Appendix 1. Phylogenetically relevant characters
1.
Shape of the posterior region of the head capsule: (0) rounded; (1) flattened behind the
compound eyes. The posterior region of the head of Culicoides (Ceratopogonidae) [359], Chironomus
(Chironomidae) [41], Chaoborus (Chaoboridae), Androprosopa (Thaumaleidae), Mischoderus [58],
Protoplasa (Tanyderidae) [94], Bibio (Bibionidae), Exechia (Mycetophilidae), Coboldia (Scatopsidae),
Psychoda (Psychodidae), Ptychoptera (Ptychopteridae), Deuterophlebia (Deuterophlebiidae), Edwardsina
(Blephariceridae) [28], Nymphomyia (Nymphomyiidae) [50] and members of Tipulomorpha [58] is
rounded. The head capsule is also rounded in Nannochorista (Nannochoristidae, Mecoptera) [80] and
Micropteryx (Micropterigidae, Lepidoptera) [171]. The head capsule in Culex, Aedes, Anopheles, Culiseta
(Culicidae) [59], Corethrella (Corethrellidae), Dixa (Dixidae) [41], Wilhelmia (Simuliidae) [95], Axymyia
(Axymyiidae) [31], Macrocera (Keroplatidae), Mayetiola (Cecidomyiidae) [30], Spathobdella (Sciaridae),
Sylvicola (Anisopodidae), Tabanus (Tabanidae) [98], Pachygaster (Stratiomyiidae), Stilpnogaster
(Asilidae), Hemipenthes, Bombylius (Bombyliidae) [60], Drosophila (Drosophilidae) [97], Eristalis
(Syrphidae) [59], Caurinus (Boreidae, Mecoptera) [81], Panorpa (Panorpidae, Mecoptera) [115], Merope
(Meropeidae, Mecoptera) [82] and Mengenilla (Mengenillidae, Strepsiptera) [84] is flattened behind the
large compound eyes. The head of Ctenocephalus (Siphonaptera) is not flattened behind the eyes [83].
2.
Head capsule densely covered with microtrichia: (0) present; (1) absent. The head of Tipula,
Limonia, Cylindrotoma, Trichocera, Culex, Aedes [58], Corethrella, Chaoborus, Androprosopa, Axymyia
[31], Exechia, Mayetiola [30], Coboldia, Spathobdella, Sylvicola, Psychoda, Nymphomyia [50], Edwardsina
[28], Stilpnogaster and Drosophila [97] is densely covered with microtrichia. In Deuterophlebia all parts
except the ventral side are covered with microtrichia [28]. In Ptychoptera all parts with the exception of the
vertex are covered with microtrichia. The head capsule in Bibio is covered with longer setae, but without
microtrichia. In Pachygaster the head capsule is covered with long setae, but no microtrichia are present.
After Beutel & Baum [80] the head capsule in Nannochorista is densely covered with thin and very short
setae (coaded as 0). Large parts of the head in Mengenilla and other Strepsiptera are densely covered
with microtrichia [84]. The head of Caurinus [81], Merope [82] and Ctenocephalus [83] is not covered with
microtrichia.
3.
Orientation of the head: (0) orthognathous; (1) prognathous. The head of Wilhelmia [95],
Culicoides [113], Chironomus (Chironomidae) [41], Deuterophlebia, [28], Edwardsina, Axymyia [31],
Macrocera, Mayetiola [30], Coboldia, Spathobdella, Psychoda, Tabanus [98], Stilpnogaster, Hemipenthes
[60], Drosophila [97], Eristalis [59], Caurinus [81], Nannochorista [80], Merope [82], Ctenocephalus [83]
and Micropteryx [171] is orthognathous. Whereas the orientation of the head is prognathous in Culex,
Aedes, Corethrella, Chaoborus, Androprosopa, Dixa [41], Mischoderus, Protoplasa [94], Bibio, Sylvicola,
Ptychoptera, Nymphomyia [50], Bombylius [60], Panorpa [115] and members of Tipulomorpha [58]. The
head in Mengenilla is subprognathous ([84] fig 1c) (coaded as 1).
4.
Frontal apodeme: (0) present; (1) absent. A frontal apodeme between the antennal bases is
absent in Deuterophlebia [28] and also missing in Nymphomyia [50], Chaoborus, Androprosopa, Tipula
(Tipulidae) [58], Pedicia, Bibio (Bibionidae), Axymyia [31], Coboldia, Sylvicola, Psychoda, Stilpnogaster,
1
Eristalis (Syrphidae) [59], Hemipenthes, Bombylius (Bombyliidae) [60], Drosophila [97], Ctenocephalus
(Siphonaptera) [83], and also in members of all groups of Mecoptera with the notable exception of
Nannochoristidae ([80]: fig. 5, fap, [115, 120]). It is also absent in members of Strepsiptera [84]. A small
apodeme-like structure is present in Trichocera (Trichoceridae) [58], Cylindrotoma and Pachygaster
dorsally between the antennal bases. It is distinctly developed in Edwardsina, Limonia (Tipulidae),
Macrocera,
Exechia,
Mayetiola
[30],
Mischoderus
(Tanyderidae),
Ptychoptera,
Culicoides
(Ceratopogonidae), [113], Corethrella, in representatives of Culicidae [59, 69, 99, 100], and in Wilhelmia
(Simuliidae) [95]. Bonhag [98] described a median inflection between the antennae (m) in Tabanus,
probably this is the frontal apodeme (coaded as 0).
5.
Frontoclypeal-/epistomal suture: (0) present as a strengthening ridge; (1) present as a joint;
(2) absent. Both regions are fused in Deuterophlebia, Edwardsina [28], Nymphomyia [50], Axymyia [31],
Spathobdella, Ptychoptera, Culicoides [359], Dixa [41], Simulium ([95]: fig. 1) and Ctenocephalus [83]. The
suture is present in Corethrella, Chaoborus, Androprosopa, Limonia, Cylindrotoma, Pedicia, Trichocera,
Bibio, Exechia, Mayetiola [30], Coboldia, Sylvicola, Psychoda, Eristalis [59], Bombylius [60], Tabanus [98],
Mischoderus and Protoplasa [94]. It is distinctly developed in Tipula paludosa [58], whereas it appears to
be absent in Tipula reesi [122] and Tipula sp. [202]. The suture is also present in members of Mecoptera
[80-82, 115, 120] and in Micropteryx [171]. A joint between the clypeus and frons in adults of Culicidae [8,
59, 99] is a potential autapomophy of the family. The frontoclypeal suture is absent in Strepsiptera, with
the exception of Protoxenos and Mengea [84]. In Drosophila the suture is present as a membrane (coded
as 0) [97].
6.
Clypeus: (0) subdivided into ante- and postclypeus; (1) undivided. A transverse suture
separating an anterior anteclypeus from the posterior postclypeus is present in the groundplan of Diptera
according to Crampton [126]. However, the division is absent in Deuterophlebia, Edwardsina [28] and
Nymphomyia [145], and is also almost generally missing in other groups [41, 50, 59, 60, 70, 98, 94, 95 97,
113]. A transverse clypeal furrow is present in adults of Culicidae (e.g. Culex, Culiseta, Aedes; 59, 99,
100], and also in Nannochorista [80] and Caurinus [81]. It is absent in other groups of Mecoptera [82, 115,
120] and also in Siphonaptera [83] and some other groups of endopterygote insects [80, 84, 171].
7.
Rostrum: (0) absent or very short; (1) distinctly developed. A distinctly developed rostrum is
present in adults of Tipula, Limonia [58], Cylindrotoma, Pedicia, Nymphomyia [50], Chaoborus, Dixa [41],
Edwardsina, Sylvicola [70], Mischoderus and Protoplasa [94]. It is very short in Corethrella, Androprosopa,
Trichocera [58], Bibio, Culicoides [113], Ptychoptera (Ptychopteridae) [70], in representatives of Culicidae
[59, 99, 100], and in Wilhelmia [95]. It is formed by a completely foldable rostral membrane in adults of
most groups of Brachycera ([59]: fig. 34, Eristalis [Rstr]; [60]: fig 5, Hemipenthes, Bombylius; [97]: fig. 1,
Drosophila; [70]: figs 62-65, 69, 72, Symphoromyia, Rhagio, Sepsis, Fucellia) (see also next character). A
rostrum is completely absent in Deuterophlebia [28], Axymyia [31], Macrocera, Mayetiola [30], Coboldia,
Spathobdella, Psychoda, Stilpnogaster, Dioctria (Asilidae) [70], Chironomus [41], Tabanus [98],
Pachygaster, representatives of Mycetophilidae (Exechia, [70] Mycetophila, Mycomya), Nannochorista
2
[80], Caurinus [81], Ctenocephalus [83], in members of Strepsiptera [84] and in Micropteryx [171]. A
different type is present in adults of most groups Mecoptera [81, 82, 115, 120, 316].
8.
Sclerotisation of the rostrum: (0) only sclerotised dorsally; (1) sclerotised on all sides; (2)
completely membranous. The rostrum of Tipula, Cylindrotoma, Limnophila [70] and Nymphomyia [50] is
entirely sclerotised. It is formed by the ventrally inflected lateral parts of the clypeus. Only the dorsal side
is sclerotised in Limonia, Erioptera, Dicranomyia [70], Chaoborus, Dixa [41], Pedicia and Edwardsina, and
this is also the case in Mischoderus and Protoplasa [94]. It is completely membranous in Sylvicola [70]
and representatives of Brachychera ([97] Drosophila; [70] Symphoromyia, Rhagio, Sepsis, Fucellia; [59]
Eristalis; [60] Hemipenthes, Bombylius). In Merope the rostrum is covering the mouthparts and is not
sclerotized on all sides [82] (coaded as 0).
9.
Reduced number of antennomeres: (0) not reduced; (1) reduced (≤6). The antenna of female
Culicoides composed of 13 flagellomeres [359], in Chironomidae of 14 segments [286], in Wilhelmia 10
segments [95], in Ptychoptera of 15 segments and in Caurinus of 16 segments [81] and is not reduced.
The number of antennomeres is also not reduced in Mischoderus [58], Protoplasa (16) [94], Axymyia (17)
[31], Exechia (16), Mayetiola (16) [30], Coboldia (10), Spathobdella (13), Sylvicola (16), Bibio (9),
Psychoda (9), Corethrella (16), Chaoborus (15), Androprosopa (12), members of Culicidae [59, 69, 99,
100] and Tipulomorpha [58], and also in members of Mecoptera (Nannochorista [32], Merope [29]: [80,
82]) and in Micropteryx [171]. The antennae of Tabanus [98] and Pachygaster comprise 7 segments. The
antenna is 5-segmented in Nymphomyia [50] and 6-segmented in Deuterophlebia [28]. The number of
antennomeres is reduced in Stilpnogaster (5) and Drosophila (6) [97]. In the groundplan of Strepsiptera
the antenna is composed of 8 antennomeres, whereas only 6 antennomeres are present in Mengenilla
[84, ] (Pohl et al. 2005). The number of antennomeres is reduced in Hemipenthes and Bombylius [60]. The
antenna of Eristalis composes of scape, pedicil and the first flagellum with an arista [59]. The antenna of
Ctenocephalus is composed the scape, pedicil and a clave, which consists of 9 segments [83] (coaded as
0).
10.
Insertion of antennae: (0) frontally, not adjacent medially; (1) frontally, adjacent in midline;
(2) laterally, widely separated. The antennae insert on the dorsal side of the head and are widely
separated In Deuterophlebia [28], Bibio and Micropteryx [171]. They are also widely separated in Caurinus
[81] and Ctenocephalus [83]. They inserted frontally, but widely separated in Psychoda (coaded as 0) and
Chaoborus. The antenna insertions lie frontally between the compound eyes in Edwardsina, Nymphomyia
[50], Mischoderus, Tipula, Limonia [58], Pedicia, Axymyia [31], Macrocera, Exechia, Mayetiola [30],
Coboldia, Spathobdella, Sylvicola, Protoplasa [94], Ptychoptera, Culicoides [113], Corethrella, in
representatives of Culicidae [59, 69, 99, 100], Simuliidae [95], Tabanidae [98], Stilpnogaster,
Stratiomyiidae, Bombyliidae [41], Drosophila [97] and also in Nannochorista [80], Panorpa [115], Merope
[82] and Mengenilla [84]. This is also the case in Trichocera [58], Chironomus [41], Androprosopa, Dixa
[41], Cylindrotoma and Eristalis ([59] fig. 31), where they are medially adjacent. The antennae inserted
laterally, close to the anterior margin of the compound eyes in Protoxenos (Pohl et al. 2005).
3
11.
Shape of the antennae: (0) filiform; (1) moniliform; (2) club-shaped; (3) flabellat. The antenna
is filiform in the most adults of Diptera [28, 58-60, 94, 108, 113]. In Nannochorista [80], Panorpa [115] and
Caurinus [81] the antenna is also filiform. In Wilhelmia [95], Axymyia [31], Coboldia, Bibio and Micropteryx
[171] the antenna is moniliform and is club-shaped in Nymphomyia, Tabanus [98], Pachygaster,
Drosophila [97] and Eristalis [59]. In Mayetiola the antennae are bead-like in males (coaded as 0) [30].
The antenna of Mengenilla is flabellat [84]. In Ctenocephalus the last 9 segments together form a clubshaped structure [83] (coaded as 2).
12.
The first flagellomere of the antenna: (0) not enlarged; (1) enlarged. The first flagellomere of
the antenna is enlarged in Nymphomyia [145], Macrocera, Tabanus [98], Bombylius [60], Drosophila [97],
Eristalis [59] and Micropteryx [171]. The flagellomere is not enlarged in Deuterophlebia [28], Protoplasa
[94], Mischoderus, Ptychoptera, Tipula, Limonia, Cylindrotoma, Trichocera [58], Pedicia, Corethrella,
Chaoborus, Androprosopa, Dixa [41], in members of Culicidae [59, 99, 100], in Wilhelmia [95], Culicoides
[113], Bibio, Axymyia [31], Exechia, Mayetiola [30], Coboldia, Spathobdella, Sylvicola, Psychoda,
Stilpnogaster, Pachygaster, Hempenthes [60], Nannochorista [80], Panorpa [115], Caurinus [81],
Ctenocephalus [83] and Mengenilla [84].
13.
Pedicellus with Johnston’s organ: (0) absent; (1) present. The pedicellus contains a sense
organ in members of Culicidae [59, 99, 100], Corethrellidae, Chaoborus, Androprosopa, Ceratopogonidae,
Chironomidae [113] and Drosophila [111]. A Johnston’s organ is also present in Wilhelmia [95], Caurinus
[81], Ctenocephalus [83] and males of Mengenilla [84]. The sense organ is absent in Protoplasa [94],
Mischoderus, Ptychoptera, Axymyia [31], Mayetiola [30], Coboldia, Sylvicola, Deuterophlebia, Edwardsina
[28], Nymphomyia [94], members of Tipulomorpha [58], in Eristalis [59], Stilpnogaster, Pachygaster and
also in Nannochorista [80] and Merope [82]. A sense organ is present on the second flagellomer in Bibio,
but absent on the pedicellus.
14.
Last segment of the antenna: (0) not elongated; (1) elongated. The last antennal segment is
extremely elongated in males of Deuterophlebia [28, 272, 273] and also elongated in Nymphomyia ([145]
fig. 16, 26, 29). It is thick and elongated in Eristalis [59], Drosophila [97] and also enlarged in
Stilpnogaster, Pachygaster, and Bombylius [60]. The segment is not elongated in members of most other
groups of Diptera [30, 31, 58-60, 94, 95, 98, 99, 113], in Strepsiptera [84] and Merope [82]. This condition
is unknown in the rest of Mecoptera [80, 81].
15.
Ocelli: (0) present; (1) vestigial or absent. Ocelli are absent in adults of Deuterophlebia [28],
Tipula, Limonia, Cylindrotoma, Pedicia, Culex, Anopheles [58], Corethrella, Chaoborus, Androprosopa,
Dixa [41], Culiseta [59, 100], Wilhelmia [95], Mischoderus, Protoplasa [94], Ptychoptera, Macrocera,
Exechia, Mayetiola [30], Psychoda, Caurinus [81], Merope [82] and also in Ctenocephalus [83] and in
fossil and extant strepsipterans [84] (Pohl et al. 2005). Three ocelli are present on the vertex in
Edwardsina, Trichocera [58], Bibio, Axymyia [31], Coboldia, Spathobdella, Sylvicola, Stilpnogaster,
Pachygaster, Eristalis [59], Drosophila [97], Exoprosopa [41], Nannochorista [80] and Panorpa [115].
