The Heterotricha group in New Zealand ( Diptera : Sciaroidea )

Die Heterotricha-Gruppe sensu Chandler ist in Neuseeland mit funf Arten vertreten. Neben Anisotricha novaezealandiae (Tonnoir) sind dies Anisotricha similis sp. n. sowie die einer neuen Gattung, Insulatricha gen. n., zugeordneten Arten I. catrinae sp. n., I. chandleri sp. n. und I. hippai sp. n. Neuseeland erweist sich damit als ein rezentes Verbreitungszentrum fur die Heterotricha-Gruppe. Sowohl Anisotricha als auch Insulatricha gehoren zum Kern der moglicherweise nicht monophyletischen Gruppe, stehen aber in keinem Schwestergruppen-Verhaltnis. Die nachste Verwandtschaft von Insulatricha ist mit der Gattung Chiletricha Chandler in der sudlichen Neotropis zu finden, wahrend die Schwestergruppe von Anisotricha zunachst nicht erkennbar ist.StichworterDiptera, Sciaroidea, Heterotricha group, taxonomy, new genus, new species, New Zealand.Nomenklatorische Handlungensimilis Jaschhof, 2004 (Anisotricha), spec. n.catrinae Jaschhof, 2004 (Insulatricha), spec. n.chandleri Jaschhof, 2004 (Insulatricha), spec. n.hippai Jaschhof, 2004 (Insulatricha), spec. n.Insulatricha Jaschhof, 2004 (Sciaridae), gen. n.


Introduction
The Hetmhirha group has recently been subject of a comprehensive review by CHAN-D I E R (2002).According to this review, 17 living species classified in nine genera occur in all blogeogr3pt-u~ regions of the world except the Nearctic, and three species are known from Eocene amber.CHANDLER (2002) excludes from the Heterotn'cha group Pterogvztrus elongata FREESMN, 1951a view followed hereand fuaher proposes the inclusion of .Ilarrgas exiiis KOV-XLEV, 1986a suggestion rejected here.CHANDLER (2002) states that there is notlung to indicate that M. exilis was not alhed to Hetemtricha; on the other hand, indcation that LW. eexrfis, only fragmentary known from a Cretaceous amber fossil, actuall) belongs to this group appears very weak.
The presence of the Hetern~ticha group in New Zealand wns first noticed in the cstcnsive work on the country's fungus gnats by TOKNOIR & ED\Y:\RDS (1927).The only species htherto h o u -n , HeternLricha novaexea/andiae TONNOIR, has recently become type o f the monoq-pic genus Anisohicha CHANDI.F.R, 2002.In the course of a study srill in progress on various groups of the Sciaroidea in Kew Zealand  , 2003), the author became aware of the presence of several more Hetemhicha-like species; these are described and discussed in the following.Initially a further intention was to make a computerized parsimony analysis of the phylogenetic and biogeographic relationships of the world species belonging to the Heterohicha group.However, soon it became apparent that such a venture would require the microscopic re-examination of almost all the species previously described, in particular the terminaha of their males.Apart from that, the author learned of the appearance of new Hetemtncha material from other parts of the world, including the first Australian species (COLLESS, in litt.; CHAKDI.ER, in litt.), an African species close to the genus Rbyrlchoheterohicha FREEMAN, 1960 (HIPP.~, in litt.) and females of the Japanese Sciaropotajaponica CHANDLER, 2002 (KALLWZIT, pers. comm.).The study of such an extensive additional material was clearly beyond the manageable scope of an investigation that primarily aims to introduce and consider the affilrations of the New Zealand representatives.As described in the following, New Zealand's remarkable sciaroid fauna includes a total of Gve Hetmmiha-like species, two belongtng to the genus Anisotricha and three classified in a new genus, In~~~larniha gen.n.This makes New Zealandbesides southern South America and Africa -one of the recent dstribution centres of the Hetemtricha group and raises the number of named living species to a worldwide 21 for the time being.Clearly the previous idea that the Hetemhiha-like flies are poor in species a n d extremely mrel;.fcouritl IS outdated.Instead, the group appears to be quite diverse and regularly distributed in the temperate zones, particularly in those of the southern hemisphere.As already suggested by CHAVDLER (2002), future research will certainly reveal the Heterotricha group, or parts of it, to deserve the status of a distinct family within the Sciaroidea.

