Molecular Phylogeny and Redefined Generic Limits of Calathea (Marantaceae)

Finn Borchsenius

Molecular Phylogeny and Redefined Generic Limits of Calathea (Marantaceae) Author(s): Finn Borchsenius, Luz Stella Suárez Suárez, and Linda M. Prince Source: Systematic Botany, 37(3):620-635. 2012. Published By: The American Society of Plant Taxonomists URL: http://www.bioone.org/doi/full/10.1600/036364412X648571 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Systematic Botany (2012), 37(3): pp. 620–635 © Copyright 2012 by the American Society of Plant Taxonomists DOI 10.1600/036364412X648571 Molecular Phylogeny and Redefined Generic Limits of Calathea (Marantaceae) Finn Borchsenius,1,5 Luz Stella Suárez Suárez,2,4 and Linda M. Prince3 1 Department of Bioscience, Aarhus University, Ny Munkegade 114, Building 1540, DK-8000 Aarhus C, Denmark. 2 Universidad Nacional de Colombia, Instituto de Ciencias Naturales, Ciudad Universitaria, entrada calle 53, edificio 425, Bogotá, Colombia. 3 Rancho Santa Ana Botanic Garden and Claremont Graduate University – Botany, 1500 North College Avenue, Claremont, California 91711-3157, U. S. A. 4 Current address: Universidad de Los Llanos, Departamento de Biologı́a, Km 12 vı́a Puerto López, Metá, Colombia. 5 Author for correspondence (finn.borchsenius@biology.au.dk) Communicating Editor: Allan J. Bornstein Abstract—Calathea, with an estimated 285 species, is the largest genus of Marantaceae and an important component of Neotropical herbaceous diversity. The genus is also of high importance for horticulture as species are cultivated for their showy, patterned leaves. Previous molecular phylogenetic studies indicated that the genus is polyphyletic, but have not provided a basis for redefining generic limits due to incomplete taxon sampling. To address this problem we analyzed DNA sequence data from three plastid markers (matK with flanking 30 trnK intron, trnL intron and trnL-trnF intergenic spacer) and one nuclear marker (ITS) under a maximum parsimony criterion for a large and representative taxon sample covering all previously proposed infrageneric entities, and representing the full range of morphological variation known in the genus. Our results confirm that Calathea is polyphyletic. One clade, including subgenus Calathea, the C. lanicaulis group, and the genus Sanblasia, is sister to a clade formed by Ischnosiphon and Pleiostachya. The genus Monotagma is placed as sister to this clade. The remaining species form a second strongly supported clade as sister to a clade containing these other genera. Based on these findings Calathea is recircumscribed in a narrow sense and Sanblasia is placed in synonymy. The genus Goeppertia is resurrected and redefined to include all members of the second Calathea clade. Morphological characters defining each genus are provided. A total of 246 new combinations are made. Keywords—Goeppertia, ITS, matK, maximum parsimony, Sanblasia, trnL-F region. would remain monophyletic if Ischnosiphon, Pleiostachya, and Sanblasia were segregated based on morphological interpretation. The character that delimits Calathea from these genera is a tri-ovulate ovary, considered by Andersson to be plesiomorphic relative to the uni-ovulate condition present in the other three genera. Phylogenetic analyses based on molecular data have indeed confirmed this suspicion (Andersson and Chase 2001; Prince and Kress 2006a, 2006b). In these studies, one group of Calathea species forms a strongly supported clade sister to Ischnosiphon and Pleiostachya. The genus Monotagma is placed as sister to this clade, but with weaker support. The remaining species of Calathea constitute a second, strongly supported clade as sister to a clade containing these other genera. The exact delimitation of the two Calathea clades and the number of species in each has not yet been established, but it is clear that the second one, including species from three of the four subgenera in Schumann’s infrageneric classification (see below), is the largest. A fifth genus of the Calathea clade, Sanblasia, has not been sampled in any phylogeny. It has inflorescences that are intermediate between Calathea and Ischnosiphon (Andersson 1984, 1998), but its exact relationship to these genera is unknown. Calathea presents considerable variation in characters, including shoot architecture, arrangement and structure of inflorescence bracts, length of cymule axes, structure and position of bracteoles, and the presence or absence of an outer staminode (Kennedy et al. 1988; Andersson and Chase 2001). Beginning with Körnicke (1858, 1862), this variation has inspired the establishment of a number of infrageneric classifications, often containing many of the same basic groups (Table 1). These have, however, never been tested systematically using cladistic methods. The most comprehensive classification was proposed by Schumann (1902) in his family monograph. Schumann took his starting point in Körnicke’s pioneering work, but also included classification elements adopted from later schemes proposed by Bentham Marantaceae, with approximately 550 species, is the second largest family in the Zingiberales. The family is particularly diverse in the Neotropics where it is represented by an estimated 450 species. Phylogenetic studies at the family level (Andersson and Chase 2001; Prince and Kress 2006a) have shown Neotropical diversity to primarily involve two large groups: the Maranta clade with nine genera and ca. 70 species is concentrated in southeastern Brazil; and the Calathea clade with five genera and ca. 370 species is distributed throughout the Neotropics but with highest diversity in northwestern South America. Within the Calathea clade, Calathea G. Mey. is by far the largest genus with an estimated 300 species (Andersson 1998). Govaerts and Kennedy (2012) list 285 accepted names. These numbers clearly make Calathea the largest genus in the family. The genus is also important for horticulture as indoor plants and in tropical garden landscaping due to their variegated leaves with spots or bands of white, orange, or red, and often bright purple underside. Several species have been used as model organisms for studies of reproductive ecology, plant physiology, and demography (e.g. Kennedy 1978; Horvitz and Schemske 2002; Claßen-Bockhoff and Heller 2008; Matlaga and Sternberg, 2009; Swenson 2009; Maron et al. 2010). Other genera of the Calathea clade are less diverse. Monotagma K. Schum. has 37 species distributed from Central America to Brazil (Hagberg and Eriksson 2011); Ischnosiphon Körn. has 35 species concentrated in the western Amazon and subAndean region (Andersson 1977); Pleiostachya K. Schum. and Sanblasia L. Andersson are both monotypic (Andersson 1998). Calathea was described by Meyer (1818) to accommodate a group of species formerly included in Maranta L. The name is derived from Greek “calathos,” which means basket or container, as Meyer noted that some species were used for this purpose by indigenous people of northern South America (Kennedy 1978). Prior to the advent of molecular phylogenetics, monophyly of the genus was generally accepted. However, Andersson (1981) questioned whether Calathea 620 2012] BORCHSENIUS ET AL.: PHYLOGENY OF CALATHEA 621 Table 1. Comparison of previously proposed infrageneric classifications of Calathea. Note that, in contrast to later authors, Körnicke (1858, 1862) did not assign a formal rank to his groups. Names placed on the same row refer to corresponding entities. Körnicke (1858, 1862) Bentham (1883) Petersen (1890) Schumann (1902) Eucalathea Section Eucalathea Section Distichae Subgenus Eucalathea Anguste vel brevissime spicatae Section Macropus Section Tubispatha Subgenus Macropus Pseudophrynium Section Pseudophrynium Section Scapifoliae Section Comosae Subgenus Pseudophrynium: - Series Scapifoliae - Series Comosae Grandiflorae Section Breviscapus Section Nudiscapae - Series Nudiscapae Section Rhizanthae - Series Rhizanthae Pseudophrynium, group Pusillae Genus Monostiche Section Microcephalum Section Monostiche [included in section Nudiscapae] Section Monostiche (1883) and Petersen (1890). He arranged the 104 known species of Calathea into four subgenera: Eucalathea Körn, Macropus Benth., Pseudophrynium Körn., and Microcephalum Benth. Subgenus Eucalathea included species with distichous inflorescence bracts and laterally compressed inflorescences; subgenus Macropus included those with distichous bracts and cylindrical inflorescences; subgenus Pseudophrynium included species with spirally arranged bracts and large inflorescences with more than five bracts; those with small inflorescences and five or less bracts were placed in subgenus Microcephalum. The largest subgenus, Pseudophrynium, was further subdivided into four series corresponding to sections established by Petersen: Scapifoliae Eichler ex Petersen with tall flowering shoots bearing one or more cauline leaves; Comosae Petersen with sterile bracts apically in the inflorescence; Nudiscapae Petersen without sterile bracts and lacking cauline leaves; and Rhizanthae Eichler ex Petersen with inflorescences borne on separate shoots. A fifth series, Polystachyae K. Schum., was added in print to accommodate a single Brazilian species, C. polystachya K. Schum., with multiple inflorescences arising from the leaf axils. Recent investigations have shown that this species is identical to Maranta purpurea S. Vieira and V. C. Souza (2008:137) described from Mato Grosso in Brazil (S. Vieira, pers. comm.). Section Polystachyae is therefore not considered in this paper. An updated version of Schumann’s classification was presented by Loesener (1930), who included 130 species. The most recent contribution to our understanding of the infrageneric relationships in Calathea was made by Kennedy et al. (1988), who divided 64 Ecuadorian species into eight groups. Four of these corresponded to sections established by Bentham or Petersen (Breviscapus, Calathea [as Eucalathea], Comosae, Microcephalum), while the rest were new informal groups: 1) the C. lanicaulis group with nine species resembling those of section Calathea, but with sessile or shortpeduncled inflorescences and spirally arranged bracts (versus distinctly peduncled with distichous bracts in section Calathea; 2) the C. marantifolia group, including six species from subgenus Pseudophrynium series Scapifoliae with terminal inflorescences on an elongated shoot bearing one to four cauline leaves and bracts often broader than long and ovate to depressed-ovate; 3) the C. ornata group, including six species from Schumann’s series Nudiscapae that, in their young Kennedy et al. (1988) Section Calathea Calathea lanicaulis group [included in sect. Comosae] - Series Polystachyae Subgenus Microcephalum Calathea marantifolia group Section Comosae Calathea capitata group Section Breviscapus Calathea ornata group [one treated species placed in the Calathea ornata group] [not treated] Section Microcephalum [included in series Comosae] [not treated] stage, have leaves with delicate white and purple dots and stripes that usually disappear in fully grown plants; and 4) the C. capitata group, a segregate of four species from section Comosae with sterile bracts on top of the inflorescences similar in shape and color to the fertile ones (versus dissimilar in section Comosae as circumscribed by Kennedy et al. 1988). The work of Kennedy et al. (1988) undoubtedly represents the best hypothesis for monophyletic