Periplaneta​ ​americana

​ ​Invertebrate​ ​Anatomy
Terminology​ ​and​ ​phylogeny​ ​used​ ​in​ ​these​ ​exercises​ ​correspond​ ​to​ ​usage​ ​in​ ​the
Invertebrate​ ​Zoology​ ​textbook​ ​by​ ​Ruppert,​ ​Fox,​ ​and​ ​Barnes​ ​(2004).

Systematics
Arthropoda​ ​ ,​ ​Mandibulata​ ​ ,​ ​Tracheata,​ ​Hexapoda​ SC​
P​ sP​
​ ,​ ​Insecta​ C​​ ,​ ​Dicondylia,
Pterygota,​ ​Metapterygota,​ ​Neoptera,​ ​Blattaria​ O​​ ,​ ​Blattoidea​ SF​
​ ,​ ​Blattidae​ F​​ ,
Blattinae
Arthropoda
​ ​Arthropoda,​ ​by​ ​far​ ​the​ ​largest​ ​and​ ​most​ ​diverse​ ​animal​ ​taxon,​ ​includes
chelicerates,​ ​insects,​ ​myriapods,​ ​and​ ​crustaceans​ ​as​ ​well​ ​as​ ​many​ ​extinct​ ​taxa.​ ​ ​ ​The
body​ ​is​ ​segmented​ ​and​ ​primitively​ ​bears​ ​a​ ​pair​ ​of​ ​jointed​ ​appendages​ ​on​ ​each​ ​segment.
The​ ​epidermis​ ​secretes​ ​a​ ​complex​ ​cuticular​ ​exoskeleton​ ​which​ ​must​ ​be​ ​molted​ ​to​ ​permit
increase​ ​in​ ​size.​ ​ ​ ​Extant​ ​arthropods​ ​exhibit​ ​regional​ ​specialization​ ​in​ ​the​ ​structure​ ​and
function​ ​of​ ​segments​ ​and​ ​appendages.​ ​The​ ​body​ ​is​ ​typically​ ​divided​ ​into​ ​a​ ​head​ ​and
trunk,​ ​of​ ​which​ ​the​ ​trunk​ ​is​ ​often​ ​itself​ ​divided​ ​into​ ​thorax​ ​and​ ​abdomen.
​ ​The​ ​gut​ ​consists​ ​of​ ​foregut,​ ​midgut,​ ​and​ ​hindgut​ ​and​ ​extends​ ​the​ ​length​ ​of​ ​the
body​ ​from​ ​anterior​ ​mouth​ ​to​ ​posterior​ ​anus.​ ​ ​ ​Foregut​ ​and​ ​hindgut​ ​are​ ​epidermal
invaginations,​ ​being​ ​derived​ ​from​ ​the​ ​embryonic​ ​stomodeum​ ​and​ ​proctodeum
respectively,​ ​and​ ​are​ ​lined​ ​by​ ​cuticle,​ ​as​ ​are​ ​all​ ​epidermal​ ​surfaces.​ ​ ​ ​The​ ​midgut​ ​is
endodermal​ ​and​ ​is​ ​responsible​ ​for​ ​most​ ​enzyme​ ​secretion,​ ​hydrolysis,​ ​and​ ​absorption.
​ ​The​ ​coelom​ ​is​ ​reduced​ ​to​ ​small​ ​spaces​ ​associated​ ​with​ ​the​ ​gonads​ ​and​ ​kidney.
The​ ​functional​ ​body​ ​cavity​ ​is​ ​a​ ​spacious​ ​hemocoel​ ​divided​ ​by​ ​a​ ​horizontal​ ​diaphragm
into​ ​a​ ​dorsal​ ​pericardial​ ​sinus​ ​and​ ​a​ ​much​ ​larger​ ​perivisceral​ ​sinus.​ ​ ​ ​Sometimes​ ​there​ ​is
a​ ​small​ ​ventral​ ​perineural​ ​sinus​ ​surrounding​ ​the​ ​ventral​ ​nerve​ ​cord.
​ ​The​ ​hemal​ ​system​ ​includes​ ​a​ ​dorsal,​ ​contractile,​ ​tubular,​ ​ostiate​ ​heart​ ​that
pumps​ ​blood​ ​to​ ​and​ ​from​ ​the​ ​hemocoel.​ ​ ​ ​Excretory​ ​organs​ ​vary​ ​with​ ​taxon​ ​and​ ​include
Malpighian​ ​tubules,​ ​saccate​ ​nephridia,​ ​and​ ​nephrocytes.​ ​ ​ ​Respiratory​ ​organs​ ​also​ ​vary
with​ ​taxon​ ​and​ ​include​ ​many​ ​types​ ​of​ ​gills,​ ​book​ ​lungs,​ ​and​ ​tracheae.
​ ​The​ ​nervous​ ​system​ ​consists​ ​of​ ​a​ ​dorsal,​ ​anterior​ ​brain​ ​of​ ​two​ ​or​ ​three​ ​pairs​ ​of
ganglia,​ ​circumenteric​ ​connectives,​ ​and​ ​a​ ​paired​ ​ventral​ ​nerve​ ​cord​ ​with​ ​segmental
ganglia​ ​and​ ​segmental​ ​peripheral​ ​nerves.​ ​ ​ ​Various​ ​degrees​ ​of​ ​condensation​ ​and
cephalization​ ​are​ ​found​ ​in​ ​different​ ​taxa.
​ ​Development​ ​is​ ​derived​ ​with​ ​centrolecithal​ ​eggs​ ​and​ ​superficial​ ​cleavage.
There​ ​is​ ​frequently​ ​a​ ​larva​ ​although​ ​development​ ​is​ ​direct​ ​in​ ​many.​ ​Juveniles​ ​pass
through​ ​a​ ​series​ ​of​ ​instars​ ​separated​ ​by​ ​molts​ ​until​ ​reaching​ ​the​ ​adult​ ​size​ ​and
reproductive​ ​condition.​ ​ ​ ​At​ ​this​ ​time​ ​molting​ ​and​ ​growth​ ​may​ ​cease​ ​or​ ​continue,
depending​ ​on​ ​taxon.
Mandibulata
​ ​Mandibulata​ ​includes​ ​arthropods​ ​in​ ​which​ ​the​ ​third​ ​head​ ​segment​ ​bears​ ​a​ ​pair​ ​of
mandibles.​ ​ ​ ​As​ ​currently​ ​conceived​ ​this​ ​taxon​ ​includes​ ​myriapods,​ ​hexapods,​ ​and

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crustaceans.​ ​ ​ ​ ​Appendages​ ​may​ ​be​ ​uni-​ ​or​ ​biramous​ ​and​ ​habitats​ ​include​ ​marine,
freshwater,​ ​terrestrial,​ ​and​ ​aerial.
Tracheata
​ ​Myriapods​ ​and​ ​hexapods​ ​share​ ​tracheae​ ​and​ ​a​ ​single​ ​pair​ ​of​ ​antennae​ ​and​ ​are
sister​ ​taxa​ ​in​ ​Tracheata.​ ​ ​ ​Crustaceans,​ ​which​ ​have​ ​gills​ ​and​ ​lack​ ​tracheae,​ ​are
excluded​ ​and​ ​form​ ​the​ ​sister​ ​group.
Hexapoda
​ ​The​ ​body​ ​is​ ​divided​ ​into​ ​three​ ​tagmata;​ ​head,​ ​thorax,​ ​and​ ​abdomen.
Appendages​ ​are​ ​uniramous​ ​and​ ​a​ ​single​ ​pair​ ​of​ ​antennae​ ​is​ ​present.​ ​ ​ ​Three​ ​pairs​ ​of
legs​ ​and​ ​two​ ​pairs​ ​of​ ​wings​ ​are​ ​found​ ​on​ ​the​ ​thorax​ ​of​ ​most​ ​adults.​ ​ ​ ​Hexapod​ ​legs​ ​are
uniramous​ ​although​ ​there​ ​is​ ​increasing​ ​evidence​ ​that​ ​they​ ​evolved​ ​from​ ​multiramous
appendages​ ​of​ ​their​ ​ancestors.​ ​ ​ ​Gas​ ​exchange​ ​is​ ​accomplished​ ​by​ ​trachea.​ ​ ​ ​Excretory
organs​ ​are​ ​Malpighian​ ​tubules​ ​and​ ​the​ ​end​ ​product​ ​of​ ​nitrogen​ ​metabolism​ ​is​ ​uric​ ​acid.
There​ ​is​ ​relatively​ ​little​ ​cephalization​ ​of​ ​the​ ​nervous​ ​system.​ ​Insects​ ​are​ ​gonochoric​ ​with
copulation​ ​and​ ​internal​ ​fertilization.
Insecta
​ ​Most​ ​hexapods​ ​are​ ​insects.​ ​ ​ ​A​ ​few​ ​hexapod​ ​taxa​ ​(orders)​ ​lack​ ​wings​ ​and​ ​have
primitive​ ​mouthparts​ ​recessed​ ​into​ ​the​ ​head​ ​and​ ​belong​ ​to​ ​Entognatha,​ ​the​ ​sister​ ​taxon
of​ ​Insecta.​ ​ ​ ​Insects​ ​have​ ​ectognath​ ​mouthparts​ ​and​ ​the​ ​adults​ ​(imagoes)​ ​of​ ​most​ ​taxa
have​ ​wings.
Pterygota
​ ​Pterygotes​ ​are​ ​the​ ​winged​ ​insects.​ ​These​ ​insects​ ​are​ ​derived​ ​from​ ​a​ ​winged
common​ ​ancestor.​ ​ ​ ​Adults​ ​of​ ​most​ ​taxa​ ​have​ ​wings​ ​although​ ​they​ ​have​ ​been​ ​lost​ ​in
some.
Blattaria​ O​
​ ​Cockroaches​ ​are​ ​fast​ ​running​ ​insects​ ​probably​ ​closely​ ​related​ ​to​ ​termites
(Isoptera).​ ​Some​ ​primitive​ ​taxa​ ​have​ ​endosymbiotic​ ​gut​ ​zooflagellates​ ​similar​ ​to​ ​those​ ​of
termites.​ ​The​ ​tarsi​ ​are​ ​five-articulate.​ ​The​ ​oval​ ​body​ ​is​ ​dorsoventrally​ ​depressed​ ​and
most​ ​the​ ​head​ ​is​ ​hidden​ ​from​ ​dorsal​ ​view​ ​by​ ​the​ ​large​ ​pronotum.​ ​ ​ ​The​ ​antennae​ ​are
long​ ​and​ ​filamentous.​ ​Most​ ​taxa​ ​have​ ​wings,​ ​in​ ​at​ ​least​ ​one​ ​sex,​ ​but​ ​they​ ​are​ ​not​ ​always
functional.​ ​Eggs​ ​are​ ​enclosed​ ​in​ ​a​ ​proteinaceous​ ​ootheca​ ​which​ ​may​ ​be​ ​gestated
internally,​ ​brooded​ ​externally,​ ​or​ ​deposited​ ​without​ ​further​ ​maternal​ ​attention.​ ​ ​ ​Most
cockroaches​ ​are​ ​tropical​ ​and​ ​several​ ​species​ ​are​ ​pests​ ​in​ ​or​ ​near​ ​buildings​ ​especially​ ​in
warm​ ​climates.​ ​ ​ ​It​ ​is​ ​often​ ​found​ ​in​ ​homes​ ​and​ ​in​ ​my​ ​experience​ ​in​ ​the​ ​southeastern
US,​ ​it​ ​is​ ​common​ ​in​ ​zoology​ ​laboratories,​ ​especially​ ​in​ ​those​ ​in​ ​older​ ​buildings.

Biology
Periplaneta​ ​americana
​ ​ ​ ​Periplaneta​ ​americana​,​ ​the​ ​American​ ​cockroach,​ ​the​ ​largest​ ​widespread​ ​North
American​ ​cockroach,​ ​reaches​ ​44​ ​mm​ ​in​ ​length.​ ​It​ ​is​ ​dark​ ​reddish-brown​ ​with​ ​a​ ​pale

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peripheral​ ​band​ ​around​ ​the​ ​pronotum.​ ​ ​ ​Adults​ ​have​ ​large​ ​wings​ ​and​ ​can​ ​fly,​ ​although
they​ ​do​ ​not​ ​often​ ​do​ ​so.​ ​Nymphs,​ ​of​ ​course,​ ​are​ ​wingless.​ ​It​ ​is​ ​easily,​ ​albeit
unintentionally​ ​and​ ​regretfully,​ ​spread​ ​by​ ​human​ ​commerce​ ​and​ ​is​ ​worldwide​ ​in​ ​warm
climates​ ​and,​ ​in​ ​heated​ ​buildings,​ ​cosmopolitan.​ ​Periplaneta​ ​is​ ​nocturnal,​ ​negatively
phototactic,​ ​and​ ​prefers​ ​dark​ ​warm,​ ​moist​ ​habitats.​ ​It​ ​is​ ​acutely​ ​sensitive​ ​to​ ​vibrations
and​ ​is​ ​one​ ​of​ ​the​ ​world’s​ ​fastest​ ​running​ ​insects,​ ​as​ ​you​ ​know​ ​if​ ​you​ ​have​ ​tried​ ​to​ ​stomp
one​ ​in​ ​the​ ​kitchen​ ​late​ ​at​ ​night.​ ​Periplaneta​ ​americana​ ​is​ ​one​ ​of​ ​several​ ​cockroaches
found​ ​near​ ​(peridomestic)​ ​or​ ​in​ ​(domiciliary)​ ​human​ ​habitations.​ ​Such​ ​insects​ ​are
referred​ ​to​ ​as​ ​synanthropic​ ​(=​ ​with​ ​man).​ ​ ​ ​Several​ ​of​ ​the​ ​most​ ​common​ ​North​ ​American
synanthropic​ ​species​ ​are​ ​discussed​ ​below.​ ​Any​ ​of​ ​these​ ​species​ ​could​ ​be​ ​used​ ​for​ ​this
exercise​ ​but​ ​other​ ​species​ ​of​ ​Periplaneta​ ​differ​ ​the​ ​least​ ​from​ ​this​ ​account​ ​and​ ​for​ ​our
purposes​ ​are​ ​interchangeable​ ​with​ ​P.​ ​americana​.
​ ​The​ ​life​ ​cycle​ ​consists​ ​of​ ​egg,​ ​numerous​ ​nymphal​ ​instars​ ​(6-14),​ ​and​ ​the​ ​adult,
or​ ​imago.​ ​Blattarians​ ​are​ ​paurometabolous​ ​with​ ​nymphs​ ​closely​ ​resembling​ ​small​ ​adults
except​ ​for​ ​the​ ​absence​ ​of​ ​wings.​ ​ ​ ​Paurometabolous​ ​insects​ ​have​ ​no​ ​pupa​ ​or
metamorphosis.​ ​The​ ​eggs​ ​are​ ​laid​ ​in​ ​a​ ​tough,​ ​sclerotized​ ​protein​ ​case,​ ​the​ ​ootheca,
normally​ ​containing​ ​16​ ​eggs.​ ​Females​ ​average​ ​one​ ​ootheca​ ​per​ ​month​ ​for​ ​most​ ​of​ ​the
year​ ​but​ ​may​ ​exceed​ ​that​ ​average​ ​during​ ​periods​ ​of​ ​peak​ ​reproductive​ ​activity.​ ​Nymphs
lack​ ​functional​ ​wings,​ ​of​ ​course,​ ​but​ ​wingpads​ ​appear​ ​in​ ​the​ ​4​ th​ ​ ​ ​instar​ ​and​ ​gradually
increase​ ​in​ ​size​ ​with​ ​each​ ​molt.​ ​ ​ ​The​ ​life​ ​span​ ​is​ ​2-3​ ​years​ ​with​ ​only​ ​the​ ​last​ ​year​ ​being
spent​ ​as​ ​an​ ​adult.​ ​Adults​ ​live​ ​for​ ​about​ ​a​ ​year.
​ ​This​ ​omnivorous,​ ​opportunistic,​ ​scavenging​ ​species​ ​feeds​ ​on​ ​almost​ ​anything
organic,​ ​plant​ ​or​ ​animal,​ ​but​ ​prefer​ ​starches​ ​and​ ​sugars.​ ​Adults​ ​and​ ​nymphs​ ​have
similar​ ​feeding​ ​niches.
​ ​Cockroach​ ​population​ ​density​ ​is​ ​controlled​ ​naturally​ ​by​ ​several​ ​species​ ​of
parasitoid​ ​wasps​ ​including​ ​Evania​ ​and​ ​Aprostocetus​.​ ​ ​ ​Both​ ​are​ ​egg​ ​parasitoids​ ​in​ ​which
the​ ​female​ ​wasp​ ​oviposits​ ​in​ ​the​ ​roach​ ​ootheca​ ​where​ ​the​ ​wasp​ ​larvae​ ​hatch​ ​and​ ​feed
on​ ​roach​ ​eggs.​ ​Evania​ ​is​ ​a​ ​distinctive,​ ​small,​ ​black​ ​wasp​ ​with​ ​a​ ​laterally​ ​flattened
triangular​ ​abdomen​ ​that​ ​waves​ ​flag-like​ ​(hence​ ​the​ ​common​ ​name​ ​“ensign"​ ​wasp)​ ​as
the​ ​adult​ ​walks.​ ​ ​ ​Adult​ ​Evania​ ​are​ ​about​ ​1.5​ ​cm​ ​in​ ​length.​ ​ ​ ​The​ ​female,​ ​with​ ​great
difficulty,​ ​inserts​ ​one​ ​egg​ ​in​ ​each​ ​ootheca.​ ​ ​ ​The​ ​larva​ ​hatches​ ​and​ ​consumes​ ​all​ ​16
roach​ ​eggs​ ​before​ ​achieving​ ​a​ ​length​ ​of​ ​8​ ​mm,​ ​pupating,​ ​and​ ​emerging​ ​from​ ​the​ ​now
empty​ ​ootheca.​ ​Adult​ ​Aprostocetus​ ​are​ ​tiny,​ ​about​ ​2​ ​mm​ ​long.​ ​Both​ ​species​ ​are​ ​stingless
and​ ​no​ ​threat​ ​to​ ​humans.​ ​Small​ ​wasps,​ ​especially​ ​those​ ​waving​ ​a​ ​flag,​ ​seen​ ​in
roach-infested​ ​areas​ ​should​ ​be​ ​protected​ ​and​ ​encouraged,​ ​not​ ​killed.​ ​Attempts​ ​to
control​ ​roach​ ​populations​ ​with​ ​pesticide​ ​sprays​ ​can​ ​be​ ​counterproductive​ ​because​ ​they
indiscriminately​ ​kill​ ​parasitoids​ ​in​ ​addition​ ​to​ ​the​ ​roaches.​ ​ ​ ​Roach​ ​pellets​ ​(“roach
hotels”),​ ​which​ ​are​ ​selective,​ ​are​ ​more​ ​effective.
​ ​Cockroaches​ ​are​ ​not​ ​biological​ ​vectors​ ​for​ ​human​ ​disease​ ​although​ ​they​ ​can
serve​ ​as​ ​mechanical​ ​vectors​ ​simply​ ​by​ ​harboring​ ​infectious​ ​organisms​ ​such​ ​as​ ​Ascaris
eggs,​ ​bacteria,​ ​or​ ​protozoan​ ​cysts​ ​on​ ​their​ ​body​ ​surfaces.​ ​The​ ​American​ ​cockroach​ ​is
the​ ​host​ ​for​ ​the​ ​cystacanth​ ​stage​ ​of​ ​the​ ​rat​ ​intestinal​ ​acanthocephalan,​ ​Moniliformis
moniliformis​.
The​ ​Major​ ​Synanthropic​ ​Cockroaches

