biochemical and evolutionary aspects of arthropod ... · climates. although all ferns tested...

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 cyanogenic 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 arthropod 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 suggesting 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 compounds 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 cyanogenic 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 arthropod 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


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

C

a d a, b a a a a, b, c, e a, b a, d, e a a b . a, b a, b a, b, e a, e a, d a + fungal damage e a, d a, b a a, b b 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 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

I to fern species

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 correlation 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


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 arthropod 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 community 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 feeding 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 apparently 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), however, there are very few recorded on today's ferns. Other arthropod orders,


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.


Appendix 1

Arthropod-Fern Herbivores

Arthropod taxon Fern host Locality Reference

ACARINA Chamobatidae: Chamobates sp. (= ?associate)

Britain Lawton, 1976

Eriophyidae: Eriophes (= Phyrop) Nephrolepis spp. paurop Nalepa (g) E. pterides Moll. (g) ferns; P. aquilinum

E. SP. (g) Dicranopteris sp.

Erwphyhrum sp. ,(O) Eriophyid sp. (g)

P. aquilinwn Angiopteris m c t a (Fors:) Hoffm.

Phyllocoptes dimorphus P. aquilinum Kiefer

Old World D. van Leeuwen, 1938 Tropics Brit. Isl. USA

D. Van Leeuwen, 1938 Jeppson et al., 1975

Portugal Tavares, 1930 Old World D. Van Leeuwen, 1938 Trop. Calif. Kiefer, 1940

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;

Polypodiwn pectinatum x plwmclwn

Puerto Rico DeLeon, 1%6

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; Polypodium sp.

COLLEMBOLA (=?assoc.) Sminthuridae: Bourletielln viridescens P. aquilinwn Stach, S. Gisin (F')

Dicyrtomidae : Dicyrtoma sp. P. aquilinwn

Ha., Mex. Trinidad

Brit. Is., Eur. Lawton, 1976

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 Banks (P)

Hawaii Zimmerman, 1948

M.J. Balick et al.

Appendix 1 (continued) - -

Arthropod taxon Fern host Locality Reference -

CORRODENTIA (continued)

Psocidae : Psocus distinguendus Sadleria Perkins ( P )


COLEOPTERA Alticidae ( = Chrysomelidae) : Apthom sp. P. aquilinum Manobia sp. 1 P. aquilinum M. sp. 2 P. aquilinum

Anthribidae: Homocloeus n. sp. 1 tree ferns Homocloeus n. sp. 2 tree ferns

Ormiscus floridanus (Leng) ferns Phaenorheriopsb n. sp. tree ferns Trigonorhinus ronmdatw ferns Leconte

Carabidae: Several native Hawaiian tree ferns SPP. ( 4 (assoc.)

Cerambycidae : ?Cornallb sp. P. aquilinum Sybra sp. P. aquilinum Tmesisfernus sp. 1 P. aquilinwn T. sp. 2 P. aquilinum Unidentified sp. 1-sp. 5 P. aquilinum

Cryptophagidae: Unidentified sp. (s) Cysfopteris fragilis

Curculionidae: Bagous sp.

Baris arropolifa Lea Brachyrhinus ( = Otior- hynchus) sulcarus (F.) (P) (r and fronds) Cryptorhynchinae ?sp. Cyriobagous singuloris Hulst. Dryophthorus insignis Sharp ( 4 D. modestus Sharp (d) D.pusillus Sharp Gromihrs gracilipes Sharp Heteramphus filicwn Perkins

saluinia sp.

P. aquilinwn ornamental ferns

P. aquilinum Salvinia

Cibotium spp.

Cibotium spp. Cibotium (2 or 3)

? Blechnum tree ferns

New Guinea New Guinea New Guinea

Jamaica Jamaica S. Fla. Jamaica Mass.

Hawaii

Kirk, 1977 Kirk, 1977 Kirk, 1977

Valentine (in litt.) Valentine (in litt.) . Valentine (in litt.) Valentine (in litt.) Valentine (in litt.)

Swezey, 1922

New Guinea Kirk, 1977 New Guinea Kirk, 1977 New Guinea. Kirk, 1977 New Guinea Kirk, 1977 New Guinea Kirk,' 1977

Chile (Juan J. Lawrence (p.c.) Fernandez Is.)

India Vorghese et al., 1972

New Guinea Kirk, 1977 USA Dodge et al.,

1948

New Guinea Kirk, 1977 Trinidad, No. Thomas, 1975 S. Amer. Hawaii Swezey, 1922

Hawaii Swezey, 1922 Hawaii Swezey, 1922 N. Zealand May, 1973 Hawaii Swezey, 1922


Appendix 1 (continued)


H. swezeyi Perkins (m) EIaphoglossum reticulatwn E. micradeniwn; E. gorgo- newn ; E. squamoswn

Hawaii Swezey, 1922

Heteramphus wollasfoni tree ferns Sharp (0) Homolocerus lyciformis Polystichwn ( = Ruhmora) (German) denriculatwn Pr. ; several

Cyatheaceae Homolocerus nigripennis tree ferns Bohemian H. plawnanni Voss Polystichwn denticulatum

Megacolabus bifurcatus Blechnwn capense (L.) May 6 ) Miocalles (=Micro- cryptorhynchus) abditiceps (Zimmerman)' (0) M. abnormis (Zimm.)

M. ambiguus (Zimm.)

ferns

Pterb sp.

fern

M. andersoni (Zimm.) (P) ferns M. angustatus (Zimm.) M. bicolor (Zimm.) M. brevis (Zimm.) (P) M. caudahcr (Zimm.)

Cyathea sp. Cyathea sp. ferns Blechnwn sp.

M. chaetectetoroides ?ferns (Zimm.1 (PI M. confmus (Zimm.) fern M. conwxus (Zimm.) (0 ) fern M. cookei (Zimm.) Pterir sp.

M. c u m (Zimm.) fern M. dircretw (Zimm.) Blechnum, ?ferns M. exilk (Zimm.) fern M. fosbergi (Zimm.) Cyarhea sp. M. fowawntris (Zimm.) Aspleniwn nidus

M. fulgidus (Zimm.) fern

M. gracilis (Zimm.) fern

M. hirtw (Zimm.) fern

M. impressicollk (Zirnm.) Asplenium nidus

M. irregularis (Zimm.) Cyathea sp. M. irroratus (Zimm.) fern M. mangaoae (Zimm.) Cyathea sp. M. mangarevae (Zimm.) Aspleniwn nidus

Hawaii

Brazil

Swezey, 1922

Brazil Vanin, 1976 N. Zealand May, 1973

Austral Islands Zimmerman, 1957 ( T a p 1s.)

Austral Islands (Rapa Is.) Society Islands (Tahiti Is.) Rapa Is. Tahiti Is. Rapa Is. Tahiti Is. Society Islands (HuaIhe) Rapa Is.

Tahiti Is. Tahiti Is. Austral Islands (Raivavae Is.) Rapa Is. Tahiti Is. Tahiti Is. Tahiti Is. Rapa Is. Tahiti Is.

Austral Islands (Tubuai Is.) Austral Islands (Tubai Is.) Rapa Is. Rapa Is. Tahiti Is. Rapa Is. Mangareva Is.

Zimmerman, 1957

Zimmerman, 1957

Zimmerman, 1957 Zimmerman, 1957 Zimmerman, 1957 Zimmerman, 1957 Zimmerman, 1957

Zimmerman, 1957

Zimmerman, 1957 Zimmerman, 1957 Zimmerman, 1957

Zimmerman, 1957 Zimmerman, 1957 Zimmerman, 1957 Zimmerman, 1957 Zimmerman, 1957 Zimmerman, 1957

Zimmerman, 1957

Zimmerman, 1957

Zimmerman, 1957 Zimmerman, 1957 Zimmerman, 1957 Zimmerman, 1957 Zimmerman, 1957

M.J. Balick et al.



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. M. punctipennis ferns (Zimm.) (P ) M. pusillus (Zimm.) (P) Aspleniwn nidus

Rapa Is. Rapa Is.

