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Home , Azya orbigera, Chilocorinae, Clitostethus, Coccidulinae, Coelophora, Coelophora inaequalis

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KEY TO THE COCCINELIDS (COLEOPTERA: ) ESTABLISHED IN HAWAII

JOHN R. LEEPER PO Box 13086 Las Cruces, NM USA, 88013 [email protected]

Abstract. Leeper (2015) identified 167 coccinellid as having been introduced to the Hawaiian Islands. Forty-six (46) species are considered established for which this paper provides a key.

INTRODUCTION Leeper (1976) wrote the only previous key to the Coccinellidae established on the seven habited Hawaiian Islands (Niihau, Kauai, Oahu, Maui, Molokai, Lanai, and Hawaii). This paper revises and updates Leeper’s key and is based on a survey of unpublished Hawaii State Department of Agriculture (DoA) records, material published in Hawaii and specimens and notes located at the Hawaii State Department of Agriculture, the Bernice P. Bishop Museum (BM), the University of Hawaii, Manoa Museum (UH) and Leeper (2015).

Leeper (1976) identified 38 species, two with varieties included ( inaequalis and var. novemmaculata; and Olla abdominalis and Olla abdominalis var. plagiata). Both variants are not recognized and removed from the present key. Three other species ( pumilio, Diomus near pumilio and Diomus sp.) were removed due to the lack of documentation of establishment. The names of nineteen species remain valid while the names of sixteen species required changes. Eleven species were added to the list of established species. Documentation for the above changes can be found in Leeper (2015).

METHODS and Materials The phylogeny and classification in this paper is based on the preliminary classification of Coccinellidae proposed by Kovar (1996) and further defined by Nedved and Kovar (2012). Their classification system recognizes nine subfamilies, five (, , , , Coccidulinae and Ortaliinae) of which are represented in Hawaii.

Observations of specimens in the UH and DoA collections were supplemented by insect descriptions from within the following references in constructing the keys to subfamilies, tribes, genera and species in this paper: Booth and Pope (1986); Booth, Cross, Fowler and Shaw (1995); Chapin, E. A. (1926, 1965); Chapin, J. B. (1974), Escalona and Slipinski (2010); Gordon (1970, 1972, 1976, 1977, 1978, 1982, 1985, 1997, 1999, 2000); Gordon and Gonzalez (2002); Kamiya, 1961; Leeper (1976); Pang and Gordon (1986); Pang and Slipinski (2009, 2010); Peck (2006); Pope and Lawrence (1990); Raimundo and van Harten (2002); Sasaji (1968, 1971); Slipinski (2004, 2007); Slipinski and Burckhardt (2006); Slipinski and Giorgi (2006); Vandenberg (2002).

Colors used in describing the species within this paper relied heavily upon what was observed on specimens at the UH and DoA collections. Many of the specimens were old and colors may have faded or become muted. Additionally, a few specimens within the collections appear to have been killed soon after adult emergence and had not taken on their full adult coloring.

The overall key is broken down into a series of sub-keys to subfamilies, tribes and then the species within each tribe. Gross morphological characteristics and color patterns were used in writing the keys. Examination of genitalia requires the destruction of specimens and was not possible for writing the keys. However, notations were made when authorities relied upon genitalia to differentiate between species and their papers may be relied upon to more definitively identify species.

The final step in the key for identifying a species also includes, in parenthesis, the islands on which the species has been recorded to be established on. Abbreviations for the islands are as follows: Niihau: Ni; Kauai: Ka; Oahu: Oa; Maui: Ma; Molokai: Mo; Lanai: La; and Hawaii: Ha. Lack of an island reference for particular species should not preclude it from being on an island as new island establishments through interisland commerce or natural distribution may occur.

