仅在某些不匹配模式的行上用sed替换

我有一个文件(标准FASTA格式),如下所示。我想替换所有的“-”字符,但只能替换不是以“>”开头的行。例如:

>Title1-mytitle
ACACAGCATGCTA-CTAGCAGT
>Title2
---CAGCTAGCAG-TGACGCGA
GTCAGCTGACGTAGCTAGTATA
CGATCGATCGTAGCT-GAGATT
>Title3-again-q
ATGCTAGCTGACTGATCGTCGG
ATGATGATTTG

输出如下所示:

>Title1-mytitle
ACACAGCATGCTACTAGCAGT
>Title2
CAGCTAGCAGTGACGCGA
GTCAGCTGACGTAGCTAGTATA
CGATCGATCGTAGCTGAGATT
>Title3-again-q
ATGCTAGCTGACTGATCGTCGG
ATGATGATTTG

I require a bash solution for this to be put in a longer pipeline. I was thinking along the lines of sed with a particularly smart regular expression. Do you have some ideas?

谢谢!