Hoplandria, KRAATZ, 1857

Hanley, Rodney S., 2003, Generic revision of the staphylinid beetle tribe Hoplandriini (Insecta: Coleoptera: Staphylinidae: Aleocharinae), Zoological Journal of the Linnean Society 138 (1), pp. 83-140 : 86-94

publication ID	https://doi.org/ 10.1046/j.1096-3642.2003.00060.x
persistent identifier	https://treatment.plazi.org/id/3F272601-504E-FFF1-C21E-F9F0FE0FF953
treatment provided by	Carolina
scientific name	Hoplandria
status

Treatment

GENUS HOPLANDRIA KRAATZ, 1857 View in CoL ( FIGS 2–11 View Figure 2 View Figure 3 View Figure 4 View Figure 5 View Figure 6 View Figure 7 View Figure 8 View Figure 9 View Figure 10 View Figure 11 )

Hoplandria Kraatz, 1857:4 View in CoL . Kraatz, 1859:9. Fairmaire & Germain, 1861:414. LeConte, 1861:61. Fauvel, 1866:310. LeConte & Horn, 1883:91. Sharp, 1883:219. Eichelbaum, 1909:225. Blatchley, 1910:345, 350. Casey, 1910:174. Cameron, 1919:230. Fenyes, 1918 – 21:306. Notman, 1920:719. Bernhauer & Scheerpeltz, 1926:715. Bradley, 1930:84. Scheerpeltz, 1929:20. Scheerpeltz, 1934:1667. Blackwelder, 1943:557. Black-

[1] Length of adult. [46] Width of apex of ligula of labium. [88] Presence of macrosetae on

[2] General body shape. [47] Shape of apex of ligula of labium. metasternum.

[3] General reflectivity of body. [48] Presence of spines at apex/subapex of [89] Presence of macrosetae on

[4] General punctation and ligula of labium. mesosternum.

microsculpturing of body. [49] Presence and length of medial setae [90] Tarsal formula.

[5] Pubescence concentration on head. on prementum of labium. [91] Shape of tarsal claws.

[6] Pubescence on abdominal terga. [50] Insertion of medial setae of labium. [92] Presence of empodial bristle.

[7] Pubescence on pronotum. [51] Presence of real pores in lateral pore [93] Length of empodial bristle.

[8] Pubescence on elytra. field of labium. [94] Length of tarsal segment 1.

[9] Pubescence on abdominal sterna. [52] Presence of setal pores in lateral pore [95] Length of tarsal segments 2–4.

[10] Head shape, comparison of width to field of labium. [96] Length of tarsal segment 5.

length. [53] Size and shape of median pseudopore [97] Abdominal shape.

[11] Eye size, comparison to length of field of labium. [98] Presence and location of basal head. [54] Composition of lateral pore fields. transverse depressions on

[12] Neck (present or absent). [55] Length of hypoglossal lobe of abdominal terga.

[13] Infraorbital carina development. labium. [99] Presence of macrosetae on abdomen.

[14] Number of antennomeres. [56] Size of setae of hypoglossal lobe of [100] Presence of differentiated region of

[15] Shape of antennal segments 1–3. labium. microsculpturing on anterior corner

[16] Shape of antennal segment 4. [57] Composition of setae of hypoglossal of abdominal sternum III.

[17] Shape of antennal segments 5–10. lobe of labium. [101] Setal pattern on abdominal tergum

[18] Coeloconical sensilla (present or [58] Size of each labial palpus. IX.

absent). [59] Number of segments of each labial [102] Presence of humeral carina or bump

[19] Medial pores of epipharyngeal area of palpus. on each elytron (male secondary labrum, size and distribution. [60] Composition of distal pore field. sexual characteristic).

[20] Definition of longitudinal sensory [61] Length of article 1 of each labial [103] Presence of carina or conical process field. palpus (compared to article 2). on medioapical region of each

[21] Pores between longitudinal sensory [62] Length of article 3 of each labial elytron (male secondary sexual field and lateral sclerotized areas. palpus (compared to article 2). characteristic).

[22] Mandibular asymmetry, right [63] Presence of twin pores of each labial [104] Presence of denticle, carina or bump mandible with distinct median palpus. on outer apical third of each elytron tooth. [64] Presence and number of median pores (male secondary sexual

[23] Shape of mandibular apex. of each labial palpus. characteristic).

