Oecologia (Berl.) 35, 55-89 (1978)

Biochemical and Evolutionary Aspects of Arthropod Predation on Ferns

Michael J. Balickl, David G. Furthz, and Gillian Cooper-Driver3 * ' Botanical Museum of Harvard University, Oxford Street, Cambridge, MA 02138, USA ' Department of Biology. Yale University, New Haven, CT 06520, USA ' Jhpartment of Biological Sciences, Boston University, 2 Cummington St., Boston, MA 02215, USA

Summary. The widely held assumption that very few arthropods feed on ferns was questioned following field observations of arthropod damage on ferns in the state of Veracruz, Mexico. The extent and type of damage was recorded and it was found that in a measured locality, ferns were no less attacked than the angiospermous flora. As chemistry and arthropod host relationships have been shown to be so closely intertwined, plants collected in the field were analysed for both condensed tannins and cyanoge- nic glycosides, compounds known to be effedtive deterrents in temperate climates. Although all ferns tested contained tannins these did not appear to inhibit predation. Cyanogenic glycosides were present in only 3% of the fern species analysed, and it is, therefork unlikely that they play a significant role as defensive compounds in the ferns examined. A literature search revealed a large number of ferns cited as being arthro- pod hosts. Approximately 420 named species of arthropods have been recorded, the majority of which are from the orders Coleoptera, Hymen- optera,. Lepidoptera, and Hemiptera. Both evolutionary primitive (sawflies) and advanced (moths) arthropods are reported to be present on ferns suggest- ing possible coevolution of arthropods and ferns both before and after the radiation of angiosperms.

I. Introduction

It is assumed that ferns generally are not eaten by herbivorous insects (Soo Hoo and Fraenkel, 1964; Eastopp 1973; Southwood, 1973). Indeed in their cl'assical paper on the co-evolution of bufferflies and plants Ehrlich and Raven (1964) state "In fact, very few insects feed on ferns at 'all, a most surprising and as yet unexplained fact with no evident chemical or mechanical basis ". 'Some ferns have toxic effects on both invertebrates (Carlisle and Ellis, 1968) * To whom offprint requests should be sent M.J. Balick et a]. as well as vertebrates (I.A. Evans, 1976; W.C. Evans, 1976), but as have generally managed to exploit most other toxic plants, why is it that they have been reported to avoid ferns? During field studies on the biology of ferns in the state of Veracruz, Mexico (March-April, 1976) it was noticed that many of the ferns showed a considerable amount of damage which was apparently due to arthropod feeding. It appeared, therefore, that entomologists and field ecologists may have either over-looked or ignored ferns as possible host plants for insect herbivores. Since earlier work had shown that two groups of secondary plant compounds, the tannins and cyanogenic glycosides (Cooper-Driver et al., 1977), are probably important in determining the extent of herbivore attack on a given fern species, the original observations were followed up with estimations of the amounts of these com- pounds in selected fern species. We also carried out an extensive literature search revealing a large number of references to ferns being used as host plants by arthropods, although it must be stressed that these records are not always clear as to whether the host plant was providing shelter or food. Few of these records relate to tropical flora and fauna, surprisingly in that insects are much more diverse in the tropics than elsewhere and doubtless constitute the major class of herbivorous animals (Janzen, 1975). From these preliminary observations on the degree and type of damage to Mexican ferns and from the records in the literature, it is concluded that the widely held assumption that very few arthropods feed on ferns, is not well founded. The chemical studies showed that while both tannins and cyanoge- nic glycosides were present in the ferns examined, their role as efficient feeding deterrents may not be as great as in other plant phyla (Swain, 1977), or as in temperate fern species (Cooper-Driver, 1976; Lawton, 1976). These findings are used to discuss the way in which ferns, during the course of evolution, have developed defensive strategies and in fact have co-evolved with their arthro- pod predators.

11. Materials and Methods I. Field Collection All field studies and collections were made in the state of Veracruz, Mexico near Jalopa in areas around Puente National, Misantla, Las Vigas and Perote, during March-April 1976. A total of six ecologically diverse sites were visited and every species of fern in the area was examined for insects or visible indications of damage due to insect feeding. Many of the specimans were collected, pressed, dried and examined in the laboratory. To determine the relative amount of damage to ferns and other plants, an "ecological plot", 2 by 4 m was laid out in a Liquidambar forest site at 1350 m altitude near Las Vigas. The location of the individual plants was recorded and a survey for insect damage made. Representative samples of each plant were pressed and dried and a rough estimate of the amount of damage obtained by photocopying the plants, cutting out the outline on the copy and weighing it (A, "intact plant") then removing from the copy the (white) area plainly showing damage (B) and weighing this. The percentage damage was then B/A x 100.

2. Chemical In~stigations Chemical tests were carried out using both fresh and dried material. The presence of a cyanogenic glycoside was determined using the method of Eyjolfsson (1970). Fresh frond samples in the field Biochemical and Evolutionary Aspects of Arthropod Predation on Ferns

were tested for the production of HCN by treating the material (ca. 1.5 g of terminal pinnae) with 2-3 drops of toluene in a sealed tube with a filter paper strip, which had been pre-treated with sodium picrate solution, suspended from the stopper and leaving the tube at room temperature for 24 h. Any change in the color of the papers from yellow to brown that was observed indicated that HCN had been released. Estimation for condensed tannins was carried out in the laboratory on dried specimans. The plant material was extracted twice with 80% boiling methanol and the extracts combined. Equal volumes of the extract and 5% conc. HCI in n-butanol were heated at 100" C for 40 min and the absorbtivity read at 530 nm (modification of Swain and Hillis, 1959). Quebracho tannin was used as the standard.

3. Literature Search

This was carried out using abstract literature from Biological Abstracts, Review of Applied entomol- ogy and Bibliography of Agriculture from 1930 up to the present time. In addition, the entomological libraries of the Connecticut Agricultural Experimental Station, Yale University and Harvard (Museum Comparative Zoology) were xanned for host plant data in taxonomic monogaphs or faunal surveys.

1. Survey for Arthropod Damage

Of the 137 fern species collected in the Veracruz area of Mexico, insect damage was noted in a total of 26 or approximately 19% of the total fern flora observed. Species for which damage was recorded are given in Table 1. The damage ranged from slight to that which appeared to be severe enough to have affected . the photosynthetic and reproductive capacity of the plant. Despite signs of considerable insect damage to the ferns in this tropical Mexican locality there : were very few arthropods actually found on the ferns at the time of collecting. This may be due to a predominance of nocturnal feeding, arthropod seasonality in the Mexican populations, collecting techniques, or some other factors. Subse- quent careful examination of pressed ferns from this locality revealed several different types of apparent arthropod damage. We have speculated, with each damage-type, which insect order might have caused the particular damage. These are as follows: a) small crescent notching of pinnules either at apex, base, or bases of several adjacent pinnules - Coleoptera (Curculionidae) (Fig. 1) ; b) large notching of pinnae-Lepidoptera, possibly Orthoptera (Figs. 2 and 5); c) skeletonization of upper surface of pinnule between the veins- Lepidoptera, Coleoptera (Fig. 3); d) shot-hole (upper surface), often concentrated but not always spherical -possibly Hemiptera (Homoptera) (Fig. 4); e) complete or par- tial elimination of several or many pinnules on one or both sides of the rachis- Lepidoptera (Fig. 6). Several fern species displayed fungal or viral damage and it may be that these pathogens are transmitted by arthropod vectors, i.e. aphids ' or leafhoppers, as is known to occur in other plants (Wood, 1967). The 8 square meter plot in the Liquidambar forest contained 3 species of ferns and 6 other flowering plant genera. The average percentage dafnage to the leaves of several selected individuals of these different taxa is recorded in Table 2. From these results it is apparent that ferns were preferred as a M.J. Balick et al.

Table 1. Fern species collected in Veracruz, Mexico on which arthropod damage was recorded, most of which were subsequently analysed for condensed tannins

Taxa % condensed Proposed type tannin of arthropod mg/g dry weight damage

Adiantum fenerum Sw. 8.40 C Anemin adiantifolia (L.) Sw. 7.80 a Asplenium momnfhes L. 0.90 d Blechnum varians (Fourn.) C. Chr. 5.10 a, b Ctenifis subincisa (Willd.) Ching - a a Cyrtomium juglandfolium (H + B) Moore - a Dicksonia ghiesbreghtii Maxon - a Dryopteris parallelogramma (Kze.) Alston 5.40 a, b, c, e Eluphoglosum latifoium (Sw.) J. Sm. 12.30 a, b Hypolepis reprms (L.) Presl. 0.14 a, d, e Lygodium mexicanwn Presl. 0.30 a Polypodium aureum L. 0.60 a Polypodium Ianceolarum L. - b. Polypodum loricewn L. 0.60 a, b Polypodium pectimtum L. - a, b Polypodium plebejium Schlect. +Cham. 5.10 a, b, e Polypodium puberulum Schlect. + Cham. 0.60 a, e Polystichum muricatum (L.) Fee 16.00 a, d Pteridium aquilinum (L.) Kuhn var. caudatum (L.) Sadeebeck 8.50 a + fungal damage Pteris dejlexa Link 1.80 e Tectaria heraclefolia (Wilid.) Underw. 8.10 a, d Thelypteris kunaii (Desv.) Morton 3.70 , a, b Thelypteris cheilanthoides 3.90 a Thelypteris normalis (C. Chr.) Moxley - a, b Thelypteris tetragona Sw. 5.0 , b Woodwardia martinezii Maxon. 9.0 e

These plants were not analysed for tannins food source, or at least showed a higher degree of predation, than the ac- companying angiosperms in this particular plot.