Christophers [99] described a pair of degenerated ocelli on the frons of Aedes. A pair is present
posterolaterad of the large compound eyes of Nymphomyia [50, 145]. Bonhag [98] described three
4
rudimentary ocelli on the vertex of Tabanus (coaded as 0). After Szadziewski et al. [359] Ocelli are absent
in Ceratopogonidae, whereas two are present on the frontal region in Culicoides after Gad [113]. Two
ocelli are present on the vertex in Micropteryx [171].
16.
Subdivision of the compound eyes in dorsal and ventral part: (0) undivided; (1) subdivided.
The compound eyes are undivided in the most dipteran groups [28, 30, 31, 50, 58-60, 94, 95, 97; 98, 108,
113] and also in Mecoptera (80-82, 115), Strepsiptera [84] and in Micropteryx [171]. The eyes are
subdivided in a dorsal and a ventral part in Axymyia.
17.
Coronal-/epicranial suture: (0) present; (1) absent. A coronal suture is absent in Deuterophlebia,
Edwardsina [28], Corethrella, Chaoborus, Androprosopa, Tipula, Limonia, Cylindrotoma, Pedicia,
Trichocera, Mischoderus [58], Protoplasa [94], Ptychoptera, Bibio, Axymyia [31], Exechia, Mayetiola [31],
Coboldia, Sylvicola, Psychoda, Wilhelmia [95], Tabanus [98], Pachygaster, Eristalis [59], Hemipenthes
and Bombylius [60]. It is also missing in Ctenocephalus [83], Caurinus [81] and many other
representatives of Mecoptera (120; 82). It is present in Culex, Culiseta ([59, 100] fig. 1), Aedes ([99] fig.
53/1), Chironomus ([41] fig. 12, e.s), Nymphomyia [50], Drosophila ([97] fig. 1C, premandibular suture),
and in Nannochorista [80]. In Micropterix the coronal suture is non-continuous [171]. The presence is
apparently a plesiomorphic condition preserved in Nymphomyiidae and Culicidae. A typical coronal suture
is present in Protoxenos, but strongly shortened in Mengenilla [84].
18.
Postgenal bridge: (0) present; (1) absent. A postgenal bridge is present in Deuterophlebia [28],
Nymphomyia [50], and also in Tipula [58], Limnophila [70], Aedes [99], Bibio, Axymyia [31], Stilpnogaster,
Tabanus [98], Eristalis [59], Exoprosopa [41], Rhagio [70], Ctenocephalus [83], and in representatives of
Mecoptera [e.g. 81, 82, 115, 120, 316] (with the possible exception of Nannochorista [80]). The ventral
closure of the head capsule is largely membranous in Limonia, but a narrow postgenal bridge is present
anterior to the foramen occipital (coaded as 0) [58]. The bridge is absent in Edwardsina, Erioptera,
Dicranomyia [70], Pedicia, Trichocera, Chironomus [41], Culicoides [113], Corethrella, Chaoborus,
Androprosopa, Dixa [41], Mischoderus [58], Protoplasa [94], Culiseta ([59]: fig. 4, gu, [100]), Ptychoptera,
Mycetophila, Mycomya [70], Macrocera, Exechia, Mayetiola [30], Coboldia, Sylvicola, Psychoda,
Wilhelmia [95], Symphoromyia [70] and Drosophila [97]. The head is largely membranous on its ventral
side in adults of these taxa. The ventral part of the head is closed by the lateral postgenal area and a
medial labial plate (mouthfield sclerite) in Mengenilla [84].
19.
Tentorium: (0) present; (1) absent. The tentorium is present as a more or less straight tube-like
rod in Deuterophlebia, Edwardsina [28] and many other groups of Diptera [e.g. 41, 59, 70, 95, 99, 100,
109, 113) ([94]: fig. 3). A similar condition is found in representatives of Mecoptera [80-82, 115, 120] and
Siphonaptera [83]. The tentorium is completely absent in Nymphomyia [50], Tipula [58] and in Strepsiptera
[84]. In Mayetiola [30], Cylindrotoma and Pedicia the tentorium is only a short vestigial tube (coaded as 0,
as muscles are attached). In Exechia anterior and posterior tentorial grooves are present, but the short
anterior and posterior tentorial arms are not connected with each other. A tentorium, incl. corpotentorium,
is also present in Micropteryx [171].
5
20.
Dorsal tentorial arm: (0) presen or present as a thin thread-like structure; (1) short vestigial
structure; (2) absent. The dorsal arm is completely missing in Deuterophlebia, Edwardsina [28],
Ptychoptera, Trichocera, Limonia [58], Pedicia, Cylindrotoma, Culex [109], Aedes [99], Culicoides [113],
Dixa [41], Androprosopa, Bibio, Macrocera, Exechia, Mayetiola [30], Coboldia, Spathobdella, Sylvicola,
Psychoda, Stilpnogaster, Drosophila [97], Ctenocephalus [83] and Micropteryx [171]. It is present as a
short vestige in Mischoderus, Culiseta ([59]: fig. 5, d.Ta, [100]: fig. 4), Corethrella, Chaoborus,
Chironomus ((41]: fig. 152, r.d.a.), Eristalis ([59]: figs 38, 39), Exoprosopa [41], Pachygaster and
Nannochorista ([80]: fig. 2d, dta). A very thin thread is present in Boreus according to V. Michelsen but not
identificable by Beutel & Baum ([80] see character 8). Wenk [95] described a tentorial ridge for Wilhelmia,
extending dorsad towards the antennal foramen ([95]: fig. 2). A typical, well developed dorsal arm is
apparently almost generally missing in Antliophora [80, 115, 120], but a thin, sclerotised structure is
present and connected to the head capsule in Caurinus ([81]: figs 5C, D) and Merope [82]. It is noteworthy
that dorsal arms are also present in Tabanus ([98]: fig. 5) (see char. 10).
21.
Shape of the anterior tentorial arm: (0) thick, approximately round in cross section, hollow;
(1) partly hollow; (2) massive. The tentorium of Deuterophlebia, Edwardsina [28], Mischoderus,
Ptychoptera, Trichocera [58], Bibio, Macrocera, Spathobdella, Sylvicola, Psychoda, Wilhelmia [95],
Culicoides [113], Corethrella, Chaoborus, Androprosopa, Stilpnogaster, Pachygaster, Tabanus [98];
Nannochorista and Bittacus [80] is a thick, hollow tube, and this is also the case in Culicidae [59, 99, 100,
109], where the lumen is wider anteriorly. A recognisable lumen is not present in Eristalis [59], Drosophila,
Exoprosopa [41], Mayetiola [30], Pedicia, Cylindrotoma, Boreus [80] and Caurinus [81]. The lumen of the
anterior part of the tentorium is narrow in Limonia. It widens at the level of the brain and the posterior
hollow part is approximately round in cross section [58]. In Coboldia the anterior part is also narrow and
the tentorial rod is hollow in the following part (coaded as 1). The vestigial anterior tentorial arm is round
and hollow at the beginning and norrow in the posterior region in Exechia. The anterior tentorial arms of
Ctenocephalus are missing [83].
22.
Frontotentorial muscle band: (0) absent; (1) present. A frontotentorial muscle band is absent in
all Diptera examined [28, 30, 31, 58, 59, 98, 108, 113] (with the exception of Drosophila ([111]: 19a) and
also in Caurinus [81], Boreus, Bittacus [80] and Ctenocephalus [83]. The muscle is present in
Nannochorista ([80]: fig. 5, ften).
23.
Labrum: (0) present; (1) absent. With the exception of Deuterophlebia [28] and Nymphomyia [50],
the labrum is present in all members of Diptera [e.g. 30, 31, 58, 59, 70, 94, 95, 98-100, 109], Mecoptera
[80-82, 115, 120], Siphonaptera [83] and Lepidoptera [171] examined. In Strepsiptera a well developed
labrum is present in Protoxenos, Mengea and Mengenilla, but missing in other strepsipterans (Pohl et al.
2005; [84].
24.
Clypeolabral connection: (0) separated; (1) fused. The clypeus and labrum are fused in
Edwardsina [28], and also in Ctenocephalus [83] and adults of most groups of Mecoptera [80, 115, 120]
and Strepsiptera [84]. They are recognisable as separate structures in other groups of Diptera [e.g. 30, 31,
6
58, 59, 70, 94, 95, 98, 99, 100, 109, 113], and also in Nannochorista [80], Caurinus [81], Merope [82] and
Micropteryx [171].
25.
Separation of clypeus and labrum: (0) transverse suture; (1) clypeus and labrum separated
by a membrane. Both elements are separated by an exposed membrane in Tipula, Limonia, Trichocera,
Mischoderus [58], Protoplasa [94], Ptychoptera, Bibio, Exechia, and Wilhelmia [95], whereas a suture and
an internalised membrane are present in adults of Culicidae [59, 99, 100, 109], in Culicoides [113],
Nannochorista [80], and in Caurinus [81]. A suture is also present in Dixa [41], Corethrella, Chaoborus,
Androprosopa, Pedicia, Cylindrotoma, Coboldia, Sylvicola, Psychoda and Micropteryx [171]. Clypeus and
labrum are separated by a rostral membrane in Eristalis [59], Drosophila [97] and Toxophora [70], and by
a small membranous area in Tabanus [98]. In Merope an indistinct clypeolabral suture is present [82].
26.
Fulcrum: (0) absent; (1) present. A fulcrum with lateral fulcral plates, which join the external
clypeal wall distally, is absent in Deuterophlebia, Edwardsina [28], and Nymphomyia [50], and is also
missing in Mischoderus, Ptychoptera, Tipula, Limonia, Trichocera [58], Pedicia, Cylindrotoma, Bibio,
Axymyia [31], Macrocera, Exechia, Mayetiola [30], Coboldia, Sylvicola, Wilhelmia [95], Corethrella,
Chaoborus, Androprosopa, Culicoides [113], Stilpnogaster, Pachygaster, Tabanus [98], Nannochorista
[80], Caurinus [81], Panorpa [115], Merope [82], Ctenocephalus [83], Mengenilla [84] and Micropteryx
[171]. It is generally present in adults of Culicidae [59, 99, 100, 109], and does also occur in Syrphidae
[59], Drosophila [97], Protoplasa [94] and Bombyliidae ([60] Hemipenthes, Bombylius).
27.
Labro-epipharyngeal channel: (0) present; (1) absent. The epipharynx forms a food channel in
most adults of Diptera examined, and also in Nannochorista and Siphonaptera [e.g. 31, 58-60, 80, 83, 95,
98, 107, 108, 113, 128], but is missing in representatives of Pistillifera, Boreidae [80, 81, 115] and also in
Strepsiptera [84]. The presence is a potential synapomorphy of Nannochoristidae, Siphonaptera and
Diptera. In Mayetiola and Macrocera the anterior epipharynx is lightly bent upwards, but formed not a food
channel [30]. In Exechia the food channel is completely missing.
28.
Shape of the labro-epiphayngeal food channel: (0) ventrally open; (1) closed by the sides of
the epipharynx; (2) ventrally closed by hypopharynx; (3) ventrally closed by the mandibles. The
food channel is open in Mischoderus, Ptychoptera, Limonia, Trichocera [58], Pedicia, Sylvicola, Psychoda,
Wilhelmia [95], Simulium [128], Corethrella, Chaoborus, Andropsosopa, Pachygaster, Nannochorista [80]
and Ctenocephalus [83]. It is very flat in Tipula ([58]: eph, figs 8C, D), Cylindrotoma, Bibio [80], Coboldia,
Spathobdella and Axymyia [31] and forms a closed tube in representatives of Culicidae [59, 99, 100, 107,
108] and in Stilpnogaster. In Eristalis [59] and representatives of Bombyliidae ([60] Hemipenthes,
Bombylius) the channel is ventrally closed by the hypopharynx, whereas it is ventrally closed by the
mandibles in Tabanus [98] and Culicoides [113].
29.
M. labroepipharyngalis (M. 7): (0) present; (1) absent. A paired M. labroepipharyngalis connects
the external and internal labral wall in Ptychoptera, Corethrella, Chaoborus, Androprosopa, Edwardsina
[28], Tipula, Limonia, Trichocera [58], Pedicia, Cylindrotoma, Bibio, Axymyia [31], Macrocera, Exechia,
Mayetiola [30], Spathobdella, Sylvicola, Psychoda, Tabanus [98], Eristalis [59], Hemipenthes, Bombylius
[60], Drosophila ([97] compressor muscle) ([111], 5) and Micropteryx [171]. It is absent in representatives
7
of Culicidae [59, 99, 100, 109] and Simuliidae [95], Coboldia, Pachygaster and also missing in
Nannochorista [80], Caurinus [81], Ctenocephalus [83] and Mengenilla [84]. It is present in adults of other
groups of Mecoptera [80, 82, 115, 120]. Whether M. labroepipharyngalis is unpaired in the groundplan of
Diptera as postulated by Gouin [124] appears questionable. Radial labroepipharyngeal muscles occur
secondarily in Cyclorrhapha according to this author.
30.
M. frontolabralis (M. 8): (0) present; (1) absent. Among all adult dipterans examined the muscle
is only absent in Deuterophlebia [28], Nymphomyia ([50] labrum reduced), Pachygaster and Drosophila
[111]. In Axymyia [31] and Cylindrotoma the muscle is not recocnisable in the µCT data set. In other
groups the muscle has an unusual origin on the clypeus (M. clypeolabralis) (e.g. Ptychoptera, Tipula,
Limonia, Trichocera, Pedicia, Bibio, Mayetiola [30], Macrocera, Exechia, Spathobdella, Psychoda,
Mischoderus, Culicidae, Corethrella, Chaoborus, Androprosopa, Tabanus, Hemipenthes, Bombylius; [5860, 95, 98-100, 109]. Schiemenz [59] interpreted the muscle he found in Culiseta and Eristalis as M.
epistomalabralis (M. 10). However, considering the function as labral levator it is much more plausible to
assume that it is homologous with M. frontolabralis. M. 8 is generally absent in representatives of
Mecoptera [e.g. 80, 81, 115, 120], in Siphonaptera ([83] Ctenocephalus) and in Mengenilla [84]. The
muscle is present in Micropteryx [171].
31.
Shape of M. frontolabralis (M. 8): (0) separated; (1) fused. M. clypeolabralis is separated in
Ptychoptera, Tipula, Limonia, Trichocera [58], Pedicia, Culicoides [113], Bibio, Spathobdella and
Micropteryx [171]. The paired components of the muscle are fused medially in Mischoderus, Macrocera,
Exechia, Mayetiola [30], Psychoda, Corethrella, Androprosopa, Chaoborus and members of Culicidae [59,
99, 100, 109] and in Tabanus [98] and Eristalis [59].
32.
M. frontoepipharyngalis (M. 9): (0) present; (1) absent. The muscle is absent in Edwardsina [28],
Mischoderus, Ptychoptera, Tipula, Limonia, Trichocera [58], Pedicia, Cylindrotoma, Axymyia [31],
Macrocera, Exechia, Mayetiola [30], Spathobdella, Psychoda, Culicoides [113], Corethrella, Chaoborus,
Androprosopa, Stilpnogaster, Pachygaster, Eristalis [59], Drosophila [111], Tabanus [98], and is also
lacking in representatives of Bombyliidae [60], Mecoptera [80-82, 115, 120], Siphonaptera [83] and
Strepsiptera [84]. It is present in Bibio, Coboldia, Wilhelmia [95], in adults of Culicidae [59, 99, 100, 109]
and in Micropteryx [171].
33.
Origin of M. tentorioscapalis anterior (M. 1): (0) tentorium; (1) head capsule. In Deuterophlebia
the muscle has multiple origins, three on the head capsule and one on the dorsal side of the tentorium
[28]. M1 has a widespread origin on the tentorium and also on the head capsule in Exechia. It originates
on the prefrontal region of the head capsule in Tipula, Limonia [58], Mayetiola [30], Nymphomyia [50] and
Pachygaster. In Axymyia the muscle originates on the gena. The muscle originates on the clypeus in
Ctenocephalus [83], and on the tentorium in Edwardsina, Mischoderus, Ptychoptera, Trichocera [58],
Pedicia, Cylindrotoma, Bibio, Macrocera, Coboldia, Spathobdella, Sylvicola, Wilhelmia [95], Culicoides
[113], Corethrella, Chaoborus, Androprosopa, Stilpnogaster, Tabanus [98], Eristalis [59], and also in
representatives of Culicidae [59, 99, 100, 109], Bombyliidae [60] and Mecoptera [80, 81, 82, 115]. In
Mengenilla the muscle originates also on the head capsule [84]. In Micropteryx M. tentorioscapalis anterior
8
isbipartite and originates on the tentorium [171]. The muscle is bipartite in Psychoda. One subcomponent
originates on the tentorium and the second one on the head capsule.
34.
