Carpological Analysis of Phoenix (Arecaceae): Contributions to the Taxonomy and Evolutionary History of the Genus

Home , Carpology, Orto botanico di Palermo, Phoenicites, Phoenix (plant), Phoenix acaulis, Phoenix atlantica

bs_bs_banner

Botanical Journal of the Linnean Society, 2014, 175, 74–122. With 25 ﬁgures

Carpological analysis of Phoenix (Arecaceae): contributions to the taxonomy and evolutionary history of the genus

DIEGO RIVERA FLS1*, CONCEPCIÓN OBÓN FLS2, JOAQUÍN GARCÍA-ARTEAGA2, TERESA EGEA2, FRANCISCO ALCARAZ1, EMILIO LAGUNA3, ENCARNA CARREÑO1, DENNIS JOHNSON4, ROBERT KRUEGER5, JOSÉ DELGADILLO6 and SEGUNDO RÍOS7

1Depto. Biología Vegetal, Fac. Biología, Universidad de Murcia, 30100 Murcia, Spain 2Depto. De Biología Aplicada, Escuela Politécnica Superior de Orihuela. Ctra. Beniel, Km 3,2. Universidad Miguel Hernández, 03312 Orihuela, Alicante, Spain 3Generalitat Valenciana. Conselleria d’Infraestructures, Territori i Medi Ambient. Servei de Vida Silvestre/Centre per a la Investigació i Experimentació Forestal. Avda. Comarques del País Valencià, 114. 46930 Quart de Poblet. València, Spain 43726 Middlebrook Ave, Cincinnati, OH 45208, USA 5National Clonal Germplasm Repository for Citrus and Dates, Riverside, 1060 Martin Luther King Blvd, Riverside, CA 92507-5437, USA 6Facultad de Ciencias, Campus de Ensenada, Universidad de Baja California, Ensenada, Baja California ZP 22830, Mexico 7CIBIO, Universidad de Alicante, Alicante, Spain

Received 23 July 2013; revised 17 November 2013; accepted for publication 23 February 2014

The main purpose of this study was, first, to analyse the morphology of seeds of Phoenix spp. and relevant cultivars and to assess the taxonomic value of the information generated as a means of studying the systematics and evolutionary history of the genus Phoenix. We then analysed seed morphological diversity in P. dactylifera, supported by morphotypes shared with fossil and/or archaeological materials, to advance the knowledge of the origins, history and biogeography of one of the most important cultivated palm species. The other objective was to develop a methodology for assigning different commercial seed samples and archaeological materials to determined morphotypes as a tool for their identification at the species level. Three hundred and sixty-four seed samples (3920 seeds) were analysed: 304 samples of modern Phoenix spp. (including five herbarium type specimens and eight type icons), 51 archaeological samples and nine fossil seed samples and subsamples. Information was systematized in a crude matrix with 364 units representing seed samples and 67 descriptors. Descriptors are frequencies, in percentage, for each of the 41 qualitative states and of the 26 classes that were recognized for the quantitative parameters. Analyses proceeded sequentially, starting with modern samples consisting of type specimens and botanically verified specimens. Eight species show characteristic seeds and are clearly assigned to morphotypes [P. acaulis, P. canariensis s.s., P. paludosa, P. reclinata, P. roebelenii, P. rupicola, P. sylvestris and P. theophrasti (excluding populations from Datça, Turkey)]; the other taxa are not clearly separated on the basis of the seed morphology alone. In parallel, fossil and archaeobotanical samples were analysed. There is no clear separation between fossil and archaeological samples, between different periods of the archaeological samples or geographical origins. Combination of modern, fossil and archaeological seed results in the same analysis revealed that it is possible to allocate archaeological and fossil materials to morphotypes shared with modern living Phoenix spp. All archaeobotanical samples could be classified in groups with modern seed samples. The assignment of archaeobotanical samples was made, mainly, to morphotypes of P. dactylifera. However, some samples were assigned to morphotypes of P. reclinata, P. caespitosa, P. atlantica, P. theophrasti, P. pusilla and P. canariensis. Archaeological seeds were not allocated to group 19, containing the samples of P. sylvestris, P. iberica and the Miocene fossil P. bohemica. It appears that species such as P. theophrasti, P. canariensis, P. caespitosa and P. reclinata formerly

*Corresponding author. E-mail: [email protected]

74 © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175, 74–122 CARPOLOGICAL ANALYSIS OF PHOENIX 75 had a much wider area of distribution. The morphology of two of the three Eocene samples (Phoenicites occidentalis and Phoenix hercynica) is that of P. dactylifera. Attribution and dating of these samples need to be carefully reviewed. Apparently the great diversity of P. dactylifera date morphotypes during the Neolithic was followed, during the Chalcolithic and the Bronze Age, by a remarkable constriction (bottleneck) in terms of morphological variability, which slowly recovered from the Iron Age onwards. With the currently available evidence, we cannot exclude a group ancestral to P. dactylifera in the Persian Gulf, related to the eastern chlorotype. In parallel, another group ancestral to P. dactylifera may exist in the western Mediterranean, including P. iberica, related to the western chlorotype. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175, 74–122.

ADDITIONAL KEYWORDS: Holocene – Miocene – multivariate analysis – palaeobotany – Palaeotropical – Phoenicites – Pleistocene – seed morphology – Tertiary.

INTRODUCTION the cultivated gene pool, the status of P. atlantica A.Chev. as distinct from P. dactylifera and its possible The genus Phoenix L. (Arecaceae) comprises 13 presence on the African continent (Mauritania and (Barrow, 1998) to 20 (Beccari, 1890) species of mostly Morocco) need to be clarified (Pintaud et al., 2013). tropical, dioecious palms with solitary stems, rarely Fruits (dates) normally develop after pollination, branched, or, in some species, with a short under- resulting in dates with seeds. Unpollinated female ground stem, ending in a crown of 20–150 pinnate flowers may develop seedless, poor-quality fruits that leaves. The fruits are berries (with a fleshy mesocarp are normal in other respects. Date fruits are ellipsoi- and a membranous rudimentary endocarp), known as dal to ovoid or almost cylindrical. Dimensions of dates ‘dates’, borne in clusters of tens or hundreds, which are variable, ranging from 10×5mminP. roebelenii develop from three-carpellate female flowers, in O’Brien (Iossi, Vitti & Rubens, 2006), to 75 × 35 mm which two carpels normally abort. in P. dactylifera ‘Medjool’. The phylogenetic isolation of Phoenix has long been Phoenix seeds are typically elliptical and slightly established. It has been placed in tribe Phoeniceae flattened dorsiventrally and have a longitudinal (Uhl & Dransfield, 1987; Asmussen et al., 2006) in furrow on the ventral face. On the dorsal face, the subfamily Coryphoideae, appearing to be the only operculum or micropyle appears at the middle point of palm group displaying the induplicate insertion of the seed, although, often, it can be slightly displaced leaf segments (Uhl et al., 1995; Dransfield et al., towards the proximal or distal end. Only one species, 2005). Phoenix differs from related genera of Coryph- P. paludosa Roxb., has a nearly basal operculum. The oideae by having pinnate rather than palmate leaves. rigid date seeds (because of the hard endosperm) are Although firmly anchored in Coryphoideae, Phoenix erroneously called date ‘stones’ or ‘kernels’ in the appears to be on a deep branch in phylogenetic trees, archaeobotanical literature, leading the reader to being sister to the large, pantropical tribe Trachy- misinterpret dates as drupes (Hopf, 1983; Kislev, carpeae (Dransfield et al., 2008). Molecular phyloge- Hartmann & Galili, 2004). Problems during pollina- netic dating placed the divergence of the Phoeniceae tion and fruit development can lead to the incom- lineage during the early Tertiary (Couvreur, Forest & pletely developed or abnormal seeds. Baker, 2011). Date fruits and their seeds present a set of charac- Phoenix dactylifera L. (the date palm) has the ters that are used as descriptors for the systematics of widest distribution and the highest morphological Phoenix spp. and cultivars (Beccari, 1890; Barrow, diversity in Phoenix, and it is the most numerous in 1998; IPGRI, 2005). However, some of these charac- terms of individuals and populations. Although ters, such as testa colour, endosperm colour or the hybrid origins have been proposed, molecular data presence/absence of a ruminate endosperm, cannot be have demonstrated that P. dactylifera is a true used for identification of palaeobotanical materials, species, distinct from all other species of the genus because they are lost during pre- and post-depositional (Pintaud et al., 2010). Recent genetic data and phylo- processes affecting the seeds. However, seed morphol- genetic data based on DNA sequences of the plastid ogy is taxonomically relevant and several nomenclatu- loci psbZ–trnfM and rpl16–rps3 indicate a strong ral types of Phoenix spp. are seeds or seed illustrations, geographical structure of the genetic diversity of the for example, P. pusilla Gaertn. (Gaertner, 1788–1791). date palm at all scales (local, regional, global) and the The shape of Phoenix seeds is characteristic and allows importance of isolation and intraspecific gene flow in determination of both fossil and archaeobotanical (car- shaping the present day agrobiodiversity. Although bonized, desiccated or mineralized) materials at the there is no evidence of interspecific hybridization in generic level. However, in routine identifications of

© 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175, 74–122 76 D. RIVERA ET AL. palaeobotanical or archaeobotanical date seeds, it is or Cote d’Azur, France), P. acaulis Roxb. (Kolkata, uncommon for attributions to proceed beyond the India) and P. roebelenii (Protheroe & Morris, Leyton- genus, generating a debate about whether they are stone, UK). Therefore, it is important, in sequence, to wild or cultivated (Terral et al., 2012). solve taxonomic problems and difficulties of typifica- Fossils showing affinities with Phoenix seeds have tion and then to proceed with analysing variability in been recorded from Tertiary levels of eastern Texas as populations of Phoenix, both wild and cultivated. Phoenicites occidentalis Berry (Berry, 1914), from the The first detailed descriptions of Phoenix seeds middle Eocene of Germany (Geiseltal) as Phoenix (P. dactylifera and P. pusilla) were by Gaertner hercynica Mai and Serenoa carbonaria Mai (Mai, 1976) (1788–1791), who used six characters totalling 11 and from the lower Miocene of Central Europe as states. Gaertner did not, however, use any character Phoenix bohemica Buzek (Buzek, 1977; Harley, 2006). based on the dimensions of the seeds (Table 1). Phoenix seeds were found in numerous archaeologi- The first comprehensive monograph of Phoenix, cal sites from North Africa and the Near East, in published by Beccari (1890), used the position of the levels from the 8th millennium BP onwards. In micropyle as a character for distinguishing species. In Ancient Egypt, dates were eaten fresh, dried and used that work the number of characters (quantitative and in magical compounds. One small carbonized seed qualitative) used to describe seeds is 15, totalling 38 was recovered from Predynastic El Omari, Helwan states. The most recent monograph of Phoenix by (Debono, 1948). Date seed recovered in other prehis- Barrow (1998) provides much less detailed descrip- toric settlements (Abu Umuri, Naqada) seem doubtful tions of seeds, using only ten characters and 20 states (Täckholm & Drar, 1950). Numerous finds are known (Table 1). from later periods, including a dish with date frag- The need to create a set of descriptors to distinguish ments and seed (11th Dynasty, Ani’s tomb, El Gaba- date palm (P. dactylifera) cultivars and land races, led lein) (Loret, 1892), dates [12th Dynasty, Dira Abu el researchers to use descriptive characters already pub- Naga (Schweinfurth, 1883), 19th Dynasty, Thebes; lished by Beccari (1890) and to introduce others. Nixon 21st Dynasty, tomb of Pinotem I, El Deir el Bahari], (1950), for example, used only 11 characters, with a dates mixed with other fruits (16th Dynasty, Mayana; total of 34 states; as quantitative descriptors he used 18th Dynasty, tomb of Khà, Thebes), seeds strung into only length and width. The official catalogue of stand- necklaces (New Empire and 18th Dynasty, Deir el ardized descriptors for the date palm (IPGRI, 2005) Madina), small dates and seeds (18th Dynasty, tomb (International Plant Genetic Resources Institute, now of Sennufer, Thebes), date cakes (Early Ptolemaic, Bioversity International) comprises 12 characters and Thebes) and peculiar long narrow dates (Greek, El 30 states referring to seeds (Table 1). Faiyum) (Täckholm & Drar, 1950). Terral et al. (2012) studied c. 1200 individual Phoenix is a genus of wild and cultivated plants Phoenix seeds using dorsal and lateral outlines, and that can be found in a range of different natural (from 64 equally spaced points (pseudo-homologous land- mangroves to pine forests) and anthropogenic habi- marks) were analysed. However, raphe, micropyle tats (from sandy beaches to oases). This genus is position, mucro and different superficial processes notoriously difficult to classify to the species level were excluded from this analysis. based on incomplete herbarium specimens and, even Morphological demarcation of species within in botanic gardens, outside their native habitats, indi- Phoenix involves numerous vegetative, floral, fruit viduals of Phoenix can be seriously challenging to and seed characters. Phoenix paludosa is easily dis- identify. Phoenix dactylifera is found almost exclu- tinguished by its leaflet discolour (abaxial lamina sively under cultivation, although occasionally, if con- surface greyish) and its seeds with a basal embryo. ditions are favourable, there are feral and wild Leaflets with abaxial ramenta are typically present in populations (Zohary & Hopf, 2000; Rivera et al., P. andamanensis S.Barrow, P. reclinata, P. roebelenii 2012b). Phoenix sylvestris (L.) Roxb. and P. canarien- and P. rupicola T.Anderson (Barrow, 1998). In the sis H.Wildpret are systematically exploited for their group with ramenta, the few herbarium specimens sweet sap to produce a concentrate known as palm available of P. andamanensis have seeds with a rumi- honey and the palms are often planted for that nate endosperm. Phoenix roebelenii is characterized purpose in their natural areas, making it difficult to by its small size (stems to 2–3 m tall, leaves to 1.5 m distinguish scattered crops and natural and feral long), whereas stems to 10 m tall are typical of P. rec- populations. The other species are often found in linata (clustering) and P. rupicola (solitary). Acute to cultivation as ornamental palms or as rare specimens acuminate staminate petal apices with jagged for collectors. Several species were originally fully or margins are typical of P. reclinata (Barrow, 1998). The in part described from specimens grown in botanical prominent horn-shaped swelling of the rachilla, sub- gardens and collections, including P. reclinata Jacq. tending each fruit, is exclusive to P. acaulis; in addi- (Schönbrunn, Austria), P. canariensis (Orotava, Spain tion, this species is acaulescent. Leaflets four-ranked,

Table 1. Historical review of the use of seed morphological characters and states in Phoenix. Codes: B, breadth; D, depth; L, length; TD, totalized dimensions

Gaertner (1788–1791)‡ Beccari (1890)† Nixon (1950)§ Barrow (1998) IPGRI (2005)§ This paper

L (mm) Not 8–40 mm, continuous 18–36 mm, continuous 7–30 mm, continuous Continuous 4.47–40, continuous or six states (4–10, 10–15, 15–19, 19–25, 25–32, 32–40) B (mm) Not 4.5–12 mm, continuous 6.5–11 mm, continuous 3–10 mm, continuous Continuous 1.31–13(15), continuous or six states (1–3.5, 3.5–6, 6–8, 8–10, 10–12, 12–16) D (mm) Not 4.5–10 mm, continuous Not 3–10 mm, continuous Continuous 0.95–13(17), continuous or ﬁve states (0–2.5, 2.5-4, oaia ora fteLnenSociety Linnean the of Journal Botanical 4–5.5, 5.5–7, 7–17) B/L Not Not Not Not Not 0.09–1.17, continuous or ﬁve states (0–0.2, 0.2–0.4, 0.4–0.6, 0.6–0.8, 0.8–1) D/B Not Not Not Not Not 0.32–1.5, continuous or four states (0–0.75, 0.75–0.85, 0.85–0.95, 0.95–1.5) TD (mm3) Not Not Not Not Not 36–10 134, continuous or six states (0–150, 150–300, 300–800, 800–1200, 1200–1850, 1850–10 200) OF ANALYSIS CARPOLOGICAL Overall Two states (oblong, Six states (cylindric, Seven states (narrowly Five states (narrowly Five states (ovoid, Seven states shape oblong–ovate) compressed from oblong, oblong, elongate, elongate, conical, fusiform, (ovate–triangular, elliptic, base to apex, oblong, oblong–elliptical, obovoid, ovoid, subcylindrical, oblong, cylindrical, ovate–elongated, oblong–wedge-shaped, terete) pyriform) globose*, hemispherical, 2014, , ovate–triangular, oblong–spathulate, narrowly fusiform) ovate–elliptic) elliptical, obovate–elliptical)

175 Colour Two states (reddish, Three states (greyish, Five states (light brown, Three states Three states (beige, Four states (blackish, blackish–brown) chestnut-brown, medium brown, greyish (grey–brown, grey, brown) greyish, cream, brown) 74–122 , cinnamon-brown) brown, light greyish brown, chestnut-brown, dark brown) pinkish-brown) Apex Not Two states (rounded, Four states (blunt, somewhat Three states (rounded, Not Five states (obtuse, acute, acute) pointed, somewhat broadly pointed, squared) retuse, oblique, truncate) pointed, rounded) Base Not Two states (rounded, Two states (abrupt, others) Three states (rounded, Not Four states (obtuse, acute, PHOENIX acute) pointed, squared) oblique, truncate) Apical mucro Not Not Two states (present, lacking) Not Not Two states (present, lacking) Basal mucro Not Not Not Not Two states (present, Two states (present, lacking) lacking) 77 78 .RIVERA D. Table 1. Continued

Gaertner (1788–1791)‡ Beccari (1890)† Nixon (1950)§ Barrow (1998) IPGRI (2005)§ This paper

Surface Two states (glossy and Two states (glossy, Not Two states (glossy, Not Two states (glossy and AL ET smooth, rough and rough) matt) smooth, rough and matt)

04TeLnenSceyo London, of Society Linnean The 2014 © matt)

