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Review of Vegetative Branching in the Palms (Arecaceae).

Abstract Vegetative branching is common in the palms (Arecaceae). However, current terms to describe vegetative branching diversity are not consistent and do not cover the full range of branching types. In this study vegetative branching types in the palms were reviewed and defined, and the phylogenetic distribution of palm branching types was described. Branching types were described from a literature review and field observations; 1903 species representing all 181 genera were included. Five branching types were found: lateral axillary branching, shoot apical division, false vivipary, abaxial branching, and leaf-opposed branching. Most species (55%) exhibited no vegetative branching. Lateral axillary was the most common branching type. Lateral axillary branching and shoot apical division were predicted to be the earliest-evolved branching types. The present study suggests that branching types have different evolutionary histories, and it is likely that the solitary habit is more common now than when palms initially diverged from commelinid relatives.

Keywords Arecaceae * Branching * Commelinid monocots * Monocotyledons * Palmae * Palm phylogeny * Vegetative anatomy * Vegetative propagation

Vegetative Branching in the Palms and the Need for a New Classification System

Branching is the outgrowth or division of a meristem and results in a new axis. Plants can branch sexually, producing an axis used for sexual reproduction, or vegetatively, producing a separate and genetically identical vegetative axis (Doust & Doust, 1988). The vast majority of plants display some form of vegetative branching, which results in a great diversity in plant form and architecture (Bell & Tomlinson, 1980). Plants branch vegetatively in three ways: axillary (occurring in the leaf axils), apically (at the apex of the shoot), or adventitiously (in neither of the previous two locations) (Halle et al., 1978). Axillary branching, the most common type of branching in plants, has two forms that account for much of the architectural diversity displayed in plants: long and short shoots. Short shoots are specialized units, usually producing photosynthetic or reproductive structures or spines that bear no indeterminate lateral branches (Halle et al., 1978). Long shoots grow, add height, and can proliferate to produce additional lateral axillary branches that become either long or short shoots.

Vegetative branching is common in the monocots, where it is used as a mechanism to increase in size, since most monocots lack secondary growth (Halle et al., 1978). The three main terms used to describe branching in the monocots are (1) axillary, (2) dichotomous, and (3) adventitious branching (Tomlinson, 1973). However, these terms are not consistently used in descriptions of monocot branching diversity.

While similar vegetative branching types exist in the palms (Arecaceae Bercht. & J.Presl) and their monocot relatives, the terminology to describe these types is not uniform and many terms have been applied to the same branching type (Tomlinson, 1961; Tomlinson, 1971; Fisher, 1973; Fisher & Tomlinson, 1973; Fisher, 1974; Fisher et al., 1989; Mendoza & Franco, 1998; Fisher & Zona, 2006). Detailed descriptions are often greatly simplified in the popular palm literature (Tomlinson, 1973; Dransfield et al., 2008a, b). Consequently, the current branching vocabulary for palms does not consistently and accurately describe the diversity of vegetative branching in the family.

The three vegetative branching types commonly described in palms are similar to the branching types used for the monocots (Tomlinson, 1990): axillary branching, apical dichotomous branching, and non-axillary branching. However, branching terms have also been classified depending on variation in outgrowth. Axillary branching, the most commonly described, is used to describe the formation of a primordial bud in the leaf axil at the base of orthotropic (vertical) shoots. If axillary branches grow erect immediately, they create branch types called basal suckers. If the basal sucker grows horizontally before turning to grow erect, it forms a rhizome (Tomlinson, 1990). Rhizomatous branching is occasionally classified as its own, unique branching type. Apical dichotomous branching occurs when the apical meristem of the stem bifurcates, creating two apical meristems. In palms, species differ in whether the meristem splits into two even (isotomous) or uneven (anisotomous) parts (Tomlinson & Moore, 1966; Gola, 2014). In palm literature, the term dichotomy has been used incorrectly to imply equality of outgrowth (Tomlinson. 1990). Non-axillary branching describes a branch that does not arise in the leaf axil (Tomlinson, 1973). The term, however, does not further differentiate among locations of the branch (non-apical portions of the stem, lamina or inflorescence), which can differ among taxa.

Classifying branching based on location of meristem for some types (non-axillary, dichotomy) vs. variation in outgrowth for others (basal suckers, rhizomatous) means that branching terms are defined at different points during development. Since vegetative branching terms are currently classified at different developmental levels, the phylogenetic distribution of branching in the palms cannot be appropriately described. A new, internally-consistent classification of branching is needed to understand branching type phylogeny in the palms.

Understanding the relationship between phylogeny and branching type will increase our understanding of the evolution and ecology of vegetative branching in the palms and will provide a framework for understanding branching in all monocots. The purpose of this study was to (1) identify, define and classify the types of vegetative branching in the palm family Arecaceae and (2) describe the phylogenetic distribution of these branching types in palms.

A New Vegetative Branching Classification System in the Palms

Determining Vegetative Branching Types in the Palms

The classification system used was based on vegetative branching meristem location. Each branching type was defined by (1) branching meristem (axillary, apical, nonaxillary); and, if non-axillary, (2) location of branch (inflorescence, leaf base or stem). Branching type(s) of species were identified from literature reviews of journal articles and books describing branching patterns and from analysis of living specimens in the palm collections at Fairchild Tropical Botanic Garden and Montgomery Botanical Center (Coral Gables, FL, USA) (Table 2). Branching types proposed in this study, with terms found in the literature, are presented in Table 1.

The five branching types were lateral axillary branching, shoot apical division, false vivipary, abaxial branching and leaf-opposed branching. A dichotomous key was created to facilitate understanding and recognition of each branching type (Table 3). Lateral axillary branching was defined as vegetative outgrowth of an axillary meristem on the vegetative shoot (stem) (Fig. lb). Many species display lateral axillary branching but can also not branch, presenting a solitary stem; these species were classified as having lateral axillary branching. Shoot apical division was defined as the division of the apical meristem into two equal or unequal meristems (Fig. 1c). False vivipary was defined as adventitious vegetative outgrowth of buds in the apical bracts of an inflorescence that eventually rooted in the soil and produced vegetative shoots (Fig. 1d). Abaxial branching was defined as the vegetative outgrowth of an adventitious meristem located on the abaxial surface of the leaf at the base of the leaf sheath (Fig. le). Leaf- opposed branching was defined as the vegetative outgrowth of an adventitious meristem borne on the stem, opposite the lamina and petiole and enclosed within the edges of the leaf sheath (Fig. 1f) Branching type combinations can also occur; two branching combinations were found in the palms: shoot apical dichotomy + lateral axillary branching; and false vivipary + lateral axillary branching.

The above branching type names were assigned using the uniqueness and priority principles of botanical nomenclature (Greuter et al., 1999). The term dichotomy was not used because it has been defined multiple ways and the evidence for whether shoot apical division results from an equal apical division was often lacking. Most commonly, dichotomy implies equal division of the shoot apical meristem (Tomlinson, 1990), but the term has also been defined as two independently functioning axes (Gola, 2014) or as two more or less equal axes. Thus, the term has been used to describe both a developmental process (equal division of the shoot apex) and the result of branch outgrowth. Since there was discrepancy among definitions and usage of dichotomy, the term apical division was used to describe any division of the apical meristem (uniqueness principle). The terms shoot apical division and false vivipary needed additional clarification because these terms were not clearly defined in previous literature. The term false vivipary was selected because it was first published in the grass literature to describe a phenomenon similar to what was found in the palms (priority principle) (Van der Pijl, 1982; Bell & Bryan, 2008).

Descriptions of Vegetative Branching Types in the Palms

The classification system described above was used to describe branching in palm species, genera and subfamilies. Species were recognized following the accepted species in the Kew World Checklist of Palms on February 5, 2016 (Goverts et al., 2011). The numbers of species, genera and subfamilies with each branching type were counted to determine the most abundant branching type and combination found at each taxonomic level.

In total, 181 genera (out of 181 genera in the family, 100% genus coverage), comprising 1903 species (out of 2501 species in the family, 76% species coverage), were sampled (Table 2). The five branching types described above were identified in the species considered; these were distinguished from the solitary phenotype, which had no vegetative branching. Some species displayed more than one branching type, referred to as branching combinations. Two branching combinations were observed: shoot apical division + lateral axillary; and false vivipary + lateral axillary. Most commonly, species exhibited no vegetative branching; four of the five subfamilies, 147 genera (81% of genera), and 1043 species (55% of observed species) were solitary (Table 2, Fig. la). Some species were found with a branching type or as a solitary individual (175 species, 9% of observed species).

1) Lateral axillary branching was the most widely distributed vegetative branching type in the palms; it was described in four of five subfamilies, 61 genera (34% of genera), and 646 species (34% of observed species) (Table 2, Fig. 2). Four forms of lateral axillary branching were identified: basal suckering, rhizomatous branching, aerial suckering and displaced axillary branching. Basal suckering was defined as lateral axillary branching where the branches grew orthotropically (vertically) immediately and were restricted to the base of the parent shoot. Basal suckering was the most common form of lateral axillary branching, found in at least 600 palm species. Basal suckers may be produced throughout the life of an individual or basal suckers may be produced only during certain times. For example, Plectocomia Mart. & Blume species (15 species) and two Licuala Thunb. species (L. celebica Miq. and L. gracilis Blume) produced basal suckers after a period of dormancy, usually after the death of the parent shoot (Tomlinson. 1990). Phoenix L. species produced basal suckers until they were sexually reproductive and then stopped producing basal suckers (Tisserat & DeMason, 1985).

Rhizomatous branching was defined as lateral axillary branching where branches were restricted to the base of the stem but grew plagiotropically (horizontally) for some time before growing orthotropically (vertically). At least 33 species exhibited rhizomatous branching. Rhizomatous branching was found in combination with basal suckering in two species (Acoelorrhaphe wrightii H.Wendl. ex Becc. and Cyrtostachys renda Blume).

Aerial suckering was defined as basal suckering that was not restricted to the base of the stem but also occurred on aerial portions of the stem. Wendlandiella gracilis sub. Polyclada Dammer, Linospadix apetiolatus Dowe & A.K. Irvine, Hyospathe elegans hort ex Hook. F., and Geonoma baculifera Kunth exhibited aerial suckering (Tomlinson, 1990; Chazdon, 1991). In this study, aerial suekering was placed within lateral axillary branching because the branching mechanism for aerial suekering was developmentally the same as the branching mechanism for lateral axillary branching, and species with aerial lateral axillary branching had basal suekering as well.

Displaced lateral axillary branching, found in Korthalsia Blume, was defined as vegetative axillary meristems that were initiated in the axil of the first or second leaf primordium and then were displaced during development on to the internode above or onto the base of the leaf above. The displaced lateral axillary branching type was placed within lateral axillary branching because the branching mechanism was lateral axillary and the transition out of the axil occurred after initiation of the meristem (Fisher & Drans field, 1979).

2) Shoot apical division was distributed throughout the palms, having been described in four subfamilies, seven genera (3% of genera), and 21 species (1% of observed species) (Table 2, Fig. 2). Three forms of shoot apical division were identified: isotomy, anisotomy and Nannorrhops branching.

Isotomy, which is equal apical division followed by equal growth, has been studied anatomically in three palm genera and eight species: Hyphaene Gaertn. (H. compressa H. Wendl, H. coriacea Gaertn., H. dichotoma (J.White Dubl. Ex Nimmo) Furtado, H. reptans Becc., and II. thebaica Mart.), Nypa fruticans Wurmb. and Matricaria saccifera Gaertn. (Gola, 2014). Leaf arrangement and equal forking in divided crowns of mature plants suggest isotomy, but anatomical study of shoot apical development is needed for confirmation of other species (Fisher, personal correspondence).

Anisotomy, which is unequal division followed by differential growth, was exhibited by Eugeissona Griff (E. ambigua Becc., E. brachystachys Ridl., E. insignis Becc., E. minor Becc., E. triste Griff., and E. utilis Becc.). Anisotomous division was so unequal in Eugeissona species that the division appeared to be lateral axillary branching on non-basal portions of the stem (Fisher et al., 1989). Apical division in palms has been reported to range from equal (isotomous) to unequal (aniosotomous) division. In Chamaedorea cataractarum Mart., the anisotomous division of the apical meristem occurred very early in development, and as the stems matured, the division appeared to be equal (Fisher, 1973). Only developmental studies showed that the division did not initiate equally.

Nannorrhops branching, which has not previously been recognized as a distinct branching type, was defined as equal apical division with branch-pair differentiation. For example, in Nannorrhops ritchiana H. Wendl., the apical meristem divides into one fertile and one vegetative branch (Tomlinson & Moore, 1968).

