Great Basin Naturalist Memoirs

Volume 10 A Reclassification of the Genera of Scolytidae (Coleoptera) Article 2

12-1986

A reclassification of the genera of Scolytidae (Coleoptera)

Stephen L. Monte L. Bean Life Science Museum and Department of Zoology, Brigham Young University, Provo, Utah

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Recommended Citation Wood, Stephen L. (1986) "A reclassification of the genera of Scolytidae (Coleoptera)," Great Basin Naturalist Memoirs: Vol. 10 , Article 2. Available at: https://scholarsarchive.byu.edu/gbnm/vol10/iss1/2

This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Great Basin Naturalist Memoirs by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. 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- for each -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 . 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 History my doctoral dissertation. Fortunately, for me and for science, a more limited topic was cho- Linnaeus (17.58) included five species of sen, but the suggestion was never forgotten Scolytidae in his Systema Naturae, edition 10, and has remained a lifelong objective. In all under the name Dennestcs (typographus , 1981, when I was asked by colleagues to final- micrographus poligraphus piniperda, do- , , ize a comprehensive world catalog of Scolyti- mesticus). Fabricius (1801) expanded this to dae, completion of the generic study became include 52 species under the generic names mandatory and urgent. Although this presen- Apate (5), Bo.strichiis (21), and (26). tation is a bit premature, its basic objectives Gemminger and Harold (1872) list 535 species are achieved. More thoughtful selection of in 60 genera. Hagedorn (1910a, b) lists 1,234 illustrations and greater detail in their presen- species in 115 genera. There has been no tation would have been helpful to the user. comprehensive classification or catalog of the The Scolytidae are among the more difficult family on a worldwide basis since the work of to classify; yet their tremendous eco- Hagedorn, but it is estimated that more than nomic importance is so great that immediate 6,000 biological species are currently recog- and precise identification is demanded by nized. The first generic name erected for a those industries threatened by the activities of member of this family was Scohjtiis Geofifroy these insects. Although my attraction to the (1762; Ghina 1962, International Commission Scolytidae was to their fundamental struc- on Zoological Nomenclature 1963). Since ture, diversity, ecology, and behavior from then, 488 generic and subgeneric names have theoretical and other points of view, as knowl- been proposed for the group, of which 215 are edge began to accumulate, my rural back- treated below as valid. ground and religious philosophy made me In an attempt to estimate the number of sensitive to the economic interests of others species remaining to be named from America and to a need for me to assist them with what I north of Mexico, White (1975), in his example had learned. Another need was also para- using Scolytidae, recognized 592 species in mount. Among the great frustrations in learn- 1970 and projected that there would be 650 ing are those resulting from errors that have named by the end-point year of 2040, when been transmitted from one generation to the 100% would be known. That estimate was next and, once learned, must be unlearned made prior to publication of my monograph and corrected before progress can continue. (Wood 1982). In the monograph, numerous Biological nomenclature and classification species were placed in synonymy and others have been profoundly impeded in their pro- were named or they extended their distribu- gress by this problem. Therefore, the primary tions into the United States and Ganada, leav- objectives of this study are (1) to review the ing about the same total number of species holotypes or syntypes of the type-species of all and projection as before. Meanwhile, more named genera and subgenera to make certain than half of the scolytid fiiuna of Mexico and that they are correctly placed in appropriate Gentral America was named from 1960 to genera, then find the correct generic names 1982. This rapid expansion of the fauna south for them, and review as many species as possi- of the United States made a projection for the ble and group them in those genera, and (2) to larger area meaningless at the present time. devise a scheme of classification such that oth- Few attempts to classify the higher cate- ers can identify those genera. If these objec- gories of Scolytidae on a worldwide basis have tives are achieved in at least some measure, it been published. Perhaps the first contribu- is hoped that the constant name changing and tion worthy of note was that of Ferrari (1867), duplication of effort (resulting from syn- who listed and described the genera of Tomi- onymy) that has impeded taxonomic progress cides (my Scolytinae with deleted). in my generation will be reduced to a mini- There was no attempt to actually classify the mum. This classification of categories above genera by Ferrari. Eichhoff (1878a) published the species level will be the framework on a classical monograph of this same group in which the catalog of species, now in prepara- which he presented keys and descriptions to tion, will be organized. all known genera and species. His family 1986 WOOD: Genera of Scolytidae

Tomicini included 40 genera that were dis- genera that are currently distributed (below) tributed among the subfomiUes Crypturgidae among the tribes , Dryocoetini, {Cn/pturgus, Dohirgus, Aphanarthruin, Trio- and . teranus, Pijcnarthrum), Cryphahdae (Li- Wood (1978) reviewed 15 major published parthntm, Hijpohorus, Triannoccnis, Cnj- classifications of the higher categories of phahis. Glyptodenia, Stephaiwdercs, Cos- Scolytidae, including all of those cited above, moderes), ProblechiUdae (Prohlechilus), Xy- and presented a revised and expanded classifi- loctonidae {Xijloctonus , Scohjtogenes), Pity- cation of subfamilies and tribes and tenta- ophthoridae (Pityophtlionis, Eidophidus, tively assigned 404 nominate genera and sub- Taphrorijchus), Tomicidae (Thamnurgtis, genera to his 25 tribes. That classification was Xijlocleptes, , Lepicenis, Drijocoe- adapted to his monograph of the North and tes), Hylocuridae (Hylociirits), Micracidae Central American Scolytidae (Wood 1982), (), Araptidae (Araptus), Hexacohdae which listed 1,4.33 species in 94 genera. The

{Hexacolus), Xyleboridae (, Xyle- present contribution is an expansion of that horus, Premnobiiis, Gnathotrichiis), Xylote- classification to the genus level for the entire ridae (Trypodendroti), Corthyhdae (Corfhy- world. lus, Brachyspartus, Anchonoceriis, Phtho- rius, Trypocranus, Pterocyclon), and Amphi- cranidae {Steganocranus, Amphicranus). Review of Characters Hagedorn (1910a, b) treated 115 genera and Characters Shared by Platypodidae 1,234 species in the entire family worldwide. and Scolytidae He divided Scolytidae (Ipidae, in his usage) into four non-Linnean subfamilies, based on In the early stages of this study, 1 was thor- the detailed structure of the mouthparts, that oughly convinced that Platypodidae could be contained the tribes indicated: Pilidentatae no more than a well-marked, primitive sub- (Phloeotrupinae), Spinidentatae (Eccoptogas- family of Scolytidae. Shared characters that trinae, Diamerinae, Crypturginae, Hylocuri- led to this premature conclusion included: (1) nae, Hylesininae, Ipinae, Cryphalinae), Sae- clearly formed pregular sutures extend from tidentatae (Xyleborinae, Corthylinae), and the anterior end of the median gular suture to Mixodentatae (Spongicerinae). or near the anterior tentorial pits; these su- Hopkins (1915a, b), in two papers, (a) re- tures are reinforced internally by a massive classified the subfamilies and tribes of Scolyti- internal inflection of the cuticle that occurs dae and (b) reclassified the genera and species nowhere else in the Curculionoidea; (2) the of Cryphalinae. He elevated the group to su- mandible lacks the horizontal cylinder that perfamily rank, Scolytoidea, and recognized rotates on the anterior and (heavily rein- four families within it (Ipidae, Scolytidae, forced) posterior condyles, with its cutting

Scolytoplatypodidae, and Platypodidae). His edge apical; instead, the posterior condyle is Ipidae included the subfamilies Cryphalinae, in a different position, the axis of the

Ipinae, Micracinae, Webbinae, Xyloctoni- mandibular hinge is transverse, and the cut- nae, Crypturginae, Phloeotribinae, Hylesini- ting edge is mesal (as in Anthribidae and nae, and Phloeoborinae. His Scolytidae in- Nemonychidae); (3) the rostrum is short to cluded the subfamilies Coptonotinae, Hexa- nonexistent; (4) the tibiae totally lack corbels, colinae, Bothrosterninae, Camptocerinae, but they share a common basic structure and and Scolytinae. His Scolytoplatypodidae in- are armed on the apical and lateral margins by cluded the subfamily Scolytoplatypodinae, a characteristic series of spines; (5) the eyes and Platypodidae the subfamilies Platypodi- are essentially flat against the head; (6) the nae, Genyocerinae, and Chapuisinae. It ap- antennae are mostly of the same basic genicu- pears to have been the intention of Hopkins to late type, with some members of several follow his basic classification with a series of primitive tribes having a short scape (little if papers treating in detail the various divisions any longer than the pedicel); (7) the visible of his classification, but only one paper was abdominal sterna 1 and 2 are weakly connate, published. Hopkins (1915a) reviewed the sub- except all are free in all primitive genera of family Cryphalinae, in which he included 48 Platypodidae; (8) the elytral interstriae 10 ex- Great Basin Natuiulist Memoirs No. 10

Fig. 2. Cephalic aspect of the head of a female Chaetas- tiis tuberculatus (Chapuis) (Platypodidae) showing a con-

Fig. 1. Outline of cephalic aspect of the head of Pst?urfo- spicuous "labrum ' in the median area just below the hylesinus sericeus (Mannerheim), with frontal rectangle epistomal margin arrow. (Re-drawn from Strohmeyer indicated by the square of dash lines. (1920: 15, Fig. 5).

tends to near the apex in all Platypodidae and bly homologous to it (Fig. 1). In Platypodidae in some primitive representatives of virtually the epistomal lobe is more generally present, all tribes of Scolytidae; (9) the larval characters and in Tesserocrerus and Chaetastus (Fig. 2) a are basically the same except as noted below; suture separating this lobe from the epistoma and (10) they share the same basic ecological is evident in some species. Comparable struc- niche. tures are unknown in the remainder of the Curculionoidea. Characters Not Shared with In general, have two gular sutures Other Curculionoids that are separated by a median gular sclerite. Curculionoidea, excluding Anthribidae, In Nemonychidae (Fig. 3) and some Belidae Nemonychidae, Platypodidae, and Scolyti- (Fig. 3B, and Crowson 1967) two gular sutures dae, share a unique mandibular character in are clearly indicated from the margin of the which the basal half forms a cylinder that ro- foramen magnum to the point of invagination tates on a horizontal axis between the two of the tentorium (posterior tentorial pit) articulating condyles (Fig. 8). The hypostomal where they end. In Anthribidae, the entire area adjacent to the posterior condyle is en- tentorial structure is lost and the gular sutures larged and strengthened to accommodate a are usually reduced to little more than lateral new mode of action; the cutting edge is apical irregularities on the margin of the foramen and usually takes the form of a large cusp on magnum. In Belidae there is variation from the side of the basal cylinder. In the four two widely separated gular sutures that end in families identified above, the mandibular two separate posterior tentorial pits (Fig. 3B) condyles are closer together, the oblique ac- to convergence to a single median pit, with tion is more like that of a hinge, the cutting consequent reduction in size of the gula, to edge is mesal, the mandible ends in an acute formation of a single median gular suture of point, and the hypostomal area is minimal. variable length (Fig. 3C). The allies of Belidae The Anthribidae and Nemonychidae have (Aglycyderidae and Oxycorynidae including an adult labrum. It is lost in all other Cur- Rhopalotria ) share at least part of this same culionoidea; however, in most primitive variability. The only other known members of tribes of Scolytidae (most Hylesininae, Cten- the Curculionoidea that share in similar vari- ophorini, etc.) there is an epistomal lobe that ability are members of the neotropical scoly-

resembles a small, fused labrum and is proba- 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 point of antennal articulation; they are not me, 5 had two complete gular sutures that represented from the area of antennal articu- extended to and were continuous with the lation to the anterior end of the median gular pregular sutures, 8 had the pregula elongated suture (the point marked externally where the and extending most of the distance to the tentorial apparatus invaginates). The pregular postgula, and 10 had the pregula more or less sutures illustrated by some writers on Cur- normal. Most Platypodidae (Fig. 4) have an culionidae (Hopkins 1911:Fig. 1) do not exist; unusually large pregula and a short to very they represent irregular undulations in the short median gular suture. cuticle that serve to strengthen the posterior Pregular sutures are totally absent in all (or ventral) wall of the rostrum and do not curculionoids, except for Platypodidae (Fig. qualify as sutures in any acceptable usage of 4) and Scolytidae (Fig. 5-G) and for their par- that term. tial presence in females of two (unidentified) Mouthparts have been used extensively in Australian Belidae (Fig. 3C). In these two fundamental divisions of the Curculionoidea belids the pregular sutures extend only from (Crowson 1967). For example, in Anthribidae near the anterior tentorial pits to near the and Nemonychidae the maxillary lacinea and Great Basin Naturalist Memoirs No. 10

tibia. Similar structure occurs in Histeridae, Bostrichidae, Brenthidae, or other coleopter- ous groups having a long history of occupancy of tunnels bored in wood. The tibial spines of Araucarini () 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).

Fig. 4. Posterior aspect ol head of a female ofDoliopijgus Visible abdominal sterna 1 and 2 are en- chapuisi (Duvivier) (Platypodidae). Note the very short tirely free in all primitive genera of Platypodi- median gular suture and the prominent pregular sutures. dae but are weakly connate in the higher Platypodidae, all Scolytidae, and in most higher Curculionoidea. galea form separate elements (Ting 1936: Fig. The larvae present an enigma. E,\cept in 78). The only other curculionoids sharing this the most primitive genera, the frons and character are the Tesserocerini (Platypodidae) clypeus are usually fused in Platypodidae (Fig. 6), Attelabidae, and Rhynchitidae (Ting (Browne 1972), as in Nemonychidae (Crow- 1936:Fig. 78). Similarly, the Anthribidae and son 1967), thus making them easily distin- Nemonychidae have a 4-segmented maxillary guishable from those of Scolytidae. However, palpus (Crowson 1967, Ting 1936). This char- characters have not yet been found that distin- acter is shared by all Attelabidae (Crowson guish some Scolytidae from some Curculion- 1967, Ting 1936), some Rhynchitidae, and idae (Viedma 1963). The difficulty may come one Platypodidae (Austroplatypus) (Browne more from reduction and simplification to ac- 1971a). commodate small size than from real differ- The costate lateral margins of the pronotum ences. Browne (1972) reported urogomphi- in primitive tribes of Scolytidae (Diamerini, like structures in two species of Platypodidae; Ctenophorini, Scolytini, etc.) is another indi- if correct, this is the only known occurrence of cation of primitive origin of this family within these structures in the Curculionoidea. the Curculionoidea. The tibial structure of Platypodidae and Discussion Scolytidae is unique. It appears to have been derived from a structure resembling that of Scolytidae and Platypodidae as Families Protohylastes (Fig. 7), in which the median member of three apical spines became the A position within the Curculionoidea is uni- terminal mucro of Platypodidae and the mesal versally accepted for Platypodidae and spine became the terminal mucro in Scolyti- Scolytidae, except for Schedl (1939), who gave

dae (Wood 1973a). The lateral margin is them superfamily status without explanation.

armed by one or more spines in primitive For more than a century, it was traditional to tribes; these may be replaced in specialized list Platypodidae and Scolyltidae as separate groups by socketed denticles (teeth) of setal families next to Curculionidae until Crowson origin (Wood 1978). The tibial structure of (1967:164) combined both families with Cur- and Camptocerus is a specialized de- culionidae, primarily on the basis of the ab- viation from the basic scolytid structure that sence of convenient larval characters that only superficially resembles the cossonine could separate them from that family. How- 1986 WOOD: Genera of Scolytidae

Stenoscelis

Fig. 5 A-F. Head structure of Curculionidae (Steno.icelis , ) and Scolytidae (Hijlurgops. ): 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 best, and warrants much closer examination. the comparative anatomy of the CurcuHon- Platypodidae and Scolytidae universally share oidea, I find that position untenable and sug- with one another; (1) well-developed pregular gest that the relationship of Platypodidae and sutures that are reinforced internally by mas-

Scolytidae to Curculionidae is remote, at 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 ments (Fig. 6); elsewhere in the Cur- the anterior end of the median gular suture) to culionoidea this character is shared by the or near the anterior tentorial pit adjacent to same four families (cited here in No. I). (3) An the anterior articulation of the mandible adult labrum within the Curculionoidea oc-

(Figs. 4-5), and (2) a primitive mandible that is curs only in Anthribidae and Nemonychidae; articulated and functions (about as in An- however, an epistomal lobe resembling a thribidae and Nemonychidae) in a manner fused labrum is widely represented among entirely different from that of higher cur- primitive Scolytidae (Fig. 1) and is usually culionoids (Fig. 8). present in Platypodidae (in Tesserocerus and

In view of these universally present primi- Chaetastiis , Fig. 2, it is even separated from tive characters, relict retention of other ances- the epistoma by a suture). (4) A complete gula, tral features in one or more genera of these with two gular sutures continuous with the two families takes on added significance. For pregular sutures, is present in at least five example: (1) The platypodid genus Austro- Gnathotriipes (Scolytidae) species (Wood platypus clearly has a 4-segmented maxillary 1973a), and the pregula is greatly prolonged in palpus (Browne 1971a) (known elsewhere only eight other species of this genus; the pregula in Anthribidae, Nemonychidae, Attelabidae, is greatly enlarged and the median gular su- and some Rhynchitidae). (2) The platypodid ture is short to very short in most Platypodi- tribe Tesserocerini has the maxilla clearly di- dae (Fig. 4). (5) In most Platypodidae and vided into separate lacinear and galear ele- many scolytid , Scolytini, Mi- ,

10 Great Basin Naturalist Memoirs No. 10

Sphenophormi Belidae Brenthidae

\

Fig. 8. Lateral a.spect of ape.\ of rostrum ot a representative of: Belidae, Brenthidae, Sphenophorini (Curculionidae), (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 pendent origin of this feature is probable. (9) any longer than the pedicel and definitely not The platypodid-scolytid body habitus most geniculate, and in other groups (some Phloeo- certainly is not of the Curculionidae type, tribtis, some Micracini, etc.) the club is although there is superficial resemblance to it poorly formed; very little imagination is in the Hylastini. needed to see the possibility of independent Family Scolytidae origin of the geniculate, clubbed antenna from that of other curculionoids. (6) The In the early stages of this study, I was thor- costate lateral margins and concave pleura of oughly convinced that Platypodidae could be scolyiid Diamerini, Ctenophorini, and some no more than a well-marked, primitive sub- Scolytini occur elsewhere in the Curculion- family of Scolytidae. In view of the often in- oidea only among the most primitive families. finitesimally minute features used to charac-

(7) The platypodid-scolytid tibiae totally lack terize other femilies, that position must be corbels. Their lateral margins bear true spines reexamined. Although most characters are in Platypodidae and some primitive Scolyti- shared by some members of each group, a dae. These are replaced in all higher Scolyti- sharp demarkation remains. For example, the dae by socketed denticles of setal origin that male spiculum gastrale is undeveloped in occur nowhere else in the Curculionoidea Platypodidae, but it is well-developed in

(Wood 1978); true spines also occur in some Scolytidae (Wood 1982); tarsal segment 1 is

Histeridae, Bostrichidae, and other families greatly elongated in all Platypodidae (Fig. 9), that frequent the tunnels of woodboring in- except for Protoplatijpus and Scohjtotarsus sects. (8) Although the literature states that but it is intermediate in Protohylastes and visible abdominal sterna 1 and 2 are connate Coptonotus (These four genera contain a total in all Platypodidae, this is not true. These of five rare, tropical species). The shape and segments are entirely free in all primitive gen- structure of the head, eye, antenna, prono- era; they are weakly connate in the higher tum, scutellum, elytra, and tibiae are almost Platypodidae and in all Scolytidae. An inde- 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- Fig. 10. Lateral margin near apex of protibia of Pohjgra- cally affected their morphology; this habit ap- phus rufipennis (Kirby). Note the socketed denticles. pears to have arisen independently in at least eight tribes of Scolytidae (Hyorrhynchini, Subfamilies and Tribes , Scolytini, Scolytoplatypodini, A review of characters usable in classifica- Xyloterini, Xyleborini, Cryphalini, Corthy- tion of the higher categories of Scolytidae was hni) (Wood 1982). presented by Wood (1978). Several trends The Platypodidae universally lack socketed were reported in that study: (1) the primitive tibial denticles (derived from setae) (Figs. 7, head is more or less truncate on its posterior 9); these structures are present in all higher face, the dorsomedian area is progressively groups of Scolytidae (Fig. 10) but are lacking prolonged caudad in specialized groups (Figs. in at least some genera of several primitive 13-14, and Wood 1982:Fig. 14); (2) the primi- tribes. In all primitive genera of Platypodidae tive frons is convex in both sexes (in the higher visible abdominal segments (sterna) 1 and 2 Hylesininae the male frons is variously im- are free; in the higher Platypodidae and in all pressed and the female frons is usually con- Scolytidae they are weakly connate. The pos- vex); the reverse is usually found in the terior half of elytral interstriae 10 is univer- Scolytinae; (3) the primitive eye is oval, en- sally present in Platypodidae; it is present in tire, and finely faceted; specializations in- primitive of tribes of Scolyti- members most clude elongation, emargination (including dae (Fig. 12), but it is lost in the higher mem- complete division into two parts) (Fig. 15), bers of almost all tribes (Fig. apical 11). The and enlargement of facets (apparently corre- protibial mucro of Platypodidae appears to lated with nocturnal flight habits); (4) the an- have been derived from the middle apical tennal scape primitively may have been short, spine of a tibia that of resembling Protohy- little longer than the pedicel; it is elongate or lastes (Wood 1973a); it appears to have been triangular in most groups; (5) the antennal derived from the inner (mesal) spine in funicle primitively contains seven segments,

Scolytidae (Fig. 7). In larval Platypodidae (ex- and there is a more or less orderly reduction to cept Protoplatypus and Schedlarius) the a minimum of one segment as specialization clypeus is fused to the frons (Browne 1972); in increases; (6) the antennal club varies from Scolytidae it is a separate sclerite. The list almost nonexistent (three movable segments could go on, but the above should indicate a no longer than those of the funicle in primitive close relationship between the two families ) to a simple cone-shaped struc- and the limited overlap of many characters. ture with transverse sutures to large and 12 Great Basin Naturalist Memoirs No, 10

Figs. 13-14. Lateral aspect of head in Scolytidae: 13, nigrinus Mannerheim, with the posterior face approximately truncate; 14, Xylosandnis retusus (Eich- Figs. 11-12. Lateral aspect of Scolytidae: 11, above, in- hoff), with the occipital area greatly extended. 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 the basal margins of the elytra tend to be Scolytodes pluineriae Wood. (8) costate primitively, with the Hylesininae be- coming procurved and crenulate (Fig. 16), the Strongly flattened or obliquely truncate, with Scolytinae transversely straight and longitudi- or without sutures; it is probably the most nally rounded (Fig. 17); (9) several complex variable major structure found throughout the changes occur in the meso- and metathorax family; (7) the prothorax may have (a) the that will be discussed below; (10) the male coxae widely separated in primitive groups to tergum 8 is visible and pubescent in most fully contiguous in specialized ones; (b) the groups (Fig. 22), but it is telescoped beneath 7 pleuron concave, with the lateral margins and without pubescence in Carphodicticini, acutely costate in primitive genera (Figs. 11- , Dryocoetini, Xyloterini, and Xylebor- 12), to convex, with the lateral margin un- ini; (11) the venter of the abdomen has (a) marked in specialized genera; (c) the prono- segments 3 and 4 (visible segments 1 and 2) tum longitudinally straight in dorsal profile free in all primitive Platypodidae but weakly and unarmed by crenulations in primitive connate in the higher Platypodidae and all groups to strongly arched and armed by Scolytidae and (b) limited specialized groups crenulations or asperities in specialized ones; 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 { 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 or tribe in which they are involved. However, tibiae are unusually variable and require special two of them are sufficiently complicated and im- treatment below. portant that elaboration is appropriate. These Most of the above character states are utilized occur on the metathorax and the tibiae. in the following key to subfamilies and tribes to Three primary characters of the metathorax indicate phyletic trends in the family, subfamily show important features that exhibit phyletic 14 GreatBasin Naturalist Memoirs No. 10

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 male 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, femole meeting the marginal crest of the groove only Fig. 22. Abdominal terga of Cryphohis nificollis Hop- briefly, if at all (Fig. 23, part 41). kins: male, with segments 7 and 8 visible; female, with

segment 7 visible (segment 8 is reduced in size, tele- The third significant metathoracic feature scoped beneath 7, and hidden from view). that is usable in phylogeny occurs only in the Hylesininae. A significant feature of insect trends. First, in Curculionoidea generally, in- flight is the intersegmental line between the cluding Platypodidae and primitive Scolyti- metathorax and its postnotum (derived primi- dae, the pleural suture follows a zigzag course tively from the anterior portion of the first from the pleural wing process ventrad to the abdominal segment but functionally part of point where the costal margin of the elytron the thorax), which must flex with each stroke touches the body. It then turns abruptly cau- of the metathoracic wings. This suture is dad to a point just before reaching the poste- present in all insects, including Platypodidae,

rior limits of the segment, where it turns Scolytinae, and primitive Hylesininae (Fig. mesad and continues to the pleural coxal pro- 23, parts 43, 45). In the more advanced cess. On the metepisternum at the anterior or Hylesininae, the median two-thirds of this first angle (below the pleural wing process) a intersegmental line is lost by complete fusion

carina or small, flattened spine is present that of the postnotum to the metathorax (Fig. 23, fits into a small groove on the inner (costal) part 41). This fusion is progressive, leaving a margin of the elytron just behind the humeral weak suture primitively, but it is totally oblit-

angle. This character state within Scolytidae is erated in advanced groups of this subfamily. best seen in the Hylastini (Fig. 23, parts 43- The tibiae exhibit remarkable variation in 44) and in some Hylesinini. The trend is for 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- derived resembles that of Frotohijlastes (Fig. ently, the primitive model from which the 7). The protibia of Protohylastes is slender, tibiae of Platypodidae and Scolytidae were 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- mes. Remnants of pregular sutures found in mediately basad from the tarsal insertion. females of two species of Belidae extend from Comparative studies (Wood 1973, 1978) sug- the anterior tentorial pit toward the antennal gest that the mesal (inner) apical spine lie- insertion and never converge or approach the came the apical spine in Scolytidae and that gular suture (or sutures). (2) In these two the middle spine became the apical spine in families, the mandibles come to an apical Platypodidae (Figs. 7, 9). This middle spine in point and have the cutting edge mesal as in

Scolytidae is bent laterad and is a major, iden- Nemonychidae; furthermore, the articulating tifiable element that projects beyond the level condyles are comparatively close together, of the tarsal insertion in one or more members with the posterior one more nearly lateral in of several tribes (Hyorrhynchini, Diamerini, position, and the hypostomal area behind the

Bothrosternini, Phloeotribini, Phloeosinini, posterior condyle is minimal in size to almost Hypoborini, Polygraphini, Scolytini, Cten- nonexistent. In most Curculionoidea, except ophorini, Scolytoplatypodini). All tibial spines for these two families and Nemonychidae and in Platypodidae (Fig. 9), Hyorrhynchini, Anthribidae, the basal half of the mandible Scolytini (Fig. 7), and, apparently, Cten- forms a horizontal cylinder that rotates on the ophorini (Fig. 7) are true spines. In all higher two condyles, and the cutting edge of the

Scolytidae these spines are either intermi.xed mandible is apical; the posterior condyle is with or replaced by socketed denticles of setal posterior in position and is supported by an origin (Fig. 10). The patterns of possession enlarged hypostomal area. (3) The subcortical and replacement within the family suggest habit in which mated pairs enter vital host that socketed denticles had an early mono- tissues to oviposit is shared by Platypodidae, phyletic origin within the Scolytidae, with dif- Scolytidae, and some Cossoninae (Curculion- ferential survival of this character in the pos- idae); however, the mode of excavation and terity, and do not suggest polyphyletic several structural features (gular area, tento- origins. rium, mandible, etc.) indicate that the latter Many other characters that are not men- group exhibits an evolutionary parallelism, tioned above are useful in classification and not a fundamental relationship. (4) Six genera phylogeny in more limited groups. Several of {Protohylastes, Coptonotus, Scolytotarsus, these are employed in the keys. Protoplafyptis, Mecopelmus, Schedlarius) are structurally and biologically intermediate be- Conclusions tween the Platypodidae and Scolytidae and As stated above, the traditional place of could be placed in either family. The charac- Scolytidae in classification has been as a fomily ters, reviewed in this paragraph, with primary next to the Curculionidae. However, as Cur- stress placed upon true pregular sutures, indi- culionidae became fragmented into several cate a fundamental close relationship between families, the position of Scolytidae within the Platypodidae and Scolytidae that is not shared Curculionoidea changed. Schedl (1939c) with other groups. made a separate superfamily to contain Platy- A second point of significance has to do with podidae and Scolytidae. Crowson (1967:164) the position of Platypodidae and Scolytidae reduced both to the rank of subfamily within within the Curculionoidea. Their traditional Curculionidae. position in classification has been next to the In reviewing this topic, our first point has Cvnculionidae; however, close examination been to establish that Platypodidae and suggests that such a placement is based on Scolytidae are closely related to each other superficial resemblance, not on fundamental and are derived from the same parental stock. structure. As indicated above, pregular su- Evidence for this, as stated above, comes from tures are shared, in part, only with females of the following: (1) Pregular sutures that extend two species of Belidae, and the mandibular- from the anterior tentorial pit (adjacent to the hypostomal structure is shared only with An- anterior articulation of the mandible) to meet thribidae and Nemonychidae. These indica- the anterior end of the median gular sutiue tions suggest a relationship of Platypodidae occur only in these two groups. These sutures and Scolytidae to the segment of Curculion- are reinforced internally by massive apode- oidea having two gular sutures, rather than 1986 WooD: Genera of Scolytidae 17 one. However, close inspection of Belidae sublamellate, with the three segments freely discloses at least three genera with only one movable upon one another (Fig. 16). In primi-

median gnlar suture and an elongated post- tive members of this genus iacaciae , rhodo- gula, and an entire segment of Oxycorynidae dacUjlus , etc.) the three segments are no (Rhopalotria) with one gular suture and a longer or wider than are the funicular seg- postgula of variable size. Furthermore, one ments and are as movable. I see no possibility genus of Scolytidae {Gnathot rapes) contains that the Phloeotrihus antenna could have at least five species with a complete gula con- been derived from a solid club of the type necting pregular and postgular elements, and found in most other Scolytidae. Since this is at least eight more in which the connection is the only deviant character found in this genus, almost complete. In many Platypodidae, the could it not be a relict reminder that the pregula is enlarged and is scarcely separated scolytid club was derived on an independent, from the postgula. Because such a situation parallel line from a common ancestor, and not does not occur elsewhere among those cur- Irom other curculionoid groups having culionoids having one gular suture, wisdom clubbed antennae? suggests that a search be continued for addi- Much has been said and written of the simi- tional clues to phyletic relationships. larity of the tibiae between Scolytiis (Scolyti- A movable labrum occurs among cur- dae) and many Cossoninae (Curculionidae) in culionoids only in Anthribidae and Nemonychi- curculionoid family classifications. However, dae. Remnants of a labrum are found in many it has never been pointed out that similar primitive Scolytidae (particularly Hylesininae), tibiae occur in Histeridae, Brenthidae, and and a fused labrum, complete with a transverse other coleopterous families containing groups suture, occurs in Platypodidae (especially Tesse- with a long history of subcortical existence. rocerini). 1 am unaware of similar remnants in Within the Scolytidae, the Scolytus tibia is other curculionoid families. unicjue and apparently does not represent a A 4-segmented ma.xillary palpus occurs in truly primitive condition. The truly primitive Aiistraloplatypiis (Browne 1971a:49), a fea- character state of the tibia on this phyletic line ture restricted to only the most primitive cur- occurs in Protohijlastes, with Platypodidae culionoids (see family key below). and Scolytidae derived along separate phylet- A lacinia separate from the galea is charac- ic lines from the basic structure.

teristic of all Tesserocerini (Platypodidae); Larval characters, thus far, have not been this is another feature that is restricted to the very helpful in answering questions about most primitive curculionoids (see family key scolytid phylogeny. In fact, reliable means below). have not yet been found for separating all The antenna has been used extensively in Scolytidae from all Curculionidae (Viedma the classification of curculionoid families, par- 1963, Lekander 1968). Whether this is due to ticularly character states of orthocerous vs. the absence of characters or to the lack of geniculate, apex filiform vs. clubbed (or capi- diligence in the search for characters among

tate), etc. In no curculionoid are segments 1 groups not found in Europe needs to be an- and 2 without some degree of enlargement; swered. Primitive larval Platypodidae have furthermore, in many of those traditionally the frontal sutures extending to the articulat- referred to as geniculate, the scape is not ing membrane of the mandibles, a fact not

longer than it is in some Anthribidae. In a previously noted. Browne (1972) suggested large segment of Platypodidae and many that structures found on two platypodid larvae

Scolytidae, the scape is short, little if any could be relicts of urogomphi. If so, this is the

longer than the pedicel, and often it is broadly only evidence of the existence of urogomphi triangular in shape. In virtually all cur- in the Curculionoidea. Larval Scolytidae and culionoids (except most Anthribidae), the Platypodidae are easily separated from each three apical segments of the antenna are at another by the separate (Scolytidae) or fused least slightly enlarged; although the literature clypeus and frons (Platypodidae). It is of inter- generally states otherwise, a few Anthribidae est that this same character separates larval have an abruptly enlarged club. In the genus Nemonychidae and Anthribidae (see family

Phloeotrihus (Scolytidae), the club is usually kev below). ,

18 Great Basin Naturalist Memoirs No. 10

Lower Jurassic curculionoids with a long tives. Rostrum length among Jurassic cur- rostrum have been referred to the CurcuHon- culionoids was diverse, varying from short idae by Crowson (1983), although they bear a and broad (Arnoldi 1977) to long and slender very strong resemblance to Belidae or, per- (Crowson 1983). The presence of obvious, so- haps, Eobelidae. Arnoldi (1977) named the phisticated scolytid tunnels in Lower and extinct family Eobelidae from 14 species that Middle Cretaceous bark suggests that were placed in 7 tribes in 4 subfamilies, taken the Platypodidae-Scolytidae phyletic line had from Upper Jurassic deposits of South Kaza- its origin at least in Upper or Middle Jurassic. khstan (Karastan) in the USSR. In this family The exact phyletic positions of known Jurassic the head is similar to modern Belidae except and Cretaceous "curculionid" fossils are diffi- that it is more broadly oval and the rostrum is cult to determine, because details of labrum, shorter, as in some modern male Brenthidae. mandibles, gular sutures, etc., are not suffi- The mandibles are large, with a single apical ciently well preserved to permit analysis. I point and a mesal cutting edge as in modern seriously doubt that any true Curculionidae Nemonychidae. The antennae attach either to existed prior to the Cretaceous. the middle or near the apex of the rostrum and are orthocerous, with the three apical seg- Habits and Classification ments slightly enlarged but not forming a defi- Although the Platypodidae are established nite club. Arnoldi did not mention gular su- above as a family, separate from Scolytidae, tures or a labrum. The body resembled they are significant because they represent modern Belidae except that the lateral mar- the first major paraphyletic branch from the gins of the prothorax were acutely elevated as platypodid-scolytid line of specialization. in modern Oxycorinidae and the elytra were Their adoption of the xylomycetophagous much less strongly sclerotized than in modern habit appears to have accelerated their struc- Belidae. tural and biological deviation. Only the inter- From the above, two fundamental conclu- mediate genera Protoplati/pus, Mecopelmus sions emerge. First, the Platypodidae and and Schedlarins lack the ambrosial habit {Pro- Scolytidae are very closely related to one an- tohylastes, Scohjtotarsiis , and Coptonotus other and, in fact, they intergrade to a limited are unknown biologically). Schedlarius (mon- extent. At the same time, these two families ogamous) and Protoplatypus (polygynous) are distantly related to families in quite other place their eggs in sealed niches; all other this superfamily. Second, in spite of the ap- known platypodids are monogamous and dis- parent absence of larval characters, there are tribute the eggs loosely in the parental gal- enough discordant adult characters to conclu- leries or carry them on the female frons. sively demonstrate that the Platypodidae- Mecopelmus makes a cave-type nuptial cham- Scolytidae, as one unit, do not belong to and ber (without ovipositiou galleries) in the cam- probably are not even closely allied to Cur- bium region of its host and deposits the eggs in culionidae. It is my contention that their preg- clusters therein, and the larvae form indepen- ular sutures (and accompanying internal dent mines in the cambium at least for the apodemes) prevented elongation of the ros- latter part of their development. Protoplaty- trum and forced them into an entirely differ- pus forms stellate or radiate tunnels, with a ent mode of existence from other cur- central nuptial chamber in the cambium re- culionoids (parent adults had to bore into vital gion, and the eggs are placed in regular, alter- host tissues to oviposit because they had no nate, sealed niches along the linear egg gal- rostrum and, consequently, could not follow leries; the larvae form independent mines the curculionid habit of using the rostrum to throughout their development. Schedlarius is form ovipositiou pits). They could have been xylophagous (Wood 1957a), places its eggs in derived in the Jurassic from a short-rostrumed randomly organized niches that are sealed Eobelidae-like ancestor that had fully formed with frass, and the larvae form long, indepen- pregular sutures. Belidae and related dent mines that wander aimlessly through the families, and all other curculionoids having wood. An obvious association with fungi is one gular suture, branched off later or from apparent in the vicinity of adult and larval other segments of Eobelidae or its deriva- activity, although no mycelial growth in the .,

1986 WOOD: Genera of Scolytidae 19

mines could be seen at magnification of 20X. pact (stout) body; (3) the body compaction that The habits mentioned in this paragraph also appears to have resulted in the development form the basis of habits for the more diverse of a row of coarse crenulations on the basal

Scolytidae. margins of the elytra (their function is evi- In my review of characters usable in distin- dently to add resistance when a predator at- guishing subfamilies of Scolytidae (Wood tempts to pull from a timnel the that is 1978), only one significant character gap was blocking the entrance), and (4) the straighten- found that was supported by numerous fea- ing of the pleural suture. The compaction of tures. These characters are summarized in the the body apparently also had an effect upon key to tribes below. The presence or absence the mechanism of flight and permitted fusion of pronotal asperities, the ambrosial habit, of postnotum 2 to the metathorax. Although and several features used traditionally by past use of these characters recjuires further study workers to separate multiple subfamilies of and refinement, a definite trend from the Hy- Scolytidae either have no value in characteriz- lastini-Hylesinini to Phloeosinini is indicated. ing subfamilies or have no value in phylogeny There appears to be a gradual transition in For these reasons, only two subfamilies were characters from the Hylastini to Hylesinini to recognized. Because the Hylesininae are . The Hyorrhynchini and Phrixoso- structurally and biologically less diverse and mini appear to be relicts of an earlier radiation their specializations more conservative, they of primitive stock and are quite unrelated to are treated here as more primitive than the the other three tribes. The remaining Scolytinae. hylesinine tribes have the postnotum partly to entirely fused to the metanotum and fall into

Tribes of Hylesininae three units. Tlie first includes the Old World Diamerini and New World Bothrosternini In general, the Hylesininae are phloeo- tiiat probably contain the most primitive spe- phagous. Exceptions include the twig- or cies of this subfamily. Their protibial, prono- small-branch (pith)-boring mycetophagous tal, and base of elytral structure is much more Hyorrhynchini (all three genera), Bothroster- primitive than the Hylastini-Hylesinini. nus (Bothrosternini), and Hylcops (Phloeo- Their antennal structure is much more similar sinini; last half of larval stage only); xyloph- to Coptonotiis-Protohylastes (Platypodidae), agous Dendrosimis and three known species although the Hylastini-Hylesinini club is of the large genus Chramestis (Phloeosinini); probably more similar to the ancestral struc- the myelophagous Cnesiniis, Eiipagiocerus ture. and Sternobothrus (Bothrosternini); and the The second tribal unit within the more spermophagous (Bothrosternini) highly evolved segment of this subfamily in- and one species of Phloeotrilnis (Phloeotrib- cludes the Phloeotribini and Phloeosinini. ini). The usual mating system in this subfamily Tibial structure of at least a few representa- is monogamy, with the female initiating the tives of each suggests an ancient origin. Ex- new parental gallery. Departures from this cept for the aberrant structure of the antennal system include (a) apparent male haploidy club of Phloeotribini, they probably could not (arrhenotocous parthenogenesis accompanied be separated from one another at the tribal by consanguineous polygyny) in Sueus (Hyor- level. They include species bearing the great- rhynchini); (b) monogamy, with male initia- est departures from the primitive forms in the tion of parental galleries in some Phloeosinus pleural and scutoscutellar suture patterns. and Olonthogaster (Phloeosinini); and (c) het- The absence of socketed protibial teeth in erosanguineous (superficially resembling Aricerus and the movable segments in the harem) polygyny in at least two species of New antennal club suggest that Phloeotrinini is the Guinea Olonthogaster (Phloeosinini), and in more primitive of these two tribes. all and many The Polygraphini and Hypoborini are obvi- (Polygraphini). ously specialized ancient groups that are un- These basic habits appear to be correlated related to one another but of uncertain affinity with (1) a conservative evolutionary departure to other groups. The superficial appearance of from the ancestral structure; (2) a more com- the body form of Phloeographtis (Poly- ,

20 Great Basin Naturalist Memoirs No. 10 graphini) suggests a relationship to Tomicus bigyny, etc. Xylomycetophagy and males nor- but the protibial structure of Serrastiis sug- mally associated with more than two females gests a much more ancient origin of this are unknown. Scalelike pronotal and elytral group. The Hypoborini could be aspecialized setae are a common occurrence. The evolu- paraphyletic line that branched from the tionary connection between this group of Phloeosinini. tribes and the more primitive Scolytini- Ctenophorini-Scolytoplatypodini is remote; Tribes of Scolytinae however, a connection to the more advanced

Xyloctonini-Cryphalini is clearly evident. The tribes of Scolytinae are easily clustered The third cluster of tribes within the into five major groups on the basis of several Scolytinae includes the Carphodicticini, Ip- anatomical characters. The most primitive, ini, Dryocoetini, Crypturgini, Xyloterini, and composed of Scolytini, Ctenophorini, and Xyleborini. In this group, male abdominal Scolytoplatypodini, share primitive protibial tergum 8 is reduced in size and telescoped structure without socketed denticles, a 6- or beneath 7, as in the female. In addition, the 7-segmented antennal funicle, a head without procoxae are almost always contiguous, the a caudal extension of the dorsal occipital area, occipital area of the head attains its greatest rather widely separated procoxae, etc. This prolongation, the obliquely truncate antennal group contains members (Cnemomjx, Scoly- club is developed and exploited, both het- todes) with the greatest structural similarity of erosanguineous and consanguineous polyg- all Scolytinae to primitive Platypodidae (Pro- yny are common, and xylomycetophagy is toplatypus , in this instance) and to Hylesini- universal in two tribes. Carphodicticini and nae. In fact, some Cnemomjx (galeritus and its some Dryocoetini are obviously very primi- allies) have coarse crenulations on the basal tive, but their connection to other primitive margins of the elytra; also, Protohtjiastes was Scolytinae is remote and not reflected in the thought to be allied to modern known fauna. The basal margins of (Tomicini) until the legs and antennae were the elytra are elevated and costate in Dendro- studied. The Ctenophorini habits include dicticus (Carphodicticini) as in some Hylesini- phloeophagy, xylophagi, and myelophagy; monogamy and polygyny (heterosanguineous nae. only); egg chambers of the cave, linear and The fourth cluster of tribes within the radiate types; and deposition of eggs in loose Scolytinae includes Xyloctonini and clusters in parental chambers or in individual Cryphalini. In these tribes the procoxae are niches sealed by frass. The monogamous, xy- entirely contiguous, and the antennal club is lomycetophagous Scolytoplatypodini appear flat, with the sutures on the posterior face to be a specialized Old World geographical moderately displaced toward the apex. The replacement of the Ctenophorini. The Scoly- small metepisternal spine that functions in tini are monogamous (except for a few bigy- locking the elytra in the closed position is nous Scohjtus) and phloeophagous (except for reduced and partly replaced by a small groove the xylomycetophagous Camptocerus). Cne- (Fig. 31). Segmentation of the antennal funi- momjx (Scolytini) and Scohjtodes (Ctenoph- cle is reduced, ranging from a maximum of orini) could have been derived from a com- seven to a minimum of three. Monogamy is mon ancestor. Pycnarthnim (Ctenophorini) the general habit, with consanguineous polyg- could easily be placed as a primitive tribe of yny universal in four genera of Cryphalini Hylesininae on both anatomical and biological {Cnjptocaremis, Hypothenemits, ,

bases. Periocnjphalus) . There appears to be a close The second cluster of tribes within the connection between Micracini and this group Scolytinae includes the Micracini and Cac- of tribes. The elytral locking mechanism also topinini. Advances include a more efficient suggests a remote connection between this body form (cylindrical compaction), slightly group and the Corthylini (Figs. 23, 31-32). extended dorsal occipital area of the head, The final cluster of tribes includes the Cor- reduction of funicular segments to six (rarely thylini. The flattened antennal club, with the five), universal occurrence of pronotal asperi- sutures equal on both sides, the unique elytral ties, common occurrence of xylophagy and locking mechanism, tibiae, eyes, elytra, etc.. .

