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Insights from Notothenioids (Teleostei, Perciformes)

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Insights from Notothenioids (Teleostei, Perciformes) UNIVERSITÀ DEGLI STUDI DI PADOVA Dipartimento di Biologia SCUOLA DI DOTTORATO DI RICERCA IN BIOSCIENZE E BIOTECNOLOGIE INDIRIZZO DI BIOLOGIA EVOLUZIONISTICA CICLO XXVII Life histories and evolution: insights from notothenioids (Teleostei, Perciformes) Direttore della Scuola: Ch.mo Prof. Giuseppe Zanotti Coordinatore d’indirizzo: Ch.mo Prof. Andrea Pilastro Supervisore: Ch.mo Prof. Maria. B. Rasotto Co-supervisori: Dott. Carlotta Mazzoldi Dott. Mario La Mesa Dottorando: Emilio Riginella RIASSUNTO Gli organismi presentano un’enorme diversità nelle caratteristiche del loro ciclo biologico, sia a livello intra- che interspecifico, mostrando differenze tra popolazioni che vivono in habitat diversi. Tale variabilità fenotipica nel ciclo biologico, è stata messa in relazione all’influenza dei fattori ambientali, e rappresenta il risultato della selezione naturale. Uno degli aspetti chiave dello studio dei processi che hanno portato all’evolversi di queste differenze è cercare di comprendere quali siano le variabili ambientali che influenzano la variabilità dei tratti del ciclo biologico e le relazioni che tra di loro intercorrono. Diversi studi hanno messo in luce come la temperatura e la disponibilità trofica sembrano essere i parametri che meglio spiegano l’andamento dei diversi tratti come i tassi di crescita, l’età alla maturità sessuale, la fecondità, l’investimento riproduttivo, le dimensioni delle uova e delle larve. I pesci del Sottordine Notothenioidea rappresentano un esempio unico di radiazione adattativa in ambiente marino, dominano le acque che circondano il continente antartico sia nel numero di specie, superando le 120 (47%), che in biomassa (90-95%). Durante la loro evoluzione, i nototenioidei hanno subito una profonda diversificazione di alcuni caratteri, adattandosi alle nicchie ecologiche disponibili. Le rigide condizioni dell’ambiente antartico hanno contribuito a differenziare i pesci antartici dalle altre specie ittiche, rallentandone i ritmi di crescita, modificando gli aspetti riproduttivi, lo sviluppo larvale e l’ecologia. I nototenioidei sono distribuiti esclusivamente nell’Oceano Meridionale e in aree sub-antartiche quali Sud America, Nuova Zelanda e le coste più a sud-est dell’Australia. È stata descritta una notevole variabilità dei tratti del ciclo biologico tra specie e tra popolazioni della stessa specie, legata alle diverse condizioni ambientali a cui sono sottoposti gli individui. Con queste premesse, pur essendo disponibili filogenesi recenti ed aggiornate dei pesci nototenioidei, non sono mai stati applicati metodi comparativi che, tenendo conto delle relazioni filogenetiche, permettano di valutare il peso dei diversi fattori ambientali sull’evoluzione delle diverse strategie riproduttive. L’obiettivo del progetto di dottorato è stato di indagare la variabilità di alcuni tratti del ciclo biologico di specie di nototenioidei in relazione alle variabili ambientali, quali la temperatura dell’acqua, la produttività primaria e la copertura del ghiaccio. Il metodo dei contrasti indipendenti, utilizzato per controllare per le relazioni filogenetiche, rappresenta un punto di forza poiché permette di eliminare la dipendenza dei dati ed evitare quindi possibili relazioni dovute alla storia evolutiva delle specie piuttosto che a reali adattamenti evolutivi. La ricerca ha previsto lo studio della biologia delle specie campionate e la raccolta di dati da letteratura, prendendo in esame lo sviluppo dei gameti, l’investimento gonadico, la fecondità e le dimensioni degli ovociti. I risvolti del presente lavoro sono interessanti sia da un punto di vista della biologia evoluzionistica, per il peculiare adattamento mostrato dai nototenioidei nel loro ambiente, che conservazionistico, poiché la conoscenza dei tratti riproduttivi e della loro risposta ai cambiamenti ambientali è fondamentale per una efficace gestione sostenibile delle specie sfruttate dalla pesca e per valutare la loro capacità di resilienza a superare i cambiamenti climatici in corso. Infatti, alcune specie tra i I nototenioidei, sono state soggette in passato a forte pressione di pesca che, in pochi anni, ha portato al collasso degli stock ittici. Nella presente ricerca sono stati studiati i tratti del ciclo biologico di 17 specie appartenenti a tutte le otto famiglie di nototenioidei (Papers I, II, III, IV, V e VI), fornendo informazioni nuove sulla biologia delle specie e rafforzando quelle disponibili in letteratura. La variabilità intraspecifica nei tratti del ciclo biologico è stata valutata in tre specie grazie alla disponibilità di campioni provenienti da popolazioni di aree diverse (Papers I, III e IV). I dati ottenuti dall’analisi delle singole