:root {
  color-scheme: light;

  /* --- raw palette ------------------------------------------------------ */
  --bg: #faf4e6;
  --bg-alt: #f1e8d3;
  --card: #ffffff;
  --ink: #202b24;
  --ink-soft: #4d5a50;
  --ink-faint: #7c8a7e;
  --amber: #d98a3d;
  --amber-dark: #b56c26;
  --border: #e5d9ba;
  --border-soft: #ece2ca;
  --shadow: 0 12px 30px -18px rgba(14, 51, 47, 0.35);

  /* --- semantic roles ---------------------------------------------------
     Everything below is what the rest of the sheet actually references, so
     the dark theme only has to restate these. Nothing outside this block
     should hard-code a colour. */
  --heading: #0e332f;        /* h1-h4, and headline text that reads as a title */
  --link: #175e56;           /* body links, the hero role line */
  --link-hover: var(--amber-dark);
  --teal-ink: #123e3a;       /* teal text/strokes on page or card surfaces */
  --accent: #123e3a;         /* filled teal: buttons, hover states */
  --accent-strong: #0e332f;  /* filled teal, one step darker */
  --on-accent: #ffffff;      /* text on --accent */
  --on-amber: #ffffff;       /* text on a filled amber button */
  --badge-bg: #dcece7;       /* pill and icon-badge backgrounds */
  --surface-dark: #0e332f;   /* header, footer, callout, back-to-top */
  --on-dark: #f3ede0;
  --on-dark-soft: #cfe0d8;
  --on-dark-faint: #9fb6ab;
  --crest-bg: #ffffff;       /* behind the university crests, which are dark ink */
  --tree-ink: #227a6f;       /* phylogeny branches and the decorative specimens */

  --radius-lg: 22px;
  --radius-md: 14px;
  --radius-sm: 8px;
  --font-head: "Fraunces", "Iowan Old Style", "Palatino Linotype", serif;
  --font-body: "Work Sans", -apple-system, BlinkMacSystemFont, "Segoe UI", sans-serif;
  --max: 1080px;
}

/* ---------- dark theme ----------------------------------------------------
   Two selectors, one palette: the media query is the default for anyone whose
   OS asks for dark (unless they have explicitly chosen light here), and the
   attribute selector is the in-page toggle. Keep the two blocks identical. */
@media (prefers-color-scheme: dark) {
  :root:not([data-theme="light"]) {
  color-scheme: dark;

  --bg: #101916;
  --bg-alt: #16211d;
  --card: #18241f;
  --ink: #e9efea;
  --ink-soft: #b6c5bb;
  --ink-faint: #8b9c92;
  --amber: #d9944d;
  --amber-dark: #e6b077;
  --border: #2c3b35;
  --border-soft: #25322d;
  --shadow: 0 12px 30px -18px rgba(0, 0, 0, 0.75);

  --heading: #eaf2ed;
  --link: #7fd4c2;
  --link-hover: var(--amber-dark);
  --teal-ink: #8fd8c8;
  --accent: #1e6f63;
  --accent-strong: #268275;
  --on-accent: #f2f8f5;
  --on-amber: #14201c;
  --badge-bg: #1c322d;
  --surface-dark: #0a1310;
  --on-dark: #e9efea;
  --on-dark-soft: #b6c5bb;
  --on-dark-faint: #8b9c92;
  --crest-bg: #f2efe6;
  --tree-ink: #4a9d8f;
  }
}

:root[data-theme="dark"] {
  color-scheme: dark;

  --bg: #101916;
  --bg-alt: #16211d;
  --card: #18241f;
  --ink: #e9efea;
  --ink-soft: #b6c5bb;
  --ink-faint: #8b9c92;
  --amber: #d9944d;
  --amber-dark: #e6b077;
  --border: #2c3b35;
  --border-soft: #25322d;
  --shadow: 0 12px 30px -18px rgba(0, 0, 0, 0.75);

