Mycetophagidae

Lawrence, John F., Escalona, Hermes E., Leschen, Richard A. B. & Ślipiński, Adam, 2014, Review of the genera of Mycetophagidae (Coleoptera: Tenebrionoidea) with descriptions of new genera and a world generic key, Zootaxa 3826 (1), pp. 195-229 : 209-211

Key to the world genera of Mycetophagidae

This key is based primarily on dissected specimens but characteristics of the genera Triphyllopsis Nikitsky (1989, 1993 ) and Pseudotriphyllopsis Nikitsky (1996) were extracted from the literature.

1. Head ( Figs 1C, 1K ) only slightly narrower than greatest pronotal width, more or less abruptly constricted behind eyes to form broad neck and short to moderately long temples ( Figs 7A–B , 8A–B ); pronotum more or less quadrate with very weakly defined lateral carinae, distinctly narrower than combined elytral bases ( Figs 7D , 8D ); male with pair of slender, oblique setose impressions between procoxae and anterior pronotal angles ( Figs 7E , 8E ); antennal club 2-segmented ( Figs 1K , 7C ) OR tarsi 3-3-3 ( Fig. 8C ); length less than 2 mm ......................................................... ... Bergininae ...2

- Head ( Figs 1A–B, 1D–J , 5 I –J, 5M–O) distinctly narrower than greatest pronotal width, not abruptly constricted behind eyes; pronotum ( Figs 1A–B, 1D–J , 5 M–N) with more distinctly rounded sides and distinct lateral carinae, not or only slightly narrower than combined elytral bases; male without paired ventral prothoracic impressions; antennal club with at least 3 segments ( Figs 1A–B, 1E–F ); tarsi 4-4- 4 in female, 3-4- 4 in male....................................................... 3

2(1) Temples shorter than eye or neck ( Fig. 7B ); antennal club 2-segmented ( Figs 1K , 7C ); tarsi 4-4- 4 in female, 3-4- 4 in male; dorsal surfaces clothed with distinct, thickened setae, usually more or less decumbent ( Figs 1K , 7G ); length more than 1.5 mm; North and Central America, West Indies, southern Europe, Caucasus, India, Sri Lanka, East Indies....... Berginus Erichson

- Temples much longer than eye and about as long as neck ( Figs 1C , 8B ); antennal club 3-segmented ( Fig. 8H ); tarsi 3-3- 3 in both sexes; dorsal surfaces clothed with minute, fine setae barely visible at lower magnifications ( Fig. 8D ); lateral edges of each elytron with row of deep pores ( Fig. 8G ); length less than 1.3 mm; Seychelles, Christmas I., Australia Nesolathrus Scott

3(1). Metacoxal length (parallel to long axis of body) more than half its width (perpendicular to long axis of body) ( Fig. 6C ), well-

separated from elytral epipleura; metanepisternum relatively short and broad, its length less than 2.5 times its width; metaventrite distinctly shorter than abdominal ventrite 1; sides of pronotum usually distinctly serrate ( Fig. 6D ); scutellar shield strongly transverse ( Fig. 6E ) or absent; elytral punctures large, nude and distinctly seriate ( Fig. 6G ); pterothorax and abdomen sometimes with large setose cavities ( Figs 6C, 6F ); hind wings usually absent; southern Europe, North Africa Esarcinae ................................................................................................. Esarcus Reiche - Metacoxal length (parallel to long axis of body) much less than half its width (perpendicular to long axis of body) ( Figs 9G , 11E , 14H ); large, nude, seriate elytral punctures and setose pterothoracic and abdominal cavities always absent................................................................................................. Mycetophaginae ...4

4(3). Mesepimera not reaching mesocoxal cavities, which are closed laterally by the meeting of mesoventrite and metaventrite ( Figs 13D, 13G , 15A, 15C ).................................................................................. 5

- Mesepimera reaching mesocoxal cavities ( Figs 9F–G , 11E , 12D , 14H )............................................ 8

5(4). Base of pronotum distinctly narrower than combined elytral bases ( Fig. 5 O); antennomeres 9 and 10 about 1.5 times as long as wide, antennomere 11 shorter and narrower than 9 or 10 ( Figs 2 H, 5O); mesoventrite carinate for most of its length ( Fig. 3 C); vestiture of moderately long erect and inclined setae ( Fig. 5 O); New Zealand...................... Zeclaviger gen. nov.