1986 WOOD: Genera of Scolytidae 21

characterize this unique tribe. Pliloeophagy is have a pre-Tertiary origin (Table 1, column 2). almost universal in the primitive half of tiie If the tribe at that time was local in distribu- tribe, xylomycetophagy in the advanced half tion and was represented on only one land

Both monogamy and heterosanguineous poly- mass, it was not included as pre-Tertiary even gyny are common. Consanguineous polygyny though it could have been present. A faunal apparently occurs in three species oiArapttis connection from South America to Australia There may be a remote phylogenetic connec- was regarded as early Tertiary, not just pre- tion to the Cryphalini. Tertiary. It is noted (1) that 6 of 11 tribes of Hylesininae and 7 of 14 tribes of Scolytinae Geographical Origin of Tribes are judged on this basis to be of pre-Tertiary

origin, and (2) that the only pre-Tertiary For a discussion on this topic to have any group containing species with the xylomyce- semblance of objectivity, certain premises tophagous haliit is Xyleborini. (Hylesinini is must be established. First, it is assumed that excluded, because its only xylomycetopha- continental drift did occur and, due to that

gous genus, Hyleops , has not yet fully phenomenon. South America and Africa were adopted that habit.) This does not mean that either connected at their southern ends or other xylomycetophagous groups were not were at least close enough to permit faunal present, but only that, if they were present, exchange by island-hopping until earliest Ter- they were local in distribution (on only one tiary. At that time, Australia was close enough land mass) at that time. It is further noted (3) to the southern end of South America to per- that all tribes judged as pre-Tertiai-y were mit at least limited faunal exchange. Second, phloeophagous except for Xyleborini and part it is assumed that South America was an island of Micracini. This suggests that xylophagy and during most of the Tertiary. In addition to the xylomycetophagy are of comparatively recent assumed connection to Africa in earliest Ter- origin. tiary, South America was connected to North The significant point is that the family America in early Tertiary and again in late Scolytidae was a well-established, diversified Tertiary either by a land bridge, as presently group by the beginning of the Tertiary. The exists, or by a series of islands that were close fossil engravings of scolytids in coniferous enough to permit faunal exchange. Third, it is bark from Upper Cretaceous illustrated by assumed that faunal exchange between Africa, Brongniart (1877) and described from Lower Europe, and Asia has been no problem except Cretaceous by Blair (194.3) are evidence of a as it has been affected by climate. Fourth, it is much longer family history of the group than assumed that climate has fluctuated as evi- is usually recognized. It is probable that the denced by fossil remains of magnolia trees in beetles that made those engravings were Alaska and of tropical forests where the Sahara much more similar to the Ctenophorini or, Desert is now located. Finally, it is assumed possibly, Protoplatijpiis than to modern that scolytid beetles have had habits through- Hylesininae. out their history that were essentially as seen at present. That is, groups presently re- stricted to the tropical forests have always Phylogeny been so restricted, and groups now in more temperate climates have been able to occupy As stated above, the Platypodidae-Scolyti- cooler climates for a long time. dae group are members of the Cur-

In the first column of Table I are listed all culionoidea. Within that superfamily, they tribes of Scolytidae presently recognized in form a distinctive unit that is characterized by the world fauna. From each tribe, those gen- the presence of fully developed pregular su- era were selected that now are restricted to tures. Because the only other known pregular tropical climates. If those genera are now rep- sutures within the Curculionoidea are the api- resented in both Africa and South America or cal remnants found in females of two species of if they have a very closely related geographi- Belidae, the possibility of an ancient common cal replacement genus on both continents, ancestor should be examined. Three genera of but are not found outside the tropics else- Belidae and an entire segment of Oxyco- where, then those tribes were considered to 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. + 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 Ainerica Worldwide Corthylini South America Worldwide

(African segmerIt)

+ ) if in if at the beginning of the Marked ( present recogniziiljle form prior to the beginni ig of the Tertiary' on more than one land mass. Marked (0) represented

Tertiary by a local representative or a precursor that was presei It on only one land mass of Belidae, have only one gular suture, not curculionoids known to me. Anthribidae and two as in other members of the Behdae- Nemonychidae have a maxillary lacinia sepa-

Oxycorynidae-Aglycyderidae (Proterhinidae) rate from the galea; this character is shared by group. Because such a placeinent would be a Platypodidae (Tesserocerini), Attelabidae, radical departure from the traditional position and Rliynchitidae; the lacinia and galea are of Platypodidae-Scolytidae within the Cur- fused in all other curculionoids. Most of those culionoidea (which has been next to the Cur- curculionoids having two gular sutures have a culionidae), additional supporting evidence 4-segmented maxillary palpus; Attelabidae, was sought. The following summary of infor- some Rliynchitidae, and one Platypodidae mation presented in the above sections was share that character. In Anthribidae, found. A functional, but degenerate, adult Nemonychidae, Eobelidae (fossil), Scolyti- labrum occurs in Curculionoidea only in An- dae, and Platypodidae the mandible is api- thribidae and Nemonychidae. Rudiinents of a cally pointed, the cutting edge mesal, the labrum are well marked in many Platypodidae condyles are closer together and differently and obscurely indicated in some primitive positioned, and the apical part of the hypos-

Scolytidae, but they are not indicated in other toma is of minimal size. In all other cur- 1986 WOOD: Genera of Scolytidae 23 culionoids, the basal half of the mandible subspherical to take advantage of the row of forms a cylinder that rotates on a horizontal crenulations on the bases of the elytra; axis between the condyles, the dentate cut- scolytine evolved toward cylindrical com- ting edge is distal and moves through a differ- paction that is more effective in deeper host ent arc, and the hypostoma is enlarged to give tissues where the ambrosial habit was ex- greater strength to the posterior condyle. ploited. Whereas the preponderance of adult charac- Progess toward a more nearly spherical ters support the suggested relationship, larval body form in Hylesininae was accompanied characters have not been found that separate by straightening out (somewhat) the scu- some Curculionidae fi'om some Scolytidae. toscutellar and pleural sutures on the meta- The larvae of Platypodidae and Scolytidae are thorax. This apparently modified the mecha- easily separated; however, it is by the same nism of flight such that flection between the primary character that Anthribidae and metathorax and its postnotum was no longer larvae Nemonychidae are separated (see key necessary and that the intersegmental line to families below). Obviously the Platypodi- (membrane) was lost, at least in the median dae-Scolytidae are not part of the Curculion- area. Accompanying that change were; a re- idae and occupy a more primitive position in duction in the number of segments of the phylogeny. E.xactly where they branch from antennal funicle from seven to as few as three the main curculionoid stem is not yet clear, and changes in the antennal club from cone- but it could be from an Eobelidae-like ances- shaped, with simple, transverse sutures, to tor. flattened and with or without sutures in nu- Within the Platypodidae-Scolytidae, three merous combinations. The primitive mating major phyletic units are evident. smallest The system was monogamy, with the female ini- and most distinctive paraphyletic line is the tiating the new parental gallery. Limited de- Platypodidae (about 1,000 species). They lack partures from that system include male ini- socketed tibial denticles, have the terminal, tiated monogamy and heterosanguineous protibial mucro formed from the middle one polygyny (many Polygraphini) and consan- of three apical spines (as compared to Proto- guineous polygyny (Siieiis in Hyorrhynchini). hylastes), always have the posterior half of A recognizable phyletic sequence appears to elytral interstriae 10 present, always have a progress from the primitive Hylastini through primitive mechanism for locking the elytra in Hylesinini to Tomicini. Hyorrhynchini and closed position, lack a spiculum gastrale, and Phrixosomini appear to have been derived contain members having the primitive charac- independently on separate lines from the ters cited in the above paragraph. The Scolyti- same basic stock as that group of tribes. dae contain partial or complete departures Among those tribes with a fused postnotum, from these characters. As indicated above, si.x Bothrosternini and Diamerini appear to be rare tropical genera (containing a total of geographical replacements of one another seven species) are intermediate between that arose from a common ancestor that was these families. much more closely allied to the Ctenophorini- The other two major phyletic lines include like ancestor of Scolytinae than to the immedi- the Hylesininae (about 2,000 species) and the ate ancestral stock of the Hylastini- Scolytinae (about 4,000 species). Although Hylesinini-Tomicini. Although the Phloeo- the most primitive members of the family un- tribini and Phloeosinini are closely related to doubtedly are members of the Scolytinae, one another, they (as one unit) were probably that subfamily also contains far more and derived from the Bothrosternini-Diamerini greater specializations than do the Hylesini- ancestral stock independently from the unre- nae. Because the Hylesininae are less diverse lated Polygraphini and Hypoborini. Differen- and depart less drastically from the primitive tiation of phyletic lines in the Hylesininae has form, they are considered more primitive and progressed slowly; consequently, they are dif- paraphyletic; the Scolytinae are more special- ficult to detect. In general body habitus, Pro- ized and contain the major evolutionary thrust tohylastes (Platypodidae) resembles primitive of the family. Hylesinine evolution appears to hylesininae very closely, but the legs and an- have been oriented toward a bodytype that is tennae do not. Pycnariliritm (Ctenophorini) .

24 Great Basin Naturalist Memoirs No. 10 would be placed in Hylesininae except for its purpose of studying types and comparing legs. A primitive relationship of Hylesininae them and other specimens to my material at to primitive Scolytinae and Platypodidae is museums in the United States, Canada, Mex- clearly indicated in these genera. ico, Venezuela, England, Austria, Finland, Among the Scolytinae, the Ctenophorini, USSR, India, Japan, Australia, and Papua- Scolytini, and Scolytoplatypodini are closely New Guinea. There visits included stays of a related to one another and were almost cer- month or more at the British Museum (Natu- tainly derived from a common Ctenophorini- ral History), the Schedl Collection (both at like ancestral stock. The Ctenophorini more Lienz with Schedl in 1965 and at Vienna in nearly resemble Protoplatijpus (Platypodi- 1983), and the Forest Research Institute, dae) than do other members of this subfamily. Dehra Dun, India (where more than 40,000 Some Cnemomjx (allies of galeritus, Cten- Indian specimens were sorted to species in ophorini) have crenulations on the basal mar- addition to studies of the types). It is esti- gins of the elytra as in Hylesininae. The four mated that well over 400,000 specimens of remaining clusters of tribes were probably Scolytidae and Platypodidae were examined. derived from a Ctenophorini-like ancestor in From those museums not visited in person by the following pattern. Although there is no me, types and other specimens were obtained clear primitive connection to an ancestral for study through loan. Except as noted below group, Micracini and Catopinini are anatomi- under Incertae Sedis, all existing type-species cally and biologically allied to one another. An for all named genera and subgenera were ex- advanced member of Micracini probably gave amined and, where type-specimens have rise to the Xyloctonini-Cryphalini group, then been designated, the type-specimen of those a Cryphalini-like member gave rise to the species were also studied. The basis for recog-

Corthylini. The remaining tribes, Carphodic- nition is given under each species and syn- ticini, Ipini, Dryocoetini, Crypturgini, Xylo- onym below. terini, and Xyleborini (listed in ascending or- Except at Helsinki (Finland), Leningrad der of anatomical and biological complexity) and Moscow (USSR), Bulolo (Papua-New were probably derived from another Mi- Guinea), and Sydney (Australia), where local cracini-like ancestor along the same (one) museum equipment was used, the studies phyletic hue of descent. Active major evolu- were made using an American Optical Com- tionary thrusts appear to be in progress within pany stereoscopic microscope, model 25, the Dryocoetini, Xyleborini, Cryphalini, and equipped with an occular grid, at magnifica- Corthylini, which combined contain more tions of 10, 40, SO, and 160X. All measure- than half the living species of Scolytidae. ments were made at lOX and excluded the head (in measurements of body length). Methods Authentic specimens of the type-species of several genus-group names assigned to the

Although this project was conceived in Scolytidae were not examined during the course of this study. These species fall into 1949, serious work on it was not started until 1955. From 1955 to 1965, comparative two categories: (1) those for which the types or anatomical studies were made of representa- other authentic specimens could not be found tives of more than 100 selected genera of Cur- and are presumed lost, and (2) fossil species. culionoidea, in addition to more than 60 gen- The types of four genera could not be era of Scolytidae and Platypodidae. As the found. These included: patterns of evolution began to emerge from 1. Allarthnim Hagedorn (1912:355), based those studies, a major efltbrt was made to col- on A. kolbei Hagedorn (1912:355), from Pe- lect specimens and to gather behavioral and terhafen, Deutsch New Guinea. The type(s), ecological data on species not previously 1.5 mm in length, was deposited in the Kgl. known to me. More than 2,000 species of Zoolog. Museum Berlin but cannot now be Scolytidae and 300 species of Platypodidae found. The illustration of the antenna (Hage- were collected from North and South Amer- dorn 1912:Fig. 5) is of a CrypJiahis species. It ica, Europe, Asia, Australia, and New is provisionally listed as a synonym of Guinea. In addition, visits were made for the Cryphahis . . .

1986 WOOD; Genera of Scolytidae 25

2. Bufonus Eggers (1919:231), based on B. 2. Hijlescierites Schedl (1947:29) was based obscii?-iiS Eggers (1919:231) from Amani, Ost- on H. gramdatits Schedl (1947:.30) from Baltic Afrika. The unique holotype, 1.5 mm, was lost amber. From the photograph of the holotype with the Hamburg Museum in 1944. The an- and the original description, I see no reason tennal hjnicle was described as 2-segmented, for separating this species from Hijlurgopa. the club elongate, with three transverse su- 3. Taphramites Schedl (1947:41) was based tures; the scutellum was not visible, the basal on T. gnathotrichiis Schedl (1947:42) from margins of the elytra were sharply roimded, Baltic amber. From the description, it ap- and the elytral sculpture was pronounced and pears that this species should be placed in unique. Except for the absence of crenula- Dryocoetini, probably in or near Drijocoetes tions on the basal margins of the elytra, this I have not examined the specimen. genus would be placed in Hypoborini. A more 4. Xylechinites Hagedorn (1907:120) was detailed knowledge of the fauna of East Africa based on X. anceps Hagedorn (1907:120) from is needed before it can be correctly placed in Baltic amber and was redescribed and illus- classification. trated by Schedl (1947:30-32). The descrip- 3. Toxophorus Eggers (1920b; 119; preoc- tions and photograph of the holotype indicate cupied, renamed ro.vop/if/ior(/.s Wood), based that this species is a Tomicini near or in on T. africanus Eggers (1920b: 119) from Xylecliintts . I have not examined the speci- Deutsche-Ostafrika. The unique holotype, men. 3.0 mm, was lost with the Hamburg Museum 5. Cryphalities Cockerell (1917:368) was in 1944. The antennal scape was elongate, the based on C. rugosissimus Cockerell (1917: funicle was 5-segmented, the club was flat- 368) from Burmese amber. Because no usable tened, with a suture near the apex, and the characters were included, the original de- frons was impressed above the eyes. This scription gives no clue as to where this species genus almost certainly is in the Dryocoetini, should be placed. It is assumed that Cockerell possibly near Tiarophoru.s or Xylocleptea was correct in assigning it to Cryphalini. I

4. Pseuc?o;?iicrac!.S' Eggers (1920a:36), based have not examined the specimen. on P. ekae Eggers (1920a:36) from Dares- 6. Xyleborites Wickham (1913:26) was Salaam, Ostafrika. The unique female holo- based on X. longipennis Wickham (1913:26) type, 1.5 mm, was lost with the Hamburg from the Wilson Ranch near Florissant, Colo- Museum. The frons was impressed, the an- rado. The author indicated that this species, tenna was similar to Micracis, the scape was 2.2 mm in length, resembled triangular, with long hair on the outer angle, pubescens . Essential characters were not de- and the sutural apex of the elytra mucronate. scribed. The specimen was not examined by Because only one known genus bears the com- me. bination of characters described by Eggers, Pityophthoridea Wickham (1916:18) and Adipocepludus Wickham (1916:16) are not his name is associated with those African spe- cies previously referred to the American considered to be members of the Scolytidae. genus Micracis. As such, this genus is recog- nizable and is treated below under the name Systematic Section Pseudomicracis Of the six fossil genera that have not been Although some obvious unanswered ques- previously placed in synonymy under older tions remain as to the exact position of Platy- names of modern genera, four are from Baltic podidae and Scolytidae within the Cur- amber, one is from Burmese amber, and one culionoidea, the following key is presented for is from sedimentary deposits. These include: the identification of families. The presence of 1. Carphoborites Schedl (1947:32) was complete pregular sutures and the mandible based on C. keilbacki Schedl (1947:32) and C. structure of Platypodidae and Scolytidae are posticm Schedl (1947:.33), both from Baltic obviously more primitive than the absence of amber (Charphoborites was a lapsus calami). these sutures and the specialized mandibular

From the descriptions, I see no reason to structure of Belidae and Oxycorynidae, but separate these species from Carphohorus; the single gular suture and presence of larval however, I have not examined the specimens. epipharyngial rods are more advanced than 26 Great Basin Naturalist Memoirs No. 10 the two gular sutures and absence of the rods Ithyeeridae, Apionidae, some Platypodidae, • and a few Sc 'ytidae). Larva: epipharyngeal in Behdae. The long subcortical history and rods present; maxillary palpi 1- or 2-seg- very small body size of Platypodidae and mented (3-segmented in Brenthidae and Scolytidae have had an obvious effect on their Rhynchitidae); thoracic legs absent (present anatomy that complicates interpretation of in Brenthidae and Ithyeeridae); frontal su- tures not reaching articulating membrane of their position in phylogeny. mandible (except Brenthidae, Apionidae, key to families of the Curculionoidea is This Ithyeeridae, some Plat\ podidae); two to four tentative. A thorough reexamination of the folds in each abdominal tergmn 6 Belidae-Aglycyderidae-Oxycorinidae groups 2(1). Adult; labrum distinct and separate (except is needed to determine the significance of absent in Bnichela of Anthribidae); maxillary single vs. double gular sutures, the precise palpi flexible, laeinia forming a distinct ele- ment (see also Rhynchitidae and Tessero- position of character gaps that separate cerini in Platypodidae). Larva: mandibles families, etc. Another area of concern is the with a distinct mola 3 section of the key in couplets 11 and 12. Char- — Adult: labrum fused or lost; maxillary palpi acters to separate these three taxa are weak, rigid; maxilla without a separate laeinia. but, in view of the large number of species to Larva: mandibles without a distinct mola .... 4 classified herein, perhaps the recognition be 3(2). Adult: gular sutures rudimentary, repre- of these three families is justified. Among sented only at margin of foramen magnum or more than 150 genera of Curculionidae (in the totally lost; tentorium largely or entirely obso- lete; proventriciilus developed; lateral mar- sense used here) dissected by me, I saw three gins of pronotum acute (rounded in possible divisions of this family. These in- B ruchela ); inner margin of elytra with a flange cluded (1) the Cossoninae-Rhynchophorinae, near costal margin; cavities of mesocoxae (2) the broadnose , and (3) all others. It broadly closed ouKvardly by sternum, visible abdominal sterna 1-4 connate. Larva: clypeus is recognized that some equally distinct distinct from frons, or head deeply retracted; groups may exist that were not seen by me, mandibles without ventral process in addition anatomical study but a thorough comparative to mola (includes the distinct subfamily should precede their recognition. Following Bruchelinae) Anthribidae my study, I seriously doubt that even one- — Adult: gular sutures extending from margin of third of the 75 or more subfamilies that have foramen magnum to posterior tentorial pits; been recognized within Curculionidae de- tentorium present; proventriculus indistinct; inner margin of elytra without a flange near serve this rank; most are worthy of no more costal margin; mesocoxae not or imperfectly status. than tribal closed outwardly by sternum; abdominal sterna free. Larva: clypeus usually fused to to the families of Curculionoidea Key frons; head not deeply retracted into protho- rax; mandibles with a ventral process in addi- 1. Adult: two gulav sutures present or rudiments tion to mola Nemonychidae of them evident (only one gular suture and a postgula present in several Belidae and Rlio- 4(2). Adult: gular sutures longer, extending to pos- palotria of O.xycorynidae); maxillary palpi 4- terior tentorial pits; partial pregular sutures segmented; abdominal segments free (e.xcept occasionally present in female near apex of 3-6 connate in Anthribidae); antennae usu- rostrum; at least three genera with large post- ally orthocerous, scape little longer than suc- gula and short median gular suture; antennae ceeding segments, club loosely 3-segmented. filiform or nearly so, inserted laterally remote Larva: epipharyngeal rods absent; maxillary from base of rostrum; inner margin of elytra palpi 3-segmented, rarely 2-segmented; tho- with a flange near costal margin; wing with racic legs present; frontal sutures reaching five anal veins in main group complete to articulating membrane of mandible; two folds base; tarsi normal, pseudotetramerous; lat- in each abdominal tergum 2 eral margins of pronotum transversely rounded. Lar\a: maxillary palpiger well de- — Adult: one gular suture extending from veloped; thoracic spiracle intersegmental . . . minute postgula at margin of foramen mag- Belidae num to posterior tentorial pits (usually where base of rostrum meets head); maxillary palpi — Adult: gular sutures usually short (a large 1- to 3-segmented (4-segmented in one Platy- postgula and one median gular suture in some podidae, some Rhynchitidae, and all Atte- Oxycorynidae), externally visible near fora- labidae); abdominal segments free or partly men magnum or else lateral margins of prono- connate in some groups; antennae frequently tum acute; antennae inserted at or near base geniculate, a definite club usually present (or- of rostrum; either antennae clubbed or tarsi thocerous in Brenthidae; intermediate in pseudotrimerous; inner margin of elytra with- 1986 WOOD; Genera of Scolytidae 27

out a flange near costal margin; wings with ventrad; mesepimeron flat, vertical; mese- fewer than five anal veins or else bases of pisternum moderately to greatly enlarged, elytral veins incomplete. Larva: maxillary varying from flat to inflated; eyes convex, palpiger usually not evident; thoracic spiracle never truly emarginate; pronotum never in mesothorax 5 armed by asperities. Larva: clypeus fused to frons in aU except primitive genera; frontal 5(4). Adult: tarsi pseudotrimerous, segment 2 sutures reaching articulating membrane of bilobed; antennal insertion at base of rather mandible or not Platypodidae short rostrum; antenna filiform, not clubbed;

proventriculus well developed. Larva: ma.\il- — Adult: tarsal segment 1 about ecjual in length lary palpiger not distinct to 2 or 3; tibiae armed on apical and/or lateral Aglycyderidae (=Proterhinidae) margins either by spines or socketed denti- cles, apical mucro formed by mesal element — Adult: tarsi pseudotetramerous, segments 2 (compared to Protohijlastes structure); male and .3 bilobed; antennal club 2- or 3-seg- spiculum gastrale present; antennal club vari- mented; (a large postgula and at least a partial able, with or without sutures; head narrower median gular suture present in at least Rho- than pronotum; pronotum almost never with palotria), lateral margins of pronotum acute; constriction near middle; when visible, antennal insertion at base of long rostrum, on scutellum usually flush with elytral surface, ventral face; proventriculus poorly devel- often flattened; metepisternum stout to elon-

oped. Larva: palpiger distinct . . Oxycorynidae gate, its anterior margin usually procurved,

6(1). Adult: pregular sutures present and rein- its anteroventral angle displaced slightly cau- forced internally by massive apodemal ridges; dad and distinctly extended ventrad; mese-

rostrum short to nonexistent; tibiae armed on pimeron flat, oblique; mesepisternum apical or lateral margins by a series of spines smaller, declivous cephalad; eyes more nearly and/or socketed teeth of setal origin, corbels flat against head, often emarginate to divided; never present; hypostomal area small, never pronotum often asperate. Larva: clypeus dis- armed by a large spine, posterior mandibular tinct, separate from frons; frontal sutures condyle more anterior in position; axis of never reaching articulating membrane of mandibular hinge oblique to transverse, cut- mandibles Scolytidae

ting edge on mesal margin (Fig. 8); some 8(6). Adult: labial palpi inserted ventrally on men- members (Tesserocerini) with a separate tum; head frequently constricted behind lacinia 7 eyes; antennae never geniculate, inserted — Adult: pregular sutures absent, with no indi- dorsally or laterally at base or middle of ros- cation of supporting internal apodemal struc- trum, club absent to moderately distinct; ture (longitudinal ridges giving rigidity to ros- proventriculus without selerotized plates. trum are not reinforced internally and should Larva: frontal sutures reaching articulating not be confused with true pregular sutures); membrane of mandibles 9 rostrum variable, long to very short; lateral — Adult: labial palpi inserted apicafly; antenna margins of tibiae unarmed (except for simple straight or geniculate, inserted laterally at spines in some Cossoninae), corbels com- middle or near apex of rostrum, club 1- to monly present; hypostomal area larger, com- 3-segmented, loose to compact, small to monly armed by a large spine, posterior large. Larva: frontal sutures either reaching mandibular condyle more posterior in posi- or not reaching articulating membrane of tion (Fig. 8); axis of mandibular hinge parallel mandibles 10 to body axis, cutting edge on apical margin; separate lacinia never present (except some 9(8). Adult: maxillary palpi 4-segmented, galea and Rhynchitidae and all Attelabidae) 8 lacinia distinct; mandibles often dentate on outer margin; labial palpi 1- or 3-segniented 7(6). Adult: tarsal segment 1 elongate, usually (rarely absent); elytra with a scutellary striole; longer than 2-.5 combined (except shorter in notosternal suture short, laterally reflexed; Protoplatypiis , Protohijlastes. Coptonotiis. hind wing usually witli four anal veins, radial Scolytotarsus), tibiae armed on apical and/or cell well developed; sterna 3-6 rarely free, lateral margins by one or more spines, never .3-4 connate in some, 3-6 connate in most; by socketed denticles, apical mucro formed cap piece of male tegmen simple, setose; by middle apical spine (as compared to Proto- malpighian tubules grouped three on each hijlastes), male spiculum gastrale absent; an- side of alimentary canal; more than two ovari- tennal club solid, unmarked by sutures (fee- oles in each ovary. Larva: legs absent, two or bly indicated in Coptonotiis), head about as more stemmata present on each side of head; wide as pronotum; pronotum with distinct antenna usually 2-segmented; maxillary palpi lateral constriction near middle; when visible, 2- or 3-segmented; abdominal terga with two scutellum often reduced, declivous, usually folds Attelabidae with not more than apex attaining (flush with) elytral surface; metepisternum very elongate, — Adult: maxillary palpi 3-segmented, galea and

its anterior margin straight, its anternventral lacinia fused; mandibles never dentate on angle neither displaced caudad nor extended outer margins; elytra never with scutellary 28 Great Basin Naturalist Memoirs No. 10

striole; notostemal suture well developed and Family Scolytidae Latreille usually extending almost to anterior margin; hind wing with three or fewer ana! veins, Scolytarii Latreille [1807:273, Type-genus: Scolytus Ge- radial cell not evident; sterna 3-4 connate and offrey 1762, see also China 1962] much longer than 5-6; cap piece of tegmen bilobed. Larva: rudimentary, segmented legs The family Scolytidae is comprosed of a present; stemmata absent; antenna 1-seg- group of more than 6,000 small to minute mented; maxillary palpi 3-segmented; ab- species. Whereas the preponderance of spe- dominal terga with three to four lolds cies are tropical, a few of them reach the Brenthidae northern and southern limits of tree distribu- 10(8). Adult: maxillary galea and lacinia distinct, tion near the polar regions. They are unique palpi 3- or 4-segmented; metepimeron sub- in that loosely pair-bonded adult parents bore transversely carinate and interacting with into subcortical tissues of their host (usually) humeral angles of elytra to aid in locking ely- tra in position, metepisternal locking element before mating or oviposition occur. Eggs are not developed and its interlocking flange at variously placed in the galleries, and the lar- this point on inner costal margin of elytra vae either expand the parental mines or form absent; antenna straight, club loosely 3-seg- individual galleries while the parents defend mented; abdominal sterna 3 and 4 connate, the outer entrance to the tunnel. Thus, the suture distinct. Larva: median area on poste- rior part of head overlapped by prothorax; beetles are essentially internal plant parasites maxillary palpi 2- or 3-segmented; two basal that spend virtually their entire lives secreted sensillae on labrum; frontal sutures complete within the tissues of the host. The ephemeral but indistinct; abdominal terga with two folds habitat they occupy is the unthrifty, weak- Rhynchitidae ened, or dying tissue of woody plants; only — Adult: maxillary lacinia usually fused to galea, one generation is completed in each host un- palpi 2- or 3-segmented; metepisternum cari- less a large tree that is succumbing progres- nate, fitting flange on inner margin of elytra near humeral angle, metepimeron not cari- sively is involved. Because of the very brief nate; antenna straight or geniculate, clubbed period this habitat is open to them, efficient or not. Larva: posterior part of head free in means of locating a host, recruiting a popula- median area; maxillary palpi 1- to 3-seg- tion to subdue it, and overcoming host resis- mented; no basal sensillae on labrum 11 tance are mandatory. This is accomplished 11(10). Adult: abdominal terguni 8 longitudinally through sensitivity to odors emitted by trees grooved for reception of inwardly raised ele- under stress, by a complex system of phero- vation at suture near apex of elytra; tergum 8 mones, and by mutualistic relationships with bearing a pair of spiracles; maxillary palpi 3- segmented, but largely retracted into fungi. Several of the most efficient species, palpifer; antenna not geniculate. Larva: rudi- with respect to meeting these challenges, mentary, segmented legs present; maxillary compete with man for timber and horticul- palpi 2-segmented; 3 stemmata present .... tural resources and present a very real threat Ithyceridae to human economy. Most woody plants, a few Adult: abdominal tergum 8 not grooved, spir- — herbaceous plants, and many fruits and nuts acle absent; maxillary palpi 2- or 3-seg- are infested by these insects. The adjustments mented, usually not retracted into palpifer . 12 they have made in their mating systems, uti- 12(11). Adult: either antenna not geniculate, or if lization of food resources, and adaptive radia- geniculate then trochanters long, cylindrical, tion form a classic example of biological diver- femur remote from coxa, attached to apex of trochanter; ventral surface of mentum usually sity. with projecting setae; proventriculus poorly developed. Larva: frontal sutures extending Key to the Subfamilies and Tribes of Scolytidae to articulating membrane of mandibles; ab- (Modified from Wood 1978) dominal terga with two folds Apionidae 1. Each basal margin of elytra procui-ved and — Adult: antenna usually geniculate; troch- armed by a series of marginal crenulations anters triangular, femur attached to side of (Fig, 16) (or less commonly by a continuous trochanter, sometimes almost touching coxa; elevated costa in some Bothrosternini, Di- ventral surface of mentum without projecting amerini, Polygraphini), usually with a scutel- setae; proventriculus usually well developed. lar emargination between them; scutellum Larva: frontal sutures not reaching articulat- usually small and rounded or depressed, ab- ing membranes; abdominal terga with two to sent in some groups; pronotum weakly if at all four folds Curculionidae 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.

but crenulations sometimes present on — Scutellar area of metanotum and its postno- anterolateral areas; head usually visible from tum completely fused on at least median above, somewhat wider; protibia usually third, intersegmental suture usually obsolete wider; scales or deeply divided setae a com- (Figs. 21, 23 pt. 41); scutoscutellar suture less mon feature (subfamily HYLESININAE) .... 2 strongly curved, approaching costa of scutel- lar groove more gradually and continuing Basal margins of elytra forming a straight, cephalad parallel to it for less than half length transverse line across body (Fig. 17), un- of metanotum (Fig. 23, pt. (it never armed, rarely (some Scolytini, Ctenophorini, 41) reaches margin of this groove in some groups; Cryphalini) with a weakly elevated continu- metapleural suture sometimes as described ous line; scutellum usually large, flat (rarely above, but more commonly running a more absent or highly modified in some Xyle- direct route from pleural wing process to borini); pronotum weakly to strongly de- pleural costal process, often remote from clivous on anterior half and usually armed by locked position of costal margin of elytra for many asperate crenulations, particularly on most or all of its course (Figs. 21, 23 pt. 41); median half (Figs. .31, 46, .52); head usually scutellum either not visible or if visible then partly or entirely concealed from dorsal as- funicle 5-segmented and male frons im- pect, somewhat narrower; protibia usually pressed (Bothrosternini with 6-segmented fu- narrower; scales or deeply divided setae an nicle but with a distinctive protibia. Fig. 24) 7 uncommon feature (subfamily SCOLYTl- NAE) 12 3(2). Eye entire to feebly emarginate; scutoscutel- lar suture parallel to costa of scutellar groove 2(1). Scutellar area of metanotum and its postno- for two-thirds length of notum; preco.xal ridge tum separated by a suturelike intersegmental on prothorax present (Fig. 20) or absent; an- membrane (Fig. 23, pts. 43, 45); posterior tennal funicle 5- to 7-segmented 4 part of scutoscutellar suture strongly curved divided mesad to a point near crest of scutellar groove — Eye completely by an emargination, halves widely then continuing cephalad parallel to this costa separated; scutoscutellar suture costa of scutellar groove; for about two-thirds of metanotum length remote from crenu- lations on basal margins of elytra low, often (Fig. 23, pt. 43) (except much less in Phrixoso- mini and Hyorrhynchini); metapleural suture poorly formed; precoxal ridge on prothorax descending subvertically from pleural wing never present; antennal funicle usually 6-seg- 6 process to metepisternal groove formed to mented (5-segmented in Sueus) receive corresponding costal groove and 4(3). Prothoracic precoxal area rather large, its lat- flange of elytron then abruptly angled and eral margins strongly, sharply elevated from continued caudad along this groove to a point anterior margin to coxae (Fig. 20); crenula- near pleural coxal process (Figs. 18, 23 pt. 44); tions on elytral bases usually poorly devel- scutellum visible; funicle 6- or 7-segmented oped; antennal funicle 7-segmented, club

or if 5-segmented (Sueus) then eye divided, conical, segment 1 often as long as others male frons not impressed, and antennal club combined; head somewhat prolonged, sub- symmetrical 3 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, Sueus), female frons convex; antennal scape

except introduced elsewhere; in Pinaceae . . either long or short, club aseptate; pronotum Hylastini either with or without asperities in anterolat- eral areas; southeast Asia Hyorrhynchini — Prothoracic precoxal piece small, short, its

lateral areas elevated or not; crenulations on 7(2). Lateral margins of pronotum usually sub- elytral bases more conspicuously elevated, acutely elevated, costate (as in Fig. 12); forming a definite row (except confused in mesepimeron moderately to very large, its Dactijlipalpus): antennal funicle variable, .5- dorsal portion usually grooved for reception of to 7-segmented, club weakly to moderately elytral base; scutellar shield under base of flattened; head less distinctly rostrate, male elytra large, extending posteriorly beyond frons usually impressed, eye oval to elongate, visible scutellum (Fig. 26); scutoscutellar su- entire to feebly emarginate 5 ture remote from costa of scutellar groove to

its base; apical protibia 5(4). Pronotum asperate on anterolateral areas (ex- outer angle of often cept ), prothorax with elevated with only one major recurved spine (Fig. 7);

costate ridge from coxa to anterior margin Africa, southeast Asia to Australia . . Diamerini 6- (weaker than in Fig. 20); antennal funicle or Lateral margins of pronotum rounded (Fig. 7-segmented; mesal surface of elytra at base of 16) (subcostate in a few neotropical Both- suture immediately behind scutellum with an rosternini. Fig. 11); mesepimeron not en- interlocking series of nodules and cavities larged or grooved (feebly grooved in (Fig. 25-C), this lock interrupts groove and Aricerus), scutellar shield beneath elytra visible in flange of suture (not when elytra small if present, not extended caudad beyond locked position); worldwide Hylesinini visible scutellum (Figs. 18, 21); scutoscutellar — Anterolateral areas of pronotum unarmed; suture near and parallel to costa of scutellar precoxal costa on prothorax absent; funicle 5- groove on at least anterior fourth of metan- to 7-segmented; mesal surface of elytra at su- otum 8 ture with interlocking groove and flange con- ^(y\ Outer apical angle of protibia with a curved tinuted to base without a series of nodules or bifid process, meso- and metatibiae with one cavities immediately behind scutellum (Fig. or two (usually smaller) cui-ved spines on 25-A-B); worldwide Tomicini outer apical angle extending beyond level of

6(3). Protibia armed on outer apical margin by sev- spine on inner apical angle (Fig. 24); prono- eral socketed denticles of approximately tum smooth or longitudinally strigose; funicle equal size; procoxae contiguous; frons convex, 6-segmentecl; lateral prosternal area usually not sexually dimorphic; antennal scape elon- subacutely elevated from coxa to anterior

gate, suture 1 of club partly septate; prono- margin; anterior coxae rather widely sepa- tum never armed by asperities; America, rated; crenulations on elytral bases rather Africa Phrixosomini small or (rarely) replaced by a continuously elevated costa; eye entire; America — Protibia with outer apical angle produced into Bothrosternini one conspicuous spine reaching level of tarsal insertion, outer margin without any socketed — Outer apical margin of protibia armed by sev- teeth; procoxae rather widely separated; male eral teeth of about equal size (except Aricerus frons broadly impressed (except in dwarfed 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 Fig. 27. Antenna of Vhloeotribus . A, rhododacttjlus rounded, costa obsolete; eye varying from en- (Marsham); B, caucasicus Reitter. tire to emarginate to divided 9

9(8). Scutellum visible, elytral bases notched for its yond process of inner apical angle (Fig. 7, reception; tarsal segment .3 stout, usually Camptocerus). lateral margins of pronotum somewhat bilobed (except slender in subacutely elevated, costate; pleural suture ), mesal surface of elytra at suture descending subvertically pleural immediately behind scutellum with a series of from wing process to groove receiving groove and flange interlocking nodules and cavities (Fig. 25-C) on at this 10 costal margin of elytra, point suture turns abruptly and follows groove caudad to — Scutellum obsolete, elytral bases only slightly metapleural coxal process (Fig. 23, part 46); if at all emarginate at suture (Fig. tarsal 17); funicle 7-segmented, sutures of antennal club segment .3 slender; mesal surface of elytra at strongly procurved or obsolete; Holarctic and suture usually without a special lock, groove Neotropical Scolytini and flange extend to base at position of .scutel- — Lateral margin of protibia armed by more lum (degenerate in Fig. 25-A-B) 11 than one denticle, none of which exceed or 10(9). Antennal club constricted at sutures and mov- curve toward inner apical process; pleural su- able at intersegmental lines (Figs. 16, 27); ture less strongly angulate, groove receiving Holarctic, Neotropical, and Australian flange of costal margin of elytra displaced ven- Phloeotribini trad from course followed by pleural suture — Antennal club immovably, broadly fused at (Fig. 23, part 42); lateral margins of pronotum sutures, sutures often partly or entirely obso- subacutely raised or not, antenna variable .. 13 lete; worldwide Phloeosinini 13(12). Metepisternum visible throughout its length, 11(9). Eye emarginate or entirely divided; prono- shghtly more than its dorsal half covered by tum never armed by asperities; crenulations elytra when in locked position, either with a at bases of elytra more widely distributed, conspicuous groove for reception of costal extending laterad beyond interstriae 5 (Fig. flange throughout its length or else groove

16); funicle 5- or 6-segmented; scutoscutellar represented at its anterior end by a denticula- suture passing near and parallel to costa of tion or costate remnant near anterior end of scutellar groove on anterior fourth of meta- metepisternum (Fig. 23, parts 42, 44, 46);

notum; Northern Hemisphere and Africa . . . antennal club varying from flat to obliquely Polygraphini truncate (Fig. 29) 14

