Carpophilus hemipterus (Linnaeus, 1758)

Ewing, Curtis P. & Cline, Andrew R., 2005, Key to Adventive Sap Beetles (Coleoptera: Nitidulidae) in Hawaii, with Notes on Records and Habits, The Coleopterists Bulletin 59 (2), pp. 167-183 : 167-183

C. hemipterus , C. humeralis , C. dimidiatus , S. geminata , A. concolor , C. mexicanus , and the three Epuraea species , occur in native and disturbed forests above 350 meters.

Methods and Materials

External features were examined using a Wild M5 stereo-microscope. If ventral surfaces were obscured, specimens were remounted to expose the thoracic and abdominal sterna. All characters are external and visible at 503 with the exception of microsculpture which requires 1003 magnification. Alternate characters are provided for three species for which microsculpture is included. Figures were prepared using a reticule and grid for initial sketches and Adobe Illustrator 10.0 for final plates.

Specimens Examined. Beetles were examined from the following collections: Cornell University Insect Collection (CUIC), Bishop Museum Insect Collection (BPBM), Hawaii Department of Agriculture (HDOA). Cybocephalus nipponicus

Endrödy-Younga —OAHU: Pawaa, 25.viii.1998, on hibiscus inf. w/ Pinnaspis strachani, M.E. Chun /98-276 (HDOA). Conotelus mexicanus Murray —OAHU: Waimea, 10 Aug. 1981, ex. Hibiscus fls., coll. G. Powell (HDOA). Aiea, 3-54, Ford (BPBM). MAUI: Kula, 22 July 1982, ex. Crysanthemums, coll. N. Miyahira (HDOA). HAWAII: Kona dist., Kaupulehu For. Res., 600 m, 24.III.1961, Ipomoea, L.W. Quate coll. (BPBM). Carpophilus (Urophorus) humeralis (F.) —KAUAI: NaPali-Kona For. Res., Kukui tr., 20-III-1991, el. 300–330 m, J.K. Liebherr, ex. rotten Passiflora (CUIC) . OAHU: Waianae Mts., Puu Kaua, 4-V-1995, lot02, 940–948 m, scraping moss on ohia trunks in day, J.K. Liebherr (CUIC). Waianae Mts., Mt. Kaala, Dupont tr., 16- XI-1993, 1,150 m, ex. Freycinetia, D.A. Polhemus (CUIC) . Mt. Kaala, 17-X-1992, M.J. & C.A. Tauber (CUIC). Light trap, John Rodgers, 6-58, E.J. Ford Jr. (BPBM). Puu Palikea, 3-58, E.J. Ford Jr. coll. (BPBM). Hon. 1924, Cartwright coll. (BPBM). Hon. 3 Aug. 49 (BPBM). Waikiki, VII-13-27, at light, E.H. Bryan coll. (BPBM). Insectary reared, Hon., 10 July 1977, x. pineapple, SIHAU (HDOA). MOLOKAI: Olokui, 25-X-1991, el. 1,320 m, R.G. Gillespie, ex. for. litter (CUIC). Nr. edge of Waiukolu Val., X-10-1943, N.L.H. Krauss (HDOA). LANAI: Haalelepaakai, Munroe tr., 6.0 mi, beating veg., 2-V-1993, lot 02, el. 1,025 m, J.K. Liebherr coll. (CUIC). MAUI: Puu Kukui, tr., at Puu 45039, 22-V-1997, lot08, 1,330 m el., sift moss on ohia, J.K. Liebherr (CUIC). Puu Kukui tr., E. Kualalewelewe Cabin, 14-V-1992, el. 890– 1,060 m, D.A. Polhemus (CUIC). Mt. Eke NAR, 24-V-1997, lot07, 1,360 m el., pyrethrum fog of ohia, J.K. Liebherr (CUIC). HAWAII: Keeau, 14 Dec. 1976, ground coconut husk, T. Lindsey (HDOA). Carpophilus hemipterus (L.) —KAUAI: Haena, I- 7-1944, N.L.H. Krauss (HDOA). NaPali-Kona For. Res., Kukui tr., 20-III-1991, el. 300–330 m, J.K. Liebherr, ex. rotten Passiflora (CUIC) . OAHU: Wahiawa, 31-V-09, J. Kotinsky coll., 933 (HDOA). light trap, John Rodgers, 6.58, E.J. Ford, Jr. coll. (BPBM). Manoa, 6-19-29, N.H. Krauss coll. (BPBM). Barbers Pt., 4-49, coll. Ford (BPBM). Barbers Pt., 6-49, coll. Ford (BPBM). Mt. Kaala, 18-V-1991, el. 1,210 m, J.K. Liebherr, beating ohia & tree ferns (CUIC). Waianae Mts., Puu Kaua, 4-V-1995, lot01, 940–948 m, beating ohia in day, J.K. Liebherr (CUIC). Waianae Mts., Puu Kaua, 4-V-1995, lot01, 940–948 m, beating ohia in day, J.K. Liebherr (CUIC). LANAI: 8-30- 27, pineapple fields, J.F. Illingworth coll. (BPBM). Haalelepaakai, Munroe tr., 6.0 mi, beating veg., 2-V-1993, lot02, el. 1,025 m, J.K. Liebherr (CUIC). MAUI: Makawao, 11-29-66, N. Miyahira (HDOA). Puu Kukui tr., at Puu 4503, 13-V-1992, el. 1,330 m, J.K. Liebherr (CUIC). Carpophilus ( dimidiatus ) dimidiatus (F.) —KAUAI: K-84-7, Anahola, 11-XI-84, ex. peanut in shell, D. Sugawa (HDOA). Na Pali-Kona For. Res., Kukui tr., 20-III-1991, el. 300–330 m, J.K. Liebherr, ex. rotten Passiflora (CUIC) . OAHU: Hon. 9-51, coll. Ford (BPBM). Kawaiai, 8-54, E.J. Ford coll. (BPBM). Hon., T.H., IX-37, ex. rice E.C. Zimmerman coll. (BPBM). Hon. T.H. VI-19-38, ex. rice,

