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Center for Systematic , Gainesville, Insecta Mundi Florida

December 2002

Mexican

J. B. Heppner Florida state Collection of , Bureau of Entomology, Division of Plant Industry, Florida Department of Agriculture & Consumer Services, Gainesville, FL

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Heppner, J. B., "Mexican Lepidoptera biodiversity" (2002). Insecta Mundi. 550. https://digitalcommons.unl.edu/insectamundi/550

This Article is brought to you for free and open access by the Center for Systematic Entomology, Gainesville, Florida at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Insecta Mundi by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. INSECTA MUNDI, Vol. 16, No. 4, December, 2002 171

Mexican Lepidoptera biodiversity

J. B. Heppner Florida State Collection ofArthropods Florida Department of Agriculture and Consumer Services P.O. Box 147100 Gainesville, FL 32614-7100 U.S.A.

Abstract. Being among the most habitat diverse countries in the world (and plant diversity is about 22,000 sp.), Mexico has a Lepidoptera fauna recorded at about 14,385 but is estimated to be over 22,000 species, if not much higher (some estimates go to 35,000 sp.). High Lepidoptera numbers in Mexico are also due to the large influx of tropical species from the border with Guatemala, as well as Mexican endemics. In this report, the Lepidoptera families are summarized for Mexico, giving known species and what experts estimate to be the true total for each when all have been described. Many regions of Mexico are still poorly known for smaller . Introduction brings in the highest number of species for the Mexican total, perhaps more than a third of all Mexico encompasses one of the most diverse Lepidoptera species found in Mexico. Yucatan and regions of the world, perhaps having only somewhat , separated by only 220 km., undoubtedly have a fewer habitats than does Peru, since Mexico ranges large overlapping fauna, asMiller and (1989) from desert areasjust south of California and Arizona noted for . In comparing larger rainforest to the tropical rain forests of Chiapas next to Guate- trees in Mexico with the biogeography of Lepidoptera, mala, plus great variations in topography and local about 30% of the tropical trees are distributed into habitats in between (Toledo and Ordhfiez, 1993). Guatemala, 25% range asfar south asAmazonia, 21% Along with the correspondingly diverse plants known areAntillean, and about 10% are Mexican endemics, for Mexico (about 22,000 species) (Rzedowski, 1993), with the remainder more widespread (Wendt, 1993), the Lepidoptera likewise are exceedingly numerous allof which compares favorably with the Lepidoptera and biodiverse. Thus far, about 14,385 species of figures. Lepidoptera are known for Mexico, including butter- Mexico being so exceedingly diverse in habitats, flies (. 1,800species). The following notes have been in part due to the many mountain ranges, has more organized to briefly summarize what is known thus endemic species than do most countries of the world, far of Lepidoptera in Mexico. Besides Lepidoptera and this component forms perhaps a quarter of the species described from Mexico, the Mexican totals for Lepidoptera inMexico. The least well known of each family include all species from the USA border Mexico are the northern region, near the border with and those described from Guatemala and Belize, since the , andyucatan, although for smaller these undoubtedly also occur in nearby areas of moths most all of Mexico is relatively little known. Mexico for the most part. The greatest biodiversity inMexico is in Chiapas, as The Mexican Lepidoptera fauna is particularly canbe seen in the number ofplant species (Rzedowski, species rich due to 5 main faunalelements to be found 1993) and butterflies (Llorente et al., 1996). in the region: 1) endemic species (about 25%); 2) The notes on Lepidoptera families herein are western North American species intruding south modlfied in part from the chapter on Lepidoptera from the United States border areas and also into the (Heppner, in press) in the Encyclopedia of Entomol- higher mountains of central Mexico (about 15%); 3) ogy, with emphasis on the Mexican fauna. Mexico eastern North American species intruding south from species totals are taken from the catalogs of Atlas of the United States border area of southeastern Texas Neotropical Lepidoptera (Heppner, ed., 1984,1995, and into parts of northeastern Mexico (about 5%); 4) 1996), and from other works by the author (Heppner, fauna in the Yucatan (about 15%); and 5) 1991,1998,in press) andothers. Some informationon CentralAmerican fauna in Chiapas and nearby areas Mexico Lepidoptera was also obtained from the over- of southernMexico (about 40%). The last element, the view summaries organized by Llorente et al., ed~. Central American component, also includes many (1996-2000), from sections authored by Balciizar and species that are more widespread in the Neotropics, Beutelspacher (2000a,b),Becker (2000),Davis (2000), even as far as into the Amazon, and undoubtedly 172 Volume 16, No. 4, December, 2002, INSECTA MUNDI

Le6n (2000), Llorente et al. (1996), Miller (2000), Razowski (1996), Solis (1996), and Warren (2000). Infraorder The arrangement of the classification of Lepi- Division doptera herein follows a progression somewhat in line Section Nepticulina with our knowledge ofhow the evolved, from the Andesianoidea most primitive moths that resembled Micropteri- gidae, to what are now considered the most advanced lepidopterans, the (Heppner, 1998,2000; Palaephatoidea Kristensen, 1999). Yet, advanced features are also Section Incurvariina foundin other groups, such asthe , with their Incurvarioidea unique wing locking mechanism that nearly ap- Division proaches the similar wing locking found in wasps Section Tineina (Hymenoptera), and butterflies, where some have Subsection Tineina evolved advanced features like the chordotonal organs found in wing veins of (). The taxonomic listing below gives an overview of Copromorphoidea the main groupings and superfamilies of Lepidoptera (not all groups listed occur in Mexico). Overall, the Immoidea order has 33 superfamilies and 125famhes (Heppner, 1998, 2003; Heppner, ed., 2000). Butterflies are in Pterophoroidea , including skippers which sometimes Subsection Sesiina are placed in their own superfamily, Hesperioidea. Groups preceeding Ditrysia contain the most primi- tive Lepidoptera, and among these, the first 3 subor- Section Cossina ders contain the most archaic relicts of ancient lepi- Subsection Cossina dopteran lineages still extant (many of these only occur in ancient biotic refugia such as South , Castnioidea , and Patagonia). Tortricoidea This is Entomology Contribution No. 944, Bureau Subsection of Entomology, Nematology and Plant Pathology, Calliduloidea Division of Plant Industry, Florida Department of Uranioidea Agriculture and Consumer Services. Papilionoidea Order LEPIDOPTERA Suborder Zeugloptera Micropterigoidea Sphingoidea Suborder Aglossata Agathiphagoidea Suborder Heterobathrniina There is some controversy in terms of how many Heterobathmioidea families and superfamilies there are and how they Suborder should be arranged. For example, some other recent Cohort Dacnonypha classifications for Lepidoptera use 46 superfamilies, Infraorder Dacnonypha splitting many odd families to their own monobasic Eriocranioidea superfamilies (Kristensen, 1999).However, thefoun- Infraorder Lophocoronina dations of the primitive groups (non-Ditrysian groups) Lophocoronoidea are fairly well established now, although new discov- Cohort eries can still come to light from remote tropical Subcohort Myoglossata regions of the world: for example, one small family of Infraorder Neopseustina primitive moths, the Neothoridae, was only discov- Neopseustoidea ered within the past 25 years and was based on a Subcohort single known specimen from . Also, a small Infraorder group of South American moths misplaced in Mnesarchaeoidea INSECTAMUNDI, Vol. 16, No. 4, December, 2002

