New species of Peromyia from Tasmania, with a revision of Schiner'

Peromyia K ie f f e r , 1894, a speciose, cosmopolitan genus o f micromyine cecidomyiids, is shown to occur in Tasmania. This is the first record of the genus in Australia and the first contribution to the taxonomy of Australian Micromyinae for 120 years. Nine o f the 12 species found proved to be new to science: Peromyia grovei sp. n., P nitoda sp. n., P. indecorata sp. n., P warraensis sp. n., P bashfordi sp. n., P despecta sp. n., P. debilis sp. n., P. cinalata sp. n., and P tasmanica sp. n. Four of the new species are assigned to the Peromyia dissona group, which is established for a chiefly western Pacific species radiation. Other Tasmanian peromyias are assigned to species groups that to present knowledge are widely distributed accross the Holarctic Region and occasionally beyond. The types of the Micromyinae species from Australia by Ignaz R. S c h in e r and Frederick A. A. S k u se are revised. None of the 11 species belongs to the genus Campylomyza M e ig e n , 1818, where they were originally assigned. Campylomyza grandiuscula S k u se , 1890 is actually a species of the genus Pteridomyia J a sc h h o f , 2003. The remaining species either belong to the genus Peromyia or their generic affiliation could not be determined. Peromyia aeratipennis (Sk u se , 1888) comb. n. takes priority over Peromyia muscorum (K ieffer , 1895) syn. n., and Peromyia impexa (Sk u se , 1888) comb. n. takes priority over Peromyia palustris (K ie f f e r , 1895) syn. n. Peromyia pellax (Sk u se , 1888) comb. n., syn. n. and Peromyia tilis (Sk u se , 1888) comb. n., syn. n. are identical with Peromyia impexa. Campylomyza sidneyensis S c h in e r , 1868, Campylomyza amplipennis S k u se , 1888, Campylomyza crocea S k u se , 1888, Campylomyzaperpallida S k u se , 1888, Campylomyzapersimilis S k u se , 1888, and Campylomyza vicina S k u se , 1888 are relegated to nomina dubia.