Material and methods
Most of the specimens snidied here were obtained from unsorted insect samples previously collected by New Zealand entomologists and deposited in various New Zealand institutions over the past almost 30 years (see the 'Material studied' sections of the species chapters).Additional specimens were collected by extensive Malaise trapping and sweepnemng in [2001][2002] in the course of a survey of various Sciaroidea led by the author.It should be noted that thls study is based on 199 individuals in total, whereas only a handful of individuals of one species, Anisoh'rha novae~eafandiae, was available for study to previous authors.Most of the studied specimens, including all types, are now deposited in the New Zealand Arthropod Collection, Auckland (NZAC).Several individuals of each species, including all types, were mounted in Canada balsam on microscope slides in order to examine morphological details under a compound microscope.The mounting procedure followed that of JASCHHOF (1998) for lestremiine g d midges; however, before mounting the individuals were macerated in cold 10 % KOH.Other individuals are stored in 70 % ethanol.For light microscopic studies and the preparation of drawings the author used an Olympus BX50 microscope in combination with the U-DA drawing unit.Morphological terminology, if not otherwise stated,

Description
Head: Head capsule hlgher than long; its setation moderately long and undfferentiated.
Postfrons slightly two-lobed, non-setose, with simple or slightly two-pointed frontal tuhcrc!~.F:lce large, non-sc::~sc.C1:;pcus i;ml;llCr ( ~s p c c i a l l ~, r i a ~~u u .i .r j LIIAI~ facc, bctosc, fused with face along its upper margin.Antennae longer than body in males and shorter than body in females, upturned, inserted near midheight of head.Scape somewhat conical, a little larger than pedicel, with ventral setae.Pedcel subglobular, with a few setae along distal m q n .Flagellurn with 14 flagellomeres, first flagellomere longest, terminal flagellomere longer than penultimate; each flagellomere cylindrical, practically without neck, several times as long as wide (basically longer in males than in females).In males, flagellum without microtrichia except a few on flagellomere l basally.Each flagellomere with an even cover of setae arising from membranous rings, these setae about 1.5 times as long as the diameter of the flagellomere or shorter; further with a few sensory spines.Flagellomeres l(-3) dorsally with a few setae arising from sockets.In females, cover with microtrichia on flagellomere 1 more extensive, and setae arising from sockets longer, more numerous and sometimes present also on flagellomere 4. With 3 ocelli at vertex, arranged in narrow triangle.Eyes reniform, separated at vertex for more than the width of the ocellar triangle; with h e , long interornmatidal setulae.Mouthparts well developed, with short proboscis.Labrum beak-like, well sclerotized, non-setose or setose, fringed apically.Lingua with dense f ~g e s apically.Maxlllae with stipites separate and cardo absent; lacinia well developed or largely reduced.Ma-dary palpus with 5 palpomeres, with the fist palpomere ('presegment') smooth and non-setose.Palpomeres 2-5 setose; palpomere 2 with 1-2 wart-like sensillae &stall!; palpomeres (2-)3-4 with long hyaline sensory hairs, most numerous on palpomere 3; palpomere 3 duckest and 5 clearly longest.Labium with prementum present as pair of setose lobes; premental apodeme sclerotized and with paired processes of apparently species-specific outline.Labial palpus 2-segmented, labellum 1 smaller than 2; labellum 1 nonsetose; labellum 2 with numerous, partly spine-like setae.a tegmen is present in Chihm'cha, Anisoh'rha, Siiampota and possibly others; and a tergite 9 modified in a simiiar way like that in ~n~ularncha is found in Chiktti'chu, &ynchohetemhirha, and Ki~ahcha.

