I^^^^P^MJ SMITHSONIANINSTITUTION UNITED STATESNATIONAL MUSEUMBULLETIN 244WASHINGTON, D.C.1964 MUSEUM OF NATURAL HISTORY Bagworm Mothsof theWestern Hemisphere(Lepidoptera: Psychidae) DONALD R. DAVISAssociate Curator of Lepidoptera SMITHSONIAN INSTITUTIONWASHINGTON, 1964 Publications of the United States National MuseumThe scientific publications of the United States National Museum include twoseries, Proceedings of the United States National Museum and United States NationalMuseum Bulletin.In these series are published original articles and monographs dealing withthe collections and work of the Museum and setting forth newly acquired factsin the fields of anthropology, biology, geology, history, and technology. Copiesof each publication are distributed to libraries and scientific organizations andto specialists and others interested in the various subjects.The Proceedings, begun in 1878, are intended for the publication, in separateform, of shorter papers. These are gathered in volumes, octavo in size, withthe publication date of each paper recorded in the table of contents of the volume.In the Bulletin series, the first of which was issued in 1875, appear longer,separate publications consisting of monographs (occasionally in several parts)and volumes in which are collected works on related subjects. Bulletins areeither octavo or quarto in size, depending on the needs of the presentation.Since 1902, papers relating to the botanical collections of the Museum have beenpublished in the Bulletin series under the heading Contributions from the UnitedStates National Herbarium.This work forms number 244 of the Bulletin series. Frank A. TaylorDirector, United States National Museum U.S. GOVERNMENT PRINTING OFFICEWASHINGTON : 1964 Contents FanIntroduction 1Review of the literature 4Life history 7Characters 13Classification 18Conclusions 23Checklist 26Family Psychidae 28Species of uncertain generic position 141Adult males known but not examined 141Adult males unknown 146Unidentified larval cases 156Literature cited 159Index 219 Bagworm Mothsof theWestern Hemisphere' IntroductionThis study is a preliminary revision of the tineoid family Psychidaeof North and South America (including the West Indies). A treat-ment of such scope has been attempted only once (Gaede, 1936),and it was at best a very superficial study. Other published accountsconsist mostly of original descriptions, taxonomic notes, and workstreating restricted areas. As a result, the interpretations of previousauthors have been erroneous in several instances; fm'thermore, acomprehensive key for the genera and species has never been attempted;thus, reliable identification of many of these insects has been virtuallyimpossible.The purpose of the present study is to re-evaluate the decisions ofearlier workers; to complete and, where necessary, to correct theirconclusions; and to investigate and utilize several taxonomic criteriain order to facilitate identification and a more comprehensive under-standing of the insects presently included in this family.The members of the family Psychidae present nmiierous problemsthat have greatly hindered a satisfactory treatment of the group.No single factor is unique for the family, and the close interrelationof many factors make their solution no easy task. The frequencywith which many of these problems are met in the Psychidae demon-strates why this family has been considered one of the most perplexinggroups, taxonomically, to be fomid in the Lepidoptera. Some of themore prevalent problems encountered are:1. The extreme convergence in superficial appearance among amajority of the species; or, conversely, the extreme lack of obviousspecific differentiation.2. The sometimes extreme variation of diagnostic features in bothadults and larval cases (e.g., wing venation in certain species).3. The rarity in collections of a majority of the known species, withthe consequent necessity of founding new species on the basis of one ' Modified from a doctoral dissertation submitted to Cornell University, 1962.1 2 U.S. NATIONAL MUSEUM BULLETIN 244 or a few specimens. (Undoubtedly this lack of adult specimensresults in part from the exceedingly brief period of the year duringwhich these forms may be collected.)4. The disappearance or unavailability of the type specimens, afact that is all too frequently linked with the inadequacy of theoriginal description in providing diagnostic characters that can berelied upon.5. The lack of biological data, a factor which can be of prime im-portance in a group that displays so much structural homogeneity.(This paucity in large part may be due to the extended life cycle ofthe larva, a factor which hinders breeding and rearing experiments.)6. The lack of adequate collecting data, involving not only locality(especially for types) but also the association of larval cases withtheir corresponding adult, a fact of paramount importance in no. 7.7. The erection of new species solely on the larval case or on thebasis of a few general statements concerning the larva and/or pupa.8. The polyphagous feeding habit of the larva. (This sometimesfacilitates the rearing of these insects but of course makes it impossibleto interpret precise relationships based on host preferences.)9. The possibility of hybridization between certain species. (Suffi-cient evidence is at present lacking to support this supposition.)Taxonomic keys and discussions of the larvae were omitted becauseimmature stages of only a few species were available. A preliminaryexammation of setal maps, noting color markings and variations, etc.,revealed only a few minor divergences among some of the species;thus, it is believed that larval systematics will not profoundly modifythe present classification, which is based essentially on the adult male.Future work should be concerned especially with the collecting ofall stages of these insects. This work could best be done throughrearings, a procedure that would provide not only an abundant seriesof specimens representing all stages but also would furnish materialthat definitely could be associated. Genetic information obtainedfrom attempts to cross certain species and populations may bedifficult to acquire but would be highly desirable.All the specimens used in this study were borrowed fromthe collections of the United States National Museum, the BritishMuseum, and the sources Hsted below. The abbreviations used toindicate where the material is deposited are:AMNH?American Museum of Natural History, New York, N.Y.ANS?Academy of Natural Sciences, Philadelphia, Pennsylvania.BM?British Museum (Natural History), London, England.CAS?California Academy of Sciences, San Francisco, California.CM?Carnegie Museum, Pittsburgh, Pennsylvania.CNC?Canadian National Collections, Ottawa, Ontario.CPK?Collection of Charles P. Kimball, Sarasota, Florida. BAGWORM MOTHS OF THE WESTERN HEMISPHERE 3CU?Cornell University, Ithaca, New York.DEI?Deutsches Entomologisches Institut, Berlin, Germany.DRD?Collection of the author, Washington, D.C.INHS?