2. Chemical Investigations

Out of the 100 fern species tested for cyanogenesis only 3 gave a positive result: Cheilanthes intramarginalis, C. marginata and Notholaena aurea. Two other Cheilanthes species had previously been found to be cyanogenic (Harper et al., 1976). All ferns examined in the survey had the ability to produce condensed tannins although theactual amounts of tannins in tlie species analysed varied from 0.3%-16% mgms per gram dry weight (Table 1). Levels of tannins in the plant taxa of the ecological plot are given in Table 2. There was no apparent correlation between the percentage of damage to the leaf of frond material by feeding arthropods and the percentage of tannins per gram dry weight of plant material. Figs. 1-6. Illustrations of presumed insect dal I to fern species Fig. 1. Hypolepis repans (L.) Presl Fig. 2. Polypodium Ianceolarum L Fig. 3. Adianrum renerum Sw Fig. 4. Asplenium monanthes L. Fig. 5. Thelypteris rerragona Sw. Fig. 6. Reris dejlexa Link M.J. Balick et al.

Table2 Degree of arthropod damage recorded on the leaves of plants in a 8sq. meter plot in a Liquidambar forest, Veracruz, Mexico. These were subsequently analysed for condensed tannins

Taxa % estimated damage to % condensed tannin leaf or frond area by mg/g dry weight feeding arthropods

Ferns Blechnum oarians 12 Polystichum muricafum 16 Thelypferis cheilanfhoides 38

Angiosperms Begonia sp. I 5 Begonia sp. 2 7 Gunnera mexicana 4 Rumex sp. 0 Graminae 0 Labiateae 4 Rosaceae 6

Table3. Eaten and uneaten fronds of fern species collected from the same ,populations in Veracruz, Mexico analysed for their tannin content

Taxa % condensed tannin mg/g dry weight

Eaten ' Uneaten

Adianfum tenenun Sw. 8.4 r 9.9 Lygodium mexicanum Presl. 0.3 0.35 Pteridium aquilinum (L.) Kuhn 8.5 9.3 Pteris deflexa Link 1.8 0.6 Tectaria heracleijblia (Wad.) Underw. 8.0 3.9 Thelypteris kunfhii (Desv.) Morton 3.7 3.9

Five populations of different fern species were selected at random and samples of eaten and uneaten fronds were collected for each species within the same population. The eaten and uneaten fronds were then analysed for their tannin content. Results are given in Table 3. There was no over-all correla- tion between % tannins in eaten and uneaten fronds, although there was a tendency in three of the species, for the levels to be slightly higher in uneaten than in eaten material.

3. Literature Data

The results of the literature survey are tabulated in Appendix 1. The data are alphabetically arranged (by Orders, Families, Genera and Species) according to the nomenclature of Brues et al. (1954). In addition to the fern-arthropod Biochemical and Evolutionary Aspects of Arthropod Predation on Ferns records, ecological information has been recorded wherever possible after the arthropod name. Those insect species known to feed on several non-fern hosts are indicated as being polyphagous (P), whereas those with only a few non-fern hosts as oligophagous (0). The mode of feeding is also indicated when known, i.e. sporangia feeders (s), gall-formers (g), root-stock or rhizome feeders (r), miners or borers (m), and decaying frond feeders (d). The remainder are assumed to be primary frond feeders. Wherever a record is questionable either by the au- thor's admission, or because it is a secondary predator or parasite (or otherwise), it is cited as a fern associate (assoc.) which is indicated after the specific arthro- pod binomial. For all fern-arthropod relationships listed, the locality and au- thor's name with a reference are given for more detailed reading. It must be stressed that Appendix 1 contains only recorded data and it is inevitable that there will be some cases of misidentification or synonomy of arthropods and their fern hosts, particularily in the case of older records. In addition this table does not claim to give a complete list of all insects recorded as being present on ferns; nevertheless this is the most comprehensive treatment that has been produced up to the present .time and hopefully will serve as the basis for further compilations. From an analysis of our data there appears to be a more or less even distribution of types of feeders: root feeders (3.3%), spore feeders (2.4%), miners or borers (5.9%), gall formers (4.3%) and decay feeders (4.0%). 73% are restricted to ferns whereas 27% are polyphagous or oligophagous. This indicates that most fern feeders must have had to specialize in order to have co-evolved with the chemistry of ferns.

IV. Discussion

No previous paper has presented evidence of arthropod-fern associations on such a broad scale (Appendix 1). The only extensive listing of fern insect pests is by Docters Van Leeuween (1938) in Verdoorn's Manual of Pteridology. Other surveys have been restricted to regions such as Hawaii (Swezey, 1922), North American fern aphids (Robinson, 1966), Western Hemisphere fern mites (De Leon, 1966), or to insects present on a particular fern; Simmonds (1967), Wiec- zorek (1973), Lawton (1976) and Kirk (1977) all have recorded insects present on bracken, Pteridium aquilinum. Hoshizaki (1975) provides a common name list of fern pests; however, these are primarily at the ordinal level. Approximately 420 species of arthropods have been recorded from fern hosts in Appendix 1, together with 44 fern associates and several references to unnamed insect species. It is not certain as to exactly which insects were actually feeding on the ferns rather than using them as a refuge or a transient resource; neither do we know whether the arthropods and ferns have been correctly identified in the older records. The list cannot be truly representative for all fern genera since over 22% of the records refer to the economically important fern bracken, the most widely distributed and probably the best known of all pteridophytes (Perring and Gardiner, 1976). It might be expected that there would be more data on arthropods feeding on ferns in the tropics M.J. Balick et al.

than in the temperate zones especially since pteridophytes are more abundant and more diverse in tropical areas. Nevertheless from this compilation several inferences can be made. Of the 420 arthropod species recorded on ferns, 59% constitute the more advanced Holometabola: the largest groups in number of species, are in the orders Co- leoptera, Hymenoptera and Lepidoptera. The Coleoptera exhibit a scattered diversity of fern feeders and the weevils, the most diverse family of insects, are probably quite widely associated with ferns all around the world, especially in the tropics. Zimmerman (1957) lists many Miocalles weevils from Oceania and demonstrates the high affinity of these insects for ferns. The ferns collected in Mexico also apparently showed a significant amount of weevil damage (a, Table 1). Vanin (1976) considers the Gondwanian belid weevils to be primitive not only because of their morphology but also their phytophagy of ferns and gymnosperms. There is a predominance of primitive sawflies within fern feeding Hymen- optera. Sawflies form a dominant group of the restricted fern feeding commu- nity throughout the Holartic region. Lawton (1976) found only one of the 43 "common" British seed plants had more sawflies than bracken. In contrast to most other groups of arthropods, it seems quite likely that these wasps may have co-evolved with ferns before angiosperm dominance in the late Cre- taceous. Among the Lepidoptera, the Noctuidae are the most numerous fern feeders with the genera Callopstria and Papaipema containing many specialist fern feed- ing species. Lepidoptera are considered to be the most advanced of holometabo- lous insects, known since the Cretaceous (Ehrlich and Raven, 1964; Mackay, 1970). Although Ehrlich and Raven have proposed that Lepyoptera diversity has elaborated along with the dicotyledons and that those inembers feeding on non-dicots show secondary adaptation and are probably more advanced, our records show that it is the primitive Lepidoptera, moths such as the Hepia- lidae rather than butterflies and higher moths, that are recorded as being present on ferns. The hemimetabolous Hemiptera (especially the Hornoptera) are also appar- ently important fern feeders. Aphids (Aphididae) are able to utilize a large number of divergent species as host plants suggesting that they may be able to avoid some of the chemical defenses of most plants due to their specialized piercing-sucking method of feeding. Aphids may thus not have to contend with plant toxins in high concentrations since such compounds are usually more or less absent from the phloem transport stream or occur there in much lower concentrations than in the rest of the plant (Van Emden, 1972). Eastop (1973) records 29 species of aphids, from several different aphid groups, restricted

' to ferns. He also notes that aphids, unlike most arthropods, are' less abundant and more polyphagous in the tropics, which perhaps explains why we did not find much evidence of aphid predation in our Mexican samples (d, Table 1). The oldest arthropod order recorded on extant ferns is the Orthoptera which has fossil representatives in the Carboniferous (Smart and Hughes, 1973), how- ever, there are very few recorded on today's ferns. Other arthropod orders, Biochemical and Evolutionary Aspects of Arthropod Predation on Ferns with more recent fossil history, i.e. Diptera (early Mesozoic) and Thysanoptera (Permian) also contain restricted fern feeders. The wide scattering of recorded fern arthropod hosts does not reveal an obvious association with primitive or more advanced arthropods. Plants have apparently responded to insect attack in various ways since they became estab- lished on land (Kevan et al., 1975; Swain, 1978), and thus insects and plants must be viewed as two co-evolving, competing and often mutually dependent biochemical systems (Ehrlich and Raven, 1964). In any given ecosystem the fates of the various guilds of herbivorous insects and plants are chemically intertwined (Whittaker and Feeny, 1971 ; Gilbert and Raven, 1975; Swain, 1977). Fossils of arthropods found in the early Devonian were arachnids with mouth- parts which could pierce plant stems and spores, and in this period many insects fed on spores, spore protoplasts and plant sap. The evolution of winged insects in the late Devonian was accompanied by an. increase in arborescent forms of plants and in the complexity of certain groups of secondary plant compounds. The ability to produce chemically more resistant spores and cell walls, to synthesize lignin and general purpose antibiotics and feeding deterrents was as important in the evolution of plants as changes in their anatomy and morphology (Swain, 1978). The condensed tannins are one of the most important of all plant chemical defenses; not only are they potent anti-fungal, anti-bacterial and even antiviral agents but they bestow on plants, that have the ability to synthesize them, a powerful feeding deterrent to many herbivores (Swain, 1977, 1978). All ferns, with the exception of the Ophioglossales, which have a comparatively recent fossil history, have the ability to synthesize tannins. As there is a higher propor-: tion of species containing proanthocyanidins in the Filicales than in the lower tracheophytes, gymnosperms, or angiosperms, it is likely that proanthocyanidins ' arose as the first soluble anti-fungal and insect feeding deterrent but were later reinforced or replaced in angiosperms by ellagitannins or by more selective terpenoid or other feeding deterrents. Although correlations have been reported between amounts of tannins and degree of arthropod feeding inhibition (Feeny, 1970; Lawton, 1976; Rhoades and Cates, 1976; Cooper-Driver et al., 1977); the data were from plants growing in temperate climates. Results also vary according to the time of year, tannin levels usually being higher in deciduous temperate plants towards the end of the growing season. As the tannin levels rise, the extractable protein content drops decreasing the nutritional value of the plant as a source of food. In the tropics, however, with perhaps more rapid co-evolution of both herbivore and host, it may be that insects have developed a higher degree of tolerance to tannins which may not be such an effective means of defense. This may explain why many of the ferns with high tannin levels were still subject to insect attack. Ferns do however possess a number of other defensive compounds (Swain and Cooper-Driver, 1973). The ecdysones or insect molting hormones are found inmany ferns. It has been assumed that the role of phytoecdysones is to induce metamorphosis in arthropods thus causing lethal anomolous development (He- rout, 1970). However laboratory experiments using a variety of insects i.e. Musca, Cynthia, Hyalophora, Manduca, Trichoplusia, Tribolium, Blatella, Thermobia and M.J. Balick et al.

