Appendix B. List of Characters
Morphological characters and character states used in my phylogenetic analyses: Each taxonomic category mentioned here is treated in the newly revised sense following my taxonomic changes.
1. Head, shape (dorsal and ventral views): (0) as long as wide to transverse, oval to hexagonal, not rectangular (Figures 16A and 50B); (1) elongate, slightly to moderately longer than wide (Figure 64B); (2) markedly elongate, much longer than wide (Figures 64D and 67A); (3) as long as wide, rectangular.
Head shapes of Tachyporinae are usually slightly to moderately transverse, whereas those of Mycetoporinae are usually elongate (extreme cases seen in Bobitobus; Figure 64D), with only a few exceptions.
2. Head, ocelli (dorsal view): (0) absent; (1) present.
3. Head, uniform punctation on vertex (modified from Herman [
25]: character 0) (dorsal view): (0) absent (Figures 7C, 23B, 51D and 64D); (1) fine; (2) medium (Figure 44A); (3) large (Figure 58A).
The dorsal surface of the head of Tachyporinae generally lacks punctation, however, that of Deropini is covered with moderate punctation, and of some Tachinusini (members of the former Megarthropsini) with much larger and deeper punctation.
4. Head, microsetae [dorsal view]: (0) absent (Figures 7C, 50A, 51D, and 64D); (1) present, uniformly and sparsely covered with thin, minute, and inconspicuous setae; (2) present, uniformly (at least apical half of head dorsum) and densely covered with thin, rather conspicuous setae (Figures 15B and 44A); (3) present, only partially covered; (4) present, uniformly covered with extremely short, thick, but vestigial setae (nearly absent); (5) present, uniformly covered with extremely short, thick, distinctly modified setae (Figure 58C; [
25]: Figure 185).
The surface of head is glossy and glabrous in Mycetoporinae but in Tachyporinae have a series of variations as listed above. The dorsal surface of the head is glabrous in Tachyporina, Vatesini, and most taxa of Tachinusini. It is, however, densely pubescent in Deropini and Euconosomatina.
5. Head, surface [dorsal view]: (0) even (Figures 7C and 50A); (1) weakly rough, with indentations; (2) moderately rough or tuberculate (Figure 58B); (3) dorsum with numerous, long, distinct, longitudinal carinules; (4) dorsum with deep, oblique grooves.
In Tachyporinae, the surface of head is generally even, but that of some Tachinusini (members of the former Megarthropsini) is rough (Figure 58B; [
25]: Figure 184).
6. Head, ocular seta near posterior of each eye (sensu Campbell [
42]) (dorsal view): (0) absent (Figures 16A, 51D and 64D); (1) present, thin, short (Figures 50 and 67B); (2) present, much thicker and longer, conspicuous (Figure 64A–C).
In some outgroups and Mycetoporinae, there is a pair of ocular setae near the posterior margins of the eyes. However,
Bobitobus (
B. lunulatus) lacks such setae (Figure 64D). I found a pair of short ocular setae in
Bryophacis smetanai (Figure 67B: 6-1), although it has been considered as absent [
33]. Within Tachyporinae, some members of Tachinusini (members of the former Megarthropsini) also have such setae, but they are much shorter and less conspicuous than those of typical mycetoporines.
7. Clypeal anterior margin (modified from Herman [
25]: character 5) (dorsal view): (0) various, not reflexed upward (Figure 50A); (1) only weakly reflexed upward; (2) strongly reflexed upward (Figure 58B,C).
In the limited taxa of Tachinusini, the anterior margin of the clypeus is reflexed above (see Herman [
25]).
8. Frontal shelf (shelf-like projection at each side of frons) [dorsal and lateral views]: (0) present, covering antennal insertion (Figures 44A and 50B); (1) absent (Figure 64B,D).
There are more or less developed frontal shelves in Tachyporinae, but they are absent in Mycetoporinae.
9. Antennal insertions (dorsal view): (0) partially exposed, more or less visible from above, but at least concealed partially under frontal shelf or part of frons (Figures 41J, 44A and 50B); (1) nearly fully exposed, but slightly concealed under very weakly developed projection of frons; (2) fully exposed, completely visible from above, not concealed under frontal shelf or part of frons (Figure 64B,D); (3) not visible from above, completely concealed under strongly developed frontal shelf.
The antennal insertions are more or less (= moderately) visible in Tachyporinae, whereas they are fully exposed in Mycetoporinae (spherical insertion fully or almost fully visible).
10. Antennal insertion, location (modified from Grebennikov & Newton [
2]: character 94) (dorsal view): (0) antennae inserted at, or anterior to, anterior margin of eyes (Figures 44A, 50B and 64D); (1) antennae inserted posterior to anterior margin of eyes ([
189]: Figure 1.13.1); (2) antennae vertically inserted posterior to anterior margin of eyes (Figure 26G).
In Tachyporinae, antennae are generally inserted at, or anterior to, the anterior margin of the eyes. However, the head capsule of two tachyporine genera, namely Vatesus (Vatesini) and Lamprinus (Tachyporina), are deformed in associations with their myrmecophilous lifestyles. This results in peculiar antennal insertions which are located posterior to the anterior margin of the modified, vertical eyes.
11. Midcranial suture (sensu Naomi [
200]: Figure 1B) (dorsal view): (0) absent (Figures 15A, 23A, 44A, 58C and 64A); (1) present (Figures 7A, 23B, and 50).
There is a well-developed midcranial suture on the frons in most taxa of Tachyporinae (Figure 7A), although some tachyporine genera, mostly specialized taxa such as myrmecophiles, lack such sutures. In one myrmecophilous species, Lamprinodes saginatus (Tachyporina), it is very short and indistinct, but it is considered here as present.
12. Postocular area, neck or narrowing (dorsal and ventral views): (0) absent (Figures 9A, 16A, 23A, 51A, 64B, and 67C); (1) present: head capsule very slightly narrowed at middle of postocular area (Figure 26A); (2) present, narrowed only weakly just behind eyes (Figures 8A,B, 64A and 67A); (3) present, narrowed moderately just behind eyes at sides only (Figures 50, 51B,D, 53A and 58A); (4) present, narrowed distinctly at middle of postocular area, all around (Figures 44A and 45A); (5) present, neck strongly constricted just behind eyes, distinct all around; (6) present, neck strongly constricted at middle of postocular area, distinct all around.
The conditions of the postocular areas vary greatly among the selected taxa used here. In some outgroups, there is strong neck constriction behind the eyes, e.g., Neophoninae, Pseudopsinae, and Trichophyinae. In Mycetoporinae, such constriction in the neck region is absent or alternatively weakly narrowed, starting immediately behind the eyes. Several different conditions of this character state were found in Tachyporinae. For example, the head lacks narrowing or constriction in most taxa of Tachyporina, some Vatesini, and Nitidotachinus (Tachinusini). Within Vatesini, only Termitoplus and Coproporus have a weak narrowing in the middle of the postocular area. In general, Tachinusini has a moderately narrowed neck region, starting from just behind the eyes. A similar condition was found in Deropini, but they are narrowed from the middle of the postocular area to the base of the head capsule.
13. Head capsule, with a pair of longitudinal ridges basolaterally along underside of eyes (ventral view): (0) absent (Figure 26A); (1) present, incomplete (Figures 53A and 58D); (2) present, complete (Figure 67A,C).
The complete and distinct infraorbital ridge can be found in some outgroups (i.e., Olisthaerinae, Staphylininae, and Aleocharinae) and Mycetoporinae. There is no such ridge found in Tachyporinae, although Leucotachinus and Nepaliodes in Tachinusini possess a pair of incomplete, somewhat weak wavy ridges.
14. Hypostomal sutures, including traces (sensu Naomi [
200]: Figure 1A) (ventral view): (0) curved or angulate (Figures 10E, 16B, 45A, and 53A); (1) straight (Figure 67C); (2) fused for part of their length, without any trace at least partially; (3) absent.
In some outgroups and Mycetoporinae, the hypostomal sutures are generally not curved, each forming a nearly straight line. In contrast, they are curved, or forming a distinct angle, in Tachyporinae. This condition is possibly a universal condition in the subfamily, and the hypostomal sutures in Tachyporini form a pair of strong angles with the submentum.
15. Gular sutures (sensu Naomi [
200]: Figure 1A) (ventral view) (Grebennikov & Newton [
2]: character 98): (0) well separated throughout (Figures 10E and 67C); (1) fused for part of their length.
16. Gular sutures (ventral view): (0) not markedly curved or divergent posteriorly (Figures 10E, 45A and 67C); (1) markedly strongly curved or divergent posteriorly (Figure 16B); (2) fused for part of their length.
In Euconosoma, Sepedophilus (both Euconosomatina), and Vatesus (Vatesini), the gular sutures are markedly divergent basally.
17. Postgena (ventral view): (0) without minute setae (Figure 67A); (1) with minute setae (Figures 24B and 45A); (2) with strong setae (Figure 58D).
Some members of Mycetoporinae, Vatesini, Tachinusini, and Deropini have scattered minute setae on the postgena, but they are absent in most other tachyporines.
18. Dorsal tentorial arms (sensu Naomi [
200]: Figure 1B; Naomi [
201]: Figure 2A) (dorsal view): (0) present, usually forming a pair of tentorial pits (Figures 9A and 16A); (1) absent or strongly reduced (Figure 67B).
As a general trend, the dorsal tentorial arms are strongly reduced in most members of Mycetoporinae, but I found dorsally pointed arms in a few mycetoporine genera shown in the phylogeny, namely Parabolitobius and Bolitobius. On the other hand, they are well-developed in Tachyporinae, and even reach the inner surface of the dorsum, resulting in a pair of tentorial pits in most cases.
19. Antenna, structure (dorsal and ventral views): (0) normal, moniliform to filiform (Figures 9D, 64B and 68F,G); (1) weakly modified in association with social insects (i.e., flagellomeres weakly flattened and contiguous, concealing each stem pedicels; Figure 9C); (2) strongly modified in association with social insects (i.e., flagellomeres strongly flattened and adjacent ones contiguous, concealing pedicels) (Figure 26G); (3) weakly clavate or clubbed apically (Figure 23B); (4) extremely slender, verticillate.
In Tachyporinae, the general shape of the antennae is filiform to fili-moniliform Although this condition may be considered as “incrassate” (Lawrence et al. [
202]: character 38-2; Lawrence & Ślipiński [
203]: 35), I maintain the usage both filiform and moniliform here.
Mimocyptus and
Termitoplus (both Vatesini) have weakly clubbed antennae, unusual within the subfamily.
Lamprinodes (Tachyporina) has slightly flattened, modified antennae in association with myrmecophilous lifestyles [
15]. The extreme cases, markedly telescoped, modified, myrmecophilous antennae [
15], are found in
Lamprinus (Tachyporina) and
Vatesus (Vatesini). The presence of very slender, verticillate antennae characterizes the subfamily Trichophyinae and the genus
Habrocerus (Habrocerinae) Habrocerinae.
20. Antenna, length compared to head width width (dorsal and ventral views): (0) not distinctly short, clearly longer than head width (Figures 15A and 64B); (1) distinctly short, as long as, or only slightly longer than, head width (Figure 23B).
Mycetoporinae can be characterized by the presence of long and slender antennae, but those of the tachyporine taxa vary greatly from very short (some Vatesini) to markedly long (Deropini and some Tachinusini). A notably longer condition of the antennae can be seen in semi-aquatic taxa, whereas very short antennae are generally found in myrmecophilous or termitophilous taxa, seeming to have a protective function. In Vatesini, the presence of shorter antennae, as long as or shorter than pronotal width, is a general common state in the tribe, and is shared among the crown group.
21. Antennal micropubescence distribution (dorsal and ventral views): (0) without clear borderline, density increasing apically (Figures 9C,D, 17A, 45C and 68F,G); (1) with clear borderline, lacking dense and fine recumbent setae on basal three antennomeres (a1–a3) (Figure 26D); (2) with clear borderline, lacking dense and fine recumbent setae on a1–a4; (3) with clear borderline, lacking dense and fine recumbent setae on a1–a2 (Figure 59C); (4) with clear borderline, lacking dense and fine recumbent setae on a1–a5; (5) with clear borderline, lacking dense and fine recumbent setae on a1–a8; (6) without clear borderline, almost same density from base to apex.