Zimmeman, 1957 Zimmerman, 1957

Rapa Is. Zimmerman, 1957 M. raivavaensir Preris sp. (Zimm.) p) M. rurutuensis (Zimm.) Dryopteris sp. M. setigem (Zimm.) (P) Cyathea

Austral Islands Zimrnerman, 1957 (Raivavae Is.) Rurutu Is. Zimmerman, 1957 Rapa Is. Zimmerman, 1957

M. setulosus (Zimm.) Aspleniwn nidus; Cyathea spp.

M. similis (Zimm.) (P) ferns M. sparhifer (Zimm.) Asplpnium niduc M. tenuis (Zimm.) (P) Cyarhea sp. M. tesraceu (Zimm.) fern

Rapa Is. Zimmerman, 1957

Tahiti Is. Rapa Is. Rapa Is.. Tubuai Is.

Zimmerman, 1957 Zimmerman, 1957 Z i e r m a n , 1957 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. Lea

Australia . Anonymous, 1940

Notacalles piciuentrir Polystichum vestitum (Brouu)

Canada (Camp Kuschel, 1964 bell Is., B.C.)

Oodemas aenescens Bohman (d) (P)

Ciborium spp. Hawaii Swezey, 1922

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 (introduced)

Strophosmos sp. Europe




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) Chile Trichophthalmus milt- Blechnum chilense;

omerus Blanchard PoIysfichum sp.

Elateridae : Corymbites nigricornis ferns Pz-1 Dolopius marginatus P. aquilinwn (L.) (assoc.)

Eumolpidae (=Chrysomelidae) : Unidentified sp. P. aquilinum

Galemcidae (= Chrysomelidae): Mimastra Iimbata Baly Adiantum sp.

Helodidae: Cyphon padi (L.) (assoc.) P. aquilinwn C. variabilis Thunberg P. aquilinum (assoc.)

Lathrididae :

Cartodere ruficollis P. aquilinum Marsham

Melasidae (= Eucnemidae) : Dirhagus pygmaeus (F.) P. aquilinum (assoc.)

Mycetophagidae: Unidentified sp. Cystopteris fragilis

Nitidulidae : Nesapferus monticola Cibotiwn menziesii (Sharp) (dl Proterhinidae:

Proterhinus longulur Cibotiwn chamissoi; C. Sharp (dl menziesii P. pteridis Perkins (m) Pteris sp. P. sharpi Perkins fern (non-arboreal) P. spp. ?ferns

Ptinidae : Holcobius hawaiiensis tree ferns Perkins (assoc.)

Scarabaeidae: Phyllopertha horticola P. aquilinwn (L.) P )

Brit. Is.

Engl.

Linssen, 1959

Lawton, 1976

New Guinea Kirk, 1977

Thailand Cantelo, 1965

Engl. Engl.

Engl.

Engl.

Lawton, 1976 Lawton, 1976

Lawton, 1976

Lawton, 1976

Chile (Juan J. Lawrence (p.c.) Fernandez Is.)

Hawaii Swezey, 1922

Hawaii

Hawaii Hawaii Hawaii

Hawaii

Swezey, 1922

Swezey, 1922 Swezey, 1922 Swezey, 1922

Swezey, 1922

Brit. Is.; Eur. Lawton, 1976 Wieczorek, 1973

70


M.J. Balick et al.


COLEOPTERA (continued)

Scolitidae:

Hypothenemus eruditus ferns Westw.

H. pusillus Egg. ferns

PoeciiQs preridophyrae P. aquilinum Schedl (m) Stephanoderes perhispidus ferns Egg.

Zaire Schedl, 1962

Zaire Schedl. 1962

New Guinea Schedl, 1962

Zaire Schedl, 1962

DIPTERA

Agromyzidae:

Hawaii Swezey, 1922 Agromyza undescribed Marattia doughsi SP. (4 Phytoliriomyza hilarefla Polypodium; P. aquilinum; (zett.1 ( 4 Osmunda (= Pteridomyza; Phytobia; Praspedomyza; Agromyza)

Brit. Is., Eur., Lawton, 1976 N. Amer. Wieczorek, 1973;

Hering, 1951

Brit. Is. ?Europe

P. preridii Spencer (m) P. aquilinum Lawton, 1976 Hering. 1951 Phytomyza scolopenrfii several Poiypodiaceae

R.-D. (m) Asplenium; Athyrium ; Blechmim

Hering, 1951

Anthomyiidae: Chirosia (= Pycnoglossa) Dryopreris aberrans Collin C. aibifons Tiens. P. aquilinum

, Engl.

Engl., Eur.

Wieczorek, 1973

Lawton, 1976; Wieczorek, 1973 Lawton, 1976; Wieczorek, 1973

Lawton, 1976; Wieczorek, 1973

Simmonds, 1964; Wieaorek, 1973 Lawton, 1976


C. albitarsis Zett. (m) P. aquilinum Brit., Eur., Asia

C. betuieti Ringdahl (g) Athyrium; P. aquilinum Europe

C. crassiseta Stein (m, r) P. aquilinum Eng., Eur.

C.flauipennis Fallen (m) P. aquilinum Chirosia hystricina Ron- P. aquilinum; Asplenium; dani (m) (= C. cinerosa A thyriwn; Blechnwn; Cyst- Stein; C. hystrix Brischke; opteris; Dryopteris; Matte- C. setifemur Ringdahl) uccia ; Osmunda; Polyp-

odium; Polystichum

C. parvicornis Zett. (m) Asplenium; Dryopteris; P. aquilinum

Hylemya (= Choro rophilia) Polypodiaceae; ferns signata Brisc. (g)

Britain

Britain, Eur., N. Amer.

Brit. Is., Eur. Lawton, !976; Wieczorek, 1973

Europe D. Van Leeuwen, 1938; Simmonds, 1967

Cecidomyiidae: Cecidomyia sp. (r-g) Cystopteris fragilis Felt, 1940


USA

Eng., Eur. Dasyneura firicina P. aquilinum (Kieff.) (g)




D. pferidicola Kieff. (g) P. aquilinwn; ferns

''a few spp." (g) ferns

Drosophilidae : Drosophila apicipwcta ferns; Sadleria Hardy

D. nigella Hardy (assoc.) ferns D. notha Bock (g) P. aquilinum

D. sadleria Bryan (m) Sadleria cyafheoides

Pipunculidae (assoc.-parasite): Pipwculus alienus (Hardy) ?leafhoppers over

Cibotiwn and Nephrolepis P. amplus a aid^ over Nephrolepis sp. P. cornutus (Hardy) over Nephrolepis sp. P.filicicolus Hardy over Nephrolepis exaltata P. jumtor Perkins ?leafhoppers over Nephro-

lepis and ferns P. megameris Hardy over Cibofium

P. obscuratus Hardy over Cibotium chamissai K. (assoc. with Neso- phrosyne sp. and Neso- sydne ipomoeicola Kirk)

P. timerlakei Hardy over Cibotium chamissai (assoc. with Nesophrosyne and Nesosdyne ipomoeicola)

P. swezeyi Perkins

Sciaridae : Sciara sp. (s)

leafhoppers on Nephrolepis

"young ferns"

HEMIFTERA: HETEROFTERA

Anthocoridae (assoc.): Lasiochilus denigratus Cibotiwn chamissai (White) ( 4 (0) Unidentified sp. Cyathea

Lygaeidae: Drymus ? brunneus P. aquilinwn Sahlberg Germahu costalis Van Cyafhea; ferns D- (0) G. fuliginosus Van Duzee (P )

Cyathea; ferns

G. infans Van Duzee (P) ferns Metrarga nuda White Cibofiwn ( 4 (0) M. obscura'Blackburn tree fern

Eng., Eur. Lawton, 1976; Wieczorek, 1973

Europe D. Van Leeuwen, 1938

Hawaii Hardy, 1965

Hawaii Hardy, 1965 New Guinea Kirk, 1977

Hawaii Swezey, 1922; Hardy, 1965

Hawaii

Hawaii Hawaii Hawaii Hawaii

Hawai Hawaii

Hawaii

Hawaii

Hardy, 1964

Hardy, 1964 Hardy, 1964 Hardy, 1964 Hardy, 1964

Hardy, 1964 Hardy, 1964

Hardy, 1964

Hardy, 1964

Netherlands Augustijn et a]., 1935

Hawaii

Ethiopia

Engl.

Marquesas Is.

Marquesas Is.

Marquesas Is. Marquesas Is.

Hawaii

Zimmerman, 1948

Limavuori (in litt.)

Lawton, 1976

Van Duzee, 1935

Van Duzee, 1935

Van Duzee, 1935 Zimmerman, 1948

Usinger et a]., 1959

M.J. Balick et al.



