^cAi qasin naturalist memoirs dumber 10 Brigham Young University 1986 A Rec!assff:catbn ^* '' . Genera of Scolytidae (Coleoptera) APR 10 1987 GREAT BASIN NATURALIST Editor. Stephen L. Wood, Department of Zoology, 290 Life Science Museum, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Chairman, Botany; James R. Barnes, Zoology; Hal L. Black, Zoology; Stanley L. Welsh, Botany; Clayton M. White, Zoology. All are at Brigham Young University, Provo, Utah 84602. Ex Officio Editorial Board Members. Bruce N. Smith, Dean, College of Biological and Agricultural Sciences; Norman A. Darais, University Editor, University Publications. Subject Area Associate Editors. Dr. Noel H. Holmgren, New York Botanical Garden, Bronx, New York 10458 (Plant Taxonomy). Dr. James A. MacMahon, Utah State University, Department of Biology, UMC 53, Logan, Utah 84322 (Vertebrate Zoology). Dr. G. Wayne Minshall, Department of Biology, Idaho State University, Pocatello, Idaho 83201 (Aquatic Biology). Dr. Ned K. Johnson, Museum of Vertebrate Zoology and Department of Zoology, University of California, Berkeley, California 94720 (Ornithology). Dr. E. Philip Pister, Associate Fishery Biologist, California Department of Fish and Game, 407 West Line Street, Bishop, California 93514 (Fish Biology). Dr. Wayne N. Mathis, Chairman, Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560 (Entomology). Dr. Theodore W. Weaver III, Department of Botany, Montana State University, Boze- man, Montana 59715 (Plant Ecology). The Great Basin Naturalist was founded in 1939 and has been published from one to four times a year since then by Brigham Young University. Previously unpubhshed manuscripts in English of fewer than 100 printed pages in length and pertaining to the biological natural history of western North America are accepted. 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See Notice to Contributors on the inside back cover. 12-86 1200 27830 ISSN 017-3614 9REAT BASIN NATURALIST MEMOIRS lumber 10 Brigham Young University 1986 A Reclassification of the Genera of Scolytidae (Coleoptera) CONTENTS ABSTRACT 1 INTRODUCTION 1 HISTORY 2 REVIEW OF CHARACTERS 3 DISCUSSION 6 PHYLOGENY 21 METHODS 24 SYSTEMATIC SECTION 25 Key to the families of Curculionoidea 26 Key to the subfamiHes and tribes of Scolytidae 28 Subfamily Hylesininae 34 Tribe Hylastini 34 Tribe Hylesinini 36 Tribe Tomicini 40 Tribe Phrixosomini 43 Tribe Hyorrhynchini 43 Tribe Diamerini 44 Tribe Bothrosternini 46 Tribe Phloeotribini 48 Tribe Phloeosinini 49 Tribe Hypoborini 53 Tribe Polygraphini 55 Subfamily Scolytinae 57 Tribe Scolytini 57 Tribe Ctenophorini 59 Tribe Scolytoplatypodini 61 Tribe Micracini 61 Tribe Cactopinini 65 Tribe Carphodicicini 66 Tribe Ipini 67 Tribe Dryocoetini 70 Tribe Crypturgini 75 Tribe Xyloterini 76 Tribe Xyleborini 79 Tribe Xyloctonini 84 Tribe Cryphalini 85 Tribe Corthylini 93 Subtribe Corthylina 97 Subtribe Pityophthorina 98 LITERATURE CITED 100 INDEX Ill Great Basin Naturalist Memoirs A Reclassification of the Genera of Scolytidae (Coleoptera) No. 10 Brigham Young University, Provo, Utah 1986 Stephen L. Wood Abstract. — A ta.\oiiomic revi.sion of the genera oi Scolytidae (Coleoptera) in tlie world fauna i.s pre.sented. Included are 215 valid genera and 27.3 invalid generic and subgeneric name.s. The type-species for each genus-group name was examined, including the type-specimen of the type-species in those taxa where a holotype. Icctotype, or neotype has been designated — with the exception of four contemporary genera, the type-series of which are lost, and six fossil genera for which no eflort was made to locate the types. Taxonomic keys to the families of the Curculionoidea, and to the subfamilies, tribes, and genera of Scolytidae are presented. Descriptions and citations of the original validations of the two subfamilies and 25 tribes of Scolytidae are included. For each genu.s-group name there is a citation of its original validation, type-species, and synonymy. For each valid genus there is an indication of its distribution, the approximate number of valid species, basic feeding and mating habits, and citations of pul)lished ke\'s foi' the identification of species. Of general interest to coleopterists is a review of the status and position of the families Platypodidae and Scolytidae within the Curculionoidea, including several characters not previously presented. These two families are entirely distinct from one another and from Curculionidae. The phyletic position of Pfatypodidae and Scolytidae is near the more primitive families oi Curculionoidea and quite remote from specialized Curculionidae. Because of their immense economic impact caused practical foresters to enter the fields of on world forests, forest products, and certain naming and identifying species to meet their aspects of agriculture, considerable attention own limited needs. With two pre-1960 e.xcep- has focused on the Scolytidae. Although the tions, taxonomic work on Scolytidae was done literature is filled with reports of the economic by senior foresters who lacked a fundamental ravages and of efforts made to manipulate interest or training in the study of diversity, their populations, comparatively little atten- phylogeny, or evolution. These exceptions tion was given to other aspects of their study were Chapuis, who produced a classical work prior to 1960. Since then, there has been a on Platypodidae (Chapuis 1S65) and was start- virtual explosion of information about many ing to work on Scolytidae (Chapuis 1869) be- other facets of their existence. fore jurisdictional problems arose, and Hage- Although advances in basic commimication dorn, whose brief career (1904-1912) ended involving nomenclature and classification nor- tragically. The unwillingness to accept and mally preceded the expansion of knowledge in apply the concept of evolution and to abandon other groups of organisms, this was not the the typological (morphological) species con- case with the Scolytidae. There appear to be cept by more recent influential workers also several reasons for this delay. The sheer num- delayed progress. ber of species and the urgency to do some- The present study iiad its origin in a conver- thing with the management of populations sation with C. D. Michener, at Logan, Utah, compelled local foresters to act. The un- in August 1949, during wliich he suggested a availability of trained taxonomists apparently reclassification of the genera of Scolytidae in Life Science Museum and Department of Zooloey, Bn^tlam ^'nuni; Uni/'ersitv. I*r<»\(>, I'tali S.l(i()2, I'S.A Great Basin Naturalist Memoirs No. 10 the world fauna as an appropriate subject for my doctoral dissertation. Fortunately, for me and for science, a more limited topic was cho- sen, but the suggestion was never forgotten and has remained a lifelong objective. In 1981, when I was asked by colleagues to final- ize a comprehensive world catalog of Scolyti- dae, completion of the generic study became mandatory and urgent. Although this presen- tation is a bit premature, its basic objectives are achieved. More thoughtful selection of illustrations and greater detail in their presen- tation would have been helpful to the user. The Scolytidae are among the more difficult insects to classify; yet their tremendous eco- nomic importance is so great that immediate and precise identification is demanded by those industries threatened by the activities of these insects. Although my attraction to the Scolytidae was to their fundamental struc- ture, diversity, ecology, and behavior from theoretical and other points of view, as knowl- edge began to accumulate, my rural back- ground and religious philosophy made me sensitive to the economic interests of others and to a need for me to assist them with what I had learned. Another need was also para- mount. Among the great frustrations in learn- ing are those resulting from errors that have been transmitted from one generation to the next and, once learned, must be unlearned and corrected before progress can continue. Biological nomenclature and classification have been profoundly impeded in their pro- gress by this problem. Therefore, the primary objectives of this study are (1) to review the holotypes or syntypes of the type-species of all named genera and subgenera to make certain that they are correctly placed in appropriate genera, then find the correct generic names for them, and review as many species as possi- ble and group them in those genera, and (2) to devise a scheme of classification such that oth- ers can identify those genera. If these objec- tives are achieved in at least some measure, it is hoped that the constant name changing and duplication of effort (resulting from syn- onymy) that has impeded taxonomic progress in my generation will be reduced to a mini- mum. This classification of categories above the species level will be the framework on which the catalog of species, now in prepara- tion, will be organized. History Linnaeus (17.58) included five species of Scolytidae in his Systema Naturae, edition 10, all under the name Dennestcs (typographus , micrographus , poligraphus , piniperda, do- mesticus). Fabricius (1801) expanded this to include 52 species under the generic names Apate (5), Bo.strichiis (21), and Hylesinus (26). Gemminger and Harold (1872) list 535 species in 60 genera. Hagedorn (1910a, b) lists 1,234 species in 115 genera. There has been no comprehensive classification or catalog of the family on a worldwide basis since the work of Hagedorn, but it is estimated that more than 6,000 biological species are currently recog- nized. The first generic name erected for a member of this family was Scohjtiis Geofifroy (1762; Ghina 1962, International Commission on Zoological Nomenclature 1963). Since then, 488 generic and subgeneric names have been proposed for the group, of which 215 are treated below as valid. In an attempt to estimate the number of species remaining to be named from America north of Mexico, White (1975), in his example using Scolytidae, recognized 592 species in 1970 and projected that there would be 650 named by the end-point year of 2040, when 100% would be known. That estimate was made prior to publication of my monograph (Wood 1982). In the monograph, numerous species were placed in synonymy and others were named or they extended their distribu- tions into the United States and Ganada, leav- ing about the same total number of species and projection as before. Meanwhile, more than half of the scolytid fiiuna of Mexico and Gentral America was named from 1960 to 1982. This rapid expansion of the fauna south of the United States made a projection for the larger area meaningless at the present time. Few attempts to classify the higher cate- gories of Scolytidae on a worldwide basis have been published. Perhaps the first contribu- tion worthy of note was that of Ferrari (1867), who listed and described the genera of Tomi- cides (my Scolytinae with Scolytini deleted). There was no attempt to actually classify the genera by Ferrari. Eichhoff (1878a) published a classical monograph of this same group in which he presented keys and descriptions to all known genera and species. His family 1986 WOOD: Genera of Scolytidae Tomicini included 40 genera that were dis- tributed among the subfomiUes Crypturgidae {Cn/pturgus, Dohirgus, Aphanarthruin, Trio- teranus, Pijcnarthrum), Cryphahdae (Li- parthntm, Hijpohorus, Triannoccnis, Cnj- phahis. Glyptodenia, Stephaiwdercs, Cos- moderes), ProblechiUdae (Prohlechilus), Xy- loctonidae {Xijloctonus , Scohjtogenes), Pity- ophthoridae (Pityophtlionis, Eidophidus, Taphrorijchus), Tomicidae (Thamnurgtis, Xijlocleptes, Tomicus, Lepicenis, Drijocoe- tes), Hylocuridae (Hylociirits), Micracidae (Micracis), Araptidae (Araptus), Hexacohdae {Hexacolus), Xyleboridae (Coccotrypes, Xyle- horus, Premnobiiis, Gnathotrichiis), Xylote- ridae (Trypodendroti), Corthyhdae (Corfhy- lus, Brachyspartus, Anchonoceriis, Phtho- rius, Trypocranus, Pterocyclon), and Amphi- cranidae {Steganocranus, Amphicranus). Hagedorn (1910a, b) treated 115 genera and 1,234 species in the entire family worldwide. He divided Scolytidae (Ipidae, in his usage) into four non-Linnean subfamilies, based on the detailed structure of the mouthparts, that contained the tribes indicated: Pilidentatae (Phloeotrupinae), Spinidentatae (Eccoptogas- trinae, Diamerinae, Crypturginae, Hylocuri- nae, Hylesininae, Ipinae, Cryphalinae), Sae- tidentatae (Xyleborinae, Corthylinae), and Mixodentatae (Spongicerinae). Hopkins (1915a, b), in two papers, (a) re- classified the subfamilies and tribes of Scolyti- dae and (b) reclassified the genera and species of Cryphalinae. He elevated the group to su- perfamily rank, Scolytoidea, and recognized four families within it (Ipidae, Scolytidae, Scolytoplatypodidae, and Platypodidae). His Ipidae included the subfamilies Cryphalinae, Ipinae, Micracinae, Webbinae, Xyloctoni- nae, Crypturginae, Phloeotribinae, Hylesini- nae, and Phloeoborinae. His Scolytidae in- cluded the subfamilies Coptonotinae, Hexa- colinae, Bothrosterninae, Camptocerinae, and Scolytinae. His Scolytoplatypodidae in- cluded the subfamily Scolytoplatypodinae, and Platypodidae the subfamilies Platypodi- nae, Genyocerinae, and Chapuisinae. It ap- pears to have been the intention of Hopkins to follow his basic classification with a series of papers treating in detail the various divisions of his classification, but only one paper was published. Hopkins (1915a) reviewed the sub- family Cryphalinae, in which he included 48 genera that are currently distributed (below) among the tribes Cryphalini, Dryocoetini, and Xyleborini. Wood (1978) reviewed 15 major published classifications of the higher categories of Scolytidae, including all of those cited above, and presented a revised and expanded classifi- cation of subfamilies and tribes and tenta- tively assigned 404 nominate genera and sub- genera to his 25 tribes. That classification was adapted to his monograph of the North and Central American Scolytidae (Wood 1982), which listed 1,4.33 species in 94 genera. The present contribution is an expansion of that classification to the genus level for the entire world. Review of Characters Characters Shared by Platypodidae and Scolytidae In the early stages of this study, 1 was thor- oughly convinced that Platypodidae could be no more than a well-marked, primitive sub- family of Scolytidae. Shared characters that led to this premature conclusion included: (1) clearly formed pregular sutures extend from the anterior end of the median gular suture to or near the anterior tentorial pits; these su- tures are reinforced internally by a massive internal inflection of the cuticle that occurs nowhere else in the Curculionoidea; (2) the mandible lacks the horizontal cylinder that rotates on the anterior and (heavily rein- forced) posterior condyles, with its cutting edge apical; instead, the posterior condyle is in a different position, the axis of the mandibular hinge is transverse, and the cut- ting edge is mesal (as in Anthribidae and Nemonychidae); (3) the rostrum is short to nonexistent; (4) the tibiae totally lack corbels, but they share a common basic structure and are armed on the apical and lateral margins by a characteristic series of spines; (5) the eyes are essentially flat against the head; (6) the antennae are mostly of the same basic genicu- late type, with some members of several primitive tribes having a short scape (little if any longer than the pedicel); (7) the visible abdominal sterna 1 and 2 are weakly connate, except all are free in all primitive genera of Platypodidae; (8) the elytral interstriae 10 ex- Great Basin Natuiulist Memoirs No. 10 Fig. 1. Outline of cephalic aspect of the head of Pst?urfo- hylesinus sericeus (Mannerheim), with frontal rectangle indicated by the square of dash lines. Fig. 2. Cephalic aspect of the head of a female Chaetas- tiis tuberculatus (Chapuis) (Platypodidae) showing a con- spicuous "labrum ' in the median area just below the epistomal margin arrow. (Re-drawn from Strohmeyer (1920: 15, Fig. 5). tends to near the apex in all Platypodidae and in some primitive representatives of virtually all tribes of Scolytidae; (9) the larval characters are basically the same except as noted below; and (10) they share the same basic ecological niche. Characters Not Shared with Other Curculionoids Curculionoidea, excluding Anthribidae, Nemonychidae, Platypodidae, and Scolyti- dae, share a unique mandibular character in which the basal half forms a cylinder that ro- tates on a horizontal axis between the two articulating condyles (Fig. 8). The hypostomal area adjacent to the posterior condyle is en- larged and strengthened to accommodate a new mode of action; the cutting edge is apical and usually takes the form of a large cusp on the side of the basal cylinder. In the four families identified above, the mandibular condyles are closer together, the oblique ac- tion is more like that of a hinge, the cutting edge is mesal, the mandible ends in an acute point, and the hypostomal area is minimal. The Anthribidae and Nemonychidae have an adult labrum. It is lost in all other Cur- culionoidea; however, in most primitive tribes of Scolytidae (most Hylesininae, Cten- ophorini, etc.) there is an epistomal lobe that resembles a small, fused labrum and is proba- bly homologous to it (Fig. 1). In Platypodidae the epistomal lobe is more generally present, and in Tesserocrerus and Chaetastus (Fig. 2) a suture separating this lobe from the epistoma is evident in some species. Comparable struc- tures are unknown in the remainder of the Curculionoidea. In general, beetles have two gular sutures that are separated by a median gular sclerite. In Nemonychidae (Fig. 3) and some Belidae (Fig. 3B, and Crowson 1967) two gular sutures are clearly indicated from the margin of the foramen magnum to the point of invagination of the tentorium (posterior tentorial pit) where they end. In Anthribidae, the entire tentorial structure is lost and the gular sutures are usually reduced to little more than lateral irregularities on the margin of the foramen magnum. In Belidae there is variation from two widely separated gular sutures that end in two separate posterior tentorial pits (Fig. 3B) to convergence to a single median pit, with consequent reduction in size of the gula, to formation of a single median gular suture of variable length (Fig. 3C). The allies of Belidae (Aglycyderidae and Oxycorynidae including Rhopalotria ) share at least part of this same variability. The only other known members of the Curculionoidea that share in similar vari- ability are members of the neotropical scoly- tid genus Gnathotrupes . Of the 23 species of 1986 WOOD; Genera of Scolytidae Fig. 3. Diagrams of the ventral surface of the head of (A) Cimbcris attclaboides (Nemonychidae), (B) Belus sp. (BeHdae, from Austraha), and (C) an unidentified female Belidae from Australia. Note the convergent postgular sutures and remnants of pregular sutures from the level of the antennal insertion to (or toward) the anterior tentorial pits, gs = gular suture, ptp = posterior tentorial pit. Gnathotrupes examined for thi.s character by me, 5 had two complete gular sutures that extended to and were continuous with the pregular sutures, 8 had the pregula elongated and extending most of the distance to the postgula, and 10 had the pregula more or less normal. Most Platypodidae (Fig. 4) have an unusually large pregula and a short to very short median gular suture. Pregular sutures are totally absent in all curculionoids, except for Platypodidae (Fig. 4) and Scolytidae (Fig. 5-G) and for their par- tial presence in females of two (unidentified) Australian Belidae (Fig. 3C). In these two belids the pregular sutures extend only from near the anterior tentorial pits to near the point of antennal articulation; they are not represented from the area of antennal articu- lation to the anterior end of the median gular suture (the point marked externally where the tentorial apparatus invaginates). The pregular sutures illustrated by some writers on Cur- culionidae (Hopkins 1911:Fig. 1) do not exist; they represent irregular undulations in the cuticle that serve to strengthen the posterior (or ventral) wall of the rostrum and do not qualify as sutures in any acceptable usage of that term. Mouthparts have been used extensively in fundamental divisions of the Curculionoidea (Crowson 1967). For example, in Anthribidae and Nemonychidae the maxillary lacinea and Great Basin Naturalist Memoirs No. 10 Fig. 4. Posterior aspect ol head of a female ofDoliopijgus chapuisi (Duvivier) (Platypodidae). Note the very short median gular suture and the prominent pregular sutures. galea form separate elements (Ting 1936: Fig. 78). The only other curculionoids sharing this character are the Tesserocerini (Platypodidae) (Fig. 6), Attelabidae, and Rhynchitidae (Ting 1936:Fig. 78). Similarly, the Anthribidae and Nemonychidae have a 4-segmented maxillary palpus (Crowson 1967, Ting 1936). This char- acter is shared by all Attelabidae (Crowson 1967, Ting 1936), some Rhynchitidae, and one Platypodidae (Austroplatypus) (Browne 1971a). The costate lateral margins of the pronotum in primitive tribes of Scolytidae (Diamerini, Ctenophorini, Scolytini, etc.) is another indi- cation of primitive origin of this family within the Curculionoidea. The tibial structure of Platypodidae and Scolytidae is unique. It appears to have been derived from a structure resembling that of Protohylastes (Fig. 7), in which the median member of three apical spines became the terminal mucro of Platypodidae and the mesal spine became the terminal mucro in Scolyti- dae (Wood 1973a). The lateral margin is armed by one or more spines in primitive tribes; these may be replaced in specialized groups by socketed denticles (teeth) of setal origin (Wood 1978). The tibial structure of Scolytus and Camptocerus is a specialized de- viation from the basic scolytid structure that only superficially resembles the cossonine tibia. Similar structure occurs in Histeridae, Bostrichidae, Brenthidae, or other coleopter- ous groups having a long history of occupancy of insect tunnels bored in wood. The tibial spines of Araucarini (Cossoninae) that have been suggested as ancestral to those of Scolytidae (Kuschel 1966) also are of indepen- dent origin; in fact, the scolytid denticles to which they were compared are socketed and bear no structural similarity to them what- ever. The elytral locking mechanism is basically the same throughout the Curculionoidea, ex- cept that it is radically modified in Attelabidae and Rhynchitidae in an obvious specialization (Wood 1978, and unpublished drawings). A minor departure occurs in Scolytidae (Cor- thylini) (Wood 1978). Visible abdominal sterna 1 and 2 are en- tirely free in all primitive genera of Platypodi- dae but are weakly connate in the higher Platypodidae, all Scolytidae, and in most higher Curculionoidea. The larvae present an enigma. E,\cept in the most primitive genera, the frons and clypeus are usually fused in Platypodidae (Browne 1972), as in Nemonychidae (Crow- son 1967), thus making them easily distin- guishable from those of Scolytidae. However, characters have not yet been found that distin- guish some Scolytidae from some Curculion- idae (Viedma 1963). The difficulty may come more from reduction and simplification to ac- commodate small size than from real differ- ences. Browne (1972) reported urogomphi- like structures in two species of Platypodidae; if correct, this is the only known occurrence of these structures in the Curculionoidea. Discussion Scolytidae and Platypodidae as Families A position within the Curculionoidea is uni- versally accepted for Platypodidae and Scolytidae, except for Schedl (1939), who gave them superfamily status without explanation. For more than a century, it was traditional to list Platypodidae and Scolyltidae as separate families next to Curculionidae until Crowson (1967:164) combined both families with Cur- culionidae, primarily on the basis of the ab- sence of convenient larval characters that could separate them from that family. How- 1986 WOOD: Genera of Scolytidae Stenoscelis Fig. 5 A-F. Head structure of Curculionidae (Steno.icelis , Rhyncolus) and Scolytidae (Hijlurgops. Ips): Stenoscelis brevis (Boh.) (Curculionidae). A, lateral, B, dorsal, and C, caudal, with internal tentorial and associated structure indicated by broken lines; Hijlurgups rugipennis (Mannerheim), D, lateral, E, posterior, F, dorsal, internal structure as above. Great Basin Naturalist Memoirs No. 10 Fig. 5 G-I. Ips woodi Thatcher, G, ventral, H, lateral; Rhyncolus hnowltoni Thatcher (Curculionidae), I, ventral. Abbreviations; ai — antennal insertion; aps — apodeme formed internally by pregular sutnre; dat = dorsal arm of tentorium; dfm — dorsal margin of foramen magnum; e — cervical membrane attachment; gs = gular suture; mgs = internal apodeme formed bv the median gular suture; mtp — median tentorial pillar that invaginates from the combined posterior tentorial pits; p = pregula; pap = paired apodemal plates that branch dorsad from mgs and bears the tentorial apparatus at its anterior extremities; pg = postgula; ps = pregular sutures; ptb = posterior tentorial bridge; ptp = posterior tentorial pit. Fig. 6. Maxilla of Platypodidae: Periommatus bispinus Strohmeyer (left); Chaetastus tuherculahis (Chapuis) (center); and Tesserocerus insignis (Saunders) (right). Note the separate galea and lacinea. Re-drawn from Strohmever (1914; pi. 1). ever, following more than 30 years of study of the comparative anatomy of the CurcuHon- oidea, I find that position untenable and sug- gest that the relationship of Platypodidae and Scolytidae to Curculionidae is remote, at best, and warrants much closer examination. Platypodidae and Scolytidae universally share with one another; (1) well-developed pregular sutures that are reinforced internally by mas- sive apodemal inflections of the entire cuticle, 1986 WOOD; Genera of Scolytidae Mecopelmus Coptonotus Tricolus Scolytodes Diamerus Camptocerus Fig. 7. Posterior face of prothoracic tibia: Protohylastes annosus Wood (Coptonotinae), Protoplatypus vetulus Wood (Coptonotinae), Mecopelmus zeteki Blackman (Coptonotinae), Schedtarius mexicanus (Duges) (Coptonotinae), Cop- tonotus ctjclops Chapuis (Coptonotini), Tricolus peltatus Wood (Scolytinae, Corthylini), Scolytodes sp. (Scolytinae, Ctenophorini), Diainerus impar Chapuis (Hylesininae, Deamerini), Camptocerus auricomus Blandford (Scolytinae, Scolytini). Each drawing made at different scale so as to reproduce at a uniform size. extending from the posterior tentorial pit (at the anterior end of the median gular suture) to or near the anterior tentorial pit adjacent to the anterior articulation of the mandible (Figs. 4-5), and (2) a primitive mandible that is articulated and functions (about as in An- thribidae and Nemonychidae) in a manner entirely different from that of higher cur- culionoids (Fig. 8). In view of these universally present primi- tive characters, relict retention of other ances- tral features in one or more genera of these two families takes on added significance. For example: (1) The platypodid genus Austro- platypus clearly has a 4-segmented maxillary palpus (Browne 1971a) (known elsewhere only in Anthribidae, Nemonychidae, Attelabidae, and some Rhynchitidae). (2) The platypodid tribe Tesserocerini has the maxilla clearly di- vided into separate lacinear and galear ele- ments (Fig. 6); elsewhere in the Cur- culionoidea this character is shared by the same four families (cited here in No. I). (3) An adult labrum within the Curculionoidea oc- curs only in Anthribidae and Nemonychidae; however, an epistomal lobe resembling a fused labrum is widely represented among primitive Scolytidae (Fig. 1) and is usually present in Platypodidae (in Tesserocerus and Chaetastiis , Fig. 2, it is even separated from the epistoma by a suture). (4) A complete gula, with two gular sutures continuous with the pregular sutures, is present in at least five Gnathotriipes (Scolytidae) species (Wood 1973a), and the pregula is greatly prolonged in eight other species of this genus; the pregula is greatly enlarged and the median gular su- ture is short to very short in most Platypodi- dae (Fig. 4). (5) In most Platypodidae and many scolytid Hylesinini, Scolytini, Mi- 10 Belidae Great Basin Naturalist Memoirs No. 10 Brenthidae Sphenophormi \ Fig. 8. Lateral a.spect of ape.\ of rostrum ot a representative of: Belidae, Brenthidae, Sphenophorini (Curculionidae), Rhyncolini (Curcidionidae), Platypodidae, and Scolytidae. The arrows mark the approximate position of the concealed anterior and posterior articulations ol the mandible. cracini, etc., the scape is very short, httle if any longer than the pedicel and definitely not geniculate, and in other groups (some Phloeo- tribtis, some Micracini, etc.) the club is poorly formed; very little imagination is needed to see the possibility of independent origin of the geniculate, clubbed antenna from that of other curculionoids. (6) The costate lateral margins and concave pleura of scolyiid Diamerini, Ctenophorini, and some Scolytini occur elsewhere in the Curculion- oidea only among the most primitive families. (7) The platypodid-scolytid tibiae totally lack corbels. Their lateral margins bear true spines in Platypodidae and some primitive Scolyti- dae. These are replaced in all higher Scolyti- dae by socketed denticles of setal origin that occur nowhere else in the Curculionoidea (Wood 1978); true spines also occur in some Histeridae, Bostrichidae, and other families that frequent the tunnels of woodboring in- sects. (8) Although the literature states that visible abdominal sterna 1 and 2 are connate in all Platypodidae, this is not true. These segments are entirely free in all primitive gen- era; they are weakly connate in the higher Platypodidae and in all Scolytidae. An inde- pendent origin of this feature is probable. (9) The platypodid-scolytid body habitus most certainly is not of the Curculionidae type, although there is superficial resemblance to it in the Hylastini. Family Scolytidae In the early stages of this study, I was thor- oughly convinced that Platypodidae could be no more than a well-marked, primitive sub- family of Scolytidae. In view of the often in- finitesimally minute features used to charac- terize other femilies, that position must be reexamined. Although most characters are shared by some members of each group, a sharp demarkation remains. For example, the male spiculum gastrale is undeveloped in Platypodidae, but it is well-developed in Scolytidae (Wood 1982); tarsal segment 1 is greatly elongated in all Platypodidae (Fig. 9), except for Protoplatijpus and Scohjtotarsus , but it is intermediate in Protohylastes and Coptonotus (These four genera contain a total of five rare, tropical species). The shape and structure of the head, eye, antenna, prono- tum, scutellum, elytra, and tibiae are almost equally distinctive. Details of platypodid be- 1986 WOOD: Genera of Scolytidae 11 Fig. 9. Posterior aspect of protibia o{ Platypus parallelus (Fabricius) (Platypodidae). havior are too poorly known to add conclusive supporting evidence. The higher Platypodi- dae appear to have been the most primitive segment of this phyletic line to take up the woodboring ambrosial habit, which drasti- cally affected their morphology; this habit ap- pears to have arisen independently in at least eight tribes of Scolytidae (Hyorrhynchini, Phloeosinini, Scolytini, Scolytoplatypodini, Xyloterini, Xyleborini, Cryphalini, Corthy- hni) (Wood 1982). The Platypodidae universally lack socketed tibial denticles (derived from setae) (Figs. 7, 9); these structures are present in all higher groups of Scolytidae (Fig. 10) but are lacking in at least some genera of several primitive tribes. In all primitive genera of Platypodidae visible abdominal segments (sterna) 1 and 2 are free; in the higher Platypodidae and in all Scolytidae they are weakly connate. The pos- terior half of elytral interstriae 10 is univer- sally present in Platypodidae; it is present in primitive members of most tribes of Scolyti- dae (Fig. 12), but it is lost in the higher mem- bers of almost all tribes (Fig. 11). The apical protibial mucro of Platypodidae appears to have been derived from the middle apical spine of a tibia resembling that of Protohy- lastes (Wood 1973a); it appears to have been derived from the inner (mesal) spine in Scolytidae (Fig. 7). In larval Platypodidae (ex- cept Protoplatypus and Schedlarius) the clypeus is fused to the frons (Browne 1972); in Scolytidae it is a separate sclerite. The list could go on, but the above should indicate a close relationship between the two families and the limited overlap of many characters. Fig. 10. Lateral margin near apex of protibia of Pohjgra- phus rufipennis (Kirby). Note the socketed denticles. Subfamilies and Tribes A review of characters usable in classifica- tion of the higher categories of Scolytidae was presented by Wood (1978). Several trends were reported in that study: (1) the primitive head is more or less truncate on its posterior face, the dorsomedian area is progressively prolonged caudad in specialized groups (Figs. 13-14, and Wood 1982:Fig. 14); (2) the primi- tive frons is convex in both sexes (in the higher Hylesininae the male frons is variously im- pressed and the female frons is usually con- vex); the reverse is usually found in the Scolytinae; (3) the primitive eye is oval, en- tire, and finely faceted; specializations in- clude elongation, emargination (including complete division into two parts) (Fig. 15), and enlargement of facets (apparently corre- lated with nocturnal flight habits); (4) the an- tennal scape primitively may have been short, little longer than the pedicel; it is elongate or triangular in most groups; (5) the antennal funicle primitively contains seven segments, and there is a more or less orderly reduction to a minimum of one segment as specialization increases; (6) the antennal club varies from almost nonexistent (three movable segments no longer than those of the funicle in primitive Phloeotribus) to a simple cone-shaped struc- ture with transverse sutures to large and 12 Great Basin Naturalist Memoirs No, 10 Figs. 11-12. Lateral aspect of Scolytidae: 11, above, in- terstriae 10 unites with interstriae 9 before the level of the hind coxae in Eupagiocerus dentipes Blandford;- 12, be- low, interstriae 10 continues to near apex of the elytra in Scolytodes pluineriae Wood. Strongly flattened or obliquely truncate, with or without sutures; it is probably the most variable major structure found throughout the family; (7) the prothorax may have (a) the coxae widely separated in primitive groups to fully contiguous in specialized ones; (b) the pleuron concave, with the lateral margins acutely costate in primitive genera (Figs. 11- 12), to convex, with the lateral margin un- marked in specialized genera; (c) the prono- tum longitudinally straight in dorsal profile and unarmed by crenulations in primitive groups to strongly arched and armed by crenulations or asperities in specialized ones; Figs. 13-14. Lateral aspect of head in Scolytidae: 13, Hylastes nigrinus Mannerheim, with the posterior face approximately truncate; 14, Xylosandnis retusus (Eich- hoff), with the occipital area greatly extended. (8) the basal margins of the elytra tend to be costate primitively, with the Hylesininae be- coming procurved and crenulate (Fig. 16), the Scolytinae transversely straight and longitudi- nally rounded (Fig. 17); (9) several complex changes occur in the meso- and metathorax that will be discussed below; (10) the male tergum 8 is visible and pubescent in most groups (Fig. 22), but it is telescoped beneath 7 and without pubescence in Carphodicticini, Ipini, Dryocoetini, Xyloterini, and Xylebor- ini; (11) the venter of the abdomen has (a) segments 3 and 4 (visible segments 1 and 2) free in all primitive Platypodidae but weakly connate in the higher Platypodidae and all Scolytidae and (b) limited specialized groups in which specialized features are very impor- 1986 WOOD: Genera of Scolytidae 13 O Fig. 15. Variations in eye shape (left to right): shallowly sinuate {Hijlesinus crenatus Fabricius); narrowly and shallowly emarginate (Ernoporicus cmicasicus Lindemann); strongly sinuate or broadly, shallowly emarginate {Phloeosinus bicolor BruUe); deeply emarginate (P/i/oco.sinu.s thujae Ferris): conipleteh', equally divided {Pohjgraphus pohgraphus Linnaeus); and completely, unequally divided (Sphaerotnjpcs globosiis Blandford). Figs. 16-21. Thoracic structure of Scolytidae: 16, elevated crenulations (arrow) on procurved basal margins of elytra of Phloeotrihus setulo.'ius Eichhoff; 17, weakly subcostate basal margins of elytra in Scolytodes plumeriae Wood; 18, metatergum oiHylastes nigrinus Mannerheim, arrows mark intersegmental line (left) and scutoscutellar suture (right); 19, metapleuron ofHylastcs nigrinus Mannerheim, upper arrow marks pleural suture, lower arrow marks anterior end of groove that receives costal margin of elytron; 20, ventrolateral aspect of prothorax, arrow marks acutely elevated precoxal ridge (or costa); 21, metatergum of Eupagiocerufi dentipes Blandford, pn = postnotum, is = fused interseg- mental line, ss = scutoscutellar suture (compare to Fig. 18). tant (Scohjtua, Scolytomimus , etc.); and (12) the tibiae are unusually variable and require special treatment below. Most of the above character states are utilized in the following key to subfamilies and tribes to indicate phyletic trends in the family, subfamily or tribe in which they are involved. However, two of them are sufficiently complicated and im- portant that elaboration is appropriate. These occur on the metathorax and the tibiae. Three primary characters of the metathorax show important features that exhibit phyletic 14 GreatBasin Naturalist Memoirs No. 10 male femole Fig. 22. Abdominal terga of Cryphohis nificollis Hop- kins: male, with segments 7 and 8 visible; female, with segment 7 visible (segment 8 is reduced in size, tele- scoped beneath 7, and hidden from view). trends. First, in Curculionoidea generally, in- cluding Platypodidae and primitive Scolyti- dae, the pleural suture follows a zigzag course from the pleural wing process ventrad to the point where the costal margin of the elytron touches the body. It then turns abruptly cau- dad to a point just before reaching the poste- rior limits of the segment, where it turns mesad and continues to the pleural coxal pro- cess. On the metepisternum at the anterior or first angle (below the pleural wing process) a carina or small, flattened spine is present that fits into a small groove on the inner (costal) margin of the elytron just behind the humeral angle. This character state within Scolytidae is best seen in the Hylastini (Fig. 23, parts 43- 44) and in some Hylesinini. The trend is for the suture to progressively straighten out whereas the metepisternal spine remains sta- tionary but becomes more remote in position from the changing suture (Fig. 23, part 42). The matching groove on the elytron moves somewhat dorsad and caudad, suggesting that the costal margin of the elytron extends far- ther ventrad than primitively. In Cryphalini the metepisternal spine is considerably re- duced in size, and its function in locking the elytra in closed position is partly assumed by a new diagonal groove on the metepisternum (Fig. 31). In Corthyulini, the spine is entirely lost and the groove is enlarged and extends to a more ventral position (Figs. 23, part 47, and 32). The result is that the closed elytron in this tribe now covers all but a small anterior por- tion of the metepisternum. A second significant feature of the metathorax involves progressive changes in position of the scutoscutellar suture. In Curculionoidea gener- ally, Platypodidae, and primitive Scolytidae (Hylastini and Hylesinini are examples), this su- ture reaches the margin of the scutellar groove near the anterior limits of the groove and contin- ues parallel to and very near its lateral crest for about two-thirds of the length of the groove, then the suture curves abruptly laterad to follow its usual course to the posterior margin of the segment (Fig. 23, part 43). In more advanced tribes this suture progressively straightens out, meeting the marginal crest of the groove only briefly, if at all (Fig. 23, part 41). The third significant metathoracic feature that is usable in phylogeny occurs only in the Hylesininae. A significant feature of insect flight is the intersegmental line between the metathorax and its postnotum (derived primi- tively from the anterior portion of the first abdominal segment but functionally part of the thorax), which must flex with each stroke of the metathoracic wings. This suture is present in all insects, including Platypodidae, Scolytinae, and primitive Hylesininae (Fig. 23, parts 43, 45). In the more advanced Hylesininae, the median two-thirds of this intersegmental line is lost by complete fusion of the postnotum to the metathorax (Fig. 23, part 41). This fusion is progressive, leaving a weak suture primitively, but it is totally oblit- erated in advanced groups of this subfamily. The tibiae exhibit remarkable variation in constant, conservative patterns that are valu- 1986 WOOD; Genera of Scolytidae 15 Fig. 23. Diagrams of terga and pleura of Scolytidae: 41, metatergiim and 42, pleuron ofCbramcsus hickoriae LeConte, arrow points to remnant of intersegmental suture in 41, to metepisternal spine in 42; 43, metatergum and 44, pleuron of Hylastes nigrinus Mannerheim, arrows as above; 45, metatergum and 46, pleuron o{ Cnemonyx panamertnis (Bland- ford), arrows as above; 47, pleuron of Pttyophthorus crotonis Wood, metepisternal spine is lost and is replaced by a small groove (arrow). Abbreviations: is = intersegmental suture, mn = metepimeron, ms = metepisternum, pn = postnotum, ps = pleural suture, s = metepisternal spine (part of locking mechanism for elytra), sg = scutellar groove, ss = scutoscutellar suture. able in following trends in phylogeny. Appar- ently, the primitive model from which the tibiae of Platypodidae and Scolytidae were derived resembles that of Frotohijlastes (Fig. 7). The protibia of Protohylastes is slender, with three rather small spines at the apex and 16 Great Basin Naturalist Memoirs No. 10 a very minute spine on the posterior face im- mediately basad from the tarsal insertion. Comparative studies (Wood 1973, 1978) sug- gest that the mesal (inner) apical spine lie- came the apical spine in Scolytidae and that the middle spine became the apical spine in Platypodidae (Figs. 7, 9). This middle spine in Scolytidae is bent laterad and is a major, iden- tifiable element that projects beyond the level of the tarsal insertion in one or more members of several tribes (Hyorrhynchini, Diamerini, Bothrosternini, Phloeotribini, Phloeosinini, Hypoborini, Polygraphini, Scolytini, Cten- ophorini, Scolytoplatypodini). All tibial spines in Platypodidae (Fig. 9), Hyorrhynchini, Scolytini (Fig. 7), and, apparently, Cten- ophorini (Fig. 7) are true spines. In all higher Scolytidae these spines are either intermi.xed with or replaced by socketed denticles of setal origin (Fig. 10). The patterns of possession and replacement within the family suggest that socketed denticles had an early mono- phyletic origin within the Scolytidae, with dif- ferential survival of this character in the pos- terity, and do not suggest polyphyletic origins. Many other characters that are not men- tioned above are useful in classification and phylogeny in more limited groups. Several of these are employed in the keys. Conclusions As stated above, the traditional place of Scolytidae in classification has been as a fomily next to the Curculionidae. However, as Cur- culionidae became fragmented into several families, the position of Scolytidae within the Curculionoidea changed. Schedl (1939c) made a separate superfamily to contain Platy- podidae and Scolytidae. Crowson (1967:164) reduced both to the rank of subfamily within Curculionidae. In reviewing this topic, our first point has been to establish that Platypodidae and Scolytidae are closely related to each other and are derived from the same parental stock. Evidence for this, as stated above, comes from the following: (1) Pregular sutures that extend from the anterior tentorial pit (adjacent to the anterior articulation of the mandible) to meet the anterior end of the median gular sutiue occur only in these two groups. These sutures are reinforced internally by massive apode- mes. Remnants of pregular sutures found in females of two species of Belidae extend from the anterior tentorial pit toward the antennal insertion and never converge or approach the gular suture (or sutures). (2) In these two families, the mandibles come to an apical point and have the cutting edge mesal as in Nemonychidae; furthermore, the articulating condyles are comparatively close together, with the posterior one more nearly lateral in position, and the hypostomal area behind the posterior condyle is minimal in size to almost nonexistent. In most Curculionoidea, except for these two families and Nemonychidae and Anthribidae, the basal half of the mandible forms a horizontal cylinder that rotates on the two condyles, and the cutting edge of the mandible is apical; the posterior condyle is posterior in position and is supported by an enlarged hypostomal area. (3) The subcortical habit in which mated pairs enter vital host tissues to oviposit is shared by Platypodidae, Scolytidae, and some Cossoninae (Curculion- idae); however, the mode of excavation and several structural features (gular area, tento- rium, mandible, etc.) indicate that the latter group exhibits an evolutionary parallelism, not a fundamental relationship. (4) Six genera {Protohylastes, Coptonotus, Scolytotarsus, Protoplafyptis, Mecopelmus, Schedlarius) are structurally and biologically intermediate be- tween the Platypodidae and Scolytidae and could be placed in either family. The charac- ters, reviewed in this paragraph, with primary stress placed upon true pregular sutures, indi- cate a fundamental close relationship between Platypodidae and Scolytidae that is not shared with other groups. A second point of significance has to do with the position of Platypodidae and Scolytidae within the Curculionoidea. Their traditional position in classification has been next to the Cvnculionidae; however, close examination suggests that such a placement is based on superficial resemblance, not on fundamental structure. As indicated above, pregular su- tures are shared, in part, only with females of two species of Belidae, and the mandibular- hypostomal structure is shared only with An- thribidae and Nemonychidae. These indica- tions suggest a relationship of Platypodidae and Scolytidae to the segment of Curculion- oidea having two gular sutures, rather than 1986 WooD: Genera of Scolytidae 17 one. However, close inspection of Belidae discloses at least three genera with only one median gnlar suture and an elongated post- gula, and an entire segment of Oxycorynidae (Rhopalotria) with one gular suture and a postgula of variable size. Furthermore, one genus of Scolytidae {Gnathot rapes) contains at least five species with a complete gula con- necting pregular and postgular elements, and at least eight more in which the connection is almost complete. In many Platypodidae, the pregula is enlarged and is scarcely separated from the postgula. Because such a situation does not occur elsewhere among those cur- culionoids having one gular suture, wisdom suggests that a search be continued for addi- tional clues to phyletic relationships. A movable labrum occurs among cur- culionoids only in Anthribidae and Nemonychi- dae. Remnants of a labrum are found in many primitive Scolytidae (particularly Hylesininae), and a fused labrum, complete with a transverse suture, occurs in Platypodidae (especially Tesse- rocerini). 1 am unaware of similar remnants in other curculionoid families. A 4-segmented ma.xillary palpus occurs in Aiistraloplatypiis (Browne 1971a:49), a fea- ture restricted to only the most primitive cur- culionoids (see family key below). A lacinia separate from the galea is charac- teristic of all Tesserocerini (Platypodidae); this is another feature that is restricted to the most primitive curculionoids (see family key below). The antenna has been used extensively in the classification of curculionoid families, par- ticularly character states of orthocerous vs. geniculate, apex filiform vs. clubbed (or capi- tate), etc. In no curculionoid are segments 1 and 2 without some degree of enlargement; furthermore, in many of those traditionally referred to as geniculate, the scape is not longer than it is in some Anthribidae. In a large segment of Platypodidae and many Scolytidae, the scape is short, little if any longer than the pedicel, and often it is broadly triangular in shape. In virtually all cur- culionoids (except most Anthribidae), the three apical segments of the antenna are at least slightly enlarged; although the literature generally states otherwise, a few Anthribidae have an abruptly enlarged club. In the genus Phloeotrihus (Scolytidae), the club is usually sublamellate, with the three segments freely movable upon one another (Fig. 16). In primi- tive members of this genus iacaciae , rhodo- dacUjlus , etc.) the three segments are no longer or wider than are the funicular seg- ments and are as movable. I see no possibility that the Phloeotrihus antenna could have been derived from a solid club of the type found in most other Scolytidae. Since this is the only deviant character found in this genus, could it not be a relict reminder that the scolytid club was derived on an independent, parallel line from a common ancestor, and not Irom other curculionoid groups having clubbed antennae? Much has been said and written of the simi- larity of the tibiae between Scolytiis (Scolyti- dae) and many Cossoninae (Curculionidae) in curculionoid family classifications. However, it has never been pointed out that similar tibiae occur in Histeridae, Brenthidae, and other coleopterous families containing groups with a long history of subcortical existence. Within the Scolytidae, the Scolytus tibia is unicjue and apparently does not represent a truly primitive condition. The truly primitive character state of the tibia on this phyletic line occurs in Protohijlastes, with Platypodidae and Scolytidae derived along separate phylet- ic lines from the basic structure. Larval characters, thus far, have not been very helpful in answering questions about scolytid phylogeny. In fact, reliable means have not yet been found for separating all Scolytidae from all Curculionidae (Viedma 1963, Lekander 1968). Whether this is due to the absence of characters or to the lack of diligence in the search for characters among groups not found in Europe needs to be an- swered. Primitive larval Platypodidae have the frontal sutures extending to the articulat- ing membrane of the mandibles, a fact not previously noted. Browne (1972) suggested that structures found on two platypodid larvae could be relicts of urogomphi. If so, this is the only evidence of the existence of urogomphi in the Curculionoidea. Larval Scolytidae and Platypodidae are easily separated from each another by the separate (Scolytidae) or fused clypeus and frons (Platypodidae). It is of inter- est that this same character separates larval Nemonychidae and Anthribidae (see family kev below). 