The distributional patterns of the antennal micropubescence have been used in generic classification and identification of Tachyporinae (e.g., [
91]). Although there is no setal borderline of the antennae observed in most Mycetoporinae, a rather clear setal borderline was found in
Neobolitobius, and was considered as ‘2′. In Tachyporinae, this character is highly variable among the tribes, genera, and species. Within Tachyporini, the antennae lack such borderlines, with gradually increasing density of microsetae towards the apex. In contrast, Vatesini has three different types of antennal micropubescence patterns (21: 0-2), antennae without dense and fine recumbent setae on the basal three antennomeres is the most common state in this tribe (21-1). Within Tachinusini, the antennae lack such recumbent setae on the basal four antennomeres in most cases (21-2), but
Nitidotachinus and members of the former Megarthropsini lack these setae on only the basal two antennomeres (21-3). Deropini has a densely pubescent antenna, but there is no clear borderline found in the basal third (21-0).
22. Antennomere 4. (0) without markedly long setae (Figure 68F); (1) with markedly long setae, longer than 2.2 times of width of antennomere IV.
Two subfamilies, i.e., Trichophyinae and Habrocerinae, share the characteristic verticillate antennae, bearing very long setae sparsely on each antennomere ([
189]: Figure 1.21.2).
23. Labrum, divided into two halves by medial longitudinal suture (modified from Ashe [
28]: character 104): (0) neither divided nor thinner longitudinally along midline of labrum (Figure 67D); (1) not divided, but thinner longitudinally along midline of labrum; (2) longitudinally divided along midline of labrum.
In Habrocerinae, the labrum is divided into two halves by a medial longitudinal suture ([
192]: Figure 1C).
24. Labrum, anterior margin, lateral areas, setose or spinose processes composed of filiform setulae (dorsal view) (modified from Ashe [
28]: character 1): (0) absent or very weakly developed, only sparsely distributed (Figure 67D); (1) well-developed, conspicuous (Figures 9B, 16D, 26C, 45A, and 53H).
In general, the Tachyporinae have a pair of well-developed setose or spinose processes on the anterior margin of the labrum, whereas most Mycetoporinae lack such setose processes. However, it should be tested in future if these setose processes are homologous because of the variations on the locations, size, and structures.
25. Labrum, anterior margin, medial area (dorsal and ventral views) (Ashe [
28]: character 93): (0) without row of flattened, spatulate or modified setae (Figure 67D); (1) with a row of flattened, spatulate or modified setae (Figures 9B, 16D, 26C, 45A, and 53H).
Most of the outgroups and Mycetoporinae lack a row of flattened, spatulate, or modified setae in the anteromedial margin of the labrum, but Tachyporinae typically have the peculiar, modified setae.
26. Right mandible, number of subapical teeth (modified from Ashe [
28]: character 15) (dorsal and ventral views): (0) absent (Figures 9B, 16A, 45A, and 67A); (1) present, one small; (2) present, one large; (3) present, two large.
In Tachyporinae and Mycetoporinae, there is no subapical tooth along the inner edge of the right mandible. However, four subfamilies (Phloeocharinae, Aleocharinae, Trichophyinae and Olisthaerinae) in the Tachyporine Group of subfamilies have such subapical teeth.
27. Left mandible, number of subapical teeth (modified from Ashe [
28]: character 16) (dorsal and ventral views): (0) absent (Figures 9B, 16A, 67A, and 68E); (1) present, one large; (2) present, one large, one small; (3) present, two large.
In Tachyporinae and Mycetoporinae, there is no subapical tooth along the inner edge of the left mandible. However, four subfamilies (Phloeocharinae, Aleocharinae, Trichophyinae and Olisthaerinae) in the Tachyporine Group of subfamilies have such subapical teeth.
28. Mandible, spinose molar lobe (modified from Ashe [
28]: character 17) (ventral view): (0) present (Figures 9B, 16C and 67A); (1) absent.
Except for Oxypoda sp. in the ‘higher’ group of Aleocharinae and Quedius paradisi Hatch in Staphylininae, all taxa studied here have a spinose molar lobe in each mandible. They are well-developed in Tachyporinae, but much smaller in Mycetoporinae (although it is difficult to separate the coding).
29. Maxillary palpus, structure: (0) 4-segmented; (1) 5-segmented.
The maxillary palpus is comprised of five palpomeres in Trichophyinae: the fourth maxillary palpomere is large and spindle-shaped, but the terminal, fifth palpomere is minute and hyaline [
16]. Both Mycetoporinae and Tachyporinae have 4-segmented maxillary palpi.
30. Maxillary palpomere 3, shape (dorsal and ventral views): (0) widest at or near apex (Figures 10C,D, 27C, 45D, 53C, 59A and 68A,B,D,E); (1) widest around middle (Figures 10A, 16E and 68C).
Most Mycetoporinae have an apically broadened maxillary palpomere 3. In contrast, two types of palpomere 3 were found in Tachyporinae. Within Tachyporini, all taxa, except Lamprinus and Lamprinodes have state 1. The remaining three tribes, Vatesini, Deropini, and Tachinusini, had state 0.
31. Maxillary palpomere 3, length compared to palpomere 2 [dorsal and ventral views]: (0) not distinctly shorter than maxillary palpomere 2 (>0.7×) (Figures 10A, 27B,C and 68A,D); (1) distinctly shorter than maxillary palpomere 2 (≤0.7×) (Figures 27A, 45D, 53C and 59A).
In Mycetoporinae, Tachyporini, and nearly all of the outgroups, maxillary palpomere 3 is not distinctly shorter than the maxillary palpomere 2. In contrast, some members of Vatesini (Cileoporus, Cilea, and Termitoplus), Deropini, and Tachinusini have palpomere 2 distinctly longer than 3.
32. Maxilla, maxillary palpomere 3, micropubescence setation (dorsal and ventral views): (0) moderately pubescent, with less than 20 setae (>10) (Figure 68D); (1) strikingly densely setose, covered with dozens of setae (>20) (Figures 10A, 16E, 27C and 68A); (2) only sparsely setose, with single to ten scattered stout setae, usually restricted to apex of palpomere (Figures 27A, 53C, 59A, and 68E); (3) completely glabrous (Figure 45D).
This character is variable among the members of Mycetoporinae (32: 0-2). The maxillary palpomere 3 in Tachyporini is densely pubescent, whereas the other tachyporine tribes have some variations. In most Vatesini, the palpomere is nearly glabrous, bearing a few scattered setae, but it is moderately pubescent in Mimocyptus and densely pubescent in Vatesus (in Termitoplus, they are very short, but sparsely scattered). In Tachinusini, all genera have a sparsely setose palpomere except Leucotachinus (moderately pubescent). Uniquely, palpomere 3 of Deropini is completely glabrous.
33. Maxilla, maxillary palpomere 4, shape (dorsal and ventral views): (0) moderately to strongly pointed apically, widest at or near base (Figures 10C, 16E, 27A, and 68B,D); (1) only weakly to strongly narrowing apically in apical half, greatest width in basal 1/6 to middle (Figures 45D, 53C, 59A and 68A,E); (2) widest near middle, abruptly narrowed near apex; (3) markedly slender, aciculate (Figure 68C).
Among the taxa of Mycetoporinae, three different types of the maxillary palpomere 4 were found. Parabolitobius, Bolitobius, and Bobitobus (B. lunulatus) were coded as ‘1′, while Mycetoporus and Ischnosoma have unusually slender, aciculate palpomere 4, scored as ‘3′. The rest of Mycetoporinae were coded as ‘0′ as they have strongly pointed ones. In Tachyporinae, Tachyporini and Vatesini were coded as ‘0′, whereas Deropini and Tachinusini, were scored as ‘1′.
34. Maxilla, maxillary palpomere 4, length (dorsal and ventral views): (0) short, much shorter than palpomere 3 (0.2–0.75×) (Figures 10C, 16E, 27C and 68C); (1) moderate, subequal or slightly shorter than palpomere 3 (>0.75–1.1× ≤) (Figure 68B); (2) long, clearly longer than palpomere 3 (>1.1×) (Figures 27A,D, 45D, 53C, 59A and 68E); (3) distinctly short, much shorter than palpomere 3 (<0.2×).
In Mycetoporinae, this character state was generally coded as either ‘1′ or ‘2′, with the exceptions of Mycetoporus and Ischnosoma (both coded as ‘0′). It was coded as ‘0′ in Tachyporini, whereas it was generally scored as ‘2′ for Vatesini, Deropini, and Tachinusini, except Vatesus (34-0) and Mimocyptus (34-1) (both Vatesini).
35. Maxilla, maxillary palpomere 4, maximum width in apical half (dorsal and ventral views): (0) moderately to distinctly narrower than maximum width of palpomere 3 (Figures 10C, 16E, 27C and 68C); (1) the same as, or only slightly narrower, than maximum width of palpomere 3 (Figures 45D, 53C and 68A).
This character state was generally coded as ‘0′ in Mycetoporinae, Tachyporini, and Vatesini and as ‘1′ in Deropini and Tachinusini.
36. Maxilla, maxillary palpomere 4, pores (dorsal and ventral views): (0) absent (Figures 10C and 16E); (1) present, with only a few small, inconspicuous pores; (2) present, with several to dozens of inconspicuous, or conspicuous, pores (Figures 45D, 53C, and 68E).
Nearly all members of Mycetoporinae have several to dozens of the inconspicuous, or conspicuous, pores on the maxillary palpomere 4. The presence of such pores are widely found in the Deropini + Tachinusini clade. However, the rest of Tachyporinae generally lack the pores on the palpomere 4, with some exceptions in Vatesini.
37. Maxilla, lacinia, apical unarticulated large, long spine (Grebennikov & Newton [
2]: character 124) (dorsal and ventral views): (0) absent (Figures 16E, 27B, 45D and 53F); (1) present (Figures 10C and 68A).
In Mycetoporinae and Tachyporina, there is a conspicuous, well-developed long spine located at the apex of the maxillary lacinia. In Sepedophilus (Euconosomatina), it is so small and inconspicuous that it is considered as absent here. I could not find such a long spine in the tribes Vatesini (except Mimocyptus), Deropini, and Tachinusini.
38. Maxilla, lacinia, length of at least some of teeth or spines (dorsal and ventral views): (0) longer than maximum width in larger part of apical third of lacinia (Figures 16E, 36C and 68B); (1) as long as or shorter than maximum width in larger part of apical third of lacinia (Figures 27B, 45D, and 53C).
In Mycetoporinae (except Mycetoporus and Ischnosoma) and Tachyporini, some of the teeth or spines along the inner margin of the maxillary lacinia are longer than the maximum width of the apical third of the lacinia. However, those of Vatesini, Deropini, and Tachinusini are shorter because of the shorter teeth (spines) and much broader apex of the lacinia. Among these three tribes, the only exception was Mimocyptus (Vatesini), which was scored as ‘0′.
39. Labium, labial palpomere 2, shape (dorsal and ventral views): (0) elongate to only weakly transverse (Figures 16F, 26B 45B, 53D, and 59B); (1) strongly to markedly transverse (Figures 26E,F and 67E).
All members of Mycetoporinae, except Mycetoporus, have a strongly transverse labial palpomere 2. Within Tachyporinae, the earlier-branching Vatesini and all Tachyporini have elongate to only weakly transverse ones, whereas the remaining tachyporine taxa were scored as ‘1′.
40. Labium, labial palpomere 3, shape (dorsal and ventral view): (0) widest at base (Figures 9E, 16F and 26E); (1) widest around middle or near base, not at base (Figures 26B,F, 45B, 53D,E, 59B and 67F); (2) narrowly elongate, straight, slender, subparallel-sided; (3) narrowly elongate, curved, slender, subparallel-sided; (4) nearly subparallel-sided, thick, weakly broadened apically, with truncate apex (Figure 67E); (5) distinctly expanded, crescent-shaped.
Most members of Mycetoporinae were coded as ‘1′, with two exceptions. Most Tachyporinae were scored as either ‘0′ or ‘1′. All members of Tachyporini, except
Euconosoma (Euconosomatina), were coded as ‘0′.
Euconosoma has a distinctly widened, laterally compressed, crescent-shaped labial palpomere 3 (see Campbell [
55]: Figure 2). In contrast, the tribes Vatesini, Deropini, and Tachinusini were coded as ‘1′ except
Mimocyptus (Vatesini), which was scored as ‘0′.
