Oceanic Island forests buried by Holocene (Meghalayan) explosive eruptions: palaeobiodiversity in pre-anthropic volcanic charcoal from Faial Island (Azores, Portugal) and its palaeoecological implications

UNCORRECTED PROOF Re iew o Palaeobo any and Palynology xxx (xxxx) xxx-xxx Con en s lis s a ailable a ScienceDi ec Re iew o Palaeobo any and Palynology jou nal homepage: www.else ie .com Oceanic Island o es s bu ied by Holocene (Meghalayan) explosi e e up ions: Palaeobiodi e si y in p e-an h opic olcanic cha coal om Faial Island (Azo es, Po ugal) and i s palaeoecological implica ions C.A. Góis-Ma ques a, b, ⁎, J. Rubiales c, L. de Nascimen o d, M. Menezes de Sequei a b, e, J.M. Fe nández-Palacios d, J. Madei a a aDepa amen o de Geologia, Faculdade de Ciências da Uni e sidade de Lisboa and Ins i u o Dom Luiz (IDL), Labo a ó io Associado, Uni e sidade de Lisboa, Campo G ande, 1749-016 Lisboa, Po ugal bMadei a Bo anical G oup (GBM), Faculdade de Ciências da Vida, Uni e sidade da Madei a, Campus da Pen eada, 9000-390, Funchal, Po ugal cUnidad de Bo ánica, Depa amen o de Sis emas y Recu sos Na u ales, Escuela Técnica Supe io de Ingenie ía de Mon es, Fo es al y del Medio Na u al, Uni e sidad Poli écnica de Mad id, Ciudad Uni e si a ia, Mad id, Spain dIsland Ecology and Biogeog aphy G oup, Ins i u o Uni e si a io de En e medades T opicales y Salud Pública de Cana ias (IUETSPC), Uni e sidad de La Laguna (ULL), A da. As o ísico F ancisco Sánchez s/n, La Laguna 38200, Cana y Islands, Spain eCIBIO, Cen o de In es igação em Biodi e sidade e Recu sos Gené icos, InBIO Labo a ó io Associado, Pólo dos Aço es, 9501-801 Pon a Delgada, Azo es, Po ugal ARTICLE INFO A icle his o y: Recei ed 9 Augus 2019 Recei ed in e ised o m 25 Sep embe 2019 Accep ed 26 Sep embe 2019 A ailable online xxx Keywo ds: Maca onesia Azo es a chipelago Py oclas ic densi y cu en Cha coali ied wood Mummi ied wood Palaeo ege a ion ABSTRACT In Faial Island (Azo es A chipelago, No h A lan ic Ocean), cha coali ied and mummi ied wood ossils ha e been epo ed wi hin la e Holocene (Meghalayan) py oclas ic deposi s om he Caldei a Fo ma ion. Due o hei ecen age, a de ailed s udy con eys a snapsho in o Azo ean palaeophy odi e si y and palaeo ege a ion, ca. 7–5 cen u ies be o e he a i al o Po uguese se le s o he Azo es Islands. He e we p o ide he i s de- ailed ana omical and axonomical s udy o hese wood ossils. In o al, 41 samples we e collec ed om se en locali ies, mainly om a ~ 1200y BP ignimb i e. Field wo k e ealed au och honous and pa aau och honous assemblages, wi h ee unks in up igh posi ion. The ana omical s udy o he ossil woods esul ed in he iden i ica ion o Junipe us b e i olia,Lau us azo ica,My sine e usa,Mo ella aya,Picconia azo ica,P unus lusi anica subsp. azo ica, and Vaccinium cylind aceum. Two ossil assemblages a e compa able o he p o- posed po en ial na u al ege a ion (PNV) o he Azo es. Su p isingly, P. lusi anica subsp. azo ica was he second mos abundan ossil wood sugges ing ha his ee was mo e abundan in a ecen pas in Faial Is- land and p obably in he a chipelago. This is co obo a ed by his o ical accoun s, and i s mode n sca ci y was ce ainly an h opically d i en. Iden i ying Holocene plan mac o ossils a e essen ial o p ope ly econs uc oceanic islands e es ial palaeoecosys ems, especially whe e o es s wi h high pe cen age o en omophilous axa a e unde ep esen ed in palaeopalynological limnic eco d. Fu he wo k is necessa y o econs uc Fa- ial Island and Azo es a chipelago palaeo ege a ion which is essen ial o p o ide an ecosys em base-line o es o a ion and managemen . © 2018. 1. In oduc ion The in e ac ion be ween explosi e o e usi e olcanic e en s and ege a ion may p o ide ideal aphonomical en i onmen s o plan s o be bu ied and p ese ed as cha coals o mummi ica ions (e.g. F i z, 1986; Spice , 1989; Hudspi h e al., 2010; Sco , 2010; Mus oe, 2018). Rapid bu ial o unks and s ems by ho olcanic ma e ials can p o- mo e hei py olysis esul ing in a o al o pa ial cha coali ica ion wi hou combus ion (e.g. Sco , 2010). The p oduc ob ained is cha - coal, a ai ly ine ma e ial ela i ely en iched in ca bon (e.g. Sco and Jones, 1991a; Sco and Jones, 1991b; Bi d, 2013). F om a ol- canological poin o iew, he deg ee o cha coali ica ion and e ⁎Co esponding au ho a : Depa amen o de Geologia, Faculdade de Ciências da Uni e sidade de Lisboa and Ins i u o Dom Luiz (IDL), Labo a ó io Associado, Uni e sidade de Lisboa, Campo G ande, 1749-016 Lisboa, Po ugal. Email add ess: [email p o ec ed] (C.A. Góis-Ma ques) lec ance can p o ide insigh s on he palaeo empe a u e o he ol- canic e en ha bu ied he wood (e.g. Sco and Jones, 1991b; Sco and Glasspool, 2005; Hudspi h e al., 2010). Mummi ied wood, i.e. al- mos unal e ed wood wi h p ese a ion o he o iginal ma e (Mus oe, 2018), can be p ese ed wi hin cold ph ea omagma ic deposi s (e.g. Góis-Ma ques e al., in p ess). Finally, cha coal and mummi ied wood ound associa ed wi h olcanic deposi s ha e been used wo ldwide o da e la e Qua e na y olcanic e up ions (e.g. B yson e al., 2006). F om a palaeobo anical pe spec i e, ossil cha coal p ese es ana omical cha ac e s ha can allow he iden i ica ion o gene a o species (e.g. Leny and Cas eel, 1975; Hudspi h e al., 2010; Sco , 2010; Mus oe, 2018), being di ec e idence o he e es ial palaeo- biodi e si y and palaeoecology o a gi en e i o y a a speci ic poin in ime. In he Maca onesian a chipelagos (Azo es, Madei a, Cana ies and Cabo Ve de), cha coali ied wood ossils ha e been p ima ily used o adioca bon da ing, being ei he ound wi hin limnic sedi h ps://doi.o g/10.1016/j. e palbo.2019.104116 0034-6667/ © 2018. UNCORRECTED PROOF 2 Re iew o Palaeobo any and Palynology xxx (xxxx) xxx-xxx men s (e.g. Rull e al., 2017b; Cas illa-Bel án e al., 2019), wi hin o below olcanic deposi s (e.g. Madei a e al., 1995; Geldmache e al., 2000; Ca acedo e al., 2007), o o in e Holocene palaeo i e egimes h ough mic ocha coal concen a ion in limnic sedimen a y co es (e.g. Conno e al., 2012; Nogué e al., 2013; de Nascimen o e al., 2016). Taxonomical s udies o cha coal ossils