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Research Article
Phylogenomics and a new classification of the tropical genus Heliconia L. (Monocots, Zingiberales, Heliconiaceae)
expand article infoW. John Kress‡, Tomáš Fér§, Mónica M. Carlsen|
‡ Smithsonian Institution, Washington, United States of America
§ Charles University, Prague, Czech Republic
| Missouri Botanical Garden, St. Louis, United States of America

Corresponding author: W. John Kress ( kressj@si.edu )

Academic editor: Lyubomir Penev
© 2025 W. John Kress, Tomáš Fér, Mónica M. Carlsen.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Kress WJ, Fér T, Carlsen MM (2025) Phylogenomics and a new classification of the tropical genus Heliconia L. (Monocots, Zingiberales, Heliconiaceae). PhytoKeys 251: 37-66. https://doi.org/10.3897/phytokeys.251.130409
Open Access

Abstract

Members of the genus Heliconia L. (Heliconiaceae) have evolved complex interactions with both insect herbivores and hummingbird pollinators in tropical forests and secondary growth where they are abundant and diverse. Many of these same species have also been cultivated as ornamentals around the world for hundreds of years because of their extraordinary colors and forms. Because of the large size, fleshy nature, and tropical distribution, and despite a long taxonomic history, the classification and phylogenetic relationships of species of Heliconia have not received sufficient attention to date. No complete classification has been published for the entire genus, although some preliminary attempts have been offered. In this paper we used tissue sampled from field and herbarium collections of 136 species for genomic sequencing to determine the phylogenetic patterns within Heliconia, which then served as the basis for a new evolutionary classification of the genus. This new classification, which is based on extensive field work and the phylogenomic insights provided here, includes 187 currently recognized species. The new classification of Heliconia is composed of 17 sections in five subgenera with all groups well-supported in the phylogenomic analysis. Four subgenera are each composed of two sections and one subgenus includes nine sections. One subgenus and 10 sections are described as new.

Key words

Classification, phylogenomics, target enrichment, tropical, Zingiberales bait set

Introduction

The Genus Heliconia L. – background

The Zingiberales, an order of monocots native almost exclusively to tropical habitats, are comprised of eight families, 110 genera, and over 2,600 species (Carlsen et al. 2018). Many of these species are of economic and cultural importance to humans, including major agricultural crops (e.g., bananas, in the genus Musa), culinary staples (e.g., turmeric and ginger, in the genera Curcuma and Zingiber), horticultural ornamentals (e.g., prayer plants [Calathea], globbas [Globba], cannas [Canna], and heliconias [Heliconia]), and even national symbols (e.g., the travelers tree of Madagascar, in the genus Ravenala Adans.).

Members of the genus Heliconia (Heliconiaceae) have been cultivated as ornamentals around the world for hundreds of years because of their extraordinary colors and forms, and continue to have a prominent position in the horticultural trade (Berry and Kress 1991). A professional society (www.heliconia.org) is devoted to consolidating and distributing information on these plants to scientists, amateur enthusiasts, and the commercial trade community. Many of these same species as well as others in the genus have also been shown to have evolved complex interactions with both insect herbivores (e.g., Garcia-Robledo et al. 2013a, b) and hummingbird pollinators (e.g., Temeles and Kress 2003; Temeles et al. 2012; Gowda and Kress 2013) in tropical forests and secondary growth where they are abundant and diverse.

An understanding and appreciation of the evolutionary history of Heliconia as well as species delimitations and classification will assist scientists in determining the breadth of ecological patterns in community composition, help horticulturists to promote commercial utilization of these plants, and inform conservationists in determining workable plans to protect species. Unfortunately, because of their large size, fleshy nature, and tropical distribution, the taxonomy and phylogenetic relationships of the 187 currently recognized species of Heliconia, have not received sufficient attention until now to fully understand their evolutionary relationships. Moreover, no complete classification has been published for the entire genus, although some preliminary attempts have been offered (e.g., Andersson 1992; Kress et al. 1999). Here, a phylogenomic analysis of species relationships coupled with extensive field work provide the foundation for a revised classification of Heliconia.

Taxonomy and classification – a history

The first botanical description of Heliconia was by Plumier (1703) in “Nova Plantarum Americanarum Genera”. He provided a short generic description of the genus Bihai and polynomials for three taxa. In “Species Plantarum” Linnaeus (1753) combined these three taxa into a single species, Musa bihai, retaining Plumier’s exact diagnoses and placing the “variegatis” variety first. Miller (1754, 1768) and Adanson (1763) considered these plants generically distinct from other species of Musa and used the name Bihai. In “Mantissa Plantarum”, Linnaeus (1771) also segregated M. bihai into its own genus, Heliconia L., and provided a generic description with a short description of a single species, H. bihai. Kuntze (1891; substituting the variant spelling Bihaia) and later Griggs (1904, 1915) recognized the earlier generic name and transferred all species of Heliconia known to them into Bihai. However, at the International Botanical Congress held in Vienna in 1905, Heliconia was reinstated as a nomen conservandum (Farr et al. 1979).

Around the turn of the last century, a number of workers attempted revisions or summaries of the genus, including Petersen (1890), Kuntze (1891), Baker (1893), and Schumann (1900). Griggs, one of the most knowledgeable students of Heliconia due to his study of plants in the field, subsequently published several papers on the genus (1903, 1904, 1915), eventually accepting 38 neotropical species. In the latter decades of the twentieth century a flurry of new species was recognized and described (e.g., Daniels and Stiles 1979; Abalo and Morales1982, 1983; Kress 1984; Andersson 1985a) and today 187 species are accepted here with many names in synonymy.

With regards to infrageneric classification of Heliconia, early efforts were based solely on the shape of the cincinnal bracts with later workers adding plant size, leaf orientation, and inflorescence habit and structure to devise more detailed classifications. Kuntze (1891) published the first infrageneric taxon (sect. Taeniostrobus Kuntze) above the rank of species. Baker (1893) later divided the genus into two subgenera, Platychlamys Baker and Stenochlamys Baker, recognizing 29 species in all (although some of these species are no longer accepted today). Schumann (1900) followed Baker’s classification except he placed Platychlamys in synonymy under Taeniostrobus and altered the ranks of these taxa from subgenera to sections.

Griggs (1903) recognized that infrageneric groups based upon a single character, for example cincinnal bract shape, were inadequate. He subsequently used plant habit and cincinnal-bract orientation to build a diagnostic key that included three subgenera, Stenochlamys, Platychlamys, and Taeniostrobus, which had been earlier defined by others. However, in his final publication (Griggs 1915) he replaced this classification with one consisting of two subgenera (Taeniostrobus and Stenochlamys) and six subordinate taxa of unspecified rank. Plant habit and height, inflorescence orientation, and distance between adjacent inflorescence bracts were used to recognize subgeneric groups and all 38 neotropical species known at that time were included in his treatment.

No attempts were made to revise Griggs’ classification until the latter part of the twentieth century with publications by Andersson (1981, 1985a, b, 1992), Kress (1984, 1990), Kress et al. (1993), Betancur and Kress (1995), and Kress et al. (1999). Andersson used a wider suite of morphological traits (including flower resupination and pollen features) and in his latter publications based his classifications on a cladistic analyses of these characters. His first paper (Andersson 1981) deals with the circumscription of Heliconia sect. Heliconia; the second treatment (Andersson 1985a) subdivided Heliconia subgen. Stenochlamys into six sections (Lanea, Stenochlamys, Proximochlamys, Lasia, Cannastrum, and Zingiberastrum); and in the third publication (Andersson 1985b), species were arranged into four subgenera (Taeniostrobus, Heliconia, Stenochlamys, and Griggsia), with several subgenera further divided into both formal and informal groups. In his final publication Andersson (1992) provided the most complete and detailed classification of the genus to date recognizing five subgenera, ten sections, and eight informal groups (subgen. Taeniostrobus; subgen. Heliconia [sects. Episcopalis, Heliconia, Tortex, Tenebria]; subgen. Stenochlamys [sects. Lanea, Stenochlamys, Proximochlamys, Lasia, Cannastrum, Zingiberastrum]; subgen Heliconiopsis ; subgen. Griggsia [informal groups H. griggsiana, H. pogonantha, H. longa, H. platystachys, H. rostrata, H. trichocarpa, H. obscura, and H. nutans]). Some of the informal groups established by Andersson within subgenus Griggsia had been earlier designated by Kress (1984).

Subgenus Heliconiopsis, encompassing the tropical Asian-Pacific species of the genus, has been an anomaly treated quite differently by various authors. Miquel (1859) erected the new genus Heliconiopsis for the only Asian-Pacific species known at that time. Baker (1893), Schumann (1900), and Winkler (1930) synonymized all taxa proposed for Asian-Pacific material under Heliconia bihai. Green (1969) later recognized that tropical Asian-Pacific heliconias were quite different from those in the American tropics, but considered that all populations belonged to a single polymorphic species, Heliconia indica Lam. Eventually, after extensive field work, Kress (1990) recognized six species that stretched from Samoa to Papua New Guinea and New Caledonia. He, as concurred by Andersson, considered these species to constitute a diagnosable group recognized as subgen. Heliconiopsis.

Most recently, Kress and colleagues (Kress et al. 1993; Betancur and Kress 1995; Kress et al. 1999) have proposed a series of preliminary classifications that were built on earlier taxonomies and based primarily on morphological traits. These newer classifications were a response in part to the many new species of Heliconia discovered and described from Costa Rica, Panama, Colombia, Ecuador, and Peru in the 1980s and 1990s. These categorizations were an attempt to account for the massive diversity represented by these new taxa and often focused on species from a single country. The most comprehensive preliminary classification (Kress et al. 1999) encompassed the 93 species then known from Colombia and included four subgenera and 22 sections, many indicated as “ined.” by the authors to denote that the work was in progress (subgen. Heliconia [sects. Heliconia, Episcopalis, Tortex, Farinosae “ined.”, Complanatae “ined.”, and Tenebria]; subgen. Taeniostrobus ; subgen. Stenochlamys [sects. Lanea, Stenochlamys, Proximochlamys, Lasia, Cannastrum, and Zingiberastrum], subgen. Griggsia [sects. Griggsia “ined.”, Barbatae “ined.”, Arcuatae “ined.”, Longae “ined.”, Obscurae “ined.”, Dromedarius “ined.”, Sigmoideae “ined.”, Rostratae “ined.”, Pendulae “ined.”, and Retiformes, “ined.”]). No species in subgen. Heliconiopsis were included as none are known from South America.

