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VOL. 116, PARTS 1 & 2 
29 MAY, 1992 


Contents 


Transactions of the 


Royal Society of South 
Australia 


Incorporated 


Shiel, R. J. & Koste, W. Rotifera from Australian inland waters VIII. Trichocercidae 
(Monogononta) - - - - - - - - - - - 
Arumugam, P. T. & Geddes, M. C. Selectivity of microcrustacean zooplankton by Golden 
Perch (Macquaria Ambigua) larvae and fry in laboratory studies 
Edmonds, S. J. & Smales, L. R. A new species of Acanthocephala from the Greenback 
Flounder, Rhombosolea tapirina Gunther, 1862 - = - - - 
Austin, A. D. & Wharton, R. A. New records of Subfamilies, Tribes and Genera of 
Braconidae (Insecta: Hymenoptera) from Australia, with description of 
seven new species - - - - - - - - - - 
Christophel, D. C., Scriven, L. J. & Greenwood, D. R. An Eocene megafossil flora from 
Nelly Creek, South Australia — - - = = = = - - 


Brief Communications: 

Nobbs, J. M. The response of soil nematodes to environmental stimulii in arid South 

AUStS id= os ope ee a a tS ee ee 

O’Callaghan, M. & Beveridge, I. Cysticerci of Jaenia hydatigena (Cestoda: Taeniidae) 
in an Entellus Langur (Presbytis entellus) 


PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS 
SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000 


Ji} 


79 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


VOL. 16, PART | 


TRANSACTIONS OF THE 
ROYAL SOCIETY OF SOUTH AUSTRALIA INC. 


CONTENTS, VOL. 116, 1992 


PARTS 1 & 2, 29 MAY, 1992 


Shiel, R. J. & Koste, W. Rotifera from Australian inland waters VIII. Trichocercidae 
(Monogononta) —- - - - - - - - - - - 
Arumugam, P. T. & Geddes, M. C. Selectivity of microcrustacean zooplankton by Golden 
Perch (Macquaria Ambigua) larvae and fry in laboratory studies - ~ 
Edmonds, S, J. & Smales, L. R. A new species of Acanthocephala from the Greenback 
Flounder, Rhombosolea tapirina Gunther, 1862 - 4 - = = 
Austin, A. D. & Wharton, R. A. New records of Subfamilies, Tribes and Genera of Braconidae 
(Insecta: Hymenoptera) from Australia, with description of seven new 
species - ee tot ee i a 
Christophel, D. C., Scriven, L. J. & Greenwood, D. R. An Eocene prigealoseal flora from 
Nelly Creek, South Australia - . = “ - = = 


Brief Communications: 


Nobbs, J. M. The response of soil nematodes to environmental stimuli in arid South 
Australia- = - - 
O’Callaghan, M. & Beveridge, I. Cysticerci of Taenia hydatigena kCesipae ‘Tasondae) in 
an Entellus Langur (Presbytis entellus) - - - - - 


29 


77 


79 


PARTS 3 & 4, 30 NOVEMBER, 1992 


Goonan, P. M., Beer, J. A., Thompson, T. B. & Suter, P. J. Wetlands of the River Murray 
flood plain, South Australia. 1. Preliminary survey of the biota and physico- 
chemistry of ten wetlands from Chowilla to Mannum.- - -~— - 


Roweit, A. I. Dispersed cuticular floras of South Australian Tertiary coalfields, part 2: 
Lochiel - Oh <a ee: _ 3 st? hy 


Beveridge, I. & Durette-Desset, M.-C, The morphology of Nippostrongylus magrus, a parasite 
of native Australian rodents - - 


Davies, M. Early development of Limnodynastes terraereginae and L. fletcheri (Anura: 
Leptodactylidae: Limnodynastinae) - - - - - - - 
Beveridge, I. & Durette-Desset, M.-C. A new species of trichostrongyloid nematode, Odilia 
bainae, from a native rodent, Rattus fuscipes (Waterhouse) - - = 


Brief Communications: 


Green, J. D. & Shiel, R. J. A dissection method for determining the gut contents of calanoid 
copepods 2 -, &£§ + + o Wo Hee! #3, Ue - 
Boulton, A. J. “Rollers” and “Carriers”: Field observations of carrion remoyal by trogid beetles 
(Omorgus strzeleckensis) in arid north-eastern South Australia - - 


Davies, M. & Watson, G. F, Redefinition of Uperoleia litlejohni Davies, McDonald & Corben 
(Anura: Leptodactylidae: Myobatrachinae) - ss - - - — 


Greenslade, P. New records of Mesaphorura (Collembola: Onychiuridae, Tullbergiinae) species 
from Australia, Macquarie Island and the Antarctic -  - - = 


Read, J. L. Ecological and biological notes on the rare plant Hemichroa mesembryanthema 
F. Muell (Armaranthaceae) - - - - - - - - - 


Morley, T. P. Eggs and incubation in the Australian lizards Amphibolurus nobbi and 
Eremiascincus richardsoni— - : : : - = - - - 
Johnston, G. R. Relictual population of Tiliqua scincoides (Sauria: Scincidae) in north-western 
South Australia - - - - - - - - - - - 
Edmonds, S. J. A note on Phascolosoma turnerae Rice (Sipuncula) - - - - - 


Tyler, M. J., Aslin, F. W. & Bryars, S. Early Holocene frogs from the Tantanoola Cave, 
South Australia - - * poe gee CH -> eh 


Insert tp Transactions of the Roval Suctety af South Australia, Val. Hb, parts 3 & 4, 30 Neweimber, 1992 


ROTIFERA FROM AUSTRALIAN INLAND WATERS 
VIII. TRICHOCERCIDAE (MONOGONONTA) 


BY R. J. SHIEL* & W. KOSTEF 


Summary 


Diagnostic keys are given to the genera and species of the Australian representatives of the 
Rotifera: Monogononta in the family Trichocercidae (Ascomorphella (1 sp.), Elosa (1 sp.) and 
Trichocerca (43 spp.)). All species know from Australian waters are described and figured. 
Distribution data and ecological information also are given. 

KEY WORDS Rotifera, Australia, taxonomic revision, Trichocercidae, Ascomorphella, Elosa, 
Trichocerca 


> 


Tmiasdenoiiy af ite Raval Adverse ef SAME IYO HUE 1-27 


KOTIFERA FROM AUSTRALIAN INLAND WATERS 
VUI, TRICHOCERCIDAE (MONOGONONTA) 


by R. J, Sutei* & W, KosTEty 


Summary 


SHEL, ROT & Kosre., W992) Rotilers froay Australia inland waters VIEL Triehocercidae (Monogondntay 


Trans. Ro Soc. So Aust, tbl), 1-27, 29 May 1992, 


Diaunostic keys are given to the genera and species ot the Australian represeatanves of the Rotitera, Monoganont 
wn the family Trichneercidue (Ascomerphella U1 sp.), Bloxe (lL sp, und Trichocerca (443 xpp.yr. Al) species known 
from Australian waters are described and figured, Distribution data and ecological information also are given, 


Rey Worns. Rodlera. Australia. taxonomic revision. Trochocercdae, Ascomerphella, Blase, Vricheeewrea 


Introducticn 


There are sporadic systematic references to the 
oecurrence of trichocercid rotiters im Australia (ef, 
Sluel & Kuste 1979), however a review comparable to 
that af Jennings (1903) for North America, or included 
it Koste (1978) far Europe. is lacking. A thorough 
global revision of the family using modern techniques 
(c.g. SEM) is desirable, particularly in view of recent 
evidence ol species-specificity in. rotifer trophi (e.g, 
Markevitch & Kulikava 1990), 

Trichocercid rotifers are a.common component of 
plankton and littoral microfauna! conimunitics in most 
Australian fresh waters. “The family ineludes three 
genera’ Elose, recorded from N.S.W. (Murray 1913b); 
Axycomorphella (A. valvecicola often occurs us an 
inhabitant Of Kalvey colonies in reservoirs (sec Gant 
eral, 83) and Thighaverea, Trichocerca is the mast 
diverse roller genus known from Australia (46 taxa 
recorded). Some species may be found in limnu- and 
river plankton, however they reach thelr greatest 
diversity und abundance tn littoral (vegetated) margins, 
especmilly in billibongs. Up to eight species nay 
coexist in hillabongs of the River Murray. where their 
Morphological and/or behavioural adaptations permit 
effective resource partitioning (Tan & Shicl in press}. 

This paper follows the format of earlier parts (listed 
i Koste & Shie) 1990) to review the present status of 
the family in Austratia, including available ecological 
joformation, Where type locality information was not 
avilable to us, the probable country of origin of the 
material is given in parentheses. Very little holotype 
oxiterial hay been lodged for the Rotifera in general. 


* Murny-Darling Frestiwiter Research Centre, POL Box 921. 
Albury, §.SAV., 2640 (to whom reprint requests should 
he addressed) 

1 Lodwij-Brill Strasse 5, Quukenbrliwk. D 4570, Federal 
Republic ot Germany. 


Methoey of studying Trichocercidae 

Taxonomically significant features of trichocercid 
rotifers are detailed in Fig. 1. Preserved (contracted) 
individuals generally can be identified, however 
contraction of protruding anterior spines, denticles or 
folds may vary between individuals. ‘To observe palpiar 
organs and sensors on the corona. living specimens 
are preferable. Trophi examination also is inportant 
in Species determination (Fig. 1b, ¢), For example 
Trichacerca parcellus and To mivsentas have similar 
inorphology.. but differ iw trophi, The position af the 
lateral anteniae at the from of the striated area is 
significant. By the addition of Eau de Javelle (KOHCI! 
or sodiuny hypochlorite (NaOCl!) the animal is spread 
our and. the position of the lateral antenni is 
momentanty clearer; rapid observation is necessary. 
[important in trophi analysis, are the tanubria 
(particularly terminal morphology), direction of the 
alulac. auoiber of teeth Of une) und tart. Bacessive 
exposure to hypochlorite destroys the trophy — it can 
be neutralised by dilute ucetiv acid. Toe and body 
lengths should be measured, Substyli at the toe bases 
ure sometimes stuck together by exerttions froni foar 
glands: in view of upparent varwobilily in number this 
is unimportant for identification, 

Two subgenera of Trichocerca are distinguished on 
toe morphology. 7. (Diurella) Bory de St Vincent has 
toes of Similar length or right at least ¥4 the length of 
Jeft; Tl (s. sf) Lamarck has dissimilar length toes, 
tight never mote than ‘4s the length of left. We buve 
combined the subgenera in w single dicholimous kee. 
but for convenience in comparing figured morphology, 
the two subgencra arc treated separately m= the 
systematic section, ‘To date 43 species of Trichocerce 
have been identified from Australia, most from littoral 
vegetation in billabongs oc in (he open water of 
billabongs or lakes and rivers as incursion species from 
marginal vegetation, 7) stmi/ix occurs commonly in the 
plankton of Murray-Darling reservoirs ind rivers (Shel 
eral, 1982), 


tJ 


R, J. SHIEL & W. KOSTE 


TRICHOCERCID ROTIFERS 3 


Abbreviations axed in systematic section: 

TL, = total length, BL = body length; LT/RT = 
Jett or right (oe; TR = trophi length: F = fulerum; 
LM/RM = lett or right manubtium: LR/RR = left 
or right ramus: RE/SE = resting or subitancous egy. 


Family Trichocercidae Remane, 1933 


Body ovaid tu barrel-shaped (Elia, 
Ascemorphella), cylindrical, spindle or sack-shaped 
(Trichocerca), often asymunetrical as a tesult Of tarsion 
ofthe body. foot present, short or absent; toes curved 
br straight. shon or long. setaeJike. rarely the same 
length: usually left toe Jonger: toes may be 
rudimentary: te(s) Tray have substyli; teaphi of virgate 
lype. asymmetrical: corona resembles Netonmatta type 
lef, Koste de Shie) 1991). Nomenctature follows Koste 
(1978), 


Key to penera 


I, Foul present. ties brstt-hke. Otten of considerable 
{1 | Hichocerca. Lamarck 
Fiat absentee 2 


J heer bs ime aes AscomorpaAelia Wisniewski 
Culwle sail qwes mdimentary or absent: Cerebral and 
apical tyespots. . Elisa Lord 


Avcomorphella Wisniewski 
Ascomorphette Wistewski. 1954, py 440 
(ype: Heriwigia volvoecicoly Plate, 1886. Monuty pic 
penus. 


Asomerphella vulvocicula (Plate) 
PIGS 2). 3 
Hernvtuin valvacivay Mate, 1880, p. 26, Big 1748. 
Ascomorphela valvacivuta (Plate) Wisniewski. 1953, 


i 340. ; . 

\pe lecalii: (Germany), 

Holonpe: Now desigiated, 

Deyeriptions Squat becrel-shaped body: corona 
Asplanchad wpe. apieal Geld with fingerlike palps 
heeween longer cilia: wide lips ventrally between which 
(rophi can be extruded: suture separates head from 
trunk, dunk With Jour longitudinal striae dorsally, two 
ventrally, short diverging strime between both sets: 
Paired lateral antennae single dorsal-antenna; abdomen 
with sniall terminal bulge, two foot segments. (one in 


Juveniles), with two minute toes: dorsally, small tail 


overhangs anus: Lrophe virgatc. of Teichocerca type, 
fulerum long: LR more robust, with pointed, 
somewhat elevated alulas subune! daggerhike; unct with 
three pointed weth, comb-toothed oral plate in frou 
ofunct: single medial red cerebral eye: trunspitrent 
retrocerebral suc: mastax with two asymmietric sylivary 
glands; unciliated gut: gastric glands Jarge, vitellanum 
with six nucleii; REs brownish with short. thick, 
slightly curved bristles; male with wnvaginated dorsal 
unlenna, everted penis, large brain and cerebral eve. 
Total length 20-160;m: width to 75m; trophi 32.0 
(LM 24m, RM l8um, F Isp, rami l4jum): mute 
‘hOpm. RE foam Jong. 
Distrifndiqa: Known trom Europe, N. America aod 
NZ. Obligate parasite of colonial algae (Olver 
Uroalena), Widespread i eastern Australia, reaches 
high densities during seasonal blaoms of belpex in 
reservoirs and billabongs (ct, Gant ef ef, 1983), Female 
ely cells OF colony front the inside, leaving distinetive 
damage (Fig. 3). lays eges oiide colony, 
Literatures Kaste (1978). 


Elosa Lord 
Elasee Lord. 48), py 423. 

Byer Elose worntti Lord, 189), p. 323. 

Two taxi are referred (o lose by Koste (978: 40Y), 
F typ. and vat, sprrtifene (Wisniewski), Tt is not evident 
that differences are more than ecotypic Variation. As 
‘var is without formal laxononmic status on present 
evidence. Elosu appeurs to be monotypic, 


Eloxa weralli Lord 
FIG. 2:2 
Flasa wardlii Lord, WO, 324, Fig, 19. 

‘ype locality: (London?) 

Holutype, Not designated. 

Deseriprion; Dorsally: rounded to domed, ventrally 
wider and Ilat; body broadly elongate oviform,; head 
demarcated by transverse suture; lorica smooth, caudal 
short spines may be plesent: on veniral posterior a 
semicircular or elliptical aperture with foot and we 
rudiments; corona with long eihat 1-2 pulps in apical 
field) Wophi axymmetric. virgate, with distally spatulate 
fulcrum; RM rod-shaped, LM a rod-or croaked; LR 
with horizontal aula, larger than right, suprarami 
present, yasiric glands relatively sroall: stomach und 
intestine indistinctly separated; gut cenerally orgnge 


Pig, l. Morphological tcarecs of trichoverciid] rotiters and theiv tophi. 1, Trichocerea bi 


wens (Lucks): (a) lateral, swimming: 


(pa pilpar organ: da dorsal antenna: ey eve: re rerrocerebral organ, tr trophus: $1 striated field: f foot: t toe}, Cb) teaphus. 
Jord, (ce) traphos, ventral (kd subuncus; tl lett uneus: ru righl WNCUS, 6 RAMs. Shas Supraraimas, fu fulerume: tr lett 
mavubeum: rm right manubrivm: al alule) 2.7 evindria (imboth: (a) lateral aly kuteral antenna. av anastux: di tHweatiye 


tract: ¥ vitelliriuiny) 12 fool glangls, ss substylen (by subhneuus eve, 


Sedle lines adult San. Gopht and ewes (jam 


(ch imile eges (dp resting vey, After Kose (YQ 71 


R. J. SHIEL & W. KOSTE. 


> 


Fig, 2 1. dscmmorphella welvocicnla Wiseniewskis li) fitteraly (bh) dooal; de) head woh trophus everted; (d) trophus, 2, 
Flose worralli Lord: ta) darsal: (b) laterals (Gy trophus, | utter Wulfert (960): 2 aflee Voigr (904). Seale lings: adult 
Sem, weypht lye, 


TRICHOCERCID. ROTIFERS 5 


Fig. 3) Coluny of to/vay showing resident dscomerphella 
velvacicola and extensive damage to cells: 


coloured; yilellarium wilh eight (102) nuclei: bladder 
small: cerebral eyespot to left of brain, secand eye 
displaced to right, lies in ciliary field, near “brow 
border, sometimes lost in separate pigment granules. 
Total Jength 80-l00pm (contracted 7Oum): width 
40m: trophi 30am (F 26um_ manubria 25m, rami 
Rum). 
Distribution: In wet Sphagnum, Reported from Europe, 
N, America, Australia and N.Z, Not seen in our 
collections. 


Literature: Murray (1913b); Russell (1960). 


Trichaverce Lamarck 
Trichnwerca Lamarck TOL: 394. 
Type. Trichacerca rattus (Miller) = Trichada rats 
(Miller) 1776, p. 281. 


Body elongate or squat, more or less curved, in many 
species somewhat spiraled, or bent from left (right side 
concave, left side convex), anterior end of lorica with 
multiple spines, denticles or tolds (particularly 
conspicuous afler contraction of more or less firmly 
loricale animals); posterior to these may be striated 
area associated with torsion of body, followed by keel 
or two ridges; caudally, abdomen projects further on 
left than right, footis inserted obliquely; rudimentary 
RT lies dorsally, LT often well developed ventrally: 
eye. brain and lateral antennae also asymmetrically 
placed; elongated mastax strongly asymetrical: rami 
have complex alitlae; subunei have been described in 
different species; LM always more strongly developed 
(terminal shape essential for identification): RM mostly 
rodlike. Salivary glands and retrocerebral organ are 
desenibed; intestine generally clearly separate from 
stomach, Excretory system has a few flame cells and 
protonephridial bladder, which can readily be confused 
with reservoir of foot glands: ocelli in living animals 
distinct on orat end of brain; dorsal antennae with short 
sensory setae (except in 7) evlindrica). Lateral antennae 
either at same height in last '4 of abdomen, or placed 


very asymmetrically, 7 cylindrica has been observed 
in gelatinous sheath. This pelagic species carries 
subitaneous and male eggs (both smooth shelled; 
Fig 1: 2a, b) at end of the abdomen, REs of 7 
cylindrica have blister-like blunt projections. of outer 
shell (Pig. 1) 2c). There is little ayailuble information 
on biology of the different species. They appear to be 
adapted to spectilized niches or their preferred habitat 
Planktonte species 7 capucina and 1. eylindriea suck 
out the contents of eggs of planktome rotilers, eg. 
Brachiomus and Keratella, Littoral taxa extract contents 
from algal cells, e.g. 70 lungivete. common in 
billabonys, breaks filaments of chlorophyceae (ep, 
Spirogyra) using its dorsal spine and sucks oul 
cytoplasm contents: TL yimilix grandis takes whole 
coccoid chlorophytes. e.g, Gloeocystis; TL. btdeny has 
developed a specialized pharyngeal basket to suck 
contents [rom desmids (see Pourriot 1970 tor other 
feeding specialties), 


Key to species of Trichocerca known trom Austealia 


I, Toes of similurlengih, or RT ar least “4 lengeh of | a 
LT (Ditrellas.. 

RT? ‘considerably redueed. alws ys <4 length of oe 

Ty (s. str. 


pd Lorica anterior margin witht projections. . 3 
Cusps, spines or other projections present... 4 
3. Anierior lateral tongue shaped plate presenr. . 
YT vernalis Hauer (Pig. 8: 5) 
Lateral plane absent. . 
P Re SS bllaris (Rousselet), Wi iy 4:3) 
4 Anterior margin with lateral tangs: atunpest Plate 


> tae 


Lateral plate absent. 
Dy Margin with stubby projec tions. no spiniets). ' 
TU sulecua dennines (Pig. 8) 
Margin with one leone spine. -- 
T webert Jonmings (Fig, 8:6) 
6, Margin with blunt projections, or blunt proyectients 
with Lwo dissimilar length spines . ani OR 
No blunt projections: one ar two spines or Usps 
7. Margin with blunt projections only... 0.6.4, 8 
Blunt projections, 2 dissimilar length spines, 8-0 


serrations - T rousseleti (Voig)) (Fig. 71) 
K, Dorsal keel present. ; m0 
No dorsal keel, y 
4, Toes curved ventrally: LM with distinct sinvle bend 


-1 inermis (Linder) (Fig. 5:2) 
Toes not curved ventrally; LM weakly curved 
terminally, To rautnert (Bonner) (Fig. 7:2) 
10, Toes curved ventrally: LM with double crook —. 
Tues follow body axis (necasianally lphtly curved 
terminally); ILM. only weukly bent, 
.. 2 dixon-ralli Jennings (Fig. 464) 
Ih. Lower. third of body conspicuously narrower, left 
uncus with several teeth / 
7 breve ‘Avure iGoskey (Fig, rt 1) 
Boily not conspicuously narrower) left uncus with 
single woth. T cavia (Gosse) (Fig, 4:2) 
12. Lorica anterior margin with single spine... 13 
Lorica anterior margin with two spines... -.17 


2\). 


i 
wi 


R. J. SHIEL & W. KOSTE 


Spine short (<tdyam) ee a, l4 
Spine long (l4-26n0) 
TE mmemnaia (Voigt) (Fy #4) 
Body long und slender. conspicuously narrower mn 
lower third; LM wath single or double bend, 
Body not constricted posteryorly, 1M weakly san 
TT tenitior (Gosse) (Fig. 852) 
BL> 130 toes > S0nM, 
7 tigrts (Muller) (Fig, $3) 
BL YO- (OB. locos < aSum ...- |6 
LM with double bend) left uncus single ‘toothed 
T intirmiedii (Stenroos) (Big 6:1) 
LM weakly benl; left uncus with several teeth 
7. insvlana (Hauer) (Fig. 5:4) 
Two Aapicrilar \ength spines/cusps 
Two similitr lent) spines/cusps- at ey eT 
Body squat posterior lorica overhangs, foot. . 9 
Rody long, posterior lorica does not vaverhang, ee 
reese} 


LT 434 Aiyam: RT 36- Sum: TR > 50m 
me enn a eee ws T. porcellus (Gosse) (Fig, 6: 4) 
LY < 43jin; RV < 45uny, TR < SOn.. 
VT omusculies Hauer (Fig, 6 2) 
Boxy canspicuously constricted in lower 4; anterior 
spines differen lengun 7? niyers! (Hauer) (Fi ig. 0:3) 
Body not constricted: spines similar length 
T. insignis (Herrick) (Fig. 5 
Single drs keel: posterior lotica projects aver joa 
OT hidens (Lucks) (Fig. 14) 
Double keel; posterior margin does nol project aver 
fief ate: 2 LT similis (Wierzejskiy (Fig, 7:3) 
Lorca wnlerior margin without projechons. ..23 
Anterior margin with blunt pIPHSChER, spines or 
CUSPS . 28 
Single dorsal Keel present, recs ate zk 
Double dorsal keel os. cere eee 25 
1M swith single crook: alula of LR aneled about 45° 
from TR axis. _T ratius (Miiller) (Fig. (1) 
LM with double crook: alula of LR angled > 45° 
from TR axis... flagellara Wuuer (Fig. 10:4) 
Body tapers in posterior 4: foes curved ventrally26 
Body slender or squat. not constricted Peatliticnty: 
toes follow body aus. a. eee ee 
BL > WOunG LT > > Kum. 
Aa oat JT! bieristata (Gosse) (iy, 9:2) 
BL < 10am: LT < 50pm. . tion 
wT _Mmucuse (Stukex) “(Fip. 122) 
0 
wo. ths ait (Goxse) (Fig, Wh 3) 
AL = 1am, Lt > 150m (exceeds BL). , 
rie. - T! braziltensiy (Murray) (Pig, 9:3) 
Lorici uterior margin with spine(s}  ---- am) 
Lorich marein with blunt EPH my pn 
| 


BL > > Mum: LT 


Dorsal keel present. . as 
i eee gracilis (Tessin) (Fig eh 
Dorval ‘keel absent. ee, waetcs cee ve ree A 
TL > [80em; BL > 135urr. . ori 3d 
Th © (On BL < (3sym. velek UNOPE HG ee o 
Posterior lorwa overhangs foot, LP > Fpm—- 
LD agnota Wullert (Big. 9: 1) 
Posterior lortca without overhang; LT < Tyan. - 
_T. stylata (Gosse) (Fig. 13> 5) 
LT > BL tad 072-083) ee 
_T. may Hauer Fig. (2 3) 
LT < ‘BL (rani 1X), “i 
ts r pusilla Jennings (Fig. ‘12: 4) 
Anterior margin wah single sping. 2... , ---34 
Anterior margin wilh (wo spines Li ay 


34. TL > 450pny; anterior spine tong... 
T cylindrica (imhot), (Pi 
TL < 450um: unerior spine short... 6). 3S 
45, Dorsal postenor Jorica projects over foot: RT 
> 30am fright LT patio < 3.3)... 
LF jenningst Voigt (Fig, 1:3) 
Pasterinr lirica does not overhang foot. RT < 30nm 
(ratio > 5,5) : 36 
Bi) TL > JQ, BL > » 20) kin; Lr > ‘OOpm-. 
Lo omewera (Gosse) (Fig 122 \) 
TL = 3400m: BL <200pm, LT <loOum—- 
..---T. ternis (Gosse) (Tie. Mh: 1) 
a7 Twn ilissiinilar lenetl spines. 
Two spines and dorsal cowl-like siructure. 
Po capucina Wrerzeyski & Zacharias (Fig. 8:4) 
38. Body constfieted in postenar '4: posterior wverhange 
foot, LM weukly curved terminally, ——- 
aD hosea (Stenroos) (hig, ‘13: aa) 
Body not constricted; no overhang, LM with single 
crook, JT longivena (Schrank) (Pig. Udy 


2) 
35 


Trichocerca bidens O.ucks) 
FIG. |: } 

Diuwrella hidens Lucks, (912, p. 66, Fig, 12-13, 

Trichacerea hidens. Ablstronmy 1938, p. 105, Fig. 9 8-9. 

Ape locality. Germany. 
folorpe: Not designated. 
Description: T bidens has (wheo contracted) two sharp 
similar length transversely striated cusps on dorsal 
lorica margin, with striated area beneath, bul no keel; 
may be three transverse folds in neck region; dorsal 
antenna in muddle of head; lateral antennae at same 
height on posterior ‘4 abdomen; al buse of similar 
length toes are conspicuous rather long substyli; toes 
often curved ventrally at tips, or sigmoid, Mastax with 
dorsal and ventral salivary glands, ganglion with long 
relrocerebral organs and two subcerebral glands; TR: 
rami-and ined multi-toothed at tips. Larger than, but 
often confused with T cave (Fig. 4:2). See alsa 
T callaris (Fig. 4:3). 

TL 220-240¢m (Swimming), lorica length 

175-205pm: height 65-80pm: toes 52-G6nm: TR 
65m) (F Slum; manubria 42/3)). 
Distribution: Cosmopolitan, isolated finds in. acid 
waters particularly Sphagnum Rare, Tas, Vie. 
13,0-27.0°C, pH 5.4-7.5, DO 9.2 mgt!, <10 NTU. 
Literature; Kosle (1978), Berzins (1982), Koste ef cal. 
(1988), 


Trichecerta brachyura (Gosse| 
FIG. 4:1 

Monorerca brachvura Cosse, ISL. p. 199, 

Trichicerca brachyuras Myers (9397, p. 6. 
Type locality: England. 
Holotype: Not designated, 
Description; Body squat, robust; on contraction. 
unterior Margin has stumpy projections on left side. 
folds on right; toes of similar length or only slightly 
Jifferent, right lateral antenna notably further to rear, 


TRICHOCERCID ROTIFERS 7 


left antenna approximately midway beiween it and 
dorsal antenna; TR with suprarami:; LM with crook. 
Similariues with T cava (Big. 4:2). T dixon-nuttalli 
(Fig. 4:4), 2 porcvellis (Pig, 6:4) and 7, pernafis (Fig. 
8:5). 

BL 73-ll2ym; toes 23-30/20-23nm: TR to 36am 
(in a 33pm TR, LM 264m; RM I2ym: F 26~n1). 
Dishibution: Cosmopolitan, generally solitary in 
psammon and littoral of most freshwatérs: pH tolerant. 
Uncommon; probably pancontinental, but not yet 


Wo 
v2 


al rat ya 
Sia) 


“att 
Bee 


ZT 


recorded from S.A, Oceurs in Myriophython in River 
Murray billabongs. 10.0 25.0°C, pH 5,76-7,5, 
43.5-218.0u8 em! 

Literature: Koste (1978, 1981). Koste ef al, (19831, Koste 
& Shiel (1987). 


Trichocerca eavia (Hudson & Gosse) 
TG 4:2 
Coelopi, cavia Hudson & Gosse, 1886, p, 69. Fig, 49.27, 
Iriehaverca cuvfa) Myers 1937, p. 6. 


Fur. 4. 1, Trichocerca lruchyure (Gosse): (a) lateral; (b) (rophus. 2, To cavia (Fludson & Gosse}: (a) lateral: (b) lrophus, 
3, 1. colluriy Rousselet: (a) lateral, swimming: (b) trophus, lateral, with pharyayeal busket. 4, 7 dixon-nuriadly Senmingys): 
(a) lateral; (b) trophus. 1 after Koste & Poltz (1984); 2-4 after Koste (1978), various authors, Seale ines; adulr 30m, 


trophi 10am 


ai R 


Tre foealiny © 
England. 
Holotype’ Nov designated, 
Deyeription: Body uf eontriteted animal plump, alrmst 
avo. Othe aspect ul i squalling guinea-pig” — 
PL H. Gosse if Hudsut) & Gosse [R8A), Without keel, 
loricn untertwor margin variably sqaooth and plated (in 
most cases); seen laterally, these vive impression of 
spines. Jarger on right. dorsally.as pointed projection, 
reduced onlall; foot conform, small, offset from wide 
abdomen, lateral antennae at same level: toes of similar 
length, usually crossed, Resernbles 7) bideviy (Pig, 1:1), 
BL 97-132am: TL 30-4lam: TR length 42-480. 
Distribution; Cosmopahtan, m petiphytar in plants wt 
standing waters, pH 5-10, S5-IQ0°C (Kaste 1978). 
Rare, ore record each trom NvT.. Tus... View 85-10, 
pil AN-20, DO ILO mel!. 7pS em', <1 NTU, 
Leerure. Koste (1978). Koste & Shiel W980, 1987). 


fear Sparesbrook, Epping Forest" 


trichacerva viltariy CRousseles | 
FIG 4:2 

Ramdas collariy Rousseler, 1k9¢. p. 266, Fig. |b 

Irivhoverce Collariy Myets W397, po# 
Type focali: England. 
Hololype, Not designated. 
Deseriprion: Lorica smooth to stippled with cansyerse 
bulges un neck region. contracted, head projects ut 
acute angle, somewhat tonzue-shiped; no dorsal 
rdpes, bul a constucterd area, dorsal anlennae between 
neck folds in normal position: latern! antennae ar 
approximately same height: UR with crooked fulerua, 
winglike saprarani ever rami; pharyngeal hasket 
anérior to rami apices, wo hinge palpar organs, 
Animal can bend sharply forward to poke the we Lips 
inte the mouth ares. 

BL. 204-309n01, lex 20-3 an, TR 80-85am (F 
Stam, 1.M 63ym). 
Distribution: Cosmopolitan in weird waters with 
Sphagnum, eats diatoms and desmids which wee not 
swallowed cotirely but Tripmented and sucked oat. 
Rare. several. records from) ‘Fasmania, only one from 
the tnainland (Magela Ck. NT) 22.5.24 5°C, pH 
§.2-§3, DO S8ing (', 3239aS ool, <i NTU, 
Literature: Koste (972), Koste & Shiel (1980, 1987), 


Triehocerca dixon mutallt Uernnings) 
FIG 4-4 

Onuirelte dopenucall’ dennings, (903, p. 318, Big. 4:40 44, 

Tetclineni din antalli, Downes (50, py ER Fig 19 
pe lecalitys Nouspecified .. . “cormmon in ponds 
in’ Engh” 
Holotype: Not designated 
Des rpion’ Body cylindrical, tapers posterwurly, head 
Sheath separited by bransverse constriction wilh sever 
longitudinal folds: shore dorsal furrows (containing 
dorsal antennac) correspond to swiated area of other 


1 SHIFT & W KOSTER 


species; corona wilh dorsal palpar orgun and several 
blunt protrusions: lateral antennae asynimetrical in 
posterion. Jef further torward than right: LT about halt 
BL. RT % length of Jett: TR asymmetrical: right 
malleus much redtuced; left uneus 4-toothed; eyespat 
at postecior end of brain: resembles 7) Arachiiera (Fig 
Ai). 1. rathiert (Fig. 7:2), 7 pusilla (Fig. 12:41 and 
T. stykaa (Fig, 13:5), Evidently most clusely relates 
to To preville, 

TL. lWa-kopiny BL. 90-122an, LT 41-S0um: RP 
27-284! TR 30-32ym. 
Distribution, Probably cosmopolitan an Hoodplains, 
inundation zones, periphyton. tyehoplankton, Recorded 
from Qld (Russell 961). In Jan, 989 populations were 
found in several small pools (Solomon's Jewels) and 
Lake Loune inte Walls of Jerusalem National Park. 
Tasmania. 230°C, pH 6.88-6,99, 
Comme: Conspeatieny of Techecercu inenms 
(Linder, 1904) and 7 diven-nutiall’ has been debated , 
e.g. Koste (1978). however Hauer (Y31) was convinced 
on the basis of European tinds of the Jormer which 
conformed 10 Linders deseripnony umd hot to those 
of Jennings, that two distinct taxa were involved, T. 
thers bas been distinguished to date only by a shorter 
Heht tov. 
Literamres Koste et al, (1988), 


Trichocercd evodonta (Hauer) 
HIG $i) 

Hivrella eaodwona Vauer, 1937. pp: 377.8. Pip, 25a d. 

Trichacerea enedonta: Koste 1978p. A08. (=Triehaceten 
enindinta, Berns W2. p. 7) 
Type locality; North Sumatra, hidenesia. 
Holotype: Not designated 
Desoriptiog; Body almost cylindrical. ca. 4 longer 
than wide. symmetrically curved. rotated about 90°; 
strong spine on right side of heud, teft of which a 
second, left-curving, wider spine, about SU% ity length, 
is deflected ventrally, a shovel-shaped protrusion. of 
the head macgan arises ket of the head opening, 
separated from tell anterior spine hy uv deep notch: 
single distinct fold on underside of head) well- 
Weveloped Keel slants across back from) baxe of anterior 
spine; ending al base oF foot; head eleariy demarcuted 
from rest of body only on underside: fher ties in 


direction of body axis, almost as long as Wide. ties 


ot dissimilar length, weakly curved, widely seperate. 
RT slightly longer than 44 LT; two stylets a1 base of 
each loc: TR asymmetric; manubria tod shaped 
TL 3pm; LT 62m; RT 37pm, 

Disreiburtor: Known ooly trom North Sumatra and one 
unconticmed Australian ceccird trom ihe Miorahool 
River a Ballan, Vie. (Berzins 1982), 

Comments: Wauec’s figures and desenptions itre 
reproduced here, The trophi were net figured in che 
original description. Whether this species ar 7! (D)- 
avers? Chig. 8:2), winch de resembles and which is 


TRICHOCERCID ROTIFERS. ) 


Fig. 3. 1, Trichwcerca ewodanta (Hawer)| (a) lateral, contracted: (b) head from right side: (¢) head trom left side; (d) foot 
und wes, 2, 7 inernis (Linder): (a) swimming, lateral; (b) toc; (c) lateral, contracted. 3, 7. insignis (Herrick): (a) lateral, 
contracted, (b-c) twophus: 4, 7 rasuflana (Hauer); (a-b) lateral. contracted; (c) trophus. | after Hauer (1937); 2-4 after 
Kaste (1978) (various authors), Scale lines: adult SQum, trophi JOpm. 


confirmed from Australia (Koste & Shiel 1980), is the 
record of Berzins, is unresolved. Neither figures nor 
description were given by Berzms. 


Trichacerca gracilis (Tessin) 
FIG. i1:1 

Acamhodactylus gracilis Tessin, 1890, p. 155, Fig. 2:14, 

Inchacerca gracilis: Curlin 1939, p. 36, Fig, 10a, 
Type locality: Rostock. Germany. 
Holotype: Not designated. 
Description: Head defined from trunk by suture: 
no large teeth on occipital margin, low keel on right 
side of lorica reaches end of lorica, No  trophi 


description, May be confused with the similar 
£ ternis (Big, 1:2). 

TL 210-227um; LT 81-90~m; RT 26-30ym. 
Distribution: Europe, N. & 8, America. Rare, between 
submerged plants. Single record, Solomon Dam, Palm 
Island, N. Qld (coll. P. Hawkins), 

Literature: Shiel & Koste (1985). 


Trichocerea inermis (Linder) 
FIG, $:2 
Coelopus inermis Linder, 1Y04, p. 240, Fig. 4:9: 
Trichacerca inermis; Edmondson 1936, p. 219, Fig. 28:10. 
Type locality: Lake Bret (Switzerland?). 


Kt Rh. 2 SHIEL & W. KeaSTE 


Holotype: Not designated 

Description: Resembles 1 dixen-nurtalli, T pusilla und 
T star, Stout body; light ssulare ventrally 
distinguishes bead-sheath; mumerous {olds in head- 
sheath on contraction; dorsum may be arched: suite 
palpar organ in ciliary field: PT <a bady length; RT 
‘LT; trophi not described. 

TE, 1385p, BL 9Spym; LT 30-45m) RT 12-130. 
Distritndion: \solated ovcurrences in lakes, Europe, 
N. America, Single record, Sheepwash Billabong. Yea, 
Vie 
Cimnents Edorondsott (1936) aoted that the body is 
shorter and thicker than (hat of 7. divensniialli, Rosie 
(978) noted that only the shorter right toe al trevnis 
separated the nxt. Until trophi structure ane compared, 
the status of these Iwo taxa is unresolved. 


Trichoverca insienis (Herrick) 
FIG. 5:3 

Disrella insiems Herrick, BBS, p. 50, Fig. 4, 

Trichocerca instenix: Edmondson 1936, p. 28 
Vype locality. USA. 
Halotype: Not designated, 
Deserypmion: Body clongate cylindrical, tapers (or (bat 
in posterior M4, lorie anterior projections usually of 
similar length; lorica heightslength ca, 1:5; keel begins 
between anterior teeth and runs fo caudal end: pilpar 
organ and two ciliated papillae in apical field; TR: 
Yulerum distally anchor-shaped, manaheia rod-like; 
LM curved tmwards distally; rami umd unei wath 
denticles. right ktteral antenna at the end of darsal keel: 
viteHarium with indentations; on average larger than 
other species; probably related to T. wevensi (Fiy, 6.3), 

TL 320-376. BL 200-257 yim; LT 90-152ain, RT 
SN-75yim: TR O2um (F 48ym; LM 38am; RM 12pm), 
Distribution: Europe, North and South America. New 
Zealand. In periphyton of standing and flowing water, 
uccamonally in plankcon. Paneontinental) rare; 
13.5 -31,2°C, pH 4,5-80, DO 74-84 my I!, 
4640008 erm! to 16 NTU, 
Lueraiure: Koste (1978). Stiel & Koste (1979). Kasle 
& Shiel (9846). 


Trichocerea insulana (Hauer) 
FIG, 5:4 


Dywtella insatuna Hauer, 1937, p. 378-9, Pig. 2c, 
Trichacerca tnsulana: Carlin 999, p. 45 


ype locality: Moor poul neae L.. Toba. Sumatra, 
Tndinesia 

Holorype: Not designated 

Deseriplion: Body cylindrical, dorsally humped: 
prominent! anterior margin spine tron which long. low 
keel suns diagonally almost to font bases tees ca. 14 
body length, of approximately similar leneth, sec wide 
apart ut theit base; fulcrum double-crooked, notably 
longer than manubria; LM chin, terminally shehtly 
expanded. not crooked; RM shorter than LM, roll ke, 


more deheate. Rescmibles 7. tigris (Fig. 8:3), 

BL, 92mim: foes 33 35am; BL including aoterw 
spine lOOuun: lateral height 34am; LT a8~m: RT 35 pm5 
Dixtribution: Mud, sand. periphyton; Sweden, 
Indonesia, Two records; Magelu Ok, N.T., L. 
Dulverion, ‘Tas.. 18.0°C, pH 7.2, 33305 em! 
Literature; Koste (1978, 1981), Koste & Shit (986), 


Trichocerca intermedia (Stenrogs) 
FIG) 6:1 

Coelapas intérmedias Stenroos. 1898, p. 1D. Tay. 210. 

Trichocerce: intermedia! Rdmondson t256. p, 214 

Trichocerca mona Bauer, 1956, pp. 308, Fig 29 
Te locality: Fintaod, 

Halarype: Not designated 

Description: Small species: body cylindrical! head 
sheath separated by constriction, when head 
contracted, nine folds identifiable in head sheath; single 
tooth at dorsal anterior margin just to right of midline: 
striated area extends back from base of tuoth to dorsil 
midline; foot very short. toes of equul length; lateral 
antentia Widely separated on dorsum) right in posterior 
i ofabdomen, lettin the midline between wand dorsal 
antenna: whorls is just in frontof head constriction; 
Foot gland including reservoir very long, TR nowbly 
iarge. 

BL. 9U 1O6um (without toes); toes 23-30um: TR 
33um (F 25am, LM 25zm. RM 14/l2Qyer). 
Distribution: New Zealand, Europe, Nurth Americ. 
between water plants in standing und Mowing water, 
Four Widely separated records, probably niore 
widespread; Magela Ck, NT) (as 7) tiveontecnis): 
Bromfield Swamp, Qld, and two stock dams in 
southern Tas. (the latter with cyanobacterial bhorimns) 
W.5A7S°C, 230a8 env. [3-10 NTU, 

Literuiure: Koste (1978, 1981), Green (1981), Koste & 
Shiel (1987). 


Trichaverca musculus Haver 
FIG, 6:2 
Pitrella mutseulay Huuer, 1930, py. id, Pig. 2:10. 
Trighowerca musculus Carlin 19399, @ We 
Tipe locality; Getmany. 
Huloiype: Not designated, 
Desoripuion, Body short. squat: head sheath folds 
project as “corner folds” (two dorsal and one ventral 
denticulate mocrones); keel striated approximately 
dorsum length; LT longer than right; crook of LM 
“shoe Jast” shaped; suprarant) approximately 
synimetrical, 
TL WS—170pm;, BL 8O—-I32jam. toes 30-43) 
25—35um; mastax 39—43~miF 30am, LM 30um: 
fight uncus. 13am). 
Disiritution: Europe, North America; indicator for 
oligosaprobic water: in periphyton, in pools, lakes. 
moors. Two records, Mt Kosciaske, N.S.W. and 
southwest WA, 
Literature: Bercins (1982). Koste pf vd, (1984), 


TRICHOCERCID ROTIFERS " 


Fig. 6.1, Phichoverce intermedia (Stenrous): (a) swimming, lateral: (b) trophus: (¢) left manubrium. 2. Trichacerca musculis 
Hauer: (a) lateral, contracted: (b) trophus. 3, 7 myyersi (Hauer) (a) lateral; (b) trophus, 4, 7 porcelhes (Gosse): (a) swinuning. 
lateral; 1b) trophus. Afiér Koste (1978) (variuus authors), Scale lines: adult 50 ym, irophi lOym, 


Trichocerca myersi (Hauer) 
MG. 6:3 

Diuretla myerst Hauer, 1931, p. 174, Fig. 2a b. 

Trichoverca myersi: Catlin 1939, p. 44. 
Typé locality: Germany. 
Holoiype: Not designated, 
Deseriprion; Anterior lorica spines (muerones) yery 
different in length; lorica heighti:widih 1:5; body 
fusiform, dorsal keel runs almost to height of right 
lateral antenna (two antennae at different heights); 


short head sheath, distinctly offset by suture: foot short. 
somewhat obliquely placed; LI’ weakly curved, 
reaching, about 4 body length: RT (tightly placed) to 
left, with substyli often cemented together by foot gland 
secretion and difficult to see. Trophi: LM robust rod. 
only weakly curved terminally; suprarami with pincer- 
like inward-directed apices, 

TL 270-310um; BL 180-2104m, LT 90-102um,; RT 
50-S7j:m; TR 50-6lpm; F 36um: LM 38um; LR 
21m) RR 2am: une7 yum. 


ne R, J. SHIEL & W, ROSTE 


Distrifadian: Probably: cosmapotitan, in standing 
waters, i periphyton al weedy ponds. Jabilukit, NTL, 
1. St Cluir Natl Park, Tas. , Moorabool R,, Vie.) rare; 
10.0-27.0°C, pH 54-63, DO $1 mgt!) 4225 en! 
Literature; Koste (1978), Koste & Shiel (1987), 


Trichoverce porcellay (Crosses 
PIG. 6:4 

Monacerva parcels Gosse, 1851, p. 199. 

Trichovera parcellus, Myers 1997, p, 6 
Ape locality: England. 

Holotype: Not designated, 

Descriprion: Short. plump. distinerly curved body; 
head sheath separated by constriction, lwo anterior 
dorsal cusps, right one slightly larger; when head 
slightly contracted, lips of a ventral notch un the lorica 
murgin may protrude as slight “teeth” but are not seen 
in extended animal; variable height nudge (sometimes 
absent), striated, extends backwards from the largest 
tooth; vorona with club-shaped palpar orean; fool 
sniall, partly englosed within Jorica; lwo toes, lef 
longer, ca, the same as body width, the right sumewhat 
shorter, usually held against ventral abdemnen, Trophi 
very asymmetrical, T.-M robust, crooked, RM a slender 
rod, LR aldla much longer than right. 

BL 90 -NW5pm: body width 45-48nm, toes 22-42 em; 
male 56-604m long. 32- 364m wide (a larger Torm, 
f, wigior was deseribed by Hunter, 1935), 
Distribution; Cosmopolitan, pH 6.0-6.7, 12.1-9.0°C, 
Jarger in wkaline than acid waters, Indicator for 
ohigosaphrobig waters: all forms in linoral. in 
periphyton, occasionally in tychoplanktem, May be 
puncontinental, not yet Kilown from S.A., W.A,; 
common; 8,0 22.0°C, pH 5.2 7.5, DO 61-112 mg I", 
16-1N20"8 em! <I-2O NTU 
Literathre: Evans (950, Green (198l), Koste (1981), 
Keste & Shiel (1987), 


Trichocerca roussetert (Voigt) 
FLG, 7:1 

Coelopun cousseleti Vous, O02. yo 38, 

Trichocerva rousseleti: Abtstrom 1938. p. 92. 
Type locality: Plon, Germany, 
Holonpe: Not designated. 
Deseriprion: Squat body with arched dorsum, anteriot’ 
lorica margin with 8-9 projecting serrations, the dorso- 
dextral tooth largest; striated area of other species 
replaced by furrow between dorsal teeth; head clearly 
distinguished by transverse suture; RT about 4 length 
of Jefi, slender, casily overlooked, ‘Trophi: manubria 
of similar length; loft uncus with short denticles. May 
be confused with T sae (Fig. (5). 

Th W5-IM45ym: BL 72-Naym; LT 27-32jun: RT 
1) Wp: TR 33ym (F27 em; LM Slyem:, RM 27 pins 
palpar organ to 20pm. SE 46%29 yin. 

Distribution: Previously known from Palaearctic and 
Nearcim oligusaprobie waters, with sporadic 


aecurrences in plankton of Jakes. where cpyy ane 
ultached to Melesira filaments, Population maxima in 
spring, Rare, in our collections invariably associated 
with Howing waters = Burling (N.S.W.), Goulburn 
(Vie). Murray (S.A), with Melosira blooms. 
14.0-17.9°C, pH 70-8.1, DO 8.9 98 mg. 47-365uS 
cor! 28 135 NTU. 

Lirevertures Shiel & Koste (1979), 


freehocerca rutiner? (Dinner) 
FIG, 7:2 

Divrelle dixen-nuralii; Hauee (9397/38, p. 409 

Tehacerea rater’ Rovner 1953, p 19-92. Fig. tad 
Type locadiry: Nov specified: Jake plankton, Sumatn 
and Java, 
Holotype: Not designated. 
Deyseription, Plumper than very similar 7 dixon 
nuttalit; tubular lateral antennae at simitar height, 
dorsal antenna displaced to right; LT slightly sigmoid: 
ont longer substyle, 2 length of others; robust (rupli. 
fulcrum inverted T; manubria of similar lengifi, bent 
to crooked in distal 5; left uneus With four stramy teeth 

BL 22-200jan; height 63-82¢m; LI 53 85am; RT 
29-40um; TR 36-43pm, 
Disibution: Widespread in tropics, Burope, Known 
only from dams near Chillagoe, Qld (coll BV 
Timms). 
Literature: Shiel & Kosle (1988) 


Trichocerod similis, (Wierzeyskt) 
FIG. 7:3 

Cozlopus similis Wierze|ski, 1893, » 406, 

Trichacerca similis, Edrontsun 1935, p, 303 

Trichaccrea birastris (Minkiewice); Kite 1978, pe 393-394 
Fig. 136ith. 
Type locality. Galicia, Poland. 
Holotype; Not designated. 
Description: Fusifoom body; head sheath marked hy 
disiinet suture(s); two slender. subequal occipital spines 
somewhat deflected] to right. flex with lorica 
movement, may be bent across head aperture on 
gontraction, may be knob or spinule betwee spines, 
two low keels extend back from spines, with narrow 
transversely: striate arca belween; dorsal antenna in 
strine field: knoblike left lateral antenna well betore 
midline, right lateral antenna slightly before foot: end 
of rump overhangs first foor segment, two toes 
unusually short, unequal: LT ex. 4 lorica length, RT 
Shorter; torsion of abdomen has placed RT base above 
LT, diffioute to distinguish separate toes in dorsal view! 
1-3 substyli present. Trophi: fulcrum straight, knife- 
like in lateral view; manubria asymmettival, lef more 
robust than wight; right uncus multi-toothed: 
retrocerebral organ notably large, may extend beyonu 
midline; bright red cerebral eye, cwo lateral ocelli 
visible in living specimens; salivary glands absent 


TRICHOCERCID ROTIFERS In 


Fig. 7.1, Trichocerca rousseleti (Vougu: (a) lateral, contracted; (b) head margin, (€) trophus. 2, 7 rarer (Donner): (a—b) 
literal, contracted; (c) hase of toes; (d) trophus, 3. 7) simi/ts (Wierzejskid; (a) swimming. latera]; (b) dorsal, contracted , 
{v) trephus, 4. 7° silts erandis Hauer: (a) lateral, contracted; (b) trophus. Afler Koste (1978) (various authors). Scale 
lines: adult SOgim, trophi 10um. 


TL 166-300nm; BL to MMOpm: LT 50-804m, 

RT 30-50jum; 'TR 30um (F 2i4m, LM/RM 30/24um); 
male 68—73um. 
Distribution: Important component of plankton in 
oligotrophic, huniic waters, attaches subitancous cegs 
to other plankters. Pancontinental, most common 
Trichocerca in Australian waters, planktonic m lakes, 
ponds, billubongs, stock dams, Particularly common 
in humic acid waters in western Tasmania. 
7,.0-24,5°C, pH 3.9-8.2, DO 54-116 mg I>. 
H-AWOKS cm!. 0.5120 NTU. 


Comment: A larger form often found with 7, similis 
has. similar morphology, including occipital spines. 
ft is common in Murray-Darling reservoirs, ¢.g, 
L. Hume, L. Dartmouth, where it grazes green algae 
e.g, Gloeocysris. This large form is presently regarded 
asa ssp.. T similiy grandis Hauer, 1965 (Fig. 7:4), 
It is distinguished by larger, more elongate body and 
relatively shorter toes than thé typical form, 
TL 400-525ym: LT to 44pm: RT t 28um. 

Distribution: Known from Amazonian floodplain 
waters. Rare; River Murray billabongs in Victoria, 


i R. J SHEL & W ROSI 


SS. e 
6a Res 


es. 


Fie 8, L. Tnehoveren yuleetta (Jennings): (a) Swirmmang. lateraly (h) teaphus, 2, Fre derenr er fenmioe (Hilson & Gusse: 


7 


(ap lueral th) tophus. 3, rigies (Muller): (ay iateral, contracted, (b) trophus. 4. 2 ueinefer (Voigt areval. contracted 
5. Po veehalts Hauer, Gu lutcral, contracted) (b) (nuphius, 6. TZ avebere Jennings): Gil lateral: (by sstr, front Tasmania. 
lateral, to} frophus, |, 3 Satter Raste (978) (various authwors): 2 afer Kose & Poliz (984) 6 atior Keate eal, (988). 


Seale lines; adult 50,0. traphi (jen, 


L. Pedder, Tas.. probably more widely distributed. 
15.5-20,0°C, pH 68-70, DO 10.8 me |! 

Literaivre: Koste (1978), Koste & Shiel (980, 1987), 
Koste er al. (1988). 


Trichocerca suleata (Jennings) 
FIG, 8:1 

Rattus sadoutis Semungys, 1RY4, po 20, Fur. 8. 

Trichocerca suleate: Myers (837. p, 6. 
Type locality; Lo St Clair, Michigan, U.S.A. 
Hitlotypes Not designated, 
Description. Body cylindrical. doryum arched to form 
hemisphere, two prominent sutures mark head sheath, 
which has numerous eréases. om contraction; two 
tongue-shaped fleshy projections from anterior margin 
uppeéar triangular in lateral view, right most obvious: 
shallow. transversely striate furrow alang dorsal median 
fine; foot and short, equal-length toes curved forward 
under posterior, held adpressed to body; dorsal antenna 


in Median furrow, lateral antennae al similar height 
on posterior 4 of trunk; trophi asyarmetrical, robust. 
fuleram double crooked: LM reaches end of fulcrum, 
distally dilated: RM a shorter, slender rod; ramus with 
enlarged bifurcate alula: brain wath honeycombed 
retrocerebral sac, evespot at base of brain: foor glands 
with large mucus reservotes, 

BL 9732p: Woes 13m: TR 40—-42m (F 33 um; 

LM 2800, RR t6ym). Wulfert (968) vives BL 200m: 
1oes lo 35m. 
Distribution: Possibly cosmopolitan in periphyton of 
submerged plants, Recorded by Berzins (1982) trom 
the Avoca Royer, Vie, No ecological information. not 
seen in our collections. 


Trivhacerca tenuinr (Hudson & Gosse) 
FIG. 8:2 
Caoelopus leniior Hudsim & Gosse. 1886. p. G8, Fig, 20219 
Trighocereu fenuivr: Uauer (93k p. 179, Fig. Sa 


TRICHOCERCID ROTIPERS i 


Tipe locatine No single localuy specified. “Woolston: 
Sutton Park and Colestill, Birmingham’, England 
Holotype. Not destenated, 

Description, Body elongated. euryesl, with single acute 
Ipoh we right anterior margin ot loriea: lef oF tooth 
is. low. pleated plate; transversely striated low keel 
continues obliquely from base of tooth to ca, midline: 
hewd sheath marked off by suture. has longitudinal folds 
when head withdrawn? laters! antennae at different 
heights, lel more anterior: foot short. offser from 
trunk. toes unequal — left tue ca. half lorica length, 


slightly curved ventrally, right more delicate, fall 


length of left, sabsiylt present: trophi asymmetrical; 
lulerum double crooked: LM almost fuleruny length. 
distally with 70-80° inward bend: RM fragile suntight 
rod. RR with triangular alula pointing aliriost at right 
ingles to fulcrum; gut may be orange coloured. 7 
fenuior inay be contused with T graciliv (Pig, Wel. 
T. tnsigniv (Pig, 53). 72 trrermedia (Pig. 6:1) and 7 
veriy (Pus. 8:3), can be separated readily by traphi 
morphology. 

BL 125 210m: height 40-50pm; LT 54=80pin; RT 
35 3pm; TR AQ 45pm (F 3) ym: LM/RM Shem; 
nimi I7/{Syin) subtaneous ege 65%38.«m. 
Distribarion; Cosmopolitan in detritus. in algal mats, 
iIn-periphyton, psdurimon, o¢eamonally in tyehoplankton 
a) scl) waters or in beach sand of flowing waters, also 
in moor pools. Ritre in mutiakind samples (Vic.). more 
connor tn Tas.> 80-22.0°C, pH-6.0.70, DO 10.0 1.2 
me (), W-70008 em!, LI NTU 
Literate: Rosie & Stel (987). 


Trichocerce tigris (Miller) 
FIG, 8:3 

Trivhida siuris Miller, 1786, p. 206, Fig. 29:8, 

Trecheeeeca Hiss Vore 957, pe a2l, Fie. 6312, 
Type localiine Copenbagen, Derimuark? 
Holotype Now designated, 
Description: Similar morphology to 7) tenuder. with 
elongate, curved cylindrical body, single antenor 
Hecipital (oot, longitadinully pleated head sheath. 
oblique low keel arising from) bse of tooth: differs 
in usually greater size. more prominent keel, relatively 
large foot, equal length toes, trophi structure (of, Pigs 
§:2b and &:3h). 7 rigeiy has distinctive anchor-shaped 
famubnumn, double crovked fulerum, May: alio be 
confused with 7) fasdlana, avuin readily separated on 
wopht structure (ct, Pigs 5:4, 4:3). 

TL220-300;m; BL 130-200, TR teal yr, toes 
50 — $0. 
Distribution: Cosmopolitay, isolated agcarrences. in 
wide range of water quality. io periphyton and benches 
(Kuste (978) Common tn billyborgs, Vie, Tas: 
16.0) 220°C. pH 64-74, T4=274a8 em! 
Literatures Koste & Shiel (987). 


Trichocerca ancinatd (Wait) 
FIG), 8.4 

Caclopus tacinatiis Voigt, WO2 p. 07 

Trivhavcria uncineta Carlin W3Q, p. 43. 
Type lecatity: Plin, Germany, 
Hoalotyper Not designated. 
Deseriprion) Body short. curved. anterior orice 
tanga shghtly denticulate; acute occipital curved tooth 
displaced ty night of midline: low tongue-yhaped plate 
to Tefi of taoth: ventral striae or pleats run to weak. 
suture separating heud-sheath from trunk; short foot 
with two unequal curved toes, trophy asyiinietricul: 
left uncus club-shaped with several acute teeth, riyghi 
wncus shorter, plump. bifureate: large red evespor. 

BL 65-95um; LT 12 271m, RT 10-25jan. TR 
27m; oevipital tooth 14-27 nn, 
Distribaion, Probably cosmopolitan, im periphyton, 
algal mats, psatimon, Wehoplankton of fresh waters 
Incontirmed record from Tabruhbuoca Creek. 
Barrington. N.SW. No evological informition Not 
seen in our collections. 
Literaiure: Kosie (1978), Berzios (IYR2)- 


Trichocerca vernalis Hyver 
FIG, &5 

Trichoveres vernaliy Hauer, 1936, p, G4. bie | 
Type locality; Germany 
elope: Nov desyinated, 
Deseription: Squat, vaulted to conigal body: left 
unteor Minin With ire rounded plate, foe and toes 
curved forward: shori diagonal keel from striae field: 
wes of similar length. teuphi strongly asymmetrical: 
[cA manubrium robust, with hoekey-stick’ crook, right 
tiunubriuin wa slender tragde rod with median kink; 
fulertio) double crooked: lef) uncus, with two teeth: 
subuncus with several (commonly three) tine teeth 
Resembles 7 sudeata (Pig. 8.1). 7. bractovure (Pig. 4). 
int particular the latter if trophi are left too long in 
caustic solution — Jeft manubyiual ray dissolve away 
to produce the characteristic bend of vernadic’ 11M. 

Bl. 86-138: height SI-Ssyamy, LT 26-43yn1, RT 
to 38am; TR 4) 46em On TR 45am, F 32m, RM 
20um, 1M ta 34pm), 
Disiributane Ephemeral waters. mm Europe, N 
America, Indonesia, Rare, bUllabongs and ephemeral 
pools, R. Murray and Mitts Mata Ro. View 40-2107, 
pH 6.2 685, 73-2922S8 enr!. 
Literature’ Berzins (\982), 


Trichoverca weber| (Jeaningss 
FIG. 8:6 

Dinrella weber? Jenninys, 1903, p, 309, Pays tlh dd: 
13 WG. U7, 

Trivhaeerca webert, Edmondson 1938, p. 4035 
Vpe lacality: Not specified Lake Eric and vicniey. 
Michivan, U8. A. 
olorype- Nut desiguared- 


MW Kk J. SHIEL & W) KOSTE 


Deveription: Body short, curved ware. head sheath 
indistinetly marked by sutures broad, rounded 
proyecting plate td left of head aperture: short pulpar 
organ in apical ficld, single proninent oecipital tooth 
lo cight of dorsal median line: occasionally a spinule 
between plite aml tooth, Aiygh, tp, transversely 
striated keel from median tooth to $4 lorica length: toes 
of similar length: trophiasynimetrical; LM with distil 
right-angled hend; RM ca. 4 Jeneth of LM, slightly 
sigmont rod. suprarami distinery ¢longated: toes with 
three-four inconspicuous substyli, Resembles 7) eat 
(Fig. 4:2) and T. poreellus (Fig, 6:4), 

BI 95 iS5uni, heleht to SOuar, LT 30-Saym, RT 
lo 42um: occipital spine to 12pm; frophi to SZpm, 
Distribution: Probably cosmopolinan, net recorded 
from Africa (Roste 1978), bt periphyton of literal, also 
mivors. Rare. Old, Tas,, Vic,, Tasmanian specimens 
slightly larger than hose of the mainland: Lt to 6Q,an, 
RT te 50um., Possibly ecotypic variation, 
Lirerunre: Russell (961), Koste et al. (IYssy. 


Trichaverce (8. Str) 
Trichveerca agnata Wultert 
FIG, 9:1 

Tite huverea avinata Wulferr, 1999, p. 73 78% Fig. 5. 

Type locality, “Heykasee bel Koshin’. Present Poland 
Holotype: Not designated, 
Descriptions Mediuin slender body with lightly convex 
dorsum: convex abdomen, 12 somewhat sintilar lane 
head creases without frontal process (in swimming 
animals folds not apparent): corona with one palpar 
onguin and (wo antennae; dorsal anteanae in pit. sightly 
displaced qe night: left lateral antenna medial: right 
amenna at ilistal end aFabdomen;, LT straight, about 
» BL, RT <WS5 length of lett. TR: LM longer and 
more strongly built with curved distal end; RM shorter 
more Slender, alse curved in at distal end: LR with 
longer pointed dluky RR weukly developed; supraranti 
staal; DB finned distally. 

TH. 23000, BL (contr) dum, LT 73am, RK 35,0. 
Diseribariuns Europe. Single record from the River 
Murray at Echocw, Vic. 15.0%. pH 74. DO 90 mg 
V. 

Literature: Koste (1978), Koste & Shiel (980). 


