Introduction

Parasitism, one of the most striking biological interactions across the tree of life, has driven numerous evolutionary and adaptive novelties in plants and animals. Nevertheless, parasitism has been more extensively studied in the animal kingdom (Schmid-Hempel, 2009), despite the fact that parasitic plants have developed a vast range of remarkable structural, functional, ecological, and genomic novelties to succeed.

The study of plant-plant parasitism has unveiled unique mechanisms in angiosperms, including: the evolution of structural elaborations between hosts and parasites (Thorogood et al., 2021); the recalcitrant parasite-on-parasite occurrence (Bernhardt, 1984; Krasylenko et al. 2021); the coordinate ecological, biogeochemical and physiological responses between hosts and parasites (Heckard, 1962; Matthies 1997; Amaral & Ceccantini, 2011; Ozturk et al., 2022; Zhang et al., 2023); the loss of autotrophic capabilities and the concomitant plastome reduction in the parasite (Bungard, 2004; Vogel et al., 2018; Arias et al., 2019); the fixation of highly elaborated reproductive and embryological traits to ensure fertilization and seed dispersal (Johri & Bhatnagar, 1972; Lucero et al., 2014; González & Pabón-Mora, 2017a, b; the ecological roles of parasitic angiosperms as sources of pollen, nectar, fruits and seeds to trophic interactions (Ladley & Kelly, 1996; Amico et al., 2007, 2011; González & Pabón-Mora, 2017a, b); the dynamics of genome evolution including horizontal gene transfer (Park et al., 2007; Cusimano & Wicke, 2016; Fan et al., 2016; Arias et al., 2019); the use of host flowering signals to trigger parasite flowering (Shen et al., 2020); the regulation of host-mRNA mediated by parasite microRNAs (Shahid et al., 2018); and the parasitism-driven limitation of speciation (Young et al., 1999; Hardy & Cook, 2012), among others.

Parasitic flowering plants directly or indirectly depend upon the uptake of nutrients, water, and minerals from the host. Two main categories and several subcategories have been recognized, mainly based on the degree of reduction of photosynthetic rates, the dependence of the parasite on the host, and the structural modifications that take place as an outcome of this intricate biological association:

Hemiparasitic plants (Heide-Jørgensen, 2008): Functionally, they remain autotrophic, as their vegetative organs (cotyledons, stems and leaves, and, to a less extent, bracts, bracteoles, sepals and embryos) produce chlorophyll and are therefore capable of photosynthesizing. The following two subcategories are recognized: (a) facultative hemiparasites, which depend upon connections with one or (exceptionally) more hosts, but can also survive without a host. And (b), obligate hemiparasites, which retain photosynthetic processes, but still depend on their host for survival.

Holoparasitic plants (Heide-Jørgensen, 2008): Functionally, this is the most extreme life form of parasitism in Angiosperms. They are totally heterothropic because they lack chlorophyll, and therefore depend entirely on their hosts. The following two subcategories are recognized: (a) holoexoparasites, characterized by the formation of structures than emerge aboveground from the host, which correspond to variously modified stems, leaves or inflorescence axes that are not photosynthetic; and (b) holoendoparasites, which lack typical vegetative structures (roots, stems or leaves) and macroscopic haustoria, and their reproductive structures develop inside the host, representing the most extreme event of morphological, anatomical and physiological specialization known in the plant kingdom.

The structural parasite-host connection in members of both categories is usually mediated by a highly elaborate and efficient haustorial system. This system is formed either belowground, and penetrates the host roots (i.e. the root parasites), or aboveground, and penetrates the host stems (i.e. the stem parasites). Interestingly, these two categories are mutually exclusive, meaning that there is not a single angiosperm species that parasitizes both shoots and roots (Heide-Jørgensen, 2008).

Parasitic angiosperms have evolved independently in twelve orders, ranging from facultative hemiparasites to obligate holoendoparasites (Fig. 1). The taxonomic occurrence of parasitic species on each order varies from exceptional in Lauraceae (Laurales), Ehretiaceae (Boraginales), and Convolvulaceae (Solanales), to frequent in Krameriaceae (Zygophyllales) and Orobanchaceae (Lamiales), to generalized in Apodanthaceae (Cucurbitales), Cytinaceae (Malvales), Mitrastemonaceae (Ericales) and Santalales (APG IV, 2016; Thorogood et al., 2021).

Fig. 1
figure 1

Angiosperm phylogeny (modified from APG IV) and orders having parasitic taxa. Purple arrows point to orders with facultative hemiparasites; green arrows point to orders with obligate hemiparasites; red arrows point to orders with holoparasites; gray arrows point to three orders whose parasitic members are not present in Colombia

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Globally, the contribution of parasitic flowering plants to angiosperm diversity is estimated to be nearly 1% of the total number of angiosperm species (Thorogood et al., 2021). This seemingly low percentage contrasts with the high diversity of parasitism-driven biological processes detected in angiosperms. Given that comparable accounts occur at a countrywide scale, here we aim to assess the diversity of parasitic angiosperms in Colombia, one of the biodiversity hotspots in the planet. We present an updated list of parasitic angiosperms in the Flora of Colombia, and provide maps of distribution (Figs. 2, 6, 8, 9, 11, 12, 14, 15, 16, 20, 21, 23, 24, and 25), photographs for 34 representative species (Figs. 3, 4, 5, 7, 10, 13, 17, 18, 19, and 22), and maps of two plastomes (Figs. 4O and 22J), one of them (that of Pilostyles boyacensis F.González & Pabón-Mora, Apodanthaceae; Fig. 4O) corresponding to one of the most (if not the most) reduced plastomes across flowering plants. We discuss distribution patterns of the parasitic angiosperms in the five natural regions of Colombia, and postulate that the uplift of the three cordilleras has played a positive role on the speciation of Aetanthus, Oryctanthus and Psittacanthus (Loranthaceae), Dendrophthora and Phoradendron (Viscaceae), and Castilleja and Neobartsia (Orobanchaceae). Our account resulted in a total of nine orders, 17 families, 44 genera and 246 species (69 of them endemic to the country), corresponding to approximately 1% of the total number of angiosperm species in Colombia, estimated in nearly 25,000 species (Cuatrecasas, 1958).

Materials and Methods

Natural regions of Colombia

The distribution of families, genera and species here reported is outlined on the five natural regions in Colombia (as defined by Cuatrecasas, 1958). These are:

  • Region I. The Caribbean region, including the Archipelago of San Andrés and Old Providence. It encompasses coastal vegetation, mangrove swamps, grassy or forested dunes, scattered dry forest relicts surrounded by man made grassland, scrub thickets, montane vegetation in the Serranía de Macuira (an isolated mountain massif in northern Guajira), and low- to high-mountain forests to paramo vegetation in the Sierra Nevada de Santa Marta (the largest and highest extra-Andean mountain range in the country, and one of the coastal massifs in the world).

  • Region II. The Pacific region, comprising a corridor of pluvial or rain forest that runs from the Gulf of Uraba in the north, throughout the western lowlands of the country reaching the department of Nariño, in southwestern Colombia. This region includes the Malpelo and the Gorgona islands.

  • Region III. The Andean region, including the slopes flanking the Cauca and the Magdalena valleys, the majority of the Andean forests and páramos in the country, the Serranía del Perijá (a mountain massif shared with Venezuela in northeastern Colombia), and the non-split region of the southern Andes known as the Macizo Colombiano (a mountain massif near the border with Ecuador).

  • Region IV. The Orinoquia or Llanos Orientales region, including natural savannas, man made grassland areas, gallery forests along numerous river banks north of the Guaviare river, and the eastern slopes of the Eastern Cordillera.

  • Region V. The Amazonian region, including lowland forests south of the Guaviare river, the eastern slopes of the Eastern Cordillera, and the Guiana shield tepuis of Colombia.

Maps of distribution are presented for all taxa, except for the widespread genera Psittacanthus (Loranthaceae), Dendrophthora and Phoradendron (Viscaceae), and Castilleja (Orobanchaceae) (Figs. 2, 6, 8, 9, 11, 12, 14, 15, 16, 20, 21, 23, 24, and 25).

Plant material

The present study is based on extensive herbarium searches at the Colombian National Herbarium, National University of Colombia (COL), the herbarium of the University of Antioquia, Medellín (HUA), and the Harvard University Herbaria (A, GH). Additionally, the authors have studied in the field a total of 37 species, most of which are illustrated in Figs. 3, 4, 5, 7, 10, 13, 17, 18, 19, and 22. Representative vouchers for all taxa are provided, including 72 type specimens that were examined for this research.

Results

General account of parasitic angiosperms in Colombia

Nine of the twelve orders with parasitic species recognized in the current system of angiosperm classification (APG IV, 2016; Fig. 1) are represented in the Flora of Colombia. These are the orders Boraginales (family Ehretiaceae), Cucurbitales (family Apodanthaceae), Ericales (family Mitrastemonaceae), Lamiales (family Orobanchaceae), Laurales (family Lauraceae), Malvales (family Cytinaceae), Santalales (families Balanophoraceae, Cervantesiaceae, Loranthaceae, Opiliaceae, Santalaceae, Schoepfiaceae, Strombosiaceae, Viscaceae, and Ximeniaceae), Solanales (family Convolvulaceae), and Zygophyllales (family Krameriaceae). Parasitic members of the orders Piperales (i.e. Hydnora and Prosopanche¸currently members of the Aristolochiaceae), Saxifragales (Cynomoriaceae) and Malpighiales (Rafflesiaceae) are not present in the country´s flora.

A total of 187 obligate hemiparasitic species are present in Colombia. They correspond to two species of Krameriaceae (order Zygophyllales), and all members of Santalales, including three species of Cervantesiaceae, 84 species of Loranthaceae, three species of Opiliaceae, three species of Santalaceae, two species of Schoepfiaceae, one species of Strombosiaceae, 88 species of Viscaceae, and one species of Ximeniaceae. In addition, a total of 37 species are facultative hemiparasites, corresponding to all the species of Orobanchaceae in the country. Conversely, only 19 are holoexoparasites (from families Balanophoraceae Cytinaceae, Convolvulaceae, Ehretiaceae and Mitrastemonaeae), and three holoendoparasites (all members of the Apodanthaceae).

List of parasitic angiosperms in Colombia with comments

Magnoliids

1. Laurales: One family.

Lauraceae Juss. One genus.

Cassytha Osbeck. One stem holoexoparasitic species (Fig. 2).

Fig. 2
figure 2

Map of distribution of Cassytha (Lauraceae), Krameria (Krameriaceae), Apodanthes and Pilostyles (Apodanthaceae), and Sanguisuga (Cytinaceae) in Colombia

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C. filiformis L. (J. Cuatrecasas 17919, COL). Regions II, IV, and V, at elevations between 0 and 1000 m.

2. Zygophyllales: One family.

Krameriaceae Dumort. One genus.

Krameria Loefl. Two obligate, root hemiparasitic species (Figs. 2 and 3).

Fig. 3
figure 3

Krameriaceae (Zygophyllales), obligate root hemiparasites; A. B. Krameria ixine, leaves, flowers and fruits (Bolívar: Juan Arias, 80 m.a.s.l., R. Romero-Castañeda 10,478, COL). C, D. K. spartioides, leaves, flowers and fruits (La Guajira: Uribia, 50 m.a.s.l., C. Saravia 2930, COL). Scale bars: 5 mm in A-D

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K. ixine Loefl. (Fig. 3A, B; R. Romero-Castañeda 10478, COL). Region I. Xerophytic thickets of the Caribbean plains, between 0–1000 m.

K. spartioides Klotzsch ex O.Berg (Fig. 3C, D; C. Saravia 2930, COL). Region I. Xerophytic thickets of the Caribbean plains, between 0–1000 m.

3. Cucurbitales. One family.

Apodanthaceae Tiegh. ex Takht. (Figs. 2 and 4). Two genera.

