Abstract

Dendrophthora and Phoradendron are neotropical genera of mistletoes belonging to the tribe Phoradendreae. The phylogenetic relationships between these two genera are still not entirely understood and, morphologically, Dendrophthora differs from Phoradendron solely by the presence of one, anther locule, as opposed to two in the latter. This study provides a synopsis of species of both genera for Brazil, where we revised all the names with Brazilian types or cited for the country. A total of 173 names were evaluated and the presence of 44 taxa in Brazil was confirmed: three species of Dendrophthora and 41 of Phoradendron. An identification key for the confirmed taxa is proposed and taxonomic comments are elaborated, along with an iconography, geographic distribution of the taxa, and original illustrations and photos of some species. Six new synonyms are proposed in Dendrophthora, and 12 in Phoradendron; 27 names are lectotypified.

Key words
mistletoes; parasitic plants; Phoradendreae; Santalales

Resumo

Dendrophthora e Phoradendron são gêneros neotropicais de ervas-de-passarinho, pertencentes a tribo Phoradendreae. As relações filogenéticas entre estes dois gêneros ainda não são totalmente claras, e morfologicamente, Dendrophthora difere de Phoradendron unicamente pela presença de um, ao contrário de dois, lóculos na antera neste último. Este estudo traz uma sinopse das espécies destes dois gêneros para o Brasil, na qual foram revisados todos os nomes com tipos brasileiros ou citados para o país. Para esta sinopse, avaliamos 173 nomes e confirmamos a ocorrência de 44 táxons para o Brasil, sendo três espécies de Dendrophthora e 41 espécies de Phoradendron. É proposta uma chave de identificação para as espécies confirmadas, são feitos comentários taxonômicos, junto com iconografia, distribuição geográfica dos táxons, e ilustrações e fotografias inéditas de algumas espécies. São propostos seis novos sinônimos em Dendrophthora e 12 em Phoradendron, e 27 nomes são lectotipificados.

Palavras-chave
passarinho; plantas parasitas; Phoradendreae; Santalales

Introduction

Dendrophthora Eichler and Phoradendron Nutt. are neotropical genera of mistletoes currently included in Santalaceae (sensu APG IV 2016APG IV - Angiosperm Phylogeny Group (2016) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society 181: 1-20.). Phoradendron was separated from Viscum L. by Nuttal (1848)Nuttal T (1848) Descriptions of plants collected by William Gambel, M.D., in the Rocky Mountains and Upper California. Journal of the Academy of Natural Sciences of Philadelphia, Second Series 1: 185-186. for having bilocular anthers, as opposed to multilocular anthers present in Viscum, which occurs in the temperate and tropical “Old World”. Nonetheless, it was Eichler (1868)Eichler AW (1868) Loranthaceae. In: Martius CFP (ed.) Flora brasiliensis. Fleicher, Leipzig. Vol. 5, pars 2, pp. 1-136, t.1-44. who provided a clear circumscription of Phoradendron in his treatments in Flora brasiliensis, and established the genus Dendrophthora for species with unilocular anthers.

Posteriorly, Engler (1889)Engler A (1889) Loranthaceae. Die Natürlichen Pflanzenfamilien 3: 156-198. classified both genera in tribe Phoradendreae Engler, based on the inflorescence morphology, and proposed three sections for each genus. Shortly afterwards, Van Tieghem (1896)Van Tieghem MP (1896) Sur le groupement des espèces en genres dans les Ginalloées, Bifariées, Phoradendrées et Viscées, quatre tribus de la famille des Loranthacées. Bulletin de la Société Botanique de France 43: 161-194. proposed four sections for Phoradendron, and three for species that he recognized in Dendrophthora, both based on the inflorescences structure. In the early 20th century, Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p. elaborated the monograph of the genus Phoradendron, which became the main reference for decades, where the author proposed a subdivision of the group in two (based on the presence or absence of cataphylls), later recognized as subgenera by Engler & Krause (1935)Engler A & Krause K (1935). Loranthaceae. In: Engler A & Prantl KAE (eds.) Die Natürlichen Pflanzenfamilien. 2nd ed. Vol. 16b. Wilhelm Engelmann, Leipzig. Pp. 98-203.: Aequatoriales Trel. ex Engler & Krause and Boreales Trel. ex Engler & Krause. Trelease’s work provides photographs of the nomenclatural types for most species, especially from B and W herbaria, which had part of their botanical collections destroyed during the second World War.

In the early 1950’s, C.T. Rizzini made significant contributions to the Brazilian flora, namely the Loranthaceae monograph, where several new species were described (Rizzini 1956Rizzini CT (1956) Pars specialis prodromi monographiae Loranthacearum Brasiliae terrarumque finitimarum. Rodriguésia 30-31: 87-234.). His efforts continued until 1980, and included description of species of Dendrophthora and Phoradendron for Brazil (Rizzini 1968aRizzini CT (1968a) Loranthaceae Austro-Americanae Novae. Rodriguésia 41: 7-35.,bRizzini CT (1968b) Lorantáceas Catarinenses. In: Reitz R (ed.) Flora Ilustrada Catarinense. Herbário Barbosa Rodrigues, Itajaí. 44p., 1975Rizzini CT (1975) Loranthaceae Novae. Revista de la Facultad de Agronomía Maracay 8: 83-109., 1976Rizzini CT (1978) El género Phoradendron en Venezuela. Rodriguésia 46: 33-125., 1980Rizzini CT (1980) Loranthaceae of the Central Brazil. Arquivos do Jardim Botânico do Rio de Janeiro 24: 19-50.) and Venezuela (Rizzini 1978Rizzini CT (1978) El género Phoradendron en Venezuela. Rodriguésia 46: 33-125., 1982Rizzini CT (1982) Loranthaceae. In: Luces FZ & Steyermark JA (eds.) Instituto Botánico, Caracas. Flora de Venezuela 4: 7-316. ).

Kuijt (1961, 2000Kuijt J (2000) An update on the genus Dendrophthora (Viscaceae). Botanische Jahrbucher fur Systematik Pflanzengeschichte und Pflanzengeographie 122: 169-193.) reviewed Dendrophthora and established two subgenera, Eudendrophthora Kuijt and Paradendrophthora Kuijt, delimited on the basis of flowers seriation in its fertile bracts: the first former with a single series of flowers (uniseriated) and the latter with two or more series. However, some species were not included under any of these two subgenera. For Phoradendron, Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643. makes a substantial review, including 234 species, 78 of them new, but no infrageneric organization is proposed, indicating only probable alliances of natural groups of species.

Molecular data from Der & Nickrent (2008)Der JP & Nickrent DL (2008) A molecular phylogeny of Santalaceae (Santalales). Systematic Botany 33: 107-116. demonstrated that Santalaceae was a polyphyletic group. After reorganizing the species, the remaining genera included in Santalaceae were distributed into six well supported clades, which were recognized as families by Nickrent et al. (2010)Nickrent DL, Malécot V, Vidal-Russell R & Der JP (2010) A revised classification of Santalales. Taxon 59: 538-558.. However, in APG IV (2016)APG IV - Angiosperm Phylogeny Group (2016) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society 181: 1-20. these six families are treated as one, despite the few morphological synapomorphes for those clades. In the studies of Der & Nickrent (2008)Der JP & Nickrent DL (2008) A molecular phylogeny of Santalaceae (Santalales). Systematic Botany 33: 107-116. , Nickrent et al. (2010)Nickrent DL, Malécot V, Vidal-Russell R & Der JP (2010) A revised classification of Santalales. Taxon 59: 538-558., Su et al. (2015)Su HJ, Hu JM, Anderson FE, Der JP & Nickrent DL (2015) Phylogenetic relationships of Santalales with insights into the origins of holoparasitic Balanophoraceae. Taxon 64: 491-506. and Nickrent et al. (2019)Nickrent DL, Anderson F & Kuijt J (2019) Inflorescence evolution in Santalales: integrating morphological characters and molecular phylogenetics. American Journal of Botany 106: 402-414., Viscaceae (sensu Barlow 1964Barlow BA (1964) Classification of the Loranthaceae and Viscaceae. Proceedings of the Linnean Society of New South Wales 89: 268-272. and Cronquist 1981Cronquist A (1981) An integrated system of classification of flowering plants. Columbia University Press. New York. 1262p.) remained intact as the most derived family and was resolved as monophyletic with good support, as well as Phoradendreae, a clade formed by the genera Dendrophthora and Phoradendron. In the analysis of these two genera, Ashworth (2000, 2017) confirmed the monophyly of Phoradendreae, but suggested that Phoradendron was non-monophyletic and no morphological traits or combinations of traits were found informative to characterize the three major clades. Dendrophthora was resolved as monophyletic, nested within Phoradendron, and sister to Phoradendron piperoides (Kunth) Trel. (Ashworth 2017Ashworth VETM (2017) Revisiting phylogenetic relationships in Phoradendreae (Viscaceae): utility of the trnL-F region of chloroplast DNA and presence of a homoplasious inversion in the intergenic spacer. Botany 95: 247-258.). Yet, molecular studies in Phoradendreae have poorly represented the taxonomic diversity at species level, with only 6% of the species in the group having been sampled (Ashworth 2000Ashworth VETM (2000) Phylogenetic relationships in Phoradendreae (Viscaceae) inferred from three regions of the nuclear ribosomal cistron. I. Major lineages and paraphyly of Phoradendron. Systematic Botany 25: 349-370., 2017).

In the early 21st century, ca. 240 species of Phoradendron, and 125–130 of Dendrophthora, are recognized (Kuijt 2013Kuijt J (2013) A brief taxonomic history of neotropical mistletoe genera, with a key to the genera. Blumea 58: 263-266.; Kuijt & Hansen 2015Kuijt J & Hansen B (2015) Flowering Plants Eudicots - Santalales, Balanophorales. In: Kubitzki K (ed.) The families and genera of vascular plants. Vol. XII. Springer-Verlag, Berlin, Heidelberg, New York. Pp. 1-213.; Canelón et al. 2020Canelón DS, Niño SM, Dorr LJ & Caraballo-Ortiz MA (2020) Two new species of Dendrophthora (Viscaceae) from the Venezuelan Andes. PhytoKeys 140: 1-10.; Nickrent 2020Nickrent DL (2020) Parasitic angiosperms: how often and how many? Taxon 69: 5-27.). Although molecular evidence suggests that Dendrophthora occupies a nested position in Phoradendron (Ashworth 2000Ashworth VETM (2000) Phylogenetic relationships in Phoradendreae (Viscaceae) inferred from three regions of the nuclear ribosomal cistron. I. Major lineages and paraphyly of Phoradendron. Systematic Botany 25: 349-370., 2017), Kuijt (2013)Kuijt J (2013) A brief taxonomic history of neotropical mistletoe genera, with a key to the genera. Blumea 58: 263-266. did not consider a nomenclatural modification necessary at that moment, due to limited sampling in molecular studies, and difficulties in the identifications in this group.

Phoradendreae comprises perennials hemiparasitic herbs, called mistletoes (“ervas-de-passarinho” in Portuguese) which form primary haustorial connections but do not produce epicortical roots (as in some Loranthaceae). It shows different growth patterns such as: monopodial/percurrent or sympodial/dichotomous, or, frequently, a combination of both (Fig. 1a-c). The stems may be segmented by the presence of cataphylls, which are pairs of bract or scale-like structures, free or fused, present in all internodes or only in proximal lateral internodes (Fig. 1d-e), in variable number. Cataphylls can subtend inactive buds, or buds that develop in inflorescences (fertile cataphylls). Leaves may be squamate or expanded, and exhibit high morphological variability, but are generally fleshy or coriaceous, with somewhat conspicuous veins. The inflorescence type present in Phoradendron and Dendrophthora is a clear synapomorphy for both genera, and is unique in Santalales, and in all angiosperms (Nickrent et al. 2019Nickrent DL, Anderson F & Kuijt J (2019) Inflorescence evolution in Santalales: integrating morphological characters and molecular phylogenetics. American Journal of Botany 106: 402-414.). Each segment or fertile internode of the inflorescence is formed by the union of two opposite fertile bracts from where the flowers grows (Fig. 1f). Flowers do not arise from the apical meristem, as for most angiosperms. Instead, they are formed by an intercalary meristem, located in the proximal region of each bract. Therefore, the flowers located at the apex of the fertile bract are the most developed, and those at the base the youngest (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.). The presence of segments with sterile bracts at the inflorescence base varies in number according to the species (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.). The flowers are arranged in longitudinal series, from one to many series (uni-, bi-, tri-, multiseriate) (Fig. 1g-j). In the male flower, anther is adnate to each petal. Morphologically, Dendrophthora differs from Phoradendron solely by the presence of one as opposed to two, anther locules.

More regional studies are necessary to improve the knowledge of these species, to evaluate populations in different environments, rendering the analysis of the morphological variability possible. For example, Dettke & Waechter (2014)Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985., when studying Phoradendron in Southern Brazil, proposed the reestablishment of one species, Phoradendron burkartii Rizzini & Ulib., and nine names as synonyms, after field collections and consultation of the large and small herbaria of the region.

For Brazil, Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643. lists 59 species of Phoradendron. According to Arruda et al. (2012)Arruda R, Fadini RF, Carvalho LN, Del-Claro K, Mourão FA, Jacobi CM, Teodoro GS, van den Berg E, Caires CS & Dettke GA (2012) Ecology of Neotropical mistletoes: an important canopy dwelling component of Brazilian ecosystems. Acta Botanica Brasilica 26: 264-274., the ca. 60 species of Phoradendron are distributed in all Brazilian phytogeographic domains, while Dendrophthora has only three species and a more restricted distribution. Hence, the aim of this study was to review all names cited for Brazil, obtaining a better overview of the diversity in this country. This work was, in part, made possible thanks to the availability of high definition images of the nomenclatural types, through the repatriation project of Brazilian historical collections (Reflora 2020Reflora (2020) Herbário Virtual. Available at <http://reflora.jbrj.gov.br/reflora/herbarioVirtual/>. Access on 9 April 2020.
http://reflora.jbrj.gov.br/reflora/herba... ), and recent efforts to catalog the Brazilian flora (BFG 2015BFG - The Brazil Flora Group (2015) Growing knowledge: an overview of seed plant diversity in Brazil. Rodriguésia 66: 1085-1113., 2018BFG - The Brazil Flora Group (2018) Brazilian Flora 2020: innovation and collaboration to meet Target 1 of the Global Strategy for Plant Conservation (GSPC). Rodriguésia 69: 1513-1527.).

Material and Methods

For this study, all names of Dendrophthora and Phoradendron cited for Brazil, or with a type specimen collected in the Brazilian territory, were analyzed through a personal visit to the herbaria, or analysis of high-definition images provided by the Reflora project, and consulted via the Virtual Herbarium (Reflora 2020Reflora (2020) Herbário Virtual. Available at <http://reflora.jbrj.gov.br/reflora/herbarioVirtual/>. Access on 9 April 2020.
http://reflora.jbrj.gov.br/reflora/herba... ). The following herbaria were analyzed: A, B, BM, BR, C, CAS, CEN*, CEPEC*, CGMS*, CM, COL, CORD*, DAO, E, ESA*, F, G, GH, HBG, HBR*, HUA, HUEFS*, IAC, IAN*, IBGE*, ILL, INPA*, K, L, LE, LP*, M, MA, MBM*, MG*, MICH, MO, NY, P, PH, R*, RB*, RFA, S, SI*, SP*, SPF*, TCD, U, UB*, UC, UEC*, UFMG*, US, W, WIS, acronyms according to Thiers (continuously updated). The acronyms followed by an asterisk (*) were personally visited. Specimens examined are marked with exclamation marks (!). Morphological terminology for growth patterns, cataphylls and inflorescences are in agreement with Eichler (1868)Eichler AW (1868) Loranthaceae. In: Martius CFP (ed.) Flora brasiliensis. Fleicher, Leipzig. Vol. 5, pars 2, pp. 1-136, t.1-44. and Kuijt (1961; 2003). For additional images, vouchers list, distribution and descriptions of all Brazilian species, see Dettke & Caires (2020)Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... . The illustrations were made by C.S. Caires. An index to the names treated in this synopsis can be found in the Appendix S1 (available on supplementary material <https://doi.org/10.6084/m9.figshare.16879597.v1>). The index of type collectors is presented in Appendix S2 (available on supplementary material <https://doi.org/10.6084/m9.figshare.16879597.v1>).

Results and Discussion

For this synopsis, 173 names were evaluated, of which 44 taxa are recognized for Brazil, three species in Dendrophthora and 41 species in Phoradendron. Some more morphological characteristics are shown in Figures 1k-n and some species in vivo in Figures 2a-p and Figures 3a-p.

The range of distribution of Brazilian species in the Neotropical region is summarized in Table 1. Eight species are widely distributed, including North, Central America (including Caribbean) and South America; and five species occur in both Central and South America. Almost 70% of the species have exclusive distribution in South America, with eleven endemic species to Brazil. We consider nine species as rare, with few records in Brazil, restricted only by the type or very few collections: Phoradendron bicarinatum, P. dimerostachys, P. krukovii and P. oliveirae are only known for the type material collected in Brazil; Phoradendron juruanum and P. laxiflorum also have type material collected in Brazil, but are registered too in few areas in adjacent countries; and four species, Dendrophthora fendleriana, Phoradendron argentinum, P. diminutivum and P. singulare, also occur in neighboring countries, but have few collections in Brazilian territory.

Identification key for Dendrophthora and Phoradendron species in Brazil

• 1. Plants with squamiform leaves.

  • 2. Stem dorsiventrally flattened; inflorescence with 1–2 fertile internodes; mature fruit translucent white...................2.40. Phoradendron tunaeforme

  • 2’. Stem cylindrical; inflorescence with 3–5 fertile internodes; mature fruit yellowish, orange or reddish.

