New species of Monepidosis Mamaev , 1966 and Antipodosis gen . nov . , a closely related genus from New Zealand ( Diptera , Cecidomyiidae )

Three new species of Monepidosis Mamaev, 1966, a holarctic genus of Porricondylinae (Diptera, cecidomyiidae), are described: M. heterocera sp. nov. from sweden and Germany, M. scepteroides sp. nov. from sweden, and M. shikokuensis sp. nov. from Japan. a new porricondyline genus, Antipodosis gen. nov., is introduced for eight new species from New Zealand, named A. australis gen. et sp. nov., A. elongata gen. et sp. nov., A. granvillensis gen. et sp. nov., A. pureora gen. et sp. nov., A. rakiura gen. et sp. nov., A. rotoiti gen. et sp. nov., A. rotoroa gen. et sp. nov., and A. waipapa gen. et sp. nov. Male genitalic morphology indicates that Monepidosis and Antipodosis gen. nov. are closely related, together forming the Monepidosis group of genera, which stands out from the other Porricondylini. Monepidosis spatulata spungis, 2006, a species originally described from Latvia and Lithuania, is for the first time reported to occur in Sweden.


Introduction
a recent revision of the genus Monepidosis Mamaev, 1966 recognized 14 valid species, 13 in the Palearctic and one in the Nearctic region (Jaschhof & Jaschhof 2013;Gagné & Jaschhof 2014).another species of Monepidosis was described from sweden soon after (Jaschhof & Jaschhof 2015).Previous authors expected a considerable number of unnamed Monepidosis still to be found in the holarctic realm (spungis 2006;Jaschhof & Jaschhof 2013), an expectation validated here with the description of three new species from sweden, Germany and Japan.Monepidosis are morphologically distinct from the other Porricondylini (cecidomyiidae, Porricondylinae) in that male genitalic structures, such as gonostyli, gonocoxites, parameres, and ejaculatory apodeme, are heavily armed in a specific manner (Jaschhof & Jaschhof 2013).Most notably, parameres are present as two pairs of large, sclerotized tusks interlinked with each other, and the ejaculatory apodeme is developed into a long, sclerotized rod whose apex is variously modified (Jaschhof & Jaschhof 2013: fig.88A).Genitalic structures similar to those in Monepidosis have now been found in eight different species of undescribed Porricondylinae from New Zealand, which are obviously closely related to each other and thus classified together in a discrete new genus, Antipodosis gen.nov.Monepidosis and Antipodosis gen.nov.appear to be, if not sister groups, members of a distinct natural subset of the tribe Porricondylini, called here the Monepidosis group.
In the present paper, the newly discovered taxa are described and named, so as to continue with the taxonomic inventory of Palearctic and especially swedish Porricondylinae within the framework of 'The Gall Midge Project' (http://www.stationlinne.se/sv/forskning/the-gall-midge-project,accessed on 25 Nov. 2015), and to take a first step into the study of porricondyline biodiversity in Australasia.This study shows that porricondyline interrelationships are more revealing when studied globally.

Material and methods
specimens for this study, exclusively male adults, were either picked out from general insect samples obtained by Malaise traps or were collected by sweepnet and aspirator.Most of the specimens came from various different projects initiated by the author, specifically 'The Gall Midge Project', an ongoing venture to inventory the cecidomyiidae of sweden (2015)(2016)(2017); the 'Lestremiinae of Japan ' project (1998-2000;see Jaschhof 2000); and the 'Lestremiinae of New Zealand' project (2001Zealand' project ( -2003; see Jaschhof & Jaschhof 2003).other specimens were provided by 'The swedish Malaise Trap Project' (sMTP; see http://www.stationlinne.se/sv/forskning/the-swedish-malaise-trap-project-smtp,accessed on 25 Nov. 2015); were received on loan from the senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany (sDEI); or were donated by Dieter Doczkal, Zoologische staatssammlung München, Germany.
Types and other voucher specimens of the species discussed here are available as microscopic preparations (Canada balsam inclusions) and shall be finally deposited in the following collections: Entomological collection of Kyushu University, fukuoka, Japan (KUEc); Naturhistoriska Riksmuseet stockholm, sweden (NhRs); New Zealand arthropod collection, Tamaki, auckland, New Zealand (NZac); and sDEI.
The procedures of specimen preparation and identification as well as the morphological terminology used in adult Porricondylinae were recently described by Jaschhof & Jaschhof (2013).several morphological terms, some of which specific to the porricondylines treated here, are explained in Figs