18 Great Basin Naturalist Memoirs No. 10 Lower Jurassic curculionoids with a long rostrum have been referred to the CurcuHon- idae by Crowson (1983), although they bear a very strong resemblance to Belidae or, per- haps, Eobelidae. Arnoldi (1977) named the extinct family Eobelidae from 14 species that were placed in 7 tribes in 4 subfamilies, taken from Upper Jurassic deposits of South Kaza- khstan (Karastan) in the USSR. In this family the head is similar to modern Belidae except that it is more broadly oval and the rostrum is shorter, as in some modern male Brenthidae. The mandibles are large, with a single apical point and a mesal cutting edge as in modern Nemonychidae. The antennae attach either to the middle or near the apex of the rostrum and are orthocerous, with the three apical seg- ments slightly enlarged but not forming a defi- nite club. Arnoldi did not mention gular su- tures or a labrum. The body resembled modern Belidae except that the lateral mar- gins of the prothorax were acutely elevated as in modern Oxycorinidae and the elytra were much less strongly sclerotized than in modern Belidae. From the above, two fundamental conclu- sions emerge. First, the Platypodidae and Scolytidae are very closely related to one an- other and, in fact, they intergrade to a limited extent. At the same time, these two families are quite distantly related to other families in this superfamily. Second, in spite of the ap- parent absence of larval characters, there are enough discordant adult characters to conclu- sively demonstrate that the Platypodidae- Scolytidae, as one unit, do not belong to and probably are not even closely allied to Cur- culionidae. It is my contention that their preg- ular sutures (and accompanying internal apodemes) prevented elongation of the ros- trum and forced them into an entirely differ- ent mode of existence from other cur- culionoids (parent adults had to bore into vital host tissues to oviposit because they had no rostrum and, consequently, could not follow the curculionid habit of using the rostrum to form ovipositiou pits). They could have been derived in the Jurassic from a short-rostrumed Eobelidae-like ancestor that had fully formed pregular sutures. Belidae and related families, and all other curculionoids having one gular suture, branched off later or from other segments of Eobelidae or its deriva- tives. Rostrum length among Jurassic cur- culionoids was diverse, varying from short and broad (Arnoldi 1977) to long and slender (Crowson 1983). The presence of obvious, so- phisticated scolytid tunnels in Lower and Middle Cretaceous conifer bark suggests that the Platypodidae-Scolytidae phyletic line had its origin at least in Upper or Middle Jurassic. The exact phyletic positions of known Jurassic and Cretaceous "curculionid" fossils are diffi- cult to determine, because details of labrum, mandibles, gular sutures, etc., are not suffi- ciently well preserved to permit analysis. I seriously doubt that any true Curculionidae existed prior to the Cretaceous. Habits and Classification Although the Platypodidae are established above as a family, separate from Scolytidae, they are significant because they represent the first major paraphyletic branch from the platypodid-scolytid line of specialization. Their adoption of the xylomycetophagous habit appears to have accelerated their struc- tural and biological deviation. Only the inter- mediate genera Protoplati/pus, Mecopelmus , and Schedlarins lack the ambrosial habit {Pro- tohylastes, Scohjtotarsiis , and Coptonotus are unknown biologically). Schedlarius (mon- ogamous) and Protoplatypus (polygynous) place their eggs in sealed niches; all other known platypodids are monogamous and dis- tribute the eggs loosely in the parental gal- leries or carry them on the female frons. Mecopelmus makes a cave-type nuptial cham- ber (without ovipositiou galleries) in the cam- bium region of its host and deposits the eggs in clusters therein, and the larvae form indepen- dent mines in the cambium at least for the latter part of their development. Protoplaty- pus forms stellate or radiate tunnels, with a central nuptial chamber in the cambium re- gion, and the eggs are placed in regular, alter- nate, sealed niches along the linear egg gal- leries; the larvae form independent mines throughout their development. Schedlarius is xylophagous (Wood 1957a), places its eggs in randomly organized niches that are sealed with frass, and the larvae form long, indepen- dent mines that wander aimlessly through the wood. An obvious association with fungi is apparent in the vicinity of adult and larval activity, although no mycelial growth in the 1986 WOOD: Genera of Scolytidae 19 mines could be seen at magnification of 20X. The habits mentioned in this paragraph also form the basis of habits for the more diverse Scolytidae. In my review of characters usable in distin- guishing subfamilies of Scolytidae (Wood 1978), only one significant character gap was found that was supported by numerous fea- tures. These characters are summarized in the key to tribes below. The presence or absence of pronotal asperities, the ambrosial habit, and several features used traditionally by past workers to separate multiple subfamilies of Scolytidae either have no value in characteriz- ing subfamilies or have no value in phylogeny . For these reasons, only two subfamilies were recognized. Because the Hylesininae are structurally and biologically less diverse and their specializations more conservative, they are treated here as more primitive than the Scolytinae. Tribes of Hylesininae In general, the Hylesininae are phloeo- phagous. Exceptions include the twig- or small-branch (pith)-boring mycetophagous Hyorrhynchini (all three genera), Bothroster- nus (Bothrosternini), and Hylcops (Phloeo- sinini; last half of larval stage only); xyloph- agous Dendrosimis and three known species of the large genus Chramestis (Phloeosinini); the myelophagous Cnesiniis, Eiipagiocerus , and Sternobothrus (Bothrosternini); and the spermophagous Pagiocerus (Bothrosternini) and one species of Phloeotrilnis (Phloeotrib- ini). The usual mating system in this subfamily is monogamy, with the female initiating the new parental gallery. Departures from this system include (a) apparent male haploidy (arrhenotocous parthenogenesis accompanied by consanguineous polygyny) in Sueus (Hyor- rhynchini); (b) monogamy, with male initia- tion of parental galleries in some Phloeosinus and Olonthogaster (Phloeosinini); and (c) het- erosanguineous (superficially resembling harem) polygyny in at least two species of New Guinea Olonthogaster (Phloeosinini), and in all Carphoborus and many Polygraphus (Polygraphini). These basic habits appear to be correlated with (1) a conservative evolutionary departure from the ancestral structure; (2) a more com- pact (stout) body; (3) the body compaction that appears to have resulted in the development of a row of coarse crenulations on the basal margins of the elytra (their function is evi- dently to add resistance when a predator at- tempts to pull from a timnel the beetle that is blocking the entrance), and (4) the straighten- ing of the pleural suture. The compaction of the body apparently also had an effect upon the mechanism of flight and permitted fusion of postnotum 2 to the metathorax. Although use of these characters recjuires further study and refinement, a definite trend from the Hy- lastini-Hylesinini to Phloeosinini is indicated. There appears to be a gradual transition in characters from the Hylastini to Hylesinini to Tomicini. The Hyorrhynchini and Phrixoso- mini appear to be relicts of an earlier radiation of primitive stock and are quite unrelated to the other three tribes. The remaining hylesinine tribes have the postnotum partly to entirely fused to the metanotum and fall into three units. Tlie first includes the Old World Diamerini and New World Bothrosternini tiiat probably contain the most primitive spe- cies of this subfamily. Their protibial, prono- tal, and base of elytral structure is much more primitive than the Hylastini-Hylesinini. Their antennal structure is much more similar to Coptonotiis-Protohylastes (Platypodidae), although the Hylastini-Hylesinini club is probably more similar to the ancestral struc- ture. The second tribal unit within the more highly evolved segment of this subfamily in- cludes the Phloeotribini and Phloeosinini. Tibial structure of at least a few representa- tives of each suggests an ancient origin. Ex- cept for the aberrant structure of the antennal club of Phloeotribini, they probably could not be separated from one another at the tribal level. They include species bearing the great- est departures from the primitive forms in the pleural and scutoscutellar suture patterns. The absence of socketed protibial teeth in Aricerus and the movable segments in the antennal club suggest that Phloeotrinini is the more primitive of these two tribes. The Polygraphini and Hypoborini are obvi- ously specialized ancient groups that are un- related to one another but of uncertain affinity to other groups. The superficial appearance of the body form of Phloeographtis (Poly- 20 Great Basin Naturalist Memoirs No. 10 graphini) suggests a relationship to Tomicus , but the protibial structure of Serrastiis sug- gests a much more ancient origin of this group. The Hypoborini could be aspecialized paraphyletic line that branched from the Phloeosinini. Tribes of Scolytinae The tribes of Scolytinae are easily clustered into five major groups on the basis of several anatomical characters. The most primitive, composed of Scolytini, Ctenophorini, and Scolytoplatypodini, share primitive protibial structure without socketed denticles, a 6- or 7-segmented antennal funicle, a head without a caudal extension of the dorsal occipital area, rather widely separated procoxae, etc. This group contains members (Cnemomjx, Scoly- todes) with the greatest structural similarity of all Scolytinae to primitive Platypodidae (Pro- toplatypus , in this instance) and to Hylesini- nae. In fact, some Cnemomjx (galeritus and its allies) have coarse crenulations on the basal margins of the elytra; also, Protohtjiastes was thought to be allied to Pseudohylesinus (Tomicini) until the legs and antennae were studied. The Ctenophorini habits include phloeophagy, xylophagi, and myelophagy; monogamy and polygyny (heterosanguineous only); egg chambers of the cave, linear and radiate types; and deposition of eggs in loose clusters in parental chambers or in individual niches sealed by frass. The monogamous, xy- lomycetophagous Scolytoplatypodini appear to be a specialized Old World geographical replacement of the Ctenophorini. The Scoly- tini are monogamous (except for a few bigy- nous Scohjtus) and phloeophagous (except for the xylomycetophagous Camptocerus). Cne- momjx (Scolytini) and Scohjtodes (Ctenoph- orini) could have been derived from a com- mon ancestor. Pycnarthnim (Ctenophorini) could easily be placed as a primitive tribe of Hylesininae on both anatomical and biological bases. The second cluster of tribes within the Scolytinae includes the Micracini and Cac- topinini. Advances include a more efficient body form (cylindrical compaction), slightly extended dorsal occipital area of the head, reduction of funicular segments to six (rarely five), universal occurrence of pronotal asperi- ties, common occurrence of xylophagy and bigyny, etc. Xylomycetophagy and males nor- mally associated with more than two females are unknown. Scalelike pronotal and elytral setae are a common occurrence. The evolu- tionary connection between this group of tribes and the more primitive Scolytini- Ctenophorini-Scolytoplatypodini is remote; however, a connection to the more advanced Xyloctonini-Cryphalini is clearly evident. The third cluster of tribes within the Scolytinae includes the Carphodicticini, Ip- ini, Dryocoetini, Crypturgini, Xyloterini, and Xyleborini. In this group, male abdominal tergum 8 is reduced in size and telescoped beneath 7, as in the female. In addition, the procoxae are almost always contiguous, the occipital area of the head attains its greatest prolongation, the obliquely truncate antennal club is developed and exploited, both het- erosanguineous and consanguineous polyg- yny are common, and xylomycetophagy is universal in two tribes. Carphodicticini and some Dryocoetini are obviously very primi- tive, but their connection to other primitive Scolytinae is remote and not reflected in the modern known fauna. The basal margins of the elytra are elevated and costate in Dendro- dicticus (Carphodicticini) as in some Hylesini- nae. The fourth cluster of tribes within the Scolytinae includes Xyloctonini and Cryphalini. In these tribes the procoxae are entirely contiguous, and the antennal club is flat, with the sutures on the posterior face moderately displaced toward the apex. The small metepisternal spine that functions in locking the elytra in the closed position is reduced and partly replaced by a small groove (Fig. 31). Segmentation of the antennal funi- cle is reduced, ranging from a maximum of seven to a minimum of three. Monogamy is the general habit, with consanguineous polyg- yny universal in four genera of Cryphalini {Cnjptocaremis, Hypothenemits, Trischidias, Periocnjphalus) . There appears to be a close connection between Micracini and this group of tribes. The elytral locking mechanism also suggests a remote connection between this group and the Corthylini (Figs. 23, 31-32). The final cluster of tribes includes the Cor- thylini. The flattened antennal club, with the sutures equal on both sides, the unique elytral locking mechanism, tibiae, eyes, elytra, etc.. 1986 WOOD: Genera of Scolytidae 21 characterize this unique tribe. Pliloeophagy is almost universal in the primitive half of tiie tribe, xylomycetophagy in the advanced half Both monogamy and heterosanguineous poly- gyny are common. Consanguineous polygyny apparently occurs in three species oiArapttis . There may be a remote phylogenetic connec- tion to the Cryphalini. Geographical Origin of Tribes For a discussion on this topic to have any semblance of objectivity, certain premises must be established. First, it is assumed that continental drift did occur and, due to that phenomenon. South America and Africa were either connected at their southern ends or were at least close enough to permit faunal exchange by island-hopping until earliest Ter- tiary. At that time, Australia was close enough to the southern end of South America to per- mit at least limited faunal exchange. Second, it is assumed that South America was an island during most of the Tertiary. In addition to the assumed connection to Africa in earliest Ter- tiary, South America was connected to North America in early Tertiary and again in late Tertiary either by a land bridge, as presently exists, or by a series of islands that were close enough to permit faunal exchange. Third, it is assumed that faunal exchange between Africa, Europe, and Asia has been no problem except as it has been affected by climate. Fourth, it is assumed that climate has fluctuated as evi- denced by fossil remains of magnolia trees in Alaska and of tropical forests where the Sahara Desert is now located. Finally, it is assumed that scolytid beetles have had habits through- out their history that were essentially as seen at present. That is, groups presently re- stricted to the tropical forests have always been so restricted, and groups now in more temperate climates have been able to occupy cooler climates for a long time. In the first column of Table I are listed all tribes of Scolytidae presently recognized in the world fauna. From each tribe, those gen- era were selected that now are restricted to tropical climates. If those genera are now rep- resented in both Africa and South America or if they have a very closely related geographi- cal replacement genus on both continents, but are not found outside the tropics else- where, then those tribes were considered to have a pre-Tertiary origin (Table 1, column 2). If the tribe at that time was local in distribu- tion and was represented on only one land mass, it was not included as pre-Tertiary even though it could have been present. A faunal connection from South America to Australia was regarded as early Tertiary, not just pre- Tertiary. It is noted (1) that 6 of 11 tribes of Hylesininae and 7 of 14 tribes of Scolytinae are judged on this basis to be of pre-Tertiary origin, and (2) that the only pre-Tertiary group containing species with the xylomyce- tophagous haliit is Xyleborini. (Hylesinini is excluded, because its only xylomycetopha- gous genus, Hyleops , has not yet fully adopted that habit.) This does not mean that other xylomycetophagous groups were not present, but only that, if they were present, they were local in distribution (on only one land mass) at that time. It is further noted (3) that all tribes judged as pre-Tertiai-y were phloeophagous except for Xyleborini and part of Micracini. This suggests that xylophagy and xylomycetophagy are of comparatively recent origin. The significant point is that the family Scolytidae was a well-established, diversified group by the beginning of the Tertiary. The fossil engravings of scolytids in coniferous bark from Upper Cretaceous illustrated by Brongniart (1877) and described from Lower Cretaceous by Blair (194.3) are evidence of a much longer family history of the group than is usually recognized. It is probable that the beetles that made those engravings were much more similar to the Ctenophorini or, possibly, Protoplatijpiis than to modern Hylesininae. Phylogeny As stated above, the Platypodidae-Scolyti- dae group are members of the Cur- culionoidea. Within that superfamily, they form a distinctive unit that is characterized by the presence of fully developed pregular su- tures. Because the only other known pregular sutures within the Curculionoidea are the api- cal remnants found in females of two species of Belidae, the possibility of an ancient common ancestor should be examined. Three genera of Belidae and an entire segment of Oxyco- rynidae (Rhopalotha), a supposed derivitive 22 Great Basin Naturalist Memoirs No. 10 Table 1. Estimated age of the tribes of Scolytidae, based on genera known to be restricted to tropical areas. Tribes are considered pre-Tertiary if a tropical genus or a close geographical replacement genus occurs in Africa and also in South America (Column 3). More recent postseparation (of Africa and South America) distribution is added as indicated (Column 4). Ancient Distribution added after Tribe Pre-Tertiary' origin early Tertiary Hylastini Holarctic No change Hylesinini + Old World South America (from Africa), North America (from Asia) Tomicini a. Xylechimis + South America Worldwide allies b. Dendroctonus + Worldwide in Worldwide allies Araucaria Prixosomini + Africa-South America No change Hyorrhynchini Oriental No change Diamerini Africa Southern Asia to Australia Bothrostornini South America North America Phloeotribini South America Australia (from South America); North America to Eurasia, from South America. Phloeosinini + Africa-South America Ancient to Eurasia; modern to North America Hypoborini + Worldwide (or Africa- Relicts South America) Polygraphini Africa Europe and Asia, then to North America Scolytini South America North America then to Eurasia Ctenophorini + South America North America Scolytoplatypodini Africa Southern Asia Micracini + South America- -Africa North America to Asia Cactopinini North America No change Carphodicticini + ? South America-Southern Asia Ipini + South America- -Africa Worldwide Dryocoetini + Africa Worldwide Crypturgini Africa North America, Eurasia Xyloterini Asia North America Xyleborini + Africa-South Ai :nerica Worldwide Xyloctonini Africa Asia Cryphalini + Africa-South Ai nerica Worldwide Corthylini South America (African segmer It) Worldwide Marked ( + ) if present in recogniziiljle form prior to the beginni Tertiary by a local representative or a precursor that was presei ig of the Tertiary' on more than one land mass. Marked (0) if represented at the beginning of the It on only one land mass of Belidae, have only one gular suture, not two as in other members of the Behdae- Oxycorynidae-Aglycyderidae (Proterhinidae) group. Because such a placeinent would be a radical departure from the traditional position of Platypodidae-Scolytidae within the Cur- culionoidea (which has been next to the Cur- culionidae), additional supporting evidence was sought. The following summary of infor- mation presented in the above sections was found. A functional, but degenerate, adult labrum occurs in Curculionoidea only in An- thribidae and Nemonychidae. Rudiinents of a labrum are well marked in many Platypodidae and obscurely indicated in some primitive Scolytidae, but they are not indicated in other curculionoids known to me. Anthribidae and Nemonychidae have a maxillary lacinia sepa- rate from the galea; this character is shared by Platypodidae (Tesserocerini), Attelabidae, and Rliynchitidae; the lacinia and galea are fused in all other curculionoids. Most of those curculionoids having two gular sutures have a 4-segmented maxillary palpus; Attelabidae, some Rliynchitidae, and one Platypodidae share that character. In Anthribidae, Nemonychidae, Eobelidae (fossil), Scolyti- dae, and Platypodidae the mandible is api- cally pointed, the cutting edge mesal, the condyles are closer together and differently positioned, and the apical part of the hypos- toma is of minimal size. In all other cur- 1986 WOOD: Genera of Scolytidae 23 culionoids, the basal half of the mandible forms a cylinder that rotates on a horizontal axis between the condyles, the dentate cut- ting edge is distal and moves through a differ- ent arc, and the hypostoma is enlarged to give greater strength to the posterior condyle. Whereas the preponderance of adult charac- ters support the suggested relationship, larval characters have not been found that separate some Curculionidae fi'om some Scolytidae. The larvae of Platypodidae and Scolytidae are easily separated; however, it is by the same primary character that Anthribidae and Nemonychidae larvae are separated (see key to families below). Obviously the Platypodi- dae-Scolytidae are not part of the Curculion- idae and occupy a more primitive position in phylogeny. E.xactly where they branch from the main curculionoid stem is not yet clear, but it could be from an Eobelidae-like ances- tor. Within the Platypodidae-Scolytidae, three major phyletic units are evident. The smallest and most distinctive paraphyletic line is the Platypodidae (about 1,000 species). They lack socketed tibial denticles, have the terminal, protibial mucro formed from the middle one of three apical spines (as compared to Proto- hylastes), always have the posterior half of elytral interstriae 10 present, always have a primitive mechanism for locking the elytra in closed position, lack a spiculum gastrale, and contain members having the primitive charac- ters cited in the above paragraph. The Scolyti- dae contain partial or complete departures from these characters. As indicated above, si.x rare tropical genera (containing a total of seven species) are intermediate between these families. The other two major phyletic lines include the Hylesininae (about 2,000 species) and the Scolytinae (about 4,000 species). Although the most primitive members of the family un- doubtedly are members of the Scolytinae, that subfamily also contains far more and greater specializations than do the Hylesini- nae. Because the Hylesininae are less diverse and depart less drastically from the primitive form, they are considered more primitive and paraphyletic; the Scolytinae are more special- ized and contain the major evolutionary thrust of the family. Hylesinine evolution appears to have been oriented toward a bodytype that is subspherical to take advantage of the row of crenulations on the bases of the elytra; scolytine evolved toward cylindrical com- paction that is more effective in deeper host tissues where the ambrosial habit was ex- ploited. Progess toward a more nearly spherical body form in Hylesininae was accompanied by straightening out (somewhat) the scu- toscutellar and pleural sutures on the meta- thorax. This apparently modified the mecha- nism of flight such that flection between the metathorax and its postnotum was no longer necessary and that the intersegmental line (membrane) was lost, at least in the median area. Accompanying that change were; a re- duction in the number of segments of the antennal funicle from seven to as few as three and changes in the antennal club from cone- shaped, with simple, transverse sutures, to flattened and with or without sutures in nu- merous combinations. The primitive mating system was monogamy, with the female ini- tiating the new parental gallery. Limited de- partures from that system include male ini- tiated monogamy and heterosanguineous polygyny (many Polygraphini) and consan- guineous polygyny (Siieiis in Hyorrhynchini). A recognizable phyletic sequence appears to progress from the primitive Hylastini through Hylesinini to Tomicini. Hyorrhynchini and Phrixosomini appear to have been derived independently on separate lines from the same basic stock as that group of tribes. Among those tribes with a fused postnotum, Bothrosternini and Diamerini appear to be geographical replacements of one another that arose from a common ancestor that was much more closely allied to the Ctenophorini- like ancestor of Scolytinae than to the immedi- ate ancestral stock of the Hylastini- Hylesinini-Tomicini. Although the Phloeo- tribini and Phloeosinini are closely related to one another, they (as one unit) were probably derived from the Bothrosternini-Diamerini ancestral stock independently from the unre- lated Polygraphini and Hypoborini. Differen- tiation of phyletic lines in the Hylesininae has progressed slowly; consequently, they are dif- ficult to detect. In general body habitus, Pro- tohylastes (Platypodidae) resembles primitive hylesininae very closely, but the legs and an- tennae do not. Pycnariliritm (Ctenophorini) 24 Great Basin Naturalist Memoirs No. 10 would be placed in Hylesininae except for its legs. A primitive relationship of Hylesininae to primitive Scolytinae and Platypodidae is clearly indicated in these genera. Among the Scolytinae, the Ctenophorini, Scolytini, and Scolytoplatypodini are closely related to one another and were almost cer- tainly derived from a common Ctenophorini- like ancestral stock. The Ctenophorini more nearly resemble Protoplatijpus (Platypodi- dae) than do other members of this subfamily. Some Cnemomjx (allies of galeritus, Cten- ophorini) have crenulations on the basal mar- gins of the elytra as in Hylesininae. The four remaining clusters of tribes were probably derived from a Ctenophorini-like ancestor in the following pattern. Although there is no clear primitive connection to an ancestral group, Micracini and Catopinini are anatomi- cally and biologically allied to one another. An advanced member of Micracini probably gave rise to the Xyloctonini-Cryphalini group, then a Cryphalini-like member gave rise to the Corthylini. The remaining tribes, Carphodic- ticini, Ipini, Dryocoetini, Crypturgini, Xylo- terini, and Xyleborini (listed in ascending or- der of anatomical and biological complexity) were probably derived from another Mi- cracini-like ancestor along the same (one) phyletic hue of descent. Active major evolu- tionary thrusts appear to be in progress within the Dryocoetini, Xyleborini, Cryphalini, and Corthylini, which combined contain more than half the living species of Scolytidae. Methods Although this project was conceived in 1949, serious work on it was not started until 1955. From 1955 to 1965, comparative anatomical studies were made of representa- tives of more than 100 selected genera of Cur- culionoidea, in addition to more than 60 gen- era of Scolytidae and Platypodidae. As the patterns of evolution began to emerge from those studies, a major efltbrt was made to col- lect specimens and to gather behavioral and ecological data on species not previously known to me. More than 2,000 species of Scolytidae and 300 species of Platypodidae were collected from North and South Amer- ica, Europe, Asia, Australia, and New Guinea. In addition, visits were made for the purpose of studying types and comparing them and other specimens to my material at museums in the United States, Canada, Mex- ico, Venezuela, England, Austria, Finland, USSR, India, Japan, Australia, and Papua- New Guinea. There visits included stays of a month or more at the British Museum (Natu- ral History), the Schedl Collection (both at Lienz with Schedl in 1965 and at Vienna in 1983), and the Forest Research Institute, Dehra Dun, India (where more than 40,000 Indian specimens were sorted to species in addition to studies of the types). It is esti- mated that well over 400,000 specimens of Scolytidae and Platypodidae were examined. From those museums not visited in person by me, types and other specimens were obtained for study through loan. Except as noted below under Incertae Sedis, all existing type-species for all named genera and subgenera were ex- amined and, where type-specimens have been designated, the type-specimen of those species were also studied. The basis for recog- nition is given under each species and syn- onym below. Except at Helsinki (Finland), Leningrad and Moscow (USSR), Bulolo (Papua-New Guinea), and Sydney (Australia), where local museum equipment was used, the studies were made using an American Optical Com- pany stereoscopic microscope, model 25, equipped with an occular grid, at magnifica- tions of 10, 40, SO, and 160X. All measure- ments were made at lOX and excluded the head (in measurements of body length). Authentic specimens of the type-species of several genus-group names assigned to the Scolytidae were not examined during the course of this study. These species fall into two categories: (1) those for which the types or other authentic specimens could not be found and are presumed lost, and (2) fossil species. The types of four genera could not be found. These included: 1. Allarthnim Hagedorn (1912:355), based on A. kolbei Hagedorn (1912:355), from Pe- terhafen, Deutsch New Guinea. The type(s), 1.5 mm in length, was deposited in the Kgl. Zoolog. Museum Berlin but cannot now be found. The illustration of the antenna (Hage- dorn 1912:Fig. 5) is of a CrypJiahis species. It is provisionally listed as a synonym of Cryphahis . 1986 WOOD; Genera of Scolytidae 25 2. Bufonus Eggers (1919:231), based on B. obscii?-iiS Eggers (1919:231) from Amani, Ost- Afrika. The unique holotype, 1.5 mm, was lost with the Hamburg Museum in 1944. The an- tennal hjnicle was described as 2-segmented, the club elongate, with three transverse su- tures; the scutellum was not visible, the basal margins of the elytra were sharply roimded, and the elytral sculpture was pronounced and unique. Except for the absence of crenula- tions on the basal margins of the elytra, this genus would be placed in Hypoborini. A more detailed knowledge of the fauna of East Africa is needed before it can be correctly placed in classification. 3. Toxophorus Eggers (1920b; 119; preoc- cupied, renamed ro.vop/if/ior(/.s Wood), based on T. africanus Eggers (1920b: 119) from Deutsche-Ostafrika. The unique holotype, 3.0 mm, was lost with the Hamburg Museum in 1944. The antennal scape was elongate, the funicle was 5-segmented, the club was flat- tened, with a suture near the apex, and the frons was impressed above the eyes. This genus almost certainly is in the Dryocoetini, possibly near Tiarophoru.s or Xylocleptea . 4. Pseuc?o;?iicrac!.S' Eggers (1920a:36), based on P. ekae Eggers (1920a:36) from Dares- Salaam, Ostafrika. The unique female holo- type, 1.5 mm, was lost with the Hamburg Museum. The frons was impressed, the an- tenna was similar to Micracis, the scape was triangular, with long hair on the outer angle, and the sutural apex of the elytra mucronate. Because only one known genus bears the com- bination of characters described by Eggers, his name is associated with those African spe- cies previously referred to the American genus Micracis. As such, this genus is recog- nizable and is treated below under the name Pseudomicracis . Of the six fossil genera that have not been previously placed in synonymy under older names of modern genera, four are from Baltic amber, one is from Burmese amber, and one is from sedimentary deposits. These include: 1. Carphoborites Schedl (1947:32) was based on C. keilbacki Schedl (1947:32) and C. posticm Schedl (1947:.33), both from Baltic amber (Charphoborites was a lapsus calami). From the descriptions, I see no reason to separate these species from Carphohorus; however, I have not examined the specimens. 2. Hijlescierites Schedl (1947:29) was based on H. gramdatits Schedl (1947:.30) from Baltic amber. From the photograph of the holotype and the original description, I see no reason for separating this species from Hijlurgopa. 3. Taphramites Schedl (1947:41) was based on T. gnathotrichiis Schedl (1947:42) from Baltic amber. From the description, it ap- pears that this species should be placed in Dryocoetini, probably in or near Drijocoetes . I have not examined the specimen. 4. Xylechinites Hagedorn (1907:120) was based on X. anceps Hagedorn (1907:120) from Baltic amber and was redescribed and illus- trated by Schedl (1947:30-32). The descrip- tions and photograph of the holotype indicate that this species is a Tomicini near or in Xylecliintts . I have not examined the speci- men. 5. Cryphalities Cockerell (1917:368) was based on C. rugosissimus Cockerell (1917: 368) from Burmese amber. Because no usable characters were included, the original de- scription gives no clue as to where this species should be placed. It is assumed that Cockerell was correct in assigning it to Cryphalini. I have not examined the specimen. 6. Xyleborites Wickham (1913:26) was based on X. longipennis Wickham (1913:26) from the Wilson Ranch near Florissant, Colo- rado. The author indicated that this species, 2.2 mm in length, resembled Xyleborus pubescens . Essential characters were not de- scribed. The specimen was not examined by me. Pityophthoridea Wickham (1916:18) and Adipocepludus Wickham (1916:16) are not considered to be members of the Scolytidae. Systematic Section Although some obvious unanswered ques- tions remain as to the exact position of Platy- podidae and Scolytidae within the Cur- culionoidea, the following key is presented for the identification of families. The presence of complete pregular sutures and the mandible structure of Platypodidae and Scolytidae are obviously more primitive than the absence of these sutures and the specialized mandibular structure of Belidae and Oxycorynidae, but the single gular suture and presence of larval epipharyngial rods are more advanced than 26 Great Basin Naturalist Memoirs No. 10 the two gular sutures and absence of the rods in Behdae. The long subcortical history and very small body size of Platypodidae and Scolytidae have had an obvious effect on their anatomy that complicates interpretation of their position in phylogeny. This key to families of the Curculionoidea is tentative. A thorough reexamination of the Belidae-Aglycyderidae-Oxycorinidae groups is needed to determine the significance of single vs. double gular sutures, the precise position of character gaps that separate families, etc. Another area of concern is the section of the key in couplets 11 and 12. Char- acters to separate these three taxa are weak, but, in view of the large number of species to be classified herein, perhaps the recognition of these three families is justified. Among more than 150 genera of Curculionidae (in the sense used here) dissected by me, I saw three possible divisions of this family. These in- cluded (1) the Cossoninae-Rhynchophorinae, (2) the broadnose weevils, and (3) all others. It is recognized that some equally distinct groups may exist that were not seen by me, but a thorough comparative anatomical study should precede their recognition. Following my study, I seriously doubt that even one- third of the 75 or more subfamilies that have been recognized within Curculionidae de- serve this rank; most are worthy of no more than tribal status. Key to the families of Curculionoidea 1. Adult: two gulav sutures present or rudiments of them evident (only one gular suture and a postgula present in several Belidae and Rlio- palotria of O.xycorynidae); maxillary palpi 4- segmented; abdominal segments free (e.xcept 3-6 connate in Anthribidae); antennae usu- ally orthocerous, scape little longer than suc- ceeding segments, club loosely 3-segmented. Larva: epipharyngeal rods absent; maxillary palpi 3-segmented, rarely 2-segmented; tho- racic legs present; frontal sutures reaching articulating membrane of mandible; two folds in each abdominal tergum 2 — Adult: one gular suture extending from minute postgula at margin of foramen mag- num to posterior tentorial pits (usually where base of rostrum meets head); maxillary palpi 1- to 3-segmented (4-segmented in one Platy- podidae, some Rhynchitidae, and all Atte- labidae); abdominal segments free or partly connate in some groups; antennae frequently geniculate, a definite club usually present (or- thocerous in Brenthidae; intermediate in Ithyeeridae, Apionidae, some Platypodidae, • and a few Sc 'ytidae). Larva: epipharyngeal rods present; maxillary palpi 1- or 2-seg- mented (3-segmented in Brenthidae and Rhynchitidae); thoracic legs absent (present in Brenthidae and Ithyeeridae); frontal su- tures not reaching articulating membrane of mandible (except Brenthidae, Apionidae, Ithyeeridae, some Plat\ podidae); two to four folds in each abdominal tergmn 6 2(1). Adult; labrum distinct and separate (except absent in Bnichela of Anthribidae); maxillary palpi flexible, laeinia forming a distinct ele- ment (see also Rhynchitidae and Tessero- cerini in Platypodidae). Larva: mandibles with a distinct mola 3 — Adult: labrum fused or lost; maxillary palpi rigid; maxilla without a separate laeinia. Larva: mandibles without a distinct mola .... 4 3(2). Adult: gular sutures rudimentary, repre- sented only at margin of foramen magnum or totally lost; tentorium largely or entirely obso- lete; proventriciilus developed; lateral mar- gins of pronotum acute (rounded in B ruchela ); inner margin of elytra with a flange near costal margin; cavities of mesocoxae broadly closed ouKvardly by sternum, visible abdominal sterna 1-4 connate. Larva: clypeus distinct from frons, or head deeply retracted; mandibles without ventral process in addition to mola (includes the distinct subfamily Bruchelinae) Anthribidae — Adult: gular sutures extending from margin of foramen magnum to posterior tentorial pits; tentorium present; proventriculus indistinct; inner margin of elytra without a flange near costal margin; mesocoxae not or imperfectly closed outwardly by sternum; abdominal sterna free. Larva: clypeus usually fused to frons; head not deeply retracted into protho- rax; mandibles with a ventral process in addi- tion to mola Nemonychidae 4(2). Adult: gular sutures longer, extending to pos- terior tentorial pits; partial pregular sutures occasionally present in female near apex of rostrum; at least three genera with large post- gula and short median gular suture; antennae filiform or nearly so, inserted laterally remote from base of rostrum; inner margin of elytra with a flange near costal margin; wing with five anal veins in main group complete to base; tarsi normal, pseudotetramerous; lat- eral margins of pronotum transversely rounded. Lar\a: maxillary palpiger well de- veloped; thoracic spiracle intersegmental . . . Belidae — Adult: gular sutures usually short (a large postgula and one median gular suture in some Oxycorynidae), externally visible near fora- men magnum or else lateral margins of prono- tum acute; antennae inserted at or near base of rostrum; either antennae clubbed or tarsi pseudotrimerous; inner margin of elytra with- 1986 WOOD; Genera of Scolytidae 27 out a flange near costal margin; wings with fewer than five anal veins or else bases of elytral veins incomplete. Larva: maxillary palpiger usually not evident; thoracic spiracle in mesothorax 5 5(4). Adult: tarsi pseudotrimerous, segment 2 bilobed; antennal insertion at base of rather short rostrum; antenna filiform, not clubbed; proventriculus well developed. Larva: ma.\il- lary palpiger not distinct Aglycyderidae (=Proterhinidae) — Adult: tarsi pseudotetramerous, segments 2 and .3 bilobed; antennal club 2- or 3-seg- mented; (a large postgula and at least a partial median gular suture present in at least Rho- palotria), lateral margins of pronotum acute; antennal insertion at base of long rostrum, on ventral face; proventriculus poorly devel- oped. Larva: palpiger distinct . . Oxycorynidae 6(1). Adult: pregular sutures present and rein- forced internally by massive apodemal ridges; rostrum short to nonexistent; tibiae armed on apical or lateral margins by a series of spines and/or socketed teeth of setal origin, corbels never present; hypostomal area small, never armed by a large spine, posterior mandibular condyle more anterior in position; axis of mandibular hinge oblique to transverse, cut- ting edge on mesal margin (Fig. 8); some members (Tesserocerini) with a separate lacinia 7 — Adult: pregular sutures absent, with no indi- cation of supporting internal apodemal struc- ture (longitudinal ridges giving rigidity to ros- trum are not reinforced internally and should not be confused with true pregular sutures); rostrum variable, long to very short; lateral margins of tibiae unarmed (except for simple spines in some Cossoninae), corbels com- monly present; hypostomal area larger, com- monly armed by a large spine, posterior mandibular condyle more posterior in posi- tion (Fig. 8); axis of mandibular hinge parallel to body axis, cutting edge on apical margin; separate lacinia never present (except some Rhynchitidae and all Attelabidae) 8 7(6). Adult: tarsal segment 1 elongate, usually longer than 2-.5 combined (except shorter in Protoplatypiis , Protohijlastes. Coptonotiis. Scolytotarsus), tibiae armed on apical and/or lateral margins by one or more spines, never by socketed denticles, apical mucro formed by middle apical spine (as compared to Proto- hijlastes), male spiculum gastrale absent; an- tennal club solid, unmarked by sutures (fee- bly indicated in Coptonotiis), head about as wide as pronotum; pronotum with distinct lateral constriction near middle; when visible, scutellum often reduced, declivous, usually with not more than apex attaining (flush with) elytral surface; metepisternum very elongate, its anterior margin straight, its anternventral angle neither displaced caudad nor extended ventrad; mesepimeron flat, vertical; mese- pisternum moderately to greatly enlarged, varying from flat to inflated; eyes convex, never truly emarginate; pronotum never armed by asperities. Larva: clypeus fused to frons in aU except primitive genera; frontal sutures reaching articulating membrane of mandible or not Platypodidae — Adult: tarsal segment 1 about ecjual in length to 2 or 3; tibiae armed on apical and/or lateral margins either by spines or socketed denti- cles, apical mucro formed by mesal element (compared to Protohijlastes structure); male spiculum gastrale present; antennal club vari- able, with or without sutures; head narrower than pronotum; pronotum almost never with constriction near middle; when visible, scutellum usually flush with elytral surface, often flattened; metepisternum stout to elon- gate, its anterior margin usually procurved, its anteroventral angle displaced slightly cau- dad and distinctly extended ventrad; mese- pimeron flat, oblique; mesepisternum smaller, declivous cephalad; eyes more nearly flat against head, often emarginate to divided; pronotum often asperate. Larva: clypeus dis- tinct, separate from frons; frontal sutures never reaching articulating membrane of mandibles Scolytidae 8(6). Adult: labial palpi inserted ventrally on men- tum; head frequently constricted behind eyes; antennae never geniculate, inserted dorsally or laterally at base or middle of ros- trum, club absent to moderately distinct; proventriculus without selerotized plates. Larva: frontal sutures reaching articulating membrane of mandibles 9 — Adult: labial palpi inserted apicafly; antenna straight or geniculate, inserted laterally at middle or near apex of rostrum, club 1- to 3-segmented, loose to compact, small to large. Larva: frontal sutures either reaching or not reaching articulating membrane of mandibles 10 9(8). Adult: maxillary palpi 4-segmented, galea and lacinia distinct; mandibles often dentate on outer margin; labial palpi 1- or 3-segniented (rarely absent); elytra with a scutellary striole; notosternal suture short, laterally reflexed; hind wing usually witli four anal veins, radial cell well developed; sterna 3-6 rarely free, .3-4 connate in some, 3-6 connate in most; cap piece of male tegmen simple, setose; malpighian tubules grouped three on each side of alimentary canal; more than two ovari- oles in each ovary. Larva: legs absent, two or more stemmata present on each side of head; antenna usually 2-segmented; maxillary palpi 2- or 3-segmented; abdominal terga with two folds Attelabidae — Adult: maxillary palpi 3-segmented, galea and lacinia fused; mandibles never dentate on outer margins; elytra never with scutellary 28 Great Basin Naturalist Memoirs No. 10 striole; notostemal suture well developed and usually extending almost to anterior margin; hind wing with three or fewer ana! veins, radial cell not evident; sterna 3-4 connate and much longer than 5-6; cap piece of tegmen bilobed. Larva: rudimentary, segmented legs present; stemmata absent; antenna 1-seg- mented; maxillary palpi 3-segmented; ab- dominal terga with three to four lolds Brenthidae 10(8). Adult: maxillary galea and lacinia distinct, palpi 3- or 4-segmented; metepimeron sub- transversely carinate and interacting with humeral angles of elytra to aid in locking ely- tra in position, metepisternal locking element not developed and its interlocking flange at this point on inner costal margin of elytra absent; antenna straight, club loosely 3-seg- mented; abdominal sterna 3 and 4 connate, suture distinct. Larva: median area on poste- rior part of head overlapped by prothorax; maxillary palpi 2- or 3-segmented; two basal sensillae on labrum; frontal sutures complete but indistinct; abdominal terga with two folds Rhynchitidae — Adult: maxillary lacinia usually fused to galea, palpi 2- or 3-segmented; metepisternum cari- nate, fitting flange on inner margin of elytra near humeral angle, metepimeron not cari- nate; antenna straight or geniculate, clubbed or not. Larva: posterior part of head free in median area; maxillary palpi 1- to 3-seg- mented; no basal sensillae on labrum 11 11(10). Adult: abdominal terguni 8 longitudinally grooved for reception of inwardly raised ele- vation at suture near apex of elytra; tergum 8 bearing a pair of spiracles; maxillary palpi 3- segmented, but largely retracted into palpifer; antenna not geniculate. Larva: rudi- mentary, segmented legs present; maxillary palpi 2-segmented; 3 stemmata present .... Ithyceridae — Adult: abdominal tergum 8 not grooved, spir- acle absent; maxillary palpi 2- or 3-seg- mented, usually not retracted into palpifer . 12 12(11). Adult: either antenna not geniculate, or if geniculate then trochanters long, cylindrical, femur remote from coxa, attached to apex of trochanter; ventral surface of mentum usually with projecting setae; proventriculus poorly developed. Larva: frontal sutures extending to articulating membrane of mandibles; ab- dominal terga with two folds Apionidae — Adult: antenna usually geniculate; troch- anters triangular, femur attached to side of trochanter, sometimes almost touching coxa; ventral surface of mentum without projecting setae; proventriculus usually well developed. Larva: frontal sutures not reaching articulat- ing membranes; abdominal terga with two to four folds Curculionidae Family Scolytidae Latreille Scolytarii Latreille [1807:273, Type-genus: Scolytus Ge- offrey 1762, see also China 1962] The family Scolytidae is comprosed of a group of more than 6,000 small to minute species. Whereas the preponderance of spe- cies are tropical, a few of them reach the northern and southern limits of tree distribu- tion near the polar regions. They are unique in that loosely pair-bonded adult parents bore into subcortical tissues of their host (usually) before mating or oviposition occur. Eggs are variously placed in the galleries, and the lar- vae either expand the parental mines or form individual galleries while the parents defend the outer entrance to the tunnel. Thus, the beetles are essentially internal plant parasites that spend virtually their entire lives secreted within the tissues of the host. The ephemeral habitat they occupy is the unthrifty, weak- ened, or dying tissue of woody plants; only one generation is completed in each host un- less a large tree that is succumbing progres- sively is involved. Because of the very brief period this habitat is open to them, efficient means of locating a host, recruiting a popula- tion to subdue it, and overcoming host resis- tance are mandatory. This is accomplished through sensitivity to odors emitted by trees under stress, by a complex system of phero- mones, and by mutualistic relationships with fungi. Several of the most efficient species, with respect to meeting these challenges, compete with man for timber and horticul- tural resources and present a very real threat to human economy. Most woody plants, a few herbaceous plants, and many fruits and nuts are infested by these insects. The adjustments they have made in their mating systems, uti- lization of food resources, and adaptive radia- tion form a classic example of biological diver- sity. Key to the Subfamilies and Tribes of Scolytidae (Modified from Wood 1978) 1. Each basal margin of elytra procui-ved and armed by a series of marginal crenulations (Fig, 16) (or less commonly by a continuous elevated costa in some Bothrosternini, Di- amerini, Polygraphini), usually with a scutel- lar emargination between them; scutellum usually small and rounded or depressed, ab- sent in some groups; pronotum weakly if at all declivous on anterior half usually unarmed 1986 WOOD; Genera OF Scolytidae 29 Fig. 24. Eupagiocerus dentipes Blandford, protibia: A, posterior face showing the bifid spine on the lateral, apical angle and small socketed denticle; B, an enlargement of the socketed denticle. 