    • 3. Plants dioecious; branches erect, internodes short (0.8–3.5 cm long); inflorescences triseriate, with 4–10 flowers per fertile bract...................2.16. Phoradendron fragile

    • 3’. Plants monoecious; branches pending, internodes long (usually more than 4 cm long); inflorescences biseriate, with only 3 flowers per fertile bract......................................2.34. Phoradendron poeppigii

• 1’. Plants with expanded leaves.

  • 4. Stem dorsiventrally flattened...................2.33. Phoradendron platycaulon

  • 4’. Stem not flattened, other shapes.

    • 5. Internodes with fertile cataphylls.

      • 6. Internode with two or more, pairs of fertile cataphylls.........................................................2.11. Phoradendron crassifolium

      • 6’. Internode with only one pair of fertile cataphylls.

        • 7. Leaves with pinnate veins, with only one primary vein visible.

          • 8. Leaf base generally obtuse; leaf blade symmetric; inflorescence internodes cylindrical; 3–5 flowers per fertile bract...................2.17. Phoradendron harleyi

          • 8’. Leaf base acute; leaf blade asymmetric; inflorescence internodes clavate; 7–9 flowers per fertile bract...................2.29. Phoradendron oliveirae

        • 7’. Leaves with palmate veins, with 3–5 primary veins.

          • 9. Branching sympodial, dichotomous; all inflorescence internodes biseriate.........................................................2.10. Phoradendron craspedophyllum

          • 9’. Branching predominantly monopodial, percurrent, with rare dichotomies; inflorescence internodes uniseriate, or more rarely, only the basal ones biseriate.

            • 10. Inflorescence with only a pair of sterile bracts at the base, 2 or 3 fertile internodes; mature fruits ellipsoid...................2.38. Phoradendron singulare

            • 10’. Inflorescence with 2–4 pairs of sterile bracts at the base, 5 or more fertile internodes; mature fruits globose.........................................................2.20. Phoradendron inaequidentatum

    • 5’. Internodes without fertile cataphylls.

      • 11. Cataphylls present in all internodes.

        • 12. Leaves with pinnate veins.

          • 13. Inflorescences triseriate; inflorescence internodes clavate.........................................................2.21. Phoradendron juruanum

          • 13’. Inflorescences biseriate; inflorescence internodes cylindrical.

            • 14. Branching predominantly sympodial, dichotomous; lateral internodes with only a pair of basal cataphylls; leaves with rounded or retuse apex; fruits white with closed sepals...................2.27. Phoradendron northropiae

            • 14’. Branching predominantly monopodial, percurrent; lateral internodes with 2–4 pairs of basal cataphylls; leaves with acute or acuminate apex; fruits yellowish to reddish with erect open sepals...................2.32. Phoradendron piperoides

        • 12’. Leaves with palmate veins.

          • 15. Branching predominantly sympodial, dichotomous.

            • 16. Stem with whitish surface and longitudinal fissures; leaves with acuminate apex.........................................................2.22. Phoradendron krukovii

            • 16’. Stem without this characteristic; leaves with acute or rounded apex.

              • 17. Inflorescences triseriate; 9–21 flowers per fertile bract.........................................................2.3. Phoradendron berteroanum

              • 17’. Inflorescences biseriate; only 3 flowers per fertile bract.

                • 18. Lateral internodes with 2–3 pairs of cataphylls; leaves with crenulated margin; fruits globose, white...................2.5. Phoradendron burkartii

                • 18’. Lateral internodes with only one pair of cataphylls; leaves with a smooth margin; fruits ovoid, orange or reddish.........................................................2.39. Phoradendron strongyloclados

          • 15’. Branching predominantly monopodial, percurrent.

            • 19. Inflorescences triseriate.

              • 20. Internodes with tubular cataphylls; terminal inflorescences present.........................................................1.2. Dendrophthora fendleriana

              • 20’. Internodes with free cataphylls; terminal inflorescences absent.

                • 21. Stem with two conspicuous lateral wings; inflorescences bisexual (staminate and pistillate flowers); inflorescences internode clavate......................................2.7. Phoradendron chrysocladon

                • 21’. Stem without lateral wings; inflorescences unisexual (with only staminate flowers); inflorescences internodes cylindrical.........................................................2.23. Phoradendron laxiflorum

            • 19’. Inflorescences uni- or biseriate.

              • 22. Lateral internodes with only one pair of cataphylls; leaves predominantly ovate or elliptical, less than 1.5 cm wide...................2.39. Phoradendron strongyloclados

              • 22’. Lateral internodes with 2 pairs of cataphylls; leaves predominantly elliptical or obovate, wider than 1.5 cm.

                • 23. Plants dioecious; cataphylls free; leaves elliptical, with apiculate or acuminate apex; pistillate inflorescences uniseriate only......................................2.23. Phoradendron laxiflorum

                • 23’. Plants monoecious; cataphylls fused; leaves obovate with a rounded apex; inflorescences biseriate...................2.13. Phoradendron diminutivum

      • 11’. Cataphylls present only in the proximal internodes of lateral branches.

        • 24. Epiparasite of other species of Phoradendron; stem winged, with two or four distinct wings......................................2.14. Phoradendron dipterum

        • 24’. Hemiparasite of other angiosperms; stem non-winged.

          • 25. Leaves with pinnate veins, only one primary vein conspicuous.

            • 26. Inflorescences triseriate.

              • 27. Plants dioecious; internodes with one, rarely two, pairs of cataphylls.........................................................2.18. Phoradendron hexastichum

              • 27’. Plants monoecious; internodes with generally 2–5 pairs of lateral cataphylls.........................................................1.3. Dendrophthora warmingii

            • 26’. Inflorescences biseriate.

              • 28. Branching predominantly sympodial, dichotomous...................2.37. Phoradendron racemosum

              • 28’. Branching predominantly monopodial, percurrent.

                • 29. Leaves linear-lanceolate to falcate, up to 1.5 cm wide.........................................................2.4. Phoradendron bicarinatum

                • 29’. Leaves circular, elliptical, ovate, oblong or obovate, rarely falcate, more than 1.5 cm wide.

                  • 30. Leaves with acute apex, usually acuminate...................2.41. Phoradendron undulatum

                  • 30’. Leaves with obtuse rounded apex.

                    • 31. Inflorescence with 1 or 2 pairs of sterile bracts at the base; fruits with erect open petals...................1.3. Dendrophthora warmingii

                    • 31’. Inflorescence with 4 pairs, or more, of sterile bracts at the base; fruits with closed petals...................2.35. Phoradendron pteroneuron

          • 25’. Leaves with palmate veins, with 3–7 primary veins, generally inconspicuous.

            • 32. Small leaves, maximum 2 cm long; inflorescences with only 1 fertile internode.........................................................1.1. Dendrophthora elliptica

            • 32’. Larger leaves, more than 2 cm long; inflorescences with 2, or more, fertile internodes.

              • 33. Inflorescences biseriate.

                • 34. Fruits distinctly ellipsoid...................2.28. Phoradendron obtusissimum

                • 34’. Fruits globose or ovoid.

                  • 35. Fertile bract with only 3 flowers, the apical flowers staminate; fruits with verrucose epicarp.

                    • 36. Leaves small, maximum 1 cm wide; young leaves glaucous, covered with epicuticular wax; only the apex of the epicarp verrucose......................................2.1. Phoradendron argentinum

                    • 36’. Leaves larger, more than 1 cm wide; young leaves green; the entire epicarp verrucose...................2.25. Phoradendron mucronatum

                  • 35’. Fertile bract with more than 3 flowers (if 3, all flowers pistillate); fruit with smooth pericarp.

                    • 37. Fertile bract with up to 4 flowers...................2.26. Phoradendron nigricans

                    • 37’. Fertile bract with more than 5 flowers.

                      • 38. Fertile bract with 15–23 flowers...................2.24. Phoradendron macrarthrum

                      • 38’. Fertile bract with 5–15 flowers.

                        • 39. Leaves lanceolate to oblong-lanceolate; infrutescence congested, shorter than 2 cm in length...................2.8. Phoradendron congestum

                        • 39’. Leaves of other shapes; infrutescence laxa, longer than 2.5 cm in length.

                          • 40. Young stems strongly rhombic or quadrangular; mature fruits dark-orange or reddish...................2.36. Phoradendron quadrangulare

                          • 40’. Young stems cylindrical or ellipsoid; mature fruits yellowish, pink or light-orange.

                            • 41. Inflorescences generally clavate, staminate flowers positioned at the apex of the fertile bract, foveae distinctly deep; fruits with closed petals.........................................................2.2. Phoradendron bathyoryctum

                            • 41’. Inflorescences cylindrical, staminate flowers mixed with pistillate ones in the fertile bract; foveae medium to shallow; fruits with erect open petals.

                              • 42. Small leaves; narrow, 1.5–3 cm wide...................2.13. Phoradendron diminutivum

                              • 42’. Large leaves; wide, 3–7 cm wide...................1.3. Dendrophthora warmingii

              • 33’. Inflorescences triseriate.

                • 43. Plants monoecious.

                  • 44. Leaves small (3–6 × 1–1.5 cm); inflorescence with the entire basal internode bearing staminate flowers...................2.12. Phoradendron dimerostachys

                  • 44’. Leaves large (5–15 × 2–7 cm); inflorescence internodes with staminate flowers, mixed with pistillate flowers.

                    • 45. Fertile bract with 7–9 flowers; fruits ovoid with erect opened petals.........................................................1.3. Dendrophthora warmingii

                    • 45’. Fertile bract with 12–18 flowers; fruits globose with closed petals.........................................................2.31. Phoradendron perrottetii

                • 43’. Plants dioecious.

                  • 46. Inflorescences axillary only.

                    • 47. Branches erect; young leaves translucent in sicco; short leaves, up to 5 cm long.........................................................2.30. Phoradendron pellucidulum

                    • 47’. Branches pending; young leaves opaque in sicco; long leaves, more than 5 cm long.........................................................2.15. Phoradendron ensifolium

                  • 46’. Inflorescences axillary and terminal.

                    • 48. Plants usually pending; dark brown or black in sicco; cataphylls usually with a whitish margin...................2.9. Phoradendron coriaceum

                    • 48’. Plants erect; light brown or yellowish in sicco; cataphylls without whitish margin.

                      • 49. Inflorescences with only 1 pair of sterile bracts at the base; mature fruits white.........................................................2.19. Phoradendron holoxanthum

                      • 49’. Inflorescence usually with 2–3 pairs of sterile bracts at the base; mature fruits yellow or greenish...................2.6. Phoradendron caripense

Taxonomic treatment

1. Dendrophthora Eichler in Martius, Fl. bras. 5(2): 102. 1868. Type species: Dendrophthora opuntioides (L.) Eichler in Martius, Fl. bras. 5(2): 102. 1868, lectotype designated by N.L. Britton & P. Wilson, Sci. Surv. Porto Rico 5: 259. 1924.

1.1. Dendrophthora elliptica (Gardner) Krug & Urb., Ber. Deutsch. Bot. Ges. 14(8): 285. 1896. Phoradendron ellipticum (Gardner) Eichler in Martius, Fl. bras. 5(2): 119, Fig. 37-2. 1868. Viscum ellipticum Gardner, London J. Bot. 4: 106. 1845. Type: BRAZIL. RIO DE JANEIRO: Organ Montains, on Gaylussacia, G. Gardner 437 (lectotype, hic designatus: K - barcode 00601371! (= Kew Neg 4570); isolectotypes: BM!, G!, K!, P!).

Dendrophthora elliptica var. genuina Krug & Urb., Bot. Jahrb. Syst. 24: 69. 1897, nom. illeg.

Dendrophthora rubicunda Ule, Notizbl. Bot. Gart. Berlin-Dahlem 6: 289. 1915. Type: BRAZIL. RORAIMA: II. 1910, E.H.G. Ule 8601 (holotype: B, destroyed; lectotype, designated by Kuijt (1961)Kuijt J (1961) A revision of Dendrophthora (Loranthaceae). Wentia 6: 1-145.: K - barcode 00601370!; isolectotype: IAN).

Dendrophthora hylaeana Rizzini, Rodriguésia 41: 8. 1976. Type: BRAZIL. AMAZONAS: Serra Aracá, 10.II.1975, J.M. Pires 15042 (holotype: RB - barcode 00540550!).

Dendrophthora elliptica is a monoecious species with erect habit; the growth pattern is monopodial/percurrent, and possesses cataphylls only on the lateral proximal internodes. The leaves show palmate veins and widely variable in shape and size, elliptical or obovate, and do not exceed 2 cm in length, smaller than in any Brazilian species of Phoradendron. The small inflorescence is biseriate, with 3–5 flowers in the fertile bract, and the apical one is staminate. Rizzini (1956)Rizzini CT (1956) Pars specialis prodromi monographiae Loranthacearum Brasiliae terrarumque finitimarum. Rodriguésia 30-31: 87-234. illustrates the unilocular anther of D. elliptica (Fig. 20-3). This species can be confused with some species of Phoradendron, such as P. argentinum and P. mucronatum, being distinguished from them by the smaller leaves and minute inflorescences (up to 0.5 cm in length), with only one fertile internode.

The holotype was found in two duplicates at the K herbarium and, according to the Code, are considered syntypes, and one of them is selected by us as a lectotype (Turland et al. 2018Turland NJ, Wiersema JH, Barrie FR, Greuter W, Hawksworth DL, Herendeen PS, Knapp S, Kusber WH, Li DZ, Marhold K, May TW, McNeill J, Monro AM, Prado J, Price MJ & Smith GF (2018) International code of nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Koeltz Botanical Books, Glashütten. 254p., Art. 9.11).

Iconography: Eichler (1868, Fig. 37-2), Rizzini (1995Rizzini CT (1995) Flora da Serra do Cipó, Minas Gerais: Loranthaceae. Boletim de Botânica, Universidade de São Paulo 14: 207-221., Fig. 21), Rizzini (1956, Fig. 20), Kuijt (1961, Fig. 21), Kuijt (2005Kuijt J (2005) Viscaceae. In: Steyermark JA, Berry PE, Yatskievych K & Holst GK (ed.) Flora of the Venezuelan Guayana. Vol. IX (Rutaceae - Zygophyllaceae). Missouri Botanical Garden Press, Saint Louis. Pp. 464-492., Fig. 392).

It is distributed in South America - Brazil (AM, ES, GO, MG, RJ, RR, SP), Colombia and Venezuela (Kuijt 1961Kuijt J (1961) A revision of Dendrophthora (Loranthaceae). Wentia 6: 1-145.; Dettke & Caires 2020Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... ).

1.2. Dendrophthora fendleriana (Eichler) Kuijt, Taxon 43(2): 190. 1994. Phoradendron fendlerianum Eichler in Martius, Fl. bras. 5(2): 129. 1868. Type: VENEZUELA. Prope coloniam Tovar, 1854/1855, A. Fendler 1102 (lectotype, designated by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: B, destroyed (= Trelease, 1916, Fig. 211b); lectotype, designated by Kuijt (1994)Kuijt J (1994) Typifcation of the names of new world mistletoe taxa (Loranthaceae and Viscaceae) described by Martius and Eichler. Taxon 43: 187-199.: P barcode 00756353!; isolectotypes: BR!, G!(3x), GH!, GOET!, K!, MO).

Dendrophthora fendleriana is a new occurrence for Brazil, found in an altitude area of “Pico da Neblina”, close to the border with Venezuela, where it is already registered. This species can be recognized by a predominance of monopodial/percurrent branches, and less frequent dichotomies caused by the presence of terminal inflorescences; tubular cataphylls in all internodes; leaves elliptical or circular, with palmate veins and triseriate inflorescences. It resembles D. warmingii and P. racemosum in leaves morphology, but it is differentiated by the position of cataphylls, in all internodes in D. fendleriana and present only on proximal lateral internodes in D. warmingii and P. racemosum.

Examinated material: Santa Isabel do Rio Negro, Parque Nacional do Pico da Neblina, 2.000 m, 00º47’14’’N, 66º01’26’’W, 22.IX.2012, R.C. Forzza et al. 7285 (RB).

Iconography: Trelease (1916, Fig. 211b).

It is distributed in South America - Brazil (AM) and Venezuela (this study; Kuijt & Kellog 1996).

1.3. Dendrophthora warmingii (Eichler) Kuijt, Novon 13(1): 88. 2003. Phoradendron warmingii Eichler in Warming, Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 209. 1870. Type: BRAZIL. MINAS GERAIS: Lagoa Santa, on Vochysia elliptica, 14.I.1865, E. Warming 383 (lectotype, designated by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: C - barcode 10014053! (= Trelease, 1916, Fig. 198a); isolectotype: C10014050!).

Phoradendron mairaryense Ule, Notizbl. Bot. Gart. Berlin-Dahlem 6: 291. 1920. Type: BRAZIL. AMAZONAS: Rio Branco, Serra de Mairari, XI.1909, E.H.G. Ule 8383 (holotype: B, destroyed (= F Neg 18190); lectotype, hic designatus: K - barcode 00601485!; isolectotypes: G!, L!), syn. nov.

Dendrophthora tepuiana (Steyerm.) Kuijt, Proc. Kon. Ned. Akad. Wetensch., Biol.93(2): 138. 1990. Phoradendron tepuianum Steyerm., Fieldiana, Bot.28: 222. 1951. Type: VENEZUELA. BOLÍVAR: Ptari-tepuí, 1.XI.1944, J.A. Steyermark 59688 (holotype: F! (= F Neg 53343); isotypes: K!, NY!, US!).

Phoradendron simile Rizzini, Rodriguésia 18/19: 191. 1956. Type: BRAZIL. MINAS GERAIS: Santa Luzia, Lagoa Santa, on Qualea, 11.IX.1932, H.L. Mello Barreto 6549 (holotype: RB - barcode 00540614!), syn. nov.

Phoradendron orbiculare Rizzini, Rev. Fac. Agron. Maracay 8(3): 88. 1975. Type: BRAZIL. MATO GROSSO: 450 km past Cuiabá, route to Goiânia, 30.IX.1963, B. Maguire et al. 56949 (holotype: RB - barcode 00540517!; isotype: NY!), syn. nov.

