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9-1999

Revision of the North American Species of with a Checklist of the World Species (Coleoptera: )

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

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Recommended Citation Revision of the North American Species of Amblyderus with a Checklist of the World Species (Coleoptera: Anthicidae) Donald S. Chandler Transactions of the American Entomological Society (1890-), Vol. 125, No. 3 (Sep., 1999), pp. 269-293

This Article is brought to you for free and open access by the New Hampshire Agricultural Experiment Station at University of New Hampshire Scholars' Repository. It has been accepted for inclusion in New Hampshire Agricultural Experiment Station Publications by an authorized administrator of University of New Hampshire Scholars' Repository. For more information, please contact [email protected]. Revision of the North American Species of Amblyderus with a Checklist of the World Species (Coleoptera: Anthicidae) Author(s): Donald S. Chandler Source: Transactions of the American Entomological Society (1890-), Vol. 125, No. 3 (Sep., 1999), pp. 269-293 Published by: American Entomological Society Stable URL: http://www.jstor.org/stable/25078683 Accessed: 15-08-2014 15:39 UTC

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This content downloaded from 132.177.228.65 on Fri, 15 Aug 2014 15:39:11 UTC All use subject to JSTOR Terms and Conditions Transactions of the American Entomological Society 125(3): 269-293, 1999

Revision of the North American Species of Amblyderus with a Checklist of the World Species (Cole?ptera: Anthicidae)1

Donald S. Chandler

Department of Zoology, University of New Hampshire, Durham, NH 03824

ABSTRACT

are a The North American species of Amblyderus revised, and key is provided. as a new InAmblyderus Pic is placed synonym of Amblyderus. Six species are redescribed, and one new species, A. owyhee Chandler, is described from the Columbia River Plateau. A checklist of the world species and their distribu tions is included. Lectotypes for granularis LeConte, Anthicus pallens LeConte, Amblyderus obesus Casey, Amblyderus albicans Casey and Amblyderus are parviceps Casey designated. The record of Amblyderus from Puerto Rico is a found to be due to misspelling of the genus Amblycerus Thunberg (Family Chrysomelidae, Bruchinae).

Amblyderus was proposed originally by LaFert?-S?nect?re (1849a) for two species of Anthicidae from northern Africa. Currently this genus contains 32 species found primarily in coastal dunes along the shores of the Mediterranean region, Africa, India, Sri Lanka, and North America, but also is known from dry inland dune areas in central Africa, India, and now, North America (Weissmann and Kondratieff, 1999). Amblyderus was first recognized inNorth America by Casey (1895), who provided the only revision of the genus. He described six species and added two species described by LeConte (1850) in Anthicus Paykull, which had been found along the shores of Lake Superior. There has been little subsequent work directly treating members of this genus. The genus has been treated in a key to the genera of North American Anthicidae by Werner (1964), and Downie and Arnett (1996) did provide a key to the species in Northeastern North America. A brief note by Werner (1975) placed four of Casey's names as one each to the synonyms, assigning resulting four valid species. Weissmann and Kondratieff (1999) recently described two species from Great Sand Dunes National Monument in Colorado, bringing the number of species inNorth America to six. These are the first North American species known to be restricted to a dune complex not or associated with riverine, marine, lake systems. Wolcott (1936:210; 1951:301) reported an Amblyderus species that

Scientific Publication Number 2011 of the New Hampshire Agricul tural Experiment Station

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had been found on the flowers of Randia mitis (= Randta aculeatus L.) and other trees by R. G. Oakley at Ponce, Puerto Rico. Werner (1983:232) as an association with flow questioned this extraordinary discovery, ers or trees has never been encountered for any members of this genus. not to However, he could find the specimens confirm this identifica tion. Michael A. Ivie (Montana State University) has spent consider able time working with the fauna of the Caribbean. He noted (pers. comm.) that the Oakley specimens were sent to the National Museum of Natural History (Washington, D.C) for identification, that Werner had already searched that collection, and that no Amblyderus specimens taken by Oakley could be found. However, he noticed that Kingsolver (1970) reported specimens from the National Museum of Natural History of his newly described species, Amblycerus schw?rzt Kingsolver (Bruchinae, Chrysomelidae), with the appropriate label information (Puerto Rico, Ponce, Sauri Finca, Aug. 8,1931, in flowers of Randia R. G. aculeata L., Oakley). Another species, Amblycerus cerdanicola is at Kingsolver (loc. cit.), also the National Museum of Natural History with appropriate label information, but was taken on - flowers of Cordia probably the "other flowers" originally mentioned by Wolcott. Though Wolcott had to have had the identification slips for these species (as Amblycerus sp., since they were not described species at that time) sent to him before he published his lists of the Puerto Rican fauna (Wolcott, 1936,1951), he does not mention them in the bruchine records. Therefore, the record of Amblyderus from Puerto Rico seems to be based on a misspelling for Amblycerus Thunberg, a member of the bruchine Chrysomelidae. I have been identifying members of Amblyderus for over twenty and the distributions of the eastern are now years, species recognized to be much broader than recorded in the literature. Since Amblyderus has not been revised since Casey (1895), I thought it time to place the two new species described by Weissmann and Kondratieff and a new species described here within the context of a revision of the North American taxa. This leaves only LaFert?-S?nect?re {-Thicanus Casey) as the last North American genus that has not been examined since Casey's (1895) treatment of the North American fauna.

BIOLOGY

Members of most species of Amblyderus are found in sand dunes the seacoasts or along margins of freshwater lakes and rivers. Three in are now species North America known to be associated with large interior dune masses of the Basin and are Range region, and not found near water. In all areas are necessarily adults scavengers, on or feeding dead debris that is blown about by the winds that the a or keep dunes dynamic system, on insects and other materials washed the of marine or up along margins freshwater systems associ ated with the dunes. Adults typically emerge from the sand at dusk and to or begin search for food mates in the early evening hours. In the Great Basin dune systems, adults may line up just below the crest of

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the dune to snap at debris as it blows over the dune crest (Weissmann and Kondratieff 1999), or scurry about low points in the dunes where debris collects. Along the margins of fresh and saltwater systems at of adults begin moving dusk between piles wrack, carrion, and as other debris searching for food, and then burrowing into the sand the Larvae are unknown at this time. day approaches. were Mating and other behavioral activities documented for Amblyderus trtplehornt Weissmann and Kondratieff (Weissmann and Kondratieff, 1999). The most thorough treatment of seasonality and habitat preferences of members of Amblyderus was by Ronchetti et al. (1986) for two species on the coast of Somalia. They found that the two for sea or areas species exhibited distinct preference either the margin further back in the dunes, and had their greatest abundance at differ ent times of the year which were associated with rainfall patterns.