  --heading: #eaf2ed;
  --link: #7fd4c2;
  --link-hover: var(--amber-dark);
  --teal-ink: #8fd8c8;
  --accent: #1e6f63;
  --accent-strong: #268275;
  --on-accent: #f2f8f5;
  --on-amber: #14201c;
  --badge-bg: #1c322d;
  --surface-dark: #0a1310;
  --on-dark: #e9efea;
  --on-dark-soft: #b6c5bb;
  --on-dark-faint: #8b9c92;
  --crest-bg: #f2efe6;
  --tree-ink: #4a9d8f;
}

* { box-sizing: border-box; }

html { scroll-behavior: smooth; }

body {
  margin: 0;
  background: var(--bg);
  color: var(--ink);
  font-family: var(--font-body);
  font-size: 17px;
  line-height: 1.65;
  -webkit-font-smoothing: antialiased;
  position: relative;
  overflow-x: hidden;
}

/* ---------- beetle rail: decorative specimens down the left margin ---------- */
.beetle-rail {
  display: none;
  position: absolute;
  top: 0;
  left: 20px;
  width: 72px;
  height: 100%;
  z-index: 0;
  pointer-events: none;
}
.beetle-rail span {
  position: absolute;
  color: var(--tree-ink);
}
.beetle-rail span.amber { color: var(--amber-dark); }
.beetle-rail svg { width: 100%; height: 100%; }
@media (min-width: 1360px) {
  .beetle-rail { display: block; }
}

img { max-width: 100%; display: block; }

a {
  color: var(--link);
  text-decoration: none;
}
a:hover { color: var(--link-hover); }

h1, h2, h3, h4 {
  font-family: var(--font-head);
  color: var(--heading);
  line-height: 1.15;
  margin: 0 0 0.5em;
  font-weight: 600;
}

h1 { font-size: clamp(2.2rem, 4vw, 3.1rem); }
h2 { font-size: clamp(1.5rem, 3vw, 2rem); }
h3 { font-size: 1.3rem; }

p { margin: 0 0 1.1em; color: var(--ink-soft); }

.container {
  width: 100%;
  max-width: var(--max);
  margin: 0 auto;
  padding: 0 24px;
}

section { padding: 64px 0; }
section + section { border-top: 1px solid var(--border-soft); }

.eyebrow {
  display: inline-block;
  font-family: var(--font-body);
  font-weight: 600;
  letter-spacing: 0.12em;
  text-transform: uppercase;
  font-size: 0.78rem;
  color: var(--amber-dark);
  margin-bottom: 0.6em;
}

.lede {
  font-size: 1.15rem;
  color: var(--ink-soft);
  max-width: 62ch;
}

/* ---------- buttons ---------- */
.btn {
  display: inline-flex;
  align-items: center;
  gap: 0.5em;
  font-family: var(--font-body);
  font-weight: 600;
  font-size: 0.95rem;
  padding: 0.7em 1.4em;
  border-radius: 999px;
  border: 1.5px solid var(--teal-ink);
  color: var(--teal-ink);
  background: transparent;
  transition: transform 0.15s ease, background 0.15s ease, color 0.15s ease;
}
.btn:hover { background: var(--accent); color: var(--on-accent); transform: translateY(-1px); }
.btn.primary { background: var(--accent); color: var(--on-accent); }
.btn.primary:hover { background: var(--accent-strong); color: var(--on-accent); }
.btn.amber { border-color: var(--amber); color: var(--amber-dark); }
.btn.amber:hover { background: var(--amber); color: var(--on-amber); }

/* ---------- header / nav ---------- */
.site-header {
  background: var(--surface-dark);
  color: var(--on-dark);
  position: relative;
  z-index: 5;
}
.nav-row {
  display: flex;
  align-items: center;
  justify-content: space-between;
  padding: 20px 24px;
  max-width: var(--max);
  margin: 0 auto;
}
.brand {
  display: flex;
  align-items: center;
  gap: 10px;
  color: var(--on-dark);
  font-family: var(--font-head);
  font-size: 1.35rem;
  font-weight: 600;
}
.brand:hover { color: var(--on-dark); }
.brand svg { flex-shrink: 0; }

.nav-toggle {
  display: none;
  background: none;
  border: 1px solid rgba(255,255,255,0.35);
  border-radius: 8px;
  color: var(--on-dark);
  padding: 8px 10px;
  font-size: 1rem;
}

