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of the 

Queensland Museum 


24 december, 1990 



Anstruci *..,.,. •*.!«•. ...,.•. .,,.... 1 

Introduction 1 

Abbreviations Used , . 2 

Material 2 

Previous Studies . 3 

Distribution, Habitat, Ecology and 

Conservation 4 

Patterns of Variation ....... 5 

Svstematics 14 



Ngairea gen. nov. 14 

N. domgoensis 16 

N. levicostata sp. nov. . . . . . 23 

N, murphyi . - . 26 

N. canaliculata sp. nov 27 

N. corticicola 3 1 

Mussonula 36 

M. verax 38 

Af. fallax sp. nov. 39 

Hedleyoconcha 42 

H. delta \ 43 

H. ailakeioae sp. nov 48 

H. addita 51 

Lenwebhia gen. nov '. . 52 

L. protoscrohiculata sp. nov 52 

Setomedea 57 

S. seticostata 59 

5. nudicostaia sp. nov 64 

S, janae sp. nov 69 

S. monteithi sp. nov 72 

S, sp 77 

Gyrocochlea 77 

G. vinitincta , 79 

G. convoluta 86 

G. paucilamellata sp.nov 87 

G. curtisiana * 90 

Riomphalopa gen. nov 94 

B. recava 95 

B. concinna 98 

Nautiliropa gen. nov. 101 

A r . omicron 102 

Letomola 109 

L contortus 110 

Rhophodon 1 14 

ft peregrinus 1 20 

R. consobrinus 121 

R. kempseyensis sp. nov. 125 

R, dizabethae sp. nov 129 

R. minutissimus sp. nov 132 

R. colmani sp, nov. 136 

R, bairnsdalensis 138 

Discocharopa 143 

D, aperta 143 

Cralopa , , 147 

C. siroudensis 150 

C. kaputarensis sp. nov 153 

C carlessi sp. nov 156 

Ehothera 160 

E. sericatula 161 

E. nautilodea 1 64 

E. gen'uhecata sp. nov. 166 

Coenocharopa gen. nov 169 

C, sordidus sp. nov 172 

C. macromphala sp nov, , . , 176 

C. alata sp. nov 179 

C. panicostata sp. nov 186 

C. yessabahensis sp. nov 187 

C. multiradiata sp. nov 190 

Egilomen 1 93 

E, cochlidium 193 

E. globosa sp. nov 197 

Omphalowpa gen. nov 198 

O. varicosa sp. nov 201 

Sinployea f , , , 205 

S. intensa 206 


Rotacharopa gen. nov 210 

R. annabelli sp. nov 211 

R. kessneri sp. nov 213 

R. densilamellata sp. nov 219 

Extralimital Relationships 223 

Phytogeny 226 

Biogeography 227 

Conclusions 234 

Acknowledgements 235 

Literature Cited 235 

Appendix . - 238 



Stanisic, J. Wfl 12 24: Syslemaiics and biogeography of eastern Australian Charopidae 
(Mollusca, Pulmonale) from subtropical rainforests. Memoirs of the Oueensland 
Museum. 30(1); 1-241, Brisbane. ISSN 0070,8835. 

Fifty 'and snail species in 18 genera of Charopidae from eastern coastal Australia are 
reviewed. Ngairea.Letnvebbia.Omphaloropu.Suutiiirapa, Coe/ioLharopa.Biomphalopa 
and Rotacharopa are new. Egilodonta Iredale. I$37 is placed in synonymy with 
Rhoplwdon Medley. L924. Ngatrea levieostam. N, cundliculata. Musvmula fallax. 
HeiJleyoconcha ailakctoae, Lenwehhia protascrobieulata. Setomedea monteitlu. S. 
nudivostata.S. janae, Gyrocochtea paucilamellata, Rhophodon momttssimus, R. eolmani, 
R, elizabefhae, R. kempseyensis. Cralopa carlcssi, C. kaputarensts. Egilomen globosa. 
Elsotheragenithecata.Ompfwhropavancosa.Coenncharopa sordtdin.C. maern/tiphufa. 
C- yessabahensis, C. parvicostata, C. multhadiata. C, alata. Rotacharopa autiuhcflt, R. 
L\sticri R. densilamellata (27 species) are new. Neot\pes are nominated for Helix 
murphyi Cox ; 1864, Helix omicron Pfeiffer, 1851 and Rhophodon perggrinus Medley, 
1924. All except 3 species are assigned to the Charoptftae; those 3 species, assigned to the 
Rotadiseinae. are the first record of the subfamily in Australia. 

Analysis of shell and anatomical patterns is presented: differences in shell, radular. pallial 
and genital struclures are evaluated. Patterns of variation are correlated with three levels 
of functional and ecological significance - species recognition, exploitation Ol adaplaiive 
/ones, and the need to conserve water. Morphological patterns oi Australian species are 
compared with those of New Zealand and the Pacific Islands and their relationships are 

The proposed phytogeny identifies major morphological trends and multiple origins of the 
fauna. A cludistic analysis (PAUP) utilises 75 shell and anatomical characters. A 
hiogeographic synthesis integrates distributions, ranges, and ecology with aspects of pas'. 
climatic- geologic and vegetational change. Charopici feVohltior] In Mfiteff] Australia is 
closely linked to climate induced changes in mesic communities since the Cretaceous. 
U Mollusca, Pulmonata. Charopidae, Systematics, Phylageny, BlQgeogr&phy* 

John Stanisic, Queensland Museum. PO Box 300, South Brisbane, Queensland 4101, 
Australia; 10 October 1989. 

The Charopidae is a family of small to minute, undescribed species from this region. Collecting 

radially ribbed, pulmonate land snails abundant by the author has exposed extensive radiations 

in leaf litter of eastern Australian rainforests. A of charopids in central Queensland mesic forests 

I cw species also occur in drier forests throughout and on the mountain peaks of the Ingham-Cook- 

tbe rest of the continent.Burch (1976), based on town rainforest massif in tropical Oueensland 

the earlier checklists of Iredale (1937a, 1937b. (Stanisic. 1987). Previously, the Charopidae 

1941a. 1941b), listed 1 10 species and 29 genera were regarded as a mainly cool temperate group 

of Charopidae from Australia. with greatest diversity in Victorian and Tas- 

The Australian Museum/Queensland Museum manian forests, 
east coast rainforest surveys of the mid-1970 s In general, eastern Australian charopids dis- 
accorded 3$ charopid species (28 new) from play an intimate ecological bond with ram- 
northern New South Wales, southern and central forests. 

Oueensland (Broadbent and Clark, 1976). Sur- Most of the SO species (18 genera) reviewed 

vcys by the Oueensland and Australian have subtropical distributions although it has 

Museums (198 1-1983) in subtropical rainforests been necessary to include several temperate and 

Of northern New South Wales and southern tropical taxa. The aims are to initiate a survey of 

Queensland revealed an even greater number of the subtropical Charopidae, to identify patterns 


of conchological and anatomical variation, and 
to place them in biogeographical context. The 
species reviewed here may represent only 15% 
of Australian charopids. Therefore 
phylogenetic conclusions are based on the 
analysis of a seemingly mosaic pattern of varia- 

Subtropical eastern Australia is defined as the 
coastal to subcoastal area between Dorrigo, New 
South Wales and Rockhampton, Queensland. 
Floristically, this area encompasses the distribu- 
tion of dry and moist subtropical closed forests 
(Fig- 1). 



AM Australian Museum, Sydney 

BMNH British Museum (Natural History), London 

FMNH Field Museum of Natural History. Chicago 

NMV Museum of Victoria, Melbourne 

SAM South Australian Museum. Adelaide 

QM Queensland Museum. Brisbane 

Collectors/Collection Data 

AM/QM-ABRS: Australian Biological 
Resources Survey funded field work (1981- 
1983) involving various collectors from the 
Australian and Queensland Museum. 

Earthwatch/QM: American Earthwatch spon- 
sored fieldwork involving volunteers and 
various staff of the Queensland Museum. 

Vegetation Types 

CNVF, complex notophyll vine forest; NVF, 
notophyll vine forest; MVF, microphyll vine 
forest; CMVF, complex mesophyll vine forest; 
SEVT, semi-evergreen vine thicket; SMVFF, 
simple microphyll vine fern forest; SMVFT, 
simple microphyll vine fern thicket. 

Anatomical Terminology 

DG, prostate; E, epiphallus; EC, epiphallic 
caecum; EF, epiphallic flagellum; EP, epiphallic 
pore; EPP, epiphallic pilaster; ERC, epiphallic 
retractor caecum; G, ovotestis; GD, 
hermaphroditic duct; GG, albumen gland; GT, 
talon; H\ heart; HG, hindgut; HV, principal pul- 
monary vein; I, intestine; K, kidney; KD, ureter; 
KX, ureteric pore; MA, mantle lobe; MC, mantle 
collar; MG, mantle gland; P, penis, PP, penial 
pilaster; PPM, main penial pilaster; PPT, prepu- 
tial tube; PRM, penial retractor muscle; PV, 
penial verge; S, spermatheca; SP, sper- 
matophore; SS, spermathecal stalk; UT, uterus, 
UV free oviduct; UVP, free oviduct pilaster; V, 



FIG. 1. Distribution of selected rainforest tloristic 
regions, provinces and areas in eastern Australia, a, 
humid tropica] northern region; b, subtropial forests 
of the Eungella area; c, dry to moist subtropical 
province: d, warm temperate to cool temperate 
province; e. temperate forests of the Lamington 
Plateau; f. cool temperate province; g, dry subtropi- 
cal province. (Adapted from Webb and Tracey, 

vagina; VC, vaginal caecum; VD, vas deferens; 
X, carrefour; Y, atrium; Z. digestive gland. 


This study is based on 6230 specimens mainly 
in the Queensland Museum, Brisbane and the 
Australian Museum, Sydney. Most specimens 
have been collected since 1980. 'Wet' material 
available was <10% of the total specimens; 150 
specimens (44 species) were dissected. 

The material contains a high proportion of 
juvenile and subadult specimens because in the 
early stages, when species presence was being 
established, most specimens were obtained by 
extraction from collected litter. This material 
consisted of a large number of dead juveniles 
which reflects the overall higher field mortality 
among young snails. Later collecting was con- 
centrated on live adults for dissection; so *wet' 




i ■ o \ M 

.-.I _. K-.- •• i 
W --*%? '^f- 

' ^ -J? ) 

i . ,' \_,Cr-^.— - - i .■,,-, ■ 

i ..... : . o 

. f ' 

3 }-- 

I 10.2. Distribution of major rainforest areas in north- 
ern New South Wales and southern Queensland. 

material includes a greater proportion of adults. 
However, because many species arc difficult to 
locate live in the field, individual 'wet' lots often 
contain only single or few specimens. 

Seasonal climatic effects may be a contribut- 
ing factor to the numbers oi live adults versus 
live subadults present at any time, bat the small 
amount of wet material and uncveness of collect- 
ing effort through the year prevent sensible con- 
clusions; only a few species were abundant 
enough for seasonal studies. 

Line drawings of shells and dissections were 
made by the author using a camera lucida attach- 

Measurements arc recorded using the method 
hf Solem (J 976, pp. 11-15). 


Previous studies of Australian Charopidae 
have been introductions in broader faunal check- 
lists except for works of Hedley ( 1*324). Solem 
| 1984), Smith and Kershaw (1955) and Stanisic 
(1987) who dealt specifically with Australian 
chafopids, With the exception of Hedley ( 1 SW ), 
who figured the rudula and jau of HcJ- 
(rvoctttn fn: delta (Pfeiffcr. I857) 3 and Odhner 
(1917) who figured the jaw and raduia of 

Pilshr\c/iur(/pci tumidus (Odhner, 1917). 
malaeologisis of the laic 19th and early 20th 
century used conchological details to establish 
species and genera. Smith and Kershaw (1985) 
and Stanisic ( 1 987) provided anatomical details 
for several genera and species from the Tas- 
mania-Victoria and north Queensland areas, 

Iredale (1937a, 1937b) compiled records up to 
that date. Subsequent works introducing new 
species are Gabriel (1947). Gabriel and Mac- 
pherson (1947), Cotton (1939), Iredale (1937c, 
1939, 1941) and Stanisic (1987). 

Most early shell-based descriptions are of little 
value in contemporary systematics. Only Hed- 
ley (1912. 1924) set an acceptable standard. 
Solem (1983) set a modern standard with his 
review of Pacific Island taxa; he showed the 
importance of soft parts in species diagnoses. 


Solem ( 1 9S3, p. 47) presented a full synonymy 
of the family level units available for the en- 
dodonioid complex., and in the Charopidae in- 
cluded Flammulinidae Crosse. 1 894, 
Dipnciicidac Iredale, 1937 and Hedleyo- 
conchidac Iredale. 1942. These family names 
had previously been applied to various sub- 
divisions of the Australian Charopidae by 
Gabriel (1930) and Iredale (1937a. 194 Lb 
1942). Solem (19{J3, p. 70) recognised five sub- 
families: Otoconchinae Cockerel!, 1895, 
Rotadiscinac Baker. 1927. Charopinae Hullon. 
1S.S4. Semperodoninae Solem. 1983. and Truk- 
charopinac Solem. 1 983. The Charopinae was 
broadly defined and considered amenahlc to sub- 
division subsequent to a revision of Australian 
and New Zealand taxa. 

The phylogcny developed in this study utilises 
anatomical features to define subfamilial, 
generic and species units. Specifically, the 
structure of the pallia! cavity, gross morphology 
of the reproductive system, and internal charac- 
teristics o1 (he terminal genitalia are considered 
important criteria for phvloccny. Conchological 
detail such as shell shape, ribbing, and coiling 
pattern may be convergcni and often mask quite 
crucial differences in anatomy. However, some 
conchological features including apical sculp- 
ture, microsculpture and primary shell sculpture 
show more positive correlations with underlying 
anatomical patterns. Hence it has been possible 
to produce a practical classificatory system 
which utilises shell features. Because this intro- 
ductory siudy investigates selected species from 


an area of mid-range diversity Tor the family, 
phylogcnelie gtfpS between genera are nol 
'even'; they reflect the diverse ecological habits 
of species studied, 

Genera reviewed and. where possible, 
redefined using both shell and anatomical 
lures arc:- Cralopa, Discocharopa, f (sothera, 
(Ivrocoi-hictt, MussonulUi Rhophodon, Seta- 
medea, fledleyoeoncha, Egil&men, Letomola, 

F.y.ilftioiita is synonymiscd with Rhophodon* 


In MMHii Australia, charopids arc mainlv 
found in closed forests (= rainforests). Histofl 
cal development of this association is examined 
further in the biogeographic discussion. Die 
strength of this association, antiquity of the 
Chan md long-term historical persistence 

CSic fofflsts in eastern Australia, have com- 
bined to produce a complex pattern of narrow, 
wide, disjunct and relict distributions 

During arid phases of the mid to late Tertiary, 
rainfbrestS would have acted both as refugia and 
epicentres of charopid evolution- Today all 
Australian rainforests are rcfugla (Webb and 
Trace y. IV81). 

Charopids flourish in the four major categories 
ol rainforest refugia of Webb and Tracey 
-I }:- A. large, relatively wet areas sac h us the 
Border Ranges ol soul hern (Jneensland, the Ijot 
rigo Plateau, NSW. and mountain summits such 
as Ml Wai ning, NSW, and Mt Glorious, SFQ: B, 
small, relatively dry, topographic isolates which 
include rocky limestone outcrops such as the 
Cavt neai RoeWiampton, MFO, Mi B»gg 

den. SEQ, and the Kcttl] Caves complex, 
NSW - besides providing protection from fire 
and a relatively high level of moisture, these 
areas also offer an abundant supply of calcium; 
(_'. small edaphic isolates where soil types are 
critical in determining rainforest existence - gal- 
lerv lorests growing on riverine alluvia ami the 
deciduous forests of Cooloola. SEQ, are ex- 
amples, D. mountain tops in lower rainfall aieas 
w here topography, soils and climate combine lo 
ensure persistence of closed forests - examples 
lie Runya Mountains and areas of the Manv 
Peaks Range. SEQ and Ml Dryander, MFQ. In 
NO and MEQ charopids arc mainly restricted to 
montane regions indicating that past environ- 
mental changes have been effective in 

geographically cornering of species. This al- 
tiludinal factor is kss noticeable in the subtropi- 
cal region where both lowland and foothill 
inniorcsis have a rich complement of species. 

Although rainfofttSt provides ample litter con- 
taining a \anei\ ol living spaces through which 
snails am crawl, species investigated here do not 
exploit [he Inter /one. Hcdlcxoconcha is ar- 
boreal with H. delta living on the leaves of .trees 
and shrubs, tfgatrea and Sinplovca arc semiar- 
boreal preferring lo live under the bark oil alien, 
rotting trees. Lenwebbhi has been found sitting 
in the open on roiling logs while Setomedea is 
the only group known to prefer vcrj flamp places 
in o! under logs Living under logs is the 
preferred habit for Sautilnopa, Gyrococklea, 
Biomphalopo* Elsothcriti Howchuropa, Cralopa 

$froudCf\sis t C kaputarensis. and Egilomen 
fclokosa. Species of Rht>phodofu L&fdrrtola con 
tortus and Coenochawpu yessahakensis, live 
on rock surfaces and some (e.g. C. j 1 ssabahen* 
sis) show habit related radulur specialisation. 
Coeuoi hurojHi alutu lives in moss on logs and 
rocks, while C macrotnphala and C. sordidus 
prohahlv live among the friable earth below the 
litter, as do (httphalotopu vurtctsa a ml 
l.\>tlomi-n cachtidium, Micmbabilals of the 
other species are unknown. 

Feeding habits arc unknown. Radular mor- 
phology is conservative; rarely (e.g. in 
Ntnttiltntpit oiw< ron and Nyutrea corticicofa) is 
there anv structural indication of possible dietarv 

Sympalry he- been noted in Rhophodon, 
Xi/ittftn. GyrflCOCtllea arid ( oenocharopd. 
Miciosvmpairy has been observer! between 
Ngairca dorrigoe/tsi& and Ngoireti corticicolh 
with the CW V '-pecies having been collected from 
under the same piece of bark. These examples 
arc discussed m itiOrt detail m the systematic 
rev icw under the relevant raxonomic heading. 

Species are largety restricted to rainforest and 
in some instances e.g. Rhophodon elizahethac 
M\i\Srit>/m-<h{i ntttlteostata. are confined tovery 
small patches. In many cases, particularly m 
northern NSW and southern Queensland, clear- 
ing for agriculture and other human activity has 
drastically reduced lowland and upland rain- 
forest. Whether or not species will survive this 
synanthropic fragmentation of Habitat remains to 

be seen. 

Charopids me uSfeful indicator species of bush 
quality Areas where diversity is high, orwhere 
restricted species reside have special refugial 

■; 1 1 leaner 


TABLE 1. Mean shell measurements in 49 species of east Australian Charuptdne 





3 1 '2- 

4 1/8 

J 12- 

4 _^ 4- 

6 1:8+ 















: QE 




i ? 1 



s is 



26 J 


v | | ■ 

i i g l 














Fourteen species with reduced and irregular ribbing were omitted from these figures. 


NflH t 

Size and shape - Whorl count (Table 1) is 
usually i^?^ species) 4-5; four species have <4 
whorls while 12 species have 5 or more whorls. 
Higher whorl numbers occur in several different 
lineages. Ifedlevoconcha delta has the largest 
number oi whorls with 6 18+ average: 
Rotacharopa annahelli has 5 3/4- whOrl average 
(Figs 23a, 140a); Biomphalopa recava and 
Setomedea monteithi have mean whorl counts erf 
5 1/4-: Ngairea murphyi has mean of 5 1/2- 

Disiocharopa aperia with 3 1 i ''2- whorls has 
the lowest whorl count (Fig. 93a), while 
Suiplnxea inicnsa with 3 5/S+ whorl average 
(Fig. |37a) 8 is near the minimum mean whorl 
count for Pacific Basin Sinployea (Solern. 1983, 
p. 83). The highly modified Letomola contortus 
(Fig. 67a-c) has a mean whorl count of 3 7 8-. 

Species with low whorl counts tend to be small 
in size, but the opposite does not apply to species 
with high whorl counts. Of those species with 
average whorl count >5. six arc <3 mm in 
diameter. Only Hedleyoconcha delta. H. 
ailaketoae. and Ngairea murphyi have mean 
diameters which fall into the upper quartile 
range. Significantly these three species have 
strongly keeled body whorls. Hence whorl ad- 
dition does not appear to have been a major 
factor in attaining large size. 

The largest species is Gyrocochlea vtnaincta 
while Dtscoeharapa aperia is the smallest. Be- 

tween these extremes of shell diameter a sig- 
nificant number of species fall into the 1st to 2nd 
quartile range which covers about 27% of the 
total range of variation (Table 1 ). In contrast, the 
larger species show a much greater size range, 
suggesting that repeated evolutionary experi- 
ments in size increase occurred. This conclusion 
is further supported by the many different shell 
morphotypes represented among larger species. 

Nine of the 15 largest species belong to genera 
which have an incomplete secondary ureter and 
elevated, keeled shells. Only Lenwebbia 
protascrohteuhitih and Setomedea, which show- 
several secondary conehological and anatomical 
specialisations, have incomplete secondary 
ureters and more average-sized shells. Other 
large species belong lo Gvrocoehlea, 
Nautittropa, and Elsoihera. 

Conservatism in H,D ratios (Table 1) em- 
phasises stability of shell shape. Prominently 
high spired species, Ngairea levicostata (Fig. 
be ). Xfassonula vera.x (Fig, 1 8c K Hedleyoconcha 
delta (Fig, 23c) and H. ailaketoae (Fig, 6c) have 
an above average II. D ratio lhai is >0.70. High 
H/D ratio in Egilomen globosa is due to greater 
spire protrusion, closed umbilicus and sharp 
deflection of the body whorl (Figs 131a-c). 
whereas high whorl count, narrow umbilicus and 
more protruded spire produce an above average 
H/D ratio in Roiacharopu densilamellatu (Figs 
146a-c). In Ngairea and Mussnnula, spire 
protrusion and large body whorl width con- 
tribute to increased height of shell, while in 


fh'illcyactmcho delta, above average whorl 
count is a further mitigating factor, In contrast 

the height ot'Gyrocodileo. which has B i ongly 
depressed spire, is due lu a comlutiaiiou of large 
body whorl and dramatic deflection of the body 
whorl m ibt last third Of its volution 

Umbilical width varies from closed in sunn: 
species o!" Cralopu and FAsotfuru {Figs 101b, 

104b. 105b, 10Kb) to wnjciv open iji Om- 
pluxlompa* Discodtaropa and Rftophadon [Pigs 
71b. 74b, 77b, 78b 8lb»84bv87b,93b), Anonen 

umbilicus is most common: a narrow umbilicus 
occurs main!\ in groups with altered wlunl 0OT1 
tour. Hattexixoncha. Mussonala and Ngairea 
(except V mUrphyi) h.ivo vciv small umbilici 
andangulate to keeled body whorls, ft. murphyi 
has secondarily narrow '©d whorl* and mote open 
umbilicus (Fig. 9b). In Egilomcn ghbosa (Fig. 
I3la.b), Ratadwropa dvnsitetwlttita 1 1 ig 
I46a,b). ff. ke&tneri (Fig. I43a,b). and A*, atf- 
prA&c//i (I 'i: 140a, b) with tightly coiled shells, 
whorls are globose and expanded, Constricting 
umbilicus will) then inner margins. Umbili- 
cal contour varies from narrow U-shaped. 
through wider V-shaped [0 wide saucer-shaped 
in Rhophodon kemnseyensis (Fig, 78b) and Dn- 

I iuhumpit aptTiu (Tig. 93b). 

Whorl profiles range from laterally com- 
pressed in ( OVnOChuropQ ctli/hi and Rhi)phodnn 

haimsdalensis (Figs i He, B9c) lo strongly 

keeled iti X.'jarca mttrphyt and HcdkvtHontita 
tlrha. In l.t'totnola coniortus whorl profile is 
altered In a supraperiplicral sulcus (Fig. (Wc). A 
weak supraperipheral sulcus is also present in 
} Isotih-ni nuttltludiii (I [g, IDdc), and 
Cotvocbaropo mtttromphala (Vig 1 14c). 
Salutes are generally weakly impressed al- 
| igh 1 tfopci ttrOUaVHSis has a \er\ deep 
suture [Fig. ^5d), and GyrococJtlca vinititn tu 
(Fig. 45aJ and Us congeners initially havt a 
w eaklv impressed suture which becomes deeply 
r - OWcd aS [hC laSl ^horj descends A more 

unusual development in Nuaifea caHaRculata 

and klsoflwra gpnilheCPtA is a narrow channel at 
[lie sutmal aiea (Fig. 1 2a). Apcnurul sinuses ate 

do eloped in Cralopa and Elsothem* 

Stutpturt'SpS'ti apical sculpture is prominent 
in SinplnyoQ intense Qtnphakirftpa varieo&Q* 

broad spiral eordS (Ffg. I37d) While 0. cum. .'sd 
has the cords narrower and slightly higher (Fig. 
H4e) In Si'hmti'ilat wUCVSWv and S, iqtwi 1 
the) are legulail'. . ■. ■ . , ■ . 1 . 1 r- .,-, 1 1|0&£ Of 

' (/' trrua whereas in S". montvithi (Fig, 4 lc|. 

-S nthhmstaia (Fig. 35e), and Sctomcifca sp. 
(| : j^.44e), the cords become squiggly, and vague 
curved radial ridges develop at the nuelear- 
posinuclear boundary. Radial elemenis on (he 
latter part of the protoconch are more 
pronounced in Mu^sonuiu ivn/v Iredale, [937 
(Fig, lSd) and \f Jalta.x (Fig. 20f). In 
Gyrocoditea pauctlam&ltata apical sculpture is 
Inn l\ reinailau with spiral and radial elements 
equally prominent (Fig. 51c) whereas in O'. 
vfttnittclir (1 ig, i**t ) and (7. mnvoUita (Fig 4Se) 
the radials are low weak ndges which are raised 
at their intersection w iih the spiral cords lo form 
tiny beads. In contrast. G. cunisiana (Fig. 64a) 
has apical sculptuie w hich initiallv is a series pi 
regularly arranged "pits" then becoming ir 
regularly, radially ribbed toward the end of the 
protoconch. Bfampftalopa recava (Fig. 57c), R. 

cuncinna dig 6DfcJ and \uutilimpa omicron 
(Fig. 63d) have simiiat apical pitting. A dif- 
ferent paiieru ol apical pits in tli'dfcYoa>w:ha 

/ i I ... li epci and more crowded depressions 
(i ig 23e) I ven in this case, the vague radial 
pattern ol pits indicates probable derivation from 
u typical reticulate pattern. f.oiHchhia 
(>!(>U)SL'uhu ulutit lias a similar bui finer pallcrn 
(Fig, 29g), 1)1 Ngairtia murphy] irregular radial 
wrinkles and shallow irregularly shaped pits 
eomhme(l Ig '") apparenll) j modification of 
radial ribs md spiral groove* seen in .V. tor- 

lickofaH o\. iSodlandA !i\ir<>\ltilii (I igsbd, 

15c). LpJamola contartus, has apical sculpture 
like thar <)\' \. warphxt bui w nh an underlying 
radial pattern which is fess conspicuous, produc- 
ing -i rtiiiliealc surlaCc (Fig 67d)< 

Spiral grooves on the protoconch are rare oc- 
curring in S^itttcu inrnacoht (Fig. l>e), N- 

mdliciifataiFigi 1 21). and to u lesser extent .V. 

tCQstata (rig. 6d). V. (tflrrlfiO&ittU has 

predominantly radial apical sculpture with very 

fine Spiral lines visible in the suinral area (lig 

Apical sculpture inosi comruonl) consists ol' 
prominent radial ribs anil less prominent spiral 
coids. Radials mav be crowded i!i\o/hrra, 
f. rafopa, Discochawpa), widely spaced 
I ilanu n) oi w idely spaced Initially, becoming 
more crowded toward the nuclear-poslnueleai 
junction {Rhn()luul<»i ( ni-nt)<h<trop<i). Spirals 
maj bi coDtinuous narrow, squiggly cords 
(( fiflopti. f-lsolluru, C of/uH'huropit), short, 
wrinkle-like cordlets {Rhophodon, DtS 

tochwoptfltOi inconspicuous (I gilomen), If the 

coids jic .oniiiinou-, dic\ are usually visible on 
the apices o[ the radial ribs. 


Apu nl sculpture til Roiucharopa appears 
reticulate under the light microscope hm hi: | 
complex under the SEM The radials are a series 
Of CUJVcd tidges which become broader u>-, . 
the nuelear-postnuelear sculpture (Fig. J4od). 

Spirals, while appearing contiguous, arc a series 
of short segments (Figs 140c. 146c). This apical 
sculpiure was also reported in liticrochtiropa 
mmtuluSotew (1983. p. 70). 

Poslnuclcar sculpture involves variations on 
the pattern of major radial ribs and microsculp- 
ii.e of micmradials and km spiral cords. Majoi 
departures are seen in species with an incomplete 
secondary ureter. Pcriostracal enhancements 
may he present and take the form of high blades 
on the major radials as seen in Sauiiliropa 
(micron (Tig. 63c) or the expanded peripheral 

; nsions o\ Cocnnchtjropa alata (Fig, 1 17edi), 
A more significant change takes place in 
Setomedea which has pcriostracal setae along 
ihc length of the major ribs (Figs 321 3$i\ 4 It »'. 

Major radial ribs may be broad and high as in 
Hiotnphalopa recant (Fig. 5~\\ Gyrocochica 
linitincta (Fig. 450. Eplomen cochlidium (Fig 
I28g) and Rlutp/soda/t p&tgfttoUS (Fig. 7 I f ). 
have a high pcriostracal blade as in Nautiliropa 
omtcron. Ompriuloropa varicosity and 
Rotacharopa (Figs 63e, 1341. 1401), or be 
reduced with a low pcriostracal blade expanded 
01 the shell periphery as in Coettocharopa alaia 
(Fig. I I7g). andC. panicosiata (Fig. 120d). In 
Mussonula verax (Fig. \Hc),M.fallax(F\g, 20g), 
and N^airca levtcostata (Fig. 6c). the pcriostra- 
cal blade is high and continuous above a tCW 
radial Tib. However, the blade is not vertical as 
in O. vahcosa. but instead is folded back toward 
the shall apejt, in Setomcdea the major ribs have 
two pcriostracal blades (Figs 321, 3S1') Rib 
height reduction is seen in Lenwebbia 
protest ratricufata which has very low radial Ufl 
dotations on adult whorls (Fig. 2s>f), Ngaitea 
corticicola which has widely spaced low, radial 
undulations with a deciduous pcriostracal blade 
(Fig. 15f)i Iclomola contains which has an 
overlapping shingle like arrangement of radial 
thickenings (Fig. 67c), and Coenocharopa yes- 
.,lu'n\i.\ which has very low, crowded radial 
fibs (Fig. I23g). Ngairea murphy i (Fig. 9g), v 
dprrigoensis (Fig. 3e). HcdUyoconcha rftfftfl 
(Fig. 230 and H. ailaketoac (Fig. 2nd) have 
crowded, reduced, and thread-like major ribs to 
ihc exclusion of mieroradials. 

M. .slave regular radial ribbing -\n 

lattcTTl is shown by Biompbafapa 

recava (Fig, ?1<\) and Setar 

(Fig. 3 2d). Reduction and mere-use in rib counts 
occur in both small and large \ t| several 

genera. Widely spaced ribs of filtoph, 
huitn\(ltu\'n\t<, (Fig. 89a), Egtlum&n cochhdium 
(Figs IZSa.e) and uvrocochlea pattcilatnellaui 
(Tig 51a}. crowded ribs oi' Biomphalopa concin- 
na (Fig. Mia.d), Rhophodon mututtsstmus (Fig. 
S4a), and Roiacharopa kessneri (Fig. 14 1 
Ciyrocochlcu cumsuma (Fig. 54a), FAsoth 
natitthnlcti (hg H)oa) and Kotacharopa dert- 
sitatnetlata tFig. 146a), and the exceptional 
Situation jn Rttophodon kcmpscycnsis which has 
a ribcount ol 27°-. 7 (1 -ig. 77a), are examples of 
the variation encountered. Species of 
Co&tQchdrcpu (wiih ihe exception of ( 
sahdftctms) have reduced rib counts. However, 
the deciduous. penoslracal naluu 6) Ihe rtl 
cOmbinfd with the worn slate of most of Ihc 
available material, made rib counts in ft 
species difficult. 

Reduction in primai \ sculpture may 
therefore involve reduction in rib counts through 
increased rib spacm;^, as well as a tcdm-hon |fl 
tht height ol the major ribs. 

Mierosculptutc i\pically consists of h 
crowded radial riblcts and low. narrow crowded 
spiral cords buttressing the radials on either side 

e.g. Elsothcra .uricatula ( Fig. I04f), 
Gyrocochica viniiinciu (Fig. 451). Binmphatopa 
conciuna (Fig. 60f) and Crulopa strouttc; 
<Fig. 95c). Rhophodon 3nd Rotacharopa have 
nigh micron [dials bu I very low broad spiral cords 
with small elongate beads formed at their inter- 
section (Figs 71 f. 89s< 143f. 14of); Rhophodtm 
kempsi'\ c/n is (Fig. 7 7 ■- i - and Discocha n tpa 
p_ ihe mieroradials divided 
into a scries of short segments; Sinployea interna 
tlredale, |<H] 'h:r. die nucroradialsmore thread- 
like and spiral cords conspicuous with elongate 
heads ;■' 'erections (Fig. 137c); am: 

Egitomen cochiuihun (Fig. 128g) and/f. globosa 
(Fig. 13 If), have spirals more conspicuous than 
mieroradials. This lattercondition istakenastep 
further in Qmphciiorof.m sartcosa whetC 
mieroradials arc very low and microspirals torn- 
large beads at Ihe intersection such thai intersti- 
tial areas appear to be sculptured with spirally 
arranged beads I T g 134f|, Reduction in iht; 
height of mic rotudials also occurs in 
Coenocharopa, Ilov. lie mtcroradiul clt- 

ment attains height by the addition of fine thread 
like pcriostracal blades e.g. C. alata (Fig. 1 I7g). 
Mieroradials mav be higher, equal in height or 
low i than lh€ mfcrosplral uwJs, b\t\ then; are no 
beads formed and there is no buttressing. 


More complex departures from the above pat- 
terns of microsculpture are seen in species with 
an incomplete secondary ureter. Sctomedea has 
conspicuous microradials which are higher than 
the thread-like microspirals, and although the 
microspirals continue up the sides of the 
microradials there is no buttressing (Figs 32f, 
35f)- S. janae has microradials with a scalloped 
profile (Fig. 38g) as a result of the microsculp- 
ture dipping into spiral furrows in the shell sur- 
face. These undulations are also developed in 5. 
monteithi and Setomedea sp. Lenwebbia 
protoscrobiculata has only microspiral grooves 
on the early part of adult whorls, but a combina- 
tion of fine thread-like microradials and grooves 
on the latter part of the body whorl (Fig. 29e.f)- 
Spiral grooves are present on shells of Ngairea 
corticicola (Fig. 15f) and N. canaliculata (Fig. 
12g). On the other hand N. levicostata has 
microsculpture of fine spiral lines (Fig. 6c) sim- 
ilar to Mussoniila fallax (Fig. 20g). M. verax 
appears to lack the fine spiral lines of M. fallax. 
but this needs to be confirmed. Ngairea dor- 
rigoensis (Fig. 3e) and jV. murphyi (Fig. 9a, e) 
lack microradials, but have many crowded, low, 
thread-like radials crossed by crowded thread- 
like microspirals. Low reticulate sculpture also 
occurs in Hedleyoconcha delta (Fig. 23f) and H. 
ailaketoae (Fig. 26d), where spiral cords and 
radials are fused. Hedleyoconcha exhibits an 
unusual growth-related change in sculplure; 
after about 4 1/2 whorls the reticulate pattern is 
replaced by a much less sculptured pattern of 
irregularly spaced radial growth ridges (Figs 

Apertural Barriers - Apertural barriers were 
recorded in Letomola contortus and Rhophodon. 
However, barrier formation is even more 
widespread among east-coast subtropical 
charopids. Several undescribed taxa with bar- 
riers live between Gympie and Rockhampton; 
until these are described, discussions on barrier 
formation will remain, skeletal. 

The form and surface characteristics of bar- 
riers in Letomola contortus separate them from 
those in Rhophodon. Without exception 
Rhophodon spp. have numerous parietal and 
palatal barriers whose overall effect is to reduce 
the apertural area. In contrast L. contortus has 
only a single, thin blade-like parietal barrier (Fig. 
68a), no barriers on the outer lip margin, and a 
low ridge-like basal palatal (Fig. 68b). Further- 
more, the barrier surface in L contortus has 
small rounded protrusions (Fig. 68c) whereas 
Rhophodon has narrow to broad transverse 

ridees angling up from the barrier surface (Figs 
78c, 81d»84d)< R. kempseyensis (Fig. 78c), and 
R. minutissimus (Fig. 84d), display extremes of 
variation. These basic differences indicate separate 
derivation of the barriers in the two genera. 

!n Rhophodon the number, size and shape of 
barriers as well as surface sculpture are variable. 
Generally the parietal barriers are more blade- 
like in form than the shorter crescent-shaped 
palatal barriers. The number of parietal barriers 
is least in R. bairnsdalensis which has a single 
elongate barrier (Fig. 90a), and ranges to the four 
barriers and two traces present in R. peregrinus 
(Fig. 7lg). Typically there are three blade-like 
parictals - a horizontal lamella about halfway 
down the whorl margin, and two on the lower 
part of the parietal wall, deflected downwards. 
R. elizabethae has only two parietal lamellae but 
has a well-developed, low, blade-like superior 
parietal trace. In R. peregrinus the first parietal 
is deflected upwards while the second, third and 
fourth are directed downwards (Fig. 71g). An 
accessory parietal trace is also present in R. 
consobrinus (Fig. 74h). Usually the parietal bar- 
riers have a long anterior taper projecting beyond 
the edge of the aperture and an expanded 
posterior section with an abruptly tapered 
resorption edge. R. peregrinus has recessed 
parietals which have shortened or no anterior 
tapers (Fig. 71 g). A similar shortening is seen in 
the deeply recessed second parietal o\R. minutis- 
simus (Fig. 84g). The only other notable varia- 
tion is in R. colmani which has the parietals 
greatly swollen, particularly at the posterior 
edge, thus further reducing the apertural opening 
(Fig. 87f). 

Columellar barriers range from well deve- 
loped, horizontal lamellae in R. elizabethae and 
R. colmani (Fig. 87h) to fine, almost thread-like 
in R. peregrinus and R. consobrinus. 

Palatal barriers vary from two in R. 
bairnsdalensis (Fig. 90b,c) to six in R. colmani 
(Fig. 87g). A superior palatal trace, situated very 
near to the palato-pan'etal margin, may also be 
present. The first palatal barrier is usually 
situated at the baso-columellar margin and is a 
short, high, crescent-shaped lamella. The shape 
in R. colmani is typical. Remaining palatal bar- 
riers are similar in shape though somewhat 
reduced in height. In R. peregrinus the upper 
palatals have a slightly elongate anterior taper 
(Fig. 71 h), while those in R. colmani have the 
anterior edge much more markedly tapered. The 
low thread-like upper palatal of R. bairnsdalen- 
sis (Fig. 90c) is a further variation. 



44 of 5IJ species were dissected. Species rtpl 
dissected arc MttYsunitlu \erax, Hedlcyoconchu 
ihiititfi. Rhophodon column t, DtSCQCjiarQpQ 
li/.-crtu, Elsolheru HUutdodcu. and Coetto- 
vnatopu multiradiata. Available material ol 
Mtutsonukt [alius and Ngaitcu tevicostata had 
immature reproductive systems which provided 
nc details on genital structures. Poor preserva- 
tion led in obtaining onlv fragmentary detail of 
the penis interior of Sinployea intensa and pall ial 
cavity of L'<K'tu>charopo purvtcosiata. 

Variations in the pallia! and genital anatomy 
are minor anil relied regional endemicity. With 
the Bxeeplion of Rotachawpa and the 
predominantly southern FJsitthcra the patterns 
are conservative. 

Ptilliul Curtly - Typically the pallial cavity 
has a well-developed mantle collar with mantle 
gland extending onto the paltial roof, a weakh 
hilobed kidney in which the pericardial lobe is 
vaguely triangular and much longer than the 
rectal lobe, and a complete secondary ureter with 
IhC u re lerie pore situated at the inner edge of the 
mantle collar adjacent to the anus. The bindvein 
is conspicuous, bul lacks prominent branching. 

A major departure from this pattern occurs in 
N gain:* a, Hedlevoconcha, l.emwhhtu, Mus- 
^oiudu, and Sciomcdcu which have an incom- 
plete secondary ureter (hat is a very short 
relieved lube with the ureteiie poic situated at 
the rear of the pallial cavity in the angle between 
(he kidney and the hindgut (Figs 7, Hie, 21, 24g, 
30c, 33g). Whether or not these taxa should be 
n separate snblarrnlial recognition is uneer- 

lifl and will remain so until more of the southern 
lava are dissected. 

la most species the primary ureter is a simple 
ihbe running from the kidnev apex to the rear of 
the pallial cavity. However, some taxa with an 
incomplete secondarj uretei have undergone 
Elongation of the primary Ureter This is m<<st 
obvious in Setomcdea which has the initial part 
of the primary ureter coded (Figs 33g. 3fe, 39f, 
42f) and in Lcnwchhut which has it con- 
spicuously retlexed (Fig. 30c). Less noticeable 
are Ihe subtle variations seen in \^uirca and 
MuFSikilik (Figs 4e, 13c, I6f, 21) where some 
degree Of reflexion also occurs. 

The kidnev is generally weakly bilobed with 
an elongate pericardial lobe. In Afgflirfifl mur- 
phyi I Fig M le 1 and some species of Rhophodon 
(Figs 27f, 3' le! the rectal lobe is almost entirely 
absent and the pericardial lobe is markedly elon- 
gates All these species have above average 

What C • ' , Hilobed kidneys with aln 
equal sized lobes are present m Egiiatnen (Figs 
I29d. 132Q, Stnplovcu nitcnsa (Fig, 1~<Kf) . rjtj 

Letoto fi ontprtus (Fig. b^g). Speeics 
Cralopa (Figs 9fle. 99c, 102c) have rhe rectal 

lobe abqul U3 the length of the pericardial lobe. 
A mantle gland is usually present itftbongh no 

gland wa- noiiceil in / Irdlrvtu nucha, Sinployea 
and some species of Qyro\ ocldcu. Nuutiltmpu. 

FAfQtherQ and f\ttluchuropu. 

own ■. 

Ovotestti - Usually the ovotestis consists ol 
CWft clumps of palmalcly clavatc to finger o1 alveoli embedded in the apical whorls 
of the digestive gland and oriented parallel to flic 
plane Of coiling. In species with an elevated 
spire such as those belonging to NgaiW£i i Hud- 
Icyoconcha and Lenwehhia the lobes are pi 
lionedal right angles (o the plane, of coiling (Figs 
5c lOg, 24e, 27d. 30c). Gyrocochlca vinitincta 
which hasagrcully enlarged shell with expanded 
whor ilsn has the ovotestis more at right 
angles than parallel (Fig. 46c ). 

Althougl lamps ol alveoli are normally 

present, Mime proliferai ion of cl m tips takes place 
in Ihe high spired and high whorled Ngpuren 
wurphyi I Fig. Hid), and Hcd!c\oc(tncha delta 
(Fig. 24c). Single clumps arc present in the tiny 
Sittplf.-'.'cti utiensu (Fit:. I3Xc) and / cionuda frjfl 
: ig. b^c) while according to Solem 
i WB3. p. 74) a single clump is present in the 
i Qcharopaaperta. Within a clump of 

the uvotesos the number ol alveolar lobes v. 
from many in some of llie larget species to the 
simple bilobed condition In the very smalt f)m 
phalnrcpu \>atlcQ$& (Fig. 135c), Rhophodon. 
batrnsdalenstt (Fig, 9\S) and /. nmtorms I I 
69d). In Roiachuropa the ovotestis consists ol 
two teardrop-shaped lobes (Figs I4lc, I44e. 

1 47c i representing a significant difference from 

l< niple clavatc alveoli in rhe Charopinae. 

Hermaphroditic Dua - Mom species in- 
tigatcd have a hermaphroditic duct which is a 
simple tube leading apically from the carrefour 
region; along the paneial W.bOfl margin to ihe 
ovotestis located in the apical whorls of Ihe 
digestive gland. In several species ihe tube ' v 
partially kinked. This is best illustrated in 
species ol ftcdlc\(>< nncha ( f ig% 24(1, 27e i which 

have elevated, trochoids sueiis. and also in 

Neutral dtn 'sy.i'i nsis and GyrQOOCl 
vitnttfh'in | 1 igfi -Id. 4bdl. Differences in tilt 
grcc of duel expansion, noted in some dissec- 

nons. .re m-.-s! pmh;ihl\ related to levels of 


reproduce. f t activity and have Utile pnvlelic 
relevance. The colour of the duct varies from 
Iridescent pink to iridescent green. 

Talon and Canefour - In a number of cases 
poor preservation made interpretation Of ihis 
region difficult. Typically the talon is a circular 
tusubcircular swelling situated atdpS short stout 
stalk although in some species e.g. tied* 
leytH'o/ichtt (fella, the talon is finger-like (Fig. 
27c) Elongation of the talon stalk occurs in 
G\rototfdea cunistnna (Fig. 55e). Rhophodon 

butrttsdalensfo (Fig. 91e) and Rotaehampa den" 

stlawellatu (Fig. I47d), while in Smplnvea in- 
tensa and Kguamen cochluiiutn the stalk is 
extremely short (Figs 1 38c, I29c> The car- 
refour is typically an expansion in the talon stalk 
al injunction with the hermaphroditic Jul! 

Albumen Gland - The size of the albumen 
gland 15 variable and probably reflects the level 

oi reproductive activity- Sbflpe ol the gland is 

1 cted largely by the degree of indentation by 
the spcrmathccal head and intestinal loops. 

Prostate and Uterus No species shov/ed any 
departure from the typical charopid pattern of 
fused prostate -uterus, The prostatic ab'ih U 
situated either along the full of the uterine 
chambers or compacted into a mass at the bASe 
of the albumen gland (Fig. 1 15a). 

The uterus consists of a lower ehamher \\\\U 
thick walls thrown into folds und a thinner, gbn- 
dular-walled upper chamber. In some cases this 
different Nation wiiscxternally conspicuous while 
in oihL rs, dissection was necessary to determine 
the . differentiation. 

Terminal Mate Genitalia - Differences in tins 
region include relative size and shape variation 
n the vas deferens, epiphallus and penis; shifts 
in the insertion of the penial retractor muscle. 
modification i>\~ the vas deferens-epiphalhis 
junction; positional variation in the cpiphallus- 
penis junction; addition of structures; and major 
changes in the penis interior particularly under 

' iditionsof sympatry. Because of differences 
in preservation stales between individual 
specimens, size and shape variations were not 


The penial retractor muscle is generally a shl HI 
lull originating on the diaphragm and inserting 
on the epiphallus near or at the epiphallus-penis 
junction. 'The muscle is elongate in Biotn- 
pftulopu recu\u isruiRhophtnlon < -onsohrimt^ 
differential contraction of individual specimens 
did not enuble full investigation ol" length dif- 
ferences. Major departures from this basic pat- 
tern include (he shift of the insertion point onto 

lhe penis head or in a point well along the 
epiphallus. The former condition is usually as- 
sociated with a shift of the cpiphallic entrance to 
a sub-apical position on the penis e.g. Ngairea 
murphyi (Fig. I (lb). Rhoplwdon eonsohrittus 
(Fig. 75e). Gvrucoeltlea and B mm phalopa (Figs 
4r,b. 4%. 52b, $5a,, nih), in Egilamen 
globosa (Fig. I32b), and Cralopn kaputareasis 
(Fig. 99b), the penial refractor muscle inserts c 
I 3 ol (he way along the epiphallus while in 
Sen -tmedeu nit did >\iu(li { 1 ig. 36b ), a ud tkothera 
gemtheCffta (Fig. |04a). the insertion point isc. 
I '2 way along the epiphallus In Rotacliaropa, 
the epiphallus is differentiated into ascending 
and descending arms and the retractor muscle is 
inserted cither at the point of reflexion (Figs 
144a. 147a). or at the vas deferens-epiphallus 
junction (Fig. 1 4 1 a ) . This latter shift in position 
m.i\ be related io ihe effects of increased whorl 

An unusual condition in Seto/nctlcu moniettht 
(Fig. 42a), and Gymeochlea eurtisiana (Fig. 
55a) has retractor muscle and terminal part •-[ 
epiphallus intertwined. 

The vas deferens is typically a thin tube des- 
cending from the prostate-ii Icrus lo the 
penio\ idueal angle before reflexing and giving 
rise to a more expanded epiphallus c. I/2-T/3 ol 
the v. ay along the penis length. Exceptions arc 
GvriH-oehlea which has the vas deferens ex- 
panded initially before becoming more slender 
prior to joining the epiphallus (Figs 46a. 49a, 
55a), and Cntlopa which has the vas deferens 
expanded for its entire length (Figs %a. 99a. 
LLCa). In most species the vas deferens enters 
the cpi fill alius aptealb through a simple pore. 
however in Elsoiheni set icatula ( Fig. 105b) and 
F. xLtathecatu (I ig. 109a), the vas deferens 
enters the epiphallus laterally through a large 
cup-shaped pilaster. Externally this complex 
junction Is P conspicuous swelling. 

The epiphallus is usually differentiated from 
the vas deferens ; '^ an abruptly expanded, mus- 
cular lube, generally shorter than the pcins. in- 
ternally with longitudinal pilasters, and reflexed 
before entering the penis apically. Departures 
from the general pattern include Gyroeachlea 
which has an epiphallus thai is as long as the 
penis (Figs 46a, 49a, 52b. 55a), and Ngtttrm 
fttUtphyi (Fig Ida). Rhophodon kempseyensis 
ii . 79a)i Elxaihcra gtwithecvtn (Fig. 109a), 
rcu cuuitl'cn/itia (lig I3a). SttOfiiedea 
rtlitliCQStaltt (Figs 36a.b). and CoettuJuiropa 
\i>rtlUlns (Pig I 1 2a). which have an epiphallus 
that Is much longci than ihe penis. In A*. 



kCftlpSCyHftitis and ,V. murphvi the elongate 
i'phalltifi may be a secondary modification re- 
lated to whorl count increment; in C .soniidus 
the change to longer epiphallus occurs under 
conditions ol complex congeneric sympatry and 
may be associated with species recognition; in£. 
i;i-.niihcnt(tt the enlarged epiphallus correlates 
with overall size increase. 

Lvtomtda contorms^ which has reduced whorl 
count, has elongate terminal genitalia with tonu 
slender epiphallus and penis of equivalent length 
(Fig 69a)i However, the bulbous pn>toeonch 
Indicates descent from a larger ancestor and 
elongation in this case can be interpreted lis 
secondary miniaturisation of a larger genital sys- 

The epiphallus undergoes minor structural 
modification \\ySeiomedea seticosuita which has 
the terminal pari of the epiphallus and apical 
section of tftfl penis bound in a connective sheath 
i Fig. 33a.bJ, and Sjanae which has the epiphal- 
lus incorporated into the penial apex (Fig. 39a.c). 
GyrocQckleeti Coawcharopa maaomphala 
(Fig. 1 15a). and C alata (Fig. 1 18b), have the 
terminal portion of the epiphallus bound to the 
penis sheath with thin strands of connective tis- 
sue. In t Isothera genithecata a thick muscular 
sheath covers the terminal half of the cpiphailus 
and the penial retractor muscle is inserted on the 
sheath (Fig. KKtg) In contrast to the mjjioi 
adjustments seen in Sttomcdva and 
Coenuchur^pa the cpiphallic sheath of E. 
gemth&CQM represents a major additive struo 

A more significant deviation Irom the above 
patterns is seen in Rotacharopa where the 
cpiphailus has ascending and descending arms 
of roughly equal length but with the- latter greatly 
expanded. The penial retractor muscle is in- 
serted either at the point of reflexion oral Ihevas 
dclerens-epiphallus junction (Figs 141a, 144a, 
|47a). This external change correlates with a 
significant change in internal structure. Typical- 
ly the interior walls o1 the epiphallus are sculp 
tured with simple longitudinal thickenings 
Whereas in Rotacharopa there arc large 
Iransveise thickenings (Figs 14 lb. 144b). 

Entry of the epiphallus into the penis is usualh 
apical, through n simple pore surrounded by a 
circular fleshy thickening or two fleshy lips. In 
(lyrovochlcu, Biomphalopa and some 
Sctoifwdett, the entry p°' n i ' s in ij subapical posi- 
tion, with the penial retractor muscle inserted on 
the penis head I figs 33c. 49b, 52b, 58b). These 
positional shifts are associated with elongation 

Of the penis (G^VWCOCfc/ftf, Biompha/opu) 01 
change in epiphallus structure iSciotncdca) 
are rri contrast to ihp positional shiiis related to 
increased whorl count in Vga/rea murphyi (I g 
lUbi and species rcc.. emuon in Rhophodott con- 
sobrimts (I kg 75c) Conical verges arc 
loped in Gyrocochlea MHiUnciA (Fig. 46b) 
and Ngai'rea cunaticufatG (Fig 13b). most 
probably in response to congeneric sym paters. 
A tubular verge is seen in Qmphalnmpa 

varicosa (Fig. 1 35b). 

The penis is generally a short, muscular tube 
which may have an apical soctipn expanded in 
response to complex internal pilasters tyt incor 
poration ol the cpiphailus. Exceptions include 
the elongate, sometimes refleu'd. cvlindru-al 
penis of GyracacM&i (Ffgs 40a.b. 4M d .D,i. ex- 
panded, barrel-shaped penis of Cralopa (Figs 
96at, 99a„ 102a), and tubular peni^ 

Coenocluiropa(f\gs 115a. 118a, 124a). A more 

Orjslic rtl^TlgC is seen n EftC penis of 

Rotacharopa which has the typical apical bulb 
of Other eharopnk :md an unusual basal e\n n 
sion. This basal extension, or preputial lube as 
it is here limned, is musl initieeablc in R. >; 
silamcllata (Tic E47a)i where if is almost twice 
the length of the penis proper. Function of this 
structure is unknown. 

A penis sheaih is prvsenl m Gyrocochlea. 
Cat'twchuropa and some ; ,pccies of Rhophodoti. 

The penis interior is variable between genera; 
and within genera under conditions ol en- 
generic sympatry. However, there is generally 
an apical epiphallic pore and basal longitudinal 
pilasters, sometimes with a muscular apical col- 
lar separating the penial bulb Irom ihe lower 
tubular section. 

In Sgairra ;\ muscular collar below the 
cpiphallic entrance effectively divides the penis 
into two ehambcrs (Figs 4b. 10b, 13b, 16b). 
while Hcdlcyoconcha has a grooved muscular 
thickening constricting the passage between the 
cpiphallic pore and the basal portion of the penis 
(Figs 24c. 27b). The shift in epiphallic entrance- 
to a subapical position in SelamedcQ wtteosMIA 
(Fig. 33c), . c . mameithi (Fig. 42e), and 
Rhophodon consnbrinu* (Fig. 75fc) has resulted 
in the development of an apical penial chamber 
which has developed its own pilaster structure, 
most Likely for species recognition. In contrast 
Ngairea murphyi which also has a subapical 
epiphaJlic entrance has an apical penial chamber 
without pronounced sculpture (Fig. 10b). 
SautHiropa Qmicrtm which has ii penis wftl 
apical bulb separated from the main chamber by 



a thin muscular collar (Fig. 65c) also has the 
epiphallic entrance apical, though slightly 
lateral, with the penial retractor muscle inserted 
on the penial bulb. In addition to the epiphallic 
entrance the penial bulb contains a horseshoe- 
shaped pilaster. This pattern is broadly 
equivalent to that seen in S. seticostata and R. 

Elsothera sericatula (Fig. 105e) and E. 
genithecata (Fig. 109b), have penes which are 
also internally divided by a muscular collar. The 
epiphallic entrance is apical and the region be- 
tween the collar and the epiphallic pore is large 
and sculptured with short fleshy thickenings. 
This change to two quite differently sculptured 
penial chambers in Elsothera is a major struc- 
tural change and contrasts with some of the 
species recognition changes mentioned pre- 

In Gyrocochtea, the long, cylindrical penis is 
internally differentiated into a lower portion with 
large, longitudinal pilasters and an area apicad 
of the simple epiphallic pore which has short, 
almost pustular, thickenings (Figs 49a, 52a). 
The exception is G. vinitincta with a conical 
verge (Fig. 46a). Biomphalopa recava has a 
single large longitudinal pilaster and short lon- 
gitudinal thickenings apicad of the epiphallic 
pore (Fig. 58b) but B. concinna lacks the apical 
thickenings and has a well-developed spatulate 
verge (Fig. 61b). Verges are also present in 
Setomedea nudicostata, Omphaloropa varicosa, 
and Rotacharopa kessneri (Figs 36b, 144b). 

The penis is typically sculptured with inter- 
generically-variable longitudinal pilasters which 
show marked intrageneric differences under 
conditions of sympatry. These differences may 
be subtle minor shifts in size and shape of 
pilasters as in Gyrocochtea, Ngairea and 
Setomedea, or may involve more drastic altera- 
tion of pilaster patterns as seen in Rhophodotu 
Coenocharopa, Egilomen and Rotacharopa. 
Gyrocochtea vinitincta and G. ptzucitamellata, 
are sympatric in areas of the Lamington National 
Park in the eastern Border Ranges. SEQ. G. 
paucilamellata retains the typical generic pat- 
tern of numerous anastomosing longitudinal 
pilasters (Fig. 52b) while G. vinitincta has an 
altered condition in which the pilasters are 
simple and unbranched (Fig. 46b). This change 
may seem minor but, together with the conical 
verge of G. vinitincta, would serve as an effec- 
tive species isolating mechanism. The 
microsympatric Ngairea corticicola and N. dor- 
rigoensis have numerous longitudinal pilasters. 

but these arc strongly corrugated in N. cor- 
ticicola (Fig. 16b) and simple \r\N. dorrigoensis. 
In Setomedea monteiihi, which is probably sym- 
patric with S. sp., short longitudinal pilasters are 
replaced by two short pad-like pilasters (Fig. 

More dramatic pilaster pattern alteration is 
seen in Coenocharopa sordidus and C. yes- 
sabahensis. Sympatry in Coenocharopa is high- 
ly complex with two to three species involved 
per locality. In C. yessabahensis an apical pad 
and large semi-circular basal pilaster (Fig. 1 24b) 
replace the more typical longitudinal pattern 
(Figs 115d, 118b), while the horseshoe-shaped 
pilasters of C. sordidus represent further varia- 
tion (Fig. 112a). 

In Rhophodon pilasters are typically fusiform 
longitudinal. R. peregrinus has a penis with an 
apical verge-like thickening and basal circular 
pilaster (Fig. 72b), while the sympatric R. con- 
sobrinus has longitudinal pilasters in the penis 
proper, a basal circular pilaster and shift in 
epiphallic entry (Fig. 75e). The two species of 
Egilomen are sympatric and have quite contrast- 
\x\u patterns of penial surface sculpture (Figs 
129b, 1 32b). 

In the three species of Rotadiscinae the 
change from typical longitudinal pilasters to the 
pad-like pilasters of Rotacharopa annabelli 
(Fig. 141b). is an equivalent change to that seen 
among members of sympatric species pairs in the 
Charopinae. However a sympatric partner for 
this species has yet to be found. 

Of the monotypic genera, Lenwebbia and Om- 
phaloropa have simple longitudinal pilasters 
(Figs 30b. 1 35b); Leiomola has an apical spongy 
thickening in addition to longitudinal pilasters 
(Fig. 69b); and the penis of Sinployea interna 
was too poorly preserved to accurately detail 
internal features, though a basal semicircular 
pilaster (Fig. 138b) relates to the generic pilaster 
pattern (Solem, 1983). 

In Nautiliropa omicron the penis has several 
anastomosing longitudinal pilasters and a large 
apical pocket stimulator (Fig. 65c). This con- 
trasts with the conchologically similar 
Gyrocochtea, and effectively separates 
Nautiliropa and Gyrocochtea. The complex 
stimulator is different to the simple circular and 
pad-like pilasters developed in other genera dis- 
cussed above and indicates that it probably is not 
a simple species recognition feature. 

Terminal Female Genitalia - Observed varia- 
tion was confined to differences in the compara- 
tive lengths of the free oviduct and vagina. The 



few species in which the internal wall sculpture 
was examined showed a repetitive pattern of 
longitudinal pilasters; no attempt was made at 
detailed analysis as presented for male genitalia. 
Length of structures may be affected by contrac- 
tion during preservation so comparisons are pre- 
sented qualitatively rather than quantitatively. 

In general, genera with elongate genitalia have 
long vaginae e.g. Hedleyoconcha, Gyrocochlea, 
Letomola and Rhophodon (Figs 24a, 46a, 69a, 
72a). However, length changes in the vagina 
may also be related to species recognition. 
Rhophodon consobrinus with short vagina (Fig. 
75a) in comparison with its congeners, is sym- 
patic with R. peregrinus and has altered ter- 
minal male genitalia. Coenocharopa, except C. 
sordidus (Fig. 112a), has average length 
vaginae: C. sordidus, which exists under condi- 
tions of complex congeneric sympatry, has a 
very short vagina. Short vaginae are also present 
in Ngairea and Lenwebbia (Fig. 30a). 

The free oviduct is typically a muscular tube 
equal in length to, or only slightly longer than, 
the vagina. Conspicuous exceptions are 
Ngairea (Figs 4a, 13a), Rhophodon consobrinus 
(Fig. 75a) and Coenocharopa sordidus (Fig. 
1 12a), which have much longer free oviducts. 

Atrium - The atrium does not display any un- 
usual features apart from minor variations in 
length. The shortened condition in Rhophodon 
consobrinus (Fig. 75a) and the elongate structure 
in Coenocharopa sordidus (Fig. 1 12a) illustrate 
the extent of variation. 

Spermatophore - Sperm packets were found in 
five species. The most unusual spermatophore 
was that of Setomedea monteithi, which had a 
long chitinous tail (Fig. 42b). The others con- 
sisted of a clavate structure in Nautiliropa 
omicron (Fig. 65d), an arc-shaped, terminally 
hooked packet in Gyrocochlea paucilamellaia 
(Fig. 52d), and a vaguely U-shaped packet in 
Rhophodon consobrinus and R. ketnpseyensis 
(Figs 75f, 79f). Although few were located, it is 
probably more than coincidental that the two 
Rhophodon species have similar looking sper- 


The usual pattern of radular structure is for a 
tricuspid central tooth, slightly smaller than the 
adjacent laterals, with long, lanceolate 
mesocone and short pointed ectocones; tricuspid 
lateral teeth which are similar in shape to the 
central; a transition zone of lateromarginai teeth 
in which the endocone becomes curved and elon- 

gate, the mesocone becomes shorter and the ec- 
tocone lengthens slightly; and marginals in 
which the endocone and mesocone are reduced, 
and the ectocone, and occasionally the en- 
docone, are split into a series of minor cusps. 
Setomedea nudicostata is typical (Fig. 35d,g,h). 

Variations fall into several categories. Minor 
differences in the size and shape of teeth occur 
in Biomphalopa and Gyrocochlea which have a 
slightly more elongate mesocone on the central 
and lateral teeth (Figs 45g, 48d, 57d). Elsoihera 
has a large, broadly lanceolate mesocone on the 
central and lateral teeth (Figs 104g; 108g,h); 
Egilomen and Cralopa have the central tooth c. 
1/2 the size of the laterals (Figs 98f, lOlf, 128h, 
1 3 1 h ) ; Omphaloropa has a small central tooth 
and reduced number of teeth per row (Fig. 
134d,h); Rotacharopa and Rhophodon have 
slender, lanceolate mesocones on the lateral and 
central teeth (Figs 146g, 190d). In contrast to the 
average generic pattern Rhophodon kempseyen- 
sis has the mesocone of the central and laterals 
greatly expanded (Fig. 78e). This condition is 
also seen in Coenocharopa yessabahensis (Figs 
1 23d. h). Significantly both R. kempseyensis and 
C. yessabahensis live on limestone rocks and the 
modification in cusp size probably relates to an 
associated shift in feeding habit. 

A more dramatic level of change in tooth shape 
is seen in Letomola contortus (Fig. 68d,e,f) and 
Nautiliropa omicron (Fig. 63c,f). L. contortus 
has a highly modified radula with a small central 
tooth which has the side cusps reduced to tiny 
knobs. In contrast, the first row of laterals has a 
greatly enlarged, broadly lanceolate mesocone 
and small side cusps. Furthermore there are very 
few lateral teeth, no transitional lateromarginai 
teeth, and serrate marginals (Fig. 68d). L. con- 
tortus lives on limestone and the enlarged 
mesocone on the laterals may be habitat related. 
How ever, the remaining modifications are major 
changes in radular structure which correlate with 
the many unusual conchological specialisations. 
In N. omicron the central and inner laterals have 
a long, slender, curved mesocone and almost 
obsolete side cusps (Fig. 63f), the outer laterals 
have the endocone lengthened and curved, and 
the marginals are unusual in having an elongate 
basal shaft with two cusps - the mesocone and 
endocone - in a crab-claw arrangement (Fig. 
63c), The teeth are arranged in V-shaped rows 
as opposed to the straight line arrangement in 
other genera. These features are significantly 
different to the typical pattern of other log-dwell- 
ing species reviewed. 



Some unusual radular variations also occur 
among species with an incomplete secondary 
ureter. Lenwebbia protoscrobiculata has typical 
central, lateral and marginal teeth but lacks tran- 
sitional lateromarginal teeth (Fig. 29d). Hed- 
leyoconcha delta has a typically tricuspid central 
tooth, but with a marked flare of the basal plate 
(Fig. 23g); the flared extension is less obvious in 
the lateral and marginal teeth. Mussonula fallax 
(Fig. 20d,e), has a tricuspid central tooth with a 
broad anterior extension, laterals initially 
bicuspid, outer laterals and lateromarginals 
tricuspid with the basal extension further en- 
larged, and marginals typically multicuspid with 
an enlarged basal extension. A similar pattern is 
present in the conchologically convergent 
Ngairea levicostata, although, the basal exten- 
sion is more pronounced (Fig. 6f). In N. murphyi 
the basal extension is absent in the central tooth 
but present as a conspicuous outwardly directed 
flare in the laterals (Fig. 9d,h). 

Ngairea corticicola, N. dorrigoensis and N. 
canaliculata have radulae with a small, broadly 
triangular central tooth with a short broad basal 
extension, and laterals and marginals with a dis- 
tinctly flared extension (Figs 3f, 12d, 15d,h). 

These changes to tooth structure in Mussonula, 
Hedleyoconcha and Ngairea represent a change 
in interrow support which in Hedleyoconcha and 
Ngairea correlates with a shift to semiarboreal 
and arboreal lifestyles. The preferred 
microhabitat of Mussonula is unknown. 


It is possible to group the conchological and 
anatomical patterns outlined above, into levels 
of evolutionary significance (Solem, 1978). At 
the lower end of the scale there are changes 
which are related to species recognition. These 
include differences in the internal sculpture of 
the penis, minor positional shifts in terminal 
reproductive structures, and differences in shell 
morphology and radular structure associated 
with small habitat shifts. Examples are given in 
species accounts. At a slightly higher level are 
those changes associated with colonising par- 
ticular adaptive zones. These include the 
periostracal setae of the damp habitat 
Setomedea; the reduced sculpture of the intersti- 
tial Coenocharopa; the strongly to moderately 
elevated shells of the semiarboreal and arboreal 
Hedleyoconcha, Lenwebbia and Ngairea; the 
strongly ribbed, flattened to biconcave shells of 
the under-log dwelling Gyrocochlea, 
Nautiliropa and Biomphalopa; the dentate, 

sculptured shells of the rock dwelling 
Rhophodon: and the dentate, secondarily 
modified shell of rock dwelling Letomola. Cor- 
responding anatomical changes include altera- 
tions in penis shape and internal penial structure, 
addition of major structures to the genitalia, 
changes in kidney shape, and radical modifica- 
tions in radular morphology. Above this level 
are basic changes in sculptural elements and 
large scale changes in anatomy, such as in 
Rotacharopa and possibly the group of genera 
with incomplete secondary ureter. 


Family CHAROPIDAE Hutton, 1884 

Solem (1983, p. 59) presented a detailed diag- 
nosis that is accepted herein. 

Subfamily CHAROPINAE Hutton, 1884 

A broad definition of the subfamily is 
presented by Solem (1983, p. 70). The genitalia 
of the Charopinae reviewed here show a consis- 
tent pattern of muscular epiphallus with internal 
longitudinal thickenings, reflexed before enter- 
ing the penis through a simple pore, or oc- 
casionally a verge. The penis has an apical bulb 
containing the epiphallic entrance and a lower 
basal portion with longitudinal thickenings 
which may be modified under conditions of sym- 
patic species interaction. The pallial configura- 
tion includes a kidney in which the pericardial 
lobe is usually much longer than the rectal lobe 
and the secondary ureter is complete or rarely 

Ngairea gen. nov. 

Oreokera Iredale, 1933 (part), p. 54, nom. nud.; 
Iredale, 1937a (part), p. 318, nom. nud.; Iredale, 
1941a, p. 261, nom. nud.; non Stanisic, 1987, p.2. 

For my youngest daughter Ngaire. 

Type Species 
Oreokera dorrigoensis Iredale, 1941 . 