SYSTEMATICS

The North American species are different from the Old World species, in lacking the row of large teeth along the anterior rim of the pronotal disc, which is typical of most of the Old World species, including the type species Anthicus scabricollis LaFert?-S?nect?re from the western Mediterranean. This led Pic (1911a) to create the subgenus Inamblyderus for the North American species, though it has never been used by North American workers, and has been frequently ignored by other workers on the Anthicidae. Later Krekich-Strassoldo (1931) name to elevated this the generic level without discussion, which was followed by Uhmann (1976,1978), while Bonadona (1986) treated it as a subgenus. The African, Mediterranean, and Indian species placed in the have a series of blunt teeth subgenus Amblyderus prominent, along the anterior of the setae are margin pronotal disc, the elytral often suberect, and erect tactile setae are Three Old prominent, present. World species have been placed in Inamblyderus (scabridus Krekich-Strassoldo from northeastern India, and brincki Bonadona and bigibber Bonadona from Sri Lanka), because they lack these teeth, while bigibber lacks the raised elytral setae and prominent tactile setae typical of the other Old World species. Amblyderus truncatus LaFert?-S?nect?re from Egypt has the pronotal teeth small and the elytral setae are appressed, but the tactile setae are prominent and erect. The North American species lack teeth on the anterior rim of the the setae are pronotum, elytral appressed (at over least the disc), and the tactile setae are always short and difficult to see. Beyond the presence or lack of distinct teeth on the anterior rim of the pronotum, the two groups cannot be separated. Since this of the are development pronotal teeth and setae variable within the Old World species, I think it is best to treat Inamblyderus as a junior synonym of Amblyderus until a thorough analysis of the Old World can species be pursued. The two disjunct distributions, one group centered inNorth America and the other in Africa/India/the Mediterranean, does present the

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question of whether the genus ismonophyletic. Since all species are or burrowers in sand dunes very sandy soils, the particular robust body form that is characteristic of Amblyderus might be due to conver gence based on occupation of similar habitats. Members of Anthicus a is that occupy sand dunes also have large pronotum that broadest a and often somewhat heads. anteriorly, short metasternum, triangular However, I know of no Anthicus that have the anterior face of the to anterior pronotum abruptly declivous the cervix, the face flattened, and have distinct granules or teeth across the anterior margin. This feature does seem to unite Amblyderus into a monophyletic group, though it is otherwise very close to Anthicus. METHODS

Body length is the distance from the labral apex to the elytral apex, with each of the main body parts measured separately in a lateral view and added together. Setal angles in degrees are determined as the average angle of the line drawn from the setal shaft near the base to near or are the apex in reference to the elytral surface, characterised more as: the of the seta generally 'appressed', tip slightly curved the touching body surface; 'subdecumbent', slightly raised, the gen eral of the shaft angle 10-20?; 'decumbent', clearly raised, 20-40?; 'suberect', 40-60?; or 'erect', 60-90?. A list that interpreted the cryptic label notations that Casey used to indicate localities was provided by Gloria House (Smithsonian Insitution, Washington, D.C). Complete label data is given for the new more com species and those with relatively few records; the monly collected species have only locality data given, though com are plete data available from the author. The male are extracted from genitalia specimens by softening them in hot water for 15 minutes, placing the specimens in small dishes in 70% alcohol, lifting the elytra and slitting the soft tergal sclerites, and then a of to and extract using pair forceps grasp the aedeagus. The aedeagus is then placed in 10% KOH until clear enough for illustra tion, and rinsed with 70% alcohol. The genitalia are placed in a slide mount and illustrations are made a camera glycerine using on a lucida Wilde M-10. Details of the illustrations are then checked by a examining the genitalia under Wilde M-5 APO. The genitalia are placed in small microvials in glycerine and pinned beneath the speci or are mens, simply glued to the top of the point using awater soluble glue. The following list indicates the acronyms for the curators/collec tors, and collections that furnished material for this study.

BCPM. Robert A. Cannings. Royal British Columbia Museum, Victoria, BC, CANADA. CASC?Roberta Brett and David H. Kavanaugh. California Academy of Sci ences, San Francisco, CA. CDAE?Fred G. Andrews. State California Collection of , Califor

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nia Department of Food and Agriculture, Sacramento, CA. CNCI?J. Milton Campbell, Anthony Davies, and Yves Bousquet. Canadian National Collection of Insects, Agriculture Canada, Ottawa, ON, CANADA. DSC?Private collection of author.

EGR?Edward G. Riley. College Station, TX (private collection). CID A?William H. Clark. Orma J. Smith Museum of Natural History, Albertson College, Caldwell, ID. HNHM?Z. Kaszab. Zoological Department, Hungarian Museum of Natural History, Budapest, HUNGARY. IRCW?Daniel K. Young. Dept. of Entomology, University of Wisconsin, Madison, WI. JAC?John and Alberta Carr. Calgary, AB, CANADA (private collection). KHSC?Karl H. Stephan. Red Oak, OK (private collection). LACM?Julian P. Donahue and Brian V. Brown. Los Angeles County Museum of Natural History, Los Angeles, CA. MCZC?Philip D. Perkins. Museum of Comparative Zoology, Harvard Uni versity, Cambridge, MA. NAUF?C. Dan Johnson. Department of Biology, Northern Arizona Univer sity, Flagstaff, AZ. OSUC?Charles A. Triplehorn. Department of Entomology, Ohio State Uni versity, Columbus, OH. TAMU?Edward G. Riley. Department of Entomology, Texas A&M Univer sity, College Station, TX. UAIC?Carl A. Olson. Department of Entomology, University of Arizona, Tucson, AZ. K. UMIC?Paul Lago. Department of Biology, University of Mississippi, Uni versity, MS. F. O'Brien. UMMZ?Mark Museum of Zoology, University of Michigan, Ann Arbor, MI. UNHC?Donald S. Chandler. Department of Zoology, University of New Hampshire, Durham, NH. USNM?Gloria House. Department of Entomology, National Museum of Natu ral History, Smithsonian Institution, Washington, D.C. UVCC?Ross T. Bell. of Dept. Zoology, University of Vermont, Burlington, VT.

AMBLYDERUS LaFert?-S?nect?re

Amblyderus LaFert?-S?nect?re 1849a:l. Type species: Anthicus scabricollis LaFert?-S?nect?re, subsequent designation by Jacquelin du Val, 1863:368. LaFert?-S?nect?re 1849b:62. Casey 1895:742. Pic 1911c:27. Leng 1920:163. Uhmann 1976:171; 1978:75. Bucciarelli 1980:112. Poole and Gentili 1996:50. Ambliderus Pic 1950:151 (misspelling). Pic 1911a:134 Inamblyderus (as subgenus). Type species: Amblyderus obesus Casey, original designation. Pic 1911c:27. Bonadona, 1986:61. NEW SYN ONYMY.

Inamblyderus (as genus). Krekich-Strassoldo, 1931:15. Uhmann 1976:171. Uhmann 1978:75.

Head Diagnosis.? and pronotum granulate; head triangular, slightly widest near anterior elongate; pronotum trapezoidal, margins, with basal in some margins briefly parallel species; anterior face of prono tum abruptly declivous; metasternum short, distance between meso and metacoxae less than mesocoxae. length of

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mm Fig. 1. Dorsal view Amblyderus triplehorni; specimen 59 long.

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3M

Figs. 2-8. Ventral and right lateral view male genitalia (line=0.3 mm) n. A. 2, A. granularis (LeConte). 3, A. obesus Casey. 4, A. owyhee Chandler, sp. 5, A. pallens (LeConte). 6, A. parviceps Casey, 7, triplehorni Weissmann and Kondratieff. 8, A. werneri Weissmann and Kondratieff.

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1_ . ' ' |

Fig. 9. Distribution map: A. granularis (circle), A. obesus (square), A. owyhee (triangle), A. triplehorni (oval).

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Fig. 10. Distribution map: A. pallens (circle), A. parviceps (square), A. werneri (triangle).

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Description.? Body robust, head and pronotum typically distinctly setae between Head granulate, arising granules. nearly triangular; base with vertexal angles prominent, median cleft shallow to deep; frontoclypeal suture distinct; eyes large, with short interocular setae; apical antennomeres slightly laterally flattened; maxillary palpi with mesal margins of fourth segments angularly produced; mandibles with more than ventral margin prominent dorsal margin, ventral margins slightly and broadly produced as a flange. Pronotum widest near anterior trapezoidal, margin; abruptly to declivous from anterior margin of disc collar, anterior face flat; disc gently convex to base; lateral margins nearly straight in converging toward base from prominent anterior angles; pronotal base briefly with parallel margins to form broad neck in some species; thin antebasal on sulcus lateral margins extending anteroventrally to foveae to posterodorsal procoxae. Elytra with prominent humeri; sutural striae present only in apical half of elytra; flight wings reduced in North American species. Mesocoxae large, clearly separated by extension of mesosternum which meets process of metasternum between coxae; distance be tween meso- and metacoxae less than meso short, clearly length of lateral of mesosternum coxae; margins straight; mesepimera slightly to covered exposed completely by mesepisterna, mesepimera deeply to mesocoxae. impressed, with shallow fovea dorsolateral Protibiae with at prominent mesal angulation apices. Males with sternite IX thick, with dorso-ventrally flattened expan sion at and exten base; apex broadly very shallowly forked, apical sions articulated at base and broadly widened to near apices; male protibiae sinuate on mesal margin in North American species, with thick setae brush of along sinuation. Blatchley (1910:1343) notes that Amblyderus is translated from Greek as "obtuse neck."