.nav-links {
  display: flex;
  align-items: center;
  gap: 28px;
  list-style: none;
  margin: 0;
  padding: 0;
}
.nav-links a {
  color: var(--on-dark-soft);
  font-weight: 500;
  font-size: 0.97rem;
  padding-bottom: 3px;
  border-bottom: 2px solid transparent;
}
.nav-links a:hover,
.nav-links a.active {
  color: var(--on-dark);
  border-bottom-color: var(--amber);
}
.nav-social { display: flex; align-items: center; gap: 14px; margin-left: 6px; }

/* ---------- theme toggle ---------- */
.theme-toggle {
  display: inline-flex;
  align-items: center;
  justify-content: center;
  width: 34px;
  height: 34px;
  padding: 0;
  border: 1px solid rgba(255, 255, 255, 0.28);
  border-radius: 50%;
  background: transparent;
  color: var(--on-dark-soft);
  cursor: pointer;
  transition: color 0.15s ease, border-color 0.15s ease;
}
.theme-toggle:hover { color: var(--amber); border-color: var(--amber); }
.theme-toggle svg { width: 17px; height: 17px; display: block; }
/* show the icon for the theme you would switch TO */
.theme-toggle .icon-sun { display: none; }
.theme-toggle .icon-moon { display: block; }
@media (prefers-color-scheme: dark) {
  :root:not([data-theme="light"]) .theme-toggle .icon-sun { display: block; }
  :root:not([data-theme="light"]) .theme-toggle .icon-moon { display: none; }
}
:root[data-theme="dark"] .theme-toggle .icon-sun { display: block; }
:root[data-theme="dark"] .theme-toggle .icon-moon { display: none; }
:root[data-theme="light"] .theme-toggle .icon-sun { display: none; }
:root[data-theme="light"] .theme-toggle .icon-moon { display: block; }
.nav-social a { color: var(--on-dark-soft); display: inline-flex; }
.nav-social a:hover { color: var(--amber); }

@media (max-width: 820px) {
  .nav-toggle { display: inline-flex; align-items: center; }
  .nav-links {
    display: none;
    position: absolute;
    top: 76px;
    left: 0;
    right: 0;
    background: var(--surface-dark);
    flex-direction: column;
    align-items: flex-start;
    gap: 4px;
    padding: 8px 24px 24px;
    z-index: 40;
  }
  .nav-links.open { display: flex; }
  .nav-links a { width: 100%; padding: 10px 0; }
  .nav-social { margin: 8px 0 0; }
  .site-header { position: relative; }
}

/* ---------- hero ---------- */
.hero {
  padding: 72px 0 56px;
  position: relative;
  overflow: hidden;
}
.hero .container {
  position: relative;
  z-index: 1;
  display: grid;
  grid-template-columns: 0.95fr 1.05fr;
  gap: 40px;
  align-items: center;
}
.hero-copy h1 { margin-bottom: 0.25em; }
.hero-copy .role {
  font-family: var(--font-head);
  font-style: italic;
  font-size: 1.25rem;
  color: var(--link);
  margin-bottom: 0.9em;
}
.hero-actions {
  display: flex;
  flex-wrap: wrap;
  gap: 12px;
  margin-top: 1.6em;
}

/* headline credentials, sat directly under the name */
.credentials {
  list-style: none;
  margin: 0 0 1.4em;
  padding: 0;
  display: grid;
  gap: 7px;
}
.credentials li {
  display: grid;
  grid-template-columns: 20px 1fr;
  align-items: start;
  gap: 10px;
  font-size: 0.95rem;
  color: var(--ink-soft);
}
.credentials li svg {
  width: 18px;
  height: 18px;
  margin-top: 3px;
  color: var(--link);
  flex-shrink: 0;
}
.credentials li:first-child span { font-weight: 600; color: var(--heading); }