specie, sono stati quindi utilizzati per il confronto interspecifico (Paper VI). I campioni disponibili sono stati raccolti durante due spedizioni antartiche a bordo della nave tedesca R/V Polarstern (ANT-XXVIII/4, 2012 e ANT-XXIX/9, 2014), lungo la Penisola Antartica e nel Mare di Weddell. Ulteriori campioni di quattro specie, provenienti da diverse aree (South Orkney, South Georgia, Burdwood Bank e versante ovest della Penisola Antartica) sono stati forniti da collaboratori di ricerca. I risultati ottenuti dallo studio sulle singole specie permettono di evidenziare delle caratteristiche comuni tra i nototenioidei. In tutte le specie analizzate, le femmine hanno presentato un ovario sincrono per gruppi in cui è possibile distinguere nettamente due classi dimensionali di ovociti. Le femmine sessualmente mature hanno mostrato ovociti in fase avanzata di vitellogenesi, che verranno deposti nella stagione riproduttiva in corso, e ovociti in stadio di alveoli corticali o previtellogenetici, che matureranno per l’evento riproduttivo dell’anno successivo. La presenza contemporanea di ovociti in stadio avanzato di vitellogenesi, insieme ad altri allo stadio di alveoli corticali, evidenzia un’ovogenesi di lunga durata, verosimilmente, superiore ad un anno. I testicoli sono stati descritti come di tipo lobulare unrestricted e, in individui sessualmente maturi, hanno mostrato la presenza di spermatozoi nei dotti e spermatogoni localizzati soprattutto nella parte distale. Inoltre, è emerso come il processo di maturazione degli spermatozoi sia completo prima dell'inizio del periodo di riproduzione. È possibile quindi affermare che anche la spermatogenesi, come l’ovogenesi, è un processo molto lungo e, in generale, la lunga e lenta gametogenesi osservata è probabilmente correlata alla bassa temperatura e all'alto investimento riproduttivo dei nototenioidei. Lo studio della biologia delle specie ha fornito nuove informazioni sulla fecondità (assoluta e relativa), sulle dimensioni degli ovociti e sull’indice gonadosomatico (IGS), informazioni poi utilizzate per le analisi comparative (Paper VI). La stima dell’età, attraverso la lettura degli otoliti, ha permesso di valutare il tasso di crescita, la relazione tra la taglia e l’età, l’età massima e quella alla maturità sessuale (Papers II, III, IV, V). I risultati seguono le attese, infatti, come descritto in specie di acque fredde, i nototenioidei mostrano lento accrescimento, raggiungimento tardivo della maturità sessuale (circa 50-87% rispetto alla lunghezza massima) e longevità media (fino a 24 anni). Dal confronto intraspecifico emerge come la variabilità osservata nei tratti del ciclo biologico, possa essere attribuita a fattori ambientali che agiscono su piccola scala, come ad esempio correnti fredde dovute allo scioglimento del ghiaccio e/o a caratteristiche biologiche tipiche della specie. Infatti, il tipo di uova (pelagiche o demerse), la presenza o meno di cure parentali, la distribuzione degli individui adulti e la mobilità della specie, II possono avere un ruolo chiave nell’influenzare l’areale di distribuzione e di dispersione di una specie. Il confronto tra le due popolazioni dell’icefish Chaenocephalus aceratus, una specie bentonica a bassa mobilità, con uova demersali e presenza di cure parentali, ha dimostrato una fecondità più alta nell’area con temperatura dell’acqua più elevata e che non risente delle correnti fredde che si originano nel Mare di Weddell. Non emergono, invece, differenze intraspecifiche nei tratti riproduttivi in Pleuragramma antarctica, una specie pelagica, con uova cryopelagiche (cioè deposte sotto lo strato di ghiaccio stagionale) e in Notothenia rossii, dove i lunghi tempi di incubazione delle uova pelagiche e della fase larvale, sembrano assicurare connettività tra le aree investigate. In generale le discrepanze emerse in termini di dimensioni degli ovociti e IGS, sono probabilmente legate a differenze temporali nei periodi di campionamento. Le analisi comparative a livello interspecifico hanno evidenziato relazioni tra fattori ambientali e tratti del ciclo biologico. La taglia massima, a differenza degli altri tratti considerati, non ha presentato nessuna relazione con le variabili ambientali. Sembra quindi che la regola di Bergmann, che descrive la relazione positiva tra taglia massima e latitudine, con individui più grandi in ambienti più freddi (latitudini più alte), non sia applicabile alle alte latitudini. Nei nototenioidei è emersa una tendenza significativa a produrre ovociti più grandi al diminuire della produttività primaria, che raggiunge i valori minimi procedendo verso il Polo, sebbene presenti variabilità stagionale. A livello interspecifico, quindi, si osservano ovociti di maggiori dimensioni in specie che occupano alte latitudini, a conferma di quanto precedentemente previsto dai modelli teorici generali
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