A new genus and species of Chironominae (Diptera: Chironomidae) with wood-mining larvae

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1 Blackwell Publishing AsiaMelbourne, AustraliaAENAustralian Journal of Entomology ;? Original ArticleNew genus of wood-mining Chironomidae P S Cranston Australian Journal of Entomology (2006) 45, A new genus and species of Chironominae (Diptera: Chironomidae) with wood-mining larvae Peter S Cranston* Entomology Department, University of California, Davis, CA 95616, USA. Abstract Key words Xylochironomus kakadu, a new genus and species of wood-mining chironomid, is described from northern Australia. Formerly known by a code unknown genus K1, the larvae mine soft immersed timber in tropical Australia. The basal striae on the larval mandible are observed otherwise only in Chironomus Meigen and likely close relatives currently placed in Einfeldia Kieffer. The adult male could key as a tanytarsine, and all stages superficially resemble Polypedilum Kieffer, except for the bare squama and the non-tapered male tergite VIII, and the dorsal larval head sclerites. However, analyses of a morphological data matrix derived from all life history stages support no such relationships. Instead, a relationship to Paralauterborniella Lenz and Apedilum Townes, and at one remove, to Fissimentum Cranston & Nolte and Imparipecten Kieffer is postulated. Larval wood-mining among the early branching Chironomini is evidently frequent, but taxa appear not to form a monophyletic clade. Australia, Chironomidae, evolution, wood mining, xylophagy. INTRODUCTION Mining in immersed wood by larval Chironomidae is well documented (e.g. Anderson 1982, 1989; Hoffmann & Hering 2000; Cranston & McKie 2006) and occurs across a wide range of taxa in the subfamilies Orthocladiinae (e.g. Cranston 1982, 2000; Kaufman & King 1987; Cranston & Oliver 1988) and Chironominae (e.g. Borkent 1984; Cranston & Hardwick 1996). Among such wood miners is a formally unnamed Australian taxon referred to as unknown genus K1 (Cranston 1991, 1996). In the corresponding informal vouchering system, K refers to Kakadu (National Park, Northern Territory, Australia), where larvae of this taxon mine in immersed wood in creeks and springs. The associated reared adult males exhibit a combination of features that intergrades between the tribes Tanytarsini and Chironomini, and for this reason K1 was included in phylogenetic analyses of Nandeva Wiedenbrug, Reiss and Fittkau (Cranston 1999). The addition of this taxon, as well as of previously unknown life history stages of wood-mining Asian Shangomyia Wang and Sæther (Cranston 2003), raised some interesting questions concerning larval xylophagy and the evolution of Chironomini (Cranston & McKie 2006). In the postulated phylogeny (Cranston 1999), both K1 and Nandeva associated with the wood-mining Australian Imparipecten Freeman, at a short remove from the well-known clade of wood-mining *pscranston@ucdavis.edu or leaf-chewing Stenochironomus Kieffer, Harrisius Freeman and Shangomyia, despite the lack of any obvious morphological similarities between the two clades. Here, life history stages of K1 are described, formal names are provided for a new genus and species and a morphologically derived phylogeny is discussed. MATERIALS AND METHODS Larvae were sought by prising apart immersed wood of soft decay classes of unknown species in tropical rainforest patches. Unusually for xylophagous chironomids, some individual rearing was successful. Adults were collected in two locations by sweeping riparian vegetation and at light beside the water. Extensive drift sampling did not yield pupal exuviae. Microscope slide preparation involved clearing where necessary with 10% KOH, neutralisation and initiation of dehydration with glacial acetic acid, then mounting from propan-2-ol (isopropanol) into Euparal. Pupal exuviae recovered post emergence were crumpled and are interpreted only with difficulty thus, Figure 1f is somewhat stylised. Several whole larvae were slide mounted in Hoyer s and ringed with nail varnish. Recognising that taxonomic drawings can be difficult to interpret for non-specialists (Cranston 2005), photographs using the Automontage system supplement more traditional line drawings. Morphological measurements, unless otherwise stated, are in µm. Standard terminology (Cranston 1994) is used, with the following abbreviations: Fe Femur; Le Larval exuviae; Le/Pe/m(f) Reared adult male (female) with associated larval doi: /j x