3) False vivipary has been described in two subfamilies, three genera (1% of genera), and ten species (0.5% of observed species) (Table 2, Fig. 2): Calamus Auct. ex. L. (C. castaeneus Griff, C. dianbaiensis C.F.Wei, C. gamblei Becc., C. ingens (J.Dransf.) W.J.Baker, C. kampucheaensis A.J.Hend. & Hourt, C. nematospadix Becc., and C. pygmaeus Becc.), Salacca Reinw. (S. flabellata Furtado, and S. wallichiana Mart.), and Socratea salazarii H.E.Moore (Fisher & Mogea, 1980; Baker et al., 2000; Pintaud & Millan, 2004; Rupert et al., 2012). In each account of false vivipary, different terms were used to describe the phenomenon (Fisher & Mogea, 1980; Baker et al., 2000; Pintaud & Millan, 2004; Rupert et al., 2012). The architectures of the palms with false vivipary were different, yet the branching of the inflorescence was the same-vegetative shoots formed at the apex of the inflorescence. If the shoot reached the ground, it rooted and a shoot grew upward. Calamus gamblei, C. pygmaeus and C. nematospadix are all climbing rattans (Dransfield, 1992), Socratea salazarii is an erect and usually solitary palm (Pintaud & Millan, 2004), while Salacca flabellata is an acaulesent palm (Furtado, 1949).

4) Abaxial branching was described in one subfamily (Arecoideae), two genera (2% of genera), and seven species (0.3% of observed species) (Table 2, Fig. 2). In abaxial branching, a vegetative branch originating on the abaxial surface of the leaf sheath occurred on the basal and intermediate internodes of orthotropic stems in Oncosperma Blume species and Dypsis lutescens (H. Wendl.) Beentje & J. Dransf. Species with abaxial branching usually do not display lateral axillary branching.

5) Leaf-opposed branching was described in one subfamily (Calamoideae), two genera (1% of genera), and seven species (0.3% of observed species). Leaf-opposed branching occurred on basal internodes and on aerial internodes of the stem, as in the liana Myrialepis paradoxa (Kurz.) J. Dransf. Axillary branching, leaf-opposed branching and abaxial branching are distinct types of stem nodal meristems based on location and position of the branching meristem (Fig. 2). In axillary branching, the meristem is located in the axil of the leaf. In abaxial branching, the vegetative branching meristem is located on the abaxial surface of the leaf sheath. In leaf-opposed branching, the branching meristem is borne on the stem, enclosed by the edges of the leaf sheath and opposite to the lamina and petiole.

Individuals within a species sometimes displayed more than one branching type at a time, here called branching combinations. The two branching combinations found were shoot apical division + lateral axillary and false vivipary + lateral axillary branching. Shoot apical division + lateral axillary branching was exhibited by one species of Basselinia Vieill. (Arecoideae), all 27 species of Korthalsia Blume (Calamoideae), two species of Hyphaene (Coryphoideae) and monospecific Nannorrhops ritchiana (Coryphoideae). False vivipary + lateral axillary branching was exhibited by five species of Calamus (Calamoideae), and Socratea salazarii (Arecoideae) (Table 2).

Phylogenetic Distribution of Vegetative Branching Types in the Palms

Subfamily-level and genus-level phylogenies were used to examine the phylogenetic distribution of branching types. The phylogeny from Baker et al. (2009) was selected for character mapping because it had the most recent genus-level phylogeny. Adjustments were made for new and deleted taxa (Dransfield et al., 2008a, b; Baker & Bacon, 2011; Bernal & Galeano, 2013; Baker, 2015; Noblick & Meerow, 2015). Branching types were used for character mapping, since the specific branching type was the character that was retained or lost. The Mesquite software (Maddison & Maddison, 2011), a software package used by evolutionary biologists to analyze comparative data, was used to map vegetative branching onto the published cladograms. Ancestral branching types were determined using the most parsimonious tree in Mesquite. A subfamily level cladogram was analyzed to predict the ancestral branching type for the family. Cladograms for Arecoideae, Calamoideae and Coryphoideae were analyzed to predict the ancestral branching type for each of these three subfamilies. A cladogram for Ceroxlyloideae was not included because this subfamily had no vegetative branching except for a single species, Ravenea deliculata Rakotoarin. A cladogram for Nypoideae was not included because it is monospecific (Nypa fruticans Wurmb).

At the subfamily level, lateral axillary branching and shoot apical division were predicted as the ancestral vegetative branching types (Fig. 3). The solitary state (no vegetative branching) was also an ancestral state. False vivipary evolved a minimum of two times: once in the Calamoideae and once in the Arecoideae (Fig. 3). Abaxial branching evolved a minimum of two times in the Arecoideae (Oncosperma and Dypsis). Leaf-opposed branching evolved two times in the Calamoideae, in Myrialepis Becc. and in Calamus.

The Calamoideae, the most basal and second largest subfamily (659 species), was the most diverse in vegetative branching types (Table 2); it exhibited four branching types and both branching combinations. On average, one branching type was exhibited in a genus. With three branching types, Calamus exhibited the most branching types in Calamoideae. The ancestral branching type of Calamoideae was predicted to be lateral axillary branching (Fig. 4). In the Calamoideae, more species had vegetative branching (341 species, 86% of observed Calamoideae species) than the solitary habit (58 species, 14% of observed species). Lateral axillary branching evolved a minimum of one time in the Calamoideae. In five genera, all species had lateral axillary branching; these genera were Laccosperma G. Mann & H.Wendl. (six species), Eremospatha Mann & H. Wendl. (11 species), Oncocalamus Mann & H. Wendl. (five species), Mauritiella Burret (four species), Plectocomia Mart. & Blume (15 species), and Plectocomiopsis Becc. (six species). Ten species in two genera in the Calamoideae displayed false vivipary: Calamus (eight) and Salacca (two). Shoot apical division evolved at least two separate times in the Calamoideae; species of Eugeissona and Korthalsia exhibited shoot apical anisotomy. Leaf-opposed branching, described only in the Calamoideae, was the least common branching type in the Calamoideae; Myrialepis (one species) and Calamus (seven species) were the only two genera with leaf-opposed branching. In Calamoideae, 15% of observed species did not display any branching, and two genera, Mauritia L.f. (two species) and Pigafetta (Blume) Becc. (two species) exhibited no branching.

The majority of the Coryphoideae, the third largest subfamily (492 species), were solitary, exhibiting no vegetative branching (39 genera/283 species, 74% of observed Coryphoideae species). Some members of Coryphoideae displayed lateral axillary branching (16 genera /79 species, 20% of observed Coryphoideae species) or shoot apical division (three species of Hyphaene, 0.7% of observed Coryphoideae species). One branching combination, shoot apical division + lateral axillary, was found (two species of Hyphaene and Nannorrhops ritchiana) (Table 2). The ancestral branching type of the Coryphoideae was lateral axillary branching (Fig. 5). The genus Hyphaene (eight species) exhibited the most branching types and combinations in the Coryphoideae (two types-lateral axillary and shoot apical division and one branching combination (shoot apical division + lateral axillary)). All species in subtribe Rhapidinae, except for Trachycarpus H.Wendl., exhibited lateral axillary branching: Chamaerops L. (one species), Rhapidophyllum H. Wendl. & Drude (one species), Maxburretia Furtado (three species), Rhapis L.f. (ten species) and Guihaia J. Dransf., S.K. Lee & F.N. Wei (two species). The subtribe Rhapidinae was the only Coryphoideae clade higher than genus-level where lateral axillary branching was retained throughout all species of the clade. Lateral axillary branching evolved at least 12 times, and shoot apical division evolved at least two times in Coryphoideae. There were no species in the Coryphoideae that displayed false vivipary, abaxial branching or leaf-opposed branching.

The Arecoideae, the largest subfamily (1376 species), exhibited four branching types and two branching combinations (Table 2). The majority of the Arecoideae exhibited no branching (59%, 657 observed species). Dypsis Noronha ex Mart, exhibited three branching types, which was the most genus-level branching types for the Arecoideae. The ancestral branching type of the Arecoideae palms was lateral axillary (Fig. 6). Five genera in Arecoideae had no solitary species (i.e., all species exhibited vegetative branching): Iriartella H.Wendl. (two species), Wettinia Poepp. ex Endl. (21 species), Jubaeopsis Becc. (one species), Podococcus Mann & H.Wendl. (two species) and Sclerosperma G. Mann & H. Wendl. (three species). Shoot apical division evolved at least four times, occurring in Allagoptera, Basselinia, Dypsis, and Matricaria. However, shoot apical division was not easily observed in these genera. In Basselinia, Dypsis and Matricaria, shoot apical division occurs early in development of the stem (Moore & Uhl, 1982; Fisher & Zona, 2006). Allagoptera Nees is a creeping palm and apical division occurs low to the ground. While shoot apical division was not as obvious as in Hyphaene (Coryphoideae), morphological signs of apical division (forking) are still present and observable in Allagoptera. False vivipary evolved once in Socratea salazarii. Abaxial branching evolved twice, occurring in Dypsis lutescens and Oncosperma. (Tables 4, 5, 6 and 7).

The Ceroxyloideae, the fourth largest subfamily (47 species), had one species that branched vegetatively. The ancestral state of Ceroxyloideae was no branching; 99% of species exhibited no vegetative branching. Ravenea deliculata, from the largest genus in Ceroxyloideae (Ravenea, 21 species), displayed lateral axillary branching both basally and aerially (Rakotoarinivo, 2008). On the basis of number of species, the Ceroxyloid palms exhibited fewer branching types and combinations than expected.

Nypoideae, the smallest subfamily (one species, Nypa fruticans), exhibited one branching type (shoot apical division), and the ancestral branching type was shoot apical division.

Evolutionary History of Vegetative Branching in the Palms

The phylogenetic distribution of vegetative branching types suggests that lateral axillary branching is the ancestral branching type and that branching evolved before palm divergence from immediate ancestors. Monocots evolved in the mid/late Jurassic period, about 160 million years ago. (Wikstrom et al, 2001). Recent evidence suggests palms diverged in the Turanian, about 90 million years ago (Harley, 2006). Newer findings demonstrate that palms diverged much earlier than commelinid relatives (Barrett et al., 2016). At some point between monocot evolution and evolution of the current palm species, a diversity of branching types evolved in the palms.

While fossilized remains of palms are distributed throughout the fossil record, stems are less commonly found as fossils, and multiple-stemmed fossils are missing from the literature entirely (Erwin & Stockeny, 1994; Harley, 2006). There is a form genus for palms with rhizomatous stems, Rhizopalmoxylon (Palmoxylon is the form genus for petrified wood) and there is apparently no literature on its architecture, specifically, whether there are multiple stems per individual (Harley, 2006). Nypa fruticans, a multi-stemmed palm once widespread on many continents, has fossilized pollen, fruit, and leaves but no stem fossils (Gee, 2001; Mehrotra et al., 2003). It is difficult to determine when branching evolved in Nypa, and in palms in general, without any branching or architectural information from fossils.

While the fossil record does not distinguish the ancestral branching type, it is possible to predict evolutionary trajectories for each branching type. Because of the prevalence of lateral axillary branching in commelinid relatives, as well as in the palm family, lateral axillary branching may have been present before the divergence of palms. Lateral axillary branching is a common branching type, and more common than the solitary habit, in Poaceae Barnhart (Holtuum, 1955; Ward & Leyser, 2004; McSteen & Leyser, 2005; Doust, 2007), Cyperaceae Juss. (Rodigues & Maranho-Estelita, 2009), Zingiberaceae Martinov (Bell, 1979) and Dasypogonaceae Dum. (Clifford et al., 1998); Dasypogonaceae, sister to the palms, is entirely rhizomatous. Therefore, lateral axillary branching may share a common evolutionary history throughout the commelinid relatives.

While analysis from the present review suggest that shoot apical division is an ancestral branching type, shoot apical division in the commelinids is described only in Strelitzia Banks (Strelitziaceae) (Fisher, 1976). Also, shoot apical division is not nearly as widespread through the palm family as lateral axillary branching. It is likely that shoot apical division evolved after the divergence of palms.

