Scaphisoma

Ivan Löbl & Ryo Ogawa, 2016, On the Scaphisomatini (Coleoptera, Staphylinidae, Scaphidiinae) of the Philippines, IV: the genera Sapitia ACHARD and Scaphisoma LEACH, Linzer biologische Beiträge 48 (2), pp. 1339-1492 : 1343-1351

5.2.1. Key to the Philippine species of Scaphisoma LEACH, 1815

1 Elytra with sutural striae starting posterior level of scutellar tip. Elytral punctation entirely or on parts of disc conspicuously coarse.............................................................2

- Elytra with sutural striae starting at or anterior level of scutellar tip. Elytral punctation variable, often fine .......................................................................................23

2 Elytra with coarse discal punctures to part arranged to form longitudinal or oblique rows .................................................................................................................................3

- Elytra with discal punctures irregular, not arranged to form rows.................................12

3 Elytra prevailing ochreous, each with dark transverse spot situated posterior elytral mid-length (Fig. 37)....................................................................... S. transversale nov.sp.

- Colour pattern of elytra different .....................................................................................4

4 Elytra uniformly ochreous, about as light as apical abdominal segments and lighter than thorax ( Fig. 24). Pronotum slightly darkened along basal margin ............................. ........................................................................................................ S. ochropenne nov.sp.

- Entire or prevailing elytral surface dark reddish-brown to blackish ................................5

5 Apical sixth to third of elytra light, yellowish or ochreous, well delimited from prevailing dark elytra surface and usually impunctate or very finely punctate................6

- Apical area of elytra about as dark as most of elytral disc, or distinctly darkened, or with very narrow light apical border, usually distinctly punctate ..................................10

6 Coarse elytral punctation abruptly ending at or slightly posterior elytral mid-length, punctation on most of apical halves of elytra very fine or absent ( Fig. 29). Aedeagus with apical section of parameres expanded ........................................... S. rufescens (PIC)

- Coarse elytra punctation reaching up to or onto apical third of elytra ( Fig. 32). Aedeagus with parameres not expanded apically ............................................................7

7 Aedeagus with internal sac bearing long, hook-like rod overlapping simple rod, and robust spine-like structures in basal part.................................... S. stigmatipenne HELLER

- Aedeagus with internal sac lacking rods, without robust basal spine-like structures.......8

8 Aedeagus with parameres curved, bearing membranous lobes, apical process of median lobe oblique in lateral view.................................................................................9

- Aedeagus with parameres sinuate, lacking membranous lobes, apical process of median lobe convex in lateral view ( Figs 174, 175 ) ........................... S. sinuatum nov.sp.

9 Apex of median lobe acute ( Fig. 167 ) ........................................... S. luteopygidiale (PIC)

- Apex of median lobe blunt ( Fig. 161 )................................................ S. banguiense LÖBL

10 Aedeagus with apical halves of parameres broad ( Fig. 171, 173 ). Prevailing surface of elytra reddish-brown, about as pronotum, each elytron with darkened transverse area anterior very narrow light apical rim ( Fig. 28). Pronotal punctation conspicuously coarser and denser along basal margin than on remaining pronotal surface ................................................................................... S. pseudokalabitum nov.sp.

- Aedeagus with apical halves of parameres narrow. Elytra reddish-brown to blackish- brown, not or weakly darkened subapically, usually without light apical rim. ..............11

11 Aedeagus with parameres abruptly narrowed proximally mid-length; apical process of median lobe gradually narrowed ( Fig. 177 ). Pronotum with punctation denser and coarser along basal margin than on remaining surface ......................... S. surigaosum PIC

- Aedeagus with parameres gradually narrowed toward mid-length; apical process of median lobe abruptly narrowed toward tip ( Fig. 163 ). Pronotum with almost even, sparse and very fine punctation, or with somewhat coarser punctures on basomedian area ............................................................................................. S. inexspectatum nov.sp.

12 Small species, with body 1.35-1.50 mm long. Punctation on prevailing surface of elytra coarse, with punctures to part larger than puncture intervals. Male mesotibiae not or hardly thicker than metatibiae. Females with apical sutural angle of elytra rounded..........................................................................................................................13

- Body length exceeding 1.70 mm. Prevailing or entire surface of elytra with punctures smaller than, or as large as puncture intervals. Male mesotibiae usually distinctly thicker than metatibiae. Female elytra with apical sutural angle denticulate or rounded..........................................................................................................................15

13 Hypomera with strigulate microsculpture. Metaventrite lacking antecoxal puncture rows .................................................................................................... S. furcatum nov.sp.

- Hypomera lacking obvious microsculpture. Metaventrite with antecoxal puncture rows...............................................................................................................................14

14 Parameres of aedeagus arcuate in apical halves ( Fig. 204 ). Punctation on pronotal sides much coarser and denser that on mesal area ........................... S. furcigerum nov.sp.

- Parameres sinuate in apical halves ( Fig. 207 ). Punctation on pronotal sides and on pronotal middle similar.................................................................... S. furcillatum nov.sp.

15 Pronotal punctation conspicuously dense and fairly coarse, with many punctures about as large as puncture intervals. Mesanepisterna, often also epimera and metanepisterna with distinct, comparatively coarse punctation ...... S. signaticolle nov.sp.

