THE CIID BEETLES of CALIFORNIA (Coleoptera: Ciidae)

Home , Ceracis, Ciidae, Ciini, Cis (beetle), Dolichocis, Ennearthron

CALlFURNlA INCECT SURVEY DlVlSlDN OF ENTOMOLCG’1 UNIVERSITY OF CALIFORflIP E~ERKELEY, 9rm

BULLETIN OF THE CALIFORNIA INSECT SURVEY

VOLUME 17

THE CIID BEETLES OF CALIFORNIA (Coleoptera: Ciidae)

BY JOHN F. LAWRENCE

UNIVERSITY OF CALIFORNIA PRESS THE CIID BEETLES OF CALIFORNIA (Coleoptera: Ciidae) BULLETIN OF THE CALIFORNIA INSECT SURVEY

VOLUME 17

THE CIID BEETLES OF CALIFORNIA (Coleoptera: Ciidae)

BY JOHN F. LAWRENCE ( Museum of Comparative Zoology, Harvard University)

UNIVERSITY OF CALIFORNIA PRESS BERKELEY LOS ANGELES LONDON 1974 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Advisory Editors: J. N. Belkin, R. M. Bohart, Paul De Bach, R. L. Doutt, W. H. Lange, E. I. Schlinger, D. D. Jensen

VOLUME 17 Approved for publication December 1,1972 Issued January 22,1974

UNIVERSITY OF CALIFORNIA PRESS BERKELEY AND LOS ANGELES UNIVERSITY OF CALIFORNIA PRESS, LTD. LONDON,ENGLAND

ISBN 0-520-09489-1 LIBRARY OF CONGRESS CATALOG CARD NO.: 72-619714

(i3 1974 BY THE REGENTS OF THE UNIVERSITY OF CALIFORNIA

PRINTED BY OFFSET IN THE UNITED STATES OF AMERICA CONTENTS

Introduction ...... 1 Bionomics ...... 2 Habitat and host preference ...... 2 Distribution ...... 4 Acknowledgments ...... 6

Systematics Genus Cis ...... 9 c . acritus., ...... 10 C.amedcaws ...... 10 C. angustus ...... 11 C. biurmatus ...... 12 C. crebem'mus ...... 12 C.duplex ...... 12 C. ephippiutus ...... 13 C .f uscipes ...... 13 C.hystriculus ...... 14 C.hettei ...... 14 C.marittmw ...... 14 C . megasttctus ...... 15 C.striolatus ...... 15 C.tdracentrum ...... 3.6 C .td.mta$us ...... 16 C. versicolor ...... 18 C. Vitulus ...... 3.7 Genus Dolichosis ...... 18 D.indkrMnctUs ...... 18 D.naanitoba ...... 18 GenusOTthods ...... 0. punctatus ...... 19 Genus Hadraub ...... 19 H . blukddld ...... 20 Genus Plesiods ...... P . cribrum ...... Genus Cera& ...... 20 C. californicus ...... 21 C. dixiensis ...... 22 C.Similis ...... 22 Genus Sulcack ...... 22 S.cUrtulus ...... 22 Genus Octotsmnus ...... 23 0.luevkr ...... 23

Addendum ...... 23 Subfamily Sphindociinae...... 24 Genus Sphfndocls Fall ...... 24 S . dentfcol2is ...... 24

Literature Cited ...... 25

Figures ...... 27 THE CIID BEETLES OF CALIFORNIA

BY JOHN F. LAWRENCE

INTRODUCTION

THE FAMILY C~DAE~includes about 550 described my revision cited above, but the more important char- species of minute Coleoptera, which are distributed acters are also illustrated in figures 1-19. throughout the world and are associated with the The following measurements are used in the keys: PL mycelia and fruiting bodies of wood-rotting fungi. ( pronotal length along midline, but including horns in These beetles are usually less than 3 mm long and they males); PW (greatest pronotal width); EL (greatest vary considerably in body form (elongate and flattened elytral length, just to one side of suture); EW (greatest to cylindrical or globose). Many species have become elytral width); GD (greatest depth, through elytra and adapted for boring into hard substrates and thus re- metasternum); TL (PL and EL combined). Note that semble powder-post beetles (Bostrichidae) and bark the head is not included in length measurements. beetles ( Scolytidae) .The males are often provided with The methods of citing data differ in some respects various kinds of cephalic and thoracic armature (horns, from those of previous Bulletins. All citations in synony- tubercles, plates) which resemble those of the larger mies are reduced to author, date, and page, but com- Scarabaeidae. According to Crowson (1960,1966), the plete references are given in the terminal bibliography. family should be included within the Section Hetero- Specimen repositories are given only for types, in which mera of the Cucujoidea and may be related to such case the following abbreviations are used: primitive heteromerous groups as the Mycetophagidae, BMNH, British Museum (Natural History) Tetratomidae, and pterogeniidae. phylogenetic The BYU, Brigham Young University position of the Ciidae, however, is still open to question. CAS, California Academy of Sciences The present work covers 27 species actually or CMNH, Cincinnati Museum of Natural History probably occurring within the confines of the state of MNHN, Mu&um National &Histoire Naturelle California, but it is based on a more extensive study of MZUH, Museum Zoologicum Universitatis Helsinki the North American Ciidae (Lawrence, 1971) and an USNM, United States National Museum examination of more than s0,oOO specimens from UW, University of Washington. various parts of the world. The terminology used in the California records have been reduced to locality only, keys and discussions is the same as that employed in without any further field data. These localities are listed The reasons for using the name Ciidae were given in a pre- alphabetically for each county, and the counties are also vious paper ( Lawrence, 1971 ). in alphabetical order. Since adult ciids may be collected 2 Bulletin of the California Iirnsect Survey in fungi throughout the year, as can be seen in table 1, weeks or more to become fully pigmented. It is not the exact collecting dates have been omitted. Host fungi known if adults in the field disperse immediately to are included in a separate section and other pertinent other sporophores, but in the laboratory several gen- field data are summarized in the discussion of bio- erations are easily reared on the same conk. nomics. By midsummer in most areas, large colonies of ciids Species of fungi are arranged according to the total are usually present within older fruiting bodies or in the number of records (over the entire geographic range), older portions of the larger perennial conks. Infested so that preferred hosts are listed first. An asterisk indi- conks may be recognized by the tiny exit holes and the cates at least one breeding record ( as used by Lawrence, presence of very fine, almost powdery, feces. Although 1971), while a double asterisk indicates a large number populations are smaller and development is slowed dur- of records for one host. The species concepts and gen- ing winter in most areas, larvae and adults may be eric classification are those of Lowe and Gilbertson obtained throughout the year, as may eggs and pupae ( 1961a, 1961b) and Overholts ( 1953). With the excep- in milder regions. tion of Gunoderma, the genera used are essentiallyform Because of their tendency to occur in large localized genera and do not reflect phylogenetic relationships. A populations, the Ciidae are a major food source for detailed account of polypore classification and its rela- various entomophagous arthropods working forest habi- tionship to host preference in ciids will be included in a tats. In California, trogositid beetles of the genus Tem- future publication ( Lawrence, 1973). nochila and anthocorid bugs of the genus Lyctocotis are known to be predators. A large number of parasitic BIONOMICS Hymenoptera are also associated with California ciids. Except for dispersal flights, about which little is These include Braconidae (Meteoms spp. and Euba- known, the Ciidae spend their entire lives in and around dizon spp. ) , Pteromalidae ( Jirnssoniella cuudata Ker- the fruiting bodies of Polyporaceae and other Basidio- rich), Eulophidae ( Astichus pulchdlineata Gahan ) , mycetes which grow on woody substrates. These fruit- and Bethylidae (Plustanoxus chittendendi Ashmead and ing structures, which may be referred to as sporophores, Cephalonornia perpusilla Evans), Larvae of certain brackets, or conks, are usually durable and often corky cecidomyid flies ( Lestodiplosis spp. ) may also feed on or woody in texture. Some ciids have been reported from ciid larvae. (Lawrence, unpublished.) under bark or in dead vines and twigs, but it is likely that they too were associated with fungi. Both larvae HABITAT AND HOST PREFERENCE and adults bore into and feed upon the sterile hyphae Since the Ciidae live in association with wood-rotting which comprise the bulk of the fruiting structures. fungi, they are restricted to those habitats in which Ciids are normally gregarious, and in the spring they dead wood is available and humidity is sufficiently high may congregate in large numbers on newly formed for the development and fruiting of fungi. As would sporophores. The females usually bore directly into the be expected, these beetles are particularly common in conk, while the males tend to remain more active on the the wet forests of the northern California coast and the surface. Copulation appears to take place on the surface west side of the Sierra Nevada. They also occur in the of the fruiting body, but this has been observed in only drier woodland and chaparral formations throughout a few species. The female constructs a main gallery and the state, and a few inhabit riparian situations in the hollows out egg cavities at irregular intervals along the arid regions of southeastern California. A rough idea of walls. After an egg is deposited, the cavity is closed with habitat preference is given in table 1, where species are a tightly packed plug of undigested frass. The egg- tabulated according to their occurrence in the four laying period may be long and there is an overlap in major Life Zones of the state (according to Grinnell, generations. 1935). The Life Zone concept, originally proposed by Ciid larvae are lightly pigmented and cylindrical, and Merriam (1892, 1898) on the basis of temperature normally bear two conspicuous hooks or urogomphi on criteria, has been criticized by several authors (Ken- the ninth abdominal segment. Upon hatching they bore deigh, 1932; Shelford, 1932), but the zones established directly into the fungus tissue and not through the frass in California by Hall and Grinnell (1919) have proven plug. There are five larval instars in the several species to be quite useful in discussing vertebrate distributions studied, and eclosion takes place about eight weeks and their relationships to a complex of environmental after eggs are laid; the generation time, however, may factors. The majority of Ciidae prefer the Transition be somewhat longer, since teneral adults take three Zone, but many of these extend into Upper Sonoran and hwrence: The Ciid Beetles of California 3 Boreal as well. Only a few occur in the Lower Sonoran be seen that only two species occur regularly on both and all of these are found along streambeds in associa- hardwoods and conifers, while the rest are usually asso- tion with cottonwoods. ciated with one or the other. Eight ciid species prefer The relative abundance of conifers and hardwoods in fungi growing on deciduous trees, while thirteen prefer an area is an important factor in determining the ciid those on conifers. Two of the latter, Cis creberrimus fauna (at least in California) since most species prefer and Dolichocis manitoba, are apparently restricted to fungi growing on one type of substrate. In table 1, it can conifers in California, but in eastern North America

TABLE 1. Seasonal and Ecological Distribution of California Ciidae Tree Adult Seasonal Activity Prefer- California (known collecting dates) ence Life Zones $8 gga mmhh3 %Be23 s CIID SPECIES SD>t.Crs C. acritus xx X xx C. americanus xxxxxxxxxxxx x x x xx x C. angustus xxx X X xx C. biarmatus xx xx X X C. creberrimus X ?X X? C. duplex X X X? C. ephippiatus X xxxxxx xx x xxx c. fuscipes xxxxxxxxxxxx xx (XI x xx C. hystriculus xxxxxxxxx x (X) xx xx C. levettei .?. ? P? C. maritimus X X X C.megastictus X X ?X C. striolatus X X ?X C. tetracentrum ? ? C. tidentatus xxxxxxxxxx x x xx X C. versicolor xxxxxxxxxxx xx (X) xxx x C. vitulus xxxxxxxxxxxx x x xx D. indistinctus x x xx X xx D. manitoba xx xxxxxxx ? x x xx Orth. punctatus xxxxx x xx ? ? XX? H. blaisdelli xx xxxxxxx x x x xx x x P, ctibrum xx xxxxxxxxx X x xxx Cer. californicus xxxxxxxxxxxx xx (X) xx Cer. dixiensis X X X Cer. similis ? ? s. curtulus xxxxxxxxxxxx xx (X) xxx Oct. laevis xxxxxxxxxxxx x xxx