This review suggests that the remaining branching types-false vivipary, abaxial branching, and leaf-opposed branching-probably evolved after the divergence of palms. False vivipary and leaf-opposed branching are found in commelinid relatives. False vivipary is common in the Poaceae (Chlorophytum comosum (Thunb.) Jacques, Deschampsia alpina (L.) Roem. & Schult., Festuca ovina var. vivipara L., Dactylis glomerata L., Poa x jemtlandica K.Richt.), as well as the Zingiberales (Costaceae Nakai and Marantaceae R.Br.). In Costaceae (Zingiberales) and Marantaceae (Zingiberales), bulbils are produced in the axils of inflorescence bracts (Jenik, 1994), a branching type closely related to false vivipary. Leaf-opposed branching is found in Musa L. (Fisher, 1973). However, the presence of these branching types in commelinid relatives does not mean that the ancestral palm displayed these branching types. Results from this study suggest that false vivipary and leaf-opposed branching evolved later in palm evolutionary history. False vivipary and leaf-opposed branching displayed by the palms and their commelinid relatives are most likely an example of homoplasy, and distinct evolutionary histories led to similar branching types. Abaxial branching, however, has been described only in the palms and may be a branching type unique to the family.

The evolutionary history of branching types may not be easily determined because the evolution (and loss) of branching types in the palms is continuous and occurred at different speeds among subfamilies (Faurby et al., 2016). The different evolutionary trajectories of vegetative branching in subfamilies Calmoideae and Arecoideae exemplify that evolution (and loss) of branching types is continuous and occurred at different speeds. In Calamoideae, most commonly an entire genus shares a branching type. Branching types in Calamoideae do not appear to be changing at the species level. Alternatively, in Arecoideae, species within a genus may not share a common branching type. In the Arecoideae, the genera are mostly solitary but have a few branching species. There are two distinct trajectories that could lead to a primarily solitary genus with a few branching species in Arecoideae. Either the ancestor to the genus did not branch and the ability to branch has re-evolved in a few species, or the ancestor did branch and the extant species have lost the ability to branch. Evolution of branching in palms may be influenced by differences in the ecology of different taxa.

Ecology of Vegetative Branching in the Palms

Regardless of evolutionary history, vegetative branching is less common in palms than in their commelinid relatives (Tomlinson, 1973). Like most monocots, including their commelinid relatives, palms do not produce secondary xylem (wood) from a vascular cambium, which limits their ability to make large trees. One of the main differences between palms and their close relatives, however, is their large, strong, woody trunks. Palms form a woody trunk through cell thickening and lignification on the surface layers of the cells in the outer cortex. It is possible that the lignification of the surface of the palm stem prevents activation and growth of dormant axillary buds. The lignified stem may have imprisoned the buds, and the ability to branch via axillary buds was lost over evolutionary time. Lignified stems (woody trunks) presumably have been selected because they increase fitness and the chance of survival (Schluter, 2001). Vegetative branching may be less common in the palms because there was selection for palms with thicker, taller trunks rather than thinner trunks that can branch (Henderson, 2002a).

It is important to note that all palms, even solitary palms, branch sexually. All palms have meristems that produce inflorescences. Similar branching types exist in vegetative and sexual branching in the palms. The most common type of sexual branching is axillary, exhibited by the vast majority of palms, where an inflorescence is produced from a bud in the leaf axil (Dransfield et al., 2008a, 2008b). In sexual branches, there is variation in types of displaced axillary branching that results in different branching patterns (Fisher & Maidman, 1999). In Salacca and Kerriodoxa J. Dransf., the sexual bud is borne in the leaf axil but may be captured by the subtending developing leaf, and the bud emerges through a slit on the abaxial side of the leaf sheath (Fisher & Mogea, 1980). In a few genera in the Calamoideae (Korthalsia, Calamus, Myrialepis, Plectocomia and Plectocomiopsis) the bud is displaced longitudinally and is adnate to the internode and leaf sheath above the node of origin (Fisher & Dransfield, 1977; Fisher & Mogea, 1980). In sexual apical branching, the apical meristem produces a large determinate inflorescence (i.e., Corypha L. and Tahina J. Dransf. & Rakotoarin.) (Dransfield et al., 2008a, 2008b). False vivipary is a combination of asexual and sexual branching, where the sexual branching reverts to vegetative branching in the inflorescence, using the same stem system. Abaxial and leaf-opposed sexual branching types have not been recorded in the palms.

While sexual branching is more common in the palms than vegetative branching, there are ecological benefits of vegetative branching. First, branching increases net primary productivity for the individual genet. When the palm branches vegetatively, it produces more crowns with more leaves, and the increase in leaves could increase photosynthetic potential (Duncan, 1971). In palms like Serenoa Hook.f., Allagoptera and Nypa, a creeping habit allows the stem to produce more roots (Tomlison, 1990; Fisher & Jayachandran, 1999). The creeping habit can support a greater photosynthetic potential. However, if the branching type is shoot apical division and the habit is erect (as in Hyphaene dichotoma), the trunk may be unable to support more crowns physically and physiologically.

Another ecological consequence of branching is that having multiple stems increases chances of an individual's survival in disturbance-prone environments, such as under-stories of rainforests and coastal strands (Tomlinson, 1990). Certain species of Chamaedorea Willd. and Geonoma Willd. live in disturbance-prone environments in the understory of rainforests, where falling debris poses a threat to their survival (Bullock, 1980; Clark & Clark, 1989; Chazdon, 1992; James, 2013;). A solitary palm only has one apical meristem and damage to that apical meristem results in death of the plant. In a multiple-stemmed palm, a genet can survive after damage to a single apical meristem. Thus, having multiple stems increases their chance of surviving a fallen branch or trunk of a large canopy tree. The understory palms Geonoma baculifera and Hyospathe spp., which have vegetative branching, exemplify this habit. These species are clumping palms that grow in the understory of rainforests. If damage occurs to a terminal apical meristem of these species, aerial axillary buds grow to produce plantlets. The stem eventually falls and the plantlets root, producing new ramets.

Nypa and Allagoptera also colonize environments where water level and substrate are unstable. Nypa fruticans colonizes coastal strands where water level is in constant flux and muddy banks are unstable (Tomlinson, 1990). Allogoptera colonizes sandy beaches and dunes, where water level changes daily and the dunes are likely to change shape (Dransfield et al., 2008a, 2008b). For Nypa and Allogoptera, branching is by shoot apical division on horizontally-growing stems, which allows them to form large monotypic stands. If damage occurs to a stem, such as meristem or stem rot from prolonged flooding, many other apical meristems exist that will survive and continue branching. Nypa and Allagoptera may also help stabilize these unstable environments.

While vegetative branching is a survival mechanism, as in Chamaedorea, Geonoma, Hyospathe, Nypa and Allagoptera, it is also a mechanism for clonal reproduction (Mogie, 1992). In unstable environments, such as flood plains, coastal strands and habitats with frequent fire or droughts, seed germination and establishment can be difficult. The ability of an individual to branch vegetatively and reproduce asexually ensures continued reproduction of the species into the next generation.

The Calamoideae epitomize the ecological benefit gained from vegetative branching. They are an interesting group because most species exhibit vegetative branching and climb prolifically. A major innovation in the Calamoideae was their liana habit (Gianoli, 2004; Couvreur et al., 2014). These palms climb, branch and dominate the canopy of Asian rainforests (Dransfield, 1992; Dransfield, 1997; Dransfield et al., 2008a, 2008b). Vegetative branching, therefore, allows the Calamoideae to climb through and explore the canopy prolifically. These palms colonize the canopy more efficiently than unbranched palms could. Vegetative branching allows the Calamoideae to exploit the canopy habitat; at the same time, the liana habit means that the plants do not invest in large woody trunks.

The Calamoideae also contain the greatest number of species that branch through false vivipary. False vivipary is interesting ecologically because it is only successful if the inflorescence is able to root in the forest floor, presumably when the crown is close to the ground (Bell & Tomlinson, 1980). In grasses displaying false vivipary, the inflorescence is never more than a few centimeters from the ground and the plantlet can easily reach the soil to root. In palms, false vivipary occurs on both erect (Socratea), climbing (Calamus) and acaulescent (Salacca) stems and is successful in all of these habits (Fisher & Mogea, 1980; Dransfield, 1992; Dransfield, 1997; Baker et al., 2000; Pintaud & Millan, 2004; Rupert et al., 2012). For all species that exhibit false vivpary, successful rooting of the false viviparous shoot has been described for sterns near the soil, but the exact heights have not been recorded. There are at least four possible relationships between stem height and successful false vivipary. First, there could be no relationship; false viviparous shoots could form at any height in the canopy and successfully root in the soil. No relationship between height of the shoot and successful rooting is the least likely of the scenarios, since the viviparous shoot may have a very long distance to reach the soil. Second, the false viviarpous shoots could form at any height in the canopy but not root successfully above a certain stem height (critical height). Alternatively, false vivipary may only occur on stems below a critical height, and stems of Calamus and Socratea may stop producing false viviparous inflorescences once they reach a certain height. The fourth possibility is that the viviparous shoot could abscise and fall to the forest floor. More studies on the morphology and ecology of false vivipary in the palms are needed in order to determine which mechanism occurs in which species.

Summary

This review of vegetative branching in palms reclassified vegetative branching types in the palms based on location of the branching meristem and then described variation within those types. This review demonstrated that diverse branching types exist in the Arecaceae. The phylogenetic distribution of shoot apical division, false vivipary, abaxial branching and leaf-opposed branching within the palm family and subfamilies gives insights into palm evolutionary history and ecological constraints. This review highlights how the lack of an overview of vegetative branching in the palm literature and the use of multiple, often poorly-defined terms for similar branching types has inhibited our understanding of basic palm evolution and ecology.

https://doi.org/10.1007/s 12229-018-9200-2

Acknowledgements We would like to thank Dr. Scott Zona for his extensive guidance and help editing. Dr. Jack Fisher for his help with the anatomical analysis and Dr. P. Barry Tomlinson for his help with the literature review and for anatomical insights. SME gratefully acknowledges the support of the Florida International University Dissertation Year Fellowship and the FIU International Center for Tropical Botany, both of which provided time for completing her dissertation; this research was completed in partial fulfillmet of her dissertation requirements. She also acknowledges the support of a Fairchild Tropical Botanic Garden Graduate Fellowship, which supported her during her dissertation research.

References

Abrahamson, W, G. 1999. Episodic reproduction in two fire-prone palms, Serenoa repens and Sabal etonia (Palmae). Ecology, 80: 100-115.

Bacon, C.D., & Baker, W.J. 2011. Saribus resurrected. Palms. 55: 109-116.

Baker, W.J. 2015. A revised delimitation of the rattan genus Calamus (Arecaceae). Phytotaxa, 197: 139-152

Baker, W.J., & Dransfield, J. 2002a. Calamus longipinna (Arecaceae: Calamoideae) and its relatives in New Guinea. Kew bulletin. 853-866

Baker, W.J., & Dransfield, J. 2002b. Calamus maturbongsii, an unusual new rattan species from New Guinea. Kew bulletin, 725-728.

Baker, W.J., & Dransfield, J. 2007. Arecaceae of Papua. The ecology of Papua. 6: 359.

Baker, W.J., & Dransfield, J. 2014. New rattans from New Guinea (Calamus, Arecaceae). Phytotaxa. 163(4): 181-215.

Baker, W.J., Dransfield, J., & Henderson, T.A. 2000. Phylogeny, character evolution, and a new classification of the calamoid palms. Systematic Botany, 25: 297-322.

Baker, W.J., Bayton, R. P., Dransfield, J., & Maturbongs, R. A. 2003. A revision of the Calamus aruensis (Arecaceae) complex in New Guinea and the Pacific. Kew bulletin, 351-370.

Baker, W. J., Zona, S., Heatubun, C. D., Lewis, C. E., Maturbongs, R. A., & Norup, M. V. 2006. Dransfieldia (Arecaceae)--a new palm genus from western New Guinea. Systematic Botany, 31: 61-69.

Baker, W.J., Savolainen, V., Asinussen-Lange, C.M., Chase, M.W., Dransfield, J., Forest, F., & Wilkinson, M. 2009. Complete generic-level phylogenetic analysis of palms (Arecaceae) with comparisons of supertree and supermatrix approaches. Systematic Biology, 58: 240-256.

Balaga, H.Y. 1975. Induction of branching in coconut. Philippine Journal of Biology, 4: 135-140

Banka, R., & Baker, W. J. 2004. A monograph of the genus Rhopaloblaste (Arecaceae). Kew Bulletin, 4760.

Barrau, J. 1959. The sago palms and other food plants of marsh dwellers in the South Pacific Islands. Economic Botany, 13: 151-162.

Barrett, H. C. 1973. Date breeding and improvement in North America. Fruit Variety Journal, 27: 50-55.

