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Boraginaceae - Boragineae Boraginaceae Boraginaceae - Boragineae Lorenzo Cecchi & Federico Selvi FLORA CRITICA Fondazione per la Flora Italiana D’ITALIA Boraginaceae - Boragineae Boraginaceae Boraginaceae - Boragineae Lorenzo Cecchi & Federico Selvi F F I, F FLORA CRITICA D’ITALIA Edito dalla Fondazione per la Flora Italiana con il supporto della Società Botanica Italiana e il contributo della Fondazione Internazionale pro Herbario Mediterraneo Comitato Editoriale Lorenzo Peruzzi (Pisa) (Coordinatore), Gianniantonio Domina (Palermo) (Segretario), Lorenzo Cecchi (Firenze), Giovanni Cristofolini (Bologna), Werner Greuter (Palermo), Enio Nardi (Firenze), Francesco M. Raimondo (Palermo), Federico Selvi (Firenze), Angelo Troìa (Palermo) Boraginaceae ‐ Boragineae Versione 1.0, pubblicata online il 03.08.2017 Lorenzo Cecchi 1 & Federico Selvi 2 1 Università degli Studi di Firenze, Museo di Storia Naturale, Sezione Botanica “Filippo Parlatore”, via G. La Pira 4, 50121 Firenze, Italia. 2 Università degli Studi di Firenze, Dipartimento di Scienze delle Produzioni Agroalimentari e dell'Ambiente (DISPAA), Laboratorio di Botanica, piazzale delle Cascine 28, 50144 Firenze, Italia FONDAZIONE PER LA FLORA ITALIANA Fondazione per la Flora Italiana Atto costitutivo: Palermo, Orto Botanico, 28/09/2007 Primo Consiglio di Amministrazione (2009 ‐ 2012): Carlo Blasi, Donato Chiatante, Bruno Corrias, Giovanni Cristofolini e Francesco Maria Raimondo Attuale Consiglio di Amministrazione: Carlo Blasi, Donato Chiatante, Giovanni Cristofolini, Enio Nardi, Lorenzo Peruzzi, Francesco Maria Raimondo, Maria Consolata Siniscalco. © 2017 Fondazione per la Flora Italiana, tutti i diritti riservati Composizione Grafica Gianniantonio Domina, Lorenzo Cecchi & Angelo Troia Immagine di copertina Symphytum officinale, disegno di L. Cecchi Pubblicato online il 03.08.2017 su http://www.floraditalia.it DOI: 10.17773/Fl_Ita_Boragineae1.0 Boraginaceae ‐ Boragineae 5 ______________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________ Boraginaceae Juss. BORAGINACEE Boraginaceae Juss., Gen. Pl.: 128. 1789. nom. cons., non Adans., Fam. Pl. 2: 173. 1763, (‘Borragines’) nom. illeg. Tipo: Borago L. (=) Anchusaceae Vest, Anleit. Stud. Bot.: 274, 302. 1818 (‘Anchusoideae’). Tipo: Anchusa L. (=) Buglossaceae Hoffmanns. & Link, Fl. Portug. 1: 63. 1809 (‘Buglossinae’). Tipo: Buglossum Mill. (=) Cerinthaceae Martynov, Tekhno‐Bot. Slovar: 120. 1820 (‘Cerinthoides’); Bercht. & J.Presl, Přir. Rostlin: 244. 1820. Tipo: Cerinthe L. (=) Onosmaceae Martinov in Tekhno‐Bot. Slovar: 437. 1820 (‘Onosmoides’). Tipo: Onosma L. (=) Echiaceae Raf., Fl. Tellur. 2: 61. 1837 (‘Echidia’). Tipo: Echium L. (=) Cynoglossaceae Döll, Rhein. Fl.: 406. 1843. Tipo: Cynoglossum L. BIBLIOGRAFIA LÅNGSTRÖM & CHASE 2002, HILGER & al. 2004 [Boragineae], 2005, THOMAS & al. 2008 [Lithospermeae], CECCHI & SELVI 2009 [Lithospermeae], WEIGEND & al. 2009, 2010a [Lithospermeae], NAZAIRE & HUFFORD 2012, WEIGEND & al. 2013, 2014, CHACÓN & al. 2016, Weigend & al. in KUBITZKI & al. 2016. DESCRIZIONE Erbe annue, bienni o perenni, più raramente arbusti o piccoli alberi (nelle regioni subtropicali), spesso diffusamente ricoperte da tubercoli o tricomi, non aromatiche. Foglie spiralate, raramente opposte (Asperugo), senza stipole. Infiorescenze terminali cimose, a maturazione progressiva, tipicamente scorpioidi, di rado corimboso‐panicolate. Fiori ermafroditi, tetraciclici e generalmente pentameri, talora (Anchusella) con riduzione del numero degli stami. Calice generalmente gamosepalo attinomorfo, dialisepalo solo in Cerinthe, con sepali ineguali inseriti a livelli diversi lungo l’asse fiorale. Corolla gamopetala, spesso attinomorfa, più raramente zigomorfa (nella flora italiana nei generi Anchusella, Echium L., Lycopsis e, in parte, Cerinthe L.); invaginazioni della corolla (pliche o “squame”) e peli sono spesso presenti alla fauce, con probabile funzione di organi per la presentazione secondaria del polline. Stami inseriti sul tubo corollino. Stilo lineare‐subulato, ginobasico, cioè inserito alla base del gineceo, sulla stessa superficie contigua al ricettacolo (ginobase) sulla quale sono ancorate le distinte porzioni dell’ovario; stigma clavato, capitato, bilobo o tetralobo (non in Italia: Arnebia Forssk.). Ovario supero, bicarpellare, generalmente tetraloculare e tetraspermo, nettamente