The Taxonomy and General Biology of Some Southern Australian

3ó .r-:5

BIOLOGY THE TAXONO¡4Y AND GENERAL

OF SOME (OIPIERA: nn¡¡etocnn¡) SOUTHERN AUSTRALIAI{ CHIRONOMIDAE

INGRID A. HERGSTROM B.Sc (ttons) Adel.

UNIVERSITY OF ADELAIDE DEPARTMENT OF ZOOLOGY'

L974

of Adelaide A thesis submitted to ttre University for the degree of Doctor of Philoscphy'

J DECLARATION

been accepted This thesis contains no material whictr- has at any university for the award of any other degree or diploma andtothebestofmyknowledgecontainsnomaterialprrblished reference is or written by anottrer Person' except where due

made.

(r. a. HERGSTRoM) CONTEI{TS

Page (i) SUMMARY (ii) ACKNOW'LEDGEME}CTS 1 I. INIRODUCTION 3 II. GENERAL BIOLOGY 3 I INTRODUCTION 4 2. METHODS 2.I Collecti-on 4

2.2 Emergence Data 6 2.3 Rearing 6 2.4 Behaviour 8 I 2.5 Measurement of Humidity I 2.6 lvleasurernent of Acidity I 2-7 Measurement of OxYgen Levels 9 3 COI,LECTING SITES 3.1 Site descriPtions 9 29 4 BIOLOGY OF INDIVIDUAL SPECIES SubfamilY Podonominae 30 32 Subf amilY APhroteniinae SubfamilY Diamesinae' Tril¡e HePtagYini 32 32 Sr:bf anilY Orthocladiinae 51 5. DISCUSSTON 54 III. MORPHOLOGY 54 1. INTRODUCTION 56 2. METHODS 2.I Collection of sPecimens 56 2.2 Preservation of sPecimens 56 2.3 PreParation of sPecimens 56 Page 2.4 Drawings 57 2.5 Recommendations regarding the preservation and curating of Chironomid collections 57 57 3 STRUCTURES OF TAXONOMIC IMPORTANCE 3.1 Adult 58 3.2 Pupa 65 3.3 Larva 65

4 SPECIES DESCRIPTIONS 66

Sub-familY APHROTENIINAE 68 Genus Aphroteniinae 68 Sub-family DIAMESINAE, TRIBE HEPTAGYINI 69 Genus ParahePtagYia 69

Sr¡b-f amilY ORTHOCIJIDI INAE 72 Genus Austrobrillia 76

Genus Cardiocladius 79

Genus Stfcbcladlus 80

Genus Eukiefferiella 84 86 Genus Svnorthocladius 'ee Genus Paratrictrocladius

Genus Cr 92

Genus Microcricotopus 98

Genus Nasuticladius 10r Genus Àustrocladius 117

Genus LimnoPhYes r29 Genus ParalimnoPhYes L32 Genus SParsicladius 136 143 Genus Nemoricladius

Genus KiefferoPhYes L47

Genus Parametriocn ernus L54 156 Genus Gymnolnel-r iocnemus Page

Genus Rhinocladius 160

Genus Smittia I63

Genus Parakief feriella 165

Genus Pseudosmittia 167

Genus Doloplastus r69

IV. PHYLOGENY 175

I INTRODUCTION 175

2. BRT]NDTN S USE OF HENNIGIS PIIYLOGENETTC SYSTE¡4ATTCS r78

3 PHYLOGENETIC SYSTEMATICS AND THE EVOLUTIONARY INDEX r84

4 A METHOD OF TND]CATING RELIABILITY OF PROPOSED PHYLOGENIES 186

5 PTIYLOGENETIC RE],ATTONSHIPS WITHTN TIIE SUB- FAMILY APHROTENITNAE 186

6. PHYLOGENETIC REI.ATIONSHTPS WITHIN THE SUB' FAMILY ORTHOCI,ADIINAE t87 6.1 Important Character Phylegsti.= 190

6.2 Genus Austrobrillia 193 6.3 cenus Cardiocladius ]-94 6.4 The Diplocladius-S tictocladius genus pa]-r 195

6.5 Genus Eukiefferiella 196

6.6. Genus SYnorÈhocladius l-97 6.7 Ttre Paratrichocladius groups of genera L97

6 .8 Genus Ml-crocricotopus 199 6.9 the Çb¿S.cla4ius group of genera 2OO

6. to The Nasutic lad.ius group 203

6. 11 Genus Parametriocnemus 206

6.L2 Genus jj¡g5l.!!ig 2c-7

6.13 Genus ParakfefferfellE 2U.7 Page

6.L4 The Pseudosmittia and Doloplastus pair 2o-7

6.15 Phylogenetic Tree of the Australian Orthocladiinae 208 209 V. NT]MER]CAL ANALYSIS 209 1. INTRODUCTION 2ro 2. 2IO 3.

4. 2LO vI. 2L4

References 2L7 SUMMARY

After collecting widely in the wetter regions of South Australia and with the assistance of a small amount of material collecÈed in other parts of south eastern Australia, I have provided descriptions and biological details of 73 species in 3 subfamilies of Chironomidae (Diptera : Nema- tocera), Èhe Aphroteniinae, Diamesinae (tribe Heptagyini) and Ortho- cladiinae. Of these, the single member of the Aphroteniinae and both species of Heptagyini are new species, while two genera and forty-eight species of Orthocladiinae are new.

I have examined the approach Èo taxonomy used by both the Mayr school and the Hennig school and also critically examined the methodology of Hennig's followers. Hennig's phylogenetic systematics has been adopted as the most suitable for the present work.

A proposal has been made for indicating the levelof reliability of phylogenetic estimates of relationship, and its use illustrated by a consideraÈion of relationships within the Orthocladiinae, f.or which an incomplete phylogeny has been suggested.. \ A phenogram has been prepared using the program of Lance g Williams (L967) on a computer, relating all Australian species of Orthocladiinae. This has then been compared with the proposals for phylogeny- The implications of this work in relation to biogeographical problerns are briefly examined. ':i

(i) ACKNOWLEDGEMENTS

My supervisor, Dr. D. A. Duckhouse has greatly assisted me through discussion and criticism of this manuscript: for this and for his continued encouragement, I sincerely thank hím'

I am grateful to Professor Andrewartha (recently retired) and to his successor, Mr. I. l'1. Thomas, for making available the facilities of the Zool-ogy DePartment.

some collections of specimens were made on privately owned properties and for access to these I thank Dr. D. Hea1ey, Mrs' Fisher, Mr. I. Robertson and Mï. L. E. M. Salter. Special thanks are due to Mr. and Mrs. salter for their hospitality and to Mr. salter for regularly tending my collecting apparatus over a period of a year and a ha1f.

I examined. many type specimens at c.s.I.R.O. Division of Entomolog!,

Canberra and I especially thank Dr. D. H. Colless and his staff for their assistance in this work and also for the loan of material from the Australian National Insect Collection. I examined other type specimens aÈ the laboratories of the school of public Health and Tropical Medicíne, sydney university, and I am grateful to Professor D. J. Lee for the use of these facilities. For typing the drafts and the larger part of this manuscript I

am indebted to my mother, M-rs. J. B. Hergstrom' I

SECTION I INTRODUCTION

The ísolated fauna of Australia and New Zealand poses many questions for systematists and biogeographers. Hennig (1960) outlined the problems posed by the then known Dipteran fauna of New Zealand and illustrated by example how phylogenetic systematics, his particular approach to the analysis of retationships, can provide a means of clarifying these questions and assist in answering some of them.

Hennig r,vas prevented from anal-ysíng relationships more extensively by a lack of detailed information about the groups involved. However, he exhorted specialist workers to be aware of the type of question to be answered and to approach the analysis of relationships in a way that may provide the answers- Brundin (1966) in his investigations of southern representatives of the subfamilies Podonominae, Aphroteniinae and Diamesinae of the fainily chironomidae responded to Hennig's plea and used the methods of phylogenetic systematics to analyse the relationships between species of these sub- fanilies in depth. on Èhe basis of this analysis he came to far reactring biogeographical and geological conclusions. Brundin uses Hennig's principles with great flexibility, but if his extrapolaÈions were valid then this approach would be a useful and powerful tool for solving

biogeographical Problems - If, however, his extrapolations are ulìlvarranted, care must also be taken in evaluating other works written in this style (e.g. Hirvenoja, Lg73). Considerable criticism has already been made both of Hennigrs phylogenetic systematics and of Brundin's application of these principles' I have approached the problem througlr- a consideration of some of these criticisms and by closely ana]-ysing some of Bru¡rdinrs proposals' In addition I have attempted to use Hennig's methods to construct a partial phylogeny of a second group, the Orthocladiinae' 2

A survey of the literaÈure (Brundi¡r, 1956; Edwards, 1931; Freeman' 1956, Ig5g, 1961) showed that of the remaining sr:bfanilies of Chironomid.ae, the Orthoclacliinae include genera with a distrilcution in the southern la¡d masses similar to that of tJ.e groups studied by Brundin. An analysis of the relationships within Orttrocladiinae could thus provide an j¡dication of the plausibility of Brundin's conclusions. For a nuniber of reasons it seemed wiser to choose Orthocladiinae for this analysis, rather than an entirely different familY. l. The characters of Orthoctadiinae would be somewhat si:nil-ar to those of the groups analysed by Brundin and thus migtr-t provide some clues about the accuracy of his evaluations of synapomorphies and synplesiomorphies-

2. In many cases Orthocladiinae are found in places where Podonomi¡rae, Aphroteniinae and Heptagryini would also be expected and thus while collecting Orthocladiinae, ne\¡¡ species of the latter might be taken which would provide a test of relationships proposed by Brundin. 3. A preliminary survey showed Orthocladiinae to be widespread and abundant in areas of South Australia not completely altered by human activity. So that proposals of relationships could be broadl-y based, I have co1lecÈed extensively and made investigations into tlre bioloqy (SECTION II) of as many species as possible. In proposing relationships I have also nade use of phenetic methods of analysis (SECTION V). 3

SBCTION II GENERAL BIOLOGY

1. INTRODUCTION Australian Chironomidae outside Chironominae have only rarely been collected on a large scale (Australian National Insect Collections,' Brundin 1966) and biological studies have been restrícted in scope (D. H. Edward, 1961; D. H. Edward and D. H. colfess, 1968; R. Jones, L973). At the outset of the present work almost nothing was known of the South Australian faqna and my objective has been to obtain a preliminary but comprehensive picture of ttre diversity of species associated wíth lotic habitats in this state. By collecting widely and using a variety of methods for trapping, extracting and rearing, I have attempted, to discover the immatures as well as the adults of as many species as possiJcle. In addition' observations have been made of such habitat preferences, tolerances and behaviour as might be useful in evaluating characters for phylogenetic purposes, Most of the collecting sites are, or were until recent clearing, surrounded by native woodland or forest comprising a dominant eucalypt and an associated understorey in which species diversity increases in the vatleys. Most of the areas sampled are characterised. by acidic soils and most waters are also slightly acidic. Maps I and II show average yearly rainfall and relief of the regions sampled and graphs I and 2 indicate the seasonal rainfall pattern in the wettest and the driest regions sampled' I did not attempt quantitative ecological work because of doubts as to its relevance and because of the difficulty of preserving ttre populations of species at small sites while still- carrying out meaningful sanpling' Sampling was also hampered by frogs, spiders and other consumers of ch-iro- nomids entering the traps and by the difficulty of constructing floating traps which are effective +*hroughout fluctuations in water level' 4

2 METHODS

2.I COLLECTION 2.LI Adults. Four methods of capturing adults were used.

2.LLL Sweep Net. The net used (see Fig.l) was designed by D.A.Duckhouse. Its closed end is squared, not tapered, slightly more than half the circum- ference of the frame and, fixed to the frame with one corner adjacent to the handle. V'fhen vegetation is swept with a rapid figure of eight stroke, leaves and large insects are thrown to the furthest corner while smaller insects and

seeds fall into the corner closest to the handle; in this way the more deticate insects are protected from damage. 2-LL2 Liqht Trap. The tr ap used was similar to that developed by J- T.

Headlee & T.D. Mulhern (in Peterson, 1953, P]ate 83, Fig.I) but rendered highly portable by modificatio¡rs which made it possible to run the fan and a 4.5V. bulb from a single 6V. battery. The globe lasted only about 50 hours but gave a brighter light than normal. Because large moths tended to remain on and thus block any mesh barrier preventing them from entering the collection, such barriers were soon abandoned. It was found important also to be sure that fumes of volatile collecting agents - usually alcohol - dispersed

over a wi

and readily identifiable. 'Tanglefootr proved too sticky and paper coated with castor oit was abandoned because it was consumed by birds'

2.L2 Larvae and Pupae. S amples of sr¡bstrate which could not be suciess-

fU1ty examined in the field were transported back to the laboratory at OoC. - 5oC. in a polystyrene box. Larvae extracted in the field were also kept at oo c. - 5oc. during transportation' 2.L2I Stream nets. By using these Brundin (for details see Brundin, L966,

5 p.72) rt¡as able to collect large nurnbers of immatr:res and casts which remained in good condition provided the nets were cleared every 2 hours. partly cmerged adul-ts were often prevented from completing their eclosion and the value of this method was thus enhanced. However, stream nets were less useful in South Australia. !ùater flow at most sites sampled is so slow for most of the year that catches are small and the eclosion of trapped pupae is not prevented. Vùhen flow increases, generally after rain storms, large amounts of debris are carried into the water which probably provides protection for ecloding adults as well as greatly reducing the flow through the net. 2.12 2 ExEraction from sampfes of substrate. Larvae and pupae were extracted only in ways which might provide life material for rearing and thus permit

i-mmatures to be linked wittr adults.

(a) From al mats. Iv1any larvae inhabiting algal mats and Nostoc- Lycopodium associations in running water come to the surface shortly after the material is removed to still water. They car¡ then be picked off with a Pasteur pipette.

(b) From decavinq or other org anic and particulate matter. One method

used was an adaptation of that developed by Bay (1967) for separating dipteran pupae from sand (refer to Fig.2). After the sample has been placed under the screen B, water flow is adjusted so that screen C remains submerged. Immatures and some organic particles are carried througlr B and the

i¡nmatures and. smaller organic particles pass through C Èo be collected and separated into two size classes by screens D and E. The i-mrnatures are then picked off the screens under a dissecting microscope- The success of ex-

traction depends on the larva or pupa releasing hold of substrate material and is therefore nosÈ successful with free living species. Utrfortunately speeimens separated by this means are often damaged and cannot be reared.

A second method involves covering the sample witl. water in a large flat dish through which a gentle cr.rrrent is maintained. The overflow falls a short distance onto a partly submerged fine mesh screen and imrnatures and casts coltect at the meniscus. This method decreases the amorrnt of damage Eí9. 2

a L +- A w B

A - plastic bucket /\

B nesh (1 mesh 1 crn) / - E \ C - mesh (4 meshes Ì cxn)

D .- rnesh (15 meshes 1 cm)

E - mesh (3o meshes I cm)

S -. =a*ple of sr¡bstrate L - water leve1 W- water supply

Apparatus for separating i-mmatures from substrate samples. 6 to specimens but also decreases efficiency: ít was most successful in obtaining pupal casts of semiterrestrial species-

2.2 EMERGENCE DAÎA

Emergence traps were set with the intention of (a) finding out where final instar larvae live, (b) discovering when adulÈs of the various species emerge, (c) obtaining an estimate of population density for various species.

However, information discovered about (a) and (b) may not be completely representative and no meaningful estimate of (c) could be made because:- (i) at the sites the substrate was patchy and it was not always possible

to move the trap to a completely new position each !üeek, (ii) if a trap were placed over an area through which water was movi-ng, some alteration in water flow could not be avoidedt oxygen levels and biological drift would also be altered in these areas, (iii) spiders often consumed large numbers of speci:nens from collecting vessels. It was sometimes possible to infer, from remaining wings, which species these represented. Other spiders building webs across inlets to collecting vessels must have actually excluded flies. 2-2I Flowerpot traps (Plate f). Earthenware flower pots, 12 inches in d,iameter, with a transparent plastic collecting vessel affixed to the central drain hole were inverted over the site from which samples were required. J. D. Foulk (1966) first used these pots because tre claimed their porosity red.uced condensation in the collecting vessel and spiders favoured them less than other traps. 2.22 Cone traps (plate 2). A cone of coarse plastic mestr was used to support an inside cone of black nylon material which opened into a collecting vessel. The area covered by this cone \^zas one square metre.

2.3 REARING

2. 3r Rearing of isolated j-mmatures. Larvae were kept indivíduaIly at

L2oC. in 7 mI vials or small petri dishes containi¡g a small amount of carefulty screened, dantp sulrstrate material . Many larvae died, damage'J by 7 extraction, over-exposed to chlori¡rated water during extraction, or attacked by firngi. Pupae were kept in smal-l vials on damp cotton\4ioo1 or filter paper

The connection betweerr life stages r{as accepted only when established at least once througtr rearing specimens individually-

2.32 Rearing from unsorted samples. S amples of substrate were kept at o>çygen, temperature and moisture levels similar to those in the fietd. By collecting emerged adults frequently and at each collection carefully examining the sample for cast pupal skins, occasionally witå last larval skins attached, support vras sometimes obÈained for the sequences established in

2. 31.

2.32L Terrestrial or semiterrestrial species. S amples of leaves, mats of grass, debris, lichen and ground moss were kept outside during the winter months. During the summer similar collections were kept at 15oC.; to red.uce the drying effect of the constant temperature cabinet, the containers v/ere covered with plastic as well as gauze-

2-322 tic s ecies. Material from aquatic sources was kept at I2oC.

Samples of stream-bed Nostoc-Lycopsrdium associations I green algae and midstream accumulations of dead leaves were partially submerged' while moss \¡Ias supported at \ttater level. 2.33 Fiel d rearinq. Particular larvae from aquatic sources failed to survive transportation so, at ttindmarsh-Fa1Ls, several attearpts were made to rear these in the field. The larvae were placed ín a shallow sock, R, (see Fig. 3) with the algae or aquatic plants from which they had been separated and the sock was supported just below water level by a floating plastic container. The container was divided into separate eclosion and collecting cha¡ribers, E and C, and kept ür an upright position by a tapered cylind'er of coarse plastic mestr, ¡4, one meÈre hiqh- The container with- the sock attachr:d was thus free to move up and down with fluctuations in water leveI- Un- fortunately these ctrambers could not be sufficiently braced against battering by debris carried i¡r a strong current without inordinate expense and so were abandoned. ris. 3

c 'l

E --,..-,*, \¡[¡f 1m

$ - braces

M- cyli¡¡der of strong plastic mesh (1 meshr/I.S cms) C - sol-id transparent plastic collecting charnber R - mesh sock supporting larvae E - solid transparent plastic eclosion chamber W- water level

Apparatus for rearinq larvae in the field I

2.4 BEHAVIOUR 2.4L Adults.

(i) Swarmins. f recorded the size of swarmsr their height above ground level, the nature of flight within ttrem, their attachment to markers as well- as the Èime of day when they were observed.

(ii) Mating. The onty mating observed was that resulting from swarming activity; a white plastic sheet was placed on the ground under a swarm and sometimes a pair leaving the swarm \¡ras revealed as it landed on the sheet and could thus be captured. (iii) Egg laying. The activities of females resti ng on stream banks or other moist surfaces or flying low across water were closely observed and, when necessary, the females vrere capturecl using an aspirator. (iv) Feeding. On numerous occasions the flowers of shrubs and herbs in the field were closely examined for the presence of adults attempting to collect nectar or dew. In the laboratory adults were subjected to a small degree of desiccation and then presented with fine droplets of distilled wa water. 2.42 lrunatures. Ihave recorded whether pupae and larvae are free-living or tr¡be-dwelling and, where appropriate, described the type of Èube built. The readiness of immatures to move and their ability to swim and, move through debris is noted. Feeding l-arvae were closely observed and, where large nr¡ribers of specimens were available, f examined stomach contents.

2.5 MEASUREMENT OF HUMIDITY

A sling hygrometer $¿'as used to determi¡re relative trumidities in collecting areas. No attempt was made to determine the humidiÈy of the often secluded resting places of adult flies.

2.6 MEASUREMBNT OF ACTDITY pH was determined in the field to within O.2 of a unit.

2-7 MEASURE¡4ENT OF OXYGEN LEVELS In order to determine a suitable level of aerat.ion for laboratory rearing a Titron o>(ygen electrode with a low response time was used in the field to determine oxygen levels in the various habitats at the major sites and at some minor sites. GRAPH I Monthly rainfall at Mt. Lofty sunmit, just above Botanic Gardens site.

GRAPH 2 Monthly rainfall_ at Nuriootpa, near Tanunda Creek site.

Horizontal Axis Months of tJ-e year Vertical Axis Rainfall in points 700

600

500

400

300

200

I00

300

200

100

0 JJ 9

3 COLLECTING SITES collections were taken from sixty-seven varied localities in south Australia. A general picture of the south Australian fauna was obtained' byexaminingtheseinconjunctionwithcollectionsmadebyD.A.

Duckhouse, B. Glover and in the Australian National Insect collection (ANIC). Addit,ional information from the ANIC provided data on the Australia-wide range of some species'

From the above localities, five which offered the greatest diversity of species and habitats were selected for intensive study and have been called ,major sites" the remaining sites are called 'minor sites"

3.1 SfTE DESCRTPTIONS Dailya¡rdseasonalvariationsoccurinthephysicalandchemicalen- vironment of any freshwater organism and only extensive measurement coul-d plant provide a general idea of the conditions at each site' Ho\^¡eveÏ' the influence of the community at any site will be indicatíve of the overall Thus environment and will not alter with minor environmental fluctuations' aknowledgeofplantcommunitieswj-Ilenablequickcharacterizationor evaluaÈionofcollectingsites.Ihavethereforemad.earecordofEhe (taj¡les I IV) in major plant species and associations found at each site - addition to recording some of the more important physical characteristics (197r) were corr- (ra¡Ies V - vII) . The \^¡orks of Hynes (1970) and Pennak of all sulted in selecting the latter. Maps I and II show the positions sitessanpled.(SiteabbreviatíonsareincludedinTablesl-]V). Nositesampledhad'awaterflowgreaterthan2.5lanimal groups may already have dimínished under these unnatural conditions' 3.1I Maior Sites Frequent visits were made to these sites over a period of one and a half years or more- 3.III Parawa (Section 323, Hundred of V'Iaitpingat property of L'E'M'

Salter) . The collecting site is on the southern aspect, close to the summit of the southernmost ridge of the Mount Lofty fault scarp. It is in a smafl gully bordered by eucatypts and includes some of the sor:rce springs of gully than GoId Digger,s swamp. Humidity remains 5? - 15% higher in the on ttre hiIItoP. over most of the period July 1969 - October 197I I made fortnightly net collections, and with the help of Mr. salter, weekly light trap and indicated emergence trap collections. In Fig. 4 the major habitats are by the letters A to J; the flowerpot emergence traps were placed at positions (i) to (vi), cone traps at positions I to v and substrate in Plates samples were taken from positions I to 13. The areas pictured I to V are marked on the figure' River 3.IL2 Hindmar:sh Fal1s. About 15 kms from the sea the llindmarsh about I km' passes over a series of rapids and fal-Is' dropping 90 metres in FIG. 4 I',IAP Otr' COLLECT]NG SITE AT PARAVqA Habitat: o A Hillside, sloping at 30 down to F.

B Mud.bed, compacted and extending into H.

c Streamlet, flowing over bare or moss-covered ground.

D Shaded leaf accumulations, kept moisÈ by spring water.

E Shaltow bog (Plate III, foreground) .

F Deep bog (Plate I, background) i a small spring appeared January I97O.

G Sheltered bog (plate rr) fed by strong seepage.

H Ford (Plate IV) receives water from G, ï and, C.

I Spring (Plate rV, background) in centre of deep bog overgrown by reeds.

J Ford (Plate V) , covered drrring winter, water flows between stones during sutnmer.

(i) (vi), positions of flowerpot emergence trap. I - V , positions of cone trap. I - 13 , positions of substrate samples.

rrees Water O o

.'.i:'. :. I, Grass interspersed Areas shown in plates. ffi with moss. n Herbaceous undergrowth Spring Bog * Ford Xanthorrhoea ,örm àK V Re¿ds +v

.t \ tl t!-,

.\- \t !

,t, I

,ììJ ¡ : PLATE I

Flower pot trap, in situ, Habitat E, position (ii), Parawa (winter). -À \\ PT,ATE TT

Cone trap, in situ, position V, parawa.

PI,ATE III

Habitat E (summer), Parawa

PLATE IV

Habitat .H, (winter) , Parawa

PI,ATE V

Habitat J (winter) Parawa

This collecting site includes the approach to the first series of rapids (r'ig. 5, positions 2 - 14) and the actual rapids (posit-ions 15 - 2Il. It is in a reserve where native vegetation has been preserved and both the gully slopes and floor are covered with dense undergrowth and shaded by eucalypt forest and ash trees as far as position 14. From position 15 to 21 the stream flows turbulently over projecting rock surfaces on the \^lestern. side of the gu1ly floor and the steep rock walrs shade the water course during early morni'ng and late afternoon' Net collections were made throughout the area every fortnigtrt' unfortunately emergence traps set out were eitlrer wastred away by the current or souveníred by tourists. stream nets were set out for four hour periods at positions 9, L4, 16 and 17 on a number of occasio¡rs and sub- pictured strate samples were taken from aII other positiOns. The areas in Plates Vl to VIIÍ are marked on the figure' on Trrnkalill-a Creek 3.113 Eric BonVthon Reserve. A six acre reserve of about 5 kn. from parawa. Thick undergrowttr, including stands -B_lechn\In

and Gleichenia extend right to the water's edge in a narrow,protected gulIy. Net collections were made throughout this area every tv¡o weeks. Substrate samples were taken from under stands of Gleichenia from the of a stream bank overhung bY Gleichenia or Blechnum and from the surfaces large rock protruding into a deep midstream pool' on the eastern 3. 114 Botanic Gardens Mt. Lof . This is a reserve at slopes of Mt. r,ofty and the collecting site is just below the summit the head of a narrow valley. Dense growtJrs of .Gleichenia and Þlefu almost obscure numerous springs and seepages. Run off is collected in a maintain small artificial dam which overflows continuously and helps natural vegetation in part of the lower valley' Weeklylighttrap,oilypaperandnetco]-lectionswerentadeandsub- a tiny grassy strate samples were taken from the followi.ng places: FTG. 5 MAP OF COLLECTING STTE AT HTNDMARSIT FALLS Terrestrial habitats: Position I Forested area at head of guIly 2r4ì5 Ground protected from dessi""iion by thick debris.

3 Debris collecÈed in stems of Xanthorrhoea.

tic habitats Position 6,13 Dead leaves accumulated on fallen branches and forming a sieve.

7 Tree roots from stream lloor.

I AJ-gae growinq in early spring and. late autumn on concret.e ford.

10 Bank of deep pool.

11 ,IIarb Mosses growing in splash zone.

T2 Reeds and rock surfaces carrying Simulium larvae.

L4 Stream bed with growth of Siwn Iatifolium.

15,18 Nostoc-lycgpoc!!¡4m assoc iation .

19 Vüiry filamentous algae growing on perpendicular rock face.

20 Soft sandstone buttress, algae and Lycopodium survive in crevices.

2L Seasonal rock pool.

9 ,L4 rL6 rl-'l Stream net collections.

Water Fallen vegetation

Forest with d.ense Xanthorrhoea undergrowth.

Grass Spring

Broken rocl< face

High Rock lfall A-reas shown in plates. Ford

PI,ATE VT A

Looking downstream towards position 17 (summer) Hindmarsh FalIs

PI,ATE VIb

Looking downstream towards position 17 (winter) Hindmarsh Falls /- PI,ATE VIIA

Looking upstream towards position 13 (suruner) Hindmarsh Falls

PLATE VI.Ib

Looking upstream towards position 13 (,v¡:j¡rter) Hindmarstr Fal1s

PTÄTE VIIT

Lookíng towards position 14, Hindmarsh Falls *,¡ PLATE IX

Stand of Gleichenia microphylla, Bota¡íc Gardens'

clearing near the main streamlet, a region where the streamlet is submerged beneath Blechnum debris, the margins of the dam and' from beneath

stands of Gleichenia (Plate IX). 3.II5 Torrens Lake. The River Torrens drains a considerable area of the Adelaide HiIIs and its tributaries include Deep Creek (Adelaide Hills) ancl sixth creek. In the Adelaide city area the river is dammed back to forvn an artificial lake which is draine'd each winter but refills before the

banks have dried out significantly. Every week net samples were taken from a sheltered section of 'Ehe bank near a small reed bed and samples of mud and debris were taken from

the bank.

3.L2 Minor Sites. These were visited only occasionally and the number of visits to each is indicated in the Tables v to vII. I Aldqate creek. A tributary of the River onkaparinga in a narrovl region of a valley just south of Aldgate' 2 Alnut, A dam at the headwaters of GoId Digger's Swamp; overflow runs through a striP of natural scrub' 3 Bethany. Lower regions of Tanunda Creek (q.v') where it flows through vineyards.

4 Blewitt SPrinqs. A hollow, a few acres in exÈent, contaíning a nunlcer of small springs in sandy ground; completely covered with water during winter. 5 Brownhill creek. In medium width va1ley of the northernmost branch of the creek. 6 Bul1's Creek. In medium wi

stream restricted to very narro\¡¡ channel '

7 Carey 's Gu11v. In a shallow valley 3 km- from Carey's Gullyt seepage areas covered bY forest. I Chamber's Creek. A trilcutary to the Sturt River in a large va11ey abutted by domestic gardens. MAP I SOUTH AUSTRALIA o between Lat. 34o3o' 35 30' S.

o Long. 1380 - 139 E.

Scale: I cm- = 10 Km.

Hil 20001

ffi 1500' - 20001

H looo: - 1500t

500r - 10001

o' - 500'

- 25 - Rainfall (inches) I

I I I \ ' ¿.

I 20

u 2E MAP II SOUTH AUSTRALIA between Lat. 34o3o' - 3Bo s

Long. l38o - 141.0 s

Scale: 1 cm. 50 Km. t.- ri I ---10- t¡ I I tl See Map I Y2- li'--t\..2--- - -{4 s E ua--/ --16 .-

Tl: - 20--

MQ BO $ - a24 I \ I 28* trl r' IL BL- L V L L t-M \

5oo' - 1000' o' - 500'

25 - ¡tainfal.I (inches) 13

g clarendon. A spring-fed tributary of the River onkaparinga passing betweensteepcleared'hillsfrequentedbybeefcattle. 10 cleland Reserve. The source area of First creeki forest with dense understorey covering numerous springs and seepages' llDeepCreek(SouthCoast).Asmallcreek,about2km'fromitsmouttu valley narro\¡¡, densely vegetated' region 12 Deep Creek (Adelaide Hills). A creek of the Adelaide Hills and a tributary of sixth creek, running between a market gard'en and forest. 13 Duck ponds. A tributary of Duck Ponds creek running through a deep cutting in a strallow valley. Streambed vegetation ripped away by seasonal flooding- L4 Finniss River. Just north of Gilbert's, about Io km' from mouth; I - 2m. deep, 2Om. wide with dense reed Jreds' I5 Fishers. At one of the source springs of sturt River in a small, steep, forested valleY.

16 GlenView.AseasonaltributaryoftheNorthParaRiverwhereit with energes from the wooded Barossa escarpment to cross slopes planted vines. Creek and' its L7 Greenhill Ouarries. Quarrying has denuded Second surroundsofvegetation;temporarybackwaterscontainingonly fitamentous algae. 18 Hackham. A shallow west-facing valtey with scattered springs; a tributary of the onkaparinga in a valley grazed by dairy herds' running into I9 Happv VaIlev. About IoO ¡n. from the rnouth of a stream HappyValleyReservoir.Thefinelysiltedbedisprotectedbyd-ense reeds. in a steep 20 Healev t s. A tiny spring-fed tributary of the Onkaparinga valleyr3km.westofBigg'sFlatPostOffice'Anemergencetrap frere was abandoned after a few v¡eeks because of spiders ' L4

the head of a 2T Humbuq scrub. A broad, heath-covered seepage area at shallow valleY.

22 Inman Vallev.TheInmanRiverat'GlacierRock'ianarrowchannel cut into the broad valleY floor' 23 Kenney's HiIl. A tributary of the somme River in high flat country 3 km. north of Eden ValIeY' 24 Kuitpo. Meadows creek near Kuitpo Forest headquarters, flowing slowly through pine plantations and woodland' 25 Lower Hermitase. The narrow valley of South Para River' The sandy riverbed is subject to torrential flows' 26 Marne River. A broad cutting among low hills between springton and

Eden ValIeY. 27 Moculta. Duck Ponds creek, about 2 km- east of Moculta, flowing slorvly througtr, a shallow cutting in level ground near the western

edge of the Barossa escarpmenti oçosed to strong winds' 28 lvlorialta. The narrow valley of Fourth cr:eek, below the lowest

waterfall - 29 Nietske. A tributaty or tlre North- Para River at the base of the Barossa escarpment, flowing through a shaded humus-filfed hollow' 30 North Para River. A concrete ford 3 km' south of Tanunda; water heavily contaminated with- micro-organisms from rnzine vintage' 3I Orfando. The narrow cutting of Jacob's creek where it crosses the main road, to Tanunda just north- of Rowlandrs Flat' of 32 g.. An ill-kept dam straddling a seepage area of a tributary

Deep Creek (South Coast) ' 33Reynella.Theshat]-owcuttingoftheFieldRiverafterpassing

through vineYardsr ' Boat Harbour Creek in a 34 Robertson,s. One of tkre source Seepages of broad, partiallY grazed' va1leY' Mount Lofty 35 Saunder,s creek. Descends the eastern escarpment of

ranges through a narrow, weJ'I-vegetated va'Iley' l_5

broad', partly forested' 36 Scott's Creek. One of t].e source springs in a valley. (An emergence trap set from August 1969 to April I97O was finally abandoned because of spiders ' ) from the mouth of 37 Second Valley A group of springs about 2 km' this creek resulÈ in the lower reaches, which flow through limestone rock, having a permanent supply of water' joins River Torrens in a deep 38 sixth creek. Immediately before it the cutting at the bottom of a steep-sided valley' retained 39 Springton. A spring-fed pool immediately south of Springton, by a rocky outcrop and frequented by dairy herds' bed 40 sturt River. A narro\^r, protected region of this valley; stream April' covered by roots and densely shaded from October to open forest of the 4L Tanunda Creek. The region where it leaves the plateau to pass into a d'ensely forested valley' 42 Tooperang- On Tookagerth Creek' in a shallow' cleared va11ey' Creek in the region 43 lrlaterfall GuIlv. The steep-sided va1ley of First between tfirstt and 'second' waterfalls. in a broad' shallow 44 Western HilI Creek. A tributary of Meadows Creek valley. 4 km' north of 45 Williamstown. A tributary of North Para River about by cattle' the townshiP, flowing through low hills; frequented occasionally sampling in the south East of the state involved one or twovisitstoovertwentysites'manyofwhichwereverysimilarinec- groups' ological characteristics. They have therefore been presented in TheslzmbolaccompanyingeachsiteisusedtomarkitspositiononMapll.

46 Lake Albert at Meningie (MG)' ad acent swamp lands. 47 Small claYPans associated with the Coorong or I (SY) old Main Road (OL) Willow (Wvl) , Waltowa East (mË) , SaIt Creek ' ' Tilley's SwamP (TI). T6

the usuall 4B TC s south of the sent limit of (FL)' containinq some water. num (RM), Lake Frome ion. Brownts 49 Lakes of the mulÈi le volcanic cone Mt. Gambier (tM) Lake (BL), VaIIey Lake (VL), Leg of Mutton Lake ' and flowing a few 50 e freshwa ter spr INctS q from Iimestone (PI), Ewenrs Ponds (ne¡ hundred metres to the sea. Piccaninnie Ponds present ng and earlier 51 Marshes and boqs; remnants of tlte Cooro similar formations. These have been severely reduced in extent over (Bo) the past 20 - 30 years by systematic draining' BooI Lagoon ' canunda(cN)-adeepbogofwhichthesurface20cms.driesoutin

sunmer excePt around sPrings' de epened to drain marshes. 52 Drains speciall v constructed or creeks (MQ)' Benara Creek (BN), Tantanoola Creek (TT), Mosquito Creek lVater Holes (Gvü) - recently 53 In1and spr l-nqs and sinkholes. Green dried,snuggery(SN)-aspring,BellumBellum(BE)-asin]

KEY TO TABLES I TO IV

Veqetation form:- The structural form of the natural plant community surrounding each collecting site has been named in accordance with the classification scheme set out by specht (L972, p. 32-4c). Only 7 structural forms are represented at the sites:- open forest, (1); woodland, (2r; closed scn¡b (3); open heath, (4) ¡ open scrub, (5) i coastal dune, (6); tussock grassland , (7); natural community completely destroyed, (8). Large shrubs and trees.

MYRTACEAE

(2) ¡ cosmophylla Eucalvptus : - obliqua L'H., (r); baxteri M. & 8.,

F. v. M., (1) , leucoxylon F. v M., ( 4) fasiculosa D. (6); rubidaD&M.' (7)¡ F. v. M., (5) ; camuldulensis ' odorata B. & S., (8) ; diversifolia Bo., (9) . (2) Br. (3); Acacia: - mvrtifoliâ Vt., (r) ; plcnantr,a e., ¡ armata R. , (5) rhetinoj-des S. , (9 r rupicola F. v- M., . (2) Leptospermum: - iuniperinun Sm., (1) ; pubescens L., ¡ myrsinoides euca sguarrosa s., (1 t Melaleuca squamea Lb., (!, MelaI oonn (5).

OLEACEAE Fraxinus oxvcarpa lrl . (1); OIea europea' L., Q) .

CASUARINACEAE

(I) muellerana Mi.q- , (2) - Casuarina:- stricta Ait. ' Medium and small bushes.

PROæEACEAE

(2) . Banksia:- ornata F. v. M (!, ma::ginata Cav. , Hakea:- rostrata F. V- M., (!' rugosa R. Br {2) .

I,ILLIACBAE

(I) semiplana F. v. M (2) Xanthorrhoea: - australis R. Br. ' ; riln LEGUMINOSAE Daviesia ulcifolia Sm-, (r); Platylobium obt

H., (2)i Pult,enaea (Ð. 18

(I) RHAMNACEAE Spvridium parvifolium (H.) I'. v. M., ' (1) DILLENIACEAE Fiibbertia stricta R- Br f

(1) conostephioicles (Soncl . ) EPACRTDACEAE Epacris imPressa Lb.,' ; Astroloma F. v. 14., (2) .

GOODENIACEAE Goodenia sP.

ROSACEAE Rubus fructicosus L., (I) .

L. (1) . SCROPHULARIACEAE Gratiota Þeruviana ' Umbe IIiferae. (1) Sium tatifolium BI. Q) . UMBELLIFERAE Foeni-culum vulgare M., ; ' Ferns.

sP. (2), FTLICALES POLYPODIACEAE: Blechnum sP' (!' Pleridium

Adiantum ae thiopicum L. (3). GLEICHENIACEAE: Gleichenia Iicrop}y!þ Sz. , (41 . Se.lqe P1ants. (3); Sedge plants. TYPHACEAE, (1) ; CYPERACEAE' (2) ¡ RESTIONACEAE'

JUNCACEAE, (4) . Aquatic AngiosPerm' R. Br., (1); Aquatic angiosperms. CRUCIFERACEAE : Nasturtium officinale Azolla sp' (3) LEMNACEAE, Lemna, sP-, Q) SAIVÏNIACEAE, ' '

CHLOROPÍTYTA (r); Algae. Cladophora sp- (targe strong wafled celfs),

(smaller, less robust cells) r (2) ì

Spirocrvra sP. , (3) r Chara sP- (4).

Tota I species descríbed in Total species. Thi-s is the total numher of those specie-s

SECTION III and which I have found at each- site' t9

Abbreviations of authorts names* used in Keys I to IV A.-Aitonr!'t.;8.&S.-BehrrIl'etschlechtendal'D'F'L'; Dehnhardt' F'; Bo. - Bonpland, A.t ç-av.. - Cavanilles, A' iI'; D' - Mueller; D. & M. - Deane, H. et t{aiden, J' H"; F' v' M' - F' von Lamarck' J' H. - Hooker, W- J.i Lb. - Labillardiere' J' J' H'; L' - B. der L'H. - LrHeritiet, C. L.¡ M' & B' - Maiden' J'H' et Blake1ey' !{. F.; M. - MiIIer, P'; Miq'- Miquel' F' A' !'I'¡ E:-Br'- Brown' Robert; S. - Schlechtendal, D'F' L'; Sm' - Smith' Sir J' E'; Willdenow, C' L' Sz. - Swartz, O. i We. - V{endland, H- L.; W' -

*As cited by J. M. Black. EL¡HtfliÉ|d E I I X. Ë',H o õ'-iiúotrFl F,l xq É z.;i^, Ë a FIH*iåflËHE g.-a g ä H. P' þ,S rt t; q ã ç hu< o foÉöond * 3 [äB a H ö,lËHE 5H H [*'iísi oFo o o o FI o

(¡ts@HHP(¡ Vegetation form HæHHl\)N }JN (¡ ïI- iI Ï EucaIYPtus l-ôorun|or (' 'Þ (ir')tJurNu)".'l. lJ H UN Acacia - tl-ll Nl HHfN¡Þ'ÞÈ rÞ ¡Þ H 'tl(¡.llHP ìl ïH TiIÏ LeptosPermum (, l,l (, ó(,td('ur ' H llHllHl Oleaceae tlNllll N H H H Casuarina H H H t\) É H N N) N N Banksia Þ N N t\) N trtÉ P t{ P H H F Hakea H ts H H N N t\) H N N N N N N Xanthorrtroea N t\) N H H N H (, I I r", ts (rHl I (, (¡, (, Leguminosae I lN (, (, (, P H ts H H Rannaceae H P

P ts H H H Dilleniaceae Hll P trHl P H ts P H ,P Epacridaceae ItNlì\rts N N) L r¡ll N IHHH ts Goodeniaceae HllHlllfFl lPll I Rosaceae HHHIHHHIIH llll H ScroPhulariaceae ftlfllltll H N I umbelliferae ¡l,llPlHll H H N N P N N N N Tilicales l.TNNNN N I ¡Þ È t¡) N N N N) N N H I I Sedge rÞ Plants U) È Å (^, (, Aquatic lNl N N N tNllNfNl anglosPerms

NHH y A1gae N) N I¡)N -t N(,N Hu)N (r(rN (, t¿ (¡)

(^) rÞ È È Total sPecies \¡ @ ul N) H or H (, \O \¡ \¡

oz H. òF.ÍD'àîõ'EU.Ht-P'H'trãã5iÉiÉFfoohdPZFF 5 o E. Fl ñfroS<.õ-<'.Ð3< '$Ë s? FË H g H i i;; (/) ilgr' 5 o (D t-. o IE H (n o d H o .. ír r-1 H.fö-tri'äF.*)rlÉ o 7.Fr¿ç.arolâÈrot-lH v, crËd'éaÞv) o Þ"õBffg o 3 I l-'F's rf ¡¡ - |-. o Þ E8p, Ë i4o o ts. ct Ê Fts o

N H ts form IgHNNHHNNHN ts N Vegetation (, NÞHHF N (¡ ot--iO-i(p@r'Þ Ol È (¡ EucalyPtus irorNNN (Jl H HU '¡'¡lPHl H ts N) Acacia lllUJ'l ÈÈ ¡Þ HH H ts I tl H I LeptosPermum lllHHlts-ll (, uJ (, (, Oleaceae llllllNltl H rHll P H IH H Casuarina ts ts IH N

ll N N Banksia Þl r¡11 N N IN N Þ td H F{ lHll F Hakea trl tlll H H Its N H N) N N) IN N Xanthorrhoea H

H H t, (, Leguminosae tlll I I l(, l¡J t,

H H IH H H ts Ramnaceae

H H IH H rlts H Dilleniaceae H H H lltl H IH Epacridaceae tlll N N N) H Goodeniaceae llll H ts H tlP

HtsIFI IFIHIHI rF.l Rosaceae

tllllll rlllts Scrophulariaceae

tPllPrlFll Umbelliferae H NN) tl N lN)f¡l It\)IIN N Filicales (, l\, N 19 lHl I Sedge H IH HIIHI rÞ Plants (, tÞ (¡, N Aquatic tlll N (, N NINIINI anglosPerms

N t\) N N N Algae N N NNI N HNN) (¡) (, (, UJ (/) (, (, sPecies rd-¡H Total HHNoIIJTToPHHN(¡

TZ E u)oØØ(nØE'dlogãvEEE l-r. g Þ är r s 8 Ë g* F & t R Ë o HP'rtoáEõibrii+H'Ér¡ Ff cI'J|'Täõ,FJþ35OOJHO rJ, .rro.':õ-clFÞ' F' o H. il g b' t:- s rt kúöo,lo-ÞFlor. ä R; i o rD-ói-t-lo:P,, a iiXHoïur ooo |.<)fs o o rfÞ P. J É o 0,

H N H t\) H N tl N F N NNHN) N, Vegetation form

ts H H P H H o\ 6t or o\ or ÈÈrOì oì Eucalyptus N N N (¡ (¡ N P N ¡Þ H N H llrl I Acacia È ¡Þ t¡ H H H P rlll I LepÈospermum (, t, (¡, (, HllNltlNlN INN) ! oleaceae

H P H IH H H P Casuarina

N I F] N lN) N N tll Banksia Þ u, r{ tttlll H ts H I Hakea t4 H N H tttlll N N I xanthorrhoea H N H H H (rll!l , l I I Leguminosae UJ (, (, (, H H ts I Ramnaceae

rllllt H H P I oilleniaceae P H H I Epacridaceae N l\) N H H H I Goodeniaceae

H l' H I H I I P I I I I F I I Rosaceae tll rl I scrophulariaceae

ttlflllHl IHHH I umbelliferae H NININI NIN tlll (/) I riticales NN N N rl IHIH H Sedge (, (¡, (¡) L Plants Aquatic NIN lllN , H anglosPerms

H H NJ N N Nt\)N tN .:" Algae N(, I NÞ (, N) (, (, È (¡J (, (^) È È

l¡J or Total species HUJ ot(¡)tsuHPÈ or

zz ( áãFl'l 3 u)uEb{a!'9 tr H. OJ ¡r.0rOp o rJr+oÞ 5 H[HHËffÉ r Ê HordÉ o ;,-t-.ui(act(qH o ú 't-.HOþ Fl 'AiDFl-Öi-.5ltooo- Þ o, Ht I'J p, H frl ã0rprp, vt Ø-iñ.ØoH tt pr P. o a 9E'ãc' cf H rf o ik5o¡õPr i [E# rf o o 8",ä)É4. HI ts;ç SHogã H o (n o K 5 rf Þ P. Þ Ê o o,

{N\¡È@Or()r(¡ N H H ts Vegetation form H ol ol Eucalyptus OtOrlO\Oì\O\O\O N N lllll H È (, Acacia N H H P H H I ts LeptosPermum (¡ N N (, (,

N N Oleaceae

N)tlNl N H P ts Casuarina

tllNl N N N Banksia Þ'l tl[d rd rl}.lHll H H Hakea N H ¡\' HHIHIHIIJ (, Xanthorrhoea tttlllll (, P Leguminosae

llllll H H Ramnaceae

H H H Dilleniaceae

tttlll,l H N Epacridaceae H ts Goodeniaceae

HHlllHtl H Rosaceae Scrophulariaceae

ttlllll tlll Umbelliferae

NNNNNIII N N) N Filicales NNNNNN N N a- r I I Sedge i-(r(r(/)(r(rÈ | | Þ (, Plants

N Aquatic N N N Nfll anglosPerms

N) N) N(, N tlfN P Algae t, (t N (¡, Total sPecies HHH(, N) N @ H oì tz 24

TABLE V

o Ê +JË o ..{ o H o '.{ l+{ x ..{ c) o brúr{O+J oo É Ð o ut{ o Ê lrj Jlr¡i'ldt Ét c) o rúo (d Ê H o go3i.r.9 +Jo É É c) c) o ?.c..r{J'.{ v, r{ |r ø Þ çà S.oôq_to .Fl o o) rú o X 6'á0ú4Ø o Ê{ A{ É o Í4 Maior SiÈes 4 I I 43t24 ParaI¡Ia PV'¡ 356L1,2

Hindmarsh 1,315 4,5 2,4 FalIs HF 334L5 2 2 I

Eric Bonython 4L4 Reserve EB 354L4 3r1 1 6 I 5 Botanic Park BO 4 6 I r 1,2 4 L 4 2 2 Torrens Lake TO L2235 1t

Minor Sites I I 3 Aldgate Creek AC r255L4 321 4 I 3 Alnut A 1 I 5 4 1 2 r14 I I 5 2 4 BethanY BT t 2 2 5 5 3 3

Blewitt 6 I 4 Springs Bs 4 4 4 3 3 L12 5 3 1 Brownhill I 13 Creek BH 7L3334 321 4 2 I 5 I 3r4 BuIIrs Creek BU L 2 4 4 2,3 5

5 3 I 4 2 4 Carey's GullY cG 345713

Chamber ¡ s 3 2 I 5'6 4 3,2 Creek cHL2351-34 215 42 Clarendon CLII4423 4 ¿ CIeIand t4 Reserve L458L24L15

Creek Deep 5 L4 (Soutt¡- Coast) DE r444r432r

Deep Creek 5 I 3 (Adefaide DP925613 311 uills) 4 1 DuckPonds DU 4 2 I 5 3 4 2 2 3 3 5 t 2 I 3 4r5 2 Finnis River FN I I I L 4r3 5 I I 4 2 3 3 Fishers FI 5 4 5 7 1 2 25

KEY TO TABLES V TO VII each site' syMBoL. This shows the symbol used in Maps I and II to represent are symbols for particular sites in the south East of south Australia given in the text. regularly for a period of NUMBER oF VISITS. AII major sites were visited

one and a half Years or more' or ottrer develop- CONSERVATION. Site surrounded by:- agricultr:ral land ment,(I);nativebushinterspersedwithareasofagriculturalorother development,Q)¡narrowbandofnativebush,remainderinagricultural use, Q); native bush, (4) . (2) 24 28, (3); 28 31'Glt RATNFALL (inches) - L6-20 , (I); 20 - 24, ¡ - - 3t - 3e, !!)t 39+ (6). (2) (3); ATTITUDE (fCCt). O - 25O, (1) ¡ 25o - 5oO, ¡ 5OO - 75O, (5); I25o I5oo (6) ; 15oo - 1750, 0) t 75O - 1OOO, Ø) ¡ looo - I25o, - L75O+ , (8).

(3); ¡ sgIIJs. Acid toam, (1) ; neutral loam, (2) ¡ alkaline loam, sand, G) clay, (5). water to col'lecting site:- DT STANCE FROM SOURCE. Distance from source of not at source (i.e- at springs close enough that water temperature has yet risen to ambient), (1); up to 5m'' (2) ¡ 5 - Soom'' '(3); (5) 5oom. - 5Km- , lE t greater than 5Km'' ' Flow as in Hindmarsh VOLUME. (Volume of water flow) ' Lakes or dams' (1) ' up to 2m' River, above Hindmarsh Falls: bed irregular, 5 - 7m. wide, to stream- deep: subject to winter flooding, occasionally quite damaging side vegetation, Q). FIow as ín Tunkalilla Creek at Eric Bonython also subject Reserve: bed steep-sided, I - 2m. wide, about Im. deep; wide, fed by strong spring to flooding, t1). stream bed less than lm. greatest or seepage, (4). Springs parÈIy dependent on Seasonal rain, flowatrickle,d,ampeningonlytheim¡necliatestreambed,(5)-. 26

springs or seepages where PERMANENCY (Permanency of water flow) ' usually area affected by water waÈer readily escapes from ttre source area; Iargelyunalteredbychangesinseasonalrr:n-off,(1).Areaaffectedby waterreducedduringsunmermonths,Ø.IÁ¡aterflowstopsduringsuinmer monthsi streans which reduce to poors, and springs scarcely keeping pace with evaPoration, (3)' (1); neutral (2) i pll. pH of water at collecting site:- acid' '

a1kaline, (3) - BASEMENT.Ataparticularsitewaterusuallyflowsovermorethanoneof the following base materials:- bare rocks' (1); sand' Q) ¡ sandy plant soil, (Ð; soil rich in decaying matter or covered with decaying debris,(4);rocksandrootedplantsormosses'fÞ;rootsandrooted plants' (6). 0r semi-terrestrial larvae:- covER. Ground cover avaitable for terrestrial plants' or several complete dense covering of longish grass or other inchesoforganicdebris,(1);grassshortordebrisformingathinor incompletelayer,(2);baregroundwithsparsegrassesandoccasional accumulations of fatlen leaves' f 3-) ' E}eosURE.Exposureofthesitetosrrnandwind:-lowsparseçJrasses,no Iargetr:eesrstreambanksshallow'lI);someclosewatersidevegetation' bankshigher'surroundingareawithouthighvegetation,\2),waterside open bush, (3); vegetation not dense, surrounding area covered by watersidevegetationd.ense,oftenroofingthestream;valleysheltered butundergrowthpenetrable,!l};streamorseepageareacompletely (5) overgrorttn, bush almost impenetrable' ' 27

TABLE Vf

É +JÉan o .r{ O l{ ¡1, ..{ tt{ c) o .F{ Ð oo É .tJ a) tßlFlO og o Ë t{ c.) o É c) a Ju{i¡rÚt rúO rlt Ê H u o 1Jo â Ë o c) o g si.r 2 o Þ 01 UI ti EÉ.'!Ð''{ ..t -lo q) (6 o x å 2I s ã 3 o A g{ m o f4 Sites (continued)

GIen View cv 1 1 2 6 3,4 3 4 3 3 22 GreenhilI 2 4 2 Quarries GA I I 34 1r3 3 42 3 4 2'L Hackham HA I 2 32 I1 f 213

2 2 Happy ValleY HV 3 I 33 L,2 4 4 3 2 4 415 4 3 Hea1ey's HE 4 t 45 L2 5 I I

2 3 4 Humbug Scn:b HB I 4 2,3 6 Lr2 I ,2 4 3 I 4

Inman Valley IV 10 I 42 1,5 5 2 2 I 3 3 2

2 a Keynes HilI KH 2 1 26 24 4 3 5

3 2 3 Kuitpo KU 1 4 44 15 2 2 1

Lower Hermitage LH I 2 35 I 5 2 I 216 I f 2 3

2 2 5 2 Marne MN 2 2 25 L,2 5 2 3 2 5 I Moculta 4 3 I6 3 3 4 2 3

3 4 5 4,3 3 Morialta MO I 4 46 1-3 4 3 2 I f 4 2 4 Nietsker s NI 4 2 25 3'4 3 5 3 I

3 1 2 North Para NP 1 I 23 3 5 2 2 3 I 3 Orlando OR I 2 24 3 4 4 2

1 2 I I I 4 I 4 I a I 4 54 1 3 4 2 Reynella RY 3 I 32 3 4 4 3

1 3 3r4 3 Robertson t s RB 3 2 45 1 3 4 2

Saunderts 2 3 Creek 1 4 I 2 L14 5 3 3 2 4 2 Scott's Creek sc 10 3 3 5 4 I 5 3 I 2

3 rr3 2 2 Second ValIeY SV 5 1 3 2 2rI t 3 I

2 lr3 1 2 3 Sixth Creek SX 2 r12 4 4 I 4 2 I L,2 4 2 Springton SP 2L 2 6 2 I I I 28

TABLE VTI

É o o ..{+JÉ t{ O tl{ x .FIU) 'r{ o U JJ É Ð o SrdF{C) ooOH o' É t{ É q , Jt+¡krúA Ét o rft do É d lÍ t{ lJO a É 0) o o 8ösi.È.g Él ${ o 9{ ut X E.È.JÐ'.'{ .Fl o o) rd o å s I Æ Ë 3 e A È m o Í4 Minor Sites (continued) 6 I4 Sturt River SR 2245t4 221 1 3 5 2 Tanunda Creek TD 4L 2 7 L12 3 4 3 6 2'5 3 Tooperang TP t3 3 2 1 4 4 3 ttaterfall r,4 4,2 3 Gully wr' 2446L4321 312 Williamstown fÀIT 11331443

South East of S.A. 2 I 3 Lake Albert L21 1 4 1 1 I 2 3 Small claypans t42 I 4 I I 2 3 3 2 I IJarge claypans I 2 2 I 4 I I 2 Lr4 I Crater Lakes t I 4 2 4 1 I I 5 I 3 Large springs I 2 3 t 4 1 2 I 6 I 2 Marshes I 2 4 1 4 I I 3 3 2 2 Drains I 2 4 1 4 3 5 3 2 3 Small springs I I 4 I 4 I 4 2 5 29

4. BIOLOGY OF INDIVIDUAL SPECIES

In SECTI9N III descriptions and, where necessary, redescriptions are provided of aII Aphroteniiae, Heptagyini and orthocladiinae ncw known from south Australia, as well as some from outside this state' Below' and biological details are provided, discovered during the investigations collections outlined above' SouthAustralianrepresentativesofPodonominaearementioned work' briefly because information about them relates to Brundin's lfhere sufficient information is available about a species this has been presented under three headings:- A. HabitsandHabitat-referstothehabitsofallthreelife stagesandthet}T)esofplacesinwhichtheyareusuallyfourrd. data B.L ife Cvc1e - for some species it has been possible to use from laboratory rearing to make an estimate of the minimum time required to complete the whole or part of the life cycle' Inothercasestheyeartyvariationinfieldcaptureofspecimens (e'g' indicates that the species is almost certainly uni-voltine Graph 12) N. mucronatus Graph lO) or bi-voltine (e.g' A' digitatus' ' for Howeveritisuncertainwhetheradistributionsuchasthatshown species or ParalimnoPhYes pullulus (Graph 14) indicates a uni-voltine fluctuations one which is on the wing throughout the year. In addition, inthenrr¡nbersofaspeciesaÈasinglesitecansometimesbetraced (e'g' directly to the influence of environmental factors nukiefierieLla

insol ida) . which might C. Taxonomic usefulness - any biological information provj-de an insighÈ into the function of particular morphological discussed structures and thus be of value in phyrogenetic anarysis is here. Forallspeciesrthenr¡nberofsitesfromwhichtheyhavebeen reported' l-n reported and range of months over which they have been vrrr. The south Austraria and in other states, is recorded in Table S ON D No. sites TABLE VIII J FMAMJJA I Aphrotenia australiensis 2 ParahePtaqyia plumata I Cardiocladius austratiensis .2 Stictocl-adius uniserialis 2 multiserialis 7 Eukiefferiella insolida 4 cladius bn:nnen 7 oluriserialis Pæatrichocladius 2 tl bifinestrus 4 Crico topus albitarsus 2l cinctus 4 I phaeosomata 1l il annuliventris I il hirtellus I M icrocritopus clavulus 2 Nasuticladius diplostylus 1 il fuscus 9 I eurynotus 5 I punctus 2 ll parapunctus 5 ll carinatus I ll sal-teri 3 tl brevipennis 3 It healensis 5 tl mucronatus 3 tt gloveri 3 tl longipennis I I duckhousi 15 Austrocladius teriugus 8 tl numerosus 2 tl kvpensis 1 il woodi 6 lt trichiatus L4 il diqitalis o N D JFM A M J J A S D No. M J JA S ON sites TABLE IX JFMA t I I Iyolobus 1 Lvmnophves tubercus 2 " pelurçris 22 Paralirnnophyes pu1lul-us 13 ll a-]b,i basis 3 Sparsicfadius bryonis 6 tl telmatus 4 t( pofysetosis 6 ll paucisetosis I I gramrnl-s 9 Nemoricladius diversus 4 tt lnversus 6 I lobitorus 6 Kiefferophye s invenustulus 9 il phallithrix 4 I arti-othrix 4 il semithrix 2 t! cuspistyius 3 ll Iobifer I Parametriocnemus ornaticornis 7 Gynnone triocnernus wilsoni 5 !l impectinatus 3 I parapectinatus 4 Rhinocladius tonnoiri 5 Smittia aterr lma 2 retracta 20 Parakie fferiella varigatus t Pseudosmittia' argilla I rt scutella 4 Doloplastus crassistylus 4 It curtatus 6 il culcitellus 2 n sensillus S o N D JF M A MJ JA 30

Iarge scale distribution of the species, and where relevant' the distribution of closely related species is also recorded' Abbreviations used for the states of Australia are Australian Capital Territory, ACT;

New South wales, NSW; South Australia, SAi Tasmania, TAS; victoria, VICt lrlestern Australia, VüA; Northern Territory, NT'

Remarks on the Extraction of Larvae.