Barrett, C.F., Bacon, C.D., Antonelli, A., Cano, A., & Hoffman, T. 2016. An introduction to plant phylogenomics with a focus on palms. Botanical Journal of the Linnean Society.

Barrow, S. C. 1998. A monograph of Phoenix L. (palmac: Coryphoideae). Kew bulletin, 513-575.

Beccari, O. 1902. Systematic Enumeration of the Species of Calamus and Daemonorops: With Descriptions of the New Ones. US Government Printing Office.

Beccari, O. 1914. Asiatic Palms-Lepidocaryeae: The Species of Calamus. Bengal Secretariat Press.

Beentje, H. J. 1994a. A monograph of Ravenea (Palmae: Ceroxyloideae). Kew Bulletin, 623-671.

Beentje, H. 1994b. Ravenea in Madagascar. Principes, 38: 195-203.

Bell, A.D. 1979. The vascular pattern of a rhiomatous ginger (Alpinia speciosa L. Zingiberaccae). 2. The rhizome. Annals of Botany, 46: 213-220.

Bell, A. D., & Bryan, A. 2008. Plant form: an illustrated guide to flowering plant morphology. Timber Press.

Bell, A.D., & Tomlinson, P.B. 1980. Adaptive architecture in rhizomatous plants. Botanical Journal of the Linnean Society, 90: 125-160.

Bennett, B. C., & Hicklin, J. R. 1998. Uses of saw palmetto (Serenoa repens, Arecaceae) in Florida. Economic Botany, 52: 381-393.

Bernal, R., & Galeano, G. 2010. Notes on Mauritiella, Manicaria and Leopoldinia. Palms, 54.

Bernal, R., Galeano, G., & Henderson, A. 1991. Notes on Oenocarpus in the Colombian Amazon. Brittonia, 43: 154-164.

Bernal-Gonzalez, R., & Henderson, A. 1986. A new species of Socratea (Palmae) from Colombia with notes on the genus. Brittonia, 38: 55-59.

Borchsenius, F., & Bernal, R. (1996). Aiphanes (Palmae). Flora Ncotropica, 1-94.

Borchsenius, F., Pedersen, H. B., & Balslev, H. 1998. Manual to the palms of Ecuador.

Britt, A., & Dransfield, J. 2005. Dypsis dcliculata. PALMS, 44: 191-198.

Bullock, S. H. 1980. Demography of an undergrowth palm in littoral Cameroon. Biotropica, 12: 247-255.

Chazdon, R.L. 1991. Plant size and form in the understory palm genus Geonoma: Are species variations on a theme? American Journal of Botany, 78: 680-694.

Chazdon, R. L. 1992. Patterns of growth and reproduction of Geonoma congesta, a clustered understory palm. Biotropica, 24: 43-51.

Chevalier, A. 1952. Recherches sur les Phoenix africains. Revue internationale de botanique appliquee et d'agriculture tropicale, 32: 205-236.

Clark, D. B., & Clark, D. A, 1989. The role of physical damage in the seedling mortality regime of a neotropical rain forest. Oikos, 55: 225-230.

Clifford, H.T., Keighery, G.J., & Conran, J.G. 1998. Dasypogonaceac. In Dr. Klaus Kubitzki (Ed.), Flowering Plants. Monocots: Alismatanae and Commelinanae (pp. 190-194). Berlin: Springer-Verlag.

Couvreur, T. L., Kissling, W.D., Condamine, F.L., Svenning, J.C., Rowe, N.P. & Baker, J.W. 2014. Global diversification of a tropical plant growth form: environmental correlates and historical contingencies in climbing palms. Frontiers in genetics, 5: 452-452.

Davis, T. A. 1950. Branching in some Indian palms. Indian Coconut Journal, 3: 135-145.

de Granville, J. J., & Henderson, A. 1988. A new species of Asterogyne (Palmae) from French Guiana. Brittonia. 40: 76-80.

Doust, A. 2007. Architectural Evolution and its Implication for Domestication in Grasses. Annals of Botany, 100:941-950.

Doust, J.L & Doust, L.L. 1988. Plant Reproductive Ecology: Patterns and Strategies. New York: Oxford University Press.

Dowe, J. L. 2009. A taxonomic account of Livistona R. Br.(Arecaceae). Gard. Bull. Singapore, 60: 185-344.

Dowe, J. L. 2010. Australian palms: biogeography, ecology and systematics. CSIRO PUBLISHING.

Dowe, J. L., & Ferrero, M. D. 2001. Revision of Calyptrocalyx and the New Guinea species of Linospadix (Linospadicinae: Arecoideae: Arecaceae). Blumea-Biodiversity, Evolution and Biogeography of Plants, 46: 207-251.

Dowe, J. L., & Irvine, A. K. 1997. A revision of Linospadix in Australia, with the description of a new species. Principes, 41: 192-197.

Dransfield, J. 1977. Calamus caesius and Calamus trachycoleus compared. Gardens' Bulletin, 30: 75-78.

Dransfield, J. 1978. Growth forms of rain forest palms. Tropical trees as living systems, 247-268.

Dransfield, J. 1979. A manual of the rattans of the Malay Peninsula. A manual of the rattans of the Malay Peninsula, 29.

Dransfield, J. 1981. A synopsis of the genus Korthalsia (Palmae-Lepidocaryiodeae). Kew Bulletin, 36: 163-194.

Dransfield, J. 1982. Nomenclatural notes on Laccosperma and Ancistrophyllum (Palmae: Lepidocaiyoideae). Kew bulletin, 455-457.

Dransfield, J. 1984a. Note on the genus Zalaccella (Palmae: Lepidocaryoideae)[Calamus harmandii]. Kew bulletin.

Dransfield, J. 1984b. The genus Areca (Palmae: Arecoideae) in Borneo. Kew bulletin, 1-22.

Dransfield, J. (1989) Voanioala (Arecoideae: Cocoeae: Butiinae), a New Palm Genus from Madagascar. Kew Bulletin 44 (2):191-198.

Dransfield, J. 1992. The rattans of Sarawak. Richmond: Royal Botanic Gardens Kew.

Dransfield, J. 1997. The rattans of Bruenei Darussalam. Brunei Darussalam: Ministry of Industry and Primary Resources.

Dransfield, J. 2003. Dypsis turkii. PALMS-LAWRENCE, 47: 26-30.

Dransfield, J., & Beentje, H. 1995. The palms of Madagascar. Royal Botanic Gardens.

Dransfield, J., Lee, S. K., & Wei, F. N. 1985. Guihaia, a new coryphoid genus from China and Vietnam. Principes, 29: 3-12.

Dransfield, J., Uhl, N. W., Asmussen, C. B., Baker, W. J., Harley, M. M., & Lewis, C. E. 2005. A new phylogenetic classification of the palm family, Arecaceae. Kew Bulletin, 559-569.

Dransfield, J., Rakotoarinivo, M., Baker, W.J., Bayton, R.P., Fisher, J.B., Horn, J.W., & Metz, X., 2008a. A new Coryphoid palm genus from Madagascar. Botanical Journal of the Linnean Society, 156: 79-91.

Dransfield, J., Uhl, N., Asmussen, C.B., Baker, W.J., Harley, M.M., and Lewis, C.E. 2008b. Genera Palmarum: The evolution and classification of palms. Richmond: Royal Botanic Gardens Kew.

Duncan, W.G. 1971. Leaf angles, leaf area, and canopy photosynthesis. Crop Science, 11: 482-485.

Erwin, D.M. & Stockey, R.A., 1994. Premineralized monocotyledons from the middle Eocene Princeton chert of British Columbia, Canada: Arecaceae. Palacontographica, 234: 19-40.

Essig, F. 1977. A systematic histological study of palm. I. The Ptychosperma alliance. Systematic Botany, 2: 151-168.

Essig, F. B, 1978. A revision of the genus Ptychosperma Labill.(Arecaceae). Allertonia, 1: 415-478.

Essig, F. B., Manka, T. J., & Bussard, L. 1999. A systematic histological study of palm fruits. III. Subtribe Iguanurinae (Arecaceae). Brittonia, 51: 307-325.

Evans, T. Sengdala, K., Viengkham, O.V., Thammavong, B., & Dransfield. J. 2000. Four new species of Calamus (Arecaceae: Calamoideae) from Laos and Thailand. Kew Bulletin, 55:929

Faurby, S., Eiserhardt, W.L., Baker, W.J., & Svenning, J. 2016. An all-evidence species-level supertree for the palms (Arecaceae). Molecular Phylogenetics and Evolution, 100: 57-69.

Fernando, E. S. 1983. A revision of the genus Nenga. Principes, 27: 55-70.

Fernando, E. S. 1990. The genus Heterospathe (Palmae: Arecoideae) in the Philippines. Kew bulletin, 219-234.

Fisher, J.B. 1973. Unusual branch development in the palm, Chrysalidocarpus. Botanical Journal of the Linnean Society, 66: 83-95.

Fisher, J.B. 1974. Axillary and dichotomous branching in the palm Chamaedorea. American Journal of Botany, 61: 1046-1051.

Fisher, J.B. 1976. Development of dichotomous branching and axillary buds in Strelitzia (Monocotyledonae). Canadian Journal of Botany, 54: 578-592.

Fisher, J. B., & Dransfield, J. 1977. Comparative morphology and development of inflorescence adnation in rattan palms. Botanical Journal of the Linnean Society, 75: 119-140.

Fisher, J. B., & Dransfield, J. 1979. Development of axillary and leaf-opposed buds in rattan palms. Annals of Botany, 44: 57-66.

Fisher, J.B., & Jayachandran, K. 1999. Root structure and arbuscular mycorrhizal colonization of the palm Serenoa repens under field conditions. Plant and Soil, 217: 229-241.

Fisher, J. B. and Maidman, K.J. 1999. Branching and architecture in palms: value for systematics. Pages 35-46. In A. Henderson & F. Borchsenius editor. Evolution, variation, and classification of palms. Memoirs of the New York Botanical Garden 83. New York Botanical Garden Press, Bronx.

Fisher, J. B., & Mogea, J. P. 1980. Intrapetiolar inflorescence buds in Salacca (Palmae): development and significance. Botanical Journal of the Linnean Society, 81: 47-59.

Fisher, J.B., & Tomlinson, P.B. 1973. Branch and inflorescence production in saw palmetto (Serenoa repens). Principes, 17: 10-19.

Fisher, J.B., & Zona, S. 2006. Unusual branching in Manicaria. Palms, 50: 99-102.

Fisher, J.B., Goh, C.J., & Rao, A.N. 1989. Non-axillary branching in the palms Eugeissona and Oncosperma (Arecaceae). Botanical Journal of the Linnaean Society, 99: 347-363.

Furtado, C.X. 1949. The Malayan species of Salacca. Garden Bulletin Singapore, 12: 378-403.

Gaiero, P., Mazzella, C., Agostini, G., Bertolazzi, S., & Rossato, M. 2011. Genetic diversity among endangered Uruguayan populations of Butia Becc. species based on ISSR. Plant systematics and evolution. 292: 105-116.

Galeano, G., & Bernal, R. 2013. Sabinaria, a new genus of palms (Cryosophileae, Coryphoideae, Arecaceae) from the Colombia-Panama border. Phytotaxa, 144: 27-44.

Gee, C.T. 2001. The mangrove palm Nypa in the geologic past of the New World. Wetlands Ecology and Management, 9: 181-194.

Gianoli, E. 2004. Evolution of climbing habit promotes diversification in flowering plants. Proceedings of the Royal Society of London, 271: 2011-2015.

Gola, E.M, 2014. Dichotomous branching: the plant form and integrity upon the apical meristem bifurcation. Frontiers in Plant Science, 5: 1-6.

Govaerts, R., Dransfield, J., Zona, S.F., Hodel, D.R. & Henderson, A. 2011. World Checklist of Arecaceae. Facilitated by the Royal Botanic Gardens, Kew. Published on the Internet; http://apps.kew.org/wesp/ Retrieved 2011-01-22.

Greuter, W., McNeill, J., Barrie, F.R., Burdet, H.M., Demoulin, V., Filgueiras, & Hawksworth, D.L. 1999. International code of botanical nomenclature (Saint Louis Code). St Louis, Missouri, USA: Sixteenth International Botanical Congress.

Halle, F., Oldeman, R.A.A., & Tomlinson, P.B. 1978. Tropical Trees and Forests: An Architectural Analysis. Berlin: Springer-Verlag.

Harley, M. M. 2006. A summary of fossil records for Arecaceae. Botanical Journal of the Linnean Society, 151: 39-67.

Heatubun, C. D. 2011. Seven new species of Areca (Arecaceae). Phytotaxa, 28: 6-26.