— Eye sinuate or entire; pronotum armed by a — Metepisternum largely covered by elytra, its few scattered or clustered asperities; crenula- groove for reception of costal flange obsolete, tions at bases of elytra restricted to area be- a small, transverse callus (Cryphalini, Fig. 31)

tween suture and interstriae .5; funicle 3- to or a small transverse groove (Corthylini, Figs. 6-segmented; scutoscutellar suture remote 23, part 47, and 32) at anterior end of met- from costa of scutellar groove on anterior episternum; antennal club strongly flattened; fourth of metanotum; almost worldwide .... antennal club never obliquely truncate 24 Hypoborini 14(13). Lateral margins of pronotum subacutely ele- 12(1). Lateral margins of pro- and metatibiae un- vated, basal margins of elytra usually finely armed except for a single, apical, spinelike elevated; procoxae rather widely separated process that curves toward and extends be- 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 ginate (divided in Tiarophorus , Dryocoetini, (Fig. 7, Scolytodes). usually extending be- Africa); pronotum sometimes with a raised yond tarsal insertion, posterior tibia tapered line on basal or lateral margin; prcgnlar area on apical third and armed by several small not depressed; se.xes of similar size and body socketed denticles; funicle6-or7-segmented; form (except male dwarfed and deformed in tarsi often retractible into tibial grooves .... 15 Coccotrypes and Ozopcmon); habits varied but never woodboring or mycetophagous ... 20 — Lateral and basal margins of pronotum rounded (except ); procoxae subcon- — If eye completely divided into two parts and tiguous (except most Micracini and a few antennal funicle 4-segmented then male frons Xyleborini); protibia with outer apical angle deeply excavated and male equal in size to

inconspicuous, armed by several small sock- female; if eye emarginate (or if divided and eted denticles; funicle 2- to 6-segmented; funicle 5-segmented) then male dwarfed, de- tarsi not retractible (except in Eccoptopterus) formed, and flightless and female meso- and 17 metathoracic tibiae expanded to just beyond middle then arcuately tapered to apex, its 15(14). Eye divided or nearly so by a very deep emar- apical two-thirds on outer margin armed by a gination; antennal club flat, usually enlarged, row of numerous small, closely set teeth of with sutures strongly procurved; abdomen equal size, these usually supplemented in conspicuously ascending toward apex (not al- same row by submarginal hair on posterior ways clear in Clonoxijlon). tarsi always re- face; male pronotum highly modified; pregu- tractible into tibial grooves; Africa and south- lar area depressed (except ), em Asia Xyloctonini woodboiing, mycetophagous 23 — Eves entire to shallowly sinuate on anterior margin; antennal club flat, usually smaller, 20(19). Pronotum rather strongly, laterally con- more slender, sutures variable, if present; ab- stricted on posterior half anterior half not domen horizontal; tarsi retractible or not ... 16 declivous and never armed by asperities; anterior coxae moderately separated; anten- 16(15). Antennal club with one or more sutures indi- nal club strongly flattened, marked by two cated by grooves, setae, or septae; scutellum sutures, sutures on posterior face almost large, flat; America; mostly phloeophagous, never mycetophagous Ctenophorini equal to those on anterior face; South America and India to Ceylon (Sri Lanka) — Antennal club unmarked by sutures; prono- Carphodicticini tum with sides strongly constricted on poste- rior half scutellum absent (a small scutellum — Pronotum not constricted, sides straight to present in Scolytoplattjpus congonus). Africa arcuate, anterior half usually declivous, usu-

to Asia and New Guinea; mycetophagous . . . ally armed; anterior coxae contiguous; anten- Scolytoplatypodini nal club obliquely truncate or with sutures on

posterior flice strongly displaced toward apex 17(14). Procoxae moderately separated; protibia with obsolete) 21 sides parallel, armed by denticles only on api- (rarely with sutures cal margin or posterior face; funicle 6-seg- 21(20). Eye shallowly sinuate (shallowly emarginate mented (5-segmented in one African genus); in some Acanthotomicus), its lower half dis- female frons often concave, male frons rarely tinctly narrower than above; piotibia with 3-4 concave (except two Pseudotlujsanoes); Africa socketed teeth; antennal club rarely obliquely and America, one species in Asia .... Micracini truncate (Pityokteines. ), pro- — Procoxae contiguous (except Carjihodicticini, coxae contiguous, intercoxal piece longitudi- some Xyleborini); protibia much wider api- nally emarginate to absent, never complete; cally, armed on lateral margin by several den- elytra moderately sulcate to elaborately exca- ticles; female frons rarely concave (a few Dry- vated, with lateral margin usually armed by ocoetini), male frons often concave; funicle 2- tubercles or spines; pronotum more strongly to .5-segmented, 6-segmented in Tiurophorus declivous on anterior third, asperities usually 18 larger; worldwide Ipini

18(17). Male frons strongly excavated, epistoma — Eye sharply, rather deeply emarginate (sinu- armed by a pair of (usually) fused horns of ate in Deropria), lower half usually almost enormous size (Fig. 47); funicle 5-segmented, equal in width to upper half protibiae usually club often small and feebly flattened; eye with four or more socketed teeth (most excep- small, entire; pronotum with summit near tions with lateral margins of pronotum acutely basal margin, projecting back over scutellum elevated); procoxae either contiguous or dis- in some species; western United States and tinctly, narrowly separated; elytral declivity Mexico Cactopinini flattened to convex, unarmed by spines or — Not fitting above combination of characters . 19 large tubercles; pronotum either evenly

19(18). Meso- and metathoracic tibiae more slender, arched from base to anterior margin or less more abruptly narrowed on apical fourth, lat- strongly declivous on anterior third, asperi- eral and apical margins armed by fewer, ties, when present, usually fine and abundant coarser teeth; eye sinuate to shallowly emar- (a few exceptions) 22 34 Great Basin Naturalist Memoirs No. 10

22(21). Antennal funicle 4- to 6-segmented, club ei- there has been variability in the taxonomic ther obliquely truncate or with .sutures on rank assigned to it. Most of the distinguishing posterior face strongly displaced toward apex; characters previously are found anterior half of pronotum more strongly de- employed not clivous and rather coarsely asperate (unarmed consistently throughout the group and, conse- in Tiarophorus); worldwide Dryocoetini quently, have little or no taxonomic value. — Antennal funicle 2- or 3-segmented, club with The most consistent and reliable character sutures on posterior face about equal to those available for the recognition of this subfamily on anterior face; pronotum feebly declivous is the procurved basal margins of the elytra on anterior half and unarmed (minutely gran- that are armed by a row of crenulations, and ulate in some Aphanarthrum), reticulate in many species; size small; Northern Hemi- the scutellar notch between them. The heav- sphere and Africa Crypturgini ier, more coarsely armed tibiae are distinctive

23(19). Eye always completely divided into two parts; but less reliable as a distinguishing feature. antennal funicle 4-segmented, base of club The more primitive Bothrosternini, Diame- feebly to moderately corneous, usually rini, and a few other isolated examples lack pubescent to base; male subequal in size to the specialized elytral crenulations and have female, his frons flattened or e,\cavated and anterior margin of his pronotum more broadly the basal margins elevated along a continuous rounded; male joins female in parental costa reminiscent of some Platypodidae. This gallery, reproduction always bisexual; Hol- same character also occurs in Dendrodicticus arctic and Oriental Xyloterini (Carphodicticini) and, in a greatly reduced — Eye emarginate except divided in some form, many Scolytini, Ctenophorini, and Amasa; funicle 5-segmented (3- or 4-seg- Cryphalini. On the other hand, a few mented in a few Asiatic forms); males flight- less, dwarfed, deformed, anterior slope of Cnemonijx (allies oi galeritus, Scolytini) have pronotum variously excavated; male head fully formed basal crenulations that suggest an convex; male absent from parental gallery ex- affinity with the Hylesininae. Even though cept as progeny; partly parthenogenetic, male the demarkation between subfamilies is not as haploid; almost worldwide Xyleborini sharp as some would like, the division of sub- 24(13). Costal margin of elytra slightly to moderately families is simple and the characters are reli- ascending from base of declivity to apex; basal end of metepisternum armed by a callus or able. partial groove of degenerating metepisternal spine (Fig. 31); sutures on posterior face of Tribe Hylastini antennal club more strongly displaced toward apex; funicle 3- to 5-segmented; tibiae more Hylastes LeConte [1876:387, Type-genus: Hylastes strongly flattened, usually armed by more Erichson, 1836] than four denticles; vestiture commonly in- cludes scales; eye usually entire, less com- monly emarginate; worldwide Cryphalini Description. —Frons not sexually dimor- phic; eye oval, entire; antennal scape elon- — Costal margin of elytra descending toward apex (e.xcept Brachyspartus), basal end of gate, funicle 7-segmented; precoxal ridge on metepisternum with a small, transverse prothorax strongly, acutely elevated; protibia groove (Fig. 32) (concealed when elytra in very broad, with rather numerous lateral locked position), elytra in locked position socketed teeth; scutellum visible; crenula- more completely cover metepisternum (Fig. at developed, usu- 52 part 5, and 53); sutures on posterior face of tions base of elytra poorly antennal club only slightly displaced toward ally net forming a definite row; declivital in- apex; funicle 1- to 5-segmented; tibiae more terstriae 10 sometimes present (Scieriis); slender, rarely armed by more than four sock- declivital sculpture usually simple; vestiture eted denticles; vestiture rarely includes scales usually includes scales (some exceptions); scu- (in tropical forms only); eye emarginate; al- most worldwide except Australia .... Corthylini toscutellar suture parallel to costa of scutellar groove for two-thirds length of pronotum; Subfamily Hylesininae metapleural suture descending subvertically from pleural wing process to metepisternal Hylesinen Erichson [1836:46, Tvpe-genus: Hylesinus groove (this groove interlocking with groove Fabricius, 1801] on costal margin of elytron) then continuing Most previous classifications have recog- caudad along this groove to a point near nized the Hylesininae as a major division of pleural coxal process; tarsal segment 3 wider the taxon treated here as Scolytidae; however. 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 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. . —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

America; Picea, Abies, Pinus, 2. 7-4. .3 mm .... ica where Picea grows. Both are monogamous Scierus and breed in phloem of roots, butts, and — Anterior coxae contiguous or at most with inter- stumps of standing trees or next to the ground coxal piece narrower than one-fourth width of a in the lower bole of prostrate trees. Keys: coxa; striae 10 obsolete behind level of posterior Bright (1976:41), Wood (1982:79). coxae, only striae 9 continuing caudad; elytral Type-spe- vestiture longer, more abundant, usually in- Hylurgops LeConte [1876:389, cluding at least some small scales 2 cies: Hylastes pinifex Fitch =Hyhirgops rugi- 36 Great Basin Naturalist Memoirs No. 10

pennis pinifex (Fitch), subsequent designa- Alniphagini Murayama [1963:i!i, 29, Type-genus: A/nip/i- ag».5 Swaine, 1918] tion by Hopkins 1914:123. Synonyms; Hyle- sinites Germar 1813:15, Type-species; Hyle- Description. —Frons obscurely to deeply sinites electriniis Germar, monobasic; Hijla- and extensively impressed in male, female flat stites Hagedorn 1907:117, Type-species: Hy- to variously convex; eye oval to elongate, en- lastites schellwieni Hagedorn, monobasic; tire to feebly sinuate on anterior margin; an- Myelophilites Hagedorn 1907:118, Type-spe- tennal scape very short to very elongate, fun- cies: Myelophilites duhiits Hagedorn, mono- nicle 6- or 7-segmented (except 5- to 7- basic; Hylescierites Schedl 1947:29, Type- segmented in Hylesinopsis), club conical to species; Hylescierites granulatus Schedl, moderately flattened, symmetrical or nearly monobasic]. Distribution; 7 species, 2 with so, two or more sutures indicated, except su- subspecies, in North America; 2 in N Africa; tures absent in Dactylipalpus : procoxae nar- 18 nominate species in Europe and N Asia; 7 rowly to moderately separated, precoxal ridge nominate fossil species including 1 from on prothorax moderately to very strongly, North America and 6 in Baltic amber acutely elevated; pronotum armed by a few (Oligocene). All are monogamous and breed asperities, except absent in Hylastinus; in tissues of Pinaceae; most are in phloem metascutellar area separated from postnotum roots, butts, and stumps of standing hosts, by a distinct suture; mesal surface of sutural some in logs, and at least one may occur in groove of elytra just behind scutellum inter- smaller material. Keys: Wood (1982:82) for rupted by a series of interlocking nodules and North America, PfefFer (1944) for Europe and cavities; tarsal segment 3 usually wider than 2, Africa, Tsai (1964b) for China, N and Huang often bilobed. Murayama (1963) for NE Asia. Biology. —All species are monogamous. Hylastes Erichson [1836:47, Type-species; All are phloeophagous except the xylophagous Bostrichus ater Paykull, subsequent designa- Phloeoborus and Dactylipalpus . Parental gal- tion by Westwood 1838:39 and Thomson leries are biramous or, if a well-developed 1859:146]. Distribution; 16 species in North turning niche is present, they may be mono- America; 1 in Jamaica; 17 nominate species in ramous. Eggs are deposited in niches and are Europe and N Asia; 1 in N Africa; 1 in the sealed in with frass. Larval mines usually fol- Canary Islands; fossil species include 2 from low a definite course away from the parental Baltic amber (Oligocene), 1 from Miocene of mine and rarely cross one another. Phloeo- Colorado. All are monogamous and breed in borus and Dactylipalpus appear to have a phloem tissues of Pinaceae; most are in roots, symbiotic relationship with fungi, but not in a butts, and stumps of standing trees, a few in mycetophagous sense (old parental mines are the bole of prostrate trees next to the ground; stained black and wood adjacent to larval seedlings are sometimes infested, apparently mines is discolored and in a more advanced under emergency stress, for the purpose of state of decay). maturation feeding or to wait for a suitable Taxonomy. —The occurrence of this tribe host. Keys: Wood (1982:93) and Blackman in the Eocene and its almost worldwide distri- (1941) for North America, Pfeffer (1944) for bution suggest an ancient origin. The strictly Europe, Murayama (1963) and Tsai and neotropical Phloeoborus and ethiopian (ex- Huang (1964a) for NE Asia. cept for two oriental species) Dactylipalpus Tribe Hylesinini suggest that these unique but related genera were derived from a common ancestral Hylesinen Erichson [1836:46, Type-genus: Hijlesinus African stock at or before the time of separa- Fabricius, 1801] tion of South America and Africa. The remain- Phloeotrupides Chapuis [1866:357, Type-genus; Phloeo- ing Hylesinini include representa- trupes Erichson, 1836 =Phloeoborus Erichson, American

1836] tives of Hylcsinus and , both of Phloeobori Blandf'ord [1893:426, Type-genus: Phloeo- which occur in Asia and appear to have borus Erichson, 18.36] reached North America in comparatively re- Dactyhpalpi Blandford [1893:426, Type-genus: Dactyli- cent time. The one Hijlesinus in South Amer- palpus Chapuis, 1869] liv- Hylastinides Niisshn [1912b:273, Type-genus: Hi/lasti- ica (antipodius Schedl) is quite similar to nus Bedel, 1888] ing Australian species and was apparently .

1986 WOOD: Genera of Scolytidae 37

derived from Australian stock anciently. This 2(1). Antennal club subconical, elongate, at least 1.5 times as long as wide, sutures straight to feebly suggests that Hylesimis is very old. procurved; eye shorter, less than .3.0 times as The African genera Hylesinopsis and Rho- long as wide (3.3 in Hylastinus], interstriae 10 palopselion appear to be comparatively never represented on posterior third of elytral young, and they are now going through a pe- length; body usually more slender 3 riod of rapid evolutionary change. It could — Antennal club rather strongly flattened, easily be argued that they should be grouped stouter, less than 1.5 times as long as wide, sutures weakly to into a single genus and also that both should rather strongly procurved (if almost straight, then scutellum not visible), be fragmented into many small genera. How- some species appear to have four or more su- ever, it appears that only a small fraction of the tures; eye elongate, at least 4.0 times as long as African species have been discovered. It is wide; interstriae 10 often narrowed and cari- anticipated that when more species are known nate on posterior third of elytra; body usually stouter; Africa 10 most of the apparent character gaps will have 3(2). Frons convex in disappeared, leaving only two major clusters both sexes, not sexually dimor- phic, sometimes marked by a fine, median of species. carina; antennal funicle 6-segmented (except Hijlesinus and Ficicis have been grouped 7-segmented in Hylastinus which lacks prono- into one genus by most workers. However, tal asperities) 4 when all species are examined, it is apparent — Male frons moderately to very strongly im- that two distinct clusters of species are formed pressed, female frons convex (or impressed in Neopteleobins), median carina absent on both anatomical and biological (host or poorly represented; antennal funicle 7-seg- choice) bases. Since these two cannot be com- mented (except 6-segmented in Neoptele- bined without also including a third cluster of obins), pronotal asperities always present .... 6 species (Alniphagus), I have elected to recog- 4(3) Frontal rectangle (Fig. 1) conspicuously longer nize all three as genera. Alniphagus appears than wide; pronotum unarmed by asperities; to be more nearly allied to Ficicis than to antennal funicle 7-segmented; elytral ground Hylesinus vestiture of short, rather stout hair (not sub- plumose); Europe to W Asia, introduced al- The three clusters of genera mentioned most worldwide; herbaceous to shrubby above, together with the comparatively unre- legumes; 2.0-2.5 mm Hylastinus lated Hylastinus, Kissophagus, Pteleobius, Frontal rectangle conspicuously wider than

Cryptocurus, and Neopteleobius , appear to long; pronotum armed on anterolateral areas represent remnants of a previous phyletic ra- by a few fine asperities; antennal funicle 6-seg- mented; elytral ground vestiture diation that took place anciently. Although of either plumose or scalelike setae 5 they cluster conveniently into one tribe, the 5(4). Frons with a fine, long, median carina; elytral extinction of intermediate forms makes com- ground vestiture of abundant, short, plumose ments on phylogenetic relationships difficult setae of uniform color; pronotum more slen- at this time. der, 0.95 times as long as wide; Europe, N

Africa, W Asia; Herfra; 2.0-2.2 mm . Kissophagus Key to the Genera of Hylesinini Frons without a carina; elytral ground vestiture 1. Antennal club subconical to strongly flattened, of abundant scales, their margins entire, usu- with three sutures 6- clearly indicated, funicle ally forming patterns of light and dark color; or 7-segmented (.5-segniented in some Hyle- pronotum stouter, less than 0.85 times as long sinopsis), posterior face of protibia flattened to as wide; Europe, N Africa, W Asia; Ulmus, weakly convex, smooth, punctures with their Sorbus, etc.; 1.8-2.2 mm Pteleobius lateral margins feebly or not at all tuberculate, 6(3). Male frons very strongly, extensively exca- lateral margin armed by socketed teeth; female vated to well above upper level of eyes; male pronotum never with a visible mycetangium; declivity truncate and armed above and below smaller, phloeophagous species, rarely larger by large, blunt processes; apical segments of than 5.0 mm 2 male funicle each ornamented by one or more — Antennal club conical, either without sutures very long, coarse setae; Africa (Nigeria to Tan- or with sutures 1 and 2 rather weakly indicated, ganyika); Piptadenia , 2.3-2.5 mm . . Cryptocurus funicle 7-segmented; posterior face of protibia Male frontal impression less profound, rarely moderately to strongly conve.x, rather coarsely, extending above eyes; declivity convex, one or closely tuberculate, lateral margin without more interstriae sometimes elevated and socketed teeth; female pronotum or pro- armed by smaller tubercles 7 pleuron with a conspicuous mycetangium;

larger, xylophagous, tropical species, 5.0-16.0 7(6). Funicle 6-segmented; male and female frons mm 11 impressed, strongly in male, moderately in fe- 38 Great Basin Naturalist Memoirs No. 10

<-*At- scales, and interstrial rows of longer, erect bris- .^^^^.M|^^Kfl^TiOS^s«t. tles 9 >i

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

I_ North America (Fig. 34); A/iu«; 2.1-3.4 mm . AIniphagus Wk — Male frons moderately to extensively exca- BHhh vated; protibia armed by no more than four socketed teeth, outer apical angle abrupt; an-

tennal club more strongly flattened, its apex WSi ^^^jj^^^^H^^^^^^H ^. ,^ less narrowly roimded; setae in elytral ground ^^Hh^^^^^HM \ mw\£ ^f- vestiture undivided, abundant to absent; Aus- ^^^i^^^^K «#.'?'<-. s^H| tralia to Japan and China; Fictis, rarely other mU'i hosts; 1.6-5.0 mm Ficicis ^^^R if^"'' ^s fei*- 10(2). Pronotum subtriangular, scutellum small to absent; funicle 5-, 6-, or 7-segmented; striae t fi1a'^' M often impressed, punctures usually larger; 1'<'S' Africa; 1.3-3.0 mm Hylesinopsis

— Pronotum subquadrate, scutellum rather IHH large; fimicle 6-segmented; striae usually ^^^Hm- weakly impressed, narrow, punctures fine to obsolete; Africa; 1.5-4.5 mm ... Rhopalopselion W 11(1). Antennal club with two sutures (often obscure); female proepisternum with a large myce- tangium ornamented by hair; metatarsus re- }-\, «i ' ^ traetible into tibial groove; Central and South '^h America; 5.0-16.0 mm Phloeoborus

— Antennal club unmarked by sutures; female pronotum with a median, transverse, slitlike i£ -^^^^X-' myeetangium on anterior third; metatarsus not retractible, tibial groove absent; Africa, SE Fig. 34. AIniphagus aspericoUis (LeConte), dorsal aspect Asia, Philippines; 6.0-14.0 mm . . . Dactylipalpus (After Bright 1976: 207). Hylastinus Bedel [1888:388, Type-species: male, impression not extending above upper Ips obsciiriis Marsham, original designation]. level of eyes; eye shallowly emarginate, elytral Distribution: 2 in Europe, 1 in N Africa; one of ground vestiture scalelike, costal margin near these (obscurus) has been introduced into apex descending; E Asia; 2.2-2.8 mm . Neopteleobius most temperate areas of the world. One is

— Funicle 7-segmented; female frons flat to con- phloeophagous in Cijtisus; one (obscurus) vex; male frons, if strongly concave, with exca- breeds in the roots of various clovers. Keys: vation extending above eyes; eye less strongly Reitter (1913:45). to not emarginate .- 8 Kissophagits Chapuis [1869:34, Type-spe- 8(7). Eye entire, oval, less than 3.0 times as long as cies: Hylesinus hederae Schmitt, monobasic]. wide; protibia armed on lateral margin of apical fourth by six or more closely set, socketed Distribution: 4 in Europe and neighboring teeth; body stouter; declivity more gradual, areas. All are phloeophagous and monoga- abdomen distinctly ascending to meet elytral mous. Key: Reitter (1913:44). apex; elytral vestiture of uniform length, Pteleobius Bedel [1888:392, Type-species: mostly of scales (except almost subglabrous in crenatus), almost worldwide; Fraximis and Bostrichus vittatus Fabricius, subsequent other Oleaceae; 1.7-4.8 mm Hylesinus designation by Hopkins 1914:128]. Distribu- — Eye shallowly emarginate, somewhat elon- tion: 2 in Europe, one of these is recorded gate, at least 3.3 times as long as wide; protibia from N Africa. They are phloeophagous in armed by 2-.5 socketed teeth; body more slen- Ubnus and Sorbus and are monogamous. der; declivity shorter, more abrupt, abdomen horizontal, not rising to meet elytral apex; ely- Keys: Reitter (1913:42), Balachowsky (1949: tral vestiture of ground cover of short hair or 94). 1986 WOOD: Genera of Scolytidae 39

Cryptocurus Schedl [1957c:869, Type-spe- nus Eggers 1922:165, Type-species; Pseudo- cies: Cryptocurus spinipennis Schedl, mono- hylesinus togonus Eggers, automatic; Pseudo- basic. Synonym; Hyloperus Browne 1970: phloeotribus Eggers 1933b: 18, Type- 546, Tvpe-species: Hyloperus bicornis Browne species; Pscudophlocotiihus africanus Eg- = Cryptociirus spinipennis Schedl, original gers, monobasic; Trypographus Schedl designation]. Distribution: 1 in Africa (Nige- 1950e:213, Type-species: Trypographus ria to Tanganyika). It was attracted to light and joveri Schedl, monobasic; Chilodendron was also taken from a Piptadenia buchananii Schedl 1953a:74, Type-species: Chiloden- log. dron planicolle Schedl, monobasic; Glo- Neoptelobius Nobuchi [1971:125, Type- ehicoptcrus Schedl 1954b;75, Type-species; species: Hylesinus scutidatus Blandford, orig- Glochicopterus baphiae Schedl, monobasic; inal designation]. Distribution: 1 species in E Hapalophlocus Schedl 1966b:363, Type-spe- Asia and Japan. Phloeophagous in several cies: Metahylesinus brinckei Schedl, original broadleaf hosts. designation; Hemihylesinus Schedl 1967:224, Alniphagus Swaine [1918:73, Type-spe- Type-species: Hemihylesinus endrocdyi cies: Hylesinus aspericoUis LeConte, mono- Schedl, monobasic]. Distribution; About 38 basic. Synonym: Hylastinoides Spessivitzev species in Africa (Considerable synonymy is 1919:249, Type-species: Hijlastes alni Ni- anticipated). Apparently all are phloeo- isima, monobasic]. Distribution: 2 in W North phagous and monogamous. America, 1 in Japan and E Asia. All are Rhopalopselion Hagedorn [1909:740, phloeophagous in Alnus and are monoga- Type-species; Rhopalopselion bitubercida- mous. Keys: Bright (1976:74) and Wood tum Hagedorn, monobasic. Synonym: Hapa- (1982:119) for North America, logenius Hagedorn 1912:352, Type-species: Hylesinus Fabricius [1801:390, Type-spe- Hapalogenius globosus Hagedorn, monoba- cies: Hylesinus crenatus Fabricius, subse- sic]. Distribution: About 30 African species. quent designation by Westwood 1838:39. Apparently all are phloeophagous and monog- Synonyms: Leperisinus Reitter 1913:41, amous. Type-species: Bostrichus fraxini Panzer = Phloeoborus Erichson [1836:54, Type-spe- Bostrichus varius Fabricius, subsequent des- cies; Phloeoborus rudis Erichson, subsequent ignation by Swaine 1918:70]. Distribution: 10 designation by Hopkins 1914:126. Synonym;

in North America, 1 in South America, 5 in Phloeotrupes Erichson 1836:53, Type-spe- Europe, 9 in Asia, 1 in Africa (Uganda). More cies; Phloeotrupes grandis Erichson, subse- than 20 additional nominate species occur in S (juent designation by Hopkins 1914:127]. Dis- Asia, Australia, and neighboring areas. All are tribution: About 24 species, S Mexico to N phloeophagous and are common in Argentina. All are xylophagous in rather large or other Oleaceae hosts; they are monoga- host material and are monogamous; they are mous. Keys: Wood (1982: lio) for North occasionally attracted to light. They have an America, Murayama (1963:6) for NE Asia. intimate association with fungi, but are not Fields Lea [1910:147, Type-species; Fields mycetophagous. Keys; Blandford (1897:150, varians Lea, subsequent designation by Hop- Eggers (1942:267), Wood (1982:122). kins 1914:122. Synonym: Fieiphagus Mu- Dactylipalpus Chapuis [1869:12, Type- rayama 1958:930, Type-species; Ficiphagus species; Dactylipalpus transversus Chapuis, goliathoides = Hylesinus porcatus Chapuis, subsequent designation by Hopkins 1914: original designation]. Distribution: About 16 120. Synonyms: Dactylopselaphus Gem- nominate species in the area from India and minger & Harold 1872:2678, Type-species; Japan to Australia. All are monogamous and Dactylipalpus transversus Chapuis, auto- phloeophagous in Ficus, rarely in other hosts matic; Ethadopselaphus Blandford 1896:321, (particularly Artocflrpu.s). Type-species; Ethadopselaphus cicatricosus Hylesinopsis Eggers [1920a:40, Type-spe- Blandford, original designation]. Distribu- cies; Hylesinopsis duhitis Eggers, monobasic. tion; 10 African and 2 Philippine and SE Asian Synonyms: Pseudohylesinus Eggers 1919; species. Apparently xylophagous in large host 234, Type-species; Pseudohylesinus togonus material; occasionally attracted to light. Key: Eggers, monobasic, preoccupied; Mefa/i!//esi- Eggers (1933c;200). ,

40 Great Basin Naturalist Memoirs No. 10

Tribe Tomicini ary trend that began in the Hylastini, contin- ued in the Hylesinini, and reached its greatest Tomicidae Thomson [1859: 145, Type-genus: Tomicus specialization in the Tomicini. Latreille, 1802/3] Two major clusters of genera appear within Hylurgini LeConte [1876: 373, Type-genus: Hylurgiis the tribe; first, the group of genera Latreille, 1807] Dendroctonides Niisslin [1912b: 273, Type-genus: Den- (Xylechinus, Chaetoptelius, Xylechinosomus, droctonus Erichson, 1836] Sinophloeus, Denclrotrupes, Hylurgopinus Xylechinides Niisslin [1912b: 273, Type-genus: Xijlechi- Pseudoxylechinus , and Pscudohylesinus) and, nus Chapuis, 1869] second, the Dendroctonus group of genera (, Tomicus, Dendroctonus, Hylur- Description. —Frons very weakly to mod- drectonus , and Pachijucotes). The first group erately sexually dimorphic, male usually im- appears to have radiated in a wide variety of pressed, female convex; eye oval to ovate, hosts from South America and Australia since entire; antennal scape elongate, funicle 4- to the beginning of the Tertiary. The second 7-segmented, club symmetrical, feebly to group appears to have been associated an- moderately flattened, three sutures usually ciently with Araucaria hosts and is sparsely, indicated; pronotum unarmed, except a few uniformly represented in major geographical very small asperities sometimes present in areas, except for Africa, in modern Araucaria

Xylechinosomus and some Xijlechiniis ; pro- and other Pinaceae. coxae contiguous to moderately separated, precoxal lateral costa absent; metascutellar Key to the Genera of Tomicini area separated from postnotum by a distinct 1. Metepisternal setae scalelike or plumose; an- suture; sutural groove on mesal surface of ely- tennal funicle 7-segmented 2 tra continuing to base without a series of inter- — Metepisternal setae usually hairlike (bifid in locking nodules and cavities; tibiae armed by one Hy/iirgonofus having a 6-segmented funi- socketed denticles. cle, and scalelike in Xijlechinus having a 5- segmented funicle); antennal funicle 4- to 7- Biology. —All species are monogamous; all segmented 8 are phloeophagous except for the xylophagous 2(1). Anterolateral areas of pronotum distinctly as- Pachycotes (and Hylurgonotus?). The perate (minute in India species); antennal parental galleries are usually biramous, ex- club apparently with either tivo or four trans-

, cept in Dendroctoniis, Hylurgns and some verse sutures; male frons strongly impressed . 3

Tomicus they are monoramous. Those of — Anterolateral areas of pronotum unarmed; an-

Sinophloeus, Hylurgonotus , and Pachycotes tennal club with three sutures clearly marked; are not known to me. The eggs are placed in male frons impressed or not 4 niches and packed in frass except that some 3(2). Antennal club more strongly flattened, more Dendroctonus have modified the niches into slender, at least 2.0 times as long as wide, apparently with two sutures; frontal rectangle elongate grooves into which numerous eggs at least as wide as long (0.8-1.0 times); Europe are packed in single or double rows. The larval and Asia to Australia and New Zealand; mines usually show on the inner surface of mostly in broadleaf hosts; 1.8-5.0 mm peeled bark and are oriented in a direction Chaetoptelius away from the parental mine. Symbiotic rela- — Antennal club less strongly flattened, stouter, less than 1.5 times as long as wide, apparently tionships with fungi may occur in all genera, with four sutures; frontal rectangle longer but they are adapted toward overcoming re- than wide (about 1.2 times); South America; sistance of the host and are not of a myce- Araucaria; 1.5-3.4 mm Xylechinosomus tophagous type. 4(2). Male frons strongly impressed; median fron- Taxonomy. —The worldwide distribution tal carina present; pronotum either without a of this diversified tribe suggests an ancient constriction affecting dorsal profile (Sino- phloeus) or with moderate lateral constric- origin, although only one Tertiary fossil has a tion on anterior third (Dendrotrupes) 5 been reported (Xijlechinus , Oligocene). The — Male frons convex to modestly flattened, me- most conspicuous division of the group is that dian carina present or absent; pronotum with presented in couplet 1 of the key, except that a conspicuous transverse constriction on ante- Xylechinus should be placed with the first rior third affecting dorsal profile 6 of group genera. The Tomicini represent the 5(4). Antennal club elongate, 2.0 times as long as most highly evolved segment of an evolution- wide, apparently with four or five sutures; 1986 WOOD: Genera of Scolytidae 41

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 M frons feebly to strongly, extensively exca- vated; xylophagous in Araucaria 13

'w 10(9). Antennal funicle 6-segmented, club conical . 11

— Antennal funicle 5-segmented, club moder-

Fig. 35. Xylechinus montanus Blackman, dorsal aspect ately flat 12 (After Bright 1976: 206. 11(10). Procoxae contiguous; pronotum more slen- der, 0.95-1.1 times as long as wide, only declivital interstriae 2 impressed, 3 (some- slightly constricted on anterior thrid; erect tubercles; times 1-7) armed by rounded interstrial setae abundant, confused; a short wider than long pronotum conspicuously (0.7 median carina from epistomal margin to level as wide); larger; South America; times as long of antennal insertion; Europe, W Asia; Finns; Nothophagus 2.3-3.0 mm Sinophloeus 3.1-5.3 Hylurgus

Antennal club more compact, less than 1.5 — Procoxae moderately separated; pronotum three sutures; times as long as wide, with stouter, less than 0.85 times as long as wide, declivity variable; pronotum almost as long as strongly constricted on anterior third; erect hosts not wide (0.9 times); New Zealand; interstrial setae in uniseriate rows (except 1.5-2.0 Dendrotrupes coniferous; mm confused in puellus), a fine median carina

6(4). Elytral vestiture hairhke, ground vestiture from epistoma to middle of frons (absent in moderately stout; frontal carina absent; an- puellus), Europe, Asia, N Africa; 2.5-4.5 mm tennal club slightly flattened, segment 1 one- Tomicus length; America; Ulmus, fourth of its North 12(10), Antennal club with sutures somewhat pro- 2.0-2.5 mm Hylurgopinus curved; procoxae contiguous; male frons con- Vestiture of abundant, conspicuous scales vex to weakly impressed; vestiture never (one exception with no ground setae); median scalelike in either sex; North America, Eu- frontal carina usually present; antennal club rope, Asia; Pinus, Picea, Larix, Pseudotsnga; more conical 7 2.5-9.0 mm Dendroctonus

7(6). Strial punctures small, close; interstriae two — Antennal club with sutures straight, trans- or more times as wide as striae, unarmed verse; procoxae moderately separated; male (except coarsely tuberculate and without frons rather strongly concave; female elytra ground vestiture in rugatus), erect setae with some scales (except araucariae): Aus- closer and coarser; elytral ground setae more tralia, New Guinea; Araucaria; 1.3-1.8 mm slender and appically pointed, variegated pat- Hylurdrectonus .

42 Great Basin Naturalist Memoirs No. 10

13(9). Declivital vestiture hairlike (except scalelike designation]. Distribution: 11 species in in tuberculatus), protibiae with either three North America. Phloeophagous in Abies, or four socketed teeth; male frons rather shal- Picea, Pinus, lowly impressed (except deeper in an- Pseudotsuga, and and tipoditis); South America; Araucaria; 2.7-4.6 monogamous. Kevs: Blackman (1942a; 5),

mm Hylurgonotus Bright (1969: 1.5), Wood (1982: 1.30). — Declivital vestiture always includes scales Xylechinus Chapuis [1869: 36, Type-spe- (sparse in male peregrinus); protibiae with cies: Hylesinus (Dendroctonus) pilosus three socketed teeth; male frons strongly, Ratzeburg, monobasic. Synonyms: Prunipha- more extensively impressed (except weak in peregrimts), Australia, New Zealand, and ad- gus Murayama 19.58: 930, Type-species: jacent islands; Araucaria; xylophagous; 2.3- Pruniphagus gununensis Murayama, original 4.5 mm Pachycotes designation; Squamasinulus Nunberg 1964: 431, Type-species: Squamasinulus chiliensis Chaetoptelius Fuchs [in Reitter 1913: 43, Nunberg, original designation; Xylechinops Type-species: Hylesinus vestittis Mulsant & Browne 1973: Type-species: Rey, automatic. Synonyms: Homarus Broun 283, Xylechinus australis Schedl, original designation]. 1881: 740, Type-.species: Homarus mundidns Distri- bution: 19 species in Central Broun, automatic, preoccupied; Acrantus and South Amer- ica, 2 in North America, 2 in Africa, 5 in Asia, Broun 1882: 409, Type-species: Homonis and 1 in Europe. All are phloeophagous and mundidus Broun, automatic, preoccupied; monogamous. Keys: Blandford (1897: Chaetophorus Fuchs 1912: 46, Type-species: 157), Wood (1982: 143) for North and Central Hylesinus vestitus Mulsant & Rey, monoba- America. sic, preoccupied]. Distribution: 1 species in Hylurgus Latreille [1807; 274, Type-spe- Europe and W Asia, 1 in New Zealand, about 8 in Australia and New Guinea. All are cies: Hylesinus ligniperda Fabricius, monoba- phloeophagous and monogamous. sic]. Distribution: 3 species in Europe, W Xylechinosomus Schedl [1963a: 209, Type- Asia, N Africa. All are phloeophagous in conif- species: Xylecliinus taunayi Eggers, original erous hosts and monogamous. designation]. Distribution: About 9 species in Totnicus Latreille [1802/3: 203, Type-spe- South America. All are phloeophagous in cies: Hylesinus piniperda Fabricius =Der- Araucaria mestes piniperda Linnaeus, monobasic. Syn- Sinophloeus Brethes [1922b: 433, Type- onyms: Blastophagus Eichhoff 1864: 25, species: Sinophlocus pui-teri Brethes, mono- Type-species: Dennestes piniperda Lin- basic]. Distribution: 2 species in South Amer- naeus, preoccupied, subsequent designation ica. Apparently phloeophagous in Notho- by Lacordaire 1866: 360; Myelophilus Eich- fagus. hoff 1878c: 400, Type-species: Dermestes Dendrotrupes Broun [1881: 741, Type-spe- piniperda Linnaeus, automatic]. Distribu- cies: Dendrotrupes costiceps Broun -Den- tion: 8 species in Europe and Asia. All are drotrupes vestitus Broun, subsequent desig- phloeophagous in Pinus and are monoga- nation by Hopkins 1914: 120]. Distribution: 3 mous. Keys: Murayama (1963: 35) for the Far species in New Zealand. Phloeophagous. East, Balachowsky (1949: 135) for France, Hylurgopinus Swaine [1918: 43, 74, Type- Schedl (1946b: 52) for the genus. species: Hylastes rufipes Eichhoft, original Dendroctonus Erichson [18.36: 52, Type- designation]. Distribution: 1 species in North species: Bostrichus micans Kugelann, subse- America. Phloeophagous in Uhnits and mon- quent fixation by International Commission ogamous. on Zoological Nomenclature 1974: 230]. Dis- Pseudoxylechinus Wood & Huang [1986: tribution: 16 species in North America and 2 465, Type-species: Pseudoxylechinus uni- in Eurasia. All are phloeophagous in Pinus, formis Wood & Hwang, original designation]. Picea, Larix, and Pseudotsuga and are Distribution: 7 species in Asia. All are monogamous. Keys: Hopkins (1909: 69), phloeophagous and monogamous in broadleaf Wood (1963: 26, 1982: 151). trees. Hylurdrectonus Schedl [19.3Sb: 40, Type- Pseudohylesinus Swaine [1917: 11, Type- species: Hylurdrectonus piniarius Schedl, species: Pseudohylesinus grandis Swaine monobasic. Synonym: Xylogopinus Schedl = Hylurgus sericeus Mannerheim, original 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. Fig. 36. Phrixosomu magna Blackmail, antenna. Pachycotes Sharp [1877: 10, Type-species: Pacycotes ventralis Sharp =Hylesmus pere- closely allied to Hylesinini than to other grinus Chapuis, monobasic]. Distribution: 8 tribes, this genus is quite unique. species in Australia, New Zealand, and New Phrixosoma Blandford [1897: 148, Type- Guinea. All are xylophagous in logs and species: Phrixoaoma rude Blandford, mono- monogamous. basic. Synonyms: Bothryperus Hagedorn 1909: 742, Type-species: Bothryperus psaltes Tribe Phrixosomini Hagedorn, monobasic; Neohylesinus Eggers Phrixosomini Wood [1978: 111, Tvpe-genus; Phrixosoma 1920b: 118, Type-species: Neohylesinus Blandford, 1897] quadrioculatus Eggers, monobasic; Sphaero- Eggers 1929: Type-species: Description. —Frons not sexually dimor- sinus 40, Sphaerosinus striatus Eggers, monobasic]. phic, frequently with a fine, median carina; in South eye completely divided; antennal scape elon- Distribution: 13 species Central and and 9 in Africa. All are phloeo- gate, funicle 6-segmented, club rather America in Guttiferae are monogamous. strongly flattened, slightly asymmetrical, un- phagous and Key: Wood (1982: 204). marked by sutures, except 1 partly septate; pronotum unarmed by asperities, procoxae Tribe Hyorrhynchini contiguous; scutoscutellar suture remote from scutellar groove, postnotum separated from Hyorrhynchinae Hopkin.s [1915b: 225, Type-genus: Hy- scutoscutellar area of metanotum by a com- orrhynchus Blandford, 1894] Sueinae Murayama [19.58: 7, Type-genus: Siieus Mu- plete suture; tibiae armed on lateral margin rayama, 1951] by socketed denticles. Biology. —These monogamous, phloeo- Description. —Frons sexually dimorphic, phagous, tropical species are restricted to male slightly to strongly impressed, female hosts of the Guttiferae. The parental galleries convex; eye completely divided; antennal are usually biramous, although a third egg scape either long or short, funicle 5- or 6-seg-

tunnel is not uncommon, with the galleries mented, club aseptate and either unmarked either longitudinal, transverse, or without by sutures or with two sutures; pronotum definite orientation. The eggs are deposited in armed or not; procoxae moderately to widely niches and sealed in by frass. The larval mines separated, precoxal ridge obsolete; scu- show on the inner surface of peeled bark, but toscutellar suture remote from scutellar they are almost entirely in the bark and usu- groove; scutoscutellar area separated from ally wander indiscriminantly without respect postnotum by a distinct suture; tibiae without to the grain of the wood. socketed denticles. Taxonomy. —Only one genus is known BiOLCJGY. —The species are myelomyce- (Fig. 36). It is apparently very old and repre- tophagous or xylomycetophagous and form sents a group that is otherwise extinct. They simple, monoramous tunnels in the pith or are unknown in the fossil record. The one xylem of twigs or small branches. Only two genus in South America and Africa in Gut- larvae were seen (Sueus niisimai) and these tiferae has changed so little since separation of were in the parental chamber. Males oi Sueus these land masses that division into species are very rare, dwarfed, and deformed; repro- groups is not recommended. Although more duction in tliis genus is apparently by 44 Great Basin Naturalist Memoirs No. 10 arrhenotocous parthenogenesis. In Hyor- lomycetophagous and monogamous (Nobu- rhtjnchus and Pseudohijorrhtjnchus the males chi, pers. comm.). are similar in size, shape, and abundance to Pseudohyorrhynchus Murayama [1950b: the females, suggesting a normal bisexual re- 61, Type-species: Pseudohyorrhynchus wa- lationship. dai Murayama, original designation]. Distri- Taxonomy. —A dozen species assigned to bution: 1 species from Japan. Specimens of three genera are listed in the literature. All this monogamous species attack living Corniis occur in the area from India and Japan to New twigs where they are xylomycetophagous. Guinea. They form an aberrant, relect group of uncertain affinity but are considered to be Tribe Diamerini among the most primitive of the Hylesininae. Diameridae Hagedorn Although definitely members of the Hylesini- [1909: 734, Type-genus: Di- umerus Erichson, 1836] nae, they share more primitive characters Strombophorini Schedl [1959e: 75, Type-genus: Strom- with primitive Scolytinae and Platypodidae hophonis Hagedorn, 1909] than with other members of their own sub- Sphaerotrypini Murayama [1963: 5, Type-genus: family. Sphaerotrypes Blandford, 1894]