X - any occurrenw XX - abundant (X) - rare or doubtful ? - assumed on basis of extralimital distribution 4 Bulletin of the California Insect Suruey they are commonly associated with hardwoods. ciated with conifers along the extreme northem coast. The subject of host preference in ciid beetles is a No other California Ciidae are known to breed in these complicated one which has been dealt with elsewhere fruiting bddies, although Fomes phi and Polyporus (Lawrence, 1967b, 1971, 1973; Paviour-Smith, 1960, gilvus are not uncommon in the state, and the latter is a 1969). Although few Ciidae are monophagous, some common host of certain Lepidoptera (Lawrence and have very narrow host ranges, while others prefer Powell, 1969). groups of related or similar fungi, which have been Most of the other fungi encountered in California fall placed in host preference groups by Paviour-Smith into the Ganodem applanatum group, which includes ( 1960) and Lawrence ( 1971 ) . Polyphagous species are a wide variety of fungi with at least three major types the exception, and most of these which have been ade- of fruiting bodies: (1)large, woody, perennial, lightly quately sampled show a preference for one group of pigmented, growing on conifers (Fomes pinicoh, F. fungi. annosus, F. 02piCcnal&); (2) laxge, corky, perennial, Among the California Ciidae, that with the broadest darkly pigmented, growing mainly on hardwoods host range is Cis americanus, which has been collected ( Ganoderma applanatum, G. brownii); (3) smaller, in the fruiting bodies of 24 different fungi and is known relatively soft-bodied, annual, lightly pigmented ( Poly- to breed in at least 18 of them. A second polyphagous powadustw, P. betulinus). Those ciids preferring the form is Cera& californicus,which breeds in at least 15 Fomes pfnicolatype include Cis angustus, C.biarmatus, different fungi but appears to have a preference for C. creberrimus, C. dupkx, C. megastidus, C. tridenta- species of Gunodem. Hadraule blaisddli has been tus, Dollchocis indistinctus, and D. manitoba. These are found in a wide variety of hosts and is known to infest all characteristic inhabitants of coniferous forests. Cis herbarium collections. The best example of an oligopha- americanus is often associated with F. pinicola and F. gous species is Pksioeis cribrum, which normally breeds annosus, but it is also common in Polyporus adustus in the small, whitish, globular fruiting bodies of PoZy- and a number of other hosts. Cis creberrimus in eastern porus volvatus on dead conifers. It may also be found North America breeds in a variety of fungi growing on on Polyporus anceps in the same habitats. hardwoods, including Polyporus adustus and Gam- The most obvious host preference group in this fauna ilerma lucidurn, but in the southwest it prefers the is the Polyporus versicolor group, which includes fungi lightly pigmented Fms on conifers. Species prefer- with thin, whitish, coriaceous fruiting bodies, such as ring Ganodemna applanatum and its relatives include P. versicolor, P. hirsutus, Lenzites betulina, Trametes Cis ephipplatus, C. Ievettei (in eastern North America, hispida, and related forms. The ciids preferring these at least), Cerucis caZifornicus, and C. similis (in Baja hosts are Cis fuscipes, C. tetracentrum, C. versicolor, C. California), Cis ephippiatus prefers G. upplanaturn in vitulus, C~acisdidensis, Sulcacis ~rtulirs,and Octo- coastal California, where the closely related Cis biar- temnzrs lasois. Ceracis califarnicus commonly breeds in mutus is found on Fomes pinicokz; in the Sierra Nevada, these fungi, but it is more often associated with species where Cis biamtus is absent, C. ephippiatus occurs on of Ganodem.Ceracis dixCensis is known to breed only Fomes pinkoh as well. in 2'. hispi&, while Cis tetracentrum has been collected only in P. uersicolol.. All of the fungi in this group occur DISTRIBUTION on deciduous trees and are rarely found on conifers. California Dist7ibution-The distribution of Ciidae A second group of hosts includes Polyporus parga- within the state is summarized in table 2, which shows menus, P. abietinus, and a few related forms with thin, the occurrence of each species in the various biotic coriaceous fruiting bodies having a purple or brownish areas originally proposed by Miller (1951) and map- pore surface. In California, all records are from P. ped by Hurd and Michener ( 1955). The absence of ubietfnuswhich is abundant on dead conifers. Ciids re- ciids from the Great Basin is partly an artifact of col- stricted to this fungus are Cis amitus, C.hystriculus, and lecting, while the few records for the two desert prov- c.Strioldus. inces are all from the Colorado River Basin. As wodd In eastern North America, a number of Ciidae in- be expected from the previous discussion of habitat habit the darkly pigmented, reddish-brown, woody, or preferences, the great majority of records are from the fibrous fruiting bodies of Polyporus giluus, Fonws northern coast and the Sierra Nevada, while a lesser robiniae, and their relatives. This group is represented number are from the foothills, inner coastal ranges, and in California by a single species, Cis maritimus, which soathem mountains. Very few ciids extend into the breeds in Fomes pini and Polyporus schweinitzii asso- Great Valley, and most of these occur along the major Lawrence: The Ciid Beetles of' California 5 river systems. Three species have not been recorded more likely to occur in mesic situations at intermediate from a specific locality, so that their occurrence within elevations. Finally, C. Ievettei occurs in eastern British Columbia, eastern Washingon, and the Rocky Moun- tains, and its occurrence in California would probably TABLE 2 be within the Sierran Montane region. Distribution of Ciidae in California According to Biotic Areas of Hurd and Michener ( 1955) North American Pattern and Faunal Afinities.-The majority of California Ciidae have northern distribu- Boreal Austral tion patterns usually corresponding to the Nearctic 8 Boreal of Powell (1964) or the C02 Type of Munroe B (1W).These widespread northern species may be further divided into those primarily western and rare 3 sx in the East (Cis ephippiatus, C. maritimus, Dolichocis b -2 c.. cribrum), U $ idistinctus, and Plesiocis those primarily Y eastern (Cis Zeoe#ei), and those fairly common on both Ciid c species c3 coasts (Cis americanus, C. fuscipes, C. striolatus, Or- C. acritus X? thocis pumtatus, Dolichocis manitoba, Sulcacis curtu- C. americanus xx lw, and Octotanus Zuevis). There are no species hav- C. angustus X ing the Western Boreal (or W3) type of distribution, C. biarmatus X C. creberrimus X since those occurring in the Rocky Mountains are also C. duplex X present in the Northeast. C. ephippiatus xxx A few species occur only along the Pacific Coast west C. fuscipes xx x C. hystriculus xxxxx of the Sierra Nevada. Two of these, Cis biarmatus and C. levettei ? C. hidentatus, are restricted to the immediate coast C. maritimus X from just south of San Francisco Bay to southern Alas- C. megastictus xx C. striolatus X ka. Each species is broadly sympatric with a sibling C. tetracentrum 3 species, Cb ephippiatus and C. americanus respective- C. tridentatus X ly, which has a much broader distribution. C. hystricu- C. versicolor x xx xxxx x C. vitulus xxx xx (XI Zua occurs both along the northern coast and in the Sierra Nevada; it is closely related to and possibly con- D. indistinctus xx D. manitoba xxx specific with C. hortidulw Casey, which occurs in eastern British Columbia, the Rocky Mountains and the Orth. punctatus X Northeast. A fourth Pacific coastal species is C. angm- H. blaisdelli x xx tus, which occurs in the Sierra Nevada and the Cascade Ranges; further collecting may reveal a wider distribu- P. cribrum x xx x tion. A final member of the Boreal group is C. mega- Cer. californicus x xx stictw, which is known only from a few montane Cer. dixiensis localities in northern California, but which is probably Cer. similis more widespread in the Pacific Northwest. s. curtulus xxxx MX All of the 17 species mentioned above probably have Old World af6nities, and most of them have closely ----Oct. Iaevis xxx X related counterparts in Eurasia. They would belong to X - any occurrence the modem Holarctic element of Linsley (1958). Two XX - abundant of these northem forms, Plesiocis cribrum and Dolicho- (X) - rare or doubtful mnitoba, to be more isolated and may rep- ? - assumed on basis of extralimital distribution cis appear resent older elements. The remainder of the California ciids have southern the state must be surmised on the basis of extralimital distribution patterns and Neotropical affinities. Cis distribution. Cera& similis occurs on the islands in the uituZus is practically endemic to the state, the only Gulf of California, and may be present in low dry areas extralimital record being from Oak Creek Canyon, near the Mexican border. Cis tetracentrum, on the other 6,000 feet, in northern Arizona. The species occurs in hand, inhabits mountain canyons in Arizona and is association with hardwoods in fairly wet situations 6 Bulletin of the California Insect Suruey along the coast and in the Sierran foothills. Its closest partment of Agriculture, Sacramento, Calif. (M. Was- relative, C. congestus Casey, is found in the Southeast, bauer); California Academy of Sciences, San Francis- while other members of the species group are Neotropi- co, Calif. (E. S. Ross, H. B. Leech); California Insect cal. This is the only species which may be a member of Survey, University of California, Berkeley, Calif. (J. the California faunal element of Linsley (1958), al- Powell, P. D. Hurd); Carnegie Museum, Pittsburg, Pa. though these "Californian" species are said to be asso- (E. Wallace); Cincinnati Museum of Natural History, ciated with more xeric communities. Cincinnati, Ohio (C. Oehler); J. F. Comell, Boone, Ceracis californicus is the most widely distributed of N. C.; Cornell University, Ithaca, N. Y. ( W. L. Brown, the southern forms, extending from Washington and H. Dietrich) ; Entomology Research Institute, Canada Colorado south into central Mexico, and occurring in Department of Agriculture, Ottawa, Canada (W. J. both lowland and montane situations. Cis acritus, C. Brown, E. C. Becker);'Field Museum of Natural His- creberrimus, and C. duplex are southern montane tory, Chicago, Ill. (R. L. Wenzel, H. S. Dybas); H. F. species, occurring only at higher elevations in associa- Howden, Ottawa, Canada; Illinois Natural History tion with conifers. C. creberrimus is also widespread in Survey, Urbana, Ill. (H. H. Ross, L. K. Gloyd); V. M. eastern North America, but there is some doubt that the Kirk, Brookings, S, D.; Museum National d"istoire southwestern populations are actually conspecific with Naturelle, Paris, France (A. M. Villiers); Museum of those of the East. C. acritus is closely related to the Comparative Zoology, Harvard University, Cambridge, eastern C. subtills Mellie. C.uersicolor, Hadraule blais- Mass. ( P. J. Darlington, Jr. ) ; Museum Zoologicurn Uni- delli, Cera& dixiensis, and Ceracis similis are primarily versitatis, Helsinki, Finland (M. Meinander); G. H. lowland species, although the first two occur at inter- Nelson, Kansas City, Mo.; Oregon State University, mediate elevations in the Sierra Nevada. All four ex- Cowallis, Ore. (J. D. Lattin); Purdue University, La- tend into Mexico and may inhabit fairly arid regions. fayette, Ind. (R. H. Arnett, L. Chandler); San Diego Cis tetracentrum is associated with hardwoods in the Natural History Museum, San Diego, Calif. (C. F. mountain canyons of southern Arizona and the Mexi- Harbison); J. Schuh, Klamath Falls, Ore.; Snow Ento- can highlands. These southern species are representa- mological Museum, University of Kansas, Lawrence, tives of Linsley's Sonoran and Neotropical faunal ele- Kansas (G. W. Byers); United States National Mu- ments. seum, Smithsonian Institution, Washington, D. C. (J. M. Kihgsolver, P. J. Spangler, T. J. Spilman, 0. L. Cart- Wright, D. M. Anderson); University of Alberta, Ed- monton, Alta. (G. E. Ball, D. Whitehead); University of Arizona, Tucson, Ariz. (F. G. Werner); University of ACKNOWLEDGMENTS British Columbia, Vancouver, B. C. (G. G, E. Scudder); Sincere thanks are extended to Professor E. G. Lins- University of California, Davis, Calif. (A. T. McClay, ley, under whose guidance the first part of this study R. 0. Schuster); University of Washington, Seattle, was carried out. Thanks are also given to the following Wash. (M.H. Hatch); University of Wisconsin, Madi- individuals for their help and advice during portions of son, Wisc. (R. D. Shenefelt, J. K. Ackerman); Utah thestudy: R. A. Crowson, H. B. Leech, C. D. MacNeill, State University, Logan, Utah (G. Knowlton). J. Powell, 0.W. Richards, R. F. Smith, the late R. L. Grateful acknowledgment is given to the mycologists Usinger, and P. Wygodzinsky. who have identified host fungi and given advice per- I am indebted to the following institutions and indi- taining to the systematics of the Basidiomycetes: L. viduals for allowing me to examine collections in their Bonar, 0. Fiddgo, R. L. Gilbertson, P. Lentz, J. Lowe, care: Academy of Natural Sciences, Philadelphia, Pa. 0. K. Miller, M. Nobles, R. Singer, J. A. Stevenson, and (H. J. Grant); American Museum of Natural History, I. Taveres. Thanks are also due to B. D. Burks, H. E. New York, N. Y. (J. Rozen, P. Vaurie, L. Herman); Evans, R. Gape, J. L. Herring, and W. Mason for the Brigham Young University, Provo, Utah (V. M. Tan- determination of predators and parasites. ner, S. L. Wood); British Museum (Natural History), I would also like to thank P. deBenedictis, J. A. London, England (J. Balfour-Browne, R. Pope, C. von Chemsak, J. T. Doyen, W. E. Ferguson, H. B. Leech, M. Hayek); Bureau of Entomology, California State De Lundgren, C. W. and L. B. OBrien, J. Powell, A. G. Luwence: TheCiid Beetles of California 7 Raske, G. I. Stage, and numerous other individuals who National Science Foundation Grant GB-4743, National have made a special effort to collect California Ciidae Science Foundation Fellowships, a Woodrow Wilson and their host fungi. Fellowship, and grants-in-aid from the University of I wish to thank Mrs. Celeste Green for her fine illus- California Committee on Research and the Society of trations and Mrs. Susan Duncan for the typing of the the Sigma Xi. Facilities of the Southwest Research Sta- manuscript. tion, Portal, Arizona, were made available to me by the Portions of this study were conducted with the aid of American Museum of Natural History. SYSTEMATICS