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 ​ ​Seven​ ​species​ ​of​ ​cockroaches​ ​enjoy​ ​close​ ​synanthropic​ ​associations​ ​with
humans​ ​and​ ​have​ ​been​ ​spread​ ​throughout​ ​the​ ​world​ ​by​ ​human​ ​migrations​ ​and
commerce​ ​and​ ​are​ ​now​ ​serious​ ​pests.​ ​ ​ ​All​ ​these​ ​species​ ​are​ ​thought​ ​to​ ​have​ ​originated
in​ ​Africa.​ ​Many​ ​other​ ​species​ ​are​ ​loosely​ ​associated​ ​with​ ​humans​ ​and​ ​their​ ​activities.
​ ​The​ ​American​ ​cockroach,​ ​Periplaneta​ ​americana​ ​,​ ​despite​ ​its​ ​specific​ ​and
common​ ​names,​ ​is​ ​an​ ​exotic​ ​that​ ​was​ ​introduced​ ​to​ ​North​ ​America​ ​from​ ​Africa​ ​in​ ​the
early​ ​17​ th​ ​ ​ ​century.​ ​ ​ ​It​ ​is​ ​the​ ​largest​ ​of​ ​the​ ​domiciliary​ ​species,​ ​reaching​ ​44​ ​mm​ ​in
length.​ ​ ​ ​Both​ ​sexes​ ​have​ ​fully​ ​functional​ ​wings​ ​but​ ​rarely​ ​fly.​ ​ ​ ​The​ ​pronotum​ ​is
bordered​ ​by​ ​a​ ​pale​ ​yellowish​ ​margin.​ ​ ​ ​It​ ​is​ ​easily​ ​confused​ ​with​ ​P.​ ​bunnea​.​ ​The​ ​last
article​ ​of​ ​the​ ​cercus​ ​of​ ​P.​ ​americana​ ​is​ ​over​ ​twice​ ​as​ ​long​ ​as​ ​wide​ ​(Fig​ ​16)​ ​whereas​ ​that
of​ ​P.​ ​brunnea​ ​is​ ​less​ ​than​ ​twice​ ​as​ ​long​ ​as​ ​wide​ ​(Fig​ ​14).​ ​Periplaneta​ ​americana​ ​is​ ​the
second​ ​most​ ​abundant​ ​cockroach​ ​pest​ ​in​ ​the​ ​United​ ​States.
​ ​Periplaneta​ ​fuliginosa​ ​(smoky​ ​brown​ ​cockroach)​ ​is​ ​an​ ​introduced​ ​peridomestic
species.​ ​ ​ ​It​ ​is​ ​common​ ​in​ ​the​ ​southern​ ​United​ ​States​ ​mostly​ ​found​ ​out​ ​of​ ​doors​ ​or​ ​in
crawl​ ​spaces​ ​or​ ​unheated​ ​buildings​ ​such​ ​as​ ​outhouses​ ​or​ ​garages.​ ​ ​ ​It​ ​also​ ​occurs​ ​in
buildings​ ​in​ ​the​ ​south​ ​and​ ​can​ ​be​ ​found​ ​further​ ​north​ ​in​ ​heated​ ​buildings.​ ​In​ ​some
localities​ ​it​ ​may​ ​account​ ​for​ ​almost​ ​80%​ ​of​ ​cockroaches​ ​captured​ ​near​ ​homes.​ ​The​ ​color
is​ ​uniformly​ ​dark​ ​brownish​ ​black​ ​and​ ​it​ ​lacks​ ​the​ ​pale​ ​pronotal​ ​margin​ ​of​ ​other
synanthropic​ ​Periplaneta​.​ ​ ​ ​Nor​ ​does​ ​it​ ​have​ ​pale​ ​areas​ ​on​ ​the​ ​forewings​ ​that​ ​distinguish
P.​ ​australasiae​.​ ​It​ ​reaches​ ​35​ ​mm​ ​in​ ​length,​ ​not​ ​quite​ ​as​ ​large​ ​as​ ​P.​ ​americana
​ ​Periplaneta​ ​brunnea​ ​(brown​ ​cockroach)​ ​is​ ​an​ ​African​ ​native​ ​spread​ ​by​ ​human
travel​ ​and​ ​commerce.​ ​ ​ ​It​ ​is​ ​common​ ​in​ ​the​ ​southeastern​ ​United​ ​States,​ ​sometimes
moreso​ ​than​ ​P.​ ​americana,​ ​but​ ​it​ ​is​ ​usually​ ​found​ ​outside.​ ​This​ ​species​ ​has​ ​a​ ​pale​ ​border
around​ ​the​ ​pronotum​ ​as​ ​do​ ​most​ ​Periplaneta​ ​but​ ​it​ ​lacks​ ​the​ ​pale​ ​areas​ ​on​ ​the​ ​forewings
characteristic​ ​of​ ​P.​ ​australasiae​.​ ​It​ ​reaches​ ​37​ ​mm​ ​in​ ​length.​ ​Both​ ​sexes​ ​are​ ​winged​ ​and
can​ ​fly.
​ ​Periplaneta​ ​australasiae​ ​(Australian​ ​cockroach)​ ​is​ ​almost​ ​as​ ​serious​ ​a​ ​domestic
pest​ ​as​ ​is​ ​P.​ ​americana​.​ ​It​ ​probably​ ​arose​ ​in​ ​Africa​ ​and​ ​spread​ ​throughout​ ​the​ ​tropics​ ​and
subtropics​ ​and​ ​is​ ​now​ ​circumtropical.​ ​ ​ ​It​ ​is​ ​the​ ​most​ ​common​ ​domiciliary​ ​cockroach​ ​in
Florida​ ​and​ ​although​ ​it​ ​occurs​ ​farther​ ​north​ ​it​ ​is​ ​intolerant​ ​of​ ​the​ ​cooler​ ​temperatures
tolerated​ ​by​ ​P.​ ​Americana​.​ ​It​ ​is​ ​similar​ ​to​ ​P.​ ​americana​ ​but​ ​slightly​ ​smaller,​ ​reaching​ ​35
mm​ ​in​ ​length.​ ​ ​ ​It​ ​is​ ​reddish​ ​brown,​ ​like​ ​P.​ ​Americana​.​ ​ ​ ​Unlike​ ​our​ ​other​ ​Periplaneta
species,​ ​its​ ​forewings​ ​have​ ​a​ ​pale​ ​yellowish​ ​band​ ​on​ ​the​ ​proximal​ ​anterior​ ​edge.​ ​Both
sexes​ ​have​ ​wings​ ​and​ ​fly.​ ​The​ ​specific​ ​epithet​ ​“​australasiae​”​ ​means​ ​“south​ ​Asia”,​ ​not
Australia.
​ ​Blatta​ ​orientalis​ ​(oriental​ ​cockroach,​ ​black​ ​beetle,​ ​water​ ​bug)​ ​probably​ ​originated
in​ ​North​ ​Africa​ ​and​ ​has​ ​become​ ​a​ ​common​ ​pest​ ​in​ ​the​ ​southern,​ ​midwestern,​ ​and
northwestern​ ​United​ ​States​ ​but​ ​is​ ​known​ ​as​ ​far​ ​north​ ​as​ ​southern​ ​Canada,​ ​the
Netherlands,​ ​and​ ​Britain.​ ​ ​ ​It​ ​prefers​ ​cool,​ ​moist​ ​habitats.​ ​ ​ ​Males​ ​are​ ​winged,​ ​females
have​ ​tiny​ ​vestigial​ ​wings,​ ​and​ ​neither​ ​is​ ​capable​ ​of​ ​flight.​ ​Individuals​ ​reach​ ​27​ ​mm​ ​in
length.​ ​ ​ ​Its​ ​color​ ​is​ ​dark,​ ​almost​ ​black,​ ​and​ ​it​ ​prefers​ ​moist​ ​habitats.​ ​Blatta​ ​lacks​ ​an
arolium​ ​on​ ​its​ ​tarsi​ ​and​ ​cannot​ ​climb​ ​smooth​ ​surfaces.​ ​ ​ ​The​ ​common​ ​name​ ​“water​ ​bug”
derives​ ​from​ ​its​ ​preference​ ​for​ ​wet​ ​areas.
​ ​The​ ​German​ ​cockroach,​ ​Blatella​ ​germanica​ ​,​ ​originated​ ​in​ ​Africa​ ​and​ ​traveled​ ​first
to​ ​Europe​ ​and​ ​from​ ​there​ ​to​ ​North​ ​America.​ ​It​ ​is​ ​the​ ​most​ ​abundant​ ​domiciliary

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cockroach​ ​in​ ​the​ ​United​ ​States​ ​and​ ​is​ ​an​ ​important​ ​pest.​ ​ ​ ​At​ ​15​ ​mm,​ ​it​ ​is​ ​much​ ​smaller
than​ ​the​ ​Periplaneta​ ​species.​ ​ ​ ​It​ ​is​ ​pale​ ​brown​ ​with​ ​two​ ​dark​ ​longitudinal​ ​bands
bordering​ ​the​ ​pronotum.
​ ​Supella​ ​longipalpa​ ​(brown​ ​banded​ ​cockroach)​ ​is​ ​assumed​ ​to​ ​have​ ​originated​ ​in
Africa.​ ​ ​ ​It​ ​is​ ​always​ ​found​ ​in​ ​buildings​ ​(except​ ​in​ ​Africa).​ ​ ​ ​It​ ​was​ ​originally​ ​introduced
into​ ​the​ ​Florida​ ​and​ ​spread​ ​north​ ​hitchhiking​ ​on​ ​fruit​ ​shipments​ ​and​ ​in​ ​the​ ​luggage​ ​and
automobiles​ ​of​ ​tourists​ ​returning​ ​north​ ​after​ ​a​ ​vacation.​ ​ ​ ​It​ ​is​ ​now​ ​found​ ​throughout​ ​the
contiguous​ ​United​ ​States.​ ​This​ ​is​ ​a​ ​small​ ​roach,​ ​reaching​ ​only​ ​14​ ​mm,​ ​and​ ​similar​ ​in
size​ ​and​ ​general​ ​appearance​ ​to​ ​Blatella​ ​germanica​.​ ​The​ ​forewings​ ​cover​ ​the​ ​abdomen​ ​of
the​ ​male​ ​but​ ​those​ ​of​ ​females​ ​are​ ​shorter​ ​and​ ​the​ ​posterior​ ​abdomen​ ​is​ ​left​ ​uncovered.
Two​ ​pale​ ​bands​ ​extend​ ​transversely​ ​across​ ​the​ ​dark​ ​abdomen​ ​but​ ​these​ ​may​ ​be​ ​difficult
to​ ​see​ ​with​ ​the​ ​wings​ ​folded​ ​at​ ​rest.​ ​ ​ ​They​ ​are​ ​obvious​ ​in​ ​the​ ​wingless​ ​nymphs.​ ​Supella
lacks​ ​the​ ​longitudinal​ ​stripes​ ​on​ ​the​ ​pronotum​ ​characteristic​ ​of​ ​Blatella​.​ ​The​ ​lateral
margins​ ​of​ ​the​ ​pronotum​ ​of​ ​Supella​ ​are​ ​transparent.

Laboratory​ ​Specimens
​ ​Periplaneta​ ​is​ ​available​ ​at​ ​modest​ ​cost,​ ​alive​ ​or​ ​preserved,​ ​from​ ​biological​ ​supply
companies.​ ​ ​ ​It​ ​can​ ​also​ ​be​ ​captured​ ​locally​ ​in​ ​homes​ ​or​ ​university​ ​buildings.​ ​ ​ ​Look​ ​for​ ​it
in​ ​warm,​ ​moist,​ ​dark​ ​places​ ​such​ ​as​ ​basements,​ ​drawers,​ ​wood​ ​piles,​ ​sewers,​ ​sunken
water​ ​meter​ ​enclosures,​ ​and​ ​compost​ ​piles.​ ​ ​ ​In​ ​the​ ​home​ ​it​ ​can​ ​be​ ​found​ ​wherever
there​ ​is​ ​food,​ ​including​ ​the​ ​usual​ ​debris​ ​on​ ​the​ ​kitchen​ ​floor​ ​and​ ​under​ ​the​ ​refrigerator,
on​ ​unwashed​ ​dishes​ ​left​ ​overnight​ ​in​ ​the​ ​sink,​ ​and​ ​even​ ​in​ ​your​ ​library​ ​where​ ​it​ ​eats​ ​the
bindings​ ​of​ ​your​ ​favorite​ ​books.​ ​Female​ ​Periplaneta​ ​are​ ​distinguished​ ​from​ ​other​ ​genera
by​ ​having​ ​a​ ​divided​ ​7​ th​​ ​ ​sternite.
​ ​Specimens​ ​may​ ​be​ ​dissected​ ​alive​ ​and​ ​anesthetized,​ ​recently​ ​sacrificed​ ​and
unpreserved,​ ​or​ ​preserved.​ ​ ​ ​Recently​ ​sacrificed​ ​is​ ​preferable​ ​unless​ ​you​ ​anticipate
making​ ​physiological​ ​observations,​ ​in​ ​which​ ​case​ ​the​ ​specimens​ ​should​ ​be​ ​alive​ ​and
anesthetized.​ ​Living​ ​specimens​ ​may​ ​be​ ​sacrificed​ ​in​ ​a​ ​covered​ ​dish​ ​with​ ​a​ ​cotton​ ​ball
dampened​ ​with​ ​ethyl​ ​acetate.​ ​ ​ ​Living​ ​specimens​ ​should​ ​be​ ​anesthetized​ ​using
chloroform,​ ​ether,​ ​or​ ​carbon​ ​dioxide.​ ​ ​ ​Sacrificed​ ​specimens​ ​should​ ​be​ ​immersed​ ​in
water​ ​or​ ​80%​ ​ethanol​ ​(or​ ​40%​ ​isopropanol)​ ​in​ ​a​ ​small​ ​dissecting​ ​pan.
​ ​If​ ​both​ ​external​ ​and​ ​internal​ ​anatomy​ ​are​ ​to​ ​be​ ​studied​ ​it​ ​is​ ​advantageous​ ​to
have​ ​separate​ ​specimens​ ​for​ ​these​ ​two​ ​undertakings.​ ​ ​ ​The​ ​study​ ​of​ ​external​ ​anatomy​ ​is
likely​ ​to​ ​damage​ ​the​ ​specimen​ ​to​ ​such​ ​an​ ​extent​ ​that​ ​it​ ​will​ ​not​ ​be​ ​useful​ ​for​ ​studying
internal​ ​anatomy.​ ​ ​ ​Students​ ​should​ ​have​ ​access​ ​to​ ​both​ ​sexes,​ ​either​ ​their​ ​own
specimens​ ​or​ ​by​ ​sharing​ ​with​ ​a​ ​lab​ ​partner.

Anatomy
External​ ​Anatomy
​ ​Species​ ​of​ ​Periplaneta​ ​are​ ​elongate,​ ​oval​ ​in​ ​outline​ ​and​ ​strongly​ ​dorso-ventrally
depressed,​ ​or​ ​flattened.​ ​The​ ​body​ ​is​ ​divided​ ​into​ ​the​ ​three​ ​tagmata​ ​characteristic​ ​of
insects,​ ​i.e.​ ​head,​ ​thorax,​ ​and​ ​abdomen​ ​(Fig​ ​21-1C).​ ​ ​ ​The​ ​head​ ​is​ ​inconspicuous​ ​in
dorsal​ ​view​ ​but​ ​the​ ​pronotum​ ​of​ ​the​ ​thorax​ ​is​ ​very​ ​large.​ ​ ​ ​The​ ​remainder​ ​of​ ​the​ ​thorax
and​ ​abdomen​ ​are​ ​hidden​ ​by​ ​the​ ​two​ ​pairs​ ​of​ ​wings.​ ​ ​ ​Six​ ​pairs​ ​of​ ​similar,​ ​strong,​ ​spiny

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legs​ ​are​ ​present.
​ ​Study​ ​a​ ​recently​ ​sacrificed​ ​or​ ​preserved​ ​cockroach​ ​with​ ​the​ ​dissecting
microscope.​ ​ ​ ​The​ ​specimen​ ​may​ ​be​ ​dry​ ​or​ ​submersed​ ​in​ ​liquid.
Figure​ ​1.​ ​Dorsal​ ​view​ ​of​ ​a​ ​female​ ​smoky​ ​brown​ ​cockroach,​ ​Periplaneta​ ​fuliginosa​,
from​ ​Greenwood,​ ​South​ ​Carolina.​ ​Blatt37L.gif

Head
Little​ ​of​ ​the​ ​head​ ​can​ ​be​ ​seen​ ​in​ ​dorsal​ ​view​ ​(Fig​ ​1).​ ​Position​ ​the​ ​specimen​ ​with​ ​the
ventral​ ​surface​ ​up​ ​and​ ​examine​ ​the​ ​head​ ​with​ ​the​ ​dissecting​ ​microscope.​ ​ ​ ​The​ ​head​​ ​is
flattened​ ​dorsoventrally​ ​and​ ​is​ ​more​ ​or​ ​less​ ​pear-shaped​ ​in​ ​outline​ ​with​ ​the​ ​mouthparts
extending​ ​posteriorly​ ​from​ ​the​ ​narrow​ ​end​ ​of​ ​the​ ​pear.​ ​The​ ​head​ ​is​ ​normally​ ​carried​ ​with
its​ ​flat​ ​front​ ​surface,​ ​the​ ​face,​ ​held​ ​horizontally​ ​facing​ ​the​ ​substratum​ ​(Fig​ ​2).​ ​ ​ ​In​ ​this
position​ ​the​ ​mouth​ ​opens​ ​posteriorly,​ ​the​ ​mandibles​ ​project​ ​posteriorly,​ ​and​ ​the​ ​palps
touch​ ​the​ ​coxae​ ​of​ ​the​ ​forelegs.​ ​ ​ ​When​ ​feeding,​ ​the​ ​flattened​ ​face​​ ​is​ ​held​ ​vertically​ ​with
the​ ​mouthparts​ ​located​ ​ventrally,​ ​toward​ ​the​ ​substratum.​ ​ ​ ​The​ ​flattened​ ​face​ ​is​ ​the
anterior​ ​aspect​ ​of​ ​the​ ​head​ ​even​ ​though​ ​it​ ​is​ ​oriented​ ​ventrally.​ ​ ​ ​Similarly,​ ​the​ ​thin​ ​edge
of​ ​the​ ​broad​ ​end​ ​of​ ​the​ ​pear​ ​is​ ​dorsal,​ ​even​ ​though​ ​it​ ​is​ ​normally​ ​held​ ​in​ ​an​ ​anterior
position.​ ​ ​ ​When​ ​in​ ​the​ ​feeding​ ​position​ ​the​ ​face​ ​is​ ​held​ ​anteriorly​ ​and​ ​the​ ​thin​ ​edge​ ​is
dorsal.
Cockroaches​ ​have​ ​typical​ ​unspecialized​ ​mouthparts​ ​and​ ​provide​ ​a​ ​good​ ​example​ ​of​ ​the
primitive​ ​insect​ ​mouthparts​ ​adapted​ ​for​ ​biting​ ​and​ ​chewing​ ​(Fig​ ​21-1A,B).​ ​ ​ ​The​ ​head

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can​ ​be​ ​retracted​ ​beneath​ ​the​ ​prothorax.

Figure​ ​2.​ ​Anterior​ ​view​ ​of​ ​the​ ​head​ ​of​ ​Periplaneta​ ​amricana

Head​ ​Capsule
​ ​The​ ​head​ ​consists​ ​of​ ​a​ ​complex​ ​of​ ​fused​ ​sclerites,​ ​which​ ​constitute​ ​the
epicranium,​ ​or​ ​head​ ​capsule,​​ ​associated​ ​with​ ​unfused​ ​sclerites​ ​and​ ​appendages.​ ​This
hard​ ​sclerotized​ ​capsule​ ​encloses​ ​the​ ​mouthpart​ ​muscles,​ ​brain,​ ​and​ ​anteriormost
region​ ​of​ ​the​ ​gut​ ​and​ ​provides​ ​sites​ ​of​ ​attachment​ ​for​ ​the​ ​mouthparts​ ​and​ ​antennae.
The​ ​head​ ​capsule​ ​is​ ​divided​ ​into​ ​several​ ​regions.​ ​ ​ ​Locate​ ​the​ ​large​ ​compound
eyes​​ ​on​ ​the​ ​dorsal​ ​edge​ ​of​ ​the​ ​head​ ​(Fig​ ​1,​ ​2).​ ​ ​ ​The​ ​antennae​​ ​arise​ ​on​ ​the​ ​face
immediately​ ​ventral​ ​to​ ​the​ ​eyes.​ ​ ​ ​These​ ​are​ ​useful​ ​as​ ​convenient,​ ​easily​ ​recognized
landmarks.
​ ​The​ ​epicranial​ ​suture​​ ​marks​ ​the​ ​junction​ ​of​ ​three​ ​fused​ ​sclerites​ ​that​ ​form​ ​the
face.​ ​In​ ​Periplaneta​ ​this​ ​suture​ ​is​ ​usually​ ​visible​ ​as​ ​a​ ​faint,​ ​yellow,​ ​Y-shaped​ ​line​ ​on​ ​the
face​ ​(Fig​ ​2)​ ​but​ ​sometimes​ ​it​ ​cannot​ ​be​ ​found.​ ​The​ ​two​ ​arms​ ​of​ ​the​ ​"Y"​ ​begin​ ​at​ ​the
antennal​ ​sockets​ ​and​ ​converge​ ​between​ ​the​ ​two​ ​eyes.​ ​ ​ ​From​ ​the​ ​convergence​ ​the
single​ ​stem​ ​of​ ​the​ ​"Y"​ ​extends​ ​dorsally​ ​between​ ​the​ ​eyes​ ​over​ ​the​ ​top​ ​of​ ​the​ ​head.​ ​ ​ ​It
divides​ ​the​ ​top​ ​of​ ​the​ ​head​ ​into​ ​a​ ​pair​ ​of​ ​sclerites,​ ​the​ ​epicranial​ ​plates​​ ​(Fig​ ​2).​ ​The​ ​part
of​ ​the​ ​capsule​ ​between​ ​the​ ​two​ ​arms​ ​of​ ​the​ ​“Y”​ ​is​ ​the​ ​frons​,​ ​or​ ​front,​ ​of​ ​the​ ​capsule​ ​(Fig
2).​ ​The​ ​frons​ ​is​ ​an​ ​unpaired​ ​sclerite.
Two​ ​white​ ​circular​ ​areas​ ​between​ ​the​ ​antennae​ ​and​ ​compound​ ​eyes,​ ​at​ ​the​ ​ends
of​ ​the​ ​arms​ ​of​ ​the​ ​epicranial​ ​suture,​ ​are​ ​the​ ​fenestrae​​ ​and​ ​are​ ​the​ ​vestiges​ ​of​ ​the​ ​ocelli
of​ ​other​ ​insects​ ​(Fig​ ​21-1A).​ ​ ​ ​Two​ ​additional​ ​pale​ ​circles​ ​on​ ​the​ ​frons,​ ​ventral​ ​and​ ​a​ ​little
medial​ ​to​ ​the​ ​fenestrae,​ ​mark​ ​the​ ​sites​ ​of​ ​muscle​ ​insertions​ ​inside​ ​the​ ​capsule.