Key to the Genera of Hyorrhynehini Description. —Frons dimorphic, feebly to very strongly impressed in male, flattened to 1. Antennal funicle 5-segmented, club rather weakly compressed; male dwarfed, rare, flight- convex in female; eye entire to completely less; male frons convex, with median carina; divided; antennal scape usually elongate, fu- tarsal segment 3 entire; Sri Lanka (Ceylon) and nicle 6- to 7-segmented, club flattened, su- New Guinea to Japan; female L 6-2.0 mm. male tures variable, one to several (five or more, 1.0-L3 mm Sueus mostly pseudosutures); procoxae moderately — Antennal funicle 6- or 7-segmented, club rather to widely separated; scutoscutellar suture re- strongly flattened; male subequal in size to fe- male, shares parental gallery; male frons con- mote from scutellar groove, postnotum fused cave or not, with or without a carina; tarsal seg- to scutoscutellar area of metanotum, interseg- ment 3 emarginate to bilobed 2 mental suture obsolete on median half. 2(1). Antennal funicle 6-segmented, club with two Biology. —These monogamous, phloeo- clearly marked sutures; male frons concave, phagous species are largely restricted to tropi- without a carina; tarsal segment 3 emarginate; cal and subtropical areas of the eastern hemi- interstrial tubercles minute; India to Japan. 2.7- .5.0 mm Hyorrhynchus sphere. The parental galleries are mostly biramous and either longitudinal or trans- — Antennal funicle 7-segmented, club wdthout su- tures; tarsal segment 3 deeply bilobbed; male verse. The eggs are deposited in niches and frons not impressed, with a median carina; inter- sealed in by frass. The larval mines show on strial tubercles on declivity rather coarse; Japan; the inner surface of peeled bark and tend to 3.5-3.6 mm Pseudohyorrhynchus radiate away from the parental tunnel without

Sueus Murayama [1951; 1, Type-species: respect to the grain of the wood in most spe- Sueus sphaerotrypoides Murayama = Hyor- cies; the latter parts of the tunnels in some rhynchus niisimai Eggers, original designa- species are parallel to the grain of wood. tion. Synonym: Neohyorrhynchus Schedl Taxonomy. —Seven genera represented by 1962e: 202, Type-species, Hyorrhynchus ni- about 122 species occur in Africa and south- isimai Eggers, original designation]. Distri- eastern Asia to Australia. Pseudodiamerus, bution: 2 species from Sri Lanka (Ceylon) and Pernophorus . and Stromhophorus occur ovAy New Guinea to Japan. Xylomycetophagous in in Africa. They appear to form a sister group small branches and, apparently, at least partly derived from the same ancestral stock as Both- parthenogenetic (only one series of males rosternini and, if this is the case, these groups known). have differentiated and radiated since early Hyorrhynchus Blandford [1894a: 58, Type- Tertiary. Features used to characterize this species: Hyorrhynchus lewisi Blandford, tribe vary and tend to intergrade with monobasic]. Distribution: About 10 species Phloeosinini. Although future study may re- from India to Japan. Habits have not been quire combination of these two tribes, their reported except host species include Acer, division gives a convenient break in a large

Fagus , and Macaranga; the species are xy- 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, Fig. 37. Diamerus curvifer Walker, lateral aspect of fe- median anterior area of pronotum usually finely male. asperate; antennal funicle 6-segmented; club elongate, sutures 1 and 2 feebly indicated (straight) to absent; .Mrica, SE Asia to Australia, — Antennal funicle 6-segmented; elytral vestiture with ground cover scalelike; Africa; 1.5-3.5 1. 1-2.5 mm Acacicls mm Strombophortis — Costal margin on basal fourth of elytra either normal or deeply, abruptly emarginate, when Acacacis Lea [1910: 149, Type-species: emarginate metepisternum expanded into this Acacacis ahundans Lea, monobasic. Syn- emargination; pronotum never asperate (feeble onyms: Trogloditica Sampson 1922: 148, lateral asperities in Bothrosternoidcs) .3 Type-species: Trogloditica trahax Sampson, 3(2). Costal margin on basal fourth of elytra deeply monobasic; Pseudoacacacis Schedl 1963c: emarginate (Fig. 37); suture 1 on antennal club strongly, subangulately procurved; Africa, SE 477, Type-species: Acacacis horneensis Asia to Australia; 3.0-.5.0 mm Diamerus Browne, original designation; Neodiamerus

— Costal margin on basal half of elytra almost nor- Schedl 197ib: 282, Type-species: Ncodi- mal, straight to wealily emarginate; basal mar- amcrus granidicoUis Schedl, original designa- gins of elytra either costate or armed by a row of tion]. Distribution: 5 species in Africa, 2 in Sri coarse crenulations; protibia with two to four Lanka, 2 in Malaya, 1 in Borneo, 1 in New curved spines; male frons shallowly impressed; Guinea, and 2 in Australia. All are suture 1 on antennal club weakly indicated (straight) or absent 4 phloeophagous and monogamous. Pseudodiamerus Eggers [1933b: 18, Type- 4(3). Eye more than 3 times as long as wide; funicle

7-segmented; club with suture 1 straight, fee- species: Pseudodiameriis sthatus Eggers, ble; protibia with two apical and two lateral monobasic]. Distribution: 3 species in Africa. spines; costal margin not emarginate; basal mar- All are apparently phloeophagous and monog- gins of elytra crenulate; pronotum without lat- amous. eral granules; Africa (Angola); 2.2-3.0 mm .... Pseudodiameriis Bothrosternoides Schedl [1969: 210, Type- species: Bothrosternoides malaycnsis Schedl, — Eye twice as long as wide; funicle 5-segmented, club unmarked by sutures; protibia with two monobasic]. Distribution: 1 species in apical and one small posterior subapical spine Malaya. Three specimens intercepted in and several lateral posterior granules; costal Japan and 4 in China in imported Nyatoh logs. margin shallowly emarginate at base; basal mar- Diamerus Erichson [1836: 57, Type-spe- gins of elytra strongly costate; some prothoracic cies: Hylesinus hispidus King, monobasic. granules or small asperities in lateral areas; male with median epistomal tubercle; body very Synonyms: Acanthurus Eichhofl 1886: 24, stout; Malaya; 2.3 mm Bothrosternoidcs Type-species: Acanthurus spinipcnnis Eich- - •5(1). Eye completely divided; scutellum visible, hoff Hylesinus curvifer Walker, subsequent longer than wide; pronotum unarmed by asperi- designation by Hopkins 1914: 116; Lissoclas- ties; body very stout, subglobular, Africa, SE tus Schaufuss 1905: 71, Type-species: Lisso- Asia; 1.5-5.0 mm Sphaerotrypes clastus pimelioides Schaufuss, monobasic]. — Eye strongly constricted on lower half to about Distriliution: About 11 species in Africa and one-third width of upper half, pronotum at least 28 in the Indo-Australian and Oriental areas. partly asperate; scutellum not visible; body elongate-oval 6 All are phloeophagous and monogamous.

Sphaerotrypes Blandford [ 1894a: 61, Type- 6(5). Antennal funicle 7-segmented; elytral vestiture hairlike to subplumose, Africa; 4.0 mm species: Sphaerotrypes pila Blandford, subse- Pernophonis ()uent designation by Hopkins 1914: 129. Syn- 46 Great Basin Naturalist Memoirs No. 10 onym: Parasphaerotrijpes Murayama 1958; Ta.xonomy. —The tribe is restricted to the 933, Type-species; Sphaerotrijpes controver- American tropics, with two species extending sae Murayama, original designation]. Distri- into the southern United States. It appears to bution: 8 nominate species in Africa and about be the New World counterpart of the closely 35 in the Indo-Australian and Oriental areas. allied Diamerini. The genera of Bothrostern- Pernophorus Strohmeyer [1910b; 92, ini differ anatomically from one another only Type-species; Acanthophorus hrevicollis slightly and may be recognized with difficulty Strohmeyer, automatic. Synonym: Acan- in some instances. The protibial and antennal thophorus Strohmeyer 1910a; 69, Type-spe- structure and the costate basal margins of the cies; Acanthophorus hrevicollis Strohmeyer, elytra in at least some members suggest a very monobasic, preoccupied]. Distribution; 5 primitive position in classification. Though nominate species in Africa. They are appar- primitive as a group, modern representatives ently phloeophagous and monogamous. appear to have evolved since the beginning of Strombophorus Hagedorn [1909: 740, the Tertiary to their present ecological and Type-species: Strombophorus crenatus structural status, because clearly identifiable Hagedorn, subsequent designation by Hop- related groups are not known outside of the kins 1914: 130]. Distribution; About 34 spe- neotropical realm. They appear to be more cies in Africa. They are phloeophagous and nearly allied to Diamerini and Hyorrhynchini monogamous. than to any American groups.

Tribe Bothrosternini Key to the Genera of Bothrosternini

1. Lateral of 2 Bothrosterni Blaiidford [1896a; 120, Type-genus: Both- margins pronotum rounded rostenius Eichhoff, 1868] — Lateral margins of pronotum marked by a sharply elevated, costate to subcostate line .... 3 Description. —Frons usually sexually di- 2(1). Sutures of antennal club transverse, straight; morphic; eye entire to slightly sinuate; funicle rostrum distinctly wider than distance between 6-segmented, club symmetrical, moderately eyes; pronotum either longitudinally strigose or flattened, sutures indicated; proco.xae moder- punctured; pith borers of twigs and other small ately separated; protibia bearing a bifid pro- stems; SE USA to South America; 1.6-3.5 mm Cnesisnus cess on outer apical angle exceeding inner — Sutures of antennal club strongly procurved; apical angle; pronotum unarmed; crenula- rostrum width at tip equal to distance between tions on basal margins of elytra poorly devel- eyes; frons excavated in both sexes, with a me- oped, sometimes represented by a continuous dian tubercle just above epistoma; body oval; costa; at least part of scutoscutellar suture par- seed borers; SE USA to South America, inter-

cepted elsewhere in maize, etc. ; 1.9-2.6 mm . . allel to margin of scutellar groove; postnotum Pagiocerus fused to scutoscutellar area of metanotum, Sutures of antennal club strongly procurved; suture obsolete. 3(1). pith borers in twigs and woody vines; Central Biology. —All species are monogamous, and South America; 2.1-3.8 mm . . . Eupagiocerus except that a form of parthenogenesis proba- — Sutures of antennal club transverse, straight ... 4 bly exists in Bothrosternus . All are myeloph- 4(3). Proepisternal area partly excavated, with cavity agous, except that Bothrosternus and, possi- densely filled by yellow pubescence, particu- bly, Eupagiocerus ater Eggers are myelo- larly in female; prothoracic intercoxal piece with mycetophagous. In annectens a transverse, subcarinate ridge; elytral inters- strongly carinate; ambrosia Wood typical transverse, biramous egg gal- triae usually not beetles in axial tunnels of woody vines; Central leries are formed in the cambium region of the and South America; 1.9-3.3 mm . . Bothrosternus host by parent beetles, complete with egg — Proepisternal area normal, not densely pubes- larvae briefly in niches, and the feed the cam- cent; prothoracic intercoxal ridge absent; elytral bium region before following the parent interstriae narrowly carinate from posterior part beetles to the pith region. Typically, the of disc to apex; pith borers in branches and twigs; Central and South America; 2.2-3.6 mm parental pith tunnel is biramous; the eggs are Sternobothrus deposted in clusters in the loose frass in this cavity; and the larvae feed in congress while Cnesinus LeConte [1868: 171, Type-spe- extending the parental pith tunnel. Pupation cies; Cnesinus strigicoUis LeConte, monoba- occurs in the larval frass. sic. Synonym; Neinophihis Chapuis 1869; 27, 1986 WOOD; Genera of Scolytidae 47

Fig. 38. Bothrosternini; 1-8, Cnesinus spp. cubensis , (1) 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 Fig. 39. Phloeotiibtis pilula Eiichson, dorsal aspect of and Brazil. Monogamous and some species male. (After Schedl 1953: 80). apparently with a type of parthenogenesis (in- volving consanguineous polygyny), and lanimelate segments; procoxae contiguous; myelophagous in woodv vines. Kev: Wood pronotum armed or not, its lateral margins (1982: 247). rounded; metatergum fused to its postnotum. Sternobothrus Eggers [1943: 372, Type- Biology. —All are monogamous and species: Bothrosternus cancellatus Chapuis, phloeophagous. Parental galleries are bi- original designation]. Distribution: About 10 ramous and engrave the wood rather deeply. species from Costa Rica to Bolivia and Brazil. Eggs are deposited in niches packed in frass. Monogamous and myelophagous, three spe- Larval mines follow a rather definite course cies breed in Nectandra branches. Key: Wood away from the parental tunnels and usually do (1982: 254). not cross one another; in the latter stages they may engrave the wood rather deeply. A few Tribe Phloeotribini species bore rather deeply into subsurface tis- Phloeotribidae Chapuis [1869: 42, Type-genu.s: Phloeo- sues of woody vines; one species breeds in the tribus Latreille, 1796] fruiting pods of its host (Inga). Phthorophloeides Niisslin [1912b: 273, Type-genus: Ph- Taxonomy. The tribe apparently origi- thoropliloeus Rev, 1883] — nated in South America, where a majority of Description. — Frons se.xually dimorphic, the species now occur. Aricerus and one very male variously impressed, female flat to con- primitive Phloeotribiis apparently reached vex; eye entire; funicle 5-segmented, club al- Australia very early; much later a few species most non-existent to strongly asymmetrical, of Phloeotribiis reached North America and deeply divided into three movable, sub- then spread from there to northern Asia, Eu- 1986 WOOD; Genera of Scolytidae 49 rope, and North Africa. The tribe apparently spinulosus Rey, monobasic; Elzearius Guille- was derived from the same parental stock as beau 1893: 64, Type-species: Elzearius crena- the Phloeosinini, although the relationship is tus Guillebeau, monobasic; Eulytocerus not close. The distribution, freely movable Blandford 1897; 161, Type-species: Eulyto- antennal club segments (Fig. 39), and the tib- cerus championi Blandford, monobasic; ial structure of Aricerus and at least one spe- Comesiella Del Guercio 1925: 218, Type-spe- cies oi Phloeotribus suggest an origin in early cies: Comesiella sicula Del Guercio =

Tertiary or late Cretaceous while it was still Bostriclms brevicollis kolenati, monobasic; possible to spread to Australia, but not into Neophloeotribus Eggers 1943: 349, Type-spe- Africa. The one Australian and a few Eu- cies: Phloeotribus nubihis Blandford, desig- ropean Phloeotribus (Fig. 27) have virtually nated by Wood 1983: 648; Dryotomicus Wood no club on the antenna, suggesting the possi- 1962: 76, Type-species: Dryotomus pu- bility that the club of Scolytidae could have berulus Chapuis, automatic]. Distribution: been derived independently from that of About 58 species in South America, 27 in other curculionoids. North and Central America, 10 in Europe, 4 in N Africa, 1 in Australia. All are monoga- Key to the Genera of Pliloeotribiiii mous and phloeophagous. Keys: PfefFer(1972: 1. Lateral margin of protibia without .socketed teeth, 31) for Europe, Blandford (1897: 162) for Cen- outer apical angle rather strongly produced, with tral America, 1982: 257) for North and few (about three) major serrations; lateral margin Wood of pronotum acutely elevated, subserrate; anten- Central America. nal club rather slender, almost symmetrical, ven- tral margins of segment.s not noticeably extended; Tribe Phloeosinini

phloeophagous; Australia to New Guinea; .3, 1-4.6 Phloeosinides Niisslin 273, Type-species: Phloco- mm Aricerus [1912b: ninus Chapuis, 1869] — Lateral margin of protibia with several socketed teeth, none strongly projecting beyond others Description. — Frons usually dimorphic, (one exception); lateral margins of pronotum male impressed, female flat to convex; eye rounded; ventral or lateral margin of each antennal varying from entire to emarginate to com- club segment weakly to profoundly extended into a .sublamellate process except in very primitive pletely divided; antennal iunicle 5- to 7-seg- species; phloeophagous; North and South Amer- mented, club flattened, slightly to strongly ica, Europe, N Asia, N Africa, Australia asymmetrical, with or without sutures; prono- Phloeotribus tum armed or not; tarsal segment 3 com- Aricerus Blandford [1894b: 1.33, Type-spe- pressed to broad and bilobed; scutelhun visi- cies: Aricerus chapuisi Blandford, subse- ble or not; metanotum fused to postnotum. quent designation by Hopkins 1914; 117. Syn- Biology. — All are monogamous except for onym: Hylesinosoma Lea 1910: 143, Type- a few species of polygynous Olonthogaster species: Hylesimis fici Lea =Aricerus eich- and one bigynous Chramcsus . Most are hoffi Blandford, monobasic]. Distribution: 3 phloeophagous, although and species from Australia to New Guinea. three species of Chramesus are xylophagous Monogamous and phloeophagous in Ficus and Hyleops larvae become xylomyce- limbs. tophagous in the later stages. The parental Phloeotribus Latreille [1796: 50, Type-spe- tunnels are mostly monoramous, with a con- cies: Hylesimis oleae Fabricius =Scolytus spicuous turning niche, a few are rather prim- scarabaeoides Bernard, monobasic. Syn- itively (unequally) biramous. Eggs are placed onyms: Phloiotribus Latreille, 1796: 50, inad- in niches and packed in frass. Lai'val mines vertent error in original spelling amended in tend to follow a definite course away from the Latreille 1804: 108, ruled i^y International parental tunnel and rarely cross one another. Commission on Zoological Nomenclature Taxonomy. —This tribe appears to consist 1979: 132; Phloeophthorus Wollaston 1854: of a diverse assemblage of somewhat distantly 299, Type-species: Phloeophthorus perfolia- related genera or clusters of genera that ap- tus Wollaston, monobasic; Dryotomus Cha- pear to be relicts from a former much larger puis 1869: 46, Type-species: Dryotomus pu- group. Their worldwide distribution, diverse berulus Chapuis, monobasic; Phthorophloeus structure, and possession of several primitive Key 1883: 128, Type-species: Phloeophthorus traits suggest an ancient origin that extends )

50 Great Basin Naturalist Memoirs No. 10

'^/'

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 Fig. 40. Phloeosinus punctatus LeConte, dorsal aspect of rather widely separated by distance equal to male. (After Bright 1976: 207). width of coxa; funicle 5-segmented, club sym- metrical to moderately asymmetrical; prono- tum unarmed; scutellum not visible; elytral well into the Cretaceous. Within the tribe, ground vestiture of circular (rounded) scales; two clusters of genera are apparent. Pseu- metathoracic wings apparently absent in male dochramesus and Chramesus (America) are of at least one species; phloeophagous; SE Asia; closely related to one another and are the 2.2-2.5 mm Phloeoditica most highly evolved from a structural point of — Protibia more strongly flattened, armed by view. Cladoctonus, Phloeosinopsoides, Olon- three or more socketed teeth of equal size; with other combination of characters 3 thogaster , and Phloeosinus (Fig. 40) form a second cluster of genera to which Phloeo- 3(2). Antennal club symmetrical to rather weakly asymmetrical (Fig. 42); eye emarginate, or di- cranus (Fig. 41) and Phloeodictica might form vided when funicle 5-segmented; procoxae ei- a primitive base. All are Old World genera ther separated or contiguous; body usually except that about half of the Cladoctonus spe- more slender 4 cies in tropical and a segment occur America — Antennal club very strongly asymmetrical (Fig. oi Phloeosinus has extended into North Amer- 43). sutures strongly procurved when present; ica in comparatively late Tertiary time. The eye entire; funicle 5-segmented; procoxae rather widely separated; body usually very Cladoctonus species of the Philippines (1), stout; American species 11 Africa (8), and tropical America (6) have appar- 4(3). Eye entire; funicle 6- or 7-segmented; prono- ently changed only slightly since attaining tum armed or not 5 their present generic distribution (perhaps — Eye weakly emarginate to completely divided; early Tertiary). The American Dendrosinus funicle 5-segmented (6-segmented in some and Carphotoreus and the Australian Hijleops Cladoctonus}; pronotum unarmed by asperi- are not closely related to one another or to the ties (except weakly armed in Phloeosinop- other known generic groups of this tribe. 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 Fig. 42-43. Antennae of Fhloeosinini: 42, above — Body stout, 1.6 times as long as wide; antennal Phlocosinus thiijae (Perris); 43, below, Chraniesus hicko-

club strongly asymmetrical, suture 1 septate, riae LeConte. others not indicated; posteromesal margin of pronotum slightly extended toward scutellum. color usually reddish brown; SE Asia to New Congo; 1.8 Catenophorus mm Guinea; mostly in nonconiferous hosts; 1.4-5.5 8(4). Antennal club subglobular, sutures obsolete or mm Olonthogastcr indicated by sparse setae; funicle 5- or 6-seg- — Eye less than one-half divided by an emargina- mented; procoxae contiguous; scutellum visi- tion; tarsal segment 3 broad, emarginate to ble or not; phloeophagous; tropical America, bilobed; protibia with three or more socketed Africa, Phihppines; 1.4-2.5 .... Cladoctonus mm denticles on apical and subapical margin, four — Club flattened, sutures transverse to oblique or more smaller teeth on lateral margin; vesti- when present; funicle 5-segmented; proco.xae ture more abundant; male frons usually shal- moderately separated; scutellum visible; lowly impressed, female Irons convex, often phloeophagous 8 with a median carina, vestiture inconspicuous in both sexes; color brown to dark brown; N 9(8). Pronotum finely asperate at least in lateral ar- Africa to N Asia, North America, Australia; eas; sutures 1 and 2 on antennal club trans- mostly in coniferous hosts; 1.4-4.1 verse; eye coarsely faceted, anterior margin mm (PUoeosinites Hagedorn presumably fits near shallowly, broadly emarginate; elytral ground here) Phloeosinus vestiture almost obsolete, erect setae in rows, flattened, almost scalelike, Taiwan to New 11(3). Antennal club with sutures strongly pro- Guinea; 1.3-2.5 mm Phloeosinopsoides curved, clearly marked by rows of setae and grooves; phloeophagous; S America; 1.3-2.0 — Pronotum unarmed; sutures 1 and 2 on anten- Pseudochramesus nal club oblique; eye more finely faceted, mm emargination at least one-third as deep as eye — Antennal club without sutures; mostly 2- width (completely divided in some Asian spe- phloeophagous; North and South America; 1 . cies); elytral ground vestiture usually more 2.7 mm Chraniesus abundant, erect setae slender when ground setae sparse 10 Phloeocranus Schedl [1942: 7, Type-spe- cies: Phloeocranus bruchoides Schedl, mono- 10(9). Eye usually completely divided by an emar- gination (several exceptions); tarsal segment 3 basic. Synonym: Diamerides Browne 1949; slender; protibia with two (rarely one to three) 893, Type-species: Dimnerides litseae apical and subapical socketed teeth, one or two Browne ^Phloeocranus bruchoides Schedl, others sometimes on lateral margin; vestiture original designation]. Distribution: 1 species usually less abundant; male frons usually con-

cave; female frons concave to convex, usually from India to Indonesia in Litsea . This species ornamented by a conspicuous brush of hair; is monogamous and phloeophagous. 52 Great Basin Naturalist Memoirs No. 10

Phloeoditica Schedl [1962d; 189, Type-spe- 29 species in North America, about 30 in Asia cies: Kissophagiis curtits Eggers, present des- and adjacent islands, 5 in Europe, 2 in N ignation]. Distribution: One species in SE Africa, and 2 in Australia. All are monogamous Asia. It breeds in Pongamia glabra. and phloeophagous. Most are in coniferous Dendrosinus Chapuis [1869: 28, Type-spe- hosts, especially Cupressineae. Keys: Black- cies: Hijulesiniis globosus Eichhoff, monoba- man (1942c: 400) and Wood (1982: 287) for sic]. Distribution: 10 species from USA (Flor- North America, Schedl (1950a: 36) for Eu- ida) and Mexico (Jalisco) to Argentina. All are rope, Murayama (1963: 22) for Japan. monogamous and xylophagous. Key: Wood Olonthogaster Motschulsky [1866: 401, (1982: 283). Type-species: Olonthogaster nitidicollis Mot- Hyleops Schedl [1938b: 35, Type-species: schulsky, subsequent designation by Hopkins Hyleops glabratus Schedl, monobasic]. Dis- 1914: 126. Synonyms: H olonthogaster Gem- tribution: 1 species in Australia in Araucaria minger & Harold 1872: 2676, Type-species: branches. It is monogamous and partly Olonthogaster nitidicollis Motschulsky, auto- phloeophagous. The later larval stages pene- matic; Hylediits Sampson 1921: 35, Type-spe- trate the xylem and become xylomyce- cies: Hyledius asper Sampson, monobasic; tophagous. Parental tunnels are transversely Hylitrguliis Eggers 1927c: 392, Type-species: biramous and without apparent symbiotic Hyhirgnhis summatranus Eggers, monoba- hingi. sic; Phloeosinopsis Schedl 1936a: 23, Type- Carphotoretis Wood [1973b: 171, Type- species: Phloeosinopsis annatus Schedl species: Chaetophloeus alni Bright, original = Phloeosinus spinifer Schedl, original desig- designation]. Distribution: 1 species in Mex- nation]. Distribution: About 25 species from ico (Oaxaca). It is monogamous and phloeo- SE Asia to Australia. All are phloeophagous phagous. and monogamous except for 2 polygynous Catenophorus Nunberg [1956b: 195, Type- species from New Guinea. Hosts include Lyt- species: Catenophorus congonus Nunberg, sea. Myristica , etc. original designation]. Distribution: 1 species Phloeosinites Hagedorn [1907: 119, Type- in the Congo. The habits are unknown. species: Phloeosinites relii Hagedorn, subse- Cladoctonus Strohmeyer [1911: 17, Type- quent designation by Hopkins 1914: 126]. species: Cladoctontis affinis Strohmeyer, Distribution: 8 fossil species in Baltic amber monobasic. Synonyms: Hoplites Eggers 1923: (Oligocene). The relationship of this genus to 140, Type-species: Hoplites banostts Eggers, Phloeosinus was not determined. monobasic, preoccupied; Hoplitontus Wood Pseudochramesus Blackman [1939: 87, 1961: 2, Type-species: Hoplites banosus Eg- Type-species: Chraniesus acuteclavatus gers, automatic; Hoplitophthorus Wood 1961: Hagedorn, original designation]. Distribu- 2, Type-species: Hoplitophthorus sentosus tion: 11 species in South America. The habits Wood = Hoplites interriiptus Eggers, original have not been reported. Kev: Blackman designation]. Distribution: 8 species in Africa, (1939: 88). 6 in Cuba to Brazil and Bolivia; 1 in the Philip- Chraniesus LeConte [1868: 168, Type-spe- Islands. The two species for which habits cies: Chraniesus hicoriae LeConte, monoba- are known are monogamous and phloeo- sic. Synonyms: Rhopalopleurus Chapuis phagous. 1869: 46, Type-species: Rhopalopleurus tu- Phloeosinopsoides Schedl [1964c: 317, berculatus Chapuis, subsequent designation Type-species: Phloeosinopsis triseriatiis by Hopkins 1914: 128; Thaumasinidus Reitter Schedl, automatic. Synonym: Phloeosinopsis 1913: 39, Type-species: Dendrosinus bon- Schedl 1964b: 297, Type-species: Phloeo- nairei Reitter =Chraniesus rotundatus Cha- sinopsis triseriatiis Schedl, original designa- puis, monobasic; Prochramesus Wood 1956b: tion, preoccupied]. Distribution: About 8 254, Type-species: Prochramesus anneetans species from Taiwan to New Guinea. All are Wood, original designation]. Distribution: 39 monogamous and phloeophagous. species in South America, 40 in North and Phloeosinus Chapuis [1869: 37, Type-spe- Central America and adjacent islands. All are cies: Hijlesinus thiijae Perris, subsequent des- monogamous, except for the bigynous C. in- ignation by Hopkins 1914: 126]. Distribution: 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 (Fig. 44), which is almost worldwide in the three xylophagous species. Keys; Blackmail warm areas, the remaining genera are of lim- (1938c; 536), Wood (1982; 316). ited distribution. One is American, 1 Aus- tralian, 2 Madagascaran, and 4 African of Tribe Hypoborini (1 these reaches nearby areas of Europe and Asia Hypoborinae NiissHn [1911: 376, Type-genus: Hy- in cultivated fig). It appears to be a relict poborus Erichson, 1836] group that once enjoyed much greater distri- Description. Frons dimorphic or not, — bution and diversity than at present. male impressed, female convex or less The American genus, Chaetophloeus , is strongly impressed, except in some Li- quite different structurely and biologically species is parthrum this feature reversed; eye from the remaining closely related genera. entire; funicle 3- to with 6-segmented, club Members of this tribe apparently prefer arid up to three sutures, absent; sometimes prono- or semiarid areas or habitats and tend to be variously in restricted areas; pro- tum armed rare. Since they breed in shrubs or small trees coxae contiguous; tarsal segment 3 narrow; of marginal economic importance, it is sus- scutellum not visible; crenulations on elytral pected that a majority of the species await bases not continued laterad from interstriae 5; discovery. postnotum fused to metatergum (remnants of

suture sometimes visible). Key to the Genera of Hypoborini Biology. —All are monogamous and 1. Funicle 6-segmented, club small, conical, with phloeophagous. In all except Chaetophloeus two straight sutures; Australia; Acacia, 1.3 mm parental the gallery is a simple, oval cave. Zygopliloeus Eggs are packed in frass in niches on the mar- — Funicle with 3- to 5-segments; club flattened. gins of the central cave (Liparthrum) or of the sutures present or absent 2 egg tunnels (Chaetophloeus). Larval mines 2(1). Protibia strongly flattened, rather broad, lateral radiate out from the parental chamber and apical half armed by a row of 7-10 closely set, rarely cross one another; they are visible on socketed teeth; pronotal asperities confined to the inner surface of peeled bark. two or three paired clusters on lateral thirds, each cluster containing 1-5 denticles; funicle 5- Taxonomy. This tribe is sparsely and — segmented, club clearly marked by three su- widely distributed in the warm climates tures; phloeophagous; North and South Amer- around the world. Except for Liparthrum 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- rather widely spaeed, socketed teeth; four species; Hijlesinus hystrix LeConte, monoba- pronotal asperities mostly on median third, . „ n /-^ mn/- r,r-n ^ic. Synonyms: Renocis Casey more abundant; funicle 3- to 5-segmented 3 1886; 257,

„,„, • Tvpe-species; Renocis , , 1 r 1 c .. J I u .^u 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 • North America and adjacent islands, m 1.5 mm Glochiphorus 2

„ ., . , South America. All are monogamous and — Protibia not armed on outer apical.111angle by a , , , ,^ /,r^<^ ^^r<\ phloeophagous. keys: mi major spine; antenna variable; elytra bearing 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-

sutures; basal . mount), club unmarked by mar- i i .- c r - 1 1- f , ^ ^. 1 1 ton, original designation. Synonyms: Lei- gins ot elytra a continuous costa, individual i n i i crenulations feebly indicated; anterior half of parth 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, 1 • n? 7 j .1 o 1 11 mi lulus but less conspicuous 6 Blackman, monobasic; Fluoeoc Schedl ct-v iri^„ viff 1953c; 292, Tvpe-species: Phloeochilus pala- 6(5). Funicle 4-seginented,tjiijclub 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 j . Distribution: 2 ilattened, much wider and more coarsely serrate ' o o 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 mous andi phloeophagous.i i i 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. Fig. 45. Carphoborus pirucolens Wood, dorsal aspect of female. (After Bright and Stark 1973: 155). Biology. —All are phloeophagous. Car- phobius and some Polygraphits are monoga- mous; apparently Chortastus and Serrastus the one known species of Phloeographus share this habit; Carphoborus (Fig. 45) and bears a remarkable superficial resemblance to most Polygraphus are polygynous. Parental certain Tomicus species (Hylesinini). tunnels are monoramous or biramous in The genera form a rather compact unit monogamous forms and radiate in polygynous without conspicuous divisions. Four small forms. The nuptial chamber is unusually large genera are exclusively African (Serrastus, in most species. Eggs are packed in frass in Chortastus, Phloeographus, Cardroctonus), niches. Larval mines wander considerably one is Malayan (Bothinodroctonus), and one and have a greater tendency to cross one an- is American (Carphohius). The larger genus other than in most other tribes. Carphoborus is primarily North American Taxonomy. —In my initial study of the and probably originated there from stock tribes of Scolytidae (Wood 1978), the Poly- derived from Asia. European and Asian spe- graphini were reluctantly given tribal status cies of Carphoborus appear to have been only after much hesitation and the pondering derived from the more primitive and diverse of many questions. Since then, all reserva- American fauna. The largest genus, Polygra- tions have been dismissed. Although quite phus , is largely African, with strong represen- specialized in several respects, representa- tation in Europe and Asia, and with three tives of two genera (Serrastus, Polygraphus) species in northern North America. The ab- have the outer apical angle of the protibia sence of this group in South America suggests produced beyond the tarsal insertion and are either an origin since early Tertiary, if Africa armed as in primitive representatives of other were the site, or else an origin in tropical unspecialized tribes. The tribe generally ap- Eurasia. Polygraphus reached North America pears to have been derived from the same in comparatively recent time, Carphoborus ancestral stock as the Phloeosinini, but and Carphobius much earlier. Eurasian Car- 56 Great Basin Naturalist Memoirs No. 10

phohorus were probably derived from a sec- small, more nearly conical; vestiture confined to a few interstrial setae on and near declivity; ondary radiation that moved from North phloeophagous; India to Malaya; 1.8-2. .5 mm . . America to Asia. Carphohius apparently rep- Bothinodroctonus resents a relict from a very early ancestral 7(3). Antennal club symmetrical, with three trans- stock not now represented elsewhere. verse sutures indicated; vestiture abundant, hairlike; phloeophagous; North America; 1.6- Key to the Genera of Polygraphini 2.1mm Carphohius

1, Basal margins of elytra armed by a continuous — Antennal club rather strongly asymmetrical,

costa; protibia very slender, its median apical part of suture 1 indicated, strongly oblique; mucro bent laterad, about one lateral denticle vestiture usually confined to declivity, scalelike;

present, posterior face asperate; funicle 5-seg- 5-.5. phloeophagous; Africa; 2.. 3 mm . . . Chortastus mented; club strongly flattened, moderately asymmetrical, sutures not indicated; phloeo- Serrastus Nunberg [1969: 392, Type-spe- phagous; Africa; 1.8-2.0 Serrastus mm cies: Serrastus ivoriensis Nunberg =Chor- — Basal margins of elytra serrate, armed by a row tastiis similis Eggers, monobasic]. Distribu- of crenulations; protibia more broadly flattened, tion: 2 species in Africa (Ghana to Zaire). They lateral margin armed by several socketed denti- are and phloeophagous. cles 2 monogamous Phloeographus Wood [1984: 229, Type- 2(1). Funicle 5-segmented (.5- or 6-segmented in

Polygraphus , if 6-segmented then eye com- species: Pliloeographus mamibiae Wood, pletely divided); male frons armed by a median original designation]. Distribution: 1 species pair of tubercles near upper level of eyes; vesti- in Africa (apparently Namib Desert, SW ture of abundant scales (except glabrous in some Africa). Bothinodroctonus) 3 Cardroctonus Schedl [1966b: 361, Type- — Funicle 6-segmented; eye never divided into two parts; male frons unarmed by median tuber- species: Cardroctonus orientalis Schedl, orig- cles; vestiture hairlike or sparse if scalelike .... 7 inal designation]. Distribution: 2 species in

3(2). Eye completely divided by a deep emargination Africa. (some Asian and Alrican species only emar- Polygraphus Erichson [1836: 57, Type-spe- ginate); antennal club rather strongly flattened, cies: Hylcsiuus pubescens Fabricius =Der- asymmetrical, devoid of sutures; phloeo- mestes poligraphus Linnaeus, monobasic. phagous; North America, Asia, Europe, Africa; 1..5-.5.5 mm Polygraphus Synonyms: Lepisomus Kirby 1837: 193, Type-

' — Eye less than half divided by an emargination; species: Apate (Lepisomus) rufipennis Kirby, antennal club symmetrical (or nearly so), clearly subsequent designation by Hopkins 1914: marked by sutures 4 124; Spongotarsus Hagedorn 1908: 372, 4(3). Sutures of antennal club procurved; vestiture Type-species: Spongotarsus quadrioculatus sparse, in uniseriate rows on interstriae; tropical Hagedorn, monobasic; Pseudopolygraphus Africa 5 Seitner 1911: 105, Type-species: Polygraphus — Sutures of antennal club straight or nearly so; grandiclava Thomson, subsequent designa- vestiture much more abundant; northern' hemi- sphere 6 tion by Hopkins 1914: 128; Ozophagus Eggers 1919: 234, Type-species: Ozophagus canier- 5(4). Antennal club with three weakly procurved su- tures indicated, ape.\ strongly acuminate; vesti- unus Eggers, subsequent designation by ture sparse, of uniseriate rows on declivity; de- Wood 1982: 386; Urdugraphus Beeson 1941: clivital interstriae 2 .slightly impressed, 301, nomen nudinn]. Distribution: 3 species unarmed, 1 and 3 armed by small tubercles; SW in North America, about 45 in Asia and adja- Africa; 2.2 mm Phloeographus cent islands, 6 in Europe, about 45 in Africa — Antennal club with suture 1 partly septate, 1 and Madagascar. Most are polygynous, al- weakly, 2 moderately procurved; strial punc- though monogamy occms in some Asian and tures very fine; Africa; 1.3-2.0 mm . Cardroctonus African species. All are phloeophagous. Keys: 6(4). Male frons shallowly impressed below upper level of eyes, female usually convex; vestiture of Schedl (1955a: 5) for Europe and part of Asia, abundant scales; antennal club moderately Murayama (1956: 278) for Japan, Wood (1982: large, flattened; phloeophagous; North Amer- 387) for North America. ica, N Asia, Europe, N Africa; 1.4-2.6 mm .... Carphoborus Carphoborus Eichhoff [1864: 27, Type- species: Hylesinus minimus Fabricius, mono- — Male frons profoundly excavated from eye to eye from epistoma to well above eyes, female simi- basic. Synonym: Estenoborus Reitter 1913: larly but less strongly excavated; antennal club 58, Type-species: Hylesinus perrisi Chapuis, .

1986 WOOD: Genera of Scolytidae 57 monobasic]. Distribution: 21 species in North Five clusters of relationship are recognized America, 14 in Europe and Asia, 2 in N Africa. within the subfamily. Progressing from primi-

All are polygvnous and phloeophagou.s. Keys; tive to the more specialized, these include: (1) PfefFer (1914: 169) for Europe, Balachowskv Ctenophorini, Scolytini, and Scolytoplatypo-

(1949: 143) for France, Wood (1954a: 507, dini: (2) Micracini and Cactopinini; (3) 1982: 372) for North America. Carphodicticini, Ipini, Dryocoetini, Cryp-

Bothinodroctonus Schedl [1969: 208, Type- turgini, Xyloterini, and Xyleborini; (4) Xyloc- species: Bothinodroctoniis bicinctus Schedl, tonini and Cryphalini, and (5) Corthylini. Of monobasic]. Distribution: 4 species from In- these, the Corthylini are by far the most dis- dia to Borneo and China. One species is tinctive from an anatomical point of view. phloeophagous in Odina Carphobius Blackman [1943c: .398, Type- Tribe Scolytini species: Carphohius arizonictis Blackman, Scolvtarii Latreille [1807: 27.3, Tvpe-^enus: Scohittis Ge- original designation]. Distribution: 3 species oftVoy, 1762] in North America (Arizona to Guatemala). All Camptoceridae Chapuis [1869: 49, Type-genus: Campto- are monogamous and phloeophagous in cents Latreille, 1829] . Key: Wood (1982: 369). Description. —Frons usually strongly di- Chortastus Schaufuss [1905: 15 (reprint p. morphic, male variously impressed, female 8), Type-species: Choi-tasttis cameninus flat to convex, one or both often ornamented Schaufuss, monobasic. Synonym: Afrochram- by hair; posterior face of head truncate; eye esus Schedl 1971a: 197, Type-species: oval, entire; scape short to elongate, fuiiicle Afrochramcsus bafi,nenai Schedl, original 7-segmented, club rather large, flattened, su- designation]. Distribution: 7 species in Africa. tures present or not; pronotum unarmed, its Apparently all are monogamous and phloeo- lateral margins costate; protibia (usually all phagous. three tibiae) unarmed on lateral margin, outer Subfamily Scolytinae apical angle extended into a spine curving toward and exceeding inner apical angle, Scolytarii Latreille [1807: 27.3, Type-genu.s: Scohjttis Ge- socketed denticles never present; meta- offroy, 1762] pleural suture descending subvertically to In previous classifications, this subfamily groove receiving groove on costal margin of has been divided into two or more major divi- elytra then turning abruptly and continuing sions equivalent to subfamilies. However, it parallel to groove to near metacoxal process; appears that this unit contains both the most in two genera venter of abdomen ascends con- primitive elements of the family that are most spicuously to meet apex of weakly declivous closely related to primitive Platypodidae and elytra. also the main evolutionary thrust of the platy- Biology. —All are monogamous except for podid-scolytid group. This diversity makes a few neotropical and one European bigynous

characterization of the subfamily ratlier diffi- Scolytiis . Camptocerus species are xylomyce- cult. tophagous; those in the other three genera are Members of this subfamily have the bases of phloeophagous. Parental galleries are bi- the elytra simple, forming a straight, trans- ramous, except for a few that are mono- verse line across the body. A large, flat scutel- ramous. In Camptocerus , a biramous, trans- lum is usually visible. The elytral bases are verse tunnel without niches is usually made in weakly subcostate in some Scolytini, the cambium region, then a radial egg tunnel

Ctenophorini, and Cryphalini. The body (sometimes branched) is extended from it into tends to be more elongate, with specialization the xylem. Eggs are deposited in niches in this directed toward cylindrical compaction and radial tunnel. The larval mines of phloeo- the xylomycetophagous habit. A broadly oval phagous species follow a definite course and body outline is usually confined to primitive, rarely cross one another. Camptocerus larvae phloeophagous genera. Armed elytral bases enlarge the egg niche into a cradle just large that suggest a relationship to the Hylesininae enough to accommodate the mature beetle, occur in Ctenophorini (Cnemonyx, allies of somewhat similar to some other ambrosia galeritus). 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- Fig. 46. Scolytopsis inincticoUis . female. Note emar- ginate costal margin of elytron. pear more primitive and, in some respects, overlap the other three.

