Pseudoperomyia gen. n. from Malaysia and the phylogeny of the Micromyidi

A new cecidomyiid genus Pseudoperomyia gen. n. is described from Malaysia, with 8 new species: P. acutistyla sp. n., P. bidentata sp. n., P. humilis sp. n., P. parvolobata sp. n., P. intermedia sp. n., P. longicomis sp. n., P. macrostyla sp. n., andP. platistyla sp. n. Pseudoperomyia belongs to the lestremiine tribe Micromyini with Anodontoceras as its sister group. The result of a computerized parsimony analysis for the Micromyidi largely supports a recently-published hypothesis on the phylogeny of this taxon.


Introduction
Present knowledge of the gall midges of the subfamily Lestremiinae with free-living larvae is largely restricted to the Holarctic Region. Except for a few occasional species records scattered in the literature, nothing more substantial has been published on tropical lestremiines. 26 species are recorded from the Oriental Region, most o f them from India (Gagné, 1977). They are associated with widespread Holarctic genera, but their true specific status is largely unrevised. The le stremiine fauna of South East Asia is completely unknown. During our studies of Oriental Lestremiinae, we have discovered a group of very similar species which clearly belong to the Micromyidi, as it has recently been defined (Jaschhof, 1998), but do not fit any of the recognized genera. We are assigning them to a new genus Pseudoperomyia. The aims of the present work are to describe Pseudoperomyia and its eight included species, to demonstrate the monophyly of the group, to find its sister-group, and to show the interrelations hips of its species. At the same time we test by computerized parsimony analysis the recently proposed hypothesis on the phylogeny of the whole Micromyidi (Jaschhof, 1998).

Material and methods
Most of the specimens studied here were collected during an expedition to parts of the Malaysian Peninsula in February / March 1997. Additional material came from the holdings of the Swedish Museum of N atural History, Stockholm, from previous collecting trips to Malaysia in 1992 and1994. The specimens were collected with Malaise traps, sweepnet and exhaustor, and were preserved in 70 % alcohol. To mount the specimens on microscope slides, they were first trans ferred from alcohol into a mixture of alcohol / formaldehyde (10 ml of 70 % alcohol mixed with 2-3 drops of 30 % formaldehyde) and then placed in creosote for 12-24 hours. On the slide, the hypopygium and (in some material) one wing were separately mounted in Canada balsam as well as the rest of the body. In some cases we dissected the male genitalia to scrutinize the structural details, but only when there was sufficient material available and when the hypopygium was large enough to manipulate with insect pins. All the material discussed here is deposited in the Swedish Museum of Natural History, Stock holm. The drawings were made with the aid of a drawing tube attached to a Leitz Diaplan mi croscope. The measurements were made with a micrometer slide. The morphological terminology follows Jaschhof (1998). Some o f the characters used in the parsimony analysis are explained in Appendix 1. The parsimony analysis was carried out using PAUP version 3.1.1. (Sw o ffo r d , 1993) in combi nation withMacClade version 3.06 (Maddison & M add ison, 1992). Bootstrap proportions were computed using PAUP and decay indices using PAUP in combination with AutoDecay version 3.0 (Eriksson & Wikstrom , 1995