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 The​ ​vertex​,​ ​which​ ​is​ ​a​ ​region,​ ​not​ ​a​ ​sclerite,​ ​occupies​ ​the​ ​dorsum​ ​of​ ​the​ ​capsule
between​ ​the​ ​two​ ​eyes.​ ​It​ ​is​ ​divided​ ​in​ ​two​ ​by​ ​the​ ​stem​ ​of​ ​the​ ​epicranial​ ​suture​ ​(Fig​ ​2)
and​ ​includes​ ​the​ ​two​ ​epicranial​ ​plates.​ ​ ​ ​In​ ​some​ ​insects​ ​the​ ​posteriormost​ ​region​ ​of​ ​the
vertex​ ​may​ ​be​ ​a​ ​distinct​ ​sclerite​ ​referred​ ​to​ ​as​ ​the​ ​occiput.
​ ​The​ ​clypeus​​ ​(KLIP​ ​ee​ ​us)​ ​is​ ​an​ ​unpaired​ ​sclerite​ ​on​ ​the​ ​face​ ​(Fig​ ​2,​ ​21-1A).​ ​ ​ ​It
lies​ ​ventral​ ​to​ ​the​ ​frons​ ​and​ ​the​ ​two​ ​are​ ​joined​ ​along​ ​a​ ​transverse,​ ​indistinguishable
suture.​ ​ ​ ​The​ ​clypeus​ ​is​ ​not​ ​movable.
​ ​A​ ​third​ ​unpaired​ ​sclerite,​ ​the​ ​labrum​,​ ​is​ ​attached​ ​along​ ​a​ ​transverse,​ ​movable
articulation​ ​with​ ​the​ ​ventral​ ​edge​ ​of​ ​the​ ​clypeus​ ​(Fig​ ​2).​ ​ ​ ​The​ ​labrum​ ​is​ ​equipped​ ​with
muscles​ ​and​ ​is​ ​movable.​ ​ ​ ​It​ ​forms​ ​the​ ​anterior​ ​wall​ ​of​ ​the​ ​preoral​ ​cavity​ ​into​ ​which​ ​the
mouth​ ​opens.​ ​ ​ ​The​ ​labrum​ ​is​ ​usually​ ​considered​ ​to​ ​be​ ​derived​ ​from​ ​a​ ​sclerite​ ​of​ ​one​ ​of
the​ ​head​ ​segments​ ​but​ ​some​ ​entomologists​ ​believe​ ​it​ ​to​ ​be​ ​homologous​ ​to​ ​a​ ​pair​ ​of
fused​ ​appendages,​ ​similar​ ​to​ ​the​ ​labium​ ​in​ ​this​ ​regard.
​ ​The​ ​sides,​ ​or​ ​cheeks,​ ​of​ ​the​ ​epicranium​ ​are​ ​formed​ ​by​ ​the​ ​paired​ ​genae​.​ ​In​ ​the
cockroach​ ​these​ ​form​ ​the​ ​thin​ ​lateral​ ​edges​ ​of​ ​the​ ​head​ ​ventral​ ​and​ ​posterior​ ​to​ ​the
eyes.​ ​ ​ ​The​ ​genae​ ​are​ ​formed​ ​of​ ​several​ ​indistinguishably​ ​fused​ ​sclerites.
​ ​Posteriorly​ ​the​ ​head​ ​capsule​ ​is​ ​penetrated​ ​by​ ​a​ ​large​ ​circular​ ​opening,​ ​the
foramen​ ​magnum​​ ​through​ ​which​ ​pass​ ​the​ ​gut,​ ​salivary​ ​ducts,​ ​and​ ​nerve​ ​cord.​ ​The
foramen​ ​is​ ​bordered​ ​by​ ​the​ ​vertex​ ​dorsally​ ​and​ ​the​ ​genae​ ​laterally.​ ​ ​ ​The​ ​soft,​ ​cervix​,​ ​or
neck,​ ​extends​ ​posteriorly​ ​from​ ​the​ ​foramen.​ ​ ​ ​Because​ ​of​ ​the​ ​attachment​ ​of​ ​soft​ ​tissue​ ​of
the​ ​cervix​ ​to​ ​the​ ​head​ ​capsule,​ ​the​ ​foramen​ ​is​ ​not​ ​visible​ ​externally.​ ​ ​ ​Ventrally​ ​slender,
transverse​ ​cervical​ ​sclerites​​ ​are​ ​embedded​ ​in​ ​the​ ​otherwise​ ​soft​ ​integument​ ​of​ ​the
cervix.
Antenna
​ ​The​ ​anteriormost​ ​head​ ​appendages​ ​are​ ​the​ ​two​ ​antennae.​ ​These​ ​arise​ ​from​ ​a
socket​​ ​ventral​ ​to​ ​the​ ​compound​ ​eyes.​ ​ ​ ​ ​The​ ​first​ ​article​ ​of​ ​the​ ​antenna​ ​is​ ​the​ ​scape​​ ​and
the​ ​second​ ​is​ ​the​ ​pedicel​​ ​(Fig​ ​2).​ ​ ​ ​From​ ​the​ ​pedicel​ ​arises​ ​a​ ​long,​ ​whiplike,​ ​sensory
flagellum​​ ​of​ ​75-90​ ​articles.​ ​ ​ ​The​ ​antennae​ ​are​ ​equipped​ ​with​ ​mechanoreceptors​ ​but
apparently​ ​not​ ​with​ ​chemoreceptors.

Preoral​ ​Cavity
Ventrally​ ​the​ ​mouthparts​ ​surround​ ​and​ ​enclose​ ​a​ ​chamber,​ ​the​ ​preoral​ ​cavity​,
from​ ​which​ ​the​ ​mouth​ ​opens​ ​(Fig​ ​21-7).​ ​ ​ ​As​ ​its​ ​name​ ​suggests,​ ​the​ ​preoral​ ​cavity
precedes​ ​the​ ​mouth​ ​and​ ​is​ ​not​ ​considered​ ​to​ ​be​ ​part​ ​of​ ​the​ ​gut,​ ​although​ ​both
mechanical​ ​and​ ​chemical​ ​digestion​ ​begin​ ​there.
With​ ​the​ ​specimen​ ​positioned​ ​with​ ​its​ ​ventral​ ​side​ ​up,​ ​grasp​ ​the​ ​labrum​ ​and
mandibles​ ​with​ ​a​ ​pair​ ​of​ ​fine​ ​forceps​ ​and​ ​pull​ ​the​ ​head​ ​into​ ​the​ ​feeding​ ​position​ ​(i.e.​ ​with
the​ ​anterior​ ​face​ ​anterior,​ ​and​ ​the​ ​mouthparts​ ​ventral).​ ​ ​ ​Lift​ ​the​ ​labrum​ ​and​ ​look
beneath​ ​it​ ​using​ ​a​ ​second​ ​pair​ ​of​ ​forceps​ ​or​ ​a​ ​fine​ ​needle​ ​to​ ​move​ ​structures​ ​aside​ ​as
needed.
The​ ​labrum,​ ​which​ ​you​ ​are​ ​currently​ ​holding​ ​aside,​ ​is​ ​the​ ​anterior​ ​wall​ ​of​ ​the
cavity.​ ​ ​ ​The​ ​heavy,​ ​toothed​ ​mandibles​ ​and​ ​softer​ ​maxillae​ ​are​ ​its​ ​side​ ​walls​ ​and​ ​the
labium​ ​is​ ​its​ ​posterior​ ​wall.​ ​ ​ ​A​ ​soft,​ ​tonguelike,​ ​mostly​ ​unsclerotized​ ​outgrowth​ ​of​ ​the
ventral​ ​body​ ​wall,​ ​the​ ​hypopharynx,​ ​extends​ ​into​ ​the​ ​preoral​ ​cavity​ ​immediately

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posterior​ ​to​ ​the​ ​mouth.​ ​ ​ ​Move​ ​the​ ​mandibles​ ​and​ ​maxillae​ ​aside​ ​and​ ​find​ ​the​ ​mouth
opening​ ​dorsally​ ​from​ ​the​ ​preoral​ ​cavity​ ​immediately​ ​anterior​ ​to​ ​the​ ​hypopharynx.

Mandible
The​ ​two​ ​mandibles​​ ​lie​ ​beside​ ​the​ ​mouth​ ​and​ ​form​ ​the​ ​lateral​ ​walls​ ​of​ ​the​ ​preoral
cavity​ ​(Fig​ ​2,​ ​21-1A,​ ​B).​ ​They​ ​are​ ​derived​ ​from​ ​a​ ​pair​ ​of​ ​segmental​ ​appendages.​ ​Each
mandible​ ​consists​ ​of​ ​a​ ​single​ ​article​ ​bearing​ ​a​ ​sclerotized,​ ​distal,​ ​cutting​ ​surface.​ ​The
cutting​ ​edge​ ​of​ ​the​ ​mandible​ ​consists​ ​of​ ​a​ ​lateral​ ​shearing​ ​incisor​​ ​with​ ​three​ ​or​ ​four
sharp​ ​denticles​​ ​and​ ​a​ ​more​ ​medial​ ​grinding​ ​molar​​ ​(Fig​ ​3,​ ​21-1A).
The​ ​mandible​ ​is​ ​largely​ ​hidden​ ​from​ ​view​ ​but​ ​can​ ​be​ ​glimpsed​ ​on​ ​the​ ​side​ ​of​ ​the
head​ ​ventral​ ​to​ ​the​ ​gena​ ​and​ ​lateral​ ​to​ ​the​ ​clypeus​ ​(Fig​ ​2,​ ​4).​ ​ ​ ​With​ ​forceps​ ​carefully​ ​lift
the​ ​labrum​ ​and​ ​peer​ ​beneath​ ​it​ ​to​ ​get​ ​a​ ​better​ ​view​ ​of​ ​the​ ​mandible.​ ​ ​ ​Its​ ​dark,​ ​heavily
sclerotized​ ​denticles​ ​are​ ​visible​ ​in​ ​this​ ​view.
The​ ​mandible​ ​articulates​ ​with​ ​the​ ​head​ ​capsule​ ​by​ ​two​ ​ball​ ​and​ ​socket-like
condyles,​ ​one​ ​anterior​ ​and​ ​one​ ​posterior​ ​(Fig​ ​3,​ ​4).​ ​ ​ ​The​ ​anterior​ ​condyle​​ ​can​ ​be​ ​seen
at​ ​the​ ​dorsolateral​ ​corner​ ​of​ ​the​ ​clypeus,​ ​on​ ​the​ ​face.​ ​ ​ ​The​ ​posterior​ ​condyle
articulates​ ​with​ ​the​ ​ventral​ ​corner​ ​of​ ​the​ ​gena​ ​and​ ​can​ ​be​ ​seen​ ​on​ ​the​ ​side​ ​of​ ​the​ ​head
capsule.​ ​ ​ ​Its​ ​plane​ ​of​ ​motion​ ​is​ ​transverse,​ ​perpendicular​ ​to​ ​the​ ​long​ ​axis​ ​of​ ​the​ ​body.
Movement​ ​in​ ​this​ ​plane​ ​brings​ ​the​ ​two​ ​mandibles​ ​together​ ​on​ ​the​ ​midline.​ ​In​ ​contrast,
vertebrate​ ​mandibles​ ​oppose​ ​each​ ​other​ ​by​ ​moving​ ​in​ ​the​ ​vertical​ ​plane.​ ​Observe​ ​the
mandible​ ​with​ ​the​ ​labrum​ ​moved​ ​aside​ ​and​ ​imagine​ ​the​ ​motion​ ​of​ ​the​ ​two​ ​mandibles
with​ ​respect​ ​to​ ​each​ ​other.​ ​Grasp​ ​the​ ​mandibles​ ​with​ ​forceps​ ​and​ ​move​ ​them​ ​in​ ​their
preferred​ ​plane​ ​of​ ​motion.​ ​If​ ​desired,​ ​one​ ​of​ ​the​ ​mandibles​ ​can​ ​be​ ​removed​ ​for​ ​closer
examination.
Figure​ ​3.​ ​The​ ​left​ ​mandible.​ ​A.​ ​Anterior​ ​view.​ ​B.​ ​Posterior​ ​view.​ ​Blatt39L.gif

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 Maxilla
Look​ ​at​ ​the​ ​head​ ​from​ ​the​ ​side​ ​and​ ​find​ ​the​ ​maxilla​​ ​(Fig​ ​2,​ ​21-1A,B).​ ​ ​ ​Like​ ​the
mandibles,​ ​the​ ​two​ ​maxillae​ ​are​ ​paired​ ​appendages​ ​lying​ ​beside​ ​the​ ​preoral​ ​cavity.
The​ ​maxilla​ ​can​ ​be​ ​recognized​ ​by​ ​its​ ​large​ ​5-articulate​ ​palp​ ​(the​ ​labium​ ​also​ ​has​ ​a​ ​palp
but​ ​it​ ​is​ ​much​ ​smaller​ ​and​ ​has​ ​three​ ​articles).​ ​ ​ ​Each​ ​maxilla​ ​consists​ ​of​ ​a​ ​proximal
cardo​ ​that​ ​articulate​ ​with​ ​the​ ​head​ ​capsule​ ​and​ ​a​ ​distal​ ​stipes​ ​from​ ​which​ ​arise​ ​several
processes.

Figure​ ​4.​ ​ ​ ​Oblique​ ​view​ ​of​ ​the​ ​head​ ​from​ ​the​ ​left​ ​showing​ ​the​ ​articulation​ ​of
the​ ​mandible​ ​with​ ​the​ ​head​ ​capsule​ ​of​ ​P.​ ​Americana.​ ​ ​ ​The​ ​labrum​ ​has​ ​been​ ​drawn
as​ ​if​ ​transparent​ ​to​ ​reveal​ ​the​ ​left​ ​mandible​ ​beneath​ ​it.​ ​ ​ ​The​ ​right​ ​mandible​ ​has
been​ ​omitted​ ​for​ ​clarity.

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 ​​
The​ ​cardo​​ ​in​ ​a​ ​short​ ​transverse​ ​partly​ ​sclerotized​ ​article​ ​extending​ ​laterally​ ​from
the​ ​head​ ​capsule​ ​(Fig​ ​5)​ ​ ​ ​It​ ​is​ ​inconspicuous​ ​because​ ​it​ ​is​ ​hidden​ ​by​ ​the​ ​larger​ ​stipes.
The​ ​stipes​​ ​is​ ​easily​ ​seen​ ​when​ ​looking​ ​at​ ​the​ ​head​ ​from​ ​the​ ​side.​ ​ ​ ​It​ ​is​ ​elongate​ ​and
articulates​ ​with​ ​the​ ​cardo​ ​proximally,​ ​beside​ ​the​ ​ventral​ ​edge​ ​of​ ​the​ ​eye.​ ​ ​ ​Distally​ ​three
processes​ ​arise​ ​from​ ​the​ ​stipes.
Most​ ​obvious​ ​of​ ​these​ ​processes​ ​is​ ​the​ ​maxillary​ ​palp​​ ​consisting​ ​of​ ​five​ ​articles.
The​ ​palp​ ​is​ ​sensory​ ​with​ ​chemo-​ ​and​ ​mechanorecepotors​ ​used​ ​to​ ​evaluate​ ​the
suitability​ ​of​ ​potential​ ​food.​ ​ ​ ​From​ ​the​ ​distal​ ​end​ ​of​ ​the​ ​stipes​ ​arise​ ​a​ ​lightly​ ​sclerotized,
lateral​ ​galea​​ ​and​ ​median,​ ​sclerotized,​ ​bladelike​ ​lacinia,​​ ​which​ ​bears​ ​denticles​​ ​distally.
The​ ​lacinia​ ​assists​ ​the​ ​mandible​ ​in​ ​cutting​ ​food​ ​into​ ​smaller​ ​particles.​ ​The​ ​lacinia​ ​is
completely​ ​hidden​ ​by​ ​the​ ​soft​ ​bulbous​ ​galea.​ ​The​ ​galea​ ​acts​ ​as​ ​a​ ​sheath​ ​to​ ​cover​ ​the
lacinia​ ​and​ ​because​ ​of​ ​it,​ ​the​ ​lacinia​ ​is​ ​not​ ​visible​ ​until​ ​the​ ​galea​ ​is​ ​moved​ ​aside.​ ​You
may​ ​remove​ ​one​ ​of​ ​the​ ​maxilla​ ​for​ ​closer​ ​study​ ​if​ ​desired.
Labium
​ ​The​ ​insect​ ​labium,​ ​also​ ​known​ ​as​ ​the​ ​second​ ​maxilla,​ ​is​ ​formed​ ​by​ ​the​ ​fusion​ ​of
the​ ​posteriormost​ ​pair​ ​of​ ​head​ ​appendages.​ ​The​ ​labium​ ​is​ ​the​ ​posterior​ ​wall​ ​of​ ​the
preoral​ ​cavity.​ ​It​ ​is​ ​best​ ​viewed​ ​by​ ​lifting​ ​the​ ​head​ ​to​ ​reveal​ ​its​ ​posterior​ ​surface.​ ​The
labium​​ ​consists​ ​of​ ​a​ ​large,​ ​platelike,​ ​proximal​ ​submentum​​ ​(Fig​ ​6,​ ​21-1A,B).
Articulated​ ​with​ ​it​ ​is​ ​the​ ​similarly​ ​platelike,​ ​but​ ​smaller,​ ​mentum​.​ ​ ​ ​Submentum​ ​and
mentum​ ​are​ ​the​ ​fused​ ​regions​ ​of​ ​the​ ​ancestral​ ​appendages.​ ​ ​ ​Arising​ ​from​ ​the​ ​distal

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edge​ ​of​ ​the​ ​mentum​ ​is​ ​a​ ​pair​ ​of​ ​prementa​,​ ​the​ ​homologs​ ​of​ ​the​ ​original​ ​two
appendages.​ ​ ​ ​Each​ ​prementum​ ​bears​ ​a​ ​distal​ ​median​ ​glossa​​ ​with​ ​a​ ​lateral​ ​paraglossa
beside​ ​it.​ ​ ​ ​The​ ​glossae​ ​and​ ​paraglossae​ ​are​ ​together​ ​known​ ​as​ ​the​ ​ligula​​ ​and​ ​function
in​ ​the​ ​manipulation​ ​of​ ​food.​ ​ ​ ​A​ ​3-articulate​ ​labial​ ​palp​​ ​arises​ ​laterally​ ​from​ ​the​ ​side​ ​of
each​ ​prementum.​ ​ ​ ​The​ ​palp​ ​is​ ​chemo-​ ​and​ ​mechanosensory.​ ​ ​ ​You​ ​may​ ​remove​ ​the
labium​ ​and​ ​examine​ ​it​ ​with​ ​higher​ ​magnification​ ​if​ ​you​ ​wish.
Figure​ ​5.​ ​Posterior​ ​view​ ​of​ ​the​ ​left​ ​maxilla​ ​of​ ​P.americana​ ​with​ ​the​ ​galea​ ​moved
slightly​ ​to​ ​reveal​ ​the​ ​lacinia.

Hypopharynx
​ ​The​ ​hypopharynx​,​ ​or​ ​tongue,​ ​is​ ​a​ ​long,​ ​process​ ​protruding​ ​from​ ​the​ ​ventral​ ​wall
of​ ​the​ ​head​ ​into​ ​the​ ​preoral​ ​cavity​ ​(Fig​ ​7,​ ​21-7).​ ​It​ ​is​ ​not​ ​a​ ​segmental​ ​appendage​ ​being
instead​ ​a​ ​fold​ ​of​ ​the​ ​body​ ​wall.​ ​It​ ​can​ ​be​ ​seen​ ​by​ ​looking​ ​into​ ​the​ ​preoral​ ​cavity,​ ​either
ventrally​ ​by​ ​spreading​ ​the​ ​mouthparts​ ​aside,​ ​or​ ​posteriorly​ ​by​ ​lifting​ ​(or​ ​removing)​ ​the
labium.​ ​ ​ ​The​ ​hypopharynx​ ​divides​ ​the​ ​preoral​ ​cavity​ ​into​ ​an​ ​anterior​ ​cibarium​ ​and​ ​a
posterior​ ​salivarium.
Thorax
​ ​The​ ​thorax​ ​is​ ​the​ ​tagma​ ​specialized​ ​for​ ​locomotion​ ​and​ ​as​ ​such​ ​bears​ ​three
pairs​ ​of​ ​legs​ ​and,​ ​in​ ​adults,​ ​two​ ​pairs​ ​of​ ​wings​ ​and​ ​houses​ ​the​ ​muscles​ ​to​ ​operate​ ​them

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(Fig​ ​16-2).​ ​It​ ​is​ ​the​ ​middle​ ​tagma​ ​of​ ​the​ ​body​ ​and​ ​consists​ ​of​ ​the​ ​anterior​ ​prothorax,
middle​ ​mesothorax,​ ​and​ ​posterior​ ​metathorax,​ ​of​ ​which​ ​the​ ​prothorax​​ ​is​ ​by​ ​far​ ​the
largest​ ​and​ ​the​ ​only​ ​one​ ​visible​ ​in​ ​dorsal​ ​view.​ ​ ​ ​In​ ​ventral​ ​view​ ​the​ ​prothorax,
mesothorax​,​ ​and​ ​metathorax​​ ​are​ ​all​ ​visible​ ​and​ ​easily​ ​recognized​ ​by​ ​virtue​ ​of​ ​the​ ​pair
of​ ​walking​ ​legs​​ ​carried​ ​by​ ​each​ ​segment.

Figure​ ​6.​ ​Posterior​ ​view​ ​of​ ​the​ ​labium.

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 Figure​ ​7.​ ​Anterior​ ​view​ ​of​ ​the​ ​hypopharynx​ ​of​ ​P.​ ​americana​.​ ​ ​The​ ​clypeus,​ ​labrum,

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 left​ ​mandible,​ ​and​ ​labium​ ​have​ ​been​ ​removed​ ​for​ ​clarity.

Legs
​ ​The​ ​large​ ​powerful​ ​legs​ ​are​ ​responsible​ ​for​ ​the​ ​cursorial​ ​competency​ ​for​ ​which
cockroaches​ ​are​ ​reknown.
​ ​Three​ ​pairs​ ​are​ ​present,​ ​of​ ​course,​ ​one​ ​on​ ​each​ ​thoracic​ ​segment​ ​(Fig​ ​8).​ ​ ​ ​The​ ​three
pairs​ ​are​ ​similar​ ​but​ ​increase​ ​in​ ​size​ ​from​ ​anterior​ ​to​ ​posterior.​ ​ ​ ​Each​ ​consists​ ​of​ ​a
large,​ ​flattened,​ ​proximal​ ​coxa​,​ ​a​ ​small​ ​trochanter​,​ ​a​ ​long​ ​femur​,​ ​a​ ​tibia​,​ ​and​ ​a​ ​long
tarsus​​ ​(Fig​ ​9,​ ​21-1E).​ ​ ​ ​The​ ​femur​ ​and​ ​tibia​ ​bear​ ​strong​ ​spines​.​ ​The​ ​tarsus​ ​is​ ​a​ ​series
of​ ​5-articulate​ ​tarsomeres​.​ ​Tarsomeres​ ​1-4​​ ​each​ ​bear​ ​a​ ​posterior​ ​pad-like​ ​adhesive
pulvillus​.​ ​Tarsomere​ ​5​ ​ends​ ​with​ ​a​ ​pair​ ​of​ ​tarsal​ ​claws​​ ​beside​ ​a​ ​pad-like​ ​arolium​.
This​ ​distal​ ​arrangement​ ​of​ ​claws​ ​and​ ​arolium​ ​is​ ​sometimes​ ​referred​ ​to​ ​as​ ​the​ ​pretarsus.
The​ ​arolium​ ​is​ ​an​ ​adaptation​ ​for​ ​clinging​ ​to​ ​smooth​ ​surfaces​ ​and​ ​makes​ ​it​ ​possible​ ​for
Periplaneta​ ​to​ ​climb​ ​smooth​ ​walls.

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Figure​ ​8.​ ​Ventral​ ​view​ ​of​ ​a​ ​female​ ​P.​ ​americana​ ​ ​ ​The​ ​distal​ ​articles​ ​of​ ​the​ ​legs​ ​have
been​ ​removed​ ​for​ ​clarity.​ ​S​ ​=​ ​sternite,​ ​T​ ​=​ ​tergite.

