Abstract
Shifts between pollinator functional groups can explain major changes in floral phenotype. I document a novel case of butterfly pollination in Platycoryne, an African genus that is phylogenetically embedded in the very large Habenaria clade in the Orchidaceae. Most Habenaria species have green or white flowers and many of these have been shown to be pollinated by moths, but my observations of the orange-flowered species Platycorynus mediocris in south-central Africa showed that it is pollinated diurnally by butterflies. The nectar-producing spurs of this species are c. 15 mm in length and closely match the tongue lengths of nymphalid and pierid butterflies that visit the flowers. The rostellum arms flank the spur entrance and place sickle-shaped pollinaria on the eyes or palps of the butterflies. In contrast to the highly scented flowers of moth-pollinated Habenaria species, the flowers of P. mediocris emit very little scent. Anecdotal observations indicate that several other Platycoryne species with orange flowers are also pollinated by butterflies. I conclude that the flower colouration (orange without UV reflectance) and low emission of scent in P. mediocris reflect an important historical shift to butterfly pollination in African members of the Habenaria clade.
Similar content being viewed by others
Avoid common mistakes on your manuscript.
Introduction
Evolutionary shifts among pollinator functional groups are common in plant lineages and can have major implications for the evolution of floral phenotypes (van der Niet and Johnson 2012). Identifying these shifts and their implications for trait evolution requires a combination of natural history observations, phylogenetics and studies of floral morphology, advertising and reward traits. Shifts involving moths and butterflies are of interest because these two insect groups are closely related and similar morphologically, yet have differences in sensory modalities and diel activity patterns that have major implications for the evolution of floral advertising traits. As a generalization, moth-pollinated flowers tend to be pale and scented at night, while butterfly-pollinated flowers tend to be brightly coloured and are not usually strongly scented (Faegri and van der Pijl 1979; Goldblatt and Manning 2002; Liu et al. 2022). However, moths and butterflies are both highly diverse groups, and within each group, there is considerable variation in traits such as proboscis length, visual systems and behaviour; for example, whether species settle or flutter while feeding (Mertens et al. 2021). Plants also exploit different body parts of these insects for pollen transport (Hapeman and Inoue 1997; Butler and Johnson 2020). As a consequence, there is marked variation in the floral traits of plants pollinated by Lepidoptera (Mertens et al. 2021).
Habenaria is one of the largest orchid genera worldwide and is known to be paraphyletic with several currently recognized genera being embedded within the broader Habenaria clade (Ngugi et al. 2020). One example is Platycoryne, an African genus of about 20 species, that has been shown to be phylogenetically nested within Habenaria (Ngugi et al. 2020). Earlier morphological studies also supported the inclusion of Platycoryne in Habenaria (Kurzweil and Weber 1992). The majority of Habenaria species have white or green flowers, and many of these species have been shown to be moth-pollinated. It is notable that the flowers of most Platycoryne species are orange. This trait is very rare in Habenaria and has been linked positively to butterfly pollination in the well-studied Asian species Habenaria rhodocheila (Chen et al. 2021; Zhang et al. 2021). Similarly in the orchid genus Platanthera which is not part of the Habenaria clade, moth-pollinated species tend to have white flowers (Nilsson 1978), whereas butterfly-pollinated species tend to have orange flowers (Robertson and Wyatt 1990; Hapeman and Inoue 1997). Orange flower colour is also strongly linked with butterfly pollination in other plant families (Goldblatt and Manning 2002; Kiepiel and Johnson 2014; Daniels et al. 2020; Liu et al. 2022). However, orange flowers also occur in some bird-pollinated orchids (Johnson and Van der Niet 2019) and butterflies have been shown to pollinate white-flowered species in the Habenaria clade (Pedron et al. 2012; Balducci et al. 2019b; Tan et al. 2023). It is therefore essential to conduct field-based pollinator observations and not to rely solely on floral syndromes for reaching conclusions about shifts in pollination systems (van der Niet 2021).
I investigated the pollination biology of the orange-flowered species Platycoryne mediocris in south-central Africa and also searched for anecdotal records of insect visitors to flowers of other Platycoryne species to test my hypothesis that the evolution of orange flowers in this clade of Habenaria is linked with pollination by butterflies. The aims of this study were 1) identify pollinators of P. mediocris, 2) to document floral morphology, nectar rewards and advertising traits such as colour and scent, and 3) to determine levels of pollination success and fruit set.