Fig. 4
figure 4

Apodanthaceae (Cucurbitales), stem holoendoparasites. A-D. Apodanthes caseariae (Meta: Villavicencio, 710 m.a.s.l., F. González et al. 4922). A, B, numerous floral buds emerging from the trunk of its host Casearia sp. (Salicaceae); C, flowers at preanthesis and anthesis; D, fruits. E-O. Pilostyles boyacensis (Boyacá: Chíquiza, 2480 m.a.s.l., F. González 4508). E. Severely infected stem of its host Dalea cuatrecasasii (Fabaceae). F–H. Numerous floral buds emerging from the host stems. I. Peeled-off infected host stem showing several P. boyacensis floral buds forming in the host cortex. J. Transverse section of infected host stem showing P. boyacensis floral buds in the host cortex, and emerged, fully formed flowers. K. Flowers at anthesis. L, M. SEM micrograph of staminate (L) and carpellate (M) flowers. N. Berries, top view. O. Plastome map; note reduction to only six or seven functional genes (after Arias et al., 2019). Scale bars: 1 cm in A, B; 4 mm in C, D, I, K, N; 2 cm in E–H; 6 mm in J; 100 µm in L, M

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Apodanthes Poit. One stem holoendoparasitic species.

A. caseariae Poit. (N. Pabón Mora & F. González 494, HUA; Fig. 4A–D). Regions II, IV, and V, at elevations between 0–1300 m. Primarily restricted to hosts from the genus Casearia (Salicaceae).

Pilostyles Guill. Two stem holoendoparasitic species, one endemic to Colombia.

P. boyacensis F.González & Pabón-Mora (F. González 4508, holotype HUA; Fig. 4E–O). Xerophytic thickets from the Eastern Cordillera (Region III), at elevations between 2400 and 2700 m, exclusively parasitizing Dalea cuatrecasasii Barneby (Fabaceae). Endemic.

P. cf. blanchetii (Gardner) R. Br. (I.J. Holton 84, GH, K). Region III (Western Cordillera). This species is reported based on a single historical collection made at about 1000 m of elevation on February 24th, 1853, and remains to be rediscovered in the field and accurately identified (González & Pabón-Mora, 2014a, b). A detailed examination of the host in the two sheets available indicates that it corresponds to Acaciella angustissima (Mill.) Britton & Rose (Mimosoideae).

4 Malvales: One family.

Cytinaceae

A.Rich. (Figs. 2 and 5). One monotypic genus.

Fig. 5
figure 5

Cytinaceae (Malvales), Sanguisuga caesarea, holoexoparasite in roots of Bursera tomentosa (Jacq.) Triana & Planch. (Burseraceae)(Cesar: Valledupar, 290 m.a.s.l., H. Cuadros et al. 6333). A. Underground runner (arrow) and emerging inflorescence. B. Inflorescence. C–D. Details of bracts and flowers. Scale bars: 5 mm in A, B; 1 mm in C, D

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Sanguisuga Fern.Alonso & H.Cuadros. One root holoexoparasitic species. This monospecific genus, endemic to Colombia, is the only member of the family found in South America.

S. caesarea Fern.Alonso & H.Cuadros (H. Cuadros et al. 6333, holotype COL). Region I. Restricted to a few spots of dry Bursera forests of the Caribbean plains, below 500 m of elevation. Endemic.

5 Santalales: Nine families, 28 genera, 189 species.

Balanophoraceae Rich. Six genera (Figs. 6 and 7).

Fig. 6
figure 6

Map of distribution of Corynaea, Helosis, Langsdorffia, Lophophytum, Ombrophytum, and Scybalium (Balanophoraceae), and Acanthosyris and Cervantesia (Cervantesiaceae) in Colombia

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Fig. 7
figure 7

Balanophoraceae (Santalales), root holoexoparasites. A–C. Corynaea crassa (Boyacá: Villa de Leyva, 2820 m.a.s.l., F. González 1604). A, B. Inflorescences and rhizome; C. Detail of emerging inflorescence. D–H. Helosis cayennensis (Antioquia: Puerto Triunfo, 409 m.a.s.l., F. González et al. 4705). D. Multiple inflorescences emerging at intervals from the underground runner; E. Two inflorescence buds. F–G. Detail of the distal (F) and proximal (G) regions of the inflorescence. H. Sagittal section of inflorescence. I–K. Langsdorffia hypogaea (Boyacá: Villa de Leyva, 2600 m.a.s.l., F. González 4712). I. Detail of underground runner and inflorescence buds. J, K. Top view of emerged inflorescences. Scale bars: 1 cm in A-D, J, K; 5 mm in E, H, I; 1 mm in F-G

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Corynaea Hook.f. One root holoexoparasitic species (Fig. 7A–C).

C. crassa Hook.f. (F. González 1604, COL). Region III. Restricted to primary or healthy secondary Andean forests, at elevations between 1800 and 2400 m.

Helosis Rich. One root holoexoparasitic species (Fig. 7D–H).

H. cayennensis (Sw.) Rich. ex Spreng. (F. González et al. 4705, COL). Regions II-V. A widespread species from wet lowland forests of the Pacific and Amazonian forests, gallery forests of the Orinoco plains, and sub-Andean forests, at elevations between 150 and 1120 m.

Langsdorffia Mart.. One root holoexoparasitic species (Fig. 7I–K).

L. hypogaea Mart. (F. González 4712, COL). Region III. Restricted to slightly disturbed sub-Andean to Andean forests of the Western, Central and Eastern cordilleras, at elevations between 1400 and 2750 m.

Lophophytum Scott & Endl. One root holoexoparasitic species.

L. weddellii Hook.f. (F. Sander & Co. s.n. without detailed locality, collected on Dec 28, 1890, K).

Ombrophytum Poepp. ex Endl. One root holoexoparasitic species.

O. villamariensis S. Guzm.-Guzm. (D. Pérez 001, isotype: COL). This species, endemic to Colombia, is known only from the type locality in Andean forests of the Central Cordillera of Colombia, at 2490 m (Region III). Endemic.

Scybalium Schott & Endl. One root holoexoparasitic species.

S. depressum Eichler (R. Callejas et al. 3924, HUA). Region III. Species restricted to Andean and high-Andean forests of the Western and Central cordilleras, at elevations between 2000 and 3000 m.

Cervantesiaceae Nickrent & Der (Fig. 6). Two genera.

Acanthosyris (Eichler) Griseb. One root hemiparasitic species.

A. glabrata (Stapf) Stauffer ex Govaerts (G. Rodríguez 1024, COL). Region I. Restricted to dry forest remnants of the Caribbean plains, at elevations between 30 and 40 m.

Cervantesia Ruiz & Pav. Two root hemiparasitic species, one endemic to Colombia.

C. macrocarpa Cuatrec. (J. Cuatrecasas 19330, isotype COL). Region III. Cloud, Andean forests of the Central Cordillera, at elevations between 2640 and 2740 m. Endemic.

C. tomentosa Ruiz & Pav. (E. Méndez 918, COL). Region III. Cloud, high-Andean forests of the Central Cordillera, at elevations around 3420 m.

Loranthaceae Juss. Twelve genera, 84 species, 38 endemic to Colombia (Figs. 8, 9, 10, 11, 12, 13, 14, and 15).

Fig. 8
figure 8

Map of distribution of Aetanthus and Cladocolea (Loranthaceae) in Colombia

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Fig. 9
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Map of distribution of Gaiadendron, Maracanthus and Phthirusa (Loranthaceae) in Colombia

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Fig. 10
figure 10

Large-flowered Loranthaceae (Santalales). A-D. Gaiadendron punctatum, root hemiparasite (Boyacá: Villa de Leyva, 2560 m.a.s.l., F. González 4314). A. Dwarf individuum. B. Leaves and inflorescences. C. Dichasium with opened central flower and preanthetic lateral flowers. D. Fruits and ruminate seeds. E–H. Aetanthus mutisii, stem hemiparasite on Buddleja bullata Kunth (Scrophulariaceae) (Boyacá: Aquitania, 3250 m.a.s.l., F. González et al. 4480). E. Habit; F. Inflorescences; G. Flowers visited by the hummingbird Eriocnemis sp.; H. Detail of distal portion of petals and staminal tube with nectar accumulation (arrow). I–J. A. colombianus, stem hemiparasite on Viburnum cornifolium Killip & A.C. Sm. (Boyacá: Belén, 3150 m.a.s.l., F. González et al. 4795). I. Flowers. J. Fruits. K-L. Psittacanthus corderoi, stem hemiparasite (Amazonas: La Pedrera, 100 m.a.s.l., Z. Cordero & E. Tanimuka 818; photo credit Z. Cordero). M. P. krameri (Suaza-Gaviria 11; photo credit V. Suaza), flowers. N–O. P. acinarius, stem hemiparasite, Casanare: Tauramena, 270 m.a.s.l. (F. González 4282). N. Fruits. O. Six-lobed, massive, green endosperm. P–Q. Tristerix secundus, stem hemiparasite on Ageratina baccharoides (Kunth) R.M. King & H. Rob. (Asteraceae)(Cundinamarca: Tausa, 3420 m.a.s.l., F. González 4567). P. Inflorescence; Q. Flowers visited by the hummingbird Eriocnemis vestita. Scale bars: 50 cm in A; 3 cm in B; 1 cm in C, D, G, H, J-O; 2 cm in I, P; 20 cm in E–F; 10 cm in Q

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Fig. 11
figure 11

Map of distribution of Oryctanthus (Loranthaceae, first part) in Colombia

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Fig. 12
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Map of distribution of Oryctanthus (Loranthaceae, second part) in Colombia

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Fig. 13
figure 13

Small-flowered, stem hemiparasitic Loranthaceae (Santalales). A–D. Oryctanthus callicarpus (Valle del Cauca: La Cumbre, 1650 m.a.s.l., N. Pabón-Mora et al. 382). A. Leaves and young inflorescences. B. Detail of inflorescence; arrows point to self-parasitic seedlings. C. Detail of flowers and forming fruits. D. Seedlings. E–F. Peristethium archeri (Antioquia: Santa Elena, 1890 m.a.s.l., N. Pabón-Mora et al. 381). E. Flowers at anthesis. F. Mature fruits. G-I. Passovia pyrifolia (Boyacá: Gachantivá, 2120 m.a.s.l., F. González 4713). G. Leaves and inflorescences. H. Flowers and floral visitor. I. Fruits. J–M. P. pedunculata (Boyacá: Villa de Leyva, 2820 m.a.s.l., F. González & S. González 4918). J. Epicortical roots on Quercus humboldtii (Fagaceae). K. Inflorescence. L. Dichasium with successive stages of floral anthesis and fruit formation. M. Floral visitors, Hybotidae (Diptera), two of them showed in detail (inset). N–S. Struthanthus phillyreoides (Casanare: Yopal, 350 m.a.s.l., F. González 4759). N–P. Seedlings at successive developmental stages. Q. Leaves and runner with secondary epicortical roots. R. Leaves and inflorescences. S. Detail of inflorescence with flower at anthesis. Scale bars: 1 cm in A–C, Q, R; 2 mm in D, E, H, L, S; 5 mm in F, G, I, K; 3 cm in J; 4 mm in M–P

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Fig. 14
figure 14

Map of distribution of Passovia, Peristethium (part 1) and Phthirusa (Loranthaceae) in Colombia

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Fig. 15
figure 15

Map of distribution of Peristethium (part 2), Struthanthus, Tripodanthus and Tristerix (Loranthaceae, four part) in Colombia

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Aetanthus (Eichler) Engl. Ten obligate, stem hemiparasitic species, eight endemic to Colombia (Figs. 10E–J and 26A).

A. alternifolius F.J.Roldán, Carmona, Alzate & J.S.Murillo (Hinestroza et al. 182, holotype HUA). Region III. Known only from the type specimen, collected in oak (Quercus humboldtii Bonpl.; Fagaceae) high-Andean forests of the Eastern Cordillera, nearby the Páramo de Santurbán, at elevations close to 2800 m. Endemic.

A. colombianus A.C.Sm. (F. González et al. 4795, COL). Regions I and III. Cloud Andean or high-Andean forests of the Eastern Cordillera, plus two disjunct populations in the Sierra Nevada de Santa Marta and the Serranía del Perijá, at elevations between 2100 and 3200 m. (Fig. 10I–J). Endemic.