HEMIPTERA: HETEROPTERA (continued)

Lygaeidae (continued):

Chile Kormilev, 1952 Micrymenus kuscheli Dryopterir inaequalijolia; Kormilev (0) Polystichwn vesticum

Neseir hiloensb inter- medius Usinger (0) Nysiw blackburni White (0) N. communis Usinger (P)

misc. ferns Hawaii Zimmerman, 1948

ferns Hawaii Zimmerman, 1948

Cibotiwn menziesii; Sadleria

Pterb sp. fern


Austria Strobl, 1900 Eyles et al., 1969

N. jacobeae Strobl N. liliputanus Eyles and Ashlock (0) '

N. rubescens White

N. Zealand

Hawaii Hawaii

Zimmerman, 1948 Zimmerman, 1948

ferns Sadleria Oceanides montivagus

(Kirk) Ozophora pauescens (Distant) (P) 0. trmotata Barber Sephom criniger (white) (PI Styngnocoris sabulosus Lumsden

Fla. ; W. Indies Torre-Bueno, 1946 ferns

Torre-Bueno, 1946 Zimmeman, 1948

ferns ferns; Sadleria

Ha. Hawaii

Pteris sp. Lunsden, 1939

M i d a e : Bryocorinae many unde scribed spp.

ferns; "soft ferns" Africa

Brit. Is., Eur.

D. Leston (p.c.)

Bryocorir preridis (Fallen) 6 )

Aspidiwn /ilk-mas; A fhyriwn /ilk-femina; Dryopteris; Lastraea dilotata; L. ?oreopteris; Polysfichwn; Asplenium ; Pteridium aquilinwn

Athyriwn filix-femina

Lawton, 1976; Wieczorek, 1973; Butler, 1923

Calocorb sexguttatus Fab. (assoc.)

Brit. Is.

111.

Brit. Is.

Butler, 1923

Knight, 1941

Butler, 1923

Ceratocapsus setosus Reuter

ferns

Deraeocorir ( = Camp tobrochis) Iutescens (Schill.) (assoc.) Dicyphus globulijer . (Fallen) (assoc.)

P. aquilinwn

A fhyriwn filix-femina; P. aquilinwn

ferns

Brit.. Is.

Australia

Brit. Is.

Butler, 1923

Woodward et al., 1970

Butler, 1923

Felirarus glabratus (Motsch.) b p u r (= Onychwnenus) decolor (Fallen) (P)

ferns




Lygocoris paburinus (L.) Aspidium (assoc.)

Lygus cervinus Herr.- ferns Sch. (0 ) (assoc.) L. indistinctus Taylor P. aquilinum

Mocrolophus nubilus P. oquilinm Herr.-Sch. (assoc.) Mecomma ombulans Aspidiwn filix-femina; Fallen (P) Polypodium dryopteris

Miris sp. ferns

Brit. Is. Butler, 1923


Brit. Is. Brit. Is.

Butler, 1923 Butler, 1923


Africa N. America

R. Limavuori (p.c.) Wagner et al., 1952 Monaiocoris omericana Osmwrdn cinnomeo ferns

(Wagner and Slater) Monalocoris filicis (L.) (s) Aspidium filix-mar; As-

plenium ; Athyriwn Nix- femitia; Lastraeo dilaro- la; L. oreopteris; Poly- stichm; Aspidim spinulosum

Brit. Is., Eur. Butler, 1923; Lawton, 1976; Wieczorek, 1973

Nigeria

Hawaii

Hawaii

Brit. Is.

France

Limavuori, 1975

Zmmerman, 1948

Zimmerman, 1948

Butler, 1923

Schneider, 1966

M. (= Sthemrusoides) P. oquilinrun punctipemis Linnavuori Orthotyhu kassandra Sadleria (Kirk.) (PI Pseudoclerah moroi creeping fern Kirk. (assoc.) Stenodema hokatm P. aquilinum (Fab.1 (PI Tenthecoris bicolor ornamental ferns k o t t (PI Nabidae : (assoc.) Nabb blockburni (White (PI N. curtipemis Blackburn N. Iusciosw White (P)

N. pele (Kirk.) (0) N. siloestris (Kirk.) (0)

Pentatomidae: (assoc.) Aelia acuminalo L. (P)

Jalla dwnosa L. Zicrono caeruleo L.

ferns Hawaii Zimmerman, 1948

Hawaii Hawaii Hawaii Hawaii

Z i e r m a n , 1948 Zimmerman, 1948 Zimmennan, 1948 Zimmerman, 1948

tree ferns ferns Cibotium

Cibotium

ferns, Osmtmdo regolis

P. aquilimun

ferns

Brit. Is. Brit. Is. Brit. Is.

Butler, 1923 Butler, 1923 Butler, 1923

Reduviidae (assoc.) :

Empicorb rubromoculatus ferns (Blackburn) (P) Nesidiolestes . fern

Tingidae: Corythucho podi Drake P. aquilinum


Hawaii G a p e et al., 1975

Brit. Columbia Leech, 1944

M.J. Balick et al.


Arthropod taxon Fern host Locality Reference - -

HEMIPTERA: HOMOPTERA

Aleyrodidae : Aleuroplatus kewensis Anemia sp.; Diplnziwn Trehan prolfenun ; Dryopreris

jlaccida; Nephrodiwn con- jluens; Oleandra africana

Britain (green- Trehan, 1937 house)

Aleuropreridb douglasi Cyclosoncs dentatus; Mound Pteris togoensis

West Africa Mound, 1961

A. easropi Mound A. hargreawsi Mound

?fern Bush Fern

Ghana Mound, 1961 Sierra Leone Mound, 1961 Nigeria Mound, 1%1 Calif. Essig, 1958

A. jamesi Mound Pterb togoensis Aleurorulus nephrolepidis ferns (Quaintance) A.filiciwn (Goldi) fern ? Mound, 1961

France ; USA Schneider, 1966 Trialeurodes wporarionun ornamental ferns (Westw.) (P) T. williamsi Trehan Anemia sp.; Diplnziwn Britain (green- Trehan, 1937

house) - - -

proliferum; Dryopreris flaccida; Nephrodium ~ o n z n s ; dleandra africana

Aphididae: Acyrrhosiphon (= Myzus) Adiantum concinnum; A. solani (Kaltenbach) (P) cuneatwn; Aspleniwn nidus

Cyrtomium falcarwn

Calif. Severin et al., 1950

Amphorophora ampullata Aspleniwn sp.; Cysto- Buckton preris montana; Onoclea

India, Brit., Ghosh, 1974; Minn. Patch, 1938

struthiopteris

A. a. bengalenris H.R.L. Cheilanthes sp. P. aquilin- and Basu wn; ferns

Ghosh, 1974

A. ( = Megoura) Dryopteris dihtata Gray dryopteridis Matsumura

Japan Patch, 1938

A. karoi Takahashi Polypodium ellipticum Formosa Brit., N.E. N. Amer.

Patch, 1938 Patch, 1938; Robinson, 1966

A. Iamgi Mason Aspidiwn; Aspleniwn; Onoclea sensibilk; 0. struthiopteris; Polystich- um ; Pteretis nodulosa ; Marfeucia struthiopteris

Anthracosiphoniella mama Asplenium esculenrum; larum (Basu) Athyrium sp.; ferns

India Ghosh, 1974

Aphis fabae Scopoli (P) P. aquilinum Brit. Is.

USA

Lawton, 1976

Patch, 1938

AuIacorthum dryopteridis Dryopteris austriaca (Icq.) Holman Wo. ; D./ilix-mas (L.);

Czechoslovakia Holman, 1958

A thyrium filixsfemina

A. filicis Van der Goot fern Java Patch, 1938



Arthropod taxon Fern host . Locality .Reference

A. pterinigrum Richards Preris sp. (0) A. sp. Polypodiwn sp.

Brit. Columbia Richards, 1972

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 reticulatum; 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- (Oestland) ium sp. M. cysropreris Robinson Cystopteris bulbifera (L.) M. ptericolens Patch .Arhyrium filix-femina;

P. aquilinum

M. pteridis Wilson P. aquilinum; Adiantum concinhum ; A. m a t u m ; A. spp.

Mastopoda pteridis P. aquilinum Oestland Micromyzodium h i ferns Verma

Ill., Minn. Patch, 1938

Penn. ~hbinson, 1966 , .