- Punctation on pronotum, epimera, mesanepisterna and metanepisterna fine, often hardly visible, or distinct on pronotal lobe, consisting of punctures much smaller than puncture intervals...................................................................................................16

16 Elytra, in addition to light apical fourth, lightened on subhumeral areas, with or without lightened middle parts of disc. Pronotum in both sexes bicoloured..................17

- Elytra uniformly reddish-brown to blackish between bases and light apical areas. Pronotum bicoloured or unicoloured .............................................................................19

17 Female with inner apical margins of elytra rounded, not denticulate. Male with apical process of median lobe long, widened and truncate at tip ( Figs 249, 250 ) ........................ ........................................................................................................ S. trifurcatum nov.sp.

- Female with inner apical margins of elytra denticulate. Male with tip of apical process of median lobe acute (lateral view), or apical process very short .....................18

18 Elytra each with large, well delimited light subhumeral spot. Basomedian area of pronotum conspicuously punctate. Female with apical margins of elytra weakly oblique......................................................................................... S. trimaculatum nov.sp.

- Elytra each with light subhumeral area extended apically and joint or almost joint to light apical area. Punctation on basomedian area of pronotum not or slightly coarser than that on remainder of pronotal disc. Female with apical margins of elytra conspicuously oblique ................................................................ S. tricoloripenne nov.sp.

19 Male pronotum uniformly dark brown or reddish-brown as anterior two thirds or three fourth of elytra, with or without small light laterobasal area ................................20

- Male pronotum ochreous on prevailing surface.............................................................21

20 Apical process of median lobe with ventral branch short, curved, acute at tip; dorsal branch reaching far posterior its mid-length ( Fig. 245 ) ......................... S. tridens nov.sp.

- Apical process of median lobe with ventral branch long, oblique, truncate at tip; dorsal branch reaching about to its mid-length ( Fig. 252 ) ................... S. trilobum nov.sp.

21 Male with pronotum narrowly darkened along basal margin, remaining pronotal surface as that of hypomera ochreous (Fig 38). Ventral branch of apical process of median lobe strongly inflexed ventrally, weakly curved and weakly narrowed toward blunt apex ( Fig. 235 ) ........................................................................... S. tricolor HELLER

- Pronotum darkened along basal margin, with darkened area expanded mesally. Ventral branch of median lobe moderately inflexed ventrally, strongly curved and narrowed apically ..........................................................................................................22

22 Pronotum with darkened area not or hardly extended anterior lobe. Ventral branch of median lobe of aedeagus abruptly bent, with truncate apex ( Figs 238, 239 ) ..................... ...................................................................................................... S. tricoloratum nov.sp.

- Pronotum with darkened mesal area extended at least to pronotal mid-length, or up to anterior pronotal margin (Fig. 40). Ventral branch of median lobe of aedeagus broadly bent and with acute tip ( Figs 240, 241 ) ........................ S. tricolorinotum nov.sp.

23 Elytra with sutural striae strongly converging from bases to apices, or from bases to mid-length, often angular near elytral bases; adsutural areas conspicuously broad near scutellum................................................................................................................24

- Elytra with sutural striae parallel or weakly converging from level of scutellar tip to sutural mid-length or up to apices, not angular in anterior section, often curved near bases; adsutural areas narrow near scutellum ..................................... 32

24 Elytra with lateral margin striae conspicuously coarsely punctate ................................25

- Elytra with lateral striae impunctate or finely punctate .................................................27

25 Submetacoxal areas very narrow, about as tenth of interval to apical margin; submetacoxal lines hardly convex. Elytra dark, each with light subapical area................. ............................................................................................................. S. subanun nov.sp.

- Submetacoxal areas moderately large, about as fifth to fourth of interval to apical margin; submetacoxal lines distinctly convex. Elytral coloration different...................26

26 Elytra darkened along bases, in middle and along apices (Fig. 8). Metaventrite with one central and two apicomedian impressions.................................. S. casiguran nov.sp.

- Elytra uniformly reddish or blackish-brown, light apices excepted (Fig. 9). Metaventrite with apicomedian impressions, lacking central impression.......................... ............................................................................................................. S. cuyunon nov.sp.

27 Metaventrite with antecoxal puncture row, microsculpture distinct on apicolateral parts or on entire lateral parts of metaventrite ...............................................................28

- Metaventrite without antecoxal puncture row, microsculpture evanescent near lateral margins of metaventrite .................................................................................................29

28 Prevailing body colour ochreous, metaventrite and mesoventrite dark brown to blackish, elytra darkened along suture, sutural striae, at bases and usually also in middle areas. Mesanepisterna not microsculptured. Punctation on basal part of elytra near sutural striae and on adsutural areas coarser than that on prevailing elytral surfaces..........................................................................................................................30

- Prevailing body colour reddish-brown ( Fig. 4 ). Mesanepisterna microsculptured. Punctation on basal parts of elytra, near sutural striae and on adsutural areas not coarser than on prevailing elytral surfaces.................................. S. bicoloripenne nov.sp.

29 Basomedian parts of metaventrite and sternite 1 very finely punctate. Female with acute, prominent inner apical angles of elytra ...................................... S. werneri nov.sp.

- Basomedian parts of metaventrite and sternite 1 with fairly coarse punctures. Female with rounded, not prominent inner apical angles of elytra................................................. ...................................................................................................... S. biplagiatum HELLER

30 Body light reddish-brown, apical fourth to third of elytra lighter than remaining elytral surface (Fig. 10) ............................................................................. S. dispar LÖBL

- Body dark reddish to blackish-brown, apical tenth to eight of elytra lighter than remaining elytral surface ...............................................................................................31

31 Internal sac of aedeagus with mesal row of triangular sclerites, parameres widened apically, with inner margins notched................................................ S. mindanaosum PIC

- Internal sac of aedeagus lacking row of triangular sclerites, apical parts of parameres not widened and lacking subapical notches ( Figs 196-198 ).............. S. disparides nov.sp.