Trivhaverca bicrisrara (Gosse) 
BIG 9:2 

Mastisocerca hicriviata Giaske, 8&7 p 2, Bie 3 

(ehow tied lieristata: Harring 1913, p. JOL 
Tipe locatiz: Scotland, 
Hafatype: Not designated. 
Description’ Bou slender and plump forms known: 
in corona two shift projechons: in cross section twa 
variable height ridges separated by wide depression run 
length of dorsum, veering, slightly to left. LY with 
triapical tip, median apex much longer than lalerals; 
lateral umtennae at similar height at end of abdomen, 


TR: both manubria lerminally crooked, one side 
curved, supraram lume. LR oocustonally with two- 
painted alula. Lirica finely stippled; sometimes red 
brow coloured; RE with irreq@ular “rode” 
reuiforvement between the shells. May be confused 
with 7. ecesa (Fig 12-2), but ts larger, with longer. 
more obvious Keel structure, 

TL 294 660um; BL including fuot 194.360;,0n; 0.7 
200-320um; RT 25-36nm; TR 65-79,.m (F SOun, LM 
hOpm, RM SOpm, rume 36-SOam, dae 2207)., 
Distrdurion: Cosmopolitan. indicater of aligosaprohic 
witters; isolated occurrences ia littoral in detritus 
oscasionally in tyehoplankton Rure in Murray-Durling 
Basin, also Cape York. Oli and Kakadte Natl Pk. NZT. 
$.0-24 S°C. pH 6.3 84. DO 53-130 mg I, 
59-5738 em'. 23.5-40,.0 NTU. 

Literature: Colledge (1914). Koste (1981). 


Trichocerca braciliensin (Murry) 
FIG 9:4 

Ruituhay bragiliensis Murray, Wiha, p. 244. Fig Wi 6a, b 

Trichocerca brasiliensis, Vauer i965, yp. 378, Pig. 32. 
Tre locality. Walter lily pond, Praca Republica, Rio 
de Janeiro, Brazil, 

Holotype: Not designated, 

Description: Short, stout bady with distinctive double 
keel 4-4 length of dorsum, body broadest behind 
keels, tapers to foot: keels strongly rounded; LT longer 
than hody, continues tine of body, with tmasal simon 
bend; RT “length lett); TR resembles those of 7 
mucosa and 1 bieriviate, Out whereas Ujese taxa have 
manubria with only a hintala crook. in % Aresiliensis 
both are strongly crooked (Fig. 9:°3b); RR with 
mirkedly bifurcate aula, LR with single spinelike 
prowess, 

TL 27) 275,in. contracted lores 120-122 qui, keels 
38-40ym; LT 150-153~m: RT 33 -SOum: substyles ca 
Ou; TR 57 60ani (F 40pm; mianubrs 40 and 24am; 
unel O-and (2am), 

Disiribuiion, Raye; previously recorded only from 
vegelaled waters im Suuth America. to 308°C, pil 
45-67 (Koste J978) Single record. Tasmania, several 
individuals. from ad rowdside marsh, Deloraine 
(22.10.87). 18°C, pH 6.9, 106,5eS en, 

Literatue: Hauer (1965). Koste er ef, (1988), 
Gomment: Although synonyinised Will 7. elongates yy 
Koste (1978), 7 braziliensiv” smaller size, general 
murphology and specific trapl differences 
(particularly biforedte alula of LR and more robust 
crooked LM) ap ligured, sdggest speeitic status be 
retairiect, 


Inchocerca capucina Wierzeyski & Zachattas 
FIG. 9:4 
Masngocera cupucina Wiereeyshi & Zachuriws, 1893, 
pe ede, Vig. a 
Trichieona capuilna. Haecityg 19, ps IO 


TRICHOCERCID ROTIFERS \7 


Fiy. 9, 1, Trichocerca aumata Wulfert: (a) lateral. swimming, individual from Darling R, at Bourke; (b) lateral, contracted; 
(¢) trophus. 2, 7) biertstara (Gosse); (a) contracted, lateral, from lef: (b) trophus. 3, 7) braziliensis (Murray): (a) lateral; 
(b) dorsal: (c) trophus. 4, 72 capucina Wierzejski & Zacharias: (a) lateral, swimming: (b) trophus. 1 b,c after Wulfert 
(1939), 2 after Waltert (1956): 4 after Hauer (1965); 4 after Koste (J978). Seale lines: adult SOum, trophi 1Oum-. 


ik Rk. 1 SHIEL & W OST): 


Hype localin, (Germany) 
Flalurypes Not desigaatent. 
Dexeription: Body tusilucn. head with 5 palps and two 
antennae: abdomen cylindrical, slizhily curved: eowl- 
like loricale head with vettuetible apjecs (=nucrones?) 
demurcated from trunk by Uistincl tritisverse suture; 
lateral umlennae al sinuher height; “TR almost 
symunetrical, RMDP slightly more robust; TR parts 
relatively fragiles mule loricate with large red eyespor 
und rudimentary toes; RE brown, hurd and shelled; 
TY, 250-d30jan: LE 90 2 5yam. RP 45-48am), male 
90-1O0um, RE lO8-(28 X XO 82am: SE BH WO X 
O8 Flyin: mule eae 60-04 X 70 80h. 
Distribution: Cosmopolitan. in plankton of lakes, ako 
in muer and attallassig saline winters! attaches its epes 
to other rotifers e.g, Asplonehra; leeds on eges al other 
rotifers, ey. Keratclla, sucking out the contents. 
Unewmimon; NT), Qld. Tas. Vie. 79-24.5°C. pll 
587.4, DO 5.898 te |. 38-02pS em', 10 50 
NTU 
Lirerenaes Murray (OG a), Koste (978). Koste & Shicl 
(1987) 


Trichacerca vheattond (Bauchamp) 
FIG, Ith 

Raretis avtindricus yar chatton Beauashanip, 
p S46, Mig 4 

Trivhocered chatonis Hiauee Wak. p S68 

Irichacerca Slindrica Chathnii: Koste (978, po AOS, 
We locality: Pond of the Dombes (narth of Lyvan, 
France) 
Holotspe: Not desigiuterd, 
Descrtprion’ Body cylindnieals contracted anrmal his 
numerous folds in head producing undulate or serratea 
Jrumal margin. no pronounced suture ab neck, only 
sheht- depression: disunetive churaeteristic single long 
ventrally curved spine left.of dorsal antenna ansing 
front thickened base; TR not described in Keste 978 
Distinguished tron 7. cylindrica by smaller, more squat 
hotly. shorter LT. less reduced RT and ratio of clit: U0 
(Lomas, l6nnnia To eviitdrica, < Spm ia 7 cfuation). 

Th 300-448pm. LE 15-40um; RT 24-39"m, 
anterior spine 26 Slam. 
Distribution. Tropical and subtropical waters, Europe, 
Affica., Indonesia, South Ameneu, probably. pan- 
trypical/subtopicul, pH 5,7-7.3 to 30.5°C (Koste 1978). 
Rare, confirmed Jrom ca. 4 Jogulities in the 
Kamberloy, WA, Magela Ck, NUT. and Cape York, 
Qld [collections by, Po Hawkins (Townsville), BV. 
Timms (Cooranbong) und) MAI, Tyler (Adelaide). 
IS0-29.0°C. pH 615-74. DO 37-80 mp Tl 
48-2458 cn'. 05-6 NTU Warm stenotherm? See 
comment helaw fer 4 ewinepice. 
Literatiere” Kaste & Shiel OBR), 


WOT, 


Trichoverca evlindeioa Uimbut) 
BIG. 10:2 

Mastisecerca evlindrica tinhot, WY, p47 

Tchocerd uvhudrivas Harring, 113. p, (02. 
Type localiry: Lake in the Black Forest (Germany), 
Holetype. Now designated, 
Deseriprion> Wongate, cylindrical body. very fine. 
ventrally-curved acute spine from median dorsal lorica 
margio, may he folded down and invisible in contracted 
animal: longitudinal striae in head region in latter case. 
short keel amd strimed field: LT always longer than 
body; RT reduced to rudimentary short, scaly spine: 
dorsul antenna long, rygid, lett lateral antenna in middle 
of truok— right just before foot, TR altriost symmetric, 
but LR mauve robust: bot munabria longer than 
fulcrum. curved distally, with lénestratet proxinial 
ends (big. 10:25), unital oveasionally un gelatinous 
sheath; eges carried altached ty parent, mule locless, 
RE jo striated gelavinous sheath 

TL 990-T96pn1; body 225-46sc01, LP 190-328: 
RT tw 2twiny TR 50pm (F Joum, manubria 46m); 
male $0-90mypm; male epe 52% 3500, 
Distribation: Palaeareiic aod Nearetic lakes, pools and 
moors. Sudauki (1967) recorded f evlinedetve Vrom L. 
Sorret} iy Tasman, we subsequently found it in b 
Kucumbenc. Snowy Mts, and L. Dartmouth, Vie 
12-18.0°C, pH 6.65 704, 26 47S cm, DO 7.0 my 
Jal, 
Comment) Apparently two ecologically, and 
taxonomically digtinet taxa oceuc in Australia: the 
Jurger cool water species, T cvlindrica, and the (rapical 
f chattoni. Koste (1978) synonymised the farter wilh 
the former (as a var), but on the basis of apparent 
ecological and morphological differences We retain Whe 
distinetion until SEM analysis can clarity the stulus 
of both taxa. 
Literature: Sudzuky (967), Koste & Shield (Ysa), 


Trichocerca elongata (Hudson & Gossel 
FIG, 10:3 

MOST enu elenvata Hudson & Gosse, IRBG, |). 62. Fig. 
OR, 

Tickocerce elonvatas Harring M13. p02 
Ape fovaliry. Loch near Dundee, Scotland, 
Holorype; Not designated, 
Deseripiiun, Laree species, body long and slender; 
head sheath not marked by constrictions: untenor edge 
unarinied, short double keel about  leneth; dorsal 
surface strated approximately Va length antenor loriea 
wich median furrow hack to dorsal antenna; coroni 
with sinmle palpar organ: lateral antennae at same level 
although torsion ot the body means left ania is dew 
roore ventral and right antenna closer to dorsal bine 
(only one seen iedorsal View); torso has also moved 
toe to dorsal (right)/ventral (eft) position rather than 
side by side: LL more than 4 lores length, RT 
rudimentary; TR asymmetrical. left side  mure 


TRICHOCERCID ROTIFERS 0) 


Fig. 10, 1, Trichoverca chattoni Beauchamp: lateral, swimming, 2. FE eyfindrica (imhof); (a) lateral. in gelatinous sheath: 
(bh) trophus. 3, 7 clonyara (Hudson & Gosse): (a) lateral, contracted; (b) trophus, 4, 7 flagellata Hauer: (a) lateral: 
(b) trophus: (c) dorsal, la after Beauchamp (1907); 1 after Hauer (1938), 2, 3b, 4b after Koste (1978) (various authors) 
Scale lines: adult SOpm, trophi 10jam. 


developed) LM robust. with strongly curved distal end. 
lenustrittcd proximal end: RM a delicate rod: fulerum 
double-erdoked Gtiverted T): LR chds in 3-pronged 
luli (mdidle spine can be missing); loriga miuty be 
reddish-brownish: surfive may alse appear suppled. 
Th. A08-RlOpains BE. 130-460,am, LT 160-350nm; 
RT 32-56n1 TR 70-8Oum: (LM to 63m; RM 
t5yurtr) 
Distribution; Cosmopolitan in hitarak and pyeho- 
plankton ol Ceshwaters. Pancootitiontal, corned a 
hillabougs, 1L6-27.0" pH 54-74, DO 91-98, 
OO 160048 en), <JO NTL. 
datehature. Koste (1978), Shicl & Koste (1979), 


Trlehoverca fluvellata Hauer 
FIG, 10-4 

Trchowerce Havelians Hauer, 1937, p. 284, Mig, 1. 
Type fecatio AMloali Lake. Madras. India. 
Helonpe. Nut designated, 
Deveriptions Body compact, ovoid: head sheath 
indistinetly demarcated) amerior margin riattsed ih at 
smooth curve on the nght side; the rest undulating: 
medially a smooth noteh: keel high-vauhed with wide 
stritled ured eMends fo the beginning of the short foot 
opening: left lateral antenna somewhat behind the 
middle of the abdomen: dorsal wotennma near beginning 
of second thad of trunk: LT slightly sizmoid, toe 
length: BL index ca. 1.5; CR robust with strongly 
crooked fulerum and LM: brain large with large 
ferme cerebral eye. 

BL W5 128m; LT 172-174m: RE Sum: TR 
60-72am (44am Fr 40am). 
Distributions Warm stenotherm; India. Malaysia. 
Amazon. Three localities known, Mawel Ck, NT. 
Cupe York, Qld and Lo Purrumbete. Vie 18.0°C. pH 
61, DO 67 me I. 6325.em!, 
Literaire: Koste (978), Koste & Shicl (1980), Green 
(IYRI), 


Trichocerce rnin (Gosse) 
FIG. Wh? 

Mastyecerea ternis Gusse. W8% po 860. bit. Ucts 

Hiviweru terns Hace Wl p. Wg 
Type locality, Weland (lacustrine), 
Holoipe, Not designated 
Description: Budy elongute, cylindrigaly head sheath 
scpariied from trunk by transverse fol. single very 
small spine on toned unterior margin, left lateral 
tintenna inserted higher than right: keel with striae runs 
from anteriy margin to end of abdomen; LM nat 
crooked: supranun small: left uneus bidentate. 

Th to 3000p: BL l68-198um: LT 80-100pm: 


Pig WL, Teicher graeiis (Tessioy) Go literal sveumming, (A) trophus, 2. 70 rernis (Gosse): lateral. swimming: 3. 7 


mi) R, J, SHIEL & Wo RKOSTE 


RM yam: TR 38-40p.m: OF 30am; LM 27pm; uncus 
I} jan). 

Distribution: Cosmopolitin, between water plants in 
Hitteral standing and slowly flowing waterss, sityle un- 
confirmed report from Yarra River at Warburton. Vic. 
Literature: Koste (1978) Berzins (1982), 


Trichocerca jemingsi Voigt 
FIG. 1.3 

Ratiuluy seipre Jennings, (903, p. 222, Figs 450-32 (on 
Misiiencrroy stinia Hudson & Crone. IBRG, p. G1) 

Trichevervd jennings’ Voigt, O57, p, 321. Fiz. 68:8, 
Wpe localinn: (U.S.A) 

Holotype Not designated, 

BDeseription Body elongated, curved wath dorsal 
striated keel rising tony anterior loried. passing aemss 
Jorsum diagonally right for ca. %& body Jenath: keel 
with footh at anterior énd whigh wnay extend beyond 
loriga margin in-contracted (ndividuals, head sheath 
indistinctly separated, more obvious on ventral side; 
dorsal antenna at left side of keel: lateral antenna 
approximately equal height in posterior M4 of lorie, 
foot offset from body with posterior dorsal edge of 
lorica projecting over lefl side of fhot, restricting 
movement on left side: LT 45 body length; RT 
rudimentary; trophi asymetricaly LM long. stout. 
curved: RM slender stramhe rod, 

TI, 320408pm; LT 120 77am: RT to 33m. 
Disindurion: North America. Burope, Sr Lanka. NT. 
Tux, mre: 290°C, pH 65, DO 249 met, eS em! 
Literarure: Koste (978), Koste-& Shiel (980). 


Trichacercd loneisent (Sebrank) 
FIG, Ik4 
fiinarta Jongivea Sehrank, WROZ p. 483, Fig. 2:13, 
Trichewerca longisene Barring 1903, p. Ki 

pe lecalinn: Germany? 

Holotype: Not designated, 

Descriprion: Body elongate with two long spines on 
occipital margin, rigbtalmosttwice length of Jett. with 
(wo small projections between xpmes; shallow keel 
from longest tooth to middle-of abdomen, usually with 
striae lateral antenfre i similar height im posterior 
“ whdomen; head sheath with longitudinal fiolds in 
contracted individuals: in ving aniinals. apical fold 
has dorsal elongate pulpar organ beside a membranelle, 
ventral to that two ciliated lobes: dorsal antenna beside 
keel, slightly behind constriction separating head and 
Irunk; |.T upproxamuately body length: RT rudimentary. 
Trophi; Lim long. terminally thickened and curved 
inwards; LR with long pointed aluls: saprarami with 
characterise acute apices, 


jenn! Vout 4, 7 donteisete (Schrank) (0 literal, contracted: (b) (opis. After Koste (978) (variaus aulhors). Seale 


liness adult SOvm. Graph (jen, 


TRICHOCERCID ROTIFERS 


i Rod, SHIEL & W, ROSTE 


TL S00-S75pinv. BL 200-370 am, antetior muerone 
48-550, 0T 100 225ym: TR 64 Bde, RESTO 
X WS-)27 ar, 

Osmibrarioe. Cosmopolitan in most submerged 
vegetunion zones. Punctures eells of green algal 
Hlaments and Sucks. out ccll vontents. Wiilespread, not 
yer recorded trom S.A, or WA. common: 8,5-200°C, 
pH 63-89 DO Fi-Mt me tl t6.6-92748 cm! 
4-28 NTU, 

Literature: Koste (1978). Koste & Shiel (980. 1987) 


frichocrrca macera (Hudson. & Gosse) 
VIG 124 

5 SEIN wv mideoru Hodson & Gersse. 1880. p. Ol. Pip. 
201 

Trichheerca naceras Warring IIR. p. 103 

frivhoeerca ftformiy Levander: Koybe LBL pi. 104, 
Type locality: England? 
Holotype, Not designated, 
Deseriptiun: Elongate fusiform body occasimnally 
curved. dorsal surface more convey than ventral: head 
sheath marked off by shght constniction; small tooth 
oi anterior margin rieht of midline; broad area 
corresponding to location of keel in other species may 
he noticeable bur generally 1 not: dorsal ind Jaccrl 
untwwiinge of usual uppearunee: LT suraighr, ca.) lerica 
length, RT < 4/6 its length; substyli present; when 
toe bent forward a spur-like point extends backwards 
from distal end af foot wt base of toes, not known tor 
uther species: iraphi not deserihed 

TL d40en, BL 278-330um, LT W2-40~m:; RT 
Burt. 
Disrrtbulion: Pealy pond waters, also littoral af barger 
waters, Europe, North America. Rare: N21. Qld, Vic 
Litercture: Koste 1978), Benzing (1982)_ 


Trivhecerca mucease (Stakes) 
PIG, 12:2 
Meantigocema inivasa Stokes 1896, po 17, Fig, 7h 
Trichaverva mova: Hauer 965, po 377 (=T, biertsrana 
pravesea Kosto W978). 


Tipe loeatine * ..% shallow clear-water pool 10a 
rocky wood near Trenton, New Jersey, USA. 
Holompe! Not designated 

Description. Body, seen laterally broadly oblong, head 
sheath with constriction; no anterior tecih or spines; 
deep narrow fold on ventral side when head retriceed; 
two well marked striated dorsal ridges ur keels with 
furrow between them in swimming animals one club 
shaped palpar organ on corona, two slender lateral 
rods; dorsal-antenna left of left-ridge,, in a pit: lateral 
amennt well behind midline: foot short conical; Jorica 
projects well beyond it on left, restricting toe movernent 
in that direction; LT may reach BL Vennings 19032331 
noted that the animal offen swims with the LT carried 
against the right side and appears toeless); RT 
rudimemary jodistinguishahle froin substyli; TR 
Iussive, xtrongly asymmetrical, fileruin crooked. LM 


robust, distally carved. RM much smaller. thin rod; 
LR with long downward curving alula, 

TL 300-350am, BL. 180-191 eins LT 120-150aim: TR 
63am (F 4A3~m, RR 2lpm; LR Sam; RM 37pm: LM 
asym), RE 150 X% l20um: 

Distribution: Cosmopolitaa/cosaiiropical? — urly 
record from Qld, not seen in aur collections. 
Comment. Koste’s (N78) referral of To mtecaxe tos 
vanchil morph of T de réstate as not jusafied in vicw 
of differences in morphology. im peuticular of the 
trophi, Speeilic status is retained awaiuine detailed 
SEM analysis of the trophi-of both taxa, 
Literainre: Colledge (911), Kaste (1978), 


Trichacerea muy Hauer 
FIG, tha 

Trivhoverca nins Hauer, WAR, p. 561, Tig. 86, 
Bre Locality: Tigombong, lava. 
Hetorype, Nov designated 
Pexcripiions Body al lully commmeted animal eg: 
shaped: head sheath a Series of tolds around! contracted 
head opening; head not clearly separated Trem trunk: 
no keel, but shallow groove estends dorsally tu past 
midline (my keel}; dorsal antennae is in the anterior 

art of thts groove, short, stout foot; LT stishtly kiaked 
atthe base, about 15 umes BL. RT abaut 1/5 length 
LT: TR2? 

TL (alin. BL 50-55am, LT 60. 7oam; RT loam. 
Distriburion: Cimada. Tuva, central America NT , 
‘Tas... Vie.) rare: 1I5.0°C. pH 7.4, DO 8.9 mg tl. 103us 
em, 

Literature: Koste (1978, 1981). Koste & Shiel (1980). 


Trichocered pusilla (ennmgs) 
FIG, 12:4 

Ravinlas pusities Jennings, 1903, p 338. Pigs. 8b 85. 

Trivhocerca pusilla: Harring 1913, p. 14. 
Type focaliny; U.S.A. 
Heloiype, Not designated, 
Description: Small pyriforns to fusitorm body [ef, 7. 
dixon-nurtalli (Pig. 44) and To rameri (Pig. 7:2)]: LT 
in contracted wndividuals elevated at right-angles; weak 
longitudinal striae at anterior margin} median groove 
dorsally; no suiated field, bright red cye dorsal to 
brain; right lateral wntenna shortly before foot, left at 
heginning Of Tast third of body, Trophic rami with 
robust, outwardly directed alulac, LM terminally 
weakly curved. RM shorter, rodlike, 

Total length H0-[74am: body length 69-1 15,10; LT 
to Gm: RT to WSyan,z trophi alam (F 22pm; manubris 
24/28 un: left uncus Bum: rami G/2p~m); male 60pm. 
Distribution: Cosmopolitan in plankton of lakes, ponds, 
also in moors and brackish water. Eggs often tixed te 
loricue of Brachionus species, particularly B 
angularts. Pancontinental. common in billabongs; 
105 24.5°C, pH 5.1 84, DO S.8-100 me 1, 
28-7258 enr!. 6-120 NTU. 


TRICHOCERCID ROTIFERS 23 


Fig, 12, 1, Trichacerca macera (Hudson & Gosse): lateral, contracted. 2, T inucosa (Stokes); (a) lateral, contracted; (b) 
trophus, 3. 7. mus Hauer: contracted, lateral. 4, 7. pusilla (Jennings): (a) swimming. lateral, (b) trophus: (c) fulcrum. 
lateral. Afler Koste (1978) (various authors). Seale lines: adult SOxm, trophi 10am. 


R. J. SHIEL & W. KOSTE 


TRICHOCERCID ROLTIFERS 4 


Literuivre: Bere 8 (1953), Koste (1981), Kosie & Shiel 
87), 


Trickoverea numis: (Mallet) 
FIG. 13:1 

Irichoda ratius Miller. (776, po ZBL, 

Trichacerca ratius: Harring 193, p. J04, 
Type locality. Cophehagen, Denmark. 
Holorvypes Not designated 
Description: Lortea with low or mediuae keel ca. halt 
dorsal length, may be radimentary:, wide striae-tield; 
lateral antennae in posterior tank cegion, right higher 
than lelt; RT shorter than longest substyle, “Trophi; 
fulcrum an inverted T; both manubria single-crouked, 
right more delicate, Occasionally straight; lef alula 
clongate, bifurcate, deflected ventrally, right alula 
shorter, thore rounded, animal may have reddish 
patches; red cerebraj eye: male tocless, 

TL 260-320pm, BL 150-2254m, LP 130-192,1n; 
RT to 20nm; TR 60um; SE 102X5lym; male 
60 7am. 

Distribution: Cosmopolitan, isolated o¢currences in 
littoral of standing and flowing waters, moors und 
brackish water. Common. widespread in euslern 
Austria and Tasmania, 8.0-27.0°C, pH 5.4-10,00, DO 
6.2-13.0 mg F'. 15-1080u8 cnr, 3-135 NTU, 
Comment: A-variant, TE ruts carinera (Bhrenberg) 
differs trom 7) ratius s, sir only by a wider head 
opening and higher keel; all body measurements and 
ecological ranges are within (hose of the typical fori, 
There is no evidence that this ts other than an ecatype: 
is here synonymised. 

Leterardres Koste (1978), Shicl & Koste (1979), Knste 
& Shiel (1986, 1987). 


Trichocerca roxea (Stenroos) 
FIG, 13:2 
Mustigocerca ravea Stentoos, 1898, p, 146 
Trivhoverca resea: Fadeev 1927, p. (2, Fig. 2,6-7. 
ype lecaliiy: Finland. 
Holotype: Not designated. 
Description, Dorsal margin with acute projection, af 


variable length, resembles. 7 /ongiseta; From buse of 


spine, left of dorsal antenna, shallow keel and striated 
field reach cight lateral antenna; left lateral antenna 
more posterior, retrucerebral organ with sac; sensory 
papillae in apical field: LT ca. BL, 'Trophi: falerum 
spatulate distally, LM slightly curved. distally. 

BL 260-376pm: LT W60-2)8em: RT to 40am. 


Distribution: Littoral ol lakes, moor pools in Europe. 
North America, New Zealand. Single record from 
Tas: 180°C. pH 69, 106.528 em. 

Literature: Koste et al. (1988), 


Trichocerva scipia (Hudson & Gosse) 
FIG. 13:3 


Masigocerca scipia Hudson & Gosse I8kO, p. 41, Fiz 
20511. 


Trithoewrea seipios Hacring (913, p. 104 

aon Ratiulas seyeo: Jennings J9Q3, p 322-323, Fig, 
3:50 52. (=T. jenningsi Voigt. 1957), 

Type localiry: England, “on water-moss in pools”. 
Holotype, Not designated. 

Deseriprien: Body sub-cylindrical, slightly larger al 
front, thickened, rounded posteriorly: three spines. on 
anterior Jorica margin, one occipital, two lateral, each 
continues as a low ridge onto anterior Lorlea; long low 
keel displaced to right, toe half lorica length; subsiyli 
ca. 4 toe length; mastax large. occupying > \2 body 
fength; trophi not described: conspicuous crimson 
cerebral eyespot- 

TL to 250um; BL to 148m; LT to 80. 100um. 
Distribution: England. Europe? Probably: littoral im 
habit, in pools. Recorded from three localities in 
Tasmania. 16.0-18.0°C, pH 4.97.5. 

Comment: Tt 1s unclear from the literature Whether 
species referred lo 7) ycipia are this species or Jennings 
taxon subsequently described as 7) jeaningst. Three 
populations recorded from L.. Pedder region, Tasinamia 
resemble the animal described by Hudson & Chosse 
16.0-25.0°C, pH 4.9-7,.47, 

Lirerarure; Koste er at. (1988) 


Trichacerce stylata (Gosse) 
FIG, 13:5 

Manacerca styvlatir Guase, W3l, p 90 

Trichoverea stvlai: Harring 1913, p. 105, 
Type locality... garden reservoir near London’) 
England. 
Ralerype: Not designated, 
Description: Body itreyular in form. plump, pibhous: 
integunient very flexible; contracted animal hus blunt, 
puckered head sheath. marked from trunk by distinet 
suture; no apparent ridges; lateral antennae in middle 
of trunk; cerebral eye may be papillate, LT <% BL, 
shghdy curved; RT rudimentary, ca, A length of LT, 
held appressed. easily overlooked). not substylt, trophi 
asymmetrical. 


Fig. 13.1 Trichocerca rerits (Muller): (a) typical form, tiateral; (b) trophus: 2. 7) resea (Stenrous): (a) literal, contracted; 
(b) trophus, 3. 7 seipiv (Hudson & Gosse): (a) lateral, contracted; (b) rrophus. 4, 7. seylan (Gosse): (ab) swimming, 
flown) 4 after Koste (1978) (various authors) Seale lines. adult 50am, traphy lOum 


ty R, J. SHMJEL & WL KOSTE 


TE 180-230um; BL 135 180pm; width 7pm: TR 
M-35pm: LT 45-33nm male 60um. 
Disiribation: Cosmopolitan in plankton of lakes and 
foals, where it attaches its eggs to other planktonic 
rollers, Rares Qlel. Tas... Vie, 1O.5-25.0"C, pH 7.5-8.4, 
DO 8210.0 met 440 495u8 om! 4.5-68.0 NTU, 
Lireratare, Koste (1978), Shict & Koste (1979), 


Ores species of Teichocerca 

We must stress that the Keys above (und in earlier 
parts of this series) are based on known morphology 
of routers Wwe have wdentfied, or other systemitists have 
recorded | from Australian waters, Collections whieh 
contained only single individuals not readily 
idenoliable with Known lana have been excluded. Until 
more material of these taxa bepames available they 
cannot be UWealed adequately, We are aware of at least 
three Trirhecerca taxa in logal waters Which eaanot 


be keyed successfully here, Users ol.our keys finding, 


animals whieh donot contorm to one oe tore of the 
morphological ranges for taxonumically significant 
features, in purticular the trophi, should treat them with 
auclon, Variants from the ‘nor’ should not be “shoe 


horned” into Known taxa because they ‘look a bit like’ 
the figures in an authoritative text! The wide 
distribution of such (exis, until pow northern 
hemisphere in origin, is the principal reason for the 
assumed cosmopobitan distribution of many rorifers, 
and the basis of widespread contusion in the laxonony 
of the group. 


Acknowledgments 

Thanks again to the many collectors (acknowledged 
in earlier parts) why contnbuted tauterial referred to 
in this paper. Alive Wells, N.T. Muscuin, provided 
colstructive enticisin of a draft, Thanks alse to Don 
Forth at DSIR Taupo, and La-orsri Sanoamuang. 
Deparment of Zoology. University of Canterbury 
Chnstchurch, for providing conipartive New Zealand 
muterial, The Deutschen Forschungsperieraschitf) 
comUnues lo support WE wilh microscope and 
photographic facilites, Support for fieldwork in 
Tasmami wats provided by the Australian Biological 
Resources Study, Lor-Wat Tan typed the dratl 
fMatuseript tO final form on a word-processor and 
provided corlimued competent field assistance. 


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SELECTIVITY OF MICROCRUSTACEAN ZOOPLANKTON BY GOLDEN 
PERCH (MACQUARIA AMBIGUA) LARVAE AND FRY IN LABORATORY 
STUDIES 


BY R. J. SHIEL* & W. KOSTE; 


Summary 


First-feeding golden perch larvae in experimental beakers were able to ingest only the smallest size- 
class (0.78-0.86mm) Daphnia. As the larvae grew the size of prey captured increased. First-feeding 
larvae preferred the cladocerans, Daphnia and Moina, to a similar size-class of the calanoid 
copepod Boeckella. Ten to 20 mm golden perch were non-selective for the prey species. while 
larger fry preferred Daphnia, the largest prey. The pattern of predation by larvae and fry changed 
from gape-limited to size-dependent to large-size selective, consistent with optimal foraging theory. 
Larval rearing ponds should be managed to provide small cladocerans and nauplii and early stage 
copepodites for first feeding larvae and larger zooplankters for fry greater than 20 mm. 

KEY WORDS: Golden perch, Macquaria ambigua, fish larvae, fry, prey. Moina micrura, Daphnia 
carinata, Boeckella, prey preferences, size-selective 


(Perseeitans of the Bae Sackeey ap dave O89) TAthy, 14 Ad 


SLECTIVITY OF MICROCRUSTACEAN ZOOPLANKTON BY GOLDEN PERCH 


{MACQUARIA AMBIGUA) LARVAFR AND FRY IN LABORATORY STUDIES 


by PT ARUMUGAM* & M,C. GED DEST 


Suinmary 


ARUMUGaM, Pol & Gronrs, M2 © (092) Selectivity of mierocrusticean 76oplankion by golden perch 
(Macquarie aibizne) larvie and try i laboratory stuches Yea AL See. 8) drest. W(t), 29-34, 29 May, 1997. 

First leeding golden pereh larvae in experimental beakers were able to ingest only the smallest size-cluss (0.78-0.86 
min) Daplorin, As the hivvae grew the size of prey captured increased. Kirst-tecding latvao preferted the cladncerans, 
Daphne and Medi. wcsimilir siae-chiss of the cahinoid copepod Beeckelfer, Ten to 20 mm golden perch were 
non sclecnive tnt the prey species, while larger fry preferred Duphons, the largest prey, The pauern of predation 
bv dareae aud [py chunged from sape-limited to size-dependent lo lure-size selective. Consisteat with optimal 
foraging theory. Larval rearing ponds should be managed to provide small cladocerans and nauplii and carly 
Slave copepodies fur first teeding larvae and larger voaplankters lor fry greane than 2) mine 


Koy Worun Golden perch, Macyderie cmiigua, Osh larvae, try. prey. Moma mterura,. Daphmit carina, 


Borvkella, prey preterenves, size salecqive 
Introduction 


The Australian freshwater lsh yolden perch, 
Macquaria ambigua (Richardson), ts reotinely 
spawned in hatcheries ad the larvae reared ivearthea 
ponds, which are managed to provide zooplankton 
forage for the larvae and try (Rowland 1983, |986) 
Some understanding of the diet of larvae and fry in 
these ponds his heen achieved by observing differences 
between Zooplankton communities in enclosures with 
and without fish (Arumugam & Geddes J986, (988), 
from consideration of mouth gupe and lecding 
behaviour (Arumugam é& Geddes 1987) and from 
unilysis of gut contents of larvae and fry (Culver & 
Geddes in press). However, when studying diet from 
analysis of gut contents of fish in natural habitats or 
Nursery ponds, size and’ specics preferences of fish fry 
may be obscured by turbidity (cf. Vinyard & O'Brien 
W976, Gardner 1981) and prey availability due to 
successional or seasonal patiorns of species abundance 
and sive fractions (Arumugam & Geddes 1986. 1988), 
Laboratory stucies of the prey preferences of golden 
perch will provide & better understinding of the diet 
during their Onlogeny, possibly leading Ws Improved 
pond management strategies, In the laboratory, 
combinations of selected prey can he presented under 
stindard conditions and the resulting predation can he 
presented as preference indices, 


Laboratory techniques have been used to study prey 
preferences of many planktiyorous freshwater fish (see 


* Centre of Apphed Science Studies, Universiti Pertanian 
Malaysia Campus Bintula PS. 396.97008 Bintulu. Sarawak, 
Malaysta. 

+ Pepactment of Zoology, University of Adelaide, PO Rox 
498. Adeulaide, 8. Aust, SOOL To when reprint requests 
should he directed. 


Lazear 1987 tor review), but fewer studies have been 
made on larvae and fry. Comer & Blades (1975) tound 
that the distance of Visual perception (reactive distance) 
of 8 fish was proportional ( the size of the prey, 
Lisually prey size is the major factor in determining 
preferences (Brooks 1968), but prey preference can be 
influenced by light intensity Uacabs 1978), decreasing 
prey densities (Werner & Hull 1974), escape ability 
of the prey (Drenner & MeConis 1980), prey motion. 
Visual portions of the prey (cf. Zaret 1980). and prey 
ratios and feeding durations (Gerking & Plant. 1980), 
In sdying prey preference il is necessary to consider 
bath prey size and alternale prey species which have 
different morphologies und bebaviours. Care needs to 
be given to the density and prey ratios in the feeding 
experiments. 


The objectives of the current study were to determine 
(i) the size preference und (ii) the preference among. 
some common microcrustaceans species, shown by 
golden perch (Maeyvaria ambigua) larvae and fry 
under laboratory conditions. The prey preferences of 
golden perch larvae and try would contribute further 
tw an understanding of predation by larval fish and 
would allow evalustion ot different preference models 
proposed for planktivorous fish larvae (Aarer 1980), 
Additionally. management options for the improvement 
oF golden pereh fry culture could be derved fromthis 
study. 


Materials and Methods 


Laboratory experiinenis were curred oul at the 
Inland Tinheries Research Station (IPRS), Narranders, 
New South Wales. from November ty December 1985. 


‘) PT ARUMLUGAM & M. C. GEDDES 


Amexpermental chamber in which ten replicates gould 
be run simultaneously at 20-22°C way used 
(Arumugaim 190). The chamber consisted af ten 250 
ml beakers arranged in two rows on a platfornt ina 
600 mm « 300 mm » 300 mm glitss aquarium which 
wis provided with aeration and sub-gravel filtration. 
Each beaker had a U-tube connecting the water inside 
the beaker with the water in the aquanum und 
anitaining water in whe beaker at the (5Q ml level 
AOU) wim mesh nylon sereen covered the end of the 
U tube inside the beaker to prevent the larvae, fry and 
prey items from escaping, lo each replicate, one Jarva 
or try was placed into-each heaker, A water exchange 
tate of 6 to 10 times hr! was maintained in cach 
beaker by continuous pumping of water from the 
aquarivin. This water rate ensured that the slower 
SWITMMOE prey Were NOL trapped at the nylan sereen. 


hish and zoeplanktan 

Goldet pereh of approximately 4.6, 10, 20 30 and 
38 mm standard length (SL) were used) The 4.6 mm 
larvae Were taken front an aquarium in which the eggs 
were incubated until the larvae commenced teediog. 
Fish 1-38 mm were caught from the nursery ponds. 
One larva or fry of appropriate size was placed atin 
cach beaker in the experimental chamber A plastic 
tube (diameter) 25 tam, length: 30-40) wun) was 
provided as a refuge. The larvae and try used for the 
feeding (nals were not fed during the acclimatization 
period whieh was usually less than 18 hours 

The mntcracrusticeany Moma micrura, Daphnia 
curinaie und Boeckella flavialis were either volleeted 
from the ponds at the station or culled in the 
laboratory. The total prey density used im feeding trials 
Was 90 dodividualys aa 1S0 ml ab water (600 
individuals/litre), This density was about (he maximo 
of total micrecrustaceaos ceconled ia the larval nearing 
poms (Culver & Geddes in press}, 


Prey preference trdey 
‘The prey preferences al larvae and Try were 
tlelermined using Manly’s index tor a case when prey 


density was limited (Chesson 1983), Shih Marily- 
Chesson indes lakes into account changing prey ratios 
that may occur during the feeding process if feeding 
is selective, The index range is Oto LO and the point 
ob equal preference would be the reciprocal of thy 
number of prey types. In this study, the value ts-otie- 
third (0.33) smee three prey types were used. 


Size preferences 

Three size-classes of Daplutia were used in feeding 
Lrials to determine the size preferences of golden perch 
lurvae and fry, Daplimia in the small size-class ranged 
from O78 1 104 mm, inthe medium size-class 2,00 
to 3.12 mm, snd the large class 3.68 to 5.44 mm: all 
meastired from the tip of the rostrum to the base of 
the spine (Tulle 1). Thirty Daphaia of each size-class 
were placed ina holding beaker, swivelled to mix them 
up and then gently transferred to one of the heakers 
with it fish of the appropriate size. For a particular size 
class of fish, ten feeding nals were nin simultaneously, 
euch trial Staggered at three to ten minute intervals 
For the 4.6.1mm larvae, each feeding trial was ran lor 
24 hrs! tor the larger larvae and fry. the feeding trial 
was run for (0 to 33 min depending on the feeding 
activity; trials Were terminated belore more than abouy 
30% of the prey were taken, Ac the eod of a feeding 
tial, the fish was removed and the number of Daphnia 
remaining in each size-class was recorded. Kor each 
size-class of ish, the preference index values for each 
size-cluss of Daphnia were calculated and expressed 
as Means (+ Stundarnd error). 


Species preferences 

Moina micrura Kurz. Daphnia varinaia King. and 
Boeckella fuvialis Henty were used to determine the 
species preferences of golden perch. Thirty individuals 
wf cach prey species Were used for each leeding trial 
as described for the size preference experiment. For 
DPD, carinata length was measured to the base of the 
spine: lor M. vnerura, total length was measured and 
for B fluvietis the measurement was of cephalo-thorax 
length An attempt was made to present prey of abour 


Tage |. Lenk ranges of selden perch (standard lengli) aie of small, medi and lari Dapitia, laether with the duration 


and consuniption rules af feeding truly. 


Dephtia Longtts (rim) 


Fish Length 


Consumprion Rate 


(mm Sniall Medium Large Duration (Number bh") 
4.4 4.4 O78 O86 ZH)-9 12 4 1$-5.23 24h 1.04 0.25 
W.=1K65 NIB O86 2.40) 60 4 1S-S.58 5-32 min 2-36 
19.5 22.0 080-104 316-272 4 52-3,12 10-38 min 15-48 
29.0- 33.6 O.7R-O.86 2 Ih-270) 4,22 324 {0-35 min 8-60 
$5 24414 O78 (86 DIM) 2.64 3.68%-3.90 15-25 inn RE 168 


MICROCRUSTACEAN ZOOPLANKTON BY GOLDEN PERCH (MACQUAR/4 AMBIGUA) LARVAF 3t 


Taste 2. Lenuth ranges of galden perch (standacd tenet) and Moina, Daphnia and Boeckella, together with the duration 


and comsumplion rates. of jeeding trials, 


Prey Lengths (rit) 
Daphnia 
carmaia 


Fish Length 
(mm) 


Moina 
nitcran 


Boeckella 
fluviatis 


Consumption Rate 


Duration (Number th!) 


44-48 
100-106 
V).5-22.0 
29.0-33.6 


0.35-0,50 
O.T72-0.88 
088-1 (4 
Q80-1.04 


0.78 -0 86 
0.78 -O86 
112-216 
1.36-2.56 


OA).2) 
8-17 
120-480 
60-240 


0.43-0.96 
0,96-1.20 
0.96-1,28 

112-1.28 


24h 
27535 h 
1-39 min 
1 ad min 


equal size and of a sive appropriate to the different sizes 
of the fish used in the experiment. Newly hatched 
Daphnia and Maina and copepodite stage 3 Boeckella 
were presented to the first-feeding larvae While larger 
zooplankton were presented to the larger larvae and 
fry {Fable 2). The relative sizes of the three prey 
species were similar for the 4.6 to 20 mint fry, bul for 
the 30 mm fry, large specimens of the varigus prey 
types were used resulting in the length of Daphnia 
being greater than Boeckella, which was greater than 
Moina. The first-feeding larvae were allowed 10 feed 
for 24 hrs.. the 10 mm larvae for about 3 hrs and the 
larger fry for from 10 to 39 min. (Table 2), 


Results 


Generally, the larvae and fry responded to the 
experimental chamber well and fed readily, A few of 
the 30, amd 38 mm try did not feed within the time 
duration and were replaced. Two of the 4.6 mm larvae 
died during feeding trials, with one of them haying a 
Daphnia engorged in its throat. Mortality of the prey. 
monitored by looking for dead zooplankters at the end 
of cach trial, was negligible for cxperiments up to three 
hrs duration and was less than 10% over 24 hrs. 


The duration of each feeding trial and the 
consuiniption rates in the size preference experiment 
are shown in Table 1, and the relative sizes of the three 
classes of Daphnia are presented in Fig, la. The 
consumpuion rate increased with the size of the fry. 
The 4.6 mm larvae ate 1-6 Daphiia in 24 hrs, while 
the 10, 20. 30 and 38 tam galden perch fry ale 2-26 
Daphnia in 10 ta 38 min. 


Observations of feeding behaviour were made for 
all size-classes of lish except for the 4.6 mim larvae 
which had only a low rate of predation. Most 10 mm 
larvae mitially attacked the large and medium Daphitia 
which, because of their relatively large size, usually 
escaped, On sighting a small Daphnia, the larvae 
would attack and engulf it with ease; however larvac 
continued to attack large and medium Daphnia. The 
20 mm fry would follow a Daphnia individual first 


(a) Daphnia carinata 


ee 


Boackella 
fluyialis 


Daphhla 
car{nata 


Moina 


se) misrura 


ras ~ = \ 
{2 Del 
/ | 
cin) (} aN’ | 
F Pi | 
; ee Li 
2 es 
us 
0 5 
(mm) 


Fig. 1. Relative sizes and morphology of prey used in the 
wulden perch prey preference experiments (all to same 
seale). (a) Small. medium and large size classes of Duphnia 
carinata used in the size preference experiment, (b) Size 
classes of Moina micryra, Daphnia carina and Boeckelle 
fluvialiy used in the prey preference experiment for (i) first- 
feeding golden perch larvae (4.6 mn SL), (ii) 20 mm SL. 
golden perch try and (iii) 30 mm SL golden perch fry. 


32 PT ARUMUGAM & M. C. GEDDES 


1.0 7 | | sma 
t 
(EA medium 
0.8 = large 
¥ 3 
5 06 
£& 
& ! 
5 
5 o4 
2 
a 
0,2 
a 


4.6 


Aporox. size of larvae 
and Iry 


Fig. 2, Prelerenves for different size classes of Daplinte 
connate by different size classes Of walden perch larvae: 
und fr. (Column represents mean and vertical bar 
represents a standard error Re cach mean value.) 


before attacking it- In general attacks on large Dapluiia 
by these fry were unsuccessful and. with experience 


fry usually avoided Daphnia that were too large. Pry 


of 30 mm and larger could engulf the whole size range 
ot Daphnia. 


The preferences by different sizes of golden perch 
fry for the three size-classes of Daphnia are shown 
in Fig, 2. First-fecding larvae preyed upon only the 
smull Daphuia, Preferenee for the sunall Dauphine 
decreased with increase in fry size. “Bell-shuped™ 
preference curves. with the middle size-class of prey 
most preferred. were obtained for the 20 and 30 mm 
fry. The 3% mm fry preferred the largest Daphniu and 
ate few of the smaller size-classes, 


The sizes of the ditlerent prey used in the species 
preference feeding trials for the 4.6 mos larvae and the 
20 mm and 30 mnj fry are shown in Fig. 1b. The 4.6 
tim larvae ate O-5 prey items in 24 hrs, the 10 min 
larvae ale 3-6 prey im about three rs while fry had 
a higher consumption rate (Table 2), The 10 nim larvae 
gaptured Moing and Daphnia in one attempt, while 
some ulternpty ul cupturing Bueckella were not 
successful The 20 and 30 mm fry could capture all 
three prey species with case. 


The species preferences of golden perch are shown 
in Fig. 3. The 4.6 nm larvae pretérred cladocerans 
(Moina and Daphnia) with the preference for Moin 
slightly higher than that for Daphnia whereas the 
copepud Boeckella was hardly eaten, The 10 mm and 
20 mm fry showed a similar preference lor all three 


species. In these trials. the lengths of the three prey 
species were similar (Fig, Ib). At 30 mm, the fry: 
preferred Daphnia which wus the largest of the three 
prey species (Fig. tb). The preference for Moina 
decreased with increasing fry size, lor calangids it 
increased initially and then decreased, and for Duphniu 
it varied initially and then increased with fry size, 


Discussion 


Most golden perch larvae and fry attacked large 
Daphnia on their first encounter even when they were 
wo large to be engulfed, suggesting that size is an 
important factor determining prey detection, The bias 
towards larger prey means that the probability of larger 
prey species or of larger individuals within a species 
being eaten is greater (Eggers 1977). However. the prey 
preference index is determined by a sequence of prey 
detection, pursuit and capture, Pursuit and capture 
abilities of the larvae and small fry were limired and 
so they influenced the preference recorded tor Daphnia 
of different sizes. Golden perch larvae at first feed have 
mouth gapes of only 0.5 nim (Arumugam & Geddes 
1987) and so they were able to capture and engulf only 


|_| Moina 


10 Boeckella 
ne Daphnia 
0.8 
& 
wD 0.6 
€ 
3 
Cc 
qQ 
a 0.4 
® 
ao 
0.2 
0 


Approx. size of larvae 
and fry 


big. 3. Preferences dor Moirer mieruna, Daphiia carinata and 
Boeckella fluvialis by diffetent swe classes of golden pent) 
jarvae and fry, (Column represents meun and vertical bur 
represents a standard error Jor each mean value, ) 


MIC ROCRUSTACEAN ZOOPLANKTON BY GOLDEN PERCH (MACUIMATA AATAICE a LARVAE 35 


the simallest size-class of Daphara, The middle size 
class of Dephifa was 2.8 mm long and with a head 
wilt about | nin (Pig. dial. well beyand the niouth 
gape af the first-fecding larvae, Golden perch larvae 
are clearly gape limited predators (4aret M80). The 
10 min larvae were anable tn engull the Lirgest size- 
class of Duplinier and so they too were gape limited. 
The 20 and 30. niin fry exhibited a bell shaped siae- 
preference curve, a fom associsted with size dependent 
predators (Zare 980. Seott & Murdoch 983), This 
mixiel ol predation as generally associated with 
invertebrate predators (Scot & Murdach 983), bur 
Cirilliths (973) reported low prelerence values at prey 
size exiremes in fish) Fry of Percu fevescens and 
Morotte america were shown to consistently select 
amaller Veyime aml her cooplankters than they were 
cypable of consuming (Hansen & Wahl 198t: Parrish 
& Miansral, (9). Laratetiens in atkach ability ol 
moderate sive fry along with the tnereased evasion 
ubility of lurger Daphaia would produce & preterenee 
for mid-sized Duphnia, When golden pesch fry were 
38 nun long they prelerred the largest Dupheia and 
so they were acting as $ize-selective predurors in the 
way repaned for most paniculate foeding planktivarous 
fish (Larzano 1987). Golden perch av 38 man have a 
mouth gape of justaver 5 moi and so they must have 
heen only just uble to engulf the largest! Urepiie which 
had a lengtlrof up to 3.5 min (oot including til spine! 
and a head width of about 3.5 nimi (Fig. Jb). ‘Thus, 
as golden perch larvae and fry develop they pass 
Hirnugh staves when they ure gapeimited predators, 
size-dependen} predators and predators with preference 
for largest sized prey. 


Experiments on prey prefercnee among different prey 
specres can offen be conloutded hy the diferent sizes 
of the alternative prey species. The prey presented to 
golden perch larvae had only a sniull sive range, with 
the calanoid copepod Borvkella uitennediatwe between 
the cladocerans (Fig. 0). and so the strong preference 
for cladocerans probubly relates to differenecs in prey 
beliaviour. Claduceram may be more casily detected 
hecuuse of their contrast, boty shape or puttern of 
movement (Zarei 980) and/or the calahold copepods 
may have been mure successful at eluding capture 
(Canter & Blades 1975: Drenner & MeComas 1980) 
The 10 mm Jarvae and 20 tnm fry had similar 
preferences tor all three spevies sugeesting that capture 
eMiciency wd imereuscal und that prey detection was 
similar for sinular-siaed prey from different species 
(Confer & Blades (975: Vinyard & O'Brien 19761 In 
the irtal watt the 30 mim try, the largest prey species, 
Duphnia. was preferred, suggesting that prey detection 
was the nvajor dete’minant of prey preference. 
Preferences wmong prey specics seems to vary with 
fish specws although generally cladocenans ur 
cyclopoid vopepds (Hulbert & Mulla (981; Meng & 


Orsi 1991; Parrish & Muargril 1991) scem to be 
preferred by smull fish air fry 


The resulls of the feeding behaviour and prev 
preferences experiments provide information ihat is 
relevant Wo inamaecoment al wollen peroh larval reareng 
ponds. Pirst-feeding larvae appear we byec a law 
peelerenee lor calanonl copepods compared te 
cladovenins, presumably heeause of the higher escape 
ability of the calanoids. It should be noted that the 
smallest calannids used in che present experiments were 
shige 3 copepodites and itis likely that copepod nauplii 
und-carly stage Copepodites cun be taken by the larvae. 
Rowland (1986) stated that golden perch larvae food 
Uiitially on copepad nauplii, copepodites and: soll 
copepods and cluducerans m ponds at IPRS. 
Inefficiency 1b the capture of larger calanoids and the 
limitations imposed by the sivall Wouth pape restricts 
the range of prey items that con be taken by che larvae. 
and so pond management needs Ww ensure sbundant 
small cladocerans and early stage cilanoid copepods 
Wher larvae are stocked. “The possible importance ot 
rotifers as prey for first-feeding larvae alse needs to 
be comndered. When golden perch ave 10 to 20 mm 
long they show about equal preference for Moinu 
Daphnia and Beeckella suggesting that they are 
generalist cooplinklivores, a oid strategy for a species 
from a complex and unpredictable: floodplain-river 
environment. At 30 mm, fry show a preference for the 
largest Daphma, whereas some other fish fry 
apparently select prey: thal are considerably heluw (he 
size they are capable of ingesting (Hansen & Wahl $98): 
Parrish & Margraf 1991). This ntpid developmen fren 
supe limited predation to selection of the largest prey 
is consistent with the optimal foraging Model proposed 
by Werner & Hall (1974), Furthermore Mills ui al, 
(1989) have shown that growth in age zero Percy 
fluvescens is promoted by the availability of large ste 
prey. Therefore. the management of golden perely 
rearing ponds should also aim jt providing large 
species of zooplankters for (ry 2U met and larger to 
Mitxinnze growth. 


Acknowledgments 


We thank the N-S.W, Fisheries for permyssivn (¢ 
work at IPRS. Nanmadera. the siatl of TFRS espeuially 
Stuart Rowland, Steven Thurstan und John Dinow: for 
their help and advice. IDP Australia for a Gradoute 
Fellowship to one al us (PTA.) and support to 
FIRTA, We thank Universiti Pertanian Malaysia 
Campus Bintulu and the Department of Zoolagy. ‘he 
University al! Adeltide for use af their facilities 


3 PT. ARLMUGAM & M,C CrEDDES 


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macrochirns). J. Fish Res. Bd Can. 33, 2845 2849, 

Weenrr. £. BE. & Hac. D1. (1974) Optimal foraging and 
the size selection of prey by the bluegill sunfish (Lepomis 
puicrovhirns). Fealogy 55, 1042-1052, 

ZArct. T, M. (1980) “Predation and Freshwater 
Communities.” (Yale University Press, New Haven & 
London). 187 pp. 


A NEW SPECIES OF ACANTHOCEPHALA FROM THE GREENBACK 
FLOUNDER, RHOMBOSOLEA TAPIRINA GUNTHER, 1862 


BY S. J. EDMONDS* & L. R. SMALES** 


Summary 


Aspersentis minor sp. nov. (Acanthocephala: Heteracanthocephalidae) is described from the small 
intestine of Rhombosolea tapirina Ginther, 1862 from Tasmania. Australia. It is distinguished from 
other species in the genus by the small size of the trunk. proboscis and proboscis hooks. 

KEY WORDS: Acanthocephala. Aspersentis minor sp. nov.. Australia. Rhombosolea, taxonomy. 


Nyposeetlity ol tte Reval Suererty of 8 ons. (902% GH 


WOR 


A NEW SPECIES OF ACANTHOCEPHALA FROM, THE GREENBACK FLOUNDER, 
RHOMBOSOLEA TAPIRINA GUNTHER, 1862 


by S$, J. EpMONDS* & L. R, SMALES*# 


Summary 


Tamonps, S. 4, ke SMAces. Lb, 


R, 11992) A new species of Acatithovephala from the greenback flounder 


Mivanbosolew tapoing Gunther, WO2, Trams, Ro See. S. Anayt. WG), 35-38. 29 May. 1992. 

Aspersevitiy Minol spy. Naw (Acanthocephala: Heteracunthpcephalidae) is described fram the small intestine of 
Rhamboseled taprrina Gunther, (b62 Jrom Tasmania, Australia. 1tus distinguished from orher species in the genus 
by the small vize of the trunk, proboscis und proboscis hooks, 


Key Words, Acwathocephili, dyporsenn’s miner sp. noy.. Australia. Rhamhosolee, twxononiy, 


Introduction 


About 30 small aeanthoecphalans. were collected 
trom the small intestine of the greenback flounder. 
Rhombasolea tapirind Gunther, 1862 in Tasmania by 
Dr DL, Obendorl on 16 June 1986, The collection 
contained only one quale. Theseaure considered to be 
i new species and are des¢ribed here. 


Materials and Methods 


Specimens were stained in haematoxylin hy 
conventional methods and amounted im balsam, 
Hluscrations and meastirements were anade with the au 
of wo ocular micrometer, drawing, tube und measuring 
wheel, Measurements are given in millimetres unless 


otherwise ostaicd, Where possible the range of 


medsurements is followed by the near in parenthesis. 
Type material has been deposited m the South 
Australiog Museum, Adelaide (SAM). 


Systematics 


Phylum; Acanthocephala Koelruther. 1771 

Class; Palacucanthocephala Meyer, 193] 

Fumdy: Heieracunthocephalidae Petrochenko. 1946 
Genus, Aypensentiy van Cleave. 1929 


Aspersentis minor sp. nov. 
FIGS 1-8 


Male tone specimen in poor condition): ‘Trunk 
spindle-shaped. Jengtl 1.6, maximum width in mid 
trunk region 0.28: anterior region heyring numerous 
rows of siall spines, most avticeuble and’ largest 
ventrally. Fiekl of spines extends: veatrally: for abour 
¥ quarter af hody length. but less extensive dorsally, 


“South Austratiad Museum, North Terruce. Adelaide, 
S Aust, 5000 
™ University College of Central Queensland, Reekhatprm. 
Olt 4702. 


Remainder of trunk, including peniuil region, without 
spines. Proboscis (alinost completely extended), set 
at angle to trunk, eylindrical ter clavate with armed 
section 0,23 long and 0.12 wide, bearing 14 rows of 
7-8 hooks per cow, Dorsal and ventral hooks differ 
murkedly in size and shape. the latter being mach 
larger. Short unarmed, truncated neck about O15 long, 
Two ovoid testes tandemly placed. Cement glinds, 
pyritorn) and pressed closely together but number not 
clear, Proboscis receptacle double walled. 0,42 loag 
with ganglion placed near base. Lemmisci about as long: 
as Teceptacle. Male aperture subterininal, 


Female (based on 10 mounted specimens): Truak 
spindle-shaped. length 2.4-4.1 (3.2). maxunuim width 
0,37-0.75 (0.54) in mid-(runk region: field of spines 
extends for about a quarter of body length ventrally. 
less extensive dorsully. a lew tiny cuticular spines 
present at posterior end of Some specimens. Probosets, 
placed at slight amgle i trunk, cylindrical to clavate 
0,24-0,32 (0.26) long 0.10-0.17 (014 wide and armed 
with 14-15 rows of 7-4 hooks per row. Ventral and 
dorsally placed hooks differ most nuticeahly in size 
und shape (Figs 1.2). Longest ventral hooks (third in 
row) measure 0062-0080 (0.065). longest dorsal hooks 
0,030-0,035 (0,032). Unarmed truncated neck 0.15-0.28 
Jong. Receptacle. maximum dimensions 0.52 long x 
0.19 wide. double walled, with ganglion lying neae its 
base: Genital complex tong. extending in most 
specimens about half length of trunk. Embryonated 
epus.. Slender O.A8-0.077 (0.071) * 0.012-0.019 (0,015) 
with prolongations of middte shell. Sniull terminal 
papilla (invagioule) present in most specimens. Femule 
aperture almost terminal. 


Host: greenback flounder, Rhewmboscled nipiring 
Cilinther, 1462. 

Location: small intestine 

Locality; Tasimanta, Australia 

Type specimens: Holotype mule: SAM 
Paratype female; SAM V4I5I 


V4150. 


36 $8. J. EDMONDS & L R. SMALES 


i) 


co 


w be ; 
« 6 
‘S 
& & 
——gp ‘ 


Figs 1-8. Aspersennis ninor sp. nov. O from Rhombosolea wipirina. |. Proboscis hooks, ventral row. 2. Proboscis hooks. 
dorsal row. 3. Praboscis. 4. Whole mount. 5. Cuticular spines, dorsal surtace 6, Cuticular spines, ventral surface 7. Vaginal 
region extended.. 8. Egg. Scale bars: Figs 1, 2. 5 & 6, 50 pm, Fig. 3. 0.2mm, Fig. 4. 0.5mm, Fig. 7, 80mm 
Abbreviations: b w, body wall: g p. gonopore: u. uterus: y. vagina. 


ANEW FLOUNDER ACANTHOCEPHALA 37 


Discussion 


The speanncns of Aspersendis dinar sp. lav. ost 
Closely resenible 4. avserinus. van Cleave. 1929 fron 
Antarctic fish and subsequently redeseribed by 
2daltowieckt (81) from hosts fram the Antarecde and 
sub-Anarctte islands, 4 disiriins wits alse reported 
by Joyeux & Baer (1954), Edmonds (1955, 1957). 
Golvan (1969), Zdzttowieck: & Reokosz (1986) and 
Zdznowieki (1990), Similarities include the number 
of rows Of proboscis hooks, which fir A. quvtrinus is 
13-16 (usually: 14) rows of FH) hooks and fora. viiner 
4-15 rows of 7-9, und to the size-al the embryapheres, 
witieh foc the former is (.060-0.088 2 0019-0025 and 
for the latter 0068-0077 %* OAN2-Q0M. The specimens 
differ most notably in the sizeof the trunk, which is 
4.4948.5 lor female A, anvreinus as compared with 
24-4.1 for female 4. mdier, the proboscis length, 
which for the former is O.51-0.73 und the Latter 
024-032. und the length of che largest ventral hook, 
012-015 as compared with 0.00-0.08. 


Golvan (969) vonsidered Eehinorhynchus 
meseirivichis ULinstow, 1892) to be a senior synoayin 
OFA Nas. a deteniination Unit wis followed by 
Agua (985) im his classification of the 
Acaunthoegphale, Amin, however appears lo have 
overlooked Zdzitawiecki’s (981) redeseription of A. 
ausTrOus based On tore than 1350 specimens collected 
Jrom South Georgia and the South Shethatids. 
Advitowieckt (98)) concluded that Aetinerhynchves 
(veaisn late) mevarAynchus, as described by Linstuvw, 
was “impossible (a Wentify with any ingre recently 
Jeseribed species’, Subsequently Zdzitawiecki & 
Rokosz (1986), then Zdaiiowierki (1990), on the hasty 
of the te-exuimination al eld muterial and the collection 
of new mMateriul, comirmed the validity of A. 
Mewar hiveclan ULiastow, I892) sya Bo meparhyec tis 
Linstow. 1892 Hee A onegarhyechus sense Galyan, 
S960. E. inevarhynches licks body or cuticular spines 
und fo sisyonetey is deserihed tor its proboscis hooks, 


4. anwieiws and A. mor diter trom atin both these 
characters 4 miner also differs from Helene 
thocephalus peltorhamphi (Baylis, 1944) and 4. 
kureaut Dots, (964, bot of which lack bady spines. 


In his redescription of A. cusirinus, Zdzitowieeki 
(1981) commented on the distribution of spines over 
the trunk, Because be found specimens in which tiny 
spines. offen embedded in the cuticle, were present 
erther over the middle and/or posterior regions of the 
trunk as well as larger more obvious spines on the 
aotenor trunk, he aceordingly proposed an emendation 
of the generie diagnosis, The armature on the distel 
pact of the body is particularly difficult to deterniine, 
especially th the material iy contacted or methods of 
preparation have rendered spines hard to deleet. The 
specimens of A. miner deseribed here were tound i 
have spines over the anterior trunk and spursely 
seultered, tny spines on the postefior of some 
specimeny, 


Adetowieeki (981) found considenble differences 
inthe size ol specemens trom A aiysries popubitions 
collected from South Georgia and the South Shetlands. 
He found that specimens fram South Georgin, where 
Water temperatures are higher, were on average 30% 
larger and their probyseises, eeceplucles and lenmise 
10-20% lurger (Table 1}, He suggested thar these 
differences in body dimensions nnught depend on 
diferent environmental conditions, (for example-water 
temperatures) where host populations o¢geur, Similar 
reasoning, cannol be used ro explain the differenee in 
size between the Tusmanign and South Georgi 
specimens since the annual water teniperature around 
Tasmania varies from 10-20°C and around South 
Georgia fronr 5-10°C (Plate 3 The Times Atlas of the 
World}. and the Tasmanian specnnens are not larger 
but sruuler, They are also smaller than A, aasrrdias 
described from Noronha evarobrancie Richardsen 
from Heard Isltad (Edmonds 1953), Although 
considenble variation in measurements has hee lound 


TARLE | a cemnprrivon of femele bedy measniviiedtys af Aspetsenliy austtinus Fey Cleese, 1929 fro the South Sherkanas 


and Srauth Geerua, dnd A. Taner sp. hot from Tavera, 


South Shetlunus 


494-6, 42(5, 79) 
116-1, 79) 49) 
prabascs lene 051-0, 66(0, 59) 
probosces wilt 0.29-0, 3240, 30) 
dorsal hoak length Ghasmuunn 0054-0, 064¢0, 060) 
venti Wook bongth (iaaxtmmand 0, 119-0, 1740, 1261 
neck tenet (.17-0.0810, 126) 
eye 0-060 0,088 
OOO O25) 


Trunk length 
trunk width 


book Ulsppsitian 


W416 rows ol FLL hooks raw 


South Georta Tasmania 


A.94-8 54/7 25) 
1.092.091, 73) 
67-0), 70. 70) 
(244). 4600 32) 
CHO, (64510 062) 
QO 1A. 14000) bd) 
O22, 5100.27) 

0.07 1-0,.087 

220, UZ) 025 


ZV 4A) 

(31950. 54) 

(24-0. 5200.8) 

QO, WOOL TAY 

(08 (LOSS(0.042) 
0062-1), OR(0_065) 

0, 12-0,25 

V.QO8-0, 077 
SOIL OTE 

I4 rows vit 7-9 hanks! raw 


38 §, J. EDMONDS & L. R. SMALES 


between populations of A. ausfrinus, measurements 
of A. minor clearly fall outside their range. Moreover 
the sinaller size of A. minor goes against the trend, 
established for A, avesirinus, that specimens from 
populations from hosts of warmer waters tend to be 
larger than specimens from cooler waters. Therefore 
A, minwr is considered sufficiently different to be a 
new species. 


