Baker e Dransfield, 2016

Botanical Journal of the Linnean Society, 2016.
With 7 figures
Beyond Genera Palmarum: progress and prospects in

palm systematics
WILLIAM J. BAKER FLS* and JOHN DRANSFIELD FLS
Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, UK
Received 30 October 2015; revised 2 February 2016; accepted for publication 10 February 2016
The systematic biology of the palm family (Arecaceae) is probably better known than that of any other tropical
plant family of comparable size. As a result, the palms are now regarded as a model group for tropical rain forest
research. Ten years ago, the first phylogenetic classification of palms was established as a foundation for the
second edition of the palm systematic synthesis, Genera Palmarum (GP2), which was published in 2008. Here, we
review progress in palm systematics since GP2, summarizing the latest developments in an updated palm
classification and schematic phylogenetic tree. To date, the palms comprise 181 genera and c. 2600 species. In
just 8 years, six new genera and 200 new species have been described, whereas eight genera have been sunk into
synonymy. This reflects the highly dynamic nature of systematic discovery of palms at both forest and
phylogenetic frontiers. Palm phylogenetics is a vibrant field, with new trees being generated and utilized in
increasingly innovative and ambitious ways. Existing understanding of relationships among the five subfamilies
has been confirmed and deep nodes in the subfamilies are crystallizing, especially in subfamilies Arecoideae and
Coryphoideae. We conclude that palm systematic knowledge is far from complete and that tools, such as GP2,
only stimulate further scientific research and discovery. Despite recent advances, however, many aspects of the
palm Tree of Life still remain scarcely known. The vast datasets that the phylogenomic revolution is now
bringing to bear on palms promise to elucidate many of these unknowns. The ultimate goal, a species-level
phylogenetic tree for palms, is now coming within reach. © 2016 The Linnean Society of London, Botanical
Journal of the Linnean Society, 2016
ADDITIONAL KEYWORDS: Arecaceae – biodiversity – classification – next-generation sequencing –

Palmae – phylogenetics – phylogenomics – plant taxonomy – rainforest.
INTRODUCTION & Beck, 1990), their keystone role in ecosystems

(Eiserhardt et al., 2011b; ter Steege et al., 2013) or
Systematics advances in increments, with each step
perhaps merely to their charisma and the scientific
intended as a successive approximation towards the
challenge that they present. The work of generations
optimal solution. Palms (Arecaceae) are no exception
of such palm specialists has laid the foundations of
to this trend. Such has been the success of the palm
palm biology today.
systematic endeavour that few, if any, tropical plant
families of comparable size rival palms in terms of
fundamental systematic resources (Govaerts &
Dransfield, 2005; Dransfield et al., 2008b; Baker A BRIEF HISTORY OF GENERA PALMARUM
et al., 2009; Baker & Couvreur, 2013a; Govaerts
Following the remarkable contributions of 19th and
et al., 2015). As a result, palms are now widely
early 20th century innovators, such as Martius, Grif-
regarded as a model group for tropical plant research
fith, Blume, J. D. Hooker, Beccari and Burret (sum-
(Eiserhardt et al., 2011b; Couvreur & Baker, 2013).
marized in Dransfield et al., 2008b), the modern
How have palms, which are notoriously difficult to
palm systematic era dawned with the work of Harold
study, become so well known? Put simply, over dec-
E. Moore Jr. (1917–1980). From his base in the L. H.
ades and centuries, palms have attracted specialists
Bailey Hortorium at Cornell University, Moore
who are drawn to their economic importance (Balick
became the leading light in palm research from the
1950s to the 1970s. An avid fieldworker, he explored
*Corresponding author. E-mail: w.baker@kew.org for palms across the globe, studying all but 12 of the
© 2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016 1
2 W. J. BAKER and J. DRANSFIELD
palm genera known at the time in the wild. Building Cornell University in 2004, the authors constructed
on his taxonomic experience and his collaborations a new classification of palms as a foundation for the
with Cornell anatomist Natalie Uhl, Moore aimed to new Genera Palmarum, drawing on all available
understand the evolution of palms (Moore & Uhl, phylogenetic evidence, which was published the fol-
1973, 1982) and to complete an account of all palm lowing year (Dransfield et al., 2005). The resulting
genera, a Genera Palmarum. In 1973, Moore pub- book, Genera Palmarum – the Evolution and Classi-
lished the ‘Major Groups’ paper (Moore, 1973), in fication of Palms (Dransfield et al., 2008b), now
which he proposed an informal classification compris- known widely as GP2, is not simply a revised edi-
ing 15 groups in five major lines of evolution, based tion of GP1, but a completely new account of the
on explicit interpretations of morpho-anatomical palms. At 744 pages, GP2 contains c. 40% more con-
specialization. This paper has since served as an tent than GP1, including rewritten treatments for
important baseline for a truly phylogenetic classifica- the 183 accepted genera, significant additions on
tion of palms. phylogeny, biogeography, the fossil record and pol-
In 1980, Moore died unexpectedly, leaving many len, and expanded illustrations, photographs, maps
materials for his Genera Palmarum, especially illus- and glossary.
trations, but the book itself remained essentially The phylogenetic classification of GP2 employs
unwritten. The task passed to Natalie Uhl and John monophyly as the primary criterion for the delimita-
Dransfield, who critically reassessed Moore’s group- tion of taxa and topological evidence to determine a
ings, formalizing a new classification of six subfami- logical linear taxonomic sequence. As a result, the
lies, 14 tribes and 36 subtribes, based on classification is substantially different from that of
evolutionary concepts of the time (Dransfield & Uhl, GP1. Major realignments were made in the five sub-
1986). This formed the foundation of Genera Pal- families of the new classification (Asmussen et al.,
marum, a classification of palms based on the work 2006). Coryphoideae was expanded to include Cary-
of Harold E. Moore Jr. (Uhl & Dransfield, 1987), or oteae (formerly in Arecoideae), and Chamaedoreeae
GP1, for short. The title defers to Moore’s contribu- (formerly in Ceroxyloideae under the name Hyophor-
tion, but it should be stressed that this 610-page beae) was moved to Arecoideae. Ceroxyloideae was
book was an entirely new work, with original re-delimited to include Phytelephantoideae of GP1
descriptions of all 200 genera accepted at the time, (as Phytelepheae) with tribes Cyclospatheae and
introductory chapters and other materials produced Ceroxyleae. Significant changes were made at tribal
by Uhl and Dransfield. At the time, no comparably and subtribal levels, such as the re-delimitation of
detailed or complete work for palms had been pub- Areceae and the subtribes within it. Further details
lished since the legendary three-volume Historia of the correspondence between the classifications of
Naturalis Palmarum of Martius (von Martius, 1823– GP2, GP1 and earlier schemes can be found in
1850). Its influence is clear from the amount of palm Chapter 9 of GP2.
research that it spawned, as indicated by the c. 800 Ten years on, it is time to reflect on the classifica-
citations of GP1 recorded by Google Scholar today. tion underpinning GP2 and the developments that
In a sense, GP1 became a victim of its own suc- have taken place in palm systematics since its first
cess. With the mass of new information about palms publication (Dransfield et al., 2005), as Uhl and
arising from research inspired by GP1, a complete Dransfield did 10 years after GP1 (Uhl & Dransfield,
revision became necessary (Uhl & Dransfield, 1999). 1999). Having set the scene, in the remainder of this
A version of the GP1 classification, with updated paper we review developments in two main areas, (1)
generic taxonomy, was published in Families and palm classification and (2) phylogenetics, providing
Genera of Vascular Plants (Dransfield & Uhl, 1998), an extensive bibliography of the latest palm research
but the rapid rise of molecular phylogenetics and concluding with a discussion of future prospects
pointed to the need for a much more radical revi- for palm systematics.
sion (Uhl et al., 1995; Baker et al., 1999a; Asmus-
sen, Baker & Dransfield, 2000; Asmussen & Chase,
2001). In the early 2000s, a project to produce a
PALM CLASSIFICATION
new Genera Palmarum took shape during a succes-
sion of meetings hosted by the Montgomery Botani- The delimitation of the five subfamilies, 28 tribes
cal Center. Recognizing the opportunity to build a and 27 subtribes recognized in the GP2 classification
classification around a formal phylogenetic frame- has, so far, been robust to new phylogenetic findings
work, Dransfield and Uhl drew in new collaborators, and, to date, no amendments to the higher level clas-
especially those active in the field of palm phyloge- sification have been proposed [with the exception of
netics (Conny Asmussen-Lange, William Baker, the correction of the subtribe name Linospadicinae
Madeline Harley and Carl Lewis). At a summit in to Laccospadicinae (Dransfield et al., 2011); Table 1].
© 2016 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016
PALM SYSTEMATICS: PROGRESS AND PROSPECTS 3
Table 1. Summary of change in taxon numbers in the genera have resulted from novel discoveries in the
classification of palms Column 2 indicates the taxa recog- field, phylogenetic evidence or a combination of both.
nized in the first publication of the phylogenetic classifi- In contrast, the sinking of genera has been driven
cation of palms (Dransfield et al., 2005), updated and purely by phylogenetics and findings of non-mono-
refined in Genera Palmarum (GP2; Dransfield et al., phyly. The overwhelming majority of the generic
2008b). Column 3 reflects the status of palm classification gains and losses are in palms of the Asian Pacific,
in October 2015 (see text). Species numbers derived from reflecting the relative intensity of systematic
Govaerts et al. (2015). research on palms in this region. A full classification
of palms is presented here (Table 3), updated to
2008 2015
account for all changes to date.
Subfamilies 5 5
Tribes 28 28
Subtribes 27 27 GENERA GAINED SINCE GP2
Genera 183 181
Unplaced genera 17 17 The generic treatments provided here are structured
Species c. 2400 c. 2600 to be consistent with the generic treatments in GP2,
with some abbreviations. A ‘fossil record’ section is
excluded as fossils have not been linked to any of the
Although modifications in the future cannot be ruled new genera at this time.
out, the GP2 suprageneric classification appears to
be meeting its stated goals of being phylogenetically Sabinaria R.Bernal & Galeano, Phytotaxa 144: 28
informative and stable. (2013). Type: Sabinaria magnifica Galeano & R.Ber-
At the genus level, however, there has been consid- nal (Fig. 1).
erable change. Currently, 181 genera are accepted, a Etymology: Named after the daughter of the
net decrease of only two from the 183 recognized by authors, Sabina Bernal Galeano.
Dransfield et al. (2008b), but this conceals the fact Taxonomic account and description: Galeano &
that six additional genera are now accepted, whereas Bernal (2013).
eight have been placed in synonymy (Table 2). The Distribution and ecology: One species known only
recognition of six new genera in the 8 years since from the Serranıa del Dari en in north-western
the publication of GP2 in 2008 is striking, given that Colombia, but probably also occurring in neighbour-
only seven genera were added in the 21 years ing areas of Panama. Primary, tropical rain forest
between the publication of GP1 and GP2: Voanioala between 100 m and 250 m.
J.Dransf. (Dransfield, 1989), Aphandra Barfod (Bar- Anatomy: Pollen (Bogot a-A et al., 2015).
fod, 1991), Lemurophoenix J.Dransf. (Dransfield, Relationships: Sabinaria belongs to Cryosophileae
1991), Satranala J.Dransf. & Beentje (Dransfield & (Coryphoideae). It is well supported as sister to Itaya
Beentje, 1995b), Dransfieldia W.J.Baker & Zona H.E.Moore (A. Cano, unpubl. data).
(Baker et al., 2006), Leucothrinax C.Lewis & Zona Common names and uses: Girasol (sunflower in
(Lewis & Zona, 2008) and Tahina J.Dransf. & Rako- Spanish). Leaves used occasionally for umbrellas or
toarin. (Dransfield et al., 2008a). The additional thatch.
Notes: Known only from a small, isolated area in
Table 2. Changes in accepted palm genera since the pub- the Darien Gap, Sabinaria is probably the most spec-
lication of Genera Palmarum (GP2; Dransfield et al., tacular of the new genera discovered since GP2
2008b) (Galeano & Bernal, 2013; Bernal, 2014). Colombia
has been rather well explored for palms in recent
Subfamily Genera gained Genera lost decades on account of the scientific leadership of two
of South America’s most prolific palm biologists,
Calamoideae Ceratolobus Rodrigo Bernal and Gloria Galeano. Thus, the dis-
Daemonorops covery by Norman Echavarrıa and Sa ul Hoyos,
Pogonotium botanical collaborators of Bernal and Galeano, of
Retispatha such a conspicuous and unusual new genus was
Coryphoideae Lanonia Pritchardiopsis
highly surprising. The spectacular leaves of Sabina-
Saribus Wallichia
ria are its most distinctive character, being divided
Sabinaria
almost in half by a deep median, abaxial split (in
Arecoideae Jailoloa Lytocaryum
common with other Cryosophileae, such as Cryoso-
Manjekia Solfia
phila Blume or Chelyocarpus Dammer), whereas
Wallaceodoxa
the remainder of the lamina margin is almost
Table 3. Revised classification of Arecaceae, updated to incorporate changes made since Genera Palmarum (GP2;
Dransfield et al., 2008b). In the interests of stability, the linear sequence of genera has not been revised here because of
the large amount of palm phylogenetic research in progress at the moment, and is consistent with the prevailing
accounts (Dransfield et al., 2005, 2008b; Trias-Blasi et al., 2015). Changes are indicated with superscript letters and
summarized as follows. ACalamus, expanded to include Ceratolobus, Daemonorops, Pogonotium and Retispatha (Baker,
2015; Henderson & Floda, 2015). BSabinaria, new genus (Galeano & Bernal, 2013). CLivistona, re-delimited, species
transferred to Saribus (Bacon & Baker, 2011). DLicuala, re-delimited, species transferred to Lanonia (Henderson &
Bacon, 2011). ESaribus, resurrected and re-delimited to include part of Livistona and monotypic Pritchardiopsis (Bacon
& Baker, 2011). FLanonia, new genus (Henderson & Bacon, 2011). GArenga, expanded to include Wallichia (Jeanson,
2011). HSyagrus, expanded to include Lytocaryum (Noblick & Meerow, 2015). ILaccospadicinae, subtribal name corrected
from Linospadicinae (Dransfield et al., 2011). JPonapea, expanded to include part of Drymophloeus (Zona et al., 2011).
K
Balaka, expanded to include Solfia (Zona & Baker, 2014). LVeitchia, expanded to include part of Drymophloeus (Zona
et al., 2011). MDrymophloeus, re-delimited, species transferred to Ponapea and Veitchia (Zona et al., 2011). N–PJailoloa,
Manjekia, Wallaceodoxa, new genera (Heatubun et al., 2014b).
SUBFAMILY
Tribe
Subtribe Genus
CALAMOIDEAE
Eugeissoneae 1. Eugeissona Griff.
Lepidocaryeae
Ancistrophyllinae 2. Oncocalamus (G.Mann & H.Wendl.) H.Wendl.
3. Eremospatha (G.Mann & H.Wendl.) Schaedtler
4. Laccosperma (G.Mann & H.Wendl.) Drude
Raphiinae 5. Raphia P.Beauv.
Mauritiinae 6. Lepidocaryum Mart.
7. Mauritia L.f.
8. Mauritiella Burret
Calameae
Korthalsiinae 9. Korthalsia Blume
Salaccinae 10. Eleiodoxa (Becc.) Burret
11. Salacca Reinw.
Metroxylinae 12. Metroxylon Rottb.
Pigafettinae 13. Pigafetta (Blume) Becc.
Plectocomiinae 14. Plectocomia Mart. & Blume
15. Myrialepis Becc.
16. Plectocomiopsis Becc.
Calaminae 17. Calamus L.A
NYPOIDEAE 18. Nypa Steck
CORYPHOIDEAE
Sabaleae 19. Sabal Adans.
Cryosophileae 20. Schippia Burret