Etymology.
T h e name is composed of the Latin insula, for island, and a part of the genus name Hetmmiba.Gender is feminine.
Distribution and phenology.ln~uhhkha hippaioccurs on both main islands, but its distribution area appears to be confined to the central parts of the North Island (Ohakune, Mt Egrnont) and the northern parrs of the South Island (Buller area).Adults were collected by Malaise trapping and sweepnetting in prime, mature native forests of both the podocarp/ broadleaf and southern beech type.Middle elevations (400-950 m) are possibly preferred.
Apparently the flight period is short and restricted to the early summer (November to mid-December).Numerous males but no females were collected.Insufatn'ba hiplsai was found to occur in the same sites like I.
Legs: Fore tibia with anteroapical depression (Fig. 9) bearing straight comb of 14-19 spine-like setae subequal in length, with ver!.Indistinct rounded upper rim in some &stance from the comb and short, fine setae between comb 2nd rim.Wing: Venation: tb shorter or longer than mcu, and X1 usually shorter but sometimes longer than CUP.Abdomen: Segment 8 shorter than half the length of segment 7. Terrninalia: Gonocoxites (Fig. 12) ventrally broadly joined under incorporation of sternite 9 (which sometimes stands out From thegonocoxites as triangular portion mecllobasally, see Fig. 13, with V-shaped e m q p a t i o n ventro&stally; with setae of various lengths on its ventral and dorsal surfaces, setation absent from a portion ventroproximally, and very dense around the emargnation; with setose, elongate lobes in section 3 and bare, rounded lobes dorsoproximolaterally; dorsal gonocoxal apodemes unusually wide, with short, pointed processes directed inwards.Gonostyli (Figs 14,15) large, consisdng of two large lobes subequal in size; with &stal lobe setose outside and bare inside; with proximal lobe directed inwards, its apex incised, beanng minute tooth and setose s w e h g ventru-subapically, with many very short seme outside and inside; dorso-subbasally with another small, rounded, setose lobe; basolateral apophysis rather small.Aedeagus (Fig. 16) with long sclerotized apodeme widened apically into weakly sderotized head, the latter with complicated structure that apparently is closely joined with the tegmen.Tegrnen (Fig. l 6) broad, with dear contours, distally with rounded lobe; ventral p m e d apodemes short and sderotized; dorsal pararneral apodemes short, well sclerotized and connected by sderotized transverse bridge.Tergfite 9 (Fig. 17) roughly U-shaped, with long seue on its outer surface and many short, thorn-like, two-pointed megasetae inside distally.
Legs: Fore tibia with anteroapical depression bearing comb of 9-1 1 spine-like sctae, and with the h e setae between comb and rim fewer in number than in male, or cvmpletely absent.Terrninalia (Fig. 11): Distal cercus longer than wide.With 2 large, ovoid, wcll sclerotized spermathecae bearing many pores on their surfaces.Discussion.Male terminalia in In~uhfricha chandleridiffer markedly from that in the congeneric species, I. h$pai and catrinae (see next species).S m h n g differences tie in the structure of the gonocoxites (see Fig.  Legs: Fore tibia with anteroapical depression bearing straight comb of 13-16 spine-like setae subequal in length, with very indistinct rounded upper rim in some distance from the comb and short, fine setae between comb and rim.Wing: Venation: tb longer than rncu, and .\las long as or longer than CUP.Terrninalia: Gonocoxites ventroproximally broadly joined and centrally with sternite 9 in between (distorted in slide preparations available for study); with setae of various lengths on its ventral and dorsal surfaces; with setose rounded lobes in section 3. Gonostyli turned inwards, broad proximally and wedge-shaped in dstal half, with apex apparently simple; with long setae on outer surfaces and very short, fine setae inside and terminally; basolateral apophysis rather small.Aedeagus (Fig. 18) with long sclerotized apodeme; no further details visible, but its structure apparently rather simple.Tegrnen (Fig. 18) long, with clear contours, &stall!-u-ith narrow head; dorsal parameral apodemes moderately long, well sclerouzed and connected by weak transverse bridge.Tergte 9 (Fig. 19) plate-like, with long setae on its outer surface and narrow processes disto-laterally bearing short, thornlike, nvo-po~nted megasetae.Cerci pointed apically, densely setose.Hypoproct veryweak, two-lobed, apparently bearing 1 seta each dstally.