was just found to be monotrysian and the new family Pterophoroidea Andesianidae was formed (Davis and Gentilli, 2003). Sesioidea The notes below summarize information on the Zygaenoidea 125 extant families of Lepidoptera in the world, and Cossoidea how this relates to the Mexican fauna. The species Castnioidea totals given for each family have been updated to Tortricoidea include all valid species described to the year 2000, Uranioidea plus estimates of how many occur in Mexico. In the Geometroidea world, the total number of knownLepidoptera species Papilionoidea comes to about 156,100 species. An estimated addi- Drepanoidea tional 100,000 species remain to be discovered and Bombycoidea named, mainly from tropical regions of the world, Sphingoidea including many of the most imperiled habitats. Noctuoidea During the decade from 1990-2000, about 8,500 Lepidoptera species were described from all regions, Totals for Mexico 14,383 22,440 particularly from remote areas of temperate Central Asia and from the tropical forests of Southeast Asia Primitive moths and Central Africa. Thus, the rate of new discoveries and descriptions of new species remains fairly steady, The families considered the most primitive of all averaging about 850 species per year among Lepi- lepidopterans, in the groups up to Ditrysia, include doptera. While most of these new species are moths, the first 22 families. The first 8 superfamilies are some larger butterflies are involved aswell, and even particularly primitive and of these, the first 3 super- a few swallowtails (Papilionidae) from such remote families are all relictual base lineages of the Lepi- regions as parts of China and New Guinea. Mexico doptera and also have only one family in each likewise has a great many undescribed species, par- superfamily and each suborder: ticularly among the smallest micro-moths such as (Micropterigoidea, suborder Zeugloptera), Agathi- leafminers. Biological and morphologicalinformation phagidae (Agathiphagoidea, suborder Aglossata), and can be obtained from such works as by Bourgogne Heterobathmiidae (Heterobathmioidea, suborder (1951), Eaton (1988), Hering (1925), Portier (1949), Heterobathmiina). These archaic superfamilies are Scoble (1992), Stehr (1987), Vane-Wright and Ackery all small moths (not over 16 mm in wingspan), (1984), Watsonand Whalley (1975), andYoung(1997). concentratedmost inwhat are now considered refugia Keys to families are in Heppner (1996, 1998) and regions of the world where the oldest faunas remain, Holloway et al. (1987). so-called Gondwanaland distributions, notably in In Mexico, the Lepidoptera fauna totals about South Africa, Australia, New Zealand, and Chile, 14,385 described species and probably has an actual although some are more widespread like the fauna, once all species are hscovered and described, of Micropterigidae which occur in all regions. No archa- about 22,500 species, possibly more. The table below ic moths are known from Mexico, although it is summarizes these figures for groups known to occur possible some Micropterigidae may eventually be in Mexico: found in Mexico. The other primitive groups, from Hepialoidea to Superfamily Species lrnown Expected total Incurvarioidea, include some species from Mexico, Micropterigoidea -- 1 although such leafminer families as will Eriocranioidea -- 1 have a larger fauna in Mexico once all species are Hepialoidea 15 20 described. Some of the include the largest Nepticuloidea -- 50 moths among the primitive families (up to about Tischerioidea 1 10 200mm in wingspan for some Mexican species). Incurvarioidea 16 35 Tineoidea 147 665 Gelechioidea 396 1,825 Copromorphoidea 6 10 Yponomeutoidea 35 110 Immoidea 2 5 Pyraloidea 1,375 3,045 174 Volume 16, No. 4, December, 2002, INSECTA MZTNDI

Micropterigoidea ly. None is inMexico, but it is possible that there could be unknown species in northern dry regions of Mexico. This superfamily is monobasic, including only the single family Micropterigidae. Lophocoronoidea

1. Micropterigidae (mandibulate archaic moths) Representing the infraorder Lophocoronina, this total 130species worldwide. There are 2 subfamiles. monobasic superfamily contains the small archaic Adults are minute to small and diurnal. Larvae feed family, Lophocoronidae. on , liverworts, or detritus. None is known for Mexico, but 2 species have been described from the 6. Lophocoronidae (Australian archaic sun moths) Andes, in Chile and Ecuador, while 2 species are have 6 known species in Australia. Adults are small known from the United States. and crepuscular in sclerophyll eucalyptus woods. Biologies and larvae remain unknown. No Agathiphagoidea species are known.

This is a monobasic superfamily, containing the Neopseustoidea relict family, Agathiphagidae. This first group of the Myoglossata, infraorder 2. Agathiphagidae (kauri moths) totalonly 2 known Neopseustina, includes only the single archaic family species, from Australia and . Adults are small and . diurnal. Larvae are -borers of kauri pines (Araucariaceae). No New World species are known. 7. Neopseustidae (archaic bell moths) include 9 known species (6 from Southeast Asia and 3 from Heterobathmioidea Chile). Adults are small (bell-shaped when at rest) and crepuscular or diurnal. Biologies and larvae A monobasic superfamily, including only the remainunknown, but species in Chile are thought to single family Heterobathmiidae. possibly feed on native bamboos. None is known for Mexico. 3. Heterobathmiidae (Valdivian archaic moths) include about 9 species, with 3 named thus far all Mnesarchaeoidea from southern . Adults are small and diurnal. Larvae are leafminers of trees This first superfamily of the infraorder Exoporia, (). None is known for Mexico. of the Neolepidoptera, contains a single family of primitive moths, the Mnesarchaeidae. Eriocranioidea 8. Mnesarchaeidae (New Zealand primitive moths) The superfamily Eriocranioidea contains 2 fami- occur only in New Zealand, with 8 known species. lies, together comprising the infraorder Dacnonypha. Adults are small and diurnally active in forest clear- ings and near . Biologies and larvae remain 4. (sparkling archaic sun moths) are unknown. No New World species are known. a Holarcticfamily of 25 species, about halfin Europe and half in . Adults are small and Hepialoidea diurnal. Larvae are leafminers on a variety of hard- wood trees and other plants. None is known for This superfamily comprises the second superfam- Mexico, however it is conceivable that there could be ily of Exoporia, and the first group of lepidopterans some species inMexico. with several families, although most species are in Hepialidae, the ghost moths. The superfamily com- 5. (archaic sun moths) are prises 518 known species worldwide. Only Hepialidae very similar to Eriocraniidae, and include only 4 are known for Mexico, about 15species. species, all North American except for one in the Palearctic. Adults are small and thought to be diur- 9. Neotheoridae (Amazonian primitive ghost moths) nal. Larvae are blotch leafminers on include a single species from the Mato Grosso area of () for the single species known biological- Brazil (2 more species have been discovered recently). INSECTA MUNDI, Vol. 16, No. 4, December, 2002 175

Adults are medium size. Biologies and larvae remain idae and , and forms the first group of the unknown. No species are known for Mexico. Monotrysia. The world totalis 990 species. Mexico has only 4 species recorded thus far, but the actual fauna 10. Anomosetidae (Australian primitive ghost may be over 60 species. moths) comprise a single with 1known species from Australia. Adults are small. Biologies and larvae 15. Nepticulidae (pygmy moths) comprise 868 spe- remain unknown. No New World species are known. cies from all faunalregions, although most (over 510 species) are from the Palearctic region; actual world 11. Prototheoridae (African primitive ghost moths) fauna likely exceeds 1,200 species. Two comprise 12 species from southern Africa. Adults are are used. Adults are minute and diurnally active. small. Biologies and larvae remain unknown, but a Larvae are leafminers, usually blotch mines, al- has been found in . No New World species though some also mine other plant parts. Many are known. hostplants are used. No species are actually described from Mexico, but 80 species are known north of the 12. Hepialidae (ghost moths, or swifts) comprise U.S. border and some may occur in northern Mexico about 537 species from all faunal regions, but most as well. Sampling of mines will unquestionably are from Australia, South Africa, and Chile. Adults result in the discovery of numerous Mexican species are medium size to very large and typically crepuscu- (perhaps as many as 50 species). lar or nocturnal, but a few are diurnally active. Larvae feed as borers on , trunks or under bark 16. Opostegidae (eye-cap moths) total 122 known of trees, various bushes, or grasses, or even leaf litter. species from allfaunal regions and especially Austra- This is the first family with a number of species lia; actual fauna probably exceeds 175species. Adults already described from Mexico, plus some species are minute to small and diurnally active. The few from the United Statesborder probably also occur in larvae known are leafminers, but some are stem northernMexico. The Mexican total thus far amounts borers. Hostplants are in several plant families. Four to atleast 15species, including some described from species have been described from Mexico, but more Guatemala and further south in . can be expected with thorough rearing of leafminers from all regions of Mexico (perhaps as many as 12 13. Palaeosetidae (miniature ghost moths) com- species). prise 7 species worldwide (5 in Assam, Thailand and Taiwan, 1in Australia, and 1in Colombia). Adults are Tischerioidea small and crepuscular or active during the day in dark forested areas, typically near wet moss-covered This third group of Monotrysia, the superfamily rock faces. Biologies remain unknown but larvae of a Tischerioidea, includes the single family Tischeri- species inTaiwan arethought to feed onmosses. None idae. is known from Mexico. 17. Tischeriidae (trumpet leafminer moths) total 81 Andesianoidea known species from all regions except Australia, with most species in the Nearctic (48 species); actual fauna This monobasic superfamily includes the primi- likely exceeds 125 species. Adults are small and tive Monotrysian family Andesianidae, recently de- diurnally active. Larvae are leafminers, usually trum- scribed.for species previously misplaced in Cossidae pet-shaped mines or blotch mines, on a variety of (Davis, 2003). hostplants. One species has been described from Mexico. As with other leafminer groups, many more 14. Andesianidae (Valdivianforest moths) totalonly can be expected to be found in Mexico (perhaps as 3 known species from southern Andean Nothofagus many as 10 species). forests in and Chile. Biologies remain unknown. None is known for Mexico. Palaephatoidea

Nepticuloidea The monobasic monotrysian superfamily Palaephatoidea contains one family from southern The superfamily Nepticuloidea includes two fam- refugia regions of the world (except South Africa), the ilies of the tiniest leafminer moths known, Nepticul- . 176 Volume 16, No. 4, December, 2002, INSECTAMUNDI

18. Palaephatidae (Gondwanaland moths) total 39 but of average length in females (sometimes thick- described species (28 from Chile andhgentina, and ened). Adults are usually diurnal but a few are 11 from Australia), but atleast 25 more undescribed crepuscular. Larvae are leafminers, but in later species are knownfrom Australia alone; actual fauna instars change to casebearers. Hostplants include probably exceeds 90 species. Adults are small to several plant families. Two species have been de- medium size. Biologies little known but adults pre- scribed from Mexico and there likely are more to be sumed diurnally active. Larvae tie hostplant twigs discovered, and atleast a few more can be expected to together (onverbenaceae andproteaceae). No species occur in northern Baja California where the Califor- are known for Mexico. nian fauna intrudes.