The only molecular analysis of phylogenetic relationships in Heliconia was provided by Iles et al. (2017). They demonstrated that subgenera and sections designated by earlier taxonomists as outlined above (e.g., Andersson 1992; Kress et al. 1993; Kress et al. 1999) were not monophyletic. However, with the molecular data in their analysis limited to seven plastid and nuclear markers, the support values for almost all clades throughout the tree were very low. None-the-less, this earlier analysis has provided a gene-based foundation for the more extensive genomic-based analysis in the current investigation.

Materials and methods

Taxon sampling

We included 136 Heliconia species (ca. 73% of the species in the genus) representing the entire spectrum of morphological and geographical diversity within the genus, as well as 5 outgroup genera (Musaceae: Ensete superbum (Roxb.) Cheesman and Musella lasiocarpa (Franch.) C.Y. Wu; Lowiaceae: Orchidantha chinensis T.L. Wu; and Strelitziaceae: Strelitzia nicolai Regel & Körn. and Ravenala madagascariensis Sonn.). See Suppl. material 1: table S1 for specimen information and accession numbers.

Library preparation and target loci capture

Total genomic DNA was extracted from either silica dried leaves or herbarium specimens using a modified CTAB protocol (Doyle and Doyle 1987). Libraries were constructed using the NEBNext Ultra™ II DNA Library Prep kit with Sample Purification Beads and Multiplex oligos for Illumina (96 Index Primes) (New England BioLabs Inc., Ipswich, MA, USA). Library concentration and expected size were confirmed using a High Sensitivity D1000 ScreenTape run on a 4150 TapeStation system (Agilent Technologies Inc., Santa Clara, CA, USA), and Qubit dsDNA HS Assay Kit run on a Qubit 2.0 Fluorometer (Invitrogen, Carlsbad, CA, USA). Libraries were combined in hybridization pool reactions consisting of 10–12 Heliconia species per reaction. All these steps were performed at the Laboratories for Analytical Biology (LAB) of the Smithsonian National Museum of Natural History. In solution hybridization of the library pools to the target loci was performed by Arbor Biosciences LLC (Ann Arbor, Michigan) using the myBaits Custom 1-20K kit designed for Zingiberales (Carlsen et al. 2018). Sequencing was performed either in an illumina MiSeq with v.3 chemistry kits (250 bp paired-end) at the Smithsonian or in an illumina NovaSeq S4 system (150 bp paired-end) by Arbor Biosciences (Illumina Inc., San Diego, CA).

Bioinformatic analyses

Raw paired-end reads were quality trimmed and adapters were removed using a combination of Trimmomatic v. 0.39 (Bolger et al. 2014) with the following parameters ILLUMINACLIP:TruSeq3-PE-2NEB.fa:2:30:10 LEADING:3 TRAILING:3 SLIDINGWINDOW:10:20 MINLEN:40 and Trim Galore v. 0.6.4 (https://github.com/FelixKrueger/TrimGalore) with default parameters. Cleaned reads were assembled using HybPiper v. 1.3.1 (Johnson et al. 2016). Coding sequences were extracted using Exonerate (Slater and Birney 2005) with the HybPiper retrieve_sequences.py script. From the set of fasta files resulting for each locus, we removed: 1. Loci with no data, 2. Loci containing species with extremely long branches (i.e., loci matched is 150% longer than reference loci), 3. Loci with paralogs warnings in HybPiper, 4. Loci without a single outgroup taxa represented, and 5. Loci recovered for less than 75% of the taxa of Heliconia sampled (i.e. with fewer than 102 taxa of Heliconia represented). Each of the remaining loci recovered were separately aligned with MAFFT v. 7.407 (Katoh and Standley 2013) with “--auto” parameter. Alignments were manually checked for misaligned regions and extremely gappy areas, which were either removed or fixed. Alignment statistics were calculated with AMAS (Borowiec 2016), and from the results of this analysis, we further removed each individual taxa with >50% missing sequence for a given locus and any individual locus with >50% total missing data per alignment. Single locus tree reconstruction was performed using RAxML v. 8.2.12 (Stamatakis 2014), with no partition scheme, GTRCAT model of evolution, and 500 bootstrap replicates. Coalescence based species tree reconstruction was performed with ASTRAL III v. 5.7.1 (Zhang et al. 2018) using the individual trees generated with RAxML above. Individual branches with low support in these trees were collapsed if bootstrap support was below 50%. All bioinformatics analyses were performed in the Smithsonian Institution High Performance Computing Cluster (SI HPC – Hydra) (https://doi.org/10.25572/SIHPC).

Constructing the revised classification

The resultant Heliconia species tree topology was assessed for congruence with past classifications and for major well-supported clades corresponding to diagnostic morphological traits and geographic patterns. Groups of species, which had been assigned to subgenera and sections in the earlier classifications, were assessed with respect to the newly identified clustering of taxa and clade support. Only clades with Local Posterior Probability (LPP) 0.90 or greater were considered for higher ranks at the subgeneric and sectional level.

In this phylogenomic analysis molecular data were only available for 136 of the 187 currently recognized species of Heliconia. Therefore, the remaining 51 species were placed into subgenera and sections according to their morphological characteristics only. In the case of significant conflict between the phylogenetic placement and morphological traits, taxa were tentatively classified according to their morphological characteristics. Their placement will be tested in future analyses with additional field observations and molecular data.

Geographic distribution

Distributions of species by country within each subgenus and section were determined from specimen data included in an analysis of conservation assessment (Kress et al. 2024) and as part of a taxonomic monograph of the genus (Kress unpubl.).

Results and discussion

A new genome-scale dataset for Heliconia

All raw reads newly generated for this study are deposited in NCBI BioProject accession number PRJNA1204471. The average number of trimmed, high-quality, non-duplicated reads obtained was 3,242,087 per Heliconia sample, ranging from 368,676 reads (in H. marginata (Griggs) Pittier) to 19,241,471 reads (in H. adflexa (Griggs) Standl.) (Suppl. material 1: table S1). On average 51% of these high-quality reads mapped to the target nuclear loci from the Zingiberales bait set (Carlsen et al. 2018), ranging from 19% (in H. lingulata Ruiz & Pav.) to 75% (in H. trichocarpa Daniels & Stiles). A final set of 452 high-quality target nuclear loci was used for phylogenetic tree inference after a strict and comprehensive data clean-up (see Methods above). They represent ca. 38% of the total 1,180 target loci included in the Zingiberales bait set. The number of high-quality target loci recovered for individual Heliconia species ranged from 275 in H. marginata to 448 in H. carmelae Abalo & Morales. The alignment length of each individual high-quality nuclear locus ranged from 300 to 3,496 bp (on average 1,002 bp), their total amount of missing data from 0 to 49% (on average 17%), and the proportion of informative sites varied between 0.03 and 0.48 (on average 0.14) (Suppl. material 1: table S1). Individual loci alignments are available on Dryad [https://doi.org/10.5061/dryad.mcvdnck9g].

The topology recovered in the coalescence species tree analysis using ASTRAL (Fig. 1) shows high LPP support values of 0.90–1.00 for most branches (Suppl. material 2: fig. S1). Only nine branches along the backbone of the Heliconia phylogeny were weakly supported by 0.46–0.89 LPP; these are among the shortest branches of the tree. Some shallow branches (i.e. species-pairs relationships) with LPP support values lower than 0.90 are scattered throughout the phylogeny (Suppl. material 2: fig. S1).

Figure 1. 

Circular phylogeny of species of Heliconia included in the genomic analysis with the new classification indicated. The six species with gray branches represent significant conflicts in genomic and morphological evidence, with the latter given priority in species placement in the classification (see text and Suppl. material 2: fig. S1).

A new genome-based phylogeny of Heliconia

The phylogenomic analysis of nearly three-quarters of the known species of Heliconia provides a robust evolutionary framework and a firm foundation for a newly revised classification of the genus (Figs 1, 2; see below). All five major lineages (i.e., designated here as taxonomic subgenera; Fig. 2) in the backbone of the new phylogeny of Heliconia and all 17 clades within them (i.e., designated here as taxonomic sections; Fig. 1) are highly supported, even though relationships among some of the subgenera and sections are not fully resolved (Figs 1, 2). This is a striking difference when compared to the previous phylogeny of Heliconia (Iles et al. 2017), in which the bootstrap support value for almost all clades was very low. Of the major lineages recognized here, only one (subgenus Heliconiopsis) had bootstrap support greater than 90% in Iles et al. (2017). Two of the remaining four major lineages recovered here (subgenera Stenochlamys and Heliconia) were polyphyletic in their earlier analysis, whereas the other two (subgenera Colubrosae and Taeniostrobus) had only weak bootstrap support (<70%). At the sectional level, only six of the sections recognized in our analyses (sections Heliconiopsis, Perplexa, Stenochlamys, Lasia, Longiflorae and Lanea) were supported by bootstrap values greater than 90% in their analysis. For the remaining eleven sections, most of the species recognized here were clustered together in the Iles et al. (2017) analysis, but almost all of these sections lacked even moderate statistical support. Clearly more nucleotide data were needed to resolve their basic cladistic structure than were available in that investigation. Iles et al. (2017) concluded that subgenera and sections designated by earlier taxonomists (e.g., Andersson 1992; Kress et al. 1993; Kress et al. 1999) were not monophyletic, but the authors did not propose a new classification, perhaps due to the lack of support for most clades recovered in that study.