/* ---------- tree-nav: the phylogeny-as-navigation hero centerpiece ---------- */
.tree-nav {
  position: relative;
  width: min(480px, 100%);
  aspect-ratio: 1 / 1;
  margin: 0 auto;
}
.tree-nav .hero-tree {
  position: absolute;
  inset: 0;
  width: 100%;
  height: 100%;
  color: var(--tree-ink);
  opacity: 0.85;
}
.tree-nav .tree-portrait {
  position: absolute;
  left: 50%;
  top: 50%;
  transform: translate(-50%, -50%);
  width: 30%;
  aspect-ratio: 1 / 1;
  border-radius: 50%;
  overflow: hidden;
  border: 5px solid var(--card);
  box-shadow: var(--shadow);
  z-index: 2;
}
.tree-nav .tree-portrait img {
  width: 100%;
  height: 100%;
  object-fit: cover;
}
.tree-tip {
  position: absolute;
  transform: translate(-50%, -50%);
  z-index: 3;
  display: inline-flex;
  align-items: center;
  gap: 6px;
  white-space: nowrap;
  font-family: var(--font-body);
  font-weight: 600;
  font-size: 0.85rem;
  color: var(--teal-ink);
  background: var(--card);
  border: 1.5px solid var(--border);
  border-radius: 999px;
  padding: 6px 14px 6px 10px;
  box-shadow: 0 6px 16px -10px rgba(14, 51, 47, 0.4);
  transition: transform 0.15s ease, background 0.15s ease, color 0.15s ease;
}
.tree-tip svg { width: 15px; height: 15px; flex-shrink: 0; }
.tree-tip:hover {
  background: var(--accent);
  color: var(--on-accent);
  border-color: var(--accent);
  transform: translate(-50%, -50%) scale(1.06);
}
@media (max-width: 520px) {
  .tree-tip { font-size: 0.72rem; padding: 4px 9px; }
}

.floating-icon {
  position: absolute;
  transform: translate(-50%, -50%);
  color: var(--tree-ink);
  opacity: 0.62;
  z-index: 1;
  pointer-events: none;
  display: block;
}
.floating-icon svg { width: 100%; height: auto; display: block; }
.floating-icon.wide { width: 54px; }
.floating-icon.tall { width: 32px; }
.floating-icon.amber { color: var(--amber-dark); }
@media (max-width: 620px) {
  .floating-icon.edge { display: none; }
}
@media (max-width: 520px) {
  .floating-icon { opacity: 0.38; }
  .floating-icon.wide { width: 38px; }
  .floating-icon.tall { width: 20px; }
}

/* ---------- research cards ---------- */
.research-card p.specimen-system {
  font-size: 0.82rem;
  letter-spacing: 0.04em;
  color: var(--amber-dark);
  margin-bottom: 0.9em;
}

/* ---------- study-systems cladogram ---------- */
.cladogram-figure { margin: 0; }
.cladogram-scroll { overflow-x: auto; padding-bottom: 4px; }
.cladogram {
  display: grid;
  grid-template-columns: minmax(170px, 0.85fr) minmax(330px, 1.15fr);
  min-width: 600px;
  color: var(--tree-ink);
}
.clado-tree { position: relative; }
.clado-tree svg { width: 100%; height: 100%; display: block; }
.clade {
  position: absolute;
  transform: translate(7px, -135%);
  font-size: 0.68rem;
  font-weight: 700;
  letter-spacing: 0.1em;
  text-transform: uppercase;
  color: var(--ink-faint);
  white-space: nowrap;
}
.clado-tips {
  list-style: none;
  margin: 0;
  padding: 0;
  display: grid;
  grid-auto-rows: 1fr; /* equal rows so tip centres line up with the branches */
}
.clado-tips li {
  display: flex;
  align-items: center;
  gap: 16px;
  padding: 8px 0 8px 12px;
  min-height: 78px;
}
.clado-specimen {
  width: 66px;
  flex-shrink: 0;
  display: flex;
  align-items: center;
  justify-content: center;
  color: var(--teal-ink);
}
.clado-specimen svg { width: 100%; height: auto; max-height: 58px; }
.clado-label {
  font-size: 0.97rem;
  font-weight: 600;
  color: var(--heading);
  line-height: 1.3;
}
.clado-label small {
  display: block;
  margin-top: 2px;
  font-weight: 400;
  font-size: 0.82rem;
  color: var(--ink-soft);
}
.cladogram-figure figcaption {
  margin-top: 16px;
  font-size: 0.85rem;
  font-style: italic;
  color: var(--ink-faint);
  max-width: 62ch;
}
@media (max-width: 700px) {
  .clade { display: none; }
}