KEY TO SPECIES

1 Restricted to California coastal sand dunes.2 ? Coastal and inland dune areas of southern Canada and the United States, but not the California coast.3 2(1) Larger, 3.7-4.3 mm long; pronotum with short but distinct neck at base, sides at base in dorsal view briefly parallel .obesus Casey ? 2.5-3.6 mm Smaller, long; pronotum with lateral margins evenly converg to base in . ing dorsal view, lacking parallel margins at base . parviceps Casey Lateral of 3(1) margins pronotum and elytra with long (0.15-0.20 mm) erect to suberect setae; southern Colorado. 4 ? Lateral of and margins pronotum elytra with setae appressed or de short erect pressed, only very (0.02 mm) tactile setae rising from surface . 5 Head and vertex convex 4(3) clearly granulate; pronotal disc strongly (Fig. 1) .triplehorni Weissmann and Kondratieff

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? Vertex and pronotum smoothly rugose; pronotal disc shallowly convex, more curvature slightly than that of head vertex. . werneri Weissmann and Kondratieff

5(3) Anterior face of pronotum with setae clearly suberect, all raised from mm surface; large species, 4.2-5.4 long; Idaho to southern British Co n. lumbia and Alberta.owyhee Chandler sp. ? a Anterior face of pronotum with setae appressed, rarely with few mm slightly raised setae visible; smaller species, 3.1-4.2 long.6 or 6(5) Pronotum with distinct neck at base, lateral margins parallel for 15% more of pronotal length in dorsal view; elytra shining, setae sparser and shorter, setae extending at most to about half length beyond next posterior puncture; eastern North America .granularis (LeConte) ? or Pronotum with neck at base shorter, 10% less total pronotal length in dorsal view; elytra microreticulate and dull, setae appearing denser and longer, reaching at least to second puncture posterior to setal origin . pollens (LeConte)

Amblyderus granularis (LeConte) Figs. 2, 9

Anthtcus granularis LeConte 1850:231. Type locality: Lake Superior. Lectotype here A. male designated, // [light green disc]/ granularis Lee./ Type 4930/ LECTOTYPEAnthicus granularis LeConte des. DSChandler'98// [MCZC]. are Single male and female specimens with similar locality labels desig nated as PARALECTOTYPES. LeConte 1852:103; LeConte 1866:65. Hubbard and Schwarz 1878:641, 662. Amblyderus granularis, Casey 1895:746. Pic 1911c:27. Leng 1920:163. Werner (synonymy) 1975:290. Poole and Gentili 1996:50. Downie and Arnett 1996:1189.

Amblyderus punctiger Casey 1895: 747. Type locality: Lake Superior. Holotype female, // L. Sup/ female/ Casey bequest 1925/ TYPE USNM 36620/ punctiger Csy// [USNM]. Pic 1911c:27. Leng 1920:163.

Diagnosis.? Moderate in size (3.5-4.3 mm); pronotum with broad basal neck; pronotal and elytral setae appressed; elytra shining; only species with a median band on the elytra, though not all specimens have this; aedeagus with blunt median point on broad apex; east of Rocky Mountains. 3.5-4.2 mm. Description.? Length Body with head and pronotum lightly reticulate, elytra shining; setae short and appressed over body, on 0.07 mm on Color appearing sparser elytra, long elytra. entirely dark brown for some from most specimens Canada, specimens with head and pronotum brown and elytra with brown marking forming narrow zone at base and broad from midband; specimens Mississippi have head and pronotum tan to light brown with the elytra slightly lighter and lacking any dark markings. Head with vertex coarsely granulate; base with distinct medial vertexal at cleft; angles sharp, about 80 ?; antennomeres VIII-IX about as as long wide in dorsal view, X wider than long, the rest longer than wide; gula with 2-4 long setae. Pronotum with distinct neck at base, in dorsal view lateral margins parallel for about 15% pronotal length; disc and margins granulate; a

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few tactile setae visible lateral in dorsal short, erect, along margins view, 0.02 mm long; anterior face of pronotum with setae appressed; pronotal disc shallowly convex, apex with shallow median cleft; erect setae. prosternum with several long Elytra with a few short tactile setae visible along lateral margins in dorsal view; elytral apices broadly rounded and prolonged to near suture. Males with mesal margin of protibiae nearly straight; sternite VIII broadly rounded at apex; aedeagus slightly narrowed in apical por tion to broad apex, apex with median point. This has been on Biology.? species usually taken under debris beaches of fresh-water lakes and large rivers, but is also found rarely near are brackish coastal habitats. Adults present from April to Sep tember around the Great Lakes, but most specimens were taken in May. Relationships.? This species shares with A. owyhee the distinct broad neck at the base of the pronotum, strongly granulate head and rounded to male pronotum, elytra broadly apices, protibiae with mesal and the with a margins nearly straight, aedeagus median point on a broad be the smaller apex. They may separated by size, appressed setae on the anterior and smooth pronotal face, elytra with shorter and setae of A. while A. is in sparser granularis; owyhee larger size, has raised setae on the anterior and the are pronotal face, elytra clearly microreticulate with denser and setae. longer Distribution.? Most commonly taken from around the Great Lakes, now a but known from few isolated localities throughout eastern North America. UNITED STATES. Illinois: Lake Co., Beach State Park. Indiana: Porter Co., Beverly Shores; Indiana Dunes State Park. Iowa: Des Moines Skunk nr. Co., River, Burlington. Louisiana: Natchitoches Parish, Kisatchie National Forest, Kisatchie Bayou Camp ground. Maryland: Anne Arundel Co., 10 km NE Annapolis, Sandy Point. Massachusetts: Middlesex Co., Tyngsboro, Merrimack River. Michigan: Berrien Co., Warren Sand Dunes State Park. Macomb Co., east of E of Memphis. Marquette Co., bay Marquette; Huron Moun Huron Mountain Club. tains; Muskegon Co., Muskegon. Sanilac Co., Sanilac State Park. Mississippi: Harrison Co., Newman Lumber Com Landon. New York: pany, Niagara Co., Niagara; county unknown, Lake Ontario. North Dakota: Burleigh Co. Ohio: Ashtabula Co., beach at Chestnut Grove. Vermont: Chittenden Co., Burlington. CANADA. Ontario: Lake Algoma Co., Superior Provincial Park; Michipicoten River; Wawa. Essex Co., Point Pelee. Kent Co., Rondeau Provincial Park. Lambton Grove. Co., Bright's Niagara Co., Ridgeway. Ottawa Carleton Co., Ottawa River, Deschenes Lookout. Prince Edward Co. Quebec: Chateauguay; La Trappe. Specimens in collections of: CASC, CDAE, CNCI, DSC, HNHM, LACM, MCZC, NAUF, OSUC, UAIC, UMMZ, USNM, UVCC

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Amblyderus obesus Casey Figs. 3, 9

Amblyderus obesus Casey 1895:743. Type locality: Marin County ["San Fran sea cisco"] beaches, California. Lectotype male here designated, // Cal = [with horizontal line in "C" Marin County]/ male/ Casey bequest 1925/ TYPE USNM 36616/ Amblyderus obesus Csy/ LECTOTYPE Amblyderus obesus Casey des. DSChandler '98// [USNM]. Five males and two females are as with similar collection labels designated PARALECTOTYPES. Pic 1911c:27. Leng 1920:163. Werner (synonymy) 1975:290. Poole and Gentili 1996:50.