in Maca onesia a e almos la gely absen , despi e hei po en ial o in o m pas di e si y and ecology. Recen ly, some s udies illus a e ana omically p ese ed wood (e.g. Pensa e al., 2015), bu wi hou a axonomic iden i ica ion. In he Cana ies, se e al wood ossils ha e been epo ed (Schmincke, 1967; Nogales and Schmincke, 1969; Ga cía-Tala e a e al., 1995; Rubiales e al., 2013), bu only a ossil cha coal was iden i ied om he Miocene o G an Cana ia as Pinus cana iensis Swee ex Sp eng., (Ande son e al., 2009). An h acological s udies om a chaeological si es a e known o Maca onesia (e.g. Machado Yanes e al., 1997; Machado Yanes, 2007; Huebe , 2015; As udillo, 2018). Since he 19 h cen u y, se e al na u alis s and geologis s (e.g. Webs e , 1821; Mouzinho de Albuque que and Menezes, 1826; Ha ung, 1860; Fouqué, 1873), epo ed he p esence o mummi ied and cha coali ied ee unks imbedded in eph a deposi s o almos all islands o he Azo es a chipelago (Góis-Ma ques and Menezes de Sequei a, 2015; Góis-Ma ques e al., in p ess). Since hen, and wi h he ad en o 14C adioca bon da ing, cha coal and wood ossils ha e been ac i ely collec ed o da ing olcanic e en s in he Azo ean is- lands (e.g. Sho on e al., 1968; Madei a e al., 1995; Madei a e al., 1998; Pacheco, 2015; Con e e al., 2019). Howe e , so a , a palaeob- o anical s udy o hese mac o ossils was ne e accomplished. The Azo es Islands we e se led by he Po uguese in he mid-15 h cen u y, and he ea lies desc ip ions o he Azo ean Islands (e.g. Fe nandes, 1508; F u uoso, 1590a; F u uoso, 1590b; F u uoso, 1590c) accoun o he p esence o o es s composed o lau els and coni e s, including a p obable a bo eal ex inc ‘Mal a’(Mal aceae?) in São Miguel (F u uoso, 1590a; Dias, 2007). I is calcula ed ha he clea - ance and explo a ion o Azo ean o es s o almos 600yea s by Po - uguese se le s le less han 5% o na i e o es s (T ian is e al., 2010) al hough his alue does no co espond o a p is ine ege a ion. So a , he e o s o de ine he na u al ege a ion o Azo es Islands ha e been based on his o ical desc ip ions (e.g. Dias, 2007), s udy o he su i ing na i e ege a ion (e.g. Dias e al., 2005; Elias e al., 2016), and in e ed om palaeopalynological analysis o lakes and pea lands ( an Leeuwen e al., 2005; Conno e al., 2012; Rull e al., 2017a; Rull e al., 2017b). Plan mac o ossils, such as cha coal, lea es im- p essions o adp essions which a e bu ied wi hin o below py oclas- ic deposi s, can p o ide impo an in o ma ion on pas di e si y and ecology (e.g. Bu nham and Spice , 1986; Bu nham, 1994). Howe e , he s udy o Azo ean plan mac o ossils was seldom used in his dis- cussion (Góis-Ma ques e al., in p ess; Góis-Ma ques e al., 2019a). Mo eo e , qua e na y mic o- and mac o ossils o plan s a e abundan in he Azo es and Maca onesia islands, and ha e al eady demons a ed he p esence o ex i pa ed o ex inc axa (e.g. de Nascimen o e al., 2009; Góis-Ma ques e al., 2019b). In Faial Island, plan ossils a e known since he beginning o he 20 h cen u y (Góis-Ma ques e al., in p ess). P e ious a emp s o iden i y mummi ied and cha coali ied ossil woods om he Azo es and Faial we e p o ided by Fo jaz e al. (1970). Howe e , no me hod- ology o ana omical easoning was gi en o suppo hose iden i ica- ions (Góis-Ma ques e al., in p ess). Cha coali ied and mummi ied woods ound in Faial Island ha e been used in adioca bon da ing (e.g. Madei a e al., 1995; Madei a, 1998; Pacheco, 2001; Pacheco, 2015), bu hei sys ema ic and palaeoecological alue ha e been o e looked un il now. Recen ield wo k in Faial Island p o ided p omising ma- e ials, whe e excep ionally p ese ed ossil woods (ei he mummi- ied o cha coali ied specimens) we e ound h oughou he Island (Góis-Ma ques e al., in p ess). In his pape we p o ide he i s ana omical and axonomical s udy o wood ossils o he Azo es a chipelago om cha coal samples col- lec ed wi hin he py oclas ic deposi s o he La e Holocene (Megha- layan) Caldei a Fo ma ion in Faial Island. Mo eo e , palaeophy odi- e si y and palaeo ege a ion da a a e compa ed o he Azo es Po en- ial Na u al Vege a ion (PNV) (Elias e al., 2016), and analysed in he ligh o he ac ual plan di e si y and his o ical da a. Finally, we dis- cuss he implica ions o he palaeobiodi e si y ound o he econ- s uc ion o Faial and Azo es palaeo ege a ion. 2. Geological se ing The Azo es a chipelago lies in he cen al No h A lan ic Ocean a he iple junc ion be ween he No h Ame ican, Eu asian and Nubian pla es (Fig. 1A), and is in e p e ed as he esul o his complex ec- onic se ing coupled wi h a ho spo (Gen e e al., 2003). The a ch- ipelago is composed o nine islands (wes e n g oup: Co o and Flo- es; cen al g oup: Faial, Pico, S. Jo ge and G aciosa; eas e n g oup: S. Miguel, S. Ma ia islands and Fo migas Isle s; Fig. 1B). Faial Island is 21km long and 14km wide, wi h a su ace a ea o 170km2and each- ing a maximum ele a ion o 1043m abo e sea le el (a.s.l.). Acco d- ing o Madei a (1998) he subae ial pa o he island is composed o wo cen al olcanoes and wo issu al olcanic zones (Fig. 1C): he Ribei inha shield olcano, mapped as Ribei inha Volcanic Complex, is he oldes edi ice and spans in age om 850 o 360ka (Fe aud e al., 1980; Hildenb and e al., 2012); he Caldei a Volcano, a s a o ol- cano s addling he wes e n lank o Ribei inha olcano is di ided in o a lowe uni , he Ced os Volcanic Complex (130–115ka; Hildenb and e al., 2012), ep esen ing a dominan ly e usi e basal ic phase, and an uppe achy ic uni whose p oduc s a e included in he Caldei a Fo ma ion (< 16ka BP; Madei a e al., 1995; Pacheco, 2001). The is- su al olcanic sys ems a e ep esen ed by he Almoxa i e Fo ma ion (< 46ka; Hildenb and e al., 2012) in he eas and he Capelo Volcanic Complex (~ 8ka, Di Chia a e al., 2014) in he wes , wi h he la es documen ed e up ion in 1957–1958 (Capelinhos e up ion). The Caldei a Fo ma ion (o he Uppe G oup o Ced os Volcanic Complex sensu Pacheco, 2001) has a geological eco d o 12 sub-plin- ian and ph ea omagma ic achy e e up ions younge han 16ka BP (Madei a e al., 1995; Pacheco, 2001; Pacheco, 2015), whose de- posi s p ese ed lea imp