First-quarter funds from operation was $934 million or $2.48 per share.
FFO increased approximately $150 million or $0.31 per share compared to the fourth quarter of 2020.
And in fact, the quarter was negatively impacted by approximately $0.08 per share compared to our expectations given the closures that have occurred internationally.
We also recorded additional COVID impacts in the first quarter of approximately $0.07 per share from -- based upon basically domestic rent abatements and uncollectible rents.
We generated $875 million in cash from operations in the quarter, which was an increase of 18% compared to the prior-year period.
We collected over 95% of our net billed rents for the first quarter and our in-line tenant collections are back to pre-COVID levels in the approximate 98% range.
Our operating metrics in the period were as follows: mall and outlet occupancy at the end of the first quarter was 90.8%, down 50 basis points compared to the fourth quarter of 2020.
This 50-basis-point decline for the quarter is approximately 75% -- 75 basis points less than the average historical seasonal decline from the fourth quarter to the first quarter.
Average base rents was $56.07, up 60 basis points year over year.
Leasing spreads declined for the 12 -- trailing 12 months, primarily due to the mix of deals that have fallen out, the spread calculation that have resulted in an increase to the average closing rate by approximately $8 per square foot for the trailing 12 months.
Pricing continues to improve with the average opening rate per square foot for the trailing 12 months of approximately $60 per foot.
And as you can see in the lease expiration schedule included in our supplemental, our expiring rents for the next few years are less than $60 per square foot.
We signed 1,100 leases for approximately 4.4 million square feet, and we have significant number of leases in our pipeline, our leasing volume in both number of leases in square feet was greater than the volume in each of the first quarter of 2020 and 2019.
Our global brands within SPARC outperformed their plans in March and April on both sales and gross margin, led by Forever 21 and Aéropostale.
For the two months combined, SPARC outperformed the sales plan by more than $135 million and our gross margin plan by more than $75 million.
Our company's liquidity position at Penney is strong at $1.2 billion, and balance sheet is in very good shape with leverage of less than 1.2 times net debt to projected EBITDA.
We completed $1.5 billion senior note offering at 1.96%, weighted average term of 8.4 years.
We also completed a EUR 750 million note, shouldn't say dollar, at one and one-eighths percent coupon at a term of 12 years.
We used those proceeds to completely repay the $2 billion unsecured term facility associated with the Taubman deal, as well as pay off our $550 million senior notes.
We've also refinanced six mortgages for $1.3 billion, our share of which is $589 million at an average interest rate of 3.36%.
And at the end of the quarter, with all this activity, we have $8.4 billion of liquidity, consisting of $6.9 billion available on our credit facility; $1.5 billion of cash, including our share of JV cash.
And reminder, that is net of $500 million of U.S. commercial paper outstanding at quarter end.
We paid $1.30 per share in cash, in terms of our dividend on April 23.
And then, finally, as you've seen, given our first-quarter results, we are increasing our full-year 2021 FFO guidance from $9.50 to $9.75 per share to $9.70 to $9.80 per.
This is an increase of $0.20 per share at the bottom end of the range and $0.05 at the top end of the range or a 13% -- or a $0.13 increase at midpoint and that represents a 6.5% to 7.6% growth rate compared to our 2020 results.

Summary:
First-quarter funds from operation was $934 million or $2.48 per share.
Our operating metrics in the period were as follows: mall and outlet occupancy at the end of the first quarter was 90.8%, down 50 basis points compared to the fourth quarter of 2020.
We paid $1.30 per share in cash, in terms of our dividend on April 23.
And then, finally, as you've seen, given our first-quarter results, we are increasing our full-year 2021 FFO guidance from $9.50 to $9.75 per share to $9.70 to $9.80 per.