Female.Body size: 2.7 mm. Head: Antenna u-ith 4' flagellomere almost 4 times as long as wide.Terrninalia: Distal cercus almost as long as wide.With 2 small, ovoid, well sclerotized spermathecae bearing pores on their surfaces.
Discussion.There are only two male individuals of thls species available for study which due to the long storage in ethanol could not be cleared sufficiently; that is why several d e d s of the terminaha cannot be described in as much detail as would be desirable.
Insuhtncha calrinae differs from its two congeners (in parentheses) in two points: the labrum is setose (non-setose) and the lacinia are well developed (largely reduced).Further dfferences are in the structure of the male terminalia (see under I. chandLen).
Distribution a n d phenology.rn~uLat~%-ha catrinae is known from a single locality in the central North Island (Puceora Forest), where chandLen'was foand as well.For details of the collection site, see under Types.
Etymology.I name this species after my wife, CATRIN JASCHHOF, Greifswald, in appreciation of her invaluable assistance in my various research projects on sciaroid fies.

Genus Anisotricha CHANDLER, 2002
T h s genus was recently proposed by CHANDLER ( 2002) for a single species, Heterohicha novaqcahndioe TONNOIR in TONNOIR & EDWARDS, 1927.Due to the fact that previous authors had only very few specimens of this species available for study, many details of the male terrninalia remained unknown.Females were totally missing.Based on the availabdity of much more material of both sexes, I am now in a position to complete the species and genus descriptions.Moreover, microscopic study of the male terminalia of Anisotn'cha led to the recognition of another speues very similar to novaer(eaLandiae.Thls new species, Anisohirha n'milis, is described below. Supplement to the description Head: Postfrons slightly two-lobed, non-setose, with slightly two-pointed frontal tubercle.Face large, non-setose.Clypeus smaller than and dearly separated from face, setose.Scape somewhat conical, a little larger than pedicel, with ventral setae.Pedicel subglobular, with a few setae along d i s d m a r p .In males, flagellum without microtrichia except a few on flagellomere 1 basally.Each flagellomere with an even cover of setae arising from membranous rings, these setae as long as diameter of flagellomere or shorter; further with a few sensory spines.Ragellomeres l(-3) dorsally with a few setae arising from sockets.In females, cover with microtrichia on flagellomere 1 more extensive, and setae arising from sockets more numerous and sometimes present also on flagellomere 4. Eyes with interommatidal setulae normally developed or much reduced.Labrum beak-like, well sclerotized, non-setose, fringed apically.Lngua with dense fringes apically.Masillae sdpites separate and cardo absent; lacinia well developed, fringed apically.hIasillary palpus with 5 palpomeres, with the first palpomere ('presepent?well developed and setose.Palpomere 2 with a few wan-like sensillae dstally; palpomere 3 not thickened, with short, hyaline sensory hairs accumulated in basal half; palpomere 5 estremely long, often longer than preceding two palpomeres together.Labium with prementum present as pair of setose lobes (often with only l seta each); ptemental apodeme sderotized and with paired processes.Labial palpus 2-segmented, labellum 1 smaller than 2; labellum 1 non-setose; labellum 2 with numerous, partly spine-like setae.Thorax: Postpronotum present as collar-like structure above neck.hntepronotum submangular, with a few setae.Episternum 1 with a few setae.Epimeron l small, subtrian- Legs: Coxae longer than half the height of thorax.Coxae and femora