A. engelsii Engl. (Engels s.n., lectotype P). Region III. Without exact locality at the Eastern Cordillera. Endemic.

A. megaphyllus Kuijt (F.J. Hermann 10901; holotype US). Region III. Disjunct species of high-Andean forests to subpáramos of the Central (Antioquia) and Eastern (Norte de Santander) cordilleras, at elevations between 3000 and 3100 m. Endemic.

A. mutisii Engl. (F. González et al. 4480, COL). Region III. Disjunct species from cloud, high-Andean forests, subpáramos and páramos of the Western (Valle del Cauca) and Eastern (Boyacá and Cundinamarca) cordilleras, at elevations between 3000 and 4000 m (Fig. 10E–H). Endemic.

A. nodosus Engl. (G. Lozano-C. 3684, COL). Region III. Cloud Andean forests of the Western and Central cordilleras, at elevations between 2700 and 2800 m.

A. ovalis Rusby (J.H. Torres & P. Pinto 2957, COL). Region I. Endemic to the Sierra Nevada de Santa Marta, in cloud forests at elevations between 2400 and 2600 m. Endemic.

A. sessilifolius Kuijt (J. L. Zarucchi et al. 6501; isotype NY). Region III. Endemic to Andean forests of the northern slopes of the Western Cordillera, at elevations between 1900 and 2100 m. Endemic.

A. trifolius Kuijt (F.J. Roldán et al. 1082, HUA). Region III. Andean and high-Andean forests of the northern slopes of the Central Cordillera, at elevations between 2200 and 2900 m.

A. validus Kuijt (J. Cuatrecasas et al. 12393, isotype COL). Region III. Endemic to Andean forests of the northern slopes of the Eastern Cordillera, at elevations between 2000 and 2200 m. Endemic.

Cladocolea Tiegh. One obligate, stem hemiparasitic species (Fig. 8).

C. coriacea Kuijt (F. González 3050-A, COL). Region IV. Restricted to gallery forests of the Orinoco plains, at elevations between 180 and 200 m. This is the first record of the species for the Flora of Colombia.

Gaiadendron G.Don f. One obligate, root hemiparasitic species (Figs. 9 and 10A–D).

G. punctatum (Ruiz & Pav.) G.Don (F. González 4314, COL). Regions I and III. Widespread through Andean and high-Andean forests, and páramos of the Western, Central and Eastern cordilleras, and the Sierra Nevada de Santa Marta, at elevations between 1500 and 3500 m.

Maracanthus Kuijt. One obligate, stem hemiparasitic species (Fig. 9).

M. chlamydatus (Rizzini) Kuijt (A. Sudgen 119, COL). Region I. Dry forests of the Caribbean plains, at elevations between 15–100 m.

Oryctanthus Eichler. Eleven obligate, stem hemiparasitic species, four of them endemic to Colombia (Figs. 11, 12, 13A–D, and 26B).

O. asplundii Kuijt (F. González et al. 2966A, COL). Region III. Cloud, Andean forests of the Western Cordillera, at elevations between 1500 and 1700 m.

O. callicarpus Kuijt (N. Pabón-Mora et al. 382, HUA). Regions II and III. Lowland, pluvial forests, and sub-Andean and Andean forests of the Western, Central and Eastern cordilleras, at elevations between 0 and 2050 m (Fig. 13A–D).

O. cordifolius (C.Presl) Urb. (J.N. Díaz & F. González 129, COL). Regions I (dry forests of the Caribbean plains and the Archipelago of San Andrés and Old Providence, II, and III, at elevations between 10 and 1500 m.

O. costulatus Rizzini (F.J. Roldán 1953, HUA). Regions III, IV, and V, at elevations between 100 and 1800 m.

O. florulentus Urb. (J. Cuatrecasas 26020, COL). Regions II (pluvial forests), III (sub-Andean and Andean forests of the Western, Central and Eastern cordilleras), IV (thickets and gallery forests of the Orinoco plains) and V (Amazonian forests), at elevations between 150 and 2500 m.

O. grammatus Kuijt (H.M. Curran s.n., holotype US). Region I. Caribbean dry forests, and inter-Andean xerophytic thickets of the northern Magdalena valley, at elevations below 300 m. Endemic.

O. grandis Kuijt (E.P. Killip & J. Cuatrecasas 38974, isotype COL). Region II. Pluvial forests at elevations between 10 and 1500 m. Endemic.

O. neurophyllus Kuijt (A.H. Gentry et al. 53663, isotype UC). Region II. Pluvial forests, at elevations between 50 and 500 m. Endemic.

O. perticus Carmona, F.J. Roldán, Alzate & J.S. Murillo (C.A. Loaiza 182, holotype HUA). Region III. Known only from the type locality, in sub-Andean forests of the northern end of the Central Cordillera, at elevations between 1000 and 1500 m. Endemic.

O. phthirusoides Rizzini (M. Arbeláez et al. 525, HUA). Region V. Amazonian forests at elevations between 100 and 400 m.

O. spicatus (Jacq.) Eichler (C. Sastre 2234, COL). Widespread throughout regions I, II, III, IV, and V, at elevations between 20 and 2600 m.

Passovia H. Karst. Four obligate, stem hemiparasitic species, one endemic to Colombia. (The taxonomic outline of this genus follows Caires et al. 2021). (Figs. 13G–I and 14).

P. cordata Kuijt (A.H. Gentry et al. 53723, holotype UC). Region II. Pluvial forests at elevation around 500 m. Endemic.

P. pedunculata (Jacq.) Kuijt (F. González & S. González 4918, COL). Andean forests from the Eastern Cordillera, at elevations between 1800–2600 m (Region III), in addition to three scattered locations at the Caribbean plains (Region I) and two collections from the Amazonian forests below 500 m of elevation (Region V) (Fig. 13J–M).

P. pyrifolia (Kunth) Tiegh. (F. González 4713, COL). Region I (dry forests of the Caribbean plains, at elevations below 300 m), III (Andean forests at elevations between 1800 and 2500 m), and V (Amazonian forests below 400 m) (Fig. 13G–I).

P. robusta (Rusby) Kuijt (I.F. Holton 651, K). Region III. Sub-Andean forests at about 1000 m of elevation.

Peristethium Tiegh. Six obligate, stem hemiparasitic species, two of them endemic to Colombia (Fig. 13E–F and 14).

P. aequatoris (Kuijt) Kuijt (J. Cuatrecasas 11540, COL). Region III. Eastern slopes of the department of Putumayo, at elevations between 1500–2200.

P. archeri (A.C. Sm.) Kuijt (N. Pabón-Mora et al. 381, HUA). Regions II and III. Pluvial forests and sub-Andean forests of the Western and Central cordilleras, at elevations between 80 and 2200 m (Fig. 13E–F).

P. colombianum Kuijt (D. Macías & B. R. Ramírez 5001, holotype CAUP). Region II. Pluvial forests at elevations below 800 m. Endemic.

P. lamprophyllum Kuijt (H.P. Fuchs & L. Zanella 22060, isotype F). Region II. Pluvial forests, at elevations around 50 m. Endemic.

P. leptostachyum (Kunth) Tiegh. (J.J. Triana 4586/1, COL). Regions II, III and V. Pluvial, sub-Andean and Amazonian forests, at elevations between 40 and 1800 m.

P. polystachyus (Ruiz & Pav.) Kuijt (H. Daniel 6262, COL). Regions II and III. Pluvial and sub-Andean forests at elevations between 500 and 1700 m.

Phthirusa Mart. Three obligate, stem hemiparasitic species, two of them endemic to Colombia (Fig. 9).

P. exilis Kuijt (F.J. Roldán 750, HUA). Region III. Sub-Andean to high-Andean forests at the northern end of the Western and Central cordilleras, at elevations between 1600 and 2400 m. Endemic.

P. hutchisonii (Kuijt) Kuijt (F.J. Roldán et al. 871, HUA). Region III. Sub-Andean and Andean forests of the Western, Central and Eastern cordilleras, at elevations between 1300 and 2300 m.

P. rhynchophylla (Kuijt) Kuijt (F.J. Roldán et al. 1141, HUA). Region III. Disjunct species in cloud, Andean forests of the Western (Antioquia) and the Eastern (Boyacá) cordilleras, at elevations between 1600 and 2800 m. Endemic.

Psittacanthus Mart. Thirty-seven obligate, stem hemiparasitic species, 17 of them endemic to Colombia (Figs. 10K–O and 26C).

P. acinarius Mart. (F. González 4282, COL). Regions I, III, IV and V. Sclerophyllous, lowland and gallery forests, as well as in Amazonian and sub-Andean forests, at elevations between 300 and 1000 m (Fig. 10N, O).

P. antioquiensis Kuijt (F.J. Roldán 1550, isotype HUA). Region III. Cloud, Andean forests, at elevations between 1600 and 1850 m. Endemic.

P. biternatus (Hoffmans.) G.Don. (R. Liesner 9135, COL). Region V. Sclerophyllous, Amazonian forests, at elevations between 100 and 150 m.

P. bolbocephalus Kuijt (J.L. Zarucchi 1205, LEA). Region V. Sclerophyllous, Amazonian forests, at elevations between 300 and 350 m.

P. carinatus Kuijt (E.L. Core 450, holotype US). Region III. Cloud, high-Andean forests, at elevations between 2800 and 3100 m. Endemic.

P. cinctus Mart. (B. Maguire & J. Wurdack 35668, COL). Region V. Sclerophyllous, Amazonian forests, at elevations between 120 and 500 m.

P. clusiifolius Eichler (B. Maguire 41437, COL). Region V. Sclerophyllous, Amazonian forests, at elevations between 100 and 150 m.

P. complanatus Kuijt (R. Callejas et al. 7844, isotype HUA). Region III. Cloud, high-Andean forests, at elevations between 3000 and 3200 m. Endemic.

P. corderoi F.González, F.J. Roldán & Pabón-Mora (Z. Cordero & E. Tanimuka 818, holotype COL). Region V. Sclerophyllous, Amazonian forests, at elevations around 100 m (Fig. 10K, L). Endemic.

P. crassicostatus Kuijt (F.J. Roldán 1218, isotype COL) Regions II and III. Pluvial and humid, sub-Andean forests, at elevations between 800 and 900 m. Endemic.

P. crassifolius Mart. s.l. (included P. baguensis Kuijt; see Dettke & Caires, 2021; V. Wijninga & Quintero 605, COL). Region V. Sclerophyllous, Amazonian forests, at elevations between 200 and 350 m.

P. crassinervis Kuijt (J. Cuatrecasas 9058, COL). Region III. Cloud, sub-Andean forests, at elevations between 1000 and 1300 m. Endemic.

P. cucullaris (Lam.) Blume (E. Pérez-Arbeláez 693, COL). Regions III, IV and V. Sclerophyllous, gallery forests of the Orinoco plains and Amazonian forests, and sub-Andean forests at elevations between 100 and 1500 m.

P. cyclophyllus Kuijt (P. Silverstone-Sopkin et al. 4525, holotype CUVC). Regions II, III, and V. Pluvial and cloud, sub-Andean to Andean forests, at elevations between 500 and 2600 m. Endemic.

P. dilatatus A.C.Sm. (R. Callejas et al. 8449, HUA). Region III. Cloud belts of the sub-Andean and Andean forests, at elevations between 1200 and 2800 m.

P. eucalyptifolius Blume (R.E. Schultes et al. 10210, US). Region V. Sclerophyllous Amazonian forests, at elevations between 100 and 150 m.

P. geniculatus Kuijt (A.H. Gentry et al. 22115, F). Region V. Sclerophyllous Amazonian forests, at elevations between 100 and 200 m.

P. gigas Kuijt (A.H. Gentry 48280, isotype COL). Region II. Pluvial forests, at elevations between 5 and 50 m. Endemic.

P. hamulifer Kuijt (L. Albert et al. 1627, HUA). Regions II and III. Pluvial and sub-Andean forests, at elevations between 100 and 1800 m.

P. irwinii Rizzini (R.E. Schultes & I. Cabrera 15029, NY). Region V. Sclerophyllous, Amazonian forests, at elevations between 100 and 300 m.

P. krameri Kuijt (V. Suaza-Gaviria 11, HUA). Regions II and III. Pluvial and sub-Andean forests, at elevations between 100 and 1000 m (Fig. 10M).

