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9-1997

Revision of the Genus Malporus Casey (Coleoptera: : Anthicinae)

Donald S. Chandler University of New Hampshire, [email protected]

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Recommended Citation Revision of the Genus Malporus Casey (Coleoptera: Anthicidae: Anthicinae) Donald S. Chandler The Coleopterists Bulletin Vol. 51, No. 3 (Sep., 1997) , pp. 265-275 Published by: The Coleopterists Society Stable URL: http://www.jstor.org/stable/4009420

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Revision of the Genus Malporus Casey (Coleoptera: Anthicidae: Anthicinae) Author(s): Donald S. Chandler Source: The Coleopterists Bulletin, Vol. 51, No. 3 (Sep., 1997), pp. 265-275 Published by: The Coleopterists Society Stable URL: http://www.jstor.org/stable/4009420 . Accessed: 17/02/2015 14:19

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REVISION OF THE GENUS MALPORUS CASEY (COLEOPTERA: ANTHICIDAE: ANTHICINAE)1

DONALD S. CHANDLER Department of Zoology University of New Hampshire Durham, NH 03824

Abstract The North American genus MalporusCasey is revised for the first time. Three species are found in the eastern North America, and are most commonly collected in riparian areas or in salt marshes: M. cinctus (Say), M. formicarius (LaFert6-Senectere), M. pro- perus Casey. A new species from the Southwestern United States, M. werneri, is de- scribed, and M. blandus Casey is here placed as a junior synonym of M. properus, new synonym. Anthicusfornicarius LaFert6-Senectere is designated as the type species of Malporus.

Malporus is a small, distinctive group of Anthicidae from North America, which was proposed as a new genus by Casey (1895) in his revision of the North American fauna. Casey placed four species into this group: formicarius LaFerte-Senectere, A. cinctus Say, and two species he then de- scribed, Malporus properus Casey and M. blandus Casey. The only subsequent systematic comment on this group was by Werner (1975), when he placed M. blandus as a junior synonym of M. formicarius. This is one of the few North American genera of Anthicinae that has not been revised within the last 35 years. While Malporus is not a diverse group, there have been consistent misidentifications of the few species in almost all collections, and a new species from Arizona and New Mexico has been found. This revision will clarify the species characteristics, correct the one synonymy, and provide an opportunity to summarize the greatly increased information on distribution of these species. Lengths cited in the species descriptions are based on measurement of the examined specimens of the respective species. The other measurements are based on an exemplar selected for each species.

Biological and Collection Information LeConte (1852) was the first to provide information on the habitat associ- ations of this group, when he mentioned that M. formicarius was common in salt marshes. Malporus formicarius is frequent in New England and eastern Canada under debris in sandy areas along lakes and streams, or in coastal dunes and along the margins of salt marshes. Blatchley (1910) later noted that M. cinctus was found beneath logs and "rubbish" in sandy areas near water. Members of this genus are usually associated with riparian or marshy areas. When individuals are collected at ultraviolet light, the light is usually by a stream or river. Adults are probably scavengers and micro-predators like most Anthicidae (Werner and Chandler 1995), as are the larvae. Kitayama (1982)

' Scientific Contribution Number 1949 from the New Hampshire Agricultural Experiment Station

265

This content downloaded from 132.177.228.65 on Tue, 17 Feb 2015 14:19:04 PM All use subject to JSTOR Terms and Conditions 266 THE COLEOPTERISTS BULLETIN 51(3), 1997 provides the only information on the immatures of this genus with his descrip- tion of the larvae of M. cinctus, which were found in Diptera puparia at a lake in Minnesota. Interestingly, the relatively rare M. properus was fairly commonly collected along the eastern seaboard until the early 1900's, but there is only one record of this species from east of the Appalachian Mountains (Cecil County, Mary- land, 1978) in the last 70 years. The other two eastern species may still be easily collected throughout their ranges, though M. cinctus is definitely more common in the Midwestern states, and M. formicarius more common in north- eastern North America. Blatchley (1910) indicated that the generic name may be translated to En- glish as "bad walker." It hasn't been apparent to me why Casey chose this generic name, if indeed this meaning was his intent, since the rather angular and ant-like walking gait of this group is similar to all other members of the Anthicinae.