Previous Studies 

Iredale (1933) introduced Oreokera for Flam- 
mulina cumulus Odhner, 1917 and added Helix 
corticicola Cox, 1866 (Iredale 1937a) and Ore - 



okera dorrigoensis Iredale, 1941. Stanisic 
(1987) validated Oreokera, including only F. 
cumulus, and one other north Queensland 
species. Ngairea is introduced to include//, cor- 
ticicola, O. dorrigoensis, two new species from 
southern Queensland, and 'Helix' murphyi Cox, 
1864 from southern NSW. The latter was pre- 
viously included in the punctid form-group 
Paralaoma Iredale, 1913 bv Iredale (1937a, 


Large to very large, diameter 5.46-7.40 mm, 
with 4 1/8 to 5 7/8 normally to tightly {murphyi) 
coiled whorls. Height 2.86-5.35 mm. Apex and 
spire moderately to strongly (murphyi, levicos- 
tata) elevated, body whorl descending slightly. 
Protoconch almost flat, 1 5/8 to 1 3/4 whorls. 
Apical sculpture of curved radial ribs, reduced 
(canaliculata) or modified (murphyi), with in- 
cised spiral grooves (canaliculata, corticicola), 
weak raised spiral cords (levicostatd) or without 
spiral elements (dorrigoensis, murphyi). Post 
nuclear sculpture of crowded, protectively 
sinuated radial ribs which may be reduced in 
some species (corticicola), Microsculpture of 
crowded spiral cords (murphyi, dorrigoensis), 
incised spiral grooves (corticicola, 
Canaliculatd), or fine spiral lines (levicostata). 
No microriblets present. Sculpture continuous 
on the base but reduced. Umbilicus narrow to 
widely U-shaped (murphyi), rarely closed (dor- 
rigoensis). Sutures impressed, maybe chan- 
nelled (canaliculata). Whorls weakly angulate 
to strongly carinate (murphyi), sometimes 
rounded (canaliculata). Aperture roundly lunate 
to subquadrate. Lip simple, sometimes slightly 
thickened; columella vertical, reflected toward 
umbilicus. Colour yellow-horn, with red flam- 
mulations or without (murphyi and 

Foot and tail broad, bluntly rounded posterior- 
ly; a caudal horn sometimes present {levicos- 
tata). Animal creamy-white with orange-grey 
speckling on the sides of the foot and on the 
mantle collar. Pallial roof and visceral hump 
with variable black speckling. Ommatophores 
black. Kidney broadly to narrowly (murphyi) 
triangular; rectal lobe vestigial or absent (mur- 
phyi). Ureter sigmurethrous with incomplete 
secondary branch. Ureteric opening at the rear of 
the pallial cavity in the angle between kidney and 
hindgut. Genitalia with ovotestis of two or more 
(murphyi) clumps of palmalely clavate lobes of 
alveoli, separated by digestive gland and 

oriented at right angles to the plane of coiling. 
Terminal male genitalia with muscular epiphal- 
lus, sometimes much longer (canaliculata) than 
the penis, entering the penis apically, or subapi- 
cally (murphyi). Epiphallic pore central in mus- 
cularised pilaster or opening through a conical 
verge (canaliculata). Penial retractor muscle in- 
serted on the epiphallus (dorrigoensis), penis- 
epiphallus junction (corticicola, canaliculata), 
or on the penis (murphyi). Penis with an apical 
bulb internally separated from a lower tapered, 
tubular section by a muscular collar. Internal 
penial sculpture of broad, corrugated (cor- 
ticicola, canaliculata) or slender (murphyi, dor- 
rigoensis) fleshy, longitudinal pilasters. Female 
reproductive tract without unusual features. 

Central tooth of radula broad, triangular with 
tiny ectocones, or slender lanceolate (levicos- 
tata, murphyi) with prominent ectocones. 

Distribution and Ecology 

Ngairea extends from warm temperate closed 
forests in the Illawarra region of southern NSW 
to the subtropical and temperate rainforests of 
northern NSW, and drier subtropical forests of 
southern Queensland. With the exception of N, 
murphyi and N. levicostata species distributions 
are well known. Most species live under the bark 
of fallen trees. 

N. corticicola is sympatric with N. dorrigoen- 
sis from Dorrigo to the eastern Border Ranges; 
N. levicostata is sympatric with N. dorrigoensis 
in the Koreelah State Forest region of the 
western Border Ranges. N. dorrigoensis and N. 
corticicola were collected from under the same 
piece of bark at Upper Pine Creek, Canungra, 
SEQ. In N. dorrigoensis and A r . levicostata dif- 
ferences in shell shape and radula indicate diver- 
gent microhabitat preferences. 

Patterns of Shell Variation 

Size and Shape - Adult Ngairea are some of 
the largest Australian subtropical charopids with 
mean shell diameter 5.80-6.90 mm. Average 
whorl counts range from 4 1/2- to 5 1/2-. N. 
murphyi has the highest mean whorl count (5 
1/2-), smallest mean diameter, but has a smaller 
whorl cross section. 

Reduced height in N. murphyi is the result of 
smaller shell size but N. levicostata has a taller 
shell through spire elevation. In A", murphyi and 
N. dorrigoensis decreased shell height correlates 
with keel development on the body whorl. 

The umbilical opening is smallest in species 
with a distinctive keel (N. dorrigoensis and A r . 



levicosiata) although N. murphyi has a secon- 
darily enlarged umbilicus resulting from reduc- 
tion in size of whorl cross section. 

Sculpture - Apical sculpture consists of 
curved, crowded radial ribs which may be 
pronounced, reduced or modified into vague 
radial ridges and wrinkles. Less prominent spiral 
elements may be present as incised spiral lines 
or raised cords. In N. corticicola (Fig. 15e) 
intersections of radials and spirals form a series 
of pits. 

Postapical sculpture is predominantly radial, 
with ribs varying from bold, broad, regularly 
spaced (Fig. 12a), and high, crowded (Fig. 6e) to 
very crowded (Figs 3e, 9g). Various types of 
spiral sculpture are also present and in N. cor- 
ticicola (Fig. 15f), incised spiral lines become 
notched, approximating some helicarionid taxa, 
e.g. Westracystis Iredale, 1939 and Delinitesta 
Iredale, 1933 (Solem, 1982; Stanisic, 1988). 

One of the more interesting developments in 
the shell of Ngairea is the canalised sutures seen 
in N canaliculate. Solem (1983, pp. 9, 108) 
recorded this feature in the Pacific Island 
charopid Sinployea proxima (Garrett, 1872). 
Canalisation is due to detachment of part of the 
body whorl and in N. canaliculata results in a 
relatively wide channel. 

Variation in shell morphology is greater than 
in other genera included in this study indicating 
a long, complex period of evolution. N. murphyi, 
which has developed in isolation from the main 
group of species exhibits some major departures 
from average patterns. 

Patterns of Anatomical Variation 

Variations in anatomy involve shape changes 
in the pallial structures, size differences in 
reproductive organs, species interaction 
modifications in the terminal male genitalia, and 
diet related shifts in radular morphology. 


Ngairea and Mussonula may be confused and 
are sympatric in and around the eastern part of 
the Border Ranges. Mussonula has a strongly 
exsert protoconch with prominent spiral cords 
and vague radial ridges prominent toward the 
nuclear-post nuclear shell boundary (Figs 18d, 
20f). Lenwebbia is sympatric with Ngairea be- 
tween Colosseum Creek and the Bobby Range, 
SEQ, and bears some similarity. However, it has 
a strongly punctate protoconch, adult sculpture 
of spiral grooves and fine thread-like radials 
(Fig. 29e-g), lacks the internal division of the 

penis, and has the kidney apex divided by a 
portion of reflexed ureter (Fig. 30b, e). 

1. Shell with prominent radial ribs 2 

Shell without prominent radial ribs; base al- 
most smooth (Fig. 1 5b) corticicola 

2. Sutures simple 3 

Sutures channelled (Fig. 12e) ....canaliculata 

3. Umbilicus nearly closed to slightly open; 
peripheral keel obtuse 4 

Umbilicus widely open (Fig. 9b); peripheral 
keel acute (Fig. 9c) murphyi 

4.Microsculpture of prominent spiral cords (Fig. 

3e); spire lower, mean H/D ratio 0.56 


Microsculpture of weak spiral lines (Fig. 6e); 

spire higher, mean H/D ratio 0.73 


Ngairea dorrigoensis (Iredale, 1941) 
comb. nov.(Figs 3-5; Tables 2, 3) 

Flammulina corticicola (Cox); Hedley, 1912 (part), p. 

264, pi. 7, figs 46-48. 
Orcokera dorrigoensis Iredale, 1941a,p. 265, fig. 5 


N. dorrigoensis differs from N. corticicola in 
its more prominent radial sculpture, more angu- 
late periphery, narrower umbilicus and micros- 
culpture of crowded microspiral thickenings 
(Fig. 3c). N. levicostata has a more elevated shell 
and microsculpture of fine spiral lines (Fig. 6e). 
The apical spiral cords, smaller size and lack of 
prominent spiral cords on the adult whorls of 
Mussonula Iredale, 1937 readily separate mem- 
bers of that genus from N. dorrigoensis. 

Type Material 
Holotype: AMC63517, Dorrigo Scrub, NSW., 

Height of shell 3.45 mm, diameter 6.23 mm, H/D ratio 
0.55, D/U ratio 14.49, whorls 4 1/2-. 
Paratype;AMC63513, same locality as holotype. 



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Other Materia! 

NEW SOUTH WALES: Platvpus Creek, New 
England N.P.. (1 AMCU2440, 27 Feb 1961, D.F. 
McMichael); Acacia Plateau Rd, c. 12.2 km from 
Acacia Ck - Killarney Road, SEVT/MVF (28-*2rS, 
152°24'E)(1.AMC12S560, 15 Mar 1981, J. Stanisic, 
D. Potter); Beaury S.F., c. 1 5 km W of Urbenville, 810 
m (28*27*45, 152 B 24"E) (3, AMC142475, 18 May 
1976, PH. Colman, L Loch): Marengo S.F., c. 40 km 
NW of Dorrigo, 1020 m (30'06'S, 152 n 25 E) (5. 
AMCI42476, 10Febl976J.B.Burch,P.H. Colman); 
SE ofTabulam, tributary of Busbv's Creek (29 i '02'S, 
I52M3 *E) (2, AMC137757, 29 Aug 1982, ABRS - 
AM/OM ); The Glade, Dorn^o N.P., 600 m, temperate 
rainforest (30"22"20"S, 152 rj 43'40"E ) (1, 
QMMO16440, 12 Nov 1983. ABRS - AM/OM), 
Tooloom Scrub. Beaury S.F., CNVF (28 29"S, 
152 B 24'E) (2, QMMO10939, L5 Mar 1981, ABRS - 
AM/OM): Dorrigo (6, AMC 142443); Dorrigo, under 
bark and logs (1. AMC63788, Dec [910, S.W. Jack- 
son); Dorrigo (1, AMC 1 42444, ex Helms); Moonpar 
S.F . Dorrigo (1, AMC14244L 4 Apr 1960. D.K. 
McAlpme); Booyang (= Booyong?) Richmond River 
(1, AMC57S6, ex Hedley); Washpool S.F., SW of 
Casino, dry eucalvpi forest (29M3"30"S 3 152*29*6) 
(1 ? AMC136S79, 29 Aug 19S2, ABRS - AM/QM); 
Wonga Walk, The Glade, Dorrigo N.P.. underbarkof 
fallen tree, rainforest (30*22'S, 152'44 , E) (1, 
QMM017137, 6 Mar 1987, J. Stanisic, D. Poller); 
Mills Rd, Moonpar S.F., under bark of fallen tree, 
rainforest (30 r 'LVS, 1S2 B 39'E) (1, QMM017141, 7 
Mar 1987. J. Stanisic, D. Potter). 
SOUTH EAST QUEENSLAND: c. 1 km Eof Cair's 
Lookout. Koreelah S.F., under bark of fallen tree, 
NVF (3, OMMO 1 6470, 3 May 1 986, J. Stanisic); Mt 
Glorious, litter. NVF(1, QMMOI6130, 26 Jan 191*6, 
J. Slanisic). Mt Glorious, C. 3 km SE of summit. NVF 
(27 20'S, 152\46*E) (1. QMM011992, 2 Dec 1982, 
ABRS - AM/OM): c 28.5 km from Goomburra. 
Goomburra S.F., 600 m, CNVF/Palms (27 D 59'S, 
152'21'E) (3. AMC136812, OMMO 1 2690, 7 Dec 
1981. ABRS - AM/OM); Fred's Road. Ml Mee, rain- 
forest (27"05'S, 152"43'E)(2, QMMO16460, 14 Apr 
1980. J. Stanisic. N. Hail, A. Green); Maiala N.P., Ml 
Glorious, lilter (1, QMM011852, 20 Jun 1982, MSA 
Party); Lower Ballanjui Falls circuit, Laminglon N.P., 
litter, NVF ( I , OM MO 1 6448, 3 Oct 1 986, J. Stanisic. 
D. Potter); Canungerah (= Canungra), pine scrub {5, 
AMC32997. Jul 1908, S.W- Jackson); Mt Tamborine 
(2, AMC142442); Maiala N.P., Mt Glorious, CNVF 
{1. OMMO6304. 1 Oct 1976, MJ. Bishop); Mt Mis- 
take, Goomburra S.F.. under bark of fallen trees, 
CNVF (2, 0MMO1657L 5 May I9S6, J. Stanisic); 
Running Ck, Lion's Road, via Rathdowney, under 
bark of fallen tree, rainforest near picnic area (2, 




* Jib 

" JB 

■■: ; T' V 







1 mm. 


«.r f 


FIG. 3. A^gazrea rfo/-Wgoen5w(Iredale, 1941). a-c,Dorrigo Scrub, NSW. AMC63517, holotype; d-e, c. 1 km 
E of Carr's Lookout, Koreelah S.F., NSW. QMMO16470; f, Upper Pine Ck, Canungra, SEQ. QMM016544. 
a-c, entire shell; d, apical sculpture; e, post nuclear sculpture; f, central (top left) and lateral teeth. Scale lines 
as marked. 






F1G.4. Ngaireadorrigoensis(\TQda\e,\^4\). a, c-e. Maiala N. P.. Mt Glorious, SEQ. QMMO6304. b. Upper 
Pine Creek. Canungra. SEQ. QMM016583. a, genitalia; b, details of penis interior; c, ovotestis; d. 
hermaphroditic duct; e, pallial cavity. Scale lines as marked. 



5 2 E 


j- a: 

- d 

^3 1^ 

x £ 


— ~ ; 
■-i — 

^ZB J 

2 n "> Q £ 

5 s 



■r ■J- 

- ~ 

Is it , 

x— t-, X 

0> <3 U^Sv 

^ - 

IF] £ *J 

3 32 

3S *g 








JL * 

- ? 

i=rs - |- fib 

13S* s ai ri 

« s < < j< Bo 



■6 a S 

27 D S 


29 D S 




i 52°E 



r IG, 5- Distribution ofNgtoirsutdorrigotttsfi (Iredale, 

QMM016472, 1 fatl 1480, V, Kessner): Upper Pine 
Creek, Canungra, NVF/Araucaria. under bark {15. 
QMM016S44, 1 Oct 1986, J- Stanisic, D. Potter; 3. 
QMM0155S3, 29 Sept 1986, J. Stamsic. J. QlflSCJ 



Shell depressed, large, diameter 5.71-7.40 
mm (mean 6.50 mm), with 4 1,8 to 5 1 '8 (mean 
4 1 4-) normally coiled whorls. Ape\ and spire 
moderately elevated, height of shell 3.36-4.62 
mm (mean 3.78 mm). H/D ratio 0.52-0.64 (mean 
0-58). Bodv whorl descending slightly in front. 
SP/BWW 0.17-0.39 (mean 0.3 l"). Protoconch 
with 1 5/8* to 1 3/4+ whorls, mean diameter 
]065.0|jLm at I 1/2 whorls. Apical sculpture of 

low curved radial ribs, without spiral elem 
(Fig. 3d). Post nuclear sculpture of numerous 
protractively sinuated radial ribs whose inter- 
stices are 5-7 times their width. Microsculpturc 
of crowded raised spiral cords {Fig. 3e}. Sculp- 
ture continuous on the base (Fig. 3b). Umbilicus 
small, diameter 0.30-0.68 mm. (mean 1 1.46 mm), 
D/U ratio 9.26-21*83 (mean 14.59), to closed. 
Sutures slightly impressed. Whorls flattened 
above and below an ungulate periphery (Fig. 3c). 
Body whorl keeled, more angulate in juvenile 
specimens. Aperture ovately lunate. Lip simple, 
becoming thickened at the basal margin. 
Columella thickened and reflected over the um- 
bilicus. Parietal callus white with brown flam- 
mulalions. pustulose. Colour yellow-horn with 
reddish-brown flammulations above, continued 
below the periphery and onto the base in juvenile 
specimens, but fading toward the umbilicus in 
adult specimens. Based on 16 measured aduliv 

Epiphallus internally with longitudinal 
pilasters, entering penis through a large ittUS 
cularised pilaster (Fig. 4b). Penis (Fig. 4b) a 
short cylindrical muscularised tube, more swol- 
len apically; internally with apical muscular col- 
lar consisting of 5-6 pad-like thickenings, ami 
slender longitudinal fleshy pilasters in the lower 
regions. Pcnial retractor muscle a shortened lufl 
inserting on the epiphallus near its junction with 
the penis. Free oviduct muscular rcflexed, much 
longer than vagina (Fig. 4a). 

Radula (Fig. 3f) with central tooth having a 
large, broadly triangular mesoconc and reduced 

Based on six dissected specimens (QM- 
MO6304 : QMM016471, QMMO16470, 

Range and Habitat 

Under the bark of rotting logs in moist suh 
tropical notophyll vine forests of cool wet high- 
land areas of the Great Dividing Range (from 
New England. NSW to the Mistake Mountains, 
SEQ); the Border Ranges, and the D'Aguilar 
Range. SEQ: penetrating adjacent drier subtropi- 
cal forest (low microphyll vine forest, semi- 
evergreen vine thicket) in the Koreelah Creek 
and Richmond Range areas, northern NSW. 


Northern representatives of A', dorrigoensis 
are larger than their southern counterparts and 
have the umbilicus more closed (Table 3). Pos- 
sibly the warmer climate provides a longer 
growth period in northern populations. 




.'.lull I 


- ■ " 

6 ,. „- 

: r 


• ^ ^r f W V % 





FIG. 6. Ngairea levicostata sp. nov. a-c, Along Acacia Plateau Rd, Koreelah S.F., NSW QMMO10959 
holotype;d-f, Cunningham's Gap, SEQ. QMM017153, QMMO 12684, paratypes. a-c, entire shell; d. apical 
sculpture; e, post nuclear sculpture; f, lateral teeth. Scale lines as marked. 



Ngairea levicostata sp. nov. 
(Figs 6-8; Tables 2,3) 

El ymolouy 

Latin levis, smooth; casta, rib; referring to 
reduced microsculpture on major ribs of adult 


N. levicostata is most similar to N. domgoen- 
sis but differs in having a higher spire, fewer 
whorls with different sculptural features, a 
protoconch with curved radial ridges and fine, 
weak, crowded spiral cords and in lacking the 
microsculpture of crowded wavy spiral cords 
present in A', dorrigoensis. Species o\' Xfussonula 
spp. are separable from N. levicosiaia by their 
exsert protoconchs and apical sculpture of bold 
spiral cords (Figs 18d,20f). 

Type Material 

Holotyfe: OMMO 10959, Koreelah S F . NENSW, 
along Acacia Plateau RdL 850 m (28*21 *S. [52'24'E), 
15 Mar 1981. AM/QM - ABRS 1931. Height of shell 
4. 79 mm, diameter 6.03 mm. H/D ratio 0.76, D/U ratio 
15.35. whorls 4 5/8. 

Paratyfes: Cunninghams Gap. SEO. near top, 755 
m, NX't/Araucaria (28°04 ? S, 15224'E)(3. 
QMMO 1 2684, QMMO 10994, AMCIZ.SM'J. lo Mar 
1981. AM/QM - ABRS 1VS1 ); Cunningham's Gap, 
SEQ. leaf littej (1, QMM017153. 18 Jun 1979, G. 


Shell trochoid, large, diameter 5-63-6.30 mm 
(mean 5.97 mm) with 4 1/4 to 4 5/8 (mean 4 IV) 
loosely coiled whorls. Apex and spire strongly 
elevated, height of shell 3.95^.79 mm (mean 
4.37 mm). H/D ratio 0. 70-0.76 (mean 0.73). 
Body whorl descending more rapidly, SP/BWW 
ratio i 0.36-0.44 (mean 0.40). Protoconch with I 
5/8 whorls, mean diameter 11 I9.()u.m at 1 1/2 
whorls. Apical sculpture of low curved radial 
ribs crossed by raised spiral cords (Fig. 6d). Post 
nuclear sculpture of numerous crowded, high, 
protractively sinuated radial ribs (Fig, 6e). Fine 
raised microspiral lines present but no 
microriblets. Sculpture continued onto the base 
(Fig. 6b). Umbilicus narrow U-shaped, diameter 
0.45-0.47 mm (mean 0.46 mm). D/U ratio 
12.46-13.35. (mean 12.91). Sutures impressed. 
Whorls flattened above and rounded below an 
angulate periphery (Fig. 6c). Aperture sub- 
quadrate. Lip simple with a slight thickening at 
the baso-columellar margin. Columella almost 


FIG. 7. Ngairea levicostata sp. nov. Cunningham's 
Gap, SEQ. QMM0126S4, paratype. Pallia! cavity. 
Scale line as marked. 

vertical, reflected toward the umbilicus. Parietal 
callus well developed. Colour yellow-horn with 
adhering dirt particles Based on 2 measured 

Foot and tail moderately broad, tapering 
posteriorly with a weak caudal horn. Mantle 
collar thickened with a strongly developed glan- 
dular zone (Fig. 7). Kidney vaguely triangular c. 
1/3 length of pallial cavity, rectal lobe reduced. 
Pulmonary vein inconspicuous. Reproductive 
system immature. 

Radula (Fig. 6f) with central tooth having a 
slender lanceolate mesoconc and small but 
prominent side cusps; inner laterals with slender 
mesocone. endocone absent and weakly 
developed ectocone. 

Based on one dissected specimen (QM- 


Z 1 ? 


■ " " . 

-: : 

FIG S. Distribution of Ngairca levicostata sp. nov. 







H 1^ Hv MG 

FIG. 10. Ngairea murphy i (Cox, 1864). c. 4km Eof Fitzroy Falls on Kangaroo Valley - Fitzroy Falls Rd, NSW. 
QMMO 16932. a, genitalia; b, details of penis interior; c, talon and hermaphroditic duct; d, ovotestis;e, pallial 
cavity. Scale lines as marked. 

FIG. 9. Ngairea murphyi (Cox, 1864). a-c, Mt Keira, Wollongong, NSW. AMC 142962, neotype; e-g, 
Woodhill NSW, AMC142968; d,h, c. 4km E of Fitzroy Falls on Kangaroo Valley-Fitzroy Falls Rd, NSW. 
QMMO 16932. a-c, entire shell; d, central and lateral teeth; e, spire; f, apical sculpture; g, post nuclear 
sculpture; h, lateromarginal teeth. Scale lines as marked. 



Range and Habitat 

iV. levicostaia is known from two localities on 
the cool wet highlands of the Great Dividing 
Range in northern NSW and SEQ where it is 
sympatrie with N. dorrigoensis. The micro- 
habitat is unknown. 

Rf marks 

A. Icvicostcva is erected to assist comparative 
and biogeographical discussions; its concnoiogi- 
eal features are distinctive. Although the general 
shell facies relates to Mussonula, N. Icvicostata 
is included in \gairea primarily on form of the 

Ngairea murphvi<Cox. 1864) comb. nov. 

(Figs 9-11; Tables 2,3) 

Httixmurphxi Cox, ISM. p. 3-7; </ok, ISoS. p.23. 
Parfitooftia murphxi (Co\K Iredale. |^37a, p 314; 
Iredale. 1941a. p. 264. 


A', murphxi is a southern outlier distinguished 
within the genus by its higher whorl count and 
wider umbilicus (Table 2). A. dorrigoensis 
which it most resembles, has a liny to closed 
umbilicus, weaker keel, fewer whorls and 
protoconch with curved radial ridges (Fig. 3d). 
Apical sculpture involves irregular radial 
rugosities and spiral wrinkles producing a pitted 
effect (Tig. °-f). Anatomically distinctive are the 
penis with epiphallic pore located subapieally 
and the penial retractor muscle inserted apically 

the. !t)b). 

Previous stutjies 

Cox (1864) introduced the name without an 
accompanying figure and referred to it again 
without figures in 1868. A further listing (Cow 
I Ml)*}, p. 54) was accompanied by an explanation 
for the lack of figures; "I fear "the types of this 
species have disappeared from our (Australian) 
Museum Collection. The specimens of it which 
I possessed were unfortunately smashed white in 
the artists hands" 1 . The large whorl count, small 
Size, widely open umbilicus, sharply keeled 
body whorl and reticulate sculpture, referred to 
in Cox \ original description, together with the 
type locality u\' Wollongong, NSW. effectively 
fi\ Cox's name to the species discussed below. 
The neotype was collected at Mt Keira. Wollon- 
gong, which I consider to be near the original 
type local it v, 

Iredale f 1937a, p. 314) placed H. murphxi in 

Para/aoma Iredale, 1913. without explanation, 
later (iredale, 194 la. p. 264) admitting that no 
known specimens of//, murphy i existed and that 
this decision was based on broad details of Cox's 
description. In Paralaoma the shell is much 
smaller, more fragile, with fewer whorls and 
bold spiral cords on the protoconch. Although 
morphology of AT murphyi is unusual, penial 
morphology, shell sculpture and peculiar body 
colour are considered sufficient to endorse the 
new generic placement. 

Type Material 

Nhitvpf: AMC 142962, Ml Keira, Wollongong, 
NS\v\ Collected bv C.F. 6 Ho\ 1948, 
Height ot shell 3. 7 mm, diameter 5. SS mm, H D ratio 
0.63, D/U ratio 4. 1 9. wto>rls5 

Other \lATHRr\L 

C, 4 km F ot PUTOJ FaJlS on Kangaroo Valley - 
Filzroy Falls road. NSW. temperate rainforest, under 
bart of fallen tree (5, QMMQ16932, J.Stanisic#4Jan 
] 987): WooitMil. c. 4 mis N\V of Bern . NSW. mixed 
l"orc>t, E _ide of ranue (1, AMCU:%S, W. Ponder, 
14 Nm |^70): Mt Cambewarra, nr NoWra, NSW (I 
AMC.-W964. L Price,.300ct, 1963): Mt Keira. Wol- 
longong. NSW, from rainforest at scout camp (1, 

anic (42962, C,F, McUuchlan, 6 No\ I94S>. 


Shell depressed to elevated trochoid, moder- 
ately large, diameter 5.46-6.05 mm (mean 5.80 



. i 

Fit; II, Disirihiiimn ni Vgairea murphyi (Cbx, 1864) 
in the Illawarra region. New Somh Wales 



mni) with 5 to 5 7/8 (mean 5 I/2-) tightly coiled 
whorls. Apex and spire weakly to strongly 
elevated, height of shell 2.NO-J.70 mm (mean 
3.24 mm). H/D ratio 0,49-0.63, (mean 0.56). 
Body whorl not descending in front, SP/BWW 
II 32-0.65 (mean 0.44). Protoconch with ir- 
regular curved radial ridges and pits, spiral de- 
ments absent (Fig. 9f). Post nuclear sculpture of 
numerous, crowded, protractively sinuatcd 
radial ribs (Fig. 9e, g) f whose interstices are 
ahoul 5—7 times their width. No microriblets. 
Mtcrosculpture of numerous, crowded, narrow 
•vpii-LiI cords. Sculpture continuous on the base 
but reduced (Fig, %). Umbilicus relatively wide, 
V-shaped, diameter I.28-1.5S mm (mean 1.3° 
mm). D/U ratio 3.84-4.39 (mean 4,17). Sutures 
impressed, Whorls flattened above and below a 
carinate periphery (Fig 9c) Aperture ovaiely 
lunate to subquadrate. Lip simple with a slight 
thickening at the baso-columcliar margin. 
Columella almost vertical, slightly reflected 
toward the umbilicus. Parietal callus well 
developed, white, pustulose. Colour cream v -yel- 
low horn without flammulations. Based on four 
n-i. asured adults. 

Kidney (Fig. lUe) with elongate pericardial 
lobe and no rectal lobe, 1/2 length of the pallial 
icgion. Ovotestis (Fig. Uld) three or more 
dumps of creamy-while palmaiclv clavate lobes 
of alveoli. Epiphallus (Fig. 10a) twice length of 
the penis with internal longitudinal pilasters, 
entering penis subapicallv through a muscular 
pilaster (Fig. 10b). Penis (Kig. 10b) short. 
C) ImdrieaL musculariscd. internally with an api- 
cal muscular collar and fleshy, longitudinal 
pilasters below the collar. Penial retractor 
muscle (Fig. 10a, b) inserted apieally on the 
penis. Vagina relatively long (Fig. 10a). 

Radula (Fig. 9d> h) with mesocone of central 
tooth slender lanceolate and cctoconcs small but 

Based on 2 dissected specimens (QM- 

Range 'ah Kahit.m 

M, tnttrphyi occurs in an area which has been 
^reatlv altered and disturbed by white settlement 
HJaur* 1957, pp. 191, IBS). Its distribution en- 
compasses the remnant patches of rnoisi waim 
temperate and cool subtropical vine forests of the 
Wavvarra region, NSW. from Wollongong south 
to Mt Cambewarra, near Nowra. The original 
specimens were taken from under stones; live 
material has been collected from under bark of a 
fallen tree (OM MO 16932). 


The high whorl count has not been observed in 
any other east coast charopid with a trochoid 
shell. The North Queensland helicarionid The*- 
keJotnetiSOr Iredale, 1933 has a trochoid shell, 
wide umbilicus and high whorl count but the 
similarity is convergent. The tew available 
specimens of : \\ murphyi show a great deal of 
variation in shell height brought about by chan- 
ges in spire protrusion (Table 3) 

Ngairea canalicular sp no\ . 

(Figs 12-l4;Tab^2.4i 

Flammulina vuriicicola (Cox). He J lev. 1912 (pan), p. 
2(>4, but not figure. 

Btwioi <"h_; v 

Latin canaliculus, small channel: for the chan- 
nelled sutures. 


Readily identified by its channelled sutures, 
low spire, rounded periphery and sculpture of 
low, broad radial ridgescrossedby fine, crowded 
microspiral grooves (Fig. 1 2g). / en^cbbia pro- 
toscrobicultita which is sympatric with N. 
canaliculate* has similar shell shape but differs 
markedly by having a punctate protoconch. fine 
Ihread-ttkc radial nblels on the body whorl, . 
simple sutures (Fig. 29c-g). 

Tvpr Material 

Hoiut>pi- QMMOLG454, Beauty Spot 98 3 Kroortv 
bit Tops. SFO. Collected h> G.B. Momeiih. 20 Sep 
, : Height 3. 87 mm, diameter b.55 mm. H/D ratio 

MO 17 154. 1 specimen, same data as liolotypc; 
Kroombit Tops, TA 47 creek ( I, QMMO IM67 > 9-!^ 
Dee. IVS3,G.B. Monleilh. G. Thompson); Kroombit 
lops uncle, i palm fro nils, NVP/Palms (I. 
OMMOUSm. S Mas l'VK4. J. Stanis.e. D. Poller); Ml 

Fori William, Kalpeww s,r , nvf (24*39% isr 

20'E)(2, QMM012598; AMCI36757. 4 Sept 1^82. 
ABRS - AM/QM; Camp Creek. S of Miriam Vale, SW 
side of Bobbv Range, MVF (24 a 36 , 2U"S. 151 n 33'E) 
i \ QMMQ132M AMC1?7913, 6 Sept 1982, ABRS 
- AM/QM); Rainforest walk, Kalpowar S.F , 
MVFfAraucaria, (24 C 4CS. 15J21 E) (4, AMC- 
L36742, QMM012574. 4 Sept 1982, ABRS - 
AM/QM); Bobby Range ST., NVF (24'37'S, 
15I^32'EJ "I. QMMOI2L51, AMC136509, fi Sept 
': ABRS - AM/QM); Bulbunn ( Austral) S.F., S of 
Gladstone. >50 m i?4' US, \$V29'B) U, 




€sf4s h 





0.5 mm 


FIG, 13. Ngairea canaliculuia sp. now a, e-e, c. 12.1km from Granite Creek crossing on Bobbv RanLie-Mi 
Perry Rd.SEQ. QMMO 16468.paratype. b. Mt Fori William, Kalpowar S.F.. SEQ. QMMOI6469.paratype. 
a. genitalia: b. details of penis interior; c, ovotestis: d, hermaphroditic duct: e. pallia 1 cavity. Scale lines as 

FIG. 12. Ngairea canaliculata sp. nov. a-c. Beauty Spot 9S. Kroombit Tops. SEQ. OMMO 16454, holotype; 
d-h, summit. Ml Fort William, Kalpowar S.F. ? SEQ. OMMO 16469. paratype. a-c. entire shell: d, central 
and lateral teeth: e, details of spire showing channelled sutures: f, apical sculpture: g. post nuclear sculpture; 
h, details of base and umbilicus. Scale lines as marked. 







150' : 51 C E ! 52 D E 


FIG. 14. Distribution of iXgairea canalicufaia sp. 

AMC 1 4246v S May 1975, J.B. Burch. W.F. Ponder. 
PH. Colman); Bulburin (Austral) ST., S of 
Gladstone. 540 m. CNVF-A1 lores! Ispe (24°33'S, 
151 29' E I { l -. AMC 1424<i&, J.B. Burch, W.F. Fonder. 
F.H. Colman): Boitle-lree scrub, wesi of Gladstone, 
on the ground under leaves ( L A\ir^'>JS, Aug 1 91 KB, 
S.W. Jackson); Colosseum Creek- 16 km S of Miriam 
Vale {2 AMC142445, 25 May 1958. L. Price); Kal- 
I .: .. - S i . cner MVFVAraucaria (9, QMMOUS459, 
4 Jul 1984, D, Poller, J. Slanisic, K. Emberton); Mt 
Lurcom, on rockv slope, litter, SEVT (2. 
QMM01M43. 9 May 19S4. J. Stanisic D. Poller); 
Summit, Mt Fori William. Kalpowar ST.. under hark. 
r\\> (4. OMM01646C 3 Jul I9S4.J, Stanisic, D, 
Poller. K. Emberton); c. 12.1 km from Granite Creek 
crossingon Bobby Range - Ml Penv Road, under hark 
Of fallen trees, VVF/Palms (24, OMMOIo4c.N, to 
Sepi 10X5. C Stanisic, D. Poller). 

Othfr MaTERJ vl 

SW of Miriam Vale, rainforest alone creek 
| I 23 3H"S. ISV2TB) (2, AMcf37S90. 
QMM013223, 6 Sept 1982, ABRS - AM/OM); Sum- 
mit of Mt Booroon Booroon. on rocky slope, liner. 
\\\T > A raucarta (2, QMMO\h4U. 17 Sept 1985, J, 
Stanisic. D. Poller); Kroombit Tops. 13 km W of 
"Chapman's . NVF (24'2?'S I51*02*E) (I. 

QMMO] 1222, 6 s pi [98 ! ABRS - \M/QM)S Port 

ifl (3. AMt ■ ■ : 2. [ \ COX): near Colosseum Ck. 
[6 km S Ol Miriam Vale (4. AMC 142446. 24 May 
IQ5& L. Price); Miriam Vale (I, AMC152447. ex 



Shell depressed, modci;ilcl\ huue, 5 97-7.39 
mm (mean 6.53 mm) in diameter, with 4 1/2 to 
4 3/4 (mean 4 5/8-) normally railed whmls. 
Apex and spire moderately elevated, height of 
shell 3.28-5.35 mm {mean'3.S3 mm). HD ratio 
0.53-0.83 (mean 0.59) Bodv whorl descending 
only slightly in front, SP/BWW r;itio O.IK-0 32 
(mean 0.24). Protoconch Hat. shiny, with 1 5/8 
to I 5 '8+ whorls, mean diameter 1 1 69.0p.rn at 1 
1/2 whorls. Apical sculpture of very weak, low, 
curved radial ridges crossed by irregularly 
spaced incised spiral lines (Fig. 121). Post 
nuclear sculpture of numerous, regularly spaced, 
low, broad, protractivcly sinuatcd radial ribs 
(Fig. I2e). whose interstices arc about 7-9 times 
their width. No intervening microriblets. 
Microsculpiure of incised spiral lines (Fig. 1 2g). 
Sculpture continuing onto base (Fig. 12h) with 
spirals fading toward the umbilicus. Umbilicus 
small, diameTcr 0.43-0.94 mm (mean 0.71), D/U 
ratio 7.2 1 - 1 5.63 (mean 9.40). Sutures distinctly 
channelled. Whorls rounded above and below 
the periphery (Fig. 1 2c). Body whorl without any 
not ice able angulation, even in juvenile 
specimens. Aperture ovately lunate. Lip simple, 
becoming thickened at the baso-columcllar mar- 
gin. Columella thickened, twisted and slightly 
reflected over the umbilicus. Parietal callus pus- 
tulose with some flammulations. Colour deep 
golden-horn with regularly arranged reddish- 
brown flammulations above, continuing below 
ihe periphery but lading near the umbilicus. 
Based on 21 rncasuicd specimens. 

Vas deferens (Fig. 13a) a very thin tube deli- 
cate I v convoluted along its length. Fptphallus 
(Fig. 13a) very long, muscularised, reflcxed half 
way along its length, internally with longitudinal 
pilasters, entering penis apically (Fig. 13b) 
through a conical verge (Fig. 13b). Pcnial retrac- 
lor muscle long, inserted at the epiphallus-penis 
junction. Penis (Fig. 13b) a cylindrical, mus- 
culansed tube swollen apically, internally with 
an apical fleshy collar and low. corrugated, spon- 
gy longitudinal pilasters. Free oviduct (Fig. 13a) 
strongly musculuriscd, about 3-4 times the 
length of the vagina Vagina very short, internal- 
ly with lleshv longitudinal lamellae. Atrium 
short without unusual features. 



Central tooth of radula (Fig. 12d) Willi broad 
triangular mesocone and small cctocones. 

Based on 6 dissected specimens (QM- 
MOI489G, OMM01M68, QMM016469). 

RwtiK -wv Habitat 

Ngairea canaliculata inhabils warm moist 
subtropical notophyll vine forests of me ranges 
and mountain caps between Colosseum Creek 
and Ml LuTCom, SEQ. This region also has tetge 
areas of drier sub-tropical forest (vine thickets 
and microphyll vine forests) but ;V. canaliculaiii 
shows little tendency to occur outside of the 
moister areas, The species has been collected 
live under bark and in the rolled up stems of 
discarded palm fronds. 


This species was first collected by Sidivv W. 
Jackson in 1908, but included under h hmimulinu 
coriicicola (Cox. 1806) by Hedley (1912). It is 
distinguishable from that species hv its chan- 
nelled sutures and radially sculptured base. The 
Ji:.Mii ■.'II- I .liturc is an unusual development 
seen lit a limited number of Australian, New 
Zealand and Pacific Island species. 

Ngairea corticicola (Cox. 1866) comb, now 
(Figs 15-17: Tables 2. 4) 

lh-hx vorticiaxltl Cox. 1866a. p. 37-1 ;C nx, L86S. pr, 

i" pl,7.figs7-l& 
I ':. \kera variicicoki iCovi; Iredale. 1937a, p. ."MS: 

[re-dale, I94ta»p. 264. 

( (Atr ■ i , 

\. CQrticlcala can he distinguished from A 1 . 
dorrigotnsis by its less proirjinem radial sculp- 
ture, almost smooth base, more rounded 
periphery, microsculpture ol incised Spiral 
grooves (Fig. 151) and penis with corrugated, 
longitudinal pilasters and a continuous, weakly 
muscular, apical collar V* canaliculata (Fig. 12) 
has similar microsculpture to N. corucicola but 
differs in having low broad radial ridges, fewer 
whorls, less elevated spire, channelled SUtUTCS 
mare rounded shell periphery and an epiphallus 
almost twice as long as the penis (Fig. 16a), 

.• ■< - i " '- Mi DTK 

Hedley ( 1912) attempted to clarify //c/Y.v cor- 
in tenia Cox. 1S66 but he considered specimens 
from Canungra. SEQ. to be conspecii'ic and 
figured a representative. This was the species 
later described by Iredale (1941a) as Orvokcru 

dorri$nensis. ,Y, contrrrnla occurs microsym- 
patrically with .V. dorrtgoensis at Pine Creek. 
Canungra, but the two species can be distin- 
guished by sculpture on the base of shell. Which 
is reduced in the former and strongly radial in the 

T'i i" M vn-K! \i 

\Jt TOTYP& \MCM520. Lasmore, NSW { o\ I fll 
leeiinn Height ot shell 3,87 mm. diameter 6 47 mm. 
H/l> ratio O.NI. D/U rutio I0.7K. w horls 5. 

PaMI i < 'ifivri s AMC6353 i 3 spci in eir same 

dala as kvlotypt;. 


ni w stJt'TM w \i E&i *olloh&b |1 

AMC 142431. Aug U'05): Richmond River (4, 
AMCM2432. e\ Cox. Fetterd). Rous Mill. Richmond 
Rx ,ih. AMC 142433, 1 900, S A\ . Jackson ); Lismore, 
RjchlflOnd Rv. s (% AMC 14:434, e\ Brazier); Rich- 
mond Rv. (I. AMC 142435. e\ Helms): upper Tweed 
River (7. AMC142426, eje Cox, Petierd); Lawanns 
near DorrigO 1. 1 . AMCA3BJ6. M Ward): Bomgall CI. 
Bellinger River, under hark ( I. AMC 142437. ex C..\ .; 
Opposite Kellv selection, lokm from Murwiliurnl 
in gtilh on main road i.v AMO_^S35U Red Scrub 
■ i 1 1 ■■ tiBi & hiofl S.Fh 210 m (28"3S S 
153 L9 , E)(13,AMC142473, 15 May 1976, F.H.C11I 
man I I 01 M tetania* net* Whran Wqj«i s I 
m (2fi WS< t53*19"E) M. AM< 142474., Ifi Mny 
1976. P.H. ( oinun. I. Loch); Dorrlgo Range, t-l. 
am( 142438, Apr 1957, 1 - Price) Bflllfna in fom n 
wuod M0, AMC14:43^. ex Cox): Big Scrub. WWafl 
Whiau stale Fbrest. litter, SNVF 1 1, QMMQI6441, 
15 NOV. I9K3 ABRS - AMOM1: Upper Wilson s 
Creek, liilfcr, NVF f28*32*S, L53 - 24 f E) (I. 
QMMOI&44& lo No\ 1983, ABRS - AMQM). 
Nightcap Range, under bark of laden tree [2{J S, 34 , S, 
153*2U*E] &QMM01646&, 15 Nov 1983, ABRS - 

AM'tjM 1 I || I H i|0 N IV, under |6g, I W)l 

lempcrate rainforest (3u :r:0"S 4 l52 : 43'4(fE'i (I. 
QMM014766, \2 Nov i')S3. ABRS - AM.'QM). 
Sw an's R«>jd. Bruvner Park Flora Reserve, underb.irk 
of (alien Iocs and in pahn irond:- nn ground, subtropi- 
cal rainforest (2X QMM014767, QMMQLfiT^T, 14 
Nov. 1983, ABRS - AM QM): Wonlgnolga Ck Flora 
Rtserv?, \\eddnit: Bells S.F.. under leaf in siihtropiv g| 
rainlortrsltl.AMC 1?::52. Mar I«»S3. M. Shea). Cut - 
rumhin Vallev Nature Reserve, under log inrainfo 
(3. AMCl5::54.,lat [977.M. She.,): LfsPioreS,. lb 
Lismore, under bark of fallen irecs, remnant rainforest 
(4.QMMUPI'\^M;i- l ( JS7. J. Sianisic, D. Pultcr). 
Wotnja Walk, rtw Glade^ Doiri^uN.P., under bark of 
lalkn tree m Fhrcsl (35, OMMOI7I46, fi Mar 
WS7.J. Stanistc. IX Poller |, Never Never 1'ienie Area, 







h £ r 

» * 

1 > 








FIG. 16. Ngairea coriicicola (Cox, 1866). Slopes of Mt Hobwee. Lamington N.P., SEQ. QMM016462. a, 
genitalia; b, detailsof penis interior; c T talon; a\ hermaphroditic duct; e, ovoleslis; f, pallia) cavity. Scale lines 
as marked. 

FIG. 15. Ngairea cortkicola (Cox. 1866). a-b, Lismore, NSW AMC63520, holotvpe; c, Tullawallal Circuit, 
Lamington N.P., SEQ. QMM016465: e-g. Red Scrub F.R. T Whian Whian S.F./NSW. AMC142473; d ? h, 
Slopes of Ml Hobwee, Lamington N.P., SEQ. a-c, entire shell; d. central (bottom left) and lateral teeth; e, 
apical sculpture; f, post nuclear sculpture; g, details of umbilicus; h, lateromarginal teeth. Scale lines as marked. 







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12^22 522-- ?S 2-2 32 I- S U S5 1^ to 30 a>o ^3 n c^ S bSJo ^o 











153 C S 


FIG. 17. 

Distribution of Ngairea corticicola (Cox, 

Dorrigo N.P., under bark of fallen tree, rainforest ( 1 , 
QMM017142. 6 Mar 1987, J. Stanisic. D. Potter); 
Booyong Flora Reserve, via Lismore, under bark of 
fallen trees, rainforest (21. QMMO 17005. 9 Mar 
1987. J. Stanisic. D. Potter). 

Falls Circuit. Lamington N.P.. litter, NVF (3. 
QMM016447, 3 Oct 1985, J. Stanisic. D. Potter); 
Tullawallal Circuit, Lamington N.P.. CNVF (3, 
QMMO 16465, 2 Oct 1985, J. Stanisic. D. Potter); 
Warrie Circuit. Springbrook N.P.. under bark of fallen 
tree. CNVF (4, QMM016464. 6 Apr 1986, J. 
Stanisic); Orchid Bower, Lamington N.P.. 900 m. 
CNVF (S. QMMOo264, Apr 1976, M.J. Bishop); 
Nagarigoon Falls Circuit, Lamington N.P., CNVF (2. 
QMMO16463.20Apr 19S6. J. Stanisic); Warrie N. P.. 
Springbrook. CNVF (28°0Q'S. 153*00'E) 15. 
QMMO6300,4Oct 1976, M.J, Bishop); Nagarigoon. 
Lamington N.P.. BOO m, CNVF (2S°14"S. 15.VLVE) 
(13. QMMO60S5. Apr 1976, M.J. Bishop); Ml. Hob- 
wee Circuit. Lamington N. P., litter (3, QMMO 1 6445. 

1 Oct 1985, J. Stanisic, D. Potter); Natural Bridge 
N.P.,NVF(28°13"S, 153°14'E) (1. QMMO10456, 18 
Mar 1981. ABRS - AM/QM); Tullawallal Circuit, 
Lamington N.P.. litter, NVF (1, QMMO 16446, 2 Oct 
1986, J. Stanisic, D. Potter); Coomera Falls, 
Lamington N.P.. under bark of fallen tree, CNVF (2, 
QMMO 16461,19 Apr 1986. J, Stanisic): SlopesofMt 
Hobwee. Lamington N.P., under bark of fallen trees, 
CNVF (17, QMM016462, 19 Apr 1986, J. Stanisic); 
Upper Pine Creek, Canungra. under bark, 
N\T/Araucaria (1, QMMO 16545. I Oct 1986, J. 
Stanisic, D. Potter; 7. QMMO16903. 21 Jan 1987, J. 
Stanisic, D. Potter: 3. QMM0165S4, 29 Sept 1986, J. 
Stanisic, J. Chaseling); Araucaria Track, Binna Burra, 
under bark. NVF (5? QMMO 16577. 27 Sept 1986. J. 
Stanisic): Lower Ballanjui Falls Track. Binna Burra, 
Lamington N.P., under bark (5, QMMO 16550, 25 
Sept 1986. J. Stanisic); Coomera Falls Circuit, Binna 
Burra, Lamington N.P., under bark, NVF (11, QM- 
M016574. 26^Sept 1986J. Stanisic). 


Shell depressed, large, diameter 5.88-7.82 mm 
(mean 6.90 mm), with 4 3/8 to 5 1/2 (mean 4 
3/4+) normally coiled whorls. Apex and spire 
moderately elevated, height of shell 3.36-4.62 
mm (mean 3.90 mm). H/D ratio 0.50-0.63 (mean 
0.56). Body whorl descending slightly in front, 
SP/BWW ratio 0.22-0.39 (mean 0.31). 
Protoconch with 1 5/8 to 2 whorls, mean 
diameter 1056. 8u.m at 1 1/2 whorls. Apical 
sculpture of curved radial ribs crossed by incised 
spiral lines, most noticeable near the sutures 
(Fig. 15e). Post nuclear sculpture of low, weak 
protractivcly sinuated radial ribs and ridges, oc- 
casionally with a vertical periostracal blade (Fig. 
15a). interstices 3-5 times their width and 
without radial riblets. Microsculpture of incised 
spiral grooves (Fig. 15f)- Sculpture reduced on 
base to almost smooth (Fig. 15b). Base shiny to 
dull. Umbilicus small, 0.55-1 .02 mm (mean 0.8 1 
mm) in diameter, D/U ratio 6.99-11.31 (mean 
8.61). Sutures impressed. Whorls flattened 
above and rounded below the periphery. Body 
whorl with a weak keel (Fig. 15c). becoming less 
pronounced in larger specimens. Aperture ovate- 
ly lunate. Lip simple, slightly thickened at the 
basal margin, becoming more so at the columel- 
la. Columella thickened, twisted and slightly 
reflected over the umbilicus. Parietal callus pus- 
tulose. Colour creamy-white to yellow-horn, 
upper surface with regular brown-red flammula- 
tions fading at the periphery, base creamy-white. 
Based on 34 measured specimens. 

Vas deferens giving rise to a lone, muscular 


cplphallus (f ig. 16a), Fpiphallus rcflcxing api- 
cally to enter (he penis through a simple pore 
(Fig. Ihb) surrounded by a Spongy pilasler. in- 
ternally with longitudinal pilasters. Penis (Fig. 
\hh) slender, cylindrical* muscular, slight I J 
swollen apicallv. internally with an apieal fleshy 
collar and strongly corrugated, longitudinal 
pilasters. Penial retractor muscle long, inserting 
on the epiphallus prior to its entry into the penis. 

Radula (Fig. 15d,h) with broad triangular 
mesocone and reduced ectocones on central 

Based on four dissected specimens (QM- 
MO 16462, QMMOA33U). 

Ram;f and Habitat 

A. conicicola inhabits warm subtropical 
notophyll vine forests of the coastal lowlands 
and foothills between Dorrigo and Lismorc. the 
Dorrigo Plateau, and the eastern MePherson 
Ranges. Its northern limit appears to be MtTam- 
borine, SEQ. On the basts oi material collected 
lute last century and early (his century, it would 
appear that the Big Scrub formed an important 
part of the range of A', corticicola. In spite of the 
fact that much of the Big Scrub was cleared by 
early cedar-getters and more recently for agricul- 
ture, A', corticicola is still common in some of 
the Temnants e.g. Booyong Flora Reserve, near 
Lismorc. NSW. The species lives under the bark 
of rotting logs. 


Southern representatives ot ,\\ corticicola 
(QMMOI7I46, AMC142438) have compara- 
tivcls lower whorl counts than northern 

m riTncns{Table4). 

Mussonula Iredalc, 1937 

Vfft n>nu!tt Ircdale OCwhp I J 

TYP1 5PS1 irs 

Q&liSSQttttfa \crax Iredalc 1937; by original 


Iredalc ( 1 c > 3 7 h ) related Mltssomfa to 

I'untlaoma Iredalc, 1913 which is a punetid 
grOUp V ith a spirally lirate protoconch, Although 
Iredalc ( lv37b) noted spirals in M. wnn he 

I tiled to notice (he less conspicuous apical radial 
elements referred to in rhis study (Figs 184 2Qf)- 

Combined with general shell sculpture and 

details of the radula (Fig. 2Qd, c), the apical 
sculpture places Mussonula in the Champidat 


Shell small to moderately large, trochoid, adult 
shell diamcier range 4.S 1-5.63 mm, with 3 7/8 to 
4 1/K normally coiled whorls, last whorl des- 
cending. Height of shell 3.32-4.45 mm. Apex 
and spire strongly elevated. Protoconch strongly 
exsert. Apical sculpture of numerous bold 
(vcmv) to fine {fa I lax) spiral cords, and very 
weak radial ridges which become more 
prominent near the nuclear-post nuclear junc- 
tion. Post apical sculpture of numerous, high. 
proiraclively sinuated radial ribs with periostra- 
cal blades inclined toward the shell apex. Some 
trace of weak incised spiral lines (Jullax) but no 
microriblets present. Sculpture continuous on 
the base. Umbilicus narrow U-shaped. Sutures 
impressed. Whorls rounded above and below a 
strongly angulate periphery. Aperture sub- 
quadrate. Lip simple, columella reflected slight- 
ly toward umbilicus. Colour yellow-horn to 
brown with adhering dirt particles. 

Pallia! cavity with incomplete secondary 
ureter and kidney with vestigial rectal lobe. 
Genitalia unknown. Radula as for Mussonula 
fa I lux, 

Distribution and Ecolqcr 

Mussonula extends from Byangum on the 
Richmond River, northern NSW to the Gympie 
forestry area, SEQ. It inhabits moist lowland 
notophyll vine forest in the south, and drier 
microplwll vine Fores! forilhArQUC&fict) and vine 
ihickci m the north The two species arc al- 

Little is known of the micmhabit for although 
CI". Musson collected (he type series of M, 
vcrax arboreally. and a single live specimen o! 
A/, fallax was found under timber among rocks, 
all other specimens have come from litter 
samples The trochoid shell and dark animal 
colour indicate an arboreal or semi-arboreal ex- 
istence. However the adhering dirt particles on 
the shell suggest thai it is a ground dweller. 
Possibly it lives among friable earth, between 
and under rocks, and forages for food on vertical 
surfaces above the ground. 


The extremely prominent radial ribs (Figs ISe, 
20g), strongly angulate periphery, elevated 
spire, exsert protoconch with strong spiral cords 
(Figs INd. 201"). and almost total lack of micros- 





1 — s 

": : ::::- 

e **+? 

FIG 18. Mussonula wax Iredale, 1937. a-c. Scrub at North Pine, SEQ. AMC63770. holotype; d-f, c. 2km 
E of summit. Ml Nebo, SEQ. QMM0165 1 8. a-c, entire shell; d, apical sculpture; e, post nuclear sculpture; 
f, terminal region of protoconch. Scale lines as marked. 



culpturc in Mi/ssonuUi are a combination of fea- 
tures not found in any other cast coast charopid. 
Orcokera Stanisic, 1 987, from north Queensland 
has the same shape and postnuclcar ribbing as 
MussOrtulabWX is larger and has quite prominent. 
reticulate mtcrosculpturc (Stanisic. 1987, pis 
\ c,&,2c Ax). Hedlevoconcha Pilsbry. 1893 has a 
similar shaped shell but has strong reticulate 
Sculpture and a punctate protoconch (Figs 23e, 
f: Z6d). 

Mussonula verax lrcdale. 1937 
(Figs 18-19: Tables 2.5) 

Thahtssia da Ita (Pfeiffer); Heiltey and Mutton, 1892, 

p. 553 - misfdeiflif i cai ion . 
Mu$sontita \*6rax I red ale, 1937b, p. 13, pi. I. fig. ) I. 

■ I ■ 

Mi verax can be distinguished from M. j a I tax 
by its largl i si/e and bolder, more regular, apical 
spiral cords (Ftgs 1 8'd, 20f). Nijairco levico.statu 
a similar shell to M. verax but is slightly 
i r with apical sculpture of curved radial ribs 
and weak spiral cords (Fig. 6d); A. domgoenSis 
which is sympatric with M. vcrax in the southern 
part of the D'Agu iiar Range, SEQ. is larger; has 
a less elevated spire and mieroseulpture oi 
prominent spiral cords (Fig. 3c). 

I '-H \1 \TFR[.\i 

HQiQTYPE: AMC635UI, in scrubs, arboreal North 
Pine River, South Queensland Call CT. Musson, 
Height of shell 4.54 mm. diameter 5.46 mm. H D ratio 
82, D U rotto 133, whorls 3 7/S+, 

PARATVfteS: AMC6377D, 5 specimens. Scrub at 
North Pine, SEO. Coll. by CT, Mussai*. 

Othvr M VTVRI \l 

t ; L d> Rd. Ml Mue. SEQ. rainforest, leaf tiller. 
(27 05 S. 152 43*E)(hQMMO7340 I !4Api I980.J, 
SuinWc N. Hall, A. Green); Mi Niebo, SEQ, c 2 km 

E "I summit. ^Vf/ArHtuCttrm la 27 M-'S, 

1 52*47 F) (I- DMMai65ia 13 Aug i960 I 
Sumisic); Wi, men's t amp, WraiknsS h.SEO-NVF 
litter t:d ITS. is: 7H I ) (I. QMMQU5QU. 17 Jul 
I'^o. .). Stanisic. A. Green); c. 0.5 km B of Mt Mia 
lurnolT on Ml Mia - Rilkoan Rd. SEQ tf6 a T5"S, 
152 27TM1.OMMO12320. I Sepl I9R2. AM/QM - 
amks |QK2] Maurta N,l\, Mi Glorious, SEQ, wi 

(],0MMO) !f'O2.20Jun NS2. MSA Party). 

Shell trochoid, small, diameter 5 I U-5 03 mm 


!6 D S 



f] r 152 C E 153°E 


FIG. 19. Distribution of Mussonula verax Iredale, 


(mean 5.36 mm) with 3 7 8to4(iwean4-)1ao8dy 
coiled whorls. Apex and spire strongly elevated, 
heigh! of shell ^03^-4.45 mm (mean 4.31 mm), 
II D ratio 0.79-0;82 (mean 0,81). Body whorl 
descending more rapidly. SP 13WW ratio 0.52- 
57 (mean 0,54). Protoconch strongly exsert 
with 1 I 2 to I 1 7A Whorls, mean diameter 
1371). 5 |am at 1 1/2 whorls. Apical sculpture of 
about 20 continuous spiral cords and curved 
radial ridges appearing in the las! I <S whorl (Fig. 
ISd). Post nuclear sculpture of high, crowded 
proiructivclv sinuatcd radial ribs, 90- 1 20 (mean 
104 8) on the body whorl (Fig. ISe). Ribs mm 
5.6S-0 78 (mean 6.20) Majoi ribs with slender 
backward curved pertostraeal Wades, interstices 

"-20 times their width, no radial rtbiets. Mieros- 
eulpture absent. Sculpture continuous on the 
base (Fig, 1Mb). Umbilicus very narrow. U- 
shaped, slighllj obscured, diameter 0.31-0 1] 
mm (mean 0.35 mm), D U ratio 13.29-17-19 
(mean 15.27). Sutures impressed. Whorls 
rounded above and below an ungulate periphery 
Aperture sufoquadniie. Lip simple, columella 
twisted, vertical. Siigl h I ovenngthe umbilicus. 
Parietal callus pustuiosc. well developed. Colour 

yellow-horn w itfi adhering din particles. Based 

on 4 measured adults. Anatonn unknown. 

= b $ 

2 s- 

Z « 


Range and Habitat 

Af. xerox inhabits notophyll vine forests of the 
D'Aguilar Range and drier microphyll vine 
forests of the Gympic region. SEQ. 

= ~ 


s C.J. Musson collected this species during an 

expedition to SEQ in i SS7 which was funded by 

noted malacologist Dr J.C. Cox. The species was 

|g misidentified as Thalassia delta (Pfeiffcr, 1857) 

| 3 || 5 by Hedley and Musson (1892). Iredale (1937b, 

| 3s p. 13) noted the error and erected the genus. 

Although the type locality of the species is given 
broadly as "scrubs at North Pine River", the 
3« itinerary of Musson T s trip suggests that a more 

n specific locality may be Mt Mee. SEQ. 


fi >- a s r — t 

r X 

Mussonula fallax sp. nov. 
(Figs 20-22; Tables 2, 5) 


Latin fallax. fallacious; referring to its having 
been previously identified as \f. verax. 


< A/, fallax differs from A/, veraxby havingfiner, 

-.t | |= more irregular apical spiral cords (Fig. 201*) and 

^<|H?r?i*5 pi s a smaller shell. Hedleyoconcha delta (Pfeiffcr, 

1 1 ' 1 " 1857) has a trochoid shell but is much larger with 

^ i greatly elevated spire and has postnuclear sculp- 

^* = Bc ture of low radial ribs and spiral cords in a 

1 1 < 1 5 reticulate pattern (Fig. 23e, f). 

g c k XI » ^ « i: 

5 S ■— o = . = o o 

I Type Material 

| k Hoi.OTYPE: AMC63870. No. 1 Scrub at Byangum, 

^ £ _ NSW, Cox coll. Height of shell 3.41 mm, diameter 

% i| * 12 % a s 4.32. H/D ratio 0.64. D/U ratio 6.18, whorls 4 1/8-. 

^ 5 PARATYPES: Mt Warning N.P.. NSW (28*24'$, 

I 153 16E) (1. AMC129289, IS Mar 1981, ABRS - 

k _ f ? AM/OM): Upper Coopers Ck, Huonbrook, NSW (1. 

1 | J s |3 ^ . h AMC 142449, 26 Aug 1936. A. Musgrave, E. 

5 ; i, ^ Troughton); Burleigh Heads N.P.. SEQ, vine thicket 

(4, QMM06157, QMMO6160, Oct 1976, M.J. 

„ Bishop); Terania Ck. NENSW. NVF. under piece of 

i | wood among rocks (1, QMM017323, Aug 1987, J. 

I £ *t - I I Stanisic); Richmond Rv.. NSW (1, AMC142448). 

- " - Shell small, trochoid, diameter 4.81-5.32 mm 

(mean 5.07 mm) with about 4 1/8- normally 
coiled whorls, last whorl only slightly descend- 
u| s w _ _ ing. Apex and spire strongly elevated. SP BWW 

§|§f| ratio 0.39-0.45 (mean 0.42), height 3.32-3.41 

lllll 11 1 |gi |» mm (mean 3.37). H/D ratio 0.64-0.69 (mean 

|I< i° j£< i< o.67). Protoconch exsert. slightly shiny with 1 






,: ■ ■■■■■■ 





'-■■ '■■■■MP- 



v* t 




K h HV 



FIG 21. Mussotrufa ftillax sp. nov. Teraniu Creek, 
NSW, QMM017325. puraiype. Pallial cavity. 
Seal* fine as marked. 

12-1 5/S- whorls, mean diameter 1 14ft.5jXlrt at 
1 12 whorls. Apical sculpture of 20-25 inter- 
rupted spiral cords with numerous, low. curved 
rndial ridges forming toward the end of the 
protoconch (Fig. 2(H), Post nuclear sculpture of 
numerous, high, protracm el\ sinuated radial 
ribs, with periosteal blades which are folded 
toward the apex {Fig. 20g). No microriblets. 
Microsculpture consisting of vague incised 
spiral lines (Fig. 20g), Sculpture continued on 
the base (Fig. 20b). Rib interstices variable, 5- 15 
times the width of the ribs. Umbilicus narrow, 
V-shaped, diameter 4.81-5.32 mm (mean 5.07 
mm), D/Lf ratio 5.86-6. 18 (mean 6.02). Sutures 
impressed. Whorls rounded above and below a 
strongly angulate periphery. Lip simple. 
columella dilated and slightly twisted toward the 
umbilicus. Aperture subquadrate. Parietal callus 
strongly developed. Colour brown, often 
covered with dirt particles. Based on 2 measured 

Foot and tail short and broad, bluntly tapering 
posteriorly. Body colour dark with sides of foot 
and outer margins of the sole purple-grey. Upper 
body and neck black. Ommatophores purplc- 

FIG. 20. Mussortttla falla.x ;sp. nov. a-c, No. 1 scrub 
at Bvangurn. NSW, AMC63870. hololvpc; d-e. 
Terania Ck, NSW. QMMO 17323, paratype; f-h, 
Burleigh Heads N. P., SEQ. QMMO 615 7,pawiype. 

a-c, entire shell; d, central and lateral teeth: e. mar- 
ginal teeth: I. apical sculpture; g. post nuclear sculp- 
ture; h, details of post nuclear microsculpture. Scale 
tines as marked. 

grey. Pallial roof black. Pallial ca\il\ occupying 
c. ill whorl apic-ally. Mantle collar well 
developed Wtth large glandular /one extending 
onto the lung root. Kidne\ with triangular 
pericardial lobe and rectal lobe present as a tin) 1 
vesttge abutting hindgut. Ureter sigmurethrous 
without complete secondary branch. Ureteric 
pore situated adjacent to the hindgut. anterior to 
the small rectal lobe of the kidney (Fig. 21), 
Reproductive organs immature. 

Central looth of radula (Fig. 20d.c) with lan- 
ceolate mesocone and small, pointed eetocones; 
inner laterals with endocone absent and more 
prominem ectocone. Outer laterals and mar- 
ginals typical. 

Based on one dissected subadult (QIY1- 
MO 17323). 

R\\0> wo Habitat 

M. fallax inhabits the notophyll viffe forests of 
the moist foothills of coastal northern NSW. and 
the unusual coasuil vine thicket at Burleigh 
Meads. SFQ. Its microhabitat is not fully known. 


In the absence of data on the terminal genitalia 
the two species are separated on the difference 
in protoconch sculpture. Habitat destruction, 
particular!) in the Big Scrub area of north-east- 
ern NSW, has had significant effects on the 
distribution of lit. fallax. Iredale (1941b T p. 2) 
cited specimens of M. vcrax from the Richr.; 
River, northern NSW'. These are A/, fallax. 


2 c J-i 

l e : r E 


f IG,22. Distribution of Mussomtla fallax sp. nov. 



Hedleyoconcha Pilsbry, 1893 

H 'edlevoconcha Pilsbry, 1893, p. 18; Iredale, 1937a, p. 
321; Iredale, 1937b. p. 13; Iredale, 1941a, p. 264; 
Iredale, 1942. p. 35; Iredale. 1944. p. 317; Burch 
1976, p. 133. 

Type Species 

Helix delta Pfeiffer, 1857; by original desig- 

Previous Studies 

Pilsbry (1893) placed Hedleyoconcha in the 
Flammulinidae where it remained until Iredale 
(1942) erected the Hedleyoconchidae for it. 
Sol em (1983) included Hedleyoconchidae with 
his expanded Charopidae. I follow Solem and 
consider that Hedleyoconcha can be derived 
from more generalised charopids. 


Shell relatively large, diameter 5.38- 10.08 mm 
with 5 1 /8 to 6 5/8 whorls. Spire and apex strong- 
ly elevated (Figs 23c, 26c), height of shell 4.62- 
6,47 mm. last whorl descending. Protoconch of 
1 1/2 to 1 3/4 whorls, pitted (Fig. 23e). Post 
nuclear sculpture of low, protractively sinuated, 
radial ribs and low rounded spiral cords in a 
reticulate pattern (Figs 23f, 26d). Sculpture al- 
tered after 4 whorls, continuous on base but 
much less pronounced (Figs 23b, 26b). Um- 
bilicus narrow, partially obscured by the 
reflected columella. Sutures lightly impressed. 
Whorls slightly flattened above and rounded 
below a strongly angulate to thread-carinate 
periphery. Aperture ovatcly lunate. Lip simple, 
columella dilated and slightly reflected over the 
umbilicus. Colour straw to cream-yellow with a 
darker spire. Periostracum deciduous. 

Foot and tail broad, rounded posteriorly. 
Caudal horn present. Colour creamy-white with 
some pigmentation on the pallia! roof. Om- 
matophores black. Kidney elongate, vaguely tri- 
angular with a reduced rectal lobe. Ureter 
sigmurethrous with an incomplete secondary 
branch. Ureteric opening in the angle between 
the kidney lobes. Ovotestis several clumps of 
palmately clavate lobes of alveoli, oriented at 
right angles to the plane of coiling. Terminal 
male genitalia with thin vas deferens and large, 
refiexed, muscular epiphallus. entering penis 
through a large fleshy pilaster opposite a 
grooved muscular thickening which acts as a 
collar. Penis proper with long slender corrugated 

pilasters. Penial retractor muscle inserted at the 
penis-epiphallus junction. Radula with tricuspid 
central and lateral teeth which have long 
mesocones, short endo- and ectocones, and well- 
developed anterior flares; marginals typical. 

Distribution and Ecology 

Hedleyoconcha has a classic relict distribution 
with a widespread species in northern NSW and 
southern Queensland, an akitudinally restricted 
species on the summit of Mt Bellenden-Ker in 
north Queensland and a further species on Lord 
Howe Island, off the coast of New South Wales. 
Although similar distribution patterns are seen 
among some flightless groups of insects such as 
the Peloridiidae^Monteith, 1980) this is the first 
time a land snail group has been identified as 
having such an unusual and biogeographically 
complex distribution. 


The high spire, punctate apical sculpture (Fig. 
23e) and distinctive low reticulate post nuclear 
sculpture (Figs 23f, 26d) distinguish Hed- 
leyoconcha from other subtropical and tropical 
charopids. Oreokera Stanisic, 1987 has an 
elevated spire but apical sculpture of spiral 
cords, prominent post nuclear radial ribs and 
fine, reticulate microsculpture (Stanisic, 1987, 
pis 1,2). Mussonula is much smaller and has 
prominent apical spiral cords (Figs 18d,20f)and 
crowded, high, curved radial ribs on the post 
nuclear whorls (Figs 18e. 20g). Ngairea lacks 
the punctate apical sculpture seen in Hed- 
leyoconcha and has much finer post apical sculp- 
ture (Figs3e, 6e, 9g, 12e, 15f). 

The species show minor differences in shell 
features but are readily segregated on the basis 
of geography. 


1. Shell with umbilicus wider (Fig. 26g), mean 
D/U ratio 4.60; Lord Howe Id addita 

Shell with umbilicus narrower, mean D/U 
ratio greater than 6.50 2 

2. Shell with strong reticulate sculpture on early 
whorls (Fig. 26d): protoconch weakly 
punctate (Fig. 26d); Mt Bellenden Ker, NEQ 



Shell with vseviker reticulate sculpture on 
early whorls (Fig. 231); protoconch Mronglv 
puncLitelFtg. 23c); southern Queem-Jand and 
northern New South Wales dcltu 

Hedlcvoconcha delta (Pfeiffcr. 1857) 
(Figs 23-25; Tables 2 6) 

Itedetia Pfciffcr, tfl57 ( p 3S< 

ffcfe conoidca Cox, 1864, p. 21; non Draparnaud, 

Helix fenestrate Cox, 1866. p. 374; non Sowerhv, 

Charopa {Thalasaia) delta ( Pleil icr): Tnon, 1886, p. 

215. p!, 63. fig, 76, 
Flamrnutina I Hcdlevocotuiia) delta (Pfeiffer); 

Pilsbry, K9& pp IS- 1 1 ) 
Hedk'Y'Koncha del fa (I frccfe! '' ' :■ 

321. Iredale, 1941a. p. 35. 
ficdlexnconchii duona Iredale, 1937b, p. 1 3 pi. 1 , fig, 

13: Iredale. 1 S>4 la. p. 35. tig. u. 


H. delta can be distinguished from //. 
ail&ketOQeby its larger size Jess promincntadult 
sculpture and more conspicuous apical sculp- 
ture. Anatomically the differences are minor 
with slightly more corrugated penial pilasters 
and a more prominent apical penial bulb present 
in H. tvlakefoae (Fig, 27a-b). //. delta is ftOl 
easily confused with any sympairic charopids. 
Mitssonulu vera* (Fig. 18). M. fat lax and 
\gatrea levieostata all have elevated shells ,u\e\ 
occur in the NSW-QLD border areas. However 
they have prominent, fine, crowded radial ribs 
loi the adult shell whorls (Figs 6e, ISe. 20g). 
spiral cords I with radials in N. lexieostata) On the 
protoconch (Figs 6d, 18d, 200- are smaller, and 
have a less prominent peripheral keel (Fie* i 
19c. 20c). Ngairea murphyi resembles H. deha 
n gross conchologfcal detail but is smaller, with 
more tightly coiled whorls, less elevated spire, 
more open umbilicus, finer adult sculpture and 
apical sculpture of fine pits and wrinkles (Fig 
L .J,i. b, e-g). 