Incurvarioidea 23. (shield bearer moths) total 106 species from all regions, with more than half the This superfamily is the sole group comprising species split between North America (3 1species) and section Incurvariina of the Monotrysia. The super- Australia (36 species); actual fauna probably exceeds family has 589 known species. Only 16 species are 200 species. Adults are minute to small and diurnal. known for Mexico thus far but more may be discov- Larvae make serpentine leaf mines at first, then ered. make blotch mines in later instars. Hostplants in- clude several of hardwood trees and bushes. No 19. (leafcutter moths) total about 116 Mexican species have been found but some are likely species from all regions, but most are Palearctic (64 to be discovered with further sampling of leaf mines species), divided into 2 subfamilies. Adults are small (perhaps as many as 10 species). and mostly diurnal in shaded habitats. Larvae are leafminers at first, then switch to leaf skeletonizing Ditrysia on a variety of hostplants; some are casebearers or shoot borers. Although the only Neotropical species This division of the infraorder Heteroneura com- recorded thus far are from southern South America, prises all the remaining Lepidoptera, with 20 super- a number of species from the western United States families involved for the 102families included for the are now included in this family, so some may eventu- world among these higher evolved lepidopterans. ally also be discovered in northern Mexico. Tineoidea 20. (gall moths) total only 7 species, with 5 species from southern South America and 2 The superfamily Tineoidea includes the most from South Africa. Adults are small and probably primitive of the Ditrysia. The superfamily total comes diurnal. In Argentina larvae are gall makers on to about 5,730 known species for the world. InMexico, Schinus (Anacradiaceae). No species are known for about 147 species are known, but as many as 665 can Mexico. be expected in total.

21. ( moths) total 65 species, most- 24. (tube moths) total 270 species in ly western Nearctic. There are 2 subfamilies. Adults the New World, mostly in the large genus Acrolo- are minute to small and diurnal. Larvae are seed, phus; actual fauna likely exceeds 350 species. Two stalk, or stem borers; rarely gall makers. subfamilies are used. Adults are small to medium Hostplants arevarious yucca plants (Agavaceae) and size, with rather robust bodies and usually large other species are on hardwood trees and bushes. At recurved labial palpi. Adults are mostly nocturnal, least 3 species are known from Mexico, but some but some may be crepuscular. Larvae are feeders, species known for southern Arizona and New Mexico mostly of grasses, and construct long underground also occur in northern Mexico, thus bringing the silken tubes to feedonhostplant roots. Mexico harbors Mexican total to about 14species. a number of species and more are likely to be discov- ered: the current total for Mexico is about 76 described 22. (longhorned fairy moths) comprise 295 species. species worldwide, but most are Palearctic (143 spe- cies), with 2 subfamilies; actual fauna probably ex- 25. ( moths) comprise the first very ceeds 400 species. Adults are minute to small; anten- large family of Lepidoptera, with about 2,160 de- nae extremely long (often 2x wing lengths) in males, scribed species. The actual world fauna probably INSECTAMUNDI, Vol. 16, No. 4, December, 2002 exceeds 4,000 species. The family is divided into 16 active diurnally. Larvae are leafminers, becoming subfamilies. Adults are minute to medium size and leaf skeletonizers in later instars; host records are nocturnal or crepuscular; rarely diurnal. Larval hab- mostly in and Aceraceae. The family has its vary greatly, but most are detritus feeders, some been erroneously called in recent making cases, tunnels, or silken tubes; also, odd European literature. None known for Mexico. groups are coprophagous, keratophagous,woolen feed- ers, and even myrmecophilous and termitophilous 30. (bristle-legged moths) are larvae are known. Included among Tineidae are some a small family of only 8 known species (3 in North of the most well known household species, such America, 1in Guatemala, 1in Colombia, 1in Europe, as clothes moths and grain moths. Besides the cosmo- and 3 in Asia). Adults are small and diurnal. Larvae politan household pests (clothes moths), Mexico has are leaf skeletonizers on or . about 35 known species, yet many more are to be At least one species should be in southern Mexico, expected (perhaps as many as 75 species). probably the same as the one from Guatemala.

26. (Old World spiny-winged moths) 31. (Douglas moths) comprise only 28 are a small family of 212 known species, mostly known species, mostly Palearctic (20 species). Adults African (120 species) and Oriental (66 species), now are minute to small and crepuscular or diurnal. dividedinto 2 subfamiles. Adults are small to medium Larvae are leafminers or borers in petioles or stems. size and mostly diurnally active, or may be crepuscu- Hostplants are knownin , Labiatae, and lar. Biologies are little known but some larvae report- Rosaceae. One species is known from Ecuador, but ed to tunnel in the soil, possibly feeding on roots or none have been recorded from Mexico. detritus. No New World species are known. 32. (ribbed-cocoon maker moths) 27. Psychidae (bagworm moths) total 1,001known total 247 species worldwide, with most species being species, mostly Palearctic and African, with only 88 Nearctic (103 species) or Palearctic (86 species); most knownfor the New World; actualfauna likely exceeds species are in the genus Bucculatrix. Adults are 1,200species. The family is now dividedinto 6 subfamil- minute to small and mostly diurnal. Larvae are ies. Adults are minute to medium size and mostly leafminers, with some changing to external leaf diurnal or crepuscular. Larvae are mostly leaf feeders skeletonizing in later instars, but a few are gall or feed on lichens, all making distinctive types of makers or stem miners. Pupation is in a white larval cases, or bags. Pupation is within the larval spindle-shaped cocoon with a ribbed surface, unique to case and females often remain there in a wingless or the family. Many different hostplants are known but larviform shape, using pheromones to attract the many feed on Compositae. Some species are economic. winged males. Some species are economic and many Four species are known for Mexico, but many more are general plant feeders. Mexico has about 17known are likely to be found (perhaps as many as25 species). species, but many more to be described. 33. (leafminer moths) comprise the 28. (tropical lattice moths) total major leafmining family of Lepidoptera, with about 26 species, mostly Neotropical, but recently with 1,740 species from all regions. The actual world total some Southeast Asian additions; actual fauna proba- probably exceeds 6,000 species. There are 3 subfami- bly exceeds 50 species. Adults are small to medium lies. Adults are minute to small and active diurnally. size. Adult activity mostly nocturnal, but some are Larvae are usually leafminers but change form in diurnal. Biologies unknown except for one Neotropical later instars (hypermetamorphism) from sap feeders species with casebearing larvae that feed on fungi. with reducedlegs, to later instars with a 3-proleg pair None described from Mexico, but 2 species occur in larval form, which is unique among the Microlepi- southern Mexico and another 2 remain to be de- doptera. Later instars sometimes feed externally as scribed. leaf skeletonizers. Hostplants include a great number of plant families. A few species are of economic 29. Arnphitheridae (double-eye moths) are an un- importance, such asthe leafminer (Phyllocnis- usual and small family of mostly tropical moths, tis citrella) introducedfrom China. The current total totaling 35 species, mostly Indo-Australian (1 genus for Mexico is about 13species, but numerous species occurs in Europe and another in South America). can be expected to occur in Mexico, and will be Adults are small, with eyes usually divided. Adults 178 Volu.me 16, No. 4, December, 2002, INSECTAMUNDI discovered once more leafminers are reared and the es (Gramineae) and related plant groups like Juncace- species described (perhaps as many as 450 species). ae and , but other plant families are also utilized. None recorded from Mexico, but atleast the Gelechioidea sugarcane leafminer, Dicranoctetes saccharella (Bus- ck), is likely to also occur in Mexico. This superfamily comprises a huge group of micro-moths, all having a scaled haustellum and 37. (lance-wingmoths) totalonly 11 unspined pupa. There has been considerable turmoil species, mostly Mediterranean, plus 2 are from South in recent classifications proposed for the group, either Africa. Adults are small and may be mostly crepuscu- with a large family or with conglomer- lar. Larvae are root borers as far as is known. Only ation of various groups into an enlarged family Ela- recorded hostplants are in Compositae. One species chistidae. In the classification adopted herein, there introduced into western North America for control of are 11 families recognized, the largest being Oecoph- knapweed. None recorded in Mexico. oridae and . The entire superfamily en- compasses about 18,230 described species worldwide, 38. Gelechiidae (twirler moths) are a very large but the actual fauna may well exceed 45,000 species, family, with over 4,830 species described, however, since there are huge numbers of undescribed tropical possibly with a fauna exceeding 10,000 species world- species. In Mexico, there are 396 described species, wide. arrangements have varied but now but as many as 1,825can be expected to occur there. include 4. Adults are small and mostly nocturnal but some are diurnal or crepuscular. Larvae have a range 34. Oecophoridae (concealer moths) are a large of feeding habits but most areleaf skeletonizers, using family of about 7,550 described species from all faunal a leaf fold or leaf tie as protection. A large variety of regions, with most species being from Australia; the plants are usedas hosts. Some species are economical- actual fauna may well exceed 12,000 species world- ly important. A number of species are recorded for wide. There are 10subfamilies recognized. Adults are Mexico (175 species), but the actual fauna undoubted- small to medium size and mostly nocturnal but some ly will exceed 1,200 species when all are discovered in are diurnal or crepuscular. Larvae include many leaf Mexico. litter feeders, but also leaf tiers, leaf webbers, bark feeders, and a few leafminers. Hostplants include a 39. (scavenger moths) totalover 296 large number of plant families, plus lichens, fungi, species worldwide, with many known from North and detritus or leaf litter. Few are economic. Mexico America and Europe; actual fauna probably exceeds has 172 species recorded (actualMexican fauna may 600 species. There are 2 subfamilies. Adults are small exceed 350 species). and nocturnal as far as is known. Larvae are scaven- gers or detritus feeders, sometimes feeding on plant 35. (tropicallonghorned moths) to- fruits, , or , among a number of plant tal about 1,038 described species, mostly tropical families, but few are known biologically. At least one Oriental, but also with one group in the Palearctic; species lives with coccids (Homoptera) but predation actual fauna probably exceeds 1,500 species. There on the coccids has not been confirmed. Mexico has 9 are 4 subfamilies. Adults are small and mostly diur- species recorded, but the true fauna likely will be near nal and many have the habit of holding the long 100 species. antennae together to the front when at rest. Larvae may mostly be leaf litter feeders or leaf tiers, but few 40. (casebearer moths) comprise species are known biologically. A few varied host- about 1,525species worldwide, with most being Pale- plants are recorded. None is recorded in the New arctic andinthe genus, . The actual world World (the Neotropical genus Deoclona has been fauna likely exceeds 2,000 species. Most are in sub- transferred to Oecophoridae). family Coleophorinae, while non-casebearers are in Batrachedrinae. Adults are small and may be mostly 36. (grass miner moths) comprise about crepuscular but many are diurnal. Larvae make 723 species worldwide, but most are Palearctic (472 small cases (except for Batrachedrinae), often dis- species). Two subfamilies are used (or only tribes). tinctly shaped for each species, skeletonizing host Adults are small and often crespuscular or nocturnal, , but some are seed borers, leafminers, or stalk but some are diurnal. Larvae are leafminers (some- borers, or skeletonize leaves beneath frass webs. Afew times gregarious) or stem miners, especially on grass- Batrachedrinae are predacious on scale INSECTA MUNDI, Vol. 16, No. 4, December, 2002 179