P. longerectus F.J.Roldán & Alzate (F.J. Roldán et al. 1155, holotype HUA). Region III. Cloud, Andean to high-Andean forests, at elevations between 2550 and 2700 m. Endemic.

P. micrantherus Kuijt (M. Monsalve 1383, holotype UC). Region II. Pluvial forests, at elevations around 100 m. Endemic.

P. microphyllus Kuijt (F.J. Roldán et al. 1149, isotype HUA). Region III. Cloud, high-Andean forests, subpáramos and páramos, at elevations between 2800 and 3200 m. Endemic.

P. ophiocephalus Kuijt (R.E Schultes & I. Cabrera 15787, holotype US). Region V. Sclerophyllous Amazonian forests, at elevations around 200 m. Endemic.

P. pentaphyllos Kuijt (J.L. Zarucchi et al. 6765, isotype HUA). Region III. Cloud, sub-Andean forests, at elevations between 1000 and 1100 m. Endemic.

P. peronopetalus Eichler (Fernández 2036, US). Regions III and V. Amazonian and sub-Andean forests, at elevations between 200 and 1350 m.

P. rhynchanthus (Benth.) Kuijt (J. Brand & A. Cogollo 99, COL). Regions I and II. Dry to wet forests, at elevations between 10 and 100 m.

P. robustus Mart. (M. Monsalve 173, COL). Region V. Sclerophyllous Amazonian forests, at elevations between 100 and 400 m.

P. roldanii Kuijt (F.J. Roldán et al. 796, isotype HUA). Region I. Dry forests, at elevations between 20 and 70 m.

P. rotundatus Kuijt (F.J. Roldán et al. 817, isotype HUA). Region III. Cloud, sub-Andean forests, at elevations between 1000 and 1500 m. Endemic.

P. schultesii Kuijt (R.E. Schultes & I. Cabrera 15111, isotype GH). Region V. Sclerophyllous, Amazonian forests, at elevations around 300 m. Endemic.

P. smithii Kuijt (H.H. Smith 1491, holotype NY). Region I. Dry forests, at elevations between 160 and 170 m. Endemic.

P. stergiosii Kuijt (J. Wurdack & Adderley 43213, isotypes F, US). Region V. Sclerophyllous Amazonian forests, at elevations between 100 and 150 m.

P. sulcatus Kuijt (R.E. Schultes & I. Cabrera 13351, holotype US). Region V. Sclerophyllous Amazonian forests, at elevations between 250 and 700 m. Endemic.

P. ternatus Kuijt (F. Alzate et al. 259, isotype HUA). Region III. Cloud, sub-Andean to Andean forests, at elevations between 1450 and 1750 m.

P. truncatus Kuijt (A. Rudas et al. 7153, COL). Regions II, III, V. Pluvial, sub-Andean and Amazonian forests, at elevations between 400 and 1800 m.

Struthanthus Mart. Seven obligate, stem hemiparasitic species, one of them endemic to Colombia (Figs. 13N–S and 15).

S. calophyllus A.C. Sm. (F.J. Roldán et al. 1064, COL). Region III. Sub-Andean forests, at elevations between 1400 and 1500 m.

S. marginatus (Desr.) G. Don (A. Dugand 5793, COL). Region I. Dry forests and savannas, at elevations below 400 m.

S. orbicularis (Kunth) Eicher (F.J. Roldán 1653, COL). Regions I, II and III. Savanna vegetation, gallery forests, and sub-Andean forests, at elevations between 40 and 1200 m.

S. phillyreoides (Kunth) G. Don (F. González 4759, COL). Regions I, IV. Savanna vegetation and gallery forests, at elevations around 200 m. (Fig. 13N–S).

S. schultesii Kuijt (R.E. Schultes 16996, isotype COL). Region V. Sclerophyllous.

Amazonian forests, at elevations between 150 and 300 m. Endemic.

S. subtilis Kuijt (F.J. Roldán 1222, COL). Region III. Cloud sub-Andean and Andean forests, at elevations between 1600 and 2600 m.

S. syringifolius Mart. (J. Cuatrecasas 7065, COL). Regions I and V. Dry Caribbean forests, and sclerophyllous Amazonian forests, at elevations between 100 and 240 m.

Tripodanthus Tiegh. One obligate, stem hemiparasitic species endemic to Colombia (Fig. 15).

T. belmirensis F.J. Roldán & Kuijt (R. Fonnegra & D. Tuberquia 4576, holotype HUA). Region III. Subpáramo and páramo vegetation of the Western and the Central cordilleras, at elevations between 3000 and 3130 m of elevation. Endemic.

Tristerix Mart. Two obligate, stem hemiparasitic species, one of them endemic to Colombia (Figs. 10P, Q and 15).

T. longibracteatus (Desr.) Barlow & Wiens (F. González 4270, COL). Region III. High-Andean forests of the Western and Central cordilleras (Region III), at elevations between 3100 and 3700 m.

T. secundus (Benth.) Kuijt (F. González 4567, COL). Region III. Subpáramo and páramo vegetation of the Eastern Cordillera, at elevations between 2800 and 4000 m (Fig. 10P, Q). Endemic.

Opiliaceae Valeton. One genus.

Agonandra Miers ex Benth. Three root hemiparasitic species (Fig. 16).

Fig. 16
figure 16

Map of distribution of Agonandra (Opiliaceae), Antidaphne and Eubrachion (Santalaceae), Schoepfia (Schoepfiaceae), Tetrastylidium (Strombosiaceae), Ximenia (Ximeniaceae), Mitrastemon (Mitrastemonaceae), and Lennoa (Ehretiaceae) in Colombia

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A. brasiliensis Benth. & Hook.f. (A. Dugand 5662, COL) Region I. Dry forests below 400 m of elevation.

A. peruviana Hiepko. Region V. Sclerophyllous Amazonian forests below 400 m of elevation.

A. silvatica Ducke (A. Rudas et al. 4836, COL). Region IV. Gallery forests below 210 m of elevation.

Santalaceae R. Br. Two genera (Figs. 16 and 17).

Fig. 17
figure 17

Santalaceae, stem hemiparasite. Antidaphne viscoidea (Boyacá: Arcabuco, 2690 m.a.s.l., F. González 4645, ♂, 4645 A, ♀). A. Collecting site of carpellate individual. B. Primary haustorium of carpellate individual. C. Runner with secondary haustoria of staminate individual on branch of Quercus humboldtii (Fagaceae). D, E. Leaves; note (in E) egg cluster of Archonias or Catasticta sp. (Lepidoptera, Pieridae). F-G. Proliferative inflorescences of staminate individual; note (in G) anthetic flowers with highly reduced perianth. H. Proliferative, sinuous carpellate inflorescences. Scale bars: 2 cm in B; 5 mm in C-H

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Antidaphne Poepp. & Endl. Two obligate, stem hemiparasitic species.

A. andina Kuijt (F.J. Roldan 1894, HUA). Region III. High-Andean forests, at elevations between 2800 and 3200 m.

A. viscoidea Poepp. & Endl. (F. González 4645, ♂, 4645 A, ♀, COL). Region III. Sub-Andean to high-Andean forests and páramo vegetation, at elevations between 900 and 3700 m (Fig. 17).

Eubrachion Hook.f. One obligate, stem hemiparasitic species.

E. ambiguum Engl. (J.L. Fernández 23697, COL). Region III. Andean and high-Andean forests, at elevations between 2600 and 3000 m.

Schoepfiaceae Blume. One genus (Fig. 16).

Schoepfia Schreb. Two root hemiparasitic species.

S. flexuosa Roem. ex Schult. (W. Purdie s.n., GH). Regions I and II (Uraba region). Pluvial forests, at elevations below 100 m.

S. schreberi J.F.Gmel. (J. Goudot s.n., P). Regions I and III (Magdalena valley). Dry forests, at elevations between 50 and 500 m.

Strombosiaceae Tiegh. One genus (Fig. 16).

Tetrastylidium Engl. One root hemiparasitic species.

T. peruvianum Sleumer (J.J. Pipoly 16296, MO). Region V. Sclerophyllous Amazonian forests, at elevations below 100 m.

Viscaceae Batsch. Two genera, 88 species (Figs. 18 and 27).

Dendrophthora Eichler. Twenty-two obligate, stem hemiparasitic species, three of them endemic to Colombia (Figs. 18A–F and 27A).

D. amalfiensis Kuijt (R. Callejas et al. 9135, isotype HUA). Region III. Cloud, sub-Andean forests, at elevations between 1300 and 1900 m. Endemic.

D. ambigua Kuijt (J. Cuatrecasas 18902, isotype A). Region III. Cloud, high-Andean forests and subpáramo and páramo vegetation, at elevations between 3300 and 3900 m.

D. avenia (Trel.) Kuijt (F. González 4482, COL). Region III. Cloud, Andean forests and subpáramo and páramo vegetation, at elevations between 2400–3550 m (Fig. 18A–C).

Fig. 18
figure 18

Viscaceae, stem hemiparasites. A-C. Dendrophthora avenia. A-B. Habitat at subparamo and paramo belts (Boyacá: Toca, 3400–3500 m.a.s.l., F. González 4482). C. Detail of leaf bases, inflorescences and flowers. D. D. clavata (Boyacá: Duitama, 3200 m.a.s.l., F. González 1589), individual with mixed staminate and carpellate inflorescences. E–F. D. lindeniana (Antioquia: Santa Elena, m.a.s.l., N. Pabón-Mora 482). E. Main haustorium. F. Leaves and inflorescences. G-M. Phoradendron nervosum (Boyacá: Villa de Leyva, 2450 m.a.s.l., F. González 4554). G-H. Staminate individual (G) and detail of inflorescence with flower at anthesis (top right in H). I-K. Carpellate inflorescences with mature fruits. L. Dissected fruits showing green seeds. M. Seedlings. Scale bars: 5 mm in C-E, K; 1 cm in F, I, J; 5 cm in G; 2 mm in L, M

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D. chrysostachya Urb. (E. Pérez-Arbeláez & J. Cuatrecasas 5911, COL). Region III. Cloud, high-Andean forests and subpáramo and páramo vegetation, at elevations between 3100 and 3500 m.

D. clavata Urb. (F. González 1589, COL). Regions III (sub-Andean, Andean and high-Andean forests, and subpáramo and páramo vegetation) and IV (gallery forests), at elevations between 500 and 3700 m. Likely the parasitic angiosperm with the broadest elevational distribution in Colombia (Fig. 18D).

D. costaricensis Urb. (B. Ramírez 16288, COL). Region III. Cloud, Andean to high-Andean forests, at elevations between 2400 and 3200 m.

D. elegantissima Kuijt (F.J. Roldán et al. 1895, HUA). Region III. Cloud, high-Andean forests, and subpáramo and páramo vegetation at elevations between 3000 and 3550 m.

D. elliptica Krug & Urb. (J. Cuatrecasas 25368, COL). Regions III (sub-Andean and Andean forests) and IV (gallery forests), at elevations between 850 and 2300 m.

D. fendleriana (Eichler) Kuijt (R. Callejas et al. 7907, HUA). Region III. Cloud, Andean forests, at elevations between 2000 and 2100 m.

D. lindeniana Tiegh. (N. Pabón-Mora 482, HUA). Region III. Cloud, Andean to high-Andean forests and subpáramo vegetation, at elevations between 1600 and 3300 m (Fig. 18E, F).

D. lueri Kuijt (G. Lozano 6704, COL). Region III. Cloud, Andean forests, at elevations between 2900 and 3200 m.

D. macbridei (Standl.) Kuijt (P. Franco 2752, COL). Region III. Cloud, Andean forests, at elevations between 2000 and 2400 m.

D. obliqua (C. Presl) Wiens (E. Pérez-Arbeláez 8343, COL). Region III. Cloud, sub-Andean to Andean forests, at elevations between 730 and 2900 m.

D. paucifolia (Rusby) Kuijt (F.J. Roldán et al. 826, HUA). Region III. Cloud, Andean to high-Andean forests, at elevations between 2500 and 3300 m.

