University President Richard Levin may be the longest-serving president in the Ivy League, but he’s not quite the highest-paid.

Levin’s total compensation — his base salary, benefits and deferred compensation — amounted to just over $1.6 million during the 2010 calendar year, placing him second in the Ivy League behind Columbia President Lee Bollinger, who made more than $1.9 million, according to the two universities’ most recent tax filings. Among Yale employees, Levin trails Chief Investment Officer David Swensen and Investments Office Senior Director Dean Takahashi, who earned roughly $3.5 million and $2.5 million in 2010, respectively.

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“These levels are driven by the market, by comparables in other universities,” Levin said. “Chief investment officers are in a market that pays a lot more than university administrators.”

Levin’s 2010 pay, which is determined by the Yale Corporation’s compensation committee, was about $16,000 less than in 2009, but roughly $1 million higher than it was a decade prior. Yale spokesman Tom Conroy said last year that Levin’s compensation reflects his longevity as president and the quality of his performance over his 20-year tenure.

Swensen declined to comment for this article, and Levin declined to comment on his own compensation.

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Brian Vogel, senior principal at Washington, D.C.-based consulting firm Quatt Associates and an expert in compensation of not-for-profit executives, said top administrators in higher education are often paid more the longer they hold a position, especially if liked by the institution’s board of trustees. But he added that a correlation between longevity and compensation is not always “clear cut” because when administrators do eventually step down, their replacements sometimes expect to earn a similar or even greater salary from the start.

Investment officials generally receive higher compensation than their administrative counterparts for a number of reasons, Vogel said, chief among them that the marketplace for investment and finance expertise simply pays more than the marketplace for university presidents. Investment officials also tend to have higher incentive payments, which are given when employees meet specific standards of performance.

Furthermore, the two types of employees are treated differently under tax law, Vogel said. A university president is classified as a “disqualified person” who is assumed to be able to influence his compensation, while employees in the Investments Office are not.

The University’s second highest administrator, Provost Peter Salovey, made about $560,000 in 2010.