-Illinois State Natural History Survey, Urbana, Illinois.MCZ?-Museum of Comparative Zoology, Harvard University, Cam-bridge, Massachusetts.MSU?Michigan State University, East Lansing, Michigan.PM?Museum National d'Histoire Naturelle, Paris, France.RNH?Rijksmuseum van Natuurlijke Historic, Leiden, Netherlands.RWH?Collection of Ronald W. Hodges, Washington, D.C.SNG?Senckenbergische Naturforschende Gesellschaft, Frankfurt, Ger-many.UM?University of Missouri, Columbia, Missouri.USNM?United States National Museum, Washington, D.C.ZMHU?Zoologisches Museum der Humbolt-Universitat, Berlin, Germany.ZSBS?Zoologische Sammlung der Bayerischen Staates, Mtinchen,Germany.Gross measurements were made with a standard millimeter ruler.The wing expanse was measured from the extreme apex of one wing tothat of the other side. In the larval cases, only those which weremature (i.e., which contained evidence of pupal material) were meas-ured. The diameter in all instances refers to the greatest diameterobserved along the length of the case.Except in those cases where specific aclaiowledgment is given,all photographs were made by the author with an Exakta VX Ila35 mm. camera. Drawings also were executed by the author, usuallywith the aid of a Kenivision Micro-Projector. Illustrations of somestructures (e.g., the male and female genitalia of certain species) havebeen included to show the similarities in these organs that may existbetween different species as well as the differences.Definitions of nearly all the terms used in this study may be foundin the glossary prepared by Torre-Bueno (1950). Terms relating tothe male and female genitalia were in part derived from the discus-sions of Burrows (1923), Forbes (1923), and Klots (1956). Selectedillustrations (figures 146-148, 310) have been labeled to show theprecise application of many of the terms used in the discussions onwing venation and genitalia. The terminology used in discussing thedistributions of the various species follows that of Smith (1935) andLobeck (1948).Many individuals and several institutions together have providedassistance in making this work possible. I wish primarily to expressthe gratitude I feel toward Dr. Frank Morton Jones (deceased. May1962), whose very fine collection and extensive earlier work on theNorth American Psychidae provided a firm basis for the presentrevision. In addition to his valuable collection, Dr. Jones presenteda fund of a thousand dollars to the Smithsonian Institution for future 4 U.S. NATIONAL MUSEUM BULLETIN 244 research in this family; I am very grateful for having received thisaward. Considerable appreciation must be given to Dr. J. F. GatesClarke, who first suggested this problem and helped promote andencourage my research, and to staff members of the United StatesNational Museum for the many services they have extended me.I wish to thank Dr. J. G. Franclemont, under whose guidance thiswork was performed, for his valuable advice concerning the manu-script and many perplexing problems, Dr. H. E. Moore, Jr., and Dr.J. A. Weidhaas, who also were very helpful in reading and criticizingthe paper.My sincerest appreciation is extended also to the following ento-mologists and the institutions or firms they represent for their coopera-tion in the loan of material: Dr. Harold J. Grant, Jr., Academy ofNatural Sciences; Dr. Frederick H. Eindge, American Museumof Natural History; Mr. Alan Watson, British Museum (NaturalHistory); Dr. C. Don MacNeil, California Academy of Sciences;Mr. George T. Okumura, California Department of Agriculture;Dr. Thomas N. Freeman, Canadian National Collection; Mr. HarryClench, Carnegie Museum; Dr. Giinter Petersen, Deutsches Ento-mologisches Institut; Dr. Frank W. Mead, Florida State PlantBoard; Dr. Ronald W. Hodges; Mr. Charles P. Kimball; Mr. PaulKoehler; Dr. Richard B. Selander, Illmois State Natural HistorySurvey; Michigan State University; Dr. W. T. M. Forbes, Museumof Comparative Zoology; Drs. Pierre Viette and Jean Bourgogne,Museum National d'Histohe Naturelle; Dr. A. Diakonoff, Rijks-museum van Natumiijke Historic; Dr. Heinz Schroder, Sencken-bergische Naturforschende Gesellschaft; Mr. Clyde Stephens, UnitedFruit Company; Dr. Wilbur R. Enns, University of Missouri; Dr.W. Forster, Zoologische Sammlung der Bayerischen Staates; Dr.H. J. Hannemann, Zoologisches Museum der Humbolt-Universitat.Finally I would like to express my gratitude to my sister, MissCarole Davis, for her services in typing the first di'aft of the manuscript.Review of the LiteratureThe first pubhshed account of members of this family that defi-nitely mhabit the Western Hemisphere was issued in 1827 by theRev. Lansdown Guilding, who lived for several years on the islandof St. Vmcent in the Lesser Antilles. While residing there, he con-tributed a few short reports dealing with West Indian biology. Inone, he described a new genus of psychid, Oiketicus, and two newspecies, kirbyi (q.v.) and macleayi, which he attributed to the WestIndies. BAGWORM MOTHS OF THE WESTERN HEMISPHERE 5Historically, the oldest name now considered referable to a speciesof Psychidae in this hemisphere is Thyridopteryx ephemerae-formis,which was described first by Haworth in 1803. At the tune, thespecies was thought to be native to the British Isles ; the first publishedsuggestion that the species had an American origin was not proposeduntil 1841 by Edward Doubleday. The oldest name of a psychidnow known to occur within the limits covered by this paper is that ofFumaria casta (Pallas), 1767.The bulk of the literatiu"e for this region that has accumulatedsince these earliest references consists chiefly of descriptions of newspecies. To summarize this mformation and to note the workerswho have contributed in varying degrees to the discovery of theAmerican Psychidae, an alphabetical list of these authorities follows,showing the species (as originally proposed) for which they areresponsible and the years that the names were fii*st used. Thenumber