2(1). but crenulations sometimes present on anterolateral areas; head usually visible from above, somewhat wider; protibia usually wider; scales or deeply divided setae a com- mon feature (subfamily HYLESININAE) .... Basal margins of elytra forming a straight, transverse line across body (Fig. 17), un- armed, rarely (some Scolytini, Ctenophorini, Cryphalini) with a weakly elevated continu- ous line; scutellum usually large, flat (rarely absent or highly modified in some Xyle- borini); pronotum weakly to strongly de- clivous on anterior half and usually armed by many asperate crenulations, particularly on median half (Figs. .31, 46, .52); head usually partly or entirely concealed from dorsal as- pect, somewhat narrower; protibia usually narrower; scales or deeply divided setae an uncommon feature (subfamily SCOLYTl- NAE) Scutellar area of metanotum and its postno- tum separated by a suturelike intersegmental membrane (Fig. 23, pts. 43, 45); posterior part of scutoscutellar suture strongly curved mesad to a point near crest of scutellar groove then continuing cephalad parallel to this costa for about two-thirds of metanotum length (Fig. 23, pt. 43) (except much less in Phrixoso- mini and Hyorrhynchini); metapleural suture descending subvertically from pleural wing process to metepisternal groove formed to receive corresponding costal groove and flange of elytron then abruptly angled and continued caudad along this groove to a point near pleural coxal process (Figs. 18, 23 pt. 44); scutellum visible; funicle 6- or 7-segmented or if 5-segmented (Sueus) then eye divided, male frons not impressed, and antennal club symmetrical 3 — Scutellar area of metanotum and its postno- tum completely fused on at least median third, intersegmental suture usually obsolete (Figs. 21, 23 pt. 41); scutoscutellar suture less 2 strongly curved, approaching costa of scutel- lar groove more gradually and continuing cephalad parallel to it for less than half length of metanotum (Fig. 23, pt. 41) (it never reaches margin of this groove in some groups; metapleural suture sometimes as described above, but more commonly running a more direct route from pleural wing process to pleural costal process, often remote from locked position of costal margin of elytra for most or all of its course (Figs. 21, 23 pt. 41); scutellum either not visible or if visible then funicle 5-segmented and male frons im- pressed (Bothrosternini with 6-segmented fu- nicle but with a distinctive protibia. Fig. 24) 7 12 3(2). Eye entire to feebly emarginate; scutoscutel- lar suture parallel to costa of scutellar groove for two-thirds length of notum; preco.xal ridge on prothorax present (Fig. 20) or absent; an- tennal funicle 5- to 7-segmented 4 — Eye completely divided by an emargination, halves widely separated; scutoscutellar suture remote from costa of scutellar groove; crenu- lations on basal margins of elytra low, often poorly formed; precoxal ridge on prothorax never present; antennal funicle usually 6-seg- mented (5-segmented in Sueus) 6 4(3). Prothoracic precoxal area rather large, its lat- eral margins strongly, sharply elevated from anterior margin to coxae (Fig. 20); crenula- tions on elytral bases usually poorly devel- oped; antennal funicle 7-segmented, club conical, segment 1 often as long as others combined; head somewhat prolonged, sub- rostrate, frons never sexually dimorphic; eyes 30 Great Basin Naturalist Memoirs No. 10 Fig. 25. Mesal aspect of elytral base: A-B, Pseudohylesinus nehulosus (LeConte), groove nearest pubecent surface continues to base without interruption (ignore the tiny grain of frass trapped in the groove in A); C, Phloeosinus cristatus (LeConte), groove near base interrupted by a series of interlocking nodules and cavities. entire, rather short; Northern Hemisphere, except introduced elsewhere; in Pinaceae . . Hylastini — Prothoracic precoxal piece small, short, its lateral areas elevated or not; crenulations on 7(2). elytral bases more conspicuously elevated, forming a definite row (except confused in Dactijlipalpus): antennal funicle variable, .5- to 7-segmented, club weakly to moderately flattened; head less distinctly rostrate, male frons usually impressed, eye oval to elongate, entire to feebly emarginate 5 5(4). Pronotum asperate on anterolateral areas (ex- cept Hylastinus), prothorax with elevated costate ridge from coxa to anterior margin (weaker than in Fig. 20); antennal funicle 6- or 7-segmented; mesal surface of elytra at base of suture immediately behind scutellum with an interlocking series of nodules and cavities (Fig. 25-C), this lock interrupts groove and flange of suture (not visible when elytra in locked position); worldwide Hylesinini — Anterolateral areas of pronotum unarmed; precoxal costa on prothorax absent; funicle 5- to 7-segmented; mesal surface of elytra at su- ture with interlocking groove and flange con- ^(y\ tinuted to base without a series of nodules or cavities immediately behind scutellum (Fig. 25-A-B); worldwide Tomicini 6(3). Protibia armed on outer apical margin by sev- eral socketed denticles of approximately equal size; procoxae contiguous; frons convex, not sexually dimorphic; antennal scape elon- gate, suture 1 of club partly septate; prono- tum never armed by asperities; America, Africa Phrixosomini — Protibia with outer apical angle produced into one conspicuous spine reaching level of tarsal insertion, outer margin without any socketed — teeth; procoxae rather widely separated; male frons broadly impressed (except in dwarfed Sueus), female frons convex; antennal scape either long or short, club aseptate; pronotum either with or without asperities in anterolat- eral areas; southeast Asia Hyorrhynchini Lateral margins of pronotum usually sub- acutely elevated, costate (as in Fig. 12); mesepimeron moderately to very large, its dorsal portion usually grooved for reception of elytral base; scutellar shield under base of elytra large, extending posteriorly beyond visible scutellum (Fig. 26); scutoscutellar su- ture remote from costa of scutellar groove to its base; outer apical angle of protibia often with only one major recurved spine (Fig. 7); Africa, southeast Asia to Australia . . Diamerini Lateral margins of pronotum rounded (Fig. 16) (subcostate in a few neotropical Both- rosternini. Fig. 11); mesepimeron not en- larged or grooved (feebly grooved in Aricerus), scutellar shield beneath elytra small if present, not extended caudad beyond visible scutellum (Figs. 18, 21); scutoscutellar suture near and parallel to costa of scutellar groove on at least anterior fourth of metan- otum 8 Outer apical angle of protibia with a curved bifid process, meso- and metatibiae with one or two (usually smaller) cui-ved spines on outer apical angle extending beyond level of spine on inner apical angle (Fig. 24); prono- tum smooth or longitudinally strigose; funicle 6-segmentecl; lateral prosternal area usually subacutely elevated from coxa to anterior margin; anterior coxae rather widely sepa- rated; crenulations on elytral bases rather small or (rarely) replaced by a continuously elevated costa; eye entire; America Bothrosternini Outer apical margin of protibia armed by sev- eral teeth of about equal size (except Aricerus in Phloeotribini), none of them extending be- 1986 WOOD; Genera of Scolytidae 31 Fig. 26. Diagram of Sphaerotrypes globosus Blandford, with elytra removed, showing enlarged scutellar plate that lies below the base of the elytra. yond tarsal insertion; funicle 4- to "-seg- mented; prostemal area with margins rounded, costa obsolete; eye varying from en- tire to emarginate to divided 9 9(8). Scutellum visible, elytral bases notched for its reception; tarsal segment .3 stout, usually somewhat bilobed (except slender in Chramesus), mesal surface of elytra at suture immediately behind scutellum with a series of interlocking nodules and cavities (Fig. 25-C) 10 — Scutellum obsolete, elytral bases only slightly if at all emarginate at suture (Fig. 17); tarsal segment .3 slender; mesal surface of elytra at suture usually without a special lock, groove and flange extend to base at position of .scutel- lum (degenerate in Fig. 25-A-B) 11 10(9). Antennal club constricted at sutures and mov- able at intersegmental lines (Figs. 16, 27); Holarctic, Neotropical, and Australian Phloeotribini — Antennal club immovably, broadly fused at sutures, sutures often partly or entirely obso- lete; worldwide Phloeosinini 11(9). Eye emarginate or entirely divided; prono- tum never armed by asperities; crenulations at bases of elytra more widely distributed, extending laterad beyond interstriae 5 (Fig. 16); funicle 5- or 6-segmented; scutoscutellar suture passing near and parallel to costa of scutellar groove on anterior fourth of meta- notum; Northern Hemisphere and Africa . . . Polygraphini — Eye sinuate or entire; pronotum armed by a few scattered or clustered asperities; crenula- tions at bases of elytra restricted to area be- tween suture and interstriae .5; funicle 3- to 6-segmented; scutoscutellar suture remote from costa of scutellar groove on anterior fourth of metanotum; almost worldwide .... Hypoborini 12(1). Lateral margins of pro- and metatibiae un- armed except for a single, apical, spinelike process that curves toward and extends be- Fig. 27. Antenna of Vhloeotribus . A, rhododacttjlus (Marsham); B, caucasicus Reitter. yond process of inner apical angle (Fig. 7, Camptocerus). lateral margins of pronotum subacutely elevated, costate; pleural suture descending subvertically from pleural wing process to groove receiving groove and flange on costal margin of elytra, at this point suture turns abruptly and follows groove caudad to metapleural coxal process (Fig. 23, part 46); funicle 7-segmented, sutures of antennal club strongly procurved or obsolete; Holarctic and Neotropical Scolytini — Lateral margin of protibia armed by more than one denticle, none of which exceed or curve toward inner apical process; pleural su- ture less strongly angulate, groove receiving flange of costal margin of elytra displaced ven- trad from course followed by pleural suture (Fig. 23, part 42); lateral margins of pronotum subacutely raised or not, antenna variable .. 13 13(12). Metepisternum visible throughout its length, shghtly more than its dorsal half covered by elytra when in locked position, either with a conspicuous groove for reception of costal flange throughout its length or else groove represented at its anterior end by a denticula- tion or costate remnant near anterior end of metepisternum (Fig. 23, parts 42, 44, 46); antennal club varying from flat to obliquely truncate (Fig. 29) 14 — Metepisternum largely covered by elytra, its groove for reception of costal flange obsolete, a small, transverse callus (Cryphalini, Fig. 31) or a small transverse groove (Corthylini, Figs. 23, part 47, and 32) at anterior end of met- episternum; antennal club strongly flattened; antennal club never obliquely truncate 24 14(13). Lateral margins of pronotum subacutely ele- vated, basal margins of elytra usually finely elevated; procoxae rather widely separated except contiguous in Xyloctonini; protibia 32 Great Basin Naturalist Memoirs No. 10 Figs. 28-32. Scolytidae parts: 28, Ips woodi Thatcher, ventral aspect of prothorax showing contiguous coxae, left coxa removed (ignore the plant fiber between the coxae); 29, obliquely truncate antennal club of Dnjocoetes confusits Swaine; 30A, posterior face of protibia of Scolytopluttjpus papuanus Eggers showing unsocketed spines on lateral margin, B, enlargement of spines 1 and 2; 31, dorsal aspect of posterolateral area of prothorax olCnjphalus ruficollis Hopkins, with left elytron removed to expose anterolateral area ol metathorax (arrow points to the groove that is partially replacing the metepisternal spine; 32, left pleuron ofCorthijlui panamensis Blandford, with elytron removed; upper arrow points to pleural suture, lower one to the groove that has entirely replaced the metepisternal spine. 1986 WOOD: Genera of Scolytidae 33 with prominent outer apical process recurved (Fig. 7, Scolytodes). usually extending be- yond tarsal insertion, posterior tibia tapered on apical third and armed by several small socketed denticles; funicle6-or7-segmented; tarsi often retractible into tibial grooves .... 15 — Lateral and basal margins of pronotum rounded (except Cnestus); procoxae subcon- tiguous (except most Micracini and a few Xyleborini); protibia with outer apical angle inconspicuous, armed by several small sock- eted denticles; funicle 2- to 6-segmented; tarsi not retractible (except in Eccoptopterus) 17 15(14). Eye divided or nearly so by a very deep emar- gination; antennal club flat, usually enlarged, with sutures strongly procurved; abdomen conspicuously ascending toward apex (not al- ways clear in Clonoxijlon). tarsi always re- tractible into tibial grooves; Africa and south- em Asia Xyloctonini — Eves entire to shallowly sinuate on anterior margin; antennal club flat, usually smaller, more slender, sutures variable, if present; ab- domen horizontal; tarsi retractible or not ... 16 16(15). Antennal club with one or more sutures indi- cated by grooves, setae, or septae; scutellum large, flat; America; mostly phloeophagous, never mycetophagous Ctenophorini — Antennal club unmarked by sutures; prono- tum with sides strongly constricted on poste- rior half scutellum absent (a small scutellum present in Scolytoplattjpus congonus). Africa to Asia and New Guinea; mycetophagous . . . Scolytoplatypodini 17(14). Procoxae moderately separated; protibia with sides parallel, armed by denticles only on api- cal margin or posterior face; funicle 6-seg- mented (5-segmented in one African genus); female frons often concave, male frons rarely concave (except two Pseudotlujsanoes); Africa and America, one species in Asia .... Micracini — Procoxae contiguous (except Carjihodicticini, some Xyleborini); protibia much wider api- cally, armed on lateral margin by several den- ticles; female frons rarely concave (a few Dry- ocoetini), male frons often concave; funicle 2- to .5-segmented, 6-segmented in Tiurophorus 18 18(17). Male frons strongly excavated, epistoma armed by a pair of (usually) fused horns of enormous size (Fig. 47); funicle 5-segmented, club often small and feebly flattened; eye small, entire; pronotum with summit near basal margin, projecting back over scutellum in some species; western United States and Mexico Cactopinini — Not fitting above combination of characters . 19 19(18). Meso- and metathoracic tibiae more slender, more abruptly narrowed on apical fourth, lat- eral and apical margins armed by fewer, coarser teeth; eye sinuate to shallowly emar- ginate (divided in Tiarophorus , Dryocoetini, Africa); pronotum sometimes with a raised line on basal or lateral margin; prcgnlar area not depressed; se.xes of similar size and body form (except male dwarfed and deformed in Coccotrypes and Ozopcmon); habits varied but never woodboring or mycetophagous ... 20 — If eye completely divided into two parts and antennal funicle 4-segmented then male frons deeply excavated and male equal in size to female; if eye emarginate (or if divided and funicle 5-segmented) then male dwarfed, de- formed, and flightless and female meso- and metathoracic tibiae expanded to just beyond middle then arcuately tapered to apex, its apical two-thirds on outer margin armed by a row of numerous small, closely set teeth of equal size, these usually supplemented in same row by submarginal hair on posterior face; male pronotum highly modified; pregu- lar area depressed (except Premnobius), woodboiing, mycetophagous 23 20(19). Pronotum rather strongly, laterally con- stricted on posterior half anterior half not declivous and never armed by asperities; anterior coxae moderately separated; anten- nal club strongly flattened, marked by two sutures, sutures on posterior face almost equal to those on anterior face; South America and India to Ceylon (Sri Lanka) Carphodicticini — Pronotum not constricted, sides straight to arcuate, anterior half usually declivous, usu- ally armed; anterior coxae contiguous; anten- nal club obliquely truncate or with sutures on posterior flice strongly displaced toward apex (rarely with sutures obsolete) 21 21(20). Eye shallowly sinuate (shallowly emarginate in some Acanthotomicus), its lower half dis- tinctly narrower than above; piotibia with 3-4 socketed teeth; antennal club rarely obliquely truncate (Pityokteines. Orthotomicus), pro- coxae contiguous, intercoxal piece longitudi- nally emarginate to absent, never complete; elytra moderately sulcate to elaborately exca- vated, with lateral margin usually armed by tubercles or spines; pronotum more strongly declivous on anterior third, asperities usually larger; worldwide Ipini — Eye sharply, rather deeply emarginate (sinu- ate in Deropria), lower half usually almost equal in width to upper half protibiae usually with four or more socketed teeth (most excep- tions with lateral margins of pronotum acutely elevated); procoxae either contiguous or dis- tinctly, narrowly separated; elytral declivity flattened to convex, unarmed by spines or large tubercles; pronotum either evenly arched from base to anterior margin or less strongly declivous on anterior third, asperi- ties, when present, usually fine and abundant (a few exceptions) 22 34 Great Basin Naturalist Memoirs No. 10 22(21). Antennal funicle 4- to 6-segmented, club ei- ther obliquely truncate or with .sutures on posterior face strongly displaced toward apex; anterior half of pronotum more strongly de- clivous and rather coarsely asperate (unarmed in Tiarophorus); worldwide Dryocoetini — Antennal funicle 2- or 3-segmented, club with sutures on posterior face about equal to those on anterior face; pronotum feebly declivous on anterior half and unarmed (minutely gran- ulate in some Aphanarthrum), reticulate in many species; size small; Northern Hemi- sphere and Africa Crypturgini 23(19). Eye always completely divided into two parts; antennal funicle 4-segmented, base of club feebly to moderately corneous, usually pubescent to base; male subequal in size to female, his frons flattened or e,\cavated and anterior margin of his pronotum more broadly rounded; male joins female in parental gallery, reproduction always bisexual; Hol- arctic and Oriental Xyloterini — Eye emarginate except divided in some Amasa; funicle 5-segmented (3- or 4-seg- mented in a few Asiatic forms); males flight- less, dwarfed, deformed, anterior slope of pronotum variously excavated; male head convex; male absent from parental gallery ex- cept as progeny; partly parthenogenetic, male haploid; almost worldwide Xyleborini 24(13). Costal margin of elytra slightly to moderately ascending from base of declivity to apex; basal end of metepisternum armed by a callus or partial groove of degenerating metepisternal spine (Fig. 31); sutures on posterior face of antennal club more strongly displaced toward apex; funicle 3- to 5-segmented; tibiae more strongly flattened, usually armed by more than four denticles; vestiture commonly in- cludes scales; eye usually entire, less com- monly emarginate; worldwide Cryphalini — Costal margin of elytra descending toward apex (e.xcept Brachyspartus), basal end of metepisternum with a small, transverse groove (Fig. 32) (concealed when elytra in locked position), elytra in locked position more completely cover metepisternum (Fig. 52 part 5, and 53); sutures on posterior face of antennal club only slightly displaced toward apex; funicle 1- to 5-segmented; tibiae more slender, rarely armed by more than four sock- eted denticles; vestiture rarely includes scales (in tropical forms only); eye emarginate; al- most worldwide except Australia .... Corthylini Subfamily Hylesininae Hylesinen Erichson [1836:46, Tvpe-genus: Hylesinus Fabricius, 1801] Most previous classifications have recog- nized the Hylesininae as a major division of the taxon treated here as Scolytidae; however. there has been variability in the taxonomic rank assigned to it. Most of the distinguishing characters employed previously are not found consistently throughout the group and, conse- quently, have little or no taxonomic value. The most consistent and reliable character available for the recognition of this subfamily is the procurved basal margins of the elytra that are armed by a row of crenulations, and the scutellar notch between them. The heav- ier, more coarsely armed tibiae are distinctive but less reliable as a distinguishing feature. The more primitive Bothrosternini, Diame- rini, and a few other isolated examples lack the specialized elytral crenulations and have the basal margins elevated along a continuous costa reminiscent of some Platypodidae. This same character also occurs in Dendrodicticus (Carphodicticini) and, in a greatly reduced form, many Scolytini, Ctenophorini, and Cryphalini. On the other hand, a few Cnemonijx (allies oi galeritus, Scolytini) have fully formed basal crenulations that suggest an affinity with the Hylesininae. Even though the demarkation between subfamilies is not as sharp as some would like, the division of sub- families is simple and the characters are reli- able. Tribe Hylastini Hylastes LeConte [1876:387, Type-genus: Hylastes Erichson, 1836] Description. — Frons not sexually dimor- phic; eye oval, entire; antennal scape elon- gate, funicle 7-segmented; precoxal ridge on prothorax strongly, acutely elevated; protibia very broad, with rather numerous lateral socketed teeth; scutellum visible; crenula- tions at base of elytra poorly developed, usu- ally net forming a definite row; declivital in- terstriae 10 sometimes present (Scieriis); declivital sculpture usually simple; vestiture usually includes scales (some exceptions); scu- toscutellar suture parallel to costa of scutellar groove for two-thirds length of pronotum; metapleural suture descending subvertically from pleural wing process to metepisternal groove (this groove interlocking with groove on costal margin of elytron) then continuing caudad along this groove to a point near pleural coxal process; tarsal segment 3 wider than 1 or 2. 1986 WOOD; Genera of Scolytidae 35 Biology. — All species are monogamous. They breed in coniferous hosts, usually at the base or in the roots of large trees, although some species prefer the lower surface of pros- trate logs that are in contact with the ground. Parental galleries are monoramous or bi- ramous, usually entirely in the phloem, but exposed on peeled bark. Eggs are placed in niches and are sealed in with frass. Larval mines are comparatively long and wander ir- regularly in the phloem. The species are not agressive, usually preferring unthirfty or felled trees after they have been attacked by other bark beetle species, except a few species of Hylastes have been reported to kill seedlings in nursery stock. Their role in the primary destruction of roots is unstudied. Taxonomy. — Members of this tribe are the most common fossil Scolytidae in Baltic am- ber (Oligocene). They are not always clearly separable from the Hylesinini to which they are obviously closely related phylogeneti- cally, and they appear to be of ancient origin in the family. Their specialized habits appar- ently have resulted in evolutionary paral- lelism and superficial similarity of appearance with certain Cossoninae (Curculionidae), re- sulting in an erroneous supposition that the two groups are closely related. The generic limits within the Hylastini are not sharp. Indi- vidual variation within species and the simi- larity of species within genera make specific identification in the group rather difficult. They are strictly holarctic in distribution, if introductions to southern Africa, New Zealand, and Australia are ignored, and they are confined to the Pinaceae. Key to the Genera of Hylastini 1. Anterior coxae rather widely separated by an intercoxal piece, its width at least equal to half width of a coxa; striae 9 and 10 both indepen- dently continued at least to level of abdominal sternum 4; elytral vestiture sparse, recumbent, yellow, hair- or bristlelike, never including scales; general surface of elytra and pronotum rather dull; body color reddish brown; North America; Picea, Abies, Pinus, 2. 7-4. .3 mm .... Scierus — Anterior coxae contiguous or at most with inter- coxal piece narrower than one-fourth width of a coxa; striae 10 obsolete behind level of posterior coxae, only striae 9 continuing caudad; elytral vestiture longer, more abundant, usually in- cluding at least some small scales 2 i I_ BHhh Wk WSi ^^^jj^^^^H^^^^^^H ^. \ ,^ ^^Hh^^^^^HM mw\£ ^f- ^^^i^^^^K «#.'?'<-. s^H| mU'i ^^^R if^"'' ^s fei*- t fi 1a'^' M 1'<'S' IHH ^^^H m- W «i }-\, ' ^ '^h i£ ■■ -^^^^X-' Fig. 34. AIniphagus aspericoUis (LeConte), dorsal aspect (After Bright 1976: 207). male, impression not extending above upper level of eyes; eye shallowly emarginate, elytral ground vestiture scalelike, costal margin near apex descending; E Asia; 2.2-2.8 mm . Neopteleobius — Funicle 7-segmented; female frons flat to con- vex; male frons, if strongly concave, with exca- vation extending above eyes; eye less strongly to not emarginate .- 8 8(7). Eye entire, oval, less than 3.0 times as long as wide; protibia armed on lateral margin of apical fourth by six or more closely set, socketed teeth; body stouter; declivity more gradual, abdomen distinctly ascending to meet elytral apex; elytral vestiture of uniform length, mostly of scales (except almost subglabrous in crenatus), almost worldwide; Fraximis and other Oleaceae; 1.7-4.8 mm Hylesinus — Eye shallowly emarginate, somewhat elon- gate, at least 3.3 times as long as wide; protibia armed by 2-.5 socketed teeth; body more slen- der; declivity shorter, more abrupt, abdomen horizontal, not rising to meet elytral apex; ely- tral vestiture of ground cover of short hair or scales, and interstrial rows of longer, erect bris- tles 9 9(8). Male frons weakly, very shallowly impressed; protibia armed by five socketed teeth, outer apical angle only moderately abrupt; antennal club more nearly subconical; setae in elytral ground vestiture subplumose; Japan to W North America (Fig. 34); A/iu«; 2.1-3.4 mm . AIniphagus — Male frons moderately to extensively exca- vated; protibia armed by no more than four socketed teeth, outer apical angle abrupt; an- tennal club more strongly flattened, its apex less narrowly roimded; setae in elytral ground vestiture undivided, abundant to absent; Aus- tralia to Japan and China; Fictis, rarely other hosts; 1.6-5.0 mm Ficicis 10(2). Pronotum subtriangular, scutellum small to absent; funicle 5-, 6-, or 7-segmented; striae often impressed, punctures usually larger; Africa; 1.3-3.0 mm Hylesinopsis — Pronotum subquadrate, scutellum rather large; fimicle 6-segmented; striae usually weakly impressed, narrow, punctures fine to obsolete; Africa; 1.5-4.5 mm ... Rhopalopselion 11(1). Antennal club with two sutures (often obscure); female proepisternum with a large myce- tangium ornamented by hair; metatarsus re- traetible into tibial groove; Central and South America; 5.0-16.0 mm Phloeoborus — Antennal club unmarked by sutures; female pronotum with a median, transverse, slitlike myeetangium on anterior third; metatarsus not retractible, tibial groove absent; Africa, SE Asia, Philippines; 6.0-14.0 mm . . . Dactylipalpus Hylastinus Bedel [1888:388, Type-species: Ips obsciiriis Marsham, original designation]. Distribution: 2 in Europe, 1 in N Africa; one of these (obscurus) has been introduced into most temperate areas of the world. One is phloeophagous in Cijtisus; one (obscurus) breeds in the roots of various clovers. Keys: Reitter (1913:45). Kissophagits Chapuis [1869:34, Type-spe- cies: Hylesinus hederae Schmitt, monobasic]. Distribution: 4 in Europe and neighboring areas. All are phloeophagous and monoga- mous. Key: Reitter (1913:44). Pteleobius Bedel [1888:392, Type-species: Bostrichus vittatus Fabricius, subsequent designation by Hopkins 1914:128]. Distribu- tion: 2 in Europe, one of these is recorded from N Africa. They are phloeophagous in Ubnus and Sorbus and are monogamous. Keys: Reitter (1913:42), Balachowsky (1949: 94). 1986 WOOD: Genera of Scolytidae 39 Cryptocurus Schedl [1957c:869, Type-spe- cies: Cryptocurus spinipennis Schedl, mono- basic. Synonym; Hyloperus Browne 1970: 546, Tvpe-species: Hyloperus bicornis Browne = Cryptociirus spinipennis Schedl, original designation]. Distribution: 1 in Africa (Nige- ria to Tanganyika). It was attracted to light and was also taken from a Piptadenia buchananii log. Neoptelobius Nobuchi [1971:125, Type- species: Hylesinus scutidatus Blandford, orig- inal designation]. Distribution: 1 species in E Asia and Japan. Phloeophagous in several broadleaf hosts. Alniphagus Swaine [1918:73, Type-spe- cies: Hylesinus aspericoUis LeConte, mono- basic. Synonym: Hylastinoides Spessivitzev 1919:249, Type-species: Hijlastes alni Ni- isima, monobasic]. Distribution: 2 in W North America, 1 in Japan and E Asia. All are phloeophagous in Alnus and are monoga- mous. Keys: Bright (1976:74) and Wood (1982:119) for North America, Hylesinus Fabricius [1801:390, Type-spe- cies: Hylesinus crenatus Fabricius, subse- quent designation by Westwood 1838:39. Synonyms: Leperisinus Reitter 1913:41, Type-species: Bostrichus fraxini Panzer = Bostrichus varius Fabricius, subsequent des- ignation by Swaine 1918:70]. Distribution: 10 in North America, 1 in South America, 5 in Europe, 9 in Asia, 1 in Africa (Uganda). More than 20 additional nominate species occur in S Asia, Australia, and neighboring areas. All are phloeophagous and are common in Fraxinus or other Oleaceae hosts; they are monoga- mous. Keys: Wood (1982: lio) for North America, Murayama (1963:6) for NE Asia. Fields Lea [1910:147, Type-species; Fields varians Lea, subsequent designation by Hop- kins 1914:122. Synonym: Fieiphagus Mu- rayama 1958:930, Type-species; Ficiphagus goliathoides = Hylesinus porcatus Chapuis, original designation]. Distribution: About 16 nominate species in the area from India and Japan to Australia. All are monogamous and phloeophagous in Ficus, rarely in other hosts (particularly Artocflrpu.s). Hylesinopsis Eggers [1920a:40, Type-spe- cies; Hylesinopsis duhitis Eggers, monobasic. Synonyms: Pseudohylesinus Eggers 1919; 234, Type-species; Pseudohylesinus togonus Eggers, monobasic, preoccupied; Mefa/i!//esi- nus Eggers 1922:165, Type-species; Pseudo- hylesinus togonus Eggers, automatic; Pseudo- phloeotribus Eggers 1933b: 18, Type- species; Pscudophlocotiihus africanus Eg- gers, monobasic; Trypographus Schedl 1950e:213, Type-species: Trypographus joveri Schedl, monobasic; Chilodendron Schedl 1953a:74, Type-species: Chiloden- dron planicolle Schedl, monobasic; Glo- ehicoptcrus Schedl 1954b;75, Type-species; Glochicopterus baphiae Schedl, monobasic; Hapalophlocus Schedl 1966b:363, Type-spe- cies: Metahylesinus brinckei Schedl, original designation; Hemihylesinus Schedl 1967:224, Type-species: Hemihylesinus endrocdyi Schedl, monobasic]. Distribution; About 38 species in Africa (Considerable synonymy is anticipated). Apparently all are phloeo- phagous and monogamous. Rhopalopselion Hagedorn [1909:740, Type-species; Rhopalopselion bitubercida- tum Hagedorn, monobasic. Synonym: Hapa- logenius Hagedorn 1912:352, Type-species: Hapalogenius globosus Hagedorn, monoba- sic]. Distribution: About 30 African species. Apparently all are phloeophagous and monog- amous. Phloeoborus Erichson [1836:54, Type-spe- cies; Phloeoborus rudis Erichson, subsequent designation by Hopkins 1914:126. Synonym; Phloeotrupes Erichson 1836:53, Type-spe- cies; Phloeotrupes grandis Erichson, subse- (juent designation by Hopkins 1914:127]. Dis- tribution: About 24 species, S Mexico to N Argentina. All are xylophagous in rather large host material and are monogamous; they are occasionally attracted to light. They have an intimate association with fungi, but are not mycetophagous. Keys; Blandford (1897:150, Eggers (1942:267), Wood (1982:122). Dactylipalpus Chapuis [1869:12, Type- species; Dactylipalpus transversus Chapuis, subsequent designation by Hopkins 1914: 120. Synonyms: Dactylopselaphus Gem- minger & Harold 1872:2678, Type-species; Dactylipalpus transversus Chapuis, auto- matic; Ethadopselaphus Blandford 1896:321, Type-species; Ethadopselaphus cicatricosus Blandford, original designation]. Distribu- tion; 10 African and 2 Philippine and SE Asian species. Apparently xylophagous in large host material; occasionally attracted to light. Key: Eggers (1933c;200). 40 Great Basin Naturalist Memoirs No. 10 Tribe Tomicini Tomicidae Thomson [1859: 145, Type-genus: Tomicus Latreille, 1802/3] Hylurgini LeConte [1876: 373, Type-genus: Hylurgiis Latreille, 1807] Dendroctonides Niisslin [1912b: 273, Type-genus: Den- droctonus Erichson, 1836] Xylechinides Niisslin [1912b: 273, Type-genus: Xijlechi- nus Chapuis, 1869] Description. — Frons very weakly to mod- erately sexually dimorphic, male usually im- pressed, female convex; eye oval to ovate, entire; antennal scape elongate, funicle 4- to 7-segmented, club symmetrical, feebly to moderately flattened, three sutures usually indicated; pronotum unarmed, except a few very small asperities sometimes present in Xylechinosomus and some Xijlechiniis ; pro- coxae contiguous to moderately separated, precoxal lateral costa absent; metascutellar area separated from postnotum by a distinct suture; sutural groove on mesal surface of ely- tra continuing to base without a series of inter- locking nodules and cavities; tibiae armed by socketed denticles. Biology. — All species are monogamous; all are phloeophagous except for the xylophagous Pachycotes (and Hylurgonotus?). The parental galleries are usually biramous, ex- cept in Dendroctoniis, Hylurgns , and some Tomicus they are monoramous. Those of Sinophloeus, Hylurgonotus , and Pachycotes are not known to me. The eggs are placed in niches and packed in frass except that some Dendroctonus have modified the niches into elongate grooves into which numerous eggs are packed in single or double rows. The larval mines usually show on the inner surface of peeled bark and are oriented in a direction away from the parental mine. Symbiotic rela- tionships with fungi may occur in all genera, but they are adapted toward overcoming re- sistance of the host and are not of a myce- tophagous type. Taxonomy. — The worldwide distribution of this diversified tribe suggests an ancient origin, although only one Tertiary fossil has been reported (Xijlechinus , Oligocene). The most conspicuous division of the group is that presented in couplet 1 of the key, except that Xylechinus should be placed with the first group of genera. The Tomicini represent the most highly evolved segment of an evolution- ary trend that began in the Hylastini, contin- ued in the Hylesinini, and reached its greatest specialization in the Tomicini. Two major clusters of genera appear within the tribe; first, the Xylechinus group of genera (Xylechinus, Chaetoptelius, Xylechinosomus, Sinophloeus, Denclrotrupes, Hylurgopinus , Pseudoxylechinus , and Pscudohylesinus) and, second, the Dendroctonus group of genera (Hylurgus, Tomicus, Dendroctonus, Hylur- drectonus , and Pachijucotes). The first group appears to have radiated in a wide variety of hosts from South America and Australia since the beginning of the Tertiary. The second group appears to have been associated an- ciently with Araucaria hosts and is sparsely, uniformly represented in major geographical areas, except for Africa, in modern Araucaria and other Pinaceae. Key to the Genera of Tomicini 1. Metepisternal setae scalelike or plumose; an- tennal funicle 7-segmented 2 — Metepisternal setae usually hairlike (bifid in one Hy/iirgonofus having a 6-segmented funi- cle, and scalelike in Xijlechinus having a 5- segmented funicle); antennal funicle 4- to 7- segmented 8 2(1). Anterolateral areas of pronotum distinctly as- perate (minute in India species); antennal club apparently with either tivo or four trans- verse sutures; male frons strongly impressed . 3 — Anterolateral areas of pronotum unarmed; an- tennal club with three sutures clearly marked; male frons impressed or not 4 3(2). Antennal club more strongly flattened, more slender, at least 2.0 times as long as wide, apparently with two sutures; frontal rectangle at least as wide as long (0.8-1.0 times); Europe and Asia to Australia and New Zealand; mostly in broadleaf hosts; 1.8-5.0 mm Chaetoptelius — Antennal club less strongly flattened, stouter, less than 1.5 times as long as wide, apparently with four sutures; frontal rectangle longer than wide (about 1.2 times); South America; Araucaria; 1.5-3.4 mm Xylechinosomus 4(2). Male frons strongly impressed; median fron- tal carina present; pronotum either without a constriction affecting dorsal profile (Sino- phloeus) or with a moderate lateral constric- tion on anterior third (Dendrotrupes) 5 — Male frons convex to modestly flattened, me- dian carina present or absent; pronotum with a conspicuous transverse constriction on ante- rior third affecting dorsal profile 6 5(4). Antennal club elongate, 2.0 times as long as wide, apparently with four or five sutures; 1986 WOOD: Genera of Scolytidae 41 M 'w Fig. 35. Xylechinus montanus Blackman, dorsal aspect (After Bright 1976: 206. declivital interstriae 2 impressed, 3 (some- times 1-7) armed by rounded tubercles; pronotum conspicuously wider than long (0.7 times as long as wide); larger; South America; Nothophagus ■ 2.3-3.0 mm Sinophloeus Antennal club more compact, less than 1.5 times as long as wide, with three sutures; declivity variable; pronotum almost as long as wide (0.9 times); New Zealand; hosts not coniferous; 1.5-2.0 mm Dendrotrupes Elytral vestiture hairhke, ground vestiture moderately stout; frontal carina absent; an- tennal club slightly flattened, segment 1 one- fourth of its length; North America; Ulmus, 2.0-2.5 mm Hylurgopinus Vestiture of abundant, conspicuous scales (one exception with no ground setae); median frontal carina usually present; antennal club more conical 7 6(4). 7(6). Strial punctures small, close; interstriae two or more times as wide as striae, unarmed (except coarsely tuberculate and without ground vestiture in rugatus), erect setae closer and coarser; elytral ground setae more slender and appically pointed, variegated pat- terns obscure; antennal club somewhat more flattened and apex less pointed; China; broadleaf hosts; 1.8-3.0 mm . Pseudoxylechinus — Strial punctures coarse; interstriae less than 1.5 times as wide as striae, often armed by tubercles of moderate size; elytral ground se- tae usually stout, apically rounded, and form- ing conspicuous variegated patterns; antennal club more nearly conical, segment 1 usually conspicuously longer; North America; conif- erous hosts; 2.2-5.8 mm Pseudohylesinus 8(1). Ground vestiture on elytra scalelike, metepisternal setae scalelike (Fig. 35); anten- nal funicle 5-segmented; median frontal carina present (except absent in a few South American species); procoxae rather widely separated; North and South America, Eu- rope, Asia, Africa, Australia; coniferous and broadleaf hosts; 1.5-3.5 mm Xylechinus — Ground vestiture on elytral disc hairlike, metepisternal setae hairlike (except some Hy- lurgonotus . female Hylurdrectonus) 9 9(8). Protibiae armed by five or more socketed teeth on distal and lateral margins; male frohs convex except in Hylurdrectonus, phloeo- phagous 10 — Protibiae armed by three socketed teeth (four in two Hylurgonotus) on distal margin; male frons feebly to strongly, extensively exca- vated; xylophagous in Araucaria 13 10(9). Antennal funicle 6-segmented, club conical . 11 — Antennal funicle 5-segmented, club moder- ately flat 12 11(10). Procoxae contiguous; pronotum more slen- der, 0.95-1.1 times as long as wide, only slightly constricted on anterior thrid; erect interstrial setae abundant, confused; a short median carina from epistomal margin to level of antennal insertion; Europe, W Asia; Finns; 3.1-5.3 Hylurgus — Procoxae moderately separated; pronotum stouter, less than 0.85 times as long as wide, strongly constricted on anterior third; erect interstrial setae in uniseriate rows (except confused in puellus), a fine median carina from epistoma to middle of frons (absent in puellus), Europe, Asia, N Africa; 2.5-4.5 mm Tomicus 12(10), Antennal club with sutures somewhat pro- curved; procoxae contiguous; male frons con- vex to weakly impressed; vestiture never scalelike in either sex; North America, Eu- rope, Asia; Pinus, Picea, Larix, Pseudotsnga; 2.5-9.0 mm Dendroctonus — Antennal club with sutures straight, trans- verse; procoxae moderately separated; male frons rather strongly concave; female elytra with some scales (except araucariae): Aus- tralia, New Guinea; Araucaria; 1.3-1.8 mm Hylurdrectonus 42 Great Basin Naturalist Memoirs No. 10 13(9). Declivital vestiture hairlike (except scalelike in tuberculatus), protibiae with either three or four socketed teeth; male frons rather shal- lowly impressed (except deeper in an- tipoditis); South America; Araucaria; 2.7-4.6 mm Hylurgonotus — Declivital vestiture always includes scales (sparse in male peregrinus); protibiae with three socketed teeth; male frons strongly, more extensively impressed (except weak in peregrimts), Australia, New Zealand, and ad- jacent islands; Araucaria; xylophagous; 2.3- 4.5 mm Pachycotes Chaetoptelius Fuchs [in Reitter 1913: 43, Type-species: Hylesinus vestittis Mulsant & Rey, automatic. Synonyms: Homarus Broun 1881: 740, Type-.species: Homarus mundidns Broun, automatic, preoccupied; Acrantus Broun 1882: 409, Type-species: Homonis mundidus Broun, automatic, preoccupied; Chaetophorus Fuchs 1912: 46, Type-species: Hylesinus vestitus Mulsant & Rey, monoba- sic, preoccupied]. Distribution: 1 species in Europe and W Asia, 1 in New Zealand, about 8 in Australia and New Guinea. All are phloeophagous and monogamous. Xylechinosomus Schedl [1963a: 209, Type- species: Xylecliinus taunayi Eggers, original designation]. Distribution: About 9 species in South America. All are phloeophagous in Araucaria . Sinophloeus Brethes [1922b: 433, Type- species: Sinophlocus pui-teri Brethes, mono- basic]. Distribution: 2 species in South Amer- ica. Apparently phloeophagous in Notho- fagus. Dendrotrupes Broun [1881: 741, Type-spe- cies: Dendrotrupes costiceps Broun -Den- drotrupes vestitus Broun, subsequent desig- nation by Hopkins 1914: 120]. Distribution: 3 species in New Zealand. Phloeophagous. Hylurgopinus Swaine [1918: 43, 74, Type- species: Hylastes rufipes Eichhoft, original designation]. Distribution: 1 species in North America. Phloeophagous in Uhnits and mon- ogamous. Pseudoxylechinus Wood & Huang [1986: 465, Type-species: Pseudoxylechinus uni- formis Wood & Hwang, original designation]. Distribution: 7 species in Asia. All are phloeophagous and monogamous in broadleaf trees. Pseudohylesinus Swaine [1917: 11, Type- species: Pseudohylesinus grandis Swaine = Hylurgus sericeus Mannerheim, original designation]. Distribution: 11 species in North America. Phloeophagous in Abies, Picea, Pinus, Pseudotsuga, and Tsuga and monogamous. Kevs: Blackman (1942a; 5), Bright (1969: 1.5), Wood (1982: 1.30). Xylechinus Chapuis [1869: 36, Type-spe- cies: Hylesinus (Dendroctonus) pilosus Ratzeburg, monobasic. Synonyms: Prunipha- gus Murayama 19.58: 930, Type-species: Pruniphagus gununensis Murayama, original designation; Squamasinulus Nunberg 1964: 431, Type-species: Squamasinulus chiliensis Nunberg, original designation; Xylechinops Browne 1973: 283, Type-species: Xylechinus australis Schedl, original designation]. Distri- bution: 19 species in Central and South Amer- ica, 2 in North America, 2 in Africa, 5 in Asia, and 1 in Europe. All are phloeophagous and monogamous. Keys: Blandford (1897: 157), Wood (1982: 143) for North and Central America. Hylurgus Latreille [1807; 274, Type-spe- cies: Hylesinus ligniperda Fabricius, monoba- sic]. Distribution: 3 species in Europe, W Asia, N Africa. All are phloeophagous in conif- erous hosts and monogamous. Totnicus Latreille [1802/3: 203, Type-spe- cies: Hylesinus piniperda Fabricius =Der- mestes piniperda Linnaeus, monobasic. Syn- onyms: Blastophagus Eichhoff 1864: 25, Type-species: Dennestes piniperda Lin- naeus, preoccupied, subsequent designation by Lacordaire 1866: 360; Myelophilus Eich- hoff 1878c: 400, Type-species: Dermestes piniperda Linnaeus, automatic]. Distribu- tion: 8 species in Europe and Asia. All are phloeophagous in Pinus and are monoga- mous. Keys: Murayama (1963: 35) for the Far East, Balachowsky (1949: 135) for France, Schedl (1946b: 52) for the genus. Dendroctonus Erichson [18.36: 52, Type- species: Bostrichus micans Kugelann, subse- quent fixation by International Commission on Zoological Nomenclature 1974: 230]. Dis- tribution: 16 species in North America and 2 in Eurasia. All are phloeophagous in Pinus, Picea, Larix, and Pseudotsuga and are monogamous. Keys: Hopkins (1909: 69), Wood (1963: 26, 1982: 151). Hylurdrectonus Schedl [19.3Sb: 40, Type- species: Hylurdrectonus piniarius Schedl, monobasic. Synonym: Xylogopinus Schedl 1972: 64, Type-species; Xylogopinus araucar- 1986 WOOD: Genera of Scolytidae 43 iae Schedl =Hyliirdrectoiuis corticinus Wood, monobasic]. Distribution: 3 species in Australia and New Guinea. Phloeophagous in Araucaria except one in central a.xis of leaflet; monogamous. Hylurgonotus Schedl [1951e: 448, Type- species: Hyhirgonotus bninnetis Schedl =Hylurgus titbercidatiis Eggers]. Distribu- tion: 4 species in South America. In Arau- caria, possibly xylophagous and apparently monogamous. Pachycotes Sharp [1877: 10, Type-species: Pacycotes ventralis Sharp =Hylesmus pere- grinus Chapuis, monobasic]. Distribution: 8 species in Australia, New Zealand, and New Guinea. All are xylophagous in logs and monogamous. Tribe Phrixosomini Phrixosomini Wood [1978: 111, Tvpe-genus; Phrixosoma Blandford, 1897] Description. — Frons not sexually dimor- phic, frequently with a fine, median carina; eye completely divided; antennal scape elon- gate, funicle 6-segmented, club rather strongly flattened, slightly asymmetrical, un- marked by sutures, except 1 partly septate; pronotum unarmed by asperities, procoxae contiguous; scutoscutellar suture remote from scutellar groove, postnotum separated from scutoscutellar area of metanotum by a com- plete suture; tibiae armed on lateral margin by socketed denticles. Biology. — These monogamous, phloeo- phagous, tropical species are restricted to hosts of the Guttiferae. The parental galleries are usually biramous, although a third egg tunnel is not uncommon, with the galleries either longitudinal, transverse, or without definite orientation. The eggs are deposited in niches and sealed in by frass. The larval mines show on the inner surface of peeled bark, but they are almost entirely in the bark and usu- ally wander indiscriminantly without respect to the grain of the wood. Taxonomy. — Only one genus is known (Fig. 36). It is apparently very old and repre- sents a group that is otherwise extinct. They are unknown in the fossil record. The one genus in South America and Africa in Gut- tiferae has changed so little since separation of these land masses that division into species groups is not recommended. Although more Fig. 36. Phrixosomu magna Blackmail, antenna. closely allied to Hylesinini than to other tribes, this genus is quite unique. Phrixosoma Blandford [1897: 148, Type- species: Phrixoaoma rude Blandford, mono- basic. Synonyms: Bothryperus Hagedorn 1909: 742, Type-species: Bothryperus psaltes Hagedorn, monobasic; Neohylesinus Eggers 1920b: 118, Type-species: Neohylesinus quadrioculatus Eggers, monobasic; Sphaero- sinus Eggers 1929: 40, Type-species: Sphaerosinus striatus Eggers, monobasic]. Distribution: 13 species in Central and South America and 9 in Africa. All are phloeo- phagous in Guttiferae and are monogamous. Key: Wood (1982: 204). Tribe Hyorrhynchini Hyorrhynchinae Hopkin.s [1915b: 225, Type-genus: Hy- orrhynchus Blandford, 1894] Sueinae Murayama [19.58: 7, Type-genus: Siieus Mu- rayama, 1951] Description. — Frons sexually dimorphic, male slightly to strongly impressed, female convex; eye completely divided; antennal scape either long or short, funicle 5- or 6-seg- mented, club aseptate and either unmarked by sutures or with two sutures; pronotum armed or not; procoxae moderately to widely separated, precoxal ridge obsolete; scu- toscutellar suture remote from scutellar groove; scutoscutellar area separated from postnotum by a distinct suture; tibiae without socketed denticles. BiOLCJGY. — The species are myelomyce- tophagous or xylomycetophagous and form simple, monoramous tunnels in the pith or xylem of twigs or small branches. Only two larvae were seen (Sueus niisimai) and these were in the parental chamber. Males oi Sueus are very rare, dwarfed, and deformed; repro- duction in tliis genus is apparently by 44 Great Basin Naturalist Memoirs No. 10 arrhenotocous parthenogenesis. In Hyor- rhtjnchus and Pseudohijorrhtjnchus the males are similar in size, shape, and abundance to the females, suggesting a normal bisexual re- lationship. Taxonomy. — A dozen species assigned to three genera are listed in the literature. All occur in the area from India and Japan to New Guinea. They form an aberrant, relect group of uncertain affinity but are considered to be among the most primitive of the Hylesininae. Although definitely members of the Hylesini- nae, they share more primitive characters with primitive Scolytinae and Platypodidae than with other members of their own sub- family. Key to the Genera of Hyorrhynehini 1. Antennal funicle 5-segmented, club rather weakly compressed; male dwarfed, rare, flight- less; male frons convex, with median carina; tarsal segment 3 entire; Sri Lanka (Ceylon) and New Guinea to Japan; female L 6-2.0 mm. male 1.0-L3 mm Sueus — Antennal funicle 6- or 7-segmented, club rather strongly flattened; male subequal in size to fe- male, shares parental gallery; male frons con- cave or not, with or without a carina; tarsal seg- ment 3 emarginate to bilobed 2 2(1). Antennal funicle 6-segmented, club with two clearly marked sutures; male frons concave, without a carina; tarsal segment 3 emarginate; interstrial tubercles minute; India to Japan. 2.7- .5.0 mm Hyorrhynchus — Antennal funicle 7-segmented, club wdthout su- tures; tarsal segment 3 deeply bilobbed; male frons not impressed, with a median carina; inter- strial tubercles on declivity rather coarse; Japan; 3.5-3.6 mm Pseudohyorrhynchus Sueus Murayama [1951; 1, Type-species: Sueus sphaerotrypoides Murayama = Hyor- rhynchus niisimai Eggers, original designa- tion. Synonym: Neohyorrhynchus Schedl 1962e: 202, Type-species, Hyorrhynchus ni- isimai Eggers, original designation]. Distri- bution: 2 species from Sri Lanka (Ceylon) and New Guinea to Japan. Xylomycetophagous in small branches and, apparently, at least partly parthenogenetic (only one series of males known). Hyorrhynchus Blandford [1894a: 58, Type- species: Hyorrhynchus lewisi Blandford, monobasic]. Distribution: About 10 species from India to Japan. Habits have not been reported except host species include Acer, Fagus , and Macaranga; the species are xy- lomycetophagous and monogamous (Nobu- chi, pers. comm.). Pseudohyorrhynchus Murayama [1950b: 61, Type-species: Pseudohyorrhynchus wa- dai Murayama, original designation]. Distri- bution: 1 species from Japan. Specimens of this monogamous species attack living Corniis twigs where they are xylomycetophagous. Tribe Diamerini Diameridae Hagedorn [1909: 734, Type-genus: Di- umerus Erichson, 1836] Strombophorini Schedl [1959e: 75, Type-genus: Strom- hophonis Hagedorn, 1909] Sphaerotrypini Murayama [1963: 5, Type-genus: Sphaerotrypes Blandford, 1894] Description. — Frons dimorphic, feebly to very strongly impressed in male, flattened to convex in female; eye entire to completely divided; antennal scape usually elongate, fu- nicle 6- to 7-segmented, club flattened, su- tures variable, one to several (five or more, mostly pseudosutures); procoxae moderately to widely separated; scutoscutellar suture re- mote from scutellar groove, postnotum fused to scutoscutellar area of metanotum, interseg- mental suture obsolete on median half. Biology. — These monogamous, phloeo- phagous species are largely restricted to tropi- cal and subtropical areas of the eastern hemi- sphere. The parental galleries are mostly biramous and either longitudinal or trans- verse. The eggs are deposited in niches and sealed in by frass. The larval mines show on the inner surface of peeled bark and tend to radiate away from the parental tunnel without respect to the grain of the wood in most spe- cies; the latter parts of the tunnels in some species are parallel to the grain of wood. Taxonomy. — Seven genera represented by about 122 species occur in Africa and south- eastern Asia to Australia. Pseudodiamerus, Pernophorus . and Stromhophorus occur ovAy in Africa. They appear to form a sister group derived from the same ancestral stock as Both- rosternini and, if this is the case, these groups have differentiated and radiated since early Tertiary. Features used to characterize this tribe vary and tend to intergrade with Phloeosinini. Although future study may re- quire combination of these two tribes, their division gives a convenient break in a large and diverse group. 1986 WOOD; Genera of Scolytidae 45 Key to the Genera of Diamerini 1. Eye oval, neither strongly reduced on lov\er half nor completely divided; antennal club with fewer than three sutures, ftrnicle 5-, 6-, or 7-seg- mented 2 — Eye either strongly narrowed on lower half or entirely divided into two parts; antennal club apparently with five or more sutures marked by constrictions and/or rows of setae, funicle 6-seg- mented (except 7-segmented in Pernop/iorus) . 5 2(1). Costal margin on basal fourth of elytra normal, not emarginate, metepisternum not expanded, median anterior area of pronotum usually finely asperate; antennal funicle 6-segmented; club elongate, sutures 1 and 2 feebly indicated (straight) to absent; .Mrica, SE Asia to Australia, 1. 