diff --git a/docs/modules/theme/pages/add-running-content.adoc b/docs/modules/theme/pages/add-running-content.adoc
index f94c82b54..c010d0f44 100644
--- a/docs/modules/theme/pages/add-running-content.adoc
+++ b/docs/modules/theme/pages/add-running-content.adoc
@@ -34,21 +34,26 @@ running-content:
 
 The `start-at` key accepts the following keywords or an integer:
 
-after-toc:: The running content starts after the TOC, no matter where the TOC is placed in the document.
-The `after-toc` value is only recognized if the title page is implicitly or explicitly enabled.
-If `media=prepress`, this keyword refers to the first recto page after the last page of the TOC (it will skip an empty verso page).
+after-toc:: The running content starts after the last page of the TOC, no matter where the TOC is placed in the document.
+If `media=prepress`, this keyword refers to the next recto page after the last page of the TOC.
+The `after-toc` value is primarily intended to be used with the book doctype, though it can be used with the article doctype if the title page is enabled.
+If the TOC is not enabled, or the title page for an article is not enabled, this value reverts to `body`.
+
 body:: This is the default value.
-The running content starts on the first page of the document body, which is typically the first page after the TOC if the TOC is in its default location.
-If `media=prepress`, this keyword refers to the first recto page after the last page of the TOC (it will skip an empty verso page).
+The running content starts on the first page of the document body.
+If the TOC is enabled in its default location, the first page of the document body is the next recto page after the TOC when `media=prepress` or the page that immediately follows the TOC otherwise.
+
 title:: The running content starts on the title page.
 The `title` value is only recognized if the title page is implicitly or explicitly enabled.
-toc:: The running content starts on the first page of the TOC.
-The `toc` value only applies if the TOC is in the default location (before the first page of the body).
-If the value is `toc`, and the toc macro is used to position the TOC, the `start-at` behavior is the same as if the TOC is not enabled.
-The `toc` value is only recognized if the title page is implicitly or explicitly enabled.
+
+toc:: The running content starts on the first page of the TOC, no matter where the TOC is placed in the document.
+The `toc` value is primarily intended to be used with the book doctype, though it can be used with the article doctype if the title page is enabled.
+If the TOC is not enabled, or the title page for an article is not enabled, this value reverts to `body`.
+
 [[page]]Integer:: The running content starts on the page that matches the number assigned to `start-at` (i.e., 1 is the first body page, 2 is the second body page).
+Refer to the `body` value for an explanation of where the body starts.
 For a prepress book (`doctype=book` and `media=prepress`), the running content starts on the empty verso page before the body if the value is `0`.
-When an integer is assigned to `start-at`, the title page doesn't need to be enabled.
+An integer value is recognized by all doctypes and the title page doesn't need to be enabled.
 
 TIP: To turn off the running content on TOC pages inserted by the toc macro, set the `noheader` or `nofooter` options on the macro (e.g., `toc::[opts=nofooter]`).
 
diff --git a/docs/modules/theme/pages/page-numbers.adoc b/docs/modules/theme/pages/page-numbers.adoc
index 2c91657da..997b97dcb 100644
--- a/docs/modules/theme/pages/page-numbers.adoc
+++ b/docs/modules/theme/pages/page-numbers.adoc
@@ -74,21 +74,27 @@ page:
 The `start-at` key accepts the following keywords or an integer:
 
