A new phylogeny and phylogenetic classification for the Canthyloscelidae (Diptera: Psychodomorpha)

A new phylogeny and phylogenetic classificationfor the Canthyloscelidae (Diptera:Psychodomorpha)

Dalton de Souza Amorim

Abstract: A new phylogeny and phylogenetic classification for the Canthyloscelidae (Diptera: Psychodomorpha) is pre-sented. A phylogenetic analysis of the Scatopsoidea is performed. A sister-group relationship between theCanthyloscelidae and Scatopsidae is accepted and the monophyly of the Canthyloscelidae is corroborated, including thegeneraExiliscelis, Synneuron, Hyperoscelis, andCanthyloscelis. An earlier phylogenetic analysis of the group is con-sidered, in whichSynneuronwas accepted as the sister-group of the Scatopsidae andExiliscelis was considered thesister-group ofSynneuron+ Scatopsidae. Some apomorphic similarities between the larvae of all genera ofCanthyloscelidae, especially the reduction of the head capsule, are considered true synapomorphies.Exiliscelis is con-sidered the sister-group of the rest of the family and is placed in a new subfamily, Exiliscelinae. In theCanthyloscelinae,Synneuronis the sister-group ofHyperoscelis+ Canthyloscelis. A phylogenetic classification of thegroup is proposed.Prohyperoscelis rohdendorfiKovalev, 1985, from the Middle Jurassic in Russia, is accepted as thesister-group ofCanthyloscelis.

Résumé: Une nouvelle classification phylogénique et phylogénétique pour les Canthyloscelidae (Diptera: Psycho-domorpha) est présentée. Nous avons procédé à une analyse phylogénétique des Scatopsoidea. Les Canthyloscelidae etles Scatopsidae sont déclarés groupes-soeurs et le monophylétisme des Canthyloscelidae est confirmé; le groupe com-prendExiliscelis, Synneuron, Hyperosceliset Canthyloscelis. Une analyse phylogénétique antérieure a été réexaminée;Synneurony est considéré comme le groupe-soeur des Scatopsidae etExiliscelis, comme le groupe-soeur deSynneuron+Scastopsidae. Certaines similarités apomorphiques entre les larves de tous les genres de Canthyloscelidae, particulière-ment la réduction de la capsule céphalique, semblent être de véritables synapomorphies.Exiliscelis est considérécomme le groupe-soeur du reste de la famille et classifié dans une nouvelle sous-famille, les Exiliscelinae. Chez lesCanthyloscelinae,Synneuronest le groupe-soeur deHyperoscelis+ Canthyloscelis. Une classification phylogénétique dugroupe est proposée.Prohyperoscelis rohdendorfiKovalev, 1985, du milieu du Jurassique de Russie, est reconnucomme le groupe-soeur deCanthyloscelis.

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AmorimIntroduction

There has been virtually no dispute that the Scatopsoideaconstitutes a monophyletic group within the Diptera. Mostauthors have placed the group close to other families in theBibionomorpha (Enderlein 1912; Edwards 1925; Hennig 1948,1954, 1973; Amorim 1993). Hutson (1977) did not particu-larly address the problem of the placement of the Scato-psoidea within the Diptera, but his comparisons of larvalfeatures included only families in the Bibionomorpha.Recently, Wood and Borkent (1989) proposed a quite newapproach based on larval features, and suggested that theScatopsoidea should be transferred to the Psychodomorpha.This position was later accepted by Amorim (1994).

During the nineteenth century and at the beginning of thelast century the first known genera of Canthyloscelidae wereconsidered part of the Scatopsidae. The inclusion of canthylo-scelid genera in a taxon with family status was first pro-

posed by Enderlein (1912), who gave subfamilial status toCorynoscelis(=Hyperoscelis). Later, Enderlein (1936) rankedthe taxon as a family and erected Synneuridae. Considerableconfusion concerning name priority has affected the historicaltaxonomy of the group (see Hutson 1977; but also Nagatomiand Saigusa 1984: 463–464) at the generic level and familylevel. Corynoscelidae is an invalid name. Other junior syn-onyms or invalid names have been proposed for the group(Hyperoscelidae, Hyperoscelididae, Synneurontidae, etc.).Hutson’s (1977) review solved most of these problems andindicated that Synneuridae Enderlein, 1936 would be theoldest valid family-group name and Canthyloscelidae Rohden-dorf, 1951 would also be a valid name if one separates thesegenera into two taxa with family rank. However, Nagatomiand Saigusa (1984), in their review of JapaneseHypero-scelis, followed Dr. Curtis Sabrosky’s opinion (personalcommunication in a letter) that the oldest valid family-groupname proposed for the taxon is not Synneuridae Enderlein,1936 but Canthyloscelidae, proposed by Shannon (1927),which was largely overlooked in the literature.

Two main taxonomic groups other than the Scatopsidaehave been identified and given family status in Scatopsoidea.One is composed ofHyperoscelisHardy and Nagatomi, 1960(a new name for the preoccupied generic nameCorynoscelis

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Received May 20, 1999. Accepted December 9, 1999.

D.S. Amorim. Departamento de Biologia, Faculdade deFilosofia, Ciências e Letras de Ribeirão Preto, Universidadede São Paulo, Avenida Bandeirantes 3900, 14040-901Ribeirão Preto SP, Brazil (e-mail: [email protected]).

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Boheman) andCanthyloscelisEdwards, 1922, collectivelynamed Canthyloscelidae (referred to as Canthyloscelididaeby some authors). The other includesSynneuronLundström,1910 andExiliscelisHutson, 1977, named Synneuridae. Thebest review of the taxonomic history of the group was madeby Hutson (1977) and does not need to be repeated here.