Key to Species of Malporus 1 Elytral setae appressed; punctures on head disc small, much smaller than eye facets; mesotrochanters spinose in one species; metatrochanters not m odified ------2 1' Elytral setae subdecumbent, raised to about 200; punctures on head larger, as large as eye facets; males with meso- and metatrochanters spinose or angulate- 3 2 Tactile setae on elytra at average angle of about 450; head vertex densely punctate and microreticulate between eyes; males with meso- and metatro- chanters curved on posterior margins; aedeagus with deep double incision at apex (Fig. 4) -Malporus werneri Chandler 2' Tactile setae on elytra at average angle of about 750; head vertex sparsely and finely punctate between eyes; males with mesotrochanters spinose; aedeagus without incision at apex, apex flared laterally and with median point (Fig. 3) -Malporus properus Casey 3 Elytral apices abruptly white; setae on head and pronotum subdecumbent; aedeagus with tegmen in lateral view nearly twice as thick at middle as near base, apices straight in lateral view (Fig. 1) ------Malporus cinctus (Say) 3' Elytral apices dark or diffusely paler; setae on head and pronotum ap- pressed; aedeagus with tegmen in lateral view nearly same depth through basal two-thirds, apices curved ventrally (Fig. 2) ------Malporus fornicarius (LaFerte-S6nectere)

Malporus Casey Malporus Casey 1895:65 1. Type species: Anthicus fornicarius LaFert6-Senec- tere, here designated. Blatchley 1910:1338. Werner 1963:749-750; 1964: 199. Uhmann 1976:174; 1978:76. Bousquet 1991:273. Anthicus (subgenus Malporus): Pic 1911:30. Leng 1920:163. Description. Head and pronotum with short erect tactile setae, setae appressed to subdecumbent and about as long as tactile setae; elytra with long erect tactile setae, much longer than appressed to subdecumbent setae. Head with large eyes, with very short setae between facets; frontoclypeal suture visible as a thin line; punctures on head densest anteriorly, with narrowly impunctate median line; head base broadly subtruncate; antennal segments elongate, apical four segments slightly broadened; maxillary palpi

This content downloaded from 132.177.228.65 on Tue, 17 Feb 2015 14:19:04 PM All use subject to JSTOR Terms and Conditions THE COLEOPTERISTS BULLETIN 51(3), 1997 267 with fourth segment broadly angulate on mesal face. Pronotum widest in apical half, distinctly narrower than head; strongly constricted laterally near base; apex with sharply defined small collar; distinct narrow transverse basal sulcus strongly angling antero- ventrally at lateral margins to lateral foveae near ventral margin of constriction. Meso- scutellum bluntly angulate; elytra with broad distinct omoplates; transverse postbasal impression distinct, usually strongly marked with whitish band across impression, setae in impression slightly to strongly angled posterolaterally; fine lines adjacent to suture visible only in apical third. Mesosternum with lateral margins straight, converging to acute anterior margin; mesepisterna with dorsal margins slightly convex, mesepimera barely visible as thin angle posterolateral to mesepistema; mesocoxae separated by pos- terior extension of mesosternum. Male genitalia with 9th stemite "Y" shaped, with articulated extensions at apices of short lateral arms; aedeagus with phallobase as long or longer than tegmen; 8th sternite broadly emarginate, slightly bisinuate. Remarks. Some members of Leptaleus LaFerte-Senectere share several of the features which characterize Malporus. In fact, if species such as Leptaleus valgipes (Marseul) of Japan and some eastern Oriental species are considered, I could not really separate the two genera. However, Leptaleus is a large Old World group based on the circum-Mediterranean type species Leptaleus rod- riguesi (Latreille), whose external form and aedeagal structure is different enough that I feel it would be inappropriate to synonymize the two genera, and I am not prepared to resolve placement of the many Japanese and Oriental Leptaleus at this time. Malporus may be readily separated from other New World genera such as Anthicus Paykull by the features of the deep lateral pronotal constriction and the presence of distinct, though low, elytral omoplates (a bump centered in the basal portion of each elytron). These features are shared with members of LaFerte-Senectere from the Neotropics and Australia, however Malporus differs in the possession of straight lateral mesosternal margins, which are strongly, laterally rounded in Acanthinus.