21. Trithrinax Mart.
22. Zombia L.H.Bailey
23. Coccothrinax Sarg.
24. Hemithrinax Hook.f.
25. Leucothrinax C.Lewis & Zona
26. Thrinax L.f. ex Sw.
27. Chelyocarpus Dammer
28. Cryosophila Blume
29. Itaya H.E.Moore
30. Sabinaria R.Bernal & GaleanoB
Table 3. Continued
SUBFAMILY
Tribe
Subtribe Genus
Phoeniceae 31. Phoenix L.

Trachycarpeae
Rhapidinae 32. Chamaerops L.
33. Guihaia J.Dransf., S.K.Lee & F.N.Wei
34. Trachycarpus H.Wendl.
35. Rhapidophyllum H.Wendl. & Drude
36. Maxburretia Furtado
37. Rhapis L.f. ex Aiton
Livistoninae 38. Livistona R.Br.C
39. Licuala Wurmb.D
40. Johannesteijsmannia H.E.Moore
41. Pholidocarpus Blume
42. Saribus BlumeE
43. Lanonia A.J.Hend. & C.D.BaconF
Unplaced Trachycarpeae 44. Acoelorrhaphe H.Wendl.
45. Serenoa Hook.f.
46. Brahea Mart.
47. Colpothrinax Schaedtler
48. Copernicia Mart. ex Endl.
49. Pritchardia Seem. & H.Wendl.
50. Washingtonia H.Wendl.
Chuniophoeniceae 51. Chuniophoenix Burret

52. Kerriodoxa J.Dransf.
53. Nannorrhops H.Wendl.
54. Tahina J.Dransf. & Rakotoarin.
Caryoteae 55. Caryota L.

56. Arenga Labill. ex DC.G
Corypheae 57. Corypha L.

Borasseae
Hyphaeninae 58. Bismarckia Hildebr. & H.Wendl.
59. Satranala J.Dransf. & Beentje
60. Hyphaene Gaertn.
61. Medemia W€ urttemb. ex H.Wendl.
Lataniinae 62. Latania Comm. ex Juss.
63. Lodoicea Comm. ex DC.
64. Borassodendron Becc.
65. Borassus L.
CEROXYLOIDEAE
Cyclospatheae 66. Pseudophoenix H.Wendl. ex Sarg.
Ceroxyleae 67. Ceroxylon Bonpl. ex DC.

68. Juania Drude
69. Oraniopsis (Becc.) J.Dransf., A.K.Irvine & N.W.Uhl
70. Ravenea H.Wendl. ex C.D.Bouch e
Phytelepheae 71. Ammandra O.F.Cook

72. Aphandra Barfod
73. Phytelephas Ruiz & Pav.
Table 3. Continued
SUBFAMILY
Tribe
Subtribe Genus
ARECOIDEAE
Iriarteeae 74. Iriartella H.Wendl.
75. Dictyocaryum H.Wendl.
76. Iriartea Ruiz & Pav.
77. Socratea H.Karst.
78. Wettinia Poepp. ex Endl.
Chamaedoreeae 79. Hyophorbe Gaertn.
80. Wendlandiella Dammer
81. Synechanthus H.Wendl.
82. Chamaedorea Willd.
83. Gaussia H.Wendl.
Podococceae 84. Podococcus G.Mann & H.Wendl.
Oranieae 85. Orania Zipp.
Sclerospermeae 86. Sclerosperma G.Mann & H.Wendl.
Roystoneeae 87. Roystonea O.F.Cook

Reinhardtieae 88. Reinhardtia Liebm.
Cocoseae
Attaleinae 89. Beccariophoenix Jum. & H.Perrier
90. Jubaeopsis Becc.
91. Voanioala J.Dransf.
92. Allagoptera Nees in M.A.P.zu Wied-Neuwied
93. Attalea Kunth
94. Butia (Becc.) Becc.
95. Cocos L.
96. Jubaea Kunth
97. Syagrus Mart.H
98. Parajubaea Burret
Bactridinae 99. Acrocomia Mart.
100. Astrocaryum G.Mey.
101. Aiphanes Willd.
102. Bactris Jacq. ex Scop.
103. Desmoncus Mart.
Elaeidinae 104. Barcella (Trail) Drude
105. Elaeis Jacq.
Manicarieae 106. Manicaria Gaertn.
Euterpeae 107. Hyospathe Mart.