32 Elytra dull, microsculptured ................................................................. S. opacum nov.sp.

- Elytra shiny, not microsculptured..................................................................................33

33 Mesepimera fused, indistinct .........................................................................................34

- Mesepimera distinct.......................................................................................................4 0

34 Abdominal sternite 1 with submetacoxal areas usually conspicuously large. Median part of metaventrite coarsely and densely punctate. Elytral disc often somewhat flattened, usually smooth at base and along lateral margin, often coarsely and conspicuously densely punctate posterior smooth basal area ........................................35

- Abdominal sternite 1 with submetacoxal areas moderately large, much shorter than intervals to apical margin. Median area of metaventrite lacking coarse punctures. Elytral disc convex, elytral punctation sparse and fine..................................................39

35 Abdominal sternite 1 with submetacoxal areas larger than intervals to apical margin. Antennomeres III and IV combined usually about as long as or shorter than V ............36

- Abdominal sternite 1 with submetacoxal areas as long as half of interval to apical margin. Antennomeres III and IV combined longer than V..................... S. simplex LÖBL

36 Prevailing elytral punctation fine, about as fine as pronotal punctation; elytra coarsely punctate on small, flattened and elongate lateral area of elytral disc. Antennomeres IV short, about as long as III ( Fig. 52 )................ S. subpunctulum nov.sp.

- Prevailing elytral punctation much coarser than pronotal punctation. Antennomere IV usually distinctly longer than III...............................................................................37

37 Elytral disc with sharply delimited patch of coarse and conspicuously dense punctures; humeral areas and apical third of elytra very finely punctate, appearing impunctate. Aedeagus with base of apical process of median lobe fairly wide, gradually narrowed in dorsal view, not reaching level of apical third of parameres.......... ........................................................................................................... S. subconvexum PIC

- Elytral disc with punctures distinct near apical margin .................................................38

38 Aedeagus with parameres conspicuously long and narrow, extending beyond level of tip of median lobe, latter with apical process conspicuously narrow and long, parallel-sided in lateral view ( Figs 48, 49 ) ....................................... S. subgracile nov.sp.

- Aedeagus with parameres short and comparatively wide, widened at apices, not reaching level of tip of median lobe, latter with apical process moderately long and gradually narrowed in lateral view ( Figs 50, 51 ) .......................... S. subplanatum nov.sp.

39 Aedeagus with symmetrical, lobed parameres ( Figs 125, 126 ) .... S. inconventum nov.sp.

- Aedeagus with asymmetrical, not lobed parameres, left paramere strongly widened basally............................................................................................... S. imuganense LÖBL

40 Elytra with sutural striae distinctly curved along margin of pronotal lobe and extended laterally along elytral bases to form basal striae.............................................41

- Elytra with sutural striae starting at margin of pronotal lobe, usually weakly curved, not extended along bases to form basal striae................................................................50

41 Antennomere VIII about as long as half of VII or IX and about 1.5 times as long as wide. Sternite 1 with submetacoxal lines and areas reduced, indistinct. Elytral and pronotal punctation similar, fine....................................................... S. breviatum nov.sp.

- Antennomere VIII much longer than half of VII or IX and more than twice as long as wide. Sternite 1 with submetacoxal lines and areas distinct. Elytral punctation often entirely or to large extend coarser than pronotal punctation ..........................................42

42 Antennomere III and IV similar in length......................................................................43

- Antennomere III much shorter than IV, both combined as long as or longer than V.....45

43 Antennomere VI slightly longer than III to V combined ................ S. minutipenis nov.sp.

- Antennomere VI shorter than III to V combined...........................................................44

44 Abdomen with strigulate microsculpture.............................................. S. kodadai nov.sp.

- Abdomen not microsculptured ................................................................. S. elpis nov.sp.

45 Apicolateral parts of metaventrite conspicuously coarsely and densely punctate. Body length 2.55 mm. Abdomen with distinct punctulate microsculpture ........................ ............................................................................................................ S. lienhardi nov.sp.

- Body length 1.45-2.10 mm. Lateral parts of metaventrite very finely punctate.............46

46 Apical sixth to fourth of elytra conspicuously light, reddish to yellowish, or with transverse yellowish band, remainder of elytra dark brown to blackish ........................47

- Elytra entirely reddish-brown or blackish-brown ..........................................................48

47 Body length 1.70-2.10 mm. Antennomeres III and IV combined longer than V, latter much shorter than VI ......................................................................... S. scapulare nov.sp.

- Body length 1.45-1.55 mm. Antennomeres III and IV combined about as long as V, latter as long as VI....................................................................... S. bicuspidatum nov.sp.

48 Submetacoxal lines parallel. Apex of median lobe broadly rounded in dorsal view ( Fig. 120).............................................................................................. S. scurrile nov.sp.

- Submetacoxal lines convex. Apex of median lobe acute in dorsal view........................49

49 Median lobe without prominent articular process. Parameres gradually narrowed and arcuate in dorsal view, lacking membranous lobes............................ S. biliranense LÖBL

- Median lobe with prominent, narrow and curved articular process. Parameres sinuate, with inner membranous lobe ( Figs 133, 134 ) ...................... S. pandanum nov.sp.