Schistocerca (Robbins et al., 1968; Williams, 1970; Hendrix, 1977) have not been able to substantiate the effects of ecdysones under natural conditions using whole plants. The phytoecdysones occur more commonly in those species of ferns which are considered to be advanced e.g. Polypodium and Pteridium and which, by and large, also possess a wider range of other potentially deterrent toxic compounds such as the bitter sesquiterpenoid lactones and cyanogenic glycosides. The cyanogenic glycosides break down under the action of B-glucosidase to produce hydrogen cyanide known to be toxic to many herbivores (Jones, 1973; Cooper-Driver and Swain, 1976). Although these compounds are not very common in ferns, occurring in approximately 3% of species (Harper et al., 1975), nevertheless they are effective feeding inhibitors. Of the 2 Cheilanthes and 1 Notholaena species that were found to be positive for HCN production, in this study; none of these had been attacked by insects. As hydrogen cyanide production varies with age, season and a number of environmental factors, . much more work still needs to be done before we can ascertain the importance of this particular group of chemicals to the total ecosystem.

V. Conclusion

It is apparent from the data presented that many more arthropods are in fact present on ferns, than has previously been thought, and that many have over- come the fern's chemical defense systems to utilize them as food, particular~ in the tropics where coevolution proceeds at a more rapid rate. A study of such interactions with ferns provides a good system for examining theories of coevolution since they are at the juQction of primitive vascular plants and the more specialized seed-bearing plants, and have a long fossil record from the late Devonian (Banks, 1972). A systematic search is needed to examine the species of insects which feed on different taxa of ferns and in what frequency; to determine seasonality of feeding correlated with changes in plant and insect chemistry, to analyse secondary plant compounds present in temperate and tropical fern species, and finally controlled feeding tests are needed to determine insect preference when a selection of suitable hosts is offered. It is hoped that this paper will stimulate direct field observations in both temperate and tropical areas. Data from all such observations could begin to shed new light on the important biological phenomenon of coevolution.

Acknowledgemenrs. We thank Drs. Jesus Dorantes L., Escuela de Biologics, Universidad Veracru- zana, Jalopa; Arturo Goma-Pompa, Instituto de Investigaciones Sobre Recursos Bioticos; Pilar Fernandez and Jose Sarukhan, Universidad National Autonoma De Mexico for help with the field experimenis in Mexico which were carried oui in conjunction with the course Biolog); 247, Harvard University with Iinancial support from the Atkins fund. We would also like to thank Drs. R. and A. Tryon from the Gray Herbarium, Harvard' U., C.L. Remington, Yale U., J.G. Franclemont, Cornell U., T. Swain, Boston U. for helpful discussions. Many people have helped in the compilation of the arthropod fern list. Most of their names are listed in Appendix 1 as personal communications (p.c.) or in litt. as well as D. Johnson, Ohio State and J. Slater, U. Conn. Biochemical and Evolutionary Aspects of Arthropod Predation on Ferns

Appendix 1 Arthropod-Fern Herbivores

Arthropod taxon Fern host Locality Reference

ACARINA Chamobatidae: Chamobates sp. Britain Lawton, 1976 (= ?associate) Eriophyidae: Eriophes (=Phyrop) Nephrolepis spp. Old World D. van Leeuwen, 1938 paurop Nalepa (g) Tropics E. pterides Moll. (g) ferns; P. aquilinum Brit. Isl. D. Van Leeuwen, 1938 E. SP. (g) Dicranopteris sp. USA Jeppson et al., 1975 Erwphyhrum sp. ,(O) P. aquilinwn Portugal Tavares, 1930 Eriophyid sp. (g) Angiopteris mcta Old World D. Van Leeuwen, 1938 (Fors:) Hoffm. Trop. Phyllocoptes dimorphus P. aquilinum Calif. Kiefer, 1940 Kiefer Tarsonemidae : Eotarsonemur rugosus ferns; Polypodium sp. Trinidad DeLeon, 1966 De Leon Hemitarsonemus tepid- ferns; Pteris spp. ;Aspleniwn Calif., Minn., Pritchard, 1951; ariorum Warb. bulbifem; Polysrichwn sp. France; Engl. Schneider, 1966; (greenhouse) De Leon, 1966 Tenuipalpidae: Brevipalpus sp.-complex Nephrolepis biserrata; Puerto Rico DeLeon, 1%6 Polypodiwn pectinatum x plwmclwn tree fern; Cyathea nigre- Puerto Rico, DeLeon, 1966 scens; Alsophifa borinquensis Jamaica T. guamensis Baker Aspknium nidus Guam DeLeon, 1966 T. lygodi De Leon Lygodiurn mlubile Jamaica De Leon, 1966 T. meekeri DeLeon BIechmun sernrlotum; Ha., Mex. Polypodium sp. Trinidad COLLEMBOLA (=?assoc.) Sminthuridae: Bourletielln viridescens P. aquilinwn Brit. Is., Eur. Lawton, 1976 Stach, S. Gisin (F') Dicyrtomidae : Dicyrtoma sp. P. aquilinwn England Lawton, 1976 ~ntodobryidae: Lepidocyrtur pallidus Adiantwn; Microlepia spp. Belgium ~e&ens,1967 Reuter (=L. cyaneus Nephrolepis; Pellea; Pteris Tullb.) CORRODENTIA Perientomidae: Cyptophania hirsuta Cibotium ;ferns Hawaii Zimmerman, 1948 Banks (P) M.J. Balick et al.

Appendix 1 (continued) -- Arthropod taxon Fern host Locality Reference - CORRODENTIA (continued) Psocidae : Psocus distinguendus Sadleria Hawaii Zimmerman, 1948 Perkins (P)

COLEOPTERA Alticidae ( =Chrysomelidae) : Apthom sp. P. aquilinum New Guinea Kirk, 1977 Manobia sp. 1 P. aquilinum New Guinea Kirk, 1977 M. sp. 2 P. aquilinum New Guinea Kirk, 1977 Anthribidae: Homocloeus n. sp. 1 tree ferns Jamaica Valentine (in litt.) Homocloeus n. sp. 2 tree ferns Jamaica Valentine (in litt.) . Ormiscusfloridanus (Leng) ferns S. Fla. Valentine (in litt.) Phaenorheriopsb n. sp. tree ferns Jamaica Valentine (in litt.) Trigonorhinus ronmdatw ferns Mass. Valentine (in litt.) Leconte Carabidae: Several native Hawaiian tree ferns Hawaii Swezey, 1922 SPP. (4 (assoc.) Cerambycidae: ?Cornallb sp. P. aquilinum New Guinea Kirk, 1977 Sybra sp. P. aquilinum New Guinea Kirk, 1977 Tmesisfernussp. 1 P. aquilinwn New Guinea. Kirk, 1977 T. sp. 2 P. aquilinum New Guinea Kirk, 1977 Unidentified sp. 1-sp. 5 P. aquilinum New Guinea Kirk,' 1977 Cryptophagidae: Unidentified sp. (s) Cysfopterisfragilis Chile (Juan J. Lawrence (p.c.) Fernandez Is.) Curculionidae: Bagous sp. saluinia sp. India Vorghese et al., 1972 Baris arropolifa Lea P. aquilinwn New Guinea Kirk, 1977 Brachyrhinus ( =Otior- ornamental ferns USA Dodge et al., hynchus) sulcarus (F.) (P) 1948 (r and fronds) Cryptorhynchinae ?sp. P. aquilinum New Guinea Kirk, 1977 Cyriobagous singuloris Salvinia Trinidad, No. Thomas, 1975 Hulst. S. Amer. Dryophthorus insignis Cibotium spp. Hawaii Swezey, 1922 Sharp (4 D. modestus Sharp (d) Cibotium spp. Hawaii Swezey, 1922 D.pusillus Sharp Cibotium (2 or 3) Hawaii Swezey, 1922 Gromihrs gracilipes Sharp ? Blechnum N. Zealand May, 1973 Heteramphus filicwn tree ferns Hawaii Swezey, 1922 Perkins Biochemical and Evolutionary Aspects of Arthropod Predation on Ferns

Appendix 1 (continued)