Phoradendron sulfuratum Rizzini, Revista Fac. Agron. (Maracay) 8(3): 89, Fig. 6. 1975. TYPE: VENEZUELA. BOLÍVAR: Gran Sabana, km 148 S El Dorado, 20.XII.1970, J.A. Steyermark et al. 104137 (holotype: RB (two sheets: barcodes 00540522! and 00545550!); isotype: NY!), syn. nov.

Phoradendron triplinervium Rizzini, Rev. Fac. Agron. Maracay 8(3): 90. 1975. Type: BRAZIL. BAHIA: Piau River, 150 km SW Barreiras, on Vochysia, H.S. Irwin et al. 14827 (holotype: RB - barcode 00540618!; isotypes: IAN!, K!, NY!, UB!, US!), syn. nov.

Phoradendron linguiforme Rizzini, Mem. New York Bot. Gard. 29: 33. 1978. Type: BRAZIL. AMAPÁ: Araguari River, 5 hours above Mururé River, 01º24’N, 51º57’W, on Caraipa, 23.VIII.1961, J.M. Pires et al. 50445 (holotype: RB - barcode 00540589!; isotypes: IAN, INPA!, K!, NY!), syn. nov.

Phoradendron virens Rizzini, Ernstia 32: 3. 1985. Type: BRAZIL, DISTRITO FEDERAL: Brasília, São Bartolomeu River, 01.X.1989, on Vochysia, E.P. Heringer et al. 2136 (holotype: IBGE; isotype: RB (two sheets: barcodes 00540619! and 00545552!), syn. nov. Figs. 2a-b; 4

Dendrophthora warmingii can be distinguished by the greenish to yellowish color in vivo, and light-brown to yellowish in sicco. This species presents a combination of growth patterns (monopodial/percurrent and sympodial/dichotomous) with 1–5 pairs of tubular cataphylls only on the lateral proximal internodes. The leaves present a variation of venation patterns, sometimes within the same individuals or population, of pinnate and palmate (1–5 main veins) patterns. The leave shape is obovate, elliptical or circular, some falcate, with rounded apex. Inflorescences may be tri- or biseriate, with staminate flowers mixed with, and in fewer number than, pistillate flowers. This species is often seen parasiting on Vochysiaceae species (Qualea or Vochysia spp.), and less frequently on Melastomataceae species or others, but preferably on Myrtales (Caires & Proença 2008Caires CS & Proença CEB (2008) Levantamento preliminar dos hospedeiros de Dendrophthora e Phoradendron (Santalaceae) no Distrito Federal, Brasil. Heringeriana 2: 11-22.). (See comments under D. fendleriana, P. dimunutivum, P. inaequidentatum and P. pteroneuron).

After analyzing the type material, we include here Phoradendron mairaryense and its two synonyms, P. sulfuratum and P. linguiforme (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.), under synonymy of D. warmingii. We also include the names assigned by Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643. as synonyms of D. tepuinana: P. orbiculare, P. simile, P. triplinervium and P. virens, under synonymy of D. warmingii.

Regarding Phoradendron mairaryense, the holotype was destroyed during the second World War, and, therefore, a lectotype is here designated (Turland et al. 2018Turland NJ, Wiersema JH, Barrie FR, Greuter W, Hawksworth DL, Herendeen PS, Knapp S, Kusber WH, Li DZ, Marhold K, May TW, McNeill J, Monro AM, Prado J, Price MJ & Smith GF (2018) International code of nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Koeltz Botanical Books, Glashütten. 254p., Art. 9.11).

Material selected for illustration (Fig. 4): Brasília, APA de Cafuringa, Dois Irmãos Farm, on Vochysia pyramidalis Mart., 21.VIII.2002, C.S. Caires et al. 207 (UB), on Vochysia elliptica Mart., 21.VIII.2002, C.S. Caires et al. 211 (UB). Sobradinho, margin of DF-170 road, on Vochysia pyramidalis Mart., 22.III.2002, C.S. Caires et al. 72 (UB).

Iconography: Rizzini (1956, Fig. 15, 19-2), Rizzini (1980, Fig. 9, 11), Rizzini (1982, Fig. 75), Caires & Proença (2005Caires CS & Proença CEB (2005) Viscaceae. In: Cavalcanti TB & Ramos AE (orgs.) Flora do Distrito Federal, Brasil. Vol. 4. Embrapa Recursos Genéticos e Biotecnologia, Brasília. Pp. 43-76., Fig. 1a-c), Kuijt (2005, Fig. 395), Caires (2017, Figs. 1a-b, 2a-d).

It is distributed in South America - Brazil (AM, AP, BA, DF, GO, MG, MT, PA, RJ, RO, SP, TO), Guyana and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Caires 2017Caires CS (2017) Flora das cangas da Serra dos Carajás, Pará: Santalaceae. Rodriguésia 68: 1139-1149.; Dettke & Caires 2020Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... ).

2. Phoradendron Nutt., J. Acad. Nat. Sci. Philad., ser. 2, 1: 185. 1848. Type species: Phoradendron californicum Nutt., J. Acad. Nat. Sci. Philad., ser. 2, 1: 185. 1848.

2.1. Phoradendron argentinum Urb., Bot. Jahrb. Syst. 23, Beibl. 57: 14. 1897. Type: ARGENTINA. CATAMARCA: Chacarita de los Padres, XI.1872, G.W. Hieronymus 419 (lectotype, designated by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: B, destroyed (= Trelease, 1916, Fig. 177a); lectotype, designated by Dettke et al. (2011)Dettke GA, Lima LFP & Waechter JL (2011) Phoradendron argentinum (Viscaceae), new mistletoe for the Brazilian flora and its general distribution in South America. Darwiniana 49: 86-89.: F!, = F Neg 70633!; isolectotypes: CORD!, GOET, K!). Fig. 2c

This species was first cited for Brazil by Dettke et al. (2011)Dettke GA, Lima LFP & Waechter JL (2011) Phoradendron argentinum (Viscaceae), new mistletoe for the Brazilian flora and its general distribution in South America. Darwiniana 49: 86-89., and is characterized by glaucous young leaves, small biseriate inflorescences, with only three flowers per fertile bracts (apical flower staminate), and fruit with tuberculate epicarp in the apical portion. Phoradendron argentinum is similar to P. mucronatum, which has wider leaves (more than 1 cm), non glaucous leaves and an entirely tuberculate pericarp. (See comments under Dendrophthora elliptica).

Iconography: Eliçabe (1973Eliçabe SE (1973) Las lorantáceas chaqueñas. Notas preliminares para la Flora Chaqueña (Formosa, Chaco y Santiago del Estero) 4: 1-25., Figs. 6-7), Kuijt (2003, Fig. 35), Dettke & Waechter (2014, Figs. 1a-f), Dettke & Waechter (2013Dettke GA & Waechter JL (2013) Southern Brazilian mistletoes. Rapid color guides of Field Museum #493. Available at <https://fieldguides.fieldmuseum.org/pt-br/guias/guia/493>. Access on 30 March 2020.
https://fieldguides.fieldmuseum.org/pt-b... , Figs. 54-59).

It is distributed in South America - Argentina, Bolivia, Brazil (RS), Paraguay and Uruguay (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke et al. 2011Dettke GA, Lima LFP & Waechter JL (2011) Phoradendron argentinum (Viscaceae), new mistletoe for the Brazilian flora and its general distribution in South America. Darwiniana 49: 86-89.; Dettke & Waechter 2014Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.).

2.2. Phoradendron bathyoryctum Eichler in Martius, Fl. bras. 5(2): 123, Fig. 43-2. 1868. Type: BRAZIL. PIAUÍ: “In prov. Piauhy,” VIII.1839, G. Gardner 2626 (lectotype, designated by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: W, destroyed (= Trelease, 1916, Fig. 120a); lectotype, first step designated by Kuijt (1994)Kuijt J (1994) Typifcation of the names of new world mistletoe taxa (Loranthaceae and Viscaceae) described by Martius and Eichler. Taxon 43: 187-199.: P, second step, hic designatus: P - barcode 02441995!; isolectotypes: BM!, K!(2x), ILL!, P!, US!).

Phoradendron ulophyllum Eichler in Martius, Fl. bras. 5(2): 123. 1868. Type: BRAZIL. RIO DE JANEIRO: 1833, M. Gaudichaud 573 (lectotype, designated by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: B, destroyed (= Trelease, 1916, Fig. 108a); lectotype, designated by Kuijt (1994)Kuijt J (1994) Typifcation of the names of new world mistletoe taxa (Loranthaceae and Viscaceae) described by Martius and Eichler. Taxon 43: 187-199.: BR - barcode 0000005509416!; isolectotypes: P!(3x), G-DC!(2x)).

Phoradendron alophyllum Eichler ex Glaziou, Bull. Soc. Bot. France, Mem. 3g: 609. 1913. Type: BRAZIL. RIO DE JANEIRO: Villa Nova, A.F.M. Glaziou 7663 (lectotype, hic designatus: P - barcode 05455410!; isolectotype: P05455409!).

Phoradendron psittacanthobium Rizzini, Rodriguésia 30/31: 186. 1956. Type: BRAZIL. MATO GROSSO DO SUL: Corumbá, on Psittacanthus cordatus, which on Triplaris, S.M. Moore 1019 (holotype: BM - barcode 993557!).

Phoradendron pinheirense Rizzini, Arq. Jard. Bot. Rio de Janeiro 24: 35. 1980. Type: BRAZIL. Minas Gerais: Joao Pinheiro-Cristalina road, on Pterodon emarginatus [Pterodon polygalaeflorum], 6.VIII.1979, E.P. Heringer & C.T. Rizzini 17553 (lectotype, designated by Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.: RB (two sheets - barcodes 00540606! and 00545659!); isolectotypes: UC(ex-LEA), R!).

Phoradendron paraguari Kuijt, Candollea 49(1): 268, Fig. 1. 1994. Type: PARAGUAY. PARAGUARÍ: Paraguarí, 27.VIII.1893, C.A.M. Lindman 1941 (holotype: K - barcode 00568330!; isotypes: RB (ex-R)!, S!(2x)).

Phoradendron lindemanii Kuijt, Syst. Bot. Monogr. 66: 272, Fig. 162. 2003. Type: BRAZIL. PARANÁ: Serra do Mar, forest on Atlantic slope above Bela Vista, on old road from Curitiba to Morretes, 600–800 m, 23.II.1967, J.C. Lindeman & J.H. de Haas 4641 (holotype: U - barcode 8116!).

Phoradendron pachyneuron Kuijt, Syst. Bot. Monogr. 66: 331, Fig. 208. 2003. Type: BRAZIL. PARANÁ: Morretes, Iacarehy, 17.III.1909, P.K.H. Dusén 7863 (holotype: S R-9174!; isotype: S 11-24563!). Figs. 2d; 5

Phoradendron bathyoryctum is characterized by a mixture of growth patterns (monopodial/percurrent and sympodial/dichotomous), cataphylls present only on the proximal lateral branches, circular or ellipsoidal stems, sometimes keeled on the distal portion, biseriate inflorescences with staminate flowers, apically positioned, and fruits in deep alveolus. The leaves of these species are highly variable in size and shape among populations (elliptical, ovate, oblong or falcate), and generally possessing a rounded apex. It can be confused with P. perrottetii, which has a triseriate inflorescence, or with P. coriaceum, which has smaller leaves and is dioecious. Also, it can be confused with P. macrarthrum, which has generally longer and falcate leaves, and inflorescences with 15–23 flowers per fertile bract (vs. 5–15 in P. bathyoryctum).

Kuijt’s species, P. lindemanii, P. pachyneuron and P. paraguari, were considered synonyms of P. bathyoryctum by Dettke & Waechter (2014)Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985. due to the overlapping morphology of analysed populations from southern Brazil, Paraguay and Argentina.

Two duplicates of the lectotype of P. bathyoryctum designated by Kuijt (1994)Kuijt J (1994) Typifcation of the names of new world mistletoe taxa (Loranthaceae and Viscaceae) described by Martius and Eichler. Taxon 43: 187-199. were located at P herbarium. Here, one of these duplicates is selected for a second-step lectotypification, of which the barcode is indicated (Turland et al. 2018Turland NJ, Wiersema JH, Barrie FR, Greuter W, Hawksworth DL, Herendeen PS, Knapp S, Kusber WH, Li DZ, Marhold K, May TW, McNeill J, Monro AM, Prado J, Price MJ & Smith GF (2018) International code of nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Koeltz Botanical Books, Glashütten. 254p., Art. 9.17). For P. alophyllum, the original material was found with two syntypes at P herbarium, and one of them is here selected as lectotype.

Material selected for illustration (Fig. 5): Brasília, DF-180 road of S.D.V.S.A. Farm 098-02, Struthanthus, 26.III.2002, C.S. Caires & V.C. Mendes 93 (UB), on Struthanthus, 25.VI.2002, C.S. Caires et al. 169 (UB), S.D.V.S.A. 126-02, on Magonia pubescens A-St. Hil., 26.III.2002, C.S. Caires & V.C. Mendes 81 (UB).

Iconography: Eichler (1868, Fig. 43-2), Rizzini (1956, Fig. 13), Rizzini (1968b, Fig. 9), Eliçabe (1973, Fig. 10), Kuijt (2003, Figs. 40, 162, 208, 212), Caires & Proença (2005, Fig. 2g), Dettke & Waechter (2013, Figs. 60-67), Rigon & Cervi (2013Rigon J & Cervi AC (2013) O gênero Phoradendron Nutt. (Viscaceae) no estado do Paraná. Pesquisas, Botânica 64: 15-38., Figs. 3a-c), Dettke & Waechter (2014, Figs. 1g-k).

It is distributed in South America - Argentina, Bolivia, Brazil (all states), Colombia, Paraguay, Suriname and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Waechter 2014Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.; Dettke & Caires 2020Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... ).

2.3. Phoradendron berteroanum (DC.) Griseb., Fl. Brit. W. I. 313: 1864, [berterianum]. Viscum berteroanum DC., Prodr. 4: 281. 1830 [berterianum]. Phoradendron dichotomum Krug & Urb., Bot. Jahrb. Syst. 24(1): 48. 1897, nom. illeg., non Phoradendron dichotomum Ettingsh., 1872. Type: DOMINICAN REPUBLIC. SANTO DOMINGO: C.L.G. Bertero s.n. (holotype: G-DC! (= Trelease, 1916, Fig. 238b)).

Phoradendron campinense Trel., Phoradendron 160, Fig. 244a. 1916. Type: BRAZIL. SÃO PAULO: Campinas, III.1900, J.C. Novaes 417a p.p. (holotype: US – barcode 107471!) [mixed with P. crassifolium 417b]. Fig. 2e

Phoradendron berteroanum can be recognized by its erect habit, sympodial/dichotomous branches and terete stems, large leaves with palmate veins, and long triseriate inflorescences, at axillary and terminal position. This species is often confused with P. crassifolium, which differs in presenting fertile cataphylls and only axillary inflorescences. (See comments under P. krukovii).

Iconography: Rizzini (1956, Fig. 11-1), Kuijt (2003, Figs. 43-44), Rigon & Cervi (2013, Figs. 3d,g), Dettke & Waechter (2014, Figs. 1l-o).

It is distributed in North America - Mexico, Central America (including Caribbean) and South America - Bolivia, Brazil (AC, MG, PR, RS, SP), Colombia, Ecuador, Peru and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Caires 2020Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... ).

2.4. Phoradendron bicarinatum Kuijt, Syst. Bot. Monogr. 66: 116, Fig. 47. 2003. TYPE: BRAZIL. AMAZONAS: Rio Negro, Serra Jacamim, Maia et al. 358 (holotype: MO).

Phoradendron bicarinatum was described by Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643., based only on two collections from Rio Negro. This species is distinct from other species of Phoradendron due to its leaves with pinnate veins, linear-lanceolate to falcate in shape, up to 1.5 cm wide, and short biseriate inflorescences. This species resembles P. congestum, especially by the rhombic and keeled stem, and inflorescence morphology, differing only in the veins, which are palmate in P. congestum. We would not be surprised if, in the future with new collections being made, they will be recognized as synonyms.

Iconography: Kuijt (2003, Fig. 47).

It is distributed in South America - Brazil (AM) (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.).

2.5. Phoradendron burkartii Rizzini & Ulib., Darwiniana 27: 499, Fig. 1. 1986. Type: ARGENTINA. ENTRE RÍOS: Federación, Rincón del Mocoreta y del Uruguay, 16.IV.1960, A. Burkart & J.C. Gamerro 21702 (holotype: SI - barcode 2184!; isotypes: SI!(4x)). Fig. 2f

Phoradendron burkartii was reestablished as a species by Dettke & Waechter (2014)Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.. This species is easily recognized by its sympodial/dichotomous, branches, crenulated leaf margin, and biseriate inflorescences, with only three flowers per fertile bract (apical flower staminate), located at the distal portion of the bract. (See comments under P. krukovii).

Iconography: Rizzini & Ulibari (1986, Fig. 1), Dettke & Waechter (2013, Figs. 68-72), Rigon & Cervi (2013, Figs. c-d), Dettke & Waechter (2014, Figs. 2a-e).

It is distributed in South America - Argentina, Brazil (PR, SC, RS) and Uruguay (Dettke & Waechter 2014Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.).

2.6. Phoradendron caripense Eichler in Martius, Fl. bras. 5(2): 111. 1868. Type: BRAZIL. PARÁ: usque Caripi, R. Spruce 140 (holotype: M!, = Trelease, 1916, Fig. 104b; isotype: BM).

Phoradendron distans Rizzini, Rev. Fac. Agron. Maracay 8(3): 87. 1975. Type: BRAZIL. MINAS GERAIS: Cerrado, 2 km SW of Diamantina, on Vochysia, 22.I.1969, H.S. Irwin et al. 22429 (holotype: RB (two sheets - barcodes 00556325! and 00556326!); isotypes: CAS, UC(ex-LEA), MG!, NY!, UB!).