50 Pronotum brown with large ochreous lateral spots. Hypomera ochreous. Body length 2.25 mm............................................................................................. S. pulchrum nov.sp.

- Pronotum lacking spots, pronotal colour similar to that of hypomera darker. Usually smaller species with body length inferior to 2.0 mm .....................................................51

51 Pronotum dark brown to blackish. Elytra ochreous, each with well delimited dark central spot and darkened areas along base, suture, lateral margin and apex. Entire metaventrite with strigulate microsculpture............................................ S. mirum nov.sp.

- Colour pattern of body different. Lateral parts of metaventrite or entire metaventrite often lacking microsculpture .........................................................................................52

52 Thorax, hypomera excepted, and basal halves of elytra black or blackish, apical halves of elytra ochreous to yellowish, well delimited ( Fig. 3 ) ............. S. bicolor nov.sp.

- Colour pattern of body different ....................................................................................53

53 Thorax ochreous, anterior two thirds of elytra black, most of apical third of elytra yellowish. Elytral punctation hardly visible ..................................... S. distinctum nov.sp.

- Colour pattern of body different ....................................................................................54

54 Prevailing elytral surface finely punctate, each elytron with small laterocentral area conspicuously coarsely and densely punctate................................................................55

- Elytral punctation different............................................................................................56

55 Apical areas of elytra slightly darkened. Mesepimera subtriangular, about as third of interval to mesocoxae ........................................................................... S. tagalog nov.sp.

- Apical areas of elytra not darkened. Mesepimera elongate, almost as intervals to mesocoxae .............................................................................. S. centripunctulum nov.sp.

56 Hypomera, mesanepisterna and metanepisterna conspicuously punctate, with punctures larger than pronotal punctures. Body blackish-brown....................................... ......................................................................................................... S. deharvengi nov.sp.

- Hypomera, mesanepisterna and metanepisterna impunctate or very finely punctate, with punctures not larger than those on pronotum.........................................................57

57 Body dark brown to black, elytra each with distinct subapical yellowish or ochreous fascia; area posterior subapical fascia brown to blackish, not or only somewhat lighter than middle of elytral disc ( Fig. 17 ) ...................................................................58

- Colour pattern of body different ....................................................................................60

58 Aedeagus with internal sac bearing rows of long, narrow, spine-like structures ( Figs 91, 92 )............................................................................................ S. lunabianum nov.sp.

- Aedeagus with internal sac bearing membranous, minute, mostly very short denticular and scale-like structures................................................................................59

59 Aedeagus with apical process of median lobe gradually narrowed in dorsal view, ventral side curved in lateral view, with apex acute ( Figs 72, 73 ) ........ S. binaluanum PIC

- Aedeagus with apical process of median lobe widest posterior base, sinuate in lateral view, apex blunt ( Figs 101, 102 ) ................................................... S. nabiluanum nov.sp.

60 Pronotal punctation coarse and very dense, with punctures well delimited, to part as large as puncture intervals and only slightly smaller than punctures on elytral disc. Antennomere IV short, about as long as III................................................ S. hiekei LÖBL

- Pronotal punctation entirely very fine, with punctures distinctly smaller than puncture intervals or punctures in middle of disc finer than near base; if pronotal and elytral punctation similar. Antennomere IV longer than III...........................................61

61 Submetacoxal areas reduced, about 0.02 mm wide; submetacoxal lines parallel, with coarse punctures; body length exceeding 1 mm ............................................................62

- Submetacoxal areas usually exceeding 0.02 mm; submetacoxal lines entirely or at least partly convex, often finely punctate, if submetacoxal lines subparallel, body length not exceeding 1 mm............................................................................................66

62 Abdomen with well visible strigulate microsculpture. Metaventrite with two apicomedian impressions. Aedeagus symmetrical.........................................................63 Abdominal microsculpture punctulate. Metaventrite without apicomedian impressions. Aedeagus asymmetrical ............................................................................64

63 Body length inferior 1.35 mm, prevailing body colour blackish .... S. compactum nov.sp.

- Body length exceeding 1.80 mm, prevailing body colour ochreous .................................. ......................................................................................................... S. angulosum nov.sp.

64 Aedeagus in dorsal view with right paramere evenly broad, truncate at apex ( Fig. 127 ) ............................................................................................ S. inopportunum nov.sp.

- Aedeagus in dorsal view with tip of right paramere expanded by membranose lobe ....65

65 Aedeagus in dorsal view with right paramere narrowed subapically, widened and rounded at tip ( Fig. 131 ) ....................................................................... S. palawanum PIC

- Aedeagus in dorsal view with right paramere gradually widener up to apical membranous lobe ( Fig. 129 )............................................................... S. oviforme nov.sp.

66 Antennomere VI about as long as or longer than III to V combined .............................67

- Antennomere VI shorter than antennomeres III to V combined ....................................68

67 Body length 1.15 mm, body light brown. Metaventrite with median impression .............. ........................................................................................................ S. hexamerum nov.sp.

- Body length 1.23 mm, body blackish. Metaventrite without impression........................... ....................................................................................................... S. apomontium nov.sp.

