Leaf-Mining Chrysomelids 1 Leaf-Mining Chrysomelids

Leaf-mining chrysomelids 1 Leaf-mining chrysomelids

Jorge A. Santiago-Blay Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA

“There are more things in heaven and earth, Horatio, than are dreamt of in your philosophy.” (Act I, Scene 5, Lines 66-167) “To be or not to be; that is the question” (Act III, Section 1, Line 58) both quotes from “The Tragedy of Hamlet, Prince of Denmark” by William Shakespeare (1564-1616)

Abstract into two morphological categories: the eruciform, less modi- fied type (Galerucinae and some Alticinae); and the flattened, Leaf-mining is the relatively prolonged consumption of foliar sometimes onisciform type characteristic of the Zeugophorinae, material contained within the epidermal layers, without elicit- many Alticinae, the Cassidinae, and the Hispinae. There are ing a major histological response from the plant. This type of no published data on the larval structure of leaf-mining herbivory is relatively uncommon in the Chrysomelidae and criocerines. Larval leaf-mining chrysomelids are reported to has been reported in 103 genera, representing 4% of the ap- have rather broad host-plant feeding preferences. For adults, proximately 2600 described genera and amounting to over the ranges are broader. The Index of Feeding Range (IFR) is 500 reported species, or 1-2% of the 40-50,000 described introduced herein as a scalar to quantify the feeding range of species. Larvae in the following subfamilies are known leaf- the larvae (IFRi) and adults (IFRa). For the Zeugophorinae, miners, with numbers and percentages of taxa also being in- IFRi is 2.0 and IFRa 2.9. The plant families (and genera, cluded. The subfamily Zeugophorinae consists of one genus parenthesized) most commonly reported serving as host-plants with reported leaf-mining species, or 25% of the total genera; for the Zeugophorinae are the Salicaceae (Salix and Populus), there are twelve reported leaf-mining species, about 17-20% the Betulaceae (Betula and Corylus), and the Celastraceae. of the species described in the subfamily and about 60-70 For adult zeugophorines, 55% of the reported species only described species in the Zeugophorinae, all believed to be feed on one plant genus, and 82% of the reported species feed leaf-miners. The subfamily Criocerinae comprises two genera on one plant family only. For the Galerucinae, IFRi is 1.0 and of reported leaf-miners, representing 10% of the described IFRa 2.4. The plant families (and genera, parenthesized) most genera, and two reported leaf-mining species, accounting for commonly reported serving as host plants for the Galerucinae less than 1% of the approximately 1450 species described in are the Asteraceae (several genera) and the Chenopodiaceae the Criocerinae. The Galerucinae has two reported leaf-min- (Atriplex, Chenopodium, Suaeda, etc.). For adult galerucines, ing genera, representing less than 1% of the approximately 32% of the reported species only feed on one plant genus, and 500 genera described. There are approximately 20 reported 60% of the reported species only feed on one plant family. For species of leaf-mining galerucines, accounting for less than the Alticinae, IFRi is 2.7 and IFRa (excluding the data for 1% of the approximately 7000 species described in the sub- Phyllotreta nemorum) is 3.8. The plant families most com- family. The Alticinae has nineteen reported leaf-mining gen- monly reported to be serving as host plants for the Alticinae era, representing about 3% of the approximately 500 genera are Brassicaceae, Lamiaceae, Asteraceae, Plantaginaceae, described. There are 65 reported species of leaf-mining flea- Scrophulariaceae, Polygonaceae, and Poaceae, but many more beetles, or about 1-2% of the 4000-8000 species described in families and numerous genera are reported as host plants. For the Alticinae. The Hispinae is represented by 78 genera that adult alticines, 47% of the reported species only feed on one have been reported as leaf-miners, or 40% of the approxi- plant genus, and 71% of the reported species only feed on one mately 200 genera described. There are over 400 reported plant family. For the Cryptostomes (Hispinae + Cassidinae), leaf-mining hispines, accounting for 14% of the over 3000 IFRi 1.6 and IFRa is 3.02. The plant families (and genera, species described in the subfamily. There is a single reported parenthesized) most commonly reported as host plants for the genus of leaf-mining in the Cassidinae, representing less than Hispinae in the Old World are Arecaceae (Cocos, Metroxylon, 1% of the 159 genera described. Only six species of cassidines and numerous other palm genera), Pandanaceae (Pandanus have been reported as leaf-miners, accounting less than 1% of and Freycinettia), and Zingiberaceae. Numerous Leguminosae, the 2760 species in the Cassidinae. The reported geographical Asteraceae, Poaceae, and Verbenaceae have been reported as distribution and host plants are summarized for most of the host plants for the Hispinae in the New World. For larval over 500 species of suspected or documented leaf-mining Cryptostomes, 77% of the reported species feed on one plant chrysomelids (Table 1). Larval chrysomelids can be classified species. In adult Cryptostomes, 51% of the reported species

Address for correspondence: Jorge A. Santiago-Blay, Department of Paleobiology, National Museum of Natural History, MRC- 121, Smithsonian Institution, 10th and Constitution Avenue, P.O. Box 370122, Washington, DC 20013-7012, USA. E-mail: [email protected]

New Developments on the Biology of Chrysomelidae edited by P. Jolivet, J.A. Santiago-Blay and M. Schmitt © 2004 SPB Academic Publishing bv, The Hague, The Netherlands 2 J.A. Santiago-Blay only feed on one plant species. Leaf-mining chrysomelids are Sagrinae larvae are borers into semi-rigid stems and, uni- or bivoltine, particularly in temperate zones, but they can when the larva pupates, it makes a cocoon within be trivoltine or multivoltine in the tropics. Natural biological the stem gall. Currently, only sagrine larvae of the enemies of the immature stages of leaf-mining chrysomelids are abundant but, in most cases, these organisms seem to be Old World genus Sagra and the New World genus unable to control outbreaks of leaf-mining chrysomelids. Leaf- Atalasis are known to make galls. However, except mining appears to have arisen independently multiple times in for anecdotal observations, nothing is known about the Chrysomelidae, typically from most recent common an- the larvae of Megamerus and other Gondwanian cestors which were exophytic. Leaf-mining mostly occurs in sagrines. Most of them are probably gallicolous, but the Hispinae, and it may have evolved from ancestors living there are likely to be exceptions in Australia. Me- between closely appressed leaves of monocotyledons. Several gamerus, for instance, is said to be gallicolous in leaf-mining chrysomelids, especially hispines, are economi- cally important as pests of major crops; others are used as Madagascar (Jolivet to Santiago-Blay, personal com- biological control agents of weeds. munication, July 2003). A leaf-mining organism is one in which, mini- mally, one of its life stages actively tunnels or mines An introduction to leaf-mining insects the usually flattened expansions of shoots, the leaves. Leaves of vascular plants have three parts: base, pe- What is leaf-mining? tiole, and blade (or lamina). The largest and most often flattened portions of leaves are usually known Leaf-mining is the consumption of foliar material as the blades, although graminologists (students of contained within the epidermal layers, without elic- grasses, Poaceae) also refer to entire leaves as iting a major histological response from the plant. ‘blades’. Leaf-miners overwhelmingly consume the The study of leaf-mines is termed ‘minology’ (He- relatively softer tissue, or parenchyma, contained ring, 1951). Mines may have a thin rim of callus between the epidermal layers, although some leaf- tissue (Hering, 1951). Leaf-miners consume devel- miners tend to consume vascular tissues (Hering, opmentally differentiated tissue and their tunneling 1951). activities do not elicit a major proliferation of un- Leaf-mining is largely restricted to the larvae of differentiated tissue (Connor & Taverner, 1997). about a dozen families of the four major holometab- Superficially similar-looking structures on leaves, olous insect orders: Coleoptera (Buprestidae and known as pseudomines, are caused by a variety of Chrysomelidae), Lepidoptera (Nepticulidae, Gra- microorganisms, including fungi and viruses; feed- cillariidae (sensu lato), Incurvariidae, Tisheriidae, ing activities of non-leaf-mining insects; or as a Coleophoridae, Eriocraniidae, and Opostegidae), reaction of plants to margin-feeding (Hering, 1951). Diptera (Agromyzidae, Anthomyiidae, and Tephri- Mining leaves is one of several modes that in- tidae), and Hymenoptera (Tenthridinidae). In nu- vertebrates have of feeding and living inside plants merous species of insects, including leaf-mining (endophyty). Other endophytic ways of living may chrysomelids, the pupa and to a lesser extent the elicit a response from the plant in the form of exu- adult, also dwell within the mine. The larvae of leaf- date production (Langenheim, 2003; Santiago-Blay miners produce a relatively long-lived behavioral et al., 2002) and/or gall formation. Numerous in- record of their recent, as well as ancient herbivory, sects bore or otherwise penetrate the shoot (trunk, hence providing a unique window into the nature branches, stems, buds, flowers, fruits, or seeds) or of ancient insect-host-plant associations and possible the roots, causing significant economic damage (Met- coevolution (Fossil evidence of leaf-mining insects, calf & Metcalf, 1993). below). Other insects and mites dwell inside leaves, elic- There are other chrysomelids with tunneling ten- iting a major histological response, forming enlarge- dencies (albeit, not in leaves). In Central America, ments of anomalous tissue, known as galls or cecidia Vencl et al. (2004) discovered that larvae of Neolema (singular, cecidium < Greek, kekis (κεκις), or gall; approximata Jacoby eat ovules and flower parts, and Ananthakrishan, 1984; Byers, 2002; Csóka et al., that larvae of Neolema sallaei Jacoby bore into the 1998; Darlington, 1968; Felt 1940; Labandeira & scandent stems of two species of Commelinaceae. Phillips, 1996a, 2002; Lindquist et al.,1996; Meyer, Also, one species of criocerine feeds on seeds of 1987; Shorthouse & Rohfritsch, 1992). Apparently Commelinaceae, and some species of Crioceris feed there is a report of mites as leaf-miners of forest cano- in fruits of Asparagus (Asparagaceae). Jolivet once pies in Queensland (Australia) but this find has not found a galerucine, Agetocera filiformis Laboissière, been followed-up with additional published reports. feeding on the fruits of the vine Cayratia japonica Mani (1964) indicates that “some species of … Chry- (Thunberg) Gagnepain (Vitidaceae) in Thailand. somelidae… have been reported to give rise to galls”, Other ecologically unusual feeding tendencies of nor- although no further details are given. The only known mally leaf-feeding chrysomelids have been observed, chrysomelid gall-makers are the Sagrinae and Ortho- such as members of the Lamprosomatinae feeding lema samalkotensis (Criocerinae) (Heinze 1943). on rodent excreta in the Florida Keys or galerucines Jolivet has seen many sagrine galls in Upper Volta feeding on snake wounds in Indonesia (Jolivet to in Africa; in Vietnam, he frequently saw adults simul- Santiago-Blay, personal communication, June 2003). taneously hatching at the start of the rainy season. Hering (1951) reported cases of stem-mining insects. Leaf-mining chrysomelids 3

In summary, “there are more things in heaven and 1, as their larvae feed temporarily on leaf buds and earth, Horatio, than are dreamt of in your philoso- the larger larval instars live on stems (e.g., Kalshoven, phy....” according to Hamlet. 1981; Maulik, 1938; Spaeth, 1933,1936; Uhmann 1942,1952,1963). Mariau (2001) indicates that both “To be or not to be”… a leaf-mining chrysomelid the larval and adult stages of Cryptonichini live Occasionally, it is difficult to precisely define what between the leaflets of unopened buds forming the a leaf-mining insect is. For instance, in the eumolpine spear of the leaf frond. Others species of the Cryp- tribe Spilopyrini (e.g., Stenomela, Hornius, Bohu- tonichini, such as species of Plesispa, seem to eat miljania, and probably Spilopyra), the first instar the adaxial and abaxial surfaces of the leaves, but larvae are diggers, eating their way into leaf buds mining often seems to be the case (Maulik, 1919; to feed and emerging from buds frequently (Jolivet Jolivet to Santiago-Blay, personal communication, to Santiago-Blay, personal communication, June June 2003). I have included all species of Plesispa 2003). As the larvae grow larger (approximately from for which I could find host-plant records in Table the second instar onward), they feed on the leaf 1. In contrast, species in the genus Callistola are surface (Jolivet to Santiago-Blay, personal commu- browsers, but there is one record (Cox, 1996; Cox nications, April, June 2003). I have excluded spilo- to Santiago-Blay, personal communications, May pyrines from Table 1, a compendium of the reported 2003) where a ‘Callistola sp.’ is recorded as a leaf- leaf-mining chrysomelids of the world. miner. This record is in error (Cox to Santiago-Blay, One of the most dramatic examples of behavioral personal communications, June 2003) and I have variability in leaf-mining is Clitea picta Baly (A1ti- excluded all species of Callistola from Table 1. cinae) larvae, which may bore into leaves from the In the hispine tribe Gonophorini, “larvae are, midrib, young shoots, spines and axils of branches, according to the species, either leaf-miners or free- and occasionally developing fruits (Zaka-ur-Rab, living larvae between the leaves” (Jolivet & Hawkes- 1991). Cases of organoxeny, or feeding in an un- wood, 1995). I have included as many gonophorines usual plant organ, are rare in leaf-mining Chryso- for which I could find some documentation on their melidae. Some species of Longitarsus (Alticinae) host-plant associations in Table 1. have larvae that “are root feeders, but sometimes The conceptual difficulties in defining leaf-min- [they have been] reported to be leaf-miners” (Jolivet ers are also present in some other groups of insects & Hawkeswood, 1995). (Hering, 1951). For example, in the cecidomyiids As a general rule, species in the hispine tribes (Diptera), blister galls described by cecidologists may Botryonopini, Anisoderini, Aproidini, Callispini, merge imperceptibly with leaf mines (Gagné, 1994). Leptispini, and Eurispini of the Old World Hispinae The difference evidently hinges on the degree of and in the Oediopalpini, Cephaloleiini, Hybosispini, production of cecidogenic nutritive callus or other Arescini, and Alurnini of the New World Hispinae, anomalous tissues before the structure can be termed are not leaf-miners. They have been omitted from a ‘gall’. Similarly, the “distinction between brows- Table 1. Even so, some exophytic taxa that typically ing and mining at times could be subtle” (Jolivet to are ‘strip miners’, have been reported (probably Santiago-Blay, personal communication, May 2003). erroneously) to be leaf-miners. For instance, Hering “[Leaf-scratching] larvae among Hispinae could be (1957) reports that “various species of the genera Arescus and Chelobasis were frequently found in also occasionally miners” (Jolivet to Santiago-Blay, communal mines on Heliconia, a genus of Heli- personal communication, June 2003). coniaceae. Maulik presumes that this communal living in a single mine must frequently give rise to Leaf-mining insects are only found in the four major hybridization. This seems all the more plausible since holometabolous orders leaf-mining Hispinae normally pupate within the mine and the emerging imagines, which in Coleoptera Within the Hexapoda, leaf-mining is apparently re- always remain for some time in the mine to tan and stricted to the four major holometabolous insect harden their exoskeleton necessary for chewing their orders: Coleoptera, Diptera, Lepidoptera, and Hy- way out, might thus easily find a suitable mate. If menoptera. It is estimated that 10 000 species of the emergence of different species took place in a insects are leaf-miners (Connor & Taverner, 1997). single mine simultaneously, it is very reasonable to All feed on vascular plants, including ferns, or pteri- assume that in all probability hybridization will dophytes (ferns and their allies), gymnosperms (co- ensue” (emphases added by JASB). However, Strong nifers, gnetaleans, ephedraceans, and Ginkgo), as well (1983), who has studied Chelobasis bicolor (Pic), as extinct seed-fern clades such as corytosperms makes no mention of leaf-mining on these hispines. (Rozefelds, 1988), and flowering plants (Labandeira, Neither Arescus nor Chelobasis are ‘leaf-miners’, 2002a). Species of leaf-miners are particularly abun- instead, they are ‘strip feeders’, eating on the sur- dant in numerous families of relatively basal Le- face and not between the upper and lower epider- pidoptera (Askew, 1980; Auerbach & Simberloff, mis of the leaves (Strong to Santiago-Blay, personal 1984; Connor & Taverner, 1997; Kristensen, 1999; communication, June 2003). Labandeira et al., 1994; Opler, 1973; Powell, 1980), Some members of the Old World hispine tribe where they also have left fossil evidence (Labandeira, Cryptonychini have also been omitted from Table 2002a,b). The thirteen lepidopteran superfamilies 4 J.A. Santiago-Blay

(and the 34 families parenthesized) reported to have In the Coleoptera, leaf-mining has been described leaf-mining species are, as follows: Heterobath- in at least five superfamilies, placed in nine fami- mioidea (Heterobathmiidae), Eriocranioidea (Erio- lies (parenthesized): Buprestoidea (Buprestidae, Tra- craniidae and Acanthopteroctetidae), Nepticuloidea chydinae); Bostrichoidea (Anobiidae); Cucujoidea (Nepticulidae and Opostegidae), Tischerioidea (Ti- (Nitidulidae, species of Xenostrongylus and of Anis- scheriidae), Palaephatoidea (Palaephatidae), Incur- ter); Tenebrionoidea (Mordellidae, Ptininae, Ptinus varioidea (Incurvariidae, Prodoxiidae, Adelidae, and antillanus Bellés); and the Chrysomeloidea [Chry- Heliozelidae), Tineoidea (Gracillariidae, Buccula- somelidae (details in this chapter), Platypodidae tricidae, Douglasiidae, and Roeslerstammidae), Gele- (Phylloplatypus pandini Kato), and Curculionidae chioidea (Oecophoridae, Elachistidae, Coleophoridae, sensu lato, species of Orchestes, Prionomerus, Rham- Momphidae, Cosmopterygidae, Scythrididae, and phus, Ciopus, and others; Belidae; and Attelabidae] Gelechiidae), Copromorphoidea (Carposinidae and (Connor & Tavener, 1997; Crowson, 1981; Hespen- Epermeniidae), Yponomeustoidea (Glyphipterigidae, heide, 1991; Hespenheide & Kim, 1992; Kato, 1998; Acrolepiidae, Argyrestiidae, Yponomeutidae, Helio- Lawrence, 1991; Lawrence & Britton, 1991; Lawson, dinidae, Ochsenheimerediidae, and Lyonetiidae, the 1991; Needham et al., 1928; Paulian, 1988; Philips latter often included with the Tineoidea), Tortricoidea et al., 1998: Wilcox, 1979). In the chrysomelids, or (Tortricidae), Pyraloidea (Pyralidae), and Pteropho- leaf beetles, leaf-mining has been reported for the roidea (Pterophoridae) (Connor & Taverner, 1997; subfamilies Zeugophorinae, Criocerinae, Alticinae- Heppner, 1998; Kristensen, 1999). Powell (1980) Galerucinae (Trichostomes sensu Jacoby 1908, or suggests that feeding preferences among ‘micro- Galerucinae, sensu lato), and Hispinae-Cassidinae lepidopterans’ have evolved following “specialized (Cryptostomes sensu Chapuis 1874-1875, or Cassi- larval feeding niches or horizons within communi- dinae, sensu lato Staines (2002a,b)) have leaf-min- ties… rather than along botanical evolutionary lines”. ing representatives. In Diptera, twelve higher taxa (and seventeen fam- The Coleoptera have more phytophagous species ilies, parenthesized) that have leaf-mining species than any other order of insects, yet a smaller propor- include the nematoceran Tipulomorpha (Tipulidae), tion of beetle groups have this modus vivendi than Culicomorpha (Ceratopogonidae and Chironomidae), do the Lepidoptera. Within the Chrysomelidae, only and Bibionomorpha (Cecidomyiidae and Sciaridae); the hispine chrysomelids are relatively well known. the brachyceran Empidoidea (Dolichopodidae), and This is partly attributable to the fact that some are the cyclorrhaphan Aschiza (Phoridae and Syrphidae), prominent agricultural pests. Why are there so few Calyptratae (Anthomyiidae and Scathophagidae), leaf-mining Coleoptera? Connor & Taverner (1997) Acalyptratae Diopsoidea (Psilidae), Tephritoidea suggest that lack of speciosity in leaf-mining insects (Tephritidae), Luaxanioidea (Luaxaniidae), Opomy- suggests the lack of adaptive radiation, implying that zoidea (Agromyzidae, notorious for having numer- the evolutionary benefits of being a leaf-mining ous pestiferous species), Carnoidea (Chloropidae), coleopteran have been outweighed by the drawbacks, as well as Ephydroidea (Drosophilidae and Ephy- such as an elevated incidence of parasitoidism. Ne- dridae) (Colless et al., 1991; Connor & Taverner, vertheless, there are numerous known leaf-mining 1997; Disney, 1994; Evenhuis, 1994; Labandeira, beetles that still need to be described (Hespenheide, 2003; Needham et al., 1928). These families are quite 1991). In fact, there are many more non-chrysomelid unrelated phylogenetically. Labandeira (2003) es- beetle leaf-mining taxa than hispines, at least in Cen- timates that there are at least 25 independent origi- tral America. The perception that there is a depau- nations of leaf-mining in the Diptera, when these perate nonhispine leaf-miner fauna is an artifact of cases are evaluated at the subfamilial/familial level the taxonomy of those other groups. For instance, or higher. Their mining is concentrated in herbaceous the current number of buprestids for the La Selva plants, instead of woody plants as leaf-mining insects Biological Station (Costa Rica) is 167 leaf-mining in the Coleoptera, Hymenoptera, and Lepidoptera species out of 218 total for the family (72%), more (Labandeira, 2003). Among aquatic nematocerans, than twice as many species as there are leaf-mining most host plants include the Poaceae and the Cypera- hispines. There are three major groups (Rhynchitidae, ceae, and less commonly herbaceous dicotyledoneous Prionomerinae, Tachygoninae) and one minor (Ca- plants (Hering, 1951; Labandeira, 2003). marotinae) group of weevil leaf-miners at La Selva. Leaf-mining is not as common in the other two At the moment, Hespenheide has separated 55 spe- holometabolous insect orders, namely the Hyme- cies of weevils. In the Tachygoninae, 21 of 26 spe- noptera and the Coleoptera. In the Hymenoptera, leaf- cies are undescribed; there are twelve leaf-mining mining has been reported in about 100 species of species of rhynchitids. The issue is that hispines are symphytans, including the families Argidae, Per- larger in size and relatively well-known taxonomi- gidae, and Tenthredinidae (Connor & Taverner, 1997; cally. The buprestids and weevils frequently are small Naumann et al., 1991; Smith, 1995). Most symphytan (often <3 mm) and very poorly known taxonomi- larvae are phytophagous and the Selandriinae (Ten- cally. For example, 113 of La Selva’s 167 buprestid thredinidae) are almost exclusively on ferns. The leaf-mining species are undescribed, and virtually Blasticotomidae (Hymenoptera) are petiole miners all the weevils are undescribed as well. There are of ferns (Needham et al., 1928). several very large genera of leaf-mining buprestids Leaf-mining chrysomelids 5 in the Old World, and they are relatively poorly 2002a,b; Labandeira et al., 1994; Lang, 1996; Ste- known. The question posed above cannot be an- venson, 1992; Wilf et al., 2001) and extending to swered at this moment due to the lack of sufficient the late Jurassic/Early Cretaceous boundary (;142 data (Hespenheide to Santiago-Blay, personal com- Ma) (Rozefelds, 1988), and the early Triassic to late munication, July 2003). Middle Triassic (230-225 Ma) of Kazakhstan, Aus- Why are there no non-holometabolous leaf-min- tralia, and South Africa (Anderson & Anderson, ing insects? Perhaps a combination of reasons may 1989; Labandeira, 2003, personal observations; Roze- help to explain this. Firstly, numerous hemipteroid felds & Sobbe, 1987; Zherikhin, 2002). Additional insects (heteropterous and homopterous Hemiptera examination of older fossils may reveal evidence for and their allies), many of which are phytophagous, an older origin for this modus vivendi. There is no have piercing-sucking mouthparts, and produce re- evidence in the fossil record of leaf-mining mites latively small holes to feed on cells’ protoplasts. (Labandeira to Santiago-Blay, personal communi- Undoubtedly, hemipteoid insects have mechanical cation, July 2003), although there are numerous difficulties creating holes large enough to maneu- documented fossil mites (Labandeira, et al., 1997; ver through the inside of the leaf blade. Interestingly, Petrunkevitch, 1955). just as many hispines, a number of species from There seems to be no reported evidence of leaf- several hemipteroid lineages live between appressed mining chrysomelid larvae in the fossil record (San- leaves, including taxa, such as the Termitaphididae tiago-Blay, 1994), but it is probable that mines with and Thaumastocoridae, both in the order Heteroptera patterns characteristic or suggestive of chrysomelids (Schuh & Slater, 1995), as well as numerous Coc- may have been overlooked or remain to be discov- coidea (Carver et al., 1991). And yet, they have not ered. Until recently, many paleobiologists would occupied the leaf-mine adaptive zone. Secondly, often disregard rocks containing fossil plants with many nonholometabolous (paleopteroid and ortho- evidence of damage in favor of material exhibiting pteroid) insects tend to have large appendiculate paleobotanically diagnosable and complete leaves immature stages that lack a vermiform facies, ren- (Allmon to Santiago-Blay, pers. comm., July 2003; dering leaf-mining physically difficult. Thirdly, it Labandeira to Santiago-Blay, personal communica- appears that the leaves or leaf-like structures of tion, July 2003). Recently, Wilf et al. (2000) (sum- aquatic plants, where many paleopteroid, orthopte- marized by Pennisi, 2000) described feeding marks roid, and hemipteroid insects live, are too thin for probably caused by rolled-leaf hispines (Cepha- a typical immature insect to inhabit. However, while loleiini or Arescini). These Cretaceous feeding marks, size could be a factor for the later instars, the ear- which were produced by strip feeders on mono- lier instars can be minute. Although hemimetabolous cotyledoneous plants, not by leaf-miners, are the groups evolved prior to the diversification of an- oldest known external feeding damage by chrysome- giosperms (Kukalová-Peck, 1991; Labandeira, 2002a; lids and they are represented by damage types from Labandeira & Sepkoski, 1993) and most likely were multiple life stages. There is material in the Dakota fixed on using food materials that were always avail- Formation (100 Ma) of Nebraska and Kansas which able, such as dead leaves of nonangiosperms, algae, has feeding marks that suspiciously resembles chry- or other animals, the targeted plant tissues by insects somelid damage (Labandeira to Santiago-Blay, per- that feed using the piercing-and-sucking method are sonal communication, July 2003). Also, there are a the same in gymnosperms or angiosperms (Laban- number of described Sceleonopla and other fossil deira & Phillips, 1996b). Hence, the lack of leaf- hispines in tribes known to have leaf-mining gen- mining hemipteroid insects is probably unrelated to era (Santiago-Blay, 1994; Santiago-Blay et al., 1996; gymno- or angiospermy (Labandeira to Santiago- Staines & Sanderson, 2000). The presence of leaves Blay, personal communication, July 2003). In addi- with characteristic feeding marks, particularly in the tion, other major insect orders containing leaf-mining company of a leaf-mining chrysomelid body-fossils, insects also originated prior to the diversification would be strongly suggestive evidence for the ex- of angiosperms (Kukalová-Peck, 1991; Labandeira istence of this behavior during the Cretaceous and 1997, 1998, 1999, 2002a; Labandeira & Sepkoski, perhaps the Jurassic periods. 1993) and their leaf-mines are often excellently Changes in the frequency of leaf-mining and other preserved in the Mesozoic fossil record (Anderson more host-specific types of damage have also been & Anderson, 1989; Rozenfelds, 1988; Zherikhin, considered as evidence for major and sudden changes 2002). in the patterns of insect herbivory at the Cretaceous- Tertiary extinction event (Labandeira et al., 2002a,b). Fossil evidence of leaf-mining insects Interestingly, Wilf et al. (2001) suggest that a pro- longed increase in leaf-mining from the relatively Despite the numerous biases of preservation (Cramp- humid and warm-temperate late Paleocene (;56 Ma) ton et al., 2003; Labandeira, 2002a), highly char- to the more subtropical early Eocene (;53 Ma) to acteristic mines preserved in fossil plants have been the relatively drier and subtropical middle Eocene reported for several orders of insects, including sev- (;43 Ma) may have been related to global paleo- eral families of Lepidoptera, Diptera, and for a num- climatic changes and concomitant vegetation shifts ber of unidentified insects as early as the Cretaceous, from what we now know as the central Rocky Moun- some 100 millions of years ago (Ma) (Labandeira, tain region of the USA. 6 J.A. Santiago-Blay

Introduction to leaf-mining chrysomelids sented by 78 genera which have been reported as leaf-miners, or 40% of the approximately 200 gen- Taxonomic distribution of leaf-mining chrysomelids era described. There are over 400 reported leaf- mining hispines, accounting for 14% of the over 3000 In this paper, I use the classification of Seeno & species described in the subfamily (Mariau, 2001). Wilcox (1982). Some of the drawbacks of Seeno & There is a single reported genus of leaf-mining in Wilcox (1982) are caused by the difficulties in some the Cassidinae, representing less than 1% of the 159 groups, particularly the natural delimitations, if any, genera described. Only six species of cassidines have between the Galerucinae + Alticinae (Trichostomes) been reported as leaf-miners, accounting less than and the Hispinae + Cassidinae (Cryptostomes), in 1% of the 2760 species names in the Cassidinae addition to the position of numerous other taxa, as (Borowiec, 1999). well as recent advances from more detailed studies In the Zeugophorinae, a subfamily with only a at all levels of biological resolution, from the mol- handful of described genera, all studied species of ecules to the ecosystem. Nevertheless, Seeno & Zeugophora have leaf-mining species. However, as Wilcox (1982) have served students of the Chryso- in most leaf-mining chrysomelids, “at present all that melidae well, and have presented the best compre- can be said is that the larvae of the Indian species hensive system. Until the higher classification of the of Zeugophorinae are presumably leaf-miners and Chrysomelidae becomes more stable and generally their host-plants have not been recorded.” (Verma accepted as a practical tool of communication that to Santiago-Blay, personal communication, July reflects some presumed past events, I find it useful 2003). A total of twelve species are reported to be to follow Seeno & Wilcox’s (1982) ‘catalogue clas- leaf-mining zeugophorines (Table 1). sification’ (as it is sometimes referred to) with the With regard to leaf-mining, the Criocerinae rep- appropriate and occasional caveats. resents a mysterious and fascinating group. With only Leaf-mining is relatively uncommon in the Chry- two species being reported as leaf-mining criocerines somelidae and is found in 103 reported genera, re- (Table 1), many more are expected to be documented, presenting 4% of the approximately 2600 genera particularly among minute forms that inhabit the described and amounting to over 500 reported spe- tropics (see Vencl et al., 2004, for examples of tropi- cies, or 1-2% out of the 40-50,000 species described cal endophytic criocerines). (Lopatin, 1984; Jolivet to Santiago-Blay, personal Within the Trichostomes sensu Jacoby, or Galeru- communication, March 2003). Larvae in the follow- cinae + Alticinae, the Galerucinae has two genera ing subfamilies are known leaf-miners, with num- that are known to have leaf-mining species: one bers and percentages of taxa included, as well. The species of Galerucella and all the known species of subfamily Zeugophorinae consists of one genus with the genus Monoxia. Blake (1939) and other research- reported leaf-mining species, or 25% of the total ers noted the need to understand the difficult gale- genera; there are twelve reported leaf-mining spe- rucine genus Monoxia, where most of the species cies, about 17-20% of the species described in the studied by me are leaf-miners. A total of 20 species subfamily and about 60-70 species described in the are reported as leaf-miners (Table 1). In contrast, Zeugophorinae, all believed to be leaf-miners (Jolivet in the Alticinae there are 19 leaf-mining genera to Santiago-Blay, Schmitt to Santiago-Blay, and (Aphtona, Apteropeda, Argopistes, Argopus, Chae- Verma to Santiago-Blay independent personal com- tocnema, Clitea, Dibolia, Epitrix, Febra, Hippuri- munications, May through July 2003). The subfamily phila, Longitarsus, Mantura, Mniophila, Ochrosis, Criocerinae comprises two genera, representing 10% Phyllotreta, Psylliodes, Schenklingia, Sphaeroderma, of the described genera, and two reported leaf-min- ing species accounting for less than 1% of the approx- and Throscoryssa) that are known to have some or imately 1450 species described in the Criocerinae all of their species as leaf-miners. A total of 65 (Jolivet to Santiago-Blay, Schmitt to Santiago-Blay, species are reported to be leaf-mining alticines (Table and Vencl to Santiago-Blay, independent personal 1). These genera belong in several seemingly un- communcations, May 2003). The Galerucinae has related groups of the Alticinae (Seeno & Wilcox, two reported leaf-mining genera, representing less 1982). than 1% of the approximately 500 genera described. Within the Cryptostomes sensu Chapuis, the His- There are approximately 20 reported species of leaf- pinae have 78 known or suspected leaf-mining gen- mining galerucines, accounting for less than 1% of era (Acanthodes, Acentroptera, Achymenus, Agonita, the approximately 7000 species described in the Anisostena, Asamangulia, Aspidispa, Baliosus, subfamily (Jolivet to Santiago-Blay, personal com- Balyana, Brachycoryna, Carinispa, Cassidispa, Cha- muncation, July 2003). The Alticinae has nineteen eridiona, Chalepus, Charistena, Chrysispa, Clino- reported leaf-mining genera, representing about 3% carispa, Cnestispa, Coelaenomenodera, Corynispa, of the approximately 500 general described. There Crapedonispa, Cyperhispa, Dactylispa, Dicladispa, are 65 reported species of leaf-mining flea-beetles, Dorcathispa, Downesia, Enischnispa, Euprionota, or about 1-2% of the 4000-8000 species described Freycinetispa, Gestronella, Glyphuroplata, Gono- in the Alticinae (Schmitt to Santiago-Blay, personal phora, Heptispa, Heterispa, Hispa, Hispellinus, His- communication, May 2003). The Hispinae is repre- poleptis, Isopedhispa, Javeta, Klitispa, Metaxycera, Leaf-mining chrysomelids 7

Micrispa, Microrhopala, Nonispa, Ocnosispa, Octhis- from many other papers (see References below), and pa, Octotoma, Octouroplata, Odontota, Oncocephala, unpublished data, have been used in Table 1. Oxychalepus, Oxyroplata, Pentispa, Pharangispa, Phidodonta, Physocoryna, Pistosia, Platochispa, Platypria, Plesispa (but see remarks in “To be or Life history of leaf-mining chrysomelids not to be … a leaf-mining chrysomelid” above), Polyconia, Prionispa, Probaenia, Promecotheca, The biology of several leaf-mining chrysomelids has Prosopodonta, Rhabdotohispa, Rhadinosa, Sceleo- been reported in detail. While there is considerable nopla, Spilispa, Stenopodius, Stenostena, Stehispa, variability in life history, some typical patterns and Sumitrosis, Temnochalepus, Trichispa, Uroplata, variations are described below, elaborated in Fig- Xenochalepus, and Wallacispa). A total of 410 spe- ures 1-5, and summarized in Figure 7. cies are reported to be leaf-miners (Table 1). In contrast, the Cassidinae has only one leaf- Egg mining genus, Notosacantha, which also has been placed in the Hispinae (Borowiec, 1995; Medvedev The fine ultrastructure of leaf-mining chrysomelid & Eroshkina, 1988). A total of six species are re- eggs appears to have been all but overlooked. Mem- ported to be leaf-mining cassidines (Table 1). mott et al. (1993) reported that egg structure is useful Considerably more leaf-mining chrysomelids will in distinguishing three sympatric, congeneric spe- be revealed. For example, Frost (1931) and Hespen- cies of Chalepus (Hispinae) that share three species heide (1991) reported rearing many species of leaf- of Central American Lasiacis bamboos (Poaceae) mining hispines from plants at Barro Colorado Island as host-plants. For a comprehensive discussion of (Panama) and La Selva (Costa Rica), as well as insect eggs, see Hinton (1981). hispines from several species of Cecropia (Cecro- piaceae) in central Panama. As researchers conduct Timing and location of oviposition careful field and laboratory observations and the life Like typical insects, after emerging from diapause histories of other chrysomelids are studied, particu- (if present), leaf-mining chrysomelid beetles feed and larly those in taxa previously documented as leaf- mate. Shortly after copulation, depending on the miners (e.g. the hispine Octotoma, c.f. Staines, 1989), species, eggs are laid on different locations of the more chrysomelid leaf-mining taxa will be docu- leaf, either abaxially, or adaxially, or in a cavity made mented, particularly within the hispines. I have not by the female on the blade (Chittenden, 1902; Monrós located reports of leaf-mining for the other subfami- & Viana, 1947; Sen & Chakravorty, 1970). The act lies of the Chrysomelidae, although this life habit of oviposition by several leaf-mining chrysomelids is suspected for the Orsodacninae and the Aulaco- has been observed, and it is relatively simple. Leaf- scelinae (Jolivet to Santiago-Blay, personal commu- mining chrysomelids tend to lay relatively few eggs nications, 2003). per oviposition bout on or in host-plant leaves (Chit- tenden, 1902; Ford & Cavey, 1985). For the hispine How many species of leaf-mining chrysomelids are/ Dicladispa armigera (Olivier), oviposition follows were there? shortly (less than three to four days) after copula- tion and most eggs are deposited during the first week I estimate that there are approximately 2500 spe- of adult life (Sen & Chakravorty, 1970). cies of leaf mining chrysomelids, principally located Several hispines, including Baliosus nervosus in the hispines. However, in addition to the intrin- (Panzer), Dicladispa armigera (Olivier), Promeco- sic difficulties in estimating “how many species?” theca couruleipennis Blanchard, and P. cumingi Baly (May 1990), it is also difficult to know how many lay eggs singly, in a space excavated by the female leaf-mining species there are among the Chrysome- on the upper portion of leaves or leaflets (Teixeira lidae because the larval stages, larval-adult associa- et al., 1999; West & Lothian, 1948). The hispines tions (e.g., Lee & Furth, 2000), and basic biological Microrhopala vittata (Fabricius), M. xerene (New- research on most species, particularly those with root- man), and Odontota dorsalis oviposit abaxially in feeding larvae, still remains to be done for over an small masses consisting of three to five eggs per estimated 95% of the species. In other cases, the cluster or for each of several rows (Clark, 1983; literature is not clear as to whether a species is a Needham et al., 1928). In some species, such as leaf-miner and, if so, what the host-plants of the Heterispa costipennis Boheman, females may lay as larvae and of the adults are. The works of Buhr little as one egg per leaf or several eggs, dispersed (1955,1956), Ford & Cavey (1985), Frost (1924), or clustered, per leaf (Monrós & Viana, 1947). Some Jolivet (1989a), Jolivet & Hawkeswood (1995), Mau- species of Sceloenopla (Hispinae) lay eggs, eight per lik (1931,1932,1933a,b), Needham et al. (1928), oviposition bout, inside an ootheca deposited on the Hering (1951,1957), Staines (2002b), Staines & mesophyll of the abaxial surface of young Cecro- Staines (1989,1992), and Wade (1935), among nu- pia spp. leaves (Andrade, 1984; Jolivet, 1989b). merous others, contain discussions, lists, and/or bi- Females may cover the excavations with an exudate bliographies of leaf-mining insects from different made out of chewed leaflet bits and feces (Boldt & regions of the world. Their data, as well as those Staines, 1993; Dharmadhikari et al., 1977; Zaka- 8 J.A. Santiago-Blay

Figs. 1 and 2. Eruciform larva of Monoxia spp. 1. Dorsal. 2. Ventral. Fig. 3. Sacculate or blotch mines caused by Monoxia guttulata (LeConte) on Artemisia douglasiana Besser (Asteraceae). Figs. 4 and 5. Different stages of pupation in Monoxia spp. in larval mine in a G. humilis. In Figure 5, note the presence of larval exuvium (to the left) still partially attached to pupae. Fig. 6. Side and top views of clip cages used by the author in host-plant feeding preference experiments by Monoxia beetles. ur-Rab, 1991; Fig. 7). In contrast, the hispine Odon- onstrated that this material reduces egg mortality tota dorsalis (Thunberg) lays its eggs in small groups from predators and parasitoids significantly, particu- on the lower surface of the leaves, although it also larly that of eggs located at upper tiers in M. vittata covers them with a “sticky substance partially cov- egg clusters (Damman & Cappuccino, 1991). In the ered with an excrementitious secretion” (Chittenden, case of the hispine Odontota dorsalis, it is believed 1902). According to Chittenden (1902), the secre- that the feces, with which eggs are covered, make tion “appears to possess some caustic properties, for eggs more cryptic (Wheeler, 1987). This material the place of an egg mass can always be seen on the also glues eggs to the leaf (Taylor, 1937). Never- upper side of the leaf as a small brown spot. It [the theless, natural enemies of eggs from non-leaf-mining secretion] hardens very rapidly, and becomes so chrysomelids may also be attracted by chemicals tough and firmly adherent to the eggs that these released by the eggs (Müller & Hilker, 2004). How- cannot be taken out from a mass without destroy- ever, the dark material with which female hispines ing them.” Octotoma scabripennis Guérin-Méneville and Uro- Fecal matter is used by numerous chrysomelids plata girardi Pic cover their eggs is liquid from the to cover their eggs (Müller & Hilker, 2004). For colleterial glands, not feces (Harley, 1969; Cilliers, instance, Clark (1983) suggests that Microrhopala 1987a). (Hispinae) anal secretions, likely feces, which soon darken and harden, cover and probably protect the Total fecundity and some correlates of oviposition eggs (Cappucino, 1991b; Damman & Cappuccino, The hispine Trichispa sericea Guérin-Méneville can 1991; Hodson, 1942). Field experiments have dem- lay up to 747 eggs in a season (Ravelojaona, 1970). Leaf-mining chrysomelids 9

Fig. 7. Typical life history of leaf–mining chrysomelids, with some variations (usually represented by dashed line).