Phoradendron maguirei Rizzini, Arq. Jard. Bot. Rio de Janeiro 24: 33. 1980. Type: BRAZIL. MINAS GERAIS: Serra do Itabirito, 45 km SE of Belo Horizonte, 12.II.1968, H.S. Irwin et al. 19911 (holotype: RB - barcode 00540595!; isotypes: R, NY!, UB!).

Phoradendron maguirei var. parvulum Rizzini, Arq. Jard. Bot. Rio de Janeiro 24: 34. 1980. Type: BRAZIL. MINAS GERAIS: 14 km SW of Diamantina on road to Gouveia, 5.II.1972, W.R. Anderson et al. 35491 (holotype: RB - barcode 00540597!; isotypes: CAS, UC(ex-LEA), NY!, UB!).

Phoradendron aurato-ochraceum Rizzini, Ernstia 32: 7. 1985. Type: BRAZIL. MINAS GERAIS: Diamantina, km-165 on road to Conselheiro Mata, on Lychnophora, 18.VII.1980, J. Semir et al. 234 (holotype: RB - barcode 00540571!; isotype: SPF!). Fig. 2g

Phoradendron caripense can be recognized by its erect habit and monopodial/percurrent, branches, with cataphylls only on the proximal lateral internodes; leaves generally cuneate or obovate, with a short petiole; triseriate inflorescences with 2–3 pairs of sterile bracts at the base. This species is very similar to P. dimerostachys, a monoecious species, but P. caripensis is dioecious. (See more comments under P. dimerostachys).

Iconography: Rizzini (1980, Figs. 6-1, 7-1), Kuijt (2003, Fig. 72).

It is distributed in South America - Brazil (AM, BA, MG, MT, PA, PB) (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.).

2.7. Phoradendron chrysocladon A. Gray, U.S. Explor. Exped. [Bot., Phanerogam.] 15(1): 43. 1854. Type: BRAZIL. RIO DE JANEIRO: near to Rio de Janeiro, 1838-42, Wilkes Exped. s.n. (holotype: US - barcode 00105562!; isotypes: F, GH!, P).

Phoradendron reticulatum Urb., Bot. Jahrb. Syst. 23, Beibl. 57: 12. 1897. Type: BRAZIL. RIO DE JANEIRO: Villa Nova, 7.V.1874, A.F.M. Glaziou 7664 (lectotype, designated by Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.: K; isolectotypes: C (= Trelease, 1916, Fig. 230), F, P!(3x), S!). Fig. 2h

Phoradendron chrysocladon is recognized by the monopodial/percurrent, branches, ellipsoidal and bi-keeled when young; sterile cataphylls present in all internodes; leaves with palmate and conspicuous veins; triseriate and clavate inflorescences. In nature, it presents a dark green to yellow-gold colour. It is often confused, in the collections, with P. piperoides, which has veins pinnate. (See comments under P. laxiflorum and P. undulatum).

Iconography: Kuijt (2003, Fig. 76), Kuijt (2005, Fig. 410), Rigon & Cervi (2013, Figs. 3e-f), Dettke & Waechter (2014, Figs. 2f-i).

It is distributed in North America - Mexico, Central America (inclusing Caribbean) and South America - Bolivia, Brazil (AL, BA, CE, ES, MG, PB, PE, PR, RJ, SC, SE, SP), Colombia, Ecuador, Guyana, Peru and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Caires 2020Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... ).

2.8. Phoradendron congestum Trel., Phoradendron 80, Figs. 103, 104. 1916. Phoradendron rubrum var. longifolium Eichler in Martius, Fl. bras. 5(2): 121. 1868. Type: BRAZIL. GOIÁS: “In prov. Goyaz”, G. Gardner 3764 (holotype: BR - barcode 000000550652! (= Trelease, 1916, Fig. 103); isotypes: BM!, K!(2x), P!(2x)).

Phoradendron andersonii Rizzini, Arq. Jard. Bot. Rio de Janeiro 24: 29. 1980. Type: BRAZIL. GOIÁS: Cerrado, 8 km S of Niquelândia, on Piptocarpha, 23.I.1972, H.S. Irwin et al. 34866 (holotype: RB – barcode 00540563!; isotypes: UC(ex-LEA), NY, UB!), syn. nov. Figs. 2i; 6

Phoradendron congestum is recognized by its large leaves, lanceolate to oblong-lanceolate to falcate and by short and congested inflorescences with 2–4 fertile biseriate segments (5–9 flowers per fertile bract). Under this concept, we include P. andersonii as a new synonym due to the overlapping morphological characteristics. Phoradendron andersonii was initially considered a synonym of P. perrottetii (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.), and was latter reestablished as a separate species by Caires & Proença (2007)Caires CS & Proença CEB (2007) Two new synonyms and a species reinstated in Phoradendron Nuttall (Viscaceae). Acta Botanica Brasilica 21: 379-382. due to the differences to P. perrottetii, such as the shape of young internodes (compressed in P. andersonii × cylindrical in P. perrottetii), the length of spikes (short, maximum 2.5 cm × long, maximum 7.5 cm) and type of spike (biseriate × triseriate). (See comments under P. bicarinatum).

Material selected for illustration (Fig. 6): Brasília, Parque Nacional de Brasília, margin of EPCT 001 road, on Banisteriopsis latifolia (A. Juss.) B. Gates, 11.VI.2002, C.S. Caires & V.C. Mendes 160 (UB). Sobradinho, margin of DF-150 road, inside an abandoned house lot, on Melia azedarach L., 26.III.2002, C.S. Caires & V.C. Mendes 77 (UB).

Iconography: Rizzini (1980, Fig. 6-3), Kuijt (2003, Fig. 80), Caires & Proença (2005, Fig. 2c).

It is distributed in South America - Brazil (AM, BA, DF, GO, MG, MT, PA, SP) and Ecuador (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Caires & Proença 2007Caires CS & Proença CEB (2007) Two new synonyms and a species reinstated in Phoradendron Nuttall (Viscaceae). Acta Botanica Brasilica 21: 379-382.; Dettke & Waechter 2014Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.).

2.9. Phoradendron coriaceum Mart. ex Eichler in Martius, Fl. bras. 5(2): 121. 1868. Type: BRAZIL. MINAS GERAIS: “in campis Taboleiro prov. Minarum, Alto dos Boys,” C.F.P. Martius s.n. (lectotype, designated by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: M - barcode 174063! (= Trelease, 1916, Fig. 107a); isolectotype: M!).

Phoradendron coriaceum var. quintense Urb., Bot. Jahrb. Syst. 23, Beibl. 57: 13. 1897. Type: BRAZIL. RIO DE JANEIRO: Quinta, 15.II.1870, A.F.M. Glaziou 4010 (holotype: B, destroyed (= Trelease, 1916, Fig. 107b), lectotype, designated by Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.: K - barcode 00601409!; isotypes: P!(3x)).

Phoradendron apiciflorum Rizzini, Rev. Fac. Agron. (Maracay) 8(3): 86. 1975. Type: BRAZIL. MINAS GERAIS: cerrado 4 km N of Patrocínio, 31.I.1970, H.S. Irwin et al. 25724 (holotype: RB - barcode 00540568!; isotypes: COL, F!, US!), syn. nov.

Phoradendron luisense Rizzini, Fl. Venezuela 4(2): 255. 1982. Phoradendron luisense var. retusum Rizzini, Fl. Venezuela 4(2): 257. 1982, nom. illeg. Type: VENEZUELA. FALCÓN: Sierra de San Luis, 1.5 km W of Macuquita, 600 m, 20.XII.1978, H. van der Werff & R. Wingfield 3260 (holotype: RB - barcode 00205522!; isotypes: MO!, U!), syn. nov.

Phoradendron bahiense Kuijt, Syst. Bot. Monogr. 66: 99, Fig. 38. 2003. Type: BRAZIL. BAHIA: Maracás, BA-026, 6 km SW of Maracás, 5.V.1979, S.A. Mori & T.S. dos Santos 11798 (holotype: UC (ex-LEA), isotypes: CEPEC!, NY, RB!), syn. nov.

Phoradendron irwinianum Kuijt, Syst. Bot. Monogr. 66: 248, Fig. 142. 2003. Type: BRAZIL. DISTRITO FEDERAL: 5 km W of Formosa, road to Brasília, 8.X.1965, H.S. Irwin et al. 9060 (holotype, F = F Neg 70628!; isotypes: IAN!, NY!, RB!, UB!, US!), syn. nov. Figs. 1m; 2j; 7-8

Phoradendron coriaceum is characterized by the robust aspect of stems and leaves, often exhibiting a dark brown or black color in sicco. This species presents monopodial/percurrent branches, more rarely with dichotomies; cataphylls and bracts of the inflorescences generally with whitish margins; the nodes are frequently thickened by the formation of many inflorescences. It is a dioecious species with staminate and pistillate, triseriate, inflorescences. It can be confused with P. bathyoryctum, but it is distinguished by the smaller leaves, cataphylls with whitish margins and pistillate inflorescences with all immature fruits in the fertile bract (in P. bathyoryctum, the staminate flowers of the apex are dehiscent and bracts are incomplete in fruiting).

In this treatment, we include P. apiciflorum, P. bahiense, P. irwinianum and P. luisense, as new synonyms of P. coriaceum. Phoradendron apiciflorum is distinguished by presenting a distal placement of the fruits, a condition also seen in some populations of P. coriaceum with a predominantly southern distribution (Dettke & Waechter 2014Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985., Fig. 3e). The nodes in P. apiciflorum are thickened, similarly to P. coriaceum. We also include P. irwinianum, recognized by Caires & Proença (2007)Caires CS & Proença CEB (2007) Two new synonyms and a species reinstated in Phoradendron Nuttall (Viscaceae). Acta Botanica Brasilica 21: 379-382. as synonym of P. apiciflorum, as a new synonym of P. coriaceum. Phoradendron bahiense also has a thickened node and a white-margined cataphyll; and fewer flowers per fertile, pistillate, bract (3(–5)) which are common in P. coriaceum, and therefore we attribute it to variation within the population. The Venezuelan species P. luisense also shows black color in sicco, white-margined cataphylls and similar inflorescence morphology, overlapping with P. coriaceum, for which we propose it as a new synonym.

Material selected for illustration (Figs. 7,8): Brasília, APA de Cafuringa, Campina Verde Farm, on Machaerium opacum, 11.VII.2002, C.S. Caires & C.L. Ramalho 181 (UB); 5 km W of Formosa, road to Brasília, 8.X.1965, H.S. Irwin et al. 9060 (UB).

Iconography: Kuijt (2003, Figs. 32, 38, 81, 171), Caires & Proença (2005, Figs. 2a-b,d), Dettke & Waechter (2013, Figs. 73-79); Rigon & Cervi (2013, Figs. 4i-l), Dettke & Waechter (2014, Figs. 3a-e).

It is distributed in South America - Argentina, Brazil (AL, AM, BA, ES, MG, MT, PI, PR, RS, RJ, SC, SP), Paraguay and Venezuela (Dettke & Waechter 2014Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.).

2.10. Phoradendron craspedophyllum Eichler in Martius, Fl. bras. 5(2): 124, Fig. 37-3. 1868. Type: BRAZIL. SÃO PAULO: F. Sello 155 (lectotype: B, designated by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: B, destroyed (= Trelease, 1916, Fig. 124a); lectotype, designated by Kuijt (1994)Kuijt J (1994) Typifcation of the names of new world mistletoe taxa (Loranthaceae and Viscaceae) described by Martius and Eichler. Taxon 43: 187-199.: P - barcode 00756337!; isolectotype: BR!). Fig. 2k

This species is distinct from the other species of Phoradendron for the sympodial/dichotomous branching, with one pair of fertile cataphylls, small leaves with palmate veins and terminal and axillary biseriate inflorescences, composed by three flowers, where the apical one is staminate. It may be confused with P. mucronatum or P. harleyi, which are species with monopodial, percurrent, branches and larger leaves.

Iconography: Eichler (1868, Fig. 37-3), Kuijt (2003, Fig. 82), Dettke & Waechter (2013, Figs. 80-82), Rigon & Cervi (2013, Figs. 5h-j), Dettke & Waechter (2014, Figs. 3f-j).

It is distributed in South America - Brazil (PR, RS, SC, SP) (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Waechter 2014Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.).

2.11. Phoradendron crassifolium (Pohl ex DC.) Eichler in Martius, Fl. bras. 5(2): 125, Fig. 40. 1868. Viscum crassifolium Pohl ex DC., Prodr. 4: 280. 1830. Type: BRAZIL. GOIÁS: “Serra d’Ourada,” 1839, J.E. Pohl 457 (holotype: G-DC! (= Trelease, 1916, Fig. 213a); isotype: BR!).

Phoradendron pepericarpum A. Gray, U.S. Explor. Exped. [Bot., Phanerogam.] 15(1): 742. 1854. Type: BRAZIL. RIO DE JANEIRO: Organ Mountains, Wilkes Exped. s.n. (lectotype, designated by Trelease, 1916: US - barcode 00105616!; isolectotypes: GH!, NY!).

Phoradendron crassifolium var. parvifolium Eichler in Martius, Fl.bras. 5(2): 125. 1868. Type: BRAZIL. MINAS GERAIS: “ad Copacabana, prov. Minarum, B. Luschnath s.n. (holotype: not located; lectotype, hic designatus: MO - barcode 00204716!).

Phoradendron crassifolium var. multiflorum Eichler in Martius, Fl.bras. 5(2): 125. 1868. Type: BRAZIL. MINAS GERAIS: Sabará, L. Riedel s.n. (lectotype, designated by Trelease, 1916: M). Figs. 1k; 2l

One of the most common mistletoes in Brazil, P. crassifolium, is easily recognized by the presence of 2–5 pairs of fertile cataphylls and large leaves with usually five very prominent palmate veins (3–7 primary veins). (See comments under P. berteroanum, P. inaequidentatum and P. undulatum).

Iconography: Eichler (1868, Fig. 40), Rizzini (1968b, Fig. 7), Rizzini (1982, Fig. 58), Rizzini (1995, Figs. 16-17), Kuijt (2003, Fig. 84), Caires & Proença (2005, Fig. 2f), Kuijt (2005, Fig. 400), Dettke & Waechter (2013, Figs. 83-87), Rigon & Cervi (2013, Figs. 4a-e), Dettke & Waechter (2014, Figs. 3k-p), Caires (2017, Figs. 1c-d, 2e-i).

It is distributed in Central America (including Caribbean) and South America - Bolivia, Brazil (all states), Colombia, Ecuador, French Guiana, Guyana, Paraguay, Suriname and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.).

2.12. Phoradendron dimerostachys Rizzini, Arq. Jard. Bot. Rio de Janeiro 24: 31. 1980. Type: BRAZIL. BAHIA: Serra da Água de Rega, 24 km N of Seabra on road to Água de Rega, 24.II.1971, H.S. Irwin et al. 30967 (holotype: RB - barcode 00540578!; isotypes: CAS, UC(ex-LEA), MG!, NY!, UB!).

Phoradendron dimerostachys is an intriguing species only known for the type specimen, which presents monopodial/percurrent branches; and, generally, elliptical leaves with a rounded apex, and triseriate inflorescences with the single entire, basal fertile internode bearing staminate flowers. This species is very similar to P. caripense, which is a dioecious species. However, we have little doubt that, with the appearance of new collections, they will be recognized as synonyms.

Iconography: Rizzini (1980, Figs. 6-2, 7-2), Kuijt (2003, Fig. 89).

It is distributed in South America - Brazil (BA) (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.).

2.13. Phoradendron diminutivum E.A. Kellogg, Novon 6: 41, Fig. 5. 1996. Type: VENEZUELA. AMAZONAS: Rio Negro, Rio Pasimoni, between its mouth and its junction with the Rio Baria and the Rio Yatua, on Vochysia, 23–25.VII.1984, G. Davidse 27772 (holotype: GH; isotypes: UC(ex-LEA), MO).

Phoradendron diminutivum is a curious species where cataphylls may be present, or not, on the main axis. This species has a combination of monopodial/percurrent and sympodial/dichotomous branches, elliptical or obovate leaves, biseriate inflorescences, in axillary and terminal position, and small fruits with erect petals. Phoradendron diminutivum somewhat resembles P. quadrangulare in the leaf morphology, but possesses circular or elliptical internodes, and P. quadrangulare has rhombic or quadrangular internodes, and never presents cataphylls on the main axis. The growth patterns, cataphylls, inflorescences, and fruits resemble D. warmingii, and the type specimen (not seen) was collected parasiting Vochysia species. In the Brazilian specimens, the leaves are narrower than those of D. warmingii, although some are obovate or circular (as seen in Acevedo-Rodriguez et al. 8255 (NY!).

Iconography: Kuijt & Kellogg (1996Kuijt J & Kellogg EA (1996) Miscellaneous Mistletoes Notes, 20-36. Novon 6: 33-53., Fig. 5), Kuijt (2003, Fig. 90).

It is distributed in South America - Brazil (AM) and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.).

2.14. Phoradendron dipterum Eichler in Martius, Fl. bras. 5(2): 109. 1868. Type: BRAZIL. CEARÁ: X.1838, G. Gardner 1672 (lectotype, designated by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: W, destroyed (= Trelease, 1916: Fig. 89); lectotype, first step designated by Kuijt (1994)Kuijt J (1994) Typifcation of the names of new world mistletoe taxa (Loranthaceae and Viscaceae) described by Martius and Eichler. Taxon 43: 187-199.: P, second step, hic designatus: P - barcode 002441994!; isolectotypes: BM!(2x), GH!, K!(2x), P!). Figs. 2m-o; 9

Phoradendron amplexicaule Eichler in Martius, Fl.bras. 5(2): 110. 1868. Type: BRAZIL. Without locality, 1858, M. Weddell s.n. (holotype: not located; lectotype, designated by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: G - barcode 00006176! (= Trelease, 1916, Fig. 94b)).