Although the Ciidae are the most common and confused with ciids are Pentaphyllus californicus Horn abundant inhabitants of Polyporaceae and related ( Tenebrionidae ), Eupisenus elongatus ( LeConte) wood-rotting fungi, they may occasionally be confused (Tetratomidae), and Sphindocis denticollis Fall (see with other small, fungus-inhabiting Coleoptera. The Addendum). All three have ll-segmented antennae, the following list includes some of the more obvious ex- first two have a 5-54 tarsal formula, Pentaphyllus has ternal characters which distinguish the Ciidae from the first three abdominal sternites fused, and Sphitt other small cucujoid beetles: (1)head usually declined dock has the fist two fused. and partly concealed by pronotum; (2) eyes oval, en- Ciid larvae have been illustrated by Boving and tire, and fairly coarsely facetted; (3) frontoclypeal area Craighead (1931)’ and Peterson ( 1957), and they may in male often raised to form a ridge, tubercles, or horns; be recognized by the following characters (based on (4) antennae 8- to 10-segmented,with a distinct 2- to more than half of the North American species, includ- 3-segmented club bearing large sensory organs (see ing all genera, and a number of exotic forms) : (1) body Lawrence, 1971); (5) last segment of maxillary palp cylindrical, without lateral extensions, lightly and subconical; (6) anterior edge of pronotum in male evenly pigmented, except for head, first thoracic ter- often bearing horns; (7) procoxal cavities narrowly gite, and posterior abdominal tergites; (2) head with open to closed behind, trochantins hidden; (8) elytra Y-shaped epicranial suture, 5 ocelli or less, and short never striate, epipleura narrow, extending almost to gula; (3) antennae 2-segmented, with a long, ventral, apex; (9) mesocoxal cavities not closed outwardly by sensory appendage near base of second segment and a sterna, trochantins hidden; (10) metacoxae narrow, long seta at the apex of that segment; (4) mandibles transverse, subcontiguous; (11) tarsi 4-4-4 in both bidentate, with or without “retinaculum” and mola; (5) sexes, segments simple, not lobed; (12) tibiae without maxillae with obliquely obtuse mala, and small, dorsal, apical spurs, the outer edge of protibia often expanded subapical, lobelike lacinia; ( 6 ) spiracles small, annular; and modified at apex; (13) abdomen with five freely (7) tergite 9 variously modified, but usually with two articulated sternites, first sternite without coxal lines, hook-like urogomphi; (8) sternite 9 without asperites, often with median pubescent fovea in male. Some Cali- segment 10 pygopod-like. The larvae are not treated fornia cucujoids which occur in fungi and might be further in this paper (see Addendum).

8 Lawrence: The Ciid Beetles of California 9 KEY TO THE SUBFAMILIES AND GENERA Genus Cis Latreille OF CALIFORNIA CIIDAE This is the largest genus in the family with about 350 1. Procoxae subconical, strongly projecting below intercoxal process, which does not extend to middle of named species distributed throughout the world. At coxae (figs. 1-2); metastemal suture absent; outer present, the North American fauna includes 43 species, edges of all tibiae spinose for more than half their of which 17 probably extend into California. lengths ( fig. 15);first visible abdominal sternite (111 ) in male with posteriorly-projecting, triangular flap, which partly conceals pubescent fovea (fig. 5); anten- KEY TO THE CALIFORNIA SPECIES OF CIS nae &segmented...... OROPHIINI, Octotemnw (p. 23) Procoxae transverse or globular, not projecting below 1. Elytral punctation dual, consisting of larger, shallow, intercoxal process, which extends beyond middle of nude megapunctures and smaller, deeper micropunc- coxae (figs. 3-4); metasternal suture present; outer tures, which bear bristles or hairs...... 2 edges of tibiae not spinose or bearing spines at apices Elytral punctation single, the punctures fairly uniform only (figs. 7-14); first visible abdominal sternite in in size and all or most of them bearing bristles or male simple or foveate, but without triangular flap hairs ...... 12 (fig.6) ...... CIINI,2 2. Vestiture consisting of 6ne hairs ...... 3 2. Prosternum in front of coxae almost twice as long as Vestiture consisting of short, stout bristles...... 7 intercoxal process (fig. 17); sides of pronotum slight- 3. Elytral hairs longer, more than 0.20 times as long as ly diverging towards apex; body mall (TL usually scutellar base and visible under 10 X magnification, less than 1.4 mm.), elongate and extremely flattened decumbent ...... 4 (fig. 33 ); antennae 9-segmented with a %segmented Elytral hairs very short, less than 0.15 times as long as club ...... HadrauIe (p. 19) scutellar base and not visible under 10 X magnifica- Prosternum in front of coxae not or slightly longer than tion, erect or inclined ...... 5 intercoxal process; sides of pronotum not diverging 4. Pronotal punctation finer and sparser, punctures sub- towards apex; body usually larger and less flattened; equal to eye facets and separated by 1.0 to 1.5 antennal club 3-segmented ...... 3 diameters; body usually smaller, narrower, and bi- 3. Outer apical angle of protibia expanded, rounded, and colored, brownish with black pronotum and trans- bearing several spines (figs. 13-14) ...... 4 verse elytral macula...... C. ephippiatus ( p. 13 ) Outer apical angle of protibia narrowly rounded (fig. Pronotal punctation coarser and denser, punctures larg- 7), blunt and angulate (fig. 9), or produced and er than eye facets and separated by 0.33 to 0.66 dentate (fig. 11 ), not spinose...... 5 diameter; body usually larger, broader, and more 4. Intercoxal process of prostemum laminate, less than 0.15 uniformly pigmented, brownish. ...C. biamtus (p. 12) X as wide as a procoxal cavity; vestiture consisting of 5. Outer edge of protibia irregularly notched or serrate for very short, fine hairs; antennae 8- or 9-segmented...... part of its length; pronotal punctation finer and Ceructs (p. 20) sparser, punctures smaller than eye facets and sep- Intercoxal process of prosternum not laminate, at least arated by more than 1.0 diameter; lateral edges of 0.20 X as wide as a procoxal cavity; vestiture consist- pronotum smooth; elytral punctation not distinctly ing of short, stout bristles; antennae 10-segmented...... seriate ...... C. leoettei (p. 14) suzcucis (v.._ 22 ) Outer edge of protibia simple; pronotal punctation 5. Antennae 10-segmented ...... 6 coarser and denser, punctures usually larger than eye Antennae 9-segmented ...... 7 facets and separated by less than 1.0 diameter; lateral 6. Outer apical angle of protibia narrowly rounded (fig. edges of pronotum coarsely crenulate; elytral puncta- 7); elytral suture with an inflexed margin near apex tion distinctly senate ...... 6 (fig. 16); head and pronotum in both sexes simple; 6. Anterior angles of pronotum broadly rounded; pronotal elytral punctation single and uniform; vestiture Con- punctures very dense, separated by 0.33 diameter sisting of very short, fine hairs; body elongate and or less; interspaces smooth; elytral megapunctures 4 parallel-sided...... Orthocis ( p. 18 1 times as large as micropunctures; abdominal fovea Outer apical angle of protibia usually produced and in male circular ...... C. megustictus ( p. 15) dentate (fig. 11) or blunt and agulate (fig. 9), if Anterior angles of pronotum subacute; pronotal punc- somewhat rounded, then vestiture consisting of short, tures not as dense, separated by 0.50 to 0.66 diam- stout bristles and elytral punctation seriate; elytral eter; interspaces granulate; elytral megapunctures suture without inflexed margin; head of male usually less than 4 times as large as micropunctures; abdom- bearing plates, teeth, or tubercles ...... Cis (p. 9) inal fovea in male longitudinally oval ...... 7. Outer apical angle of protibia rounded (fig. 12); body C. muritimua (p. 14) more elongate, TL/EW more than 2.25; apex of pro- 7. Anterior angles of pronotum distinctly produced and notum simple in both sexes...... Doltchocis ( p. 18) broadly rounded, the disc impressed above each Outer apical angle of protibia distinctly produced and angle (fig. 19); lateral margins of pronotum broader, dentate (fig. 8); body shorter and stouter, TL/EW easily visible for their entire lengths from above, their less than 2.20; apex of pronotum in male bearing two edges smooth or barely crenulate; prosternum in horns or tubercles...... Pledocis ( p. 20) front of coxae tumid but not carinate; TL usually 10 Bulletin of the California Insect Survey more than 2.00 mm ...... C. fuscipes (p. 13) base; lateral pronotal margins not or barely crenulate; Anterior angles of pronotum not or barely produced and body somewhat flattened ...... C. creberrimus (p. 12) subacute (fig. 18); lateral pronotal margins distinctly Elytra sparsely clothed with shorter, stouter bristles, crenulate, usually narrower; prosternum flat or which are uniformly distributed and much shorter carinate; TL usually less than 2.0 mm...... 8 than scutellar base; lateral pronotal margins distinctly 8. Prosternum in front of coxae distinctly carinate; body crenulate; body not flattened...... 15 short and broad, oval, EL/EW usually less than 1.40; 15. Body shorter and broader, EWEW less than 1.45; anterior edge of pronotum in male produced to form prosternum in front of coxae weakly carinate, the a subtriangular process (fig. 20) ...... 9 lateral portions concave; anterior edge of pronotum Prosternum in front of coxae flat or slightly tumid, not in male armed with 2 subtriangular horns...... carinate; body more elongate and parallel-sided, EL/ c. duplex ( p. 12 ) EW usually more than 1.40; anterioi edge of prono- Body longer and narrower, EL/EW more than 1.45; tum in male simple ...... 10 prosternum in front of coxae flat or slightly tumid; 9. Elytral punctation obscurely dual, megapunctures bare- anterior edge of pronotum in male simple or with 2 ly larger than micropunctures. ....c. tl.fdentatu8 (p. 16) small tubercles ...... 18 Elytral punctation distinctly dual, megapunctures 1.50 18. Elytral bristles shorter, stouter, and colorless or very to 3.0 times as large as micropunctures...... pale yellow; size larger, TL usually more than 1.85 C. americanus (p. 10) mm.; pronotal apex in male usually bituberculate; 10. Elytral punctation confused; elytral bristles shorter and male without abdominal fovea. ...C. hystriculus (p. 14) stouter, about 0.33 times as long as scutellar base, Elytral bristles longer, finer, and bright yellow; size colorless, and blunt at tip...... C. acritus ( p. 10) smaller, TL usually less than 1.85 mm.; pronotal apex Elytral punctation distinctly seriate; elytral bristles in male simple; male with abdominal fovea...... longer and narrower, usually 0.50 times as long as C. angustus (p. 11 ) scutellar base, yellowish, and acute at tip ...... 11 11. Outer apical angle of protibia not produced, rounded or obtusely angulate (fig. 9); lateral margins of pronotum broader, easily visible for their entire Cis am'tus Lawrence lengths from above; pmnotum usually lightly granulate and shiny, reddish in calor, and paler than elytra; Cis actitus Lawrence, 1971:443-444, figs. 52, 63, 73, 80, 106. frontdypeal ridge in male simple. .C. oerslcdor (p. 18) Holotype, 8, Rustler Park, 8 mi. W Portal, Cochise County, Outer apical angle of protibia at least slightly produced, Arizona (CAS). usually forming a distinct tooth (fig. 10); lateral margins of pronotum narrower, not or barely visible Geographic range,-Montane regions of southern California, for their entire lengths from above; pronotum usually Arizona, and New Mexico. distinctly granulate and dull, dark brown or black in color, as are elytra; frontoclypeal ridge in male bi- California ?eCOrds.-FREsNO Co.: 3 mi. E. Shaver Lake. KERN tuberculate...... C. striolatus (p. 15) Co.: 1 mi. SE Alta Sierra. 12. Body larger and broader, TL at least 2.00 mm. and EL/ EW less than 1.50; anterior angles of pronotum Host fungus.-Polyporus abietinus.* strongly produced and rounded; lateral pronotal margins broader, easily visible for their entire lengths Discussion.-This is the only California ciid in which from above, with a raised lip (fig. 19)...... 13 the body form is elongate, the elytral punctation is dual Body smaller or narrower, TL less than 2.00 mm. or and confused, and the vestiture consists of short, stout, EL/EW more than 1.S; anterior angles pmnotum of colorless bristles. Cis srriolatus and C. uersico2or are not or slightly produced and subacute; lateral pm- notal margins narrower, not or barely visible for their similar in appearance, but in both species the elytral entire lengths from above, without a raised lip. .... 14 punctation is seriate. C. creberrimus differs in having 13. Pronotal punctures only slightly derthan elytral single elytral punctation and longer, finer vestiture. Al- punctures; elytral bristles longer and her, more than though apparently rare, C. adtw may occur more 6 times as long as wide and acute at apex; hnto- clypeal ridge in male with 2 tiangular plates; male widely at higher elevations in southern California. It is with abdominal fovea; elytra in female more than 2 to be sought among dead coniferous trees and logs, in times as long as pronotum...... C. tetracentrum ( p. 18) the fruiting bodies of Polyporus abietinus, which are Pronotal punctures much smaller than elytral punctures; normally inhabited by C. hystriculus. elytral bristles shorter and stouter, less than 6 times as long as wide and blunt at apex; frontoclypeal ridge in male elevated and trisinuate, SO that 4 teeth are Cis americunus Mannerheim formed; male without abdominal fovea; elytral in female less than 2 times as long as pronotum...... (Fig. 20) c. UittdU8 (p. 17) 14. Elytra densely clothed with long, fine, acute bristles, Cis americanus Mannerheim, 1852:360. Lectotype, 8, Sitka which are subseriate and almost as long as scutellar Island, Alaska ( MZUH ) . Lawrence: TheCiid Beetles of California 11