Longitudinal Two states (present, Three states (present, Not Not Not Two states (present, . grooves lacking) shallow, lacking) lacking) Transverse Three states (smooth, Two states (ﬁnely Not Not Four states Three states (wrinkled, processes wrinkled, grooved) grooved, uniform) (smooth, ﬁnely grooved, uniform) wrinkled, bumpy, grooved) Micropyle One state (central) Two states (central, Four states (central, a little Two states (lateral, Three states Two states (central, basal) basal) above middle, near base, basal) (proximal, variable) central, distal) Raphe Two states (deep, Three states (shallow, Six states (medium in width Not Three states Three states (shallow, wide) narrow, wide) and depth, closed in the (shallow, V-shaped, U-shaped) center, closed, narrow and V-shaped, shallow, narrow and deep, U-shaped) open)

oaia ora fteLnenSociety Linnean the of Journal Botanical Raphe length Not Not Not Two states (full, Three states (Short, Not incomplete) Medium, Long) Dorso-ventral Not Two states (bent, Not Not Not Two states (bent, straight) curvature straight) Lengthwise Not Two states (present, Two states (present, lacking) Not Four states Two states (present, ridges or lacking) (lacking, wings, lacking) wings ridges, wings and ridges) Frequency of Not Not Not Not Three states Continuous, percentage of ridges or (lacking, winged seeds wings occasionally, frequently) Dorsal Not Three states (very Two states (present, lacking) Not Not Not furrow broad, very open, deep and narrow) 2014, , *Only in outgroups. †Beccari (1890) situates the ventral face of the seed in the zone of the micropyle, and the dorsal face opposite in the zone of the raphe; however, for Iossi et al. (2006) and IPGRI (2005) the dorsal face is the zone of the micropyle. 175 ‡Descriptions are restricted to only two species. 74–122 , §Descriptions for Phoenix dactylifera cultivars exclusively. CARPOLOGICAL ANALYSIS OF PHOENIX 79 seed glossy, chestnut brown and stems short (rarely to and cultivars in order to assess the taxonomic value 4 m) are characteristic of P. pusilla. Leaflets not four- of the generated information as a means of achieving ranked, seed matt, greyish and stems short (rarely to more promising research; i.e. a study of systematics 4 m) are characteristic to P. loureiroi Kunth. The and the evolutionary history of the genus Phoenix, group of robust tree palms with solitary trunks and, second, to compare seeds of living Phoenix spp. includes P. canariensis (to 1.2 m in diameter) and with palaeobotanical materials in an attempt to P. sylvestris (to 30 cm in diameter). However, solitary determine ancestral states. We also analysed seed stemmed P. dactylifera cultivars and seedlings are morphological diversity in P. dactylifera, supported by relatively frequent. Clustering robust palms, often morphotypes shared with fossil and/or archaeological with basal suckers, include P. atlantica A.Chev., materials, to advance knowledge of the origins, P. caespitosa Chiov., P. dactylifera L., P. iberica history and biogeography of one of the most impor- D.Rivera, S.Ríos & Obón and P. theophrasti Greuter. tant cultivated palm species. Species in this last group do not appear clearly dif- Other objectives are to develop a methodology to ferentiated and there is discussion of their affinities assign different commercial seed samples and and status (Gros-Balthazard, 2013). archaeological materials, to determined morphotypes Hybridization is a phenomenon considered to be as a tool for their identification to the species level. common among Phoenix spp. However, the allocation This will contribute to reduce the impact of misiden- on morphological grounds of individuals or populations tification in horticulture and will produce a frame- to a particular hybrid is difficult and often erroneous. work for identification and interpretation of Morphometric approaches are postulated as a tool to archaeobotanical Phoenix materials. identify hybrids (Gros-Balthazard, 2013). González, Caujapé & Sosa (2004) gathered evidence for introgres- sion in mixed populations in Maspalomas and Tafira MATERIAL AND METHODS (Canary Islands, Spain) where pure P. canariensis and P. dactylifera (cultivated and feral) individuals with PLANT MATERIAL species-specific random amplified polymorphic DNA Date palm seeds are rarely preserved as such in (RAPD) markers co-occur with morphologically inter- carpological collections or as herbarium specimens. mediate individuals in which these markers are com- For example, of the 480 sheets of Phoenix in the bined. However, natural interspecific hybridization herbarium of the National Museum of Natural has never been reported (Gros-Balthazard, 2013). History of Paris (France), only 13 contained fruits Experimental pollination of several P. dactylifera cul- mature enough to be able to extract seeds that could tivars with pollen of P. pusilla produced seedless dates. be analysed (but would destroy the fruit), and only Although seed development was noted initially, the one contained free and abundant seeds; in the Fair- breakdown of endosperm development was evident child Tropical Botanic Garden Virtual Herbarium, later on. Because of the development of disorders in the only eight of 87 specimens had ripe fruits. When endosperm development, the embryo growth and whole date fruits are preserved in herbarium speci- development also ceased (Sudhersan, Jibi & Al-Sabah, mens, the extraction of seeds for study is not possible 2010). Therefore, not all possible crosses between without destroying the fruit; therefore, we only Phoenix spp. are able to produce viable seeds and studied seeds which were free and clean. Herbarium hybrids. Interspecific pollination events may influence specimens were directly studied from the Jardín the dimensions of fruits and seeds as shown in differ- Botánico de Madrid (MA), Herbarium of the Univer- ent metaxenia experiments. For instance, P. dactylif- sidad Miguel Hernández (UMH) and Beccari’s her- era cultivars produce smaller seeds when their female barium specimens in the Botanical Museum Florence flowers are fertilized with pollen of P. canariensis or (F) or using high resolution images from the Berlin P. loireiroi (Gros-Balthazard, 2013). Therefore, it is Botanischer Garten (Röpert, 2000), Royal Botanic necessary to differentiate between hybrid seeds (modi- Garden, Edinburgh, Royal Botanic Gardens, Kew, fied by metaxenia) and seeds produced by hybrid Smithsonian Institution, Washington, and National female individuals of known parentage. For the pur- Museum of Natural History of Paris. In parallel, we poses of the present paper we name ‘hybrid’ seeds of collected seed samples from living Phoenix individu- the second type, for their parentage we relied on als in Algeria, France, Greece, Italy, Libya, Mexico, pedigree records (experimental hybrids). Hybrids Morocco, Spain, Tunisia and Yemen. Seed samples between P. dactylifera and P. canariensis have seeds of were also obtained from botanic gardens (Fairchild intermediate size between the two species Tropical Botanic Garden, Orto Botanico di Palermo, (Gros-Balthazard, 2013). Orto Botanico ‘Pietro Castelli’ dell’Università di The purpose of the present study is, first, to analyse Messina, Orto Botanico dell’Università di Catania, morphological characters of seeds from Phoenix spp. Jardín Botánico de la Universidad de Valencia) and

Table 2. List of living Phoenix seed samples. Nomenclature adopted in this paper compared with Barrow (1998) and Govaerts et al. (2011). Further information on samples and vouchers in the Supporting Information and for accessions INIA (2012) and Rivera et al. (2012a). NSAM, number of seed samples analysed; NT, number of type specimens; NI, number of type icons analysed

Species Barrow (1998) Govaerts et al. (2011) Main area NSAM NT NI

Outgroups 4–– Phoenix acaulis Roxb. P. acaulis Roxb. P. acaulis Roxb. India 5 0 1 Phoenix andamanensis P. andamanensis P. andamanensis South-East Asia 1 1 0 S.Barrow S.Barrow S.Barrow Phoenix caespitosa P. caespitosa Chiov. P. caespitosa Chiov. Arabia and Yemen 5 0 0 Chiov. (= P. arabica Burret) Phoenix atlantica ¿P. atlantica A.Chev.? P. atlantica A.Chev. Cabo Verde 4 0 0 A.Chev. Phoenix canariensis P. canariensis Chabaud P. canariensis Chabaud Canary Islands 18 0 1 H.Wildpret Phoenix dactylifera L. P. dactylifera L. P. dactylifera L. North Africa (from 164 0 1 (including P. excelsior Egypt westwards to Cav.) Algeria), Near East (Iran, Iraq, Arabia and Yemen), Spain Phoenix dactylifera L. P. dactylifera L. P. dactylifera L. Spain, Cabo Verde and 210 var. adunca Becc. North Africa Phoenix dactylifera L. P. dactylifera L. P. dactylifera L. Spain and Baja 210 var. costata Becc. California (Mexico) Phoenix interspeciﬁc ––– 1700 hybrids Phoenix iberica Not included P. dactylifera L. Spain 0 1 0 D.Rivera, S.Ríos & Obón Phoenix loureiroi Kunth P. loureiroi Kunth P. loureiroi Kunth South-East Asia and 24 1 0 India Phoenix paludosa Roxb. P. paludosa Roxb. P. paludosa Roxb. South-East Asia 5 0 0 Phoenix pusilla Gaertn. P. pusilla Gaertn. P. pusilla Gaertn. India and Sri Lanka 2 0 3 Phoenix reclinata Jacq. P. reclinata Jacq. P. reclinata Jacq. Africa 4 0 1 Phoenix roebelenii P. roebelenii O’Brien P. roebelenii O’Brien South-East Asia 8 0 0 O’Brien Phoenix rupicola P. rupicola T.Anderson P. rupicola T.Anderson India 6 0 0 T.Anderson Phoenix sylvestris (L.) P. sylvestris (L.) Roxb. P. sylvestris (L.) Roxb. India 12 0 1 Roxb. Phoenix theophrasti P. theophrasti Greuter P. theophrasti Greuter Crete and South-west 800 Greuter Turkey repositories (National Clonal Germplasm Repository scattered individuals living along the Chicamo River, for Citrus and Dates, Riverside, CA, USA). Finally, but interspersed with individuals, or small clumps, of commercial samples of dates and horticultural seeds the local varieties of P. dactylifera. This taxon has not were acquired for comparison. yet been found among the thousands of date palms in The species-level nomenclature (Table 2) follows Elche. Vegetatively, they are similar to P. theophrasti Barrow (1998) and Govaerts et al. (2011), except for Greuter and their fruits are small, rounded with thin P. iberica D.Rivera, S.Ríos & Obón and P. canariensis. ﬂesh and similar to the fruits of P. sylvestris. The Phoenix iberica was described from the Chicamo area authorship of P. canariensis is exhaustively discussed (Abanilla, Murcia), 40 km south-west of Elche, Spain. by Rivera et al. (2013a) and, in summary, it is H.Wild- Originally it was based upon one small population of pret in Chabaud, because Chabaud himself did not spontaneous palms growing in a single ravine, and accept the new species in the original publication and

© 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175, 74–122 CARPOLOGICAL ANALYSIS OF PHOENIX 81 published a description of the species by Hermann account previous studies (Beccari, 1890; Barrow, 1998; Wildpret. The type of P. caespitosa Chiov. was col- IPGRI, 2005) and observation of the samples analysed. lected in north-eastern Somalia, our samples were Terminology for characters and states follows Stearn collected in the classical locality of P. arabica Burret (1978), Barrow (1998) and IPGRI (2005). in Yemen and from plants introduced from Saudi Each of the 3920 seeds was individually described Arabia into the USA and from cultivars. Barrow using the 20 characters as given in Table 1. Quanti- (1998) and Govaerts et al. (2011) both considered tative characters were measured using a Mitutoyo P. arabica to be conspecific with P. caespitosa. Absolute Digimatic 500-202-21 digital caliper with a Three hundred and sixty-four seed samples total- precision of 0.01 mm and recorded on an Excel spread ling 3920 seeds were analysed. Also, 286 samples of sheet. Allometric relationships (B/L, D/B) and total- modern seeds were desiccated and reduced to c. 20% ized dimensions (L×B×D inmm3) were automati- moisture content using a Sicco Auto-Star Desiccator, cally calculated using formulas. and preserved with Scharlau silica gel with a humid- Qualitative characters were analysed with a bin- ity indicator (orange), 2.5–6.0 mm, at 5 °C in two ocular Olympus SZ microscope and a Philips 220CW Liebherr K42 refrigerators. These seeds were flat screen. Photographs of ventral, dorsal and lateral obtained from living Phoenix spp. and cultivars and views were taken for all samples using a Lumix FZ60 four outgroups (Euterpe Mart., Livistona R.Br., Nan- camera with a Leica DC lens. Another full set of norrhops H.Wendl. and Washingtonia H.Wendl.). Lots images was obtained using a Canon EOS 350D comprising c. 15 seeds each were analysed. Voucher camera. A second observer verified the qualitative specimens have been deposited in the UMH (Miguel data using these photographs. Hernández University) herbarium and carpological Individual seeds differ slightly in shape and dimen- collection (Table 2 and see also Supporting Informa- sions within the same palm and even the same bunch tion, Appendix S1). These samples derive from field and branchlet. Therefore, our interest was to develop collections (119), horticultural samples (64), commer- a method to compare individuals, cultivars and cial date fruits (60) and botanic gardens and reposi- species, depending on their overall seed morphology. tories (43). The samples analysed were randomly To describe sample observations and measurements selected as subsamples from samples usually contain- of the individual seeds they were converted to discrete ing 25–1000 seeds. From each sample, another sub- categories and frequencies within each of the sample of five to 25 randomly selected seeds was samples. This allowed us to compare samples in regularly sown, germinated and the plants grown in terms of not only mutually exclusive states or average the National Phoenix repository (Escuela Politécnica values, but also in terms of the proportion of seeds of Superior, Universidad Miguel Hernández and Soto I6, each sample presenting those states. Ayuntamiento de Orihuela and Confederación Hidro- To calculate frequencies in the cases of continuous gráfica del Segura, both in Orihuela, Spain) (INIA, quantitative parameters such as dimensions (length, 2012; Phoenix Spain, 2013). width and thickness), allometric relationships and the Five samples of herbarium type specimens and five totalized dimensions and to compare the samples, other relevant herbarium specimens were measured parameters were each reduced to four to six classes or (Table 2). Eight icons or figures previously designated categories. Samples were then analysed in terms of as nomenclatural types for Phoenix spp. were ana- counts (proportion of individual seeds falling within lysed, interpreted and measured (Table 2). the class) and expressed as a percentage (Tables 4 Fifty-one archaeological and nine fossil seed samples and 5). This allowed us to use quantitative and quali- and subsamples, each comprising one to 39 seeds, were tative characters together in a single matrix. measured using images and data available in the Information was systematized in a crude matrix original publication or provided by different research- with 364 units (seed samples) (Table 1 and Support- ers, herbaria, museums and repositories (Table 3). ing Information, Appendix S1) and 67 descriptors. Descriptors are frequencies, as a percentage, for each of the 41 qualitative states and 26 classes that were MEASUREMENTS AND QUALITATIVE recognized for the quantitative parameters (Tables 4 CHARACTERS ANALYSED and 5). The crude matrix is presented as Supporting We used 20 descriptive characters relating to seed. Of Information (Appendix S2). these, three are quantitative, two are allometric relationships, one is based on totalized dimensions (prism volume, defined as the length, width and thickness of DATA ANALYSES the seed) and 14 are qualitative. Recognized states of The crude matrix was used to compute a dissimilarity qualitative characters totalled 41. In selecting the matrix using Darwin 5 V.5.0.158 (2009-07-06) character set, and the states thereof, we took into (Perrier, Flori & Bonnot, 2003; Perrier &

© 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175, 74–122 82 .RIVERA D. Table 3. List of fossil and archaeobotanical Phoenix seed samples. NSEE, number of seeds analysed

Date BP Country Main area Code (years) Site Loc Materials NSEE Reference

USA North Eocene_TEX 40 000 000 Catahoula Texas Phoenicites occidentalis Berry. Type: 1 Berry, 1914 AL ET

04TeLnenSceyo London, of Society Linnean The 2014 © America formation collected by Chas. Laurence Baker

in Trinity County, Texas. From a . cut on the International and Great Northern Railroad in southern Trinity county. Mineralized Germany Europe Eocene_GEIS 35 000 000 Tagebau Geiseltal Phoenix hercynica Mai. One seed, 1 Mai, 1976 Neumark- Upper Eocene. Mineralized Süd Germany Europe Eocene_ 35 000 000 Tagebau Geiseltal Published as Serenoa carbonaria 4 Mai, 1976 SERENCAR_1 Neumark- Mai. Over 100 seeds, Upper Süd Eocene. Mineralized Czechia Europe Miocene_ 20 000 000 Tuchorice Bohemia Mineralized date-palm seeds from 10 Buzek, 1977 BOHEM1-18 the freshwater limestones, and Phoenix bohemica Buzek. Whole oaia ora fteLnenSociety Linnean the of Journal Botanical BOHEMTYPE gathering and the type specimen were analysed as two separate subsamples Greece Europe Pleistocene_ 37 000 Fira Santorini Impression of a fruit (seed 1 Friedrich, 1980; FIRA Palaeosol dimensions are inferred) from Friedrich et al., 1977 Fira Paleosol at Tera, Santorini dated Weichselian Interstadials Egypt North Africa Pleistocene_ 16 000 Kharga Kharga The occurrence of the loam beds, 4 Caton & Gardner, KHARGA1-2 Oasis containing carbonized reed stems, 1932; Gardner, 1935 and yielding also fruit seeds of a wild date (identiﬁed by Mrs Clement Reid as Phoenix sylvestris (L.) Roxb.). Carbonized. Images in Gardner (1935: Tab 32, 7–8) Fruit from mound spring 2014, , near Kharren Pakistan Central Asia Neolithic_ 8000 Mehrgarh IB Balochistan One mineralized date seed 1 Beech, 2003;

175 MEHRG1 Costantini, 1985 Pakistan Central Asia Neolithic_ 7000 Mehrgarh Balochistan One mineralized date seed 1 Beech, 2003; 74–122 , MEHRG2 IIB Costantini, 1985 04TeLnenSceyo London, of Society Linnean The 2014 ©

Libya North Africa Neolithic_LIBY1 8000 Takarkori, Fezzan, Two well-preserved, desiccated date 2 Anna Maria Mercuri, and 2 Central seeds: 1_U32 L286 camp 51 house researcher at the Sahara 59. 2_T33 L268 University of Modena and Reggio Emilia, Italy (pers. comm.) Kuwait West Asia Neolithic_SABI1 7530 Site H3 A Sabiyah Dump layers within chambers 1, 11 3 Beech, 2003 to 3 (west), and 18. A total of three context mineralized date seeds 1029 (19.7 × 8.8 × 8.3 mm, 22.5 × 9.5 × 9.4 mm, 15.0 × 7.7 × 6.4 mm) from there were hard and seed-like because

oaia ora fteLnenSociety Linnean the of Journal Botanical of the process of mineralization Israel West Asia Neolithic_ 7500 Atlit-Yam Atlit-Yam One waterlogged date seed. 1 Kislev, Hartmann and ATLITYAM Pre-Pottery Neolithic C (PPNC) Galili, 2004 Iran West Asia Chalcolithic_ 7400 Tepe Gaz Dowlatabad One carbonized seed was recovered 1 Beech, 2003; Nesbitt, TEPEGAZ Tavila from Dowlatabad plain, south of 1993; Costantini, Kerman, eastern Iran 1985 United Arab West Asia Chalcolithic_ 7120 DA11 Dalma Context 15 (a burnt layer located 1 Beech, 2003 Emirates DAL11_1 Island c. 80 cm below the present day ground surface and 25 cm above the ﬂoor level of one of the house structures: 5120 ± 170 calibrated APLGCLAAYI OF ANALYSIS CARPOLOGICAL BC) carbonized date stone United Arab West Asia Chalcolithic_ 7000 DA11 Dalma Context 15 mud brick cast 1 Beech, 2003 Emirates DAL11_2 Island United Arab West Asia Chalcolithic_ 7000 DA11 Dalma Context 15 mud brick cast 1 Beech, 2003

2014, , Emirates DAL11_4 Island United Arab West Asia Chalcolithic_ 6670 DA11 Dalma Context 4 (a redeposited sand layer 1 Beech, 2003 Emirates DAL11_3 Island just below the present day ground 175 surface: 4670 ± 130 cal BC)

74–122 , fragmentary carbonized date stone Pakistan Central Asia Chalcolithic_ 6000 Miri Qalat Makran Carbonized date seeds, from 3 Tengberg, 1999 MIRI_1 ﬂotation samples, very small compared with those from other archaeological sites. Two date seeds were found from Period II, PHOENIX three from Period IIIa and 22 from Period IV Palestine West Asia Chalcolithic_ 5600 Teleilat Carbonized date seeds 6 Nesbitt, 1993; Hopf, TELEILAT_ Ghassul 1983; Levy, 1986 1–6 83 84 .RIVERA D. Table 3. Continued

Date BP Country Main area Code (years) Site Loc Materials NSEE Reference TAL ET Iran West Asia Bronze 5000 Tell Karrana Tell One desiccated date seed 1 Costantini &

04TeLnenSceyo London, of Society Linnean The 2014 © Age_KARR_1 3 R16 Karrana (19.72 × 7.81 × 7.58 mm) was Costantini-

found on the ﬂoor in the area R16 Biasini, 1993 . of Bronze Age beds of Tell Karrana 3 Israel West Asia Bronze 4600 Jericho Jericho One desiccated date seed 1 Hopf, 1983 Age_JERICHO Iraq West Asia Bronze 4500 Queen Ur Many fragments of desiccated date 4 Beech, 2003; Renfrew, Age_UR_1–4 Pu-abi’s seed were found in Queen Pu-abi’s 1987 grave grave at Ur; only a few were complete enough to measure Oman West Asia Bronze Age_ 4500 Hili 8 IIC Al-Ain A high number of carbonized date 5 Costantini, 1985; HILI8_1–6 stones were recovered Fuller & Madella, 2001 Oman West Asia Bronze Age_ 4300 6016 QIK, Ra’s Carbonized Phoenix dactylifera 39 Costantini & Audisio, RASALJIZ_ 6018 QIP, al-Jinz fruits and seeds and Zizyphus 2000 oaia ora fteLnenSociety Linnean the of Journal Botanical 1–39 6019 RJ-3 fruits QHO-QIK, 6020 QHO, etc. Kuwait West Asia Bronze Age_ 4000 1129.AQS Failaka Carbonized date seeds 3 Rowley-Conwy, 1987 FAILAKA1-3 cited by Beech, 2003; Nesbitt, 1993; Willcox & Tengberg, 1995 Yemen West Asia Bronze Age_ 4000 Ar Raqlah Ar Raqlah Two casts of date seed were found 2 Costantini, 1991; ARRAQL1-2 in pottery (c. 10.7 mm long and Nesbitt, 1993 10.09 × 6.88 mm) Bahrain West Asia Bronze 3700 SAAR 1991 Saar The variation in length 16 Nesbitt, 1993 Age_SAAR1-13 E16:10:05 (10.40–19.50 mm) is particularly