Hutson (1977) proposed a phylogeny for the Scatopsoideaand concluded thatCanthyloscelis+ Hyperoscelis wouldcorrespond to the sister-group ofSynneuron+ Exiliscelis +Scatopsidae. In his classification, Canthyloscelidae includesthe first two of these genera, while Synneuridae includesSynneuronand Exiliscelis. Hutson’s (1977) classification,however, has Synneuridae as a paraphyletic group. The in-clusion ofSynneuronandCanthyloscelisin different familieswas proposed earlier by Enderlein (1936) and accepted byRohdendorf (1938, 1964), Hennig (1960), Mamaev and Krivo-

sheina (1969), Peterson and Cook (1981), and Evenhuis (1994).Hennig (1954) and Cook (1963) did not segregate the twogroups into separate families.

More recently, Wood and Borkent (1989) considered itmore likely that the genera included in Synneuridae andCanthyloscelidae belong to a monophyletic group, whichshould be included in a single family, Synneuridae. Larvaeof Canthyloscelis, Hyperoscelis, andSynneuronhave a greatlyreduced, membranous head capsule, with largely unsclerotizedmouthparts. The larva ofExiliscelis is still unknown. Woodand Borkent’s (1989) argument is straightforward: the Scato-psidae have larvae with a fully developed head capsule, so ifwe accept Hutson’s (1977) cladogram we must admit two orthree (assuming thatExiliscelisconforms to the other generain this aspect) independent reductions in the sclerotization ofthe larval head capsule. I agree with Wood and Borkent(1989) that it is quite unlikely that Synneuridae sensu Woodand Borkent (1989) is paraphyletic. Hutson’s (1977) discus-sion is nevertheless very useful in furnishing much informa-tion on character evolution, as well as for his solutions forthe nomenclatural problems of the group. This paper is ananalysis of the Scatopsoidea as a whole, with particular em-phasis on the generic relationships of the Canthyloscelidae.

Even when two separate families are considered, Hutson’s(1977) identification key is still the best available, compris-ing all four genera. Other published keys exclude one or twoof the presently known genera.

Material and methods

A very large number of the species of Scatopsidae, belonging tomost genera of the family, have been examined, together with spe-cies belonging toCanthyloscelis(Araucoscelis), as well asSynneurondecipiens. The taxa used as outgroups were mainly Psychodidae,Tanyderidae, Tipulidae, Anisopodidae, Bibionidae, Sciaridae, andin some cases, species of different families of Mecoptera. The dis-cussions of Hennig (1973), Matile (1990), and Collucci (1995)concerning the ground plan of the Diptera were very useful forsolving some problems of homology and character polarity.

List of characters

In the following list of transformation series, the plesio-morphic condition of each transformation series is followedby the apomorphic condition or conditions. When a lineartransformation series has more than two conditions, lettersare used to designate successive apomorphic conditions. Adiscussion follows each character or group of charactersmodifying the same structures, justifying decisions concern-ing homology, character polarity, and other problems. Thetaxonomic nomenclature used in the discussion follows theclassification proposed below for the family. Character num-bers are the same as those used on the cladogram (Fig. 1).Figures 2, 3, and 4 present the wings and male and femaleterminalia, respectively, for the genera of the family on acladogram.

Adult features

1. Median ocellus well developed / (a) reduced / (b) absent.There is some variation in the size of the median ocellus

in the Scatopsidae, but there seems to be no doubt that it iswell developed in the ground plan of the family. In the

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Fig. 1. Proposed phylogeny for relationships among the generaof Canthyloscelidae. Characters are numbered according to thelist of transformation series in the text.



Canthyloscelidae we find the median ocellus well developedin Synneuronand Exiliscelis, reduced inHyperoscelisandCanthyloscelis(Araucoscelis), and absent inCanthyloscelis(Canthyloscelis) (Hutson 1977). It is assumed here that thereduction is synapomorphic forHyperoscelis+ Canthyloscelis,with the loss of the median ocellus as a synapomorphy forCanthylosceliss.str.

2. Eye bridge absent / (a) incomplete / (b) complete.In the Scatopsidae ground plan the eyes are completely

holoptic, although some secondary reductions to a nearlyholoptic condition occurred. The fully holoptic condition inCanthyloscelidae occurs inCanthyloscelis(Araucoscelis) andSynneuron, the other two genera possessing eyes that nearlymeet above the antennae (Hutson 1977). It is quite clear that

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Fig. 2. Pictorial cladogram of the Canthyloscelidae, showing the wings of the genera and subgenera of the family. Those ofExilisceliscaliforniensisand Synneuron decipiensare modified from Peterson and Cook (1981), those ofHyperoscelis eximiaare modified fromMamaev and Krivosheina (1969), those ofCanthyloscelis(Araucoscelis) antennataare modified from Edwards (1930), those ofCanthyloscelis(Canthyloscelis) brevicornisare modified from Nagatomi (1983), and those ofProhyperoscelis rohdendorfiare modifiedfrom Kalugina and Kovalev (1985).



the incomplete eye bridge is a synapomorphy of the Scato-psoidea. It is possible to argue either that the fully holopticcondition was achieved independently inSynneuron, Canthy-loscelis(Araucoscelis), and Scatopsidae, or that it is a scatop-soid ground-plan feature that has reversed to an incompletecondition inExiliscelisandHyperoscelis(which implies thatthe number of steps is the same). The first of these possibil-ities is accepted here.

3. Well-developed clypeus separates eyes below antennae /eyes closer to each other below antennae.

The Scatopsidae have a well-developed clypeus and thecompound eyes are not close to each other below the anten-nae. InSynneuron, Hyperoscelis, andCanthyloscelisthe eyesnearly meet below the antennae (Hutson 1977), but inExiliscelisthey are well separated. The ventrally contiguous eyes aresynapomorphic for the Canthyloscelinae.