(Homoptera). Various hostplants are used. A few beneath bark), but few biologies are known. Host- species are economic. Ovovivipary has been recorded plants include , , , for a few species. Mexico has 8 known species, but , and . InMexico, there are 2 there likely are 30 or more species. species known.

41. ( moths) total 127 species 46. Alucitidae (many-plumedmoths) total about 184 worldwide, with about halffrom the Palearctic. Adults species worldwide. Adults are small; wings with all are small and mostly diurnal or crepuscular. Larvae veins as separate wing clefts to near the wing bases mostly leafminers, but some are borers inflowers and (rarely split only to center of wings). Adults active in stems, or gall makers. Hosts are only known in deep shade, or may be crepuscular. Larvae are borers . Mexico has 5 species recorded. or gall makers as far as is known. Various hostplant records are known. There are 2 species recorded in 42. Agonoxenidae (palm moths) are a small family Mexico. of 68 known species from all faunal regions. Adults are small and diurnal. Larvae are leaf skeletonizers, 47. (fruitworm moths) total about 279 or borers in leaves, stems, and fruits; rarely gall- species from all regions, but most are Australian and makers. Hostplants mostly in Rosaceae or Palmae. A South Pacific. Adults are small to medium and noc- few are economic. None recorded from Mexico but turnal or crepuscular. Larvae are borers in fruits, there likely are at least 5 species to be found. seeds, buds, or trunks and limbs, but a few are leafminers. Hosts include a variety of plants. A few 43. (cosmet moths) total over species are economic. Mexico should have about 3 1,540 species worldwide, but the extant fauna may species ranging from nearby areas in the United encompass 3,500 species. There are 3 subfamilies. States. Adults are small and mostly diurnal, but some are crepuscular. Larvae mostly leafminers or needle- 48. (fringe-tufted moths) total 102 miners, but some are borers of various plant parts; a species, with many being Palearctic (36 species) and few are predacious on Homoptera. Hosts are varied. Australian (23 species), in two subfamilies. Adults are Ovovivipary recordedin a few species. Some economic small and diurnal or crepuscular. Larvae are leafmin- species are known. A large group, with 27 species ers, leaf skeletonizers, or borers of seeds, fruits, or known for Mexico (actual fauna probably more than buds; a few are gall makers. Host records include 100 species). several plant families. One species is recorded for Mexico. 44. (flower moths) total about 523 species worldwide, but mostly known from Europe. Yponomeutoidea Adults are small and diurnal but some may be crepuscular. Larvae are skeletonizers onleaves, buds, This superfamily of varied moths includes 9 flowers, hiding under webbings. Many plant families families, some of which have been transferred from are used as hosts, plus some on lichens and mosses. Tineoidea, while other previously contained groups Not recordedfor Mexico but the probably are atleast are now in other superfamilies. About 1,853 species 10 species to be found. are known in the superfamily. In Mexico, 35 species are known and the actual fauna may be ashigh as 110 Copromorphoidea species.

This superfamily comprises 4 families related 49. Ochsenheimeriidae (cereal stem moths) in- especially by larval characters. The world fauna clude only 17species from the Palearctic (1 species is totals 623 species. In Mexico, about 6 species are from Kashmir), with 1species introducedinto North known, which may eventually total 10 species. America. Adults are small and diurnal. Larvae are leafminers, but become stem borers in later instars, 45. (tropical fruitworm moths) primarily on grasses (Gramineae), sedges (Cyperace- are a small family of 58 species, mostly tropical; ae) and rushes (). One species is economic. actual fauna probably exceeds 100 species. Adults are None is known for Mexico. small to medium and nocturnal. Larvae are leaf feeders using a leaf web, or are borers (one feeds 180 Volume 16, No. 4, December, 2002, INSECTAMUNDI

50. (sedge moths) total 431 spe- and some mine in various plant parts. Numerous cies from all regions, mostly in the genus Glyphip- plants are recorded as hosts. Several species are terix, with the largest number from the Australian- economic. None recorded from Mexico. New Zealand region; actual world fauna probably exceeds 600 species. There are 2 subfamilies. Adults 55. (Lyonet moths) total 264 species, are small and diurnal, usually in proximity to the mostly Holarctic; the actual fauna probably exceeds hostplants. Larvae are mostly borers in seeds, stems, 600 species. There are 3 subfamilies. Adults are or leaf axils, and a few are leafminers, but most minute to small and mostly crepuscular and noctur- tropical species are unknown biologically. Hostplants nal. Larvae are blotch leafminers; rarely mining are mostly sedges (Cyperaceae), rushes (Juncaceae), stems. Many plant groups are recorded ashosts. Afew and grasses (Gramineae), plus a few other plant species are economic. Mexico has at least 6 species. families. Mexico has 6 species, but specialized collect- ing should verify a fauna of at least 50 species. 56. (false diamondback moths) include 96 species, mostly Palearctic. Adults are small and 51. (diamondback moths) include 386 crepuscular or diurnal. Larvae mostly leafminers, species worldwide; actual fauna probably exceeds 600 but some are borers in seeds, stems, and flower buds. species. There are 4 subfamilies. Adults are small to Several hostplant groups are used, but mostly on medium size and mostly nocturnal or crepuscular, Compositae. Very few are economic. Mexico probably but some are diurnal. Larvae are leaf skeletonizers, has one species entering from Guatemala, and more but most remainunknown biologically. Hosts include may be found in northern areas. different plant groups. A few species are economic. Including the cosmopolitan diamondback from 57. (sun moths) are a small family of Europe ( xylostella), the family has only 3 56 species, mostly Neotropical (31 species); actual species noted for Mexico, but the actual fauna should fauna probably exceeds 100 species. Adults are small be at least 10 species. and diurnal. Larvae are mostly leaf skeletonizers, but some are borers in fruit racemes. Several plant 52. (tropical ermine moths) include 48 families are used ashosts. Mexico has 13species and species, mostly tropical and in the genus Atteva (the more are likely to be found. single partially non-tropical species known occurs across the southern United States andinto the Carib- Immoidea bean and Mexico); actual fauna probably at least 60 species. Adults are small to medium size and are This is a monobasic superfamily for the single diurnal or crepuscular. Larvae are leaf webbers and small family . It is somewhat related to leaf skeletonizers on and Simaroubaceae. Pyraloidea and may represent relatives of common Minor economic species occur on Ailanthus trees in ancestors. and the United States. At least 3 species are in Mexico. 58. Irnmidae (imma moths) comprise 246 species, all tropical, and mostly Indo-Australian and South Pacif- 53. Yponomeutidae (ermine moths) total 395 spe- ic; actual fauna likely exceeds 450 species. Adults are cies worldwide; actual fauna likely exceeds 500 spe- small to medium size and diurnal, but some may be cies. Three subfamilies are recognized. Adults are crepuscular. Larvae are leaf feeders, but only 3 small to medium size and mostly nocturnal. Larvae species have biological data. Hostplants are inMyrtace- are leaf skeletonizers and leaf webbers, but some are ae, Podocarpaceae, and Violaceae. None described leafminers or needleminers. Hosts include many from Mexico, but at least 2 species described from plant families. Afew species are economic. Mexico has Guatemala should also be found in Chiapas. 3 species recorded, but more should be found in the central mountains. Pyraloidea