2014, , and 14–16 striking and it is possible that, once large numbers of seeds are available for measurement,

175 clusters of different sizes

74–122 , conforming to varieties may become apparent. Dilmun. Carbonized 04TeLnenSceyo London, of Society Linnean The 2014 ©

Yemen West Asia Iron 2800 SI (N = 25), Raybun Phoenix seeds (20–22 × 8.2–9.3 mm 1 Levkovskaya & Age_RAYBU_1 S2 and 13–17 × 5–9 mm), pollen and Filatenko, 1992 to 12 (N = 15), leaf remains were found in the S3 site of Raybun (South Arabia). (N = 30), R Desiccated to mineralized (N = 50), Iran West Asia Neo-Elamite_ 2650 Burial Susa Dates seem to have been used as a 2 Miller, 1981 SUSA_1–2 693–6861 funerary offering in the Neo-Elamite burial 693. oaia ora fteLnenSociety Linnean the of Journal Botanical Desiccated Saudi Arabia West Asia Iron 2600 Oasis Tayma Tayma Subfossil desiccated seeds 4 Neef, Cappers & Age_TAYMA_1 Bekker, 2011 Israel West Asia Roman_MASSA_ 2120 Masada Southern Phoenix dactylifera desiccated fruits. 4 Sallon et al., 2008 1–4 District At least one seed was able to germinate Iran West Asia Parthian_SUSA 2050 Susa Shush One desiccated date seed 1 Miller, 1981 Italy Europe Roman_ 2000 Pompeii Naples House of the Ship Europa, a single 1 Meyer, 1980 POMPEI_4a carbonized seed Italy Europe Roman_ 1990 Pompeii Naples Museum inventory 50. 84 630 date 1 Wittmack, 1903 APLGCLAAYI OF ANALYSIS CARPOLOGICAL POMPEI_4b fruits 33 × 12 mm and one seed 22×8mm France Europe Roman_LATT 1950 Lattara Lattes Among the fruits and seeds burnt in 1 Rovira & Chabal, 2008 this offering are: Phoenix

2014, , dactylifera ﬁve seeds and eight fruits. Carbonized Egypt North Africa Roman_ 1950 Karanis Kom Subfossil desiccated fruits and seeds 18 Neef et al., 2011 175 KARANIS1-9 Aushin

74–122 , Spain North Africa Guanche_ 1200 Garajonay Garajonay Religious offering, four desiccated 5 Morales et al., 2011 GARAJONAY5 seeds. Guanche period Mali Tropical Middle 800 GAD 96(A) Gao One desiccated date seed 1 Fuller, 2000 Africa Ages_GAO

We should note that the comparison of carbonized materials (shaded in grey in the table) with modern materials is pending as there are serious doubts concerning PHOENIX the preservation of morphological features during charring. We are currently preparing for comparison a set of experimentally carbonized seeds of different species and cultivars. 85 86 D. RIVERA ET AL.

Table 4. Quantitative parameters analysed in modern and archaeological and fossil Phoenix seed samples. FosAr, fossil and archaeological materials; B, breadth; D, depth; L, length; TD, totalized dimensions a. Quantitative characters (continuous)

Total Modern FosAr

Parameters Min Max Min Max Min Max

L (mm) 4.47* 40.00 4.47 37.83 5.50 40.00 B (mm) 1.31* 15.00 1.31 12.87 3.20 15.00 D (mm) 0.95* 16.89 0.95 12.87 3.20 16.89 B/L 0.09 1.17 0.09 1.17 0.23 0.70 D/B 0.32 1.50 0.32 1.50 0.46 1.33 TD (mm3) 36.62* 10 134.00 36.62 3656.83 56.32 10 134.00

*These values correspond to abnormal seeds not fully developed. b. Quantitative characters in states or classes. Data in percentage of samples analysed. Dimensions in mm. B, breadth; D, depth; L, length

Type L 4–10 L 10–15 L 15–19 L 19–25 L 25–32 L 32–40

Modern 10.3 23.1 22.7 30.0 13.0 0.8 FosAr 2.5 24.3 16.5 44.9 5.6 2.8

Type B 1–3.5 B 3.5–6 B 6–8 B 8–10 B 10–12 B 12–16

Modern 1.4 12.1 25.9 43.5 16.4 0.6 FosAr 0.6 12.2 47.2 30.3 4.7 1.7

Type D 0–2.5 D 2.5-4 D 4–5.5 D 5.5–7 D 7–17

Modern 1.1 5.5 10.9 24.1 58.1 FosAr 0.0 0.6 20.1 48.3 27.6 c. Allometric characters and totalized dimensions in states. Data in percentage of samples analysed. B, breadth; D, depth; L, length; TD, totalized dimensions. Totalized dimensions in mm3.

Type B/L 0–0.2 B/L 0.2–0.4 B/L 0.4–0.6 B/L 0.6–0.8 B/L 0.8–1

Modern 1.0 26.9 56.4 14.4 1.3 FosAr 0.0 42.9 48.0 5.3 0.4

Type D/B 0–0.75 D/B 0.75–0.85 D/B 0.85–0.95 D/B 0.95–1.5

Modern 8.5 32.7 52.6 6.2 FosAr 16.7 20.2 32.6 27.2

Type TD 0–150 TD 150–300 TD 300–800 TD 800–1 200 TD 1 200–1 850 TD 1 850–10 200

Modern 4.7 6.9 19.9 19.4 33.9 15.3 FosAr 1.1 4.2 38.2 28.1 17.5 7.7

Table 5. Qualitative characters analysed in modern and archaeological and fossil Phoenix seed samples. Data in percentage of samples analysed. FosAr, fossil and archaeological materials; OC, ovate–triangular; EL, elliptic; OB, oblong; CY, cylindric; GL, globose; HE, hemispherical; FU, fusiform; AO, apex obtuse; AA, apex acute; AR, apex retuse; AB, apex oblique; AT, apex truncate; BO, base obtuse; BA, base acute; BL, base oblique; BT, base truncate

Shape OC EL OB CY GL HE FU

Modern 5.94 20.25 44.88 19.45 1.09 1.39 7.01 FosAr 3.68 16.56 28.22 50.92 0.61 0.00 0.00

Colour Blackish Greyish Cream Brown

Modern 1.20 2.05 37.03 59.90 oaia ora fteLnenSociety Linnean the of Journal Botanical Apex AO AA AR AB AT Base BO BA BL BT

Modern 83.87 12.69 0.19 0.18 2.87 Modern 44.17 15.29 3.97 36.27 FosAr 89.57 4.91 0.00 0.00 5.52 FosAr 50.31 29.45 6.13 14.11

Apical mucro Present Lacking Basal mucro Present Lacking

Modern 12.5 87.5 Modern 6.1 93.9 FosAr 0.0 100.0 FosAr 1.5 98.5 APLGCLAAYI OF ANALYSIS CARPOLOGICAL

Surface Glossy and smooth Rough and matt Longitudinal grooves Present Lacking Transverse processes Wrinkled Finely grooved Uniform

Modern 44.8 55.2 Modern 27.14 72.86 Modern 34.34 8.39 57.27

2014, , FosAr 34.36 65.64 FosAr 10.43 89.57 FosAr 57.06 1.23 41.72

Micropyle Central Basal Raphe* Shallow V-shaped U-shaped 175

74–122 , Modern 97.5 2.5 Modern 13.53 33.06 53.41 FosAr 100.0 0.0 FosAr 41.10 23.31 35.58

*Lengthwise ventral furrow.

Dorso-ventral Lengthwise crests or PHOENIX curvature Bent Straight wings Present Lacking

Modern 5.01 94.99 Modern 2.4 97.6 FosAr 0.0 100.0 FosAr 0.0 100.0 87 88 D. RIVERA ET AL.

Jacquemoud-Collet, 2006). The χ2 dissimilarity index second analysis was performed incorporating com- was calculated. This measure expresses a value xik as mercial samples of dates, and seeds intended for use its contribution to the sum xi on all variables and is a in horticulture and gardening totalling 303 units or comparison of unit profiles: samples and 67 variables or descriptors. This analysis also included several samples of immature seeds from K ⎛ ⎞ 2 dates abnormally ripened. In parallel, fossil and =−x.. xik x jk dij ∑ ⎜ ⎟ archaeological seed samples were analysed totalling x ⎝ x x ⎠ 1 ..k i j . 60 units and 52 variables. Excluded from this analy- where dij is the dissimilarity between units i and j; xik sis were variables or descriptors for which all of the and xjk are the values of variable k for units i and j; samples had zero frequency, i.e. colour of the seed. xi., xj. and x.k are the mean for units i and j or variable Finally, modern and fossil and archaeological samples k; x. is the overall mean. K is the number of variables. were included in a fourth analysis, with 364 units Dissimilarities are even and Euclidean distances. that are seed samples (Table 1 and Supporting Infor- Principal coordinates analysis (PCoA), which works mation, Appendix S1) and 67 descriptors. on dissimilarity matrices showing the distance between every possible pair of samples, was used to give an overall representation of diversity within RESULTS Phoenix seeds with the lowest possible dimensional space. This represented a step in the analysis of the SEQUENTIAL ORDER OF THE ANALYSIS structure of diversity in the genus Phoenix. The first analysis, which included only herbarium To represent individual relationships realistically, a type material, type icons and botanically verified hierarchical tree was constructed to describe the rela- samples collected in the field and in repositories, tionships between units (samples) based on the produced seven main clusters reflected in the hierar- common agglomerative heuristic that proceeds by chical phenetic tree (Fig. 1). A first branching sepa- successive ascending agglomerations. For updating rates outgroups (Euterpe, Livistona, Nannorrhops, dissimilarity during the tree construction, the Ward Washingtonia), and P. paludosa, from the seeds of criterion was adopted, which searches at each step for other Phoenix spp. Then two groups are clearly sepa- a local optimum to minimize the within-group or, rated depending on seed size. The first group, with equivalently, to maximize the between-group inertia. large seeds, includes numerous samples of P. dactyl- The distance between two elements is the weighted ifera (including the type icon) and seeds of P. atlantica square of the Euclidean distance between their (including original material collected by Chevalier), gravity centres. P. sylvestris (including the type icon), P. canariensis A weighted neighbor joining tree was used to verify var. macrocarpa H.Wildpret (material collected by the close similarities between samples. The neighbor author) and the nomenclatural type of P. iberica. The joining method proposed by Saitou & Nei (1987) uses second group includes samples from all other Phoenix the criterion of relative neighbourhood, weighted spp. Then the minute and thin seeds of P. roebelenii average for dissimilarity updating and adjustment to are separated, followed by P. rupicola and P. canar- an additive tree distance. A bootstrap value is given to iensis. A low-resolution group brings together the each edge that indicates the occurrence frequency of seeds of P. theophrasti, P. reclinata Jacq., P. caespitosa this edge in the bootstrapped trees. Bootstrap values (P. arabica) and P. pusilla. Samples of the palm with range between 0 and 100. Radial trees were drawn bluish fruits, named P. senegalensis André, fall within using Dendroscope (Huson & Scornavacca, 2012) and the groups of P. theophrasti–P. caespitosa (P. arabica) FigTree (Rambaut, 2012). and of P. pusilla–P. reclinata. In parallel, there is To verify stability of the results and to develop another group with the seeds of P. loureiroi Kunth, methodologies for allocating to morphotypes gener- P. acaulis (including the type icon), P. andamanensis ated in function of species, cultivars and archaeologi- S.Barrow (nomenclatural type) and the type icon of cal and fossil seed samples, the above methodology P. farinifera Roxb. was conducted in four stages. First, similarity was A second analysis including commercial seed calculated using only a sub-matrix formed by 165 samples resulted in an overall structure similar to the units or samples, which included the herbarium above, clearly separating P. dactylifera and related specimens, types and those samples collected in the taxa from the group of smaller seeded species. field, or from repositories, that were botanically iden- However, some major changes may be noticed. The tified, taking into account other characters of the samples attributed to P. theophrasti collected in Datça palm and 63 variables or descriptors. Excluded from and Gölköy (south-western Turkey) appear here clus- this analysis were variables or descriptors for which tered with different P. dactylifera samples (with rela- all of the samples had zero frequency. After that, a tively small seeds) from Spain and Baja California

Figure 1. Hierarchical tree calculated using the algorithm of Ward with Phoenix type specimens and botanically veriﬁed modern seed samples. Branchlets labelled gray correspond to Phoenix dactylifera.

(Mexico) and no longer with those of P. theophrasti in south-eastern Spain. Immature seeds, from unripe from Crete. Again, seeds of P. andamanensis fall fruits, or sterile seeds are abnormally small or thin, within the variability of P. loureiroi. Eleven horticul- and a group of these clusters among the samples of tural samples of P. sylvestris cluster around the type P. roebelenii and of hybrids of this with other species. of the species on a branch that contains numerous The third analysis was performed exclusively with P. dactylifera samples from Elche and other localities fossil and archaeobotanical seed samples. Obviously

© 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175, 74–122 90 D. RIVERA ET AL. the depositional and post-depositional processes near the middle of the dorsal face. Ventral raphe reduced the number of variables or descriptors avail- U-shaped (91%). Dorsoventrally straight. Not winged. able from 67 to 52. The resulting weighted neighbor Group 2: includes small seeds of P. acaulis from joining tree shows relationships with > 50% coinci- India and the type icon of the species. Within this dence of the 5000 bootstraps (Fig. 2). Surprisingly, group are also the type icons of P. zeylanica Trimen there is no clear separation between fossil and and P. pusilla Gaertn. Main descriptive parameters: archaeological samples, between different periods of Breadth/Length = 0.4–0.6 (86%). Length = 10– the archaeological samples, between geographical 15 mm (87%). Breadth = 3.5–8.0 mm. Depth = 4– origins or between forms of preservation. Percentages 7 mm. Totalized dimensions = 300–800 mm3. > 75% of coincidence are found only between seeds Oblong. Cream coloured (71%) or brown (29%). Apex from the same site, except for Bronze Age seed samples obtuse. Base obtuse (86%). Surface rough (71%), from Ras al Jinz (Oman) and Failaka (Kuwait) (both uniform. With longitudinal grooves (57%). Micropyle carbonized). near the middle of the dorsal face. Ventral raphe In the fourth analysis, with 364 samples including V-shaped (29%) or U-shaped (71%). Dorsoventrally fossil and archaeobotanical materials, PcoA and hier- bent (82%). Not winged. archical clustering using Ward’s algorithm (Fig. 3A–C) Group 3: includes small seeds of P. loureiroi from show four main clusters that include 24 groups and one south-eastern China, P. reclinata hybrids and two outgroup. Furthermore, at least eight species have samples from cultivated palms tentatively identiﬁed characteristic seeds and are clearly assigned to mor- as P. caespitosa. And one horticultural seed sample photypes [P. acaulis, P. canariensis s.s., P. paludosa, labelled as P. canariensis. Main descriptive param- P. reclinata, P. roebelenii, P. rupicola, P. sylvestris and eters: Breadth/Length = 0.6–0.8 (75%). Length = 10– P. theophrasti (excluding populations of Datça, Turkey) 15 mm (75%). Breadth = 6–8 mm (77%). Depth = 5.5– (Fig. 3, Table 6)], the rest are not clearly separated on 12.0 mm. Totalized dimensions = 300–800 mm3 (90%). the sole basis of the morphology of seeds. Elliptic (95%). Cream coloured (40%) or brown (60%). Characters and states that provide information to Apex obtuse. Base obtuse (72%) or truncate (28%). separate different groups behave differently. Some are Surface rough (63%) or smooth (37%), uniform. With infrequent: hemispheric seeds, seeds blackish, width longitudinal grooves (89%). Micropyle near the to 3.5 mm, B/L to 0.1 or, conversely, 0.8 to 1.0, and middle of the dorsal face. Ventral raphe U-shaped. serve to differentiate isolated groups, whereas others Dorsoventrally straight. Not winged. such as the U-shaped opposed to the V-shaped ventral Group 4: includes small seeds of P. reclinata, com- raphe are recurrent for differentiation of various prising the type icon of the species. It also include groups. Within each group, those states which are several Bronze Age archaeological samples from relevant for describing the group are marked in bold. Arraqis, Jericho, Saar and Hili, and Roman samples Frequency within the seeds of each group is presented from Karanis, and a sample of a bluish date known as as a percentage between parentheses. Percentages P. senegalensis. Main descriptive parameters: > 95% are not shown (see Figs 3A, 4). In the Support- Breadth/Length = 0.4–0.6 (74%) or 0.6–0.8 (24%). ing Information (Appendix S3) is attached the matrix Length = 10–15 mm. Breadth = 6–8 mm (91%). of correlation between variables. Depth = 5.5–7.0 mm. Totalized dimensions = 300– 800 mm3 (97%). Oblong (37%), Ovoid–triangular (24%). Brown coloured (modern seeds). Apex obtuse. CLUSTER I Base obtuse (67.5%) or truncate (30%). Surface rough This cluster includes small seeds, short (length usually (50%) or smooth (50%), uniform (68.75%) or wrinkled < 15 mm), with totalized dimensions from c. 150 to (31.25%). Without longitudinal grooves. Micropyle 800 mm3 and surface generally even, uniform. near the middle of the dorsal face. Ventral raphe Group 1: includes small seeds of different subspe- V-shaped (34%) or U-shaped (66%). Dorsoventrally cies of P. loureiroi from India and East Asia, with the straight. Not winged. nomenclatural type of P. andamanensis (which only Group 5: includes small seeds of P. theophrasti s.s. differs in the ruminate endosperm), several misla- (several samples from Crete and one from Gölköy) with belled commercial samples and one interspeciﬁc a Neolithic sample from Atlit Yam (Israel) and several hybrid. Main descriptive parameters: Breadth/ archaeological samples. Main descriptive parameters: Length = 0.4–0.6 (84%). Length = 10–15 mm (87%). Breadth/Length = 0.4–0.6. Length = 10–15 mm. Breadth = 3.5–8.0 mm (94%). Depth = 4–7 mm. Breadth = 3.5–8.0 mm (91%). Depth = 5.5–12.0 mm. Totalized dimensions = 300–800 mm3 (82%). Elliptic Totalized dimensions = 300–800 mm3 (70%) or 800– (32%) to oblong (63%). Cream coloured (83%). Apex 1200 mm3 (25.5%). Elliptic (63.6%), oblong (20.5%). obtuse (93%). Base obtuse (72%). Surface rough (69%), Brown coloured (modern seeds). Apex obtuse. Base uniform. With longitudinal grooves (72%). Micropyle obtuse. Surface smooth, uniform. With longitudi-

Figure 2. Tree calculated with the weighted neighbor joining algorithm. A, archaeological desiccated and brick casts (red) and fossil Phoenix seed samples (blue). B, carbonized seeds (black). Bootstrap values below 30% are omitted.