A. KEY TO THE SUBFAMILIES OF COCCINELLIDAE ESTABLISHED IN HAWAII 1a. Clypeus expanded laterally, shelf-like, under eyes; distinctly ventral insertion of antennae; tarsus trimerous or cryptotetramerous ...... Chilocorinae Sasaji, 1968: 20 (B.) 1b. Clypeus not expanded laterally, or if so, just slightly and not shelf-like ...... 2 2a (1b). Pronotum and elytra glabrous; mandible bifid at apex, basal tooth present; maxillary palpus securiform ….Coccinellinae Ganglebauer, 1899: 954, 986 (C.) 2b (1b). Pronotum pubescent, elytra pubescent or glabrous ...... 3 3a (2b). Body small to medium sized, hemispherical to elliptical; maxillary palpus thin, apical segment elongate and conical, with spindle shaped club; antennae inserted dorsally ……..Sticholotidinae Gordon, 1977: 186 (emendation) (D.) 3b (2b). Maxillary palpus broad, apical segment angled, not conical apically...... 4 4a (3b). Body minute to small size; antenna short, two thirds or less of head width, apical segment reduced with flagellomeres; eyes large; apical segment of maxillary palpus weakly securiform to parallel- sided ….. Scymninae Della Beffa, 1912: 168 (E.) 4b (3b). Antenna long, more than two thirds of head width ……….5 5a (4b). Body shape oblong oval, weakly to moderately convex, pubescent; maxillary palpus securiform; antenna long, 8-11 segmented; eyes with coarse facets; pronotum red, elytra uniformly black; length of pubescence medium with long, erected setae ………Coccidulinae Sasaji, 1968: 22 (F.) 5b (4b). Body oval to rounded, pubescence fine and short; eyes large with small to minute facets; mandible bifid at apex, basal tooth present; apical segment of maxillary palpus short and securiform; antennae inserted between eyes, not dorsally or ventrally ….Ortaliinae Kovar, 1996: 26 (G.)

B. KEY TO THE TRIBES IN THE SUBFAMILY CHILOCORINAE ESTABLISHED IN HAWAII 1a. Glabrous (round helmet shaped), shiny, lacking pubescence, males with six visible abdominal sternites, females in some species with only five visible abdominal sternites ……………….. Chilocorini Costa, 1849: 9 (B1.) 1b. Elliptical in shape, one-third longer than wide, pubescent, with five visible sternites in both males and females ...Telsimiini Casey, 1899: 16 (B2.)

B1. KEY TO THE SPECIES IN THE TRIBE CHILOCORINI ESTABLISHED IN HAWAII 1a. Tibial spurs present on legs II and III, metacoxal arcs as in or Pullus, antenna 10, 9 or 8- segmented…………………… 2 1b. Tibial spurs absent on all legs, metacoxal arcs as in or Diomus, antenna 9, 8 or 7- segmented…………………..3 2a (1a). Elytra black, each with two yellow-gold longitudinal stripes, one stripe along outer margin, stripes meeting at apex, elytral margin yellow-gold and slightly reflexed ; head and pronotum - yellow; tarsal claw simple, sometimes slightly thickened at base, antenna 8-segmented; body oval, convex; length 2.5-3.5 mm, width 1.5-2.5 mm; tibia 2 and 3 with apical spurs; tarsal claw simple; post coxal line of 1st abdominal ventrite complete, recurved; antenna 8-segmented; apical antennomere very small and embedded in penultimate one (Ni, Oa) ……. suturalis (Fabricius, 1798: 78) 2b (1a). Body dark metallic blue with prothoracic hypomera orange-brown; elytral margin slightly reflex; venter and legs orange-brown, tarsal claw with tooth in basal half, which is sometimes quadrate, sometimes acute, often very small, antenna 10-segmented, size up to 6 mm, elytra blue; body nearly hemispherical; length 4.9-5.2 mm, width 4.5-4.75 mm; all tibia without spurs; tarsal claw with tooth at base; post coxal line of 1st abdominal ventrite incomplete; antenna 10-segmented; apical antennomere not embedded in penultimate one (Ni, Ka, Oa, Ma, Mo, La, Hi) …… coeruleus Mulsant, 1850: 472 3a (1b). Antenna 7-segmented; body hemispherical; length 3.0-4.25 mm, width 2.85-3.9 mm; upper surface glabrous; elytra stronglt reflexed; postcoxal line of 1st abdominal ventrite parallel and close to posterior margin; body metallic iridescent blue-green to purple; males with anterior margin of pronotum yellow-orange, female with entire pronotum same metallic color as the rest of the body (Ka, Oa, Ma, Mo, La, Hi) …… chalybeus (Boisduval, 1835: 595) 3b (1b). Antenna 8-segmented ………4 4a (3b). Body nearly hemispherical; length 4.5-5.1 mm, width 4.1-4.7 mm; dorsum strongly convex, orange-brown (elytra may at times be yellow-brown to gray) with elytral margin strongly edge strongly reflex and black; venter uniformly orange-yellow (Oa, Ma, Mo, Hi) …. circumdatus (Gyllenhal in Schonherr, 1808: 152) 4b (3b). Body nearly hemispherical; length 3.2-4.0 mm, width 2.9-3.9 mm; dorsum strongly convex, black with prothoracic hypomera orange-brown to brown; elytral margin reflex and same color as rest of elytra; venter and legs orange-yellow to yellow-brown; head orange-yellow (Oa, Ma) ……. Chilocorus nigrita (Fabricius, 1798: 79)