Heatubun, C. D., Baker, W. J., Mogea, J. P., Harley, M. M., Tjitrosoedirdjo, S. S., & Dransfield, J. 2009. A monograph of Cyrtostachys (Arecaceae). Kew Bulletin, 64: 67-94.

Heatubun, C. D., Dransfield, J., Flynn, T., Tjitrosoedirdjo, S. S., Mogea, J. P.,, & Baker, W. J. 2012. A monograph of the betel nut palms (Areca: Arecaceae) of East Malesia. Botanical Journal of the Linnean Society, 168: 147-173.

Heatubun, C. D., Zona, S., & Baker, W. J. 2014. Three new genera of arecoid palm (Arecaceae) from eastern Malesia. Kew Bulletin, 69: 1-18.

Henderson, A. 1990. Arecaceae. Part I. Introduction and the Iriarteinae. Flora Neotropica, 1-100.

Henderson, A. 1995. The palms of the Amazon. Oxford University Press.

Henderson, A. 2000. Bactris (Palmae). Organization for Flora Neotropica.

Henderson, A. 2002a. Evolution and Ecology of palms. New York: New York Botanical Garden Press.

Henderson, A. J. 2002b. Phenetic and phylogenetic analysis of Reinhardtia (Palmae). American Journal of Botany, 89: 1491-1502.

Henderson, A. 2005. A new species of Calamus (Palmae) from Taiwan. TAIWANIA-TAIPEI, 50: 222.

Henderson, A. 2009. Palms of southern Asia. New Jersey: Princeton University Press.

Henderson, A. 2011a. A revision of Geonoma (Arecaceae). Phytotaxa, 17: 1-271.

Henderson, A. 2011b. A revision of Leopoldinia (Arecaceae). Phytotaxa, 32: 1-17.

Henderson, A, J., & Bacon, C. D. 2011. Lanonia (Arecaceae: Palmae), a new genus from Asia, with a revision of the species. Systematic Botany, 36: 883-895.

Henderson, A., & Dung, N. Q. 2013. Four new species of Calamus (Arecaceae) from Vietnam. Phytotaxa, 135: 19-26.

Henderson, A., & Galeano, G. 1996. Euterpe, Prestoea, and Neonicholsonia (Palmae). Flora Neotropica, 189.

Henderson, A., & Henderson, F. 2007. New species of Calamus (Palmae) from Lao and Myanmar. Taiwania, 52: 152-158.

Henderson, A, & de Nevers, G. 1988. Prestoea (Palmae) in Central America. Annals of the Missouri Botanical Gardens, 75: 203-217.

Henderson, A., & Steyermark, J. A. 1986. New palms from Venezuela. Brittonia, 38: 309-313.

Henderson, A., Galeano-Garces, G., & Bernal, R. 1997. Field Guide to the palms of America New Jersey: Princeton University Press.

Henderson, A., Ban, N. K., & Dung, N. Q. 2008. New species of Calamus (Palmae) from Vietnam. Palms, 52: 187-197.

Hodel, D. R. 1992. Chamaedorea palms: The species and their cultivation. International Palm Society.

Hodel, D. R., Marcus, J., & Dransfield, J. 2005. Dyspis robusta, a large new palm from cultivation. PALMS-LAWRENCE: 49: 128.

Holtuum, R.E., 1955. Growth-habits of monocotyledons-variation on a theme. Phytomorphology, 5: 399-413.

Isnard, S., Speck, T., & Rowe, N. P. 2005. Biomechanics and development of the climbing habit in two species of the South American palm genus Desmoncus (Arecaceae). American Journal of Botany, 92: 1444-1456.

Jaffre, T., & Veillon, J. M. 1989. Morphology, distribution and ecology of palms in New Caledonia. DO WE JL (ed.), Palms of the South West Pacific. Palm and Cycas Societies of Australia, Milton, 158-169.

James, A. 2013. Adaptations of an understory Geonoma. Palms, 57: 109-112.

Jeanson, M. L., & Guo, L. 2011. Arenga longicarpa, a poorly known wpecies from South China. Palms Journal of the International Palm Society, 55: 122.

Jenik, J. 1994. Clonal growth in woody plants: A review. Folia Geobotanica, 29: 291-306.

Kahn, F. 2008. El genero Astrocaryum (Arecaceae). Revista peruana de biologia, 15: 31-48.

Kahn, E, & De Granville, J. J. 1998. Astrocaryum minus, rediscovered in French Guiana. Principes. 42: 171-178.

Kahn, F., & Mejia, K. 1988. A new species of Chelyocarpus (Palmae, Coryphoideae) from Peruvian Amazonia. Principes (USA).

Kahn, F., & Millan, B. 1992. Asfrocarynm (Palmae) in Amazonia: a preliminary treatment Bulletin de l'Institut francais d'etudes andines, 21: 459-531.

Kiew, R. 1976. The genus Iguanura Bl. (Palmae). Gard. Bull. Singapore, 28: 191-226.

Kramadibrata, P. 1992. A revision of the genus Calamus (Palmae) section Macropodus sensu Furtado (Doctoral dissertation. University of Reading).

Maddison, W. P., & Maddison D. R. 2011. Mesquite: a modular system for evolutionary analysis. Version 2.75. http://mesquiteproject.org. Accessed 1 Oct 2013.

McClatchey, W. 1998. A new species of AfetroxyZon (Arecaceae) from Western Samoa. Novon, 252-258.

McSteen, P., & Leyser, O. 2005. Shoot branching. Annual Review of Plant Biology, 56:353-374.

Mehrotra, R.C., Tiwari, R.P., & Mazumder, B.I. 2003. Nypa megafossils from the Tertiary sediments of Northeast India. Gcobios, 36: 83-92.

Mendoza, A., & Franco, M. 1998. Sexual reproduction and clonal growth in Reinhardtia gracilis (Palmae). an understory tropical palm. American Journal of Botany, 84: 521-527.

Mogie, M. 1992. The evolution of asexual reproduction in plants. London: Chapman & Hall.

Moore, Jr H. E. 1978. Tectiphiala, a new genus of Palmae from Mauritius. Gcntes Herb.(Ithaca), 11: 284-290.

Moore, H. E., & Uhl, N. W. 1982. Major trends of evolution in palms. The Botanical Review, 48: 1-69.

Moore, H. E., & Uhl, N. W. 1984. The indigenous palms of New Caledonia. Allertonia, 3: 313-402.

Moya, C. E. 1997a. Considerations on Ecology and Distribution of Cuban Copemicia. In II International Symposium on Ornamental Palms & other Monocots from the Tropics 486 (pp. 149-154).

Moya, C. E, 1997b. The Cuban ornamental palms. In II International Symposium on Ornamental Palms & other Monocots from the Tropics 486 (pp. 53-58).

Noblick, L. R. 1996. Syagrus. Palm Journal. 126: 12-46.

Noblick, L. R. 2004. Syagrus cearensis, a twin-stemmed new palm from Brazil. Palms, 48: 70-76.

Noblick, L. R., & Lorenzi, H. 2010. New Syagrus species from Brazil. Palms, 54: 18.

Noblick, L.R. & Meerow, A.W. 2015. The transfer of the genus Lytocaryum to Syagrus. Palms, 59: 57-62.

Noblick, L. R., Lorenzi, H., & Souza, V. C. 2014. Four new taxa of acaulescent Syagrus (Arecaceae) from Brazil. Phytotaxa, 188: 1-13.

Pinatuad, J.C., & Millan, B. 2004. Vegetative transformation of inflorescences in Socratea salazarii. Palms, 48: 86-89.

Pinheiro, C. U. B,, Balick, M. J., & Frazao, J. M. F. 1996. Branching in Syagrus cocoides (Arecaceae) in Maranhao, northeastern Brazil. Brittonia, 48: 556.

Pintaud, J. C., & Baker, W. J. 2008. A revision of the palm genera (Arecaceae) of New Caledonia. Kew Bulletin, 63: 61-73.

Pintaud, J. C., & Stauffer, F. W. 2015. A revision of the large-flowered group of Basselinia Vieill. sect. Taloua HE Moore & Uhl (Arecaceae).

Putz, F. E. 1990. Growth habits and trellis requirements ofe climbing palms (Calamus spp.) in north-eastern Queensland. Australian Journal of Botany. 3i8: 603-608.

Rakotoarinivo, M. 2008. A remarkable Ravenea from the montane forest of Andilamena. Madagascar. Palms, 52: 11-17.

Rakotoarinivo, M., & Dransfield, J. 2010. New species of Dypsis and Ravenea (Arecaceae) from Madagascar. Kew Bulletin, 65: 279-303.

Rakotoarinivo, M., Trudgen, M., & Baker, W. 2009. The palms of the Makira protected area, Madagascar. PALMS 53(3):125-146.

Renuka, C., Bhat, K. M., Dliamodaran, T. K., Muraleedharan, P. K., Mohanan, C., & Seethalakshmi, K. K. 2001. Rattans in India: Status and opportunities. In Tropical Forestry Research: Challenges in the New Millennium: Proceedings of the International Symposium, 2-4 August, 2000, Peechi, India (p. 241). Kerala Forest Research Institute.

Rodrigues, A.C., & Maranho-Estelita, M.E. 2009. Morphoanatomy of the stem in Cyperaceae. Acta Botanica Brasilica, 23.

Rupert, N., Mansor, A., & Shah, S.A.M, 2012. New shoots from inflorescences in Calamus castaneus in Peninsular Malaysia. Palms, 56: 36-40.

Russell, T. A. 1965. The Raphia palms of West Africa. Kew Bulletin, 173-196.

Rustiami, H. 2002a. A new species of Daemonorops section Piptospatha (Arecaceae) from Siberut island, West Sumatra. Kew bulletin, 729-733.

Rustiami, H, 2002b. Two New Species of Daemonorops (Palmae) from Sumatra. Gard. Bull. Singapore, 54: 199-204.

Rustiami, H., Dransfield, J., & Fernando, E. S. 2014. Daemonorops sedisspirituum, a new species of Daemonorops Blume (Arecaceae: Calamoideae) from Java. Kew Bulletin, 69: 1-4.

Schluter, D. 2001. Ecology and the origin of species. Trends in Ecology & Evolution. 16:372-380.

Skov, F., & Balslev, H. 1989. A revision of Hyospathe (Arecaceae). Nordic Journal of Botany, 9: 189-202.

Stauffer, F. W., Asmussen, C. B., Henderson, A., & Endress, P. K. 2003. A revision of Asterogyne (Arecaceae: Arecoideae: Geonomcae). Brittonia, 55: 326-356.

Sunderland, T. C. 2012. A taxonomic revision of the rattans of Africa (Arecaceae: Calamoideae). Phytotaxa, 51: 1-76.

Svenning, J. C., & Balslev, H. 1998. The palm flora of the Maquipucuna montane forest reserve, Ecuador. Principes, 42: 218-226.

Takenaka, A,, Takahashi, K., & Kohyama, T. 2001. Optimal leaf display and biomass partitioning for efficient light capture in an understorey palm. Licuala arbuscula. Functional Ecology, 15: 660-668.

Tillich, H. J. 1998. Development and organization. In Flowering Plants' Monocotyledons (pp. 1-19). Springer Berlin Heidelberg.

Tisserat, B., & DeMason, D. A. 1985. Occurrence and Histological Structure of Offshoots and Inflorescences Produced from Phoenix dactylifera L. Plantlets In vitro. Bulletin of the Torrey Botanical Club, 112: 35-42.

Tomlinson, P.B. 1961. Essays on the morphology of palms: The habit of palms. Principes, 5: 83-89.

Tomlinson, P. B. 1967. Dichotomous branching in Allagoptera. Principes, 11: 70-72.

Tomlinson, P. B. 1971. The shoot apex and its dichotomous branching in the Nypa palm. Annals of Botany, 35: 865-879.

Tomlinson, P.B. 1973. Branching in Monocotyledons. The Quarterly Review of Biology, 48: 458-466.

Tomlinson, P. B., & Zimmermann, M. H. 2003. Stem vascular architecture in the American climbing palm

Desmoncus (Arecaceae-Arecoideae-Bactridinae). Botanical Journal of the Linnean Society, 142: 243-254. Tomlison, P.B. 1990. The structural biology of palms. Oxford: Oxford University Press.

Tomlison, P.B. & Moore Jr., H.E. 1966. Dichotomous branching in palms? Principes, 10: 21-29.

Tomlison, P.B. & Moore Jr., H.E. 1968. Inflorescence in Nannorhops ritchiana (Palmae). Journal of the Arnold Arboretum, 49: 16-34.