species groups presented to date. It is nevertheless far from complete in terms of taxon coverage and it lacks a formal phylogenetic framework. Therefore, it can not be used directly for establishing new generic boundaries. Schumann’s subgenera and series appear in several cases to be artificial and also lack phylogenetic support (Andersson and Chase 2001). Prince and Kress (2006a) concluded that Calathea was polyphyletic, but did not establish a morphological basis for redefining generic boundaries. The existing knowledge is therefore insufficient to draw a clear limit between the two large clades of Calathea species that appear in phylogenetic analyses. In addition, the status of the genus Sanblasia is uncertain. To overcome these limitations we present a phylogenetic analysis of Calathea based on plastid and nuclear genome DNA markers, sequenced for a large and representative taxon sample covering all previously proposed infrageneric entities, and representing the full range of morphological variation known in the genus. We use the results to: 1) explore clade structure in Calathea in relation to morphological variation and previously proposed classification schemes, particularly those of Schumann (1902) and Kennedy et al. (1988); and 2) redefine generic boundaries for Calathea corresponding to the two large clades of Calathea species identified by previous studies, thereby resurrecting one previously recognized genus. Materials and Methods Taxon Sampling—Based on the infrageneric classification of Schumann (1902) and the eight species groups of Kennedy et al. (1988) we devised an even sampling of species across all classificatory entities proposed in these two treatments (Appendix 1). We selected five species from section Calathea and five from the Calathea lanicaulis group as delimited by Kennedy et al. (1988). Species from these two groups formed the first of two Calathea clades in the analysis of Prince and Kress (2006a). From section Microcephalum we also included five species. For the remaining 622 SYSTEMATIC BOTANY five species groups of Kennedy et al. (1988), which all belong to subgenus Pseudophrynium sensu Schumann, we included four species from each. From Schumann’s subgenus Macropus and his subgenus Pseudophrynium series Rhizanthae we included two species from each as this was the maximum number of samples available to us. However, the two groups consist of few species and we consider this sampling adequate. Finally, we included three species with unusual morphology and of uncertain affinity: 1) C. cyclophora Baker, placed by Schumann in subgenus Eucalathea, but considered by Kennedy (1995) to belong to section Breviscapus Benth. (that section was included in Pseudophrynium series Nudiscapae in Schumann’s classification); 2) C. rufibarba Fenzl, placed by Schumann in subgenus Pseudophrynium series Nudiscapae and not discussed by Kennedy et al. (1988); and 3) C. killipii L. B. Sm. & Idrobo, a white flowered Colombian species (L. S. Suárez, pers. obs.) described after the publication of Schumann’s work and with no clear affinity to any described infrageneric entity. The total number of Calathea species sampled was 42, corresponding to ca. 15% of the estimated 285 species in the genus. As noted above, Calathea in its current circumscription is polyphyletic with Ischnosiphon, Pleiostachya, Monotagma, and possibly Sanblasia as nested elements. We therefore included representatives of these genera among the ingroup taxa. From the available material we selected six samples of Ischnosiphon covering five of the seven sections recognized by Andersson (1977) in that genus. For Monotagma we selected three and four samples, respectively, from the two sections of that genus proposed by Hagberg and Eriksson (2011). Finally, Pleiostachya pruinosa (Regel) K. Schum. and Sanblasia dressleri Andersson, the sole species of their genera, were added, bringing the total number of ingroup samples to 57. The outgroup was assembled by selecting two to four species from each of the four other major clades of Marantaceae identified by Prince and Kress (2006a), plus a representative of the African genus Haumania J. Léonard. Previous studies are ambiguous with respect to resolution of relationships between the five major clades of Marantaceae and the position of Haumania. The latter has been resolved as sister to all other Marantaceae (Prince and Kress 2006b); as a basally diverging lineage in the Calathea clade (Prince and Kress 2006a); or as a member of the African Sarcophrynium clade (Prince and Kress 2006b). The position of the Calathea clade within the family phylogeny is also disputed. Prince and Kress (2006a, 2006b) resolved this clade in two different intermediate positions with no single major clade as sister group, while Andersson and Chase (2001) and Suksathan et al. (2009) recognized it as sister to the Asian Donax clade. Our outgroup sample includes representatives of all major clades in the family and would therefore be appropriate irrespective of which of these hypotheses is correct. Voucher information for all samples and GenBank accession numbers for all sequences are provided in Appendix 1. DNA Extraction, Amplification and Alignment—We analyzed data for four loci: the plastid matK gene and flanking 30 trnK intron; the trnL intron; the trnL-trnF intergenic spacer (trnL-F IGS); and the nuclear ITS region in the ribosomal RNA gene, including part of the 18S, through ITS1, 5.8S, ITS2, and part of the 26S loci in a single sequence. A number of sequences generated in previous studies (Prince and Kress 2006a, 2006b) were downloaded from GenBank (Appendix 1). For new samples, total genomic DNA was extracted using a DNeasy Plant minikit (Qiagen, Valencia, California) following the manufacturer’s protocol. Amplification of the matK-trnK intron was obtained using the primer pair matK-19F with trnK-2R (Steele and Vilgalys 1994; Molvray et al. 2000). If this approach was not successful then the mIF forward PCR primer of Prince and Kress (2006a) was used instead. For sequencing we also used four newly designed internal primers: matK-867F 50 -TGGAGTCTTTCTTTCTTGAGCGAA-30 ; matK-988R 50 -CTTTTCCTTGATAYCGAACATAATG; matK-1336F 50 -TTATCAGATTGTGATATTATYAATCGA-30 ; matK-1639R: 50 -AATATCRAAATACCAAATACGTTCT-30 . The trnL intron and trnL-F IGS were amplified using the primers of Taberlet et al. (1991). The ITS region was amplified using the method described by Prince and Kress (2006b). Amplification was made using either Ampliqon Taq DNA polymerase or PCR Beads (Pharmacia, St. Louis, Missouri) according to the manufacturer’s directions. All PCR products were checked by agarose gel electrophoresis. When multiple bands were detected, the target band was cut out of the gel and purified using QIAquick gel extraction kit (Qiagen). The PCR products were cleaned using QIAquick spin columns and sequenced using an AB 3130xl Genetic Analyzer (Applied Biosystems, Foster City, California) at Rancho Santa Ana Botanic Garden, or sent to Macrogen, Korea for sequencing. Cloning of the primary PCR-product was necessary in a few cases to obtain unambiguous sequences (matK, n = 3; trnL intron, n = 3; trnL-F IGS, n = 3; and ITS, n = 2). Cloning was performed using the TOPO cloning kit for sequencing (Invitrogen, [Volume 37 Carlsbad, California) following the manufacturer’s instructions. At least three randomly selected clones were sequenced in each case. Contig assembly and sequence editing were performed using SequencherÔ v.4.1 (Gene Codes Corp., Ann Arbor, Michican). Edited sequences were aligned initially using Muscle 3.6 (Edgar 2004) and manually adjusted. Indel information was coded using the simple method of Simmons and Ochoterena (2000) as implemented in the program FastGap 1.2 (Borchsenius 2009; www.aubot.dk/FastGap_home.htm). A single sequence was unavailable for study [trnL intron: outgroup taxon Ctenanthe setosa (Roscoe) Eichler]. This was replaced by missing data in the final dataset. The final data matrix is deposited in TreeBASE (study number S12469). Phylogenetic Analyses—All analyses shown and discussed in this paper are based on the maximum parsimony (MP) criterion in PAUP* 4.10.b (Swofford 2002). Previous phylogenetic studies of Marantaceae have shown that MP and likelihood based methods such as maximum likelihood or Bayesian analysis yield congruent topologies, but that MP tends to give less resolved trees and lower statistical support values (Prince and Kress 2006a, 2006b; Suksathan et al. 2009; Ley and ClaßenBockhoff 2011). Since the purpose of this study was to establish a robust systematic classification we considered a conservative approach appropriate. However, to assure that results do not depend on the choice of method we also performed a Bayesian analysis of the data (details in legend to Supplemental Figure S1). To address the issue of nonidentical clone sequences we first conducted preliminary analyses of the four loci separately including all clones. In all cases we found that non-identical clones from the same sample formed monophyletic groups or were part of the same polytomy in all MP trees. We then kept the clone with the shortest branch and pruned the remaining clones from the dataset as suggested by Beilstein et al. (2008). The argument for this approach is that the clone with the shortest branch will be most similar to the common ancestor of the clone group. For matK, the trnL intron, and the trnL-F IGS, one clone was pruned from each dataset. For ITS, three clones from two taxa were pruned. All ITS sequences in the final dataset were screened for three universally conserved motifs in the 5.8S coding region (Harpke and Peterson 2008) and no mutations were detected. We therefore infer that all ITS sequences represent functional paralogues. Initial analyses showed that topologies based on individual loci were congruent, differing only in the relationships among outgroup taxa (notably the placement of Haumania) and in the relative positions of some terminals within a stable set of major clades. However, an incongruence-length difference (ILD) test (Farris et al. 1995) performed in PAUP* (command HomPart) comparing combined plastid data with the nuclear ITS locus was significant (p < 0.001; heuristic search, 500 replicates, each search consisting of 10 replicates with random addition sequence in the starting tree, holding up to 10 trees in each round). For the final analysis we therefore combined data from all four loci, but calculated Partition Bremer Support (PBS) values (Baker and DeSalle 1997) using TreeRot v. 3 (Sorenson and Franzosa 2007; http://people .bu.edu/msoren/TreeRot.html) to identify conflicting signals in specific nodes. All MP searches of individual and combined data partitions were performed using 100 replicates of heuristic search with starting trees generated by random sequence addition, holding all shortest trees. Bootstrap support values for the combined data analysis were calculated from 1,000 replicates of heuristic search with starting trees generated by random sequence addition, each consisting of 10 replicates and holding up to 10 shortest trees in each round. Results The combined dataset included a total of 4,320 aligned nucleotide and 337 indel characters, of which 751 nucleotide (17%) and 152 indel characters (45%) were potentially parsimony informative. Sequence length of matK varied about 300 base pairs depending on which primer pair could be successfully applied. Potentially informative