diff --git a/cpplint.py b/cpplint.py
index 07687d2..2f97c63 100755
--- a/cpplint.py
+++ b/cpplint.py
@@ -3943,14 +3943,15 @@ def CheckForNonStandardConstructs(filename, clean_lines, linenum, nesting_state,
         else:
             constructor_args = explicit_constructor_match.group(2).split(",")
 
-        # collapse arguments so that commas in template parameter lists and function
-        # argument parameter lists don't split arguments in two
+        # Collapse commas inside template and function parameter lists.
         i = 0
         while i < len(constructor_args):
             constructor_arg = constructor_args[i]
-            while constructor_arg.count("<") > constructor_arg.count(">") or constructor_arg.count(
-                "("
-            ) > constructor_arg.count(")"):
+            # Ignore `<`-prefixed operators when balancing template brackets.
+            while i + 1 < len(constructor_args) and (
+                re.sub(r"<<=?|<=", "", constructor_arg).count("<") > constructor_arg.count(">")
+                or constructor_arg.count("(") > constructor_arg.count(")")
+            ):
                 constructor_arg += "," + constructor_args[i + 1]
                 del constructor_args[i + 1]
             constructor_args[i] = constructor_arg
diff --git a/cpplint_unittest.py b/cpplint_unittest.py
index d568251..48fd9c4 100755
--- a/cpplint_unittest.py
+++ b/cpplint_unittest.py
@@ -1810,6 +1810,37 @@ class Foo {
                 "Constructors callable with one argument should be marked explicit."
                 "  [runtime/explicit] [4]",
             )
+            # `<`-containing operators are not confused with template brackets.
+            self.TestMultiLineLint(
+                """
+          class A {
+            A(int a, int b, int c = 1 << 1);
+          };""",
+                "",
+            )
+            self.TestMultiLineLint(
+                """
+          class A {
+            A(int a = (value <<= 1), int b = 0, int c = 0);
+          };""",
+                "Constructors callable with one argument should be marked explicit."
+                "  [runtime/explicit] [4]",
+            )
+            self.TestMultiLineLint(
+                """
+          class A {
+            A(int a, bool b = 1 <= 1, int c = 0);
+          };""",
+                "Constructors callable with one argument should be marked explicit."
+                "  [runtime/explicit] [4]",
+            )
+            self.TestMultiLineLint(
+                """
+          class A {
+            A(int a, int b, bool c = 1 < 2);
+          };""",
+                "",
+            )
             # explicit no-argument constructors are just fine
             self.TestMultiLineLint(
                 """