54. (shiny head-standing moths) This superfamily includes 3 families but mainly include 160 species, mostly from Holarctic regions; the very large family, . The superfamily actual fauna probably exceeds 450 species. Adults are comprises about 17,312 species worldwide. Some spe- small and mostly crepuscular, but many may be cialists split the group by placing each pyraloidfamily diurnal. Larvae are leafminers and needleminers, in its own monobasic superfamily (Hyblaeoidea, INSECTA MUNDI, Vol. 16, No. 4, December, 2002 181

Thyridoidea, andpyraloidea). InMexico, about 1,375 Mexico, the actual fauna probably is at least 3,000 species are known and perhaps 3,045 will eventually species. be recorded there in this superfamily. Pterophoroidea 59. (teak moths) are a small tropical family of 18 species, mostly Indo-Australian and in This superfamily includes only 3 families, but the genus (1 pantropical species is also most species are in the . About 1,309 established in southern Florida). Adults are medium species are known for the superfamily. In Mexico, 43 size; body usually robust. Adults are diurnal or species are recorded thus far, but perhaps 80 species perhaps crepuscular. Larvae are leaf rollers. Host- will eventually be found there. plants are in and . One economic species is the teak leafroller, which also 62. Tineodidae (false plume moths) include only 11 occurs in Mexico. species, all from Australia. Adults are small and possibly diurnal or crepuscular. Larvae are leaftiers, 60. (picture-winged leaf moths) total 794 but most are not known biologically. Only recorded species worldwide, nearly all tropical, with nearly hostplants are in Euphorbiaceae and Oleaceae. No half the species Indo-Australian (only a few species New World species known are in the Nearctic and Palearctic regions); actual fauna likely exceeds 1,200 species. There are 6 63. Oxychirotidae (tropical plume moths) include subfamilies. Adults are small to large and diurnal or only 6 species, all Indo-Australian and South Pacific. crepuscular. Larvae are leafrollers (one Australian Some specialists include this family as part of the species is gregarious), or borers in stems and flower Tineodidae. Adults are smalland may be crepuscular. racemes; a few are gall makers. Several hostplants of one species feeds on seeds ofwhite are used. Only a few species are economic. Mexico has (Avicenniaceae); remainder unknown biologically.No 24 species, but the complete fauna should approach at New World species known. least 45 species. 64. Pterophoridae (plume moths) comprise about 61. Pyralidae (snout moths) comprise the third 1,292 species worldwide, with about a third being largest family of Lepidoptera, with about 16,500 Palearctic; actual world fauna probably exceeds 1,800 described species, but a probable fauna of at least species. Subfamilies number 5. Adults are small, 25,000 species worldwide; 19subfamilies are used in usually with hindwings split into 3 fringed plumes; the classification. Recent specialists have talcen to forewings often entire or split into 2 parts near the splitting Pyralidae into its two major lineages, asthe termen (a few species have both wings entire). Adults separate families and Pyralidae, but this mostly nocturnal but some crepuscular or in shaded reversion to what was done over 100 years ago is areas during the day. Larvae mostly leaf feeders, or unnecessary and amply clarified by using the "group" miners and borers of various plant parts; a few are category below the family level within a single family gall makers. Hostplants include many families. Sev- Pyralidae. Adults are small to large and mostly eral species are economic. Recorded species in Mexico nocturnal, but some are crepuscular and a few are total about 43 species, while the actual fauna probably diurnal. Larvae are mostly leafrollers or leafwebbers, will exceed 80 species. but many are borers, root feeders, detritus feeders (including stored products pests), and a few are Sesioidea leafminers, plus rare myrmecophilous species, and even some aquatic groups making cases (Nymphuli- The Sesioidea include 4 families, although many nae). A large number of economic species are in this specialists maintain separate superfamiliesfor most family, including pests on virtually all crops and of them: , Sesiidae, , and forest trees. Hostplants are in a large number of plant . Brachodidae are thought to be the families. The family is exceedingly large in Mexico group, with close ties to Sesiidae. The world fauna also, with many species ranging through the Caribbe- totals 1,964known species in the superfamily. Mexico an and the Neotropics, plus many from the United has 117 species recorded and may have an actual States. The Mexico total recorded thus far is about fauna of 230 species or more. 1,350 species, but with the discovery of new species and more surveying both in southern and northern 182 Volume 16, No. 4, December, 2002, INSECTAMUNDI

65. Brachodidae (little bear moths) comprise 141 budworms; rarely leafminers (Millieriinae). Numer- species, most being Old World tropical, particularly ous hostplant families are recorded, but many in Indo-Australian (none is North American). There are Compositae andMoraceae. The Mexican fauna totals 3 subfamilies. Adults are small to medium size and about 21 described species, but the actual fauna will diurnal. Larvae are root feeders of grasses (in the probably exceed 50 species. EuropeanBrachodes, Brachodinae), using silken tubes, or borers in palm trunks and leaf stems, or other Zygaenoidea plants; some are leaftiers (Phycodinae). Hostplants are in Gramineae, and also Bromeliaceae, Melis- This superfamily contains the burnet moth and tomaceae, Moraceae, andpalmae. Afew are economic groups, with 6 families involved. The on palms. Southern Mexico has at least 1species. superfamily has 1,609known species from allregions. In Mexico, 127 species are recorded, and 150 may be 66. Sesiidae (clearwing moths) include about 1,325 the true faunal total. species from all regions; the actual number probably exceeds 1,800species. Subfamilies are 3. Adults are 69. (Mediterraneanburnet moths) small to medium size; antennae usually thickened include only 7 species, with 3 species from the Medi- and with a slight distal club (most also with unique terranean region of southern Europe and northern small terminal bristles on antennae). Wings very Africa and 4 species from South Africa. Adults are elongated and with unique wing-locking foldingwhere small and diurnal, but females are apterous and the margins meet. Adults are diurnal, particularly larviform. Larvae, upon hatching inside the female during morning hours. Many species mimic various cocoon, feed first on the dead female and then become wasps (Hymenoptera). Larvae mostly borers onvar- externalleaffeeders. Hostplants are inLeguminosae. ious plant parts, and a few are gall makers; 2 species No New World species known. are predaceous on scale insects (Homoptera). Host specificity is high for most species, but overall many 70. (burnet moths) comprise about 1,140 plant families are used as hosts. A number of species species worldwide, particularly welldevelopedin trop- are economic, including some major pests of fruit ical Asia (450 species) where also the largest species trees, forest trees, and grapevines. There are numer- occur; actual world faunalikely exceeds 1,500 species. ous species in northern Mexico that overlap at the There are 7 subfamilies. Adults are small to large, United States border, bringing the Mexico total to with antennae sometimes clubbed. Adults are diur- about 150 species (includes new species from an nal, often flying like butterflies, or commonly on upcoming monograph for Mexico; Eichlin, pers. flowers. Larvae are leaf skeletonizers, sometimes comm.), but the actual fauna should be about 180 feeding communally and usually atnight; often slug- species when all are discovered. like, with concealed head. Hostplants include various plant families. Economic species are mostly pests on 67. Urodidae (false burnet moths) totalonly about 80 grapevines in North America and Europe. InMexico, species, primarily Neotropical, but with a few species the fauna is about 88 species, with perhaps 100 in North America and in Eurasia. Adults are small species actually extant. and may be crepuscular or mostly nocturnal, but a few possibly diurnal. Larvae are leaf webbers or 71. Himantopteridae (long-tailed burnet moths) skeletonizers, but few are known biologically. Pupa- include 56 species from Africa and tropical Asia. tion is in a specialized filigreed cocoon. Hostplants Adults are small to medium size; hindwings with known in Lauraceae, Leguminosae, , extremely long tails. Adults are diurnal but few are Sapotaceae, and Theaceae. A few are minor pests. known biologically. Larvae are leaf skeletonizers, Mexico should have at least 3 species and likely as often communal; sometimes massing in the 1000s. many as 10 species. Known hostplants are in . No New World species known. 68. Choreutidae (metalmark moths) comprise 418 species worldwide ; actual world fauna probably ex- 72. (tropical burnet moths) total 138 ceeds 800 species. There are 3 subfamilies. Adults are species, mostly Indo-Australian but with a few in the small, with haustellum scaled. Adults are diurnal; southernunited States;actual world fauna probably usually near their hostplants and tend to hop about on exceeds 250 species. Adults are small to medium size leaves. Larvae mostly leaf skeletonizers, but some are and nocturnal, but some may be crepuscular. Larvae INSECTA MUNDI, Vol. 16, No. 4, December, 2002 183 are leaf skeletonizers and are colorful, but most are 77. (tropical carpenterworm moths) not known biologically. Hostplants are in families include 103species, mainly Afrotropical and Oriental, Celastraceae, Moraceae, andsapotaceae. Mexicoshould with 1species in the Palearctic region; actual world have about 7 species. fauna likely exceeds 150 species. Adults are small to medium size and may be crepuscular. Larvae noctur- 73. Somabrachyidae (Mediterraneanflannelmoths) nal borers on tree bark or in tree trunks, but most include only 5 species, with 3 species from southern species remain unknown biologically. Hostplants in- Europe and north Africa, plus 2 species from South clude various trees. A few have minor economic Africa. Adults are small. Adults are diurnal, but status. No New World species known. females are larviform and wingless (females of the South African species are unknown). Larvae are leaf 78. Cyclotornidae (Australian parasite moths) in- feeders ( laid on leaves, not in the female cocoon), clude only 5 known species from Australia, the most somewhat slug-like, with concealed head. Hosts are primitive of the Limacodiformes group of Cossoidea. grasses (Gramineae) and Compositae. No New World Adults are small. Adult activity uncertain but may be species known. crepuscular. Larvae flattened, with lateral protru- sions; highly evolved asparasites of leafhoppers, scale 74. Megalopygidae (flannel moths) are a New World insects, or psyllids (Homoptera) in early instars, and family of 263 species, mostly Neotropical; actual then aspredators of antlarvae. No New World species fauna likely exceeds 350 species. Adults are small to known. large and nocturnal. Larvae are leaf feeders, usually communal in early instars; slug-like,with concealed 79. ( parasite moths) to- head; with poison spines usually beneath long hair- tal 40 described species, with at least another 30 like setae. Alarge number of hostplants are recorded, known species awaiting naming; most diversity is in and some species are polyphagous. Afew are economic Australia. Two subfamilies are known. Adults are on forest trees and palms, but most are of medical minute to small and crepuscular or nocturnal; fe- importance due to urticating setae in adults and the males are sedentary. Larvae slug-like with rounded poison spines in larvae. Mexico has about 32 species. ; parasitic on fulgorids and (Homoptera). Mexico should have about 2 species, if Cossoidea not more.