The total number of eggs a female can oviposit during Larva her life can reach 100 or more for a D. armigera female, although most larvae die during hatching (Sen Using head capsule measurements from preserved & Chakravorty, 1970). The hispine, Promecotheca samples, Boldt & Staines (1993) found that Pentispa reichei Baly lays 16 to 25 eggs during her life suturalis has three larval instars. Other species, such (Hinton, 1981). These numbers fall well within as Dicladispa armigera (Hispinae), have four instars the range for the Chrysomelidae, 10 to 2800 ovipo- (Sen & Chakravorty, 1970). Samples from exuviae sited eggs during the life of a single female (Hinton, can be difficult to measure since the head capsules 1981). tend to separate along their epicranial sutures. How- In some species, the eggs are partially forced into ever, field observations are often carried out in situ the leaves’ tissues, through the tissue that has al- by placing the mined larvae against a light source. ready been partially eaten by the adult female (Hod- son, 1942). The hispine, Asanmagulia cuspidata Hatching and penetrating the leaf Maulik, oviposits just under the leaf-blade epidermis Larvae are the pre-eminently active mining stage of of sugar cane, Saccharum officinale Linné (Zakaur- leaf-mining insects. Newly emerged larvae may begin Rab, 1991). The hispine Platypria coronata Gué- mining a leaf on the upper (adaxial) or on the lower rin-Méneville can lay one to three eggs, all located (abaxial) epidermis, depending on the location of just under the lower epidermis. the egg from which the larvae ecclose. Newly hatched The number of eggs laid by Platrypia coronata Dibolia borealis Chevrolat (Alticinae) larvae enter (Guérin-Méneville) apparently varies, depending on leaves by making a slit on the epidermis and chew- whether it has fed recently on the leaf that is used ing their way in to inner tissues (Needham et al., as the site of oviposition on its host-plant, the le- 1928). Throscoryssa citri Maulik (Alticinae) larvae gume Pueraria phaseoloides (Roxburgh) Bentham mine leaves, entering through the lower surface of (Bernon & Graves, 1979). West & Lothian (1948) the blade and exit leaves through an exit hole on suggest that there must be intraspecific food com- the upper surface at night (Zaka-ur-Rab, 1991). Both petition in Baliosus nervosus hispines feeding on Octotoma scabripennis and Uroplata girardi (His- Tilia americana leaves. The hispine Pentispa su- pinae) penetrate the leaf promptly after hatching turalis (Baly) lays one to six eggs, most of them (Cilliers, 1987a). However, when larvae of the his- apically and adaxially on Baccharis bigelovii leaves. pines Promecotheca caeruleipennis and species of Small B. bigelovii leaves tend to receive only one the hispine genus Microrhopala hatch, they penetrate P. suturalis egg, perhaps a mechanism that reduces first through the maternally-laid exudate layer that competition for food (Boldt & Staines, 1993; Hespen- surrounds the egg and is glued to the leaf, and then heide, 1991; Teixeira et al., 1999). The chemical enter the leaf. In leaf-mining chrysomelids, such as determinants of oviposition in leaf-mining chry- the hispine Baliosus nervosus, whose eggs have been somelids appear to be unknown. deposited in an excavation made by the mother, the larvae invade the mesophyll-rich parenchyma upon hatching. Newly hatched B. nervosus larvae that were experimentally placed on an unchewed surface of 10 J.A. Santiago-Blay

Tilia americana Linné host leaves were unable to and A. biplagiatus Motschulsky occupy different penetrate the epidermis (West & Lothian, 1948). In niches of their oleacean host-plants. These species all cases, penetration of the epidermis opens a di- start mining on different surfaces of their host-plant rect avenue for the start of the leaf-mining habit. In leaves: on the upper surface for A. coccinelliformis contrast to some lepidopteran leaf-miners, all known and on the lower surface for A. biplagiatus. Occa- leaf-mining chrysomelid larvae are devoid of a case sionally, more frequently for A. coccinelliformis than (Hering, 1951). for A. biplagiatus, larvae change mines, perhaps as a result of fecal accumulation. Later, older larvae Leaf occupancy of Argopistes coccinelliformis may re-enter the host The number of mines in a leaf vary with the size leaves from the lower surface and Argopiostes bi- and the temporal availability of the leaf (Williams, plagiatus from the upper surface (Inoue, 1990b, 1989b). Microrhopala xerene (Newman) has many 1996). Similar pattern preferences have been noted more mines per leaf on the earlier germinating and for Octotoma scabripennis Guérin-Méneville and larger leaved Aster puniceus Linné (Asteraceae) than Uroplata girardi Pic (Harley, 1969). The anatomi- on the sympatric, later germinating and smaller cal and chemical determinants of this larval feed- leaved A. simplex (Willdenow). ing behavior are apparently unknown, although it The number of individuals per mine, and the func- is generally believed that larvae of leaf-mining in- tional significance of the mines, vary per species. sects tend to avoid the leaf veins (Hering, 1951). In some cases, such as that of Microrhopala xerene, the mines are large enough to contain up to four Morphological modifications immature members of the species (McCauley, 1938), In the context of evolutionary biology, adaptations while in some other species, mines may contain up are traits that are presumed to have been brought to two or three conspecifics (Harley, 1969; Lee, 1990; forth by the action of natural selection (Rose & Wheeler, 1980). In Odontota scabripennis and Uro- Lauder, 1996). Although anecdotal natural history plata girardi, the number of individuals in a mine cases certainly can be compelling and often form is inversely proportional to the size of the larval the basis for more detailed analyses, evidence for tenants (Harley, 1969). adaptation is best attained by experimental and com- The larvae of some species of leaf-mining chry- parative biomechanical studies coupled with a sound somelids stay in the same leaf during their entire phylogenetic analysis (Orzack & Sober, 2001; Ross life (Williams, 1989a); the larvae of other species & Lauder, 1996). In this strict sense, there are no change leaves, sometimes up to three times (Hering, traits among the leaf-mining chrysomelids that have 1951; Wheeler, 1987), generating a new mine for been experimentally shown to be specifically adap- each switch. For instance, several larvae of Zeugo- tive or linked with an adaptive radiation event. How- phora scutellaris Suffrian (Zeugophorinae) may find themselves living in a single chamber when several ever, many traits are compelling candidates, including independent mines become confluent. Yet, when a flattened body and spines on some adults as ad- mining is completed only one living larva is found. aptations for leaf-mining (Hering, 1951). Needham et al. (1928) do not explain whether this Leaf-mining chrysomelid larvae have a broad is caused by cannibalism, lack of sufficient food, spectrum of larval types from fully-legged eruciform or other reasons. Upon hatching, larvae of Odontota to very flat, disc-shaped, or onisciform shapes. In dorsalis penetrate one leaflet of Robinia pseudo- some genera, such as Chalepus (Hispinae) and Phyl- acacia (Leguminosae). The first hatched larvae or lotreta (Alticinae), the legs are well developed; in “leader” and the other larvae from the same clutch, the hispine Baliosus they are minute, whereas in the or ‘followers’, occupy the same mine, and they bur- zeugophorine Zeugophora and the hispine Octotoma, row communally during their first instar (Chittenden, larvae are apodous. Nevertheless, leaf-mining chry- 1902). Several days later, the larvae exit the leaves somelid larvae seem to be placed in two gross mor- and disperse, each larva mining a different leaflet. phological categories: first, the less modified or Interestingly, on occasion, two more mature Odontota eruciform type found in the Galerucinae and some horni Smith larvae have been found in the same mine Alticinae; second, the flattened type occurring among (Buntin & Pedigo, 1982). The solitary larvae of Zeugophorinae, many Alticinae, Cassidinae, and the Pentispa suturalis (Hispinae) usually mine two or Hispinae. Morphological data for leaf-mining lar- three Baccharis bigelovii leaves during their life val criocerines are not yet available. (Boldt & Staines, 1993). Platypria andrewesi The first type, the slightly or unmodified eruciform Weise (Hispinae) larvae consume foliar tissue briefly larvae, is represented by galerucines such as spe- out of several small pocket-like mines (Zaka-ur- cies of Galerucella spp. and Monoxia spp. and by Rab, 1991). In some leaf-mining insects, larvae some alticines, including Aphthona and Phyllotreta may change host-plant species during development (Böving, 1927, 1929; Grandi, 1959; Lawson, 1991). (Hering, 1951). Both galerucine and alticine larvae, which appear Intraleaf feeding preferences have been docu- only minimally modified for mining leaves (Needham mented in some species of leaf-mining chrysomelids. et al., 1928; Lawson, 1991) tend to be orthosomatic, Among a substantial number of differences (Inoue, have nine to ten abdominal segments in galerucines 1996), larvae of Argopistes coccinelliformis Csiki and ten in alticines, bear legs, display four to five Leaf-mining chrysomelids 11 palmate teeth on mandibles, and posses small, one & Viana, 1947). A flattened body may constitute an to two segmented antennae. Some hispine larvae, advantage in gas exchange. such as species of Microrhopala, have subcylindrical Hispine larvae tend to be flattened as well, al- bodies that taper posteriorly, an occiput with two beit, as a group they show the greatest range of body posteriorly projecting lobes, a fused clypeus and variation within leaf-mining chrysomelids. Hispine frons, a mandible projecting slightly over the labrum, larvae lack the forked projection in the eighth ab- absent labial palpi, postgenae that are lateral to the dominal segment present in cassidines. However, gula, antennae displaced anterodorsally, some mouth- because of its peculiar combination of traits – min- parts that are adnate and hyperhirsute, and an ex- ing larvae that lack posterior abdominal projections panded maxillolabial complex and ligula; labial palps and overall tortoise-shape body-like cassidines – that are involuted, a thorax that is only lightly scle- Notosacantha has been variously placed in the His- rotized, a constricted mesothoracic segment, the pinae or in the Cassidinae (Borowiec, 1995; Med- presence of legs, and ambulacral projections occur- vedev & Eroshkina, 1988; Rane et al., 2000; Staines, ring on the abdomen (McCauley, 1938). 2002b). Hispine larvae are approximately 5-10 mm In addition to the overall body shape, there are long, pale-colored, except for the dark eighth ab- many other modifications that leaf-mining chry- dominal segment, which tends to be darker; ortho- somelid larvae may have. When present, such modi- somatic; have four or six stemmata; and all eighth fications tend to be concentrated on the head. This abdominal segments frequently bear lateral projec- is exemplified by the second type of overall body tions (Lawson, 1991; Jolivet, 1989a). shape in larvae of leaf-mining chrysomelids, the As body forms become increasingly flattened, flattened, or depressed, type (Grandi, 1959), which particularly for the hispines, lateral processes become has two variants. broader and interlacing, making the larvae onisci- This is the first variant of the second type of leaf- form. This condition gives them appearance of water mining chrysomelid larvae and is represented by pennies (Coleoptera: Psephenidae) (Anderson et al., Dibolia femoralis Redtenbacher and Sphaeroderma 2002; Maulik, 1932; Jolivet, 1989a). However, ex- rubidum Gruells (Alticinae), as well as by some treme body flattening, is not exclusive to hispines hispine leaf-miners, such as Sceloenopla near bidens that mine leaves. Larvae of the leaf-browsing Pla- (Fabricius) which have conspicuously depressed tyauchenia latreille Castelnau, as well as species of bodies (Costa et al., 1988). In addition, these larval Arescus and Chelobasis, are extremely flattened, types have reduced body setation, legs, vestigial approximating 1 mm in thickness (Maulik, 1931, ambulatory lobes, a head capsule with prognathous 1933a). Presumably this is a restriction on the range or slightly declined mouthparts, an occipital area with of allowable body forms imposed on organisms living posteriorly expanded projections, involuted labial amid the crevices between appressed leaves over palps, and antennae that are displaced anterodorsally. evolutionary time (Maulik, 1931). Phyllotreta nemorum Linné and Psylliodes chryso- The heads of hispine larvae have a full spectrum cephala Linné (Alticinae), which also belong in this of variation, ranging from subglobular (as in spe- type, have a sclerotized plate on the last visible cies of Prosopodonta) to progressively flattened abdominal tergite (Grison et al., 1963). The hispine heads that have large posterior elongations on the Odontota dorsalis also is slightly modified but only epicranium. As flattening progresses, so does the minimally depressed dorsoventrally (Needham at el., prognathation and reduction of mouthparts and anten- 1928). nae (Maulik, 1931). Larvae of P. latreillei exhibit The second variant of the flattened larval type is a great reduction in mouthparts (Maulik, 1933a). The represented by species of the zeugophorine, Zeugo- prothorax enlarges correspondingly and tends to be phora. These larvae are apodous and have: a body more extensively connected to the head, at times by that is obviously flattened and enlarged anteriorly; heavily sclerotized tergites. Also, there is a reduc- a flattened head capsule with a prognathous or slight- tion in the meso- and metathorax, as well as greater ly declivous head that is somewhat retracted in the differentiation between the thoracic segments, which prothorax; two thorax-penetrating posterodorsolateral tend to be larger, legged, and devoid of spiracles, projections; three-segmented antennae; labrum with except for a pair between the pro- and mesothorax. long, spatulated bristles; long, depressed mandibles; The abdominal segments are generally smaller, apo- three-segmented maxillae having large, sclerotized dous, and bear spiracles. In general, leaf-mining stipites and cardines located beyond the labium; hispine larvae have reduced, fleshy legs, such as S. involuted labium with large, spatulated bristles and maculata Andrade or are apodous in the case of contiguous, reduced, or vestigial labial palpi; no species of Octotoma (Crowson, 1955; Steinhausen, coronal suture; frontal sutures in direct contact with 1966; Peterson, 1979). occipital foramen; thorax with two large apodemata; The fused, or connate, sclerotized eighth and ninth thoracic and abdominal ambulatory lobes, and a abdominal segments are common in many genera nine-segmented abdomen (Lawson, 1991; Lee, 1990; of hispine larvae, such as species of Prosopodonta Needham et al., 1928). The long thoracic setae in spp., as are the projections on the last visible body Zeugophora abnormis LeConte larvae may assist segment (Maulik, 1933b). Maulik (1931) believed larvae moving inside the mines (Frost, 1924; Monrós that the hardened eighth/ninth abdominal segment 12 J.A. Santiago-Blay may serve to further enlarge mines, since these lar- and 2 cm wide (Mariau, 1988). Gressitt (1957) re- vae are large, in order to allow for the accumula- ports that mines of several species of Promecotheca tion of detrimental feces and other frass. “By moving exceed 30 cm in length, and that a compound mine [this segment] the insect is able to make a clearing, of P. violaceae Uhmann produced by two or more or it [the frass] may thrust it outside if need be” larvae was 60 cm long. (Maulik, 1931). Several hispine larvae, such as spe- If the leaves of the host plant are elongated, mines cies of Dactylispa, have the terminal spiracles lo- may also be elongated, constrained by the veins cated on the abdomen, which are protected by the (Hespenheide & Dang, 1999). Several tropical his- distal projections, or papillae, inside the spiracular pines form serpentine mines, which are more char- opening, or the opening positioned at a very oblique acteristic of lepidopteran leaf-miners (Hespenheide angle to the longitudinal axis of the spiracular trunk & Dang, 1999). Another extreme case is the hispine (Maulik, 1932,1933b). Assamangulia cuspidata, which may produce lon- gitudinal mines of up to 20 cm long (Zaka-ur-Rab, Patterns of mines in leaf-mining chrysomelids 1991), and mines of the hispine Promecotheca papu- The dwellings made by leaf-mining larvae inside ana Csiki have measured up to 40 cm in length leaves are known as mines. Mines have several basic (Howard et al., 2001). Platypria coronata larvae mine morphologies and biological functions, such as waste leaves, starting at the site of ecclosion, avoiding large disposal and development, and those functions are veins, and forming circular mines (Bernon & Graves, often compartmentalized (e.g., the alticine Dibolia). 1979). Larvae of Pentispa suturalis (Hispinae) mine Extensive discussion of these subjects for leaf-mining small leaves and their mines tend to conform to the insects can be found in Hering (1951). shape of the leaf (Boldt & Staines, 1993). A few leaf-mining chrysomelids, such as Dibolia (Alti- Types of mines. There are many types of feeding cinae), make long serpentine mines; Hippuriphila damage done to plants, including leaf-mines (Har- and Phyllotreta (both Alticinae) make short serpen- grove, 1986; Hering, 1951; Labandeira, 2002a; Wilf tine mines (Frost 1924). et al., 2001). There is often a wealth of gross mor- Some of the most complex leaf-mines of tropi- phological patterns in the mines of leaf-mining in- cal hispines have been called ‘blotch lobulate’ (Hes- sects, and chrysomelids are no exception. A useful penheide & Dang, 1999) and ressemble the depection pictorial depiction of common, extant leaf-mine of digitate, or star mines, or asteronomes, in Hering’s damage with examples from each of the four orders (1951) classification. These mines have a central of leaf-mining insects is available online (http:// chamber, located under the oviposition place, coated www.leafmines.co.uk/index.htm). As a group, the with larval fecal material. Several compartments, or mines produced by leaf-mining chrysomelids vary lobes, are produced as the larva feeds at different in shape and dimensions, although most are of the sites. In some cases, resting and pupation may take blotch type, also known as blister type (see illus- place in lobes apparently used only for those activities tration in Wheeler, 1987). (Hespenheide & Dang, 1999). Mines of many chrysomelids are elongated and Communally-feeding larvae may change the shape serpentine, such as those of Phyllotreta nemorum of serpentine mines into those of blotch mines (Fig. (Alticinae) and of many hispines, including Octotoma 8), as occurs with the genera Heterispa and Oxyro- plicatula (Fabricius) (Needham et al., 1928). In other plata (Hespenheide & Dang, 1999). I am unaware cases, such as the galerucine Monoxia guttulata of blotched mines with compartments being trans- (LeConte) (Fig. 3) and species of Microrhopala formed into a simpler blotch mine or a blotch mines (Hispinae), their mines are more sacculate or blotchy converted to serpentine or irregular mines. While (Needham et al., 1928). In Octotoma scabripennis there can be functional compartmentalization in ser- and Uroplata girardi, mines are broader and com- pentine and in irregular mines, when there are dis- partmentalized, contain several separated feeding tinct compartments, they tend to be more frequent galleries. in blotch mines. Despite this variation, the shape of the blotch Patterns of mines in leaf-mining chrysomelids: mines produced by leaf-mining chrysomelids may sources of intraspecific variation. The shape of the be characteristic within a taxonomic group (Hodson, mines varies with developmental stage and age of 1942). For example, the mines produced by astera- the larvae. In Zeugophora annulata and in species cean-feeding larvae of Monoxia (Figs. 1 and 2) are of the fern-feeding alticine Schenklingia, mines of consistently simple, often inflated and sacculate, and early larvae are broadly linear; those of late larvae about 1 cm long at completion (Fig. 3). These kinds are wider (Lee, 1990; Kato, 1991). Kogan & Kogan of mines have also been reported for the hispine (1979) depict a wide range of variation in mine size, Pentispa suturalis (Boldt & Staines, 1993). as well as leaf area covered, on the hispine Odontota horni. In O. horni, some of the mines can be circu- Patterns of mines in leaf-mining chrysomelids: com- lar. The size of the mine is monotonally proportional partmentalization of functions. Larvae of the hispine to the size of the larvae. In hispines, for instance, Dicladispa testacea (Linné) make two mines, one mines produced in palm leaves can reach 20 cm long in which they spend the first larval stages, the sec- Leaf-mining chrysomelids 13

Fig. 8. Developmental transitions between types of leaf-mines; including: irregular, serpentine, blotched, or blotched with func- tional compartments. I am unaware of blotched mines with compartments being transformed into a simpler blotch mine or a blotch mines turning into serpentine or irregular mines, hence the absence of double arrows connecting those types of mines. While there can be compartmentalization in serpentine and irregular mines, distinct compartments tend to be more frequent on mines that resemble blotch mines. This figure is not a phylogenetic hypothesis and the arrows do not imply evolutionary transformations. ond in which metamorphosis is completed. The dis- somelidae known to contain leaf-mining taxa. I have position of the larval feces varies from species to included only traits that are relatively easy to ob- species and serves as a good example of mine com- serve and are less variable. Readers interested in partmentalization. In Zeugophora annulata (Baly), additional details should consult Cox (1996). Some feces are scattered within the mine (Lee, 1990). data for the Galerucinae originate from my studies Larvae of the hispine Platypria coronata and of the of Monoxia (Figs. 4 and 5). alticine Throscoryssa citri place their feces towards Dactylispa javaensis Maulik pupae have a spi- the center of the mine (Bernon & Graves, 1979; Zaka- racular presumably protective device on the fifth ur-Rab, 1991). However, P. caeruleipennis and P. abdominal segment. Some leaf-mining chrysomelid cumingi larvae defecate on one side of the mine and pupae bear strong, pointy prongs. The impact of leave the central space relatively free (Dharmadhikari pupae on mine enlargement, if any, is probably acci- et al., 1977; Zaka-ur-Rab, 1991). It is unclear whether dental. one side of the mine (right or left) is preferred, or whether such a preference is constant from individual Abbreviated diagnoses of the pupae of leaf-mining to individual or through time. Larvae of Dibolia spp. chrysomelids and of Baliosus nervosus defecate in the mine’s side Pupae of the Zeugophorinae are exarate, whitish, branches rather than on the central space of the mine have a setose head, possess four papillae on each (Hering, 1951; West & Lothian, 1948). Larvae of antennal segment, lack pronotal anterior tubercles, Physocoryna expansa Pic, Octhispa haematoppyga bear three apical femoral setae, have spiracles on I- (Baly), and a species of Probaenia lay their feces VII (VIII), posses relatively unsculptured abdomen, on special frass-lined resting mines (Hespenheide, have two bosses, each with four tubercles, and lack 2000; Hespenheide & Dang, 1999). In the latter three urogomphi. Pupae of the Criocerinae are exarate and species, feeding occurs in tunnels radiating from the of variable color, have a glabrous head, possess two resting area (Hespenheide, 2000; Hespenheide & to three papillae on each antennal segment, usually Dang, 1999). Some possible developmental transi- bear two pronotal anterior tubercles, lacking apical tions between mines that are serpentine, blotched, femoral setae, have spiracles on I-VII (VIII), pos- or blotched with functional compartments are rep- sess a microspiculed abdomen, lack a boss, and bear resented in Figure 8. However, this Figure is not a paired urogomphi. Pupae of the Galerucinae are phylogenetic hypothesis. exarate, whitish/yellowish, have a setose head, pos- sess two to five papillae on each antennal segment, Pupa lack pronotal anterior tubercles, bear one to three apical femoral setae, have spiracles on I-IV (V-VIII), Chrysomeloid pupae have been thoroughly studied possess a relatively unsculptured abdomen, lack a and diagnosed to the subfamily level (Cox, 1996). boss, and lack a urogomphi. Pupae of the Alticinae Most chrysomelid pupae, including those of leaf- are exarate, of variable color, have a dorsally se- mining species, are exarate, having free appendages. tose and ventrally glabrous head (and rest of body), Nevertheless, many leaf-mining hispines have (or possess two to five papillae on each antennal seg- almost) obtect pupae, presumably a trait secondarily ment, lack pronotal anterior tubercles, bear one to derived from exophytic ancestors (Cox, 1996). three apical femoral setae, have spiracles on I-IV Herein, I uniformly summarize and rearrange Cox’s (V-VIII), possess a relatively unsculptured abdomen, (1996) diagnoses for each of the subfamilies of Chry- lack a boss, and bear paired urogomphi or lack it 14 J.A. Santiago-Blay altogether. Pupae of the Hispinae are (almost) ob- ham et al., 1928). In the hispine Platypria andrewesi tect, yellow-brown, have a variably setose head, lack Weise and in P. coronata (Guérin-Méneville), pu- papillae on each antennal segment, lack pronotal pation occurs in a special leaf mine excavated be- anterior tubercles, possess one to five apical femo- fore pupation (Bernon & Graves, 1979; Zaka-ur-Rab, ral setae, bear spiracles on I-IV (V-VIII), have an 1991). Platypria coronata larvae, crawl out of the abdomen that sometimes bears two rows of spinules, feeding mine and move to the upper epidermis to- lack a boss, and possess a paired urogomphi, or lack wards the leaf’s apex following the margin of the it altogether. Pupae of the Cassidinae are (almost) leaf. Shortly before reaching the apex, the larvae obtect, yellow-brown, have a variably setose head, move mesally, crawl into the leaf, and form pupa- lack papillae on each antennal segment, lack pronotal tion mines. During this period (approximately 45 anterior tubercles, possess one to five apical femo- minutes), larvae are susceptible to predation, espe- ral setae, bear spiracles on I-IV (V-VIII), have an cially by ants. Chittenden (1902) also reported that abdomen that sometimes bears two rows of spinules, the transformation from prepupa to pupa in the his- lack a boss, and possess paired urogomphi, or lack pine Odontota dorsalis takes two to three minutes. it altogether. See also Rane et al. (2000) for addi- In the alticine Clitea picta, pupation may take place tional details on pupae of leaf-mining Cassidinae. in the mine or in the soil (Zaka-ur-Rab, 1991). Some hispine pupae, such as those of Sceloenopla Biology of the pupae of leaf-mining chrysomelids near bidens, are strongly dorsoventrally depressed, Pupation in leaf-mining chrysomelids takes place in allowing for movement within the mines excavated an often specialized leaf-mine, or in cells in the soil by larvae. Ford & Cavey (1985) reported that prongs which are lined with a smooth inner wall (Cox, 1996). on the seventh abdominal sternite of several genera According to Cox (1996), zeugophorines pupate in of North American hispines are ambulacral. Some earthen cells. Among the criocerines, pupae lay inside hispine pupae are remarkably fast movers. For in- “whitish cocoons constructed from mouth exudate stance, in the hispine genus Anisostena, pupae have … attached to the host-plant or in the soil; or in been observed to move at a rate of about 5 cm per earthen cells lined with mouth exudate”. Oulema second (Ford & Cavey, 1985). pumila, according to Vencl & Aiello (1997), one of It is not known how leaf-mining larvae arrive at only two criocerines known to be leaf-miners, ap- pupation sites located in the soil. Do they crawl down pear to produce a ‘foamy substance’ that coats the the shoot to reach the soil; do they simply drop; or pupation chambers in leaves. Additional details of do they do both? Cox (personal communication to the pupation biology of leaf-mining criocerines are Santiago-Blay, July 2003) suspects that often lar- not yet available, although it is known that other val legs, particularly of leaf-mining alticine larvae, criocerines, such as Lilioceris lilii Scopoli and Crio- are too short to be used in crawling, and it is more ceris asparagi (Linné) do pupate in the soil. For the likely that they just drop down. Pupation in the soil galerucine genus Monoxia and many hispines, inclu- involves considerable mortality risks, but no one has ding several North American taxa (Boldt & Staines, studied whether such dangers are significantly dif- 1993; Clark, 1983; Ford & Cavey, 1985; West & ferent from those present when pupating in or on Lothian, 1948), as well as Indo-Pacific palmophilous leaves, which are also exposed to parasitoids, preda- hispines (Mariau, 1988), and the galerucine genus tors, and diseases. Monoxia (Figs. 4 and 5), pupation takes place in- side the leaf mine excavated by the larvae. In other Adults leaf-mining chrysomelids, such as species of Dicla- dispa, there is a newer and shorter leaf compartment, Once adulthood is reached, some imagoes, such as known as ‘pupation mine’, in which they pupate those of the hispines Sceloenopla maculata (Olivier), (Hering, 1951). Larvae of the cassidine Notosacantha Microrhopala, and Promecotheca couruleipennis vicaria vacate their mined leaves and form a new Blanchard stay inside the larval mine for several days pupation mine in another leaf (Rane et al., 2000). before emerging from the leaf (Andrade, 1984; Clark, Once pupal cells are formed, the leaves of the 1983; Dharmadhikari et al., 1977). Adults chew a asteraceaean that harbor Microrhopala xerene his- hole in the mined leaf and emerge (Dharmadhikari pines inflate slightly and form a hard blister (McCau- et al., 1977). Among numerous leaf-miners, emer- ley, 1938). Several species of Central American gence holes are species specific features; leaf-min- hispines have resting mines in which they pupate ing coleopterans often obliterate these emergence (Hespenheide, 2000; Hespenheide & Dang, 1999). holes as they eat their way out of the mine (Auerbach Interestingly, the hispine Dicladispa testacea (Fab- & Simberloff, 1988). Once beetles emerge, they seek ricius) pupates in the “larval mine in leaf midrib” food (see Host-plant feeding preferences, below) and of Cistus sp. hosts (Cox, 1996). mates (see Reproduction, below). Baliosus nervosus On the other hand, in a significant number of leaf- total adult longevity has been estimated as eleven mining chrysomelids, such as Dibolia borealis, spe- months (West & Lothian, 1948). Adult chrysomelid cies of Argopistes (both Alticinae), and several other longevity ranges from a few days to over two years flea beetles, pupation occurs in earthen cells within (Hinton, 1981). the soil (Cox, 1996; Frost, 1924; Hering, 1957; Need- Leaf-mining chrysomelids 15

Ecology: the leaf as a habitat for survival and the hispines Octotoma scabripennis Guérin-Méne- reproduction ville and Uroplata girardi Pic for the biological control of noxious Lantana (Verbenaceae) weeds. Finding a suitable host plant for survival and repro- Beetles as well as larvae were starved – a common duction has been the subject of considerable research. part of the protocol in this type of experiments – These areas are important in the context of the greater and caged plants, usually of economic importance, understanding of insect-hostplant interactions and were exposed to the herbivores for a predetermined possible coevolution (Ehrlich & Raven, 1964). In time, usually 24-48 hours. Subsequently, the evidence simple terms, the host-seeking process can be di- of feeding and oviposition were observed on the vided into: initial (long-distance) orientation; con- plants as feeding marks or as deposited eggs, respec- tact (short-range) orientation; and behavior once tively. Cilliers (1987b) used exclusion experiments contact with the host-plant is made (Moldenke, 1971). to demonstrate that the presence of leaf-mining chry- Very little is known about these phases of herbivory somelids causes significant defoliation, decreased leaf for leaf-mining chrysomelids. size, flower production, and seed set in Lantana camara. Use of leaf-disc tests, standard procedure Host-plant feeding preferences: the patterns when the testing potential biological control of weed agents, were used by Vig (1998a,1998b) to show the When I began compiling these data in 1995, I had feeding preferences of adult Phyllotreta vitata (Alti- only partially realized the immensity of the task. cinae) to grasses (Poaceae) and crucifers (Brassica- Table 1 is a worldwide list of leaf-mining chry- ceae). An important distinction, not always made somelid taxa, including their geographical distribu- particularly in the context of applied research, is the tion, host-plants, and their taxonomic authorities, as ability of adults leaf-mining chrysomelids to oviposit well as pertinent references. It is not as comprehen- only on a limited suite of plant species although the sive as I would have liked. Nevertheless, I am cor- beetles may have fed on a larger group of plants and recting some deficiencies (listed below) and hope occurred on many additional plant species. Obser- to have additional and detailed quantitative analy- vations of species of Argopistes have shown that A. ses at a later date. Preliminary analyses of the data cocconelliformis oviposits and larvae develop only follow after mention of some associated caveats. in new leaves of their oleacean host-plants (Inoue & Shinkaji, 1989). Studies on Dicladispa armigera, Problematic data set the rice hispa, have confirmed that sitting, feeding, There are numerous difficulties with the data regard- and ovipositing host-plants need not be on the same ing the feeding behavior of leaf-mining chrysomelids. plant. Firstly, it is often unclear what records are actual Thirdly, authors frequently do not distinguish feeding events or which are cases of ‘accidental’ between adult and larval feeding. A case in point is sitting, feeding, nibbling, or ovipositing on a plant that of the Dicladispa armigera, the rice hispa, in (Harley 1969; Mullins, 1976; Razzaque & Karin, Bangladesh. Razzaque & Karin (1989) tested the 1989). feeding preferences of adult D. armigera on several Secondly, there are relatively little experimental cultivars, including corn, wheat, rice, and several and quantitative data on potential host-plant feeding weeds. While numerous plants were heavily used preferences of larvae and adult leaf-mining chryso- as settling substrates and/or food items by adult D. melids. However, experimental approaches to leaf- armigera, Razzaque & Karin (1989) reported that mining are not new. For instance, Buhr (1955) reports only rice (Oryza sativa Linné, Poaceae) served as an unsuccessful attempt to rear larvae of the Brassica- an oviposition site and as the host for larval devel- ceae colonizer Phyllotreta nemorum (Linné) on a opment. For the purposes of Table 1, when the life- noncriciferous plant, Allium moly Linné (Lilliaceae). history stage of an insect is not mentioned in a source, Dominique Mariau has spent a great part of his I have conservatively assumed that the context re- professional life in the Ivory Coast of Africa open- fers to adults. ing palm leaves with a knife to insert larvae of several Fourthly, there is rarely published mention of species of Hispinae to test whether they can com- voucher specimens for the insects or plants depos- plete development on artificially-produced mines ited in a collection, an issue that is especially acute (Jolivet to Santiago-Blay, personal communication, for evidence of leaf-mining. Vouchers would greatly June 2003). If there are experimental trials, the lit- facilitate verification of identifications by subsequent erature tends to omit whether the used leaves were researchers. unexcised and, if excised, how often they were re- Fifthly, the taxonomic status of numerous leaf- plenished. Experimental studies of feeding behav- mining chrysomelid genera, as well as of their host ior by and preferences of leaf-mining chrysomelids plants, varies greatly. In some cases, many species are relatively uncommon, unless the species in ques- remain to be described, particularly tropical leaf- tion is of potential economic importance (see ex- mining chrysomelids. amples in Cilliers, 1987b; Harley, 1969; Hodson, Sixthly, in those cases where the genera have been 1942; Kogan & Kogan, 1979; Richerson & Boldt, well-studied, I have yet to find a case where there 1995). Harley (1969) explored the possibility of using are rigorous phylogenies for both the leaf-mining 16 J.A. Santiago-Blay

10. 9.

12.

11.

Figs. 9 to 12. Distribution of reported plants serving as hosts for leaf-mining in different subfamilies of the Chrysomelidae. 9. Zeugophorinae. 10. Galerucinae. 11. Alticinae. 12. Cryptostomes (Hispinae + Cassidinae). While there are less data available for larvae than for adults, simple inspection strongly suggests that the range of plants serving as hosts for leaf-mining chrysomelid larvae is smaller than that of adults. IFR for the comparison of subfamily calculations support this: Zeugophorinae (IFRi = 2.0, IFRa = 2.9), Galerucinae (IFRi = 1.0, IFRa = 2.4), Alticinae (excluding Phyllotreta nemorum), IFRi = 2.7, IFRa = 3.8), and Hispinae (IFRi = 1.6, IFRa = 3.0). For adult zeugophorines, 55% of the reported species only feed on one genus, and 82% of the reported species only feed on one plant family. For adult galerucines, 32% of the reported species only feed on one genus, and 60% of the reported species only feed on one plant family. For adult alticines, 47% of the reported species only feed on one genus, and 71% of the reported species only feed on one plant family. For larval Cryptostomes, 77% of the reported species only feed on one plant species, for adult Cryptostomes, 51% of the reported species only feed on one plant species. chrysomelid group and its host-plant taxa. This would With the exception of four reported fern-feeding greatly facilitate tests of coevolutionary hypotheses. species of alticines (two in Schenklingia and two in Interestingly, Pasteels et al. (2003) have found out Febra) (Kato, 1991, and Samuelson, 1973, respec- that the patterns of chemical defenses of chrysomelids tively) and the Equisetum (horsetail) associate (Hip- are more conserved than insect-hostplant affiliations. puriphila moderii Linné; see Table 1 for references), Evidently, more species need to be described, al- leaf-mining chrysomelids appear to feed exclusively pha taxonomy refined even further, and revisionary on angiosperms. No chrysomelids have been reported works pursued before most of the tests can be imple- as gymnosperm miners. mented. As with leaf-mining insects as a group (Hes- Seventhly, authors often report the host plant by penheide, 1991), leaf-mining chrysomelids are be- its common name, only complicating species iden- lieved to be narrow spectrum foliovores or, as Wilcox tification. (1979) says, “They [chrysomelids] usually show Eighthly, the data are scattered over a wealth of some degree of specificity”. While leaf-mining chry- languages and countries and are often buried in somelids certainly have narrower feeding preferences taxonomic papers, making access difficult. than the Coleoptera as a group (Bernays & Chapman, Ninthly, some of the papers used have summa- 1994), numerous leaf-mining chrysomelids have rized information secondarily. Often, in the inves- more catholic preferences than usually suspected for tigative process, it is difficult to recover the primary leaf-mining organisms, frequently feeding on several data. congeneric, confamilial, or even rather distantly related plants. Hispines, which constitute most of Data analyses the leaf-mining chrysomelids, tend to be more poly- Clearly, leaf-mining is concentrated into several phagous as adults than as larvae (Ford & Cavey, tribes of what have traditionally been called ‘Hispi- 1985) and, as a subfamily, prefer monocotyledoneous nae’ (Seeno & Wilcox, 1982), now called the ‘Cas- plants (Borowiec, 1995, 1999; Jolivet, 1989a; Wilf sidinae’ (Staines, 2002b, 2004b) or, formerly, the et al., 2000). The same patterns hold true for the Cryptostomes. remaining subfamilies of leaf-mining chrysomelids. Leaf-mining chrysomelids 17

There are numerous qualitative classifications of ing is more restricted (especially for larvae), and the range of host-plant feeding by hervirores (e.g., oviposition even more. Hering 1951). In an effort to quantify the degree of The IFR has one major drawback: it does not polyphagy of a given taxon, I have created an in- easily lend itself to account for the chemical affini- dex of feeding range (IFR), which can be calculated ties of the objects being consumed (e.g., host plants). as a simple arithmetic average. For a given taxon, There are also smaller problems. For instance, what the IFR is defined as: to do when a plant is mentioned only to the level of genus? I counted it, conservatively, as one species, IFR = total number of entities (e.g. plant species) unless there are congeneric host-plants mentioned for the same leaf-mining chrysomelid. In this case, consumed by organisms in group being compared I ignored the Genus sp. What follows is a prelimi- (e.g. leaf-mining chrysomelids) / total number of nary analysis of the data compiled on host-plant consumers being compared feeding for larvae and for adults (Table 1). More detailed analyses are currently in progress. The IFR varies from 1, or strict monophagy, as in It is quantitatively clear that, for each subfamily many species of leaf-mining hispines, to a very large of leaf-mining chrysomelids, the IFR of larvae is number, illustrated by the rampant polyphagy of smaller than the IFR of the adults. For the Chryso- organisms with little discrimination whatsoever, such melidae, the overall IFR is 1.7 for larvae and 3.1 as the alticine Phyllotreta nemorum. The IFR, a scalar for adults. Most plants serving as feeding hosts of quantity (units consumed per organisms doing the leaf-mining chrysomelids are from relatively mod- consumption, e.g., host-plant species per species of ern lineages (Judd et al., 2002), just as in the Bu- leaf-mining chrysomelid), can be adjusted for the prestidae (Hespenheide, 1991). group doing the feeding (e.g., individuals, species, Zeugophorinae: IFR for larvae (IFRi) is 2.0 and genera, subfamilies, etc.). I have accounted neither IFR for adults (IFRa) 2.9. The plant families (and for plant phylogeny nor for plant chemical affinity genera, parenthesized) most commonly reported serv- in the IFR calculations presented in this work, al- ing as host plants for the Zeugophorinae are the though the former can easily be solved by differen- Salicaceae (Salix and Populus), the Betulaceae (Be- tially weighing distinct families, orders, or classes tula and Corylus), and the Celastraceae. For adult of host-plants fed upon by a given organism. The zeugophorines, 55% of the species reported only feed ‘unit’ as well as the ‘entity’ terms can also be adjusted on one plant genus, and 82% of the species reported by taxonomic rank (different species, different genera, only feed on one plant family. Figure 9 summarizes etc.), or by relative feeding (as in weighed indexes, the distribution of reported host-plants for larvae and particularly in the context of experimental trials). adults in the Zeugophorinae. As an example, this is what would have to be done Trichostomes. Galerucinae: IFRi is 1.0 and IFRa 2.4. The plant families (and genera, parenthesized) to make a simple quantitative statement about the most commonly reported serving as host plants for host-plant species feeding range for the adults of the the Galerucinae are the Asteraceae and the Cheno- reported species of leaf-mining Criocerinae (only two podiaceae (Atriplex, Chenopodium, Suaeda, etc.). For species). Firstly, obtain a total number of species adult galerucines, 32% of the species reported only that are being consumed by the Lema quadrivittata feed on one plant genus, and 60% of the species Boheman and by Oulema pumila Vencl and Aiello. reported only feed on one plant family. Figure 10 In this case, a minimum – hence a conservative summarizes the distribution of reported host plants estimate – of one if Commelinaceae and Piperaceae for larvae and adults in the Galerucinae. are counted as one species or one genus. Secondly, Alticinae: IFRi is 2.7 and IFRa (excluding the data calculate the resulting IFR. In this case, the IFR is for Phyllotreta nemorum) 3.8. The plant families 1, meaning that the leaf-mining criocerines are mo- most commonly reported to be serving as host plants nophagous at the level of resolution one is examin- for the Alticinae are the Brassicaceae, Lamiaceae, ing, in this case, species. Of course, we could have Asteraceae, Plantaginaceae, Schrophulariaceae, Poly- reached the same conclusion by simple inspection gonaceae, and Poaceae, but many more families, and of Table 1. numerous genera have been reported as host plants. Highly polyphagous species, such as the alticine For adult alticines, 47% of the species reported only Phyllotreta nemorum (Linné) have a very high IFR. feed on one plant genus, and 71% of the species One should compare similar terms (e.g., subfami- reported only feed on one plant family. Figure 11 lies of leaf-mining chrysomelids, as done below) and summarizes the distribution of reported host plants be aware of the fact that the results are as good as for larvae and adults in the Alticinae. the data upon which they are based. For example, Cryptostomes (Hispinae + Cassidinae): IFRi 1.6 Vig (1998b) and Vig & Verdyck (2001) have shown and IFRa is 3.02. The plant families (and genera, how variable feeding preferences can be in several parenthesized) most commonly reported serving as species of Phyllotreta. In addition, different activi- host plants for the Hispinae are the Arecaceae (Cocos, ties in the life of an organism may have different Metroxylon, and numerous other palms), Pandana- sets of ‘host’ ranges. For instance, while staying ceae (Pandanus, Freycinettia), and Zingiberaceae in ‘idle’ may happen almost anywhere, host-plant feed- the Old World. Numerous Leguminosae, Asteraceae, 18 J.A. Santiago-Blay