41. Labium, labial palpomere 3, maximum width (dorsal and ventral views): (0) distinctly to moderately narrower than maximum width of palpomere 2 (<0.75×) (Figures 9E, 16F, 26E and 67E); (1) almost same width or only slightly narrower than maximum width of palpomere 2 (0.75× ≤, <1.10×) (Figures 26B,F, 45B, and 59B); (2) moderately wider than maximum width of palpomere 2 (1.10–1.50×) (Figure 67F); (3) distinctly wider than maximum width of palpomere 2 (>1.5×).
Three states were found in Mycetoporinae, and they were scored from ‘0′ to ‘2′. In Tachyporinae, Tachyporini was generally coded as ‘0′, whereas Deropini and Tachinusini were mostly coded as ‘1′. Vatesini were either ‘0′ or ‘1′.
Euconosoma (Euconosomatina) has a distinctly widened labial palpomere 3, and was scored as ‘3′ ([
44]: Figure 1; [
55]: Figure 2).
42. Labium, labial palpomere 3, with a few to several thick, short peg-like spines on surface (dorsal and ventral views): (0) absent (Figures 16F and 26B); (1) present (Figures 26E and 67E).
Some genera of Mycetoporinae and Vatesini have a few to several thick, short peg-like spines on the outer margin of the labial palpomere 3 (
lp3) (e.g., Ashe [
28]: Figure 84). The location of these spines has some variations from near lateral base to the apex of lp3. However, it is difficult to assess the homology of this character state, I tentatively scored them the same state (although it is highly unlikely homologous).
43. Labium, labial palpomere 3, pores (dorsal and ventral views): (0) absent; (1) present, inconspicuous (Figures 9E, 16F, 26B, and 67F); (2) present, conspicuous (Figures 45B, 53D and 59B).
In Mycetoporinae, some Vatesini, and Sepedophilus (one species only, Euconosomatina), there are inconspicuous pores scattered on the labial palpomere 3. Contrary to these taxa, I observed much larger and conspicuous pores on this palpomere in Deropini and most Tachinusini.
44. Labium, ligula, shape (ventral view): (0) large, simple, as long as wide to transverse (Figures 9E and 45B); (1) small, simple, restricted to median area (Figure 67F); (2) pair of medial lobes moderately present (Figure 53E); (3) pair of medial lobes weakly present, with spines on the lobes (Figures 16F and 53B); (4) markedly narrowly elongate, pointed apically; (5) large, transeverse, with spines in middle.
In Mycetoporinae, the general shape of the ligula (glossae and paraglossae) is small and simple, although they occasionally have moderately developed medial premental lobes (
Bryoporus). In Tachyporinae, the ligula is much larger and wider than that of Mycetoporinae. Interestingly, some
Sepedophilus (Euconosomatina) have a pair of weakly developed medial premental lobes, occasionally together with a few conspicuous peg-like spines at each apex ([
28]: Figure 86). It should be also mentioned that
Austrotachinus ([
110]: Figure 2) and
Olophrinus (Figure 53E) (both Tachinusini) each have a pair of the moderately developed medial premental lobes, and were scored as ‘2′.
45. Labium, ligula, extent (dorsal and ventral views): (0) small, slightly extended laterally, but not beyond each labial palpus (Figure 67E,F); (1) medium to large, clearly extended laterally beyond each labial palpus (Figures 9E, 16F, 26B, 45B, 53E, and 59B); (2) elongate and very large, but limited to median area only, not extended laterally beyond middle of each labial palpus.
Mycetoporinae have a small ligula, whereas those of Tachyporinae are much wider and larger, extending laterally beyond the outer margins of each labial palpus (e.g., Ashe [
28]: Figures 84–86).
46. Labium, insertions of labial palpi (modified from Grebennikov & Newton [
2]: character 131) (dorsal and ventral views): (0) narrowly to moderately separated, closer than length of longest labial palpomere, sometimes even narrower than maximum width of labial palpomere 1 (Figures 45B, 53D, and 67E); (1) more widely separated, slightly narrower than, or subequal to, narrower, or subequal, to longest labial palpomere (Figures 9E, 16F and 26E); (2) widely separated, clearly wider than longest labial palpomere (Figures 26B,F, 53G and 59B).
In Mycetoporinae, the insertions of the labial palpi are close, much closer than the length of longest labial palpomere. Some variation was found in Tachyporinae. Within Tachyporini, they were rather widely separated in most cases (except in some Euconosomatina). In Vatesini, the labial palpi are generally widely separated, except in Mimocyptus. They are very narrowly separated in Deropini, while Tachinusini has much wider separation, except in the semi-aquatic genus Nitidotachinus.
47. Mentum, anterior margin (ventral view): (0) more or less truncate (Figure 53B); (1) broadly shallowly emarginate; (2) broadly rounded (Figure 26F); (3) broadly strongly emarginate; (4) pointed in middle.
In Mycetoporinae, the anterior margins of the mentum are always truncate. However, some Tachyporinae have a broadly rounded anterior margin of the mentum (e.g., Euconosomatina and part of Vatesini).
48. Mentum, shape (ventral view): (0) weakly to moderately transverse, <2.85× as wide as long (Figures 26F and 53B); (1) strongly transverse, ≥ 2.85 × as wide as long; (2) distinctly narrowed towards apex.
In Mycetoporinae, the ratio of the mentum is more than 2.85 times as wide as long in most cases, with the exceptions of Bobitobus and Mycetoporus. In Tachyporini, they are strongly transverse, but those of Vatesini, Deropini, and most of Tachinusini (except Olophrinus and Tachinomorphus) have a relatively transverse mentum, here considered as ‘0′.
49. Pronotum, shape (dorsal view): (0) not constricted or distinctly narrowed (Figure 64B); (1) abruptly narrowed posteriorly, moderately constricted in posterior half (Figure 44A); (2) weakly constricted laterally at base; (3) weakly constricted anteriorly and posteriorly; (4) forming unique corners along plate-like lateral projections (Figure 58C).
In Deropini, the pronotum is unusually strongly constricted in the posterior half. Furthermore, two genera of Tachinusini (Megarthropsis and Nepaliodes) have only weakly constricted pronotum along the plate-like lateral margins, differing completely from the modifications of Deropini.
50. Pronotum, maximum width, location (determined from pinned specimens) [dorsal view]: (0) at base (Figures 15C, 24D, and 51C); (1) between near base and basal 1/5 (Figures 7D and 27G); (2) between basal 1/5 and basal 1/4 (Figures 8A, 51D); (3) between basal 1/4 to middle (Figures 7B, 24A, 51A,B, 58A, 59D,E, 64B,D, 65D, and 69A); (4) in anterior half, but not near anterior margin (Figure 44A); (5) in anterior half, near anterior margin.
In Mycetoporinae, the maximum width of the pronotum is located between the basal 1/4 to middle because of the strongly rounded posterior pronotal margins. A series of variations are found in Tachyporinae: Tachyporina was coded as ‘1′, with the exception of Palporus and Symmixus, which were considered as ‘3′ and ‘2′, respectively; Euconosomatina was scored as ‘0′; Vatesini was coded with ‘0′, ‘1′, or ‘3′; Deropini was considered as ‘4′; Tachinusini was mostly scored as ‘3′, but some were considered to be either ‘2′ (Olophrinus) or ‘0′ (Tachinomorphus).
51. Pronotum, apical half, uniform punctation (modified from Herman [
25]: character 0) (dorsal view): (0) absent (Figures 7D, 24D, 51D, and 64B); (1) fine; (2) dense, but shallow (Figure 44A); (3) coarse and small but relatively deep; (4) coarse and large, deep, pit-like (Figure 58A,B).
All taxa of Mycetoporinae lack uniform punctation, and the surface of the forebody is strongly glossy. In Tachyporinae, the tribes Tachyporini and Vatesini also do not have conspicuous punctures on the forebody. However, some members of Tachinusini (= Megarthropsini) have well-developed, pit-like punctures. The pronotum of Deropini is covered with dense but shallow punctation.
52. Pronotum, surface [dorsal view]: (0) more or less even (Figures 51D and 64B); (1) rough, with indentations (Figures 51B and 58B).
Excepting some outgroups and part of Tachinusini (e.g., Herman [
25]: Figure 183), the surface of the pronotum is even dorsally without grooves. There are some variations of indentations found in limited in- and out-group taxa. Since it is difficult to assess the homology, they were treated as a single state.
53. Pronotum, microsetae, except for characteristically arranged macrosetae [dorsal view]: (0) glabrous (Figures 7D, 24D, 51D, 58A, and 64B); (1) pubescent, sparsely covered with very minute setae; (2) pubescent, densely covered with thin, moderately long setae (Figures 15B and 44A); (3) pubescent, covered with extremely short, thick, distinctly modified, vestigial setae (Figure 58C).
In Mycetoporinae, the surface of the pronotum is uniformly glabrous, with characteristically arranged, marginal macrosetae. Within Tachyporini, it is generally glabrous in Tachyporina, whereas the setae are moderately long and conspicuous in Euconosomatina. In Vatesini, it is completely glabrous, except in
Cilea. The pronotum of Deropini is densely pubescent, and genenerally glabrous in Tachinusini sens. str. Notably, however,
Nepaliodes in Tachinusini has distinctly short, modified setae on the pronotum ([
25]: Figure 186).
54. Pronotum, dorsolateral marginal areas, modification (dorsal view): (0) normal, not modified (Figures 51D and 64B); (1) weakly explanate (Figure 58A); (2) strongly explanate (Figure 58B); (3) foveate; (4) serrate.
In limited Tachinusini (= Megarthropsini), the dorsolateral marginal areas of the pronotum are weakly to strongly modified, and the most extreme case is found in
Nepaliodes ([
25]: Figure 183).
55. Pronotal apex (= anterior margin) (dorsal view): (0) truncate (Figure 44A); (1) weakly to moderately concave (Figures 7A, 18A, 51B, 27E, 54G, 59D,E, and 64B); (2) strongly concave; (3) distinctly modified, markedly strongly concave (Figure 58B).
As a general trend, the anterior margin of the pronotum is generally weakly to strongly concave in Mycetoporinae and Tachyporinae. However, it is truncate in Deropini, Vatesus gigas (Vatesini), and some mycetoporines.
56. Pronotum, anterior/front angles (dorsal and ventral views): (0) broadly rounded (Figures 51B, 58A, and 59D,E); (1) angulate (Figures 27G, 44A, 64B and 69A); (2) sharply pointed (Figure 7A, 15A, 18A); (3) narrowly rounded, strongly produced [
25] (Figure 58B,C).
The anterior angles of the pronotum vary greatly as follows: from rounded (Tachinusini), angulate (Mycetoporinae, Vatesini (except Vatesus gigas), and Deropini), sharply pointed (Tachyporini), to having a peculiar form (Nepaliodes in Tachinusini).
57. Pronotum, macrosetae [dorsal view]: (0) absent (Figures 15C and 51D); (1) present, characteristically arranged (Figures 7A, 64A,C, and 69A).
Characteristically arranged setae on the pronotal margins were found in Mycetoporinae and Tachyporina. In
Leucotachinus (Tachinusini), margins of the pronotum are furnished with series of discrete punctures, each bearing a short seta [
91], but they are minute and considered as absent.
58. Pronotum, transverse cluster of blackish to blackish-brown small pores near posterior margin [dorsal and ventral views]: (0) absent (Figures 11A, 18A,B, 27E,G, 45E, 59D,E, and 69A,B); (1) present (Figure 54G).
This character state can only be seen in slide-mounted specimens. In some Vatesini and most members of Tachinusini, there are some transversely scattered clusters of blackish brown pores located near the posterior margins of the pronotum. Other taxa have much more weakly pigmented or ambiguous pores, or even lack such pigmented pores. The structural details and the assessment of its homology are needed to be assessed in future.
59. Pronotal hypomeron, transverse ridge (ventral view): (0) absent (Figures 11A, 18B, 27F, 54G and 59D); (1) present, usually located in anterior 1/3 to 1/4 of each pronotal hypomeron (Figure 69A,B).
In Mycetoporinae and some outgroups, there is a prominent ridge located in the anterior area of each pronotal hypomeron, observable in ventral view.
