diff --git a/flask/app.py b/flask/app.py
index 1f7df2e8..6baee3a6 100644
--- a/flask/app.py
+++ b/flask/app.py
@@ -8,18 +8,18 @@
     :copyright: (c) 2015 by Armin Ronacher.
     :license: BSD, see LICENSE for more details.
 """
-
 import os
 import sys
 from threading import Lock
 from datetime import timedelta
 from itertools import chain
 from functools import update_wrapper
+from collections import Mapping
 
 from werkzeug.datastructures import ImmutableDict
 from werkzeug.routing import Map, Rule, RequestRedirect, BuildError
 from werkzeug.exceptions import HTTPException, InternalServerError, \
-     MethodNotAllowed, BadRequest
+     MethodNotAllowed, BadRequest, default_exceptions
 
 from .helpers import _PackageBoundObject, url_for, get_flashed_messages, \
      locked_cached_property, _endpoint_from_view_func, find_package
@@ -33,7 +33,7 @@ from .templating import DispatchingJinjaLoader, Environment, \
      _default_template_ctx_processor
 from .signals import request_started, request_finished, got_request_exception, \
      request_tearing_down, appcontext_tearing_down
-from ._compat import reraise, string_types, text_type, integer_types
+from ._compat import reraise, string_types, text_type, integer_types, iterkeys
 
 # a lock used for logger initialization
 _logger_lock = Lock()
@@ -1078,6 +1078,21 @@ class Flask(_PackageBoundObject):
             return f
         return decorator
 
+    @staticmethod
+    def _get_exc_class_and_code(exc_class_or_code):
+        """Ensure that we register only exceptions as handler keys"""
+        if isinstance(exc_class_or_code, integer_types):
+            exc_class = default_exceptions[exc_class_or_code]
+        else:
+            exc_class = exc_class_or_code
+        
+        assert issubclass(exc_class, Exception)
+        
+        if issubclass(exc_class, HTTPException):
+            return exc_class, exc_class.code
+        else:
+            return exc_class, None
+    
     @setupmethod
     def errorhandler(self, code_or_exception):
         """A decorator that is used to register a function give a given
@@ -1136,16 +1151,21 @@ class Flask(_PackageBoundObject):
 
     @setupmethod
     def _register_error_handler(self, key, code_or_exception, f):
-        if isinstance(code_or_exception, HTTPException):
-            code_or_exception = code_or_exception.code
-        if isinstance(code_or_exception, integer_types):
-            assert code_or_exception != 500 or key is None, \
-                'It is currently not possible to register a 500 internal ' \
-                'server error on a per-blueprint level.'
-            self.error_handler_spec.setdefault(key, {})[code_or_exception] = f
-        else:
-            self.error_handler_spec.setdefault(key, {}).setdefault(None, []) \
-                .append((code_or_exception, f))
+        """
+        :type key: None|str
+        :type code_or_exception: int|T<=Exception
+        :type f: callable
+        """
+        if isinstance(code_or_exception, HTTPException):  # old broken behavior
+            raise ValueError(
+                'Tried to register a handler for an exception instance {0!r}. '
+                'Handlers can only be registered for exception classes or HTTP error codes.'
+                .format(code_or_exception))
+        
+        exc_class, code = self._get_exc_class_and_code(code_or_exception)
+        
+        handlers = self.error_handler_spec.setdefault(key, {}).setdefault(code, {})
+        handlers[exc_class] = f
 
     @setupmethod
     def template_filter(self, name=None):
@@ -1386,6 +1406,33 @@ class Flask(_PackageBoundObject):
         self.url_default_functions.setdefault(None, []).append(f)
         return f
 
+    def _find_error_handler(self, e):
+        """Finds a registered error handler for the request’s blueprint.
+        If neither blueprint nor App has a suitable handler registered, returns None
+        """
+        exc_class, code = self._get_exc_class_and_code(type(e))
+        
+        def find_superclass(handler_map):
+            if not handler_map:
+                return None
+            for superclass in exc_class.__mro__:
+                if superclass is BaseException:
+                    return None
+                handler = handler_map.get(superclass)
+                if handler is not None:
+                    handler_map[exc_class] = handler  # cache for next time exc_class is raised
+                    return handler
+            return None
+        
+        # try blueprint handlers
+        handler = find_superclass(self.error_handler_spec.get(request.blueprint, {}).get(code))
+        
+        if handler is not None:
+            return handler
+        
+        # fall back to app handlers
+        return find_superclass(self.error_handler_spec[None].get(code))
+
     def handle_http_exception(self, e):
         """Handles an HTTP exception.  By default this will invoke the
         registered error handlers and fall back to returning the
@@ -1393,15 +1440,12 @@ class Flask(_PackageBoundObject):
 