with setae of moderate length.Tibial spurs 1:2:2, well developed.Fore tibial anteroapical depression well developed, with distinct, U-shaped proximal rim; depression bearing many setae, including those forming a comb distally.Tibiae and tarsi covered with large trichia and short setae; mid and h d tibiae and tarsi of all legs with a few short, strong projecting setae.Tarsomeres 1-5 gradually decreasing in length.Pretarsal daws small, somewhat curved, without teeth.Pulvdh well developed, as long as empodia.Empodia half as long as claws.Wing Long, i.e. more than two times as long as wide, but shorter than body.Calyptral area somewhat convex; anal area moderately developed.Membrane covered with setae with setation less dense towards wing base and anterior margin, i.e. costal cell lachng setae and basal cell with strongly reduced setation.Venation: C extending to apex of wing, en&ng at a point half way between apices of R5 and M1; Sc long, ending abruptly beyond level of Rs; Rs oblique, slightly curved, much longer than ta; ta oblique; R5 with short faint portion proximally; h1 absent; M1 +2 stem shorter than fork, the latter clearly dvergng; tb+mcu almost parallel to anterior wing margin, tb longer than mcu; CUP reaching to or going beyond half length of CuA2; A1 shorter than CUP; A2 barely traceable.With setae present along lving m a r p and on Sc, R, RI (dorsally and ventrally), sometimes Rs, R5 (dorsally and L-entrall!.), 111 +2 stem and fork (dorsally and ventrally), ta, tb (dorsally and ventrally), sometimes mcu, C d l (dorsally and venually), CuA2, and A l .Pattern of sensory pores: R, 0-1; R1 ,3-4; Rs, 1-2; R5,l-2 proximally, 1-3 distally.Abdomen: Tergites evenly covered with long setae.Tergite 1 and sternite 1 each subdi-1-ided into long anterior and short posterior plates with a membranous portion in benl-een.Tergite 1 shorter than 2, and 7 longer than 8. Sternite 1 setose.Sternite 2 with nso membranous windows anteriorly separated by sclerotized bar bearing numerous senson setulae.Alembranous portions on scleritcs 1 and 2 obviously creating a flexible zone enabling the abdomen to be bent behind sebment 1. Sternites 2-8 with long setae arranged in three irregular longitudinal rows.With six pairs of spiracles, one each on se,pents 2-7.Tergal plaques small, situated in a anterolateral position on each tergite, with their pattern 0/1/1/1/1/1/1/0.TerrninaIia.Male.Sternite 9 present as dlstinct sclerite.Gonocoxites fused ventrobasally and -clistally and having sternite 9 incorporated in between, with setose lobe in gonocoxal section 3; gonocoxal apodemes not forming a transverse bridge but interlmked through the dorsal wall of tegrnen.Gonostyli with sweeping lobes.Aedeagus consisting of a basiphallus with long sclerotized apodeme, and a sclerotized, two-part 'dlstiphallus'.Parameres intimately fused to form a tegrnen, with sclerotized 'teeth' distolaterally; with large, smooth lobe each dorso-proximolaterdy bearing many long, strong tricha, and arising from this lobe, with long, smooth, ribbon-like appendage bearing long tricha most s t r i h g terminally; tegmen dorsomedidy linked with lateral appendages of gonocoxal apodemes; dorsal 'parameral apodemes' (actually, clearly prolongations of the gonocoxal apodemes!)very long and strongly sclerotized.Tergte 9 plate-like, subtrapezoid, with slight, setose s w e h g dlstomedially.Tergite 10 absent or (if correctly identified) present as smooth, setose lobe.Cerci large, subtriangular, with fine, long setae dlstally.Hypoproct weak, paired or simple, non-setose or setose.Female.Slightly deviating from basic pattern in Sciaroidea (see under A. novaexeafandiae).With 2 sclerotized spermathecae.