Figure 2. 

Synopsis of the new classification based on the genomic analysis (Local Posterior Probability [LPP] is indicated for each branch).

The new classification of the 187 currently recognized species of Heliconia presented below takes into consideration earlier suggested classifications, but more importantly utilizes the newly generated extensive phylogenomic data set to support the proposed taxonomic groups. Additionally, most of the 51 species for which genomic data were not available have also been confidently placed in the classification based on their morphological characteristics. Only six species with molecular data proved difficult to classify because of substantial disparities between their placement in the phylogeny and their morphological characteristics. These taxa (H. hirsuta, H. lourteigiae, H. pendula, H. pardoi, H. brenneri, and H. titanum; indicated by “†” in the classification below) were tentatively classified according to their morphological characteristics. In the phylogeny recovered here, all of these taxa are positioned towards the base of their respective clades, therefore suggesting that these disparities may be due to sampling or sequencing errors and further analyses are needed to verify their placements.

More data are clearly needed to confidently place the few species with genomic data but contradictory morphological characters as well as verifying the position of the taxa lacking any genomic data. As new molecular traits become available and more detailed observations of living plants are conducted in the field in both Central and South America, it is expected that some species will shift position in the classification and that new taxa will be added as they are discovered and described. As detailed in a companion publication (Kress et al. 2024), increased environmental degradation and overexploitation of some taxa may significantly limit the time remaining for this future work without significant efforts in the protection of populations, species, and habitats.

A new classification of Heliconia

The new classification of Heliconia includes 17 sections in five subgenera with all clades supported by significant LPP support values > 0.90 (Table 1; Figs 1, 2; Suppl. material 2: fig. S1). Four subgenera (Figs 3, 4) are each composed of two sections (subgen. Heliconiopsis [sects. Heliconiopsis, Perplexae]; subgen. Stenochlamys [sects. Stenochlaymys, Angustae], subgen. Taeniostrobus [sects. Taeniostrobus, Aurantiacae], and subgen. Colubrosae [sects. Colubrosae, Curvae]). The fifth, subgen. Heliconia (Figs 4, 5), includes nine sections. Four of these sections were previously considered to form distinctive taxonomic subdivisions and are here unequivocally supported as monophyletic entities (sects. Heliconia, Lasia, Cannastrum, and Longiflorae [earlier Zingiberastrum]). The five remaining sections include species previously scattered and classified among various informal groups and unpublished subdivisions. Here these taxa are sorted into distinct lineages with formal designations (sects. Lanea, Longae, Griggsia, Episcopales, and Pastazae). Each of the nine sections in subgen. Heliconia has high LPP support (0.92–1.0) and is characterized by a suite of vegetative and reproductive traits. Former subgeneric and sectional names are retained where possible.

Figure 3. 

Representatives of three subgenera and six sections in the new classification of Heliconia A–D subgen. Heliconiopsis A, B sect. Heliconiopsis: A H. solomonensis B H. indica C, D sect. Perplexae: C H. gaiboriana D H. impudica E–H subgen. Stenochlamys E, F sect. Stenochlayms: E H. psittacorum F H. richardiana G, H sect. Angustae: G H. laneana H H. velloziana I–L subgen. Taeniostrobus I, J sect. Taeniostrobus: I H. reticulata J H. collinsiana K, L sect. Aurantiacae: K H. schiedeana L H. aurantiaca.

Figure 4. 

Representatives of two subgenera and six sections in the new classification of Heliconia A–D subgenus Colubrosae A, B sect. Colubrosae: A H. dielsiana B H. mutisiana C, D sect. Curvae: C H. burleana D H. gilbertiana E–L subgen. Heliconia E, F sect. Heliconia: E H. bihai F H. orthotricha G, H sect. Lasia: G H. julianii H H. lasiorachis I, J sect. Cannastrum: I H. deflexa J H. venusta K, L sect. Longiflorae: K H. longiflora L H. schumanniana.

Figure 5. 

Representatives of one subgenus and five sections in the new classification of Heliconia A–L subgenus Heliconia A, B sect. Lanea: A H. lingulata B H. zebrina C, D sect. Griggsia: C H. gigantea D H. griggsiana E–H sect. Longae: E H. curtispatha F H. longa G H. mariae H H. atratensis I, J sect. Episcopales: I H. episcopalis J H. rostrata K, L sect. Pastazae: K H. pastazae L H. rigida.

Table 1.
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Synopsis of the new classification of Heliconia (also see Fig. 2).

1. Heliconia L., Mant. Pl. 2: 147, 211. 1771, nom. cons.
Bihai Miller, Gard. Dict. ed. 4, 1: B1. 1754, nom. rej. (≡ Heliconia L.).
Type. Heliconia bihai (L.) L.
1.1. Heliconia subgenus Heliconiopsis (Miq.) W.J.Kress, Allertonia 6: 15. 1990.
Heliconiopsis Miq., Fl. Nederl. Ind. 3: 590. 1859.
Type. Heliconiopsis amboinensis Miq. nom. illeg. (≡ Heliconia buccinata Roxb.).
1.1.1. Heliconia section Heliconiopsis (Miq.) W.J.Kress, comb. et stat. nov.
Heliconiopsis Miq., Fl. Nederl. Ind. 3: 590. 1859.
Type. Heliconiopsis amboinensis Miq. (≡ Heliconia buccinata Roxb.).
1.1.2. Heliconia section Perplexae W.J.Kress, sect. nov.
Type. Heliconia impudica Abalo & Morales.
1.2. Heliconia subgenus Stenochlamys Baker, Ann. Bot. 7: 190, 194–200. 1893.
Type. Heliconia psittacorum L. f. (designated by L. Andersson, Opera Bot. 82: 22. 1985).
1.2.1. Heliconia section Stenochlamys (Baker) K. Schum., Engler A, ed., Pflanzenr. IV. 45: 37. 1900.
Heliconia subgen. Stenochlamys Baker, Ann. Bot. 7: 190, 194–200. 1893.
Heliconia section Proximochlamys L. Anderss., Opera Bot. 82: 73. 1985. Type: Heliconia densiflora Verlot.
Heliconia section Zingiberastrum L. Anderss., Opera Bot. 82: 100. 1985. Type: Heliconia hirsuta L. f.
Type. Heliconia psittacorum L. f. (designated by L. Andersson, Opera Bot. 82: 22. 1985).
1.2.2. Heliconia section Angustae W.J.Kress, sect. nov.
Type. Heliconia angusta Vell.
1.3. Heliconia subgenus Taeniostrobus (Kuntze) Griggs, Bull. Torrey Bot. Club 30: 643. 1903.
Bihai section Taeniostrobus Kuntze, Revisio Generum Plantarum. Pars I: 684. 1891.
Type. Bihai imbricata Kuntze (Heliconia imbricata (Kuntze) Baker (designated by L. Andersson, Flora of Ecuador 22: 11. 1985.)
1.3.1. Heliconia section Taeniostrobus Kuntze
Bihai section Taeniostrobus Kuntze, Revisio Generum Plantarum. Pars I: 684. 1891.
Heliconia section Tortex L. Anderss., Flora of Ecuador 22: 19. 1985. Type. Heliconia latispatha Benth.
Heliconia section Tenebria L. Anderss., Opera Bot. 111: 37. 1992: Type. Heliconia tenebrosa Macbr.
Type. Bihai imbricata Kuntze (Heliconia imbricata (Kuntze) Baker) (designated by L. Andersson, Flora of Ecuador 11: 19. 1985.).
1.3.2. Heliconia section Aurantiacae sect. nov.
Type. Heliconia aurantiaca Ghiesbr. ex Lemaire.
1.4. Heliconia subgenus Colubrosae W.J.Kress, subgen. nov.
Type. Heliconia dielsiana Loes.
1.4.1. Heliconia section Colubrosae sect. nov.
Type. Heliconia dielsiana Loes.
1.4.2. Heliconia section Curvae sect. nov.
Type. Heliconia burleana Abalo & Morales.
1.5. Heliconia subgenus Heliconia
1.5.1. Heliconia section Heliconia
1.5.2. Heliconia section Lasia L. Anderss., Opera Bot. 82: 78. 1985.
Type. Heliconia velutina L. Anderss.
1.5.3. Heliconia section Cannastrum L. Anderss., Opera Bot. 82: 86. 1985.
Type. Heliconia metallica Planch. & Linden ex. Hook.
1.5.4. Heliconia section Longiflorae sect. nov.
Type. Heliconia longiflora R.R.Smith.
1.5.5. Heliconia section Lanea L. Anderss., Opera Bot. 82: 23, 30. 1985.
Type. Heliconia lingulata Ruiz & Pav.
1.5.6. Heliconia section Griggsia (L. Anderss.) W.J.Kress, comb. et stat. nov.
Heliconia subgenus Griggsia L. Anderss., Flora of Ecuador 22: 42. 1985.
Type. Heliconia griggsiana L.B.Smith.
1.5.7. Heliconia section Longae sect. nov.
Type. Heliconia longa (Griggs) Winkler.
1.5.8. Heliconia section Episcopales L. Anderss., Opera Bot. 111: 34. 1992.
Type. Heliconia episcopalis Vell.
1.5.9. Heliconia section Pastazae sect. nov.
Type. Heliconia pastazae L. Anderss.

Provided below for each of the five subgenera and 17 sections are brief descriptions, taxonomic notes, designated types, and included species. “*” indicates placement based on genomic data and morphological traits (136 species); “+” indicates placement based on morphological traits only (51 species); “†” indicates conflict in placement between genomic and morphological data in which the latter evidence was given priority (six species).

Heliconia L., Mant. Pl. 2: 147, 211. 1771, nom. cons.