Origin of M. tentorioscapalis posterior (M. 2): (0) tentorium; (1) head capsule. M. 2 originate on
the genal region of the head capsule in Tipula, between the clypeus and the margin of the compound eye
([58]: 2, figs 4, 11). In Axymyia [31] and Macrocera the muscle originates also on the gena. M.
tentorioscapalis posterior takes its origin in Culicoides on the apodeme of the frontal region ([113]: fig. 21,
acc. adductor of the scape), whereas the same condition occurs in Corethrella and Androprosopa. It also
originates on the head capsule in Pedicia, Deuterophlebia, Edwardsina [28], Mayetiola [30], Nymphomyia
[50], Hemipenthes [60], Ctenocephalus [83] and Mengenilla [84]. It is difficult to distinguish between M. 2
and M. 4 in Limonia. Both muscles lie very closely together and have a nearly identical point of insertion
on the scapus and closely adjacent areas of origin on the tentorium [58]. In Trichocera the origin lies
dorsally on the anterior tentorial arm and this is also the case in Aedes [99] and Wilhelmia [95]. In Culiseta
[59, 100] it originates laterally on the anterior tentorium and it arises on the dorsal arm in Tabanus [98].
The muscle also originates on the tentorium in Ptychoptera, Chaoborus, Bibio, Exechia, Coboldia,
Spathobdella, Sylvicola and Stilpnogaster like in all adults of Mecoptera examined [e.g. 80-82, 115] and in
Micropteryx [171]. The area of origin lies on the circumocular ridge in Eristalis [59] and a similar M.
orbitoscapalis is also present in Bombylius [60]. Schiemenz [59] interpreted this muscle as M.
tentorioscapalis posterior (M. 2). However, it canot be excluded that it is in fact M. tentorioscapalis
medialis (M. 4). The muscle is bipartite in Psychoda. One subcomponent originates on the tentorium
(lateral of M. 1) and the second one on the head capsule, ventrolateral of M. 1. Cylindrotoma have one
muscle, which extends between the posteriomedial margin of the scapus and the dorsal wall of the
tentorium. It is not clear if the muscle is homologous with M. tentorioscapalis posterior or M.
tentorioscapalis medialis.
35.
Origin of M. tentorioscapalis medialis (M. 4): (0) tentorium; (1) frontal region of head capsule;
(2) genal region of head capsule; (3) on the vertex. M. tentorioscapalis medialis originates on the genal
region in Tipula ([58]: 4, figs 4, 11) and Nymphomyia [50], and on the frons in Ptychoptera, Trichocera,
Bibio, Exechia, Mayetiola [30], Spathobdella, Sylvicola, Tabanus, Pachygaster, Nannochorista and
Mengenilla [58, 80, 84, 98]. It originates on the vertex in Panorpa [115], and on the clypeus in
Ctenocephalus [83]. The muscle is absent in representatives of Culicidae [59, 99, 100, 109] and
Simuliidae [95]. It originates on the anterior tentorium in Mischoderus, Macrocera, Culicoides [113],
Corethrella, Chaoborus, Androprosopa, Hemipenthes [60], Merope [82] and Caurinus [81]. As pointed out
above, the homology of the muscle is not entirely clear in Limonia, Eristalis and Bombylius (see character
21). In Micropteryx M. tentorioscapalis medialis originates on the tentorium [171]. The muscle is bipartite
in Psychoda. One subcomponent originates on the tentorium and the second one on the frontal region of
the head capsule.
36.
Mandible: (0) present; (1) absent. The mandibles are absent in males and females in
Deuterophlebia [28], Nymphomyia [50], Chironomus [41], Corethrella, Chaoborus, Androprosopa, Dixa,
Tipula, Limonia, Trichocera [58], Pedicia, Cylindrotoma, Bibio, Axymyia [31], Macrocera, Exechia,
9
Mayetiola [30], Coboldia, Spathobdella, Sylvicola, Psychoda, Mischoderus, Protoplasa [94], Ptychoptera,
Stilpnogaster, Pachygaster, Eristalis [59], Drosophila [97], in representatives of Bombyliidae [60], and also
in Ctenocephalus [83]. They are only developed in females of Edwardsina, Symphoromyia (Rhagionidae)
[70] and Tabanidae [98], whereas they are always present in adults of Ceratopogonidae [125], Simuliidae
[95], Mecoptera [e.g. 80, 81, 115, 120], Strepsiptera (Pohl et al. 2005 [84] and in Micropterygidae [171]. In
Culicidae mandibles are present in all females and the most males, except Aedes and Ochlerotatus [59,
99, 100]. The mandible in culicid males is much shorter than in females [108].
37.
Shape of the mandibles: (0) tansformed into piercing stylets; (1) not transformed into
piercing stylets. The mandibles are transformed into piercing stylets in adults of Culicidae and
Ceratopogonidae. They are very thin and lie between the labrum and hypopharynx [59, 99, 100, 125]. The
mandibles of Wilhelmia are spoon-shaped [95]. According to Peterson [41] the mandibles are also
elongated in Tabanus, Culicoides, and females of Bibiocephalia and Blepharocera (Blephariceridae).
Piercing stylets are absent in all adults of Strepsiptera [84] and Mecoptera [e.g. 80-82, 115, 120], but the
mandibles are strongly modified, elongate and lamelliform in Nannochorista [80]. The mandibles of
Micropteryx are also not transformed into piercing stylets [171].
38.
M. craniomandibularis internus (M. 11): (0) present; (1) absent. The muscle is present in
Culiseta ([59, 100]: M. tentorio-mandibularis), Aedes [99], Culex, Anopheles [368], Culicoides [113],
Wilhelmia [95], Mengenilla [84], Micropteryx [171] and also in adults of all mecopteran groups except for
Nannochorista [80-82]. Mandibular muscles (M.11-M.13) are only present in females of Tabanus [98].
39.
M. craniomandibularis externus (M. 12): (0) present; (1) absent. M. craniomandibularis externus
is present in Aedes [99], Culex, Anopheles [368], Culicoides [113], Wilhelmia [95], Mengenilla [84],
Micropteryx [171] and females of Tabanus [98], as in all mecopteran groups with the exception of
Nannochorista [80, 82]. Schiemenz [59] described only one mandibular muscle for Culiseta (M. 11), but a
second one is mentioned by Wenk ([368] M. retractor mandibulae tentorialis) and Owen ([100] M. oculomandibularis, M. 12).
40.
M. hypopharyngomandibularis (M. 13): (0) present; (1) absent. The muscle is usually absent in
adults of Culicidae [59, 99, 100, 368], but is present in Anopheles according to Wenk [368]. It is also
present in representatives of Simuliidae ([95] 1admd), Culicoides [113], females of Tabanus [98], and in
Nannochorista, Boreus and Bittacus [80]. The muscle is unusually large In Caurinus [81]. M.
hypopharyngomandibularis is absent in Mengenilla [84], Merope [82] and Micropteryx [171].
41.
Maxilla: (0) present; (1) absent. The maxilla is absent in Deuterophlebia [28], Nymphomyia [50]
and also missing in some chironomids [308]. It is present in all other members of Diptera [e.g. 28, 30, 31,
58, 59, 70, 94, 95, 98-100, 109, 125], Mecoptera [80-82, 115, 120], Siphonaptera [83], Strepsiptera [84]
and Lepidoptera [171].
42.
Maxillary endite lobes: (0) lacinia; (1) lacinia and galea; (2) both absent.
The maxilla is
completely reduced in Deuterophlebia [28] and Nymphomyia [50]. Both endites are absent in Tipula,
Erioptera, Mycetophila and Fucellia [70], Axymyia [31], Mayetiola [30], and in the most groups of
Strepsiptera (with the exception of Protoxenos [84]). The lacinia is present in adults of most groups of
10
Diptera [e.g. 58-60, 70, 94, 98, 100, 125], and also in Siphonaptera [83, 127] and Nannochorista [80].
Both endite lobes are present in mecopterans [81, 82, 115, 120]. Lacinia and galea are present in
Micropteryx [171].
43.
Maxillary palp: (0) 5-segmented; (1) 4-segmented; (2) 3 palpomeres or less. The maxillary palp
is 5-segmented in Edwardsina [28], Tipula, Limonia, Trichocera, Mischoderus [58], Protoplasa [94],
Ptychoptera, Culicoides [125], Chironomus [41], Dixa [41], Corethrella, Chaoborus, Androprosopa,
Wilhelmia [95], Bibio, Macrocera, Mycomya, Sylvicola, Agathon [70], Nannochorista [80], Panorpa [115],
Caurinus [81], Merope [82] and Micropteryx [171]. In Axymyia the palps are also 5-segmented; however
some individuals have four or five segments [31, 369]. The palp is 4-segmented in Psychoda, Exechia and
representatives of Siphonaptera [83, 127]. In adults of Culicidae the palps can comprise 1-5 palpomeres
(e.g. [99] Aedes, 5-segmented; 69, Anopheles, 4-segmented). In Culex the palps of the females are 3segmented and those of the males 5-segmented. In Mayetiola [30] and Spathobdella the palps are 3segmented and in Coboldia and Drosophila [97] only one segment is present. The number is usually
reduced in representatives of Brachycera. The palps are 2-segmented in Symphoromyia, Rhagio and
Dioctria [70], and only one segment is present in Pachygaster, Eristalis [59], Toxophora, Sepsis, Fucellia
[70] and Tabanus [98]. The papls in males of Strepsiptera are only one-segmented [84].
44.
Last segment of the maxillary palp: (0) not elongated; (1) elongated. The last segment of the
maxillary palp is elongated in Androprosopa, Tipula, Limonia, Trichocera [58], Exechia, Mayetiola [30],
Ptychoptera and Protoplasa (94, fig. 9). The last palpal segment is slightly elongated in Spathobdella and
Chaoborus. It is not elongated in Dixa [41], Culex, Culiseta [59], Corethrella, Mischoderus, Bibio, Axymyia
[31], Macrocera, Sylvicola, Psychoda, Culicoides [113], Edwardsina [28], Tabanus [98], Pachygaster,
Drosophila [97], Caurinus [81], Panorpa [115], Nannochorista [80], Merope [82], Ctenocephalus [83] and
Micropteryx [171]. The maxillary palp of Eristalis is one segmented and the segment is slightly enlarged
[59] (coaded as 1); the same condition occours in Coboldia.
45.
Sensorial field on maxillary palpomere 3: (0) present; (1) absent. A sensorial field is present on
the maxillary palpomere 3 of Edwardsina [28], Tipula, Limonia, Trichocera [58], Ptychoptera, Bibio,
Sylvicola, Culicoides (125), Corethrella, Wilhelmia [95], Mycetophila [70] and Nannochorista [80]. It is
absent in all other representatives of Mecoptera (e.g. 115; 81; 82), in Culiseta [59], Culex, Chaoborus,
Andropsosopa, Axymyia [31], Mayetiola [30], Spathobdella, Psychoda and also in Siphonaptera (83;
Michelsen 1996a). In Exechia a sensorial field is present on palpomere 2, whereas the palps are only 4
segmented its possible that this sensorial field is homologous with the field on the third palpomere in other
groups of Diptera.
46.
Position of sensilla on sensorial field: (0) Sensilla placed in a groove; (1) each sensilla in a
single groove; (2) Sensilla exposed on the surface. The sensilla placed together in a large groove in
Edwardsina, Bibio, Culicoides (125), Wilhelmia (95, fig. 24), Mycetophila [70] and Nannochorista [80]. In
Tipula, Corethrella and Sylvicola each sensillum is inserted in an individual groove and they are exposed
on the surface of the palpomere in Limonia, Trichocera [58] and Ptychoptera. In the sensorial field in
Exechia the sensilla are placed in a groove.
11
47.
Stipes: (0) exposed; (1) internalised (cryptostipes sensu 41). The stipites are exposed in
Ptychoptera, Edwardsina [28], Trichocera, Pedicia, Mischoderus [58], Protoplasa [94], Bibio, Macrocera,
Exechia, Mayetiola [30], Coboldia, Spathobdella, Sylvicola, Psychoda, Culicoides [113], Dixa [41],
Corethrella, Chaoborus, Androprosopa and Pachygaster, Micropteryx [171] and this is also the case in
adults of Mecoptera (115; 120; 80; 81; 82), Siphonaptera [83] and Strepsiptera [84]. They are exposed in
females of Tabanus, whereas the stipites of the males are largely reduced and internalised [98]. The
stipites of Tipula, Limonia, Cylindrotoma, Culex, Anopheles [58], Culiseta (59; 100), Aedes (99), Wilhelmia
[95], Hemipenthes, Bombylius [60], Drosophila [97], Eristalis [59] and Toxophora [70] are internalised.
48.
Stipites: (0) separated; (1) partly fused; (2) completely fused. The stipites are fused in
Edwardsina, Tipula [58], Cylindrotoma, and some representatives of Brachycera (Stilpnogaster, 70, figs
66, 67, Dioctria, figs 69, Sepsis, fig. 72, Fucellia, fig. 72). They are only fused posteriorly and thus form a
Y-shaped rod-like structure in Toxophora [70], Limonia [58], Pedicia and some representatives of
Tipulidae (70, Erioptera, Limnophila, Dicranomyia). They are also partly fused in Coboldia and
Spathobdella. The stipites are not fused in adults of Ptychoptera, Trichocera [58], Dixa [41], Culicoides
[41], Corethrella, Chaoborus, Androprosopa, Protoplasa [94], Bibio, Macrocera, Exechia, Mayetiola [30],
Sylvicola, Psychoda, Ptychoptera, Mycetophila, Mycomya [70], Culicidae (109; 59; 99; 100), Wilhelmia
[95], Tabanus [98], Pachygaster, Drosophila [97], Eristalis [59], Symphoromyia, and Rhagio [70], and this
condition is also found in Nannochorista [80], Panorpa [115], Merope [82], Ctenocephalus [83], Mengenilla
[84] and Micropteryx [171]. A specific maxillolabial complex is present in Caurinus and other Boreidae.
The proximal parts of the stipites, the cardines and the anterior postmentum form a structural and
functional unit [81].
49.
Cardo and stipes: (0) not fused; (1) fused. Both proximal maxillary elements are usually fused in
adults of Diptera (e.g. 41; 94; 70; 59; 99; 95; 100; 60; 58; 28), and this is also the case in Nannochorista
[80], Caurinus [81] and representatives of Siphonaptera (Michelsen 1996a). Cardo and stipes are present
as separate structures in Trichocera [58], Dixa [41], Mycomya (70, fig. 60B), Tabanus [98], Panorpa [115],
Merope [82] and Micropteryx [171]. In Strepsiptera the cardo is vestigial or absent, because of the missing
musculature of the maxilla it is unknown if cardo and stipes are fused or the cardo is reduced [84].
50.
M. craniocardinalis (M. 15): (0) absent; (1) present. The muscle is absent in all dipteran adults
examined (e.g. 109; 98; 59; 99; 95; 100; 60; 58; 28, 2013a, b) and is apparently generally missing in
Antliophora [80]. All extrinsic and intrinsic musculature of the maxilla is absent in Mengenilla [84]. M.
craniocardinalis is present as a bipartite muscle in Micropteryx [171].
51.
Origin of M. tentoriocardinalis (17): (0) tentorium; (1) head capsule; (2) fulcral plates. The
origin lies on the tentorium in Edwardsina, Mischoderus, Trichocera [58], Pedicia, Mayetiola [30], Exechia,
Spathobdella, Sylvicola, Psychoda, Corethrella, Androprosopa, Culicoides, in representatives of Culicidae
(109; 59; 99; 100), Erioptera, Ptychoptera [70], Nannochorista, Bittacus [80], Caurinus [81], Merope [82]
and in Micropteryx [171]. The muscle is bipartite in Wilhelmia. One bundle originates lateroventrally on the
tentorium, the other one on its ventral side (95, fig. 20, 1, 2adcd). The muscle arises on the lateral wall of
the clypeus in Tipula, Limonia [58], Limnophila, Dicranomyia, Dioctria [70] and Macrocera, and from the
12
dorsal wall in Fucellia [70], in representatives of Rhagionidae (70, in Symphoromyia, Rhagio), and in
Panorpa [115]. The origin of M. tentoriocardinalis and a part of M. tentoriostipitalis lies on the clypeal
region in Boreus [80]. M. 17 originate on the gena dorsal and lateral to the anterior tentorial pit in females
and on the clypeal region in males of Tabanus [98]. In Eristalis the muscle is attached at the basal anterior
edge of the fulcral plate [59] and an origin on the fulcrum is also described for representatives of
Bombyliidae [60]. M. 17 is absent in Bibio, Axymyia [31], Pachygaster and Ctenocephalus [83].
52.