Poaceae, and Verbenaceae serve as host-plants for tion is restricted to studies on Phyllotreta armoraciae. hispines in the New World. For larval Cryptostomes, Karoly Vig has generously allowed me to borrow 77% of the species reported feed on one plant spe- from a recent paper of his (Vig, 1999). cies. For adult Cryptostomes, 51% of the species Phyllotreta armoraciae feeds on several crucif- reported only feed on one plant species. Figure 12 erous plants to the same extent as it does on horse- summarizes the distribution of reported host plants radish, Armoracia rusticana Gaertner, Mey, and for larvae and adults in the Cryptostomes. Scherbius (Brassicaceae), but it rejects more than For central European chrysomelids at least, while half the investigated species (Nielsen et al., 1979a). the genera tend to be relatively selective in their Both accepted and rejected cruciferous species con- overall habitat, their feeding preferences are broader tain glucosinolates in large quantities. Glucosinolates (Schöller, 1996). This broad host-plant feeding capa- are known as important feeding stimulants for P. bility parallels studies in phytophagy on southern armoraciae and for other species of Phyllotreta California weeds on the Ambrosiinae (Asteraceae) (Hicks, 1974). Horseradish contains mainly ally- (Goeden & Teerink, 1993): 73% polyphagous and glucosinolate but 2-butyl- and benzylglucosinolate 10.9% endophytic in leaves, although it is somewhat are detected as well, in traces. In spite of the fact difficult to directly compare these two sets of data. that Brassica nigra (Linné) Koch, Alliaria petiolata Many of the host plants listed in Table 1 are (M.B.) Cavara et Grande, Iberis umbellata Linné, common weedy plants. However, as life histories and Thlaspi arvense Linné (all Brassicaceae) have are studied in more detail, additional host plants will a very similar glucosinolate content as horseradish be found, and corrections made to previously pub- (Kjaer, 1976), P. armoraciae feed only on B. nigra. lished reports, particularly if they have emphasized Nasturtium microphyllum Bönningh (Tropaeolaceae), adults instead of larvae (c.f. Microrhopala; Clark, Sinapis alba Linné, and Sisymbrium officinale 1983). (Linné) Scopoli do not contain allylglucosinolate but, Some chrysomelids appear to be facultative leaf- under laboratory conditions, they were all eaten in miners and they tend to be (potentially or actually) appreciable amounts by P. armoraciae. Glucosinolate oli- or polyphagous. Of all leaf-mining chrysomelids mixtures isolated from N. microphyllum, S. officinale, Alyssum saxatile Linné, and from Cardamine amara listed, the alticine Phyllotreta nemorum has the Linné were more stimulatory than the glucosinolate largest number of recorded hosts (134); most host mixture from horseradish. No correlation was found plants are in the Brassicaceae. This species has been between plant acceptability and stimulatory activ- recorded on at least 110 crucifers in central Europe ity of glucosinolate mixtures isolated from the afore- and Poland (Lipa et al., 1977), although it does not mentioned plants (Nielsen et al., 1979a). mine all the species listed. Phyllotreta vittata has Usually, crucifer-feeding insects can discriminate been associated with 89 host plants from the same between different glucosinolate containing plant region, although it mines only two of them. Inter- species. According to Nielsen et al. (1979a), the estingly, in his extensive studies of P. vittata, Vig horseradish flea beetle, Phyllotreta armoraciae (personal communication to Santiago-Blay, April (Koch), cannot recognize horseradish solely by its 2003) has never seen this species mining leaves. glucosinolates content or by the hydrolysis products Similar reports exist for P. armoraciae (Vig & Ver- released from glucosinolates. In further experiments, dyck, 2001). It appears that all these species are two flavonol-glycosides were isolated from water facultative leaf-miners as well. If this is correct, the extracts of horseradish leaves. Larsen et al. (1982) ecological transition from exo- to endophyty may identified the flavonol-glycosides as 3-O-[2-O-(β- be evolutionarily simple. D-xylolpyranosyl)-β-D-galactopyranosyl]-kaempfe- rol (compound I) and it was present at high Chemical correlates of feeding behavior concentration in the leaves throughout the growing season. A second compound, 3-O-(2-O-(β-D xylol- Apparently, there are very few studies on the chemi- pyranosyl)-β-D-galactopyranosyl]-quercetin (com- cal correlates of feeding behavior in leaf-mining pound II) is less phagostimulatory to P. armoraciae chrysomelids. Some species of Phyllotreta are wide- than compound I. Combinations of allylglucosinolate spread, making one wonder what all the host plants and compound I are more stimulatory than any of may have in common. In Poland and central Europe the compounds alone (Larsen et al., 1982; Nielsen alone, over 750 species of insects have been reported et al., 1979b). for crucifers (Lipa et al., 1977). Nearly all species Flavonol-glycosides with different sugar moieties of Phyllotreta (Alticinae) feed on crucifers or on are widely distributed compounds in crucifers. It related genera in the Resedaceae and the Cappara- seems that P. armoraciae is able to distinguish ka- ceae. The only documented exception to this feed- empferol-glycosides with different sugar moieties. ing pattern on a species of Phyllotreta is P. vittula Simultaneous presence of kaempferol glycoside Redtenbacher, which feeds on grasses and cereals, (compound I) and glucosinolates could be the key but it is still attracted to crucifers (Kostromitin, 1973; stimulus determining the recognition of horserad- Vig, 1998a,b). Oligophagy is the characteristic fea- ish by P. armoraciae suggesting that other feeding ture of species of Phyllotreta, but some species are stimulants also contribute to the palatability of dif- monophagous. The remaining portion of this sec- ferent host-plant species to the horseradish flea beetle. Leaf-mining chrysomelids 19

Vig has observed Phyllotreta armoraciae feeding Biotic factors: host-plant and natural enemies on Capsella bursa-pastoris (Linné) Medic., Arabis Obviously, host plant is a major correlate for the sp., and on Alyssum saxatile Linné (all Brassicaceae), presence (and abundance) of leaf-mining chrysome- even under stressful laboratory conditions. P. armo- lids (Table 1). Most leaf-mining chrysomelids have raciae also ate small amounts of leaves from Bras- a relatively narrow host-plant range. In species of sica napus Linné, Barbarea vulgaris R.Br., and the galerucine genus Monoxia, it appears that most Alliaria petiolata (Vig & Verdyck, 2001). species are narrowly oligophagous, at least as adults. Thousands of host-plant preference studies using clip Ecological correlates of feeding by leaf-mining cages (Fig. 6) holding unexcised leaves of potted chrysomelids composites and chenopods (most reported host plants of the genus) clearly point to adult stenophagy to Numerous abiotic and biotic factors have been as- monophagy. Since over 100 plant genera were used sociated with the presence (and abundance) or ab- in these studies, and they most likely represented a sence of leaf-mining chrysomelids. Over 65 years wide spectrum of leaf structures and chemistries, I ago, Maulik (1937) insightfully discussed the im- hypothesize that both chemistry and surface mor- portance of both types of factors on hispine-host- phology are important determinants of feeding be- plant associations. In cases where the host-plant is havior. Perhaps the best demonstration of this feeding eclecticism on leaf-mining chrysomelids is the case present and the phytophagous insect absent, Maulik of a possible new species, Monoxia near inornata suggests that “the host plant can throw off its in- Blake. Experimental studies show that, as adults, this sect enemies under certain conditions”. Conversely, gum plant (Grindelia spp., Asteraceae) associate if the phytophagous insect is present and the host- feeds on every species of Grindelia tested, as well plant is absent, “the former [namely, the insect] must as on several other confamilial species in different have another host-plant on which it is able to adapt tribes of asteraceans (Santiago-Blay, 1990). How- itself”. ever, in all the cases, the leaves of the plant species fed upon by Monoxia near inornata, were relatively Abiotic factors: stress and shade coriaceous, glaucous, and of moderate thickness, An outbreak of the hispine Odontota dorsalis in the suggesting that, in this case, gross leaf morphology Appalachian mountains of southwestern Virginia is related to feeding behavior. The preference for (USA) was tentatively attributed to drought-stress, certain leaf thickness is so striking that, with some possibly because of changed physical and chemical experience, one can learn to accurately guess, within conditions. That stress made the host plants more a taxonomic range, which host plants are likely to attractive to would-be herbivores. In this case, O. be eaten by the adults of this species. In connection dorsalis fed on six sympatric trees belonging to four with the biocontrol of weeds, Harley (1969) reports different vascular plant families: Acer saccharum that starving adults, not larvae, of Octotoma scabri- (Aceraceae), Quercus prinus, Q. rubra (Fagaceae), pennis and Uroplata girardi almost choose not to Robinia pseudoacacia (Leguminosae), Crataegus feed on plants other than their hosts. coccinea, and Prunus serotina (Rosaceae). Sceloenopla maculata (Olivier) is freer to feed Abundant circumstantial evidence scattered on Cecropia lyratiloba var. nana when the Azteca throughout the literature suggests that the larvae of ants (Formicidae), which typically inhabit plants of some leaf-mining chrysomelids prefer leaves located this genus, do not fully utilize the plants (Andrade, on relatively shadier portions of plants. For instance, 1984). Leaf-mining chrysomelids have such numer- late in the 19th century, Packard (referred by Hodson, ous natural enemies that, in some cases have a sig- 1942,) reported that the foliage of host basswoods nificant impact on their populations (see Natural (Tilia americana, Tiliaceae) is destroyed by the biological enemies and other mortality factors, hispine Baliosus nervosus, with the exception of the below). [foliage] of very tall trees. Ford & Cavey (1985) report larvae of the hispine Anisostena nigrita (Oli- How do plants that are attacked by leaf-mining vier) mine blades of Schizachyrium scoparium (Poa- chrysomelids respond to herbivory? ceae) when the leaves were shaded, but not when exposed to direct sunlight (Cappuccino, 1991a,b; In some cases, leaf abscission has been reported in Damman & Cappuccino, 1991). They noted that conjunction with severe leaf-mining (West & Lo- hispine mines tend to occur in shaded or partially thian, 1948; Inoue & Shinkaji, 1989), but it has been shaded leaves. Ford & Cavey (1985) also observed argued that this is simply a generalized response to that, when host plants are located in sun-exposed leaf damage, and not a means to regulate popula- areas, mines, if any, occur in drooping or lower tions of leaf-mining herbivores (Hespenheide, 1991). branches which are more likely to contain shaded Some plants form a thin callus, or loose aggregate leaves. These observations suggest a negative rela- of parenchyma cells, as a reaction to leaf-mining tionship between sunlight exposure and presence of (Hering, 1951). If the herbivorous attack occurs early hispine mines. in the development of the leaf, serious deformation and leaf asymmetry may follow (Hering, 1951). There are some reports of ‘green islands’ caused by 20 J.A. Santiago-Blay miners’ activities in senescing leaves (Connor & identified in the source, are most abundant at the Taverner, 1997; Hespenheide, 1991; Hering, 1951). highest and wettest station, Cerro Campana (http:/ This possible cytokinin-analogue extends cell growth, /www.stri.org/tesp/Intro%20-%20Insects.htm), mat- hence a source of nourishment, after abscission. At ching well with the seasonality of many organisms times, mining insects attack fallen leaves and this in that part of the world (Leigh et al., 1996). Frost represents an intermediate between leaf-mining and (1931), who also studied hispines in Panama, ob- decomposition (Hering, 1951). served that the mines of hispines are scattered on host plants, with one or two mines on a plant. Do leaf-mining chrysomelids attack aquatic plants? According to Frost (1931), hispine mines are scat- tered, “with seldom more than one or two [mines “There are of course very few aquatic insects that each with one individual] on a single plant”, per- are specialists in feeding on individual species of haps because of their low fertility rate (Mariau, 1988). plants (a caddisfly on a red alga, a midge on a blue Nevertheless, in species of Monoxia, there are sev- green alga, maybe some other midges on [species eral mines on a plant, but just one mine per leaf (Fig. of] Potamogeton [pondweeds, Potamogetonaceae])” 3). When present, the adults are easy to collect as (Resh to Santiago-Blay, personal communication, they are frequently found resting on their host food July 2003, bracketed words added by Santiago-Blay; plant. see also Hering, 1951). While no definitive aquatic There appear to be no published studies of the leaf-mining chrysomelids appear to have been con- interaction of leaf-mining chrysomelids and endo- firmed, there are at least two reports of leaf-mining phytic fungi. However, a splendid case of Pro- species on aquatic emergent plants. Gressitt (1960) mecotheca papuana Csiki infected by a fungus is reports that Cyperispa thoracostachyi Gressitt pu- illustrated in Howard et al. (2001). Faeth & Hammon pates, “at extreme base of larval mines at base of (1996) suggest a possible relationship between en- long leaves of large sedges [Cyperaceae], often at dophytic fungi and Cameraria (Lepidoptera: Gracil- or below the surface of water in swamps”. Cox (1996) lariidae) leaf-mining larvae by differentially affecting report of a species of cryptonychine hispine, Cal- dispersion and colonization in different host-plants. listola sp. as a miner was a lapsus (Cox to Santiago- In addition to the host plant, leaf-mining chrysomelid Blay personal communication, July 2003), he meant and parasitoid systems, three trophic interactions C. thoracostachyi. Collart (1934), reports that the involving fungi may prove to be biologically inter- hispine Dicladispa viridicyeana (Kraatz) is associ- esting and to be of applied importance for some leaf- ated with the large aquatic grasses of the genus mining chrysomelid pests. For instance, Kalshoven Vossia, although he does not indicate where they live (1981) reports that the damage caused in coconut with respect to the waterline. Undoubtedly, many palms by the hispine Promecotheca soror is increased other species of leaf-mining chrysomelids feed on by the entry of spores of the fungus Pestaloptiopsis emergent plants but it remains to be seen how many, (Pestalozzia). Hering (1951) includes a discussion if any, are truly living underwater and how they cope on heterospecific interactions between leaf-mining with that environment. insects.

Influence of genetics Temporal distribution Long-term studies of the oil palm leaf-mining hispine Conspecific populations of chrysomelids have been Coelanomenodera elaedis Maulik in western Africa found to differ on their host-plant feeding prefer- have shown sudden shifts from mixed instar popu- ence. Vig (1996) suggests that some of the varia- lations to synchronized populations of one instar tion in host-plant feeding preference has a genetic during outbreaks (Bernon & Graves, 1979). For basis. Genetically-influenced changes of host-plant several decades, hispine pests of coconut and oil feeding preference may be important in determin- palms have been studied extensively by Mariau, ing the evolutionary history of a lineage of phytopha- Lecoustre, and their collaborators. They reported gous organisms. cyclical hispine population changes (Mariau & Morin, 1972), which are tracked by some of their Spatial and temporal distribution parasitoids (Lecoustre & Reffye, 1984). Modeling the population dynamics, including the potential The use of chrysomelids, including leaf-mining effects of human intervention (e.g., pesticide appli- forms, has been suggested for monitoring local spe- cations, pruning, etc.) may help predict and reduce cies richness in natural areas (Staines & Staines, great losses to these hispines (Lecoustre & Reffye, 1998). 1984). More cyclically extreme and synchronized population dynamics of chrysomelids have been Spatial distribution described by Kovalev (2004). Strogatz (2003) wrote In a long-term study of Panamanian insects, research- a thought-provoking and fascinating book discuss- ers found that leaf-mining chrysomelids are not ing a multiplicity of systems in which synchroniza- equally distributed along an intranational transect tion arises from apparent chaos. of Malaise traps. These leaf-miners, which are not Furthermore, populations of C. lameensis Berti Leaf-mining chrysomelids 21 and of C. minuta Uhmann are greatly affected by pause as beetles under debris or in the soil. In spe- air humidity and food supplies (Mariau & Lecoustre, cies of Microrhopala, some overwintering sites have 2000; Lecoustre & Reffye, 1984). In general, hispines been found near roots about 10 cm under the soil appear to be especially sensitive to humidity and surface (Clark, 1983; Ford & Cavey, 1985; Hodson, temperature in comparison to other chrysomelids. 1942; West & Lothian, 1948). Leaf-mining chrysomelids are no different from most other insects in their general life history pat- Defensive behavior and mimicry terns and voltinism. Also, they have considerable variation in the number of generations with the spe- It seems that some leaf-mining chrysomelids find cies and latitude. In temperate zones, leaf-mining mines to be a relatively safe retreat from neighbor- chrysomelids are usually univoltine (Hering, 1951). ing predators. For example, Andrade (1984) and Some cases of bivoltine, such as the galerucine Mo- Jolivet (1989b) report that leaf-mining species of noxia near inornata (referred to as possible new Sceloenopla which feed on Brazilian species of Ce- species, Santiago-Blay, 1990) occur in regions with cropia are protected from aggressive Azteca ants. a more moderate climate. Octotoma scabripennis and Mines provide an environment with proximity to food Uroplata girardi are trivoltine (Harley, 1969; Cilliers, and a hideout from some larger predators. However, 1987a). Several generations per year are possible for the mine may prove to be a trap, since it may serve Promecotheca caeruleipennis in Sri Lanka (Dharma- as a cul-de-sac from enemies small enough to get dhikari et al., 1977) and up to six for the hispine in the mine or from interactions with potentially Dicladispa armigera (Sen & Chakravorty, 1970). negatively interacting organisms, such as fungi (more Some sympatric congeneric species, such as Argo- on three-trophic interactions in Spatial distribution, pistes coccinelliformis and A. biplagiatus, differ in above). their voltinism: the former being a facultative uni- Many leaf-mining chrysomelids form what ap- voltine species (although normally univoltine), the pears to be a Müllerian mimicry complex with other latter an obligatory univoltine (Inoue, 1996). beetles, particularly with lycids and lampyrids. The mimicry complexes of chrysomelids have been sus- Biotic effects on temporal distribution of mining pected for a long time (Jolivet, 1989a; Maulik, 1919), insects. It seems that many leaf-mining chrysomelids and they include species that are leaf-miners and in temperate zones, such as Dibolia borealis, Zeugo- beetles in other coleopterous families. The similarities phora scutellaris, and Monoxia near inornata, over- involve the general body form and coloration pat- winter in the soil as adults (Needham et al., 1928; terns and/or the presence/absence of spines (Maulik, Santiago-Blay, unpublished data). There, they are 1919). Although the nature of those mimicry com- exposed to both abiotic and biotic elements, which plexes has not been experimentally tested, they are can be significant mortality factors (see Natural believed to be Müllerian (Hespenheide, 1991). How- biological enemies and other mortality factors, be- ever, while the anecdotal reports are interesting and low). West (1985) documented the negative com- compelling, I have been unable to find experimen- petitive interaction of oak-browsing lepidopterous tal evidence for these claims. I know of numerous larvae on leaf-mining lepidopterans attacking oaks. stories of similarly-colored insects allegedly form- According to West (1985), browsing larvae are more ing mimicry complexes (e.g., the widespread tropi- abundant in the spring, when the nutritional quality cal hispine Chalepus sanguinicollis (Linné) and of the foliage is higher and, later in the season, when another red and black beetle, the lycid Thonalmus the quality of oak leaves has decreased, the leaf- chevrolati Bourgeois). Experimental evidence is mining guild is more abundant. Similar multitrophic needed to evaluate all those claims. interactions remain to be discovered for leaf-min- ing chrysomelids. Natural biological enemies and other mortality factors Diapause Like many other leaf-miners, leaf-mining chryso- There is some variation in the resting stages of melids are attacked by numerous parasitoids and leaf-mining chrysomelids. Argopistes coccinelli- other biological natural enemies (Connor & Tavener, formis and A. biplagiatus (Inoue, 1990a; Inoue & 1997). Also, just like any organism, they have to Shinkaji, 1989) and many other leaf-mining chry- cope with numerous abiotic mortality factors. somelids overwinter as adults. In contrast, Octotoma scabripennis and Uroplata girardi diapause facul- Natural biological enemies tatively as adults (Cilliers, 1987a; Harley, 1969). In Leaf-mining chrysomelids have many natural bio- these cases, the exact factors involved in diapause logical enemies that regulate their populations. A are not known. Harvey (1969) speculates that de- list and/or discussion of chrysomelid parasitoids can creased autumn temperatures, shorter photoperiod, be found in Chittenden (1902), Cox (1994), Fulmek and reduced growth rate of the host plant possibly (1962), Gallego et al. (1983), Gressitt 1959, Mariau trigger diapause. (1975, 1988, 2001), Mariau & Morin (1971, 1974), Many leaf-mining chrysomelids overwinter-dia- Teixeria et al. (1999), and many others. Santiago- 22 J.A. Santiago-Blay

Blay & Fain (1994) discuss the mite associates of Eggs of the alticine Psylliodes chrysocephala chrysomelids. (Linné) suffer bacterioses and are also attacked by cantharid larvae. Their larvae are parasitized by a Some observations on general principles involved variety of hymenopterans, while the pre-pupae and in controlling leaf-mining chrysomelids by natural pupae are attacked by carabids as well as various biological enemies. More than 40 years ago, Gressitt species of nematodes. Adult P. chrysocephala are noticed a well-documented pattern of insect pests. host to fungi (Entomophtorales), gregarine protozo- Many of the most pestiferous leaf-mining chrysome- ans, and braconids (Grison et al., 1963). Mantura lids, such as several species of Promecotheca, “are chrysanthemi Kowartz, M. pallidicornis Waltl, Phyl- very scarce on their native hosts under natural jungle lotreta nemorum (Linné), Sphaeroderma rubidium conditions, but may become abundant under plan- Graells (all Alticinae) are attacked by a variety of tation circumstances or in village areas” (Gressitt, predatory beetles and/or parasitic wasps, including 1959). The fact that parasitoids seem to have broad braconids, chalcids, and ichneumonids (Fulmek, host preferences, and that they seem to be follow- 1962; Grison et al., 1963). ing the host-plants more than their herbivores, is very Various species of Promecotheca leaf-miners have useful in biological control. been controlled by a variety of parasitic hymenop- Parasitoids can use alternative insect hosts when terans in the Pacific Basin (Gressitt, 1959; Dharma- populations of economically important leaf-mining dhikari et al., 1977; Taylor, 1937). For example, P. chrysomelids, their usual hosts, are low. For this caeruleipenis Blanchard and P. papuana Csiki have reason, it is essential to provide alternative cover been controlled with Pediobius parvulus Ferrari (Eu- crops for natural enemies as a source of nectar and lophidae) and P. cumingi by Dimmnockia javanica shelter (Gallego et al., 1983). In cases of leaf-min- Ferrari (Elasmidae) and, perhaps, by Achrysocharis ing chrysomelids that oviposit into the parenchyma promecothecae Ferrari, (Eulophidae) (Dharmadhikari (Lecoustre & Reffye, 1984) and those that cover their et al., 1977). The hispines Coelaenomenodera minuta epidermally-laid eggs with a theca, parasitoids can and C. lameensis Berti and Mariau, both palm leaf- penetrate the protective layers and oviposit. Parasi- miners, are attacked by several oophagous parasi- toids seem to be effective in keeping leaf-mining toids, including Achryoscharis leptocerus Waterson populations at low numbers, including those of oil (Eulophidae) and Oligosita longiclavaita Viggiani palm leaf-miners. However, they cannot control rapid (Trichogrammatidae), as well as by several larval outbreaks (Cappucino, 1991; Mariau, 1988), in part parasitoids, including Sympiesis (Dimmnockia) abu- because parasitoids tend to have a relatively slow riana Waterson (Eulophidae), Pediobius setigerus development and life history. The importation of Kerrich (Eulophidae), Cotterellia podagrica Waterson numerous parasitoids has been far from successful, (Eulophidae), and Closterocerus africanus Waterson since it seems that they cannot adapt well to condi- (Eulophidae, perhaps also an egg parasitoid) (Berti tions outside their native range (Mariau, 1988). In & Mariau, 1999). Interestingly, the hispines Platypria one case, the effect of parasitoids and crawling pre- coronata (Guérin-Méneville) and another palm leaf- dators has been experimentally shown to be statis- miner, Coelaenomenodera perrieri Fairmaire, are tically independent of each other (Memmott et al., parasitized by a similar parasitoid complex (Mariau, 1993). There is still a lot to be learned about natu- 1988). African Balyana hispines also have numer- ral biological enemies of leaf-mining chrysomelids ous hymenopteran parasitoids (Berti & De Chenon, (Hespenheide & Dang, 1999). 1987). The larval stages of D. armigera suffer about Gressitt (1959) suggests numerous measures for 90% mortality (Sen & Chakravorty, 1970). Together the control of Promecotheca pests of palm and ca- with eggs, these two stages appear to be the most cao plantations in the Pacific Rim, including con- vulnerable in leaf-mining chrysomelids. servation and mass breeding of natural enemies; The parasite complex of the South American leaf- destruction of heavily infested host-plants (or their mining Hispolepsis spp. is quite different (Mariau, parts, unless parasitoids can be reared and the pests 1988). Thecae of Sceloenopla maculata are attacked excluded); and periodic censuses that aim to detect by chalcid wasps (Andrade, 1984). Wasp emergence early stages of infection. Hespenheide mentions that holes, possibly from mymmarids or trichogrammatids ‘unusual refuges’, which have been observed in have been reported for some undetermined tropical several tropical hispines, may reduce their probability Central American hispine eggs. Pteromalids and of being parasitized (Hespenheide, 1991). chalcids (Hymenoptera) and tachinids (Diptera) have been reported for Central American leaf-mining Some examples of natural biological enemies of leaf- hispines (Hespenheide, 1991). Additional examples mining chrysomelids. In Monoxia guttulata Blake, of parasites of leaf-mining chrysomelids can be found an unidentified tachinid as well as an unidentified in Cappuccino (1991a,b), McPheron (1985), and parasitic nematode have been detected (Santiago- Wheeler & Snook (1986). Blay, unpublished data). Together with abiotic fac- tors, these natural enemies probably contribute Other mortality factors significantly to the relatively low population num- Cappucino (1991) studied the mortality factors af- bers of this species. fecting the hispine Microrhopala vittata in south- Leaf-mining chrysomelids 23 central New York State. She discovered that early before and/or after hibernation. Both O. scabripennis leaf senescence of its host-plants, Solidago spp. and Uroplata girardi require more specific feeding (Asteraceae), and its effects on larvae, are partially and oviposition stimulants (or fewer inhibitors). responsible for the relatively low population densi- Hence, fewer plant species are acceptable for feed- ties of beetles. Larval parasitoidism by the eulophid ing and suitable for oviposition (Harley, 1969). For Chrysonotomyia spp. is another mortality factor. palm mining hispines, Mariau (1988) reports a low Exclusion experiments have shown that several Cen- fertility rate. However, occasional outbreaks of leaf- tral American species of Chalepus suffer significant mining chrysomelids do occur (C. minuta in Afri- and independent mortality from both crawling preda- can oil palms (Mariau, 1988)). tors, possibly ants, and from parasitoids (Memmott In a study of Dicladispa armigera, Sen & Chakra- et al., 1993). The coconut leaf-miner, P. cumingi, vorty (1970) found that beetles may mate for as long has been reported to be significantly controlled by as two hours, may mate more than once during a using several formulations of fungal disease (Dhar- single day, and they are polygamous/polygynous. For madhikari et al., 1977). Damman (1994) discovered Dicladispa armigera, the sex ratio is approximately that M. vittata larvae that eclose in large groups have 1:1 and the adults may live for up to two and a half a greater chance of surviving. However, once in a months (Sen & Chakravorty, 1970). mine, forming part of a large group decreases adult Kirkendall (1984) reported long postcopulatory weight. Mine initiation and larval movement to sec- escorts in Odontota dorsalis, and he hypothesized ondary mines are the most vulnerable stages in the that this behavior has evolved in situations where life history of Microrhopala vittata. the probability of encounters between the sexes is In addition to the mortality that pathogens and high and the cost of reproduction to the female is parasitoids cause, host plants of leaf-mining chry- relatively low. Males that escort females are pre- somelids respond to the attack with chemical de- sumed to have a greater probability of fathering the fenses. According to Zaka-ur-Rab (1991), when progeny from the sperm they have introduced into Clitea picta larvae penetrate the epidermis, “the site the female. (For oviposition, see Egg in Introduc- of infestation swells a little”, and the host plant, Aegle tion to leaf-mining chrysomelids, above.) marmelos Correa Serra (Rutaceae), produces exu- dates on the leaf sites chewed upon by larvae. Resin (a complex mixture of organic chemicals, especially Evolutionary and biogeography trends terpenes, insoluble in water) production is a consti- tutive plant defense against herbivores (Becerra, Leaf-mining chrysomelids, like any other special- 2003) and pathogens (Langenheim, 2003; Santiago- ized organism, represent a unique opportunity to Blay et al., 2002). There seems to be no quantita- explore the major pathways that evolution may have tive data on the mortality effect of these defenses. taken in tailoring a successful mode of life and de- Another interesting development is the use of plant tailed variations. Some leaf-mining chrysomelids are breeding to control populations of leaf-mining chry- good subjects to study the possible adaptive radia- somelids (Zheng et al., 2003). For example, larvae tion of herbivores to their host-plants, as in species of Coelaenomenodera lameensis Berti and Mariau of Monoxia (Galerucinae) (Santiago-Blay & Virkki, have “great difficulty developing on the hybrid de- 1996). In addition, several species are economically rived from the cross between E[laeis] guineensis and important (Taylor, 1937; Bernon & Graves, 1979; E. oleifera” (Mariau, 2001). Mariau, 1988; see Economic importance, below). The Many papers, and/or references therein, mention biology of leaf-mining chrysomelids is quite vari- or recommend the use of pesticides to control min- able, and I provide selected examples to show the ing chrysomelids in plantations (e.g., De & Konar, wide range of variation present. 1954; Dharmadhikari et al., 1977; Hodson, 1942; I propose a testable hypothesis that leaf mining Zabel et al., 1991). However, Kalshoven (1981) in most of the Chrysomelidae arose from ancestors warns that, in the hispine Promecotheca papuana, whose larvae were exophytic. However, in the Zeugo- the use of insecticides increases pest populations. I phorinae, leaf-mining appears to be the retained basal suspect the reason for this observation is the inter- endophytic condition. Figure 13 is a character state ference of the pesticide with the abundant agents of transition branching diagram mapped on to a recent biological control, mostly parasitic Hymenoptera. In chrysomelid phylogeny (Duckett et al., 2004). To addition, unless systemic pesticides are used, the simplify this hypothesis, the mapped character, feed- mining stages of chrysomelids are well protected ing mode (leaf-mining, in this case), is being treated inside the mines. as homologous among the lineages and as having two character states, exo- and endophyty. For a Reproduction discussion on parsimony, see Johnson (1982). Clear- ly, this needs not to be the case, at the molecular Mating behavior seems to be under more stringent genetics, morphological, or behavioral levels. Homo- control than feeding behavior. Different species of logy should be defined by ancestry and diagnosed the flea beetle Argopistes (Inoue, 1990a) may mate by criteria not related to ancestry, such as relative 24 J.A. Santiago-Blay position, development, histology, etc., as Owen and question remains: if leaf-mining is an evolutionary others said (Kaplan, 1987, 2001; Padian to Santiago- ‘dead end’, why does it keep appearing in such Blay, personal communication, July 2003). Nu- diverse groups? If the question is reworded to ad- merous pertinent and fascinating discussions on dresses the homologies related to endophyty, maybe homology at various levels of the biological hierar- we will find a better explanation. chy and in different groups of organisms can be found in Bock & Cardew (1999), Hall (1994), and Scot- Distribution of leaf-mining in the Chrysomelidae land & Pennington (2000). Except for species of Zeugophora (Zeugophorinae), Detailed analyses of what is involved in being where larvae primitively mine leaves and adults feed ‘exo’- or ‘endophytic’ will probably show that there mostly on members of the Salicaceae (Populus spp. are multiple genetic, physiological, and morphologi- and Salix spp.), the mining habit seems to have arisen cal mechanisms to attain endophyty, hence, consid- independently several times in the Chrysomelidae, erable convergence should be expected between those from ancestors feeding externally. Only two species lineages. Note the greater relative abundance of of criocerines have been reported as leaf-miners, alticine leaf-miners in contrast to the smaller rela- hence, proposing generalizations is out of the ques- tive abundance of leaf-mining galerucines within the tion. For the criocerines, Vencl & Aiello (1997) Galerucinae in the ‘Trichostomes’ (Jacoby) clade. hypothesize that endophyty is the basal condition. In the ‘cassidines’ (sensu Staines, 2002b, 2004b; This is based on the fact that endophyty is present Hispines and Cassidines, ‘Cryptostomes’ (Chapuis)) in the hypothesized outgroup of the Chrysomelidae, clade, leaf endophyty may have arisen independently including the Bruchidae and the Cerambycidae in both the Old and New World, from ancestors with (Crowson, 1981). Endophytous larvae neither larvae living between appressed leaves. In the sce- produce a larval shield (Vencl to Santiago-Blay, nario depicted in Figure 13, all subfamilies, except personal communication, May 2003), nor are they Hispinae, had most recent common ancestors with covered with a slimy protective mucilage (Crowson, exophytic, usually eruciform larvae. In the Hispinae, 1981). Larval shields and slimy covers, which are particularly those from the Old World, the fairly made out of fecal material, are produced only by depressed or onisciform larvae tend to live between exophytic larvae. Other presumed defenses of exo- appressed leaves of monocotyledoneous angio- phytic larvae include living in a hardened case (Cly- sperms. New World larval hispines have a greater trinae, Cryptocephalinae, and Chlamisinae), dorsal variety of body shapes and host-plant feeding pref- defensive glands (Chrysomelinae and some Gale- erences. rucinae), and lateral spines as well as excremental dorsal shields in Cassidinae sensu antiquo larvae, Origin and evolution of the leaf-mining habit among others. By forcing endophytic criocerine chrysomelids larvae that feed on stems to be exophytic, it can be tested whether they can produce a shield. The pro- Mines are relatively sealed from the outside, serv- duction of a shield in a normally endophytic larva ing as a locale for food and as a relatively buffered is considered strong evidence that this endophyty shelter, including from UV radiation (Connor & is a recently acquired condition, probably through Taverner, 1997). However, these cul-de-sacs where a reversal from a basal exophytic larva (Fig. 13). leaf-miners live limit the amount of food available Vencl (personal communication to Santiago-Blay, (Damman, 1994), area for waste disposal, and es- May 2003) has shown that all criocerine stem borers cape from predators and parasitoids. As a test of the tested produce shields, suggesting that their endo- possible adaptive significance of leaf-mining in the phyty is a reversed condition from an exophytic most Insecta done by Connor & Taverner (1997), the recent common ancestor. The alternative hypothesis multiple sister-group comparison method was used is that the endophytic larvae have retained the basal to assess whether leaf-mining has resulted in a greater characteristic and have independently evolved the diversification of leaf-mining lineages – a presumed capacity to produce a larval shield. In criocerines, surrogate of adaptive radiation. With the exception for example, leaf mining is considered a phyloge- of the Lepidoptera, where leaf-mining taxa are ex- netically reversed behavior. Perhaps more species ceedingly abundant, all other tested cases of sister of Oulema and of Lema are leaf-miners (Jolivet to taxa with one member being a predominant leaf- Santiago-Blay, personal communication, April 2003). miner had the leaf-mining taxon show a mediocre Vencl and Aiello, and many others (e.g., Kals- to low species richness. For instance, in the ‘Cryp- hoven, 1957) believe that endophyty, including leaf- tostomes’, or the clade formed by almost equally mining, is the basal condition for chrysomelids. speciose Hispinae and Cassidinae, leaf-mining is Schmitt (1988) hypothesized that the mining habit ubiquitous in the hispines yet almost absent in the may have been a retained synapomorphy with a cassidines. This suggests that leaf-mining, as well distant ancestor in common with the Hispinae but as leaf-galling for which parallel results have been he now considers that view very unlikely (Schmitt found, is an evolutionary ‘dead-end’ for most groups to Santiago-Blay, personal communication, April of insects (Connor & Taverner, 1997). However, the 2003). Leaf-mining chrysomelids 25

Fig. 13. Character state transitions for subfamilies of the Chrysomelidae containing leaf-mining taxa mapped into a recent phylogeny (data from Duckett et al., 2004). To simplify this hypothesis, the mapped trait, feeding mode, is being treated as a homologous character with two states: exo- and endophyty. The tick mark ‘—’ represents a synapomorphy for the subtended group, in the context of the Chrysomeloidea. A tick mark ‘—’ preceded by ‘–en’, (‘en’ means endophyty) represents a homoplasy (reversal) for the subtended group, such as tribes in the Cassidinae + Hispinae, etc. The small ovals with an associated ‘–en’ also represent a hypothetical reversal, in this case for a smaller subset, such as a few species in a genus or, less likely, all species in a genus. The number of ovals within the families or group of families is not in exact numerical scale. In the Zeugophorinae, leaf- mining appears to be the retained basal condition. Note the greater relative abundance of alticine leaf-miners, as indicated by the greater number of ‘-en’, in contrast to the smaller relative abundance of leaf-mining galerucines within the Galerucinae + Alticinae clade. In the ‘cassidines’ (sensu Staines 2002b, 2004b, Hispines + Cassidines or Cryptostomes (Chapuis)) clade, leaf endophyty may have arisen independently in both the Old and New World, from ancestors with larvae living between appressed leaves. Tribes of the Cassidinae + Hispinae having leaf-mining genera are parenthesized (and not placed inside small ovals). These include the Callohispini, Exothispini, Coelaenomenoderini, Promecothecini, Gonophorini, Oncocephalini, Hispini (Old World hispines) and the Prosopodontini, Sceloenoplini, Hispoleptini, Chalepini, Uroplatini (New World hispines). Within the Cassidines, only the Nothosacanthini has leaf mining taxa. The tribes Botryonopini, Anisoderini, Aproidini, Callispini, Leptispini, Eurispini, and Cryptonychini (Old World hispines) and the Cephaloleiini, Hybosispini, Arescini, and Alurnini (New World hispines), as well as the remaining tribes of the Cassidines, which are not leaf-miners, are omitted from the figure. While a basal division between Old and New World Cryptosomes has been indicated, this decision simply follows the traditional classification of many authors, including Seeno & Wilcox (1982). I am unaware of a comprehensive phylogeny for the group that would support this or any other global system for the Cryptosomes (Staines to Santiago-Blay, personal communication, June 2003), although there is a classifica- tion for part of the group (Borowiec, 1995). The placement of two or more taxa, represented by the smaller ovals, inside one of the larger ovals (more inclusive taxa) does not imply such less inclusive taxa are sister taxa or monophyletic. Details can be found in Evolutionary and biogeography trends.

Later in the evolution of Chrysomelidae, leaf- sheath, appressed, or rolled-leaf feeders, leaf-min- mining appears in one species of Galerucella and ers, and stem borers. Frost (1924) suggested that in many (probably all) species of Monoxia, (both hispine ancestors fed on materials in decay located genera are placed in the section Schematizites of the between the closely appressed leaves. Several spe- Galerucinae; Seeno & Wilcox, 1982). Some of the cies of Prosopodonta live between or mine closely chenopod-feeding Monoxia larvae seem to bore and appressed leaves (Maulik, 1931.) Likewise, larvae live in the unopened flower buds or fruits. Several of Gyllenhaleus spp., Cryptonychus spp., and oth- unrelated alticines are leaf-miners and in many of ers, feed on unopened leaf buds of the African and them the congeneric species are not miners. For Central American plant genus Costus spp. (Costa- example, as larvae, most species of Psylliodes are ceae) and on the African plant genus Amomum (Zin- root feeders. However, the pestiferous species P. giberaceae), respectively, penetrating at a later time chrysocephala feeds on both the petiole and the blade into the stem (Collart, 1928; Maulik, 1932; Spaeth, of crucifer leaves and later on the harder parts of 1933; Staines, 2004b). Crowson (1955) believes that the shoots (Grison et al., 1963). two ecological lineages of hispines evolved from The Hispinae + Cassidinae (or Cassidinae, sensu ancestors living between closely appressed leaves: lato of Staines 2002b, 2004b) are divided into four one lineage with free living larvae and the other with functional feeding groups: free-living leaf feeders, leaf-miners. 26 J.A. Santiago-Blay

that

Fig. 14. Hypothetical feeding ecology transitions leading to leaf-mining in the Chrysomelidae, with examples.