N.E.N. Amer- Patch, 1938; Robinson, Colo., Brit , 1966; Lawton,.l,976 W. USA Lawton, 1976; Severin,

et a]., 1950; Robinsbn, 1966; Patch, 1938

Mmn., E. Ca- Patch, 1938; Robinson, nada-Maine-N.J. 1966; Ghosh, 1974 India Ghosh, 1974

M. filicum David Adiantum tinctum; Asplen- India ium trichomonas; Lartrea sp. ; Nephrolepis sp. ;

* Pityrogramma peruvicma; Polypodium sp. ; Pteris cretica; Streptocarpus sp.

Micromyzus judenkoi Cheilanthes sp. Carver M. nigrum Van der Goot Adiantum ; ferns

Ghosh, 1974

India, Sri Lan- Ghosh, 1974 ka, Australia India, Sri Lan- Ghosh, 1974; ka, Java Patch, 1938

M.J. Balick et al.

Appendix I (continued)


HEMIPTERA: HOMOPTERA (continued)

Aphididae (continued):

Myzus (= Neomyzus) circumfkxus Buckton (P)

Adiantum concinnum; A. cuneatum ; A. hybridum; Cyrlomium falcalum; ornamental ferns ferns ferns Marsilea quadrifolia; M. vestifa

Ill., Calif., France

Severin et al., 1950; ,

Patch, 1938 ; Schneider, 1966

M.filicis Basu M. omatus Liang (P) M. persicae Sulzer (P)

India Cosmopolitan Colorado

Ghosh, 1974 Ghosh, 1974 Patch, 1938

M. polypodicola Takahashi

Dryopleris arida; D. parasilica; Polysrichum

ferns Polystfchum; Woodwardia radicans

Sumatra, For- mosa Britai3 Formosa, Japan

Patch, 1938

M. pterisoides Theobald Patch, 1938 Patch, 1938; Moritsu, 1952 Robinson, 1966

M. woodwardiae Takahashi Papuhphb sleesmani (Pepper)

N.Y., Pem.

Penralonia nigronemsa Coquerel Phylhphb fagi L.

ferns

A thyrium Jilix-femina Switzerland Cystopteris monrana USA

Patch, 1938 Patch, 1938 Rhopalosiphum s faphyleae

Koch R. nymphaeae L. Azolhfiliculoides; Marsilea' Brit, Formosa

quadrifolia; Salvinia natans Patch, 1938

Shinjia preridifoliae (Shinji) (0) Unidentified sp.

Polypodium sp. ; Pferidium - India, Japan, aquilinum ' S.E. Asia, Aus-

tralia

Ghosh, 1974'; Sorin, 1962

P. aquilinum New Guinea Kirk, 1977

Areopodidae (= Delphacidae) : Criomorphus pferidis P. aquilinum Brit. Is. (Spinola) Nesodyne (= Ilburnia) Sadleria cya fkoides; Hawaii ipomoeicola (Kirklady) (P) Cibotriun sp. N. amamau Muir Sadleria cyatheoides Hawaii N. nephrolepidis (Kirk.) Nephrolepis exaltata Hawaii Nofhorestias badia Muir ferns Hawaii N. swezeyi Muir Aspidium fern Hawaii Nesoresfias filicicola Ehphoglossum gorgoneum; Hawaii Kirk. Ciborium; ferns

Swezey, 1922

Swezey, 1922 Swezey, 1922 Swezey, 1922 Zimmerman, 1948 Swezey, 1922; Zimmerman, 1948

N. nimbota (Kirk.) Phegopreris Hawaii Swezey, 1922; Zimmerman, 1948 .

Cercopidae: . Philaenus spumarius (L.) P. aquilinum (PI

Cicadellidae : Balclurha (= ~ i c a d u G Ciborium camissai kihveae (Kirk.)

Brit. Is., Eur. Lawton. 1976; Wienorek, 1973




Arthropod taxon Fern host Locality

Calladonus commism P. aquilinum Calif. (Van Duzee) Friscanus intricafw (Bell) P. aquilinum Calif. Cicadella (= Euptetyx) ferns Ill.

flavoscuta Cica&lJa nigra (Osborn) Polystichum adiantiforme Fla. C. omani (Christian) Polystichum adinntiforme Fla.

Nesophrosyne mysines Sadleria Hawaii '

Kirk. (0)

N. obliqua Osborn (0) Sadleria Hawaii N. umbrotilis Kirk. Microlepia strigosa Hawaii

Nesophryne kaiamamao Microlepia sfrigosa Hawaii Kuk. (0 ) (= N.filicola Kirk.; N. microlepiae Kirk.)

Reference

DeLong et al., 1948

DeLong et al., 1948 DeLong, 1948

Meade et al., 1%8 Meade et al., 1968

Zimmerman, 1948

Zimmerman, 1948

Zimmerman, 1948

Zimmennan, 1948; Swezey, 1922

Cuiidae: Iolania perkinsi Kuk. ferns Hawaii Zimmerman, 1948;

Swezey, 1922 Zimmerman, 1948 Swezey, 1922

I. spp. ferns

OIiarus knonohi Kirk. tree ferns ( 4 (0) 0. filicicola Kirk. (=O. tree ferns; Cibotium montivagus Kirk.) (d)

Hawaii

Hawaii

Hawaii Swczey, 1922 Zimmerman, 1948 Swezey, 1922; Zimmerman, 1948 Zimmerman, 1948

0. haleakaloe Kirk. Cibotium chamissai Hawaii

0. hakhaku Gifford ( 4 (0) 0. immaculatus Giff. 0. Koanoa Kirk. (0) 0. Koele Gift 0. opum Kirk. ,

0. pele ,Kirk. (d) (0)

Cibotium; Sadleria Hawaii

ferns tree fern ferns

Nephrolepis exaltata

tree ferns

Hawaii Hawaii Hawaii

Hawaii

Hawaii

Zimmerman, 1948 Zimmerman, 1948 Zimmerman, 1948

Zimmerman, 1948

Zimmerman. 1948

Coccidae : ferns (including ornamentals)

Brit. Is., USA

N. Zealand

Coccus (= Lecanium hesperidum (L.) (P)

Newstead. 1903; Dodge et at., 1948

Ctenochiton depressus Cyafhea Mask.

Maskell, 1887

Eucalymnatus tessellatus ferns Hawaii (Sign.) Eulecanium (= Lecanium) Alsophila colemoi N. Zealand mori (sign.) News feadia colka;ti Blechmcm spicnnf; Poly- Bklgium Gh-4. 0 ) (P) stichum juniperinurn

Zimmerman, 1948

Maskell, 1887

Ghesquiere, 1934

Diaspididae : Calif. Essig, 1915

Fla. Dekle, 1965

Aspidiotus hederae (Vallot) (P)

ferns

Chrysomphalus ficus Ashmead

ferns

M.J. Balick et al.


Arthropod taxon Fern host -

Locality Reference

HEMIPTERA: HOMOPTERA (continued)

Diaspididae (continued) :

C. dictyospermi (Morgan) ferns Hawaii (P) Fiorinia fwriniae (Targ.) ferns Calif. ,

(PI Hemiberlesia rapax Akophila australis R. Br. Fla. (Comstock) Parlatoriaproteus (Curtis) Asplenium nidus Fla. Phenacaspis ( = Chionaspis) Asplenium N. Zealand dubia (Mask.). Pinnaspis (= Chionaspis; Adianhun sp. ; Asplenium Cosmopolitan Hemichionaspis) n i h ; A. spp.; Cheilanthes aspidistrae (Signoret) (P) sp. ; Cyrtomium falcatum

Presl. ; Davallia fejeensis 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 (Cooky)

Polinspis media Mask. ferns N. daland Pseudoparlatoria parlo- ferns Calif. torioides (Comstock) (P) P. pentagons (Targ.) (P) ferns Calif.