Arthropod taxon Fern host Locality Reference

H. swezeyi Perkins (m) EIaphoglossum reticulatwn Hawaii Swezey, 1922 E. micradeniwn; E. gorgo- newn ;E. squamoswn Heteramphus wollasfoni tree ferns Hawaii Swezey, 1922 Sharp (0) Homolocerus lyciformis Polystichwn ( = Ruhmora) Brazil (German) denriculatwn Pr. ; several Cyatheaceae Homolocerus nigripennis tree ferns Bohemian H. plawnanni Voss Polystichwn denticulatum Brazil Vanin, 1976 Megacolabus bifurcatus Blechnwn capense (L.) N. Zealand May, 1973 May 6) Miocalles (=Micro- ferns Austral Islands Zimmerman, 1957 cryptorhynchus) abditiceps (Tap 1s.) (Zimmerman)' (0) M. abnormis (Zimm.) Pterb sp. Austral Islands Zimmerman, 1957 (Rapa Is.) M. ambiguus (Zimm.) fern Society Islands Zimmerman, 1957 (Tahiti Is.) M. andersoni (Zimm.) (P) ferns Rapa Is. Zimmerman, 1957 M. angustatus (Zimm.) Cyathea sp. Tahiti Is. Zimmerman, 1957 M. bicolor (Zimm.) Cyathea sp. Rapa Is. Zimmerman, 1957 M.brevis (Zimm.) (P) ferns Tahiti Is. Zimmerman, 1957 M. caudahcr (Zimm.) Blechnwn sp. Society Islands Zimmerman, 1957 (HuaIhe) M. chaetectetoroides ?ferns Rapa Is. Zimmerman, 1957 (Zimm.1 (PI M. confmus (Zimm.) fern Tahiti Is. Zimmerman, 1957 M. conwxus (Zimm.) (0) fern Tahiti Is. Zimmerman, 1957 M. cookei (Zimm.) Pterir sp. Austral Islands Zimmerman, 1957 (Raivavae Is.) M. cum(Zimm.) fern Rapa Is. Zimmerman, 1957 M. dircretw (Zimm.) Blechnum, ?ferns Tahiti Is. Zimmerman, 1957 M. exilk (Zimm.) fern Tahiti Is. Zimmerman, 1957 M. fosbergi (Zimm.) Cyarhea sp. Tahiti Is. Zimmerman, 1957 M. fowawntris (Zimm.) Aspleniwn nidus Rapa Is. Zimmerman, 1957 M.fulgidus (Zimm.) fern Tahiti Is. Zimmerman, 1957 M. gracilis (Zimm.) fern Austral Islands Zimmerman, 1957 (Tubuai Is.) M. hirtw (Zimm.) fern Austral Islands Zimmerman, 1957 (Tubai Is.) M. impressicollk (Zirnm.) Asplenium nidus Rapa Is. Zimmerman, 1957 M. irregularis (Zimm.) Cyathea sp. Rapa Is. Zimmerman, 1957 M. irroratus (Zimm.) fern Tahiti Is. Zimmerman, 1957 M. mangaoae (Zimm.) Cyathea sp. Rapa Is. Zimmerman, 1957 M.mangarevae (Zimm.) Aspleniwn nidus Mangareva Is. Zimmerman, 1957 M.J. Balick et al.

Appendix 1 (continued)

Arthropod taxon Fern host Locality Reference

COLEOPTERA (continued) Curculionidae (continued): M. montemgus (Zimm.) fern Tahiti Is. Zimmerman, 1957 M. nitidus (Zimm.) (0) Asplenium nidus Rapa Is. Zimmerman, 1957 M. obesus (Zimm.) (0) ferns Rapa Is. Zimmerman, 1957 M. orofenae (Zimm.) Cyathea sp. Tahiti Is. Zimmerman, 1957 M. paenulatus (Zimm.) (0) Aspleniwn nidus; Cyathea Rapa Is. Zimmerman, 1957 sp., ferns M. parvur (Zimm.) Dryopreris sp. Austral Islands Zimmerman, 1957 (Rurutu Is.) M.pervirus (Zimm.) fern Tahiti Is. Zimmerman, 1957 M.proximus (Zimm.) (0) Cyathea sp. Rapa Is. Zimmeman, 1957 M. punctipennis ferns Rapa Is. Zimmerman, 1957 (Zimm.) (P) M. pusillus (Zimm.) (P) Aspleniwn nidus Rapa Is. Zimmerman, 1957 M. raivavaensir Preris sp. Austral Islands Zimrnerman, 1957 (Zimm.) p) (Raivavae Is.) M. rurutuensis (Zimm.) Dryopteris sp. Rurutu Is. Zimmerman, 1957 M. setigem (Zimm.) (P) Cyathea Rapa Is. Zimmerman, 1957 M. setulosus (Zimm.) Aspleniwn nidus; Rapa Is. Zimmerman, 1957 Cyathea spp. M. similis (Zimm.) (P) ferns Tahiti Is. Zimmerman, 1957 M. sparhifer (Zimm.) Asplpnium niduc Rapa Is. Zimmerman, 1957 M.tenuis (Zimm.) (P) Cyarhea sp. Rapa Is.. Zierman, 1957 M. tesraceu (Zimm.) fern Tubuai Is. Zimmerman, 1957 M. rhoracius (Zimm.) (0) Asplenium nidus; ?ferns Rapa Is. Zimmerman, 1957 M. tubuaiensis (Zimm.) fern Tubuai Zimmerman, 1957 M. rumidus (Zimm.) Aspleniwn nidus Rapa Is. Zimmerman, 1957 M. mrians (Zimm.) (P) Cyarhea spp. Rapa Is. Zimmerman, 1957 M. uenrralis (Zimm.) Aspleniwn nidus Rapa Is. Zimmeman, 1957 Neosyagrius cordipennis Adiantum sp. Australia . Anonymous, 1940 Lea Notacalles piciuentrir Polystichum vestitum Canada (Camp Kuschel, 1964 (Brouu) bell Is., B.C.) Oodemas aenescens Ciborium spp. Hawaii Swezey, 1922 Bohman (d) (P) 0. brunneum Perkins (d) Preris sp. Hawaii Swezey. 1922 Pentarthrum prolixwn Cibotiwn spp. Hawaii Swezey. 1922 Sharp (dl Pseudolus longulus (Boh.) Cibotium chamissoi KauIf. Hawaii Swezey, 1922 (dl (0) Rystheus notabilis Broun ? Blechnum N. Zealand May, 1973 Stenopelmus rufuuzrur Gy. Azolla S. &W.USA, Richerson et al., 1%7 Europe (intro- duced) Strophosmos sp. Europe Biochemical and Evolutionary Aspects of Arthropod Predation on Ferns

Appendix 1 (continued)

Arthropod taxon Fern host Locality Reference

Syagrius spp. Adiantum spp. Australia Linssen, 1959 S. fulvitarsis Pascoe wild and cultivated ferns; Australia, Ha- Anonymous, 1940 Sadleria cyatheoides: waii (introduced) Asplenium sp. S. intrudem Waterhouse ferns Brit. Is. (intro- Linssen, 1959 duced from Australia) Trichophthalmus milt- Blechnum chilense; Chile omerus Blanchard PoIysfichum sp. Elateridae : Corymbites nigricornis ferns Brit. Is. Linssen, 1959 Pz-1 Dolopius marginatus P. aquilinwn Engl. Lawton, 1976 (L.) (assoc.) Eumolpidae (=Chrysomelidae) : Unidentified sp. P. aquilinum New Guinea Kirk, 1977 Galemcidae (=Chrysomelidae): Mimastra Iimbata Baly Adiantum sp. Thailand Cantelo, 1965 Helodidae: Cyphon padi (L.) (assoc.) P. aquilinwn Engl. Lawton, 1976 C. variabilis Thunberg P. aquilinum Engl. Lawton, 1976 (assoc.) Lathrididae : Cartodere ruficollis P. aquilinum Engl. Lawton, 1976 Marsham Melasidae (=Eucnemidae) : Dirhagus pygmaeus (F.) P. aquilinum Engl. Lawton, 1976 (assoc.) Mycetophagidae: Unidentified sp. Cystopterisfragilis Chile (Juan J. Lawrence (p.c.) Fernandez Is.) Nitidulidae : Nesapferus monticola Cibotiwn menziesii Hawaii Swezey, 1922 (Sharp) (dl Proterhinidae: Proterhinus longulur Cibotiwn chamissoi; C. Hawaii Swezey, 1922 Sharp (dl menziesii P. pteridis Perkins (m) Pteris sp. Hawaii Swezey, 1922 P. sharpi Perkins fern (non-arboreal) Hawaii Swezey, 1922 P. spp. ?ferns Hawaii Swezey, 1922 Ptinidae : Holcobius hawaiiensis tree ferns Hawaii Swezey, 1922 Perkins (assoc.) Scarabaeidae: Phyllopertha horticola P. aquilinwn Brit. Is.; Eur. Lawton, 1976 (L.) P) Wieczorek, 1973 70 M.J. Balick et al.

Appendix 1 (continued)

Arthropod taxon Fern host Locality Reference

COLEOPTERA (continued) Scolitidae: Hypothenemus eruditus ferns Zaire Schedl, 1962 Westw. H. pusillus Egg. ferns Zaire Schedl. 1962 PoeciiQs preridophyrae P. aquilinum New Guinea Schedl, 1962 Schedl (m) Stephanoderes perhispidus ferns Zaire Schedl, 1962 Egg.

DIPTERA Agromyzidae: Agromyza undescribed Marattia doughsi Hawaii Swezey, 1922 SP. (4 Phytoliriomyza hilarefla Polypodium; P. aquilinum; Brit. Is., Eur., Lawton, 1976 (zett.1 (4 Osmunda N. Amer. Wieczorek, 1973; (= Pteridomyza; Phytobia; Hering, 1951 Praspedomyza; Agromyza) P. preridii Spencer (m) P. aquilinum Brit. Is. Lawton, 1976 Phytomyza scolopenrfii several Poiypodiaceae ?Europe Hering. 1951 R.-D. (m) Asplenium; Athyrium ; Hering, 1951 Blechmim Anthomyiidae: Chirosia (=Pycnoglossa) Dryopreris , Engl. Wieczorek, 1973 aberrans Collin C. aibifons Tiens. P. aquilinum Engl., Eur. Lawton, 1976; Wieczorek, 1973 C. albitarsis Zett. (m) P. aquilinum Brit., Eur., Asia Lawton, 1976; Wieczorek, 1973 C. betuieti Ringdahl (g) Athyrium; P. aquilinum Europe Lawton, 1976; Wieczorek, 1973 C. crassiseta Stein (m, r) P. aquilinum Eng., Eur. Simmonds, 1964; Wieaorek, 1973 C.flauipennis Fallen (m) P. aquilinum Britain Lawton, 1976 Chirosia hystricina Ron- P. aquilinum; Asplenium; Britain, Eur., Lawton, 1976; dani (m) (= C. cinerosa A thyriwn; Blechnwn; Cyst- N. Amer. Wieczorek, 1973 Stein; C. hystrix Brischke; opteris; Dryopteris; Matte- C. setifemur Ringdahl) uccia ; Osmunda; Polyp- odium; Polystichum C. parvicornis Zett. (m) Asplenium; Dryopteris; Brit. Is., Eur. Lawton, !976; P. aquilinum Wieczorek, 1973 Hylemya (= Choro rophilia) Polypodiaceae; ferns Europe D. Van Leeuwen, 1938; signata Brisc. (g) Simmonds, 1967 Cecidomyiidae: Cecidomyia sp. (r-g) Cystopteris fragilis USA Felt, 1940 Dasyneura firicina P. aquilinum Eng., Eur. Lawton, 1976; (Kieff.) (g) Wieczorek, 1973 Biochemical and Evolutionary Aspects of Arthropod Predation on Ferns

