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Famiglia Agaonidae Walker 1846 Dott Famiglia Agaonidae Walker 1846 Dott. Francesco Fiume L’inquadramento tassonomico (classificazione scientifica ) degli Agaonidae è il seguente: Superdominio Biota Dominio Eucariota Whittaker & Margulis,1978 Regno Animalia Linnaeus 1758 Superphylum Panarthropoda Nielsen 1995 Phylum Arthropoda von Siebold 1848 Subphylum Pancrustacea Zrzavý & Štys, 1997 Superclasse Hexapoda Latreille, 1825 Classe Insecta Linnaeus 1758 Sottoclasse Dicondylia Hennig 1953 Infraclass Pterygota Brauer 1885 Clade Metapterygota Börner 1909 Clade Neoptera Woodworth 1906 Clade Eumetabola Hennig 1953 Clade Holometabola Burmeister 1835 Superordine Hymenopterida Boudreaux 1979 Ordine Hymenoptera Linnaeus 1758 Clade Neohymenoptera Grimaldi & Engel 2005 Clade Euhymenoptera Engel & Grimaldi 2004 Subordine Apocrita Gerstaecker 1867 Infraordine Proctotrupomorpha Latreille 1802 Superfamily Chalcidoidea Latreille 1817 Descrizione Gli Agaonidae sono caratterizzati da un marcato dimorfismo sessuale: la femmina è un insetto di piccole dimensioni, dotato di ali con venatura marginale perpendicolare al margine anteriore, mentre il maschio è in genere privo di ali (o con ali molto rudimentali). Le zampe mediane sono visibilmente più sottili di quelle anteriori e posteriori. Le femmine hanno un ovopositore chiaramente estroflesso, in alcune specie molto lungo. I maschi hanno robuste mandibole. I caratteri diagnostici principali della famiglia Agaonidae sono di seguito sintetizzati: 1. Ala anteriore con vena marginale perpendicolarmente al margine anteriore dell'ala o quasi (nell’80% dei casi). 2. Le specie sono associate ai fichi ( Ficus spp.) nel 100% dei casi. 3. Gli arti medi con i femori e le tibie sono chiaramente molto più snelli rispetto a quelli degli arti anteriori e posteriori (per 85% dei casi). 4. Le femmine con un chiaro e attivo, a volte lunghissimo, ovopositore (nel 99% dei casi). 5. I maschi presentano le ali molto ridotte o assenti, con mandibole larghe per il combattimento. Adattamenti morfologici Le specie femminili impollinatrici del fico sono alate e generalmente scure, mentre i maschi sono per lo più privi di ali e chiari. Questa differenza di colore è probabilmente dovuta a una notevole dicotomia del ruolo del genere (sesso). Una volta che si sono accoppiati, i maschi e le femmine hanno destini diversi. In alcune specie di fico, come il Ficus subpisocarpa o il Ficus tinctoria , i maschi devono, con il loro apparato boccale, costruire un foro per consentire alle femmine di lasciare il frutto in cui sono nate. Le femmine alate possono volare per lunghe distanze prima di trovare un altro fico per ovideporre in esso, mentre il maschio muore dopo la costituzione del foro di sfarfallamento delle femmine. Poiché il frutto di fico è provvisto di un ostiolo molto stretto, le femmine hanno sviluppato adattamenti per penetrare all’interno. Infatti, le mandibole delle femmine hanno sviluppato appendici specializzate per consentire la penetrazione nei frutti di fico. Queste appendici sono adattate alle specie fico ospitante, per cui, ad esempio, per ostioli spiralati la specie impollinatrice possiede appendici mandibolari inferiori a spirale, specializzate per penetrare in quell’ostiolo ( Van Noort et Compton, 1996 ). Gli Agaonidae che non sono impollinatori hanno sviluppato impressionanti adattamenti morfologici per depositare le uova all'interno del frutto di fico dall'esterno, sotto forma di un lunghissimo ovopositore. Coevoluzione La maggior parte delle specie della famiglia Agaonidae sono impollinatori dando luogo alla cosiddetta impollinazione entomofila determinando il trasporto del polline dalle antere dei fiori allo stigma del gineceo fiorale. La pianta ed il relativo imenottero impollinatore sono completamente dipendenti l'uno dall'altro, dal momento che ogni specie di Ficus può essere impollinata solo dalla appropriata specie di insetto, e ogni specie di insetto non può riprodursi al di fuori dell’appropriata specie di pianta. È il caso, per citare l'esempio più noto, del fico comune ( Ficus carica ) e della Blastophaga psenes Linnaeus 1758. Una relazione così stretta fa supporre che entrambi i gruppi discendano da progenitori comuni: un Ficus ancestrale ed il suo insetto impollinatore, da cui per successiva speciazione si sono evolute le attuali specie. Si tratterebbe di un tipico caso di coevoluzione, anche se l'esatto meccanismo di tale fenomeno non è ancora del tutto compreso. Esistono delle eccezioni: per esempio due differenti specie africane, Ceratosolen arabicus Mayr 1906 (figura 1) e Ceratosolen galili Wiebes 1964 (figura 2), possono interagire sia con Ficus sycomorus L. sia con Ficus mucuso Ficalho 1884. Figura 1 – Femmina di Ceratosolen arabicus su Ficus sycomorus , in atto di ovideporre. 