Discussion.CH~NDT,ER'S (2002) definition of the genus Anisotrirhn was necessarily based on A. novaexeafandiae and, after study of the second species, A. sz7z~ifis (see below), can herewith be confirmed.After the addition of characters of the female, the definidon reads as follows: the antennae are upturned and with the flagellum long (plesiornorphous conditions); the clypeus is unproduced (plesiomorphous); episternum 3 is setose (apomorphous); the wing veins, including Sc, bear setae (plesiomorphous), while the setation on the membrane is somewhat reduced (apomorphous); Rs is oblique and longer than ta, with the latter also oblique (plesiomorphous); the stem of M1 +2 is rather short, i.e. shorter than the fork @lesiomorphous); the parameres are fused to forma tegmen (apomorphous); themale tergite 9 is mainly unmodified @lesiomorphous); the ovipositor is shortened (apornorphous); and two sclerotized spermathecae are present (plesiornorphous).As usual with genera of the Heterotricha group, this definition is polythetic, and autapomorphous characters are not recognizable of Anisotricha, except possibly those of the terminalia (see below).Episternum 3 is setose also in Rhynchohetemtricha, Afiotricha and Chifetniha; the setation on the wing membrane is reduced also in Rlynchoheterotn'cha and Chifetricha; and a tegmen is developed in Insufatticha, Chifetticha, Sriampota and possibly others.In Anisotricha, male terminalia are very peculiar by the presence of a well developed 'distiphallus' and trichose appendages o f the tegmen.The latter might indicate a secondary subdivision of the parameres into ventral and dorsal portions -with the appendages representing the dorsal portiona condition that is apparently absent in other members of the Heterotricha group and, if explained by secondary acquisition, an a ~t a ~o m o r p h o u s character of Anisotricha.T h e homology of the 'distiphallus' in Anisotncha is difficult to assess, but a corresponding structure seems to be present in Chifetrichapenae CHANDLER, 2002(CHANDLER, 2002: 122, Fig. 39).The ovipositor in Anisotrirha gaines its shortness mainly through the very short distal cerci, whch is unique within the Heterotrirba group as so far known, but otherwise : difficult to compare as the females in several genera are not yet known.
Distribution and phenology.Ani~ohkha nocuqeala?rriiaeappears to be confined to the northem South Island (Nelson and B d e r areas).The specimens studed here come from the -tensive southern beech forests, that are sometimes mixed with podocarp trees, of the Nelson Lakes National Park, from elevations between 450 and 700 m.Tlus area is in close distance to the species' type locahty, ,hiseed Valley near Nelson, where most of the origi- Specimen from Lake Rotoroa Length of scale bar: 0.2 mm.nal forest had to make way for human settlements.The specimens studied were Malaise trapped in mid summer @ecember/January).Males were collected 20 times more commonly than females.1n.rulatncba bippai was shown to occur at the same sites as A~~isomcha novae~ealhndiae.
Etymology.The specific name is Latin meaning 'similar', referring to the similarity with R noc.ae~olan&.

Relationships of Aaisotricha a n d Insulatn'cha within the Heterotn-cha group
The delimitation of the Hetemhcba group from other Sciaroidea is based on the corresponding wing venation of the species included; however, the so-defined group is not necessarily monophyletic (CHANDLER, 2002).When other characters are taken into consideration, a core-group consisting o f Chiletncha, Rhynchoheterotncha, Anisotricha, Heteratticha and .-!fiotn'cha is ascertained, while Nepaletricha and Kenyatricba, and even more so Sciaropota and Siarosoma appear to be as distantly related to the core as do other Sciaroidea, like Sciaridae, Ohaklinea and Colonomyia (CHANDLER, 2002).The close relationship benveen Heterotricha, Afrotricha and Chiletricha was inferred from similarities in the female terrninalia, while the structure of the male terminalia led both to the assumption that R!!nchoheterotnha and Anisotricha would belong to the same clade and to the anticipation of similar female structures (CHANDLER, 2002).Sull nothing can be said on the female structures in Rhynchoheterotricha, but in the two New Zealand genera ,~nzsotncha and Znsrilatricha the female terminalia differ from each other and from the genera referred to above -and thus do not support the assumption of closer relationshp.The distribution of other characters in Insalatricha does also not support the argumentation advanced by CHANDLER (2002: 142, cladogram Fig. 103).In detail, in Insulatrii-ha the episternum 3 is bare (character 17 in CHANDLER'S cladogram), whch contradicts its assignment to a clade (Chiletricha +fiynchoheterotn'cha)+Anisotricha; male tergite 9 bears spinose processes (or, as here called, short megasetae distolaterally) (character 14), w h c h indicates its belongng to a clade ChiletrichafIUIynchoheterotri'cha; and vein Sc is setose (character 15), which contradrcts its assignment to such a clade.As so often in phylogenetic considerations on Sciaroidea, the addition of new taxa causes conflict and more confusion rather than leading to support and better resolution.As CHANDLER (2002) states, "... strongly apomorphous characters are either autapomorphous in particular subordinate taxa ... or have developed independently in more than one family ...", and this seems to apply to the Hetemhicha group as well.