Diagnosis
Larvae and females of Antipodosis gen.nov.are unknown, so this genus is based solely on characters of males.Antipodosis gen.nov.are small, inconspicuous Porricondylini, exhibiting the characters typical of that tribe and coming under the group of genera with 14 flagellomeres and without basitarsal spines ("group aa" in Jaschhof & Jaschhof 2013).Antipodosis gen.nov.stands out from most other Porricondylini in that vein cua 1 runs parallel to cua 2 rather than approaching or joining it (fig.1a versus 1B), and the eye bridge is longer dorsally (5-10 versus 2-3 ommatidia).The structure of the genitalia is genus-specific.Most notably, the two gonocoxal apodemes, which in other Porricondylini are long bars separated from one another (fig.12), tend to merge into a single plate (fig.2); parameres are typically present as two pairs (a ventral and a dorsal pair) of strongly sclerotized tusks interlinked with each other (fig.2); and the ejaculatory apodeme, whose length exceeds that of the gonocoxites, is typically a strongly sclerotized rod with apical modifications.

Differential diagnosis
The genus Antipodosis gen.nov. is similar to Monepidosis (Jaschhof & Jaschhof 2013), differing from it as follows: the eye bridge is longer; sensory hairs (= setae with hooded sockets) on the flagellomeres are dispersed rather than aligned to form a single whorl (fig.1c versus 1D); wings are wider (with length/width ratios usually < 3.0); cua 1 does not approach but runs parallel to cua 2 (fig.1a versus 1B); the gonostylar apex bears typically a pectinate tooth (figs 2a, 7a), but never a large plate-like spine (figs 10a-B, 11B); gonocoxites have a distinct ventral emargination (fig.2a) and have no central processes (fig.10a); and the anterior portions of the gonocoxal apodemes are either short and separated (fig.6a), or merged into a single, long plate (fig.2a).

Etymology
The name Antipodosis is composed of Antipod-, from the Latin antipodes, for 'antipodes', and the ending -osis, from Monepidosis, a closely related genus.Gender is feminine.

Distribution and phenology
The genus Antipodosis gen.nov. is to present knowledge endemic to New Zealand. of eight species identified, two occur on the North Island and six on the South Island including Stewart Island.Almost European Journal of Taxonomy 192: 1-24 (2016) all the specimens known of Antipodosis gen.nov.were collected in native forest in austral summer (Nov.-Jan.).

Phylogenetic remarks
The peculiarity of Antipodosis gen.nov.that gonocoxal apodemes tend to merge into a large plate is a derived character state not known in other Porricondylini, thus an autapomorphy.from what is currently known of world Porricondylini, it appears that Antipodosis gen.nov. is most closely related to Monepidosis.compelling evidence of this relationship is provided by the parameres and the ejaculatory apodeme, whose basic structure is identical in the two genera.Both are therefore combined in what is here called the Monepidosis group.The relationships of this group to other Porricondylini remain obscure for the time being.With respect to male morphology, Antipodosis gen.nov.has retained more ancestral traits than Monepidosis.for example, in Antipodosis gen.nov. the two gonocoxites are largely separated by the ventral emargination, whereas in Monepidosis they are lengthwise connected, with the connecting bridge even further modified to bear a pair of processes.Furthermore, in Antipodosis gen.nov. the gonostylar apex is equipped with a pectinate tooth of fine, separate spines, which is the structure found in many other Porricondylinae and Porricondylini, whereas in Monepidosis the spines are merged into a large, plate-like tooth that encircles much of the gonostylus' distal half.concurrent in Antipodosis gen.nov.and Monepidosis, the basic patterns of parameres and ejaculatory apodeme undergo various modifications, which may be so pronounced that the generic affiliations of the respective species are obscured (see Antipodosis elongata gen.et sp.nov., fig.3; Monepidosis shikokuensis sp.nov., fig.12).one may argue that such extreme variations are likely to evolve over long periods of time and both Antipodosis gen.nov.and Monepidosis might be phylogenetically old lineages.