1 ur*r.+ l YZltl u "I Y z. ,I "I I" m L = Y u Y I" ,I I,. I. I" ".

Map. 1. California distribution of Cfs ammicanus Mannerheim. Map. 2. California distribution of Cis angustus Hatch. Inset: Inset: Distribution in North America. Distribution in North America.

Syfionyms.-Xestocis insokns Casey, 1898:W. Cis frosci Dury, dentatus in having the elytral megapunctures distinctly Cis m~3colZis Dury, Xsstocis minor Hatch, 19179. 1917:9. larger than the micropunctures. americtllztcS is ex- 1962:232. Xestocss oweni Hatch, 1962:!232. Xestocis striguh Cis Hatch, 1962:231. Cis hatchi Lawrence, 1971:445, replacement tremely variable throughout its range and may repre- name for Xestocis nitidus Hatch, 1962:233 (not Anobium nWum sent a complex of species with C. tridentatus. On the Fabricius, 1792). West Coast there are two fairly distinct forms which differ in size, relative abundance of megapunctures, Geographic range.-Northern part of North America, from Alaska to Nova Scotia, south to central California, northern density of elytral bristles, and width of pronotal mar- Utah, Colorado, and western No& Carolina (map 1 ). gins; these two forms may be sibling species (we Lawrence, 1971). Californiareconls.-Widespread in the Coast Ranges and in the Cis americanus inhabits a wide variety of fungi and Sierra Nevada through the northern half of the state, ranging up is known to breed in at least 17 species. In coastal Cali- to elevations above 6,000 feet on both sides of the Central Valley. fornia it commonly breeds in Polyporus adustus and Steteum himtum, while montane populations may Host fungi.-Polyponrs adurtucQ; Poltiponu betulinusQ;ster- occurinFomespinicda. eum himturn*; Fomes pi&laQ; P*m versicolorQ;hnw~ annosus*; Poria wrsiporae; Stecche+inum ochmceumQ; P2eurotw ostreatus*; PoIyponrp sulphureuso; Ganodermo tsugw'; Poly- Cis angustus Hatch porus abietinw, Trameta m&*; Gmodmma applmudumQ; Pollrpotus r-0; Polgpolus MrmtusQ; PoIyporus pmga- Cis angustus Hatch, 1962:230. Holotype, P, Stanley. British nwQ; Pol11pm sqwmos~r~;PolVponu t-; Poria Columbia (CAS). nigrescens; Fomes fomentarius; Polyporus bifmis; Phlabia merismoideS. Geographic range.-Mountains of the Pacific Coast, from south- central British Columbia to the southern Sierra Nevada (map 2). Discussion.-mis species is characterized by the Califonzia recwds.-Emnmo Co.: Lake Tahoe. FRswo Co.: short, broad form, by the carinate prosternum, by the Huntington Lake. LASEN Co.: Black's Mountain; Fa&. MADERA dual elytral punctation, and by the vestiture of short, Co.: 7 mi. NE The pines, Bass Lake, 5000'. MARIPOSACo.: 6 mi. stout bristles. It d&rs from the closely related C. ti- NW Fish Camp. PLW Co.: byonDam. TnuMn Co.: 7 mi. 12 Bulletin of the California Znsect Survey N Mineral, 6000'. TULARECo.: Dorset Camp, Sequoia National Synonyms.-Cis puberulus Mellik, 1848:358. Cis nubillus Gor- Park. ham, 1898:331.

Host fungi. - Fomes pinicold; Fomes annosus; Fomes of- Geographic range. - Eastern North America, from Vermont ficinalk. south to Florida and west to eastern Nebraska, Kansas, and Texas; montane regions of the Southwest; throughout the West Discussion.- This species is characterized by the Indies; and scattered localities in Mexico, Central and South elongate and somewhat cylindrical body form, by the America. single, coarse, and confused elytral punctation, and by California records.-Sm BERNARDINOCo.: 4 mi. E Running the moderately long and fine, yellowish bristles. It is Springs. SANDIEGO Co.: 1.5 mi. NW Mt. Laguna. most easily confused with C. hystriculus, which is usually larger, with shorter and stouter colorless Host fungi.- Fomes scler~";PolIlporus adustus'; bristles, and with no abdominal fovea in the male, but Ganodetma lucidurn"; Chnodenna upplanaturn; Polyporus hyd- the two species not likely to found in the same noides'; Fomes oficbuh"; Porin Intemorginota'; Fomes pini- are be coho; Polyporus pargumenus"; Trametes comgato"; Polyporus fruiting bodies. DoZichocis manitoh is also similar and gilws; Fomes annosus"; Ganodewna fulueUurn"; Polypofirs may be found in the same kinds of fungi, but that sanguineusO; Poria corticolaQ;Daedalea ekgrms; Lsnzites striata; species differs in having %segmented antennae, round- Plenotus sp.; Polllpofirp ~ceps;Polypotus hirsutus, Polypo~ ed protibial apices, and an anteriorly constricted pro- sulphureus; Polyporus supinus; Schizophyllum commune; Tmm- etes iaispida. notum, Cis mebewitnus Mers in bing somewhat flattened with finer and denser elytral punctation. C. un- gustus appears to be restricted to the mountains of the Discussion.-Cis crebdrnus is distingmhed by the Pacific Coast, where it occurs at higher elevations only. elongate and flattened form, by the single punctation, and by the dense vestiture of fairly long and fine Cis biarmatus Mannerheim bristles. The vestiture in this species is actually dual, consisting of erect and inclined bristles differing only (Fig. 21 ) slightly in length, but this is not obvious at lower mag- nifications. There is some doubt as to whether the Cali- Cis biarmatus Mannerheim, 1852:360. Lectotype, 8, Sitka fornia populations are conspecific with the eastern C. Island, Alaska ( MZUH ) . crebkrrimus, in which the pronotal punctation is finer Geographic range. - Pacific Coast of North America, from and the elytral bristles much shorter. The variation in southern Alaska to Marin County, California (map 3). this species or complex is discussed further in Lawrence ( 1971). California records.-Dn. NOR= Co.: Crescent City. HUM- BOLDT CO.: 20 mi. E. Arcata; Fieldbrook. MARIN CO.: 2 mi. SW Inverness. MENWINOCo.: Caspar. Cis duplex Casey

Host fungi.-Fomes pinicolu". (Fig. 22)

Discussion.-This species differs from most Califor- Cis duplex Casey, 1898:82. Holotype, $, California (USNM). nia ciids by the short, broad body form, by the distind- Geographic range. - Mountains of the southwestern United ly carinate prosternum, by the dual elytral punctation, States, from southern California east to north-central New Mexico and by the vestiture of moderately long, decumbent, and south as far as Morelos, Mexico and the southern tip of fine hairs. It may be distinguished from the closely re- Baja California. lated C. ephippiatus by the coarser and denser pronotal punctation, by the larger size, and by the uniform color- Californib records.-Sm DLEGOCo.: 1.5 mi. NW Mt. Laguna. ation. Throughout its range, Cis kmtusis sympatric Host fungi. - Polyporus anceps'; Fomes oficinalis'; Fmes with C. ephippdatus, but the two species have never pinkoh'; Ganoderma oregonenseO. been taken on the same conks (see p. 13). Discusslon.-This species is easily recognized by the Cis creberrimus Melli6 fairly short and broad form, single and confused elytral punctation, vestiture of short bristles, narrow, crenu- (Fig. 23) late lateral pronotal margins, and the sexual characters of the male, which consist of two subtriangular plates Cis cseberrimus Mellik, 1848:357, pl. 12, fig. 5. Lectotype, 0, New Orleans, Louisiana ( MNHN ) . on both the clypeus and the pronoturn. Map. 3. California distribution of Cis biarmatus Mannerheim. Map 4. California distribution of Cis ephippiatus Mannerheim. Inset: Distribution in North America. Inset: Distribution in North America. Cis ephippiattcs Mannerheim populations of C. ephippiatus are usually collected in the fruiting bodies of Gunohspp., while the sym- Cis ephippiatus Mannerheim, 1853:234. Types. Sitka Idand, patric c. bb-w is re&i&& to Fames pinkoh. Alaska (MZUH?). Montane populations of C. ephippiatus, however, occur Synonym.-Xestocis moznettei Dury, 1917: 16. on Fomes pinicola as well.