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E.C. Zimmerman coll. (BPBM). Barbers Pt., 4-60, E. J. Ford Jr. coll. (BPBM). HAWAII: Hilo, 2-25-80, Macadamia nut, S. Matayoshi (HDOA). Hilo, III-20-69, ex. Kukui nut, Ernest Yoshioka coll. (HDOA). Carpophilus ( dimidiatus ) freemani

Dobson — OAHU: Barbers Pt. , 6-49, coll. Ford ( BPBM) . Honolulu , 5-1922, Hawaiian Ids. ( CUIC) . DELAWARE: Smyrna , 7 Oct. 1952, W. A. Connell, weevil infested acorns, Ideotypes #, $, R.M. Dobson det. 18/XII/1956 ( CUIC) . Carpophilus

( dimidiatus ) maculatus Murray —OAHU: Moanalua, 3-19-23, J.F. Illingworth coll. (BPBM). Oahu S. Co., IV-05, C.I.E. coll., A. 1810, 75-5 (HDOA). HAWAII: Kona, 2,000 ft [610 m], Perkins, IX-1892 (BPBM). Kona, 2,500 ft [762 m], Perkins, VI-1892 (BPBM). Napoopoo, 8-10-19, O.H. Swezey coll. (BPBM). Carpophilus ( dimidiatus )

mutilatus Erichson — KAUAI: Mana , May 1984, ex. seed corn, coll. S. Masukawa ( HDOA) . Kekaha , 8-vii-1988, ex. sunflower plants, R. Oyama / Koa 88-18 ( HDOA) . NaPali-Kona For. Res. , Kukui tr., 20-III-1991, el. 300–330 m, J.K. Liebherr, ex. rotten Passiflora (CUIC) . OAHU: Barbers Pt., 4-68, E.J. Ford Jr. coll. ( BPBM) . Barbers Pt. 4- 52, coll. Ford ( BPBM) . Barbers Pt. , 11-59, E.J. Ford Jr. coll. ( BPBM) . Schofield , 4-60, E.J. Ford Jr. coll. ( BPBM) . MOLOKAI: Kaunakakai , 15.i.90, field #2, on corn silk inside ear shoot, P. Eichhom /90-030 ( HDOA) . Carpophilus marginellus Motschul-

sky —OAHU: Tantalus, 3-49, coll. Ford (BPBM). Wahiawa, 4-58, E.J. Ford, Jr. coll. (BPBM). same, 5-58. same, 7-52. light trap, John Rodgers, 5-58, E.J. Ford, Jr. (BPBM). Carpophilus oculatus Murray —OAHU: HIA, 29 Mar. 1978, pineapple trap, W. Nagamine (HDOA). HIA, 6 Sept. 1978, light trap, D. Henderson (HDOA). PONAPAE Is. (¼ Pohnpei, Caroline Islands): Colonia, I-13-1953, J.F.G. Clarke, rf. Pithecolobium duleis (BPBM) . Epuraea (Haptoncus) luteola Erichson —OAHU: Ewa, 19 Dec. 1973, in raisin trap, G.Y. Funasaki (HDOA). Nanakuli, 30, Aug. 1968, in honeydew melon, G.Y. Funasaki (HDOA). Hon. Airport, 1-53, coll. Bixby, org. Fiji (BPBM). Barbers Pt., 10-5-49, coll. Ford (BPBM). Schofield, Jun. 90, K. Will (CUIC).

Epuraea (Haptoncus) munda (Sharp) — KAUAI: Perkins, (771 under card) ( BPBM) . OAHU: Barbers Pt., 6-49, coll. Ford ( BPBM) . Barbers Pt., 10-4-49. light trap, John Rodgers, 5-58, E.J. Ford, Jr. ( BPBM) . Waipio, T.H. light trap, 9-57, E.J. Ford, Jr. ( BPBM) . Ewa, 19 Dec. 1973, in raisin trap, G.Y. Funasaki ( HDOA) . Epuraea

(Haptoncus) ocularis Fairmaire —OAHU: Poamoho tr., 9-III-61, L.W. Quate coll. (BPBM). Pupukea tr., III-32, O. Bryant coll. (BPBM). Koolau Ra., Ahuimanu Rd., 120 m, 2-II-1984, swarming, 0715-0745 hls, G.A. Samuelson coll. (BPBM). Halawa, XII- 10-70, light trap, W. Au coll. (HDOA). HAWAII: Waimanu Val., W. bank Waimanu str., 8.v.1986, site no.1, nr. river mouth, jetsam on boulders, S.M. Gon & J. Heer colls., Acc. #1987.051 (BPBM). Upper Hamakua, ditch tr., VIII-15-35, R.L. Usinger coll. (BPBM). Aethina (Idaethina) concolor (Macleay) —KAUAI: Kealoa lookout on Ipomoea pes-caprae , #366,7, M.A. Lachance coll. (BPBM). OAHU: Schofield, June 90, K. Will (CUIC). HAWAII: Kona, 30.v.1992, ex Coccinia fls., L. Doi/S. Matayoshi coll., 92-330 (HDOA). Kona 30.vii.1992, ex red hibiscus, S. Matayoshi/92-476 (HDOA). Omosita discoidea (F.) —MAUI: found in greasy can, Haleakala, Jl.20.919, gulch nr. Puu Nianiau, el. ab. 6,200 ft [1,890 m], Timberlake coll. (HDOA). Haleakala, Kalahaku, 9,0009 [2,743 m], VIII-1985 (BPBM). Stelidota geminata (Say) —OAHU: UH Manoa Campus, 2009 [61 m], 26 Jun. 1998, rotting tangerines on ground, C. Ewing coll., DNA vouch. #292 (CUIC). Waimanalo, 13.viii.1997, in fallen false kamani w/5 other nitidulid spp., M. Ramadan/97-330 (HDOA). MOLOKAI: Puu Lua, summit, 3,1809 [969 m], 16 Jun. 1999, sifting ohia litter, C. Ewing coll. # 98, 218069280N, 1568489480W (CUIC). Access rd. to Iliiliopae heiau, 10 Jun. 2004, 10 m, Passiflora mollissima fruit, C. Ewing coll. # 5, 218049430N, 1568489370W (CUIC). LANAI: Hauola ridge tr., 3,3609 [1,024 m], 15 Dec. 1998, C. Ewing coll. #24, sifting Pritchardia fronds and fruit, 208489330N, 1568529000W (CUIC). MAUI: Haleakala NP, Paliku cabin, el. 1,950 m, 208439160N, 1568089380W, 18-V-2001 lot03, diphacinone