Phoradendron multifoveolatum Eichler in Martius, Fl. bras. 5(2): 110, Fig. 34. 1868. Type: BRAZIL. BAHIA: in caatingas usque Pao d’Espinhos, C.F.P. Martius s.n. (lectotype, designated by Trelease, 1916: M! (= Trelease, 1916, Fig. 90)).

Phoradendron crulsii Urb., Bot. Jarb. Syst. 23, Beibl. 57: 11. 1897. Type: BRAZIL. GOIÁS: Près de Jaragua, 22.VII.1895, A.F.M. Glaziou 22021 (holotype: B, destroyed (= Trelease, 1916, Fig. 94a); lectotype, hic designatus: P - barcode 00756325!; isolectotypes: K!, P!).

Phoradendron glaziovii Urb., Bot. Jahrb. Syst. 23, Beibl. 57: 12. 1897. Type: BRAZIL. RIO DE JANEIRO: Tijuca, 27.II.1870, A.F.M. Glaziou 4004 (holotype: B, destroyed (= Trelease, 1916, Fig. 96a); lectotype, hic designatus: P - barcode 756324!; isolectotypes: K!, P!(2x)).

Phoradendron anamariae Rizzini ex Kuijt, Syst. Bot. Monogr. 66: 78, Fig. 23. 2003. Phoradendron anamariae Rizzini, Rev. Brasil. Biol. 51: 459. 1991, nom. nud. [without holotype cited]. Type: BRAZIL. GOIÁS: Alto Paraíso de Goiás, Parque Nacional Chapada dos Veadeiros, near Cachoeira de Rio Preto, 6.II.1987, J.R. Pirani et al. 1701 (holotype: UC(ex-LEA)!; isotypes: K, RB!, SPF!), syn. nov.

Phoradendron dipterum is an epiparasite of other species of Phoradendron, although it can also parasite (rarely) other species of angiosperms. It is easily recognized by its winged stem, with two or four distinct wings, absence of cataphylls in the main internodes, and triseriate clavate inflorescences with staminate flowers on the first to third positions on the fertile bract. Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643. described P. anamariae based on a single collection, and admitted that it can be a “depauperate form of the hyperparasitic P. dipterum”. We agree with that, and consider them as the same species. Phoradendron dipterum presents great morphological variation among the populations in Brazil, with variation in the size of the plant, number of flowers and color of the mature fruits. In P. dipterum, the staminate apical flowers are the first to open and soon fall off, and, for this, they are often missing from herbarium specimens. This may have led Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643. to interpret P. anamariae as a possibly dioecious species.

The lectotype designated by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p. for P. dipterum was destroyed during the second World War, and Kuijt (1994)Kuijt J (1994) Typifcation of the names of new world mistletoe taxa (Loranthaceae and Viscaceae) described by Martius and Eichler. Taxon 43: 187-199. designated a new lectotype from P herbarium. Two sheets were located at P, one of which we now select a second step of lectotypification, according to the Code (Turland et al. 2018Turland NJ, Wiersema JH, Barrie FR, Greuter W, Hawksworth DL, Herendeen PS, Knapp S, Kusber WH, Li DZ, Marhold K, May TW, McNeill J, Monro AM, Prado J, Price MJ & Smith GF (2018) International code of nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Koeltz Botanical Books, Glashütten. 254p., Art. 9.17).

For P. amplexicaule, Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643. indicated as isotypes the herbarium sheets at P and G herbaria. Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p. only indicated the specimen at G as type, which presents the amplexus base of the leaves, as described in the protologue (“basi cordata sessilibus amplexicaulibus”, Eichler 1868Eichler AW (1868) Loranthaceae. In: Martius CFP (ed.) Flora brasiliensis. Fleicher, Leipzig. Vol. 5, pars 2, pp. 1-136, t.1-44.: 110), collected in 1858. The specimen at P (Brazil, Minas Gerais, XI.1843, M. Weddell 1127a) does not correspond to original material, because the different collection date and the leaves are distinctly petiolate.

For P. crulsii and P. glaziovii, the Urban’s holotype from B herbarium was also destroyed during the second World War and fo this reason, we here designate the lectotypes for these two names.

Material selected for illustration (Fig. 9): Brazlândia, DF-180 road to S.D.V.S.A. Farm 098-02, on Phoradendron mucronatum (DC.) Krug & Urb., 26.III.2002, C.S. Caires & V.C. Mendes 92 (UB).

Iconography: Eichler (1868, Fig. 34), Rizzini (1956, Fig. 16-2), Rizzini (1995, Fig. 18), Kuijt (2003, Fig. 91), Caires & Proença (2005, Figs. 2i-j), Dettke & Waechter (2013, Figs. 88-95), Rigon & Cervi (2013, Figs. 4f-h), Dettke & Waechter (2014, Figs. 4a-d), Caires (2017, Figs. 2j-l).

It is distributed in North America - Mexico, Central America (including Caribbean) and South America - Argentina, Brazil (BA, CE, DF, ES, GO, MG, MT, PA, PE, PR, RJ, RS, SC, SP, TO), Colombia, Ecuador, Paraguay, Peru, Suriname and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Caires 2020Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... ).

2.15. Phoradendron ensifolium (Pohl ex DC.) Eichler in Martius, Fl. bras. 5(2): 114, Fig. 35-2. 1868. Viscum ensifolium Pohl ex DC., Prodr. 4: 281. 1830. Type: BRAZIL. MINAS GERAIS: Barbacena, J.E. Pohl 106 (holotype: G-DC! = Trelease, 1916, Fig.182a).

Phoradendron interruptum (DC.) B.D. Jacks., Ind. Kew. 2: 502. 1895. Viscum interruptum DC., Prodr. 4: 282. 1830. Phoradendron lanceolato-ellipticum Eichler in Martius, Fl. bras. 5(2): 114, pl. 35, Fig. 1. 1868, nom. superfl. Type: BRAZIL. GOIÁS: Padre Luiz Faria, J.E. Pohl s.n. (holotype: G-DC! = Trelease, 1916: Fig.89a); isotypes: BR!, ILL!, M!, W, destroyed (= Trelease, 1916, Fig.89b)).

Phoradendron nitidum (Gardner) Eichler in Martius, Fl. bras. 5(2): 113. 1868. Viscum nitidum Gardner in Hooker, London J. Bot. 4: 105. 1845. Type: BRAZIL. RIO DE JANEIRO: Organ Mountains, II.1837, G. Gardner 436 (holotype: BM - barcode 0049232!; isotypes: K!(2x), P, W! (= Trelease 1916Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: Fig. 123a)), syn. nov.

Phoradendron habrostachyum Eichler in Martius, Fl. bras. 5(2): 111. 1868. Type: BRAZIL. MINAS GERAIS: in silvis usque Ouro Preto, C.F.P. Martius s.n. (holotype: M - barcode 0174061! (= Trelease 1916Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: Fig. 109)), syn. nov.

Phoradendron linearifolium Eichler in Martius, Fl. bras. 5(2): 115, Fig. 36. 1868. Type: BRAZIL. RIO DE JANEIRO: near to Rio de Janeiro, L. Riedel s.n. (lectotype, designated by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: G (= Trelease, 1916, Fig. 181a); isolectotypes: K!(2x), P, US!).

Phoradendron selloi Eichler in Martius, Fl. bras. 5(2): 116. 1868. Type: BRAZIL. “Brasiliae austro-orientalis”, F. Sello 122 (holotype: B, destroyed (= Trelease, 1916, Fig. 123b)). BRAZIL. Rio de Janeiro: A.F.M. Glaziou 4013 (neotype, designated by Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.: K - barcode 00568331!; isoneotypes: P!(3x)).

Phoradendron falcifrons (Hook. & Arn.) Eichler in Martius, Fl. bras. 5(2): 134m. 1868. Viscum falcifrons Hook. & Arn., Bot. Misc. 3: 356. 1833. Type: URUGUAY. On laurels, J. Tweedie s.n. (holotype: K - barcode 00601433! (= Trelease 1916Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: Fig. 85); isotype: E!).

Phoradendron holoxanthum var. corallispicum Trel., Phoradendron 90, Fig. 122b. 1916. Type: BRAZIL. RIO DE JANEIRO: A.F.M. Glaziou 8729 (holotype: G - barcode 00301897! (= Trelease, 1916, Fig. 122b); isotypes: K!, P!(2x)), syn. nov. Fig. 2p

Phoradendron ensifolium is a dioecious species, easily recognized by the pending branches with monopodial/percurrent growth pattern, eventually with dichotomies, cataphylls only on the proximal lateral internodes, elongated, oblong, linear or falcate leaves, and triseriate, staminate and pistillate inflorescences. Some morphotypes of Phoradendron ensifolium, with smaller and wider leaves, can be confused with P. perrottetii; however, they can be differentiated by pending branches (generally erect in P. perrottetii) often with more cataphylls on the lateral proximal branches (2–5 vs. 1–2), and pistillate inflorescences with less flowers per fertile bract (4–10 vs. 12–18). (see comments under P. holoxanthum).

Dettke & Waechter (2014)Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985. expanded the circumscription of the species, including P. falcifrons, P. interruptum and P. linearifolium under its synonymy. After revising the type specimen of P. nitidum, we now additionally included them in P. ensifolium, because no sufficient morphological differences were found to sustain it as a separate species. Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643. uses the simple peduncle (one pair of sterile bracts) in P. nitidum to differentiate it from P. ensifolium, which has a composed one. Yet, as we analyze the type specimen, we find variation on the same branch, with one or two pairs of sterile bracts, as already described by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.. Also, Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p. recognized the similarity between P. nitidum and P. selloi, which was already considered synonym of P. falcifrons by Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643., differing only by the number of flowers.

Here, we also consider P. habrostachyum as a synonym of P. ensifolium. The holotype used by Eichler (1868)Eichler AW (1868) Loranthaceae. In: Martius CFP (ed.) Flora brasiliensis. Fleicher, Leipzig. Vol. 5, pars 2, pp. 1-136, t.1-44. in describing this species is very fragmented, but it presents parts of staminate and pistillate individuals, which are possible to assign to the inflorescence and leaf morphology of P. ensifolium. The stems are short, and with distinctly dilated nodes, similarly to the stems seen in the P. nitidum and P. selloi morphotypes. Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643. and Dettke & Waechter (2014)Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985. cited as P. habrostachyum, the collection of pistillate individuals Y. Mexia 5463 (F!, US!), with longer internodes and white fruits, which clearly corresponds to P. ensifolium. However, collections of pistillate individuals from Paraguay (Dettke & Waechter 2014Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.) and Brazil - Rio Grande do Sul State (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Waechter 2014Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.), do not correspond to the type of P. habrostachyum, remaining with an uncertain placement.

Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643. considered P. holoxanthum var. corallispicum as an uncertainly placed taxon, and we agree, that, indeed, it is not P. holoxanthum. The analysis of more duplicates than those available for Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643. exposed similarities of the stems, leaves and inflorescences in the type specimen of this taxon, with fruiting individuals of P. nitidum and P. selloi (Sello 122 = Trelease, 1916, Fig. 123b). Hence, we consider it as a new synonym of P. ensifolium.

Iconography: Eichler (1868, Figs. 35, 36), Rizzini (1956, Fig. 12), Rizzini (1968b, Fig. 8), Kuijt (2003, Figs. 97, 103, 163), Caires & Proença (2005, Fig. 2k), Dettke & Waechter (2013, Figs. 96-103), Rigon & Cervi (2013, Figs. 5a-g), Dettke & Waechter (2014, Figs. 4e-k).

It is distributed in South America - Argentina, Bolivia, Brazil (AL, BA, CE, DF, ES, GO, MG, MS, MT, PA, PB, PE, PR, RJ, RN, RS, SC, SE, SP), Paraguay and Uruguay (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Waechter 2014Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.; Dettke & Caires 2020Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... ).

2.16. Phoradendron fragile Urb., Bot. Jahrb. Syst. 23, Beibl. 57: 13. 1897. Type: BRAZIL. RIO DE JANEIRO: near Rio de Janeiro, XI.1879, A.F.M. Glaziou 10898 (lectotype, designated by Trelease, 1916: B, destroyed (= Trelease, 1916, Fig. 210a); lectotype, hic designatus: K - barcode 00601385!; isolectotypes: P!(2x)). Figs. 3a; 10

Phoradendron fragile is one of the small aphyllous species of Phoradendron in Brazil. This species is distinct from the others by its erect branches and short terete stems (0.8–3.5 cm long); also, they are dioecious plants with triseriate inflorescences (4–10 flowers per fertile bract) and orange and reddish fruits.

Since the lectotype selected by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p. was destroyed during the second World War, we have selected a new one from K herbarium.

Material selected for illustration (Fig. 10): Brasília, Catetinho, I.1982, B.A.S. Pereira 162 (IBGE); Estação Florestal Cabeça de Veado, XII.1980, E.P. Heringer et al. 5861 (IBGE); Várzea Bonita, II.1978, E.P. Heringer 16832 (IBGE).

Iconography: Rizzini (1956, Fig. 25), Rizzini (1980, Fig. 8), Kuijt (2003, Fig. 114), Caires & Proença (2005, Fig. 1d).

It is distributed in South America - Brazil (DF, GO, MG, RJ, SP) (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Caires 2020Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... ).

2.17. Phoradendron harleyi Kuijt, Syst. Bot. Monogr. 66: 224, Fig. 125. 2003. Type: BRAZIL. BAHIA: Rio de Contas, Pico das Almas, SE of Campo do Queiroz, 13º32’S, 41º57’W, on Rubiaceae, 24.XII.1988, R.M. Harley et al. 27359 (holotype: SP - barcode 001695!; isotypes: CEPEC!, K!, NY!, SPF!, UC(ex-LEA)).

Phoradendron harleyi can be recognized by the combination of monopodial/percurrent and sympodial/dichotomous branches, presence of one pair of fertile cataphylls in all internodes, elliptical leaves and biseriate inflorescences with 3–5 flowers per fertile bract. This species is very close to P. craspedophyllum, which has a predominantly southern distribution, is a smaller plant, with more dichotomies in the stems and persistently only three flowers per fertile bract. Perhaps future evidence will demonstrate they are the same species.

Iconography: Kuijt (2003, Fig. 125).

It is distributed in South America - Brazil (BA) (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.).

2.18. Phoradendron hexastichum (DC.) Griseb., Fl. Brit. W. I. 313. 1860. Viscum hexastichum DC., Prodr. 4: 282. 1830. Type: CUBA. LA HABANA: Havana, 1825, A. de la Ossa s.n. (holotype: G-DC! (= Trelease, 1916, Fig. 200a).

Phoradendron hexastichum var. longispicum Eichler in Martius, Fl. bras. 5(2): 129. 1868. TYPE: BRAZIL. AMAZONAS: prope San Gabriel da Cachoeira, usque Rio Negro, R. Spruce 2112 (holotype: M; isotypes: BM!(2x), G! (= Trelease, 1916, Fig. 204b) (3x), GH!, K, NY!, P!(2x), TCD!).

Phoradendron productipes Trel., Phoradendron 138, Fig. 204a. 1916. Type: BRAZIL. CEARÁ: X.1838, G. Gardner 1676 (holotype: W, destroyed (= Trelease, 1916, Fig. 204a); lectotype, hic designatus, K - barcode 0016683!; isolectotypes: BM, GH!, K!, L, NY!, P!(3x)).

Phoradendron brachyklados Rizzini, Ernstia 32: 2. 1985. Type: BRAZIL. DISTRITO FEDERAL: Brasília, 25.X.1979, E.P. Heringer et al. 2600 (holotype: IBGE!; isotypes: RB!). Figs. 3b; 11

Phoradendron hexastichum is characterized by monopodial/percurrent branches, cataphylls only on the proximal lateral branches, leaves with pinnate veins and triseriate inflorescences. This species can be confused with P. racemosum, which has dichotomous branches and biseriate inflorescences, or with P. pteroneuron, which has shorter leaves and biseriate inflorescences. (See comments at P. juruanum and P. oliveirae).

Material selected for illustration (Fig. 11): Brasília, Núcleo Rural Tabatinga, DF-250 road to S.D.V.S.A Farm 08, Sete Veredas Farm, on Richeria grandis Vahl., 6.VIII.2002, C.S. Caires et al. 189 (UB); APA de Cafuringa, Dois Irmãos Farm, on Richeria grandis Vahl., 21.VIII.2002, C.S. Caires et al. 209 (UB). Guará, EPTG road between Guará e Taguatinga, margin to Vicente Pires stream (near the bridge), on Calophyllum brasiliense Camb., 19.III.2002, C.S. Caires et al. 65 (UB).

Iconography: Eichler (1868, Fig. 43-1), Rizzini (1956, Fig. 29-2), Rizzini (1982, Fig. 64), Kuijt (2003, Fig. 133), Caires & Proença (2005, Figs. 1e-g), Kuijt (2005, Fig. 402), Dettke & Waechter (2013, Figs. 104-107), Dettke & Waechter (2014, Figs. 5e-h).

It is distributed in North America - Mexico, Central America (including Caribbean) and South America - Bolivia, Brazil (AM, BA, CE, DF, GO, MG, RR, RS, SC, SP, TO), Colombia, Ecuador, Guyana, Peru and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Waechter 2014Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.).

2.19. Phoradendron holoxanthum Eichler in Martius, Fl. bras. 5(2): 116. 1868. Type: BRAZIL. “Brasiliae austro-orientalis,” F. Sello 5847 (lectotype, designated by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: B, destroyed (= Trelease, 1916, Fig. 122a); lectotype, hic designatus: P - barcode 05010576!). Fig. 3c

Phoradendron holoxanthum is recognized by its erect habit, with a combination of monopodial, percurrent, and sympodial, dichotomous, growth patterns, cataphylls only on the proximal lateral internodes, obovate leaves and triseriate inflorescences in axillary and terminal positions. This is a dioecious species that may be confused with P. ensifolium, which has a pendulous habit, and longer and falcate leaves.

A duplicate of the F. Sello collection was found in P herbarium and, is, therefore, here designated as lectotype for P. holoxanthum, rendering the neotypification of Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643. superfluous.

