The plight of the animals stuck onboard the Spiridon II livestock carrier caused an outcry among animal welfare groups.

Turkish authorities refused the ship entry to the northwestern port of Bandirma last month, citing discrepancies in the animals' health certificates.

Uruguay's agriculture ministry said Monday that they had been disembarked safe and sound in the Libyan port of Benghazi after a total of two months at sea.

But the Animal Welfare Foundation cast doubt on the animals' wellbeing, claiming that dozens died during the month-long crossing from Uruguay to Turkey.

The charity called for an investigation into the case, which it said illustrated the dangers of transporting live animals by sea.

It said it was concerned that carcasses may have been illicitly thrown overboard.

Uruguay blamed the animals' plight on a disagreement between the cattle exporter and his Turkish customer.

The head of Ganosan, the Uruguayan cattle exporter which shipped the animals, denied any wrongdoing and instead blamed his Turkish customer.

"We sold the cattle, we put them on the ship, the man paid, he chartered the ship, the ship left, and that's where our business ended, and we have the proof," Fernando Fernandez told Uruguayan media.

The small South American nation, which has nearly four times as many cattle as inhabitants, is a major beef producer.

The shipment of 2,901 cattle left Montevideo in mid-September.

It arrived off Turkey on October 21 and remained moored off the coast for a month while the dispute over documentation dragged on.