- Base of pronotum not or barely narrower than combined elytral bases; antennomeres 9 and 10 transverse or only slightly longer than wide; antennomere 11 at least slightly longer than 9 or10; mesoventrite not carinate, sometimes with short anterior linear prosternal rest; vestiture without long, erect setae......................................................... 6

6(5). Elytral vestiture dual, consisting of longer and shorter, inclined or decumbent setae, the longer setae forming about 12 longitudinal rows separated by at least two rows of shorter setae ( Figs 1H , 3A ); apical maxillary palpomere more or less parallel-sided and not wider than subapical one ( Fig. 15B ); shortest distance between mesocoxal and metacoxal cavities distinctly greater than length of abdominal ventrite 1 at same point; hind wings well-developed; cosmopolitan ( Fig. 1H ).... Typhaea Stephens

- Elytral vestiture single, consisting of inclined or decumbent setae of more or less equal length ( Fig. 2A ); apical maxillary palpomere distinctly wider than subapical one ( Figs 13A–B ); shortest distance between mesocoxal and metacoxal cavities subequal to or less than length of abdominal ventrite 1 at the same point ( Fig. 13D ); hind wings more or less reduced (apical field shortened or absent, medial field reduced, radial cell sometimes absent).......................................... 7

7(6). Pronotum widest near middle; lateral pronotal carinae distinctly serrate; posterior pronotal angles acute; lateral edges of elytra visible from above for most of their lengths; elytral apex obliquely truncate; protarsomere 1 in male distinctly expanded; abdominal ventrite 1 in male with two large patches of setigerous spicules accompanied by glandular openings ( Figs 4C, 4 F); southern Europe, North Africa.......................................................... Typhaeola Ganglbauer

- Pronotum widest well behind middle ( Fig. 2A ); lateral pronotal carinae simple; posterior pronotal angles not acute; lateral edges of elytra not visible from above; elytral apex broadly rounded; protarsomere 1 in male not expanded; abdominal ventrite 1 in male with two slender transverse pore fields ( Figs 4A, 4 D); southern Africa................. Afrotyphaeola gen. nov.

8(4). Antennae gradually expanded apically or with weak club composed of 4 or more antennomeres ( Figs 1D–F , 5 J, 9D); elytral punctation seriate, but puncture rows sometimes not clearly visible due to dense interstrial punctation and recumbent setae ( Figs 1D–F ).......................................................................................... 9

- Antennae with distinct 3-segmented club ( Figs 1A–B, 1I –J , 2 F–G, 5I, 5M–N, 14B); punctation seriate or not............ 11

9(8). Frontoclypeal suture somewhat angulate and not deeply impressed; paired basal pronotal impressions absent; posterior pronotal angles more or less embracing elytral bases ( Fig. 5 J); shortest distance between mesocoxal and metacoxal cavities distinctly less than length of abdominal ventrite 1 at same point; tibial spurs biserrate; hind wings reduced and strap-like; Juan Fernandez Is......................................................................... Filicivora Leschen & Lawrence

- Frontoclypeal suture deeply impressed and straight or curved ( Figs 2 E, 9A); paired basal pronotal impressions present ( Figs 1E–F ); shortest distance between mesocoxal and metacoxal cavities distinctly greater than length of abdominal ventrite 1 at same point ( Fig. 9G ); tibial spurs simple ( Fig. 9C ); hind wings well-developed................................... 10

10(9). Basal pronotal impressions very small and sometimes indistinct; elytron with sutural stria; antennal club indistinct; eyes rounded; elytra usually unicolorous red or brown ( Fig. 1D ); eastern Palaearctic, Caucasus, North Africa, Japan............................................................................................... Eulagius Motschulsky

- Basal pronotal impressions large and distinct ( Figs 1E–F ); elytra without sutural stria; antennal club usually distinctly 4- to 6- segmented ( Fig. 9D ); eyes elongate-oval, usually slightly emarginate ( Fig. 9A ); elytra usually with dark and light pattern, occasionally black; widely distributed except Australia and New Zealand....................... Mycetophagus Hellwig