Malporus cinctus (Say) Figs. 2, 6 Anthicus cinctus Say 1824:278. Type locality: "United States" [types de- stroyed?]. LeConte 1852:97. LaFerte-Senectere 1847:185; 1848:86 (both as synonym of M. formicarius). Anthicus (Malporus) cinctus, Pic 1911:40. Leng 1920:163. Malporus cinctus, Casey 1895:653. Blatchley 1910:1338. Kitayama 1982:85 (larva). Bousquet 1991:273. Description. Length 4.2-4.5 mm. Head, pronotum, and elytral base orange to dark orange, rarely darker, head often diffusedly darker in apical half; yellowish band from lateral margins across transverse postbasal impression, narrowly interrupted at elytral suture; elytra dark brown from yellow band posteriorly to white-tipped apices. Setae on head and pronotum subdecumbent, angled at 25-30?. Head with fine punctures denser in apical half of vertex, about 10 in line between eyes, widely scattered in basal half, surface polished; eyes large, 0.32 mm long. Pronotal disc bearing scattered small punc- tures, basal portion of dorsum with punctures denser and coarser, extending into lateral constrictions; dorsum above lateral constrictions flat to slightly concave in lateral view. Elytra with long tactile setae erect, angled at 75-85?, 0.28 mm long; setae subdecumbent, angled at 20-25?, 0.08 mm long. Males with protibial apices truncate; meso- and me- tatrochanters broadly spinose; tegmen of aedeagus with deep paired incision at apex, outer apices straight in lateral view, tegmen in lateral view nearly twice as deep at middle as near base (Fig. 2). Distribution (Fig. 6). Specimens (95) have been seen from Canada: On-

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1 I Fig. 1. Dorsal habitus of Malporus formicarius. tario, Manitoba, Saskatchewan and the United States: Massachusetts, New York, Pennsylvania, New Jersey, Maryland, District of Columbia, Virginia, West Virginia, Ohio, Indiana, Michigan, Illinois, Wisconsin, Minnesota, Iowa, Missouri, Kansas, Arkansas, Florida, Mississippi, Louisiana, Oklahoma, Texas. Specimens have been examined from the following collections: CBB, CNCI,

v' - Figs. 2-5. Ventral view aedeagus, with lateral view of -tegmen. 2) M. cinctus; 3) M. fornicarius; 4) M. properus; 5) M. werneri.

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Fig. 6. Distribution map for Malporus cinctus.

DSC, EJF, IRCW, MCZC, EGR, TAMU, UAIC, UMIC, UMRM, USNM, UvCc. Biological and Collection Information. This species has been most com- monly collected at ultraviolet light near rivers. Two specific habitat records are from under bark of a sycamore tree, and a few larvae were found in Diptera puparia along a lakeshore (Kitayama 1982). Blatchley (1910) notes that this species is most common in sandy places near water beneath logs and rubbish. Type Designation. I could not find any of Say's original specimens in North American collections housed in the Museum of Comparative Zoology at Har- vard University. However, I agree with Mawdsley (1993) that it would be premature to designate a neotype before examining material in the European museums holding collections of workers with whom Say exchanged speci- mens. For this reason, and the fact that there has been no doubt of the identity of Say's Anthicus cinctus based on his written description, a neotype is not designated.

Malporus formicarius (LaFerte-S6nectere) Figs. 1, 3, 7 Anthicus formicarius LaFerte-Senectere 1847:86; 1848:185. LeConte 1852:97. Type locality: "Amerique boreale". Lectotype male, MNHN.

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II It

Fig. 7. Distribution map for Malporus formicarius.

Anthicus (Malporus) formicarius, Pic 1911:49. Leng 1920:163. Malporus fornicarius, Casey 1895:652. Bousquet 1991:273. Description. Length 3.4-4.1 mm. Head, pronotum, and elytral base orange to dark brown, head usually diffusely darker in apical half; straight yellowish band from lateral margins extending across transverse postbasal impression, band usually interrupted at elytral suture; elytra posterior to band dark brown, elytral apices sometimes slightly lighter. Head and pronotum with setae appressed. Head with small punctures of apical half of vertex denser, about 10 punctures in line between eyes, widely scattered in basal half, surface polished; eyes moderate in size, 0.28 mm long. Pronotal disc sparsely punctate, dorsum near base more densely and coarsely punctate; dorsum above lateral constriction straight to slightly convex in lateral view. Elytra with tactile setae erect at average angle of 800, 0.30 mm long; setae subdecumbent, average angle 20?, 0.08 mm long. Males with protibiae spinose at ventral apex; mesotrochanters bluntly angulate; metatrochanters with blunt spine; tegmen of aedeagus with deep paired incision, outer apices slightly curved ventrally in lateral view; tegmen in lateral view about same depth through basal two-thirds of length (Fig. 3). Distribution (Fig. 7). Specimens (355) examined are from Canada: Nova Scotia, New Brunswick, Quebec, Ontario, Manitoba, Saskatchewan and the United States: Maine, New Hampshire, Vermont, Massachusetts, Connecticut, Rhode Island, New York, Pennsylvania, New Jersey Delaware, Maryland, Dis- trict of Columbia, Virginia, Ohio, Indiana, Michigan, Illinois, Missouri, Kan-