T h e is no doubt that both InsilLmcha and Anisohcha arc dstinct, monophylctic groups.One should e-xpect that these two New Zealand genera are more closely related to one another than to genera outside New Zealand, but the opposite seems to be true.O n the basis of the male terrninaliain particular the structure of tergtte 9 -Ins~iia~cha appears to be most closely related to Cbihm'cha, even though several detds of the genitalic structures in the latter genus are not yet described (for instance, the outline of sternite 9 and the h e structure of the 'spines', or megasetae, on tergite 9).In contrast, Anisotri'cha with its strongly derived male genitalic structures (see, in particular, gonostyli, phallus and pararneres) holds a rather isolated position among the genera of the core group where it undoubtedly belonp to.
Position of the Heterotricha group within the Sciaroidea CHANDLER (2002) discusses in length the development of ideas on the phylogenetic relationships between family-level taxa of the Sciaroidea and makes it clear that most of the argumentation advanced by previous authors is baseddue to the scarcity of other useful characterso n larval morphology and wing venation.As regards the Heterotricha group, larvae have not yet been found, a fact which leaves almost all phylogenetic consideration to wingvenation.However, wing venation in the Sciaroidea provides a character set that is notorious for the extent of homoplasy involved.The previously practised assignment of Neterofniha to the family Diadocidiidae resulted mainly from such formal interpretations of vein patterns.To put it in drastic terms, one and the same vein pattern may be explained in different ways depending o n what interpretation best fits into a certain argumentation scheme.This is particularly true for pre-Tertiary fossils in whch the more or less com-pletely preserved wing venation often is the more easily, or even only exploitable source of information.Such fossils may be helpful in reconstructing general tendencies, or main stages in the evolution of venation (see WTILE,198l), but they may be object of speculation rather than interpretation in a particular taxonomic case.It is for such reasons that the author cannot accept the inclusion of Nangas exdis KOVALEV into the Heferomcha group (CHANDLER, 2002).Admittedly, after the study of the New Zealand representatives of the Hetemm'cha group we are not provided with better arguments in order to decide whether or not the group is monophyletic and where to place it within the Sciaroidea.At present, it appears that the group represents one or two extant lineage(s) of ancient sciaroids that have their origin in the broadly developed lurassic stock of fungus p a t s in the broadest sense.This .p u p obviously r r m e d the -&pe o t venation that is not found in other extant Sciaroidea but predominated in an earlier, Jurassic radiation of the Sciaroidea.What we learned in New Zealand is that the search for Heierotn'cha-like flies -and certainly also their larvaeis more calculable than cupected only a few years ago.These flies seem to be not so exceedingly rare if one only knows where and when to look for them.Currently we can do little more than to continue seardung for new study materialadults and larvaein order to get a more realistic idea of the variety and distribution of the characters and character states we work with.

Key to the New Zealand species of the Hererorricha group (males)
Remark: There might be species-specific cl~aracters even in females, as evidenced by the species-pair Znsufatrfcha chandfen' and cahinae, but the females of two of the five species in question are still unknown and thus not available for a comparative study.