Scolytits appears to have reached North segment 2; phloeophagous, tropical; Mexico

America from South America by the begin- (Oaxaca) and Cuba to Argentina; 1.6-3.5 mm . . Scolytopsis ning of the Tertiary, when a secondary radia- tion occurred. Elements of this radiation then — Costal margin of elytra normal (straight) and overlapping metepisternum; abdomen flexed reached Eurasia, where another radiation oc- upward from anterior margin of segment 2; curred that was based on a progressively re- phloeophagous; North and South America, Eu- duced gene pool. The number of Eurasian rope, N Asia, N Africa; 1.3-5.5 mm Scolytus species in this genus now may equal or exceed Cnemonyx Eichhoff [1868a: 150, Type- the American component, although the species: Cnemonyx galeritus Eichhoff, anatomical and biological diversity there is monobasic. Synonyms: Ceratolepis Chapuis greatly reduced. 1869: 52, Type-species: Ceratolepis jiicundus

Key to the Genera of Scolytini Chapuis, monobasic; Loganius Chapuis 1869: 52, Type-species: Loganius flavicornis Cha- 1. Scutellar area of interstriae 1 not depressed, puis, monobasic; Minuhis Eggers 1912b: 206, scutellum flush (even) with elytral bases; basal margins of elytra with a fine raised line {some Type-species: Minulus harhatus Eggers, Cnemontjx with crenulations instead), outline of monobasic; Coptodryas Schedl 1948b: 262, anterior margins form a continuous, straight, Type-species: Coptodryas hylurgoides with ventral transverse line scutellum; profile of Schedl. monobasic; Coptosomus Schedl abdomen ascending gradually 2 1952a: 363, Type-species: Coptodryas hylur- — Scutellum depressed, subtriangular, apically goides Schedl, automatic]. Distribution: 19 (posteriorly) pointed; elytral bases depressed in scutellar area, appearing emarginate in median species in USA (Florida), Mexico, and Central area; ventral profile of abdomen usually ascend- America, about 27 in South America and adja- ing abruptly at segment 2 3 cent islands. All are monogamous and 2(1). Antennal club usualK" with two or three sutures phloeophagous. Keys: Blandford (1896d: 128) clearly marked by setae; scutellum small, longer for Central America, Wood (1982: 394) for than wide, often convex; apical liiargin of meso- North and Central America. and metathoracic tibiae commonly bearing tu- bercles on anterior edge in addition to inner and Camptocerus Latreille [1829: 91, Type- outer apical spines; usually inore coarsely sculp- species: Hylcsiints aeneipennis Fabricius, tured; phloeophagous; USA and Mexico to Ar- monobasic]. Distribution: 19 species from gentina; 1.0-3.9 mm Cnemonyx Central and South Ainerica. All are monoga- — Antennal club with suture 1 (only) marked inter- mous and xylomvcetophagous. Key: Wood nally by a partial septum; scutellum flat, 1.5 or 1982: 412 for Central America. more times as wide as long; meso- and metatho- racic tibiae acutely margined on apical anterior Scolytopsis Blandford [1896d: 120, 123, edge, without supplemental denticles; usually Type-species: Scolytopsis puncticollis Bland- very finely sculptured; xylomycetophagous; ford, monobasic]. Distribution: 7 species from Central and South America; 2.6-8.5 mm Cuba and Mexico (Oaxaca) to Argentina. All Camptocerus are phloeophagous and monogamous. Key: 3(1). Basal portion of costal margin of elytra deeply, Wood (1982: 417) for Central America. broadly excised, metepisternum conspicuously expanded into this notch (Fig. 46); abdomen Scolytus Geoffrey [1762: 309, Type-spe- abruptly flexed upward at posterior margin of cies: Bostrichus scolytus Fabricius, desig- 1986 WOOD: Genera of Scolytidae 59 nated by the International Commission on Description. —Frons usually dimorphic, Zoological Nomenclature (China 1962: 3). male impressed and female flat to convex in

Synonyms: Ekkoptogaster Herbst 1793: 124, Pycnarthrum and Gymtiochilus , sexual differ-

Types-species: Bostrichus scoh/ttis Fabricius, ences obscure in Microborus , male convex designated by Hopkins 1914: 121; Copto- and female variously sculptured and orna-

gaster Illiger 1807: 321, Type-species: mented in Scolytodes ; posterior face of head Bostrichus scolytus Fabricius, designated by truncate; eye usually elongate, entire to sinu- Hopkins 1914: 118; Eccoptogaster Gyllenhal ate; scape elongate, funicle 6- or 7-seg- 1813: 346, an isotypical emendation oi Ekkop- mented, club with or without sutures; prono- togaster Herbst; Scolytochchis Reitter 1913: tum armed or not, its lateral margins costate; 23, Type-species: Ips multistriatus Marsham, procoxae widely separated; protibia with one designated by Wood 1982: 419; Riigitloscohj- or more socketed denticles on lateral margin, tus Butovitsch 1929: 20, Type-species: spine on lateral apical angle usually extending Bostrichus rugulosus Miiller, designated by beyond level of tarsal insertion; pleural suture Wood 1982: 419; Archaeoscohjtus Butovitsch about as in Scolytini. 1929: 21, 23, Type-species: Scohjtus claviger Biology. —All are monogamous, except for

Blandford, monobasic, not a genus-group a few polygynous Scolytodes . All are phloeo- name, no status; Spinuloscolytus Butovitsch phagous, except for the xylophagous Scoly- 1929: 21, 24, Type-species; Ips multistriatus todes multistriatus Wood and species that in- Marsham, present designation, not a genus- fest Cecropia leaf petioles. Parental galleries group name, no status; Tubuloscohjtus Bu- vary from a simple to an elongate cave to tovitsch 1929: 21, 33, Type-species: Eccopto- stellate in Scolytodes ; they are biramous in gaster intricatus Ratzeburg, present desig- Pycnai-thrum and Gymtiochilus and indefi- nation, not a genus-group name, no status; nite, nondirectional, and without definite pat- Butovitsch 1929: 28, Pygrnaeoscolytus 21, tern in Microborus. The eggs may be scat- Type-species: Bostrichus Fabri- pygmaeus tered loosely in the parental chamber or cius, present designation, not a genus-group placed in crude niches in Scolytodes ; definite name, no status; Pinetoscohjtus Butovitsch niches are formed in Pycnarthrum and Gym- 1929: 22, 48, Type-species: Scolytus nochilus; they were not observed in Mi- marawitzi Semenov, monobasic, not a genus- croborus. The larvae usually feed commu- group name, no status; Confusoscolytus Tsai, nally in Scolytodes; they form individual Yin, Huang 1962: 4, 14, Type-species: Ec- & mines that follow a somewhat definite direc- coptogaster confusus Eggers, monobasic]. tion in Pycnarthrum and Gymnochihis ; they Distribution 28 species in North and Central are individual and without a definite direction America, about 29 in South America, 47 in in Microborus . Symbiotic relationships with Asia, Europe, and Africa. Monogamous, N fungi were not observed. except for 1 European and about 20 Central Taxonomy. —The tribe is restricted to the and South American bigynous species, and all American tropics, except that Microborus are phloeophagous. Keys: Blackman (1934: 6) boops Blandford was introduced into tropical for North America, Schedl (1937a: 156) for Africa. Scolytodes and Microborus are closely South America and (1948a: 4) for Europe, related to one another. Pycnarthrum and Michalski (1973; 137) for Europe and Asia, Gymnochilus are remotely related to those Wood (1982: 420) for North and Central genera and to one another. Pycnarthrum America. could easily be placed in Hylesinini. This Tribe Ctenophorini tribe occupies a position intermediate be- tween the Scolytini and the more highly Ctenophoridae Chapuis [1869: 49, Type-genii.s: evolved tribes in this subfamily. In all Ctenophorus Chapuis, 1869 =Scohjtodes Fer- mem- rari, 1867] bers the outer apical angle of the protibia Probleehilidae Eichhotf [1878a: .34, 46, 167, 298, projects beyond the tarsal insertion, a primi- = Type-genus: Prohhchilus Eichhoff, 1878 tive feature shared by primitive members of GymnochiluH Eichhoff, 1867] several other tribes. It is the Ctenophorini Hexacohdae Eichhoff [1878a: .3.5, 57, .306, Type- genus: Hexacolufi Eichhoff, 1868 =Scolytodes protibia, not the type found in Scolytini, that Ferrari, 1867] appears to resemble the ancestral type of all 60 Great Basin Naturalist Memoirs No. 10 scolytids. The eye shape, usually unarmed ica, 1 (boops) was introduced into tropical pronotum, presence of interstriae 10, simple Africa. All are monogamous and phloeo- sculpture of the elytra, the elytral locking phagous. Key: Wood (1982: 452) for Central mechanism, and diversity of habits all suggest America. that, when considered as a whole unit, this is Pycnarthrum Eichhoff [1878a: 41, 104, probably the most primitive of all of the tribes Type-species: Pycnarthnim gracile Eichhoff of Scolytidae. It is clearly of neotropical origin = Hypoborus (?) hispidus Ferrari, subsequent and has spread into southern North America designation by Hopkins 1914: 128. Synonyms: only recently. This phyletic line is repre- Nemobius Chapuis 1869: 41, Type-species: in the Scolytoplatypo- sented Old World by Nemobiits larnbottei Chapuis =Hypoborus (?) dini, a group that has diverged significantly in hispidus Ferrari, subsequent designation by both structure and habits. Hopkins 1914; 128, preoccupied; Monebius Hopkins 1914: 125, Type-species: Nemobius Key to the Genera of Ctenophorini larnbottei Chapuis =Hypoborus (?) hispidus 1. Eyes elongate, approximate above and below, Ferrari, automatic; Nomebius Navas 1915: 34, coarsely faceted, shallovvly emarginate; entire Type-species: Nemobius lambottei Chapuis surface of pronotum smooth and punctured, not =Hypoborus hispidus Ferrari, automatic]. armed 2 (?) Distribution: About 15 species in USA (S — Eye oval, enitre, finely faceted; pronotum as- Florida) and Mexico to Brazil. All are monoga- perate anteriorly or, if smooth, then anterior

margin of elytra bearing a fine, raised line .3 mous and phloeophagous. Key: Wood (1982:

2(1). Antennal club subglobular, about as long as 455) for North and Central America. wide, sutures not clearly indicated; pronotum Gymnochihts Eichhoff [1868b: 399, Type- longer than wide, its lateral margins straight or species: Gymnochihis zonatus Eichhoff, feebly constricted; vestiture hairlike, usually monobasic. Problechilus Eichhoff sparse; small, slender species; phloeophagous; Synonyms: Central and N South America, Jamaica, Africa; 1878a: 46, 167, Type-species: Gymnochilus 1.1-1.5 mm Microbonis zonatus Eichlioff, automatic; Meringopalpus — Antennal club asymmetrically flattened, Hagedorn 1905: 547, Type-species: Meringo- pointed, at least 1.5 times as long as wide, su- palpus fallax Hagedorn =Gymnochihis zona- tures 1 and 2 clearly marked by setae; pronotum tus Eichhoff, monobasic]. Distribution: wider than long, its lateral margins arcuate; About 14 species in Mexico to Brazil and Bo- vestiture of abundant, short, bristlelike scales; larger, stouter species; phloeophagous; Florida livia. All are monogamous and phloeo-

and Texas to Brazil; 1.3-2. 1 mm .. , . Pycnarthrtim phagous. Key: Wood 1982: 462) for North and

3(1). Antennal funicle 7-segmented, club large, Central America. broad, usually with procurved sutures or su- Scolytodes Ferrari [1867: 77, Type-spe- tures obsolete; elytral vestiture consisting of cies; Scohjtodes laevigatus Ferrari, mono- abundant, minute hair and sparse interstrial rows of long, erect, scalelike bristles; summit of basic. Synonyms: Hexacohis Eichhoff 1868b: pronotum on basal third, asperities on anterior 399, Type-species: Hexacohis glaber Eich- area coarse; elytral base without a fine, raised hoff^, monobasic; Ctenophorus Chapuis 1869: line; phloeophagous; Mexico (Puebla) to Brazil 49, Type-species: Ctenophorus laevigatus and Bohvia; 1.5-2.3 mm Gymnochilua Chapuis =Hexacolus levis Blackman, mono- — Antennal funicle 6-s6gmented, club small, su- basic; Prionosceles Blandford 1897: 177, tures present or not; elytral vestiture sparse

(usually), hairlike; pronotal asperities fine, if Type-species: Prionosceles atratus Blandford, present; summit at middle or indefinite; basal subsequent designation by Hopkins 1914: margins of elytra marked by a fine, raised line; 128; Epomadius Blandford 1897: 179, Type- mostly phloeophagous; Florida and Mexico to species: Epomadius culcitatus Blandford, Argentina; 0.9-3.5 mm Scolytodes monobasic; Erineophilus Hopkins 1902: 34, Microborus Blandford [1897: 175, Type- Type-species: Erineophilus schwarzi Hop- species: Microborus boops Blandford, mono- kins, original designation; Hijlocurosoma Eg- basic. Synonym; Pseitdocryptiiifius Eggers gers 1940: 138, Type-species: Hylocurosoma 1919: 236, Type-species: Pseudocrijptur

Scolytoplatypini Blandford [1893: 425, Type-species: lomycetophagous. Keys: Browne (1971: 113) Scolytoplatypus Schaufuss, 1890] for Africa, Schedl (1975: 219) for the genus. Taeniocerini Blandford [1893: 428, Type-genus: Taenio- cerus Blandford, 1893 = Scolytoplatypus Schau- Tribe Micracini fuss, 1890]

Spongocerinae Hagedom [1909: 162, Type-genus: Spon- Micracides LeConte [1876: .346, :367, Type-genus: Mi- gocerus Blandford, 1893 = Scolytoplatypus Schau- cracis LeConte, 1868] fiiss, 1890] Hylocuridae Eichhoff [1878a: 298, 306, Type-genus:' Hy- locurus Eichhoff, 1872] Description. —Frons dimorphic, male concave, female convex; posterior face of head Description. —Frons usually dimorphic; truncate; eye oval, entire; scape elongate, fu- either sex may be variously impressed, sculp- nicle 6-segmented, club flattened, without tured, or ornamented by setae, female frons sutures; pronotum unarmed, usually con- often concave, male frons rarely concave (two stricted on posterior half, female with median species); dorsomedian occipital area usually mycetangium; procoxae widely separated; extended slightly caudad; eye oval to elon- protibia with outer apical process prominent, gate, entire to sinuate; scape very short to recurved, outer margin without socketed elongate, strongly flattened to slender, orna- denticles; scutellum not visible, except small mented or not; funicle 6-segmented (5-seg- in one species; basal margins of elytra usually mented in Miocryphaliis , Africa), club with or slightly elevated along a continuous costate without sutures; pronotum asperate on ante- line. rior half, lateral margins rounded; protibia Biology. —All are monogamous and .xy- usually with sides parallel, socketed denticles lomycetophagous. Eggs are deposited in usually confined to apical margin; procoxae niches. Larvae are reared in individual cra- usually distinctly separated (contiguous in dles somewhat similar to those of other am- three African genera); subplumose setae al- brosia beetles. most always present. Taxonomy. —The unique genus Scolyto- Biology. —All American genera are bigy- platypus appears to be a highly specialized nous, except that is monoga- geographical replacement of the neotropical mous; habits of African genera are largely un-

Scolytodes . It appears to be the only Old reported. Most species occur in broadleaf World representative derived from the same trees and shrubs in desert or semidesert ar- ancestral stock as the Ctenophorini. A careful eas. Stenoclyptus , Phloeocleptus , and most study of Scolytoplatypus shows quite clearly are phloeophagous; Thysa- how the transition from unsocketed tibial noes, Micracis, and are xy- spines to socketed tibial teeth (derived from lophagous; the species of Micracisella are setae) took place (Wood 1978). This transition myelophagous in small twigs. Details of their occurred well after Platypodidae had di- life cycle are poorly known, except that the verged from the main ancestral line and after development of most species is very slow. family characters for Scolytidae had been Taxonomy. —The tribe occurs in North and fixed. South America and Africa, with one species in 62 Great Basin Naturalist Memoirs No. 10

Table 2. A comparison of American and African gen- Key to the Genera of Micracini era of Micracini arranged in order of structural complex- ity, with the first member in each cluster forming geo- 1. Elytra broadly rounded behind (except sub- graphical replacements of one another on the same acinninate in Pseudothysanoes mucronatus); horizontal line. Other genera in each cluster from the protibia usually much more slender, less same continent are thought to have been derived from the strongly flattened (except ); lateral first member of that cluster. margins of antenna! club constricted at sutures 1 and (usually) 2, except when sutures absent American genera African genera (American species) or sutures procurved (African species) 2 Pseudothysanoes Lanurgus Stenocbjptus Traglostits — Elytral apices acuminate (partly lost in some Thysanoes Saurotocis MicraciseUa), usually mucronate; protibia more strongly flattened, at least apically (ex- Miocryphalus ceptions in African species); antennal club Phlococlcptiis Phloeocurus without constrictions at sutures, sutures always Hylocurus indicated on anterior face in American species, absent in some African species 7 Micracis Pseudomicracis

MicraciseUa 2(1). Pronotum longer than wide, its summit less strongly developed; protibia rather broadly flattened, sides parallel, subtruncate apically; xylophagous; USA to Costa Rica; 1.2-2.3 mm N Asia (China). Almost every American genus Thysanoes has a closely allied counterpart in Africa; obvi- — Pronotum as wide or wider than long, summit ously the generic traits were rather well estab- well developed; protibia slender, weakly or not lished before the faunas were separated prior at all flattened; mostly phloeophagous (some to early Tertiary, but some )nodification con- exceptions) 3 tinued after isolation. The Asiatic Pseu- 3(2). Procoxae contiguous; sutures of antennal club, dothysanoes mongolica (Sokanovskii) was ob- when present, strongly procurved, suture 1 attaining middle of club; antennal scape short, viously derived from American stock much less than twice as long as club, in female usually recently. Because most of the species in more as broad as long and ornamented by numerous this group breed in small twigs and branches setae; Africa 4 of trees and shrubs having little or no eco- — Procoxae moderately separated (a few subcon- nomic importance, they are regarded by tiguous); sutures of antennal club, when forest biologists as scientific curiosities and, present, straight to moderately procurved, consequently, are poorly known. never attaining middle of club; scape elongate except subgenus Aphanoclepfus (Pseudothysa- As seen in Table 2, three clusters of genera noes), its vestiture usually rather sparse; in this tribe are represented in both America America 6 and Africa. The first species in each cluster is a 4(3). Antennal club with sutures present, marked by geographical replacement of its counterpart setae; female mandibular and declivital spines on the other continent, and it is considered to absent; female frons concave to convex; 1.1- have given rise to the other genera within its phloeophagous; Madagascar and Africa; 2.7 mm Lanurgus own cluster on its own continent. Miocry-

phalus is unrepresented in America and is — Antennal club finely, uniformly, rather densely pubescent, sutures obsolete; female thought to have been derived very early from frons concave 5 a specialized representative of the ancestral 5(4). Antennal funicle 6-segmented; mandible bear- stock. The allies of this genus or its ancestor ing one or more very long, slender spines di- possibly given rise to ancestral could have rected dorsad; female declivity armed on inter-

Xyloctonini and Cryphalini. striae 1 at base of declivity by a prominent Whereas primitive Micracini appear to be spine; Africa; 2.2 mm Traglostus

allied to the ancestral stock that produced — Antennal funicle 5-segmented; mandible nor-

Carphodicticini, I see no clue as to where mal; declivity unarmed; Africa; 1.3-2.5 mm . . Miocryphalus these groups connect to more primitive

Scolytinae, except that it apparently was near 6(3). Elytral declivity in both sexes variously sculp- but never sulcate; antennal scape long or the Ctenophorini. The Micracini appear to be tured short, club with or without sutures; mostly the only possible ancestral group for the Cac- phloeophagous, a few xylophagous; North and is topinini; however, the connection remote South America, 1 in China; 0.7-2.0 mm and obscure at best. Pseudothysanoes 1986 WOOD: Genera of Scolytidae 63

— Elytral declivity bisulcate, subvertical; anten- 1 of 5 apical teeth on outer (lateral) margin; nal scape short, flattened, little if any longer scape usually less strongly expanded; antennal than pedicel, club small, widest through basal club rather broad, sutures more broadly pro- half, sutures 1 and 2 straight, indicated by rows curved; monogamous, myelophagous; North

of setae; phloeophagous; USA (California) and America to Colombia; 1.2-2.5 mm . . Micracisella Mexico (Oaxaca); 1.0-1.5 mm Stenoclyptus — Eye entire, always widely separated below; 7(1). Eye short, oval, not more than 1.5 times as long protibia more strongly flattened, all 5 teeth on

as wide (except up to 3.0 times in Phlococunis , apical margin; scape usually very strongly ex- Africa), finely faceted; antennal club small, su- panded; antennal club more elongate, sutures tures straight to weakly procurved or bisinuate usually much more strongly, narrowly arcuate; (South American Hylocunis with strongly pro- bigynous, xylophagous; North and South curved sutures and with posterior face of pro- America; 1.6-3.4 mm Micracis tibia tuberculate); protibia rather slender, wider apically 8 Lanurgus Eggers [1920a: 36, Type-species: — Eye elongate, 2.0 or more times as long as Lanurgus barhatus Eggers, monobasic. Syn- wide, coarsely faceted; antennal club rather onyms: Landolphianus Schedl 1950c; 106, sutures very strongly procurved; pro- large, Type-species: Landolphianus elongatits tibia rather strongly flattened (posterior face Schedl, subsequent designation by Schedl never tuberculate); American species 12 1962a: 38; Micraciops Schedl 1953a: 86, Type- 8(7). Antennal club unmarked by sutures, uni- species: Micraciops formly, finely pubescent, elongate, more than catenattis Schedl, subse- twice as long as wide; female declivity simple, quent designation by Schedl 1962: 38; Pseii- male declivity impressed medially, elevated dohylocurus Nunberg 1961: 613, Type- laterally and armed by large spines; Madagas- species: Pseudohylocurus caplandicus Nun- car; 1.6-2.8 mm Saurotocis berg = Laniirgus podocarpi Schedl, original — Antennal club stouter, sutures marked by rows designation]. Distribution: 24 species in of setae 9 Africa and Madagascar. 9(8). Antennal club with sutures straight, or if bisin- uate or procurved, then posterior face of pro- Traglostus Schedl [1938d: 454, Type-spe- cies: Traglostiis cxornatiis tibia armed by tubercles; American species . . 10 Schedl, monoba- sic]. — Antennal club with sutures moderately to pro- This genus is doubtfully distinct from foundly procurved; protibia slender, never Lanurgus. Distribution: 4 species from armed by tubercles on posterior face; African Kenya to South Africa. species 11 Miocryphalus Schedl [1939d: 381, Type- 10(9). Sutures 1 and 2 on antennal club straight, visi- species: Stephanoderes natalensis Eggers, ble only at margins, obsolete in central area, monobasic. Synonym: Afroniicracis Schedl club small (Fig. 47); protibia entirely unarmed on poosterior face, smooth (Fig. 47); phloeo- 1959d: 709, Type-species: Afromicracis ken- phagous; Mexico; 1.0-1.8 mm Phloeocleptus yaensis Schedl, monobasic]. Distribution: 6

— Sutures 1 and 2 on antennal club bisinuate to species in Africa. weakly or strongly procurved, clearly visible in Pseudothtjsanoes Blacknian [1920: 46, central area of anterior face, club larger; pro- Type-species: Pseudothysanoes drakei Black- tibia armed on posterior face by many tuber- cles or rugae; xylophagous; North and South man = rigidus LeConte, original America; 1.3-3.2 mm Hylqcurus designation. Synonyms: Cryptocleptes Black-

11(9). Eye larger, more elongate, 2.5-3.0 times as man 1920: 51, Type-species: Cryptocleptes long as viide; sutures of antennal club moder- dislocatus Blacknian, original designation, ately procurved; protibia slender, with denti- preoccupied; Clialcohyus Blackman 1943b: cles on apical margin; female frons convex; de- 363, Type-species: Clialcohyus securigerus clivity armed by moderately coarse tubercles in both sexes; phloeophagous; Africa; 2.5-3.0 Blackman, original designation; Bostrichips mm Phloeocurus Schedl 1951a: 21, Type-species: Bostrichips

— Eye usually smaller, oval, about twice as long spinatus Schedl, monobasic; Gretschkinia as wide; sutures of antennal club moderately to Sokanovskii 1959: 276, Type-species: Gre- profoundly procurved, protibia slender, with 1 tschkinia mongolica Sokanovskii, mono-basic; or more denticles on lateral margin; female Aphanocleptus Wood 1960b; 63, Type-spe- frons convex to concave; declivity unarmed; cies: Aphanocleptus coniferae Wood, original Madagascar, Africa; 1. 1-1.8 mm Pseudomicracis designation; C ryptulocleptus Wood 1962: 76,

12(7), Eye shallowly emarginate, often approximate Type-species: C ryptocleptes dislocatus Black- below; protibia less strongly flattened, at least man, automatic; Neoglostatus Schedl 1978: 64 Great Basin Naturalist Memoirs No. 10

1. H. equidens 2. H. medius 3. H. maequalis

=^^A -N^^ [ , ^ \, ^ —1 /

1

10. Ph. caudatus V.5«

Fig. 47. Micracini spp., elytral declivities, antennae, and protiliia: 1, Hylocurtin equidens Wood, male; 2, Hylocurus medius Wood, male; .3, Hijlocurus inaequalis Wood, male; 4, Hijlocurus effemincitus Wood, male; 5, Micracis Ugnator Blaekman (= truncatus Wood), male; 6, Pseudothysanoe.i spinatus Wood, male; 7-10, Phloeocleptus caudatus Wood, 7, 10, male, S, 9, female. 1986 WOOD: Gener.\ of Scoutidae 65

300, Type-species; Neoglostatus sqiiamosus Schedl, monobasic]. Distribution: 61 species in North and Central America, 8 in the An- tilles Islands and South America, 1 in China. All are big>'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). Fig. 48. Cactopimis desertus Bright, male. (.After Bright 1973; 155). Hylocurus Eich'hoff [1872: 133, T>pe- species: Hylocurus elegans EichhofP, mono- basic. Synonym: Micracisoides Blackman preoccupied]. Distribution: 20 species in 1920: 19, T}"pe-species: Micracis rudis North and Central America 1 of which also LeConte, subsequent designation by Wood occurs in South America (Colombia). All are 1982: 608]. Distribution: 34 species in North monogamous and mvelophagous. Kev: Wood and Central America, about 20 in South (19S2:^594). America and adjacent islands. All are bigy- Micracis LeConte [1868: 164, Type-spe- nousandxvlophagous. Kevs: Blandford (1898: cies: Micracis siituralis LeConte, subsequent 220), Wood (1982: 609). designation by Hopkins 1914: 125]. Distribu- Phloeocurus Wood [1984: 230, Type-spe- tion: 18 species in North and Central Amer- cies: Hylocunts africanus Schedl, original ica, 1 in Cuba, 2 in South America designation]. Distribution: 1 species in Africa. (Venezuela); several others have been named Pseudomicracis Eggers [1920a: 36, Type- from South America but most, if not all of species: Faeudomicracis elsae Eggers, mono- these, have been or should be transferred to xy- basic]. Although the unique holotype of the other genera. All are bigynous and lophagous. Key: (1982; 579). type-species is lost, the genus is identifiable Wood from the description. Distribution: 7 species Tribe Cactopinini in Africa and Madagascar. Cactopinae Chamberlin [19.39: 243, T\'pe-genus: Cac- Micracisella Blackman [1928b: 192, Type- topinus Schwarz, 1899] species: Micracis opacicollis LeConte, auto- matic. Synonym: Pseudomicracis Blackman Description.—Frons dimorphic, male 1920 (December): 20, Type-species: Micracis strongly impressed or excavated, with the opacicollis LeConte, original designation. epistomal margin armed by a pair of (usually 66 Great Basin Naturalist Memoirs No. 10 confluent) hornlike spines of large to enor- male slightly to moderately flattened and mous size (Fig. 48), female convex or mod- sometimes abundantly pubescent; eye short, estly impressed, epistoma unarmed by broadly oval and entire to very elongate and spines; posterodorsal area of head modestly sinuate to shallowly emarginate; scape short produced caudad; eye small, oval, entire; an- and rather stout to elongate and rather slen- tennal scape rather short, funicle 5-seg- der, funicle 5-segmented, club flattened, mented, club almost conical to strongly flat- small, and symmetrical, with transverse su- tened, sutures straight to procurved, marked tures, to moderately large and slightly asym- by rows of setae; pronotum asperate on ante- metrical, with slightly oblique sutures, su- rior slope, summit at or near posterior mar- tures marked by grooves and rows of setae, gin, sometimes projecting caudad beyond apparently not septate; pronotum elongate, basal margin; procoxae contiguous; elytral sides conspicuously constricted on middle sculpture unique, rather conservative, usu- half, pronotum unarmed; posterior face of ally sulcate on declivity, almost always coated head truncate, dorsomedian area not ex- by resinous film of host origin. tended caudad; proco.xae narrowly to rather Biology. —All are monogamous and either widely separated; protibia slender to very phloeophagous or in cactus (Cereits and allied stout and short, armed by socketed denticles genera). Those in cactus breed in dry, dead on lateral margin; scutellum visible; basal (yellowing) tissue immediately below the epi- margins of elytra rounded in two genera, ele- dermis, or in scar tissue in deep wounds (hiib- vated and carinate in one genus. bardi). One species breeds in Yucca leaves Biology. —Carphodicticus is monogamous (depressus). The parental galleries form an and phloeophagous. Numerous pairs of par- irregular cave, with egg niches. Eggs are de- ent adults appeared to use the same entrance posited individually in niches in two species; tunnel. Each pair followed a previously made some species in cactus deposit them in clus- tunnel for a short distance, then formed their ters. Larval mines may be individual or lost in own branch gallery for oviposition, such that a criss-crossing maze. Successive generations the entire system consisted of branching and have been bred in the same piece of dry cactus rebranching galleries. Eggs were deposited for four years. Symbiotic relationships with individually in niches at the cambium. Larval fungi have not been reported. mines were exposed on peeled bark and were Taxonomy. —This unique tribe is restricted rather short. The host had been felled for to the Mexican plateau region. Its nearest several months before this species attacked; a affinity to other groups appears to be with the Phloeotribus had largely abandoned the un-

Micracini, although the relationship is re- usually hot, dry tissues, but this species was mote. They are exceedingly rare. thriving. Cactopinus Schwarz [1899; 11, Type-spe- Taxonomy. —The head and pronotal struc- cies; Cactopinus hubbardi Schwarz, monoba- ture suggest that this group is very primitive. sic. Synonym: Cactopinonis Bright 1967; 918, It was probably derived from the same ances- Type-species: Cactopinus cactoplithorus tral stock as the Dryocoetini, Ipini, Cryp- Wood, original designation]. Distribution: 19 turgini, Xyloterini and Xyleborini, but at a species in W North America (California and W much earlier date. The disjunct distribution Utah to Me.xico). All are monogamous and and rarity suggest that it is a relict group that phloeophagous when in woody plants or had reached its maximum potential prior to subepidermal when in cactus {Cereus and al- the Tertiary. This tribe represents a first step lied genera only). Keys; Blackman (1938a: toward one of the three most hightly evolved 151), Bright (1967: 919), Wood (1969; 43, tribes (Xyleborini) of Scolytidae. 1982; 638). Key to the Genera of Carphodicticini Tribe Carphodicticini 1. Eye entire, broadly oval, short, about 1.3 times Carphodicticini Wood [197L 19, Type-genus: Carphod- as long as wide; scape rather short, about three icficHS Wood, 1971] times as long as pedicel; antennal club rather small, sutures transverse; elytral apex strongly Description. —Frons weakly to moder- mucronate; ventrolateral margin of declivity ately dimorphic, male strongly convex, fe- armed by a row of rather strongly elevated serra- -

1986 WOOD: Genera of Scolytidae 67

armed by a row of rather strongly elevated serra- lum visible; elytral declivity moderately sul- tions; apparently xylophagous; India and Sri cate to strongly excavated, lateral margins Lanka (Ceylon); 1.3-2.3 mm Craniodicticus usually armed by tubercles or spines; vesti- — Eye sinuate to emarginate, elongate, more than ture hairlike. twice as long as wide (not visible on type of Dendrodicticus), elytral apex rounded, not mu- Biology. —All are phloeophagous. Most cronate; lateral margin of declivity uniformly are heterosanguineously polygynous, al- elevated, not armed by serrations 2 though some Acanthotoniicus are monoga- 2(1). Basal margins of elytra rounded, not elevated; mous. Hosts include members of the antennal club rather large, distinctly asymmet- Pinaceae, except Acanthotoniicus breeds pri- rical, sutures slightly oblique; procoxae more narrowly separated; protibia short (only slightly marily in various angiosperm trees. Eggs are longer than antennal club), stout (almost half as deposited in niches. Larval mines are individ- wide as long); phloeophagous; South America ual and rarely cross one another; they are (Venezuela); 2.0-2,4 mm Carphodicticus exposed on the surface of peeled bark. The life Basal margins of elytra rather strongly, acutely — cycle is comparatively short, with two or more elevated along a continuous costa; antennal club generations per year apparently the normal rather small, symmetrical, sutures transverse; procoxae more widely separated; protibia habit. longer, much more slender; habits unknown; Ta.\ONOMY. —In most tribes of Scolytidae South America (Argentina); 2.0 mm the sexes are easily distinguished by a differ- Dendrodicticus ence in the number of abdominal terga. The Craniodicticus Blandford [1895: 317, female tergum 8 is reduced in size and is Type-species: Craniodicticus miicronatus telescoped beneath tergum 7 such that it is Blandford, monobasic]. Distribution: 1 spe- ordinarily hidden from view. A visible tergum cies in S India, 1 in Sri Lanka (Ceylon). They 8 is present in the male. However, in Carpho- apparently are xylophagous, since one sample dicticini, Ipini, Dryocoetini, Crypturgini, Xy- was "removed from wood" and the other from loterini, and Xyleborini a small tergum 8 is ' a creeper called "jungle rope. visible in both sexes. The general body Carphodicticus Wood [1971: 19, Type-spe- habitus, shape of the eye, basic type of anten- cies: Carphodicticus cristatus Wood, original nal club (trend toward being obliquely trun- designation]. Distribution: 1 species in South cate), and tibiae are also shared by these

America (Venezuela). It is monogamous and tribes. The Ipini occupy a position between phloeophagous; it utilized the entrance tun- Carphodicticini and Dryocoetini in which nels of a Phloeotribus species in order to gain these characters were being formed. In Den- access to the phloem. drochihis they are so poorly expressed that Dendrodicticus Schedl [1958a: 37, Type- they are detected only with difficulty. How- species: Dendrodicticus argentiniae Schedl, ever, that genus does exhibit a stage in transi-

monobasic]. Distribution: 1 species in South tion to the circumtropical Acanthotomicus . America (Argentina). Acanthotoniicus appears to have an ancient origin (prior to the Tertiary) and to have given Tribe Ipini rise, directly or indirectly, to Pitijogenes,

Pityokteines, Orthotomicus , and Ips. Pitijo- Ipini Bede! [1888: 386, Type-genus: Ips DeGeer, 177.5] genes appears to have arisen in Europe and Description. —Frons usually dimorphic, northern Asia from African Acanthotoniicus male convex, female variously excavated, pro- like stock and to have reached North America tuberant, or ornamented by setae; eye sinu- rather recently. Something resembling the ate, lower half usually much narrower than neotropical Acanthotomicus miniicus species above; antennal scape slender, elongate, funi- group probably gave rise in North America in cle 5-segmented, club either obliquely trun- early Tertiary to a stock that then evolved into cate or sutures on posterior face strongly dis- Pityokteines and Ips in North America and to placed toward apex; pronotum rather strongly Orthotomicus in Europe and Asia. At a later declivous on anterior half, rather closely, date, a few representatives oi Pityokteines and coarsely asperate; procoxae contiguous, inter- Ips then migrated into Europe and Asia and coxal piece deeply notched or absent; protibia one species of Orthotomicus reached North armed by three or four socketed teeth; scutel- America. The reasoning on which the above is 68 Great Basin Naturalist Memoirs No. 10

Antennal club flattened, based is (1) on the continent of origin of sev- — sutures either pro- curved or moderately to strongly bisinuate; lat- eral species groups found today, only a frac- eral margins ol elytral declivity armed by 1 to 6 tional number of those species groups now pairs of major denticles, third pair (if present) on occur in the invaded territory and (2) no spe- or incorporated into crest of lateral margin; eye cies group is found in the invaded territory usually either abnormally large or else very small 5 that is not presently also found in the area of origin. Fossils that might document this sup- 5(4). Sutures of antennal club (when present) moder- ately to very strongly procurved; eye large, very position are unknown. Dendrochihis and coarsely faceted, its width about equal to length Acanthotoinicus breed in a variety of non- of scape, its length more than twice length of coniferous hosts; , Pitijokteines, scape; Mexico to South America, S Asia to Aus- tralia, Orthotomiciis , and Ips occur exclusively in Africa; angiosperm hosts (some Asian ex- Pinaceae. ceptions); 1.4-2.7 mm Acanthotomicus — Sutures of antennal club moderately to strongly Key to the Genera of Ipini bisinuate (procurved in concinmts, mexicanus,

orientatis): eye small, finely faceted, its width 1. Eye short, oval, entire, not sinuate on anterior equal to much less than length of scape, its margin; antennal club rather small, flattened, length equal to much less than twice length of unmarked by sutures; declivity convex to scape; North America to N Nicaragua, Asia, Eu- slightly, broadly flattened, without granules or rope, N Africa; Pinaceae; 2. 1-6.9 mm Ips tubercles; elytral vestiture not abundant, in rows, interstrial setae hairlike to scalelike; pro- Dendrochilus Schedl [1957a; 70, Type-spe- tibia slender, armed by 3 socketed denticles on cies: Dendrochilus strombosiopsis Schedl, apical half of lateral marin; Africa; 1.0-2,0 mm Dendrochilus subsequent designation by Schedl 1962a: 55].