Amblyderus albicans Casey. 1895: 744. Type locality: Santa Monica sea beaches, California. Lectotype male here designated, // Cal [underlined and with "C" = short horizontal line through Santa Monica]/ male/ Casey bequest 1925/ albicans 2 PARATYPE USNM 36617/ LECTOTYPEmale Amblyderus albicans Casey des. DSChandler '98// [USNM]. Three female specimens a are as with similar similar locality label designated PARALECTOTYPES. Pic 1911c:27. Leng 1920:163.

in Diagnosis.? Moderate size (3.7-4.3 mm); pronotum with short but distinct broad basal neck; pronotal and elytral setae appressed; completely yellowish or with elytra bluish-brown; male genitalia near to in sand dunes narrowing apex sharp point; only along Pacific coast. 3.7-4.3 mm. common com Description.? Length Two color forms: pletely yellowish to orange, or with head and pronotum orange and elytra bluish-brown. Dorsum lightly microreticulate, head more pol ished and reticulation difficult to setae on see; appressed dorsum, dense on 0.09 mm on elytra, long elytra. Head smoothly roughened, granules difficult to distinguish; head base with median cleft barely indicated; vertexal angles broadly at about 80 antennomeres X wider than in rounded, ?; long dorsal view, VIII as as and IX about wide long, the rest elongate; gula lacking setae as as on long those prosternum. Pronotum with lateral margins slightly convex and converging toward base, with brief neck at base, lateral margins parallel at base for about 5% pronotal length; disc with small granules most distinct at anterior margin, granules becoming less distinct posteriorly; disc with shallowly convex, apex vague median impression; lateral mar gins in dorsal view with a few short erect tactile setae, setae 0.02 mm anterior face of with setae setae long; pronotum appressed; prosternai moderate in length, about half eye length. with some decumbent setae at humeral mar Elytra angles; lateral in gins dorsal view with scattered short tactile setae barely visible; elytral apices broadly rounded to subtruncate. Males with protibiae moderately arcuate, mesal margins with dense short sternite brush; VIII broadly truncate at apex; aedeagus narrowing toward apex, apex with small tooth. Found sea Biology.? along beaches beneath wrack or carcasses on

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the sand. Adults are present from March toDecember, with the longest series taken in June and July. Relationships.? This species is closest to A. parviceps in sharing the sinuate mar smoothly granulate head and pronotum, strongly mesal of the male rounded to subtruncate gins protibiae, broadly elytral to the It is apices, and the male genitalia narrowing apex. easily separated by the larger size of A. obesus, the brief neck at the base of the at to pronotum, and the male aedeagus abruptly narrowed the apex form a distinct point. Distribution.? Sandy beaches from southern British Columbia to the Los Angeles area. UNITED STATES. California: Humboldt Co., Clam Trinidad Samoa Dunes. Los Beach, Harbor; Eureka; Angeles Co., Point Dume; Santa Monica. Marin Co., Bolinas Point, 1.6mi W Bolinas; - Dillon Beach; Point Reyes National Seashore 10 Mile Beach, Point Reyes Beach South, RCA sand dunes; Stinson Beach. Mendocino Co., Gualala; Mackerricher State Park; Westport. Monterey Co., Del Monte; Moss Marina, Beach Road; Landing; Monterey; Seaside. San Francisco Co. San Luis 3 mi N Point Buch?n. San Mateo Obispo Co., Co., Pacifica. Santa Barbara Santa Barbara. Santa Cruz 16 mi Co., Jalama; Co., Aptos; S Santa Cruz, Sunset Beach. Sonoma Co., Sonoma Coast State Beach, South Salmon Creek Landing. Ventura Co., Point Mugu Naval Air Station; San Nicolas Island. CANADA. British Columbia Vancouver: Island, Duncan. A small series collected by H. F.Wickham is labeled as being from the Pinal Mountains in central Arizona (MCZC), and is here judged to be mislabeled. Specimens in collections of: BCPM, CASC, CDAE, CNCI, DSC, JAC, LACM, MCZC, USNM.

Amblyderus owyhee Chandler, New Species Figs. 4, 9

in size Diagnosis.? Moderate (3.7-3.8 mm); pronotum with broad basal setae on anterior face of neck; pronotum suberect, appressed over rest of at humeral body except angles; body yellowish, sometimes with brown blotches on with elytra; aedeagus sharp, short median on point broad apex; Columbia River Plateau and southwestern Canada. 3.7-3.8 mm. to Description.? Length Body yellowish orange, elytra sometimes with large lateral brown blotches, abdomen darker in specimens with elytral blotches; dorsum lightly reticulate; setae ap over on 0.08 mm pressed dorsum, elytra long and dense. Head with vertex densely and evenly granulate; base with distinct medial cleft; vertexal angles narrowly rounded at about 80 ?; antennomeres X slightly wider than long in dorsal view, VIII-IX about as as the rest than no more long wide, longer wide; gula with than two setae as as setae on long long prosternum. Pronotum with lateral margins convexly converging toward base, with short at neck base, margins parallel for about 10% pronotal length in dorsal view; disc shallowly convex and evenly granulate, apex with shallow median a impression; lateral margins with few short erect tactile setae visible in dorsal setae 0.02 mm view, long; anterior face of

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near pronotum with setae suberect, especially cervix; prosternum with about 6 long setae. Elytra with scattered short erect tactile setae visible on lateral in most setae some at margins dorsal view, appressed, slightly raised to near suture. humeral angles; elytral apices broadly rounded Males with mesal margins of protibiae nearly straight; sternite VIII with apex broadly convex; aedeagus with lateral margins straight to to median apex, apex broadly rounded point. Biology.? Adults are buried in dune slip-faces during the day; at night they run about investigating the debris that has accumulated at the bottom of the Most were in dune slip-faces. specimens taken May and June, and a smaller number were taken as late as November. Relationships.? It is closest to A. granularis as judged by the distinct basal lateral of and with pronotal neck, margins pronotum elytra only short setae tactile erect, elytral apices conjointly rounded, and male genitalia broad to the apex. This species may be separated by its larger on size, and the presence of suberect setae the anterior face of the pronotum. Distribution.? Found in dune areas of the Columbia River Plateau to southern British Columbia and Alberta. Specimens examined: Ho lotype male: UNITED STATES, Idaho, Owyhee Co.,Bruneau State Park, V-29-1986, B. F. & J. L. Carr. Lot 3 (USNM). Paratypes: 61: UNITED STATES, 3M, 3 F, Bruneau State Park, V-27-1986, B. F. & J. L. Carr, Lot 1; 5 males, 3 females, Bruneau State Park, V-27-1986, B. F. & J. L. Carr, Lot 9; 2 males, 1 female, Bruneau State Park, V-25-1986, B. F. & J. L. Carr, Lot 5; 2 males, 1 female, Bruneau State Park, V-25-1986, B. F. & J. L. Carr, Lot 7; 1 male, Bruneau Dunes, XI-24-1974, D. Giuliani; 1 male, Bruneau Dunes, IV-25-1976, A. D. Allen; 4 males, , Bruneau Sanddunes, V-24/31-1986, C and A. van Nidek; 2 males, 1 female, Bruneau Dunes State D. S. at Park, VII-17-1988, Chandler, dunes, light; 3 males, 7 females, Bruneau Dunes State Park, V-22-1970, V-23-1970, J. D. Marshall; 1 male, Sand Dune Lakes, 7 mi NE Bruneau, VI-24-1970, W. E. at Fremont 1 St. VI Ferguson, light. Co., female, Anthony Dunes, A. D. on dunes at 4 St. 15-1977, Allen, night; females, Anthony Dunes, K. on dunes at 1 St. VI-15-1977, Allen, night; female, Anthony Dunes, VII-1-1978, G. A. Shook; 1male, 1 female, 5 km N Parker, VI-6-1986, B. F. & J. L. Carr; 1 male, 3 females, 5 km N Parker, VI-7-1986, B. F. & J. L^Carr. CANADA. Alberta: 2males, 1 female, Medicine Hat, X-9-1971, B. F. & J. L. Carr, Lot 1. British Columbia: 1male, 4 females, 16 mi W & ex: river Osoyoos, VI-5-1968, Campbell Smetana, debris. Paratypes in collections of: CASC, CID A, CNCI, DSC, JAC