characters were nearly equally distributed among the combined plastid (435 characters) and nuclear partitions (468 characters). An MP search resulted in 27 trees of equal length (consistency index 0.58, retention index 0.78). The strict consensus tree (Fig. 1) was almost fully resolved with most ingroup nodes receiving > 80% Bootstrap Support (BS). The Calathea clade sensu Prince and Kress (2006a), excluding Haumania, was recovered as monophyletic with low BS (65%). Ischnosiphon, 2012] BORCHSENIUS ET AL.: PHYLOGENY OF CALATHEA 623 Fig. 1. Strict consensus of 27 shortest trees resulting from maximum parsimony analysis of combined data from chloroplast and nuclear markers. Numbers above branches denote bootstrap support. Numbers below branches denote partition Bremer support values for combined plastid loci and nuclear ITS, respectively. Inset upper left corner shows one of the shortest parsimony trees with branch lengths. An asterisk after the name of C. lutea marks that this is the type of Calathea. Pleiostachya, and Calathea species belonging to subgenus Calathea and the C. lanicaulis group formed a clade with 100% BS. Ischnosiphon was paraphyletic with respect to Pleiostachya, which was placed as sister to I. leucophaeus (Poepp. & Endl.) Körn. with 87% BS, and this species pair was sister to all other Ischnosiphon species sampled. Monotagma was resolved as monophyletic with 100% BS and placed as sister to the Ischnosiphon-Pleiostachya-Calathea clade. The latter relationship, however, had low BS (76%). Sanblasia formed a clade together with three members of subgenus Calathea. The clade was supported by 100% BS, but PBS revealed a strong conflict between plastid (–13) 624 SYSTEMATIC BOTANY and nuclear (+39) data for this node. The cause of conflict was the position of C. marantina (Willd. ex Körn.) K. Koch, which in analysis of plastid data alone was placed as sister to C. crotalifera S. Watson outside the clade including Sanblasia. Nuclear data supported the same topology as combined data. The remaining species of Calathea formed a clade with 90% BS (Calathea I, Fig. 1). Within this clade, C. straminea Petersen was placed as sister to all other taxa, which in turn formed five major clades (Fig. 2), each with 98–100% BS. Two clades corresponded to the C. ornata and C. marantifolia groups of Kennedy et al. (1988) (Fig. 2). One corresponded to subgenus Microcephalum of Schumann (1902), but also included a previously unplaced taxon, C. killipii. The last two clades corresponded roughly to series Comosae of Schumann (1902) and section Breviscapus of Kennedy et al. (1988), but also included a number of taxa from other Schumann categories. Calathea cyclophora, which Schumann placed in subgenus Calathea, was sister to all other species of the Comosae clade. Species of subgenus Macropus and subgenus Pseudophrynium series Rhizanthae of Schumann (1902) were included in the Breviscapus clade. Negative PBS values were detected at a few nodes for both data partitions (Fig. 1). The most important conflict concerns the nodes resolving C. villosa (Lodd. ex. G. Don) Lindl. and C. pavonii Körn. as basally diverging lineages within the Breviscapus clade. Plastid data place C. pavonii in a clade with C. varians and C. rufibarba, while [Volume 37 nuclear ITS data place C. villosa and C. pavonii as sister species with 100% BS in a more deeply nested position within the clade. Bayesian analysis yielded a tree identical to the MP strict consensus tree except for some minor differences (Supplemental Figure S1). Thus Calathea pavonii was placed as sister to C. rufibarba and C. varians (K. Koch & Mathieu) Körn. in a more deeply nested position in the Breviscapus clade and the resolution of species relationships in the Microcephalum clade was lower. Discussion Our results support the findings of earlier studies (Andersson and Chase 2001; Prince and Kress 2006a) that Calathea as currently circumscribed is polyphyletic. It should be redefined to include only the Calathea II clade (sensu Prince and Kress 2006a; Fig. 1) corresponding to the species of subgenus Calathea, the C. lanicaulis group (Kennedy et al. 1988), and Sanblasia dressleri. The type of the genus, Maranta casupo Jacq. (= Calathea lutea (Aubl.) E. Mey. ex Schult.), belongs to this clade and the type species was sampled in our study. Subgenus Calathea (sensu Schumann 1902) has traditionally been defined by having distichous bracts and usually laterally compressed inflorescences (Körnicke 1858; Bentham 1883; Schumann 1902), whereas the nine species in the C. lanicaulis group of Kennedy et al. (1988) usually have spirally arranged bracts and cylindrical inflorescences. No Fig. 2. Calathea clade I. Detail of strict consensus of 27 trees resulting from maximum parsimony analysis of combined data. Numbers above branches denote bootstrap support. Labels at right margin indicate classification of species according to Schumann (1902)/Kennedy et al. (1988). An asterisk after the label signifies that the taxon was not included in the Flora of Ecuador treatment by Kennedy et al. (1988), but placement in their classification was deduced with certainty from morphology. When this was not possible the taxon is labelled as unplaced and highlighted by gray typeface. 2012] BORCHSENIUS ET AL.: PHYLOGENY OF CALATHEA species of the latter group were known to Schumann and the combination of characters presented by these species thus cannot be accommodated anywhere in his classification. Inflorescence and flower morphology of Sanblasia have been described as intermediate between Calathea and Ischnosiphon (Andersson 1984). The genus has spirally arranged bracts that are narrow and involute as in Ischnosiphon and interphylls are missing, which is rarely the case in Calathea. On the other hand, flowers are tri-ovulate and subtended by channeled bracteoles, characters found only in Calathea. Andersson speculated that Sanblasia might be a “survivior of an experimenting calatheoid stock that gave rise to the more successful genus Ischnosiphon.” Our results suggest that it is a derived Calathea species that has evolved an unusual, Ischnosiphon-like inflorescence morphology. Based on literature descriptions and our own observations, a common feature for all three groups making up Calathea II, and a synapomorphy for the genus Calathea in a strict sense, are compound inflorescences that usually consist of two to several, similar, partial inflorescences that may have individual peduncles (Schumann’s subgenus Calathea) or are sessile and more or less congested (C. lanicaulis group and Sanblasia). Some species, however, have simple inflorescences (e.g. C. timothei H. A. Kenn.). Another synapomorphy of Calathea s. s. is that the corolla lobes are always reflexed or rolled back. Finally, most species are relatively large plants with tall flowering shoots and inflorescences placed well above the base of the plants. Based on these morphological criteria the number of currently accepted names in Calathea s. s. is 37. Our findings strongly support Calathea II as sister to Ischnosiphon and Pleiostachya. We find P. pruinosa K. Schum. in a position as sister to I. leucophaeus with 87% bootstrap support. Ischnosiphon leucophaeus is a member of section Papilioderma L. Andersson (1977) together with six other species. Andersson and Chase (2001) found two accessions of Pleiostachya to form a monophyletic group sister to a clade consisting of two species of section Papillioderma, but noted that inference was limited since they did not include any members of the other sections of Ischnosiphon. Prince and Kress (2006a) sampled five species of Ischnosiphon from four sections and did not find indications of a sister relationship between P. pruinosa and I. leucophaeus. Our results show some conflict between data partitions in the resolution of the Ischnosiphon-Pleiostachya clade. Nuclear ITS supports the topology found in the combined analysis. Plastid data alone (tree not shown) placed P. pruinosa in a clade with I. hirsutus Petersen, I. leucophaeus, and I. obliquus (Rudge) Körn., with only 69% bootstrap support. We conclude that further analysis is necessary to finally determine whether Pleiostachya should be included in Ischnosiphon or not. The position of Monotagma is also somewhat uncertain. In our strict consensus tree it is sister to the Calathea IIIschnosiphon-Pleiostachya clade, but support is low (76% BS). Similar low support values for this relationship have been found in previous phylogenetic studies (Prince and Kress 2006a, 2006b; Suksathan et al. 2009). One explanation might be that early diversification of the main lineages within the Calathea clade occurred rapidly. We did not find any indication that the African genus Haumania should be part of the Calathea clade as suggested by Prince and Kress (2006a). Instead it clusters with the two representatives of the likewise African Sarcophrynium clade [Megaphrynium macrostachyum 625 (K. Schum.) Milne-Redh. and Hypselodelphys hirsuta (Loes.) Koechlin; Fig. 1], albeit with no bootstrap support. A detailed survey of the structure and ontogeny of the hooded staminode in Marantaceae conducted by Pischtschan et al. (2010) suggested that one particular type of structure (“thumb type”) might be restricted to the Calathea II clade, Ischnosiphon, and Pleiostachya. However, included in their samples are three species that now, based on genetic and/or morphological data, clearly can be assigned to the Calathea I clade: C. microcephala (Poepp. & Endl.) Körn., sampled in this study; C. picturata K. Koch & Linden, placed in subgenus Macropus by Schumann (1902), but from its morphology and genetics clearly belongs to the Comosae clade (Schumann 1902; Andersson and Chase 2001; pers. obs.); and C. veitchiana Veitch ex Hook. f., likewise a member of section Comosae (Schumann 1902; pers. obs.). The thumb type hooded staminode, as defined by Pischtschan et al. (2010), thus seems to occur in the entire Calathea clade, possibly excluding Monotagma, which has not been investigated. Calathea I—The Calathea I clade (sensu Prince and Kress 2006a, Fig. 1) contains the largest number of species. It includes all of Schumann’s subgenera except subgenus Calathea (Fig. 2). Synapomorphies for this large clade include a simple inflorescence (with a few exceptions as discussed below) and corolla lobes straight to spreading, never reflexed or recurved as in Calathea II. Plants also tend to be smaller in size than those of Calathea II, though some members of the Ornata and Comosae clades can be large. Members of section Breviscapus sometimes have compound inflorescences composed of several, similar, partial inflorescences, like most species in Calathea II, but in that case the inflorescence bracts are always spirally arranged, never distichous, and the inflorescences are borne basally among the leaves, not in an elevated position on the flowering shoot. Characters that uniquely identify the two groups are not easy to define. They are, however, usually fairly easy to recognize in the field based on combined characters related to habit, inflorescence structure, and flower morphology. Within Calathea I our results reveal a clear and wellsupported clade structure showing a high degree of correspondence to previously proposed infrageneric groupings (Fig. 2). All species groups of Kennedy et al. (1988) are monophyletic. Less supported is Schumann’s (1902) classification. Subgenus Pseudophrynium series Nudiscapae