 after-toc:: The integer page numbering starts after the last page of the TOC, no matter where the TOC is placed in the document.
-The `after-toc` value is only recognized if the title page is implicitly or explicitly enabled.
-If `media=prepress`, this keyword refers to the first recto page after the last page of the TOC (it will skip an empty verso page).
+If `media=prepress`, this keyword refers to the next recto page after the last page of the TOC.
+The `after-toc` value is primarily intended to be used with the book doctype, though it can be used with the article doctype if the title page is enabled.
+If the TOC is not enabled, or the title page for an article is not enabled, this value reverts to `body`.
+
 body:: This is the default value.
-The integer page numbering starts on the first page of the document body, which is typically the first page after the TOC if the TOC is in its default location.
-If `media=prepress`, this keyword refers to the first recto page after the last page of the TOC (it will skip an empty verso page).
+The integer page numbering starts on the first page of the document body.
+If the TOC is enabled in its default location, the first page of the document body is the next recto page after the TOC when `media=prepress` or the page that immediately follows the TOC otherwise.
+
 cover:: The integer page numbering starts on the front cover page of the document.
 The `cover` value is only recognized if the document has a front cover page (i.e., `front-cover-image`).
+
 title:: The integer page numbering starts on the title page.
 The `title` value is only recognized if the title page is implicitly or explicitly enabled.
-toc:: The integer page numbering starts on the first page of the TOC.
-The `toc` value only applies if the TOC is in the default location (before the first page of the body).
-If the value is `toc`, and the toc macro is used to position the TOC, the `start-at` behavior is the same as if the TOC is not enabled.
-The `toc` value is only recognized if the title page is implicitly or explicitly enabled.
+
+toc:: The integer page numbering starts on the first page of the TOC, no matter where the TOC is placed in the document.
+The `toc` value is primarily intended to be used with the book doctype, though it can be used with the article doctype if the title page is enabled.
+If the TOC is not enabled, or the title page for an article is not enabled, this value reverts to `body`.
+
 Integer:: The page numbering starts at the specified page of the body (i.e., 1 is the first body page, 2 is the second body page).
-For a prepress book (`doctype=book` and `media=prepress`), the page numbering starts on the empty verso page before the body if the value is `0`.
+Refer to the `body` value for an explanation of where the body starts.
+For a prepress book (`doctype=book` and `media=prepress`), the page numbering starts on the verso page before the body if the value is `0`.
 An integer value is recognized by all doctypes and the title page doesn't need to be enabled.
 
 Let's start the integer page number on the title page of a document.
diff --git a/lib/asciidoctor/pdf/converter.rb b/lib/asciidoctor/pdf/converter.rb
index f29fd3e18..a11a69106 100644
--- a/lib/asciidoctor/pdf/converter.rb
+++ b/lib/asciidoctor/pdf/converter.rb
@@ -169,7 +169,7 @@ def convert_document doc
         ink_cover_page doc, :front
         has_front_cover = page_number > marked_page_number
         doctype = doc.doctype
-        if (has_title_page = (title_page_on = doctype == 'book' || (doc.attr? 'title-page')) && (start_title_page doc))
+        if (has_title_page = (title_as_page = doctype == 'book' || (doc.attr? 'title-page')) && (start_title_page doc))
           # NOTE: the base font must be set before any content is written to the main or scratch document
           font @theme.base_font_family, size: @root_font_size, style: @theme.base_font_style
           if perform_on_single_page { ink_title_page doc }
@@ -184,7 +184,7 @@ def convert_document doc
           font @theme.base_font_family, size: @root_font_size, style: @theme.base_font_style
         end
 
-        unless title_page_on
+        unless title_as_page
           body_start_page_number = page_number
           theme_font :heading, level: 1 do
             ink_general_heading doc, doc.doctitle, align: (@theme.heading_h1_text_align&.to_sym || :center), level: 1, role: :doctitle
@@ -195,10 +195,10 @@ def convert_document doc
 
         indent_section do
           toc_num_levels = (doc.attr 'toclevels', 2).to_i
-          if (insert_toc = (doc.attr? 'toc') && !((toc_placement = doc.attr 'toc-placement') == 'macro' || toc_placement == 'preamble') && !(get_entries_for_toc doc).empty?)
+          if (toc_at_top = (doc.attr? 'toc') && !((toc_placement = doc.attr 'toc-placement') == 'macro' || toc_placement == 'preamble') && !(get_entries_for_toc doc).empty?)
             start_new_page if @ppbook && verso_page?
             add_dest_for_block doc, id: 'toc', y: (at_page_top? ? page_height : nil)
-            @toc_extent = allocate_toc doc, toc_num_levels, cursor, (title_page_on && theme.toc_break_after != 'auto')
+            @toc_extent = allocate_toc doc, toc_num_levels, cursor, (title_as_page && theme.toc_break_after != 'auto')
           else
             @toc_extent = nil
           end
@@ -210,7 +210,7 @@ def convert_document doc
             min_start_at = 1
           end
 