Description
Male. Head: postfrons prominent, without setae or scales. Ocelli 3 in number or absent. Postocel lar bristles present. Eye bridge complete, 2-7 facets wide, at vertex little wider than laterally. With 1 or 2 rows of postocellar bristles. Occiput and clypeus densely covered with scales and sometimes additionally with setae. Antennae 2 + 12-segmented. Scape and pedicel of same size, with scales and setae. 1st flagellomere slightly elongated and covered with scales basally. Neck of 4* flagellomere slightly shorter to slightly longer than node. Node of subglobular type, as long as wide or slightly longer; with setae and sensory hairs in specific irregular or whorl-like arrangements, without crenulate whorls (exceptionally with occasional indistinct traces); with sensory spines and characteristic small bottle-shaped or conical sensillae distally. Distal flagellomeral nodes becoming more slender and necks longer. Palpus with 3 segments; 1st segment shortest (despite the indistinct basal stem), with scattered sensory hairs, long setae and scales; distal segments more slender, ter minal segment as long as to obviously longer than segment 2 and tapering distally, both segments with spine-like setae and scales, 2 nd segment sometimes with a few additional sensory hairs. Thorax: anterior pronotum' with a row of long setae. Scutum with long setae laterally and along parapsidal sutures in well defined stripes; lateral setae irregularly arranged and numerous (exten ded type, Fig. 8D) or more or less row-like and few in number (reduced type, Fig. 2F). Scutellum with 1 transverse row of long setae interrupted medially. Legs: densely covered with scales, among them additionally with a few setae; scales becoming increasingly shorter and broader on distal parts of legs. Claws crescent-shaped, those on middle legs with 2-4 small teeth mesally. Empodium narrow, as long as claws or slightly shorter. Wing: membrane densely covered with small scales; macrotrichia ventrally on R, R , and 2/3 of CuA-stem proximally; pattern of sensory pores: R j 3 (occasionally 4), rs 1, r-m 1; Sc distinct, reaching level of R,; strong anterior part of C running approximately 7 times the vein width beyond the intersection with R,, followed by distinct break; R , up to 2 times the length of rs; r-m short, usually about as long as rs; CuA -fork with angle of 85-90°; M m and CuA, usually not reaching wing margin; anal margin slightly convex. Haltere with stem strong, half the length of the ovoid knob; knob densely covered with scales and with a few setae on stem. Abdomen: segment 1 non-setose. Tergites 2-6 covered with scales, tergites 7-8 bare except for some lateral scales, and with strongly sclerotized anterior margin; intersegmental membrane with ovoid patch of scales; stemites densely covered with scales (exceptions: see under species des criptions). Hypopygium: very diverse; in general gonocoxites united ventrobasally, with setae and scales ventrally; gonostylus usually with apical tooth-like structure; genital rod present; tegmen mem braneous except for parameral apodemes, partly with characteristic nose-shaped projections dorsally; tergite 9 stripe-like with 1 row of long setae or strongly modified with horn-like projec tions, narrowed distal margin and reduced number of setae; tergite 10 bilobed, ovoid or elongated, partly with sclerotized arc dorsally, usually with setae; stemite 10 weakly membraneous, bilobed, without setae. Body length: 1.0-1.4 mm. Female: unknown.

Biology
Largely unknown. All the specimens were collected along forest margins or inside lowland and mountainous primary forest up to 1750 m. It is very probable that one of the species emerged from humus soil (see P. humilis). Only one of the 8 species was at all common.

Etymology
With their small body size, together with the subglobular flagellomeral nodes, specimens of Pseudoperomyia resemble Peromyia in general appearance under the stereo microscope, and the generic name is derived from this.