Many hours vrere spent examining sr:bstrate material by microscope and pulling decaying material apart with forceps, but immatures of three of the most interesting genera, Nasuticladius Austrocladius and Kiefferophyes, eluded discovery. However, at the end of my invesÈigations' while searching through moist but not saturated debris, and this time' pulling apart all stems, I found a single chironomid larva insid'e a small piece of tough, relatively undecayed Xanthorrhoea stem. unfortunately it before was probably damaged from this extraction and died of fungus attack developing further. It is possible that larvae of all three of the above genera are to be found in this kind of place'

SUBFAMILY PODONOMINAE

TRIBE PODONOMTNI

Genus PODONOMUS PhiliPPi

Podonomus ,derwentsis Brur¡din Adults were reared from moss, taken during the wettest months

from Èhe splash zone of rocks at positions Il, lla and llb at Hindmarsh Falls. Similar niches are occupied by many other members of this çfenus'

Genus PAROCHLUS Enderlein

Parochlus bassianus Brundin and, P. r!e¡! Brundin Larvaeandpupaofbothspecieswerefoundallyearroundat Botanic Gardens, among debris of twigs, leaves and broken Blechnum stems here taken from under dense stands of tea-tree and Blechnum' Temperatures slow enough reached I20c during late summer and the water flow is always of bassia-qus that suspended material flocculates around debris' Adults 31

round' at Botanic Gardens' h7ere taken in uniformly high numbers aII year EricBonythonReserveandParawa.P.riekiwastakenonlyatBotanic Both species Gardens, with highest numbers in the months February-April' were observed swarming at' dusk' Theim¡Ûaturesofthesespeciesarethusto]-erantofhigherwater although Brundin temperatures than others of this genus yet reported and, plainly not lovers reports them from stream net collections, in sA they are ofrapidlymovingwaters.Bothspeciesappeartohavecolonizedthistype perhaps' of habitat more successfully than any orthocladiinae except'

Parametr iocnemus ornaticornis (rieffer) .

Genus PODONOMOPSIS Brundin

PodonomoPsis evansi Brundin Adults were taken at Nietskers, Moculta' Keynes HiIl and the summer Hackham sites at which water flow all but ceases during by sparse months and the remaining pools are protected from the sun only vegetation.Thisspeciesthusappearstobeabletotoleratewarm'still vraters and is unique in what Brundin has described' as a "strictly

rheobiontic genus" -

Fromtheabovediscussionsitseemsreasonabletosuggestthat in some whire stream nets may be an efficient way of capturing immatures that a environments, the data from them should not be used as evidence species is necessarily rheophile. It is likely that many captures made bythesenetsaretheresultofspecimensbeingwashedoutoftheir preferred habitats by rains or changes in water level' 32

SI'B-FAMILY APHROTENI INAE

Genus APHROTENIA Brundin Aphrotenia austral.ien sis sp.nov. Adult males were netted during september in Eric Bonython Reserve at the perimeter of a large dense stand of Gleichenia circinata. Following Brundin's discoveriesr (1966, p. 331) debris from upstream rock faces' as well as other suïfaces was examined for immatures, but without success' Distribution.SA;otherspeciesinSouthAfrica.

SUB-FAMII,Y DIAMES TNAE

TRIBE HEPTAGYINI

Genus PARAHEPTAGYIA Brundin

Paraheptaqyia Plumata sP.nov- Adults were taken twice at light, in september and November, at Botanic Gardens. A third probable record is of a mutilated male from a stream net sample at Hindmarsh FaIIs in July. The present populations are therefore probably small, but may be remnants of a wider ranging population. Distribution. sA; other species from ACT' NSVü' TAS, and south America'

SUB-FAMILY ORTHOCLADIINAE

Genus CARD]OCLADIUS Kieffer Cardiocladius australiensis Freeman

Habits and Habitat. Known larvae of Cardiocladius species prey on Simulium larvae, and Tonnoir (Lg22) remarked of this species "larvae predacious on Simulium larvae" ' Adu It australiensis have been taken in other states, as well as at Hindmarsh Falls in November-December' a j'n However time when simulium larvae are still developingt aE least sA' larvae on a search for australiensis larvae among early instar Simulium view of rocks and water plants at Hindmarsh Falls proved fruitless' In the large number of simul-ium larvae present it is surprising that only 33 a single adult was netted' it is Taxonomi c usefulness. For c. australiensis larvae to survive the probably necessary for the gravid female to deposit eggs on or near with rov¡ water surface. rt is thus interesting that adults have a wing low speeds Aspect Ratio, which would allow greater manoeuverability at in a high humidity (see SECTToN rv) - The female of australiensis in conunon with other Cardiocladius species, has gonocoxite IX extended posteriorly' perhaps to act as a positional guide during oviposition in somewhat difficult circumstances' Distribution. SA, NSW, ACT

Genus STICTOCLADIUS

Stictoc Iadius uniserialis Freeman and S . multiserialis Freeman uniserialis into a Habits and Habitat. The emergence of a single female trapoverposition(ii)atParawaindicatesthat,asinEuropeanSpecles' Adults have been the immatures live in still or slowly-frowing water. takenamongdensewatersidevegetationatEricBonythonReserveandat parawa (Habitat G.). LifeCycle.onlyafewspecimenswereobtaínedinSA,buttakenoverits but midsummer and known Australian range uniserialis is on the wing in aII multiserialis from April to october' reguire manoeuver- Taxonomac usefulness. Here again mating and egg-laying ability in an high humidity and Aspect Ratios are low' in multiserialis marginally lower. Líke Cardioc ladius australiensis, c multiserialis females have abdominarextensions.rnthiscase.howeverritisthetergiterather than gonocoxite which is modified' S' uniserialis is probalrly more wing in ecologically tolerant than mul-tiser-ialis. It is captured on the structures' most seasons and, has less special.ised wing and female tergal Distribution. AcT, NStr{, SA' TAS' vlc. other species in New ZeaLand, South America and EuroPe. 34

Genus EUKIEFERIELLA

Euki efferiella insolida (Skuse) Freeman

Habits and Habitat. As in rnost European species of Eukiefferiella on finding immatures of insolida are rheophile and depend for survival algae' submerged, well-aerated attachment sites such as filamentous no net captures Chara and other water plants. Thus, at Hindmarsh Falls, still be were made when filamentous alga was absent, but adults could at l2oc' reared from vigorously aeraÈed samples of Nostoc-Lycopodium kept Inaddítion,atEricBonythonReservewherewaterflowiss]-owerand in low little filamentous alga develops, insolida was captured only the bog nurnbers. E. insolida was absent from Botanic Gardens and from or poots of the south East, where suitable attachment sites are al¡sent poorly aerated. ThecaptureofadultsatsecondValleyandnearthelvlarne They cart river indicates that immatures tolerate alkaline conditions' Falls also withstand the battering of torrenÈial currents at Hindmarsh (positionlg)andtheabrasionofthesandybedsoftheMarneRiverand

Saunderts Creek. 8 weeks Life cycle (Graph 3). Laboratory rearings show that more tha¡r occurs aII year is needed for development at 12oC. Eclosion probably round. is exceptionally Taxonomrc Usefulness. The aspect ratio of female insolida water.s edge. Iow, again probably to allow Successful oviposition at the Distribution. sA, NSW. Other species in Europe, canada, scandinavia'

Genus SYNORTHOCLADIUS Thienemann

Svnorthocladius brunnenr sP. nov. material' Habits and Habitat. Adults have been reared from a variety of alloriginallysubmergedinslowlyflowingwater.Capturesofadults available were restricte¡l to sites where slowly flowing water is widely all year round- No swarming was observed' GRAPII 3 Er:kiefferiella insolida. Net capÈures at Hindmarsh Fa11s, l'97L

GR,APH 4 Synorthocladius brunnen. Net captures at Eric Bonython Reserve' I97O

GR.APH 5 Cricotopus annuliventris. otal captr:res , L97O

GRAPH 6 Cricotopus parbicinctus. Net captures at Eric Bonython Reserve, I97O

Horizontal Axis Months of the year

Vertical Axis Nr:mbers of specimens Èaken 25

t5 lo

o N D 0

20 4 I5

o D A M J J o

0 J F M A H J J o 35

Life CvcIe. Adults were netted irregularly, usually in low numbers' (see Graph 4). Taken over all sites they were on the wing in all months but December and January. It is probable that adults are on the wing in low numbers all Year round. Distribution. SA. Other species from North America and Europe'

Genus PARATRICHOCLADIUS

Paratrichocladi us p1uriserialis Freeman and P. bifenestrus sP.nov.

Habi ts and Habitat. Forsyth (I97f) reports larvae of pluriserialis in New zealand living a few cenÈimetres below the surface anìong Potamoqeton, Nymphacea and Salvinia species in shallow' stagnant or flowing water. In sA pluriserialis adults have been taken only where

reed beds or Bl echnum are associated with still or gently flowing water (including Mt. Gambier lakes, River Murray, inland springs). P. bífenestrus

was taken only at Hindmarsh Falls and Eric Bonython Reserve, places where flow rates are higher Èhan in the above: probably immatures of bifenestrus require higher oxygen concentrations than those of pluriselÞI:Ls. Both males and females were taken at light; no swarming \^/as observed' Although adults appeared to remain close to the reed beds even at dusk, a single capture from aerial plankton at 105m. above the ground at Keith shows they may readilY be carried aloft' Life cycle. Adults of both species are on the wing during most months and in contrast with Forsyth's observation, I detected no increase in

numbers of pluriserialis during spring'

TaxonomicUsefulness.FittkaureportsEuropeanspeciesofthisgenus as being restricted to cold, high altitude waters. The presence of the Iarger wing microtrichia in Australian species may indicate adaptation to a higher humiditY environment' SA; Distribution. P . pluriserialis SÀ, NSV'I, VTC' TAS'; P . bifenestrus other species NT, Europe, Nth. America. scaircinavia, south Africa'

South America. 36

Genus CRICOTOPUS van der VÙulP

Adults of C. albitarsis sp.nov., C. phaeosomatus sp.nov', and C. hirtellus were captured during the spring months and albitarsis was present during February-March also. However numbers taken were too low to make any generalizations.

Cr parbicingEus' sP.nov- alga in slow Hab its and Habitat. Larvae \¡tere found among filamentous currents and backwaters where they live in short tubes and feed from raufwuchs,. In still \^later in the laboratory they were observed crawling it' Adults suspended upside down from the surface film and feeding from were not observed swarming' of the Life cycle (Graph 6). Adults were captured during aII months year'butnumbersv7eremuchhigherduringseptemberandoctober,when take filamentous alga is present and flow rates are diminishing' Larvae for three or more than a month to complete development and pupae live four days at t5oc. larvae Taxonomr c Usefulness. In groups such as Dixidae and Culicidae which feed fron the surface film, have mouth parts with speciatly Since developed brushes which allow feeding currents to be set up' mouth parts parbicinctus larvae can also feed from the surface film their probable that must to some extent be adapÈed for Èhis. rt also seems other Cricotopus larvae use this feeding method in similar circumstances larval Consideration should be given to this before characteristics of genus' mouth parts are used as evidence of near relationship in this

Cricotopus annuliventr l-s sp.nov. collections Habits and líabitat- Adults emerged, with icinctus, from produced of filamentous algae; unfortunately no larvae reared separately annuliventris adults. 37

( 5) Life cvcte. Adults were netted irregularly at several sites Graph october' and. were taken in all months, with a maximum in sepÈember or

As a genus, the abundarrce o f Cricotopus in South Australia

appears to be affected most by the availability of filamentous algae; earlyinwinter,beforerockpoolshavefilledbutafterSummer when they accumuration of argae have been rvashed away is the only period

are a.bsent.

Genus MICROCRICOTOPUS Thienemann & Harnisch Microcricotopus clavulus sp.nov.

Habi tsand.Habitat.AdultswererearedfrornNostocandfromsr'rbmerged tree roots collected at Hindmarsh Falls and constantly aerated' stream net collections at Èhis site yielded clavulus pupal casts' The single specimen reported from south east of Morgan most probably originated' in waters associated with the River Murray- Thus the habitat range of this Australian species probably does not fall outside that of other with plants members of this genus, all of which are found in association in lakes and flowi¡rg streams' Life cycle. fnsufficient data are available to allow any estimate' Distribution. sA. other species found in most parts of the world'

Genus NASUTICLADIUS Freeman

The species fuscus, diplostylus, parapunctus' salteri in healensis qloveri, longipennis and duckhousi, newly descrilced the months sEcrroN rrr have been captured in very low numbers during to be indicated in Table VIII. Immatures of these species are probably and composition' forrnd among rotting vegetation of differing moisture levels GR,APH 7 Nasuticladius eurynotus. Total captures, Botanic Gardens, J-97O

GRAPH 8 Nasuticladius punctus. Total captures, Botalic Gardens, I97L

GR;APH 9 Nasuticladius carínatus. Total captures, Hindmarsh Falls, L97O

GRAPH 10 Nasuticladius mucronatus. Total captures, Botanic Gardens, L97I

GRAPH 11 Austrocladius kypensis . Totat captures, L97I¡ captures at Paratùa, shaded.

GRAPH 12 Austrocladius diqitatus. Total captures, Torrens Lake, 1970

Horizontal Axis Months of the year Vertical Axis - Numbers of specimens taken 20 25 7 10 20 l5 l5 IO

5 TO

5 0 M A M

o À M J J IO 11

I 0 45 Ii 40

9 30 30

25 25

20 20 15 I5

IO l0

5 5

J o 0 A u J J A 38 Nasuticladius eurynotusr sp.nov-

Habits and Habitat. Adults v¡ere reared from a muddy debris of Blechnum and Leptospermum stems ahd leaves collected from Botanic Gardens and kept at ;^2o1. At Eric Bonython Reserve and Botanic Gardens they were seen swarming over markers and were al-so taken at light. The wide varj-ety of localities from which this species has been taken suggests that inunatures are tolerant of a considerable rançte of temperatures'

Life Cyc1e. ( Graph 7) Adults vrere netted from April to August at Eric

Bonython Reserve but only from April to June at Botanic Gardens. The species may be univoltine but Table VIII shows captures from all sites collectively, and considering the fact that captures may be spread over five months at a single locality it is probable that adults are on the wing all the year but that they are usually present in numbers too iow to be registered.

DistribuÈion. SA' ACT, NSI^I.

Nasuticl-adius punctus_r sp.nov.

Habits and Habitat. Probably less toleranÈ than salteri since it has been recorded only from the Southern Mt. Lofty Ranges' Life Cycle. (Graph 8) Captures at Eric Bonython Reserve and Parawa are restricted to a period of 2 months but Table VIII again shows total captures spread over a much longer period'

Nasuticladius carinatus sp. nov.

Habits and Habitat. Total- and stratified samples of leaf and plant debris were taken from below a slow hillside seepage at Hindmarsh Falls (positions 4,5). Adults emerged from the total samples and from the separated middle strata, where the immatures \^lere protected' from dessication but not subject t-o irnmersion itl seeping water. The immatures must thus be considered semi-terrestrial'' During May and June innumerable smalt svrarms were seen that 39 v¡ere strongly attracted to markers' of being Life Cvcle. ( Graph 9) This species also gives the impression univo.ltine but total captures (Table VIII) suggest a continuous low adult population.

Nasuticladius brevipenlis, sP.nov.

Habits and Habítat. The presence of this species at Torrens Lake as wel-l as Eric Bonython Reserve and Botanic Gardens suggests the immatures live in poorly aerated water or mud, among plant stems'

Nasut icladius mucronatus sp.nov.

Ha-l¡its and Habitat. Adults vtere reared from a section of permanently

sodden grassy bank taken from Botanic Gardens. Adults were taken at

light and on oilY PaPer.

Life Cvcle. Graph IO

Genus AUSTROCLADIUS Freeman

The new sPecies woodi , tubercus hindmar shi and lyolobus were VIII' taken in very low nunÙcers during the months indicated in Table the Live, unidentified femaIe Nasuticladius were observed closely under in microscope while being offered a variety of native flowers, honey

water and water. They accepted only water'

Austrocladius terjugus Freeman markers' ÏIab its and Habitat. Adults s\{arm during mid morninçt over the months Life Cvcle. Adults are on the wing in low numbers d'uring Ju:e to SePtember-

Distribution. SA, NSI^I , ACT ' VIC ' 40 Austrocladius numerosus Freeman

Habits and Habitat. Adults v¡ere taken in emergence traps on the drier part of the hitlside aÈ Para\^¡a (position (vi) ) and over a pool at Scottrs

Creek which was lined with dead leaves. One adult was netted at Salt creek where the only water supply is quite brackish. Immatures must thus be tolerant of a wide range of conditions. Numerous sl¡ilarms \dere seen' always in shaded P1aces. Life Cyc1e. Adults \¡/eïe taken between August and October in SA and have been recorded in May and December in other states. They are probably on the wing in low numbers, all year round'' Distribution. SA, NSW, ACT' VIC, TAS'

Austrocladius kypensis' sp.nov.

Habits and Habitat. Most specimens were taken from reed beds. Life cycle. (Graph 11) Captures at Parawa and Eric Bonython Reserve were restricted to May, June and July. However a single specimen taken in Botanic Gardens in April suggests that the species is on the wing all year round, although it remains possible that it is univoltine at each site.

Austrocladius d r-g itatus, sp. nov. Habits and Habitat. During JuIy, August and september adults were reared from hillside divots taken at Parawa and kept as in 2.321- Captures at

Tilleyrs Swamp and Canunda, where in general only moist soil is available in January and February indicate that some stages of the life cycle must be tolerant of low moisture levels. swarming was observed on numerous occasions. Life cvcle. Adults were taken in large numbers between April and september. Graphs of captures at Parawa and Eric Bonython Reserve show Torrens a bimodality símilar to that shown in the graph of captures at Lake(Graph12).Itispossiblethattherearereallytwospecies 4I involved here, but more likely that under more favourable conditions during the wetter months, the autumn generation completes its development quickly. The fact that Torrens Lake was drained during June without apparently affecting the distribution of catches supports the proposal that immatures tolerate semi-terrestrial moisture levels.

Austroclad.ius trichiatus Freeman

Habits and Habitat. Adults emerged into a trap placed over saturated organic debris in the small clearing at Botanic Gardens. Life cvcle. Adults \^rere taken in low numbers from April until October with a peak in late April. The species is probably always on the wing in

low nurnbers. Distribution. SA' VIC.

Genus LIMNOPHYES Eaton

European and North American species of this genus are reported

from ponds, moist earth and the beds of streams. The Australian species

seem to occupy similar Places'

Limnophves tubercus sp. nov. Alltwelvespecimensv¡eretakenatHindmarshFalls,soitis probalrle that thj-s species is rheophile'

Lirnnophyes Pe1urgis, sP.nov. the banks of fine tlab it and Habitat. Immatures live anong reed beds in was taken' silty mud which dominate both sites from which the species Larvae are excellent crawlers but poor swintners' on Life Cvcle A sample of mud taken from the Happy Valley site 2g.vi.Lg6gyieldedlargenumbersofadults3-4monthslaterinoctober be at and November. The time for completion of one cycle must therefore 42 least three months. Catches v¡ere too small to altow the distribution of generations throughout the year to be determined''

It is significant that pelurgis has a much higher aspect ratio than tubercus. Muddy areas suiÈable for pelurgis larvae are likely to vary in distribution from season to season and effective utilization of these will require dispersive flight, for which the high aspect ratio wing will be advantaçJeous. fn contrast larvae of tubercus appear to inhabit streams which do not alter in posiÈion from season to season. In the larva of pelurgis one of the ventral mandibular setae is developed into a fine flagellate brush. This structure has not previously been recorded in Limnophyes and is probably an adaptation either to moving or obtaining food in a muddy environment.

Genus PARALIMNOPHYES Brundin

The Australian species tend to be more limnic than their

European counterparts, which occur in ponds, dried out ditches and small moorland Pools. P- pullulus is al¡undant at sites where albibasis is absent but it also occurs where albibasis is numerous.

Paralimnophyes albibasis (Freeman)

Habit and Habitat. Inmatures are free-Iiving among well oxygenated, moist, decaying organic matter in a wide vari.ety of hal¡itats (e'g' Habitat D, Parawa; midstream leaf accumulations, Hindmarsh ralls') ' Larvae can move very rapidly through detritus but swim poorly' They feed on taufwuchst. Adults were taken at second Valley, indicating a tolerance of hard waters, so the absence from collections made in the south-east of the state may reflect the lack of essentia-I ions i¡r the highly leached

and coastal soils. GRAPH I3a Pa¡a@phyes_ albibasis. Total cap es, Parawa, L97O - L97L

GRAPH 13b Para@phyes- albilcasÍs. Total captures, Botanic Gardens, L97O - 1971

GRAPH 14 Paralimnophyes pullulus. Total captures, Torrens Lake, 1970

Horizontal Axis Months of the year

Vertical Axis Nr¡mbers of specimens taken 13. 2

35 14

20 l5

0 M À 43

Adults do not swarm but large numbers of males and females v¡ere often seen to fì-y slowly and. randomly in sheltered regions when the air was still. Both sexes have been taken at light and on oily paper.

Numerous dead females were collected in stream nets at Hindmarsh Falls which suggests that eggs are laid in, or very close to, running water. Life Cycle. (Graph 12) Adults were taken all year round, but in smaller nuÍibers during December and January. Rearing of larvae indícates a minimunì Iarval life of two weeks. The pupal stage lasts 3-4 days' Dístribution. Widespread in the wetter regions of NSW, VIC, TAS, SA'

Paralimnophye s pullulus (Skuse)

Habit and Habitat. Immatures are free-living in mats of Cladophora and.

Chara that occur in sandy beds of sluggish creeks, in moorland pools and in lakes. Swarming v¡as not observed but large numbers of both sexes \¡¡ere captured at Brown's Lake (a lake inside a volcanic cone, Mt. Gambier) flying

above short Cyperacea. Both sexes were taken at 1i9ht. Life Cvcle. (craph 13) The pupal stage takes 2-3 days- Adults have been captured at all times of the year but there is insufficient date to disclose any pattern of emergence.

Distribution. SA' VIA, NSIrI and VIC-

Genus SPARSÍCLADIUS gen.nov.

Compared with Paralimnophyes, Sparsicladius has a restricted distribution which probably results from the smaller range of ha-bitats occupied by larvae.

Sparsicladius bryonis, sp.nov.

Habits and Habitat. Immatures live where water seeps through a substrate covered with short grasses and moss. Such seePage areas were larger in Ig7L, due to greater rainfall, and this probably explains the large catches

in that year, especially from the híllside (Habitat A) at paratva. 44

swarms are small and move rapidly backwards and forwards within a metre from the ground. Life. Cvcle. Adults are in the wing in uniformly tow numbers throughout the year. Distribution. SA' ACT.

Sparsicladius tel-matus sp.nov.

Habits and Habitat. Immatures seem to prefer saturated ground where lhe

\^rater is still . Adults l^lere never taken in traps over seeping water and

none h¡ere taken at Hindmarsh Falls, but numerous adults emerged from

the compacted mudbed (Habitat B) at Parawa, during March and April and

hundreds were Èaken over still bog pools at canunda in the south-east of the state.

swarms are similar to those of bryonis and the two species are often captured together. Both sexes were taken at light. Life Cvcle. Adu]ts are on the wing in uniformly low numbers in all

months but JanuarY.

Distribution. SA.

sparsicladius paucisetosis, sP.nov.

Habits and Habitat. Adults \¡/ere reared from grassy divots taken at parawa duri¡g August and September: immatures probably prefer cond'itions similar to telmatus. Life cycle. AIl records are from the latter half of the year: thís

may be a univoltine species with the major emergence in August or

September.

Sparsicladius polysetosis, sP.nov.

Habits and Habitat, Probably similar to paucisetosis since it was tal

in emergence traps at Parawa (Habitat A) ' Life cycle. Adutts were taken in autumn as well as spring: they may be 45 on the wing throughout the Year.

Genus NEMORICLADIUS çJen.nov. N. diversus, sP.nov. r N. inversus, sP'nov', N' lobitorus sp. nov.

Ha-bits ancl Habitat. Rearing and emergence records suggest that these species occupy slightly different but closely similar habiÈats. N. díversus has been reared from moist leaf litter and from ground moss, taken from weII shaded sites (Parawa, position 15; Hindmarsh Fall-s' positions 1I, 11a, 11b), from moist debris found among Xanthorrhoea stems and from mushrooms (ACT). Emergence traP records confirm that the immatures are restricted to shaded regions of ground litter, while the only swarms observed were close among vegetation' N. inversus was reared from water-soaked divots of moss and grass taken from the open hillside at Parawa. lvhen swarms of N. inversuj; were observed over this hilrside, N. diversus was swarming among shaded vegetation. The absence of lobitorus from emergence traps suggests it may prefer drier or better drained conditions, and the fact that adults were captured at Hind.marsh Falls at site I, on a number of occasions when it was absent from ottrer sites, tends to confirm this' from March Èo Life Cyc1e. N. diversus and N. Iobitorus are on the wing september, while inversus has been recorded from -tr4ay to september' All cases of three may occur year round, but this seems less likely in the inversus and lobitorus than for d,iversus (Graphs 15 & 16) .

GenuS KIEFFEROPHYES FTCEMAN Kief ferophyes invenustulus (Skuse)

Habits and Ha-bitat. Adu1ts emerged into traps over the hillside (Habitat A) at Parawa, and were reared from divots kept as described in and orre 2.32L. On several OCcaSiOns sr^IaÏms were observed about midday gusty wind' sv¡arm was not deterred by heavy rain interspersed with GRAPH 15 Nemoricladius diversus. Total captures at Botanic Gardens (BO) r Eric Bonython

Reserve (ne¡, Hindmarsh Falls (HF) and. Parawa (ew¡, 1970.

GRAPH 16 Nemoricladius lobitorus. Tota1 captures, Botanic Gardens, I97O.

GRAPH 17 Kiefferophyes phallithrix. Total captures, Hindmarsh FalÌs I L97L

GRÀPH 18 ,, Parametriocnemus ornaticornis. Captr:res at Parawa at light (shaded) and by netting

(black) , 1970 - L97L.

Horizontal Axis Months of Èhe year

Vertical Axis Nurnbers of specimens taken fleo lee Elxr Øew 15

15 16 17 IJ

20 18 15 l0

o J F H ll J A o D ^ 46

LÍfe Cycle. Adults were taken in all months except April and May with a peak in July and August; they are probably on the wing all year round' Distribution. SA' VIC, NSW' TAS-

Kiefferophyes phallithrix' sp. nov.

Habits and Habitat. Swarms were observed on a number of occasions, and unlike mosÈ orthoclad swarms are very compact: the darting of individuals is unusually rapid, and only a short distance is traversed between each change of direction. Life Cvcle. Adults were taken from late May to late November but were more numerous in the later months (craph 17) -

Distribution. SA. No deductions can be made about the remaining newly described species of this genus, semisetosis artiosetosis and cuspistylus or about lobifer Freeman, since captures were so few-

Genus PARAIVIETRIOCNEMUS Goetghebuer

Parametriocnemus ornaticornis (Kieffer)

Habits and l{abitat. Larvae are bright green and are readily found close to the surface among boggy accumulationsof decaying platrt material (Parawa sites E (b),February to Aprit, Botanic Park site). They build 'fluffy' tunnel-s (which are easily destroyed) by gluing together pieces of suspended. detritus, sometimes using algae or decaying plant fibres as a framework. The tunnel is about Ib times the length of the larva and before pupation the larva builds a wall across it about 3/4 of. the distance from one end, and attaches itself to the wa]l. The cast larval skin remains attached to the walI and the pupa lives in the rest of the tunnel until it comes to the surface to eclode.

Emergence trap records show that P. ornaticornis ceases to emerge from an area whj-ch has become too deeply covered by water' 47

Life Cvcle. Adults are on the wing all year rou¡rd' at Parawa and Eric indicate Bonython Reserve. seasonal absences from other sites probably a shift in the population (Graph 18) ' Distribution. SA, NSW, ACT' WA. Other species from South Africa' North America and EuroPe.

Genus GYMNOI'IETRIOCIIEMUS Goetghebuer

G\¡mnome triocnemus wilsoni Freeman Parawa in emergence Habits and Habitat. Numerous adults were taken at traps situated over areas rich in decaying plant matter ancl overlain by (ii) (iii) from only a few millimetres of running water (positions V, and February to June, positions (iv) (v) and, (vi) from June to November) ' (i) (iv) (v) and Adults \,¡ere reared from samples taken at positions ' ' (vi)at Parawa and from Habitats 2 and 5 at Hindmarsh Falls' A continual supplyofrunningvraterisprobablyessentialtothesurvivalofthis in the species, si-nce although it is distributed widel-y at such sites south Mt. Lofty Ranges it is absent from large and otherwise suitable areas in the South East of the state' In winter months adults sv¡arm in bright sunshine from midday to just before dusk, but svTarms have also been seen during heavy misty rain at the Barossa Valley site of Nietskers' G. wilsoni is often associated with sparsicladius telmatus' Life Cvcl-e- Adults were taken in traps and by netting all year round; any part no separated larvae produced witsoni adults so no estimate of of the life cYcle is available' Distribution. SA, v¡estern VÏC' GRAPH 19 Gymnometriocnemus impecti-:ratus- Total captures at Botanic Park (dark) and other sítes (shailed) : a. ]-97L b. L970

GRAPH 20 Rhinocladius tonnoíri. Net captures, Botanic Park' 1970.

Horizontal Axis Months of ttre year Vertical Axis Numbers of specimens taken 5C t5 19. 20 lo 45 40

35 o

b 30 30

25 25

20 20 I5 I5 l0 10

5 5

o o o J A s 48

Gymnome triocnemus impectinatus' sP.nov.

Habits and Habitat. Adults have been reared from leaf litter with very low moisture content. Swarming occurs during the early morning over markers.

Lr- fe Cvc1e. At any particular site adults are on the wing for about six weeks between May and. July. As shown in Graphs 19 a and b, emergences at

Botanic Gardens were about six weeks earlier in I97I, than in 1970, a pattern repeated at other sites and probably reflecting the influence of

the earlier rains in 1971. G . impectinatus is perhaps stimulated to complete its development by suiÈable increases in moisture level.

Distribution. SA.

Taxonomic Usefulness. The fact that G. wil.soni adults swarm successfully during rain suggests they are highly resistant to wetting. It is possible that the long wing macrotrichia by supporting any surface film, prevent the wing membrane from wetting even under extreme conditions. This

function may also be important in other highly setose species, in particular, other members of this genus.

Genus RÉIINOCLADIUS Edwards

Rhinocladius tonnoiri Freeman

Hal¡its and Habitat. Adults have been reared from 'fungir at CSIRO laboratories in Canberra and I have reared them from leaf matter collected from the water's edge at Parawa (Position 15) and kept at laboratory temperatures for three weeks in August. It is possible, considering the slight morphological differences between specimens from these two areas' that more than one species is involved. If not, then it see¡ns that immatures are adaptable' although restricted to areas v¡here moisture remains even in the driest months. Adults were seen in loose swarms over markers at Botanic Gardens and Parawa during late afternoon' 49

Life Cvc1e. ( Graph20)AdultsareonthewingfromJulytoearlyoctober andthepatternofcapturessuggeststhatthespeciesisunivoltine. Distribution. SA, TAS, VIC' Other species in South America'

Genus SMITTIA Holmgren Smittia aterrima (Meigen) Immaturesofthisspeciesareterrestrialandadultshavebeeir the southern part captured widely in suburban Adelaide and through most of of the state, especíatly during the wetter months'

Smittia retracta Freeman Fewspecimensweretaken,butcapturessuggesttheremaybetwo generationsperyear,oneonthewingfromseptembertoNovember,theother from JanuarY to March.

Genus PARAKIEFFERIELLÌ\ Thienemann Paraki efferiella var tus sP.nov. Larvae for'lnd j'n Habits and Habitat. This is a rheophile species ' temporarytubesamongfilamentousalgae,Nostoc-Lycopodiumassociations, decayingleaves,matsofCharaandsubmergedtree-Ïootshaveproduced varr atusadults.Thelarvaefeedon'aufwuchs'.Untileclosionthepu'oa larva. swarminA-occurs remains in a tube constructed by the finar instar for several hundred at midday or dusk and swarms stretching conÈinuously Both sexes hrere metres were once observed at midday at Brownhill creek' ttre rapids at taken at light. Females resting in the splash zone of flights across Hindmarslr Falls (Site 11) vlere observed to make short quieter water, during which ttrey appeared to be ovipositing' TheabsenceofthisspeciesfromnorthernsouthAustralia,tlre sites in the south- I"lurray valley and the large expanses of sÈill \'rater at eastofthestate,andthelownumbersta]

is available. 50

Life cvcle. Adults were taken i¡r aII months except January and in peak numbersfromJunetoSeptember.NetcapturesgivenoindicaÈionofthe numberofgenerationsperyear.tlowever,materialwastakenfromamid- streamleafaccumulationatlii-ndmarsh-Falls(Site13)ontwooccasions, 14.iv.L972and,24-v.LgTzrandadultsemergedfrombotb'collections throughouttheperiod24.v.lg72Eo13.vi.IgT2,peakemergenceofmales must there- occurring 2 - 3 days before that of fe¡naIes. The life cycle synchronisation of fore-be at least 6 weeks rong and there may be some the life cycle not detected by netting'

Distribution. Known onlY from SA' divided into ttre subgetrera Taxonomrc Usefulness. Par akiefferiella has been (Brundin 1956) Paral

Genus PSEUDOSMITTIA Goetghebuer BothsouthAustralianrepresentativesofthisgenusarenewand argilla' sp' nov' each is confined to a single locality. Pseudosmittia between rarge brackish was taken at dusk from large s\^rarms rfear a claypan lakesinthesouthEastofthestate.I¡nrnaturesmaybeterrestríalor: in very low nurnbers lal

New South hlales, cras sipennis is a marine sPecies. 51

It is possible that more extensive collecting will reveal genus other members of this genus, each confined to a small area: the could prove a useful indicator of terrestrial, semiterrestrial or other conditions in Australia. Eric Bonython Reserve is only a few kilometres from the sea' along a fairly weII defined valley. Since a]l other Australian species just possible have been captured at or near bodies of saline water, it is that the specimens from Eric Bonython Reserve were blown upstream from nrore coastal areas-

Genus DOLOPLASTUS Skuse

Dolop lastus sensillus was found resting on mossy rock surfaces just

above water level (síte 11.b) at Hindmarsh Falls' lmmatures are probably found here also. However, relationships between species of this genus are uncertain and the remaining species may be found in quite different places'

5. DTSCUSSION A total o1 67 species in 2I genera of orthocladiinae was taken from the 67 sites at which collections were made. Four of the major sites support an outstanding diversity of species (see Table I) in 15, L7, 19 and 16 genera at Botanic Gardens, Eric Bonython Reserve, Hindmarsh FaIIs

and Parawa respectively. v{ith the excep tion of PodonomoPsis evansi species of podonominae, Aphroteniinae and Heptagyini are each restricted to one or more of tl¡e four major sites and, with the exception of Pseudosnittia argilllr every species collected was represented at one of tJ:ese sites ' Examination of Tables VIII and IX reveals that between the months of April

and september upwards of 30 species are on the wing. During this time individual corlections at each major site yield between 10 and 15 species. since most minor sites were visited at least once between April and

september some comparison between these and major sites is possible' It

can be seen from Tables I - IV that, together with Parawa, the three intensively sampled reserve areas (Botanic Gardens, Eric Bonython Reserve 52

fewer visits' and Hind,marsh Falls) are important: even allowing for the for no other sites show anln,rhere near the same diversity of species ' perhaps the greatest factor permitting such diversity is the amount of natural ground cover in the form of dense native herb growth and leaf litter. AlthoughmoreSpeciesarerepresentedatParawathanatanyother site,theyarerestrictedtofewergenera.Membersofeachgenustendto Falls have similar habitats so the greater number of genera at Hindmarsh of sub- and Eric Bonython reserve probably reflects the greater diversity with strates (and thus habitats) available at these sites as compared Parawa parawa and Botanic Gardens. The higher species : genus ratio at j.ndicates that a greater diversity of niches is available in certai¡r site is on a habitats. rt is worth noting here that, arthough the Parawa

grazhng property, the proprietor has completely preserved the natural pasture vegetation in seepage areas and refrained from using chemical altered than treatmenÈs near them. The Parawa site has probably been less may be species either Hindmarsh Falls or Eric Bonython Reserve. There preserved at Parawa which are no longer founcl elsewhere' species in south restricted to major sites and which may be usefur, particurarry Australia, as environmentl'1 indicaÈors are listed below' Aphroteni a australiensis Austroc ladius kvpensis

Par IA plumata woodi

It Cardiocladius australiensis híndmarshi Stictoclad ius multiserialis lyolobus tl uniserialis ves tubercus Paratrichocladius bifinestrus Sparsicladius gramanrs. Nasuticladius fuscus Kieffe istl'lus

Nrr diplos tyl-us Pseudosmittia scutella

I parapunctus

il duckhousi TABLE X

Nrunbers of sPecies and genera for which information is available and which are Present in each of the major tyPes of habitats'

Genera

Aquatic 10

Semiterrestrial 13 6

Terrestrial. . .. 4 2 53

The difference in diversity among the major sites is probably a result of the availability of more variation within particular habitats and 3'I' and differing environmental influences such as discussed above in The above discussion indicates the desirability of establishing a reserve along the crest of ttre ridge on which Para\'{a is situated' because fauna and flora it seems reasonable to suppose that in many other native groups the diversity of species and genera will parallel that of the Orthocladiinae. Five genera are particularly remarkable for their wide distribution in tïre areas sampled. Thus Paralimnophyes is found in most regions (25 (24 each\ Nasuticladius sites) , followed by Austroclad,ius and' Cricotopus ' (14) and Kieffe rophves (13). The abundance of Cricotopus spp. rs paralleled in other geographical regions and in south Australía is related to a certain resource - the avairability of filamentous algae. The wide from immatures of distribution o f Austrocladius however seems to result p'40) certain species tolerating a wide range of conditions (e'g' numerosus ' It is significant that the latter four of the above genera belong to group, which two groups' the C haetocladius group and the NasuÈicladius include most of the genera found in Australia' available' Table X shows that, of ttre species for which information is the greatest nunl¡er is to be found in semiterrestrial habitats' This partic- environment may present more constant conditions than fresh waters' ularly in arid Soutfr Australia' as weII The greatest numbers of insects from other dipteran families but as from sub-family chironominae were collected in october - March' of during the winter months collections consisted almost enÈirely Orthocladiinae. It appears that Orthocladiinae may have taken advantage of scarcity the milil winters of much of south eastern Australia, the relative to diversify of most other insects including predators having allowed them into many semiterrestrial and ter'restrial habitats' 54

SECTTON III MORPHOLOGY

I INTRODUCTION After nore than a century of investigation it í.s nol¡r generalry agreed that the family chironomidae can be considered separate from ceratapogon- question of idae and divisible into a number of sr:b-families' l}rere is no the separate status of the four sub-families, Chironominae, Tanypodinae' Aphroteniina and Pod,onominae. Ho\¡/ever a number of genera previously included with the orthocladiinae have been proposed as separate sub- fanilies by recent workers. Thus Brundi¡¡ (1966) recognises the large genus proposing D:grnCSg (Meigen) Pagast as sub-family Diamesinae and' in it also, Telma toqeton schiner as the sister group of Diamesinae elevates of sub- according to Hennig's principles of classification' to the status later famity, Tel-matogetoninae. His recommendations have been adopÈed by \^rorkers (e.g. SaeÈher 1969) ' genera Those dealing with immatures have proposed elevation of other to sub-family status (e.9. Clunio Haliday' Corynoneurinae), but these proposals have not been adopted in recent papers on adults' Ïnthisthesisthegroupsinquestiondonotplayanimportantpart clunio and I have retained the conception of orthocladiinae as including and the C"ry"""eqg V'linnertz group of genera' Brund'in(1956)proposedadivisionoforthocladiinaeintotwotribes provided Orthocladini and Metriocnemini, but on ttre basis of the discussion

in SECTION IV, Part 6, I have not adopted these names' grouPs which he Freeman in 1956 included in his genus Nanocladius 6 Nanocladius considered insufficiently documented to warrant separation: -

Thien. , Akief fe::ielIa Kieffer, Eukief f eriella Thienemann, Microcrico In the Thien., Parakieff eriella Thien and CamP tokiefferiella Goetgh. as well as adults recog- same year Brundin, with information on immatures Thien., Parakig[Ee.r- nised Camptokiefferiella as a synorrYm of Krenosmittia, 55 ietta and Microcricotopus as separate genera, and Eukiefferiella and Akiefferielta (together) as a single genus Eukiefferiella; he did not mention Nanocladius. In 196I Freeman acknowledged Brundin's separation of Eukiefferiella from Nanocladius but declined to attempt further clarifi- r¡r¡der Microcri- cation of the issue. From Èhe discussion in sECTTON III cotopusitcanbeseenthatNanoctadiusisaseparategenus;itis as recognised unrepresented in Australia. I have retained' the al¡ove genera by Brundín (1956). Attempts have recently been made to establish the homologies of the structures found in diverse insect groups (l¡atsuda 1958, 1965; Smith 1969) andschlee(1968),Saether(I971b)andHirvenoja(1973)haveattemptedto standardise relate these and other evaluations to chironomidae, and thus to terminology of the various tife stages. I have adopted their recommendations in manY cases. survey of Acknowledgment is due to Freeman (1961) for his useful Australian fauna Australian forms then available. sr:bsequent works on the (O' g' Edward' 1961) have dealt with the western Australian fauna in general south Australia (Glover' and the Tanytarsini (sub-family chironominae) of

1973). 56

2 ¡4ETHODS

2.L COLLECTION OF SPECTMENS Collecting methods have been described' above'

2.2. PRESERVATION OF SPECTMENS Following the recommendations of Freeman (1955) early collections vrere preserved by pinning. Howeveï, it was soon discovered that while dried specimens could rarely be identified on the pin, specimens preserved in spirit could be identified i¡nmediately because the terminalia and head appendages remained undistorted. Contrary to Freeman's clai-rn, spirit specimens from earlier collections showed tittle loss of vestiture, but showed increasing transparency of the exoskeleton. However, colour is of minimal value in Orthocladiinae and therefore all lat'er specimens were preserved in sPirit.

The arguments against the use of pinned specimens have recently been concisely set out (Sch1ee 1966) and will not be repeated here. It is notabl-e however that a number of inaccuracies or omissions in Freemanrs (196I) descriptions could not have occurred had spirit material been

examined.

2.3 PREPARATTON OF SPECI},ENS

All types of new species and specimens of previously narned species

have been presented mounted on sfides as follows' one wj-ng, including squama, was removed and placed in glacial acetic acid while the remainder of the specimen was heated for a short time in a I : I mixture of 53 aqueous KOH and 50% afcohol. The specimen was stained lightly with acid fuchsin'in glacial acetic acid and placed, with the unstained wing, in clove oi]. After clearing and dissection, head, thorax, legs and wing were each carefully arranged in a separate tiny drop of canada balsam on a slide and the balsam was then allowecl to dry completely. Later a second drop of balsam was added and a snall- piece of cover-slip placed over the part. Pupae were similarly treated but 57 usuallyrequirednostaining.Larvalandpupalcasesandlarvae\i{ere treated for I-3 days in cold KoH. The heads of larvae were left unstained'' on microscope cases and bodies \¡¡ere stai¡red lightly and similarly set up slides.

2.4 DRAWINGS necessary' Ìta-/e The types of new species and of otlrer specimens, where beendrawndirectlywithonlygrosslydisplacedpartsorbrokensetae corrected. AND CURATING OF 2.5 RECOMMENDATlONS REGARDÍNG THE PRESERVATION

CHIRONOMTD COLLECTlONS (a)Specimenscapturedinthefieldshouldbepreservedj-nspirit before tþeY dry out' (b) For at Ieast two reasons all types should be mounted on slides' (i) It takes little longer to prepare a whole permanent mount thantoremoveÈheterminalia,makeapermanentmountofthem in alone and attach this mount to the pin. specimens treated the latter way are often damaged during transport or when the terminalia are removed from a rusty pin for drawing' (ii) Adult types, accompanied by larval- and pupal casts preserved in alcohol, as left by Brundin, become very transparent intimeandconsequentlydifficulttoextractfromthevials in which they are preserved' Losses may sometimes result'

3 STRUCTURES OF TAXONOMTC IMPORTANCE Manyofthesearedescribedinearlyworks.MorerecentlySchlee (196g), Saettrer (1969, 1971b) and Hirvenoja (1973) have described new by saether characters and evaluated estabtished ones. Taxonomic works of setae and ex- Lg6g,1971c) and Hirvenoja (1973) have included counÈs are constant tensive measurements, but, while some of these measurements give species diagnoses in some genera or species, neither of these workers ormentionwhetherthemeasurementsareusefulforidentification. geogrçrphicarly Extensive measurements seem to me justifiabre only where variablepolymorphismsarebeinginvestigatedorwherelackofother 58 data poinÈs to their necessitY '

3.1 ADULT The structures discussed betow are those introduced or renamed in this thesis, or those named in recent works. Terms long in use have been explained by Freeman (1955), Schlee (1966) , Frommer (l'967), Saether (197Ib) and Hirvenoja (1973). when dis- 3. If Co 1our, qeneral shape and sture. These are described tinctive. AII are fairly uniform in the groups studied and colour is more dependent on the age of a specimen than its identity.

3.I2 Heacl ca ule. In some species the brow is extend'ed' down along themidline(rig.L7Ð¡thisdevelopmentisseparatefromthedevelopment of frontal tubercles- 3.12t setae. Following Saether (1971b) and Crampton (1942) I distin- gur sh vertical setae (v) and postorbital setae (po), but have not separated the verticals into'inner'and'outer'groups (see Fig' 79)' 3.r22 Eves. These are reniform with varying degrees of dorsal elongation. The elongation is estimated as the nr:nrber of facet rvídths between two lines drawn parallel to the midline: one through the tateral edge of the antennal foramina, the other through the medial extremity of the elongation (FiS. 79, Iines A and B respectively) ' rn some species the (nig' setae which form eye pubescence are arranged distinctively 65).

3.L23 Frontal tubercfes, sensifla and ocelli. In some Chir'onominae the brow carries on each side of the midline a protrusion known as a frontal tubercle. Hlxnps are found in similar positions in some Austra- lian orthocladiinae and are sometimes extended ventrally to encroach ort the antennal foramina (Fi9.111) . I have called these humps frontal tubercles (ft)and they are probably homoiologies (Hennig, L966' p'I17)' rather than homologies of the structures in Chironominae' In both Chiro-

nominae and orthocladiinae these frontal tubercles, when present, each caïry a small terminal circle of unsclerotised cuticle, here called a 59 frontal sensillum (fs). In species withouÈ tubercles the sensilla may that still be present in place of the tubercle (Fig. 79'). It is possible (1928) his these sensilla are vestigial ocelli as suggested by Jobring in study of Ceratapogonidae and by Saether (I971b) '

3.13 Mouth 3.131 The cibarial pump (fi 9. 79). Saether (197Ib) after Petersen (1916) and Tokunaga (1932) has identified schlee's 'kopfinnenplatter as the scterotised floor of the cibarial pump and the 'porus' as the orifice or the opening of the sal.ivary duct (after snodgrass, 1935) ' The shape and the and concavity of the dorsal border, the orientation of the cornua thickness of the tormae may vary and the structure is figured where divergence from Fig. 79 is significant' some gene!-a 3.L32 Mouth-part ratio. This ratio is introduced because in from the southern continents the clypeus and mouth-parts are extended' Distance from Èhe dorsal border of clypeus 79 M.R. = to of labial Fi width of head at wídest Par t.

There are two reasons for choosing the head width as a comparison' (i)Iftheextendedmouth-partsareforprobingtheireffectiveness witl be limited bY head width' (ii)Whentheheadissetonaslide,thismeasurementisleast

Iikely to 'be affected by orientation' that it is 3.133 The tentorium (Figs. 8, 18) ' Care has been taken correctly oriented before drawing' 3.I34 MaxilIary 1p (the p"Ip). Saether (1971b) following Tokunaga rather (1939) and Jobling (1928) considers that this has five segments' in than four- I have adopted this proposal- Saether points out that groupsrelatedtoChironomidae,palpalsensillaarefoundonsegrnent3 andthepalpiferislackingandthatthisisthecaseinChironomidae a1so,on1y if the ¡-'alp is considered as S-segmented' Thepalpshavebeenleftattachedtotheheadcapsuletoavoid.dis- it difficult tr:rbi-ng the alignment of internal structures. This makes 60

formulae have therefore to measure the first and second segments. Parpal i.e. not been calculated using Segment 3 as the unit for comparison, 1 : 2 z 4 z 8 : 8, but O'25 : o'5 : I ¿ 2 z 2' In this way the three mostvariableandaccuratelymeasurablesegmentsaredisplayedinthe most easilY comParable form' Acampaniformsensillumisfounddistallyonsegment2;hyaline sensillaoccurdistallyonsegment3andsometimesonsegment4(see

Figs. 79, e3).

