Marco Uliana

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Marco Uliana CONTENTS RIASSUNTO .............................................................................................................. 3 ABSTRACT ............................................................................................................... 5 INTRODUCTION ....................................................................................................... 7 MATERIALS AND METHODS .................................................................................. 10 Sources of study material .............................................................................. 10 Microscopy and imaging ............................................................................... 10 Phylogenetics ................................................................................................ 11 Evaluation of chromatic conditions and level of approximation .................. 12 COLOURS AND CHROMATIC EFFECTS IN BEETLES ............................................... 13 Colours producing devices: pigments ............................................................... 13 Darkening and sclerotisation of the cuticle .................................................. 13 Physical colours ................................................................................................ 14 Physical colours: multilayer reflectors.............................................................. 16 Broadband reflectors .................................................................................... 18 “Pointillistic” colour mixing ........................................................................ 18 Physical colours: diffraction gratings................................................................ 19 Quasi-ordered scattering .............................................................................. 20 Physical colours: photonic crystal structure and scales .................................... 20 Tridimensional non-crystalline structures and whites .................................. 22 Reversible colour change .................................................................................. 23 PATTERN EVOLUTION AND EVO -DEVO ASPECTS................................................. 25 Anatomy and colour pattern.............................................................................. 25 Muscles insertions and melanization ............................................................ 25 The elytron: vein patterns, punctuation and sculpture ................................. 26 Secondary sexual characters and colours .................................................... 30 Hair and scales .................................................................................................. 32 Convergence in colour patterns of sympatrically occurring beetles ................. 35 STRUCTURAL AND EVOLUTIONARY ASPECTS IN CHRYSOLINA ............................. 39 The Chrysomelidae: a taxonomic and phylogenetic outline............................. 39 Taxonomy of the Chrysomelinae and of Chrysolina (s.l.) ............................. 40 Chrysolina and Oreina phylogenetics ........................................................... 42 1 Chrysolina phylogeny........................................................................................43 Chromatic patterns in Chrysolina......................................................................46 Main colour ...................................................................................................46 Elytra/forebody relations ...............................................................................47 Physical patterns ...........................................................................................50 Box: The fastuosa like pattern and its variations………………………… 54 Pigmentary patterns ......................................................................................57 The origin of physical colours and the evolution of the black phenotypes .......59 CONCLUSIONS .......................................................................................................63 LITERATURE ..........................................................................................................68 APPENDICES ........................................................................................................100 Appendix 1. Species used in the phylogenetic reconstruction ........................100 Appendix 2. Characters coding .......................................................................101 Appendix 3. Data matrix. ................................................................................105 RINGRAZIAMENTI ...............................................................................................107 2 RIASSUNTO I colori e i pattern cromatici presenti sui tegumenti degli insetti rappresentano da tempo un soggetto di interesse per numerosi ricercatori. Tuttavia, nonostante pattern cromatici complessi si riscontrino presso numerosi ordini di insetti, lo sforzo di ricerca è stato distribuito in modo eterogeneo, concentrandosi in particolar modo sui lepidotteri. Grazie a una lunga tradizione di ricerca, infatti, i complessi disegni che si sviluppano sulle ali delle farfalle sono oggi conosciuti con notevole approfondimento negli aspetti biochimici e ultrastrutturali, nei meccanismi di sviluppo e nelle loro basi genetiche, nella composizione ed evoluzione degli elementi discreti che li compongono, nonché negli aspetti di interazione con l'ambiente. In tempi più recenti, anche i pattern cromatici che caratterizzano i ditteri drosofilidi sono stati studiati in dettaglio negli aspetti strutturali, morfogenetici ed evolutivi. Meno noti sono i fenomeni cromatici che si riscontrano presso altri gruppi di insetti, fra cui i coleotteri, ordine a cui si ascrivono circa 350.000 specie note. In particolare, sono stati ampiamente trascurati studi comparativi, interpretativi o evolutivi condotti a livello interspecifico e paragonabili a quelli condotti sugli altri gruppi citati. L’esplorazione della bibliografia specifica, infatti, ha permesso di individuare solo una scarna serie di lavori a riguardo, alcuni dei quali risalenti ai primi del ‘900. Questi si rivelano del tutto insufficienti a fornire un quadro generale sui fenomeni che governano i pattern cromatici dei coleotteri. Ad oggi non abbiamo che pochissime notizie sui meccanismi di controllo e di sviluppo dei loro pattern cromatici, sulle loro eventuali relazioni con le venulazioni delle ali (ampiamente verificate in lepidotteri e ditteri) o con altri elementi morfoanatomici, così come sulle loro capacità e tendenze evolutive. Con questa tesi si sono voluti muovere i primi passi verso una riduzione dell’enorme lacuna di conoscenza ora delineata. Lo studio è stato condotto su due livelli diversi. Ad un primo livello, di carattere più generale, si sono voluti esplorare, dopo una accurata valutazione della letteratura, alcuni fenomeni di interesse generale, a cominciare dall’analisi di quelli più basilari: quelli, cioè, che determinano la produzione dei colori. Sono stati privilegiati, in questo caso, i diversi fenomeni che determinano la produzione di colori fisici in quanto, oltre a collegarsi direttamente con la seconda parte dello studio, sono quelli che presentano le maggiori potenzialità evolutive. Questa sezione ( Colours and chromatic effects in beetles ) si basa principalmente su un’accurata rassegna e analisi critica della bibliografia, che è stata integrata con alcune osservazioni originali. Nella sezione successiva (Pattern evolution and Evo-Devo aspects ) si indagano alcuni aspetti dei pattern cromatici secondo una prospettiva “Evo-Devo” (così come viene comunemente detta la moderna la biologia evoluzionistica dello sviluppo), in particolare discutendo le relazioni che essi intraprendono con elementi anatomici del tegumento. In questi capitoli viene suggerito che le attività di induzione/controllo/alterazione dei pattern (sia pigmentari che di origine fisica) prodotte dagli elementi morfologici dell’elitra (punti, strie, vene) rivestano un ruolo di prim’ordine nello sviluppo e nell’evoluzione dei pattern. Lo stesso viene 3 suggerito a proposito delle inserzioni muscolari, per le quali viene verificata la capacità di induzione di un pattern cromatico, confermando e ampliando la documentazione di un importante dato bibliografico risalente ai primi del 1900, ma in seguito apparentemente dimenticato. Infine, viene discusso un interessante, ricorrente fenomeno di convergenza cromatica intraspecifica legata alla distribuzione geografica, per il quale sembrano improbabili spiegazioni legate alla selezione darwiniana. Nella seconda parte della tesi (Structural and evolutionary aspects in Chrysolina) si è affrontata una ricostruzione dell’evoluzione dei pattern cromatici all’interno di un gruppo di coleotteri. A questo scopo, è stato individuato come utile candidato il genere Chrysolina (Chrysomelidae Chrysomelinae) inteso in senso lato, cioè comprendente alcuni generi affini di incerta collocazione tassonomica. La possibilità di accedere ad ampie collezioni entomologiche mi ha permesso di condurre una vasta ricognizione sulla quasi totalità dei sottogeneri e delle specie esistenti a livello mondiale. Le condizioni cromatiche osservate su decine di migliaia di esemplari sono state ricondotte a un numero limitato di pattern cromatici e mappate all’interno di una tabella sinottica con risoluzione a livello di sottogenere (attualmente si considerano validi circa 65 sottogeneri per il solo genere Chrysolina
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