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11 -- . 1 Fig. 8. Distribution map for Malporus properus.

sas, Nebraska, Iowa, Minnesota, South Dakota. There is a single record from Arizona (1 male, Pinal Mountains, Wickham, USNM), which may be a mis- labeled specimen. Specimens seen are in the following collections: CBB, CDAE, CNCI, DSC, EJF, EDUM, EGR, FSCA, IRCW, MCZC, NAUC, REN, PES, UAIC, UMIC, UMRM, UNHC, USNM, UVCC, VPIC, WSUC. Biological and Collection Information. This species is often found beneath washed up coastal and lake drift, and has been particularly common in salt marshes (LeConte 1852:97). It has also been found in gopher burrow mounds, in old beaver lodges, on willows, under grass debris, under stones or boards, and in leaf litters. Only a few specimens have been collected at ultraviolet light. Type Designations. The lectotype (female) here designated bears the labels //Amer. bor./ Ex. Musaeo Mniszech/ Museum Paris ex Coll. R. Oberthur Laf- erte/ TYPE/ Formicarius (dig) mihi Am.Bor./ LECTOTYPE Anthicus formi- carius LaFerte des. DSChandler 94'll [MNHN]. There are an additional male and female with similar label information designated as paralectotypes [MNHN]. Say's (1824:279) "var. a" of A. cinctus is probably this species.

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I Fig. 9. Distribution map for Malporus werneri.

Malporus properus Casey Figs. 4, 8 Malporus properus Casey 1895:653. Type locality: Keokuk, Iowa. Blatchley 1910:1338. Lectotype male, USNM. Anthicus (Malporus) properus, Pic 1911:68. Leng 1920:163. Malporus blandus Casey 1895:654. Type locality: Willets Point, Long Island, New York. Holotype female, USNM. New Synonymy. Anthicus (Malporus) blandus, Pic 1911:37. Leng 1920: 163. Description. Length 3.8-4.2 mm. Head, pronotum, and elytral base orange to brown; straight yellowish band from lateral margins across transverse postbasal impression rarely interrupted at middle, orange and indistinct in a few specimens; posterior portion of elytra usually brown, rarely partially orange. Head and pronotum with setae appressed. Head with small punctures denser in apical half of vertex, about 10 punctures in line between eyes, widely scattered in basal half, surface polished; eyes moderate in size, 0.30 mm long. Pronotumwith scattered fine punctures on disc, dorsum in basal half and lateral constrictions microreticulate and roughened, lacking distinct punctures; dorsum between constrictions slightly convex in lateral view. Elytra with long tactile setae erect at 70-80?, 0.21 mm long; setae appressed, 0.06 mm long. Males with protibial apices truncate; mesotrochanters with small basal spine on posterior margin; metatibiae slightly sinuate; tegmen of aedeagus not divided apically, apex flared laterally and with median point (Fig. 4).

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Distribution (Fig. 8). Specimens (66) have been seen from the United States: Connecticut, New York, Pennsylvania, New Jersey, Maryland, District of Columbia, Virginia, Tennessee, Ohio, Illinois, Missouri, Kansas, Arkansas, Oklahoma, Texas, and Mexico: Nuevo Leon. The specimen from Nuevo Leon has the following label data: Santa Rosa Canyon, 14.8 mi W Linares, 2400', Route 80, VII-6/7-1966, G. E. Ball & D. R. Whitehead, blacklight (UAIC). Specimens are in collections of: CNCI, DSC, EJF, IRCW, MCZC, TAMU, UAIC, UMRM, USNM. Biological and Collection Information. Few collecting data are associated with this species. A few specimens have been collected in coastal beach drift and in washed-up debris along a lakeshore. One specimen was found in a "dead rodent trap", and the Mexican specimen was collected at ultraviolet light. Blatchley (1910) collected one specimen by sifting leaves. In the late 1800's this species was frequently collected in the mid-Atlantic states. How- ever, I have only seen one record from east of the Appalachians obtained during the last 70 years, and that was from Cecil County in northeastern Mary- land. Type Designations. The lectotype (male) here designated bears the labels //Ia (with red dot before "I")/ d/ Casey bequest 1925/ TYPE USNM 36541/ properus Csy/ LECTOTYPE Malporus properus Casey des. DSChandler '94/. A paralectotype male from Indiana is also designated. Both specimens are members of M. properus. Werner (1975) placed M. blandus as a junior synonym of M. fornicarius, but the holotype of M. blandus is clearly conspecific with M. properus based on the short appressed elytral setae and other characteristics.