B2. KEY TO THE SPECIES IN THE TRIBE TELSIMIINI ESTABLISHED IN HAWAII 1a. Body oval; length 1.85 mm, width 1.5 mm; posterior margin of fifth abdominal sternite of male feebly bisinuate; metasternum with short, median, finely impressed line moving from posterior margin forward; dorsal pubescence gray to white, very fine and suberect; epistomal hairs with yellow tinge; width between eyes about half the entire head width; anterior margin of clypeus narrowly yellow- brown; punctures on thorax and abdomen more coarse than on dorsum (Oa) …….Telsimia nigra (Weise, 1879: 149) 1b. Body oval; length 1.6 mm, width 1.2 mm; posterior margin of fifth abdominal sternite of male not bisinuate; metasternum without impressed line; dorsal pubescence grey, fine and suberect; venter dark brown; antenna, mouth parts and legs brownish-red; head with short gray to golden hairs; width between eyes less than half the entire head width; anterior margin of clypeus black (Oa, Mo, La, Hi) …….Telsimia nitida Chapin, 1926: 131

C. KEY TO THE TRIBES IN THE SUBFAMILY COCCINELLINAE ESTABLISHED IN HAWAII 1a. Body length 3.0 mm or greater; gena extending beneath eye; eye usually finely faceted; mandible 2 teeth at apex; clypeus narrower than frons, apex usually emarginate with anterolateral angle; anterior border of pronotum deeply excavate around head …….Coccinellini Weise, 1885: 7 (C1.) 1b. Body length 3.0 mm or less; gena not extending beneath eye; eye coarsely faceted; mandible with 5 teeth; clypeus wider than frons, apex without produced anterolateral angle; anterior border of pronotum feebly emarginate, partially concealing head ……..Psylloborini Casey, 1899: 73, 100 (C2.)

C1. KEY TO THE SPECIES IN THE TRIBE COCCINELLINI ESTABLISHED IN HAWAII 1a. Tarsal claws cleft near apical 1/3; form slender; legs distinctly visible beyond body in dorsal view ...... 2 1b. Tarsal claws with subquadrate basal tooth; or if tooth median, then form rounded; legs barely visible beyond body in dorsal view ...... 3 2a (1a). Length 4.2-7.3 mm, width 2.5-4.9 mm; pronotum black with two yellow commas, corners and anterior margin yellow; face about half black, half yellow; elytra orange, elytral markings extremely variable, most commonly with twelve spots arranged in a 2-4-2-2-2 pattern sometimes appearing as a 2- 4-2-2 pattern (Ni, Ka, Oa, Ma, Mo, La, Hi) …….. Hippodamia convergens Guerin, 1842: 321 2b (1a). Length 4.0-7.0 mm, width 2.8-5.0 mm; pronotum black, commonly with two yellow dots, at least front corners yellow, anterior margin not yellow; face about two thirds black, one third yellow; elytra orange, elytral markings extremely variable, most commonly lacking spots (Oa, Ma, Hi) …………………… Hippodamia quinquesignata ambigua LeConte, 1852: 131

NOTE: Gordon (1985: 727) states the two above species “are very difficult to distinguish without referring to male genitalia.”

