Biodiversity Data Journal 9: e62839 CO) doi: 10.3897/BDJ.9.e62839 open access Research Article A preliminary world checklist of fern-mining insects Jie Yang*, Xuexiong Wang, Kevin Jan Duffy§, Xiaohua Dai*! + Leafminer Group, School of Life Sciences, Gannan Normal University, Ganzhou, China § Institute of Systems Science, Durban University of Technology, Durban, South Africa | National Navel-Orange Engineering Research Center, Ganzhou, China Corresponding author: Xiaohua Dai (ecoinformatics@gmail.com) Academic editor: Torsten Dikow Received: 06 Jan 2021 | Accepted: 15 Mar 2021 | Published: 25 Mar 2021 Citation: Yang J, Wang X, Duffy KJ, Dai X (2021) A preliminary world checklist of fern-mining insects. Biodiversity Data Journal 9: e62839. https://doi.org/10.3897/BDJ.9.e62839 Abstract Compared to the leaf-miners and stem-miners on flowering plants, the miners on ferns (including both Lycopodiophyta and Polypodiophyta in the broad sense) are less known. Knowledge of miners and their host plants is essential to fully understand plant-insect interactions. Although there are many scattered records on fern miners, a worldwide checklist has not been reported. We provide a preliminary checklist of fern-mining insects and their host plants worldwide. Altogether, we found records for 128 species and 18 families of fern miners, mainly that feed on Dennstaedtiaceae, Equisetaceae, Polypodiaceae and Aspleniaceae. Fern miners belonged to four orders: Diptera (51 species; 39.8%), Coleoptera (33 species; 25.8%), Lepidoptera (28 species; 21.9%) and Hymenoptera (16 species; 12.5%). They are primarily known from the Palaearctic Region, Nearctic Region, Neotropical Region and Oriental Region. Keywords fern, leaf miner, host plant, plant-insect interactions, plant apparency hypothesis © Yang J et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 2 Yang J et al Introduction Leaf/stem miners are endophagous insects whose larvae feed on parenchyma or epidermal cells and form visually distinctive feeding tunnels, i.e. ‘mines’ on the leaves or stems (Dai et al. 2018, Eiseman 2020b, Liu et al. 2015, Sinclair and Hughes 2010). The mines can provide useful hints on insect species identities, insect life histories, insect behaviour and insect-plant interactions (Dai et al. 2018). Fewer miner groups (e.g. gracillariid moths, agromyzid flies and leaf-mining chrysomelids) can utilise over 100 plant families (Dai et al. 2017, De Prins and De Prins 2020, Santiago-Blay 2004, Spencer 1990), which are mainly angiosperm families, such as Fagaceae and Myrtaceae (Dai et al. 2018). Ferns (Pteridophyta, including both Lycopodiophyta and Polypodiophyta in the broad sense) are the second largest group of vascular plants, just after angiosperms (Dai et al. 2020, Schneider et al. 2004). With lower nutrition, higher defensive chemicals and no flowers, the interspecific associations between ferns and insects are often overlooked (Mehltreter et al. 2010, Weintraub et al. 1995). However, ferns used to be the primary food resource for insects before the thriving of angiosperms (Mehltreter et al. 2010). In fact, fossil records indicate that ferns and insects have co-evolved for at least 300 myr (Chandra and Srivastava 2003). Some ferns have nectaries and domatia, which could attract ants to be bodyguards (Mehltreter et al. 2010). Moreover, some insects mimic the soral crypsis of ferns to escape from their natural enemies (Barker et al. 2005, Patra et al. 2008). Some researchers hypothesise that fern-feeding insects should have fewer species, genera and families than those of seed plants (Weintraub et al. 1995), while others suggest that the richness of fern-feeding insects is largely underestimated (Auerbach and Hendrix 1980, Mehlitreter et al. 2010). The possible underestimation might be deduced from the following facts: (1) many fewer investigations have been performed for wild ferns than for cultivated ferns or invasive ferns (Fountain-Jones et al. 2012); (2) many more fern herbivores have been discovered in the comprehensive screening of bio-control agents for pest ferns (Mehltreter et al. 2010); (3) no noticeable difference has been found between leaf herbivory loss of ferns and that of seed plants (Chandra and Srivastava 2003); and (4) the possible biases of plant apparency (i.e. body size, distribution range and individual numbers (Dai et al. 2017)) are not considered for phytophagous insects on ferns in comparison to those on seed plants (Auerbach and Hendrix 1980). Fern-feeding insects could be classified as generalists and specialists. Most fern-feeding generalists tend to be classified as leaf-chewing or sap-sucking, while most specialists are classified as leaf-mining, gall-forming or spore-feeding (Mehltreter et al. 2010). By far, the miners on ferns are much less known than those on seed plants; although there are scattered records on publications and websites (De Prins and De Prins 2020, Edmunds et al. 2020, Ellis 2020, Eiseman 2020b, Pitkin et al. 2019, Santiago-Blay 2004, Spencer 1990), few comprehensive reviews on fern miners have been provided and a worldwide checklist has not been reported. In this study, we will compile a preliminary checklist of fern-mining insects and their host plants throughout the world, which could provide meaningful information to the study of plant-insect—-environment interactions. A preliminary world checklist of fern-mining insects 3 Material and methods The names and hosts of fern miners were obtained from websites, books and articles. Most publications were retrieved from the Web- of Science — (https:// www.webofknowledge.com) and Google Scholar (https://scholar.google.com), while the others were obtained from reference lists of the websites and retrieved publications. According to the Taxonomic Name Resolution Service (http://tnrs.iplantcollaborative.org/ TNRSapp.html), the host fern's scientific names were verified and corrected. The number of species in a fern family was obtained from the Catalogue of Life (http:// www.catalogueoflife.org/). Based on two recent mega-trees (Smith and Brown 2018, Zanne et al. 2014), 'GBOTB.extended.tre' is the latest and largest dated phylogenetic tree for vascular plants, with 74533 species, 10587 genera and all extant 479 families (Jin and Qian 2019). The R package 'V. PhyloMaker' (Jin and Qian 2019) can bind undetermined plant taxa to the backbone phylogeny of 'GBOTB.extended.tre’ and generates the customised tree we needed (Dai et al. 2020). Here, we obtained the phylogenetic tree of our host fern families using the above method. Bivariate linear regression was fitted with Past 4.04 (Hammer et al. 2001). Both leaf-miners and stem-borers have been found in the same insect family (e.g. Buprestidae, Cossidae and Blasticotomidae) or even in the same genus (e.g. Amauromyza, Melanagromyza, Phytomyza and Zygoneura) (Eiseman 2020b, Hering 1951, Shcherbakov 2006, Woodley and Janzen 1995). Occasionally, the same species could change their feeding habits from leaf-mining to stem-mining or stem-boring, when the younger larvae transform into the older larvae, when the leaf is too small to offer enough food or when leaves and stems do not differ significantly (Eiseman 2020b, Hering 1951). Such phenomena can be found in Heliozela hammoniella (= Heliozela_ betulae) (Heliozelidae), Marmara spp. (Gracillariidae), Ophiomyia spp. (Agromyzidae), Phyllocnistis spp. (Gracillariidae), Scaptomyza graminum (= Scaptomyzella incana) (Drosophilidae), Zygoneura calthella (Sciaridae), Temnosira czurhini (Pallopteridae) and many other species (Eiseman 2020b, Ellis 2020, Hering 1951, Kato 2002). There are transitions amongst leaf-mining, stem-mining, leaf-boring and stem boring (Hering 1951). Moreover, most ferns are herbaceous, with developed parenchyma in the stems (Crang et al. 2018). Therefore, we incorporated fern borers into fern miners for this article (Suppl. material 1). Some suspected insect species are not included in this study (e.g. Correia et al. 2020, Santiago-Blay 2004). The miners’ biogeographical regions followed Juan J. Morrone's system (Morrone 2002). For detailed information about fern miners associated with each host plant species, the original sources should be consulted. 4 Yang J et al Results We recorded 128 species and 18 families of fern miners (Table 1; Suppl. material 2), including Agromyzidae, Anthomyiidae, Drosophilidae, Chironomidae, Pallopteridae, Buprestidae, Chrysomelidae, Curculionidae, Crambidae, Noctuidae, Tineidae, Tortricidae, Cosmopterigidae, Gelechiidae, Hepialidae, Psychidae, Blasticotomidae and Tenthredinidae. They were primarily distributed in the Palaearctic Region, Nearctic Region and Oriental Region of the Northern Hemisphere and the Neotropical Region between the Tropic of Cancer and the Tropic of Capricorn (Table 1). One explanation for this distribution pattern could be that the land area in the Northern Hemisphere is almost double that of the Southern Hemisphere. Another reason might be that the investigations on leaf-mining insects and their host plants are more thorough in the Northern than in the Southern Hemisphere (Sinclair and Hughes 2008, Sinclair and Hughes 2008, Sinclair and Hughes 2010). Table 1. A preliminary world checklist of fern miners and their biological information. Miner family Miner species Host fern Biogeographical References regions Diptera Agromyzidae Chromatomyia Cheilanthes Oriental Region Spencer 1990 cheilanthus Garg*" virga-aure Agromyzidae Chromatomyia dorsata Asplenium Palaearctic Region Ellis 2020, Spencer 1990 Hendel? ceterach Agromyzidae Chromatomyia Dryopteris lacera_ Palaearctic Region Sasakawa 2010, Spencer dryoptericola 1990 Sasakawa? Agromyzidae Chromatomyia Asplenium Palaearctic Region dryoptericola pinnatifidum Sasakawa Agromyzidae Chromatomyia Lepisorus Palaearctic Region dryoptericola thunbergianus Sasakawa Agromyzidae Chromatomyia Lepisorus Palaearctic Region Sasakawa 2010 masumiae Sasakawa thunbergianus Agromyzidae Chromatomyia scolopendri Robineau Desvoidy* Asplenium ruta- muraria Palaearctic Region Agromyzidae Chromatomyia Asplenium Palaearctic Region scolopendri Robineau — scolopendrium Desvoidy Agromyzidae Chromatomyia Asplenium Palaearctic Region scolopendri Robineau _ septentrionale Desvoidy Civelek 2002, Dempewolf 2001, Ellis 2020, Sasakawa 2010, Spencer 1990 Miner family Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae A preliminary world checklist of fern-mining insects Miner species Chromatomyia scolopendri Robineau Desvoidy Liriomyza equiseti de Meijere® Liriomyza occipitalis Hendel® Liriomyza nordica Spencer* Liriomyza virgo Zetterstedt’ Liriomyza virgo Zetterstedt Liriomyza virgula Frey® Phytoliriomyza clara Melander Phytoliriomyza cyatheae Spencer Phytoliriomyza cyatheae Spencer Phytoliriomyza diplazii Sasakawa Phytoliriomyza felti Malloch Phytoliriomyza felti Malloch Phytoliriomyza felti Malloch Phytoliriomyza felti Malloch Phytoliriomyza felti Malloch Phytoliriomyza felti Malloch Phytoliriomyza flavopleura Watt Phytoliriomyza flavopleura Watt Host fern Polypodium vulgare Equisetum arvense Equisetum arvense Equisetum sp. Equisetum fluviatile Equisetum palustre Equisetum arvense Pteridium aquilinum Cyathea dealbata Cyathea smithii Diplazium Asplenium pinnatifidum Asplenium platyneuron Pellaea