-          if title_page_on
+          if title_as_page
             zero_page_offset = has_front_cover ? 1 : 0
             first_page_offset = has_title_page ? zero_page_offset.next : zero_page_offset
             body_offset = (body_start_page_number = page_number) - 1
@@ -223,8 +223,10 @@ def convert_document doc
               when 'title'
                 running_content_start_at = 'toc' unless has_title_page
               when 'toc'
-                running_content_start_at = 'body' unless insert_toc
+                uses_start_at_toc = true
+                running_content_start_at = 'body' unless toc_at_top
               when 'after-toc'
+                uses_start_at_after_toc = true
                 running_content_start_at = 'body'
               end
             end
@@ -243,8 +245,10 @@ def convert_document doc
               when 'title'
                 page_numbering_start_at = 'toc' unless has_title_page
               when 'toc'
-                page_numbering_start_at = 'body' unless insert_toc
+                uses_start_at_toc = true
+                page_numbering_start_at = 'body' unless toc_at_top
               when 'after-toc'
+                uses_start_at_after_toc = true
                 page_numbering_start_at = 'body'
               end
             end
@@ -298,12 +302,17 @@ def convert_document doc
           end
 
           if (toc_extent = @toc_extent)
-            if title_page_on && !insert_toc
-              if @theme.running_content_start_at == 'after-toc' || @theme.page_numbering_start_at == 'after-toc' # rubocop:disable Style/SoleNestedConditional
-                last_toc_page = toc_extent.to.page
-                last_toc_page += 1 if @ppbook && (recto_page? last_toc_page)
-                num_front_matter_pages[0] = last_toc_page if @theme.running_content_start_at == 'after-toc'
-                num_front_matter_pages[1] = last_toc_page if @theme.page_numbering_start_at == 'after-toc'
+            if title_as_page && !toc_at_top && (uses_start_at_toc || uses_start_at_after_toc)
+              if uses_start_at_toc
+                toc_offset = toc_extent.from.page - 1
+                num_front_matter_pages[0] = toc_offset if @theme.running_content_start_at == 'toc'
+                num_front_matter_pages[1] = toc_offset if @theme.page_numbering_start_at == 'toc'
+              end
+              if uses_start_at_after_toc
+                after_toc_offset = toc_extent.to.page
+                after_toc_offset += 1 if @ppbook && (recto_page? after_toc_offset)
+                num_front_matter_pages[0] = after_toc_offset if @theme.running_content_start_at == 'after-toc'
+                num_front_matter_pages[1] = after_toc_offset if @theme.page_numbering_start_at == 'after-toc'
               end
             end
             toc_page_nums = ink_toc doc, toc_num_levels, toc_extent.from.page, toc_extent.from.cursor, num_front_matter_pages[1]
@@ -2384,11 +2393,15 @@ def convert_toc node, opts = {}
         if ((doc = node.document).attr? 'toc-placement', placement) && (doc.attr? 'toc') && !(get_entries_for_toc doc).empty?
           start_toc_page node, placement if (is_book = doc.doctype == 'book')
           add_dest_for_block node, id: (node.id || 'toc') if is_macro
-          toc_extent = @toc_extent = allocate_toc doc, (doc.attr 'toclevels', 2).to_i, cursor, (title_page_on = is_book || (doc.attr? 'title-page'))
-          if title_page_on && @theme.page_numbering_start_at == 'after-toc'
-            new_start_page_number = toc_extent.to.page + 1
-            new_start_page_number += 1 if @ppbook && (verso_page? new_start_page_number)
-            @index.start_page_number = new_start_page_number
+          toc_extent = @toc_extent = allocate_toc doc, (doc.attr 'toclevels', 2).to_i, cursor, (title_as_page = is_book || (doc.attr? 'title-page'))
+          if title_as_page
+            if @theme.page_numbering_start_at == 'toc'
+              @index.start_page_number = toc_extent.from.page
+            elsif @theme.page_numbering_start_at == 'after-toc'
+              new_start_page_number = toc_extent.to.page + 1
+              new_start_page_number += 1 if @ppbook && (verso_page? new_start_page_number)
+              @index.start_page_number = new_start_page_number
+            end
           end
           if is_macro
             @disable_running_content[:header] += toc_extent.page_range if node.option? 'noheader'
diff --git a/spec/index_spec.rb b/spec/index_spec.rb
index e3889a311..88479fc41 100644
--- a/spec/index_spec.rb
+++ b/spec/index_spec.rb
@@ -525,6 +525,38 @@
     (expect terms).to eql %w(anchor AsciiDoc Asciidoctor authoring)
   end
 
