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    Immature Insects Volume 2 Edited by Frederick W. Stehr Department of Entomology ‘Michigan State University T PUBLISHING COMPANY [ca] ex Credits Figure 33.1 reproduced by permission of the Commonwealth Agricultural Bureaux, Farmham Royal, England Figures 34.1, 34.2, 34.4, 345, 34.6, 34.7 and 34.69 & b from the Insects of Australia (Melbourne University Press). Figures 34.136-34.140 from Dybas, H. D. 1976, The Larval Characters of featherwing and limolodid beetles and theit family relationships in the Staphylinoidea (Coleoptera: Puliidae and Limulodidae), Fieldiana Zoology, 70:29-78, Figures 34.698 and 34.709, Reprinted with permission from the Annals of the Entomological Society of America; copyright 1969 and 1964 by the Entomological Society of America. Figure 36,la-c. Reprinted by permission of the editor from the Journal of Medical Entomology, 1976, Volume 13:107-111. Many Figures used in Chapter 37 are taken from the Manual of Nearetic Diptera Volumes 1 and'2 (McAlpine, et al, 1981, 1987), and ate reproduced by permission of the Minister of Supply and Services, Canada. Figures 37,78, 37.79, 37.150, 37.151, 37.153e-h; Key Figures 37 numbers 32, 35, 37 and St from Merrtt-Cummins: An Introduction to the Aquatic Insects in North America, 2nd Edition. © 1984 by Kendall Hunt Publishing Company, Dubuque, IA. All rights reserved. Cover: Larvae of a tortoise beetle, Physonota alutacea Boheman (Chrysomelida Cassidinae), on wild olive, Cordia boisseri, near Weslaco, Hidalgo Co., Texas, October 1983. Photograph courtesy of Charles W. Melton. Copyright © 1991 by Kendall/Hunt Publishing Company Library of Congres Catalog Card Number: 85-81922 ISBN 0-8403-4639-5 Allrights reserved, No part ofthis publication may be reproduced, Stoced in a retrieval system, of transmitted, i any form or by any ‘means, electronic, mechanical, photocopying, recording, or otherwise, ‘without the prior writen permission of the copyright owner, Printed in the United States of America wo87654321 List of Contributors D.M. ANDERSON, Systematic Entomology Laboratory, USDA, US. National Museum, Washington, D.C. 20560 JR. BAKER, Department of Entomology, North Carolina State University, Raleigh, NC 27650 R. S. BEAL, Jr, Department of Biological Sciences, Northern ‘Arizona University, Flagstaff, AZ 86011 E.C. BECKER, Biosystematics Research Institute, Agri- culture Canada, Ottawa, Ontario, Canada KIA 0C6 RT. BELL, Department of Zoology, University of Vermont, Burlington, VT 05405-0083 Y. BOUSQUET, Biosystematies Research Institute, Agri- culture Canada, Ottawa, Ontario, Canada KIA 0C6 D.E. BRIGHT, Biosystematics Research Institute, Agricul~ ture Canada, Ottawa, Ontario, Canada KIA 0C6 H,P, BROWN, Department of Zoology, University of Okla- hhoma, Norman, OK 73019 D.C. CARLSON, Environmental Specialist, 5229 Butter- ‘wood Cirele, Orangevale, CA 91702 E. LUNA DE CARVALHO, R. do Mercado 28, 2725 Al- ueirao, Portugal K.W. COOPER, Professor Emeritus, Department of Bi- ‘ology, University of California, Riverside, CA 92521 G_A.DAHLEM, Department of Entomology, Michigan State University, East Lansing, Michigan 48824 D.S. DENNIS, 5875 E. Weaver Circle, Englewood, CO soul JR. DOGGER, Systematic Entomology Laboratory, USDA, US. National Museum, Washington, D.C. 20560 H.S. DYBAS (deceased), Field Museum of Natural His- tory, Roosevelt Rd. at Lakeshore Dr., Chicago, IL 60605, RE. ELBEL, Department of Biology, University of Utah, Salt Lake City, UT 84112 B, A. FOOTE, Department of Biological Sciences, Kent State ‘University, Kent, OH 44240 D.E. FOSTER, Department of Entomology, Iowa State University, Ames, 1A 50011 J.H. FRANK, Department of Entomology, University of Florida, Gainesville, FL 32611 B.S. HEMING, Department of Entomology, University of | Alberta, Edmonton, Alberta, Canada T6G 2E3 D.H. KAVANAUGH, Department of Entomology, Cali- fornia Academy of Sciences, Golden