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170 bait sta., Liebherr ( CUIC) . HAWAII: Kohala For. Res. , 1,122 m, 14 Oct. 1997, fogging M. polymorpha, D. Gruner & D. Polhemus colls., GRU0724-006 ( BPBM) , same, 12 Oct. 1997, GRU0584-006. Stelidota chontalensis Sharp —OAHU: Kapiolani Park, el. 5 ft [2 m], 11–12.I.1992, W.D. Perreira coll., Loise Dillingham Mem. Found., 92-357 ( BPBM) . same (HDOA). Kaimuki , el. 200 ft [61 m], BWS, 13-XI-1992, W.D. Perreira coll., stuck on apple maggot trap treated with TML ( BPBM) . Phenolia (Aethinodes)

attenuata (Reitter) —KAUAI: Princeville, night at hotel light, 02.viii.2000, via R.T. Furumizo (BPBM). OAHU: U.H. Manoa campus, 2009 [61 m], 26 Jun. 1998, rotting tangerines on ground, C. Ewing coll. DNA vouch. #284 (CUIC). U.H. Manoa, 18:VI.98, ex. rotting fruits, W. Nagamine (HDOA). Waimanalo, 7.vi.1998, ex ripe bananas, L. Nagasawa/98-144 (HDOA). MOLOKAI: Access rd. to Iliiliopae heiau, 10 Jun. 2004, 10 m, Passiflora mollissima fruit, C. Ewing coll. # 5, 218049430N, 1568489370W (CUIC). Phenolia (Lasiodites) limbata tibialis (Boheman) —OAHU: UH Manoa Campus, 2009 [61 m], 26 Jun. 1998, rotting tangerines on ground, C. Ewing coll., DNA vouch. # 285 (CUIC). Manoa, UH, 8.vi.1998, on mandarin oranges, W. Nagmine/L. Nagasawa/98-146 (HDOA). Mt. Tantalus (Manoa cliffs tr.), XI-1-00, M.J. & C.A. Tauber colls. (CUIC). MOLOKAI: Access rd. to Iliiliopae heiau, 10 Jun. 2004, 10 m, Morinda citrifolia fruit, C. Ewing coll. # 5, 218049430N, 1568489370W (CUIC). MAUI: Kahalui airport, low spot nr. wetland # 3, 208549260N, 1568259520W, 29.III.2000, MV light #2, Keawe woodland, keawe, Sesuvium , Paspalum , Pluchea, F.G. Howarth, D.J. Preston, G.A. Samuelson, K. Martz, F. Starr colls. (BPBM). HAWAII: Waikea, 30.V.96, ex. guava, H. Hirae (HDOA).

Defining Characters of Adventives and Endemics. Endemic Hawaiian Nitidulidae are morphologically diverse and difficult to separate from some of the adventives. Conotelus mexicanus is contained in the same subfamily, e.g., Cillaeinae , as all of the endemics ( Kirejtshuk 1986), and shares many character states with them. The form of the abdominal paratergites most clearly separates C. mexicanus from the endemics. The Carpophilus species are the most difficult adventives to identify and much of the following discussion of characters concerns them. In addition to the adventives, four endemics are illustrated. The habitus illustrations of Cillaeopeplus staphyliniformis Ford , and Gonioryctus arduus Sharp illustrate the range of body forms ( Figs. 3, 4 ), the labrum of Apetasimus pleomelarrosus (Ewing, manuscript name) is the most incised of the endemic forms ( Fig. 14 ), and Prosopeus floricola (Blackburn) shows the relative lengths of the abdominal sternites ( Fig. 15 ). The morphological features relevant to the following key are here reviewed.

Body Form. Length variable between 1 and 10 mm, with most between 2.5 and 4 mm. Body shape varies dorsally from ovoid to elongate, and laterally from convex to flattened.

Coloration. Coloration varies from relatively constant to highly variable. Carpophilus in the dimidiatus group (sensu Connell 1977) present in Hawaii (e.g., dimidiatus , freemani , maculatus , and mutilatus ), are particularly variable, with some individuals having distinct elytral markings and others nearly unicolorous. Teneral adult specimens usually have less pronounced or absent markings.

Head. The labrum is important for separating adventives from endemics ( Figs. 14, 16–18 ). In most endemics the labrum is truncate to slightly convex, lacks a prominent apical comb of setae, and has a small medial tooth. A few are emarginate with a moderately prominent medial setal comb, the most extreme example is illustrated ( Fig. 14 ). Nearly all adventives have an incised, bilobed labrum, one important exception being O. discoidea , which approaches the condition of some endemics but can be distinguished by the prominent setal comb continuing laterally ( Fig. 18 ). Carpophilus dimidiatus and C. freemani are distinguished by the form of the third and eighth antennomeres respectively. Carpophilus dimidiatus has the third antennomere

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staphyliniformis ; 4) Gonioryctus arduus ; 5) Aethina (Idaethina) concolor ; 6) Carpophilus

hemipterus; 7) Carpophilus oculatus ; 8) Carpophilus ( dimidiatus ) dimidiatus ; 9) Epuraea

(Haptoncus) ocularis, arrow indicates greatest length of elytron; 10) Omosita discoidea ;

11) Stelidota geminata ; 12) Stelidota chontalensis ; 13) Phenolia (Aethinodes) attenuata . mpt

medial margin of paratergites; v and vi, actual abdominal tergites 5 and 6; mp male pygidium; as

anal sclerite; fp female pygidium. Scale bars ¼ 1.0 mm.