+  it 'should adjust start page number for prepress book when page numbering starts at toc and toc macro is used' do
+    pdf = to_pdf <<~'END', pdf_theme: { page_numbering_start_at: 'toc' }, analyze: true
+    = Book Title
+    :doctype: book
+    :media: prepress
+    :toc: macro
+
+    [dedication]
+    = Dedication
+
+    Credit where credit is due.
+
+    toc::[]
+
+    == First Chapter
+
+    ((apples))
+
+    == Second Chapter
+
+    ((bananas))
+
+    [index]
+    == Index
+    END
+
+    index_title_text = (pdf.find_text 'Index')[-1]
+    (expect index_title_text[:page_number]).to be 11
+    index_lines = pdf.lines pdf.find_text page_number: 11
+    (expect index_lines).to eql ['Index', 'A', 'apples, 3', 'B', 'bananas, 5']
+  end
+
   it 'should adjust start page number for prepress book when page numbering starts after-toc and toc macro is used' do
     pdf = to_pdf <<~'END', pdf_theme: { page_numbering_start_at: 'after-toc' }, analyze: true
     = Book Title
diff --git a/spec/running_content_spec.rb b/spec/running_content_spec.rb
index 205a9df8d..9b34f6aaa 100644
--- a/spec/running_content_spec.rb
+++ b/spec/running_content_spec.rb
@@ -403,6 +403,27 @@
       (expect pgnum_labels).to eql [nil, nil, '1', '2', '3']
     end
 
+    it 'should start running content at toc in body of book when start at is toc and macro toc is used' do
+      pdf = to_pdf <<~END, pdf_theme: { running_content_start_at: 'toc' }, enable_footer: true, analyze: true
+      = Document Title
+      :doctype: book
+      :toc: macro
+
+      == First Chapter
+
+      toc::[]
+
+      == Second Chapter
+
+      == Third Chapter
+      END
+
+      pgnum_labels = (1.upto pdf.pages.size).each_with_object [] do |page_number, accum|
+        accum << ((pdf.find_text page_number: page_number, y: 14.263)[-1] || {})[:string]
+      end
+      (expect pgnum_labels.slice 0, 5).to eql [nil, nil, '2', '3', '4']
+    end
+
     it 'should start running content after toc in body of book when start at is after-toc and macro toc is used' do
       filler = (1..20).map {|it| %(== #{['Filler'] * 20 * ' '} #{it}\n\ncontent) }.join %(\n\n)
       pdf = to_pdf <<~END, pdf_theme: { running_content_start_at: 'after-toc' }, enable_footer: true, analyze: true
@@ -786,6 +807,33 @@
       (expect pgnum_labels).to eql [nil, '1', '2', '3']
     end
 
+    it 'should start page numbering at toc in body of book when start at is toc and toc macro is used' do
+      pdf = to_pdf <<~END, enable_footer: true, pdf_theme: { page_numbering_start_at: 'toc' }, analyze: true
+      = Book Title
+      :doctype: book
+      :toc: macro
+
+      == Dedication
+
+      To the only person who gets me.
+
+      toc::[]
+
+      == Acknowledgements
+
+      Thanks all to all who made this possible!
+
+      == Chapter One
+
+      content
+      END
+
+      pgnum_labels = (1.upto pdf.pages.size).each_with_object [] do |page_number, accum|
+        accum << ((pdf.find_text page_number: page_number, y: 14.263)[-1] || {})[:string]
+      end
+      (expect pgnum_labels.slice 0, 5).to eql [nil, 'ii', '1', '2', '3']
+    end
+
     it 'should start page numbering after toc in body of book when start at is after-toc and toc macro is used' do
       filler = (1..20).map {|it| %(== #{['Filler'] * 20 * ' '} #{it}\n\ncontent) }.join %(\n\n)
       pdf = to_pdf <<~END, enable_footer: true, pdf_theme: { page_numbering_start_at: 'after-toc' }, analyze: true