Geographic range.-Western North America, from southern Cis fuscipes Mellid Alaska south in California to Alameda County and the southern Sierra Nevada, and in the Rocky Mountain Region to northern (Figs. 24 and 25) Nevada and Colorado. Also known from Vermont, New Hamp- shire, and the Gasp6 Peninsula of Quebec (map 4). Cis fuscipes Mellik, 1848:271, pl. 2, fig. 23. Lectotype, 0, Boston (MNHN). Cdijornia recor&.-ALAMEDA co.: Berkeley. Dn. NORTECO.: 18 mi. S. Klamath. -NO Co.: Huntin@m Laore. HUMBOLDT Synmymc.-Cis atripennis Me&& 1848:258, pl. 2, fig. 15. Cis Co.: 20 mi. E Arcata; 8 mi. S KorbeL Mnam Co.: Alpine Lake; cheuroktti MellM, 1848:MQ.Cis dubfus Mellih, 1848:273. Cis S. P. Taylor State Park. Mec~ocmoCo.: 1 mi. N Albion; Little cwolinos Casey, 1898:78. Cis impreaa Casey, 1898:79. Cis River; Mendocino. PLUMASCo.: Budc's Lake; 7 mi. NE Buck's pnuenS Casey, 1898:78. Lake. Geogmphic range. - Widespread and abundant throughout Host fungi.-Gamderma appknatumQ; Fomap ptnicoh'; most of northern and eastern No& America, from northem Ganoderma brmii"; Fomea annosus, PO- W&W, P&- British Columbia, south to Los AngeIes County, California, east porus resinosw, Polypars .pulphum, Poria OerSipMa. across Canada to Nova Scotia, and south through eastern and midwestern United States (east of the 100th meridian) to Discussion.-This species closely resembles Cis biar- southern Texas and Florida (map 5). Also known from Cuba, mutus, but may be distinguished by the finer and spars- Madeira, and Hawaii. er pronotal punctation, smaller size, and normally bi- Caljjomiu records. - ALAMEDA Co.: Berkeley; Oakland. colored elytra. In California, it OCCUTS both along the CONTRACOSTA Co.: Tiiden Park, Berkeley Hills. HUMBOLDT northem coast and in the Sierra Nevada. The coastal Co.: Garbedq 8 mi. S KorbeL Los ANGELFS Co.: (no speci6c 14 Bulletin of the California Insect Survey locality). MmCo.: Alpine Lake; Carson Ridge; Inverness; 1 guese Pass, 7 mi. SE Pine Flat; Round Meadow, 64W, Giant mi. SE Inverness; 2 mi. N Invemess; Lagunitas; Mill Valley; Forest. TUOLUMNECo.: Dardanelle; Leland Meadow; 7 mi. NE Muir Woods; S. P. Taylor State Park; Tomales; Taylorville. Strawberry. MEND~CINOCo.: 2 mi. N Piercy. MONTXREYCO.: Big Sur; Carmel; Monterey. NAPACo.: Calistoga; Napa. PLUMASCo.: Host fungi.-Polyporus abietinw4*; Poria cinerascens; Poria 7 mi. NE Buck's Lake. SACRAMENTOCo.: Sacramento. SAN uersipora. FRANCISCOCo.: San Francisco. SAN MATEO Co.: Hillsborn; Salada Beach. SANTACLARA Co.: 2 mi. SW Los Gatos. SANTA Dism&on.-This species may be distinguished by CRUZ CO.: Ben Lomond. SHASTACo.: 1.5 mi. S Castella; Viola, 4000'. SISKNOUCO.: Dunsmuir; Hamburg; Weed. the elongate, subcylindrical body form, single, coarse, and confused elytral punctation, vestiture of short, Host fungi. - Polarpofirr uersicoloroo; PowhirsutwQ; colorless bristles, distinctly dentate protibial apex, and Lenzites betulina"; Polllponrs pubescensO; Polyporus conchiferQ; lack of an abdominal fovea in the male. Cis angustus is Ganodermu browniio; Polyporus adustwo; Po- squanw- usually smaller, with yellowish bristles and an abdomi- SUP;Daehlea atnbigtq Fomes fiaximphilur, Fomes pinicoh; Ganodenna applanatum; Po+ subectp. nal fovea in the male, while C. creberrirnus is flatter, Discussion.-This species may be distinguished by with finer and denser punctation and much longer and the fairly large size, dual and subseriate elytral puncta- finer vestiture. tion, moderately long elytral bristles, produced and Cis hystriculus is known only from the Pacific states, rounded anterior pronotal angles, broad lateral pro- but it is closely related to and possibly conspecific with notal margins, and impressed pronotal disc in the male C. hmidulus Casey, which occurs throughout the (fig. 24). Cis uersicolor and C. SttiOZutus are usually Rocky Mountains and the northeastern part of the con- smaller and more elongate, with narrower pronotal tinent ( see discussion in Lawrence, 1971 ). margins and differently constructed anterior angles, while C.americanus and C. tridentatus have a carinate Cis levettei (Casey) prostemum and different sexual modilkations in the male. Xestocip levettei Casey, 1898:83. Holotype, 8, Indiana? (US NM). Cis fuscipes is most common in northern coastal Cali- fornia and is practically absent from the Sierra Nevada Geographic mnge.-Widespread in North America east of the and the southern part of the state. It prefers humid 100th meridian, from Newfoundland south to Alabama and west areas where it occurs in Polyporur uersicolor and its to Manitoba, Kansas, and Texas; in the western part of the continent, recorded from Alberta, northwestern Colorado, eastern. relatives. It is parthenogenetic throughout most of its British Columbia and Washington, and California. range, but in California many populations are bisexual; the biology of this species is further discussed in Law- CaZiforniorecord-No specific locality ("Cal."). rence ( 1967a). Host fungi.-Ganodma applanatum" *; FmfomentMiwQ; Fomes pinicola"; Ganoderma tsugae*; Polyporus uersicobr; Cis hystriculus Casey Ganodem lucidurn*; Polyporus betulinusQ; Polyporus pargam- enus; Polyporus pubescens; Stereum ostrea; Daedalsa confra- (Fig.26) gosao; Polgporus resfnosus";Poria nigrescens'; Fomes robiniae; Lenzites betulina; Polyporus adustus; Polyporus squamosur; Cis hystriculus Casey, 1898i82. Holotype, 8, Lake Tahoe, Cali- Poria vitrea. fornia (WSNM). Discw&m.-The of Cis kuettei in the Cali- Geographic mnge.-Westem British Columbia, Washington, inclusion and Oregon, south along the Caiifomia coast and through the fornia fauna is based on a single specimen from "Cal." Sierra Nevada to the Transverse Ranges in the southern part of which might be mislabelled. The species does occur in the state (map 6). eastern Washington, however, and may extend into northern California. It resembles C. mufitimus but dif- Califonziarecotds.-AwMEDa Co.: Berkeley; Oakland. ALPINE Co.: Ebbett's Pass, 8730'. EL DORAWCo.: Lake Tahoe. FRESNO fers in having much her and sparser pronotal punda- Co.: Hudrieberry Meadow; Huntington Lake; LaLeshore; 3 mi tim. E Shaver Lake. KERN CO.: 1 mi. SE Alta Sierra; 11 mi. W Frazier Park. MARINCO.: Alpine Lake; 1 mi. SE Inverness; 2 Cis maritimus (Hatch) mi. SW Inverness; Lagunitas; Muir Woods. MARIPOSACo.: 6 mi. NW Fish Camp. MEND~CXNOCO.: Caspar; 2 mi. N Piercy; (Fig. 27) Van Damme State Park. MONTEREYCo.: Cannel. R~~~RSIDECO.: 2 mi. NE Idyllwild. SAN MA- Co,;Montara. SANTACRUZ Xestocis muritimus Hatch, 1962:233. Holotype, 8, Ocean Park, CO.: Ben Lomond. SHASTACo.: Castella. TTJLARECO.: Portu- Oregon ( UW ) . Lawrence: TheCidd Beetles of California 15

Map. 5. California distribution of Cis fusdpes Mel€i& Inset: Map. 6. California distribution of Ckr hysttfcuhs Casey. Inset: Distribution in North America. Distribution in North America.

Geographic range.-Pacffic Coast from extreme northwestern Geographic range. - Known only from montane regions in California to southwestern British Columbia. Also known from northern California. south-central Manitoba, California records.-CuvEzw Co.: Calaveras. GLENNCO.: California record.-Dm. NORTECo.: Crescent City. Plaskett Meadows, 6200'. PLUMASCo.: Bucks Lake.

Host fungi.-Po2yporus schweinitziiQ;Fms phiQ. Host fungi. - Fomes annosus'; Fomes pinicola; Polypm sulphurmu. llfadon.- This species resembles Cis hettei, C. biarmatus, and related forms with a carinate proster- DiscucFsion.-This species resembles Cis mritimus in num, but it may be distinguished by its vestiture of very general form, vestiture of short, fine hairs, and coarse, short and fine hairs, the finely granulate and shiny pro- dense pmtal punctation; it may be distinguished, notum with large dense punctures, the produced and however, by the rounded anterior pronotal angles, larg- acute anterior pronotal angles, and the longitudinally er elytral megapunctures, and round abdominal fovea oval abdominal fovea in the male. Except for the Mani- in the male. Cfs bhmatus is also similar, but differs by toba record, Cis maritim is known only from the nar- the her punctation and longer, decumbent hairs. row, humid, coastal strip extending from northwestern Cis megastichcs occurs at higher elevations in the California to British Columbia. This appears to be the conifer forests of northern California, but it does not only California ciid which breeds in the reddish-brown appear to be common. fruiting bodies of Fomes pi& and Polypmschwelnit- zti. Cis striohtus Casey

Cis striolato Casey, 1898:79. Holotype, '2, Salida, Colorado megastictus Lawrence Cb (USNM). Cis megastictus Lawrence, 1971:485-7, figs. 39, 55, 84, 74,78. Synonyms.-Ckr fiaterna Casey, 1898:80. Cis madlenta Casey, Holotype, 8,Bucks Lake, Plumas County, California (CAS). 1898: 80. 16 BulEetin of the California Insect Suwey Geographic range.-Widespread across the noithem part of North America from the Mackde District, Northwest Temtory, tu Nova Scotia, south into the Sierra Nevada and Rocky Mountains, in the Midwest as far as Kansas, and on the Atlantic Coast as far as northern Florida.

Cdifornin records.-Er. Do- Co.: Lake Tahoe. TUOLUMNE Co.: 21 mi. NE Strawberry.

Nost &ngi.-P- Obiet61uPo*; Poltrporus p~gmn~nu~'~; Daedaleo ~ntcoh~;Po- hirsutw*.

D~cussion.-Cis sOviolatus resembles C. versicolor m its elongate, somewhat flattened form, dual and distinctly senate elytral punctation, and vestiture of short brides; it differs from that species, however, in having the protibial apex at least slightly produced and angulate, the lateral pronotal margins narrower, the pronotal surface more granulate and the color dorm. In C. actitus the elytral punctation is confused and the elytral bristles are usually shorter and colorless.

Cis tetracentrum Gorham - I. (Fig. 28) IC.. 1. u - A I; A I. _-?I

Cis tetracentrum Gorham, 1886:357. Lectotype, $, Northern Map 7.California distribution of Cb tridentatus Mannerheim. Sonora, Mexico ( BMNH ) . Inset: Distribution in North America. Synonym-Cis arizonae Dury, 1917:s. California records.-Dm NORTE Co.: Crescent City. HUM- BOLDT Co.: 20 mi. E Arcata; Fieldbrook. MARIN CO.: Alpine Geographic range.-Mountains of southem Arizona and Cali- Lake; Invemess; 1 mi. SE Invemess; 1 mi. NW Invernas; fornia south through the Mexican highlands as far as central Lagunitas; Muir Woods; Tocaloma; Woodacre. ME"O CO.: Veracruz. 1 mi. N Albion; Caspar; Fort Bragg; Pigmy Forest. MONTEREY Co.: Big Sur; Camel. SANTACLARA CO.: Los Gatos. SANTA CaIijornia record.-No specific locality ('Tal.''). CRUZCo.: Ben Lomond. Host fungi.-Fomes piniwkP *; Pmla cineracetwo; Cano- Host fungus.-Polyporus versicoloro. dewnu oregonewe*; Polyporus sulphureusa; Poria carbonka'; Tranwtes sepium' ; Ganoderma applanatum; Pleurotus ostreatus; Discusslon.-Cis tetracentrum is doubtfully included Polyporus gilous. on the basis of an old Morrison specimen labelled "Cal." It should be sought in the fruiting bodies of Poly- Discussion.-This species differs from the closely re- porus vsrsicolor or related fungi growing on hardwoods lated Cis umericunus in having obscurely dual elytral at intermediate elevations in the southern California punctation, with the megapunctures barely larger than mountains. the micropunctures. Cis versicolor Casey CL tridentatus Mannerheim (Fig. 29) Cis tridentatus Mannerheim, 1852:380. Syntypes, Sitka Island, Alaska (MZUH). Cis versicolor Casey, 1898:80. Holotype, 9,Calaveras County, California ( USNM 1. Synonyms.-Xesto& ednae Hatch, 1982:%32.Xestods rej2exus Hatch, 1962:232. Geographic range.-Extreme southern Oregon, south throughout most of California, west of the Sierran crest, into Baja Cali- Geographic range.-Pacific Coast from southern Alaska to fornia and east through Arizona into New Mexico and western Monterey County, California (map 7). Texas (map 8). Lawrence: The Ciid Beetles of California 17

- =.,-...%%.-J 1 -.$.I- ," ,. ,l. /. ,<> 1Y 6.w u tn - I. ,. m. ,- ". ". -

Map 8.California distribution of Cis versicolor Casey. Inset: Map. 9. California distribution of Cis uitulus Mannerheim. Inset: Distribution in North America. Distribution in North America.

California records.-Common and widespread through much Cis vctuZus Mannerheim of the state in both arid and mesic environments, from low to intermediate elevations, ranging up to above 5,000 feet in south- (Fig. 30) ern California. Apparently lacking from the Mojave Desert except along the Colorado River and from the east side of the Sierra Cis vitulus Mannerheim, 1843:299. Types, California (MZUH?). Nevada, but lack of records from the northern counties probably represents sampling error. Synonyms.-Cis cmeN Dalla Torre, 1911:8, replacement name for Cls illustris Casey, 1898:81 (not Brow, 1880). Host fungi.-Polyporus versicoloroO; Trarnetes hispiduo; Palp porua hirsutuao; Lenxites betulina'; Gamderma brownii; Poly- Geographic range.-California, from Del Norte County to San pmcinnabarinus; SchizophyUum commune. Diego County and north-central Arizona (map 9 ) .