108. Euterpe Mart.
109. Prestoea Hook.f.
110. Neonicholsonia Dammer
111. Oenocarpus Mart.
Geonomateae 112. Welfia H.Wendl.

113. Pholidostachys H.Wendl. ex Hook.f.
114. Calyptrogyne H.Wendl.
115. Calyptronoma Griseb.
116. Asterogyne H.Wendl. ex Hook.f.
117. Geonoma Willd.
Table 3. Continued
SUBFAMILY
Tribe
Subtribe Genus
Leopoldinieae 118. Leopoldinia Mart.

Pelagodoxeae 119. Pelagodoxa Becc.
120. Sommieria Becc.
Areceae
Archontophoenicinae 121. Actinorhytis H.Wendl. & Drude
122. Archontophoenix H.Wendl. & Drude
123. Actinokentia Dammer
124. Chambeyronia Vieill.
125. Kentiopsis Brongn.
Arecinae 126. Areca L.
127. Nenga H.Wendl. & Drude
128. Pinanga Blume
Basseliniinae 129. Basselinia Vieill.
130. Burretiokentia Pic.Serm.
131. Cyphophoenix H.Wendl. ex Hook.f
132. Cyphosperma H.Wendl. ex Hook.f
133. Lepidorrhachis (H.Wendl. & Drude) O.F.Cook
134. Physokentia Becc.
Carpoxylinae 135. Carpoxylon H.Wendl. & Drude
136. Satakentia H.E.Moore
137. Neoveitchia Becc.
Clinospermatinae 138. Cyphokentia Brongn.
139. Clinosperma Becc.
Dypsidinae 140. Dypsis Noronha ex Mart.
141. Lemurophoenix J.Dransf.
142. Marojejya Humbert
143. Masoala Jum.
LaccospadicinaeI 144. Calyptrocalyx Blume
145. Linospadix H.Wendl.
146. Howea Becc.
147. Laccospadix H.Wendl. & Drude
Oncospermatinae 148. Oncosperma Blume
149. Deckenia H.Wendl. ex Seem.
150. Acanthophoenix H.Wendl.
151. Tectiphiala H.E.Moore
Ptychospermatinae 152. Ptychosperma Labill.
153. Ponapea Becc.J
154. Adonidia Becc.
155. Balaka Becc.K
156. Veitchia H.Wendl.L
157. Carpentaria Becc.
158. Wodyetia A.K.Irvine
159. Drymophloeus Zipp.M
160. Normanbya F.Muell. ex Becc.
161. Brassiophoenix Burret
162. Ptychococcus Becc.
163. Jailoloa Heatubun & W.J.BakerN
164. Manjekia W.J.Baker & HeatubunO
165. Wallaceodoxa Heatubun & W.J.BakerP
Table 3. Continued
SUBFAMILY
Tribe
Subtribe Genus
Rhopalostylidinae 166. Rhopalostylis H.Wendl. & Drude