is polyphyletic and its species are nearly equally divided between the Ornata clade and the Breviscapus clade. Subgenus Macropus is paraphyletic. Pseudophrynium series Rhizanthae is also polyphyletic. One species from that series, C. straminea, forms a basally diverging branch sister to all other species of Calathea I. The other species, C. varians, belongs to the Breviscapus clade. The position of Calathea straminea is interesting as the species has an unusual morphology. Schumann (1902) placed it in subgenus Pseudophrynium series Rhizanthae because its inflorescences are borne on separate, leafless shoots. He noted, however, that it is distinct from all other Calathea species due to its solitary, pedunculate flower pairs and rudimentary bracteoles, and considered that it might represent a separate genus. We believe it is better included in the same genus as the other species of Calathea I. Two other species, C. cannoides (Nicolson, Steyerm. and Sivad.) H. A. Kenn. and C. zingiberina Körn., have similar morphological characteristics (Kennedy 1990) and we expect that these will form part of the same clade as C. straminea. All 626 SYSTEMATIC BOTANY three taxa were considered members of the C. ornata group by Kennedy (1990). A characteristic feature of several clades in Calathea I is that they consist of a core group corresponding to a previously recognized classification system, but also some lineages with completely different morphologies (Fig. 2). For example, we find that C. killipii forms a basally diverging lineage in the Microcephalum clade. That species is a large, erect plant with a robust, turbinate inflorescence found in Panama and Colombia at elevations up to 1,600 m (L. S. Suárez, pers. obs.), and bears little morphological resemblance to other members of the clade. The only obviously shared character is white flower color. A similar result is the placement of C. cyclophora as sister to all other species of the Comosae clade with 98% BS and positive PBS values from both plastid and nuclear data partitions (Fig. 1). Calathea cyclophora does not possess the sterile, apical inflorescence bracts that characterize other Comosae species. Schumann (1902) considered C. cyclophora a member of Calathea subgenus Calathea as he erroneously interpreted the bracts as being distichous, while Kennedy (1995) considered it closely related to C. maasiorum H. A. Kenn., which she placed in section Breviscapus. A third example comes from our Breviscapus clade (Fig. 2). It contains a core group corresponding to section Breviscapus of Bentham (1883) and Kennedy et al. (1988), which are small plants with basal inflorescences. It also includes, however, species of subgenus Macropus and Pseudophrynium section Rhizanthae of Schumann (1902), with elongate flowering shoots and cylindrical inflorescences (e.g. C. pavonii, C. varians, and C. villosa). The presence of early diverging lineages with variable morphology nesting within otherwise homogeneous and species-rich clades is an interesting feature that could provide opportunities for gaining further insight into evolutionary pathways leading to successful radiations. Nees von Esenbeck (1831) described the genus Goeppertia based on a heterogeneous assemblage of taxa including Calathea, Maranta, and Monotagma species: Maranta zebrina Sims [= Calathea zebrina (Sims) Lindl.]; M. bicolor Ker Gawl [= M. cristata Nees & Mart.]; M. spicata Aubl. [= Monotagma spicatum (Aubl.) J. F. Macbr.]; and Goeppertia blanda Nees [= Calathea blanda (Nees) Steud.]. Kennedy (1978) lectotypified Goeppertia with Maranta zebrina, arguing that this taxon was the only one mentioned by Nees that lived up to his own diagnosis stating “Capsula trisperma.” Calathea zebrina is a well known species from Brazil, commonly cultivated and traded, which based on morphology (Sims 1817; S. Suárez pers. obs.) clearly belongs to our Breviscapus clade. It has an inflorescence of the same type as C. standleyi included in our study (Fig. 2). Schumann (1902) placed C. zebrina in Nudiscapae. Molecular data analyzed by Andersson and Chase (2001) placed it in an unresolved clade together with species from both our Scapifoliae and Breviscapus clades. Goeppertia is the oldest generic epithet attached to the Calathea I clade. We conclude that the name Goeppertia must be resurrected and applied to all species of that clade. An emended generic description is provided at the end of this paper. We have presented a well resolved and robust phylogeny of Calathea and related genera that allows us to redefine generic boundaries within the group. We recircumscribe Calathea to include the species formerly placed in subgenus Calathea, the C. lanicaulis group, and Sanblasia. The remaining species are transferred to a redefined Goeppertia. Our results [Volume 37 indicate only small conflicts between topologies derived from analysis of plastid and nuclear loci, and all major clades are supported by both data partitions. The clear and well supported resolution of major clades in Goeppertia provides a good basis for establishing a revised infrageneric classification of that genus. However, a few critical issues need to be addressed first. Some additional species with unclear affinities should be sampled for DNA analysis. An example is the species pair C. dicephala (Poepp. and Endl.) Körn. and C. pearcei Rusby. These were placed by Schumann (1902) in subgenus Pseudophrynium series Scapifoliae, but they have a different morphology from the remaining species of that group. We suspect they may instead be members of the Microcephalum clade. Finally, the morphological heterogeneity of several clades in Goeppertia represents a particular challenge in the search for synapomorphies that can be used to circumscribe revised infrageneric entities. Taxonomic Treatment 1. Calathea G. Mey. Prim. Fl. Esseq. 6. 1818.—TYPE: Maranta casupo Jacq. (lectotype: designated by S. Leman, Bull. Sci. Soc. Philom. Paris. 1820: 7. 1820) [= Calathea lutea (Aubl.) E. Mey. ex Schult.]. Sanblasia L. Andersson, Nordic J. Bot. 4: 21. 1984.—TYPE: Sanblasia dressleri L. Andersson. Herbs, usually with several basal and one or more cauline leaves. Leaves uniformly green or paler below, but never patterned. Inflorescence terminal on a leafy shoot, compound, usually composed of 2-several similar, partial inflorescences, rarely simple; bracts distichous or spirally arranged, persistent. Flower-groups brachyblastic, rarely sub-dolichoblastic, 2-, or rarely 3-flowered; interphylls usually present; bracteoles (0–)1–2(–4), membranaceous. Flowers open; corolla tube elongate, lobes reflexed or recurved; outer staminode 1 (rarely 0); ovary with 3 fertile locules. Fruit usually 3-seeded, dehiscent, obovoid, with persistent sepals; surface smooth; seeds with basal, white or rarely colored, aril. About 37 species from Mexico and the Carribean islands to Brazil and Bolivia. 2. Goeppertia Nees. Linnaea 6: 337. 1831.—TYPE: Maranta zebrina Sims (lectotype: designated by H. A. Kennedy, Univ. California Publ. Bot. 71: 32. 1978) [= Goeppertia zebrina (Sims) Nees]. Endocodon Raf., Fl. Tellur. 4: 49. 1838.—TYPE: Maranta zebrina Sims [= Goeppertia zebrina (Sims) Nees]. Psydaranta Neck. ex Raf., Fl. Tellur. 4: 53. 1838.—TYPE: Maranta comosa L. f. [= Goeppertia comosa (L. f.) Borchs. & S. Suárez]. Zelmira Raf., Fl. Tellur. 4: 50. 1838.—TYPE: Phrynium violaceum Roscoe [= Goeppertia violacea (Roscoe) Borchs. & S. Suárez]. Monostiche Körn, Gartenfl. 7: 88. 1858.—TYPE: Phrynium coloratum Hook. [= Goeppertia colorata (Hook.) Borchs. & S. Suárez]. Thymocarpus Nicolson, Steyerm. & Sivad., Brittonia 33: 22. 1981.—TYPE: Thymocarpus cannoides Nicolson, Steyerm. & Sivad. [= Goeppertia cannoides (Nicolson, Steyerm. & Sivad.) Borchs. & S. Suárez]. Herbs with basal and/or cauline leaves. Leaves uniformly green or with various types of patterning, this sometimes 2012] BORCHSENIUS ET AL.: PHYLOGENY OF CALATHEA present only in juvenile plants. Inflorescence terminal on a leafy shoot or more rarely borne on a separate leafless shoot, simple, or rarely compound and composed of up to 9 similar, partial inflorescences; bracts spirally arranged, persistent or soon decaying, the distal ones sometimes sterile. Flowergroups brachyblastic, rarely sub-dolichoblastic; 2- or rarely 3-flowered; interphylls usually present; bracteoles (0–)1–4, membranaceous or claviculate, the first two usually median and the remaining (if present) lateral. Flowers open or remaining closed, rarely self-fertile; corolla tube elongate, lobes straight or spreading; outer staminode 1 (rarely 0); ovary with 3 fertile locules. Fruit 3-seeded, dehiscent, obovoid or obpyramidal, usually with persistent sepals; surface smooth, muricate, or verrucose; seeds with basal white aril. About 248 species from Mexico and the Carribean islands to Argentina and Paraguay. Acknowledgments. Participation of Stella Suárez in this study was facilitated by a grant from the COIMBRA University group allowing her to stay and work in Aarhus for three months in 2008. Permits for collection and exportation of silica gel dried leaf samples of Colombian Marantaceae were kindly provided through the project “Marantaceae en la jurisdicción de Corantioquia,” financed by the Corporacion del Noroeste Antioqueño. The Amazonian Institute of Scientific Investigation – Sinchi, through the program “Inventarios florı́sticos en áreas estrategias de la Amazonia colombiana,” supplied us with valuable samples. We are also grateful to the Lyon Arboretum, in particular Karen E. Shigematsu, for allowing access to material of C. pavonii and C. varians. Carla Black, John Mood, Julio Betancur, Dairon Cárdenas, Rodrigo Bernal, Zaleth Cordero, Linda Milena Torres, Jaime Navarro, and Aida Vasco further supplied collections for this study. Gloria Galeano helped with advice and logistic support. 