Eriococcidae: Eriococcus insignis New- Pteris sp. ; Pteridium Europe. W. Asia stead (P) aquilinum

Lecaniidae : CeroplatesjloridensB ferns (including orna- Calif. Comst. (P) mentals) C. rubens Mask. Asplenium; Elaphoglossum Hawaii

reticulorum ; ferns Pulvinaria floccifera ferns Calif., Ore, B.C. Westw. P. mammeae Mask. (P) ferns Hawaii P. psidii Mask. (P) ferns Hawaii Saissetia cofleae Wlk. (P ) Adiantwn spp. ; Asplenium Cosmopolitan (= Lecanium hemisphaeri- spp. ; Pteris spp. ; Nephro- mm Uarg.)) lepis; ferns

Zimmerman, 1948

Dekle, 1965

Dekle, 1965 Maskell, 1887

Dekle, 1965 ; Wieczorek, 1973; Newstead, 1901 ; McKensie. 1956 ; Werner, 1931

Dekle, 1965; Zimmerman, 1948

Maskell, 1887 McKensie, 1956

McKensie, 1956

Hoy, 1963

Essig, 1915; Dodge et a]., 1948 . Zimmerman. 1948

Steinweden, 1946

Zimmerman, '1 948 Zimmerman, 1948 Newstead, 1903; Cory, 1945; Evans, 1942; Speyer et al., 1938; Essig, 1915; Zimmer- man, 1948




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 ( R i m ) (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 I Newstead

Ricaniidae : Scolypopa australis N. Zealand Cumber, 1966 Walker (P)

ferns

HYMENOPTERA

Blasticotomidae: ~lasficotoma Pliceri Klug Athyrium filk-femina; Dry- Palearctic (m) (g) opreris spp. ; Matteuccia

srruthiopteris (L.) ; P. aquilinwn; Polystichum sp.

Cynipidae : Unidentified sp. (g)

Formicidae (assoc.- " nectary " feeders) :

P. aquilinurn

ferns ; P. aquilinum

Tenthredinidae : Aneugmenus (= Selandria) A fhyrium; Dryopteris; coronatus (Klug) P. aquilinum

A.fivipes (Norton) P. aquilinum

A. furstenbergensis (Konow) A. padi (L.)

P. aquilinum; ferns

P. aquilinum ; ferns

A. starnineipes Klug (g) P. aquilinum

Aneugmenus temporalis P. aquilinum; ferns (Thornson)

Europe

Cosmopolitan

Engl., Eur., Siberia USA

Brit. Is., Eur.

Brit. Is., Eur., Siberia' Eur., Brit Coi.

Brit. Is., Eur., Siberia

USA

Benson, 1951 ; Schedl, 1973

D. Van Leeuwen, 1938

W.L. Brown (p.c.); Lawton, 1976

Lawton, 1976; Wieczorek, 1973 .

Hogh, 1966

Lawton. 1976; Wieczorek, 1973 Lawton, 1976; Wieczorek, 1973 D. Van Leeuwen, 1938; Ross, 1932

Lawton, 1976; Wiecwrek, 1973

Hogh, 1966

M.J. ~ a l i c k et a].

Appendii 1 (continued)


HYMENOPTERA (continued)

Tenthrdinidae (continued):

Emprin excisa Thornson P. aquilinum Brit. Is., Eur., (P) Caucasus Hepfamelus ochroleucus Aihyrium spp. ; Blechnum; Brit., Eur. Stph. (m) Polypodium vulgare ; P. sp. ;

Pferidium aquilinum

Pseudohemi&xonus sharpi ferns (Caeron) Strombocerus delicalulus Athyrium; Dryopleris; (Fallen) Mar teuccia; Onoclea;

Polypodium ; Polystichum ; Pteridium aquilinum

Slrongylogasfer contigua P. aquilinum (Konow) S. dirtam Norton P. aquilinum S.Jlicb Klug P. ap4"Iinum ; ferns

S. lineata (Christ) Dryopferis; Matleucia ; P. aquilinum ; ferns

S. manrlata (Klug) Afhyrium; P. aquilinum

S. mix& (Klug) Afhyrium; Dryopferb; P. aquilinum

S. multicinctus Norton P. aquilinum

S. struthiopteridis Matfeuccia (Forsius) S. tibialis Cresson P. aquilinum

S. xanfhoceros (Stephens) P. aquilinum

S. spp. P. aquilinum Tenthredo balieata Klug P. aquilinwn (PI T. colon Klug (P) P. aquilinum

Tenthredo ferruginea Schrank (P) T. Iioido L. (P)

T. spp.

LEPIDOPTERA

Arctiidae :

Arctia caja (L.) (P) Dincrkia pteridis H. Edwards (P)

P. aquilinum

P. aquilinum

P. aquilinum

P. aquilimm

P. aquilinum

Europe

Brit. Is., Eur., Kamchatka

Lawton, 1976; Wiecwrek, 1973

Lawton, 1976; Wieczorek, 1973; Stansfield, 1933

Wieczorek, 1973


Eur.-N. Africa- Lawton, 1976; Kamchatka Wieczorek, 1973 Calif.-Brit. Col. Beer, 1955

Europe-Japan Wieczorek, 1973 Brit. Is., Em.- Lawton, 1976; Central Asia- Wieczorek, 1973 Japan Brit. Is., Eur. Lawton, 1976;

Wieczorek, 1973 'Brit. Is., Eur.- Lawton, 1976; ' Kamchatka Wieczorek, 1973

USA Hogh, 1966

, Europe Wieczorek, 1973

Calif.-Brit. Col. Beer, 1955

Engl., Eur.- Lawton, 1976; China Wieczorek, 1973

USA Hogh, 1966 Europe Wieczorek, 1973

Brit Is., Eur.- Lawton, 1976; Asia-Kamchatka Wieczorek, 1973

Brit. Is., Em.- Lawton, 1976; Asia-Kamchatka Wieczorek, 1973

Brit. Is., Eur.- Lawton, 1976; Siberia Wieczorek, 1973

USA Hogh, 1966

N. Amer.

N. Amer.

Tietz, 1972

Tietz, 1972




D. oagans (Boisduval) (P) Pteris sp. N. Amer. Tietz, 1972 Rhyparioides metelkana Nephrodium fhelypteria Lederer

Rumania Leestmans, 1975

Spilosoma hrieum (Hufnagel) (P)

P. aquilinum Brit. Is. Lawton, 1976

Gelechiidae : Europe Wieczorek, 1973 Depressaria impurella P. aquilinum

Tr. (P) Palfodora cytisello Curt. P. aquilinum (g)

Engl., Ire., Eur. Lawton, 1976; Wieczorek, 1973

Geometridae: Cinquilia catenaria Dennsfaedtia punctilobula (Dm& (PI (Michx.)

N. Amer. Tietz, 1972

. Gueneria basinria Wlk. Dryopieris yoaboracensis (L.)

Homochlodes (= Lozo- P. oqruoqruIinum gramma) fritilloria (Guen.)

N. Amer. Xetz, 1972

E. Can.-Wisc.- Forbes, 1948; N.C.; Ha. Kimball, 1965

H. lacfispargaria (Walker) P. aquilinum E. Canada Ferguson, 1975 Lithina (= Apaecaria; various ferns Lozogramma) S U ~ U ~ ~ U U M (Wlk.)

E. Can.-N.J.- Forbes, 1948 wisc.

Petrophora (=Lithim; P. aquilinum; Dryopteris Phasiane petraria Hub.) chlorosara (hopoli)

Brit. I s ; Eur. . Lawton. 1976; Wieczorek, 1973

Philedia puncromacularia Pteris sp. (Hulst)

N. Amer.

Hawaii

Tietz, 1972

Scotorythra rara (Butler) ferns; Cibofium (P)

Swezey, 1922; 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 DeGeer (r) H. gracilis Grote (r) ferns

Brit. Is., Eur. Lawton, 1976

Maine-Mass.- Forbes, 1923 Colo. Brit. Is.; Eur. Lawton, 1976 Engl.; Scotl. Lawton, 1976

H. hectus L. (P) (r) P. aquilinum

H. sylvinus (L.) ( P ) (r) P. aquilinum

Lycaenidae: . Hyoochrysops theon Drynuria quercifolia (L.) Australia medocus (Fruhstorfer) J. Smith

Noctuidae: Achea janata L. (P) Polypodium sp. Hawaii Actebia fennica Dennstaedfia punctilobula N. Amer. (Tauscher) (P) (Michx.)

Daniels, 1976

Anonymous, 1945 Tietz, 1972

M.J. Balick et al.