Sceloenopla af. bidens larvae are typically found noses based on morphological studies (Rowe, 1988), on shoot-leaf junctions (‘axils’), as well as on min- can be documented. For the time being, I have as- ing leaves (Costa et al., 1988), suggesting external sumed that the inferences concerning their species foliovory as a possible evolutionary pathway to leaf- status are correct. mining. This is not surprising since Hering (1951) Another striking aspect of leaf-mining hispines described cases of leaf-mining insects that eat their of the Indo-Pacific region is their speciosity in a few way through a stem in moving from leaf to leaf. Other host-plant families (Arecaceae, Pandanaceae, and insect miners move to non-leafy parts of the shoot. Zingiberaceae), genera, or species (c.f. Table 1). This is because leaves may be too small or atrophied Gressitt (1957, 1963) and Monteith (1970) pointed in those plant species for the insects to complete, this out many years ago. Gressitt suggested that or even undergo, development (Hering, 1951). speciosity was related to: 1. geographical isolation Figure 14 summarizes the hypothetical ecologi- (allopatric speciation model, Mayr, 1970); 2. low cal transitions, from a browsing to an obligatory leaf- population numbers (low effective population size mining larva. (Ne), random genetic drift, shifting balance and their impact on speciation and evolution, Wright, 1968- Why is leaf-mining absent in most of the 1978); 3. changing environment (and possible se- Chrysomelidae? lection, Darwin, 1859; Fisher, 1999); and 4. ongoing genetic recombination due to rejoining of formerly With exception of hispines, leaf-mining is relatively isolated populations (e.g., New Guinea). Gressitt uncommon in the Chrysomelidae. A rapid perusal (1957) noted also that in “many New Guinea insect of Table 1 shows the tremendous speciosity of leaf- groups… [there are] one or two widespread forms, mining hispines in the Indo-Pacific region. These with other species differentiated in montane areas”. groups need to be studied carefully, including the Hawkeswood & Takizawa (1997) suggest that the use of experimental approaches, such as mating colder climate of Australia, not the lack of suitable experiments which are so commonly done for labo- hosts, is largely responsible for the relative paucity ratory-reared Drosophila (Diptera), so that their of hispines in Australia compared to neighboring genealogical relatedness, often inferred from diag- New Guinea. Gressitt (1957) discusses the bioge- Leaf-mining chrysomelids 27 ography of hispines in numerous landmasses of the 19th century. Among the seventeen leaf-mining insect Pacific Basin. He suggests that because hispines tend species found, three were beetles, one of which was toward monophagy and are mediocre flyers, most the hispine, Baliosus nervosus. The isolation of small of their diversity in this region is caused by vicariant Quercus host trees decreased the susceptibility of instead of dispersal events. leaf-mining insects to parasitoidism (Faeth & Sim- Numerous species of leaf-mining chrysomelids, berloff, 1981). However, that decrease was not fol- sometimes congenerics, occur sympatrically, even lowed by population increases because on small in the same host-plant species (Riley & Enns, 1979). isolated Quercus trees leaf-mining insects are re- Except for the case of Odontota mundulus (Sander- cruited from neighboring host-plants. Although in son) and O. scapularis (Olivier), there are no ob- the studies of Auberbach & Simberloff (1984, 1988) servations on how often reproduction isolation is B. nervosus was one of the rare taxa, those authors broken down in heterospecifics. In the case of the concluded that similar relative diversities of leaf- congeneric species of Odontota, Riley & Enns (1979) miners on host plants are partially determined by report mating between O. mundulus and O. sca- the presence of taxonomically-related host plants in pularis, but no hybrids that could be recognized the neighborhood. Those plants supply both new leaf- externally have been found. The genetic and other mining recruits, as well as natural biological enemies biological correlates of these patterns are unknown. (Faeth & Simberloff, 1981; Faeth et al., 1981). In addition, other factors, such as the biology of the Biogeographical patterns leaf-miner species and abiotic factors, are also im- portant correlates of population abundance and di- Leaf-mining chrysomelids are distributed worldwide. versity. At the broadest scale, leaf-mining chrysomelids tend After a multi-year study of insects colonizing to be more speciose in the tropics, undoubtedly due Polygonum perfoliatum Linné, Wheeler & Mengel to the presence of the mostly tropical hispines in those (1984) concluded that polyphagous insects are the latitudinal ranges, particularly the Oriental biogeo- first to colonize a plant that is new to an area. There- graphic region (Anand, 1984; Gressitt et al., 1961). after, oligophagous insects of taxonomically-related This pattern is also followed by leaf-mining Bu- plants colonize the new host-plant. Parasitoids ap- prestidae (Hespenheide, 1991). Perhaps the only pear to be tracking host plants closer than the in- exception to this is the apparent scarcity of leaf- sects they parasitize (Auberbach & Simberloff, 1984, mining forms in australotropical zones, but this may 1988; Hespenheide, 1991; etc.). be caused by the lack of extensive surveys of South America and Africa. Evidence that Wilf, Cúneo, and Labandeira are currently garnering suggests that the Economic importance lineages of Neotropical leaf-miners are ancient, as are the forests upon whose foliage they fed, and that As a group, leaf-mining chrysomelids vary in their those lineages extend, minimally, to the late Pale- economic importance as herbivorous biocontrol ocene-Early Eocene (circa 55 Ma). In addition, the agents of weeds or as pests of important crops. Neotropical site being explored by them, located in Argentina, is the single deposit with the greatest Leaf-mining chrysomelids as agents of weed diversity and number of leaf-mines in the fossil biocontrol record, with the possible exception of the Dakota Formation (mid Cretaceous, ;100 Ma) of Nebraska The relative success of some chrysomelids in con- and Kansas (Labandeira to Santiago-Blay, personal trolling weeds (DeBach & Rosen, 1991; Goeden & communication, July 2003). Andrés, 1999) has alerted students of this family to At the scale of the ecosystem, in the discussion the possibility of using host specific leaf-mining of the distribution of leaf-mining buprestids in and chrysomelids to attempt to control some weeds. For Guanacaste (both in Costa Rica, Central America), example, Octotoma scabripennis, Uroplata girardi, Hespenheide (1994) suggests that both historic bio- and a few other hispines have been used success- geography and recent climatic conditions explain the fully for biocontrol of the weed, Lantana camara higher diversity in leaf-miners in the more stable and (Verbenaceae) (Cilliers, 1977, 1983, 1987a,b; Harley, southern (closer to South America) tropical lowland 1969; Staines, 1989; Tucker & Singh, 1993; Winder rainforest, La Selva (Costa Rica), in contrast to the & Harley, 1982; Winder et al., 1984). Although highly seasonal and northern tropical lowland dry Winder & Harley (1982) gave a relatively low weight forest, Guanacaste, also in Costa Rica. to leaf-mining as an attack type on species of Lan- At the scale of individual plants, Janzen (1968) tana, a combination of factors have brought this weed suggested that host plants are analogous to the real under control in some parts of the world. More re- islands of the island biogeography theory. Several cently, Broughton (2000) critically reviewed the studies have discussed biogeographical patterns of literature on the biocontrol of L. camara and con- mining insects on species of oak (Quercus). The cluded that the hispine Uroplata girardi is the most species of oak studied were introduced to northern successful biocontrol agent. Florida (United States) during the second half of the There can be drawbacks to relying on a limited 28 J.A. Santiago-Blay number of biocontrol agents. In this case, general- it is important to consider the effect that biocontrol ist predators, including spiders, predatory hetero- agents may have on economically-important crops pterans, neuropteran larvae, and ants attack U. girardi (Hilgendorf & Goeden, 1981). and O. scabripennis in several parts of the world. Also, low temperatures negatively affect these two Leaf-mining chrysomelids as pests biocontrol agents, hence their effectiveness is some- what reduced because their populations are lowered About a dozen species of leaf-mining chrysomelids, during the winter. Although a cool-adapted biotype particularly hispines (Anand, 1984; Maulik, 1919), of U. girardi has been introduced to Australia, no are very important economically. The damage they data appear to be available on their relative estab- inflict consists mainly of eating away leaf tissue. In lishment success (Broughton, 2000). Consequently, the case of rice, two species of leaf-mining hispines additional efforts have been undertaken to identify have been implicated in the transmission of a phy- multiple host-specific, compatible biocontrol agents topathogenic virus. The cultivars affected and their of weeds, including herbivores, such as leaf-min- pestiferous leaf-mining chrysomelids are briefly dis- ing chrysomelids and pathogens (Harley et al., 1995; cussed below. Gillett et al., 1991; Goeden & Ricker 1974, 1975, 1976a,b,c; Wheeler & Mengel, 1984). Regrettably, Palms (Arecaceae) the biology of numerous, non-economically im- In some cases, attack by leaf-miners is devastating portant insects remains unknown, even if they are to the host-plants, such as oil palms, Elaeis guineen- included in studies on their potential economic sig- sis, and coconut palms, Cocos nucifera (Bernon & nificance (Goeden & Teerink, 1993). Graves, 1979; Dharmadhikari et al., 1977; Mariau, According to Tucker & Singh (1993), “A num- 1988). For instance, the sometimes cyclical outbreaks ber of leaf-mining beetles (Chrysomelidae) have been of Coelaenomenodera elaeidis Maulik and of nu- used successfully in Australia and Hawaii in con- merous other palm hispines reduce foliage and oil trolling lantana, Lantana camara, and could be in- production of the oil palm, Elaeis guineensis (Bernon troduced into Florida. However, such introductions & Graves, 1979; Lecoustre et al., 1980) and of the are frequently delayed or denied by various federal coconut palm, Cocos nucifera Linné (Howard et al., or state committees that must evaluate the risk-benefit 2001; Mariau, 2001, 2004). Lepesme (1948) and picture. There has been some opposition to the in- Howard et al. (2001) review insects on palms world- troduction of biological control agents due to the fear wide. that the introduced insects or pathogens will attack Foreign exploration-importation efforts have taken other plants once their primary food source has been place to control pestiferous leaf-mining chrysomelids consumed. Also, a weed of economic importance to (Cochereau, 1972). Two relatively successful cases one may be a desirable plant of value to others. In are the control of the coconut leaf-mining hispines, resolving such conflicts, the economic impact of the Promecotheca spp. and Brontispa longissima Gestro. weed in croplands must be compared with the nega- In both these cases, imported eulophids have con- tive aspects indicated by those who oppose its con- tributed to the relative success of the foreign impor- trol by such introductions.” tation of natural biological control agents (Cochereau, Many European herbaceous crucifers are attacked 1972). Biological control of Coelaenomenodera ela- by the alticine Phyllotreta nemorum Linné in early eidis was attempted in West Africa by introducing growth stages. This causes such damage to the plants a Malagasy larval parasitoid (Chrysonotomyia sp.) that they cannot recover at later growth stages (He- of a congeneric hispine. These efforts failed to control ring, 1951). However, interest in using P. nemorum hispine populations in the wild since it appears that and other insect herbivores to control cruciferous the parasitoid could not adapt to the fact that C. weeds has continued (Lipa et al., 1977). Sometimes, elaeidis larvae die right after being parasitized, re- trees considered ‘less desirable’, such as Ostrya ducing the availability as a food source for the in- virginiana (Miller) K. Koch., are attacked by Baliosus ternally developing parasitoid larvae (Mariau, 1988). nervosus hispines sparing the more desirable host Similar relative failure stories in long term biologi- plant (and ornamental tree) Tilia americana of some cal control hold true for several species of Prome- of its ravages. cotheca spp. and for Hispolepsis subfasciata (Mariau, In contrast, many leaf-mining chrysomelids have 1988). minimal impact on plant populations (Hespenheide, Augmentation of native parasitoids has also been 1991; Hespenheide & Dang, 1999). For example, considered to control C. elaeidis. It has been noted while at least ten species of Dibolia alticines are leaf- that C. elaeidis and another hispine, Platypria co- miners in Europe, their economic impact seems mi- ronata, a leaf-miner of a legume cover crop in oil nimal (Grison et al., 1963). Likewise, a species of palm plantations, share some parasitoids (Bernon & Monoxia, probably M. grisea, was found in densi- Graves, 1979). Perhaps one of the best-known cases ties of up to 100-500 in 3 × 3 × 3-feet plots of Ar- of biological control is that involving the control of temisia tridentata Nutall (Asteraceae) plants in 1961 Promecotheca caerulipennis by the mite Pyemotes but, “none of the plants that were heavily attacked ventricosus (Newport) in Fiji (Taylor, 1937). Other in 1961 showed adverse effects in 1962”. When cases of mite-chrysomelid associations are summa- biological control of weeds is being contemplated, rized in Santiago-Blay & Fain (1994). Leaf-mining chrysomelids 29

Rice, Oryza sp. (Poaceae) Collecting and rearing leaf-mining chrysomelids Dicladispa armigera (Olivier) is a pest of rice (Rawat & Singh, 1980; Razzaque & Kari, 1989; Sen & As long as the host plants are known, the ease of Chakravorty, 1970) and can cause severe damage collection and rearing of leaf-mining insects, while to this crop. Several species of hispines, including in the mines, has been repeatedly noted (e.g., Ford Dicladispa sp., Trichispa sericea Guérin-Méneville, & Cavey, 1985; Hespenheide, 1991; Hering, 1951; and the exophytic galerucine Sesselia pussila (Ger- Kato, 1991; Lee & Furth, 2000). However, at times, staecker) are vectors of the rice yellow mottle virus it is difficult to find adults or mines in large num- (RYMV) in Africa (Banwo et al., 2001a,b). Other bers. As collecting, rearing, and associating life stages chrysomelids reported as vectoring viruses are listed are achieved, future studies on leaf-mining chrysome- in Crowson (1981). lids should concentrate on answering some potential- ly interesting ecological and evolutionary questions. Soybean, Glycine max Linné (Leguminosae) If there is a lack of environmentally-controlled The leaf-mining hispines, Odontota horni and Ba- facilities, plant cuttings can simply be placed in a liosus nervosus, have been suggested as potential container (e.g., a plastic bag containing a piece of pests of soybean, Glycyne max, particularly if they wet cotton suffices, provided with regular air cir- are present in conjunction with other crop pests culation, or a tightly sealed wire mesh) away from (Buntin & Pedigo, 1982; McPherson & Ravlin, 1983: direct sunlight and low temperatures (Gressitt, 1959). Wheeler & Stimmel, 1983). A multiherbivore attack, Placing cuttings in a container with water or trans- including a major pest, such as the leaf-mining dip- planting whole plants, while keeping everything in teran Liriomyza trifolii (Burgess), and unnamed chry- enclosures, is another easy method to rear leaf-mining somelids, can exacerbate risks of yield reductions insects. Sealed containers (bags, cups) with adequate (e.g., Phaseolus vulgaris Linné in Cuba, Heyer et ventilation not only allow collection of the emerg- al., 1989). ing adults but also of parasitoids. See Ford & Cavey (1985) for additional details. Other cultivars Mass-rearing of leaf-mining chrysomelids and of Adult alticines, such as Sphaeroderma rubidum, their natural enemies has been improved by ‘experi- have been reported as a pest of artichoke (Cynara mental minology’, the use of ‘artificial mines’ (Gal- cardunculus Linné) in the western Mediterranean lego et al., 1983; Hering, 1951). This technique has basin (Grison et al., 1963). Chaetocnema tibialis been used for at least a century (Hering, 1951) by Illiger, a flea beetle, is a pest of sugar beets in some many, including by Mariau in Africa. Artificial mines parts of Europe (Zabel et al., 1991). Major infesta- are created by inserting a long and thin object, such tions of Citrus spp. by alticine, Throscoryssa citri, as a needle or knife, inside a leaf, and forming a larvae may cause severe defoliations (Zaka-ur-Rab, cavity that mimics a mine. Some species, such as 1991). Several species of Dactylispa hispines are the hispine Promecotheca cumingii seem to accept considered pestiferous in cichona (probably Cinchona these human-created dwellings. However, the authors officinales Linné, Rubiaceae), kapok (probably Ceiba admit that “constant practice is needed to perfect the sp., Bombacaceae), coffee (Coffea arabica, Rubia- procedure”. ceae), and maize (Zea mays, Poaceae) in southeast- ern Asia and eastern Africa (An et al., 1985; De & Konar, 1954). Acknowledgments

Ornamental trees I am profoundly indebted to the ‘learning commu- The hispine Odontota dorsalis has been reported to nity’ which helped me during the preparation of this be a major pest of Tilia americana trees in Wash- paper. Dr Charlie L. Staines (Edgewater, MD) his ington, DC (Chittenden, 1902). At that time, chemical for his critically constructive reading of several it- control measures, some of which would be consid- erations of this paper, starting in the mid-1990s when ered unacceptable by today’s standards, were rec- I failed to complete it for Volumes 3-5 of this se- ommended. Odontota dorsalis attacks on Robinia ries. At that time, the entire manuscript was only pseudoacacia are so severe that, “leaves are turned 10-15% of its current length. Conrad C. Labandeira, brown as if scorched by fire” (Needham et al., 1928). Pierre Jolivet, and Michael Schmitt, the latter two Numerous hispines attack palms, many of which are co-editors of the current volumes, reviewed this paper becoming widespread because they are used as or- and made numerous suggestions. Also, through his namental trees. Some palms have become invasive – often hilarious - emails, Jolivet contributed many (Svenning, 2002) and there is no research yet on how of the examples used in this paper, as well as some their leaf-mining complex would change as they are of the history of studies on insect-plant interactions, introduced into new localities. including a never-published paper by S. Maulik (1945). I am fully aware that Table 1 is not com- prehensive. For example, there are chrysomelids in New Zealand that seem to be miners, but there are no biological data on them (Jolivet to Santiago-Blay, 30 J.A. Santiago-Blay personal communication, June 2003). However, I software I used for the diagrams herein included. have decided to “keep to an imperfect solution” and April Pumala (Pennsylvania State University, Great hope that more additions and corrections can be made Valley, Malvern, PA) retyped the original version to this work by other workers. of Table 1, whose electronic version I could not Numerous other colleagues and friends helped in locate. Fortunately for her, at the point Ms Pumala the preparation of this chapter. Krishna K. Verma re-entered it, Table 1 was only about one-third of (Durg, Chattisgargh, India) reviewed the section on the length it is now. Shane Guan (United States Fish Zeugophorinae and made recommendations. Karoly and Wildlife Service, Washington, DC), Rudolf Fury, Vig (Savaria Múzeum, Kisfaludy, Hungary) offered Professor Junfeng Zhang (Najing Institute of Geol- numerous suggestions and literature pertaining to ogy and Paleontology, People’s Republic of China), alticines, particularly species of Phyllotreta. I am Angela Stavropoulos, and a translator of Chinese indebted to Vig for letting me paraphrase a large documents who prefers to remain anonymous, aptly portion of his 1999 paper in the section on Chemi- and promptly translated some of the papers (or parts cal correlates of feeding behavior. Michael Cox of them) written in languages I cannot read. With (Department of Entomology, The Natural History the exception of Professor Zhang, the translators Museum, London) reviewed the section on Abbre- were located through the generous help of Ms viated diagnoses of pupae of leaf-mining chrysome- Angelia Bland and Ms Amy Lemon (Behind-The- lids. Henry Hespenheide (Department of Biology, Scenes Volunteer Program of the Visitor Informa- University of California, Los Angeles, CA) and tion and Associates’ Reception Center (VIARC) at Vincent Resh (Department of Environmental Sci- the Smithsonian Institution). Alison L. Armocida ences, Policy, and Management, University of Cali- (University of Maryland, Baltimore County, Balti- fornia, Berkeley, CA) commented on evolutionary more, MD), William Bell (Tulane University, New aspects of leaf-mining on insects. Kevin Padian Orleans, LA), Gloria Friar (Program for Deaf and (Department of Integrative Biology and Museum of Hard of Hearing of Prince George’s County, MD), Paleontology, University of California, Berkeley, Lester Guthrie (Montgomery College, Rockville, CA) provided me with pertinent literature on parsi- MD), Katie Kan (William H. Hall High School, West mony and homology. Donald Kaplan (Department Hartford, CT), Joyce Meng (Thomas Jefferson High of Plant and Microbial Biology, University of Cali- School for Science and Technology, Alexandria, fornia, Berkeley, CA) provided insightful views on VA), Bethany Sadlowski (Arlington, VA), Adam Toy the concept of homology. The number of leaf-min- (Buffalo State College, Buffalo, NY), and Sarah A. ing rhynchitids (twelve) is courtesy of Robert Hamil- Wyatt (College of William and Mary, Williamsburg, ton (Loyola University, Chicago, IL). Dr. Warren VA) helped me with the bibliographical, editing, and/ Allmon (Paleontological Research Institution/Mu- or proof-reading aspects of this paper. I welcome seum of the Earth, Ithaca, New York, USA) com- additions and corrections to this database. All mis- mented on the practice of discarding plant fossils takes are my responsibility. not showing diagnozable traits, even if they have evidence of herbivory. Staff members at the Bio- Sciences Library of the University of California at References Berkeley (particularly Nancy Axelrod, Norma Kob- zina, and Ingrid Ratcliff) and at the National Museum References that could not be located or translated by the time of Natural History in Washington, DC (particularly of printing are indicated by an asterisk (*). Richard C. Greene, David T. Steere, Jr., Claire Cla- tron, Martha A. Rosen, Robert J. Skarr, Courtney Abdullah, M. and Qureshi, S.S. 1969. A key to the Pakistani A. Shaw, and Ruth F. Schallert) made it possible to genera and species of Hispinae and Cassidinae (Coleoptera: find numerous citations, including most of the ob- Chrysomelidae), with description of new species from West scure ones. Dr Dan Nicolson (Department of Sys- Pakistan including economic importance. Pakistan Journal of Scientific and Industrial Research, 12:95-104. tematic Biology, Division of Botany, National An, S.L., Kwon, Y.J. and Lee, S.-M. 1985. 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A bibliography of New Staines, C.L. 1986a. A revision of the genus Brachycoryna World Hispinae (Coleoptera: Chrysomelidae): Addenda. Mary- (Coleoptera: Chrysomelidae: Hispinae). Insecta Mundi, 1:231- land Entomologist, 3:147-151. 241. Steinhausen, W.R. 1966. Vergleichende Morphologie des La- Staines, C.L. 1986b. New combination and new synonymy in brum von Blattkäferlarven. Deutsche Entomologische Zeit- North American Stenopodius (Coleoptera: Chrysomelidae: schrift (Berlin) N.F., 13:31.3-322. Hispinae) with a taxonomic note on Uroplatini. Proceedings Steinhausen, W.R. 1978. Bestimmungstabelle fur die Larven der of the Entomological Society of Washington, 88:192. Chrysomelidae (partim). In: Klausnitzer, B. (ed) Ordnung Staines, C.L. 1989. A revision of the genus Octotoma (Coleoptera: Coleoptera (Larven). The Hague, Dr. W. Junk Publishers. pp. Chrysomelidae, Hispinae). Insecta Mundi, 3:41-56. 336-343. Staines, C.L. 1991. Generic reassignment of Anisostena cham- Stevenson, J. 1992. 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A new species of leaf-mining University Press. 370 pp. Oulema from Panama (Coleoptera: Chrysomelidae; Crio- Tutin, T.G., Heywood, V.H., Burges, N.A., Moore, D.M., Val- cerinae). Journal of the New York Entomological Society, entine, D.H., Walters, S.M. and Webb, D.A., with the assis- 105:40-44. tance of Chatter, A.O., DeFilips, R.A. and Richardson, I.B.K. Vencl, F.V., Levy, A., Geeta, R., Keller, G. and Windsor, D.M. (eds) 1976. Flora Europaea. Vol 4. Plantaginaceae to Com- 2004. Observations on the natural history, systematics and positae (and Rubiaceae). Cambridge, Cambridge University phylogeny of the Criocerinae of Costa Rica and Panama. In: Press. 505 pp. Jolivet, P., Santiago-Blay, J.A. and Schmitt, M. (eds) New Tutin, T.G., Heywood, V.H., Burges, N.A., Moore, D.M., Val- Developments in the Biology of Chrysomelidae. The Hague, entine, D.H., Walters, S.M. and Webb, D.A., with the assis- SPB Academic Publishing. pp. 423-454. tance of Chatter, A.O. and Richardson, I.B.K. (eds) 1980. *Vig, K. 1989. Kártevo´´ földibolha fajok (Phyllotretra spp.) Flora Europaea. Vol 5. Alismataceae to Orchidaceae (Mono- rendszertana, alaktana és életmódja (Coleoptera, Chryso- cotyledones). Cambridge, Cambridge University Press. 452 melidae, Alticinae) [Taxonomy, morphology and biology of pp. Phyllotreta pests]. Unpublished PhD Thesis, Keszthely–Szom- Uhmann, E. 1931. Die Hispinen des Musée du Congo Belge. 3. bathely. (In Hungarian) Teil. Oncocephalini, Exothispini, Coelaenomenoderini, Gono- Vig, K. 1998a. Data to the biology of Phyllotreta vittula (Redten- phorini, Cryptonychini, Hispini ohne die Gattung Hispa. 33. bacher, 1849) (Coleoptera: Chrysomelidae: Alticinae). Mede- Beitrag zur Kenntnis der Hispinen (Col. Chrys.). Revue de delingen van de Faculteit Landbouwkundige en Toegepaste Zoologie et Botanique Africaines, 21:74-88. Biologische Wetenschappen Universiteit Gent, 63:357-363. Uhmann, E. 1934. Hispinen-Minen aus Costa Rica. 48. Beitrag Vig, K. 1998b. Host plant selection by Phyllotreta vittula (Redten- zur Kenntnis der Hispinen (Col: Chrysomelidae). Arbeiten bacher, 1949). In: Biondi, M., Daccordi, M. and Furth, D.G. über physiologische und angewandte Entomologie aus Ber- (eds) Proceedings of the Fourth International Symposium on lin-Dahlem, 1:272-277. the Chrysomelidae. 20th International Congress of Entomol- Leaf-mining chrysomelids 41

ogy. Turin, Museo Regionale di Scienze Naturali. pp. 233- Wilcox, J.A. 1979. Leaf Beetle Host Plants in Northeastern North 351. America (Coleoptera: Chrysomelidae). Biological Research *Vig, K. 1999. Biology of the horse-radish flea beetle, Phyllotreta Institute of America, Inc. Kinderhook, World Natural History armoraciae Koch (Coleoptera: Chrysomelidae) in Hungary, Publications. 30 pp. Central Europe. In: Sobti, R.C. and Yadav, J.S. (ed) Some Wilcox, J.A. (no date) a. Zeugophorinae. Chrysomelidae. Re- Aspects on the Insight of Insect Biology. Chandigarh, Panjab gion I. Northeastern United States and Canada. The North University. pp. 5-20. American Beetle Fauna Project. Biological Research Insti- Vig, K. 2000. Data on the biology of the turnip flea beetle, tute of America. No pagination. Phyllotreta nemorum (Linnaeus, 1758) (Coleoptera, Chryso- Wilf, P., Labandeira, C.C., Johnson, K.R., Coley, P.D. and Cutter, melidae, Alticinae). Mededelingen van de Faculteit Land- A.D. 2001. Insect herbivory, plant defense, and early Ceno- bouwkundige en Toegepaste Biologische Wetenschappen zoic climate change. Proceedings of the National Academy Universiteit Gent, 65:201–212. of Sciences USA, 98:6221-6226. Vig, K. 2003. Biology of Phyllotreta species. This Volume. Wilf, P., Labandeira, C.C., Kress, W. J., Staines, C.L., Windsor, Vig, K. and Verdyck, P. 2001. Data on the host plant selection D.M., Allen, A.L. and Johnson, K.R. 2000. Timing of the of the horseradish flea beetle, Phyllotreta armoraciae (Koch, radiations of leaf beetles: hispines on gingers from the latest 1803) (Coleoptera, Chrysomelidae, Alticinae). Mededelingen Cretaceous to Recent. Science, 289:291-294. van de Faculteit Landbouwkundige en Toegepaste Biologische Wilf, P., Labandeira, C. and Miljour, B. 2001. Field guide to Wetenschappen Universiteit Gent, 66:277–283. insect damage types on compressed fossil leaves. Version 0.0. Virkki, N. and J. A. Santiago-Blay. 1998. Chromosome num- Template courtesy of M. Metz and R. Foster. The Field bers in 71 Puerto Rican species of leaf beetles (Coleoptera: Museum. Chicago. 4 sheets. Chrysomelidae). Journal of Agriculture of the University of Williams, C.E. 1989a. Damage to woody plants by the locust Puerto Rico, 82:69-83. leafminer, Odontota dorsalis (Coleoptera: Chrysomelidae), W3T Tropicos (Missouri Botanical Garden, St Louis, MO). http:/ during a local outbreak in an Appalachian oak forest. Ento- /mobot.mobot.org/W3T/search/vast.html mological News, 100:183-187. Wade, J.S. 1935. A contribution to a bibliography of the de- Williams, C.E. 1989b. Host plants of Microrhopala xerene scribed immature stages of North American Coleoptera: cereal (Newman) (Coleoptera: Chrysomelidae) in southwestern Vir- and forage insect investigations. Bureau of Entomology and ginia. The Coleopterists’ Bulletin, 43:391-392. Plant Quarantine. United States Department of Agriculture, Williams, C.E. 1991. New England Aster, Aster novae-angliae: Washington, DC, USA. E-358. 114 pp. a new host record for Microrhopala xerene (Coleoptera Chry- Ward, C.R., O’Brien, C.W.O., O’Brien, L.B., Forster, D.E. and somelidae). Proceedings of the Entomological Society of Huddleston, E.W. 1977. Annotated checklist of New World Washington, 93:790. insects associated with Prosopis (Mesquite). Technical Bul- Winder, J.A. and Harley, K.L.S. 1982. The effects of natural letin 1557. Agricultural Research Service. United States Department of Agriculture. 115 pp. enemies on the growth of Lantana in Brazil. Bulletin of West, A.S. and Lothian, T.M. 1948. The basswood leafminer, Entomological Research, 72:599-616. Baliosus ruber (Weber), (Chrysomelidae, Hispini), in the Winder, J.A., Harley, K.L.S., and Kassulke, R.C. 1984. Uroplata Rideau Lakes Region. 78th Report of the Entomological lantanae Buzzi & Winder (Coleoptera: Chrysomelidae: His- Society of Ontario. pp. 62-65. pinae), a potential biological control agent of Lantana camara) West, C. 1985. Factors underlying the late seasonal appearance in Australia. Bulletin of Entomological Research, 74:327-340. of the lepidopterous leaf-mining guild on oak. Ecological Wright, S. 1968-1978. Evolution and the Genetics of Popula- Entomology, 10:111-120. tions. Vol 1 (1968). Genetic and Biometric Foundations, 470 Wheeler, A.G. 1980. Japanese pagodatree: a host of locust leaf- pp. Vol 2 (1969). Theory of Gene Frequencies, 512 pp. Vol miner, Odontota dorsalis (Thunberg) (Coleoptera: Chryso- 3 (1977). Experimental Results and Evolutionary Deductions, melidae). The Coleopterists’ Bulletin, 39:95-98. 614 pp. Vol 4 (1978). Variability Within and Among Natu- Wheeler, A.G. 1987. Locust leafminer, Odontota dorsalis (Thun- ral Populations. Chicago, University of Chicago Press. 580 berg) Coleoptera: Chrysomelidae. Regulatory Horticulture pp. Entomology (Pennsylvania Department of Agriculture. Bu- Würmli, M. 1975. Gattungmonographie der altweltlichen His- reau of Plant Industry) Circular No. 115, 13:15-17. pinen (Coleoptera: Chrysomelidae: Hispinae). Entomologische Wheeler, A.G. and Mengel, S.M. 1984. Phytophagous insect fauna Arbeiten aus dem Museum G. Frey 26:1-83. of Polygonum perfoliatum, an Asiatic weed recently intro- Yu, P.Y. 1993. Coleoptera: Hispidae - Anisoderinae, Callispinae, duced to Pennsylvania. Annals of the Entomological Soci- Cassidinae. In: The Series of the bioresources expedition to ety of America, 77:197-202. the Longqi Mountain Nature Reserve: Animals of Longqi Wheeler, A.G. and Snook, W.A. 1986. Biology of Sumitrosis Mountain. Beijing, China Forestry Publishing House. pp. 374- rosea (Coleoptera: Chrysomelidae), a leafminer of black 379. (In Mandarin) locust, Robinia pseudoacacia (Leguminosae). Proceedings of Zabel, A., Kostiç, M. and Manojloviç, B. 1991. The possibility the Entomological Society of Washington, 88:521-530. of chemical control of Chaetocnema tibialis Illig. (Coleoptera: Wheeler, A.G. and Stimmel, J.F. 1983. The phytophagous and Halticinae) on sugar beet. Zaštita Bilja (Plant Protection, predacious arthropod fauna of soybean in Pennsylvania. Mels- Institute for Plant Protection and Environment. Belgrade, heimer Entomological Series, 33:31-38. Serbia), 42:337-344. Wilcox, J.A. 1954. Leaf beetles of Ohio (Chrysomelidae: Co- Zaka-ur-Rab, M. 1991. Leaf-mining Coleoptera of the Indian leoptera). The Ohio State University Vol 8, No. 3. Bulletin subcontinent. Journal of Entomological Research, 15:20-30. 43. pp. 353-506. Zheng, Y., Harman, D.M. and Swartz, H.J. 2003. Resistance to Wilcox, J.A. 1975. Family 129. Chrysomelidae: Checklist of the locust leafminer (Coleoptera: Chrysomelidae) in black locust. Beetles of North and Central America and the West Indies. Journal of Economic Entomology, 96:53-57. Compiled and edited by Ross H. Arnett, Jr. Vol 8. The Leaf Zherikhin, V.V. 2002. Insect trace fossils. In: Rasnitsyn, A.P. Beetles and the Bean Weevils. Gainesville, Flora and Fauna and Quickle, D.L.J. (eds) History of Insects, pp. 303-324. Publications. 166 pp. Dordrecht, Kluwer Academic Publishers. 42 J.A. Santiago-Blay and others, bore the internodes of bamboos Lasiochila /names.html; Plants Database (United States Department of (2000); Germplasm Resources Information Network (GRIN) l distribution. The geographical distribution given is the maximum are given only to genera. Common names avoided as much et al. somelidae of the world. This list is not exhaustive. Host plant data r r, in several cases, there are different views about the nomenclatural ars-grin.gov/npgs/tax; Halladay & Beadle (1983); Harvard University ni, Promecothecini, Gonophorini, Oncocephalini, Hispini (Old World they are only temporary herbivores on the buds, and larger larval scini, and Alurnini (of the New World Hispinae) Botryonopini, e to assume leaf-mining for all genera in the tribes Prosopodontini, to find published host plant association data. The suffix ‘-ceae’ is used modern names for the geographical regions of world, particularly in the tribe Cryptonychini), it appears that some of species can, on os (Missouri Botanical Garden, St Louis, MO) http://mobot.mobot.org/ e), Lamiaceae (= Labiatae), and Poaceae Gramineae). I have retained nts where these insects mined from those larvae are exophytous. n leaf-mining hispines live between the appressed or in rolled leaves of (1998) was used to complete the information for authors of chrysomelid -Blay, personal communications, April 2003). Beetle and plant synonymies (Maulik, 1932), ailed analysis of the data is in progress. d the most up-to-date name for each taxon by using revisionary works and en noted with the phrase ‘unable to find name’. As much as possible, host to facilitate location for non-specialists. In the Hispinae, genera are listed those cases, the notation ‘larvae’ (implying that adult also feeds on d (and recommend readers to do the same) that host plant records are, as far the taxon’ are leaf miners’, I have entered a taxon as miner if references . It is important to recall that, in general, host plant ranges of adults are broader et al. Estigmena e.g., Plesispa sp.) as I was unable to find more detailed data. Members of the Orsodacninae and Genus’ http://www.uk.ipni.org/index.html or http://www.us.ipni.org/index.html; Krüssmann (1984); Mabberly 1 ) and of some hispine genera ( (1961), as well the website, http://www.infoplease.com/countries.html, proved very useful for that task. Staines (2003b) Phyllotreta , et al. Epitrix

e.g., (1964, 1968, 1972, 1976, 1980); IPNI (International Plant Names Index) et al. Uhmann, 1957, 1958b, 1964). To save space, I have omitted subgeneric and subspecific ranks. Schmitt e.g., . Taxa, geographical distribution, host plants (arranged alphabetically by family), and selected references of leaf-mining Chry catalogues ( names. Except for hispines, the scientific names of leaf beetles are listed alphabetically, by genus, within subfamily in order Table 1 Scientific names and authors of species are given as completely updated I could. In all cases, an effort was made to fin for most chrysomelids, particularly larvae, are not known. with tribes, for each the Old and New World. In numerous cases, data are given only genera (‘ Aulacoscelinae are not included as there no records of them leaf miners (Jolivet & Hawkeswood, 1995; Jolivet to Santiago have been omitted to save time and space. Blackwelder (1982) numerous other works were used update names geographica reported in references found, however, this may vary depending on host plant distribution. As much as possible, I have used the for those in the Pacific Region, all extracted from web. Gressitt provides coordinates for some of those islands or archipelagos. Bailey (1976); Brummitt & Powell (1992); Everett (1980); Greute Taxonomy (Agricultural Research Service, United States Department of Agriculture) http://www.ars-grin.gov/npgs/tax, http://www. (1968); Heywood Further studies will clarify many of those records. In cases where less species specific statements are made, such as “name on other congenerics have pointed out the leaf mining habit, except for hispines where it appears that is relatively saf instars live on stems. Actually, many hispines, such as the species in tribes Oediopalpini, Cephaloleiini, Hybosispini, Are Anisoderini, Aproidini, Callispini, Leptispini, and Eurispini (of the Old World Hispinae) are not leaf-miners. Many of those no their host plants (Maulik 1933a, b), thus, they are not included in this table. Other hispines, such as species of W3T/search/vast.html; were used to complete or correct nomenclatural (including authorship) information for host plants. Howeve status of insect and/or plant names. I was unable to find the author(s) thirteen scientific names; those names have be plants listed are those of the larvae but, when stated by authors, I have annotated host-feeding stage association. In host plant) or ‘adult’ has been added next to the plant, if source makes such difference. If not stated, I have assume as known, for adults. In the case of several alticine genera ( occasion, be leaf-miners and I have, very reluctantly, included them in Table 1. However, have not distinguished the host pla Sceloenoplini, Hispoleptini, Chalepini, Uroplatini (New World hispines), as well Callohispini, Exothispini, Coelaenomenoderi hispines) (Seeno & Wilcox, 1982). The Old World hispine tribe Cryptonychini has been omitted as their larvae feed on leaf buds, (Kalshoven, 1957), and they are not listed. Numerous other suspected leaf-miners have been excluded because I unable (1987); Munz & Keck (1973); Quattrocchi (1999); Plant Names (Australian National Botanical Gardens) http://www.anbg.gov.au/anbg Agriculture, Natural Resources Conservation Service) http://plants.usda.gov/cgi_bin/topics.cgi; The Trustees (1993), and Tropic for plant families, as in Apiaceae (= Umbelliferae), Arecaceae Palmae), Asteraceae Compositae), Brassicaceae Crucifera the use of Leguminosae, instead using names Caesalpiniaceae, Fabaceae, and Mimosaceae. Some hispine host plant data possible, but, if listed, they appear in quotation marks, and I have given the best approximation of a scientific name possible than those of larvae. Some illustrations (= illustr.) adults, and/or immature stages, feeding damage are noted. A det Leaf-mining chrysomelids 43 ek 1986; õ 1928 (illustr.); Riley and Enns 1979; 1928; Wilcox, no date, a (illustr.) 1928 et al. et al. et al. 1928 1928, Wilcox, no date, a (illustr.) et al. et al. ‘1959’ (1960) Blake 1939 (illustr.) (illustr.), Nakane 1955 (illustr.) 1948, Kaszab 1962, Medvedev and Zaitzev 1978, Steinhausen 1966 Hering 1957, Jolivet to Santiago-Blay (pers. comm., June 2003) Jolivet 1977, and Hawkeswood 1995 Steinhausen 1978 (illustr.); Wilcox, no date, a (illustr.) sp. Linné Pagony 1993 (illustr.); Pál and József 1977; Surányi 1942; Szontagh spp. Populus Michaux, 1900; Erdös 1935; Hering 1957; Jolivet 1948; Kaszab 1962; Koch Linné (illustr.); Jolivet 1948; Medvedev and Zaitzev 1978; Ma s sp., Clark 2000; Needham P. tremula Linné, Böving and Craighead 1931 (illustr.); Buhr 1955; Cavey 1994; Csiki Euonymus Juglans Michaux, Buhr 1955, 1956; Cox 1996 (illustr.); Hering 1957; Kaszab 1962 spp. Chenopodium sp. (several 1992; Lawson 1991 (illustr.); Lopatin 1984; Medvedev and Zaitzev sp. (Betulaceae), Buhr 1955, Csiki 1900, Grandi 1959 (illustr.), Hering 1957, Jolivet Populu Euonymus P. nigra Salix , Populus Salix Linné, P. alba sp. (Chenopodiaceae) P. virginiana Linné, sp. (several species 1982; Szontagh and Tóth 1977 (illustr.) sp. (several species Linné sp., Corylus spp., sp. (Santalaceae) P. grandidentata Blume, Salix (larvae),