essions/adp essions, cha coali ied and mum- mi ied wood, and palaeosols (e.g. Madei a e al., 1995; Pacheco, 2001; Góis-Ma ques e al., in p ess). One o he mos exp essi e olcanic e up ions o he Caldei a Vol- cano was he sub-plinian C11 e en (Pacheco, 2001; Pimen el e al., 2015), 14C da ed a ca. 1200 o 1000y BP (Madei a e al., 1995; Pacheco, 2015), ha lead o he o ma ion o a calde a in he sum- mi o Faial Island (Fig. 1C). Pimen el e al. (2015) econs uc ed he C11 e en h ough a de ailed s a ig aphic s udy o he py oclas ic deposi s, di iding i in o h ee membe s, ep esen ed by dis inc a- cies: (i) he B ejo Membe co esponding o he ini ial ph ea omag- ma ic phase, mainly composed o ash u s co e ing a palaeosol wi h associa ed ossili e ous laye s (see Góis-Ma ques e al., in p ess); (ii) he In e no Membe ep esen ing he sub-Plinian phase, p oduc- ing coa se pumice all deposi s; and (iii) he Ced os Membe ep e- sen ing he inal phase, wi h he collapsing o he sub-plinian e up- i e column o e he no h and eas e n sec o s o he island, o ming py oclas ic densi y cu en s (PDC) ha engul ed Faial's palaeo eg- e a ion. The high empe a u e o he PDC and he consequen ign- imb i e deposi s, p oduced he py olysis o he bu ied ege a ion, p e- se ing hem as cha coal (Góis-Ma ques e al., in p ess). Two o he si es, sampled nea he Caldei a im (no 6 and 7 in Fig. 1C), a e ph ea omagma ic b eccias om p e ious explosi e e up ions, mos p obably om he C7 o C8 e en s (see Pacheco, 2001; UNCORRECTED PROOF Re iew o Palaeobo any and Palynology xxx (xxxx) xxx-xxx 3 Fig. 1. Azo es a chipelago and Faial Island geological se ing. A, Loca ion and ec onic se ing o Azo es a chipelago; B, Azo es a chipelago ba hyme y (Digi al ele a ion model gene a ed in GeoMapapp (h p://www.geomapapp.o g/); Ryan e al., 2009), wi h he oldes known subae ial ages o each island; C, Simpli ied geological map o Faial Island, adap ed om Madei a and B um da Sil ei a (2003), showing he si es sampled (black s a s): 1. Fajã da P aia do No e beach; 2and 3. Sou h o Cascalho de Cima; 4. Cancelas ash qua y; 5. Cabouco Velho; 6and 7. Wes and Eas Calde a im. Pacheco, 2015), ha con ain mummi ied ee unks (e.g. Madei a e al., 1995; Góis-Ma ques e al., in p ess). 3. The ege a ion o Faial Island The wo d ‘Faial’is he Po uguese name o a o es domina ed by Mo ella aya (Ai on) Wilbu , a bo anical and ege a ional opon- omy ha di ec ly e lec s he ege a ion encoun e ed by he 15 h cen- u y Po uguese se le s in he island. The i s w i en imp ession o Faial Island ege a ion was gi en by F u uoso (1590b), bo n in Pon a Delgada (São Miguel Island) ca. 1522, meaning ha hese a e di ec obse a ions, by him o o ally ansmi ed om he i s se le s. He s a es ha hal o he island was co e ed by o es s and desc ibes he p esence o endemic sh ubs and ees s ill ound oday in he is- land (e.g. Vaccinium cylind aceum Sm. and Lau us azo ica (Seub.) F anco). The Azo es a chipelago, including Faial's ege a ion has been de- sc ibed by se e al au ho s (Seube and Hochs e e , 1843; D oue , 1866; T elease, 1897; Guppy, 1914; Tu in, 1953; Danse eau, 1970; Ri as-Ma ínez e al., 2001; Ri as-Ma ínez e al., 2002; Dias e al., 2005; Cos a e al., 2012; Fe nández-P ie o e al., 2012). Recen ly, Elias e al. (2016) published a po en ial na u al ege a ion model (PNV) whe e eigh ege a ion bel s we e p oposed o he Azo ean a chipelago. Fo Faial Island, Elias e al. (2016) p opose he occu ence o i e a i udinal ege a ion bel s, om sea le el o moun ain op: 1.E ica-Mo ella coas al woodlands (be ween 0 and 100ma.s.l.), domina ed by E ica azo ica Hochs . and M. aya;2.Pic- conia–Mo ella low-land o es (be ween 100 and 300m a.s.l) domi- na ed by Picconia azo ica (Tu in) Knobl. and M. aya.; 3.Lau us sub- mon ane o es (be ween 300 and 600ma.s.l.), domina ed by L. azo - ica, wi h he co-occu ence o P. azo ica and M. aya on he lowe pa and Junipe us b e i olia (Seub.) An oine and Ilex pe ado Ai on subsp. azo ica (Loes.) in he uppe pa ; 4.Junipe us–Ilex mon ane o es (be ween 600 and 900ma.s.l.), domina ed by J. b e i olia and I. pe ado subsp. azo ica;5.Calluna–Junipe us al imon ane sc ub- land, domina ed by Calluna ulga is (L.) Hull and J. b e i olia (abo e 900ma.s.l.) on moun ain idges. Cu en ly, he landscape in Faial is domina ed by pas u es, exo ic ee plan a ions (p edominan ly C yp ome ia japonica D. Don) and in- asi e species (e.g. Pi ospo um undula um Ven ., Hedychium ga d- ne ianum Sheppa d ex Ke Gawl., Hyd angea mac ophylla (Thunb.) Se .) (e.g. Ja dim e al., 2007). Faial's na i e ege a ion is p esen ly e- s ic ed o small pa ches on he Eas slope and inside he calde a, and coas al woodlands in he Sou h coas o he island (Dias e al., 2007), wi hin Faial Na u e Pa k. UNCORRECTED PROOF 4 Re iew o Palaeobo any and Palynology xxx (xxxx) xxx-xxx 4. Ma e ials and me hods Plan ossils we e collec ed om di e en pa s o Faial island (Góis-Ma ques e al., in p ess) in July o 2016. P e iously known lo- cali ies (e.g. Fo jaz, 1960; Fo jaz e al., 1970; Madei a e al., 1995; Madei a, 1998; Pimen el e al., 2015) we e p ospec ed o cha coal- i ied o mummi ied wood. Addi ionally, new locali ies such as oad cu s wi h isible cha coali ied unks we e also sampled. Cha coali- ied wood was s udied and iden i ied unde a e lec ed ligh mic o- scope (LM) (see Leny and Cas eel, 1975), and each sample s ud- ied in ans e se su ace (TS), adial longi udinal su ace (RLS) and angen ial longi udinal su ace (TLS) sec ions. These we e compa ed o he e e ence collec ion o Maca onesian woods a he ‘Uni e si- dad Poli écnica de Mad id’a las and keys dealing wi h Maca onesian wood ana omy (e.g. Pe aza and Lopez de Roma, 1967; Schweing ube e al., 2011; Fe ei a e al., 2012; Schweing ube e al., 2013), online da abases ‘InsideWood’(Wheele , 2011: h p://insidewood.lib.ncsu. edu/sea ch) and ‘The Xylem Da abase’(Schweing ube and Landol , 2010: h ps://www.wsl.ch/dend op o/xylemdb/). Fu he mo e, se- lec ed samples we e spli in o TS, RLS and TLS, moun ed on me al s ubs, gold spu e coa ed ( o p e en sample cha ging) and obse ed on a JEOL JSM-5200LV scanning elec on mic oscope (SEM) a he Mic oscopy Facili y o he Facul y o Sciences o he Uni e si y o Lisbon (FCUL-MF). Wood ana omy desc ip ions and measu emen s ollow he In e na ional Associa ion o Wood Ana omis s (IAWA) lis o mic oscopic ea u es o so wood (Baas e al., 2004) and ha dwood (Wheele e al., 1989). Sys ema ic axonomy ollows Menezes de Sequei a e al. (2012). Cha coal samples a e deposi ed in he palaeob- o anical collec ion o Madei a Uni e si y he ba ium (UMad-P). 