H- delta is microsympatrie with several 
helicinids - Pleuropoma jana (Iredale, 1937) 
from the Port Macquarie area. NSW and P. 
dvaytonensis (Pfeiffer. 1857) from far northern 
NSW and southern Queensland. Both species 
have elevated shells similar in colour to //. delta 
and live on the leaves of low shrubs. However 
they may be distinguished in having strongly 
incised spiral grooves on the protoconch and 
adult whorls, and no umbilicus. The introduced 

and iV "I'spread Pacific helicarionid Coneup 
la ialrul(/\(i (GOtildi I851J is also microsym- 
patrie with //. delta in parts of northern NSW 
and southern Queensland but it is much smaller, 

transpareni vrllow-horn jfl colour, and has a 
microsculpture of numerous fine incised Spiral 



//. delta was originally described from the 
Drayton Range (= Toowoomba). SEQ. Cox 
(1864, 1866) introduced names for two NSW 
populations (eonotdea ftlld fawitrata) which h> 
later (Cox. 1868) synonymised with H. delta. 
Iredale 11937a) placed them in synonymy with 
Helix KUHuenS Cox. 1872, a helicarionid, then 
(1937b] realised the error but considered the 
NSW specimens specifically separate as //. 
duona on Ihe basis of their broader base, nar- 
rower umbilicus and weaker sculpture. 

Id this study spc cime n s of Hedlevoa mcha w ere 
dissected from numerous localities (including 
Wcntworth Falls, west of Sydney. NSW) and no 
perceptible differences in genital anatomy were 
noted. Differences m shell size and sculpture are 
mosaic and fall within the range of asinglewddcly 
distributed species. Hence Iredales separation is 
considered unsustainable and H. duona is placed 
in synonymy with //. delta. 


Lectotvpe; BMNHI98224& Drayton Ranee. N 

Australia Collccledhv Mr Siutchbury.ex II. Cuming 
OOllei IJOJl I 01 Shell i'.f'4 nun. dumeter 

mm, H/D ratio D/TJ ratio 7.67. whorls h J, ,4 

P\rai fctot> pfs: BMNH 1 982246, 2 specimens 
same collection data as leetotvpe. 

Hoti'UviT ol liif'ia: AMC63S26. Terriguh QCBl 
Gosford. NSW. No other data. Height of shell 6.30 
mm, diameter 840 mm. H'D ratio 0-75, D/U ratio 

10.2 1. whorls 6. 

t Mill R M ■ :. : I ,i 

MEW SOUTH WALES; Frank ttemL rerrigfl] ro 
lord, in nenih. on trees (2. A MC 142450. S.W. lack- 
s"ii, j.i Mji l953);DwigogS ( F„ i 

on ralien tree ( I. AMC142455, W. Dowling. 19 .lun 
1962): Usmore (1. AMC142458, luwenln v, i 
down Wentwonb Falls, rainforest bv ereek (I, AM 
C142453, D.K. McAIpinel: Sassafras Gullv, 
Springwood (2, AMC142452, C'.F. McLauehhm/s 
MAS 1 548): Doirigo Scrubs, on underside of leaves on 
i2, AMC142454 Dc< I^JO, S.W. Jackson); 
Byr«>n Bay scrubs {7 AMCl42457i L904.S.W, 
sunt: ! I.4 km S ol' Nvnihoida. on iiralton-Armidalt- 







/ % 









FIG. 24. Hedleyoconcha delta (Pfeiffer. 1 857). Dandabah. Bunya Mts. SEQ. OMMO606S. a, genitalia; b, details 
of penis/epiphallus region; c, details of penis interior; d, hermaphroditic duct; e, ovotestis; f. talon; g, pallial 

FIG 23. Hedleyoconcha delta (Pfeiffer, 1857). a-c, Drayton Range, North Australia. BMNH 1982246, 
lectotype; d. Terrigal, near Gosford, NSW. AMC63526, holotype of duona: e-h. Dandabah, Bunya Mts. SEQ. 
QMMO6068. a-d, entire shells; e, apical sculpture; f, post nuclear sculpture: g, central and lateral teeth; h. 
marginal teeth. Scale line as marked. 




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S> S*lt, > 300m 

U9»E 15CE lM'S I53*n 153"E 154°E 

FIG, 25 Distribution til \ Uedlexoameha delta (Ffeif- 
■ 1857), 

Rd. in rainforest beside roadi I. AMC1424SO .:i)M a \ 
: 176. P H, Coltuan & 1. Loch); WilsOtl River Reserve. 
via Belhnnrv (I. AMCI4245I, 25 S*p! L§Sl, O.K. 

MtAljJimj?): Richmond Rv (I amci424oi. o.x 

coll); Purl Stephens, on island in port {7. ANIC63730, 
Oct 1S92. e\ Bland, CX Cox)\ Port Macu,uaric (4. 
AMC36565. c\ Hedlevi, Col S Cave, t arrai S.l in 
|ilt$r entrance (I, AMC121W3, 2(J Fab 1980 
PH. Colman). Carta! I SVfti t arrfli S.F.. in liiler vine 
forest (S. AMCI21701. 21 Feb |980 PH Colman): 
'loonumbar Forest Road c. 34 km from Kyogle. 
■loonumbar S.F.. CNV4 (2B"33*S, 152'45'E) (1. 
\MC12850L 15 Mar 1981, AM/QM - ABRS). 
NcwfiM Fall& Oorrigo N.'P. <3ir24S. 15245T-) (1. 
AMCI2S337. 12 Mai IDS1, ABRS - AMQM); 
Acacia Plateau Road. C. 1 2.2 km from Acacia Ck-Kil- 
Lunev Road. Koreelah S.F., SEVT (28 2 i'S, 
2~4E,| (3. AMCI285f>2, OMMO109G0. [5 Mar 

1981. AM/QM - ABRS): Ml Warning N.P. (2S°24"S. 
]5346"L)(1, AMC12V290. 18 Mar 1981, AM/OM - 
ABRS »; Tooloom Scrub, Beaury S.F., NVF. (28*23*5 
L52*24"£) (6; AMC128538. QM MO 10941, 15 Mar 
I 98 I . AM/QM - ABRS); Wiangane S.F . c. 25 km Nb 
Kvogle. 1000 m (2S C 23'S, 1534ln'E) (2, 
AMC 4 42480, 17 Mav 1976, P.H. Colman. I. Loch); 
Koreelah Cfc, Beaury S.F.. c. 15 km W of'Urhenville, 
530 m (28*21*5. I52 d 2JTE)(3,AMC142477, 18 May 
1976, P.H Colman, 1. Loch); Cherry Tree North S.F. 
c.40kmvv at Ctaitia, 400 rn (28*54*5, 152"4yE)(24, 
A\K 142478. 19 Mav 1976, PH. Colman, I. Loch); 
Beaury S.F.. c. 15 km W of Urtalvflk, SKI m 
,■-> -7 S. 152"24 hll 7 .AM(" 142479, lSMa> 1 975 
P.M. Colman, 1. Loeh);0"Donnc1I Ck, nr Eden Creek, 
m N oj Kyogk f i. AMC131453, 13 May 1051, 
N. Moulds. C. Smothers); Ounmbali Scrubs, nr Go*>* 
ford | 1. AMCl52Io3.Gct I 904. S.W, Jackson); Nevv- 
, Rfitl^l I. AMC 152162, 1904.S.\V..lackson): Moonie 
Moome Creek, nr Goslotd (3. AMC4 42933, on tree 
trunk in rainforest, 6 Dec 1979 B Day; I, 
AMC142M_M.2o.1an I9S4. D. McAlpine); Wenlvvonli 

i-;, lis, Blue Mountains(2 AMC142938, i£Oct I 1 i 

D.K. McAlpine); Oibrallat Range (1.. QMMQ1 7256, 
Ml Suv 1980, G B Monteith; 2. QMM017257, 10 
Nov 1980, R. Raven); Boovong Scrub, via Lismore 
( :,S j>rs. 153 JO'! ML QMM06337, 14 Mav 1977, 
M ,1 Bishop); Richmond Range S.F. [2fl"41 S. 
15244 Fj N\ 14 17 n\1M(.)o7oo. I o Apr 1976. M J. 
Bishop); Llppct W ilsoiFs Creek (28'32'S, 153'24'E), 
in lilici (1. QMM0167»;i, [ft Nov 1983, AM/OM 
ABRS}i BeHfngen River (30"27"S, 152'32'E). rain- 
foresi ingulbt(l,QMMOlOB35.Mai 1981. AM/OM 
- ABRS)' Wemwnnh Falls (2, AMC26350. e.x C. 
Lascron); Shcrrard Falls. Dorrigo, 600 m. rainforest, 
in lite (I QMMOI6777 13 Nw\ l^S3. AM/QM - 
ABRS); Swans Rd, Banner Park Plata Reserve, lillet. 
NVF (/•'■ QMM01G783, 14 Nov [983, AM/OM 
ABRS): Whian Whian S.F., c. 7 km N Ol DorToueJihv. 
NVF (1. QMMP16788. 15 Nov los3. AM/OM 
ABRS): Sea Acres Nature Reserve, Port Macquane. 
littoral rafnforesi withpalmstLOMMOU^o^, odcc 

I9B3. P.M. Colman); BvrrillCreeUrilnr HJ tfl Melv 
hin forest road (2s:7S, 1554 2 l.)( I .OMMOHM07, 
Mai 1981 AM OM - ABRS), 

Ol'LFNSl AND: Near monument. Cunningham fiap 
N P N\4- (2.SM14S. I55"34"E)(.L AMCT286H, 16 
Mar iVtfl, *\M QM-ABRS); linnyj Mountains N f'.. 
il. \M( 14:4(4. 8 Feb |V61j D.F. McMicliacI): 
Coolabunia pine scrubs, nr Kingaroy, on pine trunks 
(2. OMC 14240(1, Jun I9(IS. SAV. Jackson: 5, AM- 
(142402. Jun Helms. SAW Jackson): 1 Ullc 
NertmgCVeekP,AMC55505,ex Brazier) Dflridabah. 
BunyaMls(26.ANH I360I7,OMM-OI2295 ( 31 Sepi 

los:, AM OM-ABRS). Halts Plain. E ol Emu Vale. 



CNVF. crawling on log (1, QM MO 1 6898, 4 May 
1986. J. Stanisic. J. Chaseling); Binna Burra, 
Lamington N.P.. crawling on 'cordylines' (4, QM- 
MO 14976. 10 Mar 1986. D.G. &. N.G. Poller): Ml 
Hob^ee. Laminglon N.P. (28°13*S. ]53"13'E), 1150 
m. MFF. (1. OMMO6077. Apr 1976. M.J. Bishop); 
Toowoomba (27 = 34'S, 15 1°57'E) rainforest. ( I , QM- 
M06276, 25 Sept 1976, R. Raven); Dandabah, Bunya 
Mts (26°53"S, 15I°35*E) Araucaria/NVF (90, QM- 
MO6068. FMNH 206312, 5 Mar 1976. M.J. Bishop); 
Upper Pine Creek, via Canungra, N\'¥/Araitcaria. 
litter (2. QMM016541. 1 OcM986. J. Slanisic. D. 
Potter); Tullawallal Circuit. Binna Burra, Lamington 
N.P., NVF. under logs (LQMMO 16736, 2 Oct 1985. 
J. Stanisic, D. Potter, J. Chaseling); Dandabah, Bunya 
Mts. behind picnic area (26 Q 53'S, 151 J 36'E)(1 1,AM- 
C136617. 31 Sept 1982, AM/QM-ABRS). 

Shell trochoid (Fig. 23c). diameter 6.22-10.08 

mm (mean 7.77 mm) with 4 5,cS-6 5/8 (mean 6 
1/8+) normally coiled whorls, last whorl des- 
cending. Apex and spire strongly elevated. 
SP/BWW ratio 0.36-0.88 (mean1l66), height 
5.04-7.98 mm (mean 6.01 mm), H/D ratio 0.71- 
0.8S (mean 0.78). Protoconch of 1 1/2-1 5/8 
whorls, mean diameter 875. 0fxm at 1 1/2 whorls. 
Apical sculpture of irregular pits and vague 
radial rugosities (Fig. 23e). Postnuclear sculp- 
ture of low spiral thickenings crossed by 
numerous low protractively sinuated radial ribs 
in a reticulate pattern (Fig. 231"), becoming less 
pronounced after the first four whorls (Fig. 23a). 
Sculpture continued on the base but reduced 
(Fig. 23b). No additional microsculpture. Um- 
bilicus small, diameter 0.62-1.54 mm (mean 
0.89) D U ratio 6.55-1 1.28 (mean 9.0 1 ), partial- 
ly obscured by the reflected columella. Sutures 
lightly impressed. Whorls rounded above and 
below a strongly angulate to thrcad-carinate 
periphery. Aperture ovately lunate. Lip simple, 
columella dilated and reflected over the um- 
bilicus. Parietal callus well developed. 
Periostracum strongly deciduous. Colour (with 
periostracum ) creamy-yellow; (without) 
whitish-pink, often with a darker brown to pink 
apex and spire. Based on 20 measured adults. 

Genitalia with epiphallus (Fig. 24b) rcflexing 
before entering the penis apically through a fleshy 
pilaster (Fig. 24c). Penial retractor muscle insert- 
ing adjacent to the penis.epiphallus junction. 

Penis (Fig. 24c) long and slender, muscular, with 
an apical bulb containing a grooved, muscular 
thickening. Penis proper with spongy lon- 
gitudinal pilasters. Free oviduct thick, mus- 
cularised, longer than vagina (Fig. 24a). Vagina 
short, internally with longitudinal pilasters. 

Radula (Fig. 23g, h) with tricuspid central and 
lateral teeth which have long lanceolate 
mesocones and well developed anterior flares. 

Based on 6 dissected specimens (QM- 
MO6068, FMNH206312). 

Range and Habitat 

H. delta inhabits cool subtropical notophyll 
vine forests of the Great Dividing Range from 
Dorrigo, NSW to the Bunya Mountains, SEQ; 
the humid warm notophyll vine forests of the 
coastal region of northern NSW; warm 
temperate rainforests in the Gosford and Blue 
Mountains area of central NSW; and some mar- 
ginally drier subtropical forest (microphvil vine 
forest, semi-evergreen vine thicket) in southern 
Queensland. The distribution of//, delta at the 
southern end of its range is not well known. 


H. delta has a caudal horn. This structure is 
typical of some New Zealand arboreal charopids 
and is common in Australian helicarionids. 

The shell of H. delta has an apical portion of 
about 4 whorls in which the sculptural features 
arc quite prominent, but on the remaining whorls 
sculpture is almost absent. A similar pattern is 
seen in//, ailaketoae. This is not gerontic growth 
and ils significance, if any. is unknown. 

Hedlevoconcha ailaketoae sp. nov. 

(Fig. 26a-e; Tables 2. 6) 


For Dr Aila Keto in recognition of her efforts 
for rainforest conservation. 


H. ailaketoae is distinguished from H. delta by 
its smaller size, fewer whorls, less elevated shell, 

weaker apical sculpture, and more prominent 
reticulate sculpture on postnuclear whorls (Fig. 
26a-d). The sympatric Oreokera cumulus 
(Odhner. 1^17) has an elevated, keeled shell but 
is larger, has fewer whorls, apical spiral cords. 

FIG. 26. a-e. Itcdlexoconcha ailaketoae sp.nov. a-c. Summit. Ml. Bellenden Ker. NEQ. QMM011254. 
holotype; d-e. Mt.' Bellenden Ker, NEQ. QMMO15X02. QMM01I273. paratypes. a-C, whole shell: d. 
apical-postapieal sculpture; e, central and lateral teeth, f-h, Hedlevoconcha addiia Iredale. 1944, AMC634S8. 





2 mm 





FIG. 27. Hedleyoconcha ailaketoae sp.nov. Mt. Bellenden Ker, NEQ. QMMO15802, paratype. a, genitalia; b, 
details of penis interior; c, talon; d, ovotestis; e, hermaphroditic duct; f, pallial cavity. 



< ■ 

u b i nda 

Sj RSfttt. > 500m 

^MW^I >.Vs" 

■ ' ■■ ■ 


» 30' f 

riG. 28. Distribution vWcdkyoeonchaatlakrtU'!* 
no v. 

and postnuclear sculpture of prominent radial 
ribs with reticulate mierosculpturc (Stanisic. 
I9g7,pls 1.2). 

Type Matekiai 

Hoi otvpf: QMMOJ1254, Ml Bellenden Kei. NPQ. 
Summit. 1560 m. SMVFT, Collected Earthwatch/- 
QM. l 7 -24 Oct 1981. Height of she!! 5.13 mm. 

Ii icier 6.81 mm, H/D ratio 0.75. D/U ratio ^ 4^. 
whorls 5 I/S+. 

ParaTVPFS: Ml Bellenden Ker. NEO- Summit T"V 
Mn. 1560 m UTtG'S, 1 45*5rE) rainforest (2, QM* 
MO 15802, 29 Apr 1**83; G.B Mnmeith. D. Yeales): 
Mi Bellenden Ker. NEO. summit. 1560 m. SMVFT 
(I.QMMO) 1273, 1-7 Nov 1981, Eanhuaich/QM). 

t tally covered by columella. Sutures lightly im- 
pressed. Whorls rounded above and below a 
thread-carinate periphery. Aperture ovate ly 
lunate. Lip simple, columella dilated and 
reflected overthe umbilicus. Parietal callus well- 
developed. Periostracum deciduous. Colour 
(with periostracum) straw yellow, spire darker. 
Based on 2 measured adults. 

Epiphallus (Fig. 27a) long, entering penis api- 
cally through a fleshy circular pad. Penis (Fig. 
27b) shorter than epiphallus, slender with cor- 
rugatcd. longitudinal pilasters I Fig. 27b), apical- 
ly with a penial bulb containing entrance of 
epiphallus (Fig. 27b) and a muscular grooved 
collar. Penial retractor muscle inserted at the 
penial bulb/epiphallus junction. Free oviduct 
museularised, much longer than vagina (Fig. 
27a). Vagina short with longitudinal pilasters 

Radula (Fig. 23c) with tricuspid central and 
lateral teeth which have long mesocones and 
strongly developed anterior flares. No data avail- 
able on marginal teeth. 

Based on 1 dissected adult (QMMO15802). 


//. ailakeioae has only been found to dale in 
simple microphyll vine fern thickets and forests 
on the summit of Ml. Bellenden Ker. NEO. Its 
microhabitat is unknown. 


Discovery of H. ailakeioae on Ml. Bellenden 
Ker was unusual considering that its ne<. 
relative is almost 1400 km to the south. Pre- 
vioush Slanisic (19S7) reported on the al- 
liludinalK restricted eharopids from this reg i 

DlACC- ! 

Shell trochoid (Fig. 2oc) 7 diameter 5 .97-6.81 
mm (mean 6.39 mm) with 5 f S normally coiled 
whorls. Apex and spire stronglv elevated, 
SPBWW ratio 0.54-0.63 (mean 0.61). heigh; 
4.62-6. 13 mm (mean 4.88 mm) H/D ratio 0.75- 
0.77 (mean 0.76). Protoconch 11.2 normally 
coiled whorls, mean diameter %5|xm at 1 12 
whorls. Apical sculpture of weak irregular pits 
(Fig. 26d). Postnuelear sculpture of prominent, 
spiral cords crossed by crowded, productively 
sinuated radial ribs in a strongly reticulate par- 
tern which becomes less pronounced after c. 4 
whorls (Fig 26b). Sculpture continuous On 
base but somewhat reduced. No additional 
microsculplure. Umbilicus verv small, diameter 
ii 72 mm, D U ratio 8.3Q-9-47 (mean N.S9), par- 

Hedlevoconchu addita Ircdalc, 1944 

'(Fig. 26f-h; Tables 2 6) 

th-tilcxnconcita addita Iredale. 1944. p |9. 
fig. J9 


In general appearance. H. addita bears some 
resemblance to ff. delta* The two type specimens 
are worn and only a few traces of vaguely reticu- 
late adult sculpture are visible. However, reticu- 
late postnuelear sculpture is not sufficient to 
conclusively link this species with the two main- 
land congeners. The keel of//- addita is much 
mote pronounced than in either H, delta or //. 
ailnketoae and may indicate that the overall 
similarity affll addita La these species isconvei- 


gent, but in the aosence of other evidence, the 

original allocation of Ircdalc f 1944) is retained. 

Tvpf Matfrtai 

I.flTOTYPE: AMC634S8. Ml Gower, SSE. Lord 
Howe Island, 
PARALECTOTVPE:, AMCI501N3, same COllCCliOn data 

as leclotype. 


Shell moderately large, trochoid (Fig, ZGg), 
diameter 6 89-6, MS mm (mean 6,94 mm) with 6 
to 6 t/8 (mean 6+) normally coiled whorls, the 
last whor! descending slightly. Apex and spire 
strongly elevated. SP/BWW ratio 0.63-0.67 
(mean D.65), height 5.3^-5 .46 mm (mean 5.42 
mm). H/D raiio 0.77-0.79 (mean 0.7N). 
Protoconch of 1 3/4 whorls, mean diameter 
S;^2.5[.im al I 1/2 whorls, Apical sculpture worn 
on mailable specimens. Post nuclear sculpture 
also mostly worn. Body whorl with remnant 
patches of periostracum and sculpture consisting 
of low crowded spiral cords and weak protec- 
tively sinuatcd radial growth ridges. Sculpture 
continuous on the base as crowded spiral cords 
and weak protractively sinuated radial growth 
ridges (Fig. 26g). No additional microsculpture 
visible. Umbilicus narrow. V-shaped, only 
slightly Covered bv the columella (Fig. 26h), 
diameter 1.48-1.54 mm (mean LSI mm). D U 
ratio 4.53-4.66 (mean 4,60). Sutures only slight- 
ly impressed. Whorls rounded above and below 
a strongly earinate periphery. Aperture quadrate 
to roundly lunate. Lip simple, columella dilated 
and partially reflected over the umbilicus. Parie- 
tal callus developed. Periostracum deciduous. 
Colour (with periostTucum) straw yellow, 
(without pcnosiraenm) chalky whitish-pink, 
w ilh a darker spire. Based on 2 measured adults. 

Anatomv unknown. 

Lenwebbia gen. now 

i i \ SlOl t)OY 

I nr Mr I m Webb in recognition ol hispJOl I I 
irii UUiiks Oti rainforest evolution. 

I | Si MIIS 

/ (.'/mi rhhtii protowroht tit lata sp 


Shell small moderately elc* mi i (j ig. 29c) 

average diamctet 1,3$ mm and relatively few 
(average 4 f/84) rounded whorls. Protoconch 

sculpture ^\' numerous dimples and vague radial 
ridges (Fig. 29g). Adult sculpture of spiral 
grooves, broad radial undulations and very fine 
thread-like radial ribs (Fig. 29e,f). Base with 
incised spiral lines and weak, raised radial 
growth lines. Sutures impressed. Umbilicus 
small (Fig. 29b). Lip simple, columella dilated 
and deflected toward the umbilicus 

Foot and tail broad, bluntly tapering posterior- 
ly. No caudal horn or foss. Body colour (in 
preservative) grey. Ommatophores black- Pallia I 
roof with some black speckling. Kidney orange, 
triangular, c. 12 length of the pallial cavity. 
Rectal kidney lobe reduced. Ureter sig- 
murethrous with an incomplete secondary arm 
and reflexed primary arm (Fig. 30e). Ureteric 
opening located at the rear of the pallial cavity 
in the angle between the kidney and the hindgut. 
G\ otcstis ( Fig 30c) several clumps of palmafcly 
elavatc lobes of alveoli, oriented at right angles 
10 'he plane of coiling. Terminal male genitalia 
with short muscular epiphallus (Pig. 30b) enter- 
ing penis apically through a simple pore (Fig. 
30b). Penis elongate, internally with few, large 
longitudinal pilasters (Fig. 30b). Free oviduct 
(I ig. 30a) longer than vagina, otherwise typical. 
Radula (Fig. 29d) with mesocone of central and 
lateral teeth long, lanceolate. Marginal leeih 


Icnu-fbhia differs from \gairca in its smaller 
size, mure weakly sculptured shell and more 
strongly pitted apex (Fig, 29g). Analumically, 
the relieved primary ureter (Fig. 30e) and bright- 
ly coloured kidney ol r f.cnwchhhi are dislinctb B. 
In the VludloGap area. SFQ. Lcmxcbhui is sym- 
patic wlthC OetiOCharOpavihG&t worn she! Is are 
grossly similar. However Cocnocharopa has an 
ape\ vs ifh curved radials anil raised spiral cords 
(Fig* 1 I li- M4f. 120r-h).._tdult sculpture of fine 
radiah- and spiral cords (Figs illg. 1 14g, I20d). 
ami more shouldered whorls. The syfllpatl 
helicarionid Turocystis Ircdalc. [937 ( has a 
protoconch with incised spiral lines and adult 
• Liilpturc of incised spiral grooves which have 
KB il.n noiefies along their length. 

Lenwebbia prntivscrobiuilal m now 
(Figs 29-31; rabies T . \ 


I aim tcrbbicultfc a little ditch; for the 
p nclatc protoconch. 

* § 

* 5 

5 2 
^ < 


z ~ 





£, protoscrobiculata most closely resembles 

Ngairea canaliculata which is sympatric with 

the former in the northern parts of its range. N. 

canaliculate lacks the punctate protoconch of L. 

;k \% protoscrobiculata, has prominent post nuclear 

S e s a radial ribs (Fig. 12a), channelled sutures (Fig. 

12e) and flammulated shell. In the southern part 
of its range L. protoscrobiculata is sympatric 
with Coenocharopa macromphala and C. par- 
vicostatu and may be confused with worn dead 
shells of these latter two species. However 
Coenocharopa has apical sculpture of curved 
radial ribs and low spiral cords and is smaller 
(Figs 1 14e-g; 120d-h). Anatomically a complete 
secondary ureter (Figs 1 15a, 12 la) is sufficient 
to distinguish these species from L. 


- 13 

- - - 


" ^ 

*s 3 §5 S S a ;i Type Material 

HOLOTYPE; QMMO 17048. c. 8 km from Kalpowar 

; on Fireclay Rd, Kalpowar S.F.. SEQ (24 E 43'S. 

15 1 "2 1 *E i Leaf litter, MVFMjwikw/a Collected by J. 
Slanisic. A. Green. 15 July 1980. Height of shell 2.94 
mm. diameter 4.4^mm, H/D raiioO. 63. D'U ratio5J1, 
whorls 4 3 s. 

QMMOI7047. same data as holotvpe. 36 specimens; 
Mt Mudlo. Kilkivan S.F. (26°01"S, I52°13 , E) leaf 
litter. NAT Araucaria (34. QMMOS359. 17 Jul 1980, 
J. Slanisic. A.Green); Mudlo Gap, Kilkivan S.F., litter 
MVF/Arauearia (3, OMMOI6S43. 5 Jul 1^84, J. 
Slanisic, D. Potter. K.Emberton): Mudlo Gap S.F. 
(26'01'S, 152M4 E) MVF /Araucaria (7. QM- 
MOI2V.5. \MCl3665, 2 Sept |982, AM/QM- 
ABRS): Clifton Ranee S.F.. SW of Fairlies Knob, 
Araucaria/fAVf (26*34'S, 1521 fr'E) (2. QMMO- 
12386, AMCI366S1. 2 Sept 1982, AM/QM-ABRS); 
Dawes Range. MYF, Araucaria (24 C 28'S, 1 31 WE) 
(2,OMMni2h52.4 Sepl 1982, AM QM-ABRS); c.4 
km below summit of Mt Fort William on Ml Fori 
William Rd. Kalpowar S.F.. liner, under rocks, logs, 
MV¥tAraucaria (7, QMMOI70I0. QMM016839, 3 
Jul 1984. J. Slanisic. D. Potter, K. Emberton); rain- 
forest walk, Kalpowar S.F. (24^4 i'S. I51*21'E) 
MVF/Araucaria (1. QMMO 12578. 4 Sept I9S2, 
AM/OM-ABRS); base of Mt Woowoonga. (25°26*S, 

|| ! c :(ihT| MVF/Araucaria (5. QM MO 12447, 

- = - - - = AMC136720. 3 Sept 1982. QM, QM-ABRS); Mt 

* ? Goonaneman, viaChilders, rainforest. 670 m, litter (2, 

0MMO17258. 3-7 Nov MW). R. Raven, V. Davies); 

ijj B |g s 5 Ml Goonaneman. rainforest (2, OMMO 17259. 6 Feb 

'■zll |: h f= ±1 1981, Q3. Monteiib); Mt Colosseum. 6 mis S of 

|| i\ || 31 MiriamVale(l2.AMC14292a.6lun 1964, L. Price); 

= ' * 5 -~ *° KalpOWST S.k, NE Of Monlo. SEQ (24 : 42'S, 




je — 

2 = 
< £ 






= 1 

=1 ; 

* f" 


~ _ 

~ * 

= rH 

t 7 

= = 






c * 











#**v *.-* 







K H 


FIG. 30. Lenwebbia protoscrobiculata sp. nov, c. 4km below summit of Mt. Fort William on Ml. Fort William 
Rd, Kalpowar S.F., SEQ. QMMO17010, paratype. a, genitalia: b. details of penis interior: c, structure of 
ovotestis; d. details of hermaphroditic duct and talon: e. pallial cavity. Scale lines as marked. 

FIG. 29. Lenwebbia protoscrobiculata sp.nov. a-c, c. 8km from Kalpowar on Fireclay Rd, Kalpowar S.F., SEQ. 
QMMO17048, holoiype; d, Summit of Mt Fort William. Kalpowar S.F., SEQ. QMMO 17010, paratype; e-g, 
Mt Mudlo, Kilkivan S.F., SEQ. QMM08359, paratype. a"-c. entire shell: d. lateral and marginal teeth: e, early 
post nuclear sculpture; f, late postnuclear sculpture: g, apical sculpture. Scale lines as marked. 




25 D S 


27 Q S 



153 D i 

FIG. 3 1 . Dislribulion of Lenwebbia protoscrobiculata 
sp. no\. 

15 r:rE) vine thicket scrub (S.QMMO 13 U<\ \ Sepi 
1982. AM QM-ABRS); Camp Creek. SW side ol 
Bobbv Range. S of Miriam Vale. SEQ (24!36'20"S. 
!5r33'E)U.QMM0132N7, 6 Sept |982i AM GM- 
ABRS): Limestone Ck. W. of Childers, SEQ 
C5M5 , 30"S. IsrSS'EJ (2, (JMMOJ30W. 3 Sept 

OTHEB M \tfri\i 


(24^32*5, i5i ■ : ainfo«5t{I,QMMO|7260, S 

Mar 1977, QM party); summit Ml Buoroon BoOtOOn, 
SW nt Mnium Vale. \'\\'F Araucatia, litter (?, 
OMMOIhShl. 17 Sept 1985. J. Sumsic D. Potter); 

Daw« Rangfc MVFMrriMctfrfA [24*28'S, isro7'E) 

(I, AMCl ^6781-4 Sept 1982. AM/QM-ABRS). 


Shell small, depressed, diameter 3.96-4.69 
mm (mean 4.35 mm) with 3 7/8 to 4 3 8 (mean 
4 i 8+) normally coiled whorls. Last whorl des- 
cending Ortly very slightly. Apex and spire 
elevated (Fig. 29c), SP/BWW ratio 0.12-0 2 S 
(mean 0.19) height 2*56-3.15 mm (mean 2.79 
mm). H/D ratio 0.61-0.71 (mean 0.64). Apical 
sculpture highly modified, consisting of 
numerous, crowded dimples with a few, vague. 
radial mainly at the terminal portion of the 
protoconch (Fig. 29g). Protoconch of 1 1/2 to 1 

12+ whorls, mean diameter 9Uo.9u,m at J 12 
whorls. Post nuclear sculpture of spiral grooves, 
closely spaced, occasionally interrupted and 
crossed by weakly protractively sinuated radial 
ridges (Fig. 29e). Secondary sculpture of fine, 
crowded, radial growth ridges particularly on the 
body whorl (Fig. 29ft. Umbilicus small, barely 
covered by the reflection of the columella, 
diameter 0.62-0.82 mm (mean 0.75 mm), D/U 
ratio 5.34-6.52 (mean 5.85 mm). Sutures im- 
pressed. Whorls rounded. Aperture roundly 
lunate. Lip simple, columella dilated, slightly 
thickened and deflected toward the umbilicus. 
Parietal callus not strongly developed- Base 
gloSSV, munded. with a sculpture o\^ incised 
Spirtl striae and weak radial growth lines- Colour 
yellow -brown with a darker, grey spire in some 
specimens. Based on 24 measured adults. 

Genitalia with epiphallus entering the penis 
through a simple pore (Fig. 30b). internal I v w nh 
longitudinal pilasters. Pen is a muscular tube with 
spongy longitudinal pilasters (Fig. 3(Jb). Penial 
retractor muscle inserting at the penis epiphallus 
junction. Free oviduct (Fig. 30u) a long, thick- 
ened muscular tube, internally with longitudinal 

Radula (Fig. 29d) with tricuspid central and 
lateral teeth in which the mesocone is long and 

Based on 3 dissected adults (QMM017U1U 
AMC 142928). 

Rasgeand Habitat 

L. protoscrobiculata inhabits the drier sub- 
tropical microphyll vine forests [whhAraucarta 
emergents) between Mudlo Gap and Bobby 
Range, SEQ, and rarely, moister subtropp a! 
notophyll vine forests where these are in jux- 
taposition with drier rainforest. Little is known 
about the microhabitat of L protoscrobiculata 
because most available material consists of dead 
shells collected from litter. However, some live 
specimens have been taken from the upper sur- 
face of rotting logs. The comparatively reduced 
sculpture of L. protoscrobiculata is not indica- 
tive of life under logs but rather suggests that the 
species lives in curled lea\es. or under bark of 


At first glance L. protoscrobiculata appears 
more like a helicarionid than a charopid. 
Reduced radial sculpture (Fig. 29a), shell colour, 
and coiling pattern are not typically charopid. 
However the peculiarapical sculpture (Fig. 29g). 


i ff iculatc adult microscuipture (Tie. 29f) and 
anatomy (Fig. 30), leave no doubts about its true 

The incomplete secondary ureter (Fig. 3<ie) is 
inconsistent with living in drierhabitats but in the 
case of L- protoscrohtculata the primary ureter is 
lengthened b) re 1 1 e ■'• i<W at the kidney apex. 

/. . protoserohiculuiti a| fpc ars to be a derivative 
of the Ngairea group. The distinctive punctate 
apex, incised spiral grooves an the post nucicai 
Shell and coiling pattern oi reduced number of 
rounded whorls can all be readily derived from 
Ngairea, Anatomically, the differences from 
Ngairea are not great. 

Setomedea Iredalc. 1941 

. nu-a'at IreUate. t933, p. 53 nom. nud., trcdale, 
1937a, p, v<*. num. nudL; Iredalc t94la, p. 267; 
Iredule. L94tb, fl 1: Hurch. 1976, p. 133 

1 H't/fa (t I t ! - ersba* 19561) p 7, 

I'vpf Spfcifs 

Staerta seiicosnua Hod lev, l l) 24; by original 

!. ignation. 

PftFvmcsSit D|] ■ 

Iredale (1933) introduced Scionwdca for 
Suteria setkostata Hedlcy. 1924 without anj 
accompanying description. Subsequently 
lie (1941a, p. 267) provided a barely mini- 
mal description and I consider that the name 
should dale from Ibat publication. FrC* 
(1937a, p. 329) included 'Endodonta' aculeata 
licdley. IS99 from Wollongongi NSW without 
giving reasons- Sciomcdca, herein, include 

o.siata. three new species from Queensland. 
and a possible new species hum the Atlu 
Tableland. 'Endodontic aculcata Medley, 1 89*3 
is excluded to be dealt with later. 


Small to medium sized Charopidae. adult shell 
diameter 256-5.7] mm. with 3 5/8 to S 5/8+ 

normalK in loosclv [nudicostata] coiled wiiotIs. 
Height o\' shell 1.36-3.37 mm. Apex and spire 
^lightly (seiicoshihi) to strongls [inoiticiiin) 

elevated- Body whorl descending In front, 

t'rotoconch ^hiny. rounded, with I 3/8 to 1 
whorls. Apical sculpture of numerous slighllj 
[seticostQJQ* janac) to strongly (nudicosnon 
motilpitfri) squiggly spiral cords with a few 
ue radial undulalions mainly near the eml 0l 
the protoeonch. Posinueleai sculpture complex. 
of broad spiral furrows (except in nudicostata) 

and numerous, moderateh spaced, COiilp 
radial rihs farmed by the enlargement of - 
radial riblets. The major ribs have periostracal 
blades scalloped (caused hy rhe spiral undula- 
tions in the shell surface) or normal (nudicos 
laut). with or without [nudieosutta] regul 
spaced slender setae along their length. Mice 
culptiire Df fine, crowded radial riblets. 7-15 
between each pair of major ribs, and numerous, 
crowded spiral cords on the radial ribs. Sculpture 
COniinVOUb or fhe base. Umbilicus wide U 
shaped to narrow V-shaped (nudicostata). Su- 
tures impressed. Whorls shouldered above and 
rounded below a slightly compressed periphery. 
or rounded abQVC and flattened at the basal mat 
gins (nudicos(ata). Aperture roundly lunate. Lip 
simple, columella Slig)Ul) thickened, Panefa! 
callus developed Coloui yellow-horn to 
brownv-vellow with reddish-brown flammula 
tions and" suffusions. 

Foot short and bioad. bluntly tapeted 
posteriorly. No caudal horn or foss developed. 
Coloui ol animal fm preservative) creamy; dark 

in nudicostata, Ommatophores i in preservative) 
black (fietit OStaMt, nudicostata) or white {jco. 
motvcilhi) Kidney broadly triangular to elon- 
gate. Ureter sigmurethrous. Secondary ureter in- 
complete with ureteric opening ai rear of the 

paliiaf cavitj j ru to the hindgut Initial p 
of the primarj ureter rellexed and coiled about 

l.idnev apev CUotcstis two clumps of pal 
matel} davalc lobes of alveoli located in the 
apical w horls of the digestive gland or above the 
stomach {nudicastata}^ oriented parallel to the 
plane ol coiling. 

Terminal map genitalia with a muscularised 
cpiphallus which ma\ bfi short (mtmteil 

vraielv Iqng, very Lang {nudtcostata)^ or 
modified and partial 1 11 | rated into the 

penial apex Epiphaltic pore simple, 

entering bulb ol penis apieally or subapicallv 
[seticostafu, montcithi). or entering di recti \ i 
lower penial chamber {janac). Penial retractoi 
muscle inserting on the penial apex {montcitht), 
on iiie cpiphallus near the penial bulb, or on the 
epip&allus well removed from the pem 
{nuditostata). Penis with an apical bulb 
separated bom the main penial chamber by a 
muscular collar, or without a muscular collar 
[ntuUcOXl(Ha)< Apical bulb with pilasters (mOn 
let flu. WliCQSt&ta), verge (nudicoMata) or with 
incorporated cpiphallus {janac). Penis proper 
with Mes|.\ longitudinal pilasters or low lOfl 
gitudinal pustular thickenings {nudicostata). 

Raduii with iricuspid central tooth that li 




ri^' -c — ~ ^. 

— x 

jo g - 



> z 

u. < 

C z 

H 5 

5 3j«2 

2 s 

3 ? ^~ 

-Si. -3 i 

--a ~ c 

ft II 

slender, lanceolate mesocone and smaller 
pointed ectocones. Otherwise typical. 

Distribution and Ecology 

Setomedea has a disjunct distribution from 
Dorrigo, NSW, to Shiptons Flat, near Cooktown, 
north Queensland. It inhabits moist temperate 
and subtropical rainforests of northern NSW and 
southern Queensland, subtropical forests of the 
Mackay/Prosperine area, and humid tropical 
forests of north Queensland between Tully and 

Setomedea lives under and in logs and oc- 
casionally may be found under bark with 
Ngairea. Species show a preference for very 
moist habitat where decomposition is well ad- 
vanced. Large numbers of individuals can be 
found under the same log or piece of bark. 

Patterns of Shell Variation 
The umbilicus is typically wide U-shaped. 

Departures from this pattern are seen in S. 
I s ^ % nudicostata (Fig. 35b) from SEQ and Setomedea 

sp. (Fig. 44b) from the Atherton Tableland, north 

Variation in postnuclear sculptureis small. 

The most dramatic change is the loss of perio- 

stracal setae and spiral furrows in Setomedea 

nudicostata (Fig. 35f), 
Pcriostracal setae are rare among the 

Charopidae and only a small number of 

Australian and New Zealand taxa posses them. 

On their ow r n they do not indicate phylogenetic 

relationship. However in the case of Setomedea, 
M _ anatomical patterns unite the group, and indicate 

1 - I S |5 5 that this character was derived from a common 
| e, $i S; :-2^ ancestor. 

Patterns of Anatomical Variation 
£-5 I £ ? The incomplete secondary ureter of 

l-E sjg %i s2M Setomedea is similar to that found in Oreokera, 

Medley oconcha, Mussonula, Lenwebbia and 
Ngairea. However, the apical region of the kid- 

2 ? | | | ney has the pericardial lobe well developed (Fig. 
i ,1*1 i«U! 36e). The initial section of the ureter is reflexed 
* ^s 55, ǣǣ and coiled around the tip of the kidney giving the 

kidney apex an S-shaped appearance. While 
some genera (e.g. Lenwebbia), may have the 
initial part of the ureter reflexed, only Setomedea 
has this unusual coiling. 

The terminal male genitalia of Setomedea 
show variations in the internal structures of the 
penis, shape and size of the epiphallus, and the 
insertion of the penial retractor muscle. Typical- 
ly the epiphallus is a short, reflexed, muscular 



tube entering the penis either apically or subapi- 
catly (Figs 42a. 33b). In 5. janae pari of the 
epi phallus is incorporated into the penia! com- 
plex (Fig. 39c) and may represent spatial adjust- 
ment in response to reduced whorl count. 

The apical and female genitalia of Setomedea 
arc without unusual features except in S. 
nudicostata which has the free oviduct very long 
and the ovotestis situated in the digestive gland 
just above the level of the stomach. The shift in 
position of the ovotestis from apicad in the diges- 
tive gland is probably related to reduced whorl 


Setomedea can be distinguished from other 
subtropical and tropical charopids with spirally 
Urate protoconchs by size, shell colour, and adult 
sculpture - Oreokera has a much larger trochoid 
shell (Stanisic, 1987, pis 1,2); Sinployea has a 
smaller, monochrome brown shell with more 
numerous, simple, radial ribs (Fig. 137a, e); and 
Omphaloropa is smaller in size, has broader 
apical cords, wide cup-shaped umbilicus and 
few. widely separate major ribs on the body 
whorl (Fig. 134a,b,e). 


1 .Shell with periostracal setae 2 

Shell without periostracal setae (Fig. 35a. f) 

J. Shell wiih baser coiling (Fig, 44a); fewer 
whorls (Fig- 44a); mean whorl count 3 7/8+ 
(Table S) Setomedea sp. 

Shell wiih tighter coiling; many whorls; 

mean whorl count greater than 4 1/2 3 

3 epical sculpture of regular spiral cords 4 

Apical sculpture of squiggly Spiral PC 
(Fij£.4lef montehhi 

A Major Tibs with simple periostracal blades (Fig. 
320 » seticostata 

Major rib& With scalloped periostracal blades 
(Tig. Rjjlg) janae 

Setomedea seticostata (Hedlev, 1924) 
(Figs 32-34; Tables 8. 9) 

Stturui SCTfCQStGM Hedlev. 1924, p. 221. pi. 32. figs. 

Setomedea seticostata (Hedlev): Iredale. 1937a, p. 

32ft Iredale. 1941 a. p. 268,'fig, 6; Iredale, 1941b 

P. I. 
SeiomeJia fsic) seticostata (Hedlev); Kershaw, 



5. seiieostaia is characterised by a flammu- 
iated shell with spirally corded protoconch and 
adult sculpture of complex radial ribs, 
microspiral cords, micruradials, spiral grooves 
and prominent periostracal setae. S. janae has 
fewer whorls with strongly scalloped periostra- 
cal blades and more prominent radial grooves 
(Fig, 38a. e-g); 5. montelthi is larger with 
squiggly apical cords and a more protruded spire 
(Fig, 41c. e). Anatomically the incomplete 
secondary ureter and coiled, reflexed primary 
ureter immediately distinguish S. seticostata 
from other sympatric charopids reviewed in this 

Previous Studies 

Hedley (1924) tentatively assigned this 
species to the New ZealandSwtena Pilsbry, 1892 
on the basis of overall similarity to Suteria ide 
(Gray, 1850) which also has periostracal setae. 
However, different apical sculpture of 5. ide 
indicates that setal processes in the two species 
are convergent. 

Type Matfriai 

HOLOTVPH: AMC6351 I. Dorrigo, NSW, under logs. 
Collected b> S.W.Jackson. Nov. 1910. Heightof shell 
1.87 mm. diameter 3. 58 mm, H/D ratio 0.52. D/U ratio 
2.90, whorls 4 5/8+. 

PARATYres: AM036S5, ? specimens, same locvin . 
data as holotype. 

Other Material 

NEW SOUTH WALES: Clarence River (>. 
AMC 142467. Cox colUiTooloom Scrub. Beaurv S F. 
CS^rS. 152 t 24"t)- NYF (<. QMMOIIN37. 
AMCt28533 > 15 Mar 1981, AM/QM - ABRS); 
Woodenbone Ck. S slope of Mt Clunie (2B*2D V S, 
152 t 30'F). rainforest (15, QMMO10979. 
AMCI2S-S5, 15 Mar 1981, AM/QM - ABRSV W 
slope Wilsons Peak, top pf Condamine River valley 
(28°t6'S, l52"28 , E)(2,AMCt28635, 16 Mar 1981, 
AM/QM - ABRS): N of Dorrigo. c. 1.5 km along 








dK p RM 


FIG.33. Setomcdea seticostata { Hedlev. 1 924). a. b. e, g. c. 1 7km from Tannv morel, on T;tnn\ mord-Cambubal 
S.F. Rd. SEQ. QMMO12705; c, Cunninghams Gap^EC. 0MMO12678' d. f, c. 1km Eof Carrs Lookout. 
Koreelah S.F.. SEO- QMMOI6477. l( . genitalia: b T details of penis epiphallus junction: c. details of penis 
inlerior: d. hermaphroditic duct: e, ovotestis; 1". talon; g. palhal eavity. 

FIG. 32. Setomedva sef/cortata (Hedley, \ l )24). a-c, Dorrieo. NSW UVIC635 1 1 . holotvpe; d. Acacia Plateau, 
Koreelah S f .- NSW. QMMO10952; e-t'. Ml Glorious. SEQ. GMM014164: c. 1km E of Can's Lookout, 
Koreelah S.F.. SEQ. QMM016477. a-c. entire shell; d. whole shell show ing setae: e. apical sculpture: f. post 
nuclear sculpture: g. central and lateral teeth: h. marginal teeth. Scale lines as marked. 





26 3 S 





152 a E 

153 D E 

154 °E 

FIG, "U. Distribution o\' Setowt'tJrn Wt'tCOVata ( Med- 
ley, 1^4) 

Kelly S Road (3CTtfri 152 4] F). lainloreM (0. 
AMC12832Q, 1 J Mar 196-1, AM/QM- ABRS); SEof 

rflbuton, tributai j of Busbj 9 C k 1 20 02 S 

I5:"43L) (12. QMMGI2944, AMC137759, - y Aug 
1982, AM/OM - ABRS): Ml Warning (28°24*5 
l5.ri7M. ratoforesi (22. QMMGitW99. AM- 
C1293U2, IV Mar 1981. AM/QM - ABRS);Toonum- 

t>ar S.t .. C. 34 km Irom Kvoglc on Toonumbar loresi 
road (2iT3J"S, !52"45"i-) (3 QMMO1090! 
AMC L2S40S. M Mar 1981, AM/OM -ABRS). Wian- 
gflrie S.f ■ c 25 1cm NE of Kvogle (28 U 2VS 
iio'L), 1000 01 (7, AMC1424K1/17 May 1976. 
PH. Coin, .in I i oi ni: Red Scrub F.R.. Whian Whian 
S.F {2J&°38*S, I5319'E), 210m(l,AMCI424X4, |j 

Mav 1976, P.H. Caiman, t. Loch); Dorrtgo (23. 

AMC63760« SAW Jackson): Koreelah S,F„ Acacia 
Plateau. (28 C 2TS, 152 : 24"E). SFVTMVF (47, 
AMC128554,QMMOI0952, 15 Mar I981.AM/UM 
- ABRS), Beaury S.F., C. 15 km W of Urbenvdle 
u 21'S, I52 L "24'F) .810 m (2, AMC 1424X3, !XM.i\ 
1976. P.H. Colman, I. iochi; Rocky Ck, Red Scrub 

F.R.. (28 '38'S. 153*20 E), rainforest (9, 
OM MO 10404. AMC I 28865. 20 MflZ 1 9$ I . AM/QM 
-ABRSjiTerani* Ck{28 P 34'S < L53*)9"£),340m(li 

AMCI424X2, 16 May 1975, P.H. Colman. L Loch). 
Bullina (2. AMC 1 1 UTS, ex C, Hedley); Tweed River 
(4. AMC 142485. ex Cox. ex Pellerd); Murwillumbah. 
under togN(l,AMC2S488.3 Dec 1907. C.F. Laseron); 
Lismore (2, AMC5784, ex C Medley); Domgo (5. 
AMC63772. S.W. Jackson); Bryon Bay f3. AM- 
C4312, f\ Cox); Big Scrub, Murwillumbah (3, AM- 
C103795. Lower, ex Cox); Richmond River, Lismore. 
in scrub under decayed timber (I, AMC63N72. ex 
Brflzier); Sherrard Falls, Dorrigo. warm temperate 
rainlt)reM.lilier,t)()0m(l,QMMOI(i7 7 ?i. J Nov 1983, 
AM OM - ABRS): c. 1.5 km E along Kelly's Rd in 
dense lainloiesi. Cloud's Creek S.F.. (30'07'S, 
i 4 IT) (3 QMMOKI773, AMC128320, Ma. 
1981. AM/OM - ABRS); along Croftby Rd, lop of 
Condamine River Valley, (2S"I5'S, IS2*2$'E) (I. 
QMMOI0540. Mar 1981, AM/OM - ABRS); c. 5.5 
km E Of Dorrigo, |30 : 2-VS. !52*44'£) (4. 

QMMO10796, AMC128351, Mar 1981. AM/OM - 
ABRS I98i)$ Tooloom Scrub, Beaury S.F.. under 
logs, NYF (V QMMO 10900,4 May 1986. J. Stanisic. 
J. Ctuttellng); Ml Glennie 16 km E of Woodenbong, 
900 m (3, OM MO I 1 933, 24 Oct 1 981 G.B. Momeith, 
D. Yeates); c. I 5 km from summit of Mt Warning. 
(28*24'S, 153' ln , E)(3.QMMO10515.AMCI2931 Q , 
Mar 1981, AM/QM - ABRS): Mt Warning, CNVF 
24 5, 153'16 , E)(2.0MMO12737.AMC136831 5 

15 Dec 198 1, AM OM- ABRS); Swan's Rd, Bruxner 
Park Flora Resei ve. subtropical rafofOre&f, under logs 
[6, OMMOI4768, 14 Nov 19X3, AM/QM - ABRS). 

raJlb hack. Bitina Burra. Laming (tin N.P., under logs 
(3, QMMO 1655 1,25 SejH 198$, J. Stanisic) 1 Burleigh 
He. ids N P , under logs, vine thicket (2. QM- 
MO 16922. 21 Jan 1987, J. St.tnisic. D, Potter); c, 3 km 
SF ur'sumnulMl Glorious. NVF(27 U 2()'S. I5246'E) 
(3, QMMOU99D, QMMOI1994. AMC136419, 2 
D^c 19S2. AM/QM -ABRS): c, 25,2 km from Goom- 
burfa. Goomburra S.F.. N V 'ViPu\n\sfAntuctiria 
(27°59*5< 152 C 21'F)(5,OMMOI 1121 AMU28618, 

1 6 Mar 1 98 1 ; 7. QMMO I 269 1 . 7 Dec 1 9R I , AM/QM 
- ABRS); Curtis Fall; I irnni. Vii I Wfboi \tit t U " 
CNVF I 1, QMMO 16436, 23 Feb 1986, J, SlanMc I 
Cbasding); Mt Hobwcc Circuit. Binnu Burra., 
l.ammglonN.P . liiter NVF( 10, QMMO I SSON, I Oct 
10X5. L SlaniMc, J. Chaselmg, D. Puller); Tullawallal 
Circuit. Binna Burra. I.amingHin N.P.. under logs, 
N\ F (9. OMMOUw37. 2 Oci 1&SS, 1. Stanisic. D. 
Potter. J. Chaseling): Border BiOAfl Burra, 
Laminglon N.P., under logs, NVF (5, OMMOW.isT I 
in Mar tos4. J, Stanisic); Maiala N.P.. Mt Glorious, 
NVF(6,QMMOll603,QMMO!1S4X, 20 Jun l L .'S2. 

o-l dV 

| -'. - - <-. e * 9 


MSA parlv): Koreelah S.F.. Acacia Plateau, under 
logs, NVFMVF (I. QMM0167%, 4 May 1986, J. 
Sianisic. .1. Chaseling): Lower Ballanjui Falls Circuit. 
Binna Burra. Lamington N.P., NVF, litter (7, QM- 
M016S92, 30 Sepl 1985, J. Stanisic. D. Poller, J. 
Chaseling); O'Reillvs, Laminglon N.P.. (28 14 S. 
153'15'Fi MVT iT\ QMMOHI427. 17 Mar [981, 
AM/QM - ABRS); Ml Glorious, NVF, litter (3. QM- 
MOI4164. 28 Nov UJ83, J, Stanisic); Araucaria 
Track, Binna Burra, Lamington N.P., under logs, NVF 
(7.Q\1MOlh578. 27 Sept 1986, -L Stanisic); c. 1 km 

»eS xs E of" Carrs Lookout. Koreelah S.F., under logs, NVF 
(10,OMMOlM-7.?Ma> 1 986 J, SUni§ic; J. Chasel- 
ing); Border Track, Binna Burra, Laminglon N.P.. 

§g under logs. NVF. (9, QM MO 16877. J. Sianisic. 31 

il p Mar 1984); Hall's Plain, E of Emu Vale, under logs. 
CNVT, (4, 0MMO16S95. 4 May 19S6, J, Sianisic^ J. 

~_ Chaseling) Cunninghams Gap. (1, QMMO6070, 21 

Mar 1477. M.J. Bishop): Binna Burra. Lamington 

5*1 1 * 

5 -4 i o i 

A 3 3 


g N.P., CNVF, 85(1 m (2SMTS, I53M2'E) (L QM- 
MO6073. Apr 1 976. M.J. Bishop); The Knoll N.P., Ml 
Tamborine. palm gully (27*55'S, ISSMO'E) (LQM- 
MO6305. 3 Aug 1976. M.J. Bishop); initial part of 
Kueebam Caves Walk, Binna Burra, Lamington N,P. 
under logs. NVT Cv QMMOU.XM. y Mir 1984, I. 
Stanisic); Cunninghams Gap, near monument. 755 m, 
NVT (28'04'S, I52*24'E) (8. QMM012678, AM- 
C 136807. 7 Dec 1 98 L .L Sianisic, 2. AMC1 28605. I 6 
Mar 1981, J. Sianisic. D. Poller; 3. QMM0169U1, 5 
Ma\ 1^86. .1. Stanisic. J. Chascling); Upper Pine 
Creek, via Canungra, under bark of fallen tree, rain- 
forest wilh Araucaria (6, QM MO 16904, J. Sianisic, 
D. Potter, 21 Jan 1987)' Coomera Falls Track. Binna 
Burra Laminglon N.P.. NVF. under log (1. QM- 
MOIo5"5. 26 Sept 1986, J. Stanisic): Nagarigoon 
Falls Track. Binna Burra. Lamington N. P., under logs, 
NVT (1. OMMO 16883. 20 Apf I9S6. J. Sianisic J. 
Chaseling); c. 17 km from Tanny morel, on Tan- 
nymorel-Cambubal S.F. Rd. 1000 m. MVF, Araucaria 
(3. OMMOI2705. AMC1368M), 7 Dec 1981. 
AM/QM - ABRS IMS! >; Ml Glorious, in litter and 
under lugs, NVF (3. QMMO]M2X. 26 Jan L986, I. 
Sianisic J. Chascling); Natural Bridge N. P.. (28 D 13"&, 
I5314T). NVF (7, QMMO 10452. QMMO10461. 
AMC129265, |8 Mar 19NL AM QM -ABRS). 


Shell strongly depressed, diameter 3.41-4.47 
nlffl (mean 3.91 mm), with 4 5 S to 5 1 4 (mean 
5-) normally coiled whorls. Last whorl descend- 
ing slightly mote rapidly. Spire elevated, apex 
--! 5 5 S#f uf Is 5 Slightly sunken to elevated, SP.'BWW ratio 
c i l}I El I Ml is it n II J 4-0.24 (mean 0.20). height 1. 75-2.34 mm 
j|f||| i| § f|j| |1 || || || (mean 2.05 mm) , R D ratio 0.47-J1.59 (mean 
I* * B * J = ~ * * 0.52). Protoconch glossy, of 1 1/2 to 1 5/8 

z - 



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whorls, mean diameter 628. 6|xm at 1 1/2 whorls. 
Apical sculpture of continuous low, narrow 
spiral cords and a few curved radial growth 
ridges (Fig. 32e). Post nuclear sculpture of 
prominent radial ribs, regularly spaced, 61-94 
(mean 72.6) on body whorl (Fig. 32f). Ribs/mm 
1 . 1 5- 1 .54 (mean 1 .32). Major ribs slightly scal- 
loped, formed by two raised radial ribs the first 
of which has periostracal setae at regular inter- 
vals along its length (Fig. 32f). Microsculpture 
of fine radial riblets, 6-12 between each set of 
major ribs, whose interstices are 6-8 times their 
width, and fine, crowded spiral cords which are 
also visible on the apical surface of the radials 
(Fig. 32f). Shell surface with weak spiral fur- 
rows. Sculpture continuous on the base (Fig. 
32b). Umbilicus large, diameter 1.15-1.54 mm 
(mean 1.32 mm), D/U ratio 2.81-3.13 (mean 
2.98), U-shaped. Sutures impressed. Whorls 
shouldered above and rounded below a slightly 
compressed periphery. Aperture roundly lunate. 
Lip simple, columella slightly thickened. Parie- 
tal callus developed. Colour yellow-horn with 
zigzag reddish-brown flammulations which are 
continued onto the base. Based on 13 measured 

Genitalia with cpiphallus (Fig. 33a, b) short, 
muscular, reflexed before entering the penis sub- 
apically through a simple pore (Fig. 33c). Penis 
(Fig. 33c) pear-shaped with apex sheathed to 
terminal part of epiphallus. Penial caecum 
present. Apical bulb of penis containing a fleshy, 
horseshoe-shaped pilaster. Lower penial cham- 
ber separated by a muscular collar and contain- 
ing low fleshy pilasters (Fig. 33c). Penial 
retractor muscle inserting on the epiphallus. 

Central and lateral teeth (Fig. 32g.h) with long, 
lanceolate mesocone. No data on marginal teeth. 

Based on 8 dissected specimens (QM- 
M016477, QMM0127Q5, QMM012678. QM- 

Rangfand Habitat 

5. seticostata inhabits warm temperate forests 
and cool subtropical vine forests of the Great 
Dividing Range (from Dorrigo, NSW to the Mis- 
take Mountains, SEQ), and the highlands of the 
McPherson Ranges. The species also occurs in 
the coastal, moist warm subtropical vine forests, 
from the Bellinger River, NSW. to the 
Queensland border, and the unusual coastal vine 
thickets at Burleigh Heads, SEQ. The northern 
limit of distribution is the warm subtropical 
notophyll vine forest of Mt Glorious in the 

southern D'Aguilar Range, SEQ. Like Ngairea 
dorrigoensis, S. seticostata has been collected in 
drier subtropical forests (vine thickets and 
microphyll vine forest) in the northern part of the 
Richmond Range, NSW. It lives under and in 
moist rotting logs and forest debris. 


Early collections of S. seticostata- include 
many localities in and around the Big Scrub 
region of northern NSW. Today S. seticostata 
survives in remnant pockets of this once large 
rainforest such as the Red Scrub Flora Reserve, 
NSW (28°38 , S, 153°19'E). The microhabitat is 
the damp, decayed recesses of rainforest floor 
debris which are generally ignored by other 
charopid species. Periostracal setae may be an 
adaptation for surviving in these sticky, damp 
areas by minimising particle adherance. There is 
geographic variation in shell parameters but it is 
not possible to infer trends from the limited adult 
material available. Although 355 specimens 
were available for study only 13 were adult and 
suitable for statistical analysis. 

Setomedea nudicostata sp. nov. 
(Figs 35-37; Tables 8, 10) 


Latin mtdus, naked; and costata, having ribs; 
referring to the absence of periostracal setae on 
the major ribs. 


5. nudicostata lacks periostracal setae. It is 
further distinguished in the genus by low whorl 
count (3 7/8+), small size (diameter = 2.86 mm), 
narrow V-shaped umbilicus (diameter =0.86 
mm), flammulatcd shell, apical sculpture of low, 
squiggly spiral cords, and numerous curved 
complex radial ribs on the adult whorls. The 
sympatric Gyrocochlea curtisiana has a flam- 
mulated shell and adult sculpture of prominent 
curved radials (Fig. 54a) but differs in being 
larger (diameter = 5.78 mm), having an average 
4 5/8+ whorls which are strongly shouldered, a 
convex pitted spire, and very wide umbilicus 
(diameter = 1.86 mm). Sinployea intensa (Fig. 
137a, b), is smaller, has fewer but more regular 
apical spiral cords, more numerous, simple 
radial ribs on the adult w r horls. and a 
monochrome brown shell. Omphaloropa 
varicosa (Fig. 134e,f) has regular spirals on the 
protoconch but is much smaller with relatively 




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FTG. 36. Setomedea nudicostata sp. nov. a, c-e, Granite Creek, on Bobby Range Rd. 5.2 ml E of Bulburin 
Forestry Station, SEQ. AMC142934, paralypc; b, Mt. Booroon Booroon, Bobby Range. SEQ. QMM016593, 
paratype. a. genitalia; b. details of penis interior and penial retractor muscle insertion; c, talon and her- 
maphroditic duct; d, ovotestis; e. pallial cavity. Scale lines as marked. 

FTG. 35. Setomedea nudicostata sp.nov. a-c. Foothills of Bobby Range, Granite Ck, SEQ, QMM01516S. 
holotype; d. g-h. Granite Ck on Bobby Range Rd, SEQ. AMC 142934, paratype; e-f. Mt Booroon Booroon, 
Bobby Range, SEQ- OMMO 16593, paratype. a-c. entire shell; d, marginal teeth: e. apical sculpture; f. posl 
nuclear sculpture; g, central and lateral teeth; h, details of marginal teeth. Scale lines as marked. 




2<t r 35' 

■ I 

ratr.r« -,:■■■:■:; 

«U. > 400m 

*% ^&$ y,t ' ^'^v^l 

151 to-] 

FIG. 37. Distribution of Sctomcdca nudicusttih 
niiv, m the Bobby Range area. SEQ. 

large umbilicus, and adult sculpture ol Few very 
prominent radial ribs with interstices which arc 
strongly reticulate. 

type Material 

Hoi.otype: QMM015 168, foothills of Bobby 
Range, Granite Creek, SEQ (24*30. 7'S, 15130,3'E), 
under Leaf litter. Collected by Terry Carless, 4 Decem- 
ber 198?. Height Qf she!! I 3b mmajiameter 2,68 mm, 
H/D ratio 0.5L D,U ratio 1.5s. whorls 3 5/8. 

PaRaTYPES: Gfani'te Creek on Bobby Range Road, 
5.2 mi!es E ol Bulburin Forestry Stn, SEQ (2. 
AMG142934, 8 May 1975. J.B. Bureh): Ml Booroon 
Booroon, Bobby Range, SEQ. bast, NVFalongcreek, 
under logs (3. QMMQ 16593, 17 Sepi 1986* J. 
Stanisic. D. Potter); c. 17,9 km E of Builyan, on 
Builyan-Bulburin S.F. Rd„ Bulburin S»F« SEQ. 
{24'33\S, 15 1-27'E), under bark, MVF (12 r 
QMM021S62. 2° Jun 198Q, J. Stanisic. D. Poller, J. 

Othfr Matf.rial 
Graeemere. near Rockhaniptun 

C1404Q8, 18S7. C.E. Musson). 



Shell small, diameter 2.56-3.11 mm (mean 
2.S6 mm) Of 3 5/8 to 4 1/4 (mean 3 7/S 4) loosely- 
coiled whorls. Apex and spire moderately 

elevated, SP/BWW ratio 0.21-0.32 (mean 0.27), 
height 1,36-]. 70 mm (mean 1.55 mm). H/D ratio 
0.51-0,57 (mean 0.55). Last whorl descending 
only slightly or not at all. Protoconch exscrt, 1 
3/8 to 1 1,2 whorls, mean diameter 719|xrn a) 1 
1/2 whorls. Apical sculpture (Fig. 35q) of fine 
spiral cords with curved radial ridges toward the 
terminal part of the protoconch. Post nuclear 
sculpture (Fig. 35f) of prominent widely spaced, 
protractively sfnuated radial ribs, 70-89 ribs 
(mean 80) on the body whorl. Ribs/mm 8.30- 
9.33 (mean 8.92). Microsculpture (Fig. j5() of 
fine radial riblets. 4-9 between each pair of major 
ribs, crossed by closely spaced, fine spiral cords. 
Sculpture continuous on base (Fig. 35b). Um- 
bilicus wide, broad V-shaped, diameter 0.72- 
0.97 mm (mean 0.83). D/U ratio 3.22-3.55 (mean 
3.44). Sutures strongly to deeply impressed. 
Whorls rounded above and flattened below a 
rounded periphery. Aperture roundly lunate. Lip 
simple, columella only slightly dilated. Parietal 
callus moderately developed. Colour yellow- 
horn with reddish-brown flammulations in a 
radial pattern. Based on 4 measured adults, 

Ovotestis two clumps of palmately clavatc al- 
veoli, embedded in the digestive gland just above 
the stomach. Epiphallus internally with lon- 
gitudinal thickenings, entering penis apically 
through a simple pore (Fig. 36b). Penis (Fig. 
3bb) short, cylindrical, apically with a transver- 
sely sculptured, spatulate verge. Remainder of 
penis with low pustules arranged longitudinally 
(Fig. 36b). Penial retractor muscle originating on 
the diaphragm and inserted on the epiphallus. 

Central and lateral teeth of radula (Fig. 35g) 
tricuspid with long lanceolate mesocone and 
shorter ectocones. 

Based on 2 dissected specimens (AM- 
C 142934. QMMO 16593). 

RANGE and Haritvt 

S. nudicosiata is known from a handful of 
specimens found in watfll moist noiophyll vine 
forests of the Bobby Range area, SEQ. The 
single specimen collected by C.T. Musson from 
Graeemere near Rockhampton, SEQ, has been 
included because of its historical significance. 
However, I consider that the locality is wrong 
and probably the result of a labelling error. S. 
nudicostatu lives under logs and bark. 


The absence of penostracal setae and altered 
coiling pattern in 5. nudicostam are major depar- 
tures from typical Setonwdca. 



Setomedeajanaesp. nov. 

(Figs3,v-4ri, Tablets, 10) 


For Jan Chaseling who accompanied mc on 
numerous collecting trips. 

( "Mt'ARlSQVS 

The squiggly, apical spiral cords, curved radial 
ribs, periostracal setae and ttammulated g] 
Combine to separate S. janae from othci sym- 
patic charopids. 'Cyrocochlea udoidea (For- 
k's.. 1851) from Port Molle (= Aitlie Beach). 
MEQ, is larger with monochrome brown shell. 
depressed spire, simple conspicuous radial ribs, 
greater whorl height, and has no petiositaeul 
setae, Biampkalapa recsva (Fig. 57u,b.c» from 
the Eungella region, has a much larger, bicon- 
cave, flammulated shell with apical sculpture of 
vague pits, and no periostracal setae. .S. jtinuc can 
be distinguished from its congeners by its small 
whorl count, almost regular spiral cords (Fig 
*8e), and strnngls scalloped periostracal blades 
on major radial ribs (Fig 381", g). 

Typf Mateuiai 

bMiniMT: AMC I424h^ Ml DryanuVr, ill mil. 
ol Proserpine, MFQ. Collected by I . Price. 16 June 
i «5S. Height 1 ,83 mm. diameter 3 71, D/U ratio 2.S2. 
H D raiit>lj.4y. whorls 4 J/?. 

P \ratypes: Mt Dr>jndcr. Ml miles E o! Prospering. 
MEQ (I. AMC150W2, lo Jun l^5S. I.. Price. 3, 
AMCI424oS. 4 Jul 1964.1 Prico; Eungella Range. 
SO miles Wot Macka.v, MEQ (2, AMC 142470, Sept 
1957. L. Price); Olmara Hills, near Eungella \V of 
Yl.tekav MEQ. rainforest (1. AMC 142488, Apr 1975. 
D.*L McAlpine); Dalrymple Heights. Eungella N.P . 
MEQ (21 : U2~S. 148 36*E) lOOfl m NVF (5. 
OMMOri371. Nov 1976. M.J. Bishop). 

Other Material 

Ml William N.P.. N of Mackay MEQ, 1120 m 
i Tors. 148 B 36'E)(5. AMCI424S7. 27 Apt r- 
LB. Burch. W.F. Ponder. P.H Colmtni Dalrymple 
Heights N.P.. NW of Mackay, MEQ. 1000 m (7 
AMC 14248b. J.B. Burch. W.K, Pondrv, P H < nlman 
■ ■■pr 1975): Mi Macartney Caihu ST.. MEQ. *tfck 
luushinus (20 51*5, 148'33'E) rainforest. 750 m [2, 
M01 155, 22 Apr 1979, G.B. Moolejth); Mount 
William, Eungella N.P., MEQ, rainforest. ilOU m, 

suck bfushings (2i = u2 , s, i4sT36'E) (I. qm- 

M017156, I*' Apr 1979, G.B. Montciih). 