3a (1b). Body rounded, convex; orange to yellow-brown; elytral suture not black; elytral margin reflexed; eighteen black spots on elytra arranged in a 5-6-5-2 pattern (Ma, Hi) ………… (Boisduval, 1835: 604) 3b (1b). Elytral spots not as statement ...... 4 4a (3b). Elytron yellow to orange-brown with black sutural margin and with black spots that may be reduced in size and somewhat indistinct or may be large and coalesce ...... 5 4b (3b). Elytron with color pattern not resembling that of Coelophora ...... 6 5a (4a).Pronotum yellow to orange-brown, two black spots reduced to short lines at front of elytron and a small spot in the middle third of the middle elytron; elytra reflexed area black; scutellum yellow with black edges (Ka, Oa, Ma, Hi)……….. Coelophora atrolineata Fairmaire, 1881: 491 UH 5a (4a). Pronotum black with orange-brown to yellow-brown markings across anterior margin; elytra orange-brown to yellow-brown; elytral suture black; elytra reflexed with nine irregular spots which may coalesce with each other or with elytral margins; scutellum black (Ni, Ka, Oa, Ma, Mo, La, Hi) ……………Coelophora inaequalis (Fabricius, 1775: 80) 6a (4b). Pronotum black with a large, subtrapezoidal or triangular white spot on each anterolateral angle (apical margin of pronotum sometimes narrowly pale); epipleuron nearly horizontal, not descending externally ...... 7 6b (4b). Pronotum not as above; epipleuron horizontal or descending externally ...... 8 7a (6a). Pronotum orange-brown to yellow-brown without markings; elytra orange-brown to yellow- brown with ten black spots encircled by lighter orange-brown to yellow-brown rings, spots arranged in a 4-4-2 pattern (Ka, Oa, Ma, Mo, Hi) ………Bothrocalvia pupillata (Swartz, in Schonherr, 1808: 184) 7b (6a). Body oval, widest at center; pronotum black with yellow hypomera; head black with 2 well separated pale spots; elytra orange-brown to yellow-brown with seven black spots in a 1-2-2-2 pattern, sometimes appearing as a 1-4-2 pattern, spot one located on elytral suture behind two yellow areas (Ka, Oa, Ma, Mo, La, Hi) ….Coccinella septempunctata Linnaeus, 1758: 365 8a (6b). Postcoxal line on first abdominal sternum with oblique dividing line; epipleuron strongly, abruptly descending externally; punctures on elytron fine, nearly invisible; coloring highly variable (gray to tan); pronotum with seven black spots; elytra with sixteen black spots in an 8-6-2 pattern; spots may merge. Another more distinctly different color from exists which is mostly black with yellow markings across margin of pronotum and two yellow to orange elytral spots (Ka, Oa, Ma, Mo, La, Hi) ...... Olla v- nigrum (Mulsant, 1866: 64) 8b (6b). Length 3.2-6.5 mm, width 2.9-5.1 mm; pronotum mostly black with lateral pale spot enclosed by black area; elytron amber to orange- red; postcoxal line on first abdominal sternum without oblique dividing line; apex of mesosternum truncate (Oa) ...... sanguinea sanguinea Linnaeus, 1763: 10

C2. KEY TO THE SPECIES IN THE TRIBE PSYLLOBORINI ESTABLISHED IN HAWAII 1. Body more elongate than oval; length 1.75-3.0 mm, width 1.40-2.35 mm; mandible with 5 teeth, 2 apical, 3 internal; body gold color; elytra with four brown spots, middle two on lateral edge of elytra with lighter brown areas coalescing (Oa, Ma) ……..Psyllobora vigintimaculata (Say, 1824: 96)

D. KEY TO THE TRIBES IN THE SUBFAMILY STICHOLOTIDINAE ESTABLISHED IN HAWAII 1a. Antennae 8 or 9 segments; antennal club composed of a single, knife-shaped segment; dorsal surface very sparsely pubescent; 5 abdominal segments with 5th segment longer than segments 2, 3 and 4 combined; prosternum expanded to conceal mouthparts; elytral epipleura with two notches to receive legs at rest; underside of body with depressions to receive the femora; femora very broad and flat ………..Serangiini Blackwelder, 1945: 450 (D1.) 1b. Antennae 10 or 11 segments; antennal club composed of more than 1 segment; 5 abdominal segments with 1st and 5th segments longer than segments 2, 3 combined; elytral epipleura with without notches to receive legs at rest …………. Weise, 1901b: 430 (D2.)

D1. KEY TO THE SPECIES IN THE TRIBE SERANGIINI ESTABLISHED IN HAWAII 1a. Body oval, dark brown to black; length 1.4-1.5 mm, width 1.1-1.18 mm; antennae 9 segmented with anterior segment forming club; head very finely and sparsely punctured, lacking or almost lacking pubescence; 1st and 5th abdominal sternites larger than sternites 2- 4; male legs and head yellow-brown, female darker reddish-yellow; tarsi 3 segmented; elytral epipleura with two notches for reception of legs (Ka, Oa, Ma, Hi) …… catalinae (Horn, 1895: 83) 1b. Body hemispherical, sparsely pubescent, shiny black to dark brown, length 1.25-1.5 mm, width 1.0- 1.25 mm; brown to amber spot on each elytra that may be difficult to see, head in front and femora, antennae, mouth parts, tibiae and tarsi amber; tarsi 4 segmented; dorsum strongly convex; 1st abdominal sternite larger than sternites 2- 5; elytral epipleura extending past first abdominal sternite; elytral epipleura with two notches for reception of legs (Oa, Ma) ……Serangium maculigerum Blackburn, 1892: 73