5. Resul s 5.1. Field sampling Se en locali ies we e sampled o ossil woods, anging in ele a- ion om 20m o 977ma.s.l. (Fig. 1C). In o al, 41 samples we e col- lec ed (Table 1), 38 cha coali ied wood samples ela ed mainly o he 1200y BP C11 ignimb i e deposi (Madei a e al., 1995; Pimen el e al., 2015) and h ee mummi ied wood samples om he calde a im ela ed o he Caldei a Fo ma ion (< 16ka) ph ea omagma ic de- posi s. As numbe ed in Fig. 1C: 1. Fajã da P aia do No e beach: loca ed on he No h coas o he island a an ele a ion o ~ 22 o 30ma.s.l., a sec ion o he sea cli cu in he C11 Ced os Membe ig- nimb i e (Fig. 2B) supplied cha coali ied unks (Fig. 2E). The occu - ence o cha coal a ha loca ion was al eady men ioned by Fo jaz e al. (1970); 2and 3. Sou h o Cascalho de Cima: along oad cu s, a ele a ions be ween 190 and 280ma.s.l., he exposu es o he C11 ign- imb i e con ain cha coali ied unks (Fig. 2F); 4. Cancelas ash qua y: loca ed in he no heas pa o he island a 365ma.s.l.; his aban- doned qua y (Fig. 2A) exposes a palaeo o es bu ied by he C11 ig- nimb i e c ea ing a ossil o es wi h cha coali ied ee unks (Fig. 2C–D, G), some s ill in up igh posi ion and p ese ing he ba k; de- gassing pipes om py olysis o he wood a e common (Fig. 2C); 5. Cabouco Velho: loca ed in he no heas pa o he island a an el- e a ion o 510ma.s.l., his ignimb i e ou c op also supplied cha coal samples; locali ies 4 and 5 we e some o he si es o sampling o cha - coal o 14C da ing by Madei a e al. (1995) which yielded an a e - age adioca bon age o 1200y BP; 6and 7. Wes and Eas Caldei a im: in hese si es, close o he calde a im, mummi ied coni e ous wood, s ill p ese ing he a oma, we e en ombed by ph ea omagma ic explosion b eccias. The eas si e (Fig. 3C–E) ou c op, a an ele a- ion o 897ma.s.l., displays 4 b eccias sepa a ed by paleosols o e o sional uncon o mi ies; a wood sample om he palaeosol unde lying he op b eccia was adioca bon da ed a 1660±45y BP (ICEN-994), while he wes si e (Fig. 3A–B) co esponds o a small s eam alley, a an ele a ion o 977m, ha exposes a b eccia co e ing a palaeosol ich in unks, one o which was da ed a 1820 ± 45y BP (ICEN-972) (see Madei a e al., 1995 o u he de ails). 5.2. Sys ema ic palaeobo any The iden i ica ion o ana omical ea u es on cha coali ied wood p esen s some challenges (Falcon-Lang e al., 2012) due o he ho- mogeniza ion o cell walls and he sh inking o cellula s uc u es due o py olysis occu ing a empe a u es abo e 300–325°C (Sco , 2010 and e e ences he ein). Ana omical cha ac e s, easily desc ibed in ex an woods by di e en ial wall s aining o cell con en (e.g. ax- ial pa enchyma), a e no easily disce ned o e en possible o see in cha coals. Mo eo e , as poin ed ou by Falcon-Lang e al. (2012), he cha coali ica ion p ocess leads o opaci y o cellula walls, p e en ing he obse a ion o se e al ana omical cha ac e s. The desc ip ions p e- sen ed he e conside hese limi a ions and a e based on LM and SEM obse a ions. Di ision:Spe ma ophy a Willk., 1854 Class:Pinopsida Bu ne , 1835 O de :PINALES Go ozh., 1904 Family:CUPRESSACEAE G ay, 1821 Genus:Junipe us L., 1753 Junipe us b e i olia (Seub.) An oine, 1857 (TS: Figs. 4A–C; RLS: Figs. 4D–F; TLS: Figs. 4G–I) Ma e ial: Fai-21; Fai-28U; Fai-02A; Fai-02B; Fai-11. Desc ip ion in IAWA code (Baas e al., 2004): 3, 40, 42, 44, 55, 58, 59, 80, 87, 93, 96, 98, 102, 107. Desc ip ion:G ow h ings: so wood wi h dis inc g ow h ings bounda ies, ab up ansi ion be ween ea lywood and la ewood (Fig. 4A–B); T acheids: pi ing p edominan ly unise ia e (Fig. 4D, F), in- e cellula spaces p esen bu a e, la ewood acheid wi h hin-walls, acheid pi s wi h o us ex ensions and no ched bo de s (Fig. 4F); Ax- ial pa enchyma: indis inguishable and ans e se end walls no ob- se ed; Ray composi ion and size: acheids absen , ay pa enchyma wi h smoo h walls (Fig. 4E); ays unise ia e up o 4 cells (Fig. 4G–I), a e age ay heigh 63.8μm (min=26.98μm; max=101.36μm); C oss- ield pi ing: usually 2, cup essoid. Compa ison and ema ks: In he Azo es, wo na i e coni e s a e known, he endemic J. b e i olia and na i e Taxus bacca a L. (Menezes de Sequei a e al., 2012). Wood ana omy o he s udied cha coals a e compa able o he Maca onesian Junipe us desc ibed by de Palacios e al. (2014). I di e s om T. bacca a due o he absence o helical hickenings in acheid cells (e.g. C i ella o and Schweing ube , 2013). Acco ding o de Palacios e al. (2014), acheid pi s wi h o us ex ensions and no ches in he ou e bo de , di e en i- a e J. b e i olia om o he Maca onesian Junipe us species. Unde e- lec ed LM and SEM hese cha ac e s we e di icul o obse e. Mo e- o e , he wa y laye and he nodula ans e se end walls o axial pa enchyma cells we e also no obse ed,mos p obably due o ho- mogeniza ion du ing py olysis o he cell walls (Fig. 3C). Al hough cha coali ica ion p e en s he iden i ica ion o ana omical key cha ac- e s, his ossil wood mos p obably belongs o J. b e i olia. As a no e, his ee occu s in he Azo es om 0 o 1500ma.s.l. (Elias and Dias, 2014). This ecological ampli ude led o he ecogni ion o in aspeci ic axa by Elias and Dias (2014). The iden i ica ion o Azo ean in aspe- ci ic axa ho ough wood ana omy seems imp obable as he ecogni- ion o speci ic axa is al eady challenging (de Palacios e al., 2014 and e e ences he ein). Class: Magnoliopsida B ongn.,1843. O de : LAURALES Be ch old and P esl, 1820. UNCORRECTED PROOF Re iew o Palaeobo any and Palynology xxx (xxxx) xxx-xxx 5 Table 1 Lis o sampled specimens in Faial island p esen ed in his s udy wi h espec i e in o ma ion on loca ion, age and iden i ica ion. Field numbe P ese a ion Collec ion Locali y Coo dina es (WGS84) Al i ude e o Age (14C) Volcanos a ig aphy Iden i ica ion Pla e Fai-02A Mummi ied UMad-P Eas im o he calde a 38°34′54”N; 28° 42′05”W 897m 10m 1820±45y BP c7(?) o c8 (?) c . Junipe us b e i olia (Seub.) An oine Fai-02B Mummi ied UMad-P Eas im o he calde a 38°34′54”N; 28° 42′05”W 897m 10m 1820±45y BP c7(?) o c8 (?) c . Junipe us b e i olia (Seub.) An oine Fai-09 Mummi ied? UMad-P Fajã da P aia do No e beach 38°36′35”N; 28°45′24”W 22m 7m ~ 1200y BP c11 No s udied Fai-10A Cha coal UMad-P Fajã da P aia do No e beach 38°36′36”N; 28°45′18”W 30m 17m ~ 1200y BP c11 Lau us azo ica (Seub.) F anco Fai-10B Cha coal UMad-P Fajã da P aia do No e beach 38°36′36”N; 28°45′18”W 30m 17m ~ 1200y BP c11 Lau us azo ica (Seub.) F anco Fai-10C Cha coal UMad-P Fajã da P aia do No e beach 38°36′36”N; 28°45′18”W 30m 17m ~ 1200y BP c11 Lau us azo ica (Seub.) F anco Fai-10D Cha coal UMad-P Fajã da P aia do No e beach 38°36′36”N; 28°45′18”W 30m 17m ~ 1200y BP c11 Lau us azo ica (Seub.) F anco Fai-11 Mummi ied UMad-P Wes im o he calde a 38°35′06”N; 28°43′38”W 977m 5m 1660±45y BP c9 c . Junipe us b e i olia (Seub.) An oine Fai-21 Cha coal UMad-P Cabouco Velho 38°35′47”N; 28°40′54”W 511m 15m ~ 1200y BP c11 Junipe us b e i olia (Seub.) An oine Fai-28U Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 Junipe us b e i olia (Seub.) An oine Pla e I Fai-28A Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 Lau us azo ica (Seub.) F anco Fai-28C Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 Lau us azo ica (Seub.) F anco Fai-28D Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 Lau us azo ica (Seub.) F anco Pla e II Fai-28E Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 Lau us azo ica (Seub.) F anco Fai-28F Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 Lau us azo ica (Seub.) F anco Fai-28T Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 Lau us azo ica (Seub.) F anco Fai-28W Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 Lau us azo ica (Seub.) F anco Fai-28AA Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 Lau us azo ica (Seub.) F anco Fai-28M Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 Mo ella aya (Ai on) Wilbu Pla e III Fai-28J Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 My sine e usa Ai on Pla e V Fai-28B Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 Picconia azo ica (Tu in) Knobl. Fai-28H Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 Picconia azo ica (Tu in) Knobl. Fai-28I Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 Picconia azo ica (Tu in) Knobl. Fai-28K Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 Picconia azo ica (Tu in) Knobl. Fai-28L Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 Picconia azo ica (Tu in) Knobl. Fai-28R Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 Picconia azo ica (Tu in) Knobl. Fai-28S Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 Picconia azo ica (Tu in) Knobl. Pla e VII Fai-28G Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 P unus lusi anica L. Subsp. azo ica (Mouill.) F anco Pla e VI Fai-28O Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 P unus lusi anica L. subsp. azo ica (Mouill.) F anco Fai-28P Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 P unus lusi anica L. subsp. azo ica (Mouill.) F anco UNCORRECTED PROOF 6 Re iew o Palaeobo any and Palynology xxx (xxxx) xxx-xxx Table 1 (Con inued) Field numbe P ese a ion Collec ion Locali y Coo dina es (WGS84) Al i ude e o Age (14C) Volcanos a ig aphy Iden i ica ion Pla e Fai-28Q Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 P unus lusi anica L. subsp. azo ica (Mouill.) F anco Fai-28V Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 P unus lusi anica L. subsp. azo ica (Mouill.) F anco Fai-28X Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 P unus lusi anica L. subsp. azo ica (Mouill.) F anco Fai-28Y Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 P unus lusi anica L. subsp. azo ica (Mouill.) F anco Fai-28Z Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 P unus lusi anica L. subsp. azo ica (Mouill.) F anco Fai-28N Cha coal UMad-P Cancelas ash qua y 38°35′12”N; 28°38′53”W 365m 7m ~ 1200y BP c11 Vaccinium cylind aceum Sm. Pla e IV Fai-29A Cha coal UMad-P Sou h o Cascalho de Cima 38°37′57”N; 28°42′54”W 191m 5m ~ 1200y BP c11 P unus lusi anica L. subsp. azo ica (Mouill.) F anco Fai-29B Cha coal UMad-P Sou h o Cascalho de Cima 38°37′57”N; 28°42′54”W 191m 5m ~ 1200y BP c11 P unus lusi anica L. subsp. azo ica (Mouill.) F anco Fai-29C Cha coal UMad-P Sou h o Cascalho de Cima 38°37′57”N; 28°42′54”W 191m 5m ~ 1200y BP c11 P unus lusi anica L. subsp. azo ica (Mouill.) F anco Fai-30A Cha coal UMad-P Sou h o Cascalho de Cima 38°37′32”N; 28°42′49”W 280m 27m ~ 1200y BP c11 Lau us azo ica (Seub.) F anco Fai-30B Cha coal UMad-P Sou h o Cascalho de Cima 38°37′32”N; 28°42′49”W 280m 27m ~ 1200y BP c11 Lau us azo ica (Seub.) F anco Pla e II Family: LAURACEAE Juss., 1789. Lau us azo ica (Seub.) F anco, 1960 (TS: Figs. 5A–B; RLS: Figs. 5C–D; TLS: Figs. 5E–G) Ma e ial: Fai-10A, Fai-10B, Fai-10C, Fai-10D, Fai-28A, Fai-28C, Fai-28D, Fai-28E, Fai-28F, Fai-28T, Fai-28W, Fai-28AA, Fai-30A, Fai-30B. IAWA code (Wheele e al., 1989): 1, 4, 9, 10, 13, 22, 25, 31, 41, 49, 53, 61, 65, 69, 98, 106, 112, 115. Desc ip ion:G ow h ings: ha dwood wi h dis inc g ow h ings bounda ies (Fig. 5A); Vessels: wood semi- ing-po ous, essel g oup- ing soli a y o 2–4 essels, ou line essel ound o angula (Fig. 5A–B), mean angen ial diame e o essel lumina be ween 50 and 100μm (X=100.5μm; n=30), 40–100 essels pe mm2 (X=41.3 essels pe mm2); essel elemen leng h mean≤350μm (X=234μm; n=35); essels wi h simple pe o a ion pla es (Fig. 5F–G), in e essel pi s al e na e and small (4–7μm) (Fig. 5G); Ves- sel– ay pi s ounded o i egula ; Fibe s: ibe s simple and sep a e, anging om hin (in ea ly wood) o hick-walled in la ewood (e.g. Fig. 5D, F); Rays: ay wid h 4 cells, heigh <500μm (Fig. 5C–D), ays pa enchyma cells wi h small pi s (Fig. 5E), p ocumben wi h squa e ma ginal cells (Fig. 5H–I). Compa ison and ema ks: he only known na i e Lau aceae