Components of external genitalia in both sexes of ectognathous insects are homologous in all orders, item by item. They consist of presumed telopodites (gonostyli: female ovipositor sheaths, male claspers) and possible endiles (gonapophyses: female ovipositor, male aedeagus) (in (gono) coxites of abdominal segments VIII and IX. One or both pairs of gonapophyses and usually a pair of gonostyli are present in at least 1 sex of all insect orders, and these gonopods are homodynamous with api:eiu‗ages or derivatives on other body segments. Gonopods are annulated or segmented, and primitively identical in both sexes. Gonapophyses IX are rotated 180° laterally on their long axes, fused with each other along their new dorsal margins to form a bridge (notum), and either gonapophysis IX interlocked on its new ventral margin with a gonapophysis VIII. The interlock provides a track (olistheter) along which the gonapophyses can be moved. Rotation of IX had inverted rami and internal structure. The notum formed through fusion of the dorsal (euventral) rami of gonapophyses IX is a landmark in both sexes of nearly all insects. Articulation of gonocoxite VIII has shifted from tergite VIII to tergite IX immediately proximal to the insertion of gonocoxite IX to provide coordination between interlocked gonapophyses. Types of coordinated gonopod movement possible if gonapophyses IX are fused dorsally and interlocked ventrally with gonapophyses VIII include: (1) gonapophyses IX provide a stationary track for alternate thrusts of VIII (most common condition); (2) gonapophyses IX move synchronously in 1 direction as both gonapophyses VIII either move together in the other (some female Microcoryphia, hemipteroids), or remain stationary (some female Microcoryphia, Odonata); or (3) all gonapophyses move together in the same direction (some female Diptera, Coleoptera). Many female insects and all nnlcs have disengaged gonopods, and different mechanisms have evolved. Female insects retaining both sets of gonapophyses have faced 2 restrictions: (1) the distance VIII can be thrust in relation to IX is relatively small, and (2) little vertical leverage (as opposed to back-and-forth motion) is possible. The first has been ameliorated in the only Holometabola retaining the entire primitive ovipositor (Hymenoptera) by a change in articulation: gonocoxite IX has shifted from tergite IX to gonocoxite VIII. The 2nd restriction has been reduced in orthopteroids through articulation of the independently movable gonostyli IX with the independently movable gonapophyses VIII. Gonapophyses IX are reduced in higher orthopteroids and many Holometabola. Gonopods of somite VIII have been lost in male Pterygota (gonapophyses VIII are retained but disengaged from IX in Machilidae), and gonapophyses IX have been reduced to 1 segment (radix). Gonapophyses IX often are detached from gonocoxites IX, and the gonocoxites may be disarticulated from tergite IX. Olistheter components of either gonapophysis IX (rhachis) have fused mesally in many Pterygota to make a ventral floor and form an intromittent tube from the appendages. This cylinder may redivide between the fused rhachies and the ventral rami to either side, and the 2 rhachies left as a separate mesal sclerite. Upon reconstitution of this tube a 2nd time in some Holometabola through overgrowth of the ventral rami, the rhachies became internal within the gonapophyses. Gonostyli usually retain their musculature and typically have been modified into clasping organs. External genitalia of males may be augmented by coxosternal or tergal derivatives, fragmenta of gonapophyseal rami or gonocoxites, or by cerci. Gonodal gonoducts of both sexes appear to have opened originally through paired papillae on somite VIII, and accessory gonoducts, on IX. Papillae are outgrowths of muscles and membranes between the gonocoxites and coxosternites, and homologues of coelomoducal ampullae of other arthropods. Gonoducts have coalesced and migrated in various directions due to development of articulating gonopods. Papillae may remain as separate labia in females or phalli in males; in the latter, they are fused usually into a median penis. Both may supplant appendicular genitalia.
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