68 Elytra each with large reddish or ochreous subbasal spot, dark brown to blackish in middle of disc, either with entire apical area ochreous to yellowish, or with light subapical spot or fascia. Pronotum reddish-brown to blackish ......................................69

- Elytra with different colour pattern ...............................................................................71

69 Antennomere IV very short, slightly longer than III, V much longer than III and IV combined .................................................................................... S. quadrimaculatum PIC

- Antennomere IV much longer than III, V not or slightly longer than III and IV combined .......................................................................................................................70

70 Entire apical third of elytra ochreous to yellowish ( Fig. 18 ). Median lobe of aedeagus trifid............................................................................................... S. luteomaculatum PIC Elytra each with transverse, light, translucent fascia posterior mid-length and narrowly lightened apices, very dark in-between........................ S. subfasciatum PIC

71 Mesanepisterna, lateral parts of metaventrite and of sternite 1 iridescent, with distinct strigulate microsculpture. Body length 1.85 mm .............................. S. iridescens nov.sp.

- Ventral side of body not iridescent, mesanepisterna not microsculptured .....................72

72 Inner apical angle of elytra denticulate. Body length 1.9 mm. Abdomen with strigulate microsculpture .................................................................... S. dentipenne LÖBL

- Inner apical margins of elytra not denticulate. Body length usually inferior to 1.9 mm. Abdominal microsculpture present or absent ........................................................73

73 Body light brown, elytra each with large spot covering most of disc ( Fig. 20 ). Small species 1.0-1.10 mm long, pronotal and elytral punctation very fine. Submesocoxal and submetacoxal areas similar, with strongly convex lines........... S. maculosum nov.sp.

- Colour pattern of body different. Species often larger than 1.1 mm, elytral punctation often distinctly coarser than pronotal punctation, submesocoxal and submetacoxal areas usually dissimilar, with moderately convex lines .................................................74

74 Elytra dark brown to blackish, each with well delimited, yellowish transverse fascia situated posterior elytral mid-length ................................................ S. fascipenne nov.sp.

- Colour pattern of elytra different ...................................................................................75

75 Abdomen with strigulate microsculpture. Metaventrite with or without microsculpture ...............................................................................................................76

- Abdomen not microsculptured, or with punctulate microsculpture, metaventrite not microsculptured .............................................................................................................9 9

76 Metaventrite with antecoxal puncture row.....................................................................77

- Metaventrite without antecoxal puncture row ...............................................................86

77 Metaventrite with mesal stria ...................................................... S. hamatum nov.sp.

- Metaventrite without median stria .................................................................................78

78 Elytra with punctures on middle part of disc coarse, to part as large or larger than puncture intervals. Elytra ochreous, lighter than pronotum, yellowish near apices ........... ........................................................................................................... S. confusum nov.sp.

- Elytral disc with discal punctures smaller that puncture intervals. Elytra often dark and not yellowish in apical section ................................................................................79

79 Small species, body 1.0-1.30 mm long. Median lobe of aedeagus symmetrical, strongly bent apically, near tip vertical to aedeagal axis, parameres lobed........................ ................................................................................................................ S. javanum LÖBL

- Medium-sized species, body 1.5-1.8 mm long ..............................................................80

80 Aedeagus with symmetrical median lobe ......................................................................81

- Aedeagus with asymmetrical median lobe ....................................................................82

81 Median lobe bifid, with dorsal branch subtriangular, ventral branch gradually narrowed. Internal sac lacking apical bunch of long spines.................... S. boettcheri PIC

- Median lobe trifid, with dorsal branches wide and weakly sclerotized, ventral branch abruptly narrower toward tip. Internal sac with bunch of long spines ( Figs 180, 181 ) ...... ............................................................................................................ S. acutulum nov.sp.

82 Aedeagus with parameres lacking lobes, apical process of median lobe with asymmetrically bent tip ( Figs 182, 183 ) .....................................S. apomontanum nov.sp.

- Aedeagus with lobed parameres ....................................................................................83

83 Apex of median lobe broad. Parameres each with subbasal and apical lobes................84

- Apex of median lobe narrow. Parameres each with single, apical lobe .........................85

84 Tip of apical process of median lobe vertical to axis of basal bulb; inner margin of parameres crenulate ( Figs 225, 227 )............................................... S. spatuloides nov.sp.

- Apical process of median lobe obliquely inflexed; inner margin of parameres not crenulate ( Figs 219, 220 ) ........................................................................ S. duplex nov.sp.

85 Tip of apical process of median lobe narrowed, parameral base not wider than parameral lobe in lateral view ( Figs 216, 217 ) .................................... S. bilobum nov.sp.

- Tip of apical process of median lobe widened, parameral base wider than parameral lobe in lateral view ( Figs 221, 222 ) ................................................. S. nishikawai nov.sp.

86 Male mesotarsomeres 1 about twice as wide as apices of mesotibiae. Punctures on elytral disc sharply delimited. Internal sac of aedeagus lacking sclerites ( Figs 83, 84 ) .......................................................................................... S. densepunctatum nov.sp.

- Male mesotarsomeres 1 about as wide as, or narrower than apices of mesotibiae. Punctures on elytral disc often poorly delimited. Internal sac of aedeagus often bearing sclerites.............................................................................................................87

87 Elytral disc with patch of dense and coarse punctures, contrasting prevailing very fine and sparse elytral punctation ..................................................................................88

- Elytra lacking distinctive patch of coarse punctures....................................................89 t

88 Elytral patch of coarse and dense punctures situated on lateral part of disc ...................... ...................................................................................................... S. simplexoides nov.sp.