Discussion
Pseudoperomyia can easily be distinguished from other related genera of the Micromyidi by the combination of the following derived characters: 1. the lack of sensory pores on R s (with simulta neous presence of 1 pore on r-m), 2. the absence of macrotrichia on R h 3. the presence of sub globular flagellomeral nodes lacking crenulate whorls and with a rather centrally placed neck, 4. the occurrence of bottle-shaped and conical flagellomeral sensillae, 5. the lack of postfrontal setae, 6. the presence of nose-like and dorsad-directed projections of the tegmen in at least some of the included species, and 7. the stripe-like shape of tergite 9 which is distinctly narrower than the distal margin of gonocoxites in some of the included species. The bottle-shaped and conical sensillae as well as the nose-like tegmen projections are unique characters found nowhere else within the Lestremiinae. Pseudoperomyia differs from the other genera of the Micromyini (sensu Jaschhof, 1998) by each of the characters 1, 2 and 4-7. The globularisation of the flagellomeral node with simultaneous reduction of crenulate whorls (character 3) also occurs in Anodontoceras. Peromyia species, which are superficially similar to Pseudoperomyia and share characters 2,3 and 5 in some of the species, have 1 sensory pore on R s (whilst lacking the /•-m-pore).
Tegmen with waist-like constriction; genital rod very short, much shorter than tegmen; projections of tergite 9 very short (Fig. 8B)   Lobes of tergite 10 distinct, elongated and with distal horn-like prolongations; gonostylus with subapical lobe (Fig. 4A)    Description M ale. Head: ocelli absent. Eye bridge 5-6 facets wide. With 1 row of postocular bristles, 1-2 weaker bristles in a short second row situated more ventrally. Occiput densely covered with scales or setae (totally absent in the specimens available). Flagellum with more than 11 segments (broken in the specimens available). Neck of flagellomere 4 longer than node (Fig. ID); node with a double whorl of setae (or scales?, broken) basally and with irregularly arranged sensory hairs of different lengths, the most proximal sensory hairs shortest and forming a rather distinct whorl; among the hair-like sensillae with a few sensory spines and 1 inconspicuous bottle-shaped sensilla distally (lacking from flagellomere 6 on). Terminal palpal segment a little to obviously longer than segment 2; 2nd segment with a few sensory hairs basally. Thorax: arrangement of lateral scutal setae of the reduced type (see Fig. 2F). Legs: claws of middle legs with 2 small teeth mesally. Empodium approximately as long as claws. Wing: Rj=l l A-2 rs; CuA -fork approximately rectangular. Hypopygium: gonocoxites ( Fig. 1A) setose ventrally (scales broken if present); distal margin with broadly U-shaped emargination. Gonostylus (Figs. 1 A, C) widest in proximal third, strongly tapering to tip; with apical fingernail-like tooth (appears to be a pointed tooth in lateral view) and 2 subapical spines clearly shorter than tooth; with long setae. Tegmen (Fig. IB) broadest at base, tapering to narrow tip; apex with a ventrally-directed, cap-like projection; parameral apodemes not reaching basal gonocoxal margin. Genital rod (Fig. IB) sclerotized, narrow and with indistinct invagination near apex. Tergite 9 with broadly rounded distal margin and 1 transverse row of setae. Tergite 10 bilobed ovoid, with a few weak setae. Stemite 10 indistinct, bilobed without setae. Body length: 1.2 mm.

Discussion
Within Pseudoperomyia only P. acutistyla and P. bidentata lack ocelli. Both species can easily be distinguished by the shape of gonostylus and tegmen (see Figs. 1B-C and 2B-C).

Description
Male. Head: ocelli absent; with a narrow sclerotized spot on vertex instead (Fig. 2E). Eye bridge 4-5 facets wide. With 1 row of postocular bristles, 2-3 weaker bristles in a very short second row situated more ventrally. Last flagellomere with small second node or not. Neck of flagellomere 4 little longer than node (Fig. 2D). Node slightly longer than wide, with a double whorl of long setae basally and with irregularly arranged sensory hairs of different lengths, the most proximal sensory hairs forming a rather distinct whorl; among the hair-like sensillae with a few sensory spines and 1 inconspicuous bottle-shaped sensilla distally. Terminal palpal segment a little to obviously longer than segment 2; 2nd segment with a few sensory hairs basally. Thorax: arrangement of lateral setae of the reduced type (Fig. 2F). Legs: claws of middle legs with 2 small teeth mesally. Empodium as long as claws. Wing (Fig. 2G): R,=VA rs; CuA-fork rectangular. Hypopygium: gonocoxites ( Fig. 2A) covered with setae and scales ventrally; distal margin with broadly U-shaped emargination. Gonostylus (Figs. 2A, B) slender, evenly tapering to tip; with apical tooth-like structure with 2 points (in lateral position often appearing to be a simple tooth) and 2 subapical spines little shorter than tooth; with long setae and some scales proximally. Tegmen (Fig. 2C) broad and short; distal margin truncate; parameral apodemes not reaching basal gonocoxal margin. Genital rod (Fig. 2C) weakly sclerotized, very broad and with broadly funnel like invagination running into the distolateral margin of tegmen. Tergite 9 with broadly rounded distal margin and 1 transverse row of setae; anterior membraneous area partly with additional long setae. Tergite 10 strong, especially distinct along lateral margins; bilobed and elongated, with weak setae. Stemite 10 indistinct, bilobed, without setae. Body length: 1.2 mm. Discussion P. bidentata is easily distinguished from the other species by the short, broadly truncate tegmen combined with a very broad genital rod with a wide funnel-shaped invagination. It is necessary to have a specimen in a suitable position on the slide to make out the two points of the apical tooth of the gonostylus.