atropurpurea Pellaea glabella Asplenium rhizophyllum Woodsia obtusa Microsorum Notogrammitis billardierei Biogeographical regions Palaearctic Region Nearctic and Palaearctic Regions Palaearctic Region Nearctic Region Nearctic and Palaearctic Regions Nearctic and Palaearctic Regions Palearctic Regain Nearctic Region Neozelandic Region Neozelandic Region Oriental Region Nearctic Region Nearctic Region Nearctic Region Nearctic Region Nearctic Region Nearctic Region Neozelandic Region Neozelandic Region References Eiseman 2020b, Ellis 2020, George 2014, Lonsdale 2017, Spencer 1990 Ellis 2020, George 2014, Spencer 1990 Eiseman 2020b Eiseman 2020b, Ellis 2020, George 2014, Lonsdale 2017, Spencer 1990 George 2014 Eiseman 2020b, Spencer 1990 Spencer 1976, Spencer 1990 Spencer 1990 Eiseman 2020b, Spencer 1990 Spencer 1990 Miner family Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Agromyzidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Miner species Phytoliriomyza flavopleura Watt Phytoliriomyza flavopleura Watt Phytoliriomyza hilarella Zetterstedt Phytoliriomyza hilarella Zetterstedt Phytoliriomyza hilarella Zetterstedt Phytoliriomyza kuscheli Spencer Phytoliriomyza pteridii Spencer Phytoliriomyza pulchella Spencer* Phytoliriomyza sp1 Phytoliriomyza sp2 Phytoliriomyza tearohensis Spencer Tropicomyia angioptericola Shiao Tropicomyia polyphaga Spencer Tropicomyia sp1 Tropicomyia sp1 Tropicomyia sp2 Chirosia albifrons Tiens Chirosia albitarsis Zetterstedt Chirosia asperistilata Suwa Chirosia asperistilata Suwa Yang J et al Host fern Asplenium flaccidum Asplenium oblongifolium Asplenium Pteridium aquilinum Polypodium Histiopteris Pteridium aquilinum Pteridium aquilinum Marattia* Cyathea Cyathea dealbata Angiopteris lygodiifolia Nephrolepis Pleopeltis Asplenium auriculatum Angiopteris evecta* Pteridium aquilinum Pteridium aquilinum Dryopteris monticola Dryopteris crassirhizoma Biogeographical regions Neozelandic Region Neozelandic Region Palaearctic Region Palaearctic Region Palaearctic Region Oriental Region Palaearctic Region Nearctic Region Oriental Region Neotropical and Andean Regions unknown Palaearctic Region Oriental Region Afrotopical Region Afrotopical Region Oriental Region Palaearctic Region Palaearctic and Oriental Regions Palaearctic Region Palaearctic Region References Ellis 2020, Lawton 1982, Lawton 1976, MacGarvin et al. 1986, Rigby and Lawton 1981, Spencer 1990 Spencer 1990 Gerson 1979, Mcgavin and Brown 1986, MacGarvin et al. 1986, Spencer 1990 Eiseman 2020b Spencer 1990 Spencer 1990 Spencer 1976, Spencer 1990 Shiao and Wu 2005 Spencer 1990, Shiao and Wu 2005 Spencer 1990 Spencer 1990 Lawton 1976, MacGarvin et al. 1986 Ellis 2020, Gerson 1979, Lawton 1976, Mcgavin and Brown 1986, Suwa 1984, Suwa 1999 Suwa 1999, Suwa 2005 Miner family Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae A preliminary world checklist of fern-mining insects Miner species Chirosia cinerosa Zetterstedt? Chirosia cinerosa Zetterstedt Chirosia cinerosa Zetterstedt Chirosia crassiseta Stein Chirosia filicis Huckett Chirosia filicis Huckett Chirosia flavipennis Fallen Chirosia gleniensis Huckett Chirosia gleniensis Huckett Chirosia gleniensis Huckett Chirosia griseifrons Séguy Chirosia griseifrons Séguy Chirosia grossicauda Strobl'° Chirosia grossicauda Strobl Chirosia grossicauda Strobl Chirosia histricina Rondani" Chirosia histricina Rondani Chirosia histricina Rondani Chirosia histricina Rondani Chirosia histricina Rondani Host fern Pteridium aquilinum Matteuccia struthiopteris Athyrium filix- femina Pteridium aquilinum Osmunda claytoniana Osmundastrum cinnamomeum Pteridium aquilinum Onoclea sensibilis Woodsia areolata Woodsia virginica * Dryopteris* Athyrium filix- femina Asplenium Pteridium aquilinum Dryopteris* Osmunda regalis Asplenium Blechnum spicant Pteridium aquilinum Dryopteris filix- mas Biogeographical regions Palaearctic Region Palaearctic Region Palaearctic Region Palaearctic Region Nearctic Region Nearctic Region Nearctic and Palaearctic Regions Nearctic Region Nearctic Region Nearctic Region Palaearctic Region Palaearctic Region Palaearctic Region Palaearctic Region Palaearctic Region Nearctic and Palaearctic Regions Nearctic and Palaearctic Regions Nearctic and Palaearctic Regions Nearctic and Palaearctic Regions Nearctic and Palaearctic Regions References Ellis 2020, Kwon and Suh 1982, Suwa 1999 Brown and McGavin 2007, Ellis 2020, Gerson 1979, Lawton 1976, Mcgavin and Brown 1986 Eiseman 2018, Eiseman 2020b Eiseman 2020b, Eiseman 2018, Lawton 1976, Suwa 2013 Eiseman 2020b, Eiseman 2018, Eiseman 2020a Ellis 2020, Suwa 1999 Ellis 2020, Gerson 1979, Lawton 1976, MacGarvin et al. 1986, Mcgavin and Brown 1986, Suwa 1999 Brown and McGavin 2007, Ellis 2020, MacGarvin et al. 1986, Mcgavin and Brown 1986, Suwa 1999 Miner family Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Anthomyiidae Drosophilidae Drosophilidae Drosophilidae Drosophilidae Drosophilidae Chironomidae Pallopteridae Lepidoptera Miner species Chirosia histricina Rondani Chirosia histricina Rondani Chirosia histricina Rondani Chirosia histricina Rondani Chirosia montana Pokorny Chirosia nigripes Bezzi Chirosia pusillans Huckett Chirosia pusillans Huckett Chirosia spinosissima Malloch Pegomya cedrica Huckett Pegomya disticha Griffiths Pegomya glabra Stein Drosophila apicipuncta Hardy Drosophila diminuens Hardy* Drosophila sadleria Bryan Scaptodrosophila notha Bock Scaptodrosophila sp. Bryophaenocladius furcatus Kieffer Temnosira czurhini Ozerov Yang J et al Host fern Matteuccia struthiopteris Polypodium Athyrium filix- femina Cystopteris fragilis Cystopteris fragilis Pteridium aquilinum