A re-examination of A. ausfrinus from Heard Island 
shows that in addition to the spines on the anterior body 
surface reported by Edmonds (1955), small spines are 


also present on some other regions of the wunk. a 
feature previously overlooked. 


Analysis of the ratios of male to lemale A. austrinus 
sp. (Zdzitowiecki 1981; Zdzitowiecki & Rokosz 1986) 
has shown that there is both a twofold predominance 
of females over males and a difference of preferred 
location in the host, males preferring the posterior half 
of the smal] intestine and females the large intestine. 
A similar difference may explain why the ratio of mules 
to females of A. miner collected from the greenback 
Nounder was. 1:30, 


References 


AMIN, O. (1985) Classification, 4 DWT. Crompton & B.B. 
Nicol (Eds) “Biology of the Acanthocephala” (Cambridge 
University Press. Cambridge) 

Eomonps. S. J, (955) Acanthocephala collected by the 
Australian National Anlarcuc Research Expedition on 
Heard Island and Macquarie Islund during 1948-50. Trans. 
R, Soe. S. dust 78, 141-144, 

(1957) Acanthocephala. BAN. ZARB. 1929-1931. 
Reports - Ser. B, (Zool, & Bat.) 6(5), 91-98 

Goivas, Y. J. (1969) Systematique des Acanthocephales 
(Acanthocephala Rudolphi, 180, Premi¢re partie. Lordre 
des Palacacanthocephala Meyer, 1931, premier fusicule, la 
super-famille des Echinorhynchoidea (Cohbold, 1876) 
Golvan et Hou. 1963. Mei Mus. natn. List. nat., Paris, 
Ser, A, 57, 1-373. 

Joyrux, C. EB. & Barr, J. G. (1954) Cestodes et 


Acanthocephales recoltes par M- Patrice Paulian aux es 
Kerguelen et Amsterdam 1951-1952. Meém, Inst. Scient, 
Madagascar. Scr. A. 9, 23-40, 

THe Times Attas OF THE Wort, (1981) Comprehensive 
Edition (Times London), 

Z2DAITOWIECKI. K- (1981) Redeseriplion of Aspersentis 
austrinus van Cleave. 1929 (Acunthucephata). Acre, Peresit. 
Pol, 28(7). 73-83. 

(1990) Re-examination of five Antarctic and sub- 
Antarctic digenean and acanthocephalan species from 
Professor Szidat'’s collection. [hid 35(1), 31-36. 

____ & RoKkosz, B. (1986) Prevalence of acanthoce- 
phalans in fishes of South Shetland. (Antarctic) [, 
Aspersentis austrinus Van Cleave, 1929 and remarks on 
the validity of Heteracanthocephalus hureaui Dolltus, 965 
Thid. OT), lal YL, 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


VOL. 116, PART 2 


NEW RECORDS OF SUBFAMILIES, TRIBES AND GENERA OF 
BRACONIDAE (INSECTA: HYMENOPTERA) FROM AUSTRALIA, WITH 
DESCRIPTION OF SEVEN NEW SPECIES 


BY A. D. AUSTIN* & R. A, WHARTON** 


Summary 


Three subfamilies of Braconidae are recorded from Australia for the first time with the description 
of the following new species from Queensland: Ecnomios stenosoma sp. noy. (Ecnomiinae), 
Histeromerus clavatus sp. noy. (Histeromerinae), and Meteoridea anic sp. nov. (Meteorideinae). 
The tribe Muesebeckiini is also specifically recorded from Australia for the first time with the 
description of Paroligoneurus pallidus sp. nov. (Ichneutinae) from the Northern Territory, 
Queensland and New South Wales, as is the genus Chrysopophthorus Goidanich, with the 
description of C. hageni sp. noy. (Euphorinae) from South Australia. A new species of the 
Australian endemic subfamily Mesostoinae from South Australia and Victoria, Mesostoa kerri sp. 
noy., is described, as is a new species of the little-known genus Calohelcon Turner (Helconinae) 
from central Australia, C. dangerfieldi sp. nov. The diagnoses, biogeography and biology of these 
taxa are discussed and notes are provided on the euphorine genus Stenothremma Shaw, previously 
thought to be rare within the Australian fauna. Keys to species are provided for the genera 
Histeromerus Wesmael, Mesostoa van Achterberg and Culohelcon Turner. 

KEY WORDS Hymenoptera, Braconidae, Ecnomiinae, Histeromerinae, Meteorideinae, 
Euphorinae, Helconinae, Mesostoinae, Ichneutinae, Muesebeckiini, new species. 


Traps ob tie Rove! Sector ofS Aen (OS, Hay dy, db at. 


NEW RECORDS OF SUBIAMILIES, TRIBES AND GENERA OF BRACONIDAE 
(INSECTA: HYMENOPTERA) FROM AUSTRALIA, WITH DESCRIPTION OF 
SEVEN NEW SPECIES 


by A.D. AUSTIN® & Ro A. WHARTONT 


Summary 


AUSTIN. A. Do & Wears, R.A, (1992) New records of subfamilies, rribes and generd of Braconidae (Insecta: 
Hymenoptera) from Australi, with deseniption of seven new species. Trans Ro Seo S$ dase W6(2). 41-63 20 
May, 1992. 

Thiwe subhaiilies of Braconidae are recorded fog Austratia fon the frst te with the description of tte fotlew itis 
Hew species from OQveensland: Feradeioy sterroyenea sp, gay, (Renominae). Histeranieruy efaveties sp. Ov. 
(Histeromerinae), and Mereornleat anic sp, wey, (Meteorideinac), he tribe Muesebechiing is alse speerfieitly 
reeorded fon Australia foc the first Gime with the description of Aurolipeneurus palkdus sp tow (Tehnetitinue) 
from the Northern Territory. Queensland and New South Wales. us is the genus Chrwopophthoruy Goidinich, 
with the deseriprion of C Aagenr sp, nov, (Eupborinac) from South Australia, A new species of the Australian 
endemic sublunily. Mesostarmae from South Auwimiieaund Virwria, Meyostod Kerrd yp, nov. os deagiibed, ds 
is W Mew Species vt the little known gents Calohelcon Turner (Heleoninve) from central Australia. (2. dangerfieldt 
ap now ‘The diawytoses, bioweorrmuphy and biology of these taxuoure discussed ard notes are provided on the euphorine 
genus Senotiremnme Shasy, previously thought to be rare within the Australian Raina. Keys to species ace proved 


for The Gonnth AN ieramerus Wesmact, Méeyostoe van Achtorhers and Calaheleon ‘Varner. 
a e 


Key Workts 


tiyvmenoplery, Borevnidac, Eqnomiinag, Histeromerngc, Meteondemac, Buphorliae, 


Helconinae. Mesostoinae. lehneutinae, Mueseheekiiii, new species. 


introduction 


The Braconidae is one of the largest families of 
parasine Hymenoptera, Its members ure ceto- and 
endoparasitoids of a wide range of msec hosts, in 
particular Jarval stages of Lepidoptera, Coleoptera and 
Diptera. Although the family has been extensively 
studied chewhere, the tauna of Australia remains 
poorly known, despite the existence of a relatively large 
number of cademic subtamilies and genera. Indeed, 
the majority of subtamilies in Australia have nor been 
revised, and most genera and species aré known only 
from their original early desenptions (the majority 
described prior to the 1920's). Recent taxoaumic work 
onderaken by us on the Microgastrinae and Alysiinae 
(Austin & Dunyerfield 1992; Wharton in prep.) 
indicates that for these two subfamilies. less than 10% 
of Australian species are described. and this is likely 
io be the general situation across the whole family. 
Until the Braconidae ave better surveyed at the generic 
level. quesuions regarding the evolution and biegce- 
graphy of the Australian fauna cannot begin to be 
addressed. Here we make a contribution in this regard 
by reporting on a number of significant taxa that were 
discovered when sorting material in major Australian 


~ Department af Crop Protesuion, Waile Campus, University 
oF Adelaide, Glan Osmond S. Aust, 5004. 

““ Department ol Butomology, Texus Ad Mo University, 
Colleve Siation, TX 77843 2475, USA. 


collections, in particular the Australaa National Insect 
Collection, Canberra and the Department of Primary 
Industries Cotleckon, Brisbune. Seven species are 
newly described, three representing the first record 
tron) Ausuatia of ise subfamilies Eenomiiae, Hister- 
omerinae and Meteorideinue, and one representing the 
first description of aa Austratian species. from the 
ighneutne tribe, Muesebeckimi. The relationships. 
diagnoses and biogeography of all taxa are discussed 
and noles ure provided on their biology where avail- 
uble. Keys to species are provided for the peneri 
Histeromerus Wesmael, Mesostoe yan Achterberg and 
Caloheleom Turner 


Abbreviations for collections are: ARTIC, American 
Entomological Institue. Gainesville; ANIC, Australian 
Nauional Inscet Collection, Canberra; BMNH, The 
Natural History Muscum, London: CNCI, Canadian 
National Collection, Ottawa; HNHM., Hungarian 
Natural History Museum, Budapest; QDPI, 
Queensland Department of Primary Industries. 
Brisbane: RMNH, Rijksmuseum van Natuurlijke 
Historie. Leiden, TAMU, ‘Texas A & M University. 
Callege ‘Station: USNM, United States National 
Museum, Washington, D.C; WARI, Waite Agricultural 
Research Instiute, Adelaide. Terminology for 
morphology and sculpturing pattern follow Gauld & 
Bolton (988) aid Wharton (1977, 1986). respectively. 
while that for venation fallows van Achterberg (1979) 


Nz AT? AUSTIN AR 


Treatment of species 
Subfamily Eenomimay van Achterhera 


Commnens” The systemilic position af the only 
included venus, Bevery Maso, bis been the subject 
of some dehyte: Misen (1979) noted us superfictal 
feseniblance to Micragastringg, but excluded it un the 
basis of the sclerolined distal fidial seqtor al the frre 
wing and the arrangement of wbloninal spiracles. 
Mason (1979) emphasized seven other vhuraclers which 
suggested a relanonship with Cryits Haliday. and thus 
included Eenriivay iy the Orgilinae. Van Achterbent 
(985, IYSR) pul Eeromins v0 its own subfamily. and 
suggested thar it is best placed halfway between 
Cheloninwe and Nevneurmnac. Yau Achterberg (1985) 
excluded Eenamios from the Ornilinae because “the 
Orpilinae lack yeut -SR of fore Wing, have myrgmial 
cell long and rather narrow, presence of vein CUID of 
fore wing. convex face. large hind coxue. and small 
plical lobe of hind wing” The fee is actually as convex 
in Kemomios an as i cnany Crgius. The remaining 
features. are as suitible forunguing against a relationship 
bemween Acieros and Cheloninae us they ure for 
ierguing avainst the relationship between Ecrewnilos and 
Orsrlrwe. Furthermore, Eenenmios lacks three at the 
(hur synapomorphies proposed by van Achterbere 
(1984) for the Cheloninac-Microgastrinae Hneage. The 
placement of Eenymniny is thas sail unsettled, as noted 
ty Quicke & van Achterberg (1990), We prefer to treat 
i provisionally as intermediate between Orgilinae and 
the chelonine-meroeastrine lineage based largely on 
wing venation patterns. However, lhe presence of it 
transverse postecutellat plate wn the species described 
below opens up the possibility for relatuonshipss wih 
lhe Alacinae nd Kuphorinac, where similarly reduced 
venation accurs. 


Mason (1979) and van Avhierbers (1985) provide 
detailed lists of characters defining and dilferentisting 
fenonnoy. The naterial available to Us, representing 
at least faur species, lureely color Witt the original 
deseriplion and redeseription. butthere are important 
exceptions. ‘The Jollowmy remurks are theretore 
provided to supplement previously published 
information. Maxillacy palps are interspecifically 
variable. cither 4- or 5- segmented. Antennul seginents 
are also variable in number both imtra- and 
interspecifically. most Nagellameres have placodes in 
Iwo ranks, and the apical flugellamere is spinose at 
the up. Although van Achterberg (985) states thar the 
apical antennal segment backs an apical spine, Mason 
(1979) correctly notes thal il is acutely pointed (Fig. 
§), und tly os true. far all species exumined 

Van Achterbere (I985) described the pronotum 


dorsally ay having w late, deep, Transverse pronape. 
However. Lin is not ihe sae stvucture us the isolate 


A. WHARTON 


pit tound. for ¢xemple, in same species of optines. 
alysines, and rogadines. itis actually part of a comply 
scrics of pits or- depressions firming a crenulate sulcus 
Which sepunites uw weakly nsed posterior median aren 
fromm weakly raised antero-lateral areas (Pig. 1). ‘There 
is considerable yarialion in pronotal sculpture pnd the 
pattern should prove useful an delinimg species or 
spewits-2eoups.. Both Mason (1979) and van Achterberg 
(1985) desenbed uw conspicuous projection in the middle 
of the anteri-lateral margin of the proootum (in larcral 
view). This feature. though very well developed in & 
papuensixs Mason, is weak or virtually absent in some 
nother species, and is thus of questionable value tor 
generic or sublumily chaructensanion. In enemies the 
“projection” iy actually ihe ventral portion of an inden 

jatiomin the thin, antertor murgin of the pronotun. 
The occipital carina Fits inte this indeniation when the 
head ts retracted. The structure ts thus different in bath 
appearance and function From the anglar projection 
of the margit seen iy some species ef Crysis, 


The -carinave untero-lateral margin of the mesiinotal 
disc has some potential fur charactermsing higher tnx 
il accurately deseribed. Mason (1979) correctly notes 
its presence in the type species, bul wan Achterberg 
(1985) claims that i) is absent in Fromt al the tegula. 
Itis well developed and complete trom the base of the 
notadl to well past thetegula in all Australian species 
we eXamned (Fig. 2, arrowed), The transscutul 
articulation (Fig. 1, arrowed) i8 alscr distinet along the 
anterior margin of ihe scuto-seutellar suleus im all 
species, hut indistinct laterally, In & papwensis, the 
scutellum lacks a Wiedian postscutcllar plate (rransverse 
sculellar depression seasa var Achterbere). In the 
species described helow, however, & small one is 
present similar in shape and position to chat im 
Sigalphus bicalor Cresson and some Centistini. The 
bicuringle median portiod of the metanotuny 16 also 
similac to that of signlphines and many centistines, bul 
this patiern ts repeated in several other braconid 
subfamilies. A brow prapodeal urcela ix present 
posteriorly in all species, but variously shaped. and 
often largely obscurcd by nugose sculpture. Mason 
(1979) noted ridges on the dorsi) surface of the hind 
coxa in his deseription af E, papuensix, AU species 
haveat lousta singhe ridge in this position, suggesting 
4 Synapomorphy fir the genus. ‘The venation of the 
short, broad fore and hind wings (Figs 3, 4) is alsa 
diagnostic for Ecnemins, hax been udequately 
characterised by previvws authors. and is esscatially 
uniform mall species, I-SR varizs in length amung 
species, and its presence may not be sufficiently 
reliable for subtamilial diagnosis. 


The new species described helinw as a predictable 
range extension for Ecvomies from Papua New Guinea 
imo nuMbeTD Queensland. We have had un opportunity 
to eximing i single female from Somalia (CONCH 


NEW RECORDS OF SUBFAMILIES, TRIBES AND GENERA OF BRACONIDAE 


4a 


Figs |-5, Kenomtios stenusama sp, noy,, @ holotype. 1, dorsal view of body ((ransscutal articulalion, arrowed); 2. lateral 
view of body (carinale antero-lateral margin of mesonotal disc. arrowed): 3. tore wine; 4. hind wing: 5. distal lagellomeres 
oF antenna. Svales: Figs 4 = 0.5 mm; Fig, 5 = 125 um. 


répresenting an undescribed species and seven 
specimens from Queensland and Northern Territory 
(ANIC). which differ primarily in volour from E- 
papHensis and the species described below, Based on 
this distribution pattern, Acremios should eventually 
be found tin India as well as other Inda-Australian 
localities. 


Eenamios stenusoma sp. nov. 
FIGS J-5 


Matertal exeniinedd. Halowypes 2. ANIC, Queensland, 
"IS 16-5 144.59 B 14 km W by N of Hope Vale Mission Q 


TAQ May (981 1, D, Naumann ex ethanol’ Paralypes: 
Queensland, 3 9 -G. Rex Range Lookout. via Julauten. 
16°30°S. 145°25'E,9.x1-2.511, 1981, malaise trap (QDPI); 1 
9.1 or, 1I5°087S, 145909’, 3 km NE Mt Webb, 1-3.x.1980, 
J, C. Cardule, ex ethanol, collected at light (ANIC. TAMU); 
1oo. 1547'S, 145*14°E, Shiptons Flat, 16-18, 1981, 1, D, 
Naumann, ex ethanol (WARI), 1 o, 1717S, 145°20°R, 
Millstream Falls Nat, Pk, 24-25.v.1980, 1. D. Naumann & 
dC. Cardale. ex aleohal collection (TAMU), 


Female 

Head, \,05 1.15 broader than mesonotum (between 
tegulue), face 1.45-1.60 x wider than high; malar 
sulcus restricted to a weak impression extending less 
than half distance from eye to mandible: malar space 


14 AD AUSTIN & R 


about 2 © basal Width of mandible: mandibular teeth 
minute, dorsal tooth nearly 2 * Jonger than Yeatral 
tooth, clypeus with shallow widely spaced punctures; 
head otherwise hugely smuottoand polished. wath fine 
hair punetores: ocelly similar tok. papiensix, thiugh 
Jerald ocelli slightly (nore distant from eyes; antenna 
28- to 30-sewmented, 


Mesusome $85 2,08 % Janger than high: wedth 
between tegula 0.93-L.15 * height; pronowm in dorsal 
view with (hin medially. cmarginate anterior border and 
sinall posterior median plate and transverse crenulate 
sulcus curving anteriorly im front ol median plale, 
indentation and associated angular protrusion along 
antero-vetiiral migrgin ol pronotune weak. barely 
evident in some specimens: lulerdl margin OF mesonotal 
dise sharply carinate. the sculpture extending from base 
of notaut beyond legula to postenor margin of busul 
wing pad: dise unitorntly and densely short-sctose and 
finely punctate, notuli narrow, very shallow, erenulabe- 
rugulose, converging postoriarly to form i large 
crescenteshaped, ruguilose paleh as i &, paputensiy: 
apical margin of scuictlum sculplured medially (Pig. 
L) giving the appearaove of ¢ transverse postscutellar 
plate; seuloseutellary sdleus, paraseutellar fields. 
Menino, mesopleuron und menipleuron us in é 
papeensis, bub withshghtly weaker sculpture, aalerior 
portion of propodeum ragulose, median longitudinal 
carina usually absent. lateral longitudmal cartnuc 
usually Weak, sometimes slmost indistinguishable 
amongst background sculpture, smaller posterioc 
deeliveus partion more finely sculptured, marked 
anteriorly by broadly bowed transverse earma: tind 
cuss Shorter (han peoole. wath a strong diagonal carim 
dorsally extending nearly (rom base to apes and with 
2 shorter weaker curinae adjacent to this, 


Fy wir, (i 3) As tor A. papuensis except as 
lallows: 243 M varlible. short usin A. papaernsix 
in one paratype brit approximately equal tn length to 
25R—M urother speciumeds. ISR+M acining trom |-M 
heal partsbgma, With SR nearly absent in several 
specimens: 2-CUl 2,20+0.35 » longer than } OU; 
2-14 often represented by a short tubular spur at 
evtreme buse. otherwise almest completely indicia. 
guishable 


Melayews, Petiole With length slightly shorter than 
(O75 O85 X) apical width; melasoma shape, sculpture. 
und setal panierns otherwise as in E. papacass, 


Colowr Lightyellow-brown, ovellar triangle, most of 
T20-3 und at leastapical margins uf subsequent tera 
darker than rest of body: scutellan region often suffused 
Wit browis anlenna aid apiges of Sth tarsal segments 
browns mnouliparhs while, except lor tip of mandible 
which is reddish 


Leagil. 22-2. on. 


4 WIARTON 


Male 

Besentiilly as for fernale except ocellai field anid petiole 
possibly a litle broader bul insulfictent midterial lr 
udequate comparison, 


Biology. Unktiown. 


Diagnosiv: This species is readily wWentihed on the 
basis af is pale coloration. Both &) pepimensiy and an 
undescribed species from Queenshand are largely dark 
brown to black, &, seaesead also has a narrower 
body, wath the fiead somewhat broader than the 
mesonotum, The mexonotum: is broader thin the heated 
1B, PUPHenses. 


Sublimmily Iohinentnae Poerster 
Tribe Mueseheckiini Mason 


Crmments. Mason (969) pluved they tribe in the 
Ichneurinae. Me included six genera, three of whieh 
were transterred from the Microwustrinae. ‘Phos 
placement was overlooked by Shenefelt (1973) bur 
aecepted by other workers (0.2. Marsh 178. van 
Achterherg 184) except Tobias & Betokubytskiy (98ly 
who later transferred the Muesebeckiini tthe 
Mirucinse on the basis of host relationships anil 
similariey in reductions of venation, palp seaments. and 
male penitalia (Tobias 1986; Belokobylskiy 1989) 

The relationship between fedinentes Nees yon 
Fsenbeck vind the Muesehechiini is based largely on 
(he puture oF the sharply bent basal vei (eM) of the 
lore wig, & feature not shared by olfier ichneulines 
such as dehaentidea Ashmead and Preverops Wesmacl. 
In Mires Haliday. | -SR generally Toros a sharp angle 
with the parastignu, but the rescmblince between this 
and the condition im ichneutines ts super fiend, and dies 
Hot suppert the inglusion of Meusebeekiin jn 
Miraeinie. Mason (969) lists six other churucterstics 
Shared by dedmenies und Muesebeckiines, but rane of 
these is unique to this clade The Muesebeckiini lack 
the most significant synapomorphy of the Miieragastwine 
aroup ol subtamilics (i) whieh Miray belongs); the 
placement of the spiracle in the membranous eeral 
poruood the lirst tergum, Additionally, the Magello- 
meres are not fixed in number as they are in Micro- 
sustrinae and Miracinmie. On the basis of sprracular 
placement, we include miracines within she mieropas- 
trine group. and phice Adeliinae and Ivhneutinae (the 
latter Including Muesebeckiini) with the Neoneurinae 
und Chelomimie as a sister-proup of this elide 
Meusebeckiines are most readily tecowinzed by the 
venation puller vf the fore wing (bin. 7), 

‘The tube Muesebeckiini ws represented ui Austria 
we the widespread genus Parelignveuras Mueseheck 
One species is deseribed here. bur others will 


SEW RECORDS OF SUBFAMILIES. TRIBES AND GENERA OF BRACONIDAT. 45 


undoubtedly be found with mare intensive collecting. 
As interpreted here, Pareligonewruy is ia lire genus, 
wilh numerous undescribed species inthe New World. 
De Saeger (1944), workitg on the tropical West Atnican 
fauna, has deseribed the largest number of species. but 
propodeal differences sugpest thal ut least some of hes 
specics belong elsewhere. Nixon (1965) previously 
noted the occurrence of Pardlizoneuras ja Queenstand, 
but did dob desenbe any species. Risbee (1951) 
described a species reared fron) Agromyzidage in 
Senegal. but this is an opine. Known busts of true 
Paroligoneurus are leal-muining Lepidoptera. 
Mugsebyck (1931) noted. the close resemblance between 
Olvoneyrns Szépligeti and Purotigoneurus im his 
original description of the latter. He differentiated the 
two solely on the basis of the relatively bare eyes and 
reduced number of Mupellameres in Parolivaneurus, 
hut noted that P jelinsond Mueésebeck had a few 
seullered eye hairs, Subsequently, De Saeger (1944) 
described some species of Purnligeneuras with 
seattered eye hairs und Belokobyiskij (986) deserhed 
W species.of Olizonenrns with hairy eyes and relatively 
few (21-23) flagellomieres. Mason (1969) did not discuss 
eitlier venus when he transferred them to 
Mueseheckiini, bat presented a key te genera in whieh 
he sepurnted Ovmonenrny and Paroliganearus on the 
basis OF whether or not the eyes were hairy. Belokab- 
ylsky (O86) noled Hil previous characterisations were 
Inpdequate for distinguishing these two gener, He 
therefore added a clypeal character, und modified the 
tradimonal eye and antennal diagnosis, The mocdentely 
hairy eyes of the species deseribed below further 
cimphusize the weakness of this character state for 


separating Olfeonenrus and Parvliwoncurus, and we: 


suigsese Una at shoulul be abandoned entirely. Although 
the type species of Oligoneuris is very distinetive, with 
its large size, relauively large number of Magellomeres, 
curinate propodeum and petiole, and broad second 
tentum. other species which bave been assigned to 
CligoneHrus possess only one ov two ol’ these traits, 
atid otherwise resemble the type species of Paraliy- 
wreuras. \ revision of the large Neotropical fauna is 
needed helore the genera can be adequately defined: 
Until this can be uecomplished. we believe that the 
best character for separating the two is the propodeal 
sculpture. admittedly a weak feature. The elypeus is 
evenly rounded in Oligorenrus concolor Szepligeti and 
£ jeohiseni Muesxebeck,, and thus cannot be used for 
separating the two genera. Members of the distinctive 
Holurctic species-group with anedially protruding 
clypeus more closely resemble, Farolivonerrtis: than 
Oliyicurus, based om propodeal sculpture and. the 
shape of tergite 2, The placement of two such species 
in Olfgoneurus (Bulukobylskij 1986) thus needs to be 
reassessed 


Pigs G 1b Porelgenenras pallides sp, nov. O holutype. 6, 
antenna: 7 dere wings 8. hind wang Sh anterior view ol 
fetid) (0, Th and 1243 of metusoma. 1h. Pak eaweurie 
pallidus sp. nev. 2 parutype, pasterstotsal view ot 
melasma showing medial pit in TA and T7 Scales: Figs 
4-4 — 0.5 mm Figs 9-H — 250 jun, Abbreviittions tor 
Fig, G; length of antenna relative to body fi = heits! oy 
= mesasonias wil = metayqma. 


Paroligoneurus pallidus sp. (wy. 
FIGS 6-1 


Material examined. Holotype: 3 ANIC, Norther Territory 
“12068 34.04% Cooper Ck, 19 kn Eby Sat MU, Borradiily, 
N.T, 9-10 Naw 19720. C, Curdale’) Paraty pes; 6 G same 
Jata ws holuivpe (ANIC. TAMU, WARI): #9. | unkown 
gen, LP4Q0'S. 132°54'E. Magela tk. & km SSL oF 
Mudginberri H.8., -8xi 1972, IoC. Curdale (ANIC, JAMU. 
WARI), | 6%, 12°27'S, 95°55 FE) Neartady Warde Dyobkene, 
Kuhadu Nal Ph, 27401980, 1. DB Naumann (ANIC) 

Queensland; 2G), fron Range, Cage York Pen... 
26-31. 1971, 5. R. Momenth tANIC). ) ov, sume dam exeept 
Qe l97P (ANIC): 1 9. 15403 'S, 4500'R, 3 kin NB ML 
Weblt 30.iy, 3.198). 1. DL Nauimitm, vx ethanal, colléctee 
at light (ANIC), 2 cree, Henrietta Ch, Paliierston Nat Pk 

23.v.1970, § Ro Curtin (ANIC): | oy 1272'S, 43 20° E, 
(3 km ENE Mt Tozer, J4.y7i. 1986, JC. Cundile, at MV tight 
(ANIC), Loot, 1816'S, 44°59° HR, km Woby § of Hope 
Vale Mission. 7-lv O81, 1D. Naumann, e& ethanol (ANIC). 


Female 

Head. Frons bare medially, head otherwise densely 
vetose: in dorsul view 2.05-2.20 “4 wider than 
Masui length, wader at eyes Chan wt temples, in 


46 AOD, AUSTIN & R.A, WHARTON 


literal Wew eye 2.85.08 % longer than iemple: height 
af head between apex of clypeus and base oF antenna 
LO-L 2 S narrowest width of face; eyes hairy (Pig. ¥); 
Jrote-clypead suture padistinel, elypeus thus nar clearly 
separated Jom duce. clypeus weakly convex in profile. 
ventral thargin Sharp, evenly convex, bering a line 
of long erect seract malar space short, in frontal view 
distinctly shorter thar basab welth of mandible. matar 
suture sharp. deep: untenna I8-seymented, slightly 
Jonver than body: Hagelluny browdest at middle. 
anidually narrowing apically ancl basally: first 
fliagellomere 120-LAS c¢ longer than second: fifth 
Nigellomnere about 2,5 2 longer thiea nud-width; tabu 
palp 4-segmeuted, the third segment mimite. 


Mesosomia, Short and broad, 13-15 % longer thin 
high, dbout as wide as high: pronotiin laterally often 
oollupsed in dricdmuaterial thus giving mesosoniut the 
appeuniice of being depressed. imesonotal dise weakly 
convex. neady Mats scuteltom (at, mesonotum densely 
anu untformty cover with short setae and assoc tated 
Weak punctures. notaull absent externally. bur visible 
internally as thin, ark streaks beneath pale 
mlcgument: sculellum densely setose faterally, nearly 
bare medially: propadeum polished, umsculptured, 
covered with sekue, these less densely spaced chan on 
mesondum: mesopleuren und anckipleuran polished, 
uaseulpturcd: hind femur brow. 26-3,0 % longer than 
mid wlth, 


Fore wing. Stigma very large. about 2% longer than 
broad, roughly 2 longer than melacurpus: arising, 
Sightly distad of midpome fully selerotised. pigmented 
portion of weakly curved + about hall length ol 
iretcurpis. but distitedy longer than piginented, 
selerulixed stub of 2—SR + M: anterior portion ul basal 
vein sharply bent distally: 


Merson Petiole nearly Wut. with very low weak 
dorsal cringe basin of spiracles, otherwise without 
sculpture: peuole broadest at spiracles. strongly 
narrowed towards base vind apex. base and upex of 
approsnmately equal width. width at spiracles 15-18 
*& width al apex, leneth Ll-}S & width al spiracles. 
‘12 bare. polished. unsculptured. with trapezoidal 
medhin scleriie, its apex roughly 2% wider Uren us 
base, Th ynd T2 with broad weakly scleratised arsias 
between median selerite and laterotergites: hypopyaiywm 
large. uhoul 2.42.5 4 donger than petiole. gradually 
narrowing over posterior halfto a weakly pointed upesxs 
ovipuNilor sheath (total length) nearly 2» longer than 
petiole (when dead), visible portion normally slightly 
Jonger Ulan petiole. with Veotral row of apical setae 
extending slightly more than half way towards base, 


Colour, Yellow to orange; fhee varying from dark 
Orange Wr varyegated orange and brown) remainder of 
heath and pool mele usually browey seupe ane 


pedicel. usually entire tirst flagcllomere and sometimes 
hase Of second flagellomere yellow, remainder of 
antenna browo; ovipositor sheaths black. 


Bady fengit, 1,7-2.1 nm 


Male 

As tor female excepr as lollows: median Hagellomeres 
more slender, Magellun thas less obviousts tapered 
towards apex, both Te and T7 with a deep median pit 
(Fig, 1). 


Referred matertat wnained, Queensland, | QB, 1, Bald 
Min iveu vin Ein Vile (ANIC); Uy Brarnstom Beach 
(ANIC): 2 cece, Brisbane (TAMU); 24 9, Saar, 12 km 
NW Brisbane. (TAMU: HL ero, Bunya Mt (ANIC. WARD: 
3 7 0 Camp Mountain (QDPI): 3 oe et. from Range (ANIC); 
2 eo, Me Tamborine (QDPI. 2 oo, Poluma Din 
(ANIC), New South Wules, 2 G9. Scots Hua, nea WMierell 
Ck (ANIC. WARD, 


Biology, Unknown. 


Diauiasts: This species is readily rceognised by its 
generally pale coloration, all congeners. having 
distinctly darker bodies. ‘Vhe metacarpus is shore 
relative lo B johaseni, and the transverse radial vein 
arises newrer the midpoint al the stigma, The venstion 
of P paltidus thus more closely resembles thut of the 
AMtrotropical A witlei De Sueger. Additionally. the 
ovipositor is longer than in all eongenene species. ‘Lhe 
antunoue of the Kogwn Afrotrapicul species are 19- to 
20-seginented, but the antennae are 18-segmeneed in 
hoth PE. pattidus and Po jekhasani, 


Discussion’ The distinetly setose eyes of P pallidus 
necessitate a clarification of the definition tor 
Paraleneurns, Mason's (M69) use oF selose cyes lot 
sepaniting genera inthe Muesebeckiini, and especially 
for separating Olwenetrus trom Parvliwonenruy, heeds 
clariticulinm, Nearly all species oF Raralipercaras hove 
af least some setae on the eyes. and several shortsetae 
are readily visible medially on the eye of /) jolmvont 
the type species. The number, size, ahd artangemen 
OL sche ConsUlule an important character set lor species 
level discrimination in Porvligenenres, Deep inedian 
pus. though not previpusly described. are found ina 
number ul touesebeckiines. ‘They ure usually Ineated 
an tentites 6 and/or 7, und accur in anly mates 


The lype series ov P pallidus has been restricted to 
the imuerial from Northen Territory and fir Noch 
Queenstand because of colour differences in the 
material from south-eastern Queensland. The more 
southerly specimens are generally darker, wah most 
of the melasomidirk brown. Elowever, there is some 
overlap, and there are insotticient representatives at 
ott Sexes front any one lacality tor advyuately assess 
whether or nl miles are Uurker [han fermnles. 


NEW RECORDS OF SUBFAMILIES, 
Subtainily Wisteromerinae Fahringer 


Chairs; The monolypic, Hixrerunenys Wesmael, 
has been vaniously treared over the years Lntil 
feeently, most 20th century authors placed 
Sisteromerns in the Doryetnae due ta the presence 
of StOuL selue or peys anothe fore Ubige. Falicinger 
(930) was the first to isolate ibus 4 separule tribe within 
the Doryctinue. Van Achterberg (1976) initially 
uansferred Histeramerus to the Bracuninge, but soon 
realised that it was misplaced. Van Achterberg (984) 
subsequently regarded itas a separate subfamily wath 
# asic Eroup relationship to Ypsistocerinae + 
Mesostoimue. This placement is based on the shared 
presence of a Mattened petiole, compressed hun 
fomora, dnd Jocation of the metasomal spiracles in the 
epiplcuron (van Achterberg 1984, 1988). Additionally, 
the transscutal articulation ts absent. In the daryeline 
Rhaprracentrus Marshall, however. the gaster is 
similarly shaped. with the spiracles located on the 
epipleuron, ‘The hind femora are also flattened in 
Shoprrecentrus (though penerally not as much as in 
Histeremeras) and the venation  sinmlar Phe petiole 
shape und absence of che propleural flange, the 
cpienomial (=prepectal) carina, and the transscutual 
wiculation are thus more useful features tor separating 
Historomierns trom doryetines, Additional features are 
discussed by Quicke & van Achterberg (1990). who 
consider the Histenamerinad to be one of the most basal 
groups of Braconidae. The subfamily is readily 
recognised by the exceptionally Jong hind basitarsus, 
oddly shaped head with long teoiples and yery shan 
face. short antennae. and chivate fore (hia with slaul 
sete ol\stere! (0 4 large patch along the dorsal (or 
duler) surface, 

This is the first species of Histeromterus described 
from quiside the Holurclic Region, 


Key to known species of Histeromerus 


1. Vein m-cu just aniefureal: fore bbw abruptly widened (Fig. 
171 [Austentia] . -- HL clavetis sp. nev. 
Vein m-cu pstfurcal: lore Uibia more cradually enlarged 
{Figs IW, WW). whet ale ee — 4 


2 Presternum yellow, anienna wilh 1S or [ower segments; 
sitiall species (alwut 25 py it tenethy |Nearcticl 
-fA conalensis Ashmead 
Prosternurm brown; antenna with 17-20 seynients; (iter 

species (at ledst 3. ni in lengthy |Pulueureniey 
H, eystacinny Wesmuel 


Histerowmerte clavatas sp. nov, 
FIQS 12 17 


Hwowpe 2 ANIC Queensland. "12.435 143. 16F QD A 
kin ENE Mr, ‘Torr H-lh duly 986.4, C Cardate Malaise 
trapethywnal 


TRIBES AND GENERA OF BRACONIDA 0 


emale 

Head, W254 broader (han meseholun [belwvecn 
teguluc); temples typically produced in ulpsul view 
2.25 x longer than eye: malar space abit hull eve 
height; length of frons (between anterii beellus and 
antennal secket) 17 & width of oeetlar treld: trons. 
vertex, temple and pena unsculplured, setae largely 
absen| on gen. sotal bases separated by length of setae 
on temple and vertex, more closely spaced an frons: 
face about equal in height to clypeus, about 4.3 x 
wider than high; face transversely strigose. Witt row 
of deep punctures laterally, extending (hrugh mitlar 
region: clypeus deeply punctate: untenna shen, about 
equal jn Jength to metusema, 15 seemented, ull 
Alagellomeres with multiple placodes, pulps 5- and 
3-segmented: apical setac on labial palp longer than 
3rd segnicat of palp. 

Mesosoma. Pronowin in dorsal view a narrow 
unsculptured band; pronotum laterality weakly rugulose 
except along anterior margin; mesopotum without 
notduli: density of setae on anterior dectivity similar 
that on frons, less dense on median part of dise and 
largely absent laterally; scute-scutellar sulcus unscalp: 
tured, without anterior demarcation, the mesonotal dine 
sloping gradually to form a. depression along anterior’ 
margin of scutellum; propadeum unsculptured; meso- 
plevron bulging, strongly convex, subalar depression 
deep. nitrrow, unsculpturcd; mesoplouron jacking 
erenulate posterior margin. metapleuron weakly 
wrinkled dorse-posterorly and ventrally 

Leys, As in other species of Aisteronterns: lind femur 
mare strongly compressed than mid terme: fore cose 
hmiadly contiguous; hind coxa long, abour 0.75 
length of hind temur: outer surface of tore Libia with 
short thick setae.on its apical half: fore femur weakly 
grooved ventrally for reception of tibia. fore tibia 
abruptly broadened over apical half (Fig 17), 
narrowing slightly trom mid tibia ty base. hind 
basitarsus very weakly curved. ghour2 \ longer than 
conibined length of ainsi 2-3. slightly inflated ever hasal 
half, 


Wires. Stgroa short, broad, nearly hemispherical. 
about 2.4 & longer (hin broad; r vertical; 3-SR about 
5.6% longer than r, subequal to SRI; SRI moderately 
eurved, reaching metacarpus somewhat before wing 
lip; 2nd submarginal cell brodder distally than proxi 
inally. m-cu antefureal by about Q.3 & its lengua; cu 
postfurcal; CUlbh.completely absent, Ist subdiscal cell 
{hus open at lower distal comer; M+CUL tubular and 
Pagniented except ul extreme bases IA+ZA Unekened. 
in region of barely visible 24; hind wing with I-M 
about 1.1 longer than M—CU; m-cu long, pigmented 
but spectral; Ir-ny shorter than cu-a: Ri of bind wing 
distinetly shorter than SC+RL. 

Metasome Petinle nearly fat. without dorsal ar lateral 
carinae and associated pits: ovipositor siongly 


Ag A. D. AUSTIN & R. A. WHARTON 


Figs 12-19. isteromerus clavaius sp. nov., 9 holotype. 12, lateral view of body, 13, fore wing, 14, hind wing: 15, hind 
leg: 16. anterior view of head: 17, fore tibia. 18. Aisteromerus canadensis Ashmead, 9 , fore tibia. 19, Histeromerus mystacinus 
Wesmael. ¢. fore tibia. Scales: Figs 12-14 = 0.5 mm; Fig. 15 = 375 wm; Figs 16-19 = 250um. 


NEW RECORDS OF SUBPAMILIES. 1RIBES AND GENERA OF BRACONIDAL WY 


compressed, blide-like. cleeper basilly. tapering 
distally. walhoul obvious tiedd- exscrted portion wbeut 
equal in lengilh Wirnesosintia) ovipositar sheath densely: 
selise, the sete longer than sheath width. 


Colmer Dark brawn: ptosterntun. anterian margin of 


pronoun, padps, first antennal segments Uargely), 
mich amd Hind coxae, tibiae. and all bar extreme up ol 
Wrst Yellows fore CONG, Mesosternum, and fernora 
variously yellow-brown, ovipasitar sheath whitish, 
wilh upied one-fifth brown: wings hyaline with 
infumuate streak aloag LM: microtrichia on membrane 
Very short ind thick, giving wing w spotted appesrunge, 
Berly length, 2.4 om, 


Male 
Unknown. 


Biviogy Unborn. bur host records of previously 
desenbod species indieine parasitism of coleoprenin 
larvie in Wordy steris Or bracket tungi- 


agrasig: This species is thast casily identified by 
its Genution, with ecu enterme the lirst submurginal 
cell, the Isl subdiscal cell oped in the lower discal 
coroer taniueh complete loss of €7Uth, and the vertical 
positidn on Bath Wo wlstecintty and AL cacdensis 
have M-cu puntfreal, (LIB present al least as a stub, 
vod rincliyous. The Australian species is otherwise 
very sume lo A. cartidensiy and Ho mystacitaes, as 
noted ubove im the nuinber of unique features used te 
Vefing the subfamily, Both AL lavas and HW. 
cunedensix are sroall species, with fewer flagellameres 
und paler coloration thaty A. rayaeacuins, The apical 
sewc on the palps are also longer and cu-a is postfuccal 
in the Iwo smaller species. 

Discusyie: The holotype has a spurious vein in the 
second submarginul cell pf the fore wing (Fig. 13). 
Anomalous vention fas also been recorded lor ff 
mvstacinus (Marshall 1885, 1888). Marshall's specimen 
showed tices of a seeond recurrent vein (2im-ou), 
producing a pattern similar to that in Apecsy Mason, 
The latter bowever, has thy petiole and prepectal carina 
more typical of duryetings than Hisreramerns. 


Subltimily Euphorinae Foerstec 
Commivnis: Shaw (1985) has provided substuntial 
suppon tor the ¢lade camposed of Srenothreniuna Shaw, 
Hesmielia Foerster, Chrwepaphthorus Goidamch, and 
drideiis Marshall, Although all but Wésiidelia are well 
represented if Australia (Shenetelt 1969; Mucdlesion 
983; Shaw 1984, this study). Clay sepophihorus has 
not been previausly reported front the continent (sed 
Masow 1964), und until recently relatively tew 
Swenetiremma have been knowh, Additional 
intoriariany Of these veneru iy presented here, 


Stenorhremma Shaw 
FIG, 24 


Comment, Amongst the most commonly cnacounered 
members of he Kuphomnue in Auscahan collections 
ae vannus species ol Srenerhremma, The genus was 
recenuy deseribed trom Australia am! New Caledonia 
(Shaw 19841, based on three species, However, mast 
Austailian species are undescribed (ep, approximarely 
20: 30 new species in ANTC) and the raterial ae hand 
considerably broadens the delinition originally 
provided by Shiw (484, 1985). Since Sretertirenile 
is such a promuient member ol the Australian 
cuphorine fauna, and bevause nates vannur be readily’ 
identified using exisuny keys, we Take Wis opportunity 
la present additiinal morphological di Hosts for 
Shenethronmae are unknown, but iwe undesembed 
apecies (ANIC) have been swept from aeuria und 
Eucalypius, respectively 


Stiw (1984, 1985) places Stenothrenimy within the 
Aridelus-Wexmaclig Chevsopaphithorns \ineage, and 
provides wset of synapomorphies for this group, The 
tnast useful of these for identification purpuses is the 
Jong, almost uniformly narrow (apical width Jess thin 
3 basal width) petiole which is completely: fuged 
ventrally troi) base to apex, This feature, (gether with 
the completely developed, tubular SRI+3—Sr and 
(-SR+M of the lore wing, are sufficient for placement 
of all Australian species in this lineage, The median 
Frontal carina, whieh) Shaw (984) hiss ay 4 
synapomoerphy for this grup of genera. is absent in 
some of the undescribed spewies of Stesorreanmae and 
weakly developed in others. de is) more strongly 
Ueveluped in larger species 


Some New and Ok World tropical species of 
Meleorus Haliday mght be confused with members 
of the Aridelus lineage. and care niust be taken ty avoid 
this error, In these species, the apes of the petiole is 
often less than 3 wider than the base, In ull cases, 
however, the sides of the petivlar remum are widely 
separated at least on the apical hind. Additionally, as 
noted by Shaw (985), the mandibles in Mereorns are 
broadly overlapping relative to the sickle-shaped 
mandibles of Stenathresuna, The petiolar nd 
mandibular characters are not always readily visible 
on pinned Specimens, Within the Aride/iy linesge, 
Anidelas is easily identified on the tasis of the reticulate 
or yericuhie arcolave sculpiuiring of the nicsonelum. 
The mesonocura of Stenethremme varies From punchile 
to finely granular, Both Weswarla ond 
Chyvsopophtharus have M+CUL at least partly 
descleronsed or absent, In all species of Stenothreonna , 
M—CUI is tubular throughout, and prowiles the most 
readily Observed character for separation (rom these 
iwo genera (ef, Pigs 24. 24). Austealian species of 
Chrysopaphitianes kniwn tw us have the basal half of 


an A. 1. AUSTIN & R.A. WHARTON 


ihe fore wing yellow, and MPCUL though appearing 
weakly developed because of the pale coloration, 1s 
actually tubular vet ty basal and apical quarter, and 
nebulous only near its mid length. The Australi 
Chesyopepithoruy are thus very similar to 
Stenerhrenina, Shaw (984, 985) hats cinphiasised the 
compressed mensoma in defining Svenurhrenmer, bal 
iis deste & fot useful for males, and varies 
gonsidirably in dried females, depending un the quality 
and manner of preservation (e.g, the metasoma of 
criticul-point dred specimens i frequently blowted 
melher than compressed), 

Shiw (85) provides au excellent character set for 
amlysis of cupharine phylogeny, Shiw's data fhe 
Merotirenma should be modified as follows, bused 
enomuternd avintable tous including all undeseribed 
species: 

Charter |. ocular sete: present in some species, 
Hhsent in athers. 

Character 4. medio frontal carina: extendhay nevely 
taantenorocellus besanie spevies, shoctand weak 
if others. dbsent in Some 

Charaeter & apical Navellomere: pomted in most 
species exumined.. bur rounded in-ac lease two 
species, 

Chunucter 13, nak suture: present br nearly all species 
examined, but weak uncldifficult to see ait several. 

Charaetrer 16. feial setae’ variable among species, 
dither obscuring face or not (as noted by Shaw 
(984) jh bis original deserptions of the species. 
but not reflected in the coding for this character 
in Shaw (1985), 

Character (7. shape of lower elypeal margin rounded 
(strongly CONVEX) In Mtest species, but nesrly 
Truneure in at least one species, The medutlly 
indented eundition given by Shaw W985) forother 
members of the Aridelis Tincage does not hold tor 
Iwo of the Australian Aridedes Species. and the 
indentation iv the Chaywsepephihoris species 
described beluw is barely percephble Tn these 
species, the clypeus vacres tront mare or Less 
Lraneate Lo CONVEN 

Character 19, onexdlary palps: O-segmented in several 
af’ the species exumined. 

Character 25, lews; the ditference belween the legs of 
Chinsupophtherus and those of small yellow, leypect 
Species ol Stenarlremme my very slight. 

Character 26, mesonotal sculpture: varies from finely 
writin tea finely punctate, The imbricitte 
Microscuiptare of the mesosoma whieh Shaw (984) 
noted as ii dnusgal leatire characterising, 
Stenfhrenund 1s absent ma few species. 

Character 36, neice length/fwidth; shart und broad 
in some species, moderately slander (enpth 3-6. & 
Taxi width) la others. very Jong and slender 
(lenetl) greater than 6 maximum width) inone 


species; bath character states used by Shaw (19851 
ute therefore applicable. 

Chavaeter 44, radial cells the distinee between the end 
of the radius.aind the wink (pis quite yairkible. and 
this variation is not itdequalely reflected in the 
character states uscd by Shuw ORS). 

Churucter 62. tergne 2+3 length’ the ditlerence 
between Steawdiremme aod Chavvepophihuras are 
clearly evident in fumules, but considernibly lexs so 
in males, 

Character 65, Lateral surare between tergites 2-32 this 
feature is present intl leaxt the Australi specics 
ot Chevsepophionis, although usually not as 
clewrly evalent asin Srevedinemia. Ww ts better 
developed im males thai lemales, 

There is litthe doubt that Stevotrenime beloogs i 
the Aridefusdincage, and although its exact placement 
theremm iy nday tess corkuin. dur dnitlysis does suppor 
Shaw's (1984) hypothesis af relationships, Shitw (1985) 
rents Srenethremma as Ube sister-proup of 
Wesmudelia + Chevsopophthoras+aridelus, With the 
new dala presented shove, we find that Lo of ube five 
chameters supporting the Hesmaela t Chavsapople 
thorws—Aridelusy clade (numbers 17 amd 19) de not 
hold, and the other three (nrumbers-62.-63 and 65) loan 
a single character complex associated with terga 2-3 
We treal (his entire characier complex asia chine, wath 
the plesiomorphie: stare found in Srenartivemme, and 
the ypomorphie suite found in dridefuy, ‘The condition 
in females of Chrysopoplithorus and Kesmedelia 0s 
definilely more like Aridefes than Steethrenen. and 
supplics Ue sule supporting feature forthe Hesmaela 
+ Chryvvopophitlerus--Anidelis elude. The problem of 
using charicters | and 36 to umambiguously support 
the Wevnuehet Chosapophithorus clade leaves. the 
partially desclerotised M+CO] as its strongest 
synapomorphy, Aridelay has a large number of 
wuopomearphics (Shaw 1985), emphasising its 
separation from the other gener, tlowever, Ue 
relationships among the other three genera are now 
less clear, Information on the hosts of Swe rene 
may help solve (his problem. for iP the tests are 
nevropteroid rather thane heniipteroid this would 
support a Stenotrenuna + Chrysepoplithorns clade 


Chrysopophiherus hageni sp ny. 
HGS 20-23 


Mirterial ovaaiined Holotype 4, ANIC. “SOL TH 
AUSTRATJA Adelaide Mian 129, 1900. Whitrloir, 
Poratypes) 1, 3 ore, sume dali as hololype (TAMU, 
WARE) 


Female 
‘ened. Transeerse. ucdorsal view 1.7 9 broader than 
mid lengthy |e broader than mesonotum between 


NEW RECORDS OF SUBFAMILTES. TRIBES AND GENERA OF BRAC ONIDAL 4l 


Pigs 20-24. Chrvseinaphthorns hagenisp. vow. Q batotype 
20, dorsal View of heads 20, anterior wew af lower heads 
22 Jateril view of body: 23. fore wing, 24, Steverhremea 
sp. lore wing, Scales = 05 min 


tevulae; eyes bulging, in dorsi) view 3.2 % longer than 
temples; temples convex. receding behind eye, densely 
coversd with shart decumbent setae; ocellar field 
small, widely separated from eye (Pig. 20); posterior 
ocelli separated by ubout 2.5 % their diameter: elypuus 
very broad (Fig. 21): apical margin thin, broadly and 
weakly truncate medially, very weakly emarginistc 
centrally; smooth, nearly impunctate dorsally, weakly 
transversely aciculo-punctate along apical margin face 
punichie medially, uansversely striate just below 
untennul bases; frons, vertex, and temples panctate; 
punctures narrowly sepanited, almost coalescing 
medially on frons. more widely separated (1-3 & their 
duumeter) on temples and vertex; malar space ruguluse; 
amenni 2-segmented: first Nagellomere about 5 
longer than wide, seeond flagellomere about 45 x 
longer than wide; filth Nagellomere about 1.4 * longer 
than wide: tirst flagelomere 2.6 * longer than fifth. 


Mesosoma. Pronotum aciculate laterally, mesonotal 
dise punctate, punctures weak (shallow) and less 
Uensely spaced on lateral labes thun oa median lobe, 


more densely spaced medially on anterior decliviry 
than on dise; notaull crenulate, distinct hough shallow. 
norrow anteriorly, converging and broadening 
posteriorly, the (wo sides separated posteriorly by a 
low median ridge; notauli nobextending to preseuteltar 
pity seute-scutellar suleus with median ridge only 
slinhtly beuer developed than lateral ridges: scutellum 
covered with shallow punctires, Tatert) cianzins 
carinaté only at extreme base, propodéum uniformly 
reticulate, without distinct carmae. shallowly 
excavated, mesopleurul dise polished. with diagonal 
row Of scutlvred punelures. otherwise smooth; precoxal 
swleus shallaw, punetate and irregularly alveolate, 


Forewing Secondsubmarginal cell subquadrangutir; 
2-SR and tm separated at the radial sector by about 
4% their width: 3-SR oearly equal in length tor, 


Mersome, Petiolcas long as mesoxomuat. (h2 % longer 
than width at spiracle. slightly deeper at spiracles that 
iW dpex and base, width at spiracle about 1.5 % width 
al base; petiole without sculpture laterally, evipesitur 
sheath about 0.8 x length of penole. 


Calour Yellow-orange: propodeum, metanotum and 
mirgins of sculellum varivusly brown to dark browa: 
T2 and apical one-quarter of ovipositor sheath dark 
brown to black, remainder ol ovipositar sheath, 
oviposttor, petiole, legs, most of pronotum, elypeus 
ventrally, and mast ol mouthparts (except red 
mandibular teeth) white to yellow-white: antenna 
yellow basally, apical sever Magellomeres brown, 
darkening towards tip; fore wing venation yellow basad 
of stigma. stigma and yeiny bordering second 
submarginal cell brown; base of meneapus vellow. 
Body feneth. 3.3-3.4 mim. 

Male 

As for female except as follows: éye smaller, in dursil 
view 18 1.9 s longer than temple; posterior ocelli 
separated by about 1.5 x their diameter: untenuae 21- 
to 22-seemented: filth fayellomere 2.6-2.8 » longer 
than wide: first flagellomere 13-16 * lunger than 
fifth, scuto-scutellar sulcus with median ridge distinctly 
better developed than lateral ridges.in 2 of 3 specimens: 
one male with distinet lateral carinae bordering median 
excavation of propadeum; 3-SR of Jere wing absent 
or nearly so, the second submiirginal cell decidedly 
petiolate in one specimen: petinle shorter, about 0.8 
* length of mesosoma,.6.4-7.7 % longer than width 
at spiracte. 

Referred meterial cvamined® ACT 15, to, Canberra 
(TAMU) 

Biology; Unknown. Other members of the genus ate 
parasitoids of adult Cheysopidiac. 


Diagnosis; This species ruins to Coupled 5 uv Mason's 
(1964) key to species, bused on the browd apd very 


52 A. Lt AUSTIN & R.A. WHARTON 


shallowly enmirginule clypeus, The distinetive 
seulpluring of the norauli. the polished median region 
af die mesopleucon, and the pattern of dark brown 
Mmartkiigs or the body readily separate this species fram 
all previously described Chrysopophiharus As Mascon 
(464) notes. C erieniafis Mason trom Singapore has 
wiumberof unusual features, This Australian species 
shures none of these and is thus not closely related to 
Co arientaliy. 


Discussion: The Wwospecunens from Canberta closely 
resemble those from Adelaide, but the fernale petinte 
is Shghtly shorter. and the clypeus is mure extensively 
punctate dorsally. We have seen un additional spectes 
from Queenstand (ANIC), but as atis thus far known 
only From males, if is not described here, The species 
wo onamed Jor Ken Hagen, in recognition of his 
cantzibutions to ehrysopid biolugy. 


Subfamily Mesostainae van Achterberg 


Comments: This small endemic subfamily wus 
reviously. known from only three species and very 
little material. Following reeogmiion of the sublaniily 
by van Achterberg (1975) and descripnan of the frst 
species, Mesostoa compressa van Achterberg tron 
Perth, Quicke & Huddleston (1989) described a secon 
species from Adelaide, M, austin? Quicke & 
Huddleston, These authors also pluced Tobias 
Monospecifie subtamily Praonopterinac (Tobias Ss) 
asa junior synonym of Mesostoinag, bul maintained 
Proonypterus (aeviy Tobias, trom Jervis Bay, ACCT. 
as a separate genus bused primarily on differences in 
wing venation. 

Meinbers of the Mesostamue show a general resemn- 
blance ta some cyclostoine braconids, purticularty 
certain doryetines. cxothecines and hormimes, but they 
cun be usually Separated from these axa by the labrum 
being only slightly depressed, fore Ubi evenly and 
finely setose, and antennal Muyellomeres (atenedt, 
However, the species deseribed below brings (wo of 
these characters into question, in that the lybrum ts 
strongly depressed.and oval in shape und the mandibles 
are curved distally'to form a subeyelostome nieutl (Pig. 
37), and the fore tibia has (wo rows of spines (Fig. 
340) The reengnition of these characters tor Mesostoe 
requires further interpretation, but muy incieute a much 
closer relationship with the Daryctinae than has pre- 
viously been postuluted (van Achierberg 1984; Quicke 
& yan Achterbere 1990), 


Key to known species al Mexasto 


|. Occipisl taring absent) prepedeum smomh, without 
medial longitudinal strigose sculpturiys, scutuin with only: 
a trace unterively ofa medial loopitudind groove [feriate 


anlenna with 12 Npellameres| M vanipirien ste 

vin Achtorburg 
Oceipial earn presear | Pigs 34, 36). propodeuny sirigose 
in medial forysitudinal dine (Fives 3, 33). seuluer with 
atmnat coniplete medial longitudinal groove (Figs 30, 32) 


2 
2 Qripestiov present (Pins 25. 26) emule)... .0 004 
Oyiposilor absent (inabe)- E 4 


3. Antenna with  Magellomeres.. 00.02... ML austiene 
Quieke & Huddleston 

Anlunna with (9 flagellomeres (lig. 261 M, Kerr? 
4p. How 


4. Posterfor halfet seatunm with some longitudinal rupose 
striate sculpturing Iatenuly, Ceansscutal articutation present 
hut tan, M. ans Quicke & Huddleston® 
Postenor hall of sentura vietually coumplealy smooth 
franssculy| articulation abwent (Pig, 32) M, Kevey 

ap. nove 


* malesaf bath these lwo species have [4-19 Hhpetlomenrs 


Mesostoa kerri sp. wov. 
FIGS 25.40 


Malerial examined Hototype: F, ANIC. Seurh Austraba 
“8. Austs Reedy Creek, J717S, HOSE. 1th Qe 1997, Austin 
& Dangerfield ex twig, gill of Banksia omurginatd’, Puratypes 
S29). 34d cro, sume data as holotype (29 9, 2 orate 
each im AEIC, ANIC. BMNH, CNCI, HNHM, ODPL, 
RNMH,USNM)5 22,5 ce TAMU; ALO, oor, 
2 of cach pold-coated. WARI), 74 9 @. 3K ceo S. Atist 
Reedy Crk., 3.x.1953. emerged from galls on Buitkyiv spr. 
dried from extended alcohol stonwe (0G, 10 Geer TAME, 
64 Po, 28 oo WART), 


Female 

Head. In dorsal view postenioe part of head broadly 
emarginate. distinetly Cruncate so that angle between 
vertex and oecipur is approximately. 90°, occipital 
ourina fine but complete throughout: vertex, temples 
and trons mostly smooth with very sparse short setae 
ovellar triangle obtuse, urea witha aud around trutagle 
faintly strigose; oeclli of equal size, ritig af distance 
belween poslenor ogelli lo shortest distince: Wo eye 
margin O8:11 (Pig, 34); frons broadly depressed, 
widest parcof head behind eyes.t-c, temples extending 
laterally pant linc.of eyes: faccund malar region rugose 
to Striate-rugose, with long seattered setae; lace evenly: 
convex, mitia of width of face to head (2.004 3), ratin 
of eye height to height of head (measured an avdline 
from) margin of labrum) (20:37): face slightly 
depressed ab epistimial suture so Ut clypeuy protrudes 
outwards. shghtly (best seen in antera-lateral view! 
lower margin of clypeus slightly conver and wrinkled; 
Jabrum depressed and aval in shape, mandible curved. 
inwards in distal halfte form subevolostome condition 
(scen bestun antera-ventral view); amtenin with 19 
flagellomeres, relative tenyths of Magellomeres: 4 


NEW RECORDS OF SUBFAMILIES. TRIBES AND GENERA OF BRACONIDAE 


vy 
lay 


Pigs 25-29 Mesostnd kerri sp. nov, 9 Nolotype, 25, dorsal view of body; 26, lateral view of boely; 27. tore wu: 28. hind 
wing; 29, 0°, paratype, dorsal view of head and mesosoma, Scales — 0.5 mn 


(.3:LbLI11), proximal 6-8 flagellomeres with very 
Sparse setae, more distal flagellomeres becoming 
progressively more setose; distal 6-7 flagellomeres 
about 15 * longer than wide. 

Mesxosoma. Moderately dorso-ventrally flanened (seen 


in lateral yiew), about 2 X as long as high: sculum 
narrower than head, as wide as long, medial 
longitudinal line depressed to form a shallow groove 
extending almost to posterior margin of scutum, 
amerior part of groove smooth, posterior part with few 


> 4 


fine Jongitudinal striae merging with surrounding 
aculplurmy; antenu-hiteral shoulders al seutun finely 
guise, posterior margin smooth, rest finely rugose- 
sirfiale Mwnlenier pil, narcuwing into fine rugese- 
Punetite Lracts posteriorly which indice position of 
notiul, ouler side of these wacts bordered by smooth 
strip; whole surace covered with Short seme. 
transscutal urdctdution distinct (Piz. 30): seuto- 
semtelliay suleus strongly curved posteriorly sind tantly 
crenulate. this sulcus separatinp distinct subtriangutar 
uxilive; medial scurellim smooth with finely striate 
lutea! borders, Virtually hairless and oval in shapes 
lator statcHum finally strigose to smooth: propodeum 
with percurrent thedial longitudiniel band of fine 
sirivose seulpiuring, postero litem corners smooth, 
rest ut propodoum very finely sirtale to rugase-strinte, 
with sone very fine background punetition (Pip. 3): 
i lateral View pronctum finely tugose medially sur- 
rounded by fine striate sculpturings extending tw 
mnareniss Mesopleucon smooth und bare except tor 
pogulose epicnemial arcu: precoxal sulcus indivated by 
He verte strate sculpturing: metipleuron rugulose 
on ventral ball, smoot dorsally; outer surtace of fore 
lihia with irregulitr double now ob spines (Fig. 39). 
Wings, Generally the same as M4, cendind and dittering 
from AL compressa inthe fore wing as follows: [—-M 
hropdly and liintly Sinuate: anterior partot I—~SR+M 
bent: 213 -M slightly curved hasally, subdiscal cell 
widening distally, 

Merisoma. As longus head and mesasoma combined, 
petrole (PT) about as loo os tradi width agrnss 
position ob spiracles. with fine longitudinal stmae. 
T2+Tthe largest tergite, about 0.6%) as long ay P47; 
sulure between T2 aad ‘T3 rdieated by fine transverse 
lines T2—TS smooth with single transverse nw of fine 
lidirs, OWpostlor and sheaths abour one-third length of 
Metso, sheaths with long sparse sete throuphout, 
Colour, Henl aml mesasoma dark brown ta black: 
mindibles yellow with dark Up: legs brown with hehter 
Buns al porots. femora sbehily darker: etusama and 
ovipesitor sheaths dark brown to black with enteror 
sermiies sometinies dirk yellow: brown, Winps hyaline, 
Sip pule, 

Bod\ lengih mean 2.6 1mm (runpe 2.4.2.9 n=13) 


Mele 

Sinitir to decale bul differing us dollows; length 2,3 
(ranve 1.9 2.72, n=5); posterior ovell7 minute Pig, 36), 
sOnTcHMes whysent.antenns with 1-20 fayellmeres, 
bruchypiterous (Fig 29) lore Wings rechingitar 
TEHCHID CO ulerion iKitgin AP propndeun. hase of 
wire durkly sclerobsed, rest white tn coloue 
nembranous and withour venation: hind wing minal, 
dhout ball teneth of fore wings mesosoma generally 
narrower, sean broader and more (ulcate anter- 
ivy. squarish at shoulders, Smooth in poster bale 


ND AUSTIN & ROA WHARTON 


transverse scutellur suture absent (Ps. 32). medial 
scutellum more elongate; fore tibia withour distinct 
spines on Outer surfice (Pip. 40}, metasomis longer than 
hedd — mesosoma (Gfh4 3): ‘TL broader aeross 
position of spitactes (han Jong (20:14): sutdee heqweer 
T2 and 13 complete and membranous, (hese and other 
Tergires subequal intength: T2-T6 smooth, witha lew 
scattered minui hirs. 