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172 philus ( dimidiatus ) dimidiatus , labrum; 17) Aethina (Idaethina) concolor , labrum; 18) Omosita

discoidea , labrum; 19) Carpophilus marginellus ; 20) Carpophilus ( dimidiatus ) mutilatus ;

21) Phenolia (Aethinodes) attenuata ; 22) Stelidota geminata ; 23) Carpophilus hemipterus . i–v,

visible abdominal sternites 1–5; mes, metepisternal suture; axs axillary space; mcxl metacoxal lines

of first visible abdominal sternite; mic medial impunctate carina of mesosternum. Scale bars;

Figures 14, 16–18 ¼ 0.20 mm; Figures 15, 19–23 ¼ 1.0 mm.

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view showing triangular areas on visible abdominal sternite 1; 26) Carpophilus ( dimidiatus )

freemani , antenna; 27) Carpophilus ( dimidiatus ) dimidiatus , antenna; 28) Carpophilus

(Urophorus) humeralis, Prothorax ventral, prosternum shaded; 29) Epuraea (Haptoncus) munda ,

dorsal, arrow indicating greatest length of elytron; 30) Phenolia (Lasiodites) limbata tibialis ,

dorsal, elytra. viii, antennomere 8, iii, antennomere 3, ama, anterio-medial area; la, lateral area.

Scale bars; Figures 24, 25, 28–30 ¼ 1.0 mm; Figures 26, 27 ¼ 0.25 mm.

longer than the second ( Fig. 27 ), though depending on the viewing angle they may appear equal. Carpophilus freemani is distinguished by the width of the eighth antennomere which is 1/2 or more as wide as the ninth ( Fig. 26 ).

Pronotum. The anterior margin in dorsal view varies between convex, truncate and emarginate. The lateral margins in dorsal view are evenly curved to sub-parallel with posterior angles obtuse to rectangular. The convexity of the pronotum separates the Carpophilus species and A. concolor from the remaining species. Viewing the specimen anteriorly, the pronotum at the margin is either convex for its entire length ( Fig. 24 ) or concave laterally and explanate, at least posteriorly. Pronotal punctation can be highly variable within Carpophilus and important in separating species. It is difficult distinguishing between a pronotal disc that is deeply or shallowly impressed and densely or sparsely punctate with limited reference material. Only the least variable area, medial and anterior to the center of the disc, is referred to in this manuscript. The microsculpture between the pronotal punctures separates the three Epuraea species , though it can only be seen distinctly with a 1003 magnification. The pronotum of E. munda appears more shining than the other two at lower magnification, however the shape of the posterior margin of the elytron is a more definitive feature ( Fig. 29 ).

Prosternum. Punctation of specific areas of the prosternum separates Carpophilus species ( Fig. 28 ).

Scutellum. The shape of the scutellum separates C. mexicanus and the three Epuraea from the remaining species. The former species has the apex sub-truncate ( Fig. 2 ) and the latter have the sides nearly straight and the apex acute ( Figs. 9 , 29 ). The

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174 remaining species have the sides curved and the apex obtuse to broadly rounded ( Figs. 3–8, 10–13 ).

Elytra. The elytra are frequently shortened in sap beetles, and traditionally the number of exposed abdominal tergites have defined genera and subfamilies within Nitidulidae . Within the endemic forms there are species with three, two, and a few with only part of one abdominal tergite exposed. This character is extremely unreliable because the abdomen may distend and retract during preservation. It is included in the key as a secondary character. The arrangement of the punctation into striae, and elevation of the interstices to form costae are important for identifying species of Phenolia .

Mesosternum. The posterio-medial area of the mesosternum is densely and coarsely punctate in C. humeralis and C. hemipterus , and the punctures are interrupted by an impunctate longitudinal carina in the latter ( Fig. 23 ).

Metasternum. The axillary space is a triangular area ( Figs. 19–23 ) formed when the posterior margin of the mesothoracic coxal cavity diverges from the cavity and meets the metepisternal suture ( Fig. 19 ) posterior to the cavity.

Abdomen. The dorsal aspects of each segment are numbered from 1–6, the terminal tergite in females, and penultimate in most males, is the pygidium (¼ tergite 7) ( Fig. 9 ) (only tergites 5 and 6 and the pygidium are referred to in the key). Most males have a small, exposed eighth tergite, the anal sclerite ( Figs. 2, 3, 5, 9 ), and in females the eighth tergite is membranous and internal. The segments are numbered differently on the ventral surface. Actual sternites 1 and 2 are not visible hence the numbers of the visible sternites are two less than the equivalent actual dorsal sclerites. The visible sternites are numbered 1–5, with terminal sternite 5, the hypopygidium, being associated with the pygidium (tergite 7). The form of the paratergites is the most reliable character for separation of endemics from adventives ( Figs. 2–4, 7 ). In endemics paratergites 5 and 6 are broad (especially apically), flattened, and the medial margin is strongly sinuate ( Figs. 3, 4 ). In species with only the pygidium exposed it is necessary to lift an elytron to reveal this character, and if the abdomen is retracted the strongly angled basal portion may be hidden within the previous segment. Aethina concolor , both Stelidota species , and P. attenuata have metathoracic coxal lines on visible abdominal sternite 1 divergent and nearly reaching the posterior margin ( Figs. 21, 22 , 25 ).

Legs. Cybocephalus nipponicus (and all Cybocephalinae ) have a 4-4-4 tarsal formula, other Nitidulidae have 5-5-5. Males of Stelidota have the metathoracic femora and sometimes the mesothoracic femora modified, in some species it is enlarged distally and/or sinuate.