When​ ​flexed,​ ​the​ ​femur​ ​fits​ ​neatly​ ​into​ ​a​ ​recess​ ​in​ ​the​ ​surface​ ​of​ ​the​ ​coxa​ ​(Fig​ ​8).
The​ ​articulations​ ​between​ ​coxa​ ​and​ ​trochanter​ ​and​ ​between​ ​femur​ ​and​ ​tibia​ ​are
dicondylic​ ​joints​ ​which​ ​restrict​ ​movement​ ​to​ ​a​ ​single​ ​plane​ ​(as,​ ​you​ ​may​ ​recall,​ ​does​ ​the
dicondylic​ ​joint​ ​between​ ​the​ ​mandible​ ​and​ ​head​ ​capsule).​ ​The​ ​coxa-body​ ​articulation​ ​is
monocondlyic​ ​and​ ​permits​ ​movement​ ​in​ ​a​ ​variety​ ​of​ ​planes​ ​as​ ​is​ ​the​ ​tibia-tarsus
articulation.​ ​Dicondylic​ ​joints​ ​are​ ​functionally​ ​similar​ ​to​ ​the​ ​hinge​ ​joints​ ​in​ ​your​ ​knee​ ​and
elbow​ ​whereas​ ​monocondylic​ ​joints​ ​function​ ​like​ ​ball​ ​and​ ​socket​ ​joints​ ​in​ ​your​ ​shoulder
and​ ​hip.
Wings
​ ​The​ ​meso-​ ​and​ ​metathorax​ ​of​ ​adults​ ​of​ ​both​ ​sexes​ ​of​ ​Periplaneta​ ​each​ ​bear​ ​a
pair​ ​of​ ​large​ ​wings​.​ ​ ​ ​The​ ​anterior​ ​mesothoracic​ ​wings,​ ​or​ ​forewings​​ ​(=​ ​wing​ ​covers,
often​ ​known​ ​as​ ​tegmina​ ​in​ ​orthopterans​ ​and​ ​blattarians),​ ​are​ ​obvious​ ​in​ ​dorsal​ ​view
whereas​ ​the​ ​equally​ ​large​ ​metathoracic​ ​wings​ ​(​hindwings​)​ ​are​ ​almost​ ​entirely​ ​hidden
by​ ​the​ ​forewings​ ​(Fig​ ​10).​ ​The​ ​heavier,​ ​parchment-like​ ​forewings​ ​protect​ ​the​ ​more
delicate​ ​hindwings.​ ​ ​ ​The​ ​hindwings​ ​are​ ​membranous​ ​and​ ​fold​ ​in​ ​pleats​ ​when​ ​at​ ​rest
under​ ​the​ ​forewings.​ ​The​ ​folded​ ​wings​ ​completely​ ​cover​ ​the​ ​dorsal​ ​surface​ ​except​ ​for
the​ ​head​ ​and​ ​pronotum.

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 Fig​ ​9.​ ​Ventral​ ​view​ ​of​ ​the​ ​right​ ​midleg​ ​of​ ​P.​ ​americana

​​
​ ​Use​ ​forceps​ ​and​ ​your​ ​fingers​ ​to​ ​extend​ ​the​ ​wings​ ​and​ ​examine​ ​their​ ​shape​ ​and
texture.​ ​ ​ ​Note​ ​the​ ​way​ ​in​ ​which​ ​the​ ​hindwing​ ​folds​ ​when​ ​at​ ​rest.
​ ​Cockroach​ ​nymphs,​ ​like​ ​immatures​ ​of​ ​other​ ​insects,​ ​lack​ ​wings,​ ​but​ ​like​ ​other
pauro-​ ​and​ ​hemimetabolous​ ​insects,​ ​develop​ ​wing​ ​pads​​ ​in​ ​older​ ​instars​ ​(Fig​ ​11).​ ​The
wing​ ​pads​ ​are​ ​transformed​ ​to​ ​functional​ ​wings​ ​by​ ​the​ ​last​ ​molt.
Thoracic​ ​Segments
​ ​The​ ​thorax​ ​consists​ ​of​ ​three​ ​segments​ ​but​ ​your​ ​view​ ​is​ ​obscured​ ​dorsally​ ​by​ ​the
wings​ ​and​ ​ventrally​ ​by​ ​the​ ​walking​ ​legs,​ ​especially​ ​their​ ​coxae.​ ​ ​ ​Each​ ​segment​ ​has​ ​the
expected​ ​complement​ ​of​ ​sclerites​ ​consisting​ ​of​ ​dorsal​ ​tergite​ ​(or​ ​notum),​ ​lateral
pleurites,​ ​and​ ​ventral​ ​sternites.
​ ​The​ ​ventral​ ​surface​ ​of​ ​the​ ​thorax​ ​consists​ ​of​ ​large​ ​expanses​ ​of​ ​unsclerotized,
flexible,​ ​white​ ​exoskeleton​ ​in​ ​which​ ​are​ ​embedded​ ​myriad​ ​hard,​ ​golden​ ​brown,​ ​sclerites.
Anchor​ ​the​ ​specimen​ ​on​ ​its​ ​back​ ​with​ ​a​ ​#1​ ​stainless​ ​steel​ ​insect​ ​pin​ ​through​ ​the​ ​margin
of​ ​the​ ​pronotum.​ ​ ​ ​Move​ ​the​ ​legs​ ​aside​ ​as​ ​needed​ ​to​ ​reveal​ ​the​ ​ventral​ ​surface​ ​of​ ​the
thorax.​ ​ ​ ​The​ ​proximal​ ​end​ ​of​ ​the​ ​coxa​ ​of​ ​each​ ​thoracic​ ​limb​ ​articulates​ ​with​ ​a​ ​complex
of​ ​sclerotized​ ​pleurites​​ ​(Fig​ ​12,​ ​21-1D).​ ​ ​ ​On​ ​the​ ​midline,​ ​between​ ​the​ ​right​ ​and​ ​left
pleurites,​ ​lies​ ​a​ ​series​ ​of​ ​thoracic​ ​sternites.​ ​ ​ ​The​ ​prosternum​​ ​is​ ​a​ ​small,​ ​triangular,
median,​ ​unpaired​ ​sclerite​ ​in​ ​the​ ​center​ ​of​ ​the​ ​prothorax,​ ​between​ ​the​ ​clusters​ ​of
prothoracic​ ​pleurites.​ ​The​ ​mesosternum​​ ​is​ ​more​ ​complicated​ ​than​ ​the​ ​prosternum​ ​and

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consists​ ​of​ ​several​ ​sclerites​ ​including​ ​a​ ​large​ ​bilobed​ ​anterior​ ​plate​ ​followed​ ​by​ ​two
slender​ ​posterior​ ​sclerites.​ ​ ​ ​The​ ​metasternum​​ ​is​ ​similar​ ​to​ ​the​ ​mesosternum​ ​except
that​ ​the​ ​bilobed​ ​plate​ ​is​ ​completely​ ​divided​ ​into​ ​a​ ​pair​ ​of​ ​plates.
Figure​ ​10.​ ​Dorsal​ ​view​ ​of​ ​a​ ​female​ ​P.​ ​fuliginosa​ ​with​ ​the​ ​wings​ ​removed​ ​or
extended​ ​to​ ​reveal​ ​the​ ​dorsal​ ​surface​ ​of​ ​the​ ​thorax​ ​and​ ​abdomen.​ ​The​ ​abdomen
has​ ​been​ ​stretched​ ​slightly​ ​to​ ​reveal​ ​tergites​ ​8​ ​and​ ​9​ ​which​ ​are​ ​normally​ ​hidden
under​ ​tergite​ ​7.​ ​blatt46L.gif

​ ​The​ ​thorax​ ​is​ ​equipped​ ​with​ ​two​ ​pairs​ ​of​ ​spiracles​ ​which​ ​open​ ​into​ ​the​ ​tracheal
respiratory​ ​system.​ ​The​ ​spiracles​​ ​are​ ​lateral​ ​and​ ​belong​ ​to​ ​the​ ​mesothorax​ ​and
metathorax​ ​although​ ​they​ ​have​ ​migrated​ ​anteriorly​ ​(Fig​ ​12).​ ​ ​ ​Consequently,​ ​the
mesothoracic​ ​spiracle​​ ​is​ ​on​ ​the​ ​posterior​ ​prothorax​ ​and​ ​the​ ​metathoracic​ ​spiracle​​ ​is
in​ ​the​ ​membrane​ ​between​ ​the​ ​mesothorax​ ​and​ ​metathorax.
​ ​Study​ ​the​ ​dorsal​ ​thorax​ ​by​ ​moving​ ​the​ ​wings​ ​aside.​ ​ ​ ​The​ ​pronotum​​ ​is​ ​not
obscured​ ​by​ ​wings​ ​and​ ​is​ ​always​ ​easily​ ​seen.​ ​ ​ ​It​ ​forms​ ​a​ ​large​ ​shield​ ​behind​ ​the​ ​head
and​ ​may​ ​be​ ​mistaken​ ​for​ ​the​ ​head​ ​by​ ​the​ ​uninitiated.​ ​ ​ ​It​ ​is​ ​heavily​ ​sclerotized​ ​and​ ​dark
in​ ​color​ ​with​ ​a​ ​pale​ ​border.​ ​ ​ ​The​ ​mesonotum​​ ​(Fig​ ​10)​ ​is​ ​a​ ​transverse,​ ​lightly​ ​sclerotized
plate​ ​covering​ ​the​ ​dorsal​ ​surface​ ​of​ ​the​ ​mesothorax.​ ​The​ ​metanotum​​ ​is​ ​similar​ ​and
covers​ ​the​ ​mesothorax.
Abdomen
​ ​The​ ​abdomen​​ ​is​ ​the​ ​largest​ ​of​ ​the​ ​three​ ​tagmata​ ​and​ ​contains​ ​most​ ​of​ ​the

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viscera.​ ​ ​ ​Its​ ​segmentation​ ​is​ ​conspicuous​ ​both​ ​dorsally​ ​and​ ​ventrally​ ​once​ ​the​ ​wings
have​ ​been​ ​removed.​ ​ ​ ​The​ ​abdomen​ ​consists​ ​of​ ​11​ ​segments​ ​of​ ​which​ ​the​ ​11​ th​ ​ ​ ​is
reduced​ ​and​ ​fused​ ​with​ ​the​ ​10​ th​ ​ ​ ​so​ ​that​ ​a​ ​maximum​ ​of​ ​ten​ ​segments​ ​are​ ​distinct​ ​and
visible​ ​(Fig​ ​21-1F).​ ​Further,​ ​some​ ​of​ ​the​ ​segments,​ ​especially​ ​in​ ​females,​ ​are​ ​highly
modified​ ​or​ ​in​ ​unexpected​ ​locations​ ​and​ ​are​ ​not​ ​immediately​ ​visible​ ​and​ ​countable.
Figure​ ​11.​ ​A​ ​late​ ​instar,​ ​24​ ​mm​ ​Periplaneta​ ​nymph.​ ​Blatt47L.gif

​ ​The​ ​first​ ​seven​ ​segments,​ ​known​ ​as​ ​the​ ​pregenital,​ ​or​ ​visceral,​ ​segments​ ​are
similar,​ ​unspecialized,​ ​visible​ ​and​ ​countable.​ ​ ​ ​Segments​ ​9​ ​in​ ​males​ ​and​ ​8-9​ ​in​ ​females
are​ ​the​ ​genital​ ​segments​ ​and​ ​are​ ​modified​ ​for​ ​reproductive​ ​functions.​ ​ ​ ​Segment​ ​10​ ​(and
the​ ​vestigial​ ​11)​ ​are​ ​postgenital​ ​and​ ​are​ ​also​ ​modified.
​ ​Dorsally​ ​each​ ​segment​ ​is​ ​covered​ ​by​ ​a​ ​sclerotized​ ​tergite​​ ​and,​ ​similarly,​ ​is
covered​ ​ventrally​ ​by​ ​a​ ​sclerotized​ ​sternite​.​ ​ ​ ​Consecutive​ ​sternites​ ​articulate​ ​with​ ​each
other​ ​by​ ​unsclerotized​ ​articular​ ​membranes​​ ​(Fig​ ​15).​ ​ ​ ​Laterally,​ ​tergites​ ​and​ ​sternites
are​ ​connected​ ​by​ ​unsclerotized​ ​flexible​ ​pleura​.
​ ​Examine​ ​the​ ​dorsum​ ​of​ ​the​ ​abdomen​ ​and​ ​find​ ​the​ ​abdominal​ ​tergites​​ ​(Fig​ ​10).
Those​ ​of​ ​segments​ ​1-7​ ​are​ ​similar​ ​in​ ​size​ ​and​ ​easily​ ​distinguished​ ​but​ ​tergite​ ​8​ ​is
smaller​ ​and​ ​tergite​ ​9​ ​smaller​ ​still.​ ​ ​ ​These​ ​two​ ​tergites​ ​are​ ​usually​ ​telescoped​ ​under
tergite​ ​7​ ​and​ ​may​ ​be​ ​difficult​ ​to​ ​see​ ​until​ ​you​ ​lift​ ​tergite​ ​7.​ ​Tergite​ ​10​​ ​is​ ​a​ ​large
posteriorly​ ​bilobed​ ​plate​ ​overhanging​ ​the​ ​anus​ ​(Fig​ ​10).​ ​ ​Tergite​ ​11​ ​is​ ​fused​ ​with​ ​tergite

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10.​ ​The​ ​bilobed​ ​posterior​ ​extension​ ​of​ ​tergite​ ​10​ ​is​ ​sometimes​ ​referred​ ​to​ ​as​ ​the
epiproct​,​ ​which​ ​is​ ​homologous​ ​to​ ​the​ ​tergite​ ​of​ ​segment​ ​11​ ​(Fig​ ​10,​ ​21-1F).
​ ​Study​ ​the​ ​ventral​ ​surface​ ​of​ ​the​ ​abdomen​ ​(Fig​ ​12,​ ​H).​ ​ ​ ​Sternite​ ​1​ ​is​ ​a​ ​small
sclerite​ ​lying​ ​on​ ​the​ ​midline​ ​between​ ​the​ ​hindleg​ ​coxae.​ ​ ​ ​Sternites​ ​2-6​​ ​are​ ​elongate,
more​ ​or​ ​less​ ​rectangular,​ ​sclerotized​ ​plates​ ​covering​ ​the​ ​ventral​ ​surface​ ​of​ ​the
abdomen.​ ​ ​ ​Sternites​ ​7-9​​ ​of​ ​males​ ​resemble​ ​sternites​ ​2-6​ ​but​ ​the​ ​posterior​ ​sternites​ ​of
females​ ​are​ ​very​ ​different.​ ​ ​ ​In​ ​females​ ​the​ ​posteriormost​ ​visible​ ​sternite​ ​is​ ​the​ ​large,
highly​ ​modified​ ​sternite​ ​7​ ​(Fig​ ​12,​ ​13).​ ​ ​ ​Sternites​ ​8,​ ​9​ ​are​ ​internalized​ ​under​ ​tergite​ ​7
and​ ​cannot​ ​be​ ​seen​ ​without​ ​dissection.
Figure​ ​12.​ ​Ventral​ ​view​ ​of​ ​a​ ​female​ ​P.​ ​fuliginosa​ ​with​ ​the​ ​right​ ​legs​ ​removed​ ​to
reveal​ ​the​ ​ventral​ ​body​ ​surface​ ​and​ ​its​ ​sclerites.​ ​ ​ ​ ​The​ ​position​ ​of​ ​the​ ​abdominal
spiracles​ ​under​ ​the​ ​overhang​ ​of​ ​the​ ​preceding​ ​sternite​ ​is​ ​indicated.​ ​Blatt48L.gif

​​​

​ ​The​ ​anus​​ ​is​ ​on​ ​segment​ ​11​ ​and​ ​lies​ ​ventral​ ​to​ ​the​ ​epiproct​ ​(tergite​ ​11).​ ​It​ ​is
flanked​ ​by​ ​a​ ​pair​ ​of​ ​triangular​ ​sclerites,​ ​the​ ​paraprocts​​ ​(Fig​ ​14,​ ​21-11B,​ ​21-12B,C)
which​ ​are​ ​modifications​ ​of​ ​sternite​ ​11.​ ​ ​ ​Sternite​ ​10​ ​is​ ​vestigial.
​ ​Most​ ​abdominal​ ​segments​ ​lack​ ​appendages​ ​but​ ​segment​ ​10​ ​bears​ ​a​ ​pair​ ​of
large,​ ​segmented​ ​cerci​,​ ​which​ ​are​ ​visible​ ​in​ ​both​ ​dorsal​ ​and​ ​ventral​ ​views​ ​(Fig​ ​8,​ ​13,
21-1F).​ ​ ​ ​Embryologically​ ​the​ ​cerci​ ​arise​ ​as​ ​the​ ​appendages​ ​of​ ​the​ ​11​ ​segment​ ​but​ ​in
adults​ ​are​ ​attached​ ​to​ ​segment​ ​10​ ​since​ ​segment​ ​11​ ​has​ ​fused​ ​with​ ​segment​ ​10.​ ​The
cercus,​ ​although​ ​segmented,​ ​does​ ​not​ ​contain​ ​intrinsic​ ​muscles.​ ​Extrinsic​ ​muscles,
however,​ ​extend​ ​from​ ​the​ ​cercus​ ​to​ ​the​ ​abdomen​ ​so​ ​it​ ​is​ ​capable​ ​of​ ​motion.​ ​ ​ ​Cerci​ ​bear
vibration-sensitive​ ​hair​ ​sensillae​ ​sensitive​ ​to​ ​air​ ​movements,​ ​including​ ​sound,​ ​and
perhaps​ ​to​ ​ground​ ​vibrations.
​ ​Eight​ ​pairs​ ​of​ ​spiracles​ ​are​ ​present​ ​in​ ​the​ ​abdomen​ ​and​ ​are​ ​much​ ​smaller​ ​than
the​ ​thoracic​ ​spiracles.​ ​The​ ​abdominal​​ ​spiracles​​ ​are​ ​located​ ​laterally​ ​on​ ​the

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anterior-dorsal​ ​corner​ ​of​ ​the​ ​pleura​ ​of​ ​the​ ​first​ ​eight​ ​abdominal​ ​segments​ ​(Fig​ ​12,​ ​14,
21-1D,F).​ ​ ​ ​Lift​ ​the​ ​free​ ​posterior​ ​edge​ ​of​ ​a​ ​sternite​ ​and​ ​look​ ​at​ ​the​ ​antero-lateral​ ​corner
of​ ​the​ ​following​ ​tergite​ ​to​ ​see​ ​the​ ​spiracle.​ ​It​ ​will​ ​be​ ​a​ ​small​ ​white​ ​oval.​ ​ ​ ​Abdominal
spiracle​ ​1​ ​is​ ​on​ ​the​ ​lateral​ ​edge​ ​of​ ​tergite​ ​1​ ​rather​ ​than​ ​at​ ​the​ ​corner​ ​(Fig​ ​12).

Genital​ ​Segments
Female
​ ​Study​ ​the​ ​dorsum​ ​of​ ​the​ ​posterior​ ​abdomen​ ​of​ ​a​ ​female.​ ​ ​ ​The​ ​tergites​ ​of​ ​the
genital​ ​segments​​ ​(abdominal​ ​segments​ ​8,​ ​9)​ ​are​ ​reduced​ ​and​ ​their​ ​tergites​ ​are​ ​largely
hidden​ ​by​ ​the​ ​overhang​ ​of​ ​tergite​ ​7​ ​(Fig​ ​10).​ ​ ​ ​Tergite​ ​10​​ ​(fused​ ​with​ ​tergite​ ​11)​ ​is​ ​a
large,​ ​thin,​ ​fan-like​ ​plate​ ​extending​ ​posterior​ ​to​ ​the​ ​genital​ ​segments.​ ​It​ ​overhangs​ ​the
anus​ ​and​ ​genital​ ​pouch,​ ​which​ ​will​ ​be​ ​described​ ​shortly.​ ​ ​ ​In​ ​P.​ ​americana​ ​tergite​ ​10​ ​is
cleft​ ​posteriorly​ ​(Fig​ ​10).
Figure​ ​13.​ ​Ventral​ ​view​ ​of​ ​the​ ​posterior​ ​end​ ​of​ ​the​ ​abdomen​ ​of​ ​a​ ​female​ ​of​ ​P.
fuliginosa​.​ ​Blatt49L.gif

​ ​Look​ ​at​ ​the​ ​ventral​ ​surface​ ​of​ ​the​ ​abdomen​ ​(Fig​ ​13).​ ​Sternites​ ​8​ ​and​ ​9​ ​cannot​ ​be

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seen​ ​externally.​ ​ ​ ​Sternite​ ​7​ ​(=​ ​hypogynum)​ ​is​ ​much​ ​larger​ ​and​ ​more​ ​heavily​ ​sclerotized
than​ ​other​ ​sternites​ ​and​ ​posses​ ​a​ ​pair​ ​of​ ​clamshell-like​ ​posterior​ ​valves,​ ​the​ ​subgenital
plates​ ​(also​ ​referred​ ​to​ ​variously​ ​as​ ​sternites​ ​7​ ​¢​ ​or​ ​7a).​ ​ ​ ​Although​ ​at​ ​first​ ​the​ ​plates
may​ ​appear​ ​to​ ​be​ ​independent​ ​of​ ​sternite​ ​7,​ ​they​ ​are​ ​in​ ​fact​ ​continuous​ ​with​ ​it​ ​and
connected​ ​by​ ​a​ ​short​ ​isthmus​ ​(Fig​ ​13,​ ​14).​ ​ ​ ​The​ ​isthmus​ ​is​ ​flexible​ ​and​ ​the​ ​plates​ ​can
move​ ​apart​ ​during​ ​copulation​ ​or​ ​to​ ​permit​ ​extrusion​ ​of​ ​the​ ​ootheca​ ​as​ ​it​ ​is​ ​formed.
​ ​Hold​ ​the​ ​specimen​ ​upright​ ​and​ ​focus​ ​on​ ​the​ ​posterior​ ​end.​ ​ ​ ​This​ ​is​ ​much​ ​easier
if​ ​you​ ​first​ ​separate​ ​the​ ​posterior​ ​abdomen​ ​from​ ​the​ ​remainder​ ​of​ ​the​ ​body​ ​(Do​ ​not
remove​ ​the​ ​posterior​ ​abdomen​ ​if​ ​you​ ​plan​ ​to​ ​use​ ​this​ ​specimen​ ​later​ ​for​ ​study​ ​of​ ​internal
anatomy).​ ​ ​ ​Make​ ​the​ ​separation​ ​between​ ​segment​ ​6​ ​and​ ​7.​ ​ ​ ​You​ ​can​ ​now​ ​stand​ ​the
posterior​ ​abdomen​ ​upright​ ​on​ ​the​ ​wax​ ​of​ ​the​ ​dissecting​ ​pan​ ​and​ ​you​ ​will​ ​not​ ​have​ ​to
hold​ ​it​ ​in​ ​position.​ ​ ​ ​Focus​ ​on​ ​the​ ​posterior​ ​end​ ​of​ ​the​ ​abdomen​ ​(Fig​ ​14).
Figure​ ​14.​ ​Posterior​ ​view​ ​of​ ​the​ ​abdomen​ ​of​ ​a​ ​female​ ​P.​ ​fuliginosa​.​ ​Arrows​ ​indicate
movement​ ​of​ ​the​ ​subgenital​ ​plates.​ ​Blatt50L.gif