2 Figura 2 – Femmina di Ceratosolen galili in atto di entrare all’interno del fico attraverso l’ostiolo,una stretta apertura all’apice. Biologia Le specie delle famiglia degli Agaonidae sono sempre associati con i fichi; molti agiscono come impollinatori di varie specie di Ficus spp. ( Agaoninae ), mentre altri sono probabilmente parassitoidi di impollinatori o di specie che producono galle di diverse parti del fico. I fichi e la specie ad essi associati, costituiti dagli impollinatori della sottofamiglia Agaoninae sono totalmente dipendenti gli uni dagli altri, dal momento che i fiori di fico possono essere impollinati solamente da specie altamente specifiche di Agaoninae e specie di Agaoninae non possono produrre progenie di fuori del fico appropriato e specifico. Il rapporto è quindi molto stretto ed i due gruppi probabilmente sono discendenti di un antenato-fico comune e del corrispondente impollinatore-vespa. Tassonomia La famiglia Agaonidae comprende tradizionalmente 71 generi e 723 specie, suddivisi in 6 sottofamiglie. In realtà esistono divergenze tra gli entomologi sugli esatti confini della famiglia. Recenti studi filogenetici, basati sulle analisi degli acidi nucleici, indicherebbero che solo le specie della sottofamiglia delle Agaoninae dovrebbero essere classificate come Agaonidae mentre Sycoecninae , Otitesellinae e Sycoryctinae andrebbero incluse nella famiglia Pteromalidae . Ancora incerta, secondo tali studi, la collocazione di Sycophaginae ed Epichrysomallinae . In linea di massima dovremmo distinguere la famiglia Agaonidae in nove sottofamiglie: 1. Tetrapusinae Rasplus & Van Noort 2010 2. Kradibiinae Rasplus & Van Noort 2010 3. Blastophaginae Kirchner 1867 4. Sycoecinae Hill 1967 5. Sycophaginae Walker 1875 6. Sycoryctinae Wiebes 1966 7. Otitesellinae Joseph 1964 8. Epichrysomallinae Joseph 1964 9. Agaoninae Walker 1846 I generi appartenenti alle sottofamiglie sono di seguito indicati. 3 Alla sottofamiglia Tetrapusinae appartiene il genere: a. Tetrapus Mayr 1885. Alla sottofamiglia Kradibiinae appartengono i seguenti generi: a. Ceratosolen Mayr 1885 b. Kradibia Saunders 1883 Alla sottofamiglia Blastophaginae appartengono i seguenti generi: a. Wiebesia Boucek 1988 b. Blastophaga Gravenhorst 1829 c. Valisia (Wiebes, 1993) A. Cruaud et al., 2009 d. Dolichoris Hill 1967 Alla sottofamiglia Sycoecinae appartengono i seguenti generi: a. Sycoecus Waterston 1914 b. Seres Waterston 1919 c. Philocaenus Grandi 1952 d. Crossogaster Mayr 1885 e. Diaziella Grandi 1928 f. Robertsia Bou ček 1988 Alla sottofamiglia Sycophaginae appartengono i seguenti generi: a. Anidarnes Bou ček 1993 b. Apocryptophagus Ashmead 1904 c. Eukoebelea Ashmead 1904 d. Idarnes Walker 1843 e. Pseudidarnes Girault 1927 f. Sycophaga Westwood 1840 Alla sottofamiglia Sycoryctinae appartengono i seguenti generi: a. Apocrypta Coquerel 1855 b. Arachonia Joseph 1957 c. Bouceka Koçak & Kemal d. Critogaster Mayr 1885 e. Dobunabaa Bou ček 1988 f. Parasycobia Abdurahiman 1967 g. Philotrypesis Foerster 1878 h. Sycoscapter Saunders 1883 i. Watshamiella Wiebes 1981 Alla sottofamiglia Otitesellinae appartengono i seguenti generi: a. Aepocerus Mayr 1885 b. Comptoniella Wiebes 1992 c. Eujacobsonia Grandi 1923 d. Grandiana Wiebes 1961 e. Grasseiana Abdurahiman 1968 f. Guadalia Wiebes 1967 g. Heterandrium Mayr 1885 h. Lipothymus Grandi 1922 i. Micranisa Walker 1875 j. Micrognathophora Grandi 1922 k. Otitesella Westwood 1883. l. Philosycella Abdurahiman 1976 m. Philosycus Wiebes 1969 n. Walkerella Westwood 1883 Alla sottofamiglia Epichrysomallinae appartengono i seguenti generi: a. Acophila Ishii 1934 4 b. Asycobia Bou ček 1988 c. Camarothorax Mayr 1906 d. Epichrysomalla Girault 1915 e. Eufroggattisca Ishii 1934 f. Herodotia Girault 1931 g. Josephiella Narendran 1994 h. Meselatus Girault 1922 i. Neosycophila Grandi 1923 j. Odontofroggatia Ishii 1934 k. Parapilkhanivora Farooqi 1973 l. Sycobia Walker 1871 m. Sycotetra Bou ček 1981 Alla sottofamiglia Agaoninae appartengono i seguenti generi: a. Pleistodontes Saunders 1883 b. Nigeriella Wiebes 1974 c. Agaon Dalman 1818 d. Paragaon Joseph 1959 e. Allotriozoon Grandi 1916 f. Courtella Kieffer 1912 g. Elisabethiella Grandi 1928 h. Alfonsiella Waterston 1920 i. Waterstoniella Grandi 1921 j. Eupristina Saunders 1882 k. Pegoscapus Cameron 1906 l. Platyscapa Motschulsky 1864 m. Deilagaon Wiebes 1977 Con riferimento alla tassonomia della famiglia Agaonidae, va precisato che questa è cambiata più volte dalla sua comparsa nelle classificazioni scientifiche e dopo i lavori di Francesco Walker nel 1846 (Walker, 1846) è stata descritta nuovamente partendo dal genere Agaon . Per molto tempo le sottofamiglie Epichrysomallinae , Otitesellinae , Sycoecinae , Sycoryctinae , Sycophaginae , e Agaoninae sono state le suddivisioni della famiglia (Bou ček, 1988). I recenti
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