Malporus werneri Chandler, new species Figs. 5, 9

Description. Length 3.8-4.1 mm. Head, pronotum, and elytral base bright orange to orange brown; jagged yellowish band extending from lateral margins across posterior portion of postbasal impression, with narrow dark brown triangular line on anterior margin of band near lateral margins; elytra posterior to band dark brown. Setae on head and pronotum appressed. Head with vertex between eyes microreticulate and more dense- ly punctate, about 12 punctures in line between eyes, posterior half of vertex with fine sparser punctures; eyes large, 0.31 mm long. Pronotum with disc sparsely and finely punctate, dorsum in basal half coarsely punctate and microreticulate, extending laterally into constrictions; dorsum above lateral constrictions flat in lateral view. Elytra with suberect to erect tactile setae most commonly at angles of 40-50?, 0.22 mm long; setae appressed, 0.07 mm long. Males with protibiae truncate at apices; meso- and metatro- chanters with posterior margins smoothly rounded, lacking spines or angulations; tegmen of aedeagus with deep paired incision at apex, outer apices straight in lateral view; tegmen same depth through basal two-thirds of length in lateral view (Fig. 5).

Specimens Examined. Holotype: c, Arizona, Santa Cruz Co., Canelo, III-28-1970, K. Stephan (FSCA). Paratypes: Arizona: 1 9, Cochise Co., Guadelupe Canyon, 30 mi E Douglas, VII-25-1985, P. K. Lago (DSC); 1 9, Yavapai Co., Ash Fork, VI-30-1991, W. F Chamberlin (TAMU); 2 8, Ar[izona], Horn Collection (MCZC); 1 ?, Yuma Co., Yuma [locality incor- rect?], Horn Collection (MCZC). New Mexico: 1 c, Grant Co., Silver City, VII-1973, R. Lenczy (DSC); 1 9, Luna Co., Deming, VII-22, Hubbard & Schwarz (USNM). Distribution. The eight specimens seen are from the middle elevations of

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Acknowledgments

I have kept records of my identifications of members of this genus submitted to me from individuals and institutions for over twenty years, and recently borrowed material from most major collections. I would like to thank all who have cooperated with the loan of the material in their care. I would like to particularly like to thank those who loaned me the types in their care: Gloria House, Smithsonian Institution, Washington, D.C.; Philip D. Perkins, Harvard University, Cambridge; and Nicole Berti, Museum National d'Histoire Natu- relle, Paris. All individuals and their associated collections are as follows: CBB, Cheryl B. Barr, Berkeley, CA (private collection); CDAE, Fred G. An- drews, California State Collection of , Plant Pest Diagnostics Cen- ter, Sacramento, CA; CNCI, J. Milton Campbell, Anthony Davies, and Yves Bousquet, Canadian National Collection of , Agriculture Canada, Ot- tawa, ON, Canada; DSC, private collection of author; EDUM, Robert E. Roughley, Department of Entomology, University of Manitoba, Winnipeg, MB, Canada; EGR, Edward G. Riley, College Station, TX (private collection); EJF, Everett J. Ford Collection (Field Museum of Natural History, Chicago); FSCA, Michael C. Thomas, Florida State Collection of Arthropods, Division of Plant Industry, Gainesville, FL; IRCW, Daniel K. Young, Department of Entomology, University of Wisconsin, Madison, WI; JAC, John and Alberta Carr. Calgary, AB, Canada (private collection); MCZC, Philip D. Perkins, Museum of Comparative Zoology, Harvard University, Cambridge, MA; MNHN, Nicole Berti, Laboratoire d'Entomologie, Museum National d'Histoire Naturelle, Paris, France; NAUC, C. Dan Johnson, Department of Biology, Northern Arizona University, Flagstaff, AZ; PES, Paul E. Skelley, Gainesville, FL (private collection); REN, Robert E. Nelson, Department of Geology, Colby College, Waterville, ME (private collection); TAMU, Edward G. Riley, Department of Entomology, Texas A&M University, College Station, TX; UAIC, Carl A. Olson, Department of Entomology, University of Arizona, Tucson, AZ; UMIC, Paul K. Lago, Department of Biology, University of Mis- sissippi, University, MS; UMRM, Robert W. Sites, Department of Entomol- ogy, University of Missouri, Columbus, MO; UNHC, Donald S. Chandler, Department of Zoology, University of New Hampshire, Durham, NH; USNM, Gloria House, Department of Entomology, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C.; UVCC, Ross T. Bell, De- partment of Zoology, University of Vermont, Burlington, VT; VPIC, Michael Kosztarab, Virgina Museum of Natural History at Virginia Tech, Virginia Poly- technic Institute and State University, Blacksburg, VA; WSUC, Richard S. Zack, James Entomological Collection, Department of Entomology, Washing- ton State University, Pullman, WA. Tess Feltes provided the habitus illustration of Malporus formicarius. Spec- imen collection records are kept in a database file, and may be borrowed if desired. I would like to thank J. E Burger and P. C. Johnson, University of New Hampshire, for reviewing the manuscript.