2 228 P S Cranston and pupal exuviae; LR Leg Ratio: Tarsomere 1 length: Tibia length; Pe Pupal exuviae; Ta 1 5 Tarsomeres 1 5; Ti Tibia; VR. Venarum Ratio: length of wing vein Cu : length of wing vein M. Xylochironomus gen. n. Genus K1. Cranston (1991, 1996, 1999). Type species. Xylochironomus kakadu sp. n., by present designation and monotypy. Generic diagnosis Larval Xylochironomus have a distinctive 5- or 6-segmented antenna with large Lauterborn organs alternate either on an antennal segment interpreted as the 2nd, or if the latter is considered as medially divided, on the apices of the 2nd and 3rd segments. Xylochironomus resembles Polypedilum nubifer (Skuse) in this feature, although the antenna of the latter is interpreted as having 4 or 5 segments, with Lauterborn organs on a divided/undivided 2nd, and with only 2 more apical segments. The mentum of Xylochironomus, with recessed paired median teeth, and the semilunar ventromental plates are distinctive and characteristic. Resemblance to larvae of Polypedilum Kieffer is countered by the presence of a discrete clypeus (dorsal head sclerite 1) with cephalic seta S3 lying lateral to the sclerite, and the presence of this discrete sclerite 1 (in contrast to Polypedilum bearing S3 on the antero-lateral corner of a frontoclypeal apotome and lacking sclerite 1). The striate mandibular base occurs otherwise only in Chironomus Meigen and immediate relatives. In pupal Xylochironomus the few-branched thoracic horn, distinct anterior and posterior spinule bands on tergites III VI, and 3,3,4,4 pattern of taeniae on segments V VIII resembles the heterogenous Polypedilum, but the medially divided tergal II hook row is unknown in Polypedilum. In the adult the microtrichiose ( hairy ) wing combined with a bare squama occurs elsewhere only in the tribe Tanytarsini, but genitalic structure and the oblique angle between RM and R 4+5 indicate that Xylochironomus does not belong to this tribe. The non-tapered abdominal segment VIII precludes allocation to Polypedilum. Generic description Larva (4th instar) medium-sized, up to 6 mm long, with ventral head length up to 520 µm. Pink-red coloured; head capsule golden yellow with medium-brown gula, mentum and inner mandibular teeth. Occipital margin slightly darkened. Dorsal surface of head (Figs 1c,2e). Frontal apotome tapered posteriorly, broad anteriorly, without frontal pit; setae S3 outside lateral margin of separate clypeus; labral sclerites 1 and 2 complete, 3 5 less distinct, 4 comprised of dissociated granules. Antenna (Figs 1b,2f) 6-segmented if faint division of subbasal segment exists just distal to basal Lauterborn organ (Fig. 2f); 2nd more distal Lauterborn organ also large, located at apex of 3rd segment rotated at 90 to basal organ (semialternate); if antenna is treated as 5-segmented, then 2nd segment with Lauterborn organs near midpoint and apical. Combined 2nd and 3rd antennal segments slightly shorter than penultimate, apical short. Style almost hyaline, subequal in length to Lauterborn organ, originating on antenna opposite to origin of basal Lauterborn organ (i.e. at apex of presumptive 2nd antennal segment, if considered 6-segmented). Ring organ poorly delimited; seta absent. Blade extending distinctly beyond subapical segment. Labrum (Fig. 2g,h). SI apically and subapically plumose, SII apically plumose, longer than SI; SIII simple, short, located between bases of SII; SIVa and b normally developed. 