Origin of M. tentoriostipitalis (M. 18): (0) tentorium; (1) head capsule. M. tentoriostipitalis
originates on the clypeus in Tipula and Limonia [58] and also on the head capsule in Mischoderus. The
origin lies on the tentorium in Ptychoptera, Trichocera [58], Pedicia, Cylindrotoma, Culex (109), Culiseta
(59; 100), Aedes (99), Culicoides [113], Corethrella, Wilhelmia [95], Bibio, Macrocera, Exechia, Mayetiola
[30], Coboldia, Spathobdella, Sylvicola, Psychoda, Edwardsina [28], Tabanus [98], Pachygaster, Eristalis
[59], Bombylius, Hemipenthes [60], Micropteryx [171] and also in Nannochorista [80] and other members
of Mecoptera (80; 81; 82). In Ctenocephalus and Axymyia the M. tentoriostipitalis is absent (83;
Schneeberg et al. 2013b). The muscle originates on the head capsule in Drosophila (97 [maxillary
muscle]), but the homology of the muscle is not entire clear.
53.
M. craniolacinialis (M. 19): (0) present; (1) absent. The muscle is present in members of
Culicidae (109; 59; 99; 100), in Culicoides [113], Wilhelmia [95], Sylvicola, Hemipenthes, Bombylius [60],
Drosophila (97 [maxillary retractor muscle]; 111 [4]), and only in females of Tabanus [98]. M.
craniolacinialis is also present in Nannochorista [80], Caurinus [81], Bittacus [80], Panorpa [115], Merope
[82], Ctenocephalus [83] and Micropteryx [171]. M. 19 is missing in Mischoderus, Ptychoptera, Tipula,
Limonia [58], Pedicia, Cylindrotoma, Corethrella, Androprosopa, Bibio, Macrocera, Axymyia [31],
Mayetiola [30], Exechia, Coboldia, Spathobdella, Psychoda, Edwardsina [28], Pachygaster, Eristalis [59]
and Boreus [80].1
54.
M. stipitopalpalis externus (M. 22): (0) present; (1) absent. M. stipitopalpalis externus is present
in members of Tipulomorpha [58], in Ptychoptera, Culiseta (59; 100), Culicoides [113], Corethrella,
Chaoboridae, Wilhelmia [95], Mischoderus, Bibio, Macrocera, Exechia, Spathobdella, Sylvicola,
Psychoda, Edwardsina [28], and is also well developed in members of Mecoptera (115; 80; 81; 82),
Siphonaptera [83] and Lepidoptera [171]. The muscle is absent in Aedes (99), Androprosopa, Axymyia,
Coboldia, and all examined members of Brachycera (111; 98; 59; 60).
55.
M. stipitopalpalis internus (M. 23): (0) present; (1) absent. M. stipitopalpalis internus is present
in Edwardsina [28], Tipula [58], Pedicia, Cylindrotoma, Macrocera and Coboldia but absent in
Mischoderus, Ptychoptera, Limonia, Trichocera [58], Culex (109), Culiseta (59; 100), Corethrella,
Androprosopa, Wilhelmia [95], Axymyia [31], Exechia, Sylvicola, Spathobdella, Psychoda, Tabanus [98],
Pachygaster, Eristalis [59], Hemipenthes, Bombylius [60], Drosophila [111], Nannochorista, Boreus,
Bittacus [80], Caurinus [81], Merope [82] and Micropteryx [171]. Christophers (1960) described a muscle
in Aedes extending from the anterior region of the stipes to the dorsal base of the palp. It functions as
levator of the entire palp and is only present in males (99, 13). Two muscles are present in the groundplan
13
of Diptera according to Hennig (1973) and Matsuda (1965). This condition is only verified for Edwardsina
and Tipula.
56.
M. palpopalpalis maxillae primus (M. 24): (0) present; (1) absent. In Mischoderus, Tipula,
Limonia, Trichocera [58], Pedicia, Cylindrotoma, Aedes (99), Culiseta (59; 100), Corethrella, Chaoboridae,
Androprosopa, Wilhelmia [95], Macrocera, Exechia, Sylvicola, Edwardsina [28], Tabanus [98] and also in
Nannochorista, Bittacus [80], Panorpa [115], Merope [82], Ctenocephalus [83] and Micropteryx [171] M.
palpopalpalis maxillae primus is well developed. Whereas the muscle is absent in Ptychoptera, Axymyia
[31], Coboldia, Spathobdella, Psychoda, Eristalis [59], members of Bombyliidae [60], Drosophila [111],
and in Caurinus [81] and Boreus [80].
57.
M. palpopalpalis secundus (M. 25): (0) present; (1) absent. This muscle is absent in
Chaoboridae, Androprosopa, Edwardsina, Tipula, Limonia [58], Pedicia, Wilhelmia [95], Axymyia [31],
Exechia, Coboldia, Spathobdella, Sylvicola, Psychoda, Tabanus [98], Pachygaster, and Eristalis [59], and
is also lacking in representatives of Bombyliidae [60] and Mecoptera (80, Nannochorista, Boreus, Bittacus;
81, Caurinus; 82, Merope) (with the exception of Panorpa, 115) and Siphonaptera [83]. A muscle with an
origin on the dorsolateral basal margin of palpomere 1 and an insertion on the lateral basal margin of
palpomere 3 is present in Mischoderus, Trichocera [58] and Corethrella. It is likely homologous with M.
stipitopalpalis secundus (coaded as 0). M. palpopalpalis secundus is present in Ptychoptera, Culiseta (59;
100), Cylindrotoma and Macrocera. The muscle is present in Micropteryx [171].
58.
Labium: (0) present; (1) absent. In all dipterans (e.g. 109; 94; 98; 70; 59; 95; 99; 100; 125; 58,
2013a, b), mecopterans (115; 120; 80; 81; 82), siphonapteran [83] and Lepidoptera [171], with the
exception of Deuterophlebia [28], Nymphomyia [50] and some Chironomidae (308), a labium is present. In
Mengenilla the labium is not developed as a separate sclerite and is integrated into the mouthfield sclerite
[84].
59.
Postmentum: (0) present; (1) absent. The postmentum is reduced or completly fused with the
prementum in members of Tipulomorpha [58], Mischoderus, Ptychoptera, Wilhelmia [95], Dixa [41],
Chaoboridae, Androprosopa, Edwardsina, in Bibio, Axymyia [31], Macr1ocera, Exechia, Mayetiola [30],
Coboldia, Spathobdella, Sylvicola, Psychoda, Stilpnogaster, Pachygaster, Tabanus [98], Hemipenthes,
Bombylius [60], Drosophila [97], Eristalis [59] and also in Mengenilla [84]. It is well developed in Culic1ides
(113, fig. 16). In Nannochorista the postmentum is indistinctly separated from the prementum [80]. In
Panorpa [115], Caurinus [81], Merope [82], Ctenocephalus [83] and Micropteryx [171] the postmentum is
also present.
60.
Dorsal surface of the anterior labium with distinct concavity: (0) present; (1) absent. The
dorsal concavity is absent in Tipula [58], Bibio, Axymyia [31], Macrocera, Exechia, Mayetiola [30],
Spathobdella, Pachygaster, and Mengenilla [84]. The lateral premental walls are slightly bent upwards in
Edwardsina [28], Mischoderus, Limonia, Trichocera [58], Pedicia, Cylindrotoma, Culicoides [113],
Androprosopa, Ptychoptera, and distinctly in Coboldia, Sylvicola, Psychoda, Corethrella, Chaoborus and
representatives of Culicidae (109; 107; 59; 99; 100). In adults of Culicidae the anterior labium forms a
concavity for the reception of the piercing mouthparts in their resting position. A similar dorsal concavity is
14
also present in adults of Simuliidae (95; 128), Bombyliidae [60] and Siphonaptera [83], and also in
Tabanus [98], Eristalis [59] and Nannochorista [80]. It is absent in Caurinus and other mecopterans (115;
120; 81).
61.
Prementum: (0) without median ridge; (1) with median ridge. A median ridge of the prementum
is absent in Tipula [58], Pedicia, Cylindrotoma, Bibio, Axymyia [31], Exechia, Mayetiola [30], Ptychoptera,
Tabanus [98], Pachygaster, Bombylius (60: fig. 2c, pr) in representatives of Culicidae (107; 59; 99; 100),
and in Siphonaptera [83], Nannochorista and most other groups of Mecoptera (120; 80) (a strongly
developed posteromedian apodeme is present in Caurinus, 81). The ridge is present and the prementum
W-shaped in cross section in Limonia, Trichocera [58], Macrocera, Coboldia, Spathobdella, Sylvicola,
Psychoda, Culicoides (113, fig. 27), Corethrella, Chaoborus, Androprosopa, Mischoderus and
Hemipenthes (60: fig. 1c, pr). It is T-shaped in cross section in representatives of Simuliidae (95; 128). A
median ridge is also present in Eristalis, but small and inconspicuous (59, fig. 56). The prementum is not
recognisable as a separate structure in Mengenilla [84].
62.
Number of labial palpomeres: (0) 2; (1) 3; (2) 5. The labium is largely reduced in Deuterophlebia
[28] and Nymphomyia [50] and the palps are completely absent. In Mengenilla the palps are also
completely missing [84]. It is almost generally 2-segmented in adults of Diptera [e.g. 30, 31, 58, 59, 69, 70,
94, 95, 98, 99, 113] and this is also the case in Mecoptera [80, 81, 82, 115, 120] and also in Micropteryx
[171]. Schiemenz [59] and Owen [100] described 3-segmented palps for Culiseta, whereas they were
interpreted as 2-segmented by Harbach & Kitching [69]. Only a single sclerite is present in Anopheles. It is
divided dorsally, which suggests that it is a product of fusion and also 2-segmented [69]. The palps are 5segmented in representatives of Siphonaptera [127]. Tokunaga [50] described a small membranous
appendage at the entrance of the mouth opening in Nymphomyia, it is possible that this are the reduced
labial palps.
63.
Labialpalps modified as labellae: (0) present; (1) absent. The 2-segmented labial palps
(Crampton 1942) are modified as thickened labellae in all adults of Diptera with a preserved labium [e.g.
30, 31, 58-60, 69, 107; 98; 99; 95; 100; 128; 125;]. This is not the case in the other antliophoran lineages
(e.g. 80; 81; 82) and in Micropteryx [171].
64.
Pseudotracheae on the internal side of the labellae: (0) absent; (1) present. The membranous
mesal sides of the labellae are equipped with two rows of pseudotracheae in Tipula [58]. This is rarely the
case in the nematoceran groups. They occur in Tipulini, and in some members of Mycetophilidae (some
Sciophilinae and Mycetophilinae, e.g. Exechia) and Ptychopteridae [70], but are usually missing. Within
Brachycera pseudotracheae are more widespread and more complex. Two pseudotracheal collecting
channels are present on the anterior edge of the labellae of Eristalis and about 40 pseudotracheae on the
mesal wall ([59] fig. 54). The number of pseudotrecheae varies within Bombyliidae ([60] fig. 4) and
Tabanidae [98]. Within Brachycera they are lacking in representatives of Asilidae [70]. Pseudotracheae
are presen ton the labellum of Drosophila [97].
65.
Simple furrows on the mesal sides of the labellae: (0) present; (1) absent. Simple furrows are
present on the mesal side of the labellae of Androprosopa, Trichocera [58], Pedicia, Cylindrotoma,
15
Axymyia [31], Sylvicola, Psychoda and some Culicids (Culex, Anopheles, Culiseta). They were referred to
as pseudotracheae by Owen [100] but it was already demonstrated by Schiemenz [59] that their
ultrastructure is distinctly different, i.e. that inner strenghtening rings are absent and also secondary
channels (e.g. [131]: figs 3, 4, [132]: fig. 2a, Chrysomya). The furrows are absent in Corethrella,
Chaoborus, Mayetiola [30], Spathobdella, Stilpnogaster and Nannochorista [80]. In Culicoides the wall of
the second segment of the “labella are thin and much folded at the sides” [113]. It is quite possible that
these are furrows (coaded as 0).
66.
Scales on labial palps: (0) absent; (1) present. Scales are almost generally missing in Diptera
[30, 31, 58-60, 69, 94, 95, 98, 125, 128]. They are also absent in Merope [82]. They occur in some
representatives of Culicidae on the external side of the labial palps ([69]: fig. 9A, B, Toxorhynchites,
Tripteroides) and on the mesal side in Nannochorista [80]. Scales are also present on the labial palps in
Micropteryx [171].
67.
Glossa and paraglossa: (0) absent; (1) present. Glossa and paraglossa are generally absent in
all groups of Antliophora [e.g. 58-60, 80-82, 95, 100, 120, 128] and also in Strepsiptera [84]. They are
present in Micropteryx [171].
68.
Retractor of the prementum (Mm. 28-30): (0) one muscle; (1) two muscles; (2) absent; (3)
three muscles. One large retractor is generally present in representatives of Diptera [30, 31, 58-60, 70,
95, 98, 99, 100, 109, 113] and Mecoptera [80-82, 115, 120]. It is probably M. tentoriopraementalis inferior
(M. 29) or a product of fusion of both tentoriopremental muscles [80]. A premental retractor is absent in
Siphonaptera [83, 127]. In Mengenilla the extrinsic musculature of the labium is completely missing [84].
Three muscles are present in Micropteryx [171]. Two retractor muscles are present in Drosophila after
Miller ([111]: 1, 2).
69.
M. palpopalpalis labii primus (M. 35): (0) present; (1) absent. The muscle is present in
Mischoderus, Ptychoptera, Edwardsina [28], Cylindrotoma, Axymyia [31], Macrocera, Exechia, Culicoides
[113], Androprosopa and Micropteryx [171]. A muscle originating on a median longitudinal premental ridge
and inserting on the mesal basal margin of palpomere 2 occurs in Simuliidae [95, 123], and a similar
muscle is also present in representatives of Bibionidae, Chironomidae, Rhagionidae, Asilidae and
Sphaeroceridae [70]. It is absent in Limonia, Trichocera [58], Pedicia, Coboldia, Spathobdella, Sylvicola,
Psychoda, Corethrella in representatives of Culicidae [59, 99, 100, 107], Pachygaster, Eristalis [59],
Hemipenthes, Bombylius [60], Drosophila [111], Tabanus [98], Fucellia [70], and in representatives of
Mecoptera [80-82, 115], Siphonaptera [83] and Strepsiptera [84].
70.
M. frontohypopharyngalis (M. 41): (0) present; (1) absent. The muscle is absent in
Deuterophlebia, Edwardsina [28], Nymphomyia [50], Limonia, Trichocera [58], Pedicia, Cylindrotoma,
Bibio, Axymyia [31], Macrocera, Exechia, Mayetiola [30], Spathobdella, Sylvicola, Psychoda, Wilhelmia
[95], Culicoides [113], Corethrella, Androprosopa, Pachygaster, Stilpnogaster, Drosophila [97] and in
representatives of Bombyliidae [60]. A small bipartite muscle is present in Tipula and Mischoderus. It
extends from the lateral clypeal wall to the lateral wall of the hypopharynx [58]. The homology with M.
frontohypopharyngalis is questionable. A muscle connecting the postfrontal ridge and the dorsal fulcral
16
apophyses is present in representatives of Culicidae [59, 100, 109] and in Eristalis [59]. It is likely that it
mainly stabilises the cibarium and it is probably homologous with M. frontohypopharyngalis. However,
Schiemenz [59] assumed its homology with M. frontobuccalis lateralis (M. 47) (Culiseta and Eristalis) and
a similar muscle is present in Aedes. It originates on the median frontal ridge and is inserted on the lateral
horn of the cibarium ([99]: fig. 67/1: 21). A muscle, which connects the clypeofrontal ridge and the anterior
surface of the pharynx, is present in Tabanus ([98] fig. 10: 22), but the homology with M. 41 is also
questionable. In Coboldia a muscle connecting the head capsule (ventral of the circumocular ridge) with
the lateral wall of the hypopharynx, it is probably homologous with M. 41. M. frontohypopharyngalis is
present in Ctenocephalus [83], Mengenilla [84], Nannochorista, Boreus, Bittacus, Merope and Caurinus,
but absent in Panorpa [80-82, 115, 120] and Micropteryx [171].
71.