1-2.5 mm Acacicls — Costal margin on basal fourth of elytra either normal or deeply, abruptly emarginate, when emarginate metepisternum expanded into this emargination; pronotum never asperate (feeble lateral asperities in Bothrosternoidcs) .3 3(2). Costal margin on basal fourth of elytra deeply emarginate (Fig. 37); suture 1 on antennal club strongly, subangulately procurved; Africa, SE Asia to Australia; 3.0-.5.0 mm Diamerus — Costal margin on basal half of elytra almost nor- mal, straight to wealily emarginate; basal mar- gins of elytra either costate or armed by a row of coarse crenulations; protibia with two to four curved spines; male frons shallowly impressed; suture 1 on antennal club weakly indicated (straight) or absent 4 4(3). Eye more than 3 times as long as wide; funicle 7-segmented; club with suture 1 straight, fee- ble; protibia with two apical and two lateral spines; costal margin not emarginate; basal mar- gins of elytra crenulate; pronotum without lat- eral granules; Africa (Angola); 2.2-3.0 mm .... Pseudodiameriis — Eye twice as long as wide; funicle 5-segmented, club unmarked by sutures; protibia with two apical and one small posterior subapical spine and several lateral posterior granules; costal margin shallowly emarginate at base; basal mar- gins of elytra strongly costate; some prothoracic granules or small asperities in lateral areas; male with median epistomal tubercle; body very stout; Malaya; 2.3 mm Bothrosternoidcs •5(1). Eye completely divided; scutellum visible, longer than wide; pronotum unarmed by asperi- ties; body very stout, subglobular, Africa, SE Asia; 1.5-5.0 mm Sphaerotrypes — Eye strongly constricted on lower half to about one-third width of upper half, pronotum at least partly asperate; scutellum not visible; body elongate-oval 6 6(5). Antennal funicle 7-segmented; elytral vestiture hairlike to subplumose, Africa; 4.0 mm Pernophonis Fig. 37. Diamerus curvifer Walker, lateral aspect of fe- male. — Antennal funicle 6-segmented; elytral vestiture with ground cover scalelike; Africa; 1.5-3.5 mm Strombophortis Acacacis Lea [1910: 149, Type-species: Acacacis ahundans Lea, monobasic. Syn- onyms: Trogloditica Sampson 1922: 148, Type-species: Trogloditica trahax Sampson, monobasic; Pseudoacacacis Schedl 1963c: 477, Type-species: Acacacis horneensis Browne, original designation; Neodiamerus Schedl 197ib: 282, Type-species: Ncodi- amcrus granidicoUis Schedl, original designa- tion]. Distribution: 5 species in Africa, 2 in Sri Lanka, 2 in Malaya, 1 in Borneo, 1 in New Guinea, and 2 in Australia. All are phloeophagous and monogamous. Pseudodiamerus Eggers [1933b: 18, Type- species: Pseudodiameriis sthatus Eggers, monobasic]. Distribution: 3 species in Africa. All are apparently phloeophagous and monog- amous. Bothrosternoides Schedl [1969: 210, Type- species: Bothrosternoides malaycnsis Schedl, monobasic]. Distribution: 1 species in Malaya. Three specimens intercepted in Japan and 4 in China in imported Nyatoh logs. Diamerus Erichson [1836: 57, Type-spe- cies: Hylesinus hispidus King, monobasic. Synonyms: Acanthurus Eichhofl 1886: 24, Type-species: Acanthurus spinipcnnis Eich- hoff - Hylesinus curvifer Walker, subsequent designation by Hopkins 1914: 116; Lissoclas- tus Schaufuss 1905: 71, Type-species: Lisso- clastus pimelioides Schaufuss, monobasic]. Distriliution: About 11 species in Africa and 28 in the Indo-Australian and Oriental areas. All are phloeophagous and monogamous. Sphaerotrypes Blandford [ 1894a: 61, Type- species: Sphaerotrypes pila Blandford, subse- ()uent designation by Hopkins 1914: 129. Syn- 46 Great Basin Naturalist Memoirs No. 10 onym: Parasphaerotrijpes Murayama 1958; 933, Type-species; Sphaerotrijpes controver- sae Murayama, original designation]. Distri- bution: 8 nominate species in Africa and about 35 in the Indo-Australian and Oriental areas. Pernophorus Strohmeyer [1910b; 92, Type-species; Acanthophorus hrevicollis Strohmeyer, automatic. Synonym: Acan- thophorus Strohmeyer 1910a; 69, Type-spe- cies; Acanthophorus hrevicollis Strohmeyer, monobasic, preoccupied]. Distribution; 5 nominate species in Africa. They are appar- ently phloeophagous and monogamous. Strombophorus Hagedorn [1909: 740, Type-species: Strombophorus crenatus Hagedorn, subsequent designation by Hop- kins 1914: 130]. Distribution; About 34 spe- cies in Africa. They are phloeophagous and monogamous. Tribe Bothrosternini Bothrosterni Blaiidford [1896a; 120, Type-genus: Both- rostenius Eichhoff, 1868] Description. — Frons usually sexually di- morphic; eye entire to slightly sinuate; funicle 6-segmented, club symmetrical, moderately flattened, sutures indicated; proco.xae moder- ately separated; protibia bearing a bifid pro- cess on outer apical angle exceeding inner apical angle; pronotum unarmed; crenula- tions on basal margins of elytra poorly devel- oped, sometimes represented by a continuous costa; at least part of scutoscutellar suture par- allel to margin of scutellar groove; postnotum fused to scutoscutellar area of metanotum, suture obsolete. Biology. — All species are monogamous, except that a form of parthenogenesis proba- bly exists in Bothrosternus . All are myeloph- agous, except that Bothrosternus and, possi- bly, Eupagiocerus ater Eggers are myelo- mycetophagous. In Cnesinus annectens Wood typical transverse, biramous egg gal- leries are formed in the cambium region of the host by parent beetles, complete with egg niches, and the larvae feed briefly in the cam- bium region before following the parent beetles to the pith region. Typically, the parental pith tunnel is biramous; the eggs are deposted in clusters in the loose frass in this cavity; and the larvae feed in congress while extending the parental pith tunnel. Pupation occurs in the larval frass. Ta.xonomy. — The tribe is restricted to the American tropics, with two species extending into the southern United States. It appears to be the New World counterpart of the closely allied Diamerini. The genera of Bothrostern- ini differ anatomically from one another only slightly and may be recognized with difficulty in some instances. The protibial and antennal structure and the costate basal margins of the elytra in at least some members suggest a very primitive position in classification. Though primitive as a group, modern representatives appear to have evolved since the beginning of the Tertiary to their present ecological and structural status, because clearly identifiable related groups are not known outside of the neotropical realm. They appear to be more nearly allied to Diamerini and Hyorrhynchini than to any American groups. Key to the Genera of Bothrosternini 1. Lateral margins of pronotum rounded 2 — Lateral margins of pronotum marked by a sharply elevated, costate to subcostate line .... 3 2(1). Sutures of antennal club transverse, straight; rostrum distinctly wider than distance between eyes; pronotum either longitudinally strigose or punctured; pith borers of twigs and other small stems; SE USA to South America; 1.6-3.5 mm Cnesisnus — Sutures of antennal club strongly procurved; rostrum width at tip equal to distance between eyes; frons excavated in both sexes, with a me- dian tubercle just above epistoma; body oval; seed borers; SE USA to South America, inter- cepted elsewhere in maize, etc. ; 1.9-2.6 mm . . Pagiocerus 3(1). Sutures of antennal club strongly procurved; pith borers in twigs and woody vines; Central and South America; 2.1-3.8 mm . . . Eupagiocerus — Sutures of antennal club transverse, straight ... 4 4(3). Proepisternal area partly excavated, with cavity densely filled by yellow pubescence, particu- larly in female; prothoracic intercoxal piece with a transverse, subcarinate ridge; elytral inters- triae usually not strongly carinate; ambrosia beetles in axial tunnels of woody vines; Central and South America; 1.9-3.3 mm . . Bothrosternus — Proepisternal area normal, not densely pubes- cent; prothoracic intercoxal ridge absent; elytral interstriae narrowly carinate from posterior part of disc to apex; pith borers in branches and twigs; Central and South America; 2.2-3.6 mm Sternobothrus Cnesinus LeConte [1868: 171, Type-spe- cies; Cnesinus strigicoUis LeConte, monoba- sic. Synonym; Neinophihis Chapuis 1869; 27, 1986 WOOD; Genera of Scolytidae 47 Fig. 38. Bothrosternini; 1-8, Cnesinus spp. , (1) cubensis Blackman, dorsal aspect, (2) same, female head, (3) panainen- sis Blackman, dorsal aspect, (4) same, male head, (5) robai Blackman, male head, (6) setulosus Blandford, female head, (7) costulatus Blandford, dorsal aspect, (8) same, female head, (12) hlachmani Schedl, dorsal aspect; 9-11, Bothroster- nusfoveatus Blackman, (9) dorsal aspect of female, (10) female head, (11) propleuron of female. (After Blackman 1943: pi, 15). 48 Great Basin Naturalist Memoirs No. 10 Type-species: Nemophilus strigillatiis Cha- puis =Cnesinus strigicoUis LeConte, subse- quent designation by Hopkins 1914: 125]. Distribution: About 101 species from the USA to Argentina. All are monogamous and myelophagous except that one is partly phloeophagous. Keys: Wood (1968b: 88, 1982: 209). Pagiocerus Eichhoff [1868a: 148, Type- species: Pagioccrus rimosus Eichhoff = Bostrichus frontalis Fabricius, subsequent designation by Hopkins 1914: 126]. Distribu- tion: About 5 species are known from the USA to Argentina, 1 is occasionally intercepted worldwide in large seeds, including maize. All are monogamous and spermophagous. Eiipagiocerus Blandford [1896d: 1.33, Type-species: Eupagiocerua dentipes Bland- ford, monobasic. Synonym: Nemopagiocerits Schedl 1962a: 85, Type-species: Eupagio- ceriis nevermanni Schedl =Eupagiocerus ater Eggers, monobasic]. Distribution: 4 spe- cies from Mexico (Chiapas) to Venezuela and Peru. All are monogamous and myelophagous in woody vines. Keys: Wood (1965: 31, 1982: 249). Bothrosternus Eichhoff [1868a: 150, Type- species: Bothrostenms truncatus Eichhoff, monobasic]. Distribution: About 12 species from Mexico (Veracruz) and Jamaica to Peru and Brazil. Monogamous and some species apparently with a type of parthenogenesis (in- volving consanguineous polygyny), and myelophagous in woodv vines. Kev: Wood (1982: 247). Sternobothrus Eggers [1943: 372, Type- species: Bothrosternus cancellatus Chapuis, original designation]. Distribution: About 10 species from Costa Rica to Bolivia and Brazil. Monogamous and myelophagous, three spe- cies breed in Nectandra branches. Key: Wood (1982: 254). Tribe Phloeotribini Phloeotribidae Chapuis [1869: 42, Type-genu.s: Phloeo- tribus Latreille, 1796] Phthorophloeides Niisslin [1912b: 273, Type-genus: Ph- thoropliloeus Rev, 1883] Description. — Frons se.xually dimorphic, male variously impressed, female flat to con- vex; eye entire; funicle 5-segmented, club al- most non-existent to strongly asymmetrical, deeply divided into three movable, sub- Fig. 39. Phloeotiibtis pilula Eiichson, dorsal aspect of male. (After Schedl 1953: 80). lanimelate segments; procoxae contiguous; pronotum armed or not, its lateral margins rounded; metatergum fused to its postnotum. Biology. — All are monogamous and phloeophagous. Parental galleries are bi- ramous and engrave the wood rather deeply. Eggs are deposited in niches packed in frass. Larval mines follow a rather definite course away from the parental tunnels and usually do not cross one another; in the latter stages they may engrave the wood rather deeply. A few species bore rather deeply into subsurface tis- sues of woody vines; one species breeds in the fruiting pods of its host (Inga). Taxonomy. — The tribe apparently origi- nated in South America, where a majority of the species now occur. Aricerus and one very primitive Phloeotribiis apparently reached Australia very early; much later a few species of Phloeotribiis reached North America and then spread from there to northern Asia, Eu- 1986 WOOD; Genera of Scolytidae 49 rope, and North Africa. The tribe apparently was derived from the same parental stock as the Phloeosinini, although the relationship is not close. The distribution, freely movable antennal club segments (Fig. 39), and the tib- ial structure of Aricerus and at least one spe- cies oi Phloeotribus suggest an origin in early Tertiary or late Cretaceous while it was still possible to spread to Australia, but not into Africa. The one Australian and a few Eu- ropean Phloeotribus (Fig. 27) have virtually no club on the antenna, suggesting the possi- bility that the club of Scolytidae could have been derived independently from that of other curculionoids. Key to the Genera of Pliloeotribiiii 1. Lateral margin of protibia without .socketed teeth, outer apical angle rather strongly produced, with few (about three) major serrations; lateral margin of pronotum acutely elevated, subserrate; anten- nal club rather slender, almost symmetrical, ven- tral margins of segment.s not noticeably extended; phloeophagous; Australia to New Guinea; .3, 1-4.6 mm Aricerus — Lateral margin of protibia with several socketed teeth, none strongly projecting beyond others (one exception); lateral margins of pronotum rounded; ventral or lateral margin of each antennal club segment weakly to profoundly extended into a .sublamellate process except in very primitive species; phloeophagous; North and South Amer- ica, Europe, N Asia, N Africa, Australia Phloeotribus Aricerus Blandford [1894b: 1.33, Type-spe- cies: Aricerus chapuisi Blandford, subse- quent designation by Hopkins 1914; 117. Syn- onym: Hylesinosoma Lea 1910: 143, Type- species: Hylesimis fici Lea =Aricerus eich- hoffi Blandford, monobasic]. Distribution: 3 species from Australia to New Guinea. Monogamous and phloeophagous in Ficus limbs. Phloeotribus Latreille [1796: 50, Type-spe- cies: Hylesimis oleae Fabricius =Scolytus scarabaeoides Bernard, monobasic. Syn- onyms: Phloiotribus Latreille, 1796: 50, inad- vertent error in original spelling amended in Latreille 1804: 108, ruled i^y International Commission on Zoological Nomenclature 1979: 132; Phloeophthorus Wollaston 1854: 299, Type-species: Phloeophthorus perfolia- tus Wollaston, monobasic; Dryotomus Cha- puis 1869: 46, Type-species: Dryotomus pu- berulus Chapuis, monobasic; Phthorophloeus Key 1883: 128, Type-species: Phloeophthorus spinulosus Rey, monobasic; Elzearius Guille- beau 1893: 64, Type-species: Elzearius crena- tus Guillebeau, monobasic; Eulytocerus Blandford 1897; 161, Type-species: Eulyto- cerus championi Blandford, monobasic; Comesiella Del Guercio 1925: 218, Type-spe- cies: Comesiella sicula Del Guercio = Bostriclms brevicollis kolenati, monobasic; Neophloeotribus Eggers 1943: 349, Type-spe- cies: Phloeotribus nubihis Blandford, desig- nated by Wood 1983: 648; Dryotomicus Wood 1962: 76, Type-species: Dryotomus pu- berulus Chapuis, automatic]. Distribution: About 58 species in South America, 27 in North and Central America, 10 in Europe, 4 in N Africa, 1 in Australia. All are monoga- mous and phloeophagous. Keys: PfefFer(1972: 31) for Europe, Blandford (1897: 162) for Cen- tral America, Wood 1982: 257) for North and Central America. Tribe Phloeosinini Phloeosinides Niisslin [1912b: 273, Type-species: Phloco- ninus Chapuis, 1869] Description. — Frons usually dimorphic, male impressed, female flat to convex; eye varying from entire to emarginate to com- pletely divided; antennal iunicle 5- to 7-seg- mented, club flattened, slightly to strongly asymmetrical, with or without sutures; prono- tum armed or not; tarsal segment 3 com- pressed to broad and bilobed; scutelhun visi- ble or not; metanotum fused to postnotum. Biology. — All are monogamous except for a few species of polygynous Olonthogaster and one bigynous Chramcsus . Most are phloeophagous, although Dendrosinus and three species of Chramesus are xylophagous and Hyleops larvae become xylomyce- tophagous in the later stages. The parental tunnels are mostly monoramous, with a con- spicuous turning niche, a few are rather prim- itively (unequally) biramous. Eggs are placed in niches and packed in frass. Lai'val mines tend to follow a definite course away from the parental tunnel and rarely cross one another. Taxonomy. — This tribe appears to consist of a diverse assemblage of somewhat distantly related genera or clusters of genera that ap- pear to be relicts from a former much larger group. Their worldwide distribution, diverse structure, and possession of several primitive traits suggest an ancient origin that extends 50 Great Basin Naturalist Memoirs No. 10 '^/' Fig. 40. Phloeosinus punctatus LeConte, dorsal aspect of male. (After Bright 1976: 207). well into the Cretaceous. Within the tribe, two clusters of genera are apparent. Pseu- dochramesus and Chramesus (America) are closely related to one another and are the most highly evolved from a structural point of view. Cladoctonus, Phloeosinopsoides, Olon- thogaster , and Phloeosinus (Fig. 40) form a second cluster of genera to which Phloeo- cranus (Fig. 41) and Phloeodictica might form a primitive base. All are Old World genera except that about half of the Cladoctonus spe- cies occur in tropical America and a segment oi Phloeosinus has extended into North Amer- ica in comparatively late Tertiary time. The Cladoctonus species of the Philippines (1), Africa (8), and tropical America (6) have appar- ently changed only slightly since attaining their present generic distribution (perhaps early Tertiary). The American Dendrosinus and Carphotoreus and the Australian Hijleops are not closely related to one another or to the other known generic groups of this tribe. Fig. 41. Phloeocranus bruchoides Schedl, lateral aspect of female (tibial denticles omitted). Key to the Genera of Phloeosinini 1 . Interstriae 10 continued to declivity (to level of visible sternum 3); protibia with lateral apical angle produced slightly and armed by three socketed teeth; humeral angles of elytra strongly produced cephalad, largest crenula- tions at humeral angles; eye deeply emarginate (three-fourths divided); funicle 5-segmented; scutellum not visible; phloeophagous; India to Malaya; 2. 5-3 5 mm Phloeocranus — Interstriae 10 not continued behind level of metacoxae; lateral apical angle of protibia not produced (except Phloeoditica ) 2 2(1). Protibia slender, outer apical angle armed by two closely set, projecting, socketed denticles, one smaller tooth on lateral margin; procoxae rather widely separated by distance equal to width of coxa; funicle 5-segmented, club sym- metrical to moderately asymmetrical; prono- tum unarmed; scutellum not visible; elytral ground vestiture of circular (rounded) scales; metathoracic wings apparently absent in male of at least one species; phloeophagous; SE Asia; 2.2-2.5 mm Phloeoditica — Protibia more strongly flattened, armed by three or more socketed teeth of equal size; with other combination of characters 3 3(2). Antennal club symmetrical to rather weakly asymmetrical (Fig. 42); eye emarginate, or di- vided when funicle 5-segmented; procoxae ei- ther separated or contiguous; body usually more slender 4 — Antennal club very strongly asymmetrical (Fig. 43). sutures strongly procurved when present; eye entire; funicle 5-segmented; procoxae rather widely separated; body usually very stout; American species 11 4(3). Eye entire; funicle 6- or 7-segmented; prono- tum armed or not 5 — Eye weakly emarginate to completely divided; funicle 5-segmented (6-segmented in some Cladoctonus}; pronotum unarmed by asperi- ties (except weakly armed in Phloeosinop- soides) 8 1986 WOOD; Genera of Scolytidae 51 5(4). Funicle 7-segmented; bases of elytra produced anteriorly, each strongly procurved; scutellum visible, elytra! notch for its reception very deep, narrow; procoxae very widely separated; pronotum armed or not; body stout, usually black; xylophagous; Mexico to South America; 3.0-5.0 mm Dendrosinus — Funicle 6- or 7-segmented; elytral bases not produced; scutellar notch at sutural base of elytra not unusually deep or narrow; procoxae contiguous to moderately separated; smaller . . 6 6(5). Procoxae contiguous; pronotum unarmed; fu- nicle 7-segmented. scutellum obsolete; tarsal segment 3 broad, strongly bilobed; phloeo- phagous except larx'ae xylomycetophagous in later stages; Australia; Araucaria , 2.4-4.0 mm Hyleops — Procoxae separated by about half width of a coxa; funicle 6-segmented; pronotum armed by fine asperities; scutellum visible; tarsal seg- ment 3 narrow; phloeophagous 7 7(6). Body more slender, 2. 1 times as long as wide, antennal club almost symmetrical, three su- tures indicated, 1 partly septate; posteromesal margin of pronotum almost straight; Mexico; AlniiSi 2.5-2.7 mm Carphotoreus — Body stout, 1.6 times as long as wide; antennal club strongly asymmetrical, suture 1 septate, others not indicated; posteromesal margin of pronotum slightly extended toward scutellum. Congo; 1.8 mm Catenophorus 8(4). Antennal club subglobular, sutures obsolete or indicated by sparse setae; funicle 5- or 6-seg- mented; procoxae contiguous; scutellum visi- ble or not; phloeophagous; tropical America, Africa, Phihppines; 1.4-2.5 mm .... Cladoctonus — Club flattened, sutures transverse to oblique when present; funicle 5-segmented; proco.xae moderately separated; scutellum visible; phloeophagous 8 9(8). Pronotum finely asperate at least in lateral ar- eas; sutures 1 and 2 on antennal club trans- verse; eye coarsely faceted, anterior margin shallowly, broadly emarginate; elytral ground vestiture almost obsolete, erect setae in rows, flattened, almost scalelike, Taiwan to New Guinea; 1.3-2.5 mm Phloeosinopsoides — Pronotum unarmed; sutures 1 and 2 on anten- nal club oblique; eye more finely faceted, emargination at least one-third as deep as eye width (completely divided in some Asian spe- cies); elytral ground vestiture usually more abundant, erect setae slender when ground setae sparse 10 10(9). Eye usually completely divided by an emar- gination (several exceptions); tarsal segment 3 slender; protibia with two (rarely one to three) apical and subapical socketed teeth, one or two others sometimes on lateral margin; vestiture usually less abundant; male frons usually con- cave; female frons concave to convex, usually ornamented by a conspicuous brush of hair; Fig. 42-43. Antennae of Fhloeosinini: 42, above Phlocosinus thiijae (Perris); 43, below, Chraniesus hicko- riae LeConte. color usually reddish brown; SE Asia to New Guinea; mostly in nonconiferous hosts; 1.4-5.5 mm Olonthogastcr — Eye less than one-half divided by an emargina- tion; tarsal segment 3 broad, emarginate to bilobed; protibia with three or more socketed denticles on apical and subapical margin, four or more smaller teeth on lateral margin; vesti- ture more abundant; male frons usually shal- lowly impressed, female Irons convex, often with a median carina, vestiture inconspicuous in both sexes; color brown to dark brown; N Africa to N Asia, North America, Australia; mostly in coniferous hosts; 1.4-4.1 mm (PUoeosinites Hagedorn presumably fits near here) Phloeosinus 11(3). Antennal club with sutures strongly pro- curved, clearly marked by rows of setae and grooves; phloeophagous; S America; 1.3-2.0 mm Pseudochramesus — Antennal club without sutures; mostly phloeophagous; North and South America; 1 . 2- 2.7 mm Chraniesus Phloeocranus Schedl [1942: 7, Type-spe- cies: Phloeocranus bruchoides Schedl, mono- basic. Synonym: Diamerides Browne 1949; 893, Type-species: Dimnerides litseae Browne ^Phloeocranus bruchoides Schedl, original designation]. Distribution: 1 species from India to Indonesia in Litsea . This species is monogamous and phloeophagous. 52 Great Basin Naturalist Memoirs No. 10 Phloeoditica Schedl [1962d; 189, Type-spe- cies: Kissophagiis curtits Eggers, present des- ignation]. Distribution: One species in SE Asia. It breeds in Pongamia glabra. Dendrosinus Chapuis [1869: 28, Type-spe- cies: Hijulesiniis globosus Eichhoff, monoba- sic]. Distribution: 10 species from USA (Flor- ida) and Mexico (Jalisco) to Argentina. All are monogamous and xylophagous. Key: Wood (1982: 283). Hyleops Schedl [1938b: 35, Type-species: Hyleops glabratus Schedl, monobasic]. Dis- tribution: 1 species in Australia in Araucaria branches. It is monogamous and partly phloeophagous. The later larval stages pene- trate the xylem and become xylomyce- tophagous. Parental tunnels are transversely biramous and without apparent symbiotic hingi. Carphotoretis Wood [1973b: 171, Type- species: Chaetophloeus alni Bright, original designation]. Distribution: 1 species in Mex- ico (Oaxaca). It is monogamous and phloeo- phagous. Catenophorus Nunberg [1956b: 195, Type- species: Catenophorus congonus Nunberg, original designation]. Distribution: 1 species in the Congo. The habits are unknown. Cladoctonus Strohmeyer [1911: 17, Type- species: Cladoctontis affinis Strohmeyer, monobasic. Synonyms: Hoplites Eggers 1923: 140, Type-species: Hoplites banostts Eggers, monobasic, preoccupied; Hoplitontus Wood 1961: 2, Type-species: Hoplites banosus Eg- gers, automatic; Hoplitophthorus Wood 1961: 2, Type-species: Hoplitophthorus sentosus Wood = Hoplites interriiptus Eggers, original designation]. Distribution: 8 species in Africa, 6 in Cuba to Brazil and Bolivia; 1 in the Philip- pine Islands. The two species for which habits are known are monogamous and phloeo- phagous. Phloeosinopsoides Schedl [1964c: 317, Type-species: Phloeosinopsis triseriatiis Schedl, automatic. Synonym: Phloeosinopsis Schedl 1964b: 297, Type-species: Phloeo- sinopsis triseriatiis Schedl, original designa- tion, preoccupied]. Distribution: About 8 species from Taiwan to New Guinea. All are monogamous and phloeophagous. Phloeosinus Chapuis [1869: 37, Type-spe- cies: Hijlesinus thiijae Perris, subsequent des- ignation by Hopkins 1914: 126]. Distribution: 29 species in North America, about 30 in Asia and adjacent islands, 5 in Europe, 2 in N Africa, and 2 in Australia. All are monogamous and phloeophagous. Most are in coniferous hosts, especially Cupressineae. Keys: Black- man (1942c: 400) and Wood (1982: 287) for North America, Schedl (1950a: 36) for Eu- rope, Murayama (1963: 22) for Japan. Olonthogaster Motschulsky [1866: 401, Type-species: Olonthogaster nitidicollis Mot- schulsky, subsequent designation by Hopkins 1914: 126. Synonyms: H olonthogaster Gem- minger & Harold 1872: 2676, Type-species: Olonthogaster nitidicollis Motschulsky, auto- matic; Hylediits Sampson 1921: 35, Type-spe- cies: Hyledius asper Sampson, monobasic; Hylitrguliis Eggers 1927c: 392, Type-species: Hyhirgnhis summatranus Eggers, monoba- sic; Phloeosinopsis Schedl 1936a: 23, Type- species: Phloeosinopsis annatus Schedl = Phloeosinus spinifer Schedl, original desig- nation]. Distribution: About 25 species from SE Asia to Australia. All are phloeophagous and monogamous except for 2 polygynous species from New Guinea. Hosts include Lyt- sea. Myristica , etc. Phloeosinites Hagedorn [1907: 119, Type- species: Phloeosinites relii Hagedorn, subse- quent designation by Hopkins 1914: 126]. Distribution: 8 fossil species in Baltic amber (Oligocene). The relationship of this genus to Phloeosinus was not determined. Pseudochramesus Blackman [1939: 87, Type-species: Chraniesus acuteclavatus Hagedorn, original designation]. Distribu- tion: 11 species in South America. The habits have not been reported. Kev: Blackman (1939: 88). Chraniesus LeConte [1868: 168, Type-spe- cies: Chraniesus hicoriae LeConte, monoba- sic. Synonyms: Rhopalopleurus Chapuis 1869: 46, Type-species: Rhopalopleurus tu- berculatus Chapuis, subsequent designation by Hopkins 1914: 128; Thaumasinidus Reitter 1913: 39, Type-species: Dendrosinus bon- nairei Reitter =Chraniesus rotundatus Cha- puis, monobasic; Prochramesus Wood 1956b: 254, Type-species: Prochramesus anneetans Wood, original designation]. Distribution: 39 species in South America, 40 in North and Central America and adjacent islands. All are monogamous, except for the bigynous C. in- comptus, and all are phloeophagous except 1986 WOOD: Genera of Scolytidae 53 Fig. 44. Liparthrum spp.: 1, outline of dorsal aspect o{ arizomcum Wood; 2, outline of dorsal aspect oi albosctum Bright; 3, outline of antenna of albosetum . (After Bright 1968: 639). for the myelophagous C, qitadridens and three xylophagous species. Keys; Blackmail (1938c; 536), Wood (1982; 316). Tribe Hypoborini Hypoborinae NiissHn [1911: 376, Type-genus: Hy- poborus Erichson, 1836] Description. — Frons dimorphic or not, male impressed, female convex or less strongly impressed, except in some Li- parthrum species this feature is reversed; eye entire; funicle 3- to 6-segmented, club with up to three sutures, sometimes absent; prono- tum variously armed in restricted areas; pro- coxae contiguous; tarsal segment 3 narrow; scutellum not visible; crenulations on elytral bases not continued laterad from interstriae 5; postnotum fused to metatergum (remnants of suture sometimes visible). Biology. — All are monogamous and phloeophagous. In all except Chaetophloeus the parental gallery is a simple, oval cave. Eggs are packed in frass in niches on the mar- gins of the central cave (Liparthrum) or of the egg tunnels (Chaetophloeus). Larval mines radiate out from the parental chamber and rarely cross one another; they are visible on the inner surface of peeled bark. Taxonomy. — This tribe is sparsely and widely distributed in the warm climates around the world. Except for Liparthrum (Fig. 44), which is almost worldwide in the warm areas, the remaining genera are of lim- ited distribution. One is American, 1 Aus- tralian, 2 Madagascaran, and 4 African (1 of these reaches nearby areas of Europe and Asia in cultivated fig). It appears to be a relict group that once enjoyed much greater distri- bution and diversity than at present. The American genus, Chaetophloeus , is quite different structurely and biologically from the remaining closely related genera. Members of this tribe apparently prefer arid or semiarid areas or habitats and tend to be rare. Since they breed in shrubs or small trees of marginal economic importance, it is sus- pected that a majority of the species await discovery. Key to the Genera of Hypoborini 1. Funicle 6-segmented, club small, conical, with two straight sutures; Australia; Acacia, 1.3 mm Zygopliloeus — Funicle with 3- to 5-segments; club flattened. sutures present or absent 2 2(1). Protibia strongly flattened, rather broad, lateral apical half armed by a row of 7-10 closely set, socketed teeth; pronotal asperities confined to two or three paired clusters on lateral thirds, each cluster containing 1-5 denticles; funicle 5- segmented, club clearly marked by three su- tures; phloeophagous; North and South Amer- ica; 1. 1-2.. 5 mm Chaetophloeus 54 Great Basin Naturalist Memoirs No. 10 — Protibia slender, lateral margin armed by about ChaetopMoeus LeConte [ 1876; 382, Type- four rather widely spaeed, socketed teeth; species; Hijlesinus hystrix LeConte, monoba- pronotal asperities mostly on median third, . „ n ■ /-^ mn/- r,r-n more abundant; funicle 3- to 5-segmented 3 ^ic. Synonyms: Renocis Casey 1886; 257, „,„, , , 1 r • 1 c .. J I u .^u Tvpe-species; Renocis heterodoxus Casey, 3(2). Antennal lunicle 5-segmented, club rather i n j i? n ir.i-T broad, sutures ver>' broadly procurved; protibia monobaSlC; Pseudocrtjplmllis Swaine 1917; slender, with one large spine on outer apical 20, Type-specieS; Pseudocrijphalus brittaini angle and four small, socketed denticles on lat- Swaine =Renocis heterodoxus Casev, original eral margin; crenulations on basal margins of designation]. Distribution; 17 species in elytra feeble; body subglabrous; Madagascar; xt ii a ■ i i- , . i i n • 1.5 mm ■ Glochiphorus North America and adjacent islands, 2 m „ ., . , .111 South America. All are monogamous and — Protibia not armed on outer apical angle by a , , , ,^ mi /,r^<^ ^^r<\ major spine; antenna variable; elytra bearing phloeophagous. keys: Blackman (1940: 376), setae ' .4 Wood (1982: 349). 4(3). Funicle 4- or 5-seginented, club with sutures Crijphyophthorus Schedl [1953c: 294, obscure, club more broadly oyal .5 Type-species: Cryphyophthoriis eggersi — Funicle 3-segmented, club usually more slen- Schedl, original designation]. Distribution: 1 der 7 in Sumatra (Indonisia), 1 in Madagascar. .5(4). Antennal funicle 4-segmented (appearing 3-seg- Liparthriltn WoUaston [1854: 294, Type- mented, but 4- or possibly .5-segmented on slide species: Lipaithrum hifuberculatum VVollas- mount), club unmarked by sutures; basal mar- . ■ ■ i i ■ ■.- c r ■ f , ^ ^. - 1 ■ 1- 1 1 ton, original designation. Synonyms: Lei- gins ot elytra a continuous costa, individual i n i i crenulations feebly indicated; anterior half of part h rum Wollaston 1854: 294, invaHd error pronotum asperate; striae not impressed, punc- in spelling, amended by Wollaston (1864: tures coarse, deep, wider than interstriae; 265), International Commission On Zoological Madagascar; 1.6 mm Cryphyophthorus Nomenclature (1981: 64) ruled this to be an — Funicle 4- or 5-segmented; crenulations on invalid spelling; Erineosimis Blackman 1920: basal margins ofelytra well-formed, strial punc- 53 Type-species: Erineosinus squcimOSUS tures smaller; pronotal asperities olten present, mi 1 • n? 7 j .1 o 1 11 but less conspicuous 6 Blackman, monobasic; Fluoeoc lulus Schedl ct-v iri^„ tjiij viff 1953c; 292, Tvpe-species: Phloeochilus pala- 6(5). Funicle 4-seginented, club devoid ol sutures; 1 11 11 r ; meso- and metatibiae slender, about equal to f/""'* Schedl, original designation; Phloeo- protibia; phloeophagous; S USA to N South frypetus Wood 1960a: 16, Type-species: America, S Europe and N Africa to China and Phloeotrypetus palauensis Wood, Original Micronesia; 0.8-1.5 mm Liparthrum designation; Dacryophthorus Schedl 1971b; — Funicle 5-segmented, club with three obscure 281, Type-species; Dacn/ophthorus brincki sutures: meso- and metatibiae rather strongly ^ ^ ii • • 1 i • i.-" 1 r^- l -u 1.- r» n ,, J 1, 1 J 1 .' Scnecil, original designation . Distribution: 2 ilattened, much wider and more coarsely serrate ' o o j than protibia; phloeophagous; S Europe, N species in South America, 7 in North and Africa to Asia Minor; 1.1-1. 4mm Hypohorus Central America; 11 in the Canary and other 7(4). Antennal club with three distinct suture.s; meso- Atlantic islands, 6 in Europe, 1 in China, 4 in and metathoracic tibiae slender, about equal to the Indo-Malayan area, and 1 in Micronesia, protibia; phloeophagous; Africa; 1.2-1.8 mm . ^11 are monogamous and phloeophagous. StyaCOptmUS j.gy^. g^J^pJJ ^g^g^. 33^ f^^. jj^^ g^^^^^^ y^^^^ — Antennal club long and slender, sutures not in- ^^gg-j: 364) for North and Central America. dicated; scape ornamented by a tutt of long hair; t^ . i no-./^> /-n t^ discal interstriae 2 on basal fourth with one tu- Hypoborus Enchson [1836: 62, Type-spe- bercle greatly enlarged in male, 3 with four cies; Hypoborus fici Erichson, monobasic], tubercles on posterior half and upper half of Distribution: 1 species from S Europe, N declivity; phloeophagous; Africa; 1.6-2(1 mm ^^^^^..^ ^,^J gyy ^^j^ j,^ cultivated fig. MonOga- Dacniostactus i i i i mous and phloeophagous. Zygophloeus Schedl [1958b; 215, Type- Styracoptinus Wood [1962: 77, Type-spe- species; Zygophloeus australis Schedl, mono- cies: Styracopterus miirex Blandford, auto- basic]. Distribution: 1 species in Acacia in matic. Synonyms; Styracopterus Blandford Australia. 1896c; 323, Type-species: Styracopterus Glochiphorus Strohmeyer [1910c: 126, murex Blandford, monobasic, preoccupied; Type-species; Glochiphorus globosus Stroh- Afrotrypetus Bright 1981b: 113, Type-spe- meyer, monobasic]. Distribution: 1 species in cies; Afrotrypetus euphorbiac Bright, original Madagascar. designation]. Distribution; About 5 species in 1986 WOOD; Genera of Scolytidae 55 S Africa. All are apparently monogamous and phloeophagous. Dacryostactus Schaufuss [1905: 79 (reprint p. 3), Type-species; Dacryostactus kolbei Schaufuss, monobasic]. Distribution; 1 spe- cies in Africa. Monogamous and phloeo- phagous. Tribe Polygraphini Polygraphidae Chapuis [1869: 48, Type-genus: Polygra- phus Erichson, 18.36] Carphoboridae Niisslin [1911: 376, Type-genus: Car- phoborus EichhofF. 1864] Description. — Frons dimorphic, male variously impressed and sparsely pubescent, female concave to convex and usually orna- mented by conspicuous setae; eye emarginate to completely divided; antennal fimicle 5- or 6-segmented, club slightly to strongly flat- tened, symmetrical to strongly asymmetrical, with or without sutures; proco.xae contiguous; tarsal segment 3 slender; pronotum unarmed; scutellum not visible; crenulations on bases of elytra either individual (separate) or repre- sented by a continuous costa (Serrastiis), con- tinued to humeral angle; metanotum fused to its postnotum. Biology. — All are phloeophagous. Car- phobius and some Polygraphits are monoga- mous; apparently Chortastus and Serrastus share this habit; Carphoborus (Fig. 45) and most Polygraphus are polygynous. Parental tunnels are monoramous or biramous in monogamous forms and radiate in polygynous forms. The nuptial chamber is unusually large in most species. Eggs are packed in frass in niches. Larval mines wander considerably and have a greater tendency to cross one an- other than in most other tribes. Taxonomy. — In my initial study of the tribes of Scolytidae (Wood 1978), the Poly- graphini were reluctantly given tribal status only after much hesitation and the pondering of many questions. Since then, all reserva- tions have been dismissed. Although quite specialized in several respects, representa- tives of two genera (Serrastus, Polygraphus) have the outer apical angle of the protibia produced beyond the tarsal insertion and are armed as in primitive representatives of other unspecialized tribes. The tribe generally ap- pears to have been derived from the same ancestral stock as the Phloeosinini, but Fig. 45. Carphoborus pirucolens Wood, dorsal aspect of female. (After Bright and Stark 1973: 155). the one known species of Phloeographus bears a remarkable superficial resemblance to certain Tomicus species (Hylesinini). The genera form a rather compact unit without conspicuous divisions. Four small genera are exclusively African (Serrastus, Chortastus, Phloeographus, Cardroctonus), one is Malayan (Bothinodroctonus), and one is American (Carphohius). The larger genus Carphoborus is primarily North American and probably originated there from stock derived from Asia. European and Asian spe- cies of Carphoborus appear to have been derived from the more primitive and diverse American fauna. The largest genus, Polygra- phus , is largely African, with strong represen- tation in Europe and Asia, and with three species in northern North America. The ab- sence of this group in South America suggests either an origin since early Tertiary, if Africa were the site, or else an origin in tropical Eurasia. Polygraphus reached North America in comparatively recent time, Carphoborus and Carphobius much earlier. Eurasian Car- 56 Great Basin Naturalist Memoirs No. 10 phohorus were probably derived from a sec- ondary radiation that moved from North America to Asia. Carphohius apparently rep- resents a relict from a very early ancestral stock not now represented elsewhere. Key to the Genera of Polygraphini 1, Basal margins of elytra armed by a continuous costa; protibia very slender, its median apical mucro bent laterad, about one lateral denticle present, posterior face asperate; funicle 5-seg- mented; club strongly flattened, moderately asymmetrical, sutures not indicated; phloeo- phagous; Africa; 1.8-2.0 mm Serrastus — Basal margins of elytra serrate, armed by a row of crenulations; protibia more broadly flattened, lateral margin armed by several socketed denti- cles 2 2(1). Funicle 5-segmented (.5- or 6-segmented in Polygraphus , if 6-segmented then eye com- pletely divided); male frons armed by a median pair of tubercles near upper level of eyes; vesti- ture of abundant scales (except glabrous in some Bothinodroctonus) 3 — Funicle 6-segmented; eye never divided into two parts; male frons unarmed by median tuber- cles; vestiture hairlike or sparse if scalelike .... 7 3(2). Eye completely divided by a deep emargination (some Asian and Alrican species only emar- ginate); antennal club rather strongly flattened, asymmetrical, devoid of sutures; phloeo- phagous; North America, Asia, Europe, Africa; 1..5-.5.5 mm Polygraphus ' — Eye less than half divided by an emargination; antennal club symmetrical (or nearly so), clearly marked by sutures 4 4(3). Sutures of antennal club procurved; vestiture sparse, in uniseriate rows on interstriae; tropical Africa 5 — Sutures of antennal club straight or nearly so; vestiture much more abundant; northern' hemi- sphere 6 5(4). Antennal club with three weakly procurved su- tures indicated, ape.\ strongly acuminate; vesti- ture sparse, of uniseriate rows on declivity; de- clivital interstriae 2 .slightly impressed, unarmed, 1 and 3 armed by small tubercles; SW Africa; 2.2 mm Phloeographus — Antennal club with suture 1 partly septate, 1 weakly, 2 moderately procurved; strial punc- tures very fine; Africa; 1.3-2.0 mm . Cardroctonus 6(4). Male frons shallowly impressed below upper level of eyes, female usually convex; vestiture of abundant scales; antennal club moderately large, flattened; phloeophagous; North Amer- ica, N Asia, Europe, N Africa; 1.4-2.6 mm .... Carphoborus — Male frons profoundly excavated from eye to eye from epistoma to well above eyes, female simi- larly but less strongly excavated; antennal club small, more nearly conical; vestiture confined to a few interstrial setae on and near declivity; phloeophagous; India to Malaya; 1.8-2. .5 mm . . Bothinodroctonus 7(3). Antennal club symmetrical, with three trans- verse sutures indicated; vestiture abundant, hairlike; phloeophagous; North America; 1.6- 2.1mm Carphohius — Antennal club rather strongly asymmetrical, part of suture 1 indicated, strongly oblique; vestiture usually confined to declivity, scalelike; phloeophagous; Africa; 2.. 5-.5. 3 mm . . . Chortastus Serrastus Nunberg [1969: 392, Type-spe- cies: Serrastus ivoriensis Nunberg =Chor- tastiis similis Eggers, monobasic]. Distribu- tion: 2 species in Africa (Ghana to Zaire). They are monogamous and phloeophagous. Phloeographus Wood [1984: 229, Type- species: Pliloeographus mamibiae Wood, original designation]. Distribution: 1 species in Africa (apparently Namib Desert, SW Africa). Cardroctonus Schedl [1966b: 361, Type- species: Cardroctonus orientalis Schedl, orig- inal designation]. Distribution: 2 species in Africa. Polygraphus Erichson [1836: 57, Type-spe- cies: Hylcsiuus pubescens Fabricius =Der- mestes poligraphus Linnaeus, monobasic. Synonyms: Lepisomus Kirby 1837: 193, Type- species: Apate (Lepisomus) rufipennis Kirby, subsequent designation by Hopkins 1914: 124; Spongotarsus Hagedorn 1908: 372, Type-species: Spongotarsus quadrioculatus Hagedorn, monobasic; Pseudopolygraphus Seitner 1911: 105, Type-species: Polygraphus grandiclava Thomson, subsequent designa- tion by Hopkins 1914: 128; Ozophagus Eggers 1919: 234, Type-species: Ozophagus canier- unus Eggers, subsequent designation by Wood 1982: 386; Urdugraphus Beeson 1941: 301, nomen nudinn]. Distribution: 3 species in North America, about 45 in Asia and adja- cent islands, 6 in Europe, about 45 in Africa and Madagascar. Most are polygynous, al- though monogamy occms in some Asian and African species. All are phloeophagous. Keys: Schedl (1955a: 5) for Europe and part of Asia, Murayama (1956: 278) for Japan, Wood (1982: 387) for North America. Carphoborus Eichhoff [1864: 27, Type- species: Hylesinus minimus Fabricius, mono- basic. Synonym: Estenoborus Reitter 1913: 58, Type-species: Hylesinus perrisi Chapuis, 1986 WOOD: Genera of Scolytidae 57 monobasic]. Distribution: 21 species in North America, 14 in Europe and Asia, 2 in N Africa. All are polygvnous and phloeophagou.s. Keys; PfefFer (1914: 169) for Europe, Balachowskv (1949: 143) for France, Wood (1954a: 507, 1982: 372) for North America. Bothinodroctonus Schedl [1969: 208, Type- species: Bothinodroctoniis bicinctus Schedl, monobasic]. Distribution: 4 species from In- dia to Borneo and China. One species is phloeophagous in Odina . Carphobius Blackman [1943c: .398, Type- species: Carphohius arizonictis Blackman, original designation]. Distribution: 3 species in North America (Arizona to Guatemala). All are monogamous and phloeophagous in conifers. Key: Wood (1982: 369). Chortastus Schaufuss [1905: 15 (reprint p. 8), Type-species: Choi-tasttis cameninus Schaufuss, monobasic. Synonym: Afrochram- esus Schedl 1971a: 197, Type-species: Afrochramcsus bafi,nenai Schedl, original designation]. Distribution: 7 species in Africa. Apparently all are monogamous and phloeo- phagous. Subfamily Scolytinae Scolytarii Latreille [1807: 27.3, Type-genu.s: Scohjttis Ge- offroy, 1762] In previous classifications, this subfamily has been divided into two or more major divi- sions equivalent to subfamilies. However, it appears that this unit contains both the most primitive elements of the family that are most closely related to primitive Platypodidae and also the main evolutionary thrust of the platy- podid-scolytid group. This diversity makes characterization of the subfamily ratlier diffi- cult. Members of this subfamily have the bases of the elytra simple, forming a straight, trans- verse line across the body. A large, flat scutel- lum is usually visible. The elytral bases are weakly subcostate in some Scolytini, Ctenophorini, and Cryphalini. The body tends to be more elongate, with specialization directed toward cylindrical compaction and the xylomycetophagous habit. A broadly oval body outline is usually confined to primitive, phloeophagous genera. Armed elytral bases that suggest a relationship to the Hylesininae occur in Ctenophorini (Cnemonyx, allies of galeritus). Five clusters of relationship are recognized within the subfamily. Progressing from primi- tive to the more specialized, these include: (1) Ctenophorini, Scolytini, and Scolytoplatypo- dini: (2) Micracini and Cactopinini; (3) Carphodicticini, Ipini, Dryocoetini, Cryp- turgini, Xyloterini, and Xyleborini; (4) Xyloc- tonini and Cryphalini, and (5) Corthylini. Of these, the Corthylini are by far the most dis- tinctive from an anatomical point of view. Tribe Scolytini Scolvtarii Latreille [1807: 27.3, Tvpe-^enus: Scohittis Ge- oftVoy, 1762] Camptoceridae Chapuis [1869: 49, Type-genus: Campto- cents Latreille, 1829] Description. — Frons usually strongly di- morphic, male variously impressed, female flat to convex, one or both often ornamented by hair; posterior face of head truncate; eye oval, entire; scape short to elongate, fuiiicle 7-segmented, club rather large, flattened, su- tures present or not; pronotum unarmed, its lateral margins costate; protibia (usually all three tibiae) unarmed on lateral margin, outer apical angle extended into a spine curving toward and exceeding inner apical angle, socketed denticles never present; meta- pleural suture descending subvertically to groove receiving groove on costal margin of elytra then turning abruptly and continuing parallel to groove to near metacoxal process; in two genera venter of abdomen ascends con- spicuously to meet apex of weakly declivous elytra. Biology. — All are monogamous except for a few neotropical and one European bigynous Scolytiis . Camptocerus species are xylomyce- tophagous; those in the other three genera are phloeophagous. Parental galleries are bi- ramous, except for a few that are mono- ramous. In Camptocerus , a biramous, trans- verse tunnel without niches is usually made in the cambium region, then a radial egg tunnel (sometimes branched) is extended from it into the xylem. Eggs are deposited in niches in this radial tunnel. The larval mines of phloeo- phagous species follow a definite course and rarely cross one another. Camptocerus larvae enlarge the egg niche into a cradle just large enough to accommodate the mature beetle, somewhat similar to some other ambrosia beetles. 58 Great Basin Naturalist Memoirs No. 10 Taxonomy. — Except for the Eurasian ele- ment of Scolytiis , members of this tribe are exclusively American. Their obvious origin is neotropical. Some members of this group re- semble the primitive ancestral line that prob- ably gave rise to the Hylesininae; in fact, some Cnemonyx have crenulations on the basal margins of the elytra. They also appear closely allied to the stock from which Platypodidae diverged. The four genera assigned here form a compact group, although the Cnemonyx ap- pear more primitive and, in some respects, overlap the other three. Scolytits appears to have reached North America from South America by the begin- ning of the Tertiary, when a secondary radia- tion occurred. Elements of this radiation then reached Eurasia, where another radiation oc- curred that was based on a progressively re- duced gene pool. The number of Eurasian species in this genus now may equal or exceed the American component, although the anatomical and biological diversity there is greatly reduced. Key to the Genera of Scolytini 1. Scutellar area of interstriae 1 not depressed, scutellum flush (even) with elytral bases; basal margins of elytra with a fine raised line {some Cnemontjx with crenulations instead), outline of anterior margins form a continuous, straight, transverse line with scutellum; ventral profile of abdomen ascending gradually 2 — Scutellum depressed, subtriangular, apically (posteriorly) pointed; elytral bases depressed in scutellar area, appearing emarginate in median area; ventral profile of abdomen usually ascend- ing abruptly at segment 2 3 2(1). Antennal club usualK" with two or three sutures clearly marked by setae; scutellum small, longer than wide, often convex; apical liiargin of meso- and metathoracic tibiae commonly bearing tu- bercles on anterior edge in addition to inner and outer apical spines; usually inore coarsely sculp- tured; phloeophagous; USA and Mexico to Ar- gentina; 1.0-3.9 mm Cnemonyx — Antennal club with suture 1 (only) marked inter- nally by a partial septum; scutellum flat, 1.5 or more times as wide as long; meso- and metatho- racic tibiae acutely margined on apical anterior edge, without supplemental denticles; usually very finely sculptured; xylomycetophagous; Central and South America; 2.6-8.5 mm Camptocerus 3(1). Basal portion of costal margin of elytra deeply, broadly excised, metepisternum conspicuously expanded into this notch (Fig. 46); abdomen abruptly flexed upward at posterior margin of Fig. 46. Scolytopsis inincticoUis . female. Note emar- ginate costal margin of elytron. segment 2; phloeophagous, tropical; Mexico (Oaxaca) and Cuba to Argentina; 1.6-3.5 mm . . Scolytopsis — Costal margin of elytra normal (straight) and overlapping metepisternum; abdomen flexed upward from anterior margin of segment 2; phloeophagous; North and South America, Eu- rope, N Asia, N Africa; 1.3-5.5 mm Scolytus Cnemonyx Eichhoff [1868a: 150, Type- species: Cnemonyx galeritus Eichhoff, monobasic. Synonyms: Ceratolepis Chapuis 1869: 52, Type-species: Ceratolepis jiicundus Chapuis, monobasic; Loganius Chapuis 1869: 52, Type-species: Loganius flavicornis Cha- puis, monobasic; Minuhis Eggers 1912b: 206, Type-species: Minulus harhatus Eggers, monobasic; Coptodryas Schedl 1948b: 262, Type-species: Coptodryas hylurgoides Schedl. monobasic; Coptosomus Schedl 1952a: 363, Type-species: Coptodryas hylur- goides Schedl, automatic]. Distribution: 19 species in USA (Florida), Mexico, and Central America, about 27 in South America and adja- cent islands. All are monogamous and phloeophagous. Keys: Blandford (1896d: 128) for Central America, Wood (1982: 394) for North and Central America. Camptocerus Latreille [1829: 91, Type- species: Hylcsiints aeneipennis Fabricius, monobasic]. Distribution: 19 species from Central and South Ainerica. All are monoga- mous and xylomvcetophagous. Key: Wood 1982: 412 for Central America. Scolytopsis Blandford [1896d: 120, 123, Type-species: Scolytopsis puncticollis Bland- ford, monobasic]. Distribution: 7 species from Cuba and Mexico (Oaxaca) to Argentina. All are phloeophagous and monogamous. Key: Wood (1982: 417) for Central America. Scolytus Geoffrey [1762: 309, Type-spe- cies: Bostrichus scolytus Fabricius, desig- 1986 WOOD: Genera of Scolytidae 59 nated by the International Commission on Zoological Nomenclature (China 1962: 3). Synonyms: Ekkoptogaster Herbst 1793: 124, Types-species: Bostrichus scoh/ttis Fabricius, designated by Hopkins 1914: 121; Copto- gaster Illiger 1807: 321, Type-species: Bostrichus scolytus Fabricius, designated by Hopkins 1914: 118; Eccoptogaster Gyllenhal 1813: 346, an isotypical emendation oi Ekkop- togaster Herbst; Scolytochchis Reitter 1913: 23, Type-species: Ips multistriatus Marsham, designated by Wood 1982: 419; Riigitloscohj- tus Butovitsch 1929: 20, Type-species: Bostrichus rugulosus Miiller, designated by Wood 1982: 419; Archaeoscohjtus Butovitsch 1929: 21, 23, Type-species: Scohjtus claviger Blandford, monobasic, not a genus-group name, no status; Spinuloscolytus Butovitsch 1929: 21, 24, Type-species; Ips multistriatus Marsham, present designation, not a genus- group name, no status; Tubuloscohjtus Bu- tovitsch 1929: 21, 33, Type-species: Eccopto- gaster intricatus Ratzeburg, present desig- nation, not a genus-group name, no status; Pygrnaeoscolytus Butovitsch 1929: 21, 28, Type-species: Bostrichus pygmaeus Fabri- cius, present designation, not a genus-group name, no status; Pinetoscohjtus Butovitsch 1929: 22, 48, Type-species: Scolytus marawitzi Semenov, monobasic, not a genus- group name, no status; Confusoscolytus Tsai, Yin, & Huang 1962: 4, 14, Type-species: Ec- coptogaster confusus Eggers, monobasic]. Distribution 28 species in North and Central America, about 29 in South America, 47 in Asia, Europe, and N Africa. Monogamous, except for 1 European and about 20 Central and South American bigynous species, and all are phloeophagous. Keys: Blackman (1934: 6) for North America, Schedl (1937a: 156) for South America and (1948a: 4) for Europe, Michalski (1973; 137) for Europe and Asia, Wood (1982: 420) for North and Central America. Tribe Ctenophorini Ctenophoridae Chapuis [1869: 49, Type-genii.s: Ctenophorus Chapuis, 1869 =Scohjtodes Fer- rari, 1867] Probleehilidae Eichhotf [1878a: .34, 46, 167, 298, Type-genus: Prohhchilus Eichhoff, 1878 = GymnochiluH Eichhoff, 1867] Hexacohdae Eichhoff [1878a: .3.5, 57, .306, Type- genus: Hexacolufi Eichhoff, 1868 =Scolytodes Ferrari, 1867] Description. — Frons usually dimorphic, male impressed and female flat to convex in Pycnarthrum and Gymtiochilus , sexual differ- ences obscure in Microborus , male convex and female variously sculptured and orna- mented in Scolytodes ; posterior face of head truncate; eye usually elongate, entire to sinu- ate; scape elongate, funicle 6- or 7-seg- mented, club with or without sutures; prono- tum armed or not, its lateral margins costate; procoxae widely separated; protibia with one or more socketed denticles on lateral margin, spine on lateral apical angle usually extending beyond level of tarsal insertion; pleural suture about as in Scolytini. Biology. — All are monogamous, except for a few polygynous Scolytodes . All are phloeo- phagous, except for the xylophagous Scoly- todes multistriatus Wood and species that in- fest Cecropia leaf petioles. Parental galleries vary from a simple to an elongate cave to stellate in Scolytodes ; they are biramous in Pycnai-thrum and Gymtiochilus and indefi- nite, nondirectional, and without definite pat- tern in Microborus. The eggs may be scat- tered loosely in the parental chamber or placed in crude niches in Scolytodes ; definite niches are formed in Pycnarthrum and Gym- nochilus; they were not observed in Mi- croborus. The larvae usually feed commu- nally in Scolytodes; they form individual mines that follow a somewhat definite direc- tion in Pycnarthrum and Gymnochihis ; they are individual and without a definite direction in Microborus . Symbiotic relationships with fungi were not observed. Taxonomy. — The tribe is restricted to the American tropics, except that Microborus boops Blandford was introduced into tropical Africa. Scolytodes and Microborus are closely related to one another. Pycnarthrum and Gymnochilus are remotely related to those genera and to one another. Pycnarthrum could easily be placed in Hylesinini. This tribe occupies a position intermediate be- tween the Scolytini and the more highly evolved tribes in this subfamily. In all mem- bers the outer apical angle of the protibia projects beyond the tarsal insertion, a primi- tive feature shared by primitive members of several other tribes. It is the Ctenophorini protibia, not the type found in Scolytini, that appears to resemble the ancestral type of all 60 Great Basin Naturalist Memoirs No. 10 scolytids. The eye shape, usually unarmed pronotum, presence of interstriae 10, simple sculpture of the elytra, the elytral locking mechanism, and diversity of habits all suggest that, when considered as a whole unit, this is probably the most primitive of all of the tribes of Scolytidae. It is clearly of neotropical origin and has spread into southern North America only recently. This phyletic line is repre- sented in the Old World by Scolytoplatypo- dini, a group that has diverged significantly in both structure and habits. Key to the Genera of Ctenophorini 1. Eyes elongate, approximate above and below, coarsely faceted, shallovvly emarginate; entire surface of pronotum smooth and punctured, not armed 2 — Eye oval, enitre, finely faceted; pronotum as- perate anteriorly or, if smooth, then anterior margin of elytra bearing a fine, raised line .3 2(1). Antennal club subglobular, about as long as wide, sutures not clearly indicated; pronotum longer than wide, its lateral margins straight or feebly constricted; vestiture hairlike, usually sparse; small, slender species; phloeophagous; Central and N South America, Jamaica, Africa; 1.1-1.5 mm Microbonis — Antennal club asymmetrically flattened, pointed, at least 1.5 times as long as wide, su- tures 1 and 2 clearly marked by setae; pronotum wider than long, its lateral margins arcuate; vestiture of abundant, short, bristlelike scales; larger, stouter species; phloeophagous; Florida and Texas to Brazil; 1.3-2. 