Discussion.-This species may be distinguished by California records.-Widespread at low to moderate elevations the elongate and somewhat flattened form, dual and throughout much of cismontane California. This species is most seriate elytral punctation, vestiture of short bristles, common in mesic situations along the northern coast and in the foothills of the Sierra Nevada, ranging up to about 3,000 feet rounded or angulate protibial apex, shiny pronotum elevation in the north and to above 5,000 feet in the mountains with fairly broad lateral margins, and lack of fronto- of the southern counties. clypeal in the male. It is normally bicolored, tubercles Host fungi.-Polyporus versicoloroo; Lenzftes bstulinaO; Poly- with a reddish pronotum and black or brown elytra. In porus adustus'; Schizophyllum commune. collections, it may be confused with bicolored specimens of SutwciS curhclus, but that species differs in Discussion.-Cis vilulus may be distinguished by the having single and confused elytral punctation, narrow fairly large and robust form, single and confused elytral pronotal margins, and spinose protibial apices (see also punctation, vestiture of moderately long bristles, broad generic key). lateral pronotal margins with a raised lip, produced and Cis uersicoIot is a common inhabitant of Polyporus rounded anterior pronotal angles, and strongly tumid versicolor and its relatives which usually occur on dead prosternum. The male is characterized by having two hardwoods. 18 Bulletin of the California Insect Survey stout pronotal horns, an elevated, trisinuate fronto- It appears to be distributed throughout the range of D. clypeal ridge, and no abdominal fovea. Cis fuscipes ap- manitoba, but it is much less common. proaches it in size and shares the same habitat but differs in having dual and subseriate elytral punctation. Dolichocis manitoba Dury Cis tetracentrum differs by the characters given in the key and is unlikely to be collected in the same areas. (Fig. 32) Dolichocis manitoba Dury, 1919: 158. Holotype, 0, Aweme, Genus Dolichocis Dury Manitoba ( CMNH ). This is a small group of species placed in the genus Geographic range.-Northern and montane regions of North Enneurthron by European workers (Lohse, 1967) but America, from the northern coast of British Columbia to New differing from the E. muturn (Gyllenhal), by Brunswick and south to the central California coast, the southern type, Sierra Nevada, southeastern Utah, and northern Pennsylvania the narrower body form and simple protibial apices. At (map 10). least two Old World species are included, as well as the two North American forms, both of which occur in CaZifmia rec~rds.-coastaiTransition Zone situations and in California (Lawrence, 1965; 1971). the Coast Ranges and Sierra Nevada from the foothills to above 6,000 feet elevation. KEY TO THE CALIFORNIA SPECIES OF DOLICHOCIS Host fungi.-Pomes pfnimZd*; Polypom betulinus'; Polv- POTUS par game nu^; Fmsrmfiosup'; Daednlsa ~nioolor';F-s 1. Pronotal punctation coarser and denser, the punctures officincrlis'; Gonodetma op~lanatum'; PO~~~OTUStdphW-*; more than 0.29 times as large as scutellar base and Fomes f0mSfim-w;Polyporus ad&w; Polyponu tulipifefae; usually separated by less than 0.33 diameter; elytral Po* uolootur. punctures distinctly larger than pronotal punctures; elytral bristles 3 to 4 times as long as wide and about Discussion. - Dolichocis manitoh may be distin- 0.33 times as long as scutellar base; vertex of male with median, raised, pubescent fovea; abdominal fovea guished by its elongate, cylindrical form, 9-segmented of male margined and located in center of sternite HI. .. . antennae, rounded protibial apex, anteriorly constricted D. doba pronotum, coarse and dense punctation, which is single bnotal punctation her and sparser, the punctures less and confused, and vestiture of moderately long bristles. than 0.25 times as large as scutellar base and usually In the male, the frontoclypeal ridge bears two separated by more than 0.33 diameter; elytral and sharp pronotal punctures subequal in size; elytral bristles tubercles and the vertex has a median, raised, pube- 2 to 3 times as long as wide and about 0.17 times as scent fovea. D. indistinctus differs in having finer and long as scutellar base; vertex of male simple; abdom- sparser punctation, much shorter and stouter bristles, inal fovea of male not margined and located anterad and no fovea on the head of the male. Sulcacis curtuhs of center...... D. indistinctus resembles this species in having a cylindrical form and Dolichocis idistinctus Hatch coarse, dense punctation, but the antennae are 10- segmented and the protibial apices spinose. Dolichocis indistinctus Hatch, 1962:234. Holotype, 8, Stanley, Dolichocis rnanitoba is common and widespread British Columbia ( CAS ) . throughout the coniferous forests of western North America and is less common in the northeastern part of Geographic rrmge.-Knoum from scattered localities through- its out the northem and montane parts of North America, from the continent. In California, preferred host fungus British Columbia to the Gasp15 Peninsula of Quebeo and south is Fomes pinicola. into the Sierra Nevada, Chiricahua Mountains of southeastern Arizona, and the Green Mountains of Vermont. Genus Orthocis Casey Cdifornia records.-Dm NORTE Co.: Crescent City. MENDO- CINO CO.: Caspar. LARIPOSACo.: 6 mi. NW Fish Camp. This genus, considered a subgenus of Cis by Lohse includes 38 described species and many more Host fungi.-Fomes pinicola'; Trametes setialis6; Fomes of- (1967), ficinalis. undescribed forms from various parts of the world (Lawrence, 1965; 1971). Unlike most Cis, members of Discusslon.-This species differs from D. manitoba in the genus Orthocis always lack the teeth, tubercles, having finer and sparser punctation, shorter, stouter horns, or other armature on the head and pronotum of elytral bristles, and na fovea on the vertex in the male. the male. The single California species is easily recog- Lawrence: The Ciid Beetles of California 19

wrlrh *, - *- ,. I? I" ,,. I"

Map 10. California distribution of Dolichocis manitoba Dury. Map 11. California distribution of Hadraule bhisdelli (Casey). Inset: Distribution in North America. Inset: Distribution in North America (isolated dots are probably all herbarium records ) . nized by its large size (TL usually more than 1.75 mm) the body form is elongate, the lateral pronotal margins and characteristic form and vestiture (see below). broad and subparallel, the prosternum slightly tumid, the protibial apex narrowly rounded, the elytral punc- Ortbcis punctatus ( Melli6) tation single and confused, and the vestiture consists of very short, fine hairs. In addition, the elytral suture has (Fig. 31 ) an inflexed margin near the apex and the male bears no tubercles or horns on the head or pronotum. It is easily Cis punetatus Mellik, 1848: 337. Holotype, 8, North America distinguished other members of the family and is (MNHN). from more likely to be confused with Eupisenus elongatus Syronyrn.--Orthocis atemima by.1898: 84. (LeConte) or Sphindocis denticoUis Fall, which are presently included in the family Tetratomidae (see Geographic range.-Widespread across North America from Addendum). the edge of the Beaufort Sea (Mackenzie Wct, Northwest Territory) to the Island of Newfoundland, south on the Pacific Orthocis punctatus occurs throughout the northern Coast to Santa Barbara, California, through the Rocky Moun- and montane regions of North America, where it prob- tains to northeastern New Mexico, into the Black Hills of South ably feeds on the mycelium and fruiting bodies of Dakota, and through the eastern and midwestern states from Auricularia auricula and its relatives. In California, it New England to Florida and west to Kansas and Texas. has been collected only along the northern coast, and it California records.--AL~mA Co.: Berkeley; Oakland Hills. does not appear to be common. MARIN Co.: Cypress Ridge; Fairfax; Muir Woods. MONTEREY Co.: Cannel; Monterey. SAN MATEO Co.: Crystal Lake. SANTA Genus Hadraule Thomson BARBARACo.: Santa Barbara. SANTA CLARA Co.: Stanford This genus includes a few very small, flattened University. SONOMACo.: Sobre Vista. species in which the prosternum is much longer than its Host fungi.-Auricularia auriculao. intercoxal process (fig. 17). In addition to H. blaisdelli, Discussion.-This is the only California ciid in which there are two species in the northeastern part of North 20 Bulletin of the California Znsect Suruey America, one of which also occurs in the Palearctic similar to that in Ceracis, while the protibial apex re- region (see Lawrence, 1971). Casey included blaisdelli sembles that found in Cis and Ennearthron. Plesiocis in a separate genus, Maphoca, on the basis of the appar- cribrum is not obviously related to any New World or ently 2-segmented antennal club, but the reduction of European species, and its closest relatives may occur in the first club segment appears to be the only significant the Oriental Region. difference between Casey's species and the type of Hudraule, H.elongdula (Gyllenhal) . Plesiocis cribrum Casey

Hadraule bluisdelli (Casey) (Fig. 34) Plesiocls cribrum Casey, 1898: Holotype, $, Mokelumne (Fig. 33) 87. Hill, Calaveras County, California (USNM).

Maphoca blakrdelli Casey, 1900: 165. Holotype, 0,Mokelumne Geographic range.-Northern and montane regions of North Hill, Calaveras County, California ( USNM). America from British Columbia to the Gasp6 Peninsula of Que- bec, south to the Laguna Mountains in California, the edge of Geographic range.-Westem America, from southern North the Colorado Plateau in Arizona and New Mexico, the Great British Columbia to southern California, east through Utah, Lakes Region, and the Shenandoah Mountains of Virginia Arizona, and New Mexico into Texas, and south into Mexico. (map 12). Also recorded from Michigan, Iowa, Ohio, Massachusetts, Con- necticut, and Florida ( map 1I ). California records.-Throughout most of the Boreal Zone areas of the state, from coastal and foothill Transition Zone situations California records.-C.u.avERlls Co.: Mokelumne Hill. EL to above 7,000 feet in the central Sierra Nevada and above 8,000 DORADOCo.: 2 mi. N Placerville. KERN Co.: 8 mi. W Bakers- feet in southern California. field; Kernville. Los ANGEL= Co.: 1 mi. NE Crystal Lake; Pasadena. MAFIIN CO.: Alpine Dam; Alpine Lake. MENWCINO Host fungi.-Polyporus voluatusoa; Polyporus anceps'; Poly- Co.: Laggett; 4 mi. W Leggett; 2 mi. N Piercy. RIVERSIDECO.: pow alboluteus. 4 mi. E Blythe; Palm Springs. SAN DIEGOCo.: 1.5 mi. NW Mt. Laguna. SAN LUISOBISPO Co.: 5 mi. SE Nacimiento Dam. Dlscussion.-This species is characterized by the re SANTACLARA CO.: 2 mi. SW Los Gatos. SVASTACO.: (no specific locality). SISIUYOUCo.: (no specific locality), YOU, CO.: Davis. bust form, 9-segmented antennae, dentate protibial apex, coarse, dense, and confused punctation, and Host fungl.-lenzites saepiari4.; Po- adustus"; Trametes vestiture of short bristles. The male has 4 sharp teeth sepium; Polyporup muns;ii; Pol- uersicolor; DaedoEea con- on the frontoclypeal ridge and 2 weak protuberances fragosa'; Fames imnosusa; PoryponrP cmoep~.; Polyporut bi- formis'; Trametes hispido'; Fomes robiniae. on the apex of the pronoturn. Cis vitulus may be confused with P. Mibrum on the basis of size, general form, Discussion,-This species is easily distinguished from and the sexual modifications, but individuals of the other California Ciidae by the very small size, elongate former are distinguished by the 10-segmented anten- and flattened form, 9-segmented antennae with a 2- nae, finer punctation, especially on the pronotum, segmented club, seriate elytra1 punctation, and by the broader lateral pronotal margins, and lack of an ab- structure of the prothorax (see generic key). The color dominal fovea in the male. In Dolichocis manitoba the is similar to that of Cis versicolor, with a reddish pro- antennae are 9-segmented and the punctation is similar, notum and brownish or black elytra. but that species differs in the more elongate form, Hadrab blaisdelli occurs naturally throughout the rounded protibial apices, and modifications of the head Pacific Coast and southwestern states, but it has also in the male. been recorded as a herbarium pest in various parts of the country. In California, it is known from scattered Plesiocis cribrum is very common throughout west- localities along the coast, in the Sierra foothills, and in ern North America wherever conifers occur and is also both desert and montane portions of southern Cali- known from the northeastern states. Although the pre- fornia. ferred host is Polyporus volvatus, the species also breeds in Polyponrs anceps-the two fungi often grow Genus Pk.viocis Casey together on dead conifer logs.