Diagnosis
A combination of genitalic characters is specific to A. australis sp.nov., as follows (fig.2a-B). of the gonocoxites (Fig. 2A), the processes are small but prominent (↓); the ventral emargination, whose shape is reminiscent of an inverted bell, has a broad glabrous rim that almost reaches the anterior gonocoxal margin (↓), and the apodemes are merged into a large subrectangular plate (↓).The ejaculatory apodeme has a distinct subapical extension (↓, Fig. 2B).

Etymology
The name australis is a Latin adjective meaning 'southern', which refers to the distribution of this species in the two southern main islands of New Zealand.

Etymology
The name elongata, a Latin adjective meaning 'elongate', refers to the genitalia of this species.

Etymology
The name granvillensis, an adjective, is derived from Granville state forest in the West coast region of south Island, which is the type locality of this species.

Diagnosis
A combination of genitalic characters is specific to A. pureora gen.et sp.nov., as follows (fig.5a-B). of the gonocoxites (Fig. 5A), the processes are small and subtriangular (↓), the large ventral emargination is perfectly U-shaped (↓), and the apodemes are merged into a large plate with rounded margins (↓).The base of the ejaculatory apodeme is strongly widened (↓, Fig. 5B), which is unique in Antipodosis gen.nov.

Differential diagnosis
Antipodosis pureora gen.et sp.nov. is similar to both A. australis gen.et sp.nov.and A. rotoroa gen.et sp.nov.The three species differ from each other in the ejaculatory apodeme, which is strongly widened basally in A. pureora gen.et sp.nov.(fig.5B), markedly widened subapically in A. australis gen.et sp.nov.(fig.2B), and two-pointed apically in A. rotoroa gen.et sp.nov.(Fig. 8B).Other interspecific distinctions are described under each of the species.

Etymology
The name pureora is from the Maori language, referring to Pureora forest, one of the largest intact tracts of native podocarp forest in the North Island and collection site of this species.

Other characters
Body length.about 1.7 mm.
head.Eye bridge 4-5 ommatidia long dorsally.Apices of antennae missing.Neck of fourth flagellomere 1.7 times longer than node.Palpus longer than height of head.

Etymology
The species epithet rotoiti is from the Maori language, meaning 'the little lake', with reference to Lake Rotoiti in northern south Island where this new porricondyline was collected.

Other characters
Body length.2.1 mm.head.Eye bridge 4 ommatidia long dorsally.Circumfila on flagellomeres 1-14, often with short posterior extensions.Neck of fourth flagellomere 1.1 times longer than node.Palpus 1.5 times longer than height of head.

Diagnosis
The gonocoxal processes are more prominent in A. rotoroa gen.et sp.nov.(↓, Fig. 8A) than in any other Antipodosis gen.nov.The gonostylus is slightly curved, tapered towards the apex, and equipped with a long pectinate tooth (↓, Fig. 8A).The apex of the ejaculatory apodeme is two-pointed (↓, Fig. 8B).At the bases of the dorsal parameral tusks are small accessory processes, which are weakly sclerotized and blunt-ended (↓, Fig. 8B).