1 strong, plumose chaeta. Seta praemandibularis short, simple. Labral lamellae narrow, bilobed but without indication of median division. Pecten epipharyngis comprising 3 separated scales, median with 2 3, lateral with 4 6 blunt teeth. Chaetulae laterales 6 7, weakly serrate, 2 simple chaetulae basales displaced more anteriorly. Premandible (Fig. 2g,i) with 2 narrow apical teeth with dense, dorsomesally orientated, very strong setal brush(es) arising from tubercular common stem, directed oro-ventrally (arrowed in Fig. 2i). Mandible (Figs 1d,2c). Lacking dorsal tooth; with short apical tooth, 3 distinct inner teeth and an innermost 4th weakly delimited from the mola; with basal striae (arrowed in Fig. 2c) as in Chironomus. Pecten mandibularis reduced to 3 fine setae. Seta subdentalis inserted on ventral surface, simple, extending to tip of 3rd inner tooth Molar margin with 2 3 fine serrations. Seta interna very well developed, with 4 major branches. Mentum (Figs 1a,2a). Anterior edge shallowly convex, without distinction between ventromentum and dorsomentum at anterior margin of mentum; with 8 evenly brown coloured teeth on each side, the innermost pair sharply depressed relative to large 2nd. Ventromental plates (Fig. 2a,b) separated medially by width of 4 median teeth, somewhat semilunar-fanshaped, medial corner directed posteriorly, anterior margin smooth; striae relatively uniform-length ridges across much of plate. Setae submenti simple. Maxilla (Fig. 2d) with notable lanceolate, medial-directed lacinial chaeta that meets its opposite chaeta antero-dorsal to mid-mentum. Abdomen. Lateral and ventral tubules absent. Anterior parapods with dense, fine, simple claws; posterior parapod claws sparse, golden, simple. Procercus weakly pigmented, small, slightly higher than wide, bearing 6 7 subequal anal setae. Pupa medium-sized, approximately 4 mm long. Thorax, cephalic area and anterior abdominal segments pale, all apophyses weak, golden brown. Cephalothorax. Frontal seta taeniate, arising from membrane, cephalic tubercle absent; apotome smooth without frontal warts. Pedicel sheath smooth. Thoracic horn with 4 6 fine branches, one appearing spinose; basal ring (Fig. 1e) small, oval, with 1 round tracheal bundle. Median suture spinose medially only, without tubercle. Prealar tubercle absent. One dorsal taeniate, 1 lateral hair-like antepronotal seta; 2 narrowly taeniate precorneals; dorsocentral Dc 1 very close to Dc 2, separated from closely approximated Dc 3 and Dc 4, all subequal and narrowly taeniate.

3 New genus of wood-mining Chironomidae 229 (a) (c) (b) Fig. 1. Xylochironomus kakadu Cranston sp. n. Larva: (a) mentum and ventromental plates; (b) antenna; (c) frontal apotome and labral sclerites; (d) mandible. Pupa: (e) basal ring of thoracic horn; (f) abdomen, dorsal view; (g) comb of abdominal segment VIII. (e) (d) (g) (f) Abdomen (Fig. 1f,g). Tergites I, VIII and anal tergite bare, VII with or without anterior and central shagreen; II VI with strong anterior transverse band of strong spines some 4 6 spinules deep; central area sparsely spinulose (sometimes denser on V and VI) to showing few weak spinules (III IV); posterior 1/3 of III VI with band of small spinules broader in posteriormost transverse band, narrower anteriorly and partly divided into 2 areas, VII with or without slight anterior and central shagreen Tergite II hook row discontinuous, c. 65% tergite width, comprising pale brown hooks on each side. Conjunctives bare except for posterior tergal armament running onto anterior conjunctive on III VI. Sternites bare. Pedes spurii A strong on IV, very weak on V and VI; pedes spurii B well developed on II. Postero-lateral corner of segment VIII with 1 3 large, dark, major teeth amid few to many smaller teeth (Figs 1g,3a). Setation. Segment I, D and V setae not visible, without L setae; II IV with 4D, 2 3V, uncertain number of weak L setae with only L2 developed; V and VI with uncertain D and V setal numbers, 3 taeniate L setae; VII with 4 or 5D, 3V, 4 taeniate L setae, VIII with 1D, 2V, 4 taeniate L setae. O setae apparently absent. Anal lobe elongate oval, with fringe of uniserially inserted, taeniate setae, extending to apex and sometimes around to inner subapical margin. Dorsal