[24] Size of condylar molar patch. [65] Shape of apical margin of mentum. [105] Presence of spine at posterior corner

[25] Composition of condylar molar patch. [66] Shape of antero-lateral angles of of abdominal sternum III (male

[26] Arrangement of condylar molar patch. mentum. secondary sexual characteristic).

[27] Subcondylar molar patch (present or [67] General distribution of sensory pores [106] Presence of spine at posterior corner absent). of mentum. of abdominal sternum IV (male

[28] Size of mandibular ‘velvety patch. [68] General shape of pronotum (dorsal secondary sexual characteristic).

[29] Composition of mandibular ‘velvety view). [107] Presence of longitudinal carina on patch. [69] Specific size of pronotum. abdominal tergum IV (male

[30] Setae on ventral aspect of outer basal [70] General shape of pronotum (lateral secondary sexual characteristic). angle of each mandible. view). [108] Presence of longitudinal carina on

[31] Length of each lacinia. [71] Setation of pronotum. abdominal tergum VII (male

[32] Shape of apex of each lacinia. [72] Orientation of setation of pronotum. secondary sexual characteristic).

[33] Length of teeth on adoral margin of [73] Visibility of hypomeron of pronotum [109] Presence of two transverse rows of each lacinia. in lateral view. macrosetae near apex of abdominal

[34] Placement of teeth/hairs on adoral [74] General size of elytra. tergum VIII (male secondary sexual margin. [75] Shape of margin of apico-lateral angle characteristic).

[35] Number of distinct regions of teeth/ of elytron. [110] General shape of aedeagus.

hairs on adoral margin. [76] Size of elytra. [111] Presence of ventral projection on

[36] Number of spinose setae on dorsal [77] Presence of microsetae on each aedeagus.

surface of each lacinea. elytron. [112] Presence of spinules/sclerites on

[37] Width of each galea. [78] General distribution of microsetae on internal sac.

[38] Size of membranous area at apex of each elytron. [113] Length of apical lobe of each each galea. [79] Orientation of microsetation on each paramere.

[39] Composition of setae at apex of each elytron. [114] Setation at apex of apical lobe of each galea. [80] Presence of medial carina on paramere.

[40] Number of articles of each maxillary mesosternum. [115] Shape of anterior margin of palpus (including pseudosegment). [81] Width of separation of mesocoxal paramerite.

[41] Setation of articles 2–3 of each cavities. [116] Modifications to apex of condylite of maxillary palpus. [82] Length of mesocoxal process. paramerite.

[42] Setation of article 1 of each maxillary [83] Presence of isthmus between meso- [117] Length of condylite.

palpus. and metasternal processes. [118] General shape of spermatheca.

KEY TO THE GENERA OF THE TRIBE HOPLANDRIINI OF THE WORLD

1. Mesocoxal cavities broadly separated by broad meso- and metasternal processes ( Fig. 3A View Figure 3 ) ..................................... 2

1¢. Mesocoxal cavities continuous, only partially separated by meso- and metasternal processes; subtribe Platandriina (part) ( Figs 16A View Figure 16 , 21A View Figure 21 ) ................................................................................................................. 4

2. Apex of galea of maxilla with rows of long to very long hairs, typically giving appearance of flowing hairs; subtribe Platandriina (part) ( Fig. 19F View Figure 19 ) ................................................................................................... 3

2¢. Apex of galea of maxilla with rows of short to very short hairs, giving appearance of stubble ( Fig. 6F View Figure 6 ) ............................................................................................................................................................................. 7

3. Mesosternum with a distinct medial carina, not reaching apex of mesosternal process ( Fig. 25A View Figure 25 ); apices of abdominal terga each with 2–4 very large, black, heavy setae ( Fig. 25B View Figure 25 ); abdominal tergum IX without complete crescent-shaped setal pattern; males with abdominal tergum VII with two narrow denticles along midline; known only from the Oriental Region (mainland of Malaysia) .......... Paroplandria

3¢. Mesosternum without a medial carina ( Fig. 20A View Figure 20 ); apices of abdominal terga with typical macrosetae, not large, black, heavy setae; abdominal tergum IX with distinct crescent-shaped setal pattern ( Fig. 20B View Figure 20 ); males without secondary sexual features; known only from the Neotropical Region ( Costa Rica).......................................................................................................................... Ligulata