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Amblyderus pallens (LeConte) Fig. 5,10

Anthicus pallens LeConte 1850: 231. Type locality: Lake Superior. Lectotype male here designated, // [light green disc]/ A. pallens Lee./ Type 4929/ LECTOTYPE Anthicus pallens LeConte des. DSChandler '98// [MCZC]. Three males and two females with similar locality labels are designated as PARALECTOTYPES. LeConte 1852:103; 1866:65. Hubbard and Schwarz 1878:641, 662. Amblyderus pallens, Casey 1895:747. Blatchley 1910:1343. Pic 1911c:27. Leng 1920:163. Downie and Arnett 1996:1189. Werner (synonymy) 1975:290. Poole and Gentili 1996:50.

Amblyderus arenarius Casey 1895:748. Type locality: Newport, Rhode Island. Holotype male // R.I/ male/ Casey bequest 1925/ Type USNM 36621/ arenarius Csy// [USNM]. Pic 1911c:27. Leng 1920:163.

in size Diagnosis.? Moderate (3.1-4.2 mm); pronotum with broad basal neck setae over short; body appressed; elytra microreticulate and dull; aedeagus with short, blunt point on broadly truncate apex; body yellowish; North America except the Pacific Coast states/prov inces. 3.1-4.2 mm. to Description.? Length Body yellow light yellow dorsum head brown; lightly reticulate, occasionally smooth; setae over dense on 0.07 mm on appressed body, elytra, long elytra. Head with of vertex often Description.? granules smoothly rounded, most distinct at basal margin, surface smooth to lightly reticulate between granules; base with distinct median cleft; vertexal angles abruptly rounded, varying from nearly parallel to eyes to converging at about antennomeres all than wide in 85?; longer dorsal view, only X as wide as with 0-2 setae. nearly long; gula long Pronotum with lateral convex near margins slightly to base, mar gins parallel for short distance at base in dorsal view, neck about 10% disc with most distinct near pronotal length; granules apex and along lateral few short tactile setae visible margins; along lateral margins in dorsal setae 0.02 mm view, long; anterior face of pronotum with setae disc anterior appressed; pronotal shallowly convex, margin slightly at with 2-4 depressed middle; prosternum long setae. Elytra with setae appressed to depressed; few short, erect, tactile setae near visible along lateral margins base in dorsal view; elytral to near suture. apices broadly rounded Males with mesal margins of protibiae clearly sinuate; sternite VIII and at broadly shallowly emarginate apex; aedeagus with apex broadly truncate, with short medial tubercle at apex. are Biology.? Adults found in the sand beneath sprawling plants, in grass roots 4-8 cm beneath the dune surface, and beneath debris in areas dune and sandy beaches (Blatchley 1910). Specimens have been taken from to February November, but most records are from May to July. It is similar to A. werneri in a Relationships.? sharing short basal

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neck of the pronotum, pronotal disc broadly convex, distinct medial at to cleft the head base, elytral apices conjointly rounded the suture; and broad to the These two be aedeagus apex. species may separated a median as by the pronotal apex exhibiting shallow impression, do A. obesus and A. and the has lateral parviceps, aedeagus prominent angles at the apex. Distribution.? Dune areas throughout the United States and south ern Canada except for the Pacific Coast. UNITED STATES. Arizona: Coconino Co., Grand Canyon National Park, Colorado River mile #138L. Colorado: Alamosa Co., Great Sand Dunes National Monu Bent Florida: ment; Co., Hasty. St. Johns Co., Vilano Beach. Volusia Co., New Smyrna Beach. Georgia: Chatham Co., Tybee Island. Glynn St. Simons Island. Illinois: Cook Co., Co., Chicago. Lake Co., Beach State Park. Indiana: Porter Co., Pine; Tremont. Louisiana: Natchitoches Par., Kisatchie National Forest, Red Dirt WMA. West Feliciana Par., 2 mi Maine: W Jackson, Vaughn's Bayou. Cumberland Co., Portland. Massachusetts: Barnstable Co., Barnstable; Chatham; Provincetown. Dukes Martha's Island. Essex Co., Chilmark, Vineyard Co., Ipswich; Lynn Beach; Nahant. Hampden Co., Chicopee. Middlesex Co., Harwich; Lowell; Tyngsboro, Merrimack River. Plymouth Co., Marion. Michi gan: Marquette Co., bay east of Marquette, Lake Superior; Huron Mountains. Mason Oceana Co., Ludington. Co., Silver Lake State Park. Mississippi: Adams Co., Homochitta National Forest. Harrison Co., Landon, Newman Lumber Company. Lafayette Co., Oxford. Wilkinson Co., 4 mi W Wilkinson. Missouri: St. Francis Co., Bonne Terre. Ne braska: Valentine. New Cherry Co., Hampshire: Rockingham Co., Seabrook. New : S. M. Newcastle; Jersey Cape May Co., Beach; Anglesea. Ocean : Co., Barnegat Bay; Island Beach State Park. New Mexico Otero White Sands National Monument. New York : Co., Oswego Co., Granby Center. Suffolk Co., Riverhead. North Carolina: New Hanover Co., Wrightsville Beach. Oklahoma: Latimer Co., 4 mi W Red Oak. South Carolina: Charleston Isle of Charleston. Co., Palms, Horry Co., Myrtle Beach. South Dakota: Hamlin Co., Poinsett Lake. Texas: Bexar Co., Fort Sam Houston, San Antonio. Comanche Co., Desdomona. Lee Co. Randall Co., Palo Duro State Park. Ward Co., Monahans Sands. Vir Accomack Island. ginia: Co., Assateague Northampton Co., Cape Charles. Princess Anne 15 km E at Co., Norfolk, Lynnhaven Inlet. Wisconsin: Bayfield Co., Bayfield. CANADA. Alberta: 7 mi S Em W. 4th press; Twp. 41, Range 5, Meridian; Twp. 22, Range 1, W. 4th th Meridian; Twp. 5, Range 7, W. 4 Meridian. Manitoba: Aweme; E. 1st Rosser; Twp. 1, Range 13, Meridian; Twp. 8, Range 16, W 1st Meridian. New Brunswick : Kouchibouguac. Ontario: Essex Co., Point Pelee. Kent Co., Rondeau Provincial Park. Manitoulin Co., Manitoulin Island, Providence Bay. Ottawa-Carleton Co., Ottawa River, Deschenes Lookout. Prince Edward Co. Quebec: Berthierville; La Trappe; Thun der River (now Rivi?re-au-Tonnerre). Saskatchewan: 13 mi E Saska nd rd toon; Twp. 17, Range 29, W. 2 Meridian; Twp. 17, Range 1,W. 3 Meridian. Specimens in collections of: CASC, CNCI, DSC, EGRC, IRCW, JAC, KHSC, MCZC, OSUC, UAIC, UMIC, UNHC, USNM

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Amblyderus parviceps Casey Fig. 6,10

Amblyderus parviceps Casey 1895:744. Type locality: southern California. Lecto type male here designated, // SCal/ male/ Casey bequest 1925/ TYPE USNM 36618/ parviceps Csy/ LECTOTYPE Amblyderus parviceps Casey a des. DSChandler '98// [USNM]. A single male with similar locality label a is designated as PARALECTOTYPE. Pic 1911c:27. Leng 1920:163. Werner (synonymy) 1975:290. Poole and Gentili 1996:50. near Amblyderus gracilentus Casey 1895:745. Type locality: San Francisco [San Leandro], California. Holotype male, / / Cal. [with vertical line through the = "C" San Leandro, Alameda Co.]/ male/ Casey bequest 1925/ TYPE USNM 36619/ gracilentus Csy.// [USNM]. Pic 1911c:27. Leng 1920:163.