Referred marerial examined: Victoria. 4 Woe. Soe, 
Melbourne, 78 1904, Banksia pally (AMINE) 


Biology. This species is assovied with galls on the 
outer branches of Banks utareteea, a relytonship 
wiht (his plant genus that may be general for all 
Mesostou spp. given that Mf. ascii has alse been thus 
reured, However. the exuct host iy not yet known. bul 
presuntably at isthe primary pall former or one of the 
several Tisects that inhabit Berdsra galls, sdeh as 
cureultonid beetle larvae, 


Diseasion: This species is much closer to MM, ausdnrt 
than itis to A compressva. the huter species bas the 
head and seutund more extensively sculptured with 
conirser Curved striae, The scutuni of MW, erumiprensa, 
although baying the posterto-medial purt fattened, 
lacks a longitudinal groove, the propedeunt is smoot, 
the demale antennue only have 12 fugellomeres. and 
the lateral fields of the scutellum are striate In 
comparinon. Af cusvind and M. Kervt venerally have 
the fave rugose 10 rupose-striate and dre scutuns linely 
Nigose-striate, the seutum with a longitadinal groove. 
the propodcum. medially strigose, the female antennae 
with a greater number of Mugelomeres, and the lateral 
fields of The scutelhunr sinouth or haothy striate, M, 
austin’ and M) keer? differ substantially only inthe 
quniber a} antennal Hayellorercs tor the lemate ane 
more subtly on the degree of sculpturing on the beuil 
und seutuin, with WM. Aerrd generally being Jess 
extensively sculptured. 


As pointed out by Quicke & Huddlestin (989) the 
Presence of absenge of an occipital Carina ts often Used 
ase primary distinguishing, character at the generic 
level. andl inthis respect there is Some justification for 
placing M1. auatiné and M. kerri ina separate venus 
trom M. compressa, However. until more material of 
lis fare sublimily becomes available Where 15 little oy 
oo advantage in arranging the lour known species yt 
Mesostoinue in three separate periera 


This species ty hunted after Professor Allen Kert. 
inaugural head of the Department of Crop Protectian 
a the Waite Institute. and one of Australis leading 
SCUCTITLSES. 


NEW RECORDS OF SUBFAMILIES, TRIBES AND GENERA OF BRACONIDAE 


> 


Figs 30-33. Mesostou kerri sp. nov., 2, paratype. 30, dorsal view of scutum and scutellum, 31, dorsal view of propodeum 
32, 33. &. paratype. 32. dorsal + of seuturm and scutellum; 33, dorsal view of propodeum and Tl. Scales = 100 pm 


AD. AUSTIN & R. A, WHARTON 


Figs 34-40. Mesostoa kerri s , paratype. 34, dorsal view of head; 35, antenor view of head. 36. 37, or, paratype. 
36. dorsal view of head) antero-ventral view of head (N.B. transverse lines on { are due to specimen charging). 
: , paratype, 38, tip of ovipositor, 39, fore tibia. 40. Oo, paratype. fore tibia. Scales: Figs 34-37 = 100 um; 
Figs 3840 = 50 pm. 


NEW RECORDS OF SUHEAMILIES 


Subfamily Meteurideniae Capek 


Comments: This small sublimily is defined by its 
biology (gregarious binal-pupal enduparastoids. ol 
Lepidoptera) and highly modified melusoma (Nixan 
94). Capek (1970) separated the numiiate genus, 
Meteandea Ashmead, from the Diaspilini on the basis 
of larval murpholugy and biology, and placed it in a 
sublioly of its owe, Lntil tecently only two gentns 
had been deseribud, Meveoridea and Bencana Nixon, 
Shenefel) & Mueseheck (1957) redeseribed the 
previousty poorly characterised Merearidea, amd 
synunyinised Henan with it. This synonymy was 
accepted by Cupek (1970), Vary Aciterberg (1984), 
however. umphied that the two were distinct, but has 
since reversed his opinion (van Achterberg 1990). Lo 
nddition to the Australian species described below, we 
hive examined maternal of Meteoridea from West 
Atrica and Nort Amencs. In North Anmencan 
material. (he mexhiin Jobe an the apical margin of che 
elypeus ts more tooth-like than in the Australian and 
West Alricun species. Additionally, the deep basal pits 
of the petiole (dersope) are more laterally displaced 
in North American species, and mot visible in dorsal 
view, However. we do not consider these differences 
sufficiently clear-eut dor separating Benda from 
Meteoridea Van Achterherg (1990) has recently 
deseribed a third venus of Meteondetnae trom New 
Zealand, Prenkia van Achterberg, which bas a number 
of unustial features that abigail. at least superficially, 
wath rhe Agutlidinae and Sigalphinac. Proakie difters 
substantially from Meteardea in that it has a smooth 
propoadcunk dorsope absent. fourth tergile depressect. 
Ipre wing vein P-SR presence and vertical. 6 short. 
Me+CUl uuselerotsed. and bind wing marginal cell 
slender. 

‘The species of Metevridea described here is the first 
record for the subfamily trun che Australian Continent 
Although van Achterberg (1984) hus previously stated 
that the Meteorideinne jire “restricted to the 
subtopics. the description of MA. compressiventers 
Shenelell d& Muesebeck from Wisconsin. U.S.A. 
IShenelelt & Meusebeck 1957) and A eretefiernvalie van 
Achterberg from New Zealand, clearly shaw that the 
sublumily extends into more temperate exons. 


Meteoricdea anie sp. wv, 
FIGS 4t-44 


Mutctialevanined. Holotype: S. ANIC, Qucenshind, “15038 
WS.090 Akm NE Mt. Wehb 4 Chet. XO 1 C Curdale 
ex orhanol” Paratype. 39 4 IST'S, 148° E, Mt Webb 
Nal Pk. 26-30.in 180. SC. Curdale, ex ethanol (ANIC, 
WARD: 5 99, Ie S, (45° RB Shain W by N Rounded 
Hill ae Hope Vole Mission. 2 W980 JC. Curdale. ex ethanol 
(ANIC. TAMLL WART): 1 oy. LS kan St Kurinda, 
In (7980 J DL Nauman & ho. Cardale FANTOD 


PRIBES AND GENERA OF BRACONIDAE ry 


~! 


Forrale 

Heel dn duirsal wiew wider than seutun temples 
hroaus eves hulbows and glabrous: ocelli forming a 
compact (inngle. distance between posterior ocelli 
much shorter (han uishince from then tu margin of 
eye: neciput. vertex. trons aod temples smooth and 
shining, excep for few tmy punctures assheiajed with 
Hecusional Fine setae; head in antenar yiew almost 
circular; fice strongly convex, with broad medial 
longitudinal ridge und scullered punctures associated 
with long fine setae: cpistomal suture impressed: 
elypeus convex: wilh scatleread punctures and slightly 
up furned lower margin; mulir space Sinall. margin 
adjacent to mandible slivhily: convex: aunlennal sockets 
with mised margins; aniennu 3l-segmented. all 
flagellomeres longer than wide. reuchify ay Tar ais 
posterior edge of tnetasoma 


Mesesoma Pronolum with large dorsepe. in lateral 
view with mediostigquanal line crenulate, posterior and 
ventral margins ¢renulate; seutum smooth with 
nceasional scatters! punctures and assxo¢iated: fine 
selae: malauli percurrent and crenulate, anterior 
decliwus partcns broadly eremulate: transscutal 
wrtCUltON Staight, scula-seutelbir suleus. comprising 
2 or 3 deep fiwere; scutellum convex, smooth anil 
shiny, cxeept for a few scattered punctures und 
assuciated long setae: lateral helds of scutetum faintly 
Suriatc: posterior manzin of scutellunt smooth though 
sometimes with fuint medial rupasity: metanotum with 
2 prominent medial longitudinal carinae ant less 
chistinet curmae laterally; propodedl carinve sometimes 
sumewhat irregular bot always forming & dustiact areal 
and enclosed lateral and posterior gress whieh are 
punctate br rugose-punctate> surface of propolam and 
metapleuron covered with long fine sete: precoxal 
Sulcus and pleural suture faintly crenulate: flange above 
epiencnugl arca carmate (see van Achterherg, J974), 
margined hy crénulate or fivedlate inopressions, 


Wings, Fore wing with vein I—M slightly buwed, 4 
ementiog from mil peint of stigma, Cilla strongly 
wiched husally: subbasal cell narrowed slightly at 
middle; subdisew) cell widened distally; find wang 
L Sound 2-M indicated by short pigmented spurs 
busally, rest of these veins desclerotised: M+CL 3 » 
us long as L Mi I-I1A deselerotised 

Metasoma, Abimost as lang as head and mesosoma 
combined; petiole (TH slightly constricred behind 
spimetes then widening slightly in posterir ball 
Widest 2crogs pusterinr margin. 2.5 © lunver than 
witle. with distiine( anten-lateral pits. dorso-lateral 
margins distineth curtoate, dorsal surface 
longiludinally striate wilh punchile to ragosc-punctire 
background sculpturing: T2 and all other ntetasomal 
fereues smooth and shiny wilh aecastumal scattered 
hairs concentrated laterally and on posterior leriles: 
Posterior Mast terBite somewhat extended distally anc 


5h A.D. AUSTIN & R. A. WHARTON 


Pies 41-4. Metcoridea anie sp. nov, 3. holotype. 41, dorsal view of body. 42, lateral view af body: 43, tore wing, 44. 
hind Wine. Scales = OS mini. 


SLEW RECORDS OF SUBPAMILIES. TRIBES AND GENERA OF BRACONIDAF su 


hiaerally lo form a capsule enclosiag ovipusitor; 
oyipesior and sheaths hidden 


Colonr Body including legs unitormly yellow. seape 
amd pedicel yellow, Mugellomeres brown; mandibles 
darkened distully,; wings hyaline. venation evenly 
coloured. sigma translucent yellow-brown. 


Mute 
Unknown. 


Biolign: Unkiown 


Dingnosis: The uoitormly vellow body separates this 
species from all but Ad, resraced (Granger) frowi 
Miatditgasear, Has nearly wenticat to (he later, differing 
Only on nner sculptural features of the pettoles, 


Subfimily Helconioue Foerster 


Commenty; The telconines represent a rather clrverse 
assemblige of taxa whieh, ever inthe strict sense (ie 
with the removal ot Cendcoetuy Huhday. into a separate 
subfarnily —Szepliget: 1902), muy sull be polyphylene 
Oral best paraphyletic (Quicke & van Achterberg 
1990). Van Avhterberg (1983) reecoenised four wibes, 
Heleonin Ashmead, Brulleiini van Achterberg. 
Diospilind Foerster and Brachistint Foerster, all of 
which ure represented in the Australian tauna 
(Brullenot onty by undeseribed species). OF these the 
Heleorini is the most diverse, with four of five 
recorded genera endemic to Australia. Heleon Nes 
von Esenbeck is Virtually cosmopolitan in distribution, 
while dasrroheleon “Tarner, Peraheleou Kokujey. 
Thichtwhelcon ‘Larner and Calehelcon ‘Tarner are 
Known only from maintind Australia and ‘Tasmania. 
Collectively, they are represented by nite deseribed 
§peeics, wilh the first three genera not having been 
weatcd since their original deseriptions (Kokuyey 1901: 
Turner WIS). Calohelcon bas recently been redesxcrihed 
aod discussed by Quicke & Holloway (990), The tribe 
Heleonini has been defined by the présence of the 
following characters: froos wilh a medial longiudinal 
carina (hamella). hind femur rugose ventrally: 
Propodedt spudcle situated medially. and lore wing 
veins 1-SR und 2A present (van Achterberg 1983). 
AS is true of many ofthe Australian helconines which 
huve been placed in the Helconini, Calokeleen as 
unusual in several respects, The species of Caloheleon 
and Techiohetcan which we have examined havea very: 
smnoath body and so lack aprecoxal sulcus.and carinate 
ac rugose propadeim. Calofefeon is particuhtrly 
rerparksible on (hat We first thetasomal tere cs 
entire so ith lo appear bitlaled (Tips 48, 31), Quirke: 
&  Molloway (991) also sture thal 


Caloheleon bas retained a nurber of plestomerphie 
characters, in particulara darge number of humuly, the 
presence of hind wing vein meu. andthe presence of 
a costal cell in (he Jore wing, Clearly, the definition 
of Heleonim used by vain Achterherg (O83) Mast be 
feassessed i the light of the Australian latina. but Unis 
ginnot be wccomplished until the rich helconine fauna 
of this continent has been more thoroughly desenbed. 


We deserthe below a third species of Cufohelcar 
from central Australia, where the genus has previously 
been known only fron the castern coast) margin of 
the vontinent. The inclusion of this species extends the 
limits of the gens slightly and requires the diagnosis 
of Caloheleon presented tn Quicke & Holloway (991) 
ta be modified os follows: frons with median longi 
wudiowl! curmoa varying from well-developed to reduced 
or nearly absent: propoded! spurucle circulue or slightly 
elliptical; fore wing with costal cell open for-about two- 
thirds of lengih of yes C and Se+Re Rs to almost 
glosed over; hind wing with vein ni-cu present or 
absent: hamuli number varmable (5-9). owipositer as 
long as or longer than body. Cufohelean shires. i 
number of feanures with Urichidehelomi, but is readily 
scpriraled by (he intlated, nearly bare first metasonnl 
lerpite, 


Host records for the Helconinae show thot they hive 
only been reared as. endopurastoids of coleopleran 
larvae. We treat with seepticism the record for 
obsenripenniy Turner in Quicke & Holloway 991) 
(QO. ANIC “probing tree trunk with cossid lurve") 
its evidence that the host biology of this genus de pacts 
from’ thal known for other helconine genera. Tn aur 
experience, Aucalypris and vlcarier trees ean be heavily 
Infested With both coleaptecan and lepdaptenin lat vac. 
and so observed ovipositer prohinge is likely to he 
Maccurale oyu necthiod ob assuctating polental husty, 


Key to knewn species af Crlokelcoun 


1} Dorsal surtiee at Th ty hiteral view Convesty routed 
in anterior pare and flauened pastenorly Cag, AB) 
evipositor longer then berly: body & agar lonwtih or 
PSE YTAE se stators yoatese: odten it leen nip CO. cfngertivhl sp. ny, 
Dorsal surtive of TH ro latent view wilh hice honp ta 
anivriar halband weakly rounded posteriorly (Fig. St, 
Ovipositor as longais bodys body abuue 1 tim ia letigrh 
or longer, a : 4) 3 


ie 


Later) martios at“) in dorsal view conistrieted Ht antertt 
parts seucuin wd 15-19 black, wins yellowish basally, 
artw-browr apically C ehycuipenais Turder 
Lateral nrarginys of Th im dorsith vies only slietily 
constricted in anenor pari scum and TS TY orang, 
Witte evenly light brow 


—.0C padei Quieke & Holloway 


60 A.D, AUSTIN & R, A. WHARTON 


Figs 45-47. Cafohelvoa dangerfieldi sp. new. G holovype. 45, dorsal ew of head and mesosoma: 46. dorsal wiew of metasemi. 
47. or. doesul view of metasoma, Scales; Fig, 45 = U8 mm: Figs 46.47 = 0.75 mm, 


Calohelcon dangerfieldi sp. nov. 
FIGS 45-50 


Material exmatined, Uolotype: 2. AEIC, Northern Teeritory, 
-Yuenduma N.E, Australia August’) no collector or date 
given, 


Fenutle 

Head, Completely smooth and shiny: temples and face 
with minute punctures and associated fine setae; vertex 
and [rons virtually bare; in dorsal vicw occipital carina 
angled slightly so as to be obtusely pointed medially; 
in lateral view occipital carina extending ventrally ta 
meet hypostomal carinuy ocelli foriung equilateral 
Inangle, distance between posterior ovelli slightly less 
than distince from them to eye margin (2.0:2.3), in 
anterior view verlex convexly rounded so that lateral 
acelli are above dorsal margin of the eyes: face evenly 
convex. pode between antennal sockets extending 


dorsally into short faint carina which fades. out before 
reaching frons; eyes more than half height of head 
(2.5:4.0 — measured in lateral view trom vertex ly base 
of mandible); malar sulcus absent; clypeus moderately 
uansverse, slightly less than 2 * wider than long: 
mandibles short, only overlapping shghtly: antennae 
reaching to about midpoint of T2+-T3, 41-segmented 


Mesosoma. Slighily narrower than head: pronotum 
well exposed dorsally, coarsely crenulate around 
pronope. crenulule fine lading on smoth lateral 
pronotum, laiero-anterior margin of pronotum finely 
crenulate; scutumm, seutellum and propodeum smooth 
and shiny, with a few scattered fine setae: antero-lateral 
margins of scutum slightly emarginate at point of 
notaull; notaul: crenulate and reaching posteriorly to 
about middle of scutum; scutellar sulcus developed as 
2 deep foveae; flange above epicnemial area carinate 


NEW RECORDS OF SUBFAMILIES, TRIBES AND GENERA OF BRACONIDAE 6l 


Figs 48-51, Calohelvon dungerfieldi sp. nov., 2 holotype. 48, lateral view of head, mesesyma and anterior metasoma (carinate 
flange above epicnemial area, arrowed): 49, fore wing: 50, hind wing. 51. Calohelvon obscuripennis Turner, @ , Vatenil 
View of head. mesosoma and anterior metasoma. Scales: Figs 48-50 = 1.0 mm; Fig. 5! = 1S mm, 


to A.D. AUSTIN & R, A, WHARTON 


(Fig, 48. arrowed) and reaching anteriorly to (ouch 
dorsal part oF prepectal carina, 

Wingy. Costal cell of fore wing indistinet; m-cu much 
shorier than 1—M so thal discal cell narrows distally, 
1—SR4 M sinuate: 1 SR very short almost obliterated, 
2-SR -M 0.67™ as long as 2 -SR: 3—SEK as long us. 
rm: SRI straight, hind wing without vein m-co 
urising from 2—-M: Ri with 5 hamuh. 

Metasome, Th longer than T24-13, in dorsal view 
browdenng posteriorly. with broad shallow medial 
longitudinal depression in anterior one third, hteral 
Murgins virtually straight. i lateral view convesly 
rounded in anterior part and flattened postericly. 
lacking large antero-Jateral pity: suture between T2 und 
T3 faints oviposttor fonger than body (75°60) 
Colour, Head, mesosorta ineluding coxac and T2—TB 
orange-brown: antennae and legs black; latero-anterier 
hallo pronotunt black: propleura yellow-brown; wings 
evenly and darkly infuscate; TL white: SE white with 
2 broad dark transverse hands: laterotergites of T2 and 
13 and posterior sterniles black; ovipositar brown, 
sheuthy black, 

Body lenprh. TO mm. not including ovipesiter. 


Aterle 

As for demale except as lollows: slightly larger in size. 
hody length 8.0 mim: Th lyrger, in dorsal view wader 
than rest of metasoma, Jateral onurins rounded: 
2-SR+-M of fore win almost as long os 2—-SK; costal 
cell Slightly more Obvious; Mange ubove epicnemial 
grea not reaching anteriorly ws far as dorsi part of 
prepectal carina, lateral pronotum more extensively 
black, anlerion mesopleuron and distil ball of all coxae 
black. 


Biology: Unknown. 


Referred material examined: South Auacralia. {, Dalhousie 
Springs, 7903, G A. Holloway (ANIC). 


Diaenasis: This species is naust easily idenutied by 
the shape of Tl, the crenulate notauli, lack of medial 
sculpturing and a carina on the face and frons. shape 


of Lhe pronolum (in Jateral view), fore wing venation 
number of hamuli and length af the ovipositor 
Although this is the ficst record ofa male for the genus, 
we have noc included the single male specimen in the 
type series because there 1s a possibility that the slighi 
differences between the sexes. deseribed here are 
representative oF lwo species. not mntmspeedie sexual 
dimorphism. Until more matertil becomes available 
this problem will not be satisfactorily resolved. 


Riymalogy. ‘Vhs species. is named after Paul 
Dangerfield in recognition of (he iustrations he has 
Prepared for us. 


Sublannly Alysitge Stephens 


Comments: In a recently published paper by the 
authors revising the Australian members of the Tribe 
Dacnusini (Whartun & Austin 1991), several type- 
sciling errors. were overlooked which could result in 
significant taxonomic contusion, We therefore take this 
Oppertumity to correct the most serious of these. as 
follows: 1) p. 198. line 30. subheading “'Chaenusi 
mgricapinis should read “'Charehus nigvivapitiy 23 
p. 201, line 501 or 2” shauld read") of 2") and) 3) 
p- 205, line V7, “'arealis”’ should read “erectarts 


Acknowledgments. 


We thank Jan Naumann und Jo Curdale (ANIC), 
Betun Cantrell (QDPI), Paul Marsh (Washington, 
B.C), Jend Pupp (Budapest). David Wahl (AETC). 
Torn Huddleston (BMINH) and Mike Sharkey (Ottawe) 
for the Joan of material. Paul Dangerfield for techmical 
assistance aod for the line drawings, and Leon 
Praetorius for assistance with the dmiwings. of 
Aisteramerus legs, This work was supported by a grant 
from the Australiqn Biological Resources Study 
Participulory programme to A.D.A. and was 
undertaken while R. AW. was a Distinguished Visiting: 
Research Scholar at the Ware lostitut from October 
1989 to March 1990. 


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_____ (1986) The braconid genus A/vseee (Hy menopwny! a 
description of the subgenera and a revision of (he subgenus 
Alivia Syst. Bet. UL. 453-504, 

& Austin, A. D. (1991) Revision of Austrafian 
Dacnusint (Hymenoptera: Braconidae: Alysinae), 
pirasitoids af Cyclorrhaphous iptera, A dase ear Soe 
30, 193 206, 


AN EOCENE MEGAFOSSIL FROM NELLY CREEK, SOUTH AUSTRALIA 


BY D. C. CHRISTOPHEL*, L. J. SCRIVEN* & D. R. GREENWOOD** 


Summary 


Clay from the Eyre Formation in Nelly Creek in far north South Australia contains the first Middle 
Eocene mummified leaf flora reported from the interior of Australia. The 269 leaves collected are 
placed in 16 parataxa, with one angiosperm parataxon of unknown affinity providing 64% of the 
flora. Eleven of the 16 parataxa can be assigned to extant families which include Proteaceae, 
Myrtaceae, Araucariaceae, Podocarpaceae, Casuarinaceae and Lauraceae. Myrtaciphyllum 
eremeaensisi sp . nov. is formally described. 

Comparison with Middle Eocene megafossil floras suggests that the Nelly Creek flora is 
taxonomically distinct and physiognomically more sclerophyllous than the other south-eastern 
Australian floras. However. Sampling programs in extant rainforests and other Eocene deposits 
suggest that the number of parataxa (16) recorded at Nelly Creek from this first collection will 
likely increase markedly with further collections. 

Comparison with the silcrete floras of northern South Australia, in particular the Poole Creek flora. 
Demonstrates that while some taxa, including a possible Proteaceae infructescence, are common to 
both deposits, the majority of both floras do not correspond. 

KEY WORDS: Fossil, Eocene, Nelly Creek, Silcrete, Myrtaceae, 


Jimbo tlie Revell Sveti Stee aU TOO) 65 Tt 


AN EOCENE MEGAFOSSIL FLORA FROM NELLY CREEK, SOUTH AUSTRALIA 


by D.C. CHRisTopHel® L. J. Scriven & DR. GREENWOOD** 


Summarys 


fem roniin. DOC. Semivbs, Lot & Grheawonn, DR, (897) Ag Eocene meyalionl Too ham Selly Creek, 


South Austealia, fea, AL Sie 


A lest, H16( 2), 68-76, 29 May, 1992, 


Clay fan the Eyre hormanon in Selly Creek in far north South Australia contin tbe first Mickle Bayeue 
cohimomitiee ted Horceeperted [one minor al Austatia. The 269 Leaves collected aire pliwel in 16 piranina, 
Wh Ghe angiosperm patatisen of unknown affinity providing 647 of the teen, Gheven of the 16 parutaygecay 
beassienvd (oes tinit Binities witch include Pretcqeeue. Myrhiceac, Araucarlacede, Podocarpacend, Castarinaccae 
and) Lauraceae. Muriceydviian eremmecensts sp. mos, in tormiully deseribed, 

Comparison wah Widdle Rovene megatossl floras suuvests the the Nelly Creek Mortis waonganeally ascii 
und pbysiagnontically more selerophyllous than the other south-eastern Amstrad Hone However scimpliiy 
PrOLTaNS In SANE cintorcsts vad arher Lacene deposits suggest thar (he numberoal parutixa (lo) reeurded ctl 
Nelly Creek Irani this Bese volleetion will likely merase markedly with futher colleetions 

Comparison with the silerene Hos of nother Seuth Australi. i parriculir the Poole Creek ora, dernoistnates 
that Whike sont Lika. ieliding & possible Protaucede miciiefescence. rc goon La bath slepersits, Me mayo ily 


Of both Maris dha nut conespond 


Wi Worbs, Torxsil., Boceng, Selly Creck, Silent. Myfievie 


Introduction 


The Middle Eoeone flora of Austeilit is well known 
from megifossil localities in seuth eastern Austeatis, 
These inclade the Anglesea flora (eg. Christophel 
WR: Christophel & Lys 986, Christophel eral. Ws, 
Hill 980: Rowett & Christophel 190), the Golden 
Grove Mord (Barrett & Chostophel 1990, Christophel 
& Greenwood 1989). the Maslin Bay flora (Lange 
1970; Christophel & Blackburn 1978: Blackburn 1981) 
and the Nerriga flora (Hull 1978, J9B3), These oeeur 
neur the coast, With the first three considered to he 
lowland and the Jitat (Nerriga) to be upland (Pig. L), 
All of these Hort have been interpreted as represensing 
tropieal to sub-tropical (or very warm leniperate at the 
mvunial) rainforest communities (Christophel 1989; 
Christophel & Greenwood 1989) ind all contin well 
preserved. compressed er munimitied leaves, Wlowine 
MaXUNUT potential for Witerprektion. 

A second Source of data on Early ‘Tertiary orgs in 
southern Australia comes from the extensive 
Impression and cast floras of mtetior southern and 
central Australia, collectively known as the silerele 
Norns, While known for a relatively long time 
(Chapman 1937), theses Horas have aot played amiajoe 
rofe I ierpreting Tertiary vegetation because of whe 
lack of stratigraphic control, More recently, Ambrose 
eral, (1979) sugested an Eocene age for some of these 
sileretes. including those deseribed by Lange (1978) 


* Peportutent of Boliny. University of Adelaide, GRO, Box 
498, Adelaule. S. Ausr S004 

 Gelogy Depaciment. Ciniveoaty ol Saskatehewarn 
Saskatoon. Saskitchewan, Canada S76 OW, 


contuning leptospermotd Myriuceae Truitt. Recent 
work hy Greenwood ere (9909 described a Mor 
from the Poole Creek area of South Australia ancl 
placed the age ay Middle Hocene based on 
lithostratigeephic vorrelutioos. Fossil pollen has nol 
been preserved in these silerctes and hence palynology 
GOUT not be used to confirm their awe. 

The discovery in 1986 by R. Callen of foysiiterous 
clays in northern South Australia whi¢h contained wet] 
preserved. compressed and mummified leaves. and 
which was Interpreted as Middle Roeene (Alley 
989°) was important for several reasons. Firstly. it 
greatly extends the geographic range of well-preserved. 
Middle: Eogene megilossil floras. Secondly, it provides 
hiostratwraphically datable evidence fora truly infand, 
lowland flora of that age, and finally, it provides the 
possibility of betier chronolomeal control uver the 
imerpretation of the numerous silerete floras of the 
interior (Ambrose wed. 1979: Greenwood cf al. 1990) 
The aims of this report are therefore to provide a 
pliminacy description ofthe megatassil flora of Nelly 
Creek. to formally describe a new species of Myrtaceae 
front this deposit. and to conipare the parataxu front 
Nelly Creek with the known silcrete elements, 


NCinecAwoop. D Ro Caripy ROA & Aticy NOP ftgon) 
the correladen and depositional environment of Tertiary 
strate bested wo racnotlords an the sauthecn Lake Eyre Bust. 
South Australia, SoA, Depurtrnent af Mines and Freres 
Report 9O45, IST. Plates 1-7, 

ley, NF (989) Prelintinary Palynalosical chitin of 
macenfloras Irem Eyre Formation, Nelly Creek, Luke kyre 
Basin, Depr of Mines and bnergy of Sou Australia Rept 
Bh Nu, 89/46 


60 


@ NELLY CREEK 


Db, C, CHRISTOPHEL. L. J, SCRIVEN. & D, R, GREENWOOD 


@ GOLDEN GROVE 


MASLIN BAY 


Fig. 1. Map.of southeastern Australia showing the location of Nelly Creek and other Middle Eocene plant megafossil Incalities. 


Materials and Methods 


The Nelly Creek flora is contained in the Eyre 
Formation located at 29°19'S, 137°18" E, approximutely 
| km south of the southern shore of Lake Eyre South 
(Fig. 1), The deposit consists of sands, silts and grey, 
carbonaceous clays forming a portion of the bed of 
Nelly Creek. Overlying strata consist of partially 
silicified sediments, disaggregated sands, and a salt- 
pan crust. The deposit is restricted to the stream hed 
and is only accessible when little or no water is found 
in Nelly Creek. The fragility of the material, as well 
ag the terrain and general inaccessibility have severely 
hinted the amount of material collected to date, The 
extent of the fossiliferous. clay horizon outside the 
stream channel is unknown, although fossiliferous clays 
have been intersected in a. number of bore holes mi the 
region (N. Alley pers. comm,), The width of the 
deposit within the stream bed is less than 3 metres, 
and its thickness less than | metre. 

The high water table and the high salt coment of the 
ground water result in freshly excavated, moist blocks 
drying quickly With a sale crust. Most southern 
Australian clays containing mummified leayes can be 


disaggregated by immersion in approximate 7% wiv 
H,0, which has been heated. Salt in the Nelly Creek 
matrix interferes with disaggregation, and maceration 
is only successful if the blocks have been. either 
presoaked in distilled water to remove some of the salt, 
or if a detergent such as Quaternary O is added 10 the 
maceration mixture. Approximately 40% of the leaves 
obtained from a given macerate are translucent (Fig. 
2, A-C) while other specimens are black/opaque and 
much more brittle, Leaf remains obtained in this 
fashion are contained in. complete cuticular envelopes 
and treatment with hot H,O, (e.g. Scriven & 
Christophel 1990) followed by staining in crystal violet 
yields clean, easily photographed cuticle specimens, 
Cuticles from this deposit prepare easily and are in 
a better staie of preservation than those of any other 
Eocene deposit previously examined by the authors, 
Two frequent causes of cuticular abrasion or fragility 
are alkalinity of the matrix (or ground water) and 
presence of excessive fungal activity during, or prior 
to; fossilization. Based on the excellent preservation, 
bath of these factors were either absent or minimal 
during the burial and subsequent fossilization of the 
Nelly Creek leaves. All specimens figured im this paper 


AN EOCENE MEGAFOSSIL FLORA FROM NELLY CREEK, S.A, 67 


Fig, 2. Selected leaves from Nelly Creek deposit. A = specimen NCIOOO and is an example of Parataxon | (affinities unknown): 
B = specimen NCIOI2 and is an example of Parataxon 2 (Proteaceae), C = NCIOI7 and is an example of Parataxon 3 
(Myrtaceae) — it is the holotype of Myrtaciphyllum eremeaensis; Scale bars = 1 em, 


have been mounted in phenol glycerin jelly and are 
housed permanently in the Palacobotany Collection, 
Botany Department, Adelaide University. 

The Middle Eocene age is based on the well 
preserved pollen flora contained within the sediments 
(Alley 19897), As he reported, the Nelly Creek 
palynoflora correlates with the Lower Nothofigicdites 
asperus Zone of Stoyer & Partridge (1973) and with 
the Proteacidites pachypolus Zone of Harris (1971), and 
the Nelly Creek flora is correlative with the floras at 
Maslin Bay and Golden Grove, 


Floristics of the locality 


Collections made by N. F. Alley in 1986 and by the 
first and third authors and others in 1988 have been 
macerated to yield a collection of 220 broadleafed 
specimens (each representing 50% or more of a leaf) 
and numerous small specimens including Gynmostoma 
(Casuarinaceae) twigs, Podocarpaceae twigs, and 
various unidentified fruits and seeds. Broken leaves 
and detrital sievings from the macerations have also 
been kept for dispersed cuticle analysis. 


TABLE |. Leaf megafossil composition of the first blocks 
maceraled from the Nelly Creek clay lens. 


Purataxon Number of Specimens Affinities 


Number (% of the Flora) 

1 172 (64) UNKNOWN 

2 16 (6) PROTEACEAE 

3 1S (5.5) MYRTACIPHYLLUM 

4 3. (1) AGATHIS 

5 1 (0.5) LAURACEAE 

6 1 (0.5) BRACHYCHITON 

7 2 (1) PROTEACEAE 

8 3. (1) MONOCOT 

9 3. () PROTEACEAE 

10 tL (0.5) UNKNOWN 

ll 1} (0.5) UNKNOWN 

12 1 (0,5) PROTEACEAE 

13 tL (0.5) UNKNOWN 

14 25 (9) PODOCARPACEAE 

15 14 (5.5) GYMNOSTOMA SP A 

16 1 (3) GYMNOSTOMA SP B 
TOTAL 269 (100) 


D. C. CHRISTOPHEL, L. J. SCRIVEN & D. R. GREENWOOD 


68 


ty 


AS BOCENE MEGAPOSSU 


The 220 specimens recovered vould be divided into 
1B paritae based on bath muemimerphologieal features 
and cuticular struetures (fable 1), The analysis of the 
material clearly showed a damimance of the sample 
by one paralaxon (Paratixon 1. Fig. 2A, 3A-B). The 
leaves of Paratixon b ure generally microphylls with 
a lew beang classed as notophylls (sense Webh 1959). 
Allure entire margined and most display apices with 
4 general ovale to elliplicul shape, Following the 
descriptive terminology of Hickey (1979), primury 
venation of Parataxon | is pinnate with brochidodro- 
mous secondary venation and reticulate lertiary yeins. 
Civnerally, five orders of vein branching are present. 

The cuticle of Puraiwkxon | is hypestomaric with 
numerous slomates displaying either two or three 
subsidiary cells, Both abaxial and adaxial surfaces are 
densely covered with simple, collured trichomes (Fig. 
3A-B), This collared appearange Could represent torn 
tissue fram glandular wpices on the trichomes. bur the 
general exeellent preservation of the cuticles and. the 
large number of specimens sampled does not support 
that hypothes)s 

IU has fot been possible to determine the affinilics 
ot ihis dominant paicdtaxon, Several large Australian 
ruinforest families ean be easily recounised by their 
cuticular siructure. Such identifying features have been 
discussed for (he Lauraceae (Hill 1986), Proteaceae 
(Lange 1870) and Myrtaceae (Christophel & Lys 1984). 
Theretore. its possible to eliminate these families in 
the Weniification provess. However, several other large 
tunities, inclading the Fabaveue, Euphorbiaceae, 
Sapindaccae and Gleaceae all have taxa with leaves 
upproxumaling lhe venation puter and general 
macromorphology of the Nelly Creek dominant. 11 is 
ulsa possible thar the fossil could represent a (amily 
or lower level (axon Which is now eXtipet, and thus 
my reasonable maich could be forthcoming. 

There ure pwelye orber broadleafed  paratixa. 
Parstaxon 2.16 a lobed, servate Ileal which, while guite 
luge, is very brinle and has not been recovered as a 
coiplete leal (Fig, 2B), [usually vecurs as a pinnalely 
lobed specimen with (tree apparent lobes, These lobes 
are Inothed near their apex. Secondary venation is 
hrochidodromeus near the base of cach labe und 
semicraspedodromous near the apex when teeth are 
present. Culicles prepared from these leaves show that 
Ue leah is hypostamatic with numerqus slomtes on 
the abaxial surface possessing @ paracyle subsidiary 
cell arrangement (Fig. 3C) Vhs, coupled with the 
nunerous four-celled trichome buses observed an both 
surfaces Of the leal (Pig. 30, D), places the parttaxon 
m the Proteaeeae. While more detailed comparisons 


PLORA FROM RELLY CREEK, SA ow 


will be required for final identifieation, preliminary 
CRU athion indicates wu Simalarily in structure to several 
spegies OF Greville. 


The vein pallern of Parauimon 3 showing aumerous, 
close spyeed. high angle secondary veins forming a 
dhstinet intermargifal vein sugpests thal the paralexon 
belongs in the Myrtaceae (Fig. IC). Examination ol 
the ciitigle-confirms this With the presence of diagnostic 
lid or cupping cells on both surfaces (Fig 2B, F). This 
leaf type is one of the most variable insize and Shape. 
However, Christophe! & Lys W986) demonstrated that 
such interspecific variation i$ comiton within the 
family, They also demonstrated thal no obviuus foliar 
character or suite of characters defined genera within 
the family. and that the capsular fruited taxa and berry 
frumted taxa often numerically clustered together (were 
murphologicully similar). This interesting to mote that 
While the Netly Creck Myrlaceue bear some general 
similitrity (o genera of both capsular fe.g 
Lophoestemon) and berry Sruited tow. Syeveten) groups 
inthe family, there ms no close similurin to Eied/ yrs, 
The Nelly Creek Myrueiphylliim is tarmally deseribed 
in the Jollowmg section, Capsular Fruits with likely 
Mynaceae affinities have been recovered from a Nelly 
Creck macerate (Fi. SC), and will be deseribed when 
more material becomes available. 


The remaining 10 purataxa are all relatively rare in 
those samples processed to dare, All but four ure 
represented by only one specimen. Same of these 
remaining paratuxa are distinetive. and assignable lo 
families. und in some canes genera, 40 are worth 
discussing jp the overall floristic contextof the paper, 
The fipst of these broadleafed taxa |g assignable wo 
Avaihis (Araucarniceae) based an general form ane 
culicular siructure (ee. Hill & Bigwood 1987: Srockey 
& Ko 1986), 


A eoupurisun of all the leaves examined (220 
hroadlested and 49 microphyllous) can be seen Ih Table 
|. Paralaxon 5 (one specimen) cun be placect in the 
Lituriceue based on the size and nature af the stomatal 
ledges and alse the subsidiary cells (Fig. 4C_ 0) Hill 
IYK6). Parataxon 6 (one specimen) cun be assigned to 
Brachychiton (Sterculiacese) based on the hair bases 
and stomatal artaliecent, Because the specimen is 
Irigmentary (ane lobe) very bithle can be said as to its 
specific affinities. Jiterestingly, three of the otber 
parutaka ((Wo non-entire and one entire margin) can 
he placed in the Proteaceue. A final paratiaxen 
(Paralaxon §) is represented by tree speciinens and 
iy clearly a munocolyletioan based on the parallel 
yenatfon and the stomatal type 


Fig. 3. Cuticles of leaves ilustriied in Pig, 2. A = abasial cuticle of NCTONO (Pardon TB = adaxial cuticle ol NCIOOD, 
C= dbaxiil curicle of NCI (Panitixon 2); B= adaxial cuticle of NCIOI2; E = abaxial cuticle of NCTON (Holotype 
al Myriad ersmeaensis); F = adaxial cuticle of NCIOV, Insert for P isan enlarged view of u lid vell: Seale bars 


= 5 pm excep the insert where bar = 2.3 pm 


D, C. CHRISTOPHEL, L. J, SCRIVEN & D. R. GREENWOOD 


eo 
¥ 


AY 


“ 


hee te 


ae rm cre 
as vee 


AN EOCENE MEGAFOSSIL FLORA FROM NELLY CREEK, 5.A, 71 


Fig, 5. Miscellaneous structures from the Nelly Creek and Poole Creck deposits; A = NCISOL 


twig of Parataxon 14 


(Podocurpaceae) from Nelly Creek X¥; B = possible Proteaceae infructescence from Poole Creek Silcrete deposit X2: 


In addition to the 13 broadleafed parataxa, three 
microphyllous parataxa were Collected, These include 
one conifer and two distinet species of Gynoestomea 
(Cusuarinaceae). Based on macromorphological 
features, the conifer could be either Cupressaceae or 
Podocarpaceae (Fig. 5A). However, the cuticle clearly 
shows that this parataxon helongs to the Podocarpaceae 
(Fig. 4F), 


Approximately 20 twigs were recovered which were 
assignable to Gyminesroma, It has been shown that 
cuticle features are distinclive in extant species of this 
genus (Dilcher ef aul, 1990; Scriven & Christophel 
1990), and examination of the Nelly Creek specimens 
revealed thal two species were present. A cuticle of 
one of the two Nelly Creek types is shown in Fig, 4E. 
No fertile material has been recovered thus far. 


Although the taxonomic study of the flora is 
preliminary, 12 of the 16 parataxa recognised can be 
assigned to some formal taxonomic level. This means 
that at least a very generalized comparison may be 
mude with other floras and with modern vegetation 
types. 


C = NCI500 — leptospermoid fruit (Myrtaceae) from Nelly Creek X10. 


Taxonomic Description 


Order: Myttales 
Family: Myrtaceae 
Genus: Myrtaciphyllum Christophel & Lys, 1986 


Myrtaciphyllum eremeaensis sp. nov. 
FIGS 2C, 3E-F 


Diagnosis 

Architectural features: leaf shape elliptic, ovate or 
obovate. Size range: 3.5-13 cm long by 1,5-4 em 
maximum width. Leaf tip acute or attenuate, rarely 
acuminate. Leaf base acute, rarely obtuse. Primary 
venation pinnate, secondary veins straight, 
brochidodromous with a prominent intermarginal vein, 

Cuticular features: leaves hypostomatic, stomatal 
complex anomocytic, with between three and six 
subsidiary cells (three or four most common), 
Anticlinal epidermal cell walls angular — straight to 
slightly curved. Cells of both upper and lower 
epidermis equal sized; no striations visible on periclinal 
walls. Simple hairs infrequent (less than three per 
mim) on both surfaces. Hydathodes rare on lower 


Fig. 4. Miscellaneous cuticles from Nelly Creek parataxa, A = abaxial cuticle of NCIO03 (Parataxon 7 — Proteaceae); 
B = adaxial cuticle of NCI003; C = abaxial cuticle of NCIOIN (Parataxon 5 — Lauraceae): D = adarial cuticle of NCION, 
E = cuticle of NCI301 (Gynnestonia sp A — Parataxon 15); F = cuticle of NCI302 (Podocurpuceae — Purataxon 14): 


Scale bars = 5 yum. 


72 DC. CMRISTOPHEL. L J SCRIVEN & D RL GREENWOOn 


(abaxial) epidermal surface, apparently absent on 
adaxial (upper) surtece, 1d cells numerous on both 
surfaces With S-shaped to straight sinus showing no 
bended thickening or pecforadons (Pig. 3F-inser).. 
Epidermal cells Surrounding lid cells frequenily 
inndified inte a somewhat radial pattern — particularly 
on upper epidermis. 

Holeiype. Specimen NC 1017, housed in. the 
Palacohormy Collection, Botany Department, Adelaice 
University, as one uunuimified leaf and one euticle slide 
(NCC IOI7) 

Type dawaliys Nelly Creek, S.A. (29°19'S. 
J37°18' FB) 

Collector; D.C. Chitistophel 

Etymology: trom Eremean, reterring to the large, 
central Australian arid vegetution province used by 
1, A. S. Johnson and B. Boggs as a distributional 
region for Australian Myriaceae and Priteaceae (e.g, 
Johnson & Briggs 1981). The type locality occurs 
within this region 

Deseription of Holotype: Collected tn 1988. Leal 
Shanty dong by 22 min wide at position at maximum 
width, Effiptic. symmetreal with ullenuale apes (apex 
angle 35°) and acule base (57°), Secondary veins are 
straight. average angle 37° 

Cudele iypical lor jhe species. Stamates located yin 
ubaxaal surface with three To six subsidiary eelts. 
Average length of steanates 20 em (meant L/W = 1.0). 
Anticlinal epidermal cell walls angular — straight to 
slightly curved with no thickening or beading, Cells 
Of buth abaxial and adaxigl curigles equal size (mean 
20 » 20 im — runge IS—30 jun), No striations visible 
on periclinal walls Simple hairs rare an bath surfaces. 
Larve jnulticellular halt bases present, ind ne 
hydathodes visible on sample prepared from hulolype. 

Lid cells numerous an both surlices with S-shaped 
Lo struhe sinuses. showing no headed thickening or 
perfarations. Epidermal cells surrounding lid cells 
frequently. nodified inte radial pattern. Lid veil size 
= 20 = 20 pm, Density of lid cells 2 per JON 2 100 
fant section, 

Comparison will arher species: The first two species 
described lor (he genus could fot be distinguished by 
Jeaf arohiwerural tearunes, and cuticular characters were 
used (Christuphel & Lys 1986). The same situation 
applics to Miytaciphyllum eremvacnsts, as the 
Specimens weluded overlap both previously described 
species in maciomorphological und venation features. 
In cuticular features, however, M, eremeuensiy is 


distiet from Mo wneduleatee from the Tocene of 


Angléeses in that i) Hicks the sinuous anticlinal walls 
of the epidermal cells exhibited by the latter xpevies. 
M. eremensis- differs trom M, dewglasii froty Anglesea 
in having, numerous lid cells on both surfaces as 
compared (ou vomplete lack of lid-cells in the latter 
SpeLies. 


Physioznomic Interpretation 


Christophe) & Greenwand (J989)_ in discussing litter 
Jeposition in Australian rainioresis, demeonstruted that 
there was 2 predictable physiognomic signature for the 
forest types categorized by Webb (19591. OF the 22U 
broudlewed specimens recovered from Nelly Creek 
samples. fis possible to oteatsure (or estimate) the 
length und muadoum width for approximately 160), 
Results showed that there were no mesophylls present, 
while approximately 20% of the leaves (40% of baxal 
were nolophylls and 80% uf the leaves (60% of the 
tast) Were fucrophylls. The diserepaney berween 
ypecies dnd total loaves refleets the high frequency ol 
one microphyl! parataxon and the attendant rarity of 
most other parulaxa, This single paratuxon dontinations 
was reflected tw lesser degree im the margin type 
pemeniwe. with 88% of leaves (75% taxa) entire- 
margined. IF Nelly Creck leaf lenyth. maxinium width 
and position of maximum width are superimposed an 
the box dlagram of physiognomue signatures from 
Christophel & Greenwood (1988, Fig. 3) it bevames 
appurent that the Nelly Creck Nora does oot resemble 
Golden Grove or Anglesea, having much smaller leaves 
than cither of them. Even remembering the cuveut 
concerning inlerpretauion of small sample qumbers. 
there are several interesting subjective observations that 
ean be coupled with the above physiognoone data, 
Unlike the other Middle Eocene deposits mentioned 
curlier, there in no evidence oF drip tips in the Nelly 
Creek Mora. Additionally. very few germlings (senna 
Lange 1976) are present on lewt cuticles. and in peer 
the leaves from Nelly Creek can be considered more 
sclerophyllous. This includes such featares as penerally 
thicker cuticles, denser trichomes, and smaller, jon 
coriaceous or woody leaves, These fealures would (end 
tO sugvest u drier (or ceraiuly nrore seasonally dry) 
climate than the ther reported Eocene megulissl 
deposits, or allematively a much more depauperate soul 
nutrient level (Beadle 1963), 


Comparison with other Eocene Floras 


‘The first Impression af the Nelly Creek flora with 
ils tolul of 16 parataxa is one of clear dominance and 
low diversity. However, same af this can be most likely 
attributed to the small sample size and limited portion 
of the clay lens sampled) For the better known 
Australian Bocene lloras, the diversity is higher. For 
exumple. the most thorgughly studied clay lens at 
Atlesea has over 40 parataya (Christophel er a/ 
1987), Golden Grove hus over 30 parataxa (Barrett & 
Christophe! (990) und Maslin Bay is estimated at 
approximately 200 paratixa (Chinstophel & Blackburn 
1976) or perhaps as tow ws IS0¢L. S. Scriven unpubl, 
ita} 


AN FOCFENE MEGAFOSSIL FLORA FROM NELLY CREFK SA 73 


Tage 2. heed! Hitter composition af (we one meire square 
qeadrats ul Noah Creek, 


Quadrat One 
Lowt Numbers 


Quadrat Two 


Spevies Preven 
Leaf Numbers 


WM) A 9%) 

Cerarmperalun 

ite repretaalteen 68 13951 SB (29) 
Acacia 

dilacucarpi 29 (14) 7h UN) 
Buckinghania 

fevetginificre 21 (10) 301 PS) 
Lindsayenyrntas 

brachyandrus 14M) 77 4) 
Meralleasmia 

sensall flares 12 iG) iO (ay 
Charieras unytes. 12 (4) 4 (7%) 
Beilschodia 

olivanlra 4 14) $124) 
ul Draspyras 

hebecarpa R (dl 7 (3.5) 
Dissiliuriu 

laxinorviy 6 Oh 7 445) 
Fucntia hortensts 5 (2) 3 (4) 
Xenihesteman 

chrysanitits 3 (tN) I) 
Francisvodendron 

faurifelivn 441.5) 4 (2 
Oritey sp. tov 2h | (0.51 
Syunweiaen rarande 2 1 (os) 
Sarrepreryy alt, 

mantyane 2h + tbs 
Cullaphvtiin 

castialianiun 1 (sy fi 
Svoxyiuen 

ervihrocalyy (U5) ‘| 
Unknown A 2 j{1) iY) 
Unknown & } (th5) ui) 
Unknown C 0) 4+ (2 
Uidkiown D 0 2 tl) 
Unknown i) 1 (03) 
Unknown b Q) L WS) 
POVAL TAXA 19 18 
MtAL LEAVES DO i tM)) 202 (100) 


IUts possible to test the relanonship berween sample 
size and diversity in both fossil deposits and extant 
rainforests where the diversity ix known, When four 
random samples of 250 leaves each were tuken from 
the Anglesea lens mentioned above, the mean diversity 
was 18 43 (D. Christophe! unpubl. data). Similarly, 
recent collections from an extant Gyniestomut 
community ona hall hectare island in Noah Creek in 
the Daintree region of north Queensland showed that, 
alihough 75 different tree species occurred on the 
island, two litter samples (containing 200-300) leaves) 
had a diversity of less than 20 species per sample 
(Tuble 2), Therefore, the 16 prelimiaury parataxa 
recovered it Nelly Creek could easity represent less 
than hallof the expected rotal diversity for the fossil 


flora, and an even smaller traction of (he diversity of 
the forest from which it was derived. 


A more accurate estimation of the diversity of a Mora 
may be had trom a study of dis dispersed culicle 
(Rowett & Christophe! 1990). Samples of ¢hay frou 
Nelly Creck had un average diversity of 26 cuticle 
purataxa, while similar sized samples from Golden 
Grove yielded 25.32 parataxa (A. Rowell pers, 
¢omm.). Rowett reported that the samples were 
dominated by Tragments of Myrtaceae leaves — 
probably all belonging to Myrtaciphyllan eremuensis. 


At higher taxunonite levels. the flora has many 
elements common io other Middle Eocene floras. 
Golden Grove. Anglesea and Nelly Creek contain 
abundant (greater than 10%) Myrtaceae leaves. 
Similarly, Anglesea has approximately the same 
percentage of the flora made up of Proteaceae species 
as does Nelly Creck Gwnnesroma is found at Nelly 
Creck, Anglesea. Nerriga uod Maslin Bay. 
Brachychiten is known fron allot the Eocene localities 
except Nerriga, and Agathis is found at Mastin Bay 
and Nelly Creck, One interesting Moristic ditference. 
however, is in the representation of the Lauraccue. 
At Maslin Bay, Nerriga, Anglesea and Golden Grove 
this family 3s both plentiful. and diverse, white at Nelly 
Creck only one leaf has been recovered. Similarly, 
the Elacocarpaceae (atl. Sloanea/Elaeocarpus), which 
is well represented at Anglesea, Golden Grove and 
Maslin Bay. has not been recovered at Nejly Creek, 


At the specific Jevel, the differences are more 
pronounced. The entire murgined microphyll 
(Parataxon 1) at Nelly Creek is not known from any 
ther locality, Parataxon 2 (lobed Proteacede) tn also 
absent drom all other floras, The twa Nelly Creek 
Gvemustoma species ure taxonomically distinet from 
the comunon species at Anglesea. ‘The Podocarpaceac 
paralaxon at Nelly Creek is different to any reported 
froma the other localities, Comparisons of the 
Brachychiton and Agathis species have yet w be made. 
The Nelly Creek Mvyrraciphyllan species is definitely 
diferent from either species al Anylesen, 


While. brie comparison of the foliar physiognomy 
was made in the preceding section, the generalization 
cun be made that the floristi¢ elements at Anglesea, 
Golden Grove and Maslin Buy all show more. tropical 
and/or high moisture regime teatures. ‘These chree 
Middle Eocene deposits all have teaves with drip tips, 
prolific. high rank germiings, and noticeable quantities 
of leaves in Webb's (1959) mesophyll size class, in 
direct contrast to Nelly Creek. Although certain 
Gondwanic familics are shared between Nelly Creck 
and the other Middle Bocene deposits (e.. Myrtaceae. 
Proteaceae, Casuarinuceae, Podocurpaceue), the 
spccilic Horistic Composition and the physioguoniic 
signature is differem for Nelly Creek. 


i4 2 C. CHRISTOPHEL. | 


Cumparison with Silerete Floras 


Dacly studies ot the silerele deposits of northern 
Sih. Australia concentrated on the deseription ond 
ewnlutinnary iaportinee af some of the plants 
(Chapruan 937, Litnee (978, 182; Ambrose ev a/ 
970). More feven studies have attempted to address 
the importual aspects of the stratigraphy of the deposits 
and oF their comparative farstics (Greenwood eal. 
1890). 4 muyor problent with these silcrete floras has 
been Wie lack of sifatigeaphic continuity with ditt 
strata, aml thus (he ave has been di feult wo determine, 
“Vhis tas been bighlighted by Ambrose ef al, (1979), 
where ad possible Miocene dge is suggested carly in 
the paper and biter in the sume paper an bavcene age 
in supported, Inthe regent stidies Greenwood ef al, 
COMO) found that there were twer distitet macrofessil 
tHlorus inthe Poole Creek silerete lacality, and tiesed 
An there Gaxanonie eimpostian and nthe stritigraphy 
of nes -by sedimentary units they determined Unt une 
flora was restricted to the upper Eyre Formation 
(Middle Paceneh and the other to the Phadnis 
hurmation (Oligo-Miocene) sediments. Comparison 
benveen the Eyre Formation sileret# Turd (Eocene) and 
the Nelly Creek fossils of pulynologically determined 
Middle Fucenc age therelere becomes tnportant 

Prelinvinory comparisons do mot result in the definite 
conelision that the Eyre bormation silereses and the 
Nelly Creek flora represent the sume vegelatin, 
However. see common taxa are present and the 
COMPATISON Mose ceraMly needs to he mide more 
figourously When additional Nelly Creek material is 
available. In sapportal the Carrelation, Ciwyurastorned 
4s conunon in the silerele and al Nelly Creek. Plowever, 
commonly ees us ferale infructeseences in the 
former and only as vegeluudve remains im the batter, 
Laid reproductive oyaterial is revovered (rom Nelly 
Creek. vonspecificity cimmot he determined, Similatly, 
a lubed Proveaeeue leal very similar (oO Nelly Creek 
Parutixon 2 commonly occurs in the siluretes. Two 
wither pardtuxa from Nelly Creek, a narrow linear, 
enlire-niacsined Peoleaceac leaf and Brachyehiia also 
weeur inthe silercte. Again, lurthor studies are required 
lo determine conapeelfierty — particularly as (he Nelly 
Creck Brachveliion 1 unly a smyle lobe and henee 
even wrudimenniry charderer, Hike the number of lobes 
Prencot. cannot be compured, 


Within the siléretes there also. commonly occurs a 
flattened, woody reproductive structure (Fig. 5B). 
Bascu on silerete impressinis alone. the structure has 
not heen Wentitishle. and has aot been recorded from 
any other published fossil plant locality known to the 
nuthors. However ane specimen of this structure has 
now been recovered from a Nelly Creek mucerate, 
Alfhouyh (he specimen has fragmented. 1 ein be seen 
(hal eacth of the woody wedpes consists of two 


J SCRIVEN & D R. GREENWOOD 


Hattenud, appressed worxly bracts, Move detailed stddy 
is still required, but it would appear (hat the structure 
hus some similarity to @ Proteaceae cone — c.2 hke 
the Nattened infructescence ot /sapogon ur Lrvend rs. 
in the original paper describing silerste miteriul. 
Chapman (1937) Hgured a specimen und labelled its 
a “Banksia flowering tip” Thit specimen, however. 
does not resemble those ciseussed here. 

Some credence is given to the interpretation of the 
woody reproductive structures as Banksiae Tribe 
(Proteaceae) infructéscences hy the fact that the silcrete 
Moras contain serrate foliage ienthed by Greenwood 
eral, (990) as Banksiaeformis Hill & Christophel, 
which could fiave affinities with Dryandra 
Unfortunately, no sich Banksiaeforms leaves have 
been recovered at Nelly Crock. 

Other evidence does not support the correlitian. In 
addition lo the connmon Bartkyicefarmis leaves in the 
suleretes. other roothed Jeaves with possible affinities 
to either the Cunonucese of Elacwcarpaveae ure 
teparted from there (Greenwood eral. 1990) and are 
missing from Nelly Creek. OF particular mitecesc, the 
narrow, Sometimes filleate Myrtaceue leaves which hear 
similarity to Seeadynas, while common ote sileretes, 
arm also absent from Nelly Creck. Finally, the 
dominant, brochidod conus Parataxon J from Nelly 
Creek has nol been reported mm the silerete deposits 
(Greenwond ef af 1990). 

Phystownoimically, the sileretes contain larger leaves 
than Nelly Creek hits thus far yieldedl. and alsow higher 
percenlage of non-entire mare ined leaves. It is reported 
(N_F Alley pers. comm, | that blocks of clay wath larbe 
Jegves pecling: off therm were ungurthed on an eurhy 
expedition te the lovulity. Untortunately these blocks 
did no} survive transport ta Adeliide, and our more 
recent mater fas not contained such leaves, However, 
this serves [oO tlusinite the potentially mosaic 
distrmbutiine af taxa within the clay, and also highlights 
the need for additional collections. His certainly the 
cuse, lmwever, that some of the selerophyllaus nature 
of the Nelly Creek leaves is mirrored in the portion 
of the Poole Creek silcrete flora consatered ly 
Greenwood et af. (1990) to be Middle Bocene, 


Discussion 


The potential othe Nelly Creek (hora wradd to out 
hroader knowledge of Middle Eocene Australia Norns 
tas heen menuoned inthe torroduction, Examination 
av that flora more elosely hay emphasized this 
importance, Firstly, the preliminary taxonomic: aysesh: 
mene has shown that the flora has a very different 
composition to that of the other well known Australian 
Middle Bovenc microfossil floras. While seme of the 
major Gondwame fanvilies. mcluding the Protewcese. 


AN EOCENE MEGAPOSSIL FLORA FROM NELLY CRELK, S.A tn 


Myrticese, Casuarinaceae, Podocurpiceac and Arau- 
cumuceae are present im all Moras of thal age, the 
veneric und specific eomposition of the floras is 
differcot. Physiognoutcally, the Nelly Creek (lori is 
different from the Golden Grove. Anglesea. Mastin 
Bay nd Nerriga floras, being decidedly smaller leafed 
and lacking the numerous raintorest indicators (drip- 
Lips etc.) shown by those lloras. The inland position 
ol the locality is perhaps responsible for the difference 
in floristics and physiognomy seen at Nelly Creek, and 
ollr_averview OF Middle Eocene Austratia must be 
tempered Accordingly. 


The potontial importance of the Nelly Creck locality 
to our understaiding of South Austrtlia’s silerete floras 
hust also be cmphasived, While the evidence for 
positive correlation is poor Ute presence of certain 
udicstor Gx, such as ihe disk-shaped woody 
reproductive structure and the jucrow lobed Proteaceae 
Jeat in both deposits and inno others, must coritinty 
be taken as cneourugement tor further collections and 
Comparisons. 


Greenwood eral. (19%) suggested thal the ussumed 
Eocene elements of the silcrete floras mizht well 
represent deciduous seasonal vegetation types mixed 
with a wetter riparian element such as thase associated 
with jaonsoonal vine thickets in Queensland today, 
Such an interpretation for the Nelly Creek locality ix 
consistent with both the Known. elements of the Mora 
undalsa the physiognomie interpreéition, and w nore 
thorough search of modern forest types of this 
description will be made in the hope of identity ing 
further clements in the Nelly Creek Mora 
particularly (he dominant parutaxon. 


Acknowledgments 


We wish fo thank DrN. Fh Alley, 5. A, Department 
wf Mines and Energy aud Dr A. 1, Rowell, Botany 
Department, Adelaide University, for their assistance 
in the original colleen OF material for this study. and 
lor-cnitically reading the manuscript, The projeet was 
funded by ARC Grant A38931389 t D.C. Christophel. 


References 


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CHasyorHel. DC, (1984) Carly Tertiary Proteaceae: ‘Whe 
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(1989) Lvolutian of the Austatian Nora through the 
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_ & Bracwavry, D.'T. 1978) The Tertiary meaatassil 
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~ —— & Grebswoor, DR, (988) A comparison al 
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a 1989) Changes in climate and yeyelation 

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~ Ubatias. WOR a& Svank, AOR, (1987) The 

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__ & Lys. 8. Do (1986) Manimified leives of tear new 
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THE RESPONSE OF SOIL NEMATODES TO ENVIRONMENT STIMULII 
IN ARID SOUTH AUSTRALIA 


BY JACQUELINE MARY NOBBS 


Summary 


Teasecrons af tre Rovel Soatety of 3, Aust, (W992). W6¢2). FIAT 


BRIEF COMMUNICATION 


THE RESPONSE OF SOTL NEMATODES TO ENVIRONMENTAIL. STIMULII IN ARID 
SOUTH AUSTRALIA 


Nemiltodes ure known to form an yahydrobietic coiled state 
in response to desiccation in sil! Prony investigation of 
nematodes occurring inatnd areas (Mojave Deen, Nevada. 
U.S.A.7) the anhydrobietie stile was also found tw be 
represented by “cothng™ The activity of nematodcs can then 
be related to form. with “toiled” nematodes being inuctive 
or anhydrobioti¢ and “straight” nematodes being active, A 
prelimimary study was set up to investigate if “coiling” was 
a good indicalur of nematode achivity within arid soils, Also 
under investigation was the overall effect of environmental 
stimuli og the different nenwtode trophie groups. 


A site located on Plumbago Stanon pastonl property (near 
Yunta, South Austrilia) was selected lar sampling. The 
vegetation vonsisted of a low Chenoped shrubland dontinated 
by Atriplex Vesiceria. Soil sayuples (n=10) were taken every 
lwo months trom August 1985 (M2) to Octaber 986 (MIG) 
toa depth of 25 cor, Nematodes were extracted frony SQ tl 
of Sou. per sumple. 