Gate Park, San Francisco, CA 94118 M. KOGAN, Section of Economic Entomology, Ilinois Nat- ural History Survey, Champaign, IL 61820 D.M. LABELLA, Department of Entomology and Nema- tology, University of Florida, Gainesville, FL 32611 J.F, LAWRENCE, Division of Entomology, CSIRO-Box 1700, Canberra City, A.C.T. 2601, Australia F. A. LAWSON, 4210 Grays Gable Rd, Laramie, WY 82070 L. LESAGE, Biosystematics Research Institute, Agriculture Canada, Ottawa, Ontario, Canada KIA 0C6 J.E. LLOYD, Department of Entomology and Nematology, University of Florida, Gainesville, FL 32611 D.R. MILLER, Systematic Entomology Laboratory, 11B111, ‘ARS USDA, Beltsville, MD 20705 H.H. NEUNZIG, Department of Entomology, North Car- ‘lina State University, Raleigh, North Carolina 27650 A.F. NEWTON, Jr, Field Muscum of Natural Histor Roosevelt Ra. at Lakeshore Dr, Chicago, IL 60605 L. B.O'BRIEN, Laboratory of Aquatic Entomology, Florida ‘A-& M University, Tallahassee, FL 32307 G.S. PFAFFENBERGER, Department of Life Sciences, Eastern New Mexico University, Portales, NM 88130 H, REICHARDT (deceased), Museo de Zoologia, Univer- sidade de Sdo Paulo, Sao Paulo, Brazil R. B. SELANDER, Department of Entomology, University of Illinois, Urbana, TL 61801 P. J. SPANGLER, Department of Entomology, Smithsonian Institution, Washington, D.C. 20560 T.J.SPILMAN, Systematic Entomology Laboratory, USDA, US. National Museum, Washington, D.C. 20560 xv LIST OF CONTRIBUTORS TA. STASNY, Department of Entomology, West Virginia University, Morgantown, WY 26505 FW. STEHR, Department of Entomology, Michigan State University, East Lansing, MI 48824 M.B. STOETZEL, Systematic Entomology Laboratory, IBIL1, ARS, USDA, Beltsville, MD 20705 C.A. TAUBER, Department of Entomology, Cornell Uni- versity, Ithaca, NY 14853, H. J, TESKEY, Biosystematics Research Institute, Agricul- ture Canads, Ottawa, Ontario, Canada KIA 0C6 F.C. THOMPSON, Systematic Entomology Laboratory, USDA, US. National Museum, Washington, B.C. 20860, MANUEL G. DE VIEDMA, (deceased) Universidad Pol tecnica de Madrid, Escuela Tecnica Superior de, Ingen- ieros de Montes, Madrid, Spain Q. D. WHEELER, Department of Entomology, Cornell Uni- versity, Ithaca, NY 14853 T.R. YONKE, Department of Entomology, University of Missouri, Columbia, MO 65211 D.K. YOUNG, Department of Entomology, University of Wisconsin, Madison, WI $3706 Introduction This introduction is intended to supplement that of Volume 1. Volume 2 is being published about three years after ‘Volume 1 (1987). The original idea for putting these books together was generated about 20 years ago and evolved along, the way. It may seem unusually long, but 15-20 years seems tobe 2 rather “normal” gestation period fora large work on larvacin North America, since Biving and Craighead (1931) indicate that they began working together on their publica- tion on Coleoptera larvae in L915 when they both realized the other was working independently ona similar project, and AAlvah Peterson (1948 and 1951) began working on his two volumes in the 1930s. The importance of immatures cannot be overempha: ‘zed, since they are found just about everywhere, cause much ‘more economic damage overall than adults, are key indica- tors of soil and water quality, are important decomposers. are key items inthe diets of many vertebrates, especially young birds and fish, and are essential components of terrestrial and fresh water ecosystems. In addition, they have unusual uses such as forensic entomology (Smith 1986), and the data from immatures have made and will continue to make major con- tributions toward better classifications and better phylo- ‘genies of insect taxa. ‘The original classification of any group of insects has almost always been based exclusively on adults but classifi- cations and phylogenies will always be better when all pos- sible data including that from immatures, are used in their construction. A summary of the taxonomic