in parentheses after the author's name indicates the numberof species that are considered synonyms in the present study (thisdoes not include those species mentioned in the two appendices) . Barnes, William (0), Thyridopteryx alcora, 1905.Barnes, W., and Benjamin, Foster (2), Manatha jonesi, 1922; Eurukullaruspolingi, 1924; Oiketicus honniivelli, 1924.Barnes, W., and McDunnough, James (0), Manatha nigrita, 1913; Prochaliapygrnaea, 1913; Apterona fragilis, 1916.Berg, Frederico (0), Oiketicus geyeri, 1877; Oeceticus westwoodii, 1882;Oeceticus platensis, 1883.Butler, Arthur (0), Hyaloscotes fumosa, 1881; Thanatopsyche canescens, 1882.Clemens, James (0), Solenobia tvalshella, 1862.Davis, Don (0), Lumacra haitiensis, 1964; L. hyalinacra, 1964; L. quadri-dentata, 1964.Dognin, Paul (0), Animula limpia, 1894.Dyar, Harrison (1), Platoeceticus aphaidropa, 1914; P. symmicta, 1914;Chalia zacualpania, 1916; Pachytelia pithopoera, 1923; Zamopsycheco7nmentella, 1923; Pachytelia lepidopteris, 1926.Edwards, Henry (0), Psyche coniferella, 1877; P. fragmentella, 1877; Oiketicusdavidsoni, 1877; Thyridopteryx meadi, 1881.Esper, Eugen (0), Bonibyx pulla, 1785.Felder, Rudolph (0), Psychoglene basinigra, 1874.Freeman, Thomas (1), Hyaloscotes sheppardi, 1944.Gaede, Max (0), Oiketicus mexicanus, 1936; Thyridopteryx seitzi, 1936.Grossbeck, John (1), Thyridopteryx pallidovenata, 1917.Grote, August (0), Hymenopsyche thoracicum, 1865; Oiketicus abbotii, 1880.Grote, A., and Robinson, Coleman (0), Psyche confederata, 1868.Guilding, Lansdown (0), Oiketicus kirbyi, 1827; 0. macleayi, 1827.Hampson, George (0), Thanlopsyche apicalis, 1904.Hawortli, Adrian (0), Sphinx ephemeraeformis, 1803.Herrich-Schaffer, Gottlieb (1), Animula dichroa, 1856; Oeceticus fulgurator,1856.Heylaerts, F. J. M. (0), Chalia rileyi, 1884; Eumeta brasiliensis, 1884;Manatha edwardsi, 1884; Chalia kUnckelii, 1901. 6 U.S. NATIONAL MUSEUM BULLETIN 244Jones, Frank (2), Eurycttarus tracyi, 1911; Psyche cacocnemos, 1922; P.celibata, 1922; Oiketicus toumeyi, 1922; Thyridopteryx vernalis, 1923;Dendropsyche burrowsi, 1926; Oiketicus dendrokotnos, 1926; Platoeceticuscongregatus, 1945; P. watsoni, 1945.Koehler, Paul (4), Chalia rebeli, 1924; Oiketicus elegans, 1931; Platoeceticusrugosus, 1931; P. tandilensis, 1931; Oiketicus horni, 1938; Chaliadispar, 1939; Chlanialicheniphilus, 1939; Clania dagiierrei, 1939; Cochlio-theca fiebrigi, 1939; Platoeceticus chaquensis, 1939; P. ho^fmanni, 1939;Oiketicus liseri, 1939; 0. oviformis, 1939; Zamopsyche haywardi, 1939;Clania borsanii, 1953; C yamorkinei, 1953; Oiketicus ginocchionus, 1953.Lucas, Pierre (1), Oiketicus poeyi, 1856.Moeschler, Heinrich (0), Psijche surinamensis, 1878.Packard, Alpheus (2), Oeceticus coniferarum, 1864; Platoeceticus gloverii,1869; Psyche carbonaria, 1887.Pallas, Peter (0), Phalaena casta, 1767.Philippi, Rudolph (0), Psyche chilensis, 1860.Schaus, William (6), Oiketicus jonesi, 1896; Chalia tristis, 1901; C vigasi,1901; Oiketicus orizavae, 1901; Oiketicus specter, 1905; Platoeceticusmarona, 1905; Thanatopsyche thoracica, 1905; Thyridopteryx microptera,1905; Platoeceticus costaricensis, 1911; Chalia pizote, 1927.Siebold, Carl (0), Psyc/ie AeKx, 1850.Townsend, Charles (0), Oiketicus townsendi, 1894.Vazquez G., Leonila (5), Eurikuttarus hoffmanni, 1941; Oiketicus assiinilis,1942; 0. fasciculatxis, 1942; 0. multidentatus, 1942; 0. ochoterenai, 1942;0. sinaloanus, 1942; 0. mortonjonesi, 1949; 0. zihuatanejensis, 1951.Weyenbergh, Hendrik (1), Oiketicus tabacillus, 1884; Psyche bergii, 1884;P. burmeisteri, 1884; P. cassiae, 1884.Zeller, Phillip (1), Oiketicus gigantea, 1871.In the United States four workers have been prominent in theliterature of this family. Dyar (1914-26) and Schaus (1896-1927)were concerned primarily with the Neotropical species, but Dyarwrote one paper (1923) that briefly treated several North Americanbagworms. Schaus described the greater nmnber of species (10),but at least six of these represent synonyms. Barnes (1905-24),in collaboration with Benjamin and McDunnough, and especiallyJones (1911-45) focused their attention on the Nearctic species. Thelatter author, through extensive collecting and in several short butinformative papers, has contributed more toward the understandingof this diflBcult family than any other entomologist of the hemisphere.In Mexico the efforts of Vazquez (1942-53) should be mentioned;her major work (a doctoral thesis) is entitled "Estudio Monografico delas Psychidae de Mexico." Relying in part on the early collectionsof Roberto Miiller and Charles Hoffman as well as her own, Vazquezhas described a total of eight new species from Mexico, of which fivehave been synonymized here. Twenty-one species are recognizedby Vazquez as occurring in Mexico, but approximately nine of thesenames (including the five mentioned above) have been reduced tosynonyms. BAGWORM MOTHS OF THE WESTERN HEMISPHERE 7Comparatively little work has been accomplished in South Americaby entomologists of that area, and nearly all that has been done is theresult of German nationals residing in Argentina. Two of the earliestworkers were Berg (1877-83) and Weyenbergh (1884). Within recentyears Koehler (1924-53) has presented several short papers on thesystematics of Argentine psychids. He has described a total of 17new species, but, unfortunately, many of these are not represented byadult males, and so they have been treated in the appendices of thispaper.Four other European entomologists need to be mentioned.Heylaerts (1881-1901), although not primarily interested in Americaninsects, described four new species of bagworms which were sent tohim from this hemisphere. The identities of three have been posi-tively established; that of the fourth (rileyi) remains highly uncertain.Heylaert's major effort in this family culminated in his "Essai d'uneMonographic des Psychides" (1881), but this was concerned mainlywith the Em'opean species. The family Psychidae was catalogedfor the world fauna by Dalla Torre and Strand (1929). This referenceis most useful, but it has become somewhat outdated and it includesseveral genera (e.g., Acrolophus) which are no longer assigned to thePsychidae. The only previous comprehensive treatment of thisfamily for the Western Hemisphere was that of Gaede (1936). In thiswork 17 genera and 68 species (including one genus and species sincereferred to the Cossidae) were considered in an extremely superficialmanner. Life HistoryThe interesting life histories of the various members