4. Mesosternum without a distinct medial ridge ( Fig. 22A View Figure 22 ); each labial palpus with three distinct articles; male secondary features not as below .............................................................................................................. 5

4¢. Mesosternum with a distinct, heavy, medial ridge extending to apex of mesosternal process, ridge with numerous, small hairs ( Fig. 18E View Figure 18 ); each labial palpus with articles 1 + 2 fused, only two recognizable segments present (males with secondary sexual features of abdominal tergum VII with four small, narrow denticles along midline and tergum VIII with distinct longitudinal row of setiferous denticles) ( Fig. 18F View Figure 18 ); known only from Afrotropical Region ( Madagascar) Ditropandria subgenus ..................................................................................................................................... Pedandria

5. Males with secondary sexual features somewhat variable, but without a medial, longitudinal carina on either abdominal terga VII or VIII; body very small, length 1.2–2.3 mm; eyes average, 0.4–0.6 times length of head............................................................................................................................................ 6

5¢. Males with abdominal tergum VII with narrow median longitudinal carina and tergum VIII with more or less wide median longitudinal carina; body size average, length 2.4–4.3 mm; eyes small, 0.2–0.4 times length of head; known from Nearctic Region; frequently found in flowers of various shrubs ........ Platandria

6. Abdominal tergum IX without complete crescent-shaped setal pattern, setae absent basally along midline; eyes average, 0.5 times length of head; males without secondary sexual features; known from Nearctic Region (Arizona, North Carolina, Texas, Virginia, northern Mexico); likely found in association with flowers ....................................................................................................................................... Microlia

6¢. Abdominal tergum IX with a distinct crescent-shaped setal pattern; eyes average to large, 0.5–0.7 times length of head; males tergum IV with two small denticles on each side of midline ( Fig. 23F View Figure 23 ) and terga VII– VIII with prominent small, narrow setiferous denticles ( Fig. 23G View Figure 23 ); known from the Oriental Region (Taiwan) and the Palaearctic Region ( Japan) ................................................................ Omoplandria

7. Mesosternum with median longitudinal carina extending to apex of mesosternal process; tarsal formula 5-5-5; males with secondary sexual features, including sinuate areas of apico-lateral angles of elytra often produced into heavy carina-like structures; known from Oriental Region; subtribe Pseudoplandriina ....................................................................................................................... Pseudoplandria

7¢. Mesosternum without median longitudinal carina; tarsal formula 4-5-5 (or 4-4-4 rarely); males without sinuate areas of apico-lateral angles of elytra produced into heavy carina-like structures; subtribe Hoplandriina ................................................................................................................................... 8

8. Pronotum with outer apical angles not sinuate, apical area along midline not extended posteriorly; males with secondary sexual features highly variable, including various combinations of elytra with humeral area with carina or bump, medioapical region with carina or conical process, outer apical third with denticle, carina or bump; abdomen with posterior corners of sterna III and IV produced into spines, tergum IV with raised area or carina, and tergum VII with a narrow posteriomedian longitudinal carina; known from throughout the New World .................... Hoplandria s.l.

8¢. Pronotum with outer apical angles sinuate, apical area along midline extended posteriorly; males with variable secondary sexual features, but without corners of abdominal sternum IV produced into spines and without modifications to tergum IV..................................................................................... 9

9. Extended area along midline apex of pronotum with rounded angles ( Fig. 12H View Figure 12 ); medial setae of prementum with insertions widely separated ( Fig. 12F View Figure 12 ); males with secondary sexual features exclusively limited to each elytron, with small denticle on medioapical area ( Fig. 12I View Figure 12 ), and abdominal tergum VII with narrow posteromedian longitudinal carina or conical carina ( Fig. 12N View Figure 12 ); elytra without modifications of humeral areas; spines absent from abdominal sterna III and IV; tergum IV without modification; known from Neotropical Region ( Ecuador) ......................... Leptandria

9¢. Extended area along midline apex of pronotum with distinctly sharp angles ( Fig. 15A View Figure 15 ); medial setae of prementum with insertions nearly adjacent ( Fig. 13G View Figure 13 ); males with secondary sexual features variable, including medioapical region of elytra with a short, wide carina ( Fig. 13A View Figure 13 ), and corners of abdominal sternum III produced into spines ( Fig. 15E View Figure 15 ); known from Neotropical Region ( Ecuador and Peru)........................................................................................................................ Heliconandria

welder, 1952:191. Moore & Legner, 1975:438. Seevers, 1978:141. Klimaszewski, 1984:10. Génier & Klimaszewski, 1986:202. Klimaszewski & Peck, 1986:83. Génier, 1989:8. Downie & Arnett, 1996:536. Hanley & Ashe, 1998:184. Klimaszewski, 2000:49. Ashe, 2001:304. Hanley, 2002a:301. Hanley, 2002b:121. Hanley, in press b.