- Elytral patch of dense and coarse punctation situated on on inner anterior and middle parts of disc ....................................................................................... S. obscurum nov.sp.

89 Antennomere VI as long as antennomeres IV and V combined and as long as VII........... .................................................................................................... S. hexameroides nov.sp.

- Antennomere VI shorter than antennomeres IV and V combined and usually either shorter or longer than shorter than VII ..........................................................................90

90 Pronotum and most of elytra black, apical fourth of elytra yellowish ...........................91

- Pronotum and elytra blackish to ochreous, elytra uniformly or almost uniformly coloured .........................................................................................................................92

91 Body length hardly 1.3 mm. Submetacoxal areas as fourth of intevals to apical margin of sternite 1............................................................................ S. maramag nov.sp.

- Body length hardly 1.9 mm. Submetacoxal areas as tenth of intevals to apical margin of sternite 1........................................................................................... S. alticola nov.sp.

92 Body length 2.2 mm. Pronotal and elytral punctation evenly fine. Metaventrite lacking microsculpture .................................................................... S. glabrellum nov.sp.

- Body length 1.0-1.7 mm. Elytral punctation usually distinctly coarser than pronotal punctation. Mesal part of metaventrite often with strigulate microsculpture.................93

93 Aedeagus with asymmetrical parameres ( Figs 155, 156 ) ........ S. dissymmetricum nov.sp.

- Aedeagus with symmetrical parameres .........................................................................94

94 Aedeagus with conspicuously lobed parameres.................................. S. laminatum LÖBL

- Parameres of aedeagus not lobed...................................................................................95

95 Aedeagus minute, 0.28 mm long, with parameres strongly narrowed posterior basal third (dorsal view) ( Fig. 99 )........................................................... S. montivagum nov.sp.

- Aedeagus 0.46-0.90 mm long, with parameres usually not narrowed posterior basal third ...............................................................................................................................96

96 Apical process of aedeagus lacking subapical denticles ................................................97

- Apical process of aedeagus with two subapical denticles (lateral view)........................98

97 Elytral disc punctation even fine, punctures not well delimited, much smaller than puncture intervals; internal sac of aedeagus with flagellum ( Fig. 69 )................................ ....................................................................................................... S. caudatoides nov.sp.

- Elytral disc with punctation uneven, punctures near inner basal angles to part about as large as puncture intervals and much coarser than on apical halves; internal sac of aedeagus without flagellum ( Fig. 74, 75 ) ......................................... S. bolmarum nov.sp.

98 Metaventrite with submesocoxal lines convex. Internal sac of aedeagus with two pairs of similar, horn-like sclerites ( Fig. 215 )......................................... S. luzonicum PIC

- Metaventrite with submesocoxal lines parallel. Internal sac of aedeagus with single pair of horn-like sclerites ( Fig. 212 ) ................................................ S. jankodadai nov.sp.

99 Metaventrite with antecoxal rows of coarse puncture....................... S. conflictum nov.sp.

- Metaventrite without antecoxal puncture rows............................................................100

100 Minute species, body length not exceeding 1.0 mm. Submetacoxal areas very narrow, smaller than submesocoxal areas. Elytra very finely punctate........................101

- Body length exceeding 1.1 mm ...................................................................................103

101 Antennomere IV about 1.3 times as long as III. Aedeagus in lateral view with straight parameres ( Fig. 56 )................................................................. S. blefusca nov.sp.

- Antennomere IV about twice as long as III. Aedeagus in lateral view with curved parameres ....................................................................................................................102

102 Aedeagus in dorsal view with parameres lobed subapically and abruptly narrowed near tip ( Figs 59, 60 ) ............................................................................... S. liliput nov.sp.

- Aedeagus in dorsal view with parameres not lobed, gradually narrowed apically ( Figs 63, 64 ) ....................................................................................... S. nanellum nov.sp.

103 Punctation on apical areas of elytra notably denser and coarser than on prevailing elytral surface. Aedeagus symmetrical, internal sac bearing robust sclerites ( Figs 65, 66 ) ......................................................................................................... S. lineare nov.sp.

- Elytral punctation not becoming coarser and denser near apices. Aedeagus symmetrical or asymmetrical, internal sac lacking robust sclerites .............................104

104 Sternite 1 with submetacoxal areas conspicuously large, as long as or exceeding interval to apical margin .......................................................................... S. sagax nov.sp.

- Sternite 1 with submetacoxal areas distinctly shorter than interval to apical margin...105

105 Apical fourth of elytra, abdomen and appendages uniformly or almost uniformly ochreous, much lighter than thorax or elytra ...................................... S. diversum nov.sp.

- Colour pattern of body different ..................................................................................106

106 Aedeagus symmetrical. Pronotum and elytra light reddish-brown or brown...............107

- Aedeagus asymmetrical. Body colour dark brown or blackish....................................109

107 Elytral punctation notably coarser than pronotal punctation. Submesocoxal lines parallel. Mesepimera about twice as long as intervals to mesocoxae. Apical process of median lobe short ( Figs 164, 165 )................................................... S. inflexum nov.sp.

- Elytral punctation hardly coarser than pronotal punctation. Submesocoxal lines convex. Mesepimera about as long as or slightly shorter than intervals to mesocoxae. Apical process of median lobe long.............................................................................108

108 Aedeagus with parameres very narrow, filiform, extended proximally to meet at proximal side of minute basal bulb.......................................................... S. sexuale LÖBL

- Aedeagus with parameres fairly wide, not filiform and not extended proximally; basal bulb conspicuously elongate........................................................ S. ilonggo nov.sp.