of this familydemonstrate many phenomena that are unique or nearly so for theLepidoptera. To supplement the present author's limited obser-vations and conclusions acquired from the rearing of four species,the works of several entomologists have been summarized ratherfreely in the following discussion.Distribution.?Present records indicate this family is widelydistributed in the Americas, from as far north as Banff in Alberta,Canada (approximately 53? N. latitude), southward to the island ofNavarino at the extreme tip of South America (55? S. latitude). Inaltitude these insects are known to range in the New World from nearsea level to as high as 2200 meters in the Colombian Andes.The flightless condition of the female psychid appears to be almosta contradiction of the very extended distribution that several specieshave attained; however, if one examines more closely their variousmeans of dispersal, the phenomenon can be better understood. Oneimportant present day means of dispersal for bagworms is the acci- 8 U.S. NATIONAL MUSEUM BULLETIN 244dental shipment of infested plant stock by human agencies as attestedby the relatively recent introductions of tlu-ee old world species(Apterona crenulella f. helix, Epichnopterix pulla, and Fumaria casta)into widely diverse regions of the Western Hemisphere. In addition,Solenobia walshella possibly may have been introduced into thiscountry from Europe during colonial times.Certain biological adaptations of the Psychidae make such acci-dental transportation very possible: the ability of the larva to resiststarvation for long periods of tune, the polyphagous feeding habit(includmg, sometimes, the ability to subsist for brief periods on thedried leaves of its own case), the habit of firmly attaching the bagto almost any support for pupation, the high reproductive potentialof a single female, and the occurrence of parthenogenesis in a fewspecies. The last factor evidently has played a major role in theestablishment of Apterona crenulella f. helix in this country.Theoretically, the successful introduction of a single egg is all that isnecessary for a parthenogenetic insect such as A. c. form hslix tobecome naturalized in certain favorable habitats.Another means of dispersal of bagworms involves birds. Anunusual observation by J0rgensen (1954) needs more investigation.He noted the viability of 30 or 40 larvae from eggs of Acanthopsycheatra L. that reportedly had passed through the digestive tract of aEuropean robin (Erithacus rubecula). Mature females of this psychidwere fed to these birds, and larval emergence from the fecal droppingswere later observed. Small larval cases of other bagworms also havebeen reported attached to the legs of birds.Wind dispersal of not only newly emerged larvae but also of variouslater stages may be an important factor over short distances. Younglarvae, on emerging from the egg, often will spin silken threads andengage in an activity generally referred to as "ballooning," the com-mon practice of many species of spiders. Strong breezes are capableof carrying these tiny larvae with their attached threads over shortdistances. Windstorms would be even more effective and couldalso aid in the dissemination of the more matm'e stages. Seitz (1912)has reported finding, on the treeless pampas of Uruguay, large branchesof trees with living bagworms attached that obviously had been trans-ported several miles by a violent storm (pampero).Egg.?The eggs, which may number from 200 for some smallerspecies (Tutt, 1900) to as many as 6756 for the female of Oiketicuskirhyi (Stephens, 1962), almost invariably are deposited in the larvalcase, usually within the pupal shell that the female frequentlyabandons. The length of this stage varies considerably dependingon the species; some (e.g., Thyridopteryx ephemeraeformis) pass thewinter in this state. Stephens (1962) has reported that the egg stage BAGWORM MOTHS OF THE WESTERN HEMISPHERE 9 of Oiketicus kirbyi from Costa Rica is rather constant; hatchingusually occurs within 29-33 days.Larva,?The length of time required for this stage varies alsowith the species and sometimes considerably within a species.Several bagworms overwinter in this stage (usually as a last instarlarva), but the shortest tmie spent by any species as a larva is approxi-mately two to three months. The larval period of some of the largertropical species is evidently very extended. 0. kirbyi (in Costa Rica)has been observed by Stephens to require as long as 382 days formaturity, with considerable variation in the length of time. Con-sidering such information, it is perhaps superfluous to mention thatrarely is there more than one brood per year, but a second generationmay be completed by some psychids inhabiting the warmer climates.Very little is known about the nmnber of larval instars, owing inpart to the concealed nature of the moulting process (performedentirely within the case) and to the immediate eating of the cast skinby the larva. Jones and Parks (1928) briefly mention that thereare "four or more molts." An actively feeding bagworm, whenpreparing to shed its cuticula, normally will attach its case andtightly close the anterior end as if preparing for pupation. Stephens,in recording the number of tunes this interruption occmTed, concludedthat 0. kirbyi underwent approximately 12 to 20 molts during itslarval history.Apparently this family does not contain a single species which dem-onstrates host specificity. The polyphagous habit is well representedthroughout the family, recorded hosts of many bagworms beingnumerous and widely varied. The more primitive members of thePsychidae, the so-called Micro-Psychina, are more prone to be scav-engers, feeding on lichens and other low plant forms, and even resortingto an insectivorous diet at times. These tendencies also are carriedover to the Macro-Psychina to some extent, but in this group thehigher plant forms are preferred.Even though the host range of most bagworms is frequently ratherextensive, several species show a great reluctance to change food plantsabruptly within the larval life. For many of these insects there seemsto be a very definite preference for a single food plant, usually thehost upon which the young larva first commenced to feed. Jones andParks (1928) stated their belief that "food-plant choice by the younglarvae rests on an inherited preference as well as on early proximityand reluctance to change. In this way impermanent food-plant racespersisting for a number of years