Anno Accademico 2011–2012 UNIVERSITÀ DEGLI STUDI DI TRIESTE FACOLTÀ DI SCIENZE MATEMATICHE, FISICHE E NATURALI CORSO DI LAUR

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Anno Accademico 2011–2012 UNIVERSITÀ DEGLI STUDI DI TRIESTE FACOLTÀ DI SCIENZE MATEMATICHE, FISICHE E NATURALI CORSO DI LAUR Anno accademico 2011–2012 UNIVERSITÀ DEGLI STUDI DI TRIESTE FACOLTÀ DI SCIENZE MATEMATICHE, FISICHE E NATURALI CORSO DI LAUREA IN SCIENZE BIOLOGICHE CURRICULUM: Biodiversità degli ecosistemi terrestri e marini “Strategie sessuali e capacità cognitive degli uccelli giardinieri” Laureanda: Degano Eleonora Relatore: Avian Massimo Correlatrice: Chiandetti Cinzia 1 CLASSIFICAZIONE GENERI: SCIENTIFICA: Ailuroedus Dominio: Eukaryota Amblyornis Regno: Animalia Archboldia Phylum: Chordata Chlamydera Classe: Aves Prionodura Ordine: Passeriformes Ptilonorhyncus Famiglia: Ptilonorhynchidae Scenopooetes Sericulus 1 Introduzione alla specie 2 Costruzione del pergolato 3 Adattamento al fuoco 4 Investimento parentale e fitness 5 Modelli di selezione sessuale 6 L’ipotesi del maschio brillante 7 Illusione ottica e la visione negli uccelli 8 Il concetto di arte 9 Abilità cognitive e di apprendimento 10 Studi sulla cognizione e dimensione del cervello Fig.1: Un esemplare femmina di Ptilonorhyncus violaceus 2 1 - Introduzione alle specie Gli Ptilonorhynchidae o uccelli giardinieri (in inglese “bowerbirds”) sono una famiglia appartenente all’ordine dei Passeriformi. Ne conosciamo 20 specie, delle quali 10 vivono solamente in Nuova Guinea, 8 in Australia e 2 in entrambe le regioni. Il territorio abitabile dagli uccelli giardinieri copre circa due terzi della superficie di queste due regioni. L’habitat naturale comprende principalmente foreste umide, ad altitudini svariate sino a picchi di 4000 m (come nel caso dell’uccello giardiniere di Archbold, Archboldia papuensis). Molte specie sono localizzate in areali precisi come l’uccello giardiniere crinito (Serikulus bakeri), confinato sui Monti Adelbert in Papua Nuova Guinea, gli uccelli giardiniere dorati (Prionodura newtoniana) e gli uccelli gatto dentati (Scenopooetes dentirostris), esclusivi delle foreste pluviali del Queensland sul tavolato di Atherton. Altre specie, come l’uccello giardiniere flammeo (Sericulus aureus) e il giardiniere maggiore (Chlamydera nuchalis), occupano invece areali continui ed estesi. Gli uccelli giardinieri hanno dimensioni variabili da quelle di uno storno a quelle di un corvo, con piedi massicci e becchi pesanti. Tendenzialmente sono lungoìhi circa 30 cm coda compresa; un maschio pesa tra 173-290 g, una femmina 170-258 g. La colorazione del piumaggio è variabile e la maggior parte degli esemplari maschi presenta tinte vivaci: oro brillante, arancio e nero (uccelli giardinieri flammei, uccello giardiniere crinito e uccello giardiniere di Archbold), blu-nero iridescente, giallo brillante e oliva nell’uccello giardiniere dorato, con creste colorate che vengono erette durante il display, giallo e arancio brillante nell’uccello giardiniere flammeo (Sericulus aureus) (Fig.2). Fig.2: Un uccello giardiniere flammeo presso il suo pergolato 3 Altre specie come il maggiore, il pettofulvo (Chlamydera cerviniventris) e il pettogiallo (Chlamydera lauterbachi), i quali vivono principalmente nascosti tra la vegetazione erbacea e nei boschi più aridi, presentano un abito dai toni grigi/marroni con piccole creste rosate sulla nuca; queste colorazioni più discrete sono tipiche anche degli esemplari femmine di molte specie in quanto si prestano alla mimetizzazione nell’ambiente. Gli uccelli giardinieri si nutrono principalmente di frutti, fiori, foglie, insetti (cicale, maggiolini ed altri artropodi) e talvolta di piccoli animali. Molte delle specie (principalmente quelle costruttrici) si riuniscono in stormi durante l’inverno e possono assaltare frutteti e piante erbacee rivelandosi una vera calamità (Frith 1991, Rowland 2008). 2 - Costruzione del pergolato I due rappresentanti più antichi del gruppo, l’uccello gatto macchiato (Ailuroedus melanotis) l’uccello gatto verde (Ailuroedus crassirostris) sono monogami, e la coppia alleva i piccoli insieme dopo aver costruito un nido; il modello è quello seguito da circa il 90% degli uccelli, che solitamente porta ad un incremento della fitness di entrambi, maschio e femmina. Un lascito all’apparenza inutile di quello che probabilmente è un ignoto antenato (forse un uccello lira del genere Menura) è riscontrabile in un comportamento mantenuto dalle coppie di uccelli gatto: essi allestiscono una rudimentale area circolare di foglie capovolte che utilizzano per esibirsi. Diversamente da questi ultimi, la prerogativa della maggior parte degli Ptilonorhynchidae è la costruzione di complessi pergolati e piattaforme allo scopo di attirare il maggior numero di femmine possibili con finalità riproduttive; delle 20 specie di uccelli giardinieri conosciute, sappiamo che 16 sono solite costruire tali strutture, e che sono