P. nigra Linné, P. canadensis Michaux, Salix Rydberg, sp. (Betulaceae), sp. (Celastraceae) Jolivet 1977, Lee 1990 (illustr.), Medvedev and Zaitzev 1978 sp. (Celastraceae), sp., Ehrhart (Betulaceae), Linné, Populus Linné (Santalaceae) Jolivet 1977, to Santiago-Blay (pers. comm., June 2003) Populus sp., Chenopodium , P. nigra Linné, Linné, Santalum Linné (Salicaceae)Needham 1924, Frost Beta vulgaris Marshall Marshall, Corylus ceae Monrós sp. (Piperaceae) Vencl and Aiello (1997) (illustr.) P. tremula Populus Linné Linné, sp. (Lythraceae) 1996 (illustr.); Hering 1951, 1957 (illustr.) Cox sp. (Salicaceae) sp. (Salicaceae) Balsbaugh and Hays 1972, Wilcox, no date, a (illustr.) Needham sp. (Salicaceae) Needham sp., sp., Populus Linné, sp. (Salicaceae) Tripterygium Tripterygium Euonymus sieboldiana sp., Santalum album Populus alba Lythrum Atriplex album Populus Populus P. deltoides P. deltoides Populus alba mentioned by Buhr 1955, 1956, all larvae) Salicaceae Populus tremuloides Salix Populus acuminata (larvae), (larvae), ‘climbing vines’, possibly species in theCelastraceae or Sapindaceae Betula Reid 1989 tremula P. nigra (Salicaceae) Populus mentioned by Buhr 1955, all larvae) Salicaceae Betula verrucosa Populus alba Linné, species mentioned by Buhr 1955, all larvae,Needham (illustr.); 1978 Salicaceae) Corylus avellana Peperomia Siberia, Asia mountains, Great Britain Great Britain, Asia including east Africa, sp., and Japan (Salicaceae), Argentina Commelina Boheman (LeConte) Canada, United States (LeConte) western North America (Oke) Australia (Duftschmidt) Europe, Great Britain 2 quadrivittata Crotch central to eastern United States Zeugophora Vencl and Aiello central Panamá ) Jacoby India Fabricius Canada, United States, Europe, ) herein included. Madagascar, India, China, (Juglandaceae), Suffrian United States, central Europe, (Marsham) Europe, Great Britain, Asia (Baly) eastern Asia, Japan sp. Old World (mostly tropics) Crotch central to eastern United States Crotch United States Power central Europe, especially in angularis 3 Fall Canada, United States Neolema ( Pedrillia Z. annulata Z. andrewesi Taxon AuthorTaxon ZEUGOPHORINAE Zeugophora abnormis Geographical Distribution Reported Host Plants Author FamilyReferences Selected Galerucella pusilla Monoxia Z. atra Z. consanguinea Z. flavicollis Z. puberula Z. scutellaris Zeugophora vitinea Zeugophora Z. varians Z. turneri Both subgenera ( and Z. subspinosa CRIOCERINAE Lema GALERUCINAE Oulema pumila 44 J.A. Santiago-Blay circa 1973 et al. 1928 et al. 1990, Kondratieff to San- 1973, Santiago-Blay 1990, et al. et al. 1990’s), Lawson 1991 (illustr.) circa 1998 Blake 1939 (illustr.) Santiago-Blay (unpl. data) 1931 (illustr.); Essig 1958; Needham Blake 1939 (illustr.) Blake 1939 (illustr.), Santiago-Blay and Virkki 1996 Blake 1939 (illustr.), Cavey to Santiago-Blay (pers. comm., 1990’s), Santiago-Blay (unpl. data) Blake 1939 (illustr.) sp., Blake 1939 (illustr.), Kirk and Balsbaugh 1971 Standley Atriplex Linné Blake 1939 (illustr.), Cooley 1916 sp., Banham 1962, Blake 1939 (illustr.), Halford sp. Chenopodium Pursh) Blake 1939 (illustr.); Böving 1929 and Craighead Grindelia (Pursh) Blake 1939 (illustr.), Hatch 1971, Smith 1930, Santiago-Blay (unpl. Linné tiago-Blay (pers. comm., Beta vulgaris sp. larvae Blake 1939 (illustr.), Cranshaw ex (Poaceae) Beta vulgaris

Chenopodium Pursh (Asteraceae), data) Quercus Artemisia sp. (Salicaceae) (Pallas

Besser larvae, Artemisia Blake 1939 (illustr.), Santiago-Blay (unpl. data), R. Brown, (Torrey) S. Wats (larvae), tiago-Blay and Virkki 1996 (Pursh) Britton and Rusby, Chenopodium album (Pursh) Dunal, A. Gray (Brassicaceae), Blake 1939 (illustr.), Hatch 1971, Santiago-Blay (unpl. data), San- Nuttall, sp. (Chenopodiaceae), (Bentham) D. Dietrich Blake 1937 (illustr.), 1939 (illustr.) Amaranthus watsonii Iva axillaris Populus Medicago sativa (Pursh) Nutall, Hooker and Arnold (Asteraceae)Halford (illustr.), 1939 Blake sp. (Asteraceae) larvae, (Linné) Moench sp. (Asteraceae) Blake 1939 (illustr.) Linné (Bataceae) Blake 1939 (illustr.) sp. (Asteraceae), Gutierrizia sarothrae Linné, sp. (Chenopodiaceae) sp. (Asteraceae) sp., sp. (Chenopodiaceae) Blake 1939 (illustr.), Fall 1927 sp. (Asteraceae) sp. (Chenopodiaceae) larvae, Grindelia Chenopodium Linné (larvae) Chenopodiaceae, ‘a wild album desert plant’ Chrysothamnus nauseosus Britton Chrysothamnus Grindelia humilis Grindelia squarrosa Solidago (Amaranthaceae) Atriplex semibaccata (Fagaceae), ‘ground cherry’, nut’, ‘gum’, ‘Gipsey flower’, ‘hackberry’ sp. (Chenopodiaceae) larvae; Beta vulgaris (Chenopodiaceae), Atriplex canescens Atriplex Linné, Sorghum bicolor Artemisia tridentata Solidago Artemisia douglasiana sp. (Asteraceae) and Virkki 1996 Batis maritima Chenopodiaceae Atriplex barclayana sp. Atriplex Beta vulgaris Linné, Chenopodium (Chenopodiaceae), (Leguminosae) Lepidium alyssoides Atriplex confertifolia Gutierrezia sarothrae Gutierrezia Artemisia Chrysothamnus Linné (Chenopodiaceae) Unknown Maes and northern Mexico California, Mexicoor (Chenopodiaceae) United States California, islands in Gulf of California (Mexico) Baja California (Mexico)Rusby, and Britton Blake) United States inornata Jacoby Guatemala, Nicaragua Blake western United States Blake western United States Blake western United States (LeConte) western United States Blake southwestern United States (LeConte) western United States (LeConte) western United States and Blake western United States Blake southwestern United States Blake southwestern United States Blatchley southeastern United States Blake United States LeConte southwestern United States Blake northern Mexico, Baja Blake Santa Inez Island, Gulf of Blake western Canada and Blake western United States sp.(near M. M. consputa M. minuta Monoxia M. brisleyi M. debilis M. obesula M. elegans M. grisea M. guttulata M. inornata M. batisia M. beebei M. obtusa M. apicalis M. pallida M. puberula M. schizonycha M. semifasciata M. sordida Leaf-mining chrysomelids 45 1999 et al. 1977 et al. 1995; Konstantinov and Vandenberg 1996 Chen 1934; Gressitt 1963; Inoue 1990a, 1990b, 1996; Lee 1992 Hering 1957 Hering 1957, Koch 1992 sp., sp., Hering 1957, Koch 1992 sp., sp., Buhr 1956; Hering 1951 (illustr.), 1957; Jolivet and Hawkeswood sp., Plantago Prunella purpurea O. sp., Plantago

Stachys Beta Linné Chen 1934, Zaka-ur-Rab 1991 L. sp., sp., Buhr 1955, 1956; Hering 1957; Kaszab 1962 (illustr.); Koch 1992; sp. (G. Don) Ligustrum Ajuga Ajuga sp., sp. Lamarck Carr, sp., Sibthorpia Linaria Galeopsis W. T. Aiton, Bellis sp. (Asteraceae),Teixeira 1942; Surányi Digitalis Syringa reticulata Digitalis sp. (Chenopodiaceae), sp. (Primulaceae), sp., Lamium Linné (Oleaceae) sp., Teucrium sp. (Plantaginaceae), sp., Saxifraga granulata Plantago F. mandshurica fortunei Satureja sp. (Lamiaceae), Linné, sp. (Schrophulariaceae) sp. (Scrophulariaceae) Cirsium E. hypericifolia x sp. (Lamiaceae), sp., Maximowicz, Osmanthus fragans Ajuga L. lucidum O. heterophyllus (Torrey) S. Wats, unable to find name, sp., Primula sp. (Onagraceae), Linné, Kickxia sp. (Asteraceae), Plantago (Linné) Schreber, Blume, S. vulgaris Teucrium Linné, Miers (Solanaceae) Rhinanthus glaber Linné, Veronica Scrophularia Chenopodium Veronica sp., Prunella Siebold and Zuccarini, japonica Linné, Circaea Prunella Osmanthus sp., sp. (Euphorbiaceae) Grandi 1959, Kaszab 1962 sp., Thunberg, Hasskarl, sp., sp., Galeobdolon sp., Bellis perennis Linné, sp., Solidago sp. (Leguminosae) Ward sp. (Saxifragaeae), (Hasskarl) Mouille, sp. (Ranunculaceae)1957 Hering sp. (Lamiaceae), sp. (Scrophulariaceae) R. graminis Linné, sp. (Primulaceae), sp., Digitalis reptans sp., sp.,

sp., Stachys (Blume) H. Hara, ilicifolius Origanum sp. (Plantaginaceae), japonicum L. obtusifolium ovalifolium (Thunberg) Loureiro, P. S. Green, Ajuga chamaepitys Veronica (Plantaginaceae) Bellis Saxifraga Rhinanthus Fraxinus japonica Euphorbia (Euphorbiaceae) Verbascum Prosopis Euphorbia hirta Lamiaceae Atriplex confertifolia vulgaris Lycium pallidum Ajuga sp., Veronica Lamium Linné (larvae) (Saxifragaceae), Pedicularis (larvae), Rhinanthus Anemone Aster Teucrium scorodonia (Lamiaceae), Centaurea nigra Ajuga lanceolata Primula Linné, Britain China Ruprecht var. southwestern United States (Linné)Great Europe, Motschulsky eastern Siberia, Korea, Japan, Koch Europe 2 Duvivier India, China Gebler eastern Europe Marsham Europe, Great Britain sordida Alluaud Europe, Great Britain sp. Europe, north Africa Illiger Europe, Great Britain nr. Argopistes biplagiatus A. splendida Apteropeda ALTICINAE Taxon AuthorTaxon Aphthona cyparissae Geographical Distribution Reported Host Plants Author FamilyReferences Selected A. nigrilabris Apteropeda bellidiastrum Monoxia A. globosa A. nigritarsis A. orbiculata 46 J.A. Santiago-Blay 1928 (illustr.); Riley and Enns 1979; et al. 1963, Hering 1957, Kaszab 1962, Koch 1992, Lee and 1991 et al. et al. 1996, Kaszab 1962, Koch 1992, Medvedev and Zaitzev 1978 Hering 1957, Kaszab 1962, Medvedev and Zaitzev 1978 (illustr.), Wilcox 1954 Inoue and Shinkaji 1989; 1990a, 1990b, 1996; Lee 1992; Böving and Craighead 1931 (illustr.); Byers 2002; Clark 2000; Lawson Hering 1951, 1957 (illustr.); Kaszab 1962; Koch 1992; Steinhausen 1966; Surányi 1942 Böving and Craighead 1931 (illustr.); Hering 1957; Kaszab 1962; sp. Jolivet and Hawkeswood 1995 sp. x1973 Samuelson sp. sp. Furth 2000 sp., sp. Hering 1957, Kaszab 1962, Koch 1992 Salvia Linné, sp. Linné, Grison sp., Euphorbia Veronica Linné, Grandi 1959 (illustr.), Hering 1957, Kaszab 1962, Koch 1992 Linné Osmanthus Linné, sp. (Rutaceae) Jolivet and Hawkeswood 1995 Linné Zabel Clematis Stachys Eresmostachys Clematis Chenopodium Linné, Osmanthus Trollius major Linné (Solanaceae) Lamium purpureum

Jacquin, sp., sp., P. Eryngium campestre Marrubium sp. (Plantaginaceae) 1991 (illustr.); Needham C. maritima sp., S. nemorosa Linné, sp. (Fagaceae), Linné (Boraginaceae) Cox 1996 (illustr.), Hering 1957, Koch 1992 Beta sp. (Rutaceae) Salvia nemorosa Linné, Thunberg, sp. (Lamiaceae), Ballota Zanthoxylum Linné, (Jacquin) R. Brown(illustr.) 1963 Selman Triticum vulgare Linné, sp. (Asteraceae), Linné (Brassicaceae), Zaka-ur-Rab 1991 S. verticillata Linné, sp., (Linné) Trev., Citrus Linné (Leguminosae), sp., Plantago Quercus sp. (Apiaceae) Lopatin 1984 Jacquin, C. vitalba (Linné) Correa Serra (Rutaceae) Cox 1996, Chen 1934, Zaka-ur-Rab 1991 Linné, Ranunculus Linné (Polygonaceae) Koch 1992, Vig to Santiago-Blay (pers. comm., May 2003) Salicornia europaea (G. Don) P. S. Green, Linné, Cynara Teucrium Linné, Linné, Solanum melongena sp. (Phytolaccaceae), pseudochamaedrys Citrus Dcne., sp.,

Carr (Oleaceae) Linné, sp., sp., Eryngium Marrubium sp. (Alismataceae), sp., sp. (Lamiaceae) Linné, Galeopsis tetrahit (Lamiaceae) Leucas martinicensis (Lamiaceae) Stachys officinalis S. pratensis (Lamiaceae) (Schrophulariaceae) Salvia austriaca Veronica Clematis flammula C. recta (Chenopodiaceae) Plantago lanceolata heterophyllus Aegle marmelos Aegle Tymus Stachys Cynoglossum officinale Marrubium vulgare Atriplex hastata Alisma album Ligustrum japonicum fortunei (Ranunculaceae) Cirsium P. rugelii sp., Linné, Ballota nigra Phytolacca Pulsatilla Poaceae(Poaceae), Rumex crispus Cox (Euphorbiaceae), Brassica campestris Crotolaria juncea Oryza sativa (Ranunculaceae), mountains especially in and western Europe (Japan), Taiwan, southeastern former Soviet Union Linné, Canada (introduced) Asia, Micronesia (Gyllenhall)Asia Europe, Csiki Korea, Japan, Ryukyu Islands Letzner central and southern Europe Koch Europe, western part of Germar southern and central Europe Chevrolat central United States (Koch)Britain Great Europe, Redtenbacher central Germany, southern (Marsham) Europe, Asia, Morocco, and Baly India, China Bach central and western Europe, Selman Ethiopia Baly India, Sri Lanka Illiger Europe, Asia sp. Old World sp. southeast Asia D. heringi D. foersteri D. femoralis Clitea picta Clitea Dibolia borealis D. cynoglossi D. cryptocephala Argopus ahrensi A. coccinelliformis Argopus D. depressiuscula C. concinna C. tibialis C. basalis Chaetocnema aridula Leaf-mining chrysomelids 47 1928 1928, Vig to Santiago- 1928, Wilcox 1954 et al. et al. et al. n 1973 Frost 1924, Hering 1957, Konstantinov and Vandenberg 1996 Blay (pers. comm., May 2003) Lawson 1991 (illustr.), Needham Lawson 1991 (illustr.) Buhr 1955, Hering 1957, Kaszab 1962, Koch 1992 Cox 1996 (illustr.); Balsbaugh and Hays 1972, Böving Craighead sp. 1984, Medvedev and Zaitzev 1978 R. Linné, Linné sp. and Zaitzev 1978 (illustr.), Needham Satureja Linné, 1931 (illustr.); Clark 2000; Needham R. Linné sp. Hering 1957, Kaszab 1962, Koch 1992 Linné, Buhr 1956, Hering 1957, Kaszab 1962 (illustr.), Koch 1992, Vig to sp., Linné, Physalis sp., S. carolinense Prunella vulgaris crispus Schultes, Buhr 1955, 1956; Hering 1957 (illustr.); Kaszab 1962; Koch 1992; Rumex acetosella Linné,

Equisetum et sp. (Boraginaceae), Buhr 1955, Hering 1957 (illustr.), Kaszab 1962, Koch 1992, Lopatin sp. (Lamiaceae) Buhr 1956, Hering 1957, Kaszab 1962, Koch 1992 S. tuberosum Eryngium R. Mentha aquatica Nepeta pannonica sp. (Lamiaceae) Santiago-Blay, (pers. comm., May 2003) R. obtusifolius sp. (Lamiaceae) Schultes (Arecaceae), Byers 2002, Cox 1996 (illustr.), Kaszab 1962 Medvedev Mentha Miller, sp. (Ranunculaceae) et S. nemorosa R. obtusifolius acetosella sp., sp.,

sp. (several other species sp. (several species P. Miller, Salvia Stachys Linné, Linné (Adantiaceae) Samuelson 1973 Linné, Wood, Roemer Stachys Linné (Equisetaceae), Frost 1924, Jolivet and Hawkeswood 1995, Konstantinov Torrey, Rumex Rumex Linné, Symphytum Clematis Roemer Rumex crispus Lamium Torrey, sp. (several species mentioned sp. (Cistaceae) Hering 1957, Koch 1992 sp., Leonorus Linné, Linné, Plantago sp. (Davalliaceae) Samuelso sp., sp., sp., genera in the Asteraceae, Boraginaceae, Buhr 1956, Jolivet and Hawkeswood 1995 sp. (Plantaginaceae), sp., sp. (Brassicaceae), sp., sp. (Dipsacaceae), Linné, Mentha Nepeta S. melongena R. altissimus sp. (Polygonaceae) Vandenberg 1996 hymenosepalus

Succisa Pulmonaria (Lamiaceae), mentioned by Buhr 1955, all larvae Plantaginaceae), Ranunculus Rumex Convolvulaceae, Dipsacaceae, Lamiaceae, Linaceae, Plantaginaceae, Ranunculaceae, Schrophulariaceae, Solanaceae, and Thymelaceae. Linné (Lamiaceae) is a doubtful record. (Polygonaceae), ‘rye’ Equisetum arvense Eryngium campestre Linné, Linné, Galeopsis Nephrolepis Polygonum arvense Polygonum Rutabaga hymenosepalus S. verticillata Solanum americanum (Solanaceae) Acrostichum aureum scutatus Brunella Salvia pratensis Lycopersicum esculentum Sabal serrulata Linné, by Buhr 1955, all larvae, Lamiaceae) Stachys recta mentioned by Buhr 1956, all larvae) Polygonaceae Plantago Linné, R. (Polygonaceae) Helianthemum Caucasus, Mongolia, Siberia (Equisetaceae), Daghestan, Kazakhstan, Turkey Europe (Apiaceae) almost all Europe, Turkey, Europe), Armenia, Caucasus, Kowarz Europe, Great Britain Linné northeastern United States, Scopoli Europe, Great Britain, Asia Harold New World, cosmopolitan? Bryant Fiji sp. Palearctic, North America Curtis Europe, Great Britain sp. worldwide Numerous Koch Europe Crotch eastern and central United States (Letzner) Europe (except northern Redtenbacher central and southern Europe Clark Fiji Illiger central Germany to southern sp. Holartic, Africa, Central America Longitarsus luridus Longitarsus Mantura chrysanthemi Hippuriphila D. timida Dibolia F. venusta Febra insularis Taxon AuthorTaxon D. occultans Geographical Distribution Reported Host Plants Author FamilyReferences Selected D. schillingi Epitrix cucumeris Hippuriphila modeeri D. rugulosa M. floridana M. matthewsi 48 J.A. Santiago-Blay a et al. 1977; et al. 1963 (illustr.); M. subobtusata et al. (Gyllenhal 1813) 1928 1928 1928 M. obtusata et al. et al. et al. 2001 Buhr 1955, 1956; Cox 1996 (illustr.); Grison Kaszab 1962 (illustr.); Hering 1957 Lipa Lopatin 1984; Medvedev and Zaitzev 1978 (illustr.); Surányi 1942; 1963; Hering 1957; Kaszab 1962 (illustr.); Lopatin 1984; Medvedev Buhr 1955, 1956; Gressitt and Kimoto 1963; Hering 1957 (illustr.); sp., Vig 1989 (illustr.), 2000 Linné, sp., and Zaitzev 1978 (illustr.); Wilcox 1954; Vig 1999; Verdyck (Linné) Digitalis Andrzeiov- Gaertner, sp. Hering 1957, Kaszab 1962 (illustr.), Koch 1992 Bigelow Needham Rumex sp. Linné, Kaszab 1962 (illustr.); Koch 1992; Lopatin 1984; Surányi 1942 Arabidopsis spp. (several Linné Nasturtium Linné, Armoracia sp., Anchonium Cardamine amara Brassica napus A rusticana A. arenosa Anastatica Grande, Böving and Craighead 1931 (illustr.); Buhr 1955; Grison Alyssoides arduini sp. (Polygonaceae) Cox 1996 (illustr.), Kaszab 1962, Koch 1992 rusticana Linné (Rubiaceae), Rheum tanguticum Teucrium

et Sisymbrium officinale (Linné) Scopoli Rheum Anagallis C. edentula sp. (Plantaginaceae) sp., Rheum sp., R. crispus Usteri, Plantago lanceolata Linné (Hypericaceae), Hering 1957, Koch 1992 purpurea Rumex R. crispus Barbarea stricta Gaertner (Cistaceae), Buhr 1956, Konstantinov and Vandenberg 1996, Surányi 1942

Linné (Brassicaceae) Needham Linné, Linné (larvae), Linné, Linné (Brassicaceae) Needham Linné, Linné (Solanaceae) Linné, R. Brown, Armoracia Nutall, Linné (Anacardiaceae), A. hirsuta Plantago Alyssum Alliaria spp. (many species listed in Buhr (Linné) Koch, Galium verum Linné, Linné, Boissier (larvae), Linné (larvae), Murray, Digitalis Linné (larvae), Bönningh (Tropaeolaceae) sp., (M. B.) Cavara sp. (larvae), Arabis Linné, sp. (Polygonaceae) Lopatin 1984 acetosa B. nigra Sinapis alba sp. (larvae) (Polygonaceae) spp. (several species mentioned by Buhr Buhr 1956; Gruev and Doeberl 1997 consider sp. (Polygonaceae)(illustr.) 1957 Hering media

Arabis alpina hiercochuntica Scopoli (larvae), (larvae), 1955, all larvae), Barbarea vulgaris Linné, Linné, Solanum dulcamara Lepidium virginicum elichrysifolium Aethionema Meyer, and Scherbius P. Pistacia lentiscus sp. (Scrophulariaceae), Hypericum perforatum Anagallis arvensis (Primulaceae), Alliaria ta Armoracia lapathifolia (Linné) Scopoli (Brassicaceae), microphyllum (Lamiaceae), Aecidium virginicum Cakile americana Hooker (Brassicaceae) Maximowicz ex Balfour , species mentioned by Buhr 1956, all larvae), Rumex acetosa (Polygonaceae) Teucrium scorodonia 1956, all larvae) PolygonaceaeHelianthemum vulgare of synonym Polygonum aviculare Rumex Rumex Rumex Rumex Polygonum Rumex Polygonum aviculare conglomeratus northern Africa, eastern Asia Fritsch (larvae), eastern Asia Gaertner, Mey, and Scherbius, Europe, Great Britain Europe, Great Britain to Vietnam New World, Africa, China, Europe China (Crotch)States United Koch Europe Illiger central and south eastern (Crotch) eastern United States Jansson Waltl Europe, Great Britain Lopatin Central Asia (Koch) United States and Canada, (Linné) Europe, Great Britain, (Gyllenhall)Britain Great Europe, Schaeffer southeastern United States Linné Europe, Great Britain, Siberia, sp. Palearctic, a few species in Phyllotreta aenicollis Ochrosis ventralis P. armoraciae P. nemorum P. chalybeipennis P. liebecki Mniophila muscorum Mantura M. subobtusata M. mesasiatica M. obtusata M. pallidicornis M. rustica Leaf-mining chrysomelids 49 B. sp., sp., sp., B. sp., rapa O. E.

E. Berteroa Linné Diplotaxis sp., B. sp., sp. (Zap.) Brassica Goldbachia Euclidium sp. (several Linné Erucastrum sophia Calepina Linné

Camelina sp., E. helveticum B. auriculata sp., E. tenuissimum Berteroa sp., Cardaria draba (Linné) De sp., Eruca sp., sp. (many species Hirschfeldia Erophila Conringia Hesperis (Juslen) DeCandolle spp. (numerous sp., E. hieraciifolium Zap. (larvae), B. napus Linné (larvae), Barbarea unable to find name B. chinensis Bunias Cheiranthus alpinus sp., Descurainia Draba Cochlearia officinalis E. pieninicum D. muralis Biscutella Reichenbach (larvae), Euclidium Descaurainia sp. (several species listed DeCandolle (larvae), Halacsy (larvae), Linné, sp., Ehrhart (larvae), (Miller) Aschers (larvae), sp., Brassicella erucastrum Linné (larvae), Heldreich and Sartorelli Erysimum sp. (several more species listed D. tenuifolia (Linné) Koch (larvae), Erysimum cheiranthoides (Linné) Cr. (larvae), Cardaminopsis Crambe Braya sp., C. kewensis Cochlearia E. hugaricum DeCandolle (larvae), B. verna Cardamine amara Linné (larvae), sp., sp., sp., Linné (larvae), Linné (larvae), (Linné) R. Brown (larvae), Cheiranthus C. senoneri Diplotaxis DeCandolle (larvae), spp. (several species listed in Buhr 1955, (Willdenow) O. E. Schulz (larvae), Linné (larvae), E. diffusum B. nigra B. oleraceae R. Brown (larvae), (Linné) DeCandolle (larvae), var. erigerifolia Brassica ski (larvae), Hugueninia tanacetifolia vulgaris listed in Buhr 1955, all larvae), Eruca sativa B. Fedtschenko (larvae), incana Schulz(larvae), (several species listed in Buhr 1955, all larvae), Erucaria myagroides gallicum Erucastrum (larvae), (Jacquin) DeCandolle (larvae), Linné (larvae), linifolium Pawlowski (larvae), species listed in Buhr 1955, all larvae), syriacum laevigata Biscutella auriculata Linné laevigata campestris listed in Buhr 1955, all larvae), Linné, Camelina sativa Capsella all larvae), Cardamine (Linné) Desvaux (larvae), Linné (larvae), (larvae), (larvae), Linné (larvae), (larvae), (larvae), cretacea Candolle (larvae), (larvae), in Buhr 1955, all larvae), (Linné) Webb (larvae), Coronopus in Burh 1955, all larvae), Taxon AuthorTaxon Geographical Distribution Reported Host Plants Author FamilyReferences Selected 50 J.A. Santiago-Blay S. Linné S. Ricotia Raphanus spp. spp. Matthiola L. ruderale Tovaria spp. Limnanthes glabra L. densiflorum Linné (Linné) Bentham S. officinale Lepidim (Leyss.) Bess. sp. (more species Petrocallis pyre- Turritis Linné (larvae), Hooker (larvae), sp. (many listed in Lesquerella Rorippa amphibia Willkomm and Lange Sisymbrium Linné (larvae), Malcolmia (Linné) Allard (larvae), ex sp., Raphanus raphanistrum sp. (many species Sweet (larvae), Linné (larvae), Iberis sp., Sinapis sp. (several species listed Isatis tinctoria R. palustris Linné (larvae), sp. (Brassicaceae), sp., Gynandropsis gynandra Sisymbrium altissimum (Linné) Bess. (larvae), (Linné) Desvaux (larvae), sp., Kremeriella cordylocarpus Jaubert and Spach (larvae), L. heterophyllum Linné (larvae), Neslia paniculata Cosson Cleome Sibthorp and Smith (larvae), Peltaria Sinapis alba Reseda S. orientale Turritis Lepidium R. sativus Lunaria Linné (larvae), sp., sp., Linné (larvae), unable to find name (larvae), (Linné) R. Brown (larvae), Linné, sp., Isatis L. perfoliatum R. sylvestris S. loeselii Matthiola annua Rapistrum rugosum Hutchinsia Linné (larvae), spp. (other species mentioned by Buhr Ruíz and Pavón (larvae) Touvariaceae, sp. (numerous species listed by Buhr Thysanocarpus curvipes DeCandolle (larvae), R. Brown (larvae), Myagrum (larvae), (Cosson and Dur.) Maire (larvae), campestre Schaeder (larvae), Linné (larvae), (several species listed in Buhr 1955, all larvae), Lobularia maritina Lobularia (several species mentioned by Buhr 1955, all larvae), (larvae), (larvae), Buhr 1955, all larvae), Hutchinsia alpina sp., Desvaux (larvae), naica Linné (larvae), spp. (several species mentioned by Buhr 1956, all larvae), (Linné) Briquet (larvae) Capparaceae, spp. (several species listed in Buhr 1955, Limnanthaceae), in Buhr 1956, all larvae, Resedaceae), pendula Capparis rupestris C. spinosa listed by Buhr 1955), Linné (larvae), R. sylvestre lunaria (Linné) Bess. (larvae), listed by Buhr 1956), (larvae), strictissimum (several species listed by Buhr 1956, all larvae), Texiera glastifolia Thlaspi 1956), Turritis glabra (larvae), Rorippa 1956, all larvae), arvensis (Linné) Scopoli, Leaf-mining chrysomelids 51 1977 et al. 1963, Lopatin 1984, Medvedev 1963, Hering 1957, Kaszab 1962 1928 et al. et al. et al. 1963 (illustr.), Hering 1957, Kaszab 1962, et al. Cox 1996 (illustr.), Grison Kaszab 1962 (illustr.), Kalshoven 1981 (illustr); Lipa Buhr 1955, 1956; Böving and Craighead 1931 (illustr.); Cox 1996 (illustr.), Grison Clark 2000, Needham Kato 1991 (illustr.) Kato 1991 (illustr.) spp. spp. Buhr 1955, 1956; Cox 1996 (illustr.); Hering 1957 B. sp. spp. Steinhausen 1978 (illustr.) Linné, Rorippa Rapistrum prothifolia majus

Capsella T. minus

sp., sp. Cox 1996 (illustr.), Grison sp., Eruca sativa Lunaria spp. (several C. Thlaspi Linné, DeCandolle, Buhr 1955, 1956; Clark 2000; Hering 1957; Kaszab 1962 sp., not leaf miners Loxogramma sp. (Poaceae) Lunaria ex sp., arvensis sp.,

Linné (larvae), Zea Pr., Lunaria Raphanus sativus Linné, Barbarea Agropyron Hyoscamus Bunias Tropaeolum Cakile amara Raphanus Sinapis

sp., B. oleraceae Smith (larvae), sp., and sp., Linné, arvense sp., Linné (Brassicaceae) Hering 1957, Kaszab 1962, Koch 1992

Aiton f., Andrzeiovski Linné, Linné, (Linné) DeCandolle, (Thunberg) Ching, Crambe microphyllum Linné,

Setaria (Linné) Medikus, Cardamine Bunias niger Brassica (Brassicaceae) both larvae (illustr.) Thlaspi

sp., sativus sp. (Asteraceae)1957 Hering Makino (Polypodiaceae) sp. (several species listed in Buhr 1956,

sp., sp., sp., elliptica (Linné) Allard,

sp., sp. (Poaceae) and numerous plants, Chen 1934; Medvedev and Zaitzev 1978; Vig 1996, 1998 sp. (Resedaceae), Linné, rapa leaves. Lepidium virginicum Tropaeolum aduncum Linné (larvae) Reseda Brassicaceae Berteroa incana Setaria particularly of the Brassicaceae and Poaceae. Vig (1998) reports that larvae are Linné (Brassicaceae) Brassica napus bursa-pastoris but feed on the surface of Hordeum Colysis Brassica Raphanus all larvae) Brassicaceae (several species mentioned by Buhr 1956, all larvae) Brassicaceae Biscutella laevigata Lemmaphyllum salicifolia (Don) Pr. (Polypodiaceae) (Brassicaceae) Alliaria officinalis Brassica Cardamine species listed in Buhr 1955, all larvae), (several species listed in Buhr 1955, all larvae), Raphanus perenne Sinapis Barbarea vulgaris Centaurea Hyoscyamus America DeCandolle eastern Asia Africa, Asiaeastern Asia, North Linné (Tropaeolaceae), several other genera of and Zaitzev 1978 Great Britain, north Africa,western Asia (Solanaceae) Great Britain (several species listed in Buhr 1955, all larvae), United States Africa, and southeastern Russia, (Linné) Europe, Ireland, Great Britain, Takizawa Japan AbeilleAfrica north (Crotch) central and eastern United States hiranoi (Linné) central and southern Europe,

Kutschera United States, Europe, northern (Illiger) central and southern Europe, (Fabricius) central Europe, and Chen Japan (Redtenbacher) Europe, Asia Fabricius Alps (Europe) (Fabricius) Europe, Great Britain, northern vittula . Taxon AuthorTaxon Geographical Distribution Reported Host Plants Author FamilyReferences Selected P. zimmermani P. undulata P. vittata P Psylliodes chrysocephala S. sauteri P. marcida P. napi P. toelgi Schenklingia P. erythroceros P. hyoscyami 52 J.A. Santiago-Blay et al. 1963 (illustr.); et al. Hering 1957 (illustr.); Kaszab 1962 Hering 1957, Jolivet and Hawkeswood, Konstantinov Vandenberg Böving and Craighead 1931 (illustr.), Buhr 1955, Grison Samuelson 1973 sp., sp., jacea sp. sp., 1996, Lee and Furth 2000

sp. Cirsium sp., (Linné) sp., Petasites C. sp. sp. sp., sp., Cirsium Hudson 1963, Hering 1957, Kaszab 1962 Cynara sp., sp., sp. (many Sprengel, Buhr 1955, 1956; Grandi 1959 (illustr.); Grison sp., sp. Smilax sp. sp., C. pycno- Galactites Carduus sp. (Musaceae), sp., sp. Artocarpus DeCandolle Selman 1963 (illustr.) (larvae), (Linné) Scopoli, Clematis Silybum

sp. (Poaceae), sp., Carthamus Scilla Onopordum Psychotria Centaurea Miscanthus Cirsium C. crispus Musa Onopordum Scabiosa Cynara sp., Centaurea Bidens Akebia Wedelia biflora sp., C. lanceolatum sp., Carduus sp. (Commelinaceae), sp., sp., sp., sp., Onopordon sp. (Pandanaceae) Smilax sp., sp. (Onagraceae) pycnocephalus Linné, unable to find name Monrós and Viana 1947 (illustr.)