60. Pronotal hypomeron, width in basal third (ventral view): (0) narrow (Figures 45E, 69A, B); (1) moderate (Figures 11A and 54G); (2) wide (Figures 18B, 27G, and 59D,E).
The maximum width of the pronotal hypomeron greatly varies among the studied taxa. For example, it is narrow in Mycetoporinae and Deropini; moderate in most Tachyporina (except Symmixus) and most members of Tachinusini; and wide in Symmixus, Euconosomatina, Vatesini, and some Tachinusini.
61. Pronotal hypomeron, postcoxal process (ventral view): (0) absent, at most only very feebly projecting inward (Figure 69A); (1) weakly developed (wider than long), with rounded apex (Figure 27E); (2) weakly developed (wider than long), with truncate apex (Figure 18A); (3) weakly developed (wider than long), with pointed apex; (4) moderately developed (as long as wide to slightly longer than wide) (Figures 18B and 45E); (5) fully developed (clearly longer than wide), usually with rounded or narrowly rounded apex (Figures 11A, 54G and 59D,E).
A series of variations were found in the lengths and structures of the pronotal postcoxal processes on the pronotal hypomeron. They are completely absent in Mycetoporinae, whereas they are weakly developed in Vatesini, moderately to strongly projecting in Tachyporina and the clade Deropini + Tachinusini. Interestingly, there are some variations found in Euconosomatina. In Sepedophilus littoreus, it is strongly tapered with a thin attachment. In this study, the attachment was not considered as a special feature because it is fully concealed under the procoxae, rather than forming a part of the lobe-like structure. The significant variations of the pronotal postcoxal processes found in the studies taxa suggests that their assessment of homology is needed. Nonetheless, it was not possible to do so in this study due to the difficulty for assessing such a morphologically diverse in- and out-group taxa.
62. Pronotal hypomeron, pronotal postcoxal process and its related areas, base: (0) without suture; (1) with suture that separates postcoxal process from rest of pronotum, resulting in a slightly movable process.
The base of the postcoxal process and related areas of the pronotum is separated by a suture from the rest of the pronotum in Olisthaerinae (see Newton et al. [
16]: Figure 63.22).
63. Prosternum, transverse sternacoxal ridge (
tsr) (sensu Clarke [
204]: Figure 2E), medio-lateral areas (ventral view): (0) ridge distinct, moderately distant from anterior margin of prosternum (Figures 11A, 45E, 52A, 54A, 58D and 59D); (1) ridge very close to anterior margin of prosternum (Figures 18C, 24A and 27E); (2) ridge indistinct, trace only, moderately distant from anterior margin of prosternum (Figure 69A,B); (3) ridge far distant from anterior margin of prosternum.
In Mycetoporinae, the transverse sternacoxal ridge (sensu Clarke [
204]: Figure 2E) is merely a trace with a row of setae, which is moderately distant from the anterior margin of the prosternum (Figure 69A,B). Tachyporina, Deropini, and most members of Tachinusini have the prominent (not reduced) sternacoxal ridges moderately distant from the anterior margins of the prosternum. In contrast, Euconosomatina and Vatesini have the ridges located very close to the anterior margins of the prosternum.
64. Prosternum, strongly developed prosternal process (ventral view): (0) absent (Figures 18A, 52A, 54A, and 69A); (1) present, partially separating procoxae (Figures 58D and 59D); (2) present, fully separating procoxae.
In Mycetoporinae and Tachyporinae, the medial prosternal process is usually at most only feebly developed. However, some Tachinusini (members of the former Megarthropsini) have a relatively developed prosternal process partially separating the procoxae.
65. Furcasternum, posterior margin (ventral view): (0) various, moderately produced posteriorly, shallowly emarginate to broadly rounded (Figures 11A, 18C, 45E, 54A, and 69A); (1) laterally expanded, contiguous with pronotal projections of hypomeron (Figure 18A).
Uniquely in Tachyporinae, a few Sepedophilus (Euconosomatina) have a laterally expanded, modified posterior margin of the furcasternum, thereby closing the procoxal cavities.
66. Procoxal cavities (ventral view): (0) externally open behind (coxa only partially encircled by pronotal postcoxal process and modified furcasternum) (Figures 11A, 18C, 27E, 54G, and 59D); (1) externally completely closed behind, coxa fully encircled by postcoxal process of pronotum and modified furcasternum (Figure 18A); (2) nearly fully or fully open behind, lacking pronotal postcoxal process (Figure 69B).
In several outgroups and Mycetoporinae, the procoxal cavities are fully opened behind because of the absence of the pronotal postcoxal processes. However, in Tachyporinae, they are not fully opened behind because of the weakly to strongly developed pronotal postcoxal processes. In three species of Sepedophilus (Euconosomatina), they are fully encircled by the postcoxal process of the pronotum and the modified furcasternum; thus, they are completely closed behind.
67. Mesospiracular peritremes (msp) (sensu Blackwelder [
30]: Figure 3A,E), degree of sclerotization [ventral view]: (0) only weakly sclerotized or even membranous (Figure 69A,B); (1) partially well sclerotized; (2) fully well sclerotized, conspicuous (Figures 18A and 52B).
In Mycetoporinae, they are only weakly sclerotized and membranous, but those of most Tachyporinae are large and well sclerotized, except Leucotachinus (Tachinusini) (the limited areas with spiracles are sheet or plate-like, extending inward, whereas the out-groups and Mycetoporinae have smaller sheet or plate-like fields, replaced with membranes).
68. Scutellum, prescutoscutellar suture (pss) (sensu Blackwelder [
30]: Figure 4A,C) lying transversely [dorsal view]: (0) located near base (Figure 11E); (1) located apically, near middle of scutellum (Figure 69D,E).
In the two subfamilies Mycetoporinae and Staphylininae (
Quedius), there are two transverse ridges, namely: basal carina (Figure 69D,E,
bc) and apical carina (Figure 69D,E, ac) (sensu Campbell [
32]). The structure of basal carina has been used as one of the diagnostic features of the genera of Mycetoporinae [
32,
33]. I consider the position of apical carina (or prescutoscutellar suture, sensu Blackwelder [
30]: Figure 4A,C) as a subfamily-level character state. In Mycetoporinae, apical carina is distant from the base of the scutellum and located apically near middle of the scutellum, whereas that of Tachyporinae is lying near the base of the scutellum.
69. Elytra (as a pair), overall shape (dorsal view): (0) narrowed posteriorly (Figures 15C, 24D and 51C); (1) widest in middle (Figures 21A and 33B); (2) posteriorly broadened (Figures 51A and 66A).
In Mycetoporinae, the elytra are posteriorly broadened (although Parabolitobius is nearly parallel sided). Within Tachyporinae, they are narrowed posteriorly, or widest around the middle of the elytra, in Tachyporini and Vatesini. The elytra are posteriorly broadened in Deropini. Although there are some variations, posteriorly broadened elytra are most common in Tachinusini.
70. Elytron, ratio of length/width: (0) > 1.8× (Figures 44C, 69F); (1) 1.3–1.8× (Figures 11E and 18F); (2) < 1.3× (Figure 30A,B).
The ratio of the elytral maximum length and width were measured usually from slide specimens. A narrowly elongate elytron was found in Mycetoporinae, and therefore, coded as ‘0′. Most taxa of Tachyporini have moderately long elytra, and were scored as ‘1′. Two types of elytra were found in Vatesini: moderately long in earliest diverging genera (
Cileoporus and
Cilea, and short in the rest of the tribe. Within the Deropini + Tachinusini clade, the general state is having moderately to strongly elongate elytra. Of note,
Sepedophilus nigeriensis (= ‘
Urolitus’
nigeriensis; Euconosomatina) was measured using a habitus drawing from the original description [
54].
71. Elytron, longitudinal rows of setigerous punctures (excluding ground setae) (dorsal view): (0) absent (Figures 11E, 24D, 30A and 51A,C); (1) present, less than three rows of setigerous punctures; (2) present, three rows of setigerous punctures (Figure 65A); (3) present, more than three rows of setigerous punctures (Figure 65D).
In Mycetoporinae, there are a few to several longitudinal rows of setigerous punctures on each elytron, although they are sometimes difficult to determine (for
Parabolitobius, see Li et al. [
152]: Figures 8 and 12). In this study, each puncture bearing a stout seta was considered for this character state.
72. Elytra, evenness (excluding sutural edge of each elytron) (dorsal view): (0) even (Figures 18F, 30A and 51A); (1) uneven, wrinkled; (2) uneven, with indistinct rows of punctures without deep grooves (Figure 65D); (3) uneven, with distinct rows of punctures with weak to deep longitudinal depressions (Figure 51D); (4) uneven, weakly costate; (5) distinctly uneven, strongly costate.
The elytral surface greatly varies from even to uneven because of the presence or absence of indentations. Since the difficulty of the accurate assessment of homology, they are scored as a single character here. In some genera of Mycetoporinae, there are indistinct rows of punctures without deep grooves. Within Tachyporinae,
Olophrinus (Tachinusini) is an unusual genus that has several longitudinal striae on each elytron based on distinct discal rows of punctures [
126,
127]. In
Olophrinus, each puncture is connected with a longitudinal dent-like impression, forming a shallow groove for each puncture row. This condition is probably not homologous to that in
Neophonus, but both tentatively scored as the same state due to the difficulty of the accurate assessment of the homology. The extinct tachyporine genus †
Protachinus from the Late Jurassic Talbragar Fish Bed in New South Wales, Australia, has similar elytral longitudinal striae [
52]. The rest of the extant and extinct tachyporines lack such rows of longitudinal striae on the elytra.
73. Elytra, punctation (including setigerous punctures) (dorsal view): (0) absent or fine (Figures 18F, 24D, 30A and 51A); (1) shallow, pit-like (Figures 44C and 51D); (2) deep, pit-like (Figures 58A,C and 60D); (3) somewhat indistinct, mostly distorted (Figure 65A,D).
Some members of Mycetoporinae have somewhat indistinct, mostly distorted punctation on the elytra. In Tachyporinae, they are generally smooth without conspicuous punctation. However, those of Deropini are covered with somewhat deep punctures. In Tachinusini, the general punctation on the elytra is either absent or shallow, but some have deep punctation and Olophrinus has longitudinal rows of the punctures as mentioned above (character 72).
74. Elytra, ground microsetae (excluding rows of setigerous punctures) (dorsal view): (0) glabrous (Figures 24D, 30A, 51A, and 65B); (1) vestigial (Figures 8D, 25A and 58C); (2) pubescent (Figures 7B, 11E, 15C, 18F, and 44C).
The elytra of Tachyporini are generally covered with microsetae, whereas the remaining tachyporines lack such elytral microsetae. It should be mentioned, however, that Deropini is densely pubescent, and some Tachinusini have very short, modified setae.
75. Elytron, blackish macrosetae on median area (excluding lateral, inner, or posterior margins) (dorsal view): (0) absent (Figures 30A and 65B); (1) present (Figures 7B, 8A, and 11E).
In most taxa of Tachyporina, the blackish macrosetae are located on the median area of each elytron, but they are absent in the two myrmecophilous genera: Lamprinodes and Lamprinus.
76. Elytron, blackish macrosetae along lateral margin [dorsal and lateral views]: (0) absent (Figures 15C, 24D, 30A, and 65A); (1) present (Figures 7D, 11E, and 18F).
Within Tachyporinae, the members of Tachyporina and a few
Sepedophilus species of Euconosomatina have a row of blackish macrosetae located along the lateral margin of each elytron. Additionally, an enigmatic tachyporine genus
Tachinoporus has a row of five setae laterally on each elytron [
11].
77. Elytron, blackish macrosetae along posterior margin [dorsal and lateral views]: (0) absent (Figures 15C, 30A, and 65A); (1) present (Figures 11E and 18F).
Within Tachyporinae, the members of Tachyporina and Sepedophilus (scriptus species group only; Euconosomatina) have blackish macrosetae arranged along the posterior margins of the elytra. Although I considered them present here in the myrmecophilous Lamprinodes of Tachyporina, they are short and rather inconspicuous.
78. Elytra, sutural edge, longitudinally raised suture [dorsal view]: (0) absent or only feebly raised (Figures 7D, 24D, and 51A,C); (1) present, moderately somewhat broadly raised (Figures 65C and 66A); (2) present, markedly strongly raised.