11(8). Pronotum widest at base, which is as wide as or wider than combined elytral bases ( Figs 1I –J ); posterior pronotal angles more or less embracing elytral humeri ( Figs 11C–D ); pronotal hypomera ( Figs 12F, 12H ) and elytral epipleura ( Fig. 12I ) usually strongly oblique or almost vertical, not visible in lateral view; pronotal disc with pair of elongate, slit-like, basal impressions, usually darkly pigmented ( Fig. 2 C); metaventrite not shorter than first abdominal ventrite ( Fig. 12D ); frontoclypeal suture not deeply impressed, forming a fine curved or angulate line ( Fig. 11A ); scutellar shield widest at base and broadly rounded at apex ( Fig. 12K ); antennal club usually slender, often with antennomeres 9–11 at least slightly elongate ( Fig. 11I ); tibial spurs usually bipectinate and sometimes highly complex ( Figs 11B, 11G –HN, 12B); body more or less flattened, often with complex, dual vestiture ( Figs 11A, 11I , 12A ).................................................................. 12

- Pronotum usually widest anterior to base ( Figs 1G–H, 1L , 5 I, 5N); IF pronotum widest at base, THEN posterior pronotal angles not embracing elytral humeri ( Figs 1A , 5 M) OR metaventrite distinctly shorter than first abdominal ventrite; base of pronotal disc without or with subcircular paired impressions; frontoclypeal suture usually more deeply impressed; elytral epipleura more or less flattened, often partly visible in lateral view; scutellar shield usually widest behind base and apically truncate ( Figs 10G , 14D ), body usually more convex with simple vestiture........................................... 13

12(11). Outer edge of protibia lined with closely-spaced, spine-like setae; protibial apex with one long bipectinate spur and another much shorter spur ( Figs 2 I, 12B); apical maxillary palpomere slightly longer than preapical one and strongly obliquely truncate

so that outer edge is acute ( Fig. 12C ); apical labial palpomere as wide as or slightly wider than preapical one and attached at its apex ( Fig. 12C ); elytra with distinct rows of punctures ( Figs 1I , 12J ); Madeira, Japan, Australia.......... Litargops Reitter - Outer edge of protibia lined with finer, less closely spaced setae ( Fig. 11G ); protibial apex with two bipectinate spurs ( Figs 11G–H ); apical maxillary palpomere distinctly longer than preapical one and not strongly, obliquely truncate ( Fig. 2 K); apical labial palpomere distinctly narrower than preapical one and attached subapically ( Fig. 2 J); elytra punctures not forming distinct rows ( Fig. 1J ); widely distributed........................................................ Litargus Erichson

13(11). Body about 2.5 times as long as wide ( Fig. 1G ); lateral pronotal carinae dentate ( Figs 10B, 10E ); sides of elytra parallel for most of their lengths; elytral punctation distinctly seriate ( Fig. 10F ); abdominal ventrites 2–4 in male each with a pair of small, circular setose patches near midline ( Fig. 10H ); Chile and Argentina............................ Pseudochrodes Reitter

- Body shorter and broader ( Figs 1A–B, 1L , 5 I, 5M–N); lateral pronotal carinae simple or minutely denticulate ( Figs 3 D, 14E); sides of elytra more or less curved for most of their lengths; elytral punctation seriate or not; abdominal ventrites in male without paired, subcircular setose patches..................................................................... 14

14(13). Elytra with distinct puncture rows....................................................................... 15

- Elytra without distinct puncture rows (occasionally with indistinct or incomplete rows)............................. 16

15(14). Shortest distance between mesocoxal and metacoxal cavities distinctly less than length of abdominal ventrite 1 at same point; sides of elytra distinctly converging beyond middle; elytra with short scutellary striole; pronotum at least 0.65 times as long as wide; hind wings absent; abdominal ventrites in male without pore fields; central Asia.............. Triphyllopsis Nikitsky

- Shortest distance between mesocoxal and metacoxal cavities distinctly greater than length of abdominal ventrite 1 at same point; sides of elytra more or less parallel, converging only at posterior fourth; elytra without scutellary striole; pronotum less than half as long as wide; hind wings well-developed; abdominal ventrites 1 to 3 in male each with a pair of transversely oriented V- or U-shaped pore fields (as in Fig. 4B ); Japan...................................... Triphyllioides Miyatake