Iconography: Kuijt (2003, Fig. 137), Dettke & Waechter (2013, Figs. 108-113); Dettke & Waechter (2014, Figs. 5i-o).

It is distributed in South America - Brazil (RS) (Dettke & Waechter 2014Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.).

2.20. Phoradendron inaequidentatum Rusby, Bull. Torrey Bot. Club 27: 137. 1900. Dendrophthora inaequidentata (Rusby) Trel., Phoradendron 218. 1916. Type: BOLIVIA. LA PAZ: Guanai, V.1886, H.H. Rusby 1544 (lectotype, hic designatus: NY - barcode 00285343!; isolectotypes: F!, GH!, NY!(2x), PH!, US!).

Phoradendron prancei Rizzini, Rev. Fac. Agron. Maracay 8(3): 89. 1975. TYPE: BRAZIL. AMAZONAS: km 10 between Manaus and Caracarai, near Igarape Leão, on Moraceae, 19.X.1966, C.K. Allen et al. 2713 (holotype: RB (two sheets - barcodes 00540608! and 00545660!); isotypes: K, MO!, NY!, R, U, US!).

This species is easily recognized by the presence of one pair of fertile cataphylls, leaves with palmate veins, the presence of axillary and terminal inflorescences, uniseriate (except for the biseriate basal segment) and by large fruits with opening petals. It can be confused with P. crassifolium, which presents more than two pairs of fertile cataphylls, biseriate inflorescences and smaller fruits, and also with Dendrophthora warmingii, which doesn’t have fertile cataphylls and presents bi-triseriate inflorescences. (See comments under P. nothropiae).

In what concerns P. inaequidentatum, the type collection has three syntypes at NY herbarium and one of which we have selected as a lectotype (Turland et al. 2018Turland NJ, Wiersema JH, Barrie FR, Greuter W, Hawksworth DL, Herendeen PS, Knapp S, Kusber WH, Li DZ, Marhold K, May TW, McNeill J, Monro AM, Prado J, Price MJ & Smith GF (2018) International code of nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Koeltz Botanical Books, Glashütten. 254p., Art. 9.11).

Iconography: Kuijt (2003, Fig. 139), Kuijt (2005, Fig. 412), Dettke & Waechter (2014, Figs. 6a-d).

It is distributed in Central America - Panama, and South America - Bolivia, Brazil (AC, AM, AP, BA, MT, PA, RO, RS), Colombia, Ecuador, French Guiana, Guyana, Peru and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Waechter 2014Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.).

2.21. Phoradendron juruanum Kuijt, Syst. Bot. Monogr. 66: 251, Fig. 145. 2003. Type: BRAZIL. AMAZONAS: Basin of Rio Jurua, Manariao, B.A. Krukoff 4592 (holotype: MICH; isotypes: F!, GH!, K!, LP!, MICH, MO!, NY!, S!, U!, UC!, US!).

Phoradendron juruanum is characterized by monopodial/percurrent branches, with cataphylls in all internodes, leaves with pinnate veins and triseriate clavate inflorescences. It is similar to P. hexastichum in the morphology of leaves and inflorescence, but differs in the presence of cataphylls in all internodes, absent in the main branches of P. hexastichum. Also, it looks like P. piperoides in growth pattern and leaf morphology, differing in the inflorescence arrangement: triseriate in P. juruanum, and biseriate in P. piperoides. (See comments under P. oliveirae).

Iconography: Kuijt (2003, Fig. 145).

It is distributed in South America - Brazil (AM) and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.).

2.22. Phoradendron krukovii Kuijt, Syst. Bot. Monogr. 66: 256, Fig. 150. 2003. Type: BRAZIL. AMAZONAS: basin of Madeira River, Humaita, near Tres Casas, IX–X.1934, B.A. Krukoff 6514 (holotype: BM - barcode 0049240!; isotypes: A!, GH!, IAN!, K!, MICH!, MO!, NY!, S! U, US!).

This species is distinct from the other species of Phoradendron for the predominance of sympodial/dichotomous branches, cataphylls present in all internodes, leaves with palmate veins and acuminate apex, and biseriate inflorescences with 3–4 flowers per fertile bract. However, Phoradendron krukovii bears an older stem with whitish appearance, and longitudinal and elongated fissures, which is unique in the genus. Phoradendron krukovii resembles P. berteroanum and P. burkartii, which do not have a fissured stem, and have acute or rounded leaves, in contrast with the acuminate apex in P. krukovii.

Iconography: Kuijt (2003, Fig. 150).

It is distributed in South America - Brazil (AM) (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.).

2.23. Phoradendron laxiflorum Ule, Verh. Bot. Vereins Prov. Brandenburg 48: 158. 1906. Type: BRAZIL. Amazonas: Juruá River, Santa Clara, Oct 1900, E.H.G. Ule 5251 (holotype: B, destroyed (= Trelease, 1916, Fig. 236); lectotype, hic designatus: MG - barcode 005223!; isolectotypes: CORD!, K!, L).

Phoradendron laxiflorum is easily recognized by its monopodial/percurrent branches, cataphylls in all internodes, leaves with acuminate apex, and three conspicuous palmate veins, uniseriate pistillate inflorescences and triseriate staminate inflorescences. This species closely resembles P. piperoides, except for the leaves with pinnate veins and biseriate inflorescence in P. piperoides. Phoradendron laxiflorum may also be confused with P. chrysocladon which is a monoecious species, with clavate triseriate inflorescence.

As Urban’s holotype at B was destroyed, we now designate the MG sheet as a lectotype.

Iconography: Rizzini (1956, Fig. 17-5), Kuijt (2003, Figs. 156-157).

It is distributed in South America - Bolivia, Brazil (AM), Colombia, Ecuador and Peru (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.).

2.24. Phoradendron macrarthrum Eichler in Martius, Fl. bras. 5(2): 107, 124, Fig. 43-3. 1868. Type: BRAZIL. GOIÁS: Conceição, G. Gardner 3765 (lectotype, first step designated by Kuijt (1994)Kuijt J (1994) Typifcation of the names of new world mistletoe taxa (Loranthaceae and Viscaceae) described by Martius and Eichler. Taxon 43: 187-199.: P, second step, hic designatus: P - barcode 756307!; isolectotypes: BM, G (= Trelease, 1916, Fig. 184a), K, P!, W destroyed (= Trelease, 1916, Fig. 194b)).

Phoradendron harmsianum Ule, Notizbl. Bot. Gart. Berlin-Dahlem 6: 290. 1915. Type: BRAZIL. AMAZONAS: gallery forest near São Marcos on Rio Branco, I.1909, E.H.G. Ule 7889 (holotype: B, destroyed (= F Neg 18188); lectotype, hic designatus: K - barcode 00601401!; isolectotypes: IAN, L, MO), syn. nov.

Phoradendron macrarthrum can be distinguished by the combination of the monopodial/percurrent and sympodial/dichotomous growth patterns, cataphylls only on the proximal lateral branches, distal portion of young stems keeled and flattened, obovate to linear-lanceolate leaves with rounded apex, and biseriate inflorescences with 15–23 flowers per fertile bract.

This species may be confused with P. bathyoryctum, which has less than 15 flowers per fertile bract; or with P. perrottetii, which bears triseriate inflorescence. Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643. correctly called for further studies in the species delimitation, as its distribution is probably wider than presented here. After analyzing the type, we here include P. harmsianum as a new synonym of P. macrarthrum, as we have observed that it is morphologically closer to this species than to P. perrottetii, as Kuijt had suggested (2003).

Iconography: Eichler (1868, Fig. 43-3), Kuijt (173).

It is distributed in South America - Brazil (AM, GO) (this study; Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.).

2.25. Phoradendron mucronatum (DC.) Krug & Urb., Bot. Jahrb. Syst. 24: 352. 1897. Viscum mucronatum DC., Prodr. 4: 282. 1830. Type: DOMINICAN REPUBLIC. C.L.G. Bertero s.n. (holotype: G-DC).

Phoradendron cearense Eichler in Martius, Fl. bras. 5(2): 118. 1868. Type: BRAZIL. CEARÁ: G. Gardner 1675 (lectotype, designated by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: W, destroyed (= Trelease, 1916, Fig. 176a); lectotype, first step designated by Kuijt (1994)Kuijt J (1994) Typifcation of the names of new world mistletoe taxa (Loranthaceae and Viscaceae) described by Martius and Eichler. Taxon 43: 187-199.: P, second step, hic designatus: P - barcode 00756305!; isolectotypes: BM(2x), G!(4x), ILL, K!(2x), L, NY, P!(2x)).

Phoradendron cearense var. minor Eichler in Martius, Fl. bras. 5(2): 118. 1868. TYPE: BRAZIL. Ceará: G. Gardner 1669 (lectotype, first step designated by Kuijt (1994)Kuijt J (1994) Typifcation of the names of new world mistletoe taxa (Loranthaceae and Viscaceae) described by Martius and Eichler. Taxon 43: 187-199.: P, second step, hic designatus: P - barcode 005456217!; isolectotypes: BM(2x), G! (= Trelease, 1916, Fig. 175b)(3x); GH, ILL, K, P!(2x), W destroyed (= Trelease, 1916, Fig. 175a).

Phoradendron emarginatum Martius ex Eichler in Martius, Fl. bras. 5(2): 118, Fig. 38-1. 1868. Phoradendron mucronatum var. emarginatum (Martius ex Eichler) Rizzini, Rodriguésia 46: 72. 1978. Type: BRAZIL. MINAS GERAIS: M. Claussen 252 (lectotype, designated by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: G - barcode 0020029!; isolectotypes: P!, K, S).

Phoradendron emarginatum var. minor Eichler in Martius, Fl. bras. 5(2): 119. 1868. Phoradendron minor (Eichler) Trel., Phoradendron 117, Fig. 172a. 1916. Type: BRAZIL. PIAUÍ: “super arbores campestres inter Olho d’Agoa et Alegre, praedia,” C.F.P. Martius s.n. (holotype: B, destroyed; lectotype, designated by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: M (= Trelease, 1916, Fig. 172a).

Phoradendron tetragonum Ule, Notizbl. Bot. Gart. Berlin-Dahlem 6(59): 290. 1915. Type: BRAZIL. AMAZONAS: near São Marcos, Rio Branco, on Leguminosae, I.1909, E.H.G. Ule 7890 (holotype: B, destroyed (= F Neg 18192); lectotype, hic designatus: K - barcode 00601483!; isolectotypes: IAN, MO, L, UC).

Phoradendron macedonis Rizzini, Rodriguésia 30/31: 163, Fig. 29-1. 1956, in clavi, nomen nudum. Type: BRAZIL. MINAS GERAIS: Ituiutaba, São Vicente, 26.VI.1949, A. Macedo 1893 (RB).

Phoradendron cuspidatum Rizzini, Rodriguésia 30/31: 182. 1956. Type: BRAZIL. SÃO PAULO: Campinas, Riqueza Farm, J. Egidio, 6.XI.1938, J. Gregorio s.n. (lectotype, hic designatus: IAC 002838; isotypes: IAC, IAN!, SP!), syn. nov. Figs. 1l; 3d; 12

Phoradendron mucronatum is one of the most common species of mistletoes in the Brazilian flora. This species is recognized for presenting cataphylls only on the proximal lateral internodes, leaves with palmate veins, generally obovate with an acute, emarginate or retuse apex, biseriate inflorescences with three flowers per fertile bract, and an entirely tuberculate pericarp. It may be confused with P. argentinum, which has narrower leaves and only the apical, tuberculate, pericarp, or with P. craspedophyllum, which has fertile cataphylls. (See comments under D. elliptica).

Kuijt (1994)Kuijt J (1994) Typifcation of the names of new world mistletoe taxa (Loranthaceae and Viscaceae) described by Martius and Eichler. Taxon 43: 187-199. correctly designated lectotypes for P. cearense and for P. cearense var. minor at P herbarium. However, we have found duplicates of these two collections in this herbarium. Hence, we here carry out a second-step lectotypification, selecting one of the duplicates for each name, in accordance with Art. 9.17 of the Code (Turland et al. 2018Turland NJ, Wiersema JH, Barrie FR, Greuter W, Hawksworth DL, Herendeen PS, Knapp S, Kusber WH, Li DZ, Marhold K, May TW, McNeill J, Monro AM, Prado J, Price MJ & Smith GF (2018) International code of nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Koeltz Botanical Books, Glashütten. 254p.). For P. tetragonum, which holotype at B was destroyed in the second World War, we here designate a lectotype.

The name Phoradendron macedonis is cited only in the identification key and illustration by Rizzini (1956, Fig. 29-1). Therefore, it is not a validly published name. At RB herbarium, we have located two duplicates, carrying the author’s handwriting and similar to the illustration of P. mucronatum.

Phoradendron cuspidatum was considered a synonym of P. quadrangulare by Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.. Analyzing the type specimen, we conclude that it represents P. mucronatum and, therefore, we propose it as a new synonym. Since two syntypes of Rizzini’s name were located at IAC, it was necessary to choose a lectotype.

Material selected for illustration (Fig. 12): Sobradinho, DF-205, inside a property, on Aspidosperma subincanum Mart., 2.IV.2002, C.S. Caires & V.C. Mendes 96 (UB).

Iconography: Eichler (1868, Fig. 38-1), Kuijt (2003, Fig. 191), Caires & Proença (2005, Fig. 2e), Dettke & Waechter (2013, Figs. 114-117), Rigon & Cervi (2013, Figs. 6f-j), Dettke & Waechter (2014, Figs. 6e-h), Caires (2017, Figs. 2m-p).

It is distributed in North America - Mexico, Central America (including Caribbean) and South America - Argentina, Bolivia, Brazil (AM, BA, CE, DF, ES, MA, MG, MS, MT, PA, PB, PE, PR, RJ, SP, TO), Colombia, Guyana, Paraguay and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Waechter 2014Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.; Dettke & Caires 2020Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... ).

2.26. Phoradendron nigricans Rizzini, Rodriguésia 30/31: 187. 1956. Type: BRAZIL. RIO DE JANEIRO: Morro Queimado, 13.V.1945, P. Occhioni 212 (holotype: RB (three sheets - barcodes 00540601!, 00545656! and 00545655!); isotype: RFA).

Phoradendron irwinii Rizzini, Arq. Jard. Bot. Rio de Janeiro 24: 33. 1980. Type: BRAZIL. MINAS GERAIS: Serra do Espinhaço, 4 km N of São João da Chapada, 23.III.1970, H.S. Irwin et al. 28178 (holotype: RB (two sheets - barcodes 00540587! and 00545648!); isotypes: F, IAN!, NY!, UB!, US!).

Phoradendron nigricans can be recognized by its monopodial/percurrent branches with cataphylls only on the proximal lateral internodes, rhombic and flattened young branches, leaves with palmate veins, elliptical in shape and that in sicco are characteristically black, and biseriate inflorescences with three flowers per fertile bracts, all pistillate.

The type collection of P. nigricans, Occhioni 212, is sometimes misinterpreted as having been collected in Distrito Federal, the state where the capital of Brazil (Brasília) is located today; but this sample was actually collected in the state of Rio de Janeiro, capital of Brazil until 1960. Therefore, the label on the sheet should include the information “Distrito Federal” or “ex-Distrito Federal”.

Iconography: Kuijt (2003, Fig. 198).

It is distributed in South America - Brazil (BA, DF, ES, MG, PB, PE, RJ) (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.).

2.27. Phoradendron northropiae Urb., Mem. Torrey Bot. Club 12: 33, Fig. 4. 1902. Type: BAHAMAS. ANDROS ISLAND: Conch Sound, J.I. Northrop & A.R. Northrop 551 (holotype: B, destroyed (= Trelease, 1916, Fig. 189a); lectotype, designated by Gillis (1976)Gillis WT (1976) The mistletoes of the Bahamas. Phytologia 33: 361-368.: F! (= F Neg 63689); isolectotypes: GH!, K!, NY).

Phoradendron excisum Rizzini, Rodriguésia 30/31: 192. 1956. Type: BRAZIL. MARANHÃO: Mongunça Island, VII.1914, A. Lisboa 20 (holotype: RB (two sheets - barcodes 00545646! and 00540581!); isotypes: F!, MBM!, MO, NY!).

Phoradendron northropiae is distinguished by the presence of cataphylls on all the internodes, a combination of monopodial/percurrent and sympodial/dichotomous branches, obovate leaves with rounded or retuse apex, and biseriate inflorescences. This species resembles P. racemosum and, in fact, Gillies (1976), when studying the Bahamas flora, considered this species a synonym of P. racemosum. We agree with Kuijt’s (2003) circumscription, which considers these two species as distinct. In P. racemosum, the dichotomies are much more frequent and present cataphylls only on the lateral branches. Thus, the leaves are larger than in P. nothropiae, and generally elliptical with an acute apex. Phoradendron northropiae may be confused with P. inaequidentatum, which presents fertile cataphylls (vs. absent in P. northropiae), generally four pairs of cataphylls on the internodes (vs. one pair), leaves with veins palmate (vs. veins pinnate) and inflorescences entirely biseriate (vs. bi-uniseriate).

Iconography: Urban (1902; Fig. 4), Rizzini (1956, Fig. 10-2), Rizzini (1982, Fig. 66), Kuijt (2003, Fig. 201).

It is distributed in Central America (including Caribbean), and South America - Bolivia, Brazil (GO, MA, MT, PA), Guyana, Peru and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Caires 2020Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... ).

2.28. Phoradendron obtusissimum (Miq.) Eichler in Martius, Fl. bras. 5(2): 134m. 1868. Viscum obtusissimum Miq., Linnaea 18: 602. 1844. Type: SURINAME. “Para Superiore,” IX.1844, H.C. Focke 1019 (holotype: U!; isotype: K!).