Diagnosis.? Smallest species (2.5-3.6 mm); pronotum with lateral margins evenly converging; setae appressed over body; apical third of to acute aedeagus evenly tapering point; sand dunes of Pacific Coast. Description.? Length: 2.5-3.6 mm. Body varying from all light to or abdomen orange-brown elytra partially entirely brown, usually brown, yellow-brown in the lightest specimens; dorsum with surface smooth between punctures and granules, faint microreticulation evi dent on some setae on on specimens; appressed dorsum, dense elytra, 0.08 on long elytra. Head smoothly roughened, granules difficult to distinguish; head base with medial cleft barely indicated, vertexal angles narrowly at about 85 antennomeres IX-X about as as rounded, angles ?; long wide in dorsal view, rest elongate, X sometimes wider than long; gula with at most two setae little more than half long setae, eye length. Pronotum in dorsal view with lateral convex margins slightly and converging toward base, lacking basal neck; disc with small granules most distinct near anterior indistinct mar margin, posteriorly; lateral in a gins dorsal view with few short erect tactile setae, setae 0.02 mm long; anterior face of pronotum with setae appressed; pronotal disc with shallow median at shallowly convex, impression apex; prosternum a with few setae little more than half eye length. with some setae at Elytra suberect humeral angles, tactile setae along lateral margins not visible in dorsal view; elytral apices each broadly rounded to subtruncate. Males with mesal margins of protibiae shallowly concave through most of length; sternite VIII slightly bi-emarginate at broad apex; with lateral to an aedeagus margins evenly narrowing acute apex. Biology.? Found in sand beneath low, spreading plants on dunes set back from the ocean. Adults have been taken in all months of the year. A small series of specimens from Pasadena (MCZC) and one from Pomona (CASC) represent the only localities not close to the seashore. Closest to A. obesus in Relationships.? sharing the indistinct gran ules of the head and or pronotum, separately rounded subtruncate elytral apices, shallow median indentation of the head vertex, and the

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lateral margins of the aedeagus narrowing apically. Both species share with A. setae of anterior pallens the appressed the face of the pronotum, and the shallow median of the impression pronotal apex. Separated from A. of a basal on obesus by the lack neck the pronotum, smaller size, and the the to lateral margins of aedeagus evenly converging the apex. Distribution.? Coastal sand dunes of California from just north of San Francisco to San Diego. UNITED STATES. California: Alameda San Leandro. Los El Co., Alameda; Angeles Co., Segundo sand dunes; Hermosa; Naples; Playa del Rey; Pasadena; Pomona; Redondo; Santa Monica. Monterey Co., Asilomar; Carmel; Del Monte; Little Sur Creek beach, 7 mi NW big Sur; Moss Landing; Pacific Grove; Pismo Beach; Point Sur; Seaside dunes. San Diego Co., Imperial Beach; San Diego. San Francisco Co. San Luis Obispo Co., Dune Lakes, 3 mi S Oc?ano; Morro Bay State Park; Pismo Beach. Santa Barbara Co., Carpinter?a; Santa Barbara. San Mateo San San Pedro Co., Mateo, Valley. Sonoma Co., Jenner; mouth of Russian River. Ventura Co., San Buenaventura State Beach; Point Mugu Naval Air Station; Ventura Beach; Zuma Beach. Specimens in collections of: CASC, CDAE, DSC, LACM, MCZC, USNM.

Amblyderus triplehorni Weissmann and Kondratieff Figs. 1, 7, 9

Weissmann and Amblyderus triplehorni Kondratieff 1999:138. Type locality: Alamosa Colorado, Co., Great Sand Dunes National Monument. Holotype male, USNM [not examined].

Diagnosis.? Largest species (4.8-6.1 mm); pronotum with lateral convex to on margins slightly base; setae suberect anterior face of erect to suberect lateral pronotum, along elytral margins; pronotal disc with lateral strongly convex; aedeagus margins evenly converg to rounded ing narrowly point; yellowish-brown; southern Colorado. 4.8-6.1 mm. to Description.? Length Body yellow orange-brown, sometimes with large undefined brown blotch centered at middle of each elytron; dorsum lightly reticulate, faintest on head and prono dorsum with setae 0.1 mm tum; appressed, elytra densely setate, long. Head vertex with coarsely granulate; base with shallow medial vertexal cleft; angles narrowly rounded, at about 85 ?; antennomeres VII about as as VIII-X long wide, wider than long, rest elongate; gula setae. lacking long Pronotum with lateral margins shallowly convex to base; disc with at granules largest middle; lateral margins with many short and long raised setae visible in dorsal view, setae longest anteriorly, longest setae from 0.20-0.28 mm varying long; anterior face of pronotum with dense suberect disc setae; pronotal strongly convex; prosternum with six or more long setae. with setae on disc some setae on Elytra appressed, erect lateral at humeral more mov margins angles, becoming gradually depressed ing posteriorly along lateral margins; elytral apices broadly

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subtruncate. Males with protibiae strongly sinuate on mesal margins; sternite VIII broadly and shallowly emarginate at apex; aedeagus with lateral to Weissmann and margins evenly converging pointed apex. Kondratieff (1999) note that males are typically darker than females. Biology.? Adult behavior is covered by Weissmann and Kondratieff (1999). Adults come out of the sand early in the evening to feed on dead insects just below the dune crest or in debris pockets lower down on are most common in the dunes. Adults early summer, but may be active until September. This is not close to of the Relationships.? species particularly any other species. It is readily distinguished by the strongly convex pronotal setae on the anterior face of the disc, suberect pronotum, strongly granulate head and pronotum, and the aedeagal apex is lengthily pointed. known from Great Sand Dunes National Monu Distribution.? Only ment in Colorado. Additional records to those in Weissmann and Kondratieff (1999): Great Sand Dunes, VI-1964, R. Lenczy; Great Sand Dunes E. C at National Monument, 7600', VII-24-1968, Becker, night on near same mi NE sand scurf pea; locality, 25 Alamosa, VIII-14-1966, K. same T. area. S. Hagen; locality, VIII-18-1965, C. Emmel, sand dune Specimens in collections of: CASC, CNCI, LACM, UAIC, USNM.

Amblyderus werneri Weissmann and Kondratieff Figs. 8,10

Amblyderus werneri Weissmann and Kondratieff 1999:141. Type locality: Colo rado, Alamosa County, Great Sand Dunes National Monument. Holotype male, USNM [not examined].

in Diagnosis.? Moderate size (3.7-3.9 mm); pronotum with lateral to disc margins evenly converging base; pronotal shallowly convex; setae suberect on anterior face of erect to pronotum, suberect along lateral elytral margins; aedeagus with short, broad point on broadly rounded apex; yellowish; southern Colorado. mm. Description.? Length 3.7-3.9 Head and pronotum orange to light yellow-brown, elytra similarly colored to largely dull brown, abdomen orange to brown; dorsum lightly reticulate, reticulation on setae over most of strongest elytra; appressed dorsum, elytra setae 0.1 mm on densely setate, long elytra. Head with granules most distinct lateral to midline, base with distinct medial cleft; vertexal angles narrowly rounded, angles at about 85?; antennomeres X slightly longer than wide, rest clearly with 0-1 setae. elongate; gula long Pronotum over evenly granulate disc; lateral margins strongly convex in anterior half, evenly converging toward base posteriorly, slightly curved laterally at base, lacking neck at base; lateral margins with setae visible clearly granulate, long along anterolateral margins in dorsal setae 0.16-0.20 mm view, longest long; anterior face of