Shell depressed, diameter 3.41-3.71 mm 

(mean 3.h3 mm), with 4 12 to 4 3.4 (mean 4 
5.H-) normally coiled v, horlv I;isl whorl 0. 
cending more rapidly. Spire elevated, apex flat 
to slinhtlv elevated (Fie. 38b). SP/BWW ratio 
0.04-0.17 (mean 0.12)7 height 1.66-2.08 mm 
(mean 1-87 mm). HD ratio 0.49-0.58 (mean 
0.5 1 ). Protoconch shiny, ot 1 3/8 to 1 5/8 whorls, 
mean diameter n24.2|j.m a! 1 1/2 whorls. Apical 
sculpture (Fig. 38e) of continuous, low, narrow 
spiral cords, only Slightly squiggly: and irregular 
curved radial tibs. Pos! nuclear sculpture (Fig. 
38f) "I deep spiral furrows and prominent com- 
plex radial libs, regularly spaced, 49-80 (me;u. 

59,7J on body whorl, Ribs/mm 4.57-6.87 (mean 
5.22). Microsculpturc (Fig. ISjg) of crowded, 

scalloped tadiaJ nblcts. 7-14 belvscen each . 
of major ribs, crossed by crowded microspiral 
cords, which continue onto the sides of the 
radials. Major radial ribs with setal processes ;n 
regular intervals. Sculpture and setae continuous 
on the base. Interstices of the major ribs 7-8 

times their width Umbilicus (Fig 38b) wide, 
U-shaped, diameter 1.01-1.31 mm (mean 1.18 
mm). D I .: ratio 2.68-3.68 (mean 3,1 1). Sutures 
impressed. Whorls (Fig. 38c) shouldered al- 
and rounded below a slightly compressed 
periphery. Aperture roundly lunate. Lip simple. 
columella slightly thickened. Parietal callus 
developed- Colour hrownish yellow-horn with 
irregular darker brown suffusions, continuous on 
the base. Based on 7 measured adults. 

Genitalia with epiphallus (Fig. 39c) shortened 
and partially incorporated into the penial bulb. 
opening into the penis apicallv through a simple 
pored ig. V)c) surrounded by a muscular collar. 
Lower penial chamber with many raised, cor- 
rugated, fleshy, longitudinal pilasters (Fig. 3 
Peiu; Loi muscle inserted on the epip' 

Radula (Fig. 3Sd. h) without unusual features 
Based on 7 dissected specimens (AM- 
C1424S6, AMCI42487. QMMQ6371). 

! I V I'll \HITAT 

S. janae is restricted lothe montane re fug:. 
the Clarke Range in the Eungella area (= Dal- 
rvmple Heights), MEQ, with an outlier popula 
tiun on Mt Dryander, north of Proserpine, MEQ. 
S. janm- prefers moist warm subtropical 
aotaphyll vine forest. Unfortunately no altitude 
data arc available for the records from Ml. 
Dryander w here several forest types are present 
The micmhabitat of S. janae is not known but it 
has been collected from slick brushitigs by Dl 
GM. Montcith in two separate instances. 





«»i -&?■ 






FIG. 39. Setomedca janae sp. nov. Dalrymple Heights N.I'.. NW of Mackay. MEQ. AMC142486. a, genitalia; 
b, ovoteslis; c, details of penis interior; d, hermaphroditic duct; e, talon; I', pallial cavity. Scale lines as marked. 

FIG. 38. Setomedca janae sp.nov. a-c. Mt Dryander 10 mis E of Proserpine, MEQ. AMCI42469, holotype; 
C-g, Euneella Range, 50 mis \V of Mackay, MEQ. AMC 14247(1. paratype; d, h, Dalrymple Heights, Eungella 
N.P., MEQ. QMM0637K paratype. a-c,'entire shell; d. central and lateral teeth; e, apical sculpture; f, post 
nuclear sculpture; g, post nuclear region showing spiral furrows: h. lateral teeth. Scale lines as marked. 




S.janae is closely related ioS. seticostata. The 
sheathing oi' part of the epiphallus and apical 
penial bulb (Fig. 33b) in 5. seticostata tan be 
readily translated into the compacted epiphal- 
lus ■ penial bulb complex of the fcwer-vvhortedS. 
j7WMe(Fig. 39c). The apical sculptures < Figs 32e, 
38e) of the two species are alike and quite dif- 
ferent from the squiggly apical cords seen in 
other Seiomedea. The exaggerated scalloping of 
the periostraca! blades in S.janae (Fig. 38g) can 
be derived from the 5. seticostata condition by 
simple increase in the intensity of the spiral 

Setomedea monteithi sp, now 
(Figs 41^3; Tables 8, 10) 


For Dr G.B. Monteilh in recognition of his 
efforts in collecting material used for this study. 


S. monteithi is the largest species of the genus 
with mean whorl count 5 J/4-, average shell 
diameter 4,60 mm, and average shell height 2 t 7S 
mm. The large size, squigglv apical cords, tight 
coiling pattern, and weakly scalloped periost- 
eal rib blades, distinguish S. monteithi from its 
congeners. S. settct>stata (Fig. 32) is smaller with 
regular apical cords; S. janac (Fig. }$) has fewer 
whorls, strongly scalloped pcriostracal blades 
and regular apical cords; and Setomedea sp. (Fig. 
44) is smaller, with looser coils and narrower 
umbilicus. Onokcra cumulus Odhner, 1917 and 
O. nimbus Stanisic. 1987 have high spired, 
keeled, tlammulated shells with strong radial 
sculpture and no pcriostracal setae (Stanisic. 
1987, pis 1,2). Biomphalopa cone in na has a 
smaller, tesselated, biconcave shell with 
prominent radials and wide umbilicus (Fig. 6U). 
'Endodonta' intermedia Odhner, 1917 has a 
very small monochrome brown shell with more 
crowded radial ribs and no setae. 

TYfL Material 

HOLOTYPE: AMCU24SW, Twelve Mile Scrub, near 
A, ion. BtoomfleW River. NFO 00 in (is S0"S. 

I45 c IM'F). Collected bv P.H. Colman. 25 Oci. 1975. 

FIG. 40, Distribution Oi St-trtmcdca janae\. in 
mid-east Queensland. (Pr = Prosperine; Ma = Mack- 
ay; Sa = Sarin a) . 

Height ot shell 2.94 mm. diameter 4. 60 nun, II D ratio 
0,64, OH ran.- 3 20, whorls 5+ . 
AMO50090, 5 Specimens, S^rnfi ioealiu data as 
holoiype; Mi Sorrow, Cape Tribulation. 30Q-SQ0 m. 
r;.mt'orest(5.0MMOl7I6J. 150et 19S0. G.B. Mon- 
teilh); Ml Lewis, c. 17.8 km along Julatten-Mi Lewis 
Rd. MFF. litter to. QMMOI47M, 27 Jun 1983, J. 
Stanisic. D. Potter): Gap Creek. Ayton, Blonmfield 
River, lowland rainforest (4. AMC142472, R. Rad- 
nell. 1^75); Frit/ Creek S.F., near Ayton, Bloomlield 
River. 30 m (15*50*5. 145 21 F) {2S. AMCISOOM^. 
24 Oct 1975, P.H. Colnun); Mt Finia) S.F., near 
As ton. Bloomlield River, 2#5 m ( 15*50*5, j 45 "20'E) 
(7. AMC150086, 25 Oct 1475, P-H. Culman); Char- 
nollan Creek Crossing, on Ravenshoe-Tullv Falls Rd, 
NVF ( I 7 ; 42'S, 145 5l X) (X OMMOI4754, 28 Sept 
i, J. Stanisic); Mt Battle FrfiTC, summit centre 

FIG 41. Setomedtui mottttHthi ^p nov, a-c. Twelve Mile Scrub, near Avion. Bloomlield River, NFQ. AMC 
142489, holoiype; d. g. Frilz Creek S.F M near Avion. Bloomfiehi River. NEC AMC 150089, puraiypj; c-F 
same data as holOtypC, A\1( 140090, paratypc. a^C entire shell; d. marginal teelh;e. apical sculpture; F, post 
nuclear sculpture: g, central and lateral lech. Scale lines as marked 













FIG. 42. Setomedea monteithi sp.nov. a, b, d, f, Mt Finlay S.F., near Ayton, Bloomfield River, NEQ. 
AMC150088, paratype; c, Fritz Creek S.F., near Ayton, Bloomfield River, NEQ. AMC150089, paratype; e, 
Mt Bellenden Ker, NEQ. QMM011346, paratype. a, genitalia; b, spermatophore; c, ovotestis; d, talon; e, 
details of penis interior; f, pallial cavity. Scale lines as marked. 





H5 ft E 


FIG. 4-3. Distribution of Seiomedea mnntdifii sp.nov. 
(squares), and Setomedca sp. (dots) in north-east 
Queensland. (Mo = Mossman; Ca = Cairns; In = 
Ingham: Tu = Tully). 

peak. 1540 m. SMVFT (irM'S, 145 J 48'E) (14, 
QMMOl 141 1. 7-8 Nov 1981. Earthwatch/QM); Ml 
Bellenden Ker, 500 m, CMVF (12. QMMOl 1324, 
25-31 Dei 1 ^M. Earthwatch/QM); Ml Bellenden Ker, 
1054 m, SNVF(I50, QMM011346, 1-7 Nov 1981, 
Earthwatch/QM); Mt Finnigcin, 37 km S of Cooktown, 
S50- 1100 m, rainforest (7, QMMO 15482. 1 9-22 Apr 
1382, G.B. Wpntellfa, D. Yeates, D Cook); Mt Bartle 
Frere. NW to centre peak. 1400-1500 m (1, 
QMM015495. 24 Sept 1981, G.B. Monleith, D. 
Cook); Mt Bellenden Ker. summit TV Stn. 1560 m, 
SMVFT{I7 r lfVS, 145°5rE)(l,QMMO!5805, 1-7 
Nov 1981 Earthwatch/QM); Shiptnn's Flat, c. 25 km 
S of Cooktown, 250 m (1. QMM015477. 22 Apr 
1982, G.B. Monleith, D. Yeates, D. Cook) 

Othfr Material 

summit, 1560 m. SMVFT (18, QMMO U 2nd. 17-24 
Oct 1981, Earthwatch/QM); Leichh3rdt Creek Cross- 
ing, on Mt Lewis Rd, Mt Lewis, under logs. SMVFF 
(16 C 36'S. 145 a 16'E)(l.QMM014755. 1 Oct I9S0J. 

Stanisic), 2 km N of Mt Lewis, rainforest. 1000 m. 
litter (1. QMM015436, 9 Sept 1981, G.B. Monleith, 
D. Cook); Noah Creek, via Cape Tribulation, rain- 
forest [1. QMMO17160, 16 Oct 1980, G.B. Mon- 
leith): c; 2.5 km N of Mt Lewis. 1040 m (1, 
QMM015431. D. Yeates, G. Thompson, 3 Nov 
1983): Charmillan creek crossing, on Ravenshoe - 
Tully Falls Rd. NVR litter (3. QMMOI6231, J. 
Stanisic, 28 Sept 1980): Mt Bartle Frere, west face, 
700-1000 m (1. 0MMO17I64, G.B. and S.R. Mon- 
teith. 7 Oct 1980); North Bell Peak. c. 20 km S of 
Cairns, *5 0-1 000 m (1. QMMOI5493, G.B. Mon- 
teith, D. Yeates. G. Thompson. 13 Oct 1982); hill 
above Daintree River valley, via Daintree CMVF 
(16 C 12'S, 145 c 16e)(t.QMMO16270,2Oct 1980, J. 
Stanisic); Mt Bartle Frere, walking (rack below NW 
peak. c. 1000 ra, SNVF (1, QMMOl 1380, 8 Nov 
1981, Earthwatch/QM); Mt Bellenden Ker. summit 
TV stn. 1560 m, rainforest (17 lo'S. 145 51'E) p, 
QMMOI5M10. J-* Nov 1981. Earthwatch/QM). 
Cape Tribulation, lowland rainforest (I, 
OMM017162. 12-190ct I980,G.B Monteith); 2 km 
N of Cape Tribulation, 50 m, rainforest, stick brushing 
<16"05"S. 145 a 28 E) (1. QMM015525. 2 Oct 1 
G.B. Monleith, D. Yeates. G. Thompson), Mossman 
Gorge (2. AMC142471. 26 Nov 1963. D.F. Mr- 
Michael, J. C. Yaldwvn). 


Shell medium to large, diameter 3.62-5. 71 mm 
(mean 4.6 mm) with 4 1/2 to 5 5/8 (mean 5 1/4-) 
normally coiled whorls, last whorl descending 
much more rapidly. Spire and apex weakly to 
stronglv elevated, SP/BWW ratio 0,05-0 3* 
(mean 0.26), height 2.04-3.37 mm (mean 2.78 
mm), H/D ratio 0.50-0.83 (mean 0.61). 
Protoconch glossy, of 1 1/2 to 1 5/8+ whorls. 
mean diameter 725.1 (xm at 1 1/2 whorls. Apical 
sculpture (Fig. 41e) of numerous squiggly spiral 
cords and a few. weak, curved radial growth 
ridges. Post nuclear sculpture of complex major 
radial ribs and broad spiral grooves. Ribs 
regularly spaced, 50-107 (mean 71,50) on (he 
body whorl. Ribs/mm 4.12-6.71 (mean 4.91). 
Major ribs consisting of two riblcts with high 
periostracai blades, the one on the apical side 
with long periostracai setae arranged at regular 
intervals along its length. Ribs appearing scal- 
loped due to broad spiral undulations in the shell 
surface. Mierosculpture (Fig. 4 If) of numerous 
radial riblcts, 7-15 between each pair of major 
ribs, and fine, closely spaced spiral cords which 
continue onto the radial ribs Interstices of major 
ribs c\ 6-8 times their width. Sculpture continued 
on the base. Umbilicus wide U-shaped, diameter 



i mm 

^^^ :.■■■■ 


1 mm 



FIG, 44. Setomedea sp. a-c, Majors Mtn, 7km SE Ravenshoe, NEQ. QM MO 1 5503; d-f, c. 10 mis E of 
Ravenshoe, on Palmerston Hwy. NEQ. AMC142490. a-d, entire shell; e, apical sculpture; f, post nuclear 
sculpture. Scale lines as marked. 



0.92-1 .85 mm (mean 1.36 mm), D/U ratio 3.04- 
3.92 (mean 3.41). Sutures strongly impressed. 
Whorls shouldered above and rounded below a 
slightly compressed periphery. Aperture roundly 
lunate. Lip simple, weakly thickened and twisted 
at the columella and slightly detached in larger 
individuals. Parietal callus well developed. 
Colour yellow-brown with darker brown zig-zag 
suffusions arranged in a radial pattern, con- 
tinuous on base. Based on 19 measured adults. 

Genitalia with epiphallus (Fig. 42a) short, 
muscular, forming a half loop around the penial 
retractor muscle before entering penis sub-api- 
cally through a simple pore (Fig. 42b). Penial 
retractor muscle long, inserted adjacent to the 
penis/epiphallus junction (Fig. 42e). Penis (Fig. 
42e) pear-shaped, internally with epiphallic 
entrance surrounded by two spongy pilasters; a 
muscular collar separating the apical bulb from 
the lower chamber; and two flattened, fleshy 
pilasters lower down. 

Central and lateral teeth of radula (Fig. 41d,g) 

Based on 8 dissected specimens (AM- 

Range and Habitat 

S. monteithi inhabits the humid tropical region 
of north Queensland between Tully and Shiptons 
Flat, south of Cooktown. It is found from near 
sea level to the mountain summits where vegeta- 
tion varies from complex mesophyll vine forest 
to cool subtropical notophyll vine forest and 
montane microphyll vine fern thicket. 5. mon- 
teithi is well represented in museum collections 
but its distribution is still inadequately known on 
the Atherton Tableland and towards the southern 
end of its range. Further field work should con- 
centrate on establishing the degree of sympatry 
with Setomcdea sp. The microhabitat of 5. mon- 
teithi is similar to that of 5. seticosiata i.e. in moist 
rotting logs and moist accumulated forest debris. 


The penis (Fig. 42e) shows some similarity to 
that of S. seticosiata in having an apical bulb 
with verge-like pilaster, muscular collar and 
lower chamber with fleshy longitudinal 

Setomedea sp. 

(Figs 43^4) 

Lake Barrine, Atherton Tableland, rainforest (1. 

QMM017163, 8 Oct 1980, G.B. Monteith); Majors 
Mtn, 7km SE of Ravenshoe, 1000-1100 m (I, 
QMMO15503, 4 May 1983, G.B. Monteith, D. 
Yeates); c. 10 mis E of Ravenshoe, on Palmerston 
Hwy, under fallen timber in rainforest (1, 
AMC142490, 28 Dec 1979, 1. Loch, J. Farquharson). 


Compared with S. monteithi the shells of 
Setomedea sp. have a larger number of ribs, 
smaller umbilicus (diameter 0.72 mm), less 
elevated spire (spire protrusion 0.17 mm) and 
overall smaller shell with diameter 3.62 mm and 
height 2.13 mm (based on QMMO15503). 

While this species is not formally described at 
this time, it is listed in order to assist with the 
biogeographical discussion. The reasons for the 
narrow distribution of Setomedea sp. on the 
western edge of the Atherton Tableland are not 
clear. However, this region does appear to have 
status as a refugium of unusual nature. The 
primitive camaenid land snail, Craterodiscus 
pricei McMichael, 1959 and the New Guinea 
related dasyurid mammal Antechinus leo Van 
Dyck, 1982, are two examples of animals with 
small circumscribed distributions in this area. 

Gyrocochlea Hedley 1924 

Gyrocochlea Hedley, 1924 (part), p. 215; Iredale, 
1937a (part), p. 322; Iredale, 1941a (part), 
pp. 267ff; Kershaw, 1955, p. 29; Burch, 1976, 
p. 132. 

Type Species 

Helix vinitincta Cox, 1868; by original desig- 

Previous Studies 

Gyrocochlea originally included several east- 
coast charopids characterised by a shell with 
sunken spire, wide umbilicus, "smooth" 
protoconch and prominent crowded radial ribs, 
plus Helix stroudensis Cox, 1864 which has an 
exsert spire and closed umbilicus. Iredale 
(1937a) added an additional four species and 
(1941a) removed H. stroudensis. Currently 14 
species are included (Burch, 1976). Anatomical 
studies of seven species show Gyrocochlea to be 
polyphyletic. Gyrocochlea is restricted to G. 
vinitincta (Cox, 1868), G. convoluta Hedlev, 
1924, G. curtisiana (Hedley, 1912) and G. 
paucilamellata sp. nov. 'Gyrocochlea' omicron 
(Pfciffer, 1851) belongs in Nautiliropa gen. 



tiov ; (jyrocorhlea rccava ( Medley. 1912) and 
'Gymcoehlea' voncinna Hedley, 1924 arc 
placed in Biomphalopa gen. nov. 


Adult shell diameter 4.71-8.57 mm. wuh 4 l/s 
Ki 4 J/4 tightly to moderately tightly {cunisiana) 
coiled whorls, last descending more rapidly 
Apex and early spire strongly lu weakly (cur- 
tisiana) concave. Protoconch of 1 3,'S to 1 5/8 
whorls. Apical sculpture of fine crowded spiral 
cords and numerous similarly spaced, Weakly 
curved radial ribs forming fine heads at their 
intersection, or more widely spaced spiral cords 
and weakly curved, radial ribs which arc partial- 
ly (used and worn, giving Ihe ;ipe\' a pitted ap- 
pearance [curtistUHaJ. Post-nuclear sculpture of 
crowded (vtnitutciii, curusiurta), moderately 
spaced (convaluta) or widely spaced 
\pQitahvuettata), prominent, weakly protec- 
tively srnuated radial ribs. Microseulpture of 
numerous radial rihlets and more crowded 
micrnspirul cords which "buttress" the 
miiToruduK. Microspirals continuous on the 
rnajoi radiate. Umbilicus wide. U-shaped, or V- 
shaped {curtisianay Sutures deeply impressed. 
Whorls rounded below, and strongly to weakly 
(curtisiana) shouldered above a We ail} rounded 
(< urfisuma) to flattened periphery. Aperture 
ovately lunate Lip simple. Colour light-yellow 
horn to dark reddish-brown, occasionally (cv/r- 
tisiana) with red flammuialions. 

Animal with long, slender foot, often darkly 
pigmented. Pallia! cavity with bold black speck- 
ling on the palhal roof, Kidney with elongate 
pericardial lobe and rectal lobe reduced. Ovoics 
lis oriented al right angles (vimtincta) or parallel 
to the plane of coiling. Male genitalia with long 
coiled epiphallus and long tubular penis with 
sheath. Epiphallus entering penis subapically 
rhrongh a conical verge (viniiificta) ot simple 
pore. Terminal portion of epiphallus connected 
10 the penis sheath by fine connective strands. 
Penis internal I J with large longitudinal pilas ert 
and with, or without (yiniiincta)^ smaller 
crowded pilasters near and above the epiprir.lhc 
pore. Female genitalia with long vagina and 
shorter I rce oviduct . Radula without unusual fea- 


(7v/-ocor/;/o?iih.ibds dry to hum id subtropical 
vine forests of the McPhcrson Ranges. SEQ, the 
Big Scrub area northern NSW, and humid 
notophyll vine fOrftSI and adjacent drier 

microphyll vine forest of the Burnett - Many 
Peaks - Calliope Ranges area of southern 
Queensland. Gyrocochlea prefers to live under 

Pattfrvs or Variation 

Species o\' Gynnochlca arc rcmarkablv con- 
servative with regard to shell features differing 
only in si/e and intensity of radial ribbing except 
lor G curiisPana which has a less sunken spire, 
modified apical sculpture, more ribs/mm on the 
adult whorls, less crowded microsculpture,more 
rounded whorl contour and V-shaped umbilicus. 
The pitted apex in (J, ruritsiana is derivable by 
partial fusion and reduction in height of the equal 
spiral-radial elements. The same effect is seen in 
some unrelated northern Australian (Solem. 
I 9X4) and New Guincan (Solcm, 1970) genera. 

Anatomically (iyrororhlea is equally conser- 
vative. There is some difference in length of 
reproductive organs and minor changes in the 
penial pilasters. Only G. vimtincta differs 
dramatically in having a conical verge and no 
pustular zone apicad of the epiphaliic entrance, 
probably because wi its svmpalry with G. 
pnudhinv:!hio G. unutfu t<> has ihe two lobes 
or the DVOtCSttS at right angles to the plane Of 
coiling but not in the same plane as is seen In 
\>xuirca, On'okaa. and Uetilexoamcha. The 
lobes are siluaicd ill ihe apical and basal margins 
ol die whorl rather than cent ralh . This positional 
change is just a different wa\ of dealing with 
space changes brought about by size increase. 

Conservatism in anatomy is probably related 
to ihe small, mainlv allopatric ranges of in- 
dividual species which minimise interspecific 


Conehologicnlly GyrocQctUvU is most similar 
to blautiliropa with which it is svmpatric in the 
eastern McPhcrson Ranges on ihe NSW/QLD 

bordci. Both genera have biconcave shells and 
prominent ,»osi nuclear radialsbul di Her in sculp- 
ture and coiling pattern. In the McPherson Ran- 
ges species otCyrororh/ra have apical sculpture 
o\ raised spiral cords and radial ribs in a 
crowded, reticulate pattern producing a beaded 
appearance (Figs 45c, 48e. 51c), adult sculpture 
of major ribs which are formed from several 
minor ribs and lacking a periostracal blade (Figs 
I i 48f. 5 I f), and a body whorl which descends 
rapidly in front (Figs 45c, 48c. 51c). On the other 
hand Xamilimpo has apical sculpture of regular- 
ly arranged pit5 (Fig. 63d), adult sculpture of 



prominent radial ribs which are formed by ex- 
pansion of a single minor rib and with a high 
perfostracal blade (Fig. 63e), and a body whorl 
which docs not descend (Fig. 64f). Anatomically 
the elongate genitalia, tubular, sheathed penis 
and longitudinal penial pilasters of Gyrococh lea 
contrast with the compact genitalia, pear-shaped, 
sheathless penis and complex pilaster arrange- 
ment of Nautiliropa. 


l. Apical sculpture beaded 2 

Apical sculpture of regularly arranged pits 
(Fig. 54c) curfisianQ 

2. Post nuclear sculpture of crowded ribs (Fig. 
45a); shell larger (mean dianielei > 7.00 
mm) i vmitincia 

I'im nuclear sculpture nfnuire widely spaced 
ribs; shell smaller (mean diameter < 7,00 
mm) 3 

3 Rjbs on botty whorl very widely spaced (Fig. 
49a); mean ribs/mm 1.56 pauviUtmclforo 

Rihs Otl body whorl moderately spaced ( Fit;. 
4Na):meunnlis/mm 3.92, , , ,. COlWOlHtA 

Gvrocochlea vinitincta (Co\, 1868) 
(Figs 45^7; Tables! 1.1 2 j 

//« H.x vinitincta Ox, 1868 p IS, pi I. figs fi, (>a 
| h[n'<„npholus vittitinctus (Cox); Tryon, 1XKS, p. 1 15. 

pi. 24, (>gsXS.S9 ( 
Ktottnmftlniltri vtnititictus (Cos): Ancey, 1888. p. 

tjjdottontd vinitincta {Cos); Shi)l<:\ ( |92l, p 34. 
GyrocochlcA vinitincta (Cox); Hedlcv, 1924, p. 218, 

pi. 31. figs 28-31; Iredale, l'>37a, p. 322; Iredale, 

I'-Ul,, p t 2$8, Tig. ft; Kershaw. 1956*1, p. L3S. 


The large size, simple sutures, beaded apical 
sculpture, and strong!) depressed spire differen- 
tiate this species from the sympatric ELsothera 
xcniihccalo. which has a flat to slight!)* elevated 
spire with strong curved radial ribs on the apex, 
channelled sutures and very inflated body whorl. 
G. vinitincta can be distinguished from 0\ 
paiuilawcilrtdh and G. convoluta by its larger 

size and more numerous radial ribs. Nautiliropa 
omicrnn has the general shell form of G. 
vinitincta but is smaller, with greater number of 
ribs/mm, body whorl that does not descend, and 
flammulated shell. 

Type Material 

Holotvpe: AMC63490, Upper Richmond River. 
NSW. Height ol shell 4.87 mm, diameter 7.98 mm, 
H/D ratio 0.61. D/U ratio 3.<) l >, whorls 4 1/2-k 

ParatypE; AMC103624, same daia as huloiypc. 

Other Material 

Richmond River. NSW (5, AMCI39754; 3. AM- 
C 139769, Pellerd, ex Cox); Byron Bay, NSW (5, 
AMC1 3975?. S.W. Jackson ); Big Scrub, MurwilUim- 
bah, NSW (I, AMC63S37. Lower, ex Cox); Lismure. 
NSW (?, AMCI39752, ex C. McLauchlan, Bent 
Cull); Red Scrub Flora Reserve, Whian Whian S.F.. 
\S\V (28 3S\S. 15319 E), 210 m (3, AMC13975I. 
15 May !«J7o, P.H. Colman. I. Loch); Terania Creek. 
NSW(2S ; 34\S, I53 E T9'E),(3.AMC153725. 16 May 
1 976, P.H. Colman. I. Loch); Mt. WarningN.P., NSW 
(2S l 24-$. 153'16'E) NVF/Palms (2, QMMO104S7, 
Mar 19S1, AM QM-ABRS; 2. QMMOI2735, 
AMC136830, 15 Dec 1981, AM QM-ABRS). Upper 
Tallebudgera Ck. SEQ. 600 m (2. QMM01599D, 9 
Dec 1984, G.B. Montciih); Warrie Circuit, 
Springbrook N.P.. SEQ, CNVF, under log in lifter (1, 
QMMO16902, 6 Apr 1986, J, Sianisic) i. 1.6 km N 
ol* Numtnbah Valley Turnofl, on Burleigh- 
Springbrook Rd, SEQ, NVF/Palms, litter (2, QM- 
MOI7319, 17 Dec 1980. J. Stanisic); Natural Bridge 
62ftt,.Oci 197^ M. J. Bishop). 
LAMINGTON N P., SEQ; Border Track, near (T 
Reillys NVF. under logs (3. QMM01o*75, 31 Mar 
i" 84, ' i Loistrti Orchid Bower, Biima Burra* NVF, 
under stone (I. QM M0 16864. 19 Apr 1986. J. 
Sianisic. J. Chaseifilg); ntlllaWafW Circuit, Binna 
Buira.NVF.undeil.^' (jMMOU,739,2 0cl 1985, 
J. Sianisic, D. Poller, I. Chaseling); Border Track. 

B Bui a KVF. under log (3. QMM016869, 10 

Mar I9S4. J. Stannic): Araucaria Track. Binna Burra. 
NVF, under logs (I. QMMQ16B64, 19 Apr 1986, J. 
Stanisic. J, Chaseling): Tullawallal Circuit. Binna 
Burra. NVK under logs (7. QM MO 1 6739, 20ct 1985, 
J. Stanisic. D. Potter. J. Chaseling); Border Track, 
Binna Buna. NVF. under log (3, QMMOI6X6V, 10 
Mar 19S4, .1. Stanisic); Araucaria Track. NVF, under 
logs (3, QMMOT6579, 27 Sept 1986, J Stanisic): 
O'Reillys (28^14'S. IS3M5 B S) M FF (3, QM- 
MOHI43I. 17 Mar 1981, AYI/QM-ARKS): BuitU 
Burra (28"13'S, L33*I2*E) »5I1 m. CNV! <i. 
QMMO6078, Apr (976, M L Btewfi 










ml $k 


FIG. 46. Gyrocochleavinitincta (Cox, 1868). Mt Warning N.P., NSW. QMM012735. a, genitalia; b f penis 

interior; c, ovotestis; d, hermaphroditic duct; e, pallia! cavity. Scale lines as marked. 

FIG. 45. Gyrocochleavinitincta (Cox, 1868). a-c. Upper Richmond R, NSW. AMC63490, holotype; d, g-h, 
c. 1.6km Nof Numinbah Valley turnoff, on Burleigh-Springbrook Rd, SEQ. QMM017319; e-f" O'Reillys, 
Lamington N.P., SEQ. QMMO 10431. a-c, entire shell; d, jaw; e, apical sculpture; f, post nuclear sculpture; 
g, central and lateral teeth; h, marginal teeth. Scale lines as marked. 



3 =4 

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Fig 47. Distribution of Gyrocochlea xinitincta (Cox, 


Shell diameter 7.22-8.57 mm (mean 7.91 
mm), with 4 1/4+ to 4 5/S (mean 4 3/8+), tightly 
coiled whorls, last descending more rapidly. 
Apex and early spire (Fig. 45a) strongly con- 
cave, height of shell 3.95-4.87 mm (mean 4.52 
mm). H/D ratio 0.53-0.63 (mean 0.57). 
Protoconch of 1 1/2 to 1 3/4 whorls, mean 
diameter 896.7 (xm at 1 1/2 whorls. Apical sculp- 
ture (Fig. 45e) of fine, crowded spiral cords and 
numerous, weakly curved radial ribs, which are 
raised at their intersection with the spiral cords 
creating a beaded effect. Post nuclear sculpture 
(Fig. 45f) of numerous, high, crowded, protrac- 
tively sinuatcd radial ribs, 106-153 (mean 
1 1 8,8) ribs on the body whorl. Ribs/mm 4.22— 
5.68 (mean 4.77). Microsculpture (Fig. 451*) of 
fine radial riblets, 6-12 between each pair of 
major ribs, continuing onto the sides of the major 
ribs, and numerous fine microspiral cords which 
are not raised at their junction with the 
microradials. Umbilicus (Fig. 45b) relatively 
wide, U-shaped, diameter 1 .79-2.26 mm (mean 
2.00 mm). D/U ratio 3.46-4.32 (mean 3.97). 
Sutures (Fig. 45a) weakly impressed, becoming 
deeper on the body whorl. Whorls rounded 
below and strongly shouldered above a weakly 
compressed periphery. Aperture ovately lunate. 
Lip simple, columella not expanded. Parietal 
callus developed. Colour yellow-brown to dark 
reddish-brown. Based on 6 measured specimens. 

Genitalia with ovolcstis (Fig. 46c) oriented at 

-J < 



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FIG 49. Gyrocochlea convoluta Hcdley, 1924. Along tributary of Iron Pot Ck, 1km N of Toonumbar, NSW. 
QMMO 10927. a, genitalia; b, penis interior; c, hermaphroditic duct; d, pallial cavity; e, ovotestis clump. 
Scale lines as marked. 

FIG 48. Gyrocochlea convoluta Hedley, 1924. a-c, Beaudesert. Old. AMC5815, holotype; d, g-h ? Along 
tributary of Iron Pot Creek, 1km N of Toonumbar, NSW. QMMO10927; e-f, Mt Lindesay. Richmond R, 
NSW. AMC39750. a-c, entire shell; d, lateral teeth; e, apical sculpture; f, post nuclear sculpture; g, cusp 
angles on lateral teeth; h, lateromarginal and marginal teeth. Scale lines as marked. 



right angles to the plane ot coiling, one clump on 
the apex and one on the base of the whorl and 
directed toward each other. Epiphallus (Fig. 46b) 
kinked and entering the penis subapically 
through a conical verge (Fig. 46b). Penis (Fig. 
46b) a long* slender, muscuhinsed tube covered 
with a thin penis sheath. Upper portion ot the 
epiphallus connected to penis sheath by short 
fibres (Fig. 46a). Internally the penis has several 
longitudinal spongy thickenings (Fig. 46b). 
Radula without unusual features, 
Based on 2 dissected adults (QMM012735). 

Range and Habitat 

G, vinitinaa occurs in the Big Scrub area of 
northern NSW, the eastern MePhcrson Ranges, 
SEQ. and some adjacent mountains and valleys 
of the Scenic Rim. It prefers cod! lo warm sub- 
tropical notophyll vine forests and docs not in- 
habit adjacent drier vine forests wllh Anttteurtu 
rmergents. Specimens have been taken from 
under logs and under stones, often in pairs. 


G. vinitincta and G. paucilamcllaia are sym- 
patrie in parts of rhe Lamington Plateau. SEQ, 
and the structural differences m their penes 
probably provide species recognition features 
for the snails. 

Gyrocoehlca convoluta Hedlcv. 1^24 
( Mgs 4S-S0; Tables IE 1*2) 

Qyroeachka convoluta Hedley, L924(pan), p 216, 

pl. 2<>, tigs !-<)■ hetfafe, LM7a; P- 322 


G Convolute differs from G. paucilamcllaia in 
being slightly larger (Table 1 1) and in having a 
greater number of ribs on the body whorl. 
Anatomically the differences arc minor. (7. am- 
voluta (Fig. 49b) has more irregular longitudinal 
pilasters in the lower part of the penial chamber 
and a larger pustular zone apicad of the epiphal- 
lic pore than G. pauciIameHutu (Fig. 52b). G, 
convotuta is easily distinguished from G. 
vUHtittcta by its smaller size and more widely 
spaced radial ribs, and \romNauiilm>panmicron 
by its fewer radial ribs, larger umbilicus and 
monochrome brown shell nn which i he !;isl 
whorl descends. 


Gyrocoehlca convoluta was bused on the 
holotypc and specimens from Camerungo (= 

Uantinerai, SEQ, The latter material has not been 
found and efforts to collect at Canungra yielded 
only G. paucilameiUila\ Medley probably had 
both species. 

I vrr Material 

HoLOTYr-K AMC5S15. Beaudfisert, Old. Height of 

shell 3.36 mm, diameter 5.88 m, H/D ratio 0.57, D/u 

ratio J. 14, whorls 4 5/8. 

OrurR Materwu. 

Ml. Lindesay, Upper Richmond River, NSW (3. 
AMCl397S0,27Apjl95B;L Price), l. 1.7km along 
Mcintosh's Rd, nr Back Crock. I: side of Richmond 
Range, NtNSW [28 25'S, tSlM^'S), broad- 
leaf/srlempbyll (I. QMMO10970, 15 Mar 1981. 
AM/OM-AHKS); along irihmar\ of Iron Pot Creek. I 
km N oi raonumbaf, NSW |28"34'S, 152-450) (1. 
QMMOUW27, Mar N81, AM OM-ABRS). 


Shell large, diameter 5.46-6.05 mm (mean 
5.78 mm), with 4 t/2- to 4 3/4 (mean 4 5/8+) 
tightl) coiled whorls, last descending more 
rapidly. Apex and early spire (Fig. 48a) strongly 
concave, height ol shell 3t03-3.78 mm (mean 

3 36 mm). II/D ratio 0.55-0.63 (mean 0.58). 
Pmtoconch of I 1/2 to I 5<H+ whorls, mean 
diameter 7S5.5 fim at 1 1 ,2 whorls. Apical sculp- 
ture (Fig. 4Sc) \)\~ fine spiral cords and fine low 
radial thickenings, raised at their junction creat- 
ing a beaded effect. Poslnuclear sculpture (Fig. 
4Sf) of numerous, widely spaced, high, strongly 
protectively sinuated radial ribs. 58-83 (mean 
71.5) ribs on the body whorl. Ribs/mm 5.33- 

4 36 (mean 3i92). Microsculpturc (Fig. 48F) of 
fine radial riblets, IS— 30 between each pair of 
major ribs, and very fine, crowded spiral cords. 
Umbilicus (Fig, 48b) U-shaped, diameter 1 .62— 
1,S7 mm (mean 1.70 mm). D/U ratio 3.14-3.55 
(mean 3.40). Sutures impressed, becoming deep 
on the last whorL Whorls rounded below and 
strongly shouldered above a slightly flattened 
periphery (Fig. 48c). Aperture ovately lunate, 
Lip simple. Parietal callus developed, radially 
sculptured. Colour golden brown. Based on 4 red adults. 

Genitalia with epiphallus long, musculuriscd, 
bound for part of its length to the apical part of 
the penis (Fig 49tt) ( internally with longitudinal 
thickenings, entering penis subapically through 
a lleshy pore (Fig. 40b). Penis (Fig. 49b) long, 
tubular, without expanded apical portion, inter- 
nally with thin fleshy longitudinal pilasters (Fig. 
49b) which become less regular in the lower part 








FIG 50. Distribution oiGyrocochlcn convoluta Hed- 
!e\\ 1924. 

of the chamber, and a small pustuiose zone above 
the epiphallic pore. Penial retractor muscle in- 
serting apically on the penis. 

Radula (Fig.'48g, h) typical. 

Based on 1 dissected specimen (GMMO- 

Range and Habitat 

G. convoluta inhabits notophyll vine forests of 
the Upper Richmond Range. NSW, and gallery 
forests of creeks which flow from these high- 
lands. No data is available on microbabitat al- 
though shell form suggests that it lives under 


G. convoluta is a conehologicalty distinctive 
Species and differs only slightly from G. 
paucilamellata in genital anatomy 

Ctvrocochlea paucilamellatii sp. nov 

(Figs 51-53; Tables 11,12) 

iiyrocochlca convoluta Medley, 1 924 (part), p. 216. 


Lai'mpaua, few; lamella, plate; for the sparse 
number of radial ribs on the adult whorls. 


The brown shell, widely separated radial ribs. 
biconcave shell and wide umbilicus distinguish 

G. paucitamellata from its congeners and other 
charopids in the border ranges area. G. vinitincta 
is much larger with more numerous radial ribf 
(Table 1 1 ). G. convoluta has more than twice as 
many ribs on the body whorl. Anatomically G. 
ptiiicilawellata (Fig 52b) differs from G. 
\inhincia (Fig. 46b) by lacking a penial verge 
and in having an apical pustular zone in the 
penis. Nuultliropu otrucroru which also is sym- 
patic with G. paucilamellata, has many more 
radial ribs and a flammulated shell. Anatomical- 
ly A', omicron has a penis which lacks a sheath. 
Contains complex pilasters, and is pear-shaped 
(Fig. 65c). 

I i re Materiaj. 

Holotype- QMMO 17321. Upper Pine Creek, 
Canungra, SEQ. H\ T Y iAraucoria, underbark and logS- 
Collected by J. St^nisic, D. Potter, 1 Oct I98bl 
Para TYPES: QMM01o54fi t 21 specimens, same data 
as holotvpe; Kweebani Caves Walk, Binna Burra, 
Laminglon N.P., SEQ, NVF, under logs (1, 
QMMO"l6NN9. 3d Sepi 1985, J. Stamsie, D. Poiter.J. 
Chaseling); Mt Tamborine, SEQ, under bark in leaf 
titter (4, QMMO 17320, VK 7299. 3 Nov 1979, V 
Kessner); Upper Pine Creek. Canungra, SEQ, 
NVF/Araucaria, under log (1. QMMO 16585. 29 Sept 
1986. J.Stanisic : J, Chaseling). 

I >; \\ [N0SI5 

Shell large for family, diameter 5.04—5.55 mm 
(mean 5.35 mm) with 4 1/4- to 4 5/8 (mean 4 
1 ,2-) tightly coiled whorls. Apex and early spire 
(Fig. 51a) strongly concave. Height 3.19-3.45 
mm (mean 3.31 mm), H/D ratio 0.58-0.65 (mean 
0.62). Last whorl descending. Protoconch of 1 
1/2+ to 1 5;$± whorls, flat, mean diameter 745.3 
p,m at 1 12 whorls. Apical sculpture (Fig, 511") 
finely reticulate, consisting of numerous. 
crowded fine spiral cords crossing numerous 
crowded, weakly sinuated radial ribs. Post apical 
sculpture (Fig. 5 it) of lew. bold, protractively 
sinuated radial ribs, 27-40 (mean 32.7) ribs on 
the body whorl. Ribs/mm 1.57-2.52 (mean 
I .%). Microsculpture (Fig, 5 If) of fine, crowded 
radial riblets, 30—45 between each pair of major 
ribs, and very fine inconspicuous spiral cords, 
only visible between the microradials. Um- 
bilicus (Fig. 51b) wide. U-shaped, diameter 
1.4°--!. 75 mm (mean 1.65 mm). D/U ratio 3.13- 
3. 38 (mean 3.25). Sutures impressed, forming a 
deep furrow as the body whorl descends. Whorls 
rounded below and strongly shouldered above a 
weakly rounded periphery (Fig. 51c). Aperture 
QVatcly lunate. Lip simple. Colour deep reddish- 









PRM " 

FIG. 52. Gyrocochka paucilamellata sp. nov. Upper Pine Ck. Canungra, SEQ. QMM016546, paratype. a, 
genitalia: b, penis interior; c, talon and hermaphroditic duct; d. spermalophore; c, ovoteslis: f, pallial cavity. 
Scale lines as marked. 

FIG. 51. Gyrocochlea paucilamellata sp. nov. a-c. Upper Pine Creek, Canungra, SEQ- QMM017321, 
holotype: d, g-h, same data as holotype. QMM016546, paratype: e-f, MtTamborine, SEQ. QMMO17320, 
paratype. a-c, entire shell; d, jaw; e, apical sculpture; f, post nuclear sculpture; g, central and lateral teeth; h, 
marginal teeth. Scale lines as marked. 



21 n ; 

152 D E 


154 "3 

FIG 53. Distribution of Gyrocochlca paua/amcllata 
sp. nov. 

brown to light-yellow horn. Based on 3 
measured adults. 

Genitalia with cpiphallus long, muscular, and 
strongly coiled, entering the penis subapically 
through a simple pore (Fig. 52b) surrounded by 
a fleshy collar. Epiphallus partially bound (api- 
cally) to the penis sheath. Penis (Fig. 52b) elon- 
gate, tubular, mtiseularised. internally with 
longitudinal pilasters (Fig. 52b) in a regular pat- 
tern, and smaller pilasters apicad a\~ the epiphal- 
lic pore. Penial retractor muscle inserting 
apical I y on the penis. Spermatophore subclavatc 
with a long recurved chitinous tail at one end 

Radula typical. 

Based on 4 dissected specimens (QMMO- 

Range ,\\n Habitat 

(i. pauctlamcllata occurs in the eastern part of 
the Lamington Plateau. SEQ. and in the Dar- 
lineton-Canungra Range area as far north as Mt 
Tamborinc. SEQ in cool subtropical to warm 
subtropical notophyll vine forest- sometimes 
with Araucaria emergents (Upper Pine Creek, 
SEQ1 The species has been collected from under 
fallen bark and logs. In spite of relatively con- 

centrated collecting in the Lamington Plateau 
area, it has been found there only once. 


(1. paucilamellaw has a distribution quite dis- 
tinct from that of 0. convoluta and is geographi- 
cally isolated from it by the Albert River Valley. 

(ivrocochlea curtisiana (Hedlcy, 1912) 
' (Figs54-5o; 64a-c: Tables 11,12} 

Endotlontu iuloidea var. curtisiana Hedley, 1912, p. 

264, pi. Q , figs 4W-51. 
Cj txa •'• fifed curtisia&a (Wedlev); Iredule. 1937a. p. 
' 323. 


6\ curtisiana is distinguished in the genus by 
its almost Hat spire, apical sculpture of regularly 
arranged pits, often flammulated shell and large- 
number of ribs on the body whorl. Sautiliropa 
omicron has apical sculpture of regularly ar- 
ranged pits, flammulated shell and large number 
of ribs on the body whorl, but has a very 
depress.: J spire and a body whorl which does not 

i 1 - . imi SSTUDTES 

Hedlcy (1912) described G. curtisiana as a 
variety oi 'Helix' ntioidca Forbes, 1851 from 
Port Molle. Central Queensland, following com- 
parison of specimens with the type of//, mlnidea 
In- E.A.Smith {Hcdley. !912,p. 265). Hedley (in 
Hedlcy and Musson. 1892) had included this 
variety with x Charopa anticron{Pfe\ftex^ 1851). 
The Two species arc similar in shell appearance 
but are anatomically distinct. Hedlcy (1924) 
listed a large number of species under 
G\ rocochlea ' without including his new variety 
but recorded 'Gyrocochlca' omicron as having a 
distribution as far north as Miriam Vale, SEQ. 
There is little doubt that these early published 
records of 'G\ omicron from the Miriam Vale 
aii I were based on specimens oi G. curtisiana. 

Tvpf M\ rt rim 

HOI onTF: AMI 329y0, "found under log in forest 
just outside Springs Scrub". Ml I atcom. Port Curtis. 
SEQ. C olleeted bv SAW Jackson. Aug 1908, Heigh) 

FIG 54. tfVracachlea curtisiana (fledlev. 1912). a-c. Granite Creek, foothills of Bohbv Ranue, SEQ. 
OMMQU7I6: d-h. Kroombu Tops, SEQ. QMM01-65H QMM01M5& a- e. entire shell; d. central and 

lateral teeth; e, apical sculpture; f, post nuclear sculpture; g. central and lateral teeth; h. marginal teeth. Scale 
In i t! marked. 












FIG 55. Gyrocochtea curtisiana (Hedky, 1912), Kmombit Tops. SEQ. QMM0165I7. a, genitalia: b, penis 
interior: c. hermaphroditic duct; d, ovplestis; e, talon: 1, patlikl cavity. Scale lines as marked. 



2 3 n 3 

:5 P S 

26 C S 



15I C E 



Pitt 5h DiMrihulion afGyracochk'u vuriisuuut (lied 

lev. i" i | 

of shell 2.52 mm. diameter 5.2^ mm, H/D raSfo 0.48, 
D EJ ratio 3.4&.wfanris4 1/4 

i ii k Material 
Summit. Mi Booroon Booronn. SW o! Miiam Vale. 
SEQ. WW ArauairuL liner (4. QMMOKnSM), 17 
Sepl 1985, -I- St anisic, D. Potter); Gr^niic Creek, 
toolhills ttf Bobby Range. SbO (Z4*3fc7"S, 
151 303'fe), under branch on ground (I, QMMO- 
! "lo.-Dec 14,S3/I.OkIcss)u\ FN km Fof Builyan 
railway station, on Builyan-Gladstone Rd, SEQ 
(24*32'S. 15 T24' E)ram!oresl ne\! torn erhanUitler 
(12. QMMOI0319, IS Jul 1980. J. Stanisic. A. 
Green); Dawes Range, SEQ. MVK Araucaria 
[11 JN'S. kSIWE (47, QMM012642, QM- 
MU12667, AMU36771. AMC136796,4Sepl 1982, 
OM.'OM-ABRS): base, Mt Booroon Booroon. S\V of 
Miriam Vale, SHO, rainforest along creek, under lOgg 
(3. QMMOL6863* QMM016594r 17 Sepl 1985, J, 
Sianisic. D. Potter); Kroombit "fops. c. 13 km W of 
"Chapmans" SEQ (24*25*$, 15 1 U'T) \IVF (12, 
QMMO 12223. QMMGT2224, AMCJ3<. 552. AMC- Sept 1982, AM/QM-ABKN), Ml Fort Wit 
|iam Kalpowar ST., SEQ, NVF (2, QMMQ12597. 
AMC 1 36756. 4 Sept 1982, AM/QM-ABRS); C 4 km 
be low summit Of Ml Fort William, Kalpuwar ST., 
SEO. Wy^'Arauciiria. under laga and rocks (2. 
QMMO1701 1, 3 Jul I9S4. J. Stanisic. D. Potter. K. 
Emberton): Mt Fort William, Kftlpwai S F SCO- 

CNYF, 533 m under IcjgS (2, QMMQ16S22, 3 Jul 
1984. J- Stanisic. D, Potter. K. Emberlon): kroombii 
Tops. SEQ, open fore*! (3, OMM016455. 3(1 Sepl 

I9S5, G«B. Montchhjc Kroambft Tops, SEO. 
MVF7NVF. undcrpalm fronds and logs (30, QMMM 
16517. in VI jv I984i-J, Sianisic, D. Potter). 


Shell large, diameter 4.71-5.97 mm (mean 
5 .38 mm) with 4 MS to 4 3/4 (mean 4 1 2-) tightly 
coiled whorls. Last whorl descending in from, 
Apex and carlv spire (Fig. 54a) almost flat, 
SP/BWW ratio 0,04-0.05 (mean 0.04), u. 
depressed Height of shell 2.52-3.28 mm (mean 
2.84 mm). H/D ratio 0.47-0.65 (mean 053). 
Proioconch of 1 318 to 1 5/8 whorls, mean 
diameter 718. S (am al 1 1,2 whorls. Apical sculp 
ture (Figs 54c, 64a) of curved radial ridges 
crossed by very low spiral cords which are par- 
tially fused and worn, producing 3 pitted ap- 
pearance. Post nuclear sculpture (Fig. 54i') of 
numerous, crowded. protracti\ ely sinuated 
radial ribs, 93-159 (mean 125. N) on the body 
w htort. Ribs mm 6. 1 3-9. 12 (mean 7.42). Micros- 
culpture (Fig. 54f) of fine thread-like radial 
riblets. 7-15 between each pair of major ribs and 
low crowded spiral cprih which buttress the 
microradials on either side. Microspirals con 
tinUOUS on the major radials. Umbilicus (Fig. 
54b) wide V-shaped, diameter 1.45-1.92 mm 
(mean lafW mm). D 1 J ratio 2.76-3.52 (mean 
Sutures deeply impressed. Whorls 
rOUndcd below and shouldered above a weakly 
rounded periphery (Fig. 54c). Aperture broad, 
ovaiely lunate. Lip simple. Parietal callus 
prcseni. Colour variable, light-yellow horn with 
red chevron- 1 ike markings, to dark-brow n 
Based on 1 7 measured adults. 

Epiphallus (Fig. 55a) long, muscular, entering 
penis suhapically. Upper portion of epiphallus 
connected to penial sheath (Fig. 55a) by long 
fibres. Penis (Fig. 55b) long, muscular, internal- 
ly with several longitudinal pilasters, more 
slender and more crowded upieally. Epiphallus 
entering through a pore (Fig. 55b) surrounded by 
a spongy Thickening. Penial retractor muscle 
(Fig, 55a) long, twisted around the cpipha 
inserted on the penis. Vagina long, internalb. 
with fleshy longitudinal pilasters. 

Radula without unusual features. 

Based on two dissected specimens (OMMO- 

R w,r ,\\T)fiAi-;h , i 
6, atrtisiana is confined to the Burnctt-Cal 


Hope-Many Pcaks-Dawcs Ranges, SEQ, with an 

5 outlier population at Mt Larcom. As with other 

* as i ? f s Gyrocochlea, G. curtisiana occurs outside of 

larger rainforest areas and has been collected in 
, remnant rainforest at Builyan. SEQ, and 'open 

5 5 ? forest* at Kroombit Tops. G. curtisiana favours 

||- s si = s warm subtropical notophyil vine forests but may 

be found in adjacent microphyll vine forest (with 

? S2 £ S gif REMARKS 

1 " - ?5 The pitted apical sculpture, almost flat spire. 

rounded body whorls and flammulatcd shell of 

G. curtisiana (Fig. 54a,c,e) are departures from 

the average Gyrocochlea pattern. However geni- 

s st s $ 53 tal anatomy, in particular the form and internal 

characteristics of the penis (Fig. 55b), is typical. 

i is 

5 -r * ;2 ~ 




§3 i 

a = 

S-m 3 — . r* _ » r- 



Biomphalopa gen. nov. 


From the Greek omphalos, meaning um- 
bilicus; referring to biconcave appearance. 

Endodonta recava Medley. 1912. 

Q s 

si tags Previous Studies 

= 1 ^£~ Hedley (1924) introduced Gyrocochlea for 

I charopids with biconcave shell and prominent 

5 o radial sculpture on adult whorls, and included G. 

s ^ concinna Hedley, 1924 and Endodonta recava 

1 SS 1 S SS Hedley. 1912. These species, characterised by 

1 boldly ribbed, tightly coiled biconcave shells on 
g x which the protoconch is macroscopically smooth 

| _ and glossy, are here separated to a new genus. 

g Diagnosis 

2 Shell diameter range 2.64^4.69 mm. with 4 1,8 
* £- to 5 3/8 tightly coiled whorls, last descending 
I |j . 2 5 s s - e more rapidly. Apex and early spire deeply 

** sunken. Apical sculpture u\' regularly arranged 

pits and vague radial ridges. Post nuclear sculp- 

2 ture of high, bold, crowded, protractively 

| x ^ ^ ^ igo sinuated radial ribs. Microsculpture of fine radial 

*5 * * *^ riblets and equally fine spiral cords, continuous 

on the surface of the major radials. Umbilicus 
° | wide and cup-shaped. Sutures becoming deeply 

furrowed as the last whorl descends. Whorls 
strongly shouldered above and rounded below a 
laterally compressed periphery. Aperture acute 
ovately lunate. Colour cream v-v\ hite with darker 
!l!l II |_B| *I |H radial suffusions. 

ilil 11 ^11 Animal with long slender foot and varying 

|3 < u° 5-- degrees of black speckling on the visceral coil. 



Kidney with pericardial lobe long and slender, 
rectal lobe reduced. Male genitalia with epiphal- 
lus shorter than penis, muscular, entering penis 
sub-apically through a simple pore. Penis 
without a sheath, either tubular with a large 
central longitudinal pilaster and a group of 
smaller apical longitudinal pilasters (recava)* or 
with a swollen apical section containing a large 
corrugated verge (concinna). Terminal female 
genitalia with very long vagina and free oviduct. 
Radula with tricuspid central and laterals in 
which mesocone is long, slender lanceolate. 

Distribution and Ecology 

Biomphalopa recava is found in subtropical 
notophyll vine forests of the foothills and 
uplands of the Eungella Range whereas B. con- 
cinna occurs mainly in notophyll vine forests of 
the Cardwell Range and montane vine fern thick- 
ets and vine forests of the Bellenden Ker Range. 
Both species live under logs. 


Biomphalopa is distinguished from Gyro- 
cochlea by its smaller size (Table 11), greater 
number of tightly coiled whorls, bolder radial 
ribs and wider, cup-shaped umbilicus. 
Nautiliropa also has a sunken spire with pitted 
apical sculpture (Fig. 63d) but is larger, has 
fewer, loosely coiled whorls, finer and struc- 
turally different radial ribbing (Fig. 63e) and a 
body whorl which does not descend (Fig. 63c). 
Anatomically the terminal male genitalia of 
Biomphalopa bear some similarities to those of 
Gyrocochlea but the lack of a penis sheath and 
presence of a single longitudinal penial pilaster 
contrast with Gyrocochlea (Fig. 46b). The super- 
ficially similar Nautiliropa has a pear-shaped 
penis with quite complex pilasters (Fig. 65c). 
Biomphalopa can be readily derived from 
Gyrocochlea and common ancestry is probable. 

Biomphalopa recava (Hedley, 1912) 

comb. nov. 

(Figs 57-59; Tables 1 1,13) 

Endodonia recava Hedley, 1912, p. 267, pi. 10, figs 

Gyrocochlea recava (Hedley); Hedley, 1924, p. 217; 
'iredale. 1937a, p. 323. 


B. recava differs from B. concinna in its larger 
size, greater number of whorls, less crowded ribs 

and wider umbilicus. The penis of B, recava has 
a large central pilaster and group of smaller 
apical longitudinal thickenings in contrast to the 
slender central pilaster and large corrugated api- 
cal pilaster of B. concinna. 

Type Material 

holotype:: AMC32992, Finch Hatton, W of Mack- 
ay, MEQ. Scrub. Collected by S.W. Jackson, 2 Sept 
1908. Height of shell 2.68 mm, diameter 4.60 mm, 
H/D ratio 0.58, D/U ratio 2.40, whorls 5 3/8. 

Other Material 

Eungella Range, 50 miles W of Mackay, MEQ (2, 
AMC140230, Sept 1957, L. Price); Finch Hatton N.P., 
W of Mackay, MEQ (21°04'S, I4S n 38'E), 180 m, (2, 
AMC153726, 29 Apr 1975, J.B. Burch, W.F. Ponder, 
P.H. Colman); Digging's Rd, Eungella N.P., MEQ 
(21°09'S, 148°29"E) NVF (10, QMM011757, 
QMMO13073, 8 Jul 1982, J. Stanisic); Palm Walk, 
Eungella N.P.. MEQ (2r00'S, HS^O'E) 800 m, 
CNVF (1, QMM06345, Nov 1976, M.J. Bishop); 
Dalrymple Heights, Eungella N.P., MEQ, NVF, 1 000 
m (2, QMM06367, Nov 1976, M.J. Bishop; 1, 
AMC153727, 28 Apr 1975, J.B. Burch, W.F. Ponder, 
P.H. Colman); Broken River, Eungella N.P., MEQ 
(21°10 , S, 148°30'E) NVF (1, QMM011715, 5 Jul 
1982, J. Stanisic); Finch Hatton Gorge, c. 10 km N of 
Finch Hatton, MEQ (2T04'S, 148°38'E) NVF (1, 
QMMOl 1 749, Jul 19S2, J. Stanisic). 


Shell diameter 4. 1 7-4.69 mm (mean 4.49 mm) 
5+ to 5 3/8 (mean 5 1/4-) very tightly coiled 
whorls, last descending very rapidly. Apex and 
early spire (Fig. 57a) deeply sunken, height 
2.13-2.68 mm (mean 2.43 mm). H/D ratio 0.51- 
0.58 (mean 0.54). Apical sculpture (Fig. 57e) of 
pits and vague radial ridges. Protoconch of 1 5/8 
to 1 3/4+ whorls, mean diameter 671.0 jim at 1 
1/2 whorls. Postnuclear sculpture (Fig. 57f) of 
high, crowded, strongly protractively sinuated 
radial ribs, 81-100 (mean 91.7) ribs on the body 
whorl. Ribs/mm 6.17-7.47 (mean 6.67). Micros- 
culpture (Fig. 57f) of fine radial riblets, 10-25 
between each pair of major ribs and fine spiral 
cords which are continuous on the apices of the 
radial ribs. Umbilicus (Fig. 57b) very wide, cup- 
shaped, diameter 1.87-2.00 mm (mean 1.93 
mm). D/U ratio 2.23-2.40 (mean 2.32). Sutures 
impressed becoming deeply furrowed as the last 
whorl descends. Whorls (Fig. 57c) shouldered 
above and below a laterally compressed 
periphery. Lip simple. Colour creamy-white 
with brown to reddish-brown radial flammula- 






* ! 

^ ^ ; 1%, 

• '< 


.ft, i , 

/ *. 





FIG. 58. Biomphalopa recava (Hedley, 1912). Digging's Rd, Eungella N.P., MEQ. OMM011757. a, 
genitalia; b, penis interior; c, ovotestis; d, hermaphroditic duct; e. ovoteslis: f, lalon and carrefour; g T pallial 
cavity. Scale lines as marked. 

FIG.57, Biomphalopa recava (Hedlev, 1912). a-c. Finch HaitoiuW of Mackay, MEQ. AMC32992, holotype: 
d. h, Palm Walk, Eungella N.P.", MEQ. QMM06346; e-g. Digging's Rd, Eungella N.P., MEQ. 
QMMOl 1757. a-c. entire shell; d, central and lateral teeth: e, apical sculpture; f. post nuclear sculpture; %, 
details of major radial rib; h, marginal teeth. Scale lines as marked. 



FIG. 59. Distribution of Biomphalopa recava (Hed- 
ley, 19 12) in mid-east Queensland. (Pr = Proserpine; 
Ma = Mackay; Sa = Sarina). 

tions. Parietal callus strongly developed. Based 
on 3 measured adults. 

Genitalia with epiphallus (Fig. 58b) entering 
penis subapically through a simple pore (Fig. 
58b). Penial retractor muscle inserting apically 
on penis head. Penis (Fig. 58b) swollen with a 
large central, longitudinal corrugated pilaster 
and shorter longitudinal pilasters apically. Free 
oviduct short, muscularised. Vagina long, inter- 
nally with longitudinal pilasters. 

Radula (Fig. 57d, h) typical. 

Based on 4 dissected specimens (QMMO- 

Range and Habitat 

Under logs in warm subtropical notophyll vine 
forests of the Eungella area (southern extension 
of the Clarke Range) and Finch Hatton Gorge, 

MEQ. It has not been found in the drier rain- 
forests of Ml Dryander, near Proserpine, MEQ. 


B. recava is distinguished by a biconcave shell 
with prominent radial ribs and tight coiling. 
'Gyrocochlea' iuloidea (Forbes, 1851), from 
Port Molle (=Airlie Beach), has a depressed spire 
and wide umbilicus but is larger with more loose- 
ly coiled whorls, apical sculpture of spirals and 
radials, broad V-shaped umbilicus, more 
rounded periphery, more prominent microspirals 
on the body whorl and monochrome, usually 
dark-brown, shell 

Biomphalopa concinna (Hedley, 1924) 
comb. nov. (Figs 60-62; Tables 11, 13) 

Gyrocochlea concinna Hedley, 1924, p. 215, pi. 29, 
' fig 1-3; Iredale, 1937a, p. 322. 

See under B. recava. 

Type Material 

HolotypE: AMC8902, Cardwell, NEQ. Height of 
shell 1.58 mm, diameter 2.64 mm, H/D ratio 0.60. 
D/U ratio 2.57. whorls 4 1/8. 

Other Material 

Kirrama Range, via Kennedy. NEQ, 500 m, rainforest 
(1, QMM017322, 2 Oct 1980, G.B. Monteith); Ken- 
nedy S.F.. NEQ (18°LVS, 145°46'E), 900 m (1, 
QMM06417, Aug 1977. M.J. Bishop); Dunn Creek, 
on Kirrama S.F. road, Cardwell Range, NEQ, NVF, 
(1, AMC142965, 27 Sept 1980. I. Loch); Mt Bellen- 
den Ker, NEQ, just below summit, c. 1500 m, under 
bark on rotting log, MVFT (2, QMM014878, 4 Jul 
1983, J. Stanisic, D. Potter); Mt Bellenden Ker, NEQ 
(base, 17, QMMOl 1297; 500 m, 2, QMM011326; 
1054 m. 5, QMMOl 1343; summit. 5, QMMOl 1255; 
Earthwatch/QM, 1981); Mt Bartle Frere, NEQ, walk- 
ing track below NW peak on western side, c. 1000 m, 
SNVF(2, QMM011381.8 Nov 1981, Earthwatch/- 


Shell diameter 2.64-3.45 mm (mean 3.08 
mm), 4 1/8 to 5 1/8 (mean 4 5/8+) tightly coiled 

FIG. 60. Biomphalopa concinna (Hedlev. 1924). a-c, Kennedy S.F., NEQ. QMM06417; e-g, Mt Bellenden 
Ker, NEQ. QMM0I4878, QMMOl 1297. QMM011255; d.'h, Cardwell. NEQ. AMCS902, holotype. a-c. 
d, h. entire shells; e, apicalsculpture; f, post nuclear sculpture; g, central and lateral teeth. Scale lines as marked. 







FIG. 61. Biomphalopa concinna (Hcdley. 1924). Ml. Bellenden Ker. NEO. QMMOl 1255. a. genitalia; b, 
penis interior; c. hermaphroditic duct; d. talon; e, interior of free oviduct; f. g. ovoleslis; h. pal Hal cavitv. Scale 
lines as marked 







■ l 






FIG. 62. Distribution of Biamphalopa concinna 
(Hedley. 1924) in north-east Queensland. (Ca = 
Cairns: In = Tngham; Tu =Tully). 

whorls, last whorl descending more rapidly. 
Apex and early spire (Fig. 60a) strongly 
depressed, height 1.58-1.87 mm (mean 1.71 
mm) H'D ratio 0. 53-0. 60 (mean 0.56). 
Protoconch glossy, 1 5/8 to 1 7/8 whorls, mean 
diametcr600.5 umat 1 1/2 whorls. Apical sculp- 
ture (Fig. 60e) of radially arranged rows of pits. 
Post nuclear sculpture (Fig. 60f) of crowded, 
high, protractively sinuated radial ribs, 80-95 
(mean 84.8) ribs on the body whorl. Ribs/mm 
7,84-9.63 (mean 8.82). Microsculpturc (Fig. 
50f) of fine radial riblets, 6-20 between each pair 
of major ribs and fine crowded spiral cords. 
continuous on the apices of the radial ribs. Um- 
bilicus very wide, cup-shaped, diameter 1.03- 
1.27 mm (mean 1.19 mm), D/U ratio 2.28-2.80 
(mean 2.58). Sutures impressed, deep on the last 
whorl. Whorls rounded below and shouldered 
above a strongly compressed periphery (Fig. 
60c). Lip simple, columella not expanded. Parie- 
tal callus strongly developed. Colour creamy- 
white to light-brown, with darker brown radial 
suffusions. Based on 4 measured adults. 

Genitalia with large, muscular epiphallus 
which reflexes before entering the penis subapi- 
cally (Fig. 61a). Penial retractor muscle inserted 
apically on the penis. Penis (Fig. 61b) with a 
large apical bulb, otherwise tubular, internally 

with an apical, fleshy swollen pilaster (Fig. 61b) 
adjacent to the simple cpiphallic entrance. Penis 
proper with a single, thin, almost lamellate, lon- 
gitudinal pilaster (Fig. 61b) and a few smaller 
thread-like longitudinal ridges, otherwise 
smooth. Free oviduct a short muscular tube. 
Vagina long, internally with very prominent cor- 
rugated longitudinal pilasters. 

Radula (Fig. 60g) typical. 

Based on two dissected specimens (QM- 
MO 11255). 

RANor and Habitat 

Under logs and under bark on rotting logs in 
the Kirrama Range, NE o\ Ingham, and the Bcl- 
lenden Ker Range, SE of Cairns, NEQ; it prefers 
montane notophyll vine forests. 


The verge in the penis indicates the possibility 
of sympatric relatives. 

Nauliliropa gen. nov 


Referring to the nautiloid-Hkc appearance -l 
the shell. 

Type species 
Helix omicron Pfe iffcr, ] 85 1 . 


Moderately large size. c. 4 1/2 tightly coiled 
whorls, the last inflated and not descending. 
Apex and early spire (Fig. 63a) strongly 
depressed. Apical sculpture \yi low radial ribs 
and low spiral cords, modified to produce a 
pattern of regularly arranged pits (Fig. 63d). Post 
nuclear sculpture (Fig. 63c) of numerous, 
protractively sinuated radial ribs formed by the 
periostracal extension of a single underlying 
radial thickening. Microsculpturc of crowded 
microspirals and microradials. Umbilicus wide, 
I -shaped. Sutures impressed, whorls rounded 
below and shouldered above a rounded 
periphery (Fig. 64f). Aperture ovately lunate, lip 

Animal with long, slender foot, weakly 
speckled with darker pigmentation on the lower 
spire. Pallial cavity elongate with complete sig- 
murethrous ureter. Kidney with long pericardial 
lobe and vestigial rectal lobe, Epiphallus (Fig. 
65b) not bound to the penial sheath, entering 
penis apicaliv through a Fleshy pilaster. Penis 
(Fig. 65c) pear-shaped with a muscular collar 



and a complex pilaster pattern. Pcnia! bulb con- 
taining cpiphallic entrance and horseshoe- 
shaped pilaster. Lower peniai chamber with api- 
cal stimulator (Fig. 65c) and longitudinal thick- 
enings. Terminal female genitalia short. Radula 
[Fig. 63c,f) with hook-like tricuspid central, 
bicuspid claw-like laterals without ectocone, and 
bicuspid marginals with long basal shafts and 
short endo- and ectocones. Teeth rows in B V 


\\mitlirofui omicron is removed from Gyro 
cochlrti becjuse of basic differences in majorrib 
StrUCtUtiG and coiling pattern- Gxroi ochleu has 
major ribs composed of several small ribs, and a 
final whorl which descends (Fig. 45c). whereas 
Naulitiropa has a last whorl which coils more 
tightly without descending, and major ribs com- 
posed of a single rib with a large pcnoslracal 
blade l Tig. 63c). Although the modified apical 
sculpture oi NautiUropa is also present in 
(iymoh hli'itCLirttsiana (Hedlcy, 1912) this fea- 
ture has evolved independently in several 
lineages of Charopidae. 

Viuliliropu omicron (Pteiffer, 1S5I) 

comb- nov. 
(Figs r.;.. Md-f, 65, 665 Tables 11.14) 

HckiomtcumPkiUa. 183*1. p, 128; I854.p.457. pi. 

!5xligs M-l7;Cox, IShSp.lS, pi. H), lig. I . 
Helix nmmuititonics Reeve. 1S54, pi. IS'l. sp. I24fi 
■:!>'<! 0M3CfOf\ (Fielder)- Medley. Isl74 t p. 

216. pi. 30, fig* 16-18; Iredale. 19374, p. 323: 

hedrfe, r5WS«up.2G& 


JV. vmlcran is most liable to be confused with 
species of Gyrocochlea which are sympatric 

with it in eastern parts of the McPherson Ranges, 
Q I he Ilammulated shell is good for field 
identification. However, rare monochrome 
brown specimens of N. omtcmn need to be dis- 
tinguished bv the more crowded nuiiul ribs, 
smaller size, pitted apical sculpture and non-des- 
cending body whorl. (7. cutttstdtut has a ilammu- 
lated shell with crowded radial ribs (Fig. Ma) but 
lacks the sirongly depiessed spue ol ,\. omicron. 