Introduction
Cecidomyiids belonging to the subfamily Micromyinae are tiny midges with mycetophagous larvae that develop in humous soil and rotting plant debris, typically dead wood. Their pre ferred habitat are humid forests worldwide. Most of the 500 or so named species were described from the Holarctic Region. Elsewhere micromyines received much less attention by taxonomists, Australia being no exception. O f many of the named species nothing more is known than male adult anatomy, which is used for species identification and classification, and fragmentary data on phenology and distribution. To look on the bright side, the Micromyinae of the Holarctic Region have been subject to comprehensive taxonomic revision in recent years, resulting in a natural clas sification of the subfamily ( S k u s e , during his short, prolific preoccupation with Australian Diptera in the late 1880s (cf. M a rk s 1991), was the first entomologist to address the taxonomy of Australian Cecidomyiidae and Micromyinae. It is notable that his "Diptera of Australia" (Sku se 1888), intended to become a series of family monographs, started with just that group of delicate midges, considering he had the choice among plenty of unworked families. Sku se (1888, 1890) distinguished among 11 spe cies of Micromyinae, all from Sydney and its environs, including one species that S c h in e r (1868) had described earlier. Both S c h in e r and Sku se assigned their species to the genus Campylomyza M e ig e n , 1818 -a taxon then equivalent to the present supertribe Micromyidi now contain ing some 30 genera. Modern workers find the original descriptions of S c h in e r ' s and S k u s e ' s Campylomyza entirely inadequate for the identification of species and the genera they actually belong to. Consequently, in the World Catalog of Cecidomyiidae they are listed among the "un placed species of Lestremiinae" (G a g n é 2004: 36). Moreover, it seemed unlikely that there was a true Campylomyza among, because this genus is at present chiefly Holarctic in distribution. The only true Campylomyza known to occur in Australasia is Campylomyza flavipes M e ig e n , 1818, a widespread Holarctic species introduced into New Zealand (Ja sc h h o f & Ja sc h h o f 2003). To shed light on the dubious Campylomyza species from Australia I endeavored to examine their type specimens. The results are presented in this article.
According to Au st in et al. (2004), Cecidomyiidae are among the seven major families in the Australian Diptera fauna that are taxonomically orphaned. In fact, there has been some progress in recent years in exploring Australia's diversity of gall-inducing Cecidomyiinae (cf., for in stance, K o l e s ik & V e e n str a -Q uah 2008), but A u st in et al. are perfectly right with respect to the non-phytophagous species classified in the subfamilies Lestremiinae, Micromyinae and Porricondylinae. S k u s e ' s late 19th century contribution to the Micromyinae of Australia has in deed had no successor ever since (cf. Bu g l e d ic h 1999). So this fauna, that I estimate to contain a few hundred species, many to be classified in new genera, remained virtually unknown to this day. Peromyia, a cosmopolitan, open-ended genus (cf. B ic k e l 2009), remained hitherto unreported from Australia and Tasmania, even though peromyias are among the most common cecidomyiids there if one may judge from Malaise samples taken in forests (Ja sc h h o f , pers. obs.). This present first account of Peromyia in Tasmania deals with only a small fraction of the Micromyinae of Australia. It is my intention to follow this by further small contributions. To proceed in such a piecemeal fashion might be considered by some to be inadequate for documenting a fauna that is unworked even at the level of genus. However, the funding needed for a more thorough study ap pears unobtainable at present, even though such a study was feasible within a modest time-frame (cf.  (2009), which is also the reference regarding Micromyinae morphological terminology and classification. Morphology was studied by compound microscope at 400 times magnification. Holotypes of the new species are deposited with the Australian Museum Sydney (AMS), paratypes with the AMS, NHRS and Senckenberg Deutsches Entomologisches Institut Müncheberg (SDEI), other voucher specimens with the NHRS and SDEI.
Information on a syntype of Campylomyza sidneyensis S c h in e r was provided by Dr Peter K o l e s ik , who had this specimen on loan from the Naturhistorisches Museum Wien (NHMW). S k u s e ' s Micromyinae (and 1 species of Lestremiinae) were with Dr Michel Baylac, Muséum National d'Histoire Naturelle, Paris (M NHN), on loan from the Australian National Insect Collection Canberra (ANIC). The specimens, apparently in the original state as S k u se had prepared and labelled them, were glued on small squares of cardboard and attached to insect pins together with two labels. Type specimens were labelled as such and found to be tagged with information on locality, occasionally also date and collector, in accordance with the information given in the original publications. The Sku se material, as I obtained it from Baylac, contained only one speci men per species labelled "type", which is inconsistent with the catalog by Bu g l e d ic h (1999) who usually refers to two syntypes -a discrepance I am unable to explain. G a g n é (2004) is certainly correct in referring to the original specimens as "holotypes", since S k u s e , in his original publica tions, writes in each case in the singular form of the "specimen" from which he drew the species description. I soaked the dry specimens in water until they were released from the glue, tried with little success to macerate them with lactic acid, dehydrated them in ethanol