Athyrium filix- femina Matteuccia struthiopteris Pteridium aquilinum Equisetum hyemale Equisetum hyemale Equisetum Sadleria Sadleria Sadleria Pteridium aquilinum Parablechnum wattsii Adiantum Huperzia serrata Biogeographical regions Nearctic and Palaearctic Regions Nearctic and Palaearctic Regions Nearctic and Palaearctic Regions Nearctic and Palaearctic Regions Nearctic and Palaearctic Regions Palaearctic Region Nearctic Region Nearctic Region Nearctic and Palaearctic Regions Nearctic Region Nearctic Region Nearctic Region Nearctic Region Nearctic Region Nearctic Region Australotropical and Australotemperate Regions Australotropical and Australotemperate Regions Nearctic and Palaearctic Regions Palaearctic Region References Eiseman 2020b, Eiseman 2018 Ellis 2020, Suwa 1999 Eiseman 2018, Eiseman 2020a, Eiseman 2020b Eiseman 2020b, Eiseman 2020a Michelsen and Palmer 2020 Michelsen and Palmer 2020 Michelsen and Palmer 2020 Magnacca et al. 2008, Magnacca and O'Grady 2014, Maunsell et al. 2016 Magnacca and O'Grady 2014 Magnacca et al. 2008 Maunsell et al. 2016 Maunsell et al. 2016 Eiseman 2020b Kato 2002 Miner family Crambidae Crambidae Crambidae Crambidae Crambidae Crambidae Crambidae Crambidae Noctuidae Noctuidae Noctuidae Noctuidae Noctuidae Noctuidae Noctuidae Noctuidae Noctuidae Tineidae A preliminary world checklist of fern-mining insects Miner species Albusambia elaphoglossumae Solis & Davis Albusambia elaphoglossumae Solis & Davis Eudonia zophoclaena Meyrick Scoparia illota Philpott Scoparia molifera Meyrick Siamusotima aranea Solis & Yen Siamusotima disrupta Solis Undulambia polystichalis Capps Hydraecia micacea Esper Papaipema inquaesita Grote & Robinson Papaipema pterisii Bird Papaipema pterisii Bird Papaipema speciosissima Grote & Robinson Papaipema speciosissima Grote & Robinson Papaipema stenocelis Dyar Pseudobryomima fallax Hampson Pseudobryomima muscosa Hampson Psychoides filicivora Meyrick’? Host fern Elaphoglossum conspersum Elaphoglossum biolleyi Pyrrosia eleagnifolia Pyrrosia eleagnifolia Pyrrosia eleagnifolia Lygodium flexuosum Lygodium Rumohra adiantiformis Equisetum Onoclea sensibilis Pteridium aquilinum Matteuccia struthiopteris* Osmunda regalis Osmundastrum cinnamomeum Woodwardia virginica Pellaea andromedifolia Polypodium californicum Asplenium adiantum-nigrum Biogeographical regions Nearctic Region Nearctic Region Neozelandic Region Neozelandic Region Neozelandic Region Oriental Region Palaearctic Region Nearctic Region Palearctic Region Nearctic Region Nearctic Region Nearctic Region Nearctic Region Nearctic Region Nearctic Region Nearctic Region Nearctic Region Palaearctic Region References Solis et al. 2005a Patrick 2015 Patrick 2015 Patrick 2015 Solis et al. 2005b Solis et al. 2017 Gerson 1979 Ellis 2020 Bird 2012 Bird 2012, Hinz and Zahniser 2015 Schweitzer 2012 Hinz and Zahniser 2015, Lafontaine and Schmidt 2010, Oppenheim et al. 2018 Chaloux, Andrea 2012 Eiseman 2020b Eiseman 2020b Gaedike 2019, Kim and Bae 2007 10 Miner family Tineidae Tineidae Tineidae Tineidae Tineidae Tineidae Tineidae Tineidae Tineidae Tineidae Tineidae Tineidae Tineidae Tineidae Tineidae Tineidae Tineidae Tortricidae Tortricidae Tortricidae Miner species Psychoides filicivora Meyrick Psychoides filicivora Meyrick Psychoides filicivora Meyrick Psychoides filicivora Meyrick Psychoides filicivora Meyrick Psychoides filicivora Meyrick Psychoides gosari Kim & Bae Psychoides gosari Kim & Bae Psychoides gosari Kim & Bae Psychoides gosari Kim & Bae Psychoides gosari Kim & Bae Psychoides phaedrospora Meyrick 13 Psychoides verhuella Bruand‘4 Psychoides verhuella Bruand Psychoides verhuella Bruand Psychoides verhuella Bruand Psychoides verhuella Bruand Apoctena taipana Felder & Rogenhofer Celypha tiedemanniana Zeller'® Philocryptica polypodii Watt Yang J et al Host fern Asplenium ceterach Asplenium scolopendrium Asplenium trichomanes Dryopteris filix- mas Dryopteris aculeata Polystichum setiferum Athyrium yokoscense Dryopteris setosa Dryopteris chinensis Dryopteris crassirhizoma Dryopteris saxifraga Aspleniaceae Asplenium ceterach Asplenium ruta- muraria Asplenium scolopendrium Asplenium trichomanes Pteridium aquilinum Pyrrosia eleagnifolia Equisetum Pyrrosia eleagnifolia Biogeographical regions Palaearctic Region Palaearctic Region Palaearctic Region Palaearctic Region Palaearctic Region Palaearctic Region Oriental and Palaearctic Regions Oriental and Palaearctic Region Oriental and Palaearctic Regions Oriental and Palaearctic Regions Oriental and Palaearctic Regions Palaearctic and Oriental Regions Palaearctic Region Palaearctic Region Palaearctic Region Palaearctic Region Palaearctic Region Neozelandic Region Palaearctic Region Neozelandic Region References Kim and Bae 2007 Gaedike 2019, Kim and Bae 2007 Ellis 2020, Gaedike 2019, Heckford 2004, Kim and Bae 2007, Muus 2015 Patrick 2015 Ellis 2020 Patrick 2015 Miner family Cosmopterigidae Cosmopterigidae Cosmopterigidae Cosmopterigidae Cosmopterigidae Gelechiidae Gelechiidae Hepialidae Hepialidae Psychidae Hymenoptera Blasticotomidae Blasticotomidae Blasticotomidae Blasticotomidae Blasticotomidae Blasticotomidae Blasticotomidae Blasticotomidae A preliminary world checklist of fern-mining insects Miner species Hyposmocoma ( Euperissus) ekaha Swezey Hyposmocoma ( Euperissus) trivitella Swezey Hyposmocoma ( Euperissus) trivitella Swezey Hyposmocoma ( Euperissus) trivitella Swezey Hyposmocoma ( Euperissus) trivitella Swezey Monochroa harrisonella Busck Paltodora cytisella Curti Endoclita excrescens Butler* Triodia sylvina Linnaeus* Apterona helicoidella Vallot Blasticotoma atra Zhelochovtsev Blasticotoma filiceti Klug