Previous Studies 

Early records of Nautilimpa omicron (Pteii- 

fer, 1 851) cited by Cox (1868), Medley and Mus- 

son (1892) and Hedley (1924), included records 

yrac$c)\Iea curtisiana which is restricted to 

a small area from south of Miriam Vale to Mt 
LarcoiTL SEQ. Attempts to locate the type of A r . 
emu ron have been unsuccessful. Most probably 
il was housed in the Stettin Museum and 
destroyed during the World War II. The initial 
description and subsequent figures (Pfeiffer. 
1S54) leave no doubts as to the identity of the 

i vn Math w 

NprrryPF: QMMO27290 (ex OMMOlOSOl) Mt 

Warning N.P., NSW (2tf24 s. ^ r.), topofroaU 
in rainforest. Height 3.03 mm. diameter 5.71 mm, 
WO raiifl D.53. D/U ratio 3'iS. tt bofls 4 .VS-. 

Mini k MATERIA! 

\V side ct Somerset Dam, NW pf Brisbane. N r E of 
Esk. SEQ. open scteropbvll on hillside (2, 
AMC53175, 6 Sepi 1970, w.f .' Ponder .v P. Marsh); 
NW 0l Brisbane- SLO. l.aeeys ( 'k |UM S ul Raynbtrd 
'. k open Ur\ sderoprn II and ianhma along creek. 

" L4 & 152*43 E)(ZAMCi37792,QMM012976, 
JO Ane I IJ S2, AM QM'ABRSl; c 15 km from 
Legume on Acacia Plateau Rvl., Koreelah Ck S.T., NE 
NSW BOG m. NYI MVF (Ml. AMC136S24, 
OMMO 12721 S Dee 1081, AM UM - ABRSJ; 
■ lOuUKlL Park. SEQ{25 a S7 > S, 153 (fVE) SNVF 
tan sand (7. AMC 1 3641(7, AMC136490, 
OMM012111.0MMO121]3.t)Sep[ 1W. AM/QM 

ABRS); Coolomong Sanctuary, upper Ourrumbin 
Vallev W of Tweed Hds, SEQ, raintna-M reserve, litter 
{I. AMC137778, 30 Aug 1982. AM/QM - ABRS): 
Macpherson Ra. SEQ (1, AMC13977S, lower, ex 
Gqx); Mi Tatnbanne, SEQ (2. AMC63795, J- Sim- 
mondsi: Brisbane. SEO(2. AMC 139767, ex Cox); off 
trib. lo Back Ck.. Unumgar S.F., NE NSW (2& B 26'S, 
152 42*E)(1, AMC128584. 15 Mar 1981, AM/QM - 
ABRS); Ml Nebo. W uf Brisbane, SEO (1, 
AMC14043G, 23 Mar I943, G.P. Whitley); Natural 
Bridge N.P.. SEO, MVF CS'l-'S, t53*l4'6) (X, 
AMC129269, AMCI2927U QMMO10457, 
QMMOHI4(>0, IS M.,r Nsi, AM/QM - ABRS); Mt 
Tambumie. SEQ. (I, AMC!4U43y, J.A. Stmmomls); 
Nth P. ne River, SEQ, in scrubs ( ID, AMO03605, ex 
Cuv); I.iitle Yahba Ck., Imbil S.F., SEQ, NVF 
I £6*28*8. t52 - 38 , E)(2 ( AMCl36448.QMMCH204I, 
7 Sept IV82, AM CM ABRS); sidetrack off Mt 
Archer KdMtMceS.F. SEQ, 

NVVfAraucflria t27°04'S. 152MTE) (9, 
ANfCl36523.QMMOl2ISJ.31 Sepi )982,AM/QM 
- ABRS); Diiililab*h, Bunya Mis N.P.. SEQ, behind 
picnip jrea Ch'M s. ISJ^fi'EJ (JB3 AMCI366Q7, 
QMMOJ2283, ^l Sept 14S1 AM.QM- ABRS);Mur- 
gviilfltnbflh, ^SW (J, AMC139765, Lower, ex Cox); 
Ml I intlesay. Upper RtChmOTld R NSW (2, 




y i 




U < 


S- £ E 



c 4 

I 3t 




ij-i e-i 



^cc C^ -^ 




ffi c 




a 5 

= 2 



c -r: 



s a 

• -■ 










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£ 3* 





' M ' f*' 

c'A! #:■ r ir xi 


FIG. 63. Nauiiliropa omicron (Pfeiffer, 1851). a-b. Mi Warning N.P., NSW. QMMO27290. neolvpe; c, f. 
Ml Glorious, SEQ. QMM016127; d-€, Kenilworth S.F., SEQ."QMM09688. a-b 3 entire shell; c, marginal 
teeth; d. apical sculpture; e, post nuclear sculpture; f, central and lateral teeth. Scale lines as marked. 












FIG. 64. a-c, Gyrocochlea curtisiana (Hedley, 1912). a, Kroombit Tops, SEQ. QMMO 16453; b-c, Springs 
Scurb, Mt Larcom, SEQ. AMC32990, holotype. a, terminal part of protoconch; b, c, entire shell, d-f, 
Nautiliropaomicron (Pfeiffer, 1851). d-e, Mt Glorious, SEQ. QMM016127; f, Dandabah, Bunya Mts, SEQ, 
QMM012283. d-f, entire shell. Scale lines as marked. 




FIG. 65. Nautiliropa omicron(? feiffer, 1851). a-g,h,Mt. Glorious, SEQ. QMM016127. f, Dandabah,Bunya 
Mts, SEQ. QMM012283. a, genitalia; b, details of penial complex; c, penis interior; d, spermatophore; e, 
hermaphroditic duct; f, talon and carrefour; g, ovotestis; h, pallial cavity. Scale lines as marked. 



AMC139761, 27 Apr 1958, L. Price); Norlh Pinr R , 
SEQ (2. AMC139752, Lower- ex Cfcx); faced R. 
NSW C6. AMC139768, Peiierd. ex C ox). CafflSAfra, 
S6Q (1 . AMG»S<J. SAV. Jackson); Pine R., SCO (2. 
AMC264, 1587, Mu550tl)i (Jootnburra S,R, Mistake 
MtSSEQ(27'59'S, 152*21 E)(6,AMa28G19,QM« 
MO 111 22, 16 Mar 1981, AM OM - ABRS); c 
km from Gnnmburra, Goom hurra S.F.. SEQ e>00 m. 
t nvi- Patms<2T59 1 S l52"2rE)(l7 3 AMC|36SR 
QMM012&92. QMMOI2rVJS. 7 Dec I9SL AM QM 

- *BRSj 

Comlamine R Valley, C 9 km NE of Killarne\, 

nsw. along strtam (28M8 s. i>: 2£'£) (3. 

AMCI2SW7.QMMOHU15. IftMar l^.Sl.QM.QM 

- ABKS). MontvlUt k± neat Kondalilla Falls N.P. 
SEO. vin« scrub on roadside (2h l 40\S. IS2*52E^5, 

\MC 15(vH4. QMM01l'>s?..SS q >i ios:. ,\M QM 

ABRS): Clarence R., NSW (3, AMC 139758. ex Pel- 
u-ui). c 3,5 km NW Mi Pleasant, on Dftyboro - Mi 

.ml Rd StQ. dr\ and wet sderoplnil<27 OS'S. 

| ,!■;-, QMMO 12)24. 30 Scpi 
l9tS2,AM/gM-ABRS);s:. 1 km S Kjxnhirtl Ck. Rd., 
or l aeevx Ck. Rd.. Laccys Ck. SEQ. open forest 
(27 UV 152'43'E) (10, AMC13M7h. QMMO- 
nioii; AMCI364E2, OMMOI2UMI. SO Sept 1982, 
AM/QM - ABRS); Benaikin S> , neai BlacM [fell 
I (B \M< 139757 Teh |96f, D.F. McMichoei); 
Dandahah. Bunya Mts N.P.. SEQ. behind picnic area 
■ WS. I5]'30"E)(4. \MC13oo00.QMMOi: 

AM QM- ABRS); Clarence R.. NSW 

AMCI39766, ex Cox); Tooloom Scrub, Beaury 

: c. S.3 km from Woodenbong. Legume Rd.. NSW, 

MYT cs:o s, 152 B 24 > E),(1,AMC1285?4 ! 15 Mar 

1981, AM.QM - ABRSt; Cunninghams Gap N.P.. 

SEQ. near monument. MVI p8"W S 152 14 a E) (6, 

\MCI2y.tiS. OMMOHWi. I ft Mai Wfll, AM QM 

ABRSt; S\V ot Oympie, SEQ, NE slopes Ol Mt. 

MoorodTeerai, remnant Ihtckel amongst hoop pine 

intaliaii £6*l6'S, 152*33 I I I \MC\ SS1Q5, 

4O133HQMM013356, 7SejM I&82, AM QM 

- ABKS). S of Gympie, SEQ enlrancc to ST. of 
> abba Ck Rd near Dwver Ck., renmattl vine thicket 
ifong gully <2<OS'S, 152 3S'F) ( i. AMC13K1II. 

AMCI3M i:. QMMOI3367, QMMOI QtiQ, 7 Scpl 

1 AM OM - ABRS); Richmond R.. NSW (I. 

AMCi3''7'vi. L90Q.-S.WJackson)iMi Warning N P 

NSW. (op of road in rainl'oresl <?S 24'S. 133 !7'i) 
I", AMC129304, (.)MM() lii-oii |Q M;, r losl 
AM QM - ABRS); Bhekhul! Range, c. 7.4 km S ol 
BetiarkirtSEQ,MVFMAtfiaw/a(2G 53'S. 152 II'E) 
107, AM( "13foM, AMCU6571, OMMOI223S. 
QMM012245. 31 Sepi 1981, AM/QM - ABKS). 
0'R«Hlv*S I-.minoton N.P '- I I | i 2 H N"S, 
~ IS Lno.nMMOIu4.-w P Mar I'JS) AM 
ABRS)- Ml Warning N.P.. NSW. tuar Ivi-.c of Mt 

(28 "4 s, 153 *WE) <2 QMMOi048& Mat l$8l, 
AM OM - ABRS): ToonumruT Forcsl Rd , Tbominr- 

aai ST .. e 34 km !rom K\oHl \|, NSW, CN\T 

(2S"3.VS. 153 i 'i H I.MMMOloui;:. |4Mar I" 

AM/QM ABRS); Hon I bS F ,NE ?JSW, era Acacia 

Plaieau Rd. C, I-.- km Irom Acacia Ck - Kilkr 
Rd NSW smi m (28"2rS. 152 24 I:; 
SEVT MVP/A rtiucarla \ ?, QMMO 10V4S 
AM( t 18561, 13 Mai 1981, AM/QM - ABRS); Ml 
GU.riouv c. 3 km SI, al Mimmii SUO (27*20"S 
'- : 4f\ E) (3. OMMOI 1991. QMMO] V 
AMCI S6521 - Dei 1982 am QM ABRS). 
Pomona SI.. SI ol Pomona. SEQ (26*2 
IS2 541 Ml OMMOI2U(I^. SSrpi I0S2_ AM'OM- 
ABRS 1982): Mi C.uvta N.P.. SEQ (25 ft 49'$, 
L52 : _ I iMVr\\T.|/r;//( (1 ™ ^OMMDPiC- 
7SCpl I «C, AM/QM ABRS). 
Byron Ck crossing. Byron S.F., c\ S t km MW of Mt 
Pleasant SEQ(27 fc 07 k S, 1 52 -l. ; 1 If'.OMMOl ' ! 
AMCl3ASflfl. 311 Sepi 1982 AM.OM - ABRS) 
I mi in ii l 1 'i, i i i 'Mp N.P. StQ "f monument 
'>4 : S. 152 34> i i I v QMM012677. 
AVK 136806. 7 Dft i 11 - M ' IM - ABKS); Mt 
WarnineN.i 1 .- NSW ( ;s 24'S, k5? >fi L) C NAT- ( I ( 
QMM<112X^ lyOrc losi.AMtjM \BRs). s.a 
Synipic, SEQ. eitirancc it- Efrooloo S.F. on Gil 
Rtiad, MVF [2fi - >0 V 152 4t ]0"E) ( I, 
QMMO] : - Sj pi 1982. AM/QM -ABRSfcUpp 

Booloomha Ck. C , 3/i. SJ SL(.) (2f.4l S, 

15: 37 E)(l-OMMOl3386.8Sepi 1 982. AM QM 
ABRS): Boomb.ota N.P Ml Nebo, SEO, (li 
QMMO6071. 14 MM B'7c. M.J. Bishop); Dandahah, 

Bunya Wis N P SI Q I t7 S3 s. 151*33 I I 

\TttUCaria NVI , Ol, MMMOoOS:, 5 Mar I^7h, M..I. 

BisImi ii Binna Bi n I gco i N.P.. SEQ, 85 n 

CNM (Z ijMM(ioiis3, Apr 1976, MJ. Bish.-pi: 
MantTiua N.f'.. Ml GluWtuift. St O, ^etseleroph) 11 1 1. 
QMM06263. Any 1976, M I Bishop); Ml Bynffl 
s.f ..sho.;- mi s. 152 30 E^2.QMM062?7 
1976, H J. Bishop) V«i ciorimis, SEQ (27*20^, 
152 W] id QMMOh293, I :6 ■" fl M J Bishop); 

Mai i ■ ! Pa >■- i jMtMumr, SEQ. NVF >. ; 

QMMO6306. U Aug 1576, MJ. Bishop); Bo.miixtna 

n p m, webo si q ^1 sclcrophyll I ! 

1:"J-' 1 Kl.UMMO(i344. -■ &. MJ. Htshopl. 

Fred's Rd, Mi.Mcc SEQ (27 o-vs |s7 4V(,)in. 
OMNKr.vlS, I I Apr l'-'KO. J. Slanisic. N. Hall. A 
Green}: Kenilwonb SF. SLtj i"f< 4u'S. I5J - 
Amucaria.{7+iMMWi Ia> I9HQ.J.SUU1 

a Green): Maiale N r. Mi Glorious SEQ. CBVI 

(2Z OMM01lr t f)4. OMMOI 1S47. 2£1 Jun 1^S2. 
MSA p.Mui, SpjCCl '•<Kip,S[-0(l,UMM(" 1-8 ■■: 
Aug i'-':v'. A. Rdxereld*). 

Mi Ncbo si q nr loll> ' ■> I nakoul NVF (I. 
QMM012771. 2 Dec UK2. J. SunUc); K L Bilwttrili 



25 ? S 






■ Mnryborough 

v 0&lt. > 300m BLisrore 

151°E 152°E 



FKi. 06. [Jislribulion OfNauritiropa nmicron (Pfetf- 
ler, IK5I). 

S.F SEQ,C ().3kmNofLillle Yabba Ck on Mulenv 
tti keruUnrih Rd. SEO (26'37'S. 152^42'EJ (3, 
OMMOI.UV,. iQMav 1483, K. Collin*. J. Stanisic): 
Ml Glorious, SEQ, NVF (6. QMMOI4K.S >; Nov 
!" J Mumsie); Boombana N.P., Mi Nebo, SEO, 
NVF (| QMMO15Q60, 5 Apr 1984, J. Sttmsi I 
MdeUttCk Off Ml Archer Rd, Ml Me« S.F., SF.Q 
(27 W U4"S, 15^41 , E)(m 3 QMMOl5074.6Sepl ''■' : 
J. Stannic F). Potier); Fred's Rd. Mt Mec, SFO, NVF 
(27 OS'S, !52'43'^lt2.QMM015985 l 28 0ct (984, 
i.smiiimv, G, Annabel!); Ml Glorious. SEO.. NVF (3. 
UMMOl598^280ct 1084, J Stanisic. G. Annabel I), 
Ml OluriOUS, SEQ. NVF (IK QMMOJM27, 2f. Jan 
198ft. J Stannic. J. Chaselinea 
Ml Tamborlna, SEQ. Curtis Falls Circuit. CNVF, 1 1 
QMM016437, 23 Feb ls>Ho, J. Stanisic, J. Chasehng): 
Upper Pine Ck,. Canungra, SEQ, NVfjArftticaHa, 1 1 . 
UMMOI6547. I Ocl 1986 I, Sianisic. D. Potter); 
Redwood Park, Toowoomba. SEQ, NVF (7, 
QMM01&572, in Sept 1966, J. Sfcnfs* i. fcfffft Iffo 
Track. Binna Hurra, l.aminglon N.F.. SEQ. NVF (2. 
OMM01657K. OMMOHoHii. %1 Sept 1986, J. 
si 1 1 1 i s i c > ; TullawaHal Circuit. Binrta Butro 
UndiAglOll N P.. SEO. NVF(7, QMMG1673S. 2 0ct 
l 1 - 1 M I Stanisic, D. Poller, J. Chaselme). Naluial 
UrnUieN P.. SEQ. on roadside e. 3011 m N of entrance. 
SNVF (5, QMMOIOXIS, 24 Mar L984, X. Stanisic); 
Blackball Ranee. SEQ., on • >i- B UCG Highway 

\t\ I- \wiHwrm\3\ 0MMO1G Zfi : 2 May I9B4,J, 

SLanisic. D. Potter). Mt GlOOOUS SI 0. CNVF (4, 
(jMVlf JlfiSV, 5 Apr i i ;S4, .F Stani-ac); Dandabah, 
Bunja Mis N.P.. SEQ. NVt (14, OMMO10S47 12 
Ma) 1984 I- Stnnisic. Potter): Big Falls Circuit, 
Bunya Mis N.P.. SEO. SNVF i i'J, QMMG16850. 5 
Nov |*J85 T I Stanisic D. Poller): Cherry Plain - 
YVeslcoll Plain Circuit, Bunya Mts N.P.. SFO. 
NVF/MVF |lb, 0MMOlhS3'\ fi Nov 1985, E 
Stanisic. 0. Potter); Mi Hobwee Circuit. Binna Burra. 
i .nt.i.i ■■.-.onN P. SEQ NVF (2. QMM016S65. ) Oct 
[985, I Suuusic. 0. Poller, J. Clpseltng); Border 
I tack, Binna Burra, Lamingion N.P-, SEQ, NVF (8, 
QMMOloS'O, 10 Mar 1984. J. Stanisic). 
Tullauallal Circuit, nr antarctic beeches, Binna 
Burra. I amington N.P.. SEQ. NVF ( I. QMM016S74. 
I I Mar I984< J.Stanisic); Border track, nr O'Reillys. 
Lammgton N.P.. SEQ. NVF (7. QMM016876, 31 
M.n |994 T j, Stanisic); initial part of Kweebani Caves 
Walk. Binna Burra. Lumingiun N.P.. SFO- NVF (4, 
QMM016885. ^ Mar [984, J. Sianisic); Kenikvorlh 
S.F.. SEQ (26 a 37'S, 1.52*42 '£) { I, OMMO 18530, 22 
May 1 9S0. J. Sianisic. A. Gieen): Mudgeeraba, SEQ, 
(3, QMMOIS53I, C..1. Wild); Cunningham's Gap. 
SEO. (4. QMMOKS532. OMMOI-SS34. ISJun 1979, 
vitnahel!); Mt Who. c 2 km east of summit, SEQ. 

(152*47 F. 2 ' ~. J "S). HVF/Arflttcatla, ( i . 

QMM01H533, 13 Aug PWU. Stanisic); Koondai-1 
Lookout. Bunva Mis N. P.. SEO (26: 53'S. 1SI°33T.) 
{I OMM01B595, 2 Mar 1976, MJ. Bishop): Mary 
Can-across Reserve. WalCH] SEO. ' *-" "46'S, 
152*51*6)* (I QMMOIS54V). 22 Mav I9K0. .1. 
Sianisic. A. Green): c. 1,6 km N Ol Nummbah valley 
tumofl '"i Burleigh lo Springbrook Rd, SEQ, NVF 
Willi palms (I. QMMOI8550, H Dec 1980, J. 
Stamsk ] 


Shell large, diameter 4,20-5-97 mm (mean 

5.24 mm) with 4 to 4 58+ (mean 4 3/8+) tightly 
coiled whorls, the last very inflated and not des- 
cending in front (Fig. 641"). Apex and early spire 
(Figs o3a. 64d) stmnglv depressed, height 2.35- 
3. II mm (mean 2.lS2 mm). FED ratio 0.48-0 57 
(mean 0,54), Protoconch o( i 3.8 to 1 5/8 whorls 
(mean I 3 N+) with mean diameter 597.5 p.m at 
I I 2 whorls. Apical sculplure (Fig. 63d) of low, 
curved radial ridges and low spiral cords which 
are partially fused and worn producinga strongly 
pitted appearance. Post nuclear sculpture (Fig, 
63e) Of numerous, crowded, thin, productively 
sinuated radial ribs. 72-167 (mean i 13.6) ribs on 
the body whorl. Ribs mm 5,03-9.79 (mean 
6 90) Ribs with a high periostracal blade, often 
lost in dead or slightly worn specimens, Micros- 



culpiurc (Fig. roc) strongly reticulate, with fine 
microradials. 6— )5 between each pair of m&ji i 
ribs and lower, crowded spiral cords which but- 
tress the microradials on either side- Umbilicus 
1 1 ig- 63b, 64e) wide, U-shaped, diameter 1.23- 
L64 mm (mean 1.47 mm). D/U ratio 2.02-4.22 
mi ,m 3.58). Sutures impressed, whorls rounded 
below and shouldered above a rounded 
Liinhen (Fig. o4f). Last whorl CO i led more 
tightly Aperture ovately lunate only slightly in- 
clined from the shell axis. Lip simple, columella 
not expanded but slightly deflected toward um- 
bilicus. Parietal callus well de\ eloped. ColQUl 
light-horn to light-brown, often with darker 
radial flammulations arranged in a ^ig-zag pat- 
tern. Based on 31 measured adul 

Genitalia with cpiphallus (Fig. 65b) mus- 
eularised. looped around the perna! retractor 
muscle before entering penis apically. Fpiphallie 
pore (Fig. 65c) between two swollen pilasters. 
Penis i Fig, 65C?) pear-shaped with a distinct api- 
cal bulb, separated from the lower penial cham- 
ber by ;tconstrictingmuscular collar. Penial bulb 
internally with apical to slighth lateral epiphal- 
lic entrance and horseshoe-shaped pilaster. 
Lower penial chamber with s large apical pocket 
stimulator (Fig. 65c), and anastomosing spongy 

Penial retractor muscle short, inserting on 
penis apex adjacent to epiphallus. Vagina long 
internally with fleshy longitudinal thickem. 
Spermatophore, simple, clavaie. 

Radula as for genus. 

Based on five dissected adults (QMMO 16 1 27. 
QM MO 122X3). 

■ wnH tfil 
Highlands of the Great Dividing Range be- 
tween Tooloom, NF. NSW and the Bunya Mis. 
SEQ; the northern end of the Richmond Range, 
the McPherson Ranges south To the Tweed 
River, in the east: and the coastal and adjacent 
ranges north to Ml Guyra, SEQ. N. omirmn 
shows preference tor notophvll vine forests but 
also occurs in adjacent microphyl] vine forests. 
wet sclerophyll forests, and in gallery rainforest 
along watercourses, ft owicron lives under logs 


,Y, nmicmn is one k)\' the most conspicuous. 
widespread and frequently encountered of the 
subtropical mierosnails. However most o\ the 
637 Specimens available were SUhaduil reflect* 
ing the high proportion of mid-year, litter- 
eblfccted material. 

The modified radula ol \ omicron is unparal- 
leled in Australian eharopids. Radulae of Ihe 
souths -. eharopids Mufathena forde.i 
(Bra/ier 1871 J, Stcnucapha hamtltoni (<'<i\ 
I868J and Thrya^onti diemcnc/isis [Cox, 1868) 
have typicai central and lateral teeth (Smith and 
Kershaw, 1985) but marginals show modifica- 
tions broadly similar to those in V. omicron. \1. 
fordei and S, hamiltoni have strongly slanted, 
slender, unieuspid marginals while T. cliemt-twn 
sis has bicuspid marginals. In If. omicron the 
chance from average charopid patterns may be 
the result of character displacement under con 
dilions o\' microsympatn with the eonchulogi- 
calls similar Gyrocochlcu- 

Specimens from eastern localities, in par- 
ticular those from the D'Aguilar Range, SEQ, 
have more crowded ribs than those from the 
Great Dividing Range [Table 14). These dif- 
ferences may be related to local environmental 


Letomuhi Iredale. 1941 

iciomola Irtdalt. 1941a, p 2fi7. 
LelomUla {ftritit) KerfihaM L956h, p. £. 

TVP€ SPEj it - 

fth&phad&n ca/UatiuS lledley. 1924 in 
original designation. 

PkFVKH sSTl D ' 

Iredale (194 1a) introduced Letomola as 
monotypic. Smith and Kershaw ( 1979) included 
the Tasmanian species He!t.\' barrenensis Pel- 
terd. 1879 without giving reasons and it is here 

Small shell (mean diameter 2.05 mm), with 
few (mean 3 7/8-) tightly coiled whorls. Spire 
f Fig. 67a) depressed. Apical sculpture (Fig. 67dl 
oi' irregular malleations and ridgelets. 
Postnuclear sculpture of weak growth ridges and 
vague spiral cords. Umbilicus (Fig- 67b) very 
wide, cup-shaped (mean D/LT ratio 2.68). Sutures 
Impressed Whorfc rounded below the peripberv, 
flattened at the basal margin with a strong 
suprapenpherai sulcus above (Fig, 67c). Aper- 
ture ovately lunate with one parietal and one 
basal barrier (Fig. 6Sa. bj. Lip simple, margins 
convergent. Colour brown w ith lighter streaks. 
Animal with foot and tail broad, bluntly rounded 
posteriorly. I Ivc animal dark-grey, particularly 
i in ihe sides o! ihe fnof above the pedal grooves, 


neck and mantle collar. Pallial cavity elongate 
with bilobed kidney, pericardial lobe reflexed. 
Genitalia with unusual features. Ovotestis a 
single, bilobed clump of follicles (Fig. 69c, d) 
embedded in the digestive gland. Terminal 
genitalia elongate. Epiphallus (Fig. 69a) a long 
slender tube, entering penis apically adjacent to 
an apical pilaster. Penis (Fig. 69b) slender with 
longitudinal pilasters. Penial retractor muscle 
inserted at the penis/epiphallus junction. Radula 
(Fig. 68d-f) narrow with few lateral and marginal 
teeth rows; central tooth very small, mesocone 
reduced; laterals tricuspid with much enlarged 
mesocone; marginals becoming multicuspid, 
serrate, with all cusps reduced in size. 


L. contortus is conchologically distinctive 
with wide umbilicus (Fig. 67b), large, finely 
malleate protoconch (Fig. 67d), reduced adult 
sculpture (Fig. 67e, f), strong sulcus on the body 
whorl (Fig. 67c), and two apertural barriers (Fig. 
68a, b). It is sympatric with Rhophodon 
kempseyensis (Fig. 77) which has a wide um- 
bilicus and brown shell with white streaks but 
differs in having a radially lirate protoconch, 
extremely crowded ribs on the body whorl (mean 
ribs/mm 33.34), and more numerous apertural 

Letomola contortus (Hedley, 1924) 
(Figs 67-70; Tables 11, 14) 

Rhophodon contortus Hedley, 1924, p. 220, pi. 32, 

figs 35-37; Kershaw, 1955, p. 30. 
Letomola contortus (Hedley); lredale, 1941b, p. 2; 

Iredale, 1941a, fig. 6. 
Lctomala (sic) contortus (Hedley}; Kershaw, 1956b, 

p. 9. 

Type Material 

Lectotype: AMC28494, Sherwood, Macleay River 
NSW. e\ Technological Museum (= collected by C. 
Laseron) Height of shell 0.80 mm , diameter 1.81 mm, 
H/D ratio 0.44. D/U ratio 2.84, whorls 3 5/8-. 

P,\R\LECTOTYPE: AMC150095, same data as lec- 

Other Material 

Yessabah Cave, W of Kempsey, NSW (31 WS, 
I52°42'E), on roof and walls of limestone cave and in 
floor litter (200+, AMC1 19349, 9 Oct 1979, P.H. 
Colman, J. Stanisic); Yessabah Cave, via Kempsey, 
NSW (31°05"30"S. 152°4rE) vine thicket, on lime- 

stone rocks and in litter (54, QMMO17014. 3 Mar 
1987, J. Stanisic, D. Potter, P.H. Colman). 


Shell glossy, minute, diameter 1.81-2.24 mm 
(mean 2.05 mm) with 3 1/2 to 4+ (mean 3 7/8-) 
tightly coiled whorls. Apex and spire depressed 
with each whorl higher than the preceding whorl, 
except for the body whorl which descends in 
front. Height 0.80-1.03 (mean 0.91 mm). H/D 
ratio 0.40-0.5 1 (mean 0.45). Protoconch of 1 3/8 
to 1 1/2- whorls, large, bulbous. Apical sculpture 
(Fig. 67d) of irregular malleations and ridges. 
Post nuclear sculpture (Fig. 67e) of broad radial 
ridges crossed by weak spiral cords, especially 
noticeable near the sutures. No additional 
microsculpture. Umbilicus (Fig. 67b) wide, shal- 
low, cup-shaped, diameter 0.64-0.86 mm (mean 
0.77 mm). D/U ratio 2.42-3.03 (mean 2.68). 
Sutures impressed, becoming a deep furrow as 
the last whorl descends. Whorls rounded below 
the periphery with a prominent supra-peripheral 
sulcus. (Fig. 67c). Aperture irregular ovately 
lunate with one parietal and one basal barrier. 
Basal barrier more deep-seated than parietal. 
Parietal barrier (Fig. 68a) extending 1/8 whorl, 
horizontal anteriorly, becoming angled 
downward posteriorly. Basal barrier (Fig. 68b) 
1/8 whorl in length, following angle of coiling, 
tapered at either end. Lip simple, columella only 
slightly dilated, margins convergent. Parietal 
callus weakly developed. Colour deep reddish- 
brown with irregular white radial streaks, brown 
on the base. Based on 13 measured adults. 

Epiphallus-vas deferens junction valvular. 
Epiphallus (Fig. 69b) entering penis apically. 
Penial retractor muscle inserted on the epiphal- 
lus-penis junction. Penis (Fig. 69b) a long tube 
with a slightly expanded apex. Epiphallic 
entrance a simple pore adjacent to a short 
tongue-like pilaster. Penis proper sculptured 
with longitudinal pilasters. Free oviduct and 
vagina long tubes without unusual features. 

Radular characteristics as given in generic 

Based on 6 dissected specimens (AMC- 
119349, QMMO 170 14). 

Range and Habitat 

From the limestone outcrop at Yessabah, west 
of Kempsey, NSW where the vegetation is semi- 
evergreen vine thicket. In spite of several collect- 
ing efforts it has not been found further westward 
in the limestones of the Carrai State Forest. L. 





FIG. 67. Letomola conforms (Uc6\ey, 1924). a-c, Sherwood, Macleay R, NSW. AMC119349. a-c, entire 
shell; d, apical sculpture; e, post nuclear sculpture; f, sculpture on base. Scale lines as marked. 






FIG. 68. Utomola conlortus (Hedley, 1924). Yessabah Cave, W of Kempsey. NSW. AMC1 19349, 
QMMOl 7014. a, parietal barrier: b, basal barrier; c, microdentieulatinn on basal barrier: d, radula; e, central 
and lateral teeth; f, marginal teeth. Scale lines as marked. 






FIG. 69. Letomola contortus (Hedley, 1924). Yessabah Cave, W of Kempsey, NSW. AMC1 19349. a, 
genitalia; b ? penis interior; c, d 7 details of ovotestis; e, hermaphroditic duct; f, talon and carrefour; g, pallial 
cavitv. Scales lines as marked. 




■ ■ ■ 


FIG. 70, 

DiMribulion al f.ctomotu cenionus (Hedley, 

comonus has been collected live on limestone 
rocks alter rain and presumably feeds on the 
microflora growing on ihe rock surface. 


L amtortus displays a number of departures 
from typical charopid patterns. The malleated 
protoconch (Fig, 67d) and suprapcnphcral sul- 
cus (Fig. 67c) arc unusual features in the context 
of trie cast-coast species reviewed herein. Sulci 
are seen in some Pacific Island Endodontidac 
and Charopidac, and according !o Solum (1983. 
p. 9) they arc usually associated with keel forma- 
tion. In /.. conforms the sulcus indirectly con- 
stricts the size of the aperture. Anatomically the 
slender reproductive organs are more typical oF 
the Punctidae. However, the radii la is typically 
charopid even though the strong reduction in size 
of the central tooth and serrate appearance of the 
marginals are unusual. Omphaloropa vuricosu 
has a grossly similar radula - feu teeth per row 
and reduced central tooth. 

Rhophodon Ilcdley. 1924 

Rhvpkoifoti ttedteV, 1924, p. &Jft Ircdaie. {937a, p^ 
32<>; Ircdaie. 1941b, p. 2; Kershaw. 1955, p. 30; 
Hurch, I976„p.133. 

lodmiA l^dale, l<?37a, p. 328; Kershaw, 1956a, 
p 142' Bureh, !976_ p .132. 

Typf SpfcIfs 

Rhophodon peregnmts Hedley. 
original designation. 

924; bv 

Previous Stuqh s 

Heillcv ( 1924, p. 219) introduced Rhophodon 
for charopids with apertural barriers. Apertural 
barriers among Australian Charopidac are rela- 
tively rare but several generic groupings have 
been proposed including Deniherona Ircdaie. 
1933 (type species: Helix dispar Brazier, 1671); 
Bischoffena Ircdaie, 1937 (type species: Helix 
bisrhoffensis Pctierd, J 879); Egilodonia Iredale. 
1937 (type species: Charopa baimsdalcnsis 
Gabriel/1930); and Letonwla Iredale 194 t (type 
species: Rhophodon conforms Medley, 1924). 
Rhophodon is expanded to include seven 
species of very small to minute charopids with 
apertural barriers and shells which have a 
predominantly radially lirate protoconch. post- 
nuclear sculpture o\' prominent radial ribs, and a 
wide saucer-shaped umbilicus. 


Very small to minute Charopidac. diameter 
range" 1.29-2-98 mm. Whorls 4 1/4 to 5 1/2-, 
tightly to very tightly coiled, last descending 
more rapidly. Apex and early spire depressed 
{kanpscvcnsis, pcrcgrinus) to very slightly 
elevated Protoconch exsert of 1 1/2 to I 3/4 
whorls. Apical sculpture of numerous 
moderately crowded to very crowded (more so 
toward the protoconch - adult whorl boundary) 
weakly curved radial ribs and very weak thread- 
like spiral cords and cordlets. Post-nuclear 
sculpture ol lew. widelv spuced {rolmani, 
h.innsdalcnsis) to many, crowded {kempsewn- 
\/\), thin, protractively sinuated to straight radial 
ribs, Microsculpture of numerous, crowded 
i adial I iblets, and widely spaced (kcmpseyensis) 
to crowded, low rounded spiral cords which are 
raised at their intersection with the radial riblets 
to form an elongate bead. Umbilicus wide, cup- 
shaped. Aperture ovatclv lunate, with few 
{htiirnsdalensis) to many {consohrinus) barriers. 
Parietal barriers consisting ol one to four 
(generally three) creseentic lamellae, with or 
without additional thread-like traces. Palatal 
barriers three to six (usually four) creseentic 
lamellae, with ot without a supeuor trace 
Columellar lamellae sometimes present. Pallia] 
region elongate with almost umlobed kidney in 
which the pericardial lobe is well developed, 
reflexcd at the tip, and the rectal lobe is reduced 
to a vestige abutting the hindgut. 



Ovotcstis two small clumps Of finger-like 
lobes embedded in the subapieal whorls gf the 
digestive gland. Terminal male genitalia with 
cpiphallus entering the penis apicalty or subapi- 
eally (eonsobrinus) through twp fleshy lips sur- 
rounded hy ;i muscular collar. Penis with an 
apical bulb containing epiphallic pore, some- 
times expanded and with longitudinal pilasters 
(eonsobrinus)- Penis with (peregrmus, con- 
sohrtrttts, kempseyensis) or without a sheath, m 
lcrnally with small, fleshy, longitudinal pilasters 
and basal, circular, stimulatory pads (con- 
v.o/j/v>7//s- pcregrinus). Penial retractor muscle 
Inserting at the penis/cpiphallus junction. 
I em;ile reproductive system elongate with long 
\ agifla and free oviduct. Raduia without unusual 
features except in kempseyensis. 


Rhophodon is iound from Bairnsdale, Vic- 
toria, to the Bulhurin State Forest, near Miriam 
Vale, SBQ. Species have very small ranges 
Some are known only from closed forest but 
others live in fringe rainforest areata and lime- 
stone refugia. R. pcregrinus and R, consohntms 
are symputne in the eastern MePherson Ranges, 
SFQ. and some adjacent areas including the 
coastal vine thicket at Burleigh Heads SEQ 
Differences in penial morphology between the 
fwo species are probably species recognition 
features. Species live in a variety of 
microhabitats and have been collected from 
kaves in litter, under logs, and from moss and 
Inur accummulated on rock surfaces. R 
0(*rdgrtnU& has also been found in an Ay/lcntum 
fern on a tree. 

pattfrnso! sheli Variation 

Size and Shape - Rhophodon ineluues st ime i if 
the smallest charopids found in eastern Australia 
with mean shell diameters 1.45-2.66 mm. The 
largest Spftcifift, R. ke/np$cvcnsi.\. is confined to 
the limestone outcrop between Yessabah and 
Carrai, west of Kcmpscy. NSW. where the readi- 
ly aVaiUbk calcium probably enables greater 
size Otherwise size variation appears to be 
mosaic in nature. 

Height is correlated with diameter (mean H/D 
ratio for the seven species ranges 0,40-0 .46) and 
is reflected in conservative shell shape. Spire 
protrusion ts minimal. Body whorl width shows 
some reduction in R. rolrnani and R. mttmus- 
simus correlating With smaH size. Mean whorl 
counts range from 4 1/2- to 5-. Species with the 
largest whotl counts are R. kempsey crisis: R. 

batrn.sdalensis trum limestone habitat in 
southern NSW; and R, eUzabethae from calcium 
rich aiaueanan \ me forests ,H SEQ, 

SCulplurB - Shell sculpture is complex but 
variation is relatively simple, The most sig- 
nificant differences are in the number and spac- 
ing of U\c radk] ribs and il is possible to divide 
Ihe species into three groups on this basis - few. 
bold, widely spaced ribs {R. Nomsdulensis. R., Figs 87a, 89a); few. narrower, 
moderate]', spaced ribs (R. mmuli^imus> R 
i'!;zobL';:>o.-. Figs S I a, K4a ); and very many. fine, 
extremely crowded ribs (R. peregrmus, R. con- 
sobnnus, R kempseyensis. Figs 71a, 74a, 77a). 
Diilercneesin microsculpturc are minor with the 
more wioci\ spaced microspiral cords of R. 
bempseyenxis [Fig. 77c) being an extreme 
modification which may relate to the larger 
whorl size of that species- Minor differences 
occur in number of apical ribs and intensity of 
apical cording. 

Apcnitral Barriers - Variation in the number 
of parietal barriers ranges bom a single elongate 
barrier (Fig, 90a) to six harriers and traces (Fig. 
7lg). The average pattern is for three elongate 
harriers, expanded posteriorly and deflected 

The average pattern oi palatal barriers is Eat 
four major barriers to which a superior trace nia; 
be added. Major deviations are a single barrier 
and trace (Fig. V()b, c). Ihree barriers (Fig K4h) 
and six barriers (Fig. 87a). In general fta bar- 
riers are short crescent-shaped lamellae. In R. 
colmani (Fig. K7g) the barriers arc expanded 
with grcatlY elongate anterior Hades 

A colutneJlar barrier ma; be present and is 
usually short crescem-shHped and may be sup- 
plemented with an accessory trace (Fjg, 74 

Patterns of AKatGMICAI .Variation 

Stability of the Rhophodon morphotypc, 
which is evident in the COflChological patterns. 
extends hi Ihe analomv Apart liom differences 
related to species size e.g. relative lengths of the 
genital and pallial structures, the most notable 
variation occurs in the male terminal genitalia. 

Mole genttuhu - Typically the penis hiss an 
apical bulb which contains the epiphallic 
entrance and a main penin! chamber with few to 
several fusiform, longitudinal, fleshy pilasters. 
1 -xceplions arc R eonsobrinus (Fig 75e) and/?. 
peregrinus [Pig. 72b) which have altered penial 
surfaces, These, two species arc sympatnc and 
the differences in penial morphology are probab- 
ly species recognition leatures. The presence of 



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a penis sheath correlates with large size and 
greater intensity of radial ribbing. The epiphallus 
is normally long and muscular but in R. kemp- 
seyettsis (Fig. 79a) it is very long, correlating 
with increased whorl count. 

Radula - The Rhophodon radula shows little 
interspecific variation reflecting overall 
similarity in species microhabitats. Size dif- 
ference between the central tooth and laterals is 
exaggerated in R. kempseyensis (Fig. 78e). The 
large broad lanceolate mesocone of the laterals 
is a feature seen in other charopids (Figs 68e, 
I 2 3 h ) sympatric with R. kempseyensis and ap- 
pears to be correlated with life on limestone. 


Rhopiwdon differs from the conchologically 
similar Egilomen Iredale, 1937 in having aper- 
tural barriers and in details of microsculpture. 
The microspiral cords in Egilomen are more 
prominent than those of Rhopiwdon and are 
crossed b\ fine thread-like radials contrasting 
with the high microradial ribs of Rhopiwdon. 
Anatomically the pallial configurations of 
Rhophodon and Egilomen are distinct. The 
strongly bilobed condition of the Egilomen kid- 
ney contrasts with the almost unilobed kidney of 
Rhophodon. Although the penial complexes Of 
the tw r o genera are grossly similar, the tubular 
penis and stout muscular epiphallus of Egilomen 
are important differences from Rhophodon. 


*• I- Nji<-«7 r. ""- r-i ~ ■- — 

1 .Ribs on body whorl very widely spaced (Figs 

87a<89a)^i, ' . ,.2 

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2, Aperture with three swollen parietal barriers 
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Aperture With a single lamellate parietal bar- 
rier (Fig. 90a) baimsdalensis 

3. Major radial ribs hold, moderately spaced 4 

Major radial ribs finer, very crowded 5 

4 r Shet| vcrv SnTftll (mean diameter 1 .52 mm); 
second parietal barrier much shorter than first 
and ihird mintnis.simus 

Shell larger (mean diameier 2.47 mm). flam- 
mulaied: second parietal barrier elongate 





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FIG. 72. Rhophodon peregrinus Hedley, 1924. On border track c. 6km from O'Reillys, Lamington N.P.. SEQ. 
QM MO 16595. a. genitalia; b. penis interior; c. ovoteslis; d. hermaphroditic duct: e. talon and carrefour region: 
f, pallia] cavity. Scale lines as marked. 

FIG, 71. Rhophodon peregrinus Hedlev , 1924. a-c, Tweed River. NSW. AMC153719. neotvpe; d, c. 6km 
from O'Reilly's on border track. Lam'ington N.P.. SEQ. QMMOI6595; e-h. Tweed River. NSW. AMC- 
95838. paralype. a-c T entire shell; d. marginal teeth; e. apical sculpture; f, post nuclear sculpture; g, parietal 
barrier; h. palatal barriers. Scale lines as marked. 



with '»n anterior taper; Bobby Range. SEQ 


5. Major radial ribs extremely crowded (mean 
ribs/mm 33,34); shell larger (mean diameter 
2.66 mm) kmpstipensfs 

Major radial ribs less crowded (mean 
ribs/mm < 30.00); shell smaller (mean 
diameter<2.40mm) 2 

6. Parietal barriers consisting of four lamellae and 
iwo [races (Fig. 71g); three palatal barriers 
(Fig. 71 h). percgrinus 

Parietal barriers consisting of three lamellae. 
and rarely, two traces (Fig. 74h); five palatal 
lamellae (Fig. 74!) consobrinus 


2 9 r - ; .r 

L < srcre 

, c^Jalt. > 500m 




i l°E 

Rhophodon peregrinus Hcdlcv, 1924 
(Figs 71-73; Tables 15, 16) 

Rhophodon percgrinus Hedley. 1924, p. 220, pi. 32. 
figs 38-40; Iredale. 1937a,"p, 329; Iredale, 1941a, 
p.268, fig. 6; Iredale. 1941b 7 p. 2. 

FIG, 73. Distribution of Rhophodon per cgrinus Hed- 
ley, 1 Q 24. 

River, NSW. Collected W. Pellerd, ex Cox Coll. 
Heightof shell 0.S6 mm, diameter 1.99mm. H/D ratio 
D/U ratio 3.43. whorls 4 7, K-, 


R. percgrinus is conchologically similar to R. 
consobrinus but is distinguished by the more 
crowded ribs and slightly depressed spire and 
greater number of parietal barriers. The two 
species are sympatric and have quite different 
penial morphology. These peniai surface dif- 
ferences are effective species recognition fea- 

R. ke?npseyensis is larger with relatively 
greater number of whorls and almost twice as 
many ribs on the body whorl (Fig. 77a). Other 
Rhophodon species have markedly fewer ribs on 
the body whorl than R. percgrinus (Table 15). 

Previous Studies 

Although Hedley (1924) referred to a type 
specimen collected by W. Petterd in the Cox 
collection, the only "type" material of R. 
percgrinus located in the Australian Museum 
v. as a lot of 38 specimens (AMC95838) from the 
type locality, labelled paralectotypes although 
this designation has no basis in literature. Also 
in the Australian Museum type collection was a 
vial labelled "Rhophodon percgrinus ... Type 
lost ...". 

Type Material 
Neotype AMCI53719 (ex AMC9583N). Tweed 

Other Materia! 

AMC958?& 37 specimens, same collection data as 
neotvpe; Burleigh Heads N. P.. SEQ, vine thicket, litter 
(1. QMM014U6, 6 Jan 19S4, J. Stanisic; 25, QM- 
M06165, Oct L97G, MJ. Bishop); c, 6 km from 
O'Reilly's, on border track, l.amington N.P., SEQ. 
NVF, on rocks (7, GM MO 16595, 31 Mar I9S4, J 
Stanisic); Tooloom Scrub. Beaury S.F., NE NSW 
(28"29 S. IS2°24'E) NVF (2. QMMO10938. 15 Mar 
1WSI, AM QM - ABRS); Burleigh Heads (5, AM- 
C55506. ex J. Brazier); Byron Bay scrub. NSW (2, 
AMC63S47, Lower, ex Cox); Beaurv S.R, c. 15 km 
W of Urbenville. NSW Sin m (2S^21'S. \5T2VE) 
(27, in litter and in A spknium on tree, AMC152160, 
AMCI52T61, 18 May 1976, P.H. Colman, I. Loch); 
Kureelah S.F.. c. 1 2.2 km from Acacia Ck - Killarnev 
Rd, NSW SEVT'MVF (2S^21S. 152'24E) (4, QM- 
MOI0956, AMC12S554, 15 Mar 1981, AM/QM - 
ABRS); Richmond Rv. Lismore, NSW, in scrub under 
decayed leaves and dead limber (3, AMC63S72, ex 


Shell minute, diameter 1,79-2.22 mm (mean 
1.93 mm) with 4 3/8 lo 5 1/4+ (mean 4 5/8+) 
tightly coiled whorls, the last descending more 
rapidly. Apex and early spire depressed to 
strongly depressed. Height of shell 0.74-0.95 
mm (mean 0.85 mm). H/D ratio 0.41-0,47 


(mean 0.43). Protoconch exsert, 1 1/2 to 1 3/4 
whorls, mean diameter 4 1 0.9 ja.m at 1 1 12 whorls. 
Apical sculpture (Fig. 71e) of curved, broad 
radial ribs and weak spiral wrinkles, ribs becom- 
ing more crowded toward the nuclear/post 
nuclear junction. Postnuclear sculpture (Fig. 
71f) of numerous, very crowded, high, slightly 
protractively sinuated radial ribs. 112-189 
(mean 156.9) ribs on the body whorl. Ribs/mm 
18.85-30.44 (mean 25.93). Apices of ribs with 
spiral sculpture. Microsculpture (Fig. 7 If) of 
fine radial riblets and conspicuous low spiral 
cords which are raised at their junction with the 
radial riblets producing a beaded effect. Um- 
bilicus (Fie:. 71b) wide, cup-shaped, diameter 
0.74-0.90 mm (mean 0.82 mm). D/U ratio 2.22- 
2.54 (mean 2.36). Sutures impressed; whorls 
shouldered above and rounded below a weakly 
rounded periphery. Aperture ovatcly lunate. 
Numerous barriers present. Parietal barriers 
(Pig- 71g) consisting of four crescent-shaped 
lamellae and two thread-like traces. First barrier 
large, deflected upwards; second, third and 
fourth without marked anterior taper, directed 
downwards. Superior and inferior traces may be 
present. Palatal barriers consisting of three cres- 
cent shaped lamellae (Fig. 71h) and a superior 
trace (Fig. 71g) high up on the palatal margin. 
Columellar barrier small, thread-like. Lip 
simple, columella not expanded or reflected. 
Parietal callus developed. Colour white to 
straw-yellow with brown radial streaks and suf- 
fusions. Based on 16 measured adults. 

Genitalia with vas deferens/cpiphallus junc- 
tion (Fig. 72a) complex, valvular. Epiphallus 
(Fig. 72a) a muscular tube continuing to the 
penis head where its coils before entering the 
penis apically. Penis (Fig. 72a) an almost 
cylindrical, muscularised tube with a thin sheath 
and head slightly expanded, tapering as it nears 
the atrium, internally with a simple epiphallic 
entrance (Fig. 71b) surrounded by a muscular 
collar. Interior of the penis with large, apical, 
longitudinal pilaster, a basal doughnut-shaped 
pilaster, and general wall sculpture of spongy 

Radula (Fig. 71d) typical. 

Based on 5 dissected adults (QMM0 16595). 

Range and Habitat 

Cool subtropical notophyll vine forests of the 
McPherson Ranges, SEQ, westward to the Great 
Dividing Range on the NSW/QLD border; coas- 
tal rainforests at Byron Bay, NSW and Burleigh 
Heads, SEQ. An earlv record at Lismore, NSW, 

needs to be confirmed. R. peregrinus has been 
collected in litter, among moss on rocks, among 
accumulated debris, and unusually, in an 
epiphytic fern growing on a tree. 


Lack of material makes a study of interpopula- 
tional variation impossible at this stage. How- 
ever. a single specimen (Table 16) from the 
Koreelah State Forest, NSW, in the western part 
of its range, differs in rib spacing from speci- 
mens further east. 

Rhophodon consobrinus Hcdley, 1924 
(Figs 74-76; Tables 15, 16) 

Rhophodon consobrinus Hedley, 1924, p. 220, pi. 31, 
figs 32-34; Iredale. 1937a. p. 329; Iredale, 1941b, 
p. 2. 


R. consobrinus can be distinguished from R. 
peregrinus by its modified penis which has the 
apical bulb reflcxed and expanded to include 
longitudinal pilasters (Fig. 75c). Conchological 
differences are minor except for the disposition 
of aperlural barriers. R. consobrinus (Fig. 74h) 
has fewer parietal barriers than R. peregrinus 
(Fig 7 1 g). R. kempseyensis (Fig. 77a), has al- 
most twice as many ribs on the body whorl, is 
larger and has a higher whorl count than R. 
peregrinus. Other species can be differentiated 
by their smaller rib counts (Table 15). 

Type Material 

LECTOTYPE: AMC 19922, Richmond River. NSW. 
Presented W.F. Petterd. Height 0.95 mm., diameter 
2.18 mm., H/D ratio 9.43. D/U ratio 2.59, whorls 4 

Othfr Material. 

Burleigh Heads N.P., SEQ, vine thicket (7, 
QMMO 16581, 29 Sept 1 986, J. Stanisic, J. Chaseling; 
2, QMMO6160, Nov 1976, M.J. Bishop; under logs, 
7. QMMO 16922, 21 Jan 1987, J. Slanisic, D. Potter): 
Upper Pine Creek. Canungra, SEQ. NVF/Araucaria 
(on rocks, 20. QMMO 16548. 1 Oct 1986, J. Stanisic, 
D. Potter; on moss covered rock, 7, QMMO 1 6906, 21 
Jan 1987, J. Stanisic, D. Potter; under logs, 7, QM- 
M016586, 29 Sept 1986, J. Chaseling, J. Stanisic); 
Binna Burra, Lamington N.P., SEQ. (28 D 12'S, 
153*1 TE) NVF/Palms (1. QMMO 10446, 18 Mar 
1981, AM/QM - ABRS): Red Scrub Flora Reserve, 
Whian Whian S.F., NSW, 210 m (28°38 4 S, 153 19*E) 
(1. AMC152157, 15 Mav 1976. P.H. Colman, I. 



I mm 

1 * if 




WIU ■ 




FIG. 75. Rhophodon consobrinus Hedlev. 1924. a-e. g. h. Upper Pine Ck. Canunnra. SEQ. QMMO 16548, 
OMM0 16586, QMMO 16906, f, Burleigh Heads N.P., SEQ. QMMO 16581. a. genitalia; b, vas deferens - 
epiphallus junction; c. details of penis - epiphallus junction; d. ovoteslis; e, penis interior; f. spermalophore; 
g. talon, hermaphroditic duct and carrefour region; h. pallial cavity. Scale lines as marked. 

FIG. 74. Rhophodon consobrinus Hedlev, 1924. a-c. Richmond River, NSW. AMC19922. lectotvpe; d-e, 
h-i. Upper Pine Creek, Canungra. SEQ. QMMO16906, QMM016548; f-g. Natural Bridge N.P., SEQ. 
QMMO10455. a-c, entire shell; d, central and lateral teeth; e. marginal teeth; F, apical sculpture; g, post 
nuclear sculpture; h. parietal barriers; i. palatal harriers. Scale lines as marked. 



." E 


alt. > 500m 
N\\vocv\ LV^i l 


^^ z. 

1 I 

FIO. 76. Distribution Q\ RhOptwifon VOf\ pbtiHW 
Hedley. I92r4 

r.och); Rocky Ck., Red Scrub Flora Reseo e. NSW 
t :s 3S s, 153 I • I ..lintoreM (5, OMViOKi 1 ' 

\M( CSSM. AM QM - ABRSj: Tyagarah (1, AM* 
C15215S. Lower, ex Coxji ITwced River, NSW (3 
AM( 5310, ex Cox )■ Cape Bryon. NSW M 
AMCI4(M,S0, 1 May r95S, L, Price): uppci h 
River, NSW (I, AMCI52I59, Ptiitrd. ex Coxl. 
Natural BndgeN.P..NSW.NVH.:.X'I.VS. I53U F) 
^QMMO10455,AMC!29266, IS Mar I9KI. AM/QM 
- ABRS): Kichmoiul Rv. Iismorc, NSW, m &CTUl 
under decayed leaves and dead timba f. I . AM Co ij 

- . Bffl [fcf)« 

DlrSGN i 

Shell minute, diameter 1,95—2.43 mm (ftreatt 

2.35 mm) with 4 3/*+ to 5 ITS- (mean 4 3/4-) 

tightly coiled whorls, the last descending I 

rapidly. Early spire and apex weakly to 
moderately depressed (Fig. 74a). Height of shell 
0.82-1 .07 mm (mean 0.94 mm), H/Dralio 0.40- 
0.44 (mean 11.42). Protoconch exsert. of I J/2 to 
I 5/8 whorls, mean diameter 457.0 ( xm at 1 12 
whorls Apical sculpture (Fig. 740 of curved, 
broad radial ribs, becoming more crowded 
toward the nuelear/postnuelcar shell junction, 
and weak spiral wrinkles. Postnuclear sculpture 
(Fig. 7_Sg) at numerous, crowded, broad, slightly 
protectively sinuate radial ribs ? 109-1 So (mean 
143.4) ribs on the body whorl- Ribs/mm 16.93- 
24. S3 (mean 20.30). Apices of ribs rounded and 
5i ulpturcd with radial lines. Microsculpture 
( Fig. 74g)of fine radial riblets. 5-9 between each 
pair ot major ribs, and very low spiral thicken- 

ings, which are raised at their junction with the 
radial riblets to form elongate beads. Umbilicus 
i Fig, 74b) wide, cup-shaped. diametcr().76-1.07 
mm (mean 0.88mm'). D/U ratio 2.23-2. 82 (mean 
2.5b). Sutures impressed. Whorls (Fig. 74c) 
rounded below and shouldered above a slightly 
compressed periphery. Aperture ovately lunate. 
Numerous barriers present, extending posterior- 
ly for about 18 whorl. Parietal barriers (Fig. 
74b) three, long, crescent-shaped lamellae, and 
several, variable, thread-like traces. First and 
second parietals large, crescent-shaped, second 
dc fleeted downwards; third low. directed 
downwards. Parietal traces may occur above the 
first parietal lamella and between the first and 
second parietal. Palatal barriers (Fig. 74c) five. 
Ihc fifth crescent-shaped almost touching the 
first parietal lourth. almost equal in size to the 
fifth, also crescent shaped, third smaller, crCS- 
ccnt-shaped. sometimes bifid apically; second 
if) ■; first basal, thin, small. A superior palatal 
trace may he present at the panelo-palalal mar- 
gin (Fie 74h) Colnmcllar lamella small, Lip 
simple, columella not expanded or deflected. 
Parietal callus developed.. Colour light slraw- 
vetlow with reddish-brown radial streaks and 
suffusions. Based on S measured adults 

dcniialia with penial retractor muscle inserted 
at cpiphallus.'pcnis junction (Fig. 75a). Penis 
(Fig. 75c) luhuiar. re flexed c. 1,3 way along its 
length, with a thin sheath, and complex internal 
sculpture. Fpiphallus entering penis through 
two thickened lips (Fig. 75c ) adjacent to several 
high longitudinal pilasters. Remainder of penis 
chamber wall with broad low pustules giving 
way to low thin ridges basally. Base of the lower 
mber with a stimulatory doughnut-like pad. 
Spermatophore (Fig. ^^t) hook-shaped. 

Radula (Fig. 74d,c) similar to that of R. 

Based on 13 dissected specimens (QMMO- 
[6548, OMMO105S1. QMMO16906, OMMO- 
J6922>QMMO 16586?). 

Rangi- .\\n Habit vi 

Moist, warm lowland notophyll vine forests of 
the Big Scrub area, northern NSW and eastern 
McPherson Ranges and the coastal vine scrubs 
at Byron Bay, NSW. and Burleigh Heads, SEQ. 
R, coHSohrimn has been collected in litter, under 
logs and on moss covered rocks. 


Although R. pen\^nnu\ and R. consobrinus 
have not been found microsympatrically. their 


overlapping ranges and convergent microhabitat 
preferences make this a distinct possibility. Cer- 
tainly, penial differences suggest imcrosym- 


Rhophodon kempsevensissp. uov 
(Fig. 77-80; Tables 15. 16) 


For the Kempsey region. 

r ompxrisons 

This species is recognizable by its lustrous 
golden brown shell which has numerous, ex- 
tremely crowded radial ribs. It has the highest rib 
count of any east Australian ehuropid. The 
species closest to R. kempseyensis are R. 
peiL'tfrirtus and R. ccmsobrinus but they have 
onlyc. huirihe numberofribson the body whorl 
an smaller, and have relatively fewer whorls 
(Table 15). iciomola contort us which is sym- 
patic with R. kempseyensis, has ;t wide um- 
bilicus and two apertutal barriers- one basal and 
one parietal (Fig- 68a. b). Sculpturally /.. con- 
tortus differs from R. kcmpcvaisis in having a 
more bulbous protoconch (Fig. 67d) with ic- 
regulai malleations and t idges, and adult whorls 
(Fig. 67e, f) with low, broad radial ridges and 
weak spiral cords* 

HoinrM'i-: QMMOI70U,, Natural Arch. C;irr;ii 

s.t.,NS\v. 30 a 59*S, 1 52*21' E. limestone onmoss) 

rocks. Collected by J. Stanisic. D. Poller. P.H. Col 
man. 3 Mm L$87, Height of shell ' 13mm, diameter 

2.77. H/D ratio 0.41, D/tJ raiifl 2.37, whorls 5 I .'"S+ 

Paratyphi QMMO1701S. 1 06 specimens, same 

collection d.ita as holotvpe: Limestone c;ives al 
NaMltal Arcti. W. of KempttV, NSW (30"51 
i52':i'h) (150+, AMC153720, 10 Mar iWi. 
W. Ponder. O Griffiths); Haydonville, \V. ol 
Kempsey. S. side ol 'Natural Arch*, under rock over- 
ling (4 specimens. AMC152156, 20 Jan 1982. e.v 
A.B. Rose. M. Dudkin); Yessabuh CaVe, W ol 
Kempsey. in leal and soil litter (31^06'S, 152^42 L) 
(5(1+ specimens. AMC1 1 9351 < 9 Ocl I979.1.5uni»c, 

P II Colman). Natural Arch. Carrai Ca\cs. httei (70-f 

specimens, AMCl52155,2?Feb I980\P u. Oilman). 


Shell minute, diameter 2.47-2. OK mm (mean 
:.oo mm) with 4 3/4- to 5 12- (mean 5) lightly 
coiled whorls. Spire and apex depressed. List 

Whorl descending more rapidly, Height 1,03- 

1.17 mm (mean 1,07 mm), H/D ratio 0,38-0.43 

mm (mean 0.40 mm). Protoconch of 1 1/2 to I 
5 s whorls exsert, mean diameter 440.8 |j.m at I 
12 whorls. Apical sculpture (Fig. 77d) of 
curved radial riblcts. more crowded al nuclear- 
posi nuclear boundary, and fine spiral wrinkles. 
Posl-nuclear sculpture (Fie. 77c) of numerous, 
crowded, protectively sinuated radial ribs, 23(V- 
330 (mean 2^7.7) ribs on the bodv whor 
Ribs/mm 28,63-38,19 (mean 33,34). Micros 
culpture (Fig. 7 7e)of crenulate radial riblcts, 2- J 
between each pair of major ribs and very low, 
widck spaced, rather inconspicuous spiral 
cord scd at their junction with the 

mieroradials in produce a weak beaded effeci 
continuous on the apices of the radial Tibs. Um- 
bilicus (Fig. 77b) wide, cup-shaped* diameter 
1 .03- 1 .2 L » mm ( mean 1.14 mm ). D U rat io 2.2 1 - 
2.44 mm (mean 2.33 until. Sutures impressed. 
Whorls rounded above and below the periphery 
(Fig. 77c), shouldered al the upper margin, flat- 
tened below . Aperture ovotely lunate with seven 

barriers. Pour palatal knnersli \g 78a), lourih 
and third large creseent-shaped; second smaller. 
crcsceni-.hiped at the basal matgin; I irsi simil.r 
in size to second, situated at the baso-colunv l 
margin Three parietal Earners (Fig, 78b). First 
and second elongate crescent-shaped lamellae: 
third a low elongate blade. Lip simple, columella 
no! thickeneo, nlmosl vertical, not reflected 
Parietal callus present. Colour light straw-yel- 
low with broad reddish-brown radial streaks , 
suffusions. Based on 1 1 measured adults. 

Genitalia with Vas dcfeieus epiphallus jun: 
ion valvular, Epiphatlus (Fig. 79a) long, thick 
muscular, coiled; internally with longitudinal 
thickenings, c uiering penis apically through 
thickened pilasters (Fit* 79b). Penis (Fig. 79b) 
a long, muscular tube with u thin sheath, inter- 
nally with a primary longitudinal pilaster, a 
medial, spherical thickening and secondare 
smooth longitudinal thickenings. Penial retrac- 
tor muscle shorl, originating from the diaphragm 
and inserting on the epiphallus just prior to i 
entry into the penis. Spermatophore (Fig. 19f) 
hook-sbap : 

Radula (Fig. 78c, f) with very long lanceolate 
mesoconc on laiernK 

Based on three dissected adults (GMMO- 

Ranoi and Habitat 

LimestOne outcrops between Vessabah arid 
Carrai. west of Kempsey. NSW in evergreen 
vine thicket derived lioin the surrounding wan-, 
temperate rainforests. Five specimens have 





e 'A- 

1 mm 

FIG. 77. Rhophodon kempseyensis sp. nov. a-c, Natural Arch. Carrai S.F., NSW. QMMO17016, holotype; 
d-f, same data holotype. QMMO170I5, AMC 153720, paraty pes. a-c. entire shell; d, apical sculpture; e, post 
nuclear sculpture; f, base. Scale lines as marked. 




FIG. 78. Rhophodon kempseyensis sp. nov. Natural Arch, Carrai S.F., NSW. QMMO17015, paratype. a, 
palatal barriers; b. parietal barriers; c. microdenliculation on surface of third palatal barrier; d, radula; e. central 
and lateral teeth; f, marginal teeth. Scale lines as marked. 








FIG. 79. Rhophodon kempseyensis sp. nov. Natural Arch, Canai S.F., NSW. QMMO17015, paratype. a, 
genitalia; b, penis interior; c, hermaphroditic duct; d. ovotestis; e, talon and carrefour; f, spermatophore; g, 
pall ial cavity. Scale lines as marked. 





"jalt. > 500m 


I ■ I ( . 80 n is ! ri hi 1 1 i ( m of Rfwphodon kempscyensis sp 

been collected from the surface o! mossy lime- 
stone rocks. 

The lateral teeth in which the mesocone is 
conspicuously large, are similar to those in the 
syrnpatric Letomola eoniorius and Coetio- 
c'haropa yessahahensis which also have been 
collected live on the limestone rocks. This con- 
vergence may reflect similar feeding habits. R. 
kempscyensis is closely related to/?, peregrin us 
arid R. eottsobrinus probably derived by habitat 

Rhophodon elizabethae sp nov 
(Figs 81-83: Tables i- s , 16) 

1 or Elizabeth Chaseling. 


Small size, flammulated shell with bold radial 
ribs, wide umbilicus and number of apermral 
barriers distinguish this species from anv sym- 
patic charopid species. R. minutissimtts is most 
similar to R. ehzabellwe bui has fewei whorls, 
•-mailer umbilicus, and a smaller monochrome 
golden brown shell In addition R ehzuheihue 
(I ig, 81 h) has a greater number of palatal bai 
riers and, unlike R. minuiissimus (Tie 34g.h). 
does not have the second parietal barrier shor- 


I LGfTYPE! QMM017315, Granite Creek, foothill* 
01 I -uhhy RangC; SBQ. 24 '30.7*S. 15I3U.3 E. under 
leaf liner, Collected bV f, Carless. 7 Dec 1983. 
Height of shell III mm, diameter 2.73 mm, H'D ratio 
0.41. D.'L " ratio 2.76. whorls 5 I/4+. 
PARATVPE8: OMM0147I4. I specimen, same data 
as holoivpv; summit Ml Booroon Booroon, SW of 
Miriam Vale, SEQ, MVFMraucami, litter, on and 
among rocks (17. QMMOI6590. QMM016S5S, |7 
Sepi 1985. J- Stanisic. D. Potter); base. Ml Booroon 
Booroon, SEQ, KVF along creek, under logs (3. 
QMMOI6592. 17 Sepi 1985. J. Stanisic. D. Potter). 


Shell minute, diameter 2.26-2.73 mm (mean 
2.47 mm), with 4 3/8 to 5 1/4+ (mean 5-) tightly 
coiled whorls, last descending slightly. Apes 
and earlv spire (Fig. 81a) flat to very slight! v 
elevated, SP/BWW ratio 0.05-0. 19 (mean (1 1 2 I 
height 0.94-1,24 mm (mean 1.10 mm). II/D 
ratio 0.41-0.48 (mean 0.45). Protoconch of 1 
5/8 to I 3 4 whorls, mean diameter 520.1 p,m at 
1 12 whorls. Apical sculpture (Fig. 81e) of 
curved radial ribs, becoming more crowded at 
the nuclear-post nuclear junction. Post nuclear 
sculpture (Fig. SI f) of bold, widely spaced, 
protractivcly sinualed radial ribs. 52-82 (mean 
65.2) ribs on the bodv whorl. Ribs/mm 6.93- 
10.21 (mean 8.38). Microsculpture (Fig. Slf)oi' 
fine, thread-like radial riblcls. 10—1 8 between 
each pair of major ribs, and fine crowded, spiral 
cords which are continuous on the apices of the 
radial ribs. Umbilicus (Fig. Sib) wide, cup- 
shaped, diameter 0,92-1,13 mm (mean 1.0 1 
mm). D/U ratio 2.07-2:76 (mean 2,46). Sutures 
impressed, whorls rounded below and strongly 
shouldered above a slightly flattened periphery 
(Fig. Sic). Aperture ovately lunate with three 
parietal, five palatal and one columellar barrk r. 
tttal 1 mi riers blade-like, consisting of two 
lamellae and an elongate, rhread-like superior 
trace First barrier elongate, crescent-shaped. 
abruptly tapered at each end; second with 
anterior taper and an expanded- downwardly 
deflected, posterior section. Columellar barrier 
low. crescent-shaped, horizontal. Five palatal 
barriers (Fig. 81 h) consisting of four lamellae 
and one trace Lamellae short, expanded. Supc- 
rtor trace sometimes present, short, close to the 
palato-parietal margin. Lip simple; columella 
without noticeable thickening. Parietal callus 
developed. Colour light straw-vellou with va 
able brown radial sireaksand stillusions. Based 
OH 9 measured adults. 







tf ' 





FIG. 82. Rhophodon elizabethae sp. nov. Summit, Mt Booroon Booroon, SW of Miriam Vale. SEQ. 
QMMO16590, paratype. a, genitalia; b, penis interior; c, hermaphroditic duct; d, ovotestis; e, pallial cavity. 
Scale lines as marked. 

FIG. 81. Rhophodon elizabethae sp. nov. a-c. Granite Creek, foothills of Bobbv Range, SEQ. QMM017315. 
holotype; d-h. Summit, Mt Booroon, Booroon, SW of Miriam Vale, SEQ. QMMO16590, QMM016858, 
paratypes. a-c, entire shell; d, microdenticulations on parietal barrier; e, apical sculpture; f, post nuclear 
sculpture; g, radula; h, palatal barriers. Scale lines as marked. 



Genitalia with epiphallus (Fig. 82a) reflexed 
before entering the penis apical ty through a 
simple pore (Fig. 82b) surrounded by a fleshy 
pilaster. Penis (Fig. 82b) simple with apical bulb 
containing epiphallic entrance and lower cham- 
ber containing a single large, spongy, lon- 
gitudinal pilaster, several shorter pilasters, and 
low pustules. No penial sheath. Peniai retractor 
muscle a short tuft inserting apically on penial 
bulb adjacent to epiphallus, 

Radula(Fig. 8tg)tvpical. 

Based on 2 dissected adults (QMiMO 16590). 

range and habitat 

Under logs, on rocks and amongst litter in 
moist notophyll vine forest of the Bobbv Range. 


R. eltzabethae has a very restricted range com- 
pared to other charopids. lis closest relative 
appears to be R. m'muiissunus. 