@media (max-width: 860px) {
  .hero .container { grid-template-columns: 1fr; }
  .hero-art { order: -1; }
}

/* ---------- section headers ---------- */
.section-head {
  display: flex;
  align-items: flex-end;
  justify-content: space-between;
  gap: 24px;
  margin-bottom: 32px;
  flex-wrap: wrap;
}
.section-head .icon-badge { margin-bottom: 14px; }
.section-head h2 { margin-bottom: 0; }
.section-head .lede { margin: 0.55em 0 0; }
/* a second heading inside a section: same scale, just more air above it */
.section-head.subhead { margin-top: 52px; }

.icon-badge {
  width: 46px;
  height: 46px;
  border-radius: 12px;
  background: var(--badge-bg);
  color: var(--teal-ink);
  display: inline-flex;
  align-items: center;
  justify-content: center;
}

/* ---------- research grid ---------- */
.research-grid {
  display: grid;
  grid-template-columns: repeat(3, 1fr);
  gap: 24px;
}
.research-grid.cols-2 { grid-template-columns: repeat(2, 1fr); }
.research-grid.cols-3 { grid-template-columns: repeat(3, 1fr); }
@media (max-width: 900px) {
  .research-grid,
  .research-grid.cols-2,
  .research-grid.cols-3 { grid-template-columns: 1fr; }
}
.research-card {
  background: var(--card);
  border: 1px solid var(--border-soft);
  border-radius: var(--radius-lg);
  padding: 28px 26px;
}
.research-card .icon-badge { margin-bottom: 16px; }
.research-card h3 { margin-bottom: 0.5em; }
.research-card p { margin: 0; font-size: 0.98rem; }

/* jekyll-scholar bib.html output (publications section) */
.bibliography { list-style: none; margin: 0; padding: 0; }
.bibliography li {
  padding: 22px 0;
  border-top: 1px solid var(--border-soft);
}
.bibliography li:first-child { border-top: none; }
.bibliography .title { font-family: var(--font-head); font-size: 1.15rem; color: var(--heading); font-weight: 600; margin-bottom: 0.3em; }
.bibliography .author { color: var(--ink-soft); margin-bottom: 0.25em; }
.bibliography .author em { color: var(--teal-ink); font-style: normal; font-weight: 600; }
.bibliography .periodical { font-style: normal; color: var(--ink-faint); margin-bottom: 0.5em; }
.bibliography .periodical em { font-style: italic; }
.bibliography .periodical .vol { font-weight: 600; }
.bibliography .links { display: flex; flex-wrap: wrap; gap: 8px; }
.bibliography .links a {
  display: inline-flex;
  align-items: center;
  gap: 6px;
  font-size: 0.85rem;
  font-weight: 600;
  color: var(--link);
  border: 1px solid var(--border);
  border-radius: 999px;
  padding: 5px 13px;
}
.bibliography .links a:hover { background: var(--accent); color: var(--on-accent); border-color: var(--accent); }
.bibliography .author strong { color: var(--heading); font-weight: 700; }

/* ---------- timeline (education) ---------- */
.timeline { list-style: none; margin: 0; padding: 0; position: relative; }
.timeline::before {
  content: "";
  position: absolute;
  left: 23px;
  top: 6px;
  bottom: 6px;
  width: 2px;
  background: var(--border);
}
.timeline-item {
  position: relative;
  display: flex;
  gap: 20px;
  padding-bottom: 30px;
}
.timeline-item:last-child { padding-bottom: 0; }
.timeline-icon {
  width: 48px;
  height: 48px;
  border-radius: 50%;
  background: var(--crest-bg);
  border: 2px solid var(--border);
  display: flex;
  align-items: center;
  justify-content: center;
  flex-shrink: 0;
  z-index: 1;
  padding: 8px;
}
.timeline-icon img { width: 100%; height: 100%; object-fit: contain; }
.timeline-body h3 { margin-bottom: 0.15em; font-size: 1.1rem; }
.timeline-body .program { color: var(--link); font-style: italic; margin-bottom: 0.2em; }
.timeline-body .dates { font-size: 0.85rem; color: var(--ink-faint); text-transform: uppercase; letter-spacing: 0.05em; }