The modified Bacrmann’s tunnel technique’ was used to 
extract the differen! nematode teophic groups. Afier extraction 
(over a three day period), the neniwitodés wete heat killed and 
fixed in 2% formatia (4096 formaldehyde) ‘The different 
tropluc proups were then counted. The tmopie proups 
coasisted of:- ommivures (mainty dorylaims), baytenial feeders 
(mainly chabditids), fungal feeders (mainly aphelenchs and 
tylenchy) and plant parasites (mamly Tlenchortyng dus tobari 
Suucr& Annells, 1981 and Teletylenchus aatitars (Collbran. 
191) Siddiqi, 1963). The exteaction efficiency was found ta 
be-aboul 63% and the counts were adjusted accordingly. 


1200 » omnlvoras = 0 
bartarial feedars =U 
jungel feeders =A ca 
plary paragtes = 7 a 
Ad Kata] pemalegee & © , 
5 / wa 
a 
ano + f/ 


mean no. nematodes / £0 mi snil 


month 


Fig. 1. Mean numbers of omnivores (o), bactenul feeders (77), 
fungal feeders (A). plant parasites (V7) and total nematodes 
(%) extracted from 50 ml of soil (n=10) using the modified 
Baermann’s funnel technique from samples collected every 
{wo months tron August 1985 (2) tw October 1986 116), 


Anhydrobiotic (“cated”) and active (“strarhr") nematodes 
Were extricted lising the Hot Formalin method? The method 
involved killing the nematodes in the soil with hor formalin 
(90°C). then separating the nematodes Irom the soil asing 
a solution containing Separan NP | 10% (Mow Chemical Ltd) 
(Q75g/V tap water). The neumtodes were then separated inte 
“coiled” and “seraizhe” forms and counted, The extraction 
efficiency of the Hot Formalin method was found to be #hout 
H% and the counts were adjusted secordingly Due to loss 
of marerial st was not possible lo extract nematodes tram the 
August 1985 (M2) and February 1986 (M8) sumples using 
the Hot Formalin method. The Hot Formadin metheut tended 
lo exiraet ntere nematodes than the ntodified Baemann funnel 
technique possibly because the Hot Forinitlin method extracted 
diteetly from soil while the modified Baermann’s funnel 
Methad relied on movement of nematodes into a collecting 
dish, 


As with other arid regions. bacterial feeders were the 
most abundant trophic group found inthe samples thraughout 
the sampling penod (Fig. 1). The other trophic gruups 
occurred In much Jower numbers, From August 1Y85 (M2) 
to April (986 (M19) the total mean number of nenvatodes 
extracted was relatively constant, averaging around 200 300. 
Over the sume period the mean nuriber of “coiled” nematodes 
was much gregier thin “straght” nermatedes (Fig, 2). 
fluctuating between 300 (M4) und 600 (MO) with the morn 
number -of “wiraight” nematodes remaining fairly constant 
throughout 


‘coiled’ = @ 

\ 

streignt) = © 
qwial av 


1500 


1200 


£00 


mean no. nematodes /50 ml soil 


month 


Fig. 2. Mean numbers of “coiled” (@), “Straight” (il) and 
total nematodes (4) extracted from 50 ml of soil (n=10) 
using the Hot Formalin method from sumples collected 
every two months from October 1985 (4) to October (986 
(16) excluding February 1986 (8), 


TS BRIEF COMMUNICATION 


However, from June 1986 (M12) to October 1986 (M16) 
there Wis ain increase in. the mean number of alt frophic sroups 
(except plant parasites), with the bacterial feeders showing 
the greatest Inercase, During the same period there was also 
a large increase inthe mean number of “straight” nematodes 
with a sharp deercase in numbers of ‘cailed” nematodes (M12) 
which remained fairly constint afterwards, The change in Jon 
of the nematodes was, therefore. closely correlated wilh the 
Increase in numbers of nematodes, particularly the bacterial 
feeders. The chinye in form and inerease in numbers of 
nentaivdes could reflect Increased activity of (he ami¢ro-flora 
within the sail ecasystent, 


Rainfall muy have heen the trigger lor the ingreased activity 
of the nematodes, The region under study usudlly has the 


“a 
o 


fj * 


an i 


| ® hs \ 
Le A A INC 
i ow \ / 
‘ VAX 


monthly ranfall 


ov 25 4 8 68 7 @ 3 45°12 75 74 15 16 
month 


Fig. 4. Monthly rainfall (mm) recorded at Plumbigo Stition 
homestead from July 1985 (1) to Oetaher 1986 (16). 


'Demeure, ¥., Ereckman, D, W. & Van Gundy. S, D. 
(1979) Journal of Nematology I, 189-195, 


“Freckman, D. W. & Mankau, BR. (1986) Pedobiulogi:n 24, 
120-142. 


‘Schindler A. F. (1961) Plant Disease Reporter 45, 747-748, 


+Freckman, D. W., Kaplan, D. T, & Van Gundy, S. D. 
(1977) Journal of Nemarology 9. 176-181. 


*Freckman, D. W. & Mankau, R- (1977) Feology Bulletin 
(Stockholm) 25, 511-S]4. 


highest rainfall and lowest temperatures diving the months 
April ky Qetober and the dricst and hottest months fron 
November i March. Fig, 3 shows the rainfall recorded over 
the sample period at the homestead of Plumbago Station (about 
(+ kim from the sample sue), Over the first lO months of 
sampling there were large Muetuarions in riamfall while the 
final six months hada more even distybulion, The Goal niemth 
had (he highest natal of the sampling period. The more 
sustained period of cainta) aver the list six nisattis.of sampling 
was mutehed with inereased numbers of nematodes and 
increased numbers of “straight” wr detive nemafades. 


In other arid areas the activity of the nematudes was found 
to be exhibited ax a “pulse” phenomenon®, with an 
environmental ‘trigger’ (ic, rainfall) causing rapid: mereaae 
in numbers flowed by a capid decrease when the soil dries 
out. The bicterial feeders were particularly well adapted ta 
a-cyele of dehydration aad rehydration, The rapid response 
of the food source (bacteria) to appropriate environmental 
slimulil and the short lite cycle of the nematades (im some 
cages only 6-7 days) allows bacterial feeders ly increase in 
numbers when conditions are favourable. In this study, the 
environmental “trigwer” Was found 16 be raintall. 


Notrient urnover in soils of other arid regions were found 
fo-he influenced by nematodes as consumers of bacterta and 
yeast (during the first stages of decomposition) and fungi (as 
decomposition advanced)’, Further stiidies on the role of 
nematodes. in nutrient turnover may be helpful when (naking 
at the ceology a? arid region siils and may be useful in 
assessing the impact of overgrazing and mining, on soil 
ecology. Nematodes could be used ts manitor Jevels of 
microbial activity within the sail.as activity of nematodes can 
be measured through extraction of “coiled” and. “straight” 
forms. which could telleetactiviry of the find source, 


°Whitford, W. G., Freeckman, D, W., Elkins, N. Z., 
Parker, L. W., Parmalee, R.. Phillips, J. & Tucker; 5. 
(1981) Soil Biology and Biochemistry 13, 417-425. 
'Whitford, W. G., Freckman, D. W., Santos. P. E., 
Elkins, N. 4. & Parker, L. W. (1982) #7 DW Freckman 
(Bd.) “Nematades in soil ecasystems” 98 116. 

‘Santos, P. KF. Phillips, J. & Whitford, W. G. (1981) 
Lvoluyy 62. 664-664, 

Whitford, W, G., Freckman, D. W., Parker, L. W. 
Schaefer, D:, Suntos, PK & Stwinberger, ¥. (1982), Proc 
VII International Collaquium of Soil Zoology, Louvaim-ti- 
Neure (Belgium) 49-59, 


JACQUELINE MARY NOBBS, CABI Institute of Parasitology, St, Albans, Herts, U.K., Al4 OXU, 


CYSTICERI OF TAENIA HYDATIGENA (CESTODA: TAENIIDAE) IN AN 
ENTELLUS LANGUR (PRESBYTIS ENTELLUS) 


BY MICHAEL O’ CALLAGHAN 


Summary 


Transactions af the Reval Sucteiy of S Aust (992), WO), 74, 


BRIEF COMMUNICATION 


CYSTICERCI] OF TAENIA HYDATIGENA (CESTODA: TAENIDAE) 
IN AN ENTELLUS LANGUR (PRESBYTIS ENTELLUS) 


In May 1982 an adult female Entellus langur monkey 
(Presbytiy entellus) died al the Adelitide Zoological Gardens 


following rupture of the uterus in the terminal stages of 


pregnancy. During autopsy three large cysts 17-31 em in 
diameter were found attached to the greater omentum. Each 
cyst contained a single cestode cysticercus with a large bladder 
16cm in diameter (Fig. 1), 

The rostella were dissected from each seolex and mounted 
en face in DeFaure’s medium for examination of the rostellar 
hooks, Each rostellum wits armed with [4 large hooks 
183-189 jam (mean 188 am) in length and fourteen small hooks 
129-144 pm (mean 132 pin) in length, morphologically 
resembling (hose of Tenia Avdatigena' (Rigs 2, 3). In 
addition the size of the hooks conformed lo measurements 
of adult specimens of the same species from dogs in 
Australia? and iLwas concluded that the cysticerci recovered 
rom the monkey are metacestodes of Taenia hydatigena. The 
scoleces have been deposited in the Australian 
Helminthological Collection, South Australian Museum 
(AHC.S 42153) 

The metacestode of 77 Avelatigena has an exceptionally wide 
host range, principally in artiodactyls but occasionally also 
in perissadactyls, rodents, lagomorphs, marsupials and 
primates 4. Abuldase (1964) listed species of Cercupithicus, 
Mavacus, Papie and man as hostsof T Aydativena, however 
his and subsequent reports of this parasite in primates™ do 
not record Preshytiy as a host. Cysts from Macacus 
cynomolgus in Vietnam recently described us 7 yaigoni appear 
to be similar to 7) Aydativena’. Recently, coenuti 
morphologically similar to 7) mutticeps or T. serialis were 
reported in Presbyis obscura raised in captivity ina number 
of 2005 in the U.S.A,4 

The more familiar langurs belonging to the venus Presbvris 
are from India and Southeast Asia, P enrellus (the Hanuman, 
Sacred or Entellus Jangur) the largest of the langurs, is tative 
to India, Sri Lanka and Nepal and is known to live close to 
towns and villages, The langur examined at the Adelaide Zoo 
was imported from Sri Lanka ten years prior to its death and 
may hive acquired the infection either here or overseas, There 
was no pathological reaction associated with the presence of 
the parasite, This finding constitutes a new host record tor 
7. hydatigend., 


Figs 13, 1, Cyst of Tien hyeetigena trom the zreater 
omentunt of Prexbytis entellus; 2 & 3, rostellar hooks (Bar 
seale = 0.10 mm). 


'Verster; A. (1969) Onderstepoort J. vet. Res. 36, 3-58. 
“Beveridge, I. & Gregory G. G. (1976) Aust. vet, J, 52. 
369-373. 

‘Abuldase, K. 1. (1964) Tueniata of animals and man and 
diseases caused by them. Essentials of Cestodology Vol, 4, 
Ed, K. 1. Skrjatbin. Akademia Nauk SSSR- English 
Translation by Israel Program for Seientific Translations, 
Jerusalem, 1970. 


“Presidente P. J. A. (1979) Int. J. Parasitol. 9, 251-355, 
‘Myers, B. J. & Kuntz, R. F. (1965) Primates 6, 137-194, 
"Kuntze, R. E.. & Myers, B, J. (1967) Primates 8, 83-88. 
"Le Van Hoa. (1964) Bull, Soc. Path, Exot. 57, 23-27, 


‘price, T. C,, Dresden, M. H., Alvarsdo, T., Flanagan, 
J, & Chappell, C. L. Am. J. trop, Med, Hyg, 40 (4), 
514-520 


MICHAEL O'CALLAGHAN, Central Veterinary Laboratories, Department of Agriculture, GPO Box 167], 
Adelaide, 8. Aust. 5001 and JAN BEVERIDGE, The University of Melbourne, Veterinary Clinical Centre, 


Princes Highway, Werribee. Vic. 3030. 


VOL. 116, PARTS 3 & 4 
30 NOVEMBER, 1992 


Contents 


Transactions of the 


Royal Society of South 
Australia 


Incorporated 


Goonan, P. M., Beer, J. A., Thompson, T. B. & Suter, P. J. Wetlands of the River Murray 
flood plain, South Australia. 1. Preliminary survey of the biota and 
physico-chemistry of ten wetlands from Chowilla to Mannum. - = 

Rowett, A. I. Dispersed cuticular floras of South Australian Tertiary coalfields, part 2: Lochiel 

Beveridge, I. & Durette-Desset, M.-C. The morphology of Rppastrong yas Pear a 
parasite of native Australian rodents - - - - 

Davies, M. Early development of Limnodynastes terraereginae and L. fletcheri (Anura: 
Leptodactylidae: Limnodynastinae)  - - - - - - - 

Beveridge, I. & Durette-Desset, M.-C. A new species of trichostrongyloid nematode, Odilia 
bainae, from a native rodent, Rattus fuscipes (Waterhouse) - - 


Brief Communications: 
Green, J. D. & Shiel, R. J. A dissection method for determining the gut contents of calanoid 
copepods - - - - > = - - - > = 
Boulton, A. J. “Rollers” and “Carriers”: Field observations of carrion removal by trogid 
beetles (Omorgus strzeleckensis) in arid north-eastern South Australia 
Davies, M. & Watson, G. F. Redefinition of Uperoleia littlejohni Davies, McDonald & 
Corben (Anura: Leptodactylidae: Myobatrachinae) - - = - 
Greenslade, P. New records of Mesaphorura (Collembola: Onychiuridae, Tullbergiinae) 
species from Australia, Macquarie Island and the Antarctic- = 3 
Read, J. L. Ecological and biological notes on the rare plant Hemichroa mesembryanthema 
F. Muell (Armaranthaceae) - - - - - - - - - 
Morley, T. P. Eggs and incubation in the Australian lizards Amphibolurus nobbi and 
Eremiascincus richardsoni - - - - - - - - - 
Johnston, G. R. Relictual population of Tiliqua scincoides (Sauria: Scincidae) in north- 
western South Australia- - - - - - - - - - 
Edmonds, S. J. A note on Phascolosoma turnerae Rice (Sipuncula)- - - - - 
Tyler, M. J., Aslin, F. W. & Bryars, S. Early Holocene frogs from the Tantanoola Cave, 
South Australia - - - - - - - - - - - 


PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS 
SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000 


81 
95 


109 


li7 


123 


129 


133 


137 


141 


145 


147 


149 
151 


153 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


VOL. 116, PART 3 


WETLANDS OF THE RIVER MURRAY FLOOD PLAIN, 
SOUTH AUSTRALIA. 
1, PRELIMINARY SURVEY OF THE BIOTA AND PHYSICO-CHEMISTRY 
OF TEN WETLANDS FROM CHOWILLA TO MANNUM. 


BY P. M. GOONAN, J. A. BEER, T. B. THOMPSON & P. J. SUTER* 


Summary 


Qualitative data were collected on the water chemistry and aquatic invertebrate fauna from ten 
wetlands between Chowilla and Mannum on the River Murray flood plain in South Australia. 
Sites were separated into two main groups that corresponded to freshwater wetlands connected to 
the River Murray, and wetlands with TDS concentrations >1000 mgL" that were isolated from the 
main channel. Wetlands with TDS concentrations <1000 mgL’' were generally low in nutrients, and 
characterized by the dipteran C/adotanytarsus sp. and the shrimp Paratya australiensis. The more 
saline wetlands were high in nutrients and characterized by the presence of dipterans such as 
Proclaims sp., Ephydridae and Culicidae. 

Phosphate and nitrogen concentrations from most sites exceeded critical levels for eutrophication. 
Nutrient enrichment was indicated by the high chlorophyll concentrations recorded from most 
wetlands. These results indicate that nutrient levels entering the flood plain need to be reduced to 
minimize the risk of nuisance algal blooms during low flow conditions. 

KEY WORDS: Wetlands, River Murray, biota, aquatic invertebrates, physico-chemistry. nutrients, 
salinity, multivariate analysis, South Australia 


Transacriens of the Revel Soden of NS Aust W982). M63). BL 


WETLANDS OF THE RIVER MURRAY FLOOD PLAIN, SOUTH AUSTRALIA. 
1, PRELIMINARY SURVEY OF THE BIOTA AND PHYSICO-CHEMISTRY OF 
TEN WETLANDS FROM CHOWILLA TO MANNUM. 


by P, M. Goonan, J. A. Beer, T. B, THOMPSON & P. J. SUTER* 


Summary 


Goonan. P.M. Beer J A THompson. TOR, d& Suter. PY, (1992) Wetlands of the River Murray tloext pln, 
South Australia, 1. Prelimimtry survey of the biota und physice-chemisn'y of ren weelands from Chowilla ky Mannum. 
Trans. R. See. S. Alist, W6(3), 81-94, 3 November, 1992, 

Qualitanve data were collected on the water cherusiry and aquatic invertebrate fauna trom ten wetlands between 
Chowilla and Mannum onthe River Murray fload phon in South Australia. Sites Were Separated into two main 
Efalips thal corresponded ty freshwater wetlands connected to the River Murmay, and wetlands @ith TDS 
Epnicentratians > 1000 mgh! whit were Isolated from the main channel, Wotlands with TDS. concentrarians 
<(000 ing" were generally low in nutrients, and characterized by (he diptenin Chitoruivrersies sp and the 
shrimp Pararva wusrriiensis. The more saline wetlands were high in qutirionts and characterized by the presence 
of diprerans such as Prowlaedias sp., Ephydridue and Culicidae, 

Phosphite und nitrogen comcentrations from most sites exceeded eritical levels (or culrophication. Nutrient 
conchment was indrewed by the hiwh chlorophyll concentrations recorded from mose wetlands These results 
indicate that nutrient levels entering the Mood plain need bs be reduced i minimize the risk of nujsance algal 


blooms diting low flaw conditions. 


Key Woods: Wetlands. Rivery Munay.biotl, aguatic invertebriles, physico-chemistry. nurricnts, saliiits multivariate 


analysis, South Australia 


Introduction 


Over 1600 wetlands are distributed throughout the 
River Murtay Mood plain, lower lakes and Coorong 
in South Australia (Pressey 1986), Whereas many of 
these Were included in 4 recent survey: of River Murray 
wellands (Thompson }986), lithe has been published 
on therr biota and physico-chemistry, Thompson (1986) 
provides some information on the water quality and 
dominant fora and fauna of the 248 wetlands included 
in his study, Gedeles (984a d& bh, 1988) gives a detailed 
account of the limnology of Luke Alexandrina over 
several years, whereas O'Malley & Sheldon (1990) 
describe the results of a survey of the biological 
communities of the Chowilla Mood plain, Birds have 
been described from some areas (Tubbs 192k: Sehodde 
& Glover 1955; Mack I961; Cox 1973; Simpson 1973a) 
and Simpson (19736) discussed the distribuhon of the 
mammals, reptiles and amphibians between Mildura 
aml Renmark. Lioyd & Walker (1986) reported the 
distribution and conservation status of the syoall 
freshwater fish throughout the lower River Murray 
fland plain. 

This paper presents the results of u preliminary 
survey conducted during May-June 1991} on the aquatic 
invertebrate assemblages and physico-chemist’y of LO 
wetlanus distributed from Chowilla to Mannum. The 


~ Ausiratian Centre for Water Quality Research, Private Mant 
Bag, Salisbury, S. Aust, S108 


“Linyd, L., Moller, J, & Balla, S. (1984) “Berny Fvaparation pene ration 
Basin Study" (Dept Zoology. Univ. of Adelaide, Unpubl, 
Report tor NPWS.) 


aims of the survey were to describe and compare the 
limnology of flood plain wetlands with different 
hydrology and geomorphology, including anabrahches, 
swamps and Jukes. The emphasis of the work wes (o 
sludy the biota and water chemisiry of regulated 
wetlands. focussing on evaporation basins. This survey 
1s part of a larger study which ats ta (1) generate a 
comprehensive baseline and comparative database on 
the aquatic biota and physico-chemistry of selected 
wetlands throughout the River Murray flood plain in 
South Australia, and (2) investigate the effects of 
various changes in the hydrological management of 
regulated wetlands, 


Materials and Methads 


Selection of study sites 

The location of study sfles wits based on those 
previously investigated by Thomspon (1986) and Lloyd 
etal, (1984)" to enable some comparison with the 
available data from previous surveys. Additional sites 
were sampled [rom sonic wetlands to examine 
between-site variation, 

Pilby Crock was the only Wetland included in this 
survey not previously studied by the above workers, 
Sites were located on either side of a causeway which 
restricted water flow, enubling comparison between 
two sites in closé proximity with different hydrology. 


Herlands surveved 
‘The wetlands sampled in this study were distributed 
from the Chowilla flood plain te north of Mannurni. 


x2 P.M, GQONAN, J A. BEER, T, B THOMPSON & FB J. SUTER 


TABLE 1, Cacetion and physical characteristics of the 10 wetlands sarveved alang the River Marvay Pov plate in Saath 


Avsinulta during May-June 1990) 


Siw Notsi" Wetland Acronym Location Area (ha) Hydrology and Geomerpholagy 
1.2 Pilhy Creek PILC 33"590'S  $40°53' Ri 5 Permanent flood plain 
anabranch 
Clover Lake chai 3490'S 140S46/E 140 Intermittent Mood plain swarnp 

4 Lake Merreti LMER 34°08 HO4S EF 3960, Perinmcnt regulated tlood plata 
Iuke 

i? Pisher Ch Evap. Basin DISC 3414'S A 260 Permanent regulated Howd plat 
Anabruyeh 

§ 10 Kitarapko Evap, Basi} KAI'S, KRADN 34°26 8) 140°38°E 42,360 Complex of permanent 
regulated flood plain lakes 

Ul Berry yap. Busty BERB S491R'S dBA 325 Permanent regulittyd flood plain 
4Wwant? 

1-15 Ramco Lagoon RAML 3440'S 130°95'E 1 Permanent regulated Mood plain 
lake 

10. 2] Devon Downs North DEVD 34°96'S 130°S6'E 120 Permanent Nood plain lake 

4 Worngulla Lagoon WONT. 3443'S 139°33'E 120, Pertpanent Mood plain swap 

16-18 Like Carlet IL.CAR 3492'S 139°S1'E 330 Permanent food plain swamp 


“Sie S from Lake Woolpoluol (34 "02" 8. 140743" Ri was omitted ax sumples were not preserved. 


Details of the location, area. hydrology and 
geomorphology are given in Table 1. 


The location of sampling sites is shawn in Fig. 1. 


Specific site coordinutes and Ueseriptions of the 


deminant vegetation are given in Appendix 1, Each 
site was designated wath an acronym and number 


Collection and analysis of samples 


At cach site, the sampling area Consisted ol a 20 im 
section of shoreline representative of thar part of the 
wetland, Sites were sampled in May-June 1990 daring 
a rise in the River Murray hydrograph. with a flow 
of about 30000 ML/D recarded atthe §. Aust. border 
(Unpubl, Murray-Darling Basin Commission records), 


Field measurements mide at cach site were pH (1C1 
21L portable pH meter), conductivity (ICT 303 ATC 
conductivity meter), waler temperature, dissolved 
oaygen (YST model 58 dissolves! oxygen meter). and 
Secchi disc transparcacy. Surface water samples were 
collected and stored in air-tree, wurtight bottles on ive 
betore laboratory anglyses for nutrients (mtroyen, 
phosphorus and carbon fractions), pesticides, and 
Toujur iony (Gat. Mg*>. Na! K°, COP, HEO,, 
$O/-and Cr), 


Analyses of NH,. oxidised nitrogen (NO_N), 
flissolved reactive phosphorus (DRP), 807° and Cl 
were made using a Skalar automated flaw analyser. 
while HCO”, CO,? and alkalinity were determined 


7Anon (989) “Analytical Methods Manual Tnarganic 
Chemistry” (Saue Wir Laboratory, EB. & WS Bepr, §& 
Aust. S§.W.L. Report No. 30.) 
‘Anon (1990) “Anilytival Methods Minual Organic 
‘Chanustry” (Stute Water Laboratory. CE. & WS; Dept, 3- 
Aust, SW.L, Report Na 324 


using titration against a HCL, siandard solution. Total 
Kjeldahl nitrogen (TKN) and total phosphorus (TP) 
anulyses were made with 4 Technicon autoamilyser and 
Spectrophotonicter. Cations were amelysed using @ 
Labtest model Y-25 inductively, coupled plasma 
emission spectrometer fitted with a polychromator 
Dissolved and total organic carbon were measured with 
a flame ionization detector. Pesticides were extracted 
in hexane and analysed using a Varian 3300 gas 
chromatograph. All procedures are described in detail 
in two methods manuals produced by the E. & W.S. 
Department. South Australia”, 


Aquatic invertebrales were sampled from the littoral 
zone at cach site using a 30s sweep sample with a 
200 win. mesh dip net. Samples were preserved in 5% 
formalin and returned to the laboratory for sorting and 
identification, Invertebrates were identitied to the 
lowest practical taxonomic level using CSIRO (1970), 
Smith & Kershaw (1979), Williams (19808). Matthews 
(980, 982), Smirnov & Timms (1983), Wiederholm 
1983). Merritt & Cudimins (984). Hawking (1986), 
and several unpublished keys prepared by one of us 
(PS). A voucher collection is maintained for all taxa 
recorded from the River Murray Mood plain in South 
Australia uf the BE. & WS. Dept, State Water 
Laboratory, Rolivar, S, Aust. 


Water samples tor analysis of chlorophyll were 
processed in the field by passing a known volume of 
water through a 1.2 pm Whatman GF/C filter disk. 
‘The GF/C filter was placed in a centrifuge tube 
contaming 95% ethanol, which was then wrapped in 
alfoil and stored on ice, Samples were centrifuged und 
then analysed in the laboratory using a Pye SP8-100 
tUtraviolet spectrophotometer at wavelengths of 750), 
665 and 649 no, Chlordphyll a and b concentrations 


SOUTH AUSTRALIAN WETLANDS 83 


'S 
|» 
ani 
4° 2 ; 
s 12 Murra ENMARK, | 
2 13008 Y BENMARIT. 
N e 15 “WAIKERIE 7 | 
BERRI 2% 
C) 
1° | 
BLANCHE TOWN, 10, 
9° 8 | 
O) 
LOXTON 
SWAN REACH | 
e 1 
21\ \= 
20" ie 
194 IS 
SOUTH AUSTRALIA iS 
18 00! 716 | 
MANNUM,, | 
MURRAY BRIDGE | 
i) 
Lake 
Alexandrina 
— 
=) 
km 0 10 20 30 40 50 km 
———— 
SCALE 


Fig. |. Map of the River Murray flood plain in South Australia with site locations and numbers. 


were calculated using the cquations developed by 
Wintermans & de Mots (1965). 

Collections of macrophytes and riparian vegetation 
were made at each site (see Appendix 1), and 
representative samples retained as voucher specimens. 
Identifications were made according to Aston (1973) 
and Jessop & Tuelken (1986). 


Data analyses 
All biological analyses were based on the 
presence/absence of the aquatic invertebrates recorded 


from the 20 sites sampled. The sampling technique 
used in the survey resulted in the collection of many 
semi~aquatic and terrestrial species that were associated 
with vegetation in the littoral zone. These were omitted 
from the analyses. 


Sorensen’s index of community similarity (cf. 
Hellawell 1978) was used to group the sites on the basis 
of the composition of the fauna at each site. Clustering 
of sites was summarized in a dendrogram showing the 
degree of similarity in aquatic invertebrate composition 
among sites. 


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SOUTH AUSTRALIAN WETLANDS 85 


Ihe difference in aquatic invertebrate specres 
composition within and among wetlands was analysed 
by multivariale procedures, Relalionships among sites 
were examined by the ordination procedure of 
deirended correspondence analysis (Hill & CGsuch 
1980: Gauch 1982), using the program DECORANA 
(Hill 19798), Salinity measurements were 
superimposed onto the DECORANA plots to reveal 
relationships helween community composition and 
salinity (ef. Williams. e7 al. 1990). The hierarchical 
classification procedure of two-way indicatar species 
analysis (Gauch & Whittaker J98}; Gauch 1982), using 
the prograin TWINSPAN (Hill 19796), was curried aul 
to group similar sites together in clusters. Indicator 
Specres refer to the prclerential taxa used by 
TWINSPAN to distinguish the clusters. ‘The 
TWINSPAN program was run using the default 
options. 


‘The data generated by ‘Thompson (1986) and Lleyd 
er al. (984)! were not included in statistical 
comparisons with the results from the present survey 
due io differences in the methods and objectives of each 
study. Only general trends in the dats from these earlier 
stunties are discussed. 


Resulls 


Haren chenistry' 

The physico-chemical data are given in Tables 2 and 
4 As the profiminary survey consisted of only one 
sample per site, m1 dita are available concerning 
fluctuations of the various physico-chemical paramercrs 
with season and changes in water level. Consequently. 
only major trends in the data will be highlighted a¢ this 
sage. 


tonic voncentration 

Willams (1967) classified any water with o 
ecoboentratian of total dissolved solids (PDS) greater 
than 3000 mal.” as “saline” Based on this definition, 
Berri Evap, Basin, Ramco Lagoon and Pilby Ck 
(PILCI) were saline when sampled. Other wetlands 
to approach this level included Claver Lake. Disher 
Ck Evap, Basin (DISC7), and Katarapko Evap. Basin. 
Converting wnic concentrations ante ionic equivalents, 
waters [rom these wetlands. were dominated by sedium 
and chloride, and had tonic stoichiometries similar to 
thal af seawater (i.e. Nat > Mg?* > Catt > KS 
Cl > $80 > HCO, ), The ofily deviations in ionic 
trends among this group of wetlands were Katarapka 
Evap. Basin and Clover Lake. which had anionic 
stoichiomernies similar tothe more freshwater graup. 

The remaining wetlands had TDS concentnitions of 
Jess than 1000 mgL7, Sodiuny and chloride were also 


the dominance iens, although they represented smaller 
fractions of the total cations and anjons respectively, 
Cusjonic stoichiometry was the same as the mure saline 
wetlands, but the anionic stoichiometry differed in thar 
bicarbonate dominated sulphate ion (ie. Clo > 
HCO; > SO." > CO,*). 


Jonic composition 

An inverse relationship was evident between sodium 
jon and calcium/magnesium ions, with sodiurn 
hecoming more duminant with increasing TDS, 
Potassium ions represented very low fractions of the 
total cations from all wetlands. The proportion oi 
chloride to total anions increased with increasing TDS, 
while the proporuion of bicarbonate decreased. 
Sulphate contributed 10-24% of the total anions, with 
the higher proportions generally being recorded fron 
the more saline wetlands. Carbonate ions were detected 
from the more alkaline wedands [pH 8.8-9.4), reflecting 
\he effect of pH on the dissolved CO, equilibrium 


Nutrients 

Ammonia was present in higher concentrations than 
NON oat most sites. although at sume the 
ooncentralions af bath forms of dissolved nitrogen were- 
negligible Ge, <0.01 mgL'), The highest NH, levels 
were recorded from three of the more saline sites 


(PILC! RAMLIS, BERBIN. The highest NO,-N 


concentration was tecorded at DEV D2. which also 
had a high NH, concentration compared with the 
ollier Freshwater siles. TKN values were generally 
higher at the more saline sites. although low 
concentrations were recorded irom Disher Ck Fvap 
Basin. 

DRP levels were relatively low at al) welland sites, 
hut were highest at the more saline sites nf PILCI, 
RAMLIS, CLOL3 and BERBIL. ‘Total phosphorus 
conceniratiims showed & similar trend as DRP and 
TKN levels, with the more saline wetlands generally 
having higher concentrations of phosphorus than the 
reshwater weulands. 

Nitrogen was mostly present as organic forms at all 
wetland sites. ILis difficult t¢ commer on.phesphorus. 
however, as only the dasselved fraction of the total 
reactive phosphorus was measured during this study. 
Despite ths, the DRP results indicate that inorganic 
phosphorus was a significant comributer tn total 
phosphorus for RAMLIS. DISC4, aid PILC2, 

The sites DEVD21, KATS, KATN, DISC?, LCAR 
and WONL were depleted of both oitrogen and 
phosphorus in dissolved inorganic torms. ‘The latter 
three sites also had the lowest TKN and TP 
conccatrations rcconded during the survey. 


Organic carbon 
Concentrations of total organic carbon (TOC) and 
dissulyed organic curbon (DOC) were highest al the 


86 P. M. GOONAN. J. A. BEER, T. B. THOMPSON & P. J. SUTER 


TABLE 4. Chlorophvil concentrations recorded from 17 sites from Chowilla to Mannum in South Australia, (Units in pg!) 


Site* Chlorophylla Chlorophyll b Site No. Chlorophyll a Chlorophyll 
PILE 1 114.8 31.4 BERB 11! 60.0 22.8 
PILE 2 17,2 5.8 RAML 12 32.3 7.0 
CLOL 3 42.1 10.6 RAML 13 9.8 3.9 
LMER 4 27.7 7.6 RAML |4 99.6 37.1 
DISC 6 17.1 4.1 RAML 15 255.8 83.8 
DISC 7 37.2 16,9 LCAR 16 1.2 0.2 
KATS 8 39.1 9.6 DEVD 20 10.8 2.5 
KATS 9 69.3 15.8 DEVD 21 3.5 0.6 
KATN 10 44.1 94 


“Dala not available for LCARI7, LCARIS and WONL. 19. 


TABLE 5, Occurrence of aquatic invertebrate taxa from 20 sites surveyed from Chowilla to Mannum during May-June 1990, 


Taxon Occurrence Total No. of 
(Site No.) Occurrences 
TURBELLARIA 2 l 
GASTROPODA 
Unidentified snail 6 1 
Potomopyrgus niger 16,18 2 
Ferrissia petteradt 4,16,19 3 
Physa acuta 4,9,10,12,15,16.17,18,19,20 10 
Isidorella newcombj 16 1 
BIVALVIA 
Sphaerium tasmanicum 16 1 
OLIGOCHAETA 1,2,4,8,12,13.14.15,16,17,19,21 12 
CRUSTACEA 
OSTRACODA 2,3,4,6,9,10,11.12,13,14,15,16,17,19,20,21 16 
COPEPODA : HARPACTICOIDA 
Attheyella australica 1 ] 
COPEPODA : CYCLOPOIDA 3.6,7,8, 1 1,13,14,17,18,19,20 ‘| 
COPEPODA ; CALANOIDA 2.3,4,6,7,8,9,10,11,16,17,18,19,20,21 15 
AMPHIPODA 
Afrochilionia australis 9.11,12,13,14,15,16,19,21 9 
ISOPODA 
Austroargathona picta 6 ] 
CLADOCERA 
Levdigia australis 3 l 
Ilyocryptus sp. 17,18 2 
Daphnia lumholtzi 2,16 2 
D. carinata l ] 
Daphniopsis pusilla il 1 
Ceriodaphnia sp. 17,19 2 
DECOPODA 
Macrobrachium australiense 20 1 
Paratya australiensis 2,4,6,8.9, 10.13, 14, 16,17,18,19,20,21L 14 
ARACHNIDA 
HYDRACARINA 4,6,13,15,16,17,18 vi 
INSECTA 
EPHEMEROPTERA 
Cleeun flaviatile 19 1 
Tasmanoceenis tillyatdi 9,16 2 
ODONATA 
ischnura heterostricta 4,9,10, 16,19 5 
Austrolestes sp. 21 1 
Juvenile Zygoptera 2 | 
HEMIPTERA 
Anisops sp. 3 | 


Anisops thienemanni 4,8,9,10,14,19,20 7 


SOUTH AUSTRALIAN WETLANDS 


37 


Taxon 


Occurrence 
(Site No.) 


Total No. of 
Occurrences 


Micronecta robusta - M. gracilis 


M. annae 
Agraptocorixa eurvnome 
Hydrometra sp. 
Mesovelia sp. 


COLEOPTERA: HYDRAENIDAE 


Ochthebius sp. 


COLEOPTERA: HYDROPHILIDAE 


Hydrophilid larvac 
Berosus sp. larvae 
Aydrochus sp. 
Helochares australis 


COLEOPTERA: DYTISCIDAE 


Sternupriscus sp. 


LEPIDOPTERA: PYRALIDAE 


Pyralid larvae 


TRICHOPTERA: LEPTOCERIDAE 


Triplectides sp. 
Juvenile leptocerid 


TRICHOPTERA: ECNOMIDAE 


Ecnomus pansus 


TRICHOPTERA: HYDROPTILIDAE 


HAydroptila acinacis 


DIPTERA; CHIRONOMIDAE: TANYPODINAE 


Proctadius sp. 


DIPTERA: CHIRONOMIDAE; CHIRONOMINAE 


Chironomus cloacalis 
C, duplex 
Dicrotendipes sp. 
Chironomus tepperi 
Cladopelma sp. 
Kiefferulus intertinctus 
Polypedilum sp. 

P. nubifer 
Parachironomus sp. 
Cryptochironomus sp. 
Cladotanytarsus sp. 
Tanytarsus, sp.4 

T. barbitarsus 


DIPTERA: CHIRONOMIDAE: ORTHOCLADIINAE 


Corynoneura sp. 
Cricotopus sp. 

C. athitibia 
Limnophyes sp. 
Parametriocnemus sp. 


DIPTERA: CERATOPOGONIDAE 


SR" sp.! 
SR sp.6 
SR sp.8 
SR sp.l6 
SR sp.18 


DIPTERA: PSYCHODIDAE 
DIPTERA: STRATIOMYIDAE 


DIPTERA: TABANIDAE 


DIPTERA: SCIOMYZIDAE 


DIPTERA: EPHYDRIDAE 
DIPTERA: MUSCIDAE 
DIPTERA; CULICIDAE 


DIPTERA: DOLICHOPODIDAE 


3,.4,6.7,10, 11.12.13, 14.15.16, 18,19,20,21 


16.19 
1,3,10,12,13,14, 19.20 
16 

16 


6,15,16,20 


2,4.9,16,21 
18 


16 

16 
1,3,4,6,7.11,12.13.14.19,20 
15,17,18 

12,13 


3,8,9 
2.4,11,14,15 


vB 
‘ 
8,9, 10,14,19,2] 


1, 
3 
4. 
t 
3 
4. 
3, 
4, 10,16,17,19,20,21 
I 

l, 


a 
411,21 


- 


cea 
on 


8.9,10,16,19,21 
9.10,16,19,20,21 
20 


y 


tO, 


1001.12.13 ,15 
1,13, 14,16,19 


4 


See feoae ae fh Us 


ae 


0,11, 13,14,15,20 
3 


—foOooe We ON ek DR DWN 


NVARAVeK Abu 


“SR — refers to voucher specimens in the collection at the State Water Laboratory, Victoria. 


88 PM. GOONAN, J. A. BEER. T. B. THOMPSON & PT SUTER 


more saline wetlands, The fraction of TOC represented 
by DOC varied from 39-50% at KAUS and KAIN 
97% al RAMLI3, 


Pesticides 
No pesticides. were detecicd in any of the water 
sumples (detection limit of 0.02 pgl'). 


Chlorophyll concentrations 

The concentration of chlorophy!l was high at most 
welland sites (Table 4). indicating that significant 
phytoplankton production was occurring during the 
sampling period. Chlorophyll @ concentrations varied 
considerably among wetlands, canging from 1.2 pg" 
at LCARI6 to'255.8 pal! at RAMLIS. Chlorophyll 6 
followed a similar trend. 

Chlorophyll concentrations also varied markedly 
within wetlunds. The most noted difference oceurred 
at Rameo Lagoon where chlorophyll a ranged [rom 
98 pel.’ at the more sheltered western site 
(RAMLI3) to 255.8 peL.! al the exposed, downwind 
site (RAMLIS). Similar wends occurred at Pilby Ck, 
Devon Downs Nth. Kiturupko Evap: Basin, und 
Dishers ‘Ck Evap. Basin, where differences in the 
morpholngy of the wetland, water flow, and the 
dominant wind direction may result in large variations 
in chlorophyll concentrations within wetlands, 


Agnaric jivertebrate composition 

Seventy-eight aquate invertebrate Laxa were recorded 
from the 20 sites (Table 5). Insect taxa predominated 
(69%), and the most diverse component of the fauna 
were dipterans with 32 species, including 19 species 
of cvhironomids, Crustacea contributed 18% and 
Gastropoda 6% of the total taxa recorded. 

Ostracod taxa were the most widespread (16 sites), 
followed by Micronecta robusta-M. gracilis (1S), 
valanoids (13). Pararve austrufiensts (14), oligochactes 
(12), cyclopoids (11), Pracluediny sp. (I) and Plryse acute 
(10). In contrast. 31 taxa were recorded from only one 
SIN. 

The taxonomy for many invertebrate groups. is 
incomplete (Williams 19806; Campbell 1981, Bennison 
ef at. 1989), making it difficult 10 assign some 
specimens below the generic or family level. 
Consequently, not all taxa were identified to species, 
which underestimates the species composition and 
richness of some sites. 

LCARI6 had the highest species mehness with 30 
taxa and DISCT the lowest with 7 taxa, Considerable 
variation occurred within wetlands, paruicularly Lake 
Carlet where WW and 30 taxa were recorded From 
the three sues sampled. Of the wetlands that were 
sunipled from more than one site, Lake Carlet was the 
most diverse with a total of 36 taxa. followed by Ramon 


PILC1 
KATN10 
DEVD20 
WONL19 
LMER4- 
KATSB 
KATS 
DEVD21 
LOAR16 
LOARI7 
LCAR18 
PILG2 
RAML13 
RAML14 
RAML12 
BERB11 
AAML15 
CLOLS 
DISCE 
DISC7 


100 80. 60 40 20 0 


% SIMILARITY 


Fiz, 2, Dendrogram produced by Sorensen's similarity 
evellicients of 20 sites based on the aquatic favertebrate 
data. 


Lagoon (29), Devon Downs Nth (25), Katarapko Evap. 
Basin (23), Pilby Ck (18). and Disher Ck Evap. Basin 
(14). 


Groupings of the sites 

Cluster analysis initially separated the sites inte twa 
main groups that generally correspond (is more suline 
wetlands with TDS concentrations >1000 mgL! and 
less saline wetlands with TDS <10XX) ngl! (Fig. 2). 
Exceptions included the clustering of the saline 
anabranch PILC\ and sites from Katarapko Eyap, Basin 
with the freshwater group, and DISC6 and RAMLI2 
with the more saline wetlands, 

Within the more saline group, sites from within the 
same wellund were more similar to each other (hun 
sites from different wetlands. In the freshwater group, 
however, sites trom the same wetland did nen 
necessarily cluster together. indicating that some 
heterogeneity existed within some wetlands (e.g. Devon 
Dawns Nth). 


Multivariate analyses 
The DECORANA ordinations of the samples are 
iustrated in Fig, 3, and show the centroids for cach 


SOUTH AUSTRALIAN WETLANDS 


300 


AXIS 1 


Fiy, 3. DECORANA ordination of sites based on the aquatic 
invertebrate daw, with TWINSPAN groups stiperimposed. 
{Gigenvalues; Axis 1 = 0.46. Axiy 2 = 0.28, Axes in 
standard deviation units). 


of the 20 sites (i.c, the avérage score for each axis). 
Superimposing the TWINSPAN groups. onto the 
ordination plots results in two groups that also 
correspond ta more saline wetlands with TDS 
concentrations >1000 mgL! and freshwater wetlands 
with TDS <1000 mgL!. This trend was confounded 
by the inclusion of sites from Katarapka Evap. Basin 
in the freshwater group, and DiSC6, RAMLI2 and 
DEVD?20 in the more saline proup. 

The two sites from Pilby Ck were outliers on the 
ordination analysis and tended to “compress” the other 
sites on the second ordination axis. Deletion of these 
sites. from. subsequent analyses did not alter the 
onentalion or spacing of sites appreciably, so the 
original results based on all sites are presented herein. 

The projection of sites onto the first ordination axis 
is shown with their TDS concentrations in Fig. 4a. 
Sites to the left were characterized by having freshwater 
with TDS <1000 mgL! and were connected lo the 
River Murray (Table 6), These included the permanent 
flood plain Jakes and swamps, and two sites from 
regulated wetlands. The freshwater site from Pilby Ck 
also grouped with the other freshwater wetlands despite 
being isolated from the main channel when sampled. 
Sites with TDS. concentrations between 
1000-2999 mgL! formed intermediate proups. 
Katarapko Evap. Basin and Disher Ck Evap. Basin 
(DISC7) were connected to the River Murray through 
their regulating structures when surveyed, while Clover 
Lake was isolated due to its Jovation high on the fload 
plain. Sites to the right were saline with TDS 
>3000 mgL.' and were isolated from the River 


wo 
400 
300478 
(Ae 
r ; 
” 
>» 200 
< 
100 P 
‘ F 
ay 
i) 
0 2000 4000 6000 B000 «625000 
TDS (mgl) 
400 
300 
N 
“ 
> 200 
<< 
100-4) 2 
ay 
10] Sa 
Qo 2000 4000 6000 8000~ 25000 
TDS (mgt) 


Fig. 4, Scattergram of DECORANA ordination from Fig. 3 
showing TDS concentration recorded at each site. 
ta) Axis | vs TDS (b) Axis 2 vs TDS 
The groups enclosed in dotted lines are described in the 
classification. of sites in Tabte 6. (Ordination axes in standard 
deviation units, TDS in mgL‘. LCAR {7 and LCAR 18 
ainitted due to absence of chemical data). 


Mutray. These included the western reach of Pilby Ck 
(PILC!) and Ramco Lagoon in one group, and the 
hyper-saline Berri Lyap. Basin in the most extreme 
group. 

The same general pattern resulted when the points 
from the second ordination axis were plotted against 
their TDS concentrations (Fig. 4b), although PILC2 
split from the other freshwater wetlands. and the two 
intermediate groups merged together. 

Superimposing the nutrient data onto the ordination 
plots revealed a similar. though Jess distinct, gradient 
between wetlands with/without any connection to the 
River Murray, TP showed increasing concentration 
with isolation from the River Murray along the first 
ordination axis, but no interpretable pattern for the 
second axis. The remaining physico-chemical variables 
displayed no obvious pattern along either axes. 


) P.M. GOONAN. J, A, BEER, T. B. THOMPSON & P, J. SUTER 


TABLE. 6. Classifrearicm ef werkurd sites: based vn their TDS concentration and connection ja the River Murray. 


TDS imet!) 0 - 999 1000 — 2999 3000 - 9999 10000 > 
Fresh Waters Saline Highly Saline 

Connected tu 4.6.12.1619.20.21 7,8,9.10 = 

Raver Murray (GROUP A} {GROUP ©) 

Isolated from 2 3 1.13,14.15 i 

River Murray (GROUP Bj (GROUP D) {GROUP EB) (GROUP F) 


The TWINSPAN classification (Fig. 5) describes a 
similar pattern to the ordination results and highlights 
the indicator taxa thal are unique to cach grouping. 
The freshwater group was characterized by {he dipteran 
Cladetanviarsus sp. and the shrimp Puratya 
australiensis. The more saline group was distinguished 
by the presence of the dipterans Prielacius sp.. 
Ephydridae and Culicidae- 


Discussion 


Water chemistry 

Like most inland waterbodies in Australia, all 
wetlands included in the present study were dominated 
by sodium and chloride (Williams & Wan 1972). The 
differences in ionic concentration and dominance 
between. wetlands were largely the result of dilution 
and concentration, The freshwaler group were 
permanent waterbodies connecicd to the mainstream, 
where water level fluctuations are less extreme than 
in the more saline group of isolated wetlands. The 
regulated wetlands, ephemeral swamp, and saline reach 
of the Pilby Ck anabranch had higher salinities due 


Cladotanytarsus sp. 
Paratya austelionsis 


to the effect of evapoconcentration. Seepage of saline 
groundwater and the inflow of saline irrigation water 
also added to the high levels of dissolved salts in the 
evaporation basins and Ramca Lagoon (Unpubl. 
E. & W.S. Dept records). Recemt and proposed 
changes in the management of these wetlands by the 
use of out of the flood plain evaporation basins (e-g. 
Noora, Stockyard Plains) and groundwater interception 
schemes. should lead to a reduction in salinity of these 
wetlands in the long-term. We should note, however, 
that meun salinity levels would probably need to be 
reduced to at most 4000 mgL" before significant 
changes in the biota of these wetlands would be evident 
(see Centre for Steam Ecology 1989* for references). 

Comparison of TP and TKN concentrations recorded 
inthis study (Table 2) with Wetzel's (1975) classification 
of lake productivity (after Vollenweider 1968), reveal 
that the 10 wetlands were eutrophic or hyper-eutrophic 
with respect to TP, and meso-eutrophi¢ or eutrophic 
with respect to TKN, Levels of DRP and 


{Centre for Steam Ecology (1989) “Biological Effects of 
Saline Discharges t0 Streams and Wetlands,” (Chisholm Inst. 
Tech., Unpub!. Report for Salinity Bureau, Vict.) 


Procladius sp, 
Ephydridaa 
Cullcidae 


Hyocrypls sp, 


Turbetleria 


Polamopygus niger 


Hydracarina 


Ferrissia pettardi 


| 


LGARV PLC? LMERA KATSB WONLIS 
LOARIS KATS® 

KATNID 

DEVD24 


LCARIG 


Altheyelta australis 


} 


Leyd/g/a sp. 


Afrachiltonia australe 


Physa acuta 


bisce 
bpISc? 


BERB1 
RAMLI3 
DEVO20 AAMLI4 


RAMLI2 
RAMLI5 


CLOL3 PILG1 


Fig. 5. TWINSPAN classification of sites. based on the aquatic iwertebrate data. Indicator specics names ure included with 


each dichotomy, 


SOUTH AVISTRALIAN WETLANUS oI 


NO,-N. however, were venerally low, suggesting that 
MOST AP [he hulreals Were i particulate furs thal muy 
be Unavailable ta phytoplankton (Smith 982: Geddes 
I9R4a), 


Nutifent concentrations of she wetlands reflec those 
in the lower River Murray. Values of TP, DRE, TKN, 
And NO,-N were withit the ranges reported from 
Lock 5 ty Murray Bridge (MucKay era/ (988) with 
some excepoons, These included the higher TP 
concentration from Clover Lake and the higher TKN 
cAncentrations from Bern Evap, Basin, Clover lake, 
Ramo Lagoot and Katarapko Eyap, Basin. These ure 
shallow and/or reewluled wetlands subject co 
comsiderable evapuration, resulting in high 
conventrations Gf nutrients and (lissalved Salts by 

upocancentration, 


Conpansen ol netnent levels with other werlunds 
fram the River Mucray ts difficult 4s few studves have 
been published im the chenusiry of these waters. Shicl 
{I9RU) reported the nutrient concentrations fram three 
billaborgs hear Wodonga during 1975-77, and found 
thal nitrile varied from 2-685 me Noor and 
phosphate from. U-624 mg Pov’. Nutrient. dewels 
trom Lake Alexandrina (Geddes 484i, Lake Hume 
dnd Lake Mulwala (Walker & Hillman 1977: Brymer 
1982), Murrumbidgil Swamp and Lake Merrimaject 
(Brigas er al. 1985) were ull within Shiels ranges, 
Larze Muctuations: in nutrient concentrations were 
recorded from cach wetland. In the present study, 
nitrile and dissolyed phosphate concentrations were 
hw compared co Shyel’s 1980) values. Law Jevels of 
inurganie nitrogen relative 10 the high TKN 
conventritivins. indicate that N was cither present in 
We sedinienis of hid been assimilaced hy 
phytoplankton. The high chlorophyll concentrations 
(Table 4) Hom nist sites support Ihe latter suggestion 
Based on Wetrels (1975) chlorophyll a categories. 
LOARI®A. DEVD21, RAMLI3 and DEVD20 were 
mesco-culrophic, whereus the other sites were eutrophic. 
Although meaningful critical concentrations of 
nutrients have not been. defined for Australian waters 
(Wood 1975, Cullen 1986), the Jlood pluin wetlands 
included in the present survey were clearly enriched 
m both N jind PP Future work will deterinine whether 
the high levels of mutiients and algal biomass ire 
sustained, as this could result ip the alteration af 
phytoplunkton comoiunities i favour nuisance species 
of <yanobacteria (Walker & Hillman 77), 


Aqtitie invertehrnes 

The aquatic jwertebrate fauna was diverse (lable 
3) cunsidering the sitall Number ol samples collected 
wu! (hat sampling occurred during the cool, wee 
manths of May-June. At least 78 taxa wene recondoul 
fromthe JO wetlands, with insects und crustaceans 
Sumitating the invertebrate Communities it every site. 


‘The majority of insects were dipterans (32 taxa). 
hemipterans 17) and eulewpienins 16), Sites From the 
permanent Treshwaler lakes and swamps (LCARIG, 
LMER4 and WONLI9) had the most (isa, while 4 
permanent regulated wetland (DISC7) had the least. 

The itnpublished database compiled by Thompson 
(1986) contains remarkably few records mf invertebrates 
and aquatic mucrophyles from the weilands included 
i his study. This was prohably due. in part. to the 
high flows af very tuchid waner from ihe Darling Rivet 
inte the River Murnay at the end ot 1983 (MacKay ef 
af. 1988), resulting in most wetlands being Wirbidl when 
sumpled by Thompson in 1983-4. Apart from noting 
ostrcods Irom DISC? no new data could be derived 
from this database. 

Lhoyd ev el. (1984)! collected 71 aquatic invertebrate 
tuxa during a 12 noni study of the fluctuations in the 
aquatic invertebrate commiunities and water chempitry 
of theee WeUlands, including Berm and Disher Ck Evap. 
Basins. Comparison of results from the same time ol 
the year show that the fauna and water chemisity have 
nut chinyed appreciably ar BERBI. while the urtiticw! 
manipulation of water levels in DISCE6 led to # lewer 
salinity. and amore diverse fauna in the present study. 
A total of 28 taxa were found ar BERBIU by. Lioyd e7 
ai, (984)! with 12 Juxa being recorded during May 
IS84, The same faunal assemblage was present during 
May 1990, with the addition of Afrochittiania australis, 
Daphyiopsis pusilla and Micronecta rabysta-M- 
geaciliy. OV the 35 laxa recorded from DISC6 in the 
eurhier work, only four were found during May WS4. 
In May 198) 1) species were collected, dominated by 
crustaceuns and dipterans, Future work will determine 
whether the seasonal trends deseriheal by the earlier 
study are maintwined. This will provide.a uscful means 
of predicting how conservative are the diflerent 
parameters thal were mensured in (hese evaporation 
basins, and establish a database upun which any 
changes tn the management of these wetlands can be 
compared, 

Lloyd & Boulton (1990) recorded 96 niseroinver- 
lebrile taxa during # recent short-term survey of 13 
wetlands from the Chowilla flood plain. Wetlands were 
sumpled as river levels fell in October G88 As in the 
present study, Most Wika Were insects. with drpterans 
14] taxa) dominating the ana. Few crustaceans were 
collected. partly becuse a larger meshed dip net was 
used and did noc sample the microvrusiaecans, The 
mayor diffcrence in the fauna between the tw studies 
was the large number of beetles (22 taxa) recorded by 
Lloyd & Boulton (1990), Dytiscids and bydrophitids. 
are fooxt Commonly collected during spring-sumnmer 
Frond most inland waterbudies (Matthews 180, 1982), 
with shallow temporary. wetlands often having a variety 
of species (Lloyd & Boulton 90; pers, dbsy,), The 
(ining of ow survey may. account for the fewer species 
of beetles recess, 


9) bh M. GQONAN, | A REFER, | 


Crnrparison of the faunal communities al the 20 sites 
using DECORANA ordination (Fig, 4 and Tuble 6) 
illustrated the Unpertinge- af connection ta the River 
Murray on both the water chemistry sad uquatic 
invertebrate assemblapes, Wellanils/stes wilh clirect 
Connection to the River Murray were charucteriaed by 
Jow salinities (TDS concentration <JOUW meL'1. 
veherally low nutrient concentrations, and the presence 
ol the dipienan Cleeeremvrasis Sp. und the shrimp 
Purarve custrationsix, Wettands/sitey that were isolated 
from) the mut channel formed a second group. 
characterized by higher swlinities (TDS. comeentrtion 
> 1000 mgt!) fish nutrient congentrations, unl the 
presence of dipterun larvae such as Procladtiny sp. 
Fphydridae and Culicidae, 

The sites hnsullocuies! by the anulyses deserve special 
Mention. Hydrological muimpulations of three regulated! 
wetlands: prior to jhe survey, conlounded the salinivy 
gradient. Kutarupke Evap. Basin, Disher Ck Teun, 
Basin, and the inlet site at Rameo Liyoun were 
recewing wales trom the River Murray when sampled, 
as their Tewulafing structures had been opened twa, 
seven und 10 days respectively, prior ta sampling 
Cnpabl. 1. & W.S. Dept records). Salinity ceadings 
trom these wetlinds indicute that some mixing and 
tlilulinn hil occurred in Kutarapko Evap, Basin and 
Dishee Ck Evup. Basin (Unpubl, 2. & Wis. Dept 
recurdst. while Intle to no flushing had oecurred 
beyortd the inlet/outlet site it Ramco Lagoon (Table 
2), This wppeurs Wo have altered the fauna of Katurapko 
Evap. Basin to resemble a more restiwater assemblage 
ol invertebrates, Sites trom Disher Ck Evan, Basin und 
Romeo Lazoon, hiwever, retained invertebrate 
asscrnbliges typleghot the more saline sites/werlands. 

the high nutrient conveninuiiins tecirded ul 
DEV O20 may have contributed to gn dssembhige of 
Invertebrates typical of saline conditions, lespate bay iy 
a TDS concentaition of only 360 mgl ! ‘This site was 
heavily grazed by sheep, with Lhe stock having diccct 
wecess to te waterbody. Biological decomposition of 
the mitnure in the water could have produced the high 
NH, concentration, whieh would) then oxidise to 


K. LHUMPSUN & PJ. SUTER 


NO -N by bacterial uction (Bayly & Williams 19731. 

Sites (rom Pilby Ck tended to forni outlier positions 
in the daw analyses. emphasizing a difference in the 
fauna fram dhis anabranch compared with the other 
wethinds, Pilby Ck is a Small anabranch in the Chowilla 
region, characterized by narrow banks with River Red 
Goms. extending over the widler. As none af the other 
weullunds resembled this macrohabitat, the 
Uistinetiveness of this Wetlind within the analyses wars 
not remarkable. The causeway across Pilby Ck has 
clearly reduced the water quality of the western reach 
wo favour organisms adapted to suline, organically 
enriched conditions. Placement of a culvert with it 
regulator under the causeway would provide a suinple 
incuns of manipulating water levels to reduce the 
salinity and mulrient concentrations of the western 
reseh 

These are the preliminary results of an ongoing study 
of the water chepnisery and biota of Hood plain wetlands 
in South Australia. ‘Uhey provide an initial database 


and demonstrate the influence of the River Murray oa 


the waler chemistry und aquatic invertebrates of the 
wetlands sampled, Future work will describe the 
influence of season, flow and regulation on the 
limnology of some of these wetlands, and provide 
wuilelines for thé management of wetlvnds throughout 
the Murrav-Darling Aue pliin. 


Acknowledginents 


This work was funded by the Munay-Darling Basin 
Ministerial Council theaugh the Natural Resources 
Manigement Strategy. Chemical analyses were 
conducred by various stalf trem the State Water 
Laboratory, E. de WS, Dept. Mrs R. Cowdrey and 
Ms S. Boyd prepared the tables, and personnel from 
Dralting Services, E..& W.S. Depr, drew the figures. 
Mr G, Wood, Dr D. Steftensen, Mr P. Christy and tuxy 
anonymous referees provided valuable comments on 
an earlier dralt of the munuscript. 


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od 


DISPERSED CUTICULAR FLORAS OF SOUTH AUSTRALIAN 
TERITARY COALFIELDS, PART 2: LOCHIEL 


BY A. I. ROWETT* 


Summary 


Dispersed cuticles were recovered from two lithotypes (Facies Ia, Ila) within the G seam of the 
Kooliata Coal Zone of the Lochiel Coalfield. The floras are distinct. The younger lithotype (IIa) 
contains a monospecific flora, represented by a robust, coriaceous non-stomatiferous cuticle 
whereas the older lithotype (la) contains thirty-seven cuticle types. The major contributors are 
Agathis (Araucariaceae) which dominates the flora and Podocarpaceae, Proteaceae and Myrtaceae. 
KEY WORDS: Palaeobotany, Tertiary, Eocene, dispersed cuticles, Lochiel, South Australia. 


Hansaenevis af te Raval saci af S$. Aust (992), W604), 95-100 


DISPERSED CUTICULAR FLORAS OF SOUTH AUSTRALIAN 
TERTIARY COALFIELDS, PART 2; LOCHTET 


by A. lL. RoWeT1= 


Summary} 


Rowrtr, A. 1 (1992) The dispersed cuticular Moras of South Austrian Tertiary Coalfields, Part 2, Lachiel. 
Truss. R. Soe, & Aust, W6(3), 95-107, 30 November, 1992. 

Dispersed cuticles were recovered from two lithotypes (Facies fa, Hn) within the G secant of the Kool Coal 
Zone of the Lochiel Cosifield. The floras are distinct. The younper Jithotype (Ifa) contamns # monospecific Mora, 
represented by a robust, coriaceous non-stomatiferous cuticle whereas the older lithotype (la) contains thirty- 
yéven culicle types, The mayor contributors ans daddys (Arwucarigceac) whieh dominates the flora ant 


Pouncarpaceae, Proteaceae and Myrtacenc. 


Key Woros Paulneobotany, ‘Tertiary, Hovene, dispersed cuticles, Lochiel, South Australia, 


Introduction 


This ts the second paper on the dispersed cuticular 
floras of Seuth Australian Tertiary coalfields und 
follows the furmat used In Rowen (1991), 

The Lochicl coalfield, located [30 km north of 
Adelaide (Fig. 1), at the head of the Gulf St Vingent 
(33°56', 138°l0"), is one of five separate lignite deposits 
Within the Nonhem St Vincent Basin, i.e, the Beaufort. 
Bowmans, Clinton and Whitwarta deposits. [i was first 
discuvered in 1982 by peologists of the Electricity Trust 
of South Australia. 

The Northern St Vincent Basin is characterised by 
north-south Lending faults, considered responsible for 
controlling Tertiary sedimentation (South Australian 
Department of Mines and Energy 1987), The 
Ardrossan and Whitwarta Faults (Fig. 1) delineate the 
Lochiel deposip in the west and east. 


The Lochiel coal-bearing sediments are members of 


the Clinton Formation which has been subdivided into 
three units in the northernmost part of jhe basin, trom 
the base. the Bumbonga Sands, Condowie Silt and 
Kooliata Coal Zone (ETSA 1988'). Small lignite 
lenses occur in the Bumbunga Sands but are of little 
economic importance, Three mayor lynite seams (F, 
G and H seants) occur in the Kootiata Coal Anne 
{Kremor & Springbett 1992). which average 
thicknesses of 2. 6&5 and 2.5 metres respectively, 
Carbonaceous lacustrine silt, sand and clay of the 
Conduwie Silt separate the Bumbunya lignites from 
hose of the Kogliata Coal Zone, Uncodsolidated 
Oligucene clay, silt and water-saturated sand of the 
Warrindi and Tarefla Sifts, ranging in thickness from 
20-70 m, uncontormably overlie the lignite- 
Palynological evidence from these lignites suggests 
a Late Kocene-Oligocene age (Harris 1965, 1971; Alley 


* Dept of Botany, University of Adelaide, G, P.O. Box 498. 
Adeluide, S. Aust. SO0L 

' Figure on pave Sof ETSA 1988, Loohiel reporr shows E-W 
sindigraphic cross-section of eposit, 


& Lindsay 1991 pers. comm.). The pulynoflora from 
the Bumbunga lignite are the time equivalents of the 
Middle and Upper Nothufigidites asperus Zones of 
Stover & Partridge (1973, 1983), which are Late Eocene 
to Early Oligocene in uge (Kremor & Springbett 1992). 
The Kooliata lignite is somewhat younger, probably 
Early Oligocene, and the palynoftoras are time 
equivalent to the Lower Proteacidites teberculatus 
Zone (Stover & Partridge 1973. 1983), 


Materials and Methods 


Lignite samples were recovered from a trial pit (Fig, 
1) excavated during the initial tesource assessment in 
1987, Only the lignite scams of the Kooliata Coal Zone 
were exposed inthe pit (Springben pers. comm.) but 
limited access (I hour) to University of Adelaide, 
Botany Department collectors prevented 
comprehensive sampling, Sampling was therefore 
undertakeo of those lignite seams noted to contain 
considerable amounts of heavily carbonised dispersed 
cuticle and wood fragments. The two samples selected 
jor this study were taken from Facies Ja and Ma of the 
central G seam (Springbett pers. comm; Fig. 2). 

