and cross it on foot

Find the UNESCO World Heritage plaque — Spain's only Industrial Heritage listing

Photograph the 1893 ironwork from the Portugalete waterfront promenade

The Vizcaya bridge was conceived, designed and built by private initiative between 1887 and 1893 and opened on 28 July 1893 — the first transporter bridge of its type anywhere in the world, and one of only eight still standing. Alberto de Palacio y Elissague designed it, with the French engineer Ferdinand Arnodin repeatedly mediating when Palacio and the builders fell out; Arnodin went on to build the very similar transporter bridge at Rochefort. It is still a working toll crossing: a gondola slung under the span carries vehicles and passengers between Portugalete and Las Arenas across the mouth of the Bilbao estuary.

Vizcaya Bridge: The World's First Transporter Bridge

Stand between the puente viejo and Calatrava's footbridge and work out how many centuries are in view

Find out what Alfonso XI granted the town in 1335 and what the bridge had to do with it

Walk the old quarter stacked up the hill above the port

Watch the fleet unload — this is one of the main fishing ports on the Cantabrian coast

Walk round to Arrigorri beach at the mouth of the ría

Alfonso XI granted Ondarroa the rights to the traffic over the wooden bridge across the Artibai in 1335, and the stone bridge that replaced it in the 18th century — the puente viejo — is still standing. Beside it Santiago Calatrava built a white asymmetric footbridge in the 1990s, so a working Basque fishing town has a 14th-century charter, an 18th-century bridge and a piece of late-20th-century engineering all within a hundred metres. The port is among the most important on the Cantabrian coast and the old quarter is packed onto the slope above it.

Ondarroa: A Bridge by Calatrava in a Fishing Town

Surf the beach, or take a lesson on it, which is what most people here are doing

Come in September for the Rip Curl Pro, which counts towards the world tour

Walk the whole three kilometres of sand and work out why it is called the Queen of Beaches

Find the Iñurritza biotope at the east end, one of very few surviving dune systems on this coast

Find the Palacio de Narros and work out which queens summered in it

Zarautz has the longest beach in the Basque Country at nearly three kilometres, which is why it is called the Queen of Beaches and why it has been the centre of Basque surfing for decades — the Rip Curl Pro Zarautz in September counts towards the world tour. Isabel II summered at the Palacio de Narros facing the sand, and so later did Queen Fabiola of Belgium. At the eastern end, round a golf course, the Iñurritza protected biotope has dunes, marsh and cliff together, one of very few preserved dune systems left on this coast.