Mycetoporinae have a longitudinally raised suture along the inner edge of each elytron, whereas such raised sutures are absent in Tachyporinae.
79. Elytra, outer margin, epipleural gutter (see Herman [
25]) [dorsal view]: (0) absent (Figures 7B, 15C, and 44C); (1) present, very narrow, inconspicuous (Figures 7D, 51A,D, and 65D); (2) present, narrow (Figures 24D and 58A); (3) present, wide, prominent, and resulting in lateral margins of elytra being concave and/or strongly explanate (Figure 58C).
A very narrow, inconspicuous epipleural gutter on the lateral margin of each elytron is found in Mycetoporinae, Tachyporina (except Palporus), some Vatesini, and most of Tachinusini. The remaining Vatesini have a narrow epipleural gutter along each elytral outer margin. Such epipleural gutter is absent in Deropini, Euconosomatina, a few members of Vatesini, and Palporus (Tachyporina). The strongly expanded, markedly wide epipleural gutters are found in limited members of Tachinusini (= Megarthropsini).
80. Elytral epipleural ridge/keel (Ashe [
28]: character 46; Grebennikov & Newton [
2]: character 165) (lateral view): (0) present (Figures 52C and 64C); (1) absent.
The elytral epipleural ridge/keel is absent in the outgroup subfamilies Aleocharinae (Gymnusa and Oxypoda), Phloeocharinae (Phloeocharis), Trichophyinae (Trichophya), and Staphylininae (Quedius).
81. Elytra, folded inward elytral epipleuron, degree of development (ventral view): (0) not or only feebly folded; (1) moderately folded (Figures 18F, 52B,C and 60D); (2) strongly folded (Figure 24A); (3) very strongly, distinctly folded (Figures 24B, 30A,B and 58D).
In Mycetoporinae, the elytral epipleuron is only weakly folded inward, whereas those of Tachyporinae are clearly folded, with a few variations as follows: only feebly folded (Deropini); moderately folded (Tachyporini and most Tachinusini), strongly folded (Vatesini and part of Tachinusini).
82. Elytra, lateral and posterior margins (dorsal view): (0) normal, thick (Figure 15C); (1) thin, blade-like (Figures 25A and 30B,C); (2) of normal thickness on lateral margins, but exceptionally thin in posterior areas, resulting in semitransparent posterior margins of the elytra (Figure 8E).
Vatesini and the two genera of Tachinusini have thin, blade-like edges of the lateral and posterior margins of the elytra. In Symmixus (Tachyporina), the posterior areas of the elytra are remarkably thin, resulting in semitransparent posterior margins.
83. Elytra, posterolateral margin (modified from Ashe [
28]: character 45) (dorsal view): (0) not sinuate (Figures 11E, 15C, 24D, 30B, 51C, 58A, 65B, and 69F); (1) weakly to moderately sinuate; (2) strongly sinuate (Figures 25A, 30C and 58C).
The posteriorly sinuate margin of the elytron is one of the characters which defines the subfamily Aleocharinae. However, this condition can also be found in Trichophyinae, Vatesus (Vatesini), and some Tachinusini (members of the former Megarthropsini).
84. Subapical bulge on each elytron near the posterolateral corner (sensu Hansen [
204]) (dorsal view): (0) absent; (1) present.
The presence of a subapical bulge near the posterolateral corner on each elytron is considered one of the possible unique synapomorphies of Silphidae [
205].
85. Mesoventrite, longitudinal median carina (ventral view): (0) absent (Figures 11D, 24A,B, 46A, 54F and 70A,D); (1) present, on main part of mesoventrite (Figures 19A, 24C, 28B, 54C,D, 58D and 60A); (2) present, only on mesoventral process, not on main part of mesoventrite (Figure 28C).
There is no longitudinal median carina in Mycetoporinae, whereas this character may have independently evolved several times in Tachyporinae based on the phylogenetic trees presented in this study. In Euconosomatina, all of the
Sepedophilus taxa studied here have a longitudinal carina on the mesoventrite, but
Euconosoma lacks this derived character. Within Vatesini, four genera, namely
Cilea,
Mimocyptus,
Termitoplus, and
Coproporus, have such a carina, whereas it is completely absent in
Cileoporus,
Vatesus, and
Coprotachinus. In
Cilea (Figure 28C) and
Pseudotachinus [
193,
206], however, the carina is found only on the mesosternal process and its basal area, and therefore I have scored them as “2”. Notably,
Tachinoproporus (Vatesini) has a very strongly developed, plate-like carina. Among the studied taxa of Tachinusini, seven genera, i.e.,
Tachinomorphus,
Olophrinus,
Austrotachinus,
Lacvietina,
Megarthropsis,
Peitawopsis and
Nepaliodes, each have a longitudinal carina on the mesoventrite, and I have scored them as “1”. I could not find this type of carina in Tachyporina and Deropini.
86. Mesoventrite, transverse, long, and straight ridge, lying near base of intercoxal process (ventral view): (0) absent; (1) present (Figures 52A and 54E).
This character was only found in Leucotachinus (Tachinusini).
87. Mesocoxae, separation (modified from Ashe [
28]: character 58) (ventral view): (0) narrowly to moderately separated (Figures 19A, 24B, 28C, 46A, 54D and 70A,E); (1) partially contiguous (at least 1/5 of mesocoxal length) (Figures 52A, 54E and 70D).
In Mycetoporinae, only Parabolitobius and Mycetoporus have partially contiguous mosocoxae, whereas they are narrowly separated, divided by the intercoxal processes of the mesoventrite in the remaining mycetoporines. Within Tachyporinae, contiguous mesocoxae are quite unusual and only seen in Coprotachinus, Mimocyptus (both Vatesini), and part of Leucotachinus (Tachinusini).
88. Metaventrite, intermesocoxal process, length (modified from Ashe [
28]: character 116) (ventral view): (0) short to subequal to that of intercoxal process of mesoventrite (Figures 19A, 24A, 28C, 46A, 52B, 54F, and 70E); (1) much longer than intercoxal process of mesoventrite; (2) extremely short, only slightly protruding apically (Figure 70A,D); (3) absent or nearly absent, only feebly confirmed (Figure 54E); (4) distinctly short, but with developed and only shallowly depressed isthmus (sensu Maruyama [
207]: Figure 7B) projecting apically (Figure 28D); (5) absent or only feebly discernible, but with developed and only shallowly depressed isthmus (sensu Maruyama [
207]: Figure 7B) projecting apically (Figures 24B,C and 28A,B).
Within Mycetoporinae, only
Mycetoporus has a distinctly short, nearly absent, intermesocoxal process of the metaventrite. Most tachyporines have well-developed intermesocoxal processes, but they are nearly absent in some genera of Vatesini, and part of
Leucotachinus (Tachinusini). Additionally, there is a developed isthmus (sensu Maruyama [
207]: Figure 7B) in some genera of Vatesini and
Nitidotachinus (Tachinusini).
89. Metaventrite, intermesocoxal pit (Herman [
25]: character 12) (ventral view): (0) absent; (1) present (Figure 60B).
A deep metaventral pit adjacent to the apex of the mesoventral process is an autapomorphy of
Lacvietina (Tachinusini) [
25].
90. Metendosternite, anterior arms (Naomi [
208]: Figure 1B) (dorsal and ventral views): (0) thin, inconspicuous (Figure 54H); (1) relatively thick; (2) thick, well-developed, sometimes lamellate (Figures 11B, 18D, 28G, and 69G); (3) absent.
The metendosternite is well-developed and strongly lamellate in Mycetoporinae. However, there are a few variations in Tachyporinae from non-developed (most members of both Deropini and Tachinusini) to well-developed and slightly lamellate (most members of both Tachyporini and Vatesini).
91. Metendosternite, posterolateral arms (Naomi [
208]: Figure 1C) (dorsal and ventral views): (0) absent (Figures 11B, 28G and 54H); (1) present, with developed lamellae (Figure 69G).
Mycetoporinae have a pair of heavily lamellate posterolateral arms in the metendosternite, connected with the anterior (distal) arms by a large lamella. However, I could not find such arms in Tachyporinae.
92. Metendosternite, median process (Naomi [
208]: Figure 1H) ([
28]: character 61) (dorsal and ventral views): (0) absent (Figures 11B, 28G, 54H and 69G); (1) present (Figure 28I).
Some outgroup taxa have a single, apically pointed process in the middle of the metendosternite. However, this median process is absent in both Mycetoporinae and nearly all taxa of Tachyporinae. Uniquely, Coprotachinus (Vatesini) has a short, undeveloped, median process on the metendosternite. This is the only example among the studied tachyporines.
93. Legs, form: (0) normal, typical form (Figure 29A); (1) modified, myrmecophilous form (e.g., flattened, robust, with developed lamellae with protective function, see below) (Figures 12A and 29D).
Within Tachyporinae, a similar type of modified legs was found in the two myrmecophilous genera
Lamprinus (Tachyporina) and
Vatesus (Vatesini). In both genera, all legs are short and thick, with strongly developed lamellae extended along the inner margin of each femur. The tibiae are also somewhat flattened and laterally expanded. Furthermore, tarsi are short and thickened, each tarsus appearing compact, reinforced. These leg modifications are known to be adaptations to a myrmecophilous lifestyle, and are considered to perform a protective function against ants [
15].
94. Legs, length: (0) moderate to long; (1) distinctly short, compact.
The legs of Mycetoporinae are generally very long, but some variations are found in Tachyporinae. For example, Deropini and Tachinoporus (Vatesini) have very slender and long legs. Such long legs were also observed in Sepedophilus (Euconosomatina), particularly the hindlegs in which the hindtarsi are exceedingly long. In contrast, the crown genera of Vatesini have distinctly short and compact legs. Two genera of Tachyporina (Lamprinus and Lamprinodes) also possess thick and strong, but distinctly short, legs in association with myrmecophily.
95. Tibia, distributed stout spines (modified from Ashe [
28]: character 47) (dorsal and ventral views): (0) present, conspicuous, irregularly distributed (Figures 12A, 28E, 52C, 66A and 70C); (1) present, inconspicuous, regularly distributed in rows; (2) absent.
It is noteworthy that the mega-diverse ‘higher’ group of Aleocharinae lacks irregularly distributed stout spines on the tibiae, together with some members of the Tachyporine Group of subfamilies, namely Phloeocharinae (Phloeocharis) and Trichophyinae (Trichophya). All members of Mycetoporinae and Tachyporinae have irregularly distributed, tibial spines. Only a single spine on each tibia was found in Mimocyptus (Vatesini), but it was scored here as ‘0′.
96. Procoxa, size (dorsal and ventral views): (0) large, as same as or slightly smaller than profemur (Figure 52A); (1) medium, much smaller than profemur (Figure 45E); (2) very small, globular; (3) distinctly large, much larger than profemur (Figures 12A, 24C, 29A, and 70B).
Mycetoporinae can also be characterized by the presence of markedly large, expanded procoxae. Within Tachyporinae, most tachyporines have large and well-developed procoxae, but Deropini have much smaller ones in proportion to their profemora. The myrmecophilous genus Lamprinus (Tachyporina) and the members of Vatesini possess very large procoxae.
97. Protibia, outer margin with longitudinal row of close-spaced spines (modified from Ashe [
28]: character 100) (dorsal and ventral views): (0) absent (Figure 12A); (1) present (Figure 17B,C).
This character state is a unique synapomorphy of Euconosomatina, with the only probable exception being the monobasic fossil genus †
Palaeosepedophilus from Eocene Baltic amber [
82].
98. Metacoxae lateral half, size (including dorsal lamellae; see
Figure S2) [ventral view]: (0) small (Figures 46B and 58D); (1) medium, much smaller than meso- and metathorax combined (Figures 11C, 15D, 18E, 24A, 52B, 54B, and 60E); (2) markedly large, moderately smaller to nearly as same size as meso- and metathorax combined (Figure 70A and
Supplementary Figure S2A).
See morphological definition of
Supplementary Figure S2A. The metacoxae of Mycetoporinae are markedly large and conspicuous, whereas they are medium in most taxa of Tachyporinae. However, Deropini and
Nepaliodes (Tachinusini) have smaller metacoxae.
99. Metacoxae, dorsal lamellae (or plates, dlmtc) on metacoxae, lateral areas (ventral view): (0) absent or only weakly developed (Figure 46A); (1) moderately developed (Figures 54B and 60E); (2) well-developed (Figures 18E, 24A, 28E, and 70C).