Type species: Hoplandria ochracea Kraatz, 1857 [= Hoplandria lateralis ( Melsheimer, 1844) ] (designated by Casey, 1910 by subsequent designation).

Diaboligenus Bierig, 1939:26 View in CoL . Pace, 1990:174. Hanley & Ashe, 1998:184.

Type species: Diaboligenus primus Bierig (1939) by monotypy.

Diagnosis: This genus is distinguishable from the other genera of Hoplandriina by the following combination of characters: body small to very large, 1.7–13.5 mm in length; eyes small to large, 0.3–0.7 times length of head; insertion of medial setae of prementum nearly adjacent to widely separated; pronotum with outer apical angles not sinuate, apical area along midline not extended posteriorly, generally distinctly oval in shape; tarsal formula 4-5-5; males with secondary sexual features highly variable, including various combinations of each elytron with humeral area with carina or bump, medioapical region with carina or conical process, outer apical third with denticle, carina or bump; abdomen with posterior corners of sterna III and IV produced into spines, tergum IV with raised area or carina, and tergum VII with a narrow posteriomedian longitudinal carina.

Description: [1] Lengths of adults 1.7–13.5 mm. [2] Body ( Figs 6A View Figure 6 , 10A View Figure 10 ) fusiform to broadly oval; [3] surface typically glossy with [4] punctation and microsculpturing in many species; [5] pubescence generally infrequent on head and [6] abdominal terga, variably dense on [7] pronotum, [8] elytra, and [9] abdominal sterna.

Head: ( Figs 6A View Figure 6 , 10A View Figure 10 ) [10] Distinctly broader than long. [11] Eyes small to large, 0.3–0.7 times length of head. [12] Neck absent. [13] Infraorbital carina strongly developed and complete to absent. Antenna with [14] 11 antennomeres; [15] articles 1–3 elongate, 1.5–2.5 times longer than wide; [16] article 4 quadrate; [17] articles 5–10 short, 1.3–2.0 times wider than long, [18] coeloconical sensilla present or absent within article 11.