Key to Adventive Hawaiian Nitidulidae

1) Body length small,; 1 mm; hemispherical, head deflexed, not visible from above; capable of rolling into a ball; shining with elytra black in both sexes, female with pronotum and head black, males with pronotum and head pale brown; setae and punctation fine and inconspicuous; tarsi 4-4-4; ( Fig. 1 ) - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Cybocephalus nipponicus Endrödy-Younga

1 9) Body length larger,.1.8 mm; never hemispherical, head visible from above; color and setae variable; tarsi 5-5-5; ( Figs. 2–13 ) - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 2

2) Body form elongate, narrow, convex, and attenuate posteriorly; scutellum transverse with apex sub-truncate; dorsum dark brown to black, sometimes elytra light; three abdominal terga exposed; pygidium elongate; ( Fig. 2 ); length 3.2–4.0 mm, width; 1 mm - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Conotelus mexicanus Sharp

2 9) Body compact to moderately elongate, never exactly as above; elytra variable, complete ( Figs. 10–12 ), or exposing only a small portion of pygidium at suture

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( Fig. 13 ), or shortened and exposing up to three terga ( Figs. 3–9 ); scutellum obtuse, triangular, or broadly rounded, never sub-truncate apically - - - - - - - - - - - - - - - 3

3) Paratergites 5 and 6 broad, flattened, and strongly sinuate along medial margin ( Figs. 3, 4 ); labrum usually truncate or broadly rounded, sometimes with concave emargination which is interrupted by a small medial tooth with short setae inside ( Fig. 14 ); metasternal axillary space absent; elytra shortened, usually with three abdominal terga exposed ( Figs. 3, 4 ); usually with visible sternite 2 shortest, 3 and 4 sub-equal and each twice as long as 2, at least laterally ( Fig. 15 ), rarely with abdomen shortened exposing only 1 or 2 abdominal segments beyond elytra and visible sternites 2-4 subequal - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - endemic Hawaiian species

3 9) Paratergites less broad, sloping laterally, and never strongly sinuate along medial margin ( Figs. 6, 7 ); labrum bilobed, usually with a distinct medial notch or incision with sides convex ( Figs. 16, 17 ), if emargination shallow and concave then lobes with fringe of long setae ( Fig. 18 ), never with medial tooth; axillary space usually present ( Figs. 19–22 ), sometimes small ( Fig. 23 ); elytra shortened or complete; visible abdominal sternites various, 2 and 3 short ( Fig. 23 ) or 2-4 subequal ( Fig. 21 ) - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 4

4) Pronotal lateral margins not explanate, disc convex ( Fig. 24 ) - - - - - - - - - - - - - - - - - - - - - - - 5

4 9) Pronotal lateral margins always explanate near posterior angle, or for entire length, disc depressed - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 13

5) Pronotum widest at base, anterior margin slightly more than half as wide, sides evenly curved; prosternum with long golden setae medially; metathoracic coxal lines forming faint raise triangles on visible abdominal sternite 1 ( Fig. 25 ); eyes conspicuously setous; unicolorous, black to dark brown; dorsal surface clothed with conspicuous golden setae; pygidium with deep pits along basal margin; ( Fig. 5 ); length 3.4–3.6 mm - - - - - - - - - - - - - - - - - - - - Aethina (Idaethina) concolor (Macleay)

5 9) Pronotum quadrate to slightly narrowed anteriorly, sides sub-parallel; prosternum at most with sparse setae; visible abdominal sternite 1 without raised triangles; eyes without conspicuously setae; unicolorous or with markings, dark brown to light brown; dorsal surface with setae various; pygidium without basal pits; ( Figs. 6–8 ) - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 6

6) Visible abdominal sternites 2-5 subequal; mesosternum behind prosternal process flat, with large coalescing punctures; three abdominal terga typically exposed; dorsum light brown to black with reddish tinge, a small pale area on humeri; shining; sparsely covered with inconspicuous dark and light setae; length 3.0–4.5 mm - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Carpophilus (Urophorus) humeralis (F.)

6 9) Visible abdominal sternites 2 and 3 very short, 4 as long as 2 and 3 combined ( Fig. 23 ); mesosternal disc unmodified or with median impunctate carina; two abdominal terga exposed; surface, color, and pubescence variable - - - - - - - - - - - - - - - - 7

7) Mesosternum behind prosternal process strongly punctate, divided longitudinally by a narrow median longitudinal impunctate carina ( Fig. 23 ); elytra with light maculae on humeri and basally extending anteriorly and laterally, dark areas laterad of scutellum convergent posteriorly along midline; ( Fig. 6 ); length 2.0–4.0 mm - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Carpophilus hemipterus (L.)

7 9) Mesosternum behind prosternum not divided longitudinally by a narrow median impunctate carina; coloration and pubescence variable - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 8

8) Antennomere 8 half or more as wide as 9 ( Fig. 26 ); pronotal disc with sparse and definite punctures, separated by 1–2 diameters; lateral area of prosternum ( Fig. 28 ) weakly punctate and granular; dorsum red-brown with dull yellow elytra, elytra often with pronotal color on lateral margins, apex, and bordering scutellum (similar to C. dimidiatus , Fig. 8 ); length 1.8–2.9 mm - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Carpophilus ( dimidiatus ) freemani Dobson

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176 8 9) Antennomere 8 less than half as wide as 9 ( Fig. 27 ); punctation and color variable - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 9

9) Axillary space large, margin directed posteriorly, meeting metepisternal suture beyond middle ( Fig. 19 ); pronotum with faint microsculpture, surface shining; pronotum and elytra unicolorous, head slightly darker, light brown to dark reddish brown; clothed with inconspicuous light setae; length 2.7–3.5 mm - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Carpophilus marginellus Motschulsky

9 9) Axillary space smaller, posterior margin following mesocoxal cavity for some distance anteriad ( Fig. 20 ), meeting metepisternal suture at or before middle; pronotal disc with granular microsculpture; seta color variable - - - - - - - - - - - - - - - - - - - 10

10) Pronotum with sparse punctures on disc anterio-medial of center, separated by 1– 3 diameters; dorsum light to dark brown, elytra with variable light areas, in heavily sclerotized individuals extended anteriad along suture prior to apex, past scutellum, and laterally along base to humeral angle; length 2.5–3.4 mm - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Carpophilus ( dimidiatus ) maculatus Murray

10 9) Pronotum with punctures more dense on disc anterio-medial of center, most separated by no more than 1 diameter; color variable 11

- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -

11) Anterio-medial area of prosternum ( Fig. 28 ) with punctures obsolete; in heavily sclerotized individuals each elytron with central dark macula surrounded by red to yellow area, a dark band extended from the humeral angle along the lateral margin and around the posterior margin, sometimes pattern indistinct, predominantly yellow with light brown maculae; ( Fig. 7 ); length 2.9–3.5 mm - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Carpophilus oculatus Murray

11 9) Anterio-medial area of prosternum ( Fig. 28 ) with punctures deeply impressed; color variable 12

- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -

12) Lateral area of prosternum ( Fig. 28 ) with punctures deeply impressed, interspaces shining; male anal sclerite and female pygidium evenly rounded; antennomere 3 longer than 2 ( Fig. 27 ); head and pronotum evenly reddish brown, infrequently lighter at center of pronotal disc, elytra with background lighter, dark areas often present adjacent to scutellum, laterally, apically, and rarely at humeri forming a vague light V-shaped transverse band, abdomen dark brown; covered dorsally with conspicuous pale setae; ( Fig. 8 ); length 2.2–3.2 mm - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Carpophilus ( dimidiatus ) dimidiatus (F.)

12 9) Lateral area of prosternum ( Fig. 28 ) with punctures obsolete, irregular, interspaces dull with granular microsculpture; male anal sclerite and female pygidium acuminate; antennomere 2 equal to or longer than 3; head and pronotum light reddish-brown, often darker at center of pronotal disc, (elytra similar to C. dimidiatus , Fig. 8 ), abdomen dark brown; elytra with light and dark setae; length 2.0–3.4 mm - - - - - - - - - - - - - - - - - - - - - - - - - - Carpophilus ( dimidiatus ) mutilatus (Erichson)

13) Pygidium usually exposed (may be deflexed and not visible in dorsal view) ( Fig. 9 ); body small;2.0 mm; scutellum acutely triangular ( Figs. 9 , 29 ); elytral punctation not organized in distinct striae; color predominantly yellow, may have some darker maculae on elytra and on anterior and posterior margins of pronotum ( Fig. 9 ) - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 14

13 9) Pygidium concealed or at most partially exposed between elytral apices ( Fig. 13 ); body larger.2.5 mm; scutellum obtuse ( Figs. 10–13 ); elytral punctation and setation usually arranged in distinct striae; color dark to light brown with light brown to yellow markings - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 16

14) Each elytron longest near suture ( Fig. 29 ); pronotum with punctation deep, interspaces shining with microsculpture consisting of very faint discontinuous lines; uniformly brownish-yellow; length 2.0–2.4 mm - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Epuraea (Haptoncus) munda (Sharp)

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14 9) Each elytron longest medially ( Fig. 9 ); pronotal punctation very shallow, less shining with microsculpture between punctures consisting of fine parallel lines or fine granular mesh; color variable - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 15

15) Pronotum slightly explanate at hind angles, microsculpture consisting of fine parallel lines, disc convex; dorsum unicolorous, brownish-yellow; length 2.0–2.4 mm - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Epuraea (Haptoncus) luteola Erichson

15 9) Pronotum distinctly explanate at hind angle, microsculpture fine granular mesh, disc less convex; pronotum with background color yellow with dark marks on anterior and posterior margins, each elytron with dark mark at base, apex, and a transverse mark at middle, infrequently marks lighter or absent in teneral individuals; ( Fig. 9 ); length 2.0–2.4 mm - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Epuraea (Haptoncus) ocularis Fairmaire

16) Elytral punctation and setation not forming distinct striae; labrum with shallow, concave emargination and long setal comb ( Fig. 18 ); pronotum dark brown on disc, margin lighter and explanate, elytra variegated, color from dark to light brown; ( Fig. 10 ); length 2.8–3.4 mm - - - - - - - - - - - - - - - - - - - - - - - - - Omosita discoidea (F.)

16 9) Elytral punctation and setation forming distinct striae; labrum distinctly notched medially ( Fig. 16 ); color variable - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 17

17) Axillary space with margin meeting metepisternal suture near or before middle ( Fig. 22 ); length 2.0–3.1 mm - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 18

17 9) Axillary space with margin paralleling metepisternal suture to metacoxae, forming narrow raised area ( Fig. 21 ); length.4.5 mm - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 19

18) Pronotum somewhat narrowed posteriorly, strongly narrowed anteriorly, and anterior margin deeply emarginate; lateral elytral margins widely explanate in dorsal view; males with both mesothoracic tibia strongly sinuate and metathoracic tibia enlarged in apical 2/3; ( Fig. 11 ); length 2.5–3.1 mm - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Stelidota geminata (Say)

18 9) Pronotum not narrowed posteriorly, sides parallel in basal 1/2, less narrowed anteriorly, and anterior margin shallowly emarginate; elytral lateral margins narrow in dorsal view; males with only metathoracic tibia distinctly enlarged in apical 1/2; ( Fig. 12 ); length 2.0–2.5 mm - - - - - - - - - - - Stelidota chontalensis Sharp

19) Metathoracic coxal lines on visible abdominal sternite 1 obliquely divergent, nearly reaching posterior margin of sternite ( Fig. 21 ); elytra distinctly costate basally; elytral coloration distinctive; ( Fig. 13 ); length 4.8–5.5 mm - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Phenolia (Aethinodes) attenuata (Reitter)

19 9) Metathoracic coxal lines on visible abdominal sternite 1 closely following metathoracic coxal cavity; elytra vaguely costate basally; elytral coloration distinctive; ( Fig. 30 ); length 5.6–8.0 mm - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Phenolia (Lasiodites) limbata tibialis (Boheman)

Notes on Adventive Nitidulidae

Subfamily Cybocephalinae

Cybocephalus nipponicus Endrödy-Younga, 1971: 244 .

Distribution. Oahu, Maui, Hawaii.

Cybocephalus spp. are predatory on scale insects and whiteflies and have been used in biological control, though no record exists of direct introduction into Hawaii for this purpose. Cybocephalus nipponicus was first reported from Maui and Hawaii from specimens collected in 1989 (Beardsley and Tsuda 1992). The original identification was based on external characters. Dissection of male genitalia and comparison with illustrations in the original description (Endrödy-Younga 1971) confirms the identification. The species description was based on specimens from Japan,

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178 China, India, Ceylon, Singapore, Palau Islands, and Mariana Islands (Endrödy-Younga 1971).