3.L4 Antennae. l:962),Frorruner(L967)and 3.141 The eI.Jobling!g2s),Fittkau Saether(1969)havepointedoutthatthelargeglobularsegmentcarrying with the pedice! the flagellunr and often calÌed the scape, is homologous ofottrerfamities.Thetruescapeisreducedtoaringofcutic]ebasal primitive species' to the pedicel, in all but the females of some more 3.I42 FI ].lomeres.Saether(I971b)saysthatbecausethesectiorrsof should be called not the fIagellum are not independently musculated, they

segments but flaqeflomeres (after Imms' 1938) ' and named the numerous 3.143 Sensilla. Early in this study f described but not alluded sensory structures found on the antennae of chironomidae tobyearlierworkers.SincethenSchlee(1968)andserra-Tosio(i97o) has proposed a set of have díscussed these structures in detail and each (e.g. Kellog, L97o, Mclver, nanes. Ho\^¡ever, work on mosquito antennae 1970)hasshownthatsuchstructurescannotbecorrectlynamedwithout to make electron detaired historogy. since r have not had. an opportunity microscopestudieslhavetakencarenottointroduceunsubstantiated

homologies by careless choice of names' former term Fibrillae (federbuschborsten) ' I have adopted the fortheveryIongbristlesoftheflagellomelespresent'throughoutthe in dou-bt' Diptera and' whose homologies must be very little (¡'iqs. 25, L64) . C and eD line sens setae 80 , bases' often peg-shaPed- Type C setae are stout, sometimes with swollen 6I within the same sex of a particular species they are uniform in shape and size.TypeDsetaearenarrowbasallyandtapertolongpoints.Typec g setae can be present on all flagellomeres in the but in the ó there are usually onty 2 or 3 on the 2nd and 3rd flagellomeres, with several more neartheapexoftheantenna.TypeDsetaearefoundinbothsexesina group covering the tip of the last flagellomere' serra-Tosio (1970) reports 'sensilles chetiformes larges' from some basal flagellomeres in Diamesini whose form suggests they are homologous et'roites" with my Type c setae. However he reports 'sensilles chetiformes setae similar in form to.my Type D setae, from both the basal flagello-

meres and the aPex of the antenna' setae In many orthocladiinae it ís difficult to see all the hyaline of the basal flagellomeres and it is possible that the third of my Type C setae is really a Type D seta. However in aII species where the Type C (it possible of setae have been modified all three have the same form is influences) course that boÈh c and D seta have been modified by the same ' and schlee distinguishes only between those setae on the terminal segment both those on the basal segments, but his drawing (1968,Fig' 87) shows

Type C and Type D sensory setae at the apex' near Sunken setae (rig. 25 ) Conical setae in narrow pits' one of the the end of each of fJ-agerlomeres r and 2 and a pair at the tip last f1agellomere. pits on Ring-shaped setae (rig- 25 ) Peg-1ike seÈae in broader the borders of the antennal furrow at the end of the last flagellomere' setae' In some species they are encircled by a ring of minute to the 3.15 Thorax (fig. 37 ) Thoracic sclerites are named according is here the system of Matsuda (1970) and saether (I971k). The pronotum mesonotum the antepronotum, the sternopleuron the pre-episternum and the

scutum. 3.16 l{ings. TíIlyard's modifications Qg24) to the Comstock-Needham g and the fold system of naming wing veins has been adopted (rig. ) 62 under cu, is called cu, as suggested by l,indeberg (1966). The Aspect Ratio is inÈroduced as a guide to the aerofoil characteristics of the wing (see .IV.6.I) and is abbreviated Asp. R. Between specimens of the 23' same sPecies, Aspect Ratio has been found to vary no more than which is no greater than the er:ror incurred in completely measuring the same specimen on a number of occasions' (H#HL. ) Aspect Ratio =

Wingareavlasmeasuredwithaplanimeter.AspectRatiohasbeenpre- ferred to a simple length:breadth ratio since a check revealed that while the latter remained constant, Aspect Ratio could vary up to half a unit. Positions of wing sensilla and microtrichiation are described as they appear when viewed at 2OOx magnification; microtrichia may be ,invisibler'small or large and their punctures in the wing membrane may

be minute or plain. Descriptions of venation have been abbreviated

where possible, ê.9. in the phrase '*4*U ending closer to Mt*rr ,closer ao tl*2' means 'closer to the level of ML*2 than to the level of Cur.' 3.r7 Abdomen. v,Ihere distinctive, colour patterns, patterns of abdominal setation and patterns of spots of unsclerotised or heavily sclerotised cuticle are descríbed'

3. t8 Terminalia. from the dorsal- 3.18I Male- (r'ig. SZ )- AIl drawings have been made view. Attaclredtotheninthsegmentate2largemovableventrallobes' the qonocoxites (Saether 1971b): the basal band of sclerotisation (saether 197Ib) sometimes found on these is called the coxaPodeme ' of the loth sternite' The s ternapodeme (Saether 197Ib), all that remains which is a band of sclerotisation with lateral and transverse arms Iink.stlregonocoxitesdorsally.oneachsi.deadiagonalsclerotised 63 rod, the phallapo4erne (Saether 1971b, Abul Nasr I95O), Iies ventral to gono- the sternapodeme and projects back into the basal region of the coxite. The phatlapodeme is sometimes expanded medially and can be supported from the junction of the sternapodeme and gonocoxite by a short sclerotised bar. (Fig'97b) when the sac containing the intromittent organ is everted support for it can be seen that the phalrapodeme twists out to provide is the ventral and lateral parts of this sac, while the dorsal region partlystrpportedbytheninthtergite:whenretractedthesacforms the ,aedeagal lobes' of saether. In some orthocladiinae the intro- with a nr-ittent organ carries an apical u-shapéd area of sclerotisation free-standing sclerotised hook through which the ejaculatory duct apparentlyopensandwhichlhaveca].ledthepenapodeme('penis', Edwards 1931)*- As a measure of the size of the penapodeme relative to the reproductive system the Penapod'eme Ratio has been calculated:

Lenqth of penapodeme (see rig. 97a) P.R. = Length of gonocoxite Thephallapodemeandthepenapodemebothmakeupapartofthepenis sheath and shourd therefore, accordíng to Abur-Nasr, be considered parts of the phallapodeme. However the apical structures are so distinct here that a separate nane is desirable' Hirvenoja (1973) extensively examined the homologies of chironomic since it terminalia. He called the phallapodeme the 'pars basalis' some makes up the basal part of the intromittent organ. In 'primitive' into species the medial parts of the phallapodeme have differentiated medial distinct structures which he named the lateraf, intermediate and parameres.Heproposesthesethreearehomologouswithcertainlobes foundonthei¡rnersurfaceofthegonocoxiteinless'prímitive.

least * I have not examined this structure histologically but at in larger sPecies it is hollow' 64 species and names the latter accordingly. Hirvenojars derivation of the particular homologies must be considered as tentative and, since the inner tobes of the gonocoxite play no part in the eversion of the intromittent organ, I have not adopted his names but instead have called them generally 'inner lobes' and described their positions where necessary. The penapodeme is not at all related to the pars ventralis described by Hirvenoja as the latter lies ventrally between the bases of the gonocoxites- 3.182 Femafe (fig. f58)-- Drawíngs have been made from ventral view unless otherwise indicated. The posterior border of sternite vIII is slightly indented and on each side of this indentaticn the sternite bears a nr¡nber of setae and a sclerotised band of cuticle. The spermathecal opening is bordered by a fringe of hairs, the genital comb (Hirvenoja 1973). The membran- ous area immediately posterior of sternite vIlI carries numerous setae (wensler and is often shaped into a pair of protrusions, the egg guides

& Rempel , Lg62; valvulae inferioÏes' Hirvenoja ' L973) ' segrment fX consists of a tergite and sternite as wel-l a lateral sclerite (ninth lateral tergite, Wensler & Rempel' L962) ' Hirvenoja

has named the sclerotised band.s of sternite vIII and these lateral scleriËes of segment IX, gonocoxites. There is some dor:bt whether these structures are homologous v¡ith the structures which are known in males and in primitive groups as gonocoxites and I have adopted Hirvenoja's terminology only with reservation' Tergite IX may have a medial cleft and is often distended unÈil, from dorsal view, it partiatly or completely obscures tergite X' (Iateral sternite IX is reduced to a pair of internal scl-erotised bars apophyse, Hirvenoja, L973) ' (post- Bet\deen the bases of the cerci is a small scterotised plate genj-talplate,Wensler&Rempel,Lg62).Inthemoreprintitiveortho- cladiinae d,iscussed below this plate forms a strongly sclerotised bar' 65

Iinking the opposite sides of the tenth tergite and must be considered' as tentatively suggested by Hirvenoja (19?3), as the tenth sternite' The internal sclerotised apoderne which supports the oviduct is almost certainly homologous with simil-ar structures in other groups and so has been called the vaginal apodeme. In many species it is broadened basally and linked tightly with what remains of the ninth sternite. The spermathecal ducts sometimes have a distinctive basal broadening. (nigs- 95, Io5)

3.2. PUPA

3.2r C othorax In naming the cephalothorax setae I have followed Brundin,s proposats but incorporated modifications similar to those adopted above for the thorax of the adult. Thus the cephalothorax has anterior and posterior frontal setae (nig. 69), anterior ancl posterior antepronotal setae, three precorneal setae, scutal setae L - 4 and varying numbers of postnotal setae' Hirvenoja (1973) proposed that the setae of the pupa shoul-d be

named after setae which occur in similar positions in the adult. I^lhile any attempt to simplify naming systems must be welcomed, f have not adopted his proposal since similarity in name carríes an irnplication of

homolog.y which has not here been established and especiaÌIy because confusion might easily arise if structures of the pupa and adult were

to carry the same names. 3.22 Abdomen. I have followed Pagast (1947, PP. 5O4-5O5) a¡rd Zavrel (Lg42) in naming the structures of the abdomen with the exception that'

as suggested by Hirvenoja (1973, p. 33), the dorsal and ventral setae

of seg:ment VIII are separated ínto rnedial and lateral groups rather than numbered, because their homologies with the setae of oÈher segment-q cannot be readilY ascerta-ined.

3.3 LARVA Hirvenoja (1973) has discussed structures of tlre larva at great length; I have adopted his proposafs- 66

4 SPEC IES DESCRIPTIONS

vlith the exception of Paraheptagyia ennea and the two species of Austrobril-lia, aII species described below are representecl in South Australia. Descriptions have atl been made from slide material and of necessity, only some of the diagnoses provided include characters which are detectable under high magnification. The number of slide mounted specimens examined is shown in brackets at the beginning of each description and, when variation between specimens is small, only the average measurement is given. If for some measurements of slide the nunrl¡er of measurements available is different from tlre number The only specimens examined, this is indicated in the appropriate place' exception is the aspect ratio, where because of the time involved, two specimens were usuaIIY measured' this Drawings are accompanied by a scale line. In drawings of wilgs scale line is 5oou long and in all other drawings it is 5o9 long' I have collected all specimens not otherwise attributed' Abbrevia- tions of institutions from which specimens or types were obtai¡red for examination are as follows'

MM Macleay Museum, University of Sydney;

NM National Museum of Victoria, Melbournei

SPHTM-SchoolofPublicHealthandTropicalMedicine' UníversitY of SYdneY. 67

KEY TO SUB-FAMILIES OF THE CHIRONOMIDAE

I True base of M3*4 presetrt 2

True base of M3*4 absent 5

2(I) R2*3 forked TANYPODINAE *2*¡ either simple and unforked or absent 3

3(2) R2*3 .o*pletely absent, radial veins well separated; postnotal furrow absent PODONOMTNAE

R2+3 and distinct 4 "i*ple

4(3) Postnotum lacking median longitudinal furrow; male antenna witlrout plumes TELMATOGETONAE and ORTLOCLADIINAE Genus Clu¡tþ Haliday

Postnotum with well developed tongitudinal furrow DIAMESINAE

5(1) Rr*, absent APHROTENIINAE Rr*, Present 6

6(5) Ratio of anterior basitarsus to tibia (i'e' L.R.) Iess than I; tibial- combs of posterior 4 leqs not composed of short basally fused spinutes; male stytes folded inwards " ' ORTHOCLADITNAS

L.R. nearly always more than 1, very occasionally about I; tibial combs formed of short basally fused spinules; male styles directed rigidly backwards (with possible exception of Parviterqum gen. nov.) CIIITIONOMINAE

(afte: Freeman 1961) 68

SUB-FAMILY APHROTENITNAE

Genus APHROTENIA Brundin Aphrotenia Brundin, 1966, P. 338'

Type species A . tsitsikamae Brund. Brundin reported this genus from South Africa only' The discovery of this Australian species thus represents a large extension of the range of the genus. Brundin,s 'Key to the known genera and species of the sub-faraily Aphroteniinae' (1966, p. 338) can be altere

Imagos L(2) Cross-vein r-m short, forming a dis+-inct angle to R4*5t acrostichal setae reaching scutellum; free end of costa . APhrotenia Brund shorter than R.I- la (Ib) Tergite IX sÈraight cut posteriorly, basistyle without lobe. South Africa A tsitsikamae Bruncl Ib (Ia) Tergite IX rounded posteriorly, basistyle with small lobe- South Australia A. aust-r-eliellliq sP. nov.

Aphrotenia australiensis sp' nov'

Separated from African species since macrotrichia cover most of wing since membrane, tergite IX rounded distally' Resemb Ies Paraphrotenia scutal process overhangs head and gonocoxite has inner lobe' Brow drawn MALE (2) . Head. Post orbitals present, verticals absent' well down slightly lateral to midline. Eyes shortened, frontal sensil-la separated. Dorsal border of cibarial. pump straight. M.R. o.3, tentorium narro'Â/. Sensil-la apparently absent fronì palps, palpal formula O.2:1:1: L.67 z 6.2' segment5curvedoutwards' æU1with14 flagellotneïes, 2 and' 3 eachwith 2 TypeC setae' 2- 12 fusiform; 5 - 12 twice as long as bro¿rd, 13 twice as long as 14' At least 2 rows of Fis. 6 Aphrotenia australiensis sp.nov. male, wing.

tl Fis. 7 terminalia.

Fig. I Paraheptagyia plumata, sp. nov- male, tentorium.

Fig. 9 ll wing

Fig 10 I terminalia.

Fíg. 11 Par l_a @, sp. nov. male, antenna (inner surface of left antenna).

Fig. 12 ¡t ma1e, terminalia. o

(L ) f¡ (¡. ) ù qr, ', v v;

7 10

t 9 3t¿

\ 1 ,t Y \\ \ I 1'

11 12 69 fibritlae on each flagellomere. Thorax. Antepronotal lobes small, each with 5 lateral setae, scutal process projecting somewhat over the head. Acrostichals extend only two thirds of distance to scutell-um, only 8 dorsocentrals, 6 pre-alars, I supra-alar, 5 large and 2 small scutellars

(rig. mm. squana with 4 in a single row. V,Iings 6). Length I.O ' ".i... Macrotrichia on all veins and spread over all cells but those of the costa' Rr. R4*5 at level of cur, costa produced half-way to wing R and ".ding tip. Legs. L.R. O.35. Abdomen. Tergites with 4 paírs of setae in a single row, sternites with one pai-r. Terminalia (FiS. 7). Posterior border of tergite fX large and swollen. Lateral arms of sternapodeme Iong, transverse arm absent, phatlapodemes absent. Gonostyle with small inner 10be built from the conical socles of numerous setae. TYPES. Holotype d and single d paratype from Eric Bonython Reserve, South Australia, 18.xi. 1970-

ST'B-FAMILY DIAMESINAE

TRIBE HEPTAGYINT

Genus PARAHEPTAGYIA Brundin

Paraheptagyia Brundin, L966, p. 379.

Type species P . cinerascens Edw. Brundin's ,Keys to the known genera and species of the tribe Heptagini' (1966 p. 377) can be altered to accommodate the new Austral-ian species by the alteration of two clauses and the addition of two coupletg'

The key then becomes, using Brur¡d'in's terminology:

Imagos 5 (8) plume of male antenna normally developed; A.R greater than 1; legs with long hair 5a 5a (5b) Male without tarsal beard; dististyles with terminal tooth. . P. pltxnltg sp.nov. 5b (5a) MaIe with tarsal beard; dististyles without terminal tooth P. cinerascens Edv¡' 70

L4 (3) Tilcial spurs faírly small, inner spur of hind tilcia only slightly longer than ti-bial diameter; heel present . LAa 14a (I4b) Dististyles with terminal tooth, heet of moderate síze, male antenna lO-segmented - - . P' ennea sp' nov' :lfib (14a) Dististyles without terminal tooth, heel small, male antenna l4-segmented . P. andina Brund. Paraheptagryia Plumata, sP. nov. Separated from other Australian species by complete complement of flagellomeres, normal- antennal plumage and lack of tibial bands'

MALE (2). Head. post orbitals a¡rd. verticals fine, scattered. Frontal sensilla absenÈ, brow slightly drawn down. Eyes truncated across dorsal border, cibarial pump with dorsal border deeply, rectangularly hollowed. M.R. O.3O; tentorium, see Fig. 8. segment 3 of palp with large group of fine, slightly capitetlate sensilla, patpal formula O.3 : 0.6 : I : 1.1 : L.7. Antenna. A.R. 1.40 mm., 13 flagellomeres present, plume well developed, flagellomeres 2 and 3 with tiny Type c setae (16¡), 2 bulbous and 2 - L2 broader than 1ong. Last flagellomere slightly clubbed at apex with numerous Type D setae (1O¡r). Thorax. About 2I small anterior pre-alar setae along posterior border of humeral pits, not continuous v¡ith pre'-alars. Legs. L.R. 0.6. Tibial spurs It times as long as tibial diameter' Middl-e and hind legs with two pairs of ventral spinules on first tarsus, one pair on second tarsus. Tarsus 4 longer than 5, sole on 4 only slightly over-

mm. squama with 50 - 60 setae' hanging 5. vüing. (nig. 9). Length 2.5 ' anal lobe large, distinct. Punctures of microtrichia conspícuous, macrotrichia on base of R, R, and R4+5t five sensilla irregularly spaced basally on R, three similarly placed on RI' *2*3 meets costa half-way between R, and R4+5, *4*5 .tding closer to Ml*2, costa produced one third of distance to wing tip. cu, curved gently downwards, An extending nearly to wing margin. Abclomen. Tergites and posterior sternites clothed in fj-ne setae, Terminalia (riq- 10). Anal point short, squared' Sternapodeme evenlyarched'cornersindistinguishable,onlyslendersclerotised 71

Iink with triangular phallapodeme, but meeting dorsal surface of gono= coxite in slightly sclerotised ridge. Inner lobe drawn out and with numerous stout setae. Gonostyle with moderate heel, slight dorsal srvel-]- ing and small Èerminal sPine. TypES. Holotype d Botanic Gardens, South Australia, 22.ix.1971. paratypes. Botanic Gardens, ld (at light) 4-xi-L97O¡ Hindmarsh Fa]Is, 1ó (damaged, in stream nets) , 2L'vii'1971' Paraheptag:Yia ennea, sP. nov.

Separated from other Australian species by reduced number of flagr- ellomeres and from south American species by its larger heel. MALE(1).Similartoplumatabut:-Head.Brownotdrawndown,eyes normal. Segment 3 of palp with large group of curved sensilla, palpal formula o.4 z o.7 : I : I : I.5. Antenna (Fig' 11)' A'R' I'3' only 9 flagellomeres, fibrillae reduced in size and nunù¡ers, pedicel with single fibrilla. Flagellomere 1 medially constricted' flagellomeres 1' 2 and 3 (8¡r) each with single sunken setae, 2 and 3 each with small Type C setae ' 2 - 7 broader than long. Last flagellomere with severaf basal fibrillae' apex clubbed and hollowed with one stout bristle, numerous small Type c setae and 3 ring setae. Thorax. Antepronotal lobes large, not tapering medially. About Io anterior pre-alar setae along posterior border of humeral pit, almost continuous with pre-alars. Legs. L"R. 0.6, tibial spurs as long as tibial diameter. Middle and hind legs with a pair of stouÈ spinelets ventralty on end.s of first and second tarsi' vling'

Length 2.4 mm., squama with about 50 setae. Punctures of microtrichia conspicuous' macrotrichia on bases of R and R*; only two wing sensilla' positioned as in Orthocladiinae' O2*3 meets costa closer to Rr' R4+5 ends closer to tL*2- Costa produced one fifth of distance to wing Tergites tip. Base of Må*4 *""t" M3*4 slightly distal of pfk" Abdomen' Terminal-ia. and posterior sternj-tes with uniform covering of fine setae' (Fig.L2).Sternapodemewithsmalldistinctcorners,extendedonto ridge' dorsal surface of the inner lobe of the gonocoxite in scferotised 72

Inner lobe smaller than plumata. Gonostyle with less distinct heel and larger dorsal swelling than plumata' TypES. Holotype d from Lee's creek A.c.T. alt. 2500 ft.) 3I.x.L962'

(I.F.B. Common and M'S' UPton) ' The affinities of these species are extensívely discussed below

(sEcrroN rv, 2.) . SUB-FAMILY ORTHOCI,ADIINAE unless otherwise stated, the males of all species described below postorbitals complete' have the following features in common:- Head' Row of (nig' Segments 2 and 3 cibarial pump wit]. evenly curved dorsal border 79)' of palp with single basal sunken seta. Labium with numeÏous 'gustatory' F'lagello- sensilla. Palpal sensilla on inner distal surface' Antenna-' D setae slender meres 2 and 3 broader than 1ong, type c setae slender, type at the base of with short whip-like apices. Wings with two sensilla, one Sternites witl- R2*3 R4+5, the ottrer near the base of R' Abdomen' RI, "td rv or minute mediar and 1aterar setae on segments rr-rv or v, sternites V - VII rrniformlY setose' (1956) Genera The key provided below is based on that of Freeman ' representedinAustraliabutnottreatedindetai].inthisth-esisare bracketed.

KEYTOAUSTRALTANGENERAOFTHEORTHOCI'ADIINAE utd I Minute species, wing length t-2 mm; Rf T¿*5 fused to each other and almost entlrery entirely 2 fused to costa to form a thickened "clavus" 3 Clavus absent, veins not fused 2(L)Eyespubescent,posteriortibiasimple(ThienemanniellaKieffer) witl¡- Eyes bare, posterior tibia obtiquely truncate and a conspicuous apicar projection ?:.l|"ut "idt (corynoneura winnertz)

3(1)VÍingswithmacrotrichiaonmembraneatleastatapex4 8 Wing membrane quiLe bare of macrotrichia Edwards 4 Labial- palps elongated, to form a sucking tr¡l¡e "Rhínocladius 5 Labial PalPs not extended 73

5(4) Acrostichal bristles present 6 Acrosticha-l- bristles absenÈ 7

6(s) Eyes with well-formed dorsal na-rrow portions' posteriorforklongParametriocnemusGoetghebue::

Eyes without dorsal narrow portions, posterior fork shorter (Metriocnemus vlulp) (Allometriocnemus Freeman) 7(5) Squama with strong complete fringe Goetghebuer Squama bare or with 2-3 bristles riocnemus B(3) Eyes pubescent, squama usual-Iy fringed 9

EYes bare, except in Smittia. and Pseudosmittia and here squama bare T2 Edwards 9(8) V'Iings heavilY Patterned Stictocladius lo Vlings Plain ventrally, wings with R 10(9) Eyes displaced et Harn. nearly fused with Rn*U fiT3ro"ri Thien I1 Eyes normal or slightly elongate dorsally' large specl-es punctures 1I(IO) Scutum with distinct dorsocentral bristles' distinct,humeralpitslargeParatrichocladiusSantosAbreu smalI, punctures Dorsocentrals ltlulP minute cricotopus van der distinct puncÈures 12(8) Microtrichia with t3 PenaPodeme PresenÈ

Microtrichiaofwingmembraneminute,prrnctures I6 tíny, penapodeme someÈimes present 13(12) outer spur of rear tibia sometimes greater than length of inner, squamal fringe half Freeman comPlete Austrocladius less than half lengttr of inner' Outer spur L4 squamal fringe reduced membrane large 14(13) Microtrichia of wing Spars icladius gen. nov. Punctures fine distance, punctures plain' Microtrichia 15 costa Produced with single strong bristle 15(14) Apex of antenna es Brundin anal Point bare Par absent, anal Point Apical bristle Eaton absent Limnophves 74

16(12) Squama either partially or completely fringed, pronotum often quite large L7

Squama guite bare, pronotum usually more reduced 2t

17(16) Outer spur of hind tibia at least half length of inner t8

Outer spur less than half length of inner I9

18(f7) Fourth tarsal segmenÈ cordiform; styles simple Cardiocladius Kieffer

Fourth tarsal segrment cylindrical; styles double Austrobrillia Freeman

19(17) AnaI point rounded, covered with stout curved bristles Kieff s Freeman

AnaI point otherwj-se 7-c

20(I9) Anal point bare, curved, keel like Nemoricladius gen. nov-

Anal point reduced' often to ridge; if long then wings darkened Nasuticladius

2I(16) Antenna with stout black bristle at aPex Smittia Holmgren

Antenna without tt¡is bristle 22

22(2L) Distinct Pulvilli Present (Camptocladius WulP)

Pulvilli absent 23

23(22) Costa retracted, ending well before Ievel of apex of M3*4, posterior fork short, accessory (Orthosmittia Goetghebuer) fork present

Costa ending on level with or beyond aPex of Mr*n posterior fork longerrno accessory forK 24 75.

24(23) Scutal humP Present 25 Scutal hump absent, number of reduced, flagellomeres flagellomeres Do]-op1?Ftus Skuse longer than broad

25(24) AnaI point short, broad' bare Parakief ferie lla Ttrien.

Anal point a.bsent or tinY, setose Pseudosmittia Goetghebuer 76

Genus AUSTROBRÍILI-A Freeman

AUS trobrillia Freeman, L96L' P'640' Freeman 1-96L TIT)e species Austrgbrillia longipes ' ThefollowingredescriptionhasbeenmadewithreferencetoFreeman'Stype and Members of this genus are large series and to the new species collessi' palliil compared with most Orthoclad'iinae' long and fine' uniserial except for M.ALE. Head. Post orbitals and verticals agroupclosetothemidline;frontalsensillaabsent.Eyesbare,dorsal pump with deeply curved dorsal border' elongation 7 facets wide, cibarial than half length of antennae' M'R' very corners directed inwards' Palps more Flagellometes 2-L2 broader than long' Iow. Tentorium as in (Fig'13) ' Antenna' lastflagellomerewith3typeCsetae(35p)and2subterminalsunkensetae. Thorax.Antepronotumreducedrlobeswellseparated,5tororaterarsetae; acrostichalsabsenÈ,dorsocentralsirregularl-ybíserial,beginningjustbefore humeralpits;6pre-alars,lsupra-alar;scutellarsin2irregularrows.Legs" L.R.l.Reartibiawithouterspur3/4lengthofinner,basalhalfofeachv¡j.th stoutpoints¡combnotdense,aÏranged'in2arcs.Pulvllliabsent.Vüing. Squamawithnumeroussetae,anallobepresent.Macrotrichiaonbasalveinarrd prain. singre sensirlum on r-m' *rn, tto=tt R. Microtrichia strong, punctures tip' Pfk of yf+Z' Costa produced nearly to wing to Rr, R ¿+sending distally An ending far distal- of r-m' close to level of r-m' c5 øently curved'

}bdomen.Tergitesdenselyanduniformlysetose.Terminalig.TergiteVffl short.Analpointbare,sternapodemewithshorttransversearmiphallapodeme Iongandthin,penapodemeabsent;inner-lobesetose,finger-like.Gonostyle withoutspirre,twolobes:-onedorsal,bare,withstoutpointloflgr'entral with setae concentrated at tiP'

FEM.ALE Like male but:- wide' eyes with dorsal elongation 5 facets Heacl. Post-orbitals absent' 2 - 5 eachwiÈh 3 type c setae (28p) 6 wj-t-h 6 rywith 6 flagerlomeresi multiserial' Wings-' Microtrichia setae distally' Thor:ax" Dorsocentrals Fis. 13 Austrobrillia collessi sp. nov. maIe, tentorium.

il Fig. L4 wl-n9

Fig.15 il tt tt terminalia

Fig. La I female, gonocoxite rX and egg guide.

Fig. tf Austrobrillia (?) longipes Freeman, female, terminalia, inseÈ showing dorsal view.

Fig. 18 Cardioc ladius australiensis Freeman, ma1e, tentorium'

f 14

! ? 13 18 a !t v

a aa a a aa aa a

a a

-. ç 17

16 77 Ionger, punctures more conspicuous. Macrotrichia on basal vein R1 and R4*5.

Terminalia. Sternite VII with numerous setae on eact¡- side, qonocoxite \/II1 scarcely sclerotized, egg guides sickl.e shaped. Tergite I'X with complete medj'ai cleft, not obscuring tergite X. Sternite X strongly sclerotízed., lirrking eacTr side of tergite. Cerci elongate, sinuous- Vaginal apodeme narro\rr, not expandeá basally, not projecting into segment VII. Three spermattrecae:- two 1atera1, sac-like with long tapering necksi one medial, spherical, with narro\^/ neck, all ducts have some sclerotization.

KEY TO SPECIES OF THE GENUS AUSTROBRTLLTA

Males. !{ing narrow (Asp.R. 5.05), inner lobe closely

adpressed to gonocoxite collessi, sp.nov. l{ing broader (Asp.R. 4.41 , inner lobe standíng out from gonocoxite Iongipes Freeman

Females.

Egg guide ending with knob, wing narrow collessi, sp-nov.

Egg guide ending with point, wing broad ? lonqlipes Freeman

Austrobrillia collessi, sp.nov. Separated fron longipes by its narrow wing, closely adpressed inner lobe of male and knobbed egg guide of female.

MALE (].) Head. Segrnent 3 of palp with short rod-like sensilla t25p) scattered widely over distal two-thirds. Palpa1 formula 0-25 : 0.34 : 1 : I : l-.6. Antenng. A.R. I.73. Flagellomeres 1 - 4 with 2 tlpe C setae (35p); l-ast flagellomere with 3 ring setae. Wi"g-. Length 3.3mm. Asp. R. 5.05 (FiS. 14).

Squama with 17 fine medium length setae. AnaI lobe smalJ-, macrotrj-chia on R4*5. Pfk just distal of r-m. Terminal-ia. (Fig-15, partly evagj-nated) Sterna- podeme with corners drawn out. Inner lobe closely adpressed to gonocoxite. Both lobes of gonostyle paral-1eJ--sided, ventral one with a few sub-terminal- setae. 7B

FEMÀLE ( 1). Like male but:- Head. Verticals more numerous. Third palpal segment with sensilla forming clusters distally. Palpal formula 0.25 z O.4 : I : I : 1'6' Antenna. Flagellomere 6 with single basal type C seta. Flagellomere formula 1.3 : 1 z I.2 : I.55 : 1.45 : 2.05. Wing. Lengt]rl 4.4 Ím. Asp.R. 4.7

Terminalia. Compared with lonqipes, gonocoxite IX larger t egg guide knob- 1 shaped (FiS fA\. Sternite X narrower and tergite IX smaller. Types. Holotlpe d and I paratype, Mt. Beauty, Victoria' 26.x.1961, (D.H. Colless)

Austrobrillia longipes Freeman

Austrobrillia q Freeman, 1961 P.640; Fig.5b. (25¡t) MALE (A',) . Head. Segrment 3 of palp with short rod-like sensilla scattered widely over distal third. Palpal formula (1) 0.3 :0.43 : 1 : 1 z I.4. Ante¡ila A.R. 1.70. Flagellomeres I - 5 with 2 tlpe c setae (35p); last flagel-lomere with 4 ring-shaped setae. v{ing. Length 2.8 - 3.8 mm. Asp. R. 4.43. Squama with about 20 long fine setae. AnaI lobe larger tha:r in collessi. Pfk at level thall of r-m . Ternuinalia. (Freeman 1961 Fig. 5b) Anal point longer and sharper in collessi. Finger-like inner lobe offset from gonocoxite. Dorsal lobe of gonostyle broadened, ventral one with isolated seta inset midway along' TYPES. I have examined the holotype d t Harz Mountains, Tastnania' Dec ' 1922 (4.L. Tonnoir) and the parat)æe series' separated Thre type series of longipes contains only males, captured in widely regions of Tasmania and Victoria. A female specimen from Both\^/ell' Tasmania (I3.ii.L964), A.L. Dyce & M.D. Murray) has wing Ineasurements compatible with

longipes males a¡d resembles them with the follcwing exceptions i-

Head. palpal formula 0.4 : 0.6 : 1 : 1.1 : 1.6. Flagellomere 6 with nwterc¡us ring setae. Flagellomere formu]a 1.05: I z I.2: !.2 z L.2 z L.4. Vüing' shaped, Length 3.4 mm. AsP. R- 3.8. Terminalia. el g"i;â) Egg-guides sickle gonocoxite IX small , sbernite X brrrad' It is possible th.t -@lÆ- an<1 col-lessi represent the oppos-ite end's of a cline or are the products of recent geographical isolation since a.11 Fís. 19 Cardiocladius australiensis male, ventral surface, tarsus 4.

Fig. 20 Diplocladius uniserialis Freeman, ma1e, tentorium.

Fig. 2I lt il terminalia.

Fis. 22 I I female, terminali-a.

Fig. 23 Diplocladius multiserial-is Freeman, female, tergites VIfI and IX. a

I l-¡

21 20

¡.,

t4 û' Ò

23 79 coll-essi Iongipes specimens originate south of the Great Dividing Range while is from the north of this Range'

Genus CARDIOCLADIUS Kieffer Freeman 1961rp'638' Cardio cladius Kieffer, LgL2 a.p.22; Brundin, 1956,p.66;

Type specaes Cardioc ladius ceylanicus Kieffer, L9I2.

Cardiocladius australiensis Freeman p'639' Ca rdiocladi us australiensis Freeman 1961, markings, coxite Freeman saYSt ,,Dark brown, shoulders pruinose, legs without Iobe righÈ-angled, styles with small aPical sPine" ' dorsal border, elongra* MALE ( 21 . Head. verticats absent. Eyes with straight border' tion 2 facets wide; cibarial pump with slightly V-shaped dorsal- sensilfa (20p) segment 3 of palp slightly curved with 2 clusters of capitellate ' : : I.37 ¿ 2.26. one dorsal and one ventrali palpal formula 0.37 : 0.56 1 M.R.O.33.TentoriumasinFig.IS.Ant-enna.A.R.1.15(I).Flagellomeres2_7 with 2 sunke¡l broader than long; type D and c setae absent; last flagellomere B setae' setae and 4 ring setae. Thorax. Antepronotum with row of -'l-ateral biserial beginning well Humeral pits long, oval; dorsocentrals irregularly extend half way after pits; acrostichals beginning close to pronotum, minute' to scutellum; pre-alars uniserialr shoÏt; large group of strong supra-a1ars; t4(Fig"19L scutellars muttiserial. Legs. L.R. O'5' All legs with cordifrom t3't3 and tn minutely and inter-segrmentalmembransbetween t, and trr t, and ventral surface setose. Middle and rear tarsi with 2 short terminaf spines on oft'lÈ,andt,.Spursofreartibiasub-equal'bothmorethanhalfcovered in 2 crescents' with accessory setae; comb with about 15 slender setae arranged Wing.Length2.Omm.ASp.R.3.62.Squamawithmorethan40setae;r.acro- *Z*3 ends before trichia on Rrr R only, microtrichia minute punctures visiblet meetingcosta,R4+5endingcloserto}a'*,icostanotprod'uced;pfkjustdistal setose' of r-m; CuI strai-ght' e¡ag*"".' TergiÈes densely' uniformly narroh/' Terminalia (l'reeman 1961 Fig.5a) Anal point absent' SternaBocleme penapoden"e cornerspointed¡phallapodemewithstoutbaseandrectangularplates; 80 small' absent. Inner Iobe right-angled; gonostyle boat-shaped, spine (A'L' TypES. I have exa[ined the holotype d , Larnceston, Tasmar¡ia oct' l-922 p seen. t"""-"-rt, held by SPHTM and paratypes at AIüIC' Additional s ecimens South Australia: Hindmarsh Falls, ió, t8.xi.L97O. Australian CaPitaI (r.E.B. Common) Territory: B1ack Mountain I d , (at tight) , 13.xii. 1960 ' DescriPtions of other members of this genus aïe poor so only tentative (see rV'6'3) suggestions can be made about the affini tie s of australiensis 'The the presence of tarsal spines is unusual and may indicaie rel'ationship to

Canadi analbiplumus(Saether]9T2).Ho\^/ever,sincemostworkershaveuseddried specimens, these spines may have been overlooked in other species'

Genus STICIOCI,ADIUS Edwards sri ctocrad.ius Edwards, 1g3r, p.279 (as subgenus of spaniotomS) i Freeman, 1961. p.640 (as subgenus of Diploclad ius)

DipIoc ladius (rieffer) Brundin, 1956, P'70 Type species Spaniotoma (stictoc ladius) pulchripennis Ed,wards species of Edwards (1931) described from South America three new Diplocladius SparriotomaEdw.whichheseparatedfromtheEuropeansub-gen us placed them in a new sub- because of their patterned wings and banded legs' He differences in genus stictoctadius. Brundin (L966r PP.428,433) recognised basic and Sti-qlog-lg1:iuq * the arrangement of the leg sheaths of the pupae of Diplo-clad:ll¡s IV con- Tlhe arralysis of the phylogeny of these groups as presented in section firms their status as separate genera' referring r have redescribed two Australian species of this genus after the additional to the type series l'reld at sPHTM and to permanent mounts of similar to each other and specimens listed. All the Australia¡r species are very others (see r-v'6'4) are probably closer to the New Zealand species than to any ' STICTOCLADIUS KEY TO THB MATES OF AUSTRALIAN SPECIES OF THE GENUS (after Freema¡¡ L967, P'641) anteriorly, I. Dorsocentral br:istles uniserial posteriorry as well as legs usually with some indication of pale band on tibiae uniserialis Freeman. 81

Dorsocentral bristles tri- or quadriserial posteriorly, legs usually al-I dark 2 pale 2 v,ring markings formed of 2 broad bands, neither containing areas (not reported from South Australia) " "' victoriensis Freeman Outer wing marking split into two by large pale spots in the cells ...multiserialis Freeman sri Iadi us uniserialis (Freeman) comb.nov- Diploclad ius uniserialis Freeman, 1961, P' 643, Ej-q. 6d. Separated.fromotherAustralianspeciesbyitspatternedlegsand distinctive shoe-shaped inner lobe of gonocoxite' pubescence MALE (4\. Head. Verticals and frontal sensilla absent, êYe uniform. cibarial pump very broad with deeply curved dorsal border' Ten- torium as in Fig. 20. Segment 3 of palp with a short tapered sensillum (11¡); palpal formula O.25 : O'5 : I z I'2 z 2'L; M'R' o'25' Antenna' A.R. o.g. Fragellomeres 2 and 3 with 2 smalr type c setae; last flagelro- D setae and mere slightly clubbed with several type C setae, numerous type 4 ring setae. Furro\^r extending to tip. Thorax. Antepronotum with 5 smal-l lateral setae. Humeral pits largei acrostichals minute; dorsocentrals uniserial, punctures conspicuous, beginning after pits; 2 pre-alars' punctures conspicuous; scutellars small, uniserial. Legq. L'R' O'74' A1I tilciae whitish in basal half; comb of rear tibia irregular, both spurs small, slender, half covered with accessory setae. t'ling. Length L'7 - present 2.O mm., Asp. R. 3.6- Squama bare, anal lobe small' Macrotrichia only on base of R; microÈrichia invisible, punctures minute, pigmented to form two Èransverse bands narrower than in the female. R,*, alwaYs sixth closer to R4*5i R4+S ending just befot" M3*4i costa produced one gentlY of the distance to wing tip; pfk slightly distal of r-m; Cu, dis- curvedi An ending under r-m. a¡É"m"g. Tergj-tes v¡ith setae tríbuted in lateral and medial strips and acrr¡ss posterior margin' Terminalia (Fi9. 2L) - AnaI point absent' Sternapodeme broad''corrìelîs 82

phallapodeme treavily drawn up and strongly reinforced, articutation with rnner lobe sclerotised; phallapodeme with narrow plates; penapodeme absent. dorsal shoe-like in outline with anterior ventral 'heel' and posterior 'toe''; l-obe bul-i:ous' inner lobe of gonostyle slender, barerending in a slight knob; ot-1ter setose, with reduced sPine'

FEMALE 2). As for male but :- 5 flagellomeres: Head. Palpal formula 0.25 : 0.5 : 1 z l--2: 1.85' 'Antenna with setae' 2 4 1 and 2 with 2 small type C setae, 5 with 4 type C and 3 ring - slightlybroaderatmiddle,5tapereddistally.wi"q.Length]-.S9mm,Asp.R. 3.3, pattern as in Freeman 196I Fig' 6 d . Terminalia CFiS. 22) - Ste¡:nite vLtI VIII with about 25 setae along medial posterior border' Gonocoxite int'o poilt narro\¡/, heavily sclerotized; egg guide with long setae' extended gonocoxite IX distally. Tergite IX with medial cleft, oLrscuring tergite xi setose cone' reduced, with small proximat bald knob. st.ernite X with small projecting into segment cerci srightry exÈended ventralry. Vaginal apodeme not eminence broacl, VII , basalty one tenth as wide aS Segment viII. Spermathecaf ducts heavity setose, without basal broadening' Nov- 1922 [A'r" TypES. r have examined the holotype d , Mt. !{e1lington, Tas- Tonnoir) and paratype series, held by SPHTM' Reserve, 1ó ,8'iv', Additional specimens seen. South Australia: Eric Bonytlron ld , l.vii. I97o¡ Parawa, 19 (emergence trap) 12'viii'l-97O' Victoria: ¡Iayers Creek, ld , l8.viii.Lg64 (D'H' Colless) '

Stictocladi us multiserialis (Freeman) comb' nov. Diplocladius muitiserialis Freeman, Lg6L, p'641, Figs' 5c' 6b' Separatedfromremainderofgenusbyitsmultiserialdorsocentral posterior lobes of bristles. Female separated from uniseralie by its large

tergite l-X .

MALES (3). Similar to uniseuäIis but Head.PatpalforrnulaO.25:0.5:1:1:1.5.M.R.0.39.A¡¡l_ennL"A.R.1.0, rvith 15 20 terminal segment evenly roundec distal1y. Thorax' Antepronotum - beginning after humeral lateral setae. Acrostichals absent; dorsocentrals 83 pits, bíserial initially, expanding to multiserial; 10 - 20 pre-alars; scutellars numerous, fine, scattered. Legs. L.R.0.67. Wing. Length 2.2 nm., Asp. R. 3.38; microtrichia pigrmented, forming 3 transverse bands, narrov¡er than in female. Terminalia. (Freeman 1961, Fig.5b) Phallapodeme hooked posteriorly, plates tríangular. Inner lobe narrow parallel-sided, directed posteriorly; gonostyle as in uniserialis.

FEMALE (1). Like male' but :-

Head. Palpal segment proportions 0.25 : 0.5 : I z L.2 = L.4. AAtenna with 5 flagellomeresr l and 2 with 2 small type C setae,3 and 4 slightly flask-shaped, 5 with at least 3 tlpe C setae, 3 ring setae and a terminal bristle; ântennal

Vling. Length 2.1 mm. Asp- R. 3-26, formula 1.67 : I : I.L1 : I.33 z 2.L. ' pattern as in Freeman 1961 Fig. 6b. Terminalia. Sternite VIII with about 25 setae scattered along medialposterior border. Gonocoxite VIII narrow, egg guide smaller than in unise:zÞ_Iie. Tergite IX drawn out on each side of medial cleft into narrow bulb extending further than cerci, completely obscuring tergite X. Cerci narro\¡r, extending further ventrally than in uniåerälis.

Vaginal apodeme projecting one quarter into segrnent VII, basally one seventh as wide as segfnent VIII. Spermathecal ducts without basal broadening, area of spermathecal eminence larger than in uniserialis.

TYPE_S. I have examined the holotype d of Manly, N.S.Vü. June 1946, (O.t¡.

Colless) arrd the paratype series held by SPHTM. Additional ctmens seen. South Australia: Eric Bonython Reserve, ló 8"iv. Ig7O, ld , 1.vii.L97Oi Parawa, Id (at lightl 27.v.1970' (7 A single incomplete male specimen from lrlalpole, Vlestern Australia .x-L97O ' D.H. Colless) has uniserial dorsocentrafs, an inner lobe sirnilar to multiserialis and a style with a broader dorsal lobe but narrower ventral lobe than either uniserialis or muttiserialis; it must be considered a separate species but a complete description rnust av¡ait further material . Fig. 24 Eul

Fig. 25 tl apex of antennar C-typeCsetae; R - ring setaei S - sunken setae.

Fig. 26 wrng.

t¡ Fig. 27 lr terminalia (right hand side of tergite IX removed).

(view Fig. 28 femal.e, terminalia somewhat ventro-lateraI)

rig. 29 il il pupa, thoracic horn and precorneal setae.

FiS. 30 I abdomen, armature of right hand side of tergite III.

Fig. 3I I ll ventral (I.h.s.) and dorsal (r.h.s.) views of segment VIII and anal lobes. 24

25 t i ,.1 a I

I i r,tl

(/ 'iíi U

29 30 31 84

Genus EUKIE!-FERIELLA Thienemann Freeman L96L, P'643' Eukiefferiella Thienemann 1926 P'325¡

Nanoc Iadius Freeman L956, P'388' icalcar Potthast, I9l5 Ty'pe sPecres DactY Iocladirrs

Freeman Euki efferiella insolida (Skuse) p'644' Fi9' 7a' Eukieff eriella insotida Freeman, Lg6l-, 1889 Orthocladi US insolida Skuse' ' P'258' costa Àlightbrownspecieswithbroadtwistedtiponlastflagellomere, produced,*4*5"}'o'tened.Pupawith2groupsof3-Tstrongorallydirected IV' IV - V and V - VÏ' hooks on each of intersegiments III - Thesingletypeofthisspeciesissomewhatdamagedandtheterminalia havebeenlost;thisredescriptionhasthereforebeenmadewiththeaidof

specimens from South Australia' postorbitals and frontal sensilla absent' Segme-nt MALE ( 3). Head. Verticals, Palpa1 3ofpalpstoutwith3slendercapitellatesensillainSeparatepits. M'R' O'26t tentorium as in Fig' 24' formula 0.55 : o'55 : I z i.'25 z L'75 ' (15¡r); last 2 and 3 each with 2 type C setae Antenna. A.R. 0.75. Flageltometes flagellomereapicallyclubbedandtwistedandwithnumerousslendertypeDsetae (L2ll)(Fis.25).Thorax.Antepronotallobesjustcontiguousfromdorsalvierv, withoutlateralseÈae.Acrostichalsminute,dorsocentralssmallanteriorly, largerposteriorly,beginninglevelwithhumeralpits;4scutellars.LegS. least half covered v¡ith accessory setae' L.R. 0.8. Spurs of rear tibiae at mm' Asp' R' A'Ll' (Fis- 261 Length t'5 - 1'7 comb of stout setae' vüings ' smalr, basar vein with singre macrotrichia- Squama with 5 setae; anar robe with it in costa' R4*5 turned up less merged with R¿+5 and ending R Z+3 more or attip,finishingjustbeforelevelofM'*2lcostaslightlyproduced;pfkending Abdomeg' Tergites densely just distal of r-m, An ending at level of r-m' v'¡ith (FiS. 27). Anal point absent' sternapodeme uniformlY setose Termina lia broadtransversealm,cornersreinforcedandextendedintopoints.Phallapodene robe lightly sclerotised, penapodene absenl-' wíth smalr elongate prate, aedeagar inner sclerotised' gonocoxites wit-htwo Membrane at junction of gonocoxites 85 lobes,onedorsal,tongue-Iiketheotherventral,smaller.Gonostyleslightly tapered distallY.

FEM.ALE (I)-. Like male but 3- Head..BrowthickenedrpalpalformulaO'40:0'60:1zL'422'r.'Antenna with5flagellomeres,land2stout,lastwithapexnarrow,drawnout;flagello- merest-5eachwithpairoftypeCsetae;5thwithnumeroustypeDsetae,2 Iargering-shapedsetaebasally,2sunkensetaeattip.Flagellomereforrnula O'55' Wings' Length 1'4mm' Asp' R' I.4: I: 1.05 : 1.3 z 2.L. Legs with L'R' few small setae' 6 - 8 with many' Terminalia' 2.9. Abdomen. Sternite 5 with a side' egg guide evenly (Fiq.2g). sternite VIII vrith about 8 setae on each rounded.TergitelXsmall,rounded,gonocoxitelXwithshortsetaeextended disÈallyhalfwaytoleveloftipsofcerci.SterniteXdrawnintobroadridge' cerciglobularfromdorsalaspect.Vaginalapodemeprojectingshortdistance intosegmentVfl,basallyoneseventhaswideassegmentVlll.Spermathecae ducts well before spermathecal more heavily sclerotized than ductsi 'uniting

opening. horn (25O - 30Ou) with slender distal PUPA ( 3). C rax. Thoracic portionl'T5timeslengthofswollenbase,spinulesabsent(Fig.29).Three pair 2/3 Lengl¿h of thoracíc horno precornear setae arranged in triangle, anterior 3rdshort.Twoanteriorandlposteriorante-pronotalsetae,each2/3l.engiLht ofthoracichorn;2anteriorand2posteriorshortscutalsetae.Abdo]nen. of enlarged spinerets (Fig'30) on Tergites rr - vrr with 4 or 5 irregurar rows of alally d'irected spines on posterior anterior border and 3 - 4 íxtegular rows tergites III & IV' IV & V and V & VI border. on each side of midline between Setae O1 - O5 presentr Dn small on is a group of strong orally directed hooks' lstsegment,becominglargeronsubsequentsegrmentsuntilonVlllitis Setae tt - present: L, and flattened and half length of thoracic horn. "4 L,situatedveryclosetogetheronsegmentslT-VII,allLsetaeclcseto vIIf with - t4 half length of posterior border in segment VIIIi segment 'z thoracichorn,LrandLrthickrspinelike'S'Lerniteswithseveralrowsof spineletsonposteriorborder,setaeVt-u¿present.Anallobes(Fj-9.31)wittr Iateral posterior borclers curved towards ventral surface where there i1"". to," o' 3 strong curved setae, as long as the lobes themselves. TypES. I have exarnined the holotype ó, Middle Harbour Sydney (MM). Additional specimens seen. South Australia. Eric Bonython Reserve ód99, May - Dec., Lg7O, L97I¡ Hinclmarsh p¿115 ód99 d April - Dec. Lg6g, LglO, L97L (some reared from algae) ; lnman Valley I r 3.ix.L969¡ Marne River, 2ô,28.ix.1969; Parawa, Ió, f.ix.1970, ld L2.vií., 2ó 9.ix., Id l7.ix.1971 (at light); Second Vat1ey, Id , 30.vii.L969, Iô 2o.viii.1969; sturt River, ld , 4-ix.1969.

The structure of the newly described pupa confirms Freeman's (1961) designation of insolida as a member of the brevicalcar group of Eukiefferiella. It is most simiLar in both adult and pupa to the European member B. stJ,I:L:þr Goetgh.

Genus SYNORTHOCLADIUS Thienemann Svnorthocladius Thieneman 1935 p.2LI. Synorthocladius, Brundin 1956 p.91. Type species Synorthocladius semivirens (Kieff.)

S orthocladius br:unnen sp . nov.

Separated from other Synorthocladius species by shape of gonostyle and presence of penapodeme in male and by presence of prothoracic horn a¡d scutal microsculpture in the pupa. All S)morthocladius species are distinguished by the presence of 2 tiny flattened setae high over the scutum of the adult. L/3 MALE ( 13) . Head. Post orbitals present, verticals of medium size reachíng dista¡ce to nridline. Frontal tubercles small. IuI.R. o.25. segrnent 3 of palp rvit-h several tapered sensilla (L2lt). Palpalformula 0.4 : 0.5 : I t L.2 z L.7. Nelno" A.R. 0.6.Flagellomeres 2 and 3 with 3 type C setae (25y)i last flagellomere slightly clubbed distally with 2 tlzpe C setae (15p), numerous type D setae (I57:l and 3 ring setae. Thorax. Antepronotal lobes only just contigrrous; 6 lateraf setae. Dorsocentrals begilning after humeral pits; 3 pre-a1ars, 4 scutellars' wing_. Length L.2 - 1.65 mm. Asp. R. (2) 4.L7. Squama with 3 - I fine, nedium Iength setae, anal l.obe small. Macrotrichia on Rl, R2*3 ending close ao *4*5, Fig. 32 Synorthocladius brunnen, sp. nov. male, terminalia.

Fig. 33 tt il female, terminalia

rig 34 |l I pupar thoracic horn.

rig. 35 I anal lobes

Fig. 36 Paratrichocladius luriserialis Freeman male, tentorium.

Fig. 37 tl " thorax. aaII- anterior anepi- sternum II ae II anepimeron II ap antepronotum as anapleural suture cI-III - coxae eIII episternum III h humeral pit ha haltere malI median anepi- sternum If palI posteríor anepi- sternum Pe pre-episternum pn postnotum s- scutum sc scutellum slrs2 - anterior & posterior spiracles. wb wing base. 32 ,:.,'... \ I t ;3

! ( u ìr* ¡ g

u lJ

(/ 3s

.o O o o oO

a o

Ca.

33 U 37

òl ù \U ù {

34 36 87 *4*5 closer ao *l*2rcosta only very slightly produced' Cu, bent evenly "rding downwards. An ending distal of r-m. Legs. L.R. 0.6, rear tibia slightly flared distally, comb strong, accessory setae on inner spur only. Abdomen. Tergites each with two irregular transverse rows of setae. Terminal-ia (Fig.32)' Anal point bare, of variable síze, sternapodeme evenly arched. Phallapodeme tríangular. p.R. 0.18. Inner lobe broad-based, finishing in right-angled bare corner. Gonostyle with bare preterminal hump'

FEMAIE 4 . Like male but 2- HeaÈ. verticals absent. segment 3 of palp with one sens-illum. Palpal formula 0.3 : 0.5 : I : 1.35 z 2.O4. Antenna with 5 flagelÌomeres. AIl flagellomeres with 2 type C setae (18F - 28P). Ftagellomere formuta 1'4 : 1 : 1'1 z L'2 : 2'3'

Vfling. Length I.27 - I.33- AsP. R . 3.1. Terminalia (FiS.33). Sternite VIII with row of 5 - B setae on each sidergonocoxite VIII narrow, heavily sclerot-ized' egg guides'Iarge, rounded. TergiÈe IX without med.ial cleft, noÈ obscuring tergite X , gonocoxite fX extended distally, knob-Iike' Sternite X with broad setose swelling' cerci somewhat elongate. Vaginal apodeme basally one quarter as wide as segment VIII , extending half its length into segment vII spermathecal du':ts with basal arrow-shaped broadening.

PUPA (r). cephalothorax (Fíg.34). Thoracic horn (29Op) slender, slightly

expanded distallyr covered with small points. Three precorneal setae 90p, 100]r' 50¡ respectively. one anterior antepronotal seta (80¡), four scutal setae (40p)' (Fig'35)' Scutum and vling covers covered with small protuberances. Abdomen Tergites with srnall spinelets, concentrated along posterior border ancl enlarged into a row of heavily sclerotize

TypES. Holotypeó an<1 ld paratype, Eric Bonython Reserve South Austral-ia' g.ix.:-:gTo.Ftrrtherparatypes,EricBonythonReserveld,IT.ll.,ld,S.iv.,

26 , l.vii., 2d , f .ix., 3d , 23.ix.1970, 1ó (reared from detrit-us on submerged 8B upstream rockface) 16.xi.I97O, 1d 3I.iii.1969; Hindmarsh Fatls 3d 25. iii .r910, 2 ó, 8.ix.I97I. Addítional cimens seen. South Australia. Botanic Gardens, 1ó 10.iii.I91L; Eric Bonython Reservei t5ó , I8.iv., 1d , 22.iv., 4ó , 2O-v., l.4ô , 28.vii., ld,23.viii.,5ó, f.ix.,5d, 18.xi.,3d ,2.xii.r97oì 2Lô, 13.vii.,3d , 26.viii., fOd , 8.ix., tBó , 6.x., 126, 3O.xi.I97I; Hindmarsh ralls dd Feb.r

April, June - Dec . Lg7o, 6ó , 3O.vi., tô zø.viii., 9d , 6.x. L97L¡ Parawa Id (reared from algae) 8.ix.I97I.

The presence of a prothoracic horn and exterrsive microsculpture in the pupal stage is atypical of the genus Synorthocladius. The discovery of further species with these characteristics may h¡arrant the construction of a new genus.

Genus PARATRTCHOCLADIUS Santos Abreu

Paratrichocladius Santos AbreuI9IS; Hirvenoja, 1973 p.88 Syncricotopus Brundin L956, p.106. Freeman L96L, p.645.

I}pe spe cies Chironomus rufiventris Meigen, 1830' sensu

Edwards, 1929. The following modifications and additions to descriptions of this genus provided by Brundin (1956) and Hirvenoja (f973) are made on the basis of my examination of Australian species.

MALE. Head. Post orbitals and verticals strong, extend to midline where 2 setae are closely adjacent; frontal sensilla absenti segment 3 of palp with broad Èapered sensilla in individual pits on slightly clilated inner distal surface; tentorium, see Fig. 36. Antenle. A.R. about I.25¡ flagellomeres 2 arrd 3 each with 3 sl-ender type C setae; Iast flagelIomere only slightly clubbed. Thorax_. Antepronotum with numerous lateral setae. Dorsocentrals beginning after humeral pits, punctures conspicuous. Legs. L.R. 0.6 - O.7¡ rear tibia

expanded distaltyi inner spur sinuous, with basal accessoriesi comb with strong setae. Wings. AnaI lobe large; microtrichia of variable sizet R2*3 ending midway between R, and R4*5r *4*5 .rding midway between M1*2 .td M3*4, cu' slightlv curved; An ending weII distal of r-m. Abdomen. Tergites with setae in two 89 broad rows, sternites L - 4 irregularly sclerotised, 5 - I uniformly setose and heavily sclerotised. Terminalia. Stern apodeme with broad lateral arms, corners drawn up into knobs and reinforced. Penapodeme absent. FEMALE. As for male but :- Antenna with 6 flagellomeres. Setae of abdominal sternites restricted to a medium strip. Egg guide broad, crescent shaped. Terminatia. Tergite IX not obscuring X . Gonocoxite IX reduced, cerci with ventral portion elongated anteriorly. Vaginal apodeme not expand.ed basally, not projecting into segrment

VII . Spermathecal ducts more heavily sclerotized than in any other genus I arranged in s- shape, without basal arrow-head expansions.

Paratrichocfadius pluriserialis (Freeman) comb.nov.

Syncricotopus pluriserialis Freeman 196I, p.645- Svncricotopus pluriserialis Fors yth 1971, p.119. Trichocladius pIuriserialis Freeman 1959, P.414- A dark species, separated. from the African micans Kieffer by its multiserial dorsocentral bristles and from bífinestrus by its single large h.umeral pit and exceptionally Iarge cerci in the female.

MALE (4). Head. Eyes with dorsal elongaÈion 4 facets wide; palpal formula 0.5 : 0.5 : 1 : 1.5 : 1.8. M.R. 0.3. Antenna. Flagellomeres 2 - 5 broader than longi last flageltomere with 7 - I type C setae and one ring seta with círc1et, ending in blunt point. Thorax (Fig.37). Acrostichals numerous, biserial; humeral pits largerdorsocentrals irregularly biserial in South Australian specimens. Four pre-alars, 12 scutellars in single row' !'ling' Length L.4 - 2.Omm., AsP. R. 4.O9. Squama with 11 strong setae, anal lobe exaggerated; macrotrichia present on basal vein, R and Rr' Microtrichia invisible, punctures plain; cosÈa not producecl; pfk at level of r-m' Abdomen' Sternites fII and IV with one posterior medial seta. Terminalia. Tergite IX with median posterior indentation; phatlapodeme with oval plates; inner lobe triangular, point curved posteriorly. Distal inner surface of gonostyle drawn ventrally into rounded setose ridge.