Mouthparts: Labrum ( Figs 2A View Figure 2 , 4A View Figure 4 , 6B View Figure 6 , 8A View Figure 8 , 10B View Figure 10 ) with epipharyngeal area with [19] medial pores large, numerous, more or less uniformly distributed in [20] typically poorly delimited, longitudinal sensory field; [21] pores between longitudinal sensory field and lateral sclerotized areas absent ( Figs 2B View Figure 2 , 4B View Figure 4 , 6C View Figure 6 , 8B View Figure 8 , 10C View Figure 10 ). Mandible ( Figs 2C,D View Figure 2 , 4C,D View Figure 4 , 6D,E View Figure 6 , 8C,D View Figure 8 , 10D,E View Figure 10 ) [22] asymmetrical, right mandible with a distinct median tooth; [23] apex more or less acute and curved adorally; [24] condylar molar patch small, width less than 1/5 of basal mandibular width, [25] composed of very small denticles, [26] densely arranged in irregular transverse rows; [27] subcondylar molar patch absent; dorsobasal ‘velvety patch’ [28] absent to large, size up to wider than half of mandibular base, generally composed of [29] numerous, very fine hairs or spinules; [30] ventral aspect with outer basal angle with 0–4 setae. Maxilla ( Figs 2E View Figure 2 , 4E View Figure 4 , 6F View Figure 6 , 8E View Figure 8 , 10F View Figure 10 ) with [31] length of lacinia generally shorter than galea, [32] lacinia acute apically, [33] teeth on adoral margin short to moderate in length, about 3–5 times longer than wide, and [34] more or less closely placed, into [35] two or three regions and [36] 2 large spinose setae on dorsal surface; [37] galea narrow to wide, narrower to wider than lacinia at widest area, and rounded apically, [38] membranous in apical 1/2–1/3, [39] densely covered with rows of very short, fine hairs giving stubble-like appearance; [40] maxillary palpus with 4 articles and a moderately defined apical pseudosegment, [41] articles 2 and 3 with numerous long setae, [42] articles 1, [43] 4 and [44] pseudosegment without setae. Labium ( Figs 2F View Figure 2 , 4F View Figure 4 , 6G View Figure 6 , 8F View Figure 8 , 10G View Figure 10 ) with ligula [45] elongate, as long or longer than labial palpal segments 1 + 2, [46] slightly broadened apically with [47] shallow to deeply forked apex, [48] most commonly with two apical and two subapical spines; [49] two short or long medial setae of prementum present, generally longer to shorter than ligula, [50] insertions of setae almost adjacent to widely separated; [51] real pores and [52] setal pores present; [53] median pseudopore field narrow and more or less linear, [54] lateral pore field most commonly with a single setose pore and two asetose pores; [55] hypoglossal lobes ( Figs 2G View Figure 2 , 4G View Figure 4 , 6H View Figure 6 , 8G View Figure 8 , 10H View Figure 10 ) generally long, typically reaching beyond base of ligula, [56] with very long (about five times longer than width of lobe), comb-like internally curved setae along [57] entire length of adoral margin. Labial palpus ( Figs 2F View Figure 2 , 4F View Figure 4 , 6G View Figure 6 , 8F View Figure 8 , 10G View Figure 10 ) [58] elongate, overall length 4–15 times longer than greatest width, with [59] 3 distinct articles, [60] with distal pore field composed of one spine and two large pores, with small sensory pegs inside in Hoplandria s. s.; [61] article 1 2.0–7.0 times longer than article 2, [62] article 3 about 2.3–7.0 times longer than article 2; [63] twin pores and [64] two median pores present. Mentum with [65] apical margin flattened to slightly concave with [66] antero-lateral angles obtusely rounded and often extended anteriorly; [67] many sensory pores typically distributed in basal 2/3, however, numerous taxa with pores distrubuted along each side of midline.

Thorax: Pronotum ( Figs 6A View Figure 6 , 10A View Figure 10 ) [68] subquadrate to transverse, variable in size, [69] typically 1.2–1.4 times wider than long, [70] more or less convex to flattened. Setae [71] generally densely distributed, [72] typically directed posteriorly at midline on disc, often with differentiated macrosetae near lateral margins. [73] Hypomera strongly inflexed, not visible in lateral aspect. Each elytron [74] moderately broad, commonly wider apically than basally; [75] apico-lateral angles sinuate; [76] elytra together about 1.3–1.8 times as wide as long; [77] microsetae numerous, [78] generally uniformly distributed and [79] directed posteriorly. Mesosternum ( Figs 3A View Figure 3 , 5A View Figure 5 , 7A View Figure 7 , 9A View Figure 9 , 11A View Figure 11 ) [80] without medial carina. Mesocoxal cavities ( Figs 3A View Figure 3 , 5A View Figure 5 , 7A View Figure 7 , 9A View Figure 9 , 11A View Figure 11 ) [81] widely separated by meso- and metasternal processes by about 1/5 length of coxal cavities. Mesosternal process [82] longer than metasternal process, extended to basal 1/5 of coxal cavities; [83] meso- and metasternal processes typically separated by very short isthmus or contiguous, isthmus absent; [84] mesosternal process rounded or flattened at apex. Metasternum [85] shorter to longer than width of mesocoxae, [86] without medial carina; [87] metasternal process generally flattened at apex. Macrosetae [88] present or absent on metasternum; [89] absent from mesosternum. Legs [90] with tarsal formula 4-5- 5; [91] tarsal claws ( Fig. 7C View Figure 7 ) long and slender, [92] with single empodial bristle, [93] empodial bristle distinctly shorter to distinctly longer than tarsal claws; [94] article 1 of hind tarsus about 1.3 times length of article 2; [95] articles 2–4 subequal in length; [96] article 5 subequal in length to combined lengths of articles 2–4.