in he Azo es is he endemic L. azo ica (Menezes de Sequei a e al., 2012), al hough o he Maca onesian and Eu opean Lau aceae we e in o- duced in se e al Azo ean islands (e.g. Oco ea oe ens (Ai on) Baill., Lau us nobilis L.; see Sil a e al., 2005). Maca onesian Lau aceae wood ana omy is e y simila (excep o Apollonias ba bujana (Ca .) Bo nm.; see Lou y, 2009). The s udied samples a e compa able o L. azo ica wood ana omy (e.g. Pe aza and Lopez de Roma, 1967; Lou y, 2009; Schweing ube and Landol , 2010; Schweing ube e al., 2011; Wheele , 2011; Ma os e al., 2019), pa icula ly he es- sel angen ial lumina leng h and dis ibu ion and g ouping, he mean essel leng h, he p esence o simple pe o a ion pla es, and sep a e ibe s. In some specimens, la ewood ibe walls a e hick, wi hou isible lumina o sep a and o ally i i ied (e.g. Fig. 5D). Lou y (2009) s a es he p esence o scala i o m pe - o a ions in L. azo ica, howe e we did no obse e his cha ac e in ou samples. The p esence o ex inc o ex i pa ed Lau aceae in he Azo es, al hough imp obable, as he i s his o ical desc ip ions only men ion he p esence o L. azo ica (F u uoso, 1590b; F u uoso, 1590a; F u uoso, 1590c), is no impossible. Lea ossils p esumably belonging o Oco ea oe ens (Ai on) Baill. ha e been desc ibed o São Miguel (Diniz, 1962), al hough hei axonomical a ini ies s ill need o be e ised (see Góis-Ma ques e al., in p ess). Today, L. azo ica oc- cu s in all Azo ean islands (Sil a e al., 2010), being an impo an ee in he ex an Azo ean na u al o es s (e.g. Ja dim e al., 2007; Elias e al., 2016). O de : MYRICALES Rich., 1835. Family: MYRICACEAE Rich. ex Kun h, 1817. Mo ella aya (Ai on) Wilbu , 1994. (TS: Figs. 6A–C; TLS: Figs. 6D–H; RLS: Figs. 6I–K) Ma e ial: Fai-28M. IAWA code (Wheele e al., 1989): 1, 5, 7, 10, 11, 12, 14, 15, 20, 21, 24, 25, 31, 41, 49, 53?, 57, 63, 65, 69, 97, 106, 112. Wood desc ip ion:G ow h ings: ha dwood wi h dis inc g ow h ings bounda ies (Fig. 6A); Vessels: wood di use-po ous (Fig. 6A), adial pa e n, wi h clus e s common (Fig. 6A); ou line angula (Fig. A–C), pe o a ion pla es simple and scala i o m wi h ≤10 ba s (Fig. 6B–C), in e essel pi s scala i o m and al e na e (Fig. 6G–H), al- e na e pi s minu e o small; essel– ay pi s appa en ly simple and ounded; mean angen ial diame e o essel lumina be ween 50 and 100μm (X=57μm; n=32), anging om 38.08 o 70.72μm, 40–100 essels pe squa e millime e (X=47 essels/mm2; n=141); essel elemen up o 340μm (maximum heigh measu ed); scle o ic yloses p esen (Fig. 6A); Fibe s: ibe s pi s common in bo h a UNCORRECTED PROOF Re iew o Palaeobo any and Palynology xxx (xxxx) xxx-xxx 7 Fig. 2. Selec ed ou c ops and associa ed cha coals om Faial Island. A, Cancelas ash qua y ou c op; B, Fajã da P aia do No e beach ou c op; C, cha coal (s a ) om Cancelas ash qua y associa ed wi h degassing pipes (a ow; pocke kni e ~ 20cm); D, cha coal om Cancelas ash qua y showing he ba k a ached (coin diame e : 1,6cm); E, cha coali ied s em ound in Fajã da P aia do No e beach; F, Cha coali ied unk ound in Cascalho de Cima locali y; G, unks om Cancelas ash qua y (scale ba in cm). dial bu a e in angen ial walls, sep a e, hin- o hick-walled (Fig. 6E); Rays: ays wid h 1–3 cells, he e ocellula (Fig. 6D–E), heigh <1mm, ay cells pi ed (Fig. 6B, F and J), body ay cells p ocumben wi h 1–3 squa e ma ginal cells (Fig. 6I–J), pe o a ed ay cells a e, p esen ing scala i o m pe o a ions (Fig. 6K); Rays pe mm: 4–12/mm. Compa ison and ema ks: The wood ana omy o M. aya was s udied and illus a ed by a ious au ho s (e.g. Ca lquis , 2002; Schweing ube and Landol , 2010; Schweing ube e al., 2011; Wheele , 2011). Sample Fai-28M bes i s wi hin he wood ana omy o M. aya, especially o he p esence o essels wi h scala i o m pe o a ions wi h < 10 ba s. Ca lquis (2002) published a de ailed s udy on My icaceae wood ana omy. Mo ella aya is compa able o ou sample, showing simila mean angen ial diame e o essel lu- mina (54μm), compa able essels pe squa e millime e , al hough lowe (34 essels/mm2), numbe o ba s on essel pe o a ions simi- la (9 ba s), and simila ay ana omy and size. G ow h ings bound- a ies a e isible in ou sample, bu Ca lquis (2002) did no obse e hem in M. aya om Hawaii. Howe e , Schweing ube e al. (2011) and Pupo-Co eia (2005) illus a e specimens om Madei a Island wi h clea g ow h ings. M. aya is p esen in all Azo ean islands (Sil a e al., 2010). O de : ERICALES Be ch . & J. P esl Be ch old and P esl, 1820. Family: ERICACEAE Juss., 1789. Vaccinium cylind aceum Sm.,1819. (TS: Figs. 7A–C; TLS: Figs. 7D–H; RLS: Figs. 7I–K) Ma e ial: Fai-28N. IAWA code (Wheele e al., 1989): 1, 4, 6, 7, 11, 12, 14, 17, 20, 21, 26, 30, 36, 37, 40, 50, 53, 63, 65, 69, 98, 102, 103, 106, 107, 112. Wood desc ip ion:G ow h ings: ha dwood, dis inc g ow h ings bounda ies; Vessels: wood di use-po ous (Fig. 7A–B), essels in an- gen ial, diagonal and adial pa e n and clus e s common (Fig. 7A); essels ou line angula (Fig. 7B), pe o a ion pla es scala i o m wi h 20–40 ba s (Fig. 7G–H, 7K), in e essel pi s scala i o m o oppo- si e and small (7–10μm) (Fig. 7G); essel– ay pi s wi h dis inc bo - de s simila o in e essel pi s in size and shape h oughou he ay cell (Fig. 7C); essel elemen s wi h helical hickenings h ough he UNCORRECTED PROOF 8 Re iew o Palaeobo any and Palynology xxx (xxxx) xxx-xxx Fig. 3. Selec ed ou c ops and associa ed mummi ied wood om Faial Island. A, Mummi ied woods c . Junipe us b e i olia (a ows) ound in he wes pa o Calde a im, ca. ea ly 1990 (Pho o by José Madei a); B, same ou c op in 2016 s ill wi h mummi ied wood; C, ph ea omagma ic deposi s in he Eas pa o he Calde a im, wi h ou ph ea omagma ic b eccias and associa ed mummi ied wood (pho o by José Madei a ca. ea ly 1990); D, same ou c op in 2016; E, example o mummi ied woods ound below b eccia no 4 da ed a 1820±45y BP. body (Fig. 7E, G–H), mean angen ial diame e o essel lumina be- ween ≤50μm (X=26.31μm; n=30), anging om 18.36 o 38.42μm, ≥100 essels pe squa e millime e (X=254 es- sels/mm2; n=762); mean essel elemen leng h anging 350–800μm (X=573.7μm; n=11); Fibe s: ibe s pi s