— Eye more elongate, sinuate on anterior margin; Distribution: About 10 species from Africa. antennal club usually larger, marked by sutures Pityogenes Bedel [1888: 397, Type-species; (except absent in many Acanthotomicus), de- Dermestes chalcographus Linnaeus, original clivity usually excavated and variously armed by designation. Synonyms: Eggersia Lebedev tubercles or spines 2 1926: 121, Type-species: Bostrichus biden- 2(1). Elytral declivity rather narrowly bisulcate, lat- tatiis Herbst, present designation; Pityocera- eral margins rather broadly elevated, rounded, genes Balachowsky 1947: 44, Type-species: and armed by not more than .3 pairs of denticles; lower margin of declivity rounded; usually Bostrichus quadridens Hartig, original desig- smaller than 3 mm 3 nation]. Distribution: 7 species in North — Elytral declivity broadly, rather deeply exca- America, 18 in Europe, Asia, and N Africa. All vated, margins acutely elevated and armed by 3 are heterosanguineously polygynous and or more pairs of denticles (1 to 6 pairs in tropical phloeophagous. Keys: Swaine (1918; 104), Acanthotomicus), lower margin of declivity with an acutely elevated, transverse ridge separating Bright (1976: 150), and Wood (1982: 650) for declivital excavation from apical margin (Fig. North America, Reitter (1913: 97) and Pfeffer 49); usually larger than 3 mm 4 1946: 112) for Europe, Stark (1952: 272) for 3(2). Prosternal intercoxal piece short, obtuse; female USSR, Schedl (1962c: 134) for Europe and frons sometimes deeply, rather narrowly exca- Asia. vated; male declivity with 2 or 3 pairs of en- Pityokteines Fuchs [1911: 33, Type-spe- larged denticles; antennal club compressed, 2 sutures visible on apical third of posterior face; cies: Ips curvidens Germar, subsequent des- North America, Europe, Asia, N Africa; Pina- ignation by Hopkins 1914: 127. Synonymy: ceae; 1.8-3.7 mm Pityogenes Ortliotomides Wood 1951: 32, Type-species: — Prosternal intercoxal piece long and acutely ta- Orthotomicus lasiocarpi Swaine, original des- pered; female frons convex, never excavated; ignation]. Distribution: 6 species in North male declivity more narrowly impressed; female America, 3 in Asia and Europe. All are het- frons and anterior pronotum with dense, long phloeopha- vestiture (2 American exceptions); North Amer- erosanguineouslv polvgvnous and ica, Asia, Europe; Abies; 1.6-3.0 mm gus. Keys: Swaine (1918: 123), Bright (1976: Pityokteines 145), and Wood (1982: 656) for North Amer- 4(2). Antennal club obliquely truncate, sutures re- ica, Reitter (1913: 102) for Europe, Bala- curved; third (lowest) major denticle not on lat- chowskv (1949: 255) for France, Stark (1952: eral margin of elytral declivity, displaced mesad 421) for the USSR. from margin, eye ofnormal size; North America, Ferrari Type-spe- Asia, Europe; Pinaceae; 2.2-4.3 mm Orthotomicus [1867: 44, Orthotomicus 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 Taplirorychini Reitter [1913: 29, Type-genus; Taphro- rychus Eichhoff, 1878] spelling Onthotomicus Ferrari (1867: 44), a lapsus calami that was corrected by Ferrari Description. —Frons usually sexually di- Neotomiciis (1869: 256). Synonym: Fuchs morphic, male convex to variously impressed, 1911: Type-species: laricis 33, Bostrichus female convex to flattened or with elevations, Fabricius, designation subsequent by Hop- variously ornamented by setae in many spe- kins 1914: 125]. Distribution: 1 species in cies; eye distinctly emarginate to divided; an- 10 in Asia, North America, about Europe, and tennal scape slender, elongate, funicle 4- to Africa. All are heterosanguineously polygy- N 6-segmented, club obliquely truncate to nous and phloeophagous. Kevs: Reitter (1913; strongly flattened, if flattened then sutures for Europe, (1949: for 108) Balachowsky 268) variously procurved to obsolete, and, on pos- France, Stark for the (1952: 407) USSR. terior face, strongly displaced toward apex; Acanthotomicus Blandford [1894a: 89, pronotum arched from base or not, if anteri- Type-species: Acanthotomicus spinosus orly declivous then declivity usually involving Blandford, monobasic. Synonym: Miinips Eg- more than anterior half, armed or not, if gers 1932: 33, Type-species: Ips pilosus Eg- armed then asperities small, usually abun- gers, original designation; Isophthorus dant; procoxae contiguous to narrowly sepa- Schedl 1938c: 173, Type-species: Isophthorus rated; protibia with lateral margin armed by quadrituherculatus Schedl, designated by four to several socketed teeth {Chiloxylon and 1980: 89]. Distribution: 10 species in Wood a few Coccotrypes have three); scutellum visi- Central and South America, more than 20 in ble; elytral declivity usually convex, some- Africa 14 in to and about China and Japan times shallowly sulcate or variously flattened, Australia and the Philippines. All are sometimes armed by small granules; vestiture phloeophagous; of 13 species studied in na- hairlike. ture by me, 1 was monogamous and 12 het- Biology. —Apparently all are polygynous, erosanguineously polygynous. Key: Wood with heterosanguineous polygyny occurring (1982: 664) for Central America. in all except the consanguineous Dryo- Ips DeGeer [1775: 190, Type-species: coetiops, Coccotrypes (Fig. 50), and Ozope- Tomicus tijpographus =Deniwstes tijpogra- mon. Most are phloeophagous, Dactylotrypes phus Linnaeus, subsequent designation by and some Coccotrypes are spermophagous, Bergroth 1884: 230. Synonyms: Cumatotomi- and at least one species {Dryocoetiops cof- cus Ferrari 1867: 44, Type-species: Bostri- fcae) is myelophagous. Most eggs are placed chus stenographus Duftschmidt = Dennestes in individual niches. Most larvae form inde- sexdentatus Boerner, subsequent designation pendent mines, but some feed in congress. by Hopkins 1914: 119; Cijrtotomicus Ferrari Taxonomy. —This is a large, rapidly evolv- 1867: 44, Type-species: Bostrichus acumina- ing group; consequently, generic boundaries tus Gyllenhal, subsequent designation by are not always clear. Three major groups of Hopkins 1914: 120]. Distribution: 25 species genera are apparent; (1) Dryocoetiops, Coc- in North and Central America, about 18 in cotrypes, and Ozopemon, (2) Dactylotrypes, Asia, Europe, and N Africa; at least 1 has been Dendrocranuhis , Xylocleptes, Thamnurgus, introduced to Australia and the Philippine Is- Tiarophorus, Triotemnus . etc., and (3) Cyr- lands. All are heterosanguineously polygy- togenius, , , etc. nous and phloeophagous. Numerous species Although structural diversity may not war- belonging to other genera have erroneously rant it, these three species groups will be been assigned to this genus at one time or discussed separately. In group 1, reproduc- another. Keys: Schedl (1950b: 69) for Europe tion in all species involves arrhenotocous and Asia, Hopping (1963-1965) and Wood parthenogenesis. Males, if known, are (1982: 669) for North and Central America. dwarfed, deformed, flightless, and haploid. Tribe Dryocoetini With about two possible American exceptions

in Coccotrypes , all species originally oc- Dryocoetoideae Lindemann [1876: 165, Type-genus: curred in the Ethiopian, Oriental, or Aus- Dryocoetes Eichhoff, 1864] Thamnurginae Niisslin [1911: 377, Type-genus: Tham- tralian realms. Many spermophagous Coc- nurgus EichhofiF, 1864] 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- ferent that there is no possibility of a close ical distributions. Of the dozen or so Ameri- relationship between these groups. can species, all but two are known to have Key to the Genera of Dryocoetini reached America through commerce, and it is

presumed that in time these two will be found I . Protibia armed on lateral margin by 5 or more to have foreign origins. socketed teeth (rarely 1 or 2 reduced, if so, then sutures on anterior face of antennal club Group 2 is one of the few scolytid groups to procurved, except only 2 teeth present in invade herbaceous plants (mostly Cucur- Dolurgocleptes); posterior face of antennal bitaceae and Euphorbiaceae). A thorough club with only one suture indicated (except 2 in some Taphronjchus), sutures on anterior study oi Thamnurgus , Xijlocleptes , and Taph- face either procurved or club obliquely trun- ronurgus will probably find that intergrada- cate, suture 1 more commonly on apical half; tion between them is complete and, conse- male of normal size, joins female in new quently, that they must be combined into one parental gallery 2 genus. Group 2 undoubtedly is the most an- — Protibia armed on lateral margin by 2 to 4 cient of this tribe, li Dendrocranulus (Ameri- socketed teeth; posterior face of antennal club marked by 2 sutures, suture 1 on anterior face can) was separated from the almost indistin- usually on basal third, both sutures either guishable Xylocleptes (African) early by straight or recurved; lateral margins of prono- Tertiary, then the origin of their common an- tum usually acutely or subacutely elevated at cestor and of the tribe must be pushed back least near base (some exceptions); male dwarfed, of abnormal shape, flightless, never into the Cretaceous. Early members of this joins female in new parental gallery (except group probably gave rise to groups 1 and 3. unknown in Chiloxylon ) 15 Group 3 appears to have originated in the 2(1). Antennal funicle 4- or 6-segmented; prono-

Old World, with Dryocoetes and Lymantor tum unarmed, feebly if at all declivous on less reaching northern North America rather re- than anterior fourth; eye deeply emarginate cently. Schedl repeatedly called attention to to divided; male frons moderately to strongly impressed 3 the similarity between some of the small, — Antennal funicle 2- to 5-segmented; anterior slender Cyrtogenius and ; how- half of pronotum declivous and armed by as- ever, the antennae, elytral locking mecha- perities (except feebly declivous and unarmed nism, and other characters are so totally dif- in Tlunnnnrgtis) 5 72 Great Basin Naturalist Memoirs No. 10

3(2). Antennal funicle 6-segmeiited; pronotum — Antennal club distinctly longer than scape, conspicuously longer than wide, teebly de- more broadK' rounded, sutures rather weakly clivous on less than anterior half; sutures on to very strongly procurved; male frons indis- antennal club procurved; procoxae narrowly tinctly impressed; S Europe, Africa; Cucur- separated; Africa; Euphorbiaceae; 1.4-.5.5 bitaceae, etc.; 2.0-2.7 mm Xylocleptes mm Tiarophorus 9(6). Sutures of antennal club obscure, strongly — Antennal funicle 4-segmented, sutures of procurved, basal area of club not corneous; club straight or slightly recurved (when body more slender, pronotum with asperities present); pronotum feebly to moderately de- usually restricted to anterior half; America; clivous on more than anterior third; procoxae Cucurbitaceae; 1.2-2,7 mm .. Dendrocranulus contiguous 4 — Sutures of antennal club on apical fourth, re- 4(3). Eye divided; protibia rather broad, margin curved, basal three-fourths of club thickened, with only 2 socketed teeth; antennal club corneous; body rather stout; asperities extend membranous and pubescent, with 1 suture to base of pronotum; Canary Islands; Dra- indicated at middle on front and back; prono- caena draco; 1.8-2.0 mm Dactijlotrypes tum smooth, shining throughout, very feebly 10(5). Antennal funicle 4-segmented, sutures of declivous on anterior half; striae not im- club very strongly procurved; male frons shal- pressed, punctures small, in rows, scarcely lowly to moderately impressed, female frons larger than those of interstriae; Madagascar; less distinctly impressed; slender species with 1.9 mm Dolurgocleptes punctures on elytral disc confused; partly xy- — Eye sinuate on anterior margin; lateral mar- lophagous; North America, N Asia, Europe; gin of protibia armed by 4 or 5 socketed teeth; 1.6-2.0 mm . Lymantor strial punctures usually larger, deeper, dis- — Antennal funicle 5-segmented (rarely 2-, 3-, tinctly larger than those of interstriae; Africa or 4-segmented in New Guinea and Indonesia to India; Euphorbia; 1.2 mm Triotemnus species), sutures on club procurved or not; 5(2). Protibia more broadly flattened on at least male frons usually convex, female frons fre- apical half, lateral margin armed by 7 or more quently pubescent 11 socketed teeth; male Irons variously im- 11(10). Pronotum with summit distinctly elevated pressed, female flat to convex and orna- near middle, often a moderate, transverse im- mented or not; sutures on antennal club mod- pression behind summit; antennal club estly to profoimdly procurved, except strongly flattened, basal area slightly cor- recurved in some Thamnurgus , scutellum neous, suture 1 distinct, straight to strongly very small, not Hat; in stems of Cucurbitaceae bisinuate, its lateral extremities reaching or Euphorbiaceae 6 basal fourth, median portion never exceeding — Protibia less strongly flattened on apical third, middle of club, 2 sutures on posterior face; lateral margin armed by 5 socketed teeth meso- and metatibiae more slender; hair on (rarely 3 to 6); male frons convex (rarely feebly pronotum and elytra fine, unusually long; Eu- impressed), female variously modified and rope and Asia; broadleaf hosts; 1.8-3.2 mm frequently ornamented by hair; sutures on Taphrorychus antennal club recurved, straight, procurved, — Pronotum with dorsal profile evenly arched, or obsolete; scutellum rather large, flat; in summit inconspicuous or else on basal fourth; phloem or xylem of trees and shrubs 10 basal area of antennal club less strongly flat- Groove face of metatibia for re- 6(5). on posterior tened, more strongly corneous or if not cor- ception of tarsus poorly developed, short, oc- neous then sutures obsolete and pubescence cupying less than apical third; Europe and extending to base; meso- and metathoracic Asia to Africa 7 tibiae usually more broadly flattened; prono-

— Metatibial groove conspicuous, occupying tal and elytral setae of more normal length . . 12 posterior more than apical threcrfourths on 12(11). Antennal funicle 2-segmented, procurved face; America and Canary Islands 9 corneous area of club occupying more than

7(6). Pronotum less distinctly declivous on anterior three-fourths of basal area; procoxae contigu- third, punctured to anterior margin, margins ous; New Guinea to New Britain Island; 1.2- of some punctures feebly to finely asperate; 1.4 mm Dendrographus antennal club rather small, sutures straight to — .Antennal club 3-, 4-, or 5-segmented; usually recurved; frons feebly impressed; male Eu- much larger 13 rope and SW Asia to N Africa; mostly in Eu- 1.3(12). Body very stout, 2.0-2.1 times as long as wide; phorbiaceae; 2.0-3.0 mm Thamnurgus pronotum rather coarsely asperate to base, — Anterior third of pronotum finely asperate, summit on basal fourth; antennal club either punctures obsolete in asperate area 8 devoid of sutures or with 1 on basal fourth,

8(7). Antennal club shorter than scape, 1 . 3 times as almost straight except recurved at margins; long as wide, sutures weakly procurved; male procoxae narrowly separated; declivity frons rather strongly impressed; SE Europe; strongly arched, convex, apical fourth exceed- Clematis; 1.8-2.0 mm Taphronurgus ing vertical and slightly undercutting central 1986 WOOD; Genera of Scolytidae 73

area; New Guinea to Borneo; 1.6-1.8 mm . . . Malaya to Fiji; broadleaf hosts; 3.3-5.5 mm Peridryocoetes Ozopemon

— Body more slender; pronotum u.sually with — Lateral margins of pronotum subacutely ele- punctures on at least part of basal half, summit vated on more than basal half; frons com- indefinite, not on basal fourth 14 monly convergently aciculate; anterior half of pronotum declivous or not; elytral declivity 14(13). Antennal club with suture 1 weakly to pro- usually convex, rarely impressed, granules foundly procun'ed, rarely with sutures absent absent or inconspicuous; mesocoxae rather and pubescent to base; funicle of small species vndely separated by distance two or more with as few as 3 segments (most 4- or 5-seg- times greater than width of pedicel; punc- mented); proco.xae narrowly separated, rarely tures on elytral disc almost always in rows intercoxal piece longitudinally emarginate; (some exceptions); or sper- commonly with ventrolateral margin of de- mophagous; introduced almost worldwide; clivity slightly elevated or armed; Micronesia ' apparently no endemic American species; to Africa; mostly non-coniferous hosts; 1.3-3.0 broadleaf hosts and palm fruits; 1.3-3.7 mm Cyrtogenius mm Coccotryupes — Antennal club with suture 1 recurved, always present, never pubescent to base; procoxae Tiarophorus Schreiner [1882:246, Type- contiguous, intercoxal piece always longitudi- nally emarginate or absent; ventrolateral mar- species: Tiaropltorus clongatus Schreiner, gin of declivity never acutely elevated or spe- monobasic. Synonyms; Hypaspistes Hage- cially armed; North America, Asia, Europe, N dorn 1908: 374, Type-species: Hypaspistes Africa; mostly coniferous hosts; 1..5-5.1 mm camerunus Hagedorn, monobasic, preoccu- Dryocoetes pied; Oi-thaspistes Hagedorn 1909b: 733, 15(1). Protibia rather broad apically, lateral apical Type-species; Hypaspistes camerunus Hage- angle abrupt (almost 90 degrees), 1 denticle on this angle, another on apical margin, and a dorn, automatic; Pseudothamnurgus Eggers third on lateral margin one-fourth tibia length 1912a: 115, Type-species: Thamnurgus scru- from apical angle; funicle 4-segmented, club tator Pandelle, subsequent designation by constricted at partly septate suture 1 (not ac- Schedl 1961: 738], Distribution: 13 species in tually obhquely truncate), suture 2 indicated Africa S Europe. by setae; habitus resembling Dryocoetes, and uniseriate interstrial setae almost scalelike on Dolurgocleptes Schedl [1965a: 61, Type- declivity, declivity very steep; strial punc- species: Dolurgocleptes malgassicus Schedl,

tures coarse, deep; Brazil; 1.7 mm . Chiloxylon monobasic]. Distribution: 1 species in Mada- — Protibia narrowed near apex, outer apical an- gascar. gle not abrupt; funicle 4-segmented, club Triotemnus WoUaston [1864: 264, Type- never septate, obliquely truncate or nearly so species: Triotemnus subretusus Wollaston, 16 monobasic. Synonym: Cladoctoproctiis 16(15). Procoxae narrowly to moderately separated; Schedl 1975: 454, Type-species: Cladocto- anterior margin of pronotum usually armed by serrations (absent in some laevis); prono- proctus scrafa Schedl, original designation]. tum usually more coarsely asperate in ante- Distribution: 8 species in N Africa and the rior areas, its summit more definite and near Canary Islands, 2 in India. Most species breed middle; India and Sri Lanka (Ceylon) to in the stems of Euphorbia, Delphinium, Philippines; broadleaf hosts; 1.7-3.5 mm . . . Dryocoetiops Aconitum, Teucrium, and Bupierum . Key: Peyerimhoff(1949). — Procoxae contiguous, intercoxal piece either longitudinally notched or absent; anterior Thamnurgus Eichhoff [1864: 40, Type- margin of pronotum unarmed, pronotum usu- species: Bostrichus euphorhiae Kiister, sub- ally more finely asperate to unarmed anteri- sequent designation by Hopkins 1914; 130]. orly, its summit less definite and usually well Distribution: About 40 species in Asia, Eu- behind middle 17 rope, and Africa. A thorough review of the 17(16). Lateral margins of pronotum obscurely to species of this and the ne.xt two genera would subacutely elevated only near base; frons probably result in combining into one never convergently aciculate; anterior half of them pronotum always strongly declivous and as- genus. Keys: Reitter (1913: 85) for Europe, perate; elytral declivity either moderately im- Balachowsky (1949: 165) for France. pressed on central half and with interstrial Taphronurgus Reitter [1913: 84, 90, Type- tubercles or discal punctures strongly con- species: Thamnurgus exul Reitter, monoba- fused; mesocoxae subcontiguous, separated by distance equal to width of antennal sic]. Distribution: 1 species in SE Europe to pedicel; phloeophagous; Indonesia and SW Asia. In Clematis. It is probable that this . .

74 Great Basin Naturalist Memoirs No. 10 genus will eventually be combined with Dendrographtis Schedl [1964c; 310, Type- Thamnurg,its species; Pelicerus pygmaeus Eggers, original Xylocleptes Ferrari [1867: 37, Type- designation. Synonym: Protopityophthorus species: Bostrichiis bispinus Duftschmidt, Schedl 1973: 73, Type-species: Protopityoph- monobasic. Synonyms: Xestips Hagedorn thorus durus Schedl =Pelicerus pygmaeus 1912: 353, Type-species: Xestips marginatus Eggers, original designation]. Distribution: 1 Hagedorn, monobasic; Hijlonitis Nunberg species in New Guinea and New Britain Is- 1973: 16, Type-species: Hijlonius hrunneus land. Nunberg, original designation]. Distribution; Peridryocoetes Wood [1984; 230, Type- About 20 species in S Asia, S Europe, and species; Ozodendron nitens Schedl, original Africa. Key: Reitter (1913; 91) for Europe. designation, also cited but not described as a Dendrocranulus Schedl [1937a: 165, Type- new genus by Schedl 1964a; 243]. Distribu- species: Dendrocranulus tardus Schedl, sub- tion: 3 species from Indonesia and New sequent designation by Schedl 1938c: 169]. Guinea. Distribution: 22 species in North and Central Cyrtogenius Strohmeyer [1910c; 127, America, about 15 in South America and adja- Type-species; Crytogenius bicolor Stroh- cent islands. Some species are monogamous, meyer, monobasic. Cyrtogenius Strohmeyer some are heterosanguineously polygynous; all (1911: 16) a valid emendation oi Kyrtogenius bore in dving stems of Cucurbitaceae. Key: Synonyms: Carposinus Hopkins 1915a: 9, 47, Wood (1982: 708) for North and Central Type-species; Carposinus pini Hopkins, orig-

America. inal designation: Orosiotes Niisima 1917: 1, Dactylotrypes Eggers [1927a: 37, Type- Type-species: Orosiotes kumatoeti,sis Ni- species: Dactylotrypes tiyttenboogaarti Eg- isima, monobasic; Metahylastes Eggers 1922: gers =Xyloterus longicoUis Wollaston, mono- 165, Type-species: Metahylastes africanus basic]. Distribution: 1 species in the Canary Eggers, monobasic; Pelicerus Eggers 1923: Islands in fruits of Phloenix canahensis and 216, Type-species: Lepicerus nitidus Hage-

Dracaena drace ; it may have been introduced dorn, original designation; Eulepiops Schedl into France (Balachowsky 1949: 186). The 1939b; 344, Type-species: Eulepiops glaber male apparently joins the female in the Schedl, original designation; Ozodendron parental gallery. Schedl 1957a: 13 and duplicate description by Lymantor Lovendal [1889; 25, 68, Type- Schedl 1964a: 243, Type-species: Pelicerus species: Lymantor sepicola Lovendal =Tomi- grandis Beeson, monobasic; Mimidendrulus cus cort//t Ferris, monobasic]. Distribution; 2 Schedl 1957a; 68, Type-species; Mimiden- species in North America, 2 in Asia and Eu- dridus movoliae Schedl, monobasic, the spe- rope. All are heterosanguineously polygynous cies designated by Schedl (1961: 732) was not and xylophagous in small, dead branches of an original species and is an invalid designa- living trees. Keys: Reitter (1913: 91) for Eu- tion; Carpophloeus Schedl 1959b; 143, Type- rope, Wood (1982: 707) for North America. species; Carpophloeus rugipennis Schedl, Taphrorychus Eichhoff [1878a: 49, 204, monobasic; Taphroborus Nunberg 1961: 617, Type-species: Bostrichiis bicolor Herbst, Type-species; Taphroborus vaticae Nunberg, subsequent designation by Hopkins 1914: original designation; Artepityophthorus 130. Synonyms: Saliciphilus Sokanovskii Schedl 1969: 157, Type-species; Artepityoph- 1954; 17, 20, Type-species: thorus aries Schedl, monobasic; Taphro- machnovskii Sokanovskii, original designa- stenoxis Schedl, nomen nudum, for Taphro- tion; Pseudopoecilips Murayama 1957: 614, stenoxis tenins Schedl, nomen nudum]. Type-species: Pseudopoecilips mikuniyamen- Distribution: About 16 species in Africa, more sis Murayama, original designation; Taphrot- than 40 in the area from India and Japan to erus Schedl 1965b; 341, Type-species: Australia and Micronesia. They are Taphroterus primitus Schedl, monobasic]. phloeophagous and heterosanguineously Distribution: About 12 species in Asia, Eu- polygynous. rope, and N Africa. All are phloeophagous. Dryocoetes Eichhoff [1864; 38, Type- Keys: Reitter (1913: 94), Balachowsky (1949; species; Bostrichus autographus Ratzeburg, 193), PfeflFer (1962: 241). subsequent designation by Hopkins 1914; 1986 WOOD; Genera of Scolytidae 75

121. Synonyms: Anodius Motschulsky 1860: gers = Spermatoplex rhizophorae Hopkins, 155, Type-species: Bostrichus autographiis subsequent designation by Wood 1982: 731; Ratzeburg, subsequent designation by Wood Hyphaene Hagedorn 1913: 254, nomen 1974: 232, suppressed by International Com- nudum. Type-species: Hyphaene guineensis mission on Zoological Nomenclature 1979: Hagedorn, nomen nudum = Bostrichus car- 149; Dryocoetinus Balachowsky 1949: 180, pophagus Hornung, no status]. Distribution: Type- species: Bostrichus villosiis Fabricius, More than 100 species have been assigned to original designation]. Distribution: 7 species this genus, mostly from Africa, S Asia, and in North America, about 80 in Asia, Europe, adjacent areas. Species in most other areas and Africa have been assigned to this genus, have been introduced through commerce. All but more than half of them have been trans- are consanguineously polygynous and phloeo- ferred to other genera. All are heterosan- phagous or spennophagous; a few species may guineously polygynous and phloeophagous. assume both feeding habits. This e.xceedingly

Keys: Reitter (1913: 75) for Europe, Mu- difficult genus is in a state of ta.\onomic chaos. rayama (1957: 594) for Japan and vicinity. Key: Wood (1982: 732) for North America. Bright (1963: 107) and Wood (1982: 724) for North America. Tribe Crypturgini

Chiloxylon Schedl [1959c: 550, Type- Crypturgi LeConte [1876: 374, 387, Type-genus; Cryp- species: Chiloxylon nifulus Schedl, monoba- turgus Erichson, 1836] sic]. Further study of this genus is needed; it Description. —Frons usually not dimor- may be allied to Dendroterus of the Corthylini phic, male sometimes slightly impressed, fe- (Pityophthorina), rather than to Dnjocoetes. male convex; eye deeply emarginate, except Distribution: 1 species in Brazil (Matto sinuate in Deropria ; antennal scape moder- Grosso). ately long, slender, funicle 2- to 3-segmented, Dryocoetiops Schedl [1957a: 13, Type- club comparatively small, two sutures on api- species: laevis Strohmeyer, origi- Ozopemon cal half, 1 sometimes septate, both frequently nal designation]. Distribution: About 15 spe- absent; pronotum usually unarmed; procoxae cies in SE Asia, Sri Lanka (Ceylon), Indone- contiguous; protibia flattened, lateral margin sia, and New Guinea. One species observed armed by several socketed denticles; scutel- in nature appeared to be consanguineously lum visible; elytral punctures in rows or con- polygynous (or some form of parthogenesis) fused, setae hairhke; anterior surfaces and and was myelophagous. (frequently) elytra uniformly reticulate. Ozopemon Hagedorn [1908: 382, Type- Biology. —They are monogamous and species: Ozopemon regius Hagedorn, mono- phloeophagous except that those species asso- basic]. Distribution: About 27 species in SE ciated with Euphorbia bore throughout Asia and Indonesia to Fiji. All are consan- subepidermal tissues of recently killed stems, guineously polygynous and phloeophagous. apparently with little regard to the cambium. Coccotrypes Eichhoff [1878a: .308, Type- usually utilizes the entrance tun- species: Bostrichus dactyliperda Fabricius, nel of another insect. Larval mines are inde- subsequent designation by Hopkins 1914; pendent. Details of the habits have not been 118. Synonyms: Poecilips Schaufuss 1897: studied. 110, Type-species: Poecilips sannio Schau- Taxonomy. —This tribe apparently was fuss, monobasic; Cryphaloides Formenek derived from the same ancestral stock that 1908: 91, Type-species: Cryphaloides don- gave rise to the Dryocoetini, although the isthorpei Formenek = Bostrichus carpopha- true affinities have not been worked out. It gus Homung, monobasic; Thamnurgides appears to have arisen well into the Tertiary in Hopkins 1915a: 45, Type-species: Thamnur- the Ethiopian realm where at least some gides persicae Hopkins =Coccotrypes adoena members of five of the six known genera occur Blandford, original designation; Spernm- today. Crypturgus appears to have reached toplex Hopkins 1915a: 48, Type-species: North America from Asia rather recently. Spermutoplex rhizophorae Hopkins, original Dolurgus (North America) appears to be a

designation; Dendrurgus Eggers 1923: 144, primitive relict of an earlier radiation that is Type-species: Dendrurgus rhizophorae Eg- now extinct except for this species. 76 Great Basin Naturijlist Memoirs No. 10

Key to the Genera of Crypturtiini Coleobothrus Enderlein [1929: 144, Type- species: Coleobothrus jandiacus Enderlein, 1. Antennal funicle 3-segmented, club with su-

tures 1 and 2 recurved, clearly marked by original designation]. Distribution: 2 species grooves and rows of setae, 1 almost at middle of in the Canary Islands and 2 in Africa. Key: pronotum than wide, a club; longer unarmed, Menier (1973: 208). distinct constriction on anterior third; elytra Aphanarthrum WoUaston [1854: 292, rather coarsely striate, strial setae largely obso- lete, interstrial setae short; W North America; Type-species: Aphanarthrum euphorbiae Abies, Picea, Pinus, 1.6-1.9 mm Dolurgus Wollaston, monobasic]. Distribution: About

— Antennal funicle 2-segmcnted, club with no 30 species in the Canary Islands, Africa, and more than 1 suture indicated or if 2 present then India. Key: Schedl (1959a: 56). club constricted at septate suture 1 2 Deropria Enderlein [1929; 143, Type-spe- 2(1). Antennal club rather strongly constricted at cies: Aphanarthrum elon^atum Eggers, origi- middle at suture 1; elytral confused or punctures nal designation]. Distribution: 1 species in the striae, if indicated, not impressed, strial punc- Canary Islands. tures little larger than those of interstriae .3 Crypturgiis Erichson [1836: 60, Type-spe- — Antennal club not constricted at middle, suture cies: Bostrichus pusillus Gyllenhal, desig- 1 obsolete or indicated only by a partial, internal septum 4 nated by Thomson 1859: 147]. Distribution: 3 species in North America, 10 in Asia, Europe, 3(2). Elytral declivity strongly flattened to concavely Africa, and adjacent islands. All are impressed; pronotum wider than long, weakly il N monog- at all declivous on anterior half, never armed by amous and phloeophagous. Keys: Reitter granules, anterior margin unarmed; N Africa, (1913: 61) for Europe, Swaine (1918: 54) and Euphorbia; 1.6-2.6 mm Coleobothrus Wood (1982: 740) for North America, Schedl — Elytral declivity convex; pronotum usually (1946: 4) for Europe and Asia, Bright (1976: longer than wide, anterior half more strongly 114) for Canada. declivous, often armed by granules, anterior Cisurgus margin elevated as a costa, serrate, or dentate; Reitter [1894: 59, 65, Type-spe- 1.3- cies: filumReitter, monobasic]. Dis- Canary Islands, Africa to India; Euphorbia ; Cisurgus 2.3 mm Aphanarthrwn tribution: 6 species in SW Asia, S Europe, N

4(2). Antennal club unmarked by sutiu'es, anterior Africa, and the Canary Islands. Key: Schedl half of pronotum declivous and armed by coarse (1959: 28). asperities, anterior margin serrate; eye entire to feebly sinuate; Canary Islands; host unknown; Tribe Xyloterini 1.5-2.0 mm Deropria

— Antennal club with suture 2 marked at apex, if Xyloteroideae Lindemann [1876: 165, Type-genus: present, suture 1 sometimes indicated by an Xi/loterus Erichson, 1836 = internal septum; pronotum weakly declivous, Stephens, 18,30] unarmed, anterior margin never serrate; eye Trypodendron Tredl [1907: IS, Type-species: Trypoden- deeply emarginate 5 dron Stephens, 1830]

5(4). Remnants of sutures 1 and 2 usually present in antennal club; elytral striae more strongly im- Description. —Frons dimorphic, male pressed, punctures rather coarse, always in weakly to very strongly impressed, female rows; elytral vestiture on disc short, longest se- convex; eye completely divided; antennal tae shorter than distance between interstrial scape long, funicle 4-segmented (Fig. 51), rows; North America, Asia, Europe. N Africa, club without sutures, basal area sometimes Pinaceae; 0.9-1.5 mm Crypturgus corneous, derived from obliquely truncate — Antennal club usually without indications of su- type; procoxae contiguous, proepimeron of tures; elytral striae usually not impressed, punc- tures much smaller and usually contused; discal female with mycetangium; pronotum asper- vestiture longer, longest setae distinctly longer ate on anterior slope; protibia flat in male, than distance between interstrial rows; Canary inflated and armed by small unsocketed denti-

Islands, S Europe, N Africa; Euphorbia , 1.1-1.7 cles on posterior face in female; scutellum mm Cisurgus visible; elytra conservatively sculptured. Dolurgus Eichhoff [1868a: 147, Type-spe- Biology. —All species are monogamous cies: Hylastes pumilus Mannerheim, monoba- and xylomycetophagous. The male joins the sic]. Distribution: 1 species in W North Amer- female in parental galleries. The eggs are de- ica (Alaska to N California). It is monogamous posited in niches above and below in the egg and phloeophagous. 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 — Basal area of antennal club broadly, weakly pro- newly transformed adult beetle. The brood curved or else not at all corneous; protibia flat- tened and unarmed on posterior face in both emerges through the parent entrance tunnel. sexes; anterior margin of pronotuni procurved Temperate species overwinter in litter on the and armed by a series of teeth in both sexes .... 2 forest floor. 2(1). Basal area of antennal club not thickened or Taxonomy. —This is a small group consist- corneous, uniformK' pubescent to base; males ing of three small genera. They appear to have and females equal in size; anterior margin of been derived from the same parental stock pronotum slightly produced at median line in male; long axis of proepimeral excavation of fe- that gave rise to the Xylehorini. They appar- male longitudinal, short to long, narrow to ently originated in Asia, with Trypodendron rather broad; SE Asia; broadleaf hosts; 2.6-3.5 extending westward into Europe and east- inm Indocryphalus ward in late Tertiary to northern North Amer- — Basal area of antennal club subcorneous, thick- ica. (North America) and In- ened, weakly prociu'ved on anterior face; male docryphalus (Asia) were derived from the distinctly smaller than female; anterior margin of male pronotum as in female except some teeth same parental stock, with Xyloterinus appar- reduced or absent; proepimeral excavation of ently reaching North America in the warm female transverse, rather large, broad; North period that preceded the last ice age. America; broadleaf hosts; 2.7-3.7 mm . Xyloterinus

Key to the Genera of Xyloterini Trypodendron Stephens [1830: 353, Type- (Modified from Wood 1957b: .344) species: Dennestes domesticiis Linnaeus, subsequent designation by Westwood 1838: 1. Basal area of antennal club distinctly subcor-

neous, its apical margin strongly, rather nar- 39 and Thomson 1859: 146. Synonym: Xy- rowly procurved on anterior face; protibia thick- loterus Erichson 1836: 60, Type-species: ened, posterior face tuberculate in female, Bostrichus lineatus Olivier, subsequent des- flattened and usually finely tuberculate in male; ignation by Thomson 1859: 146]. Distribu- male frons broadly, deeply excavated from epis- tion: 5 species in North America, about 9 in toma to vertex, convex in female; male prono- tum subquadrate, anterior margin straight to Asia and Europe. All are monogamous and slightly recurved and unarmed, female anterior xylomycetophagous. Keys: Schedl (1951c: 86) margin procurved and armed by several teeth; for Europe and Asia, Wood (1957b: 345, 1982: long axis of proepimeral excavation of female 747) for North America. longitudinal and very narrow; North America, N Asia, Europe; coniferous and broadle;if trees; Indocryphalus Eggers [1939: 5, Type-spe- 2. 7-4.6 mm Trypodendron cies: Indocryphalus malaisei Eggers =Xy- 1986 WOOD: Genera OF ScoLYTi DAE 79 loterus intermedius Sampson, original desig- on the forest floor. At least one species nation. Synonym: Dendrotrypum Schedl (Xylehorus dispar) passes through a definite 1951c: 76, Type-species: Xijlotcrua accris Ni- dipause in the adult stage during the winter isima, subsequent designation by Browne months. 1970: 76, Type-species: Xyloterus aceris Ni- Ta.XONOMY. —The worldwide circumtropi- isima, subsequent designation by Browne cal distribution of this tribe, with a few species

1970: 562]. Distribution: About 9 species in occurring in temperate areas, suggests that it Asia (India to Japan). Apparently all are is at least moderately old. The occurrence of monogamous and .xylomvcetophagous. Key: the same species groups of in Schedl (1951c: 77).' Catalog: Browne 1970: Africa and South America indicates that the 562). basic characters of groups within that genus Xyloterinus Swaine [1918: 44, 83, Type- had been fixed by early Tertiary. However, species: Bostrichiis politus Say, original des- the distributions of other groups and the large ignation]. Distribution: 1 species in E North number of species in the tribe indicate that

America. It is monogamous and .xylomyce- very rapid, recent evolution is in progress. In tophagous. this tribe, arrhenotocous parthenogenesis is universal. This suggests a possible relation- Tribe Xyleborini ship to the higher Dryocoetini; however, a

relationship to the Xyloterini is more likely. Xylebori LeConte [1876: .346, 358. Type-genus: Xtjle- borus Eichhoff, 1864] Because of the recent, active evolution affect- Webbinae Hopkins [1915b: 224, Type-genus: Webbia ing this group, generic limits are not clearly Hopkins. 1915] defined. Common and particularly confusing features of this tribe are size races within what Description. —Body dimorphic, male otherwise appears to be a single species; in dwarfed, deformed, flightless, eye reduced in some cases these behave as entirely different size, often aberrant in shape (Fig. 52); frons species and in others, involving those same convex, unadorned; eye emarginate to di- forms, there is total intergradation of the two vided in a few oriental forms; antennal scape sizes. This is probably a product of this type elongate, funicle 5-segmented except 3- or of mating system that will be more fully under- 4-segmented in a few oriental forms, club stood when details of their habits are obliquely truncate except basal corneous area known. The generic classification of this tribe pre- reduced or absent in some genera; pronotum sented below is tentative and flawed; how- asperate on anterior slope (a few exceptions, ever, it is presented as a first attempt to orga- especially in male), procoxae varying from nize a very large and difficult group even contiguous to widely separated; scutellum though a third of the species in the tribe were varying fi-om large and flat to modified to ab- not studied. It is hoped that it might give at sent; elytra variable, conservatively to elabo- least limited direction to those who will con- rately sculptured; meso- and metatibiae flat, centrate more particularly on this group. broad, tapered on distal third, lateral margin armed by a row of numerous, small, closely Key to the Genera set socketed teeth, these usually alternating of Xyleborini (Females only) with marginal or submarginal setae in more highly evolved forms; meso- and metatarsi I. Basal segment of labial palpus eylindrical, retractible into tibial grooves. only slightly wider than segment 2 or 3, none of segments ornamented by a special tuft of Biology. —Consanguineous polygyny is setae; pregula and adjacent surfaces flush with universal, apparently all males are haploid, general contour of ventral surface of head; deformed, and flightless. All are xylomyce- antennal club rather strongly flattened, with- tophagous. Eggs are deposited in clusters in out visible sutures, corneous area reduced, the parental tunnels. The larvae usually ex- usually pubescent to base; lateral margins of tend the parental galleries or feed exclusively pronotum acutely elevated, pleural area con- cave (on transverse axis); anterior margin of on the fungal mycelium in the parental tun- pronotum unarmed; protibia inflated, armed nels. The brood emerges through the parental on its posterior face by minute tubrercles; entrance tunnel. Temperate species may Africa; 2 species introduced into America; overwinter either in the brood host or in litter 1.4-4.7 mm Premnobius 80 Great Basin Naturalist Memoirs No. 10

— Basal segment of labial palpus enlarged, con- margin usually acutely costate as on segment

spicuously wider than segment 2 or 3, its pos- 1; Mexico to South America (1 species intro-

terior face usually flat, one or more segments duced into Africa); 2. 1-.5.3 mm . . Dryocoetoides ornamented by specialized setae; pregula and — Protibia with posterior face flat, unarmed, adjacent areas usually conspicuously im- metatibia almost never with more than 7 sock- pressed below general contour of head 2 eted teeth 6 2(1). Antennal club with sutures 1 and 2 rather 6(5). Declivity commencing at or anterior to mid- strongly procurved, both segments 1 and 2 dle of elytra, its lower half transversely, corneous and mostly glabrous e.vcept at su- broadly impressed and either flat or shallowly tures; protibia slender, almost cylindrical, concave; if discal interstrial punctures unise- posterior face armed by tubercles; lateral riate then declivital surface covered by dense, margins of pronotum acutely elevated, confused, small scales, if declivital setae hair- pleural area transversely concave; body very like then discal interstrial punctures rather slender; anterior margin of pronotum armed dense, confused; India to Philippines and by 2 or more very coarse serrations; domicile New Guinea; 2.2-5.0 mm Leptoxyleborus parasites of other ambrosia beetles; Mexico to Brazil; 2.7-6.0 mm Sampsonius — Elytral declivity usually convex, commencing either posterior to middle or strongly tapered — Antennal club obliquely truncate or nearly so, on posterior half never with a strong trans- sutures (when visible) on or very near margin verse impression; elytra never ornamented of corneous area, recurved (except pubescent by scales 7 to base in some oriental forms); protibia usu- ally more strongly expanded on apical hall; 7(6). Posterior fourth ot elytra comparatively lateral margins of pronotum rounded (except broad, rather broadly rounded behind, suture Cnestiis. Webbia); "parasitic habit unknown never emarginate; declivital interstriae 1 to 3 3 similar, tubercles minute, if present; body comparatively stout, less than 2.6 times as 3(2). Scutellum visible, moderately large, its sur- long as wide; Mexico to South America; intro- face flush with adjacent surface of elytra; ven- duced to Africa; 1.7-3.0 mm Theoborus tral margin of metiifeniur either rounded or rather obtusely angulate, on its posterior face — Posterior third of elytra attenuate or acumi- groove for reception of tibia usually clearly nate, narrowly rounded behind, suture often indicated on distal half 4 emarginate; 1 or more declivital interstriae sometimes armed by small denticles; body — Scutellum either not visible or () slender, at least 2.6 times as long as wide; base ol elytra at suture notched thereby ex- Mexico and South America, Africa to SE Asia; posing cone-shaped scutellum that appears 1.5-5.0 mm Coptoborus displaced cephalad, or (couplet 20) scutellum visible on anterior face of declivital slope of 8(4). Protibia with posterior face inflated and elytral bases; scutellar area usually with abun- armed by numerous fine tubercles; posterior dant setae associated with mycetangiuni; face of antennal club either with or without a metafemur usually more strongly flattened, suture; elytral declivity and at least part of

its (longitudinal) ventral margin attenuately, disc with interstriae carinate (carinae some- very acutely angulate (except Schedlia), times reduced to rows of tubercles), strial groove for reception of tibia visible only near spaces between carinae usually granular or apical joint 16 dull, with punctures usually obsolete; SE Asia to Australia; 1.3-3.1 mm Arixyleborus 4(3). Posterior face of antennal chib marked by 2 sutures on apical third (suture 2 poorly repre- — Protibia with posterior face almost flat, sented in some Coptoborus), anterior face smooth; elytra with different sculpturing .... 9 with apical portion convex (or concave only 9(8). Antennal club with segment 2 on anterior face compara- distad from segment 2), segment 2 usually conspicuous, sometimes rather large, tively large, sclerotized, protibia armed by 6 apical margin of segment 1 on both faces or teeth lateral 7 socketed on margin, metat- rounded, often inconspicuous or absent on ibia with to 9 teeth; anterior coxae 6 socketed anterior face, almost always visible on subapi- always contiguous .5 cal area of posterior face; procoxae always con- — Posterior face of antennal club with no more tiguous, intercoxal piece longitudinally emar- than 1 suture visible at or very near apex ginate, posterior element of intercoxal piece (usually none), apical portion of anterior face never inflated or armed, mesocoxae usually

usually flat to concave, segment 2 (it visible) more widely separated by distance greater not corneous; number of metatibial teeth vari- than thickness of scape 10

able; anterior coxae contiguous or separated . 8 — Segment 1 of antennal club corneous, its dis- 5(4). Protibia with posterior face inflated and tal margin very acutely elevated into a contin- armed by numerous fine tubercles; metatibia uous costa (forming a complete circle) extend- usually with 8 or 9 socketed teeth; segment 2 ing from anterior face to apex, suture almost on antennal club usually forming a complete, never visible on posterior lace; procoxae vary-

oblique annulus, on anterior face its apical ing from contiguous to widely separated, if 1986 WOOD; Genera of Scx)lytidae 81

contiguous then posterior intercoxal piece 14(13). Lateral margin of protibia armed by 9 to 12 sometimes inflated and armed; mesocoxae socketed teeth; elytral declivity moderately to usually subcontiguous, usually separated by very strongly sulcate on at least basal half,

distance less than thickness of scape 13 lateral margins armed by at least 1 major spine and several smaller tubercles; antennal funi- 10(9). Pronotal asperities extending to base, includ- cle .5-segmented; Central and South America; ing most of discal area (except numerous 2.4-4.5 mm Taurodemus African, etc. species without discal asperi-

ties); anterior margin of pronotum never — Lateral margin of protibia armed by 4 to 7 armed by a definite row of serrations; lateral socketed teeth; elytral declivity usually not margin of porotibia armed by 7 to 8 socketed conspicuously sulcate, lateral margins not

denticles, metatibia by 8 to 11 denticles; conspicuously armed by spines or tubercles . 1.5 pantropical; 1.9-4.2 Ambrosiodmus mm 15(14). Lateral margins of pronotum rounded (sub- — Pronotal asperities confined to slightly more acute in morigerus); funicle 5-segmented; than anterior half; lateral margin of metatibia corneous area (in central portion) occupying

armed by more than 11 socketed denticles . . 11 basal third of antennal club; eye about one- third divided by an emargination; protibia 11(10). Elytra obliquely, abruptly truncate, usually armed on lateral margin by 4 to 6 socketed with an acutely, distinctly elevated circumde- teeth; pantropical, 1.3-5.0 mm ... clivital, subcircular costa, face of declivity flat to concave (when costa incomplete then an- — Lateral margins (and usually basal margin) of tennal club pubescent to base); discal inters- pronotum subacutely elevated, costate; an- triae with punctures uniseriate, occasionally tennal funicle 4-segmented (some 5-seg- slightly confused on some interstriae (strongly mented?); corneous area (in central portion) confused in superficially similar mancus occupying less than basal fourth of antennal

group oi Xijlosandrus), antennal club usually club; eye feebly if at all emarginate; protibia

pubescent to its base, more strongly flat- armed on lateral margin by 6 to 8 socketed tened; eye deeply emarginate to entirely di- teeth; SE Asia to Australia; 2.0-4.0 mm ....