Figure 3. Hierarchical tree calculated using the algorithm of Ward with Phoenix type specimens and botanically veriﬁed modern seed samples, commercial samples and archaeological and fossil Phoenix seed samples. A (page 93). Cluster I. Small Phoenix species from India and East Asia. Cluster II. Phoenix roebelenii from Mekong river basin. Cluster III. Thin seeds of Phoenix dactylifera from the Near East and North Africa (p.p.). B (page 94). Cluster III. Thin seeds of Phoenix dactylifera from the Near East and North Africa (p.p.). Cluster IV. Phoenix dactylifera cultivars, and P. atlantica (p.p.). C (page 95). Cluster IV. Phoenix dactylifera cultivars, P. sylvestris, P. iberica, P. canariensis and P. atlantica (p.p.). Cluster V. Phoenix paludosa. Cluster VI. Outgroup. p.p. = pro parte. ▶

nal grooves (81.8%). Micropyle near the middle of the raphe shallow (75%). Dorsoventrally straight. Not dorsal face. Ventral raphe shallow. Dorsoventrally winged. straight. Not winged. Group 6: includes small seeds of P. pusilla, the type icon of P. farinifera [according to Barrow, (1998) CLUSTER II and Govaerts et al. (2011) a synonym of P. pusilla] This cluster includes small seeds of P. roebelenii and and seeds of P. loureiroi (P. hanceana Naudin) and immature seeds of other species with totalized dimen- P. roebelenii hybrids. It also includes several archaeo- sions < 300 mm3. logical samples and Eocene fossils from Geiseltal See Figures 3A and 4. named Serenoa carbonaria (Mai, 1976). Main descrip- Group 9: almost exclusively includes small imma- tive parameters: Breadth/Length = 0.4–0.8 (78%). ture seeds of P. caespitosa and P. reclinata. It also Length = 4–15 mm. Breadth = 3.5–6.0 mm (85.4%). includes one archaeological sample. Main descriptive Depth = 4.5–5.5 mm (80%). Totalized dimen- parameters: Breadth/Length = 0.4–0.6 (76%) or 0.6– sions = 150–300 mm3 (71.2%) or 300–800 mm3 0.8 (24%). Length = 4–10 mm. Breadth = 3.5– (25.1%). Elliptic (48.5%), oblong (25%). Cream (50%) 6.0 mm (77.6%). Depth = 2.5–4.0 mm (86.7%). or brown (25%) coloured (modern seeds). Apex obtuse. Totalized dimensions = 0–150 mm3 (89.3%). Ellip- Base obtuse (66.7%) or truncate (33.3%). Surface tic (80%). Brown coloured. Apex obtuse (80%). Base smooth (75%) or rough (25%), uniform. Without obtuse (60%) or truncate (40%). Surface smooth (80%) longitudinal grooves (66.7%). Micropyle near the or rough (20%), uniform (80%) or wrinkled (20%). middle of the dorsal face. Ventral raphe U-shaped Without longitudinal grooves. Micropyle near the (83%). Dorsoventrally straight. Not winged. middle of the dorsal face. Ventral raphe U-shaped. Group 7: exclusively includes small seeds of P. rupi- Dorsoventrally straight. Not winged. cola. Main descriptive parameters: Breadth/ Group 10: exclusively includes small seeds of P. roe- Length = 0.4–0.6. Length = 10–15 mm (75%) and belenii (from East Asia) and hybrids with other species. 15–19 mm (25%). Breadth = 6–8 mm (85.8%). Main descriptive parameters: Breadth/Length = Depth = 5.5–7.0 mm (82.7%). Totalized dimen- 0.4–0.6. Length = 4–10 mm. Breadth = 3.5–6.0 mm. sions = 300–800 mm3 (84.9%) or 800–1200 mm3 (11%). Depth = 2.5–4.0 mm (91.8%). Totalized dimen- Oblong. Greyish coloured. Apex obtuse. Base sions = 0–300 mm3. Oblong. Cream (60%) or Brown obtuse (66.7%) or truncate (33.3%). Surface rough, (40%) coloured. Apex obtuse. Base truncate. uniform. With longitudinal grooves. Micropyle near Surface smooth, uniform. Without longitudinal the middle of the dorsal face. Ventral raphe shallow grooves. Micropyle near the middle of the dorsal face. (83%). Dorsoventrally bent (90%). Not winged. Ventral raphe U-shaped. Dorsoventrally straight. Group 8: includes small seeds of P. canariensis cul- Not winged. tivars, one sample of P. caespitosa (P. arabica) and Group 11: exclusively includes thin immature archaeological samples from the Guanche Period seeds of P. dactylifera. Main descriptive parame- (Canary Islands, Spain) and Roman Karanis. Main ters: Breadth/Length = 0–0.4. Length = 15–32 descriptive parameters: Breadth/Length = 0.4–0.8. mm. Breadth = 1.0–3.5 mm (83.1%). Depth = 0– Length = 10–15 mm (75%) and 15–19 mm (25%). 2.5 mm (77.5%). Totalized dimensions = 0– Breadth = 6–8 mm (54%) and 8–10 mm (38%). 150 mm3 (80.6%). Fusiform (50%), cylindric (25%) Depth = 5.5–7.0 mm (60%) and 7.0–12.0 mm (40%). or oblong (18.8%). Brown coloured. Apex acute Totalized dimensions = 300–800 mm3 (56%) or 800– (75%). Base obtuse (60%) or truncate (40%). Surface 1200 mm3 (44%). Elliptic (30%), oblong (45%). Brown smooth (80%) or rough (20%), wrinkled (75%) or coloured. Apex truncate. Base truncate (80%). uniform (25%). Without longitudinal grooves. Surface smooth (50%) or rough (50%), uniform (90%) or Micropyle near the middle of the dorsal face. wrinkled (10%). With longitudinal grooves (63%). Ventral raphe U-shaped. Dorsoventrally straight. Not Micropyle near the middle of the dorsal face. Ventral winged.

3 Cluster I (Groups 1 to 8)

8 Ie

9 Cluster II (Groups 9 to 11) 10

Cluster III (Group 12)

Figure 3. See caption on previous page.

Cluster III (Groups 12 to 13)

Cluster III (Groups 14 to 17)

18 Cluster IV (Group 18)

Figure 3. Continued

Cluster IV (Groups 18 to 23)

Cluster V (Group 24) 24

Cluster VI (Group 25) 25

Figure 3. Continued

© 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175, 74–122 96 .RIVERA D. TAL ET Table 6. Summary of the seed morphology of Phoenix species (excluding P. dactylifera and its hybrids). Morph, morphotype 04TeLnenSceyo London, of Society Linnean The 2014 ©

P. P. . Characters/ P. P. pusilla P. pusilla P. theophrasti P. theophrasti canariensis canariensis states P. loureiroi andamanensis P. acaulis A B P. reclinata A Type Datça P. rupicola A B Type P. caespitosa P. atlantica P. iberica P. sylvestris P. roebelenii P. paludosa

B/L 0.4–0.8 0.4–0.6 0.4–0.8 0.4–0.6 0.4–0.8 0.4–0.8 0.4–0.8 0.6–0.8 0.4–0.6 0.4–0.8 0.6–0.8 0.4–0.8 0.4–0.8 0.4–0.8 0.4–0.8 0.4–0.6 0.6–1 L mm 4–15 10–15 4–15 10–15 4–15 10–15 10–19 10–19 10–19 10–19 10–19 10–15 15–25 15–25 15–25 4–10 4–10 B mm 3.5–8 6–10 3.5–8 6–8 3.5–8 6–8 6–10 6–10 6–10 6–10 6–10 6–10 6–10 8–10 8–12 6–10 3.5–8 D mm 4–7 4–5.5 4–7 5.5–7 2.5–5.5 4–7 5.5–12 5.5–12 5.5–12 5.5–12 7–15 5.5–7 5.5–12 7–15 7–15 2.5–5.5 2.5–5.5 TD mm3 150–800 300–800 300–800 300–800 150–800 300–800 300–1200 300–1200 300–1200 300–1200 800–1850 300–800 800–1850 800–2500 800–2500 100–300 150–800 Outline Elliptic, Oblong Oblong Oblong Elliptic, Ovate– Elliptic Oblong Oblong Elliptic, Elliptic Elliptic Ovate- Elliptic Ovate- Oblong Hemispheric oblong oblong triangular, Oblong triangular, triangular, elliptic, oblong elliptic, oblong oblong Colour Cream Cream Cream Brown Brown Brown Brown Brown Greysh Brown, Brown, Brown Brown Brown Cream Cream, Blackish, cream cream brown greyish Apex Obtuse Obtuse, Obtuse Obtuse Obtuse Obtuse Obtuse Obtuse Obtuse Truncate Obtuse Truncate, Obtuse Obtuse Obtuse Obtuse Obtuse

oaia ora fteLnenSociety Linnean the of Journal Botanical acute obtuse (acute) Base Obtuse, Obtuse Obtuse Truncate, Obtuse Truncate, Obtuse Obtuse Obtuse Obtuse Obtuse Truncate Obtuse Obtuse Truncate Truncate Obtuse truncate (acute) obtuse obtuse (truncate) (truncate) (truncate) (truncate) (obtuse) Surface Rough Rough Rough Rough, Smooth Smooth Smooth Smooth Rough Rough Smooth Rough Rough Rough Rough Smooth Smooth (smooth) (smooth) smooth (rough) (smooth) (smooth) Transverse Uniform Uniform Uniform Uniform Uniform Uniform Uniform Uniform Uniform Uniform Uniform Uniform Uniform Uniform Uniform Uniform Uniform processes (wrinkled) Longitudinal Frequent Always Always Not Frequent Not Always Always Always Always Always Always Not Sometimes Frequent Not Not grooves Micropyle Central Central Central Central Central Central Central Central Central Central Central Central Central Central Central Central Basal Ventral U-shaped U-shaped U-shaped U-shaped U-shaped U-shaped Shallow V-shaped Shallow U-shaped U-shaped Shallow U-shaped U-shaped U-shaped U-shaped Shallow furrow (V-shaped) (shallow) (V-shaped) (V-shaped) (V-shaped) Dorso-ventral Not Not Frequent Always Not Not Not Not Frequent Not Not Not Sometimes Not Not Not Not curvature Wings Not Not Not Not Not Not Not Not Not Not Not Not Not Not Not Not Not Morph/ 1 1 2 2 6 4 5 18 7 8 21 3 18 19 19 10 24 group 2014, , 175 74–122 , CARPOLOGICAL ANALYSIS OF PHOENIX 97

CLUSTER III P. atlantica. It also includes numerous archaeological This cluster includes modern, archaeological and samples (Neolithic to Middle Ages) and Eocene fossils fossil samples that fall within the variability of from Geiseltal named P. hercynica (Mai, 1976). Main P. dactylifera. See Figures 3A, B, 4 and 5. descriptive parameters: Breadth/Length = 0.2–0.4 Group 12: includes large seeds of P. dactylifera cul- (81.6%). Length = 15–25 mm (84%). Breadth = 6– tivars from Spain (40 samples), Baja California 10 mm (73%). Depth = 5.5–12.0 mm (83.7%). Total- 3 (Mexico) (10) and the Near East (10). It also includes ized dimensions = 300–1850 mm . Oblong (17.9%), two archaeological samples from Roman Karanis. cylindrical (59.1%) or fusiform (23.1%). Cream Main descriptive parameters: Breadth/Length = 0.2– (20.8%) or brown (37.5%) coloured (modern seeds). 0.6. Length = 19–32 mm. Breadth = 8–12 mm. Apex obtuse (37.8%) or acute (62.2%). Base truncate Depth = 7–12 mm (82.9%). Totalized dimen- (15.3%), obtuse (8.3%), acute (76.7%). Occasionally sions = 1200–2500 mm3. Oblong (20.5%), cylindrical mucronate (17.5%). Surface smooth (33.3%) or rough (54.5%), fusiform (20.3%). Cream (16.8%) or Brown (66.7%), wrinkled (70.8%), or uniform (29.2%). (80%) coloured. Apex obtuse (76.7%) or acute (20%). Without longitudinal grooves. Micropyle near the Sometimes (30%) mucronate. Base truncate (49.4%), middle of the dorsal face. Ventral raphe V-shaped obtuse (22.6%), acute (23.7%). Surface smooth (21.4%) (16.7%), U-shaped (25%) or shallow (58.3%). Dors- or rough (78.6%), wrinkled (36.4%), ﬁnely grooved oventrally straight. Not winged. (40.6%) or uniform (23%). Without longitudinal Group 15: exclusively includes thin seeds of P. dac- grooves. Micropyle near the middle of the dorsal tylifera (from the Near East) and several archeologi- face. Ventral raphe V-shaped (60.5%) or U-shaped cal samples from the Neolithic to Roman period. Main (33%). Dorsoventrally straight. Occasionally winged descriptive parameters: Breadth/Length = 0.2–0.4 (6%). (81.6%). Length = 19–25 mm (94.3%). Breadth = 6– Group 13: includes the largest seeds of P. dactylif- 8mm (90.5%). Depth = 5.5–7.0 mm (86.1%). Total- 3 era (from Baja California, Mexico), one archaeological ized dimensions = 300–1200 mm (94.3%). Oblong sample from Roman Karanis and the large Eocene (35.7%), cylindrical (57.1%). Cream (7.1%) or brown fossil seed Phoenicites occidentalis (from Texas, USA) (35.7%) coloured (modern seeds). Apex obtuse. Base (Berry, 1914). Main descriptive parameters: Breadth/ truncate (12.4%), obtuse (60.7%), acute (27.1%). Length = 0.2–0.4. Length = 32–40 mm (88.9%). Surface smooth (71.4%) or rough (28.6%), wrinkled Breadth = 8–16 mm. Depth = 7–12 mm. Totalized (33.3%) or uniform (66.7%). With longitudinal grooves dimensions = 1850–2500 mm3. Oblong (33.3%), (21.4%). Micropyle near the middle of the dorsal face. cylindrical (66.7%). Brown coloured (modern seeds). Ventral raphe V-shaped (42.8%), U-shaped (7.1%) or Apex obtuse (88.9%) or truncate (11.1%). Base trun- shallow(50%). Dorsoventrally straight or bent (9.1%). cate (22.2%), obtuse (44.4%), acute (33.3%). Surface Not winged. rough, wrinkled (33.3%), or uniform (66.7%). Group 16: exclusively includes seeds of P. dactylif- Without longitudinal grooves. Micropyle near the era (from Spain, North Africa and the Near East) and middle of the dorsal face. Ventral raphe V-shaped several archeological samples from the Neolithic to (66.7%) or shallow(33.3%). Dorsoventrally straight. Roman period. Main descriptive parameters: Not winged. Breadth/Length = 0.2–0.6. Length = 19–32 mm Group 14: almost exclusively includes elongated (91.8%). Breadth = 6–10 mm. Depth = 5.5–12.0 mm. 3 and relatively small seeds of P. dactylifera (mainly Totalized dimensions = 800–1850 mm (79.6%). from West Asia but also from North Africa), and Ovate–triangular (23.1%), cylindrical (76.9%). Cream

Figure 4. Main types of living Phoenix seed samples (1) Cluster I. Group 1. A, P. loureiroi Europ 101. B, P. loureiroi Europ 105. C, P. loureiroi Europ 13. D, P. loureiroi Europ 4. E, P. loureiroi Rare 6. F, P. loureiroi Sandeman 4. Group 2. G, P. acaulis Europ 106. Group 3. H, P. caespitosa Acaulis 1, I, P. loureiroi Usda 3. G, Group 4. J, P. reclinata Jbo- tanico 1. Group 5. K, P. theophrasti Elaguna 1. Group 6. L, P. andamanensis Olocan 2. M, P. loureiroi (P. hanceana) Riverside 37. N, P. loureiroi USDA 4P. O, P. pusilla Sun 1. Group 7. P, P. rupicola Keni 1. Q, P. rupicola Kpr 4. R, P. rupicola Rare 3. Group 8. S, P. canariensis Cespinardo 3. T, P. canariensis Cespinardo 5. U, P. canariensis Kpr 1. Cluster II. Group 9. V, P. caespitosa (P. arabica) Joe 13. Group 10. W, P. roebelenii Olocau 1. Group 11. X, P. dactylifera Jaravia 5. Cluster III (p.p.). Group 12. Y, P. dactylifera ‘Amir Hajj’ Riverside 8. Z, P. dactylifera ‘Dayri’ River- side 35. AA, P. dactylifera ‘Medjool’ Riverside 12. AB, P. dactylifera ‘Pyarum’ Berlin 2. AC, P. dactylifera ‘Redondos’ Mercen 6. AD, P. dactylifera Israel 2. AE, P. dactylifera Olivar 1. AF, P. dactylifera Orisa 5. AG, P. dactylifera SIBC 13. AH, P. dactylifera SIBC 15. AI, P. dactylifera ‘Tenats’ Elche 1. Scale bars in mm; 5-mm grid. Photographs A–AI, Joaquín García.

Figure 4. See caption on previous page.

Figure 5. See caption on next page.

Figure 5. Main types of living Phoenix seed samples (2) Cluster III (p.p.). Group 14. A, P. atlantica Cabo Verde 1. B, P. dactylifera ‘Abada’ Riverside 7. C, P. dactylifera ‘Deglet Nour’ Deglet 1. D, P. dactylifera ‘Khisab’ Riverside 34. E, P. dactylifera ‘Khudari’ Arabia 3. F, P. dactylifera Libya 1. Group 15. G, P. dactylifera ‘Khadrawy’ Riverside 15. H, P. dactylifera ‘Khir’ Riverside 4. Group 16. I, P. dactylifera ‘Bentamoda’ Riverside 6. J, P. dactylifera ‘Halawy’ River- side 25. K, P. dactylifera Fuentes 3. Cluster IV. Group 18. L, P. atlantica Cabo Verde 2. M, P. atlantica Cabo Verde 3. N, P. canariensis var. porphyrococca Lisboa 1. O, P. canariensis var. porphyrococca Riverside 1. P, P. dactylifera ‘Barhee’ Barhee. Q, P. dactylifera ‘Zahidi’ Riverside 36. R, P. dactylifera SIBC 18. S, P. dactylifera × P. iberica IslaPlana 1. T, P. dactylifera × P. iberica SIBC 23. U, P. theophrasti ‘Datça’ Turquia 2. Group 19. V, P. dactylifera Fuentes 1. W, P. dactylifera Parque 1. X, P. sylvestris ‘Robusta’ Keni 6. Y, P. sylvestris Rare 2. Z, P. sylvestris Riverside 42. AA, P. sylvestris Riverside A1. Group 20. AB, P. dactylifera ‘Badrayah’ Riverside 18. AC, P. dactylifera ‘Deglet Beida’ Riverside 2. AD, P. dactylifera ‘Hayani’ Riverside 30. AE, P. dactylifera Alcudia 1. AF, P. dactylifera SIBC 14. Group 21. AG, P. sylvestris × P. pusilla Keni 4. Group 22. AH, P. dactylifera ‘Asrashi’ Riverside 32. Cluster V. Group 24. AI, P. paludosa Rare 5. Scale bars in mm. 5-mm grid. Photographs A–AI, Joaquín García.