Figs 3, 4, 5

Bihai Miller, Gard. Dict. ed. 4, 1: B1. 1754, nom. rej. (≡ Heliconia L.).

Type

Heliconia bihai (L.) L.

Heliconia subgenus Heliconiopsis (Miq.) W.J.Kress, Allertonia 6: 15. 1990.

Figs 3A–D

Heliconiopsis Miq., Fl. Nederl. Ind. 3: 590. 1859.

Type

Heliconiopsis amboinensis Miq. nom. illeg. (≡ Heliconia buccinata Roxb.).

Description and taxonomic notes

Medium- to large-sized rhizomatous herbs with Musa-like habit. Inflorescence erect or pendent, with peduncle, rachis and cincinnal bracts green to variously colored with red and yellow; cincinnal bracts distichous or spirally arranged, horizontal to deflexed. Flowers with diurnal or nocturnal anthesis, resupinate and held at right angles to bracts or not resupinate and fully or partially enclosed in bracts; perianth uniformly curved, essentially green or white to green to yellow, variously shaped; ovary green to yellow to orange. Fruits bright red, orange, or blue, glabrous to tomentose. subgenus Heliconiopsis originally included only the six species native to the Asian-Pacific tropics (defined here in sect. Heliconiopsis), but now encompasses at least four additional species from South America contained in sect. Perplexae. Although the latter species are quite distinctive from the paleotropical taxa in inflorescence and flower morphology and color, the phylogenomic analysis provides strong support (LPP = 1.0) for the inclusion of these disparate species in a single subgenus, which is sister to all other heliconias. The morphological characters that link these two sections are not obvious.

Distribution

Tropical Asia-Pacific and Andean South America (Colombia, Ecuador, Fiji, Indonesia, New Caledonia, Papua New Guinea, Samoa, Solomon Islands, Vanuatu).

Heliconia section Heliconiopsis (Miq.) W.J.Kress, comb. et stat. nov.

urn:lsid:ipni.org:names:77355024-1
Figs 3A, B

Heliconiopsis Miq., Fl. Nederl. Ind. 3: 590. 1859.

Type

Heliconiopsis amboinensis Miq. (≡ Heliconia buccinata Roxb.).

Description and taxonomic notes

Medium- to large-sized rhizomatous herbs with Musa-like habit. Inflorescence erect or pendent, with peduncle, rachis and cincinnal bracts green-colored; cincinnal bracts distichous or spirally arranged. Flowers with diurnal or nocturnal anthesis, not resupinate and fully or partially enclosed in bracts; perianth uniformly curved, essentially green; ovary green to yellow to orange. Fruits bright red or orange, glabrous to tomentose. This section includes all six known species native to the Asian-Pacific tropics from Samoa in the South Pacific to New Caledonia. The sectional clade has 1.0 LPP support in the phylogenomic analysis.

Species

*Heliconia indica Lam. (syn.: H. buccinata Roxb.); *H. lanata (Green) W.J.Kress; *H. laufao W.J.Kress; *H. paka A.C.Smith; +H. papuana W.J.Kress; *H. solomonensis W.J.Kress.

Distribution

Tropical Asia-Pacific (Fiji, Indonesia, New Caledonia, Papua New Guinea, Samoa, Solomon Islands, Vanuatu).

Heliconia section Perplexae W.J.Kress, sect. nov.

urn:lsid:ipni.org:names:77355025-1
Figs 3C, D

Type

Heliconia impudica Abalo & Morales.

Description and taxonomic notes

Medium-sized rhizomatous herbs with Musa-like habit. Inflorescence erect, with peduncle, rachis and cincinnal bracts variously colored with red and yellow; cincinnal bracts distichous or spirally arranged, horizontal to deflexed. Flowers with diurnal anthesis, resupinate; perianth uniformly curved, white to green to yellow, variously shaped; ovary green to yellow. Fruits blue, glabrous. This section includes four species native to Colombia and Ecuador, which were included in subgen. Stenochlamys sect. Lanea by earlier authors (e.g., Andersson 1992; Kress et al. 1999). According to the genomic data, a fifth species, H. brenneri Abalo & Morales, is also placed in this section. However, in this classification it is placed in subgen. Heliconia sect. Longae where it shares morphological traits with H. foreroi Abalo & Morales and H. attratensis Abalo & Morales with erect, red, distichous cincinnal bracts and non-resupinate flowers. Section Perplexae has 1.00 LPP support as a monophyletic group and 1.0 LPP support as sister to sect. Heliconiopsis in the phylogenomic analysis.

Species

*Heliconia gaiboriana Abalo & Morales; *H. impudica Abalo & Morales; *H. virginalis Abalo & Morales; *H. willisiana Abalo & Morales.

Distribution

Tropical Andean South America (Colombia, Ecuador).

Heliconia subgenus Stenochlamys Baker, Ann. Bot. 7: 190, 194–200. 1893.

Figs 3E–H

Type

Heliconia psittacorum L. f. (designated by L. Andersson, Opera Bot. 82: 22. 1985).

Description and taxonomic notes

Small- to medium-sized rhizomatous herbs with Musa-like (rarely Zingiber-like) habit. Inflorescence erect, with peduncle, rachis and cincinnal bracts of various colors from red to orange to yellow; cincinnal bracts distichous, generally long and tapering. Flowers with diurnal anthesis, partially to fully resupinate and held at right angles to bracts; perianth short-tubed, angular in cross-section, generally straight to slightly curved, free sepal generally only slightly curved, white to green to yellow, sometimes with green tips; ovary green to yellow to orange to red. Fruits blue, glabrous. The original Heliconia subgen. Stenochlamys of Baker included 17 species (note that some of these species are not recognized today). Andersson’s (1985a) subgen. Stenochlamys was much expanded into a large and complex taxonomic entity with six sections. In the present classification only 15 species are recognized in two sections (sects. Stenochlamys and Angustae) of the subgenus. Most of the remaining species are now found in subgen. Heliconia (sects. Lasia, Cannastrum, and Longiflorae). The monophyly of subgen. Stenochlamys has 0.96 LPP support in the molecular analysis and is sister to the remaining three subgenera of Heliconia.

Distribution

Tropical Central and South America (Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Honduras, Nicaragua, Panama, Paraguay, Peru, Suriname, Trinidad & Tobago, Venezuela).

Heliconia section Stenochlamys (Baker) K. Schum., Engler A, ed., Pflanzenr. IV. 45: 37. 1900.

Figs 3E, F

Heliconia subgen. Stenochlamys Baker, Ann. Bot. 7: 190, 194–200. 1893.

Heliconia section Proximochlamys L. Anderss., Opera Bot. 82: 73. 1985. Type: Heliconia densiflora Verlot.

Heliconia section Zingiberastrum L. Anderss., Opera Bot. 82: 100. 1985. Type: Heliconia hirsuta L. f.

Type

Heliconia psittacorum L. f. (designated by L. Andersson, Opera Bot. 82: 22. 1985).

Description and taxonomic notes

Small- to medium-sized rhizomatous herbs with Musa-like (rarely Zingiber-like) habit. Inflorescence erect, with peduncle, rachis and cincinnal bracts of various colors from red to orange to yellow; cincinnal bracts distichous, generally long and tapering, usually held erect. Flowers with diurnal anthesis, fully resupinate and held at right angles to bracts; perianth short-tubed, angular in cross-section, generally straight, free sepal generally only slightly curved, yellow to orange (sometimes green or white) with green tips; ovary green to yellow to orange to red. Fruits blue, glabrous. Six of the nine species currently placed in this section were included in Andersson’s sect. Stenochlamys (H. hirsuta L. f., H. densiflora Verlot and H. sylvestris (Gleason) L.B.Smith were placed elsewhere). In the current analysis H. hirsuta, despite its position in the molecular analysis as sister to sect. Heliconia, is placed in this section based on the resupinate flowers with green-colored tips and distichous cincinnal bracts. Similarly genomic data placed H. lourteigiae Emygdio & Santos in sect. Stenochlamys, but morphological traits more strongly ally this species with others in sect. Taeniostrobus. A LPP support of 0.99 demarcates sect. Stenochlamys in the present classification.

Species

*Heliconia acuminata L.C.Rich.; *H. brachyantha L. Anderss.; *H. densiflora Verlot; *H. hirsuta L. f. †; *H. psittacorum L. f.; *H. richardiana Miq.; *H. sylvestris (Gleason) L.B.Smith; *H. tarumaensis Barr.; +H. timothei L. Anderss.

Distribution

Tropical Central and South America (Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Honduras, Nicaragua, Panama, Paraguay, Peru, Suriname, Trinidad & Tobago, Venezuela).

Heliconia section Angustae W.J.Kress, sect. nov.

urn:lsid:ipni.org:names:77355026-1
Figs 3G, H

Type

Heliconia angusta Vell.

Description and taxonomic notes

Small to medium-sized rhizomatous herbs with Musa-like habit. Inflorescence erect, with peduncle, rachis and cincinnal bracts colored primarily red; cincinnal bracts distichous usually held erect at 60–90 degrees to horizontal. Flowers with diurnal anthesis, partially resupinate; perianth straight to uniformly curved, white to green to yellow, triangular in cross-section; ovary yellow to red. Fruits blue, glabrous. The six species united here in subgen. Stenochlamys sect. Angustae were formerly distributed across three sections by Andersson (1985a 1992; sects. Stenochalmys, Tortex [now Taeniostrobus], and Lasia). Section Angustae with 1.0 LPP support as a monophyletic group in the molecular analysis is sister to sect. Stenochlamys.

Species

*Heliconia angusta Vell.; +H. dasyantha Koch & Bouché; *H. farinosa Raddi; *H. laneana Barreiros; *H. sampaioana L. Emygdio; *H. velloziana L. Emygdio.

Distribution

Tropical Northern South America (Brazil, French Guiana, Suriname).

Heliconia subgenus Taeniostrobus (Kuntze) Griggs, Bull. Torrey Bot. Club 30: 643. 1903.