D2. KEY TO THE SPECIES IN THE TRIBE STICHOLOTIDINI ESTABLISHED IN HAWAII 1a. Body hemispherical, convex; head, pronotum and elytra densely pubescent; antennae with 10 segments, 3-segmented club; abdomen with 5 visible sternites (Oa, Mo) ….. sp.

NOTE: The UH collection contains a single specimen with a note that 6 specimens had been sent to Ken Hagen for identification. There was an accompanying note that R. D. Gordon had identified the specimens as “Pharoscymnus sp. possibly undesc.” This is also corroborated in Beardsley’s (1980: 179) note. The BM collection contains one specimen from Molokai.

1b. Body hemispherical, convex, dark brown to black with four reddish to orange-brown spots on elytra in a 2-2 pattern, first two spots may coalesce at elytral suture; length: 1.6-2.1 mm; width: 1.2-1.8 mm; dorsal surface densely and coarsely punctured; antennae with 11 segments with terminal 3 segments forming a weak club; outer margin of elytra expanded laterally; tarsal claws simple (Oa, Ma, Mo, La, Hi) ……Sticholotis ruficeps Weise, 1902: 511

E. KEY TO THE TRIBES IN THE SUBFAMILY SCYMNINAE ESTABLISHED IN HAWAII 1a. Clypeus not emarginate around the antennal bases; prosternum arcuately produced, partly concealing the mouthparts ………….Stethorini Dobzhansky, 1924: 20 (E1.) 1b. Clypeus emarginate around the antennal bases; prosternum not arcuately produced, mouthparts exposed …….2 2a (1b). Elytra glabrous …………..Hyperaspidini Costa, 1849: 9, 64 (E2.) 2b (1b). Elytra pubescent ………3 3a (2b). Tibial spurs absent; antenna 11 segmented; coxal arch (line) of first abdominal sternite fused to second, not recurved anteriorly ………..Diomini Gordon, 1999: 3 (E3.) 3b (2b). Coxal arch (line) of first abdominal sternite failing to reach basal margin, recurved anteriorly … Costa 1849: 9 (E4.)

E1. KEY TO THE SPECIES IN THE TRIBE STETHORINI ESTABLISHED IN HAWAII 1. Clypeus not emarginate around the antennal bases; prosternum arcuately produced, partly concealing the mouthparts; body oval, dark brown to black, length 1.02-1.5 mm, width 0.78-1.14 mm; dorsum pubescent; distal tip of femora, tibiae and tarsi yellow-brown; mouthparts and antennae yellow- brown; face black or yellow-brown (Ka, Oa, Ma, Hi) …… siphonulus Kapur, 1948: 314

E2. KEY TO THE SPECIES IN THE TRIBE HYPERASPIDINI ESTABLISHED IN HAWAII 1a. Body oval, somewhat convex; length 2.35-2.6 mm, width 1.8-2.0 mm; pronotum entirely black or with yellow lateral margins; elytra with yellow stripe along anterior (outer) margins, not extending to elytral apex (Oa, Ma, Hi) ……… inflexa Casey, 1899: 126 1b. Pronotum and elytra with yellow spots ……2 2a (1b). Body oval, convex; pronotum with large yellow makings on the lateral margins, elytra with four yellow spots in a 2-2 pattern (Oa, Ma, La, Hi) ……..Hyperaspis silvestrii Weise, 1909: 205 2b (1b). Body oval, convex; length 2.3-2.7 mm, width 1.7-2.3 mm; frons, clypeus, labrum, antennae and palpi yellow in both sexes, rest of head colors sexually dimorphic; elytra with ten yellow spots in a 4-4-2 pattern (Ka, Oa, Ma, Mo, La, Hi) ……Hyperaspis pantherina Fursch, 1975: 43