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GenBank numbers are listed in the following order: matK plus flanking 30 trnK intron; trnL intron; trnL-F IGS; ITS. Note that the name Calathea toroi S. Suárez, validly published in Caldasia 32: 296 (2010), has not been registered in the International Plant Names Index (IPNI; www.ipni.org) and is not listed by Govaerts and Kennedy (2012). Calathea— Calathea altissima (Poepp. & Endl.) Horan., S. Suarez 2367 (COL), JQ341333, JN413119, JQ341216, JQ341268. Calathea attenuata H. A. Kenn., S. Suárez 2609 (COL), JQ341335, JN413121, JQ341218, JQ341270. Calathea bella (W. Bull) Regel, W. J. Kress 02-7178 (US), AY140278, JN413143, AY140357, JQ341292. Calathea capitata (Ruiz & Pav.) Lindl., S. Suárez 2247 (COL), JQ341336, JN413122, JQ341219, JQ341271. Calathea crotalifera S. Watson, S. Suárez 2589 (COL), JQ341337, JN413123, JQ341220, JQ341272. Calathea curaraya H. A. Kenn., D. Cardenas 21047 (COL), JQ341338, JN413124, JQ341221, JQ341273. Calathea cyclophora Baker, D. Cardenas 18779 (COL), JQ341339, JN413125, JQ341222, JQ341274. Calathea ecuadoriana H. A. Kenn., W. J. Kress 01-6966 (US), AY140269, JN413126, AY140348, JQ341275. Calathea foliosa Rowlee ex Woodson & Schery, B.Hammel 11993 (DUKE), AY140270, JN413127, AY140349, JQ341276. Calathea fucata H. A. Kenn., S. Suárez 2627 (COL), JQ341340, JN413128, JQ341223, JQ341277. Calathea guzmanioides L. B. Sm. & Idrobo, S. Suárez 2322 (COL), JQ341341, JN413129, JQ341224, JQ341278. Calathea gymnocarpa H. A. Kenn., W. J. Kress 99-6402 (US), AY140271, JN413130, AY140350, JQ341279. Calathea hagbergii H. A. Kenn., S. Suárez 2325 (COL), JQ341342, JN413131, JQ341225, JQ341280. Calathea inocephala (Kuntze) T. Durand & B. D. Jacks., D. Cardenas 18688 (COL), JQ341343, JN413132, JQ341226, JQ341281. Calathea killipii L. B. Sm. & Idrobo, S. Suárez 2318 (COL), JQ341344, JN413133, JQ341227, JQ341282. Calathea lanata Petersen, L. M. Torres 139 (COL), JQ341345, JN413134, JQ341228, JQ341283. Calathea latifolia (Willd. ex Link) Klotzsch, S. Suárez 2406 (COL), JQ341346, JN413135, JQ341229, JQ341284. Calathea leonia (Sander) K. Schum., R. Bernal 3897 (COL), JQ341347, JN413136, JQ341230, JQ341285. Calathea loeseneri J. F. Macbr., W. J. Kress 99-6594 (US), AY140273, JN413137, AY140352, JQ341286. Calathea lutea (Aubl.) E. Mey. ex Schult., L. M. Torres 140 (COL), JQ341348, JN413138, JQ341231, JQ341287. Calathea marantina (Willd. ex Körn.) K. Koch, S. Suárez 2646 (COL), JQ341349, JN413139, JQ341232, JQ341288. Calathea micans (L. Mathieu) Körn., S. Suárez 2251 (COL), JQ341350, JN413140, JQ341233, JQ341289. Calathea microcephala (Poepp. & Endl.) Körn., S. Suárez 2642 (COL), JQ341351, JN413141, JQ341234, JQ341290. Calathea mishuyacu J. F. Macbr., D. Cardenas 21142 (COL), JQ341352, JN413142, JQ341235, JQ341291. Calathea pavonii Körn., Lyon Arboretum L-78.0725, voucher K. M. Nagata 3004 (HLA), JQ341353, JN413144, JQ341236, JQ341293. Calathea petersenii Eggers, S. Suárez 2652 (COL), JQ341354, JN413145, JQ341237, JQ341294. Calathea pluriplicata H. A. Kenn., W. J. Kress 99-6399 (US), AY140280, JN413146, AY140359, JQ341295. Calathea plurispicata H. A. Kenn., S. Suárez 2644 (COL), JQ341355, JN413147, JQ341238, JQ341296. Calathea propinqua (Poepp. & Endl.) Körn., S. Suárez 2662 (COL), JQ341356, JN413148, JQ341239, JQ341297. Calathea rufibarba Fenzl, W. J. Kress 01-6856 (US), AY140359, JN413149, AY140360, AY673048. Calathea silvosa J. F. Macbr., S. Suárez 2246 (COL), JQ341357, JN413150, JQ341240, JQ341298. Calathea standleyi J. F. Macbr., S. Suárez 2628 (COL), JQ341358, JN413151, JQ341241, JQ341299. Calathea straminea Petersen, S. Suárez 2673 (COL), JQ341359, JN413152, JQ341242, JQ341300. Calathea timothei H. A. Kenn., J. Betancur 12348 (COL), JQ341360, JN413153, JQ341243, JQ341301. Calathea toroi S. Suárez, S. Suárez 2317 (COL), JQ341334, JN413120, JQ341217, JQ341269. Calathea undulata (Linden & André) Linden & André, S. Suárez 2248 (COL), JQ341361, JN413154, JQ341244, JQ341302. Calathea utilis H. A. Kenn., W. J. Kress 99-6389 (US), AY140282, JN413155, AY140361, JQ341303. Calathea varians (K. Koch & Mathieu) Körn., Lyon Arboretum L-79.0444, photo voucher K. Shigematsu s. n. (AAU), JQ341362, JN413156, JQ341245, JQ341304. Calathea variegata (K. Koch) Linden ex Körn., S. Suárez 2647 (COL), JQ341363, JN413157, JQ341246, JQ341305. Calathea villosa (Lodd. ex G. Don) Lindl., R. Dressler 2912 (COL), JQ341364, JN413158, JQ341247, JQ341306. Calathea vinosa H. A. Kenn., W. J. Kress 77-0879 (DUKE), AY140284, JN413159, AY140363, JQ341307. Calathea warszewiczii (L. Mathieu ex Planch.) Planch. & Linden, L. Conde 8 (DUKE), AY140285, JN413160, AY140364, AY673049. Other ingroup taxa— Ischnosiphon hirsutus Petersen, S. Suárez 2664 (COL), JQ341365, JN413161, JQ341248, JQ341308. Ischnosiphon leucophaeus (Poepp. & Endl.) Körn., W. J. Kress 99-6377 (US), AY140299, JN413162, AY140380, JQ341309. Ischnosiphon macarenae L. Andersson, S. Suárez 2601 (COL), JQ341366, JN413163, JQ341249, JQ341310. Ischnosiphon obliquus [Volume 37 (Rudge) Körn., D. Cardenas 18695 (COL), JQ341367, JN413164, JQ341250, JQ341311. Ischnosiphon puberulus Loes., W. J. Kress 99-6383 (US), AY140300, JN413165, AY140381, JQ341312. Ischnosiphon rotundifolius (Poepp. & Endl.) Körn., W. J. Kress 99-6379 (US), AY140301, JN413166, AY140382, JQ341313. Monotagma juruanum Loes., S. Suárez 2656 (COL), JQ341368, JN413167, JQ341251, JQ341314. Monotagma laxum (Poepp. & Endl.) K. Schum., W. J. Kress 99-6381 (US), AY140309, JN413168, AY140392, AY673058. Monotagma papillosum Hagberg & R. Erikss., W. J. Kress 94-6411 (US), AY140310, JN413169, AY140393, JQ341315. Monotagma secundum (Petersen) K. Schum., D. Cardenas 18707 (COL), JQ341370, JN413171, JQ341253, JQ341317. Monotagma smaragdinum (Linden & André) K. Schum., W. J. Kress 99-6380 (US), AY140312, JN413172, AY140395, JQ341318. Monotagma tomentosum K. Schum. ex Loes., S. Suárez 2368 (COL), JQ341371, JN413173, JQ341254, JQ341319. Monotagma tuberosum Hagberg & R. Erikss., S. Suárez 2665 (COL), JQ341369, JN413170, JQ341252, JQ341316. Pleiostachya pruinosa K. Schum., Cultivated in Botanic Garden Mainz, voucher R. Classen-Bockhoff 5224 (MJG), JQ341372, JN413174, JQ341255, JQ341320. Sanblasia dressleri L. Andersson, C. Black 10 and 11 (US), JQ341373, JN413175, JQ341256, JQ341321. Outgroup— Ctenanthe setosa (Roscoe) Eichler, W. J. Kress 99-6385 (US), AY140288, —, AY140368, AY673051. Halopegia blumei (Körn.) K. Schum., P. Suksathan 3429 (AAU), JQ341323, JN413108, JQ341206, JQ341258. Haumania sp., D. J. Harris 6672 (E), AY140293, JN413109, AY140374, AY673053. Hypselodelphys hirsuta (Loes.) Koechlin, A. Ley 269 (WAG), JQ341324, JN413110, JQ341207, JQ341259. Maranta arundinacea L., S. Johannsen 21 (AAU), JQ341325, JN413111, JQ341208, JQ341260. Marantochloa filipes (Benth.) Hutch., A .Ley 262 (WAG), JQ341326, JN413112, JQ341209, JQ341261. Megaphrynium macrostachyum (K. Schum.) Milne-Redh., A. Ley 260 (WAG), JQ341327, JN413113, JQ341210, JQ341262. Phrynium interruptum (K. Schum.) Suksathan & Borchs., P. Suksathan 3409 (AAU, QBG), JQ341328, JN413114, JQ341211, JQ341263. Phrynium pubinerve Blume, F. Borchsenius 675 (AAU), JQ341329, JN413115, JQ341212, JQ341264. Schumannianthus dichotomus (Roxb.) Gagnep., F. Borchsenius 666 (AAU), JQ341330, JN413116, JQ341213, JQ341265. Stachyphrynium repens (Körn.) Suksathan & Borchs., F. Borchsenius 667 (AAU), JQ341331, JN413117, JQ341214, JQ341266. Thalia dealbata Fraser, F. Borchsenius 671 (AAU), JQ341332, JN413118, JQ341215, JQ341267. Appendix 2. New combinations of Goeppertia. Nomenclature and original reference citation for basionyms follow Govaerts and Kennedy (2012). Goeppertia ackermannii (Körn.) Borchs. & S. Suárez, comb. nov. Calathea ackermannii Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 133. 1862. Goeppertia acuminata (Steyerm.) Borchs. & S. Suárez, comb. nov. Calathea acuminata Steyerm., Fieldiana, Bot. 28: 161. 1951. Goeppertia aemula (Körn.) Borchs. & S. Suárez, comb. nov. Calathea aemula Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 131. 1862. Goeppertia affinis (Fenzl ex Regel) Borchs. & S. Suárez, comb. nov. Calathea affinis Fenzl ex Regel, Gartenflora 28: 294. 1879. Goeppertia albertii (Pynaert & Van Geert) Borchs. & S. Suárez, comb. nov. Maranta albertii Pynaert & Van Geert in ? Calathea albertii (Pynaert & Van Geert) L. H. Bailey & Raffill in L. H. Bailey, Stand. Cycl. Hort. 2: 621. 1914. Bailey and Rafill stated that their name was based on Maranta albertii Pynaert & Van Geert, but did not mention where that name had been published. IPNI does not include Maranta albertii or Calathea albertii. Govaerts and Kennedy (2012) do not provide a reference for the presumed basionym Maranta albertii either. Given this unusual situation we have chosen to follow the citation provided by Govaerts and Kennedy (2012), but further investigation is warranted. Goeppertia albovaginata (K. Koch & Linden) Borchs. & S. Suárez, comb. nov. Phrynium albovaginatum K. Koch & Linden, Wochenschr. Vereines Beförd. Gartenbaues Königl. Preuss. Staaten 8: 369. 1865. Calathea albovaginata 2012] BORCHSENIUS ET AL.: PHYLOGENY OF CALATHEA 629 (K. Koch & Linden) K. Schum. in H. G. A. Engler (ed.), Pflanzenr., IV, 48: 99. 1902. Goeppertia bellula (Linden) Borchs. & S. Suárez, comb. nov. Calathea bellula Linden, Cat. Gén. 89: 2. 1872. Goeppertia allenii (Woodson) Borchs. & S. Suárez, comb. nov. Calathea allenii Woodson, Ann. Missouri Bot. Gard. 29: 331. 1942. Goeppertia bracteosa (Rusby) Borchs. & S. Suárez, comb. nov. Calathea bracteosa Rusby, Mem. New York Bot. Gard. 7: 220. 1927. Goeppertia allouia (Aubl.) Borchs. & S. Suárez, comb. nov. Maranta allouia Aubl., Hist. Pl. Guiane 1: 3. 1775. Calathea allouia (Aubl.) Lindl., Edwards’s Bot. Reg. 14: t. 1210. 1829. Goeppertia brasiliensis (Körn.) Borchs. & S. Suárez, comb. nov. Calathea brasiliensis Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 118. 1862. Goeppertia altissima (Poepp. & Endl.) Borchs. & S. Suárez, comb. nov. Phrynium altissimum Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 20. 1837. Calathea altissima (Poepp. & Endl.) Horan., Prodr. Monogr. Scitam.: 13. 1862. Goeppertia brevipes (Körn.) Borchs. & S. Suárez, comb. nov. Calathea brevipes Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 140. 1862. Goeppertia angustifolia (Körn.) Borchs. & S. Suárez, comb. nov. Calathea angustifolia Körn., Gartenflora 7: 87. 1858. Goeppertia brunnescens (K. Koch) Borchs. & S. Suárez, comb. nov. Phrynium brunnescens K. Koch, Wochenschr. Vereines Beförd. Gartenbaues Königl. Preuss. Staaten 7: 277. 1864. Calathea brunnescens (K. Koch) K. Schum. in H. G. A. Engler (ed.), Pflanzenr., IV, 48: 99. 1902. Goeppertia annae (H. A. Kenn. & Marcelo) Borchs. & S. Suárez, comb. nov. Calathea annae H. A. Kenn. & Marcelo, Phytologia 82: 96. 1997. Goeppertia buchtienii (Pax) Borchs. & S. Suárez, comb. nov. Calathea buchtienii Pax, Repert. Spec. Nov. Regni Veg. 7: 107. 1909. Goeppertia applicata ( Jacob-Makoy ex E. Morren) Borchs. & S. Suárez, comb. nov. Calathea applicata Jacob-Makoy ex E. Morren, Ann. Hort. Belge Étrangère 24: 228. 1874. Goeppertia burle-marxii (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea burle-marxii H. A. Kenn., Canad. J. Bot. 60: 2365. 1982. Goeppertia argyraea (Körn.) Borchs. & S. Suárez, comb. nov. Calathea argyraea Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 142. 1862. Goeppertia cannoides (Nicolson, Steyerm. & Sivad.) Borchs. & S. Suárez, comb. nov. Thymocarpus cannoides Nicolson, Steyerm. & Sivad., Brittonia 33: 24. 1981. Calathea cannoides (Nicolson, Steyerm. & Sivad.) H. A. Kenn., Phytologia 69: 375. 1990. Goeppertia amazonica (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea amazonica H. A. Kenn., Selbyana 15: 63. 1994. Goeppertia argyrophylla (Linden ex K. Koch) Borchs. & S. Suárez, comb. nov. Maranta argyrophylla Linden ex K. Koch, Berliner Allg. Gartenzeitung 25: 243. 1857. Calathea argyrophylla (Linden ex K. Koch) L. H. Bailey & Raffill in L. H. Bailey, Stand. Cycl. Hort. 2: 624. 1914. Goeppertia arrabidae (Körn.) Borchs. & S. Suárez, comb. nov. Maranta tuberosa Vell., Fl. Flumin. 1: 4, t. 13. 1829. Calathea arrabidae Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 144. 1862. Körnicke (1862) made the combination Calathea tuberosa (Vell.) Körn. based on Thalia tuberosa Vell., Fl. Flumin. 1: 4, t. 18. 1829. [= Goeppertia tuberosa (Vell.) Borchs. & S. Suárez, see below). Therefore, a new name was assigned for Maranta tuberosa when transferred to Calathea. Goeppertia atropurpurea (Matuda) Borchs. & S. Suárez, comb. nov. Calathea atropurpurea Matuda, Anales Inst. Biol. Univ. Nac. México 27: 359. 1957. Goeppertia attenuata (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea attenuata H. A. Kenn., Nordic J. Bot. 6: 146. 1986. Goeppertia bachemiana (E. Morren) Borchs. & S. Suárez, comb. nov. Calathea bachemiana E. Morren, Ann. Bot. Hort. 25: 271. 1875. Goeppertia bantae (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea bantae H. A. Kenn., Canad. J. Bot. 64: 1321. 1986. Goeppertia baraquinii (Lem.) Borchs. & S. Suárez, comb. nov. Maranta baraquinii Lem., Ill. Hort. 15: t. 542. 