common in bo h adial and angen ial walls, sep a e, hin- o hick-walled; Rays: ays la ge han 4–10 se ia e, heigh >1mm al hough smalle ays occu (Fig. 7D, F); ay cells wi h pe o a ed pla es (Fig. 7F), body ay cells p ocumben wi h one o 2–4 ows o up igh ma ginal cells (Fig. 7I-K). Compa ison and ema ks: Fai-28N is he single sample ha p e- sen s essels wi h > 20 scala i o m pe o a ions. Due o simila ana omical cha ac e s, essel angen ial lumina size and simila (al- hough lowe ) essels/mm2, his specimen seems o co espond o Vaccinium L. The Azo ean endemic V. cylind aceum s ill lacks a o - mal ana omical desc ip ion. Howe e , he sample s udied bes i s he Madei an V. padi olium (=‘V. made ense’) in he “Xylem Da a Base” (Schweing ube and Landol , 2010). Phylogene ically, hese species a e sis e axa, clus e ing oge he wi hin he Vaccinium Andean clade (see K on e al., 2002), and mos p obably sha ing iden ical wood ana omy. The essels pa e n in TS, lumina angen ial essel size and essels/mm2app oxima e mo e o V. padi olium, essels wi h heli cal hickenings h ough he body and simple pi s h ough he ay cell walls. I di e s in he numbe o ba s in he scala i o m pe o a ion, ha ing a highe numbe o ba s (> 20) han p oposed o V. padi- olium (X=3 ba s, anging om 1 o 6; Lens e al., 2004). Howe e , V. padi olium illus a ed by Schweing ube and Landol (2010) shows scala i o m pe o a ion up o 15 ba s (Lens e al., 2004), and simi- la o he ones ound in Fai-28N. Ano he simila wood is Ilex pe - ado Ai on subsp. azo ica (Loes.) Tu in, al hough he angen ial essel leng h is highe (50–100μm) and essel/mm2is lowe (40–100 essel/ mm2) (Baas, 1973). Nowadays he endemic V. cylind aceum occu s in all Azo ean islands including in G aciosa Island whe e i was in o- duced (Sil a e al., 2010). O de : PRIMULALES Lindl., 1833. Family: MYRSINACEAE R. B ., 1810. My sine e usa Ai on, 1789. (TS: Figs. 8A–C; RLS: Figs. 8D–G; TLS: Figs. 8H–J) Specimen: Fai-28J. IAWA code (Wheele e al., 1989): 2, 5, 7, 9, 12, 13, 20, 22, 24, 31, 40, 48, 62, 63, 65, 69, 98, 102, 105, 110, 114. Wood desc ip ion: G ow h ings: ha dwood wi h bounda ies ague, p esen ing angen ial and adial auma ic canal spli s ( UNCORRECTED PROOF Re iew o Palaeobo any and Palynology xxx (xxxx) xxx-xxx 9 Fig. 4. Junipe us b e i olia (Seub.) An oine ana omy (Fai-28U) SEM pho og aphs in TS, TLS and RLS. A, Gene al TS iew showing he g ow h ing bounda ies (a ows); B, TS, de ail o ea ly wood o la e wood ansi ion, C, TS, de ail showing he homogeniza ion o he cell walls caused by cha coali ica ion abo e > 300°C; D, RLS gene al iew; E, RLS, c oss- ield pi s appa en ly cup essoid, wi h wo pe c oss ield, F, RLS, de ail o acheids showing unise ia e bo de ed pi s G, gene al TLS iew; H, TLS, showing unise ia e 1–4 ay essels, I, TLS, de ail showing a unise ia e 3 ay essel. Fig. 8A–B, D–E); Vessels: wood di use-po ous, a ely mainly soli- a y o a ely in adial mul iples o 2–4 (Fig. 8A–B), ou line angula (Fig. 8C), pe o a ion pla es simple, in e essel pi s minu e (≤4μm), scala i o m o al e na e (Fig. 8F), essel- ay pi s simila o in e es- sel pi s in size and shape h oughou he ay cell, mean angen ial di- ame e o essel lumina be ween ≤50μm (X=29.4μm; n=31); 20–40 essels pe squa e millime e (X=29.67 essels/mm2; n=228); essel elemen leng h no measu ed; ibe s: ibe s wi h dis inc ly bo de ed pi s in bo h adial and angen ial walls, sep a e, hin- o hick walled (Fig. 8F), ibe leng h no measu ed; Rays: 4–10 se ia e (Fig. 8G), heigh >1mm (Fig. 8D–E) ay cells up igh and squa e, shea h cells p esen (Fig. 8H–J), ≤4 ays/mm. Compa ison and ema ks: he wood ana omy o his single spec- imen clea ly i My sine L., and mos p obably ep esen ing he en- demic azo ean M. e usa (Menezes de Sequei a e al., 2012). De- ailed wood ana omy o M. e usa was ne e in es iga ed, bu close axa we e analyzed and illus a ed in de ail (e.g. Lens e al., 2005; Schweing ube e al., 2013; de Luna e al., 2018), and showing a compa able wood ana omy. Sha ed cha ac e is ics wi h My sine spp. wood a e he p esence o mul ise ia e ay cells and up igh cells, es- sels mos ly soli a y, angula (al hough de Luna e al., 2018, desc ibes species anging om ci cula o o al) and wi h a small angen ial diame e (< 50μm), essels wi h al e na e pi s and simple pe o a ions, ibe s sep a e and wi h simple pi s (e.g. Lens e al., 2005). Radial auma ic spli s, iewed in TS and TLS (Fig. 8B, D) occu h oughou he sample, mos p obably auma ic gas pipes o med by he ins an and in ense hea om he engul ing PDC. My sine wood shows cha ac e is ics simila o paedomo phic wood (e.g. Ca lquis , 1962; Ca lquis , 1974; Ca lquis , 2001), which is closely associa ed wi h plan s wi h insula woodiness (e.g. Ca lquis , 1974; Dulin and Ki cho , 2010). Rema kably, he sample Fai-28J esembles in TS and TLS he wood ana omy desc ibed o some lobelioid Hawaiian woody Campanulaceae (see Ca lquis , 1969). O de : ROSALES Be ch . & J.P esl Be ch old and P esl, 1820. Family: ROSACEAE Juss. 1789. P unus lusi anica L. subsp. azo ica (Mouill.) F anco. (= P unus azo ica (Mouill.) Ri as Ma . & Lousã & Fe n.P ie o & E.Días & J.C.Cos a & C.Aguia ) (TS: Figs. 9A–C; RLS: Figs. 9D–F; TLS: Figs. 9G–I) Ma e ial: Fai-28G, Fai-28O, Fai-28P, Fai-28Q, Fai-28V, Fai-28X, Fai-28Y, Fai-28Z, Fai-29A, Fai-29B, Fai-29C. IAWA code (Wheele e al., 1989): 1, 4, 6, 7, 10, 11, 12, 13, 22, 24, 30, 36, 37, 40, 50, 52, 62, 66, 68, 98, 107, 115. Wood desc ip ion: G ow h ings: ha dwood, dis inc g ow h ings bounda ies (Fig. 9A–B); Vessels: semi- ing-po ous, essels in an- gen ial, diagonal and adial pa e n, essels in mul iples o 4 o UNCORRECTED PROOF 16 Re iew o Palaeobo any and Palynology xxx (xxxx) xxx-xxx Fig. 11. The po en ial na u al communi ies o Faial Island and he Cancelas ash qua y paleophy odi e si y. A, Po en ial na u al ege a ion map o Faial Island based on Elias e al., 2016; B, paleophy odi e si y and paleoecological in e p e a ion o he Cancelas ash qua y cha coal assemblage. The plan ossils p ese ed wi hin deposi C11, which is be ween 1200y BP (Madei a e al., 1995) and 1000y BP (Pacheco, 2015), a e impo an es