D. pearcei (Rusby) Kuijt (S. Díaz-Piedrahíta 4232, COL). Region III. Cloud, Andean to high-Andean forests and subpáramo and páramo vegetation, at elevations between 2450 and 3700 m.

D. squamigera (Benth.) Benth. & Hook.

f. ex Kuntze (K.T. Hartweg 1046, holotype K). Region III. Subpáramo and páramo vegetation, at elevations between 3100 and 3300 m.

D. stricta Rusby (H.H. Smith 1289, holotype NY). Region I. Cloud, montane forest relicts in the Sierra Nevada de Santa Marta, at elevations around 2200 m. Endemic.

D. subtrinervis Urb. (J.H. Torres 680, COL). Region III. Cloud, Andean to high-Andean forests, and subpáramo and páramo vegetation, at elevations between 2800 and 3500 m.

D. sulcata Kuijt (R. Callejas et al. 8498, isotype HUA). Region III. Cloud, sub-Andean to Andean forests, at elevations between 1450 and 1840 m. Endemic.

D. tenuifolia Kuijt (B. Ramírez 4029, HUA). Region III. Cloud, high-Andean forests at elevations around 3000 m.

D. thomasii Kuijt (S. Espinal et al. 2400, HUA). Region III. Cloud, high-Andean forests at elevations between 3300 and 3700 m.

D. virgata (Trel.) Kuijt (R. Callejas et al. 6282, HUA). Region III. Cloud, Andean to high-Andean forests and subpáramo and páramo vegetation at elevations between 2400 and 3600 m.

Phoradendron Nutt. Sixty-six obligate, stem hemiparasitic species, 17 of them endemic to Colombia (Figs. 18G–M and 27B).

P. anceps (Spreng.) M.Gómez (F.J. Roldán 1284, HUA). Region III. Sub-Andean forests, at elevations between 1400 and 1500 m.

P. antioquianum Kuijt (F.J. Roldán 1224, holotype HUA). Region III. Sub-Andean forests, at elevations between 1550 and 1850 m. Endemic.

P. apertiflorum Kuijt (G. Galeano et al. 2274, holotype COL). Region V. Sclerophyllous Amazonian forests, at elevations between 350 and 500 m. Endemic.

P. applanatum Kuijt (O. Haught 4181, COL). Region I. Xerophytic thickets, at elevations between 50 and 500 m.

P. balslevii Kuijt (G. Lozano 4108, COL). Region III. Andean forests, at elevations between 1800 and 2500 m.

P. berteroanum Griseb. (L. Albert 163, COL). Regions I, II, III. Dry forests, pluvial forests and sub-Andean to Andean forests, at elevations between 350 and 2000 m.

P. betancurii Kuijt (O. Haught 2437, holotype COL). Region III. Sub-Andean to Andean forests, at elevations between 700 and 2200 m. Endemic.

P. brittonianum Rusby (C.A. Loaiza et al. 138, HUA). Region III. Sub-Andean forests, at elevations between 1400 and 1500 m.

P. buritacanum Kuijt (C. Barbosa et al. 354, isotype COL). Region I, Montane forests of the Sierra Nevada de Santa Marta, at elevations around 1100 m. Endemic.

P. chrysocladon A.Gray (J. Cuatrecasas 12514, COL). Regions II and III. Pluvial forests to Andean forests, at elevations between 30 and 2400 m.

P. colombianum Kuijt (J.M. MacDougal & F.J. Roldán 3543, isotype HUA). Region III. Sub-Andean to high-Andean forests, at elevations between 1650 and 3000 m.

P. crassicarpum Kuijt (F.J. Roldán 801, holotype HUA). Region III. Sub-Andean forests, at elevations between 500 and 1000 m. Endemic.

P. crassifolium Trel. (J.M. Idrobo 10576, COL). Andean forests (Region III), gallery forests (Region IV) and Amazonian forests (Region V) at elevations between 100 and 2000 m.

P. dipterum Eichler (F. González 4445, COL). Region III. Sub-Andean to Andean forests, at elevations between 1800 and 2600 m.

P. eggersii Urban (O. Haught 4020, COL) Regions I and III. Xerophytic thickets (Region I) to sub-Andean forests (Region III), at elevations between 100 and 800 m.

P. englerianum Patsch. (G. Lozano-Contreras 904, COL). Xerophytic thickets (Region I) to Andean forests (Region III), at elevations between 100 and 2400 m.

P. exiguum Trel. (H.H. Smith 1281, isotype NY). Region I. Xerophytic thickets, at elevations between 10 and 200 m.

P. fasciculatum Kuijt (F.J. Roldán 864, HUA). Region I. Dry forests and xerophytic thickets, at elevations below 800 m.

P. fonsecanum Kuijt (O. Haught 3992, isotype COL). Region I. Dry forests and xerophytic thickets, at elevations between 100 and 200 m. Endemic.

P. gracilicarpum Kuijt (R. Callejas 8905, holotype HUA). Region III. Sub-Andean forests, at elevations between 1000 and 1100 m. Endemic.

P. haughtii Kuijt (O. Haught 6302, isotype COL). Region III. Sub-Andean forests, at elevations between 450 and 1000 m.

P. herbert-smithii Trel. (H.H. Smith 1283, isotype COL). Regions I, III. Dry forests (Region I) and sub-Andean forests (Region III), below 1200 m in elevation.

P. heterostachyum Kuijt (N. Contreras & D.L. Echeverri 353, isotype HUA). Region III. Sub-Andean forests, at elevations between 1700 and 1900 m. Endemic.

P. hexastichum (DC.) Griseb. (F.J. Roldán et al. 781, HUA). Region III. Sub-Andean and Andean forests, at elevations between 800 and 2000 m.

P. inaequidentatum Rusby (R.E. Schultes 3625, COL). Sub-Andean forests (Region III) and sclerophyllous Amazonian forests (Region V), at elevations between 100 and 2100 m.

P. killipii Kuijt (E.P. Killip 10102, holotype NY). Region III. Andean forests, at elevations between 2000 and 2700 m. Endemic.

P. larense Kuijt (F.J. Roldán et al. 1087, HUA). Region III. Sub-Andean forests, at elevations between 900 and 1200 m.

P. laxiflorum Ule (J. Cuatrecasas et al. 27192, COL). Region V. Amazonian forests, at elevations between 100 and 500 m.

P. linearifolium Eichler (R. López 3822, HUA). Region V. Amazonian forests, at elevations below 300 m.

P. longissimum Kuijt (F.J. Roldán 791, isotype HUA). Region I. Dry forests and xerophytic thickets, at elevations below 30 m. Endemic.

P. lorifolium Kuijt (F.J. Roldán 794, holotype HUA). Region I. Dry forests and xerophytic thickets, at elevations below 30 m.

P. madisonii Kuijt (B. Madrigal 667, COL). Region II. Pluvial forests, at elevations between 10 and 400 m.

P. mairaryense Ule (A. Fernández 6946, HUA). Region V. Amazonian forests, at elevations between 50 and 120 m.

P. megaphyllum Kuijt (J.C. Mutis 161, US). Region III. Cloud, Andean forests, at elevations between 2000 and 2500 m. Endemic.

P. metense Kuijt (O. Haught 2530, isotype COL). Regions III (Sub-Andean forests) and IV (savannas and gallery forests), at elevations between 400 and 1600 m.

P. mucronatum (DC.) Krug & Urb. (H.H. Smith 1285, COL). Regions I and III. Dry forests and sub-Andean forests, at elevations between 0 and 1000 m.

P. nervosum Oliv. (F. González 4554, COL). Regions I and III. Dry forests and sub-Andean to high-Andean forests, at elevations between 100 and 3000 m (Fig. 18G–M).

P. nitens Kuijt (J. Zarucchi et al. 6740, HUA). Region III. Sub-Andean forests, at elevations between 600 and 900 m.

P. obtusissimum Eichler (A. Chaparro 928, COL). Sub-Andean forests (Region III) and Amazonian forests (Region V), at elevations between 100 and 2100 m.

P. palaephyllum Kuijt (J.M. Idrobo & R.E. Schultes 638, holotype COL). Region V. Sclerophyllous, Amazonian forests, at elevations between 100 and 300 m. Endemic.

P. parietarioides Trel. (F. González et al. 2263, COL). Sub-Andean to high-Andean forests (Region III) and Amazonian forests (Region V), at elevations between 500 and 3000 m.

P. pellucidulum Eichler (G. Lozano-Contreras et al. 2736, COL). Region I. Dry forests, at elevations between 50 and 200 m.

P. perrottetii (DC.) Eichler (J.C. Mutis 161A, MA). Regions III and V. Amazonian and sub-Andean forests, at elevations between 900 and 1400 m.

P. piperoides (Kunth) Trel. (E. Pérez-Arbeláez & J. Cuatrecasas 8112, COL). Regions II, III and V. Pluvial, sub-Andean, and Amazonian forests, at elevations between 50 and 1500 m.

P. planiphyllum Kuijt (J. Cuatrecasas 19973, F). Region III. Cloud Andean forests, at elevations between 1600 and 2000 m.

P. platycaulon Eichler (G. Lozano-Contreras et al. 433, COL). Regions IV and V. Gallery forests of the Orinoco plains and Amazonian forests, at elevations between 100 and 600 m.

P. pteroneuron Eichler (R. Fonnegra et al. 4754, HUA). Sub-Andean forests (Region III) and gallery forests and savanna vegetation (Region IV), at elevations between 600 and 1800 m.

P. quadrangulare (Kunth) Griseb. (E.P. Killip 33956, COL). Regions I-III. Dry thickets (Region I), pluvial forests (Region II), and sub-Andean forests (Region III), at elevations between 50 and 1200 m.

P. racemosum Krug & Urb. (G. Lozano-Contreras 6359, COL). Region V. Sclerophyllous, Amazonian forests, at elevations between 300 and 900 m.

P. restrepoae Kuijt (C. Restrepo 05, isotype COL). Region III. Andean forests at elevations around 1700 m. Endemic.

P. retrophyllum Kuijt (C. Luer et al. 10242, holotype LEA). Region III. Cloud, high-Andean forests and subpáramo vegetation, at elevations between 3100 and 3200 m. Endemic.

P. robaloense Woodson ex Rizzini (O. Haught 4348, COL). Region I. Xerophytic thickets, at elevations between 10 and 50 m.

P. robustissimum Eichler (F.J. Roldán et al. 1338, HUA). Region III. Sub-Andean forests, at elevations between 600 and 1300 m.

P. roldanii Kuijt (F.J. Roldán et al. 861, isotype HUA). Region III. Sub-Andean forests, at elevations between 1000 and 1200 m. Endemic.

P. rugosum Kuijt (A. Chaparro 899, holotype COL). Region III. Sub-Andean forests, at elevations between 1450 and 1550 m.

P. singulare Kuijt (S. Castroviejo et al. 12,057, isotype COL). Region V. Sclerophyllous thickets at the summit of the Sierra de Chiribiquete, at elevations between 500 and 800 m.

P. squamiferum Kuijt (A.H. Gentry et al. 15,440, isotype COL). Region III. Sub-Andean forests, at elevations around 1900 m. Endemic.

P. staphylinum Rizzini (J. Cuatrecasas 9669, COL). Region III. Sub-Andean forests, at elevations between 1800 and 1900 m.

P. strongyloclados Eichler (report based on Kuijt, 2003). Region V.

P. tardispicum Kuijt (F.J. Roldán et al. 635, HUA). Regions II and III. Pluvial and sub-Andean forests, at elevations between 100 and 800 m.

P. triflorum E.A.Kellogg (A.H. Gentry 40464, COL). Region II. Pluvial forests, at elevations between 50 and 400 m.

P. trinervium (Lam.) Griseb. (G. Davidse & J.S. Miller 26383, MO). Regions I and V. Disjunct species from dry Caribbean forests, and sclerophyllous Amazonian forests, at elevations between 5 and 100 m.

P. undulatum (Pohl ex DC.) Eichler (E. Pérez-Arbeláez & J. Cuatrecasas 8112, COL). Regions I and III, at elevations between 800 and 3500 m.

P. uniseriale Kuijt (F.J. Roldán et al. 732, holotype HUA). Region III. Cloud, sub-Andean forests, at elevations between 1000 and 1400 m. Endemic.