surface lacking taeniate seta. Genital sac reaching no farther than apex of anal lobe in either sex. Adult Antenna. Male with 13 flagellomeres, antennal ratio female with 5 flagellomeres, AR c Head. Eye bare, with long parallel-sided extension in near median contact. Frontal tubercle very small; 6 8 uniserial temporal setae Palp developed normally, palpomeres (Pm) 3 and 4 (visible 2nd and 3rd) subequal, about half length of apical Pm 5. Thorax. Antepronotum widely separated medially; scutum not over-reaching antepronotum, tubercle absent. Thoracic setae long; acrostichals biserial, strong, extending to anterior 1/3 of scutum from the antepronotum, dorsocentrals uniserial,

4 230 P S Cranston a b c d e f g h i Fig. 2. Xylochironomus kakadu Cranston sp. n. Larva: (a) mentum; (b) ventromental plate; (c) mandible (striae arrowed); (d) maxilla (lacinial chaeta arrowed); (e) dorsomedian head sclerites; (f) antenna; (g) labrum; (h) epipharynx; (i) premandible (pmd) (brush base arrowed). humerals and antepronotals absent, prealars 4, scutellars unserial. Wing (Figs 3b,4a). Membrane densely setose, slightly more so distally. Costa not extended, R 2+3 indistinct, perhaps running very close to R 1 ; R 4+5 ending slightly proximal to wing apex, distal to end of M 3+4 ; RM angled obliquely to R 4+5. A false vein anterior to Cu indicated by aligned, more densely placed setae. FCu distal to RM. Anal lobe weakly developed. Squama bare. Legs. Apex of fore tibia with well developed rounded scale, without spur (Fig. 3c). Combs of mid and hind legs (Fig. 3d) occupying 2/3 circumference, slightly separated, each leg with long spur on inner comb, outer without spur. Tarsomeres lacking sensilla chaetica. Pulvilli about half length of claw, without major branching. Abdomen with scattered long setae, segment VIII not contracted anteriorly. Male hypopygium (Figs 3e,f,4b,c). Anal tergite trapezoidal with moderately pigmented V-shaped tergal bands, separated medially or meeting faintly. Few long median anal tergite setae, posterior margin of anal tergite with row of 8 12 near continuous or medially separated apical setae Anal point (Fig. 3e) well developed, arising on dorsal surface of, and set off from tergite IX, hyaline, narrow, parallel-sided until slightly dilated subapically. Superior volsella (Fig. 4c) with distal part digitiform, sinuous, bare except for 2 strong subbasal setae, one medio-dorsal, the other medio-lateral; volsellar basal lobe swollen, with 2 strong setae both directed medio-ventrally. Median volsella absent. Inferior volsella cylindrical, slightly clubbed apically, extending to

5 New genus of wood-mining Chironomidae 231 b a c g Fig. 3. Xylochironomus kakadu Cranston sp. n. Pupa: (a) comb of abdominal segment VIII. Adult male: (b) wing; (c) fore tibial apex; (d) hind tibial apex; (e) anal point of hypopygium; (f) hypopygial apodemes (lateral orientation). Female genitalia: (g) dorsal; (h) ventral. d f e h mid-gonostylus; microtrichiose only ventrally, with dorsal setae in about 3 rows of setae on stem, denser on clubbed apex, without special elongate posteriorly directed apical seta. Gonostylus well delimited from gonocoxite, somewhat broader at base and slightly narrowed medially, tapered apically with apical medially directed small tooth bearing a seta that is more stout than other setae on subapex and median margin of gonostylus. Sternapodeme an inverted U -shape (Fig. 4b). Female genitalia (Figs 3g,h,4d,e). Notum conventional with short broad rami almost as long as the fused section. Gonocoxapodeme curved, the pair not meeting medially. Coxosternapodeme IX sinuous, well sclerotised. Gonapophysis VIII with dorsomesal lobe curved around the posteromedian corner of the coxosternapodeme, ventro-lateral lobe small, short, microtrichiose basally but without group shagreen, with longer scales medio-apically; apodeme lobe associated with darkly sclerotised apodeme. Labia hyaline, without microtrichia. Gonocoxite IX small, not laterally extended, with sparse setae. Tergite IX large, broad, undivided, setose. Postgenital