         .. versionadded:: 0.3
         """
-        handlers = self.error_handler_spec.get(request.blueprint)
         # Proxy exceptions don't have error codes.  We want to always return
         # those unchanged as errors
         if e.code is None:
             return e
-        if handlers and e.code in handlers:
-            handler = handlers[e.code]
-        else:
-            handler = self.error_handler_spec[None].get(e.code)
+        
+        handler = self._find_error_handler(e)
         if handler is None:
             return e
         return handler(e)
@@ -1443,20 +1487,15 @@ class Flask(_PackageBoundObject):
         # wants the traceback preserved in handle_http_exception.  Of course
         # we cannot prevent users from trashing it themselves in a custom
         # trap_http_exception method so that's their fault then.
-
-        blueprint_handlers = ()
-        handlers = self.error_handler_spec.get(request.blueprint)
-        if handlers is not None:
-            blueprint_handlers = handlers.get(None, ())
-        app_handlers = self.error_handler_spec[None].get(None, ())
-        for typecheck, handler in chain(blueprint_handlers, app_handlers):
-            if isinstance(e, typecheck):
-                return handler(e)
-
+        
         if isinstance(e, HTTPException) and not self.trap_http_exception(e):
             return self.handle_http_exception(e)
 
-        reraise(exc_type, exc_value, tb)
+        handler = self._find_error_handler(e)
+        
+        if handler is None:
+            reraise(exc_type, exc_value, tb)
+        return handler(e)
 
     def handle_exception(self, e):
         """Default exception handling that kicks in when an exception
@@ -1470,7 +1509,7 @@ class Flask(_PackageBoundObject):
         exc_type, exc_value, tb = sys.exc_info()
 
         got_request_exception.send(self, exception=e)
-        handler = self.error_handler_spec[None].get(500)
+        handler = self._find_error_handler(InternalServerError())
 
         if self.propagate_exceptions:
             # if we want to repropagate the exception, we can attempt to
diff --git a/tests/test_user_error_handler.py b/tests/test_user_error_handler.py
new file mode 100644
index 00000000..78f4de3c
--- /dev/null
+++ b/tests/test_user_error_handler.py
@@ -0,0 +1,113 @@
+# -*- coding: utf-8 -*-
+from werkzeug.exceptions import Forbidden, InternalServerError
+import flask
+
+
+def test_error_handler_subclass():
+	app = flask.Flask(__name__)
+
+	class ParentException(Exception):
+		pass
+
+	class ChildExceptionUnregistered(ParentException):
+		pass
+
+	class ChildExceptionRegistered(ParentException):
+		pass
+
+	@app.errorhandler(ParentException)
+	def parent_exception_handler(e):
+		assert isinstance(e, ParentException)
+		return 'parent'
+
+	@app.errorhandler(ChildExceptionRegistered)
+	def child_exception_handler(e):
+		assert isinstance(e, ChildExceptionRegistered)
+		return 'child-registered'
+
+	@app.route('/parent')
+	def parent_test():
+		raise ParentException()
+
+	@app.route('/child-unregistered')
+	def unregistered_test():
+		raise ChildExceptionUnregistered()
+
+	@app.route('/child-registered')
+	def registered_test():
+		raise ChildExceptionRegistered()
+
+
+	c = app.test_client()
+
+	assert c.get('/parent').data == b'parent'
+	assert c.get('/child-unregistered').data == b'parent'
+	assert c.get('/child-registered').data == b'child-registered'
+
+
+def test_error_handler_http_subclass():
+	app = flask.Flask(__name__)
+
+	class ForbiddenSubclassRegistered(Forbidden):
+		pass
+
+	class ForbiddenSubclassUnregistered(Forbidden):
+		pass
+
+	@app.errorhandler(403)
+	def code_exception_handler(e):
+		assert isinstance(e, Forbidden)
+		return 'forbidden'
+
+	@app.errorhandler(ForbiddenSubclassRegistered)
+	def subclass_exception_handler(e):
+		assert isinstance(e, ForbiddenSubclassRegistered)
+		return 'forbidden-registered'
+
+	@app.route('/forbidden')
+	def forbidden_test():
+		raise Forbidden()
+
+	@app.route('/forbidden-registered')
+	def registered_test():
+		raise ForbiddenSubclassRegistered()
+
+	@app.route('/forbidden-unregistered')
+	def unregistered_test():
+		raise ForbiddenSubclassUnregistered()
+
+
+	c = app.test_client()
+
+	assert c.get('/forbidden').data == b'forbidden'
+	assert c.get('/forbidden-unregistered').data == b'forbidden'
+	assert c.get('/forbidden-registered').data == b'forbidden-registered'
+
+
+def test_error_handler_blueprint():
+	bp = flask.Blueprint('bp', __name__)
+	
+	@bp.errorhandler(500)
+	def bp_exception_handler(e):
+		return 'bp-error'
+	
+	@bp.route('/error')
+	def bp_test():
+		raise InternalServerError()
+	
+	app = flask.Flask(__name__)
+	
+	@app.errorhandler(500)
+	def app_exception_handler(e):
+		return 'app-error'
+	
+	@app.route('/error')
+	def app_test():
+		raise InternalServerError()
+	
+	app.register_blueprint(bp, url_prefix='/bp')
+	
+	c = app.test_client()
+	
+	assert c.get('/error').data == b'app-error'
+	assert c.get('/bp/error').data == b'bp-error'
\ No newline at end of file