109 Aedeagus with parameres strongly expanded..............................................................110

- Aedeagus with parameres not expanded............................................ S. duplicatum LÖBL

110 Aedeagus with left paramere abruptly narrowed and curved in apical section .................. ............................................................................................................ S. anomalum LÖBL

- Aedeagus with left paramere broad and oblique in apical section ........ S. ramosum LÖBL

Figures

Figs 169 - 176: Aedeagi of Scaphisoma; (169), S. ochropenne nov. sp., in lateral view, scale = 0.2 mm; (170), ditto, apical process of median lobe, lateral view, scale = 0.05 mm; (171, 172), S. pseudokalabitum nov. sp., in dorsal and lateral views, scale 0.1 mm; (173), paramere in ventral view, scale = 0.1 mm; (174, 175), S. sinuatum nov. sp., in dorsal and lateral views, scale 0.1 mm; (176), S. stigmatipenne HELLER, in lateral view, scale = 0.1 mm.

Figs 162 - 168: Aedeagi of Scaphisoma; (162, 163), S. inexspectatum nov. sp., in dorsal and lateral views, scale = 0.1 mm; (164, 165), S. inflexum nov. sp., in dorsal and lateral views, scale = 0.1 mm; (166), ditto, parameres in ventral view, scale = 0.1 mm; (167), S. luteopygidiale (PIC), in lateral view, scale = 0.1 mm; (168), S. ochropenne nov. sp., in dorsal view, scale = 0.2 mm.

Figs 157 - 161: Aedeagi of Scaphisoma in dorsal and lateral views; (157 - 159), S. hamatum nov. sp., internal sac extruded, scale = 0.2 mm; (159), ditto, apical part of median lobe with internal sac, scale = 0.1 mm; (160, 161), S. banguiense LÖBL, scale = 0.1 mm.

Figs 177 - 183: Aedeagi of Scaphisoma; (177), S. surigaosum (PIC), in lateral view, scale 0.2 mm; (178, 179), S. transversale nov. sp., in dorsal and lateral views, scale = 0.1 mm; (180, 181), S. acutulum nov. sp., in dorsal and lateral views, scale = 0.2 mm; (182, 183), S. apomontanum nov. sp., in dorsal and lateral views, scale = 0.2 mm.

Figs 201 - 206: Aedeagi of Scaphisoma; (201, 202), S furcatum nov. sp., in dorsal and lateral views, scale = 0.2 mm; (203), ditto, internal sac, scale = 0.1 mm; (204, 205), S. furcigerum nov. sp., in dorsal and lateral views, scale = 0.2 mm; (206), ditto, internal sac, scale = 0.2 mm.

Figs 207 - 211: Scaphisoma; (207, 208); S. furcillatum nov. sp. aedeagus in dorsal and lateral views, scale = 0.2 mm; (209, 210), S. glabrellum nov. sp., aedeagus in dorsal and lateral views, internal sac extruded, scale = 0.2 mm; (211), ditto, apical part of male sternite 6, scale = 0, 2 mm.

Figs 247 - 250: Aedeagi of Scaphisoma; (247), S. tridens nov. sp., internal sac, scale = 0.1 mm; (248, 249), S. trifurcatum nov. sp., aedeagus in dorsal and lateral views, scale = 0.3 mm; (250), ditto, apical part of median lobe, oblique view, scale = 0.1 mm.

Figs 242 - 246: Aedeagi of Scaphisoma in dorsal and lateral views, in lateral view without right paramere; (242, 243), S. tricoloripenne nov. sp.; (244, 245); S. tridens nov. sp.; (246), ditto, right paramere in ventral view. Scale = 0.3 mm.

Figs 251 - 255: Aedeagi of Scaphisoma; (251, 252), S. trilobum nov. sp., in dorsal and lateral views, scale = 0.3 mm; (253, 254), ditto, internal sacs, in dorsal and lateral views, scale = 0.15 mm; (255), S. trimaculatum nov. sp., aedeagus in dorsal view, scale = 0.3 mm.

Figs 234 - 237: Aedeagus of Scaphisoma tricolor HELLER; (234), in dorsal view; (235), in lateral view, without proximal part of basal bulb, scale = 0.2 mm; (236, 237), internal sac in dorsal and lateral views, scale = 0.1 mm.

Figs 238 - 241: Aedeagi of Scaphisoma in dorsal and lateral views; (238, 239), S. tricoloratum nov. sp., in lateral view, without right paramere; (240, 241), S. tricolorinotum nov. sp. Scale = 0.3 mm.

Figs 1 - 6: Habitus of Scaphisoma; S. anomalum LÖBL (1); S. bibiranense LÖBL (2); S. bicolor nov. sp. (3); S. bicoloripenne nov. sp. (4); S. bicuspidatum nov. sp. (5); S. binaluanum PIC (6). Scale: 1.0 mm.

Figs 194 - 200: Aedeagi of Scaphisoma; (193), S. dispar LÖBL, in lateral view, without proximal part of basal bulb, scale = 0.05 mm; (195), ditto, internal sac in dorsal view, scale = 0.05 mm; (196), S. disparides nov. sp., in dorsal view, scale = 0.1 mm; (197), ditto, in lateral view, without proximal part of basal bulb, scale = 0, 0 5 mm; (198), ditto, internal sac in dorsal view, scale = 0.05 mm; (199, 200), S. distinctum nov. sp., in dorsal and lateral views, scale = 0.2 mm.