This superfamily includes about 2,054 species in 80. (tropical slug moths) in- the world. In Mexico, 154 species are known and as clude 84 Neotropical species (1 species intrudes from many as 175 may occur there. Mexico into southern Arizona). Two subfamilies are known. Adults are small to medium size. Adult 75. Cossidae (carpenterworm moths) total 682 spe- activity uncertain; possibly only nocturnalor crepus- cies worldwide; actual fauna probably exceeds 750 cular. Larvae slug-like,often with translucent gelat- species. There are 5 subfamilies. Adults are small to inous wart-like surface; feeding as leaf feeders (early very large and nocturnal. Larvae are borers in trunks instars as leaf skeletonizers), but few are known and limbs. Hostplants are recorded in a large number biologically. Various hostplants are used and some of plant families. A number of species are economic larvae are polyphagous. Few have any economic pests of forest trees. In Mexico there are about 60 status. The Mexican fauna totals about 12 species. species. 81. (slug caterpillar moths) total 1,104 76. Dudgeoneidae (Dudgeoncarpenterworm moths) known species worldwide, the largest family of Cossoi- includes only 6 species in the single genus Dudgeo- dea, mostly tropical and especially biodiverse in the nea, with 2 species from Africa, 1from India, and 3 Oriental ; likely world total is near 1,600 from Australia. Adults are medium size; abdomen species or more. Adults are small to medium size. with small tympana1 organs. Adults nocturnal asfar Adults perhaps only nocturnal; many with unique as is known. Larvae mostly unknown, but one Aus- resting postures. Larvae slug-like and mostly tralian species is a stem borer on Rubiaceae. No New polyphagous leaf feeders, usually with an extensive World species known. array of poisonous stinging spines dorsally. Large numbers ofhostplants used. Few species are economic other than medically as stinging , but 184 Volume 16, No. 4, December, 2002, INSECTA MUNDI

palm defoliators can be a problem in the tropics. species, but the actual fauna probably exceeds 1,000 Mexico has about 80 species. species.

82. Chrysopolornidae (African slug caterpillar Calliduloidea moths) are a small African family of about 30 known species. Two subfamilies are known. Adults are medi- This first superfamily of the Bombycina includes um size and nocturnal. Larvae leaf-feeding and slug- 3 families of ancient lineage with many primitive like, with small spines; often colorful. Hostplants features: Ratardidae, Pterothysanidae, and Callidul- include Celastraceae. No economic species are known. idae. MareOld World and total 131known speciesfor No New World species known. the superfamily.

Castnioidea 85. Ratardidae (Oriental parnassian moths) are -like moths of the Himalayas and Oriental This superfarnily contains only the single family tropics, with 10 describedspecies,plus afew more still . undescribed. Adults are medium sized (29-87 mm wingspan), but males often much smaller than fe- 83. Castniidae (giant butterfly moths) total 170 males. Adults are diurnal. Larvae are bark feeders, known species, mostly Neotropical but with some but biologies and larvae are mostly unknown. No New species also in the Indo-Australian region; likely World species known. world total may exceed 180species. Three subfamilies are known. Adults are medium to large size and 86. Pterothysanidae (parnassianmoths) include 19 diurnal or crepuscular. Larvae are borers of monocot species from southern Africa (7 species) and Southeast plants, including grasses (Gramineae), Cyperaceae, Asia (12 species). There are two subfamilies. Adults Bromeliaceae, Marantaceae, Musaceae, and Palmae, are medium size and diurnal, possibly also crepuscu- among others. A few are economic on banana plants, lar. Biologies and larvae remain unknown. No New various palms, and sugarcane. Mexico has 14species World species known. recorded out of 134Neotropical species described thus far. 87. (Old World butterfly moths) in- clude 102 species, mostly tropical Oriental, in 2 Tortricoidea subfamilies. Adults are medium size and diurnal, flying like some butterflies in quick bursts to alter- A monobasic superfamily exclusively for the fam- nate leaf perches; resting position with wings held ily . In the past, the tortricid subfamilies together and upright. Larvae largely unknown, ex- were often considered separate families, such as cept for 2 species which are leafrollers of ferns. No Olethreutidae and Chlidanotidae, and even groups New World species known. like Ceracidae and Cochylidae which now are only tribes. Uranioidea