and mounted them on microscope slides following the method by J a sc h h o f & Ja sc h h o f (2009). I did not study three other Micromyinae specimens from S k u s e ' s material, which did not bear identification or type labels. J a sc h h o f , M.: New species of Peromyia from Tasmania The Micromyinae from Australia described by S c h i n e r and S k u s e O f the 3 female adult syntypes in the NHM W referred to by Bugledich (1999) nothing re mains. Kolesik (in litt.) describes the "specimen" he had on loan from Vienna as follows: "One pin with 3 paper labels and one plastic triangular specimen holder. No insect was attached to the specimen holder. One label reads "Campylomyza sidneyensis", one reads "X" and one is a simple red square." The other two syntypes, provided they once existed, are not present nor registered as loan in the NHM W (Sehnal, in litt.) and I regard them as lost. Schiner's description of this species is by modern standards useless for identification. Therefore I consider Campylomyza sid neyensis Schiner to be a dubious name, or nomen dubium.
Campylomyza aeratipen n is Skuse, 1888: The holotype is labelled "Campy: aeratipennis ? TYPE. F. A. A. Skuse" and "Eliz: Bay. Dec 87. M ". Preserved are the head with 7 flagellomeres of one of the antennae, the thorax and the abdo men. The antennal structure is typical for a Peromyia. The nodes of the antennal flagellomeres are bicolored, the basal halves darker than the apical halves (Jaschhof 1998: fig. 219h). Two sclerotized, disk-shaped spermathecae are present (l. c.: fig. 219i). These characters in combina tion are typical of Peromyia muscorum (Kieffer, 1895). I have no doubt that Peromyia aeratipennis (Skuse) and Peromyia muscorum (Kieffer) are identical, so the former name becomes valid for this common, widespread species. The species is apparently widely distributed on the Australian continent as is indicated by specimens I have seen from Queensland (unpubl.). As shown below Peromyia aeratipennis occurs also in Tasmania.
Campylomyza am plipennis Skuse, 1888: The cardboard holder labelled "Cy: amplipennis ? TYPE. F. A. A. Skuse" and "Mid: Harb. S." is devoid of the specimen. S k u s e ' s original description does not contain any information that might help to recognize this species, and even its generic affiliation remains uncertain. Consequently, I relegate this name to nomina dubia.
Campylomyza crocea Skuse, 1888: The holotype is labelled "Campy: crocea ? TYPE. F. A. A. Skuse" and "Eliz: Bay. S." The speci men is strongly shrivelled and lacks most of the appendages. It doubtless belongs to a species of Peromyia, but I am unable to detect any species-specific characters. S k u s e ' s description is also useless for identification, so Campylomyza crocea is another nomen dubium.
Campylomyza gran d iu scu la Skuse, 1890: The holotype specimen is labelled " C grandiuscula ? TYPE. F. A. A. Skuse" and "Aug. 21". Preserved are the thorax, some of the legs and the abdomen, including terminalia. The holotype specimen, labelled "Campy: impexa ? TYPE. F. A. A. Skuse" and "Eliz: Bay.", is almost complete, but strongly shrivelled. Male genitalic characters, which are clearly visible, indi cate Campylomyza impexa is identical to the species European workers know as Peromyia palustris (K ie f f e r , 1895). So Peromyia impexa (Sku se ) becomes the valid name of this common, wide spread species. I have seen specimens from Queensland (unpubl.) and Tasmania (see below).
C am pylom yzapellax Skuse, 1888: The holotype specimen, labelled "Campy: pellax ? TYPE. F. A. A. Skuse" and "Eliz: Bay. M /S", is strongly shrivelled, but almost complete except for the apices of the antennae and one wing that are lacking. The terminalia, which are well preserved, indicate that Campylomyza pellax is identi cal with Peromyia impexa (Sk u s e ) (see above).
Campylomyza p e rp a llid a SKusE, 1888: The holotype is labelled "Cy: perpallida ? TYPE F. A. A. Skuse" and "Sydney. S". Preserved are the head without antennae, the thorax and one wing that lacks the apical third. These fragments indicate the specimen belongs to a species of Peromyia, but any structures that might provide species-specific characters are absent. Sk u s e ' s species description is also useless for identification. Therefore I regard Campylomyza perpallida as a nomen dubium.
C am pylom yzapersim ilis Skuse, 1888: The holotype, labelled "Cpy: persimilis ? TYPE F. A. A. Skuse" and "Sydney.", is almost com plete, but strongly shrivelled. It belongs to a species of Peromyia, but which species is impossible to determine because characters of species-specific merit are absent. The species description by S k u se is also uninformative with respect to species-specific features. Consequently, Campylomyza persim ilis is another nomen dubium. J a sc h h o f , M.: New species of Peromyia from Tasmania Campylomyza su b tilis Skuse, 1888: The holotype is labelled "Campy: subtilis ? TYPE. F. A. A. Skuse" and "Eliz: Bay. M /S". The specimen is strongly shrivelled and lacks he head and wings. The gonostyli are not preserved, but the remaining genitalic structures indicate that Campylomyza subtilis is identical with, and conse quently another junior synonym of Peromyia impexa (Sk u s e ) (see above).
Campylomyza vicin a Skuse, 1888: The holotype of this species is labelled "Campy: vicina ? TYPE. F. A. A. Skuse" and "Eliz: Bay. M /S". Preserved are the head without antennae, the thorax and the abdomen including a frag ment of the terminalia, all strongly shrivelled. The specimen belongs certainly to a species of Peromyia, but the preserved fragments are insufficient for species identification. Sk u s e ' s species description is also of no help in this context. Therefore I relegate Campylomyza vicina to nomina dubia.