Blasticotoma filiceti Klug Blasticotoma filiceti Klug Blasticotoma filiceti Klug Blasticotoma filiceti Klug Blasticotoma filiceti Klug Blasticotoma filiceti var. pacificus Malaise Host fern Asplenium nidus Elaphoglossum aemulum Elaphoglossum gorgoneum Elaphoglossum crassifolium Elaphoglossum reticulatum Pteridium aquilinum Pteridium aquilinum Equisetum arvense Equisetum arvense Polypodium unknown Pteridium aquilinum Dryopteris Polystichum Matteuccia struthiopteris Athyrium alpestre Athyrium filix- femina unknown Biogeographical regions Oriental Region Oriental Region Oriental Region Oriental Region Oriental Region Nearctic Region Palaearctic Region Palaearctic Region Palaearctic Region unknown unknown Palaearctic Region Palaearctic Region Palaearctic Region Palaearctic Region Palaearctic Region Palaearctic Region unknown References Kawahara et al. 2011 Kawahara et al. 2011 Eiseman 2020b Lawton 1976, Rigby and Lawton 1981 Correia et al. 2020, Grehan 1989 Correia et al. 2020, Grehan 1989 Alders and Gielis 1999 Taeger et al. 2010, Wikipedia 2019 Ellis 2020, Liston 2007, Novgorodova and Biryukova 2011, Shcherbakov 2006, Shcherbakov 2008, Taeger et al. 2010, Wikipedia 2019 Taeger et al. 2010, Wikipedia 2019 11 12 Miner family Blasticotomidae Blasticotomidae Blasticotomidae Blasticotomidae Blasticotomidae Blasticotomidae Blasticotomidae Blasticotomidae Blasticotomidae Blasticotomidae Tenthredinidae Tenthredinidae Tenthredinidae Tenthredinidae Tenthredinidae Tenthredinidae Tenthredinidae Tenthredinidae Tenthredinidae Tenthredinidae Coleoptera Miner species Blasticotoma nipponica Takeuchi Blasticotoma smithi Shinohara Blasticotoma warabii Togashi Bohea abrupta Maa Paremphytus ostentus Brues Runaria flavipes Takeuchi Runaria hunannica Wei in Wei & Nie Runaria punctata Wei in Wei & Nie Runaria shaanxinica Wei in Wei & Nie Runaria taiwana Shinohara Aneugmenus coronatus Klug Aneugmenus coronatus Klug Aneugmenus coronatus Klug Aneugmenus coronatus Klug Heptamelus dahlbomi Thomson Heptamelus ochroleucus Stephens 16 Heptamelus ochroleucus Stephens Heptamelus ochroleucus Stephens Heptamelus ochroleucus Stephens Heptamelus ochroleucus Stephens Yang J et al Host fern unknown unknown unknown unknown unknown unknown unknown unknown unknown unknown Pteridium aquilinum Dryopteris filix- mas Polystichum setiferum Athyrium filix- femina Athyrium filix- femina Blechnum spicant Matteuccia struthiopteris Dryopteris dilatata Polypodium vulgare Athyrium filix- femina Biogeographical regions unknown unknown unknown unknown unknown unknown unknown unknown unknown unknown Palaearctic Region Palaearctic Region Palaearctic Region Palaearctic Region Nearctic and Palaearctic Regions Nearctic and Palaearctic Regions Nearctic and Palaearctic Regions Nearctic and Palaearctic Regions Nearctic and Palaearctic Regions Nearctic and Palaearctic Regions References Wikipedia 2019, Taeger et al. 2010 Taeger et al. 2010, Wikipedia 2019 Taeger et al. 2010, Wikipedia 2019 Taeger et al. 2010, Wikipedia 2019 Taeger et al. 2010, Wikipedia 2019 Taeger et al. 2010, Wikipedia 2019 Taeger et al. 2010, Wikipedia 2019 Taeger et al. 2010, Wikipedia 2019 Taeger et al. 2010, Wikipedia 2019 Taeger et al. 2010, Wikipedia 2019 Benes 2014, Ellis 2020, Schwarz 2005 Vikberg and Liston 2009 Ellis 2020, Shcherbakov 2008, Vikberg 2017, Vikberg and Liston 2009 Miner family Buprestidae Buprestidae Buprestidae Buprestidae Buprestidae Buprestidae Buprestidae Buprestidae Buprestidae Buprestidae Buprestidae Buprestidae Buprestidae Buprestidae Buprestidae Chrysomelidae Chrysomelidae A preliminary world checklist of fern-mining insects Miner species Endelus bakerianus Obenberger Neotrachys bellamyi Hespenheide Neotrachys bicolor Hespenheide Neotrachys bordoni Cobos Neotrachys caerulea Hespenheide Neotrachys concinna Fisher Neotrachys cyanipennis Fisher Neotrachys estebana Kerremans* Neotrachys fennahi Thery Neotrachys gleicheniae Hespenheide Neotrachys hoffmani Fisher Neotrachys mariae Hespenheide Neotrachys resplendens Hespenheide Neotrachys segregata Waterhouse Neotrachys solisi Hespenheide Febra insularis Bryant Febra ovata Bryant Host fern Lygodium microphyllum Gleichenia glauca Cnemidaria petiolata Cyatheaceae Cyatheaceae Cyatheaceae Cyatheaceae Dicranopteris Cyatheaceae Gleichenia Cyatheaceae Gleichenia Cyatheaceae Gleicheniaceae* Gleichenia Acrostichum aureum Angiopteris evecta Biogeographical regions Oriental Region Neotropical Region Neotropical Region Neotropical Region Neotropical Region Neotropical Region Neotropical Region Neotropical Region Neotropical Region Neotropical Region Neotropical Region Neotropical Region Neotropical Region Neotropical Region Neotropical Region Oriental Region Oriental Region 13 References Kalashian 2013, Goolsby et al. 2003, Mehltreter et al. 2010 Hespenheide 2006 Hespenheide 1982 Hespenheide 1982 Hespenheide 1982 Hespenheide 1982, Hespenheide 2006 Hespenheide 2006 Hespenheide 1982 Hespenheide 1980 Hespenheide 1982 Hespenheide 1980, Hespenheide 1982 Hespenheide 2006 Hespenheide 1982 Hespenheide 1982 Hespenheide 2006 Samuelson 1973, Santiago-Blay 2004 Samuelson 1973, Nadein 2013, Jolivet 1991 14 Miner family Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Miner species Febra venusta Clark Halticorcus bhaumiki Basu et Sengupta’” Halticorcus bhaumiki Basu et Sengupta Halticorcus bhaumiki Basu et Sengupta Halticorcus bhaumiki Basu et Sengupta Halticorcus bhaumiki Basu et Sengupta Halticorcus bhaumiki Basu et Sengupta Halticorcus bhaumiki Basu et Sengupta Halticorcus bhaumiki Basu et Sengupta Halticorcus bhaumiki Basu et Sengupta Halticorcus bhaumiki Basu et Sengupta Halticorcus hiranoi