significance of the characters of immature insects whichis still useful was pro- vided by Emden (1957). If we believe that a classification is 4 framework within which all of the data about organisms should be fled, then we must use all the available data in building that framework, if it is 10 be useful to all who seek tofile or retrieve data Its obvious that most larvae of the Holometabola are adapted for living in very different habitats than the adults, and have evolved substantially different structures and be- hhaviors than the adults. Despite this fact, there is relatively lite discordance between suites of adult and larval charac: ters, and many times the immatures have pointed the way toward the correct placement of a puzzling species or group, whether as a part of an existing taxon or a8 a new one. ‘At higher levels, the classification of the major subor- ders of the Hymenoptera, Diptera and Coleoptera works well for adults and larvae, and characters of immatures and adults have been used in developing the suborders. However, in the Lepidoptera, the two major suborders, Monotrysia and Di: tnysia, do not separate as cleanly, perhaps because of the ex- ‘treme specialization of many monotrysian larvae for a diversity of stem-boring and leaf-mining habits. Occasion- ally, as in the Siphonaptera in this book, there is some dis- agreement about classifications based on adult or larval ‘characters, but there can be only one classification and it must include all life stages, o any differences must eventually be resolved. ‘A teview of the contributions of the use of immature stages to classification and phylogeny would be a long chapter in itself, but a'small sampling follows. including some con- tributions by contributors to these volumes. Edmunds and Allen (1966) have discussed the significance of larvae in the study of Ephemeroptera, where larvae are more readily available and where identification of larvae is needed much ‘more often than adults. Immature Coccoidea (Homoptera), especially crawlers and second instar males, have been of great value in the study oftheir classification and evolution (Miller and Kosztarab 1979; Howell 1980), especially since the adult female scales are heavily sclerotized and have relatively few characters. For the Trichoptera, Wiggins (1981) has pre- sented a strong review of the relevance of immatures to thei systematics. Lepidoptera larvae, especially first insta, have been widely used in the establishment of classifications and phylogenetic relationships (Hinton 1946), and the placement ‘of some groups has only been resolved after the larvae have becomie known (Kristensen and Nielsen 1983). In the Co- leoptera, Crowson (1955) used larval data and all other ava able data in arriving at his classification, and he reiterated the theory that the larval and life cycle data were the best evidence for believing that the Coleoptera are most closely related to the Neuroptera, particularly the Megaloptera. Hy- ‘menoptera larvae have long been used, and Yuasa's larval work (1923) made an early contribution to symphytan sys- tematics and evolution. Among the Apocrita, Michener (1953), Evans (1964), Bohart and Menke (1976), McGinley (1981) and Carpenter (1982) have all made extensive use of| Iarvae in arriving at better classifications. Individuals, both amateur and professional, have made important contributions to our immature eallections by theit long and dedicated work in rearing many species. The asso- ciation of immatures with adults is without doubt one of the ‘most important contributions that can be made by ama- teurs—it ean be done with minimal equipment and some genuity, nd without the aid of a good microscope as long as the immatures are properly killed, labeled and preserved. It ‘must be emphasized that immatures of any life stage should rot be placed with other immature stages or adults unless the association is known to be correct. All key