4. Antennal flagellum with 16 articles / (a) 14 articles /(b) 10 articles (Hutson 1977).

A reduction from 16 flagellomeres (the Diptera groundplan condition) to 14 is apomorphic, but whether it is a synapo-morphy of the Scatopsoidea is difficult to determine. ThePsychodidae, for example, retain 16 flagellomeres, the num-ber also seen in the Tanyderidae, but in other basal families ofDiptera the number of articles in the flagellum is reduced.I tentatively accept here the reduction to 14 flagellomeres assynapomorphic for the Scatopsoidea. Within the group,Synneuronhas 10 flagellomeres, the Scatopsidae have 10 atmost, and the remaining genera of Canthyloscelidae retain14. The reduction from 14 to 10 articles in the group is con-sidered homoplasious betweenSynneuronand Scatopsidae.

5. Gena short / well developed (Hutson 1977).I agree with Hutson’s (1977) position that the well-developed

gena in the Scatopsoidea is apomorphic. In the ground planof the Psychodidae the gena is quite small. Hutson (1977)indicates that this would be an autapomorphy ofCanthylo-scelis s.str. In Canthyloscelis(Araucoscelis), on the otherhand, there is some development of the gena, although itis not as conspicuous as that in the other subgenus. Theapomorphic condition is here restricted toCanthyloscelis(Canthyloscelis).

6. Maxillary palpus with sensory pit / sensory pit lost.A sensory pit is present on the third article of the

maxillary palpus of different families of Diptera (e.g.,Tipulidae, Sciaridae) and is present in that condition inExiliscelis. The Scatopsidae retain a sensory pit on the singlearticle of the maxillary palpus, so it is reasonable to accept itas part of the Scatopsoidea ground plan.Synneuron, Hypero-scelis, andCanthyloscelisdo not have the pit on any of thearticles. The loss of the sensory pit is considered here to be asynapomorphy of the Canthyloscelinae.

7. Distal three palpomeres elongate / short and round.This reduction in size and modification of the shape of the

distal palpomeres inSynneuron(Hutson 1977) is an autapo-morphy of the genus.

8. Thorax elongated / stout.Hutson (1977) indicates that the elongated thorax would

be a synapomorphy of his “Synneuridae” plus the Scatopsidae.However, an elongated thorax is a plesiomorphic condition,found in different basal groups of Diptera. The inversion ofcharacter polarity would show the apomorphic condition shared

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Fig. 3. Pictorial cladogram of the Canthyloscelidae, showing the male terminalia of the genera of the family. Those ofExiliscelis andSynneuronare modified from Hutson (1977) and those ofHyperoscelisare modified from Nagatomi and Saigusa (1984). Gc,gonocoxite; Gs, gonostyle; Ae, aedeagus; Su, surstylus; S9 and S10, sternites 9 and 10; T8 and T9, tergites 9 and 10.



by Hyperoscelis+ Canthyloscelisand different subgroups ofScatopsidae.

9. Anterior thoracic spiracle within membrane / onanepisternum.

10. Anterior thoracic spiracle within membrane / on epimeron I.Hutson (1977; his character 9 in Fig. 26 and on p. 99)

proposed that the anterior spiracular sclerite “on a separatesclerite” would be a synapomorphy ofExiliscelis+ Synneuron+Scatopsidae. This seems to be an incorrect interpretation.The Scatopsidae clearly have the spiracle on a dorsal exten-sion of epimeron I in the Aspistinae and Ectaetiinae; thissclerite is partially divided in the Psectrosciarinae (as cor-rectly interpreted by Hutson 1977) and completely dividedin all Scatopsinae. InCanthyloscelisthe spiracle is in itsoriginal position, but on a large plate that is more likely anextension of the anepisternum. This condition is found inSynneuron, but the plate around the spiracle seems to bemore sclerotized than the rest of the anterior margin of theanepisternum. This may be the reason for Hutson’s (1977)interpretation of the genus as having the spiracle on a

separate sclerite. Insofar as I can interpret Hutson’s (1977,Fig. 1) drawing of the thorax ofExiliscelis, the genus hasthe plesiomorphic condition, with the anepisternum restric-ted to a more posterior position, epimeron I not developeddorsally (as in the Scatopsidae), and the spiracle situated onthe membrane. I do not have this information forHypero-scelis. I prefer two different transformation series. One is inthe ground plan of the Scatopsidae, in which the develop-ment of epimeron I is dorsal, involving the spiracle. Theother would be a synapomorphy of the Canthyloscelinae,with the anepisternum including the spiracle.

11. Suture of fusion of meron to thoracic sclerites clear,sclerites clearly separated / suture not conspicuous, meronincorporated into metathorax.

I disagree with Hutson’s (1977) interpretation of the evo-lution of the meron in the group. He proposes that a small me-ron would be a synapomorphy of (Exiliscelis+ Synneuron+Scatopsidae). However, the size of the meron inExiliscelisisvery similar to that seen inCanthyloscelis, and the meron inSynneuronis even larger than that inCanthyloscelis. An apo-morphic condition is found in the Scatopsidae, in which the

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Fig. 4. Pictorial cladogram of the Canthyloscelidae, showing the female terminalia of the genera of the family (all drawings are modi-fied from Hutson 1977). AnM, anal membrane; Ce, Ce1, and Ce2, cercus and articles 1 and 2; S7, S8, S9, and S10, sternites 7–10;T8, T9, and T10, tergites 8–10.



suture of the fusion of the meron to the thoracic sclerites ismuch less evident than in the Canthyloscelidae.

12. Posterior femur and tibia unmodified / posterior femurswollen, tibia curved (Hutson 1977).

The apomorphic condition of this transformation series isexclusive to Hyperoscelis and Canthylosceliswithin theCanthyloscelidae.