Phoradendron acinacifolium Eichler in Martius, Fl. bras. 5(2): 117, Fig. 37-1. 1868. Type: BRAZIL. RIO DE JANEIRO: M. Gaudichaud 574 (lectotype, designated by Kuijt (1994)Kuijt J (1994) Typifcation of the names of new world mistletoe taxa (Loranthaceae and Viscaceae) described by Martius and Eichler. Taxon 43: 187-199.: P, second step, hic designatus: P - barcode 00756338!; isolectotypes: G!, P!).

Phoradendron craspedophylloides Trel., Phoradendron 92, Fig 124b. 1916. Type: BRAZIL. ESPÍRITO SANTO: Vittoria, F. Sello s.n. (holotype: B, destroyed (= Trelease, 1916, Fig. 124b). BRAZIL. PIAUÍ: near Areias, on Inga, G. Gardner 2622 (neotype, first step designated by Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.: K, second step, hic designatus: K - barcode 0016719!; isoneotypes: BM, G!, K!, P!, US!).

Phoradendron reductum Trel., Phoradendron 93, Fig. 127b. 1916. Type: PARAGUAY: Süd Paraguay, IX.1982, O. Kuntze 151 [as Kuntze 15 in Trelease, 1916] ((lectotype, designated by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: B, destroyed (= Trelease, 1916, Fig. 127b); lectotype, designated by Dettke & Waechter (2014)Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.: NY - barcode 00285364!; isolectotype: US!). Fig. 3e

This species is characterized by a combination of branches with growth pattern monopodial/percurrent and sympodial/dichotomous, cataphylls presents only on the proximal lateral internodes, leaves generally obovate, inflorescences biseriate and a remarkable ellipsoidal fruit. Phoradendron obtusissimum is similar to P. singulare in growth pattern, leaves and fruit morphology, but the latter is distinct by the presence of fertile cataphylls and uniseriate inflorescences. (See comments under P. pellucidulum).

Kuijt (1994)Kuijt J (1994) Typifcation of the names of new world mistletoe taxa (Loranthaceae and Viscaceae) described by Martius and Eichler. Taxon 43: 187-199. designated a collection at P herbarium as lectotype of P. acinacifolium, but we have located two duplicates in this herbarium. Hence, we have selected one of them for a second-step lectotypification (Turland et al. 2018Turland NJ, Wiersema JH, Barrie FR, Greuter W, Hawksworth DL, Herendeen PS, Knapp S, Kusber WH, Li DZ, Marhold K, May TW, McNeill J, Monro AM, Prado J, Price MJ & Smith GF (2018) International code of nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Koeltz Botanical Books, Glashütten. 254p., Art. 9.17). The same process was followed for the neotype designated by Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643. for of P. craspedophylloides, housed at K herbarium, and here we designated a second-step neotypification.

Iconography: Eichler (1868, Fig. 37-1), Eliçabe (1973, Fig. 5), Rizzini (1982, Fig. 52), Kuijt (2003, Fig. 203), Kuijt (2005, Fig. 406), Dettke & Waechter (2013, Figs. 118-123), Rigon & Cervi (2013, Figs. 6c-e), Dettke & Waechter (2014, Figs. 6i-m), Caires (2017, Figs. 3a-e).

It is distributed in Central America and South America - Argentina, Bolivia, Brazil (AC, AL, AM, BA, CE, DF, ES, GO, MA, MG, MS, MT, PA, PB, PE, PI, PR, RJ, RO, RR, SE, SP, TO), Colombia, Ecuador, Guyana, Paraguay, Peru, Suriname and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Waechter 2014Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.; Dettke & Caires 2020Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... ).

2.29. Phoradendron oliveirae Kuijt, Syst. Bot. Monogr. 66: 329, Fig. 206. 2003. Type: BRAZIL. PARÁ: Maracanã, forest of “IDESP” Training Center, 13.XII.1977, E. Oliveira 6742 (holotype: RB (two sheets - barcodes 00540603! and 00545658!); isotype: MBM!).

Phoradendron oliveirae is a curious species, known only for its very fragmented type collection. It is very similar to P. piperoides as both species present monopodial/percurrent branches, leaves with pinnate veins, similar position and shape of the cataphylls, and biseriate inflorescences. However, P. oliveirae differs in the presence of fertile cataphylls, which have never been reported to P. piperoides. This species also resembles P. hexastichum and P. juruanum, which bear triseriate inflorescences (vs. biseriate in P. oliveirae). We have decided to keep, for the time being, P. oliveirae as a distinct species, in the absence of sufficient data to make other decision. The paratype (M. Silva 1053 (RB - barcodes 206875 and 206088) clearly does not belong to the same taxon, because it doesn’t bear any fertile cataphylls, and presents dichotomous branches; therefore, it is better placed in P. racemosum.

Iconography: Kuijt (2003, Fig. 206).

It is distributed in South America - Brazil (PA) (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.).

2.30. Phoradendron pellucidulum Eichler in Martius, Fl. bras. 5(2): 112. 1868. Type: BRAZIL. AMAZONAS: San Carlos usque Rio Negro, R. Spruce 3480 (lectotype, designated by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: W, destroyed (= Trelease, 1916, fig. 121); lectotype, designated by Kuijt (1994)Kuijt J (1994) Typifcation of the names of new world mistletoe taxa (Loranthaceae and Viscaceae) described by Martius and Eichler. Taxon 43: 187-199.: P - barcode 002441991!; isolectotypes: K!(2x)).

The species is recognized by its erect, monopodial/percurrent branches with cataphylls only on the proximal lateral internodes, short internodes, usually ellipsoidal and flattened on the nodes, leaves with glistening surface, young leaves translucent and inflorescences triseriate. It may be confused with P. obtusissimum, which bears biseriate inflorescences, and a remarkable ellipsoidal fruit (vs. globose).

Iconography: Dettke & Waechter (2014, Figs. 7a-f), Kuijt (2003, Fig. 217), Kuijt (2005, Fig. 401).

It is distributed in South America - Brazil (AC, AM, AP, BA, DF, MG, MT, PA, PE, RR, SC, SE), Colombia, Guyana, Paraguay, Peru and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Waechter 2014Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.; Dettke & Caires 2020Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... ).

2.31. Phoradendron perrottetii (DC.) Eichler in Martius, Fl. bras. 5(2): 112. 1868. Viscum perrottetii DC., Prodr. 4: 280. 1830. Type: FRENCH GUIANA. G.S. Perrottet 228 (holotype: G-DC - barcode 006223! (= Trelease, 1916, Fig. 119a); isotype: G-DC!).

Phoradendron perrottetii var. parvifolium Eichler in Martius, Fl. bras. 5(2): 113. 1868. Type: BRAZIL. PIAUÍ: G. Gardner 3763 (lectotype, designated by Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.: P; isolectotype: BM!).

Phoradendron perrottetii var. curvifolium Rizzini, Rev. Brasil. Biol. 51: 458. 1991. Type: BRAZIL. MINAS GERAIS: Águas Santas, A. Castellanos 25420 (holotype: RB - barcode 001208764! (ex-GUA); isotype: NY!). Figs. 1n; 3f-g; 13

This species is easily recognized by the monopodial/percurrent branches with cataphylls only on the proximal lateral internodes; large, oblong or falcate, leaves; and long triseriate inflorescences. It can be confused with P. bathyoryctum and P. macrarthrum, but both these species possess biseriate inflorescences. (See comments under P. congestum and P. ensifolium).

Phoradendron harmsianum was considered a synonym of P. perrottetii by Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643., but presents biseriate inflorescences, and therefore, it is placed in P. macrarthrum in this taxonomic treatment.

Material selected for illustration (Fig. 13): Brasília, Núcleo Bandeirante, margin lagoon, very anthropic, on Tapirira guianensis Aubl., 14.III.2002, C.S. Caires & Gonçalves 59 (UB); Planaltina, Córrego Vargem de Trás, near the old Cerâmica Reunidas Dom Bosco, on Tapirira guianensis Aubl., 10.VIII.2001, C.S. Caires & V.C. Mendes 5 (UB).

Iconography: Rizzini (1995, Fig. 20), Kuijt (2003, Fig. 221), Kuijt (2005, Fig. 404), Dettke & Waechter (2014, Figs. 7g-k).

It is distributed in South America - Argentina, Bolivia, Brazil (AM, BA, DF, GO, MA, MG, MS, MT, PA, PB, PE, PR, RR, SC, SE, SP, TO), Colombia, French Guiana, Guyana, Peru, Surinam and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Waechter 2014Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.; Dettke & Caires 2020Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... ).

2.32. Phoradendron piperoides (Kunth) Trel., Phoradendron 145, Figs. 217-222. 1916. Loranthus piperoides Kunth, Nov. gen. sp. 3: 443. 1818. Viscum piperoides (Kunth) DC., Prodr. 4: 281. 1830. Type: COLOMBIA. CAUCA: Popayan, 1871, F.W.H.A. Humboldt & A.J.A Bonpland s.n. (holótipo: P-HBK! (= Trelease, 1916, Fig. 217a).

Phoradendron schottii (Pohl ex DC.) A. Gray, U.S. Explor. Exped. [Bot., Phanerogam.] 15: 742. 1854. Viscum schottii Pohl ex DC., Prodr. 4: 281. 1830. Type: BRAZIL. RIO DE JANEIRO: Organ Mountains, 1828, H.W. Schott s.n. (holotype: G-DC).

Phoradendron piperoides f. compositum Trel., Phoradendron 148. 1916, [composita]. Type: BRAZIL. RIO DE JANEIRO: Cabo Frio, in restinga, E.H.G. Ule 4800 (holotype: B, destroyed). Fig. 3h-i

Phoradendron piperoides is one of the most common Brazilian mistletoes, easily recognized by its monopodial/percurrent branches, sterile cataphylls present in all internodes, veins pinnate and generally elliptical leaves, and biseriate inflorescences. In herbarium collections, it is often confused with P. crassifolium, which has fertile cataphylls and leaves with palmate veins. (See comments under P. chrysocladon, P. juruanum, P. laxiflorum and P. oliveirae).

Iconography: Eichler (1868, Fig. 41), Rizzini (1956, Fig. 14), Eliçabe (1973, Fig. 8), Kuijt (2003, Fig. 223), Caires & Proença (2005, Fig. 1i), Kuijt (2005, Fig. 407), Dettke & Waechter (2013, Figs. 124-129), Rigon & Cervi (2013, Figs. 6a-b), Dettke & Waechter (2014, Figs. 8a-f), Caires (2017, Figs. 3f-i).

It is distributed in North America - Mexico, Central America (including Caribbean) and South America - Argentina, Bolivia, Brazil (all states), Colombia, Ecuador, French Guiana, Paraguay, Peru, Surinam and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Caires 2020Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... ).

2.33. Phoradendron platycaulon Eichler in Martius, Fl. bras. 5(2): 108. Fig. 33. 1868. Type: BRAZIL. PARÁ: Santarém, R. Spruce 228 (lectotype, designated by Trelease, 1916: M - barcode 00174058! (= Trelease, 1916, Fig. 227b)).

This species is the only one within the genus bearing the expanded leaves, which are characteristic of Phoradendron, with flattened stems. Also, P. platycaulon has a pair of cataphylls in all the internodes, and biseriate inflorescences with three flowers per fertile bract (where the apical one is staminate). (See comments under P. tunaeforme).

Iconography: Eichler (1868, Fig. 33), Kuijt (2003, Fig. 226), Kuijt (2005, Fig. 405).

It is distributed in South America - Bolivia, Brazil (AM, BA, MG, MT, PA, RO, RR), Colombia, Peru and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.).

2.34. Phoradendron poeppigii (Tiegh.) Kuijt, Acta Bot. Neerl. 10: 199. 1961. Dendropthora poeppigii Tiegh., Bull. Soc. Bot. France 43: 182. 1896. Type: BRAZIL. “Brasilia borealis”, E.F. Poeppig s.n. (holotype: P - barcode 005339370!).

Phoradendron poeppigii is an aphyllous species of the Amazon rainforest, characterized by terete and pendulous stems with long internodes (usually longer than 4 cm); biseriate inflorescences, with only 3 flowers per fertile bract, and the apical flower staminate.

Iconography: Kuijt (2003, Fig. 227).

It is distributed in South America - Brazil (AM, PA) and Guyana (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.).

2.35. Phoradendron pteroneuron Eichler in Martius, Fl. bras. 5(2): 127. 1868. Type: BRAZIL. Rio de Janeiro: “crescit circa Rio de Janeiro”, A.F.M. Glaziou 1462 (lectotype, first step designated by Kuijt (1994)Kuijt J (1994) Typifcation of the names of new world mistletoe taxa (Loranthaceae and Viscaceae) described by Martius and Eichler. Taxon 43: 187-199.: BR, second step, hic designatus: BR - barcode 0000005629589!; isolectotypes: BR!(2x), P!(2x)).

Phoradendron rugulosum Urb., Bot. Jahrb. Syst. 23, Beih., 57: 13. 1897. Phoradendron warmingii var. rugulosum (Urb.) Rizzini, Rev. Brasil. Biol. 51: 459. 1991. Type: BRAZIL. RIO DE JANEIRO: “Aux Orgues”, Organ Mountains, A.F.M. Glaziou 4012 (holotype: B, destroyed; lectotype, designated by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: C - barcode 10014049! (= Trelease, 1916, Fig. 198b); isolectotypes: C!, K!, P!(2x)). Fig. 3j

Phoradendron pteroneuron can be recognized by the monopodial/percurrent growth pattern, with cataphylls only on the proximal lateral branches, leaves with pinnate veins and a conspicuous central vein, and biseriate inflorescences. This species may be confused with D. warmingii, P. northropiae and P. racemosum. It differs from D. warmingii by presenting four, or more, pairs of sterile bracts in the inflorescences (vs. one, or two, in D. warmingii) and by the closed petals in the fruit (vs. opened). It differs from P. northropiae and P. racemosum, by the rarity of dichotomous branches (vs. frequent dichotomies in P. northropiae and P. racemosum). It also differs from P. northropiae by the presence of cataphylls only on proximal lateral internodes (vs. presents in all internodes in P. northropiae).

Kuijt (1994)Kuijt J (1994) Typifcation of the names of new world mistletoe taxa (Loranthaceae and Viscaceae) described by Martius and Eichler. Taxon 43: 187-199. designated the collection from BR herbarium as lectotype of P. pteroneuron. However, we have located three duplicates at BR, one of which we select for a second-step lectotypification, according to the Code (Turland et al. 2018Turland NJ, Wiersema JH, Barrie FR, Greuter W, Hawksworth DL, Herendeen PS, Knapp S, Kusber WH, Li DZ, Marhold K, May TW, McNeill J, Monro AM, Prado J, Price MJ & Smith GF (2018) International code of nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Koeltz Botanical Books, Glashütten. 254p., Art. 9.17).

Iconography: Rizzini (1956, Fig. 16-1), Kuijt (2003, Fig. 231).

It is distributed in South America - Bolivia, Brazil (AM, AL, AP, BA, CE, ES, MG, MS, MT, PA, PE, RJ, SP), Colombia, Surinam and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Caires 2020Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... ).

2.36. Phoradendron quadrangulare (Kunth) Griseb., Fl. Brit. W. I. 711. 1864. Loranthus quadrangularis Kunth, Nov. Gen. Sp. [H.B.K.] 3: 444. 1818. Viscum quadrangulare (Kunth) DC., Prodr. 4: 283. 1830. Type: COLOMBIA. CUNDINAMARCA: near to Pandi and Fusagasuga, on Guazuma, IX.1795, F.W.H.A. Humboldt & A.J.A Bonpland s.n. (holotype: P-HBK - barcode 00322225! (= Trelease, 1916, Fig. 155a).

Phoradendron affine (Pohl ex DC.) Engl. & Krause, Nat. Pflanzenfam. 16b: 191. 1935. Viscum affine Pohl ex DC., Prodr. 4: 281. 1830. Type: BRAZIL. MINAS GERAIS: São João Baptista, J.E. Pohl 544 (holotype: G-DC! (= Trelease, 1916, Fig. 165a); isotype: ILL!, W, destroyed (= Trelease, 1916, Fig. 165b)).

Phoradendron microphyllum (Pohl ex DC.) Trel., Phoradendron 113, Fig. 164a. 1916. Viscum microphyllum Pohl ex DC., Prodr. 4: 283. 1830. Phoradendron rubrum var. microphyllum (Pohl ex DC.) Eichler in Martius, Fl. bras. 5(2): 120. 1868. Type: BRAZIL. H.W. Schott s.n. (holotype: G-DC), syn. nov.

Phoradendron liga (Gillies ex Hook. & Arn.) Eichler in Martius, Fl. bras. 5(2): 134m. 1868. Viscum liga Gillies ex Hook. & Arn., Bot. Misc. 3: 355. 1833. Type: ARGENTINA. SAN JUAN: near to Los Cerrillos de San Juan, J. Gillies s.n. (holotype: K – barcode 00601434! = Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p., Fig. 179); isotype: E!).

Phoradendron rubrum var. longispicum Eichler in Martius, Fl. bras. 5(2): 121. 1868, [longispica]. Phoradendron piauhyanum Trel., Phoradendron 110. 1916. Type: BRAZIL. PIAUÍ: G. Gardner 2617 (lectotype, designated by Trelease, 1916: W, destroyed (= Trelease, 1916, Fig. 157a); lectotype, first step designated by Kuijt (1994)Kuijt J (1994) Typifcation of the names of new world mistletoe taxa (Loranthaceae and Viscaceae) described by Martius and Eichler. Taxon 43: 187-199.: P, second step, hic designatus: P - barcode 005447871!; isolectotypes: BM, G!(4x), K!(2x), US!).

Phoradendron wiesnerianum Trel., Phoradendron 109, Fig. 156. 1916. Type: BRAZIL, CEARÁ: Villa do Crato, G. Gardner 1674 (holotype: W, destroyed (= Trelease, 1916, Fig. 156; lectotype, hic designatus: BM - barcode 0049235!; isolectotypes: BM!, G!, K, NY!, P!(3x)).