Abdomen: ( Figs 3B View Figure 3 , 5B View Figure 5 , 6A View Figure 6 , 7B View Figure 7 , 9B View Figure 9 , 10A View Figure 10 , 11B View Figure 11 ) [97] Fusiform, tapering apically to broadly pointed apex; [98] terga III – V ( VI slightly) with moderate to deep transverse basal depressions on anterior portion. [99] Terga and sterna with prominent macrosetae. [100] Anterior corner of sternum III with distinct region of differentiated microsculpturing, generally delimited by very fine carina, area appearing irridescent in many species. [101] Tergum IX with distinct crescentshaped setal pattern.

Secondary sexual characteristics: Males highly variable, including more or less distinct major and minor forms. Minor forms generally similar to females and typically indistinct externally. Major forms highly variable among species with morphological modifications on elytra in form of [102] humeral carina or bump; denticle, [103] carina or conical process on the medioapical region ( Figs 6A View Figure 6 , 10A View Figure 10 ); [104] well-developed denticle, carina or bump on the outer apical third ( Fig. 5F View Figure 5 ). Abdomen: posterior corners of [105] sternum III and [106] IV produced into long spines often reaching the posterior quarter of tergum IV and V, respectively; [107] tergum IV with more or less U-shaped ( Fig. 6A View Figure 6 ) or narrow longtitudinal carina, often appearing iridescent; [108] tergum VII with narrow posteromedian longtitudinal carina ( Figs 6A View Figure 6 , 10A View Figure 10 ); [109] apex of tergum VIII with two transverse rows of large macrosetae.

Aedeagus: ( Figs 3C View Figure 3 , 5C View Figure 5 , 7D View Figure 7 , 9C View Figure 9 , 11C View Figure 11 ) Highly variable, with bulb of median lobe [110] more or less elongate, subequal to or greater than tube, [111] with distinct ventral projection; internal sac [112] typically with large spinules; [113] paramere with apical lobe of parmerite not greatly extended beyond velum, [114] typically with 2–3 setae at apex, 2 most commonly; [115] paramerite with anterior margin generally slightly to distinctly concave; condylite with [116] apex unmodified to heavily sclerotized, [117] generally shorter to subequal in length to apex of paramerite.

Spermatheca: ( Figs 3E View Figure 3 , 5E View Figure 5 , 7F View Figure 7 , 9E View Figure 9 and 11E View Figure 11 ) [118] Generally L- shaped with three primary divisions: basal bulb, [119] generally simple and rounded at base; neck, [120] significantly bent, commonly at 90∞ angle or greater; and tube, [121] typically membranous, [122] more or less straight.

Habitat: Génier (1989) reported that species of Hoplandria are mostly saprophytic, with some records of adults collected at light traps. Anecdotal collection data suggests that adults of species of Hoplandria s.l. feed on dipteran larvae within various decaying organic materials, especially dung and carrion. Recently, the late instar larvae of H. (Lophomucter) klimaszewskii Génier were described from the US (M. Thayer, J. Ashe & R. Hanley, pers. comm.). These morphologically distinct larvae were generally collected from under oak logs where they typically were very active and fast-moving. Reared individuals were offered Drosophila rearing medium as a food source, but tended to display little interest in eating it. After several days, one larva began to eat Drosophila adults before finally pupating. No other immature stage of any other species of Hoplandria has been described and the mating behaviour of adults remains unknown.

Comments: Hoplandria s.l. is a taxonomically huge genus with hundreds or even thousands of undescribed species especially from the Neotropical Region; 95 species are currently described. Even though some species of Hoplandria are described from outside the New World, my examination of many of these species indicates that they represent other genera, or other aleocharine tribes in some instances. The morphological variation represented within Hoplandria s.l. is extremely variable, especially the secondary sexual features in the males. The function of these features is unknown; however, it seems likely that many structures are used in either male–male competition or resource guarding, while others are associated with glands or outlets of glands.

KEY TO THE SUBGENERA OF THE GENUS HOPLANDRIA 1. Pronotum scarcely to moderately pubescent, often with bare longitudinal area on disc ( Fig. 10A View Figure 10 ); pronotum most often with 6 macrosetae on disc ( Fig. 10A View Figure 10 ); major males with humeral carina on each elytron ( Fig. 10A View Figure 10 )..................................................................................................................................................... 2

1¢. Pronotum moderately to densely pubescent, pubsence generally evenly distributed ( Fig. 6A View Figure 6 ); disc of pronotum typically wthout macrosetae ( Fig. 6A View Figure 6 ); major males without humeral carina on elytra ( Fig. 6A View Figure 6 ) ............................................................................................................................................................ 3

2. Bulb of aedeagus shorter than tube ( Fig. 9C View Figure 9 ) .............................................................................. Lophomucter Notman

2¢. Bulb of aedeagus subequal to or longer than, tube ( Fig. 11C View Figure 11 ) ............................................................ Platonica Sharp

3. Tergum III 3.0–4.5 times wider than length along midline ( Fig. 6A View Figure 6 ); discs of terga IV– VI typically pubescent with fine round punctures ( Fig. 6A View Figure 6 ); body typically yellowish to light brown ................... Hoplandria s. s.