(30.8%) or brown (46.2%) coloured (modern seeds). seeds). Apex obtuse. Base truncate (36.5%), obtuse Apex obtuse (61.5%) or acute (37.9%). Base oblique (58.6%). Surface smooth (29.4%) or rough (70.6%), (81.3%). Surface smooth (46.2%) or rough (53.8%), wrinkled (51.9%) or uniform (48.1%). With longitudi- wrinkled (55.5%), or uniform (44.5%). Without longi- nal grooves (19.8%). Micropyle near the middle of the tudinal. Micropyle near the middle of the dorsal face. dorsal face. Ventral raphe V-shaped (52.8%), Ventral raphe V-shaped (30.7%), or U-shaped (69.3%). U-shaped (36.1%) or shallow (11.1%). Dorsoventrally Dorsoventrally straight. Not winged. straight or bent (1.7%). Rarely winged (1.5%). Group 17: almost exclusively includes archaeologi- Group 19: includes the seeds of P. sylvestris (from cal, cylindrical but dorsoventrally ﬂattened seeds India) and the type icon of this species, the type of from Iron Age Raybun (South Arabia) and one sample P. iberica, several samples of P. dactylifera from Spain of P. dactylifera from Baja California (Mexico). Main and Baja California (Mexico) and the whole sample of descriptive parameters: Breadth/Length = 0.2–0.6. Miocene fossil seeds named P. bohemica (Buzek, Length = 15–25 mm. Breadth = 6–10 mm. Depth = 4– 1977). Main descriptive parameters: Breadth/ 7 mm. Depth/breadth < 0.75. Totalized dimen- Length = 0.4–0.8. Length = 15–25 mm (81.2%). sions = 300–1850 mm3. Oblong (10%), cylindrical Breadth = 8–12 mm. Depth = 7–12 mm. Totalized (90%). Brown coloured (modern seeds). Apex obtuse. dimensions = 800–2500 mm3. Oblong (37.7%), ellip- Base obtuse. Surface smooth, wrinkled (10%), or soid (52.9%). Cream (64.6%) or brown (69.8%) col- uniform (90%). Without longitudinal grooves. Micro- oured (modern seeds). Apex obtuse. Base truncate pyle near the middle of the dorsal face. Ventral (42.1%), obtuse (55.9%). Surface smooth (9.9%) or raphe shallow. Dorsoventrally straight. Not winged. rough (88.1%), wrinkled (28.2%), or uniform (71.8%). With longitudinal grooves (57.7%). Micro- pyle near the middle of the dorsal face. Ventral raphe CLUSTER IV V-shaped (19.2%), U-shaped (80.8%). Dorsoventrally This cluster includes somewhat rounded seeds of straight. Not winged. P. dactylifera, P. atlantica, P. iberica, P. sylvestris and Group 20: exclusively includes seeds of P. dactylif- a few archaeological samples and one Miocene fossil era (from Spain, Baja California, North Africa and the sample. Near East) and three archeological samples, two from See Figures 3B, C and 5. Roman Karanis and one from Iron Age Tayma. Main Group 18: includes seeds of P. dactylifera (from descriptive parameters: Breadth/Length = 0.2–0.6. Spain, North Africa and the Near East), several Length = 19–25 mm (83.6%). Breadth = 8–12 mm. hybrids of P. dactylifera, two samples of P. atlantica Depth = 7–12 mm (92.2%). Totalized dimen- from Cabo Verde, two of P. senegalensis, one of sions = 1200–2500 mm3 (94%). Oblong (92.4%). P. canariensis var. macrocarpa collected by Hermann Cream (30%) or brown (60%) coloured (modern seeds). Wildpret in Tenerife, two samples from Turkey Apex obtuse (92.2%). Frequently mucronate labelled by Professor Esener as P. theophrasti and one (29.6%). Base truncate (39.2%), obtuse (44.3%), acute archeological sample from Neolithic Takarkori (16.7%). Surface smooth (13.3%) or rough (86.7%), (Libya). Main descriptive parameters: Breadth/ wrinkled (76.7%), or uniform (23.3%). Without lon- Length = 0.4–0.6 (93.9%). Length = 15–25 mm gitudinal grooves. Micropyle near the middle of the (91.2%). Breadth = 6–10 mm. Depth = 5.5–12.0 mm. dorsal face. Ventral raphe V-shaped (56.7%), Totalized dimensions = 800–1850 mm3. Oblong. U-shaped (43.3%). Dorsoventrally straight. Rarely Cream (30.2%) or brown (69.8%) coloured (modern winged (4.9%).

Group 21: almost exclusively includes seeds of from the littorals of India and South-East Asia. The P. canariensis (from Canary Islands, Spain) compris- most typical character is the basal position of the ing the type icon of P. canariensis and seeds collected micropyle. See Figures 3C and 5. by Hermann Wildpret in Tenerife and now in FI, and Group 24: exclusively includes seeds of P. paludosa two P. sylvestris × P. pusilla hybrids. It also includes (from mangroves of South-East Asia). Main descrip- the single type specimen of the sample of Miocene tive parameters: Breadth/Length = 0.6–1. Length = fossil seeds named P. bohemica (Buzek, 1977). Main 4–10 mm. Breadth = 3.5–8 mm. Depth = 2.5– descriptive parameters: Breadth/Length = 0.6–0.8 5.5 mm. Totalized dimensions = 150–800 mm3. (90.8%). Length = 10–19 mm. Breadth = 8–12 mm. Hemispherical. Blackish (81.7%) or greyish Depth = 7–12 mm. Totalized dimensions = 800– (18.3%) coloured. Apex obtuse. Base obtuse. 1850 mm3. Elliptical. Cream (20.6%) or brown Surface smooth, uniform. Without longitudinal (79.4%) coloured (modern seeds). Apex obtuse. Base grooves. Micropyle basal. Ventral raphe shallow. truncate (26.1%), obtuse (73.3%). Surface smooth Dorsoventrally straight. Not winged. (78.9%) or rough (21.1%), uniform. With longitudinal grooves. Micropyle near the middle of the dorsal CLUSTER VI face. Ventral raphe U-shaped. Dorsoventrally straight. Not winged. This last cluster includes the largely variable in Group 22: almost exclusively includes seeds of dimensions but otherwise uniform outgroup with P. dactylifera (from Spain and the Near East) compris- globose seeds of Nannorrhops, Euterpe, Livistona and ing the type icon of the species and original material of Washingtonia. See Figure 3C. P. dactylifera var. adunca Becc. (now in FI). It also Group 25: outgroup, exclusively includes seeds of contains the sample of seeds and dates bought by Nannorrhops, Euterpe, Livistona and Washingtonia. A. Chevalier in the market of Praia (Cabo Verde) Main descriptive parameters: Breadth/Length = 0.6– and named P. atlantica (now in P). Main descriptive 1.0. Length = 4–15 mm. Breadth = 3.5–16.0 mm. parameters: Breadth/Length = 0.2–0.6 (93.8%). Depth = 2.5–12.0 mm. Totalized dimensions = 0– 3 Length = 15–25 mm (94.3%). Breadth = 6–12 mm. 2500 mm . Globose. Brown coloured. Apex Depth = 7–12 mm (85.3%). Totalized dimen- obtuse. Base obtuse. Surface smooth, uniform. sions = 800–2500 mm3 (90.6%). Ovate–triangular. Without longitudinal grooves. Micropyle basal. Cream (40%) or brown (60%) coloured (modern seeds). Ventral raphe shallow. Dorsoventrally straight. Not Apex obtuse (59.1%) or acute (40.9%). Often mucro- winged. nate (38.4%). Base truncate (42.3%), obtuse (37.9%), acute (19.3%). Surface smooth (6.7%) or rough DISCUSSION (92.3%), wrinkled (37.8%), finely grooved (14.2%) or uniform (48%). Without longitudinal grooves. OVERALL PATTERNS OF GROUPING AND MORPHOTYPES Micropyle near the middle of the dorsal face. Ventral IN MODERN SPECIES raphe V-shaped (33.3%), or U-shaped (66.7%). Often In general, each of the groups described corresponds dorsoventrally bent (22.9%). Not winged. to a characteristic morphotype. Some species have Group 23: exclusively includes seeds of P. dactylif- morphologically homogeneous seeds and all samples era var. costata Becc. (from Spain, North Africa and studied of the same species are included in a single Baja California). Main descriptive parameters: morphotype. By contrast, other species show a great Breadth/Length = 0.4–0.8. Length = 15–25 mm morphological variability in their seeds, which are (87.8%). Breadth = 10–12 mm (88.9%). Depth = 7– integrated into different groups, corresponding to dif- 12 mm. Totalized dimensions = 1200–2500 mm3 ferent morphotypes. Phoenix dactylifera seeds have (94.4%). Ovate–triangular (80%), elliptical (15.6%). the highest variability (Tables 6 and 7). Cream (33.3%) or brown (66.7%) coloured (modern seeds). Apex obtuse (80%) or acute (20%). Occa- Cluster I sionally mucronate (17.8%). Base obtuse (64.4%), Phoenix loureiroi was lectotypified with the specimen acute (33.3%). Surface rough, wrinkled (33.3%), or Pierre 4832 (FB-I) from Mount Kuang Repen in Cam- uniform (66.7%). Without longitudinal grooves. bodia (Barrow, 1998), but it only consists of leaflets Micropyle near the middle of the dorsal face. Ventral and flowers. Most samples of P. loureiroi are in and raphe V-shaped (33.3%), U-shaped (33.3%) or shallow form the majority of group 1 (Cluster I) (Table 6). (33.3%). Dorsoventrally straight. Winged (88.9%). However, a few, probably of hybrid origin and particularly those named P. hanceana from Hong Kong CLUSTER V and the Philippines (now a synonym of P. loureiroi), This cluster includes exclusively an extremely homo- are in groups 3 and 6 (Cluster I). No regular geneous group of small rounded seeds of P. paludosa pattern was detected for seed morphology variation

Table 7. Summary of the seed morphology of Phoenix dactylifera and its hybrids. Numbers after names of the species refer to morphotypes/groups, 12 to 17 belong to Cluster III and 18 to 23 belong to Cluster 4 TAL ET Characters/ P. dactylifera P. dactylifera P. dactylifera P. dactylifera P. dactylifera P. dactylifera P. dactylifera P. dactylifera P. dactylifera P. dactylifera P. dactylifera P. dactylifera P. dactylifera × P. dactylifera × 04TeLnenSceyo London, of Society Linnean The 2014 © states 12 13 14 15 16 17 18 19 20 22B Type 22C adunca 23 costata P. canar 18 P. iberica 18 . B/L 0.2–0.6 0.2–0.4 0.2–0.6 0.2–0.4 0.2–0.4 0.2–0.6 0.4–0.6 0.4–0.8 0.2–0.6 0.4–0.6 0.2–0.6 0.4–0.8 0.4–0.6 0.4–0.6 L mm 19–32 32–39 15–32 19–32 19–32 19–32 15–25 15–25 19–25 15–25 15–25 15–25 10–19 15–25 B mm 8–12 10–12 6–10 6–8 6–10 8–12 6–10 8–12 8–12 10–12 6–12 8–12 6–10 6–10 D mm 5.5–15 7–17 5.5–15 4–7 5.5–15 4–7 5.5–15 7–15 7–15 5.5–12 5.5–12 7–15 7–15 5.5–12 TD mm3 1200–2500 1850–2500 800–1850 800–1850 800–1850 800–1850 800–1850 800–1850 800–2500 1200–1850 800–2500 800–2500 300–1200 800–1850 Outline Cylindric, Cylindric Cylindric, Cylindric Cylindric Oblong Oblong Elliptic Oblong Ovate– Ovate– Ovate– Oblong Oblong oblong oblong triangular triangular triangular, (elliptic) elliptic Colour Brown Brown Brown, Brown Brown, Brown Brown, Brown Brown Cream Brown, Brown, Brown Brown (cream) cream (cream) cream cream (cream) (cream) cream cream (cream) Apex Obtuse Obtuse Acute Obtuse Obtuse Obtuse, Obtuse Obtuse Obtuse Acute Acute Obtuse Obtuse Obtuse (acute) (obtuse) (acute) truncate (obtuse) (acute) Apex Frequent Not Sometimes Not Not Not Not Sometimes Frequent Always Sometimes Sometimes Not Not mucronate Base Truncate Acute Acute Acute Oblique Truncate, Obtuse Obtuse Obtuse Truncate Obtuse Obtuse Truncate Obtuse oaia ora fteLnenSociety Linnean the of Journal Botanical (obtuse, (truncate) (truncate, (truncate, obtuse (truncate) (truncate, (truncate) (acute) (obtuse) acute) obtuse) acute) acute) Base Sometimes Not Frequent Not Frequent Not Sometimes Not Not Not Sometimes Not Not Not mucronate Surface Rough Rough Smooth, Smooth, Smooth Smooth Rough Rough Rough Rough Rough Rough Rough Rough (smooth) rough rough (rough) (smooth) (smooth) (smooth) (smooth) Transverse Wrinkled, Uniform Wrinkled, Wrinkled, Wrinkled, Wrinkled Wrinkled, Wrinkled, Wrinkled Finely Wrinkled Uniform Uniform Wrinkled processes ﬁnely uniform uniform uniform uniform uniform grooved (uniform, (wrinkled) (uniform) grooved (wrinkled) ﬁnely grooved) Longitudinal Not Not Not Not Not Not Not Sometimes Not Not Not Not Not Not grooves Micropyle Central Central Central Central Central Central Central Central Central Central Central Central Central Central Ventral V-shaped V-shaped Shallow V-shaped, U-shaped, U-shaped V-shaped V-shaped V-shaped, U-shaped U-shaped, U-shaped, Shallow U-shaped furrow (U-shaped) (V-shaped) shallow V-shaped (U-shaped) (U-shaped) U-shaped (V-shaped) V-shaped V-shaped, (U-shaped) shallow Dorso-ventral Not Not Not Sometimes Not Not Not Not Not Not Frequent Not Not Not

2014, , curvature Wings Not Not Not Not Not Not Not Not Not Not Not Frequent Not Sometimes 175 74–122 , CARPOLOGICAL ANALYSIS OF PHOENIX 103

Figure 6. Phoenix andamanensis S.Barrow. A, Holotype, Ellis 14189 (K). B, Phoenix loureiroi from Batanes Island (Philippines) (type specimen of P. hanceana var. philippinensis Becc., FI-B). Scale bars in mm. B, photograph Teresa Egea.

Figure 7. Phoenix pusilla Gaertn. (p.p.)A,Phoenix pusilla Gaertn. Lectotype in Gaertner (1788–1791): table 9. B, Phoenix zeylanica Trimen. 1. Trimen (1898: plate 95). among different varieties recognized in this species by manenis (OLOCAN 02) falls within the variability of horticulturists or Barrow (1998). P. loureiroi and the studied seeds are not ruminate. For the type of P. andamanensis, Barrow (1998) For P. pusilla. Barrow (1998) mentioned Gaertner named Ellis 14189 (K), which is labelled as the holo- (1788–1791: fig. 9) as a lectotype. The icon depicts one type and contains numerous desiccated fruits and fruit and one seed (Fig. 7A). The seed falls in group 2 seeds (Fig. 6A). It is in group 1 (Cluster I) (Table 6) (Cluster I) (P. pusilla A, Table 6) with P. zeylanica, with numerous P. loureiroi samples. The ruminate which Barrow (1998) and Govaerts et al. (2011) endosperm (Barrow, 1998) was verified in this sample included in P. pusilla, and with several P. acaulis and in one sample of P. andamanensis from Rutland samples. Weighted neighbor joining shows as the (Andaman) Rogers (FI-B); both show deep brown closest sample the icon of P. zeylanica from Trimen rumina in transverse section. Weighted neighbor (1898) (Fig. 7B) (TRIMCEYHAN) (32% in 5000 boot- joining shows as the closest modern sample P. lourei- straps). Barrow (1998) typified P. zeylanica with roi from Batanes Island (Philippines) (type specimen Thwaites C.P. 3172 (K), which contains no seeds. of P. hanceana var. philippinensis Becc., FI-B) Phoenix farinifera, also a synonym of P. pusilla (Fig. 6B) (62% in 5000 bootstraps). No molecular evi- according to Barrow (1998) and Govaerts et al. (2011), dence was presented for this species by Pintaud et al. was typified with plate 74 of Roxburgh (1796) by 2010, 2013). As the character of rumination can break Barrow (1998) and the seed shown in this icon down and, in a few recorded species (e.g. Nypa fruti- (Fig. 8A) is in group 6 (Cluster I) (P. pusilla B, cans Wurmb and Ptychococcus paradoxus (Scheff.) Table 6) with two modern samples of P. pusilla (SUN Becc.), seeds can be homogeneous or ruminate, asso- 03 and SUN 01) and several from P. loureiroi with ciated with the low resolution obtained from P. lourei- Iron Age Raybun archaeological samples and an roi; further studies are necessary to ascertain the Eocene fossil labelled Serenoa carbonaria (Table 8). status of P. andamanensis as a species. The specimen Weighted neighbor joining shows as the closest labelled in the collection of Tomás Font as P. anda- sample P. pusilla (SUN 03) (Fig. 8B) (59% in 5000

Figure 8. Phoenix pusilla Gaertn. (p.p.)A,Phoenix farinifera Roxb. Holotype?, Roxburgh (1796: table 74 ). B, Phoenix pusilla (SUN 03), 5-mm grid. B, photograph Joaquín García.

Table 8. Ancestral states according to the morphology of the Tertiary fossil seeds analysed. For the purpose of comparison colour is not analysed because it is lost or strongly changed during fossilization and mucro is not because of the fragility of this appendix. For the purpose of comparison Serenoa repens (W.Bartram) Small, seeds are described

Phoenix Phoenicites Serenoa Characters / states bohemica Phoenix hercynica occidentalis carbonaria Serenoa repens

B/L 0.4–0.8 0.2–0.4 0.2–0.4 0.4–1 0.5–0.6 L mm 10–25 19–25 32–39 4–15 17–20 B mm 8–10 3.5–6 12–16 3.5–8 9–11 D mm 7–(17) 7–(17) 7–(17) 4–7 9–11 D/B 0.85–0.9 0.95–1.5 0.95–1.5 0.85–0.9 0.9–1.1 TD mm3 800–1850 800–1200 1850–2500 150–800 1300–2500 Outline Elliptic Cylindric Oblong Elliptic, globose, Elliptic ovate–triangular Apex Obtuse Acute Obtuse Obtuse Obtuse (acute) Base Obtuse Acute Obtuse Truncate Obtuse Surface Rough Rough Rough Smooth Smooth Transverse processes Wrinkled Wrinkled Wrinkled Uniform Uniform Longitudinal grooves Not? Not? Not? Not? Not Micropyle Central Central Central Central Basal Ventral furrow U-shaped U-shaped Shallow U-shaped Not Dorso-ventral curvature Not Not Not Not Not Wings Not Not Not Not Not bootstraps). Therefore, it seems that P. pusilla is poly- According to Barrow (1998), the lectotype of P. rec- morphic with regard to seed morphology. This poly- linata is an icon published in Jacquin (1801: plate 24). morphism is not found in the homogeneous cluster The seed shown in this icon (Fig. 10A) falls in group 4 obtained for the four samples of the species studied by (Cluster I) with several modern samples of P. recli- Pintaud et al. (2010). nata (Fig. 10B) and archaeological samples from North Barrow (1998) mentioned plate 273 in Roxburgh Africa and West Asia, including Bronze Age casts of (1820) as the type of P. acaulis. This plate includes an date seeds from Ar Raqlah (Yemen). Several samples image of a seed (Fig. 9A). It falls in group 2 that are probable hybrids of P. reclinata with other (Cluster I) with several P. acaulis samples (Table 6), species fall in groups 1, 3 and 6 (Cluster I). Pintaud and the types of P. pusilla and P. zeylanica. Weighted et al. (2010) studied numerous samples from different neighbor joining shows two P. acaulis fromB&T East African countries (but not from West and South Seeds (ACAULISB&T_1 and 3) (Fig. 9B) (53% in 5000 Africa), which cluster with a bootstrap value > 70%. bootstraps) as the closest modern samples. Pintaud These appear close to the P. loureiroi cluster, but also et al. (2010) studied two samples cultivated in the with low bootstrap values. USA, seemingly from seeds collected in India, and the The holotype of P. theophrasti is in the herbarium unrooted neighbor joining tree based on simple Greuter (PAL-Gr), but an isotype with numerous sequence repeat (SSR) markers clustered these fruits and seeds is at K. Several seed samples from samples with P. caespitosa and P. sylvestris, in con- the classical locality of Vai (Crete, Greece) and one trast to our results. from Gölköy (Turkey) fall in group 5 (Cluster I)

Figure 9. Phoenix acaulis Roxb. A, neotype Roxburgh (1820: table 273). B, sample from north-east India (ACAULISB&T_1). Scale bars in mm. B, photograph Diego Rivera.

Figure 10. Phoenix reclinata Jacq. A, lectotype Jacquin (1801: table 25). B, Jardín Botánico (Valencia, Spain). 5-mm grid. B, photograph Joaquín García.

