2018 ACTA ENTOMOLOGICA 58(1): 187–193 MUSEI NATIONALIS PRAGAE doi: 10.2478/aemnp-2018-0016 ISSN 1804-6487 (online) – 0374-1036 (print) www.aemnp.eu SHORT COMMUNICATION Presence of uradenia in male adults of the genus Dismegistus (Hemiptera: Heteroptera: Parastrachiidae) Dominique PLUOT-SIGWALT1) & Jerzy A. LIS2) 1) Museum national d’Histoire naturelle, Département Adaptation du Vivant, UMR 7179 MECADEV (mécanisme adaptatif et évolution), 57 rue Cuvi- er, CP 50 (Entomologie), F-75231 Paris, France; e-mail: [email protected] 2) Department of Biosystematics, Opole University, Oleska 22, PL-45-052 Opole, Poland; e-mail: [email protected] Accepted: Abstract. Uradenia, often referred to as paragenital glands, are usually voluminous paired th 8 June 2018 exocrine glands located ventrally in the abdomen mostly on the intersegmental membrane Published online: between abdominal segments (= urites) VII–VIII or VIII–IX, depending on sex or the taxon. 20th June 2018 They have been previously recorded from eight pentatomomorphan families belonging to Coreoidea, Lygaeoidea and Pyrrhocoroidea (Hemiptera: Heteroptera), found either in males, females or both sexes, and were thought to be absent in Pentatomoidea. We report here the fi rst instance of uradenia in a pentatomoid genus, the African Dismegistus Amyot & Serville, 1843 (Parastrachiidae). Only the male adult possesses uradenia located on the intersegmental membrane of segments VIII–IX. The only other genus of the family, Parastrachia Distant, 1883, as well as other examined genera belonging to pentatomoid families possibly related to Parastrachiidae (Cydnidae, Thyreocoridae), do not possess uradenia. The uradenia of Dismegis- tus exhibit the same fundamental structure as in other trichophoran families but differ by their dorso-lateral position (instead of ventral), and also by the paired orifi ces (instead of unpaired and median). The implications of the presence of uradenia within member of a pentatomoid genus are briefl y discussed. Key words. Hemiptera, Heteroptera, Pentatomoidea, Parastrachiidae, Dismegistus, Parastra- chia, morphology, abdominal segment VIII, ectodermal gland, uradenia http://zoobank.org/urn:lsid:zoobank.org:pub:B7323537-1320-4E65-A6CD-1448C32A1010 © 2018 The Authors. This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Licence. Introduction 1965), “ventral uradenial scent glands” in Coreidae (STAD- The term “uradenia” – from the Greek, ourà and adên DON 1986), and fi nally “Uradenien” (KALLENBORN 2005). meaning: “posterior part” and “gland” – has been proposed In her well-documented comparative study, THOUVENIN by THOUVENIN (1965) to name paired ventral abdominal (1965) showed that the paired uradenia are present in eight glands which are present on the last abdominal segments trichophoran families (Coreoidea: Alydidae, Coreidae, (= urites) in several representatives of Lygaeoidea, Pyr- Stenocephalidae; Pyrrhocoroidea: Largidae, Pyrrhocori- rhocoroidea and Coreoidea. Previously, these glands were dae; Lygaeoidea: Lygaeidae, Rhyparochromidae), either in variously referred to as “Oeldrüsen” in Pyrrhocoris Fallén, male, or in female, or in both sexes. Glands open directly 1814 (MAYER 1874, 1875), “Drüsenschlauch” in Lygaeus to the exterior through the intersegmental membrane Fabricius, 1794 (LUDWIG 1926), “paired accessory glands” between abdominal segments VII–VIII or VIII–IX, or on in Dysdercus Guérin-Méneville, 1831 (GUPTA 1951), “pa- abdominal segment IX (see Table 1), usually laterally in ragenital glands” in females and “subgenital glands” in the female, in the mid-ventral line in the male through males of Oncopeltus Stål, 1868 (BONHAG & WICK 1953), a single orifi ce, both glands being united just before the “paragenital glands” in Trichophora (ŠTYS 1962), “sac- outlet into a short common efferent duct. In addition to like structures” in males of Dicranocephalus Hahn, 1826 detailed morphological data, THOUVENIN (1965) gave some (LANSBURY 1965), “glande à huile” in Pyrrhocoris (MERLE information on the histological structures of the glands in Pluot.indd 187 20.6.2018 17:34:55 188 PLUOT-SIGWALT & LIS: Presence of uradenia in Dismegistus (Heteroptera: Parastrachiidae) Table 1. Taxonomic distribution of the male and female uradenia in the main trichophoran families according to THOUVENIN (1965) and this study. Superfamily Taxon Female uradenia Male uradenia Genera or tribes explicitely examined Coreoidea Alydidae absent VII–VIII, lobate, single median orifi ce Coreidae VIII–IX, bi- or multilobate, paired VII-VIII, tubular or lobate, single Puppeia, Dalader, lateral orifi ces median orifi ce Holopterna Coreidae: Petascelini VII–VIII, branched, paired lateral VII–VIII, multibranched, paired Petascelis, Oxypristis (partim) orifi ces orifi ces + lateral gl. IX on each side of the phallus Coreidae: Petascelini VIII–IX [no data] Petascelisca (partim) Rhopalidae absent VII–VIII, lobate, single median orifi ce Stenocephalidae absent VII–VIII, sac-like, paired median Dicranocephalus orifi ces in a wide inter-segmental fold Pyrrhocoroidea Pyrrhocoridae IX, branched with basal reservoir, absent Pyrrhocoris, Dys- paired lateral orifi ces dercus, Dermatinus, Callibaphus Largidae: Larginae IX, reservoir + many diverticula, absent Largus, Stenomacra paired lateral orifi ces Largidae: Physopeltinae absent absent Lygaeoidea Lygaeidae VIII–IX near IX, tubular apically VIII–IX, tubular or clavate, single Lygaeus, Oncopeltus, swollen, paired lateral orifi ces median orifi ce Lygaeosoma, Nysius Rhyparochromidae IX, paired orifi ces (save unpaired in [no data] Beosus, Megalonotini, (mentioned as Lygaeidae) Cleradini), tubular, apically globose Myodochini Pentatomoidea Parastrachiidae absent VIII-IX, branched with basal reser- Dismegistus voir, paired orifi ces absent absent Parastrachia remaining taxa absent absent Remarks. In her study, THOUVENIN (1965), unfortunately, did not provide a list of the species she examined. She only mentioned in the text some names of species or genera having glands and certainly not all the taxa (apparently numerous) she examined; the genera or tribes mentioned in her paper are included in the table. both sexes. Histological and histochemical aspects of the Therefore, the discovery in adult males of the genus female uradenia were also reported by CHEVAILLIER (1965) Dismegistus Amyot & Serville, 1843, of a paired abdomi- and FARINE (1988) in female Dysdercus. For the sake of nal gland exhibiting morphological characteristics of the completeness, we must add that CARAYON (1954) also uradenia previously described within other trichophoran described paired ventro-abdominal glands in the males of subfamilies, was unexpected. It is interesting especially Anthocoridae (Scolopini), on sternite IV, rarely V; these because the phylogenetic position of Dismegistus within glands were later recognized as uradenia by him. CARAYON Pentatomoidea has been a matter of debate for a long time (1972) considered that they were serially homologous of and still remains somewhat enigmatic (LIS et al. 2017). the uradenia described by THOUVENIN (1965) in trichopho- The African genus Dismegistus includes six described ran families. aposematic species, and was originally placed in Cydni- Very little is known about the function of these glands. In dae Sehirinae by AMYOT & SERVILLE (1843). Later, on the vivo, the secretion has an oily and yellowish aspect at least basis of morphological characters, it was transferred to in Coreidae and Pyrrhocoridae (MAYER 1874, 1875; MERLE Pentatomidae Pentatominae Strachiini (SIGNORET 1881), 1965; DPS pers. observations) and also in Anthocoridae to Pentatomidae Asopinae (BERGROTH 1923) jointly with (CARAYON 1954). Only male uradenia in coreid bugs (species the Asiatic genus Parastrachia Distant, 1883, including of Pachylis Le Pelletier & Serville, 1825, Euthochtha Mayr, only two aposematic species and having been placed into 1865, and mainly Leptoglossus Guérin-Méneville, 1831 various pentatomid groups (Asopinae, Pentatominae, were studied (as “ventral abdominal gland”) and evidence Tessaratomidae, Cydnidae) (see SCHAEFER et al. 1988 and for male-produced sex pheromones has been demonstrated SWEET & SCHAEFER 2002) until LESTON (1956), after a large (ALDRICH & YONKE 1975; ALDRICH et al. 1976, 1979, 1982; comparative study of morphological characters including GOUGH et al. 1985; ALDRICH 1988; WANG & MILLAR 2000). male and female genitalia, proposed the placement of The secretion of the male uradenia releases species-specifi c Dismegistus among the Cydnidae Sehirinae. After having volatile compounds that may act as long-range attractants rectifi ed several of Leston’s misinterpretations, DOLLING for the female; it may have the odour of cherries, vanilla, (1981) concluded that Dismegistus cannot belong to Cyd- cinnamon and rose (ALDRICH 1988). nidae. More recently, PLUOT-SIGWALT & LIS (2008) showed Uradenia are considered as lacking within the Penta- the great similarity in the structure of the spermatheca in tomoidea (THOUVENIN 1965, STADDON 1979, PAVIS 1987). Dismegistus and Parastrachia, suggesting close relation- Pluot.indd 188 20.6.2018 17:34:55 Acta Entomologica Musei Nationalis Pragae, volume 58, number 1, 2018 189 ships between these genera. In a phylogenetic analysis of mental or intersegmental origin of the glands, i.e., uradenia the Pentatomoidea based on morphological and molecular VII–VIII or VIII–IX open on intersegmental membrane characters, GRAZIA et al. (2008) recognized the group of delimited by abdominal segments VII–VIII