E3. KEY TO THE SPECIES IN THE TRIBE DIOMINI ESTABLISHED IN HAWAII 1a. Length 1.25-1.5 mm, width 0.75-1.0 mm; entire body and legs tan to yellow-brown to light amber; pubescence short, yellowish-white (Ka, Oa, Ma, Mo, La, Hi) …..Diomus debilis (LeConte; 1852: 137) 1b. Length 1.75- 2.5 mm, width 1.25-2.0 mm; predominantly black, with black elytral suture; edges of pronotum yellow-brown to orange, remained black; a yellow-brown to orange band covering each elytra, but not reaching elytral margin; leg one and face black in females; leg one orange-brown, face yellow-brown to orange in males, legs two and three black in both sexes (Ka, Oa, Ma, Mo, La, Hi) ….. (Blackburn, 1889: 196)

E4. KEY TO THE SPECIES IN THE TRIBE SCYMNINI ESTABLISHED IN HAWAII 1a. Head with mouthparts directed postero-ventrad in repose, concealing prosternum; basal antennal segment strongly enlarged ...... 2 1b. Head with mouthparts not concealing prosternum; basal antennal segment not strongly enlarged ...... 3

Note on differentiating Nephaspis bicolor from Nephaspis indus: Gordon (1997: 44) states “Males are difficult to separate, and genitalia usually must be examined except for those males of N. indus that have a brown basomedian spot, which is never present in N. bicolor. Nephaspis bicolor is larger than N. indus, but there is enough overlap in size to make this character unreliable for distinguishing the two species.”

2a. (1a). Elytra black except apical margin yellow; head, pronotum, mouthparts, legs, 2 apical abdominal sternites yellow in male; female similar except head black and pronotum broadly dark; venter coarsely punctured; length 1.4-1.5 mm, width 1.0-1.1 mm (Oa) ………….Nephaspis bicolor Gordon, 1982: 335 2b (1a). Color yellow except elytra brown to dark brown with apex yellow; abdominal sternites 1-4 dark brown to reddish brown; female similar to male in color except clypeus yellowish-red, frons and vertex brown, pronotum dark brown with anterior edge yellow; length 1.3-1.4 mm, width 1.0-1.1 mm (Ka, Oa, Ma, Mo, La, Hi) ……..Nephaspis indus Gordon, 1996: 43 3a (1b). Antenna 9-segmented, basal 2 segments enlarged; length 1.8-2.3 mm, width 1.1-1.5 mm; head, pronotum and legs yellow-brown; eltyra brown with apex yellow-brown; tibiae simple, slender, spurs absent; tarsi three segmented, claw with subquadrate basal tooth (Oa) ...... Sasajiscymnus anomalus (Chapin, 1965: 210) 3b (1b). Antenna 10- or 11 -segmented ...... 4 4a (3b). Prosternum without carinae, or at most with short ridges next to coxal cavities ...... 5 4b (3b). Prosternum with distinct carinae on intercoxal projection, carinae usually reaching anterior margin of prosternum ...7 5a (4a). Body oval, moderately convex, length 1.8 mm, width 1.2 mm; antenna 11-segmented with small symmetrical club; abdominal postcoxal lines curved throughout; elytra mostly dark brown to black, each with a single large yellow-brown to orange-red spot covering about one-third the total area; venter and legs orange-brown (Ka, Oa, Ma, Hi) …….Nephus roepkei (Fluiter, 1938: 49) 5b (4a). Antenna 10-segmented with large asymmetrical club; abdominal postcoxal lines more or less parallel to the hind sternal margin …………6 6a (5b). Body oval; length 1.4-2.1 mm, width 1.0-1.6 mm; elytra mostly dark in color, yellowish spot on apical third (Ni, Ka, Oa, Ma, Mo, La, Hi) ……Scymnobius bilucernarius (Mulsant, 1850: 997) 6b (5b). Body oval; length 1.4-1.8 mm, width 1.0-1.4 mm; male pronotum yellowish-tan, female pronotum brown with pale edges; elytra yellowish-tan with dark J-shaped marking from central elytra margin laterally, dark spot above curve of J (Ka, Oa, Ma, Mo, La, Hi) …..Scymnobius galapagoensis (Waterhouse, 1845: 41) 7a (4b). Color extremely variable from black to yellow-tan, postcoxal line incomplete, apex recurved, directed toward base of first sternum; antennae 11-segmented (Subgenus Scymnus) (Oa, Hi) ...Scymnus ambulans Blackburn, 1895: 252 7b (4b). Postcoxal line complete, recurved, extending to base of first sternum (Subgenus Pullus) ……..8 8a (7b). Body oval, length 1.8-2.2 mm, width 1.1-1.4 mm; head and prothorax yellow-brown to amber, apical 1/3 of elytra yellow-brown to amber (Oa) ………Scymnus (Pullus) dorcatomoides Weise, 1879: 151 8b (7b). Not as above ………9 9a (8b). Body oval, length 1.9-2.2 mm, width 1.2-1.35 mm; head black; pronotum black with lateral angles reddish-yellow to entirely black; anterior margin also black, elytra yellow-brown to amber with black V-shaped mark, point of V 2/3 way or less to apex of elytra ; all legs with dark brown to black femur, tibiae and tarsi lighter in some specimens; sternal segments entirely black (Ma, Hi) ……………….Scymnus (Pullus) suturalis Thunberg, 1795: 106 9b (8b). Not as above ……10 10a (9b). Body oval, widest anterior to middle of elytra; length 1.75-2.4 mm, width 1.45-1.8 mm; head yellow-brown to amber; pronotum yellow to amber with central black spot in shape of dome originating at distal end of prothorax and apex not reaching the front, front margin amber; amber with black triangular area along elytral suture, viewed dorsally as a V-shaped area beginning at base and narrowing as extending anteriorly; leg 1 entirely yellow, legs 2 and 3 yellow with femur reddish-yellow sometimes darker (Ka, Oa, Mo, La, Ma, Hi) …. Scymnus (Pullus) loewii Mulsant, 1850: 980 10b (9b). Not as above ……11 11a (10b). Body oval, length 1.8-2.2 mm, width 1.4-1.75 mm; pronotum yellow to amber on sides with a median black parabolic spot reaching anterior edge, anterior edge may have very narrow margin with color similar to the sides; elytra black except for very narrow amber apex; legs yellow-brown to amber (Oa, Ma, Hi) ……….Scymnus (Pullus) horni Gorham, 1897: 229 11b. (10b). Not as above ……12 12a (11b). Body oval, length 1.6 mm, width 1.25 mm; head yellow to yellowish-brown; pronotum dark brown with anterolateral angle and anterior margin yellowish brown; elytra with 2 yellow to amber spots, anterior spot may expand and connect with posterior spot; legs amber with dark brown spot on femur (Oa, Hi) …..Scymnus (Pullus) latemaculatus Motschulsky, 1858: 121 12b (11b). Head color variable from yellowish-brown to dark brown to black; pronotum may be yellowish-brown with small dark brown to black spot at scutellum to almost the entire pronotum dark brown to black with a yellowish-brown edge at apex; legs yellowish-brown, elytral margins brown to black surrounding a yellow-brown longitudinal band (Oa, Mo, Hi) …..Scymnus (Pullus) uncinatus Sicard, 1924: 532