This genus contains the single species P. cribrum, Genus Ceracis Melli6 Hatch's P. spenceri having been transferred to Ennear- thron (Lawrence, 1971). The prosternal structure is This genus includes 42 described species, mostly Lauwence: TheCiid Beetles of California 21

1 uw- =I Y * ,. “I t. 36s t”

Map 12. California distribution of Plssiods cribrum Casey. Inset: Map 13. California distribution of Cerocis californfcus (Casey ). Distribution in North America. Inset: Distribution in North America. from the New World. Of these, u) occur in America Cera& californicus (Casey) north of Mexico (Lawrence, species 1!367b; 1971). Two (Fig. 35) occur in CaUfomii, one generally distributed and the other known only from the vicinity of the Colorado Ennearthron californicum Casey, 1884:36.Holotype, 8, Califor- River, A third species, C. sirnilis Horn, is known from nia (USNM). islands in the Gulf of California and may occur in the Synonyms.-Ennearthron conuergens Casey, 1898: 89. Ennear- state. thron discolor Casey, 1898: 89. Ennearthron grossulum Casey, 1898: 89. Ennearthron coloredense Dury, 1917: 22. Ennearthron KEY TO THE CALIFORNIA SPECIES OF CERACIS oregonus Dury, 1917: 22.

1. Antennae %segmented; pronotal punctation coarser and Geographic range. - Western North America, from Seattle, denser, punctures usually more than 030 times as Washington, east to western Nebraska, south along the Pacific large as scutellar base and separated by less than 1.5 Coast to southern Cabfomia and through the Great Basin and diameters; occurring throughout state...... Rocky Mountain Regions to southern Arizona and New Mexico; c. calrfonricus (p. 21 ) extending into Mexico as far as Baja California Sur and southern Antennae %segmented; pmdpunctures usdy less Sindoa (map 13 1. than 1.5 times as large as scutellar base and separated by more than 1.5 diameters; extreme southern part California records. - Widespread in cismontane area of the ofstate ...... 2 state at low and intermediate elevations, ranging up to 2. Body smaller and narrower, TL usually less than 1.30 feet in southern California. mm. and EWEW usually more than 1.40; elytra uniformly black; apex of prowtum in male simple or Host fungi, - Porrrpofiu eho;PO- OdUFtUS.; weakly emarginate; abdominal fovea in male trans- Crmoderma bnnonii8;Ganoderma sp.*; Lenzites bstulina;Gano- versely oval ...... c. diriensis (p. 22) &m applanutum*; Tmmetes hispida*; Pleurotw ostreatUc*; Body larger and broader, TL usdymom than 1.30 mm. pa&w0; steccherinm ochraceum*; Polyporus &WS; and EWEW usually less than 1.40; elytra usually red- GaMdefiM Iucidumo; GmMdsrmo lobaturn*; Polyponrs par- dish in part; apex of pronotum in male bearing an gamenuso; Daedulea unioolor*; Fomes -*; Foms~*- emarginate lamina; abdominal fovea in male circular.. .. nophi2u.P; canodetma oregoneme; Fames μ Trametes c. similis (p. 22) mollfs. 22 Bulletin of the California Znsect Survey Discussion. - Cera& californicus may be distin- Genus Sulcacis Dury guished by the elongate, cylindrical form, 9-segmented antennae, concave prosternum with a laminate inter- Sukacb contains four Palearctic species and two coxal process, spinose protibial apices, coarse, dense North American forms, one of which extends into Cali- punctation, and vestiture of short, fine hairs. The other fornia. The spinose protibial apices are similar to those two Ceracis which might coexist with this species in the of Cerucis species, while the structure of the proster- extreme southern part of the state both have 8-seg- num and the type of vestiture are more like that in the mented antennae and finer punctation. The only other genus Makacocis. elongate and subglabrous ciid in the California fauna is Orthocis wnctutus, which dif€ers in having 10- segmented antennae, rounded protibial apices, a slightly tumid prosternum with a broader intercoxal process, and a flatter form (see also generic key). In California Ceracis californicus breeds in a variety of fungi asso- Cis curtuh Casey, 1898: 83. Holotype, 8, New York (USNM). ciated with hardwoods. Synonyms.-Cis montaw Casey, 1898: 82. Cis SOTTOT Casey, 1898: 83. Cis c&dricus Dury, 1917: 8. Sulcacis niger Dury, Ceracis didensis (Tanner) 1917: 21. Cis criddlsi Dwy, 1919: 158. Geographic range. - Widespread through the northern and Octotemnus didensis Tanner, 1934: 57. Holotype, 8, Zion Na- montane regions of North America, from the northern mast of tional Park, Utah (BYU). British Columbia to southern Quebec and New England, south to San Diego County, California, the mountains of southem Geographic range.-Southwestern North America, from south- Arizona and northern Mexico, and the states of Nebraska, Illinois, em Utah to Guadalajara, Mexico, and from Colorado River in and North Carolina ( map 14 ) . California to the Big Bend Region in Texas. California records. - Common and widespread throughout California record.-RrclERsrm Co.: 4 mi. E Blythe. much of the state except in the more arid regions east of the Sierra Nevada and in the deserts. This species occurs in Upper Host fungi.-Trametes hispidaoo; Ganoderma sp. Sonoran and Transition Zone situations at low elevations, ranghg up to moderate elevations (5,oocrS,000 feet) in the mounbins Discussion. - This species is characterized by the of both central and southern California. small size, &segmented antennae, and he pronotal punctation. It is usually found breeding in the fruiting Hart fungi. - Po2yponu uersiwloro0; PoIypmus hisdo; bodies of Trametes hfspidu growing on cottonwoods PO- 4duptUpO; Lenzites betho; Trmnetes heo; Pleumtus ostreotus; Powcinnabarinus*; Fomes ignfaiu9; along dry river beds. Schizaphyllwn commune*; Gonodemur applrmcrtum; Polypmur gilow; Polvpotus uulpinus; Steccherinum ochracsum; Stsrercm Ceracis sirnilis Horn hirsutum.

Ceracis simflis Horn, 1894: 391. Syntypes, Coral de Piedra, Discussion.-This species may be distinguished by Sierra el Taste, Baja California Sur (CAS). the subcylindrical form, 10-segmented antennae, spinose protibial apex, biconcave prosternum with a fairly Geogmphtc mnge.-Baja California and Nayarit, Mexico, broad, tapering intercoxal process, coarse, dense, can- south to El Salvador. fused punctation, and vestiture of yellowish bristles California rew&.-None. To be expected in extreme southern which vary somewhat in size. Species of Cera& have California much her vestiture, 8- or 9-segmented antennae, and laminate intercoxal process, while in Dolichocds the Host fungi.-GanodsmM zonatumo; Ganodemur sp.O; &no- protibial apex is rounded, the prosternum is slightly demapplanatumO . tumid, and the antennae are 9-segmented. Cis hystri- Discussion.-This species is fairly common in Baja culus differs in having the protibial apex distinctly den- California and on various islands in the Gulf of Califor- tate and the prosternum tumid. nia; its occurrence in the state is doubtful. It differs Although SUMcurtulus occurs at fairly high alti- from C. CaZifmicus in having &segmented antennae tudes in the Sirma Nevada, it is not usually associated and a somewhat shorter body form, and fiom C. &- with conifers but rather is an inhabitant of Polyporus ensis in the larger size, reddish co&%ion, and different uersicobr and various other fungi growing on hard- sexual modifications. woods. Lawrence: The Ciid Beetles of California 23

v ." m ," L" w ," tn u

Map 14. California distribution of Sulcacts curtulus (Casey). Map 15. California distribution of Octotemnus his Casey. In- Inset: Distribution in North America. set: Distxibution in North America.

Genus Octotemnus Mellik ville; 3 mi. N Placedle. HUMBO~Co.: Eureka; Garberville; 8 mi. S Korbel. MAFUN Co.: Alpine Lake; Inverness; 1 mi. SE Octotmus is primarily an Old World genus, with Inverness; Lagunitas; Ma Vdey; Muir Woods; Redwood Can- only one American occurring throughout yon; S. P. Taylor State Park; Taylorville. MKNDOCP~OCo.: Van North species, Damme State Park MONTEREYCo.: Big Sw, Carmel. PLVMhs the northern part of the continent. It is easily distin- Co.: 7 mi. NE Buck's Lake. SAN MATEO Co.: Marilama Point. guished from most American ciids by the tribal char- SANTA CLARACo.: Los Gabs; 2 mi. SW Los Gab; San Jose; acters given in the key on page 9. The only other North 2 mi. E Sveadal. SANTACRUZ Co.: Big Basin; Ben hmond; American orophiine is Rhopcrlodontus americanus Redwood Glen; Santa Cruz Mountains. SISKIYOUCo.: Dunsmuir; Weed. SONOMA Co.: hcan Ma;Glen Ekn. TaaRg w.: Lawrence from Wisconsin ( Lawrence, 1971). Dorset Camp, Sequoia National Park.

Octotemnus hisCasey Hod fungi. - Polwwur vedt~lor~~;Pofypmur himrhrs'; Polflpmpubsscensf; Lenzites betditw.; Polyporup eonchveP; ( Fig. 37) kodenna ap&matum0; PoryporuS abietinw; Pdgponrs adw tu; Polyponu aZbeUw; Polgpom gatnctimcs; PolIlpwur par- ganaenus; Polyporw Bqwmosus; Polyporus sulphureus; Stereum Octotemnus loevkr Casey, 1898: 91. Holotype. 8, %ode Island (USNM). sp.

syonym.-obotemmls denudatw casey, 1898: 91. Discussion.-- species is characterized by its oval body form, &segmented antennae, spinose tibiae, finely Geographic range.-Widespread throughout the northern part punctate, almost glabrous surface, and by other fea- of North America, from southern Alaska to Quebec and Nova tures mentioned in the generic key. It appears to prefer Scotia, south along the Pacific Coast to Monterey County, Cali- fornia, in the Sierra Nevada to Tulare County, in the Midwest humid areas where it is a common inhabitant of Poly- to southern Iowa and Kansas, and on the East Coast to Alabama pmvetsicolor and its relatives. (map IS). ADDENDUM Calffornfa wcor&--ALmmA Co.: Berkeley; Oakland; Straw- berry Canyon. CONTRACOSTA Co.: Tilden Park, Berkeley Hills. The monotypic genus Sphindocis Fall was removed DELNORTE Co.: Pahick Creek. EL DORAWCo.: 2 mi. N Placer- from the Ciidae (Lawrence, 1971) and tentatively 24 Bulletin of the California Insect Survey placed in the Tetratomidae. Further studies of this and sided), but differs from that and all other ciid species related heteromerous Coleptera since the completion of by the following characters: (1) antennae 11-segment- this manuscript have convinced me that Sphindocis ed, without large sensory organs on the club; (2) tibi- must be returned to the family Ciidae and should be inal spur present; (3) abdominal sternites I11 and IV con- cluded in this paper, since it represents a subfamily en- nate; (4) lateral margins of pronotum undulate, form- demic to California. Although most of the resem- ing several weak teeth; (5)trochanters strongly hetero- blances between Sphituhcis and the Ciidae are prob- meroid (obliquely attached to femur so that part of ably primitive with the Heteromera, at least one, the femoral base is attached directly to coxa); (6) basal distinct laciniar lobe of the larval maxilla, appears piece of aedeagus almost as long as tegmen, with two to be derived and argues for a common ancester apical condyles; median lobe membranous with two of the two groups, which is not at the same time the an- lateral struts; and (7) metendosternite with distinct cestor of any other heteromeran. Since the genus dif- median stalk. Sphindocis larvae differ from those of fers from all other ciids with respect to several funda- other ciids by having 3-segmented antennae, biforous mental characters (listed in Lawrence, 1971: 427, table spiracles, and a row of asperites on ventrite IX. Other 1), a new subfamily is being proposed (Lawrence, in characters are discussed by Lawrence (1971 and in press) for its inclusion; the remaining ciids (Ciini and press). Sphindocis denticollis also resembles the pise- Orophiini) comprise the subfamily Ciinae. nine tetratomid Eupisenus elongcrtus (Monte), but it differs from that species in having only four tarsal seg- Subfaimily Sphindociinae Lawrence ments on all legs, the first two visible abdomnal ster- Genus Sphindocis Fall nites connate, a completely hidden prothoracic troch- Sphindocis deraticouis Fall antin, and a fovea on the first visible abdominal sternite of the male. Sphindocis denticoufs Fall, 1917: 171. Holotype, 8, Alameda Co., California ( MCZ). This species is most commonly found in the fruiting bodies of Trametes sepium growing on dead madrone Geographic range.-Coastal California from northern Mendo- (Arbutus Menziesii) branches lying on the ground. cino County to Alameda and San Mateo Counties. One series of larvae was collected on Poria cinerascens a ( , California records.-ALaMEm Co.: no specific locality. MARIN growing on Douglas fir log Pseudotsuga Menziesii) CO.: Alpine Lake; Mill Valley; Mt. Tamalpais. MEND~CINOCO.: but none could be reared throdugh to the adult stage. Fort Bragg; 4 mi. E Fort Bragg; 3 mi. S Leggett; 4 mi. W Leg- A single pupa was cut out of madrone wood beneath a gee 1 mi. N Piercy; 2 mi. N Percy; 4 mi. W Willits. SAN MATEO T. sepium fruiting body, and it may be that pupation Co.: Corte Madera Creek; Kings Mtn. Road, Woodside. takes place only within the wood. Sphindocis denticollis is a very distinct and probably Host fungL-Trametes sepiumO; Po& cinermcens*. relict form which appears to be endemic to the mixed Discussion.-This species resembles a large Orthocis Transistion Zone forests of the northern California pnctatus (34 mm. long, subglabrous, and parallel- coast. LITERATURE CITED