Methods
Study species
Platycoryne mediocris Summerh. has a wide distribution in south-central Africa that encompasses Zambia, Zimbabwe, northern South Africa, Malawi, the Democratic Republic of Congo, Burundi and Tanzania. All parts of the flower including the ovary are bright orange (Fig. 1). Plants are 15–50 cm tall, and in the population that was studied, plants produced a mean (± SE) of 4.11 ± 0.10 flowers (n = 26).
Study site
Observations and trait measurements were carried out at Mutinondo Wilderness (12°27′18"S, 31°17′28"E) in central Zambia in 2022 and 2023. The study population of P. mediocris consisted of about 200 plants growing on a large granite inselbergs. A general account of the flora at the study site is given by Vollesen and Merrett (2020). Voucher specimens (Bytebier and Wightman 3819) are deposited in the Bews Herbarium (NU), the Kitwe Forest Herbarium (NDO) and Meise Botanic Garden Herbarium (BR).
Pollinator observations
Pollinator observations were carried out from 09h00 to 16h00 on 20–23 February 2022 and 7–11 February 2023. Plants were also observed intermittently in the evenings from 18h00 to 23h00, while I was conducting a separate study of moth visitors to flowers of Chamaepentas nobilis (Rubiaceae) which grows intermingled with P. mediocris at the study site. I also used four passive infrared (PIR) motion-activated cameras (Bushnell Nature View 119740) to make indirect observations of flower visitors over the same time period as the direct observations. Insects observed on the flowers were photographed for later identification, and a sample of each species was captured for morphological measurements (proboscis length and distance between inner and outer surfaces of the eyes) and confirmation of the identity of pollinaria they were carrying.
Floral traits
To assess traits that could influence the morphological fit between flowers and pollinators, I measured floral spur length, the length and width of the sepals and petals, the length and spacing of the rostellum arms, the stigma size, and the dimensions of pollinaria for one flower on each of 15 plants. I also measured the volume and concentration of nectar in these flowers using 5-µL pipettes and a 0–50% hand-held refractometer. Spectral reflectance of flowers was measured using an Ocean Optics S2000 spectrometer following the methods described by Johnson and Anderson (2002). Floral scent was analysed using headspace collection and gas chromatography–mass spectrometry (GC–MS) using the apparatus and methods described by Johnson et al (2020). I took four headspace samples from 12 plants (three plants per sample) by enclosing them in polyacetate bags (Kalle, Germany) and sucking air from the bags through an absorbent filter for 60 min at 200 ml per min. Compounds present in similar amounts in a control ambient sample were excluded from analysis. Emission rate of volatiles emitted was calculated from injection of a known amount of methyl benzoate as described by Johnson et al (2020).
Pollination and fruiting success
To assess the level of pollination success in the population, I examined 46 flowers on 24 plants and recorded the number of pollinaria removed and the number of pollen massulae deposited on stigmas. I counted the number of pollen massulae in pollinaria from eight flowers and used these data to calculate pollen transfer efficiency (the proportion of pollen removed from flowers that is deposited on stigmas). When examining flowers, I noted whether pollinia of wilted unvisited flowers remained enclosed within the anther as this would prevent autonomous self-pollination. I also calculated the percentage of flowers that set fruit for 26 plants sampled four weeks after the end of flowering.
Results
Pollinator observations
Flowers of P. mediocris were visited exclusively by medium-sized nymphalid and pierid butterflies (Fig. 2, Table 1, Video S1). I directly observed 28 butterflies feeding on flowers in the population, and a further two butterflies were recorded using camera traps (Video S1). In all cases, the butterflies probed the spur while hanging from the flower, using their feet to grip the ridged ovary or the reflexed lateral sepals. Six butterfly individuals, representing three species, were confirmed to carry pollinaria (Table 1). Pollinaria were mostly attached to the eyes of butterflies, but some were attached either to the palps or between the palps (Fig. 2). In one case, a pollinarium was attached to the legs of a butterfly. The pollinaria on butterflies were confirmed as those of P. mediocris on account of their length and distinctive sickle shape with the massulae segments lined up on one side of the caudicle. The pollinaria attached to butterflies curl forwards over the palps, such that the massulae segments are pressed against the stigma when the tongue is fully inserted. The mean (± SE) proboscis length of butterflies captured on the orchid was 13.1 ± 0.35 mm.