F. KEY TO THE TRIBES IN THE SUBFAMILY COCCIDULINAE ESTABLISHED IN HAWAII 1a. Body oval; dorsal surface dark greenish-black; male head yellow, female head greenish-back; abdomen with 5 visible sternites ...... Azyini Schilder and Schilder, 1928: 217 (F1.) 1b. Body elongate or oval; dorsal pubescence composed of dense, decumbent hairs with sparse, erect hairs intermixed; head partly concealed beneath pronotum; antenna long, slender, 11-segmented, club serrate; abdomen with 6 visible sternites ...... Coccidulini Costa, 1849: 9, 104 (F2.)

F1. KEY TO THE SPECIES IN THE TRIBE AZYINI ESTABLISHED IN HAWAII 1a. Body oval, length 2.9-4.25 mm, width 2.5-3.45 mm; male head yellow, female head greenish-black; dorsum greenish-black, each elytron with round discal spot; legs same amber to orange-brown color as abdomen; elytral epipleura with deep impressions for reception of legs two and three; pubescence bicolored on elytra, black setae form a single large circular spot on each elytron (Ni, Ka, Oa, Ma, Mo) ………. orbigera Mulsant, 1850: 930

F2. KEY TO THE SPECIES IN THE TRIBE COCCIDULINI ESTABLISHED IN HAWAII 1a. Body oval, length 2.6-3.7 mm, width 1.9-2.3 mm; head, pronotum and elytra entirely black; lateral border of pronotum and elytron discontinuous; elytral pubescence of uniform length and appearance; ventral surface and leg black except abdominal sternites yellow or red (Ni, Ka, Oa, Ma, Mo, La, Hi) …… forestieri (Mulsant, 1853: 158, 286) 1b. Body elongate, oval, length 1.7-2.85 mm, width 1.35-2.0 mm; pronotum light reddish-brown to amber, elytra dark reddish-brown with faint, green metallic tint; elytral pubescence of two lengths: short, fine golden-brown setae and longer erect setae; ventral surface including legs amber (Oa, Ma, Mo, La, Hi) …. (Blaisdell, 1892: 286)