Size of M. clypeopalatalis (M. 43): (0) long series of bundles; (1) bipartite, strongly
developed; (2) not enlarged. The muscle is not enlarged in Deuterophlebia [28]. M. clypeopalatalis is a
long series of bundles in Edwardsina [28], Ptychoptera, and also in Tipula, Limonia, Trichocera [58],
Pedicia, Cylindrotoma, Bibio, Axymyia [31], Macrocera, Exechia, Mayetiola [30], Coboldia, Spathobdella,
Sylvicola, Psychoda, Mischoderus, Stilpnogaster, Pachygaster, Tabanus [98], Eristalis [59], Hemipenthes,
Bombylius [60], Drosophila ([111]: 12), Culicoides [113], Corethrella, Androprosopa and most members of
Culicidae [59, 100, 109]. A similar condition is found in Panorpa [115], Boreus [80], Merope [82],
Ctenocephalus [83] and Mengenilla [84]. Christophers [99] described a bipartite muscle for Aedes, and a
similar condition is present in Wilhelmia [95] and Nannochorista [80]. It is composed of three
subcomponents in Caurinus and is not enlarged [81]. Tokunaga [50] described a large muscle with three
subcomponents for Nymphomyia ([50] figs 1, 7). In Micropterix the muscle is not enlarged ([171]: 45, fig.
17).
72.
Mm. frontobuccalis anterior/posterior (Mm. 45/46): (0) both present; (1) one muscle; (2)
absent. M. frontobuccalis posterior (M. 46) is the only dorsal precerebral dilator in Deuterophlebia [28],
Tipula, Limonia [58], Cylindrotoma and representatives of Bombyliidae [60]. Both precerebral dorsal
dilators are present in Edwardsina, Trichocera [58], Axymyia [31], Macrocera, Sylvicola, Psychoda,
Mischoderus, Corethrella, in representatives of Culicidae [59, 95, 99, 100, 109], and in Tabanus [98],
Pachygaster, Drosophila [111], Caurinus [81], Merope [82], Ctenocephalus [83] and Micropteryx [171].
Both muscles are absent in Eristalis [59], Nannochorista [80] and Panorpa [115]. A series of bundles is
present between the frontal ganglion and the brain in Nymphomyia [50] and also in Culicoides [113]. It
probably comprises both muscles. Only one muscle is present in Boreus, Bittacus [80], Androprosopa,
Exechia, Mayetiola [30], Bibio, Coboldia, Spathobdella, Pedicia, Stilpnogaster and Mengenilla [84].
73.
M. tentoriobuccalis anterior (M. 48): (0) present; (1) absent. The muscle is absent in
Deuterophlebia [28], Nymphomyia [50], Eristalis [59], Nannochorista, Bittacus [80], Panorpa [115],
Caurinus [81], Merope [82], Ctenocephalus [83] and Mengenilla [84]. Szucsich & Krenn [60] described a
protractor of the fulcrum in Hemipenthes and Bombylius, which is possibly homologous with M. 48 (mgc).
A muscle which extends from the ventrolateral wall of the anterior pharynx, below the frontal ganglion, to
the circum ocular ridge is present in Edwardsina. This muscle is probably homologous with M.
17
tentoriobuccalis anterior. M. 48 is present in Mischoderus, Tipula, Limonia, Trichocera [58], Pedicia,
Cylindrotoma, Ptychoptera, Bibio, Macrocera, Exechia, Axymyia [31], Mayetiola [30], Coboldia,
Spathobdella,
Sylvicola,
Psychoda,
Wilhelmia
[95],
Corethrella,
Chaoboridae,
Androprosopa,
Stilpnogaster, Pachygaster, Tabanus [98], Culicoides [113], and in representatives of Culicidae [59, 95,
99, 100, 109], Drosophila ([111]: 9) and also in Boreus [80]. The muscle is present as an unpaired muscle
in Micropteryx [171].
74.
Postcerebral pharyngeal pump (Mm. 51/52): (0) present; (1) absent. A strongly developed
postpharyngeal pump is present in Deuterophlebia, Edwardsina [28], Nymphomyia [50] and also in Tipula,
Limonia, Trichocera, Ptychoptera, Mischoderus [58], Pedicia, Cylindrotoma, Bibio, Axymyia [31],
Macrocera, Exechia, Mayetiola [31], Coboldia, Spathobdella, Sylvicola, Psychoda, Wilhelmia [95],
Culicoides [113], Corethrella, Chaoboridae, Androprosopa, Stilpnogaster, Pachygaster, Tabanus [98], and
does generally occur in Culicidae [59, 99, 100, 109]. It is also present in Nannochorista, Boreus, Bittacus
[80], Panorpa [115], Merope [82], Mengenilla [84], Micropteryx [171] and Siphonaptera [83], but is missing
in Eristalis [59], Drosophila [97] and representatives of Bombyliidae [60]. A bipartite M. verticopharyngalis
and a moderately developed M. tentoriopharyngalis posterior is present in Caurinus [81]. It is functionally
replaced by the labro-epipharnygeal pump in Cyclorrhapha according to Gouin [124].
75.
M. anularis stomodaei (M. 68): (0) enclosing the lateral and ventral wall of the posterior
pharynx; (1) ring muscle. Complete ring muscles are restricted to the oesophagus in Deuterophlebia
[28], and also in Corethrella, Mischoderus, Tipula, Trichocera [58], Pedicia, Cylindrotoma, Mayetiola [30],
Macrocera, Exechia, Coboldia, Spathobdella, Sylvicola, Psychoda, Stilpnogaster, and Nannochorista [80].
The bundles enclose only the ventral and lateral walls of the posterior pharynx and of the anteriore
pharynx in Edwardsina. Postpharyngeal ring muscles are present in Limonia [58], Aedes [99], Wilhelmia
[95], Tabanus [98], Nymphomyia [50] and Panorpa [115]. The postpharyngeal ring musculature is
completely absent in Mengenilla [84].
76.
M. hypopharyngosalivarialis (M. 37): (0) present; (1) absent. A salivary pump is absent in
Deuterophlebia [28], Axymyia [31] and Limonia [58]. M. hypopharyngosalivarialis is present in Edwardsina,
Tipula, Trichocera, Mischoderus [58], Pedicia, Cylindrotoma, Bibio, Macrocera, Exechia, Mayetiola [30],
Coboldia, Sylvicola, Psychoda, Culicoides [113], Corethrella, Androprosopa, Chaoborus, Drosophila
([111]: 13) and Stilpnogaster. It connects the ventral wall of hypopharynx with the dorsal wall of the salvary
duct. An extrinsic dilator is also present in adults of Culicidae [59, 99, 100, 109], Simuliidae [95],
Tabanidae [98], Syrphidae [59], Bombyliidae [60], and Nannochorista [80]. M. hypopharyngosalivarialis is
bipartite in Nymphomyia. It originates on the ventral membrane of the basipharynx and inserts on the
salivarium ([50]: fig. 1). It is also bipartite in Micropteryx [171]. A typical M. hypopharyngosalivarialis is
absent in Mecoptera (excluding Nannochoristidae, e.g. [80]), but a strong intrinsic muscle of the wall of the
salivary duct [115, 81] may be a derivative of this muscle.
77.
Exposure of the head: (0) fully exposed; (1) moderately retracted into prothorax, not fixed in
this position; (2) strongly retracted, fixed in this position. The head capsule of Tipula [27], Limonia
[62] and Tabanus [46] is strongly retracted and fixed in this position. In Axymyia [227], Dicranota [46] and
18
Odontomyia [46] the head is moderately retracted into the prothorax and not fixed in this position. The
head of Deuterophlebia [48], Nymphomyia [29], Protanyderus [32], Trichocera [45], Bibio [46, 75],
Phaenobremia [47], Bittacomorpha [72], Dixa [66], Simulium [46], Dasyhelea [135], Anopheles, Culiseta
[73], Androprosopa, Chaoborus [134, 238], Corethrella, Chironomidae [155], Anisopodidae [45], Exechia,
Blephariceridae [63], Psychodidae [45], Therevidae [46], Nannochorista [136], Panorpa [141] and
Nosopsyllus [139] are fully exposed.
78.
Orientation of the mouthparts: (0) prognathous or slightly inclined; (1) orthognathous. The
head of Deuterophlebia [48], Nymphomyia [29], Protanyderus [32], Tipula [27], Limonia [62], Trichocera
[45], Axymyia [227], Dicranota [46] Bibio [46], Phaenobremia [47], Bittacomorpha [72], Dixa [66], Simulium
[46], Dasyhelea [135], Anopheles [73, 154], Culiseta [73], Androprosopa, Chaoborus [134, 238],
Corethrella, Chironomidae [74, 124], Anisopodidae [45], Exechia, Blephariceridae [63], Psychodidae [45],
Therevidae [46], Tabanus [46], Odontomyia [46], Nannochorista [136] and Nosopsyllus [139] is
prognathous. The head is orthognath in Panorpa [141].
79.
Paired dorsolateral incisions: (0) absent; (1) present, short; (2) present, deep, at least
reaching anterior half of head capsule. In tipulid larval head have paired dorsolateral incisions;
separating a dorsomedian fragment of the head capsule from the remaining parts this condition is also
present in Limonia [27, 46, 61, 62]. They are present and short in Dicranota [46]. The incisions are absent
in larvae of Deuterophlebia [48], Nymphomyia [29], Protanyderus [32], Trichocera [45], Bibio [46], Axymyia
[227], Phaenobremia [47], Bittacomorpha [72], Dixa [66], Simulium [46], Dasyhelea [135], Anopheles [73,
154], Culiseta [73], Androprosopa, Chaoborus [134, 238], Corethrella, Chironomidae [74, 124],
Anisopodidae [45], Exechia, Psychodidae [45], Therevidae [46], Tabanus [46], Odontomyia [46],
Nannochorista [136], Panorpa [141] and Nosopsyllus [139]. The merging of the head with the thorax and
the anterior abdominal segment in Blephariceridae (only partly in Edwardsiinae) [63] is probably correlated
with tendency to form a hemicephalous condition somewhat similar to that of tipuloid larvae, and short and
broad dorsolateral incisions are present in Liponeura ([63]: fig. 2). However, the head is eucephalic,
without incisions in Edwardsina and Anispous ([63]: figs. 3, 7).
80.
Ventromedian incision of head capsule: (0) absent; (1) present; (2) triangular median
membranous sinus; (3) ventral head capsule entirely sclerotised. A very deep ventromedian incision
is present in tipulid larvae and larvae of Limonia [27, 46, 61, 62, 64, 152] and also in larvae of Dicranota
[46] and Trichocera [45]. The incision is absent in Deuterophlebia [48], Nymphomyia [29], Protanyderus
[32], Bibio [46, 75], Axymyia [227], Exechia, Phaenobremia [47], Bittacomorpha [72], Dixa [66], Dasyhelea
[135], Anopheles [73, 154], Culiseta [73], Androprosopa, Chaoborus [134, 238], Corethrella, Anisopodidae
[45], Psychodidae [45], Nannochorista [136], Panorpa [141] and Nosopsyllus [139]. A triangular median
membranous sinus is present in Simulium [46]. The ventral head capsule is entirely sclerotized in
Therevidae [46], Tabanus [46] and Odontomyia [46]. The ventral head capsule is also reduced in some
Blephariceridae ([63] but not in the groundplan, see Edwardsina, fig. 3).
81.
Externolateral plates with growth lines (intermolt cuticle deposition): (0) absent; (1) present.
Extensive externolateral plates with a zonal structure or growth lines are probably generally present in
19
larvae of Tipuloidea [27, 46, 62, 64], Deuterophlebiidae [16, 48], Blephariceridae [16] and in Protanyderus
[32]. The growth lines are absent in Nymphomyia [29], Bibio [46, 75], Axymyia [227], Exechia,
Phaenobremia [47], Bittacomorpha [72], Dixa [66], Simulium [46], Dasyhelea [135], Anopheles [73, 154],
Culiseta [73], Androprosopa, Chaoborus [134, 238], Corethrella, Chironomidae [74, 124], Anisopodidae
[45], Psychodidae [45], Tabanus [46], Odontomyia [46], Nannochorista [136], Panorpa [141] and
Nosopsyllus [139].
82.
Split cranial setae: (0) absent; (1) present. Split cranial setae are absent in Nymphomyia [29],
Protanyderus [32], Tipulomorpha [e.g., 27, 64], Bibionidae [46, 75], Exechia, Phaenobremia [47],
Ptychopteridae [72], Dixidae [66], Chironomidae [74, 124], Corethrella, Anisopodidae [45], Blephariceridae
[63], Therevidae [46], Tabanus [46], Odontomyia [46], Panorpa [141] and Nosopsyllus [139]. They are
present in Deuterophlebiidae [48], Axymyia [227], Culicidae [see 27], Androprosopa, Chaoboridae [238],
Simulium [46], Ceratopogonidae [64, 261], Psychodidae [64, 261] and Nannochorista [136].
83.
Frontoclypeal suture: (0) present; (1) absent. A frontoclypeal suture is generally lacking in
dipteran groups (e.g., 8, 27; 29; 32, 45-48, 63, 64, 66, 72-74, 124, 134, 135, 154, 260, 261] and also in
Siphonaptera [139]. A distinct, unsclerotised clypeal area is present in Pediciidae and other larvae of
Tipuloidea ([46, 62]: fig. 93, Tricyphona), but this is not equivalent the frontoclypeal suture [27]. A
frontoclypeal suture is present in Axymyia ([227]: fig. 3A, ts) and Panorpa ([141]: Epi.N., fig. 1). A
frontoclypeal suture is absent in Nannochorista, but an internal ridge is present, but it is morphological
rather a transclapeal sulcus than a true frontoclypeal suture ([136]: tcs, fig. 2).
84.
Coronal suture: (0) present at last 25% as long as the wall of head capsule; (1) present, less
than 25% of dorsal wall of head capsule; (2) absent. A long coronal suture is absent in most larvae of
Tipulomorpha [45, 62, 64], in Chironomidae [74, 124], Anopheles (73; 154), Culiseta (73), Dixa [66],
Corethrella, Simulium [46], Dasyhelea [135], Phaenobremia [47], Exechia, Psychodidae [45], Tabanus [46]
and Nosopsyllus [139]. However it is present and long in Pediciidae (46: “…deep median phragma
developed along the line of the coronal suture...”, fig. 18) and does also occur in Eriopterinae (62: figs 14,
21, 26) (see 27). In Protanyderus [32], Androprosopa, Bibio (46; 75), Axymyia (227 [fig. 3A, cs]),
Anisopodidae (45: fig. 2, c.s.), Blephariceridae (63 [co.ec.li.]), Nannochorista (136 [ecl, fig. 2]) and
Panorpa (141 [Cor.N., fig. 1]) the coronal suture is present at last 25% long as the head capsule. In
Chaoborus the coronal suture is present following Förster (2012 [cs, fig. 2B]), but absent after Schremmer
(1950). In Bittacomorpha [72] the coronal suture is a remnant line and in Nymphomyia it is present, but
short (29; fig. 3B). It is absent in Deuterophlebia [48] and Odontomyia [46]. The coronal suture is present,
but indistinct and short in Nymphomyia [29].
85.
Shape of frons: (0) V-shaped; (1) U-shaped. The frons, if present as a recognisable delimited
structure, is V-shaped in Nymphomyia [29], Tipulomorpha (62; Ooserbroek & Courtney 1995; 27),
Bibionidae (46; 75), Axymyia [227], Exechia, Blephariceridae [63], Chironomidae [74, 124], Anisopodidae
[45], Psychodidae [45], Dixa [66], Nannochorista [136] and Panorpa [141]. And U-shaped in Chaoborus
(238) and Dasyhelea [135]. In Deuterophlebia [48] and Bittacomorpha [72] frons and clypeus are
completely fused and the frons is not recognizable as a separate sclerite.
20
86.
Clypeus divided in ante- and postclypeus: (0) divided; (1) undivided. The clypeus is divided in
two areas in Limonia, Axymyia [227], Nannochorista ([136]: acl, fcl, figs. 1A, 2) and Panorpa ([141]: A.Cly.,
P.Cly., fig. 1). In Deuterophlebia [48], Nymphomyia [29], Protanyderus [32], Tipula [27], Dicranota [46],
Trichocera [174], Bibio [46, 75], Phaenobremia [47], Bittacomorpha [72], Dixa [66], Simulium [46],
Dasyhelea [135], Anopheles [73, 154], Culiseta [73], Androprosopa, Chaoborus [134, 238], Corethrella,
Chironomus [74, 124], Exechia, Anisopodidae [45], Psychodidae [45], Blephariceridae [63], Tabanus [46],
Odontomyia [46] and Therevidae [46] the clypeal region is undivided.
87.
Premaxillary suture and side plates: (0) absent; (1) present. A premaxillary suture and side
plates are present in Tipulidae, Limoniinae, Cylindrotomidae ([46]: fig. 14; [27, 62]) and in Protanyderus
[32]. The suture is absent in Dicranota [46], Deuterophlebia [48], Nymphomyia [29], Bittacomorpha [72],
Phaenobremia [47], Dixa [66], Simulium [46], Dasyhelea [135], Anopheles [73, 154], Culiseta [73],
Androprosopa, Chaoborus [134, 238], Chironomidae [74, 124], Exechia, Anisopodidae [45], Psychodidae
[45], Therevidae [46], Axymyia [227], Tabanus [46], Odontomyia [46], Nannochorista [136], Panorpa [141]
and Nosopsyllus [139].