Butterfly visitors to flowers of Platycoryne mediocris (a–e) and Platycoryne sp. (f). a Male Acraea natalica probing a flower. b Acraea caldarena carrying pollinaria. c Female Acraea natalica carrying pollinaria. d Catopsilia florella probing a flower. e Mylothris rueppellii probing a flower. f Eurema hecabe carrying pollinaria on its eyes. Scale bars = 5 mm, except b = 2 mm. Arrows indicate pollinaria attached to the heads of insects. Photographs: Steve Johnson (a–e) and Craig Peter (f)
Floral traits
The mean (± SE) length of the floral spur of P. mediocris was 15.7 ± 0.25 mm which is slightly longer than the mean proboscis length of the butterfly flower visitors (t = 6.15, P < 0.001). The length of the rostellum arms was 1.41 ± 0.06 mm, and the distance between the rostellum arms was 1.8 ± 0.10 mm which is less than the distance between the mean (± SE) outer margins of the butterfly eyes (3.6 ± 0.21 mm), but greater than the distance between the inner margins of their eyes (1.46 ± 0.08 mm). Pollinaria were c. 3.7 ± 0.19 mm in length, with the caudicle measuring 1.7 ± 0.13 mm in length and the pollinium 2.0 ± 0.11 mm in length. Other floral trait dimensions are provided in Table S1.
The volume of the standing crop of nectar in flowers was 1.1 ± 0.19 µL with a sugar concentration of 22.8 ± 1.20%. The mean height of the standing crop nectar column from the tip of the spur was 4.4 ± 0.65 mm which means that it was accessible to butterflies with proboscides of 11.3 mm or longer.
Flowers of P. mediocris absorb ultraviolet light and show an increase in reflectance around 550 nm resulting in the orange appearance (Fig. 3).
Flowers have no discernible scent to humans. Analysis revealed only trace emissions of three compounds (mean percentage of total peak area ± SE): benzaldehyde (68.2 ± 6.8%), linalool (12.0 ± 2.2%), and phenylethyl alcohol (19.8 ± 8.4%). The mean (± SE) emission rate of volatiles was 0.55 ± 0.1 ng/flower/hour.
Pollination and fruiting success
Of the 46 flowers surveyed, 47.8% were pollinated with an average of 12.2 ± 3.1 massulae (range: 0–70) deposited per stigma, 69.6% had at least one pollinarium removed, and 63% of all pollinaria were removed. Each pollinarium contained 178.6 ± 5.4 massulae. Pollen transfer efficiency in the population was calculated as 4.9%. At the end of the flowering season, 88.1 ± 3.9% of the flowers on 26 plants set fruit.
Discussion
The results of this first detailed study of pollination in Platycoryne are consistent with the hypothesis that orange-flowered Platycoryne species are pollinated by butterflies. In addition to the observations of butterflies visiting P. mediocris that are reported here, I have seen Acraea butterflies visiting the orange flowers of P. buchananii in northern Zambia, and they have also been photographed on the orange flowers of P. guingangae in Gabon (Ramette 2023) and P. pervillei in Zimbabwe (Fibeck and Phiri 1998). Acraea natalica, the most common visitor of the flowers of P. mediocris (Table 1), was also the only recorded species to visit flowers and carry pollinaria of P. pervillei in Zimbabwe (Fibeck and Phiri 1998). A pierid butterfly Eurema hecabe has also been observed by C. Peter (pers. comm.) on the orange flowers of an unidentified Platycoryne species in Angola and also carried pollinaria on its eyes (Fig. 2F). The spur length of this orchid was c. 13 mm, closely matching the mean (± SE) proboscis length of three captured E. hecabe butterflies which was 12.1 ± 0.17 mm (C. Peter, pers. comm.).