FEMALE (3). Head. Verticals absent except for single seta near midline; Fig. 38 Paratrichocladius pluriserialis (Freeman) female, terminalia, ventral (1.h.s.) and dorsal (r.h. s. ) views.

il FÍ9. 39 I pupar proÈhoracic horn.

Fig. 40 tl pupa, dorsal view (r.h. s. ) tergite III.

rig. 4L Paratrichocladius bifinestrus sp. nov. male, wing'

It il F.ig. 42 tt terminalia (half tergite IX removed) . 39 0o o @o o I

\ U l'-t

I )

40 42 \ I I I t'f I tl ìl' ,r , I tl 90

indented; eyes witir cibarial pump with dorsal border deeply and rectangularly :1.I : L.7. dorsal elongation 2 facets wide; palpal formula 0.5 : 0.5 : l Antenna.TypeCsetaepresent,lonflagellomerel,2on2,3on4a:rd5'6 onlastflagellomere;antennalformulal'I:I:I'1:1¿2'O'Thorax'Only (r'ig.38) . Sternite 3 pre-alars. Wing Length 2'2mm' Asp' R' 3'4' Terminalia plate; vIIl with 1I setae on each side; sternite X with rounded' setose spermathecae drop-shaped, basally spinulose' PUPA(3).Cephalothorêä.Thoracichornspherical,3precornealsetae(].20p)set 2 and 2 posterior antepro- above small wart-tike protuberance (Fig.39) ' a¡lterior (70p) Abdomen' notal setae (7OF), I anterior and 3 posterior scutal setae ' tergite as in spineleÈs arranged in Trich,ocladius pattern (Thienemann' L944) ' rig. O;spineleÈslargeonsegmentslll-Vl,setaeL,andLnenlargedonseg- spines, somewhat longer ments vrrr and rl. Anat lobes with 3 slightry curved than the final segment (Forsyth, L97L' Eig'4 (S))' the series from the TypEq. I have not examined the type specimen but have seen eastern states identified by Freeman (1961) ' lld 29 3O'vii'1969; Additional sÞec imens seen. South Australia: Alnut' ' ' ló 28.i', ld , 2o'v'I970; Eric Bonython Reserrre' 1ó Botanic Gardens, ' ' 22'íx" 3d 19 30'xi'r971; 2g.vii., ld , 2.xíL.rglo, 5ó 3r.iii" 3d , 29 ' ' ', Parawa, 7d . I9.vii" Ió , 2o'viii' Marne River, Id , 9.v::i-., 2ó , 9.X. L969¡ ', (at light) Id (at light) ld , L9.íx.I969, 2ô , 26.v., 2ó , L2.v.t97O; Id 'L7.íj-. ' ld 19 , 7'viii'1969 IO.iii., Ió , 6.x.lg7¡¡ Q 9ð , 3O.vii.Lg6g; Murray Bridge, ' (T'C'R'Vfhi'te) (8. Glover); Keith, 1ó (aerial plankton sampte at lO5m') Nov '1964 ' (J. Martin); Stratford' Victoria: ¡4elbourne Botanic Gardens ód 99 , 15.iv.1963 tó , 28.x.L96L (D.H- colless) ' AdultsofthisspeciesweredescribedfromNewZeal.and(1959)and Australia(197I)byFreeman.Forsyth(197I)hasdescribedthehabitat'eçJgmaSS' larvaandbriefly,thepupaoftheNewZealandform.TheNewZealandpupa rslender' thoracic horns and' differs from thesouth Australian one in having Zealand specimens as "claspette while Freeman describes both Austral'ian and New similartomicarjs,,,Australianspecirnens,particularlythoseinSouth dclaspette', it i-s possible that the Australia tend to have a more poin¡s¿ so 9l adultsactuallyrepresenttwoseparatespecies.Ifnot,thedivergenceheremay provide an interesting problem in cytogenetics' TheaduttsandatleastÈheAustralianformofthepupaindicatethat of the genus; it maY lie pluriserialis lies close to rufiventus the type equally close to the African mícans'

Paratrichocladius bifines trus sP. nov.

Immediatelydistinguishedfromallotherorthocladiinaelhaveseenby cuticle which form the humeral pits' similar Elne 2 large ovals of unsclerotised tomicansinformationofinnerlobeandgonostylebutdifferinginhaving multi-serial dorsocentral bristles'

MÀLE (3). Head.Eyeswithdorsalelongation5facetswide;dorsalborderof indentation; palpal formula' o'3 : 0'5 : cibarial pump with shallow rectangular ending in short point with I : 1.2: I.90. Antenna' Last flagellomere D setae, 4 rLng setae and 2 sunken setae' severar type c setae and numerous type Thorax.Acrostichalsfewinnumber,humeralpitsconsistingof2Largeovalsi dorsocentrarsbi-triserial;3pre-ararsiscuterlarsnumerousrfine'inan irregularpattern.Wing.(Fig.al)Length2.Oorun.,AsP.R.4.42.Squamawith numerousshortfinesetae,analloberight-angled;macrotrichiapresenton plain' microtrichia smallt R4+5 basal vein and R; punctures of microtrichia endingclosertoleve]of*3*4;costaproducedL/5ofd,istancetowingtip;pfk .lig.42). Phallapodeme with bent very slightlY distal to r-m' Termina lia crescent-shapedplatesiinnerlobewithsinuousposteriormargínandrectangular baremedialknob'gonostylewithconcaveinnersurfaceandstrongterminalspine. to midline; cibarial PumP with shallow FEMALE 2 Head. Verticals extend of eyes 4 facets wide; segment 3 of curved dorsal border; dorsal elongation O'3 : 0'5 : I z L.2 2.O. Antenna. palp with I sensilla (I8p)' palpal formula C setae p}¡t) t last flagellomere with Flagellomeres I - 5 each with 2 type z 2'6' formula l-' 4 : I r. 15 l-.15 z I-29 numerous type D setae (381¡) i antennal ' macrotrichia on R' Rt and R : coSta Vüing. Length 2.O5mm' Aspect Ratio 3'5; 4+5' tip' pfk at level of r-m' Terminalia, produced one quarter of distance to wing irr gonocoxite IX slightly smaller than Sternite I with I setae on eactr side' 92

in pluriseràlis-' r¡luriseräIisr sternite X flat,setose. cerci smaller than spermathecae almost twice as long as broad' 22.ív.1970' Types. Holotypsó, Eric Bonython Reserve, south Australia sB'iv'J-971-' Paratypes.Eric Bonython Reserve, 29 ' Bonython Reserve' 1ó Additional specimen S SEEN: South Australia: Eric ' 2'i'x'L97o¡ ró 4'ii" 25.ii.,ró, f.iv., 39, 22-ív-, ró,28'vii' ' Ló ' ' Falls' ló, I7.ii., 2ó, 28.iv-, Ló, 2'v', 2ô' 19' 3o'xi'L971; Hindmarsh Ió , 22.íx.I971. AsingleSpecimen(probablyrepresentingathirdspecies)caughtbyM. Mendumon2L.vii.lg6gatBullitaHomestead,NorthernTerritory'16'075'I30'26Il'! :- differs from bifinestrus in the following details 3 of palp with Head. ïnner plate v¡ith evenly scooped dorsaf border, segment only2sensilla.Wing.Asp.R.4.60,puncturesofmicrotrichiaminute, with broader plates' *4*5 .tding at level of M3+4. Terminalia. Phallapodeme the tendency towards The structure of the inner lobe of bifinestrus and related specimen smaller microtrichia on the wing membrane in the closely indicateaffinitiesoftheAustralianfaunawiththeAfricanmicans.

Genus CRICOTOPUS van der $lulP L973'p'13I' CrL cotopus van der Walp 1874tp'L32; Hirveno)a

Tlpe sPecres Chironomus tibialis Meigen, 1818,

Sub-genus CRÍCOTOPUS s'str' Hirvenoja

cricotopus (cricotopllq) Hirvenoja L973'p'135' the basis of species Hirvenoja (1973) revised the genus Cricotopus and on Cricoto pus s.str. and from the West Palearctic, divided it into two suJrgenera' s.str.. On the basis Isocladius. AII Australian species belong in Cri.c into a number of groups' of afl 3 life stages Hirvenoja divided crico-qs€us s'str' itarsis fits well- Of the 5 species from South Australia described below' alb arbicinctus into his festivellus group. The remaining four species' together tith Freeman, aeosomatus annuliventris and hirtell.¿s "fbitibi" group' are all very similar and are close to the bicinctus 93 Rather than adapt Hirvenoja's keys I have constructed a single key to aII Cric species described from Australia. It is based on descriptions of adults provided below and by Freeman (196I) and on descriptions of larvae and pupae provided below and by D.H. Edward (1961) '

KEYToTHEAUSTRALIANSPECIESoFTHEGENUScRIcoToPUS

MALES

1. 'Tibia, tarsi and a-bdomen without pale markings' A.R. greater than 1.5 phaeosomatus sP.nov. Tibia, tarsi and abdomen with pale markings' A.R. l-ess than 1.5 2 2. (1) Tibia and tarsi completely without dark pigment' abdominal segments I, TV pale; hump on gono- nov. style absent albitarsis ' sp. Tj-biae with pale bands; hump on gonostyle present"" 3 3(2) All tibia with Pale bands 4 Anteriorandmiddletibiaewithpaleband.sinbasal

half, abdomen with pale markings on segments I' (Skuse) IV and V . annuliventris

5 4 (3) All tibia with broad pale barrds

Rear tibia with narrower pale band, hump on gonostyle large hirtellus sp. nov.

5 (4) Inner lobe of gonocoxite narrow, segments II (walker) and IIï of abdomen without pale markings albitibia Inner lobe of gonocoxite broad, segrments II sP.nov- and III of abdomen with pale markings oarbicínctusr

Known

1. Prothoracic horn sfender . " ' 2 Prothoracic horn bulbous, pedes spurii B on (vlalker) segments II & fII of abdomen albitibia

2 (1) Pedes spurii B on segments 1I and III of annuliverrtris (Skuse) abdomen Fig. 43 Cricotopus albitarsis sp. nov. male, tergites fII & IV .

Fig. 44 il terminalia.

Fig.45 tt tt female, tergites III & IV.

Fig. 46 Crícotopus parbicinctus, sp. nov. male, tergites III & IV.

Fig. 47 tl terminalia.

Fig. 48 I female, antenna.

Fig. 49 ll tergites IIf 6( IV. 43 45

\ 44 48

46 c c

49 t ¡

lñ?rr, Jfl, r,r,,,, t, lt h I l.' ttt '47 \ :il 94

parbicinctusr sP.nov' Pedes sPurii B on segment II onlY

Kn own Larvae 2 I. Mandible with serrations on inner surface albitibia (InIal.ker) Mandible without serrations on inner surface 2(L) Mandible with wrinkl-ed outer surface' more than parbicinctus sP.nov. than 3 serrations on inner surface only Mandible with smooth outer surface ' (Skuse) 3 serrations on inner surface annuliventris

cricotopus ( Crico us) albitarsis sp.nov. Australian species by Close to festivellus Kieffer, separated from other complete lack of leg pigirnentation '

MAËE (Z .Head.Verticalsetaeandfrontalsensillaabsent.Byeswithdo:lsal single sensillum (ra¡i¡ in pit, elongation 3 facets wide. Segment 3 of palp with A'R' I'35 - t'55' palpal formula O.4 : O'5 : I : 1'35 : I'5' M'R' O'3' Anten¡e' Flagellomeres2-4with3tlzpeCsetae(28p).Lastflagellomereslightly clubbeddistallyr\^/ithnumerousDsetae'3ringsetae'Thorax'4prealars' yellovl . wJ'g. Length scutellars urriserial . LegS. I,.R. o.57, all legs pale lobe large' Macrotrichia 1.53mm., Asp. R. 3.9. Squama with 4 small setae, anal ending leve1 with Mr*n, Pft ends almost under r-m' on basal vein & R. 1*S 4Þ9o*egTergiteslandlVpaleyellow,remainderbrown.TergiteslllandlV (Fig'43) with u¡riserial lateral setae, posterior cross-row reduced ' corners with large points' Te rninalia (îLg.AiJ. Sternapodeme evenly arched' rounded point with very small phatlapodemes small. Inner lobe drawn into bare, spine small' ventral secondary lobe. Gonostyle without distal hump'

FEM.ALE (I). Similar to mafe but :- Preapj-cal seta Head. PaIpaI formul-a O'4 : O'5 : l: I'3 z 2'3' Arrtenna' small,formulal.85:lzI'L4zL'L4:1'1422'7'Abdomen'Tergj'teIIf IX mo

il Fig. 51 il pupar prothoracic horn.

lt Fig. 52 ! pedes spurii B. rig.53 tt It /t' tergite VIII and anal lobes.

lt Fig. 54 n Iarva, hypochilum

Fig. 55 I mandíbIe

Fig. 56 cricotopus phaeosomatus sp. nov. male' wing

It tl & IV. Fig. 57 tergites III 50 I {

I t I \ I l

L-

53 51

54 52

t( ì (( ,,t(( tt ,ll 56 t tì ''1,,t' -¡ ( ¡l I

57 95

Typ_ES. Holotyped and Id , 1? paratype, Torrens Lake, Adelaide, I0.ii.1971. Additional imens seen. Blewitt Springs , !ó , 19 , 22'ix'1971; Hindmarsh Fa1ls, L61¡.iv.Ig71i parawa, Lô ,8.ix.L91I¡ Torrens, I7ð , 39 ,10.ii., 1d, 24.ii., 3ó , 3.iii., Id , 25'iii'1971'

CricggoPus (Cricotopus) parbicinctus sp' nov' very similar to bicinctus and atbitibia but male with broader inner lobe from on gonocoxite and hump on gonostyle; pupa with pedes spurii B absent and more than 3 segment III; Iarva with smaller medial tooth on hypochilum serrations on inner margin of mandible' MÄLB(6).Head.Verticalsextendtomidline,eyeswithdorsalelongation5 facets wide. segrment 3 of palp with 3-4 clubbed sensilla, patpal formula type O.4 : 0.5 : I ¿ L.2 z 2.O. Antenna. A.R. L.4, flagellomeres 2-4 with 3 type D c setaer last flagellomere slightly clubbed distally, with numerous all setae. Thorax. Four prealars, scutellars uniserial' Legs' L'R 0'75' tibiae with pale bãnds. v,ring. Length 2.o - 2.5 mm. Asp. R. 4'l. - 4'4' Squama R' with 10-15 long setae, anal lobe disÈinct. Macrotrichia on basal vein and of *4*5 .rding midway between ML+2, M3*4t pfk ending s1i9ht1y distal r-m' Abdomen. Tergite I, anterior parts of II' III' fV and V yeì-Iow in most of specimens. Tergites IIf and rv at least with lateral and posterior rows phalla- setae (rig. 46) . Terminalia. (Fig-47) Corners of sternapodeme large' gonostyle podeme large, contoured.. fnner lobe broad, terminal portion bare, breadth of with targe dorso-distal hump, spine small. specimens vary in the theinnerlobeandthesizeofthedorso-distalhump.

FEMALE Q\. Like male but :- (Fi9.48)' Head. PalPaI formula 0.25 : 0'5 I I.2 z 2.I. Antenna Single large preapical seta; flagellomere formlrla 1'8 : 1 : 1.1 : 1.1 2.5. Abdomen. not obscuríng Tergites IIf and IV as in Fig .49. Terminalia (Fig.50). Tergit IX sper- tergite X , medial cleft complete. Gonocoxite IX rnoderately extended'

mathecae with basal broadening, ducts almost striaght' horn slender (Fig'5I) PUPA Q\. CCPhA lothorax. Frontal setae absent, thoracic (Fig'52) with d.istal spinules. Abdomen. Pedes sprrrii B on segment II only ' 96

AnaI lobes see fig.53. LARVA(2). nypochilum (r'ig.54) with narrower medial tooth than bicinctus. Mandible with about 5 serrations on inner surface and r^rith wrinkled outer surface. Otherwise similar to bicinctus'

TypES. Holotypeó , Leg of Mutton Lake, Mt. Ganbier, 26-viii.1969. Paratypes: Eric Bonython Reserve, 1ó, 8.iv.I97O; Hindmarsh Falls, Ló , 19 , 4'xi'I970;

Inman Valley, ló (reared from a19a)r. 11.x. I97l-¡ Para\^ra,Ió r 9'ix'I97O, 1d (reared from alga) II.x-I97L. Additional specimens seen- Botanic Gardens, L6 , 2'x'L969, !ô ' 6'x'L97L' Id, I7.ii. L97I¡ Brownhill Creek, 2ó , 39 , 4ix. , Iô , 2O-viii.l969i Clarendon, Ló 19 Ló , 49 ,17.ix. 1969¡ Deep creek (Adelaide Hills) 3d , 49 , 4'ix' , , ' 7.ix.1969¡ Duck Ponds 1d, 3?, 28.ix.1969¡ Eric Bonython Reserve 1d, 22'íx'Ig6g' dd 99 ró, rA.L, ró, É ,8iv,3ó r lg, 2o.v.L97o,26, Lg, r7'iii'r97L, Aug - Dec, Lg6g-L97L; Fishers, 26 , L7.ix., 1d, 7.xi.L969¡ Greenhill Quarries 1d (¡eared from alga) 20viiil969; Hindmarsh Falls, ód99 Aug - Dec, 1969-71 qó, (many reared from alga); Inman Vatley,ddgg Aug - Nov 1969, Nietskes 19, (at light.) 9.viii. 1969¡ t'Iarne, 46 , 29 ,9.viii.1969; Moculta, Ló , 9.viii'1969¡ Morialta, 46 , 7.xi. L969¡ Parawa, 1ó , 22.!v., 2ô , f .ix. L97o, l9 , 26'v'J'g7J- , ód 99 sept - oct 197r (many at light and. reared from arga) ¡ Q,LQ , 3O.vii 'L969i ód99 Saunders Creek, I I , 25.x.I969¡ Second Valtey, Aug - oct 1969; Tanunda dd99 Creek, 3 d, 1? , 9.viii. 1969,5'e¡¡q¡t Feb. Lg7O, Lg7L,29 16.iv.Ig7O,

Se^ot - Dec 1970. Adult specimens attributed here to parbicinctus vary considerablY in degree of pigmentation and shape of the inner lobe and may represent a composite group.

Cric S (Cricotopus) phaeosomatus, sp.nov. Very similar to cinctus but A.R. much higher, no tibial strípes or

abdominal markings. absent' MALE (I). Head. Verticals short, extend to midline; frontat sensilla groups Eyes with dorsal elongation 2 facelLs wide. segment 3 of palp with 2 of 3slightlyclutrbedsensilla,palpalformulaO'25:0'5:1:0'9zI'2' Fig. 58 Cricotopus phaesomatus sp. nov- male, terminalia

Fig.59 Cricotopus annuliventris Skuse, ma1e, wing

il ll " tergites IIr & IV.

Fig. 6I n lt I terminalia.

'l tl tt f agello- Fis. 62 female antennal ' meres I and 2-

I I t! III e IV. Fig. 63 tergites rig. 64C ricotopus hirtellus sp.nov. ma1e, abdominal setation' I 62 ( t ût "t,,.

ço c o 58 C¿ gu - o a o gv a goo çt)

Ò o a, 6 o ô oê o

60 63

61 L¿ ( ç 59

// 97 (18P), last Antenna. A.R. L.87, flagellomeres 2 and 3 with type c setae flagellomere slightly clubbed at apex, with type D setae (34)r) ' Thorax' Four prealars, scutellars uniserial. Legs. L.R o.60 all legs dark brown' vüing' anal (Fig.56) Length 2.4mm,Asp.R.4.4. Squama with about 13 short stout setae' lobe large. Macrotrichia on basal vein and R' *4*5 fitishing closer to M1+2' pfkjustdistalofr-m.e¡a"*9.(Fig.57)Completelybrown. Terminalia. lober (Fig.58). Sternapodeme with small corners' gonocoxite with broad inner similar to parbicinctus, gonostyle with slight dorso-distal hump' TypE. Holotyped from Leg of Mutton Lake, Mt. Gambier viii.1969. gills) Additiona I specimens seen. Deep Creek (Adetaide 7ó , 19 , 4.Lx'l-969¡ Moriarta, rd , 4.ix.1969; Nietske's (at light) 9d , 9'viii'1969'

Sttc-s< 7 Cricotopus annuliventris Freeman

Cricotopu s annuliventris, Freeman 196 L, p.oqo, Orthocladius annul-i ventris Skuse, 1889, P.255. Separated from other Cricotopus species by pale markings on anterior and' middle tibiae, hurnp on gonostyle large' MALEc).Head.Verticalsrfrontalsensilla'absent;eyeswittr-dorsalelonqa- (18F) palpal tion 4 facets wide. Segment 3 of patp with 7 - I sensilla ' 3-5 formula 0.4 : O-5 : 1 : 1'l- : L.7. Antenna. A.R- I.O Flagellomeres Legs' with3 type C setae. Thorax. Four prealars, scutellars uniserial' (Fis'59) Length L.R 0.65 First and second tibiae with yellow bands. wigg' (Fi9'60)' l-.8 - 2.O mm, Asp. R. 4.L. Squama with about 6 short setae' Abdomen. fateral ro¡¿¿ Tergites I, base II, Tv and v yellow. Tergites III and IV with (Fig.6I) Gonocoxite with and incomplete posterior row of setae' Terminalia. broad rounded in'er robe. Gonostyle with rarge dorsodistal hump.

FEMALE (I) Like mare but Erq'62' Head. PaIPal formrrla 0.35 : 0'5 : 1 : 1.3 : 2.2. Antenna. As in j Gonocoxj-te IX larger Þ.bdomen Tergites fIl and IV as in Fig'63' Terminal a. narror^ter' ttran in P-eLrbicilgtus, basal broadening of spermatliecae I6'ivrdd 99 Sept' - Dec. 1970; Additional s cirnens s een. Botanic Gardens , Iê , 98

Brownhill creek, ã , 7.x!.I969¡ Deep Creek (Adelaide Hills) 7 ô , 69 , 17.ix. 1969¡ Eric Bonython Reserve, 1ó , It.ii, 1d 2O.u.t 1d , 28.vLL, 1d , 9.ix. Lg7o, 4 ó, 7.iv, 4 ð, LA.x.197ri Fishers, ú 17.ix, 1d , 1? , 1'x'1969; Glenview, 3d 9.viii.1969¡ Hindmarsh Fall-s, ld , l.vii., 29 , 23.íx., 2ô , A.xi.1970' Id , 2L.íx.L97I¡ Nietske's, 1ó , 25.x.L969¡ Parawa, Ló , 2O.viii.Lg6g,ld , 19 , 2g.i,, 2ó , 2.xii.:Ig7O, Ló , I2.viií, ld , 29 , 2I.ix.I97L¡ Scott's Creek Id , 4ixL969; Springton, Id r 9.viii.1969; Torrens (at lightl 7ó,79 ,14.x-L97I.

Cri (Cricotopus) hirtell-us sp. nov.

Separ ated from parbicinctus by the smaller pale bands of the rear tibia, Iack of pale markings on segment II of abdomen and nìore extensive abdominal setation.

MALE (I). Head. Verticals small, extend. to midline, frontal sensilla absent

Eyes with dorsal elongation 3 facets wide. segment 3 of palp with group of tapered sensilla, palpal formula 0.4 : 0'6 :1 : L'2:1'9' M'R' 0'30' Antenna A.R. l.30.Flagellomeres without type C setae.Last flagellomere slightly cltrbbed at apex. Thorax. Three prealars, scutellars biserial. lving. Length 2'3mm'

Asp. R. 4.1. squama with numerous setae, anal lobe Iarge. Macrotrichia on basal vein and Rr. R4*5 .td" midway between MI+2, tg*¿ipfk ends not far distal of r-m. Legs. L.R. 0.6. Anterior and middl-e tibiaewith distinct yellow ring; ring on rear tibia less distinct. Abdomen. (Fig.64) ' Tergites with dense lateral setae and 2 sparse transverse rov¡s; tergite I, base II, IV and base V yellow. Terminalia. Similar to parbicinctus but gonostyle with Iarger

hump. TypE. Holotype d from Hindmarsh Fallsr South Australia, 4.xj-.I91O'

Genus M]CROCRICOTOPUS Thienemann & Harnisch

Micror:ricotoPus Thienemann & Harnisch, L932 P.I37.

Nanocladius Freeman, 1956, P.338 (in Part)

Microcri cotopus, Brundin, L956, P.L2O lvlicrocrico Fittkau & Lehmann, l97O

f\zpe species M. bicolor ZetE. 1838 Fig. 65 Microcrico topus clavulus sp. nov. ma1e, facets of eye.

lt il Fig. 66 tt wing

tt il Fig. 67 abdominal tergites III, IV and V.

Fig. 68 I ll terminalia.

Fig. 69 il pirpa, head.

ll Fig. 70 I thoracic horn a¡d precorneal setae.

Fig. 7I tl il I abdominal segments II - VI, dorso- laÈeral víew. a

a rir . i'

i.j

:¡r ;.i ¡ d't a {: ç,1 t.,. lt,

,,.: :'.:

68 67

o 4 11 65 /l )r' 69

/ ,,.. I / 7 l\

ú)

70 71 99

Freeman (1956) proposed that, since the single female holotype

Nanocladi us vitellinus Kieff. is very similar to MicrocricotoPus sPecies' MicrocricotoPus must fall as a synonym' However' Fittkau & Lehmann (1970) otopus examined a male of tl. vitellinus and found it differed from aI1 Microcric species in having an hairy anal point and so the name Microcricotopus- has been retained. Fittkau and Lehmann also determined that only four species yet described are l¡us I4icroc ricotopus species; the Australian species makes a fifth' The key to Microcricotopus species devised by Fittkau and Lehmann can be amended in the following way to include M. clavulus' sp.nov.

Males. 3. Tergite setae of segments II - VII in a single transverse rovl

Tergite setae segments II - VIII in a nunrl¡er of irregular rowsrA.R. 0.7 - 1'0 M. rectinervis Ki-ef f . 4. A.R. 0.5 - 0-6. Abdominal tergites III - V with 2 lateral setae M. parvulus Kieff. A.R. about 0.7. Abdominal tergites IÌI - V with 3 lateral setae (rig'67) M. clavulus sP" nov.

Pup_ae 1. Thoracic horn narrow, tube-Iike, tapered distallY to a Point 4 4. Tergites V & VI with distinct medial regions of spines. Intersegmental membrane III - IV with reverse sPines M. bicolor Tergite V with very few medial spines, intersegmental

mernbrane III - IV and IV - V with reverse spines

(Fig. 71) M. clavul.us sp.nov.

Microcricotopus clavulus, sP.nov. parvulus Male dj-stingn:ished by heavily clubbed antenna and separated from by slightly higher 4.R., presel)ce of 3 lateral setae on abdominal tergites III - V and rov¡ of stout setae anterior of anal point' r00

PupaseparatedfrombicolorbysmallerthornyareaonsegmentsVand

VI and by intersegmental setae on segment V' MALE 2).Head..Frontalsensillaabsent.EyepubescenceasinFig.65.M.R. distal surface' 0.33. Segment 3 cf palp with one tapering sensilla on expanded : I : 1'57 z 2'3' palp more than half length of antenna, palpal formula O'5 : 0'6 (25p, 25y, 10¡:); Antenna. A.R. 0.6g. Flagelromeres 2 and 3 with 3 type c setae D setae (3oP) at strongly rast flagerl-omere without c setae and with about 20 type 5 lateral setae' clubbed distal end of antenna' Thorax' Antepronotum with hump with 2 Dorsocentrals begin after humeral pits bases conspicuousi scutal I'25 mm' Asp' tiny setaei one pre-alar; 2 scutellars. VÍing. (rig.66) Length R.3.SO.Squamawith3setae,anal]-obeobtuse.Macrotrichiaonthebaseof Costa produced R, R2*3 merged with Rn*U, R4+5 ending just proximal of t3*4' onequarterofdistancetowingtip.Cu'evenlycurved,Anendingdistalof r-m.Legs.L.R.0.6Reartibiaslighttyflared.Bothspurswithbasal one transverse row accessories, inner spur sinuous' Abdomen' Tergites with transverse of setae (Fig.67). Terminalia. (Fig.68) Tergite 9 with a short posteriorrowofstoutsetaewithlargebases.Analpointpresseddown with knobbed against intersegmental membrane. sternapodeme strongly arched' absent' Inrler corners, phaltapodeme with large triangular plates. Penapodeme boat-shaped lobe pointed, with uniform scattering of stout setae; gonostyle butdistaltoothcarriedonalargeextendedbase. tubercles (Fig.69) PUPA (1). Cepha lothorax. Frontal plate with 2 pairs of ' on large Thoracic horn tapering gradually to a point' 2 pre-corneal setae (rig'70) Anterior double-headed wart-like protuberance, 1 on smaller tubercle ' setae antepronotal setae (3OO¡r¡ on large tubercles, 2 posterior antepronot'al II (60;:), 4 scutal setae (60F, 110F, 60¡r, 60p) ' Abdomen' (rig'71) Segment hooks' with long posterior dorsal tubercle with 3 rows of orally-directed' SegmentllwithalargepedesspuriiB,segmentsll-Veachwith3Lsetae of segrnetrts carried on large tubercles. All segments wjth seta" D1 - D4" D4 rows of large II and III on smali tubercles. Segments III - vI with several medial group of anally-directed spines along posterior bord.er, V and VI with 101 smaller spines. Interseg;rnental membrane IfI - IV and IV - V wi-th orally- directed spinules. Segment VIf with 4 and segment VIII with 5 enlarged and flatenned L setae. AnaI lobes each with 18 ftattened setae and 3 long setae. .Iypes. Holotyped , Hindmarsh Falls , 20.v.1970.

Paratvpes. 1ó and pupal caste (reared from Nostoc) , Id (reared from submerged tree roots), Hindmarsh Falls, 3I.iv.L97I-

Addi tional specimens seen: South Australia, 35m. S.E. of Morganr ló , 5'v'1968

(I.F.B. Common and M.S. Upton). . A sing1e male from Berry Springs, Northern Territory 15.vi.L964 (K.R. Norris) belongs to a separate but closely related species.

C.no" NASUTÏCLADIUS Freeman Nasuticladius Freeman, l-96L, p.650 f!'pe species Nasuticladius tonnoiri Freeman, l-96I. MALE. Head. Verticals moderatel-y long, usually reaching to midline; postorbitals longer, heavier than verticals; frontal sensiLla usually present' Eyes bare, slightly elongated dorsally, cibarial purnp with curved dorsal border' clypeus and mouthparts elongated in many species, M.R. usually greater tÏran 0.30, tentorium as in Fig. 88. Maxilliary palps with varying numbers of slightly clubbed sensilla (fO - 151) on ditated inner distal surface of

segment 3, not in pit. Antelnag. FJ.agel1omeres 2 and 3 broader than fonçf, Iast flagelfomere clubbed distally; 3 basally narrowed type C setae present on flagellomeres 2 and 3 and, in some species, on subsequent flagellomeres. Last flagellomere dist-ally with type C setae, nulnerous type D setae as weII as 3 or 4 ring-shaped setae with long projecting central peg. Thorax. Antepronotum normally developed, lobes just contiguous, lateral setae small, slender; scutal process large; acrostíchals minute, decurnbent, irregularly biserial, beginning weII forward and ending half way along the scutumt dorsocentrals usually uniserial; single supra-alar setai scutellars uniserial, usualty 6' Legs' ReaI tibia slightly expanded distally, outer spur less than half length of itrner' inner always with basal taperiug accessory setae ¡ AiJn tarst¡s longer than 5th'

wils_r" squama with a variable number of average sized setae; anal lol:e IO2 present. Basal vein antf R, with macrotrichia; microtrichia minute, darkened to form pattern in some species, punctures small, R2*3 parallel to and cfc¡se to R4*5 at level midway between M3*4 ttd RI, joining costa indistinctly; "tding *I*2. Costa always produced; pfk distal of r-mi Cu1 bent down l¡ut tip turned parallel to wing margin; An ending distaL of r-m. Terminalia' Anal ridge sometimes extended. into a pointrSth sternite joined to coxapodeme; phallapodeme straight with lightly sclerotised plate; penapodeme usually presenti gonostyle boat-shaped but often drawn out into a ridge along dorsal border. FEMALE. Similar to male but :- Head. Frontal sensilla further apart. Antenna with 6 or occasionally 5 flagetlomeres. where 6 flagell.omeres present, 2-5 of equal length, shorter than I and 6. lst - 5th each wíth one pair and 6th with at least 2 pairs of type c setae ¡ 2nð. - 5th flask-sha.ped, bulbs up to half length of segrment; lst and 2nd each with a single sunken seta. where onry 5 flagellomeres present, lst is at least twice length of 2nd, bears 2 pairs of type c setae and 2 sunken setae, and shows other signs of being the result of recent fusion' Terminalj-a. SterniteVlllwith 10 - 20 setae on each side' Egg guides with Z- enlarged' shaped medial border, in contact distatly. Tergite IX only slightly usually with distinct medial clefti gonocoxite IX not produced distally; sternite X with a narrow ridge. Vaginal apodeme one fifth segment width' projecting weII into segrment VII; spermathecal ducts lightly sclerotised with single looP, exPanded basallY' of the 15 species described below, the females of only 4 of them tonnoir:L Freem., dÍplos!¿]¡¡e sp.nov-, cari¡a'Egg sp'nov' and salteri sp'nov' have been positively identified. There are several groups of unidentified females'

each with different palpal and antennal- fonnulae and different numÌ¡ers of to squamal and pre-alar setae. variations in leg formulae overlap too often offer any guide to affiliatíons with particufar males. One of the unidentifiecl groups has an additional pair of large type c setae on each antennal segment'

(Fic;.84) . Terminali a are ofthe general pattern illustratecl for N. salteri

Freeman (I96f) namecl NasuLical-dius to include 5 species whicìr he considered formed a series. Below f have re

KEY TO SPEC]ES OF THE GENUS NASUTICLADIUS I Apex of antenna rounded or finishing in a point' not extended 4 Apical Point of antenna extended 2 2(l) Costa produced to wing tip, palpal segments 3 and 4 nov with sensilla mucronatus sp. Costa not reaching wing tip, palpal segment 4 without sensilla 3 punctus sP. lìov. 3(2) A.R. greater than 1, P'R' greater than 0'3 , parapunctusr sP.nov A.R. Iess than 1, P'R' less than 0'3 4(1) Long proboscis present (M'R' greater than 0'8) 5 Proboscisshortorabsent(M.R.lessthan0.8) 6 niger Freemall 5(4) Abdomen dark, anal point short' pubescent

Abdomen banded, anal point absent' anal ridge tonnoiri. Freeman hairy 7 6 @) lilings with dark markings Ì1 Wings without dark markings wilsoni Freematr 7(6) AnaI Point verY long, M'R' IoY I AnaI Point short, M'R' high diplostvlus, sp. no\'. 8(7) Gonostyle with rod-Iike basal lobe 9 GonostYle rvithout basal lobe ater Freeman 9 (8) V,fings extensivelY suffused 10 Wings on1Y s1ight1Y suffused 10(9) lnner lobe divided into anterior' middle and posterior sections *itthf_ Freeman fru.t¿r sp"nov- Inner fobe divided into anterior & posterior sections 12 11(6) Short proboscis present (MR' greater than 0'4) 104

t4 Mouth-parts not or only slightly extend'ed 13 t2 (11) Segrment 4 of palp shorter than 3 or 5 3' Segment 4 ofpalp shorter than 5' Ionger than carinatus sp. nov. sensilla on segment 3 onlyt P'R' Iess than 0'25 inner lobe f3(12) Segments 3 & 4 of patp with sensilla' salteri, sP. nov- large, P.R. greater than O'25 lobe small' Segment 4 of¡alp without sensilla' inner eurynotus, sP.nov' P.R. less than 0.25 very large lonqipe NNIS, sp. nov. t4 (II) Costa produced to wing Èip' penapodeme Costanotreachingwingtip,penapodemenormal and distal sections 15 (I4) Inner lobe divided into proximal Inner lobe not divided to 16(15) AnaI ridge narrovt, inner lobe restricted T7 anterior two-thirds of gonocoxite AnaI ridge broad, inner lobe extends nearl)'to duckhousi sp" nov. insertion of gonostYle process' 17(16) Inner lobe with smaIl, bare knob-like brevip ennls, sp. nov. penaPodeme very small Inner lobe with bare cornert penapodeme of gtover!, sP"nov. moderate size

Nasuticladi us tonnoiri Freeman

Nasuticladius tonnoiri Fremman, L96L, P'650' SeparaÈedfromglgbytackofanalpointandbareSquamaandfromall other species by its long proboscis' strong; frontal sensilla absent; eyes with M.ALE (1). Head. Verticals long, dorsalelongation2facelswide,cibarialpumpdeeplycurvedondorsalborder, extended to form proboscis' M'R' arms curved j-nwarcls; clypeus and' mouthparts 0.84¡palpalformulanotdetermirrablebutsegiment3withlO-15sensilla. A'R' l.O; flagel.lomexes 2-12 scattered randomty over distal surface' Anten'na" spatuJ'ate type c setae @ig'72)i each with 2 and last with s (37p) tapered 10s

3 ring setae' last flagellomere slightly clubbed and pointed disÈally with begin just Thorax. Antepronotal lobes with 4 lateral setae, dorsocentrals Legs' L'R' o'75' forward of humeral pits; 6 or more pre-alars; 8 scutellars' setae. !.Iing. rear tibia with uneven d.ense comb, both spurs with accessory Lengthl.Bmm.AspectRatio4.T;squamabare,anallobedistinct.Macrotrichia closer to *1*2' Terminalia" ot R4+5, costa produced to wing tip' R4+5 finishing with broad lateral arms' AnaI point ridge-Iike, ending abruptly; sternapodeme plates, P.R. o.23. corners drawn out. Phallapodeme with curved' triangular for eurynotus, spine rnner lobe broader distatly, right-angled; gonostyle as small. stripes separate' FEMALE. Freeman says, "Very similar to male but thoracic segiments 3 - 6 with shoulders and background yellow; antennae with 7 segments' long narrow necl

seen the ó paratype.

Nasuticladi.us ni Freeman

Nasuticladius niqer Freeman, I96L, P'651' Bothsexeswithlongproboscis'squamawithnumeroussetae,malewith unsclerotised pubescent anal point' MALE(I).Head-Cibarialpumpwithdeeptycurveddorsalborder'armscurved long proboscis' M'R' inwards. clypeus and mouthparts (Fig.73) drawn out into srightly pointed sensilfa; about r. segment 3 of palp with numerous, scattered, flagellomeres palpal formula 0.4 ;0.7: 1 : 0.9 ¡ L.25. Antenna. A.R. 1.5; spatulate (32y) type c 2 - 4 broader than longi 2 - 8 each with 3 tapered distally and with 5 setae. Last flagellomere slightly ctu'bbed and pointed typeCsetae(32p)numeroustypeDsetae(38P)andsinglesmallringseta. Dorsocentrals biserial' Thorax. Antepronotal lobes with 4 - 5 lateral setae' L'R' beginning anterior of humeral pits; 3 pre-alars, B scuteliars' ætr-' o.76|reartibiawithunevendensecomb,bothspurswithaccessoryseta.e. vLi'g.Lengthl.95mm.,Asp.R.4.8.Squamawith12setae;anallobeobtuse' procluced to distinct. Macrotrichia on R4+5, R2."3 ending closer to R4+5, costa Fig.72 Nasutícladius tonnoiri, Freeman, spatulate type C seta.

Fig.73 Nasuticladius niger Freeman, male, mouth parts.

Fis. 74 Nasuticladius diplostylus sp. nov. male, wing.

Fig.75 1l il tt terminal-ia.

Fig. 76 I female, terminalia. It a\

t t I t (

I 75

73 72 :-'- |

f r1 ¡ I :a J a a. ì¡ t ..aÙ '¡ ,

?6 t06 wing tip, R4*5 .tding closer ao tt*2 than to Mr*, . Terminalia. Anal Point short, unsclerotised, pubescent- Sternapodeme slender, evenly curved' cornels reduced to thickenings; phallapodeme with contoured triangular plates; P'R' O.26. Inner lobe large, rounded; gonostyle with- small subterminaf bare peak' outer surface sworren sinuously r/3 of distance from base, spine normar. TypES. I have made a slide of the holotype ó and returned it to SPHTM'

Nasuticladius dipfostylus' sP. nov-

Mouthparts slightly extended; wing pattern not dark, area sma1l, penapo- basal deme of medium size, claspette with 2 lobes, gonostyle with' rodlike process, reminis cent of Diplocladius- to midlj-:re; MALE (3). Head. verticals small, decumbent, frontal sensilla close pal-p eyes with dorsaf elongation l facet wide, M.R. O.47'¡ segments 3 and 4 of : I : 0'87 ì each with 2 stout rod-Iike setae in pit; palpal formula 0.3 t o-7 small dent on outer surface, Ia'st 1.9. @.. A.R. I.O, pedicel with single flagellomeredistatJ-ytaperedwith4-5typecsetae(25P)aswellasa 4 lateral nrunber of type D setae lZS¡r) . Tlrorax. Antepronotal lobes with 6 scutellars' seÈae, dorsocentrals begin level with- h-umeral pits, 3 pre-alars, Legs. L.R. 0.6, both spurs of rear tibia with- accessory setae, comb wittr few setae. w!rg. (rig.7a) Length I.25 - l-.35rnm. Asp- R. 4.i - squama with 3 closer to setae, anal lobe very small' Macrotrichria ot R4*5t RZ*3 meets- costa distance to wing R4+5, running with it for some distancei costa produced 3/4 of along tip. R4*5 ..dÍng closer to t3*4 ttran to *I*2' Micrctrictria darkened Crig.- 75) ML+2, uncler Cu1, above pfk and half-way along anal lobe' Termi¡alia. Anat point small-, bare, rounded wittr Iongitudinal ro"r of setae' coxapodeme joined with 9th sternite, sternapodeme evenly rounded-, corners reduced, P'R' phallapodeme with contoured plates, sclerotised at anterior extremities' proximal' one o.27. Inneï lobe with two broad setate digit-like lobes' one distat.GonostylewithinnerL,orderdrawnoutproximallyintoalong border unsclerotised rod-Iike process as long as the style itself, outer artcl surr- produced at distai end to form ror'rnded setose corneÏ' Spine strong

ounded by large ridqe. 107

FEMALE (1). As for male but :-

M.R. O.4B¡ second segment of palps with 3 - 4, third with single slender sensilla; palpal formula 0.4 : O.7 : I z L.2 z 2.I. Antenna with 6 flagellomeres, rst fusif ormt 2- 5 frask-shaped with neck" tnota"ã-r-oasar bulbs, 6th tapered to poínÈ. One pair of type C setae on flagellomeres 2 - 5' 6th with 2 pairs, reaching to IeveI of segrnent tip. Flagellomere formula I-22 I: 1 : I : 1 : L.45. Thorax. Dorsocentrals begin close to antepronotum. Wing.Length

1.2Omm, Aspect Ratio 3.9. Costa produced righÈ to wing tip, microt'richia darkened over slightlY larger areas. Terminalia. (Fig. 76) Sternite VIII with 15 setae on each side. Tergit IX not enlarged, median cleft distinct. Sternite X with broad ridge, cerci rectangular from dorsal view. Vaginal apodeme basally one third width of segment VIII' projecting one quarter into segrment vII. spermathecal ducts with basaÌ thickening. TypES. Holotype d , Parawa, South Australia, 16.vi.197L. Paratypes: South Australia; Eric Bonython Reserve, 2d , 22.iv.1970. 19 , f.iv.I91O.

Nasuticladius fuscus sp. nov. Mouth-parts slightly extended, wing pattern not dark, area smalfi separated fro* @.!y.lus by larger penapodeme and simple style- MALE (2) Head. Frontal sensilla carried on slight trumps, eyes with dorsal elongation 2\ facets wide. Clypeus a¡¡d mouthparts slightly extended, M'R' O.42 - 0-46. Segirnents 3 and 4 of palp with 3 and 2 lanceolate sensifla respectively; palpal formula O.25: 0.5 : 1 : 0.78 : I.43. Antenna' A'R' l-'00, flagellomeres 2 - 7 each r'¿ith 3 type C setae (32F), last flagellomere slightly (32y) type D setae t32p) and clubbed distally with 4 - 5 type C setae ' numerous 2-3 ríng setae. Thorax. Antepronotal lobes with 4 lateral setae, dorso-. centrals u¡riserial beginning level with humeral pits. Six pre-alars, 6 scutellars. Legs. L.R. O.'IO, comb of rear tibia sparsely setose' I{it9-.. Length 1.45mm, Asp. R. 4.8. Squama with 3 setae, anal lobe small but distinct' macrotrichia on R4+5, R2*3 *."ts costa half-way between Rt and R4*5' costa produced nearly to wing tip, *4*5 .tding closer to level of *I*2 than to *3*4'

t"ticrotrichia stightly darkened in same regions as i-n N' 9:gt"=t:lt=' Fig.77 Nasuticladius fuscus sp. nov. ' male, terminalia.

Fig 78 Nasuticladius eurynotus sp. nov., male, terminalia.

Fig. 79 NasuÈicladius nov., ma1e, head, - lines used in determining width of dorsal extension of eye' - campaniform sensillum. - frontal sensillum - hyaline sensilla - length of mouth Parts, - post orbital setae.

Fíg. Bo Nasuticladius punctus, sp.nov. ma1e, apex of antenna, D- tYPeDseta.

Fig. 81 I ma1e, anal point of terminalia. I I

L I I v ou J

u ú - u

77 78 rrl þe I I

l--J ---o

1 n I 79 80 108 Abdomen. Sternites 2-5 with paired sclerotised. areas as in eurynotus, sp.nov.

Terminalia. (Fig.77) Anal point round.ed, sclerotised a¡¡d. setose. Sternapodeme

strongly arched, corner processes smaIl, phallapodeme with plate drawn into 3

sclerotised points, P.R. O.42. Inner lobe large, right-angled., ventrally contoured; gonostyle with dilated dorsal surface.

TYPES. Holotyped and Id paratype (at light) Parawa, South Australia 23.Lx.L97O and 15.vi.1970 respectively. Nasuticl-adius eurlmotus, sp.nov.

A ì-ight brown species with costa produced to wing tip, mout-hparts only moderately extended. Inner lobe small, broad; recognised by distinctively dilated. dorsal surface of gonostyle.

MALE (9). Head. Dorsar border of cibarial pump deeply curved., M.R. o.44. Eyes witl: dorsal elongation 2 facets wide; segment 3 of palp with 2 groups of 2-3 slightly curved rod-Iike sensilla (15¡r), palpat formula 0.25 : 0.5 : I :0.9 : 1.4. Antenna. A.R. I.3 - I.7, flagellomeres 2 - 8 each with 3 type C setae (30¡r), la.st flagellomere tapered, not pointed, with type D setae (1 p). Tlrorax. Antepronotal lobes with 4 setae on enlarged lateral corners. Dorsocentrals irregularly bj.- serial , begin fonr.rard of humeral pits; 5 or 6 pre-alars arranged in 2 consecutive crescents; I scutellars. Leg. L.R. O.7 - 0.8, rear tibia not expanded distally, both spurs with accessory setae, comb setae long, dense. Wing. Length L.6 - 2.Ot mm. Asp. R. 4.5. Squama with 4 - I strong setae; a¡al- Iobe large, distinct.

lvlacrotrichia on RI and R4+5, Rr*, ending closer to *4*5, costa ¡íroduced to lving tip. !Ld"*"".. Sternite I without markings, II - V each with 2 pairs of lightly sclerotised spots, the anterior pair being closer together, V sometimes with narrow median strip of sclerotisation with 6 - I short strong setae, VI and VIr with increasingly broader median strips of sclerotisation with numerous short

strong setae. Terminalia. (Fi g.7e\ Anal ridge broad, tip covered with medium

length setae. Sternapodeme with broad Lateral arrns, corners rounded anC reinfcrced, phallapodeme with triangular p1ate, P.R. 0.15. Inner lobe smal1, broadr gonosc-vie dilated on dorsal- surface and with triangular cross-section. Spine small, subterminal.

Specimens from New South Wales have a M.R. cl-<¡se to 5.0 and vary in the number of flagellomeres carrying sensory setae. r09

TypES: Holotype ó and I d paratype from Botanic Gardens, South AustraLia 19.v.I97O. Further paratypes. South Australia: Botanic Gardens, ló , I5.v.I9ZO (at light) , 26 , 3t.iii.Ig7A, L 6 , (reared from Blechnum debris) f .iv. L97O¡ Eric Bonython Reserve, 1ó, 26.v., Ló , 3.vi.I97O; Healey's Ió , 29.iv.I97o. AddiÈional specimens seen: South Australia: Alnut, Id , 30.vii.1969; Botanic Gardens, ód (swarms) f .iv. - 3.vi.1970, 2I.ív. - 2.ví.L97L; Eric

Bonython Reserve, ód (swarms) l-4.iv. - L2.viii. L97o, 28-iv. - f .ix.I97L¡ 2d Ið Healey's 3ó , 29.iv.I97o; Hindmarsh FaIIs, td , f iv', ' 2o'v" ' 3.vi. L97O¡ 9d , 26.v., 2ó , 9.vi .L97I¡ Parawa, dd (some at light) 27.v'- 15.viii.LglO, 14.iv. - 12.viii.Lg7I; Tanunda Creek, Ió , 9.viii.1969' Alligator Gorge, ld , 4-v.1968 (Common & upton) - Australian capital Territory: Black Mountain, 3ó , 2.vj-., 2L.vii., 6.x.1965. (I.F-B.CÔrrunon) : Corree Creek, Id , f .iv. I96L¡ Mt. Gingera, ld , 14-iv.1968 (O.tt.Cottess); Gibralter creek, ld , 18.x.L967 (2. Liepa). New south vtales: sawpit Creek, Mt. Kosciusko, Id , Io-ix-1960 (z'r'iepa) ' Nasuticladius punctusr sP. nov. separated from other Nasuti- Both- punctus and parapunctus Sp.nov. , aTe clad.ius by the extended point of the 1asÈ flagellomere and the short' bare' rounded analpoint. N. punctus has a higher A.R- and P.R. than parapunctus' MAI,E(7).Head.(r'ig.79).Verticalssmall,singleseÈaclosetomidlineon each side, remainde:: finish half way to midline. Brow slighÈly rounded' Eyes with dorsal elongation 1 facet wide. M.R. o.32- PaIpaI segment 3 with 2 tod- like sensilla (fSp¡, palpal formula o'25 : o'5 : I : I : I'3' Antenna' A'R' (2Oy),Iast 1.O - 1.3, flagellomeres 2 - 9 each with 3 type C setae flagell-o- mere hrith extended narrow point (Fig.8O), type D setae (3O¡) ' Thorax' Ante- pronotal Lobes with g lateral setae. Dorsocentrals begin at level of humeral pits, 2 - 3 pre-alars. Legs. L.R. 0.6, comb of rear tibia with derrse setae' vriqg. Length 1.55 - I.8Omm., Asp. R. 4.27¡ squalna with 3 - 5 setae; anal lobe present. costa produced half-way to wing tip. Abdomen-' sternites simi- seg- lar Eo eurynotus except that spots of scl-erotisation are smaller and (niq. menLs I-5 bear narrow lateral bands of light sclerotisatio n. Terminalia gl_). Anal ridge broad. with small,barerround anal point; sternapodeme broad, 110 corners rounded and reinforced. Phatlapodeme with contoured Èriangular plate' P.R. O.40¡ inner lobe broad with bare corner, çJonostyle with dorsal ridge bulbous distally, spine subterminal. TypES. Holotype d an¿ f d paratype from Iight trap, Botanic Gardens South Australia, I5.v.I91O- Further paratypes: Parawa, Ió , 30.vii.1969; Aldgate Creek, ld , 3'vi'I969i Eric Bonython Reserve, 1ó , 26.v.197I'

Additional sPecimens seen. south Australia: Eric Bonython Reserve, 5d , 2O.v.I97O, 3ó , 26.v.L97L¡ Hindmarsh Fa11s, 1ó , 2O.viií -L969, Ió , 22'iv', 5 ô, 3.vi. Lglo, 1d , :I2.v.l:97:-¡ Inman Valley, ld , 3.ix.L969¡ Parawa' 5d r 20.v. r 13ó (at light), 27.v.Ig7Ot dd (at fight) 5.v. - 30.vi.I97L'

NasuticLadius ar ctus sp. nov.

Refer to comments on N- punctus. for MALE (4\. Like puqctus but verticals form a complete ro\¡/ of normal size genust M.R. 0.35. Artte"na-. A.R. 0.60, flagellomeres 2 - 10 eactr with 3 type 4'I' C setae. Legs. L-R. 0.55. Wing. Length I-25 - l-.35mm., AsP' R' Terminalia. P.R. O-20. is only Among specimens allotted to Èhis species is one in which A'R'

0.3 and segments 4 and 5 of palp are joined together in a large bulb' TypES. Holotype ó and 3 ó paratypes, Botanic Park' Mt. Lofty, 19.v.I97I. Further paratype: Hindmarsh Falls, 1d , 26'v'1970' L9'v" 2ó Àdditional spe cimens seen. South Australia: Botanic Park, 6ó ' 26.v.L97I.

Nasuticladius carinatus, sP.nov'

costa produced to wing típ, inner lobe extended into distinctive bare in sharp thumb like projection, gonostyle with dorsal ridge ending distally corner. 2 facets MALE (8). Head. Frontal sensilla absent, eyes with dorsal elongation sensilla wide, M.R.0.40. PaIpaI seg:rnent 3 with 3 groups of 2 or more rod-lìke A.R. 1.0 - l'5t (zOP), patpal formula O'3 : O'5 : 1 : 1'I z l-.'4 Antenna. Fig. A2 Nasuticl-adius carinatus sp. nov., ma1e, terminalia.

!'ig. 83 Nasuticladj-us salteri sp. nov., male, terminaLia

Fig. A4 ll female, terminalia.

Fig.85 Nasuticlad.ius brevipennis sp. nov., male, wing

rig. 86 il Í I terminalia. .::.... ä 82 É

,t It r, a o / a, I o o l, !,¡ 83

,l\ì

a 7 a I a a ¡

l--¡ 86 84 lII flagellomeres I - 5 each with 3 type c setae, lOJ¡, 33)r, 48y, 30P' 26¡-1 1on9 respectively. Last flagellomere with 3 type c setae (17F), type D setae (35p)' Thorax. tuìtepronotal lobes with 2 - 5 lateral setae- Dorsocentrals begin well forward of humeral pits; 6 pre-alars, Legs. L.R. 0.5 - 0.9. Rear tibia with t'65 accessory setae on both spurs, comb straight, setae sparse. vling. Length - 2.oo. ASp. R. 5.00. Squama with 4 short strong setaer anal lobe small, distinct. Macrotrichia ot R4+5, R2+3 joining costa closer to *4*5r costa produced to wing tip- Abdomen' Sterni tes similar to eurynotus but 5th always sclerotised along midventral line' Terminalia. (!'ig.82) Anal ridge sparsely setose, pointed. sternapodeme narro\^l, strong, without corners, phallopodenie with rectangular plate, P.R. O.L2. Tnner lobe short, thumb-li-ke' bare distally" Gonostyle boat-shaped, with dorsal ridge ending distally in sharp corner and row of medium setae along inner surface' formuLa 0'3;0'5: FEMATE (1) . Like male but palps with 3 groups of sensilla, palpal I : 1.I : 1.75. Antenljr, vrith 6 flagelloneres, 1st - 5th fiask-shaped with lot1g bulbs, 6th tapering gradually to a point, flagellomere forrnula 1 : 0'8 : 0'8 : 0.8 z I.25. Terminali-a. Sternite Vrlr with f5 setae on each s-ide' gonocoxite VIII moderate, genital comb with short setae. Gonocoxite IX enlarged' cerci bulbous. TypES. Holotype d from Hindmarsh Fallsr South Austrafia, July I97O¡ Paratypes l¿l 5.vi.\91a" south Austrafia: Botanic park, ld , 15.v., ld , 21-.v., ld , 28-v-, April - August Ad-dítional sP ecimens seen. South Australia: Botanic Park ðd 99 Lg|O, I97I (some at light or sticky paper); Eric Bonython ReserYe dd 99 3.vj-. - l.vii .L97O, 3 ó 9.vi. L97L¡ Happy VaIIey, 2 ð t t.vii,L969; Hindmarsh (a't Falls, dd 99 , May-June L97O, L9lLi Parawa, 2 ð , 6.v.I97O, 16 light), 16.vi.L97I.