1 mm .. , . Pycnarthrtim 3(1). Antennal funicle 7-segmented, club large, broad, usually with procurved sutures or su- tures obsolete; elytral vestiture consisting of abundant, minute hair and sparse interstrial rows of long, erect, scalelike bristles; summit of pronotum on basal third, asperities on anterior area coarse; elytral base without a fine, raised line; phloeophagous; Mexico (Puebla) to Brazil and Bohvia; 1.5-2.3 mm Gymnochilua — Antennal funicle 6-s6gmented, club small, su- tures present or not; elytral vestiture sparse (usually), hairlike; pronotal asperities fine, if present; summit at middle or indefinite; basal margins of elytra marked by a fine, raised line; mostly phloeophagous; Florida and Mexico to Argentina; 0.9-3.5 mm Scolytodes Microborus Blandford [1897: 175, Type- species: Microborus boops Blandford, mono- basic. Synonym; Pseitdocryptiiifius Eggers 1919: 236, Type-species: Pseudocrijptur'nous; most are phloeo-phagous, a few (about 6) are .xylophagous, a few bore in leaves, etc. Key: Wood (1982: 511) for North and Central America. Stenoclyptus Blackman [194.3b: 3.56. Type- species: Steiioclyptus rhois Blackman = Tseu- dothysanoes sulcatus Bruck, original designa- tion]. Distribution: 2 species in USA (Cahfomia) to Me.\ico (Puebla). Both are big\- nous and phloeophagous. Kev. Wood (1982: 556). Saurotocis Wood [1984: 229, Type-species: Micracidendron tomicoides Schedl, original designation]. Distribution: 2 species in Mada- gascar. Thysanoes LeConte [1876: 369, Type-spe- cies: Thysatwes firnbricornis heConte, mono- basic]. Distribution: 13 species in USA to Costa Rica. Key: Wood (1982: .557). Phloeocleptus Wood [1956a: 147, Type- species: Phloeocleptiis caudatus Wood, origi- nal designation]. Distribution: 11 species from North America (Mexico) to Central America (Costa Rica). All are big\'nous and phloeophagous. Key: Wood 1982: 570). Hylocurus Eich'hoff [1872: 133, T>pe- species: Hylocurus elegans EichhofP, mono- basic. Synonym: Micracisoides Blackman 1920: 19, T}"pe-species: Micracis rudis LeConte, subsequent designation by Wood 1982: 608]. Distribution: 34 species in North and Central America, about 20 in South America and adjacent islands. All are bigy- nousandxvlophagous. Kevs: Blandford (1898: 220), Wood (1982: 609). Phloeocurus Wood [1984: 230, Type-spe- cies: Hylocunts africanus Schedl, original designation]. Distribution: 1 species in Africa. Pseudomicracis Eggers [1920a: 36, Type- species: Faeudomicracis elsae Eggers, mono- basic]. Although the unique holotype of the type-species is lost, the genus is identifiable from the description. Distribution: 7 species in Africa and Madagascar. Micracisella Blackman [1928b: 192, Type- species: Micracis opacicollis LeConte, auto- matic. Synonym: Pseudomicracis Blackman 1920 (December): 20, Type-species: Micracis opacicollis LeConte, original designation. Fig. 48. Cactopimis desertus Bright, male. (.After Bright 1973; 155). preoccupied]. Distribution: 20 species in North and Central America 1 of which also occurs in South America (Colombia). All are monogamous and mvelophagous. Kev: Wood (19S2:^594). Micracis LeConte [1868: 164, Type-spe- cies: Micracis siituralis LeConte, subsequent designation by Hopkins 1914: 125]. Distribu- tion: 18 species in North and Central Amer- ica, 1 in Cuba, 2 in South America (Venezuela); several others have been named from South America but most, if not all of these, have been or should be transferred to other genera. All are bigynous and xy- lophagous. Key: Wood (1982; 579). Tribe Cactopinini Cactopinae Chamberlin [19.39: 243, T\'pe-genus: Cac- topinus Schwarz, 1899] Description. — Frons dimorphic, male strongly impressed or excavated, with the epistomal margin armed by a pair of (usually 66 Great Basin Naturalist Memoirs No. 10 confluent) hornlike spines of large to enor- mous size (Fig. 48), female convex or mod- estly impressed, epistoma unarmed by spines; posterodorsal area of head modestly produced caudad; eye small, oval, entire; an- tennal scape rather short, funicle 5-seg- mented, club almost conical to strongly flat- tened, sutures straight to procurved, marked by rows of setae; pronotum asperate on ante- rior slope, summit at or near posterior mar- gin, sometimes projecting caudad beyond basal margin; procoxae contiguous; elytral sculpture unique, rather conservative, usu- ally sulcate on declivity, almost always coated by resinous film of host origin. Biology. — All are monogamous and either phloeophagous or in cactus (Cereits and allied genera). Those in cactus breed in dry, dead (yellowing) tissue immediately below the epi- dermis, or in scar tissue in deep wounds (hiib- bardi). One species breeds in Yucca leaves (depressus). The parental galleries form an irregular cave, with egg niches. Eggs are de- posited individually in niches in two species; some species in cactus deposit them in clus- ters. Larval mines may be individual or lost in a criss-crossing maze. Successive generations have been bred in the same piece of dry cactus for four years. Symbiotic relationships with fungi have not been reported. Taxonomy. — This unique tribe is restricted to the Mexican plateau region. Its nearest affinity to other groups appears to be with the Micracini, although the relationship is re- mote. They are exceedingly rare. Cactopinus Schwarz [1899; 11, Type-spe- cies; Cactopinus hubbardi Schwarz, monoba- sic. Synonym: Cactopinonis Bright 1967; 918, Type-species: Cactopinus cactoplithorus Wood, original designation]. Distribution: 19 species in W North America (California and W Utah to Me.xico). All are monogamous and phloeophagous when in woody plants or subepidermal when in cactus {Cereus and al- lied genera only). Keys; Blackman (1938a: 151), Bright (1967: 919), Wood (1969; 43, 1982; 638). Tribe Carphodicticini Carphodicticini Wood [197L 19, Type-genus: Carphod- icficHS Wood, 1971] Description. — Frons weakly to moder- ately dimorphic, male strongly convex, fe- male slightly to moderately flattened and sometimes abundantly pubescent; eye short, broadly oval and entire to very elongate and sinuate to shallowly emarginate; scape short and rather stout to elongate and rather slen- der, funicle 5-segmented, club flattened, small, and symmetrical, with transverse su- tures, to moderately large and slightly asym- metrical, with slightly oblique sutures, su- tures marked by grooves and rows of setae, apparently not septate; pronotum elongate, sides conspicuously constricted on middle half, pronotum unarmed; posterior face of head truncate, dorsomedian area not ex- tended caudad; proco.xae narrowly to rather widely separated; protibia slender to very stout and short, armed by socketed denticles on lateral margin; scutellum visible; basal margins of elytra rounded in two genera, ele- vated and carinate in one genus. Biology. — Carphodicticus is monogamous and phloeophagous. Numerous pairs of par- ent adults appeared to use the same entrance tunnel. Each pair followed a previously made tunnel for a short distance, then formed their own branch gallery for oviposition, such that the entire system consisted of branching and rebranching galleries. Eggs were deposited individually in niches at the cambium. Larval mines were exposed on peeled bark and were rather short. The host had been felled for several months before this species attacked; a Phloeotribus had largely abandoned the un- usually hot, dry tissues, but this species was thriving. Taxonomy. — The head and pronotal struc- ture suggest that this group is very primitive. It was probably derived from the same ances- tral stock as the Dryocoetini, Ipini, Cryp- turgini, Xyloterini and Xyleborini, but at a much earlier date. The disjunct distribution and rarity suggest that it is a relict group that had reached its maximum potential prior to the Tertiary. This tribe represents a first step toward one of the three most hightly evolved tribes (Xyleborini) of Scolytidae. Key to the Genera of Carphodicticini 1. Eye entire, broadly oval, short, about 1.3 times as long as wide; scape rather short, about three times as long as pedicel; antennal club rather small, sutures transverse; elytral apex strongly mucronate; ventrolateral margin of declivity armed by a row of rather strongly elevated serra- 1986 WOOD: Genera of Scolytidae 67 armed by a row of rather strongly elevated serra- tions; apparently xylophagous; India and Sri Lanka (Ceylon); 1.3-2.3 mm Craniodicticus — Eye sinuate to emarginate, elongate, more than twice as long as wide (not visible on type of Dendrodicticus), elytral apex rounded, not mu- cronate; lateral margin of declivity uniformly elevated, not armed by serrations 2 2(1). Basal margins of elytra rounded, not elevated; antennal club rather large, distinctly asymmet- rical, sutures slightly oblique; procoxae more narrowly separated; protibia short (only slightly longer than antennal club), stout (almost half as wide as long); phloeophagous; South America (Venezuela); 2.0-2,4 mm Carphodicticus — Basal margins of elytra rather strongly, acutely elevated along a continuous costa; antennal club rather small, symmetrical, sutures transverse; procoxae more widely separated; protibia longer, much more slender; habits unknown; South America (Argentina); 2.0 mm Dendrodicticus Craniodicticus Blandford [1895: 317, Type-species: Craniodicticus miicronatus Blandford, monobasic]. Distribution: 1 spe- cies in S India, 1 in Sri Lanka (Ceylon). They apparently are xylophagous, since one sample was "removed from wood" and the other from a creeper called "jungle rope. ' Carphodicticus Wood [1971: 19, Type-spe- cies: Carphodicticus cristatus Wood, original designation]. Distribution: 1 species in South America (Venezuela). It is monogamous and phloeophagous; it utilized the entrance tun- nels of a Phloeotribus species in order to gain access to the phloem. Dendrodicticus Schedl [1958a: 37, Type- species: Dendrodicticus argentiniae Schedl, monobasic]. Distribution: 1 species in South America (Argentina). Tribe Ipini Ipini Bede! [1888: 386, Type-genus: Ips DeGeer, 177.5] Description. — Frons usually dimorphic, male convex, female variously excavated, pro- tuberant, or ornamented by setae; eye sinu- ate, lower half usually much narrower than above; antennal scape slender, elongate, funi- cle 5-segmented, club either obliquely trun- cate or sutures on posterior face strongly dis- placed toward apex; pronotum rather strongly declivous on anterior half, rather closely, coarsely asperate; procoxae contiguous, inter- coxal piece deeply notched or absent; protibia armed by three or four socketed teeth; scutel- lum visible; elytral declivity moderately sul- cate to strongly excavated, lateral margins usually armed by tubercles or spines; vesti- ture hairlike. Biology. — All are phloeophagous. Most are heterosanguineously polygynous, al- though some Acanthotoniicus are monoga- mous. Hosts include members of the Pinaceae, except Acanthotoniicus breeds pri- marily in various angiosperm trees. Eggs are deposited in niches. Larval mines are individ- ual and rarely cross one another; they are exposed on the surface of peeled bark. The life cycle is comparatively short, with two or more generations per year apparently the normal habit. Ta.\ONOMY. — In most tribes of Scolytidae the sexes are easily distinguished by a differ- ence in the number of abdominal terga. The female tergum 8 is reduced in size and is telescoped beneath tergum 7 such that it is ordinarily hidden from view. A visible tergum 8 is present in the male. However, in Carpho- dicticini, Ipini, Dryocoetini, Crypturgini, Xy- loterini, and Xyleborini a small tergum 8 is visible in both sexes. The general body habitus, shape of the eye, basic type of anten- nal club (trend toward being obliquely trun- cate), and tibiae are also shared by these tribes. The Ipini occupy a position between Carphodicticini and Dryocoetini in which these characters were being formed. In Den- drochihis they are so poorly expressed that they are detected only with difficulty. How- ever, that genus does exhibit a stage in transi- tion to the circumtropical Acanthotomicus . Acanthotoniicus appears to have an ancient origin (prior to the Tertiary) and to have given rise, directly or indirectly, to Pitijogenes, Pityokteines, Orthotomicus , and Ips. Pitijo- genes appears to have arisen in Europe and northern Asia from African Acanthotoniicus - like stock and to have reached North America rather recently. Something resembling the neotropical Acanthotomicus miniicus species group probably gave rise in North America in early Tertiary to a stock that then evolved into Pityokteines and Ips in North America and to Orthotomicus in Europe and Asia. At a later date, a few representatives oi Pityokteines and Ips then migrated into Europe and Asia and one species of Orthotomicus reached North America. The reasoning on which the above is 68 Great Basin Naturalist Memoirs No. 10 based is (1) on the continent of origin of sev- eral species groups found today, only a frac- tional number of those species groups now occur in the invaded territory and (2) no spe- cies group is found in the invaded territory that is not presently also found in the area of origin. Fossils that might document this sup- position are unknown. Dendrochihis and Acanthotoinicus breed in a variety of non- coniferous hosts; Pityogenes, Pitijokteines, Orthotomiciis , and Ips occur exclusively in Pinaceae. Key to the Genera of Ipini 1. Eye short, oval, entire, not sinuate on anterior margin; antennal club rather small, flattened, unmarked by sutures; declivity convex to slightly, broadly flattened, without granules or tubercles; elytral vestiture not abundant, in rows, interstrial setae hairlike to scalelike; pro- tibia slender, armed by 3 socketed denticles on apical half of lateral marin; Africa; 1.0-2,0 mm Dendrochilus — Eye more elongate, sinuate on anterior margin; antennal club usually larger, marked by sutures (except absent in many Acanthotomicus), de- clivity usually excavated and variously armed by tubercles or spines 2 2(1). Elytral declivity rather narrowly bisulcate, lat- eral margins rather broadly elevated, rounded, and armed by not more than .3 pairs of denticles; lower margin of declivity rounded; usually smaller than 3 mm 3 — Elytral declivity broadly, rather deeply exca- vated, margins acutely elevated and armed by 3 or more pairs of denticles (1 to 6 pairs in tropical Acanthotomicus), lower margin of declivity with an acutely elevated, transverse ridge separating declivital excavation from apical margin (Fig. 49); usually larger than 3 mm 4 3(2). Prosternal intercoxal piece short, obtuse; female frons sometimes deeply, rather narrowly exca- vated; male declivity with 2 or 3 pairs of en- larged denticles; antennal club compressed, 2 sutures visible on apical third of posterior face; North America, Europe, Asia, N Africa; Pina- ceae; 1.8-3.7 mm Pityogenes — Prosternal intercoxal piece long and acutely ta- pered; female frons convex, never excavated; male declivity more narrowly impressed; female frons and anterior pronotum with dense, long vestiture (2 American exceptions); North Amer- ica, Asia, Europe; Abies; 1.6-3.0 mm Pityokteines 4(2). Antennal club obliquely truncate, sutures re- curved; third (lowest) major denticle not on lat- eral margin of elytral declivity, displaced mesad from margin, eye of normal size; North America, Asia, Europe; Pinaceae; 2.2-4.3 mm Orthotomicus — Antennal club flattened, sutures either pro- curved or moderately to strongly bisinuate; lat- eral margins ol elytral declivity armed by 1 to 6 pairs of major denticles, third pair (if present) on or incorporated into crest of lateral margin; eye usually either abnormally large or else very small 5 5(4). Sutures of antennal club (when present) moder- ately to very strongly procurved; eye large, very coarsely faceted, its width about equal to length of scape, its length more than twice length of scape; Mexico to South America, S Asia to Aus- tralia, Africa; angiosperm hosts (some Asian ex- ceptions); 1.4-2.7 mm Acanthotomicus — Sutures of antennal club moderately to strongly bisinuate (procurved in concinmts, mexicanus, orientatis): eye small, finely faceted, its width equal to much less than length of scape, its length equal to much less than twice length of scape; North America to N Nicaragua, Asia, Eu- rope, N Africa; Pinaceae; 2. 1-6.9 mm Ips Dendrochilus Schedl [1957a; 70, Type-spe- cies: Dendrochilus strombosiopsis Schedl, subsequent designation by Schedl 1962a: 55]. Distribution: About 10 species from Africa. Pityogenes Bedel [1888: 397, Type-species; Dermestes chalcographus Linnaeus, original designation. Synonyms: Eggersia Lebedev 1926: 121, Type-species: Bostrichus biden- tatiis Herbst, present designation; Pityocera- genes Balachowsky 1947: 44, Type-species: Bostrichus quadridens Hartig, original desig- nation]. Distribution: 7 species in North America, 18 in Europe, Asia, and N Africa. All are heterosanguineously polygynous and phloeophagous. Keys: Swaine (1918; 104), Bright (1976: 150), and Wood (1982: 650) for North America, Reitter (1913: 97) and Pfeffer 1946: 112) for Europe, Stark (1952: 272) for USSR, Schedl (1962c: 134) for Europe and Asia. Pityokteines Fuchs [1911: 33, Type-spe- cies: Ips curvidens Germar, subsequent des- ignation by Hopkins 1914: 127. Synonymy: Ortliotomides Wood 1951: 32, Type-species: Orthotomicus lasiocarpi Swaine, original des- ignation]. Distribution: 6 species in North America, 3 in Asia and Europe. All are het- erosanguineouslv polvgvnous and phloeopha- gus. Keys: Swaine (1918: 123), Bright (1976: 145), and Wood (1982: 656) for North Amer- ica, Reitter (1913: 102) for Europe, Bala- chowskv (1949: 255) for France, Stark (1952: 421) for the USSR. Orthotomicus Ferrari [1867: 44, Type-spe- cies: Bostrichus laricis Fabricius, subsequent 1986 WOOD: Genera of Scolytidae 69 Fig. 49. Ips spp. : 1, perturhatus (EichhoR), male, 2, same, male head; 3, .same, male declivital spines; 4, same, temale declivital spines; 5, hunteri Swaine, male; 6, woodi Thatcher, male declivity; 7, same, female head; 8, same, female dechvity; 9, pilifrona utahemis Wood, male; 10, same, female head; 11, woodi, female. (After Hopping 1965: 536). 70 GreatBasin Naturalist Memoirs No. 10 designation by Hopkins 1914: 126, original spelling Onthotomicus Ferrari (1867: 44), a lapsus calami that was corrected by Ferrari (1869: 256). Synonym: Neotomiciis Fuchs 1911: 33, Type-species: Bostrichus laricis Fabricius, subsequent designation by Hop- kins 1914: 125]. Distribution: 1 species in North America, about 10 in Asia, Europe, and N Africa. All are heterosanguineously polygy- nous and phloeophagous. Kevs: Reitter (1913; 108) for Europe, Balachowsky (1949: 268) for France, Stark (1952: 407) for the USSR. Acanthotomicus Blandford [1894a: 89, Type-species: Acanthotomicus spinosus Blandford, monobasic. Synonym: Miinips Eg- gers 1932: 33, Type-species: Ips pilosus Eg- gers, original designation; Isophthorus Schedl 1938c: 173, Type-species: Isophthorus quadrituherculatus Schedl, designated by Wood 1980: 89]. Distribution: 10 species in Central and South America, more than 20 in Africa and about 14 in China and Japan to Australia and the Philippines. All are phloeophagous; of 13 species studied in na- ture by me, 1 was monogamous and 12 het- erosanguineously polygynous. Key: Wood (1982: 664) for Central America. Ips DeGeer [1775: 190, Type-species: Tomicus tijpographus =Deniwstes tijpogra- phus Linnaeus, subsequent designation by Bergroth 1884: 230. Synonyms: Cumatotomi- cus Ferrari 1867: 44, Type-species: Bostri- chus stenographus Duftschmidt = Dennestes sexdentatus Boerner, subsequent designation by Hopkins 1914: 119; Cijrtotomicus Ferrari 1867: 44, Type-species: Bostrichus acumina- tus Gyllenhal, subsequent designation by Hopkins 1914: 120]. Distribution: 25 species in North and Central America, about 18 in Asia, Europe, and N Africa; at least 1 has been introduced to Australia and the Philippine Is- lands. All are heterosanguineously polygy- nous and phloeophagous. Numerous species belonging to other genera have erroneously been assigned to this genus at one time or another. Keys: Schedl (1950b: 69) for Europe and Asia, Hopping (1963-1965) and Wood (1982: 669) for North and Central America. Tribe Dryocoetini Dryocoetoideae Lindemann [1876: 165, Type-genus: Dryocoetes Eichhoff, 1864] Thamnurginae Niisslin [1911: 377, Type-genus: Tham- nurgus EichhofiF, 1864] Taplirorychini Reitter [1913: 29, Type-genus; Taphro- rychus Eichhoff, 1878] Description. — Frons usually sexually di- morphic, male convex to variously impressed, female convex to flattened or with elevations, variously ornamented by setae in many spe- cies; eye distinctly emarginate to divided; an- tennal scape slender, elongate, funicle 4- to 6-segmented, club obliquely truncate to strongly flattened, if flattened then sutures variously procurved to obsolete, and, on pos- terior face, strongly displaced toward apex; pronotum arched from base or not, if anteri- orly declivous then declivity usually involving more than anterior half, armed or not, if armed then asperities small, usually abun- dant; procoxae contiguous to narrowly sepa- rated; protibia with lateral margin armed by four to several socketed teeth {Chiloxylon and a few Coccotrypes have three); scutellum visi- ble; elytral declivity usually convex, some- times shallowly sulcate or variously flattened, sometimes armed by small granules; vestiture hairlike. Biology. — Apparently all are polygynous, with heterosanguineous polygyny occurring in all except the consanguineous Dryo- coetiops, Coccotrypes (Fig. 50), and Ozope- mon. Most are phloeophagous, Dactylotrypes and some Coccotrypes are spermophagous, and at least one species {Dryocoetiops cof- fcae) is myelophagous. Most eggs are placed in individual niches. Most larvae form inde- pendent mines, but some feed in congress. Taxonomy. — This is a large, rapidly evolv- ing group; consequently, generic boundaries are not always clear. Three major groups of genera are apparent; (1) Dryocoetiops, Coc- cotrypes, and Ozopemon, (2) Dactylotrypes, Dendrocranuhis , Xylocleptes, Thamnurgus, Tiarophorus, Triotemnus . etc., and (3) Cyr- togenius, Dryocoetes, Lymantor , etc. Although structural diversity may not war- rant it, these three species groups will be discussed separately. In group 1, reproduc- tion in all species involves arrhenotocous parthenogenesis. Males, if known, are dwarfed, deformed, flightless, and haploid. With about two possible American exceptions in Coccotrypes , all species originally oc- curred in the Ethiopian, Oriental, or Aus- tralian realms. Many spermophagous Coc- cotrypes have been transported through 1986 WOOD: Genera of Scolytidae 71 Fig. 50. Coccotrypes rhizophorae Hopkins; 1, entrance tunnel in Rhizophora mangle seedling; 2 and 3, larvae in host stem; 4, adult female. (After Woodruff 1970: 1). commerce far beyond their original geograph- ical distributions. Of the dozen or so Ameri- can species, all but two are known to have reached America through commerce, and it is presumed that in time these two will be found to have foreign origins. Group 2 is one of the few scolytid groups to invade herbaceous plants (mostly Cucur- bitaceae and Euphorbiaceae). A thorough study oi Thamnurgus , Xijlocleptes , and Taph- ronurgus will probably find that intergrada- tion between them is complete and, conse- quently, that they must be combined into one genus. Group 2 undoubtedly is the most an- cient of this tribe, li Dendrocranulus (Ameri- can) was separated from the almost indistin- guishable Xylocleptes (African) by early Tertiary, then the origin of their common an- cestor and of the tribe must be pushed back into the Cretaceous. Early members of this group probably gave rise to groups 1 and 3. Group 3 appears to have originated in the Old World, with Dryocoetes and Lymantor reaching northern North America rather re- cently. Schedl repeatedly called attention to the similarity between some of the small, slender Cyrtogenius and Pityophthorus ; how- ever, the antennae, elytral locking mecha- nism, and other characters are so totally dif- ferent that there is no possibility of a close relationship between these groups. Key to the Genera of Dryocoetini I . Protibia armed on lateral margin by 5 or more socketed teeth (rarely 1 or 2 reduced, if so, then sutures on anterior face of antennal club procurved, except only 2 teeth present in Dolurgocleptes); posterior face of antennal club with only one suture indicated (except 2 in some Taphronjchus), sutures on anterior face either procurved or club obliquely trun- cate, suture 1 more commonly on apical half; male of normal size, joins female in new parental gallery 2 — Protibia armed on lateral margin by 2 to 4 socketed teeth; posterior face of antennal club marked by 2 sutures, suture 1 on anterior face usually on basal third, both sutures either straight or recurved; lateral margins of prono- tum usually acutely or subacutely elevated at least near base (some exceptions); male dwarfed, of abnormal shape, flightless, never joins female in new parental gallery (except unknown in Chiloxylon ) 15 2(1). Antennal funicle 4- or 6-segmented; prono- tum unarmed, feebly if at all declivous on less than anterior fourth; eye deeply emarginate to divided; male frons moderately to strongly impressed 3 — Antennal funicle 2- to 5-segmented; anterior half of pronotum declivous and armed by as- perities (except feebly declivous and unarmed in Tlunnnnrgtis) 5 72 Great Basin Naturalist Memoirs No. 10 3(2). Antennal funicle 6-segmeiited; pronotum conspicuously longer than wide, teebly de- clivous on less than anterior half; sutures on antennal club procurved; procoxae narrowly separated; Africa; Euphorbiaceae; 1.4-.5.5 mm Tiarophorus — Antennal funicle 4-segmented, sutures of club straight or slightly recurved (when present); pronotum feebly to moderately de- clivous on more than anterior third; procoxae contiguous 4 4(3). Eye divided; protibia rather broad, margin with only 2 socketed teeth; antennal club membranous and pubescent, with 1 suture indicated at middle on front and back; prono- tum smooth, shining throughout, very feebly declivous on anterior half; striae not im- pressed, punctures small, in rows, scarcely larger than those of interstriae; Madagascar; 1.9 mm Dolurgocleptes — Eye sinuate on anterior margin; lateral mar- gin of protibia armed by 4 or 5 socketed teeth; strial punctures usually larger, deeper, dis- tinctly larger than those of interstriae; Africa to India; Euphorbia; 1.2 mm Triotemnus 5(2). Protibia more broadly flattened on at least apical half, lateral margin armed by 7 or more socketed teeth; male Irons variously im- pressed, female flat to convex and orna- mented or not; sutures on antennal club mod- estly to profoimdly procurved, except recurved in some Thamnurgus , scutellum very small, not Hat; in stems of Cucurbitaceae or Euphorbiaceae 6 — Protibia less strongly flattened on apical third, lateral margin armed by 5 socketed teeth (rarely 3 to 6); male frons convex (rarely feebly impressed), female variously modified and frequently ornamented by hair; sutures on antennal club recurved, straight, procurved, or obsolete; scutellum rather large, flat; in phloem or xylem of trees and shrubs 10 6(5). Groove on posterior face of metatibia for re- ception of tarsus poorly developed, short, oc- cupying less than apical third; Europe and Asia to Africa 7 — Metatibial groove conspicuous, occupying more than apical threcrfourths on posterior face; America and Canary Islands 9 7(6). Pronotum less distinctly declivous on anterior third, punctured to anterior margin, margins of some punctures feebly to finely asperate; antennal club rather small, sutures straight to recurved; male frons feebly impressed; Eu- rope and SW Asia to N Africa; mostly in Eu- phorbiaceae; 2.0-3.0 mm Thamnurgus — Anterior third of pronotum finely asperate, punctures obsolete in asperate area 8 8(7). Antennal club shorter than scape, 1 . 3 times as long as wide, sutures weakly procurved; male frons rather strongly impressed; SE Europe; Clematis; 1.8-2.0 mm Taphronurgus — Antennal club distinctly longer than scape, more broadK' rounded, sutures rather weakly to very strongly procurved; male frons indis- tinctly impressed; S Europe, Africa; Cucur- bitaceae, etc.; 2.0-2.7 mm Xylocleptes 9(6). Sutures of antennal club obscure, strongly procurved, basal area of club not corneous; body more slender, pronotum with asperities usually restricted to anterior half; America; Cucurbitaceae; 1.2-2,7 mm .. Dendrocranulus — Sutures of antennal club on apical fourth, re- curved, basal three-fourths of club thickened, corneous; body rather stout; asperities extend to base of pronotum; Canary Islands; Dra- caena draco; 1.8-2.0 mm Dactijlotrypes 10(5). Antennal funicle 4-segmented, sutures of club very strongly procurved; male frons shal- lowly to moderately impressed, female frons less distinctly impressed; slender species with punctures on elytral disc confused; partly xy- lophagous; North America, N Asia, Europe; 1.6-2.0 mm . Lymantor — Antennal funicle 5-segmented (rarely 2-, 3-, or 4-segmented in New Guinea and Indonesia species), sutures on club procurved or not; male frons usually convex, female frons fre- quently pubescent 11 11(10). Pronotum with summit distinctly elevated near middle, often a moderate, transverse im- pression behind summit; antennal club strongly flattened, basal area slightly cor- neous, suture 1 distinct, straight to strongly bisinuate, its lateral extremities reaching basal fourth, median portion never exceeding middle of club, 2 sutures on posterior face; meso- and metatibiae more slender; hair on pronotum and elytra fine, unusually long; Eu- rope and Asia; broadleaf hosts; 1.8-3.2 mm Taphrorychus — Pronotum with dorsal profile evenly arched, summit inconspicuous or else on basal fourth; basal area of antennal club less strongly flat- tened, more strongly corneous or if not cor- neous then sutures obsolete and pubescence extending to base; meso- and metathoracic tibiae usually more broadly flattened; prono- tal and elytral setae of more normal length . . 12 12(11). Antennal funicle 2-segmented, procurved corneous area of club occupying more than three-fourths of basal area; procoxae contigu- ous; New Guinea to New Britain Island; 1.2- 1.4 mm Dendrographus — .Antennal club 3-, 4-, or 5-segmented; usually much larger 13 1.3(12). Body very stout, 2.0-2.1 times as long as wide; pronotum rather coarsely asperate to base, summit on basal fourth; antennal club either devoid of sutures or with 1 on basal fourth, almost straight except recurved at margins; procoxae narrowly separated; declivity strongly arched, convex, apical fourth exceed- ing vertical and slightly undercutting central 1986 WOOD; Genera of Scolytidae 73 area; New Guinea to Borneo; 1.6-1.8 mm . . . Peridryocoetes — Body more slender; pronotum u.sually with punctures on at least part of basal half, summit indefinite, not on basal fourth 14 14(13). Antennal club with suture 1 weakly to pro- foundly procun'ed, rarely with sutures absent and pubescent to base; funicle of small species with as few as 3 segments (most 4- or 5-seg- mented); proco.xae narrowly separated, rarely intercoxal piece longitudinally emarginate; commonly with ventrolateral margin of de- clivity slightly elevated or armed; Micronesia to Africa; mostly non-coniferous hosts; 1.3-3.0 mm Cyrtogenius — Antennal club with suture 1 recurved, always present, never pubescent to base; procoxae contiguous, intercoxal piece always longitudi- nally emarginate or absent; ventrolateral mar- gin of declivity never acutely elevated or spe- cially armed; North America, Asia, Europe, N Africa; mostly coniferous hosts; 1.. 5-5.1 mm Dryocoetes 15(1). Protibia rather broad apically, lateral apical angle abrupt (almost 90 degrees), 1 denticle on this angle, another on apical margin, and a third on lateral margin one-fourth tibia length from apical angle; funicle 4-segmented, club constricted at partly septate suture 1 (not ac- tually obhquely truncate), suture 2 indicated by setae; habitus resembling Dryocoetes, uniseriate interstrial setae almost scalelike on declivity, declivity very steep; strial punc- tures coarse, deep; Brazil; 1.7 mm . Chiloxylon — Protibia narrowed near apex, outer apical an- gle not abrupt; funicle 4-segmented, club never septate, obliquely truncate or nearly so 16 16(15). Procoxae narrowly to moderately separated; anterior margin of pronotum usually armed by serrations (absent in some laevis); prono- tum usually more coarsely asperate in ante- rior areas, its summit more definite and near middle; India and Sri Lanka (Ceylon) to Philippines; broadleaf hosts; 1.7-3.5 mm . . . Dryocoetiops — Procoxae contiguous, intercoxal piece either longitudinally notched or absent; anterior margin of pronotum unarmed, pronotum usu- ally more finely asperate to unarmed anteri- orly, its summit less definite and usually well behind middle 17 17(16). Lateral margins of pronotum obscurely to subacutely elevated only near base; frons never convergently aciculate; anterior half of pronotum always strongly declivous and as- perate; elytral declivity either moderately im- pressed on central half and with interstrial tubercles or discal punctures strongly con- fused; mesocoxae subcontiguous, separated by distance equal to width of antennal pedicel; phloeophagous; Indonesia and Malaya to Fiji; broadleaf hosts; 3.3-5.5 mm Ozopemon — Lateral margins of pronotum subacutely ele- vated on more than basal half; frons com- monly convergently aciculate; anterior half of pronotum declivous or not; elytral declivity usually convex, rarely impressed, granules absent or inconspicuous; mesocoxae rather vndely separated by distance two or more times greater than width of pedicel; punc- tures on elytral disc almost always in rows (some exceptions); phloeophagus or sper- mophagous; introduced almost worldwide; ' apparently no endemic American species; broadleaf hosts and palm fruits; 1.3-3.7 mm Coccotryupes Tiarophorus Schreiner [1882:246, Type- species: Tiaropltorus clongatus Schreiner, monobasic. Synonyms; Hypaspistes Hage- dorn 1908: 374, Type-species: Hypaspistes camerunus Hagedorn, monobasic, preoccu- pied; Oi-thaspistes Hagedorn 1909b: 733, Type-species; Hypaspistes camerunus Hage- dorn, automatic; Pseudothamnurgus Eggers 1912a: 115, Type-species: Thamnurgus scru- tator Pandelle, subsequent designation by Schedl 1961: 738], Distribution: 13 species in Africa and S Europe. Dolurgocleptes Schedl [1965a: 61, Type- species: Dolurgocleptes malgassicus Schedl, monobasic]. Distribution: 1 species in Mada- gascar. Triotemnus WoUaston [1864: 264, Type- species: Triotemnus subretusus Wollaston, monobasic. Synonym: Cladoctoproctiis Schedl 1975: 454, Type-species: Cladocto- proctus scrafa Schedl, original designation]. Distribution: 8 species in N Africa and the Canary Islands, 2 in India. Most species breed in the stems of Euphorbia, Delphinium, Aconitum, Teucrium, and Bupierum . Key: Peyerimhoff(1949). Thamnurgus Eichhoff [1864: 40, Type- species: Bostrichus euphorhiae Kiister, sub- sequent designation by Hopkins 1914; 130]. Distribution: About 40 species in Asia, Eu- rope, and Africa. A thorough review of the species of this and the ne.xt two genera would probably result in combining them into one genus. Keys: Reitter (1913: 85) for Europe, Balachowsky (1949: 165) for France. Taphronurgus Reitter [1913: 84, 90, Type- species: Thamnurgus exul Reitter, monoba- sic]. Distribution: 1 species in SE Europe to SW Asia. In Clematis. It is probable that this 74 Great Basin Naturalist Memoirs No. 10 genus will eventually be combined with Thamnurg,its . Xylocleptes Ferrari [1867: 37, Type- species: Bostrichiis bispinus Duftschmidt, monobasic. Synonyms: Xestips Hagedorn 1912: 353, Type-species: Xestips marginatus Hagedorn, monobasic; Hijlonitis Nunberg 1973: 16, Type-species: Hijlonius hrunneus Nunberg, original designation]. Distribution; About 20 species in S Asia, S Europe, and Africa. Key: Reitter (1913; 91) for Europe. Dendrocranulus Schedl [1937a: 165, Type- species: Dendrocranulus tardus Schedl, sub- sequent designation by Schedl 1938c: 169]. Distribution: 22 species in North and Central America, about 15 in South America and adja- cent islands. Some species are monogamous, some are heterosanguineously polygynous; all bore in dving stems of Cucurbitaceae. Key: Wood (1982: 708) for North and Central America. Dactylotrypes Eggers [1927a: 37, Type- species: Dactylotrypes tiyttenboogaarti Eg- gers =Xyloterus longicoUis Wollaston, mono- basic]. Distribution: 1 species in the Canary Islands in fruits of Phloenix canahensis and Dracaena drace ; it may have been introduced into France (Balachowsky 1949: 186). The male apparently joins the female in the parental gallery. Lymantor Lovendal [1889; 25, 68, Type- species: Lymantor sepicola Lovendal =Tomi- cus cort//t Ferris, monobasic]. Distribution; 2 species in North America, 2 in Asia and Eu- rope. All are heterosanguineously polygynous and xylophagous in small, dead branches of living trees. Keys: Reitter (1913: 91) for Eu- rope, Wood (1982: 707) for North America. Taphrorychus Eichhoff [1878a: 49, 204, Type-species: Bostrichiis bicolor Herbst, subsequent designation by Hopkins 1914: 130. Synonyms: Saliciphilus Sokanovskii 1954; 17, 20, Type-species: Hypothenemus machnovskii Sokanovskii, original designa- tion; Pseudopoecilips Murayama 1957: 614, Type-species: Pseudopoecilips mikuniyamen- sis Murayama, original designation; Taphrot- erus Schedl 1965b; 341, Type-species: Taphroterus primitus Schedl, monobasic]. Distribution: About 12 species in Asia, Eu- rope, and N Africa. All are phloeophagous. Keys: Reitter (1913: 94), Balachowsky (1949; 193), PfeflFer (1962: 241). Dendrographtis Schedl [1964c; 310, Type- species; Pelicerus pygmaeus Eggers, original designation. Synonym: Protopityophthorus Schedl 1973: 73, Type-species: Protopityoph- thorus durus Schedl =Pelicerus pygmaeus Eggers, original designation]. Distribution: 1 species in New Guinea and New Britain Is- land. Peridryocoetes Wood [1984; 230, Type- species; Ozodendron nitens Schedl, original designation, also cited but not described as a new genus by Schedl 1964a; 243]. Distribu- tion: 3 species from Indonesia and New Guinea. Cyrtogenius Strohmeyer [1910c; 127, Type-species; Crytogenius bicolor Stroh- meyer, monobasic. Cyrtogenius Strohmeyer (1911: 16) a valid emendation oi Kyrtogenius . Synonyms: Carposinus Hopkins 1915a: 9, 47, Type-species; Carposinus pini Hopkins, orig- inal designation: Orosiotes Niisima 1917: 1, Type-species: Orosiotes kumatoeti,sis Ni- isima, monobasic; Metahylastes Eggers 1922: 165, Type-species: Metahylastes africanus Eggers, monobasic; Pelicerus Eggers 1923: 216, Type-species: Lepicerus nitidus Hage- dorn, original designation; Eulepiops Schedl 1939b; 344, Type-species: Eulepiops glaber Schedl, original designation; Ozodendron Schedl 1957a: 13 and duplicate description by Schedl 1964a: 243, Type-species: Pelicerus grandis Beeson, monobasic; Mimidendrulus Schedl 1957a; 68, Type-species; Mimiden- dridus movoliae Schedl, monobasic, the spe- cies designated by Schedl (1961: 732) was not an original species and is an invalid designa- tion; Carpophloeus Schedl 1959b; 143, Type- species; Carpophloeus rugipennis Schedl, monobasic; Taphroborus Nunberg 1961: 617, Type-species; Taphroborus vaticae Nunberg, original designation; Artepityophthorus Schedl 1969: 157, Type-species; Artepityoph- thorus aries Schedl, monobasic; Taphro- stenoxis Schedl, nomen nudum, for Taphro- stenoxis tenins Schedl, nomen nudum]. Distribution: About 16 species in Africa, more than 40 in the area from India and Japan to Australia and Micronesia. They are phloeophagous and heterosanguineously polygynous. Dryocoetes Eichhoff [1864; 38, Type- species; Bostrichus autographus Ratzeburg, subsequent designation by Hopkins 1914; 1986 WOOD; Genera of Scolytidae 75 121. Synonyms: Anodius Motschulsky 1860: 155, Type-species: Bostrichus autographiis Ratzeburg, subsequent designation by Wood 1974: 232, suppressed by International Com- mission on Zoological Nomenclature 1979: 149; Dryocoetinus Balachowsky 1949: 180, Type- species: Bostrichus villosiis Fabricius, original designation]. Distribution: 7 species in North America, about 80 in Asia, Europe, and Africa have been assigned to this genus, but more than half of them have been trans- ferred to other genera. All are heterosan- guineously polygynous and phloeophagous. Keys: Reitter (1913: 75) for Europe, Mu- rayama (1957: 594) for Japan and vicinity. Bright (1963: 107) and Wood (1982: 724) for North America. Chiloxylon Schedl [1959c: 550, Type- species: Chiloxylon nifulus Schedl, monoba- sic]. Further study of this genus is needed; it may be allied to Dendroterus of the Corthylini (Pityophthorina), rather than to Dnjocoetes. Distribution: 1 species in Brazil (Matto Grosso). Dryocoetiops Schedl [1957a: 13, Type- species: Ozopemon laevis Strohmeyer, origi- nal designation]. Distribution: About 15 spe- cies in SE Asia, Sri Lanka (Ceylon), Indone- sia, and New Guinea. One species observed in nature appeared to be consanguineously polygynous (or some form of parthogenesis) and was myelophagous. Ozopemon Hagedorn [1908: 382, Type- species: Ozopemon regius Hagedorn, mono- basic]. Distribution: About 27 species in SE Asia and Indonesia to Fiji. All are consan- guineously polygynous and phloeophagous. Coccotrypes Eichhoff [1878a: .308, Type- species: Bostrichus dactyliperda Fabricius, subsequent designation by Hopkins 1914; 118. Synonyms: Poecilips Schaufuss 1897: 110, Type-species: Poecilips sannio Schau- fuss, monobasic; Cryphaloides Formenek 1908: 91, Type-species: Cryphaloides don- isthorpei Formenek = Bostrichus carpopha- gus Homung, monobasic; Thamnurgides Hopkins 1915a: 45, Type-species: Thamnur- gides persicae Hopkins =Coccotrypes adoena Blandford, original designation; Spernm- toplex Hopkins 1915a: 48, Type-species: Spermutoplex rhizophorae Hopkins, original designation; Dendrurgus Eggers 1923: 144, Type-species: Dendrurgus rhizophorae Eg- gers = Spermatoplex rhizophorae Hopkins, subsequent designation by Wood 1982: 731; Hyphaene Hagedorn 1913: 254, nomen nudum. Type-species: Hyphaene guineensis Hagedorn, nomen nudum = Bostrichus car- pophagus Hornung, no status]. Distribution: More than 100 species have been assigned to this genus, mostly from Africa, S Asia, and adjacent areas. Species in most other areas have been introduced through commerce. All are consanguineously polygynous and phloeo- phagous or spennophagous; a few species may assume both feeding habits. This e.xceedingly difficult genus is in a state of ta.\onomic chaos. Key: Wood (1982: 732) for North America. Tribe Crypturgini Crypturgi LeConte [1876: 374, 387, Type-genus; Cryp- turgus Erichson, 1836] Description. — Frons usually not dimor- phic, male sometimes slightly impressed, fe- male convex; eye deeply emarginate, except sinuate in Deropria ; antennal scape moder- ately long, slender, funicle 2- to 3-segmented, club comparatively small, two sutures on api- cal half, 1 sometimes septate, both frequently absent; pronotum usually unarmed; procoxae contiguous; protibia flattened, lateral margin armed by several socketed denticles; scutel- lum visible; elytral punctures in rows or con- fused, setae hairhke; anterior surfaces and (frequently) elytra uniformly reticulate. Biology. — They are monogamous and phloeophagous except that those species asso- ciated with Euphorbia bore throughout subepidermal tissues of recently killed stems, apparently with little regard to the cambium. Crypturgus usually utilizes the entrance tun- nel of another insect. Larval mines are inde- pendent. Details of the habits have not been studied. Taxonomy. — This tribe apparently was derived from the same ancestral stock that gave rise to the Dryocoetini, although the true affinities have not been worked out. It appears to have arisen well into the Tertiary in the Ethiopian realm where at least some members of five of the six known genera occur today. Crypturgus appears to have reached North America from Asia rather recently. Dolurgus (North America) appears to be a primitive relict of an earlier radiation that is now extinct except for this species. 76 Great Basin Naturijlist Memoirs No. 10 Key to the Genera of Crypturtiini 1. Antennal funicle 3-segmented, club with su- tures 1 and 2 recurved, clearly marked by grooves and rows of setae, 1 almost at middle of club; pronotum longer than wide, unarmed, a distinct constriction on anterior third; elytra rather coarsely striate, strial setae largely obso- lete, interstrial setae short; W North America; Abies, Picea, Pinus, 1.6-1.9 mm Dolurgus — Antennal funicle 2-segmcnted, club with no more than 1 suture indicated or if 2 present then club constricted at septate suture 1 2 2(1). Antennal club rather strongly constricted at middle at suture 1; elytral punctures confused or striae, if indicated, not impressed, strial punc- tures little larger than those of interstriae .3 — Antennal club not constricted at middle, suture 1 obsolete or indicated only by a partial, internal septum 4 3(2). Elytral declivity strongly flattened to concavely impressed; pronotum wider than long, weakly il at all declivous on anterior half, never armed by granules, anterior margin unarmed; N Africa, Euphorbia; 1.6-2.6 mm Coleobothrus — Elytral declivity convex; pronotum usually longer than wide, anterior half more strongly declivous, often armed by granules, anterior margin elevated as a costa, serrate, or dentate; Canary Islands, Africa to India; Euphorbia ; 1.3- 2.3 mm Aphanarthrwn 4(2). Antennal club unmarked by sutiu'es, anterior half of pronotum declivous and armed by coarse asperities, anterior margin serrate; eye entire to feebly sinuate; Canary Islands; host unknown; 1.5-2.0 mm Deropria — Antennal club with suture 2 marked at apex, if present, suture 1 sometimes indicated by an internal septum; pronotum weakly declivous, unarmed, anterior margin never serrate; eye deeply emarginate 5 5(4). Remnants of sutures 1 and 2 usually present in antennal club; elytral striae more strongly im- pressed, punctures rather coarse, always in rows; elytral vestiture on disc short, longest se- tae shorter than distance between interstrial rows; North America, Asia, Europe. N Africa, Pinaceae; 0.9-1.5 mm Crypturgus — Antennal club usually without indications of su- tures; elytral striae usually not impressed, punc- tures much smaller and usually contused; discal vestiture longer, longest setae distinctly longer than distance between interstrial rows; Canary Islands, S Europe, N Africa; Euphorbia , 1.1-1.7 mm Cisurgus Dolurgus Eichhoff [1868a: 147, Type-spe- cies: Hylastes pumilus Mannerheim, monoba- sic]. Distribution: 1 species in W North Amer- ica (Alaska to N California). It is monogamous and phloeophagous. Coleobothrus Enderlein [1929: 144, Type- species: Coleobothrus jandiacus Enderlein, original designation]. Distribution: 2 species in the Canary Islands and 2 in Africa. Key: Menier (1973: 208). Aphanarthrum WoUaston [1854: 292, Type-species: Aphanarthrum euphorbiae Wollaston, monobasic]. Distribution: About 30 species in the Canary Islands, Africa, and India. Key: Schedl (1959a: 56). Deropria Enderlein [1929; 143, Type-spe- cies: Aphanarthrum elon^atum Eggers, origi- nal designation]. Distribution: 1 species in the Canary Islands. Crypturgiis Erichson [1836: 60, Type-spe- cies: Bostrichus pusillus Gyllenhal, desig- nated by Thomson 1859: 147]. Distribution: 3 species in North America, 10 in Asia, Europe, N Africa, and adjacent islands. All are monog- amous and phloeophagous. Keys: Reitter (1913: 61) for Europe, Swaine (1918: 54) and Wood (1982: 740) for North America, Schedl (1946: 4) for Europe and Asia, Bright (1976: 114) for Canada. Cisurgus Reitter [1894: 59, 65, Type-spe- cies: Cisurgus filumReitter, monobasic]. Dis- tribution: 6 species in SW Asia, S Europe, N Africa, and the Canary Islands. Key: Schedl (1959: 28). Tribe Xyloterini Xyloteroideae Lindemann [1876: 165, Type-genus: Xi/loterus Erichson, 1836 =Trypodendron Stephens, 18,30] Trypodendron Tredl [1907: IS, Type-species: Trypoden- dron Stephens, 1830] Description. — Frons dimorphic, male weakly to very strongly impressed, female convex; eye completely divided; antennal scape long, funicle 4-segmented (Fig. 51), club without sutures, basal area sometimes corneous, derived from obliquely truncate type; procoxae contiguous, proepimeron of female with mycetangium; pronotum asper- ate on anterior slope; protibia flat in male, inflated and armed by small unsocketed denti- cles on posterior face in female; scutellum visible; elytra conservatively sculptured. Biology. — All species are monogamous and xylomycetophagous. The male joins the female in parental galleries. The eggs are de- posited in niches above and below in the egg tunnels. The larvae enlarge the niches into 1986 WOOD: Gener.\ of Scolytidae 77 lITbetulae l8.T.retu5um IMIineatum 20.T. rufilarsis 2I.T.scabricollis 22.)(. politus Fig. 51. Xyloterini spp.: 1, Xyluterinua politus (Say), outline of female; 2 and 5, Trypodcndron Uneatum (Olivier), female; .3, T. lineatum , male; 6, Indocnjpludus aceris (Niisima), female; 7, T. lineatum , male; 8, T. lincutum , female; 9, X, politus, female protibia; 10, T. hetulae Swaine, male head; 11, X. politus, male head; 12, T. rufitarsis (Kirby), female; 13, T. rufitarsis, male; 14, X. politus, female antenna; 15, T. lineatum female antenna; 16, X. politus, male antenna; 17-22, male genital capsiil, dorsal a.speet, a.s labeled. 78 Great Basin Naturalist Memoirs No. 10 Fig. .52. Xijlebonis spathipennis Eichhoft: A, outline of male, dorsal aspect; B, dorsal aspect of female; C, female declivity (setae omitted). cradles just enough to accommodate the newly transformed adult beetle. The brood emerges through the parent entrance tunnel. Temperate species overwinter in litter on the forest floor. Taxonomy. — This is a small group consist- ing of three small genera. They appear to have been derived from the same parental stock that gave rise to the Xylehorini. They appar- ently originated in Asia, with Trypodendron extending westward into Europe and east- ward in late Tertiary to northern North Amer- ica. Xyloterinus (North America) and In- docryphalus (Asia) were derived from the same parental stock, with Xyloterinus appar- ently reaching North America in the warm period that preceded the last ice age. Key to the Genera of Xyloterini (Modified from Wood 1957b: .344) 1. Basal area of antennal club distinctly subcor- neous, its apical margin strongly, rather nar- rowly procurved on anterior face; protibia thick- ened, posterior face tuberculate in female, flattened and usually finely tuberculate in male; male frons broadly, deeply excavated from epis- toma to vertex, convex in female; male prono- tum subquadrate, anterior margin straight to slightly recurved and unarmed, female anterior margin procurved and armed by several teeth; long axis of proepimeral excavation of female longitudinal and very narrow; North America, N Asia, Europe; coniferous and broadle;if trees; 2. 7-4.6 mm Trypodendron — Basal area of antennal club broadly, weakly pro- curved or else not at all corneous; protibia flat- tened and unarmed on posterior face in both sexes; anterior margin of pronotuni procurved and armed by a series of teeth in both sexes .... 