data must be with all ife stages, as well as an entry or a special label listing all the associated life stages. Log books are very useful, but tend xvl—_ INTRODUCTION to become misplaced, lst or unavailable, so as much data as is reasonable must be on the label(s). Ideally, al stages would be stored together, but this may not be practical ifthe adults are best preserved pinned and the immatures are best pre- served in aleohol. A list of useful publications on techniques is given in Chapter 2 in Volume 1, to which can be added Steyskal etal. (1986). We have a long way to go before the immatures are as well known as the adults. Those who study or are interested inimmatures are urged to collect and rear larvae, or toobtain ges from females when feasible and rear larvae (this isthe best technique since the association is certain and larvae of | all instars can be preserved). All life stages (including the cast larval and pupal skins) and the associated adults should bbe deposited in major museums for future study, and publi- cations should require that voucher specimens of newly de~ scribed life tages must be deposited in major museums. An increased knowledge of immatures will strengthen the sci cence of entomology in many ways, but especially inthe eco- ‘omic, ecological, systematic, and evolutionary arenas. Literature cited: Bohart, R. M.,and A. S. Menke. 1976. Sphecid wasps of the world, A’generic revision. Berkeley and Los Angeles Univ. Calif. Press. 695 pp. Baving, A.G., and F.C. Craighead. 1931. An illustrated synopsis of the principal larval forms of the order Co- leoptera. Brooklyn Entomol. Sor. 351 pp. Carpenter, J. M. 1982. The phylogenetic relationships and natural classification of the Vespoidea (Hymenoptera), ‘Systematic Entomology 7:11-38. Crowson, R. A. 1955. The natural classification of the fam- ilies of Coleoptera. N. Lloyd, London. 187 pp. Edmunds, G. F,, Jr. and R. K. Allen. 1966. The significance ‘of nymphal stages in the study of Ephemeroptera. Annals Ent. Soc. Amer. 59:300-303. Emden, F. I, van, 1957. The taxonomic significance of the characters of immature insects. Annu. Rev. Ent. 2:91~ 106, Evans, H. E, 1964, The classification and evolution of digger ‘wasps as suggested by larval characters (Hymenoptera Sphecoidea). Ent, News 75:225-237. Hinton, H.E. 1946. On the homology and nomenclature of setae of lepidopterous larvae, with some notes on the phylogeny of the Lepidoptera. Trans, Roy Ent. Soc. Lond. 9731-37. Howell, J.0. 1980. The value of second-stage males in ar- moured scale insects (Diaspididae) phyletics. Israel J. Ent, 1487-96 Kristensen, N.P. and B.S. Nielsen. 1983. The Hetero- bathmia life history elucidated: immature stages con- tradict assignment to suborder Zeugloptera (Insecta, Lepidoptera). Sonderdruck aus Zeitschrift {Ur Zoo! Systematik u Evolutionsforschung 21:101-24. McGinley, R. J. 1981. Systematics of the Colletidae based on ‘mature larvae with phenetic analysis of apoid larvae (Hymenoptera: Apoidea) J. Kan. Ent. Soc. 53:539-552. Michener, C. D. 1953. Comparative morphological and sys- tematic studies of bee larvae with @ key to the families cof hymenopterous larvae. Univ. Kan. Sci Bull. 35:987- 102. Miller, D. R. and M. Kosztarab. 1979. Recent advances in the study of scale insects. Annu. Rev. Ent. 24:1-27. Peterson, A. 1948, 1951, Larvae of insects. An introduction to Nearetic species. Printed for the author by Edwards Bros. Ann Arbor, Mich. Pt. 1, 315 pp. Pt. 2, 416 pp. ‘Smith, K. G, B. 1986. A manual of forensic entomology: Cor- ‘ell Univ. Press, thaca, N.Y. 205 pp. ‘Steyskal, G. C., W. L. Murphy and E. M, Hoover (eds.) 1986, Insects and mites: techniques for collection and preser- vation. US. Dept. Age., Agr. Res. Ser, Mise, Publ. 1443, 103 pp. Wiggins, G. B, 1981. Considerations on the relevance of im- mature stages to the systematics of Trichoptera. Proc. 3rd Symp. on Trichoptera, G. P. Moretti (ed.). Series Entomologica 20:395-407, ‘Yuasa, H. 1923 (1922). A classification ofthe larvae of the ‘Tenthredinoidea, Il, Biol. Monogr. 