See morphological definition of
Supplementary Figure S2B. The metacoxae of the studied taxa are usually transverse, and at least partially, plate-like. The ‘dorsal plate’ of the metacoxae is generally large and well-developed in Mycetoporinae, Tachyporini, and Vatesini, but they tend to be rather small in size among the members of the Deropini + Tachinusini clade.
100. Metacoxae, ventral lamellae (or plates, vlmtc) on metacoxae (modified from Ashe [
28]: character 73) (ventral view): (0) absent (Figures 28H and 70A,C); (1) present, lateral areas, only weakly lamellate, not projecting posteriorly, limited to mesial margins; (2) present, lateral areas, only weakly lamellate, projecting posteriorly, but not developed along mesial margins; (3) present, mesial inner areas, moderately to strongly lamellate, projecting posteriorly, as well as weakly to moderately developed lamellae along lateral margins, partially concealing metatrochanters (Figures 12B, 18E, 28E, 29B, 46B, 54B and 60E); (4) present, very widely expanded, markedly developed.
See morphological definition of
Supplementary Figure S2B. Three of these character states (100-1, 100-2, 100-4) only occur in outgroup taxa (see
Supplementary Figure S3). The metacoxa of
Olisthaerus substriatus Paykull, 1790 [
85] (Olisthaerinae) is shown in Cai et al. ([
209]: Figure 3I). Compared to the transversely developed ‘dorsal plate’, the ‘ventral plate’ is found near the inner area of each metacoxae, usually partially concealing the metatrochanters. Several morphological variations of the ventral plates were found as listed above. Mycetoporinae lacks the ‘ventral plates’ on the metacoxae, whereas all studied taxa of Tachyporinae, except part of
Vatesus (Vatesini), have developed lamellae on each metacoxa. These lamellae project posteriorly and are weakly to moderately expanded along the outer margins of the metacoxae in Tachyporinae, but such lamellae are completely absent in
Vatesus gigas.
101. Metatibia, metatibial spurs at apex, length in comparison with metatarsomere 1 (dorsal and ventral views): (0) short, less than 2/5 of metatarsomere 1 (Figures 17D and 45F); (1) long, clearly longer than 2/5 of metatarsomere 1 (Figures 28E, 29C, 54I, 60C, 66A and 69C), but shorter than metatarsomere 1; (2) long, much longer than metatarsomere 1; (3) absent.
Mycetoporinae were scored as either ‘0′ or ‘1′. The latter may be a common state in this subfamily. Regarding Tachyporinae, this character is relatively stable within certain tribes. Most members of Tachyporini and all Deropini were coded as ‘0′. Compared to these tribes, Vatesini and Tachinusini were scored as ‘1′ in this study.
102. Tarsal formula (modified from Ashe [
28]: character 50): (0) 5-5-5; (1) 3-3-3.
Both Mycetoporinae and Tachyporinae have a typical 5-5-5 tarsal formula, but
Symmixus (Tachyporina) appears to have 4-segmented tarsi because of the shortened fourth tarsomeres [
22].
103. Metatarsus, length (dorsal and ventral views): (0) long, more than half length of metatibia, but shorter than whole length of metatibia (Figures 7E, 25C, 28E, 29B, and 52C); (1) markedly long, more than whole length of metatibia (Figures 15D and 66A); (2) short, shorter than half length of metatibia (Figure 44C).
The metatarsus of Mycetoporinae is markedly long, exceeding the entire length of metatibia. Within Tachyporinae, such long metatarsi were found only in Euconosomatina. In contrast, some taxa in the clade Deropini + Tachinusini have much shorter metatarsi and were therefore scored as ‘2′.
104. Metatarsomere 3, structure: (0) simple, without ventrally modified projection (Figure 60C); (1) complex, with ventrally modified projection.
Symmixus (Tachyporina) has a ventrally modified projection on metatarsomere 3 ([
22]: Figure 18).
105. Abdomen (dorsal view): (0) largely exposed dorsally; (1) moderately exposed dorsally; (2) slightly exposed, largely concealed by elytra.
Within Tachyporinae, only Leucotachinus (Tachinusini) has the abdomen largely concealed by long elytra.
106. Abdominal segments V–VI, including paratergites, posterolateral margins (dorsal and ventral views): (0) normal, not extending posteriorly (Figures 7E, 15D and 44C); (1) pointed, slightly but clearly extending posteriorly (Figures 25A,B and 28F); (2) not determined (paratergites absent, not fused with tergite and sternite, separated by sutures); (3) not determined (paratergites absent, completely fused with their tergite and sternite, forming ring-like abdominal segments).
The posterolateral margins of the abdominal segments V–VI are slightly but clearly extended posteriorly in most Vatesini, as well as in Lamprinus (Tachyporina) and Nepaliodes (Tachinusini).
107. Intersegmental membranes, between abdominal segments III–VI, pattern [dorsal and ventral views]: (0) without ‘brick-wall’ pattern (Figure 12C); (1) with deformed, or somewhat irregular ‘brick-wall’ patterns; (2) with transverse, somewhat rectangular ‘brick-wall’ patterns (Figure 71A,B).
In most outgroup taxa and Mycetoporinae, there is a clear ‘brick-wall’ pattern (sensu Newton et al. [
16]) found in the intersegmental membranes between abdominal segments III–VI. However, I have never seen such patterns in Tachyporinae based on the examined material.
108. Abdominal tergite III (dorsal view): (0) well sclerotized; (1) weakly sclerotized or membranous.
The weakly sclerotized abdominal tergite III is considered to be one of the 13 possible unique synapomorphies of Silphidae [
205].
109. Abdominal tergites IV–V, macrosetae (dorsal view): (0) absent (Figures 25A and 44C); (1) present, only on posterolateral edges of each tergite (Figures 7E and 15C); (2) present on middle and posterolateral edges of each tergite (Figures 66A, 71B).
Most members of Mycetoporinae have several macrosetae both on the middle of abdominal tergites IV–V and their posterolateral edges. In Tachyporini, these macrosetae are located only on the posterolateral edges of each tergite. Meanwhile, they are completely absent in the rest of Tachyporinae, except for Nitidotachinus (Tachinusini), Leucotachinus (Tachinusini), Tachinoporus (Vatesini), and Vatesus berghoffae (Vatesini).
110. Abdominal tergites IV–VI in posterior half (dorsal view): (0) lacking setigerous V-shaped punctures (Figures 7E, 25A, and 44C); (1) covered with setigerous V-shaped punctures (Figures 66A and 71B).
Most mycetoporines have setigerous V-shaped punctures covering abdominal tergites IV–VI, at least in the posterior half of each tergite. This type of pore is not common in Tachyporinae, and it is only seen in a few genera of Vatesini and Tachinusini.
111. Abdominal tergite IV, width (dorsal view): (0) weakly to moderately transverse, less than 2.5 times of its length; (1) strongly transverse, more than 2.5 times of its length.
The maximum width of the tergite IV in Vatesini and Tachinusini is more than 2.5 times wider than its length. However, the remaining Tachyporinae and Mycetoporinae each have a narrower, less transverse, tergite IV.
112. Abdominal tergites VIII–IX (Ashe [
28]: character 105): (0) not incorporated into genital apparatus; (1) incorporated into genital apparatus.
The abdominal tergites VIII–IX are incorporated into the genital apparatus in Habrocerinae.
113. Abdominal tergite VIII (male/female), posterior half (excluding macrosetae) [dorsal view]: (0) covered with minute, vestigial microsetae, lacking V-shaped punctures (Figure 31D); (1) covered with minute, vestigial microsetae in V-shaped punctures; (2) covered with short to moderately long setae, lacking V-shaped punctures (Figures 12E, 19C, 31C, 44C, and 71C); (3) covered with short to long setae in V-shaped punctures; (4) covered with short, but strong, thick setae, lacking V-shaped punctures; (5) absent, glabrous (Figure 55C).
In Mycetoporinae, tergite VIII is covered with V-shaped punctures, each usually bearing a short to long seta. However, some mycetoporines lack such V-shaped punctures. In contrast, there are several variations found in Tachyporinae. For example, Tachyporini is generally covered with vestigial to moderately long setae, lacking V-shaped punctures. In Vatesini, tergite VIII is usually covered with minute, vestigial microsetae, lacking V-shaped punctures, with the exceptions of Coprotachinus, Mimocyptus, and part of Vatesus. A similar pattern was found in Tachinusini, which are covered with the minute, vestigial microsetae, lacking V-shaped punctures, with the exceptions of Nitidotachinus, Leucotachinus, Peitawopsis, and Tachinomorphus. Within the Deropini-Tachinusini clade, Deropini is the only lineage having short to moderately long setae, lacking V-shaped punctures. It should also be noted that the ground setae of Megarthropsis (Tachinusini) are almost lacking. Because of the male abdominal modification, only the female of Nomimocerus (Habrocerinae) was considered for this character.
114. Abdominal tergite VIII (male/female), macrosetae (dorsal view): (0) with several or more macrosetae (Figures 19C and 71C); (1) with only a few to several distinct, thick, and long macrosetae (excluding sensory setae, see Yamamoto & Maruyama [
210]: Figure 14; occasionally covered together with a few to several small setae) (Figures 31A, 44C and 55C); (2) macrosetae absent (Figure 31B).
Mycetoporinae and Tachyporini have several or more macrosetae on tergite VIII, although these macrosetae in Mycetoporinae are rather difficult to distinguish from the other ground setae. The tachyporine tribes Vatesini, Deropini, and Tachinusini are mostly scored as ‘1′, with the exceptions of Vatesus gigas (Vatesini) and Nitidotachinus (Tachinusini). Because of the abdominal modifications, only the female of Nomimocerus (Habrocerinae) was considered for this character. Silpha (Silphidae) was coded here as ‘2′, as I considered them absent because they are too thin to be considered macrosetae.
115. Abdominal tergite VIII (male/female), longest seta, length (dorsal view): (0) shorter than tergite VIII (Figures 12E and 31A); (1) much longer than tergite VIII.
In Cileoporus (Vatesini), the longest seta on tergite VIII exceeds the entire length of tergite VIII. Tachinoporus (Vatesini) also have long macrosetae on tergite VIII (Figure 39A, D).
116. Abdominal tergite VIII, male, posterior margin (dorsal view): (0) truncate, rounded, or weakly pointed (Figure 70C); (1) lobes developed (Figures 31B and 55C); (2) not confirmed due to highly modified structures of genital segments.
Simple structures were generally found from the male abdominal tergite VIII in the selected outgroups and Mycetoporinae, as they lack developed lobes along the posterior margins of this segment. Contrary to these examples, two tachyporine tribes, namely Vatesini and Tachinusini, have more or less developed lobes. The remaining tachyporine tribes, i.e., Tachyporini and Deropini, have a simple posterior margin of tergite VIII as in most outgroups. In Austrotachinus (Tachinusini), the apex of tergite VIII is emarginate, but has a pair of weakly developed ‘lobes’ laterally.
117. Abdominal tergite VIII, female, posterior margin (dorsal view): (0) truncate, rounded, or weakly emarginate medially; (1) pointed medially; (2) deeply notched or emarginate medially; (3) lobes present (Figures 12E, 19C, 31A and 44C).
The posterior margin of the female tergite VIII is simple in the outgroups as well as Mycetoporinae, whereas Tachyporinae, except for part of Deropini, has more or less developed lobes along the posterior margin of this segment.
118. Abdominal tergite IX, male, ventral struts (modified from Ashe [
28]: character 71) (dorsal and ventral views): (0) absent (Figures 20F and 33A); (1) present (Figure 72C); (2) not confirmed due to highly modified structures of genital segments.
The ventral struts here refer to the structure of anteriorly directed extra extensions of male tergite IX as figured in Klimaszewski et al. ([
196]: Figure 3.5). In most cases, the presence of ventral struts is tightly associated with the following character 119 (i.e., 119-0), but they are different characters because I here considered extra extensions only and did not consider medial connection of the anteroventral margins of the halves of tergite IX. In some outgroups and Mycetoporinae, a pair of ventral struts were confirmed for tergite IX, although they are only weakly developed in two mycetoporines,
Bryoporus rufescens and
Carphacis. This structure is uniformly absent in all tribes of Tachyporinae.