88.
Strengthened margins of externo-lateralia: (0) absent; (1) present. The externo-lateralia are
strengthened in Tipulidae, Limoniinae and Cylindrotomidae [27, 62], but not in other larvae of Diptera [e.g.,
29, 32, 46, 48, 73, 74, 134, 154].
89.
Dorsal endocarina: (0) absent; (1) present. A distinctly developed dorsal endocarina is present in
Tipula ([27]: fig. 2A, ec), Limonia [62] and also in larvae of Pediciidae ([61, 46]: fig. 18; [62]: fig. 93),
Eriopterinae ([62]: 93, fig. 21), Protanyderus [32], Bibio (see char. 12 in [27]) and Axymyia (293 [fig. 2]).
The endocarina is absent in Deuterophlebia [48], Nymphomyia [29], Bittacomorpha [72], Phaenobremia
[47], Trichocera [45], Dixa [66], Corethrella, Simulium [46], Dasyhelea [135], Anopheles (73; 154), Culiseta
(73), Androprosopa, Chaoborus (134; 238), Chironomidae (74; 124), Exechia, Psychodidae [45],
Blephariceridae [63], Therevidae [46], Tabanus [46], Odontomyia [46], Nannochorista [136], Panorpa [141]
and Nosopsyllus [139].
90.
Tentorium: (0) present; (1) absent. The tentorium is completely absent in larvae of Tipula [27] and
also in larvae of Nymphomyia [29], Dicranota [46], Androprosopa, Exechia and Axymyia [227]. Whereas
the tentorium is present in Deuterophlebiidae [48], Limoniidae, Tanyderidae [32], Bibionidae [46, 75],
Sciaridae [47], Trichoceridae [45], Ptychopteridae [72], Dixidae [66], Corethrella, Simuliidae [46],
Ceratopogonidae [135], Culicidae [73, 154], Chaoboridae [134, 238], Chironomidae [155], Anisopodidae
[45], Blephariceridae [63], Therevidae [46], Tabanidae [46], Stratiomyiidae [46], Nannochoristidae [136],
Panorpidae [141] and Nosopsyllus [139]. In Psychodidae the tentorium mainly vestigial (coaded as 0) [45].
91.
Dorsal tentorial arm: (0) present; (1) absent. Dorsal arms are absent in Deuterophlebia [48],
Limonia, Protanyderus [32], Bibio [75], Bittacomorpha [72], Dixa [66], Simulium [46], Anopheles ([73, 154]:
fig. 2b, 4), Culiseta [73], Chaoborus [238], Corethrella, Chironomidae [155], Psychodidae [45], Therevidae
[46], Tabanus [46], Odontomyia [46] and Nosopsyllus [139]. They are present in Trichocera ([45]: te.d.),
Dasyhelea ([135]: fig. 3, ttd), Anisopodidae ([45]: fig. 2, te.d.) and Panorpa [141]. Dorsal arms are also
21
absent in Nannochorista, but a vestige of the dorsal arm is present in form of a slendet tissue strand
([136]: dta, fig. 5).
92.
Tentorial bridge: (0) well developed and sclerotised; (1) partly reduced with thin median
connection; (2) absent. The tentorial bridge is apparently present in Deuterophlebia [48], Bibio [75],
Nannochorista ([136]: tb, fig. 5) and Nosopsyllus ([139]: fig. 1, t.b.). It is absent in Limonia, Protanyderus
[32], Bittacomorpha [72], Trichocera [45], Dixa [66], Simulium [46], Dasyhelea [135], Anopheles [73, 154],
Culiseta [73], Chaoborus [238], Corethrella, Chironomidae [155], Psychodidae [45], Blephariceridae [63],
Therevidae [46], Tabanus [46] and Odontomyia [46]. It is well developed in Panorpa [141]. The tentorial
arms are connected by a thin median connection in Anisopodidae [45] and Paenobremia [47].
93.
Anterior tentorial arms: (0) present; (1) vestigial or absent; (2) arising from paraclypeal
phragma and strongly developed. Anterior tentorial arms are absent in Bibio [75], Simulium [46] and
partly in Chironomidae [74, 124]. Whereas they are present in Limonia, Deuterophlebia [48], Protanyderus
[32], Trichocera ([45]: te.a.), Paenobremia ([47]: fig. 15, BAT), Bittacomorpha [72], Dixa [66], Dasyhelea
([135]: fig. 3, 5, tta), Anopheles [73, 154], Chaoborus ([238]: t, fig. 6C), Corethrella, Culiseta [73],
Anisopodidae [45], Psychodidae [45], Blephariceridae [63], Odontomyia ([46]: tentorium, fig. 33A),
Nannochorista ([136]: ata, fig. 5), Panorpa ([141]: v.T.A., fig. 11) and Nosopsyllus [139]. The anterior
tentorial arms are strongly developed and arising from the paraclypeal phragma in Therevidae ([46]: fig.
31B) and Tabanus [46].
94.
Cuticular lense: (0) present; (1) absent. A cuticular lense is generally lacking in nematoceran
larvae with developed larval eyes [e.g., 8, 9, 29, 46, 47, 72, 74, 124, 134, 135, 238] and also in
Androprosopa, Axymyia [227], Exechia and Protanyderus [32]. It is absent in Nannochorista [136]. A
convex cuticular lens is present in Limonia, Odontomyia [46] and Panorpa [141].
95.
Lateral eyes: (0) simplified compound eyes; (1) several stemmata; (2) eyes spot; (3) absent.
A single or bipartite larval eye is found in Tipulidae [e.g., 46]. Eye sports occur in Deuterophlebia [48],
Nymphomyia [29], Trichocera [45], Simulium [46], Bibionidae [46, 75], Paenobremia [47], Ptychopteridae
[72], Dixidae [66], Dasyhelea [135], Culicidae [see 27], Chironomidae [74, 124] and Nannochorista [136].
Simplified compound eyes are present in Chaoboridae (134; 238) and Panorpa [141]. Larval eyes are
completely absent in Dicranota [46], Limonia [65], Axymyia [227], Protanyderus [32], Anisopodidae [45],
Psychodidae [45], Blephariceridae [63], Therevidae [46], Tabanus [46] and Nosopsyllus [139]. Eye spots
are present in Odontomyia [46].
96.
Articulation of labrum: (0) free; (1) partially fused with head capsule; (2) completely fused
with head capsule. A free labrum is present in larvae of Tipula and Limonia [e.g., 46], also in larvae of
Pediciidae [46], Ptychopteridae [72], Chironomidae [155], Culicidae [73, 154], Androprosopa, Chaoboridae
[134, 238], Dasyhelea [135], Dixa [66], Anisopodidae [45], Psychodidae [45], Protanyderus [32],
Therevidae [46], Nannochorista [136], Panorpa [141] and Nosopsyllus [139]. The labrum in Bibionidae [46,
75] and Axymyia [227] is partly fused with the head capsule. The labrum of Deuterophlebia [48],
Nymphomyia [29], Trichocera [45], Paenobremia [47], Simulium [46], Tabanus [46] and Odontomyia [46] is
completely fused with the head capsule.
22
97.
Shape of labrum: (0) transverse; (1) narrow and conical. A transverse labrum is present in
Deuterophlebiidae [48], Tanyderidae [32], Tipulidae [16, 27], Limoniidae [16], Bibionidae [46, 75], Exechia,
Paenobremia [47], Ptychopteridae [72], Dixidae [66], Simuliidae [46], Ceratopogonidae [135], Culicidae
[73, 154], Chaoboridae [134, 238], Chironomidae [74, 124], Nannochorista [136], Panorpa [141] and
Nosopsyllus [139]. It is narrow in Trichocera [45], Dicranota [46] and Blephariceridae [63]. The labrum is
conical in Nymphomyia [29], Androprosopa, Axymyia [227], Therevidae [46], Anisopodidae ([45]: fig. 18,
lb.), Psychodidae ([45]: fig. 9, lb), Tabanus [46] and Odontomyia [46].
98.
Subdivision of labrum: (0) absent; (1) present. A distinctly tripartite labrum as it is present in
Tipulidae ([46, 61]: Holorusia, [158]) is a potential autapomorphy of the family. The labrum is distinctly
narrowed (see previous character) or transverse and undivided (e.g., Pediciidae) in the other tipuloid
groups [e.g., 27, 46, 62, 152]. The labrum is undivided in Deuterophlebiidae [48], Nymphomyia [29],
Trichoceridae [45], Tanyderidae [32], Bibionidae [46, 75], Exechia, Axymyiidae [227], Ptychopteridae [72],
Paenobremia [47], Anisopodidae [45], Chironomidae [74, 124], Culicidae [73, 154], Androprosopa,
Chaoboridae [238], Dixa [66], Simulium [46], Dasyhelea [135], Psychodidae [45], Blephariceridae [63],
Therevidae [46], Tabanus [46], Odontomyia [46], Nannochorista [136], Panorpa [141] and Nosopsyllus
[139].
99.
Labral brush with dense field of hairs: (0) absent; (1) present, without specific arrangement;
(2) specifically arranged labral brush; (3) complex arrangement of different types of hairs; (4)
macrosetae. Dense fields or brushes of microtrichiae are present on the labrum of larvae of Tipuloidea
(e.g., Gonompeda, Cylindrotoma Macquart, 1834, Holorusia, less strongly developed Dicranota [46, 152,
153], Tipula [27]), Trichocera [45], Dasyhelea [135], Corethrella, Anisopodidae [45], Blephariceridae [63]
and Psychodidae [45]. Labral brushes are absent in Bibionidae [46, 75], Exechia, Therevidae [46],
Tabanus [46], Odontomyia [46], Nannochorista [136], Panorpa [141] and Nosopsyllus [139]. The labral
brushes consist of a complex arrangement of different types of hairs in Chironomus [74, 124]. In Dixa [66],
Simulium [46], Culiseta ([73]: fig. 19), Androprosopa, Chaoborus [238], Anopheles ([73]: fig. 25, [154]: LPi,
fig. 2a), Protanyderus ([32]: pb fig. 2) and Axymyia ([227]: fig. 5B) a labral brush with a specific
arrangement is present. The labrum of Bittacomorpha ([72]: lm, fig. 5) is bearing two tufts of longer setae
on the apex. The labral brush consists of macrotrichia in Deuterophlebia [48] and Nymphomyia [29].
100. Labral teeth: (0) very small or absent; (1) present. Strongly sclerotised labral teeth are usually
absent in Nymphomyia [29], Tipuloidea [e.g., 27, 46, 153], Trichocera [45], Androprosopa, Exechia,
Axymyia [227], Psychodidae [45], Dasyhelea [135], Blephariceridae [63], Therevidae [46], Odontomyia
[46], Panorpa [141] and Nosopsyllus [139], but occur in larvae of Cylindrotomidae (152) and Anisopodidae
[45]. Distinct teeth are absent in Deuterophlebiidae [48], Protanyderus [32], Bibionidae [46, 75],
Ptychopteridae [72], Chironomidae [74, 124], Culicidae [73, 154], Chaoborus [134, 238], Dixa [66],
Simulium [46], Tabanus [46] and Nannochorista [136].
101. Connection of torma with labral sclerite: (0) tormae firmly connected with labral sclerite; (1)
articulated. Tormae articulating with the labral sclerite occur in Tipulomorpha (e.g., Pediciidae,
Limoniinae) [16, 27, 45, 62], Culicidae [46, 73], Chironomidae [74, 124], Dixa [66], Simulium [46],
23
Androprosopa, Ceratopogonidae
[16],
Axymyia
[293],
Psychodidae
[45],
Blephariceridae
[63],
Nymphomyia [29] and Bittacomorpha [72]. The tormae are connected with the labral sclerite in Bibio [46,
75], Paenobremia [47], Anisopodidae [45], Nannochorista [136] and Panorpa [141]. After Courtney [48]
the tormae are fused articulated or fused with the anterolateral margin of the clypeolabrum in
deuterophlebiid larvae.
102. M. labroepipharyngalis (M. 7): (0) present; (1) absent. M. 7 is absent in Deuterophlebia [48],
Nymphomyia [29], Tipula [27], Dicranota [46], Bibio (75), Exechia, Paenobremia [47], Axymyia [227],
Dasyhelea [135], Anopheles [154], Culiseta [73], Androprosopa, Chaoborus [134, 238], Corethrella, Dixa
[66], Simulium [46], Odontomyia [46] and Therevidae [46]. The muscle is present in Bittacomorpha [72]:
lmcp, fig. 14), Protanyderus [32], Chironomidae [74, 124], Tabanus ([46]: labral compressor),
Nannochorista [136], Panorpa ([141]: M.labr.ep.) and Nosopsyllus ([139]: c.l.).
103. M. frontolabralis (M. 8): (0) present; (1) absent. M. 8 is absent in Deuterophlebia [48],
Nymphomyia [29], Protanyderus [32], Tipula [27], Dicranota [46], Bibio [75], Exechia, Paenobremia [47],
Axymyia [227], Bittacomorpha [72], Dasyhelea [135], Anopheles [154], Culiseta [73], Dixa [66], Simulium
[46], Androprosopa, Chironomidae [74, 124], Therevidae [46], Tabanus [46] and Odontomyia [46]. M.
frontolabralis is present in Chaoborus ([134]: M. abductorepipharyngis, [238]: M. frontolabralis),
Corethrella, Nannochorista [136], Panorpa ([141]: M.retr.labr.) and Nosopsyllus ([139]: a.l.).
104. M. frontoepipharyngalis (M. 9): (0) present; (1) absent. M. 9 is present in Deuterophlebia ([48]:
M61, M61’, M63), Nymphomyia ([29]: 9, fig. 5), Protanyderus [32], Tipula [27], Dicranota ([46]: messorial
muscle), Bittacomorpha ([72]: lml, lmm, fig. 14), Dixa ([66]: mto), Simulium ([46]: messorial muscle),
Corethrella, Dasyhelea ([135]: messm) Anopheles ([154]: EphH, LbM, dDil), Culiseta ([73]: messorial
muscle, median palatal muscle), Androprosopa, Chaoborus ([134]: M. adductorepipharyngis, [238]: M.
frontoepipharyngalis), Chironomidae [74, 124], Exechia, Axymyia ([227]: fig. 8C-F), Tabanus ([46]:
messorial muscle), Nannochorista [136] and Nosopsyllus ([139]: p.l.). It is absent in Bibio [75],
Paenobremia [47], Therevidae [46], Odontomyia [46] and Panorpa [141].
105. Movable premendible (messores): (0) absent; (1) present, separated from basal part by
weakly sclerotised zone; (2) present as strongly developed movable messorial arms. A movable
premandible is absent in Nymphomyia [29], Tipulidae and other subgroups of Tipulomorpha [27, 46], with
some possible exceptions in Limoniidae (Hexatominae, Eriopterinae [16]). The premandible is also absent
in Deuterophlebia [48], Protanyderus [32], Bibionidae [46, 75], Exechia, Axymyia [227], Chaoborus [134,
238], Corethrella, Blephariceridae [63], Therevidae [46], Tabanus [46], Odontomyia [46], Nannochorista
[136], Panorpa [141] and Nosopsyllus [139]. In Chironomidae [155], Androprosopa, Dasyhelea ([135]: fig.
11, mess), Dixa [66], Anisopodidae ([45]: fig. 19, prmd.), Psychodidae ([45]: fig. 48, pr.md.), Trichocera
[45] and Bittacomorpha ([72]: dmd, fig. 9) a premandible, which is separated from the basal part by a
weakly sclerotized zone, is present. Premendibles are present in Culicidae [46, 73, 154]. It is present as a
strongly developed messorial arm in Simulium [46].
106. Exposure of anterior epipharynx (cibarial roof): (0) not or only slightly exposed; (1) largely
exposed. The anterior epipharynx is largely exposed in Deuterophlebiidae [48], Nymphomyia [29],
24
Protanyderus [32], Tipulidae [27], Trichocera ([45]: fig. 7, ep.), Bibionidae [46, 75], Anisopodidae [45],
Psychodidae [45], some Blephariceridae ([63]: Fig. 4, Edwardsina), Axymyia [227], Ptychopteridae [72],
Chironomidae [74, 124], Corethrella, Anopheles [154], Dixa [66], Simulium [46], Odontomyia [46],
Therevidae [46] and Nannochorista [136]. The epipharynx is slightly exposed in Androprosopa. It is not
exposed in Chaoborus [134, 238], Dasyhelea [135], Dicranota [46], Tabanus [46], Panorpa [141] and
Nosopsyllus [139].