di‐ o tetralobato; frutto eremocarpico, cioè suddiviso in 2‐4 nucule di‐ o monosperme (mericarpi) separate dalla ginobase e recanti una chiara cicatrice in corrispondenza della superficie di attacco. Germinazione epigea. NUMERI Nonostante il numero cromosomico sia noto ad oggi per non più del 35% dei CROMOSOMICI taxa descritti per la famiglia, e per specie e sottospecie in larga maggioranza ______________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________ 6 Flora Critica d’Italia ______________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________ europee e nordamericane, la gamma di variazioni del complemento cromosomico risulta assai ampia (Weigend & al. in KUBITZKI & al. 2016). I numeri cromosomici di base più comuni sono x = 6, 7 e 8; più raramente si riscontrano valori di x = 4, 9 o 10, o altri ritenuti di origine secondaria: x = 11, 12, 13, 14, 15, 17 e 19. Frequente è la formazione di complessi poliploidi, sia all’interno di generi che, più raramente, all’interno della stessa specie, che in tal modo si trova a includere diversi citotipi; altrettanto ben documentata è la tendenza alla disploidia discendente o la presenza di B‐cromosomi. Tra le specie italiane, il numero cromosomico più basso, 2n = 12, è condiviso da alcune specie diploidi di Onosma L., mentre quello più elevato, 2n = 144, è presente in alcuni citotipi di Symphytum tuberosum, interpretabile sia come dodecaploide in base 12 (GRAU 1968b, 1971) che come 18‐ploide in base 8 (MURÍN & MÁJOVSKI 1982). BIOLOGIA Piante quasi esclusivamente entomofile, di rado ornitofile (alcune specie di RIPRODUTTIVA Lithospermum L. americane) o cleistogame (alcune popolazioni di Neatostema apulum (L.) I.M.Johnst., ma carattere non fissato geneticamente; oss. pers.); prevale l’allogamia, spesso associata a polimorfismo stilare (ercogamia); sono altresì molto diffusi anche i meccanismi di autofecondazione. Dispersione dei semi principalmente per esozoocoria, grazie alla presenza di strofioli lipidici alla base dei mericarpi (“elaiosomi”) attrattivi per le formiche (mirmecocoria: la maggior parte delle Boragineae), o di appendici uncinate (glochidie) atte ad aderire al vello dei mammiferi o al piumaggio degli uccelli e variamente distribuite sulla superficie dei mericarpi o su quella esterna del calice caduco (nei generi di Boragineae sudamericane Moritzia DC. ex Meisn. e Thaumatocaryon Baill. e nella maggior parte delle Cynoglosseae); più raramente si ha dispersione per anemo‐ o idrocoria. DISTRIBUZIONE Famiglia che annovera circa 85 generi e 1600‐1700 specie, a distribuzione pressochè cosmopolita, ma con ampie lacune distributive nelle regioni a clima equatoriale e tropicale umido; i suoi principali centri di diversità specifica si trovano nelle aree a clima sub‐tropicale o temperato arido, cioè nelle regioni meridionali del regno olartico (sottoregno tetidico, regioni mediterranea e irano‐turanica) e nel regno capense, ma importanti aree di più recente differenziazione si hanno anche in Nord‐ e Sudamerica, Australia e Africa tropicale. Si contano in Italia 31 generi, 26 dei quali con almeno una specie autoctona. NOTE SISTEMATICA In contrasto con APG (2009, 2016), ma in linea con l’aggiornamento già acquisito sull’APG website (STEVENS 2016) e con le più recenti sinossi edite dal Boraginales Working Group (boraginales.myspecies.info/; CHACÓN & al. 2016, LUEBERT & al. 2016), la famiglia è qui trattata in senso stretto, separatamente da altri gruppi di pari rango nell’ambito dell’ordine Boraginales. Si tratta di una scelta diversa da quella cautelativamente adottata nella prima sinossi tassonomica per questa Flora (CECCHI & SELVI 2014), indotta dalla sempre più approfondita conoscenza della filogenesi dell’ordine Boraginales Juss. ex Bercht. & J. Presl (WEIGEND & al. 2014). Secondo la maggior parte degli specialisti del gruppo, il sistema di più famiglie è infatti l’interpretazione più appropriata e coerente col grado di effettiva separazione evolutiva tra i diversi gruppi (Fig. 1). Le Boraginaceae vengono a loro volta suddivise in tre linee monofiletiche fondamentali, corrispondenti alle sottofamiglie Echiochiloideae Weigend, Boraginoideae e Cynoglossoideae Weigend (con 8 tribù che da sole comprendono oltre l’80% delle specie note; CHACÓN & al. 2016; Fig. 2); nella flora italiana si trovano rappresentanti di 7
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