Phoradendron stenophyllum Trel., Phoradendron 81, Fig. 106b. 1916, non Viscum stenophyllum Sprengel, 1824 [“1825”]. Type: BRAZIL. BAHIA: Machado Portello, 19–23.VI.1915, J.N. Rose & P.G. Russell 19908 (holotype: US - barcode 00731090! (= Trelease, 1916, Fig. 106b); isotype: NY!).

Phoradendron martianum Trel., Phoradendron 114, Fig. 166. 1916. Type: BRAZIL. ALAGOAS: G. Gardner 1321 (holotype: G - barcode 006233! (= Trelease, 1916, Fig. 166a); isotypes: BM!(2x), G!, ILL!, K!(3x), P!(2x)).

Phoradendron lineolatum Rizzini, Rodriguésia 30/31: 184. 1956. Type: BRAZIL, CEARÁ: Dias da Rocha 118 (holotype: RB - barcode 00540588!).

Phoradendron fusco-erubescens Rizzini, Arq. Jard. Bot. Rio de Janeiro 24: 32. 1980. Type: BRAZIL. GOIÁS: Chapada dos Veadeiros, 20 km N of Alto do Paraíso, on Byrsonima, 20.III.1971, H.S. Irwin et al. 32748 (holotype: RB - barcode 00540583!; isotypes: CAS, MO, NY!, UC(ex-LEA)), syn. nov.

Phoradendron saxicola Rizzini, Arq. Jard. Bot. Rio de Janeiro 24: 37. 1980. Type: BRAZIL. MATO GROSSO: Chapada dos Guimarães, road Buriti to Cuiabá, on Byrsonima, 22.X.1973, G.T. Prance et al. 19292 (holotype: RB - barcode 00540611!; isotypes: INPA!, NY!, R), syn. nov. Figs. 3k; 14

Phoradendron quadrangulare is one of the most common species of mistletoes in the Brazilian flora. This species, according to our observations, has predominance of monopodial/percurrent branches, cataphylls only on the proximal lateral internodes, young branches generally rhombic or quadrangular, elliptical or obovate leaves, biseriate inflorescences, and globose fruits with vibrant orange or reddish color. (See comments under P. diminutivum).

Dettke & Waechter (2014)Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985., through the observation of several populations in Brazil, and adjacent countries (Argentina, Paraguay and Uruguay), included P. affine and P. liga under synonymy of P. quadrangulare. In the present study, we include three additional names: P. microphyllum, which was recognized as a separate species by Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643., and P. fusco-erubescens and P. saxicola, considered synonyms of P. trinervium (Lam.) Griseb. (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.). P. quadrangulare is a widely distributed species, morphologically variable across its distribution range, presenting delicate to more robust plants, depending on the habitat. However, the characteristics listed above are constant in all distributional records. It likely constitutes the same taxon, or a species complex with P. trinervium and P. rubrum (L.) Griseb., which occurs in Central (including Caribbean) and northern South America, but additional studies at population level are necessary to confirm this statement. Phoradendron cuspidatum, previously considered a synonym of P. affine (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.), is here placed under synonymy of P. mucronatum.

Material selected for illustration (Fig. 14): Brasília, Asa Norte, Estação Experimental de Biologia-UnB, on Anadenanthera, 9.XI.2001, C.S. Caires & V.C. Mendes 26 (UB); Chácara Sossêgo, DF-238 road to Unaí, on Anadenanthera, 21.I.2002, C.S. Caires et al. 40 (UB). São Sebastião, DF-251 road to Unaí/PADDF, over bridge of São Bartolomeu River, on Anadenanthera, 16.IV.2002, C.S. Caires & V.C. Mendes 144 (UB).

Iconography: Eichler (1868, Fig. 38-2), Rizzini (1956, Fig. 29-3), Eliçabe (1973, Fig. 9), Rizzini (1980, Fig. 10), Rizzini (1995, Fig. 19), Kuijt (2003, Figs. 16, 161, 185, 234), Kuijt (2005, Fig. 399), Dettke & Waechter (2013, Figs. 130-136), Rigon & Cervi (2013, Figs. 7e-f), Dettke & Waechter (2014, Figs. 8g-l), Caires (2017, Figs. 1e-g, 3j-m, 4a-b).

It is distributed in North America - Mexico, Central America (including Caribbean) and South America - Argentina, Bolivia, Brazil (all states), Colombia, Ecuador, Guyana, Paraguay and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Waechter 2014Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.; Dettke & Caires 2020Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... ).

2.37. Phoradendron racemosum (Aubl.) Krug & Urb., Bot. Jahrb. Syst. 24: 46. 1897. Viscum racemosum Aublet, Hist. pl. Guiane 2: 895. 1775. Viscum flavens Swartz, Prodr. 32. 1788, nom. superfl. Phoradendron flavens (Swartz) Griseb., Fl. Brit. W. I. 313. 1860. Type: (lectotype, designated by Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.: Plate 258, Fig. 4, in C. Plumier (ed. J. Burman), Plantarum americanarum, 1760).

Phoradendron pennivenium Eichler in Martius, Fl. bras. 5(2): 128, Fig. 42. 1868 (excl. syn. De Candolle), non Viscum pennivenium DC. (= Oryctanthus florulentus (Rich.) Tiegh.). Type: GUIANA, 1821, G.S. Perrottet s.n. (lectotype, hic designatus: P - barcode 05447927!). Fig. 3l

This species is distinguishable by the sympodial/dichotomous growth pattern of branches, cataphylls only on the proximal lateral branches, leaves with pinnate veins and rounded apex, and biseriate inflorescences. Phoradendron racemosum can be mistaken with P. hexastichum, which has monopodial percurrent branches and triseriate inflorescences, or with P. northropiae that has cataphylls in all internodes and smaller leaves. (See comments at D. fendleriana and P. pteroneuron).

The interpretation of Phoradendron pennivenium has been controversial, because of a confusion between Perrottet’s collections. It is clear that De Candolle (1830)De Candolle AP (1830) Loranthaceae. Prodromus Systematis Naturalis Regni Vegetabilis 4: 277-320. described Viscum pennivenium based on a collection of Oryctanthus florulentus (Rich.) Tiegh. (isotypes seen: GUIANA, Perrottet s.n. P - barcodes 05455466! and 05455466!), considering the morphological characteristics described in the protologue (De Candolle 1830De Candolle AP (1830) Loranthaceae. Prodromus Systematis Naturalis Regni Vegetabilis 4: 277-320.: 282-283): “ramulis tetragonis subteretibus”, “spicis axillaribus solitariis” and “baccis oblongis”. However, Eichler (1868)Eichler AW (1868) Loranthaceae. In: Martius CFP (ed.) Flora brasiliensis. Fleicher, Leipzig. Vol. 5, pars 2, pp. 1-136, t.1-44. used another unnumbered collection of Perrottet from Guiana, of the same year (1821), to describe P. pennivenium in Flora brasiliensis. This specimen does not have the characteristics described by De Candolle, and represents P. racemosum. After excluding De Candolle’s synonym, Eichler’s names are recognized and attributed to him, according to the Code (Turland et al. 2018Turland NJ, Wiersema JH, Barrie FR, Greuter W, Hawksworth DL, Herendeen PS, Knapp S, Kusber WH, Li DZ, Marhold K, May TW, McNeill J, Monro AM, Prado J, Price MJ & Smith GF (2018) International code of nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Koeltz Botanical Books, Glashütten. 254p., Art. 48.1). We, here, select a lectotype from P herbarium, among the syntypes listed by Eichler (1868)Eichler AW (1868) Loranthaceae. In: Martius CFP (ed.) Flora brasiliensis. Fleicher, Leipzig. Vol. 5, pars 2, pp. 1-136, t.1-44. for P. pennivenium.

Iconography: Eichler (1868, Fig. 42), Kuijt (2003, Fig. 237), Kuijt (2005, Fig. 409).

It is distributed in Central America (including Caribbean) and South America - Brazil (AC, AM, AP, BA, CE, DF, GO, MG, MT, PA, RR, TO), Colombia, French Guiana, Surinam and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.).

2.38. Phoradendron singulare Kuijt, Syst. Bot. Monogr. 66: 424, Fig. 265. 2003. Type: COLOMBIA. CAQUETA: Sierra de Chiribiquete, Campamento Norte, 01º07’N, 72º50’W, on Clusia, S. Castroviejo et al. 12057 (holotype: HUA - barcode 000217!).

Phoradendron singulare is a conspicuous Amazon species that is characterized by monopodial/percurrent branches; presence of fertile cataphylls, located on the upper or middle portion of the internodes; and uniseriate inflorescences in both axillary and terminal positions. It resembles P. obtusissimum due to the similarity of the obovate leaves, and ellipsoid fruits with opening petals, but is distinguished by the fertile cataphylls (vs. sterile) and uniseriate inflorescences (vs. biseriate).

Iconography: Kuijt (2003, Fig. 265).

It is distributed in South America - Brazil (AM) and Colombia (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.).

2.39. Phoradendron strongyloclados Eichler in Martius, Fl. bras. 5(2): 109. 1868. Type: BRAZIL. PERNAMBUCO: Itamaracá, G. Gardner 1029 (holotype: B, destroyed; lectotype, designated by Trelease (1916)Trelease W (1916) The genus Phoradendron. A monographic revision. University of Illinois, Urbana. 224p.: GH - barcode 0035780! (= Trelease, 1916, Fig. 224a); isolectotypes: BM!(2x), E!, K!(2x), P!(3x), NY).

Phoradendron gardnerianum Urb., Bot. Jahrb. Syst. 23, Beibl. 57: 11. 1897. Type: BRAZIL. PIAUÍ: Serra da Batalha, G. Gardner 2885 (holotype: B, destroyed (= Trelease, 1916, Fig. 223a); lectotype, hic designatus: BM - barcode 00993575!; isolectotype: K!).

Phoradendron caesalpiniae Ule, Bot. Jahrb. Syst. 42: 200. 1908. Type: BRAZIL. BAHIA: Caldeirão, X.1906, on Caesalpiniaceae, E.H.G. Ule 7243 (holotype: B, destroyed (= Trelease, 1916, Fig. 225b); lectotype, hic designatus: K - barcode 0016535!; isolectotypes: G!, HBG!). Figs. 3m; 15

Phoradendron strongyloclados is characterized by a mixture of monopodial/percurrent and sympodial/dichotomous branches, one pair of cataphylls on all the internodes, small leaves, generally ovate or elliptical less than 1.5 cm wide, biseriate inflorescences with three flowers per fertile bracts,the apical one staminate, and ovate fruits with opening petals.

The holotypes of P. gardnerianum and P. caesalpiniae at B herbarium were destroyed during the second World War, and so, we designate here the lectotypes for both names.

Material selected for illustration (Fig. 15): Brasília, basin of São Bartolomeu River, V.1981, E.P. Heringer et al. 6991 (IBGE); Jardim Botânico de Brasília, VIII. 1990, P.E.A.M. Oliveira 1563 (HEPH); 7 km W of Brasília, near Sobradinho, 30.IX.1965, H.S. Irwin et al. 8832 (UB).

Iconography: Rizzini (1982, Fig. 74), Kuijt (2003, Fig. 273), Caires & Proença (2005, Fig. 2h), Kuijt (2005, Fig. 397).

It is distributed in South America - Bolivia, Brazil (AM, AP, BA, DF, GO, MA, MG, MS, MT, PA, PB, PE, PI, RJ, SE, TO), Colombia, French Guiana, Guyana, Surinam and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Caires 2020Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... ).

2.40. Phoradendron tunaeforme (DC.) Eichler in Martius, Fl. bras. 5(2): 108, Fig. 32. 1868. Viscum tunaeforme DC., Prodr. 4: 284. 1830. Type: BRAZIL. Serra de San Feliz, Musquitos, J.E. Pohl s.n. (holotype: G-DC; isotypes: BR!, ILL!, M, P). Fig. 3n-o

Phoradendron tunaeforme is an aphyllous plant, clearly distinguished from the other species of Phoradendron by the flattened and pendulous stems, and by the short inflorescences, with only one or two fertile biseriate segments. It can be confused with P. platycaulon, for the identical stems, but the expanded leaves and larger inflorescences of P. platycaulon clearly separates them.

Iconography: Eichler (1868, Fig. 32), Kuijt (2003, Fig. 293), Caires & Proença (2005, Figs. 1j-k), Caires (2017, Figs. 3n-q, 4c-f).

It is distributed in South America - Brazil (AM, AL, BA, CE, DF, GO, MG, PA, PB, PE, RN, SE, SP) and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Caires 2020Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... ).

2.41. Phoradendron undulatum (Pohl ex DC.) Eichler in Martius, Fl. bras. 5(2): 122, Fig. 39. 1868. Viscum undulatum Pohl ex DC., Prodr. 4: 282. 1830. Type: BRAZIL. MINAS GERAIS: Barbacena, 1828, J.E. Pohl s.n. (holotype: G-DC! (= Trelease, 1916, Fig. 190a); isotype: BR!).

Phoradendron hamatifolium Rizzini in Reitz, Flora Ill. Catarinensis 1: 20. 1968. Type: BRAZIL. SANTA CATARINA: Blumenau, Morro Spitzkopf, 20.III.1952, R. Reitz 4633 (holotype: HBR 4757!; isotypes: RB!, MO).

Phoradendron uleanum Steyerm., Fieldiana Bot. 28(1): 223. 1951. Phoradendron macrophyllum Ule, Notizbl. Bot. Gart. Berlin-Dahlem 6: 291. 1915, nom. illeg., non Phoradendron macrophyllum (Engehnann) Cockerell, 1900, nom. illeg.; non Phoradendron macrophyllum Spreng., 1825. Type: BRAZIL. RORAIMA: E.H.G. Ule 8603 (holotype: B, destroyed (=F Neg 18191); lectotype, designated by Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.: K!; isolectotypes: IAN!, MO). Figs. 3p; 16

Phoradendron undulatum is easily distinguished from other species of Phoradendron by monopodial/percurrent branches, with cataphylls only on the proximal lateral internodes, veins pinnate and long-ovate elliptical leaves, with an acute or acuminate apex, and biseriate inflorescences. In collections, it is often confused with P. crassifolium, which has fertile cataphylls and leaves with palmate veins, or with P. chrysocladon, that has cataphylls in all internodes, leaves with palmate veins and triseriate inflorescences.

Material selected for illustration (Fig. 16): Brasília, Mansão Dois Irmãos, near road to FAL, Styrax ferrugineus Nees. & Mart., 5.II.2002, C.S. Caires & V.C. Mendes 44 (UB); São Bartolomeu River, X.1979, E.P. Heringer 2137 (IBGE). Planaltina, CPAC - reserva cerradão, on Emmotum nitens (Benth.) Miers, 2.IV.2002, C.S. Caires et al. 110 (UB).

Iconography: Eichler (1868, Fig. 39), Kuijt (2003, Figs. 294, 295), Caires & Proença (2005, Fig. 1h), Kuijt (2005, Fig. 411), Dettke & Waechter (2013, Figs. 137-140), Rigon & Cervi (2013, Figs. 7a-b), Dettke & Waechter (2014, Figs. 8m-p).

It is distributed in North America - Mexico, Central America (including Caribbean) and South America - Bolivia, Brazil (AM, BA, DF, ES, GO, MG, MT, PA, PE, PR, RJ, RR, RS, SC, SP), Colombia, Ecuador, Peru and Venezuela (Kuijt 2003Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643.; Dettke & Waechter 2014Dettke GA & Waechter JL (2014) Estudo taxonômico das ervas-de-passarinho da Região Sul do Brasil: II. Viscaceae (Phoradendron). Rodriguésia 65: 955-985.; Dettke & Caires 2020Dettke GA & Caires CS (2020) Santalaceae. In: Flora do Brasil 2020 - under construction. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB215>. Access on 9 April 2020.
http://floradobrasil.jbrj.gov.br/reflora... ).

Excluded names for the Brazilian flora

Phoradendron bilineatum Urb.

This species does not occur in Brazil. The two vouchers listed for Brazil by Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643. are refer to other species: C.A. Cid Ferreira et al. 1494 (NY!, RB!, US!) is Phoradendron macrarthrum and G.T. Prance et al. 4370 (US!) is Phoradendron racemosum.

Phoradendron brittonianum Rusby.

This species does not occur in Brazil. Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643. cited a possible record, based on the collection G. Martinelli et al. 12087 (MO!), which actually represents Phoradendron pteroneuron.

Phoradendron herbert-smithii Trel.

This species does not occur in Brazil. The only collection cited by Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643., B. Maguire & C.K. Maguire 40236 (US!), is actually a pistillate individual of P. coriaceum.

Phoradendron polygynum (H. Karst.) Eichler.

This species does not occur in Brazil. The voucher cited by Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643. for Brazil, B.E. Dahlgreen 888 (F!, NY!) is Phoradendron bathyoryctum.

Phoradendron trinervium (Lam.) Griseb.

In this treatment, we consider that this species does not occur in Brazil. Species documented by Kuijt (2003)Kuijt J (2003) Monograph of Phoradendron. Systematic Botany Monographs 66: 1-643., with Brazilian types (P. fusco-erubescens and P. saxicola), have been proposed here as synonyms of Phoradendron quadrangulare.

Acknowledgements

The authors are grateful to all researchers and herbaria involved in the Reflora project; curators of the visited herbaria; Jhon Steven Murillo Serna, for kindly providing images from the HUA herbarium; and Dra. Rafaela Campostrini Forzza, curator of RB Herbarium, for kindly providing information about types of Rizzini’s names. The authors are also grateful to several colleagues for sending images of specimens: Antonio Sérgio Farias, Isa Lucia de Morais, Jair Faria, Jéssica Lira, Juliano Ricardo Fabricante, Leonardo Jales Leitão, Maria Auxiliadora Milaneze-Gutierre, Maria Rosa Zanatta, Sandro Lucas Xavier Tobias and Sérgio Augusto Bordignon. This is publication 29 in the Parasitic Plants Research Group technical series.

References

Supplementary Material

See supplementary material at <https://doi.org/10.6084/m9.figshare.16879597.v1 >

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