Figs 3I–L

Bihai section Taeniostrobus Kuntze, Revisio Generum Plantarum. Pars I: 684. 1891.

Type

Bihai imbricata Kuntze (Heliconia imbricata (Kuntze) Baker (designated by L. Andersson, Flora of Ecuador 22: 11. 1985.).

Description and taxonomic notes

Small- to large-sized rhizomatous herbs with Musa-, Canna-, or Zingiber-like habit. Inflorescence erect (rarely pendent), with peduncle, rachis and cincinnal bracts of various colors from pink to red to orange to yellow; cincinnal bracts distichous or spirally arranged. Flowers with diurnal anthesis, not resupinate and fully or partially enclosed in bracts or fully resupinate and held at right angles to bracts; perianth uniformly curved (rarely s-shaped), white to green to yellow; ovary green to yellow to red. Fruits blue, glabrous. subgenus Taeniostrobus in the current classification is made up of two sections (sects. Taeniostrobus and Aurantiacae) and includes species from four earlier designated subgenera (subgens. Taeniostrobus, Heliconia, Stenochlamys, and Griggsia), although the majority of species were included in Andersson’s subgen. Heliconia sect. Tortex. Few, if any, morphological traits are shared by all species across subgen. Taeniostrobus. The very high strong support (LPP = 1.0) in the phylogenomic analysis is primary evidence for uniting these species into a single monophyletic subgenus with two somewhat distinct sections.

Distribution

Tropical Mexico, Central, and South America (Belize, Brazil, Colombia, Costa Rica, Ecuador, El Salvador, French Guiana, Guatemala, Honduras, Mexico, Nicaragua, Panama, Peru, Suriname, Venezuela).

Heliconia section Taeniostrobus Kuntze

Figs 3I, J

Bihai section Taeniostrobus Kuntze, Revisio Generum Plantarum. Pars I: 684. 1891.

Heliconia section Tortex L. Anderss., Flora of Ecuador 22: 19. 1985. Type. Heliconia latispatha Benth.

Heliconia section Tenebria L. Anderss., Opera Bot. 111: 37. 1992: Type. Heliconia tenebrosa Macbr.

Type

Bihai imbricata Kuntze (Heliconia imbricata (Kuntze) Baker) (designated by L. Andersson, Flora of Ecuador 11: 19. 1985.).

Description and taxonomic notes

Medium- to large-sized rhizomatous herbs with Musa-like habit. Inflorescence erect (rarely pendent), with peduncle, rachis and cincinnal bracts of various colors from pink to red to orange to yellow; cincinnal bracts distichous or spirally arranged. Flowers with diurnal anthesis, not resupinate (or rarely so), and fully or partially enclosed in bracts; perianth uniformly curved (rarely s-shaped), white to green to yellow; ovary green to yellow to red. Fruits blue, glabrous. Species in this section can be quite variable in most reproductive features, although almost all have non-resupinate flowers enclosed in the cincinnal bracts. The 0.99 LPP support identifies this clade as distinct from its sister taxon sect. Aurantiacae. The genomic data place H. lourteigiae L. Emygdio & Santos in sect. Stenochlamys, however, the shape and orientation of the cincinnal bracts and non-resupinate, enclosed flowers suggest that this species is more properly placed in sect. Taeniostrobus.

Species

*Heliconia atropurpurea Daniels & Stiles; *H. barryana W.J.Kress; *H. beckneri R.R. Smith; *H. bella W.J.Kress; *H. bourgaeana O.G. Peters.; *H. champneiana Griggs; *H. clinophila R.R.Smith; *H. collinsiana Griggs; +H. cucullata W.J.Kress & L.Anderss.; +H. darienensis L.Anderss.; *H. faunorum W.J.Kress & L.Anderss.; *H. gracilis Daniels & Stiles; *H. ignescens Daniels & Stiles; *H. imbricata (Kuntze) Baker; *H. irrasa R.R.Smith; *H. lankesteri Standl.; *H. latispatha Benth.; *H. librata Griggs; *H. lindsayana W.J.Kress; *H. lophocarpa Daniels & Stiles; *H. lourteigiae L.Emygdio & Santos †; *H. lutea W.J.Kress; +H. monteverdensis Daniels & Stiles; +H. mooreana R.R.Smith; *H. nutans Woodson; *H. reticulata (Griggs) Winkl.; +H. rodriguezii Stiles; +H. sarapiquensis Daniels & Stiles; *H. secunda Daniels & Stiles; *H. spathocircinata Aristeg.; +H. tenebrosa Macbr.; *H. thomasiana W.J.Kress; *H. tortuosa Griggs; *H. umbrophila Daniels & Stiles.

Distribution

Tropical Mexico, Central, and South America (Belize, Brazil, Colombia, Costa Rica, Ecuador, El Salvador, French Guiana, Guatemala, Honduras, Mexico, Nicaragua, Panama, Peru, Suriname, Venezuela).

Heliconia section Aurantiacae W.J.Kress, sect. nov.

urn:lsid:ipni.org:names:77355027-1
Figs 3K, L

Type

Heliconia aurantiaca Ghiesbr. ex Lemaire.

Description and taxonomic notes

Small- to medium-sized rhizomatous herbs with Musa- to Zingiber-like habit. Inflorescence erect, with peduncle, rachis and cincinnal bracts red to orange; cincinnal bracts spirally arranged (rarely distichous). Flowers with diurnal anthesis, fully resupinate and held at right angles to bracts; perianth uniformly curved, green to yellow; ovary green to yellow. Fruits blue, glabrous. The five species united here in subgen. Taeniostrobus sect. Aurantiacae were formerly placed in two sections by Andersson (1992; sects. Lanea and Zingiberastrum [now Longiflorae]). The species in this section, with primarily spirally arranged bracts, resupinate flowers, and some with Zingiber-like leaf arrangement, are distinct from taxa in sect. Taeniostrobus and are supported in the molecular analysis with 0.92 LPP support as a monophyletic group sister to that section.

Species

*Heliconia adflexa (Griggs) Standl.; *H. aurantiaca Ghiesbr. ex Lemaire; *H. crassa Griggs; *H. schiedeana Kl.; *H. spissa Griggs.

Distribution

Tropical Mexico and Central America (Belize, Costa Rica, Guatemala, Honduras, Mexico, Nicaragua).

Heliconia subgenus Colubrosae W.J.Kress, subgen. nov.

urn:lsid:ipni.org:names:77355028-1
Figs 4A–D

Type

Heliconia dielsiana Loes.

Description and taxonomic notes

Medium- to large-sized rhizomatous herbs with Musa-like habit. Inflorescence pendent or erect, with peduncle, rachis, and cincinnal bracts usually entirely red, sometimes green distally (rarely yellow); cincinnal bracts distichous or spirally arranged, horizontal or reflexed, usually elongate and tapering. Flowers with diurnal anthesis, fully or partially enclosed in bracts or resupinate and held at right angles to bracts; perianth s-shaped (sigmoid) or c-shaped (sharply curved), free sepal extended or not and reflexed, fused sepals reflexed at tips or not, generally yellow, sometimes white to green; ovary yellow to white (rarely green). Fruits blue, glabrous. The numerous species placed here in subgen. Colubrosae were formerly included in either subgen. Griggsia (with pendent inflorescences) or subgen. Stenochlamys (with erect inflorescences) by both Andersson (1985a, 1992) and Kress et al. (1999), but these two subgenera were never closely associated. The 1.0 LPP support in the molecular analysis strongly unites the species in these two subgenera into a single monophyletic lineage.

Distribution

Tropical Central and South America (Bolivia, Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Guyana, Panama, Peru, Venezuela).

Heliconia section Colubrosae W.J.Kress, sect. nov.

urn:lsid:ipni.org:names:77355029-1
Figs 4A, B

Type

Heliconia dielsiana Loes.

Description and taxonomic notes

Medium- to large-sized rhizomatous herbs with Musa-like habit. Inflorescence pendent, with peduncle, rachis, and cincinnal bracts usually entirely red, sometimes green distally (rarely yellow); cincinnal bracts distichous or spirally arranged, horizontal or reflexed, usually elongate and tapering. Flowers with diurnal anthesis, fully or partially enclosed in bracts; perianth s-shaped (sigmoid), free sepal extended and reflexed, fused sepals reflexed at tips, generally yellow, sometimes white to green; ovary yellow to white. Fruits blue, glabrous. The species placed in sect. Colubrosae were formerly included in subgen. Griggsia by both Andersson (1992) and Kress et al. (1999) with the majority of species placed in either the informal H. trichocarpa group or H. obscura group by Andersson (1992) and the unpublished sects. Sigmoideae, Pendulae, Obscurae, and Dromadarius by Kress et al. (1999). Section Colubrosae is the largest group of species in the genus with pendent inflorescences and the 1.0 LPP support in the molecular analysis strongly unites monophyletic sect. Colubrosae sister to sect. Curvae. The genomic data place H. pendula Wawra in sect. Heliconia, however, the pendent inflorescence with spirally arranged bracts, extended and reflexed free sepals, and fused sepals reflexed at the tips suggest that this species is more properly placed in sect. Colubrosae.

Species

+Heliconia badilloi Abalo & Morales; +H. berriziana Abalo & Morales; +H. caquetensis Abalo & Morales; +H. chrysocraspeda Abalo & Morales; *H. colgantea R.R.Smith ex Daniels & Stiles; +H. combinata Abalo & Morales; *H. dielsiana Loes.; +H. estiletioides Abalo & Morales; +H. fernandezii Abalo & Morales; +H. fredberryana W.J.Kress; *H. huilensis Abalo & Morales; +H. intermedia Abalo & Morales; +H. laxa Abalo & Morales; +H. lentiginosa Abalo & Morales; *H. lozanoi Abalo & Morales; *H. maculata W.J.Kress; +H. mucilagina Abalo & Morales; *H. mutisiana Cuatrecasas; +H. nariniensis Abalo & Morales; *H. necrobracteata W.J.Kress; +H. nitida Abalo & Morales; *H. obscura Abalo & Morales; *H. obscuroides L. Anderss.; *H. oleosa Abalo & Morales; *H. pendula Wawra †; +H. peteriana Abalo & Morales; *H. reptans Abalo & Morales; *H. riopalenquensis Dodson & A. Gentry; +H. robertoi Abalo & Morales; *H. robusta Pax; *H. sclerotricha Abalo & Morales; +H. signa-hispanica Abalo & Morales; *H. talamancana Daniels & Stiles; *H. trichocarpa Daniels & Stiles; +H. villosa Kl.