Van der Pijl, L. 1982. Principles of dispersal. Berlin: Springer-Verlag.

Van Valkenburg, J. L. C. H., & Dransfield, J. 2004. Hyphaene guineensis. Palms. 48: 10-16.

Van Valkenburg, J. L. C. H., Sunderland, T. C. H., Banak, L. N., & Issembe, Y. 2007. Sclerosperma and Podococcus in Gabon.

Van Valkenburg, J. L. C. H., Sunderland, T. C. H., & Couvreur, T. L. P. 2008. A revision of the genus Sclerosperma (Arecaceae). Kew Bulletin, 63: 75-86.

Ward, S., & Leyser, O. 2004. Shoot branching. Current Opinion in Plant Biology, 1:73-78.

Wikstrom N., Savolainen V., & Chase M. W. 2001. Evolution of the angiosperms: Calibrating the family tree. Proceedings of the Royal Society of London. Biological Sciences, 268: 2211-2220.

Zimmermann, M. H., & Tomlinson, P. B. 1967. Anatomy of the palm Rhapis excelsa, IV. Vascular development in apex of vegetative aerial axis and rhizome, Journal of the Arnold Arboretum, 48: 122-142.

Zona, S. 1999. Revision of Drymophloeus (Areceacca: Arecoideae). Blumea-Biodiversity, Evolution and Biogeography of Plants, 44: 1-24.

Appendix
Table 4 Genera, species counts and references for the five
branching types and their combinations for subfamily Calamoideae.
Number under species counts give specics reviewed or
examined/number of species in genus

A.                Species    Number of specics that exhibit...
                  count
Genus                        No          Lateral
                             branching   axillary

Calamus           263/521    40          210

Eleiodoxa         1/1                    1

Eremospatha       11/11                  11

Eugeissona        6/6

Korthalsia        28/28

Laccosperma       6/6                    6

Lepidocaryum      1/1                    1

Mauritia          2/2        2

Mauritiella       4/4                    4

Metroxylon        7/7        6           1

Myrialepis        1/1

Oncocalamus       5/5                    5

Pigafetta         2/2        2

Plectocomia       15/15                  15

Plectocomiopsis   6/6                    6

Ruphia            19/20      8           11

Salacca           22/23                  21

Calamoideae       399/659    58          292

A.                Number of specics that exhibit...

Genus             Shoot apical   Shoot apical       FALSE
                  dichotomy      dichotomy with     vivipary
                                 lateral axillary

Calamus

Eleiodoxa

Eremospatha

Eugeissona        6

Korthalsia                       28 *

Laccosperma

Lepidocaryum

Mauritia

Mauritiella

Metroxylon

Myrialepis

Oncocalamus

Pigafetta

Plectocomia

Plectocomiopsis

Ruphia

Salacca                                             2

Calamoideae       6              28                 2

A.                Number of specics that exhibit...

Genus             False vivipary   Abaxial   Leaf-
                  with lateral               opposed
                  axillary

Calamus           2                          6

Eleiodoxa

Eremospatha

Eugeissona

Korthalsia

Laccosperma

Lepidocaryum

Mauritia

Mauritiella

Metroxylon

Myrialepis                                   1

Oncocalamus

Pigafetta

Plectocomia

Plectocomiopsis

Ruphia

Salacca

Calamoideae       2                          7

A.                References

Genus

Calamus           Beccari, 1902; Beccari, 1914;
                  Dransfield, 1977; Dransfield, 1979;
                  Fisher & Dransfield. 1979;
                  Dransfield, 1982; Dransfield, 1984a;
                  Kramadibrata, 1992; Dransfield, 1997;
                  Evans et al., 2000; Rcnuka et al., 2001;
                  Baker & Dransfield, 2002a; Baker &
                  Dransfield, 2002b; Rustiami, 2002a;
                  Rustiami, 2002b; Baker et al., 2003;
                  Dransfield et al., 2005;
                  Henderson, 2005; Baker &
                  Dransfield; 2007; Henderson &
                  Henderson, 2007; Henderson et al.,
                  2008; Henderson, 2009;
                  Sunderland, 2012; Henderson & Dung,
                  2013; Rustiami et al., 2014;
                  Hcatubun et al., 2014; Baker &
                  Dransfield, 2014

Eleiodoxa         Dransfield et al., 2008a. b

Eremospatha       Dransfield et al., 2008a, b

Eugeissona        Fisher etal., 1989;

Korthalsia        Dransfield, 1981; Fisher &
                  Dransfield, 1979

Laccosperma       Dransfield et al., 2008a, b

Lepidocaryum      Dransfield et al., 2008a, b

Mauritia          Dransfield et al., 2008a, b

Mauritiella       Bernal & Galeano, 2010

Metroxylon        Barrau. 1959; McClatchcy, 1998

Myrialepis        Dransfield, 1982

Oncocalamus       Dransfield et al., 2008a, b

Pigafetta         Dransfield et al., 2008a, b

Plectocomia       Dransfield, 1982

Plectocomiopsis   Dransfield et al., 2008a, b

Ruphia            Russell. 1965; Fisher et al., 1989

Salacca           Fisher & Mogea, 1980

Calamoideae

Table 5 Genera, species counts and references for
the five branching types and their combinations for
subfamily Copryphoideae. Number under species give
species reviewed or examined/number of species in genus

B.                    Species   Number of species that exhibit...
                      count
Genus                           No          Lateral
                                branching   axillary

Acoelorrhaphe         1/1                   1

Arenga                20/24     3           17

Bismarkia             1/1       1

Borassodendron        2/2       2

Borassus              5/5       5

Brahea                11/11     10          1

Caryota               14/14     11          3

Chamaerops            1/1       1

Chelyocarpus          4/4       2           2

Chuniophoenix         2/2                   2

Coccothrinax          50/53     46          4

Colpothrinax          3/3       3

Copemicia             22/22     21          1

Corypha               5/5       5

Cryosophila           10/10     10

Guihaia               2/2                   2

Hemithrinax           3/3       3

Hyphaene              7/8       2

Itaya                 1/1       1

Johannesteijsmannia   4/4       4

Kerriodoxa            1/1       1

Lanonia               8/8       1           7

Latania               3/3       3

Leucothrinax          1/1       1

Licuala               60/162    38          22

Livistona             27/27     27

Lodoicea              1/1       1

Maxburretia           2/3                   2

Medemia               1/1       1

Nannorhops            1/1

Phoenix               13/13     6           7

Pholidocarpus         6/6       6

Pritchardia           30/30     30

Pritchardiopsis       1/1       1

Rhapidophyllum        1/1                   1

Rhapis                10/10                 10

Sabal                 14/14     14

Sabinaria             1/1       1

Saribus               1/1       1

Satranala             1/1       1

Schippia              1/1       1

Serenoa               1/1                   1

Tahina                1/1       1

Thrinax               3/3       3

Trachvcarpus          10/10     10

Trithrinax            3/3       2           1

Wallchia              8/8       1           7

Washingtonia          2/2       2

B.                    Number of species that exhibit...

Genus                 Shoot       Shoot apical   FALSE
                      apical      dichotomy      vivipary
                      dichotomy   with lateral
                                  axillary

Acoelorrhaphe

Arenga

Bismarkia

Borassodendron

Borassus

Brahea

Caryota

Chamaerops

Chelyocarpus

Chuniophoenix

Coccothrinax

Colpothrinax

Copemicia

Corypha

Cryosophila

Guihaia

Hemithrinax

Hyphaene              3           2

Itaya

Johannesteijsmannia

Kerriodoxa

Lanonia

Latania

Leucothrinax

Licuala

Livistona

Lodoicea

Maxburretia

Medemia

Nannorhops                        1

Phoenix

Pholidocarpus

Pritchardia

Pritchardiopsis

Rhapidophyllum

Rhapis

Sabal

Sabinaria

Saribus

Satranala

Schippia

Serenoa

Tahina

Thrinax

Trachvcarpus

Trithrinax

Wallchia

Washingtonia

B.                    References

Genus

Acoelorrhaphe         Personal observation

Arenga                Dransfield et al., 2008a, b;
                      Jeanson & Guo, 2011

Bismarkia             Dransfield et al., 2008a, b

Borassodendron        Dransfield et al., 2008a, b

Borassus              Dransfield et al., 2008a, b

Brahea                Dransfield et al., 2008a, b

Caryota               Dransfield et al., 2008a, b;
                      Personal observation

Chamaerops            Dransfield et al., 2008a, b:
                      personal observation

Chelyocarpus          Kahn & Mejia.1988;
                      Dransfield et al., 2008a, b

Chuniophoenix         Dransfield et al., 2008a, b;
                      Personal observation

Coccothrinax          Henderson et al., 1997;
                      Henderson, 2005; Moya, 1997b

Colpothrinax          Dransfield et al., 2008a. b;
                      Personal observation

Copemicia             Henderson et al., 1997; Moya, 1997a

Corypha               Dransfield et al., 2008a, b;
                      Personal observation

Cryosophila           Dransfield et al., 2008a, b;
                      Personal observation

Guihaia               Dransfield et al., 1985;
                      Dransfield et al., 2008a, b;

Hemithrinax           Dransfield et al., 2008a, b

Hyphaene              Moore & Uhl, 1982;
                      van Valkenburg & Dransfield, 2004

Itaya                 Dransfield et al., 2008a, b;
                      personal observation

Johannesteijsmannia   Dransfield et al., 2008a, b

Kerriodoxa            Dransfield et al., 2008a. b;
                      Personal observation

Lanonia               Henderson & Bacon, 2011

Latania               Dransfield et al., 2008a, b;
                      Personal observation

Leucothrinax          Dransfield et al., 2008a, b;
                      Personal observation

Licuala               Henderson et al., 1997;
                      Takcnaka et al., 2001;
                      Dransfield et al., 2008a, b;
                      Henderson et al., 2008;

Livistona             Dransfield et al., 2008a, b;
                      Dowe, 2009

Lodoicea              Dransfield et al., 2008a, b

Maxburretia           Dransfield et al., 2008a, b;
                      Henderson, 2009

Medemia               Dransfield et al., 2008a, b

Nannorhops            Tomlison & Moore Jr., 1968

Phoenix               Davis, 1950; Chevalier, 1952;
                      Barrow, 1998; Dransfield et al.,
                      2008a, b; Barrett, 1973

Pholidocarpus         Dransfield et al., 2008a, b

Pritchardia           Dransfield et al., 2008a, b;
                      Personal observation

Pritchardiopsis       Dransfield et al., 2008a, b

Rhapidophyllum        Dransfield et al., 2008a, b;
                      Personal observation

Rhapis                Dransfield et al., 2008a, b;
                      Personal observation

Sabal                 Dransfield et al., 2008a, b

Sabinaria             Dransfield et al., 2008a, b

Saribus               Bacon & Baker, 2011

Satranala             Dransfield et al., 2008a. b

Schippia              Dransfield et al., 2008a, b

Serenoa               Fisher & Tomlison, 1973;
                      Bennet & Hicklin, 1998;
                      Abrahamson, 1999;
                      Personal observation

Tahina                Dransfield et al., 2008a, b

Thrinax               Dransfield et al., 2008a, b

Trachvcarpus          Dransfield et al., 2008a, b

Trithrinax            Dransfield et al., 2008a, b;
                      Personal observation

Wallchia              Dransfield et al., 2008a, b;
                      Personal observation

Washingtonia          Henderson et al., 1997;
                      Dransfield et al., 2008a, b

Table 6 Genera, species counts and references for
the four branching types and their combinations for
subfamily Ceroxyloideae. Number under species counts
give species reviewed or examined/number of species in genus.