Takizawa'® Halticorcus hiranoi Takizawa Halticorcus kasuga Nakane Halticorcus kasuga Nakane Halticorcus kasuga Nakane Halticorcus kasuga Nakane Halticorcus platycerii Lea Halticorcus platycerii Lea Halticorcus sauteri Chen‘? Yang J et al Host fern Nephrolepis Pteris vittata Ampelopteris prolifera Cyclosorus Christella dentata Nephrolepis cordifolia Nephrolepis exaltata Adiantum lunulatum Drynaria propinqua Pyrrosia adnascens Microsorum scolopendria Lemmaphyllum microphyllum Loxogramme salicifolia* Lepisorus thunbergianus Lepisorus onoei Lemmaphyllum microphyllum Pyrrosia linearifolia Platycerium alcicorne Asplenium nidus Colysis elliptica Biogeographical regions Oriental Region Palaearctic and Oriental Regions Palaearctic and Oriental Regions Palaearctic and Oriental Regions Palaearctic and Oriental Regions Palaearctic and Oriental Regions Palaearctic and Oriental Regions Palaearctic and Oriental Regions Palaearctic and Oriental Regions Palaearctic and Oriental Regions Palaearctic and Oriental Regions Palaearctic Region Palaearctic Region Palaearctic Region Palaearctic Region Palaearctic Region Palaearctic Region Australotropical, Australotemperate and Palaearctic Regions Australotropical, Australotemperate and Palaearctic Regions Palaearctic Region References Samuelson 1973, Santiago-Blay 2004 Isowa and Kojima 2011, Konstantinov and Prathapan 2008, Patra and Bera 2007 Kato 1991, Santiago-Blay 2004 Isowa and Kojima 2011 Hawkeswood 2003, lsowa and Kojima 2011, Sinclair and Hughes 2010 Kato 1991, Santiago-Blay 2004 Miner family Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Chrysomelidae Curculionidae Curculionidae Curculionidae Curculionidae Curculionidae Curculionidae Curculionidae Curculionidae Curculionidae Curculionidae A preliminary world checklist of fern-mining insects Miner species Halticorcus sauteri Chen Hippuriphila babai Chujo* Hippuriphila canadensis Brown* Hippuriphila catherinae Barr* Hippuriphila equiseti Beller et Hatch* Hippuriphila modeeri Linnaeus Hippuriphila modeeri Linnaeus Hippuriphila modeeri Linnaeus Bagous claudicans Boheman Bagous lutulentus Gyllenhal#° Grypus brunnirostris Fabricius?! Grypus brunnirostris Fabricius Grypus brunnirostris Fabricius Grypus equiseti Fabricius?2 Grypus equiseti Fabricius Grypus equiseti Fabricius Grypus equiseti Fabricius Stenopelmus rufinasus Gyllenhal * possible host fern or miner Host fern Leptochilus ellipticus Equisetum Equisetum arvense Equisetum Equisetum arvense Equisetum arvense Equisetum fluviatile Equisetum palustre Equisetum fluviatile Equisetum fluviatile Equisetum arvense Equisetum fluviatile Equisetum ramosissimum Equisetum arvense Equisetum palustre Equisetum pratense Equisetum sylvaticum Azolla Biogeographical regions Palaearctic Region Palaearctic Region Nearctic Region Neotropical Region Nearctic Region Palaearctic Region Palaearctic Region Palaearctic Region except for Central and South America, all the world except for Central and South America, all the world Nearctic Region Nearctic Region Nearctic Region Nearctic and Palaearctic Regions Nearctic and Palaearctic Regions Nearctic and Palaearctic Regions Nearctic and Palaearctic Regions Nearctic, Afrotopical and Palaearctic Regions 15 References Correia et al. 2020, Poinar 2014 Correia et al. 2020, Poinar 2014 Correia et al. 2020, Poinar 2014 Correia et al. 2020, Poinar 2014 Biological Records Centre 2020, Ellis 2020, Santiago-Blay 2004 Ellis 2020, Gosik et al. 2019 Ellis 2020, Gosik 2009, Gosik et al. 2019, Gosik and Wanat 2014 Ellis 2020, George 2014 Ellis 2020, George 2014, Gosik et al. 2019 Center et al. 2002, Hill and Cilliers 1999, Richerson and Grigarick 1967 16 Yang J et al Synonyms: 1 Chromatomyia cheilanthus Garg = Phytomyza cheilanthus Garg; 2 Chromatomyia dorsata Hendel = Phytomyza dorsata Hendel; 3 Chromatomyia dryoptericola Sasakawa = Phytomyza dryoptericola Sasakawa 4 Chromatomyia scolopendri Robineau-Desvoidy = Phytomyza scolopendri Goureau = Phytomyza elegans Goureau = Phytomyza nevadensis Strobl = Chromatoinyia nevadensis Strobl; 5 Liriomyza equiseti de Meijere = Liriomyza kenti Spencer; 6 Liriomyza occipitalis Hendel = Liriomyza bruscae Hering; 7 Liriomyza virgo Zetterstedt = Liriomyza arcticola Spencer = Phytomyza pallicornis Zetterstedt; 8 Liniomyza virgula Frey = Liniomyza larissa Hering; 9 Chirosia cinerosa Zetterstedt = Pycnoglossa cinerosa Zetterstedt; 10 Chirosia grossicauda Strobl = Chirosia parvicornis nec Zetterstedt; 11 Chirosia histricina Rondani = Chirosia setifemur Ringdahl = Pycnoglossa hystrix Brischke = Pycnoglossa hystricina; 12 Psychoides filicivora Meyrick = Teichobia filicivora Meyrick; 13 Psychoides phaedrospora Meyrick = Mnesipatris phadrospora Meyrick; 14 Psychoides verhuella Bruand = Teichobia verhuellella Herrich-Schaffer = Lambrosetia verhuellella Stainton; 15 Celypha tiedemanniana Zeller = Olethreutes tiedemanniana Zeller Kuznetsov = Syricoris tiedemanniana Zeller; 16 Heptamelus ochroleucus Stephens = Melicerta ochroleucus Stephens; 17 Halticorcus bhaumiki Basu et Sengupta = Schenklingia bhaumiki Basu and Sengupta; 18 Halticorcus hiranoi Takizawa = Schenklingia hiranoi Takizawa; 19 Halticorcus sauteri Chen = Schenklingia sauteri Chen; 20 Bagous lutulentus Gyllenhal = Bagous glabrirostris var. nigritarsis Thomson; 21 Grypus brunnirostris Fabricius = Curculio Inspectionatus Herbst; 22 Grypus equiseti Fabricius = Grypidius equiseti Fabricius; Fern miners belong to four orders: Diptera (51 species; 39.8%), Coleoptera (33 species; 25.8%), Lepidoptera (28 species; 21.9%) and Hymenoptera (16 species; 12.5%) (Fig. 1; Suppl. material 2). In general, dipteran leaf miners are dominant in herbaceous