(Table 6) with a waterlogged date seed recovered from are the only members of group 7 (Cluster 1) Neolithic Atlit-Yam (Israel) (Fig. 11A). Weighted (Table 6). Pintaud et al. (2010) found four samples of neighbor joining shows as the closest living relatives this species from India and Bhutan in the same to the Neolithic seed two samples of P. theophrasti distinct cluster of the unrooted neighbor joining tree (66% in 5000 bootstraps), from Vai and Gölköy based on SSR markers with a bootstrap value of 92%, (Fig. 11B, C). Seeds from Datça and one sample from coinciding with our results and underlining the iso- Gölköy (both Turkey, but the last probably misla- lation of this taxon. belled for Datça) labelled P. theophrasti fall in According to Barrow (1998), the lectotype of group 18 (Cluster IV) (Table 6), with numerous P. caespitosa is the specimen collected in Somalia, P. dactylifera cultivars suggesting these are not ‘Scorasar’ valley, 1 July 1924, Puccioni & Stefanini P. theophrasti, but of hybrid origin, or simply that 672 (738) (FT) with a female inﬂorescence and a leaf these are feral P. dactylifera. Pintaud et al. (2010) fragment, but with no seeds. Phoenix arabica was showed that samples of this species from Crete and treated as a synonym by Barrow (1998). We could not Turkey fell in the same distinct cluster of the analyse seeds from East Africa (Somalia and Dji- unrooted neighbor joining tree based on SSR markers bouti), although two specimens from cultivated plants with a bootstrap value of 84%, coinciding with our (group 3) may belong to this species. A herbarium results for the Crete and Gölköy samples. specimen of P. arabica (FAIRCHILD 1) (Fig. 12A) For P. rupicola, Barrow (1998) mentioned as origi- from the Fairchild Tropical Botanical Garden nal material India, West Bengal, Sivoka, Teesta (Florida, USA) from a palm introduced there from valley, February 1867, Herb. Sikkimense T. Anderson Saudi Arabia falls in group 8 (Cluster I) (Table 6) s.n. [CAL (sterile material), K (not yet in the digital with several P. canariensis samples, one P. loureiroi herbarium)]. Several modern samples of this species sample and three archaeological samples. Weighted

Figure 11. Phoenix theophrasti Greuter. A, one complete waterlogged date kernel PPNC (6000–5200 BC) found in Atlit-Yam at structure 20 (Kislev et al., 2004). B, seeds from Gölköy (Turkey) Elaguna 02. C, seeds from Vai (Crete, Greece) Europ 07. Scale bars in mm. 5-mm grid. A, photograph Anat Hartmann-Shenkman; B–C, photographs Joaquín García.

Figure 12. Phoenix caespitosa Chiov. (P. arabica Burrett). A, PLEI_KHARGA (Gardner, 1935). B, Fairchild 1. Scale bars in mm. B, photograph Diego Rivera. neighbor joining shows a Pleistocene sample from Cluster II Kharga Oasis (Egypt) (50% in 5000 bootstraps) The holotype specimen of P. roebelenii is sterile (Fig. 12B) as the closest specimen. However, as this (Barrow, 1998), and thus seeds could not be analysed. last material is carbonized, further analyses will be Modern seed samples from different origins form the required with experimental charring of modern seeds. distinct group 10 (Cluster II) (Table 6). Seeds are Several, immature P. caespitosa (P. arabica) seeds small and thin and have a widely open ‘U’-shaped from Yemen form group 9 (Cluster II). Pintaud et al. ventral raphe (Fig. 4W). Pintaud et al. (2010) showed (2010) showed that the only sample of this species that four samples of cultivated specimens of this from Somalia fell close to the clusters of P. acaulis species fell in the same distinct cluster of the and of P. sylvestris in the unrooted neighbor joining unrooted neighbor joining tree based on SSR markers tree based on SSR markers with a bootstrap value with a bootstrap value of 79%, coinciding with our > 70%. Pintaud et al. (2013) showed that samples of results and underlining the isolation of this taxon. this species fell in an early branching position in the haplotype network reconstruction based on plastid Clusters III and IV sequence data related to the ‘Phoenix dactylifera The type specimen of P. iberica (herbarium MUB) clade’ and to the branch of P. reclinata. Further includes seeds (IBTYPE_1) that fall in group 19 studies with additional samples will be required to (Cluster IV) with several P. dactylifera/P. iberica clarify the status of this taxon. samples from south-eastern Spain, P. sylvestris

Figure 13. Phoenix sylvestris L. A, lectotype, Rheede (1682: plate 25). B, sample from India (Keni 05). C, sample from India (Keni 06). 5-mm grid. B, C, photographs Joaquín García.

Figure 14. Phoenix canariensis H.Wildpret. A, lectotype, Chabaud (1882: fig. 68). B, Wildpret (FI). C, archaeological material from Garajonay. B, photograph Teresa Egea. C, photograph Jacob Morales. samples including the type and the whole sample of Moore (1971) designated Chabaud (1882): figs 66– ten seeds from Miocene P. bohemica (FOSS 68 as the lectotype of P. canariensis. Particularly MIOCBOHEM1-10). Thus, distinction of this species interesting for the present study is Chabaud (1882): is not achieved, but the analysis places P. iberica in a fig. 68, which depicts a seed (Fig. 14A) and presents group sharing numerous ancestral states (Fig. 3C, further implications for the taxonomy of the species Table 6). No molecular evidence was presented for (Rivera et al., 2013b). This seed (CHABATYPE) falls this species by Pintaud et al. 2010, 2013). in group 21 (Cluster VI) with the sample of Moore & Dransfield (1979) designated plates 22 to P. canariensis seeds sent from Tenerife by Hermann 25 of Rheede (1682), depicting Katou-Indel, as the Wildpret to Odoardo Beccari in December 1886 (FI) lectotype of P. sylvestris. Plate 25 includes an image of (FIWILDP_1) (Fig. 14B), several modern samples a seed (Fig. 13A), which falls in group 19 from the Canary Islands and the Iberian Peninsula (Cluster IV) (Table 6) with all the modern samples of (P. canariensis B, Table 6), two hybrids and the type P. sylvestris, several P. dactylifera/P. iberica samples specimen of the sample of ten seeds from Miocene from south-eastern Spain (including the holotype of P. bohemica (FOSS MIOCBOHEM1-10). Thus, spe- P. iberica) and the whole sample of ten seeds from cific distinction is supported and the analysis places Miocene P. bohemica (FOSS MIOCBOHEM1-10). P. canariensis in a group sharing numerous ancestral Weighted neighbor joining shows as the closest speci- states (Fig. 3C, Table 6). However four small-sized mens a pair of P. sylvestris samples from Darjeeling seed samples from cultivated P. canariensis (Espi- (India) (KENI 5 and 6) (Fig. 13B, C) (44% in 5000 nardo, Murcia, Spain) fall in group 8 (Cluster I) bootstraps). Molecular evidence was presented for (P. canariensis A, Table 6) with four date seeds recov- this species by Pintaud et al. 2010, 2013), but the ered from a Guanche religious offering site at Gara- cluster had a bootstrap value < 70%. jonay (Canary Islands, Spain) (Fig. 14C) (Morales,

Figure 15. Phoenix atlantica A.Chev. and P. dactylifera var. adunca O.Becc. A, P. dactylifera var. adunca from Algeria, holotype (FI-B) (FIBECC_2). B, P. atlantica. Sample collected by Chevalier in the market of Praia (Cabo Verde) (P). C, P. atlantica. São Vicente island (Cabo Verde). Scale bars in mm. A, photograph Teresa Egea; B, photograph Muséum national d’Histoire naturelle, Paris; C, photograph Diego Rivera.

Navarro & Rodríguez, 2011), other archaeological fall in group 18 (Cluster IV). Exhaustive sampling samples from Roman Karanis (Egypt) and Chalco- in the ensemble of Cabo Verde islands will be lithic Tepe Gaz Tavila (Iran) and one modern sample required to delimit the morphology of seeds within of P. caespitosa (P. arabica). It is important to note this species clearly. Pintaud et al. (2013) included two this dimorphism in samples of P. canariensis, particu- samples of this species in the ‘Phoenix dactylifera larly when the only archaeological sample studied is clade’ based on plastid sequence data. not in the group of the type of the species. However, Moore & Dransﬁeld (1979) designated Kaempfer it is not surprising because P. canariensis shows a (1712): plates 1 and 2 depicting Palma hortensis mas et high degree of morphological and molecular variabil- foemina as the lectotype of P. dactylifera. Plate 2 ity within and between each one of the seven Canary includes an image of a seed (Fig. 16A). This seed Islands (P. Sosa, pers. comm.). Pintaud et al. (2010) (KAEMPTYPE_1315) falls in group 22 (Cluster IV) showed that four samples of cultivated specimens of (P. dactylifera 22B Type, Table 7). Weighted neighbor this species fell in the same distinct cluster of the joining shows as the closest specimen a P. dactylifera unrooted neighbor joining tree based on SSR markers sample from Ricabacica (south-eastern Spain) with a bootstrap value of 99%, underlining the isola- (Fig. 16B) (64% in 5000 bootstraps). Modern samples tion of this taxon. of this species are predominant in Clusters III and Phoenix atlantica was typiﬁed by Chevalier 45839 IV, and several immature seeds fall in Cluster II. (P) from Algodeiro (Sal Island, Cabo Verde) (Barrow, This species shows the highest variability in terms of 1998). However, the two sheets under this number at seed shape and dimensions in Phoenix and it is related P seem to contain more than one specimen and no with at least two different sets of ancestral states seeds. Phoenix seed samples from Cabo Verde show (Fig. 3B, C, Tables 7 and 8). At least 12 different high morphological variability, and attribution to morphotypes can be distinguished (Table 7). This could either P. dactylifera or P. atlantica is difficult. These be interpreted as a consequence of different selective do not form a distinct group. The type specimen of pressures during domestication, an indicator of poly- P. dactylifera var. adunca from Algeria (FI-B) phyletic origin of P. dactylifera or both. Terral et al. (FIBECC_2) (Fig. 15A) falls in group 22 (2012), using a different methodology, recognized ten (Cluster IV) (Table 6) with a sample from Praia morphotypes. Although the methodology employed market (São Tiago island, Cabo Verde) collected by and the analysed samples are different, there is in part Chevalier (P) (Fig. 15B), which is original material of a correspondence between our groups (12–20 and P. atlantica, and two fruit samples bought at the 22–23) and those morphotypes of Terral et al. (2012): market of Mindelo by J. Meseguer (São Vicente island MT1 with 17 (doubtful); MT2 with 22 (doubtful); MT3 (Cabo Verde) (Fig. 15C), the type icon of P. dactylifera with 14; MT4 with 19 and 18 p.p.; MT5 with 12; MT6 and several P. dactylifera cultivars from Spain, two with 15 p.p. and 18 p.p.; MT7 with 15 p.p.; MT8 with from Socotra and one from West Asia (P. dactylifera 16 (doubtful); MT9 with 18 p.p.; and MT10 with 20. ‘Asrashi’). The seeds are small and often regularly Pintaud et al. (2010) studied 20 samples from different dorsoventrally bent. Other samples from Cabo Verde countries that clustered with a bootstrap value > 70%.

Figure 16. Phoenix dactylifera L. A, lectotype Kaempfer (1712: table 2). B, cultivar from Ricabacica (Murcia, Spain) (PERICA_01a). Scale bars in mm. 5-mm grid. B, photograph Joaquín García.

These appear close to the P. theophrasti and P. canar- tylifera and P. sylvestris, P. canariensis, P. theophrasti iensis clusters, but also with low bootstrap values. (from Datça, Turkey) and other taxa which were not Pintaud et al. (2013) included samples of this species included in the study of Terral et al. (2012). with P. atlantica and P. sylvestris in the ‘Phoenix dactylifera clade’ based on plastid sequence data. Phoenix dactylifera var. adunca. The type specimen TERTIARY FOSSILS from Algeria (FI-B) consists of three seeds Phoenicites occidentalis (Berry, 1914: 403–406). (FIBECC_2) (Fig. 15A) and falls in group 22 (Cluster III, group 13) (Table 8). Type: collected by (Cluster IV) with three modern samples from São Laurence Baker in Texas, from a cut on the Interna- Vicente and São Tiago islands (Cabo Verde) (P. atlan- tional and Great Northern Railroad in southern tica/P. dactylifera) and one from West Asia (P. dactyl- Trinity County. The outcrop is referred to the Cata- ifera)(P. dactylifera 22C adunca, Table 7). houla formation, which in this region is of Late Phoenix dactylifera var. costata. The type specimen Eocene or Early Oligocene age. The type is shown in from Valencia (Spain) (FI-B) (FIBECC_3) falls in Figure 17A. (Berry, 1914: fig. 1). The finding con- group 23 (Cluster IV) with two modern samples tained both large and small seeds, and a cast of entire from Santomera (Murcia, Spain) and Baja California fruit of a Phoenix-like palm (Berry, 1914, 1937). It (Mexico). Seeds are small and often regularly winged was supposed to be in the Oscar M. Ball Collection (Table 7). Texas Natural Science Center, The University of Texas at Austin, or in the Smithsonian Paleontologi- Cluster V cal collections (there under USNM number P38340). The lectotype of P. paludosa, is an unpublished plate However, the type specimen is lost and, thus, our under No. 1193 of Roxburgh’s Flora Indica (K) analysis is based on the figure and original descrip- (Barrow, 1998). Modern seed samples of P. paludosa tion. Weighted neighbor joining shows Phoenix dac- form a distinct Cluster V (group 24) (Figs 3C, 5 AI, tylifera SIBC 11 from San José de Comondú (Baja Table 6), which is characterized by the basal opercu- California, Mexico) (Fig. 17B) as the closest living lum or micropyle. Pintaud et al. (2013) showed that sample (41% in 5000 bootstraps). However, this coin- samples of this species fell in an early branching cidence is merely accidental on palaeogeographical position in the haplotype network reconstruction grounds, as Phoenix is clearly a recently introduced based on plastid sequence data, related to P. roebele- genus in America (Rivera et al., 2010, 2013b). Total- nii. We find this species isolated from the rest in ized dimensions present an extremely high value many aspects (habitat, seed morphology, leaf colour). (Fig. 18). This is the only discovery of fossilized seeds, Working with outlines, Terral et al. (2012) separated related to Phoenix, across America and, although it P. theophrasti, P. canariensis, P. caespitosa, P. sylves- could not be directly studied (because it is lost), the tris and P. reclinata from a group of uncultivated (and material must be regarded as doubtful. there undetermined) Phoenix specimens from Oman. Phoenix hercynica (Mai, 1976: 102–103). At the same time, this group differed from several (Cluster III, group 14) (Table 8). Type: Tab. II, fig. 1 P. dactylifera cultivar groups. Our analysis confirms (Holotype). Open-cast mining Neumark-Süd the separation of P. reclinata, P. caespitosa and typical (Geiseltal), Stream-chute NS 22, one semi-carbonized P. theophrasti from Vai. However, with a wider sam- specimen, complete and well preserved, leg. Chrobok pling of P. dactylifera cultivars, our analysis shows a 195. It was not deposited in the Geiseltal Museum of more complex pattern of relationships between P. dac- the Martin-Luther-Universität Halle-Wittenberg,

Figure 17. Phoenicites occidentalis Berry. A, OCT_TEX (Berry, 1914). Phoenix dactylifera. B, SIBC 11. Scale bars in mm. 5-mm grid. B, photograph Joaquín García.

Figure 18. Totalized dimensions of Phoenix seeds (mm3) vs. time.

However, the type specimen is probably at Berlin. Tab. II, ﬁg. 10 (Holotype). Opencast mining Neumark- Seed long–elliptical, 20 × 6 mm, with deep ventral Süd (Geiseltal), Stream-chute NS 22, leg. Chrobok furrow (Fig. 19A, compared with P. dactylifera, 1965. The sample consists of more than 100 seeds Riverside-7, Fig. 19B and Riverside-5, Fig. 19C). fossilized in lignite. This and other samples from Weighted neighbor joining shows no living samples chute NS 37, consisting of 15 specimens, leg. Chrobok closely related to this fossil. A peculiar feature of this 1964/65, and from chute NS 38, four specimens, leg. fossil is the extraordinarily high depth/breadth ratio Chrobok 1965, were not deposited in the Geiseltal- (D/B = 1.33). This feature is relatively common among museum Zentralmagazin Naturwissenschaftlicher fossilized materials (33% D/B > 0.95) and is rare Sammlungen der Martin-Luther-Universität (ZNS) among living species (6% D/B > 0.95). Halle-Wittenberg and are presumably at Berlin. Fossil seeds published as Serenoa carbonaria (Mai, However, the specimens are not available. Although 1976: 106) (Cluster I, group 6) (Table 8). Type: described under Serenoa, the seeds, globose, ovoid or

Figure 19. Phoenix hercynica Mai. A, FOSS EOC_GEIS holotype (Mai, 1976: table 2, ﬁg. 1). Phoenix dactylifera. B, Riverside-7. C, Riverside-5. Scale bars in mm. 5-mm grid. B–C, photographs Joaquín García.

Figure 20. Serenoa carbonaria Mai. FOSS_SERENCAR_1. A, holotype (Mai, 1976: table 2, fig. 10). B, specimen from chute NS 22 (Mai, 1976: table 2, fig. 11). C, specimen from chute NS 22 (Mai, 1976: table 2, fig. 12). D, specimen from chute NS 38 (Mai, 1976: table 2, fig. 13). ellipsoidal (Fig. 20A–D), 6–12 × 4.5–7 mm, with Tuchorice near Zatec, north-western Bohemia, Czech micropyle dorsal, almost basal, are morphologically Republic, freshwater, often travertine-like limestone, Phoenix. The Geissel fossils are c. 50 Myr old. This Tuchorice Basin, Most Formation, Burdigalian, i.e. fossil falls in Cluster I, group 6, with samples of Eggenburgian (Lower Miocene) (Buzek, 1977; Kvacek several Phoenix spp. from East Asia (P. loureiroi, et al., 2004). Holotype deposited in the Department of P. pusilla and interspecific hybrids). Palaeontology, National Museum Prague (Fig. 21C, Phoenix szaferi Bakowski (1967). The variably D). Paratype: Specimen No. TU-2 shown in Buzek sized, peculiarly shaped, bulbous sandy bodies con- (1977). Fossil materials consist of ten fairly well- tained in coalified driftwood from the Lower Oligo- preserved seed casts or seeds replaced by crystallized cene (Rupelian) of the Tatra Mountains in Poland, calcite and approximately ten fragments (Buzek, originally described as fruit bodies of a new palm 1977). Seeds are oblong or ovoid, 1.2–2.0 × 0.8– species and then established as P. szaferi (Bakowski, 1.0 cm, with a longitudinal furrow on the ventral side 1967), were reinterpreted as a maze of internal and a single circular pore of germination on the moulds of siphonal tubes of the wood-boring bivalve opposite side, in some cases with transverse striae mollusks of the family Teredinidae (Teredolites clava- around the furrow (Fig. 21A, C, D). Weighted neigh- tus) (Radwanski, 2009). The grape-like ‘fruits’ do not bor joining shows as the closest living sample show any Phoenix seed-like material. Fuentes 1 (Fig. 21B) (61% in 5000 bootstraps) for the Phoenix bohemica (Buzek, 1977: 160). (Cluster IV, whole sample. However, weighted neighbor joining group 19 whole sample, group 21 holotype) indicates a group of P. canariensis samples as the (Table 8). Type: Specimen No. TU-8 on Tab. II, fig. 8. closest living sample to the holotype when analysed Road cut below the garden at house no. 80, village separately (38% in 5000 bootstraps). The holotype is

Figure 21. Phoenix bohemica Buzek. MIOCENBOHEM1_10. A, specimens 1–10, including two paratype and eight holotype. Details 11–12 of specimen 2. Thirteen dorsal of specimen 8. Fourteen dorsal of specimen 9 (Buzek, 1977). Scales, 10 mm. Phoenix dactylifera. B, Fuentes 1. Scale bars in mm. 5-mm grid. Phoenix bohemica. MIOCBOHEMTYPE. C, ventral view. D, dorsal view. Scale bars in mm. C–D, photographs Jiri Kvacek.