Distribution

Tropical Central and South America (Bolivia, Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Guyana, Panama, Peru, Venezuela).

Heliconia section Curvae W.J.Kress, sect. nov.

urn:lsid:ipni.org:names:77355030-1
Figs 4C, D

Type

Heliconia burleana Abalo & Morales.

Description and taxonomic notes

Medium- to large-sized rhizomatous herbs with Musa-like habit. Inflorescence erect with peduncle, rachis, and cincinnal bracts entirely red or yellow; cincinnal bracts distichous or spirally arranged, generally horizontal, elongate and tapering. Flowers with diurnal anthesis, resupinate and held at right angles to bracts; perianth c-shaped (sharply curved), free sepal extended and reflexed, fused sepals not reflexed at tips, yellow, or white to green; ovary yellow or green. Fruits blue, glabrous. For the most part the species placed here in sect. Curvae were formerly included in subgen. Stenochlamys sect. Lanea by both Andersson (1985a, 1992) and Kress et al. (1999). One species (H. sanctae-martae L. Anderss.) was included in sect. Cannastrum by Kress et al. (1999). The 1.0 LPP support in the molecular analysis together with the floral features listed here unite these four species into a single monophyletic lineage at the sectional level sister to sect. Colubrosae.

Species

*Heliconia aristeguietae Abalo & Morales; *H. burleana Abalo & Morales; *H. gilbertiana Abalo & Morales; +H. sanctae-martae L. Anderss.

Distribution

Primarily Tropical Andean South America (Colombia, Ecuador, Panama, Peru).

Heliconia subgenus Heliconia

Figs 4E–L, 5A–L

Description and taxonomic notes

Medium- to large-sized rhizomatous herbs with Musa-, Canna-, or Zingiber-like habit. Inflorescence erect or pendent, with peduncle, rachis and cincinnal bracts of various colors from red to orange to yellow to green; cincinnal bracts distichous or spirally arranged, congested or widely separated. Flowers with diurnal anthesis, not resupinate and fully or partially enclosed in bracts or fully resupinate and held at right angles to bracts; perianth with short to elongate tube, straight to uniformly curved to s-shaped (sigmoid), white to green to yellow to pink to red, glabrous to sometimes hairy; ovary white to green to yellow to red, generally glabrous. Fruits blue, glabrous (rarely hirsute). The size and complexity of subgen. Heliconia has varied with author, including from four (Andersson 1992) to six (Kress et al. 1999) sections. The molecular data provide 0.9 LPP support for a newly defined subgen. Heliconia, which includes many taxa formerly placed in subgenera Stenochlamys and Griggsia, e.g., from sects. Lasia, Cannastrum, Zingiberastrum (now Longiflorae), and parts of Lanea, as well as the informal species groups and unpublished sects. Griggsia, Barbatae, Arcuatae, Longae, Rostratae, Pendulae, and Retiformes. The wide variance in morphological traits characterizing species in subgen. Heliconia in part accounts for this complex arrangement of taxa. In the current classification nine sections, each supported by 0.92–1.0 LPP, are recognized. Four sections comprise an unresolved polyphyletic assemblage within the subgenus while the other five sections form a monophyletic group with 1.0 LPP support.

Distribution

Tropical Mexico, Central America, South America, and the Caribbean (Argentina, Barbados, Belize, Bolivia, Brazil, Colombia, Costa Rica, Dominica, Dominican Republic, Ecuador, El Salvador, French Guiana, Grenada, Guadeloupe, Guatemala, Guyana, Haiti, Honduras, Jamaica, Martinique, Mexico, Montserrat, Nicaragua, Panama, Paraguay, Peru, Puerto Rico, Saba, St. Eustatius, St. Kitts & Nevis, St. Lucia, St. Vincent & the Grenadines, Suriname, Trinidad & Tobago, Venezuela).

Heliconia section Heliconia

Figs 4E, F

Description and taxonomic notes

Generally large-sized rhizomatous herbs with Musa-like habit. Inflorescence erect, with peduncle, rachis and cincinnal bracts of various colors from red to orange to yellow to green, margins often green; cincinnal bracts distichous, congested and usually overlapping. Flowers with diurnal anthesis, not resupinate and fully enclosed in bracts; perianth generally with elongate tube, c-shaped (sharply curved) to s-shaped (sigmoid), hidden within bract with only apex protruding, white with green apex, glabrous; ovary white, glabrous. Fruits blue, glabrous, protruding from bract on elongated pedicel at maturity. Two species were placed in the genomic analyses as basal taxa in the clade including sect. Heliconia. However, morphological features indicate that H. hirsuta L. f. is more properly including in sect. Stenochlamys and H. pendula Wawra in sect Colubrosae. The close relationship among the species in the current sect. Heliconia has long been recognized by all authors. The 1.0 LPP support for this monophyletic group in the molecular analysis confirms this observation.

Species

*Heliconia aurea Rodríguez; *H. bihai (L.) L.; *H. caribaea Lam.; *H. lennartiana W.J.Kress; *H. orthotricha L. Anderss.; *H. rodriguensis Aristeg.; *H. stricta Huber; *H. wagneriana O.G.Peters.

Distribution

Tropical Mexico, Central America, South America, and the Caribbean (Barbados, Belize, Bolivia, Brazil, Colombia, Costa Rica, Dominica, Dominican Republic, Ecuador, French Guiana, Grenada, Guadeloupe, Guatemala, Guyana, Haiti, Honduras, Jamaica, Martinique, Mexico, Montserrat, Nicaragua, Panama, Peru, Puerto Rico, Saba, St. Eustatius, St. Kitts & Nevis, St. Lucia, St. Vincent & the Grenadines, Suriname, Trinidad & Tobago, Venezuela).

Heliconia section Lasia L. Anderss., Opera Bot. 82: 78. 1985.

Figs 4G, H

Type

Heliconia velutina L. Anderss.

Description and taxonomic notes

Medium-sized rhizomatous herbs with Musa-like habit. Inflorescence erect, with peduncle and rachis usually yellow, cincinnal bracts red, often yellow near rachis, usually hirsute; cincinnal bracts distichous, widely spaced, elongate and tapering. Flowers with diurnal anthesis, fully exposed at maturity, resupinate and held at nearly right angles to bracts; perianth with short tube, straight to uniformly curved, yellow to green (rarely red), glabrous; ovary green to yellow, glabrous. Fruits blue, glabrous (rarely hirsute). sect. Lasia was placed in subgen. Stenochlamys originally by Andersson (1985a, 1992) and followed by Kress et al. (1999). In the present classification molecular data support its inclusion in subgen. Heliconia although few morphological traits support that placement and only two of the four species in the section were included in the analysis. The 1.0 LPP support clearly unites these two species in a monophyletic section, but its relationship to other sections in the subgenus is equivocal at this time.

Species

+Heliconia estherae Abalo & Morales; *H. julianii Barreiros; *H. lasiorachis L. Anderss.; +H. velutina L. Anderss.

Distribution

Tropical South America (Bolivia, Brazil, Colombia, Ecuador, Peru, Venezuela).

Heliconia section Cannastrum L. Anderss., Opera Bot. 82: 86. 1985.

Figs 4I, J

Type

Heliconia metallica Planch. & Linden ex. Hook.

Description and taxonomic notes

Medium- to large-sized rhizomatous herbs with Canna- to rarely Zingiber-like habit. Inflorescence erect, with peduncle, rachis and cincinnal bracts usually red (rarely yellow to green); cincinnal bracts distichous, generally separated. Flowers with diurnal anthesis, fully exposed at maturity, resupinate and held at right angles to bracts; perianth with short to medium-sized tube, straight to uniformly curved, yellow (rarely pink), sometimes with green towards apex, glabrous; ovary green to yellow to red, sometimes with green apex, glabrous. Fruits blue, glabrous. Section Cannastrum was placed in subgen. Stenochlamys originally by Andersson (1985a, 1992) and followed by Kress et al. (1999). In the present classification molecular data support its inclusion in subgen. Heliconia although few morphological traits support that placement. The 1.0 LPP support clearly unites the species in this section as a monophyletic group. The genomic data place one species, H. pardoi Abalo & Morales, in sect. Griggsia, but the vegetative, inflorescence and floral traits suggest its inclusion in sect. Cannastrum. The relationship of this section to others in the subgenus is equivocal at this time.

Species

+Heliconia berryi Abalo & Morales; *H. calatheaphylla Daniels & Stiles; *H. deflexa Daniels & Stiles; *H. golfodulcensis Daniels & Stiles; *H. mathiasiae Daniels & Stiles; *H. meridensis Kl.; *H. metallica Planch. & Linden ex. Hook.; +H. mincana Abalo & Morales; +H. montana Abalo & Morales; *H. osaensis Cuf.; *H. pardoi Abalo & Morales †; *H. vaginalis Benth.; +H. venusta Abalo & Morales; *H. wilsonii Daniels & Stiles.

Distribution

Tropical Mexico, Central America, and South America (Belize, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, El Salvador, Guatemala, Honduras, Mexico, Nicaragua, Panama, Peru, Venezuela).

Heliconia section Longiflorae W.J.Kress, sect. nov.

urn:lsid:ipni.org:names:77355031-1
Figs 4K, L

Type

Heliconia longiflora R.R.Smith.