16(14). Posterior pronotal angles at least slightly acute and sometimes embracing elytral bases; basal pronotal foveae absent; shortest distance between mesocoxal and metacoxal cavities distinctly less than length of abdominal ventrite 1; scutellar shield strongly transverse; hind wings absent........................................................................... 17

- Posterior pronotal angles right or obtuse, not embracing elytral bases; basal pronotal foveae usually present; shortest distance between mesocoxal and metacoxal cavities distinctly greater than length of abdominal ventrite 1; scutellar shield slightly transverse; hind wings present.............................................................................. 18

17(16). Mesoventrite carinate; posterior pronotal angles not or only slightly embracing elytral bases ( Fig. 1A ); prosternal process extending well beyond procoxae, narrowly rounded at apex; prosternum in male with pubescent fovea (as in Fig. 14E ); abdominal ventrite 1 in male with pair of U-shaped pore fields (as in Figs 4B, 4 E); Caucasus................. Triphyllina Reitter

- Mesoventrite not carinae, with small subtriangular prosternal rest; posterior pronotal angles distinctly embracing elytra; prosternal process not extending beyond procoxae, truncate at apex; prosternum and abdominal ventrite 1 in male simple; China................................................................................ Pseudotriphyllopsis Nikitsky

18(16). Antennomeres 9 and 10 slightly longer than wide and slightly expanded apically with sensory field at apex, 11 distinctly longer than wide and distinctly longer than 9 or 10 ( Fig. 2 F); mesoventrite more or less concave, with small anterior subquadrate prosternal rest; vertex in male with pubescent fovea ( Figs 2 E, 5M); prosternum and abdomen in male simple; southern North America and Central America.......................................................... Neotriphyllus gen. nov.

- Antennomeres 9 and 10 transverse ( Fig. 2 G); mesoventrite with linear or triangular prosternal rest ( Figs 14H , 15 E); vertex in male without pubescent fovea; abdomen in male with pore fields ( Figs 4B, 4 E).................................... 19

19(18). Posterior pronotal angles broadly rounded ( Figs 2B , 15 D); mesoventrite with large triangular prosternal rest occupying most of ventrite ( Fig. 15 E); abdominal ventrite 1 in male with very slender, curved linear pore fields extending from anterolateral angles to mesal fourth ( Fig. 15 G); length less than 1.3 mm; North and Central America................. Thrimolus Casey

- Posterior pronotal angles not broadly rounded ( Figs 3 D, 14E); mesoventrite with linear or small, triangular prosternal rest ( Fig. 14H ); abdominal ventrite 1 in male with shorter and broader linear or C-shaped pore fields ( Figs 4A–B ); length greater than 1.3 mm; Holarctic and Notogean........................................................................ 20

20(19). Antennomeres 9 and 10 slightly transverse, slightly expanded apically, with sensory fields at apex; anterior edge of mesoventrite with 2 small carinae; prosternum in male without setose fovea; abdominal ventrites 1–4 in male each with pair of Vshaped or linear pore fields; habitus ( Fig. 1L ); Eurasia........................................... Triphyllus Dejean

- Antennomeres 9 and 10 distinctly transverse and widest at middle, with sensory fields often occupying half of antennomere ( Figs 2 G, 14B); anterior edge of mesoventrite with short linear or narrowly triangular elevation ( Fig. 14H ); prosternum in male with setose fovea ( Fig. 14E ); abdomen with U-shaped or linear pore fields on ventrite 1 only ( Figs 4B, 4 E)............. 21

21(20). Antennomere 11 distinctly narrower than 9 or 10 ( Fig. 2 G); prosternal process parallel-sided, slightly emarginate at apex; procoxal cavity without slit-like extension; Eurasia and eastern North America.................... Pseudotriphyllus Reitter

- Antennomeres 9–11 of equal width; prosternal process slightly expanded almost to apex, then narrowed and apically truncate ( Fig. 14E ); procoxal cavity with short, curved, slit-like extension ( Fig. 14E ); New Zealand, Madagascar and Chile................................................................................. Nototriphyllus gen. nov.

