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The Cuticle Micromorphology of Extant and Fossil The cuticle micromorphology of extant and fossil plants as indicator of environmental conditions : A pioneer study on the influence of volcanic gases on the cuticle structure in extant plants Antonello Bartiromo To cite this version: Antonello Bartiromo. The cuticle micromorphology of extant and fossil plants as indicator of environ- mental conditions : A pioneer study on the influence of volcanic gases on the cuticle structure inextant plants. Paleontology. Université Claude Bernard - Lyon I, 2012. English. NNT : 2012LYO10017. tel-00865651 HAL Id: tel-00865651 https://tel.archives-ouvertes.fr/tel-00865651 Submitted on 24 Sep 2013 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Università degli Studi di Napoli “Federico II” Scuola di Dottorato in Scienze della Terra Dottorato di Ricerca in Analisi dei Sistemi Ambientali “XXIV Ciclo” Tesi preparata in cotutela con: Université Claude Bernard Lyon 1 Ecole doctorale E2M2 Evolution Ecosystèmes Microbiologie Modélisation Thèse de Doctorat en Paléonvironnements et Évolution Thèse en Biologie et Science de la Terre The cuticle micromorphology of extant and fossil plants as indicator of environmental conditions. A pioneer study on the influence of volcanic gases on the cuticle structure in extant plants Bartiromo Antonello 2011 RELATORI/DIRECTEURS DE RECHERCHE: Dott.ssa Guerriero Giulia Maître de conférences Guignard Gaëtan COORDINATORE DEL DOTTORATO: Prof. Barattolo Filippo CORRELATORE: Dott.ssa Barone Lumaga Maria Rosaria COMMISSIONE/JURY: Raschi Antonio, Prat Daniel, Guignard Gaëtan, Guida Marco DATA D’ESAME/DATE DE SOUTENANCE: 14/02/2012 The cuticle micromorphology of extant and fossil plants as indicator of environmental conditions. A pioneer study on the influence of volcanic gases on the cuticle structure in extant plants Antonello Bartiromo [email protected] Il nostro destino è inestricabilmente legato a quello di tutte le specie e di tutti gli ecosistemi della Terra. Niles Eldredge, 2000 Alla mia Terra, che non merita tutto questo … The cuticle micromorphology of extant and fossil plants as indicator of environmental conditions. A pioneer study on the influence of volcanic gases on the cuticle structure in extant plants INDEX Riassunto 7 Résumé 9 Abstract 11 Keywords 13 Laboratories 13 General Introduction 14 CHAP. I Influence of volcanic gases on the epidermis of Pinus halepensis Mill. in Campi Flegrei, Southern Italy: A possible tool detecting 17 volcanism in present and past floras 1.1. Introduction 17 1.2. Material and methods 20 1.3. Results 24 1.3.1. Sulphur measures 24 1.3.2. Scanning electron microscopy observations 24 1.3.3. Transmission electron microscopy observations 27 1.4. Discussion 30 1.4.1. Environmental response of Pinus halepensis to volcanism 31 1.4.1.1. Epicuticular and epistomatal wax 31 Stomatal aperture 31 Volcanic toxic compounds and wax alterations 33 Other sulphur considerations 35 Wettability of leaf surface 36 Microenvironment 37 1.4.1.2. Cuticular membrane (CM) + cell wall (CW) 38 Pinus halepensis cuticle type 38 CM + CW thickness 38 CM + CW development 40 4 The cuticle micromorphology of extant and fossil plants as indicator of environmental conditions. A pioneer study on the influence of volcanic gases on the cuticle structure in extant plants CM and CW ageing 40 Calcium-oxalate deposits 41 1.4.2. Potential application for extant and fossil material 42 The cuticle micromorphology of in situ Erica arborea L. CHAP. II exposed to long-term volcanic gases in Phlegrean Fields, 46 Campania, Italy 2.1. Introduction 46 2.2. Material and methods 48 2.2.1. Plant material and sites description 48 2.2.2. Gas vent 49 2.2.3. SEM, TEM and EDS preparations 50 2.2.4. Gas concentration measurements in air and soil 51 2.2.5. Statistical analysis 52 2.3. Results 52 2.3.1. Energy diffractive X-ray analysis with SEM 52 2.3.2. Scanning electron microscopy observations 53 2.3.3. Trasmission electron microscope observations 55 2.4. Discussion 58 2.4.1. Chemical and SEM considerations 58 2.4.2. TEM considerations 60 CHAP. III An Early Cretaceous flora from Cusano Mutri, Benevento, 63 southern Italy 3.1. Introduction 63 3.2. Geological setting 63 3.2.1. Stratigraphy 65 3.3. Material and methods 68 3.4. Systematic palaeontology 69 3.5. Taphonomic and palaeoecological remarks 88 3.5.1. Taphonomy 88 5 The cuticle micromorphology of extant and fossil plants as indicator of environmental conditions. A pioneer study on the influence of volcanic gases on the cuticle structure in extant