C.              Species   Number of species that exhibit...
                count
Genus                     No          Lateral
                          branching   axillary

Ammandra        1/1       1

Aphandra        1/1       1

Ceroxylon       12/12     12

Juania          1/1       1

Oraniopsis      1/1       1

Phytelephas     6/6       4           2

Pseudophoenix   4/4       4

Ravenea         21/21     20          1
Ceroxyloideae   47/47     46          1

C.              Number of species that exhibit...

Genus           Shoot apical   Shoot apical       FALSE
                dichotomy      dichotomy with     vivipary
                               lateral axillary

Ammandra

Aphandra

Ceroxylon

Juania

Oraniopsis

Phytelephas

Pseudophoenix

Ravenea
Ceroxyloideae

C.              Number of species that exhibit...

Genus           False vivipary   Abaxial   Leaf-opposed
                with lateral
                axillary

Ammandra

Aphandra

Ceroxylon

Juania

Oraniopsis

Phytelephas

Pseudophoenix

Ravenea
Ceroxyloideae

C.              References

Genus

Ammandra        Dransfield et al.. 2008a, 2008b

Aphandra        Dransfield et al., 2008a, 2008b

Ceroxylon       Dransfield et al.. 2008a, 2008b

Juania          Dransfield et al., 2008a, 2008b

Oraniopsis      Dransfield et al., 2008a, 2008b

Phytelephas     Dransfield et al.. 2008a, 2008b

Pseudophoenix   Dransfield et al.. 2008a, 2008b

Ravenea         Beentje, 1994a; Beentje. 1994b:
Ceroxyloideae   Dransfield et al., 2008a, 2008b;
                Rakotoarinivo, 2008;

Table 7 Genera, species counts references for the four
branching types and their combinations for subfamily Arecoideae
(A. Acanthophoenix-Beccariophoenix, B. Bentinckia--Drymophloeus,
C. Dypsis--Leopoldinia, D. Lepidorhachis-Prestoea,
E. Ptychococcus--Voaniola, F. Wallaceodoxa--Bodyetia.
Number under species counts give species reviewed or
examined/number of species in genus

D.                Species     Number of species that exhibit...
                  count
Genus                         No branching   Lateral axillary

Acanlhophoenix                3

Acrocomia         8/8         8

Actinokentia      2/2         2

Actinorhytis      1/1         1

Adonidia          1/1         1

Aiphanes          23/29       11             12

Allagoptera       5/5         1

Archontophoenix   6/6         6

Areca             37/46       24             13

Asterogyne        5/5         5

Astrocuryum       32/38       24             8

Attalea           66/66       66

Bactris           72/79       5              67

Balaka            9/9         9

Barcella          1/1         1

Basselinia        14/14       10             3

Beccariophoenix   3/3         1

Bentinckia        2/2         2

Brassiophoenix    2/2         2

Bwretiokentia     5/5         5

Butia             17/20       14             3

Calyptrocaix      21/26       12             9

Calyptrogyne      10/17       10

Calyptronoma      3/3         3

Carpentaria       1/1         1

Carpoxvlon        1/1         1

Chamaedorea       91/104      73             17

Chambeyronia      2/2         2

Clinosperma       4/4         4

Clinostigma       11/11       11

Cocos             1/1         1

Cyphokentia       2/2         1              1

Cyphophoenix      4/4         4

Cyphosperma       5/5         5

Cyrtostachys      5/7         1              4

Deckenia          1/1         1

Desmoncus         24/24                      24

Dictyocaryum      3/3         3

Dictyospertna     1/1         1

Dransfieldia      1/1                        1

Drymophloeus      5/7         5

Dypsis            160/167     63             90

Elaeis            2/2         2

Euterpe           7/7         1              6

Gaussia           5/5         5

Geonoma           39/68       14             25

Hedyscepe         l/l         1

Hetemspathe       22/41       18             4

Howea             2/2         2

Hydriastele       30/49       16             14

Hvophorbe         5/5         5

Hyospalhe         2/5                        2

Iguanura          25/33       15             10

Irartea           1/1         1

Iriartella        2/2                        2

Jailoloa          1/1         1

Jubaea            1/1         1

Jubaeopsis        1/1                        1

Kentiopsis        4/4         4

Laccospadix       1/1                        1

Lemurophoenix     1/1         1

Leopoldinia       2/2         2

Lepidorrhachis    1/1         1

Linospadix        7/7         1              6

Loxococcus        1/1         1

Lytocaryum        4/4         4

Manicaria         2/2         1

Manjekia          1/1         1

Marojejya         2/2         2

Masoala           2/2         2

Nenga             4/5         1              3

Neonicholsonia    1/1         1

Neoveitchia       2/2         2

Nephrosperma      1/1         1

Normanbya         1/1         1

Oertocarpus       9/9         8              1

Oncosperma        6/6

Orania            18/18       18

Parajubaea        3/3         3

Pelagodoxa        1/1         1

Phoenicophorium   1/1         1

Pholidostachys    4/8         4

Physokentia       7/7         7

Pinanga           60/139      12             48

Podococcus        2/2                        2

Ponapea           4/4         4

Prestoea          10/10       2              8

Ptychococcus      2/2         3

Ptychosperma      19/29       7              12

Reihardtia        6/6         1              5

Rhopaloblaste     6/6         5              1

Rhopalostylis     2/2         2

Roscheria         1/1         1

Roystonea         10/10       1

Satakentia        1/1         1

Sclerosperma      3/3                        3

Socratea          5/5         4

Solfia            1/1         1

Sommieria         1/1         1

Syagrus           36/61       23             12

Synechanthus      2/2         1              1

Tectiphiala       1/1                        1

Veitchia          11/11       11

Verschaffeltia    1/1         1

Voaniola          1/1         1

Wallaceodoxa      l/l         1

Welfia            1/1         1

Wendlandiella     1/1                        1

Weltinia          21/21       19             2

Wodvetia          1/1         1

Arecoideae        1112/1376   657            423

D.                Number of species that exhibit...

Genus             Shoot apical   Shoot apical       False vivipary
                  dichotomy      dichotomy with
                                 lateral axillary

Acanlhophoenix

Acrocomia

Actinokentia

Actinorhytis

Adonidia

Aiphanes

Allagoptera       4

Archontophoenix

Areca

Asterogyne

Astrocuryum

Attalea

Bactris

Balaka

Barcella

Basselinia                       1

Beccariophoenix

Bentinckia

Brassiophoenix

Bwretiokentia

Butia

Calyptrocaix

Calyptrogyne

Calyptronoma

Carpentaria

Carpoxvlon

Chamaedorea       1

Chambeyronia

Clinosperma

Clinostigma

Cocos

Cyphokentia

Cyphophoenix

Cyphosperma

Cyrtostachys

Deckenia

Desmoncus

Dictyocaryum

Dictyospertna

Dransfieldia

Drymophloeus

Dypsis            6

Elaeis

Euterpe

Gaussia

Geonoma

Hedyscepe

Hetemspathe

Howea

Hydriastele

Hvophorbe

Hyospalhe

Iguanura

Irartea

Iriartella

Jailoloa

Jubaea

Jubaeopsis

Kentiopsis

Laccospadix

Lemurophoenix

Leopoldinia

Lepidorrhachis

Linospadix

Loxococcus

Lytocaryum

Manicaria         1

Manjekia

Marojejya

Masoala

Nenga

Neonicholsonia

Neoveitchia

Nephrosperma

Normanbya

Oertocarpus

Oncosperma

Orania

Parajubaea

Pelagodoxa

Phoenicophorium

Pholidostachys

Physokentia

Pinanga

Podococcus

Ponapea

Prestoea

Ptychococcus

Ptychosperma

Reihardtia

Rhopaloblaste

Rhopalostylis

Roscheria

Roystonea

Satakentia

Sclerosperma

Socratea

Solfia

Sommieria

Syagrus           1

Synechanthus

Tectiphiala

Veitchia

Verschaffeltia

Voaniola

Wallaceodoxa

Welfia

Wendlandiella

Weltinia

Wodvetia

Arecoideae        13             1

D.                Number of species that exhibit...

Genus             False vivipary   Abaxial   Leaf-opposed
                  with lateral
                  axillary

Acanlhophoenix

Acrocomia

Actinokentia

Actinorhytis

Adonidia

Aiphanes

Allagoptera

Archontophoenix

Areca

Asterogyne

Astrocuryum

Attalea

Bactris

Balaka

Barcella

Basselinia

Beccariophoenix

Bentinckia

Brassiophoenix

Bwretiokentia

Butia

Calyptrocaix

Calyptrogyne

Calyptronoma

Carpentaria

Carpoxvlon

Chamaedorea

Chambeyronia

Clinosperma

Clinostigma

Cocos

Cyphokentia

Cyphophoenix

Cyphosperma

Cyrtostachys

Deckenia

Desmoncus

Dictyocaryum

Dictyospertna

Dransfieldia

Drymophloeus

Dypsis                             1

Elaeis

Euterpe

Gaussia

Geonoma

Hedyscepe

Hetemspathe

Howea

Hydriastele

Hvophorbe

Hyospalhe

Iguanura

Irartea

Iriartella

Jailoloa

Jubaea

Jubaeopsis

Kentiopsis

Laccospadix

Lemurophoenix

Leopoldinia

Lepidorrhachis

Linospadix

Loxococcus

Lytocaryum

Manicaria

Manjekia

Marojejya

Masoala

Nenga

Neonicholsonia

Neoveitchia

Nephrosperma

Normanbya

Oertocarpus

Oncosperma                         6

Orania

Parajubaea

Pelagodoxa

Phoenicophorium

Pholidostachys

Physokentia

Pinanga

Podococcus

Ponapea

Prestoea

Ptychococcus

Ptychosperma

Reihardtia

Rhopaloblaste

Rhopalostylis

Roscheria

Roystonea

Satakentia

Sclerosperma

Socratea          1

Solfia

Sommieria

Syagrus

Synechanthus

Tectiphiala

Veitchia

Verschaffeltia

Voaniola

Wallaceodoxa

Welfia

Wendlandiella

Weltinia

Wodvetia

Arecoideae        1                7         0

D.                References

Genus

Acanlhophoenix    Dransfield et al., 2008a, 2008b

Acrocomia         Dransfield et al., 2008a, 2008b

Actinokentia      Dransfield et al., 2008a, 2008b

Actinorhytis      Dransfield et al., 2008a, 2008b

Adonidia          Dransfield et al., 2008a, 2008b

Aiphanes          Borchsenius & Bemal. 1996;
                  Henderson et al., 1997

Allagoptera       Tomlison, 1967

Archontophoenix   Personal observation

Areca             Dransfield, 1984b;
                  Henderson, 2009;
                  Heatubun, 2011;
                  Heambun et al., 2012

Asterogyne        Henderson & Steyermark, 1986;
                  de Granville & Henderson, 1988;
                  Stauffer et al., 2003;
                  Dransfield et al., 2008a, 2008b

Astrocuryum       Kahn & Millan, 1992;
                  Henderson et al., 1997;
                  Borchsenius et al., 1998;
                  Kahn & de Granville, 1998;
                  Kahn, 2008

Attalea           Dransfield et al., 2008a, 2008b

Bactris           Tomlison. 1990; Henderson et al.,
                  1997; Henderson, 2000

Balaka            Dransfield et al., 2008a, 2008b

Barcella          Dransfield et al., 2008a, 2008b

Basselinia        Moore & Uhl, 1984; Essig et al.,
                  1999; Pintaud & Baker, 2008;
                  Pintaud & Stauffer, 2015

Beccariophoenix   Dransfield et al., 2008a, 2008b

Bentinckia        Dransfield et al., 2008a, 2008b

Brassiophoenix    Dransfield et al., 2008a, 2008b

Bwretiokentia     Dransfield et al., 2008a, 2008b

Butia             Gaiero et al., 2011

Calyptrocaix      Dowe & Ferrero, 2001

Calyptrogyne      Henderson et al., 1997;
                  Dransfield et al., 2008a, b

Calyptronoma      Dransfield et al., 2008a, 2008b

Carpentaria       Dransfield et al., 2008a, 2008b

Carpoxvlon        Dransfield et al., 2008a, 2008b

Chamaedorea       Fisher, 1974; Model, 1992

Chambeyronia      Dransfield et al., 2008a, 2008b

Clinosperma       Dransfield et al., 2008a, 2008b

Clinostigma       Dransfield et al., 2008a, 2008b

Cocos             Balaga. 1975; Dransfield et al.,
                  2008a, 2008b

Cyphokentia       Moore & Uhl, 1984;
                  Jaffre & Veillon, 1989

Cyphophoenix      Dransfield et al., 2008a, 2008b

Cyphosperma       Dransfield et al., 2008a, 2008b

Cyrtostachys      Dransfield, 1978;
                  Heatubun et al., 2009

Deckenia          Dransfield et al., 2008a. 2008b

Desmoncus         Putz, 1990; lsnard et al., 2005;
                  Tomlinson & Zimmerman, 2003

Dictyocaryum      Henderson, 1990;
                  Dransfield et al., 2008a, 2008b

Dictyospertna     Dransfield et al., 2008a, 2008b

Dransfieldia      Baker et al., 2006

Drymophloeus      Zona, 1999

Dypsis            Dransfield & Beentje, 1995;
                  Fisher and Maidman, 1999;
                  Dransfield, 2003;
                  Britt & Dransfield, 2005;
                  Hodel et al., 2005;
                  Rakotoarinivo et al., 2009