The genus Peromyia in Tasmania
With 150 named species, including one Tertiary fossil, this cosmopolitan genus is the largest of the subfamily Micromyinae. The fact that most Peromyia species were described from the Holarctic Region is due to too little study elsewhere ( A key to the Peromyia species in Tasmania is not presented at this stage, as any key for randomly caught specimens is likely to fail due to the large number of species not yet described and named.

P e ro m y ia d is s o n a species group
Diagnosis: Male antennal flagellomeres bear 2 whorls of hair-shaped translucent sensilla, one distal whorl of long sensilla and one basal whorl of shorter sensilla (Fig. 1A). Position of hair-shaped sensilla at the very bases of the flagellomere nodes is an unusual character state among peromyias, even though there are instances of sensilla that irregularly intermingle with the short subbasal setae that are universally present (see, for instance, Fig. 8A). Postocular bristles are usually arranged in 2 rows. Male terminalia have a diverse, often unusual structure, which is less pronounced in the representatives from Tasmania. This group gathers several species of previously uncertain affin-  Peromyia grovei is characterized by a combination of male genitalic characters as follows. The ven tral emargination extends to the midlength of gonocoxites (Fig. 1B), the gonostyli are flattened and excavated mesally (Fig. 1B), and the tegmen has dark lateral stripes (Fig. 1C). Description: Male. Body size: 0.7-0.8 mm. Head: Postfrons asetose. Eye bridge 2-3 ommatidia long. Postocular bristles in 2 rows. Neck of fourth antennal flagellomere longer than node (Fig. 1A). Node sub globular, with 1 basal whorl of short hair-shaped sensilla, short subbasal setae, 1 mesal single to triple whorl of long setae and 1 distal whorl of long hair-shaped sensilla. Maxillary palpus 4-segmented, all segments with hair-shaped translucent sensilla. Wing: AntC ending at juncture with R5 or extending slightly beyond R5. ApicRl = 3 times the length of Rs. Legs: Pretarsal claws without teeth.
Female and immature stages. Unknown. Etymology: The species is named to honour Dr Simon J. G r o v e, Conservation Biologist with Forestry Tasmania, Hobart, whose broad, keen interest in the Natural History of Tasmania helped with the initiation of this study.  Peromyia nitoda is characterized by the following male genitalic characters in combination: the ventral emargination extends to the midlength of gonocoxites ( Fig. 2A), the gonostyli are thick throughout their lengths and slightly curved ( Fig. 2A), and the tegmen has dark mesal stripes (Fig. 2B).
Description (mentioned are only those features which differ from that in P. grovei): Male. Head: Maxillary palpus 4-segmented, first to third segments with hair-shaped translucent sensilla.
Female and preimaginal stages: Unknown.