Nasuticladius salterì., sp. nov. wings segrments 3 and 4 of paLp with sensilla, apex of antenna pointed, without pattern, costa produced to wing tip, itlner lot¡e of male distinctive'

penapcdeme of average size for genus' MALE(13).Head.Vert.icafslong,frorrtalsensil]-aabsent.Eyeswithdorsal II2 elongation 2 facets wide. M.R. 0.45. Segments 3 and 4 of palp each with gToup of 6 clubbed sensilla (2O¡t), palpal formula 0.3 :0.5 :1 : 0.75 :1. Antenna. À.R. L.2 - I.55, flagellomere 1 with singl-e small type C seta, 2 - 12 each with 2 - 4 type C setae, Iongest ( 5p) on flagellomere 4, last flagellomere ending with sharp point, 3 ring setae. Thorax. Antepronotal lobes with 7 small laÈeral setae, dorsocentrals begin at level of humeral pits, 5 - 7 pre-a1ars. Legs. L.R. 0.65 - O.75, rear tibia with accessory setae on both spurs, comb in two arcs, dense. I{ing. Length I.75 - 2.00mm., AsP. R. 4.8. Squama with 6 long setae, anal lobe distinct Macrotrichia on both R, .td R4*5. *2*3 ending closer to Rrr costa produced to wing tip. Abdomen. Sternites similar to

us except that VII is completely sclerotised. Terminalia. (Fig.83) ' AnaI ridge sharp, bare at distal tip. Sternapodeme with scarcely distinguish- able corners, phallapodeme with short base, plates large and heavily contoured' p.R. O.2 - 0.35. Inner lobe proximal, rounded, gonostyl-e dilated on dorsal surf ace, tooth su.bterminal .

FEMALE 2 . Like male but palpal formula 0.3 : 0.5 : I = O.77 ¿ I.24. Antenna. with 6 flagellomeres, lst - 5th flask-shaped with long bulbs, 6th taperi-ng gradually to point, flagellomere formula 1.o ; 0.85 : 0.85 : 0.85 : 1.54. Terminalia. (¡íg.84) Sternite VI]I with 20 setae on each side' Tergite IX short, gonocoxite IX slightly produced, with 2 - 3 long setae distally; celci hemispherical from dorsal víew. Vaginal apodeme basally one third width of segment VII, projecting about one sixth of length into segment vTf. TypES. Holotype d and Id and 29 paratypes, Brownhill Creekr South Australia, 22.!v.Lg72. Further paratypes South Australia: Aldgate Creek, 2d , 3'vi'1969¡ Botanic Gardens, Ió ,2.vi.r 1ó (at light) 2.vii.Lg7O,ld ,19.v-1971; Deep Creek (Adelaide Hilis), tó , 29.vi.I96g; Eric Bonython Reserve , Ló' 26'v'I97Li

Hindmarsh Falls, Ió , 20-v-l-91 o¡ Para\'Ia' 1d , 28'v'1970' Additional specimens seen. south Australia: Botanic Gardens, 2ó ,27.v.,3ó t 3.vi., Ió , I.vii .Ig7O, Ló , 19.v. L97I¡ Brownhill Creek,dd , 22.iv.1912; Erj-c Bonython Reserve, 2d , 2O.v.1970; Hindrnarsh Falls, 2d ,2O-v.L97O¡ Parawa, 2ó , 2o.v., 2ó , 3.vi.rg7o, 2ð , 28.iv., 1ó , r2-v-, Ló , 26'v'r91T' 113

Nasuticladius brevipennis, sp. nov.

Costa produced only half of distance to wing tip, penapodeme small for genus, distinguished by small bare knob on inner lobe.

MÀLE (5). Head. Verticals short, reach 3/4 of. distance to midline. Eyes with dorsal elongation 2 facets wide. Frontal sensilla absent, M.R. O.24, segment

3 of palp with only 2 rod-like sensilla, palpal formula O.4 : O.5 : I : 1.1 : I.7. Antenna. A.R. 1.O, type C setae 2O¡r, type D 25¡t long, 3 ring setae. Thorax. Arrtepronotal lobes with 2 - 4 Lateral setae. Dorsocentrals beginning forward of humeral pits, 4 pre-alars, 4 scutellars. Legs. L.R. 0.60, 4th tarsus equal to 5th. Vling (rig. 85) Length 1.35 - 1.45 mm., Asp. R. 4.43.

Squama with single seta, anal lobe obtuse, veins bare, R2*3 *""ting costa closer to *4*5. Costa produced. half way to wing tip. Terminalia (Fis. 86) .

Anal ridge reduced, pu-bescent, with small anal point; sternapodeme of mediun:r thickness, strongly arched, corners pointed and reinforced. Phallapodeme with curved plates, P.R. O.O2¡ inner lobe thorn-like, directed. posteriorly, distally with small bare knob. Gonostyle with crescent-shaped cross-section, spine surrounded by snall ridge. A single specimen from Eric Bonython Reserve has type C setae on 4th flagellomere and inner lobe with larger, longer bare point. TYPES. Holotype ó and I ó paratype, Botanic Gardens, South Australia,

21.iv.1970. Further paratypes. South Australia: Eric Bonython Reserve l-ó , 26.v.I97L, Botanic Gardens fd , 2L.iv.I97o, Id , 26.vii.I97o. Additional specimens seen. South Australia: Botanic Gardens, ód (swarm). 2L.v., # (sticky paper) 25.viii.I97a¡ Torrens Lake, Id , 24.ii.1971.

Nasuticladíus healensisr sp. nov.

Separ ated from other Nasuticladius since inner lobe of gonocoxite divided into proximal and distal sections.

MALE (3). Head. Verticals reach half way to midline, eyes not elonga.ted dorsally, IvI-R. about O.3, segment 3 of palp with 2 groups of cl-ubbed sensil, l-a

2Oy), palpal formula variable but each segment longer than preceed.ing one" Fig. 87 Nasuticladius healensis sp. nov. male, terminalia

Fig. 88 Nasuticladius mucronatus sp. nov. male, tentorium.

Fig- 89 lt terminalia.

Fig. 90 Nasuticladius gloveri sp. nov. male r gonocoxite and style.

Fig 91 Nasuticladius longipennis sp. nov. male, terminalia

Fig. 92 Nasuticladius duckhousi sp. nOv. male, gonocoxite and stYle. r

a J v J 87 o a t a

89 88

./ 90 I I t a, C t J I l,

3 ö

aa a ô ô GÙ ór r¡ 92 r14

Ar¡tenna. A.R. 1.35 - 1.65, rast fragerlomere with 2 type c setae lzsy)' pointed apically, type D setae 2o.¡: 1orrg. Thorax. Antepronotal lobes n<¡t begin at contiguous, 3 - 5 lateral setae, scutal process small-' Dorsocentrals 0.7, }roth level of humeral pits, 4 prealars, 4 scutellars. LJSE.. L.R" 0.6 - rear tibiar spurs with accessory setae. comb straight, setae sparse' wing-' LengthL.4_I.95mm.,AsP.R.about4.4,Squanawith8setae,anallobe to present. Macrotrichia on base of Rrr costa produced one third of distance but V alwaYs wing tiP' Cul distinctlY bent. Abdomen. Sternites like eurYnotus similar to TI. Ternuinal-ia. (Fig.87) AnaI ridge tapered to bare tip' sterna- Phallapodeme with podeme of varialcle breadth, corners rounded and reinforced' into two contoured triangular plate P.R. 0.2 - 0.3. fnner lobe divided ventral and portionsr oÍrê proximal, dorsal with corner, the other distat, spine subterminal' rounded. Gonostyle boat shaped, may have slight dorsal ridge; Paratypes TypES. Holotype ó from Hindmarsh Falls, south Australia B'ix'1971' 1ó 29'iv'1970' South Australia: Orlando, Ió , 9.viii,1969; Healey's , considerable As índicated in the description the specimens exallined show specíes' variation i.n a number of characters; they may represent 3 differenct treated as a untir further specimens can be examined, however, they are better single species. Nasuticfadius mucronatus sP. nov. wing tip' inner lobe Antenna with apicat point extended, costa produced to of distinctive shape, penapodeme small for genus' MALE0).Head-Eyeswithdorsalelongation3facetswide'M'R'O'34'ten- pairs of rod-like sensilla torium see Fig.88. Segments 3 and 4 of palp with 2 (15¡),pa1palformulaO'252O'4:l:O'9:I'I'Antenna-'A'R'I'3-I'6' (2qù Iast flagellomere with two type flagellomeres 2 - 5 with 3 type c setae ' 2O¡r long 4 rlng setae' C setae (2Op) and apical point drawn out, type D setae ' begin forward Thorax. Antepronotal lobes v¡ith 4 lateral setae, dorsocentrals L.R. 0.6.7, xear of humerat pits, about 7 pre-alars, 8 scutellars. I,e-gs-. Length r'56 - l'80 mnL' tíbia' comb in two arcs, with long dense setae. vling. but distinct'macrotrichia Asp. R. A.Lg, squama with 1 - 6 setae, anal lobe small IJ.J present on Rl .td R4*5, costa produced right to wing tip. Abdomen. Sternites similar to eurynotus. Terminalia. (rig.89) AnaI ridge with short point sparsely covered with microtrichia, sternapodeme narrov/r corners sharp, reinforcedi phallapodeme with large plate, P.R.O.07. Inner lobe right-ang1e'J v¡ith bald corneri gonostyle wiÈh slightly dilated inner surfacej spine subterminal. TypES. Holotyped a¡dó paratype, Hindmarsh Falls¡ South Austrafia &ivl970, Further paratyPes: South Australia: Botanic Gardens, Lô , 15.v.L97O¡ Eric Bonython Reserve , Ló , 22.ív-1970. Addition al scecimens seen. South Australia: Botanic Gardens , Lô , 8.iv.1970 (emergence trap), dd 16.iv. - I5.v.Ig7O (some at light)rIó ,31.iii.L97O (reared from moist Blechnum debris), dó 31.iii. -26.v.L97l-¡ Eric Bonython ó Reserve, 3 d, L2.v., 1 ó, I5.v. I97O¡ Hackam, 3 d, I.viii. L969¡ Parawa, 5 27.v.I97o (at light), Lô, 9.vi-, dd , 12.viii-L97]-.

Nasuticladius qloveri.s p. nov.

Brow cut off squarely, cibarial pump with rectangular dorsal border, inner lobe with bare corner.

MALE G\. Head. Brow squared across midline, cibarial pump with rectangular indentation in dorsal border. Eyes without dorsal elongation. Segment 3 of palps with about 7 clubbed sensilla ín two groups, palpal formula 0'5 z O'75: 1 : l : 1.35. Antenna. A.R. 0.8 - 1.1, flagellomeres 2 - 12 with 2 type c setae (25p), Iast flagellomere with 3 type C setae (251t), type D setae(fSp)ana 4 l:arge ring-shaped setae. Thorax. Ante-pronotal lobes with 2 - 3 lateral setae, dorsocentrafs begin just forward of humeral pits, 4 - 5 pre-alars. Legs. L.R. 0.5, rear tibia with accessory setae on both spurs, comb straight, setae sparse. wing. Length 1.6 - 1.65mm, Aspect Ratio 4-53, squama with 3 or 4 setae, anal lobe obtuse. Macrotrichia on RI, R2*3 ending closer to R4+5' costa produced 2/3 d,istance to wing tip. Abdomen. Sternites 1 - 6 without sclerotisation, Sth with B - 10 evenly distributed setae, 6th and 7th with onll' medían strip free of sclerotisation. Terminaria. (rig.90) Anal ridge shar¡r'

sternapodeme broad, corners large, rounded and reinforced, phallapodeme with IIb parallet-sided plates, P.R. o.28¡ inner lobe r¿ith- proximal bare corner; gonostyle with concave inner surfacei spine termj-nal' TypEs. Holotype d from Hindmarsh Farls, South Australia, 26'v'I97L' paratypes. SouËh Australia: 2 ô, Botanic Park, 28.v.197O, 1 d , Eri'c

Bonython Reserve , 26-v.1970' Additional specimens seen. Soutt¡. Australia: Eric Bonython Reserve' ]d

26.v.L97L, 3 d, 9-vi-L97I- Nasuticl-adius longipennis, sP. nov-

Mouth parts shortened, costa produced to wingtip' penapodeme very 1on9, projecting into segment VlI. eyes MALE (4\. Head. verticals finish at varying distance from midline, with dorsal elongation 1 facet wide, M.R. o.25, segment 3 of paip r¡¡ith group of 3 capitellate sensilla on outer surface, one on inner surface, palpal formula o.3 : 0.65 : I : O-82 z I.2. Antenna. A-R. O-7 - O'B' Iast flagellomere not clubbed distally but tapered to a point r¿ith- 4 tíng setae and 3 tlpe c setae, both. type c and D setae 25u 1on9. Thorax' Antepronotal Iobes wittr 3 - 4 lateral setae, dorsocentrals beqin at level of humeral pits, 2 pre-alars. Legs. L.R. O-58, accessory setae of rear tibial spurs Asp' small, comb straigh-t, setae sparse. viing. Lengttr 1-35 - 1.55 mm' on R. 4.55. Squama witt¡- single fine seta, anal lobe present. Macrotrichia basal vein on1y, R2+3 ends closer to Rtr costa produced to wilg tip' Terminalia (Fig. 91) - Anal ridge narro\4r, anal point a small- bare knob' pha1la- Sternapodeme narrow, corners with. smaLl points slighÈIy reinforced' bare podemes with curved plates, P-R- O.7O. Inner lobe small, digitate, distäIly, gonostyle drawn into seta*covered peak distally' spine average' sub-terminal. TYPES. Holotype d and I d paratype, Eric Bonyttron Reserve, south Austral-ia'

l-ó Ma1' 26.v.Ig1L. Further paratypes. south Australia: Hindmarsh Fall-s' '

Lg7o, Botan-i-c Gar

Nasuticladius duckhousi sp. nov. Recognisedbybroadanalridgeandbroadlybasedinnerlobe. eyes withouf MALE (3). Head. single vertícal seta on each side of midline, dorsal elongation. M.R. 0.35. Segment 3 of palp with 2 rod-like sensilla' palpal formula 0.4 : 0.85 :1 : I'I :1'45' Antenng' A'R' 0'80 - 0'90' Flagellomeres 2 - I each with 3 type C setae (I8¡r) r last flagellomere with lobes with rounded tip and 4 - 5 type c setae (13¡:). Thorax. Antepronotal 3-6 lateral setae. Acrostichals absent, dorsocenÈrals beginning above humeral tibia pits, 3 prealars, 4 scutellars' Legs' L'R' 0'40' both spurs of rear with accessory setae, tibial comb sparse. ÍIing- Length, 1'25 nun' *2*3 *"t s costa Asp. R. 4.2. squama with 4 short setae, anal lobe small' closer ao *4n5, R4+5 finishing above M3+4' costa produced L/3 of' distance towingtip,pfkwelldistalofr_m.Terminafia.Analridgebroadwith Phallapodeme v¡ith numerous setae. sternapodeme thickened, without corners' along gonocoxite large rounded. plate, p.R. 0.1g. rnner lobe stout, extencling nearlytoinsertionofgonostyle.Gonostyleboat-shapedfromdorsalview (FiS' 92)' but drawn up ínto a broad ridge on dorsal surface TYPES.Ilolotypedandlóparatype,EricBonythonReserve,southAustralia,

8. ix. 1971 Reserve' Id Additional sPecimens seen. South Australia: Eric Bonython 8. L97r. f . ix. Lg7o, Ló , 23-ix.L97o, t5ó , ix.

Genus AUSTROCLADIUS Freetnan

Austrocladius Freeman, 196I , P'647 ¡ Fig. B p.283 (in Spaniotoma ( orthocladius) Edwards' 1931, Part)

T\zpe species Campto cladius teriugus Skuse' 1889- to midline wirere MAï,E. Head. Postorbitals and verticafs strong' reaching absent' eyes often 2 or more setae are closely adjacent. Frontal sensilla punp with evenly bare, slight dorsal elongation sometimes present. Cibarial than 0.4' Palp curved dorsal border! tentorium as in Fig. Lo2; M.R. greater than 4th, 3rd more than half length of antenna, segments 3 a¡rd 5 longer cushioneddistallyofwhere4thisinsertedandwithslightlyclubbedsensi]-Ia 2 and' 3 with on disÈal surface. Antenna. A.R. greater than t, flage1lomeres than long; at least 3 type C setae with slightly bulbed bases ¡ 2 - 6 broader D setae as last flagellomere slightty clubbed distallyr with both type c ancl Thorax' well as 3 - 4 ring setae without circlets, sunken setae large. from dorsal view Antepronotal lobes weII developed, junction often obscured acrostichals by scutal process; lateral setae present. Humeral pits oval; Dorsocentrals minute, decembent biseríal and beginning close to pronotum' punctures conspic- strong, erect, beginning just anteriorly of humeral pits, group setae on a raised, uousi in some species the row terminates with a of pre- pre-scutellar tubercle, (Fig.1o3) Pre-alars numerous, single supra-alar' Legs' L'B-' episternal setae sometimes present, scutellars multiserial' greater than 0.5, rear tibia expanded distally and with adpressed accessory two crescents; outer spur setae at least on basal half of both spurs, comb in about 2rnm' Asp' R' often almost as large as inner (Fis.ro4). !'ling' Length high.Sguamawithnumeroussetae,anaflo]¡edistinct.Macrotrichiaonbasal small punctures' vein, basalty on R, Rt and distally ot R4*5' Microtrichia not far plain; R2*3 ttt" midway beÈween Rt and R4+5' costa produced' Pfk finishing at leveL dístal of r-m, cur curved down very srightly distarly, An ofpfk.Abdomen.Tergitesdensely,t..niformlysetose.Sternitelwithout median area sclerotisation or setae. Sternites fITIII & IV with sclerotised sternite v with numerous and number of setae becoming progressively larger, sclerotised and setae and almost completely sclerotised, Vf-vlllcompletely Anal ridge generally setae spreading to cover whoLe of segmentvlll'TeÏTornglaa' small,penapodemepresent,sternapodemenotbroad;spinesmall. fEMAtE. Like rnale but :- always cylindrical and rvith Antenna with 5 flagetlomeres' lst often longest, adpressed type c setae are stight media,L constriction where up to 3 srnall Flagellomeres 1 - 4 found. sunken seta present on each half of segment' r19

Abd'omen' each with t pair, 5th with 2 or more pairs of larger type c setae' setation than male' Bands of sclerotisation broader, less heavy and with less those occupied sternites r - v with unsclerotised spots in same position as by sclerotised sPots in l{asu t-icladius and Kieffe es. Terminalia- TergitelXexpandeddistally,oftenobscuringtergiteXfromdorsalview' sternite X with mediat cleft presenti gonocoxite IX not drawn out distally' genital comb broad ridge. Cerci hemispherical from dorsal view' Setae of mínute. contrary to Examination of slide material of this genus reveals that, the genus does have the observations Freeman (1961) made on pinned material, presence of snall the chaetocladius-1ike characters of strong vertical setae, acrostichal setae and a slight dorsal elongation of the eyes' However' A'R' greater than 1' Austrocladj-us remains separate from chaetocladius since scutellarsmultiserial,penapodemepresentandanalpointabsent.fn setåe addition Austrocladius species have a wetl produced costa, sternopleural

and most species have pre-scutellar tubercles'

KEYToTHEAUSTRALIANSPECIESoFTHEGENUSAUSTRoCLADIUS 1. Outer spur of rear tibia more than half length of inner 2 Outer spur equal to or less than hatf length of inner 4 sguamal 2 (1) ïnner lobe and gonostyle densely setose' trichiatus Freeman setae verY long Inner lobe and gonostyle not heavily setose'

squama with setae of moderate length penapodeme large 3 (2) Inner lobe cupped posteriorly, Inner lobe small, hooked; penapodeme small than 1'5 4 (1) Inner lobe distinct, A'R' greater lnner lobe sroall, inclistj-nct, A'R' less than 1'5 Fig.93 Austrocladius terjugus Freeman, male, apex of palpal segment 3 showing two configurations of palpal sensilla (a anil b).

Fig. 94 male, terminalia rig.95 t! female, base of spermathecal duct.

Fig. 96 Austrocladius rrg3ry, male, wing.

Fig. 97 It lt ll a Terminalia with intromittent organ invaginated; b Terminalia with intromittent organ evaginated: a - anal point Cx - gonocoxites Lb - inner lobe Pe - penapodeme Ph - phallaPodeme St - sternapodeme A/B- Penapodeme Ratio (P.R.) o o ( ( ,Í r I L¡ û 99_ c o o 94 L \ù

f.Ñ ¡\

A B 93

-st i:T .i..i:i{:,,. l. Pc A !, - -'Ph ---a

B ---Lb (b) 97 I20

5 (4) Inner lobe almost free 1yolobus, sp.nov Inner lobe normal 6

6 (s) Squama with setae of varYing size,

penapodeme present numerosus Freeman

Squama with numerous long setaet

penapodeme absent hindmarshi sp. nov.

7 (4) Squama with short setae, numerous sensiÌla

on segment 3 of PaIP, usually with pre- episternal setae terjugus Freeman

Squama with medium-Iong setae, pre-episternals

absent I

8 (7) Scutal tubercle distinct, carrying numerous setae inner lobe with bare corner tubercus sp. nov Distinct tubercle absent inner lobe with small bare digit-like proj ection diqitatus ' sp. nov.

Austrocladius terìuqus Freeman

Austrocladius Freeman, 1961 , P.648, Fig. 8a.

Campto cladius ter-iuqus Skuse, 1889, p-262. A large species, separated from remainder of genus by small inner lobe and short squamal setae and from all other Orthocladiinae since sensilla oÍ palps carried in dilatable saucer-Iike depression. Penapodeme bulbed posteriorly. Al.L South Australian specimens with pre-episternal setae.

MÃLE (4). Heqd. Eles wÍth dorsal elongation 1 facet wide, M.R. 0.53. segrnent 3 of palp with l-o - 15 slightly clubbed sensilla ps¡t) close together on outer distal surface; cuticle carrying sensilla can be retracted in a saucer- Iike depression (Fig.93a) or expanded to follow the contour of the palp (FiS' 93b); palpal formula 0.2 : O.4 : I : 0.9 : 1.3. Antenna. A.R. 1.0 - l-.3; flagellomeres 1, 2, 3 and sometimes 4 with 3 type c setae (2Oy) t last flagellome:re with up to I type C setae (2O¡t), type D setae 25p Long ' Ilgg-+å' setae' Dorso- Àntepronotal lobes just separate from dorsal view, 6 lateral in group of centrals uniserial beginning at level of humeral pits, finishing pre-arars; 14 scuterlars, 2 - 4 setae without tubercle; 10 pre-episternals, 14 Ratio 5'0' squama biseriar. Legs. L.R. 0.6 wing. Length r.9 - 2.2 rnm. r Aspect to *1*2' costa only with 8 - IO strong short setae' R4*5 finishing closer (Fig.94) Anal ridge broad; sternapodeme narrow' sIight1Y Prod'uced'. Terminalia. plates; penapodeme corners slightly reinforced; phallapodeme with triangular only slightly' sparsely bulbed posteriorly; P.R. o.2. Inner lobe projecting subtenninal' protected setoser gonostyle stout, outer surface contoured; spine by ridge.

IEMALE (3). Like male but :- : I : O'90 : I'5' Antenna'' Head. I4-R. about O'5, patpal formula O'3 : O'5 setae; flagelromere formula Flagetlomere 5 with 2 or 3 pairs of stout tlpe c I.8:l:1:1:L.2.Thorax.Prescutellargroupofdorsocentralsabsent. each side; egg-guide bulbous, Terminalia. sternite Vrrr with r0 moderate setae on X from dorsal view' medial with pectinate medial border. Tergite IX obscuring knob' vaginal apodeme one cleft broad; gonocoxite IX with small proximal bald vIlI spermathecal sixth into segment vIÍ, base one sixth as wide as segiment ' ducts broadened basally (Fig'95) ' of fentales from near TlpES. r have examined the lectotype and cotlpe series Sydney in the Macleay Museum collection' Botanic Gardens' Id (at light) Additional sÞe cimens seen. South Australia: ' (oily paper) t7'ix'1970; 15.vii.1970, 2ó (oiIy paper) 25.viii.I97O, td Reserve, Id 19 l5'vii" Brownhil1 Creek, 2d , 2O.vii.I969; Eric Bonython ' ' 29 4'xl'j'Lg6g; Hindmarsh Falls' 1d 2ó , 29 , 28.vii.197Oi Fisher's' ld ' ' l.vii.:tglo,26,39,24.vi.I971;Parawa,19,3O.vii.Lg6g,19,(atlight)' Salt Creek' Id I 15.vii., ld (at light), 5'viii ':|g7}ì Q' L9 '30'vii 'Lg6g; Marne, Nietske's' 2g.viii.1969; Glen view, Hunrbug scrub, Lower Hermitage, Springton,TanundaCreek,Inrilliamstown'numerousspecimensg-lO.viii.L969. In addition' those Victoria: Swa¡ Reach, Id , 22.vii.I964 (O.H. Colless) '

examined bY Freernan (1961) ' LlA

Austrocladius numerosus (Skuse) Freeman

Austrocladius numerosus Freeman' 196l , P.649' Orthocladi us numero SUS Skuse, 1889, P.256' great variation in lengths of Separated from remainder of genus by lobe' squamal setae and by shape of inner elongation 2 facets wide' M'R' o.4L. MÄLE (4). Head. Eyes with dorsal Seg.ment3ofpalpdrawnintopointdistallyandwithuptoTtaperingsensilla (L2¡l)inindividualpiÈs,spreadwidelyoverinnerdistalsurfaceTsegment4 insertedsubterminally;palpalformulaO'25:O'43:I:1:1'3'Antenna' C setae (8Jr)' Thorax' Antepronotal A.R. 1.9; Iast flagellomere with 4 type biseriat, finishing with group of 4 tobe wiÈh 7 lateral setae; dorsocentrals -5setaeonpre-Scutellartubercle;about20pre-alars;aboutlOpre-epister- nalsonallsouthAustralianspecimens'14scutellars,multi-serial.!e99. 2'3mm' t Aspect RaÈio 5'15' Squama L.R. 0.78. V'iing' (Fig'96) Length 2'O - withabout16longandmediumlengÈhsetae,anallobedistinct.Rn*,endin9 half way to wing tip' Terminalia. closer to level of M1+2r costâ produced (Fig.97)Anatridgeslight,coveredwithrnediumlengthsetae.Sternapoderne withslightlyreinforced,squaredcorners,phallapodemervithtriangularplates' setose; gonostyle narrorq with p.R. 0.16. rnner lobe large, triangular, finery slight dorsal bulge and sub-terminaf spine' and the d cotype series from Lawson' TYPES. r have examined the d lectotype Museum' New South Wales, in the Macleay Botanic Parkr fó (at light ) Additional specimens seen. South Australial dd (swarm) 2'x'L969¡ Eric Bonython Reserve' % ' 9.ix. L97O¡ carey's Gully, ód (swarm) 2'x'L969; Parawa' fd (at lighi) 23.i-x.I97O¡ Fisher's, N , LJ'ix' ' 26.viii. :-9]L¡ Robertson", fó 28.vii., 1d (emerged into hillside trap) , ' Id (emerged into trap over Pool) I 2g.x.I969; Scott's creek, 4d , 4'viíi' Lg6g' 14.viii.|969;Williamstown,dó(swarm)IO.viii.Lg6g.AustralianCapital (D'H' colless) ' Territory: Mt. Gingera, ld ' 2o'xi'1960 Fig. 98 Austrocladius kypensis, sp. nov., male, wing.

Fig. 99 tl ll terminalia

Fig. loO il female, terminalia

Fig. 1O1 Austrocladius woodi, sp. nov., male, gonocoxite.

Fig LOz Austrocladius trichiatus sp. nov., male, tentorium.

Fig. to3 It thorax

Fig. 1o4 tt comb and spurs of rear tilcía.

Fig.. 1O5 female, base of spermathecal duct. 96 I 102 1 05 98

ù 101 r¡ T l'

( \ ! l¡ OO C at a ù 3ð' ¡ y¡ ¡---¡ åB Þ Þ 0 I þ 99 o ¿ +c ¿þ & OL o a 104

a tJ rJ

o a o 100 o oo o o o oo 103 t¿J

Austrocladius kvp ensl-s, sp. nov. lobe, outer Separated from other Austrocladius by hoIlowed, setose inner spur of rear tibia almost as large as inner' wide' M'R'0'5' segment MALE Ø\. Head. Eyes with dorsal elongation 2 facets inner distal 3 of palp with about 10 rod-Iike sensilla (8F) close together on palpal formula surfacei segment 4 inserted just before tip of segment 3; 2 - 5 e-ach o.2 : 0.5 : I : 0.9 : 1.35' Antenna. A.R. I.8 - 2.2¡ flagellomeres Antepronotal lobes with 3 type c setae; last flagellomere with L or 2' Thorax' just. contiguous, 9 setae on lateral corners. Dorsocentrals uni-biserial' pre-alars, 18 beginning a]¡ove humeral pits, ending in a grcup of 3; about 14 spur only scutellars, scattered. Legs' L'R' O'75 - O'90' outer tiL'ial wing' (Fig'98) slightly shorter than inner, both with small accessories' 15 tong strong setae' Length 2.7 - 2.9rnm., Aspect Ratio 5.15. squama with t2 - plain. *4*5 midvray between M]*, membrane microtrichia large, punctures "tding tip, cul evenly curved -rd M3*4, costa produced one third of distance to wing narrow' evenly downwards. Terminalia. (Fig.99) Anal ridge low; sternapodeme arched;phallapodemewithovalplates'P'R'O'4'Innerlobelarge'cuppedon posterior surface; style with dorsal border drawn into ridge'

FEMALE (4). Like male but : I'5' Ant-e-nF' Head. M.R. about O.57¡ palpal formula O'3 : O'5 : I : I 4 with 2 and 5 Flagellomeres l, 2 artd 3 each with 3 type c setae distally, setae rtom 2 closely adjacerrt with up to 9r scattered over length; in some cases with much enlarqed peg; bases have joined distally, About 6 ring setae, one : I .27 . Terminalia. (Fig.100) flagellomere formula l'8 : I : O'82 " O'82 s-shaped medial sternite vIII with 16 setae on each side; egg guide with vaginal apodeme border. Tergite IX not completely obscuring X ; cleft broad' VIII. projecting one sixth into segment VII, base one sixth as wide aS Segment

Spermathecal ducts basally narrowed' south Austr:alia' TypES. Holotype d and 29 paratypes, Eric Bonython Reserve, Id t9'vj-j''1969r 2O.v.1970. Further paratypes. South Australia: Parawar ' rd ¡ 2O,tv., L6 , 29 , 28viir97o, 1ó 2lvii.r971' Parawa' 2 ó (at light) 1 Ç Additional specimens seen. Souttt Australia: ' 4'vii'' rod 19 2L'vii'1971' 27.v.Lgl(.,3d,3.vi.,ld,2O-vi'' 26 ' 1? ', ' ', Id !4.v.I962 Australian Capit:'I Territory: Black Mountain (light trap), ,

(I.F.B. Common) .

Austroc Iadius woodi sp. nov.

SeparatedfromtheremainderofAustrocladiusbysnallinnerlobe,Iow P.R. and large outer spur of rear tibia' MÃLE(1).Head.Eyeswithoutdorsalelongation.Segment3ofpalpwith5or over distal surface' palpal formula 6 rod-like sensilla (I3]r) ' scattered O.2:0.35:I:1.05:1'I8'Antenna'A'R'L"73'flagellometes2-Bwith 2 type c setae progressively smaller type c setae, Iast flagellomere with (22¡l)andtaperedoffgracluallytoaroundedpoint.Thcrax.Antepronota]- Iobesslightlyseparatedfromdorsalaspect,3fateralsetae.Dorsocentrals finishing with a group of 3 multi-biserial, beginning to close to anteplonotum' pre-episternars; 14 scutellars' setae on prescutefrar tubercre ¡ 20 pre-alars, 4 uniserial.LegS.L.R.O.5;outerspurgreaterthanhalflengthofinner. !Íing.Lengthl.87nrm.,AsP.R.4.7.Squamawitlr12ntediuml.engthsetae,R4+5 endingcloser.o*I*2,costaproducedonethirdofdistancetowingtip. moderately broad' Terminalia. (Fig.lol) Anal ridge shallow' sternapodeme with triangular plates; corners slíghtly pointed and reinforced. Phallapodeme style boat-shaped.. P.R. O.09; inner lobe small, broad and slightly hooked;

FEMALE (1). Like male but :- LLIa 3/4 of distance along Head. 14.R. 0.47, segment 3 of palp with 5 - I0 sens : I'25: I'5' Anlennal on outer surface; palpal formula O'4 : O'5 :1 VII] with about 20 formulal.6:I:1:Iz2' 2. Terminalia. Sternite Tergite IX not setae on eac}r side, egg guide with broad medial lobe. projecting one fifth into obscuring X , medial cleft slight. vaginal apoderne SegrmentVll,baseonethirdaswideassegmentVlll.Spermatheca].ducts without basal thickening' Austra-lia' 3f iii'197t' TypES. Holotype d and I paratype from Parawa, South r25 Austrocl-adius trichiatus Freeman

Austrocladi us trichiatus Freeman, P'649' by a lcng A large dark hairy species separated from other Austrocladius and wedge-shaped squamal fringe, small anal lobe, an almost free inner tobe gonostyle. MALE(6).Head.Eyeswithdorsalelongation2facetswide'M.R.0.40Tentor- scattered over ium as in Fig.1o2. Segment 3 of palp with long slend'er sensilla anttgc' A'R' inner dist.ar surface; parpar formura 0.2 :0.3 : r z o-7:0.9. last flagellornere 1.75. Flagellomeres 2 and 3 each with 3 type c hyaline setae; Antepronotal tapered distally with 3 - 4 type C setae up to 28p long' Thorax' ending with group lobes not contiguous , 4 Iat:eral setae. Dorsocentrals uniserial l8 pre-alars; 12 pre- of 4 - 5 setae on pre-scuteflar tubercle (Fis'103); about Legs' (Fig'r04) episternals; 18 scutellars, multiserial, pattern distinctive' L.R.0.75'outerspurtwothirdslengthofinner.Wing.Length2.I-2.9mm.; length; anal lobe small' Aspect Ratio 5.1. Squama with 10 strong setae of r:neven closer ao *1*2; costa produced' hatf way to wing tip' Terminalia. R4*5 "rdng a narro\^/ even arch; (Freeman t L96L, rig.8c). Anal ridge shallow; sternapodeme almost free' phallapodeme wiÈh oblong plates, P.R. o.L2. Inner lobe digit-Iike' gonostyle broad basally' bare along posterior border, othenvise densely setose; wedge-shaped, densely setose; spine sharp' subterminal'

FEMALE Q'l . Like male but :- Antenna' Flagello- FIead. MR. O.54, pa1pal formula O'2 : O'3 : I : O'8 : l'3' meresL_4becomingincreasinglyflask-shaped,tlrpeCsetaestout,last : 1 z I'7'l' TLoraå flagellomere with 3 pairs; flagellomere formula 1'54 : I : I on each side; Four sternopleurats. Terminalil. sternite VIII with 25 setae eggguideindentedonmedialborder.TergiteTxwithslightmedialcleft. base one seventh as wide as Vaginal apodeme projecting half into segment vII, (Fis'1c5) segment vIII. spermathecal ducÈs wjth butbous basal thickening ' a¡ci TypES. Holotype ó , Walhalla, Victoria, April 1930' F'E' Vlitson' examined returned to N.M- 2 ó 3'iv'7962i Additiona 1 spe cimens seen. South Australia: Aldgate Valley ' ' Fis. tO6 Austrocladius diqitatus sp. nov., male, wing.

FíS. l-o7 lt I terminalia.

Fig. tOB Austrocfadius tubercus, sp. nov- maIe, gonocoxite.

Fig. Io9 Austrocladius hi-ndmarshi, sP. nov- | male, terminal-ia.

Fig. 110 Austrocfadius lyol-obus sp. nov-, male, gonocoxite. 106

!- \ Itl a, (. I 9'l t, b 108 ,. v o ú J tt r, r,

107

\ e tlt e \ (/ It vbu t VJU 109 110 base one fifth as wide as segment VIII. TYPES. Holotyped from Parar¡¡a, South Australia 19.vii.1969. Paratypes. Brownhill Creek, ld, 29.vii.1969; Deep Creek (Ade1aide Hills), Ló , 29.ví.1969;

Humbug scrub, Ió ,lO.viii.1969; Hindmarsh Fall-s, Id, 24.vLj-.1970, orlando, 7ó, 9.viii.1969; Springton, ld, lO.viii.L969, Torrens Lake' 2ó, 2Q, 15.v., 2ó,26.viii.I970. Additional specimens seen. South Australia'. Alnut, 2ô , 30.vii.1969; Brorqnhill Creek ód (swarm) 29.vii.L969¡ Botanic Gardens, 2ð , 26.viii.797Li Carey's Gu1ly, óé99 (swarm) l.vii.L97O¡ Eric Bonython Reserve' 6d ,22.ív.r 19¿1, 28.vii. Ig7o, 3ô ,I2.v., 26 , 3.viii.IglL; Hindmarsh Falls 8d r 29.vii.I97o, ld , 26.v., Lô , 21.vii. , Iô , 26.viii,1971; Keynes Hill , 26 , 9.viii.1969r Marne Rivert 20ó, 9.viii.L969¡ orlandor 2ó ,9.viii.L969¡ Parawa,2d, 30.vii.1969, dd?9 (swarms) May 197I, Id (reared from hillside divots) 26.viii .I97L; Second Valley ld 30.vii. , Id 2O.viii. L969¡ Torrens Lake dó99

(swarms) May Lg7o, May-June Lg7I, Ió , l2.viii., 1d , 25.viii.L97I. Austroclad :'-us tube.rcus sp. nov.

Separated from otlrer Austroclaclius by nature of inner lobe and well developed prescutellar tubercfe.

MALE (1). Head- M.R. O.4. Segment 3 of palp with 2 tapered sensilla, palpal formula O.2 z 6.4: I : O.9 : 1.O5. Antenna. A.R. L.45, flagellomeres 2 - 5 each with 3 type C setae, last flagellomere not clubbed., tip rounded. Thorax. Antepronotal lobes wíth numerous lateral setae. Dorso- central-s multi-biserial ending in tubercle with numerous setae; 16 pre- alars; scutel-lars in 2 rows. Legs. L.R. O.78, comb straight, setae dense. !'lings. Length 2.O5 mm. Asp. R. 4.45. Ànal lobe small, squama with 1O moderately strong setae. *4*5 ending closer to leveL of Ml+2, costa produced half way to wing tip. Terminalia. Sternapodeme with s1iøh.tly thickened corners, phallapodeme with narrow pI.ates, P.R. O"28. Inner lobe small with rounded bare corner (fig. 1O8), Gonostyle with slightly concave inner surface drawn up dorsally. TYPE. Holotype d from Orlando, 9.viii.1969. I¿ö

Austrocl-adius hindmarshir sp. nov. Similar to terjugus but separated frorn it by the structure of the inner lobe of the gonocoxite and, the spine.

MALE (1). Head. Frontal sensil-Ia well separated, M.R. O.43. Segments 3 and 4 of palp each with clump of up to IO slightly capitellate sensilla, palpal formula O.3 : O.6 : I : 1.25 : I.45. Anterta. A.R. 2.OO. Flage11o- meres 5 - 8, 10 - 12 each with 3 type C setae (28¡t), 2 - 12 broader than long; last flagellomere with 4 type C setae (22y). Thorax. Antepronotun with several lateral setae. Dorsocen+-rals tri-uniserial, ending in tubercle, 16 pre-alars, numerous biserials in 2 rows. Legs. L.R. O.68. Wings. Length 1.58 mm. Asp. R. 4.6. Anal lobe very small , squama w-ith 16 long strong setae. *4*5 ending midway between Mln2 .td M3+4, costa produced I/3 distance to wing tip. Terminalia (nig. 1O9). Sternapodene with rounded, extended corners, phallapodeme with oblong plates, penapoderue absent. Inner lobe broad, rounded. Gonostyle with surface drawn up distally into a ridge which partly obscures the spine. TYPE. Holotype d from Hindmarsh Falls, 20.v.I97o. Austrocladius lyolobus, sP. nov- Readily recognised since inner lobe of gonocoxite almost free, but, unlike that of trichiatus carries only a few short setae.

MALE (t). Head. Frontal sensilla present. Cibarial pump with straight dorsal border. Segrment 3 of palp with several slight.Iy clubbed sensilla (rrp¡, palpal formula O.3 : O.4 : I : 1.15 : I.45. M.R. O.32. Antenna. A.R. I.g. Flagellomeres 2 - 12 broader than J-ong, last ftagellomere with 4 - 5 type C setae (2O1¡ and 4 large ring setae. Thorax. Antepronotum with 6 - 7 lateraL setae, dorsocentrals multiserial, uniserial punctures smal-I, 9 pre-alars, many scutellars in single :cow. Wilgs. Length 1'37nim'

Asp. R. 4.5. Squama with to strong setae, anal lobe smal-l. Rn*u ending closer ao trnn, costa slightly produced. _l"gr. L.R. 0.62, comb without 2 crescents. Termina]-ia (riS. IfO). Sternapodeme highly arched, corners just thickened. Phallapodeme with triangular plates. P.R. O.15- Irtner Fig. 111 Linnophyes tu-bercus sp. nov., male, head.

Fig. II2 thorax, showing spatulate setae.

Fig.1I3 I l! wing

Fis.1I4 il terminalia

Fig. 1I5 Limnophyes pelurgis, sp nov.. male, setae of humeral pit.

Fig. 116 il wr-ng

Fig. LL7 tl It terminalia Íf

.74

117 111

112 0

a '--->p o a

=æa

àà I llò ì

o 114

113

I 15

116 lobe almost free. Gonostyle boat-shaped. TYPE. Holotype ó from Botanic Gardens, 24.xí.I97O'

Genus LfMNOPHYES Eaton

Limnophyes Eaton, 1875, p. 60; Brundin, 1956, p- 13I; Freentan, L96L, p. 654¡ Saether, 1969, P. IO1.

Type species Limnophyes pusillus Eaton. In redescribing this genus Brundin (1956, p.13I) omitted to comment on a number of characters. The following additions are made on the basis of my examination of Australian species. MALE. Head. Cibarial pump with curved dorsal border; palps with sensilla on slightly dilated inner distal surface of segrment 3. Vlingr. Macrotrichia present only on basal vein and R.

Two species, Paralimno s pullulus (Skuse) and P. albibasis (Freeman), were incorrectly placed by Freeman (1961) (see p- I32) .

Limnophyes tubercus, sp. nov. A btack species; flagellomeres almost globular, fibrillae welf dis- played; numerous spatuÌate setae on humeral and posterior regions of scutum; separated from all other Orthocl.adiinae by its large frontal tul¡ercIes.

MALE (4¿. Head (fig. I11). Verticals medium síze, uniserial, exLend to midline. Frontal tubercles large flattened, U-shaped, finely setose, with terminal sensilla. M.R. O.33. Segment 3 of palps with one straight sensillum without basaf pit, palpal formuLa o.3 : o.4 : I : I.5 : 2.O. Antenna. A.R. O.4. Flagellomeres I - 12 91obu1ar, 2 and 3 each with 2 type c setae. Last flagelLomere slightly clubbed ancl pointed apically, with 5 - 6 type C setae and 3 ring setae" Thoraë (Fig. L!2). Antepronotal lobes with about 6 setae on lateral corners at least 2 of the¡n lanceolate. Dorsocentrals irregularly biserial, begin close to antepronotum; humeral area covered with spatulate setae about 9O¡r long and a bi-tri.serial row of similar setae medial of dorsocent:rals on posterior quarter: of scutum; 130 posterior anepisÈernum II wiÈh 2 seËaer pre-episternum with 6 - B setae; 9 pre-alars, 6 scutellars. Legs. Rear tibial spurs with small accessory setae. L.R. about O.5. Wing (FiS. II3). Length I.1 - I.35 mm. Asp. R.3.6.

Sguama with 2 setae. Microtrichia very sturdy giving wing a dark appearance, costa produced one fifth of distance to wing tip, *4*5 fitishing at of M^... Abdomen. Tergites uniformly setose. Sternite I without Ievel 5++ setae but medial setae increasing on subsequent segments to cover whole of sternite VIII. Terminalia (Fig. 1I4). AnaI point absent, sternapodeme with distinct corner processes. Gonocoxite with 2 inner lobes, one large, dorsal, ttre other smaller, ventral. Phallapodemg witJ. large rounded plates. penapodeme absent. Gonostyle boat-shaped, spine normal. TYPES. Holotype d and f ó paratype, Hindmarsh Falls, South Australia, 1ó I8.xi .Ig7O. Further paratypes: Hindmarsh Falls, 1ó , 25.iii., '

15. vii .L97o .

Additional spe cimens seen. South Australia: Hindmarsh Falls, 4ó 20.viii,, 2ó , r.x., Id , 15.x., ld , 31-iii-1970. Limnophyes pelurgis, Sp. nov. A black species with slightly fusiform flagellomeres and a distinctive ro\¡¡ of 3 tiny spatulate setae just above the humeral pit. Frontal tubercles much smaller than in tubercus-

MALE (3). Head. Single vertical seta on each side of midline. Brow drav¡n out into slight tubercles with terminal sensilla, M.R. O.35. Segment 3 of palpwith2or3sensilla;palpalformulaO'4:O'5:I:l'I5:L'7' Antgnna. A.R. 0.6. Flagellomeres I - 12 fusiform; Io - t2 incompletely separated¡ 2 and 3 broader than ]ong¡ 2 - 5 each witjl 3 type c setae. Last flageflomere clqbbed apically. Thorax. Antepronotum wiÈh 3 lateraf setaei dorsocentrals biserial, beginning close to pronotum. Row of 3 - 4 spatulate setae (33¡¡) just above humeral pits (nig. II5); spatulate setae (46p) in single row parallel to and medial- of posterior dorsocentral.s; pr:e- fI episternum with row of setae at anterior margint posterior anepj-sternrrm Fig 1.18 L ES pelurgis, female, terminalia.

Fig. II9 tt larva, ventral view of head.

Fig. L2 { ll larva, labrum, dorsal view.

Fis. L2D I tt anal segments.

Fig. I22 Paral-imnorrhves pullulus (Skuse), ma1e, wing. 118 : 120

oQo I 1n:

119

122 L3r with single seta near base; anepimeron II witJ- single seta near mi-ddle;

6 pre-atars, one anterior and one posterior groupt 6 scutellars. W.ing (rig. 116). Length 1.3 - 1.6 mm., Asp. R. 5.O. Squama with 2 setae, anal lobe very small. Microtriclria stout, giving wing dark appearance, costa produced one third of distance to wing tip. Rn*U finishing at level midway between Ml_*2 .rd t3*4. Legs. Spurs of rear tibia without accessory setae, L.R. O.5. Abdomen. Tergites uniformly setose. Sternite I without setae, but medial setae increasing on subsequent segments to cover whole of sternite VIII. Terminalia (Fi9. l-I7). Anal point absent. Sternapodeme with corners extended. Phaltapodeme with cu,rved triangular plates, penapo* deme present, P.R. O.2. Gonocoxite with 2 inner lobes' one dorsaL pointed' one ventral, reduced. Gonostyle boat-shaped with dorsal distal border produced into a coïner. Spíne large and tooth-l-ike.

FEMALE ( 3). Like male but:-

Head. Segment 3 of palp with single tapering sensillum, curved at tip; palpal formula O.4 : O.5 : 1 z I.25: I.75. Antenna. All 5 flagellomeres with 2 type C setae (22¡t); flagellomeres f and 5 cytindrical, 2 - 4 flask- shaped, necks short; flagellomere formula 1 : O.9 : I : I z L.7. Thorax. row Antepronotal lobes with 4 small setae on lateral corners and "';; setae along dorsal third. Legs. L.R. 0.6. V'Iing. Length I.27 mm, Asp' R. 3.8. Macrotrichia on R, Rl utd R4*5. Terminalia (nig. II8). SÈernite VIII with 6 - I setae on each s-ide. Egg guide flat, evenly rounded' Tergite IX with median furrov/, gonocoxite IX narrow, produced posteriorly, with 2 - 3 strong setae. Sternite X with small ridge, cerci globular frotn dorsal aspect. Vaginal apodeme projecting one sixth into segment VII, base one fifth as wide as segment VIII.

LARVA (2). Up to 4 mm. long, colourless. Head (fig. 119) as long as broad; single eye spot almost divided into sma1l anterior and large posterior sections. Antennal formul-a I: o.23: O.ll: o.o4: o.o1 , antennal seta on basal segment broad, almost as long as Jcasal segment.

Mandjlcle with 5 teeth; internal- setae with numerous fJ-agelfa; realî setae l.32

(Rb, Hirvenoja Lg73) normal but in addition there is a depression on the ventral surface bearing numerous fine flagella. Labrum (fig. 12O) with (shown IV, V slender. seta I flattened, seta II also in Fig. 1I9) ' III' Remainder as shown in figures. Setae of abdominal and thoracic segments minute; anal papillae and pre-anal setae as shown in Fig. I2L. TypES. Holotype d and ló, 29 paratypes (reared from silt) Happy Valley, October, Lg6g. Further paratypes: Happy Va1ley ld, 19, October, 1969¡ Torrens Lake, Id, I5.vii.L97o.

Genus PARALII.O{OPHYES Brundin (emended) Paralimnophyes Brundin, 1956' p. L29.

.éLimnophves (l-ongiseta) Thienemann, L944, p. 571. Camptocladius (Hexatomus group) Thienemann, I92I, p- 8Io. Type species Paralimnophves hydrophila Goetgh. (longiseta Thien. c Kieff.) In his description of this genus Brundin omitted to comment on a

number of characters. The following additions are made on the basis of my examination of Australian species. MALE. Head. FronÈal sensilla present; segment 3 of palp with sensilla on ditated inner surface. Antenna. Flaqellomeres 2 - L2 slightly fusiform; flagellomeres 2 and 3 broader than long, each with 2 tapered type C setae; last flagellomere slightly clubbed distally with several type C and numerous type D setae. Thorax. Sing]e supra-alar seta' no pre-episternal-s, 6 scutellars. Wing. Macrotrichia on basal vein and R, microtrichia strong, punctures large. Abdomen. Tergites uniforrnly setose; sternite I without setae, sternite II with 2 setae; subsequent sterniÈes with increasing numbers. sternite VIlt completely setose. Terminalia. SternaPodeme with coïners drawn out. Gonocoxite v¡ith two inner lobes: one dorsal and' proximal, the other ventral and distal.

Freeman (196f) placed the two species pullulus and albibasis (redescribed bel<¡w) in Limnophyes þecause on the dried material he examined the antenna lackecl a straight apical bristle. Ilowever' in fresh material Fig. l-23 Paral s pullulus, ma]e, terminalj.a.

Fig. I24 tl tt female, terminalia.

Fig. J-25 Paralimnophyes albibasis, Freeman, male, wing.

Fig. L26 tl It il terminalia.

Fig. I27 I pupa: ventral- view (I.h. s. ) and d.orsal view (r.h.s.) of anal lobes and pre-anal segment. 123 )

09 1

0 t, l¡ 126 l¡ -/' \1 ,tl tI

I I ì tt I ,ì I ,tì \ I I I ,lr I I , t I I

\ 1 t\ r7,

124 127

125 133 t¡.is bristle is present, although easily lost. The presence of an apical bristle together: with an anal point and an only mod,erately developed pro- nottur indicates Paralimnophyes is the more appropriate genus. The exceptionally long precorneal setae of the pupa as well, as the purple colour and. long lateral setae of the larva support this designation.

Paralímnophy es pul-l-ulus (Skuse) comb. rrov. Orthocladius pullulus Skuse, 1889, p. 259.

Limnophyes pullufus Freeman, 1961' P. 655. A black species, separated from most Orthocl-adiinae because the gonostyle of the mafe is extended well beyond the spine. Segment 4 of palp with smaller proximal sclerotised point than al-bibasis. Female cerci with 3 setae, one and a half times tength of cercust pupa with two of precorneal setae flattened.

MALE (3). Head. Single medial vertical seta, frontal sensiffa on short, broad tr¡bercles. Palpal seqment 3 with 4 close-set straight sensory setae (l3P), without pits; segment 3 inserted beyond proxiroal horny point of segment 4. Pal¡ral formuta O.4 : O.5 : I z L.2:2. Antgnna. A.R. 0.6. Last flagellomere with 4 large ring setae. Thoral<. AcrostichaLs minute' decumbent, biserial. Dorsocentrals begin at leve1 of humeral pits, 3 pre- alars. Wing (r'ig. L22) . Length I.6 - 1.8 mm. Asp. R. 5'O' Squama with 3 setae. AnaI lobe very small, microtrichia 3¡r long. costa produced half way to wìng tip. cu, slightly curved, tip turned parallel to r^zing ma:rgin'

Legs . L.R. o.5 - o.6. Terminalia (r'ig. 123). Anal ridge prominent, anaf point bare, one third length of gonostyle. Gonocoxite with large dorsal lobe, smaller, rounded ventral lobe. Phallapodeme with almosÈ circular plates. Gonostyle extended beyond spine into broad point.

FEMALE (2). Like male but:- Head. Frontal sensilta widely separated, segment 3 of palp nearly spherical' with 3 slightly cfubbed sensory setae. Palpal for¡nula O-4 : 0.6 : I z I'2: twc 2.5. 4qtenna with 5 flagellomeres, 2 - 4 swollen medially, I - 4 with largetypeCsetaeandonerowoffibriltae'6thwit.h3typeCSetae, 134 two ring setae, numerous type D setae and a short subterminal bristle. Ftagellomere formula I : O.g : O.9 : O.8 '. L.4. Thorax with dorsocentrals biserial- beginning close to antepronotum. Legs . L.R. O.54. Terminalia (Fig. L24). Sternite VIII with sparsely setose genitaÌ comb' e99 guj-de elongate. Tergite IX small, gonocoxite fX extended into narrow posterior lobe. Tergite X with broad ridge; cerci rectangular from dorsal view, 3 long setae on outer distal corners. Vaginal apodeme projecting one quarter into segment vII, base one quarter as wide as segment vIfI. pUpA (2). Cephalothorax.. Thoracic horn reduced to cone IO¡r high. Three pre-corneal setae: anterior, 22O¡t Iong, flattened, 5¡ across at base; medial, I5O¡:, flattened; posterior, AOF. One anterior and one posterior pronotal seta, 4 mesonotal setae. Abdomen. Tergites covered wíth strong spinelets; along posterior border of each segment, 3 or 4 rows of these are extended into long points which overhang the following segment. Sternites covered with short microtrichia. Setae ,l - ,4 as long as or longer than segment length. Segment 8 with additional L seta close to I'4- Anal lobes as in P. afbibasis (Fi9. r27).

IARVA. Described by D. H. Edwards (thesis, 1962, p. 24) . No further material available- TYPES. Holotype I from Sydney (examined L972) M.¡4. Additional specímens seen. south Australia: Alnut, dd99, 30.vii.L969¡ Bool ód99 Lagoon, 15d,27.viii.L969¡ Botanic Park, ió,9.xii.I97O¡ Brown's Lake ' 26.vi.ii.Lg6g; Duck Ponds, L6, 28.íx.I969¡ Ewen's Ponds, #, 26.viii.1969¡

Hindmarsh Falls, Iô, 23.i.L97O¡ Inman Valley, 3ó, 3.ix.L969¡ r'eg of Mutton

Laker 6d, 26.viii.:-:969, fó (reared from algae) 15.ix.L969¡ Kuitpo, 2d , I.vii.1969; Moculta, ód99, Aug. - Sept. 1969, Ló (in pupal case) 299

(reared from Cha¡e) 3O.viii .1969; Marne, dó99, 9.viii .Lg6g; Morialta, 1ó ' 4.ix.Lg6g, Id, 7.xt.L969¡ Nietske's 2ð (light trap), 9.viii.L969¡ piccaninnie Ponds, Ió, 26.viii.1969; Parawa, ß, 19.vii., Id, 3O.vii.1969¡

Q., 1d, 30-vii.1969; Reynelfa, 2ó , 7 -xí-1969; Tantanoola Creek, 3d,

26.viii .Lg6g; Tanunda Creek, 2ó, g.viii.Lg6gi Torrens take ód99 r I35

Mar. - April, Sept. - Nov., L97Q¡ Valley Lake, 1ó, 19, 26-viii.I969- River Broughton, Merriton, 2ó, 26.viii.1969 (8. Glover). Western Australia: Pemberton, Id (at light) , 6.x.I97O (o- H. Colless). Paral-imnophyes alblbasis (Freeman), comb. nov.

Lfunnophyes afbibasis Freeman, 1961, P. 656. A black species, quickly distìnguished by its white wing bases and Iarge sclerotised point on irurer proximal surface of palpal segment 4. MaIe with row of 3 small distally directed setae on inner surface of gonocoxite, female with longer setae on cerci than pullulus, precorneal setae of pupa not flatÈened.