G. KEY TO THE TRIBES IN THE SUBFAMILY ORTALIINAE ESTABLISHED IN HAWAII 1a. Body medium to large, length 2.5-5.0 mm; antennal insertion concealed; antenna 8-segmented, thin, basal 2 segments enlarged, club 3-segmented but not enlarged; apical segment of maxillary palpus large, strongly securiform; 2 tibial spurs; 6 visible sternites ……Noviini Ganglbauer, 1899: 954 (G1.) 1b. Body small to medium-sized, length 2.0-3.5 mm; head not covered by pronotum; antenna 8- segmented, first segment enlarged, about as wide as long, club 3-segmented but not enlarged; tibia without spurs …..Ortaliini Mulsant, 1850: 892 (G2.)

G1. KEY TO THE SPECIES IN THE TRIBE NOVIINI ESTABLISHED IN HAWAII 1. Body elongate; head black; pronotum red except for black band along anterior, elytra red with black elytral suture and black markings, nearly parallel sided, widest at middle; length 2.65-4.18 mm, width 2.0-3.33 mm (Ni, Ka, Oa, Ma, Mo, La, Hi) …..Rodolia cardinalis (Mulsant, 1850: 906)

G2. KEY TO THE SPECIES IN THE TRIBE ORTALIINI ESTABLISHED IN HAWAII 1. Head yellow in male, black in female; yellow pronotum with a median black marking; elytra black; antenna 11-segments; abdomen with 6 ventral segments (Ni, Ka, Oa, Ma, Mo, La, Hi) …Apolinus lividigaster lividigaster (Mulsant, 1853: 285) 2. Body elongate oval, hairy, length 3.4-4.5 mm, width 2.4-3.1 mm; head, prothorax, tip of elytra and abdomen amber to reddish-yellow; mesosternum and metasternum, legs one yellow-brown to orange in males, dark in females; legs two and three dark; pronotum reddish yellow; anterior margin of prosternum lobed, extended forward, at least partially concealing mouthparts; antenna 10 segmented; tarsi 3 segmented; abdomen with 6 ventral segments (Ni, Ka, Oa, Ma, Mo, La, Hi) ….. Cryptolaemus montrouzieri Mulsant, 1853: 140

DEDICATION AND ACKNOWLEDGMENTS This article is dedicated to my wife, Catherine Lindburg Leeper, whose interest, support and encouragement made its writing possible.

It is also dedicated to my graduate professors (John W. Beardsley (my major professor), Henry Bess, Franklin Chang, Mercedes Delfinado, Frank H. Haramoto, D. Elmo Hardy, Wallace C. Mitchell, Ryo Namba, Toshiyuki Nishida, Martin Sherman and Minoru Tamashiro). They provided a well- rounded, diverse grounding in entomology that proved invaluable throughout my professional career.

Bernarr Kumashiro (DoA) and Luc LeBlanc (UH) played an instrumental roles allowing access to the insect collections under their responsibility and in critically reviewing the key.

Gratitude is extended to the Department of Entomology, Plant Pathology and Weed Science at New Mexico State University, Las Cruces, New Mexico for the appointment to an Affiliate Faculty position and providing office space and faculty privileges.

Gratitude is also extended to the College of Tropical Agriculture and Human Resources at the University of Hawaii, particularly to Dr.s Kenneth Grace, Mark Wright, Dan Rubinoff and Ms. Lynnet Higuchi for facilitating a visit to Hawaii to work on this paper. I also wish to thank the Bernice P. Bishop Museum and the Hawaii State Department of Agriculture and their staffs for making their coccinellid collections available for examination.

This work also required considerable use of library services and I wish to recognize the support received from Joyce Komraus, Charles Smith and Charlotte Zimmerman of the Michael Nivison Public Library in Cloudcroft, NM; the Interlibrary Loan Department, Zuhl Library at the New Mexico State University, Las Cruces, NM; and the exceptional services provided by Jodie Mattos of the Hawaiian Collection, Hamilton Library at the University of Hawaii, Manoa.

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