Differential diagnosis
see A. australis gen.et sp.nov.(fig.2) and A. pureora gent.et sp.nov.(fig.5) for two species with genitalic structures superficially similar to that found in A. rotoroa gen.et sp.nov.

Etymology
The name rotoroa, which means 'the long lake' in the Maori language, refers to Lake Rotoroa, whose vicinity provides the only habitat yet known of this porricondyline.

Diagnosis
This is the only species of Antipodosis gen.nov., whose gonostyli have no apical structure, such as a pectinate or solid tooth (↓, Fig. 9A).As another peculiarity, the apical portion of the ejaculatory apodeme is conspicuously enlarged and spoon-shaped (↓, Fig. 9B).

Etymology
The name waipapa, from the Maori language, refers to Waipapa Reserve in Pureora forest where the type specimens of this species were collected.

European Journal of
Wing.Length/width 2.5.a short M present at wing margin.legs.acropods missing, so length of empodia unknown.terminalia (fig.9A-B).Ninth tergite rounded posteriorly.Gonostylus flattened, rounded apically (fig.9a).Gonocoxites (fig.9a): true processes missing, medial bridges with membranous outgrowths that extend into the ventral emargination; emargination small, perfectly U-shaped, with glabrous rim basally; apodemes short, bilobed anteriorly.Ventral parameral tusks smaller and less strongly bent than dorsal tusks; parameral apodemes large, connected ventrally by weakly sclerotized bridge (fig.9B).Base of ejaculatory apodeme flattened; apical extension covered by membranous cap (Fig. 9B).Mamaev, 1966 a bibliography including the synonymy of the genus Monepidosis was given in the revision of Porricondylinae by Jaschhof & Jaschhof (2013).species of Monepidosis can be distinguished from each other using characters of the male gonocoxites, or more specifically, the processes at the ventroposterior gonocoxal margin, which vary in size, shape and position.There are two kinds of processes: the gonocoxal processes, which are outgrowths of the medial bridges, and the central processes, which are outgrowths of the ventral bridge (Jaschhof & Jaschhof 2013;this paper, fig. 10a).apart from genitalic structures, the morphology of male Monepidosis is extremely uniform.females and larvae of most Monepidosis species, including the species treated here, are unknown.
Their apices are typically rounded (fig.10a), rarely more pointed (fig.10B), and membranous rather than sclerotized.The central processes, separated medially by a V-shaped space, are parallel-sided, unicolored, and end in small, strongly sclerotized hooks (↓, Fig. 10A).The bases of both gonocoxal and central processes are occasionally interconnected by a sclerotized rim, which is very pronounced in the specimen depicted in Fig. 10B (↓).

Remarks on intraspecific variability
Most of the specimens studied have rounded gonocoxal processes that arise from small depressions (Fig. 10C), so show a combination of the character states figured in A and B. The specimens depicted in a and B were initially suspected to belong to different species, until it became obvious that they represent the extreme ends of the range of variation described above.I believe that most of this variation is artificial, i.e. dependent on how a specimen is positioned in the balsam drop.

Etymology
The name heterocera is derived from the Greek words heteros, for 'different', and keras, for 'horn', meaning 'with different horns', which refers to the gonocoxal processes that distinguish this species from its closest relatives.4A).

Etymology
The name scepteroides, an adjective, refers to the similarity of this species to M. scepteri.

Distribution and phenology
Japan (shikoku).Known from a single specimen collected in evergreen secondary growth in November.

Remark
Monepidosis spatulata, a species described on the basis of specimens found in Latvia and Lithuania (Spungis 2006), is here reported for the first time from Gotland, Sweden.