fossilized by calcium carbonate (CaCO3) (J. Kvacek, samples of P. theophrasti collected at Vai and Prevali pers. comm.). (Crete) and Gölköy (Turkey) and one archaeological Neolithic seed from Atlit Yam (Israel). Weighted neighbor joining indicates date seed from Queen PLEISTOCENE SEEDS Pu-abi’s grave (at Ur, Iraq) as the closest sample (52% At Kharga Oasis (South Egypt, West of Nile River) in in 5000 bootstraps). Late Pleistocene deposits (16 000 BP), dated cultur- ally to Aterian (early Upper Palaeolithic) loam beds containing carbonized reed-stems and yielding fruit HOLOCENE ARCHAEOLOGICAL REMAINS seeds of a wild date (identified by Mrs Clement Reid All archaeobotanical samples could be classified in as P. sylvestris) have been reported (Caton & Gardner, groups with modern seed samples. The assignment of 1932) (Fig. 12A). The seeds fall in group 8 archaeobotanical samples was made, mainly, to differ- (Cluster I) with several P. canariensis samples, one ent morphotypes of P. dactylifera (Clusters III and P. caespitosa sample [(P. arabica) FAIRCHILD 1] IV). However, some samples were assigned to groups (Fig. 12B) and three archaeological samples. with P. reclinata, P. caespitosa, P. atlantica, P. theo- However, as the material from Kharga is carbonized, phrasti, P. pusilla and P. canariensis. Archaeological further analyses will be required with experimental seeds were not allocated to group 19 containing charring of modern seeds. samples of P. sylvestris, P. iberica and Miocene fossil Fira palaeosol has been dated to c. 37 000 BP. P. bohemica. In general, it appears that some species During fieldwork in 1975–1978, fragments of pinnate such as P. theophrasti had a much larger area than at Phoenix-like foliage, casts of spines and a single present (reaching at least Israel and Palestine, and impression of a Phoenix fruit (12 × 5.4 mm, giving perhaps Iraq) and were collected and deposited in 11×5mmasputative dimensions for the seed) iden- contexts that have allowed their preservation. Some- tified as P. theophrasti were recovered (Friedrich, thing similar happened with P. caespitosa and P. recli- Pilcher & Kussmaul, 1977; Friedrich, 1980). This nata. By contrast, in the eastern and western ends of sample forms group 5 (Cluster I) with date seed the current range of P. dactylifera, we currently find from Queen Pu-abi’s grave (at Ur, Iraq), from Chal- P. sylvestris and P. iberica, respectively, which do not colithic Teleilat Ghassul (Palestine) and from Neo- appear in the archaeological record. This is logical as Elamite burial 693–6861 at Susa (Iran), and includes we have not been able to study archaeological samples

© 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175, 74–122 CARPOLOGICAL ANALYSIS OF PHOENIX 113 from these areas. Phoenix canariensis has two mor- (Cluster III) of P. dactylifera, but some samples were photypes, of which only Group 8 (Cluster I) includes assigned to groups with P. caespitosa, P. theophrasti archaeological materials from the Guanche period and P. canariensis (Cluster I). (Canary Islands) and others from North Africa (Egypt) Four samples (two carbonized and two brick casts) and the Middle East, suggesting that this morphotype from Dalma Island (United Arab Emirates) exclu- was much more extensive than at present. sively form group 15 (Cluster III), with a single desiccated seed from Bronze Age Tel Karrana (Iran) Neolithic (c. 8000–7500 BP) and two modern P. dactylifera samples from West All studied Neolithic seeds are desiccated or water- Asia. We note here the coincidence of seed imprints logged. Neolithic samples from different origins (Fig. 3, and carbonized materials from Dalma in the same Table 3) show different shapes and relationships, but group, as it shows how different forms of conservation they only fall in groups 14–16 (Cluster III) of P. dac- have not changed the morphological relationships tylifera, group 18 (Cluster IV) of P. dactylifera, between them in this case. P. atlantica and P. canariensis var. macrocarpa and, in one case, group 5 with P. theophrasti (Cluster I). Bronze Age (5000–3700 BP) The coincidence of a waterlogged date seed Bronze Age seed samples are brick casts (one site), (11.8 × 6.5 × 5.6 mm) (Fig. 11A), which was recovered desiccated (three sites) or carbonized (four sites). from Neolithic Atlit-Yam (Israel) with several modern Bronze Age samples from different origins (Table 3) samples of Phoenix theophrasti from Crete (Greece) fall in group 15 (Cluster III) of P. dactylifera, but and Gölköy (Turkey) in group 5 (Cluster 1) merits some samples were assigned to groups with P. recli- mention. Weighted neighbor joining shows two nata and P. theophrasti (Cluster I). modern samples of P. theophrasti from Vai and Gölköy Bronze Age casts of date seeds from Ar Raqlah (59% in 5000 bootstraps) as the morphologically (Yemen) fall in group 4 (Cluster I) with several closest samples (Fig. 11B, C). This would confirm the archaeological samples from North Africa and West identification of the seed by Kislev et al. (2004) as Asia and the type icon of P. reclinata. Many fragments P. theophrasti. of date seeds were found in Queen Pu-abi’s grave at A mineralized seed recovered from Mehrgarh IB Ur (Iraq), but only a few seeds were complete enough (Balochistan, Pakistan) in layers dated at c. 8000 BP to measure; these fall in group 5 (Cluster I) with is almost identical to another (Mehrgarh IIB) recov- Pleistocene, Chalcolithic and Neoelamite samples, ered in layers c. 1000 years more recent (Costantini, close to modern P. theophrasti samples. 1985; Beech, 2003), both in group 14 (Cluster III). A single desiccated seed from Tell Karrana (Iran) Two modern seed samples of P. dactylifera cultivars falls in group 15 (Cluster III), with several samples also belong to group 14 [‘Abada’ which originated in from Chalcolithic Dalma Island (United Arab Emir- California and related to ‘Amir Hajj’ from Iraq and ates), other Neolithic to Roman samples and five ‘Deglet Beida’ from Algeria (Cao & Chao, 2002], as do modern P. dactylifera samples from West Asia. A des- ‘Horra’ from Tunisia, 39 seeds from Bronze Age Ra’s iccated date seed from Jericho (Israel) falls in al-Jinz (Oman), three from Bronze Age Failaka group 4 (Cluster I), with several Bronze Age (Kuwait), and one from Middle Ages from Gao (Mali). archaeological samples (seed casts) from Yemen and Three seeds from Neolithic Sabiyah (Kuwait), dated modern samples of P. reclinata. Carbonized date seeds c. 7530 BP, form group 16 (Cluster III), that also from Ra’s al-Jinz (Oman) fall in group 14 include the sample from Roman Masada (Israel) and (Cluster III), with a carbonized seed from Bronze several modern samples of P. dactylifera cultivars Age Failaka (Kuwait). These are almost identical, and from south-eastern Spain, West Asia and North thus weighted neighbor joining shows a high coinci- Africa. One seed from Neolithic Takarkori (Libya) dence (93% of 5000 bootstraps). falls in group 15 (Cluster III) close to a modern sample of P. dactylifera ‘Khadrawy’. Other seed from Iron Age onwards (2800–800 BP) Neolithic Takarkori (Libya) in group 18 (Cluster IV) Iron Age and later seed samples are desiccated (eight appears to be related to modern samples of P. theo- sites) or carbonized (two sites from Europe). Iron Age phrasti (populations from Datça, Turkey), P. atlantica and later samples from different origins (Table 3) fall and P. dactylifera (from south-eastern Spain, Baja in groups 14–17 (Cluster III) and group 20 California and West Asia). (Cluster IV) of P. dactylifera, but some samples were assigned to groups with P. theophrasti, P. atlantica, Chalcolithic (c. 7400–5600 BP) P. caespitosa, P. canariensis, P. reclinata and P. pusilla Chalcolithic seed samples are brick casts (one site) or (Cluster I). carbonized (three sites). Chalcolithic samples from Desiccated dates from a Neo-Elamite burial at Susa different origins (Table 3) fall in group 15 (Iran) fall in group 5 (Cluster I) with one cast from

Pleistocene Fira (Santorini, Greece) and Bronze Age ANCESTRAL TRAITS Ur (Iraq) and modern samples of P. theophrasti. Many date seeds recovered from Raybun (Yemen) dated The oldest Phoenix-related seeds (excluding the prob- from 2800 BP onwards almost exclusively form lematic Phoenicites occidentalis) were recovered in group 17 (Cluster III) with one modern P. dactylif- Central Europe (Phoenix hercynica, Serenoa carbon- era sample and fall in group 15 (Cluster III) aria). However, with the available evidence we could (IA_RAYBU_4) with other archaeological and modern not determine if these are only isolated samples of a P. dactylifera samples from West Asia and group 6 previously widespread genus in the Late Cretaceous (Cluster I) with samples of P. pusilla, P. loureiroi or are part of an original nucleus from which they hybrids and Eocene Serenoa carbonaria. could diversify and colonize other territories. Subfossil desiccated seeds from Iron Age Tayma We found four seed morphologies clearly deﬁned in (Saudi Arabia) fall in group 20 (Cluster IV) with the Tertiary (Table 8). However, estimating the occur- archaeological samples from Roman Karanis (Egypt) rence of evolutionary events (interpreted as specia- (KARANIS_9 and 6) and modern samples of P. dac- tion and/or evolutionary radiation) would be tylifera from south-eastern Spain, Baja California, unrealistic according to the results and without con- West Asia and North Africa. A date seed from Par- sidering other sources of evidence including biology of thian Susa (Iran) falls in group 14 (Cluster III). the species, vegetative morphology and molecular Desiccated date seeds from Roman Masada (Israel), markers. Morphotypes are not species and thus which are well known for the extraordinary persis- simply tell us that during the Tertiary a relatively tency of their ability to germinate (Sallon et al., 2008), high morphological diversity was present. fall in group 16 (Cluster III) with modern samples Totalized seed dimensions show a slight tendency to of P. dactylifera from North Africa, West Asia and present a wider range of values, from the Miocene to south-eastern Spain. Weighted neighbor joining the present, but, logically, could be because modern shows P. dactylifera ‘Halawy’ from Iraq as the closest materials are best represented. Phoenicites occiden- modern sample (44% in 5000 bootstraps). talis (Eocene of Texas, USA) has dimensions that are Subfossil desiccated date seeds from Roman abnormally large (Fig. 17, Table 6). Karanis (Kom Aushin, Egypt) show different morpho- For the purpose of representing the different pos- logical patterns. One sample (KARANIS_1) falls in sible ancestral lines based on the fossils studied, we group 4 (Cluster I) with several archaeological compared the 24 groups and the outgroup (described samples from North Africa and West Asia, including above) with the four seed samples of the Tertiary and Bronze Age casts of date seeds from Ar Raqlah one Pleistocene sample, using the mean values of the (Yemen), and the type icon of P. reclinata. Another percentages of presence in the samples for each group sample (KARANIS_3) falls in group 8 (Cluster I) (rows) and each of the 67 states analysed (columns). with Pleistocene seeds from Kharga (Egypt), other The resulting hierarchical tree calculated with the archaeological samples and several modern samples Ward method (Fig. 22) shows clustering patterns of of P. caespitosa (P. arabica) and P. canariensis. Other modern and archaeological samples around four dif- samples (KARANIS_6 and 9) fall in group 20 ferent morphologies of fossil materials. (Cluster IV) with archaeological desiccated seed Fossil samples fall in Cluster I (Eocene and Pleis- samples from Iron Age Tayma (Saudi Arabia) and tocene), Cluster III (Eocene) and Cluster IV modern samples of P. dactylifera from south-eastern (Miocene). Archaeological samples fall in Clusters I– Spain, Baja California, West Asia and North Africa, IV. No fossil remains (seeds) have been found related and KARANIS_7 and 8 fall in group 12 (Cluster III) to P. paludosa (Cluster V). with modern samples of P. dactylifera from south- Small Eocene (Serenoa carbonaria) and Pleistocene eastern Spain and Baja California. Another sample, (Phoenix sp.) fossil seeds appear in Cluster I (Fig. 22) KARANIS_5, forms group 13 (Cluster III) with a associated with small seeded Phoenix spp. from modern sample of P. dactylifera from Baja California South-East Asia and India (P. loureiroi and P. anda- (Mexico) (SIBC 06) and the Eocene fossil P. hercynica. manensis, groups 1, 3 and 6; P. acaulis group 2, Four date seeds recovered from a Guanche religious P. pusilla, groups 2 and 6, P. rupicola group 7 and offering at Garajonay (Canary Islands, Spain) P. roebelenii group 10), Tropical Africa and the Near (Morales, Navarro & Rodríguez, 2011) fall in group 8 East (P. caespitosa, group 3 and P. reclinata, group 4), (Cluster I) with other archaeological samples and eastern Mediterranean (P. theophrasti, group 5) and several modern samples of P. caespitosa (P. arabica) the Canary Islands (P. canariensis, group 8). In the and P. canariensis. A single seed from Middle Ages light of the historical palaeogeography of the Medi- Gao (Mali) falls in group 14 (Cluster III) with terranean Basin (Rögl, 1998), land bridges for conti- modern samples of P. dactylifera from south-eastern nental migrations connected Europe and Asia since Spain, North Africa and Baja California. the Aquitanian (Early Miocene), allowing the migra-

Figure 22. Hierarchical tree calculated with the Ward method representing relationships between Tertiary and Pleis- tocene fossil samples and each one of the 25 groups recognized. tion of Phoenix. With the collision of the Arabian and and 22). The specimen designated as the type for the Anatolian plates in the late Burdigalian (Early species P. bohemica (Buzek, 1977: 160) shows closer Miocene to Mid Miocene) a Eurasian–African land similarity with the type icon of P. canariensis (Canary bridge opened for terrestrial plant migrations, which Islands). could explain the above relationships. The uplift of The attribution of Serenoa carbonaria to Phoenix is the Canary Islands took place from the Miocene made on the base of the raphe and other characters of onwards (Anguita & Hernán, 2000), allowing the colo- the fossil seeds that are present in modern and fossil nization by plants from nearby Atlantic North Africa. Phoenix but not in modern Serenoa seeds (Table 8). It The abnormally large Eocene seed from Texas falls is likely that other fossilized seeds from the Western in Cluster III (group 13). This morphotype appears and Eastern Hemispheres labelled Serenoa would be clearly separated from the rest of samples in more directly related to Phoenix than to modern Figure 22. In the same cluster, the other Eocene seed Serenoa spp. sample, from Geiseltal (P. hercynica), appears with To clarify the possible geographical patterns, we numerous archaeological samples and modern P. dac- map the palaeontological and archaeological sites tylifera cultivars (groups 14–17) (Fig. 22). yielding the different samples studied: Tertiary and Miocene fossil seeds from Tuchorice (P. bohemica) Pleistocene (Fig. 23A) and Holocene (Fig. 23B). fall in Cluster IV (Fig. 22), with numerous modern P. dactylifera cultivars from south-eastern Spain, North Africa, West Asia and Baja California APPROACH TO THE ANCESTRY OF (groups 12, 18–20 and 22–23), and Phoenix spp. from PHOENIX DACTYLIFERA the Mediterranean (P. iberica, group 19; P. theo- Although human selection practices, multiple hybridi- phrasti from Datça in Turkey, group 18), India (P. syl- zation events, geographical diffusion of varieties (fol- vestris, group 19) and Canary and Cabo Verde Islands lowing human migration routes), adaptation, etc., (P. canariensis, group 21 and P. atlantica, groups 18 may have completely blurred the evolutionary signal

Figure 23. Maps of paleontological and archeological sites yielding Phoenix seeds. A, Tertiary and Pleistocene. B, Holocene.

(evolutionary radiation, ancestral traits, ancestry, the Western Mediterranean, including P. iberica etc.), which could be deduced from measurements of related with the Western chlorotype. However, the the seed in P. dactylifera, the molecular evidence sug- direct wild ancestor of the date palm is still unknown gests that hybridization played a limited role in the and this issue remains unsolved, although we can emergence of P. dactylifera, and geographical patterns assume that the direct wild ancestor of the date palm of chlorotypes in the ‘Phoenix dactylifera clade’ possessed small and elliptic seeds. (Western and Eastern) (Pintaud et al., 2013) present notable coincidences with the main clusters of the GEOGRAPHICAL GROUPS analysed seeds. We find a surprising morphology in two of the three Major geographical patterns Eocene samples (Phoenicites occidentalis and Phoenix When analysing the geographical origins of samples hercynica) that could be linked with Phoenix. Both coinciding in clusters and groups, several patterns of fall in Cluster III, and their morphology is that of relationships became evident (Fig. 24). To clarify these P. dactylifera. Their shape (seed elongated and large) geographical patterns we summarized the distribution is common and exclusive to modern date palm culti- areas of the different species studied (Fig. 25). A group vars. Unfortunately, type material of both fossils was brings together south-eastern Spain, North Africa and inaccessible or lost. Attribution and dating of these Baja California (Fig. 24). This group comprises P. dac- samples needs to be carefully reviewed. tylifera local cultivars and suggests the original prov- Miocene fossil seeds from Tuchorice (Phoenix enances of the palms introduced in America were bohemica) show a close morphological relationship Spain and North Africa (as documented by Rivera (group 19, Cluster IV) with various samples of et al., 2013b). Another group clearly links archaeologi- P. canariensis, P. iberica, P. dactylifera land races of cal samples and West Asia, based on the association of south-eastern Spain and P. sylvestris. However, the archaeological materials with modern samples of lack of archaeological materials assigned to this mor- P. dactylifera from Iraq, Iran and the Arabian Penin- photype suggests that it has not contributed much to sula. However, as pointed out by Terral et al. (2012), the mainstream of modern commercial P. dactylifera morphological diversity appears to be only slightly cultivars. It should be underlined that archaeological structured according to the geographical origin of P. dactylifera samples represent almost exclusively, in P. dactylifera cultivars. The pairwise relationship, con- geographical terms, the Eastern chlorotype of cerning seed morphology, of P. theophrasti–P. reclinata Pintaud et al. (2013). Overall, Cluster IV shows is reflected in the association of Eastern Mediterra- remarkable geographical overlap with the Western nean to Tropical Africa. India, East Asia and Macaro- chlorotype of Pintaud et al. (2013). nesia appear, each one, relatively isolated. The Neolithic samples from different origins (Fig. 3, Table 3) present different Phoenix dactylifera morpho- Phoenix dactylifera in Baja California (Mexico) types (Table 7), but Neolithic samples fall only in At present, the major extant palm groves of Spanish groups 14–16 (Cluster III) and group 18 origin in the Americas are situated in Baja California. (Cluster IV) and Chalcolithic and Bronze Age Documentary evidence about the introduction of date samples fall in only group 15 (Cluster III) of P. dac- palms during the first century of the missions (i.e. the tylifera. Iron Age and later samples fall in 18th century) in Baja California is contradictory. groups 14–17 (Cluster III) and group 20 However, date palm cultivation was well established (Cluster IV) of P. dactylifera. Apparently, the great in several mission areas by the 1750s. (Rivera et al., diversity of P. dactylifera morphotypes during the 2013b). Preliminary molecular results show little Neolithic was followed, during the Chalcolithic and genetic diversity and suggest the introduction the Bronze Age, by a remarkable constriction (bottle- occurred once in one mission (presumably Loreto) and neck) in terms of morphological variability, which from there to the rest (Rivera et al., 2010). slowly recovered from the Iron Age onwards. This All Baja California seed samples fall in groups 12, bottleneck could be related to the genetic bottleneck 13 and 17 (Cluster III) and groups 18, 19 and 20 occurring between the 8th and 5th millennium BP in (Cluster IV), which include P. dactylifera cultivars the domestication model proposed for the date palm and related species. Seed morphology does not show a by Pintaud et al. (2013). geographical pattern in Baja California. Samples fall With the currently available evidence, we cannot in six of the 12 groups recognized for Clusters III and exclude the proposal made by Terral et al. (2012) IV. A few samples coincide in one group (Fig. 3), e.g. concerning a group ancestral to P. dactylifera in the group 12 (Cluster III): SIBC 3 (San Ignacio), Persian Gulf, related to the Eastern chlorotype of SIBC 15 (Loreto), SIBC 19 and 20 (La Purísima), Pintaud et al. (2013). Furthermore, in parallel SIBC 11, 12 and 22 (San José de Comondú). Thus, another group ancestral to P. dactylifera may exist in morphological diversity is relatively high, particularly

Figure 24. Hierarchical tree calculated with the Ward method representing relationships between large geographical zones based in the coincidence of samples of known origin in each one of the 25 groups recognized.

in San Ignacio. Concerning coincidences with species morphotypes represent the variability of P. dactylif- and cultivars from abroad it is to be noted that the era. Regarding the existence of continuity in time, relative morphological similarity between SIBC 11 nine morphotypes include only modern seed (San José de Comondú) and Phoenicites occidentalis samples, two morphotypes include modern and fossil from Eocene of Texas, the weighted neighbor joining samples, nine morphotypes include modern and tree shows 52% coincidence with 5000 bootstraps. archaeological samples and four morphotypes However, a factorial analysis (PCoA) shows this coin- include both modern and archaeological samples and cidence is superﬁcial. SIBC 20 (La Purisima) closely fossils. resembles Orisa 4 (Orihuela, Spain), and SIBC 23 Eight species have characteristic seeds and are (San Ignacio) coincides with Isla Plana 1 (Cartagena, clearly assigned to morphotypes [P. acaulis, P. canar- Spain). In relation to archaeobotanical materials, iensis s.s., P. paludosa, P. reclinata, P. roebelenii, SIBC 6 (San Ignacio) and SIBC 11 (San José de P. rupicola, P. sylvestris and P. theophrasti (excluding Comondú) show coincidences with Karanis 5 (Roman populations of Datça, Turkey)], but the others cannot period, Egypt). Overall, samples show high coincidence be clearly separated on the sole basis of the morphol- with samples from Elche and Abanilla–Fortuna ogy of seeds. (south-eastern Spain). Seeds from Mulegé (SIBC 16, Geographical patterns are detected for main groups 17) show coincidences with samples from the Near and clusters in the genus Phoenix. However, morpho- East. logical diversity and geographical origin of P. dactylifera cultivars are not clearly related. A major geographical western/eastern pattern related with CONCLUSIONS previously described chlorotypes is detected in P. dac- Seed morphology including shape or outline, total- tylifera. ized dimensions, superﬁcial processes, shape of With the currently available evidence, we cannot raphe, apex and base and position of micropyle, is exclude a group ancestral to P. dactylifera in the taxonomically useful. Twenty-four morphotypes were Persian Gulf, related to the Eastern chlorotype. Fur- differentiated into three major clusters. Of these, 12 thermore, in parallel, another group ancestral to

Figure 25. Distribution maps of Phoenix species. A, 1. P. canariensis;2.P. atlantica;3.P. iberica;4.P. theophrasti.B, 1. P. reclinata;2.P. sylvestris.C,P. dactylifera.D,P. caespitosa.E,P. loureiroi, 1. ‘Pedunculata’; 2. ‘Humilis’; 3. ‘Loureiroi’; 4. ‘Hanceana’. F, P. pusilla; 1. ‘Zeylanica’; 2. ‘Farinifera’; 3. P. acaulis.G,1.P. roebelenii;2.P. andamanensis. H, 1. P. paludosa. 2. P. rupicola.