often result." This reluctance of alarva to feed successively on different plants probably varies with eachspecies. The present writer, in rearing numerous larvae of 0. platensis,observed that the insects feed indiscriminately on a wide variety of 10 U.S. NATIONAL MUSEUM BULLETIN 244plants {Ostrya, Robinia, Ulmus, Vinca minor, etc.) without anynoticeable cessation in feeding from the changing of hosts.The construction of a portable case by the larva is one of the mostobvious features of the family. This unusual habit, however, is notrestricted to the Psychidae, but it also occurs in the closely relatedfamily Tineidae as well as in such groups as the Coleophoridae andLacosomidae. The basic construction of the case is of silk, which isspun from the labial glands of the larva. Various types and arrange-ments of ornaments usually are applied to this foundation. Bothextremities of the case are open and are usually very flexible for ashort distance from either end, thereby greatly facilitating the closingof the two apertures by the larva inside. The overall appearance ofthe case may var}^ considerabl}^ among different species of psychids,and it frequently may be diagnostic for a given species. In severalinstances, however, the architecture of the larval bag definitely isknown to be of no specific importance. The Macro-Psychina fre-quently differ from the more primitive psychids in having the larvalcases often covered with rather large fragments of plant material; onthe other hand, the sacks of the Micro-Psychina are more tineid-likein construction: the exterior is nearly bare or it is covered often withminute fragments of lichens and/or soil particles.Newly emerged bagworms very soon begin to construct somewhatconical cases that at first are carried upright. As the larva grows,the bag is enlarged correspondingly and, because of the increasedweight, is carried in a pendant manner in which the insect clings tosome support by means of its well-developed thoracic legs. The analprolegs play an important role in retaining a firm grasp on the case.The anterior four pairs of abdominal prolegs, having lost a majorpart of their function, are reduced to mere vestiges.Pupa.?All members of this family undergo pupation within thelarval case. In preparing for pupation, the mature larva first firmlysecures the upper end of the case to some support, thereby sealingthe normal entrance. The larva of both sexes then reverses itselfwith the result that the head is directed toward the lower end of thecase, and it pupates in this position. The male pupa of the Macro-Psychina is approximately half the size of the female, and it pos-sesses the normal sheaths for the various body appendages; that ofthe female is very simple, with the leg and antennal sheaths beingpresent only in the more primitive group. The length of time in thisstage varies considerably within a species, between sexes, and amongthe species. In those that do not hibernate in this stage, the intervalfor the pupal instar may range from less than a week for some speciesto as long as 111 days for the male of 0. kirhyi (Stephens, 1962).Adult.?Immediately before the emergence of the adult, the malepupa gradually pushes itself partly out of the bag. Soon after leav- BAGWORM MOTHS OF THE WESTERN HEMISPHERE Hing the pupal shell, the male is ready for flight, and it may be ob-served clinging to the pupal remains or to the lower part of the caseexercising its wings. The wing scales of several species are attachedvery loosely and often are easily lost at this time, leaving a portionof the wing membrane transparent. The flight time of the male mayfollow either a diurnal, crepuscular, or nocturnal habit, depending onthe species. The response to light also varies accordingly; somespecies are somewhat phototropic while others almost never occurnear lights.For a majority of the members of the Macro-Psychina (those with-out functional legs), the initiation of the adult stage in the femaledoes not involve an emergence from the larval case; in fact, seldomdoes this form leave the pupal shell until after copulation has oc-curred. In some of the more primitive members of this group (e.g.,Fumaria) and in the Micro-Psychina, the females possess a normalcomplement of legs, and they are thus capable of leaving both thepupal shell and the larval case. After emerging, these forms usuallymay be observed clinging to the exterior of the case awaiting copula-tion with a male. In the genera Diplodoma and Narycia, whichpresently are considered by most authorities to be the most primitivemembers of the family, the female is fully winged and presumablygoes through the same procedure in emerging as does the male.More than likely adult males are attracted to receptive females byscent. The distance for this reception may be rather considerable,and in some instances it is believed to be as much as a mile or so.Observations by Hardenburg (1917) on Acanthopsyche junodi sug-gested that females had been visited by males which "could only havecome from a couple of miles distant."The actual mating procedure varies somewhat in this family,depending upon the location of the emergent female. In the lowerforms, as stated previously, copulation with the male occurs outsidethe larval bag. In the higher psychids, mating has to occur withinthe case, and the entire abdomen of the male is highly specializedfor this requirement. The typical procedure for such forms may besummarized as follows. After locating a case containing a receptivefemale, the excited male assumes a position at the lower end of thebag and begins to probe with its abdomen into the aperture of thecase. Normally the only rupture in the female pupal case occurs atthe anterior end,whichprojectsdownward, as did the malepupa. Thus,the abdomen of the male must first pass into this opening and extendthe entire length of the female's body (fig. 145) before coming intocontact with the female genitalia. This remarkable feat is accom-plished primarily by the telescopic action of the male abdomen,which is capable of extending itself to three times its normal length.693-052?64 2 12 U.S. NATIONAL MUSEUM BULLETIN 244The intersegmental membranes of this region are very developed andflexible, providing the necessary surface area for expansion. Relyingchiefly upon pneumatic pressure from within its inflated body, themale extends its abdomen the necessary distance for eventual copula-tion. During actual union the moth becomes quiescent and remainssuspended, head