The evolution of orange butterfly-pollinated flowers in Platycoryne in Africa parallels that of the independent evolution of orange flowers in the lineage leading to the Asian species Habenaria rhodocheila which is also pollinated by butterflies (Chen et al. 2021; Zhang et al. 2021). The importance of colour for attraction of butterflies has been demonstrated in numerous experiments (Chen et al. 2021; Kiepiel and Johnson 2021). There is a strong overall association between orange-red flowers and butterfly pollination in Africa (Newman et al. 2012; Kiepiel and Johnson 2014; Butler and Johnson 2020; Mertens et al. 2021). The butterfly Catopsilia florella which was one of the species observed to carry pollinaria of P. mediocris was also observed to visit the orange flowers of Gloriosa superba at the study site, and butterflies are known to visit flowers of G. superba across its extensive range in Africa (Daniels et al. 2020).
A notable result was the almost complete lack of scent emission by flowers of P. mediocris. The emission rate of 0.55 ng per flower per hour is at least 100-fold less than values reported for moth-pollinated species in the Habenaria clade—321–2551 ng/fl/hr for H. clavata (Johnson et al. 2020), 120 ng/fl/hr for Bonatea polypodantha (Balducci et al. 2020) and 152.5 ng/fl/hr for Bonatea steudneri (Balducci et al. 2019a). Other authors have reported only emission rates per inflorescence for moth-pollinated Habenaria species, but when divided by the reported average number of flowers, these give similarly high rates of emission—c. 4566 ng/fl/hr for Habenaria limprichtii (Tao et al. 2018) and c. 2083 ng/fl/hr for Habenaria epipactidea (Peter et al. 2009). Conversely, a low rate of 28.8 ng/fl/hr was reported for Bonatea cassidea, which is a butterfly-pollinated member of the Habenaria clade (Balducci et al. 2019b).
Pollinaria of P. mediocris were attached the eyes or palps of butterflies (Fig. 1B, C). This is also a common mode of pollinaria placement in the genus Habenaria and is associated with short diverging rostellum arms that flank the entrance to the spur (Singer and Cocucci 1997; Pedron et al. 2012; Zhang and Gao 2017; Tao et al. 2018). The pollinaria of P. mediocris are sickle-shaped and on withdrawal, project forward and slightly downwards such that the massula segments are pushed against the stigma. The high percentage of flowers of P. mediocris that were pollinated and set fruit is not unusual for orchids that produce nectar rewards (Tremblay et al. 2004). The values are also consistent with pollination rates of “50–80%” reported for a population of P. pervillei in Zimbabwe (Fibeck and Phiri 1998). The pollen transfer efficiency of 4.9% that was recorded in the study population is lower than the average of c. 10% for orchids with massulate (sectile) pollinia (Johnson and Harder 2023). This may reflect that many of the pollinaria attached to butterflies were out of alignment (Fig. 2B) which is likely due to physical interference when large number of pollinaria become attached to the same insect (Harder et al. 2021).
Although it is clearly apparent that a shift took place from moth to butterfly pollination in the Habenaria clade that includes Platycoryne, more detailed phylogenetic studies and pollinator observations are required to infer whether this transition was directly from moth to butterfly pollination or involved an intermediate stage with a different pollination system. There are some Platycoryne species (e.g. P. macroceras, P. isoetifolia) with pale white or greenish flowers which suggest that they may be moth-pollinated, and a more detailed phylogeny is therefore needed to reconstruct the pollination system of the original ancestor of the Platycoryne clade.