Linné (larvae), sp., sp. (Salicaceae), Lilium Sch. Bip. (Asteraceae) Selman 1963 (illustr.) Farfugium Linné, Linné (larvae), C. oleraceum sp., Coffea Carduus Serratula chrysanthemifolia sp., Bernhardi (larvae), Circaea

Cirsium sp., Commelina Arctium Sasa Carthamus Linné (Asteraceae) Zaka-ur-Rab 1991 Salix Carduus Lappa Miller or Schrank Freycinetia scolymus Andropogon acanthium sp., Linné (larvae),

sp. (Pandanaceae), sp.,

(Linné) Scopoli, sp. (Moraceae), sp., Linné (larvae), schimperi sp., sp. (Asteraceae), sp. (Asteraceae) C. nutans sp., sp. (Arecaceae),

sp., sp., (Linné) Scopoli, C. scabiosa sp. (Rutaceae) Zaka-ur-Rab 1991 sp. (Leguminosae), Senecio Cynara Panicum Artocarpus Freycinetia (Ranunculaceae), (Rubiaceae), (Smilacaceae), various Zingiberaceae Citrus Guizotia sp., (Asteraceae), Vigna (Liliaceae), (Lardizabalaceae), Dichrocephala Centaurea angustifolia Dichrocephala Arctium Guizotia Carduus Linné, Carduncellus Quetzalia uruquensis (Celastraceae) (Asteraceae) Bidens pilosa sp., Ageratum (larvae), acanthoides Onopordon Serratula cephalus Scopoli Non Hill, C. palustre more listed in Buhr 1955), Serratula Ponapea arvense (Dipsacaceae), Arctium minus northern Africa Ethiopia Ethiopia Germany, Great Britain, South America Eastern Carolines),Solomons Islands DeCandolle (Asteraceae) larvae, (Moraceae), Selman Maulik India Spaeth Argentina sp. Worldwide, absent from bipunctatum

Jacoby India (Fabricius)Britain Great Europe, Gressitt Micronesia (Central and 4 Selman Graells southern Europe up to central Throscoryssa citri HISPINAE S. rubidum Sphaeroderma Acanthodes unca S. brevicornis S. guizotiae Sphaeroderma S. wedeliae S. testaceum Leaf-mining chrysomelids 53 Monrós and Viana 1947 (illustr.), Staines 1993 Thomas and Werner 1981 1979 Uhmann 1968 sp., Maulik 1937 sp. sp., Coelogyne sp. sp., Schizachy- sp., sp., Combretum sp., Pandanus Phalaenopsis sp. (Leguminosae), sp., Pandanaceae)1989 Anand Sorghum (Linné) Chase Monrós and Viana 1947 (illustr.) Panicum Hyparrhenia Humboldt, Bonpland Maulik 1929 (illustr.), Staines 1993 sp., sp., Arundina (Swartz) Rydberg Staines 1994c sp. (Arecaceae), Jolivet 1989a, Staines 2003b, Uhmann 1953 sp., sp., larvae Kalshoven 1957 (Scheele) Walpers Ford and Cavey 1985 (illustr.), Riley Enns 1982, Staines 1994c, (Linné) Linné (Poaceae) Riley and Enns 1982, Staines 1994a (Linné) Linné (Poaceae) Staines 1994a, Thomas and Werner 1981 Trinius (Poaceae) Lopatin 1984 Pandanus Phalaenopsis E. Morren (Bromeliaceae) Monrós and Viana 1947 (illustr.) Isoberlinia Jacquin (larvae) Noguera 1988 (illustr.), Staines 1994b Andropogon Linné (Poaceae) larvae Cox 1996 (illustr.), Ford and Cavey 1985 Riley Enns sp. (Vitaceae), several genera sp., Coelogyne sp. (Bignoniaceae), Bambusa (Michaux) Nash (Poaceae) larvae, Phoenix Sporobolus Valota insularis Miscanthus sp. (Orchidiaceae) sp. (Orchidiaceae), sp., sp. (Poaceae) Jolivet 1989a sp. (Arecaceae) larvae Kalshoven 1957 sp., Cissus Panicum latifolium sp. (Pandanaceae) larvae Kalshoven 1957 sp. (Orchidiaceae) larvae Kalshoven 1957 sp., sp., virgatum (Poaceae) larvae Maulik 1937, Kalshoven 1957 sp. (Ochnaceae), sp.,

sp., Dendrodium Tripsacum dactyloides larvae ‘sweeping grasses of glade communities’ Malvastrum auranticum rium scoparium (Poaceae) Olyra and Kunth (Poaceae) both larvae Bothriochloa saccharides Panicum maximum Paspalum Spathoglottis (Pandanaceae), sp., Tripsacum dactyloides Loudetia Rottboelia (Poaceae), of Zingiberaceae. Staines (2003b) has been unable to confirm records in the Zingiberaceae. Panicum Lophira Phragmites Ananas macrodontes Phragmites communis Metroxylon Stereospermum sp. (Combretaceae), Coelogyne Spathoglottis Metroxylon Pandanus Arundinacea Africa ‘grasses’ ArgentinaUnited States Poaceae (larvae) (Malvaceae) adults; MexicoMexico (Poaceae) United States and Mexico larvae, ‘probable grass feeder’ Java Borneo ? orchid (Orchidaceae) Asia, Africa Java bamboo Java Thompson French Guyana to Argentina Zoubkoff central Asia (Newman) central and eastern United States (Gestro) Uhmann Monrós and Viana Pic Paraguay and Argentina sp. central Asia (Guérin-Méneville) western and southern (Gestro) Sumatra (Baly) Java Schaeffer central United States (Baly) India ‘screwpine’ (possibly (Spaeth) (Maulik) Brazil (Horn) southern United States and (Smith) central and western

Uhmann Staines southern United States and (Olivier) southern Canada and most of sp. A. missionensis A. gracilis A. kansana A. nigrita A. bondari A. cyanea A. bicoloriceps A. bicolor Anisostena ariadne Acmenychus Taxon AuthorTaxon Acentroptera basilica Geographical Distribution Reported Host Plants Author FamilyReferences Selected Achymenus inermis Agonita bicolor A. decorata A. fossulata A. suturella A. undata Agonita A. pallipes A. fuscipes A. spathoglottis 54 J.A. Santiago-Blay 1928 et al. 1963 (illustr.) 1963 (illustr.) Linné, Anand 1989, Kalshoven 1957, Kimoto 1999 (illustr.), Maulik 1937; sp. and Abdullah and Qureshi 1969, Jolivet 1989a sp., Jolivet 1989a, Staines 2002b

sp., Sporobolus S. sp., Jolivet 1989a sp., sp. (larvae), Kalshoven 1957, 1981; Maulik 1919, 1937; Needham sp., , Pinanga sp. (Pandanaceae)(illustr.) 1963 Gressitt sp. A. Gray Staines 1994c, Thomas and Werner 1981 Paspalum Oryza Calamus sp. (Arecaceae) Gressitt 1963 (illustr.) Oryza ex sp., sp., Schizachrium sp. (Flagellariaceae), Linné (Poaceae) Gressitt and Kimoto 1963, Kalshoven 1957 Linné (larvae), Humboldt, Bonpland and Staines 1994b Bothriocloa sp., Freycinetia Saccharum officinarum unable to find name Gressitt 1963 (illustr.) Bentham Korthalsia sp. (Araceae) , ‘small palm with Gressitt 1960a (illustr.), 1963 (illustr.) Panicum sp. (Araceae), sp., sp. (Arecaceae) Gressitt 1963 (illustr.) Linné, Flagellaria Miscanthus Daemoronops Linné (larvae), sp., sp. (Poaceae) sp. (Poaceae) Linné (Poaceae) (larvae), ‘also … on (Arecaceae) Gressitt 1957, 1963 sp. (Pandanaceae) sp. (Flagellariaceae)(illustr.) 1963 Gressitt sp. (Poaceae) larvae Zaka-ur-Rab 1991 sp., ‘palm’ (Arecaceae)(illustr.) 1963 Gressitt sp. (Arecaceae) larvae(illustr.) 1963 Gressitt Tripsacum sp., sp., sp. (Arecaceae) larvae(illustr.) 1963 Gressitt sp., ‘rattan’ (Arecaceae)(illustr.) 1963 Gressitt sp. (Arecaceae)(illustr.) 1963 Gressitt sp. (Poaceae) sp., Tridens Dieffenbachia (Arecaceae), Freycinetia Dieffenbachia ‘rattan’Korthalsia Daemonorops Gressitt Saccharum officinarum Pinanga Saccharum officinarum Saccharum Olyra Valota Oryza sativa Saccharum Oryza sativa Korthalsia Korthalsia beccarii (Arecaceae) larvae Calamus Daemonorops Pinanga pinnae irregularly arranged’ (larvae), ‘small palms’, ‘rattan’ (Arecaceae) Flagellaria Calamus Korthalsia ‘rattan’ Gressitt spontaneum wild species of cane and bamboo’, on ‘other grasses’ (larvae) Bothrichloa Bambusa Panicum leucophaeum Kunth (Poaceae) larvae Acacia constricta sp., (west Papua) Mexico, El Salvador (Leguminosae) adults, Maulik Afganistan, Thailand, India Weise Brazil, Paraguay, Argentina Gestro New Guinea Gressitt New Guinea Gressitt New Guinea Gressitt New Guinea Gressitt New Guinea sp. Asia cuspidata (Horn) southwestern United States, Gressitt New Guinea ‘rattan’ larvae Gressitt 1963 (illustr.) Gressitt New Guinea

Gressitt New Guinea ? Gressitt New Guinea Gressitt New Guinea (Zehntner) Australia, Java Gressitt New Guinea ‘rattan’ (Arecaceae), Gressitt New Guinea sp. New Guinea sp. Canada to Argentina Gressitt New Guinea Gressitt New Guinea, Japen Island ‘rattan’ (Arecaceae) Gressitt 1963 (illustr.) Gressitt New Guinea ‘palms and rattans’ (Arecaceae)(illustr.) 1963 Gressitt Gressitt New Guinea Gressitt New Guinea ‘slender pinnate palms’(illustr.) 1963 Gressitt Uhmann Taiwan Gressitt New Guinea A. sedlaceki A. striata A. subviridipennis A. wilsoni Aspidispa A. rotanica A. rattana A. horni Asamangulia A. wakkeri Aspidispa albertisi A. bicolor A. daemonoropa A. maai A. palmella A. calami A. flagellariae A. papuana A. pinangae A. ifara A. korthalsiae A. lata Asamangulia Anisostena A. prompta A. perspicua Leaf-mining chrysomelids 55 1981, Ford and et al. 1928 (illustr.), Nicolay and et al. 1928 et al. Kogan and 1979, Needham Chittenden 1902, Frost 1924, Jones and Brisely 1925, Maulik 1937, Maulik 1937 Auerbach and Simberloff 1988, Balsbaugh Hays 1972, Chittenden sp. A. A. Walter, Loes. in malus Linné, Weiss 1918 (illustr.), Riley and Enns 1979, Robert 1947 Nee Walter West and Lothian 1948, Wilcox 1954

Linné Cavey 1985 (illustr.), Frost 1924, Hargrove 1986, Hodson 1942, Prunus C. (Miller) K. Citrus alba Urtica sp., Amelanchier

Glycine max Platymenia Malus C. integerrimus Q. acutissima Linné (adults), sp. agrifolia Michaux,

sp. (Rhamnaceae) Needham Cham., (Asteraceae), Monrós and Viana 1947(illustr.) Betula

Linné, (Linné) Osbeck Castanea crenata spp. (Tiliaceae), Prunus americana sp. (adults) 1902, Faeth and Simberloff (1981), Robinia pseudoacacia (Linné) Moench, Lamarck (Linné) Linné f. sp. adults (Salicaceae), Tilia Acer Quercus Bureau and K. Schumann Monrós and Viana 1947(illustr.) Linné (adults), Corylus americana Guaiacum sp. (Poaceae), A. nigrum Ostrya virginiana Ceanothus Sprengel (Malpighiaceae) Maulik 1937; Uhmann 1934 (illustr.), 1937 Salix incana Carpinus caroliniana Miller, (Swartz) Kuntze unable to find name,

Gray (larvae), Linné, probably C. sinensis (Linné) Persoon, sp., Linné (Rosaceae), sp. ‘white oaks’ adults DeCandolle Hippocratea griesebachi Linné, Phaseolus lunatus arbutifolia Olyra sp. (adults) (Leguminosae),

Linné, Cordia salicifolia Eupatorium agerateroides Alnus Kunth (Boraginaceae), Cassia nictatans Linné (adults), Linné, (Linné) Med. (adults), P. virginiana Q. nigra Carpinus P. malus Pyrus (Aiton) Willdenow, Quercus and Arnold, , sp. (adults), arbutifolia unable to find name Guazuma ulmifolia . Linné (adults), Robinia M. sylvestris

sp. (adults), sp. (adults) (Ulmaceae) negundo

(Zygophyllaceae) (Sterculiaceae) larvae, Dioclea divaricata (adults), aurantium Meibomia axillaris (Leguminosae), (Urticaceae), foliosa Banisteria argentea larvae, canadensis (adults), Ulmus Vernonia sororia (Fagaceae), (Linné) Merill, P. vulgaris Linné, Aronia Linné, Marshall, (adults) (Rutaceae), Tilia americana polystachya (Asteraceae), serrulata Carruth rubrum (Aceraceae), Ceanothus fendleri Arrabidaea coleocalyx (Bignoniaceae), Betula Corylus Koch (Betulaceae), probably Siebold and Zuccarini, (larvae), (adults), Acer Engler and Prantl Celastraceae) Bignoniaceae Paraguay, Argentina Bolivia Mexico Hooker States (Horn) southwestern United States and Weise Brazil, Paraguay, Argentina (Baly) Brazil (Baly) Costa Rica, Guatemala unidentified Bignoniaceae (larvae) Hespenheide and Dang 1999 (Chapuis) Caribbean? (unlikely), Brazil, Uhmann (Panzer) southeastern Canada and United B. productus B. schmidti B. parvulus Taxon AuthorTaxon Baliosus californicus Geographical Distribution Reported Host Plants Author FamilyReferences Selected B. conspersus B. nervosus B. duodecima 56 J.A. Santiago-Blay 1991, Jolivet 1989a, Staines 2002b et al. Staines 1986a Staines 1986a, Noguera 1988 sp., sp. sp. Jolivet 1989a; and Hawkeswood 1995; Mariau 1975 (illustr.), sp. sp. Cordia Alnus sp. sp. Staines 1986a sp., sp., (Greene), sp. Gillett sp. Ulmus Raphia sp. Guacoma sp. (Hooker) Molina sp. (Betulaceae), Malus Jatropha sp., Quercus Citrus Acer sp. C. sanguineus sp. (Urticaceae), sp. (Rosaceae), Artemisia sp., Hymenoclea Robinia sp. (Ehretiaceae), sp., Guaiacum sp. (Malpighiaceae), Desmodium Corylus (Bertoloni) Kuntze Hespenheide and Dang 1999 Urtica M. sativa Medemia K. Schumann (Rubiaceae), Rubus Greene, sp. (Asteraceae), sp., Douglas (Rhamnaceae) Ceanothus sp., Nuttall, sp. (Tiliaceae), sp., Cordia sp., A. Gray, unable to find name (Urticaceae) Hespenheide and Dang 1999 sp., Castanea sp. (Salicaceae), Banisteria Tilia Ameliancher D. Don, Holocarpa heermannii Linné, sp. (Rubiaceae), (Asteraceae) (adults) Riley and Enns 1979, Staines 1986a sp. (Acanthaceae), sp. (Malvaceae) Monrós and Viana 1947 Ceanothus cuneatus Meibonia Urera Vernonia sp. (Celastraceae), Bauhinia Pyrus Salix Carpinus sp. and other bignoniaceans, sp., Torrey and Gray (Asteraceae) (both N. L. Burman (Malvaceae) adults, sp., sp. (Arecaceae) Mariau 2001 sp., sp., sp. (Verbenaceae), C. velutisinus sp. (Poaceae), (all adults) Franseria dumosa monogyra adults) Erigeron sp. Artemisia tridentata (Asteraceae) (both adults) Sphaeralcea (Ulmaceae), (Sterculiaceae), (Arecaceae) 1988 Hemizona (Asteraceae), Rhamnaceae (all adults) Staines 1986a Media elegans Ceanothus lucodermis Lippia (Zygophyllaceae) Cocos Dioclea Prunus Basanacantha (Rutaceae), Cocos nucifera (Leguminosae), Sida acuta Olyra (Rhamnaceae), (Euphorbiaceae), (Fagaceae), Basanacantha spinosa (Aceraceae), Hippocratea Urera bogitaense larvae Odontonema tubaeforme (Acanthaceae) larvae Odontonema sp. (Boraginaceae), Betula Arrabidaea western United States Pursh, Mexico Rocky Mountains Madagascar Brazil, Paraguay, Bolivia,Argentina ‘Guayabo silvestre’ Monrós and Viana 1947(illustr.) Van Dyke western United States Berti and (Crotch) eastern and central United States Pic Bolivia, Paraguay, Argentina (Baly) Costa Rica, Panama, (Horn) Canadian and United States Weise western United States and (Crotch) southwestern Canada and sp. 1’ Costa Rica sp. 3’ Costa Rica sp. New World sp. west Africa, Madagascar Baliosus Baliosus B. longula B. melsheimeri B. montana B. notaticeps Brachycoryna dolorosa B. hardyi Balyana mariaui Desmier de Chenon Balyana B. viridanus ‘ ‘ Baliosus Leaf-mining chrysomelids 57 1928, Riley and Enns 1979 . 1993, Staines 1996, Uhmann 1934 (illustr.) 1991; Maes 1998; Moldenke 1971; Noguera 1988 et al et al. et al. Moldenke 1971, Staines 2002b Hespenheide and Dang 1999, Jolivet 1989a, Maulik 1937, Staines 2002b (illustr.), Uhmann 1934 1937 (larvae) Memmott Butte 1968b (illustr.), Chittenden 1902, Ford and Cavey 1985 Needham L. sp. Linné sp. Maes 1998, and Staines 1991, Moldenke 1971, Noguera 1988 sp., sp. sp. Gillett ) sp. (Gill. sp. Bunchosia Panicum Malpighia Sida Linné (all Davidse, Abelmoschus (Linné) sp. Butte 1968b (illustr.), Maes Sida (Linné), Alcea rosea P. oligosanthes Cervantes, (illustr.); Staines 1986a, 1991, 1996 (illustr.) Bauhinia Baccharis (Kunth) sp.), sp., Humboldt, Oryza sp., P. microcarpum Waltheria Linné, S. rhombifolia Abutilon lignosum Dicanthelium N. L. Burman (larvae), Abutilon americanum Schott (Araceae) adults Moldenke 1971 ( Phaseolus vulgaris Waltheria americana Rose (larvae), Lasiacis Malvastrum coromandeli- sp. (Leguminosae) Jolivet 1989a Lamarck, Abutilon Aloysia gratissima Lasiacis nigra Linné, Abutilon Linné (larvae), A. americanum Humboldt, M. americanum L. ruscifolia S. spinosa sp., Panicum sp. (Leguminosae), Linné (larvae), Gossypium hirsutum Sida acuta A. peduncalarae Galactia P. nitidum sp. (Malvaceae), Monarda citridora sp., sp., Linné (adults), (Linné) Moench, Hackel, sp. (Leguminosae), sp. (larvae), sp. (Asteraceae), sp., sp. (Zingiberaceae)1989a Jolivet sp. (Zingiberaceae) larvae Maulik 1937, Kalshoven 1957, Stanes 2003b sp. (Lamiaceae), Gossypium Sweet (adults), Alcea , Phaseolus Malpighia glabra (Linné) Garcke, sp. (Poaceae) adults, Sphaeralcea Linné (Sterculiaceae) adults Artemisia (Sterculiaceae) Bunchosia costaricensis esculentus (Malvaceae) previous five host plants of larvae, Zea sp. (Malpighiaceae) larvae (Leguminosae) adults, Linné (Cavanilles) G. Don, Sweet (adults), Bonpland and Kunth, S. cordifolia (adults and larvae), Dunbaria Buhinia ungulata Monarda Curcuma Curcuma and Hooker) Troncoso (Verbenaceae) adults Chusquea Hitchcock (larvae mine procerrima (Poaceae) Philodendron anisostomum Phaseolus Panicum clandestinum Linné, Malvastrum Baccharis thesioides Muhlenberg, adults), nus Schultes (adults), (Poaceae) Argentina (Linné) southern United States to Nicaragua Colombia, also in Jamaica (Asteraceae), Baly tropical Africa, Asia, Australia Chapuis Mexico, Belize, Guatemala,

Uhmann] sp., sp. sp. Asia (Newman) southeastern United States ‘everglades grasses’ Butte 1968b (Chapuis) Mexico, Guatemala, Nicaragua sp. Congo, China, Vietnam Baly Mexico to Colombia Jacoby Mexico Guérin-Méneville southern United States to (Olivier) eastern half of the United States sp. [monotypic genus, Central America Brachycoryna Carinispa C. nevermanni Cassidispa Chalepus acuticornis Chaeridiona metallica Chaeridiona Taxon AuthorTaxon B. pumila Geographical Distribution Reported Host Plants Author FamilyReferences Selected C. amabilis C. amicus C. bacchus C. bellulus (data from several subspeciesincluded) C. bicolor (Poaceae) both adults 1998, Maes and Staines 1991, Noguera 1988 (illustr.) 58 J.A. Santiago-Blay 1993 et al. 1977 et al. 1993 et al. Staines 2002b, Ward Ford and Cavey 1985 (illustr.), Thomas Werner 1981 Monrós and Viana 1947 (illustr.) Maulik 1937, Sanderson 1967, Virkki and Santiago-Blay 1998, Wilcox Memmott Hespenheide and Dang 1999, Memmott L. L. sp. sp. sp., Bureau Hystrix Olyra sp. Meas 1998; Maes and Staines 1991; Uhmann 1934 (illustr.), 1937 Betula sp. Hackel, Hackel, sp. Centaurea sp. (Araceae), Jolivet 1989a, Maes 1998, and Staines 1991, Passoa 1983, Actinostemon (Poaceae) adults Monrós and Viana 1947 (illustr.) Verbesina

(Linné) Chase Commelina Paspalum densum Panicum Chusquea Bromelia Hitchcock, 1975 sp., Muehenberg, sp. (Convolvulaceae), Swartz (Poaceae) all Monrós and Viana 1947 (illustr.) sp. (Celastraceae), sp. (Bignoniaceae), L. procerrima L. procerrima Lamarck Bert. (Verbenaceae) all Philodendron Kunth, sp. (Asteraceae), Urban (Asteraceae) adults, Hespenheide and Dang 1999, Moldenke 1971 Rose (Tiliaceae) Noguera 1988 M.T. Masters. Monrós and Viana 1947 (illustr.) (Linné), unidentified Poaceae (Humboldt, Bonpland, and (Linné) Nees. (Poaceae) unable to find name Monrós and Viana 1947 (illustr.) Ipomoea Cham. (Boraginaceae), sp. (larvae), , Arrabidaea coleocalyx Eupatorium Celastrus Valota insularis Davidse, Davidse, Wedelia Arrabidaea sp., sp. (Aristolochiaceae), Elymus villosus Vitex cymosa sp. (Combretaceae), sp. (Sterculiaceae), sp. (Leguminosae) adults, flowers of Maes 1998, and Staines 1991, Passoa 1983 Panicum molle (Kunth) Hitchcock (Poaceae) (Kunth) Hitchcock (Poaceae), sp., nigra sp. (Brassicaceae), sp., Lasiacis

Sorghastrum setosum sp., Moench (Poaceae) sp. (Aceraceae), Cyanus (Euphorbiaceae) all adults sp. (Poaceae) Valota insularis unidentified species of Leguminosae (adult) Noguera 1988

Brassica (Bromeliaceae), Terminalia (Commelinaceae), (Betulaceae), Aristolochia sp., Vernonia Acer Phaseolus Zea Bromus patula adults Guazuma (Poaceae) (larvae), Aristolochia elegans (Aristolochiaceae) and K. Schumann (Bignoniaceae), sp. (Poaceae), Trichachne insularis adults Bromelia caragua ruscifolia ruscifolia unidentified Tiliaceae (larvae) Lasiacis Olyra adults Cordia salicifolia Heliocarpus pallidus Paspalum quadrifarium sp., Panicum leucophaeum (Poaceae) Kunth) Alefeld (Leguminosae) adults, unidentified Poaceae (larvae) Lasiacis nigra Verbesina greenmani Benthamantha mollis Paraguay, Argentina (Bromeliaceae), West Indies, South America Poiret, Florida (United States), Uhmann southern Brazil, Bolivia, Baly Mexico Uhmann Paraguay and Argentina Pic Bolivia, Paraguay, Argentina (Chapuis) Mexico, Guatemala, Nicaragua Baly Mexico an (Chapuis) Brazil, Paraguay, Argentina Baly Mexico, Costa Rica Uhmann Nicaragua and Costa Rica Baly Mexico, Guatemala (Chapuis) Brazil and Paraguay (Crotch) United States sp. Canada to Argentina Baly Costa Rica C. verticalis C. walshii Chalepus C. schmidti C. subcordiger C. sanguinicollis australis C. horni C. parananus C. hepburni C. placidus C. putzeysi C. cordiger C. consanguineus C. digressus C. sanguinicollis (Linné) Leaf-mining chrysomelids 59 1986 et al. (illustr.), 2001; Mariau and Morin 1971, 1974; 1971 (illustr.); Maulik 1931 Morin and Mariau 1971; Uhmann 1968 Bernon and Graves 1979; Berti Mariau 1999; Chen Maulik 1937 sp. Maes 1998, and Staines 1991, 2002b (illustr.) sp. Berti and Mariau 1999, Collart 1934, Jolivet 1989a, 1988 sp., Monrós and Viana 1947 sp. sp., sp., sp. sp., Vitex Apios Pueraria sp., sp., Zea sp., Glycine Malus sp. Coffea Olyra sp., sp. (Poaceae) Jolivet and Hawkeswood 1995, Staines 2002b Roystonea Theobroma sp., sp., Cassia sp., sp., Cymbosema Mucuna Desmodium sp., Desmodium sp., sp. (Poaceae) Staines 2002b sp., Wendland Ramos 1996 (illustr.) Aloysia Chusquea sp., sp., Olyra sp. (Rubiaceae) Valota ex. sp., Zea Prosopis sp., Falcata sp., Actinostomon sp. (Fagaceae), Elaeis sp., Lasiacis Elaeis guineensis sp., Crataegus sp., Berg. (Poaceae), Bentham, Coffea sp. (Leguminosae), sp. (Zingiberaceae) sp., sp., sp., Vogel (Leguminosae) sp., Canavalia sp., Benthamantha Meibomia Schrader Quercus Jacquin (Arecaceae) Collart 1934, Lepesme 1947, Mariau 2001, Uhmann 1931 Jacquin (Arecaceae) Jacquin (Arecaceae) Berti and Mariau 1999 (illustr.), 2001 2001 1988, Mariau Vicia sp. (Sapindaceae), Cymbosema Cymbosema Bambusa sp., other palms (Arecaceae) (illustr.); Cox 1996 Lepesme 1947 Mariau 1988 sp., Brachiaria Cocos sp. (adults), Paspalum Amonum sp., Cocos sp., Paspalum Dolichos sp., Hystrix sp., sp.) Cerasus (Arecaceae) Mariau 1988, 2001 sp., sp.,

sp., sp., Elaeis sp., sp., sp., Paullinia sp., sp. sp. (Ehretiaceae), sp. (Poaceae) Jolivet 1989a sp. (Rosaceae), Bauhinia Lathyrus Robinia Amomum Cocos Elaeis guineensis Elaeis guineensis Elaeis guineensis (Staines 2003b has been unable to confirm record of (Arecaceae), (Euphorbiaceae), sp., Bambusa Oryza Jacquin, Borassus Crotularia Cordia Dioclea Elymus sp., Calopogonium sp., Pithecelobium Panicum (Poaceae), sp. (Sterculiaceae), Tiliaceae, Pyrus Panicum Bambusa vulgaris Borassus adults, sp. (Verbenaceae) Paspalum conjugatum (Rubiaceae) adults (Poaceae) Centrosema (Leguminosae), Leguminosae Centrosema pubescens Desmodium discolor French to Ghana) Madagascar Africa, Madagascar Colombia to Argentina Costa Rica to Argentina Guyana, Suriname, Guyana, Brazil, Peru Colombia to Peru ) Maulik west and central Africa (Fabricius) Trinidad, Tobago, Colombia, 5-6 Maulik Brazil (Fabricius elaiedis Maulik Argentina

sp.

Berti and Mariau Ivory Coast Gestro Zaire

sp. sp. Sierra Leone (Africa) Uhmann west Africa (from Cameroon Fairmaire sp. C. speciosa Coelaenomenodera C. minuta C. perrieri Chrysispa C. lameensis Taxon AuthorTaxon Geographical Distribution Reported Host Plants Author FamilyReferences Selected Clinocarispa humeralis Charistena ruficollis Charistena Coelaenomenodera Cnestispa acuminata Cnestispa darwini Cnetispa 60 J.A. Santiago-Blay 1986 (illustr.), Gressitt and Kimoto 1963 1985, Kalshoven 1957, Nakane 1955 (illustr.), Tan 1993 et al. et al. Cox 1996 (illustr.), Maulik 1932 (illustr.) Kalshoven 1957 Chen Staines 2002b De and Konar 1954, Kimoto 1999 (illustr.), Zaka-ur-Rab 1991 (illustr.) A. sp. Jolivet 1989a F. sp., Blume., An sp. Kalshoven 1957 Ohwi, ‘bamboo’ Gaertner, Kalshoven 1957

Sorghum sp. Gressitt 1960a (illustr.), and Samuelson 1988 (illustr.) Sterculia Linné Maulik 1937 Saccharum ex sp. (‘he- Linné Prunus Isodon ) Maximowicz, . glabre sp. (Pandanaceae) sp. (Poaceae) Maulik 1937 Hymenache (larvae),

Miller, Thoracastachyum Prunella vulgaris Filipendula Q. myrsinaefolia sp., Linné (Poaceae) larvae Kalshoven 1957 ) both larvae

Oryza sativa Mitter var Pandanus (Linné) Brotero, sp. (Arecaceae) Gressitt and Samuelson 1988 (illustr.) Panicum Schultes, Schultes Thoracastachyum Lantana camara (Blume) Bentham and Hooker Schultes (Poaceae) larvae Kalshoven 1957 Beauvois (Poaceae) Collart 1934; Maulik 1932 (illustr.), 1937 Carr, Merrill (Rubiaceae) adults Gressitt 1957, Kalshoven 1957 Nakai, (Lamiaceae), Banks and Soland. Phyllostachys bambusoides Scleria sp. (Malpighiaceae Malus pumila Linné, sorghum Linné (adults) (Poaceae), sp. (Cyperaceae) Cox 1996 (illustr.); Gressitt 1957 (illustr.), 1960a sp.,

sp. (Poaceae sp. (Poaceae) Staines 2002b lilacina Filipendula multijuga (Pallas) (adults), (Thunberg) Kudo, Persoon (Poaceae) sp. ‘le genus’, (Fagaceae), Rivinus, Metroxylon sp., ‘sedge’ (Cyperaceae) Gressitt 1957 (illustr.), 1960a (illustr.) sp. (Rosaceae) sp. (seedlings) larvae, ‘glagah’, ‘lalang’ Kalshoven 1981 Kurz (Rubiaceae) ex Plectronia horrida f. Nertera depressa Stigmaphyllum Saccharum Saccharum Hypolytrum Scleria vulgare Linné var. (Poaceae), Gardenia augusta Saccharum spontaneum Zea adults (Poaceae) Bambusa blumeana spontaneum (Poaceae) Bambusa blumeana Arundinaria pygmaea simonii Siebold and Zuccarini, (Verbenaceae) Bambusa blumeana (Poaceae) both larvae Oryza sativa Melinis minutiflora (Cyperaceae) Hypolytrum ‘sur cacaoyer’ Andropogon Quercus acutissma ye-zi genus’), Rosa Uhmann 1968 Quercus inflexus palmata sp. (Sterculiaceae) adults Island) Java Trinidad Java Thailand, Laos, Vietnam Java Maulik (Klug) Zaire Gressitt Solomons Islands (Guadalcanal) Gressitt Malaita (Solomons Islands) ‘palm’ (Arecaceae), (Duvivier) ‘United Provinces’, India sp. Trinidad to Brazil (Guérin-Méneville) Philippine Islands (Gestro) India ‘grasses’ (Poaceae), Gressitt Solomon Islands (Santa Isabel ‘palms’, (Gestro) India, Burma (or Myanmar), (Chapuis) Philippine Islands (Solsky) Korea, Japan, China, Siberia Guérin-Méneville Philippine Islands, Java (Gestro) Ethiopia, Uganda, west Africa Gressitt Solomon Islands (Guadalcanal) Gressitt New Guinea ‘large-leaved shrub’ Gressitt 1963 (illustr.) sp. Brazil sp. Solomon Islands (Gestro) (Gestro) (Gestro) Chûjô Japan, China Gestro Java Corynispa Craspedonispa Cyperispa hypolytri C. palmarum C. scleriae D. bakeri D. balyi D. bipartita D. issikii D. discalis D. infuscata D. chapuisi D. cladophora D. debilis D. dilaticornia Craspedonispa saccharina (includes two subspecies listed in Gressitt 1960a) C. thoracostachyi (including two subspecies listedin Gressitt and Samuelson 1988) Cyperispa Dactylispa aculeata (Cyperaceae) larvae, D. albopilosa D. angulosa D. aspera D. brachycera Leaf-mining chrysomelids 61 1986 (illustr.), Gressitt and Kimoto 1986 (illustr.), Nakane 1955 (illustr.) et al. et al. 2001a 1986 (illustr.), Gressitt 1957, Kalshoven Kimoto 1985, Gressitt and Kimoto 1963 1985, Chen et al. et al. et al. et al. Cox 1996 (illustr.), Gressitt and Kimoto 1963, Kalshoven 1957 Kalshoven 1957, 1981 (illustr.); Maulik 1937; Staines 2003b Uhmann 1968 sp. Linné sp. B. sp. Saccharum Cinchona Thunberg, Cinchona McClure, (illustr.) sp. (Zygophyllaceae) Maulik 1937 (Makino), Lingmania Curcuma S. spontaneum Q. glauca sp. (Poaceae) Uhmann 1968 (Thunberg) Schottky, 1963, Hayashi 1986 Lingmania cerocissima Trimen , J. König (larvae), 1999 (illustr.) McClure, larvae Kalshoven 1957 (Linné) Beauvois (adults), Chen Carr, (Loureiro) Raeuschel, ex. Linné f. (larvae), Linné (larvae) (Rosaceae) Kalshoven 1957 (Thunberg) Makino (Poaceae) Gressitt and Kimoto 1963, Kalshoven 1957 (F.Schmidt) and other An Guaiacum Linné (Sterculiaceae) larvae1968 1931, Uhmann Stapf (Poaceae) larvae Uhmann 1968 Linné (larvae) (Poaceae) Siebold and Zuccarini, An (Linné) Gaertner (Bombacaceae) Kalshoven 1957, 1981 (illustr.); Kimoto 1999 (illustr.) sp. (Sterculiaceae) larvae B. subspinosa sp. (Lytraceae) adults, sp. (larvae) Poaceae sp. (Anacardiaceae) Gressitt 1963 (illustr.) Linné (Poaceae) Banwo Linné (larvae), Anropogon Moens spp. (larvae), Munro, sp. (Poaceae) Cox 1996 (illustr.), Maulik 1932 Uhmann 1968 sp. (Rubiaceae), ‘salam utan’, Cox 1996 (illustr.), Kalshoven 1957, Maulik 1931 (illustr.) Zea mays , L. chungii Sinobambusa tootsik Helicteres sp. (Rosaceae) Gressitt and Kimoto 1963, Maulik 1937 aceae Asteraceae Panicum Rubus moluccanus Setaria chevalieri Rubus Ceiba pentandra Anthraxon hispidus larvae Lagerstroemia Oryza sativa ledgeriana Gardenia ‘kerema’ (Myrtaceae?) all larvae seedlings (Rubiaceae) larvae, (Zingiberaceae) adults Petasites japonicus Theobroma cacao McClure (larvae), ?Sinobambusa ? Semecapus Bambusa multiplex Bambusa officinarum (adults), Castanea crenata tuldoides Rottboellia exaltata Quercus acutissima Castanopsis cuspidata Imperata cylindrica Panicum palmifolium Nigeria, Uganda, Java India, southern China Vietnam, China, Sumatra adults, Tanzania Sumatra, Java Java Congo, Fernando Poo, Spanish Guinea, Nigeria, Natal (South Africa) Sulawesi (= Celebes Island) China, Sumatra, Java, Borneo, Thailand, Laos, Vietnam, (Baly) Korea, Japan, China Gressitt New Guinea Gestro Democratic Republic of (Motschulsky) Guinea, Gabon, west Africa, Maulik India ‘plum’ (Rosaceae), (Gestro) (Chapuis) tropical Africa Commelinaceae Uhmann 1931, 1968 (Gyllenhal) tropical Africa Cyperaceae, Uhmann China (Gyllenhal) tropical Africa Po Maulik eastern Himalayas, northern (Ritsema) Thailand, Cambodia, Laos, (Gestro) Malacca (Malaysia), Maulik (Kraatz) Tropical Africa (Weber) India, Burma (or Myanmar), Gestro China ‘bamboo’ (Poaceae) Gestro Korea, Japan, China, Siberia Weise Uhmann China D. nemoralis D. puncticollis D. melanaria D. pallipes D. parbatya D. pubicollis D. leonardi D. luhi D. manterii D. kamarupa D. masoni D. semecarpus D. sjoestedti D. spinigera D. spinulosa D. subquadrata D. kaulina D. lenta Taxon AuthorTaxon D. javaensis Geographical Distribution Reported Host Plants Author FamilyReferences Selected D. spinosa 62 J.A. Santiago-Blay 1986 et al. 1985; Chen et al. 2001b, Collart 1934, De and Konar 1954, Jolivet et al. 1989a, Staines 2003b, Uhmann 1953 1980; Razzaque and Karim 1989; Zaka-ur-Rab 1991 (illustr.); Cox 1996 Kalshoven 1957, 1981 Kimoto Abdullah and Qureshi 1969; An sp., Malus sp., sp., sp. sp., sp. sp., sp. Banwo sp., Gardenia sp., Ceiba sp., sp. Mnesithea sp., sp. sp., sp., Digitaria sp. Plectranthus Hyparrhenia Leersia sp., sp., Eleusine Zizania Echinochloa (Linné) 1999 (illustr.); Maulik 1919 (illustr.), 1931, 1937; Rawat and Singh Phoenix Arundinaria Q. serrata Phaseolus sp. Scleria Castanopsis Panicum sp. (Rosaceae) Linné (larvae), sp., Rottboellia sp., Lobelia Dalbergia Leptochloa sp. sp., sp., Coffea Theobroma Roth, Urelytrum sp., Sinobambusa sp., Petasites sp. Callipedium sp., Melinis Leucosidea sp. (Rubiaceae), sp., Gaertner, sp., sp., Hevea sp., sp. (Verbenaceae), Rubus Zea sp., Isodon sp. (Onagraceae), sp., Linné (Poaceae) Kalshoven 1957 sp., Blume (Fagaceae) sp., sp., sp., sp., sp., sp. (Betulaceae), sp., Oryza E. crusgalli sp., Digitaria sp., Nertera (Blume) Bentham and Hooker Kalshoven 1957 Hymenachne Artemisia Commelina Linné (Cyperaceae), Leersia Castanea Oryza sativa Anthrascon Piliostigma Jussiaea D. setigera Setaria Rosa sp., Callicarpa sp., Sterculia Triticum Pennisetum Cinchona sp., Croton Cynodon Vossia Loudetia Phyllostachis sp., Fischer ex Turczaninow ( var. Rehd and Wilson, Carpinus sp., sp. (Lamiaceae), sp., sp. (Anacardiaceae), sp., sp. Filipendula Q. variabilis sp., sp. (Bombacaceae), Swartz, (Zygophyllaceae). Apparently, also in sp., sp., sp., Eleusine indica sp., sp., sp., sp., sp. (Fagaceae), sp. (Rosaceae), ‘almond leaf’ Maulik 1937 Persoon, (Linné) Link, Kurz (Rubiaceae) larvae, ‘tauluan’ Prunella Plectronia Imperata Oplismenus Durio Prunus ex Q. mongolica grosseserrata Thunberg, Saccharum officinarum (Euphorbiaceae), Quercus sp., Desmodium (Leguminosae), Andropogon Bambusa Dactyloctenium Enchinochloa sp., Lignania sp., (Asteraceae), sp., (Cyperaceae), Prunus Plectronia horrida f. (Rubiaceae?) larvae (Arecaceae), Semecarpus Paspalum Saccharum Sporoblus (Campanulaceae), (Commelinaceae), Vetiveria (Poaceae), sp., Canthium hexandra colona sp., Helicteres ciliaris (Sterculiaceae) Guaiacum some Acanthaceae, Malvaceae, and Marantaceae chichona, kapok, coffee, and maize Cyperus rotundus Sumatra, Java, Indonesia China, Taiwan, Malaya, Beauvois, Thailand, Laos, Vietnam, Burma (or Myanmar), (Olivier) Pakistan, India, Nepal, Weise Java Maulik India sp.World Old Gestro Java D. sumatrana D. vestita D. vethi Dactylispa Dicladispa armigera Leaf-mining chrysomelids 63 et al. 1986 (illustr.), Cox 1996 Jolivet 1989a 1957, 1960a et al. Samuelson 1988 (illustr.) Bordy 2000 (illustr.); Buhr 1955; Cox 1996 Grandi 1959 ? C. sp. Jolivet 1989a Oryza sp., sp. Chen e sp., sp. Zea Linné, sp., Zea mays Aralia sp., Eleusin (Poaceae) Gressitt 1960a, 1963 sp. (Lobeliaceae),

sp. (Asteraceae), Jolivet 1989a, Staines 2003b, Uhmann 1953 sp. (Poaceae), Saccharum Dactyloctenium Rhynchosia sp., Sinocalamus Mnesithea Grewia sp. (Malvaceae), sp., Sorghum sp., ‘palms’ Gressitt 1957 (illustr.), 1960a 1963 (illustr.); and sp., sp., Zizania sp., Linné (Poaceae) adults Kalshoven 1957 sp. (Cistaceae) (illustr.); Hering 1957 Maulik 1919, 1937; Needham ? sp. (Zingiberaceae) Gressitt 1957 (illustr.), 1960a, 1963 (illustr.); Staines 2003b Petasites sp., Linné (larvae), sp., Lobelia sp. C. monspeliensis sp., sp. (Poaceae) larvaesp. (Poaceae) larvae Maulik 1937, Kalshoven 1957 Kalshoven 1957 sp. (Zingiberaceae) Echinochloa Cistus Paspalum Cynodon Costus sa sp.) Poaceae1993 Yu Malvastrum ‘and other grasses’ Dalbergia Linné, Vossia Daemonorops sp., sp. (Poaceae) larvae Kalshoven 1957 Saccharum sp., Leptochloa Costus sp., sp. (Leguminosae) Uhmann 1953 Pennisetum sp. (Rosaceae), sp., Linné (Poaceae) Abdullah and Qureshi 1969, Maulik 1937 Linné, sp., sp. (Poaceae) larvae, ‘grass’, Bambu sp., Bambusa Bambusa sp., (Arecaceae) larvae Maulik 1937, Kalshoven 1957 sp. ‘sorghum’ (Poaceae) Anand 1989 sp.,

sp. (Araliaceae), sp. (Cistaceae), sp. (Cistaceae) Hering 1957 sp., Digitaria Panicum Paspalum sp. (Araliaceae), Crotalaria Aralia Cistus Crotalaria (Leguminosae), sp., Leersia sp., Leucosidea (Arecaceae) Calamus (Arecaceae) ‘grasses’Cistus Zea mays Cistus albidus salvifolius Gressitt (Poaceae) ‘bamboo’ ‘bamboo’ Sinocalamus bamboo Saccharum spontaneum Sorghum Oplismenus sp. Callipedium Vetiveria (Tiliaceae), Oryza Bambusa Saccharum officinarum Linné and other Poaceae, ‘paddy’ New Guinea) Africa, Turkey, Syria 1928 Bougainville Islands, Algeria, northern Java India region on east), Burma (or Myanmar) Gressitt Bismark Archipelago, Maulik Java (Guérin- Weise China (Gestro) east Africa (Motschulsky) Gestro Java, Sumatra bamboo (Arecaceae) larvae Maulik 1937, Kalshoven 1957 Spaeth Java (Chapuis) (Péringuey) southwest Africa Gressitt New Guinea Gressitt China, Vietnambamboo ( Vietnambamboo Gressitt China, sp. Africa sp. Old World (Guérin-Méneville) New Guinea, Brittain, (Linné) southern Europe, Canary Weise Java (Brullé) Canary Islands (Spain) sp. (some species are miners) Indo-Australian region bellicosa (Weise) New Guinea (Chapuis) India (including Assam ‘leaves of apple tree’ (Rosaceae) Maulik 1937

D. D. linnei D. occator D. vicinalis Dicladispa in Gressitt and Samuelson 1988) Enischnispa calamivora (includes two subspecies listed( Ireland New D. kapauku D. striaticollis D. testacea Taxon AuthorTaxon Geographical Distribution Reported Host Plants Author FamilyReferences Selected D. vandykei [prob. Méneville)] Downesia bambusae D. javana D. marginicollis D. perniciosa D. sumatrana D. cyanipennis D. dama D. fabricii Dorcathispa Downesia D. alternata 64 J.A. Santiago-Blay 1928, Riley 1985 (illustr.) et al. Riley 1985 (illustr.), Noguera 1988 sp. Jolivet 1989a and Jolivet 1989a, Riley 1985, Staines 2002b

’ sp. Gressitt 1963 (illustr.) sp., Kalshoven 1957, Staines 2003b, Uhmann 1955 (illustr.) Digitaria sp., sp. Kalshoven 1957, Maulik 1937 (Poaceae), (Fournier) Roxburgh Kalshoven 1957, Staines 2003b

(Zingiberaceae) Gressitt 1963 (illustr.)