Figures

FIGURES 1 A – L, habitus, dorsal: A, Triphyllina lederi Reitter, length 2.0 mm, © K. V. Makarov; B, Pseudotriphyllus colchicus (Reitter), length 2.3 mm, © K. V. Makarov; C, Nesolathrus sp. (Australia: QLD: McIlwraith Range), length 1.2 mm, © CSIRO Australia; D, Eulagius ussuriensis Nikitsky, length 4 mm, © K. V. Makarov; E, Mycetophagus decempunctatus Fabricius, length 4.2 mm, © K. V. Makarov; F, Mycetophagus irroratus Reitter, length 3.8 mm, © K. V. Makarov; G, Pseudochrodes suturalis Reitter, length 2.5 mm, © CSIRO Australia; H, Typhaea stercorea (Linnaeus), length 3.1 mm, © CSIRO Australia; I, Litargops maculosus Reitter, length 3.4 mm, © K. V. Makarov; J, Litargus connexus Geoffroy, length 3.5 mm, © K. V. Makarov; K, Berginus sp. (Panama: Paraiso), length 1.4 mm, © CSIRO Australia; L, Triphyllus bicolor Fabricius, length 3.9 mm, © K. V. Makarov.

FIGURES 7 A – H, Berginus sp. (Panama): A, head, dorsal; B, head, ventral; C, antenna; D, prothorax, dorsal; E, prothorax, ventral; F, mesotibia and mesotarsus; G, elytron; H, portion of mesothorax and metathorax, showing lateral closure of mesocoxal cavity. All figures © CSIRO Australia.

FIGURES 8 A – H, Nesolathrus sp. (McIlwraith Range, Queensland, Australia): A, head, dorsal; B, head, ventral; C, mesotibia and mesotarsus; D, prothorax, dorsal; E, prothorax, ventral; F, pterothorax, ventral with mesocoxae removed; G, left elytron, lateral; H, antenna. All figures © CSIRO Australia.

FIGURES 5 A – H, mycetophagid male genitalia: A – B, Nototriphyllus araucania sp. n.: A, basale (phallobase) and apicale (partly fused parameres); B, penis; C – E, Neotriphyllus confusus (Horn): C, basale and apicale; D, penis; E, segments IX and X with long spiculum gastrale; F – H, Zeclaviger explanatus sp. n.: F, basale and apicale; G, penis; H, segments IX and X with short spiculum gastrale. FIGURES 5 I – J, habitus, dorsal: I, Nototriphyllus hispidellus (Broun), length 1.9 mm, © Landcare New Zealand; J, Filicivora chilensis (Philippi & Philippi), length 5.3 mm, © CSIRO Australia. FIGURES 5 K – L, Afrotyphaeola natalensis sp. n.: K, basale and apicale; L, penis. FIGURES 5 M – O, habitus: M, Neotriphyllus confusus (Horn), length 3.4 mm, © Landcare New Zealand; N, Nototriphyllus araucania sp. n., length 2.8 mm, © CSIRO Australia; O, Zeclaviger explanatus sp. n., length 1.6 mm, © Landcare New Zealand.

FIGURES 6 A – G, Esarcus leprieuri Reiche: A, head, ventral; B, anterior portion of head; C, pterothorax with right mesocoxa and left metacoxa removed; D, prothorax, ventral, with right coxa removed; E, scutellar shield; F, abdominal ventrites; G, right elytron. All figures © CSIRO Australia.

FIGURES 9 A – G, Mycetophagus punctatus Say: A, head, dorsal; B, prothorax, ventral; C, mesotibial apex; D, antenna; E, prothorax, dorsal; F, portion of meso- and metathorax, showing laterally open mesocoxal cavity; G, pterothorax, ventral. All figures © CSIRO Australia.

FIGURES 11 A – I, Litargus irregularis (Sharp): A, head, dorsal; B, mesotibia and mesotarsus; C, pronotum, dorsal; D, prothrax, ventral; E, pterothorax, ventral, with right meso- and metacoxae removed; F, antenna; G, protibia and protarsus; H, apex of protibia; I, area of dorsal head surface. All figures © CSIRO Australia.