167. Hedyscepe H.Wendl. & Drude
Verschaffeltiinae 168. Nephrosperma Balf.f.
169. Phoenicophorium H.Wendl.
170. Roscheria H.Wendl. ex Balf.f.
171. Verschaffeltia H.Wendl.
Unplaced Areceae 172. Bentinckia Berry ex Roxb.
173. Clinostigma H.Wendl.
174. Cyrtostachys Blume
175. Dictyosperma H.Wendl. & Drude
176. Dransfieldia W.J.Baker & Zona
177. Heterospathe Scheff.
178. Hydriastele H.Wendl. & Drude
179. Iguanura Blume
180. Loxococcus H.Wendl. & Drude
181. Rhopaloblaste Scheff.
A B
Figure 1. Sabinaria magnifica, Serranıa del Dari en, Colombia. A, Habit, with the eponymous Sabina Bernal Galeano
providing scale. B, Infructescence, rachis bracts parted to reveal fruits. Photographs: W. J. Baker.
A B
Figure 2. Lanonia gracilis, cultivated, Kebun Raya, Bogor, Indonesia. A, Habit. B, Inflorescence. Photographs: W. J.
Baker.
entire, being only shallowly lobed by short splits. In Taxonomic account and description: Henderson &
addition, the lamina is glossy green above, but Bacon (2011).
silver-white below. Distribution and ecology: Eight species, six from Viet-
Sabinaria most closely resembles its sister genus nam, two of which reach into adjacent Laos and China,
Itaya as they both bear a biseriate perianth, gynoecia respectively, one from Hainan and one from Java. Pri-
comprising a single carpel and androecia with numer- mary, tropical rain forest from low elevation to 1700 m.
ous stamens. Sabinaria is, however, distinguished Anatomy: Not studied.
from Itaya by its unisexual flowers, the pistillate flow- Relationships: Lanonia belongs to Livistoninae
ers being located at the base of the lower rachillae (Coryphoideae: Trachycarpeae). It is strongly sup-
only, and the persistent rachis bracts that enclose the ported as a monophyletic genus that is moderately to
rachillae (especially at the base) and tightly ensheath strongly supported as sister to Johannesteijsmannia
the developing fruit. In contrast, Itaya has hermaph- H.E.Moore (Henderson & Bacon, 2011; Bacon, Baker
roditic flowers and deciduous rachis bracts. The flow- & Simmons, 2012a; Bacon et al., 2013a).
ers of Sabinaria are also unique in Cryosophileae in Common names and uses: Hat palm or la non
that the two whorls of the perianth, although distinct, (Vietnam). At least one species [L. centralis
are connate at a single point of contact. (A.J.Hend., N.K.Ban & N.Q.Dung) A.J.Hend. &
C.D.Bacon] used for making conical hats in Vietnam.
Lanonia A.J.Hend. & C.D.Bacon, Syst. Bot. 36: For further details on common names and uses, see
887 (2011). Type: Lanonia acaulis (A.J.Hend., Henderson & Bacon (2011).
N.K.Ban & N.Q.Dung) A.J.Hend. & C.D.Bacon Notes: Lanonia was described as a segregate of the
(basionym: Licuala acaulis A.J.Hend., N.K.Ban & Asia-Pacific genus Licuala Wurmb. by Andrew Hen-
N.Q.Dung) (Fig. 2). derson and Christine Bacon (2011). Comprehensive
Etymology: Based on a Vietnamese local name la phylogenetic research on Trachycarpeae (Henderson
non, meaning hat palm, referring to its use in the & Bacon, 2011; Bacon et al., 2012a, 2013a) revealed
manufacture of the typical conical hats of the region. that a group of species described in Licuala is in fact
A B
Figure 3. Saribus. A, S. merrillii, habit, cultivated. B, S. rotundifolius, trifurcate inflorescence base, cultivated, Kebun
Raya, Bogor, Indonesia. Photographs: C. E. Lewis and W. J. Baker.
more closely related to Johannesteijsmannia than it early 20th centuries, remarkably, the remaining four
is to Licuala, and required recognition at generic are recent discoveries described since 2007.
rank. This was surprising because it showed that the
unique Licuala-type leaf is not restricted to a single Saribus Blume, Rumphia 2: 48 (1838). Lectotype:
genus as previously thought, but shared among two Saribus rotundifolius (Lam.) Blume (basionym: Cory-
monophyletic lineages that are not sister taxa, i.e. pha rotundifolia Lam.) (Fig. 3).
Lanonia and Licuala s.s. Licuala-type leaves are, in Pritchardiopsis Becc., Webbia 3: 131 (1910). Type:
essence, typical induplicately divided, palmate cory- Pritchardiopsis jeanneneyi Becc. [= Saribus jeanne-
phoid leaves that are further subdivided to the base neyi (Becc.) C.D. Bacon & W.J.Baker].
into wedge-shaped segments by a number of deep, Etymology: Latinization of a Moluccan plant name,
secondary, splits along abaxial folds (Dransfield saribu.
et al., 2008b). The leaves of Lanonia are further Taxonomic account and description: Eight species
characterized by the central segment being split to treated in a monograph of Livistona R.Br. (Dowe,
the apex of a short costa, which bears a pulvinus-like 2009) and a ninth in Pritchardiopsis (Hodel & Pin-
structure on its abaxial surface. taud, 1998). A description of the genus is not
A number of reproductive characters further dis- currently available.
tinguish Lanonia from the superficially similar Distribution and ecology: Nine species, six endemic
Licuala and its sister genus Johannesteijsmannia. to Papuasia (New Guinea and Solomon Islands), one
Lanonia is typically dioecious, often with sexually endemic to the Philippines, one endemic to New Cale-
dimorphic inflorescences, and bears staminate flow- donia and one species that is widespread in central
ers in pairs or clusters and solitary pistillate flowers. Malesia (Philippines to Raja Ampat). Various kinds of
The staminate flowers contain dorsifixed, non-versa- tropical, ever-wet forest from lowlands to 1300 m.
tile anthers, whereas the perianth of the pistillate Anatomy: Leaf (Tomlinson, Horn & Fisher, 2011).
flowers reflexes in fruit, the calyx splitting into six Relationships: Saribus belongs to Livistoninae
lobes after anthesis. Although half of the included (Coryphoideae: Trachycarpeae). It is strongly sup-
species are based on taxa described in the 19th and ported as a monophyletic genus and is resolved as
A B
Figure 4. Jailoloa halmaherensis, Halmahera, Indonesia. A, Habit. B, Fruit. Photographs: C. D. Heatubun.
sister to Pholidocarpus Blume, although with only taking priority. It should be noted that, in the
weak support (Bacon & Baker, 2011; Bacon et al., most recent monograph of Livistona (Dowe, 2009),
2012a, 2013a). the species now included in Saribus were high-
Common names and uses: Numerous local names lighted as a distinct group of close relatives,
reported by Dowe (2009). Uses include leaves for although a connection to Pritchardiopsis was not
thatch and edible palm hearts. Saribus rotundifolius made.
is an important ornamental in the tropics. The group is unique among other genera of Livis-
Notes: Although the generic name Saribus is toninae in its trifurcate inflorescences, which com-
far from new (Blume, 1838), the concept of this prise three (sometimes two) equal primary axes
resurrected genus, as proposed by Christine Bacon within a common prophyll (in contrast with the sin-
and William Baker (Bacon & Baker, 2011), is sig- gle axis of all other Livistoninae). In addition, the
nificantly different from any used previously. As mature fruits of Saribus spp. are orange, orange–
with Lanonia, the evidence for the addition of a brown or red, whereas Livistona fruits are green,
further coryphoid genus in Livistoninae stemmed blue, purple, brown or black. A number of leaf
from phylogenetic studies of Trachycarpeae (Bacon anatomical characters have also been identified for
et al., 2012a, 2013a), which showed that Livistona Saribus (Tomlinson et al., 2011).
(sensu Dransfield et al., 2008b; and Dowe, 2009) The removal of Saribus accentuates the disjunct
was not monophyletic, but divided into two sepa- distribution of Livistona s.s., which occurs in
rate clades. The first clade comprised the majority South-East Asia only west of Wallace’s Line, Aus-
of Livistona as sister to all other genera of Livis- tralia, the horn of Africa and the Arabian Peninsula
toninae. The remaining Livistona spp. formed the (Bacon & Baker, 2011).
second, smaller clade, which was resolved as sister
to Pholidocarpus. This segregate group, in which Jailoloa Heatubun & W.J.Baker, Kew Bull. 69
the New Caledonian monotypic genus Pritchardiop- (9525): 5 (2014). Type: Jailoloa halmaherensis (Hea-
sis was also nested, was thus delimited as a new tubun) Heatubun & W.J.Baker (basionym: Ptychos-
generic concept for which two genus names already perma halmaherense Heatubun) (Fig. 4).
existed: Saribus, typified on S. rotundifolius (Lam.) Etymology: Refers to Jailolo, an earlier name for
Blume, and Pritchardiopsis, the former, older name the island of Halmahera.
A B
Figure 5. Manjekia maturbongsii, Biak, Indonesia. A, Habit. B, Inflorescence. Photographs: W. J. Baker.
Taxonomic account and description: Heatubun, taining the sister genera Veitchia H.Wendl. and
Zona & Baker (2014b). Balaka Becc. (Alapetite et al., 2014). The three new
Distribution and ecology: One species from Halma- genera reflect this phylogenetic topology and mor-
hera. Heath forest on ultramafic rocks at c. 550 m. phological disparities, whilst maintaining taxonomic
Anatomy: Not studied. stability of existing genera (Heatubun et al., 2014b).
Relationships: Jailoloa belongs to Ptychospermati- The three new genera are similar to Adonidia in
nae (Arecoideae: Areceae). It is moderately supported their endocarp and seed characters (terete seed with
as sister to Manjekia W.J.Baker & Heatubun (Alape- straw-coloured fibres adhering to the endocarp),
tite, Baker & Nadot, 2014). which have been considered to be systematically use-
Common names and uses: None recorded. ful (Zona, 1999), but they are abundantly distinct in
Notes: The publication of any new palm genus is a other respects.
significant event, but the simultaneous description Jailoloa is a slender palm of ultramafic heath for-
of three new genera is unprecedented in recent dec- est, known only from one site in a nickel mining con-
ades (Heatubun et al., 2014b, c). Jailoloa, Manjekia cession in Halmahera. It is distinguished from other
and Wallaceodoxa Heatubun & W.J.Baker were genera of Ptychospermatinae by its recurved leaves
described as a result of fieldwork in eastern Indone- with leathery ascending leaflets, purple inflorescence
sia led by Charlie Heatubun and William Baker to axes and flower and orange–yellow fruit. Ptychos-
smaller islands to the north and west of the western perma, the most superficially similar genus, does not
end of New Guinea. The first two genera were ini- have ascending leaflets or the same combination of
tially described in Ptychosperma Labill. and Adoni- inflorescence and fruit colours, and usually has
dia Becc., respectively, based on morphological and ridged endocarps and seeds.
DNA evidence (Heatubun, 2011a; Zona et al., 2011;
Baker & Heatubun, 2012). These decisions were Manjekia W.J.Baker & Heatubun, Kew Bull. 69
reconsidered as a result of subsequent work on the (9525): 9 (2014). Type: Manjekia maturbongsii
molecular phylogenetics of Ptychospermatinae, (W.J.Baker & Heatubun) W.J.Baker & Heatubun
which placed the three taxa with Adonidia in a (basionym: Adonidia maturbongsii W.J.Baker &
paraphyletic group at the base of a clade also con- Heatubun) (Fig. 5).
A B
Figure 6. Wallaceodoxa raja-ampat, Waigeo, Indonesia. A, Habit. B, Inflorescence. Photographs: C. D. Heatubun.
Etymology: Based on a local name Manjek (Biak Wallaceodoxa Heatubun & W.J.Baker, Kew Bull.
dialect). 69 (9525): 9 (2014). Type: Wallaceodoxa raja-ampat
Taxonomic account and description: Heatubun Heatubun & W.J.Baker (Fig. 6).
et al. (2014b). Etymology: The name commemorates Alfred Russel
Distribution and ecology: One species from Biak Wallace (1823–1913), renowned British naturalist,
Island. Lowland rain forest on limestone up to 170 m. explorer of the Malay Archipelago and Amazon, co-
Anatomy: Not studied. discoverer of the theory of evolution by natural selec-
Relationships: Manjekia belongs to subtribe Pty- tion and author of the first field guide to palms (Wal-
chospermatinae (Arecoideae: Areceae). It is moder- lace, 1853). Suffix –doxa means ‘glory’.
ately supported as sister to Jailoloa (Alapetite et al., Taxonomic account and description: Heatubun
2014). et al. (2014b).
Common names and uses: Manjek (Biak dialect). Distribution and ecology: One species from the
The stems are used in building construction. Raja Ampat Islands (known only from Gag and Wai-
Notes: Manjekia is a moderately robust, solitary geo). Lowland rainforest on limestone up to 50 m.
palm that is recorded from several lowland rainforest Anatomy: Not studied.
localities on limestone in Biak Island. Its arching Relationships: Wallaceodoxa belongs to Ptychosper-
leaves bear broadly lanceolate, pendulous, praemorse matinae (Arecoideae: Areceae). It is moderately sup-
leaflets with concave, praemorse apices and its inflo- ported as sister to Adonidia (Alapetite et al., 2014).
rescence axes are white or greenish white. Manjekia Common names and uses: Gulbotom (Wayaf or
bears some similarities to Adonidia, in which it was Gebe dialect). The stem is used in building con-
originally described, such as the white inflorescences struction and the seed is consumed as a betel nut
branched up to four orders, the red fruit and similar substitute.
seed and endocarp morphology, but its leaves do not Notes: Wallaceodoxa is a rare palm of the Raja
resemble those of Adonidia, which have ascending, Ampat Islands (Heatubun, Lekitoo & Matani,
narrow leaflets in slightly different planes that are 2014a). Fewer than 40 adults have been recorded
less conspicuously praemorse and fewer stamens in from the islands of Gag and Waigeo, to which it
the staminate flowers (30–32, instead of 45–50 in appears to be restricted. It is a moderately robust,
Adonidia). See also notes under Jailoloa. solitary palm bearing arching leaves with narrow,
pendulous leaflets that are obliquely praemorse at appears to be supported by anatomical evidence
their apices. A pronounced layer of thick, white and (Seubert, 1996; Fisher, Tan & Toh, 2002; Tomlinson
brown–black indumentum covers the leaf sheath, & Spangler, 2002; Tomlinson et al., 2011).
petiole and rachis. The inflorescences are greenish
white with thick rachillae and relatively crowded flo-
ral triads. The staminate flower contains a short, PRITCHARDIOPSIS (CORYPHOIDEAE: TRACHYCARPEAE:
ellipsoid pistillode, rather than the elongate, bottle- LIVISTONINAE).
shaped pistillode that is typical of many Ptychosper- Saribus includes the sole species of the monotypic
matinae. With the exception of seed and endocarp Pritchardiopsis (Bacon & Baker, 2011; Bacon et al.,
characters, Wallaceodoxa does not obviously resem- 2012a). As Saribus is an earlier generic name,
ble its sister genus Adonidia, or indeed any other Pritchardiopsis has therefore been placed in syn-
genus in the subtribe. See also notes under Jailoloa. onymy. For further discussion, see the treatment of
Saribus above.
GENERA LOST SINCE GP2 WALLICHIA (CORYPHOIDEAE: CARYOTEAE).