MAI,E (3). Head. Single medial vertical seta present, frontal sensiJ-l-a on small tubercles. Segment 3 of palp with 4 straight sensilla often incom- pleteJ-y separated from segment 2, inserted beyond proximal point of segment 4. Palpal formula 0.6 : O.9 : I : I.3 | 2.L. Antenna. A.R. 0.6. Last flagellomere with 5 ring setae. Thorax. Acrostichals minute, decr:nbent' biseriat; dorsocentrals begin above humeral pits. Three pre-alars' Ialing (Fig, :.25). Length I.4 - 1.75 mm., Asp. R. 4.I. Squama with 2 setae, anal lobe sma1l. Macrotrichia on Rr, microtrichia 6p long. Costa produced half

v¿ay to wing tip, Cul slightly curved, tip turned parallel to wing margj-n.

Legs. L.R. O.6 . Terminalia, (nig. :.26) . Anal point with basal hairs; gonocoxite with equal sized dorsal a¡rd ventral inner lobes; 3 short distalJ-y directed setae on inner surface just proximal to insertion of gonostlzls' Gonostyle truncate, spine smalI, terminal.

FEMALE (3) . Like ma-le but:- Flead. Frontal sensilfa widely separated, palpal formula O.3 : 0.6 : I : L.2 z 2-O. Antenna. Similar to pullu1us, flaqellomere formula 1'O : o'7 : O.75 : O.7 : 1. Thorax. Dorsocentrals biserial , beginning cl-ose to pronotum. Terminalia. Sternite VIII with dense genital comb, egg guides elongate. Tergite IX small, gonocoxite IX broader than pullulus' Tergite X \^rith broad rioge; cerci oval from dorsal view, with 3 long distal seLae' Fig. I28 Sparsi-cladius btgis, sp. nov., male, tentorium.

Fig. L29 I tf tt termi.nalia.

ríg. 130 I lr pupa, dorsal t¡iew, anal lobes and pre-anal segment (2 views of anal setae) .

I'i.9. 131 Sparsicladius telmatus, sp. nov., rnale, wing.

Fis. L32 tl il l¡ te:rminalia.

Fig. 133 Sparsicfadius polysetosis, sp. nov., male, terminalia. 131 12E^ 132 0, L 129

'¡ yt + , ,l'r.. ,l ,l { 7 , t, r rl v YI Y I 'r , o IV v l I / I u r/ v luu V I v v \\ / \ i v I rl'rI '/ I, I r .\ I f1 { a i ì I .t ¡ . rt I 1 .\l (t t,-Lt. ¡

J I 133 ûl 130 r36

VagirralapodemeprojectsonethirdirrtosegmentVlI.,haseonequarter:as wide as segnent VIII.

PUPA '¿ Like pullulus but:- cephalothorax. precorneal setae not flattened, Iengths r8oP, I8o¡-r and 8o¡t; 2 Large anterior and 2 short fine posterior antepronotal setae' Abqglnglt-' AnaI lobes as in Fig- L27.

I,ARVA (2\ . LiKC Lj-mnophves Pelurqis but:- formula About 5 mm. Long, violet or dark blue i¡¡ colour. Head. Antenual I : O.32: O.I4 : 0.06 : O.O3, ant-ennal seta about as long as basal- segment, slender. Mandijcle \^rithout ventral seta-bearing depression. Abdomen. Lateral setae longer than segment length. Pre-anal setae more nuneTous; dorsal setae of segment X longer than posterior prolegs' TypES. I have exanrined the holotype d and 59 paratypes from National Park'

New South Vüales, August, 1953, E' J' Reye (Spnf¡l) ' Bonython Additional sp ecimens seen. South Australia: Botanic Park' Eric (in Reserve, parawa, numerous specirnens, JuIy 1969 - Mar. L972' ail traps AJ'nut and reared from most material taken where surface water flowi'ng); Id, 3O.vii.I969; Deep creek (south coasÈ) 2ó, 49, 3.ix.L969¡ Healey's ld, I8.x.L969¡ Hindmarsh Falls dd99, sept' - Nov', 1969' IgTo' L97L' 1ó'

Ló 3'ix' lô' f .iv. Lglo, 19, 17.vii -L97L¡ fnman Valley 1ó, 2O'viii' ' ', ' 59' 26'xj-'r969¡ r.x.1969; Q.,26,3o-vii.Lg6g; Robertson's 2ó, 19' 12'xr-'' Scott,s Creek, Ió , 4.í2.L969¡ Second Valiey Ió, LQ , 3o'vii' ' Id ' 20.viii. Lg69¡ Tanunda creek, l.d, 25.x.rg6g¡ Tooperang 1ó, 15-x.1969. Vicboria: Ben Cairn , L6, lo.xi 'L964, l{' Dobrotworsky'

Genus SPARSICI,ADIUS , gen' nov '

Type species Spar sicladius Þ:¿gUt sP. nov. verticals di¡ninishing in size towards ul]JÄ. Head. Post orbitals present, miclline i-¡r niclline. Frontal. sensifla absent, broTv ext-encled velttrai]-y along Tentorj'urn as in some species; eyes bare, dorsal elongation I face+- v¿ide' pits on l.is.. Lze. segment 3 of palp wit-l:' several sensilla in indiviclual t.3 I dilated inner distal surface. Antenna. A.R. variable; flagellomeres 2 and 3 with 3 type c setae (2O¡r); last flagellomere with 3 type c setae' 2 minute terminar sunken setae, 3 ::ing shaped setae; distal- third without fibrillae but clothed with nunìerous fj-ne haiirs. tþggt- r'lnteprcnotum normally developed, lobes slightly separated by larç¡e scutal process; several lateral setae. Acrostichal-s uniseri.al , short, fine and clecumberrt, beginning close to pronotum; dorsocentrals biserial anterior of humeraL pits, uniseriai at rear, punctures laIge' up to lo pre-alars; sctrtellars cotnb, cuter spur uniserial . lgg!_. Rea:: tibia expancled oistaJ'Iy, carryi-ng less than half tength of inner, inner with small basal accessory setae; tarsus 4 about equal to 5; L.R. o.5. !¡-i'rg.. I,ength o.95 - }.7 nm. Squanna ou R' with one or 6 - B strong short setae; ana-l lobe small' Macrotrichia microtrichia long, fíne, punct-.ures large, R2.i3 joining costa midway between dj-stal of Rl_*2 .rd R4+5, costa not or scarcely produced; posterior fork Sterriite r-m. Ctr' bent evenly clown, An finishing at level of r-m. aÞqg*s". along råd^ I wj-ttroul- seLae, sLezDites II - v with incr-'easj-ng nurnJcers t¡f setae point l()nq' Iine, sLe::Ðites vI - VIIT cove¡.'ec with set-ae- Ter'Finqi-ia' Anal- sclerotised; penapoileme stout, gonostyle with medj'um size sr':l¡terrninal spine'' Gonocoxite r/IIl broad' FBMA-LE.. 49LgIn1 with 5 flagellomeresi' Term-inalia' not ol>scu:r- egg guicles evenly rou¡rded. Terqite Ix with small' med-ian cleft' ing terqi-1:e X. Cercj- hemispher-ì-cal- frotir dorsai view' Vaginai apcde:me çro- vlll' jecting iD1-o segm-ent \¡Tf , base about one third as wide as segrnent Precorneal setae PilPA. Cepfiafo 'chor;rx. Prothoracic horn srn'all , wart-lil

bl ade-Ii,ke cebae. rnicro- Adults of this genus are quickly distiriEu-ishecl because ihe fj'nc¡ trichia in clistinci- punctures give tire wing a speckl-c:cl appearance ¿ì'b ic)Dx f rorn ggIq"lt"'qqo--c-Lgg4E Freentan niat¡nificatior¡' 5-i,-¡:1'=r;-i*i¿tl!Lt- -is sepa::ated anal (which must be renamed) * because it has a small pronotum and a bare point. ThepupaeofsParsicladius,althoughreadilydistinguishedbythetwcl blade-like setae of the anal lobes, show affinities with those af many semi-terrestrial and terrestrial species (see Iv.6.2). Thegenericdiagnosesofthefemaleandthepupagivenabovewill probably prove incomplete as they are based on single species'

KEY TO SPECIES OF THE C'ENUS SPARSTCI'ADIUS

2 I Squama with more than one seta, anal point large' stout 3 Squama with onlY one seta 2 (l-') Gonostyle with proximal hump, dorsocentrals bi-uniserial nov" polysetosi.s, sP. nov. Gonostyle broad, dorsocentrals multiserial sP. nov. 3 (I) Arral point large, stouti gonostyle broad' naucise+-osis, Anal point long, slender; gonostyl'e slender 4 4 (3) Inner lobe rounded, small; anal point tapered sp. nov. to sharP Point bryoryÞ, Inner lobe broad, hammer headed; anal point long, slightly flattened distally" " telmatus, sP. nov.

Sparsicf adius bryonis, sP- ncv. Separatedfromremainderofgenusbysimplestructureofinnerlobeof

gonocox-ite and qonostYle. MALE(B).Head.Brorvslightlyextended'cibar:ialpumpwithalmoststraigh-'- (11¡:) palpal dorsal border. segment 3 of patp with 6 - I clubberl sensilla ' not clubbed' formula- o.4 : O.53 : I : I : I.g. Antenua. Last flagellomere pre-a1ars Tlrorax. Antepron.otar robeswith 3 - 4lateral setae; 6-7 slencler¡ arranged in 2 crescents. f¡c+9s. Inner spur of rear tibia straight'

o chael?g iqq *Vüulkcr (1959) des cribed a subgen us P.9¡; q-c-þ3i-1t-9g leÈ+-"- f þr1 co abtrobeus. Sae tttei (tgc-g) elevated this subgenlls wiLir tYPe s¡>ec-ies therefore be rena:ne'¡" the rank- cf genus .--Pä.-..tt"etoc1 adius Freeman must 139 cornb with short closely set setae. wing. Length I.o - 1.4 mm. Asp. R'

4.27. Squam.a with singfe seta, anal lobe indistinct, *4*5.tding closer to M^ Abdomen. Tergites with 2 rows of setae Terminalia (Fi9. L29) . 3+4,. Anal point witir strong basal setae, tapering rapidly to long, fiue point' coxapodeme weakly linked with sternite X; sternapodeme narrow, corners drawn out and reinforced; phatlapodeme with long narrow platest P'R' O'15' Inner lobe evenly rounded; gonostyle boat-shaped, proxima'I inner surface dilated with 3 large setae-

PUPA (1). Cephalothorax. Prothoracic horn 4u high' Precorneal setae: anterior 3O¡rr rniddle 80¡, posterior J-O¡r. Two anterior (8Op) and one posterior (5o¡:) antepronotal setae, 2 pai'cs of scutal setae. @g:9"' posterior spinules of tergites progressively increasing in length from tergi,te I to VII. Setae of anal lobes 12¡r long¡ gonopod sheaths extendilçt well beyond anal lobes in male (nig' 13o) ' TYPES. Holotype d Botanic Gardens, south Australia' II'vi'1970' Par:rtypes' south Australia: parawa, 2ó, 3O.vii., Id, 2o.viii.1-969, Lô, (emergence trap at positiol (iv)) 2I .x.7.91O¡ Botanic Gardens, Iô, l-iv., Ió (emergence trap over grassy clearing) 28.v-Lg7O, ld with pupal case, I pupa (grass divot) 14.vi.L972. Addit_ional specimens seen. South Australia: Nuruerous specimens !7ere Èaken throughout the year from Botanic Gardens, llric Bo¡ython Reser:ve, Parawa in L969 - L972¡ Healey's,Ló,lS.x.1969.AustralianCapitalTerriiory:]3].acii Mountain , Lð, 12.xi.l-965 (o. H. Colless). Sparsicfadius t<:flC!_"s_, SP. nov. its long, heavily Quickly separated frotn all other orthocladiirrae by sclerotisecl , Èerminally broadened, anal poin+* and harruner-headed inner fobe' side of midiine' MALE (3). i]gq. Brolr' extended i¡rto smafl knob on each O'25: : segment 3 of palp with 2 - 3 stout cluJrbed sensilla, paJ-pal formula' o.5:1:l:L.2.åutellg.Lastflagellonre::ewith3extratypeCsetae' slj-ght1.yclul¡br':dapically.lÞo_Iå":.six-7pre-'alars,u:riseri;rl.I.'99:' 140

Inner spur of rear tibia stronq, sinuous, comb with long, slender setae. vting (r'ig. 13I) Length I.o - I.3 mm., Asp. R. 4.4. Squama with single setai compared with bryonis, R veins and costa end further distal and' wing tip narrovler. eboo*u". Tergites sparsely, uniformJ-y setose. Terminal'ia'

AnaJ- point long, widened distally and with small basal setae. Sternapodem-e narrow, corners reduced, transverse aÏm broader; phallapodeme with narrow triangular plater P.R. O.14. lnrrer fobe broad with hammer-Iike head; gon.o- style with inner proximal surface dil,ated to form bare peak; dorsal surface forming a narrow riclge ,tist.ally; spine subterminal .

FEM.ALE (8). Like male but:- HeaÈ. Vertical setee not reac:hing midl-ine, palpaf formtrla o.4 : 0.6 : l' : 1 : 2.2. èrt"lq,.a. Flagellomeres 2 - 4 fusiforrn, 5 strongly tapered dist"rl- Iy; 2 - 4 each with two, 5 v¡it-h 4 peg^like type c setae; 5 rvith 3 ring setae ancl 2 short terminal bristles; flagellomere formula 1.7 : I : 1'l: I : 2.O. gS.. Length about O'85 mrn', Asp' R' 3'1' Legs' L'R' O'4' Terminal-ia (f ig. I32). Sternite VIII with 4 - 7 setae on each side, gorìo- coxite I)i Ì.¡roa,dly roundeó, sternite X with sma'll- riclge. Vagina'l apodeme projecting half into segment vfl, spermathecae with basal broadening' TypES. Ho-lotype ó and Ió paratype, Parawa, South Austrafia, 19'vJi'1969' Further parabypes. Parawa, Ió, 2O.viii.1969; Blewitt Springs, 29, 2.viii.L972. Àdditiona I specirnens seen. A1nut, fd, 30.vii. L969¡ tsotanic G¿rrderls , Lô , 2ô 9'íx' 4.ii. , Ló, i-9, L2.xi.Lg7o; Eric Bonython Reselîve, ú, 28'vii' ' ' ' 26 ld, I8.xi., rd, 2.xíi.Lg7o, 7ó , 17.ij'-, 2ó , 3.iii', ld, 28'j-v" ' 26-v. rglr¡ ltearey's 26, 18'x' r9'o9; Parawa' ód99 (emergence trap a¿ habitat B) Feb. - May, dd?9 (emergence trap a.i hab-itat' Ai Auq' ód9? (¡re1:teC) l¡eL¡. - Dec.

Spa rsicladirrs polysetosis, sP. nov. po.lvsetosis pauciseLosj-s and ctrar¡j-uis share a lor:q The Lhree sPecics ' stout ;rnäl point; poiy:;etc>sis is readily separated by ttre mu]-Èi-ser'i'af 141 dorsocentralbrist.Iesandpresenceofmorethanlsquamalseta. brow MALE (5). Head.. Verticals finishing in group of 3 at mid]-ine; slightly extended ventrally. M.R. o.33. Segment 3 of palp with 3 slightJ-y capitellatesensilla;palpalforrn.ulao.3:o.5:lzl-.2=I.7.Antenn?-. A.R. l.O - 1.3. Flagellorneres 2 - l-2 broader than long. Type C setae 2 l¡roadened one third of distance from base, sunken setae on flagellomere I¡iith 2 enlarged, last flagellomere heaviÌ12 clubbed and squared distally, type c setae; type D setae 35¡r long. Thorax. Antepronotr'rm with about l-o only dorsalry directed l-aterar setae. Acrostichars biseriat, extending into half-way aloug scrrtum; dorsocentrals begirr close to pronoturrr' spreacl row; 7 LO pre-al'ars' 3 - 4 ro\^7s and contract posteriorly to a biserial - setae' 6 scutellars" Legs. Inner spur of rear tibia r+ith fine accessory Wing-. Leugth I.3 - 1.7 mm' Asp' R' 4'2' Squama with 6 or more setae' uni- *4n5 .tding closer to Ml*2, costa not produced' Abdomen' Tergites rvith fine formly, densely setose. Termrnalia (FiS. I33). Anal point large' with tri* basal setae. sternapodeme broad, corners extencleci; pharlapodeme c¡c'rflo- angular plate, P.R. o.IB. Inner lobe broad, right-angled, setose; stytewithinnersurfacesliglrtlybu]-bed,outerst¡-rfaceextendedintoa ridge. ix'197o' TypES. Holotype d and 1d paratype, Para',^¡a' South Australia, f 2'ó' 2o'viii Ló Further paratypes. South Austratia: Parawa ' 'L9()9' ' 30.vi.1971.

2ó 25. iii. , Additional speci:nens seen. South AusLralia: Bo'Eanic Gardens' ' Ù3' f ix' 6ó, 2.i,x.l-97o; E::ic Bonython Reserve, Iô ' 28'vii'197o; Parawa ' fd, 23.íx.Lg7o, !ú, 28-ix.,. 8d , 6'x', ú, r4'x'" 7-ô' 3'xi'r97L¡ 'rorreris Lake, ú,28.ii.L97L-

Spars i.cl-adius paucisetosis, sp. nov. Simi.l-artopgly¡e-lo;;:.sbutreadilyseparatedsiDceA'R'always-1-ess

than t, squalna with s:Lngfe seta'

MALE Í4J . Simitar to p-S.1.-v-åe!a-q-Ls- but:- Fig. I34 Sparsicladi.us paucisetosis, sp. nov., ma1e, apex of antenna. rig. I35 Spar sicladius qramin-is, sp. nov., malen terminalia.

Fig I36 Ne-rncricladius diversu,gr sp. nov., nrale, wing.

I Fig. L37 term-i-naIia-.

Fig. l-38 - tt female, termj-nalÍa. 136

,i1'l I

t t LJ t (t 134 t, 135

Ù ( a a aù J a a I a

c \, U u a

137 138 L42 g_q. Segment 3 of palp wi--h 3 rod-like sensil"l-a (I5¡r) , palpal formula o.3 : o.4: I : I :2. Antenlq. A.R. O.75 - O.85. T,ast flagel-Iomere tapered in two steps to fine point (rig. 134). Thcrax. Dorsocentrals u¡i- biserial . 'v\]ing. Length I.3 - t.5 mm., Asp. R. 4.25, squana with only one seta, costa procluced very slightly. Abdornen. Tergites with 2 rows of setae. Terminalia. Sternapodeme with extended corners, P.R- O"20t inner lobe right-angled. Para- TYPES. Hol-o'Uype o' , Hindmarsh FaIIs , South Australia , 22' ix' 1971' types. south Austral.ia: Botanic Gardens , 2ô , ll.vii., Id, 26.vííi., ld, 30.i.x. , Ió , 9.xi.I97o¡ Eric Bcnython Reserve, Id, L2'xi'I97o; Humbug Scrub, Ió, 26.viii.19'iO; Hinclmarsh Falls, Ió, 1.x.1969, 2ô, i-.vii.L97O¡

Parawa , Iô , 19.ix. L969- Additiona I specimens seen. south Australia: Botanic Gardens, #, 12.vj-j-i., 2ó, 17.ix. , Ió, 30.ix.I97O¡ Blewitt spriDgs, 3ó, 22.tx.L97L¡ Eric Bonl'tirotr (reared Reserve, ld, f .ix., 18ó , I2.xi.l-97O; Parawa, 4ó, 19.ix. 1969, óó from grass divots) 12"viii- - 8'i.x'I912'

Spar sicladius graminis, sp. nov. símilar to poryggtogi= but separated by higher 4.R", Iarger corllers on

sternapocieme, reduced number of clorsocentral bristles and p:roximal hurrp ort gonostyl.e.

MÀLE ( l-) . Si-milar to polvsetosis but:- fiagellcmere Heacl . PaIpaI sensilla rod-like. An+-:enna. A-R. 1..45. Last taperecl clistal-Iy, truncate. sgËgå. Dorsocentrals biserial arrter:iorIy, uniseriaL posteriorl¡'- I'Iing. Length 1'4 nun', Asp' R' 4'2' Squama with 5

medium setae. Abdomen. Tergites with sparse covering of setae. Terrninalia (nig. f35) . Anal point narrower than in po-ry¡e-!o¡5.¿ sternapocleme with corners extended and broadened, transverse arn ve::y broacl; P'R' O'15; inner lcbe as i¡r aucisetosis qonostyl.e r';j.'th inner sulrface peaked about one thii:C of disl-ance from l>ase, dorsal surface dr:awn i¡¡to a ridç¡e¡ s'oine terminal.. j.a 30'vi 1971 T):PES . IIoJ-orÌype d , Par:av¡a, Sout:ìr Atrstral ' ' ' r43

Genus NEMORICLADIUS, gen' nov'

Type s¡:ecies N':lmo::iciadius $ivffi, sP. nov- greenish col-ourati<'rn of the MALES. $trhen first captured, all show a slight remainder of the body' abdomen which conl-rasts rvith the light brorvn of the postorbital's Head. Verticals fine, short, reaching almost to midline; without- dorsal strong. Frontal sensillae present, brow norrnal; eyes bare, elongation;M.R.o.33"Segrnent3ofpalpv¡ithsensillainindividualpits oninnerdistalsurface.Anten¡q.Flagellomeres2and3with3large, di'stal}1' slightly tapered type C setae; last flagellomere sl-ightly clubbed withtypeCsetae,ringsetaealldnumeroustypeDsetae.rho'_el.Ante- process broa

KEY TO SPECIES OF THE GENUS NEI{ORICI'ADIUS Irfales I Inner lobe of gonocoxite broad 2 rnner l-ol:e of gonocoxite narrow (r'ig' L42)"" Iobitq¡9.-s-' sp. nov.

.> (1) fnner lobe oÍ- gonocoxite with ventral ridge, turned up distally; sternapodeme with large sp. nov. corners; anal point bare (r'ig' 137) diversus ' fnner lobe of gonocoxite with dorsal ridge, not turnecl upt sternapodeme with small corners; anal point rvith minul-e setae inverstts , sp. nov.

N emoricladius diversus' sP. nov.

separated frorn lobitoruå by the broad inner lobe of Lhe gonocoxite and as the from inversus by the larger corners of the sternapodeme, as v¡eli ventralrirlgeandturnedupposteriorborc]eroftheinner]-obe. sensill-a; ext-reme MALE ( 13). Hearl . segment 3 of palp wj-th 2 - 4 tapered : l-'5' palpal for:mulae O.25 : O.5 : I : I : I'2 to O'25 : O'5 : I z I'2 Antj.¡]'g.A.R.0.60-o.85.Lastf}a9ef].omerewith5ringsetae.aLJ.east 2tl,pecSetaelzoy).Iþgg.Antepronota]-lobesseparatedfrolrrd.orsal half way aspect. wing (r'ig- 136) ' Length L'25 - I'55 mm' *n*U finishing tip' betrveen *L*2, M3.r4, costa producecl 2i3 of distance to wing a.n'J rej-nforced Termi¡raLia (Fig. I37). Sternapodeme lvith strongly drawn out Inner lobe corners. Phallapoderne highly contoure<1 . P.R. abor-¡t o'1O' surface broad, posterior border curved upwards to varying degree, ventr:a-l just prox- with stighL ridge. Gonostl'1e with slight seta-rinqed depression imal of spine. clul¡be<1 seirsilla' FIIMALE (1O) . Heacl. Segment 3 of pal-p with 3 slightly ExÈremepalpal.formulaeo'5:o'6:l:1"35:I'Bc-rtoO'5:0"6:I: 1:L.4.4.try-rvith5fiagelicmeres;ftagellornerelslightlycon- type C seta stricteci medially witii a sunken seta on eacli liaff and a single c setae (s;r) on basal ha1f. Fiagel lomerjes I - 4 each rt'i.th 2 distal type (3o¡)a-¡ldsiwith4typeDsetae.Bxtremçlantennalformu].aer"43:I: t45

5 ring L.43 : I : 1.57 to I.67 : 1 : I : I : 2'25' Flagellomere 5 with present' vling' setae and numerous type D setae. Thorax. T\^IO pre-alars and' from 3 6 macro- Length I.oO - l-"25 mm. Macrotrichia on R' Rl ttd R4*5 - trichia near distal end of tto2' Terminalia (niq. 13S). Sternite VIII sclerotisa- with al:out 14 setae on each side, gonocoxite vfII with moderate tion.Eggguidesomewhatrectangular.Tergit.elXwithshallowmedian Tergite x wiÈhout cleft, not obscuring x; gonocoxite IX slightly produced' one seventh ridgei cerci oval from dorsal view. vaginal apoderne projecting openings of spern'- into segment vrr, basarly haìf as wide as segment vrrr; athecal ducts sclerotised' a small hump (IO¡) th::ee PUPA (3 cephalothorax. Thoracic horn reduced to ' (l'ig' Sing1e precorneal setae arranged in triangle each abouL 3Op 139) " setae (7o¡r) and frontal seta on praefrons (niq. t4o); 2 anterior pronotal II VIf each rvith 3 mesonotal seÈae (4op) ' Abdomen (nig' I4I) ' Tergites - the tt"¡o uniform covering of short poster:ior1y di-rected points, of wtrich posteriorrowsaregreatlylengthened;somepointsareforkedandoverhang setae and all- L the intersegmental memJrrane. D4 quite large, remaining D of minute ora]Iy- setae minute. Intersegmenj:al membrane wi.th several rows each with 3 directed hooks. Anar lobes forming part of hexagon in outline, 2 posteriorly stout cun'ed seÈae 2/3 Length of lobes, one anteriorJ-y, directed. TYP$."tlolotypeófromParawa,SouthAustralia,Ig.vii.1969.P.l-ratypes. (p' a' Duckhotrse); Bota¡ic south Austr:alia: vJaterfall Gul]-y, Ió, 13.xi.L962 29 2'viii'L972¡ cardens ld, I.iv.7970¡ Ló, 9.v'197I; Blewitt sBrings ' ' Bonython Reserve' Id' Ðeep creek (Ad.elaide ttills), f9, 29.v:r.l.g6g; Eric Lô' 25'v!i" 15.vj-i.L9'lo¡ Hindmarsh Fal]s , L6, I'vii 'I97o¡ ld' 3'vi' ' ló,8.i.x.1971,1?,24.v'Lg'72,29'(reareclfromlit'Ler)3'vi'1972'26' Keynes HiII' 1ó' reared from Xanthorrhoea steÍt debris), 10'vij-i'L972i Ig12' n (reared from lo.viii. L969¡ r)arawa' Id, 29'vii 'Lg6g, 29 ' August' ' Iitte-r: and ground moss respecti'vely) ' 2'viii'L972' Fig. 139 N_eqg¡3cladius- diversusr PuPa, thoracic horn and pre-corneal setae.

t: tt Fíg. LAO il head, ventral view,

Fig. J-4L ll ta dorsal vievr, anal. lobes an'1 pre-anal segments.

Fig. L42 Nemori.cladius l_obitorus, sP. nov., male, terminalia.

Fig. 1"43 $-eÉ@pþJes. invenustulus, Skuse, maÌe, tentorium.

FiS. I44 (see below)

Fis. L45 Kiefferophyes phalÌithrix, sp. nov., maler rflagel-lomere 3 of antenna.

Fig" 146 ll wing. 140 r30

a , I ,",' at lol l I I a !! !,", ttll a I tt a a .. t I ,,J aa a I ! t ,, , ,, at' ta ( I I attt I , a a t "'loa a

,,,,,,,i,,,,,i

a \ ¡ \ o a, 141 c ú I 42

l- 145 iæl

146 ItlC'

Additiorraf spe.cimens seel:Ì- Soutlr Australia! dd99 (some at light' some swarming), Apr:il - sept,. 1970 and 1971 from Botanic Gar:dens, Eric Bonythoil

Reserve, Hi.ndmarsh Falls and Parawa; Mt. crawford state Forest, ló, 6.viii.1968 (Col.-tess & Liepa) . victoria: oceran Grov('.j , Ló , 17"¡iii' L96o (J. l4artin) ; Tcorloo Arm, Lakes Entrance, !ô, Swan R.each, 1ó, 22.víí. 1964 (p. H. colless). Australian Capítal Territory: Paddy's River, Lô, 19, 22.víi.í.1968, bred from mushrooms - etnerged Io.xi.1968 (Z- Liepa) ' the AmonE the males there is considera-ble varj.ation in the outline of inner lobe, in palpal formula and in antennal ratio: it is possi'trl'c that this species is realty a compcsite group'

Nemo::icladj^us iuversusr sp. nov. Distinguished frorn rest of genus by narrow sternapodeme with redr¡ced corrr.ers, and from,civersus by the dorsal ridge of the inner Iobe.

MALE (2) " tlead.Segntent3ofpalpwith3slender,sli-ghtlycapiteJ-l-ate sensilla; palpal formula o.5 : o.25: I : l.o5 : 1.4. Antenna. A.R. 0.6' (28¡) type D setae (13¡r) and Ia-st ftageffoinere with type C setae ' numerous miclLi¡re, acros- 4 ring setae" rhqla". Arrtepronol-al lobes in contact al-oug ticbal l:ristles absent, humeral pits relativety large. L-qg.å' L'R' o'65 vüing. Length I.35 rnn. Termir:+lia. Anal poirrt with milute setae' s"Úern* phallaporlettte apodeme highly arched, narrower than .diversus' corners slight, with triangular plates. P.R. o.25. lnner lobe broad withr- dorsal- ri'ì;e surface rr:unded posteriorly. Gonostyle boat-shapecl, heavily curved, dorsal

extended. TypES. Holotype d , Eri-c Bonython Reserve, SouLh Austr¿rlia, I5r 'r"ii ' 1970 '

Pa.ratype. Hindr¡a-rsh Falls , L6 , 2l' ' viii ' 1971" swarmt 7\clditi"ona I specimens seeït. south Austi:alia: Botanic Gardens, (at 9'vii'1970 L6 2Q.v.:-:glt-, 4ð, 17.vi.1970, Ió, 24-vj-,Ig7}, 2ô, light) ', ', I1.ix. Lglot 2ó, 26.v.Lg'1L, 76,, 2-vi-rg7r, !ó, 12'vii'i'L97L' J'ð', 26.viii.f971; Deep C:;eek (Ac1ela.-'Lcle nills) ld, 9.viii .1969; Eric Bonvthon I'al'ls' J'd Reser:ve 1.c{, 2.6.v.1.9'iIi Éllunbr]g Sc::rr}: ld, 9'viii' 1969; }I-irrdrnarsh ' r47

12.viii.I97L; Parawa, dd (swarrn) 30.viii.1969, Id, 27 -v-L97o, 3d, (at light) 1.6.r¡i., 2ô , (at light) 3O.vi.I97L, numerous specimens reared frorn hillsi¿le divots JuIy - Sept. 1971. Victoria: Toorloo Arm, Lakes Entrance'

Ló, 22.vLi.L964; Swan Reach, Ió, 22.víi.I964 (D. H. Colfess).

Nemoricladius Iobitorus, sp. nov. Simil-ar to diversus but easily recognised by the 1ong, slender, twj,sted inner lobe"

MAI,E (3). Heacl . Segrment 3 of palp with 4 rod-like sensilla, palpal fornLu-l.a o.25: o.5 : I ¿ I.2: 1.6. altege. A.R. O.75. Last flagellomere with 2 type C se+-ae. ftt*9¡1. Antepronotal lobes well separated. Legs. L.R. 0.6' both sprrrs of rear til¡ia with small accessory setae. I^li_ry-. LengLh 1.5 rnm., *¿*5 firishing closer to level of tlnr, costa produced half way to w-ì-ng tip.

Termina-l-ia (Fig. I42). Sternapodeme thickened, wj.th re-inforced corners; phallapodeme with rectangular heavily contoured plates, P.R. O.20. l-nner Iobe narrow, almost free, curved upwards disÙalIy.

_ryIq. Holotype d, Waterfatl Gr-iJ-Iy, 13.vi.1-962 (D. A. Dttckhouse). Para- types. Bota¡ic Gardens, Iô, 16.iv.I97o¡ Parawa, Lô, l-2.vii -I97o-

Additionaf specirnens seen. South Australia: Botanic Garclerìs , Ló , 25.iii.,

1ó, 3I . j-ii.t971, numelous specimens (some from light, some from oi.ly paper) Àpril - Aug. L97O¡ Brownhill Creek, Lô,3O.vii., 26,2O.vii.1969; Eric Bonythorr Reserve , 2ó, f .i.v. , Ló, 2O.v. , 26, is.viii., 4ô, 2B.viii.I97i), 46, 12"v.. 5d, 26.v.L9'lo¡ Hindmarsh Falls , Ló, 3.vi. , 4ó (swarm) , J-.vií.1-97C, 'Lô, 2ó , 26"v.r97r¡ Farav¡a , Lô, 12.vii.1970, Lé, r7.vii-, 28-vii.l97l.

Victoria: Toorloo Arm, Lakes Entrance, L6, 22.víi.i964 (n. H. Col-less) '

Genus KIEFFBROPIIYHS Freefiân

Kieffe es I'l:eeman, 1961, P. 663

Type species Camptoc ladit'.s invenustulus Skuse' 1889.

¡ " Fïeacl.. postor:bj-tals strong; verticals clecumbent' usual-fy teacl'l 2/3 of distance to nidline¡ front.rl- sensilla on slíght hurnpsi eyeci Lrar-'e. Ten* tor1um as in !'ig. L43; segmcnt 3 of palp with sensilla (25Y) ou Írirrcr 148

distal surface often clubbed and bent to a right angle. Anteuna. Type C setae slightJ-y Lulbed basal-ty; flagellomeres 2 and 3 broader than long, each with 3 type C setae (I8¡:), last flagellomere slightly clubbed distally with 2 type C setae (2O¡t), a number of type D seiae (22Ð and at least 2 ring setae with small pegs. fhog. Antepronotim slightly reduced, lobes not contiguous; scutal process broado lateral setae present. Acrostichals nr-inute, uniserial; dorsocentrals uniserial; 3 pre-alars; scutellars uniserial, about 6. Legs (Fig. I48). Inner spur sinuous, with accessory setae; outer spur very short; setae of tibial comb parallel, evenly space-d, of uniform length (c.f. Austrocladius); tibia not expanded distally. Il¡tg.

Squama with single small seta or bare; macrotrichía on basal vein, Rr Rr; r¡icrotrichia invisible, punctures minute; R2*3 P-tullel to and ending close to R4n5r Rn*, finishing closer to leveL of M3*4; pfk sliqhtly dist-a-r- of r-m. Abdomen. Tergites with setae in 2 irregular transverse rows, sternite I without markings, II - V with 2 pairs of well separated sclerotised spots, the anter-i-or paì-r closer together than the posterior, VÍ - VfiI cornpletely sclerotised and with numeror.rs J.arge setae. Terminalia. Arlal point broad, hairy, fin-Iike in lateral view. Sternapodeme evenly arched; penapodeme present; goncstyle usually with heavily sclerotised distal process on dorsal surface; spine large.

FEM4LE. Hea

spermathecal- ducts basal.Iy br:oacienecl . r49

KEY TO THE SPECIES OI' KIEFFBROPHYES

1. Inner lobe of coxite large 2 Inner lobe of cc¡xite small 6 2 (L) Gonostyle with distal process . " " 3

Gonostyle without distal Process 5 3 (2) Distal process of gonostyle sclerotised, pointed 4 Distal process rounded, rrnsclerotised " lobifer Freeman 4 (3) Dista1 point of gonostyle extended perpendicular to style, mouth parts not extended " invenustuLus Skuse Distal point extended along axis of sty1e, mouth parÈs slightly extended cuspistylus' sp. nov'' 5 (2) Dorsocent-ral setae begin above humeral pits' anal lobe absent, A.R. low semithrixf sp' nov' Dorsocentrals begin close to antepronotum' anal lobe Present, A.R. high artiothrix sp. nov. 6 (1) AnaI lobe large, aedeagal lobes not pubescent Ieei .Freeman Anal tobe small , aecleagal lobes pubescen't phallithrjx, sP. rrov.

Kiefferophves invenustulus (Skuse) Freeman (b) Kief ferophves invenustuLus Freeman, 1961, P.664t Fig. 14(a) ' p.22,8. Campto cla-,lius invenustulus Skuse' 1889 , P.256; Kieffer, I9L7, point on the gono- Quick]-y recognised by the exaggerated sclerotised styIeofthema1e;anaIlobes1i9ht1ylargerthaning@x. bulbously on inner distal sur- SAl,Fi-!ql-. HSed. Segment 3 of palp expanded face wj-th one solitary and one group of slightly ctubbecl sensilla' Palpal fornrula o.3 : O.5 : l- .. L.2 z L.4. M.R. O.29, tentorium as in Fig' I43" AntennÈ" A.R. o.85 - 1.1 mrn. Type c setae very stout; last flagellomere Antepro- tapered to a ¡nint distally, 3 or 4 ring setae near tip. Thorax' notal lobes wibh 4 lateral setaei dorsocentrals begin forward of humeral- a few accessory setaer coKib pit.s. ægs. L"R. o.45 - 0.6¡ inner spur wì-th withr widely spaced teeth. VÊig (Freeman, L96L, Fig. I4a). Length 150

O.g - I.4 nun. Asp. R. 4.8. Squama with si.ngle seta; anal lobe very small; costa produced one third, of distance to wing tip; Cu, bent distinctl-y dor+n- wards, An finishing under pfk. Abdomen. Sternites III , IV and V each r,vittr narrow medial strip of sclerotisal-ion bearing several setae Terminal.ia (Freeman, 196l , Fig. 146). AnaI poi.nt U-shapecl , covered v¡ith stout curved setaei sternapodeme slightly narrower than semithrix. phallapodeme wi'th large rectanguJ-ar p]a-tes; P.R. O.05; inner lobe right-angled, setose; dorsal surface of gonostyle extended distally into broad curved sclerotised point; spine Paralle1-sided.

FEIUALE (4). Like male but:- sensilla'Palpal formula Eead. Segment 3 of palp with 2 groups of capitellate o.4 : O.5 : I z L.4 z 2. Ante.@ formula 1.43 : I : I : I : 1.5O. l0horSrx-' Dorsocentral-s begin close Èo pronotum; 3 pre-alars. I¡¡ig-. Length 1.1 nun' Asp. R. 3.4. Abdoten. No medial sclerotisation on sternites IrI - V' Terminalia (f ig. L44). Sternite VIII with 13 setae on each sj'de; gortocoxii:e IX wj.th short ciistal setae. Vaginal apodeme projecting one Lhird into segment VlI, base one third as wide as segment VÏIT' the Macleay Museum col-lection' rylrs_. ï have examined the I hoJ.otype in Additional srrecimens seen. Soutir Austral-ia: Afnut, J6, 30.vii..1969; Blewitt Springs, 19, 2.viii.Lgl2; Eric Bonyt-hon Reserve, J't1 , 28"vij-'L97O; (srçarm) 2O.viii' Hunbug Scrub , Ló , 9.viii .1969; Parawa, 2ô , 3O.vii., ld, '

2ó 28'vii'r97o; rd 20'i-" 2ó , L7-ix., 2ô , 4.xii.1969, 1ó, 19, rs.vii., , ' 1ó, 4.ii. , Iô, 1.?.ii.., (at light) , ió, 28.iii. (at light) , Aug. 19?1" and Aug. - Nov. 1972 numerous specimens hatched frorn grass d'ivots; Robertsons' 4ð, 7.xj..1969. 5iSfI9."Éy"o_ hal,Lithrix sp. nov. without Similar in appearance to ¿¡lvgl'ryLgtUleg: the only $!'ig*tropæs irr a large inner lobe an<1 the only orthocladid spec-tes I ha've observed v¡hich the aedeagal lobes are strongJ-y p'rbescent' oue tiqhLty I'q-LEJI f )-. lleÉ. Segnent f r-rf patp witli otie soli tary and forrnu'ia packecl qroL'lp cf 3 - 4 bent, stightly capitellate sensil-Ia; ,oalpal Fig. 144 Kief f erc.¡Phyes invenustulus female, terminalia.

Fig. T47 Kiefferophves phallithrix, male, terminalia.

Fig: 148 KiefferyFhYes arti,othr sp. nov., male, apex of rear tibia.

tl Fig. L49 ll il terrninalia

Fig" I50 Kieff es semithrix sp. nov., male, wing.

rig. 15I lr il terminalia,

Elig. 152 StejEeroplfes- cqEPl-styh., sP. nov' male, gonostyle' û t.. t_ o

148,' _¡r_J ta aa I a I ( g

Itv

-+ri /I í44

149

a a 147 a ¡ ¡--- - 151 152

150 15.1

O.3 : O.8 : I : 1.3 z I-7, M.R' O'3' An!e44q' A'R' O'75 - l-'O' Type C setae heavity bulbecl (nig" 145). Thorax. AnÈepronotal l-obes with 3 - 4 small lateral setae; dorsocentrals begin close to pronotrxl. L.egs' L.R. O.5 - 0.6. Vling (rig. 146). Length 1.I - 1.45 mm., .Ërsp. R. 4.85. Anal lobe stight; macrotrichia on R4*5t costa produced half way to vring tip; cu,. bent sl.ightly downrvardsr An finishing s]ightly distal- of pfk. Terrainalia (f i-g. :-47) . Anat point U-shapecl , covered with short curved setae; sternapodeme with lateral arms broader, corners reinforced; phallapodeme with flat triangular plates; aedeagal lobes strongly pubescr:u.l-' P"R- o.3; inner Lobe reduced to stight swelling along inside of co:'r-lLe; gonostyle boat-shaped, distal-ly with small- sclerotised satv-tooth; spine

slencler , tooth-like.

FjIMALE (3} gegg. Segment 3 of palp with 2 groups of capitellate sensi}]-a; palpal formula o"3 : O.5 : I : 1.3 z L.7. Antennal formula I.43 : 1 : I : 1 : L.2. giry-. Length 1.25 mm'' AsP' R . 3.4. Terminalia. SterniLe VIII with 9 setae on each side; gonocoxite IX closely adpressed to tergj-te x'

vaginal apodeme projecting one quarter into segment vII, basally one'cTrird as wide as segment VÏII

south Australia, !97O. Para- ry3Eg. Holotlzpe d, Eric Bonython Reserve, types. South Australia: Aldgate CreeJ<, fd, 3.vi.L969¡ Botanic Gardens fd, 26.r,iii..I9'tC¡ Eric Bonython Reserve, ld, 28.vii.L97O¡ Hi-ndmarsh I'al-1s' Parawa, 2ô, Ì9'vij-' 29 , ld (swarms) , f .ix. L97i¡ Morialta , Ló , 29-v:-.í.Ig6g; ' # , 3o.vii., Lô, 2o.vii.7.969, 1?, Aug. 1971' Addi LionaI specintens seelt. South Australia: Alnut, 9ó , 30'vii'L969; Brownhil-'l Creek, 2ó, 4.tx.L969¡ Bot-.anic Gardenr-;, d, (at liqht) ZZ'ix'i97J-' ld, (at l.ight), 3.x.i971; Deep Creek (adelaide HiIIs) , Lð, 3.ix.L969¡ Hindrnarsh FaIls, Il-tshersr swal"rns , ]-7.xi.1969; Heafeys, Bó, 18.x.196'9;

swarms , 20. vj-i. 1971 . Represeirt:atives c¡f this spec;-es vary sì.ightly in 4.R., palpal formufa r52 and nunlcer of setae depending on the locality; however, the number of specimens available from eactr site is small and further sampling coulcl reveal the existence of more thatl one species'

Kieff es ar:tiothrix sp. nov. SimilarinappearancetolLY.enustq]u€.butgonostyiewithout exaggerated tooth. to midfine; segrment MALE (6). Eeag. Verticals ending in small group close 2 of palp with group of 6 - 7 bent, capiteÌlate sensill-a; palpal formr:la O.4 : O.4: I : 1.3 : 1.6. M'R' o'30' A!e¡1lÈ' A'R' O'75 - 1"oO; fast flageltomere truncated distally. Thorax. Antepronotal l-obes rru-ith olie Iateral seta, dorsocentrals beginning at border vrith pronotum' LcgF-' L'R' o.5-o.T,reartibiaasínFig.148.wl'g.LengthL.2-1.5nm.,Asp.R. 4-S.Anallobeobtuse;macrotr:ichiaotR4+5rcostaproducedhaifvrayto wing tip; Cur slightly bent, An fi-nishing under pfk' n¡99' sternite V with medial grouP of 2 - 3 setae Terminalia (rig. f49) - AnaJ- point brcao u-shapecl, covered v¡ith shorL curved setaei sterna-podeme of even t|ickness; j,nner phaltapodeme v¡i.th large pJ-ate; P,R. O.O4¡ fobe forming right-'*angled corner with a few short se'i:ae; gonostyle boat-shaped with slight sclerotised distal tooth' 1 : 1'5 z 2'2' Antennal FETI4ALE (r). Head. Palpal fortnula O.3 z Q"l : fc¡rmulaI.43:I:1:1:L'79'VJing'LengLhl'1mm''As-p'R'4'3' Termina-Iia. stern:"ie vÏfI with 14 setae on each sicle; gonocoxite IX hatf way into larger than in remainde-r of genus; vaginal apodeme p::ojecting

segment VII, basaLly one quarter as 'øj'de as segment VIIÏ' South Australia' TYLìES-. Holotype d and 1d paraLtype taken at light' Parawa' 1ó' 20.x.Lg'll. Further paratyPes" south Ausl-ralia: Blevritt springs ' (swarm),19,2.viii-L9?2;EricBonythonReserve'-l'd'1I':i'l-9'7o'1ó' l¡af Is, 1d, 2'3.tx.L97o¡ Humbtl.g Scrub , ?Å , 9.viii. ]-972; Il:Lnc]marskr 25.vii.l97I.

25'vii'i' 2ô , Äd.c1it jona.l sÞecimens seen. Scut-l-r Austral.'i-¿ì.: Para\^¡a, -td, ,

20.x. , 26 , l¡.x.1.97I (all at light) ' r53

Kief ferophves semithrix sp. nov.

Very similar to K artiothrix separated by lower 4.R., shorter dorsocentral row and l-ack of arr anal lobe. MALE--j$-. H-edg. Segment 3 of palp with 3 - 4 se¡rsil-lae' M'R' o'35' Antenna. A.R. O.40 - O.55. þ-1"". Dcrsocentr.al-s begin at level of humeral pits. !ùing. Lenqth L.25'- L.3 mm., Asp. R. 5'I; anal lobe absent (nig. r50). Terminalia (ni_g. I5I). Anal point more pointed than in artiothrix, plate of phallapodeme small-er' TypES. HoJ..otype d and 2d paratypes from Parawa, South Austr;r1 iir' 30.vii.1969. Further parat-ypes. South Austra.lia: Eric Borrythoii Reselve, Ió, 20.v.Lg7O, 1ó, 15.vii.L97O¡ Tanuncla creek, 1ó, 1O.v-i-ii.1969. Aclditj-onaf specimens seen. south Australia: Bl-ewitt springs , llô, 2.viii.Ig72; Eric Bonython Reserve, #, 3'viii'1971-'

Kie fferophves clrsp istvlus, sp. nov.

End of gonostyle extended into large sclerotise<1 thorn, separated from invenustufus since t-horn produced al-ong axis of gonostyle' MA],E(3).Hejg.Segment3ofpalpwith3-4capitella.t-esensj-]-Ia;paIpai. formula o.3 : o.6 : 1 : 1 : 1.1-. M"R. o.38" Anler¡ne. A.R" 1.O; last flagellomere tapered to long thin point; ring-shaped setae laige' TÞc'feï-' Antepr:onotal lobes with I - 2 lal-eraL setae; dorsocentrals begin close to

O O.55. Length l'25 nun' Asp' R' 4'9' prorlotum. -LegE-. L.R. -45 - I:,ry-' ' Similar to semithrix but costa produced more' Terrninalia. Sl-r:ne of arral

point midwav between artiothrix and semithr-'i.x sternapccleme rrar-ror'¡er t-han in these; pttallapodeme vrith hea¡,,iJ-y contourecl plate; P.R. O.1; ir¡ner I'obe large with. round-ed point setose; gonostyìe wit--h dorsal surface drawn out distally into sclerotised tho::n (Fis. L52) ¡ spine tapered to a narrow base'

TYPES Holotype d-' uindmarsh Fa1ls , 4.v!i-'Lg7L; pa'ratyPe d, Hindma:rsh FaIIs, 28.vi¡l"1970, South Australi a. Àdditional imen seen. South Austral-ia: Para-wa , Ió , 2l'vii "797L' 154

Kiefferophyes lgÞifer, Freeman Kiefferophyes ]gUt¡"t Freeman, 1961, p- 664; Figs' I4c' d'

MALE 2 . Hgad.. Verticals reach only one quarter distance to micline.

Segrnrent 3 of palp with 3 - 4 rod-like sensilla, palpal fc¡rmu1a O'2 : O'5 : I : ].5 : I.5. Antenna. A.R. i.2- Thorax. Antepronotal fobes with one lateral- seta. beginning at border rvith pronotum. Legs. L.R. ""*""..*rals O.45. Wing. Length I.3 mm., Asp. R. 4.3. Squama with one seta, anal Iok:e extremely srn*all-. Macrot.richia ot R4*5, costa produced nearly to wing tip; Cu, bent sharply; An finishing d,istal of pfk. f&9gme".. Terqites ivith a¡tterior and posterior row of setae. Tgrm:Lnglia (Freemari, 1961 , F-ì-g;' 14c, d). AnaI point U-shaped, covered with stout setae. Sternapoderne narro\{, evenly arched; phallapode¡ne with t,riang:ular plates". P-R- o.08' fnner lobe triangular, slightly hooked; gonostyle vrith 1arge, flattened.- unsclerotised distal fobe. Spine subterminal' TYPES. I have examined the paratype ser.ies, 8d, Bogong Plains, Victorj'a, Jan. L92e, F.E. I'filson-

Additional s ecímens seen. South Austral-ia: Botanic Gardens, 7ó, 2.x.\i:69, dd, 2.íx. - 2I .x.L9'lo (some at tight and. on sticky paper); ilrnd:narsh t--alfs, Id, 1.x.I969; Parawa 3d, 22-Lx-, Ló, I2,-v-L97o¡ 6ð, I2'v', Id, 3o'vi', 1ó, l.3.viii. (at líght) , 2ó, 8.ix. , ró, 14-x-1971.

Genus PARAMETRIOCNEIVÍUS Goetghebuer parametriocnenìus Goetghebuer, L932, p. 22. Freeman, 1961, p. 659.

Type spe cies Metriocnemus st¡]-atus Kieffer, 1.924. Parametriocnenius ornaticorrlis {t

Paramet-::iocnernus ornatic<¡rnis Freeman, 1961, p. 660, Fig' 13c'

Metriocnemirs ornaticornis Kieffer, 191''7, p' 225"

Freeman says "Vel:y sit¡.il-ar: t.o Pafea:rct.i-.c species stylatr-rs Tiieffer, buI

small-er and" with.shorter leg ratio." The pupe. exhjhj-ts sexli¡;tl- clirnorphism. ¡.'AI.E (4)-. E.eeg. Frontal sensilla abscnt..; eyes wi.bh dorsa]. el<-lrrqiatj'on 5 facets wicle; rY"R. 'J"2/+- liegrLrerlt 3 of palp with 2 gÏoups of' 2 ov i) cl":l';bccl , 15s pointed sensilla arising from small pits on ventral surface, palpal formula o.3 : O.3 : Ì : 1.4 : 2ì palps longer than half length of antenna. Antennq. A.R. o.9 - I.2; flagellorneres 2 and 3 with 2 type c setae (25p) and segment 5 with single curved one; las'b flagellomere- heavily cl-ubbed distalJ-y with whorl of 3 or more type C setae lZCO¡r), curved right over tjp, numerous type D setae (soru) and at least 4 small ring setae (5¡r) ' Thora{'

Antepronotum with 2 or 3 lateral setae. Acrostichals biserial-; dorsocentrals with conspicuous punctures, beginning muJ-t'iserial anterio:: of pits, 'L'R' biserial at reari 5 pre-alars; B scutellars, uniserial.. Legs' O'7' Inner spur with short basal accessories, comb with few setae' WiL4f (nig' r53). Length 1.6 - 2.O5 mm., Asp. R. 4.OI, squama with up lo L2 fi:-re setae' Macrotrichia biserial on R, Rl_ utd R4n5, uniserj-al on remaíning veins and' scattered over: whole winq membrane excepL for ceLl R.. cc¡sta prod'uced one -,hird of clistance to viing tip, Cu bent slightly towards trailing sdge¡ An reaching nearly to costa. Abdomen. Tergites uniformly setose' ster:nites wi-th l¿r-.eral ancl medial seÈae increasing on consecutive segmenl-s, Vf an'å V-ì-i- uniformly setose. Terminalia (Freeman l-961-, Fig' l3c) ' Anat pojnt ur:piE- mented, bare, degree of point variable. sternapodeme only slightly srclero"' tised transversely, corners with small poi.nts' Penapodeme absent' phalla¡;o- rounded' deme with plates reduced to narrow scleroti.sed bar. Inner lobe

gonos l-yle clrawn into ridge distally '

FEMALE ( 3) . Lilce male but:- Head. verticals cl-ustered aL rnidline, segnrent 3 of palp witJ. one sJ-ightly

cl-r¡bbed sensj.ll-um (I4ir), pa.ipal formula o.4 : O.4 : I : I"5 z 2.4' Antenll with 5 flagellomeïesìi fst constricted rnedially, with I basaL type c seta

(f5,¡r) aud. 2 terminal (3O¡r) , 2'4 increasingJ-y flask-shaped, each with 2

type C seta,e (3Op), 5th wi1:h 3 long curved set-aei flaEellomere for:nula

L-54:l:1.1 : l.l:l..I.Termina1i-a(nlq.154)'St-erniteVITIwithl'4 IX setae on each sic1e, egg 9ui<1e wjth conc¿I'¿"¿ msdÍc1l rnargin' Tergite \'¡ith nur'ericus partl1, obscurirtg X, rL'recl-ia]. cl-eft cleep; qcnccoxite IX r:ounded''

stz.ong setae. fiterni*.e x with,¡:,rt ri-c1ge. c..:rci hernisl:herical- frorn dorsal Fig. 153 Parametr iocnemus ornaticornis Goetghebuer, male' wing.

Fig. L54 ll female, terminalia

Fig.155 lr pupar thoracic horn and precorneal setae.

Fig. 156 ll female pupa' anal lobes and pre-anal segments.

Fig. L57 tl rl l,arVa.

rig. 158 Glzmnome triocnemus wilsoni- Freema¡, female, terminalia. C VIII - gonocoxite VIII crx gonocoxite IX GC genital comb SP spermatheca SIX sternite IX SX sternite X V. AP vaginal apodeme 153

l.r- i a. .: ûth {i

o o l't v ó tJ ,rr u r 157 o!e u u uu v 9d

Itt 154

155

,'.;.; i': 3p_' !j-Y aP t: :f'l rit: c vilt \ tr tu J

tx ise c v

158 r56 vie\Á/. Vaginaj- apodeme not projecting into segTment VIf , base one ninth as wide as segrnenÈ VIII; spermathecal ducts not broaclened. basally.

PUPA (5). MaIe. Light green in colour. Cephalothorax (nig. 155).

'Ihoracic horn ( 2OO¡r) slender , spinulose , slighti-y cl-ubbed distally; 3 precorneal setae, mid.dle orre half lengÈh of horn; 2 anterior antepronotals' 2 anterior and 2 posterior scutals. Praefrons with a sl-ender wart-Iike seta half length of horn on each side. AbComen- (fig. l-56). Tergites and sternites II - VfI with I - 2 rows of anally directed stout scierotised spinules on postericr border; setae - and Ð, - D, Present. Ana.l- LoJ:e "t "5 broad, Iateral edges turned up dorsalJ-y with 3 strong cur:ved setae as long as anal lobe. Female. Like male pupa but sternit"e VII with deep medial indent-a-' tion, cuticle drawrr up on each side into a posteriorly directed peak, spinules absent.

I,ARVA (2). (Fig. 157a - f). Usually more than 5 m¡n. long; green-yellott' Head- Antennal forrLrula J- : O.5 : O.1 : O.35 z Q.2; mandibfe with interrra-f seta deeply sectioned; 5 teeth; sel-ae of abdom-inal. and thoracj-c segments minute; anal papillae not visible; pre-anal setae as sho\'ün in figure.