Description
Male. Head: 3 ocelli. Eye bridge 4-5 facets wide. With 1 row of postocular bristles, 2-4 weaker bristles in a short second row situated more ventrally. Last flagellomere constricted and with small second node. Neck of flagellomere 4 as long as or little longer than node (Fig. 3E). Node with a basal whorl of long setae consisting of 2-3 rows and with irregularly arranged sensory hairs of different lengths, the most proximal sensory hairs sometimes forming an irregular whorl being indistinctly crenulate posteriorly (Fig. 3F); among the hair-like sensillae with a few sensory spines and 1 inconspicuous bottle-shaped sensillum distally (lacking on distal 6 flagellomeres). Terminal palpal segment a little to obviously longer than segment 2; 2nd segment with a few sensory hairs basally. Thorax: arrangement of lateral scutal setae of the reduced type (see Fig. 2F). Legs: claws of middle legs with 2 small teeth mesally. Empodium approximately as long as claws. Wing: J?/= I'A rs\ CuA-fork rectangular. Hypopygium: gonocoxites (Figs. 3 A, C) covered with setae and scales ventrally; distal margin with broadly U-shaped emargination. Gonostylus (Figs. 3 A, C) rather small, a little flattened, tapering to tip; with a curved apical tooth and 2 subapical spines shorter than tooth; with long setae and some scales proximally. Tegmen (Figs. 3B, D) pyramid-shaped, with narrow apex (pointed or truncate); parameral apodemes not reaching basal gonocoxal margin. Genital rod (Figs. 3B, D) sclerotized, very narrow and just reaching tip of tegmen; with narrow funnel-shaped invagination apically; sometimes with a basal enlargement of various shapes. Tergite 9 with broadly rounded to straight distal margin with a slight depression medially (corresponding to weak sclerotization) and 1 transverse row of setae. Tergite 10 weak, bilobed, ovoid, with a few weak setae. Stemite 10 indistinct, bilobed, without setae (occasionally with 1 seta). Body length: 1.1-1.3 mm .