Appendix 1 (continued)

Arthropod taxon Fern host Locality Reference

D. pferidicola Kieff. (g) P. aquilinwn; ferns Eng., Eur. Lawton, 1976; Wieczorek, 1973 ''a few spp." (g) ferns Europe D. Van Leeuwen, 1938 Drosophilidae : Drosophila apicipwcta ferns; Sadleria Hawaii Hardy, 1965 Hardy D. nigella Hardy (assoc.) ferns Hawaii Hardy, 1965 D. notha Bock (g) P. aquilinum New Guinea Kirk, 1977 D. sadleria Bryan (m) Sadleria cyafheoides Hawaii Swezey, 1922; Hardy, 1965 Pipunculidae (assoc.-parasite): Pipwculus alienus (Hardy) ?leafhoppers over Hawaii Hardy, 1964 Cibotiwn and Nephrolepis P. amplus a aid^ over Nephrolepis sp. Hawaii Hardy, 1964 P. cornutus (Hardy) over Nephrolepis sp. Hawaii Hardy, 1964 P.filicicolus Hardy over Nephrolepis exaltata Hawaii Hardy, 1964 P. jumtor Perkins ?leafhoppers over Nephro- Hawaii Hardy, 1964 lepis and ferns P. megameris Hardy over Cibofium Hawai Hardy, 1964 P. obscuratus Hardy over Cibotium chamissai Hawaii Hardy, 1964 K. (assoc. with Neso- phrosyne sp. and Neso- sydne ipomoeicola Kirk) P. timerlakei Hardy over Cibotium chamissai Hawaii Hardy, 1964 (assoc. with Nesophrosyne and Nesosdyne ipomoeicola) P. swezeyi Perkins leafhoppers on Nephrolepis Hawaii Hardy, 1964 Sciaridae: Sciara sp. (s) "young ferns" Netherlands Augustijn et a]., 1935

HEMIFTERA: HETEROFTERA Anthocoridae (assoc.): Lasiochilus denigratus Cibotiwn chamissai Hawaii Zimmerman, 1948 (White) (4(0) Unidentified sp. Cyathea Ethiopia Limavuori (in litt.) Lygaeidae: Drymus ? brunneus P. aquilinwn Engl. Lawton, 1976 Sahlberg Germahu costalis Van Cyafhea;ferns Marquesas Is. Van Duzee, 1935 D- (0) G. fuliginosus Van Cyathea; ferns Marquesas Is. Van Duzee, 1935 Duzee (P) G. infans Van Duzee (P) ferns Marquesas Is. Van Duzee, 1935 Metrarga nuda White Cibofiwn Marquesas Is. Zimmerman, 1948 (4(0) M. obscura'Blackburn tree fern Hawaii Usinger et a]., 1959 M.J. Balick et al.

Appendix 1 (continued)

Arthropod taxon Fern host Locality Reference

HEMIPTERA: HETEROPTERA (continued) Lygaeidae (continued): Micrymenus kuscheli Dryopterir inaequalijolia; Chile Kormilev, 1952 Kormilev (0) Polystichwn vesticum Neseir hiloensb inter- misc. ferns Hawaii Zimmerman, 1948 medius Usinger (0) Nysiw blackburni ferns Hawaii Zimmerman, 1948 White (0) N. communis Usinger (P) Cibotiwn menziesii; Hawaii Zimmerman, 1948 Sadleria N.jacobeae Strobl Pterb sp. Austria Strobl, 1900 N. liliputanus Eyles and fern N. Zealand Eyles et al., 1969 Ashlock (0) ' N. rubescens White ferns Hawaii Zimmerman, 1948 Oceanides montivagus Sadleria Hawaii Zimmerman, 1948 (Kirk) Ozophora pauescens ferns Fla. ; W. Indies Torre-Bueno, 1946 (Distant) (P) 0. trmotata Barber ferns Ha. Torre-Bueno, 1946 Sephom criniger ferns; Sadleria Hawaii Zimmeman, 1948 (white) (PI Styngnocoris sabulosus Pteris sp. Lunsden, 1939 Lumsden Midae: Bryocorinae many unde ferns; "soft ferns" Africa D. Leston (p.c.) scribed spp. Bryocorir preridis Aspidiwn /ilk-mas; Brit. Is., Eur. Lawton, 1976; (Fallen) 6) A fhyriwn /ilk-femina; Wieczorek, 1973; Dryopteris; Lastraea Butler, 1923 dilotata; L. ?oreopteris; Polysfichwn; Asplenium ; Pteridium aquilinwn Calocorb sexguttatus Athyriwn filix-femina Brit. Is. Butler, 1923 Fab. (assoc.) Ceratocapsus setosus ferns 111. Knight, 1941 Reuter Deraeocorir ( = Camp P. aquilinwn Brit. Is. Butler, 1923 tobrochis) Iutescens (Schill.) (assoc.) Dicyphus globulijer . A fhyriwnfilix-femina; Brit.. Is. Butler, 1923 (Fallen) (assoc.) P. aquilinwn Felirarus glabratus ferns Australia Woodward et al., 1970 (Motsch.) bpur (= Onychwnenus) ferns Brit. Is. Butler, 1923 decolor (Fallen) (P) Biochemical and Evolutionary Aspects of Arthropod Predation on Ferns

Appendix 1 (continued)

Arthropod taxon Fern host Locality Reference

Lygocoris paburinus (L.) Aspidium Brit. Is. Butler, 1923 (assoc.) Lygus cervinus Herr.- ferns Brit. Is. Butler, 1923 Sch. (0) (assoc.) L. indistinctus Taylor P. aquilinum Brit. Is. Butler, 1923 Mocrolophus nubilus P. oquilinm Brit. Is. Butler, 1923 Herr.-Sch. (assoc.) Mecomma ombulans Aspidiwn filix-femina; Brit. Is. Butler, 1923 Fallen (P) Polypodium dryopteris Miris sp. ferns Africa R. Limavuori (p.c.) Monaiocoris omericana Osmwrdn cinnomeo ferns N. America Wagner et al., 1952 (Wagner and Slater) Monalocoris filicis (L.) (s) Aspidium filix-mar; As- Brit. Is., Eur. Butler, 1923; Lawton, plenium ;Athyriwn Nix- 1976; Wieczorek, 1973 femitia; Lastraeo dilaro- la; L. oreopteris; Poly- stichm; Aspidim spin- ulosum M. (= Sthemrusoides) P. oquilinrun Nigeria Limavuori, 1975 punctipemis Linnavuori Orthotyhu kassandra Sadleria Hawaii Zmmerman, 1948 (Kirk.) (PI Pseudoclerah moroi creeping fern Hawaii Zimmerman, 1948 Kirk. (assoc.) Stenodema hokatm P. aquilinum Brit. Is. Butler, 1923 (Fab.1 (PI Tenthecoris bicolor ornamental ferns France Schneider, 1966 kott (PI Nabidae : (assoc.) Nabb blockburni ferns Hawaii Zimmerman, 1948 (White (PI N. curtipemis Blackburn tree ferns Hawaii Zierman, 1948 N. Iusciosw White (P) ferns Hawaii Zimmerman, 1948 N. pele (Kirk.) (0) Cibotium Hawaii Zimmennan, 1948 N. siloestris (Kirk.) (0) Cibotium Hawaii Zimmerman, 1948 Pentatomidae: (assoc.) Aelia acuminalo L. (P) ferns, Osmtmdo regolis Brit. Is. Butler, 1923 Jalla dwnosa L. P. aquilimun Brit. Is. Butler, 1923 Zicrono caeruleo L. ferns Brit. Is. Butler, 1923 Reduviidae (assoc.) : Empicorb rubromoculatus ferns Hawaii Zimmerman, 1948 (Blackburn) (P) Nesidiolestes . fern Hawaii Gape et al., 1975 Tingidae: Corythuchopodi Drake P. aquilinum Brit. Columbia Leech, 1944 M.J. Balick et al.

Appendix 1 (continued)

Arthropod taxon Fern host Locality Reference - - HEMIPTERA: HOMOPTERA Aleyrodidae : Aleuroplatus kewensis Anemia sp.; Diplnziwn Britain (green- Trehan, 1937 Trehan prolfenun ;Dryopreris house) jlaccida; Nephrodiwn con- jluens; Oleandra africana Aleuropreridb douglasi Cyclosoncs dentatus; West Africa Mound, 1961 Mound Pteris togoensis A. easropi Mound ?fern Ghana Mound, 1961 A. hargreawsi Mound Bush Fern Sierra Leone Mound, 1961 A. jamesi Mound Pterb togoensis Nigeria Mound, 1%1 Aleurorulus nephrolepidis ferns Calif. Essig, 1958 (Quaintance) A.filiciwn (Goldi) fern ? Mound, 1961 Trialeurodes wporarionun ornamental ferns France ; USA Schneider, 1966 (Westw.) (P) T.williamsi Trehan Anemia sp.;-- Diplnziwn - Britain (green- Trehan, 1937 proliferum; Dryopreris house) flaccida; Nephrodium ~onzns;dleandra africana Aphididae: Acyrrhosiphon (=Myzus) Adiantum concinnum; A. Calif. Severin et al., 1950 solani (Kaltenbach) (P) cuneatwn; Aspleniwn nidus Cyrtomium falcarwn Amphorophora ampullata Aspleniwn sp.; Cysto- India, Brit., Ghosh, 1974; Buckton preris montana; Onoclea Minn. Patch, 1938 struthiopteris A. a. bengalenris H.R.L. Cheilanthes sp. P. aquilin- Ghosh, 1974 and Basu wn; ferns A. ( = Megoura) Dryopteris dihtata Gray Japan Patch, 1938 dryopteridis Matsumura A. karoi Takahashi Polypodium ellipticum Formosa Patch, 1938 A. Iamgi Mason Aspidiwn; Aspleniwn; Brit., N.E. Patch, 1938; Onoclea sensibilk; 0. N. Amer. Robinson, 1966 struthiopteris; Polystich- um ;Pteretis nodulosa ; Marfeucia struthiopteris Anthracosiphoniella mama Asplenium esculenrum; India Ghosh, 1974 larum (Basu) Athyrium sp.; ferns Aphis fabae Scopoli (P) P. aquilinum Brit. Is. Lawton, 1976 USA Patch, 1938