13. Tibial spurs reduced / absent.The reduction of the tibial spurs seems to be a feature

shared by the Scatopsoidea and Psychodidae. The spurs areshort but present inExiliscelis, Synneuron, andHyperoscelisand completely absent in Scatopsidae andCanthyloscelis. Twoindependent losses of the spurs in the superfamily appearlikely.

14. Tarsal claw simple / with basal prominent tooth.

15. Tarsal claw simple / with large basal lobe with smallteeth.

Two modified conditions of the tarsal claws are seen inthe Canthyloscelidae: inExiliscelis there is a basal promi-nent tooth, while inCanthyloscelisthere is a basal lobe onwhich a group of small teeth is seen (Hutson 1977). Even ifwe code these two modifications as identical in a data ma-trix, they would appear as homoplasies in the cladogram.Because they are not identical in shape they are coded differ-ently and assumed to be different autapomorphies for thegenera.

16. Wing membrane completely clear / maculate.The wing membrane of all scatopsids is completely clear

except for some entirely yellowish or brownish wings. Con-sidering the variation within the Scatopsidae and most out-groups, it is possible to infer with certainty that in theground plan of the family the wing is completely clear. Thiscondition is also seen inExiliscelisandSynneuron. Hypero-scelis and Canthyloscelishave dark markings on the wingthat correspond to a synapomorphy ofHyperoscelis. This re-fers to a transverse distal band; an additional medial bandmay appear. Figure 2 illustrates the wings of the genera ofthe family.

17. Wing membrane with macrotrichia and microtrichia /with only microtrichia, macrotrichia entirely absent.

The evolution of macrotrichia on the wing membrane inDiptera is still unclear. Basal Bibionomorpha groups (suchas the Pachyneuridae and Cramptonomyidae) have macro-trichia on the wing membrane, as do the Ptychopteridae,some Tipulomorpha (especially the Eriopterini), some Culico-morpha (such as Thaumaleidae, Ceratopogonidae, and Chiro-nomidae), and Brachycera families (such as Stratiomyidae).Within the Scatopsoidea, most Canthyloscelidae do havemacrotrichia on the wing membrane, as well as in the basalscatopsid stems. It does not seem possible now to determinewhether or not there were macrotrichia in the Dipteraground plan. Aside from this question, however, macro-trichia are quite widely distributed among members of theScatopsoidea. An optimization analysis points to the pres-ence of macrotrichia on the membrane as a feature of theScatopsoidea ground plan, so its absence within the groupwould be apomorphic. In the Canthyloscelidae this loss is

known to have occurred inCanthyloscelisand on differentoccasions in the Scatopsidae (in which the loss occurredgradually).

18. Subcostal vein (Sc) complete / incomplete.This is a synapomorphy of the Scatopsoidea. None of the

scatopsid or canthyloscelid genera have a complete Sc. Hutson(1977) proposed that the faint Sc would be a synapomorphyof his “Synneuridae” + Scatopsidae. I disagree, sinceHypero-scelishas a very faint Sc and the condition in mostCanthylo-scelisspecies is similar to the plesiomorphic condition foundin Exiliscelis and some Scatopsinae.

19. Vein R1 long, reaching C quite close to R4 / R1 shorter,reaching C in a more basal position.

In the Diptera ground plan, R4 reaches C very distally onthe wing, quite close to the wing apex, so the apomorphiccondition of the character is certainly that the vein ends in amore basal position. In the Canthyloscelidae we find R4reaching C quite separate from R1 in Canthyloscelisbutcloser to it inExiliscelisandHyperoscelis. Synneuronis notcomparable in this feature, nor are the Scatopsidae. Thiscould represent an apomorphic feature shared byExiliscelisandHyperoscelis. However, more detailed observation of thewing shows that the condition inCanthyloscelismay not beplesiomorphic: there is probably a retraction of R1 in this ge-nus (in which it ends quite close to the medial fork).

20. Vein Rs free from R1 distally / Rs and R1 partially fuseddistally (Hutson 1977).

This is an autapomorphy ofSynneuron.

21. Vein R4 present / lost (Hutson 1977).Synneuronis the only genus of Canthyloscelidae that lacks

R4. This feature is considered here to have been achievedindependently from the Scatopsidae. It is possible that thefeature is somehow related to the distal fusion between Rsand R1.

22. Vein Rs completely separate from R1 in the basal third,r1 cell angled / (a) r1 cell not too wide, angled / (b) r1 cellslender, Rs parallel to R1 from the beginning.

Exiliscelis is the only genus of Canthyloscelidae in whichRs is largely separated from R1 in its basal third, which issimilar to the shape seen in other basal families of Diptera(see Anisopodidae, Trichoceridae, Cramptonomyiidae, basalBrachycera, some Culicomorpha families, etc.). InSynneuronthe r1 cell is still angled anteriorly, although it is not partic-ularly slender.HyperoscelisandCanthyloscelishave a slen-der r1 cell, with Rs parallel to R1 right from its base, and thevery basal sector of Rs is nearly transverse.

23. Vein M1+2 not fused to Rs / (a) fused for a short distance /(b) fused for at least three times r–m length.

Fusion of the base of M1+2 to Rs is seen in all Canthylo-scelidae. The fusion is quite short inExiliscelisbut longer inall remaining genera of the family. There is a similar shortfusion in some genera of Scatopsidae, such asAspistes,Ectaetia, and part ofPsectrosciara, but in the ground plan ofthe family there is certainly no fusion. Hence, this is an ad-ditional synapomorphy of the Canthyloscelidae. The longerfusion is a synapomorphy of the Canthyloscelinae.