119. Abdominal tergite IX, male, base of ventral side (dorsal and ventral views): (0) divided by sternite IX (Figure 72C); (1) basally fused, not divided by sternite IX (Figures 20F, 33D and 61E); (2) not confirmed due to highly modified structures of genital segments.
The ventral side of the base of tergite IX is divided by sternite IX in most outgroups and Mycetoporinae. However, it is fused and not divided in nearly all taxa of Tachyporinae. In Deropini, the ventral lobes of tergite IX are not fused as figured in Smetana ([
23]: Figure 4).
120. Abdominal tergite IX, male, structure of dorsal side in basal 1/3 [dorsal view]: (0) longitudinally contiguous (Figures 20A and 33C); (1) divided apically, each side divergent (Figures 33A, 47F and 55E,F); (2) completely continuous (Figure 72A,B); (3) lobes widely separated from each other at base, each lobe directed inward toward apex, with straight sides; (4) not confirmed due to highly modified structures of genital segments.
Several configurations were found in the basal 1/3 of male tergite IX. It is completely continuous, plate-like, in Mycetoporinae, whereas two types of variations were observed in Tachyporinae: the basal 1/3 of male tergite IX is longitudinally contiguous in Tachyporini and Mimocyptus (Vatesini), whereas it is divided posteriorly in the basal area, with sides diverging in Vatesini (except Mimocyptus), Deropini, and Tachinusini.
121. Abdominal tergite IX, male, structure of dorsal side from basal 1/3 to basal half (dorsal view): (0) longitudinally contiguous (Figure 20A); (1) separated from each other (Figures 33A,C and 55E,F); (2) completely continuous (Figure 72A,B); (3) not confirmed due to highly modified structures of genital segments.
A generally similar result was obtained with the above character (character 120), although it is slightly different.
122. Abdominal tergite IX, male, each apical form [dorsal and lateral views]: (0) short, rounded to pointed apically (Figure 72C); (1) more elongate, with elongate apex (Figures 20A and 33D); (2) elongate, much more slender, forming a lobe, sharply pointed at apex (Figures 33B,C,55G and 61E); (3) elongate, markedly slender, forming two distinct lobes, each sharply pointed at apex (Figures 33A and 55F); (4) not confirmed due to highly modified structures of genital segments.
In Mycetoporinae, the apices of male tergite IX are short and pointed apically. In contrast, a total of three forms listed above were found in Tachyporinae. A pair of elongate apices on this segment were found in Tachyporini and Deropini, but all three types were found in Vatesini, here coded from ‘1′ to ‘3′. The crown group of Vatesini has elongate, lobe-like apices, whereas they are merely elongate and less pointed in the earliest diverging lineage (Cileoporus). Interestingly, the vatesine genus Cilea has two distinct lobes on each lateral half of the segment. A similar state was found in Tachinusini, and only Olophrinus has two pairs of developed lobes as in Cilea.
123. Abdominal tergite IX, male, each apical area, macrosetae (dorsal and lateral views): (0) with several or more macrosetae, uniformly covering apical areas (Figures 20A and 72A,C); (1) with numerous macrosetae, restricted only to near and/or near apex; (2) with only single to several prominent macrosetae, sometimes together with a few small setae, restricted to apex (Figures 33C, 47F, 55E,F, and 61E); (3) distinct macrosetae absent; (4) with only a few setae, but not located on apex; (5) not confirmed due to highly modified structures of genital segments.
In Mycetoporinae and Tachyporini, each apical part of the male tergite IX has several or more macrosetae, uniformly covering apical areas. However, Vatesini, Deropini, and Tachinusini have only a single to several prominent macrosetae, restricted to the apex, although they are sometimes associated with a few to several small setae. Notably, the setose male tergite IX was only found in a few tachinusine genera.
124. Paratergites on abdominal segments V–VI (modified from Ashe [
28]: character 62) (dorsal and lateral views): (0) single pair (Figures 7E, 25A and 44C); (1) two pairs (Figures 66A and 71B); (2) zero (absent), but abdominal segments are not fused (Figure 15D); (3) zero (absent), abdominal segments are completely fused, ring-like.
Members of Mycetoporinae have two pairs of well-developed paratergites on abdominal segments III–VII. Compared to mycetoporines, there were significant variations found within Tachyporinae. In general, Tachyporinae have only a single pair of paratergites for these segments, but occasionally have two pairs (‘lower’ group of Vatesini) or none (only in Euconosomatina, see also Discussion).
125. Abdominal sternite III, longitudinal median basal carina (ventral view): (0) absent (Figure 12C); (1) present (Figures 19B and 71A).
There is a longitudinal median carina found in the middle of sternite III in Mycetoporinae, Euconosomatina, Deropini, and some members of Tachinusini.
126. Abdominal sternites IV–V, punctures (ventral view): (0) absent to fine, inconspicuous (Figure 25C); (1) coarse, conspicuous, semicircular, pit-like; (2) coarse, conspicuous, V-shaped (Figure 66B).
V-shaped punctures were found in Mycetoporinae, some outgroups, and only limited members of Tachyporinae (Vatesini and Tachinusini).
127. Abdominal sternite VI, male, peg-like setae, characteristically arranged along posteromedial margin (ventral view): (0) absent; (1) present (Figure 52D).
Only two studied taxa of Tachinusini have characteristically arranged peg-like setae along the posteromedial margins of sternite VI in the male.
128. Abdominal sternite VII (male/female), basomedial margin (observable when dissected) (ventral view): (0) without a broad, semicircular protrusion; (1) with a broad, semicircular protrusion (Figure 47A).
In Deropini, there is a peculiar semicircular protrusion located along the basomedial margin of sternite VII in both sexes (Figure 47A; Smetana [
23]: Figure 1; Naomi [
43]: Figure 3B; Zhao & Li [
106]: Figure 2A). A similar, or more pointed, protrusion is occasionally found along each basolateral margin of the sternite VII in Tachinusini (Figure 61A; e.g., Herman [
25]: Figure 110), but I am not scoring them here because of the significant variation.
129. Abdominal sternite VII, male, posteromedial margin, emargination [ventral view]: (0) not emarginate (Figure 66B); (1) only weakly emarginate (Figure 55A); (2) moderately to strongly emarginate (Figures 44B, 47A, 52D, and 61A); (3) deeply emarginate, with a pair of large triangular lateral lobes.
The members of Tachinusini have an emargination along the posteromedial margin of male sternite VII. Additionally, a similar emargination was found in Cilea limbifera (Vatesini) and two Mycetoporinae, namely Bolitobius and Ischnosoma. Interestingly, in Cilea silphoides (Linnaeus) sternite VII forms two halves of the lateral lobes, resulting in a deep emargination between them. The conditions of C. silphoides and Nomimocerus (Habrocerinae) are probably not homologous.
130. Abdominal sternite VII, male, posteromedial margin, concavity (ventral view): (0) not concave (Figure 66B); (1) very weakly concave (Figure 55A); (2) moderately to strongly concave (Figures 44B, 47A, 52D and 61A).
The posteromedial margin of sternite VII in the male is usually moderately to strongly concave in Tachinusini, although only weakly concave in some members. Similar concavities were found in a few taxa of Mycetoporinae. However, they are probably not homologous to those of Tachyporinae.
131. Abdominal sternite VII, male, short stout or peg-like setae, characteristically arranged along posteromedial margin [ventral view]: (0) absent (Figures 25C, 66B); (1) present, more than three to dozens of stout setae (not modified as peg-like setae), arranged (Figure 55A); (2) present, more than several to dozens of peg-like setae, arranged, directed posteriorly (Figures 44B, 47A, 52D and 61A).
Most taxa of Tachinusini have several to dozens of characteristically arranged peg-like setae (sensu Hammond [
211]) along the posteromedial margin of male sternite VII. In some tachinusines, there are not well-developed into peg-like setae, i.e.,
Pseudotachinus and
Tachinomorphus.
132. Abdominal sternite VIII (male/female), posterior half, uniform setae (excluding macrosetae) (ventral view): (0) covered with minute, vestigial microsetae, lacking V-shaped punctures (Figures 32A, 55D, and 61B,C); (1) covered with minute, vestigial microsetae in V-shaped punctures (Figure 55B); (2) covered with short to moderately long setae, lacking V-shaped punctures (Figures 12D, 19D,E, 32E, and 71D); (3) covered with short to long setae in V-shaped punctures; (4) absent, glabrous.
In Mycetoporinae, tergite VIII is covered with short to long setae, with or without V-shaped punctures. There are several variations found in Tachyporinae. For example, Tachyporini is generally covered with short to moderately long setae, lacking V-shaped punctures. In Vatesini, tergite VIII is usually covered with the minute, vestigial microsetae, lacking V-shaped punctures, with the exceptions of Coprotachinus, Mimocyptus, and Vatesus. A similar pattern was found in Tachinusini, which are covered with minute, vestigial microsetae, generally lacking V-shaped punctures, with some exceptions. Within the Deropini + Tachinusini clade, Deropini is the only tribe having moderately long ground setae, lacking V-shaped punctures. Because of the abdominal modification, only the female of Nomimocerus (Habrocerinae) was considered for this character.
133. Abdominal sternite VIII (male/female), macrosetae (ventral view): (0) with several or more macrosetae (Figures 12D, 19D,E, 55D and 71D); (1) with only a few to several distinct, thick, and long macrosetae (excluding sensory setae, see Yamamoto & Maruyama [
210]: Figure 14; occasionally covered with a few to several small setae) (Figures 32A, 47A, 55B and 61B,C); (2) macrosetae absent (Figure 32D).
Mycetoporinae and Tachyporini have several or more macrosetae on tergite VIII of both sexes, although these macrosetae in Mycetoporinae are rather difficult to distinguish from the ground setation. Within a clade comprised of the three tribes, namely Vatesini, Deropini, and Tachinusini, this character was scored as ‘1′, with the exceptions of Vatesus gigas (Vatesini) and Nitidotachinus (Tachinusini). Because of the abdominal modification, only the female was considered for Nomimocerus (Habrocerinae). Silpha (Silphidae) was here coded as ‘2′, as I considered them absent because they are too thin to be considered macrosetae.
134. Abdominal sternite VIII, male, characteristically arranged microsetae or peg-like setae on postero-medial area (excluding macrosetae) (dorsal view): (0) absent (Figure 71E); (1) present, only at apex (Figure 71F); (2) present, numerous, longitudinally distributed (Figures 52D, 66B and 71G).
In some genera of Mycetoporinae, some characteristically arranged microsetae, or peg-like setae, are found on sternite VIII of the male. They are either restricted to the apex (Bobitobus and Bryoporus) or cover a much wider area (Bolitobius, Bolitopunctus, Carphacis, and Ischnosoma). Within Tachyporinae, only Nitidotachinus (only one species, N. tachyporoides) has such clusters of microsetae along inner margins of the deep emargination of sternite VIII.
135. Abdominal sternite VIII, male, posterior margin (ventral view): (0) truncate, rounded, only weakly emarginate, or weakly pointed (Figure 71E); (1) rather narrowly, moderately emarginate medially (Figure 71H); (2) rather widely deeply emarginate medially (Figures 19E and 32D); (3) much more deeply incised or very strongly emarginate medially (Figure 47B); (4) rather widely deeply emarginate medially, with a few feebly developed lobes laterally (Figures 32E and 55B); (5) lobes developed (Figures 32D and 61B); (6) not confirmed due to highly modified structures of genital segments.
In general, a series of simple structures were found in male sternite VIII within the selected outgroups and most members of Mycetoporinae which lack well-developed lobes along the posterior margins of male sternite VIII. However, the apex is generally more or less modified into lobes in Tachyporinae, with several variations as listed above. In Tachyporini and Vatesus (Vatesini), they were uniformly deeply, rather widely emarginate medially. In Vatesini, three types of modifications were found, namely ‘2′, ‘4′, and ‘5′. All three species of Deropini unusually lack such lobe-like modifications, and are coded here as ‘3′. The most common state of Tachinusini is ‘5′ which have well-developed lobes. Within Tachinusini, there are a few feebly developed lobes posterolaterally in Nitidotachinus, Pseudotachinus, and Leucotachinus luteonitens, here scored as ‘4′.