LEPIDOPTERA (continued)

Noctuidae (continued):

Callopislria Guenee

jloridemis Adiatum sp.; A.pednrum So. USA, Fla., Comstock, 1939; 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 C.juvenrina (Cramer) P. aquiIinum C. htreilli Dup. P. aquilinwn

C. (= Eriopus) maillordi Adianrum sp. Guenee C. mollissima Guenee C. monetifera Guenee

ferns ferns

C. purpureofatciata Pill. P. aquilinum

Ceramica pisi (L.) (P) P. aquilinum

Euplexia benesimilis McDunn. (P) E. Iucipara (L.) (P )

P. aquilinum

P. aguilinum, ferns

N.J.-Fla. Forbes, 1954 Europe Wiecwrek, 1973 Eur., N. Africa Wieczorek, 1973 Thailand Cantelo. 1965

Can.-F1a.-Colo. Forbes; 1954 E. Can.-Wise.- Forbes, 1954 Fla. Europe Simmonds, 1967 Brit. Is. Lawton, 1976 Calif., Eur. Essig, 1958

Brit. Is., Ew. Lawton, 1976; Wieczorek, 1973

Fagitana liltera (Guenee) Dryopleris rhelypteris (L.); Fla., N.J. Osmunda sp.

Habrynlhis (= Phlogo- Aspidium filix-mas; , Eur. phora) scita (Huebner) Blechmm sp. ; P. aquiIinum (PI

Kimball, 1965; D. Schweitzer (p.c.) Wieczorek, 1973

Swezey, 1922

Zimmerman, 1948

Lawton, 1976

Haliophyk (= Erio- many ferns Hawaii pygodes) euclidias (Meyrick) H. spp. several ferns; Acrostichum; Hawaii

Aspidium cyatheoides Loconobirr (= Mameslra) P. aquilinum Brit. Is. 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.




P. stenocelis (Dyar) (r) Woodwardia virginica N.E. USA Forbes, 1954 (L.) Sm.

Peridroma margaritosa P. aquilinum (Haworth) (= P. saucia Huebner) (P) Phlogophora meticulosa P. aquilinum (L-) (PI Polia adjuncta (his- P. aquilinum duval) (P) .

P. assimiIis(Morrison) (P ) P. aquilinum

N. Amer. Tietz, 1972

Brit. Is.

N. Amer.

N.Y.

Lawton, 1976

T i e y 1972

J. Franclemont (p.c.)

Nymphalidae : Metamorph stelens (L.) Blechnum sp.

Olethreutidae :

Fla. ; Cuba Kimball, 1965; Dethier, 1940

Olethreutes osmunddna Osmunda regalis Fernald (0)

Maine; Wash., Forbes, 1923 D.C.

Pyralidae : Ambia? sp. Nymphulo? novaginensis Kenrick Phlyctaenh (= Pyrausta) rhcseusalb . Psara platycapnu Meyrick Pyrausta aeglealir Walker P. descrepitalis H.-S. Samea multiplicalis Guenee

Polystichum adiontiforme

P. aquilinum

Fla. KimbaU, 1965 New Guinea Kirk, 1977

various ferns E. Can.-Fla. Forbes, 1923

P. aquilinum

Osmunda Aspidium spinulosum

Salvinia sp. ; S. rotund~olia Azolla caroliniana; Pbtia stratiotes Polystichum adiontiforme P. aquilinum

New Guinea Kiik, 1977 N.Y. Forbes, 1923 Engl. Ford, 1949 Trinidad, No. S. Thomas, 1975; Knopf Amer., Ha. et al., 1976

Undubmbia polystichalis Capps Unident*ed sp.

Fla. Kimball, 1965 New Guinea Kirk, 1977

Antherea eucalypti Nephrolepis Scott (PI

Australia French, 1933

Opogonu chlinata Meyr. Polypodium quercifolium India ( a m - )

Maxwell-Lefroy, 1971

Praecedes thecophora P, aquilinum Wals. Teichobia filicinora Meyr. Aspidium filix-mas

New Guinea

Engl.

Kirk, 1977

Hering, 1951 ; Ford, 1949

T. verhuellaella Stt. Several Polypodiaceae ; Engl. . Ford, 1949

Unidentified sp. 1 (m) P. aquilimun

Unidentified sp. 2 (m) P. aquilinum

New Guinea New Guinea

Kirk, 1977 Kirk, 1977

Tortricidae : Tortrix ciranu Fernald (P) ferns Calif. Essig, 1915

M.J. Balick et al.



LEPIDOPTERA (continued)

Yponomeutidae (= Walshiidae) : Bafrachedra sophroniella Aspidium cyafheoides Hawaii Walsingham(s) B. Iomenfella Wals. fern Hawaii B. bedeliella Wals. (s) Asplenium nidus;Elapho- Hawaii

glossurn reticukmm; Dryo- pteris parasilica

B. sp. (m) Pteris irregulnris Hawaii B. SPP. (4) (?m) Polypodium spectrum Hawaii

(?part) Euhyposmocoma ekaha Asplenium nidus Hawaii (Swezey) E. trivitella Swezey (m) Elaphoglossum reticulafum Hawaii Hyposmocoma spp. ferns (several)

ODONATA

Anfiagrion grinskrgsi BIechnum chilense Jurzitza (oviposits only) (assoc.) Megalagrwn oahuense Gleichenia linearis (Blackburn) (truly terrestrial, lives in fern litter) (assoc.)

ORTHOPTERA Acrididae: Paulinia acumimta (De Geer) Unidentified sp.

Gryllidae: Lepfogrylhu nigrolineatus Perkins (0) Para frigonidium Jlicum Perkins P. viridescens Perkins

Progmrhogryl~us atatus Brunner (P)

Tetigoniidae : Banza spp. ( P )

Salvinia

P. aquilinum

Cibotium

"tall soft ferns";

Hawaii

, C hie

Swezey, 1922

Swezey, 1922 Swezey, 1922

Swezey, 1922 Swezey, 1922

Swezey, 1922

Swezey. 1922 Swezey, 1922


Trinidad, No. S. Amer. New Guinea

Hawaii

Hawaii

"creeping fern"; Lindsaya Hawaii macraeana ? ferns ; Cibotiwn Hawaii

ferns Hawaii

Thomas, 1975

Kirk, 1977

Zimmerman, 1948

Swezey, 1922

Swezey, 1922; Zimmerman, 1948 Swezey, 1922 ; Zimmerman, 1948

Swezey, 1922




THYSANOPTERA

Phlaeothripidae: Atractothrips macrurus ferns Okajima (d) (P)

Japan (Ryukyu Okajima, 1975 Is.)

Haplothrips hawaiiensis Cibotium menziesii; tree Hawaii Moulton (d) (0) ferns H. rosai Bianchi (P) Sadleria Hawaii Kamyolrips doliicornis Sadleria Bianchi (P) K melaleuca (Bagnell) ( P ) Ciborium chamissoi Uredothrips tibialis ferns Okajima (d)

Hawaii

Zimmerman, 1948

Zimmerman, 1948 Zimmerman, 1948

Hawaii Zimmerman, 1948 Japan (Ryukyu Okajima, 1975 Is.)

Heliothrips haemor- ferns (including o m - Calif. rhoidalis Bouche (P) mentak); Pteris cretica Leucothrips (=Micro- Asplenium sp. Mich. rhrips) Ieucus Herrick L. nigripennis (Reuter) ferns (including orna- USA

mentals) ; Pteris cretica Physorhrips (= Pterido- Polypodium pteropus Moore Java thrips) pferidicola (Karny) (g)

Zonothrips osmunahe Osmunda cinnnmomaea N.J. Crawford Asplenium nidus; Pleopeltis Indonesia UnidenWied spp. (g) superfacialis Bedd.

. Essig, 191 5; Dodge et al., 1948 Herrick, 1927

Lewis, 1973

Priesner, 1938

Crawford, 1941 ; D. Van Leeuwen, 1938

References

Anonymous: Insect pests and their control. Agric. Gaz. N. S. W. 51, 15&154 (1940) Anonymous: Notes and exhibitions presented at meetings of the Hawaiian Entomological Society.

Proc. Hawaiian E n t Soc. 12,46M92 (1945) Augustijn, C., Verkade, J.: De rouwvlieg-larvae als beschadigerivan jonge varens. Tijdschr. Pliekt.

41,301-308 (1935) Banks, H.P.: Evolution and plants of the past In: Fundamentals of botany series. London: Macmil-

Ian 1972 Beer, R. : Biological studies and taxonomic notes on the genus Srrongylogaster Dahlbom. (Hymenop

tera: Tenthredinidae). Univ. Kans. Sci. Bull. 37, 223-249 (1955) Benson, R.: Handbooks for the identificati,on of British insects. VI. Hymenoptera, Symphyta.