AuIacorthum dryopteridis Dryopteris austriaca (Icq.) Czechoslovakia Holman, 1958 Holman Wo. ; D./ilix-mas (L.); A thyrium filixsfemina A. filicis Van der Goot fern Java Patch, 1938 Biochemical and Evolutionary Aspects of Arthropod Predation on Ferns

Appendix 1 (continued)

Arthropod taxon Fern host . Locality .Reference

A. pterinigrum Richards Preris sp. Brit. Columbia Richards, 1972 (0) A. sp. Polypodiwn sp. Hawaii Timberlake, 1924 Cerataphis lataniae Cyrtomium Calif. Patch, i938 (Boisduval) Idioptencr nephrelepidk Acrostichum reticulatum; Cosmopolitan Patch, 1938; Davis Adiantum capillus veneris; Severin et al., 1950; A, concinnwn; A. cuneatwn; Bonnemaison, 1937 ; A. spp.; Aspleniwn platy- Robinson, 1966; neuron; A. ruta; A. tricho- Steffan, 1962; manes L.; Ceropteris Zimmerman, 1948 calomelanos; Cererach officcinarwn Lam. and Dc.; Cibotium schredi; Cyrro- miwn falcatum; Dryopteris sp. ; Elaphoglosswn reticula- tum; Nephrolepis exaltata; N. daualloides; Onychiwn japonicum; Polypodiwn spp; Pterk cretica; P. tremula; Pteridiwn aquilinum; etc. Macromyzus polypodicola Dryopteris arida; Poly- Japan, Formosa, Ghosh, 1974 (Takahashi) stichum sp. ;ferns Sumatra, India

M. woodwardine (Takah.) Woodwardiasp.; Dryoprer- India, Formosa Ghosh, 1974 ' is monticola; D. wria; Rumora mutica; Asplenium curinrlarium; A. esculentum Macrosiphum adianti Adian tum pedotum ;Aspid- Ill., Minn. Patch, 1938 (Oestland) ium sp.

M. cysropreris Robinson Cystopteris bulbifera (L.) Penn. ~hbinson,1966 , . M. ptericolens Patch .Arhyrium filix-femina; N.E.N. Amer- Patch, 1938; Robinson,

P. aquilinum Colo., Brit , 1966; Lawton,.l,976 M. pteridis Wilson P. aquilinum; Adiantum W. USA Lawton, 1976; Severin, concinhum ;A. matum; et a]., 1950; Robinsbn, A. spp. 1966; Patch, 1938 Mastopoda pteridis P. aquilinum Mmn., E. Ca- Patch, 1938; Robinson, Oestland nada-Maine-N.J. 1966; Ghosh, 1974 Micromyzodium hi ferns India Ghosh, 1974 Verma M. filicum David Adiantum tinctum; Asplen- India Ghosh, 1974 ium trichomonas; Lartrea sp. ; Nephrolepis sp. ; * Pityrogramma peruvicma; Polypodium sp. ; Pteris cretica; Streptocarpus sp. Micromyzus judenkoi Cheilanthes sp. India, Sri Lan- Ghosh, 1974 Carver ka, Australia M. nigrum Van der Goot Adiantum ; ferns India, Sri Lan- Ghosh, 1974; ka, Java Patch, 1938 M.J. Balick et al.

Appendix I (continued)

Arthropod taxon Fern host Locality Reference

HEMIPTERA: HOMOPTERA (continued) Aphididae (continued):

Myzus (= Neomyzus) Adiantum concinnum; A. Ill., Calif., Severin et al., 1950; , circumfkxus Buckton (P) cuneatum ;A. hybridum; France Patch, 1938 ; Schneider, Cyrlomium falcalum; or- 1966 namental ferns M.filicis Basu ferns India Ghosh, 1974 M. omatus Liang (P) ferns Cosmopolitan Ghosh, 1974 M. persicae Sulzer (P) Marsilea quadrifolia; Colorado Patch, 1938 M.vestifa M. polypodicola Dryopleris arida; D. para- Sumatra, For- Patch, 1938 Takahashi silica; Polysrichum mosa M.pterisoides Theobald ferns Britai3 Patch, 1938 M. woodwardiae Polystfchum; Woodwardia Formosa, Japan Patch, 1938; Moritsu, Takahashi radicans 1952 Papuhphb sleesmani N.Y.,Pem. Robinson, 1966 (Pepper) Penralonia nigronemsa ferns Coquerel Phylhphb fagi L. A thyrium Jilix-femina Switzerland Patch, 1938 Rhopalosiphum s faphyleae Cystopteris monrana USA Patch, 1938 Koch R. nymphaeae L. Azolhfiliculoides; Marsilea' Brit, Formosa Patch, 1938 quadrifolia; Salvinia natans Shinjia preridifoliae Polypodium sp. ;Pferidium - India, Japan, Ghosh, 1974'; (Shinji) aquilinum ' S.E. Asia, Aus- Sorin, 1962 (0) tralia Unidentified sp. P. aquilinum New Guinea Kirk, 1977 Areopodidae (= Delphacidae) : Criomorphus pferidis P. aquilinum Brit. Is. (Spinola) Nesodyne (=Ilburnia) Sadleria cya fkoides; Hawaii Swezey, 1922 ipomoeicola (Kirklady) (P) Cibotriun sp. N. amamau Muir Sadleria cyatheoides Hawaii Swezey, 1922 N. nephrolepidis (Kirk.) Nephrolepis exaltata Hawaii Swezey, 1922 Nofhorestias badia Muir ferns Hawaii Swezey, 1922 N. swezeyi Muir Aspidium fern Hawaii Zimmerman, 1948 Nesoresfiasfilicicola Ehphoglossum gorgoneum; Hawaii Swezey, 1922; Kirk. Ciborium; ferns Zimmerman, 1948 N. nimbota (Kirk.) Phegopreris Hawaii Swezey, 1922; Zimmerman, 1948 . Cercopidae: . Philaenus spumarius (L.) P. aquilinum Brit. Is., Eur. Lawton. 1976; (PI Wienorek, 1973 Cicadellidae : Balclurha (= ~icaduG Ciborium camissai Hawaii Zimmerman, 1948 kihveae (Kirk.) Biochemical and Evolutionary Aspects of Arthropod Predation on Ferns

Appendix 1 (continued)

Arthropod taxon Fern host Locality Reference

Calladonus commism P. aquilinum Calif. DeLong et al., 1948 (Van Duzee) Friscanus intricafw(Bell) P. aquilinum Calif. DeLong et al., 1948 Cicadella (=Euptetyx) ferns Ill. DeLong, 1948 flavoscuta Cica&lJa nigra (Osborn) Polystichum adiantiforme Fla. Meade et al., 1%8 C. omani (Christian) Polystichum adinntiforme Fla. Meade et al., 1968

Nesophrosyne mysines Sadleria Hawaii ' Zimmerman, 1948 Kirk. (0) N. obliqua Osborn (0) Sadleria Hawaii Zimmerman, 1948 N. umbrotilis Kirk. Microlepia strigosa Hawaii Zimmerman, 1948 Nesophryne kaiamamao Microlepia sfrigosa Hawaii Zimmennan, 1948; Kuk. (0) (=N.filicola Swezey, 1922 Kirk.; N. microlepiae Kirk.) Cuiidae: Iolania perkinsi Kuk. ferns Hawaii Zimmerman, 1948; Swezey, 1922 I. spp. ferns Hawaii Zimmerman, 1948 OIiarus knonohi Kirk. tree ferns Hawaii Swezey, 1922 (4(0) 0. filicicola Kirk. (=O. tree ferns; Cibotium Hawaii Swczey, 1922 montivagus Kirk.) (d) Zimmerman, 1948 0. haleakaloe Kirk. Cibotium chamissai Hawaii Swezey, 1922; Zimmerman, 1948 0.hakhaku Gifford Cibotium; Sadleria Hawaii Zimmerman, 1948 (4 (0) 0. immaculatus Giff. ferns Hawaii Zimmerman, 1948 0. Koanoa Kirk. (0) tree fern Hawaii Zimmerman, 1948 0. Koele Gift ferns Hawaii Zimmerman, 1948

0. opum Kirk. , Nephrolepis exaltata Hawaii Zimmerman, 1948 0.pele ,Kirk. (d) (0) tree ferns Hawaii Zimmerman. 1948 Coccidae : Coccus (=Lecanium ferns (including Brit. Is., USA Newstead. 1903; hesperidum (L.) (P) ornamentals) Dodge et at., 1948 Ctenochiton depressus Cyafhea N. Zealand Maskell, 1887 Mask. Eucalymnatus tessellatus ferns Hawaii Zimmerman, 1948 (Sign.) Eulecanium (=Lecanium) Alsophila colemoi N. Zealand Maskell, 1887 mori (sign.) News feadia colka;ti Blechmcm spicnnf; Poly- Bklgium Ghesquiere, 1934 Gh-4. 0) (P) stichum juniperinurn Diaspididae : Aspidiotus hederae ferns Calif. Essig, 1915 (Vallot) (P) Chrysomphalusficus ferns Fla. Dekle, 1965 Ashmead M.J. Balick et al.