​ ​Tergites​ ​7-10​ ​dorsally​ ​and​ ​sternite​ ​7​ ​ventrally​ ​enclose​ ​a​ ​large​ ​space.​ ​ ​ ​The
space​ ​is​ ​divided​ ​into​ ​dorsal​ ​and​ ​ventral​ ​regions​ ​by​ ​two​ ​lateral​ ​sclerites,​ ​the​ ​paraprocts
(Fig​ ​14,​ ​21-11B).​ ​The​ ​anus​ ​opens​ ​into​ ​the​ ​dorsal​ ​chamber​ ​whereas​ ​the​ ​oviduct​ ​and
seminal​ ​receptacle​ ​open​ ​into​ ​the​ ​ventral​ ​chamber,​ ​which​ ​is​ ​the​ ​genital​ ​pouch​ ​(Fig​ ​15).
With​ ​forceps​ ​pull​ ​the​ ​paraprocts​ ​aside​ ​to​ ​reveal​ ​the​ ​anus​​ ​on​ ​segment​ ​10-11​ ​under​ ​the
epiproct.
The​ ​genital​ ​pouch​ ​is​ ​formed​ ​by​ ​invagination​ ​of​ ​the​ ​exoskeleton​ ​of​ ​the​ ​sternum
which​ ​brings​ ​sternites​ ​8​ ​and​ ​9​ ​deep​ ​into​ ​the​ ​pouch​ ​(Fig​ ​15).​ ​ ​ ​This​ ​is​ ​the​ ​reason​ ​you
could​ ​not​ ​see​ ​these​ ​sternites​ ​externally.​ ​ ​ ​With​ ​forceps​ ​and​ ​needles​ ​as​ ​needed,​ ​pull​ ​the

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subgenital​ ​plates​ ​of​ ​sternite​ ​7​ ​laterally​ ​and​ ​ventrally​ ​to​ ​reveal​ ​the​ ​genital​ ​pouch​​ ​(=
gynatrium).​ ​ ​ ​The​ ​genital​ ​pouch​ ​receives​ ​the​ ​spermatophore​ ​from​ ​the​ ​male​ ​during
copulation​ ​and​ ​later​ ​the​ ​ootheca​ ​is​ ​molded​ ​in​ ​it.
The​ ​most​ ​conspicuous​ ​feature​ ​in​ ​the​ ​genital​ ​pouch​ ​is​ ​the​ ​ovipositor​​ ​(Fig​ ​15).​ ​It​ ​is
a​ ​median​ ​process​ ​arising​ ​from​ ​the​ ​roof​ ​of​ ​the​ ​genital​ ​pouch​ ​and​ ​consists​ ​of​ ​three​ ​pairs
of​ ​cuticularized​ ​valvulae​​ ​(=​ ​gonapophyses).​ ​ ​ ​The​ ​valvulae​ ​are​ ​derived​ ​from​ ​the​ ​paired
appendages​ ​of​ ​segments​ ​8​ ​and​ ​9.​ ​The​ ​ovipositor​ ​guides​ ​eggs​ ​from​ ​the​ ​oviduct​ ​to​ ​the
forming​ ​ootheca.​ ​ ​ ​The​ ​male​ ​external​ ​genitalia​ ​grip​ ​the​ ​ovipositor​ ​during​ ​copulation.
Sternite​ ​8​ ​can​ ​be​ ​seen​ ​by​ ​lifting​ ​the​ ​ovipositor​ ​dorsally​ ​and​ ​looking​ ​under​ ​it​ ​Fig
15,​ ​23).​ ​ ​ ​Sternite​ ​8​ ​is​ ​a​ ​large​ ​sclerotized​ ​plate​ ​penetrated​ ​by​ ​the​ ​female​ ​gonopore
opening​ ​from​ ​the​ ​common​ ​gonoduct.​ ​ ​ ​Sternite​ ​9​ ​is​ ​at​ ​the​ ​base​ ​of​ ​the​ ​ovipositor.
Figure​ ​15.​ ​Sagittal​ ​section​ ​of​ ​the​ ​posterior​ ​abdomen​ ​of​ ​a​ ​generalized​ ​female
cockroach.​ ​ ​ ​S​ ​=​ ​sternite,​ ​T​ ​=​ ​tergite.​ ​Redrawn​ ​from​ ​Cornwell​ ​(1968)​ ​after
McKittrick​ ​(1964).​ ​Blatt51L.gif

Male
The​ ​dorsum​ ​of​ ​the​ ​male​ ​posterior​ ​abdomen​ ​is​ ​similar​ ​to​ ​that​ ​of​ ​the​ ​female.
Tergite​ ​7​ ​is​ ​large​ ​and​ ​tergites​ ​8​ ​and​ ​9​ ​are​ ​much​ ​smaller​ ​and​ ​obscured​ ​by​ ​the​ ​overhang
of​ ​7​ ​(Fig​ ​16).​ ​ ​ ​Tergite​ ​10​​ ​is​ ​a​ ​large,​ ​posteriorly​ ​cleft​ ​plate​ ​fused​ ​with​ ​tergite​ ​11​ ​to​ ​form
the​ ​epiproct.
Figure​ ​16.​ ​Dorsal​ ​view​ ​of​ ​the​ ​posterior​ ​abdomen​ ​of​ ​P.​ ​americana​.​ ​T​ ​=​ ​tergite.
Blatt52L.gif

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 ​​
The​ ​male​ ​venter​ ​is​ ​very​ ​different​ ​from​ ​that​ ​of​ ​females.​ ​Sternites​ ​8​,​ ​and​ ​9​ ​are
unremarkable​ ​and​ ​unmodified​ ​from​ ​the​ ​condition​ ​of​ ​the​ ​more​ ​anterior​ ​sternites​ ​and
visible​ ​externally​ ​(Fig​ ​17).​ ​Sternite​ ​9​ ​is​ ​the​ ​posteriormost​ ​sternite​ ​and​ ​its​ ​posterior
border​ ​bears​ ​a​ ​slender​ ​stylus​​ ​on​ ​each​ ​side.​ ​Styli​ ​are​ ​absent​ ​in​ ​mature​ ​females​ ​but​ ​are
present​ ​in​ ​nymphs​ ​of​ ​both​ ​sexes​ ​(Fig​ ​21-12B).
A​ ​pair​ ​of​ ​triangular​ ​paraprocts​​ ​is​ ​located​ ​immediately​ ​ventral​ ​to​ ​tergite​ ​10-11​ ​and
lateral​ ​to​ ​the​ ​anus​ ​(​Fig​ ​14).​ ​ ​ ​Between​ ​the​ ​paraprocts​ ​and​ ​sternite​ ​9​ ​is​ ​a​ ​large​ ​genital
pouch​​ ​in​ ​which​ ​are​ ​housed​ ​the​ ​male​ ​external​ ​genitalia.
Figure​ ​17.​ ​Ventral​ ​view​ ​of​ ​the​ ​posterior​ ​abdomen​ ​of​ ​a​ ​male​ ​P.​ ​americana​.
Blatt53L.gif

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 ​​
Anchor​ ​the​ ​posterior​ ​abdomen,​ ​ventral​ ​side​ ​up,​ ​with​ ​a​ ​pair​ ​of​ ​pins​ ​passing
through​ ​the​ ​lateral​ ​tergites​ ​of​ ​segment​ ​7.​ ​ ​ ​Lift​ ​or​ ​remove​ ​sternites​ ​8​ ​and​ ​9​ ​to​ ​reveal​ ​the
genital​ ​pouch​ ​and​ ​its​ ​contents​ ​(Fig​ ​18,​ ​24).​ ​ ​ ​The​ ​male​ ​external​ ​genitalia​ ​comprise​ ​the
several​ ​complex​ ​sclerites​ ​and​ ​soft​ ​tissues​ ​of​ ​three​ ​phallomeres​​ ​(=​ ​phallic​ ​lobes,​ ​=
gonapophyses),​ ​surrounding​ ​the​ ​male​ ​gonopore.​ ​The​ ​phallomeres​ ​are​ ​responsible​ ​for
moving​ ​the​ ​subgenital​ ​plates​ ​aside​ ​to​ ​open​ ​the​ ​female’s​ ​genital​ ​pouch​ ​and​ ​for​ ​holding
the​ ​female​ ​genitalia​ ​during​ ​copulation.​ ​ ​ ​Muscles​ ​arising​ ​on​ ​segment​ ​9​ ​extend​ ​to​ ​the
phallomeres​ ​and​ ​operate​ ​them.​ ​Phallomeres​ ​are​ ​the​ ​appendages​ ​of​ ​segment​ ​9.
Viewed​ ​from​ ​the​ ​venter,​ ​the​ ​ventral​ ​phallomere​​ ​(Fig​ ​18)​ ​is​ ​the​ ​most​ ​obvious​ ​of
the​ ​three​ ​and​ ​can​ ​be​ ​seen​ ​immediately​ ​under​ ​sternite​ ​9​ ​(which​ ​has​ ​been​ ​removed​ ​or
lifted).​ ​ ​ ​It​ ​is​ ​ventral​ ​to​ ​the​ ​gonopore​ ​and​ ​is​ ​by​ ​far​ ​the​ ​simplest​ ​of​ ​the​ ​three​ ​lobes.​ ​ ​ ​Its
ventral​ ​surface​ ​is​ ​a​ ​slightly​ ​cupped​ ​sclerotized​ ​plate​ ​but​ ​it​ ​is​ ​membranous​ ​dorsally.​ ​The
membranous​ ​ejaculatory​ ​duct​ ​ends​ ​at​ ​the​ ​gonopore​ ​on​ ​the​ ​dorsal​ ​surface​ ​of​ ​its​ ​base.
Because​ ​it​ ​bears​ ​the​ ​gonopore,​ ​it​ ​is​ ​sometimes​ ​known​ ​as​ ​the​ ​penis.
The​ ​left​ ​phallomere​​ ​is​ ​left​ ​of​ ​the​ ​ventral​ ​lobe​ ​and​ ​a​ ​little​ ​dorsal​ ​to​ ​it​ ​(Fig​ ​18).​ ​ ​ ​It
is​ ​a​ ​complex​ ​structure​ ​consisting​ ​of​ ​many​ ​sclerotized​ ​pieces​ ​including​ ​a​ ​hooked​ ​lobe​ ​(=
grumolobus,​ ​titillator),​ ​prickly​ ​lobe​ ​(=​ ​acantholobus),​ ​a​ ​sharp​ ​stylet​ ​(=​ ​acutolobus,
asperate​ ​lobe),​ ​and​ ​a​ ​pseudopenis.​ ​ ​ ​These​ ​are​ ​used​ ​during​ ​copulation​ ​to​ ​hold​ ​the

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valvulae​ ​of​ ​the​ ​female​ ​ovipositor​ ​so​ ​the​ ​spermatophore​ ​can​ ​be​ ​deposited​ ​in​ ​the​ ​correct
location​ ​(at​ ​the​ ​opening​ ​of​ ​the​ ​seminal​ ​receptacle).
Figure​ ​18.​ ​Ventral​ ​view​ ​of​ ​the​ ​posterior​ ​abdominal​ ​segments​ ​of​ ​a​ ​male​ ​P.
americana​.​ ​Sternites​ ​8-9​ ​have​ ​been​ ​removed​ ​and​ ​the​ ​phallomeres​ ​rearranged
slightly​ ​for​ ​clarity.​ ​ ​ ​Some​ ​membranes​ ​and​ ​connective​ ​tissue​ ​have​ ​been​ ​removed.
S​ ​=​ ​sternite,​ ​T​ ​=​ ​tergite.​ ​Blatt54L.gif

The​ ​right​ ​phallomere​​ ​is​ ​to​ ​the​ ​right​ ​of​ ​the​ ​ventral​ ​phallomere​ ​and​ ​gonopore​ ​and
is​ ​farther​ ​dorsal​ ​than​ ​either​ ​of​ ​the​ ​other​ ​two​ ​lobes.​ ​It​ ​is​ ​less​ ​complicated​ ​than​ ​the​ ​left
lobe​ ​but​ ​also​ ​has​ ​easily​ ​recognized​ ​sclerotized​ ​parts.​ ​ ​ ​ ​These​ ​include​ ​a​ ​piece​ ​that​ ​looks
like​ ​a​ ​crayfish​ ​cheliped​ ​(=​ ​dikella,​ ​serrate​ ​lobe)​ ​and​ ​slender​ ​hooked​ ​piece​ ​(=​ ​falx).

Internal​ ​Anatomy
​ ​The​ ​cricket,​ ​Acheta​,​ ​is​ ​recommended​ ​as​ ​an​ ​alternative​ ​for​ ​the​ ​study​ ​of​ ​internal
anatomy​ ​in​ ​preference​ ​to​ ​the​ ​cockroach.​ ​ ​ ​Even​ ​though​ ​it​ ​is​ ​smaller,​ ​it​ ​is​ ​easier​ ​to
dissect​ ​and​ ​its​ ​internal​ ​structures​ ​are​ ​easier​ ​to​ ​demonstrate.
​ ​Begin​ ​the​ ​study​ ​of​ ​cockroach​ ​internal​ ​anatomy​ ​with​ ​a​ ​fresh,​ ​undamaged
specimen​ ​if​ ​possible.​ ​ ​ ​The​ ​dissection​ ​will​ ​be​ ​made​ ​from​ ​the​ ​dorsal​ ​side​ ​so​ ​you​ ​must
first​ ​remove​ ​the​ ​wings​ ​by​ ​cutting​ ​their​ ​attachments​ ​with​ ​a​ ​pair​ ​of​ ​fine​ ​scissors.​ ​The

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dissection​ ​is​ ​facilitated​ ​if​ ​the​ ​legs​ ​are​ ​removed​ ​by​ ​cutting​ ​across​ ​their​ ​trochanters.
Organ​ ​systems​ ​will​ ​be​ ​considered​ ​in​ ​order​ ​of​ ​their​ ​appearance​ ​in​ ​dorsal​ ​dissection.
" ​ ​Place​ ​the​ ​specimen​ ​in​ ​a​ ​small​ ​dissecting​ ​pan​ ​of​ ​water​ ​or​ ​alcohol​ ​so​ ​it​ ​is
completely​ ​immersed.​ ​Insert​ ​the​ ​blade​ ​of​ ​a​ ​pair​ ​of​ ​fine​ ​scissors​ ​under​ ​the​ ​posterior
overhang​ ​of​ ​the​ ​right​ ​side​ ​of​ ​tergite​ ​7​ ​about​ ​1​ ​mm​ ​from​ ​the​ ​lateral​ ​margin​ ​of​ ​the​ ​tergite.
Insert​ ​the​ ​blade​ ​only​ ​as​ ​deep​ ​as​ ​is​ ​necessary​ ​to​ ​penetrate​ ​the​ ​exoskeleton​ ​and​ ​be
careful​ ​you​ ​do​ ​not​ ​damage​ ​internal​ ​organs​ ​with​ ​deep​ ​cuts.​ ​Cut​ ​anteriorly​ ​along​ ​the​ ​right
side​ ​of​ ​the​ ​tergites​ ​all​ ​the​ ​way​ ​to​ ​the​ ​anterior​ ​end​ ​of​ ​the​ ​pronotum​ ​(Fig​ ​19).​ ​ ​ ​Cut
transversely​ ​across​ ​the​ ​anterior​ ​margin​ ​of​ ​the​ ​pronotum,​ ​just​ ​posterior​ ​to​ ​the​ ​head,​ ​and
upon​ ​reaching​ ​the​ ​left​ ​side,​ ​change​ ​directions​ ​and​ ​cut​ ​posteriorly​ ​along​ ​the​ ​left​ ​side​ ​all
the​ ​way​ ​back​ ​to​ ​tergite​ ​7.​ ​ ​ ​Make​ ​a​ ​transverse​ ​cut​ ​through​ ​the​ ​exoskeleton​ ​across​ ​the
posterior​ ​border​ ​of​ ​tergite​ ​7.​ ​You​ ​have​ ​now​ ​cut​ ​all​ ​of​ ​the​ ​way​ ​around​ ​the​ ​dorsum.
Figure​ ​19.​ ​Dorsal​ ​view​ ​of​ ​a​ ​male​ ​P.​ ​americana​.​ ​ ​ ​Thoracic​ ​nota​ ​and​ ​abdominal​ ​terga
1-7​ ​have​ ​been​ ​removed.​ ​T​ ​=​ ​tergite.​ ​Blatt55L.gif

​​
​ ​Anchor​ ​the​ ​specimen​ ​to​ ​the​ ​wax​ ​of​ ​the​ ​dissecting​ ​pan​ ​with​ ​a​ ​#1​ ​stainless​ ​steel
insect​ ​pin​ ​through​ ​the​ ​left​ ​side​ ​of​ ​one​ ​of​ ​the​ ​thoracic​ ​segments,​ ​lateral​ ​to​ ​your​ ​incision.
Insert​ ​the​ ​pin​ ​at​ ​a​ ​45​​ ​°​ ​angle.​ ​ ​ ​Remove​ ​the​ ​abdominal​ ​and​ ​thoracic​ ​tergites​ ​beginning
with​ ​tergite​ ​7.​ ​ ​ ​Lift​ ​each​ ​tergite​ ​and​ ​remove​ ​it​ ​without​ ​removing​ ​any​ ​of​ ​the​ ​underlying
soft​ ​tissues.​ ​ ​ ​The​ ​tergites​ ​are​ ​transparent​ ​and​ ​thin​ ​whereas​ ​the​ ​underlying​ ​tissues​ ​are
opaque.​ ​These​ ​tissues​ ​include​ ​muscles,​ ​heart,​ ​and​ ​tracheae.​ ​ ​ ​Do​ ​not​ ​remove​ ​the
opaque​ ​tissue​ ​at​ ​this​ ​time.
​ ​In​ ​the​ ​abdomen​ ​the​ ​tergal​ ​muscles​​ ​form​ ​a​ ​broad​ ​thin​ ​sheet​ ​of​ ​longitudinal
muscle​ ​fibers​ ​(Fig​ ​19).​ ​ ​ ​In​ ​the​ ​thorax​ ​the​ ​tergal​ ​muscles​ ​are​ ​smaller​ ​and​ ​confined​ ​by
other​ ​muscles,​ ​chiefly​ ​those​ ​operating​ ​the​ ​legs.​ ​Running​ ​along​ ​the​ ​midline​ ​of​ ​this
muscle​ ​layer​ ​is​ ​the​ ​heart.​ ​ ​ ​If​ ​you​ ​accidentally​ ​remove​ ​the​ ​muscle​ ​layer,​ ​the​ ​heart​ ​will​ ​be
destroyed,​ ​so​ ​be​ ​careful.

