diff --git a/lib/twine/twine_file.rb b/lib/twine/twine_file.rb
index dc524ae..7a15d8f 100644
--- a/lib/twine/twine_file.rb
+++ b/lib/twine/twine_file.rb
@@ -49,11 +49,14 @@ def matches_tags?(tags, include_untagged)
     end
 
     def translation_for_lang(lang)
-      translation = [lang].flatten.map { |l| @translations[l] }.compact.first
-
-      translation = reference.translation_for_lang(lang) if translation.nil? && reference
+      # Language priority also applies to inherited translations.
+      [lang].flatten.each do |language|
+        translation = @translations[language]
+        translation = reference.translation_for_lang(language) if translation.nil? && reference
+        return translation unless translation.nil?
+      end
 
-      return translation
+      nil
     end
 
     def plural_translation_for_lang(lang)
diff --git a/python_twine/tests/test_output_processor.py b/python_twine/tests/test_output_processor.py
index a9831bd..f1c77d9 100644
--- a/python_twine/tests/test_output_processor.py
+++ b/python_twine/tests/test_output_processor.py
@@ -137,10 +137,25 @@ def test_fallback_to_developer_language(self, twine_file):
 
         assert result.definitions_by_key["key1"].translations["de"] == "value1-fr"
 
+    def test_regional_locale_inherits_generic_reference_translation(self, twine_file):
+        reference = TwineDefinition("bridge")
+        reference.translations = {"en": "Bridge", "fr": "Pont"}
+        definition = TwineDefinition("pedestrian_bridge")
+        definition.translations["en"] = "Pedestrian Bridge"
+        definition.reference_key = reference.key
+        definition.reference = reference
+        twine_file.sections[0].definitions.extend([reference, definition])
+        twine_file.definitions_by_key.update(
+            {reference.key: reference, definition.key: definition}
+        )
+
+        result = OutputProcessor(twine_file, {"include": "all"}).process("fr-CA")
+
+        assert result.definitions_by_key["pedestrian_bridge"].translations["fr-CA"] == "Pont"
+
 
 class TestPluralEdgeCases:
-    """Regression tests for plural-definition handling that previously
-    raised KeyError / TypeError before the OutputProcessor fix."""
+    """Plural definitions with partial or missing target-language forms."""
 
     def test_include_translated_with_singular_target_does_not_raise(self):
         """A plural definition whose target language has only a non-plural
@@ -165,9 +180,7 @@ def test_include_translated_with_singular_target_does_not_raise(self):
         assert "ja" not in processed.plural_translations
 
     def test_no_plural_fallback_available_does_not_raise(self):
-        """When no fallback language has plural data, the OutputProcessor
-        must skip the plural augmentation rather than assigning None and
-        TypeError-ing on `"other" not in None`."""
+        """A singular translation remains usable when no plural fallback exists."""
         twine_file = TwineFile()
         twine_file.language_codes = ["en", "fr"]
         section = TwineSection("Section")
diff --git a/python_twine/tests/test_twine_definition.py b/python_twine/tests/test_twine_definition.py
index 8bcd0c8..f44b50c 100644
--- a/python_twine/tests/test_twine_definition.py
+++ b/python_twine/tests/test_twine_definition.py
@@ -140,3 +140,37 @@ def test_reference_translation_override(self, setup_references):
         definition.translations["en"] = "value"
 