FIGURES 14 A – H, Nototriphyllus araucania sp. n.: A, head, dorsal; B, antenna; C, head, ventral; D, scutellar shield; E, male prothorax, ventral with left coxa removed, showing prosternal sex patch; F, right elytron, dorsal; G, mesotibial apex and mesotarsus; H, pterothorax, ventral. All figures © CSIRO Australia.

FIGURES 13 A – L, Afrotyphaeola natalensis sp. n., ♀: A, head, dorsal; B, head, ventral; C, prothorax, dorsal; D, pterothorax, ventral with right mesocoxa removed; E, antenna; F, prothorax, ventral; G, left mesocoxa, showing lateral closure of cavity by mesoventrite and metaventrite; H, abdomen, ventral; I, right procoxal cavity; J, mesotibial apex and mesotarsus; K, protibial apex and protarsus, L, scutellar shield. All figures © CSIRO Australia.

FIGURES 15 A – C, Typhaea stercorea (Linnaeus): A, pterothorax, ventral; B, head, dorsal; C, mesocoxa, showing lateral closure of cavity by meso- and metaventrites. Figures 15 D – G, Thrimolus minutus Casey: D, prothorax, ventral, with right coxa removed; E, pterothorax and base of abdomen, ventral; F, head, dorsal; G, left metacoxa and left portion of abdominal ventrite 1, with pore field vaguely indicated. All figures © CSIRO Australia.

FIGURES 12 A – L, Litargops sp. (Brindabella Range, ACT, Australia): A, head, dorsal; B, apex of protibia; C, head, ventral; D, pterothorax, ventral with right mesocoxa removed; E, apex of mesotibia; F, prothorax, ventral, with right coxa removed; G, mesotibial apex and mesotarsus; H, prothorax, anterior; I, left elytron, ventral; J, right elytron, dorsal; K, scutellar shield; L, antenna. All figures © CSIRO Australia.

FIGURES 2 A – B, habitus, dorsal: A, Afrotyphaeola natalensis sp. n., length 1.6 mm; B, Thrimolus minutus Casey, length 1.2 mm. FIGURES 2 C – D, Litargus balteatus LeConte, prothoraces: C, dorsal, showing paired slit-like impressions; D, posterior, showing notal attachments of posterior apodemes and cryptopleura. Figure 2 E, Neotriphyllus confusus (Horn), head, dorsal. FIGURES 2 F – H, antennal apex: F, Neotriphyllus confusus (Horn); G, Pseudotriphyllus suturalis Reitter; H, Zeclaviger explanatus sp. n. Figure 2 I, Litargops sp. (Australia: ACT: Picadilly Circus, Brindabella Range), protibial apex. FIGURES 2 J – K, Litargus balteatus LeConte: J, labium; K, maxilla. All figures © CSIRO Australia.

FIGURES 3 A – B, portion of elytral surface: A, Typhaea stercorea (Linnaeus); B, Nototriphyllus araucania sp. n. Figure 3 C, Zeclaviger explanatus sp. n., pterothorax, ventral, with right metanepisternum and right metacoxa and hind leg removed. Figure 3 D, Nototriphyllus araucania sp. n., prothorax, dorsal, with both coxae removed. Figure 3 E, Zeclaviger explanatus sp. n., left halves of abdominal ventrites 1 and 2, showing pore field in anterolateral corner of ventrite 1. All figures © CSIRO Australia.

FIGURES 4 A – C, right half of abdominal ventrite 1: A, Afrotyphaeola natalensis sp. n. with linear pore field; B, Nototriphyllus araucania sp. n., with C-shaped pore field; C, Typhaeola maculata Perris, with pores at setal bases. FIGURES 4 D – F, all five abdominal ventrites of same three species. Figure 4 G, Nototriphyllus araucania sp. n., hind wing. All figures © CSIRO Australia.

FIGURES 10 A – H, Pseudochrodes suturalis Reitter: A, head, dorsal; B, prothorax, dorsal; C, mesotibial apex; D, antenna; E, prothorax, ventral; F, left elytron, dorsal; G, scutellar shield; H, male abdomen showing setose sex patches. All figures © CSIRO Australia.