Elaeis            Dransfield et al., 2008a. 2008b

Euterpe           Henderson & Galeano, 1996;
                  Dransfield et al., 2008a, 2008b

Gaussia           Dransfield et al., 2008a, 2008b

Geonoma           Henderson, 1995;
                  Henderson et al., 1997;
                  Dransfield et al., 2008a, 2008b;
                  Henderson. 2011 a

Hedyscepe         Dransfield et al., 2008a, 2008b

Hetemspathe       Fernando. 1990

Howea             Dransfield et al., 2008a, 2008b

Hydriastele       Baker & Dransfield, 2007

Hvophorbe         Dransfield et al., 2008a. 2008b

Hyospalhe         Skov & Balslev, 1989;
                  Borchsenius et al., 1998

Iguanura          Kiew, 1976; Henderson, 2009

Irartea           Dransfield et al., 2008a, 2008b

Iriartella        Dransfield et al., 2008a, 2008b

Jailoloa          Heatubun et al., 2014

Jubaea            Dransfield et al., 2008a, 2008b

Jubaeopsis        Dransfield, 1989

Kentiopsis        Dransfield et al., 2008a, 2008b

Laccospadix       Dowe, 2010

Lemurophoenix     Dransfield et al., 2008a. 2008b

Leopoldinia       Bemal & Galeano, 2010;
                  Henderson, 2011a, b

Lepidorrhachis    Dransfield et al., 2008a, 2008b

Linospadix        Dowe & Irvine, 1997;
                  Dowe & Ferrero. 2001

Loxococcus        Dransfield et al., 2008a, 2008b

Lytocaryum        Dransfield et al., 2008a, 2008b

Manicaria         Bernal & Galeano, 2010:
                  Fisher & Zona, 2006

Manjekia          Heatubun et al., 2014

Marojejya         Dransfield et al., 2008a, 2008b

Masoala           Dransfield et al., 2008a, 2008b

Nenga             Fernando, 1983; Henderson, 2009

Neonicholsonia    Henderson & Galeano, 1996;
                  Dransfield et al., 2008a, 2008b

Neoveitchia       Dransfield et al., 2008a, 2008b

Nephrosperma      Dransfield et al., 2008a, 2008b

Normanbya         Dransfield et al., 2008a, 2008b

Oertocarpus       Bemal et al., 1991;
                  Henderson et al., 1997;
                  Dransfield et al., 2008a, 2008b

Oncosperma        Fisher et al., 1989;
                  Fisher and Maidman. 1999

Orania            Dransfield et al., 2008a, 2008b

Parajubaea        Dransfield et al., 2008a, 2008b

Pelagodoxa        Dransfield et al., 2008a. 2008b

Phoenicophorium   Dransfield et al., 2008a, 2008b

Pholidostachys    Dransfield et al., 2008a, 2008b

Physokentia       Dransfield et al., 2008a, 2008b

Pinanga           Dransfield, 1978; Henderson, 2009

Podococcus        Bullock, 1980;
                  Van Valkenburg et al., 2007

Ponapea           Dransfield et al., 2008a, 2008b

Prestoea          Henderson & deNevers, 1988;
                  Henderson & Galeano, 1996

Ptychococcus      Dransfield et al., 2008a, 2008b

Ptychosperma      Essig, 1977; Essig, 1978;
                  Dowe & Ferrero, 2001

Reihardtia        Henderson et al., 1997;
                  Henderson, 2002b

Rhopaloblaste     Banka & Baker, 2004

Rhopalostylis     Dransfield et al., 2008a, 2008b

Roscheria         Dransfield et al., 2008a, 2008b

Roystonea         Dransfield et al., 2008a, 2008b

Satakentia        Dransfield et al., 2008a, 2008b

Sclerosperma      Van Valkenburg et al., 2007;
                  van Valkenburg, et al., 2008

Socratea          Bernal-Gonzales & Henderson,
                  1986; Svenning & Balslev, 1998;
                  Pintaud & Millan, 2004

Solfia            Dransfield et al., 2008a, 2008b

Sommieria         Dransfield et al., 2008a, 2008b

Syagrus           Henderson, 1995:
                  Pinheiro et al., 1996:
                  Noblick, 1996; Noblick. 2004;
                  Noblick & Lorenzi, 2010;
                  Noblick et al., 2014;
                  Noblick & Meerow, 2015

Synechanthus      Dransfield et al., 2008a, b

Tectiphiala       Moore, 1978

Veitchia          Dransfield et al., 2008a, 2008b

Verschaffeltia    Dransfield et al., 2008a, 2008b

Voaniola          Dransfield, 1989;
                  Dransfield et al., 2008a, 2008b

Wallaceodoxa      Heatubun et al., 2014

Welfia            Dransfield et al., 2008a, 2008b

Wendlandiella     Dransfield et al., 2008a, 2008b

Weltinia          Henderson et al., 1997;
                  Borchsenius et al., 1998

Wodvetia          Dransfield et al. 2008a, 2008b

Arecoideae


Sara M. Edelman (1,2,3) * Jennifer H. Richards (1,2)

(1) Florida International University, 11200 SW 8th Street, Miami, FL 33199, USA

(2) International Center for Tropical Botany, 4013 Douglas Road, Miami, FL 33133, USA

(3) Author for Correspondence; c-mail: Sedel003@fiu.edu Published online: 18 June 2018

Caption: Fig. 1 Vegetative branching types in the palms (arrows indicate vegetative branch): a No branching type (Hyophorbe laugenicaulis) or solitary: b Lateral axillary branching (Rhapis mulifida); c shoot apical division (Hyphaene dichotoma): d false vivipary (Socratea salazarii); D. abaxial branching (Dypsis lutescens); and E. leaf-opposed branching (Myrialepis paradoxa). Arrow points to branch

Caption: Fig. 2 Plan view of a palm leaf with locations of the three distinct types of stem nodal buds and thus branching types; the stem is not drawn but the encircling leaf base is shown, a Axillary branching--the meristem arises in the axil of the leaf; b Abaxial branching-the meristem is located on the base of the leaf sheath, on the abaxial surface of the leaf; and c Leaf-opposed branching-the meristem is borne on the stem of the palm, enclosed by the outer edges of the leaf sheath and opposite to the lamina and petiole

Caption: Fig. 3 Distribution of branching types in the palm family (Arecaceae) on a sub-family level cladogram (a key to branching types; b sub-family cladogram)

Caption: Fig. 4 Distribution of branching types in the Calamoideae on a genus level cladogram. Key the same as Fig. 3a

Caption: Fig. 5 Distribution of branching types in the Calamoidcac on a genus level cladogram. Key the same as Fig. 3a

Caption: Fig. 6 Distribution of branching types in the Arecoideae on a genus level cladogram (a entire cladogram showing further break down; b Socratea-Parajubaea; c Podococcus-Clinostigma d Chambeyronia-Neoveitchia; and e Ptychospemia-Normanbya). Key the same as Fig. 3a
Table 1 Definitions of branching terms used in this study.
In the first column terms in bold are the branching terms
used in this review with synonyms (non-bold terms listed
in parentheses directly below the bold); indented below the
new term arc terms from the literature that are included in
the new term. Additional columns provide references,
definitions of the branching type, and palm examples

Term (synonym(s))    Refercnce(s)       Definition

Lateral axillary     Tomlinson 1990     Branch originates in the
branch                                  axil of the leaf

Basal sucker         Tomlinson 1990     Lateral axillaiy branch
                                        immediately grows
                                        upward, limited to
                                        basal internodes

Aerial lateral       Tomlinson 1990     Lateral axillary branch
axillary branch                         is not limited to basal
                                        intcmodcs

Dormant basal        Tomlinson 1990     Basal sucker outgrowth
suckers                                 is dormant until death
                                        of parent stem

Rhizomatous          Zimmermann         Vegetative outgrowth of
branch               and Tomlinson      axillary meristcm at base
                     1967               of stem where monopodial
                                        or sympodial units form
                                        a plagiotropic rhizome

Sympodial            Zimmermann         Vegetative outgrowth of
rhizomatous          and Tomlinson      axillary meristem at base
branch               1967               of stem where sympodial
                                        units form a plagiotropic
                                        rhizome

Monopodial           Bell & Tomlinson   Vegetative outgrowth of
rhizomatous          1980               axillary meristcm at base
branch                                  of stem where monopodial
                                        units form a plagiotropic
                                        rhizome

Shoot apical         Tomlinson 1990     Branch originates in the
division                                apical meristcm, most
(Apical                                 commonly as a division
branch)                                 of the apical meristem

Apical division      Gola 2014          More or less equal
                                        division of apical meristem,
                                        resulting in two independent
                                        functioning axes

Apical isotomy       Gola 2014          Equal division apical
                                        meristem that results in
                                        two independent functioning
                                        axes of similar size and
                                        morphology

Apical anisotomy     Gola 2014          Unequal division apical
                                        meristem that results in
                                        two independent functioning
                                        axes of different size and
                                        morphology

Nannorhops           * new term         Equal division apical
branching                               meristem that results in
                                        two independent functioning
                                        axes of different size and
                                        morphology

Adventitious bud/    Fisher 1973        Meristem not in typical
branching                               position

False vivipary       Fisher and         Adventitious vegetative
(prolification,      Dransfield 1977;   outgrowth at the shoot
vegetative           Bell and Btyan     apex of the inflorescence,
transformation       2008               growing independently
of                                      of inflorescence axis
inflorescence,
broadly
as proliferation
(sensu latu))

Proliferation        Bell and Bryan     Adventitious mcristcm
(sensu stricto)      2008               originates from vegetative
                                        material, usually leaves

Abaxial branch       Fisher 1973,       Vegetative branch meristem
                     Fisher et al.      borne on the abaxial
                     1989               surface of leaf, on the
                                        base of the leaf sheath

Leaf-opposed         Fisher and         Vegetative branch meristem
branch               Dransfield 1979;   borne on the stem opposite
                     Tillich 1998       of leaf and enclosed in the
                                        leaf sheath

Term (synonym(s))    Palm example

Lateral axillary     Serenoa repens
branch

Basal sucker         Pytchosperma
                     macarthurii

Aerial lateral       Geonoma baculifera,
axillary branch      Hyospathe elegans

Dormant basal        Plectomia spp.
suckers

Rhizomatous          Rhapis excelsa
branch

Sympodial            Rhapis exelsa
rhizomatous
branch

Monopodial           No palm example
rhizomatous
branch

Shoot apical         Hvphaene thebaica
division
(Apical
branch)

Apical division      Hyphaene coriaceae

Apical isotomy       Nypa fruticans

Apical anisotomy     Eugeissona tiistis

Nannorhops           Nannorhops richiana
branching

Adventitious bud/    Socratea salazarii
branching

False vivipary       Calamus castaneiis
(prolification,
vegetative
transformation
of
inflorescence,
broadly
as proliferation
(sensu latu))

Proliferation        No palm example
(sensu stricto)

Abaxial branch       Dyipsis lutescens

Leaf-opposed         Myrialepis
branch

Table 2 Palm subfamilies and their species counts for the
five branching types and their combinations. References for
sub-families can be found in the individual sub-family tables

Subfamily       Species count   Number of spcies that exhibit...

                                No          Lateral
                                branching   axillary

Arccoidcae      1112/1376        657        423
Calamoidcae     395/659           58        292
Ccroxyloideae   47/47             46          1
Coryphoidcae    381/492          283         92
Nypoidcae       1/1
Arccaceae       (1903/2501)     1043        646

Subfamily       Number of spcies that exhibit...

                Shoot apical   Shoot apical       FALSE
                dichotomy      dichotomy with     vivipary
                               lateral axillary

Arccoidcae      13              1
Calamoidcae      6             28                 2
Ccroxyloideae
Coryphoidcae     3              3
Nypoidcae        1
Arccaceae       21             31                 2

Table 3 Key to major branching types in the palms; palm branching
types were distinguished by location of the meristem

If there is more than one crown meristem used for branching is (a)
axillary, (b) apical, (c), adventitious (atypical position)

a. Axillary                         (1) Later axilary branching

b. Apical                           (2) Shoot apical division

c. Adventitious: bud borne on (a) inflorescence, (b) leaf sheath, (c)
sterm

a. Inflorescence                    (3) False vivipary

b. Leaf sheath, base                (4) Abaxial

c. Stern, enclosed in leaf sheath   (5) Leaf-opposed
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Author:Edelman, Sara M.; Richards, Jennifer H.
Publication:The Botanical Review
Article Type:Report
Date:Mar 1, 2019
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