1868. Calathea baraquinii (Lem.) Regel, Gartenflora 18: 99. 1869. Goeppertia capitata (Ruiz & Pav.) Borchs. & S. Suárez, comb. nov. Maranta capitata Ruiz & Pav., Fl. Peruv. 1: 3. 1798. Calathea capitata (Ruiz & Pav.) Lindl., Bot. Reg. 14: t. 1210. 1829. Goeppertia caquetensis (S. Suárez & Galeano) Borchs. & S. Suárez, comb. nov. Calathea caquetensis S. Suárez & Galeano, Caldasia 22: 9. 2000. Goeppertia cardenasii (Rusby) Borchs. & S. Suárez, comb. nov. Calathea cardenasii Rusby, Mem. New York Bot. Gard. 7: 222. 1927. Goeppertia cataractarum (K. Schum.) Borchs. & S. Suárez, comb. nov. Calathea cataractarum K. Schum. in H. G. A. Engler (ed.), Pflanzenr., IV, 48: 95. 1902. Goeppertia chimboracensis (Linden) Borchs. & S. Suárez, comb. nov. Maranta chimboracensis Linden, Ill. Hort. 17: t. 6. 1870. Calathea chimboracensis (Linden) Linden, Ill. Hort. 17: t. 6. 1870. Goeppertia chrysantha (Horan.) Borchs. & S. Suárez, comb. nov. Calathea chrysantha Horan., Prodr. Monogr. Scitam.: 13. 1862. Goeppertia chrysoleuca (Poepp. & Endl.) Borchs. & S. Suárez, comb. nov. Phrynium chrysoleucum Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 19. 1837. Calathea chrysoleuca (Poepp. & Endl.) Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 142. 1862. Goeppertia cinerea (Regel) Borchs. & S. Suárez, comb. nov. Calathea cinerea Regel, Gartenflora 25: 2. 1876. Goeppertia barbata (Petersen) Borchs. & S. Suárez, comb. nov. Calathea barbata Petersen in C. F. P. von Martius & auct. suc. (eds.), Fl. Bras. 3(3): 110. 1890. Goeppertia cleistantha (Standl.) Borchs. & S. Suárez, comb. nov. Calathea cleistantha Standl., J. Wash. Acad. Sci. 17: 250. 1927. Goeppertia bella (W. Bull) Borchs. & S. Suárez, comb. nov. Maranta bella W. Bull, Cat. 1875: 7. 1875. Calathea bella (W. Bull) Regel, Gartenflora 28: 297. 1879. Goeppertia clivorum (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea clivorum H. A. Kenn., Canad. J. Bot. 63: 1147. 1985. 630 SYSTEMATIC BOTANY [Volume 37 Goeppertia coccinea (Standl. & Steyerm.) Borchs. & S. Suárez, comb. nov. Calathea coccinea Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 39. 1944. Goeppertia divaricata (Rusby) Borchs. & S. Suárez, comb. nov. Calathea divaricata Rusby, Bull. Torrey Bot. Club 29: 695. 1902. Goeppertia colombiana (L. B. Sm. & Idrobo) Borchs. & S. Suárez, comb. nov. Calathea colombiana L. B. Sm. & Idrobo, Caldasia 5: 44. 1948. Goeppertia dodsonii (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea dodsonii H. A. Kenn., Selbyana 2: 46. 1977. Goeppertia colorata (Hook.) Borchs. & S. Suárez, comb. nov. Phrynium coloratum Hook., Bot. Mag. 57: t. 3010. 1830, comb. nov. Calathea colorata (Hook.) Benth. in G. Bentham & J. D. Hooker, Gen. Pl. 3: 654. 1883. Goeppertia communis (Wand. & S. Vieira) Borchs. & S. Suárez, comb. nov. Calathea communis Wand. & S. Vieira, Hoehnea 29: 115. 2002. Goeppertia comosa (L. f.) Borchs. & S. Suárez, comb. nov. Maranta comosa L. f., Suppl. Pl.: 80. 1782. Calathea comosa (L. f.) Lindl., Edwards’s Bot. Reg. 14: t. 1210. 1829. Goeppertia compacta (S. Suárez & Galeano) Borchs. & S. Suárez, comb. nov. Calathea compacta S. Suárez & Galeano, Caldasia 22: 12. 2000. Goeppertia donnell-smithii (K. Schum.) Borchs. & S. Suárez, comb. nov. Calathea donnell-smithii K. Schum. in H. G. A. Engler (ed.), Pflanzenr., IV, 48: 75. 1902. Goeppertia dressleri (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea dressleri H. A. Kenn., Ann. Missouri Bot. Gard. 60: 414. 1973. Goeppertia dryadica (J. M. A. Braga) Borchs. & S. Suárez, comb. nov. Calathea dryadica J. M. A. Braga, Kew Bull. 63: 309. 2008. Goeppertia eburnea (André & Linden) Borchs. & S. Suárez, comb. nov. Calathea eburnea André & Linden, Ill. Hort. 20: 171. 1873. Goeppertia ecuadoriana (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea ecuadoriana H. A. Kenn., Canad. J. Bot. 62: 15. 1984. Goeppertia concinna (W. Bull) Borchs. & S. Suárez, comb. nov. Maranta concinna W. Bull, Gard. Chron., n. s., 1: 78. 1874. Calathea concinna (W. Bull) K. Schum. in H. G. A. Engler (ed.), Pflanzenr., IV, 48: 119. 1902. Goeppertia eichleri (Petersen) Borchs. & S. Suárez, comb. nov. Calathea eichleri Petersen in C. F. P. von Martius & auct. suc. (eds.), Fl. Bras. 3(3): 108. 1890. Goeppertia concolor (Eichler ex Petersen) Borchs. & S. Suárez, comb. nov. Calathea concolor Eichler ex Petersen in C. F. P. von Martius & auct. suc. (eds.), Fl. Bras. 3(3): 126. 1890. Goeppertia elegans (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea elegans H. A. Kenn., Bot. Not. 131: 349. 1978. Goeppertia contrafenestra (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea contrafenestra H. A. Kenn., Canad. J. Bot. 62: 18. 1984. Goeppertia elliptica (Roscoe) Borchs. & S. Suárez, comb. nov. Phrynium ellipticum Roscoe, Monandr. Pl. Scitam.: t. 44. 1827. Calathea elliptica (Roscoe) K. Schum. in H. G. A. Engler (ed.), Pflanzenr., IV, 48: 75. 1902. Goeppertia coriacea (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea coriacea H. A. Kenn., Brenesia 14-15: 351. 1978. Goeppertia crocata (E. Morren & Joriss.) Borchs. & S. Suárez, comb. nov. Calathea crocata E. Morren & Joriss., Ann. Bot. Hort. 25: 141. 1875. Goeppertia cuneata (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea cuneata H. A. Kenn., Bot. Not. 129: 352. 1977. Goeppertia curaraya (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea curaraya H. A. Kenn., in Fl. Ecuador 32(224): 128. 1988. Goeppertia cyclophora (Baker) Borchs. & S. Suárez, comb. nov. Calathea cyclophora Baker, Bull. Misc. Inform. Kew 1895: 17. 1895. Goeppertia cylindrica (Roscoe) Borchs. & S. Suárez, comb. nov. Phrynium cylindricum Roscoe, Monandr. Pl. Scitam.: t. 40. 1828. Calathea cylindrica (Roscoe) K. Schum. in H. G. A. Engler (ed.), Pflanzenr., IV, 48: 83. 1902. Goeppertia densa (K. Koch & Linden) Borchs. & S. Suárez, comb. nov. Phrynium densum K. Koch & Linden, Ann. Hort. Belge Étrangère 15: 100. 1865. Calathea densa (K. Koch & Linden) Regel, Index Seminum (LE) 1866: 83. 1866. Goeppertia dicephala (Poepp. & Endl.) Borchs. & S. Suárez, comb. nov. Phrynium dicephalum Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 20. 1837. Calathea dicephala (Poepp. & Endl.) Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 127. 1862. Goeppertia dilabens (L. Andersson & H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea dilabens L. Andersson & H. A. Kenn., Nordic J. Bot. 6: 450. 1986. Goeppertia enclitica ( J. F. Macbr.) Borchs. & S. Suárez, comb. nov. Calathea enclitica J. F. Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 11: 53. 1931. Goeppertia erecta (L. Andersson & H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea erecta L. Andersson & H. A. Kenn., Nordic J. Bot. 6: 448. 1986. Goeppertia eximia (K. Koch & C. D. Bouché) Borchs. & S. Suárez, comb. nov. Phrynium eximium K. Koch & C. D. Bouché, Index Seminum (B) 1855(App.): 11. 1855. Calathea eximia (K. Koch & C. D. Bouché) Körn. ex Regel, Gartenflora 7: 87. 1858. Goeppertia exscapa (Poepp. & Endl.) Borchs. & S. Suárez, comb. nov. Phrynium exscapum Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 18. 1837. Calathea exscapa (Poepp. & Endl.) Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 122. 1862. Goeppertia exserta (Rusby) Borchs. & S. Suárez, comb. nov. Calathea exserta Rusby, Bull. New York Bot. Gard. 6: 495. 1910. Goeppertia fasciata (Linden ex K. Koch) Borchs. & S. Suárez, comb. nov. Maranta fasciata Linden ex K. Koch, Berliner Allg. Gartenzeitung 25: 243. 1857. Calathea fasciata (Linden ex K. Koch) Regel & Körn., Gartenflora 7: 348. 1858. Goeppertia fatimae (H. A. Kenn. & Marcelo) Borchs. & S. Suárez, comb. nov. Calathea fatimae H. A. Kenn. & Marcelo, Phytologia 82: 94. 1997. Goeppertia flavescens (Lindl.) Borchs. & S. Suárez, comb. nov. Calathea flavescens Lindl., Bot. Reg. 11: t. 932. 1825. 2012] BORCHSENIUS ET AL.: PHYLOGENY OF CALATHEA 631 Goeppertia foliosa (Rowlee ex Woodson & Schery) Borchs. & S. Suárez, comb. nov. Calathea foliosa Rowlee ex Woodson & Schery, Ann. Missouri Bot. Gard. 29: 332. 1942. Goeppertia inocephala (Kuntze) Borchs. & S. Suárez, comb. nov. Phyllodes inocephala Kuntze, Revis. Gen. Pl. 2: 694. 1891. Calathea inocephala (Kuntze) T. Durand & B. D. Jacks., Index Kew., Suppl. 1: 72. 1902. Goeppertia fragilis (Gleason) Borchs. & S. Suárez, comb. nov. Calathea fragilis Gleason, Bull. Torrey Bot. Club 56: 21. 1929. Goeppertia jagoriana (Regel) Borchs. & S. Suárez, comb. nov. Calathea jagoriana Regel, Gartenflora 28: 297. 1879. Goeppertia fucata (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea fucata H. A. Kenn., in Fl. Ecuador 32(224): 133. 1988. Goeppertia gandersii (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea gandersii H. A. Kenn., in Fl. Ecuador 32(224): 110. 1988. Goeppertia gardneri (Baker) Borchs. & S. Suárez, comb. nov. Calathea gardneri Baker, Bull. Misc. Inform. Kew 1895: 18. 1895. Goeppertia glaziovii (Benth.) Borchs. & S. Suárez, comb. nov. Calathea glaziovii Benth. in G. Bentham & J. D. Hooker, Gen. Pl. 3: 654. 1883. Goeppertia gloriana (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea gloriana H. A. Kenn., Selbyana 18: 35. 1997. Goeppertia grandis (Petersen) Borchs. & S. Suárez, comb. nov. Calathea grandis Petersen in C. F. P. von Martius & auct. suc. (eds.), Fl. Bras. 3(3): 124. 1890. Goeppertia granvillei (L. Andersson & H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea granvillei L. Andersson & H. A. Kenn., Nordic J. Bot. 6: 451. 1986. Goeppertia grazielae (H. A. Kenn. & Marcelo) Borchs. & S. Suárez, comb. nov. Calathea grazielae H. A. Kenn. & Marcelo, Phytologia 82: 101. 1997. Goeppertia guianensis (Klotzsch ex Benth. & Hook. f.) Borchs. & S. Suárez, comb. nov. Calathea guianensis Klotzsch ex Benth. & Hook. f., Gen. Pl. 3: 654. 1883. Goeppertia gymnocarpa (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea gymnocarpa H. A. Kenn., Bot. Not. 130: 333. 1977. Goeppertia jocosa ( J. F. Macbr.) Borchs. & S. Suárez, comb. nov. Calathea jocosa J. F. Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 11: 53. 1931. Goeppertia joffilyana ( J. M. A. Braga) Borchs. & S. Suárez, comb. nov. Calathea joffilyana J. M. A. Braga, Bradea 9: 1. 2002. Goeppertia kappleriana (Körn. ex Horan.) Borchs. & S. Suárez, comb. nov. Calathea kappleriana Körn. ex Horan., Prodr. Monogr. Scitam.: 12. 1862. Goeppertia killipii (L. B. Sm. & Idrobo) Borchs. & S. Suárez, comb. nov. Calathea killipii L. B. Sm. & Idrobo, Caldasia 5: 48. 1948. Goeppertia koernickeana (Horan.) Borchs. & S. Suárez, comb. nov. Calathea koernickeana Horan., Prodr. Monogr. Scitam.: 12. 1862. Goeppertia kummeriana (E. Morren) Borchs. & S. Suárez, comb. nov. Calathea kummeriana E. Morren, Ann. Bot. Hort. 25: 270. 1875. Goeppertia laetevirens (Huber) Borchs. & S. Suárez, comb. nov. Calathea laetevirens Huber, Bol. Mus. Goeldi Hist. Nat. Ethnogr. 4: 548. 1906. Goeppertia lagoagriana (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea lagoagriana H. A. Kenn., Nordic J. Bot. 6: 148. 1986. Goeppertia lanata (Petersen) Borchs. & S. Suárez, comb. nov. Calathea lanata Petersen in C. F. P. von Martius & auct. suc. (eds.), Fl. Bras. 3(3): 119. 1890. Goeppertia lancifolia (Boom) Borchs. & S. Suárez, comb. nov. Calathea insignis W. Bull, Cat. 1905: 2. 1905, nom. illeg. Calathea lancifolia Boom, Cat. 1905: 2. 1905. Goeppertia hammelii (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea hammelii H. A. Kenn., Selbyana 18: 33. 1997. Goeppertia lasiophylla (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea lasiophylla H. A. Kenn., Univ. Calif. Publ. Bot. 71: 63. 