God​ ​gives​ ​knowledge

 Hemal​ ​System
​ ​The​ ​hemal​ ​system​​ ​is​ ​the​ ​first​ ​organ​ ​system​ ​uncovered​ ​in​ ​dorsal​ ​dissection​ ​(Fig
19).​ ​ ​ ​It​ ​consists​ ​of​ ​the​ ​hemocoel,​ ​blood,​ ​and​ ​heart.​ ​ ​ ​The​ ​hemocoel​ ​is​ ​divided​ ​by
perforated​ ​horizontal​ ​membranes,​ ​known​ ​as​ ​diaphragms,​ ​into​ ​a​ ​shallow​ ​dorsal
pericardial​ ​sinus,​ ​a​ ​spacious​ ​perivisceral​ ​sinus,​ ​and​ ​a​ ​small​ ​ventral​ ​perineural​ ​sinus​ ​(Fig
16-7).​ ​Together​ ​the​ ​sinuses​ ​form​ ​a​ ​large​ ​partitioned​ ​blood​ ​space,​ ​the​ ​hemocoel.​ ​The
hemocoel​ ​is​ ​the​ ​functional​ ​body​ ​cavity​ ​of​ ​these​ ​acoelomate​ ​animals.
​ ​The​ ​heart,​ ​surrounded​ ​by​ ​the​ ​pericardial​ ​sinus​ ​(=​ ​pericardial​ ​hemocoel)​ ​lies
immediately​ ​under​ ​the​ ​tergites.​ ​ ​ ​Removal​ ​of​ ​the​ ​tergites​ ​opens​ ​the​ ​shallow​ ​pericardial
sinus​,​ ​although​ ​it​ ​is​ ​not​ ​at​ ​all​ ​obvious​ ​that​ ​you​ ​are​ ​looking​ ​at​ ​a​ ​blood​ ​space.​ ​ ​ ​It​ ​is​ ​the
space​ ​between​ ​the​ ​exoskeleton​ ​and​ ​the​ ​dorsal​ ​diaphragm.​ ​ ​ ​The​ ​perforated,​ ​horizontal
dorsal​ ​diaphragm​​ ​is​ ​the​ ​sheet​ ​of​ ​connective​ ​tissue​ ​with​ ​longitudinal​ ​tergal​ ​muscles​​ ​on
top​ ​of​ ​it​ ​(Fig​ ​-16-7).​ ​ ​ ​This​ ​diaphragm​ ​separates​ ​the​ ​pericardial​ ​sinus​ ​from​ ​the​ ​much
larger​ ​perivisceral​ ​sinus.​ ​ ​ ​These​ ​two​ ​sinuses​ ​are​ ​part​ ​of​ ​the​ ​hemocoel.​ ​Blood​ ​on​ ​its​ ​way
back​ ​to​ ​the​ ​heart​ ​flows​ ​through​ ​the​ ​perforations​ ​in​ ​the​ ​diaphragm.
​ ​The​ ​heart​​ ​is​ ​a​ ​longitudinal​ ​middorsal​ ​tube​ ​extending​ ​the​ ​length​ ​of​ ​the​ ​body​ ​in​ ​the
pericardial​ ​sinus​ ​and​ ​resting​ ​on​ ​the​ ​dorsal​ ​diaphragm​ ​(Fig​ ​19).​ ​ ​ ​The​ ​heart​ ​has
segmental​ ​swellings,​ ​paired​ ​segmental​ ​ostia,​ ​paired​ ​segmental​ ​arteries​ ​(unusual​ ​in
insects),​ ​and​ ​paired​ ​segmental​ ​alary​ ​muscles.​ ​ ​ ​The​ ​ostia,​ ​segmental​ ​vessels,​ ​and
muscles​ ​will​ ​not​ ​be​ ​seen​ ​and​ ​most​ ​of​ ​the​ ​swellings​ ​are​ ​inconspicuous.​ ​ ​ ​The​ ​heart
opens​ ​anteriorly​ ​into​ ​the​ ​cephalic​ ​hemocoel​ ​of​ ​the​ ​head​ ​and​ ​posteriorly​ ​bifurcates​ ​to
form​ ​two​ ​arteries.
​ ​In​ ​arthropods​ ​blood​ ​is​ ​pumped​ ​anteriorly,​ ​during​ ​systole,​ ​by​ ​contractions​ ​of
circular​ ​muscles​ ​in​ ​the​ ​heart​ ​wall.​ ​Blood​ ​leaves​ ​the​ ​heart,​ ​enters​ ​the​ ​cephalic​ ​hemocoel
and​ ​passes​ ​posteriorly​ ​through​ ​the​ ​three​ ​sinuses​ ​of​ ​the​ ​hemocoel,​ ​eventually​ ​passing
through​ ​the​ ​perforations​ ​of​ ​the​ ​diaphragms​ ​to​ ​enter​ ​the​ ​pericardial​ ​sinus.​ ​ ​ ​During
diastole​ ​contractions​ ​of​ ​radiating​ ​alary​ ​muscles​ ​cause​ ​the​ ​heart​ ​to​ ​dilate​ ​and​ ​draw​ ​blood
into​ ​its​ ​lumen​ ​through​ ​the​ ​ostia.​ ​The​ ​heart​ ​is​ ​now​ ​refilled​ ​with​ ​blood​ ​and​ ​ready​ ​to​ ​enter
another​ ​systole.​ ​ ​ ​Heartbeat​ ​reversal​ ​has​ ​been​ ​observed​ ​in​ ​cockroaches.
Respiratory​ ​System
​ ​The​ ​respiratory​ ​system​ ​consists​ ​of​ ​10​ ​pairs​ ​of​ ​spiracles​ ​which​ ​open​ ​into​ ​a
complex​ ​system​ ​of​ ​distributory​ ​tracheae​ ​which​ ​deliver​ ​oxygen​ ​to​ ​the​ ​tissues.​ ​ ​ ​In​ ​insects
the​ ​blood​ ​is​ ​not​ ​involved​ ​in​ ​oxygen​ ​transport.
​ ​Removal​ ​of​ ​the​ ​tergites​ ​exposes​ ​the​ ​network​ ​of​ ​branching,​ ​tubular,​ ​white​ ​or
silvery​ ​tracheae​​ ​(Fig​ ​19).​ ​ ​ ​Use​ ​fine​ ​forceps​ ​to​ ​remove​ ​muscles​ ​and​ ​connective​ ​tissue
covering​ ​some​ ​of​ ​the​ ​tracheae​ ​and​ ​spiracles​ ​to​ ​improve​ ​your​ ​view​ ​of​ ​them.
​ ​The​ ​tracheae​ ​arise​ ​at​ ​spiracles​​ ​(Fig​ ​12,​ ​19)​ ​in​ ​the​ ​pleura​ ​near​ ​the​ ​posterolateral
corners​ ​of​ ​the​ ​tergites​ ​and​ ​join​ ​with​ ​other​ ​tracheae​ ​to​ ​form​ ​a​ ​single​ ​network​ ​of​ ​air​ ​tubes
for​ ​the​ ​delivery​ ​of​ ​oxygen​ ​to​ ​the​ ​tissues​ ​(Fig​ ​21-10B,​ ​C).​ ​Two​ ​pairs​ ​of​ ​spiracles​ ​are
present​ ​in​ ​the​ ​thorax​ ​and​ ​eight​ ​pairs​ ​in​ ​the​ ​abdomen.​ ​No​ ​spiracles​ ​are​ ​present​ ​in​ ​the
head.​ ​ ​ ​The​ ​spiracles​ ​were​ ​seen​ ​earlier​ ​in​ ​your​ ​study​ ​of​ ​the​ ​external​ ​anatomy.
​ ​On​ ​each​ ​side​ ​the​ ​respiratory​ ​system​ ​includes​ ​three​ ​longitudinal​ ​trunks​ ​and
segmental​ ​tracheae​ ​to​ ​the​ ​three​ ​hemocoel​ ​sinuses​ ​and​ ​their​ ​viscera.​ ​The​ ​portion​ ​of​ ​the

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system​ ​you​ ​see​ ​at​ ​present​ ​consists,​ ​on​ ​each​ ​side,​ ​of​ ​a​ ​lateral​ ​longitudinal​ ​trunk
extending​ ​from​ ​spiracle​ ​to​ ​spiracle​ ​(Fig​ ​19),​ ​a​ ​series​ ​of​ ​dorsal​ ​segmental​ ​tracheae
extending​ ​from​ ​the​ ​spiracles​ ​to​ ​the​ ​heart​ ​and​ ​pericardial​ ​sinus,​ ​and​ ​a​ ​dorsal
longitudinal​ ​trunk​​ ​connecting​ ​the​ ​dorsal​ ​segmental​ ​tracheae​ ​medially.​ ​The​ ​dorsal
longitudinal​ ​trunks​ ​lie​ ​beside​ ​the​ ​heart.​ ​ ​ ​Other​ ​segmental​ ​vessels,​ ​which​ ​cannot​ ​be
seen​ ​at​ ​present.
> ​ ​1a.​​ ​Remove​ ​a​ ​short​ ​piece​ ​of​ ​trachea​ ​and​ ​make​ ​a​ ​wetmount​ ​with​ ​it.​ ​ ​ ​Examine​ ​it
with​ ​the​ ​compound​ ​microscope​ ​at​ ​100X​ ​and​ ​400X​ ​with​ ​the​ ​light​ ​carefully​ ​adjusted.
Observe​ ​the​ ​closely​ ​spaced​ ​rings​ ​of​ ​chitin,​ ​similar​ ​in​ ​function​ ​to​ ​the​ ​cartilaginous​ ​rings​ ​in
a​ ​vertebrate​ ​trachea.​ ​These​ ​rings,​ ​known​ ​as​ ​taenidia​,​ ​hold​ ​the​ ​tracheae​ ​open,
permitting​ ​air​ ​to​ ​pass​ ​unimpeded​ ​(Fig​ ​21-10A).​ ​The​ ​taenidia​ ​is​ ​actually​ ​a​ ​single​ ​helix​ ​of
chitin,​ ​not​ ​a​ ​series​ ​of​ ​rings​ ​at​ ​all.​ ​ ​ ​The​ ​helix​ ​has​ ​a​ ​very​ ​short​ ​wavelength​ ​so​ ​its​ ​coils​ ​are
adjacent​ ​to​ ​each​ ​other,​ ​like​ ​a​ ​Slinky™​ ​at​ ​rest.​ ​<
​ ​As​ ​you​ ​remove​ ​tracheae​ ​from​ ​the​ ​thorax​ ​some​ ​taenidia​ ​may​ ​uncoil​ ​and​ ​extend
as​ ​a​ ​long,​ ​tough,​ ​very​ ​fine​ ​thread,​ ​thus​ ​demonstrating​ ​that​ ​it​ ​is​ ​a​ ​continuous​ ​coil,​ ​not​ ​a
series​ ​of​ ​separate​ ​rings.
Digestive​ ​System
" ​ ​Remove​ ​the​ ​dorsal​ ​diaphragm​ ​along​ ​with​ ​the​ ​tergal​ ​muscles,​ ​tracheae,​ ​and
heart​ ​from​ ​the​ ​abdomen.​ ​ ​ ​Accomplish​ ​this​ ​by​ ​cutting​ ​with​ ​fine​ ​scissors​ ​around​ ​the
periphery​ ​of​ ​the​ ​abdomen.​ ​ ​ ​Be​ ​very​ ​careful​ ​that​ ​you​ ​do​ ​not​ ​cut​ ​deeper​ ​than​ ​the
diaphragm.​ ​The​ ​crop​ ​of​ ​the​ ​digestive​ ​system​ ​is​ ​a​ ​large,​ ​thin​ ​walled​ ​chamber​ ​filling​ ​most
of​ ​the​ ​abdomen.​ ​ ​ ​Its​ ​walls​ ​may​ ​be​ ​closely​ ​appressed​ ​to​ ​the​ ​dorsal​ ​diaphragm,​ ​creating
the​ ​risk​ ​that​ ​it​ ​will​ ​be​ ​removed​ ​with​ ​the​ ​diaphragm.​ ​Remove​ ​the​ ​muscles,​ ​heart,​ ​and
tracheae,​ ​but​ ​nothing​ ​else,​ ​from​ ​the​ ​thorax.​ ​Be​ ​careful​ ​removing​ ​tissue​ ​in​ ​the​ ​thorax​ ​that
you​ ​do​ ​not​ ​inadvertently​ ​destroy​ ​the​ ​salivary​ ​glands​ ​and​ ​salivary​ ​reservoirs,​ ​both​ ​of
which​ ​are​ ​closely​ ​associated​ ​with​ ​the​ ​walls​ ​of​ ​the​ ​anterior​ ​gut​ ​(esophagus).
​ ​The​ ​space​ ​you​ ​have​ ​uncovered​ ​is​ ​the​ ​large,​ ​spacious​ ​perivisceral​ ​hemocoel​,
in​ ​which​ ​most​ ​of​ ​the​ ​viscera​ ​are​ ​found​ ​(Fig​ ​16-7).​ ​ ​ ​Note​ ​the​ ​large,​ ​amorphous,​ ​white​ ​fat
body​​ ​in​ ​the​ ​hemocoel​ ​(Fig​ ​22).​ ​This​ ​versatile​ ​tissue​ ​occupies​ ​much​ ​of​ ​the​ ​space​ ​in​ ​the
abdominal​ ​hemocoel​ ​but​ ​is​ ​also​ ​present​ ​in​ ​the​ ​thorax.​ ​It​ ​shares​ ​many​ ​functions​ ​with​ ​the
vertebrate​ ​liver​ ​and​ ​annelid​ ​chlorogogen.​ ​ ​ ​The​ ​fat​ ​body​ ​functions​ ​in​ ​storage​ ​of​ ​lipid,
glycogen​ ​and​ ​protein​ ​reserves.​ ​ ​ ​Stored​ ​food​ ​in​ ​the​ ​fat​ ​body​ ​supports​ ​survival​ ​over​ ​long
periods​ ​of​ ​starvation​ ​and​ ​its​ ​size​ ​varies​ ​depending​ ​on​ ​the​ ​extent​ ​of​ ​starvation.​ ​ ​ ​It​ ​is​ ​a
site​ ​for​ ​intermediary​ ​metabolism,​ ​amino​ ​acid​ ​synthesis,​ ​blood​ ​glucose​ ​regulation,
vitamin​ ​synthesis,​ ​and​ ​uric​ ​acid​ ​storage.​ ​ ​ ​Some​ ​of​ ​these​ ​functions​ ​are​ ​mediated​ ​by
mutualistic,​ ​intracellular​ ​bacteria​ ​known​ ​as​ ​bacterioids.
​ ​Note​ ​the​ ​two​ ​pairs​ ​of​ ​large​ ​diameter​ ​tracheae​ ​extending​ ​through​ ​the​ ​thorax​ ​into
the​ ​head.​ ​ ​ ​One​ ​pair​ ​lies​ ​beside​ ​the​ ​heart,​ ​the​ ​other​ ​is​ ​ventral,​ ​beside​ ​the​ ​esophagus.
They​ ​will​ ​be​ ​destroyed​ ​as​ ​you​ ​remove​ ​tissues​ ​to​ ​reveal​ ​the​ ​gut.
​ ​The​ ​cockroach​ ​gut,​ ​like​ ​that​ ​of​ ​other​ ​arthropods,​ ​consists​ ​of​ ​an​ ​anterior,
ectodermal​ ​stomodeum,​ ​or​ ​foregut,​ ​a​ ​middle​ ​endodermal​ ​midgut,​ ​and​ ​a​ ​posterior
endodermal​ ​proctodeum,​ ​or​ ​hindgut.​ ​Both​ ​stomodeum​ ​and​ ​proctodeum,​ ​being
ectodermal,​ ​are​ ​lined​ ​by​ ​epidermis​ ​and​ ​exoskeleton,​ ​which​ ​in​ ​some​ ​regions​ ​may​ ​be
sclerotized.​ ​The​ ​foregut​ ​has​ ​chief​ ​responsibility​ ​for​ ​mechanical​ ​digestion,​ ​trituration,

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chemical​ ​digestion,​ ​and​ ​storage,​ ​the​ ​midgut​ ​is​ ​the​ ​region​ ​of​ ​enzyme​ ​secretion,​ ​chemical
digestion​ ​and​ ​absorption.​ ​Water​ ​reclamation,​ ​feces​ ​formation​ ​and​ ​storage​ ​occur​ ​in​ ​the
hindgut.​ ​ ​ ​It​ ​is​ ​also​ ​a​ ​storage​ ​kidney​ ​that​ ​sequesters​ ​uric​ ​acid.
​ ​The​ ​foregut​ ​consists​ ​of​ ​mouth,​ ​pharynx,​ ​esophagus,​ ​crop,​ ​and​ ​proventriculus
(Fig​ ​21-8A,​ ​16-9).​ ​ ​ ​The​ ​midgut​ ​consists​ ​of​ ​midgut​ ​and​ ​digestive​ ​ceca.​ ​ ​ ​The​ ​hindgut​ ​is
composed​ ​of​ ​an​ ​ileum,​ ​colon,​ ​and​ ​rectum.​ ​ ​ ​The​ ​ileum​ ​and​ ​colon​ ​together​ ​are
sometimes​ ​known​ ​as​ ​the​ ​intestine.
​ ​The​ ​mouth​ ​was​ ​seen​ ​earlier​ ​opening​ ​from​ ​the​ ​roof​ ​of​ ​the​ ​preoral​ ​cavity​ ​(Fig
21-7).​ ​ ​ ​It​ ​opens​ ​onto​ ​the​ ​pharynx​ ​which​ ​extends​ ​dorsally​ ​through​ ​the​ ​head​ ​capsule.​ ​ ​ ​It
will​ ​not​ ​be​ ​seen​ ​from​ ​your​ ​present​ ​viewpoint.​ ​It​ ​becomes​ ​the​ ​esophagus​ ​and​ ​exits​ ​the
head​ ​capsule​ ​to​ ​enter​ ​the​ ​thorax,​ ​where​ ​it​ ​widens.​ ​ ​ ​The​ ​esophagus​​ ​is​ ​a​ ​conspicuous
feature​ ​of​ ​the​ ​anterior​ ​thorax​ ​(Fig​ ​20)​ ​where​ ​it​ ​can​ ​be​ ​recognized​ ​as​ ​a​ ​ridged,
thin-walled​ ​tube.
​ ​A​ ​pair​ ​of​ ​white,​ ​multi-lobed​ ​salivary​ ​glands​​ ​and​ ​a​ ​pair​ ​of​ ​bladder-like​ ​salivary
reservoirs​ ​adhere​ ​to​ ​the​ ​walls​ ​of​ ​the​ ​esophagus​ ​(Fig​ ​20).​ ​ ​ ​Saliva​ ​is​ ​secreted​ ​by​ ​the
glands​ ​and​ ​stored​ ​in​ ​the​ ​reservoirs.​ ​The​ ​salivary​ ​glands​ ​extend​ ​posteriorly​ ​from​ ​mid-
prothorax​ ​through​ ​the​ ​mesothorax.​ ​ ​ ​Salivary​ ​glands​ ​and​ ​reservoirs​ ​are​ ​drained​ ​by​ ​ducts
which​ ​extend​ ​anteriorly​ ​into​ ​the​ ​head,​ ​eventually​ ​to​ ​coalesce​ ​and​ ​empty​ ​into​ ​the
salivarium​ ​of​ ​the​ ​preoral​ ​cavity.​ ​ ​ ​The​ ​walls​ ​of​ ​the​ ​ducts​ ​are​ ​reinforced​ ​by​ ​helical​ ​rings
similar​ ​to​ ​the​ ​taenidia​ ​of​ ​tracheae.​ ​Here​ ​the​ ​saliva​ ​is​ ​mixed​ ​with​ ​food​ ​particles
masticated​ ​by​ ​the​ ​mandible​ ​and​ ​maxillary​ ​lacinia.​ ​ ​ ​The​ ​saliva​ ​contains​ ​amylase.
Figure​ ​20.​ ​Dorsal​ ​view​ ​of​ ​the​ ​perivisceral​ ​coelom​ ​of​ ​a​ ​male​ ​P.​ ​americana​.​ ​ ​ ​The
dorsal​ ​diaphragm,​ ​heart,​ ​and​ ​tergal​ ​muscles​ ​have​ ​been​ ​removed.​ ​ ​ ​Abdominal
segments​ ​are​ ​numbered.​ ​ ​ ​Malpighian​ ​tubules​ ​have​ ​been​ ​shortened​ ​and​ ​reduced
in​ ​number​ ​for​ ​clarity.​ ​Blatt56L.gif

​ ​Near​ ​the​ ​junction​ ​of​ ​the​ ​mesothorax​ ​and​ ​metathorax​ ​the​ ​esophagus​ ​widens​ ​to
become​ ​the​ ​crop​.​ ​ ​ ​This​ ​thin-walled,​ ​expansible​ ​storage​ ​organ​ ​may​ ​fill​ ​the​ ​perivisceral
hemocoel​ ​of​ ​the​ ​posterior​ ​thorax​ ​and​ ​anterior​ ​abdomen.​ ​Its​ ​thin,​ ​transparent​ ​walls​ ​may
push​ ​against​ ​the​ ​body​ ​wall​ ​and​ ​dorsal​ ​diaphragm.​ ​The​ ​crop​ ​of​ ​preserved​ ​specimens
may​ ​contain​ ​a​ ​large​ ​air​ ​bubble.​ ​ ​ ​Almost​ ​all​ ​hydrolysis​ ​occurs​ ​in​ ​the​ ​crop​ ​making​ ​use​ ​of
enzymes​ ​from​ ​two​ ​sources.​ ​ ​ ​Salivary​ ​amylase,​ ​is​ ​mixed​ ​with​ ​the​ ​food​ ​in​ ​the​ ​salivarium
and​ ​then​ ​ingested,​ ​whereas​ ​enzymes​ ​from​ ​the​ ​digestive​ ​ceca,​ ​move​ ​anteriorly​ ​from​ ​the