downward, for a brief period of usually less thanten minutes (Jones, 1927). After completion of the act, the abdomenis slowly withdrawn and the male flies away, often capable of matingagain.In a few old world genera (e.g., Apterona, Lnffia, and Solenobia),parthenogenetic races are Ivnown to exist. A considerable amountof investigation of these insects has been made, primarily by Seller(1917-49) and more recently by Narbel-Hofstetter (1946-55). Ithas been suggested, although by no means proven, that partheno-genesis also may occur occasionally in some species which are normallybisexual.Egg laying commences immediately after mating. As statedpreviously, the eggs normally are deposited within the larval caseand frequently within the pupal shell. In those species thatnormally mate outside the case (e.g., Solenobia), the females, aftercopulation, introduce the abdomen into the lower opening of the larvalbag and deposit the eggs in an irregular mass, intermixed with woolyhair shed from the posterior segments of the female. Several authorshave suggested that the eggs actually are not laid but mature withinthe parent's body, the young larvae eventually bursting through thebody wall. Generalizations of this nature should be regarded withconsiderable skepticism until more careful observations can beperformed. Following oviposition, the very shrunken female usuallysucceeds in forcing her way out of the case (if she has not alreadydone so) and dropping to the ground eventually to perish.Natural enemies.?Populations of bagworms in some areas oftheir distribution apparently can be seriously affected by the predationand parasitism of diverse organisms. Previous work in this phase ofpsychid biology has been quite insufficient, and no conclusions canyet be drawn concerning such phenomena as the frequency of hostspecificity among some of the parasites. One of the few papersdevoted to this area of study has been that of Koehler (1939b), Re-cently published findings by Stephens have shown the existence ofseveral new species of parasites attacking Oiketicus kirbyi.Birds, lizards, and certain arachnids are known to prey uponmembers of the Psychidae. Birds have been noted eating both thelarvae and adult females; however, the construction of the larvalcase sometimes offers adequate protection from such dangers. Oc-casionally bagworm cases are found pierced with holes that apparently BAGWORM MOTHS OF THE WESTERN HEMISPHERE 13 were made by bii'ds (probably woodpeckers) in extracting the contents.Wolcott (1951) mentions an interesting example of predation by thelizards Anolis pulchellus and Anolis cristatellus in Puerto Rico.These animals reportedly swallow the larva inside its bag althoughthe latter, presumably, is indigestible. The present writer has noticedthat the egg-filled cases of Oiketicus platensis and Thyridopteryxephemeraeformis are frequented at times by small spiders of the familySalticidae. As the young larvae emerged, several were eaten by thepredators. Predaceous mites also have been reported among the eggmasses: these arachnids probably attack the eggs primarily. Stephensfm-ther reports that ants ("probably Solenoj^sis") will occasionallyfeed on the larvae of 0. kirbyi.The Psychidae are parasitized by various other groups. Themost common is the Ichneumonidae, which contain several species oflarval parasites. Other families of Hymenoptera that are known toattack bagworms are the Bethylidae, Braconidae, and Chalcidae.Dipterous parasites are known from the families Sarcophagidae andTachinidae. In addition to these insects, Stephens has found that afungus, Beauvaris bassiana Balsamo, and a protozoan, Nosemaspecies, are parasitic on the larvae of 0. kirbyi.In the collections of the United States National Museum there area few larval cases of Oiketicus toumeyi that have been used for nestsites by a vespid wasp, identified by Dr. Karl Krombein asPachodynerus acuticarinatus (Cameron). Mud nests consisting ofapproximately four to eight cells were constructed within the case,where the larvae of this intruder were reared. In one larval case wasfound the female pupal shell of a psychid; in another, the dried re-mains of a nearly matured larva. Since the bag of the latter wasblocked at both ends by the nests of the wasp, it is possible that thedeath of the bagworm was caused indirectly by the presence of thevisitor. The relationship of these two insects is evidently not thatof host-parasite. It is also possible that the wasp makes use of thecase after the original inhabitant has either died or pupated.Pachodynerus acuticarinatus previously has been reported (Rau,1940) utilizing for nesting sites the old, vacated cells of anotherhymenopteran, Sceliphron. CharactersThe morphological characters that are employed in this family todelimit genera and species present many problems of character evalu-ation in defining taxonomic categories. Clench (1959), in workingwith a small group of South African psychids, has accurately em-phasized this predicament. Several species or groups of species, al-though resembling each other very closely in most features and thus 14 U.S. NATIONAL MUSEUM BULLETIN 244demonstrating obvious relationships, sometimes may diverge extraor-dinarily in respect to other characters that in other families of Lepi-doptera would be considered of generic or even suprageneric impor-tance. This prominent divergence of several important charactersthroughout the Psychidae adds greatly to the difficulty of preciselydefining the family. This divergence also emphasizes the presentwidely accepted principle of systematics that the evaluation of anycharacter must be appraised for the particular group under considera-tion. Frequently this principle is applied to the family level, and oftenit may be extended to higher categories. Within the Psychidae,however, as observed in this study, the proper appraisal of supposedgeneric characters sometimes has to be decided for a given group ofspecies. Certain characters may be relied upon to define severalgenera, but these same characters may vary within other comparableunits, where they may be of no more than specific unportance.Criteria for generic limits were acquired in this study from theexamination of such groups as Oiketicus and Cryptothelea. The in-formation gained thereby was then employed in the recognition ofsupposed equivalent groups. It is obvious that more characters needto be investigated and correlated. In the African Monda group,Clench has discovered very significant