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24. Vein R2+3 present / absent.Because of the inclusion of the Scatopsoidea in the Psychodo-

morpha, the absence of R2+3 must be seen as a synapo-morphy of the group. It is present in the Psychodidae andTanyderidae (and even in other supposedly closely relatedfamilies) but absent in the Scatopsidae and Canthyloscelidae.I must agree with Hutson (1977) that the vein branchingfrom R5 is R4, not R2+3, a conclusion that differs from Mamaevand Krivosheina (1969), Nagatomi (1983), Nagatomi andSaigusa (1984), and Cook (1981). This comes almost neces-sarily from the position of the radial-sector fork, which isvery distal in the wing, even inExiliscelis, which is sup-posed here to be a genus with quite conservative wing vena-tion (that nevertheless presents its own apomorphies).

25. Vein R5 unmodified / thickened.In all Canthyloscelidae, R5 is unmodified in relation to the

ground plan of the family (which is virtually the same as theground plan of the Diptera in its shape).Canthyloscelis(Araucoscelis), however, has a thickened R5 (Hutson 1977),an obvious apomorphic condition exclusive to the subgenus.

26. Basal sector of Rs (before contact with M1+2) long(longer than R4) / basal sector of Rs very short.

This is an additional synapomorphy of the Canthylo-scelinae in the sense used here. M1+2 contacts Rs quite dis-tally and the shape of the basal sector of Rs is conservative.In the remaining genera of the family, M1+2 connects to Rsvery basally.

27. Vein bM present / absent.Like the loss of R2+3, the absence of bM must be seen as a

synapomorphy of the Scatopsoidea.

28. Free sector of M1+2 (basal to fusion with Rs) short /long.

Exiliscelis possesses a short free basal M1+2 (before thefusion between bM and Rs), which is also seen inSynneuronandCanthyloscelis. In Hyperoscelisthis basal sector of M1+2is more elongated, possibly because the cubital fork is dis-placed to a more basal position, without a correspondingshift in the base of Rs.Synneuronalso has a very basalcubital fork, but a short M1+2 base. In the Scatopsidae, bM islong and longitudinal in position because of strong displace-ment of the cubital fork to the base of the wing. However,this condition seems to have been acquired independently.Other outgroups, such as the Psychodidae, would showin their ground plan the plesiomorphic condition seen inthe Diptera. This can be considered a synapomorphy ofHyperoscelis.

29. Vein M1 curved basally / M1 stems straight from fork.This is a modification seen inSynneuronand Canthylo-

scelis. The typical anterior basal curve of M1, seen in mostDiptera families, is absent in these two groups; this is possi-bly related to the basal interruption of M2.

30. Vein M2 complete basally / incomplete basally.The basal interruption of M2 is certainly apomorphic and

shared, homoplastically, in the Canthyloscelidae bySynneuronandCanthyloscelis. Edwards (1930, text and Fig. 4) depictsthe wing of Canthyloscelis antennataEdwards with a com-plete M2, but Edwards (1930: 90) in the text and also Hutson

(1977: 78) in his key to the genera of Canthyloscelidae con-firm that the basal interruption of M2 is a key feature of thegenus.

31. Vein m-m present / absent.

32. Vein M3 present / absent.Veins m-m and M3 are absent in all members of the

Scatopsoidea and must be taken as synapomorphies of thegroup, since it is placed within the Psychodomorpha, mostmembers of which have them. The inclusion of the Scatop-soidea in the Mycetophiliformia is partially based on theabsence of M3 and m-m.

33. Contact between M1+2 and M3+4 well sclerotized / hardlysclerotized mesally.

With the loss of bM, the base of M1+2 and the base ofM3+4 (the latter is called “m-cu” by most authors) are aligned.The vein is completely sclerotized inExiliscelisandSynneuron,but is rather interrupted inHyperoscelisandCanthyloscelis.The condition seen in the latter two genera is certainlyapomorphic.

34. Vein M3+4 (m-cu) connects with CuA1 / connects withCuA.

This is an autapomorphy ofCanthyloscelis(Canthylo-scelis). Even the Scatopsidae, in which the cubital fork is ina very basal position in the wing, seem to show a “m-cu”connection to CuA1, not to CuA.

35. Vein M3+4 (m-cu) nearly horizontal / oblique or vertical.In Exiliscelisthe medial connection to CuA1 is nearly hor-

izontal. This is due to displacement of the cubital fork to thebase of the wing. As is described below, this cubital dis-placement goes even further in the Canthyloscelinae andScatopsidae. However, there is parallel displacement of theorigin of Rs to the base of the wing, so this CuA1–Rs con-nection moves from a nearly horizontal to an effectivelytransverse position.

36. Cubital fork short, shorter than CuA / (a) cubital forklonger, CuA about half length of CuA2 / (b) cubital fork verylong, CuA at most a third of CuA2.

If the Psychodidae is considered the sister-group of theScatopsoidea, the displacement of the cubital fork to thebase of the wing would be an apomorphy shared by theScatopsidae and Canthyloscelidae. If any other family in thePsychodomorpha is considered the sister-group of theScatopsoidea, the displacement would be synapomorphic forthe group. I here tentatively place the feature as a synapo-morphy for the Scatopsoidea. The second step, where thecubital fork is still more basal in the wing, seems to be asynapomorphy of the Canthyloscelinae within the family.Obviously the condition in Scatopsidae is yet moreapomorphic than that found in any genus in the Canthylo-scelidae, but would have arisen independently under thisscheme.

37. Vein CuP present / virtually nondiscernible.The CuP vein (“Cu2”) is indicated by Hutson (1977) as a

synapomorphy of his “Synneuridae” + Scatopsidae. How-ever, CuP can be clearly seen inExiliscelis. Even thoughCuP is quite faint in some basal members of the group, it

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should be considered present in the ground plan of the Scatop-sidae. On the other hand, inHyperoscelisandCanthyloscelisthe vein seems to be indistinguishable. I prefer to state thatthe nearly complete reduction of the vein is a synapomorphyof the Canthyloscelinae.