Typgq. Il,cJ-otype unavai-lable. I have examined the serj.es frcm whíctr

Freeman (1961-) made his description. Addition¿rl speci:nens seen. South Australia: Botanic Garderrs, Bric Bcnython

Reserve¡ Hincìm"rrsli Fal-l-s, Parah'a, dd99 (from afl of trar.ps) , througìrotr.'U ye¿ì"r,'

Broivnhill C:reek, l-é, 14.viii.1969; Healey's Ió, 18.x"1969; Morialta, 2ó, lI.vi. L969¡ e, Id, 14.v-iii.f969¡ Robertsons, 1d, 26.xj-.1969; Second

valley, Ló , 26.xi.1969.

Genus GYIIINOD'TETRIOCIJE¡IUS Gcetghebuer

Gynurornetri-o cnernus Goegthebuer, L932, p" 23. Bi:unclin, 1956, p. l-4O. Freeman, 1961, P- 660.

Type sepcies Metr j-ocnemus s:ubrruclus lliwards , L929 -

Gv:nnoLnetri-r>cr:emus rviJ-s;oni Freeman L57

Gvmnometriocnemus wilsoni Freeman, 196I, P.661. A yellowish species; wings completely covered. with macrotrichia; separatedfromsouthAmericanlongistylusbysingleinnerlobe. extended, squared'; YlÀLE (4). Head. Frontal sensilla present; brow s1i9ht.1y distal segment 3 of palp with I or 2 flattened sensilla on expanded inner surface, palpal formula O.2 : O'3 : 1 z L'2: 1'6' M'R' O'3' Antenna' (I8¡); A.R. o.9 - 1.I. Flagellomeres 2 and 3 each with 3 type C setae last flagellomere slightly clubbed and pointecl, distally with numerous type Dsetae(55¡r),about4typeCsetae(35p),3ringsetae'tttotgl'Antepro- notal lobes with 4 lateral setae. About 6 acrostichals; dorsoce¡rtrals biserial, beginning along border with pronotumi 6 - 7 pre-alar setae' 6 of rear tibia with small scutell-ars. _Lrg.s. L.R. 0.60 - 0.65. Both spurs accessory setae, comb strong. wing (Freeman, 1961, Fig. I2e) ' Length 1.5-I.Bmm.,Asp.R.5.o.AnallobeabsentiSquamabare.AIlcellsa'nd veinswithmacrotrichiarmicrotrichiainvisible,punctu::esminute.*2-r.3 joining costa closer to Rr; nn*5 fitish-ing before M3*4t costa produce'J

3/4 of- distance to vring tip; pfk c1iätal of r-m, cu, bent d'¿v¿nwards, medial setae Abdr:men. Tergites uniformly setose, sternites wiilr laÈeral and increasing on consecutive segments, sternites VII and VTII uniformly setose' Terminalia (Freeman, Lg6i-, Fi9' 13d) ' Anal point absent' Ster:napodeme curvedi wiÈh strong drawn out corners; plates of pha-]-lapodeme triangular' Gono- penapodeme altsent. I¡ner lobe rounded, covered with inicrotr-i't:hia' style enlargcd distallY' groups of 3 serrsilla distally; ¡'EI\ÌALE ( 2.) . Head. Segment 3 of palp v¡ith 2

palpa-l formula O-2 : O.3 : I : I'3 : 1'6' ¿^ntenng- rvith 5 flagellomeres; flagellomereformulaI.3T:1.o:I.i:o.g..L.2.FlagellornereIrvith 2 type c setae singJ-e type c seta (rB¡) half way a1on9, 1 -' 4 each with (32¡tt insertecl clcser to distal end; 2 - 4 flask-shaped. Itlg' l'ength wit'll abouL 1t3 I.3 mm., AsP. R. 3.7 . Termi¡ralia (nig. 158) . Stcrnite VIII guide evenly setae on each sicle, gonocoxite vrrr of moclerate thickrress, egg t5rJ rou-nded. Tergite IX with shalfow medial cleft; gonocoxite IX rounded' sternite X smal-I, cerci hemispherical from d.orsal víew. Vaginal apodeme projecting one eighth into segment vII, base one fifth as wide as segment VTII; spermathecal ducts rrot basally thickenecl' TypES. T have examined, the holotype d, Hamilton, Victor:i.a, 29'v3-'Lg3Al F. E. Vfilson.

Reserr¡e, I{j-ndmarsh Falls, Paravra, óó99, (some at l.ight; in emergence traps) r

a1I rnonths; Alnut , d , 29 , 3O...¡j i .Lg6g; I-Ioaley's 14d, 59, 18.x. J-969;

Hin&narsh FaIIs , Lð , 19, (reared from debris at hillside seèPrige) , 1.viii.Lg71¡ Nietske's, d, 9.viii.1969i Parawa 6.j9?, (reared from grass divots) Ju}y - lÑoverrr]cer; Q, ód99, 30.vii.1969. Austra]-ian Capj-t.,rl Territory: Black l'{ountain, 1d (at light) , J.x.196O, (1"jl .B. Common) '

Members of this species are generally lighter in colour than suggesl-ecj byFreemanandpossessastraight-ternLinalantennalseLa.

Gr¡mnometr:iocllemus impectir:atus ' sP. ilov

Read:i-l-y recognised by its stripecl abd.ornen, light cofour and restríct'ec' macrotrj-chiation of the wing memblîane' Diffe rs from pectinatus l¡reeman since inner lobe slender, pointed; spine sitnple' present; segtnent 3 of pal'o with group of MLLL 13_)_. I]ead. I'rontal sensil..l-a 3 diverging :lod-l.ike sensilla (1O¡); palpal íorntula o.2 : O"5 : I : O'8 : 2 a.nd 3 with 2 tvpe c I.1; !1"R" o.32. 4lr_tenna" A.R. O.85. l-Iagellomeres setae (r.2¡); l.ast, flage-ì-lornere slightly tapered and evenly rounde<1 dj-sLa'fil¡ C setae (22¡t) ; with nur,rerous r:ype D set;rc $2¡t) as well' as a rrllrxller of type singfe -iaterai at l-east-- one rj-nq seta. I]rqr*. AnteproDotal lobes vrith pre-al-ar se'tae; seta. Do.rsoÇentral-s begin abcve humeral pits, tmiseria,l; 3 6scutellars.r,e-s.g-.L.R"o.T.Spur.sofbothreartibiawithrrumerous Squan'a bare' accessory setrae. i*liX¡-. r-'enqth i'6 - f'7 mn''' Asp' R' 4"3' pfk ¿rnd' R' anal lobc present. Macro'Lrichia present oir membrane c1-ista1 of on Rz*.3 R. arrd, baserl vein' l'licrctricttia i'nvisíblc, puncttrres n-inute' R-,I' 4f5- Fig. 159a Gymnometriocnemus parapectinatus , Sp. nov., male, inner lobe of gonocoxite.

Fig' 159b Gymnometriocnemus impectinatus, male, termi.nalia

Fig. 160 Rtrinocladius tonnoiri, Freeman, male, mouthparts.

tl Fis. 161 I I wing.

Fig. 162 il terminafia

Fig.163 I female terminalia, egg guides and spermathecae.

Fig. L64 Smittia aterrima Meigen, ma1e, flagellomere 2 showíng tYPe C setae of antenna. 160

J 162 c bh o a J 159 å a

ll 163 164

I , I -t/ ) ,)/

<;-<-<-__ 161 r59 parallel with and cfose to Rr, costa pro

Ftr¡/LALE (I) . Like male but.:- re""d. Palpal formula O.2 : 1.5 : I : O.9 : 1.3. An!çqna with 5 flagelJ-o"' meres, I slightly constricted rnedially, a sunken seta and a type C seta on basal half , ftr-::Èher sunke¡r seta and 2 type C setae on distal half; 2 - 4 urnshapedr each vrith 2 type C setae; 5 with 4 type C setae and numerous ty-pe D setae; flagellomer:e fo::mula 1.53 : I :I : l- : 1-53- Thorax" Dorso- cen+-rals irregularly biser:ial . !{ing" Length 1"6 min., Asp. R. 3.5. Macro- tr-i-chia present along aII veins distal of r-m, on membrane Cistal- of pfk and right along trailing edge. Telryinalia. Sternite VIII with l-8 setae cn each side; gonocoxite VIII nìoderatefy broad; egg guicle som ev¿hat rectangr.r* 1ar. Tergite IX partially obscuring X, medial cl-eft deep; gotrocoxi.te IX slightJ-y extcnded. Sternite X with rj-dge, cerci tr:'-angular from dorsal vierv. Vaginal apodeme projec.Ej-ng one quarter into segrnent \/II, basalJ-12 one quarte.r a-s wi<1e ;ìs segment VIII; spermat-hecal ducts slightì.y broadener.l basal1y.

IypEÞ_. Holotype d a¡rd ld parar:ype Botanic Gardens, 2.ví.L97O. Further para.types : Botanj-c Gardens l-d , L9 , 15, v. 1970; Hindmarsh Falls , ld , July

L970- A

Reserve, I-lj.¡d¡¡arsh I¡allsf Para\4ra, ddg9 (so¡rre s\{arrns' some a-t light) , micl A¡;riI - end June I97o , L97I; Hindmarsh Fall,s , 26 , 5.vii "L97L (rearei fr"orn nroj.st lea.f

Colless a Z. L,iepa) " 160

GVrììnome tric¡cnetnu sp arapectinatusr sP. nov. winc; Separated from impectinatus by higher A'R" highly transparent membrane and more differentiated terminalia'

MAIJI Q) . Like inpectj.na tus but-: - palpal forrnula Head. segment 3 of palp with 5 diverging rod-Iike setae; 2 and 3 with O.3 : 0.6 : I : O.8 z L'2' Anqennq.' A'R' L'2' Flagellomeres distally, with I 2 type C set.ae (3o¡r); last f1-agellomere slightly tapered (28¡r) 3 ring setae' or 2 type c setae Q2¡t) , nulnerous type D setae and

I.çgF-.L.R.O.BO'Wing'Lengthl'8-I'9mm"'macrotrichia'preser:t-on heaviiy downwarda. membrane only in proximity of wirrg tip; Cua bent Terminalia (Fig- I59b). AnaI point of even width' setose' Sternapoden'e broacler than in impectinat us,cornersdrawnou-trrounCedibaseofphalla¡:o- io a hc¡ok, rrentrally wi'th deme afso b::oader. -lnner robe tapering dorsarly plane' a smalf hairy humP. Gonostyle br<>ad' in the dorsoventral Aust:ra1i¿r gPE. Holotype d arrd ld paratype' Brownhj-fl Creek' South

14. viii. 1969 . l'lorialta' 16 4'ix'1969; Additional specimens seen. South Australia: '

Second ValleY, 2#, 30'vii'1969' nar:apecti¡r¿lt--us have Ì-¡een forulcl in The two llev\¡ sPecres amÍ) ectinatus and L.--.-+- separ'atebutoverlappingregionsofsouthAustra.lia.trronTableVlllit mcrphs' llowever' coutd be suggested that they are actually trvo seasonal <;c1''¡-ecting \!¡rs ex+-ensive' the recorrls of ErgP9.9-tiLqt-qs are few although iack a coirrpound spine Both these species are very cl'ose to Pgtiæill-' but

Genus RHII']OCLADIUS Edrvard s Rhinocladius Erfwards, f931', p' 268' j-us Edwar cjs 193I Type sPecres Rhinoc -l,acJ cuii.cinus '

Rhinoc l- ad:ltr's tonnoiri Ë'reeman

RhinocJ aclius tonnoir-j- Freeman, 196f , P ' 662 ' to the typ'e I h¿rve re,cescribed this species after referring "iF('':ci'ne¡ls andaddj'.t-iorr.elspeciltenslistecl.Sincel'lravenoLexa¡rri.nedt'heSouth 161.

America_n species of this genus I have not revised Freeman's generic descrip-

tion but many of the additions below ma)' actually be gener:ic characters'

t"tAL,E_!?)_. H.gg. vertical-s large, decumbent, extend to midline, post orbita}s long, str:otig; eyes wiÈh d.orsal elongat-ion 3 faceLs wide; cibaria-l

pump with shalLow rectangular indentation along dorsal border; frontal- sensilla slightly raised, brow produced medially. Maxilfiary palps with 3 (3¡l) segments; distal surface of segment 3 with numerous digitate sensilfa ' palpal formula o.2 : 0.65 : l. Labiat palp 2.55 times head width (sl-ightly Ionger than anterior: femur) with swollen tip carrying usual gustator:y setae sir¿r'ft' and, an acld|tj-onal dor:sal row afong each shaft; inner surface of wilh row of short distally curved setae (these may function in zipper-like fashion to keep tire two halves together) (fig. l-60) . Anter:4a-' A'R' 1'I; Í"lagellomeres 2, 3 and 4 with 3 slender type c setae (r8¡r); last flagelfo-'

mere with about 7 similar type C setae, numerous type D setae and 4 ring setae without circlets; pedicel with ventral row of 3 small inde'ntations' Thorax. An+-eprono+-al lobes no::mally developed, with 4 stout lateral setae' Acrostichals u¡riser-'ial , snial-l and decu¡rl:ent; clorsocentral.s uniseri;"LJ', beginning in front of humeral. pi.ts; 4 pre-al-ars; 6 scutefl-ars, irriserial'

post not.um l-ong. T,egq. L.R. O.75 - O.8C. Iìear: tibia with outer spur haì-f length of inner, both spurs with accessory setae' collb carried on slight R. 4.55. Squama with 5 shorc flare. Ei"g. (Fig. 161) . Length I.9 mm., Asp. sretae, arral tobe small. îlicr:otrichia invisihle, pr-rnctures rninute; nlacro- trichia on R, R, ancl R4*5.^d on distal- th-ird o-i rrrernloranei r-m and i¡ase of sc each with 2 sensilla; R2*3 .tding closel- to Rt th¿r¡r to R4r-5t R4*5 of distance er:cling rni-dway betrveen *I*2 -td M3*4, costa produced two thirds to wir'g ti¡>; pfk well distal of r-in; cu, bent- tnioway along,' An ending a't ivii:ir l'o'teral level of pfk. 4Þgç.g!. Tergit-es uniformly setose. sterüites and VIIj- and medial set¿re increasing on consecuti.r'e seg$ents, sternites vII ,.rniformly setosei ster:nites f , f I, IV an<] V with 2 ¡-';lt'ches of sclei;oLi'seci cnticle, sternite l-IÏ wíth thÏee. le-r;g3rerjlg (I'iq. L62) . AnaI point arrched, coÏners slightly -rouncletcl , b::c-rad, stìtose. steruapodeme highry r62 thickened; phaltapodeme witl¡- narÏow curved plates; P.R. o'57; inner l-obe with broad triangular base, setose; gonostyle becoming rnuch larger distally, sPine verY small'

FEl"lALE Q) . Like male but:-

Ead.Segment3ofpalpwithmoresensorysetaetlranmale¡palpal formufa o.3 z O.77 : l. Antenna with- 6 flaqellomeres; flagellome::e 1 2 distal gentiy oval , 2-5 flask-shaped,. 1- 5 eachwith 2 basal and 5 type C tapered type C setae Qalt), 6 tapered and roundeC distally with c'9 : l- : I"-L : I'I4 setae and. numerous type D setae; flagellornere forr'tlrla 1.14:2.oo.ü]inglengthl.75mm.,Asp.R.4.2¡macrotriclriaína.].]cells 1 8 buÈ medial and R. Terminalia (FiS. 163). Sternite \zÏIf with - Tergite setae on each side of midlinet gonocoxj't'e VIII b::oader medially' IX with shallow medial cteft not obscuring X; gonocoxite IX b'ul"becl ' view' sternite X with moderate peak; cerci hemispherical from dorsal base one fifth v¡idth vaginal apodeme projecting two fifths into seginent vfI, equal- in length tr-'' of segment VIII. spermathecal dtrcts heaviJ-y sclerotised, spermathecae, ba.sally broadenecl' l'It. we1--linEton¡ TypES. r have examined tk¡.e hol-otype ó and l-d paratype from Belgrave' Victorj-a' Tasman,ia, Nov. Ig22 (4. L. lonuoi::), ancl the d paratype,

Sept. L928, (r. B- wilson) '

Aecies' specimens clescr:ibed here shouÌd be ccnsj-dered as a s;eparate 163

Genus SMITTIA Holmgren Smittia Holmgren, 1869, p. 47. Brundin, 1956' p' L46'

Camptocladius Skuse, 1889, p. 26L (in part) '

L929, p' 3CrO' S a¡riotoma subgenus Smittia Group B Edwards, Type species smittia breviPennis ÉIoImgren' materia-l, Freeman t96I described 3 species of this genus from Australian the two of which are represented in south Australia. After examination of lectotyPes I have made the following redescriptions from south Australian specimens. Smittig aterrima (t"teigen)

Chironomus aterrimus Meigen, 1818' p' 47 ' p' 24¡ c adius macl eavi Skuse, Ig89, p. 266. Kieffer, 1906, L9I7, p. 228 (syn. nov.) aqe Freeman, !954a, p' l-75¡ p' 348 (syn' nov'); 1961 Smittia ,1956, p. 655, Fig. l5a c b. Smittia aterrj-ma Brundin , 1956r PP' 146-7 '

Freeman says ,,Eyes finely pubescer¡t, thorax black, costa slíghtly produced, anal lobe of wing present but not prominent, anal po-int of male bare, gonostyle with prominent inner flange.'' MALE(7).Head.Verticalsmediumsize,extendinghalfwaytonid.]-ine. riyes witf. Brow extended into small medial point, frontal sensilla absent- shortdensepubescence.Segment3ofpalprvithagroupofrod-].ikeseir* A'R" silla (131¡). Palpa1 formula O'25 ¡ O'5 : 1 : O'9 : l-'4' Ð@'1' 2.O. Flagellomeres 2 and 3 each with 2 type c setae with elongate bases covering 4 normal sized cuticle perforations (fi9' 164).Last flagellomere type I) slightly clubbed distatJ-y with 2 type c setae (24Ð and numerous qhora{' setae, one of which is distinctty terminal; 3 type C setae' setae, Antepronota]- lobes contiguorrs. Scutal p]:ocess smafl, 3 latera]' Acrosticirals al:se¡t, dorsocentral,s begin at- level- of hume::al pits' 4 pre- alars, scutellar.s u¡j-serial-' Legq' L'R' O'5' Comb of re"'-r tibia densre' Fig. 164 _Ðlttia aterrima (¡,teigen), male, flagellomere 3 of antenna.

Fí9. 165 Parakiefferiell-a varigjrtusr sp. nov., male, tentorium.

Fig. 166 I I terminalia

Fig. 167 lt female, wing.

Fig. 168 ll I terminalia.

Fig. 169 pupa, prothoracic horn.

Fig. 17o ll ll pupar dorsal surface right posterior corner showing small pedes spurii B.

Fí9. 171 il pupar anal lobes- I \

165 169 \

168 170

167

¿#à ,É'1"1 J

166 l-¡ 171 L64 both spurs with basal accessories. vJiry.. Length 2"o mm', Asp. R. 4.o. Anal fobe dj-stinct. Macrotrichia on basal vein R, Rl. *2*3 utding closcr to R4*5, R4+5 finishing at levelof M3o4, costa produced' half way to wing tip. cua distinctly h¡ent. a¡4gT9". Tergites with

normal. qyPES" I have exaruined the l-ecto-'ype ó from the Mcleay l'luseurn, $r¡,';'r:1" 9ð i'7'vj-" Additio nal specirlrens seen. South Australia: Botanic Garclens , , ' Id, 2.íx"L97o¡ Duck Ponds, 2ó,9.v-iii.1969; I]inclrnarsh Falls, Ió,

j- j.. 30. .L97O; Robertsotrs , 2g. vii .Lg7o; Moculta, Id , 9 . vi. 1959 ; Parawa , Iô , vi 1d,26.xi-i.1969;Torr:ensLake,dóuay-Sept'L97o'Id'I2'viii'I97L'

Smittia retracta Freeinan S*j+tÞ retractjl Freeman, 196i, p' 667, Fi-.g' l5d c e'

Freeman Says "Black speciesr eyes bare, tibia-l hairs 1ong, costa' irr' anal rnale short and not produced, posterior fork short and relatj-vely vricle'

poi't rong ancl narro\^r. Atthou.gh shovring some resembl¿ince t<¡ -Ç:ry!pSra€t¿9.' bristle the a.bsence of putvilli and the presellce of the apical antennal place it here"" along micll-ine. Fr'ont-al MALE Õ . Hea,d. verticals absent, brow extended sensill-a aLrsent-. Byes bare. segment 3 of palp with group of 7 capì-tefiate A'R' O'75' setae. pa-Ipal f<¡rmu1a O.2 z O.4: 1.O : O.9 : l-.3' }n!ggg' (22¡t) Last Flagellorn eres 2 and 3 with 2 slender and I stout type c seta ' flagellomere with numerous type D setae. Tlbg:-gl. Ànte¡:r:onotal' lobes not' minute' contic;ur:us, 3 f ateral set¿re. Scutal process ]uarÇe. Acrosticirals pre-alars' biserial" clorsocentrals beç¡in ;ius'L before level of hur're::¿rl pits, 9 l ll r¡'m' Asp' R' 3"2' Anal- lobe !S-gå. L.R. O.6- Iif.g." l'engtìr l-'2 - ' ' proclrrcedr Cut curved slighb. R2-r3 lralf rvay between R, arrd R4+5' costa not arched' down. T,e:-mj-üa-ìj..:. Anal p.riirt bar:e, rcru¡rcl.ec, s'Lerna-Lpodente highl;z 165 broadest laterally, corners drawn out into points. Phallapodeme wíth triangular plate. P.R. O.2. Gonocoxite with slight proximal dorsal as wefl-

as dis'r:.al- ventral l-obe. Gonostyle tapered distally and drawn into a terminal- spine. gpES-. I have examined the holotype d and 2d paratypes, Narrabeen, Nevr

South lrtales, Oct. - Nov- 1956, W' w' Wirth'

Additional sPecirnens seen. South Australia: Hindmarsh FaIIs, Id, 2ô 3f iii'L97l1' 9"ix. L97o¡ Parahra' 2ú, 2o.i- , Ió, -t8'xi 'Lg7o, 7ð, 4'Lí' ' '

Genus PARAI¿.IEFFERIELLA Th-ienemann

parakiefferiefl-a T'hienemann, 1936, p. 195 (ín part). Brundin, 1956, p'148' Saether, 1969, P. 138.

Nanocladius Freeman, 1956, p' 338 (in parl-) '

Type species PaËaE¿slferie-I-!¿i 99r:enate' Edw' Parak-iefferiel-la (s- str-) atus ¡ sp. nov. À smatl species, antenna heav,ily cl'.:bbed-¿ A-Il- about o'3' Ì'Íafe rvith distinctive dark brown thorax and contrasting pale yettow abdomen. Fenrarle pale yellow with srvollen egg guides. Pupa wii-h slender spinufose thoracic hoÏn andsmalfpetlesspuriiBonahdominal-segmentsTlan'rIII' Broil squarecl off on ea'ch s;ilib d.istally witJr numerous type D setae (25¡), 2. rj-ng setae' seta' Antepronotum recluced, -lobes separated by scrrl-a1 pr:ocessi one -Lateraf dorsccentrals Acrostichals absent, scutal hr-rmp wj.th tuft of abou'c l-2 hairs; uniseriaL, berses conspicrrc.rrrs, begin afl:er humer"r:al pits, 2 pre*alars; 4 of sctrtelf¿rr selae in single Ero-nsverse row. -tjgg. L.R. o" 5. outer spìr'r rear tj-bja abotLt hatJ.f length of inner, inncr rvi-Lh ¡ruÌnerous basal accesso::,' r66

!{ings' Lerrgtl: setae, comb slightly spiral , tjlcia not expanded distalty' macrotrichia on Rr' O.95 - I.3 nun., Asp. R' 3'3 - 3"6' AnaI lobe obtuse' *4*5 slightly turned R4*5 *td base R, R2*3 u^ding close to end of R4*5' produced one third up distally, ends midway betwee" MI*2 and Mt*n' Costa Tergites sparsery' of d_istance to wing tip. cu, armost straight. Abdomen- lateral setae uniformly setosei sternit-es IfI and tV with one and two respectively,sternitesV-Vlllwithincreasinglylargernedialareasof narroiv scleroti-sation and setae. Termj.nalia (rig. 166) ' Sternapodeme transversely,cornersstronglyreinforced.Platesofphallapodemedeep]-y gonostyre onl1z curved. p.R. o.L - o.2. rnner robe squared medially, slightlY exPanded distallY' FEMALE(8).Asformalebutantennawith5flagellomeres|lst-and.2ndeach basally' 5th wibh with 2 type c setae (18¡), 3rd and 4th sl-iç;htly bulbed O'82 : L'43' 2 basal rinq setae; flagellomere formula I : O'54 : O'75: (nig - 168) . Sternite VITI with onlv 3 Wing as in Fig- 16 7 . Ter¡ninal.ia puff covered with setae on each side; egg guide expanded into rourrdc.i gonocoxite microtrichia. Tergite fx sma1lo wiLh rfistinct medial furrow' st'ernite x witÌì IX drawn into posterior lobe with single strong seta' vaginal apodenLe pro- na.rrow ridge, cerci hemispherical from clorsal- vielv' wide aS SegÎìeni. VIII. jectíng or¡e third into segirrent VII, base one fifth as

Spermathecal clucts without basaf broadening' hrorn (l'1g 1"69) slender ovoicl about PLìPJ\ ( E Cephalo thorax. Thoracic ' ' ' 3 precorneal setae 2nd L7O)1, covered distally v¡ith minute spinelets; ' prorrotal seta, 2 almost as long as horrr; 3 anterior and 2 prcste::ior with 2long setae' anterior and 3 posterior mesonota-L setae. Flîontal platr:

4Þ@ryl.SegmentsÏIandIIIwj--ulrsmaf].peclesspuriiB(l.ig.I7o). rnedielly Tergj-tes II - VfTf with anal .Ly-C1-ì-rected spinel-ets concerrtraÈed Posterj-or of these: near posterior border, greatest nunrl¡er on seglllent ITI' hoo].:s. on tergites II - IV are 3 transverse rows of crally-clirected !/4 segnent length' Setae rrl- - u5, L3 ancl T-'or Lr;-rnd L, longer:r about Arrallobesdirectedrnesally,rJoj'.ntsturncdout-.AnalstltaedistaJ.ly t67 flagellate, close together and direcl-ed laterally' 39 paratypes (reared from TYPE!-. Holotype ó (and pupal cast) and 1ó ' Further paratypes' Eric Bonython Nostoc) from Hindmarsh Falls' 3'vi'Lg72' Reserve,Ló,n,E.iv.197o;Hind'rrrarsTrFalts,16,I.vii.LgTo;tiumbug Scrub,Ió,19,9.viii.Lg6g;LowerHernitage,I9,9.viii.t969;Moculta Ió, g.viii.1969; Parawa l9 ' 3o'v'r97o' Australia: erownhilt Creek dd (swarms) Additional spec imens seen- South ' ls'vi-ii" ld 23.ix. I97o¡ Clarenclon, Iô' 5.vi. 1969¡ Botanic Garde-ns ' Ló ' ' 17.ix.I969¡CIeIandReserve,#,19,17.ii.19'7o;DeepCreek(SouthCoast) 25'x'l:9€>9i Eric Bonytltorr Rese"ve' 5ó, 3.ix. L969¡ Duck Ponds' 26 ' æ ' numerousspecimens,March-Oct'I97OandL97L¡ElvensPoncls'M''.i'/' 24.viii.1969il,ishers,Id,2.x.,ld,23.x.L969¡IJeaIey.sIó,I9(rear-.ed fromfilamentorrsaigae)6.xi.1.969¡lîindmars;lrFalfs,numerousspecit'rens' (reared from rniclstream l eaf accumula-tion) ' I"larch - Oct. 1969 - L97L¡ óó99 5ó 9'viii"1969; Inman Valley' 2.v.72 - 13. v.L972; Hrunbug Scr'¡b ' Kuitpc, 3ó, l'vii"L969; Lowcr:: numerous speci-mens, Aug. - oct. 1969; Hermi.tage,J-6,9"viii.1969'lvlarneRiver,Ló,9.vj-ii..L969¡Morialta,Ié, 23"vii''Lg7o; Parawa' numel'ous specimens 4.!x-L969¡ Torrens River ' Lô ' Aprj-t-Sept.Lg6g-L97L¡Id(emergencetrap)f.2.v.19?1;dd99(rearecl dó99 (at light) Sept' LgiI' from silt washed from leaves) August 197ft Falls , !'ô 28'ii 'L962 (p' u' Australj-an capital Terr:itory: Ginni:ld,erra '

Colless) .

Genus PSEUDOSMT:I'TtA Goel-ghebuer

p' L27 (as suirgenus of Sntittia t Pseudo smittia Goetqhebuer, 1932' p' 663 Brundin, 1956, p. 165; Freeman' 196I'

(Srii.ttia) Edwarcls (L929) ' Type sPecies Spani,otonìa il¡qgÉÂ c:f F¡:eeman (1961) ' fhe follovring key is t:':sed on that 168

KEY TO AUSTRALIAN SP ECIES OF THE GENUS PSEUDOSMITTIA

I *4*5 fo".d with costa imperfecta (Skuse)

These veins not fused 2

2. Eyes bare 4 l'reeman 3 Eyes pubescent latifurca sipennis 4 A.R. O.4 cïas

A.R. > O.5 5

5 Apex of antenna diamond shaPed, sirrgle inner lobe scutella, sP nov.

Apex of antenna normalr2'd'istinct inner lobes ar%11l1, sp. nov.

P seudosmittia argills' sP. nov. Anunremai:kablespecies'separaLedonlybyitstaperedantennaarrd terminalía. punp MALE (3). Head. Brow drawn d.own, frontal sensilla absent. cibarial with- nearly straight dorsal border. M.R" 0.36. Segment 3 of palp ;with single tapered sensillum. Palpal formula o'3 : o'5 I : I.23 1.70. Antenna. A"R, L.2- Flagellomeres 2 and 3 with 3 slender type C setae, point (l'ig' L72)¡ 2 - g broader than long. Last flagellomere tapered to a about 3 type C setae, 4 _ 5 ring setae. Thorax. Antepronotal lobes pits' contiguous, several lateral setae' Dorsccentrals begin above humeral uniserial' scutal hump with several short setae. 4 pre-alars, 6 scutellars' L.R. O.5. Wings. Length 1'50 run" Asp' R' 4'2' Squama bare' a:ral -L .. lobe stight, mac:rotrichia on basal vein a¡rd RI, RZ+3 ends closer to R4+5' between M1*z .ld *3-r4, .?:tt qriglltty prgduced' An ends R4*5 "rd" midway (nig. 173). distat of pfk. j\bclomen. Tergites uniformly seto se. Terminalia reduced' AnaI point small , rourtd, setose. Sternaporimal and riorsal , the other setc)se, distal ancl veni:ral' Gonost)¡Ie

shaped. r69

Lake TYPE.S. HolotYPe d, 2ó and 39 paratypes fro¡n Rum (claypan between Eliza and Lake St. Clair) 23'viii'1969' ód Additional spec imens seen. south Australia: Run 23.viii.1969.

Pseudosmi ttia scutella sP. nov. apical club of QuickIY distinguished bY the flattened diamond-shaped the antenna.

MALE (2 .Head.Browstraight.Frontalsensi]-laabsent.M.R.o..]9. (8P), palpal formul¿ O'3 : Segment 3 of palp with single tapered sensillum o.5:l:I¿L.4.44!_enne..A.R.o.9.FlagelJ.orneres2and3eacì].¡itlr3 distal- dia:r';;r"ì- broad-based type c setae (3OP). Last flagellomere v¡ith 6 ( setae vrit}¡ shaped club (FiS. L7ã> , numerous type D setae 3O,p) , 3 ring l-obes con- circlets and 2 mi¡:ute terminal sensill-a. Trto¡"". Antepron<¡tal begin tiguous, scutellar Process smal.]-, 3 lateral setae. Dorsocentra-fs

pre-alars' 2' above humeral pits, scutal trump with several small setae, 5 scutel].ars.LegS-.L.R.o.5,bothspursotreartibiawithaccessories. anal lobe wing. LengÈh 1.3 mm., Asp. R. 3.6. Squamar with.4 short setae' srnarl. lvlacrotrichia on l¡asal veitl Rt, R4+5, R2*3 encls closer to R4+5' R4*5'.'=levelof',].*2lCoStêsl1ghtlyprod'uced,Ànenclingdistalofprlk. llergite Termj.nal-ia. Anal point very short, rounded, setose' Abdor¡-en' phalla- setae sparse, biserial-. Sternapod,eme rectangular. corners recluceci' + (rig-' T7 s . (.ìono- podeme triangular, P.R. o.23. Inner lobe broad based )

style boat-shape-d rvj-th small tennj-nal spine' TVPES. Holotype d and ld paratype, Etíc Boriythorr Reserve, LJ"ii'I971' Reserve, ld, Ad,fi-'bional sÞec im.ens seen. sor:th Australia: Erj-c BonythOtr g.iv. Lg¡o, ld, 29.i--, 1ó, 39, 17.ii., 2ô, l9'vii' , Ló ' l-'rr'1971'

Genus ÐOLOPLAST'US Skuse 1917' p' 228¡ DolopI¿:stus Skuse, ltì8?, p- 26-J¡ Kieffer, LgO6, P' 28¡

.t'reem'ln, J-961 , P. 611 .

Type sPecr.es Dolopl-a-s tus monticc.rla Skuse. In roilescrihir,g t-he si-ng1-e represerrtative of tirj-s genus lrreemalt (-1 96I) Ej.g. L72 Pseudosmittia ar9ft., sp nov. male, aPex of antenna

I Fig. 173 il terminalia

Fis. J-74 Pseudosmittia scutella, sP. nov. maIe, terminalia

tt Fig. 175 tt apex of antenna

Fig. L76 Doloplastus crassistvlus, sp. nov. male, gonostYle

nov. ma1e, terminalia Fig. L77 Doloplastus curtatus ' sp.

Fig.178 Dolopl astus culcitellus, sp. nov. ma1e, wing

ll Fig. I79 I terminalia

flag- Fig. 18o Dolopiastus sensillus sp. nov. male, antennal ellomeres2and3and aPex of antenna.

tt gonocoxite Fig. 18tr. male, l-- t

ç a

J s, 172 ìt 174

173 176 175

,;iiÌ:iii i¡ ., nn;"

1 177 179 .l

l.------r 180

, 1y', >\¿l 181 t 178 170 expressed dor¡bt a-bout its status as a genus, separate from smittia. The discovery of the 4 new species described below, aII of which share with monticola a low 4.R., cask-shaped flagellomeres and a bare squama and 6ne of which also has a reduced number of flagellomeres' su99e sts Doloplastus is a valid genus, at least among Austral-ian species' I have modified Freeman's generic diagnosis to include ttre new species from South Australia. I4ALE. Head. Brow drawn down to varying degree. Frontal sensì lla a-bsent' AntennÈ. A.R. less tlran J-, most flagellomeres more than twice as long as broad; nunber of flagellomeres sometimes reduced. Thorax' Antepronotum slightly reduced, lobes not in contact, few lateral setae, scutal process broad. Acrostichals absent, dorsocentrals uniserial. Legs. Rear tibia with,outer spur less than half length of inner, comb wit} setae reduced in size and number, pulvilli absent, all tarsal segments cylindrical. iqiog. Length less than 1 mm.t squama bare, anal lobe very srnall or absent. l,licrotrichia invisi-ble, punctures minute. RZ*: sometimes partly merged

R Costa sometimes retracted- C.I curved down distaÌIy . Terminafia. with 4+5' Anal point sometimes present. Two inner lobes sometimes present, gonostyle variable.

KEY TO SPECTES OF THE GENUS DOLOPI,ASTUS I. Antenna with 13 flagellomeres 3 Antenna with less ttran 13 flagellomeres 2 2 (1) Costa not retracted; anal point presenti antenna with only seven flagellomeres monticola Skuse Costa retractedi anal point absent; antenna with twelve flagellomeres curtatus sp. nov. 3 (t) Costa not retracted; inner lobe small, rounded, type C setae enlarged sensillus sp. nov Costa retracted; inner lobe pointed or extended, type C setae slender 4 I¿¡

4 (3) Inner lobe extended into a cushion-Iike pad;

gonostYle boat-shaPed culcitell .us sp. nov. Inner tobe small, pointed; gonostyle stout crassistvlus sp. nov.

Doloplastus crassis tvlus, sp. nov. Distinguished by exceptionally elongate flagellomeres and stout gono- sty1e. drawn MALE ( s). Head. Verticals, post orbitats absent, brow only slightly down. M.R. o.24. segment 3 of palp with one rod-Ij-ke sensillum. Palpal formula O.3 : O.7 : 1 : 1 : 1.8. Antenna. A.R. O.5. Flageltomeres 2 and 3 each with 3 type C seta (1O¡) , 4 - 12 from 2 to 5 times as long as broad' Last ftagellomere slightty clubbed at apex with 2 type c setae, 2 ri'ng setae and several type D seta (B¡). Thorax. Two lateral setae; dorsocentrals begin after humeral pits. Three pre-alars. Legs. L.R. O'5' Wing. Length t.O mm. R¿*S .td1ng midway between Cut and M3+4' costa slightly produced. Pfk distat of r-m. Terminalia. Anal ridge alcsent;

sternapodeme highly archedi corners drawn out, lateral apodemes broad' phallapodeme with rectangular plates, penapodeme a]¡sent. Gonocoxite with small pointed inner lobe. Gonostyle stouÈ, rvith concave inner surface

(ris. l-76) . TypES. Holotype ó and ló paratype, Carey's Gully, South Australia, 2.x-Ig6g. Further paratypes. Eric Bonython Reserve, 1-ó, 2O.v.I97O¡ Parawa, Ió, 19.vii.1969. 26 L9'x'1969; Additi-ona1 ]-mens s . South Australia: Healeyrs, ' parawa, ljô, 19.vii.1969, Ló, f -iv.l97o, 3ó, 26'v'1971'

Doloplastus curtatus' sp. nov. separated from most orthocladiinae since only 12 antennal flagellomeres. Readily recognised by the transparent wings and the proxinally

expanded gonostyle, which lacks a spine' slightly drawn MALE 2 . Head. Verticals extend almost to midline, brow

down, M-R- O.3. Seglnent 3 of palp with group of 2 or 3 capitellate l-72

Sensilla (12p). Falpal formula o.5 : o.5 : 1 : 1.45 : 2.3. Antenn{. A.R. o.3. Only 12 flagetlomeres present, aII fusiform, longer than broad,; 2 and 3 ;¡ith 3 rod-Iike type C seta (I5¡r) | 4 - tI about three times as long as broad. Last flagellomere clubbed at apex, with at least 2 type C setae, 3 ring setae and numerous type D setae; furrow extending right to tip. Thorax. Three lateral setae, acrostichals absent; dorsocentrals begin close to pronotum; 3 pre-alars. Legs. L.R. O.5. Wings. Length 0.6 mm.' Asp. R. 3.3, Squama bare, anal lobe absent. Macrotrichia on R, only single rovü on costa; microtrichia minute, punctures invisible. R2*3 *tt9ed with R4*5, R4*5 *""ts costa at level of Cut, costa produced slightly distal of r-mr Cu, curved downwards, An ending under r"m. Terminalia (FiS' L77) ' Anal point al¡sent. sternapodeme squared, cornels reinforced' Phallapc'deme with elongate p1ate. P.R. O.2. Gonocoxite with curved, pointed dorsal Iobe, slightJ-y rounded ventral lobe. Gonostyle with flattened outer surface drawn into a pointed heel proximatly, spine absent' TYPES. Holotype d and one ó paratype from Eric Bonython Reserve' 3'iii'l'97}' A third specimen from Parawa, 25.iii.I97O has a slightly larger and rouncler heel and the head was lost: it may lepresent a second species. additional sp ecimens seen. South Australia: Botanic Gardens ' L36 ' 4.ii. L97a; Hindmarsh Falls , Lô , 22'ív'L97o'

,Dolop lastus cul-citellus, sP- nov.

Separated by its shortened wing venation and cushion*like inner lobe' l"fALE(5).Head.Verticalssmall,extendhalfwaytomidline.Browex- tended into moderate medial point with 2 - 3 slender bent capítellate setae(9y)rpalpalformulao'3:o'5:1:1'3:I'5'M'R'o'33'Alrtennqe' A.R. 0.6 - O.7. Flagellomeres 2 and 3 with 3 slender type c setae (2oP), 4 - L2 at reast twice as long as broail. Last flagellomere stightry clubbed atapexwith2typeCsetae,numerousDsetaeand3ringsetae,furrow extending to tip. Thorax. single lateral seta. Humeral pits sliqhtly enlarged, acrostichals minuÈe. biserial, dorsocentrals begin close to L73 pronotum, bases large, 3 pre-alar setae, 6 scutellars. Legs. L.R. O.4. Tarsus 4 shorter than 5. wing (rig. 178) . Length o-9 mm., Asp' R. 3.9.

Squama bare, anal lobe small. Macrotrichia on basal vein. Microtrichia invisible, punctures minute. R2*3 almost completely merged with Rn*r'

Rn*U ending proxi-lnal of t3*4. Costa slightly produced' Pfk distal of r-m' An finishing distal of r-m. Abdomen. Tergal setae sparse, in two transverse rows. Terminalia (r'ig. 179) - Anal ridge present' Sternapodeme rectangular, corners drawn out. Phallapodeme narrow. P.R' O'I' Inner lobe drawn into a cushion-like pad covered with fine setae' Gonostyle narrow' boat-shaped. TypES. Holotype d and 1ó paratype, Parawa, South Australia' 19'vii'L969' Further paratypes: Botanic Gardens Id, 11.vi.197oi Healey's, fd, 18.x.L969¡ Parawa, Lô, Io-xii.L969' Additiona1 specjmens seen. south Australia: Botanic Gardens, 6 , IO.iv., ld, 16.iv.I97O, 16, 24.ii., 1ó , 25.iii.197l; Eric Bonython Reserve, ld. 3.iii .IglL¡ Healey,s Id , 2.xí-Lg6g¡ Hindmarsh Falls, 13ó, I-x.I969, 6 , 28.i.L97O¡ Parawa ód ¡an. - April , Lg7o, L97L, l]ó, lO.vi.L97O¡ Robertsons' 2ó, 26.xii.1969.

DolopIas tus sensillusr sP- nov-

Separated from rest of genus by its enlarged type C setae; wing venation not reduced- Segment MALE (2). Head.. verticals extend half way to midtine, M.R. o.32. 3 of palp with 2 diverging, Iong' tapered sensilla' Palpal formul-a o'2 : o.4 : 1 : l : 1.4. Palp greater than half length of antenna. Antenna. A.R.0.6;tastflagellomereslighttyknobbedwith3distaltypeCsetae and 3 ring setae. AlI type c setae greatly enlarged (rig' 18O) ' Thorax-' 'rnedially' Acrostichals Antepronotum with 4 lateral setae, lobes in contact pre-alars' absent, dorsocentrals uniserial, beginning above humeral pits, 5 L'R' 6 scutellars. Abd.omen. TerEites uniformly, sparsely setose. Legs' o.5.Combstraightrwithfewsetae.Bothspurswith.accessorysetae, L74 inner less than half length of outer' W!ryg-' Length l'5 mm'' Asp' R' 4.L. Anal lobe small, squama bare; macrotrichia on base of Ra. Micro- trichia invisible, punctures minute. *2*3 ends closer to R4+5' R4*5 ttd" midway between Mr*, .td M3*4. Costa produced one third of distance to wing tip, Pfk distal of r-m; Cut bent, Ant distal of r-m. Terminalia. Anal point a.bsent; sternapodeme thin, evenly rounded. Gonocoxite with 2 inner lobes, one dorsal, proximal with rounded corner, the other ventral, distal, scarcely detectable (FiS. 18I). Phallapodeme with Èriangular platesr gonostyle boat-shapedt spine small' TypES. Holotype d, Eric Bonyttron Reserve, South Australia, 3f iii'1971' Paratype d, Eric Bonyttron Reserve, f ix'L97O'

Addi tíonal specimens seen. South AusÈralia: Hindmarstr Fa11s, L6, 24.v.I97O (one of hundreds roosting on moss just alrove \¡tater line) '

D. curtata resembles some species of saunderia sublette in the shape of the gonostyle and absence of a spine, but cannot be included in this genus since it has reduced wing venation and bare eyes. It resembles the sarrnderia in the reduced nr¡nloer of flagellomeres, but in saunderia reduction is due to a fusion of the basat flagellomeres and this would probably not be ttre case in curtata since sensilla are on both flageltomeres 2 and 3. Both curtata and culciÈe1Ia resemble Krenosmittia but lack Pulvil-li' jmmatrllles In Krenosmittia the anal point of the male and the are distinctive, So I have refrained from placing the new species here. 175

SECTION IV PHYLOGENY

1 TNTRODUCTTON since this study is intended as part backg'round for a historical biogeographical investigation which may conflict with wide-ranging proposals made by Brundin (1966), using Hennig's (1966) methods, the methods of taxonomic analysis needed to be carefully selected' The logicai- place to look for guidance in selecting a suitable approach is to the opponents of Brundin and Hennig. Foremost among these are the simpson-Mayr-Darlington group who claim the title phylogeneticists' However, as Darwin apparently coined the name phylogenetics for the study of relationship through descent, I will reselve 'phylogeneticist' for the Hennig school and call Mayr et aI. the 'evolutionistsr. The evol-utionists d.efine relationship in terms of the similarity of genetic material and thus, wtrile acknowledging for example that crocodi-l-ians diverged later than reptilians from the evolutionary Iine which eventually produced the birds, they none the less group crocodilians and reptil-ians together, separate from the birds. Mayr justifies this approach (Mayr,

1969)...InviewoftheimportanceofaclassificationaSaninformation from retrieval system \^7e recofimend that genetic relationships resulting parallel evolution be given the appropriate weight in the delimitation of higher Èaxa." Historicat biogeography requires the determination and expression of Lhe order of splitting of groups as well as information retrieval and, from the example above, it can be seen that Mayr's analysis guite precludes any adequate e>çression of the time of splitting' It al-so overlooks what might be an important piece of information'- i'e' birds and crocodilians form a sister group and share some features not possessed by reptilians. Hennig clefines relationship ín genealogical terms and in the above

example separates crocodilians and birds' as a 'sister-pairt, from the reptilians. His system thus provides a recognition of splitting processes' L76 as required by biogeographers, but still loses some information; information which Mayr considers more important since, as he points out, similarity of genetic content through convergence may be greater than thaÈ conferred by descent. Biogeographical conclusions will be short lived if the classification on which they are based must be restructr.rred whenever a new group is discovered. under Mayr's system the discovery' for examirle, of an intermediate between birds and crocodilians would require redefinition of one or other of his major categories. Hennig, however, has provided the taxonomist with a logical procedure by which an attempt can be made to reconstruct 1-he exact course of evolution. Evolution has proceeded along a diversity of Iines, but a proper Hennigian reconstrucÈion of it should readily accommo-

date any new discoveries without major re-arrangement. Thus the evolutionist's criticism of Brundin and Hennig is based on the relative capacities for information retrieval of the two types of classification. The evolutionists provide no logicat analysis of Hennigrs methods for reconstructing evolution (separate from making a classificatiori) and determining the type of relationships that are important in biogeography, and thus are no help in evaluating Brundin's r¡rork' provided Herueig says that evidence for genealogical relationships can be only through synapomorphies which are not a result of convergent or parallel evolution. The processes by which apomorphic characters are recognised has brought accusations of both circularity (sokal & sneath, L963; Colless, 1969) and of producing a phenetically based classification (Colless , 1969) ' Certainly at the very first approach, ânY classification wiII be phenetically based' However, when after background investigation different characters and' their character states are re-examined with reference to fossil sequences, to their possible functionrand in the light of evofutionary theory, the initiat cfassification can be modified: it is no longer strictly phenetic' L77

After this it :may be found that unconsidered characters support the groupings proposed. Further reconsideration will result in a closer appLox- imation to evolutionary reality. This process $¡as call-ed "reciprocal illunination,' by Hennig and, as Hull (1967) pointed out, is not circular. Hennig's method has also been criticised for its use of intuition'.

However, most taxonomic constructiorrs are partly i¡rtuit-ive hypotheses which are tested when a new species is found or a nevl structure investigated. popper (1959) in his analysis of scientific method says there is no logical background for the process of forming an hypothesis as there is for testing it once it has been presented. Thus there is room for intuition provided that it is carefully circumscrihed. In Hennig's analysis the only place where intuition is used is in the preliminary assessment of apomorphies and eljmination of parallelism. Hennig has attempted to regularize the use of intuition by providing copious guidetines for the recognition of true synapomorphies and by introducing an 'auxil-iary principle' (Hennig, 1966, p'I21) to cope with parall-elism. He also frequentty expresses the need fo:: caution in applying the methods he proposes. This contrasts with the relatively random use of criteria by the evolutionists (e.g. they overlook parallelism when considering certain taxa) and pheneticists (who must choose the scoring method, range and type of analysis used in their numerical studies) ' Thus from a theoretical point of view at least, Hennig's system of analysis seems more suitable for the task of biogeographical investigation than Mayr's system and the major criticisms which have been made of it trave not been substantiated. Therefore vte must look to ttre practice of Hennigrs methods to discover the procedures which have stimulated the heated argu- ment of recent years. Unfortunately Hennig's followers are not always as cautious in their use of intuition as Hennig is and neither are they always rigorous in their expositions. These workers often make assumptions abouÈ the value of synapomorphies without justifying ttrem, bu'b despite this' terms¡ express their conclusions about rerationships in highry objectíve 1.78

It is here that some criticism is justified'

IS 2 BRUNDIN'S USE OF HENNIG PHYLOGENETTC SYSTEMAÎICS

As an example I shall consider Brundinrs treatment of the genus paraheptagyia (Brundin, L966, p. 375). Brundin describes 7 species of this

t genus and divides them into the cinerascens ' group canerascens nitiscens

and andinus from South America) and the 'semiplumata' group (semiplumata

and umbracul-ata from South America and tonnoiri and tasmaniae from Australia). I have described a further two Austral-ian species of Para- heptagyia, plumata and ennea which appear to be member s of the semiplumata group. Figure IB2 is a diagram I have constructed to show the pattern of apomorphies used by Brundin to determine the relationships between the species. Species of Paraheptaqyia are listed across the top and characters I - 11 are indicated as being present in the apomorph (dark squares) or plesiomorph state (Iight squares). The characters and their states are as follows' Character l. The zone of enlarged spinelets along the anterior margins of the abdominal segments of the pupa may be black and sharply delineated (apomorph) or only faintly indicated (plesi-omorph)' character 2. Inverse spinelets may be found along the posterior margins of the alcdorninal segments of the pupa'

(i) On segment II these may be enlarged and found in a mesallY bisected black zone (apomorph) or small, uniform on all segments (P1esiomorPh). (i-i) These may be quite absent (apomorph) or small and present o¡r aLl

segments (P1esiomorPh)' Character 3. The fibrillae of the antenna of the maÌe may be reduced in (ptesiomorph)' number and size (apomorph) or the antenna may be fully plumed character 4. The enlarged terminal spine of the gonostyle of the male terminalia may be absent (apornorph) or present (plesiomorph) ' character 5. The inner surface of the gonocoxite may have a large and complex lobe, covered witl¡ humerous strong setae (apomorph) or it may be 179

Fig. Ig2. Inter-specific relationships within the genus Paraheptagvia.

Schedule of argumentation for the establishment of sister groups'

cinerascens 9p. semi-plumata gp.

tvl lJl ¡al tdl s/ Øl í¡t 7/ +J FI D mm trmm ? 1. Enlarged spinelets (P) 2é) inverse spinelets (P) u ün mm E lflil mJ ? (ii) inverse spinelets (P) ! ft ! xrl ? ? T nn mün I uuflmm 3. fibrillae (A) D un nll n xntr 4. shape of gonostyle (A) 5. inner surface of trn M mt]uil gonocoxite (a) trn m mflum 6. acrostichals (A) trf, ? mmn 7. pupal stripe (P) trx M mmm 8. heels (A) xr tr mmm 9. flagellomeres (A) MM x trxE 10. Ieg hairs mm x trxn 11. style (A) m tr ples j-omorphy m intermediate stage tr data unavailable 180

smooth and hollowed (plesiomorph) ' Character 6. Acrostichal bristles form a short row (apomorph) or a long

ro\^t (plesiomorPh) - Character 7. A marked black stripe along each lateral margir of a-bdomirlal sternite VII of the pupa is present (apomorph) or absent (plesiomorph). character 8. Exaqgerated heels on tt¡-e gonostyles are present (apornorph)

or absent (Plesj ,rorPh) . Character 9. The number of segments in ttre antenna is reduced to 13 (apomorph) or normal- (plesiomorph)' character Io. setae on the legs are conspicuously short (apomorph) or:

normal (plesiomorPh) - Character 1I. The gonostyles are Lurned up distally (apomorph) or normal

(plesiomorph) .

AII members of the semiplumata group possess the apomorphic state of characters 1, 2(i) and 3 and, according to Brundin' can therefore be considered a monophyletic group. However, most Heptagyini have black' sharply delineated. spinelets as in character I and the reduced condilion in tåe:

t cinerascens t group coutd therefore represent a return to the primitive state and ttrus an aPomorPhY. In addition, this similarity in the 'semipluma.la.' group may well be due to parallel development for, as Brundin himself states, pupal structures in chironomidae have been subject to strong environmental influences' In the

Maoridiamesa group there are similarly delineated black spinelets of slightJ-y different structure, obviously a convergent development and this' pressures as weII as their widespread presence in Heptagyini illustrates the that exist for inducing such structures to develop' Brundin rnust have discounted these possibilities in evaluating character I. Hennigrs 'auxiliary principle' suggests that the possibility of parallelism can be discounted but I consider that its use in such a doubtful si.tuation is ill-considered and that character I can be used only in 18I phenetic considerations or, if it is purety 'intuitively' weighted, as is surely the case here, it should be given 1ow value as an indicator of phylogenetic relationshiP.

The particular form of the inverse spinelets in character 2 (i) is unique and, while it is possible that environrnental pressure may have ]ed rauxilia::y to a paraltel development, it seems reasonabfe here to use the principle' to dismiss the possibility.