107. Position of antennal insertion: (0) lateral; (1) close to midline. The antenna inserted laterally in
Deuterophlebia [48], Nymphomyia [29], Protanyderus [32], Tipula [27], Limonia [65], Dicranota [46],
Trichocera [45], Bibio (75), Exechia, Paenobremia [47], Axymyia [227], Bittacomorpha [72], Dixa [66],
Simulium [46], Anopheles ([154]: fig. 1), Culiseta ([73]: fig. 19), Chironomidae [74, 124], Anisopodidae
[45], Psychodidae [45], Blephariceridae [63], Therevidae [46], Tabanus [46], Odontomyia [46],
Nannochorista [136], Panorpa [141] and Nosopsyllus [139]. In Androprosopa the antennal insertion is
slightly shifted to the midline, but still lateraly. The antenna inserted at the anterior margin of the head
capsule in Chaoborus (coaded as 1) [134, 238]. The antennal insertionis shifted closely to the midline in
Corethrella and Dasyhelea ([135]: fig. 6. ant).
108. Antennal segmentation: (0) with basal antennomere and one or several distinctly developed
distal segments; (1) appearing 1-segmented, distal segment vestigial or absent; (2) antenna
vestigial. The antenna von Tipula appears 1-segmented like in other larvae of Tipulidae, Cylindrotomidae,
Limoniinae [27, 46, 61, 152, 158], Trichocer1idae [45], Nymphomyia [29], Chironomidae [74, 124],
Culicidae [73, 154], Androprosopa, Chaoboridae [134, 238], Corethrella, Dixa [66], Dasyhelea [135],
Paenobremia [47], Bittacomorpha [72], Psychodidae [45], Blephariceridae [63], Therevidae [46], Tabanus
[46] and Odontomyia [46]. The antenna is completely missing in Bibionidae [46, 75], Exechia and Axymyia
[227]. The antenna of Deuterophlebia consists of a basal antennomere with a bifurcate distal article [48].
The antenna is composed of four segments in Simulium [46]. Whereas the antenna is two-segmented in
Anisopodidae [45], Dicranota [46], Protanyderus [32], Panorpa [141] and Nosopsyllus [139]. A threesegmented antenna is present in Nannochorista [136].
109. Distal antennal segment: (0) simple; (1) bifurcate. The distal antennal segment is simple in
larvae of Nymphomyia [29], Tipulidae [27], Limoniidae [65], Pediciidae [46], Trochoceridae [45],
Paenobremia [47], Anisopodidae [45], Chironomidae [74, 124], Dasyhelea [135], Culiseta [73],
Androprosopa, Chaoborus [134, 238], Dixa [66], Simulium [46], Bittacomorpha [72], Protanyderus [32],
Psychodidae [45], Blephariceridae [63], Therevidae [46], Tabanus [46], Odontomyia [46], Nannochorista
[136], Panorpa [141] and Nosopsyllus [139]. In Anopheles ([154]: fig. 1) and Corethrella the antenna is
simple with two thorns on the apex. In Deuterophlebia [48] the distal antennal segment is bifurcate.
110. Extrinsic antennal muscles: (0) present; (1) absent. Extrinsic antennal muscles are lacking in
Tipula like in other larvae of Tipulomorpha examined [27, 46] and are also missing in Nymphomyia [29],
Chironomidae [74], Androprosopa, Paenobremia [47], Therevidae [46] and Odontomyia [46]. One extrinsic
antennal muscle is present in Deuterophlebia ([48]: Mant), Protanyderus [32], Bittacomorpha ([72]: antm),
Dixa ([66]: mant), Anopheles [73, 154], Culiseta ([73]: Antennal muscle), Chaoborus ([134]: M. retractor
25
antennae, Corethrella, [238]: M5, M. tentorioscapalis posterior), Tabanus ([46]: antennal muscle),
Nannochorista ([136]: M1/2/3/4) and Panorpa ([141]: M.d.sc.). Three extrinsic antennal muscles are
present in Nosopsyllus ([139]: d.a., l.a., f.a.).
111. Mandibulo-maxillary complex: (0) absent; (1) present. A mandibulo-maxillary complex is absent
in all nematoceran lineages [e.g., 27, 29, 32, 45, 48, 66, 71-75, 124, 134, 135, 154, 164, 227, 238],
Nannochorista [136], Panorpa [141] and Nosopsyllus [139]. The presence [e.g., 46] is likely an
autapomorphy of Brachycera [71]. It is present in Therevidae ([46]: fig. 31C), Tabanus ([46]: fig. 27) and
Odontomyia ([46]: fig. 35A).
112. Plane of operation of mandibles: (0) horizontal or slightly oblique; (1) distinctly oblique or
vertical; (2) horizontal in first larval stage and oblique or vertical in later instars. The plane of
operation is horizontal in Tipulidae (e.g., Tipula, Tanyptera; [8, 27]), Trichocera [45], Protanyderus [32],
Exechia, Axymyia [227], Paenobremia [47], Nannochorista [136] and Panorpa [141]. The axis of
movement is slightly oblique in Bibio ([46], coaded as 0), Corethrella and Nosopsyllus [139]. The plane of
operation is horizontally in some Chironomidae, whereas the plane is oblique or nearly vertical in other
chironomid larvae [155], and also in Deuterophlebia [48], Nymphomyia [29], Bittacomorpha [72],
Dasyhelea [135], Androprosopa, Anisopodidae [45], Psychodidae [45], Blephariceridae [63], Therevidae
[46] and Odontomyia [46]. The plane is oblique in Dixa [66], Simulium [46], Anopheles [73, 154],
Chaoborus [134, 238], Culiseta [73] and Tabanus [46]. The plane of operation is horizontal or slightly
oblique in Dicranota [46].
113. Position of secondary mandibular joint: (0) not shifted posteriorly of the antennal foramen;
(1) shifted posteriorly of the antennal foramen. The unusual position of the secondary mandibular joint
posterad of the antennal articulation area is a potential autapomorphy of Tipulidae [27]. This condition is
not found in the other groups of Tipulomorpha [e.g., 46, 152]. The mandibular join is not shifted or slightly
shifted posteriorly in Deuterophlebia [48], Nymphomyia [29], Protanyderus [32], Trichocera [45],
Bibionidae [46, 75], Exechia, Axymyia [227], Paenobremia [47], Ptychopteridae [72], Anisopodidae [45],
Psychodidae [45], Blephariceridae [63], Dasyhelea [135], Chironomidae [74, 124], Culicidae [73, 154],
Androprosopa, Chaoborus [134, 238], Dixa [66], Simulium [46], Odontomyia [46], Therevidae [46],
Nannochorista [136], Panorpa [141] and Nosopsyllus [139].
114. Shape of mandible: (0) without distinctly elongated distal part; (1) distal part elongated,
sickle-shaped. The distal part of the mandible is short and stout in Deuterophlebia [48], Nymphomyia
[29], Protanyderus [32], Tipula [27], Limonia [62], Trichocera [45], Bibio [46, 75], Exechia, Axymyia [227],
Paenobremia [47], Bittacomorpha [72], Dixa [66], Dasyhelea [135], Simulium [46], Culicidae [73, 154],
Androprosopa, Chaoborus [134, 238], Corethrella, Chironomidae [74, 124], Anisopodidae [45],
Psychodidae [45], Blephariceridae [63], Therevidae [46], Tabanus [46], Odontomyia [46], Panorpa [141]
and Nosopsyllus [139]. The mandible is sickle-shaped and elongated in Dicranota [46]. The mandible is
slender and elongated in Nannochorista ([136]: Fig. 8A, B).
115. Movable lacinia mobilis: (0) absent; (1) present. An articulated lacinia mobilis is present in Tipula
[27], Chaoborus ([134, 238]: lm, fig. 5C) and Nannochorista ([136]: lcm, fig. 8A). It is absent in
26
Deuterophlebia ([48]: mdp), Nymphomyia [29], Chironomidae [155], Dasyhelea [135], Anopheles [154],
Culiseta [73], Dixa [66], Corethrella, Limonia [27], Dicranota [46], Trichocera [45], Bibio [46, 75], Exechia,
Paenobremia [47], Simulium [46], Anisopodidae [45], Tabanus [46], Therevidae [27], Bittacomorpha [72],
Protanyderus [32], Psychodidae [45], Blephariceridae [63], Odontomyia [46], Panorpa [141] and
Nosopsyllus [139].
116. Anteriorly directed cone of mesal mandibular edge: (0) absent; (1) present. An anterior
directed cone of the mandibular edge is absent in the larvae of Deuterophlebia [48], Nymphomyia [29],
Protanyderus [32], Tipula [27], Limonia (16), Dicranota [46], Bibio [46, 75], Exechia, Axymyia [227],
Paenobremia [47], Bittacomorpha [72], Dixa [66], Simulium [46], Dasyhelea [135], Culicidae [73, 154],
Androprosopa, Chaoborus [134, 238], Corethrella, Chironomidae [74, 124], Blephariceridae [63],
Therevidae [46], Tabanus [46], Odontomyia [46], Nannochorista [136], Panorpa [141] and Nosopsyllus
[139]. The cone is present in Anisopodidae ([45]: figs. 26, 29, kr.), Trichocera [261] and some
Psychodidae [261],
117. Mandibular comb on dorsal surface: (0) absent; (1) present. A mandibular comb is absent in
Tipula and in larvae of the other groups of Tipulomorpha [e.g., 27, 62, 152, 153], Trichocera [45],
Nymphomyia [29] and also in Bibionidae [46, 75], Exechia, Axymyia [227], Paenobremia [47], Dasyhelea
[135], Corethrella, Androprosopa, Protanyderus [32], Blephariceridae [63], Therevidae [46], Tabanus [46],
Odontomyia [46], Nannochorista [136], Panorpa [141] and Nosopsyllus [139], but it is present in
Deuterophlebia [48], Bittacomorpha ([72]: mdbr), Chironomidae [74, 124], Culicidae [73, 154], Chaoborus
[134, 238], Dixa [66], Simulium [46] and Anisopodidae [45].
118. Apical multitoothed mandibular comb: (0) absent; (1) present (at least in instar 1). An apical
multitoothed mandibular comb is absent in most groups of lower Diptera [27, 32, 45, 47, 66, 72-75, 124,
134, 135, 154, 173, 227] and also in Tabanus [46], Odontomyia [46], Nannochorista [136], Panorpa [141]
and Nosopsyllus [139]. It is present in Deuterophlebia [48], Nymphomyia [29], Androprosopa and some
Blephariceridae [16].
119. Subdivision of mandible: (0) absent; (1) present, distal part separated by a furrow; (2)
present, distal part separated from basal part by weakly sclerotised zone. A movable distal part of
the mandible is absent in Deuterophlebiidae [48], Nymphomyia [29], Protanyderus [32], Tipulidae [27],
Limoniidae [16], Pediciidae [46], Trichoceridae [45], Bibionidae [46, 75], Exechia, Axymyia [227],
Paenobremia [47], Dasyhelea [135], Culicidae [73, 154], Androprosopa, Chaoborus [134, 238],
Corethrella, Dixa [66], Simulium [46], Blephariceridae [63], Nannochorista [136], Panorpa [141] and
Nosopsyllus [139]. Whereas, it is present in Bittacomorpha [72] and Anisopodidae [45]. The mandible of
Tabanus [46] and Odontomyia [46] is subdivided in a distal and a basal part.
120. Insertion of adductor tendon: (0) nor or very slightly shifted anteriorly; (1) distinctly shifted
anteriorly. The mesal wall of the mandible is distinctly shortened in larvae of Tipulidae and
Cylindrotomidae (152), and this is probably a general feature in larvae of Tipulomorpha ([45]: fig. 38, [62]:
figs. 61, 128, [27]: figs. 4A, B); it also inserted distinctly anteriorly in Anisopodidae ([45]: figs. 26-31,
ap.ad.), Psychodidae [45], Dasyhelea [135], some Blephariceridae [63] and Nosopsyllus ([139]: fig. 6,
27
ad.a.). Both tendons insert approximately at the same level in Nymphomyia [29], Culicidae [73, 154],
Androprosopa, Chaoborus [134, 238], Chironomidae [46], Axymyia [227], Paenobremia [47], Tabanus
[46], Nannochorista [136] and Panorpa [141]. The adductor tendon is also shifted anteriorly in
Deuterophlebia ([48]: M21, fig. 4), Bittacomorpha ([72]: mdf, fig. 9), Dixa ([66]: see fig. 18, tamdb),
Simulium ([46]: mandibular adductor muscle), Therevidae [46] and Odontomyia ([46]: mandibular adductor
apodeme). The adductor tendon is not shifted anteriorly in Protanyderus [32].
121. M. tentoriomandibularis (M. 13): (0) present; (1) absent. M. tentoriomandibularis is absent in
Deuterophlebia [48], Nymphomyia [29], Protanyderus [32], Tipula like in all other examined larvae of
Diptera [e.g., 27, 47, 66, 72-75, 124, 134, 135, 154, 227, 238], Panorpa [141] and also in Nosopsyllus
[139]. M. tentoriomandibularis is present in Nannochorista [136].
122. Cardo: (0) mainly membranous; (1) in part represented by a clearly delineated sclerite; (2)
absent. The cardo is a distinctly developed sclerite in Tipula [27], Limonia [65], Trichocera [45],
Protanyderus [32], Bibio [46, 75], Bittacomorpha ([72]: cd, fig. 7), Dixa [66], Anisopodidae ([45]: fig. 20,
car.), Therevidae [46], Tabanus [46], Nannochorista [136], Panorpa [141] and Nosopsyllus [139]. It is a
small sclerite in Dicranota [46] and Blephariceridae [63]. The cardo is absent or completely fused with the
head capsule in Nymphomyia [29], Chironomus [74], Anopheles [73], Culiseta [73] and Axymyia [227]. It is
mainly membranous in Deuterophlebia [48]. The cardosclerite is fused with the head capsule in
Psychodidae ([45]: fig. 17, car.).
123. Endite lobes: (0) distinctly developed and separated; (1) partly fused, still recognisable as
separate structures; (2) galea and stipes completely fused or absent. Separate endite lobes were
described for Tipula species by Selke ([61]: fig. 23) but are not recognisable in Tipula montium ([27]: Fig.
4C). Separate endite lobes are present in Protanyderus [32], Anisopodidae [45], Blephariceridae ([63]: fig.
25, lac, gal) and Nannochorista [136]. Both are absent or completely fused in Nymphomyia [29], Bibio [46,
75], Paenobremia [47], Limonia [65], Dicranota [46], Trichocera [174], Bittacomorpha [72], Dixa [66],
Simulium [46], Culicidae [73, 154], Chaoborus [134, 238], Chironomidae [155], Psychodidae [45],
Therevidae [46] and Odontomyia [46]. In Dasyhelea ([135]: lob); Deuterophlebia [48], Tabanus [46] and
Nosopsyllus [139] only one maxillary endite is present. The endites are partly fused, but recognizable as
separate structures in Axymyia [227] and Panorpa [141].
124. Number of maxillary palp segments: (0) 3 or more; (1) 2; (2) 1. The maxillary palp is onesegmented in Nymphomyia [29], Tipula-larvae [27], Limonia [45] and also in larvae of Dicranota [46],
Trichocera [45], Paenobremia [47], Bittacomorpha [72], Protanyderus [32], Dixa [66], Anopheles [73, 154],
Culiseta [73], Chaoborus [134, 238], Corethrella, Dasyhelea ([135]: plm), Simulium [46], Anisopodidae
[45], Psychodidae [45], Blephariceridae [63],Tabanus [46] and Odontomyia [46]. It is two-segmented in
larvae of Bibionidae [46, 75], Chironomus [162], Deuterophlebia [48], Axymyia [227], Therevidae [46] and
Nosopsyllus [139]. The palp is three-segmented in Nannochorista [136]. The maxillary palp bears 5
segments in Panorpa [141]. Palps are completely absent in Exechia.
125. Length of single maxillary palpomere: (0) not elongated, about as long as wide; (1) distinctly
elongated. The maxillary palpomere is not elongated in Deuterophlebia [48], Nymphomyia [29],
28
Protanyderus [32], Tipula [27], Limonia [45], Trichocera [45], Bibio [46, 75], Axymyia [227], Paenobremia
[47], Bittacomorpha [72], Dixa [66], Dasyhelea [135], Simulium [46], Anopheles [73, 154], Culiseta [73],
Chaoborus
[134, 238],
Corethrella, Chironomus
[162],
Anisopodidae
[45],
Psychodidae
[45],
Blephariceridae [63], Therevidae [46], Tabanus [46], Odontomyia [46], Nannochorista [136], Panorpa [141]
and Nosopsyllus [139]. It is distinctly elongated in Dicranota [46].