38. Vein A1 distally complete / (a) very short / (b) absent.Hutson (1977) interpreted the distal interruption of A1 as

a synapomorphy ofExiliscelis + Synneuron+ Scatopsidae.This seems to follow Hennig’s (1954) statement that the lossof A1 would be synapomorphic for the Scatopsidae. How-ever,Aspistes, Ectaetia, and part ofPsectrosciarado have acomplete A1 and it should be designated a complete vein inthe Scatopsidae ground plan. The first apomorphic conditionin the Canthyloscelidae is seen only inExiliscelis. Completeloss of the vein is known only inSynneuron. A number ofother apomorphic features indicate thatSynneuronconsti-tutes a monophyletic group withHyperoscelisand Canthy-loscelis, so the reduction of A1 must be seen as a homo-plasious derivation inSynneuronand Exiliscelis.

39. Vein A1 complete basally / A1 incomplete basally.This quite unusual modification in A1 is apomorphic only

for Canthyloscelis(Canthyloscelis) (Hutson 1977).

40. Unmodified male pregenital segments 1–7 / (a) 1–6 /(b) 1–4 (Hutson 1977).

Male abdominal segment 8 is reduced in the Scatopsidaeand Canthyloscelidae. In the abdomen ofCanthyloscelis(Arauco-scelis), segment 7 is strongly modified, while in that ofCanthyloscelis(Canthyloscelis), segments 5–7 are stronglymodified (Hutson 1977). The modification of segment 7 isunderstood as synapomorphic forCanthyloscelisand themodifications of segments 5 and 6 as autapomorphic forCanthyloscelis(Canthyloscelis).

41. Sperm pump attached to male terminalia / free inabdomen (Hutson 1977).

A sclerotized male sperm pump attached to the maleterminalia is supposedly present in the Diptera ground plan(Wood 1991), and is also seen in the Mecoptera and Siphon-aptera. The sperm pump, which is present in the Tipulinae,Ptychopteridae, Anisopodidae, Scatopsidae, and Canthylo-scelidae, was certainly lost many times in the Diptera. Withinthe Scatopsoidea, the sperm pump, a sclerotized structure, ofthe Scatopsinae is detached and free in the abdomen, con-nected to the terminalia only by the ducts. Hutson (1977) in-dicates that inSynneuronalso, the sperm pump is detachedfrom the terminalia. In the maleS. decipiensI examined itseemed that the sperm pump was attached to the terminalia,although the connection between the two structures was notas strong as in other genera. I takeSynneuronto be apo-morphic for the transformation series. It is very obvious,however, that the condition inSynneuron, if its apomorphiccondition is confirmed, was achieved independently fromthat seen in the Scatopsinae.

42. Sperm pump encapsulated / not encapsulated (Hutson1977).

Hutson (1977) describes the sperm pump inCanthylo-scelisas not encapsulated, supposedly an apomorphic condi-tion.

43. Gonocoxites separated / fused mesally.

44. S9 situated anteriorly on the terminalia / S9 displaced toa posterior position on the terminalia.

45. Gonocoxites rather small / gonocoxites well developedposteriorly and laterally.

46. Gonostylus finger-shaped / slender and hardly sclerotizedposteriorly.

47. Gonostylus rather straight / folded mesally.

48. S9 rather small / projects posteriorly between the gonostyli.

49. S9 rather small / present as a flat, well-developed plateventrally on the terminalia.

50. Surstyli present, short / present, well developed.Some distinctive features of the male terminalia can be

found in the genera of Canthyloscelidae in relation to theDiptera ground plan, but they are not easy to interpret. It ispossible that mesal fusion of the gonocoxites is a synapo-morphy of the Canthyloscelidae. This is certainly found inExiliscelis and seems to be present in the terminalia of theother three genera also (Fig. 3). InExiliscelis the gono-coxites are well developed (see Peterson and Cook 1981,Figs. 4 and 5), are quite rounded, and displace the gonostylito a more posterior position in the terminalia. InSynneuronthere is a mesal plate between the gonostyli that seems to bean extended sternite 9. A posteriorly displaced and elongatedsternite 9 plate is also found inExiliscelis. Hyperoscelishasa large mesal plate on the terminalia ventrally, with thegonostylus placed lateral to it. It seems likely that posteriordisplacement of sternite 9 is a synapomorphy for the Canthylo-scelidae. If this is correct, Peterson and Cook’s (1981) indi-cation of the position of sternite 9 (“hypandrium”) would beincorrect (unless sternite 9 is extended laterally at the ante-rior extremity of the terminalia and posteriorly between thegonocoxites). The mesally projected sternite 9 inSynneuronand the large flat sternite 9 inHyperosceliswould be autapo-morphies. The gonostylus ofExiliscelis, on the other hand,is easily identifiable, having a large base and a slender, well-sclerotized distal end, certainly an apomorphic condition. Inthe other three genera, the gonostyli are more difficult toidentify. They are rather elongated, but they have a mesalfold with a different shape in each genus. This modificationseems to be synapomorphic for the Canthyloscelinae. Thequestion of the homology of the “surstyli” is a difficult oneto answer. Matile (1990: 64–65) stated that these scleritesare metameric appendages of segment 10 and would be pres-ent in the Diptera ground plan. If this is correct, their pres-ence inExiliscelis, for example, would be archeomorphic.Hence, the apomorphic condition here would be a secondarydevelopment. It can be also seen inCanthyloscelis, but is notas well characterized inSynneuronor, especially,Hyperoscelis.Here I tentatively place the feature as a synapomorphy of theCanthyloscelidae.