CERATOLOBUS, DAEMONOROPS, POGONOTIUM AND Jeanson (2011) completed a monographic study of
RETISPATHA (CALAMOIDEAE: CALAMEAE: CALAMINAE). tribe Caryoteae that points to the sinking of Wal-
The non-monophyly of the rattan genus Calamus L. lichia Roxb., as a result of it being embedded in a
has been revealed in multiple phylogenetic studies paraphyletic Arenga Labill., although formal syn-
(Baker et al., 1999b; Baker, Dransfield & Hedderson, onymy has not yet been published. Dransfield et al.
2000a; Baker, Hedderson & Dransfield, 2000b, c; (2008a, b) acknowledged that the differences between
Baker et al., 2009; W. J. Baker, unpubl. data), all of the two genera are small, the only unambiguous
which indicate that the remaining genera of Calami- character being the sepals of the staminate flower
nae are variously nested in Calamus. In GP2, being connate in a tube, rather than free and imbri-
Dransfield et al. (2008b) were reluctant to address cate (as in Arenga). Therefore, this taxonomic change
this problem taxonomically, following earlier authors is unsurprising.
(Baker et al., 2000b) who preferred to wait for further
data before proposing an alternative classification.
Notwithstanding this, a process of reduction in genera LYTOCARYUM (ARECOIDEAE: COCOSEAE: ATTALEINAE).
of Calaminae had already started, first with the sink- In GP2, the small genus Lytocaryum Toledo is distin-
ing of Calospatha Becc. into Calamus in preparation guished from other species of Attaleinae by its fruit
for GP2 (Baker & Dransfield, 2008), more recently fol- with its epicarp and mesocarp splitting at maturity
lowed by Retispatha J.Dransf. (Henderson & Floda, and its thin endocarp, by its narrow, closely spaced
2015). Baker (2015) completed this process by sinking leaflets with pale indumentum abaxially and its ver-
the remaining three genera of Calaminae (Ceratolobus satile anthers. In a recent paper (Noblick & Meerow,
Blume, Daemonorops Blume ex Schult.f. and Pogono- 2015), a strong case is made for reducing Lytocaryum
tium J.Dransf.) into Calamus, thereby expanding the into Syagrus Mart., as proposed by earlier workers
largest genus of palms from > 400 to c. 520 species. (e.g. Glassman, 1965). There is much phylogenetic
The re-delimited, broad concept of Calamus has evidence for the close relationship between Lyto-
several advantages. First, it is a stable, pragmatic caryum and Syagrus (Hahn, 2002b; Gunn, 2004;
solution to the Calamus problem that is likely to be Baker et al., 2009), with the most recent studies
robust to any future phylogenetic findings. Second, it (Meerow et al., 2009, 2015) suggesting that Lyto-
ensures that the largest genus of palms is in fact caryum may in fact be nested in Syagrus. Noblick &
monophyletic and phylogenetically sound. Third, it is Meerow (2015) also demonstrated that all distin-
more easily identified, compared with the ambigu- guishing characters of Lytocaryum are found, at least
ously defined and polymorphic Calamus s.s. and the to some extent, in Syagrus.
multiple genera nested in it. Calamus is now defined
as a genus of dioecious, pleonanthic rattan species,
usually with a swollen knee-like structure on the leaf SOLFIA (ARECOIDEAE: ARECEAE: PTYCHOSPERMATINAE).
sheath at the petiole base, with inflorescences adnate The Samoan, monotypic Solfia Rechinger was resur-
to the internode and leaf sheath above the axil of ori- rected from synonymy in Drymophloeus Zipp. on
gin, and with floral clusters almost always compris- morphological grounds (Zona, 1999) and included as
ing a functional pistillate flower and a sterile an accepted genus in GP2. Multiple molecular phylo-
staminate flower in the female plant and a solitary genetic studies have indicated that it is closely
staminate flower in the male plant. The genus also related to Balaka from Fiji and Samoa (Norup et al.,
2006; Baker et al., 2009, 2011; Zona et al., 2011). nized at subgeneric level in Astrocaryum than as a
However, a recent study of Ptychospermatinae with genus.
near-complete taxon sampling (Alapetite et al., 2014)
placed Solfia in Balaka. Consequently, Solfia was
placed in synonymy (Zona & Baker, 2014), resulting
SPECIES-LEVEL TAXONOMY
in a monophyletic if somewhat heterogeneous Balaka
(e.g. in relation to endocarp morphology), which is Although the focus of this paper is palm systematics
consistent with the rather challenging nature of gen- at the genus level and above, a note on species-level
eric delimitation across the subtribe. taxonomy is merited. Since the publication of GP2,
> 200 new palm species have been described (IPNI,
2015), with > 320 names published in total, if new
THE CASE OF HEXOPETION (ARECOIDEAE: COCOSEAE: combinations and replacement names are also con-
BACTRIDINAE). sidered. Genera that have been revised entirely (or
Hexopetion Burret, originally described by Burret at regional or infrageneric level) since GP2 include
(1934), was resurrected and expanded by Pintaud, Basselinia Vieill. (in part; Pintaud & Stauffer, 2011),
Millan & Kahn (2008) as a segregate of two species Ceroxylon Bonpl. (Sanın & Galeano, 2011), Chunio-
from Astrocaryum G.Mey. The characters presented phoenix Burret (Henderson, 2015), Cyrtostachys
to distinguish Hexopetion were multifold lateral leaf- Blume (Heatubun et al., 2009), Desmoncus Mart.
lets in adults, the lack of a sterile portion of the (Henderson, 2011a), East Malesian Areca L.
rachilla between the basal pistillate flower and distal (Heatubun et al., 2012), Eremospatha (G.Mann &
staminate flowers, white, woolly indumentum on the H.Wendl.) Schaedtler, Laccosperma (G.Mann &
rachilla between the flowers, stigmas much shorter H.Wendl.) Drude and Oncocalamus Mann & H.
than the ovary and leaf vascular anatomical fea- Wendl. (Sunderland, 2012), Geonoma Willd. (Hender-
tures. Phylogenetic evidence to support the segrega- son, 2011b), Leopoldinia Mart. (Henderson, 2011c),
tion of Hexopetion was not presented at that time. Bornean Licuala (Saw, 2012), Livistona (including
Dransfield et al. (2008b) considered that insufficient most of Saribus; Dowe, 2009), Lytocaryum (now
justification had been provided for Hexopetion to be included in Syagrus; Noblick & Lorenzi, 2010), Ora-
accepted in GP2. nia Zipp. (Keim & Dransfield, 2012), Pholidostachys
Several molecular phylogenetic studies have now H.Wendl. ex Benth. & Hook.f. (Henderson, 2012),
clarified that the two Astrocaryum spp. included in Trithrinax Mart. (Cano, Perret & Stauffer, 2013) and
Hexopetion by Pintaud et al. (A. alatum H.F.Loomis Welfia H.Wendl. (Henderson & Villalba, 2013). In
and A. mexicanum Liebm. ex Mart.) form a mono- addition, many new species have been published in
phyletic group that is sister to a clade comprising all non-monographic studies, e.g. in Areca (e.g. Hea-
remaining Astrocaryum spp. (Eiserhardt et al., tubun, 2011b; Heatubun, Iwanggin & Simbiak,
2011a; Ludena et al., 2011; Roncal et al., 2013, 2015; 2013), Calamus (e.g. Henderson & Nguyen Quoc,
Meerow et al., 2015). Thus, the argument for accept- 2013; Baker & Dransfield, 2014), Dypsis Noroha ex
ing Hexopetion or not is equivocal from a phyloge- Thou. (e.g. Rakotoarinivo, Trudgen & Baker, 2009;
netic standpoint because Astrocaryum is Rakotoarinivo & Dransfield, 2010) and Syagrus (e.g.
monophyletic whether or not the Hexopetion spp. are Noblick, Lorenzi & Souza, 2014). These publications
included. Eiserhardt et al. (2011a) emphasized this indicate that there is still much alpha taxonomic
point, but drew attention to the split between Hex- research and species discovery to be accomplished in
opetion spp. and the remainder of Astrocaryum, the palm family.
which is deeper than any other in Astrocaryum The biodiversity informatics portal Palmweb
(although it is younger than any other genus-level (Palmweb, 2015) has grown to become an important
split in the subtribe), and the biogeographical dis- internet resource for palm systematics, providing
junction between Central American Hexopetion and complete taxonomy and distribution information for
the mostly Amazonian remainder of Astrocaryum. all genera and species, based on the World Checklist
Primarily, however, the case for recognizing Hex- of Palms (Govaerts et al., 2015) and additional rich
opetion rests on a subjective judgement on the suffi- content (e.g. descriptions, images) for c. 1500 species
ciency of its morphological distinctness. (Baker et al., 2015). Palmweb data have also been
Some authors have taken up Hexopetion (e.g. Ron- provided to larger biodiversity informatics portals,
cal et al., 2013, 2015), whereas others prefer the such as eMonocot (Gardiner, Bone & Kilgallen,
broad sense Astrocaryum (e.g. Eiserhardt et al., 2013). Content from Palmweb has also been re-
2011a; Meerow et al., 2015). We concur with Meerow purposed in an application for mobile devices,
et al. (2015) that Hexopetion would be better recog- Palmworld, which brings technical taxonomic con-
tent to a broader audience through a more accessible is often called, has been a springboard for many
interface (Palmworld, 2015). important comparative studies addressing biogeogra-
phy and diversification (Baker & Couvreur, 2012,
2013a, b; Kissling et al., 2012; Eiserhardt et al.,
2013a), biome origins (Couvreur, Forest & Baker,
PALM PHYLOGENETICS
2011; Couvreur & Baker, 2013), community assembly
Palm phylogenetic research from its inception to (Eiserhardt et al., 2013b) and the evolution of floral
2008 was extensively reviewed in GP2, which also (Sannier et al., 2009; Nadot et al., 2011; Rudall,
summarized phylogenetic evidence for each taxon Ryder & Baker, 2011), vegetative (Couvreur et al.,
from genus to subfamily. Forty-six phylogenetic 2015; Thomas & Boura, 2015) and chemical (Siles,
papers were reported in GP2, more than one-half of Cela & Munn e-Bosch, 2013) traits.
which included morphological data (either exclu- Despite the wide uptake of the Baker et al. (2009)
sively or in combination with DNA data). A much ‘palm supertree’, its potential for use in macroevolu-
smaller subset of these papers, based primarily on tionary research is limited by its lack of species-level
DNA, and with broad taxon sampling at the higher detail. Some studies have identified expedient
level, were of particular importance to the formula- workarounds to overcome this problem (Couvreur
tion of the new classification (Baker et al., 2000a, c; et al., 2011, 2015; Kissling et al., 2012; Eiserhardt
Lewis & Doyle, 2001, 2002; Hahn, 2002a, b; Asmus- et al., 2013b), but, recently, a more comprehensive
sen et al., 2006; Norup et al., 2006; Savolainen et al., solution has been sought in the construction of a
2006; Trenel et al., 2007; and some unpublished complete species-level phylogenetic analysis of palms
research cited in GP2 as Baker et al., in review and (Faurby et al., in press). Faurby et al. assembled all
in prep., now published as Baker et al., 2009; Baker publicly available DNA data for palms, covering all
et al., 2011, respectively). genera and 901 species, adding morphological data
Here, we provide a synthesis of palm phylogenetic that increased the number of species for which at
developments since GP2, focusing primarily on the least one data type was available to 1255 species,
phylogenetics of higher level groups. The most representing c. 50% of the family. The complete spe-
strongly supported phylogenetic relationships among cies-level phylogenetic tree was constructed using a
the subfamilies and tribes of palms are summarized novel Bayesian approach with the placement of the
in Figure 7. remaining 50% of unsampled species informed by
taxonomic information.
The value of the phylogenetic analysis of Faurby
PALM FAMILY-WIDE PHYLOGENETICS
et al. lies primarily in its potential as a tool for evo-
The most prominent contribution to family-wide phy- lutionary, ecological and biogeographical research
logenetics since GP2 was published by Baker et al. that requires a complete tree, rather than as a
(2009), who generated the first complete genus-level truly accurate reconstruction of palm species-level
phylogenetic analysis of palms. This study brought relationships for systematic purposes. However, in
together all available major phylogenetic datasets constructing the backbone of their phylogenetic tree,
into a supermatrix comprising 14 DNA sequence Faurby et al. revisited the supermatrix of Baker
data partitions, a DNA restriction fragment length et al. (2009), reanalysing it with Bayesian methods,
polymorphism dataset and a morphological dataset in contrast with the parsimony-based ‘total evi-
covering all palm genera. These data were analysed dence’ and supertree analyses that Baker et al. had
with a variety of supermatrix and supertree meth- used. A surprising number of conflicting relation-
ods, resulting in a range of alternative topologies ships between the two studies were revealed, the
that arbitrated objectively between hypotheses more highly supported disagreements being concen-
obtained from subsets of the 16 partitions. The taxo- trated in Calamoideae, Coryphoideae, Ceroxyleae,
nomic congruence between the alternative topologies Cocoseae and Areceae. This serves as an important
was formally evaluated to achieve a ‘best estimate’ of reminder that different analytical approaches
the most consistently recovered relationships. Impor- should, where possible, be utilized in parallel to
tantly, the resulting trees were much more highly gain a rounded picture of the phylogenetic signal