P. wiensii Kuijt (report based on Kuijt, 2003). Region III. Cloud, sub-Andean forests, at elevations between 1350 and 1450 m.

P. woodsonii Trel. (J. Cuatrecasas 11312, COL). Regions II, III. Pluvial and sub-Andean forests, at elevations between 50 and 1600 m.

Ximeniaceae Horan. One genus (Fig. 16).

Ximenia Plum. ex L. One root hemiparasitic species.

X. americana L. (G. Lozano-Contreras 2944, COL). Regions I (Urabá) and II. Pluvial forests, at elevations below 100 m.

6 Ericales

Mitrastemonaceae Makino. One genus. (Fig. 16).

Mitrastemon Makino. One root holoexoparasitic species.

M. matudae Yamam. (Á. Cogollo et al. 7661, MO). Region III. Cloud, sub-Andean forests, at elevations between 1400 and 1500 m.

7 Boraginales: One family.

Ehretiaceae Mart. One genus (Figs. 16 and 19A–E).

Fig. 19
figure 19

A-E. Lennoa madreporoides (Ehretiaceae), root holoexoparasite; La Guajira: Dibulla, 20 m.a.s.l., scanning micrographs of flowers (A, B), clavate trichome of the outer perianth surface (C), and plurilocular ovary (D, E). F-M. Cuscuta corymbosa (Convolvulaceae), stem holoexoparasite (Boyacá: Corrales, 2560 m.a.s.l., F. González 4794). F. Habit. G-L. Filamentous stems; note (in K, L, arrows) lateral, discoid haustoria formed at intervals in stem portions in contact to the branch host. M. Inflorescence. Scale bars: 50 µm in A, B, D, E; 5 µm in C; 1 mm in F-M

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Lennoa Lex. One root holoexoparasitic species.

L. madreporoides Lex. (M.L. Schnetter 69, COL). Region I. Coastal thickets, at elevations below 25 m.

8 Solanales: One family.

Convolvulaceae Juss. One genus (Figs. 19F–M and 20).

Fig. 20
figure 20

Map of distribution of Cuscuta (Convolvulaceae) in Colombia

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Cuscuta L. Nine twining, stem holoexoparasitic species, two of them endemic to Colombia.

C. americana L. (J.M. Idrobo 6625, COL). Regions I and III. Dry forests and xeromorphic thickets, and sub-Andean and Andean forests, at elevations between 20 and 2500 m.

C. colombiana Yunck. (C. Saravia 2304, COL). Region I. Xeromorphic thickets, at elevations between 30 and 300 m. Endemic.

C. corymbosa Ruiz & Pav. (F. González 4794, COL). Region III. Xerophytic thickets, at elevations between 2400 and 2500 m (Fig. 19F–M).

C. grandiflora Kunth (J.J. Triana 3816/1, COL). Region III. Sub-Andean through high-Andean forests, and subpáramos and páramos, at elevations between 500 and 4000 m.

C. indecora Choisy (E.P. Killip 34700, COL). Region III. Sub-Andean and Andean forests, at elevations between 1000 and 2550 m.

C. obtusiflora Kunth (F.A. Barkley 17C336, COL). Region III. Restricted to sub-Andean forests of the Central Cordillera in Antioquia, at elevations between 700 and 1560 m.

C. partita Choisy (C. Saravia 527, COL). Region I. Xerophytic thickets below 50 m.

C. pentagona Engelm. (W.R. Jervise s.n., MO). Only one (historic) collection from the department of Antioquia, without precise locality (“near the Magdalena River”), below 100 m of elevation.

C. popayanensis Kunth (J.J. Triana 3816/3, COL). Region III. Only a single (historic) collection from Andean forests of the department of Cauca, at elevations around 1800 m. Endemic.

9 Lamiales: One family.

Orobanchaceae Vent. Eight genera, 37 species (Figs. 21, 22, 23, 24, and 25).

Fig. 21
figure 21

Map of distribution of Anisantherina and Buchnera (Orobanchaceae) in Colombia

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Fig. 22
figure 22

Orobanchaceae, facultative root hemiparasites. A. Castilleja arvensis (Boyacá: Villa de Leyva, 2460 m.a.s.l., F. González 4762). B. C. fissifolia (Cundinamarca: Guatavita, 2930 m.a.s.l., F. González 301), lobed leaves, bracts and flowers. C–E. C. integrifolia, details of bracts and flowers at lateral (C, D) and adaxial (E) view (Boyacá: Toca, 3120 m.a.s.l., F. González 4732). F–J. C. paramensis. F–G. Collection site at paramo cushion bogs (Boyacá: Sogamoso, 3875 m.a.s.l., F. González et al. 4478). H. Lobed leaves, bracts and flowers. I. Underground stolon (arrow) attached to the taproot of Plantago rigida Kunth. J. Plastome (from Fan et al. 2016). K. Lamourouxia virgata (Nariño: Túquerres, 3000 m.a.s.l., F. González et al. 4339). L–M. Neobartsia santolinifolia (Boyacá: Chiscas, 4020 m.a.s.l., F. González et al. 4751). N–S. Orobanche minor (Boyacá: Monguí, 3400 m.a.s.l., N. Pabón-Mora et al. 491). N. Habit. O. Underground stolon. P–R. Young (P) and fully formed (Q, R) racemes. S. Capsule. Scale bars: 5 mm in A-E, H, I, P–R; 1 cm in K–O; 1 mm in S

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Fig. 23
figure 23

Map of distribution of Escobedia and Lamourouxia (Orobanchaceae) in Colombia

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Fig. 24
figure 24

Map of distribution of Neobartsia (Orobanchaceae, first part) in Colombia. The widely distributed N. laniflora, N. pedicularoides, N. ramosa, and N. santolinifolia are not including in the map

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Fig. 25
figure 25

Map of distribution of Neobartsia (Orobanchaceae, second part), Orobanche, and Pedicularis (Orobanchaceae) in Colombia. The widely distributed N. laniflora, N. pedicularoides, N. ramosa, and N. santolinifolia not included in map

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Anisantherina Pennell. One facultative, root hemiparasitic species.

A. hispidula (Mart.) Pennell (C. Allen 647, MO). Region I. Xerophytic thickets and savannas, at elevations below 150 m.

Buchnera L. Eleven facultative, root hemiparasitic species, two of them endemic to Colombia.

B. jacoborum Fern.Alonso (R. Jaramillo 1297, holotype COL). Regions IV and V. Savannas and margins of gallery forests, at elevations between 250 and 700 m. Endemic.

B. longifolia Kunth (H. Daniel 500, COL). Sub-Andean forests (Region III), and savanna vegetation (Region IV), at elevations between 150 and 1900 m.

B. obliqua Benth. (P. Pinto 1766, COL). Region III. Andean forests of the Western Cordillera, at elevations between 2200 and 2300 m.

B. palustris (Aubl.) Spreng. (C. Sastre 1531, COL). Region IV. Savanna vegetation, at elevations between 120 and 400 m.

B. pusilla Kunth (J.M. Idrobo 192, COL) Regions I and III. Savannas and paddocks, at elevations between 100 and 1900 m.

B. rosea Kunth (F. González et al. 3088, COL). Regions III and IV. Savannas and paddocks, at elevations between 100 and 1500 m.

B. schultesii R.Bernal (R.E. Schultes & I. Cabrera 14211, holotype A). Region V. Borders of gallery forests and clearings in Amazon forests, at elevations around 450 m. Endemic.

B. spruceana Philcox (F. González et al. 2294, COL). Region IV. Savanna vegetation, at elevations around 200 m.

B. ternifolia Kunth (O. Haught 2369, COL). Region III. Roadsides and paddocks, at elevations around 500 m.

B. virgata Kunth (D. Rivera 1297, COL). Regions IV and V. Borders of gallery forests, savanna vegetation, and clearings in Amazon forests, at elevations between 100 and 550 m.

B. weberbaueri Diels (P. Pinto 1064, COL). Region IV. Savannas and borders of gallery forests, at elevations between 90 and 650 m.

Castilleja Mutis ex L.f. Eight facultative, root hemiparasitic species, one of them endemic to Colombia (Fig. 22A–J).

C. arvensis Cham. & Schltdl. (F. González 4762, COL). Regions III and IV. Roadsides, paddocks and grassland vegetation, at elevations between 800 and 3700 m. (Fig. 22A).

C. breviflora Benth. (J. Cuatrecasas 24600, COL). Region I (Sierra Nevada de Santa Marta). Páramos and high-páramos at elevations between 3750 and 4500 m.

C. fissifolia L.f. (F. González 301, COL). Regions I (Sierra Nevada de Santa Marta) and III. Roadsides, paddocks and grassland vegetation, and subpáramo, páramo and high-páramo vegetation at elevations between 1600 and 4350 m. (Fig. 22B).

C. integrifolia L.f. (F. González 4732, COL). Regions I (Sierra Nevada de Santa Marta) and III. Roadsides, paddocks and grassland vegetation, and subpáramo, páramo and high-páramo vegetation at elevations between 1650 and 4100 m. (Fig. 22C–E).

C. nubigena Kunth (R. Galindo 1332, COL). Region III. Borders of Andean and high-Andean forests, and subpáramo, páramo and high-páramo vegetation between 2900 and 4200 m.

C. paramensis F.González & Pabón-Mora (F. González et al. 4478, holotype COL). Region III. Páramo and high-páramo vegetation, at elevations between 3500 and 4500 m. Endemic. (Fig. 22F–J).

C. pumila Wedd. (H. Sturm 92, COL). Region III. Páramo and high-páramo vegetation, at elevations between 3900 and 4100 m.

C. virgata (Wedd.) Edwin (M. Schneider 1199, COL). Region III. High-páramo vegetation, at elevations around 4200 m.

Escobedia Ruiz & Pav. Four facultative, root hemiparasitic species, two of them endemic to Colombia. (Fig. 23).

E. brachydonta Pennell (E.P. Killip 7877, holotype PH). Region III. Roadsides and paddocks of the Western Cordillera, at elevations around 2500 m. Endemic.

E. grandiflora (L.f.) Kuntze (F. González et al. 3051, COL). Regions I-IV. Savannas, roadside, paddock, and grassland vegetation, at elevations between 150 and 2550 m.

E. parvifolia Pennell (E.P. Killip & A.C. Smith 17210, isotype NY). Region III. Known only from its type locality at the Páramo Rico/Páramo de Santurbán (Eastern Cordillera), at elevations between 3000 and 3600 m. Endemic.

E. scabrifolia Ruiz & Pav. (J.M. Idrobo & H. Weber 1348, COL). Two disjunct records at regions III (Cauca, at elevations between 1700 and 2000 m) and IV (Vichada, at elevations around 100 m).

Lamourouxia Kunth. One facultative, root hemiparasitic species (Figs. 22K and 23).

L. virgata Kunth (F. González & N. Pabón-Mora 4339, COL). Region III. Roadside, paddock and grassland vegetation, at elevations between 1100 and 3400 m.

Neobartsia Uribe-Convers & Tank. Ten facultative, root hemiparasitic species, one of them endemic to Colombia (Figs. 22L–M, 24, and 25).

N. glandulifera (Molau) Uribe-Convers & Tank (J.L. Fernández 7759, COL). Páramos of the Central and Eastern cordilleras (Region III), plus a few records at the Sierra Nevada de Santa Marta (Region I), at elevations between 3000 and 3900 m.

N. inaequalis (Benth.) Uribe-Convers & Tank (R. Guarín-M. 273, COL). Region III. Andean to high andean forests and páramos primarily from the Eastern Cordillera, at elevations between 1990 and 3560 m.

N. laniflora (Benth.) Uribe-Convers & Tank (L.E. Mora-Osejo 769A, COL). Páramos and high-páramos of the Central and Eastern cordilleras (Region III), plus a few isolate collections at the Sierra Nevada de Santa Marta, at elevations between 3000 and 4100 m.

N. laticrenata (Benth.) Uribe-Convers & Tank (L.E. Mora-Osejo 7539, COL). Region III. Restricted to the Andes of Nariño, at elevations around 4000 m.