         assert definition.translation_for_lang("en") == "value"
+
+    @pytest.mark.parametrize(
+        ("languages", "expected"),
+        [
+            (["fr", "en"], "Pont"),
+            (["de", "fr", "en"], "Pont"),
+            (["de", "en"], "Pedestrian Bridge"),
+            (["de"], None),
+            ([], None),
+        ],
+    )
+    def test_reference_respects_language_priority(
+        self, setup_references, languages, expected
+    ):
+        definition = setup_references
+        definition.translations["en"] = "Pedestrian Bridge"
+        definition.reference.translations["fr"] = "Pont"
+
+        assert definition.translation_for_lang(languages) == expected
+
+    def test_language_priority_through_reference_chain(self, setup_references):
+        definition = setup_references
+        definition.translations["en"] = "Pedestrian Bridge"
+        reference = TwineDefinition("bridge")
+        reference.translations["fr"] = "Pont"
+        definition.reference.reference = reference
+
+        assert definition.translation_for_lang(["fr", "en"]) == "Pont"
+
+    def test_empty_translation_overrides_reference(self, setup_references):
+        definition = setup_references
+        definition.translations["en"] = ""
+
+        assert definition.translation_for_lang(["en"]) == ""
diff --git a/python_twine/tests/test_twine_file.py b/python_twine/tests/test_twine_file.py
index 0af16b6..0073c04 100644
--- a/python_twine/tests/test_twine_file.py
+++ b/python_twine/tests/test_twine_file.py
@@ -7,6 +7,7 @@
 
 import pytest
 
+from twine.output_processor import OutputProcessor
 from twine.twine_file import TwineDefinition, TwineFile, TwineSection
 
 
@@ -221,6 +222,49 @@ def test_accent_symbol(self, fixtures_dir):
         assert twine_file.definitions_by_key["value_wrapped_by_accents"].translations["en"] == '`value`'
 
 class TestTwineFileOptimizations:
+    @pytest.mark.parametrize(
+        ("content", "fallback_to_default"),
+        [
+            ("[bridge]\nen = Bridge\nzh-Hans = 桥\n"
+             "[pedestrian]\nref = bridge\nen = Pedestrian Bridge\n"
+             "zh = 人行桥\nzh-CN = 人行桥\n", False),
+            ("[bridge]\nen = Bridge\nfr = Pont\n"
+             "[pedestrian]\nref = bridge\nen = Pedestrian Bridge\n"
+             "fr-CA = Pedestrian Bridge\n", True),
+            ("[bridge]\nen = Bridge\nfr = Pont\nfr-CA = Pont\n"
+             "[pedestrian]\nref = bridge\nfr = Passerelle\n", False),
+            ("[bridge]\nfr-CA = Pont\nfr = Pont\nen = Bridge\n", True),
+        ],
+        ids=["inherited-script", "inherited-generic", "alias-consumer", "regional-developer"],
+    )
+    def test_optimization_preserves_generated_translations(
+        self, tmp_path, content, fallback_to_default
+    ):
+        path = tmp_path / "strings.txt"
+        path.write_text(content, encoding="utf-8")
+        twine_file = TwineFile(fallback_to_default=fallback_to_default)
+        twine_file.read(str(path))
+        languages = [*twine_file.language_codes, "de", "fr-CA", "zh-CN"]
+
+        def translations(source):
+            processor = OutputProcessor(source, {"include": "all"})
+            return {
+                language: {
+                    key: definition.translations[language]
+                    for key, definition in processor.process(language).definitions_by_key.items()
+                }
+                for language in languages
+            }
+
+        expected = translations(twine_file)
+        twine_file.optimize_duplicates()
+        assert translations(twine_file) == expected
+
+        twine_file.write(str(path))
+        restored = TwineFile()
+        restored.read(str(path))
+        assert translations(restored) == expected
+
     @pytest.fixture
     def fixture_twine_file(self) -> TwineFile:
         twine_file = TwineFile()
diff --git a/python_twine/twine/twine_file.py b/python_twine/twine/twine_file.py
index 5308110..df223eb 100644
--- a/python_twine/twine/twine_file.py
+++ b/python_twine/twine/twine_file.py
@@ -97,30 +97,18 @@ def matches_tags(
         return True
 
     def translation_for_lang(self, lang: str | list[str]) -> str | None:
-        """
-        Get translation for a language, checking reference if not found.
-
-        Args:
-            lang: Language code (can be str or list)
-
-        Returns:
-            Translation string or None
-        """
-        # Handle both single lang and list of langs
+        """Resolve languages in priority order, including inherited translations."""
         if isinstance(lang, list):
-            for ln in lang:
-                if ln in self.translations:
-                    return self.translations[ln]
-            lang_to_check = lang
-        else:
-            if lang in self.translations:
-                return self.translations[lang]
-            lang_to_check = [lang]
+            for language in lang:
+                translation = self.translation_for_lang(language)
+                if translation is not None:
+                    return translation
+            return None
 