References
Balducci MG, Martins DJ, Johnson SD (2019a) Pollination of the long-spurred African terrestrial orchid Bonatea steudneri by long-tongued hawkmoths, notably Xanthopan morganii. Pl Syst Evol 305:765–775. https://doi.org/10.1007/s00606-019-01605-2
Balducci MG, Van der Niet T, Johnson SD (2019b) Butterfly pollination of Bonatea cassidea (Orchidaceae): solving a puzzle from the Darwin era. S African J Bot 123:308–316. https://doi.org/10.1016/j.sajb.2019.03.030
Balducci MG, Van der Niet T, Johnson SD (2020) Diel scent and nectar rhythms of an African orchid in relation to bimodal activity patterns of hawkmoth pollinators. Ann Bot (Oxford) 126:1155–1164. https://doi.org/10.1093/aob/mcaa132
Butler HC, Johnson SD (2020) Butterfly-wing pollination in Scadoxus and other South African Amaryllidaceae. Bot J Linn Soc 193:363–374. https://doi.org/10.1093/botlinnean/boaa016
Chen XH et al (2021) The pollination of Habenaria rhodocheila (Orchidaceae) in South China: when butterflies take sides. Ecol Evol 11:2849–2861. https://doi.org/10.1002/ece3.7242
Daniels RJ, Johnson SD, Peter CI (2020) Flower orientation in Gloriosa superba (Colchicaceae) promotes cross-pollination via butterfly wings. Ann Bot (Oxford) 125:1137–1149. https://doi.org/10.1093/aob/mcaa048
Faegri K, van der Pijl L (1979) The principles of pollination ecology. Pergamon, Oxford
Fibeck W, Phiri V (1998) Pollination of Platycoryne pervillei. S African Orchid J 29:11–12
Goldblatt P, Manning JC (2002) Evidence for moth and butterfly pollination in Gladiolus (Iridaceae-Crocoideae). Ann Missouri Bot Garden 89:110–124
Hapeman JR, Inoue K (1997) Plant-pollinator interactions and floral radiation in Platanthera (Orchidaceae). In: Givnish TJ, Sytsma KJ (eds) Molecular evolution and adaptive radiation. Cambridge University Press, Cambridge, pp 433–454
Harder LD, Richards SA, Agren J, Johnson SD (2021) Mechanisms of male-male interference during dispersal of orchid pollen. Amer Nat 197:250–265. https://doi.org/10.1086/712378
Johnson SD, Andersson S (2002) A simple field method for manipulating ultraviolet reflectance of flowers. Canad J Bot 80:1325–1328. https://doi.org/10.1139/b02-116
Johnson SD, Harder LD (2023) The economy of pollen dispersal in flowering plants. Proc Roy Soc B, Biol Sci 290:20231148. https://doi.org/10.1098/rspb.2023.1148
Johnson SD, Van der Niet T (2019) Bird pollination in an African Satyrium (Orchidaceae) confirmed by camera traps and selective exclusion experiments. Pl Syst Evol 305:477–484. https://doi.org/10.1007/s00606-019-01587-1
Johnson SD, Balducci MG, Shuttleworth A (2020) Hawkmoth pollination of the orchid Habenaria clavata: mechanical wing guides, floral scent and electroantennography. Biol J Linn Soc 129:213–226. https://doi.org/10.1093/biolinnean/blz165
Kiepiel I, Johnson SD (2014) Shift from bird to butterfly pollination in Clivia (Amaryllidaceae). Amer J Bot 101:190–200. https://doi.org/10.3732/ajb.1300363
Kiepiel I, Johnson SD (2021) Responses of butterflies to visual and olfactory signals of flowers of the bush lily Clivia miniata. Arthropod-Pl Interact 15:253–263. https://doi.org/10.1007/s11829-021-09813-9
Kurzweil H, Weber A (1992) Floral morphology of southern African Orchideae. II Habenariinae. Nordic J Bot 12:39–61
Liu C-Q et al (2022) Papilio butterfly vs. hawkmoth pollination explains floral syndrome dichotomy in a clade of Lilium. Bot J Linn Soc 199:678–693. https://doi.org/10.1093/botlinnean/boab074
Mertens JE et al (2021) Elevational and seasonal patterns of butterflies and hawkmoths in plant-pollinator networks in tropical rainforests of Mount Cameroon. Sci Rep 11:9710. https://doi.org/10.1038/s41598-021-89012-x
Newman E, Anderson B, Johnson SD (2012) Flower colour adaptation in a mimetic orchid. Proc Roy Soc B, Biol Sci 279:2309–2313. https://doi.org/10.1098/rspb.2011.2375
Ngugi G, Le Péchon T, Martos F, Pailler T, Bellstedt DU, Bytebier B (2020) Phylogenetic relationships amongst the African genera of subtribe Orchidinae s.l. (Orchidaceae; Orchideae): implications for subtribal and generic delimitations. Molec Phylog Evol 153:106946. https://doi.org/10.1016/j.ympev.2020.106946
Nilsson LA (1978) Pollination ecology and adaptation in Platanthera chlorantha (Orchidaceae). Bot Not 131:35–51