Discussion
P. humilis is the species of the genus that is most frequently encountered. Only males were observed (and collected in large numbers), forming loose swarms when flying in the morning over mixed accumulations of rotten leaves, wood and soil. The tegmen of P. humilis is evenly pyramid-shaped and similar to that of P. intermedia, but broader basally. The apex of the membraneous tegmen is not stable in shape and varies from pointed to truncate. The gonostyli differ in size and shape in a limited degree, and the apical tooth appears stronger, longer or more curved in some specimens. This is largely the result of different positions of these structures in the balsam mounts, but is probably also due in part to infraspecific variation.  (Fig. 4C). Node with a double whorl of setae (or scales?, broken in both specimens) basally and with irregularly arranged sensory hairs of different lengths, the most proximal sensory hairs shortest and forming a rather distinct whorl; among the hair-like sensillae with a few sensory spines and 1 inconspicuous bottle-shaped sensilla distally (absent on distal 6-7 flagellomeres). 3rd palpal segment little longer than 2 nd (Fig. 4D). Thorax: arrangement of lateral scutal setae of the reduced type (see Fig. 2F). Legs: claws of middle legs with 2 small teeth mesally. Empodium as long as claws. Wing: R ,=1 ! A rs\ CuA -fork rectangular. Hypopygium: gonocoxites (Fig. 4A) covered with setae (and scales?) ventrally; united for almost its whole length medioventrally (resulting in a very shallow emargination). Gonostylus (Fig. 4 A) with subbasal lobe dorsolaterally, near apex with a small lobe-like projection ventrally; with a slightly curved apical tooth shifted somewhat dorsad, and 2 inconspicuous subapical spines shorter than tooth. Tegmen (Fig. 4B) short and broad, parameral apodemes not reaching basal gonocoxal margin, nearly parallel-sided, w ith pointed apex forming a nose-shaped, dorsally-directed projec tion. Genital rod (Fig. 4B) weakly sclerotized, with a wide funnel-shaped invagination apically running into margin of tegmen. Tergite 9 with slightly and broadly rounded distal margin and 1 transverse row of setae. Tergite 10 distinct, bilobed, elongated with horn-like projections distally and some weak setae. Stemite 10 indistinct, bilobed, without setae. Body length: 1.0 mm.

Discussion
This is the only species of Pseudoperomyia with such distinct distal projections on tergite 10. The widely opened funnel-like genital rod is similar to that in P. bidentata, probably indicating a closer relationship between these two species than is shown by the cladogram. We did not use this character (shape of opening of genital rod) for the cladistic analysis as the definition of clear character states is not possible because of intermediate forms connecting the extremes.

Discussion
P. intermedia is characterized by the combination of a compact gonostylus with strong apical tooth, a narrow pyramid-shaped tegmen and a rather broad and long genital rod. It has the widest eye-bridge of all congeners. Since only the type-specimen is known, it is not possible to decide if the projecting basal gonocoxal margin is a stable character (or an artefact).
Pseudoperomyia longicornis sp. n.  (Fig. 6E illustrating flagellomere 1). Node with irregularly arranged short setae and short sensory hairs basally, with a double whorl of long setae distad of medial circumference, and distally with irregularly arranged sensory hairs longer than those basally; furthermore with a few sensory spines and 1 inconspicuous small conical sensilla distally (sometimes lacking). 3rd palpal segment longer than segment 2; 2nd segment with a few sensory hairs basally. Thorax: arrangement of lateral scutal setae of the extended type (see Fig. 8D) Legs: claws of middle legs with 3 small teeth mesally. Empodium 3 /i length of claws. Wing: R,=VA rs; CuA-fork rectangular. Hypopygium: gonocoxites (Fig. 6A) covered with setae and scales ventrally, distal margin with a deep U-shaped emargination, its inner margin slightly sclerotized. Gonostylus (Fig. 6A) flattened (when in lateral view); widest in proximal third and tapering to tip; in distal third with rather large tooth-like plate inserted dorsally and reaching apex of gonostylus. Tegmen (Fig. 6B) nearly parallel-sided, with rounded and hardly visible distal margin; at apex with nose-like projection directed dorsad; parameral apodemes running beyond basal gonocoxal margin. Genital rod (Fig.  6B) sclerotized, rather broad and with membraneous cap apically, sometimes with an irregularlyshaped enlargement basally.

Discussion
The species is easily distinguished from its congeners by the very long distolateral horns of tergite 9, but these are not at all clearly visible in some positions. Further specific characters are the large and distinct lobes of tergite 10 running beyond the distal margin of tergite 10 and united by a strong sclerotized arc, and the broad, plate-like apical tooth of the gonostylus. P. longicornis appears to be the sister species of P. platistyla, mainly because they share the inner sclerotization of the gonocoxal emargination. But the structure of gonostylus, tegmen, genital rod and tergite 9 of both species differ greatly (see Figs. 6A and 8A, 6B and 8B, 6C and 8B).
Pseudoperomyia macrostyla sp. n.  (Fig. 7F). Node with irregularly-arranged short setae and short sensory hairs basally, with a double whorl of long setae medially, and distally with irregularly arranged sensory hairs little longer than those basally; additionally with a few sensory spines and 1 inconspicuous small conical sensilla distally (sometimes absent). 3rd palpal segment slightly to obviously longer than segment 2; 2nd segment with a few sensory hairs basally. Thorax: arrangement of lateral scutal setae of the extended type (see Fig. 8D). Legs: claws of middle legs with 3-4 small teeth mesally. Empodium % length of claws.