Dispersed cuticles of the Lochiel deposit were 
processed and analysed using techniques outlined by 
Christophe er af, (1987), Rowett & Christophe} (1990) 
and Rowell (1991), 


Dispersed Cuticle Flora 


Analysis of samples revealed a diverse cuticular flora 
of 38 parataxa which are unevenly represented in the 
Iwo facies, Le. 1 (Facies Wa), 38 (Facies Ta), 

"The cuticle flora of the upper highly gelilied lignite 
(lla) consists solely of the parataxon No, AW 007, 
which iy very distinctive despite the absence of 
stomates, The robust, thick, cotiacemis cuticle, sinuous 


96 A. I. ROWETT 


NORTHERN 
ST VINCENT 


SNOWTOWN 
BASIN 


WHITWARTA FAULT 


Cummulative lignite 
thickness Cin metres) 


5-10 
10-15 
15-25 


Pre—Tertiary 
Outcrop 


(salt) FAULT 


“Lochiel 
Deposit 


WAKEFIELD] @ 


_ochiel 
acedan 


Gaiden Grove 


> Kingston 


Gulf St Vincent 


Anglesea 


kilometres 


Yatlourn 
ry 


LOCHIEL DISPERSED CUTICLES Y7 


epidermal cells (types 3 -4 of Wilkinson 1979) and 
distinctive trichome clusters distinguish this cuticle 
from all stomatiterous parataxa (Fig. 3). This 
monospecific Mors inay provide a useful stratigraphic 
marker in the correlation of the Lochiel ignites. 
Parataxon No. AW 007 ts also present in the underlying 
facies (Facies lit). 

The lower lignite facies. with an abundanee af wood 
ringing from twigs to large diameter logs, contuns a 


Facies lla dispersed cuticle flora characterised by a large 


Approximate Araucariiceae component (29.0%) as well as 
sampling significant Podocarpuceae (15.5%), Myrtaceae (15.2%) 
sites und) Proteaceae (12.9%) vomponents (Table 1). 


Unknown cuticle types represent a lurge percentage 
‘ of the Loehiel Mora (33.5%). Casuarinaceae. 
Facies la Eldevearpaceae, Lauraceae and Zamiaceae are poorly 
represented (Q5%, U6%, O4% and 0.2% 

respectively), 

The Araucuriaceae is represented by a single 
parataxon No, LC Q03 (Figs 4-5) with a suggested 
affinity tu the modern Avarhis. The fragmentary 
cuticles, & commen feature OF this locality, makes a 
niore definite wentification impossible, 


KOOLIATA COAL ZONE 
G Seam 


The Podocurpaceve is represented by three parataxa 
ob which No, LC 002 (99%) is most abundant (Pigs 
6-7), Stomates have a yvariuble appearance which mukes 
W difficult to assign the cuticle (ype to aw known 
padooarp genus. The cireulir appearance of numerous 
slomiates suggests a possible affinity to Falceifaliant 
(Greenwood 1987). Parataxa Nos, LC O11 und 
ABP OOF are also common, representing 4.6% and 
10% respectively, Parataxon No, LC O15 is assigned 
Pip. 2. A suinphificd Thological column of G seam ofthe to Daervearpus on the basis of smooth-walled 

Kooliata Coal Zone showing the seven lenses identified by epidermal cells and arrangement and eureular 


Springbett (LI-L7) and the approximate location af the : ; : - - 

eantpicn uscd inthis iivestigaticin within Facies iiand Ite. thickening of subsidiary cells (Pigs 8-9). Parataxon No. 
Facies Ta is an earthy textured coal contaiting more thin ABP OOL (Figs 10-11) is identified as the cuticle of 
40" relanvely ungelified woody nateriat while Facies Ikt Podocarpus plaryphyllum (Greenwood 1987), 

isa darker Hibrous, lignite consisting of gelitied twigs, The . ; . 

seam is approximately 6.3 m thick and ata depth of 30 The Myrtaceae comprises two cuticle types Nos, 


metres, LC OO) (12.0%) and LE OW (3.2%). Both appear 


TABLE |, The cut le frequenetes (Sof estant plant fianilies represented in the Wwe facies of the central lignite sean of 
He Lochiel ileposit, Families represented ane Podocarpaccue (POD), Araucariaceae (ARAUIC), Myrtaceae (MYRT), 
Blacoearpaccae (ELAEO), Proteaceae (PROT), Laurieceie (LAUR), Casuarinaccue (CAS), Zanmiaceuc (4AM), The OTHERS 
category represents all other cudele paratasa whose moderna family affinities are unknown, 

eee 


LOCALITY POD ARAUC MYRE  BLAEQ PROT LAUR CAS ZAM OTHERS 
a 


Loebiel (bacies Lay 15.8 29.0 15.2 0.6 129 (4 (5 02 257 
Loehiel (Paces Ta) - — - ~ = Wo) 


SSeS 


oA 3 ————————————————————— 

Fiz), Map showing extent aod thickness of fizmte inthe Lochiel deposit and major structural features. Inset: Map of eastern 
Australia showing the Joeation oF the Sedan, Lachiel and Kingston coal localities relative to the Tertiary megalossil localities 
of Golden Cirove (Eocene), Anglesea (Eocene) and Yallourn (Oligocene) 


A. |, ROWETT 


OM 


LOCTIEL DISPERSE CL LICH bS Ww 


morphologically similar bit can be distinguished on 
epaleniiet cell shape, stomalil urrimgement und 
epidermal cell, stomme and oil gland lid cell 
dimensions. Parataxon Noo LC OOL (Figs 12-13) fs 
vhuraviensed by rounded to undulate epidermal cells, 
AT ANTSUEV LIE {0 sTHUPOCVNG Stoniatal atramgement of 
3-5 subsidiury cells and hd cells (hat may be constricted 
allie sinus, Parataxon No LC OW onthe other hand. 

ischuracterised by undulacte to sinuous epidermal cells. 
a feveloeytie Io) stuuroeytic Stonmatal arrangement of 
3-6 durb-Stuining subsidiary vells and large lid cells 
Hor constricted atthe siaus (Rowe 191), None of these 
purdiixd ure elsely comparable with cuticles of the 
species of Myvrecplvtivn described by Christophel 
& Lys (YK) from Anglesea. 


The Proteaceae is a diverse group of ging cuticle 
types the majorly oecaring in Jow frequencies 
(20.1%). The most abundant is Banksicueplivihun all. 
BL lueve (3.1% 7. the sume cuticle type Chit doniinated 
the Late Bocene lignites of the Sedan deposit (Rowet! 
1891). Two other cominon culicle (ypes are parataxa 
Nos, LE 004 (3.0%) und LC 012 (2.8%), both assigned 
to Barksievepiolian with the former cuticle type 
idemilied as Banksiewephyilun aft. RB fastigarun 
Cookson & Duigan (Pigs 14-15), The cuticle can be 
identified by the well-defined iregles, the relatively 
low frequency of stomatiarcole. slightly raised 
slomiuti, a pale of pourly defined sabsihiary cells 
surrounded by 3-4 darkly sutined epidermal cells and 
long unicellular tiehomes with a por) Buse. The 
culicle of B fastiearuny was described by Cookson & 


Duigan (1950) from the Oligocene brown voals of 


wlll five other Species of 
Banksieaepivllum, ic RB aneustin, B acwninarint, 
B lueve, B. sfintisla am! A fastigaien, The 
distribution of these species has been discussed by 
Rowert (991) 


Purataxon No LO O12 may have an aifinity to B 
Obovant Cookson & Duigan (begs 16-17). 
Deseripnons ol A abevatin and & fastigatun are 
similar. the distinguishing feature appears to be position 
Of the stomiute. ic. slighUy raised in the former and 
slightly sunken inthe luner (Cookson & Duigan 1950). 
Another-cuticle type, parataxon Noa. LC 013 { Fig, 18), 
is also assigned lo Barkviewephytlune. Wis possible the 
Puraluxon muy represent another fragment of 
Banksteaemiytiont aa LB, ebovetune. 


Yallourn together 


Casuarimacene. Elacoeurpaceae, Lauraceae und 
Zamidceae are of minor imporkince in the Tower 
Lochiel flora, Casuuimuceae is represented by 
Gynmosten (paratixon No PM 007), which alse 
occurs in the lower lignile seam of the Sedan deposi 
(Rowerl 1991), Guwanrastena cuticles at Lochiel indicate 
that the genus was more widespread during the Late 
Bovene-Early Oligocene, not restricted to south-eastern 
Australia as would appear was the case during the 
Middle Eocene (Christophe! 1980; Rowell & 
Christophel 1990), 

Zamiaceae is represented by paratixen Noo ABD 002 
identified as Plerostontia WP zanilajiedes. Hil] (ORO) 
(Figs 120), This tentative association ty prompled by 
the rarity and poor preservation of the Lochicl 
specimens, 

Elaeocarpaceae and Lauraceae are repre pares ny 
cuticle types ABD 005 and AG O10 (Migs 21-22 a 
Figs 23-24 respectively). 


Dispersed Cuticle Descriptions 


As In Roweit (991) only paratixa of Uorystic. 
striligraphie and bixonomic significance ure described 
Some parataxa have been previously idendtied by the 
author from other Eavene loealities (Rowe & 
Christophe! 1990; Rowett 1991), These and many more 
are included in the National Energy Researetr 
Development und Demonstration Counei) relerence 
cuticle catalogue nf Australian Eovene cuticles types 
(Rowett in prep’) ALL paratzxon numbers are 
preceded by an abbreviation of the type locality 
Terminology used i these desenptions follows that 
oF Rowett (1991) which was derived front Stace (1965). 
Dilcher (1974) and Wilkinson (1979), 


Cuticle Paratixon No, AW 007 
FIG 3 


Thick, coriaceous cuucle. Non-stomaiterousy surtice 
only, Adaxial epidermal cells undulate (type 3+. 


* Austrilian Eocene med Cuncle Catalogue. Appenclis 
Aw ROWETT, ~ (990). National Enemy Research 
Development nd Derssratreion Cuuncil, Pinal Report. 
No. 3, 


Figs 3-8, 3, Pacmeu So. AW U7 Non-stomatiterous cuticle of unknown affinity. Note heavy cuticular thiekenmnpe around 


the trichonte base. Seale | em = 


obliquely Unented stolreiies arranged in cows dnd minded epidermal cells, Seale Lem = 


7O ny 4, Parataxon No. LC O03. wll, dgarhiy, Anueariacede. Shows predaminantly 


35 pm. 5, Paraiavon No LC 0048 


uth, Agathe, Afiuchniaeete Catieular thickening around stomatal pore could be interpreted as 9 Flori cine Carrowedl) 


Seale bom = 
hands consisting of lomeuniseriate rows, Seale | em = 


27 pi A. Paralaxon No LC 002 Ailrasifivine, Podoe: arpyceac Shows arrangement of stomutes in broad 
SU pm, 7, Parataxon Ne, LC 002, VFalcatiteliam, Podocarpaceae 


Shows slornsttal surranige merit wna number of cneular stomates, which may suggest an affinity tn tits genus. Seale Pern 


«40pm. &, 


§. Pantuxun No, LC OS, Bucrrearus. Podocarpaceac, Shows arrangement of stomates. Scale | cm = SO am, 


A. 1, ROWETT 


100 


LOCHILL DISPERSE CL TICLES hy 


Wilkinsea, 1979). 48 -80nm long, 40-68pn1 wide 
Anticlinal wall thin, smooth, Perichnalwall irregularly 
Ihickened. granulate fe stride, ‘Vrichome bases 
multicellukr?, common, saoilorn disiriburion, vive prse 
fo Many singke cell tichomes, appear as tuts, 
‘Lrichomes of vanable tenth. radiate out from centre 
OL Wise. 

Aff Caknown 


‘uutixon No. LO O03 
PIGS 4-5 


Cubicle 


Stomatiterous surfiee only, Epidermal cells angular 
lo rounded, arranged in longitudinal tows. 24—44y0n 
Jong, 20-36unr wide. Anticlinal wall ireeulurly 
tiekencd. Smoot te beaded, Perivlinal wall granulate 
Stonmalw loosely arranged in uniseriate rows, 20-32. 
long, 8-lojan wide. Stomutal orientation ranges trom 
parallel oblique i perpendicular to long axis ol 
epidermal cell cows. Stomuatil arrangement teuracy lic 
ta cyeloeyae, Guard cells sunken. Subsidiary celly 4-5, 
literal cells generally larger that polar cells. Anticlinal 
wall smooth to headed, Periclinal wall granulate. Guard 
cell/subsidiary cell wall heavily cutinized. slightly 
raised, Plorin ring evident. 

Affinities The parataxon is assigned to the 
Araicaraeeas, The predonmantly oblique orientation 
gt stomata, rounded epidermal cells and uniseriante 
stoma) rows Suggest an alfinity with) Agarhis 
(Cookson & Dureun l9Sh) Stuckey & Taylor 198); 
Bizwood & HV 1985, Hil] & Bigwood 1987). 


Curicle Paratwxon No, LC 002 
FIGS 6-7 


Stomititepods surface only, Epidermal cells angular, 
urrinped in lvogiudinal rows orented purallel to-vein 
direction, 32-60am long, 828m wide. Anuclinal wall 
inrepulitly thickened, smvoth be beuded. Periclinal 
wall irregulirly thickened, smooth to granulate to 
pitted, Sromata avruiged in uniseriite rows, muy be 
discoghnuous, onented parallel lovein direction. cows 
gmiuped in broad binds. 2-4 epidermal cells apart, 
Stomatal arningement tetruvyuic. Guard cells sunken 
poral thickening, Outer stomatal ledge, prominent, 


brid. Florin rig present, Subsidiary cells four (rarely 
five), polur cells wedge-shaped to raunded 
foccusionily clongatel rarely Shared. lateral cells 
creseem-shaped. Antichinul wall irreeularly thickened , 
smooth ta beaded, thickening extends aloe radial Walls 
OF polar cells. Perichiial wall iprewulurly thickened, 
Smooth to granule, 

Affi The puratwxon ts assigned to the 
Podocurpueede with a possible affinity ty faleatifalinay 
but may equally beloqg fo one of the muy extinel 
Austithan Tertuiry generis, 


Cuncle Paritaxon No. LC WS 
FIGS &-9 


Stonmuatiferous surface only. Epidermal cells generally 
rectangular, Cells 25-125 juni long. 125-27 5m wide. 
Anuclinal wall undulate. smooth or showing some 
buttressed thickening, Periclinal wall irregularly 
thickened , granulate 10 striate. Stonata i broud bands, 
onented parallel to the long asis of the leal, Stomuatal 
bunds contain on. average Six unisenate rows of 
stomata. Stomata generally separated by more than w 
single epidermal cell. Stomatal arrangement 
parate(rucytic, Guard cells slightly sunken. Subsidiary 
cells lour (rarely five), Polar cells much seller than 
lateral cells. Lateral cells arched, Anticlinal wall 
rounded. smooth to beaded thickening. Uccasignilly 
buttressed, Periclinal wall very thick. darker stating. 
Thickening extends Over guard cells. Stomatal ledge 
evident. 
ffir: The paratison is assigned to Duorveurpus 
(Podocurpaveac). Distinguishing features of this cuticle 
muinly ure related. to subsidiary cells, ic, shared polar 
subsidiary cells, thieker cuticle over the polar 
subsidiary cells und literal subsidiary cells that 
surround the base of the polar subsidiary cells. 


Cuticle Purataxon No, ABP QO! 
FIGS 10-11 


Stomatilerods sutfiee only. Rpidermal cells rectangular 
(oO rsodiinctric between stomatal rows, becoming 
elongate over veins, oriented parallel to long axis ot 
the leaf, some groups of cells obliquely oriented to 


Fiys OL 0. Paratixon No, LC 01S. Daervcurpis. Padocarpaccue, Shows elongate polar subsidiary cells and inercased thickening 
ofthe lateral subsidiary cells Within the stomatal aeranuement. Seale | om = 37 jan, 10, Paraiso No, ABP OOL Prete UPN 
ANP plaivpln tian, Podocarpacese. Shows arringement of stormites. in short unisertute rows. Nowe headed thickening 
oo gaiclio) walls oF epidermal cells and the presence pF striauons on polar subsidiary ects Seale am = SS jan th, 

‘araluxon No, ABP OO], Poderarniy at PF platphyitan Podovarpuceae, Shows stomici drrangement, Note prominent 
Hono riog surroundine stomate (arrowed). Seale bem = 35 am. 12. Paratuxvn No. LC u0l Mynaceae Shows siiuous 
cpdvrnial cells aad rindom armiigement of stumites. Note promiment guard cells und associated ‘T-shaped thickening 
alihe poles Seale }em = 40 jan. 13, Pariagen Ne. LO OOL Mynaceae. Shows ain oi} gland Jid cell (ed. a dingnestic 
euncular Levliite of the lainily Seale ben = 40 aim. 4, Paratigon Nu. LC 004. Runkyivarplivitien otf. Bo fustiguran 
Proteacede Shows somal clusters of sunken stomtes between prominent veins covered with numerous dark-saining hair 
huses, Sele bem = 80 jan, 


ROW ETT 


I, 


A. 


. 


1. 


LOCTUEL DISPERSI.D CLTICLES 3 


lon axis af the leat, 40-96 jm long, 12-36 pi wide. 
Anuctinal wall irregularly thickened. beaded lo 
bultressed, Periclinal wall irregularly thickened, 
strongly striate to reticulate. Stomata in’ distinet 


linserute rows, oriented parallel to the long axis of 


the leat, 92-144 pan long, 24-32 pm wide, Rows 2-6 
cells apart, Stofatel asrangement paniletmeytic. 
Subsidiary cells four, lateral subsichary cells renitornm 
Anliclinal wall irregularly thickened. smooth to 
buttressed. Periclinal wall treeguliarly thickened, 
smoot te retwulite, Polar subsidiary cells square to 
revlanguhir, Anticlival wall irregularly thickened, 
spat to butlressed, Periclinul wall irregularly 
thickened, reticulate, Polar subsidiary cells muy be 
shated. Florin riag prominent. 

Affinity, The pardiaten is. identical jo cuticle 
deseribed by Greenwood (1987) lor the Angleses fossil 
species Pudocarpuy playphyllam (Podocarpaceae). 
The distinetive beading of the anticlinal walls and 
stnavon ol the periclinal walls of the epidermal cells 
are diaynostic of the species. Parataxon No. ABP 002 
is (heretore identified as Podocerpuy uff. P 
platypinttin, 


Cuticle Purataxon No. TC OOF 
FIGS (2-13 


Stomatiterous surface only, Abaxial epidermal cells 
counded to undubale (3). bevoming elongate over yeins, 
12-36 Jong, §-20a wide. Anticlinal wall thin, 
smooth, Periclinal wall thin, smooth. Stomata 
randuinly oriented, uniform distribution, 20-28.m 
long, 20-24nm wide. S.1, 11.4, Stomatal arrangement 
anisoeyuc lo stiuroyylic. Guard cells not sunken, poral 
thickening. T-shaped thickening, polar rods present. 
Ovter stomatal ledge prominent. narrow, Subsidiary 
cells 4-5. Antichinal wall thin, smooth. Periclinal wall 
thin, smooth, Hydathodes cure, over veins, dimensions 
32am long. 24nm wide. Oil gland lid cells rare, 
isodiamenio, 32;au in chaneter, Constacted at sinus. 
Sinus undulate (single undulation), dark-staininag 
virculir region of thickened cuticle centres on sinus. 


Lid cell surrounded by a cyeloeyiie arnugement of s1x 
modified epidermal vells 

Affinty: The off wland id cells and general stantatal 
murphology (Rowell 991) contirnt affinity of the 
cuticle parataxon to Myrtaceae, 


Cuticle Parataxon No. LC 004 
FIGS 14-15 


Stomatiterous surface only. Abaxtal epidermal cells 
unuulate (3), beconting clongate over veins, 24-40¢m 
long, lo-32pm wide. Antichnal wall thin. smooth. 
Periclinal wall irregularly thickened. smooth to finely 
erunulate. Stomata randomly oriented, uniform 
distnbution within well defined urcoles, 16-24) long. 
12-l64m wide, Stomatal arrangement brachyparacytic. 
Guard cells sunken. 3-5 dark-staining. staurocy tically 
arranged epidermal ceils surround stonmatal apparatus, 
Raised cuticular folds that encircle and over-arch 
stomata may be present. Outer stomatal ledge 
prominent, deheate, narrow. Peristimal rim may be 
present. Subsidiary cells two, inconspicuous. Anticlinal 
wall thin, smooth. Periclinal wall irregularly thickened, 
smooth to granulate. Trichome bases vomunoy. over 
veins, poral, heavily cutinised pore. 4-6 scarcely 
modified epidermal cells surround pore, 

Affinity: The parataxon is assigned to the Proteaceae 
on the basis of the brachypuracytic stomatal 
arrangement (Cookson & Pike 1950; Blackburn 1981). 
Che rather inconspicuous. small subsidiary cells 
encircled by a staurocylic ring of dark-staining 
epidermal cells, stomutal density and sunken Momiata 
indicule an affinity w Bunkseaephvihim fastiganial 
(Cookson & Duigan 1950). i.e. Bankseuepliviane alt. 
B. fastigatum. 


Cuticle Parataxon No. LC 012 
FIGS 16-17 


Hypostomatic, Adaxial epidermal cells undulate to 
sinuous (3-4), becoming elongate over veins, 20-44 
long, 16-40; wide, Anticlinal wall thin, smooth, 
Periclinal wall thin, smooth. 


Pigs 15-20. 15, Parutiixen No, LO 004, Banksirdepiythen at&t fustigatam, Proteaceae, Shows prominent guard ccs, subsidiary 
colls dre generally inconspicuous. Scale 1 ci = 56 pm. 16, Purataxon No. LC’ 0)2, Barcksivapliylien all. Bobevanint, 
Priteaceac. Shows small clusters of raised stomales und simple, poral trichome bases scattered amungsr stomates. Svale 
fenp = 44 jun, 17. Paraiixon No, LC O12, Bankyiewephyuen wt, Bo obovatam, Proteaceae. Shows narrow light areus 
immediely inside dark-saining surmuading cells indivating the position of invenspicuous subsidiary celly. Seale | en 
= 32 jin. IR. Pardtaxon No. LC OW. Ranksicdephyllan ait, 2B ebayer, Projeaceae. Shows raised paracytic stonates 
und seatieree simple, poral trichome buses bul the absence of Jark-staining surrounding cells makes (his indentifeation 
fontative Seale Lom = 43 jun. IY, Paretaaon No. ABD 002. Plerostomautl, PR vamiofdes, Zamimeeae, Shows prominent 
cubeuhir pdges over (he antielinal wall of sinuous epidermal cells and heavily curimised stomares grouped im loosely defined 
bards. Scale } em = 100 jan. 20. Parataxon No, ABD (U2, Prerusrona alt. YP caminides, Zantiaceae. Shows “ban-bon"-like 
uppedrancy of the prominent vuter stomatal ledge. Note (eregular distribution and patterning of cuticukte ridges an epidermal 
wells Seale Lem = 53 pm. 


104 A. 1. ROWETT 


‘ 3 my 


Figs 21-24. 21, Parataxon No. ADB 005. Elaeocarpaceae. Shows high degree of ornamentation (fine striations) that characterises 
this parataxon. Scale | cm = 60 ym. 22. Parataxon No. ABD 005. Elaeocarpaceae, Shows guard cells without ornamentation, 
other than a fine apiculate outer stomatal ledge, Scale | em = 30 gm. 23, Parataxon No. AG O10. Lauraceae. Shows 
random arrangement of stomates. Scale | cm = 40 xm, 24, Parataxon No. AG O10, Lauraceae. Shows paracytic stomates, 
highlighting the prominent, narrow outer stomatal ledge, and absence of a guard/subsidiary cell wall, all of which are 


common features of the family, Scale | em = 30 pm. 


Abaxial epidermal cells rounded to undulate (3), 
becoming elongate over veins, 24 - 44m _ long, 
12-32m wide. Areoles well-defined. Anticlinal wall 
thin, smooth. Periclinal wall thin, smooth. Stomata 
randomly oriented, uniform distribution, 18-20um 
long, 1I4-20um wide. Stomatal arrangement 
brachyparacytic. Guard cells not sunken to very slightly 
raised, Outer stomatal ledge prominent, narrow. 
Subsidiary cells two. 4-5 dark-staining staurocytically 
arranged epidermal cells surround stomatal apparatus. 
Anticlinal wall irregularly thickened, smooth to 


beaded. Periclinal wall granulate. Trichome bases 
common, over veins, poral, some thickening around 
pore, up to six surrounding cells. Trichomes simple. 
unicellular, acute apex. Small poral trichomes, 
common within areoles, four radially arranged 
surrounding cells. 

Affinity: The cuticle parataxon has been assigned to 
the Proteaceae on the basis of the brachyparacytic 
stomatal arrangement. The rather inconspicuous, small 
subsidiary cells encircled by a staurocytic ring of dark- 
staining epidermal cells, stomatal density and raised 


LOCHIEL DISPERSED CUPICILES IWS 


sluimula indiewte an affinity to Banksivaeptivilare 
Phiovatiin (Cookson & Duigan 1950), Le. 
Barksicucphvilan alt. B. ebevarum. ‘The teature thal 
distinguishes this fossil species [rom the related & 
fastiganin is the superticial position of the stortata, 
ic. they ure slightly raised, 


Cuticle Parataxon No. ABN 002 
FIGS 19-20 


Hypostomatic. Adaxial epidermal cells sinuous (8), 
becuming elongale ever veins. 40-72pn1 long. 32-44dum 
wide. Anticlinal wall erregulurly thickened, smooth te 
beaded to buttressed, Periclinal wall irregularly 
thickened, striate t0 reticulate, Striations follow. cell 
outline. The curicular ornamentation may obscurecell 
outline. 

Abaxial epidermal cells sinuous (6), becoming 
elongate over veins, 44-80um long, 20-40pm wide. 
Antictinal wall irregularly thickened, thin, beaded. 
Beading imay uppeur slightly raised, Perichinit! wall 
irregularly thickened, granulate to striate, reticulate. 
Stomata randomly oriented. arranged in broad bands 
between veins, S.1. 27.3. Stomatal urrangement 
haplocheilic. Guard cells sunken, Subsidiary cells 4-5. 
Anticlinal wall regularly thickened, beaded. 
Perichual wall irregularly thickened, with prominent 
Mtriation in radiating pattern. Short cuticular folds 
common over the entire stomatal region. often 
associted with the stompte. Quice stumital ledge 
prominent, raised over lateral subsidiary cells narrows 
ty produce thin arcs over polar subsidiary cells. Gives 
outer stomatal ledge a “bon-fon-like appearance. Often 
absent trom polar subsidiary cells. 

Affinity: The cuticle has been assigned to extinct 
Prerostome (Hill 1980) of the Zamiaceae due to sinuous 
epidermal cells, cucicular ridges/folds on the abaxial 
surface. and a prominentand distinctive stomatal ledge 
(ban-bonlike appearance), The apparent regular 
venation patiern of the Luchicl specimens suggests a 
possible affinity to °F santioldes, 


Cuticle Parataxon No. ABD 005 
FIGS 21-22 


Stwomatiferous surface anly, Abaxial epidermal cells 
founded WW. undulate, becoming elongate over veins, 
l6-28am long, 8-20pm wide. Anticlinal wall thin, 
smooth. Periclinal wall irregularly thickened, smooth 
Io strate. Striations. obscure most cell detail. Stomata 
randomly oriented, uniform distfibution, 16-24 
long. 16-20um wide. S.1. 13,5. Stomatal arrangement 
slaunscytic? Guard cells not sunken, polar ruds present. 
Subsidiary cells 4-67 Outer stomatal ledwe evident, 
nairow. T-shaped thickening occasionally. present. 


Hydathodes rare. over veins, 36-44um long, 20-28 
wide. Striations radiate out trom hyduthode, 
Affini; The cuticle parataxon is assigned to the 
Elacocarpaceae On the basis hydathodes and what 
wppears to be a stuurocytic stomatal arrangement, 


Cuttele Paratuxon No. AG 010 
FIGS 23-24 


Stomatiferous surluce only, Abaxial epidermal cells 
angular in rounded, becoming elongate over veins. 
24-44ym long. 12-281 wide. Anticlinal wall thin. 
smouth. Periclinal wall thin, smooth. Stomata 20-24um 
long, 20-36um wide, randomly oriented. uniform 
disteybution, $.7_ 11.1. Stomatal arrangement paracytic. 
Guard cells slighily sunken. Guatd cell/subsidiary cell 
wall absent. Cuticular scales prominent, natrow. 
Subsidiary cells two, Anticlinal wall thin, smouih 
Periclinal wall thin, smooth. 

Affinity: The cuticle type is assigned to the Lauraceae 
due to the paracytic stomates and inconspicuous, 
sunken guard cells. 


Floristic Contparison of Samples 


The older lignite Nora comprising 38 dispersed 
culicle puralaxa is characterised by an abundance of 
Araucanaccae, Podocarpaceae, Myrtaceae and 
Proteaceae cuticles and minor occurrences (< 1%) of 
Casuarinaceae, Elaeocarparceac, Lauraceae and 
Zamiaceae cuticles which easily distinguishes a from 
the younger monospecitic flora. 

The floristic difference is-also reNlected in the twa 
lithatypes.. The lignite-of the older sample (Facies Ia) 
is defined as an earthy textured coal containing more 
than 40% relatively ungelified woody maternal 
(Springbett 1980 pers, comm.). Facies ITy isa darker 
fibrous, lignite consisting of gelified twigs (Fig. 2), 

Correlation between flora and lithotype indicates that 
the two floras were deposited in different sedimentary 
environments, and as the Kooliata Coal Zone consists 
of peatswamp and lacustrine sund and silt cycles 
(Kremor & Springbett 1992) the changes in the 
sedimentary environments are most likely due to 
fluctuations in water level. The degree of gclification 
also gives an indication of changes um water levels 
(Springbett pers. como); high degree of pelification 
— Jow water level and vice versa, 


Comparison with Other Australian Tertiary 
Deposits 


The dispersed cuticle Moras of the Lochiel depusit 
includes purataxa |hat occur in 4 number of other 


1S 4. ROWETI 


Australian Tertiary deposits. Buakreaephyiiian all. a 
Ivete. the pringipal parataxon of the Sedan lignites 4s 
Well represented at Lochiel, ‘This expands the known 
distribution Wf the species and establishes a Norastic 
link between the Northern St Vincent Basin and 
Lainibe Valley, fram where the type species was 
Griginally described. Thrs assuciaion 3s further 
strengthened by the presence of Banksicaephyllam aff, 
&, fastigaran and Banksiecephylumatf. ? B. obuvenen 
(Cookson & Duigan (950) both reported outside the 
Latrobe Valley for the first time. Burksiceepihen 
abewanen hus als been identified in the Miocene coals 
al Morwell where it occurs in low frequencies in the 
medium light coloured coals, the colour values 90-116 
(State Fleetricny Commission of Vicia Coal colour 
classification scheme, Blackburn |98S"y. All three 
cuticle types have affinities ta the ampdern genus 
Banksia (Blackburn 1985": Hill & Christaphel 1988). 

Pararaxon No. LC O|| m both the Lochicl and Sedan 
deposits bas an affinity vw the Myrtaceae, It is 4 minor 
component m the Horas of both localities but is most 
wbhundunt at Lochiel, The cuticle type has also heen 
Wentified from the Middle Eocene Maslin Bay 
secimeitts. 

The oteurrence of Prerosroma all, 7P zaminides 
(purataxon No. ABD 02) at Lochiel 1s significant in 
thal it represents the first report of Preroxtoma 
specimens oulside of south-eastern Australia and 
Tasmania. Prerostane is reported trom a number of 
Tertiary Jovalities, including Anglesea (Eocene, P 
minmiddes Hill 1980), Nerriga (Eocene, P 
anastumoesans Hill 80) Cecthyna (Ohigacene, 
Carpenter 1991) and Buekland (Eocene, Carpenter 
1991) and has a known siratigraphic range from the 
Cretaceous to Early Oligocene (Hill pets. comm). 
With such a extensive age range Tor Prerostoma 
paralaxon No, ABD 002 as of Jittle biostratigraphic 
significanec, 

Similarly, parataxon No, LC O15 with an affinity te 
Ducrvcarpus, is of Jittle biostratigraphic importance. 
BDacrrcarpis is he most common podocarp genus in 
Tertiary sediments ia south-eastern Australia which ts 
kauwn from numerous deposits ranging in age from 
Eovene to Oligocene-Miocene (Anglesea, Vegetable 
Creek, Yallourn, Bacchus Marsh in mainland Australia 
and Regatta Point, Loch Aver and Cethsna in 
Tasmania) (Hill & Carpenter 1991), Therefore cuticles 
assigned to Daerycarpus are unsuitable as 
biusiratigraphic indicators, However, indentificatian 
of Ducrycarpus cuticles at Lachiel does expand oun 
krxrwledge of the distribution of the genus during the 
Late Encene, 


' Blackburn. D. T. (£985) Palacobotany of the Yullourn and 
Morwell coal seains, Palaeahotanical Repory No. 3. State 
Hlecinieny Commission of Victoria 


‘There are a number of Angleseu parstaxa presem 
in the Lochiel flora, including No. ABP CHIL 
(Podocarpus aff. P. platyphyllumy, all representatives 
of the Elaeocarpacege and Lauraceae aml several oF 
the unknown cuticle types. 

In conclusion, analysis of the dispersed cuticle floras 
of Facies fa and Is, of che Lochiel lignite show that 
Two distinct floras exist, Te, (he diverse, Araticariaceac- 
dominated flora of the alder ungelified, woody lignite 
and the monospecific Nora (parataxon No. AW 007) 
of the younger dark, pelified lignite. The lahotype snd 
floral differences hetween lignites are most likely due 
to fluctuations in the hydrological cycle. changing From 
a Licustritte t peatswamp environment. These 
differences may prove useful in intra-betsin correlation 

The cispersed cuticle composition of the Lochiel 
lignites provides valuable information on the 
distribution of a number of Tertiary plant taxi, 
including Agathix, Barikstewephyllumn aff. B laeve, 
Banksteaephyllun aff 8B fastigatum and 
Banksieaephyllun aff, 7B obayatum, Dacrycarpus, 
Gymnostona, Podecarpys platyphyllun aod 
Pierastoma, The presence of thethree Latrobe Valley 
Banksieaephyllum species provides a interesting 
floristic link between the deposits and may he of some 
biostratigraphic significance. Rowett (1991) discussed 
this paint in relation to Banksicaephylien: afl. Bo laeve 
at.Sedan and concluded that its occurrence could cither 
imply a younger age for the deposit or an extendcdt 
lower limit 10 the age of the fossi]. These comments 
could apply to the Banksiewephyluay culicle types ul 
Lochiel. However, as the dispersed cuticle Mora is 
without any known Bocetie, Oligocene or Miocene 
indicators little can be concluded regarding the age at 
the Lochiel ligniles in addition to that provided by 
palynology. 


Acknowledgments 


The author is grateful for the support provided by 
N.E.R-D.D-C. grant No. 17174. Thanks go lo David 
Christophe] for his encouragement and assistance 
throughout this. project, tothe Electricity Trust of South 
Australia far access lo the deposit and to Andrew 
Kremot and) particularly Gavin Springbett for 
information on the Lovhiel geology and valucd 
comments on the manuscript, To Neville Alley of the 
South Australian Department of Mines and Energy for 
assistance in palynologcal siratigraphic information, 
and w Anthony Fox and Jennifer Clark fos their 
assistance wilh photography. 


LOCHIEL DISPERSED CLTICLES Kit 


References 


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Macrofossils from Tasmania. Ans), J. Bor 33, 645-656. 

BiackBurn, D. T. (1981) Tertiary Megafossil flora of Maslin 
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leaves. Alcherinva, 5. 9-28, 

Carrentir. R. J. (1991) Macracamia from the Early Tertiary 
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Aust. Syxt. Rot. 4, 433-444. 

Chkistornie. D.C, (1980) Occurrence of Casuarina 
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& Lys. S. D: (1986) Muminified leaves of two new 

species Of Myrtaceae from the Eocene of Victoria. 

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. Harris, WK, & Sywer, A. K. (1987) The 

Eocene Flora of the Anglesea Locality, Victoria. 

Alcheringa, 1, 303-323. 

. & GREENWOOD, TD. R. (1987), A Mevalossil flora 
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Cookson, LC., & Duican. §, L. (1950) Fossil Banksiewe 
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»&___ (1951) Tertiary Araucuriaceue from south- 
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Diccner. D. L. (1974) Approaches to the identification of 
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Greenwoopd, D, R, (1987) Early Tertiary: Podocarpaceae: 
Megifossils. from the Anglesea Locality, Victoria. Aust. 
J. Bor, 35, 1-133. 

Harris, W..K. (1965) Basal Tertiary microfloras. from the 
Princetown area, Victoria. Australia, Palueontographica 
B. 115, 75-106. 

——— (1971) Tertiary Siraligraphic palynology, Otway Basin. 
Mn The Otway Basin of south-east Australia. Spec. Bull 
Geol. Survy, S. Aust. & Vier. 67-87. 

Hu. R. §, (1980) Three new Eocene cyeads trom eastern 
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—— . & Bicwoob, A. J, (1987) Tertiary gymnosperms 
from Tasmania; Araucariaceae, Alcheringa MW, 325-335. 

— —— & Carventer. R, J, (1991) Evolution of Acmopvle 
and Davrycarpus (Podocarpayeae) foliage ay inferred from 
Tacrofossils in south-eastern Australia. Ausr. Syw. Bot. 4, 
449-479, 

—__ & Cueisrorue, D.C. (1988) Tertiary leaves of the 
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Kkemor. A. G. & Sprinapert, G. M. (1992) St Vincent 
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Bicentennial Monograph of Australian Coal Deposits, 
(Australian Government Publishing Services, Canberra). 
In. press. 

Rowrit, A. I. (1991) The Dispersed cuticular floras of South 

Australian Tertiary Coalfields. Part 1: Sedan. Truns. R. Soc. 

3. Aust. US (1), 21-36. 

~-— & Ciristoeuri., D.C. (1990) The Dispersed Cuticle 

Profile of the Anglesea Clay Lenses, Jn Christophel., D, 

C. & Douglas, J, (Eds) Proceedings of the 3rd International 

Organisation of Palaeobotanists Conference, (A-Z Printers. 

Melbourne, Victoria.) 

STACE, C. A. (1965) Cuticular studies as an aid to plant 
taxonomy. Bull, Br, Mus. (Nat. Hist.) Bet. 4); 1-78, 
SOUTH AUSTRALIAN DEPARTMEN! OF Mines AND KNERGY, 
Mineral Information Series (1987) Coal Deposits in South 
Australia. +27, (Woolman, S.A. Govt Printer, Adelaide. ) 

Srockey, R.A, & Tavior. T. N. (1981) Scunning electron 
microscopy epidermal patterns and cuticular structure in 
the genus Agathis. Scenning Electron Microscopy 3. 
207-212. 

Srover, L. FE, & Partertbce, A. D, (1973) Tertiary and Late 
Cretaceous spores and pollen from the Gippsland Basin. 
south-eastern Australia. Proc. R. Soe. Vier. 88(2) + 237-286, 

= ———— (1983) Spore-Pollen of the Wernllup 
Formation, Western Australia. Pedynelogy 4: 69-95, 

WILKINSON, H_ P. (1979) The Plant Surface (Mainly. Leaf) 
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of the Dicotyledons. Vol 1 2nd ed- (Clarendon Press. 
Oxford.) 


THE MORPHOLOGY OF NIPPOSTRONGYLUS MAGNUS, A PARASITE OF 
NATIVE AUSTRALIAN RODENTS 


BY IAN BEVERIDGE* & MARIE-CLAUDE DURETTE-DESSETT 


Summary 


Nippostrongylus magnus (Mawson) (Nematoda: Trichostrongyloidea) is redescribed from 
specimens from naturally-infected Rattus fuscipes from Blackwood, Victoria and from 
experimentally infected R. fuscipes and R. norvegicus. The asymmetry of the bursa, a characteristic 
of the genus, is matched by asymmetry of the spicules and genital cone. The synlophe is similar to 
that of N. brasiliensis but includes some variable features which appear to be of specific value. 
The morphological differences in N. magnus are discussed in relationship to the estimated period of 
separation from its congener, N. brasiliensis. 

KEY WORDS: Nematoda, Trichostrongyloidea, rodents, Nippostrongy/lus 


Trantseotinas of the Rovel Saciety of S. Aust. (1992). hice, O15 


THE MORPHOLOGY OF NIPPOSTRONGYLUS MAGNUS, A PARASITE OF NATIVE 
AUSTRALIAN RODENTS 


by JAN BEVERIDGE* & MARIE-CLAUDE DURETTE-DESSFTY 


Summary. 


Bevimior, | & Durerin-Drsset. M -C, (1992) The morphology ol Mippestremievies meaty, & parusite of nitive 
Australian rodents. 7rans, Ro See S. Aust, W603), 109-05, 30 November 1992, 

Nippestronavins magnus (Mawson) (Nematoda, “lrichustrongylodes) is redescribed fram specimens from 
nanuraly-brfected Kare fuseipes froay Blackwood. Vietora aad fron) experimentally infected Ri faieypes and 
R norvegicus, The asymenetry of the bursit, a characteristic of the yenus, is matched by asymmetry af the spicules 
and genial cone. The synlophe is similar ty that ot M Arusiliensis but includes some variable lealures which 
appear to be of specific value The inorphological differences in AU magus wre discussed in relanonship to. the 
estimated period of separation from ws congener, WV. brasihensis, 


Key Words, Nemaiwda, Trichostrongylenes, rodenry, Mippeseroneviias 


Introduction 


During a study of the helminth parasites of the bush 
rat. Rattus faseipex, a particular nematode species, 
Nippostromeyius magnus (Mawson) was encountered 
commonly in the duodenum. The species was 
described originally by Mawson (1961), although 
features of the complement of vuticular ridges, the 
synlophe, were not described. Some features of ps 
synlophe were described by Dtirette-Desset (1969) and 
by Vighienfels (1974), based on a small number of 
specimens, and in the latter paper their use for 
taxonomic purposes al the species level was 
considered. No magus has never been described in 
detail, and the use of the synlophe to identify species 
ol Nippastrongwus as suggested by Durette-Desset 
(970) and Lichtenfels (1974) has not been fully 
explored. 

Tt was evident therefore that a detailed redeseription 
of the nematode, particularly features of the synlophe, 
would allow a more definitive assessment of whether 
i provided useful taxonomic characteristics at the 
species level, as is the case in other trichostrongyloid 
genera. [t would also provide a basis for subsequent 
ultrastructural and life-history studies of this parasite, 


Materials ancl Methads 


Nematodes were obtaincd fiom aaturally infected 
Rattus fuscipes collected at Blackwood, Victoria 
(37°29'S, [44°19"E) and from laboratory raised R. 
fuseipes and R. norvegicus Which bad been infected 
experimentally with third-stage larvae of the species, 


* Departinent of Veterinury Science. University of 
Methourne, Parkville, Vic. 3052 

+ Biolngie parasitaire, Proustologie et Helminthologic, 
Museum national 2’Histeire naturelle, @l rue Bullion, Parix, 
W008 Pranee. 


Nematodes were collected live, washed in 0.9% saline 
and fixed in hot 70% ethanol. Additional specimens 
were fixed in 2.5% aluturaldelyde in phosphate bufter 
ut 4°C and embedded in resin, Sections bt am thick 
were stained with toluidine blue, examined under the 
light mucroscope and photographed, Whole specimens 
were examitred using Nomarski imerference contrast 
microscopy after cleuring in lactophenol and drawings 
were made with the aid of a drawing tube attached to 
un Olympus BH microscope. Apical views and trans- 
verse sections of the nematode body were prepared by 
hand using a cataract scalpel, Morphological terms for 
the complement of body ridges or synlophe and the 
numbering system tor bursal rays follow Duretie-Desset 
(1971, 1985), 

Numbering of synlophe ridges was based on relution- 
ship 1 the axis of orientation of the synlophe as 
described by Durette-Desset (1971). Ridges dorsal 10 
the axis were numbered from left to right 1. 2. 3 ete.: 
ridges ventral to the axis were numbered from lett to 
right 17.2". 3°.., , ete, Measurements are given in 
ium as the range followed by the men of five 
specimens in parembheses. 

Specimens examined have been deposited in the 
South Australian Museum (SAM), Adelaide. 


Nippostrongylusy magnus (Mawson) 
FIGS 1-20 


Austrohihgmonema magnon Mawson, WO, pp. Stb-817, fies 
46-47, trom Raviuy fusvipes, Ro rats, Ro conatus, R. 
norvexious aud Melomys ceryinipes; Duretie Desset (1969), 
p. 737, tig. 3 (as 4. magna from Ravras sp,), 

Nippostrong vias niagnas, Durette-Desset (1971), p. 88s 
Lichtetitels (974), j. 286. (an omegrter): Obendart (1979). 
Pp. 868. 896, 


Material examined: From Ratius fuscipes: adtural infections: 
Wee, 209 9, Blackwood, Vic. Joc, 49 P- 
deposited (SAM HC22477); experimental intections: 
dora. 20 9 (SAM HO22878): From Rattus nurvegicas 
experimental infections: 20cr-c° 169 9 (SAM HC22875). 


|. BEVERIDGE & M-C. DURETTE-DESSET 


WO 


SN. AtIGNUS FROM RODENTS tH 


Deveripiione Small. siniserally-coiled nematodes. red 
inculour when tive: prominent slietitly asymmetrical 
cephulic vesicle present, buccal capsule vestizial: 
Mouth opening sub-triungular, surrounded by six tiny 
labia) papillae; four double submedian papillae und 
pulred umphids present. external ii labial papillae: 
besophigus clavilorm, nerve ring in mid-ocsaphageal 
regiun, derrids dome-shaped. ui region af excretory 
puree 


Svlophe; composed uf 14 sidges in mid-body region; 
avis OF ociemation from right-ventral field to left dorsal 
field. at approximately 60° to sagittal axis (Pig. 15): 
caréne. nr cuticular swelling present in left dorsal field 
between fidges 2° jind 4: eight ridges in dorsal fiekl; 
ridges |4 diminishing in size, ridges 5 and 6 larger 
than 7 and &. six nudges jn dersal field! ndge 1” very 
large. dumishing in size to ridge 6; all ridges arise 
immediitely posierior ta cephalic vesicle excep! for 
tidges 3.2.1 which wrisc progressively between vesicle 
md excretory pore. ridges sometimes interrupted in 
wid-budy region, number ard Orientation uf ridges 
allers in posteriof extrendily at body: 


Mele, Length 3.3-4.243.7). maximum width (0-014 
(O11): cephalic vesicle 0.06-0.07 (0.065) long: 
vesophagus 036-053 (044), nerve ring O17 from 
anterior end) exeratury pore 0,29-0.32 (6.2%) trom 
unterior end: deirids 0.26-0.32 (0.29) from anterior 
end; spicules 0.50-0.54 (0.52): gubernaculumy (05 
long. Symlophe: additional ridge arises in right ventral 
field in region of spicules. between d.4§ and 0.85 fram 
posterior end. immediately anterior te bursit. addiuonal 
dorsa) ridge present, with cigit dorsal and eight ventral 
ridges; ridges reduced in size, orienution biurely 
discernible, ridges of similar size: irregular 
anastomosing and branching ul ridges seem close iv 
bursa. Bursa asymmetrical, right lateral lobe longer 
thin left; dorsal lobe reduced. Dorsal ray with tays 
Sarising acdifferent levels; lef ruy 8 more robust and 
ubising posterior ay right: major bifurcation of dorsal 
ray in posterior thitd of its lengths tays @ as long as 
internal rays (IO): latter with suggestion af secondary 
laleritl lobe; on teft, iy! 6 robust. arising clase to dorsal 
trunk, reduhing margin of bursa; rays Sand 4 slender, 
not reaching inargia ol bursa, common lateral trunk 
with proninent bulge at origin of ray. 5: rays. 3 and 
J elongate. slefider. reaching murgin of bursa: on right, 
hiy Ge short, slender, arising from Jateral trunk; ray 5 
slender, reaching matgin of bursa; ray 4 extremely 


nubust at base, extremity slender. reaching margin of 
burst; mys 3 ahd 2 stender. reaching margin of bursa, 
Genital cone prominent, clongite, conical. lightly 
sclerotised: ventral lobe simple with globoid, non- 
sclermtised apical appendage: dorsal lobe with (wo 
woequil pointed ends, typ surrounded by clonpate 
uppendage. Spiculcs elongate. wiquetrous in transverse 
section: spicule tips dissimilar; tip of left spicule 
knobbed. with ula arising near tip; tip of right spieule 
(iny. ald arising at tip: pubernaculum present. lightly 
sclerotised 


Female, Length 4.6-4.8 (4.7), maximum width in mid- 
body. region 0.12-0.14 (0:13), at posterior extremity 
O.14-0,17 (0.15): cephulic vesicle 0,06-0.08 (0.07) long: 
sesophagus 0.46-0,50 (0,48); nerve ring 0.20 trom 
unterior end: excretory pore 0.26-0.31 (0.29) from 
unterior end: delrads 0,27-0,31 (0.29) from anterior end, 
til 0.03-0.06 (0.05): vulva To posterior end Q.US-0,23 
(O10); egg 0.07-0.0%8 (0.07) « (03-005 (0.04). 
Symlophe: sante nomber of ridges in pasterior end al’ 
hudy: ridges become more prominent in region of 
nvejector. terminwte immediately umenor to yulva: 
Vdges of almost equal size. orientation almost lost in 
poslenor region. Posterior extremity of female with 
swelling af Cuticle, variible in shape, often forming 
sleeve over tip of call, Tail short. gonival. vulva close 
W anus: munodelphic, ovejector leads to short 
infundibulum, then into uterus; ep2 thineshelted, 
ellipsoidal, 


Discussion 


In spite of the fact that the sab-fanvily 
Nippostroagylinue ix cosmopolitan in distribution, and 
(hat jhe type spevies al Nigpasreenigyiie, ON, 
frasitiensis, has been widely used as a coudel in 
immunological research. few of the species as 
recognised by Duretle-Desset (1970) have been 
desertbed in detail. Features of the svnloplie in the iid. 
body region have been described fur varigus species 
by Chabaud & Duretic-Desset (1966), Dufetto-Desset 
(1969, 1970), Greenberg (1972) and Litchtentels (1974) 
Features. of the synlophe which might be useful in 
species separation have been investigated by Lichtentels 
(1974) following a detailed examinution of the synlophe 
in Jaboratory strains of No Areasiliensis tind linnited 
observations on several additional species. Equally 


—_—_——--errrn ng 


Bigs bh? Mapospeanaylay mnegaus (Mawson 1.2. cephutic eatrentity duteral view, Sowing usymamieny vf cephalic vesivic! 
2, apical view of anterior extremity: 3, cr. unterior end, lett side showing origins (arnowst qt ridges 12 and 3. 4 =. 
untenor end: right side showing origins of ridges at cephalic vesicle; 5, oF left view. al level ol spicules. 0.5 mai trom 
Posterfer end, showing origin Gierow) of additional ridge and branching and wnustomosing af ridges, 6 mid-bady rezinn 
ocr. lef side, showing diseuntimuities (arnnys) in ridge: 7 female til night se. showing sleeve furmed by cutiele 


und GeninaGon OF tidges Seale lines: Figs L2. OO min Migs 3 ME i 


Levene: 4, onphids d. derridy dee dorsal: 


ce calretary pores |, Tibial papilla> p. submmedian pupils v. vena 


Wz 


| BEVERIDGE & M-C. DURETTE-DESSET 


4, MAGNUS PROM RODENTS 113 


detailed stadies however have not heen made on) any 
coigeners. Thus, apart trom providing a basis. for 
i)irusiructural studies currently underway, she detwiled 
deseription of AW. ayy is. considered valuable as a 
compurison with studies already carried oul an AL 
Arasilen ds. 

The asymmetry of the bursa has beet noted in ewch 
cnngeéner, The bursa ts best studied in apical or ventral 
views (Durettc-Desset 1985), however, in species of 
Nippostrongyiay i issextremely difficult to upen the 
bursa. becnuse of ns asymmetry, For this reason, Jef 
and right titer viewsare provided (Figs & 4) aswell 
as an apical View (Fig, 10), which was obtained using 
live male specimen prior to fixation. The greatest 
morphological asymmetry vccuts in rays 6 and #, both 
of which ere much largeron the left side of the bursa 
than un. the right. though ray 4 is Jarger an the right 
side. Apart from the bursa utself, the spicules und 
genital cone alse exhibil some degree of asynimetry, 
The tip al the lefi spicule is much lonver and nere 
complex stvucturally than (hat of the right spicule. 
which terminates in a simple point. paralleling the 
asymmetry af the bursa. Detarls of the spicule tips have 
nul been provided tir congeners cacepe for the ups of 
the spicules oF N. drusifiensis (see Mawson WL). In 
the case oF the genial ceme. the (ventral tobe, bearing 
papilla 0. is synuneteical. while the dorsal lobe, bearing 
the paired papillae 7 1s asymmetrical. with the right 
piprile longer ind hence more pasteriur thin the left 
(Figs JA 14). Companible morphological details ure 
generally lackiny for other species, although the genial 
tone appears to be usymhimecirical alsaan Fig. IF af NV 
rausehi (see Chabaud & Duretie-Desset 1966). Some 
of these characters maty prove useful as eenerie eriteria 
When described. in all species 

The sy nlophe is deseribed fully for the first time und 
confirms the preliminury ubservaliany af Durette- 
Desset (1969) and Lichtenféls (1974). [t resembles that 
of congeners (Chalxiud & Dureite-Dessct 166: 
Dunste-Desset 1970, 1971; Cireenbere 1972) in 
possessing I4 ndgey in the mid-bady region with an 
Oblique axis of orientation directed from right-ventral 
wo left-dorsal and a consistent) wrauhient in rides size, 
The ihaparity of ridges Arise immediately posterior in 
the cephalic vesicle, with ridges |,2,3 in the lefi-dorsal 
field (ridges 224 of Lichtenfels 1974) aristag 
immedintely antetior ta the deirid (1), hallway hebween 
deirid und cephalic vesicle (2) und posterior to the 
vesicle (3), These Origins are consistent in ditles and 
foitiales afd resemble the situation found in Nv 


brasiliensis except that in N. brasiliensis, ridge | arises 
slightly more pusteriorly, at the level of the excretory 
pore (Lichtentels 1974) In (he posterior region of the 
Wile, Iwo additional Hdges appear in Lhe left-vent rat 
field. also resembling the arrangemenc deseribed i 
No daisilionsix (sex Liemenfels W742 une abaut 
OS 4AL& min from the posterior extrenvity und a seedind 
odge inthe prehursal regan, In the pastestoy revi 
of the female. the number of ridges remuins constant. 
ulthough the ndges become more similar ia size wn 
the virientitian js more difficult toestablish The extrs 
ridge described on fenwle NV. brasiliensis by Lichtentels 
(1974) ts abseot in NV, meesus. Thus the synlophe of 
No neigeus resembles that of No brasitensis very 
closely. 

The system for numbering ridges employed here 
differs [rom that used by Lichtentels (1974). Tt attenrpts 
ty show the axis of orientation and the homology of 
ridges on either side of the uxis. Irdenonsrstes that 
in both the male und fernale of Noorrayiuy. the 
asymmetry oF ridges and the sive pradient are lost iri 
the posterior parts of che body with a syminetricul 
ubrangement of almost equal sized ridges. fiestty 
arranged perpendicular to the body of the nemalnde 
This armangement would be ermsidered i “hy per- 
evolved” state in the sense of Durerte-Desset (1985). 
It is of interest that in male WW. magus. inthe posterior 
region of the body. not only is there y reduction ii size 
of body ridges and a loss of particular orientation, bul 
ilso the syminetry of the nuinber of ridges is restored 
with eight dursal and eight ventral ridges, 

Features ot the synlophe of N. magnus Which meh 
be useful at the specific level are the interruption of 
ridges in the mid-body region and the irregular 
branching and unastomosing of ridges in the region of 
the male bursa. noted by Lichtenfely (1974). In the 
present study, the interruption of ridges (Fig. 6) 
occurred in both male and female neniatodes, while 
hranching and anastomosing (Fiz 5) was seen in 
inales. Thos Lightentels’ (19741 observartons have been 
confirmed. but studies of the remaining congeners are 
required to establish their usefulness. 

Lichtenfels (974) examined specimens of laborstory 
sitams of No Srasiiensis adapted to the rat, mouse snd 
hamster and showed that the syqlophe was constatt, 
Independent. of the hast species in whieh the hematode 
developed, Although much more limited ire their extent. 
the Observation that the synlophe of No magnus is 
Mentical in specimens from the natural host. AL 


Jiwetpes, as wells in the laboratory mat, AL warvegicus, 


Figs 8-14. Nippesmongvhay meas (Mawson), ale genitatia, & bursa, lett haleral and dorsal lobes: 9 bursa. righy lateral 
did Uirsal fohess 10, Hirsa pied! view. [tt ktteral lobe on right hand sides 1, nip of right spicules IZ, ip af lett spneute 
2. genial cone latenul view: I, geniil cone and spteute nips. right ventny- literal view Sevle lines: Digs 80. U.0 mine 
Figs Heh). UOl mm, Pigures (blow ray numbering syste os ceseribed by Duretio Desset (ORS) Levend: do. dirsal: 


te, fell rh, righ, 


14 


I, BEVERIDGE & M.-C, DURETTE-DESSET 


be 


" LARD 


NO MAGNUS FRONT RODENTS 15 


adds weight io his conclusiois On the stability. oF 
synlophe charvelers in different host species. 

The alfinitics of M. mages with congenets have fot 
heen fully investigated. Mawson (1961) considered its 
differentiation from N, fyjitcns thotl as species at 
Auvtrohelignoiera Mawson, 1961) based on the shape 
al the spicules, nuinber of ridges and overall size and 
iru NV, brasiliensis, due to the greater asynimetry in 
is bursa. and the fornt of the dorsal ray. Greenbery 
(1972) provided a comparatve table of measurements 
ofall species, but not of other morphological features. 
Because of the incomplete nature of the descriptions 
uf several species. vomparisons are lirnited to the 
synlophe and bursal rays, The synlophe is apparently 
similar in most species of Nippostronevtas, but ridge | 
is substantially larger than ridge 2 in V, mages. the 
malé af Wo fyitens, and N, rausehé. with the 
qualification that A. ravsed is described a5 having I 
ridges, bur only (3 are iMustrated (Chabaud & Durette- 
Desset 1966, Fig, 2A), In the case of the dorsal ray 
OLN. mesris, the asyinetry ol nays & with a slender 
right ray arising before a more robusi lef ray resembles 
N, typicus, bul differs tram No rauschi, N. bravitiensis 
and A, djumachani which have pays & arising 
symmennically, though with the left ray more robust 
than he nght. and fromN, oyevw in which the lett 
ray 4 arises first and is more slender chan the right 
ray (Erhendova 1959: Mawson 1961; Chabaud & 
Durette-Dessel 19665; Tenora 1969), In NL wilenbengi, 
the branching pattern of the dorsal ray resembles that 
Of MW. eyyervy?, but rays 8 are slender (Greenberg 1972), 


Thus, NV tenes cat be differentiated from congeners 
by several morphological features, in addition i the 
measurements tabulated by Witenberg (1972), bur de 
features discussed indicate a close relationship with 
N. pices, also a patasiie of endemic Austratiun 
rodents, 

N. magnus wolf biogcogtaphical interest because jt 
is an endemig Australian species occutring, in various 
species of Raitus and occasionally in Melons 
cervinipes, The tll hast range may be greater than 
this as a number of endemic rodene specics in Australia 
have not yet been examined for helminth parasites 
(Magkerras 1958). The endemic species of Ratity 
probably arnved in Australia about ome millian years 
age [Waits & Aslin 1981), henee the murphological 
differentiation between N. hrasiliensix/N. raayeh) and 
N. mageys!N. rypicus has probably ocurred ever (his 
same period of time. There ure few instances where 
uv time scale can be placed on morphological 
differentiation between species of parasitic nematodes. 


Acknowledgments 


We wish to thank Christine Andersen for excelleiit 
technical assistance and Dr D. M. Spratt for comments 
car the manuscript. This work was supported 
finanelally by the Australian Research Couneil. Rats 
were trapped under a penmt from the Vietorian 
Department of Conservation and Environment (RP 
SI-NOS}, 


References 


CHazaun, Ao G & Dtircrte-Desser. M-C. (1966) 
Nippestrangyins rawscht np. Nematode parasite de 
dennopteres et consiterations sup NV, bresitiensis parasile 
cosmopalie des rats donmastiques. Arm. punialnrt. dian, 
comp 41, 243-249, 

Dugerte-Dessut, MAO. (1969) Les sysicmes daretes 
cunculaires chey Jes nemutites heligmosomes purasiles ile 
Muride’s austratiens JAid 44, 733-747 

(W970) Le gente Nippextravevius Lane. 

(Nematode-Heligmosonatide), ive. 48, 815-824 

(1971) Essai de velassification des nenatudes 
Keligmosoimes. Correlutiony-avec la paleobiogcagraphic des 
hotes. Mein. Mus. natn. His. aar.. Paris ser A. Zoologle, 
69, 1120. 

(1985) Trichostrongyloid nematodes and (heer 
Vertebrate hosts: fecanstruction of (le phylogeny of a 
pacdsiliy group, Advan, Parasitol. 24, 239-306. 

EknARDWA, Bo (1959) Osnildonema rysavit tsp. ond 
Vieuella chinensis n.sp. (Nematoda: Heligmosomuridae) 
bet Chinesischen navern. Ceska. parasinil 6, 93-96 


1923, 


Greeserrs, 7. (1972) Helminths of birds and) munimals 
from tarael TY Helivintis Crom Nesakter fdica Gray. & 
panaeeen (832 (Rodentia: Muridag), Israel) Zoal, 21. 
64370, . 

LicHTENFELS, | Ro (1974) Nunvber and distributional. ridies 
in the cuticle of Nippastronamis brasilignsts (Travansos 
1914) (Nemaioda: Hehgmosomateidead, J. Paravitel, 4, 
285-288, 

Mawson, P.M, (1961) Trichosrrongyles Iron rodents in 
Queensland with comments cin (he genus Limgdstrdetea 
(Nemutoda: Heligmesomatidae). das J. Zoal. 1 O)426 

Mackerras, M. &. (958) Carlogue of Austniliaa mammals 
and theit revorded internal parasites. UH. Butheria. Prac. 
Lan, Soe. NS. Wiles 83, 101-60, 

OveNborr. D. L, (1979) The helminth parasives of Reins 
fuseipes (Waterhouse) tran! Victoria, including description 
of wo new nematode spectes. Aust J. Zonal. 27. 867-879. 

TeENORA, F, (1969) Parasitic nemunmdes of cenaln rodents 
from Afghanistan. Keateit Cede, spol. Zoot 33, 174-192, 

Wars, CH, S & Asus, HJ. (E981) “The Rodents al 
Australia” (Angus & Robertson, Austratia), 


OOO OOO nnn 


Figs (5-20. Nippusrroigvlus mares (Mawson), synlophe. 15, male. mid-body region, 19 niva fromvanterior end of 33 mm 
worth with full complement of t4 ridges: arrow indicates axis of orientationof synlophe: 16, mate, anterior esophageal 
region prior io origin of ridges. 2-3; 17, male, O88 my [run posterior end showing additional ridge 1A), 18, male, immediately 
watcrinr to bursa, showing reduced size of rupes and additional ventral fidges, 19. female, mid-body revion, with full 
complement of 1 ridges; 20, female, 0.80 im from posterior extremity showing reduction in size of ridges bur jiaintenanice 


of sume wamber al ridges. Scale le: OOL tom 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


VOL. 116. PART 4 


EARLY DEVELOPMENT OF LIMNODYNASTES TERRAEREGINAE AND 
L. FLETCHERI (ANURA: LEPTODACTYLIDAE: LIMNODYNASTINAE) 


BY MARGARET DAVIES* 


Summary 


The development biology of Limnodynastes terraereginae and L. fletcheri is described. Life history 
data generally conform with those of congeners in the same species groups. Tadpoles show lentic 
adaptations and have a generalized body form. At 30°C L. terraereginae completed its development 
in 71 days and L. fletcheri in 60 days. 