7. 172 pp. Order Coleoptera 341: John F, Lawrence, coordinator Division of Entomology, CSIRO BEETLES" ‘The Coleoptera is the largest order of inseets, with more than 350,000 species worldwide and about 25,000 occurring in America north of Mexico. In the present book, they are placed in 156 families, 31 of which do not cccur in the United States or Canada, Larvae of Coleoptera, unlike those of some ‘other orders, have no one common name that can be applied to all forms, and various terms like grub, white grub, wire- worm, mealworm, rootworm, glow worm, round-headed borer, flat-headed borer, timber borer, water penny, etc. have been used for individual larval types. Most beetles are terrestrial, but a number of genera and some families are aquatic as larvae, some occur in water throughout most (or rarely all) of the lifecycle, and a few are aquatic inthe adult stage only. The majority of beetle larve feed on various kinds of living and dead plant tissue (roots stems, trunks, branches, logs, leaves, flowers, seeds), but many feed on fungi, carrion or dung, some are predaceous, and a few are parasitic. Many Coleoptera cause damage to agricultural crops, forests, or stored products, but very few are of medical importance DIAGNOSIS There is no single feature which will distinguish beetle larvae fram those of other insect orders and only a few which ‘may be considered universal within the group. The following list includes most characters considered of diagnostic value 1. Head capsule well-developed and usually sclerotized 2. Head usually without paired endocarinae (adfrontal ridges) forming a V or Y, and never with adfrontal areas, formed between them and the frontal sutures or eedysial lines; paired endocarinae, when present, very rarely ex: tending to the anterior part of the frontoclypeal region. 3. Antennae almost always with 4 segments or fewer and with a seasorium on the penultimate segment. 4. Number of stemmata on each side always 6 or fewer. 5. Mouthparts almost always of the chewing or orthop- teroid type with opposable mandibles moving in a trans- verse plane, and with normal palp-bearing maxillae and labium:; rarely modified to form a sucking tube bearing. stylts, or with immovable or non-opposable mandibles. 6, Median labial silk gland or spinneret always absent ‘in this section a gure reference suchas (Eig. 121 p90) refers to Key fig. 121, page 190 144 7. Legs usually with 5 or 6 segments (coxa, trochanter femur, tibia, and cither tarsungulus or tarsus and prel tarsus, the latter consisting of one or two claws) or absent; occasionally with reduced segmentation. 8, Abdomen usualy with 10 segments (occasionally fewer), without articulated appendages (cerci) on segment 10. 9, Paired abdominal prolegs almost always absent! asperity-bearing prolegs occasionally present on sterna 2-4, 3-4 of 2-5; simple prolegs rarely present on sterna 1-8; crochet-bearing prolegs rarely present on sterna 2Tor 3-7, 10, Respiratory system usually peripneusti, without fune- tional spiracles on metathorax; occasionally amphi pneustic, metapneustic, o apneustc. I. Spiracles. often with accessory openings (annular- uniforous, annular-biforous, annular-multiforous), with divided opening (biforous), or with poroid sieve-plate (cribriform); ecdysial sear, if present, never completely enclosed by sieve plate Because of the great diversity of form within the Co- leoptera, at least some beetle larvae may be confused with immatures belonging to cach ofthe other endopterygote orders and probably those ofa few exopterygote groups as well The ‘riungulinids of Strepsiptera are said to resemble the triun- gulins of Meloidae and Rhipiphoridee, but they differ from them in several respects, including the absence of mandibles, antennae, and labial alps, the lack of trochanters inthe legs, land the presence