Description and taxonomic notes

Small to medium-sized rhizomatous herbs with Zingiber-like habit. Inflorescence erect with peduncle and rachis yellow or green and cincinnal bracts red with orange to yellow towards rachis; cincinnal bracts distichous and widely separated. Flowers with diurnal anthesis, fully exposed at maturity, resupinate and held at right angles to bracts; perianth generally straight to variously curved, white to yellow to orange, sometimes green towards apex, glabrous; ovary white to yellow to orange with green apex, glabrous. Fruits blue, glabrous. The Zingiber-like habit of the shoots is a shared featured of the species in this section, which has 1.0 LPP support in the molecular analysis. Andersson (1985a) previously named this section “Zingiberastrum” and designated H. hirsuta as the type. However, H. hirsuta L. f. is placed in sect. Stenochlamys in the present classification and the section is renamed sect. Longiflorae with the designation of a new type.

Species

+Heliconia apparicioi Barreiros; *H. cordata L. Anderss.; +Heliconia ecuadoriensis (L. Anderss.) W.J.Kress, comb. et stat. nov. (Heliconia longiflora ssp. ecuadoriensis L. Anderss., Opera Bot. 82: 113. 1985. Type. Asplund 16457. Ecuador, Esmeraldas, Timbre [San Mateo], 24 May 1955 [S]); *H. longiflora R.R.Smith; *H. schumanniana Loes.; +H. tacarcunae L.Anderss.

Distribution

Tropical Central America and South America (Brazil, Colombia, Costa Rica, Ecuador, Nicaragua, Panama, Peru).

Heliconia section Lanea L. Anderss., Opera Bot. 82: 23, 30. 1985.

Figs 5A, B

Type

Heliconia lingulata Ruiz & Pav.

Description and taxonomic notes

Medium- to large-sized rhizomatous herbs with Musa-like habit. Inflorescence erect, with peduncle, rachis and cincinnal bracts of various colors from pink to red to yellow; cincinnal bracts distichous or spirally arranged, usually widely separated. Flowers with diurnal anthesis, fully exposed at maturity, resupinate and held at right angles to bracts; perianth generally with short to medium-sized tube, straight to uniformly curved, green to yellow, glabrous; ovary green to yellow to red, generally glabrous. Fruits blue, glabrous. Section Lanea includes species which were formerly placed in sects. Lanea or Cannastrum by Andersson (1985a, 1992). Here these six species comprise a monophyletic group that is strongly supported (LPP = 1.0) as sister to a lineage of species with pendent inflorescences which were formerly classified in subgen. Griggsia by Andersson (1992) and Kress et al. (1999).

Species

*Heliconia aemygdiana Burle-Marx; +H. fugax L. Anderss.; *H. gloriosa Abalo & Morales; *H. lingulata Ruiz & Pav.; *H. subulata Ruiz & Pav.; *H. zebrina Plowman, W.J.Kress & Kennedy.

Distribution

Tropical South America (Argentina, Bolivia, Brazil, Colombia, Ecuador, Paraguay, Peru, Suriname).

Heliconia section Griggsia (L. Anderss.) W.J.Kress, comb. et stat. nov.

urn:lsid:ipni.org:names:77355032-1
Figs 5C, D

Heliconia subgenus Griggsia L. Anderss., Flora of Ecuador 22: 42. 1985.

Type

Heliconia griggsiana L.B.Smith.

Description and taxonomic notes

Large-sized rhizomatous herbs with Musa-like habit. Inflorescence pendent, with peduncle, rachis and cincinnal bracts red to pink to green; cincinnal bracts distichous or sub-spirally arranged, partially separated. Flowers with diurnal anthesis, not resupinate and fully enclosed in bracts; perianth with short to medium-sized tube, uniformly curved, yellow to orange, glabrous; ovary white to red, glabrous. Fruits blue, glabrous. subgenus Griggsia was established by Andersson (1985b, 1992) to include all species of Heliconia with pendent inflorescences. Here it is recognized as a section with only three species (LPP = 1.0) and is clearly allied to the other three sections with pendent inflorescences in this subgenus. In the molecular analyses H. pardoi Abalo & Morales was indicated as allied to species in sect. Griggsia, but shares morphological features with taxa in sect. Cannastrum, where it is placed in this classification. A second species, H. titanum W.J.Kress & J. Betancur, was placed in sect. Longae by the genomic data, which may support an earlier suggestion (C. Black, pers. comm.) that this species is a hybrid with one parental species in that section. Here it is placed in sect. Griggsia.

Species

*Heliconia gigantea W.J.Kress & J. Betancur; *H. griggsiana L.B.Smith; *H. titanum W.J.Kress & J.Betancur †.

Distribution

Tropical Andean South America (Colombia, Ecuador).

Heliconia section Longae W.J.Kress, sect. nov.

urn:lsid:ipni.org:names:77355033-1
Figs 5E–H

Type

Heliconia longa (Griggs) Winkler.

Description and taxonomic notes

Medium- to large-sized rhizomatous herbs with Musa-like habit. Inflorescence generally pendent or occasionally erect, with peduncle, rachis and cincinnal bracts almost always red to rarely yellow; cincinnal bracts distichous or slowly spirally arranged, congested or widely separated. Flowers with diurnal anthesis, not resupinate and fully enclosed in bracts; perianth generally with medium-sized tube, uniformly curved to c-shaped to rarely s-shaped, yellow to white to rarely pink or red, glabrous to hairy; ovary white to yellow to red, generally glabrous. Fruits blue, glabrous. Section Longae is the second largest group of species with pendent inflorescences next to sect. Colubrosae. Species in this section have previously been informally included in the H. pogonantha, H. longa, and H. grigssiana groups by Andersson (1992) and scattered among seven unpublished sections (Arcuatae, Barbatae, Dromedarius, Obscurae, Longae, and Retiformes) by Kress et al. (1999). Three species in sect. Longae have erect inflorescences (formerly in the unpublished section Complanatae). One of these species, H. brenneri Abalo & Morales, was included in subgen. Heliconiopsis sect. Perplexae according to the genomic data, but is morphologically similar to the other two species with erect inflorescences in sect. Longae (H. foreroi Abalo & Morales and H. attratensis Abalo & Morales) sharing red, distichous cincinnal bracts, and non-resupinate flowers and is placed in this section. Tissue was not available for many of the species with pendent inflorescence in the current phylogenomic analyses and future investigations may shuffle some of these species between sect. Longae and sect. Colubrosae. Section Longae is closely allied by 0.9 LPP support to the other three sections in subgen. Heliconia characterized by pendent inflorescences (sects. Griggsia, Episcopales, and Pastazae).

Species

+Heliconia arrecta W.J.Kress & J. Betancur; *H. atratensis Abalo & Morales; *H. berguidoi R. Flores, C. Black & A. Ibáñez; *H. brenneri Abalo & Morales †; *H. carmelae Abalo & Morales; *H. curtispatha Peters.; *H. danielsiana W.J.Kress; +H. donstonea W.J.Kress & J. Betancur; *H. excelsa L. Anderss.; +H. foreroi Abalo & Morales; *H. fragilis Abalo & Morales; *H. harlingii L. Anderss.; +H. holmquistiana Abalo & Morales; *H. longa (Griggs) Winkler; +H. lutheri W.J.Kress; *H. magnifica W.J.Kress; *H. mariae Hook. f.; +H. markiana Abalo & Morales; *H. nigripraefixa Dodson & Gentry; +H. paludigena Abalo & Morales; *H. pogonantha Cuf.; *H. ramonensis Daniels & Stiles; +H. regalis L. Anderss.; *H. rhodantha Abalo & Morales; +H. samperiana W.J.Kress & J. Betancur; +H. sanctae-theresae Abalo & Morales; *H. spiralis Abalo & Morales; +H. stella-maris Abalo & Morales; *H. stilesii W.J.Kress; *H. terciopela W.J.Kress & J. Betancur; *H. vellerigera Poepp.; *H. xanthovillosa W.J.Kress.

Distribution

Tropical Central America and South America (Belize, Colombia, Costa Rica, Ecuador, Guatemala, Honduras, Nicaragua, Panama, Peru, Venezuela).

Heliconia section Episcopales L. Anderss., Opera Bot. 111: 34. 1992.

Figs 5I, J

Type

Heliconia episcopalis Vell.

Description and taxonomic notes

Medium- to large-sized rhizomatous herbs with Musa-like habit. Inflorescence pendent or erect, with peduncle and rachis generally red and cincinnal bracts red to pink with green towards tip; cincinnal bracts distichous or rarely spirally arranged, moderated separated (rarely congested). Flowers with diurnal anthesis, not resupinate and fully enclosed in bracts; perianth with medium-sized tube, uniformly curved, white to yellow, glabrous; ovary white to yellow, glabrous. Fruits blue, glabrous. Nearly all of the species in sect. Episcopales (earlier referred to as “sect. Rostratae ined.” by Kress et al. 1999) have been considered as closely related because of their shared pendent inflorescences and flower features. The placement in this group of H. episcopalis Vell. with an erect inflorescence and congested bracts suggests that this inflorescence type has an independent evolutionary origin in this lineage. The closest relatives of H. penduloides Loes. have always been puzzling, but the 0.97 LPP support clearly unites these six species into a monophyletic group.

Species

*Heliconia episcopalis Vell.; *H. juruana Loes.; *H. marginata (Griggs) Pittier; *H. penduloides Loes; *H. rostrata Ruiz & Pav.; *H. standleyi Macbr.

Distribution

Tropical Central and South America (Bolivia, Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Guyana, Panama, Peru, Trinidad & Tobago, Venezuela).

Heliconia section Pastazae W.J.Kress, sect. nov.

urn:lsid:ipni.org:names:77355034-1
Figs 5K, L

Type

Heliconia pastazae L. Anderss.