Appendix 1 (continued) - Arthropod taxon Fern host Locality Reference

HEMIPTERA: HOMOPTERA (continued) Diaspididae (continued): C. dictyospermi (Morgan) ferns Hawaii Zimmerman, 1948 (P)

Fiorinia fwriniae (Targ.) ferns Calif. , (PI Hemiberlesia rapax Akophila australis R. Br. Fla. Dekle, 1965 (Comstock) Parlatoriaproteus (Curtis) Asplenium nidus Fla. Dekle, 1965 Phenacaspis ( = Chionaspis) Asplenium N. Zealand Maskell, 1887 dubia (Mask.). Pinnaspis (= Chionaspis; Adianhun sp. ; Asplenium Cosmopolitan Dekle, 1965 ;Wieczorek, Hemichionaspis) nih; A. spp.; Cheilanthes 1973; Newstead, 1901 ; aspidistrae (Signoret) (P) sp. ; Cyrtomiumfalcatum McKensie. 1956 ; Presl. ; Davallia fejeensis Werner, 1931 Hook. ;Nephrolepis exaltata; Platycenun bifurcatum (Cav.) C. Christ; Polypodium poly- podioides; p. punctatum (L..); Polysrichum adiontiforme; P. Ionchitis (L.) Roth; Pteris spp. ; Tectaria heracleifolla (Willd) ; TheIypteris normalis (C. Christ) Pinnaspis strachani Adionturn sp.; ferns Fla., Hawaii Dekle, 1965; (Cooky) Zimmerman, 1948 Polinspis media Mask. ferns N. daland Maskell, 1887 Pseudoparlatoria parlo- ferns Calif. McKensie, 1956 torioides (Comstock) (P) P. pentagons (Targ.) (P) ferns Calif. McKensie, 1956 Eriococcidae: Eriococcus insignis New- Pteris sp. ;Pteridium Europe. W. Asia Hoy, 1963 stead (P) aquilinum Lecaniidae : CeroplatesjloridensB ferns (including orna- Calif. Essig, 1915; Dodge Comst. (P) mentals) et a]., 1948 . C. rubens Mask. Asplenium; Elaphoglossum Hawaii Zimmerman. 1948 reticulorum ;ferns Pulvinaria floccifera ferns Calif., Ore, B.C. Steinweden, 1946 Westw. P. mammeae Mask. (P) ferns Hawaii Zimmerman, '1 948 P. psidii Mask. (P) ferns Hawaii Zimmerman, 1948 Saissetia cofleae Wlk. (P) Adiantwn spp. ; Asplenium Cosmopolitan Newstead, 1903; Cory, (=Lecanium hemisphaeri- spp. ;Pteris spp. ; Nephro- 1945; Evans, 1942; mm Uarg.)) lepis; ferns Speyer et al., 1938; Essig, 1915; Zimmer- man, 1948 Biochemical and Evolutionary Aspects of Arthropod Predation on Ferns

Appendix 1 (continued)

Arthropod taxon Fern host Locality Reference

Pseudococcidae: .Geococcus radicum ferns Hawaii Zimmennan, 1948 Green (P) Pedronia spp. ferns Hawaii McKensie, 1967 Pseudococcus adoniifum ornamental ferns; ferns USA; Hawaii Dodge et al., 1948; (L.) (PI Zimmerman, 1948 P. citri (Rim) (P) ferns Calif. Essig, 1915 Pseudococcus glaucus Cyathea N. Zealand Maskell, 1887 (Mask.) (P)

' P. longispinus (Targioni- Nephrolepis exalfa&; N.sp. Calif. McKensie, 1967 Tazzetti) (P) P. obscurus Essig (P) Aspleniwn sp. Calif. McKensie, 1967 Rhizoecus pritchardi A&ntum Calif. McKensie, 1967 McKensie (P) (r) Ripersia filicola Trichomanes spicatum Brit. Is. ~ewstead,1903 Newstead I Ricaniidae : Scolypopa australis ferns N. Zealand Cumber, 1966 Walker (P)

HYMENOPTERA Blasticotomidae: ~lasficotomaPliceri Klug Athyrium filk-femina; Dry- Palearctic Benson, 1951 ; Schedl, (m) (g) opreris spp. ; Matteuccia 1973 srruthiopteris (L.) ; P. aquilinwn; Polystichum sp. Cynipidae : Unidentified sp. (g) P. aquilinurn Europe D. Van Leeuwen, 1938 Formicidae (assoc.- " nectary " feeders): ferns ; P. aquilinum Cosmopolitan W.L. Brown (p.c.); Lawton, 1976 Tenthredinidae : Aneugmenus (=Selandria) A fhyrium; Dryopteris; Engl., Eur., Lawton, 1976; coronatus (Klug) P. aquilinum Siberia Wieczorek, 1973 . A.fivipes (Norton) P. aquilinum USA Hogh, 1966 A. furstenbergensis P. aquilinum; ferns Brit. Is., Eur. Lawton. 1976; (Konow) Wieczorek, 1973 A. padi (L.) P. aquilinum ; ferns Brit. Is., Eur., Lawton, 1976; Siberia' Wieczorek, 1973 A. starnineipes Klug (g) P. aquilinum Eur., Brit Coi. D. Van Leeuwen, 1938; Ross, 1932 Aneugmenus temporalis P. aquilinum; ferns Brit. Is., Eur., Lawton, 1976; (Thornson) Siberia Wiecwrek, 1973 USA Hogh, 1966 M.J. ~alicket a].

Appendii 1 (continued)

Arthropod taxon Fern host Locality Reference

HYMENOPTERA (continued) Tenthrdinidae (continued): Emprin excisa Thornson P. aquilinum Brit. Is., Eur., Lawton, 1976; (P) Caucasus Wiecwrek, 1973 Hepfamelus ochroleucus Aihyrium spp. ; Blechnum; Brit., Eur. Lawton, 1976; Stph. (m) Polypodium vulgare ;P. sp. ; Wieczorek, 1973; Pferidium aquilinum Stansfield, 1933 Pseudohemi&xonus sharpi ferns Europe Wieczorek, 1973 (Caeron) Strombocerus delicalulus Athyrium; Dryopleris; Brit. Is., Eur., Lawton, 1976; (Fallen) Mar teuccia; Onoclea; Kamchatka Wieczorek, 1973 Polypodium ;Polystichum ; Pteridium aquilinum Slrongylogasfer contigua P. aquilinum Eur.-N. Africa- Lawton, 1976; (Konow) Kamchatka Wieczorek, 1973 S. dirtam Norton P. aquilinum Calif.-Brit. Col. Beer, 1955 S.Jlicb Klug P. ap4"Iinum; ferns Europe-Japan Wieczorek, 1973 S. lineata (Christ) Dryopferis; Matleucia ; Brit. Is., Em.- Lawton, 1976; P. aquilinum ; ferns Central Asia- Wieczorek, 1973 Japan S. manrlata (Klug) Afhyrium; P. aquilinum Brit. Is., Eur. Lawton, 1976; Wieczorek, 1973 S. mix& (Klug) Afhyrium; Dryopferb; 'Brit. Is., Eur.- Lawton, 1976; P. aquilinum ' Kamchatka Wieczorek, 1973 S. multicinctus Norton P. aquilinum USA Hogh, 1966 S. struthiopteridis Matfeuccia ,Europe Wieczorek, 1973 (Forsius) S. tibialis Cresson P. aquilinum Calif.-Brit. Col. Beer, 1955 S. xanfhoceros (Stephens) P. aquilinum Engl., Eur.- Lawton, 1976; China Wieczorek, 1973 S. spp. P. aquilinum USA Hogh, 1966 Tenthredo balieata Klug P. aquilinwn Europe Wieczorek, 1973 (PI T. colon Klug (P) P. aquilinum Brit Is., Eur.- Lawton, 1976; Asia-Kamchatka Wieczorek, 1973 Tenthredo ferruginea P. aquilinum Brit. Is., Em.- Lawton, 1976; Schrank (P) Asia-Kamchatka Wieczorek, 1973 T. Iioido L. (P) P. aquilinum Brit. Is., Eur.- Lawton, 1976; Siberia Wieczorek, 1973 T. spp. P. aquilinum USA Hogh, 1966

LEPIDOPTERA Arctiidae : Arctia caja (L.) (P) P. aquilimm N. Amer. Tietz, 1972 Dincrkia pteridis H. P. aquilinum N. Amer. Tietz, 1972 Edwards (P) Biochemical and Evolutionary Aspects of Arthropod Predation on Ferns

Appendix 1 (continued)

Arthropod taxon Fern host Locality Reference

D. oagans (Boisduval) (P) Pteris sp. N. Amer. Tietz, 1972 Rhyparioides metelkana Nephrodium fhelypteria Rumania Leestmans, 1975 Lederer Spilosoma hrieum P. aquilinum Brit. Is. Lawton, 1976 (Hufnagel) (P) Gelechiidae : Depressaria impurella P. aquilinum Europe Wieczorek, 1973 Tr. (P) Palfodora cytisello Curt. P. aquilinum Engl., Ire., Eur. Lawton, 1976; (g) Wieczorek, 1973 Geometridae: Cinquilia catenaria Dennsfaedtia punctilobula N. Amer. Tietz, 1972 (Dm& (PI (Michx.) .Gueneria basinria Wlk. Dryopieris yoaboracensis N. Amer. Xetz, 1972 (L.) Homochlodes (= Lozo- P. oqruoqruIinum E. Can.-Wisc.- Forbes, 1948; gramma)fritilloria (Guen.) N.C.; Ha. Kimball, 1965 H. lacfispargaria(Walker) P. aquilinum E. Canada Ferguson, 1975 Lithina (=Apaecaria; various ferns E. Can.-N.J.- Forbes, 1948 Lozogramma) wisc. SU~U~~UUM(Wlk.)