The use of the auxiliary princi-ple is also justified to dismiss the possibility of the reduction in antennal fibrillae being due to pa::a1lelism; reduction such as this occurs very rarely and, since close relatives of Chironomidae (e.g. Ceratapogonidae, Culicidae) have numerous long fibrillae, there is little doubt that this is a meaningful apomorphy. Ho\,vever, males of one of the new Australian species, plumata has v¡el1 developed f-ibrill-ae yet possesses other characterj-stics of the I semiplumata t group. In the absence of information about immatures of plumata, it could be suggested either that this forms a plesiomorph sister of the 'giPiujnq.t3' group or, if the adults alone are indicative, the monophyly of the 'sçm!- plumata' group rests on characters from the single adaptive complex represented bY the PuPa. Brundin next establishes the reciprocal monophyly of the 'titttun""lqr' group on the basis of its members havi¡g the apomorph state of charac+ueÏS 2(íi) and 4. The apomorph states of both these characters aIe unique within Heptagyini, and the plesiomorph degrees are widesp:lead in other Chironomi

The heels referred to in character 8 are much smaller in other !-ara- heptagyia: their exaggeration certainly seems to be apomorphic' However to entirely exclude parallelism seems unjustified since in Podonominae also the most exaggerated heels are found in the Australian species' The evidence of the new Australian species, plumata and ennea, both with heels of intermediate síze, suggests that. ttre heel could be a graded response to

some environmental variable. However, whether the enlarged heels are really

apomorphic nay be questioned since Brundin has not discussed the possibility that the heel is merely the second lobe of a plesiomorphic double gonostyle. The enlarged heels may be a ptesiomorphy of the Australian fauna: a modified

second lobe of the gonostyle retained as a response to environmental pressures similar to those which produced the lobe in Podonominae. A reductíon in the nurnber of flagellonteres (character 9) is found in tonnoiri, tasmaniae and to a greater degree in ênnea. Since this is rarely encountered in chironomid males it probably indicates relationship between these species. Brundin uses the remaining characters (tO, 11) to establish the monophyly of the semiplumata-umbraculata species pair. However, with tkre discovery of the new Australian species, it seems that the length of leg setae(characterlo)showsacontinuousvariationinthe.giP@qg' group. since the gradation is a simple one and different degrees may thus have arisen on a nr¡rlber of occasions eith-er by reversal or parallel development, it is unjustified to use one end of the gradation as evidence of monophyly of this species pair. Likewise the degree to which the gonostyle is tr:rned up shows a continuous variation in the group, particutarly with the discovery of ennea, and this character should not be used to show mono-

Phvrv- Thus Brundin has based his separation of the 'cínerascçr¡q' amd 'Eo*i* Despite pjl]Jlnata. groups on five ctraracters from two different life stages' the above criticisms it seems probable that these do form monoplryletic r83 groups. However in separating two groups withín 'semiplumata' Brundin than made use of characters whictr showed continuous variation' Rather stretching the procedures of phylogenetic analysis to propose a complete set of relationships a valid conclusion about the 'semipluma'Ee' group would be that it shows a closely interlocking pattern of relationships' AswellassettingoutaphylogenyofPg:aheptagyiaBrund.ingoes further and states whictr species or species group of each- sister-pair is the more apomorphic. For example, he says that Èhe '[email protected]"u*' group rsemiplumatat ttabove stands out as more plesiomorphic than the group all because of weII developed and normally plumose male antennae and only faintly indicated transverse zones of enlarged spinelets at ttre anterior margin of the abdominal segments in the pupa. Apomorph is, on the other of the hand, ttre absence of a terminal tooth (t seta) on the dististyles; at the same nature is perhaps also the total absence of inverse spinelets posterior margin of the abdominal segments in the pupa." Brundin does not pupal explain how he determined that the plumose antennae and enlarged groups is spinelets weigh especially heavily in deciding which of the two methodo- more plesiomorph. In this type of sÈatement he far exceeds the Iogical limits of Hennigrs system and, although Hennig says thaÈ of everi¡ sister group one of the pair will be more apomorptr ttran the other' he does not expect this will be readily detectable in practice' Paraheptaqviawaschosenfordetaileddiscussionbecauseitisthe is avail-able' only genus discussed in depth by Brundin for which new material involved in ft can be seen that much more uncertainty and intuition are close Brundin,s evaluations than the tenor of hís exposition woul'd suggest' provides no support- examination of other parÈs of his work reveals that he phylo- ing statements for many of his assessments of apomorphy or of the genetic value he attaches to them. At the level of relationships between conclusions'How- species this approach is likeIy to lead Èo readily refutable .ever;aÈthelevelofrelationshipsbetweengeneraorlargegroupsof r84 species, Brundin's conclusions are often broadly based; derived' from a consideration of characters from all life stages and are thus likely to provide a reasonable estimate of Èhe true situation despite his lack of caution. Tt is unfortunate that because Brundin has proposed relationships between all species and framed them wiÈh equal emphasis irrespecti-¡e of the reliability of evidence, he has precipitated unnecessarily strong cr:iticjsm of h.is whole work and quite unfair criticism of Hennig's 'phylogenetic systematics. '

3 PHYLOGENETTC SYSTEMATICS AND THtr EVOLUTIONARY INDEX Illies(1961)proposedaphylogenyforthesixteenfamiliesofPlecop- tera, and in order to represent the relative degree of apomorphy of each famity quantitatively he calculated an 'Evolutionary Tndex'' under this scheme rlties considered 15 characters. Beginning with the most plesiomorph the character sÈates l^Iere given a val-ue by nr:rnbering then and t'hen these values were totalled for each familv' The highest scoring family was thus the most ,apomorphicr' and had the highest 'Evolutíonary Index" Illies was considering well documented characters in which there seemed little evidence that reversals had occurred a¡rd the character phylogenies were readily determinable. For all practical purposes his

analysis was ProbablY correct' However closer examination shows that calculation of the EvoJ'utionary Index has a phenetic component, since a measure of the 'distancer f::om a hypothetical ancestor is being calculated. By this means, a group of taxa tevolved.l In contrast' can be placed in order from least 'evolvedr to most defined' Hennig says that a hypothetical conmon ancestor witf not be readil-y but by careful consideration the apornorptric forms of certain charactel:s might be separated from plesíomorphic forms occurr in that : the within a terms apomorphic and plesiomorphic are relative and stand onJ'y small group. Thus onty infrequently will it be possible to estimate which of two present-day taxa is the most apomorphic' 185

Later v/orkers (Saether L97L¡ Sch1ee 1968; Hirvenoja 1973) have taken the liberty of combining estimates of the Evolutionary Index with theír estimates of phyJ-ogeny based on Hennig's methods. However' the two estimates have different theoretical bases and would better be considered as t\¡ro separate estimates of the relationships within a group of taxa. An examination of the phylogenetic analyses of these later authors provided by shows an even more loosely documented approach than that Brund'in.Forexarnple,Hirvenojapurportstobeabletoarrangeinorder, from plesiomorph to apomorph, the character states o1 7O characters by use of of which he is able to prepare a phylogeny for 24 geneta and groups species. of Hirvenoja also proposes sequences for the evolutionary development 6l characters and, using these, he estimates the distances of 77 species of from their hypothetical ancestor. He then combines these two estimates relationships into a diagram wt¡,ich implies that the relative synapomorphy (1971) and of eactr species or group can be determined. Earlier saether Sckrlee(1968)conductedsj:nilaranalysesofothergenera,thoughvlith slightlY more documentation' Afurthercriticismoftheapproachusedbytheseworkersstemsfrom true phenetic their employment of a'relatively'phenetic approach. In a to be made' the approach where some measure of multidimensional distance is and Èhe more direction in which character states are scored is unimportant' the characters that are scored, the more accuÏately is the specíes represented. In using the Evolutionary Index, the direction of scoring is important.Ho\^Tever,onlyforafewcharacterswillthed'irectionof evolution be accurately estimable and as additional characters are will diminish evaluated the probability that this estimatj-on will be correct Theo::etically' r:ntil it is no greater than that exçected by chance alone. then, as more and more characters are evaluated the random component may a group of species outweigh the non-random and the evolutionary indices of tB6 will approactr each other. This may be happening in Hirvenoja's analysis

(I¡irvenoja, 1973, P- 56) .

PROPOSED PHYLOGENIES 4 A METHOD OF INDICATI NG RELIABILITY OF The sor¡rce of much of the practical misuse of Hennig's phylogenetic systematics is the failure of this methcd to provide the user with a ready any means of indicating his degree of conviction as to the correctness of statement about phylogeny. In order to do this at present, extensive written documentation must be provided' The major group under consideration in this work is the orthocladiinae' for which few characters are available which are of any assistance in phylogenetic analysis. Thus it is important that characters which are available should be used to the best advantage. Tn this thesis I am therefor:e proposing a system for rating the reliability of estimates of monophyly' to a nore suctr a system, if widely discussed and elaborated upon, could lead critical approach to the construction of phylogenies'

LevelA.Monophylyisbasedonthecommonpossessionofanotherwise has uníque character state, for which the possibilitl' of parallelism apparentlY been etiminated' state Level B. Monophyly is based on the colÛnon possession of a character either a which is not unique, but for which it seems possible to exclude of the reversal of evol,utíon or parallel deveropment as being the cause similaritY. state Level c. Monophyly is based on the common possession of a character develop- which is not unique, and for which neither reversal nor parallel ¡nent can be safelY excluded' Level D. ,I4onophyly is very tentatively proposed'

Y APHROTENT]NA-E 5- PHYLOGENETIC RE LATIONSHTPS V{TTHIN TIJE SUB_FAMII as Brundin used mainly pupal characteristics to establ-ish Aphrg!9qlq the pt'lp¿ the plesiomo::ph sister of Paraphroteni-a and Aphrotenie-11-g' Since ofAphrot".iq3lE!qe]gs'.l,tlresingleSouthAustralianrepresentativec¡f 187

this sub-family, is unknown, no evaluation of Brundin's proposal can be

made Lì A. australiensis the ninth tergite is not emarginated, the styles are stouter and the inner lobe is slight - characteristics which

would make it more apomorph than the two African species, accordíng Èo Brundin's criteria. Such an evaluation would not conflict with Bnmdin's proposals of trans-antarctic migration (1Se0, p' 45O) '

ORTHOCLADITNAE 6 PHYLOGENETIC REI,ATIONSHIPS IiüITHIN THE SUB-FAMILY

The phylogenetic scheme proposed by Brundin (1966, p. 424) relating

all members of the Family chironomidae is represented in the figure below' Brundin tentatively proposed Prodiamesa as the apomorph sister group of Heptagyini + Diamesini (i.e. Diamesinae) and recognised T"f*g!ggq!on. as the apomorph sister group of Diamesinae *? Prodiamesa thus elevating

TeIma tontosub-familystatusasHennig.stheoryrequires.Brurrdindid not reach firm conclusion s about Prodiamesa and he left the remaining ,Orthocladiinae' together as the apomorph sister group of Diamesinae *? prodiamesa + Telmatogetoninae, remarking that 'a hindrance for further progress is lacking information of the larvae and pupae of" ' s.9me plesiomorph genera of the southern Èemperate zone.' unfortunately some of these immatures are still unknown'

(t) a) fü 45 a) a c ç t(t N o ..1 r; a) +J .-, a) ç 4t Q) 'lJ E tú '4 ôt qt q 'Fl 'Fl ç ry o 'È1 .-, .lJo o ç E (D Ìt h q o o o +J o rd (D g ,q ç o g +J' ry f o ¡r :ÌY F a) Âr E ql & Fr o U o pr q a Pr q FJ x

\\ Rrun

The first step in further study should be the establishment of the monophyly of Èhe Orthocladiinae and if possible the establishment of its sister group. Unfortunately in adults of Orthocladiinae (the best known life stage) only the Èerminalia present characters which are of any assistance. The known larvae and pupae are of little help. In the terminalia of Austrobrill-ia collessi (Fig. 15) , the sternaPo- deme has 3 processes,. the first two, one of which curves anteriorly towards the coxapodeme and the second of which extends posteriorly on the dorsal surface of the gonocoxite, probably represent connections of the primitive loth sternite. the third process tinks the sternapodeme with the phallapodeme. In Paraheptagyia ennea (nig. L2) the second, process j-s present but less heavily sclerotised. Drawings made by Hirvenoja (1973)

show that all three processes are present in Protanypus and Prodiamesa and this is also the case in many Chironominae (e'g' Schlee, 1968, Fj-9'238) Thus, since it is widespread, the fuII d'evelopment of these three processes could be considered a plesiomorphy in this branch of Chironomid'ae' In all Orthocladiinae excePt Austrobrillia the second process ís extremeJ-y small or tacking and this reduced state is probably an apomorphy. Ho\^¡ever' only a loss of sclerotisation is involved and tl:te reducecl state is found also in

some Chironominaei it is possible that reduction has occurred more than

once within orthocladiinae. The reduction of the second process can therefore be used as evidence of the monophyly of the orthocladiinae only very cautiously (Leve1 D). In the absence of a firm indication of the monophyly of the sub-family Orthocladiinae it would be unwise to make definite proposals a-bout its sister group relationships: for the Present the tentative proposal made by Brundin (see above) must suffice. Before Brundin had adopted a strictly phylogenetic approach he divided the sub-family into two Èribes, the Orthocladirri and Metrocremini (Brundin, 1956). This dívision was based on a complex of 4 characters' each showing continuous variation within the sub-family' Genera v¡ith

gracles of these four characters tending to one extreme were clesignated 189

,OrLhoc1adini' and those of the other extreme 'Metriocreminit. However many species wi;hin the genera of these tribes occupy intermediate positions and thus subsequent workers harze not widely accepted Brundin's classific¿r- tion. Brundin also noted that, while all genera of 'Orthocladinir are limnic, many 'Metriocneminir have colonized terrestrial and semiterrestrial habitats. Let us then consider the Australian genus Nasuticladius. According to Brundin,s morphological criteria it must be considered as a it member of the 'orthocladini'. Yet, although immatures remain unknown, has been shown that they occupy semiterrestrial substrates and in this respect the genus would be better considered as belonging to the Metrioc- nemini. Thus, Brundin's division, made wiÈhout strict consicleration of possible phylogeny is not completely satisfactory when applied to Australian genera.

Edwards (1931) described species of LimnoplJel from south America (recognised as Austrocladius, Freeman, 1961) with "a median reflexed hook at the base of the hypopygium." I have called this hook a penapodeme (sEcTIoN III, 3.18I). In the famil-y chironomidae the penapodeme is found onty in the sul¡-family orthocladiinae' as restricted above' within orthocladiinae it is present in aII genera of Brundin's tribe'Metriocnemini' which are represented in Australia with the exception of Parametriocnemus genera and Gvmnometriocnemus. rt is also present in arl other southern of rMetriocnemini' yet described. An examination of the literature shows that structures which might represent incipient penapodemes have also been figured in drawings of northern representatives of chaetocladius, Met¡i9s- Parakiefferiella ]]Ë' Heleniella Gvmnometriocnemus t Psilometriocnemus of and Pseudosmittia of 'Metriocnemini', and Orthocladius and Er:kieffe::ie1la ,Orthocladini' (Brundin, 1956; Saether, 1969). It seems likely' therefore' as: well that the penapodeme -is an apomorphy shared by all 'Metriocnemini' as some ,orthocladini.' Altfrough it is possible that this structure has 190 arisen more than once as a parallel response to some kind of environmental pressure resulting from Èhe invasion of semiterrestrial habitats ' its presence in many timnic species and its, absence from Podonominae (some of which ínhabit semiterrestrial environments) suggests otherwise' The presence of a penapodeme may therefore be used as evidence that the genera in which it appears form a monophyletic group (Level B). The apparent absence of a penapodeme from northern representatives of some genera of Metriocnemini may indicate plesiomorphy, or it mdy be an oversight' Thus any conception of Brundin's 'Metriocnemini'must now be altered to inclucle Likewise Nasuticladius and PerhaPs Eukiefferiella and Orthocl-adius. below' no longer 'Orthocladini', particularllz in the tight of discussion has the meaning intended by Brundin in 1956. I have therefore abandoned both tribal names. Insubsequentpageslhavefirstlydiscussedimportantcharacter genera phylogenies and then considered separately the relationships wiÈhin I or groups of genera which are represented in Australia' Followinq this intergeneric have discussed the value of the Australian fauna in clarifying relationships within orthocladiinae'

6.1 Important Cha racter Phvlogenies The order in which character transformations have occurred can some- useful in times be inferred and the resulting character phylogeny may be determining relationships in more than one group. Below I have discussed of character the justification or otherwise for the construction of a number phylogenies. of those not discussed here, but which have been used in -eub- give the sequent sections, reference to the level of relia-bility should reader an indicati.on of the basis for a particul-ar evaluation' used by schlee Many character transformation series (including some as useful (1968) , Saether (197I) and Hirvenoja (1973) ) have been dismissed indicators of phylogeny for reasons similar to those put forward in the discussion of Brundin's work (see 2 ¿¡eove) ' 19r

the eye take 6. 1r Pubescence. Setae found between the facets of on a number of configurations within chironomidae' In the orthocladid and the much smaller Pseudosrnittia setae çJenera Cricotopus and Diplocla

D iptera : Nematocera, the absence 6 -L2 Dorsal extens ion of the eve. In In many families of an,eye bridge is apparently the plesiomorph condition' are close to Chironomidae, e.9. CeraÈapogonidae, Culicidae, the extensioirs within the found quite extensively (which suggests they are plesiomorphic they have superfamily which includes chironomidae) while in other families are lacking been established as an apomorphy. In Chironomidae extensions nost orthoclad- in most Podonominae, some Tanypodinae and chironominae and plesiomorph' iinae. I have tentatively adopÈed the reniform condition as Although the 6. 13. C setae of the antennal fl llomeres. Presence pLesiomorplry' of type c setae on all flagellomeres must be considered a dipteran since this condition is found in Podonominae and' also in other groups,thecommonpossessionoft}rereducedstatemayeasilyhaveoccurred in sensitivity to on a number of occasions since they are directly involved of monophy'ly' the environment and cannot therefore be considered' as evidence character- 6.I4 Winq char acteristics. Brundin (1966) stated that wing of any specíes istics are 'plainly not adaptive" I{owever the survival dependsonsuccessfuleclosion,mating,dispersaltoandselecÈionof fr:om the suitable oviposition sites, aJ-I of which require special abilities wings. Eclosion in running waters reguires a wing which is almost imrnediate- Iyusefulandcertainlysornevenal:ionpatternsmayirlcreasetherateat l-92 which it becomes so. fncreased setae may also prevent wetting of wing and body surfaces (e.9. Gymnometriocnemus) . The flight of midges is slow and wings are small, resulting r-n a thick borrndary layer which increases with increasing hrrmidity" The possibility of stalling increases as the borrndary layer increases since separation of laminar flow from the wing surface is more likely and drag becomes greater than 1ifÈ. Voge1 (1966) reported a low tendency to stal-I in fruit fly wings and suggested bhat the presence of microtrichia and macrotrichia serves to prevent the formation of vortices in the boundary Iayer and thus reduce the risk of separation. He points out that where only a few macrotrichia are present ttrey are found in positions most useful in this regard. Thus in Ctrironomidae macrotrichia are found arouncl the perimeter of the wing and if not over aII the wing membrane at least in the distal region near the trailing edge where they are directed in the best direction to reduce drag. În many, microtrichia are enlarged in these regions and are more developed in species which have been observed swarming in high humidity. A narrow wing is less likeJ-y to stall- than a broad wing and for a given wing area, the longest wing will be more successful, i.e. if Aspect Ratio (rengrtJ-2¡area) is high, Iess anti-stafting devices wiII theoretically be necessary, but stronger and therefore heavier wing supports witt be needed. However, contrary to the report of Vogel, an exaÌnination of the species discussed here reveals no clear relationship between Aspect Ratio and the extent of wing trichiation. Vogel has also observed that during the downstroke, fruit fl-ies employ a cambered wing profile, which theoretically gives better lift than a flat surface, while in the up stroke a fl-at profile is used. No doubt similar changes occur in other Diptera and it must be supposed that the need for this kind of wing beat will have an effecÈ on the shape of the wing and placonent of veins. particular wing characteristics may thus have been gained and lost many times in response to different en'¡ironmental influences and' the 193 possession of similar wíng characteristics should therefore not be used as evidence of monoPhYIY. A possible exception is the position of the posterior fork' where this is proximal of r-m, or base of *3*4, it is usually associated with a group of other plesiomorph characters and it is in this position in fossil specimens. Ho$rever, although it is thus reasonable to say a distal pfk is apomorph, the actual degree of distalness need not represenL an ordered sequential development, since many other factors may be influencìng vein position.

6.2 Genus AUSTROBR]LLIA. As indicated above, Austrobrillia alone among Orthocladiinae retains the primitive structure of the internal sclerites of the terminalia' posterior Freeman placed it close to Brillia because the wings have a }cng fork and-similar slope of r-m, Èhe outer spurs of the rear tibiae are long, and the gonostyles aïe double. However, all theSe character states have been established as plesiomorphies and are therefore of no value in establishing relative recency of genealogical relationship' Other plesio- (saether, morphies of Austrobrillia include the shape of the tentorium L97I) , the presence of type c setae on a large number of flagellomeres, the short ninth tergite of the male, the double gonostyle (Brundin. L966' p'8O) and the retention of ttrree spermathecae in the female (Brundin' 1966' pp' 8I'

364) .

The affinities of this plesiomorph genus will- undoubtedly become possible clearer when immatures are found. For the present, a number of relationships could be proposed on the basis of adurt morphorogy, but it is uncertain which of these is correct' The diagram below illustrates the possible affi¡rities of stro- bríllia. Character 4 refers to the fact that the sternapodeme of Austrobrillia and of many Chironominae is shortened and broadened' A similar modification in Corl¡noneura and å!hto!."ia seems like1Y to be a l-94

result of the small size of these genera. Schlee (1968) has discounted the possibility that a similar condition in some apomorph Tanypodinae is meaningful. Since Chironominae have many derivative features while AustrobriLlia is a highly plesiotnorph genus, it seems more likely that Austrobrillia will form the sister group of part of the Orthocl-adiinae, or

of all thre orthocladiinae, or of Diamesini + Orthocladiinae, than of tfie

Chironominae

m apomorph plesiomorph q !

presenr absentlillïïl fI X ffi m W Ii¡U m vein M.*n t] n fl n ffii m[il Inner lobe proximal ll distal tì Spine present absent " n n m m m m 't tr tr m m ü AnaI point present rl absent " tr T fi ü m m Sternapodeme narrow tr broad 'r r )t L:R)-T t, il tl il ry un L.R. < ft t \ ¡l \ I I I I I I

6.3 Ger¡us CARDIOCLADIUS C. australi.ensis the only Australian specieg, shares with the

Canadian al-biplu¡rus the possessi-on of tarsal spines. Hor¡Iever, si¡nilar spines are found in Heptagyini, Podonominae arld Tanypodinae as wel-I as in other farnilies of Diptera - Nematocera dnd are probably plesiomorphic' 195

Descriptions available for cardiocladius species in other regions are incompleteandnousefulapomorphiescanberecognised.Thegenusis probalrlyclosertosomerrorthernçfenerathananyrepresentedinAustralia but austral,iensis provides no new clues about theSe relationships' Possibleautapomorphiesofaustraliensisareitslackofvertical of the tarsi' setae and presence of setae on the inter-segmental membrane The large group of supra-alars are probably plesiomorphies'

6.4 The Díplocladius-S tictocladius genus Pair' group. There is no doubt that these two genera form a monophyletic spine on the gonostyle' Among shared apomorphies are the lack of a terminal pubescent eyes and a reduced or lacking anal point' ThemonophylyofStictocladiusrestsonasinleapomorphy'the presenceofpatternedwings(LevelB)whiletlrederivativeformoftfie pupa(Brundin,1966,p.433)andthewelldeveÌopedi¡rnerlobeofthegono- coxiteSuggestthatDiplocladiusisalsomonophyletic(bothLevelB). lobe of In all three Australian species of stictocLadius the ventral an apomorphic condition the gonostyle is very large and bulbous, undoubtedly (Level A) The much smaller' which establishes the group as monophyletic ' species pictus but still swollen, ventral gonostyle of the New Zealand much reduced or lacking anal Freeman a]ld lacuniferus Freeman and the very these together point of the Australian and New Zealand species suggest that formamonophyleticgroup(LevelB).Presentdescriptionsaretoo incompletetoallowanysuggestionsabouttherelationshipsofthethree species described by Edvrards from South America' Aninnerlobeofthegonostyleispresentinalmostallpenbersof thisgenuspairanditsstrongreductionoraÏ¡senceintheNewZealand. speciesisprobablyanapomorphyindicatingtheirmonophyly. can be seParated from Among the Australian sPecies, multiser ialis ninth tergite of the female uniserialis a¡rd victoriensis by the extended (apomorphyLevelA)andthemultiserialdorsocentralbristles. l-96

ThissisterpairprobablyformsthesistergroupofBrilliaand'/or other Northern genera (see Fig. I83) together, but the Australian species provid.es no new indications of these affinities'

Ia.dius Stictocladius Diploc fe ilã Ø' Q) m apomorphic *f8 g/ f *r/ c Ò/ D plesiomorphic øo ql I inner lobe u n m m normal- I reaucea mil lf rl gonostyl.e lr It tI E ml mt I normal enlarged anal !¡lr¡oint ruJ mt m m m present reduced WJ-i-tÇ m m ml m run plain ,r patterned "

6.5 Genus EUKTEF¡'ERIELI,A. As suggested by Freeman (1961, p. 644) the Australian lnsil¿gg fafl-s within the brevicalcar Kieffer sub-group of Eukieif{gg-Slls' E. insolida is most similar to styliEç! Goetgh., a species reporte'J from rrerand and other parts of Europe- rn both, the tip of the maLe antenna is heavily clubbed and twisted sideways, a signifi'cant apomorphy (LevelA).Inthepupaofbothspecies,theL-setaeofthepre-anaJ. aponorptric segment are grouped together on the posterior corners, also an of condition. similarity in the ratio of length of flagellum to length- differences base of the thoracic horn of the pupa rnay be significant, but probi:bly in the number of spinerets on the horn and abdoniinal segments resurt from different environmental pressures. rt is probable that these information two species form a monophyletic pair, but there is insufficient available to suggest what position the pair occupies wi't-hin the brev:!- calcar group. any The Austral.ian species does not aid in placing the genus in t97 phylogenetic scheme. Ho\¡Ìever the possible presence of a penapodeme in

Some European species suggests that it diverged later than Cricotopus'

6.6 Genus SYNORTHOCLADIUS The single Australian representative S. brun¡eq is typically rheophile.

Few European species have been described and the horny structures of the Australian pupa are unique in this genus. s. brunnen possesses a penapo-

d.eme and, if representative of the genus, this suggests it is more apomorphic than it has been regarded previously. The presence of two tiny stout setae high over the scutum of the adult, is an apomorphy which has previously been discounted, but which is shared with genera such as Pseudosmittia. In view of the similar reductÍon of the anal lobes of the pupa in synorthocladius and Pseudo-sln:Ltt:Ls and Èhe above characters, it seems possible that this genus diverged at an early point from the line which

Ied to many of the terrestrial genera'

genera. 6.7 The P ar atri cho c I ad ius-Cr i- co groups of Hirvenoja (Lg73) proposed that the relationships between the genera Acri Paratrichocfadius and Cricotopus 4 Halocladius + Paracladius could be represented as shown by the solid lines in the diagr:am below'

o +J (n 5 a)

ô. apomorphic o m ' td CJ tr pl.esiomorphic dorsocentrals m large' r:t minute m m LI eyes hairY bare D

egg guides tt mm m close t, seParat-e together 198

He has proposed ttre monophyly of the aratrichocladius and Cricotopus group and the structures of Australian species do not conflict with this. Hov¡ever, on the basis of the absence of an internal seta on the mandijcle o f Acricotopus and Paratrichocladius he has also claimed t-o establish the monophyly of this pair. The larva of the Austra- lian specimens of P. pluriserialis are unavailable but Forsyth distinctly illustrates the pïesence of that seta in the New Zealand specimens' doul¡t' Whether these two genera do form a monophylitic group is Èhen in Hirvenoja has only tenÈatively proposed Orthocladir'ns as a sister group of this complex, so his designation of the reciprocal monophyly of this sister group on the basis of the lack of eye pubescence seems unnecessary' 6.71 Genus CricotoPus. A portion of Hirvenoja!s diagram proposi-ng relationships among

Cricotopus is shown- &s, & ln ß ]J 5 q & plesiomorphic apomorphic N

pedes sPurii e seg. III ñ NNNN I present L-l a-bsent üu Ieg sensilla PlesiomorPh. tt apomorph. u um m lateral setae t) short n n N m (larva) normal 'l spermathecal tt mn N n n duct plesiomorph. aPomorPh. ')

He has designat-ed short lateral- setae of th-e farva and a particular formation of the spermathecal duct as apomorphies of phylogenetic value and therefo:ce must have sometrow eliminated the possibilities of these arising by parallelism. However, in bottr festivellus and trifascialuq onIY 1.99

segment II has pedes spurü- B. llirvenoja has considered this apomorphic state as arisí¡rg through parallel development but that it can be used within a monophyletic group to slrow relative apomorphy. He does noi- discuss how he excludes parallelism in one case but accepts it in a second equally doubtful case. The fact that the Australian spec ies parbicinctus lacks pedes spurii

D on segrment III of the pupa suggests that this character is fairly widespread here and that it cannoÈ be used to divide up the group of genera shown.

The remaining Austratian species are only known from ad.ults and provide no characters which suggest associations different from those proposed by Hirvenoja. 6.72 Paratr:ichocl-adius The two Australian species, pluriserialis and bifinestrus share exaggerated humeral pits and heavily sclerotisecl spermathecae, both characteristics which are not forrnd in such an extreme form elsewhere and may therefore be used to establish the monophyly of the Australian pair (Level e). Bottr the Australian species are separated from the rest of the genus by their multiserial dorsocen'tral bristl-es. Since more èhan a single row of suctr bristl-es is found in ott¡er genera of the Cricotopus group, the situation in gluriserialis and bi.Einçq_tr.us suggests these are a relatively plesiomorphic pair-

6.8 Genus Microcricotopus

The Australian MicrocricotoBus clavulus is very similar to European species and shares a long thin thoracj-c horn with b-icol€r. This character state is unique and I consider it apomorphic. A comparison of 'preferred' habitat and pupal horn shape in other members of this genus re'¡ealed no relationship between them, so the common possession of the apomorphic state is evidence of monophyly (Level A). A similar comparison of the occurrence of abdominal setae in relation to habitat showed that these were clirectly r:elated: the stronger the currents inhabitecl , the more 200 setae are to be found. Thus the different aïrangements of such setae proposal of encountered in bicolor and clavulus do not contradict the and is relationship between Èhem. M. bicoror is a widespread species reported from Japan so its near relationship to clavulus is not unexpected. It is diffícult to suggest where Microcricotopus lies in relationship toothergenera.ItsharesmanycharacterswithRheocricotopusand psectrocladius but these may not represent apomorphies' In future investigations the arrangement of eye pubescence should be considered' the rete¡r- The development of swim-plates in the pupa accompanied by tion of the three anal- setae has occurred widely in the more 'plesiomorph' It orthocladids, but thís potential seems to have been lost elsewhere' tendency to develop may be reasonable to consider that a grad'ual loss of the before cricoto- suctr a plate has occurred and that MicrocricoÞppus- diverged ry from a common line of descent' 6.9 The Chaetocladius group of genera. cladius Brundin grouped Limnophyes and Paralimnophy€q with B of the over-aLl Thienemann and Chaetocl-adius (fieffer), on the basis of apomor- similarity of ttre adults rather tLran on the common possession phies.Anexaminationofthepuparevealsthatmodificationoftheanal genera as well as lobes has occurred along similar lines in the latter two two Australian genera i-n the Canadian Parachaetocladius (Wül:

This is very unusual in Orthocladiinae. It also occurs in the Australian genus Kief ferop]'ty.s-.

6.9r Genus The Australian species ttb"t.* and pelurgis have frontal tubercles, (Level an apomorphic character wtrich makes ttrem a monophyletic group e) '

In common with a nr:nÚcer of European genera they also have scale-Iike scutal setae, but since such setae could be relatively easily J-ost it seems unwise to suggest a relationship on this basis. The penapodeme is retained by one Australian species and alt souÈh American species ' Thu's these species can be considered, more plesiomorph than most of their

European corrnterparts which have secondarily lost it'

6.92 Genus Paralimnop hves

The two Australian species are unique in having a horny protrusion of the proximal end of the third palpal segmenÈ, an apomorphy which sugqests the monophyly of this pair (Leve1 A) '

6.93 Genus Nemoriclad'ius

Adults of this genus are very simitar. The only clue to thej-r relationships is in the shape of the inner lobe of the gonostyle' In both Iobitorus and diversus it is turned up dista1ly, a peculiar development (Level B): which suggests that these two form a mono'ohyletic sister pair the longer and narrovrer lobe of lqÞilo. rus suggests that this may be the

more apomorPh of the Pair.

(Dl 5l 8l mil apomorphic il T plesiomorphic inner lobe D üil short elongate m normal ), turned up ), m m 202

6.94 Genus Sparsicladius Both S. telmatus and S. graminis have an unusual proximal lobe on the gonostyle and an inner lobe wtrictr is complex compared with that of the remaining species of the genus. These are apomorphies and may tentatively be proposed as evidence of the monophyly of this pair (t,evel c). g. polysetosis and s. paucisetgsis share an enlarged styte and anal point, both of which are apomorphies and probably separate these species as a monophyletic pair. The undifferentiated nature of the bryonis terminalia suggest that it shoulcl remain outside a group comprising the remaining species.

m apomorphic (nl lJ '"t '41 trl plesiornorphic 4l8/ IJ proximal lobe on gonostYle absent present mn ru n mu inner lobe of m mn gonocoxite f, normal t, enlarged tl anal point tr tr nmm rìormaI It enlarged t, gonostyle It t, norrna.l- enlarged ú N mm

6.95 Genus KiefferoPh-vgs Kieff es phaflithrix, artiothrix and selnithrix are the only proposed Orthocladiinae known to have bent palpal sensilla and they can be as a monophyletic group (r,evel B) within Kiefferophves' K' phallithrix lobes has basally expanded type c setae, high P.R. and pubescent aedeagal which mark it as more apomorph than the other two genera' Kiefferop hves was associated with the Chaetocladius group partly menÙcers of both because of the rrnusual development of the style in some 203 genera. If this suggestion proves correct ttren K invenustulus as plesiomorph relative to istvlus and tobifer must be considered phall ithrix, artiothrix and semithri-x- AllKiefferophyesspecieshaveadisti¡¡ctiverorrndanalpointwith group (Level A) ' strong setae wtrictl, establishes them as a monophyletic

ry o

palpal sensilla straight bent m NI N tr N type C setae IN u tr tr normal enlarged

6.IO The Nasut iocnemus and the southern This grouP comPrises Metr iocnemus and Allometriocnemus. genera NaSU ticladius Austro cladius all members of which Brundin (1956) defined a'Metriocnemus group" hadwelldevelopedacrostichalbristles,longwingmacrotrichiaandalong analpoint.Noneoftlresecharacterscanbeconsideredvaluableapo- may not be monophyletic' morptries. Therefore Èhis group as it stands BrundinseparatedGymnometriocnemusfromtheMetr.iocnemus-groupsinceits position seemed ill-defined' Austro cl.adius Nasuti- Let us consider the three Australian genera In bottr Austrocladius and NasuËi- cladius and AIIometriocnemus .Freeman ' Allometri ocnemus specimen cladiustheMouthRatioishigLr'.Inthesingle Ihaveexaminedronlythecibarialpunpremainsof'Ehemouthparts'but thisissomewhatextendedand,exceptforitshairywingsandadditional pre.alarsthisspecimencouldwellbemistakenforaNasuticladius. 204

Edwards described two sPecies of Metriocnemus subrostratus and auran- ti.acus from South America. Both are almost certainly species of AIlo- metriocnemus and both have extended mouth parts' I have examined specimens of Metriocnenus from New South Wales and found tJlat they also have extended mouth parts' Extension of the mouth parts has been reported by Brundin (1966, pp. 288, 2gg) in the very plesiomorph genera Afrochlus and Archeochlus where the whore of tl- e head including ttre mouth parts has a very primitive appearance. colless & Doìl¡l-les, Lg67, have reported extended mandibula'te mouth parts in a chironomid which they suggest is a podonomid collected in lrlestern Australia. It is possible thaÈ the proboscis is a plesiomorphy group among chironomidae. However ttre form it takes in the Nasuticladius that its is somewhat separate from those discussed above and I consider appearance in thÍs group must be an apomorphic development or re-develo'oment (Level C). In support of this relationship it shoul-d be mentioned that- in all four genera palpal segTments do not progressiveJ-y increase in length from 1 to 5r but th,at 4 is shorter than or no longer than 3. There appears to be little,funcÈional correl-ation between M.R- and' the pal'pal proportions' In Archeochlus and AfrochJ-us, the pa1pal proportions are normal and thus the unusual proportions are not encorrntered in aII species with high 14'R' (t'evel c) These unusual palpal proportions are probably an apomorphy ' A further apomorphy shared by these genera is ttre tendency for the anal point to be reduced r:ntil it is no more than a ridge' The complete

absence of the point and, the indentation of the border suctr as is

encorrntered in Eukiefferiella does not occur among them" propose It seems reasonable in the light of the above evidence to different that these four genera may constitute a monophyletic group' The tlpes of wì.ng trichiation found in these four genera indicate, however,

that eacl-r has a

It is possible that the ur¡usual genus Rhinocladius is a sister group of these four: the genetic flexibitity which allowed the development of the elongate clypeus and labial palps in the latter, has perhaps been and to ) channelled in Rhinocladius to the elongation of the clypeus alone the loss of some palpal segments. It also shares the reduction of the anal point with the above four species. of the remainder of Brundin's

Metriocnemus group, onIY Parametriocnemus is represented in Australia and' this is discussed befow. It is probable that Gym4olle:q5þslenge is the sisÈer group of

Rhinoc Iadius and Austrocladius. However further conclusions must await the discovery of immatures of Austroqladiue'

(Dt 5l ãl ry() ol8l fr/ s/

can loe divirted 6- IO. I Genus Nasuticfadius. species of Nasuticladius into two groups on Èhe basis of whether there is any development of the anal point. It is possible that the large group of species J-acking an anal point - eury4gqgq, cari.natus brevipenni s, healensis glover:l and ancl niger' duckhousi - form a monophyletic group together with tolnoi'rí pattern Ho\n/ever, i., tti" group -it is impossible to detect any consistent The of apomorphies which allows further relationships to be proposed" extension singJ-e exception is that tonnoiri and niger share an exaggerated ofthemoutlrpartsandflattenedtypeCsetae,bot}rofwhichare apomorphies indicating rnonophyly (Leve1s B and A) ' A separate çtroup 206 also without an anal point, formed by four species, punctus, paraPungttls' be founcl on mucronatus and salteri, in atl of which palpal sensill-a are to but one that both segments 3 and 4, an apomorphy not encountered' elsewhere in all almost certainly indicates monophyly (Leve1 A). In addiÈion, speciesofthisgrouptheapexoftheantennaisproducedintoashort (l'evel A) of tlie point, a characteristic which supports their monophyly ' speciesshowingananalpoint,wilsonianddiplosty.l-uqhaveasimilarly lobe c'¡r tlie enlarged stYle which, in diplostvl-us is drawn up to a second (Level B) and shares d,orsal surface. Thís pair is probabJ-y monophyletic qthi which a-gain may the possession of a patterned wing with fusçius and wings in indicate monophyly (Level D) although the presence of darkened Allometriocnemus casts doubt upon ttris evaluation' genus is largely ::esÈricted' 6-10" 2 Genus Austrocladius. Variation in this prefer not l-o propose to nunr]¡ers and lengths of various setae ancl I would recognitionofmonophylyonsuchcharacÈerswhichareliabletol:ave]reen gaìned and lost on a number of occasion's' species' pectinatus 6.10.3 Genus Gymnometr iocnemus. The three Australian structure oi parapectinatus and impectinatus- share a situilar distinctive G- wilsoni shares the terminaria and certainly form a monophyletic group. America and' wittr iÈs lack of an anal point with longiçostalis from south

6-11 Genus ParametriocneJnus . group' on Parametriocnemtls was included in Brundi¡r's 'Metriocnemus and the the bases outlíned above' However, the lack of a penapodeme a swim-plate retention of the ability of the pupa of this genus to form Brundinrs rorthoclad- suggesb that its sister group wiII be found among group or thi> ini,r.ather tLran in the proposed extended Cþitetocl-ailius

Nasu.t i cladius group. 2c-'1

6.12 Genus Smittia'

Relationshipsinthislargegroupofspecieshavebeenpoorlystudied andforthese,immaturesareimportant.Sinceonlymalesareavailable relationship' for these Australian species I will refrain from proposing any

6. r3 Genus Parakiefferiella. Brundin separated ttris genus into ParakiefferieLla s.str' and parakiefferiella sub-genus Rheosmittia, on the basis of pupal characters' p. variqatus sÏrares characteristics with both sub-genera and it is not possible to determine which of these is most likely to contain sister species of Parakiefferiella' The presence of pedes spurii B in var atus is unusuar in this genus and may indicate this is an apomorphic

specres. The niche occuPied bY P. variqatus is closer to that of Krenosmittia than to P-epbleÉEeI¿gtla s. str' and this must be considered in evaluating the phylogenetic usefulness of any character'

pair. 6.L4 The Pseudosmittia and Doloplastus ThemainStructureseparaÈingthesegeneraisthelackofascutal (lnmatures are unknown') The sj-gnifica¡lce hump in adults of noloplaq.lue. ofthisstructureisnotcompletelyunderstood. Inbothgenerathereisamarkedtendencytowardselongationofthe ratio' within orthoclad- flagellomeres and a reduction of the antennal iinaethisisaveryunusualoccurrenceandisprobablyanapomorphy condition. rt indicates rather tÏ¡an a retention of the presiomorphic

monophYlY for this genus Pair' 208

6. 15 PhyIo qenetic Tree of the Aus tralian Orthocladiinae .

The phylogenetic tree provided below is a provisional synthesis of the indications mentioned above and relates çtenera represented in Australia to each other.

Chaetoclad.ius-gP'. Smittia

Doloplastus Nasuticladius 9"P. Pseu

arakief f eriell-a

Eukiefferiella Ð¡northocl+diqs

Parametriocnenus

CracotoPE Microcr:ico t-oPus @S

Stictoclaclius

Cardioc Iadius ? Àustrobrillia

? CHIRONOI4TNAE 209

SECTION V NUMERTCAL ANALYSTS

I INTRODUCT TON not been possible Using ltennig's methods cautiously' as above' it has None-the-less it to construct a complete phyjogeny of the orthoclad'iinae' guidance to future ís desirabre that a specialist should at least give some such as mine' but workers as to the broad indications obtained from a study that it would rather than introduce a rarge element of intuition, t decided to phylogeny in the be wíser in some areas to present a first approximation analysis' form of a phenetic dend'rogram, determined by numerical Ichoseapurelypheneticapproachtonumericalanalysisratblerthana phylogeneticallybiasedonebecauseapheneticanalysisismorelikelyto is thus more likely contrast with the phylogenetic approach made above and tosuggestwhereimportantdatamayhavebeenoverlooked.Moreovertlre produced by distorting effects and implications of phylogenetic dendrograms slobodchilcoff and numerical analysis have not been completely resolved' approximations to phylogeny Johnson (1973) have atternpted t'o make numerical arrive at are and, although they conclude that the groupings they 'phylogenetíc,,theysaythattheyarenotconsideringrvhetherthephylogeny producedreflectsevolutionaryhistory,butmerelythatworkerswhoclaim tobephylogeneticistsinMayr.SsenseproducesimilarclasssificaÈions. otherprogramsrwhichattempttoproduceHennig-stylsphylogeneticdendro- grans,distinguishtheapomorphicandplesiomorphicdegreesofacharacter a practice which is on the basís of the frequency with which they occur, assrunptions made by Bruldin even more questiona.ble than lhe unwarranted (1966) and Hirvenoja (1973) (see above) ' Lance ç Vlilliams (L967 The phenetic clustering schemes developed by L97L) . etseq)havebeendiscussedatlength(SibsonL}TL,Witlíamsetal' of siloson that if a Lance and williams have acknowledged the criÈicism smallpartofthedendrogramisexcisedandrerurr,adifferentpicture maybeobtained.Inthecombinatíonofdistan""*".',''eandsorting 2lo strategy used below this fault is not great'

2. MBTHODS ThecomputerprogramusedhereisbasedononewrittenbyLance& at Adelaide williams, which is available for use on the cDC 6400 computer in this university. A concise accounÈ of the general procedure employed produced is program is given in capon & Jellet (1968). The phenogram chosen was a group based on Euclidean distance and the sorting method average,(SokalandMichener,lg5S,asmodj.fiedbyLance&V{illiams). weakly dilating Lance and llilliams (I97I) criticise this conrbination as anduntikelytoproduceaclassification.However,inthiscaseaSclose species is a representatj-on as possible of the spaEial positions of these reo.uired: one in which the discontinuities are not unduly emphasised' A strongly clustering method' is not suitable' Runsweremacleonbothstandardisedand.r:nstandardiseddata.

ChoiceofCharacters.Inthestrictestsenseatrulypheneticanalysis cannotinvolvea,choice'ofcharacters.Ihaveusedtheavai]able morphologicalcharactersexcludingexhaustivecountsofnumbersofsetae, of colorË patterns' extensive measurements or characters derived from coding on those This serection was based, not on any taxonomic grounds, but t'o that of practicality and upon the need to limit the nr¡¡rücer of characters whichtheprogramcouldaccommodate.Ninety-fivecharacterswerefinally

employed and are listed separately'

3 RESULTS is set out in The phenogram produced using unstandardised data Figures I83, 184, 185, 186'

4 DISCUSSION Ihaveonlypresentedtheresultsofthecomputerrunonunstandardisecl using standardisecl data clata in detair. The dendrogram procluced by the ruir

CHARACTERS USED IN NUMERICAL ANALYSIS Foreachcharacterlistedbe]-owthecharacberstateswerearrangedÈo separate states coded for form a logical sequence and cocled. The nunrlcer of

each character appears in brackets'

Vertical setae: Size (3) Acrostichal bristles: Disrribution (5) Seriality (3) Size (3) Postorbital setae: Size (3) Scutal hu.mp: Presence (2) Setae (2) . Brow: Shape (4) tal sensilIa: Position (5) Scuturn: Fron -11". ãf lanceolate setae(3) (5) No. of other setae(2) Shape setae(3) Pubescence (3) No. of pre-alar No. of suPra-alar setae(3) of humeral (4) Cibarial Pump: Size Pits Shape of border G) ,dorsal- Pre- isternal set-ae: (3) Mouth ratio: (6) Scutellar setae: (3) Egle' Serial-itY -lffitn of seg.4 cf' 3 e 5 (3) Number (3) cf 3 E' 4 (3) r,enitfr of seg.S ' (5) oegiee of subterminal insertion Leg: Ratio of seg. 4 on seg' 3, (3)' on seg'3(ç4) (5) Rear tibia: shape áf sensitla ' (2) No. of segments with sensilla Q) Comb densitY length of to Comb shape (2) Ratio of PaIP setae on outer sPur Q) Iength of antenna (3) ' AccessorY Accessory setae on inner sPur- (3) Shape of inner sPur (2) Antenna: Relative size of inner and' outer Rario (5) (4). Degreee of aPi.cal club (5) spurs Presence of terminal b¡listle (2) broader than Tarsi: No. of segrnents (3) (4) Stt-æ. of 4th Iong Q) No. of segments with tYPe C Presence of Pulvilli setae (5) Presence of sPines Q) No. of aPical tYPe C setae (4) Shape of tYPe C setae (5) iling: (3) Aspect ratio (5) No. of ring seÈae (4) No. of segments Q) Anal lobe, shaPe Size of squamal setae (3) No. of squamal setae (5) Pronotum: vein macrotrichia (4) Size (4) Position of (3) Position of membrane macrotrichia(3) Lateral setae (3) spliÈ (3) Size of microtrichia Size of microtrichia punctures (3) (5) Dorsocen tral hristles: End of R2,.3 beginning of row (3) End of R4n5 (5) Position of (5) Seriality (3) Production of Cost:a (4) Position of Pfk (4) Size of bases (4) Presence of terminal tubercle (2) Shape of Cu1 End of An1 (3) Presence of Pattern (2) Presence of sensorY se1*ae (3) 2L2

(continued) : Abdomen: Terminalia De" y of tergal setae (5) Thickness of sternapodemd (3) Distríl¡ution of tergal setae(3) Shape of phallaPodeme (4) Sternites with lateral setae(4) Penapodeme ratio (5) medial setae (4) No. of inner lobes (2') Sternites with (4) Sternites witl: sclerotisation(3) Hairiness of inner lobes Size of inner lobes (3) (2) Terminalia: No. of gonostYles point (2) Shape of gonostYle (5) Shape of anal (3) Size of anal Point (4) Spine, terminaliÈY Hairiness of anal Point (3) Spine, size (3) Curvature of sternaPodeme (3) Corners of sternaPodeme (3) Austrobrillia longiPes Austrobrillia collessi- Cardiocladius australiensis

Cricotopus albitarsis Cricotopus hirtellus CricotoPus Phaeosomatus Cricotopus annuliventris Cricotopus Parbicinctus Èd P arannetriocnemus ornaticornis qH.

Paratrichocladius pluriserialis P (,æ P aratrichocladius bif inestrus

Smittia retracta X Doloplastus culcitella o o Doloplastus curtatus rtÞ curtata P. Doloplastus n (D Eukief feriella insolida 0, P. Doloplastus sensillus F{ H. Pseudosmittia scutel-la la clavulus Mj-crocricotoPus H æ Parakief ferielia varigatus È Stic tocla'J.ius uniserialis

S tictocl-aCius multi-serialis Limnophyes pullulus Limnophyes albibasis Nasuticladius tonnoiri Nasuticladius niger Rhinocladius tonnoiri Pseud.osnittia argilla Smittia aterrima Gymnometriocnemus wil soni Gymnometriocnemus imPectinatus H Gym-norne triocnemus Parapectinatus F. q

H @È

Austrocladius hindmarshi Austrocladius lyolobus x o Austrocl-adius tubercus o ri woodi H' Austrocla.dius 5 É o Austrocladius digiiatus È Austrocladius terjugus P'

hd Austrocl-adius kYPensis P. Austrocladius trichiatus I H æ Austrocladius numerosus ('rl Sp arsicla.dius PolYsetosis Spars icladius Paucisetosis Sparsicladius graminis Sparsicladius bryonrs Spars icladius telmatus ParalimnophYes albitibia ParalimnophYes Pullulus

Èd H. : H @ Ul

Kief ferophYes semithrix K ief ferophYes artiothrix Kief ferophYes Phallithrix ox Kief f eroPhYes cusPistYlus o invenustr'rlus rt Kief f erophYes Fr' 3 Ë Kief feroPhYes lobitorus o È Nasuticladius duckhousi E. Synortho cladius brunnen Èd t-. Nemoricladius inversus ta ; Nemoricladius diversus æ î N emoricladius lobitorus o o É rn

zp, brevipennis Ø É É rt longipennis lr' ts. rf o H punctus F, ct Þ Þi' l-. parapunctus 5þ É p, (n hd o P. healensis ÉoPrX a ootd Þf H !toH. rt o gloveri do Ol hÞ F' salteri i-O Ftt mucronatus rr o eurynotus Ð o l-' carinatus FJ l-¡' diplostylus u t-, fuscus tq o FJ 2L3 analysis.Thisweightingisnotunreasonableevenwh'enusingaphenetic approachrsinceeachcharacterstaterepresentsanevolutionarystepr particularty in characters with d'iscreet states' provides an excerrent over- r consider that the dendrogïam presented impression is of a number of view of the subfamily orthocladiinae. The closelysimilargroupsasshownbythesmall'rangeofjuncùionvaluesover which theY merge. Asexpected,thetwogeneraAustrobrilliaandCardiocladiuslieout- sidetherangeofvariabilityoftherestofthesubfamily.Theremaining diverse group (comprising species are divided into a small-, relatively Euki efferiella Do StictocLadius Parakigfferiella, Microcricotopus' and Smittia and a larger group' tus a¡rd some members of Pseuclosmittia sub-groups rthich separate off at The larger group comprises a number of the order in which this very close junction values' indicating that e.9. Rhino cfadius is onlY verY separation occurs is not very significant' than is Austroclacliu-s' slightly closer to ttre group containing Nasuticladius 2L4

SECTION VI CO}TCLUD ING REMARKS

been made, In SECTIoNS IV and V two separate sets of proposals have between and' each on a different basis, about the possible relatj-onships wíthin genera of orthocladiinae. since the phylogeneticist must use phenetic observations as his starting point some agreement is to be it must be expected between the two proposal.s. Where disagreement arises due, to a large extent, to the different weight which the phylogeneticist of attaches to some characters, on the basis of a broader background information and a consideration of the evolutionary process' HohTever' a pin- great difference between the two should allow ttre phylogeneticist to proposals point areas which may need moÏe careful attention or in which his need special justification' Thus,incompari_rrgthetwosetsofresultspresentedaboveitmustbe as well as groups remembered that data from immatures and general biology, study, but represented in ottrer areas were avaira-bre for the phytogenetic too that this data courd not be used in the phenetic study because it was incomplete. groups represented The phenetic dendrogram shows that in the two major in Australia,the Ctraetocladius and Nasuticladius groups, the constituent in finding genera are af.I closely similar and the difficulty I experienced proposals suitable adult characters on l,¡hich to base any phylogenetíc major dis- could therefore have been expected.. The phenogram shows no it índicates agreement at the level of intergeneric relationships, but of the that attention needs to be given to the position and homogeneiÈy have been as yef Pseudosmittia an

given in SECTION IV' ThepositionofSyr.rorthoctaÈLusprovidessomewhatcfaproblembecause theadultc}raracteristícspointinquiteadifferentdi.rectionfromthe pupalonesandítmustthereforeawaitmoredetailedconsideration" rvith Nemoricladius and Na suticladius duckhousi has been linked of this species to Kiefferophyes in the ptrenogram. My alrocation \æ"ti- shows it joi'ns synortho- cradius was made tentativery and the phenogram

ciadius and Nemo ricladiusatahigherJ-evelofdissimilaritythananylink be an outrj-der of the genus Nasuti- within Nasu.bicladius. rt may stitl of the remainder of thj-s genus sJsdius. excluded because the group average

was moved away from it duri-ng clustering' 2l-6

is to be The association of Microcr icotopus with Parakiefferiella pupae are quite expected on the basis of adurt characters, but the different. VfhileithasbeenshownthatHennig'smethodsofphylogenetic of the possibil- analysis have often been used with too great a disregard this was not his ity of an evolutionary reversal, I have indicated that by both Brundin intention. ì4y above discussion of proposals put forward andHirvenojahasshownhowtheyhavemisusedHennig.snethods. be understood' in The level of phylogenetic relationship which needs to the distribution order to clarify biogeographical problems associated with closely related of biota in the southern hemisphere is not that between groups of species' species of one genus but at the very lowest, that between ProposalsrelatJ.ngsinglespecieshaveonlyledtoHennig.smethods comingintopartialdisrepute,a¡dinthisthesislhavebeenconcerned relationships. more with a critical examination of higher leveI ofthegeneradiscussed,themostinterestingarethoseoft]re group' of the Na.suticladius group, followed by those of the chaetocladius and Austroc-ladius are these çtenera, Allometriocnemus Rhinocladi.us representedinbothsouttrAmericaandAustralia.Therehasbeenno and so there indication that any of tì-ese are poryphyletic in orígin, species-groups appears to be a pattern of close refationship between areas within closely rerated genera in quite separate geographicar ' Thematerialprovidedinthisthesisandespeciallytheevidencefor close.relationshipbetweendisjunctgenerashouldprovideabasisfor region' future studies of the orthocladid fauna of the entire southerD mo::e decisively the extent to such studies would be expected to indicate as claimed so which Antarctica may have formed a centre for dispersal forcibly bY Brurrdin. 2L7

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an Schlee, D., Lg66- Preparation und Ermittlung von Messwerten L69 193' Chironomidae (Diptera) ' Gewasser und Abwasser 4I/422 - t968c. Vergleichende Merkmalsanalyse zlJx Morphologie und zugleich Phylogenie der corynotleura-Gruppe (Biptera chironomidae). eineallgemeineMorphologiederChironomiden-Imago.Stuttgarter Beitr, Naturkunde lBO: I - l5O'

Serra-Tosio, B- 1970. Morphologie generale de 1b'ntenne des des unites Diamesini males (Diptera, Chironomidae) et localisation 61= I47-L63' sensorielles - Trav. Lab. Hydrobiol. Piscicult' Grenoble

a paper by Lance and Sibson, R-, L97L. Some observations on Williams- Comp. J. LAz 156*157' 223

VI' The Skuse, F.A.A-, I8B9: Diptera of Australia' Part Chironomidae. Proc. Linnean Soc' New South lfales (Ser. 2) 4z 215 - 311'

and a Slobodchikoff, c. N. and Johnson, C.D., L973. A Phenetic a Genus of Phylogenetic Approach to tl.e Classification of Zool-' 222 28O - 294' Seed Beetles (Coleoptera: Bruchidae) ' Syst'

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1963' Principles of Numerical Sokal, R. R" and Sneath' P'H'A' ' Taxonomy, Freeman, San Francisco and London'

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Thienemann,A.1936.AlpineChironomiden(ErgebnÍssevonUnter_ oberbayern) suchungen in der Gegend von Garmisch-Par1-enkirchen, Arct¡-. HYdrobiol. 30: L67 ' 262' bekannten Larven - Lg44. Bestimmungstabellen fur díe bis )eluzi- Arch' und Puppen der Orthocladiinen (Diptera: Chironomidae) ' Hydrobiol. 392 551 0 664' -Ig54.Chironomus.Leben.Verbreitungundwirtschaftliche I 83' Bedentung der Chironomiden' Binnengewasser 20: -

studies on a new Tokunaga, M. , L932' Morphological and biological Part I' marine chironomid ffv, Pon tomvia pacifica, from Japan. KYoto lmP' MorphologY and Taxonomy' Mem. ColI. Agricult. Univ.lg: 1-56' 224

r' Flight performance of Vogel, S., 1966" Flight in Drosophila' tethered flies' J' exp' BioI' 44: 567 - 578' -:196T.FlightinDrosophila.Il.Variationsinstrokeparameters (In the Press') and wing contour' J' e>ç' BioI' -Lg6TFlightinDrosophila.Ill.Aerodynamiccharacteristicsof 46¿ 43L - 443' fly wings and' wing models' J' exp' BioI '

Dale' M'B' and Clifford' H'T' L97L Vflilliams, W. T., Lance, G' N' ' ' taxonometric strategies' Controversy concerning the criteria for Comp. J. !42 162 - J-65'

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