KEY WORDS: Anuran larvae, development, Limnodynastes terraereginae, Limnodynastes 
fletcheri, Limnodynastinae, tadpoles, foam nests. 


Transactions af the Royal Sueiety of §. Aust. (1992), Wed), 11 622. 


EARLY DEVELOPMENT OF LIMNODYNASTES TERRAEREGINAE AND L. FLETCHER! 
(ANURA: LEPTODACTYLIDAE: LIMNODYNASTINAE) 


by MARGARET Dayies* 


Summary 


Davies, M. (1992) Barly development of Limnod\nustes terraerepinae and 1, fletchert (Anura: Leptodactylidae: 
Limmnadynastinae). Trens. R. Soc, S Aust. 11604), 117-122, 30 November, (992. 

the development biology of Limnodynastes rerraereginae and Lb. flereher! is described. Lite history dats generaily 
conform with those of congeners in the same species groups. Tadpoles show fentic adaptations and have a generalized 
body form, At 30°C, L. terraereginae completed its development in 71 days and /.. flercheri in 60 dayy. 


Kry Worns: Anuran larvae, development. Limnodynastes terracreginac, Linkadynastes fetcheri, Limnodynastinac, 


(adpoles, foam pests, 
Introduction 


Limnodynastes Fitzinger comprises 12 species (Tyler 
1992) that can be grouped into three: morphotypes; 
burrowing species with short hind limbs and a rotund 
body, 29-46 mm in body length (two species: L. 
ornatus group), larger burrowing species (52-90 mm 
length) characterised by well-developed tibial glands 
(four species: L. dorsaliy group) and more streamlined 
non-burrowing, species found in marsh and flooded 
grassland, and ranging from 31-75 mm in body length 
(six species: L. tasmaniensis group) (Tyler et af, 1979). 

Limnedynastes spp: lay foamy egg masses (but see 
Roberts & Seymour 1989), but the entire developmental 
biology has been described only for L. ernatis (Tyler 
etal. 1983). Watson & Martin (1973) described tadpoles 
of L. inerioris, one of the large burrowing species and 
examined larvae of L. dumerilii, L. terraereginae, L. 
peromi, L. salmini, £, tasmaniensis and L, fletcheri 
to permit a generic definition. However, these authors 
did not describe these larvae specifically, nor did they 
provide complete developmental data lor L. interioris, 

Martin (1965) described tadpoles and early stages 
of L. dumerilti, L. peroni and L. tasmaniensis whilst 
Tyler et al. (1983) described tadpoles of /.. 
convextusculus, Here I describe the developmental 
biology of Limnodynasies ierraereginae Fry, (I., 
dorsalis group) and that of L. flercheri Boulenger, (L. 
lasmaniensis group). 


Materials and Methods 


Spawn of L. terraereginae was collected 7 kin SW 
Pentland, Qld and. transported to the University of 
Adelaide after cight days, where it was reared at 
30 +°C in aerated dechlorinated tapwaier contained 
in glass aquaria 25 x 25x 8 cm. 


* Department of Zoology. University of Adelaide, GPO Box 
49%, Adelaide, S.A. SOI. 


Spawn of £. fletcheri was obtained from individual 
adults from Deniliquin, N.S.W., that spawned in the 
laboratory. This material was reared in the same manner. 

Developing larvae were fed lightly-boiled, 
organically-grown mignonette lettuce leaves 
supplemented with SERA bioflakes pond fish food. 
Water and food were changed daily. 

Samples of eggs, embryos and larvae were collected 
as shown in Tables 1 and 2 and preserved in Tyler's 
fixative (Tyler 1962). Cuts were made with a scalpel 


TARLE 1. Dimensions of developmental stages of 
Limnodynastes terraereginae. 


Age (approx. Stage Body Length Total Length 
days, date) Xx, Tange in ¥, range in on 
parentheses parentheses 
1 20 2,37 5.24 5 
27.1.1991 (2.24-2.4) — (5,2-5.68) 
4-17 25 4.19 12 62 
28.1, 1991-12. 1.199] (2.4857.2) — (7.12-19.52) 
{7-22 26 6.69 14.08 10 
10,14.1991-15.4j 1901 (5.6-7,2) (15,84-19, 52) 
21-28 27 9.27 25,15 4 
14,41, 1991-8, iii, 1991 (704-11,.97 — ((8.4-32.13) 
28 28 8.72 34,45 } 
QALAST 
55 29 1103 29.72 2 
19,0 1991 (105-1155) (29 4-30.03) 
28-55 31 1.07 35.69 3 
21, 11.J991-19,G1, 991 (8.72-12.45)  (34.03-38.18) 
55 az 14.12 38,81 Z 
19. 1ii),1991 (13.7-14.53  (38.18-39.43) 
50-70 44 5.3 42.94 5 
14. it) I99}-7.iv.199] (14.53-15.8) — (41.5-44.6) 
55-10 36 15.51 43.77 7 
19.411.199)-7, iv, 1941 (14,.0-16.5)  (39,5-46,7) 
55-70 a7 15.98 45.B4 RB 
19.411 1991-71. 1991 (40476) — (4L749.7) 


1K M BAVIES 


in the presumptive region of the Ubral gland of L. 
terruerevinee in the Metymorphosing specimens. 

Measurements of developitiental stuges were made 
uyitig un eyepiece micrometer or vernier calipers 
(reading to 0.05 min). IMlustrations were made with the 
aid of a Wild MB stereodissecting microscope uad 
cumers Jucida. 

Developmental stages are thase ol Gosner (1960) 


Results 


Linnnidynestes terraereg ite 

Spawn was collected from a heavily-vegetated. deep. 
roadside depression al. approximately O800 on 
Mri 1991 Adults ol L. terrvereginge, had been heard 
calling at this site the previous night, just before 
midnight, under an isolated patch of vegetation, The 
foum nest was large, bul an ege coun! Was not obtained , 
nor Wete vges preserved, 

At 2130 on 28.1, 991, embryos were ut suze 25 (Fig, 
I) Adhesive glands were reduced to proniment 
pigment patches, The horny beak und one upper and 
one lower complete tooth rows were keratinized. The 
goal opening was median und the spiracle had formed, 
Eiubryos remained al stage 28 for a further 24 hr. 
Feeding had ¢ummenced (faces were in the anal tube). 
Faim pigmentition was detectable on the adhesive 
glunds, One complete and one divided upper and two 
complete lower labial tonth rows were keratinized, 

Embryo, sampled on 307.1991 and 111.1991 
rempined at stage 25. Pigmentation on the adhesive 
vhinds was detectable although considerably reduced 
in the oider embryos. Further keratinization of tooth 
rows Was not apparent. 

Adbenwwe gland pigmentation had vanished by 
411.199), although embryos were still at stage 25, The 
body was more heavily pigmented together with the tail 


Mee. 1. Eebryas of Aviendyitestes rercaereginene he Stage 25, 
bh. Stave 20 of butching. Scale hars }mm 


Wusculature and a lighter dusting oa the ral fins. A 
second divided upper fibial tooth row was keratinized. 

Embryos sampled on Zi. 98! remained at stage 25, 
the only change being an increase in pigmentation 
particularly on the tail fins. Measurements are given 
in ‘lable 4, 

A single stuuge 26 larva was sampled on 10.11.1991 and 
a second on 12.11.1991. Piymentation had increased in 
these farvac. Stave 27 larvae sampled on 14.0,1991 had 
strongly pigmented bodies, tail musculature and tail fins. 
By 15.11.1991 stage 27 larvae had a further divided upper 
und a divided lower tooth row keratinized, 

By 21, 4i:1991, larvae had reached stage 28 and 
pigmentation of the body and upper tail musculature 
had darker pigment pawhes superimposed over the 
uniform background. Most of these pigment patches 
surrounded the neuromasts of the Literal line organs. 

By 8.iH.1991, a single stige JI larva bad «well- 
differentiated. lateral line organs and the hind: limb 
paddle was pigmented dorsally, extending along the 
mediolateral surface of the hind limb bud by stage 34. 
Larvae attained stage 37 by 14.111, 1991, 

Measurements of all larvae sampled wre given in 
Table 1, 

A larva at stage 36 as shown in Fig. 2. The body 
is ovoid and widest behind the eyes, The snout is evenly 
rounded in dorsal and lateral views. The nares arc 
dorsolatcral and sessile, opening laterally. The eves ure 
lateral and telatively large, The sprracte is sinistral. 
short and ventrolateral with a small orifice directed 
posterodorsally, [f is atached along its medial edge to 
the body of the larva, It is visible from above and is 
slightly. tapered towards its orifice, 

The anal tube is medial and opens at the extremity 
of the ventral fin. ‘The tail fins are poorly arched and 
rounded terminally. The dorsal fin commences in the 
posterior (/Qof the body, being deepest about halfway 
dong ifs length, The tai) musculature tsthick, lapering 
lo a subacuminate terminus. 

Tadpoles are moderately beavily pigmented and 
chocolate markings usually surrounding, neuromast 
cells Of whe literal line are superimposed on the 
background pigmentation, The mouth is ventral and 
the oral dise is surrounded by lateral and posterior 
labial papillae interrupted by an anteromedial gap. 
There are five or six upper und three lower labial woth 
rows. The first upper and second and third lower rows 
remain undivided. The horny beak 1s moderately 
robust. ‘The oral dise of # larva at stage 36 is shown 
in Fig, 3, The frst larva reached stage 42 on 2.iv. 199] 
and stage 46. on 7.199) having taken a total of 71 days 
to metamorphose from spawning, Froglets al stage 46 
measured (8.00 mm S-¥, 

Supralubial glands were apparent at stage 42 ane 
although not apparent externally, glandular lissuc wus 
detected by cye in out skin in the region al the 
presuiiplive tibial glued, 


EARLY DEVELOPMENT OF LIMNODYNASTES SPECIES 


Fig. 2, Larva ot Limnodyerasies terracrevinae at Stage 36y ii, 


Lininadynastes. fletcheri 


Spawn was laid in the laboratory in a chamber of 
recyling, continuously-flowing water described by 
Chapman (1987), Two spawa clumps deposited 
overnight on I6/17.xi.1991 and 21/22-x1.1991 


Fig. 3, Oral dise af a larva of Livvtodvidstes terraereninge 
at Suge 36, Seale har = | mm, 


19 


lateral View: b, dorsal view. Scale bur = [0 mm. 


respectively were collected, Data are mostly derived 
from the first clump. Spitwn was laid in a foamy nest 
and twelve eggs from the second clump had a mean 
diameter of 1,39 mm (range |.28-1.56). Mean capsule 
diameter was 1.80 mm (range 1.72-1.88). 

Within 24 hours the embryos had reached stage 18 
(Fig. 4) and hatched at stage 19, 24 hr later on 
20.x1,1991 (Fig. 4), Gills were poorly developed, The 
mouth was perforated but the nares were not, Tail 
musculature was poorly delincated- 


At 48 hy, the gills had disappeared, but the embryo 
remained at stage 19, 

Embryos reached stage 25 four days after hatching. 
The snus was median. Slight protrusions of the 
adhesive glands remained. the base speckled with 
pigmentation, Over the next three days, the adhesive 
glands disappeared. The anus moved from a median 
toa dextral position and the horny beak, together with 
firstly one and thea two upper labial tooth rows and 
three lower tooth rows, keratinized. 

Measurements of stage 25 embryos are piven in 
Table 2 and one is illustrated in Fig. 4. 

Stage 26 and 27 larvae were sampled on the seven 
days after hatching. By stage 27, all tooth rows were 
keratinized. The body of the larva was irregularly 
pigmented and a faint dusting of piginent was apparent 
on the dursal tail musculature and tail fins. 


120 M. DAVIES 


Fig. 4. Eaubryos of Linmodyrastes fletcheri: a, Staze 18; b. 
Stave 19 at hatching: ¢. Stage 25, Seale bars = 1 mim. 


Larvae reached stage 30, 14 days after hatching, The 
anus was median and patches of chocolate pigmentation 
were appearing, superimposed over the background 
coloration of the body and tail. Measurements are given 
in Table 2. 

Larvae reached stage 33 23 days alter hatching and 
stage 35 after 27 days. 

A larva.at stage 37 is illustrated in Fig. 5. The body 
is avoid and widest behind the eyes. The snout is evenly 
rounded in dorsal view and slightly truncated laterally, 
The nares are dorsolateral, not clevated, opening 
anterglaterally. The spiracle is sinistral and relatively 
short and is visible from above. Ir is attached to the 


body wall along its medial edge with the diameter of 


its orifice being less than the diameter of the tuba. The 
spiracle tapers towards its orifice. The anal tube is 
median und opens along the ventral edge of the ventral 
tail fin. The tail fins are arched, the dorsal fin 
commencing in the posterior 1/10 of the body. Both 
are deepest approximately half way along their length. 
The tail fin is slightly rounded at its terminus. Tail 
musculature if moderately thick, tapering to a point 
posteriorly. Blutchy chocolate pigmentation on a cream 
background is located on the tail musculature with 
weaker meélanic patches on the fins, The body is 


TABLE 2. Dimensions pf developmental stayes af 
Limnudynastes fletcheri. 


Ave (approx, Stage Body Length Total Length n 


days. date} &, range in x, range in 
parentheses parentheses 
6-l 25 4.97 12.49 27 
23.x1, 1991-24. x1. 1991 (4.0-6,.24) — (9.76-15.2) 
10-17 26 7.01 18.07 6 
IENIA991-A4. x11. 1991 (64-80) (12.0-22.0) 
W-26 27 7.25 19,83 12 
27.81. 199)-13. xii, 1991 (651-861) (17.22-23.31) 
17-26 28 979 26.48 3 
4. xt 1991-13. x11.199] (9.1310,29) (24.9-27,.46) 
26 2p 10,49 27.73 4 
13x11, 199% (9,96-10,79)  (26.56-2%.59) 
17-26 30 H91 3071 3 
ANU T9OT-13. x11, 1991 (11,62-12.06) (29,88-31.54) 
47 Al 10:85 30.45 2 
15.1.1992 (10,3-11.4) —-(30,2-31.5) 
23-52 32 13.83 36.96 18 
9, x1) 1991-15. 7.1992 (10.8.17.43)  (27,6-46.07) 
26-47 33 13.35 37.20 0 
13, x17, 1997-15.) 1992 (12.0148) (34.4-40,26) 
47-52 34 14.36 38.83 14 
Wh xid, 991-15, 1.1992 (IRAAS.8) — (34.0-42.0) 
47-52 35 16.23 44.23 4 
8.11992. 15.1, 1992 (15,3-18.0) (406-487) 
32-70 36 17.0%, 48.32 5 
8.1.199226.1,1992 (16.0-189) — (44.2-52.4) 
70 37 18.86 $2.24 13 
26.1,1992 (175-205) (48,.4-56.0) 
70 38 19.0 57. 2 
26,1,1992 (181-199) (56.9-457.9) 
70 39 22.0 58.5 | 
26.1.1992 
710 4g 24,2 68,5 } 
264.1992 
65 42 22.0 67.9 | 
24,1,1992 
78 43 26.1 43.5 \ 
311.1992 
80 al 23.5 60.1 | 
5.41. $992 
74 44 22,9 34.8 | 
AD.i1.1992 
79 44 22.5 29.6 1 
4111992 
60 46 21.2 - i} 
141.1992 


FARLY DEVELOPMENT OF LIMNODYNASTES SPECIES i 


fi S, Larva of Linwadvnastes flercheri at Stage 37: y, lateral view: b. dorsal view, Seale bar = 10 mm, 


moderately pigmented with « puler cream 
posterodorsally. Chocolate freckles and smaller 
blotches dre superlinpased on the background 
pigmentation, 

The mouth is anteroventral, The oral disc is 
surrounded laterally by a single row and pasteriorly 
by a double row. of labial papillae. Papillue are absent 
anteromedially. There are three upper and three lower 
rows of lubial teeth. Only those rows adjacent 10 the 
beuk, which is moderately keratinized. are divided 
(Fig. 6), 

Many of the oral discs exymined were abnormal in 
development with split beaks and incomplete or 
distorted tooth rows. 

Meusurements of larvae are provided in Table 2. 

By 12.1.1992 a larva hud reuched stage 42 and by 
15.1.1992 it had reached Stage 46, 60 days after 
spawning, Apart from a further three or four 
individuals, the remainder of the spawn clump did not 
metamorphose until about 30 days later, ie., about 
1), 17.1992, 

Body length at metitmorphosis was 21.2 mm- 


Discussion 


OF the tadpoles of large burrowing species of 
Linnodynastes, only those of L. interiaris and L. 


dunwrilit have been deseribed and illustrated (Watson 
& Martin 1973; Martin 1965). Larvae af L. 
terruereyinae described here have a similar 
morphology to that of these species although the inner 
edge of the spiracle may be free in L, dimerilii and 
these larvae are usually darkly pigmented. with older 
tadpoles being generally lighter (Martin 1965). The 


Piz. ft. Oral dise of larva af Linmodvnastes fehert ar Stage 
37. Scale bur = 1 tin, 


22 M, DAVIES 


general Jentic body form of the three species is similar 
(described as generalized by Watson & Martin 1973). 

A tooth row formula of five or six upper and-three 
lower labial tooth rows is consistent within the group 
and the pattern of labial papillae is common to the three 
speciés, 

Ofthe marshy and flooded grassland species, larvae 
of L, taxmaniensix and L, peroni have heen described 
by Martin (1965) and those of L. convextevcitus by 
Tyler et al. (1Y83)- 

Tail fins of these species and of L. flewheri tend to 
be more strongly arched than those of the L. dorscliv 
group. 0. peroni has four upper ard three lawer labial 
tooth rows, L. tasmaniensis have five upper and. three 
lower rows and L. convexéuyculas has five upper and 
three lower rows. of labial teeth (Martin 1965; Tyler 
etal, 1983). Watson & Martin (1973) recorded at least 
four and usually five to six rows of upper labial teeth 
for L. peront. L. salmini, L, tasmaniensis and ©. 
fletcheri. 

The presence of only three upper tooth rows in the 
L. fletcheri examined here muy be a result of the high 
proportion of abnormal mouths in the two. spawn 
clumps reared, Only one male was present in. the 
colony of adults from which the spawn was obtained 
and given that larvae of L. rerraereginae and ©. selmini 
teared under identical conditions did not show the same 


phenomenon, it is possible that the problem is a genetic 
one, Ridges lacking in teeth were apparent in the 
L. fletcheri tadpoles and it is possible under other 
circumstances that tooth rows form on these. H is 
known that larvac reared in the laboratory tend not to 
have tooth rows that are as well developed as those 
that are collected from the field but the deficiency in 
the rearing methods has not been identified (M. Davics, 
M. J, Tyler & G. F Watson unpubl, data). 

Whilst recognising the anomaly in the tooth row 
formula recorded for L. flercheri here, larval characters 
are consistent with the species groupmys baxed on adult 
morphology. 


Acknowledgments 


Michael J, Tyler and Leanne Seller helped with the 
rearing of tadpoles. Graeme F. Watson and Keith R- 
McDonald provided field companionship and Michael 
J. Tyler collected the L. fletcheri adults and critically 
read the manuscript, Their assistance is appreciated. 
The comments of A. A. Martin and G. F Watson are 
appreciated. This study was supported by an Australian 
Research Grants Scheme grant to the author and 
M. J. Tyler. 


References 


CHAPMAN, J. E. (1987) Rearing and maintenance of the 
Australian. dituran Linwtodynagres tesmaniensis under 
Jaboratory conditions Aiiiia Jechnolagy 3&(3). 175-182. 


Gosner. K, L. (1960) A simplified table for staging anuran 
embryos and larvae with notes on identification. 
Herpersogica tb, 183-190. 


Manin, A. A. (1965) ‘Tadpoles of the Melbourne area, Fier, 
Nat. 82(5), 139-159. 


Rowers, J. BD. & Seymour, BR. §. (1989) Non-loamy exe 
masses in Limaedvadstes rasinantensis (Anuca; 
Myobatrachidue) [rom South Austealla, Copeia (1989), 
ANR-492 


Tyicr, M. J, (1962) On the preservation of anuran tadpoles, 
Aust. J. Sei, 25, 222. 

(1992) “Bneyelopedia of Australian animals, Frogs.” 
(Angus & Robertson, Pymble}. 

Croon, GA, & Davies. M. (1983) Reproductive 
biology of the frogs of the Magela Creek System. Northern 
Territory. Rec, S. Aust. Mas, W818), 415-441) 

es, Marrin, A, A, & Davies, M. (1979) Biology and 
systematics of a new hinnodynastine gents (Anuta: 
Leptodactylidac) from north-western Australia. Aast, J. 
Zaol, 27, ¥35-150. 

Warsom, GE & Manin, A.A. (1973) Life history. hirval 
morphology und relationships of Australian leptadacty lid 
frogs. Trans. BR. Soe, §. Aust 9711}. 33-45, 


A NEW SPECIES OF TRICHOSTRONGYLOID NEMATODE, 
ODILIA BAINAE, FROM A NATIVE RODENT, RATTUS FUSCIPES 
(WATERHOUSE) 


BY I. BEVERIDGE* & M. C. DURETTE-DESSET} 


Summary 


Odilia bainae sp. nov. is described from the duodenum of Rattus fuscipes (Waterhouse) from 
Blackwood, Victoria. The new species differs in having shorter spicules than any congener 
(0.26-0.28 mm), in having a symmetrical bursa and in the number of body ridges. 

KEY WORDS: Nematoda, Trichostrongyloidea, Odilia, new species, Rattus. 


Transactions of the Raval Society of S. Aust. (1992), H6(d), 123 128 


A NEW SPECIES OF TRICHOSTRONGYLOID NEMATODE, ODILIA BAINAE, FROM 
A NATIVE RODENT, RATTUS FUSCIPES (WATERHOUSE) 


by 1. BEVERIDGE* & M.-C. DURETTE-DESSETt 


Summary 


Beveripor. |, & Durerre-Desser, M.-C. (1992) A new species of trichostrongyloid nemuiude, Odilla bainae, 
from # native rodent, Rattus fuscipes (Waterhouse), Tis, RK. Soe. 8, Aust, MO(4), 123-124, 30 November, 1992, 

Crile bainge sp. now. is described from the duodenum of Rutius.fiscizes (Waterhouse) from Bhickwood. Victoria. 
The new specics differs in having shorter spicules than any congener (0.26-0,28 mm), in having a synimetrical 


bursa oid in the number of body ridges. 


Key Woros: Nematoda, Trichastrangyhomea, Qdilia, new species, Rarnis, 


Tntroduction 


Very few studies have been undertaken lo ascertain 
the parasite fauna of the various species of the genus 
Rattay endemic in Australia (Mackerras 1958). The 
fost extensive survey to date on the parasites of Rarus 
Juscipes in Victoria (Obendorf 1979) revealed several 
undescribed species of nematodes including twa which 
were described. A third species, referred to ly 
Obendorf (1979) as ‘'‘Longisiriata sp. (undescribed)” 
was found ata single locality (Blackwood) in Victoria. 
In this paper, we describe and name the new species 
as a precursor to Studies on its ultrastructure and life: 
history. 


Materials and Methods 


Nematodes collected from the dacxlenum of naturally 
infected Rattus fraveipey |rom Blackwood, Victoria 
(37°29'S, 144°19’ B) were washed in 0.9% saline and 
fixed five in either hot 70% ethanol or 2.5% 
glutaraldchyde in phosphate buffer at 4°C, Additional 
specimens were obtained from Jaboratory-reared Rarrus 
fuseipes infected either orally or subcutaneously with 
the third-stage Jarva of the nematode and from 
laboratory rats (Raftus norvegicus) infected via the 
same routes. Specimens fixed in ethanol were cleared 
in lactophenol and examined using Nomurski 
interference microscopy, Drawings and measurements 
were made using a Urawing tube attached to an 
Olympus. BH microscope. Transverse sections of the 
body of male and female nematodes were prepared 
using a cataract sculpel and were mounted in 
lactophenol for examination, An apical view of the 
cephalic extremity was prepared by the same means. 
Morphological termy for the body ridges and bursal 


“ Department of Veterinary Science. 
Melbourne, Parkville, Vic. 3052, 

+ Biologie parasitairc, Protistologie et Hélminthalogic. 
Museum national d'Histoire paturcite, 61 rue Button, 75CIS 
Paris, France, 


University of 


rays follow Durette-Desset (1985). Ridges dorsal to the 
axis-of orientation of the synlophe Were numbered from 
left to right, 1, 2,3... ete, while ridges ventral to 
the axis were numbered from left to right 1', 2!) 3! 
... &tt. Measurements were made on five male and 
five female specimens and are presented, in 
millimetres, as the range followed by the mean in 
parentheses. Specimens fixed in glutaraldehyde were 
embedded in resin, sectioned ata thickness of 1 wm, 
stained with toluidine blue and cxamined under the 
light microscope. 

Type specimens and specimens from experimental 
infections have been deposited in the collections of the 
South Australian Museum (SAM), Adelaide: 


Odilia bainae sp. nov, 
FIGS 1-18 


Longisiriaty 3p. (undeserthed) of Ohendort (9794) 


Types; Holotype male, from duodenum wf Rattus 
Juseipes (Waterhouse), Blackwood, Victoria, 15.%,1991, 
SAM V4l181; allorype Jemale, SAM HC22890; 
paratypes, Zocor, 109 9, SAM HC22879, 22843, 


Material examined; From #.  fiseipes (natural 
infection): types: (experimental infections): 260° 0, 
290 (SAM HC22881). From &. norvegicus 
(experimental infections); orc’, 179 9, (SAM 
HC22875). 


Description: Small, sinistrally-coiled nematodes, red 
in colour when live; prominent cephalic vesicle 
present, syninetrical in shape; buccal capsule vestigial, 
teeth absent, mouth opening sub-triangular, surrounded 
by six tiny labial papillae; four double sub-median 
papillac and paired amphids present, external to labial 
papillae, dorsal vesophageal gland small but distinct 
in apical Views Of head; oesophagus clavifornm; nerve 
fing in nid-oesophageal region: deirids dome-shaped, 


|, BEVERIDGE & M.-C, DURETTE-DESSET 


NEW NEMATODE FROM A RODENT 125 


if region ul excretory pore. Synlaphe composed ot 17 
longitudinal ridges in ond-body repion: axis of 
orientation (Fig 13) from right-ventral field to lefi- 
dorsal fie gt approximately 40" to horizontal: 
diminutive carene, or cuticular dilation, present on left 
dort! aspect between ridges 2° and 5 (Figs 14, 17); 
nine ridges in dorsal field, ridges | to 3 small’, ridges 
6 and 7 larger, ridges 8 and 9 small; cight ridges in 
ventral Held, ridges + and S larger than wither | to 3 
or @ to By some Variation in relative size of ridges occurs 
along body of nematode. with ridges 11) and 3(2°) 
diminishing in size in mid-body region (for example, 
compare Figs 13 and '4); most ridges arise immediately 
pestcrior to cephalic vesicle, single ridge on left side 
arises ut level of nerve ring; single ridge on right sade 
atises Postenir to excretnry pore: number and 
orientation uf ridges variable in posterior extremity of 
body. 

Male. Length 3.5-4.1 (3.8), iiaxinium widih 009-0.10 
(0.09). cephalic vesicle (04-005 (0.05) long: 
oesophagus (),3)-0,36 (0.34) long: nerve ring 0.16 from 
anterior extremity; exeretory pore 0 23-0.24 0.23) 
from anterior extremity; deirids 0.23-0.25 (0.24) from 
umtenior extremity: spicules 0.26-0.28 (0.27) long: 
2ubernaculum lightly sclerotised, visible only: in few 
specimens. 0.05-4)07 (0.061 lang. Synlephe: ridges 
branch and snustomese irregularly in posierior region 
of beidy: up to 19 ridges at level of spicules: ridges 
reduced bul relatively uniform in size, most oriental 
perpenchcular to body. synluphe exrentarion difficult 
in discern. Bursa symmetrical, dorsal lobe reduced. 
Drtsal my symmetrical, divided neat origin, lerminal 
subdivisions short, symmetrical; rays 8 wrising with 
dorsal trunk, papiliac 8 close to margin ot bursa; rays 
4. 5, A grouped yether: ray 6 slender, sharply 
recurved near extremity; ray 3 robust: ray. 4 robust, 
almost acuminate; says 2 and 3 slender, reaching 
marain of bursa, Genital cone extremely: prominent, 
lightly: sclerntised; yeniral lip conical with single apical 
pupil, dorsal lobe with paired papillace 7. Spicules 
elungaie lute, (riquetrous in transverse Section; 
similiry tips Wilh slightly: expanded flange of clear 
spicular mitenal in dorso-vental view: gubernaculum 
present: very lightly scleratised. not visible in all 
specimens, 


Female. Length 4.3-4,9 (4.6), maximum width 
0,10-0.13 (1.11): cephalic vesiele 0.04-0,06 (0.05) long: 
oesophagus 033-044 (0,38); nerve tag 0.19 trom 
anterior extremity: excretory pore 0,23-0,27 (0,25) 
from amterior extremity; deirids 0.25-0,27 (0.26) front 
antenor extremity; tail 003-007 (0.05); vulva tw 
posteriorend 0,09-0.17 (0.12). egg 0,07-0,08 (0.08) by 
0.04-0.05 (0.05). Monodelphic; infundibulum short: 
cee thin-shelled, ellipsoidal, Sytilophe: Ridges 
interrupted in posterior part of body, disappear at level 
of vulva; up to 19 ridges present ar level of 
inlundibulum. ridges reduced in size and uniform ww 
size, Most oriented perpendicular to body: onentalion 
at synlophe difficule to discern. Tail short, conical: 
vulva close ta anus. 


Discussion 


The species described above clearly belongs to the 
Nipposirongylinae Durette-Desset, 1971 in possessing 
a synlophe onemed between 45 and 67° from the 
Saginal axis and 4 latero-median pratient in ridge size, 
In possessing a Carene, or swelling on the left dorsal 
aspect of the body, with a moderately hypertrophied 
left lateral ridge and an obvious Size ditterence between 
the eft lateral rxlge and the left dorsal ridges, the 
hemitode belongs t onc of series of related genera, 
Neohelizginanelle Darette-Dessei, (971, Curnlinensis 
Travassos. 1937, Cilia Durette-Desset.. 1973 -and 
Nippostrongylus Lane, 1923 (see Durette-Desset 1983), 
The possession of a dorsal ray divided close to its origin 
and a common engin for the dorsal and externo-dorsal 
rays exclude this species from Nevheligmonella and 
CarNinensis, while (he symmetry of its bursa, and in 
Particular rays 6, exclude Jit from Nippostrongylus, 
Genera erected since the publication of the key of 
Duretie-Desset (1983), Malaisirongylus Ow Yang. 
Duretie-Desset & Ohbayashi., 1983. Rarrustmvigvlus 
Ow Yang. Durette-Desset & Ohbayashi, 1983, und 
Sabanema Gw Yang, Durette-Desset & Ohbayashi, 
1983, all differ from the species described above in 
lavking a hypertrophied lett lateral ridge (sec Ow ‘Yanp 
er ai, 1983), The species described abuve therefore 
belongs ia Onifia 


nN 


Figs 1-12 Ofilia bainae sp.nov, from Ratris fuscipes,. 1. cephulic extremity, showing cephalic vesicle: 2 apical wew of 
anterior extremity, 3, anterior end, right side, showing original ridge (arrow) at level of Ockophago-inlestinal junction: 
4. dntetior end, left side. showing origin of ridge (arrow) at mid-ocesuphugeal level) 5, posterior end pf mule, ventral 
Siew, stewing irresuluritics (9 yidges, , dorsal lobe of hursa, dorsal view, 7, lateral view of bursa showing prominent 
genital Gone; 8, genial cone, ventral View: 9. genital cone, laterat view, 10, spicule tips, ventral view, UH. sptuule tip, 


lateral »iew. 12. tail of female. 


Figs 5-9 follow ray numbering system as deseribed by Dureile-Desser (1945), 0.7 represent the pilpillae of the genital 


cond in the Sate systeu. 


Legend’ a amphid. d. dermid, &. excretory pore: |, Jabal papilla: p, sabmedian papillu, 
Seale lites figs t. 2, & 8-1, O01 jem; fies 4-4) 7 12. Ob nnt 


I BEVERIDGRE & M.-C. DURETTE-DESSET 


126 


NEW NEMATODE FROM A RODENT 127 


The venus Qvilia was established to contain seven 
species of trichostrongyloid nematodes beloneing to 
the Nippestrongylinac. parasitic i@ Australian murid 
rotlonts, The genus, Is clearly related 16 
Nippostroitevies, which is cosmopolitan in nidents, 
panticulariy Rattus spp. (see Durene-Desset (970), 
Nevhelignonella i Alrican morid dents and 
Cardivensis in holarctic arviculid and gethillid 
rodents, hut is distinguished primarily by the 
charactecistic lorm) of the dorsal ray (Durene- Desset 
1971) which is deeply divided and arises af the origin 
of the externodorsal rays. The new species desenbed 
here was inually identified as‘ Lonpistriate sp.. 
undescribed” (sexsi: Mawson 1961) by Obendorf 41979). 
The species desenbed above can be differentiated from 
all congeners by ity extremely short spicules 
(0 26-0,28 rion). lt differs from O. jrackerrasae, O 
mawsanae, O. brachybursa, O. polyrhabdote and 0 
crranielae in possessing a symmetrical caudal bursa, 
The remaining species, O. melomnwy and O. wromyys 
have it symmetrical burss. The new species also differs 
Irom all congeners in which the syploplie bis been 
described in the number of ridges in Lhe mid-body 
region, I7 in the new species compared with 14 in O 
bruchybursa, WS in O. emanuelue, 16 in O. 
mackerrasue. 15 in O. metonnvas, 36 if O. polyrhabdote 
and 21 in Oo maweinge, although G. maiwsende has 
(7 ridges in the anterior part of the body (Duretic- 
Desset 1969), The number of ndges in OL nromyes is 
nol known, Tous the muterial described confirms the 
ohservation mt Obendort (1979) that it is a new species. 
which we here name O. bainae, aller Dr Odile Bain. 
in Whose honour (he genus was erecta. 

The presence of an additional spectes of Odiia in 
un cndemic species of Rarruy is of interest in View of 
current hypotheses un the evolutionary relationships 
of the genus. Mawson (1961) observed that the 
tichostrongyloid genera present in endemic niurine 
rodents, thut is the species of Rattus, belong primarily 
10 Ninpeytronevies and slustroheligmonema Mawson, 
1961, although Austroheligmonema is now Tegarded as 
a synonym Of Nippastrengvus (sec Durene-Desset 
1971), Those nematodes present in the “Old endemic” 
rodents belonging to the sub-family Hydromyinae were 
mainly species placed in the gener: Longisrriata 
Scholz, 26 and Heligmonoides Baylis, 928, alciough 
all of them are. now included in a single genus Qailia, 
In & pe-caumination of the merphulogy af Australian 
species by Durctte-Desser (1969), a trond in syalophe 


morphology was observed from species with 4 carene, 
hypertrophied tefl, literal ridges and a size gradient 
in ridges from right to left towards synlophes sueh a5 
that found in Q. pelyrhabeete in which the number 
of ridges was increased, but their sizes diminished and 
the distinctive orientation was lost. Because thé former 
lype of synlophe occurs in genera such as 
Nippostrongylus which pecur in south-east Asia, 
Durette-Desset (1971, 1985) considered these findings 
consistent with the hypothesis that the hydramyine 
rents reached Australia with nematodes resembl ite 
Nippostrong vies, and that the genus Oellia evolved in 
isolating in the Hydromyinae. The more revent arrival 
of species of Rairus in Australia about one million yeurs 
aga (Watts & Astin 1981) probably imetroduced ur re- 
intraduced Nippastrangyius, and lead [a the 
development of the two. endemic: species. N- fypicws 
and MV. mewnys in Ruts fuscipes, According 00 this 
hupothesis. species of Ouilia present an endemic 
Australian Raltus spp onepresent transfers from 
hydromyine rodents. 

With respect in synlophe morphology, the new 
species fils within the transtiian series envisaged by 
Durette-Desset (1969). The number of syntophe ridges 
(17) ts grealer than that expected in the supposed 
ancestral slate (14) and although a size gradicnt in the 
tidges teniains, the carene is not prominent. Two 
gpecies of Odile secur in Rativs species in addition 
10 O bainee, these being OL enaruelae in R. conerac 
und Q palvhabdare in R. fescipes isyn R, assimilis). 
Tn cach instance. the species of Rartus involved is 
broudly sympatric with hydiomyine rodents, principally 
Melonzys spp. {Watts & Aslin 1981) which could have 
acted as danors in the transfer to Rattus spp At 
Blackwood, the only known locality for Q Aainue, nu 
other hydrumyine rodents other than the water rit, 
Adrian chrvsoguster, occur. This suggests that O. 
bainde js exclustvely a parasite of Ranus spp. and that 
Iranster of the species of A. Aescynes occurred some 
lime in the past, either when hydmmyine rodents 
occurred in the area, or prior to the extension of R, 
fusgipes into this region. However, of the Australian 
hydromyine rodents. only che pasasites of Myyronpys, 
Uremys and Meforws have thus far been studied, and 
for thany species and genera, there are as yet no records 
(see Mackerras 1958), Therefore any conclusions un 
the host of geographic distributions of individual 
nematode species within them need tyr be treated with 
sume canton. 


OO - ee ee ———eeeeeSSSSSESSeSeSFSseMF 


Bigs 1318. Ovilia botice sp.nov, from Rares fuseipes. Transverse sections ai different levels showing synlaphe. 19, mate, 
DAU mm fom anterior end with arrow indicating axis of orientation Of synlophe, 14, amale. mid-body pegion, 2,2 min 


Iron) interior end: 1§, mate, secrion at level of spicules, O 


26 the trary pusterjur end, 16, fenrale, 0.40 rim (rom anterior 


em); 17, female, inid-body region. | rant from anterior end: Ix, feraale, 0.20 from posterior end Seale bine. Oth mm. 


128 I. BEVERIDGE & M.-C. DURETTE-DESSET 


Acknowledgments 


We wish to thank Christine Andersen for expert 


Rats were trapped under a permit from the Victorian 


technical assistance and Dr D, M. Spratt for comments Department of Conservation and Environment 
on the manuscript. This work was supported (RP 91-095). 
financially by the Australian Research Council. 

References 


Durerrt-Disser, M.-C. (1969) Les systémes d'arctes 
cuticulaires chez les nematodes heligmosomes parasites de 
Murides australiens. Ann. parasitol. haem. comp. 44, 
733-747. 

970) Le genre Nippestrongylus Lane, 1923, 
(Neématode-Heligmousomatide). bid. 45, 815-821, 

_s (1971), Essar de classification des nématodes 
heligmosomes. Correlations avec la paléobiogevgraphie des 
hates. Mém. Mus. rain. Hist. nat., Paris sér. A. Zoologie, 
69, 1-126. 

(1983) CIH Keys to the Nematode Parasites of 
Vertebrates. No. 10. Keys to genera of the superfamily 
Trichostrongyloidea. (Commonwealth Agricultural 
Bureaux, Farnham Royal, England). 

(1985) Trichostrongyloid nematodes and their 
vertebrate hosts: reconstruction of the phylogeny of a 
parasitic group, Adven. Parasitol, 24, 239-306. 


Mawson, P. M. (1961) Trichostrongyles from rodents in 
Queensland with comments on the genus Lungistriata 
(Nematoda: Heligmosomatidac). Aust. J. Zool, 9, 791-826, 


Mackerras, M. J, (1958) Catalogue of Australian mammals 
and their recorded internal parasites. I]. Eutheria. Prac, 
Linn, Soc. N.S.Wales, 83, 101-60. 


Owenvorr, D, L. (1979) The helminth parasites of Rattus 
fuscipes (Waterhouse) trom Victoria, including description 
of two new nematode species. Aust, J. Zoal. 27, 867-879. 

Ow Yang, C. K., DureTre-DEssetT, M.-C. & OHBAYASH]. 
M. (1983) Sur les nematodes. parasites de rongeurs de 
Malaisie. Il. Les Trichostrangyloidea. Ann. parasitol. ham, 
comp. 58, 467-492. 

Waris. C. H. 8, & Astin, H. J. (1981) “The Rodents of 
Australia” (Angus & Robertson, Australia). 


A DISSECTION METHOD FOR DETERMINING THE GUT CONTENTS OF 
CALANOID COPEPODS 


BY JOHN D. GREEN 


Summary 


Tramactions af the Rival Suery of & Anse. (1992), Ubed). 129-192 


BRIEF COMMUNICATION 


A DISSECTION METHOD FOR DETERMINING THE GUT CONTENTS OF 
CALANOID COPEPODS 


‘The examination und recording ef gut contents has proved 
lo be 4 useful aid in the study of the diets of zooplankton, 
The method cangol give 2 complete picure of the diet of u 
Particular species as Some food ikem acc more delicate than 
others und are more readily broken during mastication anil 
dissolved by uagestive cnzymes. However, many algae and 
animals in the diets of copepods remain sufficiently 
undamaged, or have ailequate idowlifiable parts thar sre 
resistant 4) enzymatic breakdown, to allow 3 reasonably good 
usyessiment of diet fram pot contents analysis '244. It is 
preferable to base the analysiy on the contents of the fore-put 
(Fig. 2), it which mach of the ingested material has boon 
less allected by enzymatic action and is less compacted, than 
On the compacted and well uagested bolus in the Dind-gut, 
Or the faecal pellers. 

His nota simple matier, however, to disseot out the entire 
guCcontents of a oypepud, largely because of the :tanipulative 
delicacy required, The contents of the foregul can be 
particularly difficult ta extract in their entivety becuse of their 
diffusences. Those who have used 4 dissection method may 
thus choose 69 reniove only the hind gut balus®, The methods 
Most used for oxaminang eut contents avoid direct dissection. 
In the squash techmique, the gut contents are extracted fram 
either lwe or pieserved specimens by pressing down on & 
coverslip over the animal!*°. This method has the advaniage 
of bath releasing the material in the gut and dispersing it so 
that individual terns may be identified and counted. Another 
method is to render the whole aninzal transparent by clearing 
wt lacie acid’, or in cuparal or canada balsam afier passing 
ii through an aloghol series®. The darwhack here is that the 
gut contents are nol dispersed, and even when the put heluges 
cen be clearly seen the indivatad! food nems are musily 
Wifficult ta identify positively or us vount. One ‘wey of 
«wercoming such difficulties is ta place the specimen under 
i cover slip und erode away most of the tissues with weak 
sodium hypochlorite’. The hypochiorite is then flushed away 
before the gul contents are oxidised, and iris usitally possible 
to identify many of the individual food items by pently moving 
the cover slip, which partly redistribute the gut coments, The 
method has been used successfully is Australia ts determine 
maximum gat food-particle sizes af lhe copepods Calenwecia 
(wool, Boeckella mirada and & trierticutany in Wallerewaneg 
Reservoir*. 


During a study of carnivory by three large omnivorous 
calanuid copepods (Boeckella major, B. asemdoctelie and 
Hemibuneckella searii} trom temporary ponds an the upper 
River Murray Qowdplain, we tried ail the above nethads of 
gul contents analyses, None of them proved entirely 
satisfactory, particularly for revealing the remains of amiunvals 
in ihe gil contents. This appesred to be m@pinly due to ihe 
large size and thick bodies of the copepods, and because we 
had available only specimens preserved in 4% formalin and 
70% alcohol, The squash method appeared seasonably 
salistactary for small specimens, bu in larger animals (and 
particularty those preserves! in 4% formalin) the gur contems 
were often difficult to observe clearly amongst the mass of 
disrupted exoskelenon and musvle tissue. Clearing in tactic: 


acid was only partly successful. The copepods did mo) ehear 
very well, again apparently because of their large size. 
Whenever the food billises could be seen clearly, animal 
remains (c.g. cuticle, selue) were difficule oF ijipossible to 
recognise as they usually were Crished and compacted within 
the holus The hypochlorite erosion method was also nor 
entirely successful. Even though jHe gut cuntents could be 
partly manipulated, the fact that the gut boluses were not fully 
dispersed mide animal material difficult «a see. As well. it 
was found that bubbles of oxygen produced during (issue 
erosion accumulated within the body and obscured the gut 
contents, and thal caro had to be taken oo ensure that the jul 
contents themselves were nor axidised, 

In order to overcome these difficulties We developed (he 
lowing dissection method, which enwbley the entire gue 
contents of both small and lange copepods io be reyes 
The contents of both fiore- and hind-gutx can be cleanly 
ealraciod without interference fram moat surrouniling tissucs, 
dispersed. ond petrnanently mounted. 

Needlex for dissection are made. from 2 ém lengths ol 
1.3 mavtungsien wire, which js rigid enough uw allow same 
pressure to be applied during dissection, and may be 
sharpened ww a fine peyint. For dissecting large copepods a 
shurp enough point can be produced with a fine dlamond 
whetstone {¢ ©. “Ezelap"). For small copepods itis bemer to 
produce the desired point by erosion, cithear in molten 
NaNO,, heated over a bunscn burner ina crucible! or try 
clectredysis in 10-20% KOH. For electrolysis, he wire is 
Clamped ta cine terminal ofa 6Y alternating current electrical 
circuit (9 pricrscopellumunation transformer is suitable) and 
Wipped inte the KOH"? tn either case, the wire is moved 
m and out-of the fluid, and the depth to which the wire is 
inserted governs whether the resulting point ay ahoOr and stout 
or long and slender The sharpened noedie is then mounted 
ina holder, Satisfactory holders may be made fram pu vices 
(small Fingor-operated drill holders available from model 
shops} that have been lengthened, if necessary, by the addition 
ata section of brass cod (Fig. 1c), The uingsten needle: is 
hent at a slight angle to the axes of the holder (Fig, lami, w 
aid keeping the needle parallel to the slide surface during 
dissection. Jeweller’s forceps, with finely sharpened paints, 
are used for transfesrme copepods, or their pars, 

Dissection can he done in water, but in is casier fa more 
Viscous medium ts used, Polyvinyt alcuhol-luctophenol 
mauntiar |PVA}" is very suitable as it can be used to make 
permanent mounts of the gut contents. Lignin pink muy be 
added Wy the PVA to stain chicit 


tn 


eB —— 


Fig. LA dissecting needJe, consisting of a pin vice holding 
a finely pointed congsten needle (tn). The commercially 
avallabte pm vice has bee extended by the addition of a 
section of brass rod fe), Seale bar — Lom. 


Fig. 2. Dissevtinn of put contents. Orientaliun of specimens 
1% that for a tight handed person, A, Lateral view of 
Boeckella mapor shoring the fore gut (Fgh) and hind gut 
(hgb) boluses, The first.and second dissection cuttings are: 
shown by dashed lines J and 2, respectively, B. Orientation 
af the copepod and dissection technique for removing the 
antero-ventral portion of the prosome, The right needle is 
placed with its point between the maxilliped and first 
swimming lezs and pressed firmly down and held against 
the slide. Back and forth movements of the left needle ther 
sever the antero-ventral surface, which is pultcd away to 
the left. C. The body ready for cransfer ta.the second dep 
of PVA. The optional cut-line for rentoving the remaining 
veniral suflace is shown by 2 dashed line. DL Extraction 
pf the fore gut contents, The body is held with the left needle 
while the fore gut bolus is gently pulled cut with the right 
needle. Scale hars = t mm. 


Using the forceps, two drops of PWA are placed bna slide. 
The copepod j¢ picked up with the forceps and placed im one 
drop. in which most of Ihe disseeilon (i.e, removal pf urosome, 
antero-ventral surface and mouthparts) id done. The body is 
then transferred fo (ie second drop for the reniwya), teasing- 
out and mounting of the gul contents, 


Dissection is done using a steren dissecting microxcope al 
a4 magnification of ca. 30-40%. The copepod is orientated 
with its ventral surface partially inclined'to the left and away 
trom the dissector, and, Tora right-handed person, with its 
anterior end to the left (Fig- 2a). Firsdy the urosome atc 
terminal segment of the prosyame are wemoyed by cuiting along 
dashed line | (Fig. 2a), and then the anteto-ventral surface 
of the cephalusyme plus mouthparts, hy cutting along dashed 
line 2 (Pig. 2a). If desired, ihe swirnating linvbs (PI-P4) may 
alyu be removed (by cutting ulony dashed line 3, Fig. 2c), 
This is not absolutely necessary but may be useful if the 
ovaries are well developed. Swallen ovarioles make removal 
of the gut contents difficult, and remoyal of the swimming 
fimbs and remaining ventral surface usually results in the 
coneonyitant removal of much of the ovary Clase. 

The first-cut is made with the anima! orientated ay shown 
in Fig. 29. The body is held with the left needle and the cut 
made wih the maht needle by pressing down firovly along 
line 1, using 4 forward and backward Sawing action of the 
right needle if necessary. For the second cut, the animal is 
roorientated to the position shown in Fig. 2b, The animal is 
held with the left needle (near the base of the first antennae 
is a Suitable point) and the right needle tirmly pressed down 
over the body (Fiy. 2b), with the point of the needle between 
the maxillipeds and first pair of swimtming limbs. While the 
right needle is pressed firmly down against the slide, the 
antero- ventral surface and mouthparts are severed by buck 
and forth cutting movements by. the tp of the left needle (Fig. 
2b), The provediire usually pushes the fore-gut bolus slightly 
dorso-pesteriorly towards the rear-of the fore gut, and very 
occasionally moy result in the rear-gut bolus being extruded, 
if this happens, the rear-gut bolus can be retrieved with the 
foreeps and transferred to the second drop of PVA. The body 
should now look as shawn in Fig. 2c. If necessary, the 
swimming legs may now be removed hy cutting along line 
3 (Fig. 2c), pressing down on the body with the right needle, 

Lising the forceps, the body may now be transferred to Ihe 
second drop of PVA, atid held by the left needle with ventral 
ide uppermost and anterior end facing right (Fig, 2d), The 
fore-gut conlents we then carefully scraped out with the right 
needle (Fig, 2d), the body rotated 180°, and the reat-gut bolus 
removed in a similar manner. Finally, the body is removed 
with the forceps and discarded, 

The tood boluses may now be carefully teaged apart with 
both needles and a amall coverslip added. A 10 mm or smaller 
diameter coverslip ib beter than the standard 24 min size, 
lw reduce the area that has to be searched during Migroscope 
exumination, The gut contents can be fully dispersed by the 
application of light pressure, and perhaps also small side-to- 
side movements, to the top of the cover slip with a needle 
ur the forceps. 

The gut contents uf both small (e.g, Bverkella symmetrica, 
body. length ca. 1-1.5 mm) and large (e.g. B major, body 
length ea. 3-5 mim) freshwater calanoid copepods can be ewsily 
extracted using this disseetion method, Because both the gut 
boluses can be extmeted and teused apart we found thai the 


i 


Fig. 3, Photornicrogfaphs of animal remains and algae in dissected put contents. of Boeckelfa major A, Daphnia carinate: 
2, post-abdomen:,. b, inandibles: ¢, enricle and thoracic limbs, B. a, calanoid copepodite limbs: b. calanuid copepodinc 
niandible; ¢, calunoid nauplius; U, Téstuelineiia patina C. Reratella procurva, wophus armed. D Algae; a, Stawrastrem 


sp; b, indet, diatom 


131 


132 


method reveals animal remains in the gut contents better than 
the whole-animal squash and clearing techniques mentioned 
above. It is possible to pick out both very small animal remains 
(e.g. rotifer trophi, Fig, 3), and the diaphanous cuticular 
remnants and setae of cladocerans and copepods (Fig, 3). The 
visibility of cuticular fragments is enhanced by the lignin pink 
stain in the PVA, and also by the use of Nomarski interference 
optics. Algae, fungi, detritus and inorganic material in the 
guts are also clearly visible (Fig, 3). It is possible to make 
quantitative counts of the gut contents, 

Animals preserved in 70% alcohol proved to be easier to 
dissect than those in 4% formalin, Alcohol preservation results 
in the dissolution of much of the muscle tissue and the 
softening of the exoskeleton. The body is thus easier to sever 
and to manipulate than when preserved in formalin, and there 
is less tissue “rubbish” in the final gut contents preparation. 


‘Fryer, G. (1957) J. Anim. Ecol. 26, 263-268. 

2Gliwiez, Z. M. (1969) Ekol, Polska, Ser, A, 25, 663-708. 
Infante, A. (1978a) Arch, Hydrobiol. 82, 347-358. 
‘Kobayashi, T. (199]) Aust. J. Mar. Freshwater Res. 42, 
399-408. 

SHart, R. C. (1987) Arch. Hydrobiol. U1, 67-82. 
°Clarke, N. V. (1978) Freshwat, Biol, 8, 321-326. 

"Yen, J. (1985) Limnol. Oceanogr. 30, 577-597. 

SGreen, J. D, (1976) Arch. Hydrobiol, suppl. 50, 313-400. 
“Infante, A. (1978b) Trans. Amer. Micros. Soc. 97, 256-258. 


The dissected limbs and other body parts remaining in the 
first drop of PVA can be put to good use. The mouthparts 
can be dissected off the remmant antero-ventral surface more 
readily than they can be from the whole animal, To do this, 
the apodemes at the bases of the mouthparts are anchored 
solidly against the slide with the left needle, while the 
mouthparts are easily dissected off with the right needle. 
Moreover, egg sacs removed with the urosome can be used 
for clutch-size determinations and measurements of egg size. 


We are gratetul to Dr I.A.E. Bayly tor his helpful comments 
on the manuscript and for bringing two recondite papers to 
our attention, This work was done while JDG was on 
sabbatical leave at The Murray-Darling Freshwater Research 
Centre. Albury, N.S.W.. and the use of facilities there is 
gratefully acknowledged. 


“Cannon, H. G, (1941) J. Roy. Microscopical Soc. 61, 58-59. 

"Brady, J. (1965) Bull. World Hith, Organisation 32, 143-144. 

Frey, D. G. (1986) Cladocera analysis. Pp. 667-692 Jn 
B. E. Berglund (Ed) “Handbook of Holocene Palaeoecology 
and Palaeohydrology”. (Wiley, London). 

Chapman, M, A. & Lewis, M. H. (1976) An introduction 
to the freshwater Crustacea of New Zealand, with a chapter 
on the Arachnida by V. H. Stout. (Collins, Auckland). 


JOHN D. GREEN, Dept of Biological Science, University of Waikato, Private Bag 3105, Hamilton, New Zealand. 
RUSSELL J. SHIEL, Murray-Darling Freshwater Research Centre, P.O. Box 921, Albury N.S.W, 2640, 


“ROLLERS” AND “CARRIERS”: FIELD OBSERVATIONS OF CARRION 
REMOVAL BY TROGID BEETLES (OMORGUS STRZELECKENSIS) IN 
ARID NORTH-EASTERN SOUTH AUSTRALIA 


BY ANDREW J. BOULTON 


Summary 


freanvaetions af the Reval Societe of 5. Aas, (992), UM6idi, W354, 


BRIEF COMMUNICATION 


“ROLLERS” AND “CARRIERS: FIELD OBSERVATIONS OF CARRION REMOVAL BY 
TROGID BEETLES (OMORGUS STRZELECKENSIS) IN ARID NORTH-EASTERN 
SOUTH AUSTRALIA 


Despite substantial advarices in the fawonomy bf Australian 
beetles (Coleoptera)! litle is known about the biology and 
ecolagy of most of our species. especially those considencd 
a linle economic significance. Matthews? suggested thar 
miluralins Cun make valuable conpriburions by investigwting 
And describing aspects of (he qulural history of becthes. Often 
such obscrvutiens aro Opportunistic and represent situations 
difficull to replicate In the laborlory, With 3 minum af 
equipment, imany wsetol aspeets of Dchaviour can be now 
and simple experiments used to clarity observations. 

Duting 4 feceat lininologicul Held trip to the Coongie Lakes 
area. an appoctunity arase to study groups-of carrion beetles 
(Trogidae) transporting fhecal pellets and regurgitated cgesta 
along 4 sahd-dune an [he western share of Lake Guolangirie 
(26°93'S i40%N'E), 12 km NW bf Innomincka, In the 
(nnamurcks Reginwl Reserve. The lake filly as wu result of 
overbank Hows inthe North-West Branch of Cooper Creek 
This lake, and many omncrs nearby, owe their irregular shapes 
lo the extensive dune sysicis running approximately nogth- 
south, and vegelaled by an open wall shrubland aF Acacia 
Igulata A, Quinn, ex Benth, in the wider swales, « hummock 
grassland dothinated by Thodia baredowii F. Pritzel along 
the muyor dune systems and sand-hill cane-prass (Zivochloa 
paraioxva [R, Brt) un tie mobile dune crests? 

The beetles. were wemilied as Dworeuy srcelectensis 
(Blackburn, 1895+) described from specimens collected near 
Lake Callubonne and Strvelecki Oreck, The species ja 
widespread throughout mainlaid Australia, bevurring in law 
Tuinfall arews of all States except Victoria’, tt 1s distinguished 
trom congeners by the combination Of shape ofthe elyiral 
costa and the single large median uwith an the fore tibia, 
Sctioltz? noted thut spevimeny hud pever been collected in 
Janiary, April. June or December — ay observations were 
madé in December {early summer) 

Life history data on the Aastnilasian Trugidue are few 
although some 33 species have been described, af which (wa 
wre introduced? The only published record af a particular 
feeding habit appewrs to be tial by Leefmans® whn reported 
OL (Trax) cosraney livitg and feeding on bal euntno jn caves 
in the Celebes (Sulawesi). In Africa and America, trogid 
adults and larvae are facultative necrophages and generally 
Appear late in the succession of invasions of carcssyes in the 
arid zones af these continems, 

Tiree broad sjrategies for coprophagy and necrophagy in 
arid zones are secugnised’. The first is tu tap the latent 
moisture of the purticle and to breed as quickly a5 poysible 
before it dries (eg. dipterun fics, especially cabliphorids). 
Alternatively. the material may be comminuted and buried 
ta Conserve its Mnisture as ts the case with dung-beetlos 
(Scarabucjaae), In arid anets the lack of reliable tain leaves 
some species of dung-beeiles undeterred because they can 
hury desivested faecal pellets that rehydrate frum soi 
moisture’, Finally, some groups (lermites, tenehriontds, 
rragids) #re able op eal dry faeces ind mummified carrion, 
Gnnirgus (Thos) spp. can complete their life cycle an nly 
woul clippings ar even discarded woal clothe, 


Omorgus spp. in the Kalahar: Des¢el ace morphologically, 
behaviourally and phystoloziculy udapied w survive long 
periods of aridity interspersed with brief favourable periods 
when feeding and cupulation occur’, Adults and larvac 
quiesce underadverse ambient conditions, renewing activity 
within hours af ameliomiion, Inunsture stages develop rapidly 
(3) weeks) whereas the adult lifespan ys long enough to 
allow overlapping generations The availability of mummified 
curcusses of lange natclope for several years allows populations 
10 huikl up below the remains withnut the risks Involved in 
dispersal to seek food or mates” 

The following observations were made ever two days 
HD -AiL1991), Tracks ocansisting of numerous small 
depressions in a band 3-4 cm wide were noticed runniig east- 
west alemg a dune face soon afer stinrise (0630 hours) on 
(043,51, They Were made by Iwo beetles carryime a tapered 
dingn faecal pellet measuring approximately Jem long and 
} em in diameter (Table 1), The beetles were moving 
westward, und a light easterly treeme was blowing wher 
Jetulled observations connienced at 0645 Ir, Subsequently, 
six uther groups of heetles were discovered carrying or rolling 
egesia fram dhe base of a Jone Coolibah use (Bucwtyntns 
michutheco F von Meuller) growing Int! Away up the dune face, 
These groups, nated at about 0700 hr, when the hreest had 
strengrhened slizhdy anu swung ina northerly. were moving 
southwards and had travelled only several metres. 

The most sinking feature was the two different modes of 
carrion transport adopted by the groups of bectles. They cither 
camed or roiled the cgesta, depending upon its shape, 
[regular pieves of carrion were physically carried upan the 
hacks of the beeiles with the load being spread fairly evenly 
between imdividualy. The pieces of carrion were stabilized 
hy nodules and grooves en the pronvki aud elytra of the 
beelles. Mare cylindrical pioves were rolled ulong. Beetles 
Popped their stout forelegs against the carrion ar iweked their 
proniiom under the edge of the particle, lifling thetr heids 
while pushing forwards with their middle and hind pairs ot 
legs, Although the beetles were always on the trailing edge 
of the rolling piece of carrion, they often appeared to push 
each other forwards and @ was not unusual for & beetle lo 
walk over the backs of others ta ral! the particle. 

Ir is Important to distinguish between these twa types of 
carrion transport by tragid beetles and rhe classification of 
resource relocation that ia frequently applied 10 the majarity 
of dang beetles Dung beotles are sepurated into guilds of 
tonncllors (paracoprids), dwellers (endoaaprids) sind folliers 
(telocoprids) "| ‘Tunnellers dig burns inumediately beluw 
the dung and push pieces of it into the lunnels, Uwellers tive 
and fed within che dung heap itself, and rollers mike balls 
of dung thar they roll away from the food source and other 
beetles before concealing them in the sail. bn the case of 
the tragids that f observed, all the beetles-would be in the 
soller gpd" but exhibit either carrying or Tolling behaviour, 
In dung beetles, the guilds are species specific whereas the 
modes of particle transpxid evident in the (rogids in ny stiKly 
herth pecur in ihe one species, 


Isa 


TABLE 1, Suranary of olvervations muile on sever groups of carrion beetles at Late Goolanyirte. Means (+S6s)-ine for 
all groups Carning egest sourwieeds (group Tix exclieled beewiese it was carrung @ facoal peller west), 


Group No, of Size of Shape Transport Track Mean 
beetles burden (ent mode length (ml pradient (%) 

' 2 3 ¥ 1 Tapered Currier 3 20 
2 4 h«7 Irregular Currier 35 \) 
i 5 b«3 Cylindrical Roller 25 " 
4 3 8 xz Irregular Currier 18 ib 
5 5 5 xi rregular Catmier 4) i} 
6 3 ix 2 Cylindrical feller 42 T 
7 5 9x2 Irregular Currier as t 

Meun 4 bxa 34,33 IL€i7 
SE 0.45 LOO TR 2.95 1.56 


The inclines up which the carrion was. rofled or carried 
were measured by hulding # metre stick horizontally level 
with one end resting. on the Irack and the stk polhting along 
the path taken by the heelles The elevation of the free end 
of the stick measured the rise (in cm) aver 1 om and was 
expressed us a percentage (Table |). The maximum incline 
wp which beetles rolled carrion war (lf whereas particles 
were carried up a slope of 42%. The shape of ihe carnon 
particle appeared to determine tts mode of transport ralher 
than the slape al the incline or the number of beetles involved 
(Table 1) Jn genera), there were more beetles associated with 
larger pieces of carrion but other factors such as particle shape 
wie probably also imponaal, By placing smal! Nags along the 
tracks at 3D of 60 second intervals and measuring (he length 
of the path. 1 way able lo compare the speeds of carrion 
Lranspurt by the two different modes, At lemperatures ot 
26-28°C and traversing similar terrain. five beetles carrying 
epesta (Group 2, Table |) moved faster thana group (3) rotling 
a smaller piece (Table U). “Carriers” averaged (142 crn.s (1 
= WO determinuions, SE = (403) whereas “rollers” moved 
a O28ema! (n = 9, SE =0.02). 

The pathways were nor straight but meandered in a general 
direction, When the egestum became wedged against 4 stick 
arembedded in a hollow, the beerles soxin retraced their stepe 
andelycling around, tried aslighily new bewing, These circles 
never exceeded 30 cm indinmeter and [only found one preve 
of carrion that became ingstricably trapped in a clump of 
Enchyleena tomentosa R. Br. (Group 5), The hillowing day. 
this particle was covered in ams and there was no-sign of the 
carrion beetles nearby or burical below. 