1978. Goeppertia hieroglyphica (Linden & André) Borchs. & S. Suárez, comb. nov. Calathea hieroglyphica Linden & André, Ill. Hort. 20: 63. 1873. Goeppertia lasseriana (Steyerm.) Borchs. & S. Suárez, comb. nov. Calathea lasseriana Steyerm., Fieldiana, Bot. 28: 163. 1951. Goeppertia hirta (Ravenna) Borchs. & S. Suárez, comb. nov. Calathea hirta Ravenna, Onira 9: 46. 2004. Goeppertia hopkinsii (Forzza) Borchs. & S. Suárez, comb. nov. Calathea hopkinsii Forzza, Rodriguésia 58: 535. 2007. Goeppertia humilis (S. Moore) Borchs. & S. Suárez, comb. nov. Calathea humilis S. Moore, Trans. Linn. Soc. London, Bot. 4: 489. 1895. Goeppertia hylaeanthoides (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea hylaeanthoides H. A. Kenn., Canad. J. Bot. 75: 1356. 1997. Goeppertia latifolia (Willd. ex Link) Borchs. & S. Suárez, comb. nov. Alpinia latifolia Willd. ex Link, Jahrb. Gewächsk. 1(3): 22. 1820. Calathea latifolia (Willd. ex Link) Klotzsch in R. H. Schomburgk, Reis. Br.-Guiana: 918. 1849. Goeppertia legrelleana (Linden) Borchs. & S. Suárez, comb. nov. Maranta legrelleana Linden, Ann. Hort. Belge Étrangère 17: 104. 1867. Calathea legrelleana (Linden) Regel, Gartenflora 28: 301. 1879. Goeppertia incompta (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea incompta H. A. Kenn., Canad. J. Bot. 75: 1361. 1997. Goeppertia leonia (Sander) Borchs. & S. Suárez, comb. nov. Maranta leonia Sander, Cat. 1896: 63. 1896. Calathea leonia (Sander) K. Schum. in H. G. A. Engler (ed.), Pflanzenr., IV, 48: 90. 1902. Goeppertia indecora (Woodson) Borchs. & S. Suárez, comb. nov. Calathea indecora Woodson, Ann. Missouri Bot. Gard. 29: 333. 1942. Goeppertia leonoriae (Lascur., H. Oliva & Avendaño) Borchs. & S. Suárez, comb. nov. Calathea leonoriae Lascur., H. Oliva & Avendaño, Novon 21: 66. 2011. 632 SYSTEMATIC BOTANY [Volume 37 Goeppertia leucostachys (Hook. f.) Borchs. & S. Suárez, comb. nov. Calathea leucostachys Hook. f., Bot. Mag. 101: t. 6205. 1875. Goeppertia mandioccae (Körn.) Borchs. & S. Suárez, comb. nov. Calathea mandioccae Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 138. 1862. Goeppertia libbyana (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea libbyana H. A. Kenn., Brittonia 36: 206. 1984. Goeppertia mansoi (Körn.) Borchs. & S. Suárez, comb. nov. Calathea mansoi Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 119. 1862. Goeppertia liesneri (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea liesneri H. A. Kenn., Novon 3: 49. 1993. Goeppertia marantifolia (Standl.) Borchs. & S. Suárez, comb. nov. Calathea marantifolia Standl., J. Wash. Acad. Sci. 17: 250. 1927. Goeppertia lindbergii (Petersen) Borchs. & S. Suárez, comb. nov. Calathea lindbergii Petersen in C. F. P. von Martius & auct. suc. (eds.), Fl. Bras. 3(3): 113. 1890. Goeppertia martinicensis (Urb.) Borchs. & S. Suárez, comb. nov. Calathea martinicensis Urb., Repert. Spec. Nov. Regni Veg. 15: 103. 1917. Goeppertia lindeniana (Wallis) Borchs. & S. Suárez, comb. nov. Calathea lindeniana Wallis, Ann. Hort. Belge Étrangère 16: 200. 1866. Goeppertia matudae (H. A. Kenn. & Ganders) Borchs. & S. Suárez, comb. nov. Calathea matudae H. A. Kenn. & Ganders, Novon 21: 59. 2011. Goeppertia lindmanii (K. Schum.) Borchs. & S. Suárez, comb. nov. Calathea lindmanii K. Schum. in H. G. A. Engler (ed.), Pflanzenr., IV, 48: 175. 1902. Goeppertia mediopicta (E. Morren) Borchs. & S. Suárez, comb. nov. Maranta mediopicta E. Morren, Ann. Bot. Hort. 25: 135. 1875. Calathea mediopicta (E. Morren) Jacob-Makoy ex E. Morren, Ann. Hort. Belge Étrangère 24: 228. 1874. Goeppertia littoralis (Ledeb. ex Sweet) Borchs. & S. Suárez, comb. nov. Phrynium littorale Ledeb. ex Sweet, Hort. Brit., ed. 3: 658. 1839. Calathea littoralis (Ledeb. ex Sweet) Körn., Gartenflora 7: 88. 1858. Goeppertia loeseneri ( J. F. Macbr.) Borchs. & S. Suárez, comb. nov. Calathea loeseneri J. F. Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 11: 51. 1931. Goeppertia longibracteata (Sweet) Borchs. & S. Suárez, comb. nov. Phrynium longibracteatum Sweet, Hort. Brit., ed. 2: 494. 1830. Calathea longibracteata (Sweet) Lindl., Bot. Reg. 12: t. 1020. 1827. Goeppertia longiflora (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea longiflora H. A. Kenn., Univ. Calif. Publ. Bot. 71: 76. 1978. Goeppertia longipetiolata (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea longipetiolata H. A. Kenn., Canad. J. Bot. 61: 1432. 1983. Goeppertia louisae (Gagnep.) Borchs. & S. Suárez, comb. nov. Calathea louisae Gagnep., Bull. Soc. Bot. France 55: xlii. 1908. Goeppertia lucianii (Linden) Borchs. & S. Suárez, comb. nov. Maranta lucianii Linden, Cat. Gén. 89: 2. 1872. Calathea lucianii (Linden) Cogn. & Marchal in A. Dallière, Pl. Ornam. Feuill. 2: t. 50. 1874. Goeppertia maasiorum (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea maasiorum H. A. Kenn., Brittonia 47: 156. 1995. Goeppertia macilenta (Lindl.) Borchs. & S. Suárez, comb. nov. Calathea macilenta Lindl., Edwards’s Bot. Reg. 14: t. 1210. 1829. Goeppertia macrosepala (K. Schum.) Borchs. & S. Suárez, comb. nov. Calathea macrosepala K. Schum. in H. G. A. Engler (ed.), Pflanzenr., IV, 48: 84. 1902. Goeppertia metallica (Planch. & Linden) Borchs. & S. Suárez, comb. nov. Calathea metallica Planch. & Linden, Cat. Gén. 10: 2. 1855. Goeppertia micans (L. Mathieu) Borchs. & S. Suárez, comb. nov. Maranta micans L. Mathieu, Prospectus 1853: s.p.. 1853. Calathea micans (L. Mathieu) Körn., Gartenflora 7: 87. 1858. Goeppertia microcephala (Poepp. & Endl.) Borchs. & S. Suárez, comb. nov. Phrynium microcephalum Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 20. 1837. Calathea microcephala (Poepp. & Endl.) Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 125. 1862. Goeppertia mirabilis ( Jacob-Makoy ex E. Morren) Borchs. & S. Suárez, comb. nov. Calathea mirabilis Jacob-Makoy ex E. Morren, Ann. Hort. Belge Étrangère 24: 228. 1874. Goeppertia misantlensis (Lascur.) Borchs. & S. Suárez, comb. nov. Calathea misantlensis Lascur., Novon 6: 385. 1996. Goeppertia mishuyacu ( J. F. Macbr.) Borchs. & S. Suárez, comb. nov. Calathea mishuyacu J. F. Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 11: 54. 1931. Goeppertia modesta (Brongn. ex Gris) Borchs. & S. Suárez, comb. nov. Calathea modesta Brongn. ex Gris, Ann. Sci. Nat., Bot., IV, 41: 193. 1859. Goeppertia monophylla (Vell.) Borchs. & S. Suárez, comb. nov. Maranta monophylla Vell., Fl. Flumin. 1: 4, t. 11. 1829. Calathea monophylla (Vell.) Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 144. 1862. Goeppertia multicincta (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea multicincta H. A. Kenn., Canad. J. Bot. 64: 1323. 1986. Goeppertia neblinensis (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea neblinensis H. A. Kenn., Phytologia 69: 373. 1990. Goeppertia majestica (Linden) Borchs. & S. Suárez, comb. nov. Maranta majestica Linden, Ann. Hort. Belge Étrangère 15: 103. 1865. Calathea majestica (Linden) H. A. Kenn., Canad. J. Bot. 64: 1325. 1986. Goeppertia neoviedii (Petersen) Borchs. & S. Suárez, comb. nov. Calathea neoviedii Petersen in C. F. P. von Martius & auct. suc. (eds.), Fl. Bras. 3(3): 117. 1890. Goeppertia makoyana (E. Morren) Borchs. & S. Suárez, comb. nov. Calathea makoyana E. Morren, Ann. Hort. Belge Étrangère 22: 321. 1872. Goeppertia nidulans (L. B. Sm. & Idrobo) Borchs. & S. Suárez, comb. nov. Calathea nidulans L. B. Sm. & Idrobo, Caldasia 5: 51. 1948. 2012] BORCHSENIUS ET AL.: PHYLOGENY OF CALATHEA Goeppertia nigricans (Gagnep.) Borchs. & S. Suárez, comb. nov. Calathea nigricans Gagnep., Bull. Soc. Bot. France 50: 588. 1903. Goeppertia nigrocostata (Linden & André) Borchs. & S. Suárez, comb. nov. Calathea nigrocostata Linden & André, Ill. Hort. 20: t. 144. 1873. Goeppertia nitens (Ender) Borchs. & S. Suárez, comb. nov. Calathea nitens Ender, Gartenflora 30: 180. 1881. Goeppertia nitidifolia (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea nitidifolia H. A. Kenn., Brenesia 14-15: 350. 1978. Goeppertia nobilis (K. Koch) Borchs. & S. Suárez, comb. nov. Phrynium nobile K. Koch, Berliner Allg. Gartenzeitung 25: 147. 1857. Calathea nobilis (K. Koch) Körn., Gartenflora 7: 88. 1858. Goeppertia oblonga (Mart.) Borchs. & S. Suárez, comb. nov. Phrynium oblongum Mart., Flora 24(2 Beibl.): 59. 1841. Calathea oblonga (Mart.) Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 138. 1862. Goeppertia orbifolia (Linden) Borchs. & S. Suárez, comb. nov. Maranta orbifolia Linden, Cat. Gén. 16: 2. 1861. Calathea orbifolia (Linden) H. A. Kenn., Brittonia 34: 22. 1982. 633 Goeppertia pearcei (Rusby) Borchs. & S. Suárez, comb. nov. Calathea pearcei Rusby, Mem. Torrey Bot. Club 6: 123. 1895. Goeppertia peruviana (Körn.) Borchs. & S. Suárez, comb. nov. Calathea peruviana Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 128. 1862. Goeppertia petersenii (Eggers) Borchs. & S. Suárez, comb. nov. Calathea petersenii Eggers, Bot. Centralbl. 53: 304. 1893. Goeppertia picturata (K. Koch & Linden) Borchs. & S. Suárez, comb. nov. Calathea picturata K. Koch & Linden, Wochenschr. Vereines Beförd. Gartenbaues Königl. Preuss. Staaten 6: 346. 1863. Goeppertia pilosa (Rusby) Borchs. & S. Suárez, comb. nov. Calathea pilosa Rusby, Bull. New York Bot. Gard. 6: 496. 1910. Goeppertia pittieri (K. Schum.) Borchs. & S. Suárez, comb. nov. Calathea pittieri K. Schum. in H. G. A. Engler (ed.), Pflanzenr., IV, 48: 108. 1902. Goeppertia plicata (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea plicata H. A. Kenn., Bot. Not. 129: 354. 1977. Goeppertia poeppigiana (Loes. ex H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea poeppigiana Loes. ex H. A. Kenn., in Fl. Ecuador 32(224): 77. 1988. Goeppertia ornata (Linden) Borchs. & S. Suárez, comb. nov. Maranta ornata Linden, Fl. Serres Jard. Eur. 4: t. 413. 1848. Calathea ornata (Linden) Körn., Gartenflora 7: 87. 1858. Goeppertia porphyrocaulis (W. Bull) Borchs. & S. Suárez, comb. nov. Maranta porphyrocaulis W. Bull, Cat. 110: 7. 1875. Calathea porphyrocaulis (W. Bull) N. E. Br., Suppl. Johnson’s Gard. Dict.: 890. 1882. Goeppertia ovandensis (Matuda) Borchs. & S. Suárez, comb. nov. Calathea ovandensis Matuda, Anales Inst. Biol. Univ. Nac. México 21: 333. 1951. Goeppertia portobelensis (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea portobelensis H. A. Kenn., Bot. Not. 128: 313. 1975 publ. 1976. Goeppertia ovata (Nees & Mart.) Borchs. & S. Suárez, comb. nov. Phrynium ovatum Nees & Mart., Nova Acta Phys.Med. Acad. Caes. Leop.-Carol. Nat. Cur. 11(1): 27. 1823. Calathea ovata (Nees & Mart.) Lindl., Edwards’s Bot. Reg. 14: t. 1210. 1829. Goeppertia praecox (S. Moore) Borchs. & S. Suárez, comb. nov. Calathea praecox S. Moore, Trans. Linn. Soc. London, Bot. 4: 488. 1895. Goeppertia pachystachya (Poepp. & Endl.) Borchs. & S. Suárez, comb. nov. Phrynium pachystachyum Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 19. 1837. Calathea pachystachya (Poepp. & Endl.) Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 142. 1862. Goeppertia pacifica (Linden & André) Borchs. & S. Suárez, comb. nov. Calathea pacifica Linden & André, Ill. Hort. 19: t. 101. 1872. Goeppertia pallidicosta (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea pallidicosta H. A. Kenn., Nordic J. Bot. 6: 143. 1986. Goeppertia panamensis (Rowlee ex Standl.) Borchs. & S. Suárez, comb. nov. Calathea panamensis Rowlee ex Standl., J. Wash. Acad. Sci. 15: 4. 1925. Goeppertia paucifolia (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea paucifolia H. A. Kenn., in Fl. Ecuador 32(224): 86. 