of pair of long setae on the terminal 10th segment. The endoparasitic forms of Strepsiptere (larvae and adult females) are easily distinguished because of their ex tremely reduced body which is indistinctly segmented. The free-living last larval instar (puparium) in Mengenilidae has ‘S-segmented legs with 2 claws but differs from beetle larvae in the presence of compound eyes composed of numerous facets. Larvae of Tichoptera, Megaloptera, Raphidioptera, and [Neuroptera may be distinguished from most beetle larvae by the presence of 6-segmented legs, which in the Coleoptera ‘occur only inthe suborders Archostemata and Adephaga. The distinctive head of Cupedidae and Micromalthidae, with a ‘median endocarina, well-developed mandibular molae, and a sclerotized ligula, separate them from members ofthe above orders, while the absence of a free labrum will distinguish adephagan larvae fram those of Trichoptera, Megaloptera, ‘and Raphidioptera. Neuroptera larvae also have the labrum fused to the head capsule, but they differ from any beetle larvae in their distinctive feeding apparatus, consisting of a pair of sucking organs formed by the falciform or styliform mandibles and maxillae on each side, and by the complete absence of maxillary palps. The divided, blade-like mandi- bles of Lycidae or the perforated, sucking mangibles of Lam- pyridae and Phengodidae might be confused with neuropteran ‘mouthparts, but these groups always have maxillary plps and S-segmented legs. Six-segmented legs also occur in 2 rare families: Nannochoristidae (Mecoptera) and Heterobathmi- idac Lepidoptera}; larvac of both may be distinguished from those of Coleoptera by the combination of leg segmentation and free labrum, while the former has more than 6 stemmata ‘on each side and panorpoid spiracles (with numerous open- ings surrounding an ecdysial scar), and the latter has ad- frontal ridges, adfrontal areas, and a median labial gland, Larvae of Lepidoptera are easily confused with various beetle larvae, especially those of Chrysomelidae, but all of the former differ in having @ median Isbial gland, which is usually developed into a protruding spinneret, almost all of| them differ in having paired adfrontal ridges, with adfrontal areas formed between them and the ecdysial lines, and most of them differ in having paired, crochet-bearing prolegs on abdominal sterna 3 to 6 und 10. The median labial gland may ‘be difficult to see in some primitive forms without distinct spinnerets (Micropterigidae), and the adfrontal ridges are absent in Micropterigidae and Agathiphagidae. A number of ‘roups lack prolegs, while some also lack thoracic legs and. ‘may be confused with apodous Coleoptera larvae; al of these ‘roups, however, have the typical lepidopteran head. Several beetle larvae have abdominal proleps, but these are usually simple or asperate, without crochets, and occur on different abdominal segments than those in most Lepidoptera (1 to 9 in schizopodine Buprestidae; 2 or 3 0 7 in Hydrophilidac; 2 04, 3 to 4, or 2 to 5 in Oedemeridae; and 1 to 8 in Cureu- lionidae), Crachets occur on the prolegs of some Hydrophil- idae, but these larvae differ from any lepidopteran in having biforous spiracles and the labrum fused to the head capsule Larval Micropterigidae differ from those of other Lepidop- tera in having simple, more or less acute, mecopteran-like prolegs, which are not always distinct, on segments 1 to 8 “The lack of lepidopteran head characters makes them par- ticularly dificult to separate from beetle larvae; but the com- bination of a more or less hexagonal shape in exoss-section, a retracted head, the lack of an antennal sensorium, 3-segmented legs, and a vestiture consisting of characteristic thickened setae should distinguish them, Three-segmented thoracic legs and simple prolegs on segments | to 8 will also distinguish larvae of most Mecop- tera from those of Coleoptera, and, in addition, most