Discussion
The hypopygium of P. macrostyla is very remarkable because of the unusual modifications of the gonocoxites (Fig. 7 A) 8C) as long as node. Node with irregularly arranged setae (or scales?, broken) and short and weak sensory hairs in basal half, with a double whorl of stronger setae (or scales?) in distal half, and distally with irregularly arranged sensory hairs a little longer than those basally; additionally with a few sensory spines and 1 inconspicuous small conical sensilla distally (absent on distal flagellomeres beginning with 5th). Terminal palpal segment longer than segment 2; 2nd segment with a few sensory hairs basally. Thorax: arrangement of lateral scutal setae of the extended type (Fig. 8D). Legs: claws of middle legs with 3 small teeth mesally. Empodium approximately as long as claws.
Wing: R ,-2 rs\ CuA -fork rectangular. Hypopygium: gonocoxites (Fig. 8A) covered with setae and (probably) scales ventrally, with stronger and longer setae along distal margin; distal margin with a very deep U-shaped emargination (basomedial link narrow), its inner margin distinctly sclerotized; dorsal gonocoxal link looped in a complicated manner. Gonostylus (Fig. 8A) strongly flattened and broad (appears narrow and slender in lateral view); densely covered with setae outside and inside and with scales in proximal half; apically with flattened tooth-like structure inserted rather dorsally, with 2 short spines inside subapically. Tegmen ( Fig. 8B) with characteristic shape: parameral apodemes directed outwards and reaching basal gonocoxal margin, followed distally by a waist-like medial constriction, distal half with triangular enlargement with rounded apex, latter appearing as an indistinct nose-like projection directed dorsad. Genital rod (Fig. 8B) membraneous and very short, reaching 1/3 of tegmen length, with membraneous cap. Setae of tergite 9 (Fig. 8B) few in number (up to 3?) and reduced in extension to a small central area, on both sides of that area 1 very short sclerotized horn-like projection directed ventrad; inner (anterior) margin of tergite 9 strongly sclerotized; 2 large disto-lateral membraneous lobes linked with the distal gonocoxal margins. Tergite 10 ( Fig. 8B) distinct, elongated, bilobed and both lobes connected by horseshoe-shaped sclerotization dorsally; with a few rather long setae distally. Stemite 10 not visible. Body length: 1.3 mm.
Discussion P. platistyla is easily distinguished from its congeners by the constricted shape of tegmen, the strongly flattened gonostyli and the extremely short genital rod.