51. Female spermatheca simple / modified (Hutson 1977).An oval or spherical female spermatheca is seen in differ-

ent families, including the Scatopsidae. This condition ismodified in both subgenera ofCanthyloscelis, which have anelongated spermatheca with a medial constriction and an

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apical extension that continues into the duct. This is cer-tainly a synapomorphy of the genus.

52. Female tergites 9 and 10 separate / fused.

53. Female sternite 8 with short posterior notch separatinggonapophyses 8 / deep mesal notch separating shortgonapophyses 8.

54. Female cercus two-segmented / one-segmented.Few features can be found in the female terminalia of the

Canthyloscelidae (Fig. 4). The presence of two segments inthe female cercus is a feature of the Diptera ground plan(Hennig 1973). In the Canthyloscelidae this is modified onlyonce, in Synneuron, the only genus with a one-segmentedcercus. In the Scatopsidae, a one-segmented cercus is pres-ent only in the Aspistinae. The structure interpreted as acercus in the Scatopsinae is probably a secondarily dividedtergite 10. Fusion between tergites 9 and 10 seems to be asynapomorphy of the Canthyloscelinae. The deep mesalnotch on sternite 8 of the female terminalia is an apomorphyseen inSynneuronandHyperoscelisbut not inExiliscelisorCanthyloscelis. Here the notch is interpreted as a synapo-morphy of the Canthyloscelinae, reversed to a condition sim-ilar to the plesiomorphy inCanthyloscelis.

Immature features

55. Head capsule well developed and sclerotized / (a) headcapsule weakly sclerotized and reduced / (b) greatly reducedand unsclerotized (Hutson 1977).

56. Antennae short but normally developed / reduced, with acharacteristic posteriorly ovoid shape (Hutson 1977).

57. “Enigmatic organ” absent / present (Tonnoir 1927 inHutson 1977).

58. Cuticle with normal setae / setae on body surface re-duced and covered with spinulose areas (Hutson 1977).

59. Posterior end of body without hooks / with a pair ofheavily sclerotized hooks set on a single adanal plate(Hutson 1977).

Hutson (1977) listed a number of features of the larvae ofSynneuron, Hyperoscelis, andCanthyloscelisthat are modi-fied, and in which they differ from the larvae of Scatopsidae,Cecidomyiidae, and Mycetophilidae. The differences are notrestricted to the most conspicuous modification, the reduc-tion of the head capsule. Hutson (1977) listed other featuresknown to occur in the larvae of the three genera with knownimmatures. However, he did not consider that these modif-ications were sufficient to corroborate the hypothesis thatSyn-neuronandExiliscelisconstitute a monophyletic group withHyperoscelisand Canthyloscelis. I concur with Wood andBorkent’s (1989) position that collectively the larval featuresstrongly indicate that the three genera compose a mono-phyletic group separate from the Scatopsidae. Unfortunately,the larva ofExiliscelis is still unknown and generalizationsare merely hypothetical. We assume here provisionally thatthe apomorphic conditions proposed above for the three gen-era are part of the Canthyloscelidae ground plan.

Discussion

The hypothesis proposed here to account for the relation-ships between members of the Scatopsoidea differs from thatof Hutson (1977) in two main aspects. First, in his systemExiliscelis and Synneuroncompose a monophyletic groupwith Scatopsidae, while here these two genera compose amonophyletic group withCanthyloscelisand Hyperoscelis.In this respect the monophyly of the Canthyloscelidae fol-lows Wood and Borkent (1989). Second, some similaritiesshared byExiliscelis and Synneuronthat were consideredapomorphic by Hutson (1977) are considered here to behomoplasies or plesiomorphies.

I shall address in more detail Hutson’s (1977: 98–99,Fig. 26) discussion of the characters at these levels of hiscladogram. The synapomorphies that he proposed as a basisfor including Synneuronwith the Scatopsidae are as follows(only apomorphic conditions are given): head rounded andsomewhat flattened, eyes meet above the antennae, prothoraxstoutly developed, R4 lost, and cercus one-segmented (re-ferred to as ovipositor one-segmented). Regarding this set offeatures, I would disagree that the shape of the head is anapomorphy of these two groups and I cannot understandHutson’s (1977) description of the modification of the pro-thorax. The other three features (2b, 16, and 36 here) are notunusual in outgroups. Hence, based on the available data setfor larvae and adults, it seems more parsimonious to con-sider the similarities betweenSynneuronand Scatopsidae ashomoplastic. To accept that the considerable modificationsof the larvae shared bySynneuronand two Canthyloscelidaegenera originated homoplastically, it would be necessary tofind considerably more consistent apomorphies shared be-tweenSynneuronand Scatopsidae than are presently known.The eye bridge is certainly homoplasious betweenSynneuronandCanthyloscelis(Araucoscelis), and is also known in manyother groups of Diptera (some Cecidomyiidae, Sciaridae,Perissomatidae, etc.); R4, as discussed below, is possiblypresent in the sister-group of the Scatopsidae,Mesoscatopserohdendorfi Kovalev; the one-segmented cercus is seen innumerous other groups of Diptera.