Rhophodon minutissimus sp. nov. 
(Figs 84-86: Tables 15, 17) 

Latin minulus, little; for the small size. 

typt- material 

Holotvpe: QMM017316. c. 1 km S of Raynbird 
Creek Rd. on Laccy's Creek Rd, Laccys Creek, SEQ 
ITU% 152'43'E. Collected am QM-ABRS, 30 
Stpi 1^82. Height of shell 0.70 mm, diameter 1.29 
mm, H D ratio 0*54, D.'l) ratio 2. 10. whorls 4 5 '8. 

PARATYH-.S: OMMOi:i)94, AMC 136470. 19 
specimens, same locality data as holotvpe; Bunva Mis 
N.P.. SEO. NVF Ct8QMMO6089i 3 Mar 1976, MJ. 
Bishop)*, Dandabah. Bunva Mis N.P., SEQ, behind 
picnic area. (26'50'S, ISlWE) (5, QMMO12280, 
AMC136604, 3 1 Aug 1982. AM/QM-ABRS); 
sidetrack off Mt Archer Rd, Ml Mee S.F.. SEO 
(27WS, 152 41'F) NXT /Anmcaria (3, QMNfO- 

I507S, 6 Sept L983> J. Slamsie, D. Poller); \Vralten\s 

Camp, Wratteffs s.f, SEQ (2r> c i7"S. 152 20'E) 

NVF. lilier(3.0MMOll554, 17 Jul 1980, J. Stamsic. 
A Green); I iule Yabba Creek, Imbil ST., SEO 
(2b : :S-S. 152 C 3S'E). NVF ( I, QMMO12023. B Sep! 
[982. AM, QM-ABRS): Fred's Rd, Mt Mee, SEO 
(27 05 s. 152 43'E) rainforest. leatiitier( I, (JMMO- 
17318. 14 Apr 19SG\J. Sianisic, N. Hall. A. Green); 
Big Fa lis Circuit. Bunya Mis N.P., SEQ. SNVF. under 
logs (1, QMMG 16852, 5 Nov 1985, J. Stanisic, D. 
Potter): Dandabah. Bunva Mis. N.P.. SEQ [26"SFS 


*4*3S l S 

'. ■ 

§alt. > 400m 

^^s>$* ■ w ° ' o n g i 



FIG. 83. Distribution of Rhophodon elizabelhae sp. 

nov. in the Bobb\ Range area, SEO- 

15P35'K). S\T Aniucanu do. QMMO6069, 5 War 
IV76.M.J. Bishop). 


Shell minute, diameter 1. 29-1, 60. mm (mean 
1.52 mm) with 4 3/8 to 4 5,s (mean 4 1/2+) 
tightly coiled whorls. Last whorl descending 
more rapfdh ■ Early spire and apex (Fig. 84a) Oat 
to very slightly elevated, SP/BWW ratio 0.07- 
0.1 2 mm (mean 0.09 mm), height 0.64— 0.76 mm 
( mean 0.70 mm). HD ratio 0.40-0.54 mm (mean 
Q.46 mm). Protoconch of 1 5/8 to 1 3, 4 whorls, 
exsert. mean diameler39 1.9 fxm at 1 1/2 whorls. 
Apical sculpture (Fig. S4e) of curved radial ribs 
and spiral wrinkles. Post nuclear sculpture (Fig. 
S4f) of numerous, prominent, protractively 
sinuated radial ribs. 56-9S (mean 76.6 mm) ribs 
on the body whorl. Ribs/mm 10.9S-21.26 mm 
(mean 16.23 mm). Microsculpture (Fig. 84f) of 
fine radial riblets, 8-12 between each pair of 
major ribs, and very law, barely visible, spiral 
cords which are raised at their junction with the 
microradials to produce a beaded effect. Um- 
bilicus {Fig. 84b) wide, cup-shaped, diameter 
0.60-O.72 mm (mean 0.65 mm). DU ratio 2.09- 
2.59 mm (mean 2.35 mm). Sutures impressed, 
whorls rounded below and shouldered above a 
slightly, laterally compressed periphery (Fig. 
84c). Aperture ovate I y lunate. Barriers present, 
consisting of three palatal and three parietal 



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FIG. 85. Rhophodon minutissimus sp. nov. Big Falls Circuit, Bunya Mts N.P., SEQ. QMM016852, paratype, 
a, genitalia; b, penis interior; c, ovotestis; d, hermaphroditic duct; e, pallial cavity. Scale lines as marked. 

FIG. 84. Rhophodon minutissimus sp. nov. a-ruc. 1km Sof Raynbird Creek Rd ? on Lacey's Creek Rd ? Lacey's 
Ck. SEQ. QMMO12094, paratype. a-c, entire shell; d, microdenticulations on surface of upper parietal; e, 
apical sculpture; f, post nuclear sculpture; g. parietal barriers; h, palatal barriers. Scale lines as marked. 




°E 152°E 1d3°E 

154 C 'E 

FIG. Sn. Distribution of Rhophodon minutissimus sp. 

lamellae (Fig S4g). First and third parietal 
lamellae large with anterior edge protruding 
from aperture, expanding posteriorly: second 
deeply recessed, short, crescent-shaped. All 
directed downwards. Palatal lamellae (Fig. S4h) 
deeply recessed, short, crescentic Lip simple 
without columella expansion or reflection. 
Parietal callus developed. Colour golden- 
brown. Based on 12 measured aduhs. 

Genitalia with epiphallus (Tig. K5a") thick- 
coiled, muscular; longer than the perns, internal- 
ly with longitudinal pilasters, entering the penis 
apically through two fleshy thickenings (Fig. 
85b). Penial retractor muscle inserted at the 
penis epiphallus junction. Penis a broad mus- 
cular tube with sheath; internally with lon- 
gitudinal fleshy pilasters (Fig. 85b). 

Radula tvpical. 

Based on 2 dissected adults (QMM016852). 

Rangf and Habitat 

Notophyll vine forests of the D'Aguiiar 
Range, Bunya Mountains, and the Coast Range 

near Gvmpie. SEQ. R. mmuiisstmus has been 
collected in litter and from under logs. 


Ability to exist in drier open forest is probably 
a contributing factor to the comparatively wide 
distribution of this species. R. hairnsdalcnsis 
displays similar adaptability and breadth of dis- 

Rhophodon colmani sp. nov. 

(Figs 87-88; Tables 15, 17) 

F.TVMm! OGt 

For Mr Phillip Colman, Australian Museum. 


Small si2e ? almost flat spire, widely spaced 
ribs, open umbilicus and aperlural barriers, ef- 
fectively separate R- colmani from any sym- 
patric charopid. Egilomen cochlidium from the 
McPhersOfl Ranges is grossly similar in shell 
form, but lacks aperlural barriers, and has a 
larger shell with higher whori count, higher rib 
count and more elevated spire. 

Conchologiciilly R, colmani is most similar to 
R. bmrnsdalcnsis from southern NSW and Vic- 
toria. However the latter species is larger, with 
more ribs, wider umbilicus, greater number of 
whorls and quite different pattern of apertural 
barriers. The three parietal,. six palatal and single 
columellar barrier of R. colmani (Fig. 87f-h) 
contrast with the single parietal and two palatals 
of/?. batrnsduk'iLsis (Fig. 90a-c). 


Hoi OTYFE: QMM017314. Kenihvorth State Forest, 
SEQ (26 C 35*S S 152*36 - E) ittixod notophyll ^'ne 
forest. Collected by MJ. Bishop. IS May 1976. 
Height of shell 0.ft4 mm., diameter F56 mm, H/D ratio 
ii,4i. D/U rata 2 3Si whdrls A 5/8. 
Paratopes: QMMOhlISS, 14 specimens, same col- 
lection ti,«t.i as hnio:\pe. 


Shell minute, diameter 1.34-1.56 mm (mean 
I.45mm)with4 1/4 to 4 5 ■S(mean4 l/4-)light!y 
coiled whorls, the last descending more rapidly. 
Early spire and apev (Fig, X7a) flat to vcrv sHght- 
Iv rfevfltcd, SPBWW ratio O.U«M)."l4 mm 
(mean 0.1 1 mm), height (1.53-0.66 mm (mean 

FIG. 87, Rhophodon colmani sp. nos Kenilworth S.F., SEQ. QMM06288. para types a-e. entire shell; d. 
apical sculpture; e. post nuclear sculpture; L parietal harriers, g. palatal barriers, h. columellar barrier, Scale 

I in is marked. 



■ ■• . 












151°E 152°E 153°E 154°E 

FIG. 88. Distribulion of Rhophodon colmani sp. nov. 

0.61 mm). H/D ratio 0.38-0.46 mm (mean 0.42 
mm). Protoconch of 1 1/2 to 1 5/8 whorls, exsert, 
mean diameter 357.4 (Jim at 1 1/2 whorls. Apical 
sculpture (Fig. 87d) of curved radial ribs, becom- 
ing more numerous at the nuclear-postnuclear 
junction, and fine almost obsolete spiral threads 
and wrinkles. Postnuclear sculpture (Fig. 87e) 
of bold, widely spaced, protractively sinuated 
radial ribs, 23-29 (mean 26.4) ribs on the bodv 
whorl. Ribs/mm 5.48-6.51 (mean 5.81). 
Microsculpture (Fig. 87e) of fine radial riblets, 
17-25 between each air of major ribs, and low 
spiral cords. Umbilicus (Fig. 87b) wide, cup- 
shaped, diameter 0.51-0.66 mm (mean 0.58 
mm). D/U ratio 2.38-2.60 mm (mean 2.50 mm). 
Sutures impressed. Whorls rounded below and 
strongly shouldered above a compressed 
periphery (Fig. 87e). Aperture ovately lunate. 
Barriers numerous consisting of three large 
parietal barriers (Fig. 87f) protruding from the 
aperture, a large columellar barrier (Fig. 87h) 
situated inside the aperture margin, and six pala- 
tal barriers (Fig. 87g) set back about 1/8 whorl. 
First palatal barrier small, crescent-shaped, 
tapering rapidly anteriorly; second similar in 
shape, but with longer taper anteriorly; third and 
fourth about twice the size of the second with 
long anterior tapers; fifth almost equal in size to 
the second, rapidly tapering anteriorly; sixth 

elongate, thread-like, about twice the length of 
the others. Columellar barrier (Fig. 87h) a large 
crescent-shaped lamella situated well inside the 
aperture. First parietal barrier long expanded 
posteriorly and directed upward; second and 
third, thin anteriorly, expanding posteriorly, 
swollen. Lip simple, without columella expan- 
sion or deflection. Parietal callus well 
developed. Colour light straw-yellow. Based 
on 5 measured adults. 
Anatomy unknown. 

Range and Habitat 

Notophyll vine forest of Kenilworth State 
Forest, SEQ. No data are available on micro- 


Lack of soft parts and poor distributional data 
restrict discussion. R. colmani is most likely 
sympatric with R. minutissimus and it is tempting 
to speculate that the sharply altered shell pat- 
terns, in particular the increased number of pala- 
tal barriers, represent character displacement 
under conditions of sympatry. 

Rhophodon bairnsdalensis (Gabriel, 1930) 

comb. nov. (Figs 89-92; Tables 15, 17) 

Charopa bairnsdalensis Gabriel, 1930. p. 78, pi. 2, 

figs 11-12. 
Endodonta bairnsdalensis (Gabriel); Kershaw. 

1956a. p. 137 
Egilodoma bairnsdalensis (Gabriel); Iredale, 1937a, 

p. 329. 

Previous Studies 

Iredale (1937) included this species in a new 
genus Egilodonta based on the presence of aper- 
tural barriers. He did not give reasons for 
separating the species from the northern 
Rhophodon nor from the geographically more 
proximate Dentherona Iredale, 1933 and Bis- 
choffend If edale, 1937. While R. bairnsdalensis 
has fewer apertural lamellae than other species 
of Rhophodon, shell sculpture is comparable, as 
is the structure of the pallial cavity and terminal 
male genitalia. A review of Dentherona and 
Bischoffena is beyond the scope of this study, but 
coiling patterns and microsculptural details of 
these genera suggest that they are not related. 


R. bairnsdalensis can be separated from its 
congeners by its average size, widely spaced 










d V 


"CV • 


s|- -^ *, 

FIG. 89. Rhophodon bairnsdalensis (Gabriel, 1930). a-e. Marble ArcruS of Braidwood, NSW. AMC12676I ; 
f, near Lakes Entrance, VIC. NMVF53 1 60. a-c, entire shell; d, apical sculpture; e, post nuclear sculpture; f, 
central and lateral teeth. Scale lines as marked. 





FIG. 90. Rhophodon barinsdalensis (Gabriel, 1930). a-d. Marble Arch, Sof Braidwood, NSW. AMC 126761; 
e-f, Bairnsdale, VIC. NMVF 704. holotype. a, parietal barrier; b. basal barrier; c, palatal barrier; d, surface 
of parietal barrier. 



FIG. 91. Rhophodon hairnsdalensis (Gabriel, 1930), -In leaf litter of damp forest gully, near Lakes Entrance. 
VIC.NMVF53160. a, genitalia; b, penis interior; c, d, ovotestis; e. talon and canefour region; f, pallial cavity 
Scale lines as marked. 



FIG. 92. Distribution of Rhophodon bairnsdalensis 
(Gabriel. 1930) in south-east New South Wales anc 
Victoria. (Ca = Canberra; Co = Cooma; Ba = 

radial ribs, wide umbilicus and only three aper- 
tural barriers (one parietal, two palatal) (Fig. 
90a-c). R. colmani is smaller, with fewer widely 
spaced ribs, has a smaller umbilicus and nine 
apertural barriers. Other Rhophodon species 
have a much higher number of ribs on the body 
whorl and are more easily distinguished from/?. 
bairnsdalensis (Table 15). 'Champa' jem- 
mysensis Gabriel, 1947, 'Charopa* colliveri 
Gabriel, 1947 and 'Charopa' snowyensis 
Gabriel and McPherson, 1947 are all sympatric 
with R. bairnsdalensis and have bold radial 
sculpture on the postnuclear whorls. However 
these species lack apertural barriers. 

Type Material 

FfOLOTYPE: NMVF704, Bairnsdale, Victoria. Col- 
lected by T. Worcester. Height of shell 0.86 mm, 
diameter i.89 mm, H/D ratio 0.46, D/U ratio 2.36, 

whorls 4 7/8. 

Paratvpes: NMVF30749, 2 specimens, same data as 


Other Material 

Jemmv's Point, Lakes Entrance, VIC, (5, NMVF- 
25277, CJ. Gabriel: 6, NMVF23326; 7. SAMD5683, 
Jan 1936, C.J. Gabriel; 7, SAMD5683 C.J. Gabriel); 
Wombargo Creek, VIC, rock bank above 3000 ft (1, 
NMVF1678);Toorloo Scenic Reserve, Toorloo Arm, 

9m E of Lakes Entrance, VIC, damp forest gully, leaf 
litter (40, NMVF53367, 6 Dec 1970. D.C Long); near 
Mitchell River. 2m from Bairnsdale, VIC (5, WMVF- 
53365. C. Oke); Jemmy's Point, Kalimna, VIC, leaf 
litter at foot of cliff (6, NMVF53366, 7 Dec 1 970, D.C. 
Long): in leaf litter of damp forest gully, near Lakes 
Entrance. VIC (7, NM VF53 1 60); Toorloo Arm, Prin- 
ces Hwy, 6 mis N of Lakes Entrance, VIC (7, 
AMC153706, 6 Nov 1963, L. Price); Bridle Creek, c. 
0.5km N of Ballantyne's Gap. Buchan-Jindabyne Rd. 
VIC under leaves and logs of damp gully in gum forest 
(31. AMC153704, AMC154810, 13 Jan 1980, W.F. 
Ponder, P.H. Colman): Lakes Entrance, VIC (1, 
AMC 103564, Jan 1935, C.J. Gabriel); Marble Arch, 
S of Braidwood, NSW (35°44 , S. 149°41'E), in litter 
in deep gully on limestone (180, AMC 126761, 17 Jan 
1981. W.F. and W.F. Ponder); Yarrangobilly Caves, 
NSW, beside trail to river cave, in litter (144, 
AMC 142959, 31 Oct 1980, W.F. Ponder, J. Hall); 4 
mis E of LakeTryon turnoff on main road, near Lakes 
Entrance. VIC. in dense bush under leaves and logs on 
ground (64. AMC154770, 10 Jan 1970, W.F. Ponder. 
P.H. Colman); Leather Barrel Creek, near Murray 
River, Alpine Way, Kosciusko N.P., NSW, in open 
forest with some scrub, in leaf litter ( 19, AMC154808, 
1970, W.F. Ponder, P.H. Colman); Deep Creek, 
Kiewa Hwy, c. 5 ml N of Tawonga. VIC, in open bush 
on roadside, leaf litter (20, AMC1 54757, 18 Jan 1970, 
W.F. Ponder, P.H. Colman); Mt Drummer, VIC (21, 
AMC154728, 27 Sept 1959, D.F. McMichael). 


Shell minute, diameter 1.85-2.42 mm (mean 
2.06 mm), with 4 5/8 to 5 3/8+ (mean 5) tightly 
coiled whorls. Apex and early spire (Fig. 89a) 
depressed, flat or only slightly elevated, 
SP/BWW ratio 0.00-0.23 (mean 0.10), height 
0.76-1 .05 mm (mean 0.89 mm). HD ratio 0.39- 
0.48 mm (mean 0.43 mm). Protoconch of 1 1/2 
to 1 5/8 whorls, exsert, mean diameter 385.8 |xm 
at 1 1/2 whorls. Apical sculpture (Fig. 89d) of 
curved radial ribs, more crowded at the nuclear- 
postnuclear junction, and fine spiral threads. 
Post nuclear sculpture (Fig. 89e) of bold, widely 
spaced, slightly protractively sinuated radial 
ribs, 23-57 (mean 36.2) ribs on the body whorl. 
Ribs/mm 3.96-8.33 (mean 5.56). Microsculp- 
ture (Fig. 89e) of fine radial riblcts, 10-22 be- 
tween each pair of major ribs, and fine 
microspiral cords which are continuous on the 
apices of the radial ribs, and raised at their junc- 
tion with the microradials to produce a beaded 
effect Umbilicus (Fig. 89b) wide, cup-shaped, 
diameter 0.74-1.13 mm (mean 0.88 mm), D/U 
ratio 2.02-2.66 (mean 2.37). Sutures impressed. 



Whorls rounded below and shouldered above a 
compressed periphery (Fig. 89c). Aperture 
ovately lunate. Three apertural barriers present. 
Parietal barrier (Fig. 90a), low, elongate, thread- 
like anteriorly, protruding beyond the lip margin, 
expanded and deflected downwards posteriorly. 
Parietal trace sometimes present. Palatal bar- 
riers two. First (Fig. 90b), short, low, crescent- 
shaped situated at the baso-columellar margin; 
second (Fig. 90c) elongate, slender, located 
halfway up margin of outer lip. Lip simple with 
no columellar thickening or deflection. Parietal 
callus present. Colour golden yellow-horn to 
white with light-brown radial suffusions. Based 
on 32 measured adults. 

Genitalia with epiphallus/vas deferens junction 
valvular. Epiphallus (Fig. 9 1 a) reflexing and loop- 
ing the penial retractor muscle before entering the 
penis apically through a simple pore (Fig. 91b) 
Hanked by two fleshy thickenings, internally with 
longitudinal pilasters. Penial retractor muscle in- 
serting at the epiphallus/penis juntion. Penis (Fig. 
91b) with apical bulb. Main penial chamber inte- 
rior with a fleshy longitudinal pilaster and several 
low longitudinal thickenings (Fig. 91b). 

Radufa (Fig. 89f) typical. 

Based on three dissected adults (NMV- 

Range and Habitat 

From Bairnsdale, Victoria, north through the 
Snowy Mountains area to the Yarrangobilly 
Caves and Braidwood, southern NSW. The 
species lives in open scrub, moist forest and on 
limestone outcrops. Microhabitat appears to be 
in litter and under loss. 

(1983, p. 76) gives a full synonymy of a range of 
geographic variations. 

Discocharopa aperta (Mollendorff, 1888) 
(Figs 93-94; Tables 15, 18) 

Panda aperta Mollendorff, 1888, p. 89 

Endodonta concinna Hedlev, 1901. p, 279, pi. 48, figs 

Discocharopa concinna (Hedlev); Iredale, 1937a, p. 

Discocharopa apena (Mollendorff); Solem, 1983, pp. 

76-81, figs 35-37; Solem. 1984, pp. 164-165, figs 


Previous Studies 

Solem (1983) synonymised D. concinna with 
D. aperta, D. planorbulina (Tate, 1896) from 
Central Australia, and further species from In- 
donesia, New Hebrides, Fiji, Kermadecs, Samoa 
and the Society Islands. In spite of differences in 
apertural barriers, intensity of spiral sculpture, 
size, umbilical width and rib count. Solem 
(1983) could not effectively separate the distant 
populations and opted for a single wide ranging 
species. Solem (1984) reviewed the Australian 
distribution of this species and provided records 
from west, north-west and central Australia with 
brief mention of likely distribution limits in east- 
ern Australia. In the absence of soft parts for 
study, Solem's classification is maintained, al- 
though it is probable that the SEQ populations 
may be a separate species derived from local 
charopid ancestors. 


R. bairnsdalensis is a southern outlier 
separated from its nearest relative, R, kemp- 
seyensis, by almost 500 km. 

Discocharopa Iredale 1913 

Discocharopa Iredale, 1913, pp. 279-280; Iredale. 
1937a, p. 325; Kershaw, 1956a. p. 140; Burch, 
1976, p. 132; Solem, 1983, p. 74; Solem, 1984, p. 

Type Species 

Champa, (Discocharopa) exquisita Iredale, 
1913; by original designation. 

For diagnosis see Solem ( 1 983. p. 74). Solem 


D. aperta is distinguished by its minute, white 
shell which has normally coiled whorls, almost 
flat spire, crowded, protractively sinuated radial 
ribs, and wide saucer-shaped umbilicus (Fig. 
93). D. aperta bears a general similarity to 
Rhophodon in apical sculpture, and in particular 
with R. kempseyensis in microsculpture on 
postnuclcar whorls. However, Rhophodon has 
more shouldered, tightly coiled whorls, a cup- 
shaped umbilicus and few to many apertural 
barriers. Solem (1983), using material from the 
Kermadec Islands, showed the species to have 
terminal male genitalia without an externally 
differentiated epiphallus. While Cralopa 
Iredale, 1941 approaches this condition, the lack 
of an epiphallus is more prevalent among New 
Zealand charopids. 







^(j^™®*- |n 

FIG. 93. Discocharopaaperta (MdUcndorff, 1888). a-b, Bundaberg, Old. AMC8970Jectotype; c-f, c. 1.8km 
E of Builyan Rwy Station, on Builyan-Gladstone Rd, SEQ. QMMO17308. a-c, entire shell; d, apical 
sculpture; e-f, post nuclear sculpture. Scale lines as marked. 



22 C E 


:6 a s 


0alt. > 500m 

150 b e 151°£ : i -'"f, 

j 5-3" £ 

FIG. 94. Disirihution t>1 Disccharopa aptrta 
( Mollendortt, 1888} in southeast Queensland. 

IM'F M\lhRI \L 

Lectotypf. (designated Solem. LSB3,p. 77J; AMI' 
8970, Bundaberg, Old.- ex May. Heighl 0-61 mm, 
difltnetci \M mm. K/D ratio 0.45, D U ratio 2.SI 

pARAI fv toi vi'i s: AMC63489. 7 specimens, same 
dataas teciolvpe. 

Other Material 

C. I km Irom OtsCfl's Caves, MFC i23 r 10'S f 
150'30'S) litter, limestone outciop (16. QMMO- 
17307, 13 Jul L9S0, J. Stanisic. A, Green, G. An- 
i i,ll): Ml Archer. Bersaker Ra.AIEQ (2321 S. 
I5U 35 E), remnant vine thtekel (6. QM MO 1)922. .^< I 

Jun 1982, J. Stanisic); c. 2 km \V of Gavndah, on 
Gayndah-Hinesville Rd. SEQ t25'3SS. 15K3<vt 
bottle tree scrub/SEVT. under rocks and in litter (2 
QMMOl 1528,16 Jul 19S0,J.Stanisic, A.Green); Dnn 
Dan Scrub, SEQ. SEVT, in tiller (2. QMMO16800. 10 
May 19S4, J. Stanisic. D. Potter); Dawes Range. SEQ 
(24 C 28'S. 15 1 '07'E) MV¥fAr*ncaria (28. 
QMMOl 2666, AMC 136795. 4 Sept 1982, AM/QM 
ABRS):c. 1,8 km E ot Builvan Kwv Stn, on Builvan- 
GladstoneRd.SEQ{2422"S. 151 24'E), rainforest. j: 
nver hank litter [13, QMMO17308, 15 Jul 1980, J. 
Stanisic. A Green); Ml Biggenden. SEQ (25 a 32*S, 
15T50 E), SEVT. limestone outcrop (5, QM- 
MQ173Q9, BJul 1980. J. Stanisic, A. Green; 1, QM 
M01683S, 3 Jul 1984. J. Stanisic. D. Potter. K. 
Emberton; 14, QMMOl [958; AM CI 36396, 3 Sept 
L9B2, AMQM - ABRS); Walla Range. E of Ban Ban 
Springs.SEQ(25 38 S.15I 50E)SFVT(1.QMMO- 
12414. 3 Sept 1982, AM/QM - ABRS); Kalpowai 
S.F.. SEQ Wf&AMkcwk (2, QMMO17310. 15 Jul 
1980. J. Stanisic. A. Green. 2. QMMO17306, 4 Sep! 
1982, AM QM -ABRS); Gavial Cfc Bouldercombe, 
MEQ (23"34*S, 150'2S'E) MVF, litter (4, QMMO- 
16808, 9 Jul 1983. J Stanisic. D Putter). Coalstoun 
Lakes Rd. Coalstoun Lakes, SEQ (25 '37' S, 151"53 ? E) 
SEV1 I. QMMG12396, 2 Sept 1982. AM/QM - 
ABRS); c. 54 km S of Biioela, on Bitoela-Monlo Rd. 
SEQ (24 3 45'S, I5D*50*£) SEVT on rockv outcrop, 
litter (3. QMMOI731L 14 Jul I9S0, J. Stanistc, A. 
Green »:Mt Etna, MEQ (23'09'S. 150 27E) MVFon 
limestone outcrop, litter (100. QMMOI73I2, 13 Jul 
1980, J, Stanisic. A. Green, G.Annabell); Johannsens 
Caves, MEQ (23'09'S. l50 e 2TE)MVF on limestone 
DUiCfOPi Huer (19. QMM017313, 13 Jul 1980, J. 
Stanisic, A. Green, G. Annahell), Cammoo Caves. •.. 
24 km N of Rockhampton, MEQ l231(rs. 150*771 I 
limestone outcrop (4.AMC1521 45. W. Ponder, P.M. 
Colman. LB. Buich, 8 Sept. 1970); Glastonbury SI , 
c.0.25 km Sot" lor- ti ft (26 I3*30"S, 152 2' 

45°E) ii. QMMD 13305, 7 Sept I^.AM/QM- 
ABRS); S of Gvmpie, entrance to state forest off 
Yabba Cfc Road (26'28'S, !52\3N'E) (3, AMl\ 
138110 QMMOI33h5. 7 Sept L982, AM/QM- 
ABRSi: Ken.Kvorih SF, SEQ (26 B JTS« 152'42'E}, 
rainforest, leaf litter in. QMM0836Q, 22 Ma> 1980 
J, Stanisic, A. Greeny sidetrack oft" Mt Archer Rd, Mt 
MecSF. SEQ |2" 04 S, l52*4rE) ( 1, QMMOl 2 I 
31 AV| 1982, AM QM-ABRS): Natural Bridge NP. 
SEQ (28 13 S. 153 !4*E) NVE (4, QMM01O 
AMCi: i J2{ : .h, IK Mar 1 98 1 . AM, QM- ABRS); Fred** 
Rd, M. Nice, SEQ (27'05"S, 152 43'E) rainforest, 
litter (3. QMM08357, 14 Apr 1980, 1 Stanisic, N 
Hall, A Green 1; Yarraman Forest Drive. Yarraman 
SE. SEQ ( 2d 50 S 1 5 T 5 7 " E ) MVF (i. QMMO- 
I226L AMCI3S5K& 31 Aug 1982, AM, QM-ABRS 



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c. 15 km from Rathdowney on Boonati-Raihdowney 
Rd. SEQ. ruckv outcrop, vine thicket (28"lfS. 
152 B 46"E) <8, QMMOlOSriS. AMC128445, Mar 
| AM'CJM-ABRS); Binna Bnrra, Lamington 
N.P., SEQ (28M2'S, 153' WE) Wf Palms (1, QM- 
MO 10444. 18 Mar 1 981. AM/QM-ABRS); Koreelah 
Ck.. Beaurv S.F., c. 15 km W ol rjrbenviFle, NSW 
(2» :i'S. 15: 2irE)530mJuter(2,AMC152l51.18 
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junction with Lever's Plateau Rd, Tartar & Creek. SFO 
(28 28'S, 152 08'E), roadside gully (1, QMMO- 
I I US. 17 Mar ls>SJ AM/QM-ABRS). 


Shell minute, diameter 0,82-3 2 l ) mm (mean 
L19 mm) with 3 1/8 to 3 3/4 (mean 3 1/2-) 
loosely coiled whorls, last whorl descending in 
front. Early spire and apex (Fig. 93cj barely 
elevated, SPBWW rail' ; 1 1 N i 0,22 (mean 0. 1 1 }. 
heigbl <rf$hell 0.39-0.64 mm (mean (L49) H/D 
ratio 0.35-0.55 (mean 0.44). Protoeoneh of I 
3 S to 1 5/8 whorls, mean diameter 693.9 u.m at 
I 1/2 whorls. Apical sculpture (Fig, 93e.f) of 
widely spaced, slightly sinuous radial ribs and 
even finer, irregular spiral riblets. Postnuclcar 
sculpture of crowded, protractivcly sinuatcd 
radial ribs. o7-l 34 (mean %.70) ribs on the body 
whorl. Ribs/mm 24.25-40.74 (mean *1 75) 
Mierosculpture (Fig. 93e,f)of fine radial riblets, 
whose apices are erenulate. 4—10 between each 
pair of major ribs. Umbilicus (Fig. 93b) saucer- 
shaped, diameter 0.34-0.57 mm (mean 044 
mm), D/U ratio 2.16-3.3! (mean 2.41). Whorls 
rounded above and below the periphery. Apcr- 
lure ovalcly lunate. Lip simple. Parietal callus 
developed. Colour light straw-yellow to white. 
Based on 2S measured adults. 

No soft parts available for dissection. 

RvNJGr ,v>p Habitat 

In eastern Australia from just north of Rock- 
hampton. south to near the Queensland/New 
South Wales border. In the northern part of its 
range i.e. between Biggenden and Olsen's 
Cases, the species lives in dry subtropical 
microphyll vine forest. In the south i.e. between 
(ivmpie and Natural Bridge National Patk. f>. 
aperta inhabits warmer moist notophyll vine 
iorests. P. apcrta lives among the dtrl particles 
below the leaf litter. 

Rj \!M<US 

The decision to consider all the above 

.uncus as D. apena is made reservedly. 

There areconchological differences between the 

northern and southern populations which indi- 
cate two species. However, without soft parts. 
recognition of two species is a tenuous proposi- 
tion (Solem. 1963). P. apenu is probably a 
complex of species which represent localised 
experiments in mimanuisauon. 

Cralopa Iredalc, 1941 

o-tf/o/wrredale*l94li r ZW7 Korriiaw. lMfib + -p.8; 

Butch. 1976. p. I 


f lefts ttroydensii COX 1854 :by original desig- 

Iredale (1941a) introduced Cralopa for Helix 
stmudensis Cox, 1864 which has a small shell 
with weakly elevated spire, closed umbilicus 
and numerous curved, bold radial ribs. 
'Cralopa' int&l$A Iiciiale. 1941 was added on 
the basis ol small si/e and crowded radial 
postnuclcar sculpture. Two new species 
added here and '( /.•rvnsr/ is removed to 
Sinptoyea Solem. 198 


Shell small, diameter L66-3-63 mm. with 3 

34 h>4 12 tightly, to loosely coiled whorls, the 
last descending slowly. Larlv spire and apex 
slightly elevated. Apical sculpture of prominent 
to bold, weakly curved radial ribs and low. nar- 
row spiral cords. Adult sculpture of few {carter- 
\i) 10 mans [straudeitsis)i widely spaced 
{carlrssi). to eiovxdcd {kuputarensis), bold, 
strongly protectively sinuatcd radial ribs 
Mierosculpture of fine radial riblets and 
crowded, narrow spiral ands, continuous on the 
major ribs. Umbilicus closed m only narrowly 
open. Sutures deeply impressed. Whorls round 
to slightly shouldered above the periphery and 
rounded below. Aperture roundly lunate. Lip 
sinuous, Strongly retracted at the sutural margin 
to form an apertural sinus. Columella reflected 
over the umbilicus. 

Foot shot!, bluntly rounded posteriorly. coIout 
white, ommatophuies white. Mantle collar with 
an orange glandular section extending onto the 
pallia! roof. Kidnev weakly bilobed. wilh 
pericardial lobe longer. Ureter sigmurethrous, 
eomplcLe. w ith primary arm teflexed near origin 
Terminal male genitalia with a relatively ex- 
panded vas deferens, very large cylindrical 
penis, ^\\m\ weakly differentiated epiphallus. In- 





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3^ 55 

ternally T penis with apical epiphallic pore and 
two large, spongy longitudinal pilasters. Pcnial 
retractor muscle inserted on penis (stroudensis, 
carlessi), or epiphallus near penis (kaputaren- 
sis). Vagina relatively short. Radula with 
tricuspid central and lateral teeth; central tooth 
reduced, about 1/2 size of the laterals, cusps 
lanceolate, mesocone long; lateral teeth with 
large lanceolate mesocone. 

Distribution and ecology 

The east-west distribution of Cralopa con- 
trasts with the more common north-south pattern 
shown by other genera reviewed in this study. 

Although the extensive rainforest distribution 
of C. stroudensis appears to contrast with the 
'cornered' distributions of C. kaputarensis and 
C. carlessi, closer analysis reveals that C. 
stroudensis is also an environmentally isolated 
species in cool subtropical notophyll vine forests 
of the Great Dividing Range. 

Patterns of Variation 

Conchological variation is conservative and 
probably reflects relatively recent fragmentation 
of populations. It involves changes in size, spire 
protrusion, whorl count, rib spacing and rib num- 
ber. C kaputarensis, the smallest member of the 
genus, is a scaled down version of C. stroudensis 
with less protruded spire (SP/BWW ratios = 0.54 
and 0.99 respectively ) and more crowded ribs on 
the body whorl. C. carlessi is larger with greater 
whorl count and more elevated spire. C. 
stroudensis has the greatest number of ribs but 
C kaputarensis has most ribs/mm. C carlessi 
has the widest spaced ribs. 

An interesting aspect of the Cralopa shell is 
the development of an apertural sinus. This is 
more commonly seen in New Zealand 


Cralopa is most similar to FAsothera having 
shells with very similar sculpture, barely open to 
closed umbilicus, and relatively depressed spire. 
However, Elsothera is much larger with greater 
mean whorl count and higher rib count (Table 
19). Furthermore Elsothera has an almost 
unilobed kidney, tapered at the apex and the 
primary ureter as a straight tube (Figs 109f, 
105e). In contrast the kidney of Cralopa is 
weakly bilobed with the apex of the pericardial 
lobe and initial part of the primary ureter weakly 
reflexed (Figs 96e, 99e, 102). 'Cralopa has a 
large, cylindrical penis with two large 



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longtitudinal pilasters (Figs %b. 99b, 102b) in 
conlrasl to Efsothera which fws a pear-shaped 

penis internally divided by an apical collar (Figs 
105c. 109b). The epiphallus- vas deferens junc- 
tion of Elsotherah highly modified (Figs 105b. 
! 09a) while thai oiCralopa is scarcely differen- 
tiated (Figs 96a. 99a, 102$). Egilomen gfobosQ 
also has prominent radial ribs and a closed um- 
bilicus but differs in having beaded miciosculp- 
ture(Figs 128g, 1310- 

Cralopa stroudensis (Cox, 1864) 
(Figs 95-91; Tables 19,20) 

Wtf//XtfrtfJf^eW5isG0e, 1864, p. :0 :Tr\iin,l.SS7, p.25. 
pt 4,fig». 2S.29. 

■'. a i!roiuh'n*t.\{C^\):\ied\c\. lv>24, p. 21"'. 
pi, 11, figs 25-27; Iredale. 1937*, p. 323. 
Cnili-'pLi stroudettsis (Cox); [re dale, 194 la. p &9 

Comp i rsa • ■ 

C . stroudcttsis can be distinguished from its 
congeners by high rib count, relatively large size 
and moderate spire elevation. C. ktiputurensis IS 
smaller, has a lower rib count and almost Hat 
spire (Tig 98c), C. carlessi is larger With reduced 
Tib COUIIt and strongly elevated spire (Fig. 101c). 

tomically the differences are subtle. C 
stnnulcnsts (Fig. 9Gb] has a large cylincl 
penis with ftVO unequal, longitudinal pilasters. 
and a penial retractor muscle inserted OH the 
penis apex; C kupularensis t Fig, 99b)hasa large 
penis with two large longitudinal pilasters which 

tOIDOSe and divide into smaller pilasters 
hasdly. and a penial retractor muscle which is 
inserted on the epiphallus; C, rarlcssi (Fig. 
102b) has the penial retractor muscle inserted oi 
the penis which has ftVO slender equal pilasters 

a low central pustular swelling internally. 


Ho] di n IMO&3500. Swoud, nsw Call Rev. 
RL King Mciet'ii t bl mm, dtametei 2.56mm, H/D 
ratio 0.to3, D ' i ci f ;,) . whorls 4 

PAK\hn\ ftMC225, 4 specimens, same Jala I* 
holi p 

'■ | . i, '.I.M 

aril Falls. Dorrigo, NSW <'"' f m. v\.irm 
operate ratnfdn i titer (L QMMOI6778, 13 

Sos I'^IAMOM- ARRS);e 2H Skrrt t'rurtiGoum- 
hurr.i. na Uovimbmra S.F. road. SE0 |27*59 , S. 
152*21 l | fiOtl m CNVF/Palrm [H. QMMj 
QMMOItllS, AMC136gl6, 7 bet IWI AM I M 

ABRS}. Ycs^.ih-th 1 uriesione Cave. Sherwood Ouar- 

rv . Wol RempseyNSW (31 Ub'S, 152 42'E) ground 
later in tofcave(2 <\MCI2168G\90ct. 1979, 

P.H. Caiman J. Siamsiej; Lowct Ballanjui Falls Cir- 
cuit. Binna Burra, Lamington NT.. SEQ, NVF, litter 
{i.QMMno,:-N-..j.Si:jmMe-D. Potter J. ChassJing); 
ft Nts N.P. (27*5.VS, 1 5 1 3 3 ' E ) NVF (F QM- 
MO6072L OMM06278 3 5 Mar 1976, MJ. Bishop): 
initial pan of" Kweebani Caves walk, Binna Burru. 
Lamjiigton N.P , SEQ NVF, Under log (F QMMO- 
16888, { > Mar 1984 J. Stamsie); top of Condamine 
River Valley, SEQ (28*15*5 152*29' B) (2. QM 
MO104K). AMC12SW. Mar 1981, AM QM - 
ABRS); Cherrv Plain - WestcjiH Plain Circuit. Bunva 
Mis N.P.. SEQ. NVF M\T. under logs (43, QM- 
MO!fi857.+ 6N<n 1965, J. Startisfe D.Potter); Dan- 
± h Bunya Mts N.P., SEQ. NYF.unJcr logs (1>, 
QMMOTQ846, I J Maw I0S4-J Stajiiwc, Df. Potter); 
Big RaltS Circuit Bunva Mis N.P.. SEQ. SNVF. under 
logs [27, QMMOI6K51, 5 Nq> 1985, L Sianisie, D. 
i a ler): Hails Plain, I ol Emu Vale. SEQ, CNYF. 
undei log ii.qmvioo-.svi, a Maj iSSe I 5u i i 
.1. Chaselmgi; Cunningham^ Gap N.P.. SEQ near 
monument. 755 m 128WS. I52 3 24 , E)SNVF/Fems 
1 1, QMM012685, 7 Dee NM, AM QM - ABRS), e, 
IS km lYorn ! e^ume, on Acacia Plateau Rd, Koreefah 
S.K. SI O.. SFA'T NVF \t\F i I. QMM01271S. S 
Dee (981, \\1 QM - ABRS): Bellengen River S V . 

nsw i:Mr-':_s-s. |5J*4**E> dense rainforest d. qm- 

MOl'isu. AMC1283B9, 12 Mar, 1981, AM OM - 
ABRS r. Bunyn Mis MV, Si Q ( 1 X AM( 63869, B 
1961 D.F, MeMiehud): R.^ckv C^k. ReJ Scrub Flora 
ReserM;,NS\Vi:s ;.N's. 153 20'1 |i I U i ( I 3 J8C ' 
211 M.n 1983. AM QM - ABRS); Porl Maiquarie (1. 
\WC36564,ejLHedIeyl:Dandabah, Bunya Mis, SEQ 
S, 15 I ?h'h \ ( 15s QMMQI 22s K 

ANH i v , i i-,:, am QM - ABRS). Port 

Maoqudite 1 1- AMC63H55. ex Cux>: Koreelah S.F., 
NSW. St:\ T-MVT (2S 2 1 S. 15 2 24 E) (11, 
AMCI2S5S7, OM\, i f] | ISMarP'M AM.QM 

- abrsi. Naiural *\cch, Caitai < aves» near Kempsev, 

NSW. litter* l,AMC1^2ns flftb I9S0. P.H, Col- 
in n i 

Di \r\; st! 

Shell Lli;imeter 2.2f>-3.83 mm (mean 2,89 mm) 
With J 7 s to a 1,44 {mean 4+) loosely coiled 
whorls. Apes in h .'i i\ spire ( Fig. 95b.) slightly 
ele\aieei. SP/BWW ratio 0.14-0.25 [m i 
'i l :i ). bcighl F4.s-2.22 mm (mean 1.71 mm). 
H/D Eaiia 0.52r<» T 67 (mcai D.6G) last whorl 
descending much more rapidK. Protoconch of 
I I 2 to I 5 8 whorls, mean diameter 5''3, 3 [Jim 

at 1 12 whorls. Apical sculpiurc (Fig. ^5d) of 

numerous, high, curved radial rihs. becoming 
■-liuhth more crowded ti Ward the nuclear-i 



I mm 

f ?< 

FIG. 95. Cralopa stroudensis (Cox, 1864). a-c, Stroud, >ISW. AMC63500, holotype; d-f, Dandabah, Bunya 
Mts, SEQ. QMM012281. a-c, entire shell; d, apical sculpture; e, post nuclear sculpture; f, central and lateral 
teeth. Scale lines as marked. 


FIG.%. Craiopa stroudensis (Cox, 1864). Dandabah. Bunya MisN.P.. SEQ. QMM01228L. a. genitalia; b. 
penis interior; c, hermaphroditic duct, carrefour and talon: d, ovotestis; e, pallial cavity . Scale lines as marked. 











151 °E 152°E 153°E 154 °K 

FIG. 97. Distribution of Cralopa stroudensis (Cox, 

nuclear junction, crossed by fine spiral cords. 
Post nuclear sculpture (Fig. 95e) of numerous, 
large, strongly protractively sinuated radial ribs 
69-109 (mean 91.4) ribs on the body whorl. 
Ribs/mm 7.84-13.57 (mean 10.08). Micros- 
culpture (Fig. 95e) of fine radial riblets, 3-10 
between each pair of major ribs, crossed by 
numerous, regularly spaced spiral cords. Um- 
bilicus (Fie. 95b) very small to closed, diameter 
0.29-0.43 mm (mean 0.36 mm), D/U ratio 7.86- 

8.52 (mean 8.15). Sutures deep; whorls slightly 
shouldered above and rounded below the 
periphery (Fig. 95c). Aperture roundly lunate. 
Lip sinuous, strongly retracted at the sutural 
margin and forming an apertural sinus. Columel- 
la expanded and slightly channelled in older 
specimens, weakly reflected over the umbilicus. 
Parietal callus developed. Colour variable, 
white through straw-yellow to light-brown. 
Based on 10 measured adults. 

Epiphallus (Fig. 96a) not strongly differen- 
tiated from the vas deferens, entering the penis 
apically through a simple pore (Fig. 96b) sur- 
rounded by fleshy thickenings. Penis (Fig. 96b), 
internally with two large, unequal longitudinal 
pilasters, and a series of low, longitudinally ar- 
ranged pustulations. Penial retractor muscle in- 
serting on the penis head adjacent to the 
epiphallus. Free oviduct (Fig. 96a) very long, 
twisted and muscularised, internally with a spon- 
gy pad-like thickening. Vagina long with inter- 
nal lamellar thickenings. 

Radula (Fig. 95a) as for genus. 

Based on two dissected specimens (QMMO- 

Range and Habitat 

Cool subtropical notophyll vine forests of the 
Great Dividing Range between Barrington Tops, 
NSW and Bunya Mfs, SEQ; the McPherson Ran- 
ges of the QLD/NSW border; and the limestone 
ridge which extends intermittently from Yes- 
sabah to Carrai, W of Kempsey, NSW. C. 
stroudensis lives under logs and sometimes 
occurs in large numbers per site. 


There is significant conchological variation 
between populations. However insufficient 
material is available to investigate its extent. 
The Yessabah Cave shells (AMC121680) are 
noticeably larger, possibly due to the abundant 
supply of calcium for shell construction. 

Cralopa kaputarensis sp. nov. 
(Figs 98-100; Tables 19, 20) 

For the type locality, Mt Kaputar. 


Cralopa kaputarensis is separated from C. 
stroudensis in discussion of that species above. 
The barrel-shaped penis is a conspicuous char- 
acter which identifies C. kaputarensis. C. car- 





* '- ~'"S 

i A/ 

4 Jb 



f V v \v-it 

FIG. 98. Cralopa kaptttarensis sp. nov. a-c. Summit, Mt Kaputar. NSW. QMM016753, holotype; d-f, 
Dawson Springs, Mt Kaputar, NSW. QMMO 16742. paratypes. a-c. entire shell; d. apical sculpture; e, post 
nuclear sculpture; f, central and lateral teeth. Scale lines as marked. 



FIG. 99. Cralopa kaputarensis sp. nov. Dawson Springs, Mt. Kaputar, NSW. QMM016742, paratype. a, 
genitalia; b, penis interior; c, ovotestis; d, talon, carrefour and hermaphroditic duct; e, pallial cavity. Scale 
lines as marked. 



,1 w j 

JD* 211 5 

' . 

: : 

I St 

FIG. 100- Distribution of Cralopa kaputaraisis sp. 
nov. on ihe Ml Kaputar Plateau. 

fett/is much larger with more elevated spire and 
Hatter whorls. 

Type Material 

HoloTYPE: QMM016753, summit, Ml Kaputar, 
NSW. 1500 m, among rocks, grass and litter. Col- 
lected 9 Nov 1983, AM/OM - ABRS. Height 0.90 
mm, diameter 1 ,66 mm, H/D ratio 0.54, whorls 4+. 

Paratypes: Dawson Springs, Mt Kaputar. NSW. 
1300 m, under lo^s, in tern gully (3, OM MO 1 6742. S 
Nov [9g3, AMQM - ABRS); Joker's Spring. Mt. 
Kaputar. NSW. J 000 m, edges of rocky creek bed, 
among lichen (I, QMMOI675U. 8 Soy 1983. 

Shell diameter 1.66-1.75 mm (mean 171 
mm), with 4+ to 4 1/8+ (mean 4+) loosely coiled 
whorls, last descending slowly. Early spire and 
apex (fig, 98a) verv slightly elevated, SP/BWW 
ratio (M 5-0.19 (mean Q.17), height 0.90 mm. 
Protoconch of 1 1/2 whorls, mean diameter 
734.0 p.m at 1 1/2 whorls. Apical sculpture 
(Fig. 98d) of slightly curved, radial ribs and fine 
low spiral cords, more noticeable toward the 
sutures. Postnuclear sculpture (Fig. 98e) of 
large, crowded, strongly protractively sinuated 
radial ribs. 69-80 (mean 74.5] ribs on the bodv 
whorl. Ribs/mm 12.57-15.30 (mean 13.94). 
Microsculpture (Fig. 9Se) of fine radial riblets, 
6-10 between each pair of major ribs, crossed by 
crowded, regularly spaced, low spiral cords. 
Umbilicus (Fig. 98b) reduced to a slender crack. 

more open in sub-adull specimens. Sutures im- 
pressed. Whorls rounded above and below the 
periphery (Fig. 98c). Aperture roundly lunate. 
Lip sinuate, retracted at Ihe sulural margin to 
form an apertural sinus. Columella dilated and 
reflected over the umbilicus. Parietal callus 
well developed. Colour dark brown. Based on 
2 measured adults. 

Vas deferens a thin tube, giving rise to a weak- 
ly differentiated epiphallus. Epiphallus entering 
penis apieally through a simple pore. Penis 
broadly tubular, internally with two large lon- 
gitudinal pilasters which anastomose, and divide 
into numerous smaller pilasters basally. Penial 
retractor muscle (Fig. 99b) inserted on the 
epiphallus. Female reproductive system small 
compared with the penis. 

Radula (Fig. 9S\) with central tooth much 
smaller than laterals. Based on one dissected 
specimen (QMMO 16742). 

Range and Habitat 

Only on the Ml Kaputar Plateau, near Narrabri, 
NSW. Under logs, rocks, and among lichen on 
the edges of springs. 

C. kaputarensis is closely related to C. 

uroutlcnsis and is probably a recent derivative. 

Cralnpa earlessi sp. nov 
(Figs 101-103: Tables 19,20) 


For Terry Carless who collected specimens for 
this study, 


The flatter more loosely coiled whorls, 
elevated spire and reduced number of radial ribs 
on the body whorl characterise C. cariessL C 
kaputarensis has an almost flat spire (Fig. 98c) 
and is much smaller {Table 19), 


Holotype: AMC12353S,lnverelI, NSW. Collected 
by C.T. Musson, 1S89. Height of shell 2.04 mm. 
diameter 3 62 mm, H/D ratio 0.56. whorls 4 1/2. 

PAP VTYFES: AMCfl537l6, 2 specimens, same col- 
lection data as holotype; Hills of Glenrock Stn, NSW 
(31"*rS, 151 24I0"E) alt. c. 600 m, under logs and 
limestone slabs and litter, dry. semi-open scrub (2, 
AMCI44200, 17 j U n I984 t Fif. Colman); Worn- 
be van Caves. W 7 ol Mittagong, NSW. sleep scrub and 
grass covered limestone slope, in litter outside large 





1 mm 


1 1 • 



; ^m^ 







FIG. 101. Cralopa carles si sp. nov. a-c, Inverell, NSW. AMC1 23538, holotype; d-e, Hills of Glenrock Sin. 
NSW. AMC144200, paratype; f, Inverell, NSW. AMC3640, paratype. a-c, entire shell; d, apical sculpture; 
e, post nuclear sculpture; f. central and lateral teeth. Scales lines as marked. 




FIG. 102. Cralopa carlessisp. nov. Hills of Glenrock Stn, NSW. QMMO 144200, paratype. a, genitalia; b, 
penis interior; e. talon and hermaphroditic duct; d. ovotestis; e, pall ial cavity- Scale lines as marked. 





31 = S 



34 °S 






i : :. -- :-. 

FIG. 103. Distribution of Cralopa carlessi sp. nov. 

cave (2. AMC142967, 28 Dec 1981, M. Shea); West 
side Kowmung River at Gridiron Bends, nearTuglow 
Caves, NSW, in litter and soil on limestone bluffs {7, 
AMC124279, 12 Dec 1979, W.F. Ponder. P.H. Col- 
man); Attunga, NSW (2. AMC28496, C. Laseron); 
Inverell, NSW (3, AMC123610, C.T. Musson); In- 
verell, NSW ( 1 1, AMC3640, W.S. Duncan); c. 1 km 
N of Wee Jasper, NSW, on Careys Caves Rd, in 
limestone scrub (2. AMC142961, 20 Oct 1980, P.H. 

Other Material 

Entrance to Devil's Coachhouse, Jenolan Caves. 
NSW. litter (2. QMM017295. 13 Dec 1979. J. 


Shell diameter 2.98-3.79 mm (mean 3.30 mm) 
with 3 3/4 to 4 1/2 (mean 4 1/8) loosely coiled 
whorls, last descending more rapidly. Apex and 

earlv spire (Fig. 1 1 b) verv slightly to moderate- 
ly elevated, SP/BWW ratio G.2 1-0.64 (mean 

0.33), height 1.58-2.17 mm (mean 1.84 mm), 
H/D ratio 0.48-0.61 (mean 0.56). Protoconch of 
1 3/8 to 1 5/8 whorls, mean diameter 727.3 |xm 
at 1 1/2 whorls. Apical sculpture (Fig. lOld) of 
thin, curved, regularly spaced radial ribs, crossed 
by low spiral cords. Postnuclear sculpture (Fig. 
101c) of numerous large, strongly protractively 
sinuated radial ribs. 43-82 (mean 59.1) ribs on 
the body whorl. Ribs/mm 4.17-7.95 (mean 
5.77). Microsculpturc (Fig. 1 01 e) of fine radial 
riblets, 6-12 between each pair of major ribs, and 
fine crowded spiral cords which are higher at 
their intersection with the radials, producing a 
beaded effect. Sculpture continuous on the base. 
Umbilicus (Fig. 101b) narrow, D/U ratio 4.78- 
7.56 (mean 5.98), diameter 0.41-0.74 mm (mean 
0.56 mm), reduced to a lateral crack, or closed 
by the coiumellar reflection. Sutures impressed. 
Whorls rounded above and below the periphery 
(Fig. 101c). Aperture ovately lunate. Lip 
sinuous, thickened at the coiumellar edge. 
Columella dilated, reflected over the umbilicus, 
retracted at the suture to form a small sinus. 
Parietal callus developed. Colour yellowy- 
brown. Based on 22 measured adults. 

Epiphallus (Fig. 102a) weakly differentiated 
from vas deferens, entering penis apically 
through a pore surrounded by fleshy thickenings. 
Penis (Fig. 102b) cylindrical, without a sheath, 
internally with two slender equal longitudinal 
pilasters and an apical spongy thickening which 
may act as a verge, otherwise pustulose. Penial 
retractor muscle inserting on the penis head. 
Vagina tubular, about as long as the free oviduct. 
Atrium relatively long. 

Radula (Fig. lOlf) as for genus. 

Based on one dissected specimen 
(AMC 144200). 

Ranch and Habitat 

Confined to a series of isolated limestone out- 
crops on the western edge of the Great Dividing 
Range, from north of Goulburn to north of Tarn- 
worth. NSW; under logs and rocks and among 
soil and litter. 


Various populations of C. carlessi show some 
notable variation in shell characters (Table 20). 
However the patterns arc largely mosaic and 
probably reflect differences in isolated popula- 
tions. In contrast, specimens from the Tuglow 
Caves (AMC 1 24279) have a more elevated spire 


and more open umbilicus which arc uncorrected 
changes that may indicate speciation trends in 
ini populations, 

Elsothera Iredale. L933 

Elsothtra Iredale. W3, p. 53, I red A*. 1937a, p 
Iredale, 1941a, p. 2ft7; Kt-rshaw, I95fm, p.UO; 
Burch, 1976. p. 132. 

Type Species 
HcIlx sericatula Pfeiffei, IK50, by original 


(01 ■.% Studies 
Ehothera was introduced for Helix sericatula 
PfcitTt r. I8S0 from the Svdnev region, NSW. 
Iredale (1937a, 1939) added 9 species from 
southern Australia, and Kershaw [1956a] 
another 3 from Victoria based on shell features. 
A number of generic groups may be represented 
in Elsothera but this matter is beyond the scope 
Ol The present study. Elsothera is redefined to 
include //. sericatula and two species from 
northeast NSW and SEQ. 


Diameter ),96-&64 mm, with an average of 4 
14 to 4 3/4+ normal to loosely (genithecata) 
coiled whorls. Apex and curly spire flat 
(geniihecata) to slightly elevated Apical sculp- 
ture of CTOWdcd, weakly euivcd, high radial ribs 
and finer, more crowded. low spiral cords w hich 
Continue Omtt) the radial ribs. Radials * ague to 
absenton initial part of protoconch. Adultsculp- 
ture of numerous, verv crowded, high, protrac- 
tivcly sinuated radial ribs MiCrOSCUtplure of 
fine, high radial riblcts and equally high, narrow, 
mors wulcl\ spaced spiral cords, t Jmhtl icus 
closed 10 barely open (nauiilodca). Sutures im- 
pressed to distinctly channelled (geniihecata). 
Whorls shouldered above and rounded below a 
weakly compressed periphery, last inflated and 
sometimes (nnunloiiea) with a weak supra- 
peripheral sulcus. Lip simple, columellar margin 
reflected over the umbilicus. Colour light to 
dark-brown Wfth darker radial streaks. 

Terminal male genitalia with long, mus- 
culansed cpiphallus, sometimes partially 
sheathed [eenithecata), entering penis through a 
simple poic sUtTOUnded by a collar. Epiphallus- 
\ as deferens junction swollen, complex with vas 
entering laierallv. Penis short, tubular with api- 
<ulb, internal I v with a spongy or fleshy collar 
o restricting I he entrance to the main pen is cham- 

ber. Fenial bulb with numerous short, transverse 
(\cncatula) or longitudinal (geniihecata) 
pilasters. Penis proper with short, longitudinal 
thickenings and pustulations (geniihecata), or 
Onl) pustulations (sericatula). Vagina short, or 
long with an apical caecum {geniihecata). 
Radula with tricuspid central and lateral teeth, 
central smaller; laterals with very large broad 
lanceolate mesocone, 


Elsothtra has a wide distribution from just 
norfh of the NSW, QLD border to south of Syd- 
ney. NSW. It is one of the few groups of east- 
coast charopids which inhabits drier, open forest. 
E. seruatula inhabits wet and dry sclerophyll 
B( and /■. genithecaia, has been found in 
'open eueahpt forest' in the Upper Richmond 
Range. NSW. Ability to adapt to drier environ- 
ments gives Elsoihera the opportunity to exploit 
a vast amount of relatively snail depauperate 
habitat. The genus is quite probably more 
widespread than is indicated by the coverage 
presented here and detailed statements about 
distribution are inappropriate at this time. Life- 
styles of the three species reviewed are diverse, 
probably reflecting the overall adaptability of the 


lAsuthcra has a shell with average number of 
whorls and numerous, weakly, protectively 
sinuated radial ribs. In contrast the conchologi- 
callj similar Cralopa has fewer whorls and 
fewer, bolder radial ribs which are strongly 
protractively sinuated. Radular characteristics 
of (he two genera are similar. The constricting 
penial collar, vague penial pilasters, complex vas 
dcferens-epiphallus junction, epiphallic sheath 
and vaginal caecum of Elsothera are all sig- 
nificant differences from the Cralopa pattern of 
simple penis with large longitudinal pilasters, 
bare!} differentiated cpiphallus, and lack of ad- 
ditive structures. 


Elsothera is most likely a rainforest-adapted 
group which has managed to colonise drier 
forests. The large brown ladialls ribbed shell 
and unilobcd elongate kidney are characteristics 
of other rainforest genera such as Gyrocochlea 
and Nautiliropa. Large size, combined with a 
complete seoottdan ureter were probably im- 
portant preadaptations which have enabled £7- 
\othera to exploit drier habitats 


s $£ Elsothera sericatula (Pfciffer. 1850) 

I If (Ties 104-105; Tables 19,21) 

ffel& serfcaialttWti&Wi 1850, p. 127; Reeve, 1852, 

^ ^ pi. 132, sp. si:, 

ll* II Elsothera sericatula (Pfeiffer); Iredale, 1933,p. 53; 

fli : i i || \- \ i Iredale. 1937a, p. 324; Iredale. 1941a, p. 269, fig, 

is "51 3= '3"" 

£ | § fs j§s 5 §1 $ a i ■ I Comparisons 

; * A full treatment is deferred until a more com- 

Sj prehensive review of the genus is completed. E, 

s E sericatula resembles E. hautilodea, but differs in 

|l |l ^ having the umbilicus closed (Fig. 104b) rather 

= E 

than slightly open (Fig. 106b). T EJ biretracta 

(Mousson, 1869) from the Illawarra region, 

? ^5 g* =a NSW is conchologically similar to E. sericatula 

, , , and may prove to be conspecific. Species of 

Cralopa are similar in having a closed umbilicus 
and prominent radial sculpture but are smaller 
with fewer, bolder ribs on the body whorl. The 
lectotype was selected from a lot of syntypes in 
the British Museum. 



I 5? 


|I W |- I| If s-- Type Material 

iiiil s II '21 b- 11 i i LECTOTYPE: BMNH 1987055. Australia, H. Cuming 

§2 £ - Collection. 

g Parai hctotypfs: BMNH 1987055, 3 specimens, 

I 9 ^ is £p 

same collection data as lectotype. 

Description (of lectotype) 

Shell small, diameter 4.64 mm, with 4 1/2 
normally coiled whorls, the last descending 

z- - 

SP/BWW ratio 0.1 1, height 2.39 mm H/D ratio 

Js ^2 §3 more rapidly. Apex flat, spire slightly elevated. 

I Protoconch of I 1/2 whorls, diameter 739 jxm. 

I Apical sculpture eroded with traces of radials 

I" £e IS l! If near * ne Stiturcs (paralectotypes show a pattern 

g a j: of high, slightly curved radials and low spirals). 


Post nuclear sculpture of numerous, closely 

spaced, prominent, protractively sinuated radial 

i I jg ribs, 190 on the body whorl. Ribs/mm 13.02. 

--• $3 «J « + 1 a ? * Microsculpturc of fine radial riblets, 5-6 be- 

* *S. *5 3 'z * * 5 tween each pair of major ribs, and low crowded 

spiral cords. Umbilicus closed by reflection of 
the columellar margin. Sutures impressed. 
Whorls shouldered above and rounded below a 
slightly laterally compressed periphery. 
Aperture ovately lunate. Lip simple, columellar 
margin slightly dilated reflected over the urn- 
|§ *■*!!§ ^E |l bilicus. Parietal callus strongly developed. 
itiii I5 ^1 Colour light-grey with darker irregularly spaced 
radial markings. 

3 < 

3 5 

2 5 

sag ,1 llin c 

« -.*- 









S " m 


f ' ■,"*?* 





FIG. 105. FJsothera sericaiuia (Pfeiffer, 1850). Clwe Park, Northbridge, NSW. AMC152205. a, genitalia; 
h. details of vas deferens - epiphallus junction; c talon and carrefour; d, ovotesiis; e* details of penis interior; 

I, pallial cavity. Scale lines as marked. 

FIG. 104. F.lsnihcm scriamrfa (Pfzifter, 1850). a-c. Australia. BMNM 1987055, leciulvpe; c,g-h, Clive Park 
Northbridge, NSW. AMC152205; e-F. Shark Is, Sydney. NSW. AMC103604. a-c, entire shell; d, jaw; e. 
apical sculpture; t". post nuclear sculpture; g, central and lateral teeth: h. marginal teeth. Scale lines as marked. 



Other Material Studied 

Shark Is, Sydney NSW (13, AMC 103604. ex J. 
Braziercoll); Clive Park, Northbridge, NSW, in fairly 
dense bush (3. AMC152205, 19 Oct 1969, W.F, 
Ponder, D.P. Fairfax). 

Anatomical Diagnosis 

Terminal male genitalia with a muscular 
epiphallus (Fig. 105a) which is partly coiled 
before entering the penis apically. Vas 
deferens/epiphallus junction (Fig. 105b) com- 
plex with vas deferens entering epiphallus 
laterally through a large cup-shaped pilaster. 
Penial retractor muscle inserting at the epiphal- 
lus/penis junction. Penis (Fig. 105e) short, 
tubular with an apical bulb, internally with a 
simple epiphailic entrance (Fig. 105e) and 
numerous short transverse pilasters. A large 
spongy collar constricts the entrance to the 
lower penial chamber which has fine spongy 
pustulations arranged in vague longitudinal rows 
(Fig. 105e). Vagina short with fleshy, pustular 

Radula (Fig. 104g,h) with tricuspid central and 
lateral teeth, central smaller. 

Based on 2 dissected specimens (AMC- 

Range and Habitat 

Closed and open forest from Cronulla to Gos- 
ford, NSW, extending westward to at least the 
Great Dividing Range. E. sericatula is a com- 
paratively adaptable species and still survives in 
disturbed areas where microenvironments are 
suitable. It lives in the litter, among leaves and 
accummulated debris. 


E. sericatula has been reported from Victoria 
(Gabriel, 1 930) and Tasmania (Petterd and Hed- 
ley, 1909). However more recent surveys of 
these regional faunas (Smith and Kershaw, 1 979, 
1985) do not list this species and I regard the 
earlier records as erroneous. 

Elsothera nautilodea (Cox, 1866) comb. nov. 
(Figs 106-107; Tables 19, 21) 

Helix nautilodea Cox, 1866a, p. 47. 

Helix nautiloides Cox, 1866b, p. 696; non Ferussac. 

Helix inusta Cox. 1868, p. 13. pi. 10, fig. 3. 

Helix nautilodes (sic) Iredale, 1937a, p. 324. 
Elsothera inusta (Cox); Iredale. 1 937a. p. 324; Iredale. 
1941a, p. 269. 


E. nautilodea is conchologically similar to E. 
sericatula. However the species are allopatric 
and misidentification in the field is not likely. E. 
nautilodea has a brownish shell with very nar- 
row umbilicus whereas the shell of E. sericatula 
is light grey with darker streaks and a completely 
closed umbilicus. E. genithecata is larger with 
relatively fewer whorls, more widely spaced ribs 
and channelled sutures (Fig. 108a). Cralopa 
stroudensis is smaller with fewer and bolder ribs. 

Previous Studies 

Helix nautilodea appeared on 1 January, 1866. 
In the same year, the same article appeared in the 
Proceedings of the Zoological Society (April 
24, 1866) with the name changed to H. 
nautiloides. Subsequently Cox (1868) intro- 
duced Helix inusta as a replacement name for H. 
nautiloides after realising that the latter was 
preoccupied. I consider nautilodea to be the 
correct name for this species as there was never 
any indication by Cox that H. nautiloides was 
an emendation of the earlier name. The lec- 
totype was chosen from topotypic material in the 
Cox collection. 

Type Material 

LECTOTYPE: AMC 1 36903, Clarence River, NSW, ex 
Cox. Height of shell 2.81 mm, diameter 5.24 mm, 
H/D ratio 0.54, whorls 4 3/4-. 

Other Material 

Clarence Rv (2, AMC152170. ex C.F. McLauchlan); 
Grafton {2. AMC152171, Jan 1904, S.W. Jackson); 
Clarence Rv (6, AMCI52172, G. Thornley); South 
Grafton (3, AMC 152 173. ex Nat. Mus. Vict.); Har- 
wood Id, Clarence Rv (73, AMC152174, Feb 1961, 
A. A. Cameron); North Arm, Clarence Rv (9, 
AMC152212); on E side of tree 5 ft from ground 
Harwood Is, Clarence Rv, NSW (4, AMC152218, Jan 
1970. A.A.Cameron). 


Shell small, diameter 4.05-5.24 mm (mean 
4.43 mm), of 4 1/4 to 4 3/4- (mean 4 3/8+) 
normally coiled whoris. Last whorl descending 
rapidly. Apex and spire (Fig. 106c) almost flat, 
SP/BVVW ratio 0.02-0.12 (mean 0.09), height 
2.17-2.81 mm (mean 2.41 mm). H/D ratio 0.50- 
0.57 (mean 0.54). Protoconch of 1 1/2 to 1 5/8 
whorls, mean diameter 7 1 8.6 |xm at 1 1/2 whorls. 
Apical sculpture (Fig. 106d) of curved radials 
and low, crowded spirals. Post nuclear sculpture 
(Fig. 106e) of numerous, closely spaced, 











FIG. 106. Elsothera nautilodea (Cox, 1866). a-c, Clarence River, NSW. AMC136903, lectotype. d-e, 
Harwood Island, Clarence River, NSW. AMC152174. a-c, entire shell; d, apical sculpture; e, post nuclear 
sculpture. Scale lines as marked. 









1 Z3-Z 

1 54 °E 

FIG. 107- Distribution of EUotficra naatilodea (Cox, 
1 866). 

prominent, prolractivelv sinuated radial ribs, 
129-190 (mean (51.9) ribs on the bod\ whorl. 
Ribs/mm 1(102-1 1.53 (mean 10-94). Micros- 
culpture (Fig, Mine) of crowded spiral cords and 
thread-like radial nblets. <M3 between each pair 
of major ribs. Umbilicus (Fig IQf»b] a lateral 
crack to narrowly open, rarely completely 
closed. Sutures impressed- Whorls rounded 
below and strongly shouldered above a some- 
what laterally compressed periphery w ilh a weak 
subsutural sulcus formed in some specimens 
(Fig. lilbc). Aperture roundly lunate. Lip 
simple, columella expanded and reflected over 
theumiblicus in some specimens. Parietal callus 
well developed. Colour brown with darker 
radial streaks. Based on 11 measured adults, 

"Animal almost black, not quite, tentacles well 
dc i niv'j and same colour as animal. Foot lighter 
in colour". (Based on field notes by A. A. 
Cameron. AMC 1 52 174). No soft parts available 
For dissection. 

RANr.F AM") I lAHII \| 

In spite ol intensive collecting in northern 
NSW ovei six years, L. nauiih>dai has not been 
Ideated. Earlier collections indicate the species 
on Hanvood Island, at the mouth of the Clarence 

River. Live specimens have been taken scmiur- 

Elsatherd genithecalu sp. now 

(Figs 108-1 10; Tables 19.21) 


Latin tiiccatus. encased; referring to the 
sheathed epiphallus. 


E. genilhecata is not easily confused with sym- 
patic charopids. Large size, small number of 
whorls, channelled sutures, well developed aper- 
tural sinus and closed umbilicus are not com- 
bined in other northern NSW charopids. E. 
fuiuiitodea is smaller, has a less inflated body 
whorl and lacks sutural channelling. 
Gyrocochlea find Nautili fopa which have gross- 
ly similar adult sculpture, have a very depressed 
to sunken spire and wide L'-snapcd umbilicus. 