84. Tortricidae (leafroller moths) are a large family This superfamily encompasses both very large of 8,945 described species from all faunal regions; the and small moths among 5 families, but mostly Old actual fauna likely will exceed 12,000 species. Three World. Some of these families continue to be mixedup subfamilies are known. Adults are small (rarely with Drepanoidea and Geometroidea, and recently medium sized) and mostly nocturnal or crepuscular, many groups have been combined into Epiplemidae. but some are diurnal. Larvae mostly leafrollers, but About 814 species are known in the superfamily. In some are borers in various plant parts, including Mexico, about 30 species are known, with as many as stems, branches, flowers, and seeds; a rare few are 50 species eventually to be found there. leaf litter feeders. Most species are restricted to certain hostplants among innumerable plant fami- 88. (Oriental swallowtail moths) are a lies. Many species are economic. The Mexican fauna small family with 25 species known in 2 subfamilies, is exceedingly rich and has a large component of mostly Himalayan and East Asian. Adults are medi- species from the United States border, plus endemic um to large and diurnal. Larvae little known, but leaf species. The Mexican total thus far is about 495 feeders with waxy secretions over body. Hostplants INSECTA MUNDI, Vol. 16, No. 4, December, 2002 185 are in Clethraceae, Cornaceae, Ericaceae, Theaceae, species. The major biodiversity is in the Neotropics, and Ulmaceae. No New World species known. with over 6,500 species described, and the Indo- Australian region with about 6,670 species. The 89. Apoprogonidae (African moths) include family is dividedinto 8 subfamilies. Adults are small only a single species from South Africa. Adults are to large; a number of genera have brachypterous or medium size; antennae clubbed (hooked at tip); body apterous females. Adults mostly nocturnal, but also robust. Adults presumed diurnal, but nothing is some crepuscular and diurnal groups. Larvae mostly known of the biology or larvae. No New World species leaffeeders, typically moving in looping fashion due to known. reductions in proleg numbers. Hostplants include most plant families. Some major defoliating pests are 90. Sematuridae (American swallowtailmoths) to- known in this family. The total for Mexico should be tal 36 Neotropical species, one of which just reaches at least 2,500 species but may exceed 3,000 species. into the United States in southern Arizona. Adults are medium to large; antennae thickened, with elongated 94. Hedylidae (American butterfly moths) total only club (slightly hooked at tip). Adults nocturnal but 40 known species, all Neotropical. Adults are medium some may be crepuscular. Larvae are leaf feeders, but size and nocturnal. Larvae are leaf feeders. Host- few known biologically. Hostplants are unrecorded. plants are recorded in Euphorbiaceae, Malvaceae, Mexico has about 5 species. Streculiaceae, and Tiliaceae. Mexico has about 8 species. 91. (swallowtail moths) comprise about 120 species from all tropical regions, mostly Indo- Papilionoidea Australian (85 species); one species strays into the United States-Mexicanborder region (mainly in Tex- The butterflies, also called Rhopalocera, include 7 as). Two subfamilies are used. Adults are medium to families, including the skippers, family Hesperiidae. large and nocturnal or diurnal, with some of the Skippers are sometimes placed in their own monoba- larger diurnal species known to migrate. Larvae are sic superfamily, Hesperioidea. Extant butterflies leaf feeders. Hostplants are knownin Asclepiadaceae, number about 20,400 described species, but the actual , and Euphorbiaceae. Mexico has about 5 butterfly fauna probably willbe about 23,500 species species. worldwide when all are discovered. InMexico, 1,808 species are known thus far. 92. Epiplemidae (crenulate moths) total about 632 species worldwide, with most being Neotropical(230 95. Hesperiidae (skipper butterflies) include about species) andIndo-Australian(301 species). Two sub- 4,100 species from all faunal regions; most are Neo- families are used. Adults are small to medium size tropical, with over 2,338 species. The actual world and nocturnal. Larvae are leaf feeders or webbers. fauna probably exceeds 4,500 species. Seven skipper Hostplants are in several plant families. Mexico has subfamilies are recognized. Adults are small to medi- a fauna of about 20 species. um size and diurnal, usually with very rapid flight, but a few tropical species are crepuscular. Larvae are Geometroidea leafrollers or borers. Hostplants are primarily grass- es (Gramineae) and other monocots. Afew economic One of the largest superfamilies of Lepidoptera, species are known, particularly on rice. Mexican mainly with the large family Geometridae, but also records include 741 species. including the odd Hedylidae (some specialists place the latter family near the true butterflies, in its own 96. Papilionidae (swallowtailbutterfkes) totalabout superfamily, Hedyloidea). The superfamily totalcomes 589 species worldwide, with about 260 species being to 21,190 known species. Mexico has about 2,508 Indo-Australian, including the largest of all butter- species recorded, but perhaps as many as 3,000 flies, the . Three subfamilies are recog- species. nized. Adults are medium size to very large and diurnal. Larvae are leaf feeders; with anosmeterium 93. Geornetridae (geometer moths, also calledinch defensive glandbehind head. Hostplants include many worms) are the second largest family of Lepidoptera, plant groups. Some economic species are known, with about 21,150 described species from all faunal mainly citrus feeders. Mexico has 56 species, includ- regions; the actual fauna probably exceeds 26,500 186 Volume 16, No. 4, December, 2002, INSECTA MUNDI

ing what is the most primitive genus in the family, crepuscular activity is known in a few cases (some Baronia, in the subfamily Baroniinae. Amathusiinae and Brassolinae). Larvae are mostly leaf feeders; a few feeding gregariously. Hostplant 97. (yellow-white butterflies) total about records are among almost all higher plants. Mexican 1,275species worldwide, most being Indo-Australian records include about 440 species. (about 515 species). Four subfamilies are recognized. Adults are small to large and diurnal. Larvae are leaf Drepanoidea feeding. Various plants are utilized. Some economic species are known, particularly on cabbages amd This superfamily includes 4 families perhaps other crucifers. Mexican species total about 90 spe- most related to Geometroidea. Some recent classifica- cies. tions have greatly disorganized this superfamily, placing Axiidae in its own monobasic superfarnily 98. (gossamer-winged butterflies) total (Axioidea) and merging Cyclidiidae and Thyatiridae about 5,955species worldwide; the actual faunaprob- into as subfamilies; the latter 3 families ably exceeds 7,000 species. About 1,125 species are all have some cohesive features (e.g., abdominal Neotropical. The family has 8 subfamilies. Adults are tympana1organs) but also many unique features that small to medium size and diurnal, but a few of the provide evidence of family status. The superfarnily relict genera possibly crepuscular or only in dark comprises about 1,056known species. InMexico, only forests. Larvae mostly somewhat slug-like, with tu- 2 species are known, and perhaps 5 will eventually be bercles and short setae; head usually retractable into recorded there. thorax. Larvae feed as leaf feeders (some on other plant parts), but many are myrmecophilous and some 102. Axiidae (gold moths) are avery small family of even are carnivorous on ant larvae or homopterans. only 6 Palearctic species in the Mediterranean region. Hostplants are in a widevariety of plant families, but Adults are medium size and nocturnal. Larvae are particularly Fagaceae and Leguminosae. A few eco- leaf feeders. Hostplants are in Euphorbiaceae. No nomic species are known. Mexico has about 225 New World species are known species. 103. Thyatiridae (false owlet moths) comprise 224 99. (metalmark butterflies) total about species from all regions except Australia, but most are 1,419 species worldwide but predominate in the Neo- tropical Oriental (199 species); the actual fauna prob- tropics (1,322 species);the actual world fauna proba- ably exceeds 275 species. Two subfamilies are recog- bly exceeds 2,500 species. Five subfamilies are recog- nized. Adults are medium size and nocturnal. Larvae nized. Adults are small to medium size and diurnal; are mostly nocturnal leaf feeders. Hostplants are often with rapid flight. Larvae are leaf feeders, but recorded in a number of plant groups. Two species are many are myrmecophilous; one is known to be car- known for Mexico. nivorous on homopterans. Larvae mostly slug-like with short setae. Hostplants are recorded in numer- 104. Cyclidiidae (giant hooktip moths) are a small ous plant families. Mexico has about 255 species. family of 14 described species, all Oriental plus one species in the southern Palearctic. Adults are medi- 100. Libytheidae (snout butterflies) are a small um to large size and nocturnal. Larvae are leaf family of only 12 species, with at least one species in feeders. Hostplants recorded so far only in Alangiace- each faunal region. Adults are medium size and ae. No New World species are known. diurnal. Larvae are leaf feeders. Hostplants are in Ulmaceae. A single species is known for Mexico. 105. Drepanidae (hooktip moths) comprise 812 spe- cies worldwide, but predominately Oriental (647 spe- 101. Nymphalidae (brush-footedbutterfhes) are the cies); none is known for the Neotropics and only a few largest family of butterflies, with about 7,080 de- are in the Nearctic; the actual fauna probably exceeds scribed species worldwide; the actual fauna probably 950 species. Three subfamilies are known. Adults are exceeds 8,000 species. To represent the division of small to medium size and nocturnal. Larvae are leaf relict basal groups within the family, the subfamilies feeders. Hostplants include a variety of plants. Afew are divided into two groups: Nymphalinina, with 6 are economic. InMexico, no species have been record- subfamilies, and Satyrinina, also with 6 subfamilies. ed, although the higher mountains of northern Mex- Adults are medium to large and diurnal, although ico may harbor some relict species. INSECTA MUNDI, Vol. 16, No. 4, December, 2002 187

