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The proposed Keystone XL oil pipeline from Canada into the US has survived multiple environmental reviews, law suits, and a rejection from the Obama administration that was then overturned by President Donald Trump. Now it faces another obstacle: the pallid sturgeon.

Nebraska’s Public Service Commission on Monday granted approval for the pipeline to cross the state — the last major permit it needed — but insisted on a different route from the one preferred by Transcanada, the company leading the project.

Changing the route, according to an expert called by Transcanada to give evidence to the commission, raises the risk to the pallid sturgeon, a rare bottom-feeding fish found in US rivers.

The pallid sturgeon is an ancient species that has been around since the Cretaceous period 70m years ago, and looks a bit like a dinosaur with bony plates down its back. It described by the US Fish & Wildlife Service as “one of the least understood fish in the Missouri and Mississippi River drainages”, and its numbers have been in steep decline since the mid-1960s.

It is just one of the new issues raised by the PSC decision that could further delay the $8bn pipeline, which has already been under review for nine years.

Brigham McCown of the Alliance for Innovation and Infrastructure, a senior pipeline safety official under President George W Bush, said Nebraska’s decision was “another step forward for the project, but far from a decisive victory”.

He added: “It’s not the end of the process at all.”

One of the central issues will be whether the state department’s broadly favourable assessment of Keystone XL’s environmental impact in 2014 is still valid. The Trump administration used that statement in March as its basis for issuing the presidential permit needed for Keystone XL to proceed.

That issue is already being fought out in a US court in Montana, where environmental and Native American groups have brought cases arguing that the new administration should not have been able to rely on a study that was three years old. The change of route in Nebraska could similarly create scope for arguments that the 2014 environmental impact statement should no longer apply.

That assessment concluded that only one endangered species was likely to be “adversely affected” by the project: the American burying beetle. But Jon Schmidt, a biologist who works as a consultant on pipeline projects, testified to the Nebraska PSC that using the “mainline alternative route”, the one that the commission eventually approved, would mean crossing the ranges of four threatened and endangered species of fish: the pallid sturgeon, the Topeka shiner, the sturgeon chub, and the lake sturgeon. Those threats could be used to build a case that an additional environmental impact assessment is needed.

Changing the pipeline’s route also means Transcanada will need to reach agreements with a new set of landowners, or use the right of “eminent domain” to compel them to allow use of the land.

Doug Hayes, a senior attorney with the Sierra Club environmental group, said changing the route opened Transcanada up to “a whole new world of legal obstacles”.

Direct action by protesters can also be expected. Calls have been circulating on Facebook this week for people to prepare for “creative peaceful resistance along the pipeline route”, probably in the spring.

Transcanada responded cautiously to the PSC decision, saying it would have to assess how the cost and schedule of the project would be affected. Other proposed pipelines for exporting oil from western Canada have been running into difficulties.

In October, Transcanada abandoned its planned Energy East project, which would have been an alternative to Keystone XL. Chief executive Russ Girling told analysts earlier this month that demand for Keystone XL was as strong as when the project was first proposed back in 2008.

But Mr McCown argued there was still a good chance the pipeline would not go ahead, not so much because of environmentalists’ opposition, but because at today’s oil prices there is less demand for relatively high-cost crude from the Canadian oil sands.

“If oil were $130 a barrel, they would not be hanging around. They’d bring down the hammer and it would get done,” he said.

“If it keeps languishing in the courts for more years to come, it may reach the point where the costs exceeds the benefits.”

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