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midgut.
​ ​In​ ​the​ ​vicinity​ ​of​ ​segments​ ​5-6​ ​the​ ​crop​ ​narrows​ ​to​ ​become​ ​the​ ​short,​ ​muscular,
thick-walled​ ​proventriculus​​ ​(=​ ​gizzard).​ ​ ​ ​This​ ​region​ ​of​ ​the​ ​foregut​ ​has​ ​cuticularized
walls​ ​with​ ​heavily​ ​sclerotized,​ ​black​ ​or​ ​golden​ ​brown​ ​teeth​ ​(Fig​ ​21,​ ​21-8B).​ ​ ​ ​These​ ​may
be​ ​faintly​ ​visible​ ​through​ ​the​ ​walls​ ​of​ ​the​ ​crop​ ​but​ ​the​ ​walls​ ​of​ ​the​ ​proventriculus​ ​are
opaque​ ​and​ ​you​ ​will​ ​get​ ​a​ ​better​ ​look​ ​later​ ​when​ ​you​ ​open​ ​the​ ​gut.​ ​Esophagus,​ ​crop,
and​ ​proventriculus​ ​are​ ​regions​ ​of​ ​the​ ​foregut​.
​ ​The​ ​gut​ ​narrows​ ​abruptly​ ​posterior​ ​to​ ​the​ ​proventriculus​ ​and​ ​becomes​ ​the
midgut​​ ​(=​ ​ventriculus).​ ​ ​ ​The​ ​transition​ ​from​ ​foregut​ ​to​ ​midgut​ ​is​ ​marked​ ​internally​ ​by
the​ ​stomodeal​ ​valve,​ ​that​ ​protrudes​ ​into​ ​the​ ​anterior​ ​midgut​ ​(Fig​ ​21).
​ ​A​ ​whorl​ ​of​ ​eight​ ​long,​ ​fingerlike​ ​diverticula,​ ​the​ ​digestive​ ​ceca​,​ ​marks​ ​the
beginning​ ​of​ ​the​ ​midgut​ ​(Fig​ ​20,​ ​21-8A).​ ​ ​ ​The​ ​ceca​ ​increase​ ​the​ ​surface​ ​area​ ​for
secretion​ ​of​ ​enzymes​ ​and​ ​absorption​ ​of​ ​monomers.​ ​ ​ ​Enzymes​ ​produced​ ​in​ ​the​ ​ceca
include​ ​invertase,​ ​lipase,​ ​maltase,​ ​protease,​ ​(and​ ​perhaps​ ​cellulase​ ​in​ ​some​ ​species).
Lactase​ ​is​ ​also​ ​present​ ​in​ ​the​ ​midgut.​ ​Some​ ​of​ ​these​ ​may​ ​be​ ​produced​ ​by​ ​symbiotic​ ​gut
microorganisms​ ​and​ ​it​ ​may​ ​be​ ​that​ ​amylase​ ​is​ ​the​ ​only​ ​digestive​ ​enzyme​ ​produced​ ​by
the​ ​roach​ ​itself.
​ ​The​ ​midgut​ ​makes​ ​an​ ​asymmetric​ ​loop,​ ​curving​ ​to​ ​the​ ​right,​ ​then​ ​bending
sharply​ ​anteriorly,​ ​and​ ​then​ ​ventrally​ ​before​ ​heading​ ​posteriorly​ ​and​ ​dorsally.
Immediately​ ​after​ ​resuming​ ​its​ ​position​ ​on​ ​the​ ​midline​ ​it​ ​becomes​ ​the​ ​ileum​ ​of​ ​the
hindgut.
​ ​The​ ​transition​ ​from​ ​midgut​ ​to​ ​hindgut​ ​is​ ​marked​ ​by​ ​60-150​ ​long,​ ​slender,​ ​white,
threadlike,​ ​blind-ending​ ​Malpighian​ ​tubules​​ ​in​ ​six​ ​clusters​ ​(Fig​ ​21-8A,​ ​21-9).​ ​ ​ ​The
tubules​ ​are​ ​hollow​ ​gut​ ​diverticula​ ​that​ ​extend​ ​into​ ​the​ ​hemocoel​ ​where​ ​they​ ​are​ ​bathed
in​ ​blood.​ ​ ​ ​In​ ​Periplaneta​ ​they​ ​contain​ ​intracellular​ ​enzymes​ ​and​ ​may​ ​be​ ​more​ ​important
in​ ​digestion​ ​than​ ​in​ ​excretion.
​ ​The​ ​first​ ​region​ ​of​ ​the​ ​hindgut​​ ​is​ ​the​ ​short,​ ​narrow,​ ​muscular​ ​ileum​.​ ​It​ ​extends
for​ ​a​ ​short​ ​distance​ ​posteriorly​ ​from​ ​the​ ​midgut​ ​and​ ​then​ ​dilates​ ​to​ ​become​ ​the​ ​colon​.
The​ ​transition​ ​is​ ​marked​ ​internally​ ​by​ ​the​ ​ileocolic​ ​valve,​ ​a​ ​sphincter​ ​muscle.​ ​ ​ ​The​ ​colon
extends​ ​posteriorly​ ​to​ ​about​ ​segment​ ​7​ ​where​ ​it​ ​expands​ ​to​ ​become​ ​the​ ​rectum​.​ ​ ​ ​The
wall​ ​of​ ​the​ ​rectum​ ​bears​ ​six​ ​longitudinal,​ ​opaque,​ ​white​ ​ridges,​ ​the​ ​rectal​ ​pads​.​ ​ ​ ​The
pads​ ​protrude​ ​into​ ​the​ ​lumen​ ​of​ ​the​ ​rectum​ ​where​ ​they​ ​remove​ ​water​ ​from​ ​the​ ​forming
feces.​ ​ ​ ​The​ ​rectum​ ​efficiently​ ​reclaims​ ​water​ ​from​ ​feces​ ​when​ ​water​ ​is​ ​scarce​ ​and
conservation​ ​is​ ​advantageous,​ ​producing​ ​dry​ ​hard​ ​fecal​ ​pellets.​ ​ ​ ​When​ ​water​ ​is
abundant,​ ​however,​ ​the​ ​rectum​ ​does​ ​not​ ​remove​ ​water​ ​so​ ​effectively​ ​and​ ​produces​ ​wet
fecal​ ​pellets.​ ​ ​ ​Insect​ ​fecal​ ​pellets​ ​have​ ​characteristic​ ​patterns​ ​of​ ​longitudinal​ ​grooves
and​ ​ridges​ ​impressed​ ​on​ ​them​ ​by​ ​the​ ​rectal​ ​pads.
​ ​The​ ​rectum​ ​ends​ ​at​ ​the​ ​anus​,​ ​under​ ​tergite​ ​10​ ​between​ ​the​ ​two​ ​paraprocts.​ ​Slip
a​ ​needle​ ​under​ ​tergite​ ​10​ ​between​ ​the​ ​paraprocts​ ​and​ ​watch​ ​it​ ​appear​ ​in​ ​the​ ​rectum.
" ​ ​Use​ ​fine​ ​scissors​ ​to​ ​open​ ​the​ ​posterior​ ​crop​ ​and​ ​proventriculus.​ ​ ​ ​Inside​ ​the
proventriculus​ ​you​ ​will​ ​find​ ​an​ ​elaborate​ ​gastric​ ​mill​​ ​composed​ ​of​ ​an​ ​assortment​ ​of
sclerotized​ ​plates​ ​(Fig​ ​21,​ ​21-8A,B).​ ​ ​ ​Three​ ​sets​ ​of​ ​six​ ​plates​ ​each​ ​are​ ​embedded​ ​in​ ​the
proventricular​ ​wall.​ ​ ​ ​In​ ​a​ ​circle​ ​adjacent​ ​to​ ​the​ ​crop​ ​are​ ​12​ ​plates​ ​in​ ​two​ ​alternating​ ​sets.
Six​ ​are​ ​toothed​ ​plates​,​ ​each​ ​with​ ​a​ ​large​ ​complex​ ​tooth,​ ​and​ ​six​ ​are​ ​ridged​ ​plates​,

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each​ ​with​ ​several​ ​parallel​ ​ridges.​ ​The​ ​teeth​ ​continue​ ​the​ ​mechanical​ ​breakdown​ ​of​ ​food
particles​ ​initiated​ ​by​ ​the​ ​mandibles​ ​and​ ​maxillae.​ ​Six​ ​additional​ ​plates​ ​are​ ​arranged​ ​in​ ​a
second​ ​whorl,​ ​this​ ​one​ ​closer​ ​to​ ​the​ ​midgut.​ ​ ​ ​These​ ​lightly​ ​sclerotized​ ​plates​ ​support
soft​ ​cushions,​ ​the​ ​pulvilli​,​ ​bearing​ ​short​ ​fine​ ​setae​ ​which​ ​presumably​ ​act​ ​as​ ​a​ ​filter​ ​to
exclude​ ​large​ ​particles​ ​from​ ​the​ ​midgut.​ ​The​ ​astonishing​ ​gastric​ ​mill​ ​of​ ​the​ ​insect
proventriculus​ ​is​ ​a​ ​sight​ ​you​ ​don’t​ ​want​ ​to​ ​miss.
​ ​Continue​ ​the​ ​incision​ ​along​ ​the​ ​midgut​ ​and​ ​hindgut​ ​noting​ ​that​ ​these​ ​regions​ ​are
lined​ ​by​ ​a​ ​loose​ ​transparent​ ​peritrophic​ ​membrane​​ ​(Fig​ ​21-9).​ ​ ​ ​If​ ​food​ ​is​ ​present​ ​in​ ​the
gut​ ​of​ ​your​ ​specimen,​ ​it​ ​will​ ​be​ ​contained​ ​within​ ​the​ ​peritrophic​ ​membrane.​ ​ ​ ​The
membrane​ ​is​ ​secreted​ ​continuously​ ​by​ ​the​ ​gut​ ​epithelium.
Figure​ ​21.​ ​Interior​ ​of​ ​the​ ​proventriculus.​ ​Blatt57L.gif

​​

Excretory​ ​System
​ ​Cockroaches​ ​may​ ​employ​ ​several​ ​mechanisms​ ​for​ ​eliminating​ ​or​ ​inactivating
nitrogenous​ ​waste​ ​products,​ ​chiefly​ ​as​ ​uric​ ​acid.​ ​It​ ​is​ ​not​ ​known​ ​which​ ​is​ ​most​ ​important.
​ ​Some​ ​nitrogen​ ​may​ ​be​ ​transferred​ ​from​ ​the​ ​blood​ ​to​ ​the​ ​gut​ ​lumen,​ ​and​ ​thus​ ​to
the​ ​feces,​ ​by​ ​Malpighian​ ​tubules​ ​and​ ​the​ ​midgut.​ ​In​ ​most​ ​insects​ ​the​ ​Malpighian
tubules​​ ​remove​ ​wastes​ ​and​ ​inorganic​ ​ions​ ​from​ ​the​ ​blood​ ​and​ ​secrete​ ​them​ ​into​ ​the​ ​gut
lumen,​ ​where​ ​they​ ​become​ ​incorporated​ ​in​ ​the​ ​feces​ ​and​ ​are​ ​eliminated.​ ​Uric​ ​acid​ ​has

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not​ ​been​ ​demonstrated​ ​in​ ​cockroach​ ​Malpighian​ ​tubules,​ ​however,​ ​and​ ​it​ ​may​ ​be​ ​that,
since​ ​they​ ​are​ ​known​ ​to​ ​contain​ ​digestive​ ​enzymes,​ ​their​ ​role​ ​is​ ​in​ ​digestion​ ​rather​ ​than
excretion.
​ ​One​ ​of​ ​the​ ​functions​ ​of​ ​the​ ​fat​ ​body​​ ​is​ ​the​ ​sequestration,​ ​storage,​ ​and​ ​possibly
recycling​ ​of​ ​nitrogenous​ ​waste​ ​products.​ ​ ​ ​Specialized​ ​urate​ ​cells​ ​in​ ​the​ ​fat​ ​body
synthesize​ ​uric​ ​acid​ ​from​ ​waste​ ​nitrogen​ ​and​ ​store​ ​it​ ​indefinitely.​ ​ ​ ​Cockroaches​ ​on​ ​high
protein​ ​diets​ ​develop​ ​a​ ​hypertrophied​ ​fat​ ​body​ ​functioning​ ​as​ ​a​ ​storage​ ​kidney.​ ​ ​ ​It​ ​is
thought​ ​that​ ​endosymbiotic​ ​prokaryotic​ ​cells​ ​(bacterioids)​ ​use​ ​the​ ​stored​ ​nitrogen​ ​for
protein​ ​synthesis.​ ​Storage​ ​in​ ​the​ ​fat​ ​body​ ​is​ ​probably​ ​the​ ​most​ ​important​ ​excretory
mechanism​ ​in​ ​cockroaches.
​ ​In​ ​males​ ​of​ ​a​ ​few​ ​cockroach​ ​species​ ​nitrogen​ ​is​ ​excreted​ ​by​ ​the​ ​accessory
glands​ ​of​ ​the​ ​reproductive​ ​system.​ ​ ​ ​Some​ ​tubules​ ​of​ ​this​ ​gland​ ​absorb​ ​and​ ​store​ ​uric
acid​ ​which​ ​is​ ​then​ ​deposited​ ​in​ ​the​ ​spermatophore​ ​and​ ​eliminated​ ​during​ ​copulation.
This​ ​mechanism,​ ​of​ ​course,​ ​is​ ​of​ ​no​ ​use​ ​to​ ​females​ ​and​ ​is​ ​limited​ ​to​ ​males​ ​of​ ​only​ ​a​ ​few
species​ ​(such​ ​as​ ​Blatella​ ​germanica​ ​but​ ​not​ ​Periplaneta​).
​ ​The​ ​periodic​ ​molting​ ​by​ ​arthropods​ ​is​ ​potentially​ ​a​ ​mechanism​ ​for​ ​eliminating
unwanted​ ​materials​ ​from​ ​the​ ​body.​ ​It​ ​has​ ​been​ ​suggested​ ​that​ ​the​ ​nitrogen,​ ​as​ ​protein,
incorporated​ ​in​ ​the​ ​developing​ ​cuticle​ ​during​ ​can​ ​serve​ ​as​ ​a​ ​mechanism​ ​for​ ​eliminating
excess​ ​nitrogen.​ ​ ​ ​The​ ​incorporated​ ​nitrogen​ ​is​ ​then​ ​jettisoned​ ​with​ ​the​ ​next​ ​molt.​ ​ ​ ​This
mechanism​ ​could​ ​be​ ​effective​ ​in​ ​nymphs​ ​but​ ​not​ ​adults,​ ​since​ ​they​ ​do​ ​not​ ​molt.

Reproductive​ ​System
Female
​ ​The​ ​female​ ​internal​ ​genitalia​ ​consist​ ​chiefly​ ​of​ ​a​ ​pair​ ​of​ ​ovaries,​ ​two​ ​colleterial
glands​ ​(=​ ​female​ ​accessory​ ​glands)​ ​with​ ​separate​ ​ducts,​ ​seminal​ ​receptacle​ ​(two​ ​in
some​ ​species),​ ​and​ ​oviducts​ ​(Fig​ ​21-11A).​ ​Each​ ​ovary​ ​is​ ​a​ ​bundle​ ​of​ ​tapering​ ​tubes,​ ​the
ovarioles,​ ​which​ ​empty​ ​into​ ​an​ ​oviduct.​ ​ ​ ​Oogenesis​ ​and​ ​gamete​ ​maturation​ ​occur​ ​in​ ​the
ovarioles.​ ​ ​ ​The​ ​lateral​ ​oviducts,​ ​one​ ​from​ ​each​ ​of​ ​the​ ​two​ ​ovaries,​ ​join​ ​to​ ​form​ ​a
common​ ​oviduct​ ​that​ ​opens​ ​through​ ​sternite​ ​8​ ​into​ ​the​ ​genital​ ​pouch​ ​(Fig​ ​15).​ ​ ​ ​The​ ​duct
from​ ​the​ ​seminal​ ​receptacle​ ​opens​ ​at​ ​the​ ​end​ ​of​ ​the​ ​receptacle​ ​papilla​ ​in​ ​the​ ​pouch​ ​(Fig
23).​ ​ ​ ​During​ ​copulation​ ​the​ ​male​ ​attaches​ ​a​ ​spermatophore​ ​a​ ​sternite​ ​in​ ​the​ ​female
genital​ ​pouch.​ ​ ​ ​Sperm​ ​exit​ ​the​ ​spermatophore,​ ​travel​ ​up​ ​the​ ​receptacle​ ​duct​ ​to​ ​the
seminal​ ​receptacle​ ​where​ ​they​ ​are​ ​stored,​ ​potentially​ ​for​ ​a​ ​year​ ​or​ ​more.
" ​ ​The​ ​female​ ​reproductive​ ​system​ ​will​ ​not​ ​be​ ​visible​ ​until​ ​much​ ​of​ ​the​ ​fat​ ​body​​ ​in
the​ ​dorsal​ ​abdomen​ ​has​ ​been​ ​removed.​ ​ ​ ​Begin​ ​first​ ​in​ ​the​ ​posterior​ ​abdomen​ ​and
remove​ ​tergites​ ​7-9​ ​if​ ​you​ ​have​ ​not​ ​already​ ​done​ ​so.​ ​The​ ​fat​ ​body​ ​covers​ ​and​ ​is​ ​packed
around​ ​the​ ​colleterial​ ​glands​ ​and​ ​ovaries​ ​and​ ​must​ ​be​ ​removed​ ​carefully​ ​without
damaging​ ​either.​ ​ ​ ​The​ ​size​ ​of​ ​the​ ​fat​ ​body​ ​varies​ ​depending​ ​on​ ​metabolic​ ​and
reproductive​ ​condition​ ​of​ ​the​ ​insect.​ ​ ​ ​Reserves​ ​stored​ ​in​ ​the​ ​fat​ ​body​ ​are​ ​transferred​ ​to
the​ ​developing​ ​eggs​ ​when​ ​the​ ​female​ ​is​ ​reproductive​ ​active.
​ ​Once​ ​you​ ​have​ ​removed​ ​the​ ​fat​ ​body,​ ​the​ ​left​ ​colleterial​ ​gland​​ ​will​ ​be
conspicuous​ ​as​ ​spaghetti-like​ ​tangle​ ​of​ ​many​ ​long,​ ​slender,​ ​branching,​ ​opaque​ ​white

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tubes.​ ​ ​ ​The​ ​glands​ ​lie​ ​beside​ ​and​ ​dorsal​ ​to​ ​the​ ​hindgut​ ​and,​ ​along​ ​with​ ​the​ ​fat​ ​body,
obscure​ ​your​ ​view​ ​of​ ​everything​ ​else​ ​in​ ​the​ ​posterior​ ​abdomen​ ​(Fig​ ​22).​ ​ ​ ​The​ ​colleterial
gland​ ​is​ ​not​ ​restricted​ ​to​ ​the​ ​surface​ ​and​ ​fills​ ​much​ ​of​ ​the​ ​space​ ​between​ ​the​ ​sternum
and​ ​tergum.​ ​The​ ​smaller​ ​right​ ​colleterial​ ​gland​ ​will​ ​probably​ ​not​ ​be​ ​seen.
​ ​The​ ​two​ ​colleterial​ ​glands​ ​empty​ ​via​ ​separate​ ​ducts​ ​into​ ​the​ ​genital​ ​pouch.​ ​ ​ ​The
two​ ​glands​ ​differ​ ​dramatically​ ​both​ ​morphologically​ ​and​ ​chemically.​ ​The​ ​left​ ​gland​ ​is
much​ ​larger​ ​than​ ​the​ ​right​ ​and​ ​completely​ ​obscures​ ​it​ ​from​ ​view.​ ​Secretions​ ​from​ ​both
are​ ​released​ ​simultaneously​ ​into​ ​the​ ​genital​ ​pouch​ ​and​ ​react​ ​to​ ​form​ ​the​ ​sclerotized
lining​ ​of​ ​the​ ​pouch​ ​that​ ​will​ ​become​ ​the​ ​ootheca.​ ​ ​ ​Protein​ ​released​ ​from​ ​the​ ​left​ ​gland​ ​is
tanned​ ​by​ ​phenol​ ​from​ ​the​ ​right​ ​gland.​ ​The​ ​resulting​ ​wall​ ​of​ ​the​ ​ootheca​ ​wall​ ​is
composed​ ​of​ ​scleroprotein​ ​identical​ ​to​ ​that​ ​in​ ​cuticular​ ​sclerites,​ ​although​ ​chitin​ ​is
absent.​ ​Eggs​ ​are​ ​deposited​ ​into​ ​this​ ​lining​ ​as​ ​they​ ​leave​ ​the​ ​ovarioles​ ​and​ ​the​ ​lining
then​ ​hardens​ ​(tans)​ ​to​ ​become​ ​the​ ​ootheca.
​ ​The​ ​ovaries​​ ​lie​ ​laterally​ ​in​ ​the​ ​abdomen,​ ​one​ ​on​ ​either​ ​side​ ​of​ ​the​ ​gut​ ​and​ ​may
extend​ ​for​ ​most​ ​of​ ​the​ ​length​ ​of​ ​the​ ​abdomen.​ ​Or,​ ​if​ ​immature​ ​or​ ​inactive,​ ​may​ ​be
restricted​ ​to​ ​abdominal​ ​segments​ ​5-7.​ ​The​ ​ovaries​ ​are​ ​hidden​ ​from​ ​view​ ​by​ ​the​ ​fat​ ​body
and​ ​colleterial​ ​glands,​ ​which​ ​much​ ​be​ ​removed.
​ ​Each​ ​ovary​ ​is​ ​a​ ​bundle​ ​of​ ​large​ ​tapering​ ​tubes,​ ​the​ ​ovarioles​.​ ​In​ ​Periplaneta
each​ ​ovary​ ​consists​ ​of​ ​eight​ ​ovarioles​ ​(Fig​ ​22).​ ​ ​ ​Each​ ​ovariole​ ​is​ ​a​ ​connective​ ​tissue
tube​ ​consisting​ ​of​ ​two​ ​regions.​ ​The​ ​germarium​​ ​is​ ​the​ ​short,​ ​small-diameter,​ ​upstream,
anterior​ ​region.​ ​Oogonia​ ​in​ ​the​ ​germarium​ ​initiate​ ​oogenesis​ ​culminating​ ​downstream​ ​in
the​ ​production​ ​of​ ​oocytes.​ ​The​ ​vitellarium​​ ​is​ ​the​ ​much​ ​longer,​ ​much​ ​wider,​ ​downstream
region​ ​in​ ​which​ ​follicle​ ​cells​ ​transfer​ ​nutriment​ ​to​ ​the​ ​maturing​ ​and​ ​growing​ ​oocytes.​ ​The
diameter​ ​of​ ​the​ ​ovariole​ ​is​ ​very​ ​small​ ​in​ ​the​ ​germarium​ ​but​ ​increases​ ​downstream​ ​as​ ​the
developing​ ​gametes​ ​accumulate​ ​yolk​ ​from​ ​the​ ​follicle​ ​cells​ ​and​ ​increase​ ​dramatically​ ​in
size​ ​(Fig​ ​22).
Figure​ ​22.​ ​Dorsal​ ​view​ ​of​ ​the​ ​opened​ ​abdomen​ ​of​ ​a​ ​female​ ​P.​ ​fuliginosa​.​ ​The​ ​fat
body​ ​and​ ​colleterial​ ​gland​ ​have​ ​been​ ​removed​ ​from​ ​the​ ​left​ ​side​ ​to​ ​reveal​ ​the​ ​left
ovary.​ ​T​ ​=​ ​tergite.​ ​Blatt58L.gif

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 ​ ​The​ ​downstream​ ​ends​ ​of​ ​the​ ​eight​ ​ovarioles​ ​of​ ​each​ ​ovary​ ​coalesce​ ​to​ ​form​ ​a
lateral​ ​oviduct​ ​(Fig​ ​21-11A).​ ​ ​ ​Remove​ ​the​ ​colleterial​ ​glands​ ​from​ ​one​ ​side​ ​and​ ​find​ ​the
lateral​ ​oviduct​​ ​exiting​ ​the​ ​large,​ ​posterior,​ ​basal​ ​end​ ​of​ ​the​ ​ovary.​ ​ ​ ​Upon​ ​exiting​ ​the
ovary​ ​the​ ​oviduct​ ​passes​ ​deep​ ​into​ ​the​ ​tissues​ ​in​ ​the​ ​ventral​ ​abdomen​ ​and​ ​extends
medially​ ​to​ ​join​ ​the​ ​other​ ​oviduct​ ​on​ ​the​ ​ventral​ ​midline.​ ​ ​ ​The​ ​junction​ ​of​ ​the​ ​right​ ​and
left​ ​lateral​ ​oviducts​ ​forms​ ​the​ ​short​ ​unpaired​ ​common​ ​oviduct​​ ​which​ ​passes​ ​posteriorly
to​ ​open​ ​through​ ​the​ ​female​ ​gonopore​ ​in​ ​the​ ​genital​ ​pouch​ ​(Fig​ ​15,​ ​23).
​ ​Maturing​ ​gametes​ ​are​ ​lined​ ​up​ ​one​ ​after​ ​another​ ​in​ ​the​ ​ovariole​ ​with​ ​oocytes
maturing​ ​as​ ​they​ ​move​ ​downstream​ ​(Fig​ ​22).​ ​ ​ ​The​ ​mature​ ​egg​ ​(actually,​ ​it​ ​is​ ​a
secondary​ ​oocyte​ ​that​ ​will​ ​not​ ​complete​ ​meiosis​ ​until​ ​it​ ​is​ ​fertilized)​ ​is​ ​the
downstream-most​ ​gamete​ ​in​ ​the​ ​ovariole.​ ​ ​ ​ ​Each​ ​ovariole​ ​typically​ ​has​ ​a​ ​single​ ​mature
egg​ ​ready​ ​for​ ​fertilization​ ​and​ ​incorporation​ ​into​ ​an​ ​ootheca​ ​(Fig​ ​22).​ ​ ​ ​Eggs​ ​leave​ ​the
ovarioles​ ​via​ ​the​ ​oviduct​ ​and​ ​are​ ​fertilized​ ​by​ ​sperm​ ​from​ ​the​ ​seminal​ ​receptacle​ ​as​ ​they
enter​ ​the​ ​genital​ ​pouch.​ ​ ​ ​The​ ​ovipositor​ ​manipulates​ ​them​ ​and​ ​orients​ ​them​ ​properly​ ​in
the​ ​developing​ ​ootheca.​ ​If​ ​the​ ​eggs​ ​are​ ​mis-oriented,​ ​the​ ​nymphs​ ​will​ ​be​ ​unable​ ​to
escape​ ​from​ ​the​ ​ootheca​ ​and​ ​will​ ​die​ ​.​ ​The​ ​ootheca​ ​hardens​ ​around​ ​two​ ​rows​ ​of​ ​eight
eggs​ ​each,​ ​one​ ​from​ ​each​ ​of​ ​the​ ​16​ ​ovarioles​ ​(eight​ ​ovarioles​ ​in​ ​each​ ​of​ ​two​ ​ovaries).
The​ ​number​ ​of​ ​eggs​ ​in​ ​the​ ​ootheca​ ​depends​ ​on​ ​the​ ​number​ ​of​ ​ovarioles​ ​in​ ​the​ ​ovary
and​ ​differs​ ​with​ ​species.
​ ​Females​ ​of​ ​some​ ​species​ ​(e.g.​ ​Blatella​ ​germanica​)​ ​retain​ ​the​ ​ootheca​ ​and​ ​brood
their​ ​eggs.​ ​ ​ ​Most,​ ​including​ ​Periplaneta​,​ ​drop​ ​the​ ​completed​ ​ootheca​ ​and​ ​abandon​ ​it.