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to disc pronotum with setae suberect rarely depressed; pronotal shal at lowly convex, with faint median impression apex sometimes lack ing; prosternum with several long setae. with setae on disc some Elytra appressed; humeral angles with suberect setae visible in dorsal view, scattered suberect setae visible along lateral margins clearly visible in dorsal view; elytral apices broadly subtruncate. Males with sinuate on protibiae strongly mesal margins; sternite VIII broad, shallowly trisinuate; aedeagus broadly rounded at apex, with median point. Biology.? Most of the specimens known were taken with pitfall or were to a in traps, attracted mercury vapor lamp sand dunes. These were more specimens taken along with similarly sized A. pallens in areas to the central dune two peripheral mass, while these species do not seem to occur in the central dune mass where the much larger triplehorni is found (M. Weissmann, pers. comm.). The dunes where werneri were taken have scattered on specimens typically plants them, and be stabilized. Adults are active in summer. may partially early Relationships.? It is similar to A. triplehorni in sharing the suberect on setae the anterior face of the pronotum and on the lateral margins of the to A. elytra and pronotum, and similar pallens in the antennomeres all longer than wide, and the aedeagus broad to the apex with a median point. This species may be separated from A. triplehorni by the convex evenly pronotal disc, subtruncate elytral apices, and the lack of a basal pronotal neck. Distribution.? Only known from Great Sand Dunes National Monu ment in Colorado.

Specimens examined.?14: UNITED STATES. Colorado, Alamosa Co., Great Sand Dunes National Monument, 25 mi NE Alamosa, VI-18 1983, T. P. Sluss (CASC); Meda?o Ranch, interdunal wetland, VI-17/ 20-1998, P. M. Piedra and C Cordova, pitfall (DSC); Main Sand Mass, VI-23/25-1998, P. M. Pineda (DSC).

Checklist of the World species of Amblyderus LaFert?-S?nect?re

A. aspericollis Fairmaire, 1897:116. Madagascar. A. bigibber Bonadona, 1986:62. Sri Lanka. A. brincki Bonadona,1986:61. Sri Lanka. A. brunneus Pic, 1893:15. Italy (Calabria, Sicily). A. concolor Pic, 1933:6. India "m?ridionale." A. coronatus Pic, 1913:381. Namibia. A.fasciatus Pic, 1933:6. India "m?ridionale." A., granularis (LeConte), 1850:231. United States, Canada (eastern North America). 1895:747 punctiger Casey (synonymized by Werner, 1975). A. indicus Fairmaire, 1894:33. India (Bihar). A. latipennis Pic, 1898a:69. Eritrea. A. laxatus Bonadona, 1964:228. Chad (Tibesti), Saudi Arabia. A. longidentatus Pic and Hawkins, 1957:436. Yemen (Aden). A. longipilis Pic, 1931:11. Djibouti.

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A. macultpennts Pic, 1898b:181. Somalia, Eritrea, Ghana, Togo, Nigeria, Tanzania. A. mitis Hille, 1971:378. South Africa. A. obesus Casey, 1895:743. United States, Canada (west coast). albicans Casey 1895:744 (synonymized by Werner, 1975). A. obscuripennis Pic, 191 lb: 157. Uganda (North Buganda, Buvuma Island). new A. owyhee Chandler, species. United States, Canada (Columbia River Plateau). A. pallens (LeConte), 1850:231. United States, Canada. arenarius Casey 1895:748 (synonymized by Werner, 1975). A. parviceps Casey, 1895:744. United States (west coast). gracilentus Casey 1895:745 (synonymized by Werner, 1975). = A. sabulosus Pic, 1899:173. Israel (Jaffa Tel Aviv). A. scabricollis LaFert?-S?nect?re, 1847:368. Algeria, Spain, Corsica, Sardinia, Italy, Sicily. inhumeralis Pic, 1900:89 (as variety). A. scabridus (Krekich-Strassoldo), 1931:15. India (Orissa). A. spiniger Motschulsky, 1863:490. Sri Lanka. A. sulcithorax Pic, 1898a:69. Somalia, Djibouti, Yemen. A. tavetanus Pic, 1914:171. Kenya (Coast Province). A. thoracicus Bonadona, 1989:256. Sri Lanka. thoracinus Bonadona, 1989:256, lapsus. A. triplehorni Weissmann and Kondratieff, 1999:138. United States (Colo rado). A. truncatus LaFert?-S?nect?re, 1849:1. Egypt, Somalia. A. tuberculatus Krekich-Strassoldo, 1928:88. India (Orissa). A. villiersi Pic, 1950:151. Niger (Air). A. werneri Weissmann and Kondratieff, 1999:141. United States (Colo rado).

ACKNOWLEDGEMENTS

Iwould like to thank all of the curators and their assistants, and collectors who have sent me specimens over the years, and who are listed with the collections and acronyms earlier. Michael A. Ivie, University of Montana, provided tremendous help in resolving the misidentification of a bruchine beetle as an Amblyderus from Puerto Rico. Tess Feltes provided the habitus illustration of on a Amblyderus triplehorni, which was based specimen taken from a series of this to me lengthy species given by Charles A. Triplehorn (Ohio State Mike Weissman and Boris University). Kondratieff kindly made a gift of a nice series of Amblyderus werneri. John F. Burger and Paul C. Johnson, both of New Gerhard University Hampshire, Uhmann (Pressath, Germany), and Darren Pollock of (University Manitoba, Winnipeg, Canada) are thanked for the Gianluca Nardi reviewing manuscript. (Cisterna, Italy) graciously pro vided copies of some literature that I lacked.

LITERATURECITED

Blatchley, W. S. 1910. On the Cole?ptera known to occur in Indiana; an illustrative of the or descriptive catalog Cole?ptera (exclu sive of the occur Rhynchophora) known to in Indiana with bibliog and of new raphy descriptions species. The Nature Publishing Company, Indianapolis, Indiana, 1386 pp.

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P. sur Revue Bonadona, 1964. Note les Anthicidae Pal?arctiques. 31:225-249. fran?aise d'Entomologie Bonadona, P. 1986. Anthicidae (Cole?ptera) de Sri Lanka. Entomol?gica 30:55-75. Scandinavica Supplement nouveaux Bonadona, P. 1989. Anthicidae (Cole?ptera) des collections du Mus?um d'histoire naturelle de Gen?ve. Revue Suisse de Zoologie 96:253-276. Y. I. Bousquet, and Bucciarelli, 1980. Cole?ptera Anthicidae. Fauna d'Italia. Volume 17. Edizioni Calderini, Bologna, viii + 240 pp. Bousquet, Y. 1991. Family Anthicidae, antlike flower beetles, pp. 271 In Y. 273. Bousquet, ed., Checklist of beetles of Canada and Alaska. Agriculture Canada Research Branch, Publication 1861/E. vi + 430 pp. Casey, T. L. 1895. Coleopterological notices, VI. Annals of the New York Academy of Sciences 8:435-838. Downie, N. M. and R. H. Arnett, Jr. 1996. The beetles of North America. Volume II. : Series through x + Curculionoidea. Sandhill Crane Press, Gainesville, FL. pp. 891 1721. Fairmaire, M. L. 1894. H?t?rom?res du Bengale. Annales de la Soci?t? Entomologique de Belgique 38:16-43. Fairmaire, M. L. 1897. Mat?riaux pour la faune Col?opt?rique de la r?gion Malgache. Annales de la Soci?t? Entomologique de Belgique 41:92-119. Hille, J.C. van. 1971. Anthicidae (Cole?ptera) from northern Zululand. Transactions of the Royal Society of South Africa 39:367-391. H. G. and E. A. Schwarz. 1878. The of Hubbard, Cole?ptera Michigan. Proceedings of the American Philosophical Society 17:593-669. Jacquelin du Val, C. 1863. Pp. 353-418. In Jacquelin du Val, C and L. Fairmaire. Manuel Genera des Entomologique. Col?opt?res en d'Europe comprenant leur classification familles naturelles, la description de tous les genres, des tableaux synoptiques destin?s ? faciliter le toutes l'?tude, catalogue de les esp?ces, de nombreux dessins au trait de caract?res. Vol. 464 A. 3, pp. Paris, Deyrolle. Kingsolver, J.M. 1970. A synopsis of the subfamily Amblycerinae Bridwell in the West Indies, with descriptions of new species (Cole?ptera: Bruchidae). Transactions of the American Entomo logical Society 96:469-497. Krekich-Strassoldo, H. 1928 (1926). Beitr?ge zur Kenntnis indischer Anthiciden. Archiv f?r Naturgeschichte 92:67-105. Krekich-Strassoldo, H. 1931. Beitr?ge zur Kenntnis indischer Anthiciden II. Folia Zool?gica et Hydrobiologica 3:1-41, plates 1-3. F. T. de.1847. 32. LaFert?-S?nect?re, [Anthicidae] pp. 365-381, pl. In, H. Lucas, Exploration scientifique de l'Alg?rie pendant les Ann?es 1840,1841,1842. Histoire naturelle des animaux articul?s. Deuxi?me partie, Insectes. Sciences Physiques Zoologie, Volume II, 590 pp. A. Bertrand, Paris. LaFert?-S?nect?re, F. T. de. 1849a (1847). Livraison No. 6, Amblyderus, 3 1 In: ed.: pp., plate. Gu?rin-M?neville, M.F.E., Species et