generic characters in thethoracic anatomy as well as in the more traditional morphologicalareas of the body. Similar differences also may be present amongsome of the New World genera of Psychidae.In the following paragraphs several characters are discussed undertheir various stages in an effort to smnmarize their apphcability to thesystematics of this family.Male.?Systematic research in this family has been and still ispredominantly dependent upon the adult male. The paucity ofsufficient characters in the other stages and the extreme specializationin a majority of the females probably will always make this so.Antennae: Several characters of value have been noted on thisorgan: the presence of lateral pectinations as they may occur along theentire length of the shaft; the relative length of the pectinations whenpresent; the relative length and arrangement of the sensory hairs.One of the few characters that appears invariably to be of genericimportance in the Psychidae is the origin of the pectinations from thesegments of the shaft. The rami may originate basad (Fumaria),mesad (in most genera), or distally (as is the tendency for Hyaloscotes) . In most species the rami usually arise more basad from those segmentsnear the base of the antenna and gradually become more distad in theouter segments. Thus, as a standard in comparing this condition,observations near the middle of the antennae have been made in thisstudy. BAGWORM MOTHS OF THE WESTERN HEMISPHERE 15Palpi: For our New World psychids, these structures offer limiteduse in classification, owing to their extreme reduction in nearly all thespecies. Only in the more primitive genera (Epichnopterix or below)do they retain an observable size; they may be of generic value, if notof specific importance, in these groups.Thorax: As previously mentioned, this region of the body maybear taxonomically important characters, usually as observed in thevarying proportions and outlines of the lateral sclerites. This featurehas not been fully evaluated in the present study. Dissectionsrepresenting foiu- genera (viz, Astala, Cryptothelea, Oiketicus, andThyridopteryx) were examined. Relatively minor differences werenoted between the subalar sclerites of the mesothorax, but some ofthis may be due in part to intraspecific variation. A thorough studyof the thorax in this regard eventually should be performed.Wings: Very little divergence exists in the wing color of mostspecies of psychids. Few genera throughout the world demonstratea distinct color pattern, and in the Western Hemisphere this charac-teristic is restricted primarily to Oiketicus. Normal variation in thecolor tone of certain members of this genus (e.g., 0. kirhyi) has re-sulted in a rash of superfluous names.Of more importance than the actual wing color is the shape of thewing scales and their distribution over the wing membrane. Theshape of the scale varies according to its relative location on the wing.Those near the wing base usually tend to be the broadest and possessthe greatest number of apical teeth (if any) ; a gradual narrowing ofthe scales is evident toward the outer margin. Consequently, it isboth necessary and convenient to select a particular area of the wingto compare and discuss. In this study the discal cell of the forewingwas chosen.The wing venation is sometimes exceedingly variable in this family;however, it is believed by the present author that for many speciesthis feature has been greatly overemphasized. Most of the variationthat is observed usually involves the medial veins. Only occasionallyis the radial system also affected. The exact cause of this inconsist-ency is not definitely known, but perhaps it is caused by unusualenvironmental influences on the larva and/or pupa. In some instances,the variation that is observed may result, in part at least, from therearing of certain species. A study of the importance that rearingmay exert, however, has not been pursued. Only certain speciesexhibit pronounced variation; others demonstrate a remarkably con-stant venation. A very consistent pattern can be observed in Oiketi-cus kirhyi. The wing veins of 65 specimens (130 wing pairs) of thisspecies were examined, and only two minor variations from the norm 16 U.S. NATIONAL MUSEUM BULLETIN 244 were noted. These involved merely the connate condition (versustypically stalked) of M2 and 3 in the hindwing.Legs: Several important characters are to be noted in the legs ofthese moths. The armatm-e of the tibiae is one of the more obvious,which, imfortunately, has been neglected largely by a number ofearlier workers. The presence or absence of the epiphysis or of thesmall tibial spurs occasionally provide absolute species characters (asin 0. platensis), and frequently these are correlated with other charac-ters in order to aid in defining genera. The condition of the epiphysis(or the spurs of the tibiae) should not itself be considered a firm basisfor generic separation as some authors erroneously have believed inthe past. Examination of the interspecific variation of this structureas it occurs in such genera as Solenobia, Hyaloscotes, and Oiketicusshould make this evident.Leg ratios, involving usually the relative lengths of the tibia andtarsal segments, often can be rather useful. Characters of this sorthave been applied in the erection of the new subgenus Paraoiketicus.Tarsal claws may be of considerable taxonomic importance (Clench,1959). In the American forms, however, no divergences were notedfrom the smoothly ciu-ved and simple condition except for relativelyminor interspecific differences with regard to size.Abdomen: The main systematic importance of the abdomen lies inthe structure of the abdominal sclerites, particularly the last (eighth)sternite. This is frequently of specific importance, but, like so manyother characters, it varies, and so it should be used with the usualdegree of taxonomic discretion. In the specific descriptions of thisstudy, the eighth sternite has been discussed in connection with themale genitalia since these structures are closely associated in life andfrequently in slide preparations.Genitalia: The copulatory organs of the male offer only limitedpossibilities for specific and generic delimitation in this family, ascontrasted to their great significance in many groups of Lepidoptera.In some genera (e.g., Cryptothelea) the genitalia can be relied upon indelimiting the genus, but for many psychids of both hemispheresspecies belonging even to different genera scarcely can