Alpinia ‘rattan with Gressitt 1960a (illustr.) (Arecaceae) Jolivet 1989a

sp. Hespenheide and Dang 1999, Riley 1985 Dictyosperma

spp. Nicolaia sp. sp. sp., sp., Amomum sp., unidentified Panicum Mimosa laxiflora (?)’ (Cyperaceae) larvae Gressitt 1960a (illustr.) sp., Costus (Leguminosae), Digitaria

Mimosa? Cocos Schumann (Marantaceae) larvae(illustr.) 1963 Gressitt Linné (Poaceae) (larvae) andNeedham 1985, Cavey and Ford Valota sp. larvae, sp. (Pandanaceae) larvae(illustr.) 1996 Cox sp. Scleria sp., Eriochloa gracilis Zingiber cassumunar sp. (Arecaceae)(illustr.) 1963 Gressitt Torrey (Ulmaceae) Erichloa Daemonorops Daemonorops Merrill, sp. (Pandanaceae) larvae(illustr.) 1960a Gressitt sp. (Arecaceae) Kalshoven 1981 Scopoli, sp. (Asteraceae) Staines 2002b sp., sp. (Asteraceae) larvae Maulik 1937, Uhmann 1934 sp. (Cyperaceae) Gressitt 1963 (illustr.) Mimosa Amomum sp., sp., sp. (Zingiberaceae)(illustr.) 1963 Gressitt sp. (Zingiberaceae) larvae Gressitt 1957, Kalshoven Staines 2003b sp.,

sp. (Zingiberaceae) sp. (Zingiberaceae)(illustr.) 1963 Gressitt ? sp. (Poaceae) sp. (Zingiberaceae)(illustr.) 1963 Gressitt sp. (Zingiberaceae) sp. (Zingiberaceae)(illustr.) 1963 Gressitt (illustr.) 1963 Gressitt sp. and other gingers (Zingiberaceae) Gressitt 1963 (illustr.) sp. (Musaceae), Freycinetia Alpinia Alpinia Acanthophoenix sanguinalis Celtis pallida Digitaria Daemonorops Calamus possibly other members of the Poaceae Calamus Vernonia Vernonia Freycinetia Glycine max (Leguminosae), Panicum capillare Valota Alpinia Scleria slender pinnae’ (Arecaceae) poacean (larvae) (Poaceae) (Zingiberaceae) larvae Hitchcock (adults), Metroxylon Curcuma Zingiber Alpinia Musa Alpinia Alpinia Orchidaceae, and probably (Zingiberaceae) larvae (Zingiberaceae) Donax canniformis Amomum probably Asia, Indo-Australian regionReunion small (Arecaceae) United States, Mexico Bentham New Guinea Java Moluccas (Weise) Arizona (United States) Gressitt Solomon Islands Gressitt Biak island (New Guinea) ? Weber Sumatra, Sunda Islands, Uhmann Central America Gressitt New Guinea

Gressitt Gressitt Admiralty Islands (New Guinea) ‘small sedge, near Gressitt New Guinea ‘smooth-leaved ginger, possibly Gressitt New Guinea ? sp. United States to Costa Rica Gressitt New Guinea sp. Gressitt New Guinea ‘small palm’ Gressitt 1963 (illustr.) Baly Java (Smith) southern and western a species of Bombacaceae, Gressitt New Guinea Gressitt New Guinea sp. New Guinea sp. Madagascar, Mascareignes- sp. Mexico to Colombia Gressitt New Guinea Spaeth Java (Gestro) Baly Java Gressitt New Guinea Gressitt New Guinea (Newman) eastern United States Gressitt New Guinea Gestronella Glyphuroplata E. daemonoropa E. palmicola E. rattana Euprionota gebieni Euprionota Freycinetispa collinsi Freycinetispa G. uniformis Uroplatini reported to mine grasses and not broad leaf plants) Gonophora biakana G. scleriae G. semiviridis Glyphuroplata nigella (Only reported genus of the Enischnispa G. pluto G. bicolor G. cubicularis G. maai G. musae G. puncticollis G. bowringii G. pellucida G. sinuicosta G. taylorii G. haemorrhoidalis G. cyperaceae G. donaxiae G. integra Leaf-mining chrysomelids 65 1986 (illustr.), Collart 1928, Grandi et al. 1984, Maulik 1937, Medvedev and Zaitzev Jolivet 1989a, Staines 2002b Bordy 2000 (illustr.), Chen Maulik 1937; Noguera 1988; Staines 1996; Uhmann 1934 (illustr.), Hespenheide and Dang 1999; Maes 1998; Staines 1991; Maulik 1937; Kalshoven 1957, 1981 (illustr.); Staines (2003b); Uhmann 1955 (illustr.) Hespenheide and Dang 1999, Maes 1998, Maulik 1937, Uhmann 1934 (illustr.), 1937 (illustr.) sp. Staines 2002b sp. sp., 1959 (illustr.), Gressitt and Kimoto 1963, Hering 1957, Kaszab sp. sp., Jolivet 1989a sp., 1962, Koch 1992, Lopatin Vanda Nicolaia (Linné) Maulik 1919, Monrós and Viana 1947 (illustr.) sp., sp., Hooker, Apeiba is Nicolaia spp., 1978 Avena C. Sphaeralcea Spruce ex sp., Mimosa Guazuma Phalaenopsis sp., sp., Poa sp. 1937 Serjania Inga edulis sp., sp., Alpinia sp., Serjania sp., Phalaenopsis , Sphaeralcea sp., Helicotrichon Stenotaphrum Sida rhombifolia V. tricolor Sida Polak. (larvae), Agropyron repens Elettaria Linné, sp. (Sapindaceae) Machaerium sp., sp., Agrostis Amomum Spathoglottis sp., sp. (Orchidiaceae), Linné Guazuma sp., Cassia Arundina sp., sp. (Malvaceae), Lindley, Machaerium sp. (Sterculiaceae), Serjania sp. (Poaceae), sp. (Poaceae) Maulik 1937 Vanda Beauvois, ex sp., Miller (Leguminosae), Panicum sp. (larvae), Apeiba membranacea Sida sp. (Tiliaceae) (Linné) Gould, Dendrobium Curcuma sp. (Zingiberaceae) all larvae sp., Poa compressa (Cavanilles) Grisebach (Malvaceae), Malvastrum Setaria repens sp. (Leguminosae), Guazama sp. (Tiliaceae) Inga ., Malvastrum coromandelinus Agropyron

Inga Sorghum Linné (larvae), Triumfetta josefina sp., sp., sp. (Poaceae) Maulik 1937 sp. (Poaceae) sp Griffith sp., sp. (Poaceae) ‘beetles seen on cereals’ (Linné) Blume, Linné (larvae), sp., (Leguminosae) larvae, Sida rhombifolia sp., sp. (Poaceae) Collart 1934 sp., sp., sp. (Orchidiaceae), sp. (Musaceae), Triumfetta Elatteria Bentham sp. (Malvaceae), (Poaceae), Triumfetta Althaea Phleum Sorghum Vossia sp., Agropyron Beauvois, Elymus repens Triticum ulmifolia Indigofera bonariensis Panicum (Legumominosae), Althaea Gracke, amabilis Spathoglottis Donax Amomum sp. (Zingiberaceae) Cassia fruticosa Musa coerulea Vanda sp., Arundina grandis Martinez, Mimosa questioned by Hespenheide and Dang 1999) Cassia (Sapindaceae) larvae (record of Bolivia (Sterculiaceae), Sri Lanka Venezuela, Ecuador, Peru, Costa Rica, Panama, Colombia,(larvae), Linné Mexico to Brazil (Bohemann) Paraguay, Uruguay, Argentina Weise China ‘narrow leaved grass’ (larvae) Gressitt and Kimoto 1963 (illustr.), Kalshoven 1957 (Baly) Central America (Baly) Central Kraatz Congo (Wiedemann) Sumatra, Java, Borneo sp. Asia, Indo-Australian region

Gyllenhal United Provinces (India), ‘cholum’ sp. Mexico to Argentina (Erichson) Mexico, Nicaragua, Chapuis India sp. Linné northern Africa, Europe, Asia atra

Heterispa H. ramosa H. stygia H. viridicyanea Hispa andrewesi H. H. vinula Heterispa costipennis Heptispa limbata Taxon AuthorTaxon G. xanthomela Geographical Distribution Reported Host Plants Author FamilyReferences Selected Gonophora Heptispa 66 J.A. Santiago-Blay 1986 (illustr.); Kalshoven et al. 1986 (illustr.), Gressitt and Kimoto 1963 et al. 1985, Chen t 1963 et al. Gressitt 1957 (illustr.), 1960a, 1963 1957; Kimoto 1999 (illustr.); Maulik 1919, 1937; Tan 1993 (illustr.) Gressitt 1957, 1960a, 1963 sp., Jolivet 1989a sp., Poa Zizania Zizyphus sp., sp. Saccharum sp., sp. Jolivet 1989a sp., Jolivet 1989a sp., Linné, grass Cox 1996 (illustr.), Gressitt and Kimoto 1963 Kalshoven Triticum sp.) (Poaceae) Zea Oryza sativa Miscanthus sp., sp., Digitaria sp., sp. (Solanaceae), sp. (sugarcane), Pinanga sp., Paspalum Bambusa Aegopodium sp., sp. (Leguminosae), sp., (Poaceae), Malus Sorghum (Poaceae) Physalis

sp. (Arecaceae) Jolivet 1989a, Staines 2002b Jacquin (Arecaceae) Mariau 2001 Jacquin (Arecaceae) Mariau 2001 Jacquin (Arecaceae)2001 1988, Mariau Themedea sp. sp. (Poaceae) larvae Gressitt 1957, 1960a, 1963; Kalshoven 1957 Dactylis sp., Saccharum Blume (Arecaceae) larvae Kalshoven 1957 sp., Heteropogon Linné ‘grasses’, (Poaceae) sp. (Poaceae) An sp. (Arecaceae) larvae Kalshoven 1957 sp. (‘suan-me-mang’ genus), Abdullah and Qureshi 1969; Chen sp., Linné Robinia Cocos (Cistaceae), Panicum Metroxylon Saccharum officinarum sp. Jolivet 1989a, 1989b sp. (Poaceae) larvae Kalshoven 1957, Maulik 1937 sp., sp. (‘bai-mao’ genus),

Zizania sp. (Verbenaceae) sp., ‘bamboo’ ( sp. (Zingiberaceae) Gressitt 1957 (illustr.), 1960a, 1963 (illustr.); Staines 2003b sp. sp., sp., sp. (Arecaceae) larvae Kalshoven 1957, 1981 sp., sp., Saccharum (Poaceae) Sorghum Saccharum

Elaeis guineensis Elaeis guineensis sp. sp., (Arecaceae) Bambusa Areca bamboo (Poaceae) larvaeCecropia Alpinia Jolivet 1989a, Kalshoven 1957 Oryza Metroxylon Zizania ‘grasses’Themeda sp., Imperata Saccharum spontaneum Miscanthus Gressit Imperata spontaneum Imperata Pinanga kuhlii Agropyrum Lantana Centotheca Themeda sp., Imperata Saccharum Areca (Rosaceae), Cistus sp., Zea sp. (Rhamnaceae), Elaeis guineensis Elaeis Hainan Island Java Sumatra, Naias, Borneo, Philippine Islands, Indonesia Java Cambodia, Vietnam, China,Taiwan, Malaysia, Indonesia, (larvae), Linné (or Myanmar), Thailand, Laos, Japan, Taiwan, Siberia eastern Asia, PhilippineIslands, southern China, Hainan Island, Malaya leaf (Poaceae) 1957 Guinea (?), Buru (?) Java Indonesia Ecuador, Colombia, and Amazon region Sumatra Latin America, especially de Colombia Bondar Brazil (Gestro) Australia, New Guinea wild (Guérin-Méneville) French Guiana (Gressitt) New Guinea 5 (Germar) Australia, Papua-New (Spaeth) (Bates) Sri Lanka, India, Burma Uhmann (Chapuis) Australia, New Guinea Pic Berti and Desmier Uhmann Uhmann (Baly) eastern China, Korea, sp. Old World sp. French Guyana to Brazil (Baly) Burma, India, south ‘bamboo’, (Gestro) New Guinea sp. Java sp. Indonesia: Java and Sumatra, sp. Palearctic, Turkey, and Chenon H. subfasciata H. ollagnieri Hispoleptis diluta Klitispa opacula Klitispa Micrispa alpiniae Metaxycera subapicalis Javeta H. coarctatus H. csikii H. moerens Hispellinus J. thoracica Hispellinus albertisii H. moestus H. multispinosus J. corporaali H. callicanthus Hispa Javeta arecae Hispoleptis Leaf-mining chrysomelids 67 Jolivet 1989a, Kalshoven 1957, Kimoto 1999 1991, Maes et al. 1928 Brisley 1925, McCauley 1938, Moldenke 1971, Richerson and Boldt 1995, Riley and Enns 1979 Clark 1983 et al. Chittenden 1902, Clark 1983, Maulik 1937, McCauley 1938, Needham F. A. sp. Maes 1998, Staines 1996 sp., and Clark 1983, Hilgendorf and Goeden 1981 E. sp. Gressitt 1963 (illustr.), Staines 2003b Franseria Salvia sp. Gressitt 1963, Staines 2003b hirsutus Cavanilles,

Pityopsis acanthicarpa

Flourensia cernua Brickelia Torrey, Helianthus sp. (Zingiberaceae) Gressitt 1963, Staines 2003b Hooker and Arnold, Heliconia Encelia californica ) Heterotheca Alpinia ex sp. (larvae) Gressitt 1957 (illustr.), 1960a; Staines 2003b (Bentham) Payne Clark 1983; Goeden and Ricker 1975, 1976a, 1976b; Jones N. L. Burman Gillett sp. (Zingiberaceae) DeCandolle (adults), sp. (Poaceae) 1998, Maes and Staines 1991 Nees, Helianthus sp., Costus Torrey and A.Gray , Riley and Enns 1979, Staines to Santiago-Blay, pers. comm., July Franseria Oersted (larvae), Zea Costus A. Gray Costus Rolfe, ? F. ambrosioides Nuttall (Leguminosae; according Sida acuta (Zingiberaceae) Gressitt 1963, Staines 2003b sp. (Asteraceae (Michaux) Nutall (Asteraceae),

Helianthus (DeCandolle) Rydberg, Cavanilles (adults), (Michaux) Elliott or sp., S.F. Blake, Robinson (adults), sp. sp. (Asteraceae)sp. (Asteraceae) Clark 1983, Noguera 1988 sp. (Marantaceae) larvae(illustr.), 1996 Cox diffusus (Gray) Payne (adults), A. confertifolia sp. (adult),

sp. (Zingiberaceae) Gressitt 1963, Staines 2003b sp. (Zingiberaceae) sp. (Zingiberaceae) sp. (Zingiberaceae) larvae sp. and other gingers (Zingiberaceae) Gressitt 1963, Staines 2003b 2003b Staines 1963, Gressitt Gressitt 1963, Staines 2003b Gressitt 1963, Staines 2003b , E. farinosa sp. (Musaceae), Haplopappus squarrosus Alpinia halimifolia DeCandolle (larvae), (Hooker) Coville, Nutall confertiflora H. venetus Rafinesque, dumosa vernicosa sp., to Clark, a questionable host plant record) Salvia costaricensis (Lamiaceae) adults Helkianthius Ambrosia chenopodiifolia Alpinia graminifolia Lupinus Gossypium (Zingiberaceae) (Heliconiaceae), ? Musa Alpinia Alpinia Alpinia Alpinia Maranta Solidago drummondii Solidago sp. (Asteraceae)Solidago Doellingeria umbellata Solidago Chrysopsis 2003 Alpinia Donax cunniformis United States Nicaragua, Costa Ricaadults, (Malvaceae) (New Guinea) (Zingiberaceae) adults New Guinea and Mexico (Say) southern Canada, United States (Mannherheim) western United States Gressitt New Guinea Gressitt New Guinea ‘a smooth-leaved ginger’, ? Gressitt New Guinea Baly Guatemala to Colombia Gressitt New Guinea Baly Mexico, Honduras, Gressitt New Guinea (Olivier) southern Canada and (Motschulsky) Sri-Lanka, Java ‘ginger’ (Zingiberaceae) larvae Kalshoven 1957, Staines 2003b Schwarz eastern and southeastern Gressitt New Guinea ? (Newman) Florida (United States) Gressitt New Guinea sp. southeastern Asia Clark southeastern United States Gressitt New Britain, Ireland Gressitt New Guinea Gressitt New recognized by Clark 1983) M. rileyi M. perforata M. pulchella (including data subspecies and Mexico (larvae), M. rubrolineata M. maai M. musae M. cubicularis M. donaxiae M. pellucida M. puncticollis M. semiviridis Gressitt M. sinuicosta M. zinzibaris Micrispa Microrhopala cyanea M. erebus M. excavata (including data for subspeciesrecognized by Clark 1983) M. floridana United States M. costi Taxon AuthorTaxon M. biakana Geographical Distribution Reported Host Plants Author FamilyReferences Selected 68 J.A. Santiago-Blay et al. 1928; Riley et al. Uhmann 1937 Monrós and Viana 1947 (illustr.) 2000; Damman (1994); Ford and Cavey 1985 (illustr.); Lawson Goeden and Teerink 1993, Staines 2002b and Enns 1979 Byes 2002 (illustr.); Cappucino 1991; Chittenden 1902; Clark 1983, 1991; Maulik 1919; McCauley 1938; Needham Chittenden 1902; Clark 1983, 2000; Ford and Cavey 1985 (illustr.); 1928; Riley and Enns 1979; Williams 1989b, 1991 Goeden and Ricker 1974, 1976c; McCauley 1938; Needham S. S. sp., S. Aster sp., altisima

J. Staines 2002a (illustr.) sp. Aster Nuttall (Linné) Aiton, Willdenow Torrey and Linné, sp., Brickellia sp. (Poaceae) Monrós and Viana 1947 (illustr.) A. novae- Linné, Solidago S. laevigata sp., Solidago Helianthus S. missouriensis A. paternus Solidago Serjania asteroides spp. (Asteraceae) sp., larvae Linné,

S. graminifolia

(Linné) L’Her, sp. larvae Aiton, sp., (Linné) Beauvois Aiton, A. Jussieu Hespenheide 2000, and Dang 1999 A. Jussieu Hespenheide 2000, and Dang 1999, Maes 1998 sp., S. nemoralis A. simplex DeCandolle (larvae), (Chamisso) Monrós and Viana 1947 (illustr.) K. Schumann Cox 1996 (illustr.), Hespenheide and Dang 1999, Maulik 1937, K. Schumann Hespenheide and Dang 1999, Maulik 1937, Uhmann 1934 (illustr.) Aiton, , Paspalum Boltonia Serjania costaricensis Paullinia costaricensis (Lessing) Greene, Solidago caesia Solidago sp. (Lamiaceae) Linné, sp., Nees Solidago californica Burman f., Silphium Dicoria canescens John Donnell Smith (larvae), Smith Donnell John Hespenheide 2000, and Dang 1999 Paullinia S. juncea S. juncea asteroides Franseria sp. (Asteraceae)

Seriocarpus Bartling, s sp., Conceveiba pleiostemona Silphium laciniatum sp., A. cordifolius sp., Setaria viridis , A. patens Aster Sericocarpus Salvia Linné, sp., Linné (larvae), sp., Linné, Nuttall, sp. (Rhamnaceae) Linné, sp., A. psilostachya sp., Nees, Solidago sp. (Sapindaceae) larvae S. mollis Boltonia Linné S. canadensis S. lanceolata Seriocarpu Encelia Boltonia Chrysopsis Silphium (Asteraceae), Panicum grumosum Pithecoctenium echinatum (Bignoniaceae) larvae, Donnell Smith (Euphorbiaceae).’ Stigmaphyllum lindenianum (Malpighiaceae) larvae Stigmaphyllum lindenianum (Malpighiaceae) larvae unable to find name (larvae), Paullinia Sapindaceae Adenocalymma marginata DeCandolle (Bignoniaceae) Colubrina spinosa Colubrina (larvae), perfoliatum Linné, chilensis sp., Gray, Pithecoctenium echinatum (Bignoniaceae) larvae, Radkofer (larvae), (Sapindaceae) Aiton, (Linné) Salisbury, Nuttall, sempervirens A. puniceus Ambrosia chamissonis Cronquist, angliae Ambrosia grandiflora (larvae), Seriocarpus sp. (Poaceae), ‘boneset’, ‘bottle brush grass’, ‘box elder’, ‘boxwood’ ‘service berry’, ‘shad bush’ BSP, canadensis (Asteraceae), Panama southwestern Canada southern Canada Canada to Colombia Maulik Argentina Staines Costa Rica ‘Taken fogging Uhmann Costa Rica Uhmann Costa (Baly) Costa Rica

(Baly) Central America sp. Chapuis Argentina (Baly) Nicaragua, Costa Rica, (Weise) Paraguay, Bolivia, Argentina (Newman) United States and (Baly) Central America (Baly) Central (Fabricius) United States and Nonispa carlosbruchi Ocnosispa humerosa Octhispa bimaculata O. decepta O. elegantula O. gracilis O. haematopyga O. elevata O. elongatas M. xerene M. vittata Microrhopala Leaf-mining chrysomelids 69 et al. 1928, Staines 1989 (illustr.) et al. 1928; Staines 1989 (illustr.) et al. Moldenke 1971; Needham Balsbaugh and Hays 1972, Broughton 2000, Chittenden 1902, Clark Broughton 2000, Cilliers 1983, Maes 1998, and Staines 1991, Monrós and Viana 1947 (illustr.) sp., Staines 2002b sp. L. viridis Vigna sp. sp., Linné, Lactuca

Sesamum Cassia Linné, americana sp. Michaux, Fraxinus

Chionanthus M. Vigna sp. (adults), Fraxinus Coccoloba Salvia aterrimum Marshall, (Linné) 2000, Ford and Cavey 1985 (illustr.), Krauss 1964, Needham

Tilia Kunth; sp. (Pedaliaceae), vulgaris L. glandulosissima

Sesamum Malpighia capitata sp. (Rhamnaceae),

Adenocalymma sp. (adults), Origanum sp. (Verbenaceae) larvae (Lamiaceae), radicans sp., sp. (Verbenaceae) Warburg (Fagaceae) (illustr.), 1987b (illustr.); Harley 1969; Krauss 1964; Maes 1998,

Machaerium

Sesamum L. hispida Stizolobium Linné (adults), sp. (Ehretiaceae), sp. Phaseolus sp., Ligustrum vulgare DeCandolle (Asteraceae) Annonymous, no date; Broughton 2000; Cilliers 1977, 1983, 1987a Lespedeza Colubrina Swartz (adults), Lantana Linné (Lamiaceae), F. pennsylvanica Lantana (Linné) Linné (Polygonaceae) Sanderson 1967 Swartz (Bombacaceae) Hespenheide and Dang 1999, Wilcox 1975 Stizolobium Campsis Linné (larvae), Gray (Asteraceae) (larvae), Jones and Brisley 1925. Needham Linné; Inga Byrsonima Linné (Apiaceae) (adults), sp. (adults), sp. (Malpighiaceae), sp. (Rubiaceae) Serjania sp., (Verbenaceae) (larvae) Krauss 1964, Staines 1989

Origanum (Linné) Walpers (adults), Linné, sp. (larvae) (Leguminosae), sp. (Bombacaceae), sp., sp. (adults) (Oleaceae), sp. (adults), Miller; sp. Linné (adults), Linné (larvae? and adults), sp. (adults), sp. (larvae? and adults) sp., Mentha spicata Mentha occidentalis Quercus astriglans

Dioclea Eupatorium collinum Fraxinus Ligustrum Linné (Tiliaceae), (adults) (Lamiaceae) , Phaseolus Piper and Tracy, unguiculata (adult) (Leguminosae) (adults), orientale Lantana camara Ocimum basilicum sp. (Oleaceae) (adults) Daucus carota Perezia thurberi Aesculus (Hippocastanaceae), larvae Lantana Lespedeza virginica americana Salvia urticifolia Lantana camara Basanacantha Mentha sp. (Polygonaceae), (Leguminosae), Stigmaphyllum sp., Malpighiaceae Ochroma Coccoloba uvifera Ochroma lagopus Paullinia spp. for biological Lantana Honduras, El Salvador, adults, Nicaragua, Hawaii, Australia, adults, South Africa Linné, and Mexico eastern and southern sp. (Asteraceae), United States Seeman ex Bureau (larvae) (Bignoniaceae), 1928 (illustr.), Riley and Enns 1979, Staines 1989 (illustr.) and South Africa control of States). Introduced into Hawaii Nicaragua, Guatemala, México,and southern Texas (United sp. (Pedaliaceae) all previous ones, adults; Staines 1996 (illustr.) , Uhmann 1934, 1937 Baly Panama, Costa Rica, Honduras, Uhmann Costa Rica Horn southwestern United States Guérin-Méneville Mexico, Guatemala, Suffrian Cuba (Fabricius) Brazil, Honduras, Cuba, Weise Puerto Rico sp. Mexico to Argentina Uhmann Paraguay, Argentina O. scabripennis O. plicatula O. marginicollis O. gundlachi Octotoma championi O. spitzi Octhispa Taxon AuthorTaxon O. loricata Geographical Distribution Reported Host Plants Author FamilyReferences Selected O. nervermanni 70 J.A. Santiago-Blay 1928 et al. 1999 (illustr.) et al. 1928 Clark 2000; Ford and Cavey 1985 (illustr.); Hargrove 1986; Hodson Annonymous 2000; Butte 1968c (illustr.); Chittenden 1902 1942; Kirkendall 1984; Kogan and 1979; Lawson 1991; et al. ’ (illustr.); Wheeler 1980 (illustr.), 1987 and Mengel P. Verbena Fagus Miller, (Linné) sp. sp., Monarda Robinia grandis sp., Jolivet 1989a, Staines 2002b sp. Robinia

, Betula Lippia Ulmus sp., sp., japonica

sp. (Pedaliaceae), Linné, McPherson and Ravlin 1983; Mullins 1976; Needham Q. pedunculata sp. Prunus serotina Linné (adults), lobata sylvestris Linné, Linné, 1984; Williams 1989a Tectona

(Ehrhart) Medic, (Elliott) DeCandolle, Tectona sp. (Malpighiaceae), Quercus Meibonia rigida Eugenia ovalifolia Campsis Lespedeza sp., Mentha Sophora sp., Linné, Q. prinus sp., Sesamum ), Falcata comosa Canavalia Malus sp. (larvae), sp. (adults) (Leguminosae), Ouratea cuspidata Linné (Polygonaceae) rigidum (Linné) Fernald (adults), Butte 1968c (illustr.), Buntin and Pedigo 1982, Chittenden 1902, fruticosa

(Linné) DeCandolle, Ford and Cavey 1985, Kogan 1979 (illustr.), Needham DeCandolle (Leguminosae)1929 Maulik alba

(Linné) Merrill (larvae),

D. C. tomentosa Xanthorrhoea sp. adults (Betulaceae), Salvia Lantana Linné, calpodendron Robinia max

Lantana Stigmaphyllum Marshall (Aceraceae) Dioclea Miller (adults), (Willdenow) Owhi (adults), Wisteria (Vellozo) H.S.Irwin and Teixeira R. pseudoacacia sp., sp., bracteata (Linné) (Leguminosae) Butte 1968c (illustr.)

Gray, Amorpha sp., Quercus sp. (Asteraceae), (Linné) Merrill, Betula Linné, sp. (adults), Linné (Ulmaceae) spp. (larvae), Cavanilles (adults), sp. (Oleaceae), lobata Q. rubra

sp. (Bignoniaceae), Glycene (Loureiro) Merrill var. sp. (adults) (Fagaceae), ‘rhododendron Linné, sp. (Leguminosae) Staines 2002b Crataegus Linné, Origanum montana (Willdenow) Maesen and S. Almeida, hispida Linné (larvae), (Ericaceae), Desmodium Pueraria Quercus alba sp. (adults), sp. (larvae in all Polygonum perfoliatum adult, sp. (Verbenaceae), (Xanthorrhoeaceae) Kuntze, Laburnum C. coccinea Senna australis R.C.Barneby (Leguminosae), Cambess. (Myrtaceae), Tieghen (Ochnaceae) Glycine soja Acer saccharum Senna Ehrhart, (Leguminosae), Fraxinus Clerodendron Malus sylvestris Ehrhart, ‘some quinces’ (Rosaceae), americana Amphicarpa Desmodium canescens D. illinoense Cymbosema (Fagaceae), (Lamiaceae), umbellata Linné F. (Verbenaceae) (adults) Canavalia ensiformis Eupatorium sp., Glycine max Hayek (larvae), Tecoma eastern United States French Guyana to Argentina (Baly) Brazil (Uhmann) Arizona (United States)

sp. Maulik Brazil (Thunberg) southeastern Canada, sp. United States to Brazil Smith eastern half of the United States Octouroplata octopustulata O. dorsalis Octouroplata Odontota arizonicus O. horni Octotoma O. tesselata Leaf-mining chrysomelids 71 1928 et al. 1928 (illustr.), Riley and Enns 1979 et al. Maes 1998, Monrós and Viana 1947 (illustr.) Anand 1989, Maulik 1937, Zaka-ur-Rab 1991 Butte 1968c (illustr.), Chittenden 1902, Ford and Cavey 1985, sp. A. sp. Apios Ulmus Prunus sp., C. sp., I. sp. Kalshoven 1957, Maulik 1937, Staines 2003b serrulata sp. Linné f. Linné Needham

sp., Tephrosia sp. Rhododendron sp., Aiton sp. Glyceria Carpinus nigra sp. (Fagaceae), sp. Jolivet 1989a sp. Rubus Alnus sp.,

Meibomia Malus Canavalia sp., sp. (adults), alternifolia sp. (adults) Oryza Phaseolus

Medikus (larvae), Betula sp., sp. (Polygonaceae), (Linné) Persoon sp. (Tiliaceae), sp., Quercus Solidago Sophora Amphicarpaea Betula Curcuma (Linné) Fernald Butte 1968c (illustr.), Ford and Cavey 1985 Riley sp., Tilia sp., Cornus sp., Quercus sp., (Linné) DeCandolle, Glycine sp. (Cornaceae), sp., (Linné) Persoon Butte 1968c (illustr.), Chittenden 1902, Needham sp. (Convolvulaceae), Jolivet 1989a, Staines 2003b, Uhmann 1953 sp., Desmodium americana

Crataegus Miller (Leguminosae) larvae Hespenheide and Dang 1999, Wilcox 1975 Dioclea (Linné) Lamarck larvae, Fagus Polygonum Solanum auriculatum sp., bracteata

Robinia Cornus Alnus Linné (Leguminosae), Mina sp., Apios Amorpha sp., Eupatorium unable to find name, (Rosaceae), sp. (Dioscoreaceae), sp., sp. (Convolvulaceae) Anand 1989 sp. (Zingiberaceae) larvae. Staines Koenig ex Roxburgh (larvae) sp., sp. (adults) (Asteraceae), Moench, apios

(Willd) Trinius (Poaceae), Tephrosia virginiana sp., sp., Pyrus Desmodium (Solanaceae) all adults Cimbosema spontanea sp., sp. (Ulmaceae) Ipomoea sp. (Convolvulaceae), Dioscorea Curcuma (2003b) considers that ‘the true host plant of this species appears to be unknown.’ Ipomoea Ipomoea batatas (Poaceae), Orchidiaceae, (Zingiberaceae) Cassia fruticosa Canavalia ensiformis (Leguminosae), Ameliancher sp., sepiaria (Convolvulaceae), ‘egg plant’ (Solanaceae) larvae Ipomoea Discorea Acacia Pueraria (Asteraceae), sp. (Ericaceae), (Betulaceae), (adults) (Betulaceae), (Fagaceae), (adults) (Rosaceae) Aster (Leguminosae) Solidago (adults) (Cornaceae), tuberosa Glycine nervata (Leguminosae) Amphicarpaea (Leguminosae) larvaeTephrosia virginiana 1979 Enns Elliot, Paraguay, Argentina (Leguminosae), Ecuador, Brazil, Bolivia, Panama, Venezuela, Colombia, Archipelago, Java,Sumatra (Indonesia) (Dioscoreaceae) larvae, Orchidiaceae (larvae), United States (Aiton) Willdenow (adults), United States Gestro India, Malaysia (Baly) Costa Rica, Nicaragua, Panama (Olivier) India (Olivier) southeastern Canada and sp. Old World (Chapuis) Mexico, Nicaragua, Costa Rica, Weise India, Java (Sanderson) eastern half of the United States (Olivier) eastern and southeastern sp. North and Central America Oncocephala angulata O. dorsalis O. tuberculata Oxychalepus alienus O. anchora Oncocephala Odontota O. scapularis Taxon AuthorTaxon Geographical Distribution Reported Host Plants Author FamilyReferences Selected O. mundula O. notata 72 J.A. Santiago-Blay Hespenheide and Dang 1999, Maes 1998, Staines 1991, sp. sp., Jolivet 1989a, Staines 2002b sp. Serjania sp., sp. sp., sp. Vail sp. Gressitt and Samuelson 1988 (illustr.), Staines 2003b Serjania sp., Humboldt, Freycinetia (Leguminosae),

sp. as sp. sp., sp. (Arecaceae), Ramos 1998, Flowers and Janzen 1997, Staines 2002b sp. (larvae), Staines 1996 (illustr.), Uhmann 1937 (illustr.) sp. Dioclea Heliconia sp. (Heliconiaceae), Gressitt and Samuelson 1988 (illustr.), Staines 2003b Vernonia sp. (larvae) Elephantopus sp., Eupatorium sp. (Poaceae), C. urticifolia Canavalis ), Heliconia Heliconia sp., sp. (Malpighiaceae), Cocos Malpighia sp. (Pandanaceae), Paullinia sp., sp. (Leguminosae), Pithecocthenium sp. (larvae), sp., Clibadium Vernonia mollis B. L. Robinson (larvae), Vernonia Aublet (larvae), (Humboldt, Bonpland, and Moldenke 1971 Phaseolus Heliconia Sprengel (Malpighiaceae) larvae1937 Uhmann 1937, Maulik sp., Gray (Asteraceae) Boldt and Staines 1993 (illustr.), Cox 1996 (illustr.) sp., Eupatorium populifolium sp., Chusquea Linné, Alpinia Miller (Leguminosae) Hespenheide and Dang 1999, Maes 1998 Cassia Verbesina Malpighia sp. (Bignoniaceae) adults Hespenheide and Dang 1999, Maes 1998, Staines 1991 Cymbosema Pandanus DeCandolle, Desmodium lindheimeri Clibodium (Malpighiaceae) Sanderson 1967

sp. (Pandanaceae), ‘ginger’ Colea sp., Calea Indigofera sp., sp. (Flagellariaceae), sp., sp. sp. sp. (Heliconiaceae) Gressitt and Samuelson 1988 (illustr.) sp. (Agavaceae), sp., sp. (larvae), sp. (Poaceae), (Musaceae), Zingiberaceae Mucuna sp. (Solanaceae). Staines (2003b) sp., Pandanus Pithecoctenium Bunchosia (Asteraceae), (Bignoniaceae), Chusquea (Malpighiaceae), sp. (Sapindaceae) adults ‘unverified’. Benthamantha mollis Kunth) Alefeld, (Legumiunosae) both adults Banisteria argentea Calea axillaris Baccharis bigelovii Baccharis Eupatorium sp., (Sapindaceae) (Zingiberaceae) Solanum considers the reference to Elephantopus spicatus Elephantopus Hooker and Arnold (larvae), Verbesina costaricensis Verbesina (Asteraceae), DeCandolle, Bonpland and Kunth, Malpighia glabra Freycinetia Heliconia Cassia fruticosa Pleomele Flagellaria (Heliconiaceae), Inga Centrosema Costa Rica, Panama, Colombia Jamaica Mexico Group) Malaita) ‘banana’ Costa Rica Samuelson Solomon Islands ‘coconut palm’ (Arecaceae (Baly) Central America (Thunberg) (Baly) Costa Rica unidentified host plant (larvae) Hespenheide and Dang 1999 Gressitt Solomon Islands (Santa Isabel) (Baly) (Chapuis) Mexico to Panama Gressitt Solomon Islands (San Cristobal, ‘rattan’ (Arecaceae), (Chapuis) Mexico, Guatemala, Nicaragua, sp. Mexico to Argentina (Baly) southwestern United States (Fabricius) sp. southern United States to Peru bellicosa nr. P. explanata P. morio Oxyroplata bellicosa O. P. fairmairei P. suturalis Pentispa Pentispa collaris (includes four subspecies listedin Gressitt and Samuelson 1988) (Malaita, Santa Isabel, New Georgia Group, Florida (Pittosporaceae), (Heliconiaceae), ‘banana’ (Musaceae), ‘karo’ Pharangispa alpiniae P. cristobala P. heliconiae Oxychalepus O. posticatus Leaf-mining chrysomelids 73 1986 (illustr.), Kimoto 1999 Zaka-ur-Rab 1991 et al. Maulik 1929, 1937; Gressitt 1957 (illustr.), 1960a; and 1919, 1937; Zaka-ur-Rab 1991 Piper sp. Costus Blume Chen sp., sp., Jolivet 1989a, Staines 2002b jujuba sp.,

sp. Staines 2002b Sorghum sp., Gressitt 1957, Kalshoven 1957 sp. (Poaceae) Jolivet 1989a sp. (Piperaceae) Hespenheide and Dang 1999, Staines 2003b, Wilcox 1975 Linné, 1919, Zaka-ur-Rab 1991 Zea Dioclea Dioclea Lamarck Zizyphus Alpinia Stigmaphyllum Cassia Piper sp., sp., sativa E. lithosperma sp.,

sp. (Marantaceae), Uncaria gambier Eryhtrina Linné (Poaceae), Abdullah and Qureshi 1969; Anand 1989; Kalshoven 1957; Maulik sp. (Zingiberaceae) sp. (Leguminosae), (Roxburgh) Bentham Bernon and Graves 1979 (Linné) Brotero (larvae), Abdullah and Qureshi 1969, Anand 1989, Kalshoven 1957, Maulik (Linné) DeCandolle, Monrós and Viana 1947, Staines 1998, 2002b (illustr.) mauritiana sp. (Zingiberaceae) Jolivet 1989a Linné (larvae), Oryza

sp. (Arecaceae) Jolivet 1989a Swartz (Bombacaceae) Hespenheide and Dang 1999 Z. DeCandolle, ‘katjangen’ Lamarck, Saccharum Costus Sprengel, Calathea sp. (Malpighiaceae) larvae Hespenheide 2000 Cymbosema Cymbasema sp. (Malpighiaceae) Phaseolus Linné, sp., Zingiber sp. (Heliconiaceae), sp., sp., Phoenix sp. (Bombacaceae), sp (Leguminosae), sp. (Marantaceae), sp. (Rhamnaceae), ‘paddy’ Anand 1989, Maulik 1937 sp. (Leguminosae), sp., sp., Persoon (Poaceae), ‘wild grass’ sativa sp.,

Miquel (larvae) (Leguminosae) Andropogon sorghum Avena Saccharum officinale vulgare (Malpighiaceae) larvae Costus Andropogon Heliconia Canavalia ensiformis Canavalia Canavalia Stigmaphyllum Phaseolus Mucuna Ochroma Areca Ochroma lagopus Stigmaphyllum Calathea (Legumonosae), sp. (Piperaceae), Saccharum officinarum Erythrina ex. Cajanus indicus Tephrosia candida (Leguminosae) all larvae, Roxburgh (Rubiaceae) Zisyphus Lamarck (larvae), (Rhamnaceae) Pueraria phaseoloides (Leguminosae) Erythrina indica Malaita, Santa Isabel),New Georgia sp. ‘ginger in bush’ (Zingiberaceae) larvae Samuelson 1988; Staines 2003b Peru, Argentina and northern South America Archipelago and, more broadly SE Asia) (or Myanmar), Vietnam, Borneo, Sumatra, Java, Sulawesi (= Celebes Islands), Philippines, Guinea (Principe Island, St. Thome Is.) (or Myanmar), Thailand, Vietnam (Baly) Mexico, Costa Rica, Panama Pic southern Central America Weise Pakistan, India Weise India, Sri Lanka Maulik Solomon Islands, (Guadalcanal, Gestro Java, Sumatra sp. Solomon Islands sp. 1’Rica Costa sp. southern United States to Peru (Guérin-Méneville) Africa sp. Nicaragua to Argentina (Fabricius) India, Sri Lanka, Burma sp. East Indies (Malasyian (Guérin-Méneville) India, Sri Lanka, Burma Guérin-Méneville Colombia, Brazil, Paraguay, sp. (some species are miners ) Indo-Australian region Platochispa Phidodonta modesta Physocoryna Pharangispa Phidodonta Taxon AuthorTaxon P. purpureipennis Geographical Distribution Reported Host Plants Author FamilyReferences Selected P. scabra Physocoryna expansa Platocthispa Pistosia ‘ Platochispa championi Platypria andrewesi P. echinogale P. erinaeus P. coronata P. echidna 74 J.A. Santiago-Blay 1986 (illustr.), Gressitt and 2001; Maulik 1937; Gressitt et al. et al. 1986 (illustr.), Jolivet 1989a et al. Gallego and Abad 1985; Howard 1957 (illustr.), 1960a, 1963; Kalshoven 1981 (illustr.); Lepesme 1947; Panggoy and Pedro 1982 (illustr.) Kalshoven 1981 (illustr.), Lepesme 1947, Maulik 1937 Chen Zaka-ur-Rab 1991 Abdullah and Qureshi 1969, Chen sp., sp., s sp. sp. sp., sp., Cox 1996 (illustr.), Jolivet 1989a sp., sp. sp., Rubu nucifera sp., Desmodium sp.