3¢. Tergum III 2.0–2.5 times wider than length along midline; discs of terga IV– VI typically bare and without fine round punctures; body usually darker, light brown to dark brown ........................................................ 4

4. Size generally small (2.5–3.3 mm); general body shape narrowly elongate; major males with a single prominent denticle on outer third of each elytron ( Fig. 5F View Figure 5 ), without posterolateral carina ( Fig. 5F View Figure 5 ); abdominal tergum IV unmodified ...................................................................................... Genosema Notman

4¢. Size large (3.5–5.0 mm); general body shape fusiform; major males with a prominent denticle on apical area along the midline of each elytron ( Fig. 3F View Figure 3 ), and with a posterolateral carina on each elytron ( Fig. 3F View Figure 3 ); abdominal tergum IV with a prominent raised area along midline .............. Arrhenandria Génier

Distribution: AFROTROPICAL REGION. Madagascar. NEARCTIC REGION. Canada (Ontario, Quebec), Mexico (Coahuila, Durango, Nuevo Leon, Oaxaca, Sonora, Tamaulipas), United States (Alabama, Arizona, Arkansas, Connecticut, District of Columbia, Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Kentucky, Louisiana, Maryland, Massachusetts, Michigan, Missouri, New Jersey, New Mexico, New York, North Carolina, Ohio, Oklahoma, Pennsylvania, South Dakota, Tennessee, Texas, Virginia, West Virginia). NEOTROPICAL REGION. Argentina (Tecuman), Bolivia, Brazil (Espirito Santo), Chile, Colombia, Costa Rica, Cuba, Ecuador, Grenada, Guadeloupe, Guatemala, Haiti, Mexico, Panama, Paraguay, Peru, Puerto Rico, Saint Vincent and the Grenadines, Suriname. ORIEN- TAL REGION. China [People’s Republic] (Fujian [Fukien], Sichuan [Szechwan]), Singapore. PALAEARCTIC REGION. Japan.

SUBGENUS ARRHENANDRIA GÉNIER, 1989

( FIGS 2,3)

Hoplandria (Arrhenandria) Génier, 1989:20 View in CoL . Hanley, 2002a:304. Hanley, in press b.

Type species: Hoplandria laeviventris Casey, 1910 (designated by Génier, 1989 by monotypy).

Diagnosis: This subgenus is distinguishable from the other subgenera of Hoplandria by the following combination of characters: body moderate to large (3.5–5.0 mm); body colouration dark throughout; punctation on abdominal terga III – IV with lanceolate ridges impressed laterally and more or less widely separated; males with elytra with prominent denticles on the medioapical margins and prominent posterolateral carina; abdominal tergum IV of males with distinct swelling or flat iridescent elevation.

Description: In agreement with Hoplandria description, except for the following characters. [1] Lengths of adults 3.5–5.0 mm. [2] Body elongate with sides broadly arcuate; [3] surface more or less glossy with [4] prominent punctation and microsculpturing; [5] pubescence moderately frequent on head and [6] abdominal terga, more or less dense on [7] pronotum, [8] elytra, and [9] abdominal sterna.

Head: [10] Slightly broader than long. [11] Eyes moderate in size, 0.3–0.5 times length of head. [13] Infraorbital carina weakly developed. Antenna with [16] article 4 slightly longer than wide; [18] coeloconical sensilla present in article 11.