Petrophora (=Lithim; P. aquilinum; Dryopteris Brit. Is; Eur. . Lawton. 1976; Phasiane petraria Hub.) Wieczorek, 1973 chlorosara (hopoli) Philedia puncromacularia Pteris sp. N. Amer. Tietz, 1972 (Hulst) Scotorythra rara (Butler) ferns; Cibofium Hawaii Swezey, 1922; (P) Zimmerman, 1948 Heliodinidae : Erineda aenea Braun (s) Asplenium anguslifolium; Ohio Braun, 1918 A. acrosfichoides Erinedo elyella Busck (s) Acrostichum aureum S.W. Fla. Needham, 1947 Hepialidae : Hepiahu fusconebulosur P. aquilinum Brit. Is., Eur. Lawton, 1976 DeGeer (r) H. gracilis Grote (r) ferns Maine-Mass.- Forbes, 1923 Colo. H. hectus L. (P) (r) P. aquilinum Brit. Is.; Eur. Lawton, 1976 H. sylvinus (L.) (P) (r) P. aquilinum Engl.; Scotl. Lawton, 1976 Lycaenidae: . Hyoochrysops theon Drynuria quercifolia (L.) Australia Daniels, 1976 medocus (Fruhstorfer) J. Smith Noctuidae: Achea janata L. (P) Polypodium sp. Hawaii Anonymous, 1945 Actebia fennica Dennstaedfia punctilobula N. Amer. Tietz, 1972 (Tauscher) (P) (Michx.) M.J. Balick et al.

Appendix 1 (continued)

Arthropod taxon Fern host Locality Reference

LEPIDOPTERA (continued) Noctuidae (continued): Callopislriajloridemis Adiatum sp.; A.pednrum So. USA, Fla., Comstock, 1939; Guenee L. ; Blechnum sp.; Cyrtom- Calif. to tropics Forbes, 1954; lietz, ium sp. ;Nephrolepis sp. ; 1972 N. exaltata; Polypodium sp. ; Pteris sp. ; Wwdwardia oirginica (L.) Sm. C. granitosa Guenee ferns N.J.-Fla. Forbes, 1954 C.juvenrina (Cramer) P. aquiIinum Europe Wiecwrek, 1973 C. htreilli Dup. P. aquilinwn Eur., N. Africa Wieczorek, 1973 C. (= Eriopus) maillordi Adianrum sp. Thailand Cantelo. 1965 Guenee C. mollissima Guenee ferns Can.-F1a.-Colo. Forbes; 1954 C. monetifera Guenee ferns E. Can.-Wise.- Forbes, 1954 Fla. C. purpureofatciata Pill. P. aquilinum Europe Simmonds, 1967 Ceramica pisi (L.) (P) P. aquilinum Brit. Is. Lawton, 1976 Euplexia benesimilis P. aquilinum Calif., Eur. Essig, 1958 McDunn. (P) E. Iucipara (L.) (P) P. aguilinum, ferns Brit. Is., Ew. Lawton, 1976; Wieczorek, 1973 Fagitana liltera (Guenee) Dryopleris rhelypteris (L.); Fla., N.J. Kimball, 1965; Osmunda sp. D. Schweitzer (p.c.) Habrynlhis (= Phlogo- Aspidium filix-mas; , Eur. Wieczorek, 1973 phora) scita (Huebner) Blechmm sp. ; P. aquiIinum (PI Haliophyk (=Erio- many ferns Hawaii Swezey, 1922 pygodes) euclidias (Meyrick) H. spp. several ferns; Acrostichum; Hawaii Zimmerman, 1948 Aspidium cyatheoides Loconobirr (=Mameslra) P. aquilinum Brit. Is. Lawton, 1976 contiqua (Den. und Schlif.) (P) (assoc.) L. oleracea (L.) (P) P. aquilinum Brit. Is. Lawton. 1976 (assoc.) Orthodes crenuhta Onoclea sensibilis L. Penn. Schweitzer (p.c.) (Butler) (P) Papipema inquoesita Onoclea semibilis L. Can.-111.-Wash.. Forbes, 1954 Gr. and Robins. (r) D.C. P. n. sp. near pterisii (r) Pteretis pennsylvanica N.E. USA D. Schweitzer (p.c.) (Willd.) P. pterisii Bird (I) P. aquilinum E. Can.-N.Y.- Forbes, 1954 Pa. P. speciosissima Gr. and Osmunda regalis L.; 0. E. Can.-N.Y.- Forbes, 1954 Rob. cinnamomea Ga. Biochemical and Evolutionary Aspects of Arthropod Predation on Ferns

Appendix 1 (continued)

Arthropod taxon Fern host Locality Reference

P. stenocelis (Dyar) (r) Woodwardia virginica N.E. USA Forbes, 1954 (L.) Sm. Peridroma margaritosa P. aquilinum N. Amer. Tietz, 1972 (Haworth) (= P. saucia Huebner) (P) Phlogophora meticulosa P. aquilinum Brit. Is. Lawton, 1976 (L-) (PI Polia adjuncta (his- P. aquilinum N. Amer. Tiey 1972 duval) (P) . P. assimiIis(Morrison)(P) P. aquilinum N.Y. J. Franclemont (p.c.) Nymphalidae : Metamorph stelens (L.) Blechnum sp. Fla. ;Cuba Kimball, 1965; Dethier, 1940 Olethreutidae : Olethreutes osmunddna Osmunda regalis Maine; Wash., Forbes, 1923 Fernald (0) D.C. Pyralidae : Ambia? sp. Polystichum adiontiforme Fla. KimbaU, 1965 Nymphulo? novaginensis P. aquilinum New Guinea Kirk, 1977 Kenrick Phlyctaenh (=Pyrausta) various ferns E. Can.-Fla. Forbes, 1923 rhcseusalb . Psara platycapnu Meyrick P. aquilinum New Guinea Kiik, 1977 Pyrausta aeglealir Walker Osmunda N.Y. Forbes, 1923 P. descrepitalis H.-S. Aspidium spinulosum Engl. Ford, 1949 Samea multiplicalis Salvinia sp. ; S. rotund~olia Trinidad, No. S. Thomas, 1975; Knopf Guenee Azolla caroliniana; Pbtia Amer., Ha. et al., 1976 stratiotes Undubmbia polystichalis Polystichum adiontiforme Fla. Kimball, 1965 Capps Unident*ed sp. P. aquilinum New Guinea Kirk, 1977

Antherea eucalypti Nephrolepis Australia French, 1933 Scott (PI

Opogonu chlinata Meyr. Polypodium quercifolium India Maxwell-Lefroy, 1971 (am-) Praecedes thecophora P, aquilinum New Guinea Kirk, 1977 Wals. Teichobiafilicinora Meyr. Aspidium filix-mas Engl. Hering, 1951 ; Ford, 1949 T. verhuellaella Stt. Several Polypodiaceae ; Engl. . Ford, 1949

Unidentified sp. 1 (m) P. aquilimun New Guinea Kirk, 1977 Unidentified sp. 2 (m) P. aquilinum New Guinea Kirk, 1977 Tortricidae : Tortrix ciranu Fernald (P) ferns Calif. Essig, 1915 M.J. Balick et al.

Appendix 1 (continued)

Arthropod taxon Fern host Locality Reference

LEPIDOPTERA (continued) Yponomeutidae (=Walshiidae) : Bafrachedra sophroniella Aspidium cyafheoides Hawaii Swezey, 1922 Walsingham(s) B. Iomenfella Wals. fern Hawaii Swezey, 1922 B. bedeliella Wals. (s) Asplenium nidus;Elapho- Hawaii Swezey, 1922 glossurn reticukmm; Dryo- pteris parasilica B. sp. (m) Pteris irregulnris Hawaii Swezey, 1922 B. SPP. (4) (?m) Polypodium spectrum Hawaii Swezey, 1922 (?part) Euhyposmocoma ekaha Asplenium nidus Hawaii Swezey, 1922 (Swezey) E. trivitella Swezey (m) Elaphoglossum reticulafum Hawaii Swezey. 1922 Hyposmocoma spp. ferns Hawaii Swezey, 1922 (several)

ODONATA

Anfiagrion grinskrgsi BIechnum chilense , C hie Jurzitza (oviposits only) (assoc.) Megalagrwn oahuense Gleichenia linearis Hawaii Zimmerman, 1948 (Blackburn) (truly terres- trial, lives in fern litter) (assoc.)

ORTHOPTERA Acrididae: Paulinia acumimta Salvinia Trinidad, No. S. Thomas, 1975 (De Geer) Amer. Unidentified sp. P. aquilinum New Guinea Kirk, 1977 Gryllidae: Lepfogrylhu nigrolineatus Cibotium Hawaii Zimmerman, 1948 Perkins (0) Para frigonidiumJlicum "tall soft ferns"; Hawaii Swezey, 1922 Perkins P. viridescens Perkins "creeping fern"; Lindsaya Hawaii Swezey, 1922; macraeana ? Zimmerman, 1948 Progmrhogryl~usatatus ferns ; Cibotiwn Hawaii Swezey, 1922 ; Brunner (P) Zimmerman, 1948 Tetigoniidae : Banza spp. (P) ferns Hawaii Swezey, 1922 Biochemical and Evolutionary Aspects of Arthropod Predation on Ferns

Appendix 1 (continued)

Arthropod taxon Fern host Locality Reference

THYSANOPTERA Phlaeothripidae: Atractothrips macrurus ferns Japan (Ryukyu Okajima, 1975 Okajima (d) (P) Is.) Haplothrips hawaiiensis Cibotium menziesii; tree Hawaii Zimmerman, 1948 Moulton (d) (0) ferns H. rosai Bianchi (P) Sadleria Hawaii Zimmerman, 1948 Kamyolrips doliicornis Sadleria Hawaii Zimmerman, 1948 Bianchi (P) K melaleuca (Bagnell) (P) Ciborium chamissoi Hawaii Zimmerman, 1948 Uredothrips tibialis ferns Japan (Ryukyu Okajima, 1975 Okajima (d) Is.)

Heliothrips haemor- ferns (including om- Calif. . Essig, 1915; Dodge rhoidalis Bouche (P) mentak); Pteris cretica et al., 1948 Leucothrips (=Micro- Asplenium sp. Mich. Herrick, 1927 rhrips) Ieucus Herrick L. nigripennis (Reuter) ferns (including orna- USA Lewis, 1973 mentals) ;Pteris cretica Physorhrips (=Pterido- Polypodium pteropus Moore Java Priesner, 1938 thrips) pferidicola (Karny) (g) Zonothrips osmunahe Osmunda cinnnmomaea N.J. Crawford, 1941 ; Crawford Asplenium nidus; Pleopeltis Indonesia D. Van Leeuwen, 1938 UnidenWied spp. (g) superfacialis Bedd.

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