diff --git a/spine-unity/Assets/Spine/Runtime/spine-unity/Components/Following/BoneFollower.cs b/spine-unity/Assets/Spine/Runtime/spine-unity/Components/Following/BoneFollower.cs
index 917648037..25c686bf0 100644
--- a/spine-unity/Assets/Spine/Runtime/spine-unity/Components/Following/BoneFollower.cs
+++ b/spine-unity/Assets/Spine/Runtime/spine-unity/Components/Following/BoneFollower.cs
@@ -156,6 +156,32 @@ namespace Spine.Unity {
 
 			Transform thisTransform = this.transform;
 			float additionalFlipScale = 1;
+
+			float scaleSignX = 1;
+			float scaleSignY = 1;
+			Bone parentBone = bone.Parent;
+			if (followParentWorldScale || followLocalScale || followSkeletonFlip) {
+				Vector3 localScale = new Vector3(1f, 1f, 1f);
+				if (followParentWorldScale && parentBone != null) {
+					float cumulativeScaleX = 1.0f;
+					float cumulativeScaleY = 1.0f;
+					Bone p = parentBone;
+					while (p != null) {
+						cumulativeScaleX *= p.ScaleX;
+						cumulativeScaleY *= p.ScaleY;
+						p = p.Parent;
+					};
+					scaleSignX = Mathf.Sign(cumulativeScaleX);
+					scaleSignY = Mathf.Sign(cumulativeScaleY);
+					localScale = new Vector3(parentBone.WorldScaleX * scaleSignX, parentBone.WorldScaleY * scaleSignY, 1f);
+				}
+				if (followLocalScale)
+					localScale.Scale(new Vector3(bone.ScaleX, bone.ScaleY, 1f));
+				if (followSkeletonFlip)
+					localScale.y *= Mathf.Sign(bone.Skeleton.ScaleX * bone.Skeleton.ScaleY) * additionalFlipScale;
+				thisTransform.localScale = localScale;
+			}
+
 			if (skeletonTransformIsParent) {
 				// Recommended setup: Use local transform properties if Spine GameObject is the immediate parent
 				thisTransform.localPosition = new Vector3(followXYPosition ? bone.WorldX : thisTransform.localPosition.x,
@@ -165,6 +191,8 @@ namespace Spine.Unity {
 					float halfRotation = Mathf.Atan2(bone.C, bone.A) * 0.5f;
 					if (followLocalScale && bone.ScaleX < 0) // Negate rotation from negative scaleX. Don't use negative determinant. local scaleY doesn't factor into used rotation.
 						halfRotation += Mathf.PI * 0.5f;
+					if (followParentWorldScale && scaleSignX < 0)
+						halfRotation += Mathf.PI * 0.5f;
 
 					Quaternion q = default(Quaternion);
 					q.z = Mathf.Sin(halfRotation);
@@ -196,6 +224,8 @@ namespace Spine.Unity {
 						if ((skeletonLossyScale.y * parentLossyScale.y < 0))
 							boneWorldRotation += 180f;
 					}
+					if (followParentWorldScale && scaleSignX < 0)
+						boneWorldRotation += 180f;
 
 					Vector3 worldRotation = skeletonTransform.rotation.eulerAngles;
 					if (followLocalScale && bone.ScaleX < 0) boneWorldRotation += 180f;
@@ -207,18 +237,6 @@ namespace Spine.Unity {
 				additionalFlipScale = Mathf.Sign(skeletonLossyScale.x * parentLossyScale.x
 												* skeletonLossyScale.y * parentLossyScale.y);
 			}
-
-			Bone parentBone = bone.Parent;
-			if (followParentWorldScale || followLocalScale || followSkeletonFlip) {
-				Vector3 localScale = new Vector3(1f, 1f, 1f);
-				if (followParentWorldScale && parentBone != null)
-					localScale = new Vector3(parentBone.WorldScaleX, parentBone.WorldScaleY, 1f);
-				if (followLocalScale)
-					localScale.Scale(new Vector3(bone.ScaleX, bone.ScaleY, 1f));
-				if (followSkeletonFlip)
-					localScale.y *= Mathf.Sign(bone.Skeleton.ScaleX * bone.Skeleton.ScaleY) * additionalFlipScale;
-				thisTransform.localScale = localScale;
-			}
 		}
 	}
 }
diff --git a/spine-unity/Assets/Spine/Runtime/spine-unity/Components/Following/BoneFollowerGraphic.cs b/spine-unity/Assets/Spine/Runtime/spine-unity/Components/Following/BoneFollowerGraphic.cs
index a7d3ce13d..3520bc487 100644
--- a/spine-unity/Assets/Spine/Runtime/spine-unity/Components/Following/BoneFollowerGraphic.cs
+++ b/spine-unity/Assets/Spine/Runtime/spine-unity/Components/Following/BoneFollowerGraphic.cs
@@ -141,8 +141,33 @@ namespace Spine.Unity {
 