Figs 52 - 57: Scaphisoma; (52), antenna of S. subpunctulum nov. sp.; (53, 54), S. diversum nov. sp., aedeagus in dorsal and lateral views; (55, 56), S. blefusca nov. sp., aedeagus in dorsal and lateral views; (57), S. ilonggo nov. sp., aedeagus in dorsal view. Scale = 0.1 mm.

Figs 46 - 51: Aedeagi of Scaphisoma in dorsal and lateral views; (46, 47), S. simplexoides nov. sp.; (48, 49), S. subgracile nov. sp.; (50, 51), S. subplanatum nov. sp. Scale = 0.1 mm.

Figs 126 - 130: Aedeagi of Scaphisoma; (126), S. inconventum nov. sp., in lateral view; (127, 128), S. inopportunum nov. sp., in dorsal and lateral views; (129, 130) S. oviforme nov. sp., in dorsal and lateral views. Scale = 0.1 mm.

Figs 131 - 136: Aedeagi of Scaphisoma in dorsal and lateral views; (131, 132), S. palawanum PIC; (133, 134), S. pandanum nov. sp.; (135, 136), S. scapulare nov. sp. Scale = 0.1 mm.

Figs 14 - 21: Habitus of Scaphisoma; (14), S. furcillatum nov. sp.; (15), S. inopportunum nov. sp.; (16), S. lunabianum nov. sp.; (17), S. inexspectatum nov. sp.; (18), S. luteomaculatum PIC; (19), S. luteopygidiale (PIC); (20), S. maculosum nov. sp.; (21), S. mindanaosum PIC. Scale: 1.0 mm.

Figs 89 - 94: Aedeagi of Scaphisoma in dorsal and lateral views; (89, 90), S. lunabianum nov. sp.; (91, 92), S. maculosum nov. sp.; (93, 94), S. maramag nov. sp. Scale = 0.1 mm.

Figs 70 - 75: Aedeagi of Scaphisoma in dorsal and lateral views; (70, 71), S. apomontium nov. sp.; (72, 73) S. binaluanum PIC; (74, 75), S. bolmarum nov. sp. Scale = 0.1 mm.

Figs 101 - 107: Aedeagi of Scaphisoma in dorsal and lateral views; (101, 102), S. nabiluanum nov. sp., scale = 0.1 mm; (103, 104), S. obscurum nov. sp., scale = 0.1 mm; (105, 106), S. sagax nov. sp., scale 0.1 mm; (107), ditto, internal sac, scale = 0.05 mm.

Figs 224 - 227: Aedeagus of Scaphisoma spatuloides nov. sp.; (224, 225), in dorsal and lateral views; (226), median lobe, without expanded dorsal side of basal bulb; (227), paramere in lateral view. Scale = 0. 2 mm

Figs 219 - 223: Aedeagi of Scaphisoma in dorsal and lateral views; (219, 220), S. duplex nov. sp., scale = 0.2 mm; (221, 222), S. nishikawai nov. sp., scale = 0.2 mm; (223), ditto, apical process of median lobe in lateral view, scale = 0.1 mm.

Figs 212 - 218: Aedeagi of Scaphisoma; (212, 213); S. jankodadai nov. sp., in dorsal and lateral views, scale 0.2 mm; (214), S. luzonicum PIC, in dorsal view, scale = 0.2 mm; (215), ditto, internal sac, scale = 0.1 mm; (216, 217), S. bilobum nov. sp., in dorsal and lateral views, scale 0.2 mm; (218), ditto, internal sac, scale = 0.1 mm.

Figs 83 - 88: Aedeagi of Scaphisoma in dorsal and lateral views; (83, 84), S. densepunctatum nov. sp., scale = 0.2 mm; (85, 86), S. hexamerum nov. sp., scale 0.1 mm; (87, 88), S. kodadai nov. sp., scale = 0.1 mm.

Figs 151 - 156: Aedeagi of Scaphisoma; (151), S. iridescens nov. sp., aedeagus in dorsal view, internal sac extruded; (152), ditto, paramere in dorsal view, scale = 0.2 mm; (153, 154), S. werneri nov. sp., aedeagus in dorsal and lateral views, scale = 0.2 mm; (155, 156), S. dissymmetricum nov. sp., aedeagus in dorsal and lateral views, scale = 0.1 mm.

Figs 95 - 100: Aedeagi of Scaphisoma in dorsal and lateral views; (95, 96), S. minutipenis nov. sp.; (97, 98), S. mirum nov. sp.; (99, 100), S. montivagum nov. sp. Scale = 0.1 mm.

Figs 65 - 69: Aedeagi of Scaphisoma; (65 - 66), S. lineare nov. sp., in dorsal and lateral views; (67), ditto, paramere in ventral view; (68), internal sac and apex of median lobe in lateral view; (69), S. caudatoides nov. sp., aedeagus in dorsal view. Scale = 0.1 mm.

Figs 58 - 64: Scaphisoma; (58), S. ilonggo nov. sp., aedeagus in lateral view, internal sac extruded; (59, 60), S. liliput nov. sp., aedeagus in lateral and dorsal views, internal sac extruded; (61), ditto, parameres in ventral view; (62), ditto, paramere in lateral view; (63, 64), S. nanellum nov. sp., aedeagus in dorsal and lateral views. Scale = 0.1 mm.