Mouthparts: Labrum ( Fig. 2A View Figure 2 ) with epipharyngeal area with [19] medial pores large, numerous; [21] pores between longitudinal sensory field and lateral sclerotized areas few, irregularly arranged ( Fig. 2B View Figure 2 ). Mandible ( Fig. 2C,D View Figure 2 ) with [23] apex more or less rounded with slight curve adorally; condylar molar patch with [26] densely, irregularly arranged denticles; dorsobasal ‘velvety patch’ [28,29] absent; [30] ventral aspect with outer basal angle with two small setae; [33] teeth of adoral margin of lacinia short, each about 3 times longer than wide. Maxilla ( Fig. 2E View Figure 2 ) with teeth on adoral margin divided into [35] three more or less distinct regions; [37] galea moderately narrow, subequal to width of lacinea at widest area, and rounded apically. Labium ( Fig. 2F,G View Figure 2 ) with ligula [45] elongate, longer than labial palpus 1 + 2, with [47] moderately forked apex; [49] two long medial setae of prementum present, typically longer length of ligula, [50] insertions of setae almost adjacent. Labial palpus ( Fig. 2F View Figure 2 ) [58] elongate, overall length 5–6 times longer than greatest width, [60] with distal pore field composed of two large pores and small sensory pegs; [61] article 1 about 4.0 times longer than article 2, [62] article 3 about 3.0 times longer than article 2. Mentum with [65] apical margin concave.

Thorax: Pronotum [68] distinctly transverse. Elytra with [75] apico-lateral angles distinctly sinuate; [76] elytra together about 1.3 times wider than long. Mesocoxal cavities ( Fig. 3A View Figure 3 ) [81] widely separated by meso- and metasternal processes by about 1/4 length of coxal cavities. Mesosternal process length [82] subequal to length of metasternal process; [83] meso- and metasternal processes generally contiguous, isthmus absent; [84] mesosternal process generally rounded at apex, [85] shorter than width of mesocoxa. Macrosetae [88] present on metasternum. Legs with [93] empodial bristle distinctly shorter than tarsal claws, but over half of length of tarsal claws.

Abdomen: ( Fig. 3B View Figure 3 ) [100] Anterior corner of sternum III with distinct region of differentiated microsculpturing, generally delimited by very fine carina, area appearing irridescent.

Secondary sexual characteristics: Males highly variable, including more or less distinct major and minor forms. Minor forms generally similar to females and typically indistinct externally. Major forms highly variable among species with morphological modifications of elytra in form of [102] without humeral carina or bump; [103] prominent denticle, carina or conical process on the medioapical region; [104] without denticle, carina or bump on the outer apical third; [104a] prominent posterolateral carina on each elytron. Abdomen: posterior corners of [105] sterna III and [106] IV produced into long, bent spines often reaching posterior quarter of terga IV and V, respectively; [107] tergum IV with more or less large swollen area; [108] tergum VII with highly elevated and acute narrow carina; [109] apex of tergum VIII without two transverse rows of large macrosetae.

Aedeagus: ( Fig. 3C View Figure 3 ) Bulb of median lobe [110] more or less irregularly shaped, shorter than tube, [111] with moderately developed ventral projection; condylite ( Fig. 3D View Figure 3 ) [117] generally subequal in length to apex of paramerite.

Spermatheca: ( Fig. 3E View Figure 3 ) Basal bulb [119] generally simple and rounded at base with a distinct depression at apex.

Habitat: Specimens of H. (Arrhenandria) laeviventris , the only species in this subgenus, have been collected from various sources including: dead mice, carrionbaited pitfall traps, and through sifting unspecified leaf litter, rhododendron litter, mushroom litter, leaf litter in deep crevices and berlese samples or leaf debris, oak-hickory debris, dry oak litter, or mixed hardwood forest litter. The collecting period for this species is from February to October.

Comments: As Génier (1989) pointed out, the establishment of Arrhenandria as a subgenus of Hoplandria is based on the distinctive secondary sexual characters, especially the presence of a lateral carina on each elytron and a raised swelling on tergum IV. Both of these structures are absent in the subgenus Genosema , the sister group to Arrhenandria. In addition, the absence of microsculpure on the swollen area of tergum III of the abdomen also supports a close relationship with Genosema . Arrhenandria contains only one species, H. (Arrhenandria) laeviventris Casey from the midwestern United States.

Distribution: NEARCTIC REGION. United States (Alabama, Arkansas, Connecticut, District of Columbia, Georgia, Illinois, Indiana, Kentucky, Louisiana, Maryland, Massachusetts, New Jersey, New York, North Carolina, Ohio, Pennsylvania, Tennessee, Texas, Virginia, West Virginia).