Subfamily Cillaeinae

Cillaeinae includes one adventive and all 134 described endemics.

Conotelus mexicanus Murray, 1864: 337 .

Distribution. Kauai, Oahu, Maui, Hawaii.

Species of Conotelus inhabit fresh flowers as adults, and the larva develop in fallen corollas. Conotelus mexicanus was first reported as an unidentified nitidulid by Chilson and Ford in March 1954, and again in April of the same year ( Maehler 1955) as occurring in Capparis and cotton flowers. Soon afterward it was collected again from morning glory flowers and identified ( Bianchi 1955). A more complete survey of its host plants, distribution, and abundance was soon reported ( Nishida 1957). It has also been found in Sida fallax Walpers flowers ( Bianchi 1961) and is common in lily flowers in Honolulu (Ewing, pers. obs.). Native from Southwestern U.S. through Mexico and Central America as far south as Panama ( Parsons 1943).

Subfamily Carpophilinae

Initially it was believed that Carpophilus species caused direct damage to healthy pineapple fruit in Hawaii ( Illingworth 1929). It was subsequently shown that they are more properly defined a nuisance pest, attacking previously damaged and overripe fruit, and field refuse ( Schmidt 1935). Large populations of these species can develop quickly around fields on discarded plants after harvest, and around canneries when proper sanitary practices are not followed ( Schmidt 1935). All Carpophilus present in Hawaii are globally distributed with the exception of C. oculatus , which is likely limited to the South Pacific and Southeast Asia.

Carpophilus (Urophorus) humeralis (F.)

Nitidula humeralis F., 1798: 74.

Common name. Yellow shouldered souring beetle, pineapple souring beetle.

Distribution. Niihau, Kauai, Oahu, Molokai, Lanai, Maui, Hawaii.

Carpophilus humeralis was first reported in Hawaii in 1908 ( Sharp and Scott 1908). Only 13 specimens were collected and it was considered to be the rarest of the Carpophilus present in Hawaii at the time. Three other species, C. hemipterus , C. dimidiatus , and C. maculatus had been reported earlier ( Sharp 1878). This species quickly became one of the most abundant in the islands and has proven to be a major nuisance around pineapple fields and canneries ( Illingworth 1929; Schmidt 1935; Hinton 1945). It replaces C. hemipterus as the most abundant nitidulid in pineapple fields as the refuse proceeds into a more advanced state of decay ( Schmidt 1935). It is the most common nitidulid in sugarcane fields where it feeds on wounds created by other insects, though it is not considered a pest of cane ( Williams 1931). It can be found in decaying flowers, fruits, and other organic material from sea level to high elevation montane forests. It is also known to attack a wide variety of fruits and grains worldwide ( Hinton 1945).

Carpophilus hemipterus (L.)

Dermestes hemipterus L., 1758: 358.

Common name. Dried fruit beetle.

Distribution. Kauai, Oahu, Molokai, Lanai, Maui.

Carpophilus hemipterus was rare in the 1870’s when first collected ( Sharp 1878), but is now a common nitidulid on the islands. Along with Carpophilus humeralis , it is the

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most abundant nitidulid around Hawaii’s pineapple fields. Carpophilus hemipterus tends to be most abundant during the early stages of decomposition, and is subsequently replaced by C. humeralis ( Schmidt 1935) . It can be found in decaying flowers, fruits, and other organic material from sea level to high elevation montane forests.

Carpophilus ( dimidiatus ) dimidiatus (F.)

Nitidula dimidiatus F., 1792: 261.

Common name. Corn sap beetle.

Distribution. Midway, Laysan, Necker, Nihoa, Niihau, Kauai, Oahu, Molokai, Maui, Hawaii.

Carpophilus dimidiatus along with the closely related C. freemani , C. fumatus , C. maculatus , C. mutilatus , and C. pilosellus form the dimidiatus group ( Connell 1977). These species share habits and some of the species have been combined historically. Carpophilus dimidiatus , C. freemani , C. maculatus , and C. mutilatus are all present in the Hawaiian islands. Carpophilus dimidiatus was first collected in Hawaii by the Rev. Thomas Blackburn ( Sharp 1878). Blackburn reported that it was ‘‘almost the commonest Hawaiian beetle.’’ This species attacks many types of stored foods, especially grains and fruits (Blackburn and Sharp 1885), including bananas in Hawaii ( Swezey 1939). It can be found in decaying flowers, fruits, and other organic material from sea level to high elevation montane forests. A member of the dimidiatus group, see Hinton (1945) for an exhaustive list of habits.

Carpophilus ( dimidiatus ) freemani Dobson, 1956: 41–42 .

Distribution. Oahu.

Carpophilus freemani is not commonly collected in Hawaii. The first reported specimen was collected on Oahu in 1949 ( Ford 1961). An additional specimen identified during the course of this work was collected in 1922 in Honolulu. A member of the dimidiatus group, see Hinton (1945) for an exhaustive list of habits under C. dimidiatus .

Carpophilus ( dimidiatus ) maculatus Murray, 1864: 372 .

Distribution. Kauai, Oahu, Molokai, Lanai, Maui, Hawaii.

Carpophilus maculatus was the first species of Nitidulidae described from Hawaii. It was described from specimens collected on Oahu ( Murray 1864) and is presently recorded from all of the high islands. This species has been reported from macadamia nuts ( Browne 1939), and is a nuisance pest around pineapple fields ( Illingworth 1929). A member of the dimidiatus group, see Hinton (1945) for an exhaustive list of habits.

Carpophilus ( dimidiatus ) mutilatus Erichson, 1843:258 . (redescribed) Dobson, 1954: 397.

Distribution. Kauai, Oahu, Molokai.

Carpophilus mutilatus was first recognized as being established in Hawaii from material collected in 1952 ( Ford 1961). It is relatively common in decaying fruits and grains. A member of the dimidiatus group, see Hinton (1945) for an exhaustive list of habits under C. dimidiatus .