/* ---------- teaching / experience list ---------- */
.entry-list { list-style: none; margin: 0; padding: 0; }
.entry {
  padding: 24px 0;
  border-top: 1px solid var(--border-soft);
}
.entry:first-child { border-top: none; }
.entry-head {
  display: grid;
  grid-template-columns: minmax(0, 1fr) auto;
  align-items: baseline;
  gap: 8px 20px;
}
.entry-head h3 { margin-bottom: 0.1em; }
.entry-meta {
  color: var(--ink-faint);
  font-size: 0.88rem;
  white-space: nowrap;
  text-align: right;
}
.entry-list.compact .entry { padding: 14px 0; }
.entry-list.compact .entry-head h3 { font-size: 1.1rem; margin-bottom: 0; }
.entry-list.compact .entry-role { margin-bottom: 0; }
.entry-role { color: var(--amber-dark); font-weight: 600; font-size: 0.92rem; margin-bottom: 0.4em; }

/* ---------- cards / callouts ---------- */
.card {
  background: var(--card);
  border: 1px solid var(--border-soft);
  border-radius: var(--radius-lg);
  padding: 32px;
}

.callout {
  background: var(--surface-dark);
  color: var(--on-dark);
  border-radius: var(--radius-lg);
  padding: 44px;
  display: flex;
  align-items: center;
  justify-content: space-between;
  gap: 24px;
  flex-wrap: wrap;
}
.callout h2 { color: var(--on-dark); margin-bottom: 0.2em; }
.callout p { color: var(--on-dark-soft); margin: 0; }
.callout .btn { border-color: var(--amber); color: var(--amber); }
.callout .btn:hover { background: var(--amber); color: var(--on-amber); }

/* ---------- footer ---------- */
.site-footer {
  background: var(--surface-dark);
  color: var(--on-dark-soft);
  padding: 40px 0;
  margin-top: 40px;
  position: relative;
  z-index: 5;
}
.footer-row {
  display: flex;
  justify-content: space-between;
  align-items: center;
  gap: 16px;
  flex-wrap: wrap;
}
.site-footer a { color: var(--on-dark); }
.site-footer a:hover { color: var(--amber); }
.footer-social { display: flex; gap: 16px; }
.footer-note { font-size: 0.85rem; color: var(--on-dark-faint); }
.footer-meta {
  display: flex;
  align-items: center;
  justify-content: space-between;
  flex-wrap: wrap;
  gap: 10px 20px;
  margin-top: 18px;
}
.site-footer a.repo-link {
  display: inline-flex;
  align-items: center;
  gap: 7px;
  color: var(--on-dark-faint);
}
.site-footer a.repo-link:hover { color: var(--amber); }
.repo-link svg { width: 16px; height: 16px; flex-shrink: 0; }

/* ---------- 404 ---------- */
.error-page { text-align: center; padding: 100px 0; }
.error-page h1 { font-size: 5rem; }

/* ---------- misc ---------- */
.badge {
  display: inline-block;
  font-size: 0.78rem;
  font-weight: 600;
  padding: 3px 10px;
  border-radius: 999px;
  background: var(--badge-bg);
  color: var(--teal-ink);
  margin: 0 6px 6px 0;
}
.divider-icon { display: flex; justify-content: center; color: var(--border); margin: 8px 0 0; }

/* the address is assembled by JS; until then the link is inert, so don't
   advertise it as clickable */
a.js-eml:not([href^="mailto"]) { cursor: default; }
a.js-eml .eml-text { white-space: nowrap; }

/* ---------- back to top ---------- */
.to-top {
  position: fixed;
  right: 22px;
  bottom: 22px;
  z-index: 30;
  width: 44px;
  height: 44px;
  display: flex;
  align-items: center;
  justify-content: center;
  border-radius: 50%;
  background: var(--surface-dark);
  color: var(--on-dark);
  box-shadow: 0 10px 24px -12px rgba(14, 51, 47, 0.7);
  opacity: 0;
  visibility: hidden;
  transform: translateY(8px);
  transition: opacity 0.2s ease, transform 0.2s ease, visibility 0.2s;
}
.to-top.visible { opacity: 1; visibility: visible; transform: none; }
.to-top:hover { background: var(--amber-dark); color: var(--on-amber); }
@media print { .to-top { display: none !important; } }