Etymology:
The species name is made up of the first two letters each of the forenames of Niklas J o n ss o n , Tobias M a lm and Dawn W il lia m s, all of Naturhistoriska Riksmuseet Stockholm, Sweden, who are the collectors of a part of the peromyias studied here.  Peromyia indecorata is characterized by the following male genitalic characters in combination: the ventral emargination extends beyond the midlength of gonocoxites (Fig. 3A), the gonostyli have a simple, subcylindrical outline (Fig. 3A), and the tegmen is largely unmodified except for the apical margin of ventral plate which is occasionally discernible (Fig. 3B).
Description (mentioned are only those features which differ from that in P grovei): Male. Body size: 0.9-1.1 mm.
Female and preimaginal stages: Unknown.

Remark on variation:
There is some individual variation in the extent (depth) of the ventral emargination of gono coxites. Some gonostyli are thicker than the one depicted (Fig. 3A) and occasionally are slightly curved.  Even so, the gonostylus structure in these two species is fundamentally different: while the subbasal lobe in P. warraensis emerges from the dorsal surface of gonostylus, it emerges from the ventral surface in P. praeclara. Further differences lie in the outline of gonocoxites and tegmen. For instance, the ventral emargination of gonocoxites is shallow in P. warraensis and deep in P praeclara, and the tegmen is short and rounded in P warraensis and long and pointed in P praeclara.
Description (mentioned are only those features which differ from that in P grovei): Male. Head: Maxillary palpus 4-segmented, first segment with hair-shaped translucent sensilla.
Female and preimaginal stages. Unknown.

Etymology:
The name is derived from the type locality, Warra.   Male. Body size: 0.7-0.8 mm.
Female and preimaginal stages: Unknown.
Remark on relationships: Male morphology suggests P. bashfordi and P. obunca are very closely related to one another, pre sumably sister species. So no doubt persists any longer that P. obunca is a member of the interme d ia group to which it was tentatively assigned

by J a sc h h o f & Ja sc h h o f (2004).
Etymology: This species is named to honour Richard Ba sh f o r d , Senior Silvicultural Technician at Forestry Tasmania, Hobart, who is in charge of the invertebrate collections at Warra LTER site.

Types:
Holotype: Male, Australia, Tasmania, Warra LTER site, Manuka Road, 1 March 2005, Malaise sample FT35682, R. Bash fo r d (in AMS). Paratypes: 2 males, same data as the holotype; 4 males, same data but sample FT35684; 2 males, same data but sample FT35685; 3 males, same data but sample FT35693 (in AMS and SDEI). in particular Peromyia boreophila J a sc h h o f , 2001. The following characters in combination are species-specific for P despecta: postocular bristles are present in 2 rows; the whorl of short hair-like sensilla on the antennal flagellomeres is situated mesally, just beyond the double whorl of setae (Fig. 6A); the gonostyli are elongate and slightly curved (Fig. 6B); and the ventral plate of the tegmen is weak and comparatively long (Fig. 6C).
Wing: AntC ending at juncture with R5 or extending beyond R5 for once the vein width. ApicRl = 3.5-4.0 times the length of Rs. Legs: Pretarsal claws without teeth.
Female and preimaginal stages: Unknown. Etymology: From the Latin, meaning unnoticed.  The thorax in proportion to the head in Peromyia debilis is unusually small and the wings are rather narrow, which represents the first case of atrophy observed within the genus Peromyia. The whorl of short hair-shaped sensilla on the antennal flagellomeres is irregular and intermingles with subbasal setae (Fig. 7A), which is unusual among species of the modesta group where P. debilis is assigned. This character, in combination with curved gonostyli, is species-specific.
Female and preimaginal stages: Unknown. Etymology: The name is Latin, composed of cinis, ash, and alatus, winged, and refers to the grey-tinged wings of this species. P e ro m y ia ta s m a n ic a sp. n. Male terminalia in Peromyia tasm anica are unmistakable because of the large, subtriangular gonostyli (Fig. 9B) in combination with the large, curved tegmen lacking a distinct ventral plate (Fig. 9C). Due to the peculiar set of male genitalic characters it is difficult to place this species in any of the currently recognized species groups. Description: Male. Body size: 0.8 mm.
Female and preimaginal stages: Unknown. Etymology: The name is derived from Tasmania where specimens were collected.