vided; SE Asia to Australia; 2.4-.3. 1 mm . Amasa Cncstus

— Declivital margin rounded, a carina not indi- 16(3). Basal margins of elytra distinctly, shallowly

cated on its basal two-thirds, declivital lace emarginate in scutellar area, emargination basically convex; antennal club obliquely usually filled by dense setae, scutcllum dis- truncate (except pubescent to base in a few placed slightly cephalad into a visible, conical Xyleborus 12 process; posterior face of antennal club with-

out sutures; protibia flat, its lateral margin 2(11). Pronotum commonly subquadrate, its ante- armed by 6 to 8 socketed denticles, all on rior margin almost always unarmed, postero- apical half; strial punctures in rows; declivity lateral margin of declivit\' subacutely elevated usually armed by tubercles or spines; tropical from sutural apex to interstriae 7; strial and and temporate areas worldwide; 1.4-3.5 mm interstrial punctures usually in rows, elytral Xijleborinus vestiture comparatively sparse, confined to strial and interstrial rows; Asia and Australia — Basal margins of elytra not emarginate at su- to Africa; 2.4-4.6 mm Emcallacva ture, scutcllum not conical, with other combi- nations of characters 17 — Pronotum almost never quadrate, its anterior margin usually procurved and armed by a 17(16). Antennal funicle 5-segmented; lateral margin definite row of serrations; posterolateral mar- of protibia armed by socketed denticles on gin of declivity rounded; elytral pimctures less than apical half 18 often confused, vestiture usually more much — Antennal funicle 3- or 4-segmented; protibia

abundant; Asia to Africa; 2.1-4,1 . . . SE mm inflated but not asperate on posterior face, its Terminalinus lateral margin armed (usually to base) by sock- 13(9). Proeoxae contiguous, intercoxal piece longi- eted teeth on more than apical three-fourths

tudinally emarginate (a few individuals in /«- 23 vaniis group subcontiguous but only feebly 18(17). Pronotum asperate to base, including discal emarginate), its inflated, po.sterior element area; basal corneous area occupying almost occasionally dentate; body usually more slen- three-fourths length of club, its apical margin der, mostly 2.0 times as long as more than straight, not acutely elevated, suture 2 clearly wide; tropical areas world- and temporate indicated (club more reminiscent of Hylastcs wide; 1.7-.5.9 Xyleborus mm than oi Xyleborus). suture 1 visible near apex

— Either funicle 4-segmented or if ,5-scgmented on posterior fiice; .scutellum visible on ante- then proeoxae moderately to rather widely rior slope of elytral base; pimctiu'es on elytral separated, intercoxal piece continuous, not disc confused on basal third, in obscure rows longitudinally emarginate (a slight notch in a behind; strial and interstrial hair moderately lew Taurodemtis); body usually stouter, long; all tibiae very broad; New Zealand; 1.9

mostly less than 1 . 9 times as long as wide ... 14 mm Mesoscolytus 82 Great Basin Naturalist Memoirs No. 10

— Discal area of pronotum unarmed by asperi- — Elytra variously convex or truncate at base of ties, smooth; basal corneous area of antennal declivity, never acuminate or mucronate at club occupying less than basal half in central sutural apex; declivital margin abrupt, usually area, its apical margin recurved 19 armed; SE Asia to New Guinea and Philip- ; 1.5-3.0 mm Webbia 19(18). Posterior face of antennal club unmarked by sutures, acute costa marking apical margin of Premnobius Eichhoff [1878a: 65, 404, corneous area usually forming a complete Type-species: Fremnohius cavipennis Eich- ring; scutellum usually visible on anterior de- clivous slope of elytral margins; body stout hoff, monobasic. Synonym: Premnophilus (less than 1.8 times as long as wide), usually Browne 1962a: 79, Type-species: Xyleborus black in color; punctures on elytral disc joveri Schedl, original designation]. Distribu- dense, confused 20 tion: 24 species in Africa, 2 of them were — Posterior face of antennal club with at least 1 introduced into America. All are consan- suture visible, apical margin of corneous area guineously polygynous and xylomyceto- never costate; scutellum not visible on ante- phagous. Key: Wood (1982: 756). rior slope at bases of elytra; body more slen- der (at least 2.0 times as long as wide), color Sampsonius Eggers [1933a: 23, nomen usually yellowish to reddish brown 21 nudum, 1935: 157, Type-species: Sampsonius

20(19). Metatibia normal, socketed denticles present sexdcntatiis Eggers, original designation]. on lateral margins; metatarsus normal, nei- Distribution: 4 species in S Mexico and Cen- ther unusually long nor abnormally com- tral America, about 7 species in South Amer- pressed; declivity mostly restricted to poste- ica. All are domicile parasites of other am- rior half of elytra, convex or moderately impressed, unarmed; India to Philippines and brosia beetles and are consanguineously Australia; 4.0-6.0 mm Hadrodemius polvgynous and xylomycetophagous. Key:

— Metatibia abnormally large and broad, sock- Wood (1982: 759) for Central America. eted denticles on lateral margin obsolete; Dryocoetoides Hopkins [1915a: 10, 52, metatarsus unusually long, very strongly Type-species: Dryocoetoides guatemalensis compressed; declivity extending almost to Hopkins =Xyleborus capucinus Eichhoff, base of elytra, armed by very coarse spines; Philippines and New Guinea to Africa; 1.1-4.0 original designation]. Distribution: 2 species mm Eccoptopterus in Central America, about 22 in South Amer-

21(19). Protibia inflated and densely asperate on pos- ica 1 of which was introduced into Africa. All terior surface; anterior face of mesotibia simi- are consanguineously polygynous and xy- larly but less strongly inflated and armed; ely- lomycetophagous. Key: Wood (1982: 762) for tral disc smooth and shining, abruptly Central America. becoming rugose-reticulate on declivity, dis- cal punctures confused, declivity variously Leptoxyleborus Wood [1980: 94, Type-spe- armed by rather large spines; Sumatra to New cies: Phloeotrogus sordicauda Motschulsky, Guinea; 4.0-.5.0 mm Schedlia original designation]. Distribution: 4 species — Pro- and mesotibiae neither inflated nor in SE Asia. All are consanguineously polygy- armed on posterior face; declivity never nous and xylomycetophagous. densely, uniformly rugose-reticulate, if Theoborus Hopkins [1915a: 57, Type-spe-

armed then spines small; size much smaller . 22 cies: Theobonis theobromae Hopkins, origi- 22(21), Strial punctures in definite rows (except con- nal designation]. Distribution: 9 species in fused in bellus group); declivity convex, with- out a marginal costa, variously sculptured; Central America, several additional species in elytral scales never present; SE Asia and In- South America belong here. All are consan- donesia to Philippines; 1.4-3.4 mm guineously polygynous and xylomyce- Coptodryas tophagous. Key: Wood (1982: 770) for Central — Elytral punctures small, confused; declivity America. commencing on basal half of elytra, very Coptoborus Hopkins [1915a: 10, 53, Type- strongly concave, its marginal crest forming a blunt, elongate, circumdeclivital costa, inner species: Coptobortts emar^natus Hopkins face ornamented by abundant, small scales or =Xyleborus oespatorius Schedl, original des- longhair; Indonesia; 3.3-3.5 mm ignation. Synonym: Streptocranus Schedl Taphrodasus 1939a: 52, Type-species: Streptocranus 23(17). Elytra moderately acuminate on declivity and mirabihs Schedl, luonobasic]. Distribution: 5 mucronate at sutural apex; base of declivity species in Central America, about 10 in South rounded, unarmed; base of elytra often with a submarginal mycetangium; Malaya; 2.2 mm America, and about 6 in SE Asia and adjacent Crijptoxylehorus islands. All are consanguineously polygynous 1986 VVooD: Gener\ of Scolytidae 83

and xylomycetophagous. Key; Wood (1982: Xyleborus Eichhoff [1864; 37, Type-spe- 780) for Central America. cies: Bostricluis itKiiiograjihus Fabricius, sub- Arixyleborits Hopkins [1915a: 10, 59, Type- sequent designation by Lacordaire 1866; 381. species: Arixyleborus rugosipes Hopkins, Synonyms; Anisandrus Ferrari 1867; 24, original designation. Synonym: Xyleboriciis Type-species: Xyleborus (Apate) dispar Fab- Eggers 1923: 212, Type-species: Xylcboricus ricius, monobasic; Anaeretus Duges 1887: canaUculatus Eggers, subsequent designa- 141, Type-species: Xyleborus guanajuatensis tion by Schedl 1936b: 64]. Distribution: Duges =Bostrichus volvulus Fabricius, About 42 species in SE Asia to Australia and monobasic, neotype for type-species desig- the Philippines. All are consanguineously nated by Wood 1983: 650; Progenius Bland- polygynous and xylomycetophagous. ford 1896a: 20, Type-species; Progenius fleu- Ambrosiodmus Hopkins [1915a: 10, 55, tiauxi Blandford =Xyleborus subcostatus Type-species: Xyleborus tachygraphiis Zim- Eichhoff, subse(|uent designation by Hopkins mermann, original designation. Synonyms: 1914; 128; Hagedorn 1912: Phloeotrogits Motschulsky 1863: 127, Type- 355, Type-species; Cyclorhipidion pelliculo- species: Phlocotrogus obhquccaudata Mot- sum Hagedorn -Xyleborus prelaetus Schedl, schulsky, designated by Hopkins 1914: 127, monobasic; Heteroborips Reitter 1913; 79, International Commission on Zoological 82, Type-species: Bostrichus cryptographus Nomenclature 1979: 151, name rejected; Ratzeburg, monobasic; Xyleborips Reitter Browneia Nunberg 1963; 37, Type-species; 1913: 79, 111, Type-species; Xyleborus Xyleborus illepidus Schedl ^Pityophthorua meuscli Reitter, monobasic; Boroxylon Hop- obliquus LeConte, original designation]. Dis- kins 1915a: 10, 58, Type-species; Boroxylon tribution; 11 species in North and Central stephegynis Hopkins Phloeotrogus biden- America, about a dozen in South America, tatus Motschulsky, original designation; numerous species (40 or more) in Asia and Notoxyleborus Schedl 1934; 84, Type-species: Africa. All are consanguineously polygynous Notoxyleborus kalshoveni Schedl, monoba- and xylomycetophagous. Key: Wood (1982: sic]. Distribution; Several hundred species 781) for North and Central America. almost worldwide. All are consanguineously Amasa Lea [1894; 322, Type-species: polygynous and xylomycetophagous. Keys; Amasa thoracicus Lea, monobasic. Syn- Bright (1968; 1296, 1976; 131) for North onyms: Pseudoxylebonis Eggers 19.30; 206, America, Wood (1982; 778) for North and Type-species; Pseudoxylebonis beesoni Eg- Central America, Reitter (1913; 81) for Eu- gers, monobasic; Anaxyleborus Wood 1980: rope. 90, Type-species; Tomiciis truncatus Erich- Taurodetnus Wood [1980: 96, Tvpe-spe- son, original designation]. Distribution: cies: Xyleborus sharpi Blandford, original About 35 species in India and Malaya to Aus- designation]. Distribution; 14 species in S tralia. All are consanguineously polygynous Mexico to South America. All are consan- and xylomycetophagous. guineously polygynous and xylomyceto- Euwallacea Hopkins [1915a: 10, 54, Type- phagous. Key: Wood (1982: 778) for Central species; Xyleborus loallacei Blandford, origi- America. nal designation]. Distribution: About 50 or Xylosandrus Reitter [1913; 80, 83, Type- more species in Africa to SE Asia and Aus- species; Xyleborus morigerus Blandford, tralia. All are consanguineously polygynous monobasic. Synonym; Apoxyleborus Wood and xylomycetophagous. 1980: 90, Type-species; Xyleborus mancus Terminalinus Hopkins [1915a: 10, 57, Blandford, original designation]. Distribu- Type-species: Terminalinus terminaliae Hop- tion; 8 species in USA to South America, kins, original designation. Synonyms; Kelan- about .32 species in SE Asia to Australia and tanius Nunberg 1961; 621, Type-species: Africa. All are consanguineously polygynous Xyleborus punctatopilosus Schedl, original and xylomycetophagous. Key: Wood (1982: designation]. Distribution; About 30 or more 765) tor North and Central America. species in Africa to SE Asia and Australia. All Cnestus Sampson [1911: 383, Type-spe- are consanguineously polygynous and xy- cies; Cnestus magnus Sampson, monobasic. lomycetophagous. Synonym: Tosaxyleburus Murayama 1950a; 84 Great Basin Naturalist Memoirs No. 10

49, Type-species: Tosaxylcboriis pallidipcn- species in SE Asia to Australia and adjacent nis Murayama =Cnestus miiraijamai Schedl, islands. All are consanguineously polygynous original designation]. Distribution: About 17 and xylomycetophagous. species in SE Asia to Indonesia, the Philip- Taphrodasiis Wood [1980: 95, Type-spe- pines, and Japan. All are consanguineously cies: Xyleborus percorthylus Schedl, original polygynous and .xylomycetophagous. Kev: designation]. Distribution: 3 species in SE Nunberg (1972: 476). Asia to Indonesia. All are consanguineously Xyleboriniis Reitter [1913: 79, 83, Type- polygynous and xylomycetophagous. species: Bostrichus saxeseni Ratzeburg, sub- Crijptoxijleborus Schedl [1937b: 550, sequent designation by Swaine 1918: 50]. Dis- Type-species: Crijptoxijleborus naevus tribution: 8 species in North and Central Schedl, subsequent designation by Schedl America, at least 4 in South America, at least 1962a: 103]. Distribution: About 12 species in 20 in SE Asia to Africa. All are consan- Malaya. All are consanguineously polygynous guineously polygynous and .xylomyceto- and xylomycetophagous. phagous. Key: Wood (1982: 842) for North and Webbia Hopkins [1915b: 222, Type-spe- Central America. cies: Webbia dipterocarpi Hopkins, original Mesoscolytus Broun [1904: 125, Type-spe- designation. Synonyms: Xelyborus Schedl cies: Mesoscolijtiis inurbanus Broun, mono- 1939b, nomen nudum (Browne 1963a: 57); basic]. Distribution: 1 species in New Pseudowebbia Browne 1961: 308, Type-spe-

Zealand. It is consanguineously polygynous cies: Xyleborus trepanicauda Eggers, original and xylomycetophagous. designation; Prowebbia Browne 1962b: 208, Hadrodemius Wood [1980: 94, Type-spe- Type-species: Proivebbia subuculae Browne, original cies: Xylebonis globus Blandford, original designation]. Distribution: About 32 designation]. Distribution: About 6 species in species in SE Asia to Indonesia and the Philip- SE Asia to Indonesia and the Philippines. All pines. All are consanguineously polygynous are consanguineously polygynous and xy- and xylomycetophagous. Key: Browne lomycetophagous. (1962b: 210) to the pabo group of species. Eccoptopterus Motschulsky [1863: 515, Tribe Xyloctonini Type-species: Eccoptopterus sexspinosiis Motschulsky =Scohjtits spinosits Olivier, Xyloctonidae Eichhoff [187Sa: 171. Tvpe-genus: Xyloc- monobasic. Synonyms: Platydactyhis Eich- fonus Eichhoff, 1872] hoff 1886: 25, Type-species: Platydactylus Description. —Frons apparently not di- gracilipes Eichhoff, monobasic; Eurydactyhis moiphic, usually unadorned; eye emarginate Hagedorn 1909: 733, Platydactylus gracilipes to divided; antennal scape elongate, funicle 6- Eichhoff, automatic]. Distribution: About 12 or 7-segmented, club strongly flattened, su- species in SE Asia to Australia and adjacent tures procurved, present or obsolete, 1 partly islands, and Africa. All are consanguineously septate or not; pronotum asperate on anterior polygynous and xylomycetophagous. slope, anterior margin usually armed; pro-

Schedlia Browne [1950: 641, Type-species: coxae contiguous; scutellum large, flat; tarsi Xyleborus sumatranus Hagedorn, original retractable into tibial grooves; venter of abdo- designation]. Distribution: 5 species in SE men moderately to very strongly ascending to Asia to Indonesia and New Guinea. All are meet elytra. consanguineously polygynous and xylomyce- Biology. —All are apparently monoga- tophagous. Key: Browne (1950: 642). mous and phloeophagous. The egg galleries Coptodryas Hopkins [1915a: 10, 54, Type- are monoramous in Ctonoxijlon and biramous species: Coptodrijas confusa Hopkins, origi- in Scolijtomimus . The eggs are deposited in- nal designation. Synonyms: Microperus dividually in niches and sealed in by frass. The Wood 1980: 94, Type-species: Xyleborus larval mines radiate out from the parental theae Eggers, original designation; Adry- gallery and may be rather long. ocoetes Eggers, nomen nudum, in Schedl Taxonomy. —This is a small group of pre- (1952: 371), Type-species: Adryocoetes ni- dominantly African genera that are poorly tidus, nomen nudum, =Xtjleborus pullus known. It appears to be a primitive branch of Schedl, no status]. Distribution: About 20 the same phyletic line that gave rise to the ;

1986 WooD; Genera of Scolytidae 85

Cryphalini. Together these two tribes appear area on interstriae 1 and 2 impressed thereby causing scutellum to project slightly dorsad; to occupy a position intermediate between Africa; 1.3-2.8 mm Xyloclonus primitive Micracini and Corthyhni. A princi- pal distinguishing character oi Xyloctonini is Cryphalomimus Eggers [1927b: 174, Type- their abihty to totally withdraw the meso- and species: Cnipluilomimus striatum Eggers, metatibiae into tibial grooves. This character monobasic]. Distribution: 3 species in Africa is shared by a small group of genera in the (Congo to East Africa). Cryphalini that are allied to Scolytogenes Glostatus Schedl [1939d: 386, Type-spe- thus, tribal placement for them is dependent cies: Glostatiis decUvidepressiis Schedl, on the number of segments in the antennal monobasic. Synonyms: Ctonocryplms Schedl funicle. The limited distribution of this tribe, 1941: 398, Type-species: Ctonocryplms xyloc- accompanied by conspicuous anatomical di- tomis Schedl, monobasic; Apoglostutus veristy, makes comments on its anticjuity diffi- Schedl 1957a: 155, Type-species: Apoglo- cult. statiis acaciae Schedl, monobasic; Para- glostaus Schedl 1964c: 304, Type-species: Key to the Genera of Xyloctonini Ctonocryplms uigrivcstris Schedl, original

1. Eye entire to feebly emarginate; suture 1 of designation; Rhopalocryplms Nunberg 1967: antennal club septate, almost straight; funicle 320, Type-species: Rhopalocryphus seycleli 7-segmented; elytra declivous behind, abdo- Nunberg]. Distribution: 16 species in Africa. men raised only slightly to meet them, inters- Apparently all are monogamous and phloeo- triaecostate; Africa; 1.5-2.1 mm . Cryphalomimus phagous. — Eye moderately emarginate to divided; sutures Ctonoxylon Hagedorn [1910c: 4, Type-spe- of antennal club weakly to strongly procurved .. 2 cies: Ctonoxylon auratiim Hagedorn, subse- 2(1). Eye moderately emarginate, about one-third di- vided by an emargination; basal (usually) and quent designastion by Hopkins 1914: 119, lateral margins of pronotum rounded; funicle Schedl's (1961: 426) designation is invalid]. 6-segmented, sutures of antennal club feebly to Distribution: About 32 species in Africa. Ap- moderately procur\ed, 1 not septate; elytral de- parently all are monogamous and phloeo- clivity steep, abdomen ascending very slightly; phagous. Africa; 1.9-2.6 mm Glostatiis Scolytomimus Blandford [1895: 319, Type- — Eye more than half divided by an emargination to completely divided; basal and lateral margins species: Scolytomimiis dihitiis Blandfoi'd, of pronotum with a fine, raised line (either con- monobasic. Synonyms: Neoxyloctonus Eggers tinuous or beaded); sutures of antennal club 1923: 143, Type-species: Neoxyloctonus strongly procurved, 1 partly to entirely septate philippincnsis Eggers, monobasic; Scolyto- (when sutures reduced septum remains); abdo- clcptes Schedl 1962f: 490, Type-species: men moderately to very strongly ascending to

meet moderately to feebly declivous elytra .... .3 Scolytoniinms maculatns Beeson, original

3(2). Antennal funicle 7-segmented, club moderately designation]. Distribution: About 15 species flattened, asymmetrical, comparatively small, from India and Sri Lanka (Ceylon) to Samoa. about equal in length to scape, suture 1 with All are monogamous and phloeophagous. posterior half septate; elytral declivity rather Xyloctonus Eichhoff [1872: 134, Type-spe- steep, abdomen ascending moderately to meet cies: Xyloctonus scolytoidcs Eichhoff, mono- apex; eye always divided; Africa; 1.8-.3.6 mm . . Ctonoxylon basic]. Distribution: 14 species in Africa. All

— Antennal funicle fj-segmented, club strongly are monogamous and phloeophagous. Key: flattened, symmetrical, large, conspicuously Menier (1974: 658). longer than scape, suture 1 partly septate or not; elytral declivity short, very gradual, abdomen Tribe Cryphalini very strongly ascending to meet apex; eye di- vided or not 4 Cryphaloidea Lindemann [1876: 16.5, Type-genus: 4(3). Antennal club devoid of sutures except for Cniphalus Erichson. 1836] strongly procurved septum in posterior half of Trypophloeinae Niisslin [1911: 373, Type-genus: Try- suture 1; scutellum large, flat, subtriangular. its pophldcufi F'airmaire, 1868] surface flush with that of base of elytra, India to Ernoporinae Niisslin (1911; 375, Type-genus: Ernoponis

Philippines and Fiji; 1.0-2.4 mm . , Scohjtomimus Thomson, 1859] — Antennal club with 2 or 3 very strongly pro- Eidophelinae Murayama [1954; 200, Type-genus: Eido- curved sutures, none of them septate; scutellum phcltifi EichhofI, 1875, amended from Eidopheri- averaging smaller, subquadrate, adjacent basal 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, North America were recent dcrivitives from usually convex, usually not adorned; eye usu- the neotropical realm. The South American ally entire, shallowly emarginate in a few gen- element of Cryphalini appears to contain very era; antennal scape elongate, simple, funicle ancient, endemic elements {Acorthylus, 3- to 5-segmented, club moderately to Ncocnjphus, Stegomerus, ) strongly flattened, sutures present or obso- and more recent arrivals (Hypothenemiis, lete, 1 sometimes septate, sutures on poste- Scolytogenes). The more recent elements are rior face strongly displaced toward apex; largely confined to the tropics and are shared pronotum coarsely asperate on anterior slope with Africa; similarity of species groups sug- (Fig. 53), anterior margin usually armed, lat- gest a connection of faunas as late as early eral and basal margins usually (not always) Tertiary. The main center of distribution for marked by a finely elevated line; procoxae the tribe appears to be in the Oriental and contiguous; tibiae rather strongly flattened, Australian realms. This is also the area where their lateral margins armed by more than four rapid evolution has produced a great abun- socketed teeth; metepisternal spine usually dance of very closely related species. In this modified, often partly replaced liy a callus or area, generic limits are often obscure, modified groove; scutellum large, flush with whereas in other parts of the world they are adjacent surface of elytra; basal margins of quite distinct. This is probably the most elytra sometimes marked by a fine, raised poorly known tribe in the world and it is likely line; elytral sculpture usually conservative, to remain so until much more material is avail- their costal margins near apex usually ascend- able for study. ing at least slightly; venter of abdomen usually Key to tlie Genera of Cryphalini rising slightly to meet elytra; vestiture com-

monly includes scales. 1 . Basal and lateral niargin.s ot pronotum Biology. —Monogamous, except for con- rounded; procoxae usually narrowly sepa- rated, intercoxal piece continuous (except sanguineous polygyny in Hypothenemiis, contiguous and intercoxal piece longitudi-

Crtjptocarentts, Feriocryphahis , and Trischi- nally emarginate in Trypophloeiis, Ste- dias, and either phloeophagous or myelo- gomerus, Acoi-thijlus) 2 phagous. Mycetophagous in one species of — Basal and usually basal third of lateral margins

Hypothenemiis . Parental gallery usually of a of pronotum marked by a finely raised line; cave type, often modified into crude, elon- procoxae either contiguous or narrowly sepa- rated; eye entire or narrowly emarginate .... 8 gate, brood chambers. The eggs are deposited twice as wide, in clusters, often mixed with frass; definite 2(1). Eye shorter, less than long as entire (rarely with a few lacets absent suggest- egg niches have not been reported. The larvae ing a weak, narrow emargination) 3 may form individual mines {Trypophloeiis, — Eye moderately elongate, 2.0 or more times Cryphalus) or feed in congress as they extend as long as wide, more than half of its anterior the parental chamber (ProcrypJialus, Hy- margin occupied by a sinuation or broad pothenemiis, etc.). The brood may emerge emargination one-third as deep as width of through individual exit holes or through the eye; neotropical genera 6 parental entrance timnel, depending upon 3(2). Antennal funicle 5-segmented, club slender the larval habit. In temperate areas the winter (1.8 or more times as long as wide), 2 sutures clearly marked, its apex subacutely pointed is most commonly passed in the larval stage; (Fig. .52); elytral interstriae 10 continuing to two generations per year is the common habit. apex; North America, N Asia, Europe; Salix, Taxonomy. —The worldwide distribution Popiilus, AIniis, 1.4-2.1 nun Trypophloeiis and structural diversity of this tribe suggest a — Antennal lunicle 3- to 5-segmented, club pre-Tertiary origin. However, the prolifera- broad (less than 1.3 times as long as wide), its tion of large numbers of species, particularly apex rather Inoadly rounded, suture 1 usually marked, 2 obsolete or nearly so; interstriae 10 in Cryphalus, indicates that recent, very obsolete before attaining level of base ol ab- rapid evolution is in progress. Trypophloeiis, dominal sternum 5 4

Procryphalus, Ernoporicus , and Cryphalus 4(3). Eye entire; antennal club longer, basal area appear to have reached North America from more stnmgly flattened, suture 1 straight,

Asia rather recently. Hypoihencmus , Crypto- septate. 2 obsolete; funicle 4-segmented; fe- carenus, Trischidias , and Scolytogenes in male irons not conspicuously pubescent; 88 Great Basin Naturalist Memoirs No. 10

North America to NE Asia; , Salix. — Antennal funicle 3-segmented; vestiture of Fraxinus; 1.3-2.2 mm Procryphaliis uniseriate rows of scales on discal interstriae, a few supplemental scales on declivity; su- — Eye entire to weakly sinuate; antennal club tures on antennal club rather weakly pro- with sutures procurved or absent, 1 not sep- curved, sometimes obscure; SE Asia; tate when present .5 broadleaf trees; 1.1-1.5 mm Ernocladius 5(4). Antenna! funicle 4-segmented, club with pro- 12(8). Posterior face of metatibia (usually also curved sutures indicated by rows of setae; E mesotibia) with a groove for reception of tar- North America, N Asia, Europe; Fraxinus, sus on lateral half from apex at least two-thirds Fagus, Cornus; 1.2-1.7 Ernoporicus mm of distance toward base, grooved area

— Antennal funicle .3-segmented, club devoid of glabrous, usually with a row of setae along its

sutures; NE Asia; Euomjmus ; 1.4 mm mesal margin, tibia usually more broadly flat- Allernoporus tened, gradually tapered on its distal third, with socketed teeth more numerous and dis- 6(2). Antennal huiicle .5-segmented, club large, tributed over at least apical third (possible subcircular, about as wide as long, sutures confusion with Xyloctonini, except eye entire conspicuously procurved; discal striae not im- in these Cryphalini); male subequal in size to pressed, poorly defined (punctures often con- female, capable of flight 13 fused); Mexico to South America; vines Metatibia either without groove (lianas); 1.1-1.9 mm Stegomerus — for reception of tarsus or groove restricted to less than distal — Antennal funicle 3-segmented, club elongate, one-fifth of tibial length, setae randomly dis- at least 1.5 times as long as wide; sutures tributed on its lateral half, tibia usually sub- almost straight 7 truncate apically, socketed teeth usually re-

7(6). Antennal funicle shorter than scape, seg- stricted to apical one-fifth; male either normal ments 2 and 3 small, subequal in size; elytral or deformed 16 striae rather coarse; S impressed, punctures 13(12). Antennal club with suture 1 partly septate, 1.4-1. South America; host unknown; .5 mm suture either straight or strongly procurved, if Neocryphux straight then funicle 3-segmented); body usu- Antennal fimicle with segment 2 greatly en- ally stouter, pubescence moderately abun- larged, as long as scape; elytral striae not im- dant; groove foriTiing lateral line on pronotum pressed, strial rows usually not distinguish- rather poorly defined 14

able (at least one exception); South America; — Antennal club with suture 1 aseptate (or ob- Pruntis. etc.; 1.2-1.8 mm Acorthylus scurely septate, if suture present then funicle 4-segmented; body rather slender, elytral 8(1). Basal margin of pronotum marked by a fine, vestiture sparse, largely confined to declivity; raised line, lateral margin rounded; sutures groove forming lateral and basal raised line on on antennal club procur\'ed, usually distinct . 9 pronotum rather strongly impressed 15 — Both basal and lateral margins of pronotum 14(10). Antennal funicle 3-segmented, club with su- marked by a fine, raised line; sutures on an- tures 1 and 2 weakly procurved, clearly tennal club present or absent, variable 12 marked by setae, 1 also grooved and partly 9(8). Eye emarginate; pronotal asperities confused; septate; venter of abdomen horizontal; India antennal grooves or sutures moderately pro- to Sri Lanka (Ceylon); Euphorbia, 1.0-1.3 curved 10 mm Cryphalogenes

— Eye entire; pronotal asperities arranged in — Antennal funicle 4-segmented, club with su- concentric rows; antennal svitures rather tures strongly to profoundly procurved, 1 strongly to profoundly procurved or obsolete marked on mesal half by a septum, remaining ' 11 sutures marked by setae or obsolete (septum complete in several small New Guinea spe- 10(9). Antennal funicle 5-segmented, club with su- cies); venter of abdomen weakly to very tures 1 and 2 marked by moderately pro- strongly ascending to meet apex of elytra, curved grooves and rows of setae; pronotal apex of elytra usually ascending also; pantrop- asperities coarse, confused; eye conspicu- ical; hosts usually lianas; 1.0-2.5 mm ously emarginate; Africa; hosts unknown; 2.0- Scolytogenes 2.3 mm Stephanopodius 1.5(13). Antennal club with sutures 1 and 2 weakly Antennal funicle club with su- — 4-segmented, procurved and clearly marked by rows of se- ture 1 septate angulate; eye rather small, and tae, 1 also grooved; basal half of pronotum shallowly emarginate; SE Asia; 1.2-1.6 mm reticulate or minutely rugose, punctures Coriacephilus small to obsolete; Micronesia; hosts unknown;

11(9). Antennal funicle 4-segmented; vestiture of 1.1-1.4 mm Hemicryphalus abundant, confused scales; sutures on anten- — Antenuiil club with sutures entirely obsolete; nal club very strongly procurx'ed or obsolete; basal hall of pronotum smooth, shining, with a Europe, Asia; Tilia, Fagus, etc.; 1.1-1.5 mm few coarse punctures; India and E USSR to

Ernoporus 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- — Anterior margin of pronotum usually armed coxae narrowly separated, interco.\al piece by 1-8 serrations; antennal hmicle 3- to 5-seg- not longitudinally emarginate; eye emar- mented, when 5-segmented then suture 1 of ginate; antennal club with aseptate sutures club partly .septate; antennal club clearly clearly marked by grooves and setae 17 marked by sutures; mature color usually darker, vestiture more abundant; phloeo- — Tarsal segment 3 narrow, often laterally coin- pressed, not bilobed, procoxae contiguous, phagous, myelophagous, spermopliagous, 1 worldwide in intercoxal piece longitudinally emarginate or mycetophagous, tropical and 0.6-2.8 partly absent 18 warm temperate areas; mm Hypothenemus 17(16). Antennal funicle 5-segmented, sutures on club weakly to rather strongly procurved; 23(21). Antennal club with sutures clearly marked by phloeophagous in broadleaf trees; pantropi- rows of setae; eye entire; phloeophagous in cal; 1.1-2.8 mm trees and shrubs; E North America to Mexico and Hawaii; 0.6-1. 1 mm Trischidias — Antennal funicle 4-segmented, sutures on club recurved (occasionally weakly pro- — Antennal club without indications of sutures; curved); phloeophagous in broadleaf and eye emarginate, myelophagous in \'ines. coniferous hosts; E Hemisphere, North South America; 0.8-1.1 mm ... Periocryphaltts

America (1 dubious record from South Amer- ica); 1.0-2.8 mm Cryphalus Fairmaire [1868; 105, Type- 18(16). Antennal funicle 3-segmented, club with or .species: Bostrichiis binodiihis Ratzehurg, without sutures, never with a septum, male monobasic. Synonym; Glyptoderiis EichhofF normal, not dwarfed or flightless 19 1878a: 34, 44, 137, Type-species; Bostrichiis — Antennal funicle 3- to .5-segmented, sutures binodulus Ratzeburg, subsequent designa- always indicated on club, 1 frequently partly tion by Hopkins 1914; 122]. Distribution; 4 septate; male dwarfed, flightless 21 species in North America, about 12 in N Asia 19( 18). Antennal club with moderately procurved su- tures clearly marked by grooves and rows of and Europe. All are monogamous and setae; eye emarginate; striae obsolete, pinic- phloeophagous. Keys; Wood (1954b; 989, tures on disc confused; SE Asia to Philippines; 1982; 851) for North America, Reitter (1913; 0.8-1.3 mm Margadillius 69), Balachowsky (1949; 214), Stark (1952: — Antennal club without indications of grooves 281) for Europe and Asia. or rows of setae; eye entire, strial punctures in recognizable rows 20 Hopkins [1915a; 7, 33, Type- species; Procryphahis popiili Hopkins = 20(19). Anterior margin of pronotum armed by 0-6 serrations, asperities on anterior half of Cryphalus miicronatus LeConte, original pronotum larger, less numerous; body designation]. Distribution; 2 species in North stouter, 1.4-2.0 times as long as wide; Irons America (in Popuhts, Salix), 1 in NE Asia convex to flattened in both sexes; SE Asia to (Ussuri, in Fraxiniis). All are monogamous Hawaii; O.S-1.8 mm Ptilopodius and phloeophagous. Key; Wood (1954b; 982, — Anterior margin of pronotum armed by 10-16 serrations, asperities smaller, much more nu- 1982; 859) for North America. merous, often extending to base; body more Ernoporicus Berger [1917; 242, Type-spe- slender, 2.5-2.6 times as long as wide; female cies: Ernoporiciis spessivtzevi Berger, mono- frons rather narrowly impressed; SE Asia and basic. Synonyms: EocryphahiS Kurentzov Indonesia to Micronesia; 1.3-1.5 mm Cosmoderes 1941: 161, 230, Type-species; Eocrypliahis se- menovi Kurentzov, monobasic; Ernopocerus 21(18). Antennal funicle 3- to 5-segmented, club with or without sutures, when funicle 3-seg- Balachowsky 1949: 211, Type-species: mented then club always with suture 1 partly Ernoporus caucasicus Lindemann, subse- septate 22 quent designation by Wood 1954b: 986]. Dis- — Antennal funicle .3-segmented, club never tribution; 1 species in North America (West septate, sutures sometimes marked by rows of Virginia), about 12 in Europe and Asia. All are setae; body rather stout, 2.0-2.3 times as long apparently monogamous and phloeophagous. as wide; very small 23 Keys: Balachowsky (1949: 211), Reitter (1913; 22(21). Anterior margin of pronotum armed by 10-16 68)', serrations; antennal funicle 5-segmented, Stark (1952: 271). club never septate, sutures marked by rows of Allernoporus Kurentzov [1941: 159, Type- setae; mature body color yellowish to reddish species; Alh'rnoponis eiionymi Kurentzov, brown; vestiture usually very sparse (rare ex- monobasic]. Distribution; 1 species in NE ceptions); myelophagous; tropical America (1 introduced to Africa); 1.4-3.3 mm Asia (in Euonynuts). It apparently is monoga- Cryptocareniis mous and phloeophagous. 90 Great Basin NATUiiALiST Memoirs No, 10

Stegomerus Wood [1967; 129, Type-spe- nal designation]. Distribution: 4 species in

cies; Stegomerus vulgaris Wood, original des- India to Sri Lanka (Ceylon) in Euphorbia . All ignation]. Distribution; 5 species in Mexico are monogamous and excavate dying tissue and Central America, 1 in South America immediately under the epidermis. (Venezuela). All are monogamous and Scohjtogenes Eichhoff [187Sa; 475, 497, phloeophagous. Keys: Wood (1967: 130, 1982; 1878b: 387, Type-species; Scolytogenes dar- 855). wini Eichhoff, monobasic. Synonyms: Lep- Neocryphus Nunberg [1956a; 139, Type- icerus Eichhoff 1878a; 476, 501, Type-spe- species; Neocryphus argentinensis Nunberg, cies: Lepicenis aspericollis Eichhoff, mono- original designation]. Distribution: 2 species basic; C ryphalomorphus Schaufuss 1890; 12, in South America (Argentina). Type-species: C ryphalomorphus communis Acorthyltis Brethes 1922a: 304, Type-spe- Schaufuss, monobasic; Letznerella Reitter cies; Acot-tlnjliis asperatiis Brethes, monoba- 1913; 68, Type-species: Bostrichus jalapae sic. Synonym: Phacryliis Schedl 1938a: 24, Letzner, monobasic; Hypothenoides Hopkins Type-species: Phacryliis hosqid Schedl, 1915a: 7, 11, Type-species; Hypothenoides monobasic]. Distribution: About 5 species in parvus Hopkins, original designation; South America. All are monogamous and Ernoporides Hopkins 1915a: 8, 34, Type-spe- phloeophagous. cies: Ernoporides floridensis Hopkins Stephanopodhis Schedl [1941: 396, Type- = Ernoporides knabi Hopkins, original desig- species; Stcphanodcres dispar Eggers, subse- nation; Neocryphalus Eggers 1922: 169, quent designation by Schedl 1961: 633. Syn- Type-species; Neocryphalus usagaricus Eg- onyms; Cryphalomimus Browne 1962a; 75, gers, monobasic; Negritus Eggers 1923; 141, Type-species: Hypocryphalus ghanaensis Type-species; Negritus ater Eggers, desig- Schedl, original designation, preoccupied; nated by Wood 1982: 861; Cylindrotomicus C ryphalomiinetes Browne 1963b: 242, Type- Eggers 1936: 6.33, Type-species: Cylindroto- species: Hypocryphalus ghanaensis Schedl, micus squamulosus Eggers, monobasic; Lep- automatic]. Distribution: About 7 species in icerinus Hinton 1936: 473, Type-species: Eep- Africa. icerus aspericollis Eichhoff, automatic; Coriacephilus Schedl [1939b; 339, Type- Xylocryptus Schedl 197.5c: .352, Type-species: species: Stephanoderes coriaceiis Eichhoff, Xylocryptus papuanus Schedl, original desig- original designation]. Distribution; About 4 nation]. Distribution; 7 species in North and species in SE Asia to Philippines. Central America; about 60 in other tropical Ernoporus Thomson [1859: 147, Type-spe- and subtropical areas around the world. All all in cies: Bostrichus tiliae Panzer, original desig- are monogamous and almost bore the nation. Synonyms; C ryphalops Reitter 1889: stems of vines (lianas). Key: Wood (1960: 27) 94, Type-species; Cryphahis lederi Reitter for Micronesia, (1982; 862) for North and Cen- = Bostrichus tdiae Panzer, monobasic; Steph- tral America. anorhopalus Hopkins 1915a; 35, Type-spe- Hemicryphalus Schedl [1963b: 264, Type- cies: Stephanorhopcdus melodori Hopkins, species; Eidophelus argutus Wood, original original designation (specific name of type- designation]. Distribution; 3 species in Mi- species validly amended by Schedl 1966a: 19); cronesia (W Pacific Islands). Key: Wood Euptilius Schedl 1940b; 589, Type-species; (1960a; .32). Ernoporus concentralis Eggers, original des- Eidophelus Eichhoff [1875: 200, Type-spe- ignation]. Distribution: At least 13 and per- cies: Eidophelus iniitans Eichhoff, monoba- haps 20 species in Europe and S Asia to the sic. Synonym: Phellodendrophagus Krivolut- Philippines. All are monogamous and phloeo- skaya 19.58; 150, Type-species: Phellodend- phagus. rophagus elegans Krivolutskaya, monobasic]. Ernocladius Wood [1980: 93, Type-species: Distribution: About 8 species in SE Asia and Cryphalus corpulentus Sampson, original Indonesia. designation]. Distribution: About 4 species in Hypocryphalus Hopkins [1915a; 8, 41, S Asia to Sri Lanka (Ceylon). Type-species; Hypocryphalus rotundus Hop- Cryphalogenes Wood [1980: 91, Type-spe- kins, original designation. Synonym: Dacry- cies; Cryphalogenes cuphorbiae Wood, origi- phalus Hopkins 1915a: 8, 42, Type-species: 1986 WOOD: Genera of Scolytidae 91

Dacryphahis ohesus Hopkins, original desig- Ptilopodius Hopkins [1915a: 7, 11, Type- nation]. Distribution: About 47 species in species: Ptilopodius stephegynis Hopkins, Africa, S Asia to Australia and Samoa; 1 intro- original designation]. Distribution: 15 species duced into America {mangiferae , in Mang- in Africa, SE Asia, and adjacent islands have ifera indica ). All apparently are monogamous been assigned to this genus, some erro- and phloeophagous. neously. Key: Wood (1960a: 18) for Microne- Cryphalus Erichson [1836: 61, Type-spe- sia. cies: Bostrichus asperatus Gyllenhal, subse- Cosmoderes Eichhoff [1878a: 495, Type- quent designation by Thomson 1859: 147, lec- species: Cosmoderes inonilicollis Eichhoff, totype for type-species designated by Wood monobasic. Synonyms: Erioschidias Schedl 1972; 41. Synonyms: Pseudocryphalus Fer- 1938b: 42, Type-species: Cryphalus setistria- tus rari 1869: 252, Type-species: Bostrichus sid- Lea, subsequent designation by Wood neyanus Nordlinger, monobasic; Taenio- 1960a: 21; Dendriops Schedl 1953b: 125, Type-species: granulicollis glyptes Bedel 1888: 398, Type-species: Dendriops Schedl, Bostrichus abietus Ratzeburg =Bostrichus monobasic; Vitaderes Beeson 1941: 301, nomen nudum. Type-species: Vitaderes asperatus Gyllenhal, original designation;

, nomen nudum =Cosinoderes inonili- Cryptarthrum Blandford 1896b: 200, Type- luffa collis Eichhoff, no status]. Distribution: species: Cryptarthrum walkeri Blandford, About 22 species in Africa to SE Asia and monobasic; Allarthrum Hagedorn 1912: 355, Australia. At least 2 species were taken from Type-species: Allarthrum kolbei Hagedorn, lianas. monobasic; Ericryphalus Hopkins 1915a: 8, Crijptocarenus Eggers [1937: 79, Type- 38, Type-species: Ericryphalus henslunoi species: Cryptocarcnus diadematus Eggers, Hopkins =Hypotheneinus sylvicola Perkins, original designation. Synonym: Tachyderes original designation; Piperius Hopkins 1915a: Blackman 1943a: 35, Type-species: Tachy- 8, 39, Type-species: Piperius pini Hopkins deres floridensis Blackman =Cryptocarenus =Hypothenemus sylvicola Perkins, original seriattis Eggers, original designation]. Distri- designation; Ernocryphaliis Murayama 1958: bution: 5 species in North and Central Amer- 934, Type-species: Ernocryphalus hirosimen- ica, more than 7 in South America, 1 intro- sis Murayama, original designation; Acry- duced into tropical Africa. All are con- phalus Tsai & Li 1963: 604, 622, Type-spe- sanguineously polygynous and primarily cies: Cryphalus lipingensis Tsai & Li, myelophagous. Key: Wood (1982: 912). designated by Wood 1984: 224; Jugo- Hypothenemus [1836: 34, Type- cryphalusTsai & Li 1963: 602, 622, Type-spe- Westwood species: Hypothenemus eruditus Westwood, cies: Cryphalus piceus Eggers, designated by monobasic. Synonyms: Stephanoderes Eich- Wood 1984: 224]. Distribution: 3 species in hoff 1872: 132, Type-species: Stephanoderes North America, 1 dubious record from South chapuisii Eichhoff =Crypturgus dissimilis America, 11 in Africa, about 7 in Europe, Zimmermann, subsequent designation by more than 200 nominate species have been Hopkins 1914: 130; Homoeocryphalus Linde- reported from Asia to Australia and adjacent mann 1876: 168, Type-species: Stephan- islands. All are monogamous and phloeo- oderes ehlersii Eichhofi = Hypothenemus eru- phagous. Keys: Reitter (1913: 66) for Europe, ditus Westwood, monobasic; Adiaeretus Balachowskv (1949: 206) for France, Stark Hagedorn 1909: 744, Type-species; Adiaere- (1952: 254) for USSR, Wood (1954b: 1002, tus spinosus Hagedorn = Stephanoderes ela- 1982: 867) for North America and (1960a: 23) phus Eichhoff, monobasic; Stylotentus Schedl for Micronesia. 1939b: 380, Type-species: Hypothenemus Margadillius Hopkins [1915a: 8, 37, Type- concolor Hagedorn, subsequent designation species: Margadillius margadilaonis Hop- by Schedl 1961: 4, 48; probable synonym, kins, original designation]. Distribution: Triarmocerus Eichhoff 1878a: 42, 119, Type- About 9 species in SE Asia, Philippines, New species: Triarmocerus cryphaloides Eichhoff, Guinea, and Indonesia. The true status and monobasic, type lost; Chondronoderes extent of this genus have not been estab- Schedl 1940b: 589, Type-species: Stephan- lished. Key: Hopkins (1915a: 37). 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: original designation; Epsips Beeson 1941:

Archeophalus nataleiisis Schedl, monobasic; 287, nomen nudum. Type-species; Epsips stjl- Pachijnodcres Schedl, 1941: 393, Type-spe- varum, nomen nudum, no status]. Distribu- cies: Pachynoderes deprecator Schedl, tion: 39 species in North and Central America; monobasic; Lepiceroides Schedl 1957a; 59, several hundred nominate species from other Type-species; Lepiceroides aterrinius Schedl, tropical and subtropical areas have been as- monobasic; Ernophloeus Nunberg 1958; 484, signed to this genus, but a majority of them Type-species; Ernophloeus costalimai Nun- are now in synonymy. All are consan- berg = Stephanoderes sundaensis Eggers, guineously polygynous and myelophagous, .