Bombycoidea 1ll. (Australianlappet moths) total 100 species, all from Australia. There are two subfami- This superfamily includes many of the largest lies. Adults are medium size to large; rarely with moths and encompasses 13families and about 4,810 micropterous females. Adults nocturnal, but at least known species. The superfamily is divided into two one species with diurnal males. Larvae are leaf groups: series Bombyciformes (Carthaeidae, Euptero- feeders and generally colorful. Host plants recorded in tidae, , , , several plant families. Some species have urticating Anthelidae, and ) and series Saturnii- larval setae. No New World species known. formes (, Lemoniidae, , Oxytenidae, Cercophanidae, and ). In 112. Lasiocampidae (lappet moths) include 2,130 Mexico, 341 species are known and perhaps 350 species worldwide, with many inAfrica (790 species). species will be the final total when all are described. Subfamilies are 5. Adults are small to large; some with micropterous females. Adults mostly nocturnal 106. Carthaeidae (Australiansilkworm moths) are but some males are diurnal. Larvae are leaf feeders, a monobasic relict family in Bombycoidea with a sometimes communally in silken tent-like webbings. single species from western Australia. Adults are Hostplants are various. Some species are economic as large and nocturnal (usually flying after midnight). tree defoliators. InMexico there are about 78 species. Larvae are leaffeeders, with numerous clubbedsetae. Hostplants are only in . No New World 113. Endromidae (glory moths) are a monobasic species are known. family of 4 species, from Europe to Central Asia. There are two subfamilies. Adults are medium size. 107. (giant lappet moths) total 325 Adult males are diurnal but females are nocturnal. species worldwide (except the Nearctic), but most are Larvae are leaf feeders. Hostplants recorded in Oriental (238 species); only 4 species are recorded in Betulaceae, Caprifoliaceae, Salicaceae, Tiliaceae, and the Neotropics. Three subfamilies are known. Adults Ulmaceae. No New World species known. are small to large and nocturnal. Larvae are leaf feeders. Hostplants among numerous different plants. 114. Lemoniidae (autumn silkworm moths) total 21 Few species are economic (e.g., rice or forest pests). species, mostly Mediterranean. There are two sub- Mexico has 2 species. families. Adults are medium size and nocturnal, but some males are diurnal; flight periods are oftenin the 108. Apatelodidae (American silkworm moths) are autumn in Europe. Larvae are leaf feeders. Host- exclusively New World, and total 252 species, mostly plants mostly in Compositae and Euphorbiaceae. No Neotropical(247 species). Three subfamilies are used. New World species known. Adults are small to mediium size and nocturnal. Larvae are leaf feeders. Hostplants are in various 115. Brahmaeidae (brahmin moths) are a small plant families. In Mexico, 43 species are known. family of 28 species, mostly Palearctic and African. There are two subfamilies. Adults are medium size to 109. Bombycidae (silkworm moths) total 166 de- very large and nocturnal. Larvae are leaf feeders. scribed species, all Old World and primarily Oriental Hostplants recorded in Asclepiadaceae and Oleaceae. (146 species), with only 5 species known for Africa. No New World species known. Two subfamilies are involved. Adults are medium size and nocturnal. Larvae are leaf feeders. Hostplants 116. Oxytenidae (American tropical silkworm moths) predominate in Moraceae. The silkworm ( include 60 species, all Neotropical. Adults are medi- mori) is usedfor production. None is native to the um size to large and nocturnal. Larvae are leaf New World, but one species is domesticated worldwide feeders; some mimic snakes. Hostplants recorded in for production of silk. Rubiaceae. Mexico has about 3 species.

110. Mimallonidae (sackbearer moths) total 254 117. Cercophanidae (Andean moon moths) include species, all New World and primarily Neotropical(250 30 species of mostly austral South American moths. species). Adults are medium size and nocturnal. There are two subfamilies Adults are medium size to Larvae are leaf feeders, with larvalcases. Hostplants large and nocturnal. Larvae are leaf feeders. Host- are recorded in a number of plant families. A few can plants recorded in Celastraceae, Lauraceae, be economic. Mexico has about 20 species. 188 Volume 16, No. 4, December, 2002, INSECTAMUNDI

Saxifragaceae, and Tiliaceae. None is known from ious (especially among ) and feeding Mexico. nocturnally. Hostplants include a large variety of plant families, especially for broadleaf forest trees. A 118. Saturniidae (emperor moths) include 1,435 number of economic species are known. In Mexico, species worldwide, but are predominately Neotropical about 800 species are known. (860 species). There are 7 subfamilies. Adults are medium size to very large and mostly nocturnal or 121. Dioptidae (American false tiger moths) total crepuscular, but some are diurnal. Larvae are leaf 507 species, primarily Neotropical(505 species); actu- feeders and many are polyphagous, some being com- alfauna likely exceeds 800 species. Two subfamilies munal or gregarious; many are extremely large. are known. Adults are medium size and mostly Hostplants are extremely varied. Some species are nocturnal, but some are diurnal or crepuscular. economic for agriculture, but major urticating larvae Larvae are leaf feeders, particularly toxic plants in are involved in dermatitis and more severe allergic families like Aristolochiaceae, Euphorbiaceae, Passi- reactions, and in a few cases even from adult scales. floraceae, and Violaceae, but also on various others Mexico has about 195 species. like Fagaceae. Very few are economic. In Mexico, about 200 species are known. Sphingoidea 122. Thyretidae (Africanmaidenmoths) include 212 This is a monobasic superfamily for the hawk species, all African. Adults are medium-size. Adults moth family, . Some specialists include perhaps mostly diurnal; often wasp mimics. Larvae the family in Bombycoidea. are thought to be leaffeeders, but most species remain unknown biologically. Hostplant records include 119. Sphingidae (hawk moths) total 1,238 species Thymelaeaceae and Ulmaceae. No New World species worldwide. Tropical regions of the New World, Africa, are known. and Asia have the most biodiversity. There are three subfamilies. Adults are medium size to very large and 123. Lymantriidae (tussock moths) total2,490 spe- nocturnal or crepuscular, but some are diurnal. cies worldwide; actual fauna likely exceeds 3,000 Larvae are leaf feeders, usually with a posterior horn- species. Most of the fauna is Old World tropical (about like scolus (thus, the common name, hornworms, for 2,090 species). Two subfamilies are used. Adults are the larvae); many larvae extremely large. Hostplants small to very large and mostly nocturnal, but some recorded in numerous plant families. Some are eco- are diurnal or crepuscular. Larvae are leaf feeders, nomic. Mexico has about 201 species. sometimes gregariously. Hostplants include many different plant families. Many species are serious Noctuoidea defoliators of forest trees. About 50 species occur in Mexico. The largest superfamily, containing nearly a third of all lepidopterans, the Noctuoidea comprise 124. Arctiidae (tiger moths) include 11,155 species about 44,025 described species worldwide, mainly in worldwide, primarily Neotropical (about 6,000 spe- the families Noctuidae and Arctiidae. In Mexico, cies); actual world fauna likely exceeds 14,000 species. about 6,550 species are known, but as many as 9,500 There are 5 subfamilies among 3 groups: group species may occur there. Pericopinina (with Pericopinae), group Arctiinina (for Lithosiinae and ), and group Ctenuchinina 120. (prominent moths) total 3,562 (for Ctenuchinae and Syntominae). Adults are small species from all faunal regions, particularly from the to large; hindwings greatly reduced in some groups Neotropics (1,766 species); actual world fauna likely (wasp moths). Adults mostly nocturnal but many are exceeds 4,000 species. The subfamily classification crepuscular or diurnal (Pericopinae, Ctenuchinae, varies, but currently involves 10 subfamilies, with and Syntominae). Larvae areleaffeeders. Hostplants segregationinto 3 groups: Oenosandrinina (for Oenos- are varied among numerous plant families, including andrinae, with 3 species inAustralia), Thaumetopoein- mosses and lichens. Mexico has about 1,000 species, ina (for Thaumetopoeinae), and Notodontinina (for with many more to be described. the remaining 8 subfamilies). Adults are small to very large; some with massive bodies. Adults are mostly 125. Noctuidae (owlet moths) are thelargest family nocturnal. Larvae are leaf feeders, sometimes gregar- of Lepidoptera, with about 26,310 described species INSECTA MUNDI, Vol. 16, No. 4, December, 2002 worldwide; actual fauna likely exceeds 30,000 species. Davis, D. R., and P. Gentili. 2003. Andesianidae, Major regions of biodiversity are in the Neotropics a new family of monotrysian moths (Lepidoptera: (about 8,600 species) and the Indo-Australianregion Andesianoidea) from austral South America. In- (6,500 species). Numerous subfamilies and tribes vertebrate systematics, 17: 15-26. have been described and the classification is still in Eaton, J. L. 1988. Lepidopteran Anatomy. John flux, but 26 subfamilies are now recognized, mostly in Wiley, New York. 257pp. group Noctuinina; also segregated are Heppner, J. B. 1991. Faunal regions and the diver- (previously in Arctiidae), in group Aganainina, and sity of Lepidoptera. Tropical Lepidoptera, 2(sup- , in group Herminiinina. Adults are pl. 1): 1-85. small to very large and mostly nocturnal, but some Heppner, J. B. 1996. Keys to families of Lepidoptera. are crepuscular and several groups are diurnal. Lar- Tropical Lepidoptera, 4 (suppl. 3): 1-28. (1993) vae mostly leaf feeders, but many are borers; hermi- Heppner, J. B. 1998-inprep. Classification of Lepi- ines are litter feeders. Hostplants include numerous doptera. Holarctic Lepidoptera, 5 (suppl. 1):1-148 plant famihes, but the majority of owlet moths are not (1998); 10 (suppl. 1): 149-320(2003); 11 (suppl. 1): known biologically. Some agricultural pests are in- [in prep]. cluded in the family (e.g., cutworms, armyworms, Heppner, J. B. In press. Lepidoptera. In J. L. and others). InMexico, there are about 4,500 species, Capinera. (ed.), Encyclopedia of entomology. CRC with a likely actual fauna of perhaps 5,500 species. Press, Boca Raton. Heppner, J. B. (ed.). 1984. Atlas of Neotropical References cited Lepidoptera. Checklist: Part 1. Micropterigoidea - Immoidea. W. Junk, Leiden. 27 + 112pp. Balcazar-L., M. A., and C. R. Beutelspacher-B. Heppner, J. B. (ed.). 1995. Atlas of Neotropical 2000a. Saturniidae (Lepidoptera). In J.Llorente- Lepidoptera. Checklist: Part 2. Hyblaeoidea - B., E. Gonzhlez-S., and N. 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