/* ---------- accessibility ---------- */
.skip-link {
  position: absolute;
  left: -9999px;
  top: 0;
  z-index: 100;
  background: var(--card);
  color: var(--heading);
  font-weight: 600;
  padding: 12px 20px;
  border-radius: 0 0 var(--radius-sm) 0;
}
.skip-link:focus { left: 0; }

a:focus-visible,
button:focus-visible,
.tree-tip:focus-visible,
input:focus-visible {
  outline: 3px solid var(--amber);
  outline-offset: 3px;
  border-radius: 4px;
}
main:focus { outline: none; }

@media (prefers-reduced-motion: reduce) {
  html { scroll-behavior: auto; }
  *, *::before, *::after {
    animation-duration: 0.001ms !important;
    animation-iteration-count: 1 !important;
    transition-duration: 0.001ms !important;
  }
}

/* ---------- print: the page becomes a clean CV ---------- */
@media print {
  @page { margin: 16mm 14mm; }

  /* print is always the light theme, whatever the screen is set to */
  :root,
  :root[data-theme="dark"] {
    --bg: #fff;
    --bg-alt: #fff;
    --card: #fff;
    --ink: #000;
    --ink-soft: #222;
    --ink-faint: #444;
    --border: #bbb;
    --border-soft: #ccc;
    --heading: #000;
    --link: #000;
    --teal-ink: #222;
    --badge-bg: transparent;
    --crest-bg: #fff;
    --tree-ink: #444;
    --surface-dark: #fff;
    --on-dark: #000;
    --on-dark-soft: #333;
    --on-dark-faint: #444;
  }

  body {
    background: #fff;
    color: #000;
    font-size: 10.5pt;
    line-height: 1.42;
  }

  /* drop everything decorative or navigational */
  .print-hide,
  .theme-toggle,
  .beetle-rail,
  .floating-icon,
  .hero-tree,
  .tree-tip,
  .site-header,
  .skip-link,
  .hero-actions,
  .icon-badge,
  .cladogram-figure,
  .site-footer .footer-social {
    display: none !important;
  }

  .hero { padding: 0 0 10pt; overflow: visible; }
  .hero .container { display: block; }
  .tree-nav { width: 108pt; aspect-ratio: auto; float: right; margin: 0 0 8pt 12pt; }
  .tree-nav .tree-portrait {
    position: static;
    transform: none;
    width: 108pt;
    height: 108pt;
    border-width: 1pt;
    box-shadow: none;
  }

  .container { max-width: none; padding: 0; }
  section { padding: 10pt 0; border-top: 0.5pt solid #bbb; break-inside: auto; }
  section + section { border-top: 0.5pt solid #bbb; }

  h1 { font-size: 20pt; margin-bottom: 2pt; }
  h2 { font-size: 13pt; margin-bottom: 4pt; }
  h3 { font-size: 11pt; }
  h2, h3, .section-head { break-after: avoid; page-break-after: avoid; }
  p, li, .lede { color: #222; font-size: 10.5pt; }
  .lede { max-width: none; }
  .eyebrow, .entry-role, .specimen-system { color: #444; }

  .research-grid { display: block; }
  .research-card {
    border: 0;
    padding: 0 0 8pt;
    break-inside: avoid;
  }
  .entry, .bibliography li, .timeline-item { break-inside: avoid; }
  .entry { padding: 6pt 0; }

  .btn { display: none; }
  .bibliography .links a { border: 0; padding: 0; color: #444; font-weight: 400; }
  .bibliography .links svg { display: none; }

  .badge {
    background: none;
    border: 0.5pt solid #888;
    color: #222;
  }

  .site-footer { background: none; color: #000; padding: 8pt 0 0; margin: 0; border-top: 0.5pt solid #bbb; }
  .site-footer a, .footer-note { color: #333; }

  a { color: #000; text-decoration: none; }
}