P. dactylifera may exist in the Western Mediterra- Bakowski Z. 1967. Phoenix szaferi sp.nov. from the Podhale nean, including P. iberica, related to the Western Region and an outline of the history of the genus Phoenix L. chlorotype. Prace Muzeum Ziemi 10: 169–213. Apparently, the great diversity of date seeds mor- Barrow S. 1998. A monograph of Phoenix L. (Palmae: Cor- phology during the Neolithic was followed, during the yphoideae). Kew Bulletin 53: 513–575. Chalcolithic and the Bronze Age, by a remarkable Beccari O. 1890. Rivista monografica delle specie del genero constriction (bottleneck) in terms of morphological Phoenix L. Malesia 3: 345–416, f. 17, t. 43-44. variability of dates of P. dactylifera, which slowly Beech M. 2003. Archaeobotanical evidence for early date consumption in the Arabian Gulf. In: The Emirates Center recovered from the Iron Age onwards. for Strategic Studies and Research (ECSSR), ed. The date More detailed studies are needed of a greater palm. Abu Dhabi: Emirates Center for Strategic Studies number of clearly identified modern samples (espe- and Research, 11–31. cially with increased representation of diversity in Berry E. 1914. Fruits of a date palm in the Tertiary deposits South-East Asia) and archaeological samples (clearly of eastern Texas. American Journal of Science 37: 403–406. radiocarbon dated) to deepen the knowledge of mor- Berry E. 1937. Tertiary floras of eastern North America. phological variation patterns in time and space. For Botanical Review (Lancaster) 3: 31–46. this purpose, it is essential to increase the presence of Buzek C. 1977. Date-palm seeds from the Lower Miocene of specimens in herbaria, with well-preserved samples Central Europe. Ve˘stnik Ustrˇednîho Ústavu Geologickêho of dried seeds (not just fruit), in suitable condition to 52: 159–168. be studied (at least 30 seeds). Institutions that Cao B, Chao C. 2002. Identification of date cultivars in manage collections of modern plant material (her- California using AFLP markers. HortScience 37: 966– baria and repositories) and archaeological and fossil 968. (museums) should ensure that such materials are Caton G, Gardner E. 1932. The prehistoric geography of localizable and available to researchers. It would be Kharga Oasis. The Geographical Journal (London) 80: 369– especially useful to put online quality images of all 406. materials, especially nomenclatural types, as is Chabaud B. 1882. Le Phoenix canariensis. La Provence agri- already underway in some large herbaria. cole et horticole illustrée 19: 293–297, ff. 67–68. Costantini L. 1985. Considerazioni su alcuni reperti di palma da dattero e sul centro di origine e l’area di coltivazi- ACKNOWLEDGEMENTS one della Phoenix dactylifera L. Serie Orientale Roma 56: 209–217 + 1 tab. This research received, for the Spanish part, support Costantini L. 1991. Ecology and farming of the protohistoric from the INIA projects RF2007-00010-C03 and communities in the Central Yemeni highlands. In: Gnoli, G. RF2010-00006-C02 (European Regional Development ed. Reports and Memoirs 24: 187–204. Fund 2007–2013). For assistance in finding type and Costantini L, Audisio P. 2000. Plant and insect remains archaeological materials, we thank Dr Jiri Kvacek from the Bronze Age site of Ra’s al-Jinz (RJ-2), Sultanate of (Head of the Department of Palaeontology, National Oman. Paléorient 26: 143–158. Museum Prague), Dr Frank D. Steinheimer (Leitung Costantini L, Costantini-Biasini L. 1993. The plant Zentralmagazin Naturwissenschaftlicher Sammlun- remains. In: Wilhem G, Zaccagnini C, eds. Tell Karrana 3. gen, Martin-Luther-Universität Halle-Wittenberg), Baghdad / Mainz am Rhein: Deutsches Archaologisches Dr Mark Nesbitt (Royal Botanic Gardens, Kew), Dr Institut Abteilung – Verlag Philipp von Zabern, 237–250. Marina Clauser (Orto Botanico, Universitá degli Couvreur T, Forest F, Baker WJ. 2011. Origin and global Studi di Firenze), Dr Jacob Morales (Universidad de diversification patterns of tropical rain forests: inferences Las Palmas), Dr Anat Hartmann-Shenkman and Pro- from a complete genus-level phylogeny of palms. BMC fessor Mordechai Kislev (Bar Ilan University) and Dr Biology 9: 44–56. Maxine Kleindienst (Toronto University). Debono F. 1948. El-Omari (près d’Hélouan). Exposé som- maire sur les campagnes des foilles 1943–44 et 1948. Annales du Service des Antiquites de l’Égipte 48: 561–569. REFERENCES Dransfield J, Uhl NW, Asmussen CB, Baker WJ, Harley MM, Lewis CE. 2005. A new phylogenetic classification of Anguita F, Hernán F. 2000. The Canary Islands origin: a the palm family, Arecaceae. Kew Bulletin 60: 559–569. unifying model. Journal of Volcanology and Geothermal Dransfield J, Uhl NW, Asmussen CB, Baker WJ, Harley Research 103: 1–26. MM, Lewis CE. 2008. Genera Palmarum. The evolution Asmussen CB, Dransfield J, Deickmann V, Barfod AS, and classification of palms. Kew: Royal Botanic Gardens. Pintaud JC, Baker WJ. 2006. A new subfamily classifi- Friedrich W. 1980. Fossil plants from Weichselian intersta- cation of the palm family (Arecaceae): evidence from plastid dials, Santorini (Greece) II. In: Doumas C, ed. Thera and the DNA phylogeny. Botanical Journal of the Linnean Society Aegean World II. London: Thera and the Aegean World, 151: 15–38. 109–128.

Friedrich W, Pilcher H, Kussmaul S. 1977. Quaternary with particular reference to the Bilina Mine section. pyroclastics from Santorini/Greece and their significance for Journal of the Czech Geological Society 49: 1–40. the Mediterranean palaeoclimate. Bulletin Geological Levkovskaya G, Filatenko A. 1992. Palaeobotanical and Society Denmark 26: 27–39. palynological studies in South Arabia. Review of Palaeo- Fuller D. 2000. The botanical remains. In: Insoll T, ed. botany and Palynology 73: 241–257. Urbanism, archaeology and trade, further observations on Levy T. 1986. Archaeological sources for the study of Pales- the Gao Region (Mali). London: BAR International Series tine: the Chalcolithic Period. The Biblical Archaeologist 49: 829, 28–35. 82–108. Fuller D, Madella M. 2001. Issues in Harappan archaeo- Loret V. 1892. La flore pharaonique. 2a ed. Paris: E. Leroux. botany: retrospect and prospect. In: Settar S, Korisettar R, Mai D. 1976. Fossile Früchte und Samen aus dem Mittele- eds. Indian archaeology in retrospect. Vol. 2. Dehli: ozän des Geiseltales. Abhandlungen Zentrales Geologisches Manohar, 317–390. Institut 26: 93–149. Gaertner J. 1788–1791. De fructibus et seminibus plan- Meyer F. 1980. Carbonized food plants of Pompeii. Hercula- tarum. Stuttgart: Academiae Carolinae. neum, and the villa at Torre Annuziata. Economic Botany Gardner E. 1935. The Pleistocene fauna and flora of Kharga 34: 401–437. Oasis, Egypt. Quarterly Journal of the Geological Society of Miller N. 1981. Plant remains from Ville Royale II, Susa. London 91: 479–518. Cahiers de la Délégation Archéologique Française en Iran González M, Caujapé J, Sosa P. 2004. Molecular evidence 12: 137–142. of hybridization between the endemic Phoenix canariensis Moore HE. 1971. Phoenix canariensis and P. cycadifolia. and the widespread P. dactylifera with random amplified Principes 15: 33–35. polymorphic DNA (RAPD) markers. Plant Systematics and Moore HE, Dransfield J. 1979. The typification of Linnean Evolution 247: 165–175. palms. Taxon 28: 59–70. Govaerts R, Dransfield J, Zona SF, Hodel DR, Morales J, Navarro JF, Rodríguez A. 2011. Plant offerings Henderson A. 2011. World Checklist of Arecaceae. Kew: to the gods: seed remains from a pre-Hispanic sacrificial Facilitated by the Royal Botanic Gardens. Published on the altar in La Gomera Island (Canary Islands, Spain). In: Internet. Available at: http://apps.kew.org/wcsp/ (last Fahmy AG, Kahlheber S, D’Andrea C, eds. Windows on the accessed 2/6/2013). African past. Current approaches to African archaeobotany. Gros-Balthazard M. 2013. Hybridization in the genus Frankfurt: Africa Magna Verlag, 67–78. Phoenix: a review. Emirates Journal of Food and Agricul- Neef R, Cappers TJ, Bekker RM. 2011. Arecaceae in ture 25: 831–842. Digital Atlas of Economic Plants in Archaeology. Available Harley M. 2006. A summary of fossil records for Arecaceae. at: http://www.plantatlas.eu/pdfs/daap_arecaceae_72dpi.pdf Botanical Journal of the Linnean Society 151: 39–67. (last accessed 26/11/2012). Hopf M. 1983. Jericho plant remains. In: Kenyon KM, Nesbitt M. 1993. Archaeobotanical evidence for early Dilmun Holland TA, eds. Excavations at Jericho. Vol. V. London: The diet at Saar, Bahrain. Arabian Archaeology and Epigraphy British School of Archaeology in Jerusalem, 576–624. 4: 20–47. Huson DH, Scornavacca C. 2012. Dendroscope3–an Nixon R. 1950. Imported varieties of dates in the United States. interactive viewer for rooted phylogenetic trees and net- Washington, DC: United States Department of Agriculture. works. Systematic Biology 61: 1061–1067. doi: 10.1093/ Perrier X, Flori A, Bonnot F. 2003. Data analysis methods. sysbio/sys062. In: Hamon P, Seguin M, Perrier X, Glaszmann JC, eds. INIA. 2012. Phoenix. Available at: http://wwwx.inia.es/webcrf/ Genetic diversity of cultivated tropical plants. Montpellier: CRFesp/Paginaprincipal.asp (last accessed 20/12/2012). Enfield, Science Publishers, 43–76. Iossi E, Vitti F, Rubens S. 2006. Seed anatomy and germi- Perrier X, Jacquemoud-Collet JP. 2006. DARwin software. nation of Phoenix roebelenii O’Brien (Arecaceae). Revista Available at: http://darwin.cirad.fr/ (last accessed 26/11/ Brasileira de Sementes 28: 121–128. 2012). IPGRI. 2005. Descripteurs du palmier dattier (Phoenix dac- Phoenix Spain. 2013. Phoenix Spain Colección Nacional. tylifera L.). Rome: IPGRI. Available at: http://www.phoenix-spain.org (last accessed Jacquin NJ. 1801. Fragmenta botanica vol. 1. Vienna: 22/7/2013). Mathias Andreas Schmidt. Pintaud JC, Ludena B, Zehdi S, Gros-Balthazard M, Kaempfer E. 1712. Amoenitatum Exoticarum, Fasciculi V. Ivorra S, Terral JF, Newton C, Tengberg M, Santoni S, Leipzig: Heinrich Wilhelm Meyer. Boughedoura N. 2013. Biogeography of the date palm Kislev M, Hartmann A, Galili E. 2004. Archaeobotanical (Phoenix dactylifera L., Arecaceae): insights on the origin and archaeoentomological evidence from a well at Atlit-Yam and on the structure of modern diversity. Acta Horticulturae indicates colder, more humid climate on the Israeli coast 994: 19–36. during the PPNC period. Journal of Archaeological Science Pintaud JC, Zehdi S, Couvreur T, Barrow S, Henderson 31: 1301–1310. S, Aberlenc-Bertosi F, Tregear JW, Billotte N. 2010. Kvacek Z, Böhme M, Dvorak Z, Konzalova M, Mach K, Species delimitation in the genus Phoenix (Arecaceae) based Prokop J, Rajchl M. 2004. Early Miocene freshwater and on SSR markers, with emphasis on the identity of the date swamp ecosystems of the Most Basin (northern Bohemia) palm (Phoenix dactylifera L. In: Seberg O, Petersen G,

Barfod A, Davis J, eds. Diversity, phylogeny, and evolution Roxburgh W. 1796. Plants of the coast of Coromandel. Vol. 1. in the monocotyledons. Aarhus: Aarhus University Press, London: Bulmer and Co. 267–286. Roxburgh W. 1820. Plants of the coast of Coromandel. Vol. 3. Radwanski A. 2009. ‘Phoenix szaferi’ (palm fruitbodies) rein- London: Bulmer and Co. terpreted as traces of wood-boring terenid bivalves from the Saitou N, Nei M. 1987. The neighbor-joining method: a new Lower Oligocene (Rupelian) of the Tatra Mountains, Poland. method for reconstructing phylogenetic trees. Molecular Acta Palaeobotanica 49: 279–286. Biology and Evolution 4: 406–425. Rambaut A. 2012. FigTree v1.4 2006-2012. Institute of Evo- Sallon S, Solowey E, Cohen Y, Korchinsky R, Egli M, lutionary Biology, University of Edinburgh. http://tree.bio Woodhatch I, Simchoni O, Kislev M. 2008. Germination, .ed.ac.uk/software/figtree/ (last accessed 7/4/2014). genetics, and growth of an ancient date seed. Science 320: Renfrew J. 1987. Fruits from ancient Iraq: the paleoethno- 1464. botanical evidence. Bulletin on Sumerian Agriculture 3: Schweinfurth G. 1883. Neue Beiträge zur Flora des alten 157–161. Aegyptens. Berichte der Deutschen Botanischen Gesellschaft Rheede HV. 1682. Hortus Indicus Malabaricus 3. Amster- 1: 546–554. dam: J. van Someren and J. van Dyck. Stearn WT. 1978. Botanical Latin. Newton Abbot: David and Rivera D, Carreño E, Johnson D, Delgadillo J, Carrillo Charles. M, Obón C, Krueger R, Alcaraz F, Ríos S. 2010. His- Sudhersan C, Jibi S, Al-Sabah L. 2010. Xenic and metax- torical and molecular evidence for date palm introduction in enic effect of Phoenix pusilla pollen on certain date palm America (Phoenix dactylifera L., Arecaceae). Revista de cultivars. Acta Horticulturae 882: 297–302. Fitoterapia 10 (Suppl. 1): 108. Täckholm V, Drar M. 1950. Flora of Egypt Vol. 2. Cairo: Rivera D, Johnson D, Delgadillo J, Carrillo MH, Obón C, Fouad I University Press. Krueger R, Alcaraz F, Ríos S, Carreño E. 2013b. His- Tengberg M. 1999. Crop husbandry at Miri Qalat, Makran, torical evidence of the Spanish introduction of date palm SW Pakistan (4000–2000 BC). Vegetation History and (Phoenix dactylifera L., Arecaceae) into the Americas. Archaeobotany 8: 3–12. Genetic Resources and Crop Evolution 60: 1433–1452. Terral J, Newton C, Ivorra S, Gros M, Morais C, Picq S, Rivera D, Matilla G, Obón C, Alcaráz F. 2012a. Plants and Tengberg M, Pintaud J. 2012. Insights into the historical humans in the Near East and the Caucasus. Vols. 1, 2. biogeography of the date palm (Phoenix dactylifera L.) using Murcia: Editum & Plants and Humans. geometric morphometry of modern and ancient seeds. Rivera D, Obón C, Laguna E, Alcaraz F, Carreño E. Journal of Biogeography 39: 929–941. 2012b. Biogeografía de las palmeras datileras (Phoenix, Trimen H. 1898. A handbook to the Flora of Ceylon 4. Arecaceae) en la Cuenca Mediterránea. In: Cunill R, London: Dulau and Co. Pélachs A, Perez R, Soriano JM, eds. Las zonas de montaña: Uhl NW, Dransfield J. 1987. Genera palmarum: a classifi- gestión y biodiversidad. Sant Pere de Ribes: GRAMP, 111– cation of palms based on the work of Harold E. Moore, Jr. 117. Lawrence, Kansas: Allen Press. Rivera D, Obón C, Laguna E, Alcaraz F, Egea T, Carreño Uhl NW, Dransfield J, Davis JI, Luckow MA, Hansen KS, E, Laguna E, Santos A, Wildpret W. 2013a. A review of Doyle JJ. 1995. Phylogenetic relationships among palms: the nomenclature and typification of the Canary Islands cladistic analyses of morphological and chloroplast DNA endemic palm, Phoenix canariensis (Arecaceae). Taxon 62: restriction site variation. In: Rudall PJ, Cribb PJ, Cutler 1275–1282. DF, Humphries CJ, eds. Monocotyledons: systematics and Rögl VF. 1998. Palaeogeographic considerations for Mediter- evolution. Kew: Royal Botanic Gardens, 623–661. ranean and Paratethys seaways (Oligocene to Miocene). Willcox G, Tengberg M. 1995. Preliminary report on the Annalen Naturhistorisches Museum Wien 99: 279–310. archaeobotanical investigations at Tell Abraq with special Röpert D, ed. 2000. (continuously updated). Digital specimen attention to chaff impressions in mud brick. Arabian Arche- images at the Herbarium Berolinense. Published on the ology and Epigraphy 6: 129–138. Internet. Available at: http://ww2.bgbm.org/herbarium/ Wittmack H. 1903. Die in Pompeji gefundenen pflanzlichen default.cfm (last accessed 26/5/2013). Reste. Botanische Jahrbücher für Systematik, Pflanzenge- Rovira N, Chabal L. 2008. A foundation offering at the Roman schichte und Pflanzegeographie 33: 38–66. port of Lattara (Lattes, France): the plant remains. Vegeta- Zohary D, Hopf M. 2000. Domestication of plants in the Old tion History and Archaeobotany 17 (Suppl. 1): 191–200. World. 3rd edn. Oxford, UK: Oxford University Press.

SUPPORTING INFORMATION Additional Supporting Information may be found in the online version of this article at the publisher’s web-site: Appendix S1. List of modern Phoenix seed samples analysed from Herbaria, Germplasm repositories and Carpological Collections. Appendix S2. Crude matrix. Appendix S3. The matrix of correlation between variables.