plants 3.5.2. Palaeoecology of the Cusano Mutri sedimentary basin 89 3.5.3. Xeromorphic adaptations of the plants 90 3.5.4. Palaeoclimate and floral comparison 91 CHAP. IV Plant remains from the Early Cretaceous Fossil-Lagerstätte of 93 Pietraroja, Southern Italy, Benevento 4.4.1. Introduction 93 4.2. Geological setting 94 4.3. Material and methods 96 4.4. Systematic Palaeontology 97 4.5. Taphonomic and palaeoecological implications 111 4.5.1. Taphonomy 111 4.5.2. Palaeoecology of the sedimentary basin 114 4.5.3. Palaeoclimate and comparison with other Albian florae 114 General Conclusions 117 Acknowledgments / Ringraziamenti 120 References 121 Photos 155 6 The cuticle micromorphology of extant and fossil plants as indicator of environmental conditions. A pioneer study on the influence of volcanic gases on the cuticle structure in extant plants RIASSUNTO Lo strato acellulare che ricopre le parti aeree delle piante vascolari superiori è chiamato cuticola. Quest‟ultima funge da barriera protettiva ed è un efficace rilevatore d‟inquinamento ambientale. Lo studio delle cuticole vegetali e in particolare degli apparati stomatici delle conifere, è estesamente utilizzato quale strumento di analisi ai fini della comprensione delle caratteristiche ecologiche e paleoecologiche. E‟ interessante notare che, sebbene siano numerosi gli studi inerenti le cuticole delle piante, poco o niente è stato fatto relativamente agli effetti prodotti sulle cuticole delle piante da parte dei gas vulcanici. La Campania, con la presenza di numerose località caratterizzate da emissioni di gas di origine vulcanica (Pisciarelli, Solfatara, complesso del Somma-Vesuvio, ecc.), consente di effettuare studi di questo tipo. L‟obiettivo di questa ricerca è quello di contribuire a individuare le potenzialità delle conifere e delle angiosperme (attuali e fossili) quali indicatori ecologici utili per la comprensione delle variazioni dei parametri ambientali. A tale scopo sono state compiute osservazioni macroscopiche e microscopiche di piante vascolari in relazione all‟influenza di fattori ambientali quali: aerosol vulcanici, intensità luminosa, disponibilità di acqua e salinità. Nel corso della ricerca sono state campionate numerose aree e sono state utilizzate apparecchiature quali: microscopio ottico, SEM, TEM e EDS. La statistica è stata utilizzata per l‟analisi delle caratteristiche micromorfologiche. Le osservazioni condotte su piante attuali hanno consentito di studiare, per la prima volta, gli effetti dei gas vulcanici sull‟ultrastruttura delle cuticole della conifera Pinus halepensis [pino d‟Aleppo; siti di raccolta: Pisciarelli (presenza di gas vulcanici) e Cigliano (assenza gas vulcanici)] e dell‟angiosperma Erica arborea [siti di raccolta: Solfatara di Pozzuoli e Pisciarelli (presenza di gas vulcanici per entrambi) e Cigliano (assenza gas vulcanici)]. Le osservazioni al TEM effettuate su cuticole di P. halepensis, influenzate e non da gas vulcanici, hanno evidenziato che lo spessore totale di CM (cuticola) + CW (parete primaria) non subisce sostanziali variazioni di spessore. In particolare, la cuticola degli aghi influenzati da gas vulcanici mostra (a forti ingrandimenti: TEM) un aumento dei depositi di ossalato di calcio e un riarrangiamento delle fibrille che si dispongono parallelamente alla superficie. Le osservazioni condotte al SEM e al TEM su aghi di P. halepensis attuali hanno permesso altresì di realizzare una chiave dicotomica che consente di identificare possibili alterazioni (dovute alla presenza di gas potenzialmente tossici come quelli vulcanici) anche in cuticole di pino sub-fossili o fossili. 7 The cuticle micromorphology of extant and fossil plants as indicator of environmental conditions. A pioneer study on the influence of volcanic gases on the cuticle structure in extant plants Le osservazioni condotte su E. arborea, hanno permesso di costatare che gli spessori totali delle cuticole, “fumigated and not fumigated”, sono diversi. Comunque, in presenza di gas vulcanici lo strato esterno A2 subisce un sensibile incremento di spessore. Quest‟ultimo aumenta quando la concentrazione di CO2 in atmosfera è elevata, mentre non subisce sostanziali variazioni quando la quantità di CO2 al suolo varia drasticamente. Ciò conferma che la cuticola è il mediatore principale negli scambi gassosi tra ambiente interno ed esterno. Per entrambe le specie attuali studiate non è stata riscontrata presenza di zolfo nella cuticola, nella parete cellulare o nel citoplasma. Ciò conferma l‟ipotesi che gli scambi gassosi avvengono
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