Sect. a. London: Roy. Ent.Soc. 1951 Bonnemaison, L.: Un aphide nuisible aux fou geres nouveau pour la France. Rev. Path. Veg.

24, 163-171 (193'7) Braun, A.: New species of microlepidoptera. Can. En t 50.229-251 (1918) Brues, C., Melander, A., Carpenter, F.: Classification of Insects. Bull. M.C.Z. 108, 1-917 (1954) Butler, E.: A biology of the British Hemiptera-Heteroptera. London: Witherby 1923 Cantelo, W.: A host list of the insects of Thailand. U.S. Operations Mission to Thailand and

Dept. Agric., Royal Thai Government (1965)

M.J. Balick et al.

Carlisle, D., Ellis, D.: Bracken and locust tcdysones: their eifects on moulting in the Desert Locust. Science 159, 1472-1474 (1968)

Comstock, J.A.: Studies in Pacific coast Lepidoptera. Bull. S. Calif. Acad. Sci. 38, 34-35 (1939) Cooper-Driver, G.: Chemotaxonomy and phytochemical ecology of bracken. Bot. J. Linn. Sot.

73, 35-46 (1976) Cooper-Driver, G., Finch, S., Swain, T., Bernays, E.: Seasonal variation in secondary plant corn-

pounds in relation to the palatability of Pteridiwn aquilinwn, Biochem. Syst. Ecol. 5, 177-183 (1977)

Cooper-Driver, S., Swain. T.: Cyanogenic polymorphism in bracken in relation to herbivore predation. Nature 260, 604 (1976)

Cory, E.: Control of several scales infesting orchids. J. Econ. Ent. 38. 395 (1945) Crawford, J.: The genus Zonothrips in N. America. Proc. Ent. Soc. Wash. 43, 105-107 (1941) Cumber, R.: Factors influencing population levels of Scolypopa australis Walker (Hemiptera-Ho-

moptera: Ricaniidae). N. Zeal. J. Sci. 9, 336356 (1966) Daniels, G.: Life history of Hypochrysops theon medocus (Fruhstorfer) (Lepidoptera: Lycaenidae).

J. Austral. Ent. Soc. 15, 197-200 (1976) Dekle, G.: Arthropods of Florida and neighboring land areas. Fla. Dept. Agric. 3, Contrib. 60

(1965) DeLeon, D.: A new fern mite from Trinidad, West Indies (Acarina: Tarsonemidae). Ha. Ent.

49, 127-129 (1966) DeLong, D.: The Leafhoppers, or CicadeUidae, of Illinois (Eurymelinae-Balcuthinat), Bull. Ill.

Natur. Hist. Surv. 24.97-376 (1948) DeLong, D., Severin, H.: Characters, distribution, and food plants of leafhopper species in Than-

norettix group. =lgardia 18, 185-199 (1948) Dethier, V.: L ie histories of Cuban Lepidoptera. Psyche 47, 14-26 (1940) Docters Van Leeuwen, W.: Zoocecidia. In: Manual of pteridology (F. Verdoorn, ed.), pp. 192-195.

The Hague: Nijhoff 1938 Dodge, B., Rickett, H.: Diseases and pests of ornamental plants, 638 pp. New York: Ronald 1948 Eastop, V.: Deductions from the present day host plants, of aphids and related insects, In: Insect

plant nlationships, No. 6 (H. Van Emden, ed.), pp. 167-178. London: Symposia of the Royal Entomological Soeciety 1973

Ehrlich, P., Raven, P.: Butterflies and plants: a study in coevolution. Evol. 18, 586-608 (1964) Essig, E.: Injurious and beneficial insects of californii. Suppl. Mon. Bull., Calif. St. Comm.

Hart 1, 1-541 (1915) Essig, E.: Insects and mites of western North America, 1050 pp. New York: Macmillan 1958 Evans, I.&: Relationship between bracken and cancer. Bot. J. Linn. Soc. 73, 105-112 (1976) Evans, J.: Orchard and garden. Mites and their control. Scale insects. Tasm. J. Agric. 13, 156159

(1942) Evans, W.C: Bracken thiaminase-mediated neurotoxic syndromes. Bot. J. Linn. Soc. 73, 113-132

(1976) Eyjolfsson, R : Recent advances in the chemistry of cyanogenic glycosides. hog. Chem. Org. Natural

hods. 128, 74 (1970) Eyles, A., Ashlock, P.: The genus Nysius in New Zealand (Heteroptera: Lygaeidae). N. Zeal.

J. Sci. 12,713-727 (1969) Feeny, P.: Seasonal changes in oak leaf tannins and nutrients as a cause of spring feeding by

winter moth caterpillars. Ecology 51, 565-581 (1970) Felt, E.: Plant galls and gall makers., 364 pp. Ithaca: Comstock 1940 Ferguson, D.: Host records for Lepidoptera reared in eastern North America. U.S.D.A. Tech.

Bull. No. 1521, 49 pp. (1975) Forbes, W.: Lepidoptera of New York and neighboring states. Mem. Cornell Univ. Agric. Expt.

Sta. No. 68; Primitive forms, Microlepidoptera, Pyraloids, Bombyces, 729 pp. (1923). No. 274; Geometridae, Sphingidae. Notodontidae, Lymantriidae, 263 pp. (1948). No. 329; Noctuidae, 433 pp. (1954)

Ford, L.: A guide to the smaller British Lepidoptera, 230 pp. London: The South London En- tomological Natural History Society 1949

French, C: Victorian insects. New record of plants attacked by insects. Vict. Nat. 49, 234 (1933) Gagne, W., Howarth, F.: The cavericolous fauna of Hawaiian lava tubes: 7. Emesinae or thread-

legged bugs (Heteroptera: Reduviidae). Pac. Ins. 16,415-426 (1975)


GhesquiBre, J.: Newsteadio collarti n. sp. Coccidae (Ortheriinae) radicicole et mycophile de la fo6t de Soignes. Bull. Ann. Soc. Ent. Belg. 74, 27-32 (1934)

Ghosh, A.: Fern infesting aphids (Insects: Homoptera) in Indian. J. Hort. 31. 106109 (1974) Gilbert, L., Raven, P. (eds.): Coevolution of animals and plants. Austin-London: Univ. Texas

Press 1975 Hardy, D.: Diptera: Brachycera 11-Cyclorrhapha I. In: Insects of Hawaii, Vol. 11, 458 pp.

Honolulu: Hawaii Press 1964 Harper, N., Cooper-Driver, G., Swain, T.: A survey for cyanogenesis in ferns and gymnosperms.

Phytochemistry 15, 1764-1767 (1976) Hendrix, S.: The resistance of Pteridiwn aquilinwn (L.) Kuhn to insect attack by Trichoplusia ni

(Hub.). Oecologia (Berl.) 26, 347-361 (1977) Hering, E.: Biology of the leaf miners, 420 pp. Gravenhage: Uitgeverij Junk 1951 Herout, V.: Some relations between plants, insects and their isoprenoids. In: Progress in phytochem-

istry (L. Reinhold, Y. Liwschitz, eds.), pp. 143-202. London: Interscience 1970 Herrick, G.: Two new species of Thrips (Thysanoptera). Ent. News 38, 276-281 (1927) Heugens, A.: Lopidocyrtus pallidus Reuter in de varenteelt. Meded. Rijksfac. LandbWetensch.

Gent. 32,243-251 (1967) Hogh, G.: Local distribution of a fern sawfly Strongylogosier multicinctus (Tenthredinidae). Kans.

Ent. Soc. J. 39, 347-354 (1966) Holman, J.: Aulacorthum dryopteridis n. sp. and Aphis torquens n. sp., two new aphid species

from Czechoslovakia (Homoptera: A~hidibe). Acta Soc. Ent. Czech. 56, 221-236 (1958) Hoshizaki, B.: Fern growers manual, 256 pp. New York: Knopf 1975 Hoy, J.: A catalogue of the Eriococcidae (Homoptera: Coccoidea) of the world. N. Zeal. Dept.