plate broad, microtrichiose. Seminal capsule pale, oval, tapering to narrow neck; spermathecal duct nearly straight and ending approximated at single orifice but without fusion. Cercus well developed, quadrate in lateral aspect. Xylochironomus kakadu sp. nov. Material Holotype: Le/Pm, Australia, Northern Territory, S E, Kakadu N.P., Baroalba Springs, 1.vi.1988, Cranston. Paratypes 2Le/Pe/f, Le/Pf, 5L, m, as holotype; 3L, Le/ Pm, 15 m, S E, Radon Springs, 6.vi.1988, Cranston; L, 2 m, S E, S. Alligator R., Coronation Hill, 4 5.vi.1988, Cranston; L, Mudginberri B bong, S E, 12.vi.1989, Cranston; Queensland, Le/ Pe/f, S E, nr. Rex Creek, [Mossman N.P.], x.1998, Cranston and Dimitriadis. All type material is mounted on slides in Euparal and deposited in the Australian National Insect Collection, CSIRO Entomology, Canberra, Australia (ANIC), except for 2 male paratypes each in the Natural History Museum (BMNH), London, UK and the Zoologische Staatssammlung München (ZSM), Munich, Germany. Non-type material: several larvae from Kakadu, slide mounted in Hoyer s, including a molecular voucher. Description (largely mensural additions to genus description) Larva (n = 8) Length mm (n = 4), ventral head length µm. Antennal segment lengths ; (if fused); 25 32;

6 232 P S Cranston (a) (c) (b) (d) (e) Fig. 4. Xylochironomus kakadu Cranston sp. n. Adult: (a) male wing, setation omitted; (b) male hypopygium (left dorsal, right semi-internal); (c) superior volsella. Female genitalia: (d) dorsal; (e) semi-internal ; 8 12; blade , Lauterborn organ 20 32, style Mentum width , maximum width of ventromental plate Mandible length Respective longest anal seta Pupa (n = 3) Length mm, cephalothorax µm. Longest spine on comb 40 45, triangular. Anal lobe with uniserial taeniae. Adult male (n = 7). Pale brown, without distinct vittae. Body length mm, wing length mm. Antenna with apical flagellomere long, Fm 1 12 combined , AR Head setation: 5 9 uniserial temporal setae, clypeals. Palpomere 2 5 lengths: 35 45, , , Thoracic setation: acrostichals 12 18, biserial, in anterior 1/3 of scutum; dorsocentrals 9 16, uniserial; prealars 3 5; scutellars Wing VR ; vein setation not distinguishable from background membrane setation; squama bare. Legs with longest setae no longer than 3 times corresponding tarsomere width. LR , LR , LR Hypopygium. Gonocoxite length , gonostylus length , anal point length Adult female. Pharate/teneral condition prevents further description. Etymology After Kakadu National Park, a World Heritage listed area, where the first specimens were collected kakadu, to be treated as a noun in apposition. Ecology Larval X. kakadu were found first in immersed and rather soft wood of undetermined identity lying in a deep rock pool in

7 New genus of wood-mining Chironomidae 233 the main channel of the quite strongly flowing stream. Field notes imply larvae were below bark but not obviously mining in heart wood, which remained firm. The Mossman Gorge stream in Far North Queensland was very similar in being deeply shaded by riparian tropical/monsoonal vegetation. Substantial immersed wood was trapped in pools between riffles. Wood texture was soft; harder wood was unsampled. No other wood miners have been found to cooccur; associated chironomids in these habitats include Dicrotendipes lindae Epler, D. leei Freeman, Conochironomus australiensis Cranston, Ablabesmyia notabilis Skuse and undescribed species of Polypedilum. Adult males of X. kakadu have been found at additional sites in Northern Territory where immersed wood was abundant but larvae were not found. A subsequent collection of larval Chironomidae from immersed wood in the Alligator Rivers region, Northern Territory, by Chris Humphrey (Environmental Research Institute of the Supervising Scientist (ERISS)) revealed larval Xylochironomus mining in softer wood. The absence of pupal exuviae from drift samples taken in the region and elsewhere is surprising given the usual representation of various other members of the community in corresponding samples of larvae. Phylogeny of