plants while lepidopteran leaf miners are dominant in woody plants (De Prins and De Prins 2020, Edmunds et al. 2020, Eiseman 2020b, Ellis 2020, Pitkin et al. 2019, Spencer 1990). The life form of most extant ferns is herbaceous, which could explain why nearly half of fern- mining species are dipteran flies. A preliminary world checklist of fern-mining insects 17 Tenthredinidae Psychidae GE Diptera HE Coleoptera Wa = Lepidoptera Ga) =Hymenoptera Gelechiidae 1% Hepialidae 2% Cosmopterigidae 2% Tortricidae 2% Chironomidae Pallopteridae 1% Figure 1. EESI Percentage distribution of the fern-mining species into the four orders and the 18 families. Amongst the 18 fern-mining insect families, Agromyzidae, Anthomyiidae, Buprestidae, Chrysomelidae and Blasticotomidae had the highest numbers of species (20.3%, 14.1%, 11.7%, 10.2% and 10.2%, respectively), while the other 13 families accounted for 33.5% only (Fig. 1; Suppl. material 2). The fern families with highest numbers of host species were Dryopteridaceae (19), Polypodiaceae (18) and Aspleniaceae (15) (Fig. 2; Suppl. material 3). The fern families with the highest numbers of miner species were Dennstaedtiaceae (21), Equisetaceae (21), Polypodiaceae (20) and Aspleniaceae (14) (Fig. 2; Suppl. material 3). With 82 species and 12 families of host ferns and 67 species of fern miners, Polypodiales was the dominant host order of fern-mining insects (Suppl. material 3). 27.64 eT ae Blechnaceae AT 27.64 a Onocleaceae 2 at -_—_—_—_———— Thelypteridaceae 3/1 Cystoptendaceae W2 Dryopteridaceae 19/13 Polypodiaceae 18/20 Dennstaedtiaceae 2/21 Ptendaceae 8/6 Cyatheaceae S/11 Salviniaceae Lia) Lygodiaceae 3 Gleicheniaceae 46 Osmundaceae 70.79 7 Equisetaceae 8/21 Marattiaceae 34 Lycopodiaceae Ww Figure 2. EES The dated phylogenetic tree of host fern families generated with the R package 'V. PhyloMaker' (Jin and Qian 2019). The first number after the fern family is the number of host fern species and the second is the number of fern miner species. The length of each branch is also shown and the scale bar unit is 100 myr. 18 Yang J et al The number of host species was significantly and positively correlated with the total number of fern species at the family level (R* = 0.614, t= 5.352, P < 0.001; Fig. 3a), but the number of miner species was not significantly correlated with the total number of fern species at the family level (R@ = 0.110, t= 1.495, P = 0.152; Fig. 3b). a b the number of host fern species the number of miner species T T T T T T T T T T T T T T T T T T 250 500 750 1000 1250 1500 1750 2000 2250 i] 250 500 750 1000 1250 1500 1750 2000 2250 Total number of fern species Total number of fern species Figure 3. Linear regressions between (a) the number of host species in each fern family; (b) the number of miner species for each fern family and the total number of fern species in the corresponding family. Families without any host species were not included. Discussion In this paper, we provide a preliminary checklist about fern miners and their host plants worldwide. Table 1 summarises this checklist in terms of published information to date. However, there is also more information available on some fern-mining groups and this is summarised here: (1) Diptera: In Anthomyiidae, there is an unknown Chirosia species with Deparia acrostichoides as host in the Nearctic Region (Eiseman 2020b), while C. similata could be a possible Nearctic Pteridium borer (Eiseman 2018). In Drosophilidae, the Fuscoamoeba subgroup has many species that have been reared from rotting fern rachises (Magnacca et al. 2008). For Chromatomyia species in Agromyzidae, Kahanpaa (2014) separates Chromatomyia and Napomyza as different genera (Kahanpaa 2014) and Spencer (1990) considers that C. cheilanthus should belong to the genus Pfochomyza (Spencer 1990). Molecular phylogeny suggests that the genus of Phytomyza should include all species of Phytomyza, Chromatomyia, Napomyza and Ptochomyza (Winkler et al. 2009). However, only one fern-feeding Chromatomyia species is included in the above molecular analysis. Moreover, no Phytomyza s. s. species has previously been found as fern-mining. In this article, we rather kept the genus name of Chromatomyia and listed the Phytomyza species as the synonym of the corresponding Chromatomyia species in . A preliminary world checklist of fern-mining insects 19 (2) Lepidoptera: In Tineidae, early instar larvae of the subfamily Teichobinae are leaf miners, while their later instars feed on sporangia from a loose portable case (Gaedike 2019). An unknown species of Pyralidae has two hosts (Lygodium microphyllum and L. flexuosum) in the Oriental Region (Goolsby et al. 2003). There is an unknown moth in the Nearctic Region, which mines the leaves of Pteridium aquilinum (Eiseman 2020b), but the species name could not be confirmed. In Gelechiidae, Monochroa placidella larvae make gall-like deformities on the fronds of the bracken (P. aquilinum) (Eiseman 2020b). Eiseman (personal observations) believes that the deformities are caused by internal feeding; he has also reared an undetermined Monochroa species from larvae that similarly bored in the terminal part of the rachis and caused a gall-like deformity. (3) Hymenoptera: In Tenthredinidae, the genus Heptamelus has 36 species in the Palaearctic and Oriental Regions and their larvae are internal feeders and all probably use ferns as larva hosts (Vikberg and Liston 2009), but we cannot know with certainty which species of Heptamelus is involved, except for H. ochroleucus on Athyrium filix-femina (Vikberg 2017). With only 13 species and 3 tribes in Eurasia's temperate region, Blasticotomidae