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Literature Cited Blatchley, W. S. 1910. On the Coleoptera known to occur in Indiana; an illustrative descriptive catalog of the Coleoptera or (exlusive of the Rhynchophora) known to occur in Indiana with bibliography and descriptions of new species. The Nature Publishing Company, Indianapolis, Indiana, 1386 pp. Bousquet, Y. 1991. Family Anthicidae, antlike flower beetles [pp. 271-273]. In: Check- list of beetles of Canada and Alaska (Y Bousquet, editor). Agriculture Canada, Research Branch, Publication 1861/E, vi+430 pp. Casey, T. L. 1895. Coleopterological notices, VI. Annals of the New York Academy of Science 8:435-838. Kitayama, C. Y. 1982. Biosystematics of anthicid larvae (Coleoptera: Anthicidae). Coleopterists Bulletin 36:76-95. LeConte, J. L. 1852. A synopsis of the Anthicites of the United States. Proceedings of the Academy of Natural Science, Philadelphia 6:91-104. LaFerte-Senectere, F. T. de. 1847. Livraison No. 29, Anthicus (Troisieme Division) [pp. 85-132, plage 1]. In: Species et iconographie generique des animaux articul6s ... Premiere Partie: Insectes Col6opteres (M. F E. Guerin-M6neville, editor). Fain et Thunot, Paris. LaFert&Se'nectere, F. T. de. 1848. Monographie des Anthicus et genres voisins, Co- leopteres Het6romeres de la tribu des Trach6lides. De Sapia, Paris. xxii+340 pp, plates 17-32. Leng, C. W. 1920. Catalogue of the Coleoptera of North America, north of Mexico. Cosmos Press, Cambridge, Massachusetts. x+470 pp. Mawdsley, J. R. 1993. The entomological collection of Thomas Say. Psyche 100: 163-171. Pic, M. 1911. Anthicidae [Pars 36]. In: Coleopterorum Catalogus (S. Schenkling, edi- tor). W. Junk, Berlin. 102 pp. Say, T. 1824. Descriptions of Coleopterous insects collected in the late expedition to the Rocky Mountains, performed by order of Mr. Calhoun, Secretary of War, under the command of Major Long (continued). Journal of the Academy of Natural Science, Philadelphia 3:238-282. Uhmann, G. 1976. Die Gattungen der Anthicidae und ihre systematische Anordnung. Entomologische Blatter 72:166-182. Uhmann, G. 1978. Die Gattungen der Anthicidae und ihre systematische Anordnung. Teil 2. Die Systematische Anordnung. Entomologische Blatter 74:75-80. Werner, F. G. 1963. Fascicle 84. Anthicidae (Latreille, 1825), the ant-like flower bee- tles [pp. 747-752]. In: The beetles of the United States (a manual for identifica- tion) (R. H. Arnett, Jr., editor). Washington, D.C., Catholic University Press. xi+1112 pp. Werner, F. G. 1964. A revision of the North American species of Anthicus s. str. (Coleoptera: Anthicidae). Miscellaneous Publications of the Entomological Soci- ety of America 4:195-242. Werner, F. G. 1975. New synonymy in the Nearctic Anthicidae (Coleoptera). Pro- ceedings of the Entomological Society of Washington 77:290. Werner, F. G., and D. S. Chandler. 1995. Anthicidae (Insecta: Coleoptera). Fauna of New Zealand, Number 34, 64 pp. (Received 14 October 1996; accepted 3 January 1997)

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