Xylochironomus Xylochironomus was included (as genus K1 ) in a substantial morphological data matrix combining features of larvae, pupae and both sexes of adults, used to investigate relationships of Anuncotendipes Cranston and Nandeva Wiedenbrug, Reiss and Fittkau (Cranston 1999). Anuncotendipes evidently belongs with the Harnischia complex and is not discussed further. Nandeva, which remains unknown in the larval stage, associated with a clade comprising Australian wood miners K1 and Imparipecten Freeman (Cranston 1999) plus Nilodosis Kieffer and Fissimentum Cranston & Nolte (two taxa with distinctive cleft larval menta). The position of Nandeva varied, and in some analyses joined the monophyletic tribe Tanytarsini: most nodes were poorly supported. A critical misscoring (due to paucity and tenerality of available adults) was identified by Sæther and Roque (2004): in Nandeva the angle between wing veins RM and R 4+5, which had been scored as distinctly oblique, actually is very low at most, which is the sole diagnostic autapomorphy for Tanytarsini. However, even if rescored for this feature, Nandeva remains placed equivocally in parsimony analyses, most likely due to the lack of larval data and reduction of many features of the known stages. Interestingly, exclusion of Nandeva from analyses of Chironominae phylogeny little affects the proposed phylogenetic structure among other taxa. The ever-developing phylogenetic matrix assessed by Cranston (1999, 2003) has been expanded with additional taxa relevant to relationships of Xylochironomus, notably inclusion of Paralauterborniella Lenz, Apedilum Townes and Xestochironomus Sublette & Wirth (matrix available from author). Parsimony analysis was undertaken on this matrix of 66 generic-level taxa scored for 119 characters (of which 117 are informative) derived, where available, from all life history stages (character list available from author). Analysis used PAUP*4.0b10 installed on a Macintosh, with 1000 random addition repetitions, with all changes equally weighted and unordered. Brillia Kieffer of the Orthocladiinae is used (as outgroup) to root the tree. Over 3200 equally parsimonious minimum length trees resulted, of length 958 steps, which were visualised as a 50% majority rule tree. Nodes and relationships pertaining to Xylochironomus are summarised in Figure 5. Xylochironomus is postulated as sister group to Paralauterborniella and Apedilum. This group is sister to, sequentially, Fissimentum, Imparipecten and Nilodosis at increasing remove (Fig. 5). This clade appears in 73% of minimum length trees (MLT), with these internal relationships appearing in %. The clade appears to be sister group to the remaining Chironominae less Pseudochironomini and Nilothauma. Pseudochironomini (represented by Riethia, Aedokritus and Pseudochironomus) is paraphyletic in the MLT, but monophyletic in trees only one step longer (959 steps). Inclusion of Xestochironomus Sublette & Wirth to the matrix analysed by Cranston (2003) confirms sister group relationship to Stenochironomus postulated by Borkent (1984), in a robust clade that includes also wood-mining Australian Harrisius and Asian Shangomyia as in Cranston (2003). This clade, which is subsumed as wood group in Figure 5, remains monophyletic in analyses of the expanded morphological matrix. Based on earlier morphological analyses, Cranston and McKie (2006) noted that many early nodes in the phylogeny of the Chironomini can be reconstructed parsimoniously as being associated with the wood-mining habit. While this remains true with additional taxa included, support for critical nodes is weak (all lacking an asterisk (*) in Fig. 5 occur in less than 80% of MLT). To test certain possible relationships, Fig. 5. Cladogram of Xylochironomus gen. n. and postulated close relatives based on unweighted parsimony analysis of data matrix as available from author, wood group = Stenochironomus and relatives; connectens group = Paratendipes and relatives; Chironomini 1 = Polypedilum and relatives, Chironomini 2 = Chironomus and relatives.