resolved than other comprehensive palm phylo- within datasets. It also further highlights areas of
genetic trees available at the time (Asmussen et al., poor data sampling in the Baker et al. supermatrix,
2006). some of which have been addressed in subsequent
The results of Baker et al. (2009) were used heav- studies. Fortunately, the tree of Faurby et al. did
ily as evidence to underpin the GP2 classification, not call into question the GP2 classification, beyond
even though they were published after GP2. Since those issues that were already discussed by Baker
that time, the ‘palm supertree’ of Baker et al., as it et al. (2009).
Eugeissoneae
Lepidocaryeae CALAMOIDEAE
Calameae
NYPOIDEAE
Cryosophileae
Sabaleae
1
Phoeniceae
Trachycarpeae
CORYPHOIDEAE
Chuniophoeniceae
Caryoteae
2
Corypheae
Borasseae
Cyclospatheae
Ceroxyleae CEROXYLOIDEAE
Phytelepheae
Chamaedoreeae
Iriarteeae
Podococceae
6 Oranieae
Sclerospermeae
Roystoneeae
7 Reinhardtieae
ARECOIDEAE
Cocoseae
3
Leopoldinieae
4 Geonomateae
Manicarieae
5
Pelagodoxeae
Euterpeae
Areceae
Figure 7. Schematic tree synthesizing the phylogenetic relationships among palms (redrawn and updated from Drans-
field et al., 2008b; fig. 6.1, p. 98). All nodes are strongly supported in at least one of the most comprehensive phyloge-
netic studies of palms (strong support = ≥ 90% bootstrap or jackknife support for maximum parsimony or likelihood
analyses, ≥ 0.9 posterior support for Bayesian analyses, or supported by ≥ 5 input trees for supertree analyses). No
strongly supported conflicting relationships were identified. Relationships among subfamilies are well established
(Asmussen et al., 2006; Baker et al., 2009, 2011; Barrett et al., 2016; Faurby et al., in press). Evidence for numbered
nodes is as follows. Node 1. CSPT clade – Barrett et al. (2016), Faurby et al. (in press); lower support: Baker et al.
(2009), Bacon et al. (2012a). Node 2. Syncarpous clade – Asmussen et al. (2006), Baker et al. (2009), Bacon et al.
(2012a), Barrett et al. (2016), Faurby et al. (in press). Node 3. Comer et al. (2015, 2016), Faurby et al. (in press); lower
support: Baker et al. (2009). Node 4. Comer et al. (2015, 2016). Node 5. Core arecoid clade – Baker et al. (2009, 2011),
Comer et al. (2015, 2016), Faurby et al. (in press). Node 6. POS clade – Baker et al. (2009, 2011), Comer et al. (2015,
2016), Faurby et al. (in press). Node 7. RRC clade – Baker et al. (2011), Comer et al. (2015, 2016), Faurby et al. (in
press); lower support: Baker et al. (2009). For evidence for remaining nodes, see text.
The impact of the next-generation sequencing revo- among the genera, placing Laccosperma as sister to
lution (Pyron, 2015) is now being felt in palm phyloge- Eremospatha.
netics (Comer et al., 2015, 2016; Barrett et al., 2016,
in press; Heyduk et al., 2016). Two of these studies
(Comer et al., 2015; Barrett et al., 2016), focusing on PHYLOGENETICS OF CORYPHOIDEAE
the whole family and Arecoideae, respectively, have Significant clarification of the relationships among
already included complete or near-complete plastid the tribes of Coryphoideae has been achieved with
genome data for 62 genera of palms. These studies the publication of highly congruent phylogenetic
provide strong support for the GP2 classification and topologies of the subfamily since GP2 (Baker et al.,
relationships among the five subfamilies (Fig. 7) that 2009; Bacon et al., 2012a; Barrett et al., 2016;
have been established in earlier studies (e.g. Asmus- Faurby et al., in press). We can now be confident
sen et al., 2006; Baker et al., 2009, 2011). In addition, that the subfamily is divided into two major clades
Barrett et al. (2016) illustrated, for the first time with (Fig. 7): the CSPT and the syncarpous clades. The
genomic data, the exceptionally slow rate of plastid CSPT clade, coined here, is divided into two sub-
DNA evolutionary change that has dogged all plastid clades, one comprising the New World thatch palm
phylogenetic studies of palms (Baker et al., 1999a; clade (sister tribes Cryosophileae and Sabaleae;
Asmussen et al., 2000, 2006; Asmussen & Chase, recovered by the above authors, except Faurby et
2001; Scarcelli et al., 2011). al., in press) and the other containing Phoeniceae
as sister to Trachycarpeae. Evidence for the sub-
clades of the CSPT clade has been found previously
PHYLOGENETICS OF CALAMOIDEAE (e.g. Uhl et al., 1995; Asmussen et al., 2006), but
Little additional research on calamoid phylogenetics robust support for their sister relationship has
has been published since GP2, although several stud- appeared only in more recent studies. It should be
ies are currently in progress. The summary relation- noted that, although the supertree analysis of
ships among tribes presented in GP2 as Baker et al. (2009) resolved the relationships
(Eugeissoneae (Lepidocaryeae, Calameae)) have, described above, their supermatrix analysis placed
however, been challenged. Baker et al. (2009) found Phoeniceae sister to all remaining CSPT members,
support for this pattern from supertree analyses, but but without bootstrap support. The syncarpous
their supermatrix analysis resolved a paraphyletic clade contains four tribes, which resolve as (Chu-
Lepidocaryeae with Eugeissoneae embedded in it as niophoeniceae (Caryoteae (Corypheae, Borasseae))),
sister to Raphia P.Beauv. However, these relation- as indicated in earlier studies (Bayton, 2005;
ships are poorly supported. Others have also found Asmussen et al., 2006). The supermatrix analysis of
different relationships (Barrett et al., 2016; Faurby Baker et al. switches the placement of tribes Cary-
et al., in press), including a paraphyletic Lepido- oteae and Corypheae, but without support.
caryeae and a sister relationship between Eugeis- In the GP2 classification, Dransfield et al. (2008a,
soneae and Calameae. These findings reflect b) were unable to erect a complete classification in
ambiguity regarding the placement of Eugeissoneae tribe Trachycarpeae as a result of inadequate phylo-
and the monophyly of Lepidocaryeae that were genetic resolution, and therefore placed only some of
already prevalent in the earliest studies of the sub- the 17 genera in the two subtribes Rhapidinae and
family (Baker et al., 2000a, c; Fig. 7). These uncer- Livistoninae, leaving seven genera [six from the
tainties may be a result of the intrinsic properties of Americas, one (Pritchardia Seem & H.Wendl.) from
calamoids, such as their great age (c. 80 Mya) and the Pacific] unplaced to subtribe. We now have a
the relatively rapid succession of divergences at their much clearer understanding of the relationships
base (Baker & Couvreur, 2013a), or long branch arte- among the genera of Trachycarpeae (Henderson &
facts in species-poor lineages, such as Eugeissoneae. Bacon, 2011; Bacon et al., 2012a, b, 2013a, b; Barrett
However, the available evidence for Calamoideae is et al., 2016) that supports the monophyly of the two
based on rather limited data sampling and, where subtribes, placing Brahea Mart. sister to Rhapidinae,
deep character sampling has been achieved (Barrett and a clade of Acoelorrhaphe H.Wendl. and Serenoa
et al., 2016), taxon sampling was limited. On bal- Hook.f. as the sister group of Livistoninae. The best
ance, the deep relationships in Calamoideae remain evidence (Bacon et al., 2012a) suggests that Coperni-
an open question. cia Mart. ex Endl., Pritchardia and Washingtonia
At a lower taxonomic level, two phylogenetic stud- H.Wendl. fall outside this group as a monophyletic
ies of Ancistrophyllinae (Lepidocaryeae) have con- or paraphyletic group, with the placement of
firmed the monophyly of the three genera accepted Colpothrinax Schaedtler remaining ambiguous. The
in the subtribe (Faye et al., 2014; Faye, 2015). The relationships of the unplaced genera of Trachy-
latter study strongly resolved the relationships carpeae do not lend themselves readily to classifica-
tion without recognizing several small subtribes that Faurby et al., in press) has provided strong support
cannot be easily distinguished morphologically. For for several major clades in the subfamily (Fig. 7): (1)
the time being, the unplaced Trachycarpeae remains the POS clade (Podococceae, Oranieae, Sclerosper-
a useful and practical concept. meae); (2) the RRC clade (Roystoneeae, Rein-
The comprehensive tribe-wide research on Trachy- hardtieae, Cocoseae); and (3) the core arecoid clade
carpeae (Bacon et al., 2012a) has been complemented (Areceae, Euterpeae, Geonomateae, Leopoldinieae,
by a series of focused studies with dense species sam- Manicarieae, Pelagodoxeae). These studies do not
pling aimed at taxon delimitation (Bacon & Baker, provide consistent accounts of the relationship
2011; Henderson & Bacon, 2011; Bacon et al., among and within these groups. However, by weigh-
2012b), and the biogeography of Wallace’s Line ing up the strongly supported relationships against
(Bacon et al., 2013a), Australia (Crisp et al., 2010) the weaker hypotheses, especially from recent plastid
and the Isthmus of Panama (Bacon et al., 2013b). and nuclear phylogenomic research, a clearer picture
Consequently, Trachycarpeae now ranks as one of emerges (Fig. 7). Most studies identify Iriarteeae and
the best known groups of palms from a systematic Chamaedoreeae as the earliest branching lineages in
and macroevolutionary perspective. Beyond Trachy- Arecoideae, although all possible alternative arrange-
carpeae, Pintaud et al. (2010) used simple sequence ments are recovered, i.e. the two tribes as sisters
repeats to explore species limits in Phoenix L., which (Comer et al., 2016) or the two as a paraphyletic
confirmed the prevailing taxonomy, without inferring group with Iriarteeae (Asmussen et al., 2006; Baker
phylogenetic relationships because of the limitations et al., 2009; Comer et al., 2016; Faurby et al., in
of the markers used. Heyduk et al. (2016) have press) or Chamaedoreeae (Comer et al., 2015) as sis-
recently published a comprehensive phylogenetic ter to all remaining arecoids. The POS clade is most
analysis of Sabal Adans. in a paper that established strongly supported as sister to an RRC + core arecoid
invaluable new tools for next-generation sequencing clade (Comer et al., 2015, 2016), although moderate
in palms (see below). support has also been recovered for a topology in
which the positions of the RRC and POS clades are
switched (Baker et al., 2009; Faurby et al., in press).
PHYLOGENETICS OF CEROXYLOIDEAE In the POS clade, strong evidence points to Oranieae
When preparing the GP2 classification, all evidence being sister to Sclerospermeae (Baker et al., 2009,
converged on the same relationships among the 2011; Comer et al., 2015, 2016). In the RRC clade,
three tribes of Ceroxyloideae, placing Cyclospatheae Reinhardtieae is most closely related to Cocoseae
sister to a clade of Ceroxyleae and Phytelepheae (Baker et al., 2009, 2011; Comer et al., 2015, 2016;
(Hahn, 2002b; Asmussen et al., 2006; Trenel et al., Faurby et al., in press).
2007; Fig. 7). Subsequent studies confirmed these The relationships among the core arecoids remain
results (Baker et al., 2009, 2011; Faurby et al., in elusive, but recent phylogenomic data point to at
press). Recently, however, plastid phylogenomic anal- least one strongly supported node, placing Areceae
yses have proposed an alternative topology, placing as sister to Euterpeae (Comer et al., 2015, 2016).
Cyclospatheae sister to Phytelepheae (Barrett et al., Other proposed relationships, such as Leopoldinieae
2016). Although substantial data underpin this con- sister to all other core arecoid tribes (Comer et al.,
trasting topology, we note that this node is one of 2016) or a corky-warted fruit clade (Mani-
only a few with a bootstrap percentage < 100% carieae + Pelagodoxeae; Comer et al., 2016), remain
(although the values are still high, mostly > 95%), doubtful because of low support values and conflict.
indicating lower confidence than most other relation- The higher level relationships of Arecoideae,
ships recovered in the study. The low taxon sampling although better known now than previously, remain
(one species per tribe) is also potentially problematic a potentially fruitful focus of future research.
because long branch artefacts may be an issue in The phylogenetics of several arecoid tribes have
this group (Asmussen et al., 2006). Further taxon been studied in depth since GP2. There is much
sampling would mitigate this risk. In addition to conflict between phylogenetic hypotheses for inter-
higher phylogenetic research on Ceroxyloideae, spe- generic relationships in Chamaedoreeae (Asmussen
cies-level phylogenetic analyses have been conducted et al., 2006; Thomas et al., 2006; Cuenca & Asmus-
for Phytelepheae (Barfod, Trenel & Borchsenius, sen-Lange, 2007; Cuenca, Asmussen-Lange &
2010) and Ceroxylon (M. Sanın, unpubl. data). Borchsenius, 2008; Baker et al., 2009, 2011;
Cuenca, Dransfield & Asmussen-Lange, 2009). How-
ever, the studies with the densest data and taxon
PHYLOGENETICS OF ARECOIDEAE sampling offer moderate to strong support for a
Phylogenetic research since GP2 on Arecoideae clade comprising Chamaedorea Willd., Gaussia
(Baker et al., 2009, 2011; Comer et al., 2015, 2016; H.Wendl. and Synechanthus H.Wendl., which is sis-
ter to Wendlandiella Dammer, with Hyophorbe Laccospadicinae, Rhopalostylidinae; Baker et al.,