Al times, individual beetles would wander away trom the 
carrion or become dislodged affer a particularly vigorous roll. 
Invariably, they would circle te the downslope side nf the 
carrion and then travel with the wind, moving uphill vin 
they were directly downwind of the particle. Immediately, 
ihe beenle would turn towards the carrion and walk in a straight 
line up io it, even it the particle was out of the direct line 
of sight due to hollows aud sand ripples, | observed this 
behaviour several times, and successful reunions ocurred over 
distances of 2 m. Whe a member of the group wandered 
sway or was dislodged. the rest of the group did not alter 
behaviour und continved moving the egesta. 

Beetles were able to sense carrion from a radius of 10-15 ern 
even if upwind (as “rollers” often were) und moved directly 
uowarda the particle. ‘This was confirmed experimentally using 


both “rollers” vind “cartiers) and was possibly visual, 
Shubeck!'? otserved that carrion beetles in the family 
Silphidae could dewet odours at distances af approximately 
Lin when wr movement was negligible bul at creater distances 
(5-75 m), he concluded that ormemavion fo carrion was due 
&) randon) wandering. 

ff beetles go to such lenyths to wansport carrion, do groups 
exhibit uny possessiveness, perhaps defending cheir particles 
from other conspecifics? Anecdotal evidence from field 
observations indicates that inter and intra-specific competition 
among dung-beetles can be intense, ranging from direct 
vombat When beetles fight aver the possession of dung to 
scramble competition when the beetles’ activity at high 
densities prevents most individuals acquiring sufficient 
resources for breeding" 

What happens if a waridering beetle trying t recover itk 
catrion titds itself downwind of another group's particle and 
homes in on thal, Can a wandering “roller” readily switch 
lo"careying "if an irregular piece of carrion is encountered? 
To cxamine these possibilnics, | planned a series of transfers 
of beetles from one group to unvther, within and ycross modes 
of transport. ‘Touching the beetles caused them to feign deuill 
instintly, becoming immobile and jucking their limbs tightly 
under their body. Thus, it was necessary (0 allow them by 
walk onto a strategically placed teaf and then transfer the beetle 
quickly, placing it just downwind of the carrion in all trials, 

In all transfers. (a = 5), there: was no change in the 
behaviour of the recipient group and | was unable to detect 
any physical antagonism, Newcomers-often crawled over the 
carrion tur several seconds hefore joining their fellows either 
rolling or carrying the particle, 1 returned all! beetles to their 
uriginal posinons at the end of the experiment, where they 
resumed their behaviour, seemingly unaffected by their brit 
jronsfer: Thus, i¢ seem that groups of bectics af this species 
wre nol especially protective of their carrion resources. I 
wauld he interesting Ww add beetles continually to a particle 
lu see if [ntraspecific competition could be induced. Beeiles 
had no apparent difficulty switching modes of transpart bo 
match that of their fellows — in no case. dhe they wry to roll 
a particle that was being curried oc vice versa, Furthermure, 
L never observed a uroup ot beetles switch modes of transport 
im response tou change iit grade or subanratum particle size. 

also found an-eighth group of two dectles rolling 4 
cylindrival pitee of egesta (2 x 3 cm) southwands, Lsaenificed 
these two heetles for identification. amt will three arhers, 


they are lodged in the South Australia Muxcyirn (classifted 
wilh the Trogidae, Dr Ene Mauhews. South Australian 
Museum, pers omen.) 

Could beelles be enjiced away from their carrion by another, 
secmingly palatable piece of egestum, espectally given their 
Apparent lick of possessiveness? When this Was done, groups 
of bectles carried of rolled their particles pust the new picce, 
ever When jt was plaged in the path af ihe grup. However, 
if'4 single beetle became separated trom the group and the 
particle was pliced in is path, the beetle crawled over the 
carrion iit sinila expluralary fashion to that observed earlier 
and then proceeded to enther roll i or a burrmw below: ir. 

The inerning | made these observations tas overcust but 
Sind temperdtures Pose gradually Crom 18°C! at 0700 hr iw 
34°C by WOO he By 0900 hr (26°C), Group I carrying the 
dingo thecal pellet to the top of the dune hud burrowed below 
the pellet leaving it exposed. However, sund blown by the 
wind had ball-buried the pellet and completely obliterated 
the tracks by HPO br. At WIS hr. sant tomperiture reuichedt 
40°C and the other six groups of beetles ceased activity almost 
insiamtuineously. The wind hac Strengthened and air 
temperature was 35°, Ih all cayes, beetles either sheltenid 
below the carrios-or had burriwed into che sunu benesth the 
particle to adepth of 3-5 cr in = 3), Observations of depili 
of burrowing were restricted to groups 2 and 4 (Pearricns and 
‘iollers” respectively), For the rest of the day, the beetles 
remained inactive 

At 19/0 hr, acriviry ground several particles (2, 9, 4, 5 and 
7) resucned: Air temperature was 28°C sand temperature Was 
27°C_ and the wind had dropped. However, humidity was 
exrrecily high and intense electrical activity overhead heralded 
a thunderstorm which broke at abet 2000 hr Upto this tiitic, 
beetles in groups 2 and 3 had moved their carmun several 
Mctres gqutl, and group 4 had carried Meir particle 12,3 m 
noth, The beetles in group 7 hau iuved around in a circle 
(10. 0m distnerer) surmunding (eit carrion and then upparently: 
burrowed, leaving pock-marks several nom broad in the 
ground. Activity ceased vermpletely during the rain trom the 
thunderstorm which eflectively ended my observations. 

There were iy Cracks fr activity the follawing morning, 
which was sunny and [8%C at OX) hr with a gentle southerly, 
The fin had soaked Io £5 cm and although | way able to 
rewuver several hai-buried. bedraggled pieces ol egesta, 1 was 
unable to find any beetles even though T destructively excavated 
each spot where (he observutinns had ceased during the senn. 
JL was not clear wherher the ezesta had been buried by the 
beetles ov, mune likely. wind-blown sani! and cain. 


135 


Presumably, the beetles had either dispersed individually or 
had carried and buried the carrion i the mierverting 10 hours. 


Why do phese bectles go va such lengths ¢o transport the 
fgesta! he tdaptive explanation for this furm of behaviour 
in dung beetles is that the Action reduces compelition for che 
resource from Fivals of the same species Or other specyes that 
comune Gung. Alcack suggests thar had ihe beciles 
remained at the site of deposit, the concentration of material 
might have had 9 higher probability af attracting vertebrate 
stuvengers or anls that could consume the egesta before the 
trogids Possibly, the beetles themselves would tren be put 
al msk as a nearty food resquree. 


Another explanation has Deen applied jo dung beetles i 
the tunneller and roller guilds thal need to eranspon (he particle 
from an area where it may have fallen an ground thar is 
unsuitable for burrowiig of that is loo exposed to hursh 
umhient canditions’®. This does nor mutually exelude the 
first hypathests and may alse be a valid explanation for the 
trogid behaviour ubserved in the present study; Onan unstable 
surul ulune, buried egesta ate likely to bo exposed by wand 
whereas in areus stabilised ly vepetition, this risk is lessened 
Funher, Incal suil moisture ts likely to be greater, perhaps 
enhancing the food quality of the currion, Relative humidity 
is un imMpomant factor cuntralling the behaviour of two species 
Ot Kaluhse: Onmorgus!!, High relative humidity restricts 
texpiralory water loss, jniysoves food (moist hair and kerannt 
quality and may compensate for faecal water toss!4, Perhaps 
the trogids T observed were vransporting their particles lang 
distances until they found clumps of shrubby vegetaGun where 
telahive humidity and sand stability were high and food quality 
would be enhanced when the particle was buried. This 
hypothesis awailsy testing, 


Tam gralefal for the coprologica) encouragement by the 
other members of the expedition (Fran Sheldon, Philippa 
Kneebone, Leslie Doddridge, Wendy Marth and John Slade) 
and especially the organiser. Jim’ Puckridge, whose local 
kacwledge and enthusixsny greatly enriched the Urip, Dr E. 
G. Matthews (South Australian Museum) provided keys, 
encouragement! and helpful discussion, Prof. Jolin Alcock 
(Anzona Suite Universny) guided! any reading on animal 
behaviour, and Dr CH. Scholtz (University of Pretoria, South 
Africa) Kindly sent me reprints of his work on African 
Omurgus. E thank Dr Margaret Davies, Alice Wells and 
Shelfy Barker, Mr Jim Puckridge and James Wallan, and 
an shonymous referee for comniciits on an early drattaf this 
TaMuScOpT And dircchan to Unfimiliar liternqiee: 


References 


'CSIR( (1991) “Insects af Australia” 2nd Ba(n (Melbourne 

_University Press, Melbuurne), 

~Matthews, By G. (1981) 5A Guide w the Geaera of Beciles 
of Buuth Australia. Purt 1 Archestenity amd Adcphago.” 
Special Education Bulletin Series (Suuth Australian 
Miisenm, Adelaide), 

*¢ivittin, 'V. & MoCaskill, M. (kds) (1986) “Ailas of South 
Australia.” (South Australiag Goverment Printing Division. 
Adelaney 

“Blackburn, T. (1895) Trois R. Soc S. Aust. 19, 27 60, 

*Sechultz, ©, H. 1986) Aust J. Zool. Suppl. Ser. 125, Wy 

*Loefinaas, S, (1932) Tj. Entomul, 75, 36-43 (cited by 
Scholtz"), 


TMiatthews, E, G. (1976) “Inseer Ecology (Queenstand 
University Press, Brishane), 

"Scholtz, C, N. & Caveney, 5, (1988) L Arid knw 18, 
TH 191, 

"Scholtz, C. H, (1991), [nveriebr. Taxon, 5, 827-835. 

“Holfter, G, & Matthews, E.G. (1968) Hol Bitomol, Mex. 
12-14. 1-312 (cited by Camberfan & Hanski!'), 

"Caviteefort, ¥. S Tlanski, 1, (1990) 96-50 fn L Hanski 
& Y, Cambefort (Eda) “Dung Beelle Ecology? (Princeton 
Wnrverstty Press, New Jersey). 


*Panmaret, 4, P, Kardici, N. & Bertrund, M1, (142) 4 Agi, 
Ecol, 29, 349-356 


136 


BShubeck, P. P. (1966) J. New York Entomol. Soc. 76, 
253-265. 
i4Lumaret, J. P. & Stiernet, N. (1990) 242-254. In I. Hanski 


& Y. Cambefort (Eds) “Dung Beetle Ecology.” (Princeton 
University Press, New Jersey). 


SAlcock, J. Arizona State University, pers. comm. 


'6Hanski, I. (1990) 5-21. In I. Hanski & Y. Cambefort (Eds) 
“Dung Beetle Ecology.” (Princeton University Press, New 
Jersey). 

"Scholtz, C. H. & Caveney, S. (in press) Ecol. Entomol. 17. 

'8] oreau, M. (1990) pp 31-38. In N. E. Stork (Ed) “The Role 


of Ground Beetles in Ecological and Environmental Studies.” 
(Intercept, Hampshire). 


ANDREW J. BOULTON, Dept of Zoology, University of Adelaide GPO Box 498, Adelaide, S. Aust. 5001. 


REDEFINITION OF UPEROLIEA LITTLEJOHNI DAVIES, MCDONALD & 
CORBEN (ANURA: LEPTODACTYLIDAE: MYOBATRACHINAE) 


BY MARGARET DAVIDS & GRAEME F. WATSON 


Summary 


Transcerions Of the Rival Soviety of S Aust 992), Dory De7-au 


BRIEF COMMUNICATION 


REDEFINITION OF UPEROLEIA LITTLEJOHN] DAVIES, McDONALD & CORBEN 
(ANURA: LEPTODACTYLIDAE: MYOBATRACHINAE) 


Uperalele litlejohin was described (yon preserved material 
Inthe collections oF (he Queenshind National Parks & Willie 
Service und (he Queenstiad Museum! Foe this reason, 
brelogieal dati Were absert trom (he description, as were 
ephemercal dita such as colour in tile 

Hh February 1990 and I9¥l, we visited Oeeupatiin Ligence 
7 Burra Ranve. 26.0 kin SW: Pentland, Okt (2oMa' Ss. 
M5 °H bb now inthe White Moontiins National Park, neue 
the type Tuculity ob the species. On the Hest visit Wwe located 
a single male in exvenely dry Conditions. but did notobiain 
further biological data. A Sevond Visit coincided with heavy 
huts, endhwe Toculed I~a-chonuses af O! firlep on, calleeted 
specimens and revorded calls, We present those dat here, 

Material ts depesited in the South Austratiun Museum, 
Adclaide (SAM) and the University of Adeliide Ostealogicul 
Collection (LIAZ). Methods of meiurementand ubbreviaitions 
nthe text follow Tyler (68). Selected mrtlersal was cleared 
andestuned for osteologied! examination’? 

Tape reconings were made using a Sony TCDSPRO. 
eissette recarder (tape speed +76 cm/s) and Beyer M-&R 
Gandini) dynanie mierophone. Air wet hulls teniperatures. (ihe 
eflcchive lenperuture OF 4 tree calling on hind) were miensired. 
Ue the calling site of cach individual using an eleeronie 
thermistor thermometer (Takara Digitiulir Model D6ih, 
Recordings were analysed on a DSP 5500 digital Sond Graph 
(Ray Elenicirics Corp.) using the in-built set-up AIG, with 
Playback cn Nukomich) Drawon cassche recorder, Overall 
HPWHS In kipe speed (Le, from recording to playback) 
areestimuled al less thin Q.5%. und she frequeney responses 
of ull oudho-cleetronic components are close te linear within 
the relevant frequeney ninge (hased an the nianuliclurer’s 
spec  Heationss, 

bor cach cull, three primary aliributes were determined: 
0) durttion, as the interval front the beginning of the first 
pulse to the ene of the last pulse pinsi: Ci) nimberot pulies 
per Note (direct county, and (ily dominant (reyuency (H7), 
asthe (iid value of the spectrum of power between the 
cursors of the whole note. Two derived attributes were 
determined: () pulse reperition rate (pulses/s), calculated as 
JOO0Un-+ pulses)/durubon in pis; sind amplitude modulanon, 
palculated by Using the only incastrable characteristics of 
envelope umplitude, the nexium amplitide CY) aid the 
Minmun aniplitide: (4), according to the formula. % 
ampliude jnodilition cquals WOON Y 4", Levels of 
resolution were less (Hin Tras tin fempoetlaspeets, ind less 
thin 10 Hy dor dominant frequency, 


Uperalea liitejohnd Mavies, MeDamild & Corben 
Hpervler Hinlejoht Davies. MeDonald & Corben, 1986 p.174 


Definition A motertiely large species [mules 20-32 mim 
SHoul-vent length (SV females 23-29 mm SV] Jacking 
maxilfiry wet. fingers unwebbed. poorly triaged, hasal te 
fo toe wehbjnw, lows fringed; dermal glands prominent: 
fentoparienil fontanetle modernity extensive; carpus of 5x 
vlerients: suitcroniedial proeesses af anteriog Hyule slender: 


Hhal crest absent: Short pulsatile call of eahe pulses. wath a 
pulse reperition cate ul about (at pulsests ut Tha5C 


Metered evaniiied, UAZ AV2, AITI4, AIT), BIT. BITIS-4, 
SAM R39402-9, 26.0 kin SW Pentland. Burra Range, 
25.1199}, M. Davies, Goh Watsem, KR. McDonald: SAM 
R39801, adie locality, (07 )990, Mo Danies, KOR 
McDonald 


Eyrernal horpholoy> Lith can be added (er ihe eriginl 
Hesenprion vther than an increkse inthe sive range of pales 
(Fin, 1) 


Colne in lifer Dorsunt grey With Welldelined dark chocolate 
markings. Proninent tubercles tipped with vpricet or cream. 
Parotoid and inguinal ghinds bull yellow. supralabial gland 
erodim. Flashes in the inguinal reejon and the backs of the 
thigh pre chrome onenge” 


Oseoliis A TUrthet six specimens were examined (Mhiree 
double-stained and three singhe-stained) but variability wr 
mininal anc does pot add tr the oryinal description, 


TABLE | Characteristics af the udveriixement cally of three 
individnals of Uperoter Inteqonni, recorded on 25.1194), 26 
Hn SH of Pentland, Old. Patras of amplitude modulation 
throughout the cally are shown in Table 2. Fyscrive 
teniperanive at the calloeg ses of etel iadivielual (air wet 
Pret) iy alser listed. 


——————————— 
Attribute 4) #2 al 


Temperature 1C) 22 764 26,7 
Call duration tniss 47,92) 52,73 Gi 44 
Domuninr frequency (Fz) 2080) TUR Bb) 


Now of pulses: N i K 
Pulse repetifion rates (pulses/s) 146.1 1328 fiau 


st 


Call One call ot each of three individuals wits analysed. arn 
usunmmary of the call characteristics ts listed in Table | The 
ddvertisement call ol Uperoleia hitlejuliat os a short amen 
dunition 53,86 ms), partly pulsatile call (iiean. pulse repetition 
mile 131.27 puisxes/s) The depth of amplitude modulation 
decreases throughout the call so thar individual pulses. are 
diffieult to discern, except at the heginning of the cull, alihough 
may calls also Hala isting! final pulse (Pas. 2). 

Table 2 shows the changing levels of amplitude modalatien 
in the three calls analysed. To phe car. the call isa sharp bud 
lick, With ud pulsandé nature. repeated ara fate ot about 20 
culls/nin. Among other species ot Cpemdteia whose culls bave 
heen deseuribed. six (UL aypera, U elancditese, UL lithemeda, 
Wntimuld, Uo minima onl Uo rygesa) produce “chick” ealls 
(short calls of Tylor ey af (980) with the call of t¢ aspera 
(at 256°C) durttion 30-35 ms: frequeney 2690-2900 He. 
number of pulses 3-6) pulse repetinion rate WA eel 
pulses/s”) being diost similar we than af Oe ditlejalini 


138 


TABLE 2. Patterns of amplitude modulation (%) throughout 
calls of three individuals of Uperoleia littlejohni recorded on 
25,1.199], 26 kin SW of Pentland, Qld. 


SSeS 


Pulse No. #1 #2 #3 
a a 
| 95 75 100 

2 87 78 90 

3 66 31 36 

4 15 6 35 

¥ 13 4 19 

6 17 42 20 

27 28 29 

& 38 50 59 


60 


dB 40 


0 12.5 


kHz 


Distribution and habitat: The distribution records of Uperoleia 
litlejohni are within the Einasleigh Uplands, northern Desert 
Uplands, and the north-western Brigalow Biogeographic 
regions of Queensland”. The geology of the collection sites 
ranges through Quaternary alluvium and colluvial sands, 
Triassic sandstones, undifferentiated Palaeozoic and 
Triassic?/Permian granites, and Upper Silurian/Lower 


Devonian granodiorites. No records are known from the 
Recent (<3 My. BP), extensive basalts of the McBride Plateau 
and associated lava flows of the Einasleigh Uplands". 
Vegetation types at collection sites are predominantly Iron 
Bark and Bloodwood (sometimes with Box) woodlands and 


open woodlands with tussock grasses on granites or sandstones 


25 37.5 50 
ms 


Fig. 2. Power spectrum and wave form of an advertisement call of Uperoleia linlejolini recorded on 25.1.1991. 26 km SW 
of Pentland, Qld. at a wet-bulb air temperature of 26.2°C (#1. Table 1). Note that the ordinate of the wave-lorm display 
is not labelled because it depicts a relative linear scale in volts 


of the Einasleigh Uplands and northern Deseri 
Uplands"! *5, Laneewood (Acacia shirleyi) communities 
with an understory of Spinifex (Jriodia sp.) on dissected 
Warrang sandstone ol the Desert Uplands. and the Eucalyptus 
populnea or E. microtheca woodlands along drainage lines 
in the northwestern Brigalow belt. 

Altitudes at collection sites range from 150 m at Caerphilly 
Station (21°03, 1455 32") to approximately 1000 m near 
Herberton (17°23', 145°23'), with most records within 


‘Davies, M., McDonald, kK. R. & Corben, C. (1986) Proc. 
R. Soe. Vict. 98(2), 147-188, 

“Tyler, M. J. (1968) Zool, Verhandl. Rijksmus. Nat, Hist, 
Leiden 96, 1-203. 

‘Davis, DD. & Gore, V, R. (1947) Fieldiana Tech. 4. I-16, 

*Dingerkus, G. & Ubler, L. D. (1977) Stain Technol. 52, 
229-23). 

*Dorbuck, T, (Ed.) (1978) “The Radio Amateur’s Handbook”. 
55th Edin (American Radio Relay League, Newington). 
*Smithe, F. B. (1975) “Naturalists Color Guide” (American 
Museum of Natural History, New York). 

"Tyler, M. J., Davies, M. & Martin, A. A. (1981) Aust. 
J, Zool. Suppl. Ser. 79, 1-64, 

“Tyler, M. J., Davies, M. & Martin, A. A. (1981) Ree, 
W. Aust. Mus. 9, 147-172. 

“Stanton, J. P, & Morgan, M, G. (1977) Project RAKES 
— a rapid appraisal of key and endangered sites. Report 
No, I: The rapid selection and appraisal of key and 
endangered sites; the Queensland case study. (University 
of New England, School of Natural Resources. Armidale). 


139 


altitudes of 300-900 m. Rainfall ranges from 480-1150 mm, 
with most collection locations found within the 500-800 mm 
rainfall isohyets. Rainfall is strongly seasonal, concentrated 
in (he summer months from December to March '45, 

The recorded frogs were calling on a steep scree slope, or 
at the edge of a stream at its base. They were located at the 
base of, or between, Triodia tussocks. A second chorus was 
found around the edge of a roadside scrape, and individuals 
were calling at the bases of grass tussocks. 


“Day, R. W., Whitaker, V. G., Murray, C. G., Wilson, 
1. H. & Grimes, K. G. (1983) Geological Survey of Qld 
Publication 383, 1-194, 

"CSIRO (1967) Lands of the Nogoa — Belyando Arca, 
Queensland. Land Research Series No. 18, 

"CSIRO (1970) Lands of (he Mitchell — Normanby Area, 
Queensland. Land Research Series No. 26. 

"Perry, R. A. (Compiler) (1964) General Report on Lands 
of the Leichhardt — Gilbert Area, Quéensland, 1953-54. 
Land Research Series No. U. 

4Lee, D. M. & Gaffney, D, O, (1966) “District Rainfall 
Deciles — Australia” (Bureau of Meteorology & A.G.PS,, 
Canberra), 

"Bureau of Meteorology: (1977) “Rainfall Statistics for 
Australia”, (Bureau of Meteorology & A.G,P,S., Canberra). 


MARGARET DAVIES, Department of Zoology. University af Adelaide, G.P.O, Box 498, Adelaide, S, Aust. 5001. GRAEME 
F. WATSON, Department of Zoology, University of Melbourne. Parkville, Vic. 3052 and K. R. McDONALD. Queensland 
National Parks and Wildlife Service. Pullarenda, Townsville, Qld 4810. 


NEW RECORDS OF MESAPHORURA (COLLEMBOLA: ONYCHIURIDAE, 
TULLBERGIINAE) SPECIES FROM AUSTRALIA, MACQUARIE ISLAND 
AND THE ANTARCTIC 


BY P. GREENSLADE 


Summary 


fransactiony of the Reyal Suetery af 8. Aust. 992), 116(4), 14F-143, 


BRIEF COMMUNICATION 


NEW RECORDS OF MESAPHORURA (COLLEMBOLA: ONYCH 
TULLBERGIINAE) SPECIES FROM Sry aig MACQUARIE ISLAND AND THE 
ANTARCTI 


The Tullbergrinac is a subfamily of strongly reduced 
Collembola. lacking oelli, pigment und furca, which is 
adapted for suil living. Within thix subfiutily the genus 
Mesaphorura currently comprises oyer 20 described species 
most oF which ace-only knewn from the Northern Hemisphere. 
Specimens belonging to the genus Mesuphorura are 
commonly found in Australia in moist soil under arable and 
grazing regimes, anc also, raore rarely, (a soils under mittive 
vegemmon, Mesapharura krausbaueri Borner 1901! was 
vecorded from Australia’? and specimens determined ss 
Mesaphorura sp. krauxbaueri group were yecorded fram 
southern Australia and Macquarie Istand®, Three species of 
Mesaphanina already known from Europe have beer identified 
from Australia, and they have all probably been introduced 
relatively recently with Ruropeans. All matertal discussed is 
deposited inthe South Australian Museurn collection. 


Mesaphorura Borner, 190) 


Diagnosis: elangate poduromorph Collembola about 
500-600 ym Jong, lacking ocelli. pigment and furca, and 
possessing pscudocelli on head, thoracie and abdominal 
segments with the formula WOU/O00I0 or H/OMHOOH): antennal 
TE] organ normally with two cutved cylindrical elubs and lwo 
small pegs, only a smal) cuticular swelling in front of pegs 
without enlarged granules; ant IV without greatly enlarged 
sensilla; postantennal-organ elongzte. consisting af two parallel 
rows each OF 1D to 20 simple clongate vesicles arranged a} 
night angles iy longitudinal axis of the organ; abd VI with 
a pair of creseentic cuticular ridges anteriarly and two 
posterior anal spines. shorter than claw. 

‘Abbreviations: collectors, KK. K, King, PG. Greenslide, 
HW, H. Womersley. 


Key to Ausiratian species (after 7%) 


1, 34 4 median microchaetae in antetior row between ay 
un abd V (a, present); a, either a nticru- ora magrocheti 
on abd IV, L, present on anal lobes: a, of abd V nor 
Jisplaced anteriorly: psetdecelli on thorax close to mid 
ling, behind or between py and py... .... nti eed 
2 + 2 median microchaete in anterior raw between ay 
an abd V (a, absent); 4, a microchacts on abd IV: 1, 
missing on anal lobes: a, of abd V displaced anteriorly’ 
Picudacelli of thorax between im, and Ps...Md. entica 

- M, present on abd TV; long macrochaeta (1.6, a, on abd 
1V) over twice the length of microchaeia fay); 
macrochaéts #, o1 abd’ V is shorter than macrochaeta p.} 
Pp) 4 macro- and p, 4 microchaeta on abd IV. . 

PT ere Le ee 4 M, macrochaeta 
M. absent on abd IV; macrochaeta (1. a, on abd JV) 
only 1.8 times as long than microchacta (a,)! macrchueta 
a, on abd V longer than Py Py a Mactochaeta und p, a 
micrachaeta on abd TV... 2. ss UM. Sasi 


me 


Mesuphorura macvrochaet Rusek. 


Mesaphorura macrochaeta Rusek 1976 p33 
FIG. | 


Matcris! examined, Australian, Antarctic Territory, 
Mawson Station, pot plant soil (Coleus, Philedendron?), 
January 1989, PG, oa, 200 exs; Macquarie Island, fsthimus. 
in Breenhouse, soil and muss, 2.x11.86, PG; New South Wales, 
Armidale, Chiswick native pasture. plot 8, 2).vili.78, 26.11.73, 
KK, 2 ex; Chiswick improved pasture, ungrazed, 26,11,72, 
KK, 2 exs: Cambewarra Ranges, 10k W of Narvonu, leaf 
liter, Sept 1990, PG, | ex; South Australia, Mt Lofty Ranges, 
Bridgewater, Engelbrook Reserve, leaf liver, I6v.7), PG, 2 
exs] Belair, in inuss, April 1W38, HW, t ex; Belair, grass 
mowings, 27.v.1971, PG, 2 exs; Coorong, Coolatoo, pitfall traps 
in grass beside rowd, 28.ix-8.x,75, PG, 1 ex IS k N Mt 
Gambier, Pinus ractata leat litter, 19.v,1975, PG. 2. exs, 


Distribution: described from Canada but comunon in North 
Americt and Europe. Mesapherura macrochaen is abundant 
in improved pasture in southeastern Australia and has been 
introduced jo an Australian Antarctic ‘Territory Station and 
to Macquarie Island in imported -soil, probably frorn 
Tasmanis. 


Mesaphorura erttica EM\ts; 


Mesaphorura critica Ellis 1976 p. 230, 
FIG 2 


Material examined: South Australia, Kognamore Station, 
340 km NNE Adelaide, Black Oak Creek, leaf litter, 
75.vi. 1971, PG, 1 ex, 


Distribution: previausly only known from Europe. 


Mesaphorura yosiil (Rusek) 


Tullbergia yostit Rusek 1967 p. 191, 
FIG, 3 


Maternal examined; New South Wales, Armidale, Chiswick 
native pasture plots, plot 8 21,viu.78 KK. 2 xa; Chiswick 
improved pasture, ungrazed plots, 26.11.72, KK, | ex, 
Queensland, 17 kro cast of Killarney,. wet sclerophylt (nrest, 
teat liner, l6v.76, PG, 1 ex; Great Barrier Reef, Swain’s Reef, 
Frigate Cay, 22.yii.1I983. KK, | «x South Australia, 
Koonamore, 340 km NNE Adelaide, Black Oak Creek, leat 
litter, 25,vit.73, PG, 2 exs; 10 km N Whyullu, Middleback 
Sin, under Casuarina strict, 8.x.79, PG, | ox, 


Pistribution: Europe, Nori America, China’, Australia. 
New Culedonia® 


Figs 1-3. 1. Mesaphoruru macrochaetac Rusek. Dorsal chaetotaxy of abdomen IV-VL, 2. M. critica Ellis. Dorsal chaetotaxy 
of abdomen FV-VI. 3. M. vosiii Rusek. Dorsal chaetotaxy of abdomen IV-V1. 


Both M. yosiit and M. macrochaeta are found together in 
improved pastures in southeastern Australia, generally M. 
yosiit is found on warmer sites and M. macrochaetae in cooler, 
more southerly regions. Is likely that both were introduced 
to Australia with Europeans. Morphological differences 
between the species are given by Rusek® and are cited in the 
key, The specimen from Darlington, Western Australia, 
determined by Womersley* as M. kraushaueri, is not in good 
enough condition to be identified, but other specimens from 
Belair, South Australia, also determined by Womersley as 
M. krausbaueri, are in fact M. macrochaeta, \t seems 
probable that M. krausbaueri does not. occur in Australia. 


‘Borner, C. (1901) Zool. Anz. 24, 1-15. 

2Womersley, H. (1935) Trans. R, Soc, $.Austr, 59, 207-218, 

3Womersley, H. (1939) “Primitive insects of South Australia” 
(Government Printer, Adelaide). 

‘King, K. Greenslade, P. & Hutchinson, K, (1986) Aust. 
J.Ecol. 10, 421-427, 

Greenslade, P. & Ireson, J. E. (1986) J. Aust. ent. Soc. 
25, 273-291. 

5Greenslade, P. (1990) Pap. Proc. R. Soc. Tas. 124(1), 35-50. 
7Rusek, J. (1971) Acta ent. bohemoslay. 68, 188-206. 
5Rusek, J. (1976) Canad. J, Zool, 54 (1), 19-41. 


143 


All three species are likely to be more widely distributed 
than these scattered records suggest. In Canadian forests M. 
yosiit and M. microchaeta can occur together but have slightly 
different vertical distributions with M. macrochaeta markedly 
aggregated in the humus layer and upper soil horizon from 
Oto 5 cms in depth, and M. yosiii concentrated lower in the 
soil profile and more randomly spaced'*. In another 
Canadian forest where M. macrochaeta was absent, M. yosiii 
occupied the whole soil profile. This suggests possible 
competitive exclusion of M. yosiii by M, macrochaeta on some 
sites. Mesaphorura critica may have been included with the 
species M, yoviii in these ecological studies. In Australia, 
M. critica has been found only under arid native vegetation. 


*Rusek, J. (1986) pp 73-78. Jn Dallai, R. (Ed.) “2nd 

Eeecaasocal Seminar on Apterygota.” (University of Siena, 
y). 

Fjellberg, A. (1973) “Identification keys to Norwegian 
Gollembola®. pp 1-152. (Norwegian Entomological Society, 
As-NLH, Norway). 

NEMlis, W. N. (1976) Tijdschr, Ent, 119(8), 221-326. 

?Rusek, J. (1967) Acta ent. bohemoslov. 64, 184-194. 

3Weiner, W. & Najt, J. (1991) Mem, Mus.natn. Hist. nat. 
(A), 149, 119-130. 

Rusek, J. (1979) Acta ent. bohemoslov. 76, 1-9. 


P. GREENSLADE, c/o CSIRO, Division of Entomology, GPO Box 1700, Canberra, A.C.T, 2601, Australia, 


ECOLOGICAL AND BIOLOGICAL NOTES ON THE RARE PLANT 
HEMICHROA MESEMBRYANTHEMA F. MUELL (ARMARANTHACEAE) 


BY JOHN L. READ 


Summary 


Transaiions uf the Royal Society of S. Aust, (1992), W6(4), 145-146, 


BRIEF COMMUNICATION 


ECOLOGICAL AND BIOLOGICAL NOTES ON THE RARE PLANT HEMICHROA 
MESEMBRYANTHEMA F, MUELL (ARMARANTHACEAE) 


The succulent shrub, Hermichinad aesembrvunhema, wits 
described in 1473 after being collected in the previous yeay 
by the explorer Emest Giles in the vicinity of Lake Eyre, South 
Australia’, No further South Australian speciniens were 
collected until 1984 when 4 specimen was collected from 
Serangways Springs, 112 years after ir was first discovered?, 
[1 was suggested that the population at Stranyways was likel 
tn be the source of the specimen collected by Ernest Gitex’. 

tn October 1991 the Roxby Downs Field Najiiralist Club 
lncated H. mesembryantheme at the base of Mt Kingston in 
the Denison fanges, South Austrslia (28° O2/S, 135° 54"), 
A tollection was lodged at the State Herbgriwmn of South 
Australia. A subsequent seareh of the region by the authai 
an Apnl 1992 vevealed that the population of H. 
tiesembryantheme was apparently restricted 1 & Saal regian 
betyween the buse of Mt Kingston.and the Willparoona Springs 
fhag. 1. This population is only 5 km from the site of the 
Peake Overland Telegraph Station which was dlyo visited ty 
Ernest Giles!, thus casting doubt on the provenance. of the 
orminal collection, H, mesembryunrhema has been recorded 
from eight sites in Queensland and populations upparencly 
intermedia: between H. mesembryanthema and H. alicnadra 
have been revorded in Wesrern Austratia’, 


OGONADATTA 


eo 
WILLFAROONA 
SPRINGS 


STRANGWAYS 
SPRINGS 


MARHEE 


isa 


1 


i 
MARREE 
e. Lan | 


| 
“Gain! | broxe downs 
ih! 
| 


ADEL AIDE 


Tig, |. Major locations mentioned with respect ta Hemiehroa 
mesewitrvantheme in South Australia 


The Willparcona Springs population of ca. 150 mature 
Hemichroa mesembryanthema shrubs was located in a.sub- 
circular patch measuring ca, 150 % 160m. These shrubs 
were located 120 m from the nearest spring as detined by 
a bed of Cyperus pymnveauos. The population was 
growing on powdery satine clay soil (3.14% Cl) with a 
capping Of angular feldspar rich gronite and quartz gravel, 
approximately twa metres above the spring sediments. 
Although H, mesembryanthema was the tallest shrub in 
the region, Halosareia spp, dominated the chenopod 
shrubland, Atriplex yesiceria, Nirraria pilliardieri, 
Gunniopsis quadrifida and a Frankenia 9p were also 
prevent. Two small streams supporting predominantly 
Acacia cambagei, A. yietoriae and A. tetragonophylla 
divided the population, The remainder of the hillside above 
ihe springs was dominated by chenopod shrubland while 
the majority of the drainayeé line associated with the springs 
was unvegetated. 

AUl of the #. entesemtbryanthema bushes appeared to: be 
nature and the larvest individual was approximately 0.8m 
tall and over Lm wide. A (ull description of this species 
is presented elsewhere”. Most of the individuals were 
Nowering in April 1992 and the small whice Mowers with 
brilliant red stamens produced a beautiful perfume. Muny 
insects Including wasps, butterflies and flies were attracted 
to these flowers. The wasps, particularly a tucge black 
species, were cuvered with 4 cunsidarable Joad of yellow 
pollen and are probably an important pollinotor of H. 
mesembryanthema, 

No evidence of browsing was detected on any of the shrubs 
although cattle, donkeys und rabbits all inhabit the regian. 
This supports the observation that the Strieways 
Population of A. mesembryanthema was not tiuched by 
either catls op rabbits". Interestingly itwos presumed (hat 
A. mesembryanthema had been forced to extinction by 
domestic stock and rabbit grazing’. Although introduced 
herbivore grazing on seedlings cannot be discounted, il is 
unlikely thar the apparent rarity of this species is related 
fo grazing pressure. 

The discovery of a further population of H, 
mesembrventheme in close proximity to found. springs 
faises the possibility that there is some form of assocation 
between Al, mesembryanthena and aeesian springs, Most 
yegelation associated with springs grows directly on the 
Vent, tail of scepage zone of the spring. However, it is 
possible that AL mesembryantherma volics on certain edaphic 
or hydrological properties which are found in a zone ata 
greater distance {rom the springs. A. mesembryanthema 
is evidently not obligately: tied 10 springs in all parts of its 
Tange since the Queensland collections are from alluvial 
run on areas in hilly country, or saline areas’, The 
common factor with all of these tocalities 19 thar they all 
appear to be contined to comparatively moist regions which 
aliggest Chit wallet stress may be limiting. 

Although the Willparoana population has significantly 
increased the recorded South Australian range of A. 
mesembrynnihema , this specics siust still be regarded as 


146 


rare in South Australia, with only two known populations 
estimated to comprise a total of less than 1500 plants. The 
main population at Strangways Springs was under some threat 
from road building activities? and is potentially threatened 
by off-road driving, since the Oodnadatta track and station 
tracks pass through it. However, grazing is not believed to 
be a problem, The Strangways population also appears to have 


'Giles, E. (1899) Australia Twice Traversed, vol. 1 (Sampson 
Low, Marston, Searle & Rivington: London). 


2Chinnock, R. J. & Badman, F. J. (1986) Muelleria 6, 
205-209, 

3Palmer, J. pers. comm. 

‘Leigh, J., Boden, R. & Briggs, J. (1984), “Extinct and 
Endangered Plants of Australia.” (Macmillan, Australia). 


JOHN L. READ, PO, Box 150. Roxby Downs, S.A. 5725. 


spread to limestone mounds in recent years’ where it was not 
observed in 1984? or in 1978”. Since the discovery of the 
Queensland populations, the status of the species has been 
lifted from presumed extinct* to “poorly known”, 

Further research is required on the physiological and 
ecological requirements of H. mesembryanthema to establish 
a meaningful management plan for this rare species. 


5Symon, D. E. (1985) Botanical notes on Mound Springs and 
Bores pp 27-43. In J. Greenslade, L. Joseph & A. Reeves 
(Eds) “South Australias Mound Springs” (NCSSA, 
Adelaide). 

Briggs, J. D. and Leigh, J. L. (1989). Rare or Threatened 
Australian Plants 1988. Revised Ed. Canberra, Aust. 
N.PW’S. Special Publication No. 14. 

7Badman, FE. J. pers. comm. 


EGGS AND INCUBATION IN THE AUSTRALIAN LIZARDS 
AMPHIBOLURUS NOBBI AND EREMIASCINCUS RICHARDSONI 


BY T. P. MORLEY 


Summary 


Transactions of the Rovel Society af §. Aust. (992), U6t4), 147-148. 
BRIEF COMMUNICATION 


EGGS AND INCUBATION IN THE AUSTRALIAN LIZARDS AMPHIBOLURUS NOBBI 
AND EREMIASCINCUS RICHARDSONI 


Reproductive biology, particularly egg-faying, incubation 
and Hconate sizes, is poorly known in Australian lizards! , 
Here [ present dita on these parameters in the Australian 
lizards Amphibolurus nebbi -cogyeri and Eremiaseinens 
richardson. 

Amphibolurus nubbi Witten, 1972 includes two 
subspecies”, and all available ecological data relate to the 
nominate subspecies***>_ No-studies have’ been conducted on 
A. on. coggeri, and in South Australia the species is poorly 
collected and little known®. The Desert Banded Skinks or 
Sandswimmers (Eremiascincus spp.) aré distributed over most 
of arid and semi-arid Australia’? and are abundant in 
suitable habilals. The biology of the genus is poorly known 
and accurate reproductive data are only available for oyiparous 
E, richardson? recording its clutch:size'*. The accuracy of 
Iie ayailable information on £) fitsciolatas is in doubt, having 
been reported as a viviparous species!*. 

Since jate 1987, my collections at Swan Reach Conservation 
Park, and those wf Mark Hutchinson in Brookfield 
Conservation Park have in¢luded four gravid A. nohbi copgerl 
(Table 1) and one Eremiascincus richardsoni. After collection 
all femates were placed in individual cages, furnished with 
a hide: box and a nest box filled with moist sphagnum moss. 
Fresh water was available, and a climbing branch provided. 
Lizards were offered various insects and feeding often occurred 
until the day before oviposition, 

The epes were marked and measured, with vernier calipers, 
to the nearest 0.1 mm (Table 2) and were placed on a medinin 
of vermiculite and minwater (50:50 by weight), in a small 


Plastic container, with 12 small holes drilled into the hd to 
allow for air-exchange, The firsr two cluwhes of 4. 1. coxgert 
egps were incubated ul room ternperature (20-34°C), The 
others, and the £. richardsoni egps, were incubated in 
temperature controlled (27-31°C) snake cage. The eggs were 
checked daily, and (he medium sprayed, us necessary, with 
rainwater (hat was the same temperature. 

Each a, a. coggeri laid a clutch of 57 epgs 
(24,%1,1987-10,1.1991) (Table 1). Female SVL and ¢latch size 
were not significantly correlated (r = 0.8465, 0.1 > P > 005). 
The nest boxes were not used: all cggs were laid on the floor 
of the cage. The feraale E. richardsani (SAM R375, SVL 
90 mm) fnid four egys in the afiernoon of 18, xii.1990. These 
eges were laid in the sphagnum moss, and cach adhered to 
one other egg in the clutch. 

Five. A. 4. Oaggert eggs from the second cluich were slightly 
collapsed and peur shaped (vs. oval) upon laying, and went 
mouldy.during the first week of incubation. These eges were 
opened, prior to disposal, to establish fertility. All were 
inferule. The A. n. coggeri eggs maintained in the more 
controlled environment were more succcsstul in both hatching 
rate, and a shorter incubation period. All eggs incubated by 
this methed, successfully hatched after % = 47.25 + 2.71 
(44-50) days, whereas the eggs in the uncontrolled conditions 
took X = 62.57 + 2.71 (56-73 days), and two embryos were 
dead oe severely deformed, 

On the 27. xii.1990 it was apparent that only ovo of the E. 
richardsoni eggs (nos. 2 and 3) were fertile. They had 
increased ih size, had a pinkish tinge and bloud-vessels were 


TABLE | Source and clutch sizes for gravid female Amphibolurus nobhi coggeri, 


_ OO eee SO Rn 


Feniale BVL Locality Dute Date Clutch SAM 
No Collected Laid Siz Reg, No, 
ge 

i 30) Swan Reach CP 9.x, 1987 Z4.-xiJOR7 7 = 
2 84 Swan Reach CP 9.%,1987 28.x1, 1987 7 — 
a T4 Swan Reach CP 18,41, 1989 10.1,1990 5 R36316 
4 bi) Brookfield CP 1. xii.3990 12.17.1990 s* R36997 
5 69 Ti Tree Well 8.xi1.1977 4 RI65k7 


_—_———-- rrr 


© Two of these cggs were laid in the bag following collection. When discovered they were not viable and were discandsd. 


TABLA 2, Egy atid neonate sizes in Eremiascincus richardsoni ard Amphibolurus nobbi coggeri expretsed ast + 5D if’ 
uppropriate with range in parenthesis, 


ere 


Species Egg Sizes Neynate Sizes 
Length Width SVL TL 
a ge 


& richardsoni 18 58 9.95 32.5 78.5 
(16.5 ~ 19.5) (9.9 - 10.1) (31 - 34) 77 - 3H 

AL coggeri 160 + 1.49 912 + 05 28.53 4+ 0.74 80.67 + 2,97 
{12.1 - 18.3) (87 - 9.6) (27 = 30) (76 - B7) 


rr 


148 


forming on the inside walls. Eggs 1 and 4 had not changed 
in size or colour, and were thought to be: infertile. 
Measurements of the eggs could not be taken at this stage 
due to the adherence and shape of the mass. After 36 days 
incubation, on 23.1,199/ the shell on egy no. 2 had split. This 
was noticed at 1935 fr, bur the lizard did nel emerge until 
0315 hy the next morning. The shell on egg no. 3 was split 
at 2020 hr on 23.i.1991 and full emergence occurred at (1922 
he the next day, after 13 hours in the open egg shell. The other 
iwo eggs (I and 4) were mouldy. and were opened before 
discarding, to confirm them to be infertile. Too few eggs were 
avnluble to permit opening an egg to determine at what stage 
of embryonic development this species lays its eggs, The 
incubation period shown here is similar to that for Clenanes 
lueniolatus’, a similar sized skink, whose eggs were laid at 
stage 30” All neonates were measured at hatching (Table 2), 

Most of the 4. .n. coggeri neonates were released at the 
collection site of their respective parents. The deformed 
specimen and four neonates were placed in the South 
Australian Museum (SAM R35843-44, 36318-19 and 37951). 
The E. richardyoni neonates were maintained, 

‘To supplement the observations reported here, specimens 
held in the South Australian Museum were examined for 
gravid fernales. Greer examined all specimens of 
Eremiascineus in State Museutn collections prior W 19798, 
therefore only specimens of Eremiascincus registered after 
that year were examined, 

Only one specimen of A. n. coggeri (R 16587) had 
oviducal eggs (Table 1). The largest egg in this specimen 


‘Greer, A. E. (1990) “Biology and Evolution of Australian 
Lizards”, (Surrey Bearty & Sonos, Chipping Norton). 
*witten, G. J. (1972) Herpetologica 28(3), 191-195, 
Witten, G. J. A Study of Amphiholierus nobbi Witten, 1972. 
Unpubl. M.Sc. Thesis. University of New England. 
4witten, G. J. (1974) Aust, Zool. 18(2), 129-132. 
Switten, G. J. & Heatwole, H, (1978) Copeia 1978(2), 
362-364, 
*Houston, T. F, (1978) “Dragon Lizards and Goannas of 
Sonth Australia’. Special Educational Bulletin Series. (South 


(168 »® 82 mm) suggests that these egps wetc hear 
oviposition. The only data for clutch sizes in A. nethi are 
related to the nominate subspecies (3-4), which has a 
smaller cluich size than reported here for A. n. coggeri (4-7). 
This suggests 9 correlation between female size and clutch 
size. as A. nv. cogueri is larger than the nominate race’, The 
clutch size reported here for A. n. caggeri is, however, similar 
to thosé reported for A. muricarys and A, norrisi (3-8 and 
37 respectively)', 4. nobbi’s closest relatives", and both 
these species are repurted to be larger thin A. 9. coggeri (75 
vs 100 and 110 mim SVL respectively)’, 

No further specimens of gravid E. richardsuni were found, 
but two &. fesciolans (RA0948 and R36137) were found with 
well-developed oviducal eggs (5 and 3 respectively). These 
epgs were surrounded by a thin shell membrane, the 
appearance of which suggests thal the cggs would haye been 
voided. 

The egg-laying reported here confirms observations on 
aviparity in E. richardseni'®, and the findings from dissected 
Museum specimens supports the suggestion that previous 
reports of viviparity in E fasciolaras may be in error. 

The S.A. National Parks & Wildlife Service provided 
collecting permits, Dr Mark Hutchinson collected two of the 
specimens on which these observations were made, allowed 
me fo-examint Museum specimens and read drafts of the 
manuscript. Adrienne Edwards provided data for Muscurn 
specimens and Brian Miller assisted with weighing the 
neonates. David Langdon and Ed McAlister read the final 
drafts of the manuscript, which was typed by Judy Woolman, 


Australian Museum,, Adelaide, } 

7Cogger, H. G. (1986) “Reptiles and Amphibians of Australia” 
4th Ed. (Reed,, Sydney). 

*Greer, A. E. (J979) Ree. Aust. Mus, 32(7), 321-338, 
Taylor, J. A. (1985) Herpetologica 41(4), 408-418: 

‘OMufaure, J. P. & Hubert, J. (1961) Arch. Anat. Microscop. 
Morphol, Exp. 50. 305-328, 

“Witten, G. J. & Coventry, A, J. (1984) Proc. R, Soc. Vict. 
96(3), 185-59. 


T. B MORLEY, Roya! Zoological Society of S.A., Frome Road, Adelaide, 5, Aust. 5000. 


RELICTUAL POPULATION OF TILIQUA SCINCOIDES 
(SAURIA: SCINCIDAE) IN NORTH-WESTERN SOUTH AUSTRALIA 


BY G. R. JOHNSTON 


Summary 


Transactions of the Reyal Soeiery of 8. dase (Wor), Widids 


BRIEF COMMUNICATION 


)40- [4 


RELICTUAL POPULATION OF TILIQUA SCINCOIDES (SAURLA > SCENCIDAE) 
IN NORTH-WESTERN SOUTH AUSTRALIA 


TVhe mountin rninges Gf central Australia ure kiown to 
provide loculised, mesic retuvia for several groups al 
organisms, including pkents!, scorpions”. frogs’ and 
reptiles? While some populations isolated in central 
Australia have diverged considerably and represent endemic 
species! "and others are retictual populations of species 
occurring i other parts of Australia’? all are of 
considerable biogcographie interest, 

It fas long been recognised thar some taxa oeeurring as 
isolates th central Australia have ther closest relatives in the 
Tropical north of Australie anc may be reliets ofa more tropical 
elinnic in the pust™®. Tis now clear that there are also 
significant links between the fina and flora of (he central 
ranges and fempenwe southern Austratia’ 7, 

The senneid fizard, Tlique scunemdes. is one of the most 
firniliar species of repile in Australia, Ip South Australia it 
has hitherte been regarded as in inhabitant of the cool, wel 
southern areas, extending as a series of relictual populations 
Into the nore mesie valleys in (he Flinders and Gawler Ranges 
(Fig. H. Extralinitally this species oecurs in Victorta, eastern 
New South Wales und Queensland. and the north of the 
Northern Territory amd Western Australia®. 

This paper reports the occurrence of Tilique sctmordes in 
the arid northwest of South Australia, 740 km NNW of ibs 
previously recognised range in this state. Two specimens have 
been lodged inthe South Australian Muscum (SAM) to verily 
this repon 


Re hrti Se A 


* 
* 


L 
| 
' 
| 
| 
( 


goo 


KILOMETRES 


Mig. L. Distribution of Tiliquerscineoides in South Ausualia. 
Solid circles denote S.A, Museuin specimens. The stars 
show. the new records [rom Mimil and Pukaya, 


One specimen (SAM R33V39) wats colleeted at the base 
Of 4 Linge sranite outerop closelo Mimili tes Mount Lyenerd 
Atation) (27°OI'S.. 132°43"B). in the Everstd Rata on 
I2jy.1989, A second specimen (SAM R4dU 30) was colleeted 
at Pukatya (ex Ernitbetla Mission) (26°18'S, 2°08 Ey jie ihe 
Musgrave Range of 20,1992, Both specnpens were initially 
sighted basking among rocks and Were rerioved fron crevices 
int whieh (hey had moyed when disturhed The nicky 
outcrops of the Everard and Musgrave Rauges provide 
localised mesic refugia which are surrounded hy harsty, aed 
sandplains and mulga country, Che paves suppor very 
localised, dense stands of denver and Fics. A farther sis 
specimens were observed at Pukula frm 20 254 192. 
Furthermore, the Aboriginal people whi livesat Minit told 
me that 7 sedternedes ts cyamon in the boulders atthe bases 
of the weanitie Wills iy that irea’, 

There was no doubt as to the specific Wenety of these 
speciiens.as therr anterior lemporals were much longer that 
broad, & dhignastie charicteristic of 7 seircaides in 
Australie! AT) opher Australian species: ot Tiligieet Wate 
fragmented temporal scales, However both of rhe specimens 
collected differed front typieal southern South Ausuntlian 
populunons of 7 scinudes in severil respeets (Pig. 2) They 
were Jane anpmals (S¥l, = 280 qm. 325 moi), and 
considerdbly more mbust in bouy form (ban sauwhern 
specimens, The dorsum was pille grey With nine trrepulan, 
hrown, (ninsverse bands on the body Compared with 5-6 bands 
onthe body ofother South Australian specimens Both lacked 
the distinet black temporal streak typieul ut other Seuth 
Austrilian specimens and the tail was Variegated with bhick 
am! pale grey which tended tu form very meister bands 
proximally, Whereas 7 semeerdes trom elsewhere in South 
Australia have a series ob distinet bands on the nut 

Two other species of Tiligae oceur near Minuli ane 
Pukaga!! and both are Kiown, and distinguished from eagh 
other and from 7 semcoldes by local peaple!?  Tilique 
multijasctate ogeurs i [Frode daniinated sandy country and 
Is called “Langku"” 7ifiqne ocuipitalis occurs in mule country 
and ts called. “Hingkarkara™. Viligua veincoidies & culled 
“Tillyarka’y 

Itscems likely thal the Z srinceides Ty northwest South 
Australie ure isolated fron the restal that species’ range and 
represent the remmants of a formerly more c&ktemave 
dipttthution which may have become resirieted by merusing 
‘ridily to isolited patches of suitable habital during the 
Pleisocene?®, The possibility exists that further isolated 
populations of 7 setncondes may be found in arher merges 
which may provide suitable mesic refugit in (he aortlvoel the 
state 

The fact that 72 scimepides does pot oceor im the MacDanald 
Runges! may indicate thal this species did not cross the [race 
al sand plain and dines between the Miisurave and 
MacDonitld Ranges, Pianka" idemified ities tract of land as 
a corridor foe latitudinal dispersal of sand pho inhabiaiinp 
reptiles. The same imiet may have bees 4 barrier (0 longitudinal 
dispersal by at least some animals with different habitat 
requirements a8 shawn by (he occurrence of several species 


Fig. 2. Tiliqua scincoides from Mimili in the Everard Ranges, South Australia (SAM R33939) showing the robust habitus, 
greater number of bands on the body, indistinct temporal streak and variegated tail which distinguish central Australian 
specimens from those elsewhere in South Australia, SVL = 325 mm. 


which find their geographic limits on either side of it (e.g. 
Litoria gilleni, Pseudophryne occidentalis, Ctenotus rufescens, 
C. caudicinctus)®. 

The occurrence of 7. scincoides in central Australia 
represents a significant range extension for this species and 
provides further evidence that the mountain ranges of that 
area provide an important refuge for non-xeric adapted 
organisms. More importantly, this record represents a most 


'Jessop, J. (ed) (1981) “Flora of Central Australia.” (Reed, 
Sydney). 

Koch, L. E. (1977) Rec. West. Aust. Mus. 5, 83-367. 
Tyler, M. J. (1972) Trans. R. Soc. S. Aust. 95, 215-217. 
4Cogger, H. G. & Heatwole, H. (1981) pp. 1333-1373, In. 
Keast, A. (Ed.), “Ecological Biogeography of Australia” 
(Junk, The Hague). 

5Storr, G. M. (1990) Rec. West. Aust. Mus. 114, 547-552. 
°Crocker, R. L. & Wood, J. G. (1947) Trans. R. Soc. S. 
Aust. 71, 91-136. 

7Schwaner, T. D., Edwards, A. & Miller, B. (1985) Trans. 
R. Soc. S. Aust. 109, 55-56. 


unexpected occurrence of a very familiar species, underscoring 
our ignorance at a fundamental level about the Australian 
fauna. 

Adrienne Edwards provided data on the distribution of 7° 
scincoides in South Australia. Mark Hutchinson made helpful 
comments on the manuscript. Jenny Wendelbourne and the 
people of the Mimili and Pukatja communities are thanked 
for their hospitality. 


8Cogger, H. G. (1986) “Reptiles and Amphibians of 
Australia.” 4th edn (Reed, Sydney). 

°N. Yanima, pers. comm. 1989, 

‘OMitchell, F. J. (1950) Rec. S. Aust. Mus. 9, 275-308. 

"Shea, G. M. & Peterson, M. (1981) Aust. J. Herp. 1, 27-28. 

2Lietzke, S. & Davis, M. (1987) “A Basic 
Pitjantjatjara/Yankunytjatjara to English Dictionary”. 
(Institute for Aboriginal Development, Alice Springs). 

Martin, K. (1975) Herpetofauna 7, 6-7. 

4Pianka, E. R. (1972) Copeia 1972, 127-145. 


G. R. JOHNSTON, School of Biological Sciences, Flinders University of South Australia, G.P.O. Box 2100, Adelaide, 


S. Aust. 5001, 


A NOTE ON PHASCOLOSOMA TURNERAE RICE (SIPUNCULA) 


BY S. J. EDMONDS 


Summary 


Transactions of the Roval Society of S. Aust. (992), 16(4), 


BRIEF COMMUNICATION 


ISI. 


A NOTE ON PHASCOLOSOMA TURNERAE RICE (SIPUNCULA) 


Phascolosoma turnerae Rice, 985 was described from 104 
specimens of a sipunculan found boring into submerged wood 
and was collected at depths of 1135-1184 m in the Straits of 
Florida, south of Key West. U.S.A. and also from 366-4]2 m 
in the northern area of the Gulf of Mexico, Alabama, 
U,S.A.' This record is the first of a sipunculan living and 
boring in wood. Po turnerae is distinguished from other 
congeners by the shape of its introvert hooks (Fig. 1). and 
the structure of its body papillae. 

Phascolosome kapalun Edmonds, 1985 was described from 
three specimens dredged at 710 m_ off Sydney, N.SW.. 
Australia during a cruise of he “Kapala” in 1977". Edmonds 
in 1985 was unaware oF Rice’s 1985 species, On comparing 
material of the two species m 1988. it became clear that P. 
nrnerag and P kapalum were conspecific, the latter being 
a junior synonym, (The date of publication af P lurnerae was 
20 March 1985 and that of PB kepalin was 28 June 1985). 
No wood. however. was associated with the “Kapala™ 
specimens nor did the collection records report the presence 
of any atthe time of collection. It seems probable. then, that 
the speciniens had been dislodged either during dredging or 
sorting, 

Recently a single specimen of P mmnerve was found in some 
material sent for identification rom the Northern Territory 
Museum, Darwin. The specimen (NTM WS87) was callected 
during trawling operations of “SOELA’ in Queensland waters 
(7°S9.2'S — 17°55.8'S, 47°04,5'E = 147°01,5'E) at 
259-260 m by H, Larson, 161.1986. The collection label 


"Rice, M. E. (1985) Proc, Biol. Sou. Washington 98(1), 54-60, 


as ah 


Fig. J. Phascolusoma nirnerae, introvert: hooks fram 


Queensland specimen (Scale line = 0.05 mm). 


reports thai the specimen was collected from “a piece of rotting 
wood" and a piece of the wood was included in the collecting 
lube along with the sipunculan. 

The purpose of the present note is threefold: 1. ta record 
(he synonymy of P furnerce und P. kapatum, the former name 
having priority. 2. to confirm that P turnerue is associated 
wilh submerged wood und 3. to record the wide distribution 
oF P turnerae now reported from the Atlantic Ocean (Straits 
vf Florida and the Gulf of Mexico) and the south-west Pacitic 
Ocean (off Sydney and off the Great Barrier Reef. Australia). 


"Edmonds, S. J. (1985) Trans. R. Soe, S. Aust. 109,(2), 
43-44. 


S.J, EDMONDS, South Australian Museum, North Terrace. Adelaide. S. Aust. 5000. 


EARLY HOLOCENE FROGS FROM THE TANTANOOLA CAVE, 
SOUTH AUSTRALIA 


BY MICHAEL J. TYLER, FRED W. ASLIN AND SIMON BRYARS 


Summary 


Transactions af the Reval Society ef 8. Aust, (1992), 106(4), 153 


BRIEF COMMUNICATION 


EARLY HOLOCENE FROGS FROM THE TANTANOOLA CAVE, SOUTH AUSTRALIA 


Five extant species of frogs are. represented in the 
Quaternary record uf the southeast of South Australia! 
constituting one of the best known components of the 
Australian Qualjernary frog fauna, Here we report a further 
site and add one more species. 

In 1982.4 quantity of fossil vertebrates and invertebrates 
was recovered from a pocket of sandy clay fill at the entrance 
to the ‘luntanoola Tourist Cave at Tantanoola, S.A, The 
foulerial was uncovered in the process of excavation of the 
Noor of the-entrance to permit wheelchair accesy. Included 
in the vertebrate material were 185 frog ilia. Here we report 
the identity of the ilia and place them in the contex( of the 
Australian Quaternary record. 


rP=0,92 
v= 10,454 16Sx 


Snout-Vent Length (mm} 


(lium Length (mm) 


Fig. 1. Length of ilia of Cimnadynestes rasmaniensix plotted 
against snout to vent length. Assumed snout to venr length 
of largest and smallest representatives of the fissil material 
indicued by broken lines. t-value for slope 10.395, p<0,001, 
For x = 186, y = 41.1 (95% confidence limits = 
37.8-44.4). For x = 10.6, ¥ = 27.9 (95% confidence limius 
= 25.4-30,4). OF the 2) complete ilia in the saniple x = 
3.6 mm, $.D. = + 17, range 106-186 mm, median 
{3.4 mm. 


Five species were collected at Tantanoola. They, and the 
quantities invelved are listed in Table 1, From the maximum 
nuaober of left or of right ilia in each sample it is apparent 
that a minimum total of 100 individual specimens is included, 
The very large number of Liraendvnastes tasmaniensis 
Gunther recovered permits an accurate extrapolation of the 
size of the individuals compared with modern representatives 


"Tyler, M. J. (1977) Trans, R. Soc. S. Aust, IOL (3), 85-89. 
“Tyler, M. J. (1989) “Australian Frogs.” (Viking, O'Neil, 
Melbouriie), 


of the species (Fig. 1), The size tanges of modern individuals 
are 31,1-39.5 mm (males) and 32.0-47,2 mm (females), 
hence the fossil material clearly is comparable in size. 

The faunal composition is almost identical 1 that 
represented at Victuria Cave and Henschke’s Quarry Cave near 
Natacoorte, $.A,!  Lireria ewingi, Limnodynastes 
tasmaniensis, L. dumertlii and Crinia signifera are common 
tw the three sites. A single Gevcrinia laevis from Victoria Cave 
is not represented at Tantanoola, whilst a single Neobarruchus 
pictus at Tantangola is not ceprestnied at the other sites, so 
increasing ro six the number of taxa in the fossil record of 
the southeast. 


TABLE |: The free ilia recovered at Tantanvola Caver 


Species Toial Lefi Right Registration 
tha tia iia Numbers 
Re ee 
Limnodynastes 137 66 71 P32111, 
lasmnaniensis P32239 
Limnodynastes 2 | 1 P32237 
dumeritii 
Crinia 42 24 18 P32112, 
siynifera P32240 
Necbatrachus i ] 0 P3224] 
bicias ; 
Lirarta 2 0 2 P32238 
ewiigi 
unidentifiable 1 0 I 
oe 
‘Totals 185 92 93 


— 
“All specimens are deposited in the South Australian Museum, 


Tt has becn suggested that the accumulation of so many frogs 
in cave deposits dues not reiJect the use of caves as diurnal 
or seasonyl refuges, but rather is a consequence of the 
Jisgurgement of pelletx by owls which ate predators of 
frogs’. 


The age of the material as determined by <!" dating. of 
charcoal is 9860 + 190 yours BP. The analysis was 
undertaken by Beta Analytic Inu. (Beta reference S401; 
FW.A. “Area 2”). 


We ure indebted to J. Callaghan and J. Aslin for their 
assistance during the excavation and collection of the materia! 
reported here. The participation of S. Bryars was made 
possible by an Australian Research Council grant to Mb 
Tyler. The excavation was undertaken under Permit 636 
granted hy the National Parks and Wildlite Service ty F. W, 
Astin, and the C% daung wax funded bv the N.PW.S. 


‘Tyler, M. J. (1978) “Amphibians of South Australis.” 
(Handbouks Commitice, Adelaide). 


MICHAEL. J. TYLER, Dept of Zoology. University of Adelaide, Rox 498 GPO, Adelaide, § Aust. 5001. FRED W, ASLIN, 
29 Bhabeth Street, Mb Gambicr, S. Aust 5290 anil SIMON BRYARS. Dept of Zoology, Laiversity of Adelaide, Box 498 


GPO. Adelaide. & Aust. SOOL 


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OFFICERS FOR 1992-93 


President: 
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