Bomc, A. G. and F. C. CRAXCHFNI FALL,H. C. 1931. An illustrated synopsis of the principal larval forms of 1917. New Coleoptera. VI. Can Entomol. 49: 1S171. the order Coleoptera. [part]. Entomol. Am., N. S., GORHAM,H. S. ll(4): 258351, PIS. 86-125. 1886. Supplement to Malacodermata. In F. D. Godman and BROUN,T. 0. Salvin, eds., Biologia Centrali-Americana. Insecta. 1880. Manual of the New Zealand Coleoptera. Part I. Well- Coleoptera. Vol. 3, Part 2. London: Porter. Pp. 313- ington: N. Z. Mus. Geol. Sw.Dep. 652 pp. 360, pl 13, figs. 21-24. CASEY,T. L. 1898. On the serricorn Coleoptera of St Vincent, Grenada, 1884. Contributions to the Descriptive and Systematic Cole- and the Grenadines (Maladermata, Ptinidae. Bostry- opterology of North America. Part I. Philadelphia, 80 chidae), with descriptions of new species. Proc. Zool. PP. SOC. London, 1898: 315-343, pL 27. 1898. Studies in the Ptinidae, Cioidae, and Sphindidae of G-, J. America. J. N. Y. Entomol. Soc., 6: 61-93. 1935. A revised life-zone map of California. Univ. Calif. 1900. Review of the American Corylophidae, Cryptophagidae, Publ. ZooL, 40: 327330, 1 map. Tritomidae and Dermestidae, with other studies. J. HALL,H. M. AND J. GIUNNBLL N. Y. Entomol. Soc., 8: 51-72. 1919. Life zone indicators in California. Proc. Calif. Acad. Sd., ser. 4,Q: 37-67. &OWSON, R. A. H. 1960. The phylogeny of Coleoptera. Annu. Rev. Entomol. HATCH,M. 1962. The beetles of the Padc Northwest. Part 111: 5: 111-134. Pselaphidae Diversimmia 1. Seattle: 1966. Observations on the constitution and subfamilies of and Univ. Wash- ington Press, ix + 503 pp., 66 pls. the family Melandryidae, Em. 41: 507-513. HORN,G. H. DALLATORRE, K. W.VON 1894. The Coleoptera of Baja California. Proc. Calif. Acad. 1911. Cioidae. Pars 30. In W. Junk and S. Schnkling, eds.. Sci., ser. 2,4: 302449. Coleoptmrum Catalogus. Berlin: Junk, 32 pp. Hum, P. D. AND C. D. MICFLENW DURY,C. 1955. The megachiline bees of California (Hymenoptera: 1917. Synopsis of the coleopterous family Cisidae (Cioidae) Megachilidae). Bull. Calif. Insect Sw., 3: 1-247. of America north of Mexico. J. Cincinnati SOC, Nat. KENDnmr, s. c. Hist., 22 ( 2 ) : 1-27. 1932. A study of Meniam’s temperature laws. Wilson Bull., 1919. A new ciside genus with new species from Manitoba. 44: 129-143. Can. Entomol. 51: 158. LAWRENCE,J. F. FABRICIUS,J. C. 1965. Comments on some recent changes in the classification 1792. Entomologia Systematica Emendata et Aucta. Tomus I of the Ciidae (Coleoptera). Bull. Mus. Comp. Zool., [Pars 11. Copenhagen: Prost, xx + 33Opp. 133; 273-203. 26 Bulletin of the California lnsect Survey 1967~.Biology of the parthenogenetic fungus beetle Ckr Md,J. fuscipes A4eW (Coleoptera: Ciidae). Breviora, No. 1848. Monograhie de I?ancien genre Cis des auteurs. Ann. 258,14 pp. Soc. Entomol. Fr., Ser. 2, 6: 205-274, 313-396, pls. 1967b. Delimitation of the genus Cemcis (Coleoptera: Ciidae) 9-12. with a revision of North American species. Bull. Mus. MEFIRIAM,C. H. Comp. Zool., 138: 91-144. 1892. The geographic distribution of life in NdAmerica 1971. Revision of the North American Ciidae (Coleoptera). with special reference to the Mammalia. Proc. Biol. Bull. Mus. Comp. Zoo]., 142: 419522. SOC. Wash., 7: 1-64. 1973. Host preference in ciid beetles (Coleoptera: Ciidae) 1898. Life zones and crop zones of the United States. U. S. inhabiting fruiting bodies of Basidiomycetes in North Dep. Agric. Div. Biol. Surv. Bun. 10: 1-79. America. Bull. Mus. Comp. Zool., 145: 163-212. MILLER,A. H. In 1951. An analysis of the distribution of the birds of California. Press The larva of Sphinidocis denticollk Fall and a new sub- Univ. Calif. Publ. Zool., 50(8): 53144. family of Ciidae (Coleoptera: Heteromera). Breviora. MUNROE,E. LAWRENCE,J. F., AND J. A. POWELL 1956.?Canada as an environment for insect life. Can. En- 1969. Host relationships in North American fungus-feeding tomol., 88: 372476. moths ( Oecophoridae, Oinophilidae, Tineidae) . Bull. OVERHOLTS,L. C. Mus. Comp. Zool., 138: 29-51. 1953. The Polyporaceae of the United States and Canada. LINSLEY,E. C. Univ. Mich. Stud., Sci. Ser., Vol. 19, xiv 4- 466 pp., 1958. Geographical origins and phylogenetic affinities of the 132 pls. cerambycid beetle fauna of Western North America, PAVIOUR-SMITH,K. pp. 299420, in C. L. Hubbs, ed., Zoogeography. Am. 1960. The fruiting bodies of macrofungi as habitats for Assoc. Adv. Sci. Publ. No. 51, x + 509 pp. beetles of the family Ciidae (Coleoptera). Oikos, 11: LOHSE, G. A. 43-71. 1967. Familie: Cisidae, pp. 280-295, in H. Freude, K. W. 1969. An attempt to correct some mistakes and misconcep- Harde, and G. A. Lohse, Die Kafer Mitteleuropas. Band tions about some fungus beetles and their habitats. 7. Krefeld: Goecke and Evers, 310 pp. Entomologist, April:86-96. LOWE,J. L. AND R. L. CJLBERTSON PETERSON,A. 1961~.Synopsis of the Polyporaceae of the southeasternUnited 1957. Larvae of Insects. An Introduction to Nearctic Species. States. J. Elisha Mitchell Sci. Soc., 77:43-61. Part II. Coleoptera, Diptera, Neuroptera, Siphonaptera, 1961b.Synopsis of the Polyporaceae of the western United Mecoptera, Trichoptera. Columbus, Ohio: Author, v States and Canada. Mycologiq 53: 474511. i-416 pp. MANNERHEIM,C. C. VON POWELL,J. A. 1843. Beitrag zur Kaefer-Fauna der aleutischen Inseln, der 1964. Biological and taxonomic studies on tortridne moths, Insel Sitkha und Neu-Califomiens. Bull. SOC.Imp. Nat. with reference to the species in California. Univ. Calif. MOSCOU,l6( 2) : 175-314. Publ. Entomol., 32:1317. 1852. Zweiter Nachtrag zur Kaefer-Fauna der nord-ameri- SXELFORD,V. E. kanischen Laender des russischen Reiches. Bull. SOC. 1932. Life zones, modern ecology, and the failure of temp- Imp. Nat. Moscou, 25( 2) : 283-387. erature summing. Wilson Bull., 44: 144157. 1853. Dritter Nachtrag zur Kaefer-Fanna der nord-ameri- TANNER,V. M. kanischen Laender des Russischen Reiches. Bull. SOC. 1934. Coleoptera of Zion National Park, No. 2. Ann. Entomol. Imp. Nat. Moscou. 26( 3): 95-273. Soc. Am., 27:4349. FIGURES 28 Bulletin of the California Insect Survey

Figs. 1-2. Octotemnus hisCasey, 8,prothorax with right coxa removed (scale = 0.125 mm) . Fig. 1. Ventral view (arrow pointing to tip of intercoxal process). Fig. 2. Anterior view.

Figs 3-4. Cis fuscipes Me&&, 0 , prothorax (scale = 0.250 mm). Fig. 3. Ventral view with both coxae removed. Fig. 4. Anterior view with right coxa removed.

Fig. 5. Octotemnus laeois Casey, 8 , abdomen, ventral view ( showing covered abdominal fovea) (scale = 0.135 mm).

Fig. 6. Cis tetracentrum Gorham, 8,abdomen, ventral view (showing exposed abdominal fovea) (scale = 0.250 mm). Lawrence: The Ciid Beetles of California 29

2 30 Bulktin of the California Insect Surocly

Figs. 7-15. Right tibia of male, posterior view (all drawn to same scale). Fig. 7. Ordhocis punc- tutus ( MelliB). Fig. 8. Pkdocis cribrum Casey. Fig. 9. Cis wssicobr Casey. Fig. 10. Cis striokztus Casey. Fig. 11. CCr vitulur Mannerheim. Fig. 12. Dolichocis munitoba D~uy.Fig. 13. Cerucis CaUfmicus (Casey). Fig. 14. Sulcocfs curtulus ( Casey). Fig. 15. Octotemnur he& Casey.

Fig. 16. mho& punctatw (Mellib), male, elytra1 apices, postero-dorsal view.

Fig. 17. HudrauZe blak&Zli (Casey ), male prothorax, ventral view with both coxae removed (scale = 0.125 mm).

Figs. 18-19. Anterior angle and lateral margin of pronotum, dorsal view (scale = 0.063 mm). Fig. 18. C@ americanus Mannerheim. Fig. 19. Cis fuwipes MelliB. Lawrence: The Ciid Beetles of Califmkz 31

9 10 9 8 8

12 1 13

14 32 Bulletin of the California Insect Survey

Fig. 20. Cis amerfcanw Mannerheim, 8.

Fig. 21. Cts biarmatus Mannerheim, $.

Fig. 22. Cis dupk Casey, 8.

Fig. 23. Cis crebewimus Mellib, 8. Lawrence: TheCiid Beetles of California 33

23 34 Bulletin of .the California Insect Survey

Fig. 24. Cts fusdpa MelliB, 8.

Fig. 25. Cis fuscipes MelliB, 0.

Fig. 26. Cis hystriculw Casey, $.

Fig. 27. Cis maritimus (Hatch), 8. Lawrence: The Cifd Beetles of California 35 36 Bulletin of the. California Insect Survey

Fig. 28. Cis tetracentm Gorham, 8. Fig. 29. Cis vsrsicobr Casey, 8. Fig. 30. Cis uitulw Mannerheim, 0. Fig. 31. Orthocis punctatus (Mellih), 8, Lawrence: The Cad Beetles of California 37 38 Bulletin of the California Insect Survey

Fig. 32. Dolichocis rnanitoba Dury, 8.

Fig. 33. Hadraule blokdelli ( Casey 1, 8.

Fig. 34. Pksiocis cribrum Casey, 8.

Fig. 35. Cera& califonticur (Casey), 8. Lawrence: The Ciid Beetles of California 39

34 40 Bulletin of the California Insect Survey

Fig. 36. Sukacts curtulus (Casey), $.

Fig. 37. Octotemnus Eaeois Casey, 8. Lawrence: The Ciid Beetles of California 41