diff --git a/crates/perry-runtime/Cargo.toml b/crates/perry-runtime/Cargo.toml
index dda516c511..11e031de29 100644
--- a/crates/perry-runtime/Cargo.toml
+++ b/crates/perry-runtime/Cargo.toml
@@ -25,17 +25,22 @@ crate-type = ["rlib"]
 default = ["full", "regex-engine", "temporal", "url-engine", "string-normalize", "intl-segmenter", "intl-namespace", "global-math", "global-json", "global-reflect", "global-atomics", "global-url", "global-text", "global-websocket", "global-webcrypto", "global-webfetch", "proc-ipc", "intl-locale", "intl-datetime", "diagnostics", "mod-dgram", "mod-http2-constants", "mod-node-test", "dyn-eval", "keepalive-anchors", "alloc-mimalloc"]
 # Compile the ~490 `#[used]` keep-alive anchor statics (`KEEP_*`, `keep!`,
 # `K*`/`G*` fn-pointer statics) that pin codegen-facing `#[no_mangle]` entry
-# points as dead-strip roots. They exist for the whole-program bitcode-LTO
-# link (`PERRY_LLVM_BITCODE_LINK=1`), where the merged module could otherwise
-# internalize + drop symbols only referenced from the user's separate `.o`.
-# The CLASSIC link path never needs them: every reachable runtime symbol is
-# kept by a real undefined reference from the program's objects, and the
-# anchors only defeat `-dead_strip` (measured: a hello-world binary carries
-# the entire fs/child_process/vm/node_submodules surface through ONE anchored
-# chain — KEEP_JS_MODULE_DYNAMIC_IMPORT_DEFERRED → dynamic-import fallback →
-# js_nm_install_all → every dispatch bucket). Kept in `default` so the
-# prebuilt/full archives and `cargo test` are unchanged; the auto-optimize
-# rebuild omits it unless the bitcode link was requested.
+# points so they survive into the staticlib archive. Without these anchors,
+# rustc dead-code-eliminates `#[no_mangle] pub extern "C" fn` symbols that are
+# only called from the program's generated object files (not from within the
+# perry-runtime crate itself), producing an incomplete `libperry_runtime.a`
+# that fails to link with "Undefined symbols" when the codegen emits calls to
+# them (e.g. `js_box_release`, `js_bool_box_release`,
+# `js_closure_set_box_capture_ptr`, `js_link_path_module_parent`).
+#
+# In a staticlib archive (`.a`) the linker only pulls in object files that
+# resolve an undefined reference, so `#[used]` anchors only become
+# `-dead_strip` roots when their object file is pulled in — i.e. when the
+# program actually references a symbol from that unit. The size cost is
+# therefore limited to the transitive callees of symbols the program uses
+# (which would be kept anyway), not the entire runtime surface. Kept in
+# `default` so the prebuilt/full archives and `cargo test` are unchanged;
+# the auto-optimize rebuild always re-adds it (see optimized_libs.rs).
 keepalive-anchors = []
 # Route every Rust heap allocation through mimalloc (#62) — ~3-5× faster
 # per-alloc than macOS `malloc` on hot paths, at ~140 KB of binary. In
diff --git a/crates/perry/src/commands/compile/optimized_libs/freshness.rs b/crates/perry/src/commands/compile/optimized_libs/freshness.rs
index 8268ba68c1..543f0280e9 100644
--- a/crates/perry/src/commands/compile/optimized_libs/freshness.rs
+++ b/crates/perry/src/commands/compile/optimized_libs/freshness.rs
@@ -157,7 +157,10 @@ pub(crate) fn auto_optimized_cache_key(
                 ""
             }
         ),
-        std::env::var("PERRY_LLVM_BITCODE_LINK").ok().as_deref() == Some("1"),
+        // keepalive-anchors is now always enabled (see auto_optimized_cross_features),
+        // so the cache key always reflects anchors=true. The field stays in the key
+        // so a future change to gate it again would get its own cache dir.
+        true,
         env!("CARGO_PKG_VERSION"),
     )
 }
@@ -286,15 +289,32 @@ pub(crate) fn auto_optimized_cross_features(
     // system-allocator ranges, the feature stays force-on; the cfg gate in
     // perry-runtime remains for that future audit.
     cross_features.push("perry-runtime/alloc-mimalloc".to_string());
-    // The `#[used]` keep-alive anchors exist for the whole-program bitcode
-    // LTO path only (see perry-runtime's `keepalive-anchors` feature docs).
-    // The classic link keeps every reachable symbol via real undefined
-    // references, so the anchors are omitted there — that is what lets
-    // `-dead_strip` drop the never-imported node-module surface from small
-    // programs. Re-enable them whenever the bitcode link was requested.
-    if std::env::var("PERRY_LLVM_BITCODE_LINK").ok().as_deref() == Some("1") {
-        cross_features.push("perry-runtime/keepalive-anchors".to_string());
-    }
+    // `#[used]` keep-alive anchors must be present on EVERY auto-optimize
+    // rebuild, not just the bitcode-LTO path. The anchors pin codegen-only
+    // `#[no_mangle] pub extern "C" fn` symbols (e.g. `js_box_release`,
+    // `js_bool_box_release`, `js_closure_set_box_capture_ptr`,
+    // `js_link_path_module_parent`) that are never called from within the
+    // perry-runtime crate itself — only from the program's generated object
+    // files. Without `#[used]`, rustc dead-code-eliminates these unreferenced
+    // symbols during staticlib archive creation, so they are absent from
+    // `libperry_runtime.a` and the final link fails with "Undefined symbols
+    // for architecture arm64" when the program's codegen emits calls to them.