1 YFFMATfri \i 

Houm rr. AMCI52 105. Byangum, northern NSW, 
collected by Lower, ex Cox. Height of shell 2.77'mra. 
diameter 5. fo mm. H. H r&TJO 0.49. whorls 4 1/4, 
P\r\iap(-s: AMC153717, 2 specimens, same data 
as hfllOiypC; Red Scrub Flora Reserve. Whian Whian 
S.F.. No! ! isntfu* NSWC:- 3S"S. I 5V10 E) 210 m. 
Iii(er(:. AMC 153-71& tS MTaj 19%, P.H. Colman, I 
Loch); Ml Warning N.F., NSW, near base, (2.S ; 24'S, 
153MfVE) NAT ptfim (2, QMMOI0484, QMMO- 
10502. 19 Mar l l 'Sl. AM QM - ABRS); Richmond 
Range, S.F.. NSW (28 a 39'& !52 a 43 T E) 400 m. open 
L'ucal\p[ loreM (1. QMM0f£67, i: > Apr 1976, M.J. 
Bishopi; Big Scrub, Whian Wbjaii s.K. N rvsw 
(2ff s 3S*S, l53*ltf"jn"Ej SNVF under logs {), 
0MMO16772. 15 Nov [983. AMQM - ABRS); Ml 
l .urihorinc, SFQ. in floor litter in rainforest 
(AMC1S2164. It Sept 1979. P.H. Colmun): FJppei 
Tweed Rv. NSW 1 2. AMC. '152167. Peneni! es Cox); 
1 migrant Ck, Richmond Rv. NSW (4. AMQ5216 (, 
ex Cu\:i. B\ ran B;iv Scrubs. NSW {\, AMCI52I69, 
19(1(1. SAW Jacks,. n). 

(.)1]!FR MATf-Rl \I 

Fewer Ballanon Falls Circuit. Binna Burra, 
Lan-iingumN.P.ShQ NVK litieM I . QMMO 1 o ' 
30 Sep' WR5« 1 ManiMC. O. Potter. J Chaseling): 

FIG. IDS. Eictsilwta g&ttikecattl sp. nov„ a-Cj Byangurm NSWi AMC152)b5. holoivpe; d, g-h, Richmond 
i .NSW. OMM06267,paTalype;e-f,MlWarning.N.P,.NSW. QMMO'lUMlZ.pankl.yne. a-e. enure 
shell; d. marginal teeth; e. apical sculpture; f. post nuclear sculpture, g-h, central and lateral teeth. Scale lines 
as marked. 






*^- j 





) ^1 







FIG. 109. E Isothera genithecata sp. nov. Big Scrub, Whian Whian S.F., NSW. QMMO 16772, paratype. a, 
genitalia; b, penis interior; c, talon; d, ovotestis; e, pallial cavity. Scale lines as marked. 



1 5 2 9 £ 



FIG. 1 10. Distribution of Elsot he ra ^ftufhccalu sp. 

Clarence River, NSW (emir!) (9, AMCIDMGO, ex 



Shell large, diameter 4,37-6.64 mm (mean 
mm), with 4 to 4 5/8- (mean 4 1/4) looseU 
coiled whorls, the last whorl descending more 
rapidly and strongly inflated. Apex and spire 
flat. height 2.77-3.36 mm (mean 2.94mm). H/D 
ratio 0.47-0.63 (mean 0.53). Protoconch of 1 
12 whorls, mean diameter 821.3 n.m. Apical 
sculpture (Fig. 108c) of high, slightly curved 
radial ribs, and low crowded spiral cords. 
Postnuclear sculpture (Fig. lOSF) of numerous 
crowded, high, protractively sinuated radial ribs, 
111-153 (mean 132) ribs on the body whorl. 
Ribs/mm 5.83-8.64 (mean 7.24). Microsculp- 
ture (Fig. 1081} of fine radial riblets, 5-10 be- 
tween each pair of major ribs, and crowded spiral 
cords which continue onto the apices of the 
major radials. Umbilicus (Fig. 108b) closed or 
a very small lateral crack. Sutures deeply im- 
pressed. Whorls shouldered above and rounded 
below a slightly compressed periphery (Fig, 
lOSc). Aperture roundly to ovatcly lunate. Lip 
sinuous, simple, retracted at the suture to form 
an apertural sinus (Fig. lOSa-b). Columella ver- 
tical and thickened, reflected over the narrow 
umbilical opening. Parietal callus weakly 
developed. Colour light-brown with irregularly 
spaced dark radial streaks. Based on 4 measured 

Genitalia with vas dcfcrcns-epiphallus junc- 

tion complex; vas deferens entering almos! 
laterally through a large spongy pilaster (Fig. 
109a). Epiphallus (Fig, 109a) long, convoluted 
with a large muscular sheath for the last half of 
ils length, entering penis apically through a 
simple pore (Fig. 109b) surrounded by a circular 
collar. Penis (Fig. 10%) short, tubular with an 
apical bulb. Apical bulb internally with short, 
longitudinal pilasters and a transverse spongy- 
collar constricting entrance to lower penial 
chamber. Lower penial chamber with numerous 
short longitudinal thickenings (Fig. 109b). and 
irregular pustulations nearer the atrium. Penial 
retractor muscle inserted on the epiphallic 
sheath. Vagina (Fig. 109a) long with a caecum 
containing a large internal spongy thickening, 
and an elongate basal extension. 

Radula fFig. 108d.g,h) with expanded 
mesocones on central and lateral teeth. 

Based on 2 dissected specimens (AMC- 
152165, QMM016772). 

Ranof and Hahu m 

Warm moist subtropical noiophyll vine 
forests of the Big Scrub area between the Rich- 
mond and Tweed Rivers. NSW; cooler subtropi- 
cal forests of the Lamington Plateau, SEQ; palm 
dominated forest of Mt Tamborine, SHQ; and 
drier scrub of the Richmond Range. NSW. It 
lives under logs. 


E. genitkecaia is one of the largest charopids 
in subtropical rainforests. The channelled su- 
tures are an unusual development also seen in 
Ngairea canaliculata (Fig. 12a. e) and a numbei 
of extralimital taxa. The significance of this 
feature is not known. The well-developed aper- 
tural sinus is shared locally only with Cralppa, 
but is more common in New Zealand charopu!-.. 

Coenocharopa gen, nov, 


Latin cocno, dirt: referring to adhering dirl 
particles on shell. 


Coenocharopa sordidly sp. nov 


Shell diameter 1.91-4.43 mm, with rttfi 
whorl count 4- to 4 3/8-. Whorls normally 
coiled, last descending. 


Apc\ and spire slightly (ntaltitadiata, yes- 
\LihiiJicttsi*) to conspicuously (a lata) elevated. 
Apical sculpture of weak (\rs\ahahc/i\t\) to 
piominent, curved radial ribs which may be 
crowded to moderately crowded ialaia), more so 
;ti the nuclcar-postnuclcar junction, or regularly. 
Wldelj spaced {tmdnrudiaia). Ribs with 
prominent pcriostracal blades. Secondary apical 
sculplure of crowded, squiggly spiral cords 
which are not continuous on r he radials. 
nut K-arsculpiurcofwidely spaced (a/t/m) to 
crowded (yessaonhensis) weak radial rihs with 

rj ■uiucni periosteal blades modified into 
wing-like processes in some species (alata). 
rarely lacking (ycwahahcnMs). Mieiosculpturc 
of thread-like radials which mav have sironglv 
developed penostracal blades {tntditradiaia) and 
low, broad [alata) to high, narrow [inidhnsdtata) 
spiral cords. Din particles adhering, to shell sur- 
t'aee. Umbilicus narrow to moderately wide 
(irmcr /n{./lnda). U-shaped. Sutures impressed, 
whorls strongly lo weakly shouldered, oe- 
rasionallv with a supiaperiphcral sulcus (macm- 
mphala). Aperture roundly 10 ovalelv lunate. 
Lip simple, margins convergent Parietal callus 
strongly developed. 

Animal with fool and tail broad, bluntly 
Llwered posteriorly. Terminal male genitalia 
with slender to expanded (sortHdux} vas 
deferens. Fpiphallus sometimes partially bound 
lo penis sheaih {mucmmphulu. alula), entering 
penis apicallv through two fleshy thickenings. 
Penis large lovetj large [sardldugl, tubular, with 
a filTCath, internally with several longitudinal 
pilasters which arc modified in some species 
[sordtdus, ycssuhaiwnsis). Pen ial retractor 
muscle inserted on or very near io (he fpiphal- 
lus penis junction. 

Radula with mesocone ol the laierals greatly 
I -Miiiled in C \\. 

Oisiuiiu iim\ v\o Ecoi uov 

Warm temperate rainforests and thickelsol the 
N'essabah-Carrai limestone outcrop, cool sub- 
tropical noiophyll vine lorests i)\' the Great 
Dividing Range (from Dorrigo to the Border 
Ranges) and McPhcfton Ranges, and warmer 
subtropical notophyll vine forests (wiih 
\>,iurt'rm) in southern (Queensland. 

Inirageneric sympalry is complex when com- 
pared \\i!h other charopids examined in this 
sludy. Sympatric combinations are as lollows 
C, WrtUdtfSfl tttucromphala f. sorduJusC. 
panicosiara. ( purvtco\lsttu ('. nuuromphala. 
( . >df_aa i .ntaltuuditttaA'.midiitadiaiai.par- 

i iiosiaw, C. muhiradkitafC, ye&s&bakonsis and 
possibly C aluiu'C, pur\ic<tsiuia. No details of 
microsympatry are available although this is 
likely in the case of ( sordidus and C. itutcrtt- 
tnphala. t . \ essabah&nste has an enlarged penis 
and inodilied pemal siulace structures which 
indicate a mierosympatrie congener, probably < 
muliinidmui. Although sympatric with both C. 
muitiradiata and C pumcoslala C alata 
probable not sympatric at the microhabitut level. 

P\i !i-u\soi \ \\m\rui\ 

Shell size and shape - Whorl counts correlate 
positively '••• ilh increased sr/c i-'scepi in G fttul* 
iiradiata. Spire protrusion is proportionately 
giealest in ( . ahifa. Umbilical width is uniform 
except in C. mm nvttpha I a where it is wider (Fig. 
I 14c). 

Shell ScMptUre The prominent calcified 
sculpture of most ground-dwelling charopids 
serves lo lessen the chance of particle adherance. 

Positive coi relation between wider rib spacing 

and increased height of pcriostracal extension 

suggests that in Cocnochaiopa the periostracal 
elements may be compensating for the lack of 
primary rib sculpture. Nonetheless mosi species 
have their shells covered by a layer of adhering 
dirt parlicles Reduced sculpiurc in ( yes- 
tabahensts correlates with living on rock sur- 
| i b tt hCTC adherence of dirt particles mnv noi 
he a major problem. ( '. muitiradiata (Fig. I2bd- 
f) most closely mimics the more lypieal sculp- 
ture as in Rhophodon and Gymcochlca. 
Significantly it ts the 'cleanest* species of 
( (K'lUH inintpa. 

Genitalia - Variation in reproductive anatomy 

is confined |o terminal male genitalia. 

In Ng&iftfc Gyrocochlea and Rhophodon 
vanalion in pcnial surface among mierosym- 
patcrs involved minor changes in features of the 
apical chamber ol the penis together wiih small 
changes in shape and size ol the longitudinal 
pilasters. In con I r a>i a lie rations in 

0\ nocharqpa involve major -dupe changes in 
pilasieis. The basic patieni of internal peuial 
sculpture appears to be several longitudi 
pilasters as in C Jnacwmphala (Fig. II5d). C. 

\ordidu\, which is probably microsyrnpatric 
with this species, has a large U-shaped pilaster 
and spvcral accessory longitudinal thickenings 
(he. li:a). 

Insi uiion ol the penial Tetractor muscle dis- 
olavsonly minor shifts from the penis/epiphallus 
junction onto the epiphallus adjacent to the junc- 
tion. The epiphallus is typical, but in C. macro- 


- s _ - n = mphala and C a/fl/fl is partially bound to the 

penial sheath (Figs. 115a. 118b). 
■: 1 -] sg jg S§ jj| Radula - The enlarged mesocones of the 

central and lateral teeth of C.yessabahensis (Fig. 
123d,h) resemble those in the sympatric 
Letomoia conforms and Rhophodon kempseyen- 

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Coenocharopa is distinct among genera 
reviewed in having a basically 'smooth' shell 
with strongly developed periostracal sculpture. 
Apical sculpture resembles FAsothera and 
Cralopa but these genera differ in having very 
i prominent radial ribs on the postnuclear whorls, 

and closed or nearly closed umbilici. Letomoia 
has greatly reduced sculpture but is much 
smaller, has modified apical sculpture, wide um- 
bilicus and several apertural barriers (Figs 67, 
68). In the Mudlo Gap area, north of Gympie. 
SEQ .the conchologically similar Lenwebbia is 
sympatric with Coenocharopa. However. Len- 
webbia is distinguished by its modified apical 
sculpture which consists of irregular pits and 
wrinkles (Fig. 29g), and postnuclear sculpture 
which includes incised spiral grooves (Fig. 29e ). 
Coenocharopa shares some genital features with 
Cralopa. These include a tubular penis, simple 
epiphallic entrance into the penis through large 
fleshy thickenings, and basic penial surface 
sculpture of longitudinal pilasters (fewer in 
Cralopa) and background pustulations. 

The postnuclear sculpture in Coenocharopa is 
deciduous and often lacking in older and dead 
specimens. The following key is based on new- 
adult fresh shells and may not work for gerontie 
or litter-retrieved specimens. 


1. Shell with prominent wing-like periostracal 
processes (Fig. M7e.h); spire strongly 
elevated (Fig. 1 1 7c) aiata 

Shell without prominent wing-like periostra- 
cal processes; spire weakly elevated _ 

2. Apical sculpture of regularly spaced radial ribs, 
not crowded at the nuclear-poslnuclear 
boundary (Fig. I26d) mulliradiaia 

Apical sculpture of curved radial ribs which 
become crowded at the nuclear-postnuclear 
boundary (Fig. 1200 3 



.vPnsinuclear sculpture of regular, widel\ spaced 
radial rih$ wilh periosiracal blades (Figs 
1lle.ll4eJ20e) 4 

Poslnueleur sculpUue of low, crowded radial 
ridges without prominent periosteal blades 
1 1 ig. 1 2 ?e.g) yrssahahen^is 

■It larger, mean diameter 3.92 mm; apical 
sculpture ol" weak thread-til. e radials (Fig. 

mO tnrdldWk 

Shell smaller, mean diameter <3,00 mm; 
apical sculpture of low, prominent curved 
radial ridges (Flg4 II 4f, 1201") 5 

5. Umbilicus narrow. V-shaped, mean D/U ratio 
4 Mv, shell colour glossy yellow-horn 
., ., par\ia) , ;tatii 

diameter 4.43 mm. H/D ratio p.59, U U rAUo 3.17, 
whorls 4 1/2+. 

Mt ) l 1 575,$ specimens, same data as holotype; Freds 
Rd. MlMee(27 : (»5 S. 152 Z 45"E) ramtoresl. leal II 
[X QMM09696, 14 Apr 1980, J. Stanisic. A. Green. 
\ \\,\\\): O.ivhoro - Mr Byron Rd. at crossing of North 
Pine R)V<t(27'15*&. l52 ;, 47'E;Ueut :Iitier(l t QMMO- 
17277. 2? Apr 1980, J Stanisic); Mt Guyra N.P 
(25 4s»'S. r52*3S"E) NV?/Araucaria (U OMMO- 
12073, 7 Sept 1982, AM/QM-ABRS); c. I km from 
R ,\\ nhi rdC reek Rd. on Lacey's Ck Rd. Laceys Creek, 
open scrub, litter (I. QMM016S34, J, StHJltSJC, G. 
AnnflbclL 2 Apr 1984); t\ 1 1.3 km NE ofDandabah 
on Bunva Ml* Rd (26*50'S, 15T33"E) (1. QMMO- 
i !30 i S , 1982, AM/QM-ABRS); Upper Brook- 
field, NVT Arum ana, lilter ( 1. QMMO 17271. 3 Mar 
L081 V. DavjftS. R Ka\en; I. QMM0172hS. 1 I Dec 
1980. V.Davics. R. Raven). 

Umbilicus broader, V-shaped, mean D/U 
ratid 3 14; shell colour hurnishtd-yetlow ru 
beige mu, mmphala 

Cocnocharopa sordidus sp. noV. 

(Figs 11 1-1 13; Tables 22, 23) 

Etymol oca 
Latin sordidus, dirty: fur the dirty shell. 


C. lOrdidus is similar to C. macromphala but 
is distinguished hv lis larger size, narrower um- 
bilicus (D/U tatio 3.56 compared with 3.14 for 
C, macromphala) and much finer sculpture. C. 
sordidus (Fig. lllf 3 g) has apical sculpture of 
vague radial ridges and crowded spiral cords 
extending onto post nuclear whorls. C. macro- 
mphala (Fig. 1 14 f.g) has prominent curved api- 
cal radial ribs and more widely spaced spiral 
eouls on apex and post nuclear whorls. 
Anatomically C sordidus is distinguished from 
C macromphala by its larger terminal male 
genitalia which include a long cpiphallus and vas 
deferens, and greatly expanded penis with 
modified pilaster pattern (Fig. 112a). C. par- 
vicostata which is also sympatnc with C. sor- 
didus, has a more rounded, smaller shell, less 
elevated spire, smaller umbilicus and mure 
prominently sculptured apex. 


Holotype: QMMOI7276, Upper Brooklield, SFQ 
(27 C 30'S. 152"55'E) N\¥h\rmtcanu, litter. Collected 
h\ V. Davies. R. Raven. S Apr I 98 I . Heighl 2.h0 mm. 


Gold Creek Reserve, Upper Brooklield, SEQ, closed 
forest, liner (I, QMMO 17278, 16 Dec 1980. V. 
Davies. R. Raven); Kcnilworth S.F.. SEQ (26*35 *S, 
152 42 F)NVF(2,QMMOol52. (8 Mb) 1976, MJ. 
Bishop |. 

DtAGW i 

Shell smalt, oftep completely covered in dirt, 
depressed, diameter 3.41-4.43 mm (mean 3.92 
mm) with 4+ to 4 5?$+ (mean 4 ?>/&-) normally 
coiled whorls Apex and spire moderately 
elevated. SP BWW ratio 0. 16-0.30 (mean 0.2o j : 
height 2.09-2,64 mm (mean 2.31 mm). H/D 
ratio U.55-0. 61 (mean 0.59), Protoconch of 1 
1/2 to 1 5 8 whorls, mean diameter 709.9 |xm at 
1 12 whorls. Apical sculpture (Fig. I llf) of 
weak, curved radial ribs becoming more 
crowded at the nuclear - postnuclear boundary, 
crossed by very crowded, weakly defined. 
squiggly spirals. Protoconch often appearing 
smooth in worn shells. Postnuclear sculpture 
(Fig. 1 1 ig) of strongly bladed, widely spaced, 
protractively sinuated radial ribs. Microsculp- 
ture (Fig, 11 lg)of thread-like radials and incon- 
spicuous low spiral cords, which buttress both 
sides of the major radials. Sculpture continuous 
on the base. Umbilicus (Fig. 1 1 lb) narrow, U- 
shaped, diameter 1.01-1.40 mm (mean 1.11 
mm), D/U ratio 3.17-3.94 (mean 3.56). Sutures 
impressed. Whorls shouldered above and slight- 
ly flattened below the periphery (Fig. lllc). 
Aperture roundly lunate, margins convergent. 
Lip simple. Columella slightly dilated. Parietal 
callus prominent, uniting lip margins. Colour 



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FIG. 112. Coenocharopa sordidus sp. nov. Upper Brookfield, SEO. QMMO 17268, paratype. a, genitalia 
and details of penis interior; b. hermaphroditic duct; c. talon and carrefour; d, ovotestis; e, pallia! cavity. 

FIG. 111. Coenocharopa sordidus sp. nov. a-c. Upper Brookfield, SEQ. QMM01727fi, holotvpe;e-g, Fred's 
Rd, Mt Mee. SEQ. QMM09696. paratype: d, h, Upper Brookfield. SEQ. QM MO 17271, paratype. a-c, entire 
shell; d. central and lateral teeth: e, entire shell; f. apical sculpture: g. post nuclear sculpture; h, lateromarginal 
teeth. Scale lines as marked. 




26°S -J 








FIG. 113. Distribution of Coenocharopa sordidus sp. 

dirty vellow-horn when clean. Based on 7 
measured adults. 

Vas deferens (Fig. 112a) large, convoluted, 
giving rise to a more expanded, muscular 
epiphallus which is as long as the penis. 
Epiphallus (Fig. 112a) entering penis apically 
through a spongy pilaster. Penial retractor 
muscle inserted at the penis-epiphallus junction. 
Penis (Fig. 1 12a) with an enlarged upper cham- 
ber receiving the epiphallus and enveloped in a 
thin sheath; internally (Fig. 112a) with a very 
large spongy horseshoe-shaped pilaster opposite 
the epiphallic pore, and a series of short lon- 
gitudinal thickenings below. Lower chamber 
with irregular spongy pustules arranged in vague 
longitudinal rows. 

Radula(Fig. 1 1 ld.h)without unusual features. 

Based on 3 dissected specimens (QMMO- 
17268, QMM01727S\QMM017271). 

Range and Habitat 

Drier araucarian vine forests between Brook- 
field, Mt Guyra, and the Bunya Mountains, SEQ. 

It has been caught in pitfall traps set for ground 
dwelling spiders and probably lives among fri- 
able soil and litter. C. sordidus has been col- 
lected in degraded forest at Laceys Creek and 
along the Dayboro-Mt Byron Road, SEQ. These 
are areas with little rainforest that have been 
largely cleared for farming. 


C. sordidus is identified by its size and almost 
completely dirt-encrusted shell (Fig. 1 1 la). It is 
inconspicuous in the field. 

Coenocharopa macromphala sp. nov. 
(Figs. 114-1 16; Tables 22, 23) 


Latin macro-, large; Greek omphalos, um- 
bilicus; referring to large umbilicus. 


C. macromphala may be confused with C. 
sordidus but the wider umbilicus, smaller shell, 
more prominent apical sculpture and more wide- 
ly spaced spiral cords on the postnuclear whorls, 
distinguish it from that species. C. parvicostata 
(Fig. 120a-c) has more rounded shell whorls, 
flatter spire and much smaller umbilicus. Len- 
webbia protoscrobiculata is sympatric in the Mt 
Mudlo area, north of Gympie, SEQ. It has apical 
sculpture of irregular pits and wrinkles, and 
postnuclear sculpture that includes incised spiral 
grooves (Fig. 29e,g). 

Type Material 

Holotype: QMM017263, Upper Brookfield, SEQ 
(27°30*S. 152°55*E), litter, NVF/Araucaria. Col- 
lected by V. Davies, R. Raven. 8 Apr 198 1 . Height of 
shell 1 .83 mm, diameter 2.98 mm, H/D ratio 0.6 1 , D/U 
ratio 2.64, whorls 4 3/8-. 

MOl 1576. same data as holotype: Uppe,r Brookfield, 
CNVB4 mucaria (2, QMM017262, 3 Mar 1981. V. 
Davies. R. Raven); beside Bruce Hwy, Blackbutt 
Range, under logs, NVF/Araucaria (8. QMMO 1 6828, 

12 May 1984, J. Stanisic. D. Potter); c. 7.4 km S of 
Benarkin. Blackbutt Range (26°53"S, 152° 1 1 *E) (12, 
QMM01224LAMC136567.31 Sept 1982, AM/QM- 
ABRS); Upper Brookfield, NVF/Araucaria, litter (2, 
QMM017265, 28 Nov. 1980. V. Davies, R. Raven); 
Mt Mudlo. Kilkivan S.F., (26'01'S, 152M3 , E) 1 

FIG. 1 14. Coenocharopa macromphala sp. nov. a-c, Upper Brookfield, SEQ. QMM017263, holotype. d-h, 
same data as holotype. QMMO 17265, QMMOl 1576, paratypes. a-c, entire shell; d, central and lateral teeth; 
e, entire shell; f, apical sculpture; g, post nuclear sculpture; h, radula. Scale lines as marked. 







I mm 





FIG. 115. Coenocharopa mac rompha la sp.nov. Beside Bruce Hwy, Blackbutt Range, SEQ. QMM016828, 
paratype. a, genitalia; b, talon and carrefour; c, ovotestis; d, details of penis interior; e, pallial cavity. Scale 
lines as marked. 



■5 D S 



2S D S 



152 C 'E 



PIG, L Hi. Distribution uf Cncnocharopa macro- 
mphala sp. now 

NYh'AuuiCLUia. leal litter (4, CJMMOIK'-Ta, 17 Jul 

Or Hir R M Mh Hi \i. 

Coolabunia Pine Scrubs, Kingarov, SEO (L 
AM02999, Jun 1908, S.W. Jackson). 


Shell small, depressed, diameter 2.47-2.98 
mm (mean 2,69 mm) with 3 3/4- |o43/8- (mean 
4+) loosely coiled whorls, last descending more 
rapidly. Apex and spire (Fig- I 14c) moderately 
elevated SP BWW ratio 0.15-0.33 (mean (J. 25). 
height 1.2S-1. 83 mm (mean 1.57 mm). II D 
ratio U .52-0.64 (mean 0.58). Protoconch of 1 
3 8 to I I 2- whorls, mean diameter 047.8 u.m at 
I I 2 whorls. Apical sculpture (Fig- II If) of 
crowded, protractively sinuated radial ribs 
which become more crowded at the nuclear- 
postnuciear boundary, crossed h\ fine spiral 
COrds. Poslnuclear (Fig. I14g) sculpture of ir- 
regularly spaced. protraetively sinuated radial 
rihs with conspicuous periostracal blades thatare 
often worn, 211-30 ribs on the body whorl. 
Microisculpture (Fig. I !4g) of numerous low 
lliread-like curved radial riblcts and low 
crowded spiral cords. Umbilicus (Fig. 1 14b) 

broadly V-shaped, diameter 0.70-1.13 mm 
(mean 0.87 mm), D/U ratio 2.64-3.54 (mean 
3.14). Sutures strongly impressed. Whorls (Fig. 
1 14c) rounded above and below the periphery, 
with a supra-peripheral sulcus. Aperture round- 
ly to ovatcly lunate. Lip simple, columella di- 
lated and slightly twisted toward the umbilicus. 
Parietal callus strongly developed and connect- 
ing lip margins- Colour burnished yellowy- 
beige. Based on 1 1 measured adults. 

Genitalia with large, muscularised epiphallus 
(Fig. 1 15a); internally with longitudinal thicken- 
ings and entering the penis apically through a 
simple pore- Epiphallus partially bound to the 
penis sheath by short fibres. Penis (Fig. 1 15d) 
tubular, internally with low pustules in the apical 
portion giving rise to low, spongy, longitudinal 
thickenings in the penis proper. Penis sheath 
present. Penial retractor muscle long, inserting 
at the epiphallus/pents junction. Vagina mus- 
cular with low pustules internally. 

Radula (Fig. 114d.h) similar to that of C. sor- 


Based on 3 dissected specimens (QMMO- 
1687$ QMMOJ72G5), 

Rangf and Haritat 

Prefers drier arauearian notophyli vine forests 
between Brookficld and Mt Mudlo. SEQ. Like 
C sordid tis, C macromphala has been collected 
in pitfall traps and probably lives in friable carih 
and litter. Distribution is poorly known. 

Coenocharopa alat;i sp nov. 

(Figs 11 7- lls>: Tables 22, 23) 

Ervvioi 0G\ 

Latin alatu^, a wing; refers SO wing-like 
periostracal processes. 

( i ►MPAftfSGNS 

Live specimens arc recognised by then large 
Wing-Hke periostracal extensions (Fig. 117c, h) 
which are deciduous and often tost in dead 
specimens. Worn shells can be distinguished 
from C. macromphala by their comparatively 
higher spire and smaller umbilicus; from C, par- 
vivasiata b\ iheir strangle elevated spire and 
flaticr periphery: and from C, miduradiata bj 
their larger size and less regular sculpture. C 
a/ata has a more immediate resemblance to some 
species of the punctid genus Paralaoma lredale, 
19 13. That genus is much smaller and has apical 
sculpture of l<n\ , mended spiral cords. 



JRv ? ' 

. ■■;* ... „, *i 








f Ai 

/— ' 

. . . ■ 







FIG. 1 18. Coenocharopa alata sp. nov. a, c 12.2km from Acacia Plateau-Killarney Rd, Koreelah S.F., NSW. 
AMC 128563. paratype. b-e, Coombadjha Walk, Washpool N.P.. Gibralter Range, NSW. OMMO 17270, 
paratype. a, genitalia; b. details of penis interior: c. hermaphroditic duct; d, ovotestis;e. pallial cavity. Scale 
lines as marked. 

FIG. 1 17. Coenocharopa alata sp. nov. a-c. c. 12.2km from Acacia Plateau-Killarney Rd, Koreelah S.F., 
NSW. QMM017279, holotvpe; d, same data as holotvpe. QM MO 10961. paratype; e-g, Wiangarie S.F., c. 
24kmNofKyogle t NSW. AMCI50094, paratype; h, Gibralter Range, NSW. QMMOl 7270. paratype. a-c, 
entire shell; d, lateromarginal teeth; e, entire shell; f, apical sculpture; g. post nuclear sculpture; h, details of 
periostracal process. Scale lines as marked. 




V8 C S 



1 5 1 D E 



FIG. 119. Distribution at Coetwcharopa at at a sp. 



HOLDTVTE OMMO 17279- c. L22 ^m Imm ACadfl 
Plateau - Kilhmev Rd, Korcclah S.F.. NENSW, S50m. 
si-yi ^iniicaria, Collated is Mai i9$h am/om- 
ABRS. Heightol.>helI 1.70mm. diameter 2. 73mrrt, H D 
rauo0.63-D/UrauoJ.3iwhoris4 I 4-. 

Paratopes OMMOI096I, -\MCI2s563. 21 sprci- 
mens. same data ;i> holotype; Binna Hurra, Lamineton 
N.P.. SEQ. 850 m*. CNVF (28*13*5. I53M2E] 1 1. OM- 
km ME o| Kvogfe, m-asw. hum m (28"23'S, 
I53*(KtE) hTl?r<6. AMCI5QU94, 17 Maj W7fc P.U. 
Cnlman, I. Loch), MoonparSF.. NENS W„ O.ti km u i 
Mills RJ from Mnoftpai Rd imetseciiun. NSWtfQ'LVS. 
152 39'E>. under mo>s on log ( 1 , QM.Mt ) [ 73W, 7 Mar 
1987. 1, St anisic. D. Patter* Caornbadjtoi Walk. Wa>h- 
pnol N P Gibratter Range, NSW f2*3 28"S. 152 Ifi I I 
(i. QMM01727a 8 Mar Wfffc, J. Suumic. Puller): 
Gibraltar Ranee, N.P., \FNSVv, rainforest (I, 
OMMOt72f?V.G.MonleUn, [PNoA 1980). B«iuryS.F„ 
' km \\ ul L'rhenville. \NSWf7S 2^S. L52°5-TE). 

ilfl n I ami 154730; is M.n |97fi. p h. ( cvlrrw | 

I . ■ i |; Rod Scrub KR , Winn, Wlu.ui St' . NNSW 
[2N 38'S* 153'IU'EJ 211] m C. AMCk5.l7*| 15 Mtt\ 

I 170 I'H t_ , . 1 1 ■ 1 , ■ ■ i I ..,1,1 

I M ili R M.\ii-iO\l. 

rn i i. Cilhraltet fa ig i ■ I ''•-->. | " '..vs. 

152 17E) NVK. under muss on rock (1, 
QMMO ! 7272. 8 Mar 1 987, 1 Slanisic. D. Potter), top 
of Condamine River valley, alone Croflbv Rd 
(28 15'S. 152 : 29'E) (2. QMMO10541, Mar 1981 
AM QM-ABRS). Gibralter Range. NENSW, rain- 
forest, litter (1, QMMO 17275^10 Nov 1980, R. 


Shell minute. tdoss\ , depressed, diameter 
230-2.73 mm (mean 2.49 mm) with 5 3/4- to 4 
1/4- (mean 4-) loosely coiled whorls. Last whorl 
(in aduit specimens) descending much more 
rapidly. Apex and spire (Fig. 1 17c) strongly 
elevated, SP/BWW ratio 0,29-0.45 (mean 0.37), 
iic 1.36- 1.70 mm (mean 1. 49mm), H/D ratio 
AS-0.63 (mean 0,60). Protoconcb of 1 5/S to 
1 3 4 whorls, mean diameter 629.7 u_m at 1 12 
whorls. Apical sculpture (Fig. 1 17f) of curved 
radial ribs, becoming more crowded toward the 
nuclear-postnuclear boundary, and low spiral 
COTds. Postnuolcai sculpture (Figs. 1 1 7e,h) con- 
sisting of widely spaced, prom i ne m. protractive- 
ly sinuated radial ribs which have large 
wing-like pcriostrucul processes at the 
periphery; processes deciduous and not present 
in larger, worn specimens. Microsculpturc (Fig. 
I I7g) of numerous, fine, thread-like radials. 
crossed h\ low. crowded spiral cords. Sculpture 
continuous on the base. Umbilicus (Fig. 1 17b) 

■ ow. U-shaped, diameter 0.4 1-0. 82 mm 
(mean 0.60 mm). D/TJ ratio 3.32-5.60 (mean 
4-38). Sutures impressed, whorls strongly 
shouldered above and rounded below a flattened 
periphery (Fig, LI7c), Aperture roundly lunate. 
Lip simple, columella slightly dilated. Parietal 
callus strongly developed, connecting lip mar- 
gins. Colour yellow-horn. Often covered in dirt 
particles. Based on 3 measured specimens. 

Genitalia with a short, expanded epiphallus 
b< i nd lo ihe penis stuvih for its Entire li ngth 
(Fig. 1 18b] Ejiipballli* tubular, entering perns 
apici! : through a li' sh\ pilaster, internally \s itli 
longitudinal thickenings. Rental retractor 
muscle inserted on epiphallus prior to the 
CpiptutllUS penis junction, usually originating 
from the diaphragm" but in figured specimen 
I QMMO IlloOi j. arising Lorn tentacular branch 
of columella! muscle. Penis (Fig. 1 isb) tubular. 
swollen apically. with a sheath in rhe expanded 
upper portion; internal I) W ith apical pu&tulattl 
and has&l spongy longmulinal pilasters. 

Kadula(t ; i-."i 173) typical. 

BaStfXl on fcui dissected specimens (QMMO- 


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FIG. 121. Coenocharopa parvicostata sp. nov. Kenilworth S.F., SEQ. QMM017264, paraiype. a, genitalia; 
b, details of penis interior; c, talon; d, ovotestis; e, hermaphroditic duct; f. anterior edge of pallial cavity. Scale 
lines as marked. 

FIG. 120. Coenocharopa parvicostata sp. nov. a-c, Freds Rd. Ml Mee, SEQ. QMMO 17266, holotype; d-f, 
Mt Mudlo, Kilkivan S.F., SEQ. QMM08358, paratype; g, Diehappy S.F., NSW. AMC 128388, paratype; h, 
c. 8.3km from Woodenbong-Legume Rd, Tooloom Scrub, Beaury S.F., NSW. AMC 128532. paratype. a-c, 
entire shell; d, post nuclear sculpture; e, spire; f-h, apical sculpture. Scale lines as marked. 



10270, QMMO10269, QMM0172jB9, AMC- 


Cooler subtropical notophyll vine forests oi' 
the Greal Dividing Range between Dorrico, 
NSW and ihe QLD/NSW border, and the 
phyll vine forests (and possibly microphyll 
fern forests) of the McPberson Ranges, SEQ. 
Live specimens have been collected from among 
moss on togs and rocks. 


Probably St'totnedea' aculeata (Hedley 1 B99) 
COmifi closes! in shell form. However, the 
periosteal extensions of \S\ aculeata are more 
|j rtilcr. [DOre numerous ;ind regularly spaced 
along the major radial ribs. 

Coenocharopa parvieostata sp. nov. 
(Figs 120-122; Tables 22. 24) 


Latin parvus, small; and cosiuius rib; refers lo 
the postnuclear sculpture. 

I if ir 1 \uisn\s 

C. parvieostata is sympalric with C sunlnlns, 
C- mmrumphala and C. multiradiatu, and dead 
shells of these species are apl to be confused. 
HoWcVcr C. panhostatu can be distinguished 
from C. sanJidus by its smaller size, more 
founded whorls, less elevated spire and more 
prominently sculptured apex: from C. macro- 
mphaJa by the lack of a supraperipheral sulcus, 
less elevated spire and smaller umbilicus; from 
C. mulitradiata by the more numerous apical 
radial ribs which are crowded at the nuclcar- 
postnuclear boundary, and from CI alulu. by 
having a less elevated spire and more rounded 

Tmm M mi n U 

KOLOTYM 0WMGl f 72G6, Fred's Road. Ml Met*. 

rj 27'05'S, l5i"4S'E. ramforesL Hue? Collected 

h\ LSt;inisie, N. Half A. Green, 14 Apr ioso. Height 

ol shell I.5N mm. diameter 2.81 mm, H D ratio I l.5i i. 

01! r.iho3 34.\vruuis4 3 S. 

PARATYPHI Ml Mudlo. Kiiki\an ST . SEQ 
(2n HIS. I52 6 1TE), NVFMttUftWta k-jf lirier («, 
Q.MMOS35S, II Jul 1480, L Sunisic, A Greejii. 
Keml\surtnST,.SFO(2o \V"S. L52 d 4TE)rainFoi 
lcttfiiUtfr(l«QMMOl 7267.22 May mil. J. Sumisie. 
A. t..rtfii); e S.3 km from Wondenbnn.e - l.euume 
Road, Too loom Scrub. Beauts SI.. NENSW 

(28 Z9"S. i 5-2 24 l E0 I - I | ■ \\i* 1283 ■ 
OMMOHWh. 15 Mar I 'IS I , AM (JM-ABRS): Dan 
dabah.Bunya Mis NT.. SEQ (2d 53'S, lSr35 , E)(2, 
0\IMOl-.(i:S7_ 5 Mar 1976, M J Bishop); e. ro km fc 
of tickgtt* Mt Rfcaperte S.P.. NFNSW, (29*01*8, 
I52 3 45"'E) rafnfotfest (l, QMM016946, Mar tf?8] 
AMQM-ABRS); creek cutting, Diehappv ST.. 
NENSW (3o':s'S. I52'3N*E). broad leaf scrub (7, 
QMMO10X24. AMi: 1283845, Mar 1981 AM/QM- 
ABRSl; Newell Falls, DorrigQ NT., NENSW 
(3D 24 S. 152 45Ti NAT (6. QMMO107S2, AMC- 
128535, Mar 1981, AMQM-ABRS); Mudlo Gap 
ST.. SEQ [26 01 S, I52 D 14T» MW.Anutruria (5 
QMMCH2355, AAtC13G65o. 2 Sept 1982, AM/OM- 
abrsi: c. 0,5 km S or 'Reilbird' parking area, Kenil- 
vvnriliS.l ; ,.SEn.ra,nloreMlitter(1.0MMOl"2i- 1 
Feh IQgO. J. Siani.sie, A.Green). 


lillle Yahbu Creek. Imbil ST.. SEQ (26^S'S. 
I52'38*E) NAT (5, QMMO1202& AMC13M40, 8 
Scpi [9B2 3 AM OM-ABRS): Bellinger River. 
NFNSW (30°27*$, 152 Q 37'E> rainforest gully (3. 
QMMOIOS32. AMC12K39Q. Mar 1981. AM/QM- 
AliRS); Mi Warning N P . NFNSW. near base 
CSZTN. 15.1 tfi El NVETulrii-, i4. QMMO104S0, 
AMI '"1242s-.. Mar I^JM. AM OM-ABRS): vide track 
nil Mi Areher Ku\ Ml Mee ST.. 5EQ (27 04"S, 
152UEE). NVF/Arawwia il, OMMO15079, J. 
SLanisie. D. Poller), Munanna NT.. Mi Nebo, SFQ 
(27 2.VS. l5T47-F)\veiselerophylMFQMMO(v25S, 
Aug l l i7f>. M.J. Bishop); Wralten"S Camp, Wrallens 
S.F., SEQ (2ol7'S. l^2'2l("E) NVP. litter ( \ t}M- 
M011563, I Tin! P/XO J.Siams.c.A.GreenKt'ruip, 

Tret North s r_, e 4ii km w oj Casino, nsw 

(:S"S4\S. [52 45*1 i iODpM M.)0+,AMCI54733, J4 
Mav 1976. I' II Golman. 1. Loch); Red Scrub I I 
NENSW' (28'38'S, I53'20'*E), raint'oresi (I 

AMGI2SSM7. :o Mar 19S1. AMQM-ABRS); Mi 
Emdesay. Uppej Richmond Rv. NSW (3. 
A\K IS4734. 77 Apr l l )4X. E. Pnee); Knreelah Ck. 
ISeaury ST,, c 15 km W ol Erbenvdle. NSW 
| 16 Jri ES2-2HT)(S:. AMC154'30. ES May 1976, 

P.H. Colrnnrr. I, r oeh), 

Shell small, depressed, diameter 2.09-2.81 
mm (mean 2.30 mm) wiih ? 3/4 10 4 1/4 (mean 
4+i whorK l'i^i whorl descending, Spire (1 i 
12Ue) moderalclv elevated to depressed, 
SP 13 WAV ratio 0.16-0,32 (mean 021). Height 
i [17 -1 .58 mm { mean 1. 24 mm), H/D ratio 0.50- 
O.oJ (mean 0.57). Frotoconch of 1 1/4 to 1 5S 
uhorls. meandiameter564.5(j.mat 1 1 2 whorls. 
Apical sculpture (Fig. I20f-h) of fine, protrac- 














FIG. 122. Distribution of Coenocharopa parvicostata 
sp. nov. 

lively sinuated radial ribs, more crowded near 
the nuclear-postnuclear boundary, crossed by 
irregular spiral cords. Postnuclear sculpture 
(Fig. 120d) of regularly spaced (often obsolete) 
blade-like, protractivcly sinuated radial ribs, 
continuous on the base. Microsculpture (Fig. 
1 20d) of numerous, thread-like radial riblets and 
low, broad crowded spiral cords. Umbilicus 
wide, V-shaped, diameter 0.45-0.84 mm (mean 
0.57 mm). D/U ratio 3.34-4.62 (mean 4.16). 
Sutures impressed; whorls rounded below and 
slightly shouldered above the periphery (Fig. 
120c). Aperture roundly to ovately lunate. Lip 
simple, columella dilated and twisted toward the 
umbilicus. Parietal callus present. Colour light- 
yellow horn with a pink tinge in some specimens. 
Genitalia with epiphallus (Fig. 121a) tubular, 

internally with longitudinaly thickenings, enter- 
ing the penis apically through a simple pore 
surrounded by a muscular collar (Fig. 121b). 
Penial retractor muscle inserting on the epiphal- 
lus prior to its entry into the penis chamber. 
Penis (Fig. 121b) expanded, tubular, internally 
with crowded pustulations and weak lon- 
gitudinal pilasters. Vagina short, expanded, in- 
ternally with spongy pustulations. 

No data available on radula. 

Based on one dissected specimen (QMMO- 

Range and Habitat 

From the Dorrigo area, northern NSW to Mt 
Mudlo. north of Gympie, SEQ. It has the largest 
distribution of any Coenocharopa species and is 
found in both warm moist notophyll vine forest 
and drier araucarian notophyll vine forest. Al- 
though there is some range overlap with C. alata, 
C. parvicosiata has only been found sympatri- 
cally with that species in the Beaury State Forest, 
northern NSW. No data available on micro- 


In spite of the comparatively large number of 
dead shells available for study only one sub- 
adult specimen was suitable for dissection and 
more material is needed to confirm structures of 
the terminal male genitalia. As may be expected 
in a wide ranging species there is a degree of 
interpopulational variation in shell characters. 
Southern specimens are larger and have bolder 
sculpture on the protoconch. 

Coenocharopa yessabahensis sp. nov. 
(Figs 123-125; Tables 22, 24) 


For the type locality, Yessabah. 


Its relatively large size and postnuclear sculp- 
ture of numerous low, crowded, radial ribs dis- 
tinguish C yessabahensis from congeners. C. 
sordidus is most similar but has widely spaced 
radial ribs with conspicuous periostracal blades. 
Anatomically the penial pilaster pattern of apical 
stimulatory pad and large basal U-shaped thick- 
ening contrasts with the typical pattern of simple 
longitudinal thickenings. C. m u It irad lata which 
is possibly sympatric with C. yessabahensis, has 
regular widely spaced radial ribs, more con- 
spicuous microsculpture and is much smaller. 









9 rvfl 

« J s •■III ilr « iJ^ 



? jlP^ 






FIG. 1 24. Coenocharopa yessabahensis sp. nov. Yessabah Caves, via Kempsey, NSW. QMMO17013, 
paratype. a, genitalia; b, details of penis interior; c. ovotestis: d. hermaphroditic duel and talon; e, pallial 
cavity. Scale lines as marked. 

FIG. 123. Coenocharopa yessabahensis sp. nov. , a-c, Yessabah Caves, via Kempsey, NSW. QMM017261, 
holotype: d-h, same data as holotype. QMMO17013, AMC121680, paratypes. a-c, entire shell; d. marginal 
leeth; e. spire; f. apical sculpture; g, post nuclear sculpture; h, central and lateral teeth. Scale lines as marked. 



FIG. 125. Distribution of Coenocharopa yessabahen- 

sis sp. nov. 

Type Material 

HoLOTYPE: QMM017261, Yessabah Caves, via 
Kempsey, NENSW (31°05 , 30"S, i52 B 5l'E), vine 
thicket on limestone rocks. Collected by J. Stanisic. 
D. Potter, P.H. Colman. 3 March 1987. Height of shell 
2.22 mm, diameter 3.75 mm, H/D ratio 0.59, D/U ratio 
4.24, whorls 4 3/4. 

Paratypes: 130 live, and 20 dead adults and sub- 
adults, OMMO17013, same data as holotype; Yes- 
sabah limestone cave, Sherwood Quarry. W of 
Kempsey, NSW (31°06'S, 152°42*E), ground litter in 
and out of cave (63, AMC121680. 9 Oct 1979, P.H. 
Colman, J. Stanisic). 


Shell small, depressed, diameter 3.19-3.83 
mm (mean 3.40 mm), with 4 1/8 to 4 3/4 (mean 
4 3/8) normally coiled whorls. Last whorl des- 
cending slightly in front. Apex and spire (Fig. 
123c) onlv slightly elevated, SP/BWW ratio 
0.1 1-0.26 '(mean 0.20). Height 1.66-2.22 mm 
(mean 1.88 mm). H/D ratio 0,48-0.61 (mean 
0.55). Protoconch shiny with 1 1/4 to 1 5/8 
whorls, mean diameter 644.5 fxmat 1 1/2 whorls. 
Apical sculpture (Fig. 123f) of weak, squiggly, 
irregularly spaced spiral cords and vague, low- 
radial ribs. Postnuclear sculpture (Fig. 123g)of 
low. broad, crowded spiral cords, crossed by 
weak thread-like radial and microradial ribs, 
continuous on the base. Umbilicus (Fig. 123b) 
narrow, U-shaped, diameter 0.76-1.03 mm 
(mean 0.83 mm). D/U ratio 3,73-4.43 (mean 

4.12). Sutures strongly impressed; whorls 
shouldered above and rounded below the 
periphery (Fig. 123c). Aperture roundly lunate. 
Lip simple, slightly thickened at the columellar 
margin. Columella dilated and slightly reflected 
over the umbilical opening. Parietal callus 
developed. Colour yellowy-brown. Based on 
20 measured adults. 

Body colour dark grey, with ommatophores 
and pedal glands darker. Terminal male genitalia 
with epiphallus (Fig. 1 24a) reflexed, entering the 
penis apically through two spongy thickenings. 
Penis (Fig. 124b) thick, muscular with a thin 
sheath for its entire length; internally with a 
spongy horseshoe-shaped pilaster (Fig. 124b) in 
lower part of the penial chamber, and a small 
subcircular thickening in the apical half, other- 
wise with pustules. Penial retractor muscle in- 
serted on the epiphallus prior to its junction with 
the penis. Free oviduct short, muscular, internal- 
ly with rather vague, transverse, spongy thicken- 
ings. Vagina short, thin with internal 
longitudinal pilasters. 

Radula (Fig. 123d, h) with mesocone of the 
central and inner laterals greatly enlarged. 

Based on four dissected adults (QMMO- 

Range and Habitat 

Only on the limestone outcrop at Yessabah, 
near Kempsey, NSW, which supports a low vine 
thicket. Crawling on limestone rocks. 


C. yessabahensis. Letomola contortus and 
Rhophodon kempseyensis are restricted to the 
limestone outcrops between Yessabah and Car- 
rai, near Kempsey, NSW suggesting that these 
outcrops are important refugia for terrestrial 

Coenocharopa multiradiata sp. nov. 
(Figs 126-127; Tables 22, 24) 


Latin multi -, many; and radiata radial; refer- 
ring to the numerous radial ribs. 


Sculpturally, C. multiradiata is the most dis- 
tinctive species of Coenocharopa described 
herein. The sculpture is regular and in many 
ways simulates the more typical reticulate pat- 
tern seen in Gyrocochlea and Rhophodon. Com- 
pared with its congeners. C multiradiata has 






FIG. 126. Coenockaropa mulliradiala sp. nov. a-c. Top of Condamine River vallev on W slopes of Wilson's 
Peak along Croftby Rd. NSW. QMMO 18973, hololype; d-f, Natural Arch, Carrai Caves, NSW. AMC 
142958, paratype. a-d, entire shell; e, apical sculpture; 1", post nuclear sculpture. Scale lines as marked. 




20° S 





alt. > 500m 



1 ^2 C K 


154 a E 

FIG. 1 27. Distribution a{ CoL'twchanf pa m aid rudiata 
Sp, nov. 

more numerous, less protractively sinuated 
radial ribs and microradiats which have con- 
spicuous periostracal blades. In addition, the 
microspirals are less crowded, more slender and 
raised to produce a conspicuous reticulate pat- 
tern. C. ahtta is larger, has fewer radial ribs with 
extremely expanded periostracal processes and 
has a more reduced microsculpturc. C. parvieos- 
lata which is most apt to be confused with C. 
mtdtiradiaia. is larger with more elevated spire 
and has fewer, less regularly spaced, more 
productively sinuated radial ribs. In suhadult 
specimens, these two species can be differen- 
tiated by their apical sculpture which consists of 
few. widely spaced, well defined, radial ribs in 
('. multiradiaiu. and crowded, low ridges, be- 
coming more crowded at the nuclear- 
postnuclear boundary, in G. panicostata. C, 
yessahahensis is larger with virv crowded, 
weak, radial ribs and reduced microscuipture. 

Tvrr Material 

HolOTYPE: QMMOISW, top ol Contlamine River 

valley on W slopes of Wilson's Peak, along Crofthy 
Road,SEQ(28' ]t.\s. 152 ":iS'E). rainforest/hoop pine. 
Collected AM.QM-ABRS. 16 Mar NSL Height of 
shell ].l 7 mm.diamcter2.l6mm, H/DraliuO .54, D/U 

ratio 3 03. whorls 4 I 8- 

PaRATYWES: AMC128634. same collection data as 
K'l.i!'.. ,-;■. N I Ural Vvh,Cirr 4 ii Caves. iioarKcrnpsey. 
NSW. leal Ulter {V. AMC142V5B, 22 Feb 1980, P.H. 
Co) mail): c 6.5 km E ol 1 fckgate. Mi Pfkapene S.k. 
NSW (28 tUS 152 45 E>. riverine rainloresi (11. 
QMMGIQ893. i-l Mai |«l. am om-ABRS). 

Otiifr M-xiTftur 

I nii._'M<>rK' l\ocs 31 Nalural Arch. W of Kempsev. 
NSW (30 vrs. 15: :I'F:). under sifts ullimestone 
(7.AMC143728 10 Mar 1981 W IY-kI.t. O. Gnl- 
lilh:.): Korcflah S.K NSW. c 15,2 km from Acacia 
Plaretiy kill,irne\ R.I (2H°2| "S. I52 : 24T) 
SEYT MVF( I. AMC" 128553. 15 Mar WU I . AM OM- 


Shell diameler IV 1-2. lb mm (mean 2.03 
mm), with 3 7/8- to 4 1/8+ (meai 4-) normally 
coiled whorls. Last whorl descending slightly. 
Apex and spire (Fiu. 126c). very .lightly 
elevated. SP/BWW ratio 0.08-0. 1 6 (mean 0. 13), 
height of shell 1.07-1.19 mm (mean 1.14 mm). 
H.D ratio 0.43-U.62( mean 0.54). Protoconchof 
1 1 2 lo 1 5/8 whorls, mean diameter 511.1 |xm 
at ! 12 whorls. Apical sculpture (Fig. 126e) 
consisting of widely spaced, conspicuous, slight- 
ly curved radial ribs with raised periostracal 
blades and numerous, narrow, low spiral cords. 
Post nuclear sculpture (Fig. 12hf) of numerous, 
regularly spaced, weakly protractively sinuated 
radial ribs which have prominent periostracal 
blades that are often worn, 38—4-7 (mean 42.5) 
ribs on the body whorl. Ribs/mm 5.7S-6.96 
(mean 6,63). Microscuipture (Fig. 126f)offine 
thread-like radial ribleis. n-1 3 between each pair 
of major ribs, and fine, slender conspicuous 
spiral cords which are not raised at their junction 
with the micToradials. Sculpture continuous on 
the base. Umbilicus (Fig. 126b) narrow. U- 
shaped, diameter 0.47-0.55 mm (mean 0.51 
mm). D/U ratio 3.79-4.36 (mean 3.98). Sutures 
impressed, whorls shouldered above and 
rounded below a slightly flattened periphery 
(Fig. 126c). Aperture roundly lunate. Lip 
simple, columella not expanded. Parietal callus 
developed. Colour light-horn. Based on 10 
measured adults. 

Anatomv unknown. 


Rwge and Habitat 

T rom the top of the Richmond Range, NSW 
and from the Canal limestone outcrop near 
kempsey, NSW; in dry cool subtropical vine 
forest and warm temperate rainforest. It is prob- 
able that C. multiradiQta extends further south in 
vv arm temperate forest. No data arc available on 


C. multiradiata differs from average 
dtenocharapa patterns in a number of features. 
Nonetheless these altered character states are 
n adily derived from typical Cnenocharopa. In 
the absence of anatomical detail, placement in 
this genus is preferable to creating another 
nomenclatural entity based on shell characters. 

Egilomen lredaie. 1937 

r^ilomc/i lredaie. 1937a, p. 328: lredaie. 1941a. pp. 
267, 269; Kershau, 1455, p. 2^; Bunch, 1976. p. 

lYPF Spn ]RS 

Helix eoehlidutm ( o\ 1 868; bj original desig- 

n-'i vioi 5 ■ 

Egilomen was introduced for five species with 
bold radial ribs and wide umbilicus. Egilomen 
is herein redefined to include species that have 
apical sculpture of prominent radial ribs with 
spiral elements absent or reduced, postnucleai 
sculpture of bold radial ribs, and a microsculp- 
luic of low rounded spiral cords. 'Helix' 
i oefilidium, and a new species from northern 
NSW and SEQ arc assigned. Other species pre- 
viously allocated to Egilomen probably belong 

DlAGM i 

Shell diameter 1.62-3.07 mm. with 4 14 to 5 
1/8 tightly coiled whorls, last descending rapid- 
l\ . Apex and spire weakly to strongly (gtohostj) 
elevated. Apical sculpture of broad, bold, W idc- 
ly spaced, weakly curved radial ribs, and weak 
spiral cords. Postnuclear sculpture of broad, 
bold, moderately lo widely spaced {cochlidiwnh 
weakly protractively sinuated to vertical radial 
ribs. Microsculpture of low. rounded, crowded 
spiral cords crossed by fine, thread-like radial 
riblets. Umbilicus wide, U-shaped {eoehluliitm) 
in closed (glohosa). Sutures strongly impressed. 
Whorls shouldered above and rounded below a 

weakly compressed periphery. Up thickened, 
rarely with a distinct basal knob (some popula- 
tions of ghhosa). retracted at the suture to form 
an apertural sinus. 

Animal with foot short and broad; colour white 
without any accessory pigmentation. Pa 
cavity with complete secondary ureter and 
strongly bilobed kidney- Talon with very shoo 
stalk, subcircular. Terminal male genitalia with 
well-differentiated epiphallus and gtfflpli 
epiphallic pore. Penial retractor muscle insert- 
ing on epiphallus. Penis subcylindrical. inter- 
nally with a single, large longitudinal pilaster 
and an accessor) circular pad. Female genitalia 
with relatively long vagina and free oviduct. 
Radula with eeniml toolh much smaller than 
laterals, otherwise without unusual features. 

1 , i\,r \k,m i\s 

Conchologieally t gtlnmen rescmblesCra/ofw; 
and Rhophocfon Rhophodon differs rnOSI 
markedly in having apertural barriers. Micros- 
culpturally Egilomen (Figs 12Kg. 1311") has 
prominent, rounded, continuous spiral cords 
Crossed hv fine, thread-like radials in contrast to 
Rhophodon. which has low, barely visible spiral 
cords and prominent, high microradial ribs with 
elongate beads at their intersection (Figs 8 If. 
B7e). Anatomically the strongly bilobed kidney 
pf Egilomen (TigS i 2dd, 132f) contrasts with Ihc 
weakly bilobed. to almost unilohed, condition of 
Rhoplwdon (Figs 7 -f- 75h), Terminal male 
genitalia of Egilomen have the penial tetfaetpl 
muscle inserted on the epiphallus (Figs 129b. 
132b) whereas In Rhophadon it is always in- 
serted at the penis-epiphallus junction (Figs 7 la 

Cralopu differs in having more prominent api- 
cal spiral cords, more raised microsculpture and 
less differentiated epiphallus. 

Egilomen coehlidium (Cox. Isf S) 
(Figs lis ISO; Tables 22,25) 

tfeltxeacMMiimCm is;,s n l&pl, B»*fe. 1. 
Egtfomwi coehlidium (i n\i. Iredaje, iyJ7a, p 
lredaie. 1941a. p. ZG9. 

E. eoehlidium differs from t. globosa in its 
ilattened shell and wide umbilicus. It is most 
similar to species of Rhoph<)don but lacks a per 
tural barriers and has less crowded apical sculp- 
ture. Discocharopa aperia (I ig M) has a wide 
umbilicus and prominent radial sculpture hut is 







d* . \ 







i^fc,*-^ , 


*-■ - - ■ 


$ ■?'■[ 


f K- 


- ^ . 



FIG- \29. Egilomen coehlidium (Cox. 1868). a. c,d, Upper Pine Ck, Canungra, SEQ. QMMO16905. b, Mills 
Rd. Moonpar S.F., NSW, QMMO17290. a, genitalia: b, details of penis interior: c. talon and carrefour; d. 
pallial cavity. Scale lines as marked. 

FIG. 128. Egilomcn coehlidium (Cox. 1868). a-c. Clarence River. NSW <\MC635Q5. leclotvpe: d. h. Mills 
Rd, Moonpar S.F.. NSW. QMMO17290; e-g, The Island, off Grafton, NSW. AMC57247. a-c, entire shell; 

d. marginal teeth; e, entire shell; f. apical sculpture; g. post nuclear sculpture; h. central and lateral teeth. Scale 
lines as marked. 


much smaller, lacks regular spiral sculpture and 
1 || has finer, more crowded radial rihs. 

s a-s pi t 5 ! i 

Type Material 

Lfctotype: AMC63505, Clarence River, NSW. 
J = _ |§ Height of shell 1.25 mm, diameter!. 26 mm, H/D ratio 

p 5f j |j * | j . 0.55. D/U ratio 3.14, whorls 4 1/2. 

| = ^= == = = Parai.ECTOTYPES: AMC1537I4, 2 specimens, same 

collection data as holotype. 

Other Material 

Dorrigo. NSW ( 1, AMC63790);The Island, offGraf- 

ton,NSW (12, AMC57247); O'Reillys, Lamington 

N.P., SEQ (28°14'S, 153 D 15'E), MFF (1, 

B |s AMC 129243, 17 Mar 19S1, AM/QM - ABRS); W 

= - =~, ^p ? K E r:! s £ side Richmond Range, NSW, tributary of Duck Ck 

1 (2& a 33'S, 152 r 40"EH2. AMCl28524,"l4.Mar 1981, 

? 5 _ * rt ^_ AM/QM - ABRS); top of Condamine River Vallev. 

I tl |5 W slope of Wilson's Peak, NSW (28M6'S, 152°28'E) 

|S i Eg a ! is I| S (1, AMCJ2S637, 16 Mar 1981, AM/QM - ABRS); 


I !- '_ — 


Upper Pine Creek, Canungra, SEQ, NVF/Araucaria, 

1 5 underrock(l.QMMO16905,21Jan 1987, J. Stanisic 

c5 =-i 

S i - 5^ . **t 

» irt 3" — * * »*- 

D, Poller); Mills Rd. Moonpar S.F., NSW PO'IS'S, 

j | 152°39'E) rainforest (1, QMMO17290, 7 Mar 1987, 
J. Slanisic. D. Polter); Carr's Creek, Clarence River, 
NSW (L AMCS737, presented C. Hedley, ex Brazier, 

>-3 £ 

c = 

§jg =5 §? Rossiter. Co.x: 1, AMC152253, e\ Brazier); Cherry 

II ^ll = -s U ft a 5 II ft r Tree North S.F,, NSW (2N' J 54*S, 152 J 45'E) litter (5, 


c ^ ' " ~ '* AMCI53714. AMC154732. 19 May 1976, P.H. Col- 
man. 1, Loch); Goomburra S.F., SEQ (27 a 59'S, 
it _ B 152 D 2l'E)(l, AMC12S618, 16 Mar 1981 AM/QM - 

K •/■ .C 3C *^?i 5 "• .p .2 _ -3-4 

_ r /-u" in i ^ tn F-r-rw. r-r- 

= O 

ABRS); N of Dorrigo, Clouds Ck S.F., NSW 
(30 D 07'S. 152-41'E) dense rainforest (1, AMC- 
128319. 11 Mar 1981, AM/QM- ABRS): c. 5.5 km N 
of Dorrigo NSW <3fT23"S. 152'44'E) (t, QMMO- 
10794. Mar 1981, AM/QM - ABRS); Cunninghams 
Gap, SEQ. leaf litter (1.QMM01729 1. 18Jun"l979, 
G. Annabell); Koreelah Ck, Beauty S.F.. NSW (28*- 
S 2TS, 152 a 20*E)(3. AMC15-1735, IBM-ay 1976, P.H. 

3 I* 1= 2f Colman, F Loch). 

£ 3| 3- $7 $± 

|« % 2E 5§ c = te gq SE IB 

- - -a ~ "" ~~ " Diagnosis 

Shell diameter 1.91-3.07 mm (mean 2.35 mm) 
| _ with 4- to 5 1/8- (mean 4 1/2+) lightly coiled 

dl *l « whorls, the last descending rapidly in front. 
*~ f ; ; ^ iS ? Apex and spire (Fie. 128c) flat to slightly 

elevated. SP/BWW ratio 0.09-0.27 (mean 0. 1 8), 
height 0.95-1. S3 mm (mean 1.30 mm), H/D 

"" * x " ' ratio 0.49-0.61 (mean 0.55). Protoconch of 1 

12 to 1 5/8 whorls, mean diameter 49 1.8 (im at 
I 5 1 1/2 whorls. Apical sculpture (Fig. 12Sf) of 

ilot 2 + I- *- 1 l~ 2 5 widely spaced, slightly curved, broad, prominent 
1=1 ?§§ § J§ ^i i 5*o -o |I radial ribs, and low. inconspicuous spiral cords. 
jlpll W || || |||i| l| *1 Postnuclear sculpture (Fig. 12Sg) of broad, 
sp 5 6 5? widely spaced, protraciively sinuated radial ribs. 


I - 8 


- _ 

-■ - 

2 ^ 




20° S 


:,-. ri s 


151 D E 




FlfT 130. PiMributiort al' Egikmnvi rochlidittm (Cox. 

30-47 (moan 36. S) ribs un the body whorl. 
Ribs/mm 4, 22-5, 99 (mean 5.07), Mieroseulp- 
turc (Fig. I28g) of fine radial rihlcts, 8-12 be- 
tween each pair of major ribs, and fine crowded 
spirals. Sculpture continuous on the base. Um- 
bilicus (Fie 1 28b) ^ ide to vcrv wide, U-shaped, 
diameter 0.60-1.03 mm(mean 0.S2 mm). D/U 
ralio 2.59-3.47 (mean 2.93). Sutures impressed. 
Whorls flattened below r and rounded above a 
laterally compressed periphery (Fig. 128c). Aper- 
ture ovatelv lunate. Lip simple, thick, margins 
convergent. Columella only slightly dilated. 
Parietal callus well developed. Colour white to 
light straw-yellow, often covered in dirt particles. 
Based on 10 measured adults. 

Vas deferens a thin tube. Epiphallus (Fig 
12%). cylindrical, greatly expanded, reflexmg 
prior to entering penis apically through a simple 
pore surrounded by two fleshy lips. Penis (Fig. 
129b) short, internally with a large, apical, lon- 
gitudinal pilaster, and a basal, circular pilaster. 
Penial retractor muscle (Fig. 129b) inserted on 
the epiphallus prior to its junction with the penis. 
F rce oviduct short, muscular, internally with lon- 
gitudinal pilasters. 

Radula(Fig. I28d,h) with central tooth greatly 

Based on two dissected specimens (QMMO 

R \N(ih, \ndHahit\t 

Mainly confined to cooler subtropical 
notophyll vine forests on highlands of the Great 
Dividing Range (between Dorrigo, NSW and the. 
Mistake Mountains, SEQ), and ihe MePherson 
Ranges on the NSW/QLD border. The early 
Oration Clarence River records need to be con- 
firmed. The only data available on microhabiiai 
are from a specimen collected from soil under a 
rock at Canungra. SEQ. 


The apical longitudinal penial pilaster and 
basal penial stimulator contrast with the apical 
stimulator and basal longitudinal pilaster of E 
glohosa. This difference in penial surface sculp- 
ture is probably a species recognition change 
brought about by sympatric interaction. E. 
corhlidium is poorly represented in collection* 

Egilomen globosa sp. now 

(Figs 131-133; Tables 22, 25) 

Latin globow, rounded; referring to shell 


E. qlabosa differs from E. cocklidlurn in 

having a more globose shell with strongly 
elevated spirc.morc crowded radial ribs, more 
prominent spiral cords and closed umbilicus. /:. 
glohosa is most apt to be confused with the 
sympatric Cralopa siroudensis (Fig. 95) which 
has similar shell shape, prominent radial ribbing 
and closed umbilicus. However, C. stroudensis 
has fewer whorls, a more conspicuous apertura! 
sinus, less elevated spire, thinner lip. and a 
greater number of ribs on the body whorl. El- 
sothcra genithecatu (Fig. 108) and£. nauiiloden 
(Fig. 106), from northern NSW, have closed 
umbilici and prominent radial ribs, but are much 
larger, with Hatter spires, and comparatively less 
crowded radial ribs on the body whorl. 

Type Material 

HoiOTYPE: QMMO 17292. sidetrack off Ml AreheT 
Rd t MlMeeS.F\.SEQ27 t 04 , S. 152'4FE. Collected 
31 Sepl 1982, AM QM - ABRS. Height of shell L.87 
mm. diameter 2.47 mm. H/D ratio 0.76, whorls 4 3/4, 
ParaTYPFS: QMM012178, AMC136S21. 20 

.■■ imei - ami i olte Hon gaw $s tiolotypej mi 



Guyra N.P.. SEO CZSWS, L5:T35'E) MVF/NVF 
Plmucarto (9, QMMOI1068, AMCI36463, 7 Sepi 

iws: ,\\j QM - ABRS): nturbase. Ml Warning N.P.. 
NSW (2S 2-TS. 153"HPE] NVF/Palms 12, GMMO- 
10483 'Mm I'JSl. AM/QM - ABRS); Mr Warning. 
N;P* NSW (28*24 S. I53 B irfe)j£, AMC129305, 19 
M,it losi, AM QM - ABRS); Natural Bridge N.P . 
SEU 12SM3S. 153'14'E) NVF (8, QMMO10459, 
\M( 124270, 18 Mar 19BI, AM/QM - ABRS): 
pfa1 'Lck. Nightcap Range N. P.. NSW (2S U 34*S, 
153'19'E). sublropieal rainlorest, under bark on log 
(15. QMMO 17012, II Mar 1987, J. SranisicD.l'Ot* 
ler; 13. QMM01TM& |] Apr I9S7 I Sianisie. J 

Otiu:r Materia* 

Tnormmbar PoresJ Road, Toonumhar S.F.. NSW 
(2S 33 S.I 52 45 F- )CNVF( 1 , QMMO 1091 1,14 Mar 
1981. AM/QM - ABRS) I n forest Park, near 
Maleiiv ,.n Maleny-MorUville Rd. SEQ. NYT (I. 
QMM01 iSfclS Dec 1 98% J Slvmsic); sidetrack off 
M. ArrhcrRd.Mi Mee S.F. SEQ (27 t 04'S. !52 C 4PE] 
(2. QMMOI5075. QMM015G76, 6 Sep! 1981 J 
i. i .. D.Potler;2.0MMQl2i77,AMC136520,3(l 
Sepi I9M2, AM/QM - ABRS). Fred's Rd Ml Mee, 
SIQ i:ti;;Vs. |52°43 Ei NVf , liner (I. (JMMO- 
0( [Q !. J. Stanisie, D. Pniicrj. c. I.hkm 
N af Numinbah Vallev lurnnff on Burleigh- 
Springhrook Rd, SEO NVFPaims. lmct ( 1 QMMO- 

17293. 17 Dec 1981/, J. Standi i E(enilwort*i S.F.. 
SEO .:o "37'S. 15242*E) raioRvreftt. lilter (1. QM- 
MOi7^4.22Ma\ L980.J.Siamsit , \ "■■■ c i B3 in 
gum. NSW (E AMC152199 UwerUMuwilturnbah, 
NSW (\. AMC2S495, ev Tech Mus v. Northern 

Rivers. NSW (E AMEMS23. iMlcrd, c\ Cox 1; top ol 
idamineRrver Valley, NSW (28 I5\S, Lfi2 v 29*E) 
I I AMC128Q02. 16 Mar 1981. AM/QM - \BHS 

shell iliameter 1.62-2.47 mm (mean 2-14 mm) 
of 4 ^ B in 5 (mean 45/8+) tightly coiled whorls, 
1 I whorl descending more rapidly. Ape\ and 
spire (Fig 1 31c) moderate I v to stronelv 

li vatcd,SP/BWWnilio0.25 D S5(mean<W3i 
h ighiof shell 1.19-2.04 mm- (itican 1.64mm). 
WD ratio 0.71-0. 96 (mean (k77). ProtoeonchoJ 

I J/8 |o I j S wJioHS, mean diameter 4°-7.5 |xm 
at I 1 2 whorls. Apical sculpture (Fig. !3ie)ol 
slight) y curved, Lrt enl\ spaced, broad radial ribs. 