A. pinnata Smith Flagellaria Korthalsia Oncosperma Miquel, Kimoto 1963, Kalshoven 1957, 1999 (illustr.), Maulik 1937, Archontophoenix (Linné) sp., Metroxylon sagu Myrica Rottbóll, sp., ex. Mucuna sp., Erythrina indica sp., Korthalsia Arenga Cocos Thunberg, Caryota cumingii Brrassus flabellifer Roxburgh, Sesbania sp., sp., sp., sp., Pyrus Daemonorops Rhopaloblaste Oncosperma Areca sp., sp. (Flagellariaceae), sp. (Arecaceae) all larvae Gressitt 1963 (illustr.) Thunberg, Cajanus sp., palm similar to Gressitt 1963 (illustr.) sp., Blume sp. (Flagellariaceae), Sesbania aculeata spp., Cyrtostachys Linné (Poaceae) larvae(illustr.) 1960a Gressitt sp. (Arecaceae) larvae(illustr.) 1960a Gressitt sp., sp. (Myricaceae), Linné (Poaceae) larvae Cox 1996 (illustr.); Gressitt 1957 (illustr.), 1960a sp., Cocos (Roth) Nees (Poaceae) Rubus ellipticus Areca Areca sp., S. grandiflora sp. (Rubiaceae) Linné, Blume Erythrina Beccari (larvae), Heterospathe (Arecaceae) all larvae sp., sp., Arenga

Nypa fruticans Myrica Linné (Arecaceae) Lepesme 1947 Linné (Arecaceae) Lepesme 1947 (illustr.) Linné (larvae), Corypha elata Kunth (Arecaceae), Calamus Nypa Phoenix Pueraria Linné, Metroxylon sagu sp., Phoenix sp. Flagellaria Flagellaria ? (Arecaceae) Gressitt 1960a (illustr.) sp., Cocos Korthalsia Uncaria sp., Martinez (larvae), sp., sp., sp., sp. (Poaceae) sp., spp. (larvae), E. lithosperma sp., ex Nypa fruticans sp., sp., sp. (Rhamnaceae), sp. (Fagaceae), Rhopaloblaste Linné, Dolichos Areca catechu Saccharum Oreodoxa (Arecaceae), Metroxylon Daemonorops Calamus Cocos nucifera Saccharum officinarum Archontophoenix Cyrtostachys renda Leptochloa chinensis Korthalsia Metroxylon Oreodoxa regia indica Adonidia merrilli Areca catechu Metroxylon Saccharum officinarum Rottbóll, sp. (Arecaceae) Archontophoenix sp., Phaseolus Heterospathe (Rosaceae) larvae (Leguminosae), Ziziphus (Rosaceae), Heterospathe Quercus Cocos nucifera Calamus Dolichos lablab Erythrina Philippines, New Guinea, sp., (New Britain, Cape York (Wurmb) Merrill, Malaysian Peninsula,Sumatra, Java, Sulawesi(= Celebes Islands) Indonesia, (larvae), Linné Loddiges Bismark Archipelago, Samoa, Australia Peninsula, Malacca),(larvae), Linné New Guinea Islands) (Myricaceae) larvae, Java, Sulawesi (= Celebes Persoon (larvae) (Leguminosae), Burma (or Myanmar),China, Sumatra, Indonesia, Lamarck, Poiret, (Willdenow) Thailand, Laos, Vietnam, Maulik New Caledonia Gressitt New Guinea Gressitt New Gressitt New Guinea Gressitt New Guinea Gressitt New GuineaGressitt New possibly Spaeth New Caledonia Gressitt New Guinea Gressitt Chapuis Thailand, Malaysia, Indonesia, sp. Old World (Fabricius) India, Sri-Lanka Nepal, Maulik Malaysian Peninsula, Java sp. Indo-Australian region P. saccharivora P. ruficollis P. reichei Plesispa P. nipae P. palmella P. palmarum P. montana Platypria Plesispa cocotis P. hagenensis P. korthalsiae P. hystrix Leaf-mining chrysomelids 75 et 2001 2001; Kalshoven 1957, 2001; Kalshoven 1957; 2001; Kalshoven 1957; 2001; Kalshoven 1957; et al. et al. et al. et al. et al. 1983; Howard 1977; Howard et al. et al. 1986 (illustr.), Cox 1996 (illustr.); Gressitt 1959; Lepesme et al. 1977; Gallego Abdullah and Qureshi 1969, Cox 1996 (illustr.); Dharmadhikari Lepesme 1947; Maulik 1937 Chen 1947 (illustr.); Mariau 2001; Maulik 1929 (illustr.), 1937 sp. Jolivet and Hawkeswood 1995 sp. Gressitt 1957 (illustr.), 1960a (illustr.); Maulik 1937; Kalshoven nucifera

sp., Hespenheide and Dang 1999, Staines 2002b sp. Dharmadhikari Zea sp. Lepesme 1947; Mariau 2001 sp., Alpinia sp. Cocos larvae Metroxylon al. sp. (Arecaceae) Gressitt 1957 (illustr.), 1960a, Howard Inga Livistona sp. (Flagellariaceae) Gressitt 1957, Lepesme 1947 sp., sp., ‘unidentified Gressitt 1957, 1960a, 1963; Howard Thunberg (larvae), 1981 (illustr.); Lepesme 1947; Maulik 1919, 1929 (illustr.), 1931, Jacquin (Arecaceae) Abdullah and Qureshi 1969, Maulik 1937 Pandanus Piptocarpha Sorghum Hasskarl Kalshoven 1957 Baker (Asteraceae) Hespenheide and Dang 1999 Jacquin, sp. (Verbenaceae) sp. (Asteraceae), ex sp., Cocos ) sp. Gressitt 1957 (illustr.), 1960a sp., ) sp. (Arecaceae) larvae Gressitt 1957 (illustr.), 1960a; Howard Caryota Calamus sp. (Pandanaceae) larvae(illustr.) 1960a Gressitt Seeman and H. Wendland, Lepesme 1947; Mariau 2001; Maulik 1931, 1937 fruticans

) Don (Asteraceae) larvae Cox 1996 (illustr.) Flagellaria sp., Verlot (Bignoniaceae) Hespenheide and Dang 1999 guineensis

Endl Kunth (Arecaceae) 1937; Mariau 2001; Zaka-ur-Rab 1991 Verbena Linné, Humboldt and Bonpland (Asteraceae) Hespenheide and Dang 1999 Vernonia Linné (Arecaceae) Linné (larvae), Nypa and Mikania guineensis Linné,

pacifica sp. (large species) (Pandanaceae) larvae1963 (illustr.), 1960a Gressitt Bryantia

sp. (larvae), and other Arecaceae sp. sp. (Bignoniaceae), ( sp. (Heliconiaceae) larvae, Elaeis Pennisetum Balaka sp., sp., Ptychosperma Ptychosperma ( ( Elaeis sp. (Arecaceae) Lepesme 1947, Mariau 2001 sp., sp. (Arecaceae) larvae Gressitt and Kimoto 1963, Kalshoven 1957 nucifera sp., sp.,

Rottbóll, Oryza (Poaceae) Pollia thyrsiflora (Leguminosae), Heliconia Cocos Arecaceae Pandanus Mikania guaco Verbesina Arrabidaea Vernonia mollisima Arrabidaea chica Rolandra (Commelinaceae) larvae Piptocarpha chontalensis Cocos nucifera Pritchardia Areca catechu Cocos nucifera Pritchardia sagu Freycinetia Oreodoxa regia Cocos Balaka Cocos Cocos Balaka Areca Java Pacific islands, Philippine Islands, Solomon Tonga, Fiji, Samoa including Java, Singapore, Linné, Malayan Archipelago, other Pacific islands, Australia Philippines, Borneo, Indonesia Malik Solomon Islands (Malaita) (Baly) Costa Rica, Nicaragua Guérin-Méneville Maulik Solomon Islands (Kolombangara) Blanchard southeast Asia and many Gressitt Biak Island (New Guinea) Pic Argentina UhmannRica Costa unidentified plant (larvae)(illustr.) 1934 Uhmann Gressitt Solomon Islands Aulmann Samoa Maulik Sulawesi (= Celebes Island), (Erichson) Philippine Islands sp. 1’Rica Costa Maulik Solomon Islands Weise New Guinea, Solomon Islands sp. Nicaragua to Argentina Baly Sri-Lanka, southeast Asia Maulik Fiji Baly Australia (Cape York sp. Africa sp. India to New Guinea Zingiberaceae (questionable Staines 2003b) Gressitt 1982, Staines 2003b Spaeth Solomon Islands Uhmann Costa Rica Probaenia Taxon AuthorTaxon Polyconia Geographical Distribution Reported Host Plants Author FamilyReferences Selected Prionispa fulvicolis Promecotheca alpiniae (including two subspecies listedin Gressitt 1960a) P. antiqua P. bicolor P. bryantiae (including two subspecies Peninsula), New Guinea native palms’ (Arecaceae), (Zingiberaceae) larvae 1957 P. crenulata mentioned in Gressitt 1963)P. caeruleipennis (Pandanaceae) larvae P. pici ‘ Probaenia Probaenia armigera Prionispa P. atricornis P. callosa P. cumingii P. freycinetiae P. guadala P. cyanipes (includes two species listed in Gressit 1960a) P. lindingeri P. nuciferae P. kolombangara P. leveri 76 J.A. Santiago-Blay et al. 2001, et al. 2001 et al. 2001; Lepesme 1947; Maulik 2001; Kalshoven 1957, 1981; 2001; Kalshoven 1957; Lepesme et al. et al. et al. 1986 (illustr.), Cox 1996 Lespesme 1947 1947; Mariau 2001; Maulik 1929 (illustr.) et al. ressitt 1957, 1960a; Howard Maulik 1931 (illustr.) Chen 1937 Cox 1996 (illustr.); Gressitt 1957 (illustr.), 1959; Kalshoven Abdullah and Qureshi 1969, Lepesme 1947, Mariau 1988, Maulik Abdullah and Qureshi 1969, Gressitt 1960a, Howard Kalshoven 1957, Lepesme 1947, Mariau 2001, Maulik 1937 sp., 1981, Würmli 1975 sp., Cocheraeu 1972, Gressitt 1959 (illustr.), Jolivet 1989a, Kalshoven sp. (Arecaceae) Gressitt 1957 (illustr.), 1960a; Howard sp. Cocos

Thunberg, Elaeis Pandanus Linné, Gressitt 1957 (illustr.), 1959 1960a, 1963; Howard sp. Jacquin, 2001; Kalshoven 1957, 1981; Lepesme 1947; Mariau 2001 Pritchardia Rottbóll, Caryota sp., Elaeis sp., (Arecaceae) Gressitt 1957, 1959; Howard Heliconia

Pavón, Ravenala sp., sp., sagu sp. (Arecaceae), et

sp. (Arecaceae) G sp. sp. (larvae), sp. (Poaceae), sp. (larvae), Phoenix Calamus sp. (Pandanaceae) 1947 sp. (Pandanaceae), sp. (Musaceae), Ravenala Ruiz Nypa fruticans sp. (Arecaceae) Gressitt 1957, 1960a; Howard

Livistona Metroxylon sp. (larvae), sp., ) ) sp., Cocos Cocos Calamus Areca , Cocos nucifera Phoenix Wendland, and other Arecaceae 1937 Metroxylon Elaeis guineensis Phytelephas sp., Saccharum J. F. Gmelin, Nypa Pandanus Ravenala Pandanus et Linné, Linné, Linné, Linné, Cocos sp., Linné Linné, sp. (Arecaceae) larvae, macrocarpa sp., sp., sp. (larvae), spp. (cultivated and wild) or Cox 1996 (illustr.); Gressitt 1957 (illustr.), 1960a sp., sp. (larvae), sp. (Poaceae) sp. (Flagellariaceae), spp. (Pandanaceae) Gressitt 1957 (illustr.), 1959, 1960a, 1963 sp. (Pandanaceae) larvae Gressitt 1960a (illustr.), 1963 (illustr.) Jacquin, Balaka Linné, Ptychosperma Ptychosperma ae Lespesme 1947 (illustr.), Maulik 1931 1937 Seeman ( sp. (Zingiberaceae) ( sp., sp. (larvae), sp. (larvae), sp. (larvae) (Arecaceae) ecaceae Livingstonia Phychosperma Flagellaria (Heliconiaceae), Freycinetia Mischanthus Alpinia Areca Cocos nucifera Pandanus Cocos nucifera pacifica larvae Lepesme Metroxylon Nypa Balaka Cocos nucifera Elaeis Cocos Ptychosperma Saccharum Miscanthus Balaka Cocos nucifera larvae and some madagascariensis (larvae) (Arecaceae) Pandanus Areca cathecu nucifera guineensis Phytelephas Phytelephas Areca catechu Ysabel, New Georgia)Oceania sp. (Pandanaceae) Pacific region (Indonesia) (New Guinea) Fiji, Solomon Islands, Indo- Moluccas Islands, Sula Island New Guinea ‘unknown thick-leaved palm’ (larvae)1963 (illustr.), 1960a Gressitt Bismark Archipelago Australia, Manus, New Britain, Islands (Santa Cruz Island, Banks Island) Weise Colombia Arecace Maulik Solomon Islands Weise New Britain, Manus Spaeth Solomon Islands (Guadalcanal) sp. Africa, Asia, Indo-Australia, Gestro Vanuatu and Solomon Maulik Colombia Ar Gressitt Solomon Islands ‘small pinnate palm’ Gressitt 1960a (illustr.) Csiki New Guinea, Solomon Islands, Gressitt Solomon Islands (Guadalcanal) Uhmann Solomon Islands (Bougainville, Gressitt New Guinea Baly Philippines, Tonga, Samoa, Baly Australia Maulik Sulawesi (= Celebes Island), Prosopodonta corallina P. cordillera P. ptychospermae P. reichei Promecotheca P. salomonina P. sacchari P. straminipennis P. varipes P. papuana P. palmella P. palmivora Gressitt P. pandani P. violacea P. soror P. opacicollis Leaf-mining chrysomelids 77 1988 (illustr.), Cox 1996 (illustr.) 1985, Kalshoven 1957, Tan 1993 et al. et al. Monrós and Viana 1947 Lepesme 1947 (illustr.), Maulik 1937 Maulik 1937 Lepesme 1947 (illustr.); Maulik 1931 (illustr.), 1937 Lepesme 1947 (illustr.), Maulik 1937 Anand 1989 An Kalshoven 1957, Maulik 1937 , Linné sp. Hespenheide and Dang 1999, Maes 1998, Staines 1991, sp.). Jolivet 1989a, and Hawkeswood 1995, Staines 2002b Zea mays Schott Hespenheide and Dang 1999 sp., Jolivet 1989a Stevensonia Digitaria (Rutaceae) sativa sp. (Arecaceae) Jolivet 1989a Linné (larvae)

’ Warburg Staines 2002a (illustr.) sp. (Poaceae) sp. sp., Cuspania E. Taylor(illustr.) 2002a Staines Heliconia Andrade and Andrade 1984 (illustr.), Jolivet 1989b Oryza Esenbeckia febrifuga Oryza Pavón) Mez. Sanderson 1967 (illustr.), Virkki and Santiago-Blay 1998 Zea radiatum leaf-miners and sp. (‘ma-tang’ genus), Saccharum officinarum et Martínez sp., sp., nana Linné (larvae), ex Pourouma var. Reitz (Araceae) Costa Stevensonia Roscheria Oryza sativa (Ruiz var. Virola koschnyi sp., sp., sp., Digitaria (Araceae), Triticum Prosopodonta Miscanthus Linné ‘rice’, sp. (‘ye-gu-cao’ genus), Jacquin (Clusiaceae) larvae Hespenheide and Dang 1999, Maes 1998 sp. (‘di’ genus), sp., sp. (Araceae) adults, sp., lyratiloba sp. (larvae), host plants. Cecropia

Beauvois, eae Maulik 1931 (illustr.), 1937 recaceae recaceae (Sapindaceae) larvae(Myristicaceae) adults Philodendron radiatum 1996 Staines (Araceae) larvae Philodendron sp. (A. St.-Hil.) A. Jussieu Arecaceae Staines 2003b casts serious doubts on the association of sp. (Arecaceae) Phoenicophorium Linné ‘sugarcane’ (Poaceae) Arundinella glabra Oryza sativa Miscanthus (larvae) Poaceae Imperata Sterculia recordiana papyracea (Sterculiaceae) adults Philodendron renauxii Anthurium Cecropia (Cecropiaceae) Rapanea ferruginea Myristicaceae Clusia flava Carauta, Heliconia Phoenicophorium Linné (larvae), ‘wild grasses’ (larvae) Poaceae Saccharum officinarum Saccharum Digitaria Río Vitaco China, Hainan Island, Malaya Staines Costa Rica Maulik Seychelles Islands (Baly) Thailand, Laos, Vietnam, ‘wild grasses’ (larvae) Gressitt and Kimoto 1963, Kalshoven 1957, 1999 (illustr.) Staines Costa Rica unidentified species of Visaceae (larvae) Staines 2002a (illustr.) (Baly) Costa Rica, Nicaragua Weise [Colombia], St. Antonio, Weise Colombia Arecac (Motschulsky) Korea, Japan, China, Siberia sp. Seychelles Islands (Pacific Ocean) Maulik India Weise Colombia A sp. tropical America Arecaceae and Heliconiaceae ( Sanderson Puerto Rico Weise Colombia A Staines Costa Rica ‘Collected fossing (Fabricus) Brazil (Baly) Nicaragua, Costa Rica, Panama (Olivier) Brazil Baly Brazul, Paraguay, and Argentina (Motschulsky) Sumatra, Java sp. Asia (Baly) Costa Rica, Panama unidentified Araceae (larvae) Hespenheide and Dang 1999, Wilcox 1975 (Baly) Mexico to Panama bidens

af.

Taxon AuthorTaxon P. interrrupta Geographical Distribution Reported Host Plants Author FamilyReferences Selected P. sulphuricollis Prosopodonta S. godmani S. obscurovitatta S. pretiosa P. intercepta P. quinquelineata Rhabdotohispa R. nigrocyanea Rhadinosa fleutiauxi R. lebongensis R. parvula Sceloenopla bicolorata S. S. erudita S. lampyridiformis S. mantecada S. nigropicta S. maculata S. longula Rhabdotohispa scotti Rhadinosa 78 J.A. Santiago-Blay et al. 1928, Maulik 1937 et al. Hespenheide and Dang 1999 Jolivet 1989b Monrós and Viana 1947 Jolivet 1989a Butte 1969 (illustr.), Cavey 1994, Ford and 1985 (illustr.) 1928; Noguera 1988; Wheeler and Snook 1986 (illustr.); Wilcox 1954 1985 (illustr.); Frost 1924; Maulik 1919, 1937; Needham sp. sp. Maulik 1937 A. sp., Linné, Balsbaugh and Hays 1972; Butte 1969 (illustr.); Ford Cavey E. Cupania Ell., S. umbellata sp. sp. (adults)sp. Jolivet 1989a, Moldenke 1971 Ruprechtia Ruprechtia sp. (Clusiaceae), Cecropia (Leguminosae) Maulik 1929 (illustr.) papyracea Phaseolus

agerateroides

Persea sp. (Sapotaceae), Clusia Ryds. (larvae), sp., and others Jolivet 1989b, and Hawkeswood 1995, Staines 2002b (illustr.) Hooker and Arnold Maulik 1937, Uhmann 1937 sp. (Cecropiaceae) Jolivet 1989b A. sagitifolius sp. (Asteraceae)(illustr.) 1969 Butte Lamarck (Poaceae) and Cox 1996 (illustr.), Maulik 1937, Monrós and Viana 1947 Linné (adults), sp. (Dilleniaceae), , A. novae-angliae Sphaeralcea DeCandolle DeCandolle, sp. (Myrsinaceae), Rubiaceae, (Arecaceae), Standley var. (Linné) Elliott,

Eupatorium J. Torr. (Malvaceae)J. Torr. (Malvaceae) Staines 1986b Staines 1986b sp., sp. L. (Chenopodiaceae), Jones and Brisley 1925, Needham (Aristolochiaceae), Linné

Davilla Chrysophyllum Zexmenia (Vahl) Kubitzki (Dilleniaceae) larvae Hespenheide and Dang 1999, Wilcox 1975 (Leguminosae), Philodendron sp. (Polygonaceae) Lamarck, sp. Pourouma Rapanea grossulariaefolia emoryi emoryi sp. (Malvaceae)

sp. (Aristolochiaceae), sp. (Rutaceae), Rubiaceae, Malva sp., sp. (Poaceae) Jolivet 1989a, Staines 2002b spp. (Heliconiaceae) larvae Hespenheide and Dang 1999, Staines 2003b sp., Cocos sp. (Cecropiacee), sp., E. maculatum oleracea sp. (Poaceae) ‘bamboo’ (larvae) Cox 1996 (illustr.) Maulik 1937, Uhmann 1934 sp. (Sterculiaceae)

sp., Huber (Polygonaceae) Willdenow, divaricatus

Coccoloba Davilla nitida Cecropia Sterculia recordiana (Sterculiaceae) larvae Cyclanthaceae, Anthurium Pourouma Lonchocarpus (Araceae), (Lauraceae), Esenbeckia (Sapindaceae), Sterculia (Malvaceae) other grasses Paspalum Arustolochia Paspalum quadrifarium Spinacia latifolia Aristolochia Canavalia ensiformis larvae Sphaeralcea Sphaeralcea Althaea Eupatorium populifolium (Asteraceae) larvae Canavalia ensiformis (Leguminosae) Aster A. paniculatus Bambusa simplex Baccharis Sphaeralcea Sphaeralcea Heliconia Strophostyles helvola Linné f., Caribbean Indonesia Unknown Paraguay Costa Rica to Argentina United States Britton (Leguminosae) and Mexico imperalis

S. rudgeana

Horn southwestern United States S. Uhmann (Baly) Costa Rica and Panama Weise Peru, Uruguay (Weise) Brazil, Colombia Maulik Brazil Staines Costa Rica Unidentified species of Visaceae (larvae) Staines 2002a (Weber) North and Central America sp. 3’Rica Costa (Schaeffer) eastern and southern sp. Central, South America, and sp. North and Central America flavidus (Baly) Nicaragua, Costa Rica, Panama sp. Costa Rica to Argentina (Baly) Brazil

(Schaeffer) western United States Schaeffer western United States (Uhmann) Central America Butte Arizona (United States) sp. [prob. sp.[monotypic, (Baly) Central America Sceloenopla S. scherzeri ‘ S. unicostata S. sheppardi Sceloenopla Spilispa Stethispa crenulata Stenostena laeta Sternostena (Baly)] Stenopodius Stethispa S. fryi S. fuscicornis S. heringi S. inaequalis Uhmann]S. canavaliae sp., S. arnetti S. texanus Stenopodius Sumitrosis amica S. ancoroides S. lateralis Leaf-mining chrysomelids 79 2000, Butte 1969 (illustr.), Cavey 1994, Hespenheide and Dang 1999, Dolichos Cornus D. malus Robinia sp., Banks, Urtica Robinia

sp., S. Aiton Linné, Houttuyn, sp., sp. sp., Jolivet 1989a; Staines 2002b, 2003b sp., C. nistitans sp. Cornus Rafinesque Pyrus Desmodium sp. (Fagaceae), Cajanus Linné, (Linné) adults, Laportea Solidago Lamarck, Linné (= ‘white (Muehenberg) sp. (Chenopodiaceae), (Linné) Merrill, Oenothera hirsutus sp., S. gigantea Vernonia E. rugosum

Linné, Desmodium Linné (adults), Helianthus Linné (Cyrillaceae) Ford and Cavey 1985 (illustr.), Noguera 1988 Celastrus E. urticifolium (Linné) Willdenow sp., Quercus nictitans (Linné) Salis.,

R. pseudoacacia Miller (Rosaceae), sp., sp., malus Linné (Chenopodiaceae, Balsbaugh and Hays 1972, Buntin Pedigo 1982 (identification (S. Watts) Britton, C. rugosa laevigata

Linné, triloba Linné (Solanaceae),

Michaux (larvae), Linné (adults), Cassia Quercus alba Richard, Helianthus Glycine max sp. (Brassicaceae), Cassia Malus Chenopodium Solidago racemiflora Amphicarpaea Arabis

Linné (adults) (Leguminosae), virginiana (Wooton and Standley) W.C. Linné (Rosaceae), (Linné) DeCandolle (adults),

sp., sp., Linné (adults), sp., intermedia (Muehenberg) Wood (adults), (Linné) Weddell (adults) (Urticaceae), sp.,

novaboracensis fruticosa canadensis Eupatorium dulcamara

Rudbeckia S. graminifolia

sp., sp. (Asteraceae), Muehenberg (adults),

sp. (Cyrillaceae), sp. (Leguminosae), Aiton (Urticaceae), ‘everlasting’ Amphicarpaea bracteata Radicula Rosa Cyrila malus

sp., Linné, Solanum Robinia (Onagraceae), Linné (Leguminosae) adultsChenopodium album Maes 1998, Staines 1996 gracilis Cassia fasciculata neomexicana Martins and Hutchins, 1985), adults, (adults), Solidago (adults), ulmifolia Poiret, asperifolia Michaux, sp. (Cornaceae), oak’, Fagaceae), Vernonia E. urticaefolium Eupatorium perfoliatum (Asteraceae), Amorpha glutinosum paniculatum Desmodium Canavalia Amorpha Lespedeza pseudoacacia Malus canadensis unidentified Urticaceae Wedelia (Celastraceae), Cyrilla Aster Rudbeckia States and Mexico and States probably an error according to Ford and Cavey error, corrected by Ruesink 1984), Butte 1969 (illustr.), Clark (Baly) United States to Panama (Baly)Panama Rica, Costa Mexico, ‘unidentified Fabaceae’ (larvae) Hespenheide and Dang 1999, Wilcox 1975 sp. Canada to Argentina (Weber) Canada, eastern United Taxon AuthorTaxon Geographical Distribution Reported Host Plants Author FamilyReferences Selected S. rosea S. pallescens S. terminata Sumitrosis 80 J.A. Santiago-Blay 1995 et al. 2001a, Jolivet 1989a, Ravelojaona 1970 et al. lart 1928, Uhmann 1964 Broughton 2000, Winder and Harley 1982 Maulik 1937 Harley 1982 (illustr.), 1987b (illustr.); Harley 1969; Krauss 1964; Winder and Monrós and Viana 1947 (illustr.) Broughton 2000, Krauss 1964, Uhmann 1934 (illustr.) L. sp. Lippia sp. sp., Monrós and Viana 1947 (illustr.), Jolivet Hawkeswood 1995, sp. unable sp., Lantana L. sp., sp., Banwo Linné, sp. Linné, sp., Lantana Lantana sp. (Annonaceae) Maulik 1937, Peña sp. (Verbenaceae) Heliconia Linné (larvae), Panicum sp. (Rosaceae), Meibonia Oryza Chusquea Tectona grandis Pithecoctenium ., Lantana camara Laportea unable to find L. trifolia Robinia L. tiliaefolia sp., Wedelia paludosa Miller, sp., K. Schumann larvae, Hespenheide and Dang 1999, Maes 1998, Staines 1996, Uhmann Chamisseau, Verbena trifolia sp., Lantana et Baif Rollinia longifolia , Annona sp., (Linné) Gentry Cilliers 1983, Monrós and Viana 1947 (illustr.) Potentilla Arrabidaea coleocalyx Cham. (Aristolochiaceae) Monrós and Viana 1947 (illustr.) sp. (Poaceae) Schau., Linné (Verbenaceae) Cox 1996 (illustr.), Monrós and Viana 1947 (illustr.) Lantana camara Sch. Bip. (Asteraceae) Monrós and Viana 1947(illustr.) D. Don (Asteraceae) Maulik 1937 (Miller) N. E. Britton, and Schlecht. and Cham., Rottbóll, Poeppig, Linné Linné (Pedaliaceae), Annonymous, no date; Broughton 2000; Cilliers 1977, 1983, 1987a Poeppig, Steudel, Lespedeza Bambusa Linné, Malpighia glabra Caesaria silo Cham., L. urticifolia Pueraria sp., unidentified bignanacean, 1934 et Echinochloa Setaria (Sprengel) Briquet, sp. (Leguminosae), sp. (Commelinaceae), sp., K. Schumann (Bignoniaceae), sp., sp., sp. (Malpighiaceae) sp., Schlechtendal sp. (Sterculiaceae), Lippia alba L. micromera Linné, sp. (Poaceae), L. glutinosa sp. (Poaceae) (all adults) Staines 2002b sp. (Asteraceae) larvae, Glycine Lantana glutinosa Schlechtendal Rolandra argentea DeCandolle (Asteraceae) montevidensis Vernonia mollissima Linné f. (Verbenaceae) (larvae), Wilson, tiliaefolia Sesamum orientale Clerodendron thomsonae Linné, Commelina Pharus Acroceras Annona squamosa Macfadyena unguis-cati name, Leguminosae, Paspalum Aristolochia fimbriata Anona squamosa echinatum Lippia myriocephala camara sp. (Verbenaceae), larvae on Verbenaceae Pithecactenium echinatum Pithecoctenium (Bignoniaceae), Malpighia Phaseolus sp., Strophostyles (Heliconiaceae), (Urticaceae) Lasiacis Guazuma Lippia urticoides Vernonia molissima to find name, and other verbenaceans (Verbenaceae) A.St.Hil. (Anonaceae), Bureau and K. Schumann (Bignoniaceae) Verbena bonariensis Calea Zaire Unknown Col Hawaii, Australia, South Africa Argentina, South Africa? (Bignoniaceae), Argentina Costa Rica, Guatemala, Mexico Uhmann Brazil to Argentina (Maulik) Brazil Baly Brazil, Venezuela, Panama, Chapuis Gestro Sao Thome Is., (Olivier) ‘Guiana’, Brazil (Olivier) ‘Guiana’, (Pic) Argentina Weise Brazil, Paraguay, Argentina Chapuis Bolivia, Paraguay, (Pic) Argentina Buzzi and Winder Brazil, Mexico Chapuis Uruguay, Argentina (Guérin-Méneville) Tanzania, Madagascar (Guérin-Méneville) Tanzania, Pic Brazil, Paraguay, Argentina, Chapuis Nicaragua to Brazil U. lantanae U. mucronata U. jucunda Trichispa feae T. sericea Uroplata annonicola U. coarctata U. girardi U. fusca Temnochalepus insolitus U. atricornis U. bilineata U. bipuncticollis U. daguerrei U. fulvopustulata Leaf-mining chrysomelids 81 1928 et al. Cox 1996 (illustr.), Maulik 1919, Monrós and Viana 1947 (illustr.) Hespenheide and Dang 1999, Wilcox 1975 Butte 1968a (illustr.), Jones and Brisley 1925, Kogan sp., and Hawkeswood 1995, Maulik 1932, Staines 2002b, Winder sp. sp., Harley 1982 Inga Sida Verbena sp. Maes 1998, Maulik 1937, Uhmann 1934 (illustr.) Bignonia Gouania vulgaris

sp. Monrós and Viana 1947 sp., sp., Robinia sp., sp., sp., sp., Baccharis Nissolia Gaertner 1934, 1937 Byrsonima sp. (Annonaceae), Cox 1996 (illustr.), Flowers and Janzen 1997, Jolivet 1989a, Poeppig and Eupatorium Urera Lippia Althaea Martinez Vernonia sp. (Bignoniaceae), Phaseolus sp. (Leguminosae)Needham 1979, sp., Wisteria chinensis Coussapoa Martinez sp., sp. (Vivianaceae) sp., Rollinia Arrabidaea Calopogonium sp. (Poaceae), sp., C. villosa , Phaseolus Ortega (Convolvulaceae) Monrós and Viana 1947 (illustr.) Robinia sp., DeCandolle (adults), Hespenheide and Dang 1999; Maes 1998; Staines 1996; Uhmann Inga edulis Barb. (Leguminosae) Maulik 1937 Banisteria Caesarea Lantana Linné (Leguminosae) larvae Cox 1996 (illustr.), Monrós and Viana 1947 (illustr.) Liebmann, Jacquin (larvae), Smith (Asteraceae) adults Moldenke 1971 Panicum Kunth (Verbenaceae) Monrós and Viana 1947 (illustr.) sp., sp. (Poaceae) both adults Monrós and Viana 1947 Rolandra Synedrella nodiflora Elephantopus Pithecoctenium Standley, Linné (larvae), (Linné) Merrill, (Poaceae) Thomas and Werner 1981 Annona both adults

DeCandolle

sp. (Aristolochiaceae), Pourouma bicolor sp. (Asclepiadaceae), sp., Oryza Gill. ex Planch (Ulmaceae) adults Monrós and Viana 1947 (illustr.) sp., sp., sp., sp. (Malpighiaceae), sp. sp. (Cecropiaceae) larvae(illustr.) 2000 Staines sp. (Lauraceae), sp. (Leguminosae) adults Maes and Staines 1991, 1996 sp. (Asteraceae), Phaseolus sp., Colea Lippia geminata Clibadium aspersum Clibadium larvae (Asteraceae), (Leguminosae) larvae Melanthera nivea Aristolochia Clibadium (Sims) DeCandolle (Leguminosae), others adults pseudoacacia Digitaria Melanthera Mucuna pluricostata Cecropia Inga affinis nymphaeifolia Wedelia sp., Ocothea sp. (Verbenaceae), Phaseolus vulgaris Endlicher., (Cecropiaceae) all larvae Ipomoea heterophylla adults Olyra Nissolia fruticosa (Leguminosae) Nissolia Cecropia insignis (Urticaceae) Celtis tala sp. (Leguminosae), sp. (Malvaceae), sp. (Rhamnaceae), Glycyne max Schubertia Malpighia Argentina South America Mexico Linné, Paraguay, Argentina (Baly) Costa Rica and Panama ‘undetermined Fabaceae’ (larvae) Hespenheide and Dang 1999 Weise Paraguay, Argentina Pic Brazil, Peru, Bolivia, Weise Paraguay, Argentina (Baly) Mexico to Costa Rica (Chapuis) Costa Rica, Panama, Staines Costa Rica Weise Paraguay, Argentina congruus Jacoby Mexico laetificus (Smith) United States Chapuis Mexico to Panama Maulik Brazil (Baly) Mexico and Central America Uhmann Bolivia, Brazil, Paraguay ab. (Chapuis) Brazil, Paraguay, Uruguay, ab. (Chapuis) Bolivia, Argentina sp. Central and South America Acanthaceae, (Weise) southern United States and Taxon AuthorTaxon U. nigritarsis Geographical Distribution Reported Host Plants Author FamilyReferences Selected U. sculptilis U. sulcifrons U. uniformis Uroplata X. mucunae X. hespenheidi X. medius X. ater Xenochalepus amplipennis X. haroldi X. faustus X. guerini X. contubernalis X. erythroderus X. bicostatus fasciatus X. chapuisi X. bajulus 82 J.A. Santiago-Blay 2000 (illustr.) et al. 1947 (illustr.); Maulik 1919 (illustr.), 1937 1937; Moldenke 1971; Staines 1996; Uhmann 1934 (illustr.), 1937 sp., Bl., Kalshoven 1957, 1981; Lepesme 1947, Maulik 1937 sp. sp., Faba Oryza Mucuna sp., sp. Hespenheide and Dang 1999, Jolivet 1989a, Staines 2002b sp., Urera Schubertia (Poaceae),

sp., sp., sp. a sp. Roxburgh, Kalshoven 1981 (illustr.), Lepesme 1947 Canavalia Robinia Ipomea Olyra Thunberg, O. filamentosa Zalacca conferta Zea Dolichos sp., sp., sp., Pedley Borowiec 1999, Monteith 1991 (illustr.) Lathryrus sp., sp., sp. (Theaceae) Borowiec and Takizawa 1991, Monteith 1991

(Linné) Britton, Sterns Butte 1968a (illustr.), Ford and Cavey 1985 Theobrom sp., A. triandra Fagaceae’ (Leguminosae), (illustr.) Ridley, sp. (Ulmaceae), Barbosa Rodrigues Monrós and Viana 1947 (illustr.) Gray (Leguminosae)(illustr.) 1968a Butte (Humboldt, Bonpland, and Butte 1968a (illustr.); Flowers and Janzen 1997; Maes 1998; Maulik Linné (Euphorbiaceae) Medvedev and Eroshkina 1988 (illustr.) crassa Hooker and Arnott Monrós and Viana 1947 (illustr.) sp. (Leguminosae), (Loureiro) Merrill Rane O’Brien, ornamental palms Kuntze (Rubiaceae) Borowiec 1999 Thun. Bauhinia Phoenix Phaseolus Dioclea Centrosema macrocarpum Bambusa Celtis Inga Nypa fruticans Linné, (Leguminosae) Monrós and Viana 1947 (illustr.) ssp. figillaria Saccharum sp., sp.,

sp., sp., sp., sp. (Arecaceae) larvae Kalshoven 1957 sp. (Araceae) larvae, sp. (Arecaceae)1989a Jolivet Wisteria sp. (Leguminosae) adults Monrós and Viana 1947 (illustr.) sp., sp. (Apocynaceae) Borowiec 1999 sp., ‘date palm’ (Arecaceae) Abdullah and Qureshi 1969; Anand 1989; Kalshoven 1957; Lepesme sp. (Rosaceae), sp., Nissolia Glycine Griffith (Arecaceae) Oncosperma (Arecaceae) Phoenix Oncosperma Metroxylon Phoenix roebelinii Areca catechu Metroxylon Metroxylon (Urticaceae) Prunus (Sterculiaceae), sp., Vigna sp. (Malvaceae), sp., Panicum Anthurium (Convolvulaceae), Cymbosema ‘repollo’ (Brassicaceae), ‘frijol’ (Leguminosae) Passoa 1983 Phaseolus Lathyrus pubescens (Leguminosae) larvae Mucuna pluricostata (Leguminosae) Benthamantha mollis Phaseolus sp. Phaseolus polystachios and Poggenberg (Leguminosae) larvae Robinia neomexicana Acacia crassa Canthium inerme Bentham, ‘species of ‘wild bean vine’ (Vitaceae) all adults Cleyera japonica Carissa Phyllanthus emblica Carallia brachiata (Rhizophoriaceae) Indonesia Honduras Argentina Costa Rica Kunth) Alefeld, Argentina (Waterhouse) Australia Gestro Singapore Gestro Sulawesi (= Celebes Island) dorsalis (Baly) (Maulik) Pakistan, India (Maulik) Pakistan, 9 (Chapuis) Bruch Argentina sp.World New Pic Argentina Butte eastern United States (Crotch) southwestern United States to Borowiec and Takizawa Nepal Maulik Malaysian Peninsula, Carey Island (Boheman) South Africa, Tanzania Spaeth Socialist Republic of Vietnam Uhmann Butte Arizona (United States) (Spaeth) India (Spaeth) India, Sri Lanka sp.(some species CASSIDINAE Nothosacantha W. dactyliferae W. phoenicia appear to be leaf-miners) Wallaceana apicalis Wallacispa Wallacispa javanica X. viridiceps X. tandilensis X. trilineatus Xenochalepus X. omogerus X. potomacus X. robiniae X. signaticollis X. phaseoli N. laticollis N. nepalensis N. severini N. siamensis N. vicaria Leaf-mining chrysomelids 83 ne irdly, the Australian Plant Name Index (APNI) begins with Linnaeus me Index. However, there are problems with each of these indexes. is table as detailed data, including identification, were not provided Although it has full, parenthetic author citations of New World taxa ines (2002b). See Crowson (1955), Gressitt and Kimoto 1963, Staines e incomplete author citations, such as missing the parenthetical ic distinction of these two taxa has been repeatedly pointed out by numerous larvae in old inflorescenses and fruitlets or Chenopodiaceae. Monoxia invasive weed), it will have updated the full author parenthetic citation. A little web ‘tutorial’ e.g., 2003). ‘Trichostomes’ (Jacoby) has been used for Alticinae + Galerucinae (Schmitt to Santiago-Blay, personal communication, Ju et al., 2003; Kim et al., believe that all the species in genus are leaf miners. However, I have collected Monoxia Böving, 1927; Duckett e.g., Firstly, Index Kewensis (IK) did not include parenthetic author citations until recently. Relying on it can cause a user to hav of a basionym. Secondly, the Gray Card Index (GCI) begins after first issue Kewensis, names published 1893. names, it does not account for names published before 1893. The GCI is the best tool of plants from New World. Th and incorporates full author citations. Essentially, if the plant needs to be accounted for ( on standard reference botanical works (etc.) can be found in http://persoon.si.edu/botlinks/dhntyp.htm. workers ( 2003). in that paper. (2002b), and references therein, for a discussion on various placements of the hispines cassidines. 1. This site contains the names accumulated in three important indices: Index Kewensis, Gray Card Index, and Australian Plant Na 2. I am using the more traditional, yet artificial, separation of Alticinae and Galerucinae. The problematic phylogenet 3. Most students of 4. Hespenheide 1991 refers to about 40 other leaf mining taxa in this subfamily Mesoamerica. They have not been entered th 5. About ten described species inhabit western Africa (Mariau 1988). 6. About 30 more described species inhabit Madagascar (Mariau 1988). 7. About four species inhabit northwestern South America (Mariau 1988). 8. About ten described species inhabit southeastern Asia and the Pacific islands (Mariau 1988). 9. This genus was placed in the Hispinae by Medvedev and Eroshkina (1988) but Cassidinae Borowiec (1995, 1999) Sta