Carpophilus marginellus Motschulsky, 1858: 262 .

Distribution. Oahu, Hawaii.

Carpophilus marginellus was first collected in 1949 on Oahu though it was not immediately identified due to difficulties in separating Carpophilus species ( Ford 1959) . Carpophilus marginellus is a known pest of whole and milled grains ( Hinton 1945).

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180 Carpophilus oculatus Murray, 1864: 374 .

Distribution. Oahu.

Carpophilus oculatus was intercepted at Honolulu airport, from a departing passenger, inside seeds collected on Oahu on 2 April 1964 ( Shiroma 1966). Carpophilus oculatus is apparently rare in Hawaii, though it will attack pineapples and other fermenting fruits. Described from Bora Bora in the Society Islands ( Murray 1864), it has been reported in the Mariana, Marshall, and Caroline Island groups of Micronesia ( Gillogly 1962), and is likely more widely distributed around the South Pacific and Southeast Asia.

Subfamily Epuraeinae

Epuraea (Haptoncus) luteola Erichson, 1843: 272 .

Distribution. Oahu.

Epuraea luteola was first collected in October and November of 1949 ( Ford 1960). It is often found with specimens of E. munda and E. ocularis in lowlands as well as in native forests, and can often be collected from the decaying fruit of Freycinetia arborea Gaudichaud-Beaupre , especially in disturbed forests. It is a generalist pest of fruit ( Hinton 1945). Distribution tropicopolitan ( Parsons 1943).

Epuraea (Haptoncus) munda (Sharp)

Haptoncus mundus Sharp, 1878: 139 .

Distribution. Kauai, Oahu, Lanai, Hawaii.

Epuraea munda has only been applied to specimens collected in Hawaii, though Sharp considered it to be introduced ( Sharp and Scott 1908). Sharp (1878) expressed the opinion that it was closely related to E. testacea from New Guinea, and separated it on the basis of the more explanate pronotal margins. When a thorough revision of this genus is completed it may prove to be a synonym, though it may represent an endemic species resulting from a relatively recent colonization event. Found in the same situations as the other Epuraea species.

Epuraea (Haptoncus) ocularis Fairmaire, 1849: 363 .

Distribution. Kauai, Oahu, Lanai, Hawaii.

Epuraea ocularis was first reported by Sharp (1878) from specimens collected by Rev. Blackburn. Found in the same situations as the other Epuraea species. Distributed throughout the Orient, Southeast Asia, China, and East Africa (see Grouvelle 1913; Kirejtshuk 1998).

Subfamily Nitidulinae

Aethina (Idaethina) concolor (Macleay)

Nitidula concolor Macleay, 1873: 162 .

Common name. Hibiscus flower beetle.

Distribution. Kauai, Oahu, Molokai, Lanai, Maui, Hawaii.

Aethina (Idaethina) concolor was first reported as Macroura sp. from specimens collected at Kahuku in November 1983 (Beardsley and Samuelson 1992). This species is native to Australia where it is most commonly found in flowers of hibiscus. A tubular flower specialist, in Hawaii it has been reported from flowers of Hibiscus, morning glory, and various Cucurbitaceae . Distributed throughout the South Pacific ( Kirejtshuk and Lawrence 1999).

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Omosita discoidea (F.)

Nitidula discoidea F., 1775: 78.

Distribution. Maui.

Omosita discoidea was first collected on Haleakala ( Timberlake 1920) and identified 15 years later ( Swezey 1934). It apparently has not spread to the other islands. Unlike most of the pest species, it does not attack fruits and grains, but rather attacks dried animal tissue and other dry decaying material. Native to Europe it has a holarctic distribution, being introduced into North America ( Audisio 1993).

Stelidota geminata (Say)

Nitidula geminata Say, 1825: 181 .

Distribution. Kauai, Oahu, Molokai, Lanai, Maui, Hawaii.

Stelidota geminata was first collected on Oahu and Hawaii islands in 1997 and reported in 2004 (Ewing and Cline 2004), it has spread quickly and is present on all of the main islands. Most commonly found in decaying fruit at low elevations and in leaf litter and decaying wood in higher elevation native forests. Distributed worldwide ( Audisio 1993).

Stelidota chontalensis Sharp, 1889: 314 .

Distribution. Oahu.

Originally reported as Stelidota sp. from specimens collected on dried guava leaves at the University of Hawaii Experimental Farm at Waimanalo and in Kapiolani Park, Kaimuki, all in 1992 ( Beardsley et al. 1995). Native to Central America ( Sharp 1889).

Phenolia (Aethinodes) attenuata (Reitter)

Lasiodactylus attenuatus Reitter, 1878 (1879) : 169.

Distribution. Kauai, Molokai, Oahu.

Phenolia (Aethinodes) attenuata was first collected on Oahu in 1998 and reported in 2004 (Ewing and Cline 2004). Common at low elevations in decaying citrus and other rotting fruits. Distributed from Southeast Asia through the Malay Archipelago and at least as far east as New Guinea ( Grouvelle 1913).

Phenolia (Lasiodites) limbata tibialis (Boheman)

Soronia tibialis Boheman, 1851: 568 .

Distribution. Oahu, Molokai, Maui, Hawaii.

Phenolia (Lasiodites) limbata tibialis is the largest adventive Nitidulidae in Hawaii. First collected on Hawaii Island in 1996 and reported as Lasiodactylus sp. probably tibialis (Kumashiro and Heu 1997) , and Oahu and Maui records reported in 2004 (Ewing and Cline 2004). Often collected in the same situation with P. attenuata . Distributed throughout Africa, Asia, the Orient, and South Pacific (Kirejtshuk and Kvamme 2002).

Figures

Figs. 1–13. 1) Cybocephalus nipponicus; 2) Conotelus mexicanus; 3) Cillaeopeplus

Figs. 14–23. 14) Apetasimus pleomelarrosus, labrum; 15) Prosopeus floricola; 16) Carpo-

Figs. 24–30. 24) Aethina (Idaethina) concolor, anterior view; 25) A. (I.) concolor, ventral