126. M. craniocardinalis (M. 15): (0) well developed; (1) absent. M. 15 is absent in Deuterophlebia
[48], Nymphomyia [29], Protanyderus [32], Tipula [27], Dicranota [46], Bibio (75), Axymyia [227],
Paenobremia [47], Bittacomorpha [72], Dixa [66], Dasyhelea [135], Simulium [46], Culicidae (73; 154),
Chaoborus (134; 238), Chironomidae (74; 124), Therevidae [46], Tabanus [46], Odontomyia [46],
Nannochorista [136] and Panorpa [141]. The muscle is present in Nosopsyllus (139 [ab.m.]). One
maxillary muscle is present in Exechia, but it is not clear if it is homologous with M. cariocardinalis. In
Androprosopa two maxillary muscles with unknown homology are present.
127. M. tentoriocardinalis (M. 17) with origin from posterior tentorial arm: (0) present; (1) absent.
M. 17 is absent in Protanyderus [32], Tipula [27], Dicranota [46], Bibio (75), Paenobremia [47],
Bittacomorpha [72], Dixa [66], Dasyhelea [135], Simulium [46], Anopheles (73; 154), Culiseta (73),
Chaoborus (134; 238), Chironomidae (74; 124), Therevidae [46], Tabanus [46], Odontomyia [46], Panorpa
[141] and Nosopsyllus [139]. The muscle has its origin on the ventrolateral cranium in Deuterophlebia [48]
and Nymphomyia [29], and on the ventral wall of the head capsule in Axymyia [227]. The muscle has its
origin on the anterior tentorial arm in Nannochorista [136].
128. M. tentoriostipitalis (M. 18) with origin from posterior tentorial arm: (0) present; (1) absent. M.
18 is absent in Deuterophlebia [48], Protanyderus [32], Tipula [27], Dicranota [46], Bibio (75),
Paenobremia [47], Bittacomorpha [72], Dixa [66], Dasyhelea [135], Simulium [46], Anopheles (73; 154),
Culiseta (73), Chaoborus (134; 238), Chironomidae (74; 124), Therevidae [46], Tabanus [46], Odontomyia
[46] and Nosopsyllus [139]. The muscle has its origin on the ventral head capsule in Axymyia [227]. The
muscle is present with two subcomponents in Panorpa [141], one component has its origin on the
tentorium (M.add.st.1) and the other subcomponent on the head capsule (M.add.st.2). The muscle has its
origin on the anterior tentorial arm in Nannochorista [136]. The muscle originate on the head capsule in
Nymphomyia [29], but it is not entre clear if it is M. tentoriostipitalis or M. craniolacinialis, because the
maxillary endites are not recognizable as separate structures.
129. Muscles of the endite lobes (Mm. 20/21): (0) 2; (1) 1; (2) absent. Intrinsic maxillary muscles of
the galea and lacinia (Mm. 20/21) are missing in dipteran larvae (e.g., 73, 1949; 154, 1950; 72; 124; 75;
66; 47; Hennig 1973; 48; 135; 29; 32; 227) and also in Nannochorista [136]. One muscle is present in
Panorpa (141 [M.prom.lac.]) and Nosopsyllus (139 [l.s.]).
130. Anterior teeth of hypostomium: (0) absent; (1) present. A hypostomium equipped with triangular
teeth on the anterior margin is present in Nymphomyia [29], Protanyderus (32 [hp, fig. 1B]), Tipula [27],
Limonia [62], Dicranota [46], Anopheles (154), Culiseta (73), Corethrella and Chironomus (74). Anterior
teeth of the hypostomium are absent in Deuterophlebia [48], Trichocera [45], Bibionidae (46; 75), Exechia,
Axymyia [227], Dasyhelea [135], Chaoborus (134; 238), Androprosopa, Therevidae [46], Tabanus [46],
29
Odontomyia [46], Nannochorista [136] and Panorpa [141]. A toothed hypostomium is present in
Ptychopteridae [27] and Simulium [46].
131. Median division of hypostomium: (0) absent; (1) partly divided; (2) completely divided. The
hypostomium or hypostomal plate is medially partly divided by an incomplete triangular zone of weakness
in Tipula [27], Limonia (Lindner 1959: fig. 10, “Hypostomium”), it is also partly devided in Dixa (66; see fig.
24, 25). It is not divided in Deuterophlebia [48], Nymphomyia [29], Protanyderus [32], Trichocera [45],
Bibionidae (46; 75), Ptychopteridae [72], Culicidae (73; 154), Simulium [46], Androprosopa, Chaoborus
(134; 238), Corethrella, Chironomidae (74; 124), Anisopodidae (45, b), Psychodidae [45], Exechia,
Axymyia [227], Therevidae [46], Tabanus [46], Odontomyia [46] and Nannochorista [136]. The
hypostomium is completely divided (or reduced) in Dicranota [46].
132. Separate submentum: (0) present; (1) absent. A separate submentum is absent in
Deuterophlebia [48], Nymphomyia [29], Protanyderus [32], Tipula, Limonia [27], Dicranota [46], Trichocera
[45], Bibio (46; 75), Exechia, Axymyia [227], Paenobremia [47], Bittacomorpha [72], Dasyhelea [135],
Anopheles (154), Culiseta (73), Androprosopa, Chaoborus (134; 238), Corethrella, Dixa [66],
Blephariceridae [63], Nannochorista [136] and Nosopsyllus [139]. But present in Chironomus (74),
Simulium (46 [Fig. 9B]), Therevidae (46 [fig. 29C]), Tabanus [46] and Odontomyia (46 [fig. 32B]). The
submentum is present, but fused with the head capsule in Panorpa (141 [Subm., fig. 9]). It is present in
Olbiogaster (Anisopodidae) (Anthon 1943b) and some Psychodidae (16).
133. Hypostomal plate: (0) fused with elements of the head capsule; (1) present as a separate
structure. The hypostomal plate is fused with elements of the head capsule in Nymphomyia [29], Tipula
[27], Limonia [27], Dicranota [46], Dixa [66], Dasyhelea [135], Simulium [46], Anopheles (154), Culiseta
(73), Chaoborus (134; 238), Androprosopa, Corethrella, Chironomidae [155], Anisopodidae [45],
Psychodidae [45], Blephariceridae [63], Axymyia [227] and Nannochorista [136]. It not fused in
Protanyderus [32], Bibio (46; 75) and Bittacomorpha (72 [hst, fig. 6]).
134. Premental teeth: (0) present; (1) indistinct or absent. The prementum bears five teeth at its
anterior margin in Tipula [27], Anopheles (154 [fig 9b]), Culiseta (73 [fig. 23D]) and they are also present in
Nymphomyia [29], Chironomidae (74; 124), Androprosopa, Trichocera [45] and Therevidae [46]. In
Tabanus [46] the prementum bears two teeth. Premental teeth are absent in the larvae of Deuterophlebia
[48], Protanyderus [32], Bibio (46; 75), Axymyia [227], Anisopodidae (45, b), Psychodidae [45],
Blephariceridae [63], Dicranota [46], Chaoborus (134; 238), Corethrella, Dixa [66] and Nosopsyllus [139].
Premental teeth are not visible in Dasyhelea [135], Simulium [46], Odontomyia [46], Nannochorista [136]
and Panorpa [141].
135. Labial palp segments: (0) more than one segment; (1) 1-segmented, more or less vestigial.
Labial palps are completely absent in larvae of Nymphomyia [29], Tipulomorpha (46; 27), Trichoceridae
[45], larvae of Chironomidae (74; 124), Culicidae (73; 154), Androprosopa, Dasyhelea [135] and also in
larvae of Deuterophlebia [48], Protanyderus [32], Bibio (46; 75), Exechia, Axymyia [227], Chaoborus (134;
238), Corethrella, Dixa [66], Therevidae [46] and Odontomyia [46]. The labial palps are more ore less
vestigial in Psychodidae [45] and Paenobremia [47]. After Kramer (1954) the labial palps in Bittacomorpha
30
are much reduced and possibly represent as paired apical projections (lbp, 72). The prementum bears two
small lobes in Simulium [46], which are possibly representing the labial palps. The palps are onesegmented in Anisopodidae (45: fig. 19, la.p.; Anthon 1943b: fig. 7, la.p.), Blephariceridae (63: fig. 26,
lb.p.), Tabanus [46], Nannochorista [136] and Nosopsyllus (139 [l.b., fig. 8]). The palps are 3-segmented
in Panorpa (141 [P.lb., fig. 9]).
136. M. submentopraementalis (M. 28): (0) present; (1) absent. M. 28 is absent in Deuterophlebia
[48], Nymphomyia [29], Protanyderus [32], Tipula [27], Dicranota [46], Bibio (46; 75), Exechia, Axymyia
[227], Paenobremia [47], Bittacomorpha [72], Dixa [66], Dasyhelea [135], Simulium [46], Anopheles (154),
Culiseta (73), Chaoborus (134; 238), Androprosopa, Corethrella, Chironomidae (74; 124), Therevidae
[46], Tabanus [46], Odontomyia [46], Nannochorista [136], Panorpa [141] and Nosopsyllus [139].
137. Tentorio-premental muscles (Mm. 29/30): (0) 2; (1) 1; (2) extrinsic labial muscles are absent.
One extrinsic labial muscle (M. 29) is present in Deuterophlebia (48 [Mlm]), Nymphomyia (29 [pr]),
Protanyderus [32], Tipula [27], Dicranota (46 [labial muscle]), Bibio (46; 75 [rhy]), Paenobremia (47
[MML]), Bittacomorpha (72 [lbm]), Dixa (66 [mrtlb]), Dasyhelea (135 [78]), Simulium (46 [labial adductor
muscle]), Anopheles (154 [Fl.hph]), Culiseta (73 [labial adductor muscle]), Chaoborus (134 [M.
adductorlabii]; 238 [M13, M.tentoriopraementalis inferior/superior]), Chironomus (74), Odontomyia (46
[labial muscle]) and Nosopsyllus (139 [p.l.a.]). Extrinsic labial muscles are absent in Exechia, Axymyia
[227], Androprosopa, Corethrella, Therevidae [46], Tabanus [46], Nannochorista [136] and Panorpa [141].
138. M. praementopalpalis internus (M. 34): (0) present; (1) absent. M. 34 is absent in
Deuterophlebia [48], Nymphomyia [29], Protanyderus [32], Tipula [27], Dicranota [46], Bibio (46; 75),
Exechia, Axymyia [227], Paenobremia [47], Bittacomorpha [72], Dixa [66], Dasyhelea [135], Simulium [46],
Anopheles (154), Culiseta (73), Androprosopa, Chaoborus (134; 238), Corethrella, Chironomus (74),
Therevidae [46], Tabanus [46], Odontomyia [46], Nannochorista [136], Panorpa [141] and Nosopsyllus
[139].
139. M. hypopharyngosalivarialis (M. 37): (0) present; (1) absent. M. 37 is present in Protanyderus
[32], Tipula [27], Axymyia [227], Androprosopa, Corethrella, Bittacomorpha (72 [sldm]), Exechia,
Nannochorista (136 [37, fig. 11]), Panorpa (141 [M.dil.spp.]) and Nosopsyllus (139 [d.s.m.]), but absent in
Deuterophlebia [48], Nymphomyia [29], Bibio (46; 75), Paenobremia [47], Dicranota [46], Anopheles (154),
Culiseta (73), Chaoborus (134; 238), Dixa [66], Dasyhelea [135] and Simulium [46].
140. M. tentoriohypopharyngalis (M. 42): (0) present; (1) absent. The muscle is generally absent in
Diptera (e.g., 46; 154, 1950; 72; 75; 66; 47; Denis & Bitsch 1973; 48; 135; 27; 238; 29; 32; 227), in
Panorpa [141] and Nosopsyllus [139]. Whereas it is present in Nannochorista (136 [42, fig. 11]).
141. Pharyngeal filter: (0) present; (1) absent. A pharyngeal filter is absent in larvae of Deuterophlebia
[48], Nymphomyia [29], Tipula [27], Limonia (16), Dicranota [46], Bibio (75), Paenobremia [47], Dasyhelea
[135], Simulium [46], Chironomidae (74; 124), Androprosopa, Anisopodidae [45], Nannochorista [136],
Panorpa [141] and Nosopsyllus [139]. But it is present in Culicidae (73; 154), Bittacomorpha [72],
Trichocera (16), Protanyderus [32], Blephariceridae [63] and Chaoboridae (238). It is present Axymyia
[227]. A pharyngeal filter is present in Psychodinae, but absent in other Psychodidae (16).
31
142. M. tentoriobuccalis anterior (M. 48): (0) present; (1) absent. M. 48 is absent in Deuterophlebia
[48], Nymphomyia [29], Protanyderus [32], Tipula [27], Dicranota [46], Anopheles (154), Culiseta (Cook
1944), Chaoborus (134; 238), Dixa [66], Dasyhelea [135], Simulium [46], Exechia, Therevidae [46],
Tabanus [46] and Odontomyia [46]. The muscle is present in Bibio (75[dvphy]), Axymyia [227],
Androprosopa, Corethrella, Bittacomorpha (72 [vphm1, vphm2]), Nannochorista (136 [48, fig. 11A]),
Panorpa (141 [M.dil.l.phar.]) and Nosopsyllus (139 [part of v.d.a.]).
143. M. verticopharyngalis (M. 51): (0) present; (1) absent. M. 51 is absent in Nymphomyia [29],
Tipula [27], Dicranota [46], Simulium [46], Culiseta (73), Corethrella, Androprosopa, Exechia, Therevidae
[46], Tabanus [46] and Odontomyia [46]. The muscle is present in Deuterophlebia (48 [M86]),
Protanyderus [32], Bibio (75 [ddphy]), Axymyia [227], Paenobremia (47 [part of MDF]), Bittacomorpha (72
[phd3]), Dasyhelea (135 [oesdilm]), Dixa (66 [dlpphy]), Anopheles (154 [not mentioned in the text, but
shown in illustrations]), Chaoborus (238 [M15, M. verticopharyngalis]), Nannochorista (136 [51, fig. 11])
and Panorpa (141 [M. par.phar.ant. and post., M.occ.phar.]).
144. Posterior ventral pharyngeal dilators: (0) anterior and posterior subcomponent (M. 50 and M.
52): (1) one bundle; (2) absent. A posterior pharyngeal pump is absent in Nymphomyia [29], Tipula [27],
Dasyhelea [135], Dicranota [46], Exechia, Therevidae [46], Tabanus [46] and Odontomyia [46]. One
muscle bundle is present in Deuterophlebia (48 [M87-90]), Bibio (75 [dvphy]), Simulium (46 [ventral
pharyngeal muscle]), Androprosopa, Corethrella and Culiseta (73 [ventral pharyngeal muscle]). Both
muscles are present in Bittacomorpha (72 [phd3, vphm3]), Paenobremia (47 [MDP]), Protanyderus [32],
Dixa (66 [dvphy2]), Anopheles (154 [v.Dil. and M.51 is not mentioned in the text, but shown in
illustrations]), Chaoborus (134; 238), Nannochorista (136 [50, 52, fig. 11]), Panorpa (141 [M.tent.phar.sup.
und inf., M.gul.phar.]) and Nosopsyllus (139 [v.d.a., v.d.p.]). One bundle is present in Axymyia, but it is not
entirely clear if it is a part of M. 50 or M. 52 [227].
145. Position of brain: (0) completely or largely within head capsule; (1) partly shifted to thorax;
(2) completely shifted to thorax. The brain is completely shifted into the prothorax in Nymphomyia [29],
Tipula [27], Dicranota [46], Axymyia [227], Androprosopa, Therevidae [46], Tabanus [46] and Odontomyia
[46]. The brain is partly shifted into the prothorax in Chironomidae (46; 124), Bibio [27], Exechia and
Protanyderus [32]. The entire brain lies completely within the prothorax in Dixa [66]. The brain remains
completely within the head capsule in Deuterophlebia [48], Bittacomorpha [72], Culicidae (73), Chaoborus
(238 [b, 6B]), Corethrella, Nannochorista (136 [br, sog, fig. 11A]), Panorpa [141] and Nosopsyllus (139
[br., s.g., fig. 9]). The brain is partly shifted into the prothorax in larvae of Simulium [27].
146. Cibariopharyngeal sclerotisation: (0) absent; (1) present. A structural unit with a trough-like,
sclerotised ventral wall is formed by the cibarium and pharynx in the brachyceran groups examined by
Cook (1949). This condition is absent larvae of lower Diptera examined (e.g., 73, 1949; 72; 75; Denis &
Bitsch 1973; 48; 27; 238; 29; 32; 227) and also in Nannochorista [136], Panorpa [141] and Nosopsyllus
[139]. It is present in Therevidae [46], Tabanus [46] and Odontomyia [46].
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