This content downloaded from 132.177.228.65 on Fri, 15 Aug 2014 15:39:11 UTC All use subject to JSTOR Terms and Conditions 292 REVISIONOF AMBLYDERUS

des animaux articul?s ou iconographie g?n?rique repr?sentation avec et toutes de des genres, leur description celle de les esp?ces cette du . Premi?re Partie: Insectes grande division r?gne Fain et Paris. Col?opt?res. Thunot, LaFert?-S?nect?re, F. T. de. 1849b (1848). Monographie des Anthicus et genres voisins, Col?opt?res H?t?rom?res de la tribu des Trach?lides. De Sapia, Paris; xxii + 340 pp, pis. 17-32. LeConte, J. L. 1850. General remarks upon the Cole?ptera of Lake 201-242. In: L. R. Lake its Superior, pp. J. Agassiz. Superior: physi and with those of cal character, vegetation, , compared other and similar regions. Gould, Kendall & Lincoln, Boston. 428 pp. LeConte, J. L. 1852. Synopsis of the Anthicites of the United States. Proceedings of the Academy of Natural Sciences of Philadelphia 6:91-104. LeConte, J. L. 1866. List of the Cole?ptera of North America prepared for the Smithsonian Institution. Part I.No. 140, pp. 50-70 (pp. 1-49 published 1863). Leng, C. W. 1920. Catalogue of the Cole?ptera of North America, north x + 470 of Mexico. Cosmos Press, Cambridge, Mass., pp. Motschulsky, V. 1863. Essai d'un catalogue des Insectes de l'Ile Ceylan. (Suite). Bulletin de la Soci?t? Imp?riale des Naturalistes de Moscou 36:421-532. M. 1893. Anthicides nouveaux. Revue Linn?enne 9:15 Pic, L'?change, 16.

Pic, M. 1898a. Anthicides (Col. H?t?rom?res) africains nouveaux des collections du Mus?um de Paris. Bulletin du Mus?um d'Histoire Naturelle, Paris 4:67-72. M. 1898b. Un nouveau Pic, Amblyderus [Col. H?t?rom?res] d'Abyssinie. Bulletin du Mus?um d'Histoire Naturelle, Paris 4:181. Pic, M. 1899. Xylophilides et Anthicides. Recueillis en Orient, en mars, avril et mai 1899. Revue scientifique du Bourbonnais et du Centre de la France 12:170-178. Pic, M. 1900. Diagnoses de Col?opt?res du globe. L'?change, Revue Linn?enne 16:89-91. M. 1911a. nouveaux ou connus Pic, Col?opt?res exotiques peu (Suite). Revue Linn?enne 27:132-134. L'?change, Pic, M. 1911b. Collezioni zoologiche fatte nell'Uganda dal Dott. E. Bayon. XIII. Anthicidae (Col. H?t?rom?res). Annali del Museo Civico di Storia Naturale di Genova 45:156-159. Pic, M. 1911c. Anthicidae, Pars 36. Ed. S. Schenkung, Coleopterorum W. Catalogus. Junk, Berlin. 102 pp. Pic, M. 1913. Neue Anthicidae aus Afrika (Col.). Deutsche Entomologische Zeitschrift, pp. 377-382. M. Pic, 1914. Voyage de Ch. Alluaud et R. Jeannel en Afrique Orientale (1911-1912). Insectes Col?opt?res. IV. Hylophilidae et Anthicidae, IL A. pp. 155-194, pi. Schulz, Paris. M. 1931. Nouveaut?s diverses. Pic, M?langes exotico-entomologiques 58, 36 pp.

This content downloaded from 132.177.228.65 on Fri, 15 Aug 2014 15:39:11 UTC All use subject to JSTOR Terms and Conditions DONALD S. CHANDLER 293

Pic, M. 1933. Nouveaut?s diverses. M?langes exotico-entomologiques 62, 36 pp. Pic, M. 1950. Contribution a l'?tude de l'A?r (Mission L. Chopard et A. Malacodermoidea et Heteromeroidea. Villiers). Col?opt?res M?moires de l'Institut fran?aise d'Afrique Noire 10:145-153.

Pic, M. and C N. Hawkins. 1957. 29. Cole?ptera: Anthicidae, pp. 435 450. In: British Museum (Natural History) Expedition to South west Arabia 1937-8. Vol. 1.

Poole, R. W. and P. Gentili. 1996. Nomina Insecta Nearctica. A check list of the insects of North America. Volume 1: Cole?ptera, Information MD. Strepsiptera. Entomological Services, Rockville, 827 pp. Ronchetti, L., I. Colombini, and L. Chelazzi. 1986. Researches on the coast of Somalia. The shore and the dune of Sar Uanle. 40. Anthicidae (Cole?ptera). Monitore Zool?gico Italiano, N. S. Supplemento XXI, pp. 169-184. Uhmann, G. 1976. Die Gattungen der Anthicidae und ihre systematische Anordnung. Entomologische Bl?tter 72:166-182. Uhmann, G. 1978. Die Gattungen der Anthicidae und ihre systematische Anordnung. Teil 2. Die systematische Anordnung. Entomologische Bl?tter 74:75-80. Werner, F. G. 1964. A revision of the North American species of s. str. Anthicus, (Cole?ptera: Anthicidae). Miscellaneous Publica tions of the Entomological Society of America, 4(5): 195-242. Werner, F. G. 1975. New synonymy in the Nearctic Anthicidae (Co le?ptera). Proceedings of the Entomological Society ofWashington 77:290.

Werner, F. G. 1983. Anthicidae of the Greater Antilles, and a new species from Venezuela (Cole?ptera). Psyche 90:211-235. M. and B. Kondratieff. 1999. Two new Weissmann, J. C. species of Amblyderus from Great Sand Dunes National Monument, Colorado (Cole?ptera: Anthicidae). Entomological News 110:137-143. G. N. 1936. Wolcott, "Insectae Borinquenses." A revised annotated check-list of the insects of Puerto Rico. Journal of Agriculture of the University of Puerto Rico 20:1-600. Wolcott, G. N. 1951 (1948). The insects of Puerto Rico. Cole?ptera. Journal of Agriculture of the University of Puerto Rico 32:225-416.

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