136. Abdominal sternite VIII, male, central area (ventral view): (0) not distinctly deeply and narrowly incised (Figures 19E, 32D, 71E, H); (1) distinctly deeply and narrowly incised (Figure 32C,E and 61B); (2) distinctly deeply and relatively widely incised or emarginate (Figures 32C,E, 47B, 55B and 61B).
In some members of Vatesini, Deropini, and Tachinusini, sternite VIII in the male is deeply incised or emarginate.
137. Abdominal sternite VIII, female, posterior margin (ventral view): (0) truncate, rounded, weakly emarginate medially, or weakly pointed (Figures 19D and 71D); (1) deeply emarginate or incised medially; (2) nearly rounded, with only a pair of feebly developed projections (Figures 12D and 47E); (3) lobes present, including small internal lobes (Figures 32A, 55D, and 61C); (4) strongly pointed medially.
The posterior margin of female sternite VIII is simple in the outgroups and Mycetoporinae, whereas Tachyporinae have several variations. For example, in Tachyporini, it is merely truncate or rounded in Euconosomatina, whereas it is slightly different in Tachyporina, nearly rounded but generally having a pair of feebly developed posteriorly directed projections. In Tachyporina, a pair of similar projections was found in Deropini and Leucotachinus (Tachinusini) along the nearly rounded posterior margins. The rest of Tachinusini and all of Vatesini, have a few pairs of well-developed lobes.
138. Abdominal sternite VIII, female, arranged sensory setae along lobes or in alternative positions on posterior margin (ventral view): (0) prominent lobes absent, lacking rows of minute sensory setae (Figure 71D); (1) prominent lobes absent, with single row of minute sensory setae, distributed uniformly along posterior margin; (2) prominent lobes absent, with single row of minute sensory setae distributed along projections or in alternative positions on posterior margin, but limited to median area of posterior margin (Figures 12D, 19D and 47E); (3) prominent lobes present, inner lobes with only a pair or three minute sensory setae around apex of each lobe (Figure 32B); (4) prominent lobes present, inner lobes with rows of more than three, minute sensory setae in fan-like arrangement, located in apical area of each lobe (Figures 55D and 61C).
In the aleocharine taxa used here, a row of the minute sensory setae (sensu Yamamoto & Maruyama [
210]: Figure 14) widely covers the posterior margins of sternite VIII in the female. Compared to the outgroups and Mycetoporinae, there are a few variations found in Tachyporinae. Two tribes, Tachyporini and Deropini, lack well-developed lobes, but a single row of minute sensory setae rather narrowly covers the middle area of the posterior margins. All members of Vatesini, except
Vatesus gigas, have pairs of well-developed lobes and their associated inner lobes, with only a pair, or at most three, minute sensory setae located around the apex of each lobe. Tachinusini has a similar character state to Vatesini, but generally has more than three setae at the apex of each inner lobe. The condition of
Austrotachinus (Tachinusini) is rather difficult to interpret and was tentatively scored here as ‘4′ because it has a row of dense minute setae (although it is located between lateral inner lobes, instead of near the apices of each inner lobe).
139. Abdominal genital segments in female, general shape (excluding exposed gonocoxites and styli) (dorsal and ventral views): (0) transverse to only weakly elongate (Figures 12G, 20E, 34B, 47C and 61D); (1) moderately to strongly elongate (Figures 34A and 73B).
The outgroups and Mycetoporinae generally have moderately to strongly elongate genital segments in the female. In contrast, they are transverse to only weakly elongate in almost all members of Tachyporinae with the exception of Vatesus (Vatesini). A vatesine species, Coproporus laevis, was considered here as having narrowly elongate female genital segments.
140. Abdominal genital segments in female, basal part [dorsal and ventral views]: (0) well sclerotized (Figure 73B); (1) only weakly sclerotized and/or partially membranous (Figures 12G, 20E, 34B, 47C, and 61D).
In Mycetoporinae, the basal part of the female genital segments is well sclerotized. However, in Tachyporinae, they are only weakly sclerotized and even partially membranous in some cases, with only a handful of exceptions in this study (i.e., Vatesus in Vatesini and Leucotachinus luteonitens in Tachinusini).
Male:
141. Internal sac, spines (modified from Ashe [
28]: character 77): (0) absent (Figures 12F, 20B,C, 33E, 56A and 72E); (1) few or insignificant; (2) many and prominent.
In Trichophyinae and Habrocerinae, the internal sac includes numerous, prominent large spines. Some outgroups also have a few or insignificant spines on the internal sac. The vatesine genus Tachinoproporus has such prominent spines (Figure 42B, dsp) although this structure needs further observation.
142. Parameres, mesial area, degree of attachment to median lobe (modified from Jenkins Shaw et al. [
212]: character 67) (lateral and parameral views): (0) paramere(s) well separated from median lobe (Figures 66C, 72F); (1) parameres very closely appressed to median lobe (Figures 33E,G, 56C,D and 61F); (2) parameres absent; (3) not confirmed because of highly modified structures of genitalia.
All examined taxa of Tachyporinae in this study have a pair of parameres which are closely appressed to the median lobe of the aedeagus for nearly its whole length. In Mycetoporinae and the outgroups, they are generally well separated from the median lobe. The configuration and structure of the male genitalia of Habrocerinae are complicated, and the presence of parameres has not been confirmed in that subfamily [
192].
143. Parameres, apical inner margins (parameral view): (0) widely separated from each other (Figures 66C and 72F); (1) moderately separated (Figures 33H and 56A); (2) only narrowly separated longitudinally (Figures 12F, 20B and 33F); (3) contiguous longitudinally, nearly contiguous, or nearly fused (Figures 20C, 47F, 56B,D, and 61F); (4) completely fused, forming single structure; (5) parameres absent; (6) not confirmed due to highly modified structures of genitalia.
All members of Mycetoporinae have a pair of widely separated parameres, however, those of Tachyporinae are much more narrowly separated, with three variations from ‘1′ to ‘3′.
144. Parameres, apical half, structure (lateral and parameral views): (0) simple (Figures 20B, 33F, and 72D); (1) complex, modified; (2) very complex, multi-articulated; (3) parameres absent; (4) not confirmed due to highly modified structures of genitalia.
The general structures of the parameres in Mycetoporinae and Tachyporinae are exclusively simple, ranging from clavate to plate-like. Multi-articulated, complex parameres can only be found in Aleocharinae.
145. Parameres, shape (ventral view): (0) not flattened or plate-like ventrally (Figures 12F, 20B, 33F, 56A and 72F); (1) widely or rather widely flattened or plate-like ventrally (Figures 47F, 56B and 61F); (2) parameres absent; (3) not confirmed due to highly modified structures of genitalia.
Within Tachyporinae, the general shape of parameres is relatively diverse with several variations, though flattened or plate-like parameres can be found uniformly in the Deropini + Tachinusini clade (except Leucotachinus luteonitens in Tachinusini). The mycetoporine genus Parabolitobius has a pair of apically flattened and enlarged parameres, but it was coded here as ‘0′ based on general overall shape and structure. In Tachinomorphus (Tachinusini), parameres are flattened but fused longitudinally in basal area.
146. Parameres, velum (modified from Ashe [
28]: character 82): (0) absent (Figure 72E); (1) present; (2) parameres absent; (3) not confirmed due to highly modified structures of genitalia.
Aleocharinae has a velum on each paramere [
190], but it is completely absent, or not confirmed, in the other staphylinid groups, including Mycetoporinae and Tachyporinae.
147. Parameres, number of setae on each paramere (excluding minute sensilla): (0) zero (absent) (Figures 12F, 20B, and 33F,H); (1) present, less than five (Figure 72D); (2) present, five or more (Figure 72E); (3) parameres absent; (4) not confirmed due to highly modified structures of genitalia.
All members of Tachyporinae lack setae on each paramere, however, Mycetoporinae have a great variation in number, length, and distribution of such setae.
148. Parameres, minute sensilla or filiform setulae: (0) absent (Figures 12F, 20B, 33F and 56B); (1) present, a few to several (Figure 72D); (2) present, numerous (Figure 56C); (3) parameres absent; (4) not confirmed due to highly modified structures of genitalia.
Within Tachyporinae studied here, Tachinomorphus (Tachinusini) was the only taxon that has numerous filiform setulae on each paramere. The rest of the Tachyporinae lack these setulae. More than half of the taxa of Mycetoporinae have such minute sensilla, but they are insignificant in most cases, with only a few to several sensilla or setulae.
149. Parameres, setae, arrangement (excluding minute sensilla): (0) absent; (1) present, lacking a row of parameral setae; (2) present, each with a row of parameral setae at least partially aligned (Figure 72D); (3) parameres absent; (4) not confirmed due to highly modified structures of genitalia.
In the outgroups, parameral setae do not form a row, but alternatively are scattered on the apex of each paramere. However, Mycetoporinae have a distinct, longitudinal row of these setae, at least partially aligned.
150. Parameres, setae, location (excluding minute sensilla): (0) absent; (1) present, limited only to apex of paramere (Figure 72F); (2) present, not limited to apex of paramere (Figure 72E); (3) parameres absent; (4) not confirmed due to highly modified structures of genitalia.
The parameral setae are located in the apices of the parameres in the outgroups. In Mycetoporinae, they are either limited to the apices or are much more widely distributed on the parameres.
Female:
151. Spermatheca: (0) absent, or very weakly sclerotized, inconspicuous, oblong to cresent shaped (Figure 34A); (1) well sclerotized, comparatively complex or forming a distinctly complicated structure, usually associated with coils (Figures 56F, 61D and 73C).
In some Mycetoporinae and Tachyporinae (most taxa of Tachinusini), a well sclerotized spermatheca with a complex structure, such as basally nested coils, was found.
152. Gonocoxites I and/or II (modified from Ashe [
28]: character 89) (dorsal view): (0) present (Figure 47C); (1) present, but reduced to spinose process; (2) absent.
A single taxon used here, Oxypoda sp. (Aleocharinae: Oxypodini), does not have clearly developed, lobe-like gonocoxites. This condition is universal among the diverse members of the ‘higher’ group of Aleocharinae.
153. Gonocoxite II, shape (dorsal view): (0) slender, narrowly elongate (Figures 12G, 20E, 47D, 56E, 61D); (1) rather thick, more or less triangular (Figure 34A); (2) thick, narrowly elongate (Figures 34C and 73A); (3) thick, narrowly elongate, fused with gonocoxite I; (4) flattened, rectangular; (5) lobe-like gonocoxite II absent.
A thick and narrowly elongate gonocoxite II is uniformly found in Mycetoporinae. In Tachyporinae, it is generally much more slender, with the exception of most taxa of Vatesini and some Tachinusini. Two types of gonocoxite II were found in Vatesini: (i) rather thick, more or less triangular (Coproporus, Coprotachinus, Mimocyptus, Termitoplus, and Vatesus gigas); (ii) thick, narrowly elongate (Cilea and Vatesus praedatorius).
154. Gonocoxite II, size in comparison with gonostylus (dorsal view): (0) small (Figures 12G, 20D, 34A, 47C,D, and 56E); (1) large (Figure 73A); (2) lobe-like gonocoxite II absent.
The overall size of gonocoxite II in the outgroups, including Mycetoporinae, is usually large and prominent in comparison with the gonostylus. In contrast, they are much smaller in Tachyporinae.
155. Gonocoxite II (I and II, if they are fused together), setae [dorsal view]: (0) covered with normal to stout setae (Figure 73B); (1) covered with curved setae (strongly bent backwards) (Figures 12G, 20D, 47D, and 56E); (2) covered with both normal and curved setae (Figure 34A,C); (3) covered with long, strong setae; (4) covered with only a few scattered setae; (5) lobe-like gonocoxite II absent.
Gonocoxite II is covered with normal to stout setae in Mycetoporinae, but that of Tachyporinae are generally uniformly covered with curved setae. Although the presence of curved setae may be a universal condition in the entire subfamily (except Peitawopsis in this study), both normal and curved setae were found together on gonocoxite II in Vatesini.
156. Gonostylus (modified from Grebennikov & Newton [
2]: character 260) (dorsal view): (0) absent; (1) present, but minute and inconspicuous (Figure 73D); (2) present, large and conspicuous (Figures 12G, 20D, 34A, 47D, and 56E); (3) strongly sclerotized, relatively large, sharply pointed.
Mycetoporinae only has a pair of weakly developed, minute gonostyli, but those of Tachyporinae are much larger and conspicuous.