8 234 P S Cranston constraints were applied in PAUP*, first to test monophyly of all wood miners that is wood group (Xestochironomus, Stenochironomus, Harrisius and Shangomyia) plus Xylochironomus, Imparipecten and Amakye, an undescribed African wood-mining taxon. With this group constrained to occur, MLT length increased by 9 steps to 968 steps. Next Xylochironomus + Polypedilum was tested to the exclusion of all others, resulting in an MLT of 967 steps, 8 steps longer. Thus, neither a close relationship of Xylochironomus to Polypedilum, or to the clade of wood miners centred on Stenochironomus is maximally parsimonious (although such trees are only 1% longer). Stronger support for hypothesised relationships (and evolutionary scenarios such as refugial wood mining) (Cranston & McKie 2006) is likely to come from molecular analyses in which multiple genes are combined with morphology (GA Morse and PS Cranston in prep. 2006). ACKNOWLEDGEMENTS I thank directors and staff of ERISS (formerly OSS), principally Chris Humphrey, for recognising that monitoring activities required a taxonomic framework and for providing financial and logistic resources, to sustain such studies. Chris also took time to find fresh material for molecular studies. Professors Nigel Stork and Richard Pearson of the Rainforest CRC supported similar studies of the biodiversity of Chironomidae in the wet tropics of north Queensland. REFERENCES Anderson NH A survey of aquatic insects associated with wood debris in New Zealand streams. Mauri Ora 10, Anderson NH Xylophagous Chironomidae from Oregon streams. Aquatic Insects 11, Borkent A Systematics of the Stenochironomus complex. Memoirs of the Entomological Society of Canada 128, Cranston PS The metamorphosis of Symposiocladius lignicola (Kieffer), n. gen., n. comb., a wood-mining Chironomidae (Diptera). Entomologica Scandinavica 13, Cranston PS Immature Chironomidae of the Alligator Rivers Region. Open File Record 82. Supervising Scientist for the Alligator Rivers Region. SSARR, Darwin, Australia. Cranston PS Morphology. In: Chironomidae: Biology and Ecology of Non-biting Midges (eds PD Armitage, PS Cranston & LCV Pinder), pp Chapman & Hall, London, UK. Cranston PS Identification Guide to the Chironomidae of New South Wales. AWT Identification Guide Number 1. Australian Water Technologies Pty Ltd, West Ryde, Australia. Cranston PS Two unusual Chironomini (Diptera: Chironomidae) from Australian rainforest streams: one new genus and a neotropical genus new for the region. Australian Journal of Entomology 38, Cranston PS Austrobrillia Freeman: immature stages, and a new species from the Neotropics. Spixiana 23, Cranston PS The oriental genus Shangomyia Sæther & Wang (Chironomidae: Diptera): immature stages, biology, putative relationships and the evolution of wood mining in chironomid larvae. Raffles Bulletin of Zoology 51, Cranston PS Ancient and modern toolboxes for e-bugs. Systematic Entomology 30, 1 3. Cranston PS & Hardwick R The immature stages and phylogeny of Imparipecten Freeman, an Australian endemic genus of woodmining chironomid (Diptera). Aquatic Insects 18, Cranston PS & McKie B Aquatic wood an insect perspective. USDA Forest Service Southern Research Station General Technical Report (in press). Cranston PS & Oliver DR Aquatic xylophagous Orthocladiinae systematics and ecology (Diptera: Chironomidae). Spixiana Supplement 14, Hoffmann A & Hering D Wood-associated macroinvertebrate fauna in Central European streams. International Review of Hydrobiology 85, Kaufman MG & King RH Colonization of wood substrates by the aquatic xylophage Xylotopus par (Diptera: Chironomidae) and a description of its life history. Canadian Journal of Zoology 65, Sæther OA & Roque FO New neotropical species of Nandeva (Diptera: Chironomidae), with a phylogeny of the Tanytarsini. Tijdschrift voor Entomologie 147, Accepted for publication 13 May 2006.