is a small family in the Hymenoptera and their larvae are stem borers on ferns (Taeger et al. 2010, Wikipedia 2019, Santiago-Blay 2004). (4) Coleoptera: In Buprestidae, both Neotrachys and Endelus have fern-mining habits (Xiao 2018, Bellamy 1997). Most Neotrachys feed on the ferns of Cyatheaceae and Gleicheniaceae (Hespenheide 1980, Hespenheide 1982, Hespenheide 2006). However, some Neotrachys larvae may mine other non-fern plants. For example, N. dominicanus feeds on Arthrostylidium (Poaceae) (Meurgey 2017). The genus’ Leiopleura is morphologically similar and sometimes confused with Neotrachys, but Leiopleura feeds on Moraceae and Apocynaceae (Hespenheide 1991). Fern-feeding or not could be a clue to distinguish Neotrachys and its related genera. Although there are many publications on Endelus, only very few mention its host plants (Kalashian 2013). Dominant plant groups generally are rich in leaf miners and rich in host plants, which could be explained by the ‘plant apparency hypothesis’ (Feeny 1976). Such phenomena have been found in several other leaf-mining insects (Dai et al. 2017, Dai et al. 2018). Apart from species richness in a fern taxonomic group, the distribution range should also be considered as an important component of 'plant apparency' (Dai et al. 2017). Equisetaceae has 39 species and eight host species, and Dennstaedtiaceae has 245 species and two host species, but both families host 21 miner species, which is the highest amongst all fern families (Fig. 2; Suppl. material 3). It is Equisetaceae and Dennstaedtiaceae that strongly affected the significance of the correlation in Fig. 3b. In particular, the bracken fern ( Pteridium aquilinum), one species in Dennstaetiaceae, had 20 miner species (Suppl. material 3), which is not less than many dominant flowering plants. The bracken fern might be the only globally distributed fern and one of the most widespread vascular plants, which occurs in temperate and subtropical regions in both hemispheres (Flora of North America Editorial Committee 1993). It is used as vegetable, food or feed in many places. It is also a common invasive plant in disturbed areas (Flora of North America Editorial Committee 1993). The above features of the bracken fern make it highly attractive to both miners and researchers, thus the high number of mining species might be the combined effects of 20 Yang J et al plant apparency and sampling effects. Dryopteridaceae has 2257 species (Suppl. material 3) and also a cosmopolitan distribution, with many cultivated ornamental species (Olsen and Olsen 2007). Aspleniaceae has 855 species (Suppl. material 3) and also a worldwide distribution (POWO 2019). Polypodiaceae has 1667 species (Suppl. material 3) and is distributed nearly worldwide, but mainly in tropical areas, with some cultivated species (Simpson 2010). Both high species richness and wide geographical distribution could explain why the three families have large numbers of both host fern species and miner species. Besides P aquilinum, Equisetum arvense (Equisetaceae), Athyrium filix-femina (Athyriaceae) and Matteuccia struthiopteris (Onocleaceae) also have a high richness of miners (10, 8 and 6 species, respectively) (Suppl. material 3). The common horsetail (E. arvense) is native throughout the Arctic and temperate regions in the Northern Hemisphere (Schaffner 1930). E. arvense becomes an invasive plant in New Zealand and a systematic evaluation of its potential biocontrol agents including miners and borers has been performed (Paynter and Barton 2008). The common lady-fern (A. filix-femina) is one of the most abundant fern species in the temperate regions in the Northern Hemisphere (Adam 1995). The ostrich fern (MM. struthiopteris) is widely distributed in the temperate regions of the Northern Hemisphere (Kimura et al. 2004). However, since the checklist of fern-mining insects and the corresponding host fern species is preliminary, these patterns need further verification. As the sampling of fern miners and their hosts are insufficient in many places and some sampled records might be inaccessible, this study was only a preliminary list. We hope that this basic list can serve as an inital reference for future inventories and research on fern- mining insects. Acknowledgements This work was supported by the National Natural Science Foundation of China (31760173 and 41971059) and the Science and Technology Project of Ganzhou City. Kevin Duffy acknowledges the support of the National Research Foundation of South Africa (Grant Numbers 131604). We also thank our colleague, Dr. Zhongyang Li for his significant assistance in our understanding why some fern families host many more miners. Thanks are also given to the two reviewers (Dr. Charles Eiseman and Dr. Yume Imada) for their valuable comments, which significantly improved the manuscript. References ° Adam K (1995) Caffeic acid derivatives in fronds of the lady fern (Athyrium filix-femina). 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Nature 506 (7486): 89-92. https://doi.org/10.1038/nature12872 Supplementary materials Suppl. material 1: The feeding mode and feeding plant organ of each fern miner EE) Authors: Jie Yang, Xiaohua Dai Data type: Feeding habits Download file (16.21 kb) 28 Yang J et al Suppl. material 2: The number of fern-miners in each insect family and each insect order EE) Authors: Jie Yang, Xiaohua Dai Data type: Number of species Download file (16.04 kb) Suppl. material 3: The number of miners on each fern species, each fern family and each fern order EE) Authors: Jie Yang, Xiaohua Dai Data type: Number of species Download file (20.77 kb)