1986 WOOD: Genera of Scolytidae 93 phloeophagous, or spermophagous. Kevs: Parental galleries may be monoramous, bi- Wood (1954: 1017, 1052, 1982; 876) for North ramous, or variously multiramous. The eggs and Central America. are deposited in niches or, in , in Trischidias Hopkins [1915a: 7, 12, Type- fully formed larval cradles. The larvae form species: Trischidias georgiae Hopkins, origi- individual mines or cradles depending on the nal designation]. Distribution: 4 species in SE food habit. The domicile parasitic habit is North America to Me.xico, 1 imported to known in Corthylocurus, Tricolus, amd Am- Hawaii. All are consanguineously polygynous phicrantis and phloeophagous. Kevs: Wood (1954b: Taxonomy. —Except for two small endemic 1067, 1982: 872). genera in Madagascar {Pityodendron, Periocryphalus Wood [1971: 33, Type-spe- Sauroptilius) and one in Africa (Mhniocurus), cies: Periocnjphahis pulhis Wood, original a few primitive Pityophthoriis species in designation]. Distribution: 2 species in South Africa, and several modern Pityophthoriis in America. Both are consanguineously polygy- Eurasia, obviously derived from North Amer- nous and myelophagous in minute lianas. ica, this large, unique tribe is entirely Ameri- can. The occurrence of primitive elements of Tribe Corthylini ancient origin in Africa and Madagascar sug-

Corthyli LeConte [1876: 346, 347, Type-genus: Cort/iy- gests that basic tribal characters formed prior lus Erichson, 1836] to or early in the Tertiary. The elytral locking Pityophthoridae Eichhoff [1878a; 173, Type-genus; mechanism and antennal club are unique in Pitijophthorus Eichhoff, 1864] the family, with the Cryphalini possibly form- Araptidae Eichhoff [1878a: 30.5, Tvpe-genus; Araptus Eichhoff, 1872] ing an intermediate step between Corthylini Amphicranidae Eichhoff] 1878a: 460, Type-genus; Amp/r- and the primitive tribes of Scolytinae. icranus Erichson, 1836] On a biological basis, the tribe is readily Description. —Frons usually dimorphic, divisible into the phloeophagous Pityoph- either or both sexes feebly to strongly modi- thorina and the xylomycetophagous Cor- fied in sculpture and ornamentation; eye thylina, although anatomical characters to emarginate; scape usually elongate, strongly support that division are less definite. The flattened in some Coi'thijlus (Fig. 54), etc., tribe obviously originated in South America funicle 1- to 5-segmented, club strongly flat- and was affected by two radiations. The first tened, sutures present or obsolete, frequently occurred prior to the Tertiary and carried a

1 and/or 2 septate; when present, sutures on few species into Madagascar and Africa, while posterior face little if any displaced toward those land masses were either connected or apex; anterior slope of pronotum asperate close enough for island hopping, and took sev- (one exception in female Corthylus cecropii), eral species over the land bridge to North anterior margin frequently armed, lateral and America, where a minor secondary radiation basal margins fi-equently marked by a fine, occurred. The second major radiation oc- raised line; procoxae contiguous, tibiae rather curred in South America during the Tertiary slender, their lateral margins rarely armed by and produced the Corthylina. more than four socketed teeth; metepisternal might have reached Central America or spine obsolete, replaced by small, transverse southern Mexico over the pre-Tertiary land groove (Fig. 32), thus making it possible for bridge just prior to or during the early stages elytra in locked position to cover at least pos- of its closure. A few genera and species have terior two-thirds of metepisternum (Figs. 53- reached North America over the present post- 54); vestiture usually hairlike, scales rarely Tertiary land bridge or by island hopping in present. recent time. The occurrence oi Gnatharus, a Biology. —Monogamy and heterosan- member of the Corthylina, in China is more guineous polygyny are common throughout difficult to explain. the tribe; a few species of Araptus practice The South American element of this tribe is consanguineous polygyny. Phloeophagy pre- one of the more poorly known segments of the dominates in temporate areas, xylomyce- Scolytidae. Much remains to be learned about tophagy in tropical areas, although myelo- this remarkable group. Chiloxylon Schedl phagy and spermophagy are common. (see Dryocoetini) could belong to this tribe. 94 Great Basin Naturalist Memoirs No. 10

Kev to the Genera of Corthvlini ginate to accommodate a pair of mandibular (Modified from Wood 1982) - spines; phloeophagous; Costa Rica to Bolivia and Brazil; Asfroniiim, Spotidias; 1 3-1. 6mm 1. Phloeophagous, myelophagous, or sper- Styphlosonuj mophagous; antennal funicle 5-segmented — Sutures of antennal club straight or recurved; (except 3- or 4-segineiited in Dcndroterus, female epistoma entire, male frons without a Daciwpbthorus , and some Pitijophthorus transverse carina; mandibular spines never costatiis), club usually smaller, symmetrical; present; elytral declivity convex, interstriae prosternal intercoxal piece acutely pointed unarmed by tubercles or spines; female frons (except obsolete in Dacnophthorus); pubes- shallowly concave, bearing a brush of hair, cence usually more abundant, usually in rows armed on median fourth above eyes by an on elytra; elytral declivity mostly convex to acute, transverse carina and by a pair ofcoarse bisulcate, armature conser\'ative (subtribe tubercles in lateral areas at level of antennal Pityophthorina) 2 insertion; male frons convex, simple; Mexico; — Xylomycetophagous; antennal funicle 1- to 5- Bursera; 1.6-1.8 mm Phloeotertts segmented, club usually much larger, com- 6(2). Sutures of antennal club moderately to very monly asymmetrical; prosternal intercoxal strongly procurved, onh- suture 1 septate, or piece absent (except obtuse in Gnatho- if all external sutures obsolete then mesal half trichus, Gnathotrupes); pubescence usually of suture 1 marked internally by a strongly greatly reduced to obsolete or minute and procurved septum at least on 1 side; strongly confused (not in rows); elytral decliv- phloeophagous, myelophagous, spermopha- ity convex to truncate to deeply excavated gous; S USA to Argentina; 1.1-3.3 mm (weakly bisulcate in some Gnathotrichus), Araptus commonly with spinelike processes (subtribe Corthylina) 17 — Sutures 1 and 2 clearly, equally marked by rows of setae and grooves, straight to moder- 2(1). Basal and lateral margins of pronotum ately procurved, if procurved then both su- rounded, devoid of a fine, raised line; elytra tures at least partly septate (sutures always rather coarsely punctured (American genera), straight when both almost obsolete) 7 or veiy finely punctured (African genus), un- armed declivity steep, in American genera 7(6). Lateral margins ol pronotum rounded, with- usually subvertical and somewhat flattened out a fine, raised line 8 on lower half, almost never bisulcate; discal — Lateral margins of pronotum subacute, vestiture abundant 3 marked by a fine, raised line (rather obscure — Basal and usually lateral margins of pronotum in Spennophthorus and some Pitijophthorus marked by a finely raised line; elytral de- having pronotal asperieies in subconcentric clivilty usually more gradual, convex to bisul- rows 12

cate, often ornamented by granules or small 8(7). Sutures 1 and 2 of antennal club aseptate and denticles 6 clearly marked by grooves and rows of setae;

3(2). Antennal club pubescent to base, unmarked pronotal asperities continuing in lateral areas by sutures, funicle .5-segmented; female frons to base; larger species; spermophagous in ornamented by a brush of long hair, male cones of Piniis. at least 2 species sometimes usually with a median tubercle or longitudinal myelophagous; North America; 2.2-4.1 mm carina at upper level of eyes; elytral declivity conservatively sculptured, usually convex; — Sutures 1 and 2 both partly to completely

Africa to India and China; 1.4-2.8 mm septate or, if aseptate, then antennal club Mimiocurus largely glabrous or body size much smaller; — Antennal club with sutures clearly indicated; pronotal asperities not extending to basal American genera 4 margin; smaller species, not found in cones of conifers 9 4(.3). Antennal funicle 3- or 4-segmented, club with rather large, at least 2.5 sutures at least slightly recurved, aseptate; 9(8). Antennal club times .... 10 elytral vestiture hairlike; female epistoma not as long as funicle; monogamous species emarginate, mandible never with projecting — Antennal club comparatively small, less than spines; strong, transverse frontal carina at up- 1.5 times as long as funicle; polygynous spe- per level of eyes, when present, a female cies 11 character; phloeophagous; SW USA to 10(9). Body stouter, 2.5-2.8 times as long as wide; Panama; Bursera, 1 species in Jatropha; 1.3- elytral declivity convex; anterolateral areas of 2.9 Dendroterus mm female pronotum with a pair of large, oval, — Antennal funicle .5-segmented; secondary densely pilose areas; antennal club with only 2 sexual characters different 5 sutures; phloeophagous, larval mines resem- ble cradles (of ambrosia beetles); S USA to 5(4). Antennal club with aseptate sutures strongly Honduras; Pinus; 1.5-3.2 procurved; interstrial setae scalelike; male mm frons strongly, transversely carinate at upper — Body very slender, 3.7-3.8 times as long as level of eyes, female epistoma deeply emar- wide; elytral declivity strongly impressed; fe- , .

1986 WOOD: Genera of Scolvtidae 95

male pronotum without special pilose areas; Central and South America; Cacsalpinus antennal club with suture 3 indicated by a row etc.; 1.3-1.9 mm Spermophthorus of setae; phloeophagous, but with fungal 16(13). Sutures of antennal club moderately pro- mycelium in parental gallery; Mexico to curved, segment 1 shorter than 2 or 3; greater South America; Clematis and Bignoniaceae frontal pubescence a male character, clytral vines; 1.2-2.0 mm Dacnophthorus punctures very fine, usually confused, short 11(9). Female pregula very greatly enlarged and pubescence abundant, often .scalelike; striae bearing a rather dense, conspicuous tuft of usually obsolete; phloeophagous, monoga- very long hair, male pregula only slightly en- mous; North America to Colombia, China;

larged, usually without tuft of hair; phloeo- Quercus, rarely other hosts; 1.2-2.6 mm . . . phagous; USA (Arizona, New Mexico), Pintix, 1.3-1.8 mm Pityotrichus — Sutures of antennal club straight to modestly — Female oral region abnormally broad and, procurved, segments 1 and 2 subequal in usually, with mandibles greatly enlarged; length; pubescence usually much less abun- pregula normal, phloeophagous. Central and dant, never scalelike, strial punctures in rows

South America; Dacreoidcs. Cedrcla. Prot- or, if confused, then rather coarse; phloeo- ium; 1.3-1.8 mm Gnatholeptus phagous, monogamous or heterosanguine- America, Europe, Asia, 12(7). Pronotum weakly declivous on anterior ously polygynous; 0.9-3.9 fourth, asperities small, numerous, gradually Africa; coniferous and broadleal hosts; Pityophthorus decreasing in size toward base; head unusu- mm

ally wide, mandibles large and stout in both 17(1). Antennal funicle 5-segmented, club always sexes; eye large, coarsely faceted, one-third symmetrical, with 2 or 3 clearly marked su-

divided by a broad emargination; Madagas- tures; protibia widest near its apex, its poste-

car; 3.2-3.4 mm Vitxjodendron rior face usually flat, unarmed (a few minute — Pronotum more strongly declivous in front, granules in some Gnatbotrupes), elytral de- asperities usually not present behind summit; clivity conservatively sculptured (except in head and mandibles normal 13 some Gnathotnipes) 18

13(12). Pronotum without a transverse impression — Antennal funicle 1- to 4-segmented, club behind summit, transition between asperate commonly asymmetrical, often greatly en- and smooth areas more gradual, asperities al- larged, sutures often reduced or absent; tibiae ways confused; interstrial bristles usually variously sculptured 19

stout to scalelilce (1 exception, also, almost 18(17). Sutures of antennal club straight to moder- glabrous in Sauropfi/ius); tropical species .. 14 ately procurved, segment 1 not noticeably — Pronotum almost always with a distinct, trans- reduced in size; elytral declivity convex to verse impression behind summit, if doubtful narrowly, rather shallowly sulcate, subapical then asperities almost always arranged in con- margin near apex acutely elevated, sutnral

centric rows, interstrial setae hairlike (il stout apex entire, rather narrowly rounded behind; then pronotal asperities concentric) 16 monogamous; North and Central America; coniferous and broadleaf hosts; 2.0-4.3 mm 14(1.3). Anterior margin of pronotum unarmed; de- Gnathotrichus clivity very broadly excavated (as wide as body), its lateral crests profoundly elevated — Sutures of antennal club moderately to

and serrate, but abruptly ending before su- strongly procurved, segment 1 distinctly

tural apex; Madagascar; 3.2 mm . . . Sauroptilius smaller; elytral declivity moderately to — Anterior margin of pronotum armed by serra- strongly flattened; elytral apex at least weakly shallowly tions or a continuous costa; declivity convex to divaricate, very broadly romided to moderately sulcate, impression rarely equal emarginate behind, without a submarginal to more than half width of body; neotropical costa near apex; monogamous; Central and 15 South America; broadleaf hosts; 1..3-4.0 mm Gnathotrupes 15(14). Strial punctures rather coarse, mostly in rows, declivity moderately to rather strongly 19(17). Antennal funicle 2- to 4-segmented, club with impressed, lateral margins armed or not; 2 sutures clearly marked; elytral apex divari- frons never armed, epistomal margin with a cate (except Mctacorthylus. Glochinoccrus) small, median, premandibular lobe, polygy- commonly explanate, declivity often elabo- nous; phloeophagous, axes of tabular nuptial rately excavated and armed by spines; pro- chamber perjjendicular to cambium, longest tibia always slender, with posterior face in- axis parallel to grain of wood; Colombia, flated and tuberculate; body usually slender Venezuela; broadleaf trees, 1.3-3.1 mm .... 20 Phelloterus — Antennal funicle 1-segmented, club with 1, 2, — Strial punctures either very small or con- or no sutures; elytral apex entire (except fused; declivity not as steep, convex to shal- Brachyspartus, Corthylua cmarfiinatus), de- lowly impressed, lateral margins never clivity convex to rather weakly excavated, armed; male frons armed; spermophagous; never explanate; posterior margin ol protho- 96 Great Basin Naturalist Memoirs No. 10

racic precoxal piece transversely straight, not — Antennal funicle 2- or 3-segmented; prono- extended between coxae; protibia variable; tum and elytra cjuite different; American gen- body conparatively stout 24 era 23

20(19). Procoxae contiguous, anterior wall of com- 23(22). Elytral apex divaricate, often also explanate; bined cavities and precoxal piece transversely antennal club oval to broadly triangular; lat- straight (longitudinally very thin); protibia eral margins of pronotum usually with a fine, similar in male and female, with a longitudinal raised line (a few exceptions); body moder- marginal row of tubercles, posterior face ei- ately to very slender; mostly polygynous; ther unarmed or with a longitudinal row of up North and South America; 1.4-4.8 mm to about 4 tubercles, posterior face of female antennal club with long hair sparse to absent; — Elytral apex entire, never explanate; lateral funicle usually 3-segmented, less antennal margins of pronotum rounded; antennal club commonly 2-segmented; frons commonly more than twice as long as wide (except 1.6 with a sharply defined granular area 21 times in male Metacorthijlus), its apex nar- — Prothoracic precoxal piece moderately large, rowly rounded; body comparatively stout ... 24 posteriorly angulate, occupying anterior por- 24(23). Antennal funicle 3-segmented; female frons tion of area between coxae (except in excavated and elaborately ornamented by Gnatharus, from Tibet, a thin, transverse long hair; antennal club not sexually dimor- partition); male protibia armed by coarse mar- phic, elongate, slightly asymmetrical; prono- ginal serrations and a row of equally coarse tum and elytral disc glabrous; monogamous; S serrations on posterior fiice, female protibia Mexico to Guatemala; 3.0-3.8 mm with posterior face moderately inflated and Glochinocerus armed by numerous, confused, small tuber- — Antennal funicle 2-segmented; frons convex cles in addition to coarser marginal row; pos- and subglabrous in both sexes; antennal club terior face of female antennal club orna- asymmetrically very elongate in female, elon- mented by more abundant, long hair; frons gate-oval in male; pronotum and elytra never ornamented by a sharply defined gran- minutely, closely pubescent; monogamous; ulate area 22 Costa Rica to Colombia; 1.9-2.7 mm 21(20). Elytra broadly rounded behind, posterior Metacorthylus margin of declivity feebly if at all explanate, 25(19). Lateral margins of pronotum rounded; elytral weakly if at all divaricate; lateral margins of disc usually impunctate, declivity short, very declivity armed by as many as 3 pairs of steep, narrowly sulcate on basal third, trian- spines; antennal club oval to subtriangular, gularly impressed below, costal margins near little if any longer than wide; anteroventral apex ascending slightly; antennal club subcir- margin of prosternum flanged, bent or folded cular, syummetrical, with 2 aseptate sutures caudad away from head, usually bearing a tuft marked by rows of setae; female frons convex, of hair; pronotum usually stouter, anterior pubescence inconspicuous; monogamous; margin usually serrate; antennal funicle 3- Mexico to South America; 1.2-2.4 mm segmented; monogamous; Mexico to South Microcorthylus America; 1.5-3.9 mm Tricolus — Lateral margins of pronotum marked by a — Posterior margin of elytral declivity strongly fine, raised line (except some Corthycyclon), to profoundly explanate, weakly to pro- elytral disc usually with clearly marked, con- foundly divaricate; declivital armature vari- fused punctures, declivity convex, truncate, able, often remarkable; antennal club oval to or variously sculptured (but not as above); very elongate; anteroventral margin of female frons usually moderately to strongly prosternum fitting snugly against head (not concave, often ornamented by hair; antennal bent caudad), sparsely pubescent; pronotum club symmetrical to strongly asymmetrical, usually much more elongate, anterior margin sutures present or not 26 variously sculptured, rarely serrate; antennal funicle usually 3-segmented, occasionally 2- 26(25). Antennal club aseptate, without sutures segmented; monogamous; Mexico to .South (some species with weak, transverse grooves, America; 2. .5-8.0 mm Amphicramis without rows of setae), usually very elongate; lateral margins of pronotum either with or 22(20). Antennal funicle 4-segmented; frons with me- without a fine, raised line; posterior face of dian carina; pronotum dimorphic, female protibia inflated and tuberculate; female frons with anterior slope strongly declivous, armed broadly, evenly concave and ornamented by by numerous asperities, anterolateral angles fine hair; monogamous; Mexico to South each bearing a tuft of hair, male slope gradual, America; 1.3-2.4 mm Corthycyclon asperities greatly reduced in size and num- ber, anterior margin a strongly formed, — Antennal club with 1 or 2 sutures, its outline slightly produced continuous costa; elytral subcircular to strongly asymmetrical (if su- declivity divaricate from middle, truncated tures absent then posterior face of protibia before apex; Asia (Tibet); 2.0 mm .. Gnatharus flat, smooth); female frons variable 27 .

1986 WOOD: Genera of Scolytidae 97

27(26). Elytral apex strongly, obtusely divaricate, fe- designation]. Distribution: 15 species in male antennal club with 1 septate suture, ta- North and Central America. All are het- pered on apical half to strongly acuminate polygynous and phloeo- apex, posterior face of club without a tuft of erosanguineously long hair; female protibia inflated, armed on phagous. Key: Wood (1982: 918). posterior face; Venezuela; 2.5 mm Phloeoterus Wood [1984: 117, Type-spe- Brachyspartus cies: Phloeoterus burserae Wood, original — Elytral apex entire (a small cleft in Corthijlus designation]. Distribution: 1 species in Burs- emarginatus). female antennal club not ta- era in Mexico. pered or acuminate at apex 27 Arapfus Eichhoff [1872: 136, Type-species: 28(27). Elytral declivity narrowly, weakly sulcate (ex- cept uniformly convex in 1 species), lateral Araptus nifopalliatus Eichhoff, monobasic. margins armed by 2 or 3 pairs of pointed Synonyms: Neodi-yocoetes Eggers 1933a: 9, club granules; antennal scape elongate, Type-species: Neodryocoetes hymenaeae Eg- shaped; female frons variously impressed, Schedl with a pair of median carinae narrowly sepa- gers, monobasic; Thamnophthorus rated by a sulcus over part or all of median 1938c: 174, Type-species: Thamnophthorus line; color pale yellow to yellowish brown; volastos Schedl, subsequent designation by antennal club symmetical, broadly oval, with Blackinan 1942b: 178; Neopityophthorus 2 finely marked sutures; female protibia in- Schedl 1938c: Type-species: Pityophtho- flated, posterior face tuberculate; monoga- 180, mous; Mexico to South America; 1.5-2.8 mm rus laevigatus Eggers, designated by Wood Corthylocurus 1982: 928; Sphenoceros Schedl 1939e: 565, — Sculpture of elytral declivity convex, trun- Type-species: Sphenoceros Umax Schedl, cately concave, or variously impressed (but monobasic; Hypertensus Hagedorn, nomen never narrowly sulcate); antennal scape sub- nudum (in Schedl 1950d: 164), Type-species; quadrate, stout; female frons never with pair of median carinae; antennal club slightly to Hypertensus reittcri, nomen nudum = Sphe- profoundly asymmetrical, sutures (when noceros Umax Schedl, no status; Brachyden- present) rather strongly marked; posterior dridus Schedl 1951b: 114, Type-species: face of protibia smooth or tuberculate; Brachydendrulus eggersi Schedl, monobasic; monogamous; North and South America; 1.2- 4.3 mm Corthijlus Gnathocranus Schedl 1951b: 116, Type-spe- cies: Gnathocranus novateutonicus Schedl, Pityophthorina Corthylini; monobasic; Gnathohorus Schedl 1970a: 93, Mimiocurus Schedl [1957a: 72, Type-spe- Type-species: Breviophthorus argentiniae cies: Mimiocurus acuminatus Schedl, mono- Schedl, original designation]. Distribution: basic. Synonyms: Micracidendron Schedl .53 species in North and Central America, 1957a: 71, Type-species: Micracidendron about 50 in South America and adjacent is- montanum Schedl, monobasic; Mimiophtho- lands. Some are monogamous, some are het- rus Schedl 1957a: 77, Type-species: Bra- erosanguineously polygynous, and at least 3 chydendrulits montanu.s Schedl, original des- are consanguineously polygynous {laevigatus, ignation]. Distribution: About 8 species in costaricensis , etc.); most are phloeophagous, Africa, 1 in India, 1 in China. a few are spermophagous, and some are Styphlosoma Blandford [1904: 2.32, Type- myelophagous in vines (lianas). Key: Wood species: Styphlo.'ioina uranulatum Blandford, 1982: 928) for North and Central America. monobasic]. Distribution: 1 species in Central Conophthorus Hopkins [1915c: 430, Type- America (Costa Rica to Panama), 3 in South species: Pityophthorus coniperda Schwarz, America. They are heterosanguineously poly- original designation]. Distribution: 14 species gynous and phloeophagous. in North America. All species are monoga- Dendroterus Blandford [1904: 233, Type- mous and spermophagous in cones oi Pinus species: Dendroterus mexicanus Blandford, Key: Wood (1982: 982). subsequent designation by Hopkins 1914: Pityoborus Blackman [1922: 96, Type-spe- 120. Synonyms: Plesiophthorus Schedl 1940a: cies: Crypturgus comatus Zimmermann, 343, Type-species: Plesiophthorus perspectus monobasic]. Distribution: 7 species in North Schedl, monobasic; Xylochilus Schedl 1956: and Central America. All are monogamous 30, Type-species: Xylochilus insularis Schedl and phloeophagous in Pinus. Keys: Wood =Dendroterus sallaei Blandford, original (19.58: 47, 1982: 1147). 98 Great Basin Naturalist Memoirs No. 10

Dacnophthoriis Wood [1975; 394, Type- Pittjophthorus Eichhoff [1864: 39, Type- species: Gnathophthorus clematis Wood, species: Bostrichus lichtcnsteini Ratzeburg, original designation]. Distribution: 2 species subsequent designation by Hopkins 1914: in North and Central America, 2 in South 127. Synonyms: Trigonogenius Hagedorn America. All are monogamous and bore in the 1912: .354, Type-species: Trigonogenius fallax stems of vines (lianas) (Clematis and a species Hagedorn, monobasic; Hagedornus Lucas of Bignoniaceae). Key: Wood (1982: 1152) for 1920: 683, Type-species: Trigonogenius fallax North and Central America. Hagedorn, automatic; Mijeloborus Blackman Pityotrichus Wood [1962: 76, Type-spe- 1928a: 16, Type-species: Pitijophthorus cies: Pitijophilus harbatus Blackman, auto- ramiperda Swaine, original designation; matic. Synonym: Pitijophilus Blackman Gnathophorus Schedl 1935: 342, Type-spe- 1928a: 147, Type-species: Pitijophilus harba- cies: Gnathophorus sparsepilosus Schedl, tus Blackman, original designation, preoccu- monobasic, preoccupied; Conophthocran- pied]. Distribution: 2 species in North Amer- idus Schedl 1935; .343, Type-species: Conoph- ica (Arizona, New Mexico). Both are hetero- thocranulus blackmani Schedl, monobasic; sauguineously polygvnous and phloeo- Breviophthorus Schedl 1938c: 176, Type-spe- phagous. Key: Wood (1982: 1141). cies: Breviophfhorus brasiliensis Schedl, Gnatholeptus Blackman [1943a: 34, Type- monobasic; Pityophthoroides Blackman species: Gmitholcptus mandihularis Black- 1942b: 199, Type-species: Pityophthoroides man = Pitiiophthorus shannoni Blackman, pudens Blackman, original designation; original designation]. Distribution: 4 species Cladohorus Sawamoto 1942: 165, Type- in Central and N South America. All are het- species: Cladoboriis arakii Sawamoto, erosanguineously polygynous and phloeo- monobasic; Neomips Schedl 1954a: 37, Type- phagous, mostly in Protium. The status of this species: Neomips brasiliensis Schedl genus should be reviewed. Key: Wood (1982: = Pityophthonis dimorphus Schedl, monoba- 1142). sic; Ctenyophthorus Schedl 1955b: 26, Type- Pityodendron Schedl [19.53a: 93, Type-spe- species: Ctenyophthorus glabratulus Schedl, cies: Pityodendron madagascarensis Schedl, mono-basic; Gnathophthorus Wood 1962: 76, monobasic]. Distribution: 1 species in Mada- Type-species: Gnathophorus sparsepilosus gascar. Schedl, automatic; Hypopityophthorus Sauroptilius Browne [1970: 558, Type-spe- Bright 1981a: 14, Type-species: Pitijophtho- rus inops original designation]. Distri- cies: Xyleborus sauropterus Schedl, original Wood, bution: 225 species in North and Central designation]. Distribution: 1 species in Mada- gascar. America, about 100 in South America and adjacent islands, about 15 in Asia, 22 in Eu- Phelloterus Wood [1971: 46, Type-species: rope, 22 in Africa. Most are heterosan- Phelloterits tersus Wood, original designa- guineously polygynous, a few are monoga- tion]. Distribution: 3 species in South Amer- mous; most are phloeophagous, a few are ica (Colombia to Venezuela). All are het- mvelophagous. Kevs: Pfeffer .334) for erosanguineously polygynous' and phloeo- (1976: Europe, Blackman' (1928a), Bright phagous. (1981a), and Wood (1982: 991) for North and Central Spermophthorus Costa Lima [1929: 111, America. Type-species: Spermophthorus apuleiae Costa

Lima, monobasic]. Distribution: 1 species in Corthylini: Corthylina Central America (Costa Rica), 2 in South America. Gnathotrichus Eichhoff [1869: 275, Type- Pseudopityophthorus Swaine [1918: 93, species: Gnathotrichus corthyloides Eichhoff Type-species: Crypturgus minutissimus Zim- = Tomiciis materiarius Fitch, monobasic. mermann, original designation]. Distribu- Synonyms: Gnathotrichoides Blackman tion: 23 species in North and Central Amer- 1931b: 267, Type-species: Cryphalus sulcatus ica, 1 in South America (Colombia), 1 in LeConte, subsequent designation by Wood China. All are bigynous and phloeophagous, 1982: 1 1.55; Anci/Zof/ere.s Blackman 1938b:205, mostly in Quercus. Keys: Blackman (1931a: Type-species: Cryphalus pilosus LeConte, 225), Wood (1982: 966). original designation; Paraxyleborus Hoff- 1986 WOOD; Genera of Scolytidae 99 inann 1942: 72, Type-species; Xyleborus Gnatharus Wood & Yin [1986; 463, Type- duprezi Hoffmann =Tomicits mutcriarhis species; Gnathariis tibctensis Wood & Yin, Fitch, monobasic; Prognathotricliiis Bright original designation]. Distribution: 1 species 1972: 1678, Type-species: Prognathotrichiis in Asia (Tibet). priimis Bright, original designation]. Distri- Monarthruin Kirsch [1866: 213, Type-spe- bution; 13 species in North and Central Amer- cies: Monarthrum chapiiisi Kirsch, monoba- ica. All are monogamous and .xylomyce- sic. Synonyms; Coi-thylomimus Ferrari 1867: tophagous. Kevs: Blackman (1931b: 266), 48, Type-species; Bostrichus fasciatus Say, Wood (1982; 1156). subsecjuent designation by Hopkins 1914: Gnathotrupes Schedl [1951b: 125, Type- 118; Cosmocoryniis Ferrari 1867: 62, Type- species: Gnathotnipes bolivianiis Schedl, species; Cosmocorynus cristatus Ferrari, monobasic. Synonyms: Gnathotnjpaiuis Wood monobasic; Pterocyclon Eichhoff 1869; 276, 1968a: 9, Type-species; Gnathotrypanus terc- Type-species; Pterocyclon late rale Eichhoff, hrattis Wood, original designation; Gnatho- subsequent designation by Hopkins 1914; cortiis Schedl 1975a; 11, Type-species: 128, neotype for type-species designated by Gnathocorius caliculus Schedl, original des- Wood 1966; 25; Anchonocerus Eichhoff ignation; Gnathomhnus Schedl 1975a; 12, 1878a; 67, 431, Type-species; Anchonocerus Type-species; Giuithomimiis nothofagi ntfipes Eichhoff, monobasic; Phthohus Eich- Schedl, original designation; Gnathoglochi- hoff 1878a; 67, 433, Type-species; Phthorius nus Schedl 1975a: 16, Type-species: Gnatho- ingens Eichhoff, monobasic; Trypocranus glochinus impressus Schedl, original designa- Eichhoff 1878a: 67, 4.35, Type-species; tion]. Distribution; 5 species in Central Trypocranus cincinnatus Eichhoff, monoba- America, about 30 in South America. All are sic; Eupteroxylon Eggers 1936a: 392, Type- monogamous and xylomycetophagous. Key: species: Type-species; Eupteroxylon coma- Wood(1982; 1167) for Central America. tum Eggers, monobasic]. Distribution: 58 Tricolus Blandford [1905; 286, Type-spe- species in North and Central America, about cies; Tricoltis ovicollis Blandford, subsequent 78 in South America and adjacent islands. designation by Hopkins 1914; 131. Synonym: Most species are heterosanguineously polygy- Pterocyclonoides Schedl 1970a: 101, Type- nous, a few small species are monogamous; all species: Pterocyclonoidcs octodentatits are xylomycetophagous. Key: Wood (1982: Schedl, monobasic]. Distribution: 24 species 1207) for North and Central America. in North and Central America, about 17 in Glochinoceriis Blandford [1904: 266, Type- South America. All are monogamous and xy- species: Glochinoccrus retusipennis Bland- lomycetophagous. Key: Wood (1982: 1170) for ford, subsequent designation by Hopkins North and Central America. 1914; 122]. Distribution: 2 species in Mexico Amphicranus Erichson [1836; 63, Type- (Hidalgo) and Central America (Guatemala). species; Arnphicranus thoracicus Erichson, Both are monogamous and xylomyce- monobasic. Synonyms: Piezorhopahis Guerin- tophagous. Key; Wood (1982; 1246). Meneville 1838: 107, Type-species; Piezorho- Metacorthijlus Blandford [1904: 251, 263, palus nitidulus Guerin-Meneville =Amphi- Type-species: Metacorthyhis nigripennis craniis thoracicus Erichson, monobasic; Blandford, monobasic. Synonym: Paracor- Steganocranits Eichhoff 1878a; 460, Type- thyhis Wood 196Sa; 7, Type-species; Paracor- species; Steganocranus dohnii Eichhoff, thyhis velutinus Wood, original designation]. monobasic]. Distribution: 29 species in North Distribution; 4 species in Central America 1 of and Central America, about 23 in South which also occurs in South America (Colom- America. All are monogamous and xylomyce- bia). All are apparently monogamous and xy- tophagous; apparently some appropriate the lomycetophagous. Key: Wood (1982: 1248). tunnel of another species of Scolytidae or Microcorthylus Ferrari [1867; 58, Type- Platypodidae by driving out or killing the orig- species; Microcoiihylus parvulus Ferrari, inal occupants (domicile parasitism). Key: monobasic]. Distribution: 13 species in Mex- Wood (1982; 1186) for North and Central ico and Central America, 19 in South Amer- America. ica. All are monogamous and xylomyce- 100 Great Basin Naturalist Memoirs No. 10 tophagous. Key: Wood (1982; 1252) for North Bebger. B. M. 1916. Bark beetles of southern Ussuri and Central America. region (In Russian). Rus.skoe Entomologicheskoe Obozrenie 16: 226-248. Corthycyclon Schedl [1951b: 128, Type- Bergroth, E. E 1884. Bemerkungen zurdritten auflage species; Corthycyclon ustum Schedl, mono- des Catalogus Coleopterorum Eiiropae auctoribus V. basic]. Distribution: 6 species in Mexico and L. Heyden, E. Reitter, et J. Weise. Berliner Entomologische Zeitschrift 28(2): 225, 230. Central America, about 10 in South America. Blackman, M. W 1920. North American Ipidae of the All are monogamous and xylomycetophagous. family Micracinae, with descriptions of new spe- Key; Wood (1982: 1260) for North and Central cies and genera. Mississippi Agricultural Experi-

America. ment Station, Technical Bulletin 9, 60 p. , 5 pis. Brachyspartus Ferrari [1867: 65, Type- 1922, Mississippi bark beetles. Mississippi Agri- cultural Experiment Station, Technical Bulletin species: Brachyspartus moritzi Ferrari, 11. 130p.,28pls. monobasic. Synonym; Thyhircos Schedl 1928a. The genus Pitijophthorus Eichh. in North 1939e: 567, Type-species; Brachyspartus America. A revisional study of the Pityophthori, moritzi Ferrari, subsequent monotypy (Wood with descriptions of two new genera and seventy- one new species. New York State College of 1982: 1298). Distribution: 1 species in South Forestry at Syracuse LIniversitv. Technical Publi- America (Venezuela). cation 25: 1-159, 11 pis. Corthylocurus Wood [1966: 18, Type-spe- 1928b. Notes on Micracinae, with descriptions of twelve new species. York State College cies; Brachyspartus barbatus Blandford, New of Forestry at Syracuse University, Technical Publi- original designation]. Distribution: 6 species cation 25; 18.5-208. in Mexico and Central America, about 8 in 1931a. A revisional study of the genus Pseudopi- South America. All are monogamous and xy- hjophthorus Swaine in North America. Washing- lomycetophagous. Key; Wood (1982; 1265) for ton Academy of Sciences, Journal 21(10): 223-236, 15 pis. Mexico and Central America. 1931b. A revisional study of the genus Gnatho- Corthylus Erichson [1836; 64, Type-spe- trichus Eichhoff in North America. Washington cies; Bostrichus compressicornis Fabricius, Academy of Sciences, Journal 21: 264-276. subsequent monotypy (Ferrari 1867; 49), lec- 1934. A rivisional study of the genus Scolytus Ge- offroy (Eccoptogaster Herbst) in North America. totype designated for type-species by Wood v. S. Department of Agriculture, Technical Bul- 1974; 202. Synonyms; Morizus Ferrari, 1867; letin 431. 30 p. 69, Type-species: Morizus excisus Ferrari, 1938a. New species of Cacfopinus Schwarz(Cole- monobasic; Pseudocorthyhis Ferrari 1867: 59, optera: Scolytidae). Entomological Society of Washington, Proceedings 40(6): 151-157. Type-species: Pseudocorthyhis letzneri Fer- 1938b. Ancyloderes , a new genus of Scolytidae. rari, subsequent designation by Hopkins Entomological Society of Washington, Proceed- 1914; 128]. Distribution: 56 species in North ings 40: 204-206. and Central America, about 54 species in 1938c. The genus Chramesus LeConte in North America (Coleoptera: Scolytidae). Washington South America. All are monogamous and xy- Academy of Sciences, Journal 28: 5.34-545. lomycetophagous. Key: (1982: 1271) for Wood 1939. A new genus and three new species of North and Central America. Scolytidae from Argentina and Bolivia (Cole- optera). Revista de Entomologia 10(1): 86-96. 1940. The scolytid beetles of the genus Renocis Literature Cited Casey, with descriptions of nine new species. U. S. National Museum, Proceedings 88(3084): 373-401. Arnoldi, et al, 1977. (Title not seen). Trudy Paleont. 1941. Bark beetles of the genus Hijlastes Erichson Inst. 166: 1-203. (Partial English translation re- in North America. U, S. Department of Agricul- ceived through R. T. Thompson). ture, Miscellaneous Publications 417. 27 p. Balachowsky. a S 1947. A propos du genre Pitijonencfi 1942a. Revision of the bark beetles belonging to Bedel (Col. Scolytoidea). Societe Entomologique the genus Pscudohijlcsinus Swaine. U. S. Depart-

de France, Bulletin 52(.3): 44. ment of Agriculture, Miscellaneous Publications

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. 1971. New records and species of neotropical bark 1984. New generic synonymy and new genera of beetles (Scolytidae: Coleoptera), Part V. Brigham Scolytidae (Coleoptera). Great Basin Naturalist Young University Science Bulletin, Biological Se- ' 44(2): 223-230. ries 15(3): 1-.54. Wood, S, L . andF Hu.\nc 1986. A new genus of Scolyti-

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, 1973b. New synonymy in American bark beetles zophorae (Hopkins) (Coleoptera, Scolytidae). (Scolytidae: Coleoptera), Part 111. Great Basin Florida Department of Agriculture, Entomology Naturalist 33(3): 169-188. Circular 98. 2p.