+    //
+    // #6917 gated the anchors behind `keepalive-anchors` (bitcode-LTO only)
+    // under the assumption that "the classic link keeps every reachable
+    // runtime symbol via real undefined references from the program's
+    // objects." That assumption is wrong: the program's undefined references
+    // can only resolve symbols that are IN the archive, and the archive
+    // doesn't contain symbols that were DCE'd during archive creation.
+    //
+    // In a staticlib archive (`.a`) the linker only pulls in object files
+    // that resolve an undefined reference, so `#[used]` anchors only become
+    // `-dead_strip` roots when their object file is pulled in — i.e. when
+    // the program actually references a symbol from that unit. The size
+    // regression is therefore limited to the transitive callees of symbols
+    // the program uses (which would be kept anyway), not the entire runtime
+    // surface. Re-enable unconditionally; the bitcode-LTO path already did.
+    cross_features.push("perry-runtime/keepalive-anchors".to_string());
     // Compile OUT perry-runtime's no-op fetch stubs (`js_fetch_with_options` /
     // `js_headers_new` / `js_request_new`, gated `#[cfg(not(feature =
     // "external-fetch-symbols"))]`) whenever the program uses fetch — perry-stdlib's
diff --git a/crates/perry/src/commands/compile/optimized_libs/tests.rs b/crates/perry/src/commands/compile/optimized_libs/tests.rs
index b24c810174..cf636b0d9e 100644
--- a/crates/perry/src/commands/compile/optimized_libs/tests.rs
+++ b/crates/perry/src/commands/compile/optimized_libs/tests.rs
@@ -1053,6 +1053,60 @@ fn retain_workspace_declared_features_keeps_all_without_manifests() {
     assert_eq!(cross_features.len(), 2);
 }
 
+/// Regression: the auto-optimize rebuild must always include
+/// `perry-runtime/keepalive-anchors` in the cross-feature set. #6917 gated
+/// the `#[used]` keepalive anchors behind the feature and only enabled it for
+/// the bitcode-LTO path, under the assumption that the classic link path
+/// "keeps every reachable runtime symbol via real undefined references from
+/// the program's objects." That assumption is wrong: `#[no_mangle] pub extern
+/// "C" fn` symbols that are only called from codegen (not from within the
+/// perry-runtime crate) are dead-code-eliminated during staticlib archive
+/// creation when no `#[used]` anchor pins them. The resulting
+/// `libperry_runtime.a` is missing core symbols (`js_box_release`,
+/// `js_bool_box_release`, `js_closure_set_box_capture_ptr`,
+/// `js_link_path_module_parent`) and programs whose codegen emits calls to
+/// them fail to link with "Undefined symbols for architecture arm64."
+#[test]
+fn auto_optimize_always_includes_keepalive_anchors() {
+    let dir = tempfile::tempdir().expect("tempdir");
+    let empty_features: std::collections::BTreeSet<&'static str> =
+        std::collections::BTreeSet::new();
+    let ctx = CompilationContext::new(dir.path().to_path_buf());
+    let cross = auto_optimized_cross_features(&ctx, &empty_features, &[]);
+    assert!(
+        cross.iter().any(|f| f == "perry-runtime/keepalive-anchors"),
+        "auto_optimized_cross_features must always include \
+         perry-runtime/keepalive-anchors so codegen-only #[no_mangle] symbols \
+         survive into the staticlib archive, got {cross:?}"
+    );
+}
+
+/// The `keepalive-anchors` feature must NOT be conditional on
+/// `PERRY_LLVM_BITCODE_LINK` — the classic link path needs it too.
+#[test]
+fn auto_optimize_keepalive_anchors_not_bitcode_only() {
+    let _guard = env_lock();
+    let old_bitcode = std::env::var_os("PERRY_LLVM_BITCODE_LINK");
+    std::env::remove_var("PERRY_LLVM_BITCODE_LINK");
+
+    let dir = tempfile::tempdir().expect("tempdir");
+    let empty_features: std::collections::BTreeSet<&'static str> =
+        std::collections::BTreeSet::new();
+    let ctx = CompilationContext::new(dir.path().to_path_buf());
+    let cross = auto_optimized_cross_features(&ctx, &empty_features, &[]);
+
+    set_env_var(
+        "PERRY_LLVM_BITCODE_LINK",
+        old_bitcode.as_deref().and_then(|v| v.to_str()),
+    );
+
+    assert!(
+        cross.iter().any(|f| f == "perry-runtime/keepalive-anchors"),
+        "keepalive-anchors must be present even without PERRY_LLVM_BITCODE_LINK, \
+         got {cross:?}"
+    );
+}
+
 /// The well-known flip drops `compression-brotli`/`compression-zstd` from the
 /// stdlib rebuild on a stated premise: "The ext crate carries all codecs, so
 /// nothing is lost by dropping them here."