diff --git a/lib/templatesimplifier.cpp b/lib/templatesimplifier.cpp
index 1bf98c9dc..9183accbf 100644
--- a/lib/templatesimplifier.cpp
+++ b/lib/templatesimplifier.cpp
@@ -31,6 +31,38 @@
 #include <string>
 #include <cassert>
 
+#ifdef GDB_HELPERS
+
+#include <iostream>
+
+static void printlist(const std::list<Token *> &list)
+{
+    for (std::list<Token *>::const_iterator it = list.begin(); it != list.end(); it++) {
+        const Token *token = *it;
+        std::cout << "   ";
+        while (token && !Token::Match(token, "[{};]")) {
+            std::cout << " " << token->str();
+            token = token->next();
+        }
+        std::cout << std::endl;
+    }
+}
+
+static void printvector(const std::vector<const Token *> &v)
+{
+    for (unsigned int i = 0; i < v.size(); i++) {
+        const Token *token = v[i];
+        std::cout << "    " << i << ":";
+        while (token && !Token::Match(token, "[{};]")) {
+            std::cout << " " << token->str();
+            token = token->next();
+        }
+        std::cout << std::endl;
+    }
+}
+
+#endif
+
 //---------------------------------------------------------------------------
 
 void TemplateSimplifier::cleanupAfterSimplify(Token *tokens)
@@ -620,9 +652,16 @@ void TemplateSimplifier::expandTemplate(
 
                 // replace type with given type..
                 if (itype < typeParametersInDeclaration.size()) {
+                    unsigned int typeindentlevel = 0;
                     for (const Token *typetok = typesUsedInTemplateInstantiation[itype];
-                         typetok && !Token::Match(typetok, "[,>]");
+                         typetok && (typeindentlevel>0 || !Token::Match(typetok, "[,>]"));
                          typetok = typetok->next()) {
+                        if (Token::Match(typetok, "%var% <") && templateParameters(typetok->next()) > 0)
+                            ++typeindentlevel;
+                        else if (typeindentlevel > 0 && typetok->str() == ">")
+                            --typeindentlevel;
+                        else if (typeindentlevel > 0 && typetok->str() == ">>")
+                            typeindentlevel -= (typeindentlevel > 1) ? 2 : 1;
                         tokenlist.addtoken(typetok, tok3->linenr(), tok3->fileIndex());
                     }
                     continue;
diff --git a/test/testsimplifytokens.cpp b/test/testsimplifytokens.cpp
index 954b155a5..78b42781b 100644
--- a/test/testsimplifytokens.cpp
+++ b/test/testsimplifytokens.cpp
@@ -127,6 +127,7 @@ private:
         TEST_CASE(template33);  // #3818 - inner templates in template instantiation not handled well
         TEST_CASE(template34);  // #3706 - namespace => hang
         TEST_CASE(template35);  // #4074 - A<'x'> a;
+        TEST_CASE(template36);  // #4310 - passing unknown template instantiation as template argument
         TEST_CASE(template_unhandled);
         TEST_CASE(template_default_parameter);
         TEST_CASE(template_default_type);
@@ -2241,6 +2242,16 @@ private:
         ASSERT_EQUALS("A<'x'> a ; class A<'x'> { }", tok(code));
     }
 
+    void template36() { // #4310 - Passing unknown template instantiation as template argument
+        const char code[] = "template <class T> struct X { T t; };\n"
+                            "template <class C> struct Y { Foo < X< Bar<C> > > _foo; };\n" // <- Bar is unknown
+                            "Y<int> bar;";
+        ASSERT_EQUALS("Y<int> bar ; "
+                      "struct Y<int> { Foo < X<Bar<int>> > _foo ; } "
+                      "struct X<Bar<int>> { Bar < int > t ; }",
+                      tok(code));
+    }
+
     void template_unhandled() {
         // An unhandled template usage should be simplified..
         ASSERT_EQUALS("x<int> ( ) ;", tok("x<int>();"));