be distinguishedon the basis of this apparatus alone. This convergence of structuremay be due in part to the very similar mating habits of the membersof this family.Female.?In the more primitive psychids, various specializationshave not advanced as far as in the higher members of this family.In most of the lower forms the female has retained the antennae,functional legs, and a normally sclerotized body wall with the resultthat specific or at least generic boundaries can be determined.Furthermore, the ovipositor is reasonably elaborate and similar in BAGWORM MOTHS OF THE WESTERN HEMISPHERE 17general form to the Tineidae. In the higher psychids, however, allbody appendages have become vestigial and the body walllargely membranous. The external genitalia, in addition, vary frompoorly developed to almost nonexistent. For these reasons it isdoubtful that practical taxonomic characters, applicable on the specificlevel, ever will be discovered for many of the females. One feature ofthe female is apparently present in all known species, and thus itprovides a good character for the family as a whole: the anal hair tuft;however, it is not entirely unique but is believed to occur in certaintineid moths (e.g., Kcarfottia) . Both the internal as well as the external genitalia were examined foras many species as available. Major differences were observed amongwidely separated genera, and minor ones were sometimes found amongclosely related groups (e.g., Oiketicus and Thyridopteryx) ; however,characters for differentiating species apparently are largely absentwithin a given genus. Small differences in the external genitalia of afew species of Oiketicus, after a sufficient series has been studied, maybe determined to have reliable value; considerable variation, however,in the structm^e of this organ is known to exist as noted in approxi-mately four dissections each of 0. kirhyi and 0. platensis. No scleroti-zation is known to occur within the internal organs, but a heavilychitinized and basiphilic structure, which has not been named, wascharacteristically present in the spermathecal duct of all the psychidsexamined. This elongate structure in part may serve in keeping thelumen of this tube open. The most divergent feature of the internalgenitalia were the bifid accessory glands that, owing to their pronouncedvariability, are probably not of specific value within a particular genus.Pupa.?Certain external structures (e.g., ridges, furrows, minutespines, and anal hooks) that evidently are of taxonomic importanceare present on the male and female pupae. These have beendiscussed by Jones (1945) and heavily relied upon by Koehler(1939a). Unfortunately, the structm'es have not been investigatedsufficiently for inclusion in the present study.More attention has been given to the female pupa; consideringthe extreme reduction of the adult, adequate characters in this formwould be very desirable. In this sex, various rugosities on theposterior abdominal segments (the pygidium, in particular) havebeen employed primarily for specific identification. A review ofthe works of Jones and Koehler shows that these minor characters,like those of the larval case, have limited application in theclassification of the family as a whole. In some instances genericgroupings possibly may be indicated by pupal characters, but thishas not been confu-med. Presently it is believed that such charactersare essentially of only specific importance, and thus they should be 18 U.S. NATIONAL MUSEUM BULLETIN 244 considered supplementary to the accepted criteria for specific classi-fication and not a comparable substitute. It is the present author'sconviction that, because of the various complications involved, theerection of new species for this family should not be based primarilyon the female stages. If a species is so uncommon that adult malescannot be obtained easily, it is far better to leave the insect unnameduntil that stage can be collected and studied.Larval case.?For some time, the familiar bags which the larvaeconstruct have attracted considerable attention from various natural-ists. This has been unfortunate as some workers have described newspecies solely from such artifacts. To a large extent this incorrect pro-cedure has resulted from the comparative rarity of the adult male andfrom the misconception that the architecture of the case inevitablywas diagnostic. It is now realized by most workers that the superficialappearance of the bags may vary considerably, depending upon thehost to which a particular larva restricts itself. The basic construc-tion, moreover, is not influenced by such variation, but it may beidentical, or nearly so, for some related species. Thus, if used withdiscretion, the structure of the larval case on occasion may havespecific value, but less frequently, generic.ClassificationBecause a revision of the new world Psychidae necessarily involvesa study of the most specialized members of the family, a study of thistype can offer little toward an understanding of the classification andphylogeny of the group as a whole. Several old world genera, includ-ing some of the most primitive forms of the famUy, were examinedduring the preparation of this work. The results are as yet inconclu-sive and will be discussed more fully in subsequent papers. Thewriter's intention is eventually to revise the North AmericanTineidae and, through such a study and a more complete review ofthe old world annectant groups, to arrive perhaps at a more naturalclassification for the several families of Tineoidea.The precise limits of this "family" have not as yet been ascertained.The examinations of several authors in the higher relationships ofthe bagworms have been so superficial that diverse opinions concern-ing the classification have arisen. The majority of these earlierconcepts were reviewed adequately by Tutt (1900, pp. 117-127).The Psychidae frequently are divided into two groups: the Micro - Psychina and the Macro-Psychina. Several characters, involvingboth morphology and behavior, serve to separate the two groups(Ford, 1945). Intermediate conditions for some of these charactersexist, however, as both Tutt and Ford have pointed out; thus, thedivision is not sharply defined, but it does serve to emphasize certain BAGWORM MOTHS OF THE WESTERN HEMISPHERE 19 o art O ?go + + - O Q _ 05C.2g-gSK? i+ I i+++++++++++ + ^^ + + ++ ++ + Kg t^ t- ^ (^ K_ J^,O l^ t- t- C ^ 00 Ct- 00 t- t- t- t- 5S "^ ^ tl 00COOO 00 00 t^^oo-? -? - ^ 00 t-^107 o ^ O (>? ?.