2 2(1). Basal area of antennal club not thickened or corneous, uniformK' pubescent to base; males and females equal in size; anterior margin of pronotum slightly produced at median line in male; long axis of proepimeral excavation of fe- male longitudinal, short to long, narrow to rather broad; SE Asia; broadleaf hosts; 2.6-3.5 inm Indocryphalus — Basal area of antennal club subcorneous, thick- ened, weakly prociu'ved on anterior face; male distinctly smaller than female; anterior margin of male pronotum as in female except some teeth reduced or absent; proepimeral excavation of female transverse, rather large, broad; North America; broadleaf hosts; 2.7-3.7 mm . Xyloterinus Trypodendron Stephens [1830: 353, Type- species: Dennestes domesticiis Linnaeus, subsequent designation by Westwood 1838: 39 and Thomson 1859: 146. Synonym: Xy- loterus Erichson 1836: 60, Type-species: Bostrichus lineatus Olivier, subsequent des- ignation by Thomson 1859: 146]. Distribu- tion: 5 species in North America, about 9 in Asia and Europe. All are monogamous and xylomycetophagous. Keys: Schedl (1951c: 86) for Europe and Asia, Wood (1957b: 345, 1982: 747) for North America. Indocryphalus Eggers [1939: 5, Type-spe- cies: Indocryphalus malaisei Eggers =Xy- 1986 WOOD: Genera OF ScoLYTi DAE 79 loterus intermedius Sampson, original desig- nation. Synonym: Dendrotrypum Schedl 1951c: 76, Type-species: Xijlotcrua accris Ni- isima, subsequent designation by Browne 1970: 76, Type-species: Xyloterus aceris Ni- isima, subsequent designation by Browne 1970: 562]. Distribution: About 9 species in Asia (India to Japan). Apparently all are monogamous and .xylomvcetophagous. Key: Schedl (1951c: 77).' Catalog: Browne 1970: 562). Xyloterinus Swaine [1918: 44, 83, Type- species: Bostrichiis politus Say, original des- ignation]. Distribution: 1 species in E North America. It is monogamous and .xylomyce- tophagous. Tribe Xyleborini Xylebori LeConte [1876: .346, 358. Type-genus: Xtjle- borus Eichhoff, 1864] Webbinae Hopkins [1915b: 224, Type-genus: Webbia Hopkins. 1915] Description. — Body dimorphic, male dwarfed, deformed, flightless, eye reduced in size, often aberrant in shape (Fig. 52); frons convex, unadorned; eye emarginate to di- vided in a few oriental forms; antennal scape elongate, funicle 5-segmented except 3- or 4-segmented in a few oriental forms, club obliquely truncate except basal corneous area reduced or absent in some genera; pronotum asperate on anterior slope (a few exceptions, especially in male), procoxae varying from contiguous to widely separated; scutellum varying fi-om large and flat to modified to ab- sent; elytra variable, conservatively to elabo- rately sculptured; meso- and metatibiae flat, broad, tapered on distal third, lateral margin armed by a row of numerous, small, closely set socketed teeth, these usually alternating with marginal or submarginal setae in more highly evolved forms; meso- and metatarsi retractible into tibial grooves. Biology. — Consanguineous polygyny is universal, apparently all males are haploid, deformed, and flightless. All are xylomyce- tophagous. Eggs are deposited in clusters in the parental tunnels. The larvae usually ex- tend the parental galleries or feed exclusively on the fungal mycelium in the parental tun- nels. The brood emerges through the parental entrance tunnel. Temperate species may overwinter either in the brood host or in litter on the forest floor. At least one species (Xylehorus dispar) passes through a definite dipause in the adult stage during the winter months. Ta.XONOMY. — The worldwide circumtropi- cal distribution of this tribe, with a few species occurring in temperate areas, suggests that it is at least moderately old. The occurrence of the same species groups of Ambrosiodmus in Africa and South America indicates that the basic characters of groups within that genus had been fixed by early Tertiary. However, the distributions of other groups and the large number of species in the tribe indicate that very rapid, recent evolution is in progress. In this tribe, arrhenotocous parthenogenesis is universal. This suggests a possible relation- ship to the higher Dryocoetini; however, a relationship to the Xyloterini is more likely. Because of the recent, active evolution affect- ing this group, generic limits are not clearly defined. Common and particularly confusing features of this tribe are size races within what otherwise appears to be a single species; in some cases these behave as entirely different species and in others, involving those same forms, there is total intergradation of the two sizes. This is probably a product of this type of mating system that will be more fully under- stood when details of their habits are known. The generic classification of this tribe pre- sented below is tentative and flawed; how- ever, it is presented as a first attempt to orga- nize a very large and difficult group even though a third of the species in the tribe were not studied. It is hoped that it might give at least limited direction to those who will con- centrate more particularly on this group. Key to the Genera of Xyleborini (Females only) I. Basal segment of labial palpus eylindrical, only slightly wider than segment 2 or 3, none of segments ornamented by a special tuft of setae; pregula and adjacent surfaces flush with general contour of ventral surface of head; antennal club rather strongly flattened, with- out visible sutures, corneous area reduced, usually pubescent to base; lateral margins of pronotum acutely elevated, pleural area con- cave (on transverse axis); anterior margin of pronotum unarmed; protibia inflated, armed on its posterior face by minute tubrercles; Africa; 2 species introduced into America; 1.4-4.7 mm Premnobius 80 Great Basin Naturalist Memoirs No. 10 — Basal segment of labial palpus enlarged, con- spicuously wider than segment 2 or 3, its pos- terior face usually flat, one or more segments ornamented by specialized setae; pregula and adjacent areas usually conspicuously im- pressed below general contour of head 2 2(1). Antennal club with sutures 1 and 2 rather strongly procurved, both segments 1 and 2 corneous and mostly glabrous e.vcept at su- tures; protibia slender, almost cylindrical, posterior face armed by tubercles; lateral margins of pronotum acutely elevated, pleural area transversely concave; body very slender; anterior margin of pronotum armed by 2 or more very coarse serrations; domicile parasites of other ambrosia beetles; Mexico to Brazil; 2.7-6.0 mm Sampsonius — Antennal club obliquely truncate or nearly so, sutures (when visible) on or very near margin of corneous area, recurved (except pubescent to base in some oriental forms); protibia usu- ally more strongly expanded on apical hall; lateral margins of pronotum rounded (except Cnestiis. Webbia); "parasitic habit unknown 3 3(2). Scutellum visible, moderately large, its sur- face flush with adjacent surface of elytra; ven- tral margin of metiifeniur either rounded or rather obtusely angulate, on its posterior face groove for reception of tibia usually clearly indicated on distal half 4 — Scutellum either not visible or (Xyleborinus) base ol elytra at suture notched thereby ex- posing cone-shaped scutellum that appears displaced cephalad, or (couplet 20) scutellum visible on anterior face of declivital slope of elytral bases; scutellar area usually with abun- dant setae associated with mycetangiuni; metafemur usually more strongly flattened, its (longitudinal) ventral margin attenuately, very acutely angulate (except Schedlia), groove for reception of tibia visible only near apical joint 16 4(3). Posterior face of antennal chib marked by 2 sutures on apical third (suture 2 poorly repre- sented in some Coptoborus), anterior face with apical portion convex (or concave only distad from segment 2), segment 2 compara- tively large, sclerotized, protibia armed by 6 or 7 socketed teeth on lateral margin, metat- ibia with 6 to 9 socketed teeth; anterior coxae always contiguous .5 — Posterior face of antennal club with no more than 1 suture visible at or very near apex (usually none), apical portion of anterior face usually flat to concave, segment 2 (it visible) not corneous; number of metatibial teeth vari- able; anterior coxae contiguous or separated . 8 5(4). Protibia with posterior face inflated and armed by numerous fine tubercles; metatibia usually with 8 or 9 socketed teeth; segment 2 on antennal club usually forming a complete, oblique annulus, on anterior face its apical margin usually acutely costate as on segment 1; Mexico to South America (1 species intro- duced into Africa); 2. 1-.5.3 mm . . Dryocoetoides — Protibia with posterior face flat, unarmed, metatibia almost never with more than 7 sock- eted teeth 6 6(5). Declivity commencing at or anterior to mid- dle of elytra, its lower half transversely, broadly impressed and either flat or shallowly concave; if discal interstrial punctures unise- riate then declivital surface covered by dense, confused, small scales, if declivital setae hair- like then discal interstrial punctures rather dense, confused; India to Philippines and New Guinea; 2.2-5.0 mm Leptoxyleborus — Elytral declivity usually convex, commencing either posterior to middle or strongly tapered on posterior half never with a strong trans- verse impression; elytra never ornamented by scales 7 7(6). Posterior fourth ot elytra comparatively broad, rather broadly rounded behind, suture never emarginate; declivital interstriae 1 to 3 similar, tubercles minute, if present; body comparatively stout, less than 2.6 times as long as wide; Mexico to South America; intro- duced to Africa; 1.7-3.0 mm Theoborus — Posterior third of elytra attenuate or acumi- nate, narrowly rounded behind, suture often emarginate; 1 or more declivital interstriae sometimes armed by small denticles; body slender, at least 2.6 times as long as wide; Mexico and South America, Africa to SE Asia; 1.5-5.0 mm Coptoborus 8(4). Protibia with posterior face inflated and armed by numerous fine tubercles; posterior face of antennal club either with or without a suture; elytral declivity and at least part of disc with interstriae carinate (carinae some- times reduced to rows of tubercles), strial spaces between carinae usually granular or dull, with punctures usually obsolete; SE Asia to Australia; 1.3-3.1 mm Arixyleborus — Protibia with posterior face almost flat, smooth; elytra with different sculpturing .... 9 9(8). Antennal club with segment 2 on anterior face usually conspicuous, sometimes rather large, apical margin of segment 1 on both faces rounded, often inconspicuous or absent on anterior face, almost always visible on subapi- cal area of posterior face; procoxae always con- tiguous, intercoxal piece longitudinally emar- ginate, posterior element of intercoxal piece never inflated or armed, mesocoxae usually more widely separated by distance greater than thickness of scape 10 — Segment 1 of antennal club corneous, its dis- tal margin very acutely elevated into a contin- uous costa (forming a complete circle) extend- ing from anterior face to apex, suture almost never visible on posterior lace; procoxae vary- ing from contiguous to widely separated, if 1986 WOOD; Genera of Scx)lytidae 81 contiguous then posterior intercoxal piece sometimes inflated and armed; mesocoxae usually subcontiguous, usually separated by distance less than thickness of scape 13 10(9). Pronotal asperities extending to base, includ- ing most of discal area (except numerous African, etc. species without discal asperi- ties); anterior margin of pronotum never armed by a definite row of serrations; lateral margin of porotibia armed by 7 to 8 socketed denticles, metatibia by 8 to 11 denticles; pantropical; 1.9-4.2 mm Ambrosiodmus — Pronotal asperities confined to slightly more than anterior half; lateral margin of metatibia armed by more than 11 socketed denticles . . 11 11(10). Elytra obliquely, abruptly truncate, usually with an acutely, distinctly elevated circumde- clivital, subcircular costa, face of declivity flat to concave (when costa incomplete then an- tennal club pubescent to base); discal inters- triae with punctures uniseriate, occasionally slightly confused on some interstriae (strongly confused in superficially similar mancus group oi Xijlosandrus), antennal club usually pubescent to its base, more strongly flat- tened; eye deeply emarginate to entirely di- vided; SE Asia to Australia; 2.4-.3. 1 mm . Amasa — Declivital margin rounded, a carina not indi- cated on its basal two-thirds, declivital lace basically convex; antennal club obliquely truncate (except pubescent to base in a few Xyleborus 12 2(11). Pronotum commonly subquadrate, its ante- rior margin almost always unarmed, postero- lateral margin of declivit\' subacutely elevated from sutural apex to interstriae 7; strial and interstrial punctures usually in rows, elytral vestiture comparatively sparse, confined to strial and interstrial rows; Asia and Australia to Africa; 2.4-4.6 mm Emcallacva — Pronotum almost never quadrate, its anterior margin usually procurved and armed by a definite row of serrations; posterolateral mar- gin of declivity rounded; elytral pimctures often confused, vestiture usually much more abundant; SE Asia to Africa; 2.1-4,1 mm . . . Terminalinus 13(9). Proeoxae contiguous, intercoxal piece longi- tudinally emarginate (a few individuals in /«- vaniis group subcontiguous but only feebly emarginate), its po.sterior element inflated, occasionally dentate; body usually more slen- der, mostly more than 2.0 times as long as wide; tropical and temporate areas world- wide; 1.7-.5.9 mm Xyleborus — Either funicle 4-segmented or if ,5-scgmented then proeoxae moderately to rather widely separated, intercoxal piece continuous, not longitudinally emarginate (a slight notch in a lew Taurodemtis); body usually stouter, mostly less than 1 . 9 times as long as wide ... 14 14(13). Lateral margin of protibia armed by 9 to 12 socketed teeth; elytral declivity moderately to very strongly sulcate on at least basal half, lateral margins armed by at least 1 major spine and several smaller tubercles; antennal funi- cle .5-segmented; Central and South America; 2.4-4.5 mm Taurodemus — Lateral margin of protibia armed by 4 to 7 socketed teeth; elytral declivity usually not conspicuously sulcate, lateral margins not conspicuously armed by spines or tubercles . 1.5 15(14). Lateral margins of pronotum rounded (sub- acute in morigerus); funicle 5-segmented; corneous area (in central portion) occupying basal third of antennal club; eye about one- third divided by an emargination; protibia armed on lateral margin by 4 to 6 socketed teeth; pantropical, 1.3-5.0 mm ... Xylosandrus — Lateral margins (and usually basal margin) of pronotum subacutely elevated, costate; an- tennal funicle 4-segmented (some 5-seg- mented?); corneous area (in central portion) occupying less than basal fourth of antennal club; eye feebly if at all emarginate; protibia armed on lateral margin by 6 to 8 socketed teeth; SE Asia to Australia; 2.0-4.0 mm .... Cncstus 16(3). Basal margins of elytra distinctly, shallowly emarginate in scutellar area, emargination usually filled by dense setae, scutcllum dis- placed slightly cephalad into a visible, conical process; posterior face of antennal club with- out sutures; protibia flat, its lateral margin armed by 6 to 8 socketed denticles, all on apical half; strial punctures in rows; declivity usually armed by tubercles or spines; tropical and temporate areas worldwide; 1.4-3.5 mm Xijleborinus — Basal margins of elytra not emarginate at su- ture, scutcllum not conical, with other combi- nations of characters 17 17(16). Antennal funicle 5-segmented; lateral margin of protibia armed by socketed denticles on less than apical half 18 — Antennal funicle 3- or 4-segmented; protibia inflated but not asperate on posterior face, its lateral margin armed (usually to base) by sock- eted teeth on more than apical three-fourths 23 18(17). Pronotum asperate to base, including discal area; basal corneous area occupying almost three-fourths length of club, its apical margin straight, not acutely elevated, suture 2 clearly indicated (club more reminiscent of Hylastcs than oi Xyleborus). suture 1 visible near apex on posterior fiice; .scutellum visible on ante- rior slope of elytral base; pimctiu'es on elytral disc confused on basal third, in obscure rows behind; strial and interstrial hair moderately long; all tibiae very broad; New Zealand; 1.9 mm Mesoscolytus 82 Great Basin Naturalist Memoirs No. 10 — Discal area of pronotum unarmed by asperi- ties, smooth; basal corneous area of antennal club occupying less than basal half in central area, its apical margin recurved 19 19(18). Posterior face of antennal club unmarked by sutures, acute costa marking apical margin of corneous area usually forming a complete ring; scutellum usually visible on anterior de- clivous slope of elytral margins; body stout (less than 1.8 times as long as wide), usually black in color; punctures on elytral disc dense, confused 20 — Posterior face of antennal club with at least 1 suture visible, apical margin of corneous area never costate; scutellum not visible on ante- rior slope at bases of elytra; body more slen- der (at least 2.0 times as long as wide), color usually yellowish to reddish brown 21 20(19). Metatibia normal, socketed denticles present on lateral margins; metatarsus normal, nei- ther unusually long nor abnormally com- pressed; declivity mostly restricted to poste- rior half of elytra, convex or moderately impressed, unarmed; India to Philippines and Australia; 4.0-6.0 mm Hadrodemius — Metatibia abnormally large and broad, sock- eted denticles on lateral margin obsolete; metatarsus unusually long, very strongly compressed; declivity extending almost to base of elytra, armed by very coarse spines; Philippines and New Guinea to Africa; 1.1-4.0 mm Eccoptopterus 21(19). Protibia inflated and densely asperate on pos- terior surface; anterior face of mesotibia simi- larly but less strongly inflated and armed; ely- tral disc smooth and shining, abruptly becoming rugose-reticulate on declivity, dis- cal punctures confused, declivity variously armed by rather large spines; Sumatra to New Guinea; 4.0-.5.0 mm Schedlia — Pro- and mesotibiae neither inflated nor armed on posterior face; declivity never densely, uniformly rugose-reticulate, if armed then spines small; size much smaller . 22 22(21), Strial punctures in definite rows (except con- fused in bellus group); declivity convex, with- out a marginal costa, variously sculptured; elytral scales never present; SE Asia and In- donesia to Philippines; 1.4-3.4 mm Coptodryas — Elytral punctures small, confused; declivity commencing on basal half of elytra, very strongly concave, its marginal crest forming a blunt, elongate, circumdeclivital costa, inner face ornamented by abundant, small scales or longhair; Indonesia; 3.3-3.5 mm Taphrodasus 23(17). Elytra moderately acuminate on declivity and mucronate at sutural apex; base of declivity rounded, unarmed; base of elytra often with a submarginal mycetangium; Malaya; 2.2 mm Crijptoxylehorus — Elytra variously convex or truncate at base of declivity, never acuminate or mucronate at sutural apex; declivital margin abrupt, usually armed; SE Asia to New Guinea and Philip- pines; 1.5-3.0 mm Webbia Premnobius Eichhoff [1878a: 65, 404, Type-species: Fremnohius cavipennis Eich- hoff, monobasic. Synonym: Premnophilus Browne 1962a: 79, Type-species: Xyleborus joveri Schedl, original designation]. Distribu- tion: 24 species in Africa, 2 of them were introduced into America. All are consan- guineously polygynous and xylomyceto- phagous. Key: Wood (1982: 756). Sampsonius Eggers [1933a: 23, nomen nudum, 1935: 157, Type-species: Sampsonius sexdcntatiis Eggers, original designation]. Distribution: 4 species in S Mexico and Cen- tral America, about 7 species in South Amer- ica. All are domicile parasites of other am- brosia beetles and are consanguineously polvgynous and xylomycetophagous. Key: Wood (1982: 759) for Central America. Dryocoetoides Hopkins [1915a: 10, 52, Type-species: Dryocoetoides guatemalensis Hopkins =Xyleborus capucinus Eichhoff, original designation]. Distribution: 2 species in Central America, about 22 in South Amer- ica 1 of which was introduced into Africa. All are consanguineously polygynous and xy- lomycetophagous. Key: Wood (1982: 762) for Central America. Leptoxyleborus Wood [1980: 94, Type-spe- cies: Phloeotrogus sordicauda Motschulsky, original designation]. Distribution: 4 species in SE Asia. All are consanguineously polygy- nous and xylomycetophagous. Theoborus Hopkins [1915a: 57, Type-spe- cies: Theobonis theobromae Hopkins, origi- nal designation]. Distribution: 9 species in Central America, several additional species in South America belong here. All are consan- guineously polygynous and xylomyce- tophagous. Key: Wood (1982: 770) for Central America. Coptoborus Hopkins [1915a: 10, 53, Type- species: Coptobortts emar^natus Hopkins =Xyleborus oespatorius Schedl, original des- ignation. Synonym: Streptocranus Schedl 1939a: 52, Type-species: Streptocranus mirabihs Schedl, luonobasic]. Distribution: 5 species in Central America, about 10 in South America, and about 6 in SE Asia and adjacent islands. All are consanguineously polygynous 1986 VVooD: Gener\ of Scolytidae 83 and xylomycetophagous. Key; Wood (1982: 780) for Central America. Arixyleborits Hopkins [1915a: 10, 59, Type- species: Arixyleborus rugosipes Hopkins, original designation. Synonym: Xyleboriciis Eggers 1923: 212, Type-species: Xylcboricus canaUculatus Eggers, subsequent designa- tion by Schedl 1936b: 64]. Distribution: About 42 species in SE Asia to Australia and the Philippines. All are consanguineously polygynous and xylomycetophagous. Ambrosiodmus Hopkins [1915a: 10, 55, Type-species: Xyleborus tachygraphiis Zim- mermann, original designation. Synonyms: Phloeotrogits Motschulsky 1863: 127, Type- species: Phlocotrogus obhquccaudata Mot- schulsky, designated by Hopkins 1914: 127, International Commission on Zoological Nomenclature 1979: 151, name rejected; Browneia Nunberg 1963; 37, Type-species; Xyleborus illepidus Schedl ^Pityophthorua obliquus LeConte, original designation]. Dis- tribution; 11 species in North and Central America, about a dozen in South America, numerous species (40 or more) in Asia and Africa. All are consanguineously polygynous and xylomycetophagous. Key: Wood (1982: 781) for North and Central America. Amasa Lea [1894; 322, Type-species: Amasa thoracicus Lea, monobasic. Syn- onyms: Pseudoxylebonis Eggers 19.30; 206, Type-species; Pseudoxylebonis beesoni Eg- gers, monobasic; Anaxyleborus Wood 1980: 90, Type-species; Tomiciis truncatus Erich- son, original designation]. Distribution: About 35 species in India and Malaya to Aus- tralia. All are consanguineously polygynous and xylomycetophagous. Euwallacea Hopkins [1915a: 10, 54, Type- species; Xyleborus loallacei Blandford, origi- nal designation]. Distribution: About 50 or more species in Africa to SE Asia and Aus- tralia. All are consanguineously polygynous and xylomycetophagous. Terminalinus Hopkins [1915a: 10, 57, Type-species: Terminalinus terminaliae Hop- kins, original designation. Synonyms; Kelan- tanius Nunberg 1961; 621, Type-species: Xyleborus punctatopilosus Schedl, original designation]. Distribution; About 30 or more species in Africa to SE Asia and Australia. All are consanguineously polygynous and xy- lomycetophagous. Xyleborus Eichhoff [1864; 37, Type-spe- cies: Bostricluis itKiiiograjihus Fabricius, sub- sequent designation by Lacordaire 1866; 381. Synonyms; Anisandrus Ferrari 1867; 24, Type-species: Xyleborus (Apate) dispar Fab- ricius, monobasic; Anaeretus Duges 1887: 141, Type-species: Xyleborus guanajuatensis Duges =Bostrichus volvulus Fabricius, monobasic, neotype for type-species desig- nated by Wood 1983: 650; Progenius Bland- ford 1896a: 20, Type-species; Progenius fleu- tiauxi Blandford =Xyleborus subcostatus Eichhoff, subse(|uent designation by Hopkins 1914; 128; Cyclorhipidion Hagedorn 1912: 355, Type-species; Cyclorhipidion pelliculo- sum Hagedorn -Xyleborus prelaetus Schedl, monobasic; Heteroborips Reitter 1913; 79, 82, Type-species: Bostrichus cryptographus Ratzeburg, monobasic; Xyleborips Reitter 1913: 79, 111, Type-species; Xyleborus meuscli Reitter, monobasic; Boroxylon Hop- kins 1915a: 10, 58, Type-species; Boroxylon stephegynis Hopkins Phloeotrogus biden- tatus Motschulsky, original designation; Notoxyleborus Schedl 1934; 84, Type-species: Notoxyleborus kalshoveni Schedl, monoba- sic]. Distribution; Several hundred species almost worldwide. All are consanguineously polygynous and xylomycetophagous. Keys; Bright (1968; 1296, 1976; 131) for North America, Wood (1982; 778) for North and Central America, Reitter (1913; 81) for Eu- rope. Taurodetnus Wood [1980: 96, Tvpe-spe- cies: Xyleborus sharpi Blandford, original designation]. Distribution; 14 species in S Mexico to South America. All are consan- guineously polygynous and xylomyceto- phagous. Key: Wood (1982: 778) for Central America. Xylosandrus Reitter [1913; 80, 83, Type- species; Xyleborus morigerus Blandford, monobasic. Synonym; Apoxyleborus Wood 1980: 90, Type-species; Xyleborus mancus Blandford, original designation]. Distribu- tion; 8 species in USA to South America, about .32 species in SE Asia to Australia and Africa. All are consanguineously polygynous and xylomycetophagous. Key: Wood (1982: 765) tor North and Central America. Cnestus Sampson [1911: 383, Type-spe- cies; Cnestus magnus Sampson, monobasic. Synonym: Tosaxyleburus Murayama 1950a; 84 Great Basin Naturalist Memoirs No. 10 49, Type-species: Tosaxylcboriis pallidipcn- nis Murayama =Cnestus miiraijamai Schedl, original designation]. Distribution: About 17 species in SE Asia to Indonesia, the Philip- pines, and Japan. All are consanguineously polygynous and .xylomycetophagous. Kev: Nunberg (1972: 476). Xyleboriniis Reitter [1913: 79, 83, Type- species: Bostrichus saxeseni Ratzeburg, sub- sequent designation by Swaine 1918: 50]. Dis- tribution: 8 species in North and Central America, at least 4 in South America, at least 20 in SE Asia to Africa. All are consan- guineously polygynous and .xylomyceto- phagous. Key: Wood (1982: 842) for North and Central America. Mesoscolytus Broun [1904: 125, Type-spe- cies: Mesoscolijtiis inurbanus Broun, mono- basic]. Distribution: 1 species in New Zealand. It is consanguineously polygynous and xylomycetophagous. Hadrodemius Wood [1980: 94, Type-spe- cies: Xylebonis globus Blandford, original designation]. Distribution: About 6 species in SE Asia to Indonesia and the Philippines. All are consanguineously polygynous and xy- lomycetophagous. Eccoptopterus Motschulsky [1863: 515, Type-species: Eccoptopterus sexspinosiis Motschulsky =Scohjtits spinosits Olivier, monobasic. Synonyms: Platydactyhis Eich- hoff 1886: 25, Type-species: Platydactylus gracilipes Eichhoff, monobasic; Eurydactyhis Hagedorn 1909: 733, Platydactylus gracilipes Eichhoff, automatic]. Distribution: About 12 species in SE Asia to Australia and adjacent islands, and Africa. All are consanguineously polygynous and xylomycetophagous. Schedlia Browne [1950: 641, Type-species: Xyleborus sumatranus Hagedorn, original designation]. Distribution: 5 species in SE Asia to Indonesia and New Guinea. All are consanguineously polygynous and xylomyce- tophagous. Key: Browne (1950: 642). Coptodryas Hopkins [1915a: 10, 54, Type- species: Coptodrijas confusa Hopkins, origi- nal designation. Synonyms: Microperus Wood 1980: 94, Type-species: Xyleborus theae Eggers, original designation; Adry- ocoetes Eggers, nomen nudum, in Schedl (1952: 371), Type-species: Adryocoetes ni- tidus, nomen nudum, =Xtjleborus pullus Schedl, no status]. Distribution: About 20 species in SE Asia to Australia and adjacent islands. All are consanguineously polygynous and xylomycetophagous. Taphrodasiis Wood [1980: 95, Type-spe- cies: Xyleborus percorthylus Schedl, original designation]. Distribution: 3 species in SE Asia to Indonesia. All are consanguineously polygynous and xylomycetophagous. Crijptoxijleborus Schedl [1937b: 550, Type-species: Crijptoxijleborus naevus Schedl, subsequent designation by Schedl 1962a: 103]. Distribution: About 12 species in Malaya. All are consanguineously polygynous and xylomycetophagous. Webbia Hopkins [1915b: 222, Type-spe- cies: Webbia dipterocarpi Hopkins, original designation. Synonyms: Xelyborus Schedl 1939b, nomen nudum (Browne 1963a: 57); Pseudowebbia Browne 1961: 308, Type-spe- cies: Xyleborus trepanicauda Eggers, original designation; Prowebbia Browne 1962b: 208, Type-species: Proivebbia subuculae Browne, original designation]. Distribution: About 32 species in SE Asia to Indonesia and the Philip- pines. All are consanguineously polygynous and xylomycetophagous. Key: Browne (1962b: 210) to the pabo group of species. Tribe Xyloctonini Xyloctonidae Eichhoff [187Sa: 171. Tvpe-genus: Xyloc- fonus Eichhoff, 1872] Description. — Frons apparently not di- moiphic, usually unadorned; eye emarginate to divided; antennal scape elongate, funicle 6- or 7-segmented, club strongly flattened, su- tures procurved, present or obsolete, 1 partly septate or not; pronotum asperate on anterior slope, anterior margin usually armed; pro- coxae contiguous; scutellum large, flat; tarsi retractable into tibial grooves; venter of abdo- men moderately to very strongly ascending to meet elytra. Biology. — All are apparently monoga- mous and phloeophagous. The egg galleries are monoramous in Ctonoxijlon and biramous in Scolijtomimus . The eggs are deposited in- dividually in niches and sealed in by frass. The larval mines radiate out from the parental gallery and may be rather long. Taxonomy. — This is a small group of pre- dominantly African genera that are poorly known. It appears to be a primitive branch of the same phyletic line that gave rise to the 1986 WooD; Genera of Scolytidae 85 Cryphalini. Together these two tribes appear to occupy a position intermediate between primitive Micracini and Corthyhni. A princi- pal distinguishing character oi Xyloctonini is their abihty to totally withdraw the meso- and metatibiae into tibial grooves. This character is shared by a small group of genera in the Cryphalini that are allied to Scolytogenes ; thus, tribal placement for them is dependent on the number of segments in the antennal funicle. The limited distribution of this tribe, accompanied by conspicuous anatomical di- veristy, makes comments on its anticjuity diffi- cult. Key to the Genera of Xyloctonini 1. Eye entire to feebly emarginate; suture 1 of antennal club septate, almost straight; funicle 7-segmented; elytra declivous behind, abdo- men raised only slightly to meet them, inters- triaecostate; Africa; 1.5-2.1 mm . Cryphalomimus — Eye moderately emarginate to divided; sutures of antennal club weakly to strongly procurved .. 2 2(1). Eye moderately emarginate, about one-third di- vided by an emargination; basal (usually) and lateral margins of pronotum rounded; funicle 6-segmented, sutures of antennal club feebly to moderately procur\ed, 1 not septate; elytral de- clivity steep, abdomen ascending very slightly; Africa; 1.9-2.6 mm Glostatiis — Eye more than half divided by an emargination to completely divided; basal and lateral margins of pronotum with a fine, raised line (either con- tinuous or beaded); sutures of antennal club strongly procurved, 1 partly to entirely septate (when sutures reduced septum remains); abdo- men moderately to very strongly ascending to meet moderately to feebly declivous elytra .... .3 3(2). Antennal funicle 7-segmented, club moderately flattened, asymmetrical, comparatively small, about equal in length to scape, suture 1 with posterior half septate; elytral declivity rather steep, abdomen ascending moderately to meet apex; eye always divided; Africa; 1.8-.3.6 mm . . Ctonoxylon — Antennal funicle fj-segmented, club strongly flattened, symmetrical, large, conspicuously longer than scape, suture 1 partly septate or not; elytral declivity short, very gradual, abdomen very strongly ascending to meet apex; eye di- vided or not 4 4(3). Antennal club devoid of sutures except for strongly procurved septum in posterior half of suture 1; scutellum large, flat, subtriangular. its surface flush with that of base of elytra, India to Philippines and Fiji; 1.0-2.4 mm . , Scohjtomimus — Antennal club with 2 or 3 very strongly pro- curved sutures, none of them septate; scutellum averaging smaller, subquadrate, adjacent basal area on interstriae 1 and 2 impressed thereby causing scutellum to project slightly dorsad; Africa; 1. 3-2.8 mm Xyloclonus Cryphalomimus Eggers [1927b: 174, Type- species: Cnipluilomimus striatum Eggers, monobasic]. Distribution: 3 species in Africa (Congo to East Africa). Glostatus Schedl [1939d: 386, Type-spe- cies: Glostatiis decUvidepressiis Schedl, monobasic. Synonyms: Ctonocryplms Schedl 1941: 398, Type-species: Ctonocryplms xyloc- tomis Schedl, monobasic; Apoglostutus Schedl 1957a: 155, Type-species: Apoglo- statiis acaciae Schedl, monobasic; Para- glostaus Schedl 1964c: 304, Type-species: Ctonocryplms uigrivcstris Schedl, original designation; Rhopalocryplms Nunberg 1967: 320, Type-species: Rhopalocryphus seycleli Nunberg]. Distribution: 16 species in Africa. Apparently all are monogamous and phloeo- phagous. Ctonoxylon Hagedorn [1910c: 4, Type-spe- cies: Ctonoxylon auratiim Hagedorn, subse- quent designastion by Hopkins 1914: 119, Schedl's (1961: 426) designation is invalid]. Distribution: About 32 species in Africa. Ap- parently all are monogamous and phloeo- phagous. Scolytomimus Blandford [1895: 319, Type- species: Scolytomimiis dihitiis Blandfoi'd, monobasic. Synonyms: Neoxyloctonus Eggers 1923: 143, Type-species: Neoxyloctonus philippincnsis Eggers, monobasic; Scolyto- clcptes Schedl 1962f: 490, Type-species: Scolytoniinms maculatns Beeson, original designation]. Distribution: About 15 species from India and Sri Lanka (Ceylon) to Samoa. All are monogamous and phloeophagous. Xyloctonus Eichhoff [1872: 134, Type-spe- cies: Xyloctonus scolytoidcs Eichhoff, mono- basic]. Distribution: 14 species in Africa. All are monogamous and phloeophagous. Key: Menier (1974: 658). Tribe Cryphalini Cryphaloidea Lindemann [1876: 16.5, Type-genus: Cniphalus Erichson. 1836] Trypophloeinae Niisslin [1911: 373, Type-genus: Try- pophldcufi F'airmaire, 1868] Ernoporinae Niisslin (1911; 375, Type-genus: Ernoponis Thomson, 1859] Eidophelinae Murayama [1954; 200, Type-genus: Eido- phcltifi EichhofI, 1875, amended from Eidopheri- nae by Wood 1978; 114] 86 Great Basin Naturalist Memoirs No. 10 Fig 1 Slephanoderes female Fig 3 Stephanoderes mole Fig 2 Stephonoderes female Fig 4 Stephonoderes mole Fig. 5 Pityophtfionni Fig 8 Crypholus Fig 10 Crypfiolomorpfius Fig 6 Procrypholus fig 7 Procrypholus FiQ 9 Crypholus Fig II Cryphalomorphus Fig. 53. Cryphalini, tribal and generic characters: 1-2. Hypofhcnemus (^Stcph(Hwderes) dissimilis (Zinimermann), outline of female; 3-4, same, male; 5, Pseudopityophthonts pubipcnnis (LcConte) (Corthylini), showing consealed metepisternum and horizontal abdomen; 6, Procnjphalus uiahcnsis Hopkins, anterior aspect of antennal club; 7, same, posterior aspect; 8, Tnjpophloeus (-Cryphalus) popiili Hopkins, anterior aspect of antennal club; 9, same, posterior aspect; 10, Scolytogenes {^Cryphalomorphus) knabi (Hopkins), anterior aspect of antennal club; 11, same, posterior aspect. 1986 WOOD: Genera of Scolytidae 87 Description. — Frons rarely dimorpliic, usually convex, usually not adorned; eye usu- ally entire, shallowly emarginate in a few gen- era; antennal scape elongate, simple, funicle 3- to 5-segmented, club moderately to strongly flattened, sutures present or obso- lete, 1 sometimes septate, sutures on poste- rior face strongly displaced toward apex; pronotum coarsely asperate on anterior slope (Fig. 53), anterior margin usually armed, lat- eral and basal margins usually (not always) marked by a finely elevated line; procoxae contiguous; tibiae rather strongly flattened, their lateral margins armed by more than four socketed teeth; metepisternal spine usually modified, often partly replaced liy a callus or modified groove; scutellum large, flush with adjacent surface of elytra; basal margins of elytra sometimes marked by a fine, raised line; elytral sculpture usually conservative, their costal margins near apex usually ascend- ing at least slightly; venter of abdomen usually rising slightly to meet elytra; vestiture com- monly includes scales. Biology. — Monogamous, except for con- sanguineous polygyny in Hypothenemiis, Crtjptocarentts, Feriocryphahis , and Trischi- dias, and either phloeophagous or myelo- phagous. Mycetophagous in one species of Hypothenemiis . Parental gallery usually of a cave type, often modified into crude, elon- gate, brood chambers. The eggs are deposited in clusters, often mixed with frass; definite egg niches have not been reported. The larvae may form individual mines {Trypophloeiis, Cryphalus) or feed in congress as they extend the parental chamber (ProcrypJialus, Hy- pothenemiis, etc.). The brood may emerge through individual exit holes or through the parental entrance timnel, depending upon the larval habit. In temperate areas the winter is most commonly passed in the larval stage; two generations per year is the common habit. Taxonomy. — The worldwide distribution and structural diversity of this tribe suggest a pre-Tertiary origin. However, the prolifera- tion of large numbers of species, particularly in Cryphalus, indicates that recent, very rapid evolution is in progress. Trypophloeiis, Procryphalus, Ernoporicus , and Cryphalus appear to have reached North America from Asia rather recently. Hypoihencmus , Crypto- carenus, Trischidias , and Scolytogenes in North America were recent dcrivitives from the neotropical realm. The South American element of Cryphalini appears to contain very ancient, endemic elements {Acorthylus, Ncocnjphus, Stegomerus, Cryptocarenus) and more recent arrivals (Hypothenemiis, Scolytogenes). The more recent elements are largely confined to the tropics and are shared with Africa; similarity of species groups sug- gest a connection of faunas as late as early Tertiary. The main center of distribution for the tribe appears to be in the Oriental and Australian realms. This is also the area where rapid evolution has produced a great abun- dance of very closely related species. In this area, generic limits are often obscure, whereas in other parts of the world they are quite distinct. This is probably the most poorly known tribe in the world and it is likely to remain so until much more material is avail- able for study. Key to tlie Genera of Cryphalini 1 . Basal and lateral niargin.s ot pronotum rounded; procoxae usually narrowly sepa- rated, intercoxal piece continuous (except contiguous and intercoxal piece longitudi- nally emarginate in Trypophloeiis, Ste- gomerus, Acoi-thijlus) 2 — Basal and usually basal third of lateral margins of pronotum marked by a finely raised line; procoxae either contiguous or narrowly sepa- rated; eye entire or narrowly emarginate .... 8 2(1). Eye shorter, less than twice as long as wide, entire (rarely with a few lacets absent suggest- ing a weak, narrow emargination) 3 — Eye moderately elongate, 2.0 or more times as long as wide, more than half of its anterior margin occupied by a sinuation or broad emargination one-third as deep as width of eye; neotropical genera 6 3(2). Antennal funicle 5-segmented, club slender (1.8 or more times as long as wide), 2 sutures clearly marked, its apex subacutely pointed (Fig. .52); elytral interstriae 10 continuing to apex; North America, N Asia, Europe; Salix, Popiilus, AIniis, 1.4-2.1 nun Trypophloeiis — Antennal lunicle 3- to 5-segmented, club broad (less than 1.3 times as long as wide), its apex rather Inoadly rounded, suture 1 usually marked, 2 obsolete or nearly so; interstriae 10 obsolete before attaining level of base ol ab- dominal sternum 5 4 4(3). Eye entire; antennal club longer, basal area more stnmgly flattened, suture 1 straight, septate. 2 obsolete; funicle 4-segmented; fe- male irons not conspicuously pubescent; 88 Great Basin Naturalist Memoirs No. 10 North America to NE Asia; Populus, Salix. Fraxinus; 1.3-2.2 mm Procryphaliis — Eye entire to weakly sinuate; antennal club with sutures procurved or absent, 1 not sep- tate when present .5 5(4). Antenna! funicle 4-segmented, club with pro- curved sutures indicated by rows of setae; E North America, N Asia, Europe; Fraxinus, Fagus, Cornus; 1.2-1.7 mm Ernoporicus — Antennal funicle .3-segmented, club devoid of sutures; NE Asia; Euomjmus ; 1.4 mm Allernoporus 6(2). Antennal huiicle .5-segmented, club large, subcircular, about as wide as long, sutures conspicuously procurved; discal striae not im- pressed, poorly defined (punctures often con- fused); Mexico to South America; vines (lianas); 1.1-1.9 mm Stegomerus — Antennal funicle 3-segmented, club elongate, at least 1.5 times as long as wide; sutures almost straight 7 7(6). Antennal funicle shorter than scape, seg- ments 2 and 3 small, subequal in size; elytral striae impressed, punctures rather coarse; S South America; host unknown; 1.4-1. .5 mm Neocryphux Antennal fimicle with segment 2 greatly en- larged, as long as scape; elytral striae not im- pressed, strial rows usually not distinguish- able (at least one exception); South America; Pruntis. etc.; 1.2-1.8 mm Acorthylus 8(1). Basal margin of pronotum marked by a fine, raised line, lateral margin rounded; sutures on antennal club procur\'ed, usually distinct . 9 — Both basal and lateral margins of pronotum marked by a fine, raised line; sutures on an- tennal club present or absent, variable 12 9(8). Eye emarginate; pronotal asperities confused; antennal grooves or sutures moderately pro- curved 10 — Eye entire; pronotal asperities arranged in concentric rows; antennal svitures rather strongly to profoundly procurved or obsolete ' 11 10(9). Antennal funicle 5-segmented, club with su- tures 1 and 2 marked by moderately pro- curved grooves and rows of setae; pronotal asperities coarse, confused; eye conspicu- ously emarginate; Africa; hosts unknown; 2.0- 2.3 mm Stephanopodius — Antennal funicle 4-segmented, club with su- ture 1 septate and angulate; eye rather small, shallowly emarginate; SE Asia; 1.2-1.6 mm Coriacephilus 11(9). Antennal funicle 4-segmented; vestiture of abundant, confused scales; sutures on anten- nal club very strongly procurx'ed or obsolete; Europe, Asia; Tilia, Fagus, etc.; 1.1-1.5 mm Ernoporus — Antennal funicle 3-segmented; vestiture of uniseriate rows of scales on discal interstriae, a few supplemental scales on declivity; su- tures on antennal club rather weakly pro- curved, sometimes obscure; SE Asia; broadleaf trees; 1.1-1.5 mm Ernocladius 12(8). Posterior face of metatibia (usually also mesotibia) with a groove for reception of tar- sus on lateral half from apex at least two-thirds of distance toward base, grooved area glabrous, usually with a row of setae along its mesal margin, tibia usually more broadly flat- tened, gradually tapered on its distal third, with socketed teeth more numerous and dis- tributed over at least apical third (possible confusion with Xyloctonini, except eye entire in these Cryphalini); male subequal in size to female, capable of flight 13 — Metatibia either without groove for reception of tarsus or groove restricted to less than distal one-fifth of tibial length, setae randomly dis- tributed on its lateral half, tibia usually sub- truncate apically, socketed teeth usually re- stricted to apical one-fifth; male either normal or deformed 16 13(12). Antennal club with suture 1 partly septate, suture either straight or strongly procurved, if straight then funicle 3-segmented); body usu- ally stouter, pubescence moderately abun- dant; groove foriTiing lateral line on pronotum rather poorly defined 14 — Antennal club with suture 1 aseptate (or ob- scurely septate, if suture present then funicle 4-segmented; body rather slender, elytral vestiture sparse, largely confined to declivity; groove forming lateral and basal raised line on pronotum rather strongly impressed 15 14(10). Antennal funicle 3-segmented, club with su- tures 1 and 2 weakly procurved, clearly marked by setae, 1 also grooved and partly septate; venter of abdomen horizontal; India to Sri Lanka (Ceylon); Euphorbia, 1.0-1.3 mm Cryphalogenes — Antennal funicle 4-segmented, club with su- tures strongly to profoundly procurved, 1 marked on mesal half by a septum, remaining sutures marked by setae or obsolete (septum complete in several small New Guinea spe- cies); venter of abdomen weakly to very strongly ascending to meet apex of elytra, apex of elytra usually ascending also; pantrop- ical; hosts usually lianas; 1.0-2.5 mm Scolytogenes 1.5(13). Antennal club with sutures 1 and 2 weakly procurved and clearly marked by rows of se- tae, 1 also grooved; basal half of pronotum reticulate or minutely rugose, punctures small to obsolete; Micronesia; hosts unknown; 1.1-1.4 mm Hemicryphalus — Antenuiil club with sutures entirely obsolete; basal hall of pronotum smooth, shining, with a few coarse punctures; India and E USSR to Japan; P/if//o(/ci!rfroii; 1.2-1.3 mm . Eidophelus 1986 WooD: Genera of Scol-^tidae 89 16(12). Tarsal segment 3 rather broad, bilobed; pro- coxae narrowly separated, interco.\al piece not longitudinally emarginate; eye emar- ginate; antennal club with aseptate sutures clearly marked by grooves and setae 17 — Tarsal segment 3 narrow, often laterally coin- pressed, not bilobed, procoxae contiguous, intercoxal piece longitudinally emarginate or partly absent 18 17(16). Antennal funicle 5-segmented, sutures on club weakly to rather strongly procurved; phloeophagous in broadleaf trees; pantropi- cal; 1.1-2.8 mm Hypocryphalus — Antennal funicle 4-segmented, sutures on club recurved (occasionally weakly pro- curved); phloeophagous in broadleaf and coniferous hosts; E Hemisphere, North America (1 dubious record from South Amer- ica); 1.0-2.8 mm Cryphalus 18(16). Antennal funicle 3-segmented, club with or without sutures, never with a septum, male normal, not dwarfed or flightless 19 — Antennal funicle 3- to .5-segmented, sutures always indicated on club, 1 frequently partly septate; male dwarfed, flightless 21 19( 18). Antennal club with moderately procurved su- tures clearly marked by grooves and rows of setae; eye emarginate; striae obsolete, pinic- tures on disc confused; SE Asia to Philippines; 0.8-1.3 mm Margadillius — Antennal club without indications of grooves or rows of setae; eye entire, strial punctures in recognizable rows 20 20(19). Anterior margin of pronotum armed by 0-6 serrations, asperities on anterior half of pronotum larger, less numerous; body stouter, 1.4-2.0 times as long as wide; Irons convex to flattened in both sexes; SE Asia to Hawaii; O.S-1.8 mm Ptilopodius — Anterior margin of pronotum armed by 10-16 serrations, asperities smaller, much more nu- merous, often extending to base; body more slender, 2.5-2.6 times as long as wide; female frons rather narrowly impressed; SE Asia and Indonesia to Micronesia; 1.3-1.5 mm Cosmoderes 21(18). Antennal funicle 3- to 5-segmented, club with or without sutures, when funicle 3-seg- mented then club always with suture 1 partly septate 22 — Antennal funicle .3-segmented, club never septate, sutures sometimes marked by rows of setae; body rather stout, 2.0-2.3 times as long as wide; very small 23 22(21). Anterior margin of pronotum armed by 10-16 serrations; antennal funicle 5-segmented, club never septate, sutures marked by rows of setae; mature body color yellowish to reddish brown; vestiture usually very sparse (rare ex- ceptions); myelophagous; tropical America (1 introduced to Africa); 1.4-3.3 mm Cryptocareniis — Anterior margin of pronotum usually armed by 1-8 serrations; antennal hmicle 3- to 5-seg- mented, when 5-segmented then suture 1 of club partly .septate; antennal club clearly marked by sutures; mature color usually darker, vestiture more abundant; phloeo- phagous, myelophagous, spermopliagous, 1 mycetophagous, worldwide in tropical and warm temperate areas; 0.6-2.8 mm Hypothenemus 23(21). Antennal club with sutures clearly marked by rows of setae; eye entire; phloeophagous in trees and shrubs; E North America to Mexico and Hawaii; 0.6-1. 1 mm Trischidias — Antennal club without indications of sutures; eye emarginate, myelophagous in \'ines. South America; 0.8-1.1 mm ... Periocryphaltts Trypophloeus Fairmaire [1868; 105, Type- .species: Bostrichiis binodiihis Ratzehurg, monobasic. Synonym; Glyptoderiis EichhofF 1878a: 34, 44, 137, Type-species; Bostrichiis binodulus Ratzeburg, subsequent designa- tion by Hopkins 1914; 122]. Distribution; 4 species in North America, about 12 in N Asia and Europe. All are monogamous and phloeophagous. Keys; Wood (1954b; 989, 1982; 851) for North America, Reitter (1913; 69), Balachowsky (1949; 214), Stark (1952: 281) for Europe and Asia. Procryphalus Hopkins [1915a; 7, 33, Type- species; Procryphahis popiili Hopkins = Cryphalus miicronatus LeConte, original designation]. Distribution; 2 species in North America (in Popuhts, Salix), 1 in NE Asia (Ussuri, in Fraxiniis). All are monogamous and phloeophagous. Key; Wood (1954b; 982, 1982; 859) for North America. Ernoporicus Berger [1917; 242, Type-spe- cies: Ernoporiciis spessivtzevi Berger, mono- basic. Synonyms: EocryphahiS Kurentzov 1941: 161, 230, Type-species; Eocrypliahis se- menovi Kurentzov, monobasic; Ernopocerus Balachowsky 1949: 211, Type-species: Ernoporus caucasicus Lindemann, subse- quent designation by Wood 1954b: 986]. Dis- tribution; 1 species in North America (West Virginia), about 12 in Europe and Asia. All are apparently monogamous and phloeophagous. Keys: Balachowsky (1949: 211), Reitter (1913; 68)', Stark (1952: 271). Allernoporus Kurentzov [1941: 159, Type- species; Alh'rnoponis eiionymi Kurentzov, monobasic]. Distribution; 1 species in NE Asia (in Euonynuts). It apparently is monoga- mous and phloeophagous. 