tutte specie poligame. Questi pergolati sono talmente particolari ed atipici che per anni i ricercatori occidentali li pensarono opera di tribù delle foreste. Queste creazioni sono le più complesse che si siano mai viste nel mondo ornitologico; solamente castori ed esseri umani intraprendono opere che comportano un gran numero di fasi e gran flessibilità in termini di progettazione, materiali ed esecuzione. Tendenzialmente la maggior cura riscontrabile nella costruzione di nidi e strutture simili è volta alla mimetizzazione nell’ambiente o alla resistenza; ci sono pochissime eccezioni, ad esempio casi isolati di pivieri (Charadrius morinellus) che circondano il proprio nido con un cerchio di pietre e starne (Perdix perdix) che trascinano, con fatica non indifferente, cumuli di alghe sino ai nidi; in entrambi i casi non pare esservi un comprensibile scopo (Rowland 2008). Nelle foreste dell'Austrialia è facile dunque imbattersi nei pergolati degli uccelli giardinieri, costituiti da rametti tondeggianti posti in verticale e rivestiti con muschi e felci, decorati con conchiglie evo 4 fili colorati; nelle vicinanze degli insediamenti umani le decorazioni spesso includono oggetti come monete, mollette, penne a sfera, spazzolini da denti, tappi di bottiglie di plastica, argenteria, detriti vari, bottoni, vetri, unghie, viti, ditali e piccoli oggetti simili (Borgia 1985b, Madden 2003 e 2004). A volte le pareti del pergolato sono colorate con tinture naturali che l’animale ottiene mescolando la propria saliva a succhi prelevati da bacche, carbone vegetale o vegetazione masticata. E’ stato dimostrato che la pittura delle pareti, misurata in termini di ore spese dedicandosi a questa attività, porta ad un incremento della fitness del maschio performante con un aumento considerevole del numero di accoppiamenti (fig.3). Per quanto riguarda alcune specie, in particolare l’uccello giardiniere grigio (Ptylonorhyncus nuchalis), gli studiosi stanno ancora discutendo se il viale sia effettivamente dipinto oppure no: sebbene si siano osservati numerosissimi maschi intenti a tale operazione, il colore sembra ad occhio umano essere immutato. Va considerato, tuttavia, che gli uccelli percepiscono l’ultravioletto; se una componente della saliva dell’uccello riflette questa lunghezza d’onda, e il colore riflesso risulta attraente per le femmine, il comportamento potrebbe allora rivelarsi determinante. La saliva, inoltre, potrebbe contenere un odore attraente per le femmine che l’essere umano non è in grado di percepire, e che potrebbe rivelarsi un potentissimo additivo alla pittura. Fig.3: Probabilità binomiale che un esemplare maschio ottenga almeno una copula durante i mesi di monitoraggio del pergolato (in questo caso novembre e dicembre) come previsto dalla quantità di eventi di pittura/ora. I punti rappresentano -per ogni categoria divisa in base al numero di eventi di pittura/ora- la percentuale di maschi che ottennero almeno un accoppiamento durante il periodo di osservazione; i numeri vicino ai punti indicano il numero di osservazioni su ogni singolo esemplare della popolazione in studio. 5 I pergolati non sono nidi: sono infatti le femmine a costruire il nido vero e proprio sugli alberi, dove senza alcun contributo da parte del maschio con il quale si sono accoppiate alleveranno la prole (Borgia 1985b, Madden 2003). I più studiati e noti tra le varie specie conosciute sono gli uccelli giardinieri satinati (Ptylonorhincus violaceus), i quali costruiscono un pergolato relativamente modesto. In un primo momento avviene la scelta di un sito adeguato, un luogo aperto e pianeggiante con un buon passaggio di luce; il maschio pulisce l’area da eventuali foglie residue agendo anche sulla vegetazione circostante, in modo da rendere la zona prescelta il più luminosa e ordinata possibile: vengono rimossi tutti i detriti fino a liberare un’area di circa 1 m2. Arriva ora il momento di costruire e procedere con l’inserimento di centinaia di bastoncini e ramoscelli lasciati cadere sul terreno e debitamente calpestati in maniera apparentemente casuale su quella che in seguito sarà sede del pergolato. La forma precisa di rito di corteggiamento del maschio può essere molto diversa tra differenti popolazioni. Il tipo di struttura, in base a impostazione e resistenza, si rifà alle linee guida dei nidi dei passeracei. Un’ulteriore fase della costruzione prevede la raccolta di bastoncini lunghi 20-30 cm che vengono disposti verticalmente (utilizzo atipico da parte degli uccelli); gli eventuali ramoscelli curvati vengono posizionati in maniera tale da creare una struttura “di copertura” arcuata sul viale che sta prendendo forma; quest’ultimo ha tendenzialmente una lunghezza che va dai 37,5 cm ai 62,5 cm (Madden 2003). L'uccello giardiniere manipola accuratamente l'ambiente in modo da farlo divenire oggetto di selezione sessuale, come se fosse parte integrante del suo corpo;
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