Phylogeny of Pseudoperomyia and the Micromyidi
There is no doubt that the species of Pseudoperomyia are correctly placed within the supertribe Micromyidi. The Micromyidi are a monophylum well supported by a number of synapomorphies such as the forked CuA, the reduction of CuP and A, the reduction of ventral macrotrichia on R}, the presence of scales, the number of male flagellomeres reduced to 12, and the usually close connection of the lobes of tergite 10 with tergite 9 (Kleesattel, 1979;Jaschhof, 1998 .03 t t ro < u 'C :  Fig. 9 with branch length proportional to the number of characters that change unambiguously on branch (see scale); decay index on the left side and bootstrap proportion on the right side of each branch. Note that the number of changes involving the terminal taxa is not comparable with that elsewhere, since their autapomorphies were excluded from the analysis.
In addition, all micromyids have a pattern of sensory pores on the wing veins which differs from group to group but which is stable within each species and within each group of related species. Pseudoperomyia shares all these derived characters, which contrast with the plesiomorphous conditions of the outgroup. Questions concerning the monophyly and sister group of Pseudoperomyia were incorporated into a test of the recently-proposed hypothesis on the phylogeny of the supertribe Micromyidi (JASCH-HOF, 1998).
For the parsimony analysis, we considered a total of 47 adult male characters (Append. 1) for all the known Pseudoperomyia species and for the type species of each genus of the Micromyidi (Tab. 1). Peromyia levellei (type species of Peromyid) was replaced by Peromyia aurantiaca (type species of the synonym Joannisia), since the former is too incompletely known so far as the morphological details needed for the analysis are concerned. Anaretella defecta was selected as the outgroup. It represents the tribe Lestremiini and has mainly plesiomorphous characters within the Cecidomyiidae (Kleesattel, 1979;Jaschhof, 1998). The characters used for the computer analysis were for the most part those traditionally used for taxonomic studies within the Lestremiinae. We redefined some of them to get clear character states (see character no. 28 or 43), and we added a few characters by considering homologous structural details of the same body part independently (see, for example, 10 and 11 or 12 and 14). The obvious lack of a good number of comparable characters with precisely defined character states does not permit well-supported monophyletic groups and sister group relationships, i. e. a more stable cladogram. Adding only one additional character with obviously incorrectly evaluated states into the matrix resulted in the replacement of many dichotomies by polytomies in such a poorly supported tree. Basic comparative studies to identify homologous structures are needed in order to achieve a substantial improvement in evaluating characters such as, for example, the large number of different types of flagellomeral sensillae or the confusing structural diversity of gonostylus and tegmen. So far as these characters are concerned (no. 13,31-35 and 36-38), we were unable to find a more objective system of structural types on the basis of the information available, based on real homologies rather than on independent superficial similarities. Without any doubt there is still a large amount of information left in these structures, as is probably the case with other character complexes that have been entirely overlooked so far. A heuristic search by PAUP (random addition sequence with 1000 replicates, TBR, COLLAPSE and MULPARS options in effect, steepest descent option not in effect, all characters unweighted) resulted in the same 4 minimum length trees when all the characters were unordered (150 steps, Cl 0.45, RI 0.66) and when the multiple state characters (7, 8, 9, 14,19, 20, 32, and 43) were ordered (156 steps, Cl 0.44, RI 0.69). In all the trees (strict consensus tree Figs. 9 and 10), the species of Pseudoperomyia appear as a well supported monophyletic group (bootstrap proportion in 1000 replicates 89, decay index 4). The internal phylogeny of Pseudoperomyia is only partly resolved, as there is a basal trichotomy. Only two of the internal clades (41 and 42) have both the bootstrap proportion over 50 and decay indices more than 1.
Pseudoperomyia appears in all 4 trees as the sister-group of Anodontoceras within the tribe Micromyini (sensu Jaschhof, 1998). The sister-group relationship is rather poorly supported (bootstrap proportion <50, decay index 1), as is the case with most of the interrelationships of supraspecific taxa within the Micromyidi.
Within the Micromyini both Anodontoceras and Pseudoperomyia share the globular / subglobular type of flagellomeral node as a synapomorphous character, which also occurs through convergence in Skuhraviana (tribe Bryomyiini) and in Peromyia (tribe Peromyiini).
Without discussing all the results in detail, it is apparent that the result of the present parsimony analysis is in rather good agreement with the relevant part of a recent hand-made cladogram for the entire Lestremiinae (Jaschhof, 1998). The main differences between the two analyses involve the position of Campylomyza, previously referred to a group of genera including Neurolyga, Micropteromyia and Excrescentia (together forming the tribe Campylomyzini), and the position of Aprionus, previously assigned to the Aprionini together with Tekomyia and Mycophila. Peromyia (the only genus of the Peromyiini) was previously considered to be the sister-group of the Bryomyiini + (Micromyini + Aprionini).