-        # Check reference
+        if lang in self.translations:
+            return self.translations[lang]
         if self.reference:
-            return self.reference.translation_for_lang(lang_to_check)
-
+            return self.reference.translation_for_lang(lang)
         return None
 
     def find_plural_lang_fallback(self, fallback_langs: list[str]) -> str | None:
@@ -229,20 +217,36 @@ def get_developer_language_code(self) -> str | None:
         return None
 
     def optimize_duplicates(self):
-        # Check if translation matches value from 'ref' definition.
+        """Remove translations only when their effective fallback is identical."""
+        referenced = {
+            definition.reference for definition in self.definitions_by_key.values()
+            if definition.reference is not None
+        }
+        developer_language = self.get_developer_language_code()
         for key, definition in self.definitions_by_key.items():
-            if definition.reference is not None:
-                ref = definition.reference
-                definition.translations = {lang:value for lang, value in definition.translations.items()
-                                           if lang not in ref.translations or ref.translations[lang] != value}
-
-        """ Some regional languages have common items. Such as 'en-GB' and 'en'.
-            Deduplication: for each item and each language search the same translations
-            within fallback languages. Not all languages have fallbacks.
-        """
-        for key, definition in self.definitions_by_key.items():
-            definition.translations = {lang:value for (lang, value) in definition.translations.items() \
-                                       if not self.match_fallback_lang(definition.translations, lang, key, value)}
+            # Apply deletions one at a time so subsequent comparisons see the
+            # translations that will actually remain available.
+            for lang, value in list(definition.translations.items()):
+                inherited = (
+                    definition.reference.translation_for_lang(lang)
+                    if definition.reference is not None else None
+                )
+                if inherited is not None:
+                    if inherited == value:
+                        del definition.translations[lang]
+                    continue
+
+                # Aliases inherit exact-language values, without regional
+                # fallback. The developer language is also a terminal fallback.
+                if definition in referenced or lang == developer_language:
+                    continue
+                fallback = definition.translation_for_lang(
+                    self.fallback_languages(lang, self.fallback_to_default)
+                )
+                if fallback == value:
+                    print(f"Warning: key '{key}' in lang '{lang}' matches its fallback translation")
+                    del definition.translations[lang]
+
             definition.plural_translations = {lang:value for (lang, value) in definition.plural_translations.items() \
                                               if not self.match_fallback_lang(definition.plural_translations, lang, key, value)}
 
diff --git a/test/test_twine_definition.rb b/test/test_twine_definition.rb
index ccfbe38..1b3adb7 100644
--- a/test/test_twine_definition.rb
+++ b/test/test_twine_definition.rb
@@ -107,5 +107,31 @@ def test_reference_translation_override
 
       assert_equal 'value', @definition.translation_for_lang('en')
     end
+
+    def test_reference_respects_language_priority
+      @definition.translations['en'] = 'Pedestrian Bridge'
+      @reference.translations['fr'] = 'Pont'
+
+      assert_equal 'Pont', @definition.translation_for_lang(%w(fr en))
+      assert_equal 'Pont', @definition.translation_for_lang(%w(de fr en))
+      assert_equal 'Pedestrian Bridge', @definition.translation_for_lang(%w(de en))
+      assert_nil @definition.translation_for_lang(['de'])
+      assert_nil @definition.translation_for_lang([])
+    end
+
+    def test_language_priority_through_reference_chain
+      @definition.translations['en'] = 'Pedestrian Bridge'
+      reference = Twine::TwineDefinition.new 'bridge'
+      reference.translations['fr'] = 'Pont'
+      @reference.reference = reference
+
+      assert_equal 'Pont', @definition.translation_for_lang(%w(fr en))
+    end
+
+    def test_empty_translation_overrides_reference
+      @definition.translations['en'] = ''
+
+      assert_equal '', @definition.translation_for_lang(['en'])
+    end
   end
 end