Description and taxonomic notes

Medium-sized rhizomatous herbs with Musa-like habit. Inflorescence pendent, with peduncle, rachis and cincinnal bracts usually red with yellow distally along margins (rarely completely yellow or green); cincinnal bracts sub-distichous to spirally arranged, separated. Flowers with diurnal anthesis, not resupinate and fully to partially enclosed in bracts; perianth with short to medium-sized tube, uniformly curved, yellow to green, glabrous; ovary white to green to yellow, glabrous. Fruits blue, glabrous. Most of the species in this section have been considered as related and informally placed in the H. platystachys group (Andersson 1985b, 1992) or the unpublished sect. Pendulae (Kress et al., 1999). The strong LPP support (0.92) for sect. Pastazae is further evidence for its recognition as a distinct monophyletic lineage within subgen. Heliconia.

Species

*Heliconia abaloi G. Morales; *H. chartacea Lane ex Barreiros; *H. pastazae L. Anderss.; *H. platystachys Baker; +H. rigida Abalo & Morales.

Distribution

Tropical Central America and South America (Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Guyana, Panama, Peru, Venezuela).

Conclusions

The phylogenomic patterns of species relationships revealed here coupled with the newly proposed classification can now be used to further our understanding of the ecological patterns and interactions of heliconias in nature. Moreover, the results can aid in enhancing the economic potential of these plants in the horticultural trade by identifying prospective hybridization partners as well as species with shared cultivation and ornamental traits for introduction into sustainable commerce. As more genomic data are added to analyses and field observations of morphological features are increased, the current classification of Heliconia will undoubtedly be modified and refined as all classifications have undergone in the past.

Acknowledgements

WJK would like to thank Lennart Andersson for his camaraderie in pursuing a classification of the genus and Julio Betancur, Tunty Echeverry, Harry Luther, Cheryl Roesel, Carla Black, and Barry Hammel for their enthusiasm in the field. The authors thank Ida Lopez for her unflinching organization skills with specimen data and John H. Wiersema for his invaluable help with and knowledge of nomenclature and the botanical code. We also thank Michael Bordelon, Carla Black, Colton Collins (Plant Group Hawai’i), staff at Waimea Botanical Gardens, The National Tropical Botanical Garden, and Lyon Arboretum at the University of Hawai’i, and the MO and US herbaria for providing samples for this study. They have all contributed to this new phylogeny and classification of Heliconia. Computational resources were provided by the Smithsonian Institution High Performance Computing Cluster (SI HPC – Hydra) (https://doi.org/10.25572/SIHPC).

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

Funding for this research was provided by a Peter Buck Postdoctoral Fellowship from the Smithsonian Institution to MMC, a Small Grant from the Smithsonian’s National Museum of Natural History, and a Global Genome Initiative Grant from the Smithsonian Institute for Biodiversity Genomics.

Author contributions

Conceptualization: WJK, TF, MMC. Data curation: TF, MMC. Formal analysis: TF, MMC. Funding acquisition: MMC, WJK. Project administration: WJK.

Author ORCIDs

W. John Kress https://orcid.org/0000-0002-0140-5267

Tomáš Fér https://orcid.org/0000-0002-0126-3684

Mónica M. Carlsen https://orcid.org/0000-0002-1663-0475

Data availability

All of the data that support the findings of this study are available in the main text or Supplementary Information.

References

  • Andersson L (1985a) Revision of Heliconia subgen. Stenochlamys (Musaceae-Helico­nioideae). Opera Botanica 82: 1–123.
  • Andersson L (1985b) 221. Musaceae. In: Harling G, Sparre B (Eds) Flora of Ecuador 22: 1–87.
  • Andersson L (1992) Revision of Heliconia subgen. Taeniostrobus and subgen. Heliconia (Musaceae-Heliconioideae). Opera Botanica 111: 1–98.
  • Berry F, Kress WJ (1991) Heliconia: An identification guide. Smithsonian Institution Press, Washington, DC.
  • Betancur J, Kress WJ (1995) Distribucion geografica y altitudinal del genero Heliconia (Heliconiaceae) en Colombia. In: Churchill SP, Balslev H, Forero E, Luteyn JL (Eds) Biodiversity and conservation of neotropical montane forests. New York Botanical Gardens, Bronx, 513–523.
  • Carlsen MM, Fér T, Schmickl R, Leong-Škorničková J, Newman M, Kress WJ (2018) Resolving the rapid plant radiation of early diverging lineages in the tropical Zingiberales: Pushing the limits of genomic data. Molecular Phylogenetics and Evolution 128: 55–68. https://doi.org/10.1016/j.ympev.2018.07.020
  • Doyle JJ, Doyle JL (1987) A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochemical Bulletin 19: 11–15.
  • Farr ER, Leussink JA, Stafleu FA (1979) Index nominum genericorum. Vol. 1. Bohn. Scheltema & Holkema, Utrecht.
  • Garcia-Robledo C, Kuprewicz EK, Staines CL, Kress WJ, Erwin TL (2013a) Using a comprehensive DNA barcode library to detect novel egg and larval host plant associations in a Cephaloleia rolled-leaf beetle (Coleoptera: Chrysomelidae). Biological Journal of the Linnean Society. Linnean Society of London 110(1): 189–198. https://doi.org/10.1111/bij.12115
  • Garcia-Robledo C, Erickson DL, Staines CL, Erwin TL, Kress WJ (2013b) Tropical plant-herbivore networks: Reconstructing species interactions using DNA barcodes. PLOS ONE 8(1): e52967. https://doi.org/10.1371/journal.pone.0052967
  • Griggs RF (1904) Two new species of American wild bananas with a revision of the generic name. Bulletin of the Torrey Botanical Club 31(8): 445–447. https://doi.org/10.2307/2478895
  • Iles WJD, Sass C, Lagomarsino L, Benson-Martin G, Driscoll H, Specht CD (2017) The phylogeny of Heliconia (Heliconiaceae) and the evolution of floral presentation. Molecular Phylogenetics and Evolution 117: 150–167. https://doi.org/10.1016/j.ympev.2016.12.001
  • Johnson MG, Gardner EM, Liu Y, Medina R, Goffinet B, Shaw AJ, Zerega NJC, Wickett NJ (2016) HybPiper: Extracting coding sequence and introns for phylogenetics from high-throughput sequencing reads using target enrichment. Applications in Plant Sciences 4(7): 1600016. https://doi.org/10.3732/apps.1600016
  • Katoh K, Standley DM (2013) MAFFT multiple sequence alignment software version 7: Improvements in performance and usability. Molecular Biology and Evolution 30(4): 772–780. https://doi.org/10.1093/molbev/mst010
  • Kress WJ (1984) Systematics of Central American Heliconia (Heliconiaceae) with pendent inflorescences. Journal of the Arnold Arboretum 65(4): 429–532. https://doi.org/10.5962/p.36697
  • Kress WJ (1990) Taxonomy of Old World Heliconia (Heliconiaceae). Allertonia 6: 1–58.
  • Kress WJ, Betancur J, Roesel CS, Echeverry CBE (1993) Lista de las heliconias de Colombia con cinco especies nuevas. Caldasia 17: 183–198.
  • Kress WJ, Betancur J, Echeverry BE (1999) Heliconias – Llamaradas de la Selva Colombiana. Cristina Uribe Editores, Bogotá, Colombia.
  • Kress WJ, Fér T, Carlsen MM, Kupnick G (2024) A Conservation Assessment and Phylogenomic Evaluation of Protection Status in the Tropical Genus Heliconia L. (Zingiberales: Heliconiaceae). Plants, People, Planet [accepted].
  • Kuntze O (1891) Revisio Generum Plantarum. Pars I. Arthur Felix, Leipzig.
  • Linnaeus C (1753) Species plantarum. Vol. 2. [1957 facsimile of ed. 1, Ray Society, London.]
  • Linnaeus C (1771) Mantissa plantarum. [1961 facsimile reproduction, J. Cramer, Weinheim.]
  • Miller P (1754) The gardeners dictionary. ed. 4. Vol. I. J. & F. Rivington, London.
  • Miller P (1768) The gardeners dictionary. ed. 8. Vol. I. J. & F. Rivington, London.
  • Miquel F (1859) Heliconiopsis. Flora Indiae Batavae 3: 590.
  • Petersen OG (1890) Musaceae. In: von Martius CFP, Eichler AW (Eds) Fl. Brasil. 3: 1–28.
  • Schumann K (1900) Musaceae. In: Engler A (Ed.) Pflanzenr. IV. 45. Wilhelm Engelmann, Leipzig, 1–45.
  • Temeles EJ, Rah YJ, Andicoechea J, Byanova KL, Giller GSJ, Stolk SB, Kress WJ (2012) Pollinator-mediated selection in a specialized hummingbird-Heliconia system in the Eastern Caribbean. Journal of Evolutionary Biology 26(2): 347–356. https://doi.org/10.1111/jeb.12053
  • Winkler H (1930) Musaceae. In: Engler A, Prantl K (Eds) Pflanzenfam. , 2nd ed. 15a, 505–541.
  • Zhang C, Rabiee M, Sayyari E, Mirarab S (2018) ASTRAL-III: Polynomial time species tree reconstruction from partially resolved gene trees. BMC Bioinformatics 19(S6): 153. https://doi.org/10.1186/s12859-018-2129-y

Supplementary materials

Supplementary material 1 

Specimen information, genomic sequence data, and accession numbers for the phylogenomic analysis

W. John Kress, Tomáš Fér, Mónica M. Carlsen

Data type: xlsx

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (56.83 kb)
Supplementary material 2 

Full phylogeny of Heliconia with all species included in the genomic analysis with Local Posterior Probability (LPP) support values indicated

W. John Kress, Tomáš Fér, Mónica M. Carlsen

Data type: pdf

Explanation note: The six species with gray branches represent significant conflicts in genomic and morphological evidence, with latter given priority in species placement in the classification (see text).

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (113.36 kb)
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