Gaertn. sister to all remaining Chamaedoreeae 2009, 2011). The relationships of the ten genera that
(Cuenca et al., 2008, 2009; Baker et al., 2011). A were not placed to subtribe in the GP2 classification
recent study of tribe Iriarteeae confirms the mono- of Areceae remain obscure. Two subtribes have
phyly of all five accepted genera, and finds strong become better known through focused studies. An
support for a clade of Iriartea Ruiz & Pav. and Dic- extensive plastid DNA study has illuminated the
tyocaryum H.Wendl. resolving as sister to a second relationships among the species and genera of
clade comprising the remaining three genera (Bacon Archontophoenicinae, although questioning the
et al., in press). placement of Actinorhytis H.Wendl. & Drude in this
Cocoseae has been intensively studied since the group (Domenech et al., 2014). Two papers on Pty-
publication of GP2, with papers focusing on higher chospermatinae have substantially improved the
relationships in the tribe (Meerow et al., 2009, 2015; understanding of the generic limits and relationships
Eiserhardt et al., 2011a), providing further support of this difficult subtribe, resulting in substantial gen-
to the well-established sister group relationship eric change (Zona et al., 2011; Alapetite et al., 2014),
between Bactridinae and Elaeidinae that has been including the movement of species of Drymophloeus
reported frequently elsewhere (Hahn, 2002b; Asmus- to expanded Veitchia and Ponapea Becc., the sinking
sen et al., 2006; Baker et al., 2009, 2011; Comer of Solfia into Balaka and the establishment of the
et al., 2015, 2016). Highly incongruent relationships new genera Jailoloa, Manjekia and Wallaceodoxa
have been found among Bactridinae in the studies (Heatubun et al., 2014b; Zona & Baker, 2014).
that have sampled all genera (Baker et al., 2009; Eis-
erhardt et al., 2011a; Meerow et al., 2015), although
support is highest and most widespread in the study
PROSPECTS
of Eiserhardt et al., which is based on five plastid
DNA regions and three nuclear regions. Several Global knowledge of higher level palm systematics is
papers have focused on the phylogenetics of the com- in excellent shape. GP2 provides a robust baseline,
plex genus Astrocaryum and its implication for reflecting the achievements of the strong, collabora-
Neotropical biogeography (Ludena et al., 2011; Ron- tive research community working on palm biology,
cal et al., 2013, 2015). Considerable ambiguity also which is fuelled by regular interactions through the
remains in the intergeneric relationships of Attalei- European Network of Palm Scientists, 5-yearly
nae, although the best sampled studies all point to World Palm Symposia and other co-operations, such
the American taxa forming a clade and the African as the European Union-funded Palms project (Bal-
(Jubaeopsis Becc.) and Madagascan (Beccariophoenix slev, Macia & Navarrete, 2015). Like GP1, GP2 has
Jum. & H.Perrier, Voanioala) coming together in a been a springboard for the many new developments
sister clade or paraphyletic group (Baker et al., 2009; in palm taxonomy and phylogenetics that have been
Meerow et al., 2009, 2015; Eiserhardt et al., 2011a). synthesized here. However, numerous aspects of
Attalea Kunth has recently been the focus of species- palm systematics are still unknown and palm
level phylogenetic research (C. Freitas, unpubl. researchers must grasp new opportunities to address
research). these, so that the full potential of the palms as a
Multiple studies converge on a common set of rela- model group for tropical plant research can be
tionships among genera of Geonomateae (sister rela- achieved (Couvreur & Baker, 2013). We conclude
tionship between Welfia and Pholidostachys, here with some perspectives on how the next steps
Calyptrogyne H.Wendl. embedded in a paraphyletic in palm systematics may be taken.
Calyptronoma Griseb.), albeit with patchy support
and questions over the placement of Asterogyne
H.Wendl. ex Hook.f. (Roncal et al., 2005, 2010, 2011; IN PALM TAXONOMY, EXPECT THE UNEXPECTED
Baker et al., 2009, 2011). The addition of new Palm species continue to be discovered at a startling
nuclear data, however, places Pholidostachys as sis- rate. Even well-studied areas become hotspots for
ter to all remaining Geonomateae, followed by Wel- new discoveries because enhanced knowledge of a
fia, and Asterogyne as sister to the Calyptrogyne/ region facilitates the identification of further novel-
Calyptronoma clade (Roncal et al., 2012). ties. For example, the completion of baseline tools for
The relationships in Areceae, the largest tribe of the palms of Madagascar (Dransfield & Beentje,
all palms, remain poorly understood, to the extent 1995a; Dransfield et al., 2006) has unlocked our abil-
that relations among subtribes are scarcely estab- ity to detect yet more new taxa. The combination of
lished (with the exception of a weakly supported field and phylogenetic exploration has also been pro-
western Pacific clade) and the monophyly of some ductive at the genus level. The description of six new
subtribes remains questionable (e.g. Basseliniinae, genera so soon after GP2 was highly unexpected and
the location of further new genera cannot be ruled tools to the full, but suggests that optimism should
out. In summary, it is far too early to consider palm be tempered with realism.
taxonomy as adequately understood. We must take
all opportunities to explore palm taxonomic frontiers
A SPECIES-LEVEL PHYLOGENY OF PALMS IS WITHIN REACH
in the field and herbarium and embrace the insights
from the molecular characterization of species Species-level phylogenetic trees are in great demand
(Buerki & Baker, 2016). Without knowing our spe- for evolutionary and ecological research, and palms
cies, we are poorly equipped to study them, and even are no exception. In palms, researchers needing a
less able to protect them from extinction. complete species tree have resorted to simulating the
branches into the genus-level supertree of Baker
et al. (2009) or assembling larger trees from publi-
THE PALM TREE OF LIFE IS NOT COMPLETELY KNOWN cally available data (Couvreur et al., 2011; Kissling
No larger family of tropical plants is probably better et al., 2012; Eiserhardt et al., 2013b; Couvreur et al.,
known phylogenetically than the palms. The palm 2015; Faurby et al., in press). However, we are now
community has established a complete genus-level poised to build a species-level phylogenetic tree, not
framework (Baker et al., 2009) and a synthetic spe- based on interpolating branches, but on real data.
cies-level framework (Faurby et al., in press), each The scientific need for complete species-level phylo-
building on numerous in-depth case studies, and is genetic research is evident, to produce fundamental
now moving into the genomic era (Barrett et al., in biodiversity knowledge for pure and applied compar-
press). However, although we have reason to be con- ative research and to reinforce palms as a model
fident in many critical relationships, there are group for tropical research. The data underpinning
numerous areas of significant ambiguity in some of such an endeavour would also yield an authoritative
the most diverse and important groups of palms, genomic resource for palms, providing molecular
such as in Arecoideae and Calamoideae, and species- tools in support of identification and other applied
level phylogenetics are incompletely known. Under- uses. New phylogenomic tools now make a species-
standing the relationships of such ecologically and level phylogenetic tree for palms a tractable objec-
economically important plants is of fundamental sci- tive, but it can only be achieved within a collabora-
entific importance and must remain a high priority. tive framework, in which researchers share material
from the field, cultivation, DNA banks and herbaria,
working together on complementary protocols and
PHYLOGENOMICS WILL REVOLUTIONIZE PALM RESEARCH sharing data freely and fairly. These are the princi-
Next-generation sequencing methods are now ples of the Palm Phylogeny Working Group, estab-
becoming routinely used in phylogenetics. The lished at the World Palm Symposium in Montenegro,
impact of these methods is now being felt in palms Colombia in July 2015. The road ahead for palm sys-
as plastid and nuclear datasets of unprecedented tematic research has never been more exciting.
scale are being brought to bear on palm phyloge-
netics from the species to the family level (Barrett
et al., in press). The sequence capture methods
ACKNOWLEDGEMENTS
recently published by Heyduk et al. (2016) are par-
ticularly promising as they are ambitious and infor- We offer heartfelt thanks to the organizing commit-
mative, yet tractable. The potential of this scale of tee of the 2015 World Palm Symposium in Colombia,
DNA sequence data to inform or even overturn especially our generous local hosts, Rodrigo Bernal
palm systematics is immense. Next-generation and Gloria Galeano, for inviting our participation
sequencing methods also have the potential to and the Instituto de Investigaci on de Recursos
unlock herbaria as a source of samples for sequenc- Biol
ogicos Alexander von Humboldt and the Ben-
ing, because they are much more tolerant of tham Moxon Trust for funding our attendance. We
degraded DNA than traditional Sanger sequencing take this opportunity to recognize our extraordinary
methods (Buerki & Baker, 2016). A caveat should GP2 collaborators, Conny Asmussen-Lange, Made-
be added here that these approaches may not be a line Harley, Carl Lewis and Natalie Uhl, and the
panacea and some relationships may ultimately many palm researchers, institutions and gardens
prove to be insoluble for real biological reasons. We who have supported our work. We are grateful to
have already seen, for example, that rates of discor- Christine Bacon, Craig Barrett, Angela Cano, Jason
dance among gene trees can be extremely high, Comer, Søren Faurby, Adama Faye and Marc Jean-
undermining confidence in resolved relationships son for allowing us access to unpublished results in
(Heyduk et al., 2016; Comer et al., 2016). This the preparation of the manuscript. Wolf Eiserhardt,
should not be an obstacle to exploiting the new Christine Bacon and an additional anonymous
reviewer provided very helpful comments on the Bacon CD, Michonneau F, Henderson AJ, McKenna
manuscript. Finally, we thank Mike Fay for making MJ, Milroy AM, Simmons MP. 2013a. Geographic and
meticulous updates to the manuscript that were well taxonomic disparities in species diversity: dispersal and
beyond the call-of-duty. diversification rates across Wallace’s Line. Evolution 67:
2058–2071.
Bacon CD, Mora A, Wagner WL, Jaramillo CA. 2013b.
Testing geological models of evolution of the Isthmus of
DEDICATION Panama in a phylogenetic framework. Botanical Journal of
the Linnean Society 171: 287–300.
This paper is dedicated to French palm expert Jean-
Bacon CD, Vel asquez-Puentes F, Fl orez-Rodrıguez A,
Christophe Pintaud. News of Jean-Christophe’s
Balslev H, Galeano G, Bernal R, Antonelli A. in press.
unexpected death just days after the 2015 World
Phylogeny of Iriarteeae palms, cross-Andean disjunctions,
Palm Symposium that he co-organized brought not
and convergence of clustered infructescence morphology in
only immense sadness, but also many reflections on Wettinia. Botanical Journal of the Linnean Society.
wonderful times spent studying, debating and travel- doi:10.1111/boj.12399
ling for palms with him. Jean-Christophe was an Baker WJ. 2015. A revised delimitation of the rattan genus
extraordinary and generous individual, a selfless and Calamus (Arecaceae). Phytotaxa 197: 139–152.
open collaborator in a time when science is becoming Baker WJ, Allkin R, Barker AM, Macıa MJ, Theys A,
only more competitive and self-centred. He was a Villalba S, Gardiner LM. 2015. Bioinform atica y la
brilliant palm biologist, able to move fluidly from the familia de las palmas. In: Balslev H, Macıa MJ, Navarrete
finest detail to the biggest of ‘big picture’ ideas. As H, eds. Cosecha de palmas en el noreste de Suram erica:
an incisive critic, his views were argued meticu- bases cientıficas para su manejo y conservaci
on. Quito: Pon-
lously, but also shared kindly. Jean-Christophe gave tificia Universidad Catolica del Ecuador, 213–222.
his time, ideas and resources freely to all around Baker WJ, Asmussen CB, Barrow SC, Dransfield J,
him, seniors, peers and those fortunate enough to Hedderson TA. 1999a. A phylogenetic study of the palm
come under his wing. The palm community has lost family (Palmae) based on chloroplast DNA sequences from
a great friend who has made an indelible mark on the trnL-trnF region. Plant Systematics and Evolution 219:
our field. 111–126.
Baker WJ, Couvreur TLP. 2012. Biogeography and distri-
bution patterns of Southeast Asian palms. In: Gower D,
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Botanical Journal of the Linnean Society, 2016.