90 Great Basin NATUiiALiST Memoirs No, 10 Stegomerus Wood [1967; 129, Type-spe- cies; Stegomerus vulgaris Wood, original des- ignation]. Distribution; 5 species in Mexico and Central America, 1 in South America (Venezuela). All are monogamous and phloeophagous. Keys: Wood (1967: 130, 1982; 855). Neocryphus Nunberg [1956a; 139, Type- species; Neocryphus argentinensis Nunberg, original designation]. Distribution: 2 species in South America (Argentina). Acorthyltis Brethes 1922a: 304, Type-spe- cies; Acot-tlnjliis asperatiis Brethes, monoba- sic. Synonym: Phacryliis Schedl 1938a: 24, Type-species: Phacryliis hosqid Schedl, monobasic]. Distribution: About 5 species in South America. All are monogamous and phloeophagous. Stephanopodhis Schedl [1941: 396, Type- species; Stcphanodcres dispar Eggers, subse- quent designation by Schedl 1961: 633. Syn- onyms; Cryphalomimus Browne 1962a; 75, Type-species: Hypocryphalus ghanaensis Schedl, original designation, preoccupied; C ryphalomiinetes Browne 1963b: 242, Type- species: Hypocryphalus ghanaensis Schedl, automatic]. Distribution: About 7 species in Africa. Coriacephilus Schedl [1939b; 339, Type- species: Stephanoderes coriaceiis Eichhoff, original designation]. Distribution; About 4 species in SE Asia to Philippines. Ernoporus Thomson [1859: 147, Type-spe- cies: Bostrichus tiliae Panzer, original desig- nation. Synonyms; C ryphalops Reitter 1889: 94, Type-species; Cryphahis lederi Reitter = Bostrichus tdiae Panzer, monobasic; Steph- anorhopalus Hopkins 1915a; 35, Type-spe- cies: Stephanorhopcdus melodori Hopkins, original designation (specific name of type- species validly amended by Schedl 1966a: 19); Euptilius Schedl 1940b; 589, Type-species; Ernoporus concentralis Eggers, original des- ignation]. Distribution: At least 13 and per- haps 20 species in Europe and S Asia to the Philippines. All are monogamous and phloeo- phagus. Ernocladius Wood [1980: 93, Type-species: Cryphalus corpulentus Sampson, original designation]. Distribution: About 4 species in S Asia to Sri Lanka (Ceylon). Cryphalogenes Wood [1980: 91, Type-spe- cies; Cryphalogenes cuphorbiae Wood, origi- nal designation]. Distribution: 4 species in India to Sri Lanka (Ceylon) in Euphorbia . All are monogamous and excavate dying tissue immediately under the epidermis. Scohjtogenes Eichhoff [187Sa; 475, 497, 1878b: 387, Type-species; Scolytogenes dar- wini Eichhoff, monobasic. Synonyms: Lep- icerus Eichhoff 1878a; 476, 501, Type-spe- cies: Lepicenis aspericollis Eichhoff, mono- basic; C ryphalomorphus Schaufuss 1890; 12, Type-species: C ryphalomorphus communis Schaufuss, monobasic; Letznerella Reitter 1913; 68, Type-species: Bostrichus jalapae Letzner, monobasic; Hypothenoides Hopkins 1915a: 7, 11, Type-species; Hypothenoides parvus Hopkins, original designation; Ernoporides Hopkins 1915a: 8, 34, Type-spe- cies: Ernoporides floridensis Hopkins = Ernoporides knabi Hopkins, original desig- nation; Neocryphalus Eggers 1922: 169, Type-species; Neocryphalus usagaricus Eg- gers, monobasic; Negritus Eggers 1923; 141, Type-species; Negritus ater Eggers, desig- nated by Wood 1982: 861; Cylindrotomicus Eggers 1936: 6.33, Type-species: Cylindroto- micus squamulosus Eggers, monobasic; Lep- icerinus Hinton 1936: 473, Type-species: Eep- icerus aspericollis Eichhoff, automatic; Xylocryptus Schedl 197.5c: .352, Type-species: Xylocryptus papuanus Schedl, original desig- nation]. Distribution; 7 species in North and Central America; about 60 in other tropical and subtropical areas around the world. All are monogamous and almost all bore in the stems of vines (lianas). Key: Wood (1960: 27) for Micronesia, (1982; 862) for North and Cen- tral America. Hemicryphalus Schedl [1963b: 264, Type- species; Eidophelus argutus Wood, original designation]. Distribution; 3 species in Mi- cronesia (W Pacific Islands). Key: Wood (1960a; .32). Eidophelus Eichhoff [1875: 200, Type-spe- cies: Eidophelus iniitans Eichhoff, monoba- sic. Synonym: Phellodendrophagus Krivolut- skaya 19.58; 150, Type-species: Phellodend- rophagus elegans Krivolutskaya, monobasic]. Distribution: About 8 species in SE Asia and Indonesia. Hypocryphalus Hopkins [1915a; 8, 41, Type-species; Hypocryphalus rotundus Hop- kins, original designation. Synonym: Dacry- phalus Hopkins 1915a: 8, 42, Type-species: 1986 WOOD: Genera of Scolytidae 91 Dacryphahis ohesus Hopkins, original desig- nation]. Distribution: About 47 species in Africa, S Asia to Australia and Samoa; 1 intro- duced into America {mangiferae , in Mang- ifera indica ). All apparently are monogamous and phloeophagous. Cryphalus Erichson [1836: 61, Type-spe- cies: Bostrichus asperatus Gyllenhal, subse- quent designation by Thomson 1859: 147, lec- totype for type-species designated by Wood 1972; 41. Synonyms: Pseudocryphalus Fer- rari 1869: 252, Type-species: Bostrichus sid- neyanus Nordlinger, monobasic; Taenio- glyptes Bedel 1888: 398, Type-species: Bostrichus abietus Ratzeburg =Bostrichus asperatus Gyllenhal, original designation; Cryptarthrum Blandford 1896b: 200, Type- species: Cryptarthrum walkeri Blandford, monobasic; Allarthrum Hagedorn 1912: 355, Type-species: Allarthrum kolbei Hagedorn, monobasic; Ericryphalus Hopkins 1915a: 8, 38, Type-species: Ericryphalus henslunoi Hopkins =Hypotheneinus sylvicola Perkins, original designation; Piperius Hopkins 1915a: 8, 39, Type-species: Piperius pini Hopkins =Hypothenemus sylvicola Perkins, original designation; Ernocryphaliis Murayama 1958: 934, Type-species: Ernocryphalus hirosimen- sis Murayama, original designation; Acry- phalus Tsai & Li 1963: 604, 622, Type-spe- cies: Cryphalus lipingensis Tsai & Li, designated by Wood 1984: 224; Jugo- cryphalusTsai & Li 1963: 602, 622, Type-spe- cies: Cryphalus piceus Eggers, designated by Wood 1984: 224]. Distribution: 3 species in North America, 1 dubious record from South America, 11 in Africa, about 7 in Europe, more than 200 nominate species have been reported from Asia to Australia and adjacent islands. All are monogamous and phloeo- phagous. Keys: Reitter (1913: 66) for Europe, Balachowskv (1949: 206) for France, Stark (1952: 254) for USSR, Wood (1954b: 1002, 1982: 867) for North America and (1960a: 23) for Micronesia. Margadillius Hopkins [1915a: 8, 37, Type- species: Margadillius margadilaonis Hop- kins, original designation]. Distribution: About 9 species in SE Asia, Philippines, New Guinea, and Indonesia. The true status and extent of this genus have not been estab- lished. Key: Hopkins (1915a: 37). Ptilopodius Hopkins [1915a: 7, 11, Type- species: Ptilopodius stephegynis Hopkins, original designation]. Distribution: 15 species in Africa, SE Asia, and adjacent islands have been assigned to this genus, some erro- neously. Key: Wood (1960a: 18) for Microne- sia. Cosmoderes Eichhoff [1878a: 495, Type- species: Cosmoderes inonilicollis Eichhoff, monobasic. Synonyms: Erioschidias Schedl 1938b: 42, Type-species: Cryphalus setistria- tus Lea, subsequent designation by Wood 1960a: 21; Dendriops Schedl 1953b: 125, Type-species: Dendriops granulicollis Schedl, monobasic; Vitaderes Beeson 1941: 301, nomen nudum. Type-species: Vitaderes luffa , nomen nudum =Cosinoderes inonili- collis Eichhoff, no status]. Distribution: About 22 species in Africa to SE Asia and Australia. At least 2 species were taken from lianas. Crijptocarenus Eggers [1937: 79, Type- species: Cryptocarcnus diadematus Eggers, original designation. Synonym: Tachyderes Blackman 1943a: 35, Type-species: Tachy- deres floridensis Blackman =Cryptocarenus seriattis Eggers, original designation]. Distri- bution: 5 species in North and Central Amer- ica, more than 7 in South America, 1 intro- duced into tropical Africa. All are con- sanguineously polygynous and primarily myelophagous. Key: Wood (1982: 912). Hypothenemus Westwood [1836: 34, Type- species: Hypothenemus eruditus Westwood, monobasic. Synonyms: Stephanoderes Eich- hoff 1872: 132, Type-species: Stephanoderes chapuisii Eichhoff =Crypturgus dissimilis Zimmermann, subsequent designation by Hopkins 1914: 130; Homoeocryphalus Linde- mann 1876: 168, Type-species: Stephan- oderes ehlersii Eichhofi = Hypothenemus eru- ditus Westwood, monobasic; Adiaeretus Hagedorn 1909: 744, Type-species; Adiaere- tus spinosus Hagedorn = Stephanoderes ela- phus Eichhoff, monobasic; Stylotentus Schedl 1939b: 380, Type-species: Hypothenemus concolor Hagedorn, subsequent designation by Schedl 1961: 4, 48; probable synonym, Triarmocerus Eichhoff 1878a: 42, 119, Type- species: Triarmocerus cryphaloides Eichhoff, monobasic, type lost; Chondronoderes Schedl 1940b: 589, Type-species: Stephan- oderes magnus Eggers, monobasic; Archeo- 92 Great Basin Naturalist Memoirs No. 10 Fig. 54. Corthtjius spp. (Corthylini): A-B, subserratus Wood; C-E, scrratiis Wood; F-G, concisus Wood; H-I, procerus Bright. phaliis Schedl 1941: 392, Type-species: Archeophalus nataleiisis Schedl, monobasic; Pachijnodcres Schedl, 1941: 393, Type-spe- cies: Pachynoderes deprecator Schedl, monobasic; Lepiceroides Schedl 1957a; 59, Type-species; Lepiceroides aterrinius Schedl, monobasic; Ernophloeus Nunberg 1958; 484, Type-species; Ernophloeus costalimai Nun- berg = Stephanoderes sundaensis Eggers, original designation; Epsips Beeson 1941: 287, nomen nudum. Type-species; Epsips stjl- varum, nomen nudum, no status]. Distribu- tion: 39 species in North and Central America; several hundred nominate species from other tropical and subtropical areas have been as- signed to this genus, but a majority of them are now in synonymy. All are consan- guineously polygynous and myelophagous, 1986 WOOD: Genera of Scolytidae 93 phloeophagous, or spermophagous. Kevs: Wood (1954: 1017, 1052, 1982; 876) for North and Central America. Trischidias Hopkins [1915a: 7, 12, Type- species: Trischidias georgiae Hopkins, origi- nal designation]. Distribution: 4 species in SE North America to Me.xico, 1 imported to Hawaii. All are consanguineously polygynous and phloeophagous. Kevs: Wood (1954b: 1067, 1982: 872). Periocryphalus Wood [1971: 33, Type-spe- cies: Periocnjphahis pulhis Wood, original designation]. Distribution: 2 species in South America. Both are consanguineously polygy- nous and myelophagous in minute lianas. Tribe Corthylini Corthyli LeConte [1876: 346, 347, Type-genus: Cort/iy- lus Erichson, 1836] Pityophthoridae Eichhoff [1878a; 173, Type-genus; Pitijophthorus Eichhoff, 1864] Araptidae Eichhoff [1878a: 30.5, Tvpe-genus; Araptus Eichhoff, 1872] Amphicranidae Eichhoff] 1878a: 460, Type-genus; Amp/r- icranus Erichson, 1836] Description. — Frons usually dimorphic, either or both sexes feebly to strongly modi- fied in sculpture and ornamentation; eye emarginate; scape usually elongate, strongly flattened in some Coi'thijlus (Fig. 54), etc., funicle 1- to 5-segmented, club strongly flat- tened, sutures present or obsolete, frequently 1 and/or 2 septate; when present, sutures on posterior face little if any displaced toward apex; anterior slope of pronotum asperate (one exception in female Corthylus cecropii), anterior margin frequently armed, lateral and basal margins fi-equently marked by a fine, raised line; procoxae contiguous, tibiae rather slender, their lateral margins rarely armed by more than four socketed teeth; metepisternal spine obsolete, replaced by small, transverse groove (Fig. 32), thus making it possible for elytra in locked position to cover at least pos- terior two-thirds of metepisternum (Figs. 53- 54); vestiture usually hairlike, scales rarely present. Biology. — Monogamy and heterosan- guineous polygyny are common throughout the tribe; a few species of Araptus practice consanguineous polygyny. Phloeophagy pre- dominates in temporate areas, xylomyce- tophagy in tropical areas, although myelo- phagy and spermophagy are common. Parental galleries may be monoramous, bi- ramous, or variously multiramous. The eggs are deposited in niches or, in Corthylus , in fully formed larval cradles. The larvae form individual mines or cradles depending on the food habit. The domicile parasitic habit is known in Corthylocurus, Tricolus, amd Am- phicrantis . Taxonomy. — Except for two small endemic genera in Madagascar {Pityodendron, Sauroptilius) and one in Africa (Mhniocurus), a few primitive Pityophthoriis species in Africa, and several modern Pityophthoriis in Eurasia, obviously derived from North Amer- ica, this large, unique tribe is entirely Ameri- can. The occurrence of primitive elements of ancient origin in Africa and Madagascar sug- gests that basic tribal characters formed prior to or early in the Tertiary. The elytral locking mechanism and antennal club are unique in the family, with the Cryphalini possibly form- ing an intermediate step between Corthylini and the primitive tribes of Scolytinae. On a biological basis, the tribe is readily divisible into the phloeophagous Pityoph- thorina and the xylomycetophagous Cor- thylina, although anatomical characters to support that division are less definite. The tribe obviously originated in South America and was affected by two radiations. The first occurred prior to the Tertiary and carried a few species into Madagascar and Africa, while those land masses were either connected or close enough for island hopping, and took sev- eral species over the land bridge to North America, where a minor secondary radiation occurred. The second major radiation oc- curred in South America during the Tertiary and produced the Corthylina. Gnathotrichus might have reached Central America or southern Mexico over the pre-Tertiary land bridge just prior to or during the early stages of its closure. A few genera and species have reached North America over the present post- Tertiary land bridge or by island hopping in recent time. The occurrence oi Gnatharus, a member of the Corthylina, in China is more difficult to explain. The South American element of this tribe is one of the more poorly known segments of the Scolytidae. Much remains to be learned about this remarkable group. Chiloxylon Schedl (see Dryocoetini) could belong to this tribe. 94 Great Basin Naturalist Memoirs No. 10 Kev to the Genera of Corthvlini (Modified from Wood 1982) - 1. Phloeophagous, myelophagous, or sper- mophagous; antennal funicle 5-segmented (except 3- or 4-segineiited in Dcndroterus, Daciwpbthorus , and some Pitijophthorus costatiis), club usually smaller, symmetrical; prosternal intercoxal piece acutely pointed (except obsolete in Dacnophthorus); pubes- cence usually more abundant, usually in rows on elytra; elytral declivity mostly convex to bisulcate, armature conser\'ative (subtribe Pityophthorina) 2 — Xylomycetophagous; antennal funicle 1- to 5- segmented, club usually much larger, com- monly asymmetrical; prosternal intercoxal piece absent (except obtuse in Gnatho- trichus, Gnathotrupes); pubescence usually greatly reduced to obsolete or minute and strongly confused (not in rows); elytral decliv- ity convex to truncate to deeply excavated (weakly bisulcate in some Gnathotrichus), commonly with spinelike processes (subtribe Corthylina) 17 2(1). Basal and lateral margins of pronotum rounded, devoid of a fine, raised line; elytra rather coarsely punctured (American genera), or veiy finely punctured (African genus), un- armed declivity steep, in American genera usually subvertical and somewhat flattened on lower half, almost never bisulcate; discal vestiture abundant 3 — Basal and usually lateral margins of pronotum marked by a finely raised line; elytral de- clivilty usually more gradual, convex to bisul- cate, often ornamented by granules or small denticles 6 3(2). Antennal club pubescent to base, unmarked by sutures, funicle .5-segmented; female frons ornamented by a brush of long hair, male usually with a median tubercle or longitudinal carina at upper level of eyes; elytral declivity conservatively sculptured, usually convex; Africa to India and China; 1.4-2.8 mm Mimiocurus — Antennal club with sutures clearly indicated; American genera 4 4(.3). Antennal funicle 3- or 4-segmented, club with sutures at least slightly recurved, aseptate; elytral vestiture hairlike; female epistoma not emarginate, mandible never with projecting spines; strong, transverse frontal carina at up- per level of eyes, when present, a female character; phloeophagous; SW USA to Panama; Bursera, 1 species in Jatropha; 1.3- 2.9 mm Dendroterus — Antennal funicle .5-segmented; secondary sexual characters different 5 5(4). Antennal club with aseptate sutures strongly procurved; interstrial setae scalelike; male frons strongly, transversely carinate at upper level of eyes, female epistoma deeply emar- ginate to accommodate a pair of mandibular spines; phloeophagous; Costa Rica to Bolivia and Brazil; Asfroniiim, Spotidias; 1 3-1. 6mm Styphlosonuj — Sutures of antennal club straight or recurved; female epistoma entire, male frons without a transverse carina; mandibular spines never present; elytral declivity convex, interstriae unarmed by tubercles or spines; female frons shallowly concave, bearing a brush of hair, armed on median fourth above eyes by an acute, transverse carina and by a pair of coarse tubercles in lateral areas at level of antennal insertion; male frons convex, simple; Mexico; Bursera; 1.6-1.8 mm Phloeotertts 6(2). Sutures of antennal club moderately to very strongly procurved, onh- suture 1 septate, or if all external sutures obsolete then mesal half of suture 1 marked internally by a strongly procurved septum at least on 1 side; phloeophagous, myelophagous, spermopha- gous; S USA to Argentina; 1.1-3.3 mm Araptus — Sutures 1 and 2 clearly, equally marked by rows of setae and grooves, straight to moder- ately procurved, if procurved then both su- tures at least partly septate (sutures always straight when both almost obsolete) 7 7(6). Lateral margins ol pronotum rounded, with- out a fine, raised line 8 — Lateral margins of pronotum subacute, marked by a fine, raised line (rather obscure in Spennophthorus and some Pitijophthorus having pronotal asperieies in subconcentric rows 12 8(7). Sutures 1 and 2 of antennal club aseptate and clearly marked by grooves and rows of setae; pronotal asperities continuing in lateral areas to base; larger species; spermophagous in cones of Piniis. at least 2 species sometimes myelophagous; North America; 2.2-4.1 mm Conophthorus — Sutures 1 and 2 both partly to completely septate or, if aseptate, then antennal club largely glabrous or body size much smaller; pronotal asperities not extending to basal margin; smaller species, not found in cones of conifers 9 9(8). Antennal club rather large, at least 2.5 times as long as funicle; monogamous species .... 10 — Antennal club comparatively small, less than 1.5 times as long as funicle; polygynous spe- cies 11 10(9). Body stouter, 2.5-2.8 times as long as wide; elytral declivity convex; anterolateral areas of female pronotum with a pair of large, oval, densely pilose areas; antennal club with only 2 sutures; phloeophagous, larval mines resem- ble cradles (of ambrosia beetles); S USA to Honduras; Pinus; 1.5-3.2 mm Pityoborus — Body very slender, 3.7-3.8 times as long as wide; elytral declivity strongly impressed; fe- 1986 WOOD: Genera of Scolvtidae 95 male pronotum without special pilose areas; antennal club with suture 3 indicated by a row of setae; phloeophagous, but with fungal mycelium in parental gallery; Mexico to South America; Clematis and Bignoniaceae vines; 1.2-2.0 mm Dacnophthorus 11(9). Female pregula very greatly enlarged and bearing a rather dense, conspicuous tuft of very long hair, male pregula only slightly en- larged, usually without tuft of hair; phloeo- phagous; USA (Arizona, New Mexico), Pintix, 1.3-1.8 mm Pityotrichus — Female oral region abnormally broad and, usually, with mandibles greatly enlarged; pregula normal, phloeophagous. Central and South America; Dacreoidcs. Cedrcla. Prot- ium; 1.3-1.8 mm Gnatholeptus 12(7). Pronotum weakly declivous on anterior fourth, asperities small, numerous, gradually decreasing in size toward base; head unusu- ally wide, mandibles large and stout in both sexes; eye large, coarsely faceted, one-third divided by a broad emargination; Madagas- car; 3.2-3.4 mm Vitxjodendron — Pronotum more strongly declivous in front, asperities usually not present behind summit; head and mandibles normal 13 13(12). Pronotum without a transverse impression behind summit, transition between asperate and smooth areas more gradual, asperities al- ways confused; interstrial bristles usually stout to scalelilce (1 exception, also, almost glabrous in Sauropfi/ius); tropical species .. 14 — Pronotum almost always with a distinct, trans- verse impression behind summit, if doubtful then asperities almost always arranged in con- centric rows, interstrial setae hairlike (il stout then pronotal asperities concentric) 16 14(1.3). Anterior margin of pronotum unarmed; de- clivity very broadly excavated (as wide as body), its lateral crests profoundly elevated and serrate, but abruptly ending before su- tural apex; Madagascar; 3.2 mm . . . Sauroptilius — Anterior margin of pronotum armed by serra- tions or a continuous costa; declivity convex to moderately sulcate, impression rarely equal to more than half width of body; neotropical 15 15(14). Strial punctures rather coarse, mostly in rows, declivity moderately to rather strongly impressed, lateral margins armed or not; frons never armed, epistomal margin with a small, median, premandibular lobe, polygy- nous; phloeophagous, axes of tabular nuptial chamber perjjendicular to cambium, longest axis parallel to grain of wood; Colombia, Venezuela; broadleaf trees, 1.3-3.1 mm .... Phelloterus — Strial punctures either very small or con- fused; declivity not as steep, convex to shal- lowly impressed, lateral margins never armed; male frons armed; spermophagous; Central and South America; Cacsalpinus , etc.; 1.3-1.9 mm Spermophthorus 16(13). Sutures of antennal club moderately pro- curved, segment 1 shorter than 2 or 3; greater frontal pubescence a male character, clytral punctures very fine, usually confused, short pubescence abundant, often .scalelike; striae usually obsolete; phloeophagous, monoga- mous; North America to Colombia, China; Quercus, rarely other hosts; 1.2-2.6 mm . . . Pseudopityophthorus — Sutures of antennal club straight to modestly procurved, segments 1 and 2 subequal in length; pubescence usually much less abun- dant, never scalelike, strial punctures in rows or, if confused, then rather coarse; phloeo- phagous, monogamous or heterosanguine- ously polygynous; America, Europe, Asia, Africa; coniferous and broadleal hosts; 0.9-3.9 mm Pityophthorus 17(1). Antennal funicle 5-segmented, club always symmetrical, with 2 or 3 clearly marked su- tures; protibia widest near its apex, its poste- rior face usually flat, unarmed (a few minute granules in some Gnatbotrupes), elytral de- clivity conservatively sculptured (except in some Gnathotnipes) 18 — Antennal funicle 1- to 4-segmented, club commonly asymmetrical, often greatly en- larged, sutures often reduced or absent; tibiae variously sculptured 19 18(17). Sutures of antennal club straight to moder- ately procurved, segment 1 not noticeably reduced in size; elytral declivity convex to narrowly, rather shallowly sulcate, subapical margin near apex acutely elevated, sutnral apex entire, rather narrowly rounded behind; monogamous; North and Central America; coniferous and broadleaf hosts; 2.0-4.3 mm Gnathotrichus — Sutures of antennal club moderately to strongly procurved, segment 1 distinctly smaller; elytral declivity moderately to strongly flattened; elytral apex at least weakly divaricate, very broadly romided to shallowly emarginate behind, without a submarginal costa near apex; monogamous; Central and South America; broadleaf hosts; 1.. 3-4.0 mm Gnathotrupes 19(17). Antennal funicle 2- to 4-segmented, club with 2 sutures clearly marked; elytral apex divari- cate (except Mctacorthylus. Glochinoccrus) . commonly explanate, declivity often elabo- rately excavated and armed by spines; pro- tibia always slender, with posterior face in- flated and tuberculate; body usually slender 20 — Antennal funicle 1-segmented, club with 1, 2, or no sutures; elytral apex entire (except Brachyspartus, Corthylua cmarfiinatus), de- clivity convex to rather weakly excavated, never explanate; posterior margin ol protho- 96 Great Basin Naturalist Memoirs No. 10 racic precoxal piece transversely straight, not extended between coxae; protibia variable; body conparatively stout 24 20(19). Procoxae contiguous, anterior wall of com- bined cavities and precoxal piece transversely straight (longitudinally very thin); protibia similar in male and female, with a longitudinal marginal row of tubercles, posterior face ei- ther unarmed or with a longitudinal row of up to about 4 tubercles, posterior face of female antennal club with long hair sparse to absent; antennal funicle usually 3-segmented, less commonly 2-segmented; frons commonly with a sharply defined granular area 21 — Prothoracic precoxal piece moderately large, posteriorly angulate, occupying anterior por- tion of area between coxae (except in Gnatharus, from Tibet, a thin, transverse partition); male protibia armed by coarse mar- ginal serrations and a row of equally coarse serrations on posterior fiice, female protibia with posterior face moderately inflated and armed by numerous, confused, small tuber- cles in addition to coarser marginal row; pos- terior face of female antennal club orna- mented by more abundant, long hair; frons never ornamented by a sharply defined gran- ulate area 22 21(20). Elytra broadly rounded behind, posterior margin of declivity feebly if at all explanate, weakly if at all divaricate; lateral margins of declivity armed by as many as 3 pairs of spines; antennal club oval to subtriangular, little if any longer than wide; anteroventral margin of prosternum flanged, bent or folded caudad away from head, usually bearing a tuft of hair; pronotum usually stouter, anterior margin usually serrate; antennal funicle 3- segmented; monogamous; Mexico to South America; 1.5-3.9 mm Tricolus — Posterior margin of elytral declivity strongly to profoundly explanate, weakly to pro- foundly divaricate; declivital armature vari- able, often remarkable; antennal club oval to very elongate; anteroventral margin of prosternum fitting snugly against head (not bent caudad), sparsely pubescent; pronotum usually much more elongate, anterior margin variously sculptured, rarely serrate; antennal funicle usually 3-segmented, occasionally 2- segmented; monogamous; Mexico to .South America; 2. .5-8.0 mm Amphicramis 22(20). Antennal funicle 4-segmented; frons with me- dian carina; pronotum dimorphic, female with anterior slope strongly declivous, armed by numerous asperities, anterolateral angles each bearing a tuft of hair, male slope gradual, asperities greatly reduced in size and num- ber, anterior margin a strongly formed, slightly produced continuous costa; elytral declivity divaricate from middle, truncated before apex; Asia (Tibet); 2.0 mm .. Gnatharus — Antennal funicle 2- or 3-segmented; prono- tum and elytra cjuite different; American gen- era 23 23(22). Elytral apex divaricate, often also explanate; antennal club oval to broadly triangular; lat- eral margins of pronotum usually with a fine, raised line (a few exceptions); body moder- ately to very slender; mostly polygynous; North and South America; 1.4-4.8 mm Monarthrum — Elytral apex entire, never explanate; lateral margins of pronotum rounded; antennal club more than twice as long as wide (except 1.6 times in male Metacorthijlus), its apex nar- rowly rounded; body comparatively stout ... 24 24(23). Antennal funicle 3-segmented; female frons excavated and elaborately ornamented by long hair; antennal club not sexually dimor- phic, elongate, slightly asymmetrical; prono- tum and elytral disc glabrous; monogamous; S Mexico to Guatemala; 3.0-3.8 mm Glochinocerus — Antennal funicle 2-segmented; frons convex and subglabrous in both sexes; antennal club asymmetrically very elongate in female, elon- gate-oval in male; pronotum and elytra minutely, closely pubescent; monogamous; Costa Rica to Colombia; 1.9-2.7 mm Metacorthylus 25(19). Lateral margins of pronotum rounded; elytral disc usually impunctate, declivity short, very steep, narrowly sulcate on basal third, trian- gularly impressed below, costal margins near apex ascending slightly; antennal club subcir- cular, syummetrical, with 2 aseptate sutures marked by rows of setae; female frons convex, pubescence inconspicuous; monogamous; Mexico to South America; 1.2-2.4 mm Microcorthylus — Lateral margins of pronotum marked by a fine, raised line (except some Corthycyclon), elytral disc usually with clearly marked, con- fused punctures, declivity convex, truncate, or variously sculptured (but not as above); female frons usually moderately to strongly concave, often ornamented by hair; antennal club symmetrical to strongly asymmetrical, sutures present or not 26 26(25). Antennal club aseptate, without sutures (some species with weak, transverse grooves, without rows of setae), usually very elongate; lateral margins of pronotum either with or without a fine, raised line; posterior face of protibia inflated and tuberculate; female frons broadly, evenly concave and ornamented by fine hair; monogamous; Mexico to South America; 1.3-2.4 mm Corthycyclon — Antennal club with 1 or 2 sutures, its outline subcircular to strongly asymmetrical (if su- tures absent then posterior face of protibia flat, smooth); female frons variable 27 1986 WOOD: Genera of Scolytidae 97 27(26). Elytral apex strongly, obtusely divaricate, fe- male antennal club with 1 septate suture, ta- pered on apical half to strongly acuminate apex, posterior face of club without a tuft of long hair; female protibia inflated, armed on posterior face; Venezuela; 2.5 mm Brachyspartus — Elytral apex entire (a small cleft in Corthijlus emarginatus). female antennal club not ta- pered or acuminate at apex 27 28(27). Elytral declivity narrowly, weakly sulcate (ex- cept uniformly convex in 1 species), lateral margins armed by 2 or 3 pairs of pointed granules; antennal scape elongate, club shaped; female frons variously impressed, with a pair of median carinae narrowly sepa- rated by a sulcus over part or all of median line; color pale yellow to yellowish brown; antennal club symmetical, broadly oval, with 2 finely marked sutures; female protibia in- flated, posterior face tuberculate; monoga- mous; Mexico to South America; 1.5-2.8 mm Corthylocurus — Sculpture of elytral declivity convex, trun- cately concave, or variously impressed (but never narrowly sulcate); antennal scape sub- quadrate, stout; female frons never with pair of median carinae; antennal club slightly to profoundly asymmetrical, sutures (when present) rather strongly marked; posterior face of protibia smooth or tuberculate; monogamous; North and South America; 1.2- 4.3 mm Corthijlus Corthylini; Pityophthorina Mimiocurus Schedl [1957a: 72, Type-spe- cies: Mimiocurus acuminatus Schedl, mono- basic. Synonyms: Micracidendron Schedl 1957a: 71, Type-species: Micracidendron montanum Schedl, monobasic; Mimiophtho- rus Schedl 1957a: 77, Type-species: Bra- chydendrulits montanu.s Schedl, original des- ignation]. Distribution: About 8 species in Africa, 1 in India, 1 in China. Styphlosoma Blandford [1904: 2.32, Type- species: Styphlo.'ioina uranulatum Blandford, monobasic]. Distribution: 1 species in Central America (Costa Rica to Panama), 3 in South America. They are heterosanguineously poly- gynous and phloeophagous. Dendroterus Blandford [1904: 233, Type- species: Dendroterus mexicanus Blandford, subsequent designation by Hopkins 1914: 120. Synonyms: Plesiophthorus Schedl 1940a: 343, Type-species: Plesiophthorus perspectus Schedl, monobasic; Xylochilus Schedl 1956: 30, Type-species: Xylochilus insularis Schedl =Dendroterus sallaei Blandford, original designation]. Distribution: 15 species in North and Central America. All are het- erosanguineously polygynous and phloeo- phagous. Key: Wood (1982: 918). Phloeoterus Wood [1984: 117, Type-spe- cies: Phloeoterus burserae Wood, original designation]. Distribution: 1 species in Burs- era in Mexico. Arapfus Eichhoff [1872: 136, Type-species: Araptus nifopalliatus Eichhoff, monobasic. Synonyms: Neodi-yocoetes Eggers 1933a: 9, Type-species: Neodryocoetes hymenaeae Eg- gers, monobasic; Thamnophthorus Schedl 1938c: 174, Type-species: Thamnophthorus volastos Schedl, subsequent designation by Blackinan 1942b: 178; Neopityophthorus Schedl 1938c: 180, Type-species: Pityophtho- rus laevigatus Eggers, designated by Wood 1982: 928; Sphenoceros Schedl 1939e: 565, Type-species: Sphenoceros Umax Schedl, monobasic; Hypertensus Hagedorn, nomen nudum (in Schedl 1950d: 164), Type-species; Hypertensus reittcri, nomen nudum = Sphe- noceros Umax Schedl, no status; Brachyden- dridus Schedl 1951b: 114, Type-species: Brachydendrulus eggersi Schedl, monobasic; Gnathocranus Schedl 1951b: 116, Type-spe- cies: Gnathocranus novateutonicus Schedl, monobasic; Gnathohorus Schedl 1970a: 93, Type-species: Breviophthorus argentiniae Schedl, original designation]. Distribution: .53 species in North and Central America, about 50 in South America and adjacent is- lands. Some are monogamous, some are het- erosanguineously polygynous, and at least 3 are consanguineously polygynous {laevigatus, costaricensis , etc.); most are phloeophagous, a few are spermophagous, and some are myelophagous in vines (lianas). Key: Wood 1982: 928) for North and Central America. Conophthorus Hopkins [1915c: 430, Type- species: Pityophthorus coniperda Schwarz, original designation]. Distribution: 14 species in North America. All species are monoga- mous and spermophagous in cones oi Pinus . Key: Wood (1982: 982). Pityoborus Blackman [1922: 96, Type-spe- cies: Crypturgus comatus Zimmermann, monobasic]. Distribution: 7 species in North and Central America. All are monogamous and phloeophagous in Pinus. Keys: Wood (19.58: 47, 1982: 1147). 98 Great Basin Naturalist Memoirs No. 10 Dacnophthoriis Wood [1975; 394, Type- species: Gnathophthorus clematis Wood, original designation]. Distribution: 2 species in North and Central America, 2 in South America. All are monogamous and bore in the stems of vines (lianas) (Clematis and a species of Bignoniaceae). Key: Wood (1982: 1152) for North and Central America. Pityotrichus Wood [1962: 76, Type-spe- cies: Pitijophilus harbatus Blackman, auto- matic. Synonym: Pitijophilus Blackman 1928a: 147, Type-species: Pitijophilus harba- tus Blackman, original designation, preoccu- pied]. Distribution: 2 species in North Amer- ica (Arizona, New Mexico). Both are hetero- sauguineously polygvnous and phloeo- phagous. Key: Wood (1982: 1141). Gnatholeptus Blackman [1943a: 34, Type- species: Gmitholcptus mandihularis Black- man = Pitiiophthorus shannoni Blackman, original designation]. Distribution: 4 species in Central and N South America. All are het- erosanguineously polygynous and phloeo- phagous, mostly in Protium. The status of this genus should be reviewed. Key: Wood (1982: 1142). Pityodendron Schedl [19.53a: 93, Type-spe- cies: Pityodendron madagascarensis Schedl, monobasic]. Distribution: 1 species in Mada- gascar. Sauroptilius Browne [1970: 558, Type-spe- cies: Xyleborus sauropterus Schedl, original designation]. Distribution: 1 species in Mada- gascar. Phelloterus Wood [1971: 46, Type-species: Phelloterits tersus Wood, original designa- tion]. Distribution: 3 species in South Amer- ica (Colombia to Venezuela). All are het- erosanguineously polygynous' and phloeo- phagous. Spermophthorus Costa Lima [1929: 111, Type-species: Spermophthorus apuleiae Costa Lima, monobasic]. Distribution: 1 species in Central America (Costa Rica), 2 in South America. Pseudopityophthorus Swaine [1918: 93, Type-species: Crypturgus minutissimus Zim- mermann, original designation]. Distribu- tion: 23 species in North and Central Amer- ica, 1 in South America (Colombia), 1 in China. All are bigynous and phloeophagous, mostly in Quercus. Keys: Blackman (1931a: 225), Wood (1982: 966). Pittjophthorus Eichhoff [1864: 39, Type- species: Bostrichus lichtcnsteini Ratzeburg, subsequent designation by Hopkins 1914: 127. Synonyms: Trigonogenius Hagedorn 1912: .354, Type-species: Trigonogenius fallax Hagedorn, monobasic; Hagedornus Lucas 1920: 683, Type-species: Trigonogenius fallax Hagedorn, automatic; Mijeloborus Blackman 1928a: 16, Type-species: Pitijophthorus ramiperda Swaine, original designation; Gnathophorus Schedl 1935: 342, Type-spe- cies: Gnathophorus sparsepilosus Schedl, monobasic, preoccupied; Conophthocran- idus Schedl 1935; .343, Type-species: Conoph- thocranulus blackmani Schedl, monobasic; Breviophthorus Schedl 1938c: 176, Type-spe- cies: Breviophfhorus brasiliensis Schedl, monobasic; Pityophthoroides Blackman 1942b: 199, Type-species: Pityophthoroides pudens Blackman, original designation; Cladohorus Sawamoto 1942: 165, Type- species: Cladoboriis arakii Sawamoto, monobasic; Neomips Schedl 1954a: 37, Type- species: Neomips brasiliensis Schedl = Pityophthonis dimorphus Schedl, monoba- sic; Ctenyophthorus Schedl 1955b: 26, Type- species: Ctenyophthorus glabratulus Schedl, mono-basic; Gnathophthorus Wood 1962: 76, Type-species: Gnathophorus sparsepilosus Schedl, automatic; Hypopityophthorus Bright 1981a: 14, Type-species: Pitijophtho- rus inops Wood, original designation]. Distri- bution: 225 species in North and Central America, about 100 in South America and adjacent islands, about 15 in Asia, 22 in Eu- rope, 22 in Africa. Most are heterosan- guineously polygynous, a few are monoga- mous; most are phloeophagous, a few are mvelophagous. Kevs: Pfeffer (1976: .334) for Europe, Blackman' (1928a), Bright (1981a), and Wood (1982: 991) for North and Central America. Corthylini: Corthylina Gnathotrichus Eichhoff [1869: 275, Type- species: Gnathotrichus corthyloides Eichhoff = Tomiciis materiarius Fitch, monobasic. Synonyms: Gnathotrichoides Blackman 1931b: 267, Type-species: Cryphalus sulcatus LeConte, subsequent designation by Wood 1982: 1 1.55; Anci/Zof/ere.s Blackman 1938b:205, Type-species: Cryphalus pilosus LeConte, original designation; Paraxyleborus Hoff- 1986 WOOD; Genera of Scolytidae 99 inann 1942: 72, Type-species; Xyleborus duprezi Hoffmann =Tomicits mutcriarhis Fitch, monobasic; Prognathotricliiis Bright 1972: 1678, Type-species: Prognathotrichiis priimis Bright, original designation]. Distri- bution; 13 species in North and Central Amer- ica. All are monogamous and .xylomyce- tophagous. Kevs: Blackman (1931b: 266), Wood (1982; 1156). Gnathotrupes Schedl [1951b: 125, Type- species: Gnathotnipes bolivianiis Schedl, monobasic. Synonyms: Gnathotnjpaiuis Wood 1968a: 9, Type-species; Gnathotrypanus terc- hrattis Wood, original designation; Gnatho- cortiis Schedl 1975a; 11, Type-species: Gnathocorius caliculus Schedl, original des- ignation; Gnathomhnus Schedl 1975a; 12, Type-species; Giuithomimiis nothofagi Schedl, original designation; Gnathoglochi- nus Schedl 1975a: 16, Type-species: Gnatho- glochinus impressus Schedl, original designa- tion]. Distribution; 5 species in Central America, about 30 in South America. All are monogamous and xylomycetophagous. Key: Wood(1982; 1167) for Central America. Tricolus Blandford [1905; 286, Type-spe- cies; Tricoltis ovicollis Blandford, subsequent designation by Hopkins 1914; 131. Synonym: Pterocyclonoides Schedl 1970a: 101, Type- species: Pterocyclonoidcs octodentatits Schedl, monobasic]. Distribution: 24 species in North and Central America, about 17 in South America. All are monogamous and xy- lomycetophagous. Key: Wood (1982: 1170) for North and Central America. Amphicranus Erichson [1836; 63, Type- species; Arnphicranus thoracicus Erichson, monobasic. Synonyms: Piezorhopahis Guerin- Meneville 1838: 107, Type-species; Piezorho- palus nitidulus Guerin-Meneville =Amphi- craniis thoracicus Erichson, monobasic; Steganocranits Eichhoff 1878a; 460, Type- species; Steganocranus dohnii Eichhoff, monobasic]. Distribution: 29 species in North and Central America, about 23 in South America. All are monogamous and xylomyce- tophagous; apparently some appropriate the tunnel of another species of Scolytidae or Platypodidae by driving out or killing the orig- inal occupants (domicile parasitism). Key: Wood (1982; 1186) for North and Central America. Gnatharus Wood & Yin [1986; 463, Type- species; Gnathariis tibctensis Wood & Yin, original designation]. Distribution: 1 species in Asia (Tibet). Monarthruin Kirsch [1866: 213, Type-spe- cies: Monarthrum chapiiisi Kirsch, monoba- sic. Synonyms; Coi-thylomimus Ferrari 1867: 48, Type-species; Bostrichus fasciatus Say, subsecjuent designation by Hopkins 1914: 118; Cosmocoryniis Ferrari 1867: 62, Type- species; Cosmocorynus cristatus Ferrari, monobasic; Pterocyclon Eichhoff 1869; 276, Type-species; Pterocyclon late rale Eichhoff, subsequent designation by Hopkins 1914; 128, neotype for type-species designated by Wood 1966; 25; Anchonocerus Eichhoff 1878a; 67, 431, Type-species; Anchonocerus ntfipes Eichhoff, monobasic; Phthohus Eich- hoff 1878a; 67, 433, Type-species; Phthorius ingens Eichhoff, monobasic; Trypocranus Eichhoff 1878a: 67, 4.35, Type-species; Trypocranus cincinnatus Eichhoff, monoba- sic; Eupteroxylon Eggers 1936a: 392, Type- species: Type-species; Eupteroxylon coma- tum Eggers, monobasic]. Distribution: 58 species in North and Central America, about 78 in South America and adjacent islands. Most species are heterosanguineously polygy- nous, a few small species are monogamous; all are xylomycetophagous. Key: Wood (1982: 1207) for North and Central America. Glochinoceriis Blandford [1904: 266, Type- species: Glochinoccrus retusipennis Bland- ford, subsequent designation by Hopkins 1914; 122]. Distribution: 2 species in Mexico (Hidalgo) and Central America (Guatemala). Both are monogamous and xylomyce- tophagous. Key; Wood (1982; 1246). Metacorthijlus Blandford [1904: 251, 263, Type-species: Metacorthyhis nigripennis Blandford, monobasic. Synonym: Paracor- thyhis Wood 196Sa; 7, Type-species; Paracor- thyhis velutinus Wood, original designation]. Distribution; 4 species in Central America 1 of which also occurs in South America (Colom- bia). All are apparently monogamous and xy- lomycetophagous. Key: Wood (1982: 1248). Microcorthylus Ferrari [1867; 58, Type- species; Microcoiihylus parvulus Ferrari, monobasic]. Distribution: 13 species in Mex- ico and Central America, 19 in South Amer- ica. All are monogamous and xylomyce- 100 Great Basin Naturalist Memoirs No. 10 tophagous. Key: Wood (1982; 1252) for North and Central America. Corthycyclon Schedl [1951b: 128, Type- species; Corthycyclon ustum Schedl, mono- basic]. Distribution: 6 species in Mexico and Central America, about 10 in South America. All are monogamous and xylomycetophagous. Key; Wood (1982: 1260) for North and Central America. Brachyspartus Ferrari [1867: 65, Type- species: Brachyspartus moritzi Ferrari, monobasic. Synonym; Thyhircos Schedl 1939e: 567, Type-species; Brachyspartus moritzi Ferrari, subsequent monotypy (Wood 1982: 1298). Distribution: 1 species in South America (Venezuela). Corthylocurus Wood [1966: 18, Type-spe- cies; Brachyspartus barbatus Blandford, original designation]. Distribution: 6 species in Mexico and Central America, about 8 in South America. All are monogamous and xy- lomycetophagous. Key; Wood (1982; 1265) for Mexico and Central America. Corthylus Erichson [1836; 64, Type-spe- cies; Bostrichus compressicornis Fabricius, subsequent monotypy (Ferrari 1867; 49), lec- totype designated for type-species by Wood 1974; 202. Synonyms; Morizus Ferrari, 1867; 69, Type-species: Morizus excisus Ferrari, monobasic; Pseudocorthyhis Ferrari 1867: 59, Type-species: Pseudocorthyhis letzneri Fer- rari, subsequent designation by Hopkins 1914; 128]. Distribution: 56 species in North and Central America, about 54 species in South America. All are monogamous and xy- lomycetophagous. Key: Wood (1982: 1271) for North and Central America. Literature Cited Arnoldi, et al, 1977. (Title not seen). Trudy Paleont. Inst. 166: 1-203. (Partial English translation re- ceived through R. T. Thompson). Balachowsky. a S 1947. A propos du genre Pitijonencfi Bedel (Col. Scolytoidea). Societe Entomologique de France, Bulletin 52(.3): 44. 1949. Coleopteres: Scolytidae. Faune de France 50. 320 p. Bedel, L, E. M 1888. La Famille Scolytidae. 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Morphologische Studien uber borkenkafer. I. Die Gattungen Ipa DeCeer und Pityogenen Bedel. Habschr. techn. Hochschule Karlsruhe. Mvinchen (C. Void und S., Munchen). 45 p., .39 figs. 1912. Morphologische Studien uber Borkenkafer. n. Die europaischen Hylesinen. E. Reinhardt, Miinchen. 53 p. , 3 pis. , 82 fig.s. 1913. Footnote No. I. Page 43 in E. Reitter, Bes- timmungs-Tabelle der Borkenkiifer (Scolytidae) aus Europa und den angrenzenden Liindern. Wiener Entomologische Zeitung .32(Beilieft): 1-116. Gemminger, M . AND B. VON HAROLD 1872. Catalogus Goleopterorum, huscusque descriptorum syn- onymicus et systematicus. Scolytidae, Brenthi- dae, Anthribidae, Cerambycidae. Monachii, Munchen 9: 2669-2988. Geoffroy. E. L 1762. Histoire abregee des insectes qui se trouvent aux environs de Paris dans laliaerotrypes, Parasphaero- trypes), Sphaerotrypes , Diamerini, p. 46. Coptoborus Hopkins, Xyleborini, p. 80, 82. Coptodryas Hopkins, Xyleborini, p. 82, 84 . Coptodryas Schedl { = Cnemonyx), Scolytini, p. 58. Coptogasterl\\igerl = Scohjtus), Scolytini, p. ,59. Coptonotinae, Platypodidae, p. 3. Coptonotus Chapuis, Platypodidae, p. 10, 16, 19, 26. Coptosomus Schedl {~Cnenu>nyx), Scolytini, p. 58. Coriaccphilus Schedl, Cryphalini, p. 88, .90. coriaceus EichhoS (Stephanoderes), Coriacephihis , Cry- phalini, p. 90. corpulentus Samp.son (Cryphalus), Ernocladius, Cry- phalini, p. 90. Coit/ii/riyc/on Schedl, Corthylini (Corthylina), p. 96, 100. Corthyli LeConte ( = Corthylini), p. 93. Corthylidae ( = Corthylini), p. 3. Corthylinae ( = Corthylini), p. 3. Corthylmi, p. 34, 93. Corthylocurus Wood, Corthylini, p. 93, 97, 100. corthyloides Eichhoff ( = Tomicus materiarius Fitch), CUmthotrichus, Corthylini (Corthylina), p. 98. 114 Great Basin Naturalist Memoirs No. 10 Coiihylomimus Ferrari {=Monai-thnim). Corthylini (Corthylina), p. 99. Corilnjhis Erichson, Corthvlini (Corthvlina), p. 3, 93, 97, 100. corticinus Wood (^Xijlogopinus araucariae Schedl), Htj- Ivrdrectonus , Tomicini, p. 43. conjli Ferris {Tomicus, =Lymatitor sepicola), Dryocoe- tini, p. 74. Cosmocorynua Ferrari (-Monarthntm), Corthylini (Cor- thylina), p. 99. Cosmoderes EichhofiF. Cryphalini, p, 3, 98, 91 . costalimai Nunberg {Enwphlociis , = Stcphanoderes sun- daensis Eggers), Hijpothenemus , Cryphalini, p. 92. costaricensis Schedl, Arapfus, Corthylini (Pityophthor- ina), p. 97. costiceps Broun { — Dcndrofnipes vatitus Broun), Den- drotrupea , Tomicini, p. 42. Craniodicticus Blandford, Carphodicticini, p. 67, 67. crenatufi Fabricius, Hyleshius. Hylesinini, p. 38, .39. crenatus Guillebeau (Elzcarius) , Phloeotribus , Phloeo- tribini, p, 49. crenatus Hagedorn, Strornbophorus , Diamerini, p. 46. cristatus Ferrari (Cosmocorynus), Moiiat-thruin. Cor- thylini (Corthylina), p. 99. cristatus Wood, Carphodicticus , Carphodicticini, p. 67. Cryphalidae ( = Cryphalini), p. 3. Cryphalinae ( = Cryphalini), p. 3. Cryphalini, p. 34, 85. Cryp/ia/ites Cockerell (Cryphahni?), p. 25. Cryphalogenes Wood, Cryphalini, p. 88, 90. Cryphaloidea Lindemann (=Cryphalini), p. 8.5. cryphaloides Eichhoff {Triannocerus), Hypothcnemus , Cryphalini, p. 91. Cryphaloides Formenek ( = Coccotrypes), Dryocoetini, p. 75. Cryphalomimetcs Browne { = Stcphanopodius), Crypha- lini, p. 90. Cryphalomimus Browne ( = Stephanopodius), Crypha- lini, p. 90. Cryphalomimus Eggers, Xyloctonini, p. 85, 85. Cryphalomorphus Schaufuss {=Scolytogenes), Crypha- lini, p. 90. Cryplmlophilus Schedl { = Scohjtodes), Ctenophorini, p. 61, 62. Cryphalops Reitter ( = £riioporu.s), Cryphalini, p. 90. Cryphalus Erichson, Cryphalini, p. 3, 24, 87, 89, 91 . Cryphyophthorus Schedl, Hypoborini, p. 54, 54. Cryptarthrum Blandford (^Cryphalus), Cryphalini, p. 91. Cryptocarenus Eggers, Cryphalini, p. 20, 87, 89, 91 . C ryptocleptes Blackman { = Pseudothysanocs), Micracini, p. 63. Cryptocurus Schedl, Hylesinini, p. 37, 39. cryptographus Ratzeburg {Heteroborips), Xylcborus, Xyleborini, p. 83. Cryptoxyleborus Schedl, Xyleborini, p. 82, 84. C ryptuloclcpttis Wood { = Pseudothysanoes). Micracini, p. 63. Crypturgi LeConte ( = Crypturgini), p. 75. Crypturgidae (=Crypturgini), p. 3. Crypturginae (=Crypturgini), p. 3. Crypturgini, p. 34, 7.5. Crypturgus Erichson, Crypturgini, p. 3, 75, 76, 76. Ctenophoridae Chapuis ( = Ctenophorini), p. 59. Ctenophorini, p. 33, 59. Ctcnophorus Chapuis ( = Scohjtodes), Ctenophorini, p. 59, 60. Ctemjophthorus Schedl (-Pityophthorus), Corthyhni (Pityophthorina), p. 98. Ctonocryphus Schedl { = Glostatus), Xyloctonini, p. 85. Ctonoxylon Hagedorn, Xyloctonini, p. .33, 85, cS5. culcitatus Blandford (Epomadius), Scolytodes, Cten- ophorini, p. 60. Cumatotomicus Ferrari ( = /p.s), Ipini, p. 70. Curculionidae, p. 28. Curculionoidea, p. 25. ciirta or cui-tus Eggers (Kissophagus), Phloeoditica , Phloeosinini, p. .52. curvidens Germar (/;«■), Pityokteines , Ipini, p. 68. curvifcr Walker {Hylesiiuis, =Acanthurus spinipennis Eichhofl), Diamcrus, Diamerini, p. 45. Cyclorhipidion Hagedorn { = Xylehorus), Xyleborini, p. 83. ' Cylindrotomicus Eggers { = Scolytogenes), Cryphalini, p. 90. Cyrtogenius Strohmeyer, Dryocoetini, p. 70, 71, 73, 74. Cyrtotomicus Ferrari {-Ips}, Ipini, p. 70. Dacnophthorus Wood, Corthylini, p. 95, 98. Dacryophthorus Schedl ( = Lipartlirum), Hypoborini, p. 54. Dacryostactus Schau(uss, Hypoborini, p. 54, 55. Dacnjphalus Hopkins ( = Hypocryphalus), Cryphalini, p. 90. Dactylipalpi Blandford ( = Hylesinini), p. 36. Ductylipalpus Chapuis, Hylesinini, p. 30, 36, 38, 39. dactyliperda Fabricius (Bostrichus), Coccotrypes, Dry- ocoetini, p. 75. Dactylopselaphus Gemminger & Harold ( — Dactylipal- pus), Hylesinini, p. 39. Jnru.'i;ii Eichhoff, Scolytogenes , Cryphahni, p. 90. Dendriops Schedl { = Cosmoderes), Cryphalini, p. 91. Dcndrochilus Schedl, Ipini, p. 67, 68, 68. Dcndrographus Sched], Dryocoetini, p. 72, 74. Dactylotrypes Eggers, Dryocoetini, p. 70, 72, 74. dcclividepressus ScUed\. Glostatus. Xyloctonini, p. 85. Dendrocranulus Schedl, Dryocoetini, p. 70, 72, 74 . Dendroctonides Niisslin (=Tomicini), p. 40. Dcndroctonus Erichson, Tomicini, p. 40, 41, 42. Dcndrodicticus Schedl, Carphodicticini, p. 20, 34, 67, 67. Dendrosinus Chapuis, Phloeosinini, p. 19, 50, 51, 52. Dendroterus Blandford, Corthvlini (Pitvophthorina), p. 94, 97. Dendrotrupes Broun, Tomicini, p. 40, 41, 42. Dendrotryptim Schedl ( = lndocryphalus), Xyloterini, p. 79. Dendrurgus¥.g%eTs(=Coccotrypes), Dryocoetini, p. 75. dentipes Blandford, Eupagiocerus , Bothrosternini, p. 48. deprccator Schedl (Pachynoderes), Hypothenemus, Cry- phalini, p. 92. depressus Bright, Cactopinus, Cactopinini, p. 66, Dennestes domesticus Linnaeus ( = Trypodendron), p. 2. Dertnestes micrographus Linnaeus (= Pityophthorus), p. 2. Dennestes piniperda Linnaeus (=romicu.s), p. 2. Dennestes poligraphus Linnaeus { = Polygraphus}, p. 2. Dennestes typographus Linnaeus { = lps), p. 2. Dcropria Enderlein, Crypturgini, p. 33, 75, 76, 76. diudematus Eggers, Cryptocarinus, Cryphalini, p. 91. Diameridae Hagedorn ( = Diamerini), p. 44. 1986 WOOD: Genera of Scolytidae 115 Diamerides Browne ( = Phlococranus), Phloeosinini. p. 51. Diamerini, p. 30. 44 . Diamerinae ( = Diamerini). p. 3. Diamerus Erichson, Diamerini, p. 44, 45. 45 . dilutus Blandfmd. Scohjtomimus, Xyloctonini, p. 8.5. dimorphus Schedl [Pihjophihorus , —Neomips brasiUcn- sis Schedl), Pittjophthorus , Corthylini (Pityoph- thorina), p. 98. dipterocarpi Hopkins, Webbia, Xyleborini, p. 84. dislocatus Blackman (Cnjptocleptes), Pseudothysanoes , Micracini, p. 63. dislocatus Blackman (Cryptidocleptus), Pseiidotlujsa- noes, Micracini, p. 63. dispar Eggers (Stephanoderes). Stephayiopodius , Cr\- phalini, p. 90. dispar Fabricius (Apate, Anisandrus), Xyleborus , Xyle- borini, p. 79, 83. dissimilis Zimmermann (Cnjpturgus, =Stephanoderes chapuisi Eichhoff), Hypothenemus , Cryphalini, p. 91. dohrni Eichhoff {Steganocranus), Amphicranus , Cor- thylini (Corthyhna). p. 99. Do/urgocteptes Schedl. Dryocoetini, p. 72, 73. Dolurgus Eichhoff. Crypturgini, p. 3. 75. 76, 76. do/nestictwi Linnaeus (Dennestes), Trypodendron , Xy- loterini, p. 2, 78. donisthorpei Formenek (Cryphaloides, -Bostrichus carpophagus Hornung). Coccotrypes , Dryocoetini. p. 7.5. drakei Blackman ( = Cryphalus rigidus LeConte), Pseu- dothysanoes , Micracini, p. 63. Dryocoetes Eichhoff, Dryocoetini, p. 3, 25, 70. 73, 74 , Dryocoetini, p. 34, 70. Dryocoetiops Schedl, Dryocoetini. p. 70, 73, 75. Dryocoetoideae Lindeniann ( = Dryocoetini), p, 70. Dryocoetoides Hopkins, Xyleborini, p. 80. 82 . Dryotomicus Wood ( = Phloeotribus), Phloeotribini, p. 49. Dryotomus Chapuis ( = Phtoeotribus}. Phloeotribini, p. 49. dubius Eggers, Hylesinopsis, Hylesinini. p. 39. dubius Hagedom (Myetophilites), Hylurgops , Hvlastini, p. 36. duprezi Hoffmann (Xyleborus , Paraxijichorus , =Tomic- us materiarius Fitch), Gnathotrichus , Corthylini (Corthyhna), p. 99. durus Schedl IProtopityophthorus , =Dendrographus pygnuieus Eggers), Dendrographus , Dryocoetini. p. 74. £ccoptoga.ster Gyllenhal( = Sco/!/(u.s), Scolytini, p. 59. Eccoptogastrinae ( = Scolytini), p. 3. Eccoptopterus Motschulsky. Xyleborini, p. 33. 82, 84 . eggersi Schedl IBrachydendrulus), Araptus, Corthylini (Pityophthorina). p. 97. eggersi Schedl, Cryphyophthorus , Hypoborini, p. .54. Eggersia Lebedev ( = Pityogenes), Ipini, p. 68. ehlersii Eichhoff IStephanoderes. Homoeocryphahis , = Hypothenemus eruditus Westwood), Hypothene- mus, Cryphalini, p. 91. eichhoffi Blandford (Hylesinosoma, -Hylesinus fici), Aricerus, Phloeotribini, p. 49. Eidophelinae Murayama( = Cryphalini), p. 85. Eidophelus Eichhoff. Cryphalini, p. 3, 85, 88, .90. Ekkoptogaster Herbst { = Scolytus), Scolytini, p. 59. elaphus Kichhoii [Stephanoderes , ~ Adiaeretus spinosus Hagedorn), Hypothenemus, Cryphalini, p. 91. e/ec(rinus Germar (Hi//e.5inife.5), Hylurgops, Hyastini, p. 36. e/fgons Eichhoff. Hylocurus, Micracini, p. 65. elegans Krivohitzkaja (Phellodendrophagus), Eidophe- lus, Cryphalini, p. 90. elongatum Eggers (Aplunuirthrum), Deropria, Cryp- turgini, p. 76. elongatus Schedl (Landolphianus), Lanurgus, Micracini, p. 63. e/ongafus Schreiner, Tiarophorus, Dryocoetini, p, 73. elsac Eggers, Pseudomicracis, Micracini, p. 25, 65. Elzearius Guillebeau { = Phloeotribus), Phloeotribini, p. 49. emarginatus Hopkins { = Xyleborus vespatorius Schedl), Coptoborus , -Xyleborini, p. 82. endroedyi Schedl (Hemihylesinus), Hyle.nnopsis , Hyle- sinini, p. .39. Eocryphalus Kurentzov ( = Ernoporicus), Cryphalini. p. 89, Epomadius Blandford (^Scolytodes), Ctenophorini, p. 60. Epsips Beeson ( = Hypothenemus), Cryphalini, p. 92. Ericryphalus Hopkins { = Cryphalus), Cryphalini, p. 91, Erineophitus Hopkins { = Scolytodes), Ctenophorini, p. 60. Erineosinus Blackman {=Lipiii-thrum), Hypoborini, p. 54. Erioschidias Schedl (^Cosmoderes), Cryphalini. p, 91. Ernocladius Wood, Cryphalini, p. 88, 90. Ernocryphalus Muravann {- Crijphalus), Crvphalini, p. 91. ErnophloeiisT^unheTgi = Hypothenemus), Crvphalini, p. 92. Ernoporicus Berger, Cryphalini, p. 87, 88, 89. Ernoporides Hopkins { = Scolytogenes), Cryphalini. p. 90. Ernoporinae Niisslin ( = Cryphalini). p. 85. Ernoporus Thomson, Cryphalini, p. 85, 88. 90. eruditus Westwood. Hypothenemus, Cryphalini, p. 91. eruditus Westwood {Hypothenemus , Homoeocryphahis , = Stephanodcres ehler.Hii Eichhoff), Hypothenemus, Cryphalini, p. 91. Estenoborus Reitter ( = Carphoborus), Polvgraphini. p. .56. Ethadopselaphus Blandford ( = Dactylipalpus), Hylesin- ini. p. 39. Eulepiops Schedl { = Cyrtogenius), Dryocoetini, p. 74. Eulytocerus Blandford ( = Phloeotribus), Phloeotribini, ' p. 49. euonymi Kurentzov, Allernoporus , Cryphalini, p. 89. Eupagiocerus Blandford, Bothrosternini, p. 19, 46, 48. euphorhiae Bright (Afrotrypetus), Styracoptinus , Hypo- borini, p. 54. euphorbiae Kiister (Bostrichus), Thamnurgus, Dryocoe- tini, p. 73. euphorbiae Wollaston, Aphiniui-thrum, Crvpturgini. p. 76, euphorbiae Wood. Cryphalogenes, Cryphalini, p. 90. Eupteroxylon Eggers ( = Monarthrum), Corthyhni (Cor- thyhna), p. 99. Euptilius Schedl (^Ernoporus), Cryphalini, p. 90. Eurydactylus Hagedorn ( = Eccoptopterus), Xyleborini, p. 84. 