Abstract
Cuticular parts of the spermatheca and associated vaginal structures (chiefly the ring sclerites of the parietovaginal glands) have been examined and compared in 190 cydnid species representing 65 genera and all five subfamilies currently recognized in the family (Amnestinae, Cephalocteinae, Cydninae, Garsauriinae, Sehirinae). Four species belonging to genera formerly included within the Cydnidae (Dismegistus, Parastrachia, Thaumastella, Thyreocoris) were also examined. Morphology of the three main parts of the spermatheca [seminal receptacle (distal bulb), intermediate part (pump apparatus), spermathecal duct] is described. Four main types of spermathecae can be recognized from the distal receptacle and the intermediate part: the amaurocorine type (in Sehirinae: Amaurocorini), amnestine type (in Amnestinae), garsauriine type (in Garsauriinae), and "cydnoid" type (in Cephalocteinae + Cydninae: Cydnini, Geotomini + Sehirinae: Sehirini). No synapomorphy of these types was found which suggests that the currently conceived Cydnidae are not monophyletic. Moreover, out of these four types only the "cydnoid" is typically pentatomoidean due to the presence of an intermediate part usually well delimited by two flanges and having always an unsclerotized flexible zone as well as two internal cuticular structures (septum and fretum) partly obstructing the lumen. The simple tubular amaurocorine type is unusual and aberrant within all Pentatomoidea. The amnestine and garsauriine types display some similarities with taxa outside the Pentatomoidea, especially with some lygaeoid or coreoid spermathecae, mainly in the structure of the intermediate part not delimited proximally (absence of flanges) and devoid of the flexible zone. Within the "cydnoid" type, six spermathecal facies can be characterized principally according to the shape of both the apical reservoir along with the intermediate part, and the differentiations of the spermathecal duct. It has been impossible to find any synapomorphy for all species and for the six facies belonging to the "cydnoid" type of spermatheca. We suggest that the Cydnidae as defined presently are probably a polyphyletic group; moreover its main "cydnoid" branch, called by us Cydnidae sensu stricto (Cephalocteinae + Cydninae + Sehirinae: Sehirini) seems to be relatively recent among the Pentatomoidea. Nishadana and Nishocoris are transferred from Garsauriinae back to Cydninae: Cydnini and the tribe Amaurocorini (Sehirinae) is upgraded to a separate subfamily Amaurocorinae stat. nov. Moreover, we regard the Geotomini and the Sehirini both as non-monophyletic and we indicate that by appending them sensu lato (Geotomini "s. l.", Sehirini "s. l.")
Highlights
In Heteroptera the structure of the spermatheca, often highly complex, shows a great diversity and was found to exhibit many important characters for classification, taxonomy and phylogeny (Pendergrast, 1957).Little attention has been paid until now to the spermatheca in the Cydnidae, a family often considered a key taxon to understand the relationships and origin of the Pentatomoidea, but still an unsatisfactorily defined group of Pentatomoidea
The intention of the present study is to give a general view of the diversity and complexity of the spermatheca within the family, to compare its structure and construction to those found in other pentatomomorphan spermatheca and to evaluate the importance of the spermatheca in the systematics and phylogeny of Cydnidae
From the results presented above, one may recognize four main types of spermathecae in the Cydnidae, which can be distinguished according to the number and structure of the differentiated parts: the amaurocorine, amnestine, garsauriine types, and a multifacied pentatomoid type, named here for convenience “cydnoid” type
Summary
In Heteroptera the structure of the spermatheca, often highly complex, shows a great diversity and was found to exhibit many important characters for classification, taxonomy and phylogeny (Pendergrast, 1957). The intention of the present study is to give a general view of the diversity and complexity of the spermatheca within the family, to compare its structure and construction to those found in other pentatomomorphan spermatheca and to evaluate the importance of the spermatheca in the systematics and phylogeny of Cydnidae. The first one brings precisions about the general structure of the cydnid spermatheca and terminology used; all spermathecal parts considered in the study are listed and illustrated using the geotomine spermatheca as example, because this type exhibits the highest complexity. We discuss the significance of the spermatheca for cydnid classification and phylogeny
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