1988. Goeppertia pavonii (Körn.) Borchs. & S. Suárez, comb. nov. Calathea pavonii Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 116. 1862. Goeppertia pavonina (K. Koch & Linden) Borchs. & S. Suárez, comb. nov. Maranta pavonina K. Koch & Linden, Ann. Hort. Belge Étrangère 15: 99. 1865. Calathea pavonina (K. Koch & Linden) Petersen in C. F. P. von Martius & auct. suc. (eds.), Fl. Bras. 3(3): 128. 1890. Goeppertia prolifera (Vell.) Borchs. & S. Suárez, comb. nov. Maranta prolifera Vell., Fl. Flumin. 1: 4, t. 10. 1829. Calathea prolifera (Vell.) J. M. A. Braga, Acta Bot. Bras. 19: 766. 2005. Goeppertia propinqua (Poepp. & Endl.) Borchs. & S. Suárez, comb. nov. Phrynium propinquum Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 18. 1837. Calathea propinqua (Poepp. & Endl.) Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 122. 1862. Goeppertia pruinata (W. Bull) Borchs. & S. Suárez, comb. nov. Maranta pruinata W. Bull., Cat. 72: 6. 1872. Calathea pruinata (W. Bull) N. E. Br., Suppl. Johnson’s Gard. Dict.: 890. 1882. Goeppertia pseudoveitchiana (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea pseudoveitchiana H. A. Kenn., Canad. J. Bot. 61: 1429. 1983. Goeppertia pulchella (E. Morren) Borchs. & S. Suárez, comb. nov. Maranta pulchella E. Morren, Ann. Bot. Hort. 25: 272. 1875. Calathea pulchella (E. Morren) Körn., Gartenflora 7: 87. 1858. Goeppertia pumila (Vell.) Borchs. & S. Suárez, comb. nov. Maranta pumila Vell., Fl. Flumin. 1: 3, t. 8. 1829. Calathea pumila (Vell.) Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 138. 1862. Goeppertia regalis (Rollison ex Lem.) Borchs. & S. Suárez, comb. nov. Maranta regalis Rollison ex Lem., Ill. Hort. 2: t. 74. 1855. Calathea regalis (Rollison ex Lem.) H. A. Kenn., Acta Hort. 413: 173. 1995. 634 SYSTEMATIC BOTANY [Volume 37 Goeppertia reginae (J. M. A. Braga) Borchs. & S. Suárez, comb. nov. Calathea reginae J. M. A. Braga, Kew Bull. 63: 311. 2008. Goeppertia sousandradeana (H. A. Kenn. & Ganders) Borchs. & S. Suárez, comb. nov. Calathea sousandradeana H. A. Kenn. & Ganders, Novon 21: 63. 2011. Goeppertia robin-fosteri (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea robin-fosteri H. A. Kenn., Bot. Not. 128: 316. 1975 publ. 1976. Goeppertia sophiae (Huber) Borchs. & S. Suárez, comb. nov. Calathea sophiae Huber, Bol. Mus. Goeldi Hist. Nat. Ethnogr. 4: 550. 1906. Goeppertia robiniae (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea robiniae H. A. Kenn., Ann. Missouri Bot. Gard. 60: 419. 1973. Goeppertia sphaerocephala (K. Schum.) Borchs. & S. Suárez, comb. nov. Calathea sphaerocephala K. Schum. in H. G. A. Engler (ed.), Pflanzenr., IV, 48: 101. 1902. Goeppertia rodeckiana (K. Schum.) Borchs. & S. Suárez, comb. nov. Calathea rodeckiana K. Schum. in H. G. A. Engler (ed.), Pflanzenr., IV, 48: 115. 1902. Goeppertia splendida (Lem.) Borchs. & S. Suárez, comb. nov. Maranta splendida Lem., Gard. Chron. 1864: 414. 1864. Calathea splendida (Lem.) Regel, Gartenflora 18: 99. 1869. Goeppertia roseobracteata (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea roseobracteata H. A. Kenn., Nordic J. Bot. 6: 459. 1986. Goeppertia sprucei (Rusby) Borchs. & S. Suárez, comb. nov. Calathea sprucei Rusby, Bull. New York Bot. Gard. 6: 495. 1910. Goeppertia roseopicta (Linden) Borchs. & S. Suárez, comb. nov. Maranta roseopicta Linden, Ann. Hort. Belge Étrangère 16: 202. 1866. Calathea roseopicta (Linden) Regel, Index Seminum (LE) 1869: 12. 1869. Goeppertia squarrosa (L. Andersson & H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea squarrosa L. Andersson & H. A. Kenn., Nordic J. Bot. 6: 454. 1986. Goeppertia rossii (Lodd. ex Sweet) Borchs. & S. Suárez, comb. nov. Phrynium rossii Lodd. ex Sweet, Hort. Brit., ed. 3: 658. 1839. Calathea rossii (Lodd. ex Sweet) Körn., Gartenflora 7: 88. 1858. Goeppertia rufibarba (Fenzl) Borchs. & S. Suárez, comb. nov. Calathea rufibarba Fenzl, Gartenflora 28: 294. 1879. Goeppertia rusbyi (Loes.) Borchs. & S. Suárez, comb. nov. Calathea nigricans Rusby, Bull. New York Bot. Gard. 6: 496. 1910, nom. illeg. Calathea rusbyi Loes. in H.G.A. Engler, Nat. Pflanzenfam. ed. 2, 15a: 678. 1930. Goeppertia standleyi ( J. F. Macbr.) Borchs. & S. Suárez, comb. nov. Calathea standleyi J. F. Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 11: 54. 1931. Goeppertia stenostachys (Rusby) Borchs. & S. Suárez, comb. nov. Calathea stenostachys Rusby, Mem. New York Bot. Gard. 7: 221. 1927. Goeppertia steyermarkii (H. A. Kenn. & Nagata) Borchs. & S. Suárez, comb. nov. Calathea steyermarkii H. A. Kenn. & Nagata, Brittonia 41: 164. 1989. Goeppertia straminea (Petersen) Borchs. & S. Suárez, comb. nov. Calathea straminea Petersen in C. F. P. von Martius & auct. suc. (eds.), Fl. Bras. 3(3): 118. 1890. Goeppertia sanderiana (Sander) Borchs. & S. Suárez, comb. nov. Maranta sanderiana Sander, Cat. (Sander & Co.) 1894: 9. 1894. Calathea sanderiana (Sander) Gentil, Pl. Cult. Serres Jard. Bot. Brux.: 43. 1907. Goeppertia stromanthifolia (Rusby) Borchs. & S. Suárez, comb. nov. Calathea stromanthifolia Rusby, Bull. New York Bot. Gard. 4: 456. 1907. Goeppertia saxicola (Hoehne) Borchs. & S. Suárez, comb. nov. Calathea saxicola Hoehne, Relat. Commiss. Linhas Telegr. Estratég. Matto Grosso Amazonas 5: 24. 1915. Goeppertia subtilis (S. Moore) Borchs. & S. Suárez, comb. nov. Calathea subtilis S. Moore, Trans. Linn. Soc. London, Bot. 4: 487. 1895. Goeppertia schunkei (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea schunkei H. A. Kenn., Brittonia 34: 18. 1982. Goeppertia taeniosa ( Joriss.) Borchs. & S. Suárez, comb. nov. Calathea taeniosa Joriss., Ann. Bot. Hort. 26: 83. 1876. Goeppertia sciuroides (Petersen) Borchs. & S. Suárez, comb. nov. Calathea sciuroides Petersen, Vidensk. Meddel. Naturhist. Foren. Kjøbenhavn 1889: 329. 1889. Goeppertia tinalandia (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea tinalandia H. A. Kenn., Canad. J. Bot. 63: 1141. 1985. Goeppertia selbyana (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea selbyana H. A. Kenn., in Fl. Ecuador 32(224): 63. 1988. Goeppertia trichoneura (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea trichoneura H. A. Kenn., Univ. Calif. Publ. Bot. 71: 58. 1978. Goeppertia sellowii (Körn.) Borchs. & S. Suárez, comb. nov. Calathea sellowii Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 120. 1862. Goeppertia truncata (Link ex A. Dietr.) Borchs. & S. Suárez, comb. nov. Maranta truncata Link ex A. Dietr., Sp. Pl. 1: 26. 1831. Calathea truncata (Link ex A. Dietr.) K. Schum. in H. G. A. Engler (ed.), Pflanzenr., IV, 48: 104. 1902. Goeppertia silvicola (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea silvicola H. A. Kenn., Univ. Calif. Publ. Bot. 71: 67. 1978. Goeppertia silvosa ( J. F. Macbr.) Borchs. & S. Suárez, comb. nov. Calathea silvosa J. F. Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 11: 52. 1931. Goeppertia tuberosa (Vell.) Borchs. & S. Suárez, comb. nov. Thalia tuberosa Vell., Fl. Flumin. 1: 4, t. 18. 1829. Calathea tuberosa (Vell.) Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 124. 1862. Goeppertia singularis (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea singularis H. A. Kenn., Novon 9: 61. 1999. Goeppertia ucayalina (Huber) Borchs. & S. Suárez, comb. nov. Calathea ucayalina Huber, Bol. Mus. Goeldi Hist. Nat. Ethnogr. 4: 551. 1906. Goeppertia soconuscum (Matuda) Borchs. & S. Suárez, comb. nov. Calathea soconuscum Matuda, Anales Inst. Biol. Univ. Nac. México 21: 327. 1951. Goeppertia ulotricha ( J. F. Macbr.) Borchs. & S. Suárez, comb. nov. Calathea ulotricha J. F. Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 11: 52. 1931. 2012] BORCHSENIUS ET AL.: PHYLOGENY OF CALATHEA 635 Goeppertia umbrosa (Körn.) Borchs. & S. Suárez, comb. nov. Calathea umbrosa Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 137. 1862. R. Sweet, Hort. Brit., ed. 3: 658. 1839. Calathea villosa (Lodd. ex G. Don) Lindl., Edwards’s Bot. Reg. 20(Misc.): 61. 1834. Goeppertia undulata (Linden & André) Borchs. & S. Suárez, comb. nov. Maranta undulata Linden & André, Cat. Gén. 87: 4. 1871. Calathea undulata (Linden & André) Linden & André, Ill. Hort. 19: t. 98. 1872. Goeppertia vinosa (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea vinosa H. A. Kenn., Ann. Missouri Bot. Gard. 60: 422. 1973. Goeppertia ursina (Standl.) Borchs. & S. Suárez, comb. nov. Calathea ursina Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 70. 1940. Goeppertia violacea (Roscoe) Borchs. & S. Suárez, comb. nov. Phrynium violaceum Roscoe, Monandr. Pl. Scitam.: t. 37. 1828. Calathea violacea (Roscoe) Lindl., Bot. Reg. 11: t. 932. 1834. Goeppertia vaginata (Petersen) Borchs. & S. Suárez, comb. nov. Calathea vaginata Petersen, Vidensk. Meddel. Naturhist. Foren. Kjøbenhavn 1889: 331. 1889. Goeppertia virginalis (Linden ex Regel) Borchs. & S. Suárez, comb. nov. Calathea virginalis Linden ex Regel, Gartenflora 28: 299. 1879. Goeppertia varians (K. Koch & Mathieu) Borchs. & S. Suárez, comb. nov. Phrynium varians K. Koch & Mathieu, Index Seminum (B) 1855(App.): 12. 1855. Calathea varians (K. Koch & Mathieu) Körn., Gartenflora 7: 87. 1858. Goeppertia wallisii (Linden) Borchs. & S. Suárez, comb. nov. Maranta wallisii Linden, Ann. Hort. Belge Étrangère 17: 105. 1867. Calathea wallisii (Linden) Regel, Index Seminum (LE) 1869: 14. 1869. Goeppertia variegata (K. Koch) Borchs. & S. Suárez, comb. nov. Phrynium variegatum K. Koch, Berliner Allg. Gartenzeitung 25: 147. 1857. Calathea variegata (K. Koch) Linden ex Körn., Gartenflora 7: 88. 1858. Goeppertia warszewiczii (L. Mathieu ex Planch.) Borchs. & S. Suárez, comb. nov. Maranta warszewiczii L. Mathieu ex Planch., Fl. Serres Jard. Eur. 9: 209. 1854. Calathea warszewiczii (L. Mathieu ex Planch.) Planch. & Linden, Cat. Pl. Exot. 10: 3. 1855. Goeppertia veitchiana (Veitch ex Hook. f.) Borchs. & S. Suárez, comb. nov. Calathea veitchiana Veitch ex Hook. f., Bot. Mag. 91: t. 5535. 1865. Goeppertia whitei (Rusby) Borchs. & S. Suárez, comb. nov. Calathea whitei Rusby, Mem. New York Bot. Gard. 7: 221. 1927. Goeppertia velutina (Poepp. & Endl.) Borchs. & S. Suárez, comb. nov. Phrynium velutinum Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 19. 1837. Calathea velutina (Poepp. & Endl.) Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 127. 1862. Goeppertia widgrenii (Körn.) Borchs. & S. Suárez, comb. nov. Calathea widgrenii Körn., Bull. Soc. Imp. Naturalistes Moscou 35(1): 117. 1862. Goeppertia venusta (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea venusta H. A. Kenn., Ann. Missouri Bot. Gard. 60: 416. 1973. Goeppertia williamsii (J. F. Macbr.) Borchs. & S. Suárez, comb. nov. Calathea williamsii J. F. Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 11: 54. 1931. Goeppertia verapax (Donn. Sm.) Borchs. & S. Suárez, comb. nov. Calathea verapax Donn. Sm., Bot. Gaz. 31: 124. 1901. Goeppertia wiotii (E. Morren) Borchs. & S. Suárez, comb. nov. Maranta wiotii E. Morren, Ann. Bot. Hort. 25: 273. 1875. Calathea wiotii (E. Morren) Regel, Gartenflora 28: 298. 1879. Goeppertia verecunda (H. A. Kenn.) Borchs. & S. Suárez, comb. nov. Calathea verecunda H. A. Kenn., Bot. Not. 130: 336. 1977. Goeppertia villosa (Lodd. ex G. Don) Borchs. & S. Suárez, comb. nov. Phrynium villosum Lodd. ex G. Don in Goeppertia zingiberina (Körn.) Borchs. & S. Suárez, comb. nov. Calathea zingiberina Körn., Bull. Soc. Imp. Naturalistes Moscou 35: t. 122. 1862.

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