116 Great Basin Naturalist Memoirs No. 10 Euwallacea Hopkins, Xyleborini, p. 81, 83. excisus Ferrari (Morizus), Cotihijlus. Corthylini (Cor- thylina), p. 100. exornatus Schedl, Traglostus, Micracini, p. 63. exul Reitter (Thamnurgus) , Taphronurgus , Drvocoetini. p. 7.3. fallax Hagedorn (Meringopalpus), Gymiiochilus . Cten- ophorini, p. 60. fallax Hagedorn (Trigonogenius, Hagedornus), Pitijoph- thorus, Corthylini (Corthylina), p. 98. fallax Hagedorn (Trigonogenius). PUyophthorus , Cor- thylini (Corthylina), p. 98. fasciatus Say {Bostrichus, Corthylomimus). Monar- thrum, Corthylini (Corthylina), p. 99. /tci Erichson, Hypohorns, Hypoborini, p. .54. fici Lea (Hylesinus. Hylesinosoina , =Aricerus eiclilioffi Blandford), Aricerus, Phloeotribini, p. 49. Ficicis Lea. Hylesinini, p. .37, 38, .39. Ficipliagufi Murayama (=Fidcis), Hylesinini, p. .39. fihnn Reitter, Cisurgus, Cr\'pturgini, p. 76. fimbricornis LeConte, Thysanoes , Micracini, p. 65. flavicornis C\\Ap\x\s [Loganius] , Cnemonyx. Scolytini, p. .58. fleutiaiixi Blandford (Progenius , =Xyleborus subcostatus Eichhoff), Xyleborus , Xyleborini, p. 83. floridensis Blackman (Tachyderes , =Cryptocarenus se- riatus Eggers), Cnjptocarenus, Cryphalini, p. 91. floridensis Hopkins ( = Ernoporides knabi Hopkins), Scohjtogenes , Cryphalini, p. 90. fraxini Panzer (Bostrichus, Leperisinus. = Bostrichus t'or/us Fabricins), Hylesinus , Hylesinini, p. 39. frontalis Fabricius (Bostrichus, =Pagiocerus rimosus Eichhoff), Pagiocerus , Bothrosternini, p. 48. galeritus Eichhoff, Cnemonyx . Scolvtini, p. 20, 24, 34, 58. Genyocerinae, Platypodidae, p. 3. georgiae Hopkins, Trischidias , Cryphalini, p. 93. ghanaensis Schedl (Hypocryphalus, Cryphalomimetes), Stephanopodius , Cryphalini, p. 90. ghanaensis Schedl (Hypocryphalus, Cryplmlomimus), Stephanopodius , Cryphalini, p. 90. glaber Eichhoff (Hc.vaco/H.s), Scolytodes , Ctenophorini, p. 60. glaber Sched\ (Eidepiops), Cyriogenius, Drvocoetini, p. 74. glabratulus Schedl (Ctenyophthorus), Pitijophthorus , Corthylini (Pityophthorina), p. 98. glabratus Schedl, Hyleops. Phloeosinini, p. .52. globosus Eichhoff (Hylesinus), Dendrosinus, Phloeo- sinini, p. 52. globosus Hagedorn (Hapalogenius), Rhopalopsclion , Hylesinini, p. 39. globosus Strohmeyer, Glochiphorus . Hypoborini, p. 54. globus Blandford, Hadrodemius, Xyleborini, p. 84. GlochicopterusSc\\eA\(=Hylesinopsis), Hylesinini, p. 39. Glochinocerus Blandford, Corthylini (Corthylina), p. 96, 99. Glochiphorus Strohmeyer, Hypoborini, p. 54, 54 . Glostatus Schedl, Xyloctonini, p. 85, 85 . Glyptoderus Eichhoff (=Tri/popWoeus), Cryphalini, p. 3,89. Gnatharus Wood & Huang, CorthvHni (Corthylina), p. 96,99. Gnathoborus Schedl (=Araptus), Corthylina (Pityoph- thorina), p. 97. Gnathocorius Schedl ( = Gnathotrupes), Corthyhni (Cor- thylina), p. 99. Gnathocranus Schedl (=Araptus), Corthylini (Pityoph- thorina), p. 97. Gnathoglochinus Schedl ( = Gnathotrupes), Corthylini (Corthylina), p. 99. Gnatholcptus Blackman, Corthylini, p. 95, 98. Gnathomimus Sched\( = Gnathotrupcs), Corthyhni (Cor- thylina), p. 99. Gnathophorus Schedl (=Pityophthorus), Corthyhni (Pityophthorina), p. 98. Gnathophthorus Wood ( = Pityophthorus), Corthyhni (Pityophthorina), p. 98. Gnathotrichoidcs Blackman ( = Gnathotrichus), Corthy- lini (Corthylina), p. 98. Gnathotrichus Eichhoff, Corthylini (Corthylina), p. 3, 93, 95, 98 . gnathotrichus Schedl (Taphramites), (?==Dryocoetes, Dryocoetini), p. 25. Gnathotrupes Schedl, Corthvlini (Corthylina), p. 4, 5, 9, 17, 95, 99 . Gnathotrypanus Wood ( = Gnathotrupes), Corthylini (Corthylina), p. 99. goliathoides Murayama (Ficiphagus , =Hylesinus porca- ti(.s Chapuis), Ficicis, Hylesinini, p. 39. gracile Eichhoff ( = Hypohorus hispidus Ferrari), Pyc- narthrum, Ctenophorini, p. 60. gracilipes KichhoS (Platydactylus), Eccoptopterus , Xyle- borini, p. 84. gracilipes Eichhoff (Platydactylus, Eurydactylus) , Ec- coptopterus, Xyleborini, p. 84. grandiclava Thomson (Pseudopolygraphus), Polygra- phus, Polygraphini, p. 56. grandis Beeson (Pelicerus, Ozodendron), Cyriogenius , Dryocoetini, p. 74. grandis Erichson (Phloeotrupes), Phloeoborus, Hyle- sinini, p. 39. grandis Swaine ( = Hylurgus sericeus Mannerheim), Pseudohylesinus , Tomicini, p. 42. granulatum Blandford, Styphlosoma, Corthylini (Pity- ophthorina), p. 97. granulatus Schedl (Hylescierites), (? = Hylurgops), Hy- lastini, p. 25, .36. granulicollis Schedl (Dendriops), Cosmoderes, Crypha- lini, p. 91. granulicollis Schedl (Neodiamerus), Acacacis, Diamer- ini, p. 45. Gretschkinia Sokanovoskii ( = Pseudothy.mnoes), Micrac- ini, p. 63. guanajuatensis Duges (Anaeretus), Xyleborus , Xylebor- ini, p. 83. guatemalensis Hopkins (—Xyleborus capucinus Eich- hoff), Dryocoetoidcs, Xyleborini, p. 82. guineensis Hagedorn (Hijphacne, =Bostrichus carpoph- agus Hornung), Coccotrypes, Dryocoetini, p. 75. giwnnensis Murayama (Pruiniphagus), Xylechinus, To- micini. p. 42. Gymnochilus Eichhoff, Ctenophorini, p. .59, 60, 60. Hadrodemius Wood, Xyleborini, p. 82, 84 . Hagedornus Lucas ( = Pityophthorus), Corthylini (Pity- ophthorina), p. 98. Hapalogenius Hagedorn ( = Rhupalop.wlion), Hylesinini, p. .39. Hapalophloeus Schedl ( = Hylesinopsis), p. 39. 1986 WOOD: Genera of Scolytidae 117 hederae Schmitt (Htilesinus). Kissophagus . Hvlesinini, p. 38. Hemicryphalus Schedl, Crvplialini, p. 88, 90. Hemihi/Iesimis Schedl { = Hiilesinopsis), Hylesinini, p. 39. henshawi Hopkins { = Ericryphahis, =Hijpothrncmti>i sylvicola Perkins), Cnjphalus. Cr\plialini, p. 91. Heferofaorips Reitter ( = Xi//ffcon(,s), Xyleborini, p. 83. heterodoxus Casey (Renocis). Chaetophlocus. Hypo- borini, p. 54. heterodoxus Cisey (Renocis , = Pseudocnjphahis brittaini Swaine), Chactopbloeus , Hypoborini, p. 54. Hexacolidae ( = Ctenophorini), p. 3. Hexacolinae ( = Ctenophorini), p. 3. HexacoUnus Schedl (^ Scohjtodes). Ctenophorini, p. 60. Hexacolus Eichhoff ( = Sco/i/torfes), Ctenophorini, p. 3, 60. hicoriae LeConte, Cbramcsus, Phloeosinini, p. 52. hispidus Ferrari (Hypobonis, Monebius, =Ncmobius lambottei Chapuis), Pycnarthrum, Ctenophorini, p. 60. hispidus Ferrari {Hijpoborus, =Neinobius lambottei Chapuis), Pycnarthrum . Ctenophorini, p. 60. hispidus Ferrari (Hypobonis. Nomebius, =Nemobius /amfoortei Chapuis), Pycmirthrum . Ctenophorini, p. 60. hispidus Ferrari (Hypobonis, -Pycnarthrum gracilc Eichhoff), Pycnarthrum , Ctenophorini, p. 60. hispidus Klug(Hi//e.sin!(.s), Diamerus , Diamerini, p. 45, Holonthogaster Gemminger & Harold ( = Olontho- gaster). Phloeosinini, p. 52. Homarus Broun ( = Chaetoptelius), Tomicini, p. 42. Homoeocryphalus Lindemann ( = Hypothenemus). Cry- phalini, p. 91. HophtesEg^ers( = Cladoctonus), Phloeosinini, p. .52. HoplitontusWood( = Cladoctonus), Phloeosinini, p. .52. HoplitophthorusVi/ood( = Cladoctonus), Phloeosinini, p. 52. hubbardi Schwa.TZ, Cactopinus , Cactopinini, p. 66. Hylastes Erichson, Hylastini, p. 34, .35, 36. Hylastes LeConte (-Hylastini), p. .34. Hylastini, p. 30, .34 . Hylastinides Niisshn ( = Hylesinini), p. 36. Hylastinoides SpessWtzev {=Alniphagus), Hvlesinini, p. .39. Hylastinus Bedel, Hylesinini, p. 30, 36, 37, 38. Hylastites Hagedorn ( = HyUirgops), Hylastini, p. 36. Hyledius Sampson ( = Olonthogaster). Phloeosinini, p. .52. Hyleops Schedl, Phloeosinini, p. 19, 49, 51, .52. HylescieritesSched\(? = Hyhirgops), Hylastini, p. 25, .36. Hylesinen Erichson ( = Hylesinini), p. .34, 36. Hylesininae, p. 3, 19, 29, 34. Hylesinini, p. 30, 36. H ylesinites Germar (- Hylurgops ), Hylastini, p. 36. Hijlesinopsis Eggers, Hylesinini, p. 36, .37, 38, 39. HylesinosoDW Lea ( = Ariceru.s), Phloeotribini, p. 49. Hyk.mius Fabricius, Hylesinini, p. 2, 34, 36, 37, 38, 39. Hylocuridae Eichhoff( = Micracini), p. 3, 61. Hylocurinae ( = Micracini), p. 3. Hylocurosoma Eggers ( = Scobjtodes), Ctenophorini, p. 60. Hylocurus Eichhoff, Micracini, p. 3, 61, 63, 65. Hyloniu.':Nnnherf'(—Xylncleptes), Dryocoetini, p. 74. Hyloperus Browne {=Cryptocurus}, Hylesinini, p. .39. Hyhtrdrectonus Schedl, Tomicini, p. 40, 41, 42. Hylurgini LeConte (=Tomicini), p. 40. hyhu-goides Schedl (Coptodryas), Cnemonyx , Scolytini, p. 58. hylurgoidcs Schedl (Coptodryas, Coptosomus), Cne- monyx, Scolytini, p. .58. Hylurgonotus Schedl, Tomicini, p. 40, 42, 43. Hyhirgopinus Swaine, Tomicini, p. 40, 41, 42. Hyhirgops, LeConte, Hylastini, p. 25, 35, 35. Hylurguhis Eggers ( = Olonthogaster), Phloeosinini, p. .52. Hylurgus Latreille, Tomicini, p. 40, 41, 42. hymenaeae Eggers (Neodryocoetcs), Araptus. Corthylini (Pityophthorina), p. 97. Hyorrhynchinae Hopkins (- Hyorrhnchini), p. 43. Hyorrhynchini, p. 30, 43 . Hyorrhynchus Blandford, Hyorrhynchini, p. 43, 44, 44. Hypaspistcs Hagedorn (-Tiarophorus), Drvocoetini, p. 73. Hypertensus Hagedorn (=ArapfH.s), Corthylini (Pityoph- thorina), p. 97. Hyphacnc Hagedorn (=Coccotrypes), Dryocoetini, p. 75. Hypoborinae Niisslin ( = Hypoborini), p. .53. Hypoborini, p. 41, .53. Hypoborus Erichson, Hypoborini, p. 3, 54, 54. Hypocryphahis Hopkins, Cryphalini, p. 89, 90.. Hypopityophthorus Bright ( — Pityophthorus), Corthylini (Pityophthorina), p. 9S. Hypothcncmus Westwood, Cryphalini, p. 20, S7, 89, 91 . Hypothenoides Hopkins ( = Scohitogenes), Cryphalini, p. ■ 90. hystrix LeConte (Hylcsinus). Chaetophlocus , Hypobor- ini, p. 54. illepidus Schedl (Browneia, =Pityophthorus obliquus LeConte), Amhrosiodmus . Xyleborini, p. 83. imitans Eichhoff, Eidophclus , Cryphalini, p. 90. impressus Schedl (Gnuthoglochinus), Gnathotrupes, Corthylini (Corthylina), p. 99. incotnptus Wood, Chramcsus, Phloeosinini, p. 52. Indocryphahis Eggers, Xyloterini, p. 78, 78 . ingcns Eichhoff (P/if/iOriii.s), Monai-thrum, Corthylini (Corthylina), p. 99. inops Wood (Hypopityophthorus), Pityophthorus , Cor- thylini (Pityophthorina), p. 98. insularis Schedl (Xylochihis, —Dendroterus saUaei Blandford), Dendroterus , Corthylini (Pityophthor- ina), p. 97. intermedins Sampson (Xylotenis , = Indocryphahis mal- aisei Eggers), Indocryphahis , Xyloterini, p. 78. intcrniptiis Eggers (Hophtes, -HopUtophthorus sentus Wood), Cladoctonus , Phloeosinini, p. 52. intricatus Ratzeburg (Eccoptoguster , Tubuh)scohjtus), Scohjtus, Scolytini, p. 59. inurbaniis Broun (Apate), Mesoscohjtus, Xvleliorini, p. 84. Ipidae ( = lpini), p. 3. Ipinae (=lpini), p 3. Ipini Bedel, p. 31, 67. /;«■ DeGeer, Ipini, p. 67, 68, 70. Isofjhthorus Schvd] (~ Acanthotomicus), Ipini, p. 70. Ithyceridae, p. 28. ivoriensis Nunliei'g ( = Chortastus .similis Eggers), Serras- tus, Polygraphini, p. .59. jakipae Letzner (Bostrichus , LetznereUa ), Scob/togenes , Cryphalini, p. 90. 118 Great Basin Naturalist Memoirs No. 10 jandiacus Enderlein, Coleobothrus , Crypturgini, p. 76. joveri Schedl (Trypugraphus), Hylesinopsis, Hvlesinini, p. 39. joveri Schedl {Xyleborus, Prenuiophilus], Premtiohius , Xyleborini, p. 82. jiicundtis Chapuis, (Ceratolepis), Cneinonijx , Sc-olvtini, p. 58. Jugocryphalus Tsai & Li { = Cryphalus), Cryphalini, p. 9L kalshoveni Schedl (Notoxylcborus), Xyleborus , Xylebor- ini, p. 83. keilbachi Schedl, Carphoborites , Polygraphini, p. 25. Kelantanius Nunberg ( = Tenninalinus), .Xyleborini, p. 83. kenyaensis Schedl (Afromicracis), Miocryphabis , Mi- cracini, p. 63. Key to the families of Curculionoidea, p. 26. Key to the subfamilies and tribes of Scolytidae, p. 28. Key to the genera of Bothrosternini, p. 46. Key to the genera of Cryphalini, p. 87. Key to the genera of Crypturgini, p. 76. Key to the genera of Ctenophorini, p. 60. Key to the genera of Diamerini, p. 45. Key to the genera of Dryocoetini, p. 71. Key to the genera of Hylastini, p. 35. Key to the genera of Hylesinini, p. 37. Key to the genera of Hyorrhynchini, p. 44. Key to the genera of Hypoborini, p. 53. Key to the genera of Phloeosinini, p. 50. Key to the genera of Phloeotribini, p. 49. Key to the genera of Polygraphini, p. 56. Key to the genera of Scolytini, p. 58. Key to the genera of Tomicini, p. 40. Key to the genera of Xyleborini, p. 79. Key to the genera of Xyloterini, p. 78. Kissophagus Chapuis, Hylesinini, p. 37, 38 . knabi Hopkins (Ernoporides , =Ernoporides floridensis Hopkins), Scohjtogenea , Cryphalini, p. 90. kotbei Hagedorn (Allarthrum), Cryphahis, Cryphalini, p. 24, 91. kolbei Schaufuss, Dacryostactus , Hypoborini, p. 55. kumatoensis Niisima (Orosiotes), Cyrtogenius , Dryocoe- tini, p. 74. laevigatus Chapuis {Ctenophorus , =Hexacohts levis Blackman), Scolytodes , Ctenophorini, p. 60. laevigatus Eggers {Pityophtlwrus, Neopityophthorus), Araptus, Corthylini (Pityophthorina), p. 97. /aeuigaf us Ferrari, Scolytodes , Ctenophorini, p. 60. Wm Strohmeyer (Oiopemon), Dryocoetiops , Dryocoe- tini, p. 75. lamhottei Chapuis (Nemobius, =Hypoborus hispidus Ferrari), Pycnarthrum, Ctenophorini, p. 60. lamhottei Chapuis {Nemobius, Monebius. = Hypuboriis hispidus Ferrari), Pycnarthrum, Ctenophorini, p. 60. lambottei Chapuis (Nemobius, Nomebius. =Hypoborus hispidus Ferrari), Pijcnarthrum . Ctenophorini, p. 60. LandolphianusSc\ied\(=Lanurgus), Micracini, p. 63. Lanurgus Eggers, Micracini, p. 62, 63. laricis Fabricius [Bostrichus, Neotomicus], Orthotomi- cus, Ipini, p. 70. laricis Fabricius (Bostrichus), Orthotomicus), Ipini, p. 68. lasiocarpi Swaine (Orthotomicus, Orthotomides), Pityok- teines, Ipini, p. 68. laterale EichhoS (Pterocyclon), Monarthrum. Corthylini (Corthylina), p. 99. lederi Reitter (Cryphalus, Cryphalops, =Bostrichus tiliac Panzer), Ernoporus, Cryphalini, p. 90. Leiparthrum Wollaston ( = Liparthrum) , Hypoborini, p. 54. L(?perismu.s Reitter ( = H!/fcsmu.s), Hylesinini, p. 39. Lepicerinus Hinton ( = Scolytoge7ies), Cryphalini, p. 90. Lepiceroides Schedl (^ Hypothcnemus), Cryphalini, p. 92. Lepicerus Eichhofi( = S(:o/(/fog<;ne.s), Cryphalini, p. 3, 9L Lepisomus Kirby { = Polygraphus), Polygraphini, p. 56. Leptoxyleborus Wood, Xyleborini, p. 80, 82. Letznerella Reitter ( = ScoIytogcnes), Cryphalini, p. 90. letzneri Ferrari (Pscudocorthylus), Corthylus, Corthylini (Corthylina), p. 100. levis Blackman (Hexacolus, =Ctenophorus laevigatus Chapuis), Scolytodes , Ctenophorini, p. 60. lewisi Blandford, Hyorrhynchus. Hyorrhynchini, p. 44. lichtensteini Ratzeburg (Bostrichus), Pityophthorus , Corthylini (Corthylina), p. 98. ligniperda Fabricius (Hylesinus), Hylurgus , Tomicini, p. 42. lirrmx Schedl (Sphenoceros), Araptus, Corthylini (Pity- ophthorina), p. 97. tonax Schedl (Sp/i(?nocerus , =Hypertensus reitter Hage- dorn), Araptus, Corthylini (Pityophthorina), p. 97. lineatus Olivier (Bostrichus, Xyloterus), Trypodendron , Xyloterini, p. 78. Liparthrum Wollaston, Hypoborini, p, 3, 53, 54, 54. lipingensis Tsai & Li (Acryphalus), Cryphalus, Cry- phalini, p. 91. Li^soclastus Schaufuss ( = Diamerus), Diamerini, p. 45. litseae Browne (Diamerides , = Phloeocranus bruchoides Schedl), Phloeocranus , Phloeosinini, p. 51. Loganm.s' Chapuis ( = Cnemonyx), Scolytini, p. 58. longicollis Wollaston (Xyloterus, = Dactylotrypes uytten- boogaarti Eggers), Dactylotrypes , Dryocoetini, p. 74. lungipennis Wickham, Xtjleborites , ?=Xyleborini, p. 25. luffa Beeson (Vitaderes), Cosmoderes, Cryphalini, p. 91. Lymantor, Loyendal, Dryocoetini, p. 70, 72, 74. machnovskii Sokanoyskii (Hypothenemus , Saliciphilus), Taphrorychus, Dryocoetini, p. 74. maculatus Beeson (Scolytomimus, Scolytocleptes), Scoly- tomimus. Xyloctonini, p. 85. madagascariensis Schedl, Pityodendron, Corthylina (Pityophthorina), p. 98. magnus Eggers (Stephanoderes , Chondronoderes) , Hy- pothenemus, Cryphalini, p. 91. /nrtgrjw\ Sampson, Cnestus, Xyleborini, p. 8.3. malaisei Eggers, Imlocryphalus , .Xyloterini, p. 78. malayensis Schedl, Bothro.itcrnoides , Diamerini, p. 45. j)ia/ga.ssicH.s Schedl, Dolurgocleptes , Dryocoetini, p. 72, 74. nuimibiae Wood, Phloeographus , Polygraphini, p. 56. mancus Blandford (Xyleborus, Apoxyleborus), Xylosan- drus , Xyleborini, p. 83. mandibularis Blackman (-^Pityophthorus shannoni Blackman), Gnatholeptus , Corthylini (Pityophthor- ina), p. 98. margadilaonis Hopkins, Margadillius , Cryphalini, p. 91. Margadillius Hopkins, Cryphalini, p. 89, 91 . marginatus Hagedorn (Xestips), Xylocleptes, Dryocoe- tini, p. 74. 1986 WOOD; Genera of Scolytidae 119 marowitzi Semenov {Scohjtus. Pinetoscolytus), Scobjtus, Scolytini, p. 59. materiarius Fitch (Tomicus. =Gnathotrichus corthij- loides EichhofD, Gnathotrichus , Corthylini (Cor- thylina). p. 98. materiarius Fitch {Tomicus. Paraxijicbonis, =Xijlehorus duprezi Hoffmann), Gnathotrichus, Corthyhni (Corthyhna), p. 99. Mecopehnus Blackman, Platypodidae, p. 16, 18. melodori Hopkins (Stephanorhopahis), Ernoporus , Cry- phahni, p. 90. Mesoscobjtus Broun, Xyleborini, p. 81, S4 . Metacorthyhis Blandford, Corthyhni, p. 96, 99. Metahi/lastes Eggers ( = Cijrtogenius). Drvocoetini, p. 74. Metahylesinus Eggers { = Hylcsinopsis), Hvlesinini, p. 39. meuseli Reitter (Xyleborips), Xyleborus , Xvleborini, p. 83. mexicanus Blandford, Dendrotcrus , Corthyhni (Pityoph- thorina), p. 97. micans Kugelann (Bostrichus), Dendroctonus . Tomicini, p. 42. Micracidae ( = Micracini), p. 3. Micracidendron Schedl ( = Mimiocurus). Corthylini (Pityophthorina), p. 97. Micracides LeConte ( = Micracini), p. 61. Micracinae ( = Micracini), p. 3. Micracini, p. 33, 61 . MicraciopsSched\{ = Lamirgus), Micracini, p. 63. Micracis LeConte, Micracini, p. 3, 25, 61, 63, 65. Micracisella Blackman, Micracini, p. 61, 63, 65. Micracisoides B\ackmiin { = Hi/h>cunis). Micracini, p. 65. Microhorus Blandford, Ctenophorini, p. 59, 60, 60. Microcorthylus Ferrari, Corthylini, p. 96, 99. micrographus Linnaeus (Dermestes. Pityophthorus), Corthylini (Pityophthorina), p. 2. Micropenis Wood { = Coptodryas). Xyleborini, p. 84. mikado Blandford (Scolytoplatypus , Taeniocerus), Scobj- toplatypus, Scolytoplatypodini, p. 61. mikado Blandford (Scobjtoplatypus, Strophionocerus), Scohjtoplatypus , Scolytoplatypodini, p. 61. mikuniyamensis Murayama (Pseudopoecihps), Taphro- rychus Dryocoetini, p. 74. Mimidendrulus Schedl ( = Cyrtogenius), Drvocoetini, p. 74. Mimiocurus Schedl, Corthylini, p. 93, 94, 97. Mimiophthorus Schedl { = Mimiocurus), Corthylini (Pityophthorina), p. 97. Mimips Eggers (- Acanthotomicus), Ipini, p. 70. minimus Fabricius (Hylesinus), Carphuborus. Poly- graphini, p. 56. Minulus Eggers (Cnemonyx). Scolytini, p. 58. minutissimus Schedl ( = Hexacobnus minutissimus Schedl), Scobjtodes , Ctenophorini, p. 60. minutissimus Schedl (^ Hcohjtodes minutissimus Schedl), Scolytodes , Ctenophorini, p. 60. minutK.^imus Zimniermann {Crypturgus), Pseudopi- tyophthorus , Corthylini (Pityophthorina), p. 98. Miocryphabis Schcd\. Micracini, p. 62, 63. mirahilis Schedl (Streptocranus), Coptoborus, Xyle- borini, p. 82. Mixodentatae, p. 3. Monarthrum Kirsch, Corthylini (Corthyhna), p. 96, 99. Monebius Hophns ( = Pycnarthrum), Ctenophorini, p. 60. mongobca Sokanovskii (Grctschkinia), Pseudothysanoes , Micracini, p. 63. monibculbs Eichhoff, Cosmodercs , Cryphalini, p. 91. iiionibcollis Eichhod i = Vitaderes biffa Beeson), Cosmod- ercs, Cryphalini, p. 91. monographus Fabricius (Bostriclius), Xylebot-us, Xyle- borini, p. 83. itiontanum Schedl (Micracidendron), Mimiocurus , Cor- thylini (Pityophthorina), p. 97. inontanus Schedl (Brachydendrubis, Mimiophthorus), Mimiocurus. Corthylini (Pityophthorina), p. 97, morigerus Blandford (Xt/leborus), Xykisandrus, Xyle- borini, p. 83. moritzi Ferrari, Broc/u/.s/irtitii.v, Corthylini (Corthylina), p. 100. moritzi Ferrari (Bracbyspartus. Thyb(rcos), Brachyspar- tus , Corthylini (Corthylina), p. 100. Morizus Ferrari (--Cortliybis), Corthylini (Corthylina), p. 100. movobae Schedl (Mimidendrulus), Cyrtogenius, Dry- ocoetini, p. 74. mucronatus Blandford, Craniodicticus, Carphodicticini, p. 67. mucronatus LeConte (Cryphalus , —Procrypliabis popub Hopkins), Procryphabis, Cryphalini, p. 89. nuibistriatus Marsham (Ips, Scobjtochebis), Scolytus, Scolytini, p. 59. muhistriatus Marsham (Ips, Spinuloscobjtus), Scolytus, Scolytini, p. 59. muhistriatus Wood, Scobjtodes, Ctenophorini, p. 60. miindidus Broun (Homarns, Acrantus), Chaetoptelius , Tomicini, p. 42. mundulus Broun (Homarus ), Chaetoptelius , Tomicini, p. 42, murayamai Schedl ( = Tosaxyleborus pallidipennis Mu- rayama), Cncstus, Xyleborini, p. 84. murex Blandford, Styracopterus , Hypoborini, p. 54. murex Blandford (Styracopterus), Styracoptinus , Hypo- borini, p. 54. Myeloborus Blackman ( = Pityophthorus), Corthylini (Pityophthorina), p. 98. Mijclophilites Hagedorn (=^ Hybirgops) , Hylastini, p. 36. Myelophibis Eichhoff (-Toniici/.s), Tomicini, p. 42. naevus Schedl, Cryptoxylcborus, Xyleborini, p. 84. natalensis Eggers (Stephanoderes), Miocryphalus, Mi- cracini, p. 63. natalensis Schedl (Archaeophalus), Hypothenemus, Cry- phalini, p. 91. Negritus Eggers ( = Scobjtogenes), Cryphalini, p. 90. Nemonychidae, p. 26. Nemopagiocerus Schedl ( = Eupagioccrus), Bothrostern- ini, p. 48. Neocryphalus Eggers (-Scolytogcnes), Cryphalini, p. 90. Neocryphus Nunberg, Cryphalini, p. 87, 88, 90. Neodiamerus Schedl (^Acacacis), Diamerini, p. 45. Neodryococtes Eggers (=Araptus), Corthylini (Pityoph- thorina), p. 97. Neoglostatus Schedl ( = Pseudothy.sanoes), Micracini, p. 63. Neohylciinus Eggers ( = Phrixosoma), Phrixosomini, p. 43. Neoviips Schedl (^Pityttphtliorus), Corthylini (Pityoph- thorina), p. 98, Neophlocotrihus Eggers ( = Phloeotribus), Phlocotribini, p, 49. 120 Great Basin Naturalist Memoirs No. 10 Neopittjophthonts Schedl i=Araptuii), Corthylini (Pity- ophthorina), p. 97. Neopteleobius Nobuclii, Hylesinini, p. 37, 38, .39. Neotomicus Fuchs ( = Orthototnicus), Ipini, p. 70. Neoxyloctonus Eggers { = Scolytomimus), X.vloctonini, p. 85. nevemianni Schedl (Nemopagiocerus , =Eupagioceru'i ater Eggers), Eupagiocerus , Bothrosternini, p. 48. nigripennis Blandford, Metacorthyhis , Corthylini (Cor- thylina), p. 99. nigrivestris Schedl (Ctonocryphiis, Paraglostatus), Glo- status, Xyloctonini, p. 85. niisinuii Eggers (Hyorrhynchus), Neohyorrhynchus , Hy- orrhynchini, p. 44. niisimai Eggers {Hyorrhynchus, =Stietts aphaerotry- poides Murayama), Sueus, Hyorrhynchini, p. 44. nitens Schedl (Ozodendron). Peridryococtes , Dryocoe- tini, p. 74. nitidicollis Motschulsky {Olot^thogaster , Hutontho- gaster), Olonthogaster , Phloeosinini. p. 52. nitidicollis Motschulsky, Olonthogaster , Phloeosinini, p. 52. nitidulus Guerin-Meneville {Piezorhopalus , =Amphi- cranus thoracicus Erichson), Amphicranus , Cor- thylini (Corthyhna), p. 99. nitidus Eggers (Adryocoetes), Coptodryas, Xyleborini, p. 84. nitidus Hagedorn {Lepicerus, Pelicerus), Cyrtogenius, Dryocoetini, p. 74. Nomebius Navas { = Pycnarthrum), Ctenophorini, p. 60. nothofagi Schedl (Gnathominius), Gnathotrupcs , Cor- thylini (Corthylina), p. 99. f^otoxyleborusSc\\eA\(=Xyleborus). Xyleborini, p. 83, novateutonicus Schedl (Gnathocranus), Araptus, Cor- thylini (Pityophthorina), p. 97. nubilus Blandford (Phloeotribus, Neophloeotribus), Phlocotribus , Phloeotribini, p. 49. obesus Hopkins (Dacryphalus), Hypocryphalus, Cry- phalini, p. 91. obliquecaudata Motschulsky {Phloeotrogus ), Amhrosiod- tnus, Xyleborini, p. 83. obliquus LeConte (Pityophthorus, Browneia, =Xyle- borus illepidus Schedl), Ambrosiodmus , Xyleborini, p. 83. obscurus Eggers, Bufonus, Tribe?, p. 25. obscurus Marsham (Ips), Hylastinus, Hylesinini, p. 38. octodentatus Schedl {Pterocyclonoides) , Tricolus, Cor- thylini (Corthylina), p. 99. oleae Fabricius {Hylesinus , —Scolytus scarabaeoides Bernard), P/i/oeo*n7)!«, Phloeotribini, p. 49. Olonthogaster Motschulsky, Phloeosinini, p. 19, 49. opacicollis LeConte (Micracis), MicraciscUa , Micracini, p. 65. opacicollis LeConte (Alicracis, Pseudomicracis). Mi- cracisella, Micracini, p. 65. orientalis Schedl, Cardroctonus , Polygrapiiini, p. .56. Orosiotes Niisima( = C(/rfogPHtii.s), Dryocoetini, p. 74. Orthaspistes Hagedorn ( = Tiarophorus), Drvocoetini, p. 73. Orthotomicus Ferrari, Ipini, p. 31, 67, 68, 6S. Orthotoynides Wood ( — Pityokteines), Ipini, p. 68. ovicollis Blandford, Tricolus, Corthylini (Corthvlina), p. 99. Oxycorynidae, p. 27. Ozodendron Schedl ( = Cyi-togenius), Dryocoetini, p. 74. Ozopcmon Hagedorn, Dryocoetini, p, .33, 70, 73, 75. Ozophagus Eggers ( = Pulygraphus), Polygraphini, p. 56. Pachycotes Sharp, Tomicini, p. 40, 42, 43 . Pachynoderes Schedl { = Hypothenemus), Crvphalini, p. 92. Pagiocerus Eichhoff, Bothrosternini, p. 19, 46, 48 . palaquius Schedl {Phlocochilus), Liparthrum, Hypo- borini, p. 54. palauensis Wood (Phloeotrypetus), Liparthrum, Hypo- borini, p. 54. pallidipennis Murayama (Tosaxyleborus , =Cnestus mu- rayamai Schedl), Cnestus, Xyleborini, p. 83. papuanus Schedl (Xylocryptus), Scolytogenes , Cry- phalini, p. 90. Paracorthylus Wood { = Metacorthyhis), Corthylini {Cor- thylina), p. 99. Paraglostatus Schedl i^Glostatus), Xyloctonini, p. 85. Parasphaerotrypcs Murayama { = Sphaerotrypes), Di- amerini, p. 46. Paraxyleborus Hoffmann ( = Gnathotrichus), Corthylini (Corthylina), p. 99. parvulus Ferrari, Microcorthylus . Corthylini (Corthy- lina), p. 99. parvus Hopkins (Hypothenoides), Scolytogenes, Cry- phalini, p. 90. Pelicerus Eggers { = Cyrtogenius}, Dryocoetini, p. 74. pelliculosum Hagedorn (Cyclorhipidion , =Xyleborus perlaetus Schedl), Xyleborus, Xyleborini, p. 83. percorthylus Schedl, Taphrodasus , Xyleborini, p. 84. peregrinus Chapuis {Hylesinus, ^Pachycotes ventralis Sharp), Pachycotes , Tomicini, p. 42, 43. pei'foliatus Wollaston (Phloeophthorus), Phloeotribus, Phloeotribini, p. 49. Peridryococtes Wood, Dryocoetini, p. 73, 74. Periocryphalus Wood, Cryphalini, p. 87, 89, 93. perlaetus Schedl (Xyleborus , = Cyclorhipidion pelliculo- sum Hagedorn), Xyleborus . Xyleborini, p. 83. permirus Schaufuss, Scolytoplatypus , Scolytoplatypo- dini, p. 61. Pernophorus Strohmeyer, Diamerini, p. 44, 45, 46. perrisi Chapuis (Hylesinus, Estenoborus), Carphoborus , Polygraphini, p. 56. persicae Hopkins (Thamnurgides , =Coccotrypes advena Blandford), Coccotnjpes, Dryocoetini, p. 75. perspectus Schedl (Plesiophthorus), Dendroterus, Cor- thylini (Pit\()phthorina). p, 97. Phacrylus Schedl (-Acorthylus), Cryphalini, p. 90. Phellodendrophagus Krivolutzkaja ( = Eidophelus), Cry- phalini, p. 90. Phelloterus Wood, Corthylini (Pityophthorina), p. 95, 98 . philippinensis Eggers (Neoxyloctonus), Scolytomimus, Xyloctonini, p. 85. Phloeobori Blandford ( = Hylesinini), p. 36. Phloeoborinae ( = Hylesinini), p. 3. Phloeoborus Erichson, Hylesinini, p. .36, 38, 39. Phlocochilus Schedl ( = Liparthrum), Hypoborini, p. .54. Phloeocleptus Wood, Micracini, p. 61, 63, 65. Phloeocranus Schedl, Phloeosinini, p. 50, 51 . Phloeocurus Wood, Micracini, p. 63, 65. Phloeoditica Schedl, Phloeosinini, p. 50, 52. Phloeographus Wood, Polygraphini, p. 56, 56. Phloeophthorus Wollaston ( = Phloeotribus), Phloeotrib- ini, p. 49. Phloeosinides Niisslin ( = Phloeosinini), p. 49. Phloeosinini, p. 19,31,49. 1986 WOOD: Genera of Scolytidae 121 Phloeosinites Hagedorn, Phloeosinini, p. 51, 52. Phloeosinopsis Schedl (1936)(=Ofoi!f/ioga.ster), Phloeo- sinini, p. 52. Phloeosinopsis Schedl (1964){ = Phloeosinoides), Phloeo- sinini, p. 52. Phloeosinopsoides Schedl, Phloeosinini, p. 50, 51, 52. Phloeosinus Chapuis, Phloeosinini, p. 19, 50, 51, 52. Phloeoterus Wood, Corthylini (Pityophthorina), p. 94, 97. Phloeotribidae Chapuis ( = Phloeotribini), p. 48. Phloeotribinae (Phloeotribini), p. 3. Phloeotribini, p. 31.48. Phloeotribus Latreille, Phloeotribini, p. 10. 17, 19, 48, 49, 49 , 66, 67. Phloeotrogus Motschulsk)' (=Arnbrosiodmiis]. Xylebor- ini, p. 83. Phloeotrupes Erichson ( = Phloeoborus), Hvlesinini, p. 36, 39. Phloeotrupides Chapuis ( = Hylesinini), p. 36. Phloeotrupini ( = Hylesinini), p. 3. Phloeotrypetus Wood i=Liparthrum). Hypoborini, p. 54. Phloiotribus Latreille { = PhIoeotribus), p. 49. Phrixosoma Blandford, Phrixosomini, p. 4.3 . Phri.\osomini Wood, p. 30. 4.3 . Phthorius Eichhoff ( = Monarthrum). Corthylini (Cor- thylina), p. 3, 99. Phthorophloeides Niisslin ( = Phloeotribini), p. 48. Phthorophloeus Rey { = PhIoeotribus), Phloeotribini, p. 48, 49. piceus Eggers (Jugocnjphalus). Cryphahis. Crvphalini, p. 91. Piezorhopalus Guerin-Meneville ( = Amphicranus). Cor- thylini (Corthyhna), p. 99. pila Blandford, Sphaerotrypes . Diamerini, p. 45. Pilidentatae, p. 3. pilo.sus Eggers {Ips, Mimips). Acanthotumicus , Ipini, p. 70. pilosus LeConte (Cryphalus, Ancyloderes). Gnatho- trichus, Corthylini (Corthylina), p. 98. pilosus Ratzeburg (Hylesinus), Xylechinus , Tomicini, p. 42. pimelioides Schaufuss (Lis.wcla.stus), Diamerus. Di- amerini, p. 45. Pinetoscolytus Butovitsch ( = Scolytus). Scolytini, p. .59. pini Hopkins (Carpo.sinu.s), Cyrtogenius. Drvocoetini, p. 74. pini Hopkins (Piperus. ^Hypothenemus sylvicola Per- kins), Cryphalus. Crvphalini, p. 91. piniarius Schedl, Hylurdrectonus , Tomicini, p. 42. pinifex Fitch {Hylastes, =rugipennis pinifex Fitch), Hy- lurgops. Hylastini, p. .35, .36. piniperda Fabricius (Hylesinus . =Dermestes piniperda Linnaeus), Tomicus, Tomicini, p. 42. piniperda Linnaeus (Dermestes. Blastophagus). Tomi- cus, Tomicini, p. 42. piniperda Linnaeus (Dermestes. -Hylesinus piniperda Fabricius), Tomicus, Tomicini, p. 2. piniperda Linnaeus (Dermestes. Myelophilus). Tomicus. Tomicini. p. 42. Piperius Hopkins (^Cryphalus). Cryphalini, p. 91. Pityoborus Blackman, Corthylini (Pityophthorina), p. 94, .97. Pityoceragenes Balachowsky (-Pityogenes). Ipini, p. 68. Pityodendron Schedl, Corthylini (Pityophthorina), p. 93, 95, .9*. Pityogenes Bedel, Ipini, p. 67, 68, 6S. Pityoktcines Fuchs, Ipini, p. .33, 67, 68, 68. Pityophilus Blackman ( = Pityotrichus), Corthylini (Pity- ophthorina), p. 98. Pityophthoridae Eichhoff( = Corthylini), p. 3, 93. Pityophthoridca Wickham (not Scolytidae), p. 25. Pityophthorina, Corthylina, p. 94, 97. Pityophthoroides Blackman ( = Pityophthorus). Corthy- lini (Pityophthorina), p. 98. Pityophlhorus Eichhoff, Corthylini, p. 3, 6, 71, 93, 95, 98. Pityotrichus Wood, Corthylini, p. 95, 98. planicolle Schedl (Chilodendrun). Hylc^inopsis. Hvle- sinini, p. 39. Platydactylus Eichhoff ( = Eccoptopterus). .Xyleborini, p. 84. Platypodidae, p. 4, 27. Platypodidinae, Platypodidae, p. 3. Plesiophthorus Schedl ( = Dendroterus). Corthylini (Pity- ophthorina), p. 97. podocarpi Schedl ( = Pseudohylocurus caplandicus Nun- berg), Lanurgus. Micracini, p. 63. Poecilips Schaufuss ( = Coccotrypes). Dryocoetini, p. 75. poligraphus Linnaeus (Dermestes, ^Hylesinus pubes- cens Fabricius), Polygraphus. Polvgraphini, p. 2, 56. politus Say (Bostrichus). Xyloterinus . Xyloterini, p. 79. Polygraphidae Chapuis ( = Polygraphini), p. .55. Polygraphini. p. 31, 55. Polygraphus Erichson, Polygraphini, p. 19, 55, 56, 56. populi Hopkins ( = Cryphalus niucronatus LeConte), Procryphalus. Cryphalini, p. 89. porcatus Chapuis (Htjlesinus , =Ficiphagus goliathoides Murayama), Hylesinus. Hvlesinini, p. .39. porteri Brethes, Sinophloeus . Tomicini, p. 42. Premnohius Eichhoff, Xyleborini, p. 3, .33, 79, 82. PremnophilusBrowne( = Premnobius), Xyleborini, p. 82. pn'7ni(i(.v Schedl (Tap/irof(?ru.s), Taphrorychus , Dryocoe- tini, p. 74. primus Bright (Prognathotrichus), Gnathotrichus , Cor- thylini (Corthylina), p. 99. Prionosceles Blandford ( = Scolytodes). Ctenophorini, p. 60. Problechilidae Eichhoff ( = Ctenophorini), p. 3, 59. Problechilus Eichhoff( = G(/ninoc/ii/i/.s), Ctehophorini, p. 3, .59, 60. Prochramesus'Wood( — Chrame,sus). Phloeosinini, p. .52, Procryphalus Hopkins, Ciyphalini, p. 87, 88, 89. Progenius Blandford (=Xyleborus), Xyleborini, p. 83. Prognathotrichus Bright ( = Gnathotrichus). Corthylini (Corthylina), p. 99. Protohytastcs Wood, Platypodidae, p. 6, 10, 11, 15, 16, 17, IS, 19, 20, 23, 27.' Protopityophthorus Schedl (-Dcndrographus), Dry- ocoetini, p. 74. Protoplatypus Wood. Platypodidae, p. 10, 11, 16, 18, 20, 21,24,27. Prowebbia Browne ( = Webbia}. Xyleborini, p. 84. Pruniphagus Murayama ( = X!//ec/!in».s), Tomicini, p. 42. psaltes Hagedorn (Bothryperus). Phrixosoma, Phrixoso- mini, p. 43. Pseudoacacacis Schedl (=Acacacis), Diamerini, p. 45. Pseudocliramesus Blackman, Phloeosinini, p. 50, 51, 52. Pseudocorthylus Ferrari ( = Corthylus), Corthylini (Cor- thylina), p. 100. 122 Great Basin Naturalist Memoirs No. 10 Pseudocryphalus Ferrari {=Cryphalus}, Crvphalini, p. 91. Pseudocn/phaltis Swaine ( = Renocis), Hypoborini, p. .54. Pseudocnjpturgus Eggers {=Microborus), Ctenophor- ini, p. 60. Pseudodiamcrus Eggers, Diamerini, p. 45, 4.5. Pseudohijlesinus Eggers ( = Hij}esinopsis). Hylesiniiii, p. ,39. Pseudohijlesinus Swaine, Tomicini, p. 20, 40, 41, 42. Pseudohi/Iocurus Nunberg { = Lanurgus), Micracini, p. 63. ' Pseudohtjorrhynchus Murayama, Hyorrhynchiiii, p. 44, 44. Pseudomicracis Blackman { = MicraciseUa), Micracini, p. 65. Pseudomicracis Eggers, Micracini, p. 25, 63, 6.5. Pseudophloeotribus Eggers i=HyIesinopsis), Hvlesinini, p. 39. Pseudopityophthorus Swaine, Corthylini, p. 95, 9S. Pseudopoecilips MuTayama { = Taphrorychus), Dryocoe- tini, p. 74. Pseudopohjfiraphus Seitner l = PoIygraphus), Polygraph- ini, p. .56. Pseudothamniirgus Eggers { — Tiarophorus), Dryocoe- tini, p. 73. Pseudothy.^anoes Blackman, Micracini, p. 33, 61, 62, 63. Pseudowebhia Browne { = Wehbia), Xyleborini, p. 84. Pseudoxijleborus Wood {=Atnasa), Xyleborini, p. 83. Pseudoxijlechinus Wood & Huang, Tomicini, p. 41, 42. Pteleobius Bedel, Hylesinini, p. 37, 38. Pterocyclon EichhoS { = Monarthrum), Corthylini (Cor- thylina), p. 3, 99. Pterocyclonoides Schedl { = Tricohis), Corthylini (Cor- thylina), p. 99. Ptilopodius Hopkins, Cryphalini, p. 89, 91 . puberulus Chapuis (Dnjotomus, Drijotoinicus), Phloeo- tribus, Phloeotribini, p. 49. ;)»/«"n(/HS Chapuis (Dnjotomus), Phloeotrihus, Phloeo- tribini, p. 49. pubcscens Fabricius (Hijlesinus, =Dennestcs poligra- p/jus Linnaeus), Polygraphus , Polygraphini, p. 56. pubescens Zimmermann. Xyleborus , Xyleborini, p. 25. pudens Blackman (Pityophthoroides), Pityophthorus . Corthylini (Pityophthorina), p. 98. pucUus Reitter, Tomicus. Tomicini, p. 41. puUus Schedl (Xyleborus, Adryocoetes), Coptodnjas, Xyleborini, p. 84. puUus Wood, Periocryphalus Cryphalini, p. 93. pumilus Mannerheim (Hylastes), Dolurgus, Cryptur- gini, p. 76. puncticollis Blandford, Scohjtopsis , Scolytini, p. 58. punctatopilosus Schedl (Xyleborus, Kelantanius), Termi- nalinus, Xyleborini, p. 83. pusillus Gyllenhal (Bostrichus), Crypturgus, Cryptur- gini, p. 76. Pycnarthrum Eichhoff, Ctenophorini, p. 3, 20, 23, 59, 60,60. Pygmaeoscohjtus Butovitsch ( = Scolytus), Scolvtini, p. 59. pygmacus Eggers (Pelicerus), Dendrographus , Dryocoe- tini, p. 74. pygmaeus Eggers ( = Protopityophthorus dtirus Schedl), Dendrographus, Dryocoetini, p. 74. pyg7naeus Fabricius (Bostrichus, Pygmaeoscohjtus), Scolytus, Scolytini, p. 59. quadridens Hartig (Bostrichus , PHyoceragenes), Pityo- genes, Ipini, p. 68. quadridens Wood, Chrame.'^us, Phloeosinini, p. 53. quadrioculatus Eggers (Neohylesinus). Phrixosoma, Phrixosomini, p. 4.3. quadrioculatus Hagedorn (Spongotarsus), Polygraphus , Polygraphini, p. 56. quadrituberculatus Schedl (Isophthiirus), Acanthotomi- cus , Ipini, p. 70. ramiperda Swaine (Pityophthorus, Myelobortis), Pity- ophthorus, Corthylini (Pityophthorina), p. 98. regius Hagedorn, Ozopemon , Dryocoetini, p. 75. rehi Hagedorn, Phlocosinites , Phloeosinini, p. 52. rcitteri Hagedorn (Hypertensus, =Sphenoceros Umax Schedl), Araptus, Corthylini (Pityophthorina), p. 97. Rcnocjs Casey { = Chactophlocus), Hypoborini, p. 54. retusipetmis Blandford, Glochinoccrus, Corthylini (Cor- thylina), p. 99. rhizophorae Eggers (Dendrurgus , =Spermatoplex rhi- zophorae Hopkins), Coccotrypes, Dryocoetini, p. 75. rhizophorae Hopkins (Spennatoplex), Coccotrypes, Dryocoetini, p. 75. rhizophorae Hopkins (Spcnmitoplex, =Dendrurgus rhi- zophorae Eggers), Coccotrypes, Dryocoetini, p. 75. rhododactylus Marsham, Phtoeotribus, Phloeotribini, p. 17. rhois Blackman ( = Pseudothysanoes sulcatus Bruck), Stenoclyptus , Micracini, p. 65. Rhopalocrijphus Nunberg ( = G/o.sfafus), Xyloctonini, p. 85. Rhopalopleurus Chapuis ( = Chramesus), Phloeosinini, p. .52. Rhopalopselion Hagedorn, Hylesinini, p. .37, 38, ,39. Rhynchitidae, p. 28. rigidus LeConte (Cryphalus, = Pseudothysanoes drahei Blackman), Pseudothysanoes , Micracini, p. 63. riinosus EichhoS (-Bostrichus frontalis Fabricius), Pa- giocerus, Bothrosternini, p. 48. rotundatus Chapuis (Chramesus, =Dendrosinus bon- nairei Reitter), Chrame.nis, Phloeosinini, p. .52. rotundus Hopkins, Hypocryphalus, Cryphalini, p. 90. rude Blandford, Phrixosonw , Phrixosomini, p. 43. rudis Erichson, Phlocoborus , Hylesinini, p. 39. rudis LeConte (Micracis, Micracisoides), Hylocurus, Micracini, p. 65. rufipennisKirby (Apate, Lcpisoinus), Polygraphus , Poly- graphini, p. 56. rufipes Eichhoff (Anchonocerus), Monarfhrum , Cor- thylini (Corthyhna), p. 99. rufipes Eichhofr(H(//fl.sf('.s), Hylurgopinus, Tomicini, p. 42. rufopatiatus Eichhoff, Araptus, Corthylini (Pityophthor- ina), p. 97. ra/n/i(,s Schedl, Chiloxylon, Dryocoetini, p. 75. rugipennis Schedl (Carpophloeus), Cyrtogenius, Dry- ocoetini, p. 74. rugosipes Hopkins, Arixyleborus , Xyleborini, p. 83. rugosissiinus Cockerell, Cryphalitcs, Cryphalini?, p. 25. rugulosus Miiller (Bostrichus, Ruguloscolytus), Scoly- tus, Scolytini, p. 59. Saetidentatae, p. 3. Saliciphilus Sokanoyskii ( = Taphrorychus), Dryocoetini, p. 74. 1986 WOOD; Genera of Scolytidae 123 sallaei Blandford {Dendroterus , =Xylochilus- instilaris Schedl), Dendroterus. Corthylini (Pityophthorina), p. 97. Sampso7!ius Eggers, Xyleborini, p. 80, 82. sannio Schaufuss (Poecilip.^). Coccotnjpe.s. Dryocoetini, p. 75. sauropterus Schedl, Sauroptilius. Corthylini (Pityoph- thorina), p. 98. Sauroptilius Browne, Corthylini (Pityophthorina), p. 9S. Saurotocis Wood, Micracini, p. 63, 6.5. saxc^eni Ratzeburg (Bo.^triclnis). Xijicboriiuis . .Xyle- borini, p. 84. scarabaeoides Bernard (Scohjtus. =Hijlesinus oleae Fab- ricius), Phloeotribus . Phloeotribini, p. 49. Schedlarius Wood. Platypodidae, p. 11, 16, 18. Schedlia Browne, Xyleborini, p. 82, 84. schellwieni Hagedorn (Hi/lastitcs). Htjiuraops . Hvla.stini, p. 36. schwarzi Hopkins {Erineophibis), Scolijtodes. Cten- ophorini, p. 60. Scierus LeConte, Hylastini, p. 34, 35, .3.5. Scolytarii Latreille (^Scolytidae, Scolytinae, Scolytini), p. 28, 57. Scolytidae, p. 3, 6,27, 2S. Scolytinae, p. 3, 20, 29. Scolytini, p. 31, 57. Scolytochehis Reitter ( = Sco/!/f!/.s), Scolytini, p. 59. Scobjtocleptes Schedl {-Scobjtoinimus), Xvloctonini, p. 85. Scohjtodes Ferrari, Ctenophorini, p 20, .33, 59, 60, 60. Scolytogenes Eichhoff, Cryphalini, p. 3, 87, 88, .90. Scolytoidea, p. 3. scolytoides Eichhoff, Xyloctomis , Xyloctonini, p. 85. Scolytomimus Blandford, Xyloctonini, p. 13, 84, 85, 85 . Scolytoplatypini Blandford ( = Scolytoplatypodini), p. 61. Scolytoplatypodidae ( = Scolytoplatypini), p. 3. Scolytoplatypodinae ( = Scolytoplatypini), p. 3. Seolytoplatypodini, p. .33, 61 . Scohjtoplatypus Sch'iufufis, Scolytoplatypini. p. 113. Scolytopsis Blandford, Scolytini, p. 58, 58 . Sco/ytotor.su.s Schedl, Platypodidae, p. 10, 16, 18, 26. ■icolytus Fabricius (Bostrichus , Coptogaster), Scohjtus. Scolytini, p. .59. scolytus Fabricius [Bostrichus, Eccoptogaster), Scohj- tus, Scolytini, p. .59. scolytus Fabricius (Bo.Htrichus, Ehhoptogastcr). Scoly- tus, Scolytini, p. .59. scolytus Fabricius (Bostrichus), Scoh/tus, Scolytini, p. .59. Scolytus Geoffrov, Scolytini, p. 2, 6, 13, 17, 20, 28, .57, 58,58. scrofa Schedl (Ctadoctoproctus). Triotemnus, Dryocoe- tini, p. 73. scrutator Pandelle [Thamnurgus, Pseudolhutnniirgus), Tiarophorus, Dryocoetini, p. 73. .icutulatus Blandford (Hylesinus), Neopteleobius, Hyle- sinini, p. .39. securigerus Blackman (Chalcohyus), Pseudothysanoes, Micracini, p. 63. semenovi Kurentzov (Eocryphalus). Ernoporicus , Cry- phalini, p. 89. sentosus Wood (Hoplitophthorus , =Hoplites interruptus Eggers), Cladnctonus , Phloeosinini, p. 52. sepicola Lovendal ( = Tomicus coryli Perris), Lymantor, Dryocoetini, p. 74. .•icriatus Eggers { = Tachyderes floridcnsis Blackman), Cryptucarcnus, Cryphalini, p. 91. sericcus Mannerheim {Hylurgus, -Pseudohylesinus grundis Swaine), Pslcudohylesinus . Toinicini, p. 42. Serra.stus Nunberg, Polygraphini, p. 20, 55, 56, 56. setistriutus Lea (Cri/p/ia/i/s, Erioschidias), Cosmoderes, Cryphalini, p. 91. sexdentatus Boerner (Dermestes, Cumatotoinicus, = Bostrichusstenographus Duftschniidt), Ips, Ipini, p. 70. sexdentatus Eggers, Sampsonius , .Xyleborini, p. 82. sexspinosiis Mot.schulsky ( = Scolytus spinosus Olivier), Eccoptopterus, Xyleborini, p. 84. seydeti Nunberg (Rhopalocryphus), Glostatus, Xyloc- tonini, p. 8.5. shannoni Blackman (Pityophthorus . = Gnatholeptus mamhbularis Blackman), Gnatholeptus , Corthylini (Pityophthorina), p. 98. sharpi Blandford (Xylcborus). Taurodemus, Xvleborini, p. S3. sicula Del Guercio (Comcsiella , ^ Bo.'itrichus brevicollis Kolcnati), Phloeotribus, Phloeotribini, p. 49. sidneyanus Nordlinger (P.scudocryphalus). Cryphahis. Cr\phalini, p. 91. similis Eggers (Chorta.'itus, =Serrastus ivoriensis Nun- berg), Serrastus, Polygraphini, p. .56. Sinophloeus Brethes, Tomicini, p. 40, 41, 42. sordicauda Motschulsky {Phloeotrogtis), Leptoxylcbor- us, Xyleborini, p. 82. sparsepilosus Schedl {Gnathoi>horus), Pityophthorus, Corthylini (Pityophthorina), p. 98. sparsepUosus Schedl (Gnathophorus, Gnathophthorus), Pityophthorus, Corthylini (Pityophthorina), p. 98. SpernuitopU'x Hopkins { = Coccotrypes), Dryocoetini, p. 75. Spermophthorus Cosia Lima, Corthylini, p. 95, 98. .ipessivtzevi Berger, Ernoporicus , Cryphalini, p. 89. Sphaerosinus Eggers {=Phrixosoma}, Phrixosomini, p. 43. Sphaerotrypes Blandford, Diamerini, p. 44, 45, 45. Sphaerotrypini Murayama (=^ Diamerini), p. 44. sphaerotrypoides Murayama {^ Hyorrlujnchus niisimai Eggers), Sueus. Hyorrhynchini, p. 44. Sphcnoceros Schedl {=Araptus), Corthylini (Pityoph- thorina), p. 97. .•ipinatus Schedl (Bostrichips), Pseudothysanoes, Mi- cracini, p. 63. Spinidentatae, p. 3. spinifer Schedl (Phloeosimis , =Phloeosinopsis annatus Schedl), Olonthogastcr , Phloeosinini, p. 52. spinipennis Eichhoff (Acfliif/iia'H.v, =Hylesinus curvifer Walker). Diamerus, Diamerini, p. 45. spinipennis Schedl, Cryptocurus , Hylesinini, p. 39. spinosus Blandford, Acanthotomicus , Ipini, p. 70. spinosus Hagedorn (Adiaeretus, ^Stcphanodercs ela- phus Eichhoff), Hypothenemus , Cryphalini, p. 91. .S'pino.sH.s Olivier (Scolytus, = Eccoptopterus .sexspinosus Mot.schulsky), Eccoptopterus, Xyleborini, p. 84. Spinuloscolytus Butovitsch ( = Scolytus), Scolytini, p. 59. S])inulosus Rcy (Phloeophthorus), Phloeotribus , Phloeo- triliini, p. 49. Spongocerinae Hagedorn (=^ Seolytoplatypodini), p. 3, 61. Spongocerus Blandford (^Scolytoplatypus), Seolyto- platypodini, p. 61. 124 Great Basin Naturalist Memoirs No. 10 Spongotarsus Hagedorn {=Pobjgraphus, Polygraphini, p. 56. Squamasinulus Nunhergi -Xylechinus). Tomicini, p. 42. squanwsiis Blackman (Erineosinus), Liparthrum, Hypo- borini, p. 54. squamosus Schedl (Neoglo^tatus), Pseiidothysaiioes . Mi- cracini, p. 6.3. sqtiamulosus Eggers (Cylindrotumicus), Scolytogenes , Cryphalini, p. 90. Steganocramis Eichhoff i=Amphkrantis), Cortliylini (Corthylina), p. 3, 99. Stegomerns Wood, Cryphalini, p. 87, 88, 90. Stenochjpfus Blackman, Micracini, p. 63, 6.5. stenographus Duftschniidt {Bostrichus, Cumalotomicus , = Dermestes sexdentatus Boerner), Ips, Ipini, p. 70. Stephanoderes EichhoS ( = Hypothenenius), Crvplialini, p. 3, 91. Stephanopodius Sched\, Cryphalini, p. 88, .90. Stephanorhopalus Hopkins { = Ernoporus). Cryphalini, p. 90. stephegynis Hopkins (Boroxylon, =Phloeotrogus hidcn- tatus Motschulsky), Xylehorus , Xyleborini, p. 83. stephegynis Hopkins, Ptilopodius , Cryphalini, p. 91. Sternobothrus Eggers, Bothrosternini, p. 19, 46, 48. Streptocranus Hopkins (^Coptoborus), Xvleborini, p. 82. striatum or striatus Eggers (Hylocurosoma), Scobjtodes , Ctenophorini, p. 60. striatits Eggers, Cryphidomimus, Xyloctonini, p. 85. sfriafu.s Eggers, Pseudodianicrus, Dianierini, p. 45. striatus Eggers (Sphaerosintts), Phrixosoma, Phri.xoso- mini, p. 4.3. strigicoUis LeConte, Ctiesinus , Bothrosternini, p. 46. strigicolbs LeConte { — Nemophihts strigillafus Chapiiis), Cnesinus, Bothrosternini, p. 47. strigillatus Chapuis (=Nemophibis , =Cnesinus strigicol- lis LeConte), Cnesinus. Bothrosternini, p. 47. Strombophorini Schedl ( = Dianierini), p. 44. Strombophorus Hagedorn, Diamerini, p. 44, 45, 46. strombosiopsis Schedl, Dendrochibis , Ipini, p. 68. Strophionocerus Sampson { = Scohjtoplatypus). Scolyto- platypodini, p. 61. Stylotentus Schedl ( = Hypothenemus), Cryphalini, p. 91. Styphlosoma Blandford, Corthylini, p. 94, 97. Styracopterus Blandford { = Styracoptinus). Hvpoborini, p. 54. Styracoptinus Wood, Hypoborini, p. 54, .54 . sid?costatus Eichhoff ( = Prooe?uus fleutiauxi Blandford), Xyleborus , Xyleborini, p. 83. .su/:>refHSu.s Wollaston, Triotemnus , Dryocoetini, p. 73. stdmculae Browne {Prowcbbia ), Webbia , Xyleborini, p. 84. sulcatus Bruck (Pseudothysanoes , =Stenocleptus rhois Blackman, Stenocleptus , Micracini, p. 65. suturabs LeConte, Micracis, Micracini, p. 65. Sueus Murayama, Hyorrynchini, p. 19, 23, 29, 30, 43, 44, 44. ' sulcatus LeConte (Cryphahts, Gnathotrichoides), Gnathotrichus, Corthylini (Corthylina), p. 98. sumatranus Eggers (Hybirgidus). Olonthogaster , Phloeosinini, p. 52. sumatranus Hagedorn, Schedlia, .Xyleborini, p. 84. sundaensis Eggers ( = Ernophloeus costaUmai Nunberg), Hypothenemus, Cryphalini, p. 91. sylvarum Beeson {Epsips), Hypothenemus, Cryphalini, p. 91. sylvicola Perkins {Hypothenemus, =Ericryphabis hen- shawi Hopkins), C ryphahis , Cryphalini, p. 91. sylvicola Perkins {Hypothenemus, =Piperius pini Hop- kins), Cryphalus, Cryphalini, p. 91. Tachyderes Blackman ( = Cryptocarenus), Cryphalini, p. 91. tachygraphus ZimniermanTi, Ambrosiodmus , .Xylebor- ini, p. 83. Taeniocerini Blandford ( = Scolytoplatypodini), p. 61. Jaeniocerus Blandford {-Scolytoplatypus), Scolyto- platypodini, p. 61. Taenioghjptes Bedel { = Cryphalus), Cryphalini, p. 91. Taphramites Schedl {? = Dryocoetes), Dryocoetini, p. 25. Taphroborus Nunherg { = Cyrtogenius), Dryocoetini, p. 74. Taphrodasus Wood, Xyleborini, p. 82, 84. Taphronurgus Reitter, Dryocoetini, p. 71, 72, 73. Taphrorychini Reitter ( = Dryocoetini), p. 70. Taphrorychus Eichhoff, Dryocoetini, p. 3, 72, 74. Taphrostenoxis Schedl { = Cyrtogenius), Dryocoetini, p. 74. Taphroterus Schedl { = Taphrorychus), Dryocoetini, p. 74. forc/ii.s Schedl, Dendrocranuhis, Dryocoetini, p. 74. taunayi Eggers {Xylechinus), XylechinosotJius , Tomicini, p. 42. Taurodemus Wood. Xyleborini, p. 81, S3. tenuis Schedl {Taphrostenoxis), Cyrtogenius , Dryocoe- tini, p. 74. terehratus Wood {Gnathotrypanus), Gnathotrupes, Corthylini (Corthylina), p. 99. terminaliae Hopkins {Tenninalinus). Xyleborus, Xyle- borini, p. 8.3. Tenninalinus Hopkins, Xyleborini, p. 81, 83. tersus Wood, Phelloterus , Corthylini (Pitvophthorina), p. 98. Tes.serocerus Saunders, Platypodidae, p. 4, 9. Thamnophthorus Schedl {=Araptus), Corthylini (Pity- ophthorina), p. 97. Thamnurgides Hopkins ( = Co(?cofrypes), Dryocoetini, p. 75. Thamnurginae Niisslin ( = Dryocoetini), p. 70. Thamnurgus Eichhoff, Dryocoetini, p. 3, 70, 72, 73. Thaumasinulus Reitter { = Chramesus), Phloeosinini, p. 52. theae Eggers {Microperus), Coptodryas , Xyleborini, p. 84. Theoborus Hopkins, Xyleborini, p. 80, 82 . theobromae Hopkins, Theoborus , Xyleborini, p. 82. thoracicus Erichson, Amphicranus, Corthylini (Cor- thylina), p. 99. thoracicus Erichson {Amphicranus, =Piezorhopalus ni- tidulus Guerin-Meneyille), Amphicranus , Corthy- lini (Corthylina), p. 99. thoracicus Lea, Amasa , Xyleborini, p. 83. thujae Ferris (Hylesinus), Phloeosinus, Phloeosinini, p. 52. Thylurcos Schedl ( = Brac/H/.spar/Hs), Corthylini (Corthy- lina), p. 100. Thysanoes LeConte, Micracini, p. 61, 62, 65. Tiarophorus Schreiner, Dryocoetini, p. 25, 33, 34, 72, 73. tibetensis Wood, Gnatharus, Corthylini (Corthylina), p. 99. tiliae Panzer {Bostrichus, Cryphalops, =Cryphalus led- eri Reitter), Ernoporus , Cryphalini, p. 90. 1986 WOOD: Genera of Scolytidae 125 tiliae Panzer (Bostrichus). Ernoporus , Cryphalini, p. 90. togonus Eggers (Pseiidolujlesinus), Hylcsinopsis, Hyle- sinini, p. 39. togonus Eggers {Pseudohylesinus, Metahijtc.iinu.t), Hylc- sinopsis. Hylesinini. p. 39. Tomicidae, p. 3, 40. Tomicini, p. 30, 40. tomicoides Schedl (Micracidendron), Saurotocis, Mi- cracini, p. 6.5. Tomicus Latreille, Tomicini, p. 3, 40, 41, -^2 . Tosaxyleborus Murayama( = Cn£'.sf!(s), Xyleborini, p. 84. Toxophorus Eggers ( = Toxophlhorus). Dryocoetini, p. 25. Toxophthorus Wood, Dryocoetini, p. 25. Traglostus Schedl, Micracini, p. 62, 63 . trahax Sampson (Trogloditica). .Acacacis. Diamerini, p. 45. tran.wersus Chapuis, Dactylipalpus . Hylesinini, p. 39. transversus Chapuis (Dactylipalpus, Dactylopselaphus), Dactylipalpus , Hylesinini, p. .39. trepanicauda Eggers (Xyleborus, Pseudowebbia). Web- bia, Xyleborini, p. 84. Triarmocerus EichhofiF { = Hypothenemiis). Crvplialini, p. 3, 91. Tricolus Blandford, Corthylini, p. 93, 96, .9.9. Trigonogenius Hagedorn { = Pityophthorus). Corthylini (Pityophthorina), p. 98, Triotemnus WoUaston, Dryocoetini, p. 3, 70, 72, 73. TrUchidias Hopkins, Cryphalini, p. 20, 87, 89, 93. triseriatus Schedl, Phloeosinopsis . Phloeosinini, p. 52. triseriatus Schedl [Phloeosinopsis , 1964), Phloeosinop- sloides , Phloeosinini, p. .52. Trogloditica Sampson (^-Acacacis). Diamerini, p. 45. truncatus Eichhoff, Bothrostcrnus . Bothrosternini, p. 48. truncatus Erichson (Tomicus. Anaxyleborus). Amasu, Xyleborini, p. 83. Trypocranus Eichhoff ( = Monart/)n(m), Corthylini (Cor- thylina), p. 3, 99. Trypodendrinae Tredl ( = Xyloterini), p. 76. Trypodendron Stephens, Xyloterini, p. 3, 76, 78, 7S . TrijpographusSched\(^-Hylesinop.',is), Hylesinini, p. 39. Trypophloeinae Niisslin ( = Cryphalini), p. 85. Trypophloeus Fairmaire, Cryphalini, p. 85, 87, 89. tuberculattis Chapuis (Rhopalopleurus), Chramcsus, Phloeosinini, p. .52. tuberculatus Eggers (Hylurgus. =Hylurgonotus brun- neus Schedl), Hylurgonotus, Tomicini, p. 43. Tuhuloscolytus Butovitsch { = Scohjtus), Scolytini, p. .59. tycon Blandford (Scohjtoplatypus , Spongoccrus), Scoly- toplatypus , Scolytoplatypodini, p. 61. typographus Linnaeus (Dermestes, Tomicus), Ips, Ipini p. 2, 70. uniformis Wood & Huang, Pseudoxylechinus , Tomicini p. 42. Urdugraphus Beeson (^Polygraphus), Polvgrapliini, p .56. usagaricus Eggers (Neocryptialus), Scolytogenes Cryphalini, p. 90. u.sfum Schedl, Corthycyclon, Corthylini (Corthvlina), p 100. uyttenboogaarti Eggers (=Xyloterus longicollis Wollas ton), Dactylotrypes , Dryocoetini, p. 74. various Lea, Ficicis, Hylesinini, p. .39. varius Fabricius (Bo.itrichus , Leperisinus -Bostrichus fraxini Panzer), Hylcsinus, Hylesinini, p. 39. laticae Nunherg (Taphroborus), Cyrtogenius , Dryocoe- tini, p. 74. vclutinus Wood (Paracorthylus), Metacortlujlus, Cor- thylini (Corthylina), p. 99. I'Ciifra/is Sharp (-H!//£'siHi(.s;)cn?griii((.s Chapuis), Pachy- cotes, Tomicini, p. 43. vespatorius Schedl ( = Coptoboi'us emarginatus Hop- kins), Coptoborus, Xyleborini, p. 82. vcstitus Broun, Dcndrotrupes , Tomicini, p. 42. vestitus Mulsant & Rey (Hylesir^us, Chaetophorus), Chuetoptelius , Tomicini, p. 42. vestitus Mulsant & Rey (Htjlcsinus), Chuetoptelius, Tomicini, p. 42. villosus Fabricius {Bo.<>trichus, Dryocoetinus). Dry- ocoetes , Dryocoetini, p. 75. Vitaderes Beeson (-Cosmodcres), Cryphalini, p. 91. vittatus Fabricius (Bostrichus), Ptcleobius , Hylesinini, p. .38. volastos Schedl (Thamnophihorus) , Araptus, Corthylini (Pityophthorina), p. 97. volvulus Fabricius (Bostrichus, =Xyleborus guanaju- atensis Duges), Xyleborus , Xyleborini, p. 83. vulgaris Wood, Stcgomerus , Cryphalini, p. 90. uadi Murayama, Pscudohyonhyrichus, Hyorfhychini, p. 44, ' walkcri Blandford (Cryptcnilirum), Cryphalus, Crypha- lini, p. 91. wallacei Blandford, Euwullacea . Xyleborini, p. 83. Wehbia Hopkins, Xyleborini, p. 79, 82, 84. Webbinae Hopkins ( = Xyleborini), p. 3, 79. Xelyborus Schedl (^Webbia), Xyleborini, p. 84. Xe.'itips Hagedorn ( = Xylocleptes), Dryocoetini, p. 74. Xylebori LeConte ( = Xyleborini), p. 79. Xyleboricus Eggers (=Arixyleborus), Xyleborini, p. 86. Xyleboridae (=Xyleborini), p. 3. Xyleborinae (=Xyleborini), p. 3. Xyleborini, p. 34, 79 . Xyleborinus Reitter, Xyleborini, p. 81, 84 . Xyleborips Reitier ( = Xyleborus), Xyleborini, p. 83. Xyleborites Wickham (Xyleborini?), p. 25. Xyleborus Eichhoff, Xyleborini, p. 3, 79, 81, 83. Xylechinides Niisslin (=Tomicini), p. 40. Xylechinites Hagedorn {?=Xylechinus), Tomicini, p. 25. Xylechinops Browne { = Xylcchinus), Tomicini, p. 42. Xylechinosomus Schedl, Tomicini, p. 40, 42 . Xylechintis Chapuis, Tomicini, p. 25, 40, 41, 42. Xylochihis Schedl ( = Dendroterus), Corthylini (Pityoph- thorina), p. 97. Xylocleptes Ferrari, Dryocoetini, p. 3, 25, 70, 72, 74 . Xylocryptus Schedl { = Scolytogenes), Cryphalini, p. 90. Xyloctonidae Eichhoff( = Xyloctonini), p. 3, 84. Xyloctoninae ( = Xyloctonini), p. 3. Xyloctonini, p. 33, 84. Xyloctonus Eichhoff, Xyloctonini, p, 3. 84, 85, 85. xyloctoniis Schedl, Ctonocryphus , Xyloctonini, p. 85. Xylogopinus Sched\{ = Hylurdrectonus), Tomicini, p. 42. Xylosandrus Reitter, Xyleborini, p. 81, S3. Xyloteridae ( = Xyloterini), p. 3. Xyloterini, p. 34, 76. Xyloterinus Swaine, Xyloterini, p. 78, 79. Xyloteroideae Lindemann ( = Xyloterini), p. 76. Xyloterus Erichson ( = Trypodendron), Xyloterini, p. 76, 78. 126 Great Basin Naturalist Memoirs No. 10 jonafijs Eichhoff, Gt/7nnof/ji/us, Ctenopliorini, p. 60. zonatus Eichhoff { = Meringopalpus fallax Hagedorn), zonatus Eichhoff (Problechihis), Gijmnochihis , Cten- Gyimwcliilus , Ctenophorini, p. 60. ophorini, p. 60. Zygophloeus Schedl, Hypoboriiii, p. 53, 54. 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