N. matuy Malagón, Humberto Mend. & Uribe-Convers (H. Mendoza & E. Torres, 22013, isotype COL). Region III. Known only from its type locality in Cómbita (Boyacá), at 3528 m. Endemic.

N. orthocarpiflora (Benth.) Uribe-Convers & Tank (H.G. Barclay 4532, COL). Region III. Páramos and high-páramos of Cundinamarca, Risaralda and Nariño, at elevations between 3000 and 3700 m.

N. pedicularoides (Benth.) Uribe-Convers & Tank (J.L. Fernández 6687, COL). Páramos and high-páramos of all three cordilleras (Region III), and the Sierra Nevada de Santa Marta (Region I), at elevations between 3100 and 4250 m.

N. ramosa (Molau) Uribe-Convers & Tank (J.L. Fernández 11627, COL). Region III. Páramos and high-páramos of the Central and Eastern cordilleras, at elevations between 3150 and 4000 m.

N. santolinifolia (Kunth) Uribe-Convers & Tank (F. González et al. 4751, COL). Region III. Páramos and high-páramos of the Eastern Cordillera, plus a few isolated records in the Central Cordillera (Antioquia, Cauca, Quindío and Nariño), at elevations between 2800 and 4400 m.

N. stricta (Kunth) Uribe-Convers & Tank (J.H. Torres et al. 1325, COL). Region III. Paramos and high-paramos of the Eastern and Central cordilleras, at elevations between 3130 and 4500 m.

Orobanche L. One facultative, root hemiparasitic species (Figs. 22N–S and 25).

O. minor Sm. (N. Pabón-Mora & F. González 491, HUA). Region III. Restricted to the Eastern Cordillera, at elevations between 2500 and 3400 m. Likely a recent introduction to Colombia, where it has been collected in a few paddocks of Boyacá and Cundinamarca. So far, this species represents the only parasitic angiosperm in Colombia that could become a potential weed.

Pedicularis L. One species (Fig. 25).

P. incurva Benth. (R. Jaramillo 5876, COL). Region III. Subpáramos to high-páramos of the Western and Central cordilleras, at elevations between 2700 and 4300 m.

Discussion

Regional Distribution of Parasitic Angiosperms in Colombia

Region I: A total of 58 species of parasitic flowering plants are found in Region I. Restricted to this region are: (a) All members of the families Cytinaceae (Sanguisuga caesarea; endemic) and Krameriaceae (two Krameria species) and the genera Acanthosyris (Cervantesiaceae; one species) and Lennoa (Ehretiaceae; one species); (b) the Santalalean Aetanthus ovalis (endemic), Maracanthus chlamydatus, Oyctanthus grammatus (endemic), Psittacanthus roldanii, P. smithii (endemic) and Struthanthus marginatus (Loranthaceae); (c) Agonandra brasiliensis (Opiliaceae), Phoradendron applanatum, P. buritacanum (endemic), P. exiguum, P. fonsecanum (endemic), P. longissimum (endemic), P. loriflorum, P. pellucidum and P. robaloense (Viscaceae); (d) Cuscuta colombiana and C. partita (Convolvulaceae); and (e) Anisanthera hispidula and Castilleja breviflora (Orobanchaceae). A total of seven species from three families are endemic to Region I.

Conservation and restoration of the coastal vegetation and the dry forests, two highly disturbed ecosystems of the Caribbean region, are urgently needed to preserve the above mentioned taxa in Colombia.

Region II: A total of 35 species of parasitic flowering plants are found in Region II. Seven species of Loranthaceae found in this region (Oryctanthus grandis, O, neurophyllus, Passovia cordata, Peristethium colombianum, P. lamprophyllus, Psittacanthus gigas, and P. micrantherus) are endemic to Colombia. Two additional species of Viscaceae (Phoradendron madisonii and P. triflorum) are restricted to this region.

Region III: A total of 169 species of parasitic flowering plants are found in Region III, being by far the natural region of Colombia with the highest number of parasitic angiosperms. The following taxa are restricted to Region III: (a) The two species of Pilostyles (Apodanthaceae); (b) all the species of Corynaea, Langsdorffia, Ombrophytum, and Scybalium (Balanophoraceae); (c) the two species of Cervantesia (Cervantesiaceae); (d) most Aetanthus spp. as well as Oryctanthus perticus (endemic), Passovia robusta, Peristethium aequatoris, Phthirusa exilis (endemic), P. hutchisonii, P. rhynchophylla (endemic), Psittacanthus antioquiensis (endemic), P. carinatus (endemic), P. complanatus (endemic), P. crassinervis (endemic), P. dilatatus, P. longerectus (endemic), P. microphyllus (endemic), P. pentaphyllus (endemic), P. rotundatus (endemic), P. ternatus, Struthanthus calophyllus, S. subtilis, Tripodanthus belmirensis (endemic), and Tristerix longibracteatus and T. secundus (endemic) (Loranthaceae); (e) the three species of Santalaceae; (f) most species of Dendrophthora and Phoradendron (Viscaceae); (g) the sole species of Mitrastemonaceae; (h) most Cuscuta (Convolvulaceae) species; and (i) Buchnera obliqua, B. ternifolia, most Castilleja, Escobedia and Neobartsia species, Lamourouxia virgata, Orobanche minor and Pedicularis incurva (Orobanchaceae).

The Region contains 14 endemic species. Conservation and restoration of cloud, sub-Andean, Andean and high-Andean forests and páramo ecosystems, some of them severely disturbed and fragmented, are urgently needed to preserve all these taxa in Colombia.

Region IV: A total of 29 species of parasitic flowering plants are found in Region IV. The following species are restricted to this region in Colombia: (a) The Santalalean Cladocolea coriacea (Loranthaceae), and Agonandra silvatica (Opiliaceae); and (b) Buchnera palustris, B. spruceana and B. weberbaueri (Orobanchaceae). No endemic species are present in this region.

Region V: A total of 54 species of parasitic flowering plants are found in Region V, and 27 of them are restricted to this region: (a) the mistletoes Oryctanthus phthirusoides, Psittacanthus biternatus, P. bolbocephalus, P. cinctus, P. clusiifolius, P. corderoi (endemic), P. crassifolius, P. eucalyptifolius, P. geniculatus, P. irwinii, P. ophiocephalus (endemic), P. robustus, P. schultesii (endemic), P. stergiosii, P. sulcatus (endemic), and Struthanthus schultesii (endemic) (Loranthaceae); (b) Agonandra peruviana (Opiliaceae); (c) The sole species of Tetrastylidium (T. peruvianum, Strombosiaceae); (d) Phoradendron apertiflorum, P. laxiflorum, P. linearifolium, P. mairaryense, P. palaephyllum, P. racemosum, P. singulare, P. strongyloclados (Viscaceae); and (e) Buchnera schultesii (endemic)(Orobanchaceae). Six of these species are endemic to Colombia.

Elevational Distribution of Parasitic Flowering Plants in Colombia (Figs. 26, 27, 28, and 29)

Fig. 26
figure 26

Elevational range of genera of Loranthaceae with eight or more species in Colombia. A. Aetanthus. B. Oryctanthus. C. Psittacanthus. Purple bars correspond to endemic species to the country

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Fig. 27
figure 27

Elevational range of Viscaceae with eight or more species in Colombia. A. Dendrophthora. B. Phoradendron. Purple bars correspond to endemic species to the country

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Fig. 28
figure 28

Elevational range of Convolvulaceae (A) and Orobanchaceae (B-D) with eight or more species in Colombia. A. Cuscuta. B. Buchnera. C. Castilleja. D. Neobartsia. Purple bars correspond to endemic species to the country

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Fig. 29
figure 29

Elevational range of the parasitic Angiosperms endemic to Colombia. Color coding: blue, holoendoparasite; green, facultative hemiparasites; purple, obligate hemiparasites; ochre, holoexoparasites

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Taxa exclusive to lowlands (0–1000 m): A total of 84 parasitic flowering plants (including 24 endemic species) are restricted to lowland ecosystems in Colombia. These are: Cassytha filiformis (Lauraceae); the two Krameria spp. (Krameriaceae); Sanguisuga caesarea (Cytinaceae; endemic); Acanthosyris glabrata (Cervantesiaceae); Cladocolea coriacea, Maracanthus chlamydatus, Oryctanthus grammatus (endemic), O. neurophyllus (endemic), O. phthirusoides (Fig. 26B), Passovia cordata (endemic), Peristethium colombianum (endemic), P. lamprophyllum (endemic), Phthirusa exilis (endemic), Psittacanthus biternatus, P. bolbocephalus, P. cinctus, P. clusiifolius, P. corderoi (endemic), P. crassicostatus (endemic), P. crassifolius, P. eucalyptifolius, P. geniculatus, P. gigas (endemic), P. irwinii, P. krameri, P. micrantherus (endemic), P. ophiocephalus (endemic), P. rhynchanthus, P. robustus, P. roldanii, P. schultesii (endemic), P. smithii (endemic), P. stergiosii, P. sulcatus (endemic) (Fig. 26C), Struthanthus marginatus, S. phillyreoides and S. schultesii (endemic) (Loranthaceae); all three Agonandra spp. (Opiliaceae); the two Schoepfia spp. (Schoepfiaceae); Tetrastylidium peruvianum (Strombosiaceae); Phoradendron apertiflorum (endemic), P. applanatum, P. crassicarpum (endemic), P. eggersii, P. exiguum, P. fasciculatum, P. fonsecanum (endemic), P. haughtii, P. laxiflorum, P. linearifolium, P. longissimum (endemic), P. lorifolium, P. madisonii, P. mairaryense, P. mucronatum, P. nitens, P. palaephyllum (endemic), P. pellucidulum, P. platycaulon, P. racemosum, P. robaloense, P. singulare, P. strongyloclados, P. tardispicum, P. triflorum and P. trinervium (Viscaceae) (Fig. 27B); Ximenia americana (Ximeniaceae); Lennoa madreporoides (Ehretiaceae); Cuscuta colombiana (endemic), C. partita, C. pentagona (Convolvulaceae) (Fig. 28A); and Anisanthera hispidula, Buchnera jacoborum (endemic), B. palustris, B. schultesii (endemic), B. spruceana, B. ternifolia, B. virgata and B. weberbaueri (Orobanchaceae) (Fig. 28B).

Taxa exclusive to mid-elevations (1000–2800 m): A total of 59 parasitic flowering plants (including 29 endemic species) are restricted to this elevational belt in Colombia. These are: Pilostyles boyacensis (endemic), and P. cf. blanchetii (Apodanthaceae); Langsdorffia hypogaea, Ombrophytum villamariensis (endemic) and Scybalium depressum (Balanophoraceae); Cervantesia macrocarpa (endemic) (Cervantesiaceae); Aetanthus alternifolius (endemic), A. nodosus, A. ovalis (endemic), A. sessilifolius (endemic), A. trifolius, A. validus (endemic) (Fig. 26A), Oryctanthus asplundii, O. perticus (endemic) (Fig. 26B), Passovia robusta, Peristethium aequatoris, Phthirusa hutchisonii, P. rhynchophylla (endemic), Psittacanthus antioquiensis (endemic), P. crassinervis (endemic), P. dilatatus, P. longerectus (endemic), P. pentaphyllos (endemic), P. rotundatus (endemic), P. ternatus (Fig. 26C), Struthanthus calophyllus, S. subtilis and S. syringifolius (Loranthaceae); Dendrophthora amalfiensis (endemic), D. fendleriana, D. macbridei, D. stricta (endemic), D. sulcata (endemic) (Fig. 27A), Phoradendron anceps, P. antioquianum (endemic), P. balslevii, P. brittonianum, P. buritacanum (endemic), P. dipterum, P. gracilicarpum (endemic), P. heterostachyum (endemic), P. killipii (endemic), P. megaphyllum (endemic), P. planiphyllum, P. restrepoae (endemic), P. roldanii (endemic), P. rugosum, P. squamiferum (endemic), P. staphylinum, P. uniseriale (endemic) and P. wiensii (Viscaceae) (Fig. 27B); Mitrastemon matudae (Mitrastemonaceae); Cuscuta corymbosa, C. indecora and C. popayanensis (endemic) (Convolvulaceae; Fig. 28A); and Buchnera obliqua, Escobedia brachydonta (endemic) (Fig. 28B), and Orobanche minor (Orobanchaceae).