Pedron M, Buzatto CR, Singer RB, Batista JA, Moser A (2012) Pollination biology of four sympatric species of Habenaria (Orchidaceae: Orchidinae) from southern Brazil. Bot J Linn Soc 170:141–156. https://doi.org/10.1111/j.1095-8339.2012.01285.x
Peter CI et al (2009) Confirmation of hawkmoth pollination in Habenaria epipactidea: Leg placement of pollinaria and crepuscular scent emission. S African J Bot 75:744–750. https://doi.org/10.1016/j.sajb.2009.08.007
Ramette G (2023) Orchids from Gabon. Available at: http://orchideesgabon.e-monsite.com/. Accessed 24 Sep 2023
Robertson JL, Wyatt R (1990) Evidence for pollination ecotypes in the yellow-fringed orchid, Platanthera ciliaris. Evolution 44:121–133
Singer R, Cocucci AA (1997) Eye attached hemipollinaria in the hawkmoth and settling moth pollination of Habenaria (Orchidaceae): a study on functional morphology in 5 species from subtropical South America. Bot Acta 110:328–337. https://doi.org/10.1111/j.1438-8677.1997.tb00648.x
Tan S-L et al (2023) Swallowtail butterflies and hawkmoths contribute equally to the pollination of Habenaria dentata (Orchidaceae). Flora 300:152230. https://doi.org/10.1016/j.flora.2023.152230
Tao ZB et al (2018) Nocturnal hawkmoth and noctuid moth pollination of Habenaria limprichtii (Orchidaceae) in sub-alpine meadows of the Yulong snow mountain (Yunnan, China). Bot J Linn Soc 187:483–498. https://doi.org/10.1093/botlinnean/boy023
Tremblay RL, Ackerman JD, Zimmerman JK, Calvo RN (2005) Variation in sexual reproduction in orchids and its evolutionary consequences: a spasmodic journey to diversification. Biol J Linn Soc 84:1–54. https://doi.org/10.1111/j.1095-8312.2004.00400.x
van der Niet T (2021) Paucity of natural history data impedes phylogenetic analyses of pollinator-driven evolution. New Phytol 229:1201–1205. https://doi.org/10.1111/nph.16813
van der Niet T, Johnson SD (2012) Phylogenetic evidence for pollinator-driven diversification of angiosperms. Trends Ecol Evol 27:353–361. https://doi.org/10.1016/j.tree.2012.02.002
Vollesen K, Merrett L (2020) Field guide to the (Wetter) Zambian Miombo woodland part 1 and 2. GVPedia Communications, Zambia
Zhang WL, Gao JY (2017) Multiple factors contribute to reproductive isolation between two co-existing Habenaria species (Orchidaceae). PLoS ONE 12:e0188594. https://doi.org/10.1371/journal.pone.0188594
Zhang Z et al (2021) Pollination of the orchid Habenaria rhodocheila by the swallowtail butterfly Papilio helenus in subtropical evergreen broad-leaved forests in Southern China. Flora 274:151736. https://doi.org/10.1016/j.flora.2020.151736
Acknowledgements
Many thanks to Benny Bytebier and Nicholas Whiteman for assistance and for sharing their knowledge of the Zambian flora during the fieldwork and to Lari Merrett at Mutinondo Wilderness for counting fruits on the plants. I am also grateful to Colin Congdon and Steve Woodhall for assisting with identification of the butterfly species and to Craig Peter for sharing his observation and photographs of butterflies pollinating a Platycoryne species in Angola.
Funding
Open access funding provided by University of KwaZulu-Natal.
Author information
Authors and Affiliations
Contributions
SDJ collected and analysed the data, prepared the figures and wrote the paper.
Corresponding author
Ethics declarations
Conflict of interest
The author declares that there is no conflict of interest.
Additional information
Handling Editor: Vinãcius Brito.
Publisher's Note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Supplementary Information
Below is the link to the electronic supplementary material.
Supplementary file1 (MP4 9926 kb)
Information on Electronic Supplementary Material
Information on Electronic Supplementary Material
Online Resource 1. Video footage of a female Acraea natalica butterfly pollinating flowers of Platycoryne mediocris.
Online Resource 2. Table of floral traits measured in 15 plants of Platycoryne mediocris.
Rights and permissions
Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
About this article
Cite this article
Johnson, S.D. Butterfly pollination in Platycoryne (Orchidaceae): evidence for a key pollinator shift in the large Habenaria clade. Plant Syst Evol 310, 21 (2024). https://doi.org/10.1007/s00606-024-01895-1
Received:
Accepted:
Published:
DOI: https://doi.org/10.1007/s00606-024-01895-1