Mecop- tera larvae have panorpoid spiracles, more then 6 stemmata on each side, and no distinct cardo. Larvae of Boreidae have vestigial spiracles, no prolegs, and only 3 stemmata, but they ray be distinguished by their short, clawless, 3-segmented legs, which are very close together on the prothorax and widely separated on the meso- and metathorax ‘Various Hymenoptera larvae are easily confused with those of beetles, although all possess a median labial gland (not always easy to see) and many have 2 thoracie spiracles. Larvae of Apocrita and some Symphyta (Orussidae) are lightly sclerotized and legless, with a hypognathous head, and ORDER COLEOPTERA 145 ‘are most likely to be confused with some Meloidae, Rhipi- phoridae, and Curculionidae; in addition to the labial gland and the presence of a 2nd thoracic spiracle (in some), the for mer groups have reduced maxillary and labial palps (I-segmented or absent). Larvae of most Symphyta differ from those of Coleoptera in having paired prolegs on abdom- inal segments 2 to 7 or 8 and usually 10 (Tenthredinidae, Ar gidae, Cimbicidae, Diprionidae), more than 4 antennal Segments (Blasticomidae, Pamphilidae, Xyelidae), or more than 4 plicae on most abdominal segments (Cimbicidae, Di prionidae). The only beetle larvae with prolegs on the same Segments are Hydrophilidse, which have a fused labrum and biforous spiracles, never occurring in Hymenoptera, while those few beetle larvae with more than 4 antennal segments either have long, flagellate antennae, highly modified mouth- parts, and a terminal respiratory chamber (Helodidae) or a ‘median endocarina, -segmented legs, and mandibular molae (Cupedidae). Larvae of the last group bear a superficial re- semblance to larvae of Cephidae, and both have 4 or $ an tennal segments and a median spine (suranal process) on tergum 9. Of those Symphyta larvae with fewer antennal seg ments, fewer plicae, and no prolegs, the Xiphydriidae and Siricidae both have vestigial legs and a suranal process, much as in larvae of Mordellidae; they differ from mordellids in having elongate-lliptical spiracles. Some surface-feeding Pergidae which lack prolegs differ from any beetle larvae in having single large stemma on each side and the antennae reduced to dome-like structures; alo there is usually a smaller, but obvious, 2nd thoracic spiracle. The apodous larvae of Diptera can often be distin. guished from legless beetle larvae on the bass of their incom: plete head capsule and/or vertically oriented, non-opposable mandibles; in the Nematocera, however, most larvae have & well-developed head capsule and transverse, opposable man bles, as in most Coleoptera. Except for some aquatic forms without functional spiracles (Chironomidae, Ceratopogon- idae, Simuliidae), most nematoceran larvae may be distin- ‘Buished from those of beetles by the type and location of their spiracles, as well as by the universal absence of a spiracular closing apparatus (present in all legless Coleoptera). The presence of spiracles on the thorax and 8th abdominal sea- ‘ments only (amphipneustic system, as in Tanyderidae, Psy- ‘chodidae, Trichoceridae, etc.) and the presence of a single pair of spiracles on segment 8 (metapneustic system of Ti+ pulidae, Culiidae, etc.) are conditions known among Co- leoptera, but not in groups with apodous larvae, Bibionidac and a few other groups have a holopneustic system with 2 thoracic and 8 abdominal spiracles on each side, @ condition unknown in Coleoptera; and the remaining Nematocera with 1 petipneustc system usually differ from beetles in having the thoracic and last abdominal spiracles enlarged. The spir- acles of Nematocera are either of the panorpoid type, with 3 ring of small openings surrounding an ccdysial scar, oF they have a characterstie cribriform plate with 3 openings spir- cles are never of the biforous type (as in sphaeridiine Hy-

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