imonies o an insula e es ial paleoecosys em spa ially p ese ed as a T0assemblage (see Góis-Ma ques e al., in p ess). Plan ossils ound wi hin he C11 deposi s we e pa o a paleo o es g ow- ing 700–500yea s be o e he Po uguese coloniza ion o he Azo es Islands. These ossils a e he ideal g ound p oo s o es he Azo ean PNV bu also o econs uc an ecological baseline (see Nogué e al., 2017), essen ial o a u u e lo is ic es o a ion and conse a ion o Faial Island. Mo eo e , his is p obably applicable o o he Azo ean Islands, al hough u he paleobo anical and palynological s udies a e needed (Góis-Ma ques e al., in p ess). Paleo ege a ion ha is spa- ially p ese ed bu ime es ic ed (i.e. single olcanic e en ha led o he bu ial o insula o es s), coupled wi h spa ially es ic ed bu wi h long- e m in o ma ion, such as he palynological eco d ound in limnic sedimen s, a e essen ial o he econs uc ion o Azo ean p e-an h opic ege a ion. 6.4. An h opic in luence and ex i pa ion? Why did Azo ean P unus almos disappea , and o he ees such as L. azo ica did no ? Acco ding o F u uoso (1590a, 1590b, 1590c) ,P. lusi anica subsp. azo ica was an abundan ee in all Azo ean is- lands, g owing oge he wi h o he endemic and na i e ees. This si u- a ion is e y simila o he one ound in he cha coali ied ees in Can- celas qua y. Howe e , since he mid-15 h cen u y, he Azo es na u al landscapes we e massi ely modi ied (Dias, 2007) by logging ees o uel and cons uc ion, and opening he o es s o ag icul u e and pas- u es (e.g. Dias, 2007). Habi a educ ion is one o he main d i e s o he educ ion o his pa icula ee (Bilz, 2011). Howe e , and as al- eady poin ed ou by F u uoso (1590a), P. lusi anica subsp. azo ica is poisonous o ca le by inges ion. The symp oms and ecological clues o plan oxici y a e desc ibed in de ail: “…howe e , he oliage o ‘ginja’[P. lusi anica subsp. azo ica], on Ma ch onwa d, g owing in highlands and da kening woodlands makes hem ( he ca le) o u ina e blood.”(F u uoso, 1590a, pg. 212; ansla ion by he au ho s). P unus spp. a e poisonous o uminan s (see Wags a , 2008 and e e ences he ein), due o he p esence o cyanogenic glycosides. The ac i e con- ol o P. lusi anica subsp. azo ica o a oid animal in oxica ion and dea h could mos p obably be a cause o i s decline, along wi h he use o wood o uel. The con ol o poisonous ees is no unknown in Po ugal; o example, Taxus bacca a L. was d i en o an acu e de- cline by shephe ds (D ape and Ma ques, 2006) o his eason. O he hypo heses could be ela ed o his ee being endozoocho ic, ha ing su e ed om he nega i e e ec s o human coloniza ion. The decline o e en ex inc ion o bi ds ha ac ed as ec o s canno be uled ou , as he paleozoological ossil eco d has a eco d o ex inc axa (e.g. Rando e al., 2013; Alco e e al., 2015; Rando e al., 2017). The de- cline o P. lusi anica subsp. azo ica was mos p obably a combina ion o causes, such as logging and habi a loss, associa ed wi h he loss o seed dispe se s and an ac i e popula ion con ol, leading o he almos e adica ion o his ee. Recen ly, Ca ine and Menezes de Sequei a (2019) p oposed he exis ence o a Hooke ian sho all, an addi ion o se e al biological knowledge gaps ha ha e been p oposed (see Ho al e al., 2015 o u he de ails). The Hooke ian sho all ques ions how human impac on p is ine oceanic islands shaped he p esen di e si y pa e ns ob- se ed oday. The Faial Island ossil eco d poin s o a paleo ege a ion assemblage ha di e s om he obse ed and modeled (Elias e al., 2016) and ha is mo e simila o he his o ical accoun s (Fe nandes, 1508; F u uoso, 1590a; F u uoso, 1590b; F u uoso, 1590c). The s udy o Faial's medie al ossil o es will ce ainly con ibu e o ill he Azo ean Hooke ian sho all. 7. Conclusions Faial Island is a unique island wi hin he Azo es A chipelago o s udy p e-an h opic paleophy odi e si y and paleo ege a ion, due o plan ossil p ese a ion ha esul ed om Holocene ecu en explo- si e olcanism (especially he C11 e en ). The de ailed s udy o he wood ana omy o 41 samples allowed he iden i ica ion o 7 phane ophy es, co esponding o ex an axa. The spa ial dis ibu ion o he si es s ongly sugges s he p esence o wo ege a ion bel s. The iden i ica ion o a o es p ese ed in si u, p o- ides a unique iew in o a p is ine Meghalayan ege a ion wi hin he Azo es Islands. Unexpec edly, P unus lusi anica subsp. azo ica was he second mos abundan ossil wood ound, indica ing ha his ee was much mo e abundan in he ecen pas , as co obo a ed by his o - ical accoun s. As a concluding ema k, he ecu en explosi e olcanism in Fa- ial Island allowed he p ese a ion o ‘ ecen ’mac o ossils. These, in he con ex o a ecen ly colonized bu highly an h opically modi ied island, a e he missing link o ully app ecia e hese paleoecosys ems in a p is ine s a e, and ideal o ill he knowledge gap o he Hook- e ian sho all (Ca ine and Menezes de Sequei a, accep ed). Fu he wo k is needed in Faial Island o ha e a ull pic u e o he paleophy UNCORRECTED PROOF Re iew o Palaeobo any and Palynology xxx (xxxx) xxx-xxx 17 odi e si y and paleo ege a ion bu ied by he C11 explosi e olcanic ac i i y. Decla a ion o Compe ing In e es s The au ho s decla e ha hey ha e no known compe ing inancial in e es s o pe sonal ela ionships ha could ha e appea ed o in lu- ence he wo k epo ed in his pape . Unci ed e e ences B ongnia , 1843 B own, 1810 de Jussieu, 1789 Go ozhankin, 1904 Ho mannsegg and Link, 1809–1820 Humbold e al., 1817 Knoblauch, 1934 Lindley, 1833 Linnaeus, 1753 Smi h, 1819 on Ma ius, 1835 Willkomm, 1854 Acknowledgemen s We would like o hank Paloma de Palacios and Luis Ga cía (Uni- e sidad Poli écnica de Mad id) o p o ide access o he Maca one- sian wood e e ence collec ion; Ad iano Pimen el (Azo es Uni e si y) o indica e some key locali ies in Faial Island; A u Gil (Azo es Uni- e si y) o helping wi h GIS da a; Jo ge Capelo (Ins i u o Nacional de In es igação Ag á ia e Ve e iná ia) o ui ul discussion on Azo es ege a ion; Telmo Nunes (FCUL-MF) o SEM sample p epa a ion and imaging; Ria Mi chell o he e iew ha imp o ed his man- usc ip . 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