Sci. Ind. Res. Bull. No. 150 (1963) Janzen, D.H.: Interactions of seeds and their insect predatorslparasitoids in a tropical deciduous

forst. In: Evolutionary strategies of parasitic insects and mites (P.W. Price, ed.), pp. 154-180. New York: Plenum 1975

Jeppson, L., Kiefer, H., Baker, E.: Mites injurious to economic plants, 614 pp. Los Angeles: Univ. Calif. Press 1975

Jones, D.A.: Co-evolution and cyanogenesis. In: Taxonomy and ecology (V.H. Heywood ed.), pp. 213-242. London: Academic Press 1973

Jurzitza, G.: Antiagrion gayi (Selys, 1876) and A. grinsbergsi n. sp., two closely related species from Chile (Zygoptera: Coenagrionidae). Odonatologica (UTR) 3,221-230 (1974)

Kevan, P., Chaloner, W., Saville, D.: Interrelationships of early terrestrial arthropods and plants. Palaeontology 18, 391417 (1975)

Kiefer, H.: Eriophyid studies, VIII. Bull. Calif. Dept. Agric. 29, 21-46 (1940) Kimball, C.: Lepidoptera of Florida. Arthropods of Florida and neighboring land areas. Fla.

Dept. Agric. 3 (1965) Kirk, A.: J. Austr. Ent. Soc. 16,403-409 (1977) Knight, H.: The plant bugs, or Miridae, of Illinois, Bull. Ill. Natur. Hist. Surv. 22, 1-234 (1941) Knopf, K., Habeck, D.: Life history and biology of Samea multiplicalis (Lepidoptera). Environ.

Ent. 5, 539-542 (1976) Kormilev, N.: Los insectos de 1% Islas Juan Fernanda Lygaeidae. Rev. Chil. Ent. 2, 7-14 (1952) Kuschel, G.: Insects of Campbell Island. Curculionidae. Pac. Ins Monogr. 7. 416493 (1964) Lawton, J. : The structure of the arthropod community on bracken. Bot. J. Linn. Soc. 73, 187-216

(1976) Leech, H.: The bug Corythuca padi on bracken fern (Hemiptera: Tingidae). Proc. Ent. Soc. Brit.

Columbia 41, 36 (1944) Leestmans, S.: Discovery of a new station of Rhyparwides metalkana Lederer and new data on

its biotype in Romania (Arctiidae). Linn. Belg. 6, 102-104 (1975) Lewis, T.: Thrips, their biology, ecology and economic importance, 349 pp. London: Academic

Press 1973 Linnavuori, R.: Hemiptera of the Sudan, with remarks on some species of the adjacent countries.

4. Miridae and Isometopidae. Ann. Zool. Fenn. 12, 1-1 18 (1975) Linssen, E.: Beetles of the British Isles, Ser. 2, 295 pp. London: Warne 1959 Lunsden, W.: Natural history of Canna and Sanday, Inner Hebrides. Proc. Roy. Physical Soc.

23.33 (1939) Mackay, M. : Lepidoptera in Cretaceous Amber. Science 167, 373-380 (1970)

M.J. Balick et al.

Maskell, W.: An account of the insects noxious to agriculture and plants in New Zealand, 116 pp. Wellington: Didsbury 1887

Maxwell-Lefroy, H., Howlett, F.: Indian insect life, 786 pp. New Dehli: Today &Tomorrow 1971 May, B.: A new species of Megacolabus and descriptions of the immature stages of M. decipiens

(Coleoptera: Curculionidae). J. Roy. Soc. N. Zeal. 3, 255-262 (1973) McKensie, H.: The armored scale insects of California. Bull. Calif. Ins. Surv. 5 (1956) McKensie, H.: Mealybugs of California, 525 pp. Los Angeles: Univ. Calif. Press 1967 Meade, F., Habeck, D.: Ent. Cir. no. 78. Fla. Dept. Agric., Div. PI. Ind. (1968) Moritsu, M.: The fern aphids in Japan. Yamaguti Univ. Facul. A@. B. 3,25-28 (1952) Mound, L.: A new genus and four new species of white flies from ferns. Rev. 2001. Bot. Afric.

64, 127-132 (1%1) Needham, J.: Moth larvae living on fern spores. Proc. Ent. Soc. Wash. 49, 165-166 (1947) Newstead, R.: Monograph of the Coccidae of the British Isles, Vol. 1. London: The Ray Society

1901 Newstead, R.: Monograph of the Coccidae of the British Isles, Vol. 2. London: The Ray Society

1903 Okaijima, S.: Notes on the Thysanoptera from the Ryukyu Islands. I. Descriptions of two new

species. Kontyu 43, 13-19 (1975) Patch, E.: Food-plant catalogue of the aphids of the world, including the Phylloxeridae. Maine

Agric: Expt. Sta. Bull. 393, 3-31 (1938) '

Perring, F.H., Gardiner, B.G. (eds.): The biology of bracken. Bot J. Linn. Soc. 73, 1-284 (1976) Priesner, H.: Materialien m einer Revision der Taeniothrips-Arten (Thysanoptera) des Indo-Malayi-

schen Faunengebietes. Treubia 16,469-526 (1938) Pritchard, A.: The fern mite. Calif. Agric. 5, 10 (1951) Rhoades, D.F., Cates, R.G.: A general theory of plant antiherbivore chemistry. In: Recent advances

in phytochemistry. Vol. 10. Biochemical interaction between plants and insects (J.W. Wallace, R.L. HanseU, eds.), pp. 168-213. New York-London: Plenum Press 1976

Richards, W.: Three new species of Aulacorthum from British Columbia, with a key to the Canadian species (Aphididae: Homoptera). Can. Ent. 104, 1017-1023 (1972)

Richerson, P., Grigarick, A.: The life history of Stenopelmus ~ w f i s (Coleoptera: Curculionidae). Ann. Ent. Soc. Am. 60,351-354 (1%7)

Robbins, W.. Kaplanis, J., Thompson, N., Shortino, T., Cohen, C., Joyner, S.: Ecdysones and analogs : effects on development and reproduction oc insects. Science 161, 1 158-1 160 (1968)

Robinson, A.: Review of the fern aphids of North America with a description of a new species and a new genus. Can. Ent. 98, 1252-1259 (1966)

Ross, H.: Records of additional European sawflies in America and descriptions of new varieties in North American species. Can. Ent. 64, 247-251 (1932)

Schedl, K.: The scolytids and platypodids of Africa, 111. Rev. Ent. Mocamb. 5, 595-1352 (1962) Schedl, W.: Erster Nachweis der Farnblattwespe Blasticotony fdiceti Klug, 1934, in Osterreich

(Hymenoptera: Blasticotomidae). Arbeitsgem Osterr. Ent. 25, 114-117 (1973) Schneider, J.: Ennemis de fougeres ornamentales. Phytoma 18 (182). 2632 (1966) Severin, H., Tompkins, C. : Symptoms induced by some species of aphids feeding on ferns. Hilgardia

20, 81-92 (1950) Simmonds, S.: Chirosia crussiseta Stein (Muscidae) on bracken in Lanchashire. Ent. Mon. Mag.

100. (1 198/1199), 80 (1964) Simmonds, S. : Possibilities of bio-control in bracken Pteridium aquilinum (L.) Kuhn (Polypodiaceae).

P.A.N.S. Sect. C. Weed Contr. 13, 200-203 (1967) Smart, J., Hughes, N.: The insect and the plant: progressive palaeoecological integration. In:

Insect plant relationships, No. 6 (H. Van Emden, ed.), pp. 143-155. London: Royal Entomolog- ical Society 1973

Soo Hoo, C., .Fraenkel, G.: The resistance of ferns to the feeding of Prodenia eridania larvae. Ann. Ent. Soc. Am. 57,788-790 (1964)

Sorin, J.: Notes on the aphid Shhjia pteridgoliae Shinju (Aphididae: Homoptera). Kontyu 30, 21-26 (1962)

Southwood, T. : The insect plant relationship: An evolutionary perspective. In: Insect plant rdation- ships, No. 6 (H. Van Emden, ed.), pp. 3-30. London: Royal Entomological Society 1973

Speyer, E., Read, W., Orchard, 0.: Animal pests. Rep. Exp. Res. Sta..Cheshunt. 23, 59-65 (4938) . Stansfield, F.: Another fern pest. Brit. Fern Gaz. 6, 237-238 (1933)

biochemical and evolutionary aspects of arthropod ... · climates. although all ferns tested...

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