Discussion
The example of Monepidosis is ideally suited to demonstrate how little we know of the biodiversity of Porricondylinae in Europe, the continent regarded quite rightly as the best-investigated for this and the other subfamilies of fungivorous cecidomyiidae.Monepidosis is also exemplary for the fact that many Porricondylinae must be regarded as rare in the sense that specimens are seldom found despite great efforts to collect them.This scarcity, which cannot be explained due to the lack of autecological information, makes species inventories fruitful but time-consuming ventures.for example, a nationwide Porricondylinae inventory running in sweden since 2009 (see Jaschhof & Jaschhof 2013) found nine different species of Monepidosis to date, of which five were new to science (a tenth species, also new, is known to me from two specimens that are too poorly preserved for description).some of the morphologically most unusual Monepidosis, such as M. shikokuensis sp.nov., came to be known not from Europe but from Japan, where Porricondylinae have never been surveyed on a systematic basis, meaning that random findings contribute significantly to the overall picture we have of this genus.Antipodosis gen.nov.must be regarded as a similarly random discovery.The 13 specimens I had available to describe Antipodosis gen.nov.and its eight species were accumulated at the time of the 'Lestremiinae of New Zealand' project for curiosity's sake rather than for taxonomy.another 60 specimens from the same 'bycatch' of New Zealand Porricondylinae belong to 41 different species of 9 genera (Jaschhof, unpublished data), whereas only one porricondyline, Yukawaepidosis aliculata (Yukawa, 1964), was named from the New Zealand region in the past (Gagné 1989;Gagné & Jaschhof 2014).finally, the discovery that Antipodosis gen.nov.and Monepidosis are closely related to each other does not necessarily indicate straight affinities between New Zealand and Holarctic Porricondylinae.It remains unknown for the time being whether the Monepidosis group has representatives in southeast asia, New Guinea, New caledonia, or australia, since all these regions are practically unstudied for Porricondylinae.
Monepidosis and Antipodosis gen.nov., are largely merged into a single structure, they are described as a unit.Body length was measured from the head to the genitalia.Arrows (↓) are used to highlight important diagnostic characters in both the taxonomic descriptions and illustrations.The descriptive sections, titled here 'other characters', mention only those characters that are not referred to in the diagnoses and differential diagnoses, respectively.Taxa are treated in alphabetical order.
australis gen.et sp.nov.resembles A. pureora gen.et sp.nov. in having elongate, slender gonostyli; a long, plate-like gonocoxal apodeme; two pairs of unmodified parameral tusks, and a singlepointed, straight ejaculatory apodeme.characters to differentiate between the two species are provided by the gonocoxal emargination, which in A. australis gen.et sp.nov. is bell-shaped (fig.2a) and in A. pureora gen.et sp.nov.U-shaped (Fig.5A), and by the ejaculatory apodeme, whose base is flattened in A. australis gen.et sp.nov.(notvisible in the ventral view depicted in fig.2B) and widened in A. pureora gen.et sp.nov.(fig.
(Jaschhof & Jaschhof 2013ctinatoidesJaschhof,2013resemble each other in having moderately sized, prominent gonocoxal processes and wide, parallel-sided central processes.The two JASCHHOF M., Taxonomy of Monepidosis and Antipodosis gen.nov.speciesdiffer in minute details concerning those processes.ascharacteristics of M. pectinatoides, the gonocoxal processes are sharply pointed and sclerotized apically, and the central processes are bicolored, black laterally, white medially(Jaschhof & Jaschhof 2013: fig.89B).Further, the apical portion of the ejaculatory apodeme is more strongly bent ventrally in M. pectinatoides than in any other Monepidosis, including M. heterocera sp.nov.
Monepidosis scepteroides sp.nov. is similar to M. scepteri spungis, 2006, especially with respect to the central processes.In M. scepteroides sp.nov., as distinct from M. scepteri, the gonocoxites are wider than long instead of the reverse, and the plate-like gonostylar spine is a single piece rather than consisting of two clearly separated portions (Spungis 2006: fig.4B).Like all Monepidosis but M. scepteroides sp.nov., M. scepteri has two pairs of parameral tusks(Spungis 2006: fig.