The features proposed by Hutson (1977) to uniteSyn-neuron and Exiliscelis with the Scatopsidae also deservesome attention. His proposed synapomorphies for this groupare as follows: thorax elongated, spiracle on a separatesclerite, CuP (his Cu2) absent, anal vein absent, and Scweakly present. I believe that thoracic shape has the oppositecharacter polarity to that which he presents, the stout thoraxbeing an apomorphy shared byHyperoscelisand Canthylo-scelis, and has originated several different times within theScatopsidae. Hutson’s (1977) discussion concerning the po-sition of the anterior thoracic spiracle, as mentioned above,seems incorrect. InCanthyloscelisandSynneuronthe spira-cle is actually on a large plate formed by the anepisternum.In the Scatopsidae ground plan the spiracle is on a dorsalextension of epimeron I and inExiliscelis it is in itsplesiomorphic position, on the membrane between epimeronI and the anepisternum. Hence, two independent apomorphicconditions can be defined, one for the Scatopsidae and theother for the Canthyloscelinae. The reduction of CuP is alsonot a synapomorphy of this group: it is very faint in the

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basal Scatopsidae but visible inSynneuronand effectivelypresent inExiliscelis. It would be better to restrict the veryfaint condition of CuP to the Scatopsidae. The reduction ofA1 is indeed an apomorphic condition shared bySynneuronand Exiliscelis, but is most certainly not a feature of theScatopsidae ground plan. As mentioned above, the vein iscomplete in different basal groups of Scatopsidae, hence theplesiomorphic condition would have to be attributed to theground plan of the family. Finally, the faint Sc would bebetter seen as a synapomorphy of the Scatopsoidea. There isindeed some variation in the extension of the vein and thedegree of sclerotization in the group. InCanthyloscelis, aswell as the Scatopsinae, it seems to be slightly more devel-oped. In Exiliscelis the vein is longer but not very wellsclerotized, and inSynneuronand Hyperoscelisthe vein isactually hardly visible, so I would not conclude that there isan apomorphic condition shared exclusively byExiliscelis,Synneuron, and Scatopsidae. Again, it is difficult to acceptthat the features proposed by Hutson (1977) as synapomor-phies of his “Synneuridae” + Scatopsidae could be more re-liable indications of a common exclusive ancestry than themodifications of the larvae and the other adult features sharedby these genera withCanthyloscelisand Hyperoscelis.

On the other hand, Hutson’s (1977) conclusions about themonophyly of the genera and subgeneraHyperoscelis, Canthylo-scelis, Canthyloscelis(Canthyloscelis), Canthyloscelis(Arauco-scelis), Exiliscelis, andSynneuronand the groupHyperoscelis +Canthyloscelisare completely corroborated in this analysis.

Known fossils of CanthyloscelidaeThere is an undescribed fossil species from the Lower

Cretaceous of Australia referred to by Eskov (1992) as be-longing toSynneuronthat will not be considered here. If itsinclusion in the genus is confirmed, this would extend thedistribution of the genus to the southern hemisphere, at leastduring the Mesozoic, which would be expected, consideringthe age of the group (Nagatomi 1983). A fossil species fromthe Middle Jurassic of Siberia,Prohyperoscelis rohdendorfi,has also been described as belonging to the Hyperoscelidaeby Kovalev (Kalugina and Kovalev 1985) and reassigned tothe “Canthyloscelididae” by Evenhuis (1994) (actually an in-correct form of Canthyloscelidae; see Hutson 1977). As canbe seen from the discussion above, I accept the Canthylo-scelidae as a whole as monophyletic and I keep it as a singlefamily-rank taxon. However, I agree thatHyperoscelis+Canthylosceliscompose a monophyletic group (that corre-sponds to Hutson’s (1977) Hyperoscelidae), andProhypero-scelisindeed seems to belong to this taxon. The long fusionof M1+2 with Rs, the long cubital fork, the slender r1 cell,the alignment between the base of M1+2 and the base of M3+4(m-cu), the incomplete contact between M1+2 and M3+4, andthe shorter R1 are some of the apomorphies seen in the wingof Prohyperoscelisthat are shared with the remaining extantCanthyloscelinae, the Canthyloscelini, and the Canthylo-scelina. A1 is present and complete, reinforcing the conclu-sion that the position of this fossil species is not inExiliscelisor Synneuron. Actually, the wing venation is verysimilar to that ofCanthyloscelis. Some important differencesare that M2 is complete at the base, CuP seems to be welldeveloped, M1 is not very curved at its base, although it isnot straight as inCanthyloscelis, and M1+2 and M3+4 are not

completely aligned, the base of M1+2 occupying a slightlymore distal position. These are actually plesiomorphies ofthe fossil species in relation to modifications shared byCanthyloscelis(Canthyloscelis) andCanthyloscelis(Arauco-scelis) and it would be very reasonable to consider thatProhyperosceliscorresponds to the sister-group ofCanthylo-scelis. This adds a northern hemisphere representative to amonophyletic group previously regarded as having only asouthern temperate distribution.

Classification proposedThe classification below is a sequenced phylogenetic clas-

sification (Nelson 1972), with “plesion” used as a substitutecategory for Linnaean ranks of extinct taxa (Patterson andRosen 1977). Square brackets are used to indicate redundantnominal taxa in the classification, a convention proposed byChristoffersen (1988).Scatopsoidea

Scatopsidae Newman, 1834Canthyloscelidae Shannon, 1927

Exiliscelinae, subfam.nov. [Exiliscelis Hutson, 1977[E. californiensisHutson, 1977]]

Canthyloscelinae Shannon, 1927Synneurini Enderlein, 1936 [SynneuronLundström,

1910]Canthyloscelini Shannon, 1927

Hyperoscelina Hardy and Nagatomi, 1960 [Hypero-scelis Hardy and Nagatomi, 1960]

Canthyloscelina Shannon, 1927PlesionProhyperoscelisKovalev, 1985 [P. rohden-

dorfi Kovalev, 1985]CanthyloscelisEdwards, 1922Canthyloscelis(Canthyloscelis) Edwards, 1922Canthyloscelis(Araucoscelis) Edwards, 1930

Acknowledgements

The manuscript benefited from corrections and sugges-tions made by Dr. Art Borkent, to whom I am sincerely in-debted. An anonymous reviewer also helped with commentsand corrections. This work was supported by the Brazilianfederal science agency, Conselho Nacional de Desenvolvi-mento Científico e Tecnológico, with a research fellowship.

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A new phylogeny and phylogenetic classification for the Canthyloscelidae (Diptera: Psychodomorpha)

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