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Figure​ ​23.​ ​Dorsal​ ​view​ ​of​ ​the​ ​floor​ ​of​ ​the​ ​genital​ ​pouch​ ​with​ ​the​ ​gut,​ ​fat​ ​body,​ ​and
colleterial​ ​glands​ ​removed.​ ​Blatt59L.gif

Male
​ ​The​ ​male​ ​reproductive​ ​system​ ​consists​ ​of​ ​paired​ ​testes,​ ​each​ ​draining​ ​by​ ​a
sperm​ ​duct​ ​(=​ ​vas​ ​deferens)​ ​to​ ​a​ ​common​ ​ejaculatory​ ​duct​ ​which​ ​opens​ ​via​ ​the​ ​male
gonopore​ ​in​ ​the​ ​genital​ ​pouch​ ​(Fig​ ​21-12A).​ ​ ​ ​Associated​ ​with​ ​the​ ​confluence​ ​of​ ​the​ ​two
sperm​ ​ducts​ ​are​ ​two​ ​clusters​ ​of​ ​secretory​ ​accessory​ ​glands.​ ​ ​ ​Near​ ​the​ ​confluence,​ ​the
sperm​ ​ducts​ ​are​ ​expanded​ ​to​ ​form​ ​seminal​ ​vesicles.​ ​It​ ​is​ ​thought​ ​that​ ​spermatogenesis
occurs​ ​in​ ​the​ ​testes​ ​during​ ​the​ ​last​ ​nymphal​ ​instar​ ​and​ ​the​ ​spermatozoa​ ​stored​ ​for​ ​later
use​ ​as​ ​an​ ​imago.
​ ​The​ ​testes​ ​are​ ​located​ ​dorsolaterally​ ​in​ ​segments​ ​4​ ​and​ ​5​ ​(Fig​ ​20).​ ​ ​ ​They​ ​are
embedded​ ​in​ ​the​ ​white​ ​fat​ ​body​ ​and​ ​can​ ​be​ ​difficult​ ​or​ ​impossible​ ​to​ ​demonstrate​ ​in
adults,​ ​especially​ ​older​ ​specimens.​ ​Part​ ​of​ ​this​ ​difficulty​ ​is​ ​attributed​ ​to​ ​the​ ​alleged
(disputed)​ ​degeneration​ ​of​ ​the​ ​testes​ ​in​ ​older​ ​adults​ ​but​ ​most​ ​of​ ​the​ ​problem​ ​is​ ​due​ ​to
the​ ​fat​ ​bodies.​ ​Like​ ​the​ ​testes,​ ​the​ ​sperm​ ​ducts​ ​are​ ​difficult​ ​to​ ​demonstrate​ ​and​ ​you
probably​ ​will​ ​not​ ​see​ ​them.
​ ​Prior​ ​to​ ​copulation​ ​a​ ​spermatophore​ ​is​ ​formed​ ​in​ ​the​ ​male​ ​ejaculatory​ ​duct​ ​when
secretions​ ​of​ ​the​ ​accessory​ ​glands​ ​enclose​ ​and​ ​harden​ ​around​ ​a​ ​mass​ ​of​ ​sperm​ ​from
the​ ​seminal​ ​vesicles.​ ​During​ ​copulation​ ​the​ ​phallomeres​ ​open​ ​the​ ​female’s​ ​genital
pouch​ ​and​ ​hold​ ​the​ ​male’s​ ​genitalia​ ​in​ ​the​ ​correct​ ​position.​ ​ ​ ​The​ ​ejaculatory​ ​duct​ ​on​ ​the
ventral​ ​phallomere​ ​is​ ​held​ ​adjacent​ ​to​ ​the​ ​opening​ ​of​ ​the​ ​spermathecal​ ​papilla​ ​(Fig​ ​23)
and​ ​the​ ​spermatophore​ ​released.​ ​It​ ​is​ ​believed​ ​that​ ​the​ ​spermatophore​ ​is​ ​glued​ ​in​ ​place

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by​ ​secretions​ ​of​ ​the​ ​male’s​ ​phallic​ ​gland.​ ​In​ ​about​ ​24​ ​hours​ ​or​ ​less,​ ​sperm​ ​exit​ ​the
spermatophore​ ​and​ ​are​ ​stored​ ​in​ ​the​ ​female’s​ ​seminal​ ​receptacle.
Figure​ ​24.​ ​Dorsal​ ​dissection​ ​of​ ​the​ ​posterior​ ​hemocoel​ ​and​ ​genital​ ​pouch​ ​of​ ​a
male​ ​P.​ ​americana.​ ​The​ ​internal​ ​genitalia​ ​(accessory​ ​glands)​ ​have​ ​been​ ​removed.
Blatt60L.gif

" ​ ​In​ ​dorsal​ ​dissection​ ​your​ ​view​ ​of​ ​the​ ​male​ ​reproductive​ ​system​ ​is​ ​probably
obscured​ ​by​ ​lobes​ ​of​ ​the​ ​bright​ ​white​ ​fat​ ​body.​ ​ ​ ​Remove​ ​this​ ​region​ ​of​ ​the​ ​fat​ ​body​ ​and
tergite​ ​8-9​ ​if​ ​you​ ​have​ ​not​ ​already​ ​done​ ​so.
​ ​Following​ ​removal​ ​of​ ​the​ ​fat​ ​body,​ ​the​ ​male​​ ​accessory​ ​gland​​ ​(=​ ​mushroom
gland)​ ​is​ ​by​ ​far​ ​the​ ​most​ ​conspicuous​ ​of​ ​the​ ​male's​ ​internal​ ​genitalia​ ​(Fig​ ​20).​ ​ ​ ​It
secretes​ ​the​ ​covering​ ​of​ ​the​ ​spermatophore​ ​and​ ​is​ ​a​ ​mass​ ​of​ ​worm-like​ ​diverticula​ ​in​ ​the
posterior​ ​hemocoel​ ​under​ ​tergites​ ​7-8​ ​beside​ ​the​ ​anterior​ ​end​ ​of​ ​the​ ​posterior​ ​colon​ ​and
anterior​ ​rectum.​ ​ ​ ​The​ ​mass​ ​includes​ ​large​ ​(utriculi​ ​majores)​ ​and​ ​small​ ​(utriculi
breviores)​ ​diverticula​ ​and​ ​is​ ​usually​ ​surrounded​ ​by​ ​lobes​ ​of​ ​the​ ​fat​ ​body.
​ ​The​ ​two​ ​sperm​ ​ducts​ ​enter​ ​the​ ​gland​ ​and​ ​the​ ​seminal​ ​vesicles​ ​are​ ​enclosed​ ​in
the​ ​mass​ ​of​ ​tubules.​ ​The​ ​thick​ ​ejaculatory​ ​duct​,​ ​formed​ ​by​ ​the​ ​union​ ​of​ ​the​ ​sperm
ducts,​ ​exits​ ​ventrally​ ​from​ ​the​ ​center​ ​mass​ ​of​ ​accessory​ ​glands​ ​and​ ​extends​ ​deep​ ​into
the​ ​ventral​ ​body​ ​wall​ ​and​ ​then​ ​posteriorly​ ​to​ ​the​ ​gonopore​ ​on​ ​the​ ​dorsal​ ​surface​ ​of​ ​the

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ventral​ ​phallomere​ ​(Fig​ ​24).
​ ​Most​ ​of​ ​the​ ​phallic​ ​gland​​ ​is​ ​ventral​ ​to​ ​the​ ​nerve​ ​cord​ ​and​ ​should​ ​not​ ​be​ ​exposed
until​ ​you​ ​have​ ​completed​ ​your​ ​study​ ​of​ ​the​ ​nervous​ ​system​ ​(Fig​ ​24).​ ​ ​ ​It​ ​is​ ​deeply​ ​ventral
in​ ​the​ ​posterior​ ​abdominal​ ​cavity,​ ​ventral​ ​even​ ​to​ ​the​ ​ventral​ ​diaphragm​ ​and​ ​is​ ​a​ ​little​ ​to
the​ ​right​ ​of​ ​the​ ​midline.​ ​ ​ ​Remove​ ​the​ ​tissue​ ​ventral​ ​to​ ​the​ ​posterior​ ​nerve​ ​cord​ ​to
expose​ ​the​ ​gland.​ ​ ​ ​Its​ ​appearance​ ​is​ ​distinctive.​ ​ ​ ​It​ ​is​ ​glandular,​ ​white,​ ​and​ ​shaped​ ​like
an​ ​elongate​ ​teardrop,​ ​tapering​ ​posteriorly​ ​to​ ​form​ ​the​ ​phallic​ ​gland​ ​duct​.​ ​It​ ​is​ ​between
abdominal​ ​ganglion​ ​5​ ​and​ ​the​ ​terminal​ ​ganglion.​ ​ ​ ​It​ ​is​ ​ventral​ ​to​ ​the​ ​terminal​ ​ganglion
and​ ​the​ ​right​ ​cercal​ ​nerve.​ ​Its​ ​duct​ ​opens​ ​in​ ​the​ ​genital​ ​pouch​ ​between​ ​the​ ​pseudopenis
and​ ​acutolobus​ ​of​ ​the​ ​left​ ​phallomere​ ​(Fig​ ​18,​ ​24).​ ​ ​ ​During​ ​copulation​ ​secretions​ ​of​ ​the
phallic​ ​gland​ ​are​ ​used​ ​to​ ​attach​ ​the​ ​spermatophore​ ​in​ ​the​ ​vicinity​ ​of​ ​the​ ​opening​ ​of​ ​the
seminal​ ​receptacle.
Nervous​ ​System
​ ​The​ ​basic​ ​features​ ​of​ ​the​ ​insect​ ​nervous​ ​system​ ​are​ ​the​ ​dorsal​ ​brain​ ​joined​ ​by
circumenteric​ ​connectives​ ​to​ ​ventral​ ​nerve​ ​cord​ ​with​ ​paired​ ​segmental​ ​ganglia.​ ​It
includes​ ​both​ ​somatic​ ​and​ ​visceral​ ​components​ ​and​ ​is​ ​served​ ​by​ ​a​ ​variety​ ​of​ ​sensory
devices.​ ​ ​ ​Most,​ ​but​ ​not​ ​all,​ ​segments​ ​have​ ​a​ ​pair​ ​of​ ​segmental​ ​ganglia.​ ​Segmental
ganglia​ ​and​ ​the​ ​brain​ ​consist​ ​of​ ​paired​ ​lateral​ ​ganglia​ ​connected​ ​by​ ​a​ ​transverse
commissure.​ ​ ​ ​In​ ​cockroaches​ ​the​ ​commissures​ ​are​ ​short​ ​resulting​ ​in​ ​fusion​ ​of​ ​the
lateral​ ​ganglia​ ​so​ ​each​ ​segmental​ ​ganglion​ ​seems​ ​to​ ​be​ ​a​ ​single​ ​ganglion.
" ​ ​Remove​ ​the​ ​gut,​ ​ventral​ ​diaphragm,​ ​and​ ​muscles​ ​from​ ​the​ ​thorax​ ​and​ ​abdomen.
Remove​ ​the​ ​longitudinal​ ​body​ ​wall​ ​muscles​ ​(sternal​ ​muscles)​ ​and​ ​connective​ ​tissue​ ​as
necessary​ ​from​ ​the​ ​floor​ ​of​ ​the​ ​abdominal​ ​cavity​ ​to​ ​reveal​ ​the​ ​ventral​ ​nerve​ ​cord​​ ​(Fig
16-11).​ ​The​ ​cord​ ​consists​ ​to​ ​two​ ​side-by​ ​side,​ ​parallel,​ ​longitudinal​ ​connectives​ ​which
are​ ​united​ ​by​ ​transverse​ ​commissures​ ​passing​ ​between​ ​paired​ ​segmental​ ​ganglia​.
Notice​ ​the​ ​longitudinal​ ​ventral​ ​tracheal​ ​trunk​​ ​lying​ ​beside​ ​the​ ​ventral​ ​nerve​ ​cord.
​ ​Three​ ​thoracic​ ​ganglia​​ ​are​ ​located​ ​in​ ​the​ ​thorax​ ​(Fig​ ​25).​ ​ ​ ​From​ ​each​ ​extends
several​ ​pairs​ ​of​ ​nerves​ ​to​ ​the​ ​abundant​ ​muscles​ ​of​ ​these​ ​segments.​ ​ ​ ​These​ ​are​ ​large
ganglia.
Figure​ ​25.​ ​Dorsal​ ​dissection​ ​of​ ​P.​ ​americana​ ​T​ ​=​ ​thoracic​ ​ganglion,​ ​A​ ​=​ ​abdominal
ganglion.​ ​ ​ ​The​ ​abdominal​ ​segments​ ​are​ ​numbered.​ ​ ​ ​Blatt61L.gif

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 ​ ​In​ ​the​ ​abdomen​ ​are​ ​six​ ​much​ ​smaller​ ​abdominal​ ​ganglia​.​ ​Abdominal​ ​ganglia
1-3​ ​are​ ​scarcely​ ​wider​ ​than​ ​the​ ​double​ ​nerve​ ​cord​ ​and​ ​can​ ​be​ ​difficult​ ​to​ ​distinguish​ ​from
the​ ​cord.​ ​ ​ ​Abdominal​ ​ganglia​ ​4​ ​and​ ​5​ ​are​ ​usually​ ​readily​ ​discernable.​ ​ ​ ​The​ ​sixth
abdominal​ ​ganglion,​ ​known​ ​as​ ​the​ ​terminal​ ​ganglion​,​ ​is​ ​the​ ​largest​ ​of​ ​the​ ​abdominal
ganglia​ ​and​ ​is​ ​easily​ ​recognized.​ ​The​ ​nerve​ ​cord​ ​ends​ ​at​ ​the​ ​terminal​ ​ganglion​ ​which
gives​ ​rise​ ​to​ ​two​ ​large​ ​sensory​ ​cercal​ ​nerves​​ ​to​ ​the​ ​cerci.​ ​ ​ ​The​ ​terminal​ ​ganglion​ ​is​ ​the
coalesced​ ​ganglia​ ​of​ ​several​ ​posterior​ ​segmental​ ​ganglia.
" ​ ​Using​ ​your​ ​finest​ ​scissors​ ​and​ ​forceps​ ​carefully​ ​remove​ ​the​ ​epicranium​ ​from​ ​the
region​ ​between​ ​the​ ​compound​ ​eyes​ ​and​ ​antennae.​ ​ ​ ​Remove​ ​muscles​ ​as​ ​necessary​ ​to
reveal​ ​the​ ​bright​ ​white,​ ​dorsal​ ​brain​​ ​between​ ​the​ ​compound​ ​eyes​ ​(Fig​ ​25).​ ​The​ ​brain
consists​ ​of​ ​two​ ​cerebral​ ​ganglia​​ ​fused​ ​across​ ​the​ ​midline.​ ​Short​ ​thick​ ​optic​​ ​nerves
extend​ ​from​ ​the​ ​eyes​ ​to​ ​the​ ​brain.​ ​ ​ ​Smaller​ ​antennal​ ​nerves​​ ​run​ ​from​ ​the​ ​antennae​ ​to
the​ ​brain.​ ​ ​ ​Remove​ ​the​ ​mandibles​ ​and​ ​other​ ​tissues​ ​from​ ​one​ ​side​ ​of​ ​the​ ​head​ ​to​ ​reveal
the​ ​thick​ ​circumesophageal​ ​connectives.​​ ​These​ ​exit​ ​the​ ​ventral​ ​side​ ​of​ ​the​ ​brain​ ​and
extend​ ​posteriorly​ ​to​ ​pass​ ​around​ ​the​ ​esophagus,​ ​and​ ​then​ ​coalesce​ ​to​ ​form​ ​the
subesophageal​ ​ganglion​​ ​ventral​ ​to​ ​the​ ​gut.​ ​ ​ ​Nerves​ ​to​ ​the​ ​mandibles,​ ​maxillae,​ ​and
labium​ ​exit​ ​the​ ​subesophageal​ ​ganglion.​ ​A​ ​pair​ ​of​ ​longitudinal​ ​nerve​ ​cords​ ​exit​ ​the
subesophageal​ ​ganglion​ ​as​ ​the​ ​ventral​ ​nerve​ ​cord​ ​and​ ​pass​ ​posteriorly​ ​through​ ​the
foramen​ ​magnum​ ​to​ ​thoracic​ ​ganglion​ ​1​ ​(Fig​ ​25).

References
Anon.​​ ​1967.​ ​ ​ ​Pictorial​ ​keys,​ ​arthropods,​ ​reptiles,​ ​birds,​ ​and​ ​mammals​ ​of​ ​public​ ​health
significance.​ ​ ​ ​US​ ​Public​ ​Health​ ​Service,​ ​Communicable​ ​Disease​ ​Center,​ ​Atlanta.​ ​192​ ​pp.
Barbara​ ​KA​​ ​.​ ​University​ ​of​ ​Florida​ ​Institute​ ​of​ ​Food​ ​and​ ​Agricultural​ ​Sciences,​ ​Periplaneta
americana​ ​page:​ ​http://creatures.ifas.ufl.edu/urban/roaches/american_cockroach.htm
Bell​ ​WJ,​ ​Adiyodi​ ​KG.​​ ​1981.​ ​ ​ ​The​ ​American​ ​cockroach.​ ​Chapman​ ​and​ ​Hall,​ ​London.
Borror​ ​DJ,​ ​Triplehorn​ ​CA,​ ​Johnson​ ​NF.​​ ​1986.​ ​An​ ​introduction​ ​to​ ​the​ ​study​ ​of​ ​insects.
Saunders,​ ​Philadelphia.​ ​875pp.
Buck​ ​JB,​ ​Keister​ ​ML.​​ ​ ​1950.​ ​Periplaneta​ ​americana​,​ ​pp475-496​ ​in​ ​Brown​ ​FA​ ​(ed),​ ​Selected
invertebrate​ ​types.​ ​ ​ ​Wiley,​ ​New​ ​York.​ ​596pp.
Cornwell​ ​PB.​​ ​ ​1968.​ ​The​ ​cockroach,​ ​vol​ ​1,​ ​A​ ​laboratory​ ​insect​ ​and​ ​industrial​ ​pest.​ ​Hutchinson​ ​of
London.​ ​391pp.
Ebling​ ​.​ ​W.​​ ​1975​ ​Urban​ ​entomology​ ​Univ.​ ​Calif,​ ​Richmond​ ​CA
Furman​ ​DP,​ ​Catta​ ​EP.​​ ​1970.​ ​Manual​ ​of​ ​medical​ ​entomology,​ ​3​ rd​ ​ ​ ​ed.​ ​National​ ​Press​ ​Books,​ ​Palo
Alto.​ ​163​ ​pp.
Guthrie​ ​DM,​ ​Tindall​ ​AR.​​ ​ ​1968.​ ​The​ ​biology​ ​of​ ​the​ ​cockroach.​ ​Edward​ ​Arnold​ ​Publisher.​ ​408pp.
McKittrick​ ​FA.​​ ​1964.​ ​Evolutionary​ ​studies​ ​of​ ​cockroaches.​ ​Cornell​ ​Univ.​ ​Agric.​ ​Expt.​ ​Sta.​ ​Mem.
389.
Miall​ ​LC,​ ​Denny​ ​A.​ ​ ​ ​1886.​ ​The​ ​structure​ ​and​ ​life​ ​history​ ​of​ ​the​ ​cockroach.​ ​Lovell​ ​Reed
Company,​ ​London.​ ​224pp.
Ruppert​ ​EE,​ ​Fox​ ​RS,​ ​Barnes​ ​RB.​ ​ ​ ​2004.​ ​Invertebrate​ ​Zoology,​ ​A​ ​functional​ ​evolutionary
approach,​ ​7​ th​
​ ​ ​ed.​ ​Brooks​ ​Cole​ ​Thomson,​ ​Belmont​ ​CA.​ ​963​ ​pp.
Snodgrass​ ​RE.​​ ​1935.​ ​Principles​ ​of​ ​insect​ ​morphology.​ ​McGraw-Hill,​ ​New​ ​York.​ ​667pp.
​ ​Stang​ ​LA​.​ ​University​ ​of​ ​Florida​ ​Institute​ ​of​ ​Food​ ​and​ ​Agricultural​ ​Sciences,​ ​Evania

God​ ​gives​ ​knowledge