Taxa exclusive to subpáramo, páramo and high-páramo ecosystems (2800–4500 m). A total of 34 parasitic flowering plants (incuding 11 endemic species) are restricted to this elevational belt in Colombia. These are: Cervantesia tomentosa (Cervantesiaceae); Aetanthus megaphyllus (endemic), A. mutisii (endemic) (Fig. 26A), Psittacanthus carinatus (endemic), P. complanatus (endemic), P. microphyllus (endemic) (Fig. 26C), Tripodanthus belmirensis (endemic), Tristerix longibracteatus, and T. secundus (endemic) (Loranthaceae); Antidaphne andina (Santalaceae); Dendrophthora ambigua, D. chrysostachya, D. elegantissima, D. lueri, D. squamigera, D. subtrinervia, D. tenuifolia, D. thomasii and Phoradendron retrophyllum (endemic)(Viscaceae; Fig. 27); and Castilleja breviflora, C. nubigena, C. paramensis (endemic), C. pumila, C. virgata (Fig. 28C), Escobedia parvifolia (endemic), and nine Neobartsia spp. (Orobanchaceae; Fig. 28D), one of them (N. matuy) endemic to the Eastern Cordillera of Colombia.

Elevational Gradients and Speciation

The elevational gradients for parasitic Angiosperm genera with eight or more species in Colombia suggest that the Andean orogeny has played a role in their diversification. These are:

Aetanthus (Loranthaceae): Andean to high-Andean forest remnants above 2000 m and paramos are the preferred habitat for all the species of Aetanthus. Most of the species (four out of ten) live in cloud forest remnants between 2700 and 3200 m (Fig. 26A). Three endemic species (A. colombianus, A. megaphyllus, and A. mutisii) converge at elevations between 3000 and 3100 m, whereas four endemic Aetanthus species (A. alternifolius, A. megaphyllus, A. ovalis and A. sessilifolius) are elevational disjuncts (Fig. 26A).

Oryctanthus (Loranthaceae): Lowland forests between 200 and 400 m hold the highest (nine out of 11) number of species of this genus in Colombia, including the rainforest endemics O. grammatus (below 300 m), O. grandis (between 10 and 1500 m), and O. neurophyllus (between 50 and 500 m). The fourth endemic species (O. perticus), which grows in sub-Andean forests between 1000 and 1500 m, is an elevational disjunct with respect to O. grammatus and O. neurophyllus (Fig. 26B).

Psittacanthus: Lowland rainforests between the sea level and 250 m hold the highest (21 out of 37) number of species of this genus in Colombia, including seven endemics (P. corderoi, P. gigas, P. micrantherus, P. ophiocephalus, P. schultesii, P. smithii and P. sulcatus). Disjunct endemics grow in sub-Andean (P. antioquensis, P. crassicostatus, P. crassinervis, P. pentaphyllos and P. rotundatus), Andean (P. longerectus) or high-Andean (P. carinatus) forests. The only endemic Psittacanthus species in Colombia restricted to the páramo ecosystem are P. complanatus and P. microphyllus (Fig. 26C).

Dendrophthora (Viscaceae): High-Andean forests and páramos between 3000 and 3400 m, hold the highest (13 out of 22) number of species of this genus in Colombia. Interestingly, the three endemic species of Dendrophthora occupy lower elevations at sub-Andean and Andean forests; these are D. amalfiensis (between 1300 and 1900 m), D. stricta (c 2200 m), and D. sulcata (between 1450 and 1840 m) (Fig. 27A).

Phoradendron (Viscaceae): The elevational belt between the sea level and 500 m holds the highest (36 out of 66) number of species of this genus in Colombia, including four endemic to the country (P. apertiflorum, P. fonsecanum, P. longissimum and P. palaephyllum). These four species are elevational disjuncts with the remaining endemic species of the genus. Only P. retrophyllum appears as a high Andean disjunct at elevations above 3000 m (Fig. 27B).

Cuscuta (Convolvulaceae): The elevational distribution for Cuscuta in Colombia shows no clearcut trend for its diversification. Whereas C. colombiana, C. partita and C. pentagona are found exclusively below 300 m, C. corymbosa and C. popayanensis are disjuncts restricted to elevations above 1700 m. Altogether, the remaining species (C. americana, C. grandiflora, C. indecorosa and C. obtusifolia) occupy a broader elevational belt between 0 and 4000 m, and only overlap between 500 and 1500 m (Fig. 28A).

Buchnera (Orobanchaceae): Lowland ecosystems between 400 and 500 m hold the highest (eight out of 11) number of species of this genus in Colombia, including the two endemic species to the country, namely, B. jacoborum (between 250 and 700 m) and B. schultesii (at around 450 m) (Fig. 28B).

Castilleja (Orobanchaceae): Páramo ecosystems between 3800 and 4100 m hold the highest (seven out of eight) number of species of this genus in Colombia, including the only endemic species to the country, namely, C. paramensis, which grows exclusively in peat bogs between 3500 and 4500 m (Fig. 28C).

Neobartsia (Orobanchaceae): All ten species of this genus in Colombia occupy Páramo ecosystems above 3000 m, including the only endemic species to the country, N. matuy, which grows at elevations c 3528 m (Fig. 28D).

Elevational Ranges of Endemic Parasitic Angiosperms in Colombia (Fig. 29)

Out of the parasitic Angiosperm species endemic to Colombia, 27 of them (39.13%) grow at elevations below 1000 m, 23 (33.33%) are found between 1000 and 2000 m, 20 (28.98%) grow between 2000 and 3000 m, 12 (17.39%) occupy páramos between 3000 and 4000 m, and only one species (1.44%) reaches superpáramos above 4000 m in the country. This account strongly suggests that the number of endemic parasitic angiosperms in Colombia is inversely proportional to the elevation, regardless the type of parasitic lifeform (Fig. 29).

Lifeforms of Parasitic Flowering Plants in Colombia

The majority of parasitic flowering plants present in Colombia are stem obligate hemiparasites with various forms of haustoria, and belong to the Loranthaceae, Santalaceae and Viscaceae (Santalales; Figs. 1, 10, 13, 17, and 18). The root facultative hemiparasites, restricted to the Orobanchaceae (Figs. 1 and 22), are the second predominant lifeform. Conversely, the holoendoparasitic lifeform occurs only in Apodanthes and Pilostyles (Apodanthaceae, Cucurbitales; Figs. 1 and 4). In the remaining orders, four convergent lifeforms occur recurrently, as follows:

  1. 1.

    The twining stem holoparasitic Cassytha (Lauraceae) and Cuscuta (Convolvulaceae), both developing an adhesive disk. This remarkable example of convergent evolution has been extensively discussed in the literature. Notably, Heide-Jørgensen (1991) highlighted the most significant developmental and structural similarities of this organ in these two distantly related genera, including: the epidermal and cortical development of the adhesive disk; the trichomatose formation of the adhesive epithelium; the polysaccharide and pectinaceous nature of the sticky secretion; the development of the endophyte from the middle layers of the cortex; the collapsed parenchymatic zones of the twining stem; and the lack of Santalalean glandular tissue. Reproductive structures in both genera consist of small axillary flowers or few-flowered cymes scattered in the distalmost portions of the twigs.

  2. 2.

    The root holoexoparasitism has evolved independently in Sanguisuga (Cytinaceae), Mitrastemon (Mitrastemonaceae), all Balanophoraceae (Santalales), and Lennoa (Ehretiaceae). Convergent morphological traits include thickened underground runners, and massive, thick axes, protective scale bracts, racemose inflorescences, minute flowers with bright white, pink, or reddish perianth. Detail accounts exist on the structural connection of the Balanophoraceae species with their host roots (Kuijt & Bruns, 1987; Pellissari et al., 2022), but in general, the haustorium in this family consists of underground tuber-like structures. The vascular connection between parasite and hosts can range from simple (when the vessel elements from both species make contact) to complex (when parasitic cells reach the core of the host´s conducting tissue). The emerging body portion consists of showy but ephemeral reproductive structures either as individual flowers or as massive, racemose inflorescences aboveground. Whereas the former three taxa possess small but highly elaborate flowers and accessory extrafloral structures, Lennoa displays intricate branched inflorescences, flowers with a showy, rotaceous perianth, and proliferative carpels (Fig. 19A–E).

  3. 3.

    The root obligate hemiparasitism has evolved independently in Krameria (Krameriaceae), Gaiadendron (Loranthaceae), and all Cervantesiaceae, Opiliaceae, Schoepfiaceae, Strombosiaceae and Ximeniaceae. This lifeform is the least known in terms of development, structure and ecology, likely due to the fact that most species are medium-sized to large trees.

Colombian Species Belonging to Parasitic Orders with Unconfirmed Parasitic Lifeforms

The following species native to Colombia belong to predominantly parasitic orders but require field studies and direct verification of their purported parasitic lifeforms:

Zygophyllales:

Zygophyllaceae:

Guaiacum officinale L., from dry forests of the Caribbean plains (Region I) and the Cauca valley (Region III), at elevations below 1000 m.

Kallstroemia maxima (L.) Hook. & Arn., K. parviflora Norton, and K. pubescens (G. Don) Dandy, all from dry forests and thickets in regions I and III, at elevations below 1700 m.

Plectocarpa arborea (Jacq.) Christenh. & Byng, and P. carrapo (Killip & Dugand) Christenh. & Byng, both from dry to wet forests of the Cauca and Magdalena valleys, in regions I and II, at elevations below 1200 m.

Santalales

Aptandraceae Miers (1853).

Aptandra caudata A.H. Gentry & R. Ortiz; A. tubicina (Poepp.) Benth. ex Miers. Region III, at elevations between 400 and 900 m.

Cathedra acuminata Miers.

Chaunochiton angustifolium Sleumer; C. kappleri (Sagot ex Engl.) Ducke, C. loranthoides Benth. Regions II and IV, at elevations below 600 m.

Coulaceae Tiegh.:

Minquartia guianensis Aubl. Regions II, III and V, at elevations below 1000 m.

Erythropalaceae:

Heisteria acuminata (Bonpl.) Engl., H. asplundii Sleumer, H. barbata Cuatrec., H. concinna Standl., H. duckei Sleumer, H. insculpta Sleumer, H. latifolia Standl., H. maytenoides Spruce ex Engl., H. nitida Engl., H. ovata Benth., H. pacifica P. Jørg. & C. Ulloa, H. scandens Ducke, and H. spruceana Engl. Regions II, III (Magdalena Valley) and V, at elevations below 500 m.

Olacaceae Martinov:

Olax candida (Poepp.) Christenh. & Byng; O. inopiflora (Miers) Christenh. & Byng; O. macrophylla Benth., and O. redmondii (Steyerm.) Christenh. & Byng., all at elevations between 50 and 500 m, in regions I (Magdalena valley), II, IV and V.

Ximeniaceae:

Curupira tefeensis G.A. Black. Amazon forests (Region V), at elevations below 200 m.

Concluding remarks

The majority of parasitic angiosperm species in Colombia is concentrated in regions I (Caribbean) and III (Andean), which currently have the highest deforestation areas (c 75% and 55%, respectively) in the country (IDEAM, 2022). Given the crucial roles of most of these species providing pollen, nectar and fruits to the fauna, we call for urgent plans of conservation, recovery and protection of the natural reserves still in place on those ecosystems. In general, the presence of species of Balanophoraceae, Apodanthaceae, Cytinaceae, Loranthaceae (especially Aetanthus, Oryctanthus, Psittacanthus, and Tristerix), and Mitrastemonaceae in dry forests as well as cloud or rainforests, and endemic Orobanchaceae from paramo peat bogs serve as indicators of healthy environmental conditions and little disturbance in their ecosystems. At present, we do not detect any large-scale threat caused by any parasitic flowering plant species to the native flora or to the crops in the country. However, small-flowered Loranthaceae and species of Viscaceae can rapidly colonize fruit or ornamental trees in human disturbed environments, and awareness needs to be raised to control the excessive proliferation of these plants.