 			float scale = skeletonGraphic.MeshScale;
 			Vector2 offset = skeletonGraphic.MeshOffset;
-
 			float additionalFlipScale = 1;
+
+			float scaleSignX = 1;
+			float scaleSignY = 1;
+			Bone parentBone = bone.Parent;
+			if (followParentWorldScale || followLocalScale || followSkeletonFlip) {
+				Vector3 localScale = new Vector3(1f, 1f, 1f);
+				if (followParentWorldScale && parentBone != null) {
+					float cumulativeScaleX = 1.0f;
+					float cumulativeScaleY = 1.0f;
+					Bone p = parentBone;
+					while (p != null) {
+						cumulativeScaleX *= p.ScaleX;
+						cumulativeScaleY *= p.ScaleY;
+						p = p.Parent;
+					};
+					scaleSignX = Mathf.Sign(cumulativeScaleX);
+					scaleSignY = Mathf.Sign(cumulativeScaleY);
+					localScale = new Vector3(parentBone.WorldScaleX * scaleSignX, parentBone.WorldScaleY * scaleSignY, 1f);
+				}
+				if (followLocalScale)
+					localScale.Scale(new Vector3(bone.ScaleX, bone.ScaleY, 1f));
+				if (followSkeletonFlip)
+					localScale.y *= Mathf.Sign(bone.Skeleton.ScaleX * bone.Skeleton.ScaleY) * additionalFlipScale;
+				thisTransform.localScale = localScale;
+			}
+
 			if (skeletonTransformIsParent) {
 				// Recommended setup: Use local transform properties if Spine GameObject is the immediate parent
 				thisTransform.localPosition = new Vector3(followXYPosition ? bone.WorldX * scale + offset.x : thisTransform.localPosition.x,
@@ -175,6 +200,8 @@ namespace Spine.Unity {
 						if ((skeletonLossyScale.y * parentLossyScale.y < 0))
 							boneWorldRotation += 180f;
 					}
+					if (followParentWorldScale && scaleSignX < 0)
+						boneWorldRotation += 180f;
 
 					Vector3 worldRotation = skeletonTransform.rotation.eulerAngles;
 					if (followLocalScale && bone.ScaleX < 0) boneWorldRotation += 180f;
@@ -186,18 +213,6 @@ namespace Spine.Unity {
 				additionalFlipScale = Mathf.Sign(skeletonLossyScale.x * parentLossyScale.x
 												* skeletonLossyScale.y * parentLossyScale.y);
 			}
-
-			Bone parentBone = bone.Parent;
-			if (followParentWorldScale || followLocalScale || followSkeletonFlip) {
-				Vector3 localScale = new Vector3(1f, 1f, 1f);
-				if (followParentWorldScale && parentBone != null)
-					localScale = new Vector3(parentBone.WorldScaleX, parentBone.WorldScaleY, 1f);
-				if (followLocalScale)
-					localScale.Scale(new Vector3(bone.ScaleX, bone.ScaleY, 1f));
-				if (followSkeletonFlip)
-					localScale.y *= Mathf.Sign(bone.Skeleton.ScaleX * bone.Skeleton.ScaleY) * additionalFlipScale;
-				thisTransform.localScale = localScale;
-			}
 		}
 	}
 }
diff --git a/spine-unity/Assets/Spine/package.json b/spine-unity/Assets/Spine/package.json
index f441579df..52c0436e9 100644
--- a/spine-unity/Assets/Spine/package.json
+++ b/spine-unity/Assets/Spine/package.json
@@ -2,7 +2,7 @@
 	"name": "com.esotericsoftware.spine.spine-unity",
 	"displayName": "spine-unity Runtime",
 	"description": "This plugin provides the spine-unity runtime core.",
-	"version": "4.3.0",
+	"version": "4.3.1",
 	"unity": "2018.3",
 	"author": {
 		"name": "Esoteric Software",