Beyond Genera Palmarum: progress and prospects in

Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, UK

ADDITIONAL KEYWORDS: Arecaceae – biodiversity – classification – next-generation sequencing –

INTRODUCTION & Beck, 1990), their keystone role in ecosystems

Sabaleae 19. Sabal Adans.

Cryosophileae 20. Schippia Burret

Phoeniceae 31. Phoenix L.

Chuniophoeniceae 51. Chuniophoenix Burret

Caryoteae 55. Caryota L.

Corypheae 57. Corypha L.

Ceroxyleae 67. Ceroxylon Bonpl. ex DC.

Phytelepheae 71. Ammandra O.F.Cook

Oranieae 85. Orania Zipp.

Sclerospermeae 86. Sclerosperma G.Mann & H.Wendl.

Roystoneeae 87. Roystonea O.F.Cook

Manicarieae 106. Manicaria Gaertn.

Euterpeae 107. Hyospathe Mart.

Geonomateae 112. Welfia H.Wendl.

Leopoldinieae 118. Leopoldinia Mart.

Rhopalostylidinae 166. Rhopalostylis H.Wendl. & Drude

Figure 4. Jailoloa halmaherensis, Halmahera, Indonesia. A, Habit. B, Fruit. Photographs: C. D. Heatubun.

Figure 5. Manjekia maturbongsii, Biak, Indonesia. A, Habit. B, Inflorescence. Photographs: W. J. Baker.

Figure 6. Wallaceodoxa raja-ampat, Waigeo, Indonesia. A, Habit. B, Inflorescence. Photographs: C. D. Heatubun.

GENERA LOST SINCE GP2 WALLICHIA (CORYPHOIDEAE: CARYOTEAE).

ter to Wendlandiella Dammer, with Hyophorbe Laccospadicinae, Rhopalostylidinae; Baker et al.,

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