Postnuelcar sculpture (Fig. 13 If) of strong, 
slightly protectively sinuated radial ribs, 53-80 
(mean 64.1) ribs on the body whorl. Ribs/mm 
7.75-12.1° (mean 9.04). Rib interstices about 
4-5 times their width. Microsculpture (Fig. 
13 If) of thread-like radial nblets crossed by low, 
crowded spiral cords. Umbilicus closed or 
reduced to a lateral crack. Sutures impressed: 
whorls slightly shouldered above and rounded 
below a laterally compressed periphery. Aper- 
ture roundly lunate. Lip expanded at the basal 
and coiumcllar margins, occasionally with a 
basal denticle Columella strongly dilated with 
a ridge-like thickening, reflected over the um- 
bilicus. Lip sinuous, retracted at the suture to 
form an apertural sinus. Parietal callus strongly 
developed, Colour white to light straw -yellow 
Based op 3 4 measured adults 

Genitalia with a strongly expanded, muscular 
epiphallus (Fig 132b). Epiphallus with ascend- 
ing arm parallel to the penis and reflcxed before 
entering the penis apically. Pcnial retractor 
muscle (Fig. 132b) inserted on the epiphallus 
prior to its junction with the penis Penis (Fig. 
132b) Internally with a apical longitudinal spon- 
gy pilaster and a basal pad-like thickening. 
Radula (Fig, 131d.h) similar toE. cochtidlum 
Based on 6 dissected specimens (QMMO 

Range v\n habitat 

Warm, moist notophyll vine forests of the Big 
Scrub area. NSW and the D'Aguilar Range. 
SRQ, It is sympalric with E. cochlidium in the 
McPherson Ranges. An outlier population at Mt 
E-uvra, north of Gympie. SEQ. is the northern 
ii nil oi the species & glohosa lives-imder lags. 

Rl MARJ ■ 

Specimensot / . flobasa (QMM01206K, AM- 
CI 36463) from Mt Gtiyra are smaller than 

average an;l have an ahereel lip With a small 

Omphaloropa gen rw 

Rl v-Mi . ,. . . 

From die I ".ark 0tnph&hs t umbilicus; refer- 
ring to the prominent umWKcUS. 

FIG I II I giltmcH efo&ft* sp nos a c Skfeiraci oil Mi Arc+iei Rd Mi Mx S.T - SEQ. QMMQK7292 
Teraniu Ck. NSW. QMM< ) I 71 XX. puralype. e-f. V: \|. . :S I '.. Si :Q. QrvfM0l2l7S,paralype: 
■■ !■ i oivia n.p,. si QMMOJi2G68.parfuypc a-i cniircshclhdifnaigirtalttcihie, apical sculpture, r. pi>st 

nucli II '-'--iilpiari. : p. eiiinv shell shn\\ uie eictiliele mh lip. Ii. eeali;il r:,! ', IfcEth. Scale lines as marked. 





:■,;:; :■::■;■:■:.. 








FIG. 132. Egilomen globosa sp. nov, Terania Ck, Nightcap Range. N. P., NSW. QMMO17012, paratype. a, 
genitalia; b, penis interior; c ? talon and carrel'our; d, hermaphroditic duct: e. ovotestis; f, pallial cavity. Scale 
lines as marked. 





23 C S 


151 a E 

152 *E 



FIG 1 33. Distribution pi Egilotm'ti gfoboSQ Sp, HCfl 

TVFKSf'H if.s 
Omphaloropa varieosa sp. nov. 


Shell small, mean diameter 2.23 mm. with 
lightlycoiled\vhorls(mean4 12-), last descend- 
ing rapidly. Apex and spire moderately 
elevated. Protoconch with low spiral cords. 
Post nuclear sculpture of Very widely spaced, 
prominent, protectively sinuated radial ribs and 
microsculpture of low radial riblcts and spiral 
cords which have raised knobs at their intersec- 
tion. Umbilicus wide, cup-shaped. Sutures 
strongly impressed. Aperture roundly lunate, lip 
simple. Columella only slightly dilated. 

Terminal male genitalia with epiphallus poor- 
ly differentiated from vas deferens, entering 
penis apieally through a papillate verge. Penis 
with a sheath and internal longitudinal pilasters 
Female genitalia without unusual features, 
Radula with very small tricuspid central tooth, 
otherwise typical. 


Ompluiloropa is conchologicall y distin- 
guished by small size, apical spiral cords, few 
widely spaced radials on the adult whorls and 
wide cup shaped umbilicus. Rhophodon is also 

small, with wide umbilicus and many to few 
radial ribs on the adult whorls, but in contrast has 
curved, spaced radials on the protoconch, and 
few to many lamellate barriers in the apem 
Sinployca inlensa hasaspirally lirate protoconch 
but more crowded posmuclear radial ribs. 

Omphuluropa varieosa sp. nov. 

1 1 igS 134-136: Tables 26. 27) 


Lalin varix, dilated vein; referring to the 

prominent postnudcar radial ribs. 


Within its range. O. varieosa is most apt to In- 
confused with Sitiployea interna and species Oi 
Rhophodon. S. intensa has a brown shell with 
spiral cords on the protoconch but has very 
crowded radial ribs on the body whorl and mm h 
smaller umbilicus (Fig. 137), Sympatnc 
Rhophodon alSQ have monochrome or tlammu- 
latcd shells with wide umbilicus and few to many 
radial ribs on the adult whorls. However, in 
contrast io£X varieosa, they have a radially lirate 
protoconchs with weak spiral elements, and few 
to numerous aperturai lamellae. 

Typf Materia] 

HtoU 1 m : QMMOr>:], G^vigj freek. Boulder- 
combe. SHO [23 34'S. 15'r:,S E) MVF. in iiller. 
collected b\ J, Stanfcic, D, Putter. 9 Jul 1983. Height 
of shell 1.42 mm. diameter 2.36; H D ratio 0.60, D/U 
mil,. 2.21, whorls 4 5/H* 

MG 16805. 88 specimens, same data as holotypt: 
sidetrack off Ml Archer Rd. Mi Mee S.F* 5 {2TQ4 
l52*4rE) NYTArauearia (3. QMMO1S077, h Sepi 
[963, J SlartisicD Potter); Dawes Range (24' J 28'S, 
L51TITEJ MVF'Araucaria (600+, QMM012671, 
AMC136800; ANtC136780 QM MO 12651. 4 Sepi 
19S2. AM QM-ABRSi; Bouldercombe Falls, SEVT 
(1 QMMG1727", 24 Mar JMX3, G. Annabell); Dan 
Dan Scrub, SEVT. in litter (5, QMMO16S03. 10 May 
19S4, J. Stanisic, D. Potter); Freds Rd, Mi Mee 
(27 flS 9 S, IS2*43 a E] mblfmfcl, leaf lit<er(7_QMMO- 
K363, 14 Apr I MSI I. J, Sianis.c. N. Hall, A Green); 
ECeililWOrth S.F, (26 40S, \>2'3^h)Araucaria. leaf 
litter (15. OMM(J17:sii.:; May 19SUJ. Slanisic. A. 

Other Mates**] 

SOUTHEAST QUEENSLAND - Entrance to slate 
lores! off Yabba Creek Rd. Yabba Creek S.F. 
(2h J 2.x'S, 152*3S*E)(1, QMMO13370, 7 Sept 1?62, 




6 L* 



x - 

m - 






FIG. 135. Omphahropa varicosa sp. nov. Gavial Ck, Bouldercumbe, SEQ. QMMOI6805. paralypc. a, 
genitalia; b. penis interior; c, ovotestis; d, talon; e. hermaphroditic duct; f. pallial cavity. Scale lines as marked. 

FIG. 1?4. Omphahropa varicosa sp. nov. a-c, Gavial Creek, Bouldercombe, SEQ. QMMOI 7281. holotype; 
d-h. same data as holotype, QMMOI 6805, paratypes. a-c. entire shell; d. radula; e, apical sculpture: f, post 
nuclear sculpture; g-h, central and lateral teeth. Scale lines as marked. 














152 C E 



FTG. 136. Distribution of Omphaloropa varicosa sp, 


AM/QM-ABRS); Fred's Rd, Mt Mee (27'05'S, 

l52 = 43"E)NVF,litter(l,QMMOl5981,28 0ctl9S4, 
J. Stanisic, G. Annabell); Cunningham's Gap, near 
monument (28°04'S, 152*24'E) 755m, SNVF/ferns 
(8, AMC136808, QMM012679, 7 Dec 1981, 
AM/QM-ABRS); Little Yabba Ck, Imbil S.F. 
(26°28"S, 152 D 38'E) NVF (6, QMMO12020, 
AMC136437, 8 Sept 1982, AM/QM-ABRS); 
Montville Rd, near Kondalilla Falls N.P. (26 D 40"S, 
I52°52"E) vine thicket on side of road (I. QMMO- 
1 1979, 8 Sept 1982, AM/QM-ABRS). 

Shell diameter 2.03-2.34 mm (mean 2.23 mm) 

with 3 3/4 to 4 3/4 (mean 4 1/2-) tightly coiled 
whorls last descending rapidly. Apex and spire 
(Fig. 134c) moderately elevated, SP/BWW ratio 
0.18-0.44 (mean 0.29), height 1.03-1.42 mm 
(mean 1.26 mm). H/D ratio 0.50-0.64 (mean 
0.7). Last whorl descending more rapidly. 
Protoconch white, glossy, exsert, 1 1/2 to 1 3/4 
whorls, mean diameter 505.1 (xm at 1 1/2 whorls. 
Apical sculpture (Fig. 134e) of 12-15 low spiral 
cords. Postnuclear sculpture (Fig. 134a) of 
widely spaced, prominent, protractively 
sinuated radial ribs, 18-35 (mean 26.8) ribs on 
body whorl. Ribs/mm 2.79-4.93 (mean 3.83). 
Microsculpture (Fig. 134f) of fine thread-like 
radial riblets, 1 8-35 between each pair of major 
ribs, crossed by numerous low spiral cords with 
distinct raised knobs produced at their intersec- 
tion. Umbilicus (Fig. 134b) wide, cup-shaped, 
diameter 0.70-1.07 mm (mean 0.91 mm), D/U 
ratio 2.16-2.92 (mean 2.48). Sutures strongly 
impressed. Whorls rounded below and 
shouldered above a rounded periphery (Fig. 
134c). Aperture roundly lunate. Lip simple, 
thickened slightly. Columella only slightly di- 
lated. Parietal callus strongly developed. 
Colour brown with alternate light and dark 
brown radial streaks. Based on 29 measured 

Genitalia with a poorly differentiated epiphal- 
lus which enters the penis apically through a 
large papilla (Fig. 135a, b). Penial retractor 
muscle inserting on the penis adjacent to the 
penis-epiphallus junction. Epiphallus internally 
with longitudinal pilasters. Penis (Fig. 135b) 
pear-shaped with a distinct bulb and sheath, in- 
ternally with apical papilla and low, spongy lon- 
gitudinal pilasters. 

Based on 3 dissected specimens (QMMO- 
15077, QMMO16805). 

Range and Habitat 

Humid subtropical notophyll vine forests in 
the southern part of its range; drier microphyll 
vine forests (with Araucaria emergents) and 
semi-evergreen vine thickets in the north. O. 
varicosa has been found living among litter and 
in soil, but not under logs. 


Conchologically O. varicosa could be linked 
with Sinployea but the simple internal penial 
structures relate O. varicosa to other charopids 
reviewed in this study. 

O. varicosa may be abundant in certain 
localities e.g. 600+ specimens were collected in 



X — V 'I ■» 

j£ FK ~? ?i 3* 

the Dawes Range, SEQ (QMM012671, QM- 
MO 12651, AMC13680Q, AMC 136780) in Sep- 
tember, 1982. 

• r- 3* — f- * 

x — .— . *•' "l "", ~ *"- 

= il ,,5 -.1 -i -J *5 

Sinployea Solem, 1983 
Sinployea Solem, 1983, p.Sl. 

;r 9 *■ - r " 

* g S 5 S 

5 £ ^ = * Si = ■£ 

* I* "■= "'^ E = 

Type Species 
Sinployea peasei Solem, 1 983; by original 


■-+ u- I- ?- - « x "* 

I 15 

I i 

3: 2 

1 1 1 e i 

'." £ * s 

p f- p ; i : 

— — 3* 

5 ft* *£ &fl 



F- H! 

Fl — s £ 

2 7 

* 5 * ' 

I li 

.J: ■: 

Previous studies 

Solem (1983) introduced Sinployea for 49 
Pacific Island charopids that show a uniform 
pattern of anatomical variation. The penis inte- 
rior is characterised by an apical verge, muscular 
collar and stimulatory' pads in the lower part of 
the chamber. Conchologically Sinployea is con- 
servative with fine apical spiral cords, rounded 
whorls with radial ribs, and microsculpture of 
radial riblets and fine, low spiral cords in all 

Sinployea includes 'Cralopa' intensa Tredale, 
1941. whose characters are close to the average 
Pacific Island species. 

Distribution and Ecology 

In the Marianas in Micronesia; parts of 
Melanesia including Kiribati (= Solomon Is) and 
Vanuatu (= New Hebrides): and into Polynesia 
as far east as the Society Islands. Locally S. 
intensa is confined to subtropical notophyll vine 
forests of northern NSW and SEQ. Its northern 
limit is the Rundle Range near Rockhampton. 5. 
intensa often occurs in strand-line rainforests 
within this region e.g. at Byron Bay, NSW; 
Cooloola. SEQ; Eurimbula. near Bundaberg, 
SEQ; and Burleigh Heads. SEQ. 


Sinployea is not easily confused with other 
Charopidae. The small, reddish-brown shell, 
slightly elevated spire, apical spiral cords and 
postnuclear crowded radial ribs are distinctive. 
Setomedea is larger, has apical spiral cords, and 
more complex adult sculpture with pcriostracal 
setae on the major ribs. Cralopa has prominent 
radial ribs, narrow to closed umbilicus, apcrtural 
sinus, and apical sculpture of low, broad, radial 
ribs and weak, low spiral cords. Omphaloropa 
(Fig. 134). has apical spiral cords but few major 
radials on the body whorl, a wide cup-shaped 
umbilicus, and low microsculpture. 


df-iC r-i 


Sinplovea intensa (Iredale, 1941) 
(Figs 137-139; Tables 26, 28) 

Cralopa intensa Iredale, 1941a, p. 269, fig. 5. 

Ills; §£ Is Comparisons 

1MI ii ali S. intensa is distinguished from sympatric 

charopids by its small brown shell which has few 

whorls, small umbilicus, apical spiral cords, and 

cj |= |§ postnuclear sculpture of crowded, protractively 

: :Sj 22 -.3 = sinuated radial ribs. Setomedea nudicostata 

(Fig. 35) has a flammulated shell and squiggly 
apical spiral cords. Omphaloropa varicosa (Fig. 
134) has a brown to flammulated shell with few 
prominent radial ribs. Discocharopa aperta 
(Fig. 93) has crowded radial ribs on adult whorls 
but is smaller, white to straw-yellow in colour, 
with wide umbilicus, radially lirate apex, and 

Si n m Zi 

^:c ,_ u-. 

1 1 i?ii ^t *? complex postnuclear spiral sculpture. 

^ s s - 

Z a " 


Type Material 

Holotype: AMC63496, Bryon Bay, NSW. Height 
111? if |S 1.15 mm, diameter 2.30 mm, H/D ratio 0.50. D/U 

111! II Si! ratio 4 - 00 < whorls 3 5 / g+ - 

Other Material 
zzif, |f is AMC31650, AMC63769, 4 specimens. Cape Byron, 

1J3| £5 ^3J NSW. Collected Lower, ex J.C. Cox; Kenilworth 

^i =1 5d5 S.F.,SEQ(26°37'S, 152°42 , E) rainforest, leaf litter(l, 

0MMO17284, 22 May 1980, J. Stanisic, A. Green); 
„ _ ^ MtMudlo, Kilkivan S.F., SEQ (26 o 01'S, 152M3'E) 

•c ir. v. t/i 

2 n J -c — — 

5 g |5|l 5S IS Araucaria/NVF, litter (13, QMM017283, 17 Jul 

1980, J. Stanisic, A. Green); Kalpowar S.F., SEQ 
(24°43"S, 151 °21 'E) MVF/Araucaria, litter (12, QM- 
M017287, 15 Jul 1980, J. Stanisic, A. Green); rain- 
Is forestwalk, Kalpowar S.F., SEQ (24 4TS, 151°21'E) 
ni§ MVF/Araucaria (1, QMM012576, 4 Sept 1982, 
Q ~ "-"*» ~~ ri " d AM/QM-ABRS); Below summit, Mt Warning, NSW 

(28°24'S, 153°16'E) (1, QMMO10511, Mar 1981, 

„ g ? §$ g? gs AM/QM-ABRS); Mt. Fort William, Kalpowar S.F., 

1 1 ? e |g|| |2 R |l SEQ, 833 m, CNVF (1, QMM016824, 3 Jul 1984, J. 

-"--£ -c --= Stanisic, D. Potter, K. Emberton); Whian Whian S.F. 

N. of Lismore, NSW, rainforest (1, QMM06294, 12 

S 5? Sept 1976, M.J. Bishop); Mt Guyra N.P., SEQ 

i . + ,| f 4i j (25°49'S, 152°35'E) MVYINVYIAraucaria (2, QM- 

| * ! !| 4 5 !3| MO12075, 7 Sept 1982, AM/QM-ABRS); Burleigh 

Heads N.P., SEQ, vine thicket, under logs (2, QMMO- 
o£ 16917, 21 Jan 1987, J. Stanisic, D. Potter); Freds 

Road, Mt Mee SEQ (27°05'S, 152°43'E) NVF, litter 
(1, QMMO15980, 28 Oct 1984, J. Stanisic, G. An- 
nabel); Cooloola N.P., SEQ (25°55\ 153°08'E) 50m, 
, CNVF on sand (1, QMMO6350, 25 Oct 1976, M.J. 

Bishop); sidetrack off Mt Archer Rd, Mt Mee S.F., 

- i 

2 5 



c ?o k O £ o 

-? ss-=s 

u | I I| HI SEQ(27°04'S, 152°4rE)(l,QMMO12180, 31 Sept 

Jjo^o |o o jo \t% 1982, AM/QM-ABRS; Byangum, Tweed River dis- 


= -= E 


tricl. Nth NSW, scrub, (2. AMC63779); Cape Byron. 
If Nlh NSW, <2, AMC 140493. 1 Mav 1958, L. Price); 

S s |jjj | Bvron Bav. NSW (2, AMC140483. C.F. Mc- 

Lauchlan); South Grafton. NSW (2, AMC63785); 
Mudlo Gap. Mudlo S.F., SEQ, MVF/Araucaria, 
|i |? U (26*0 rS. 152M4"E). (55, AMCI36654- QMMO- 

, ||, H - 5 a l| s a 12352, 2 Ocl 1982, AM/QM-ABRS); Yarraman 

A -i* = a *• c - _ s £ = Foresl Drjve Yarraman S F SE q MV p, (26^50'S, 

l5rS7'EJ, (3. AMC136587, QMMQ12262, 31 Sepl 

gS jS 1982, AM/QM-ABRS); Cooloola Nai. Park, SEQ, 

i - fi. s § k £ |3fb |2 SNVFonsand<2S"57*S 3 153 ,, 06 V E)(2 I AMC136489, 

? £ 2 3 r^.£ i= QMM0121 12, 6 Sept 19S2, AM/QM-ABRS); Bobhv 

Range S.F. SEQ, NVF. (24 a 57"S 1 I5T32'E) (lo", 

g , - AMCI36505. QMM012141, 6 Sepl 1982, AM/QM- 

Sg , lz gg ABRS); Blackbutl Range, c. 7.4 km S of Benarkin. 

I S H 1 5 5 * :- g|s ss SEQ. MVF/Araucaria (26\vVS. 152M I'E). (5. 

AMC 136565, QMM012239, 3] Sept 1982, J. 
_ Stanisic. D. Potter); Ml Fori William. Kalpowar SF, 

%2 If tl 12 ii SEQ, NVF (24' 3 39'S. !51"20*E) (2, AMC136759 

'-< z 

»S 5r* ~ ~ ^ 


— ■ ^ 
3 t 

QMMO 1269Q. 4 Sepl 19S2, AM/QM-ABRS); NE of 

Monio. Kalpowar SF., SEQ. vine thicket/scrub litter 

\ 5 (24'42'S, 15r2rF)(2.AMC137851,QMM013153, 

If || is 4 Sept 1982, AM QM -ABRS); NW of Miriam Vale. 

§§ ill 2 ^is l£ g 5 5 ^i* f Colosseum Ck.. SEQ, rainforest alongcieek(2423 , S, 

15T27-E) (7, AMC137S94, QMM013264, 6 Sept 
1 982, AM/QM-ABRS); Bulburin SF. S of Gladstone, 
SFQ. 541! m, CNVF-AI forest tvpe (24\31'S. 
151 1 29 , E) (1. AMCI52213. S May 1975, W.F. 
Ponder. J B, Burch. P.H. Colman): Eurirnbula S.K.. S 
Of Gladstone, SEQ, 10 m (24 5 H'S. 15T50'E) (4, 
AMC152214.AMC152215, 7 May I975. J.B. Burch. 
W.r. Ponder. P.H. Colman); Rundle Range, SE o! 
Rockhumpton, SEQ. 30 m (2.V39S, iSO'59'E) 
(many. AMC 152216. AMC 15221 7 ft May 1975. J.B. 
Burch. W.F. Ponder P.H. Colman). 

X - _ M A f- _ 

£• - * 3E v "I - 

%\ f SJ35 I| « 5 S |S= e Diagnosis 

Shell diameter L95-2.55 mm (mean 2.31) 
with 3 1/2+ to 3 7/S- (mean 3 5. 8+) loosely coiled 

- _ g| p fg ir whorls, last whorl descending more rapidly. 

11 fi fj = |3 ^ r . , |JL |t Apex and spire (Fig. 137c) flat to slightly 

= = z ~ £ elevated, SP BWW ratio 0.02-0. 1 9 (mean 0. 1 2), 

height of shell 0.97-1.31 mm (men 1.13 mm). 

* | II D ratio 0.45-0,54 (mean 0,50). Protoconch of 

a .7 I 3'8 to 1 1/2 whorls, mean diameter 574.8 am 

? ::^r ~ z - zz^s zz at 11 2 whorls, sometimes exsert but generally 

flattened. Apical sculpture (Fig. 13-7(1] of c. 12 
glossy, evenly spaced, fine spiral cords. 
Postnuclear sculpture (Fig. 137e) of complex, 
protractively sinuated. crowded radial ribs, Sl- 
116 (mean 92.1) ribs on the body whorl. 
Ribs mm 10.65-16.79 (mean 13.1 1 ).' Micros- 
culpture (Fig. 137c) of fine radial riblcts, 5—11 
between each pair of major ribs, and low spiral 
cords which form raised knobs at their intersec- 


Z -1 

= 7 

- -i 













J T 


■C - 5- 



- 1 







— n 


!_: „ 

— •? t £ 


t ;. 





= 7 

i- 3 








5 ^ 



i ^ < 






j " 









■ -■ 


. ■:■ 





*.«% _ j% 




. *• 


Jrjjmr '" jP" 

^■^r^L^f^ < >* *I ill 


■* Ik * ^ ""■ ***■ c* 
f •* k A- 

FIG. 137. Sinplovea intensa (Iredale, 1941). a-c, Bryon Bay, NSW. AMC63496, holotype; d-e, Mudlo Gap, 
MudloS.F., SEQ. QMM012352; f-g, Cherry Tree North S.F., via Casino, NSW. AMC154769. a-c, entire 
shell; d, apical sculpture; e, post nuclear sculpture; f, central and lateral teeth; g, marginal teeth. Scale lines 
as marked. 



FIG. 138. Sinployea interna (Iredale. 1941). Burleigh Heads N.P.. SEQ. QMM016917. a. genitalia; b, d. 
details of penis; c, talon and carrefour; e, ovotestis; f, pallial cavity. Scale lines as marked. 

MEMOIRS or- the Queensland museum 





: 3° S 

29 D S 


L 50 ' E 

151»E 132°2 '- 154°E 

FIG. 139. Distribution of Smphvea interna (Iredale, 

tion with the microradials. Sculpture continuous 
on the base^ Umbilicus (Fig. 137b) wide, V- 
Bh&pcd, last whorl decoiling more rapidly. 
diameter 0.55— (3.78 mm (mean 0.66 mm). D/U 
ratio 3.17-4.00 (mean 3.78). Sutures strongly 
impressed. Whorls rounded above and below 
the periphery (Fig. 137c). Aperture roundly 
lunate. Lip simple, slightly thickened. Columel- 
la weakly twisted toward umbilicus. Parietal cal- 
lus well developed, margins convergent. Colour 
brown. Based on 17 measured adults. 

Epiphallus (Fig. I38d) short, muscular, reflex- 
ing before entering the penis apicalty, internally 
with longitudinal pilasters. Penis (Fig. 138a) 
short; externally differentiated into an upper and 

a lower chamber; without sheath. Details ot 
upper chamber and epiphallus entry unavailable. 
Lower chamber with a semicircular (horseshoe- 
shaped?) pilaster (Fig. 138b). Penial retractor 
muscle (Fig. 138a) inserting on the penis ad- 
jacent to the epiphallus-penis junction. 

Radula (Fig. 137l.ej tvpical for tienus. 

Based on 1 dissected adult (QMMOI6917). 

RANtit and Habitat 

Humid sub-tropical noiophyll vine forest in 
northern NSW and SEO; drier microphyll vine 
forest between Gympie and Rockhampton, SEQ: 
and strand line vine forests at Bryon Bay, NSW T , 
Burleigh Heads, Cooloola and Eurimbula, SEQ. 
The latter habitat preference is unusual in the 
context of Australian species but as a remnant of 
a much wider Pacific Island distribution it is not 
unexpected. S. interna lives under the bark of 
fallen trees and has the typical grey markings also 
present in semi-arboreal Pacific Island Sinployea. 


5. interna is similar to£ cutyomplwla (Solem, 
1959) from Vanuatu in external appearance of 
penis, insertion of penial retractor muscle, shape 
of epiphallus. unusual muscular collar on the free 
oviduct and pallia! configuration (Solem, 1983). 
However. S. iniensa has more postnuclear ribs, is 
smaller in si/e, and has a less elevated spire. 

Subfamily ROTADISCINAE Baker, 1927 

This tuxon was introduced (Baker, 1927) for 
endodontoid snails with incomplete secondary 
ureter, and kidney with a rectal lobe much longer 
than pericardial lobe. Solem ( 1 983, p. 70) in- 
cluded the subfamily in the Charopidae for 
species with or without a complete secondary 
ureter and weakly to strongly bilobed kidney. 
He considered thai the peculiar apical sculpture 
and unusual variations in genital morphology 
were the main distinguishing features and in- 
cluded the Northern Hemisphere Rotadiscus 
Pilsbrv, 1926, Radiadiscus Pilsbrv and Ferriss, 
191)6, RtuUoconus H.B. Baker, 1927 and 
Radiodomus H.B. Baker. 193U, together with 
Microcharopa Solem, 1983 from the Pacific 

Rutaeharopagen. nov\ 


A combination of syllables from Rotadiscinae 
and Charopidae. 



Type Species 
Rotacharopa annabelli sp. nov. 


Very small to minute Charopidae, diameter 
range 1.77-2.46 mm, with 4 5/8+ to 6 1/8 very 
tightly coiled whorls, last descending more 
rapidly. Apex and spire weakly to moderately 
elevated. Apical sculpture of short, slightly 
twisted segments, organised into spiral rows, and 
weakly curved, low radial ridges. Postnuclcar 
sculpture of very crowded (densilamellata) to 
moderately spaced (annabelli), protractively 
sinuated radial ribs. Microsculpture of very fine 
radials and low, broad spiral cords which are 
raised at their intersection with the microradials 
to form an elongate bead. Umbilicus very nar- 
row, U-shaped to almost completely closed by 
reflection of the columella. Sutures impressed; 
whorls shouldered above and rounded below a 
laterally compressed periphery. Aperture 
roundly lunate. Parietal callus weakly 

Animal with slender foot and tail. Kidney with 
well developed, vaguely triangular, elongate 
pericardial lobe and a reduced rectal lobe. 
Ovotestis two teardrop-shaped lobes lying in the 
apical whorls of the digestive gland. Terminal 
male genitalia with epiphallus differentiated into 
ascending and descending arms. Descending 
arm of epiphallus with transverse thickenings 
internally. Epiphallus entering penis through a 
simple pore surrounded by a large fleshy pilaster. 
Penial retractor muscle inserted on the epiphal- 
lus. or on the epiphallus - vas deferens junction 
(annabelli). Penis large, barrel-shaped, with 
(kessneri), or without a verge, normally with 
longitudinal pilasters which may be modified 
(annabelli). Penis with short (annabelli), long 
(kessneri) to extremely long (densilamellata) 
preputial tubular extension. Vagina long, rarely 
short (annabelli), internally with longitudinal 
thickenings. Free oviduct short. Atrium without 
unusual features. Spermatheca typical. Radula 
with tricuspid central and lateral teeth (central 
slightly smaller) which have a long slender lan- 
ceolate mesocone and short, pointed ectocones. 

Distribution and Ecology 

Rotacharopa is confined to moist notophyll 
vine forests and drier microphy 11 vine forests and 
vine thickets of south to mid-east Queensland. 
Undescribed rotadiscines. with flatter shells and 
more open umbilici, also occur in this region 
(Stanisic, unpublished). Rotacharopa kessneri 

and R. annabelli have been collected in close 
proximity to each other near Gladstone, SEQ, 
and further fieldwork may demonstate sympatry. 
Both species have modified penial surfaces 
which indicate possible microsympatry but this 
sympatry may also involve undescribed taxa. 

The two southern species, R. densilamellata 
and R. kessneri live under logs, whilst R. an- 
nabelli is a litter dweller. 


Other charopids reviewed in this study which 
have similar-sized shells lack the peculiar apical 
sculpture of Rotacharopa. Rhophodon has a 
flattened shell with wide umbilicus, apical sculp- 
ture of prominent radial ribs, and apertural bar- 
riers; Discocharopa has apical sculpture of 
prominent radial ribs and weak spiral wrinkles, 
and a saucer-shaped umbilicus; Omphaloropa 
has a very wide cup-shaped umbilicus and con- 
tinuous spiral cords on the protoconch; and 
Sinployea intensa has continuous apical spiral 
cords, fewer whorls and a more open umbilicus. 
Microcharopa Solem, 1983 from Fiji is readily 
distinguished from Rotacharopa by its lower 
whorl count, smaller size, flatter shell and very 
wide umbilicus. 


1 .Postnuclear sculpture of widely separate radial 
ribs (mean ribs/mm 6.62) (Fig. 140a) 

Postnuclear sculpture of more crowded radial 
ribs(Figs 143a. 146a) 2 

2. Postnuclear radial ribs very crowded; umbilicus 
more open (Fig. 146b), mean D/U ratio 4.93 

Postnuclear radial ribs less crowded; um- 
bilicus more closed (Fig. 143b), mean D/U 
ratio 9.71 kessneri 

Rotacharopa annabelli sp. nov. 

(Figs 140-142; Tables 26. 29) 

ETYMOl ogy 

For Graeme Annabell who provided assistance 
with collecting. 

The widely spaced ribs (mean ribs/mm 6.62). 



-J u-, 




0" 3" 


~ ^ * 
_ c x 



I- 5r 5* 

33 23 

s 9 


2 * x § « g « Sin H^ 5 g gfi g£ 

1 1 1 s ^ fill I if I 11 i I ll| i Is II 

a — — w w 

5? 9-. Id =9 Ti9 =5? _ ~«T „ „ 3 

: - 

- z 

y in 

■W — aft 

r- -h 

SS2 q3 =* 9^ 5 ^ 5 * 

0<3d =59 39 «? «? *T 

T *! 7 * J _ * J - a 3criA (N f. r!« 9*1 

il s 22 si|2 b Is s 12 $ s gii ? lit 

as 9 

t -r -T -r rt 

-T X DC 

- - 

5 U 




















5 ~ 











= s 





7 C 




— f\ 

li IS flip II I M 1 i| 1 



large whorl count, narrow umbilicus, details of 
internal penial sculpture and position of pcnial 
retractor muscle insertion separate R. annabelli 
from its congeners. 

Type Mavhrim 

Holotype. 0MMO17.UUJ. Ml Archer, Rock- 
hampton, MEQ Jeaf litter. Collected by G. Annabell. 
23DecL978. Heigttfflf shell I 23 mm. diameter 184 
mm. H D ratio 0.65. D U ralio 8.3b, whorls 6 1 8. 

■ atypes: OMM08348. 26 specimens, same col- 
lection data as holotype; Ml Archer, Rockhampton, 
MBO. about one third of the uay up (23 c 2t'S, 
1 50 r 35 *E). remnant vine thicket in gully (62.QMMO- 
II659 ? QMMOH919, 30 JUn 1982, J. Sunisic): 
Gavial Creek, Bouldereombe MEO (23 3-i'S. 
IMJ 2SL-I. MVF. litter (24, QMMO16S07, 9 .luh 
1983. J. Slanisic. D. Potter); BouJdcrcotnhc. W ol' 
Rockhampton, MEO, rainforest litter (10. QMMO- 
I2S27. 3 Feb 1980. V. Kessner); c. 7 km E Ol Ml 
Morgan. MEQ (23*38*5, 150 L 28'E) MVF/NVF, litter 
(KQMMO1730L 11 Jul 1980,1 Sianisie. A Onsen; 
5.0MM0131l7,29Jun r9&J.£tanfeic;4.QMMO- 
U63S.29JUW J982;J. Slanisic): Wside of Mt Moore. 
SW of Gladstone. MEQ. in patch of vine thicket, Inter 
(24'02'S, !.SJ s 05"E)(5,AMCLV7875,QMMO132u3. 
5 Sept 1 98:, AM/OM-ABRS). 

Other Materia! 

Caiilungal, meo (i. AMC264. ct Mowon); W of 

Gladstone. MEQ. under leaves on ground in bottle tftC 
scrub (I. AMC32996, Aug 1908, SW. Jackson). 


Shell diameter 1.77-2.36 mm (mean 2 .(10 mm) 
with 5 3/8 to 6 1/8 (mean 5 3/4-) tightly coiled 
whorls, the last descending more rapidly Apex 
and early spire (Fig. 140c) flattened to 
depressed, SPBWW ralio 0.1 1-0 ,26 (mem 
M.16). height 1.13-1.54 mm (mean 1.28 mm!. 
II (D ratio 0.52-0.69 (mean 0.63). Protoconchof 
I 5/8 to 1 3/4 whorls, mean diameter 343.5 u.m 
at 1 1/2 whorls. Apical sculpture (Fig. 140e) of 
short segments arranged in 14-1S spiral rows 
crossing a series of curved, broad radial ribs and 
ridges. Postnuclear sculpture (Fig. I40f,g) of 
hue, widely spaced, weakly protractively 
sinuated radial ribs, 34-51 (mean 41.3) on the 
body whorl. Ribs/mm 5.60-7.45 (mean 6.62). 
Microsculpture (Fig. 140f) of very fine radial 
riblets, 9-16 between each pair of major ribs, 
crossed by low spiral cords which are raised at 
theirjunction with the microradials. producing a 
beaded effect Umbilicus(Fig, I40b,h) very nar- 
row. U-shaped to completely closed by reflec- 

tion of the columellar margin, diameter 0.23- 
0.27 inm (mean 0.25 mm). DV ratio 6.85-3 54 
(mean 7.66). Sutures impressed. Whorls 
shouldered above and rounded below a laterally 
compressed periphery (Fig. 140c). Aperture 
roundly lunate. Lip simple, columella slightlv u> 
strongly reflected over the umbilicus. Parietal 
callus well developed. Colour gulden-yellow 
horn. Based on 19 measured adults. 

Epiphallus (Fig. 14 la) muscularised with as- 
cending and descending arms, internally with 
well developed transverse thickenings (Fig. 
141b). entering penis through a simple pore sur- 
rounded by a large circular pilaster. Penis (Fig. 
141b) elongate, with a short basal extension (Fig. 
141 a), internal fy with strong pustules and ir- 
regularly shaped pad-like thickenings (Fig. 
141b). Penial retractor muscle (Fig. 14 1a) in- 
serted at the epiphallus - vas deferens junction. 

Radula (Fig. 140d) with central tooth slightly 
smaller than latctals. 

Based on 3 dissected specimens (QMMO- 

Rangf and Habitat 

In leaf Titter in dr\ microphyll Nine forests and 
vine thickets between Gladstone and Rock- 
hampton. SEQ. 


The shift in insertion of the penial retractor 
muscle to the epiphallus - vas deferens junction 
is most likely related to whorl count increase. 

Rotacharopa kessueri sp. nov. 
(Figs 143-145; Tables 26, 29) 


For Vince Kessner. in recognition of his con- 
tribution to Australian malacoK 


R. kessntn differs from R, densilamellata in 
having coarser sculpture, a less elevated spire 
and narrower umbilicus. Anatomically it lacks 
the very long penial extension of R. densila 

Ivi'i MATOtlAl 
HOI oTYFT; QMMf)17302, e. 8 km from KatpowSt 

on fireclay Rd. Kalpowar S.F.. SEQ (24 - 43'8, 

l5r2i'E), M\T/Antucaria, litter. Collected by J. 
Stanisie- A. Green. L5Jul W80. Height of shell 140 
mm. diamelei 2.28 mm. H'D ratio 0.61, D/U ratio 

1 1 |2, whorls 5 i/44 











FIG. 141, Rotacharopa annabellisp. nov. c. 7km E of Mt Morgan. SEQ. QMMOM633. a, genitalia; b, details 
of penis interior; c, ovotestis; d, hermaphroditic duct; e, pallial cavity. Scale line as marked. 

FIG. 140. Rotacharopa annahelli sp. nov. a-c, Mt Archer, Rockhampton. MEQ. QMMO 17300, holotype; d, 
c. 7km E of Mt Morgan, MEQ. QMMOl 1633, paratype;e-h. same data as holotype,. QMM08348. paratype. 
a-c. entire shell; d, lateral teeth; e. apical sculpture; f, posijiuclear sculpture; g, details of microsculpture; h, 
base. Scale lines as marked. 




3 a S 

1 5 Q a S 

FIG. 142. Dislribuiion of RuUufhimpa unmihclli sp. 

Ill A 

Par VTVn : :>; QM MO I 7503, 6 specimens, sninc in 
as holoivpe; Mi fntonaneman. via Childers, SEQ, 
rainfall (3, GMMQ17304, 7^ Feb I'JM.R. Raven. 
G Mpitli " high Upon rw wbankbeside road. Canu 
Gurge, SEQ (24 41 S I5(TS8"Ehrai0|bresl, litter (2, 
0MMQI7305, 1 4 Jul J9SI. J. Scuiisie. A. Green); 
Dan Dan Scrub. SEQ. SEVT. liuei (4, QM MO16802, 

HI Ma> 1984, J, Stanisic, D. Potter); Mi Biggendcn. 
SEQ. SEVT on limestone oUlcrop, Iftter (I, QM- 
MOlf»S3n. 3 Jul l"N4, J. SUinisie. D. Poller. K. Em- 
bermn): S\V of Miriam \ r ale. QUI. Colosseum Ck. 
rslnfoKSi along creek (24 r 23'S. 13 I 2TV) [S. 

^MC137S95*QMMQ 13265, 6 Sept I982.AM/QM- 
ABRS): Mi Fort William, Kaipow; rS.F..Si 0. NVF 
t24 y 39'S.15r^0'E)(3,AMCl3675S.0MMO12599. 

4 Sep! 1982. AM QM-ABRS): Woi ( -Jiilders, Lime- 
stone Ck S.F (25' I5*S, 15E55'E) (5. *MC13783t), 
wMMiH -i:"\ 3 Sep! 1982 AM QM \BRSj. Rjin- 
toresu walk. Kalpowar S.F.. MVR SEQ (24'41'S, 

I5L2LE) (3 ( AMC136743. 0MMO12575. 4 Sept 
1982, AM. QM-ABRS). 

Otker Material 

Bulburin S.F.. S of Gladstone. SEQ. 5&0 m(24"34'S < 
15E2y'E)(l. AMCi522UVSMay 1975, J.B. Burch, 
W.p. Ponder, P.H. Culman); Bulburin ST., S of 
Gladsione, SEQ, 540 m, CNVF-A1 forest type 
(24 a 3-rS, l51 a 29 ? E)()l, AMC152211,X May 1975, 
W.F. Ponder, J.B. Burch. P.H. Colmaii); Bobbv Range 
SF, SEQ, NVF (24'37'S. I51 B 32 r E) (12, AMC- 
136504, QMM012138. 6 Sept 1982, AM, QM- 
ABRS); Clifton Range ST.. S\V of FairliesKnobN.P., 
SEQ t Vin£ifticketMrflMCd«d(2S'34"S t l52*I6 , E){2 
AMC136672, QMM012376. 2 Ocl 1982, AM QM- 
ABRS); c. 12.1 km NW Granite Creek Crossing, on 
Bobby Range - Ml Perry Rd. SEQ. NVF /palms, under 
logs (2, QMM016S44.' 16 Sep! 1QS5, J. Stanisic. D. 
Potior); summit Ml Booroon Booroon, SWof Miriam 
Vale, SEQ. MVVt'Araucariii, among rocks (5, QM- 
M016849. I 7 Sept 19S5. J. Slanisic. D. Potter). 


Shell diameter 1.85-2.46 mm (mean 2. 15 mm) 
with 4 7/8- to 5 5/8+ (mean 5+) tightly coiled 
whorls, lust whorl descending more rapidly. 
Apex flat (Fie 143c). spire slightly to moderate- 
ly elevated/ SP/BWW ratio" 0. 10-0.24 (mean 
6.15), height 1.15-1.60 mm (mean 1.31 mm). 
H,D ratio 0-57-0-65 (mean 0.61 ). Pmtoconchof 
1 5/8 whorls, mean diameter 385.7 u-m at I 1/2 
whorls. Apical sculpture (Fig. 143c) o( short 
segments arranged in spiral rows and curved, 
broad radial ridges and ribs. Postnuclear sculp- 
ture (Fig- m! fine moderately spaced to 
crowded, weakly protractively sinuated radial 
ribs, 57-146 (mean 95.8) ribs on the body whorl. 
Ribs mm S.S3-IV.32 (mean 14.19). MiCtOS- 
culpture (Fig. 143f) of tine radial riblets. 5-10 
hetween each pair of major ribs, and low spiral 
cords w bidl are raised at their junction with the 
microradials tu form d bead. Umbilicus (Fig. 
143b,h) narrow. U -shaped, diameter 0. 16-0.20 
nun (mean 0.23 mm). D/U ratio 7.23-11.90 
(mean 9.71 ) 4 Sutures impressed, whorls 
shouldered above and rounded below a broadly 
rounded periphery (Fig. 143c). Aperture broad, 
roundly lunate. Lip simple, columella margin 
Slifthth twisted toward umbilicus. Parietal cal- 

FIG. 143- Rj'jchttropu k&ssm ib$ M . c. c. Skm From Kalpow-ar on Fireelas Rd Kalpowaj 5,F.« SFQ 
OMMOP3Q2. hototypes c. I2.lkni No] Granftc Ck cmssing BoKb\ Range slq QMMOl6844;c-h, 

Col liseum ( k. SW of Miriam Vale, SEQ, QMMOl 3265. parjt\ pc. ±-c. entire she 1 ,!; d. Centra] and lateral 
■ ■ h ipfi |1 sculpture: f.posl iCl tffe & del i|s Ol |Tjicni5C0lplUf£ h. base. Se.ile lines ;;s marked. 



(K — 








FIG. 144. Rotachawpa kessneri sp. nov. Summit. Mt Booroon Booroon, SW of Miriam Vale. SEQ. 
QMM016849, a. genitalia; b. details of penis interior: e. hermaphroditic duct: d, talon and carrefour: e. 
ovolestis: f, pallial cavity. Scale lines as marked. 




26 Q S 

J i 1 .\ >\ 


1 5 : " e 



FIG 145- Distribution of Rota cha ro pa kcssncr: sp. 

lus developed. Colour beige lo light-golden 
horn. Based on 18 measured adults. 

Hpiphallus (Fig. 144a) with short, narrow as- 
cending arm and much wider descending arm 
that has (hick, transverse, fleshy pilasters inter- 
nally (Fig. 144b). Epiphallic pore (Fig. L44b) 
simple, surrounded by a weak fleshy collar. 
Penis (Fig- 144a) with a wide upper chamber, a 
low er, cylindrical section, and a basal extension 
(Fig, 144a). Internally- penis with strong pus- 
tulations arranged into longitudinal thickenings 
(Fig. 144b) and an apical conical verge Pcnial 
refractor muscle (Fig. 144a) inserting on the 
cpipnallus at the point of reflexion. 

Rsrdula (Fig. 143d) typical. 

Based on 4 dissected specimens (QMMO- 
[6849 OMMOIhS44)- 

Kangf WO M AH1TAT 

Under logs in dry microphyl! vim ■ and 

Vine thickets between Biggenden and Calliope, 
SEQ. It has also been found in the wetter Warm 
Subtropical notophyll vine forests oi the Bobby 
Range and Mt Fort William. The species appears 
lo be able to tolerate drier conditions and occurs 
ill •fringe' rainforest in the Cania Gorge, near 
Momo, SEQ. 


P. kessneri may be sympatric with R. annabcllt 
because the two species have been collected 
within only a few kilometres of each other west 
of Gladstone. SEQ 

Rotacharopa densilamellata sp, nov 

(Figs 146-14S; Tables 2o. 30) 


Latin (ienSUS, erowded; and lamella, plate; 
referring to (he perinuclear sculpture of crowded 


The small, globose shell with many tightly- 
coiled whorls, elevated spire, crowded, fine 
radial ribs, and small umbilicus, identify/?, den 
silamcltaiQ in SEQ. Anatomically the peculiar 
preputial extension of the penis (Fig. 147a) is 
distinctive. The only other species apt to be 
confused with R. densilamellata is the sympatric 
Egilomen globosa, but it has adult sculpture of 
fewer, broad radial ribs, a completely closed 
umbilicus, and a sinuate lip with apcrtural sinu>. 

Type mah-.riai 

HoldtYpE: QMM017296, Kenilworfll S,F. S SEQ 
(26 J7S, 152-42'E) Tainfore&L lear Kller, Collected 
h\ 1 Sianisic -v Green Height of shell 1.38 mm. 
diameter 1.95 mm. H/D ralio 0.71. D/U ratio 5.27, 
VtthorM -4-. 

Parav. ft Si QMM017297. 4 specimens, same col- 
lection data as h id ivpc, sidetrack off Mt Archer Rd, 
Ml Mee S.F.. SEQ (27 S 04"S. 152*41' E) 
NVK Araucatia (20, QMMOI5081, 6 Sept 1983. J. 
Slanisie. D Poller). Kemlworlh S.F., SEQ (26°35 5 
l52*3fVE) mixed NVF (12, QMM06285, 18 Way 
197^ M.E Bishop); Fred'a Rd, Mt Mc-e, SFQ 
(27*05 S. 152M3E) rainforest, leaf litter (11. 
QMM08347 14 \pt |980, J, Stamsic H, Mall, A. 
Green, o.. QMMO I 5984. 28 Od 1 984. J. Slanisic. G, 
Annabell): Little Yafrba Creek. Imbil S.F.. SEQ 
28'S, |52 B 38'E) NVF (4, QMMO12018, AMC- 
136435, 8 Sept |9R2, AM.QM-ABRS); Kenilworih 
S-F-. SEQ <2' : ' r 4ri S, 152'36'E) Araucaria, leaf litter 
(12. QMM0834o 22 May i ,J So. J Sianisic 
Green); c. 25.5 km and 28.5 km from Goatnburra OH 

OUiburrt S.F, Road, SEQ N\ I -".'Palms /AfaUt ■ 
(8, QMM012nM4_ AMC136813 7 Dee t9S I 
AMQM-ABRS; 1, QMMO 11 125. 16 Mar 198 1, 
AM QM-ABRS); Rainforest Purk\ nearMaleny. Of 
Maienv-Momville Rd. SEQ NVF. under log f l.QV 
MOi 1883 5 Dec L982 I i( c Kenilworth SJ 

SEQ (26 35'S, i 52*42 r £ | c. 80m, rainforest/eucalypt 


emergents (22. QMM062S4, 3 May 1976, M.J. 

Bishop); Cunninghams Gap N.P.. SEQ. near monu- 

%*Ji H ? : mem, 755 m. SNVF/ferns <>- D H4'S. 152"24'E) ( 104. 

AMC136805, QMMOI2676. 7 Dec 19X1. AMQM- 

ABRS); Montville Rd near Kondalilla Falls N.P.. 

|?M SEQ, vine scrub on roadside (2640 S, 152 B 52'E) (2. 

AMC13641 1. QMMOl 19S0. 8 Sepl 1982, AM/QM- 

ABRS): Cunninghams Gap N.P., SEQ, near monu- 

menl. MW/Araucaria (28°04 t S, I52°24'E) (10, 

= ! AMCI28609,QMMOI0993, 16Mar 198L AM/QM- 

- * ll e ? i s | ABRS); sidetrack off Ml Archer Rd. Ml Mee S.F., 

1 - «- * * * * - SEQ NVFMrcwcflrifl (27"04'S, 152'41'E) (6, 

1982. AM/OM-ABRS). 

j i - >^ n 2 ^ T ~ ~ "* I ^ 

J- 1-1- 1*1 - = * 

.7 « - «'*T*7 

~ i -C— r* x ^ — "" ' i ■"■ * 

Othfr Material 

Little Yabba S.F„ SEQ (26*34'S, I52°39*E) c. 160 
m. euealypl and vine forest (I. QMMO6301. 3 Ma> 
1976, MJ. Bishop); Maiala N.P., Mt Glorious, SEQ, 
NVF. leaf litter (LOMMO 11 853. 20 Jun 1982. MSA 
party); Boomhana N.P., Ml Nebo, SEQ, wet 
scierophyl](l,QMM06298,Aug 1976, M.J. Bishop); 
Campbell's Camp. Kenilworlh S.F.. SEQ (26*42*S, 

Z -- ?r^ ^S §3 = 2 

l\\ 3 | Ull j 1^237"E)CNVF 1 leariitlertl.QMMOl7298- 1 Jul 

> _ : -i 

X r ' 

i = 

1980, J. Stanisie. A. Green); Cunningham's Gap, 
SEQ. leaf litter (I. QM MO 1 7249. IS Jun 1979, G. 

*l y l» S5j %S^£z~ AnnabellKc. 0.3 km N of Little Yabba Ck. on Malenv 

Z 5 7 = % 

H =fi - " |j 325235 ^ _ r Kenilworlh Rd. Kenilworlh S.F.. SEQ (26'37'S, 


152°42*E) litler (2. QMMG13434. 19 May 1983. K, 
2 J Collins. J. Slanisic). 

1 < ll Diagnosis 

< = . — .. - _ 

: EJ r !| E ; 1 ! J r 2 Shell diameter 1.79-2. 16mm(mean 1.94mm) 

= with 4 5/8+ to 5 7 8- (mean 5 1/8+) tightly coiled 

2 * whorls, last whorl descending slowly. Spire 
z~ elevated (Rg. 146c), apex tending to be flat- 

5 1| II |5! ? § S tcncd ' SP/BWV/ ratio 0.21-0.53 ("mean 0.34). 

height 1.23-1.52 mm (mean 1.35 mm). H D 
ratio 0.67-0.78 (mean 0.71). Pmtoeonch of 1 
1 2 to I 5 8 whorls, mean diameter 428.3 p.m at 
I 1 2 whorls. Apical sculpture (Fig. 146d.e) of 
short segments arranged in 19-25 prominent 
crowded spiral rows overlying broad. Curved 
radial ribs and ridges. Postnuclear sculpture 
(I ig. !46f) of narrow, crowded, weakly protec- 
tively sinualed radial ribs, 93-206 (mean 139 ton 
the body whorl, whose intcrfjees are 4-5 limes 
their width. Ribs/mm 16.18-36.68 (mean 

KG. 146. Rottn'hiin.iptt (icnsiliimvliuhi Sj>, nov. a-c. 
Kenilworlh St .SEQ QMM( J \~y>h. holotypeid-f. 
same data as holuUpe. QMM08346. para type; g-h. 
Cunningham's Cap, SEQ- QMM012B76. paratype. 

a-c. entire shell: d. apical sculpture; e. details of apical 
spiral cords; I. pasi nuclear sculpture, g. central and 
lateral teeth: h. marginal teeth Scale lines as marked 

. - — — 

i - 

= ^ 




7 - 




















f f 





g - 







$ 7 

2 3 



> * 

; ^ 


9 1 



- 2 

^ s 




= 2 

j! 2 


5 5 














FIG. 147. Rotacharopa densilamellata sp. nov. Cunningham's Gap N. P., SEQ. QMMO 10993. QMM012676. 
paratypes. a, genitalia; b, details of penis interior; c, hermaphroditic duct and carrefour; d. talon; e, ovotestis; 
f. pallial cavity. Scale lines as marked. 








151 D E 152°E 153°E 

154 °E 

FIG, I4S. Distribution of Rtrt&charapa dcnsilamel- 
lata sp. nov. 

22.10). Microsculpture (Fig. 146f) of very fine 
radial riblets, 6-10 between each pair of ntajot 
ribs, and equally fine spiral cords, which are 
raised at their intersection with the microradials 
producing a beaded effect. Umbilicus (Fig. 
146d) narrow, U-shaped, diameter 0.35-0.51 
mm (mean 0.41 mm). D/U ratio 4.08-5.27 
(mean 4.93). Sutures impressed. Whorls 
shouldered above and rounded below the 
periphery (Fig. 146c). Aperture roundly lunate. 
Lip simple, columella slightly deflected toward 
umbilicus. Parietal callus strongly developed. 
Colour light-brown. Based on 14 measured 

Epiphallus internally with transverse pilasters, 
entering penis apically through a simple pore 
(Fig. 147b). Penial retractor muscle (Fig. 147a) 
inserted on epiphallus at the point of reflexion. 
Penis (Fig. 1 47b) thin walled, with a very long 
preputial extension (Fig. 147a). internally with a 
simple, apical epiphallic entrance, and strongly 
pustulose walls; pustules arranged in vague lon- 
gitudinal rows (Fig. 147b). 

Radula (Fig. 146g ? h) typical. 

Based on 4 dissected specimens t'QMMO 
IU993, QMMO 12676). 

R vnoi \m.i Habitat 

Disjunct distribution in cool subtropical 
notophyll vine forests of the Great Dividing 
Range in southern Queensland, and the warme* 
subtropical notophyll vine forests of the D"- 
Aguilar, Conoudale and Blackall Ranges, SEQ 
It lives under logs 


R. densilamellata has a more elevated spire 
than cither of its congeners and the longer prepu- 
tial tube may he a secondary modification related 
to this height increase. 

There appears to be considerable interpopula- 
tional variation in the shells ofR. densilamellata 
but much of the available material is subaduli 
and not suitable for statistical analysis. 


Attempts to relate the Charopidae reviewed in 
this study toextralimila! charopids remain tenta- 
tive because of the lack of knowledge about 
some of the major regional charopid faunas, 

Solem (19flJ) reviewed the Pacific Basin 
species but only briefly mentioned the large 
radiations i)\~ New Caledonia, Lord Howe Island 
and Norfolk Island. Climo (1969, 1970, 1971, 
1978. 1980, 1981, 1983) reviewed a large num- 
ber of New Zealand species, but omitted impor- 
tant comparative data on shell sculpture and 
anatomy. Additionally, a large number of New 
Zealand charopids are still undescribed (Solem, 
Climo and Roscoe, 1981. p, 454). Locally, the 
Australian charopid fauna has thus far received 
scant attention with only Solem (1984). Smith 
and Kershaw (1985) and more recently Sianisic 
(1987) presenting significant detail on a handful 
of species. Nevertheless broad comparative ob- 
servations are possible. 

shell Patterns 

The extent of conchological variation in 
Australian species is similar to that in New 
Zealand charopids but contrasts with the relative 
stability shown by Pacific Basin taxa. 

With the exception of Mlcroeharopa mimula 
Solem, 1983 and the widespread Discocharopa 
aperta (Mollendorff. 1888), apical sculpture in 
Pacific Island species consists of spiral cords. 
Patterns of apical sculpture in New Zealand 
species may be spiral, radial, reticulate, smooth 
or otherwise modified (Climo, 1969b, 19"?0, 
1981, 19K3), Among species examined in [bis 
study, a combination of curved radial ribs and 


tSOFTHl i 

I - - ..■,.... Spiral apical 

iCUlptUI lominant among Pacific Island 
species, is present oni\ in Sci&mvdea, Onn 
phuJoropa varicasa and the extra! itnitauy i 
luted ifypioyw uitrnsa. 

(he fiiiicd apical sculpiurc of Bfcmphalopa, 
Xutuiliropii omicron and Gyracachlen atrtisi- 
iin>! (Fig* 60c 63d- 54e, 57c) has noi he 

reported tOt Pacific Island Ot New Zealand 
( NarOprddC bUi is seen in the primarily Papua 
New (.imncau Pilshryvhuwpa, Solem fl" 
I ci or dud this sculptural feature for 
IHlsbrychoropu atmidus (Odhner, 1917) from 
the Northern Territory, Australia. These taxa 
may be distantly related bul considering that this 
cutptvrc is derived from the more common 
rrtteulale paLlern (see discussion tor Gyro- 

eocMeacuntsiana)* it is likely to have devciopi I 
independently m I he New Guinciin genus, ihc 
pitted apical sculpture seen in Ngairea murphyL 

I rlotnolii VOfitQFtttSi / '.'n\\ct)hi<i [)n>lns< rohtctil- 
niit and Hfdlvyuamvhu (Figs 9f, 23e, 2frd< 29g, 
fi \h isgrossb similarlothal ofthe New Zealand 
Paracharopfi Oimo, 1 983- to Ihcse eases how- 
ever, the macroscopic similarity has quite dif- 
fcrcrtt underlying mi*, restructure which 
correlates with major anaiomical dillcrenecs. 
Apiqal sculpture of Raiacharopa is identical to 

ihai in the Pacific Basin AhcmcJutmpa mimulti 
rrii 1963 and North American Radi&disctiS 
millccosiams Pilsbry and Fcrriss, 1906 although 
whorl numbers and coiling pattern indicate im- 
portant regional differences in these rotadiscines. 

I !k postnuctear sculptures Of Australian, New 
Zealand and Pacific Island species share 
similarities which are probably convergent and 
related to sharing a common niche. However, in 
contrast with the coiiseivaiisni tit Pacific Basin 
species, the Australian charopids display the same 
rich range o] variation seen in the New Zealand 
fauna. Sculptural reduction, common among New 
.V.ii.nui 'niit Australian species i-. ian in Pacific 
Island charopids. Comparison of rib counts of 
Australian (Table 1) and Pacific Island species 
(Solem, l u So. Table J ) shows that the median rib 
eounr in (he latter is greater It is premature 10 sav 
whether this will prove tfl : i H basic difference 
between these iwochaiopid faunas. 

The most unusual poslnuclear sculptural ele- 
ments among Austialian suhliopieul Charoptdue 
.ue the incised spiral lines of Ngairea. l.cnwch- 
hta and Itl lesser extent Mmsnmila- I his Icalurc, 
which, is more typical (if helicarionids. is con- 
fined JO the group iyf genera with incomplete 

ut ,, . ureters NttiMfl . . ■ . LUirnl ih .-• 
reported in any cxtralimital charopids. 

Shell shape til Australian and New Zealand 
species is mure diverse than tbalpi Pacific W i 

l :■;■■ A'hicl with a single exception, arc a 

. i i laj spread over a number 

'! i I ind (Solcm, 1983). It appears : . rl 

■ ustraKan and New Zealand charopids have had 

ei more complex evolutionary history involving 

many radiations ;_md extinctions andcr condi- 
h mtra-site diversity where competi- 
tion for food and space has led to considerable 
variation in shell form. Consequently shell 
L'ie\;uion «nd associated umbilical dosun 

more frequent occurrence in Austral ii d New 

Zealand charopids than in Pacitic Island species. 
Accordingly. Solem and Climo ( 1985) showed 
thai development oi' a keel, periostracal pi- 

nid shell coloui in New Zealand species arc 
sile-pielereuce correlated and not based on 
monophylelic assemblages. These authors also 
found thai in the de\ elopment of large sympalric 
snail CQtnmui i'. iVolutjori appears to favour 
an increase in diversity of shell shape. In such 
situations convergences u ill occur and are hi [fl- 
ighted in the Australian-New Zealand species 
pairs o\' fiedteyoconcha delta Serpfio kM 
(Cira\ , ISM) and Seiamcdea seticcstata - 
Sutcrui idc (Gray, 185(1). 

Barrier formation is extensive among Pad He 
Island and New Zealand Charopidae but occurs 
to I [C5SCI \tent in Australian subtropical taxa. 
Solem (IV73; 1983; p. 15) discussed ihe 
polyphyletic origin of barriers in Pacific Basin 
species and Climo (1978) provided evidence for 
polyphyletic origin in New Zealand speeies, 
I he lorm, number and microsculpture (Figs 7Sc, 
Bid) of the barriers in Rhaphodon make :ii, i 
comparable to the barriers in Pacific Island taxa 
such as Semperdofl Solem, 1983, and New 
Zealand Plxchndon Ancev 1 N.SN However the 
reduced structures of / ctomoh CQMartus (Fig. 
bSa.b) tire It"-'- i usik refned to Pacific taxa. 
beinu grossly similar to the barriers seen in New 
Zealand / vctala Ircdale. 1915 (Climo. 197B). 


Pallia! configurations of species reviewed are 

most similar [u ihose ol New Zealand 

I i iropidac All endemic Pacific Island species 
possess:! kidncv in which the lobes are subcecal, 
equal or the rectal lobe is much longer than the 
pericardial lolv (Solem. 1983). In contrast New 
Zealand species for which d;i!n is available 
(Climo. I'JNO. 1 983; Solcm, l 4 >N3f. havepericar- 



dial longer than rectal lobe. Leiomola con tort us, 
Egilomen cochlidium, E. globosa and Sinployea 
intensa have equal to subequal kidney lobes 
(Figs69g, 129d, 132f, 138f) while the remainder 
have the pericardial lobe much longer than the 
rectal lobe. 

An incomplete secondary ureter was pre- 
viously recorded in the Charopinae only in 
Oreokera cumulus (Odhner, 1917) and O. nim- 
bus Stanisic, 1987 from north Queensland, and 
Amphidoxa marmorella (Pfeiffer, 1850) from 
Juan Fernandez. This is a primitive condition in 
the Charopidae (Solem, 1983; Stanisic 1987). 
Its presence in genera such as Hedleyoconcha, 
Setomedea and Ngairea, which all show typical- 
ly disjunct relict distributions, is not surprising, 
and a similar condition in southern Australian 
and New Zealand species would not be unex- 
pected. Shortened secondary ureters have been 
reported for some extralimital Rotadiscinae 
(Baker, 1 927) but were not present in rotadiscine 
species reviewed here. 

Intrusive mantle-gland tissue was noted in the 
majority of species examined. This structure is 
rare in Pacific Island species and present only in 
Graeffdon graeffei (Mousson, 1869) and 
Semperdon xyleborus (Solem, 1983). Extent of 
its occurrence in New Zealand species is not 
known but it has been recorded for Charopa 
coma (Grav, 1843) and Phenacohelix pilula 
(Reeve, 1852). 

In local species the ovotestis usually consists 
of two clumps of palmately clavate to finger-like 
lobes of alveoli oriented parallel to the plane of 
coiling and embedded in the apical whorls of the 
digestive gland. Occasionally lobes assume a 
right angle orientation and may vary in size and 
number. All Pacific Island species have ovotestis 
lobes parallel to the plane of coiling. Informa- 
tion on New Zealand species is scarce although 
both parallel and right angle orientation have 
been reported. Stanisic (1987) considered that a 
right angle orientation of the ovotestis was a 
primitive character. However, from this study it 
appears that ovotestis orientation is related to 
shell height. Hedleyoconcha, Ngairea and Len- 
webbia have elevated shells and right-angle 
orientation of the ovotestis. In contrast, 
Setomedea, which possesses a primitive pallia! 
configuration, has a less elevated shell and the 
typical parallel orientation. Gyrocochlea 
vinitincta has a biconcave shell, with secondarily 
enlarged body whorls, and has the ovotestis 
oriented at right angles. 

Number and size of clumps arc variable and 

probably related to variability in whorl count and 
shell size. Ngairea murphyi and Hedleyoconcha 
delta have high whorl counts and multiple 
clumps of ovotestis (Figs 10d, 24e) while the 
small Sinployea intensa and Letomola contortus 
have single bilobed clumps (Figs 69c, 138c). 
Among Pacific Island and New Zealand species 
the 'two clump' condition is typical. Some 
Pacific Island Semperdon Solem, 1983 and 
Sinployea Solem, 1983 show reduction to a 
single clump but there are no examples of the 
increase in clumps recorded in some Australian 
and New Zealand species. 

The penis of Pacific Island species is pear- 
shaped with the penial retractor muscle inserting 
on the penis head, or epiphallus, very near to the 
epiphallus-penis junction. The penis lacks a 
sheath, and internally, has a basic pattern of 
apical verge, circular ridges and a pocket 
stimulator. Among Australian subtropical taxa 
verges are rarely present, penial collars are con- 
fined to a handful of genera and a true pocket 
stimulator is seen only in Nautiliropa omicron 
(Fig. 65c). Instead there is a simple, apical (oc- 
casionally subapical) entrance of the epiphallus 
and few to many longitudinal pilasters which 
may be modified under conditions of sympatry. 

The penes of Hedleyoconcha and Ngairea 
have an apical muscular collar (Figs 4b, 10b, 
13b, 16b, 24c, 27b) which is comparable with 
that of the New Zealand Phenacohelix pilula 
(Reeve, 1852) (Solem, 1983, p. 28). Addition- 
ally the large fleshy penial pilasters of Cralopa 
are similar to those of the New Zealand Cavellia 
sterkiana (Suter, 1891) and C. colensoi (Suter, 
1891)(Climo, 1969b). 

According to Climo (1980, 1983) Pulch- 
ridomus^Aeschrodomus, Charopa and Phenaco- 
charopa have a constricting 'sphincter* dividing 
the penis into distinctly sculptured upper and 
lower chambers. A similar pattern is seen in 
Elsothera sericatula and E. genithecata (Figs 
105e, 109b). 

Most Australian, New Zealand and Pacific 
Island species have a well defined epiphallus. 
However, some New Zealand species lack an 
externally differentiated epiphallus. A com- 
parable Australian species is Setomedea janae in 
which the epiphallus is partially incorporated 
into the penial complex (Fig. 39c). 


The typical charopid radula has tricuspid 
central and lateral teeth (usually the central tooth 
is slightly smaller) with long lanceolate 



mesocone and short pointed ectoeones. The 
marginal teeth arc multicuspid and in many in- 
stances a transitional zone of lateromarginal 
teeth is present. This pattern is the rule in Pacific 
Island and most Australian subtropical species 
Notable local exceptions nre ligifomen globosa, 
£. lochl/dinm (Figs 128h, 13 1 h). Omphaloropa 
vancosa and Lctvmoh conl&rtus (Figs 134h. 
68c) which have reduced central teeth, This 
latter condition is common in New Zealand 
species I Chrno, 1969b, 19701 

The broadly trianmilar mesocone and bicuspid 
laterals of Ngairea (Figs 3f, 9d. I2d, 15d) are 
similar to those of I he New Zealand 
PhenacoheiLx (Cumber, t%l). Significantly 
belli are arboreal to semi-arboreal However, 
presence of anlerior flares -r teeth in groups 
such OS Ngairea and Hedley&COficha has no 
known correlative in either New Zealand or 
Pacific Island taxa 

Nuut'ihr\>pa omicron has slender bonk-like 
teeth and V-shaped tOW arrangement (Figs 
63c,fji This is a majur departure I mm other 
subtropical species and also has no equivalent 
among Pacific Island or New Zealand 
Ounipidae. rV. omicrof} \& sympatne with the 
conehologically convergent Gyracochlea and 
the unusual raduln may be a local feature related 
fO intergeneric sympatry. 

In summary. Australian subtropical 
( Hiropidac display a number of conchological 
and anatomical features which suggest relation- 
ships with New Zealand eharopids but also indi- 
cate evolution in isolation for some considerable 
period, At the same lime development under 
similar environmental conditions has led lo 
many convergences in shell features am 
species of these Iwa regional charoprd faunas 
Analysis of southern Australian species will be 
critical in understanding the fulJ extent of pos- 
sible relationships. 


The fifty species o! Charopidae reviewed dis- 
play a wide variety of anatomical and con- 
chological patterns, yet ihev represent only a 
small subset of an extensive Australian cbaropid 
FaUnflj The prolific Tasmanian and Victorian 
faunas, gxusnsfve radiations in mideast and 
northeast Queensland, and a further 60-70 poor- 
ly known or undescribed subtropical species are 
still to be reviewed Hence the following discus- 
sion is preliminary. 

Ngairea, iledleyoeoneha. Mussomtla, Lvn~ 

webbia and Setomedea have incomplete secon- 
dary ureters and possess a number of con- 
en* l.'g'ial specialisations (incised spiral lines, 
broad spiral grooves, periosteal setae, punctate 
apical sculpture) which isolate them from the 
genera with the more typical charopid pallia! 
configuration in which the secondary ureter is 
complete. They probably had a common ances- 
tor. Their penes have a typical internal pattern 
of apical collar and longitudinal pilasters. This 
is well illustrated in Ngairea and Hed- 
leyoeoncha. Together with Mussonufa and/.?rc- 
wehhia these genera are further characterised by 
elevated shells with very narrow umbilici. An 
exception is A', murphy* which has secondarily 
narrowed whorls and a slightly more open um- 
bilicus. Lenwebbia can be derived from Ngairea 
through modification of shell sculpture, penis 
and kidney and shares the unusual feature of 
incised spiral lines withJVjgoirca and Mussonttla. 

S&omedea appears to represent an attempt at 
habitat specialisation among the taxa with in- 
complete secondary ureters. The relatively small 
size, peculiar coiling of the primary ureter, and 
interspecific variability in whorl numbers and 
shell coiling pattern, arc inconsistent with other 
genera in thi> group. Unlike Ngairea and Hed- 
leyoconeha fand probably Mussonula), 
SetOtnedec is not arboreal or semi-arboreal lr 
usually lives in moist rotting logs and the 
pcriostracal setae are probably an adaptation for 
preventing dirt accumulation on the shell in this 
comparatively wet mtcrohabilat. 

Among the more generalised radially-ribbed 
groups. Cralopa and FAsotliera stand apart in 
having altered penial and epiphallic complexes, 
closed umbilici and conspicuous apertural 
sinuso. This latter feature is relatively rare in 
Australian charopids and suggests a possible 
relationship with New Zealand particularly 
through CaveUia Ircdale, 1915. Cralopa and 
Etsothera arc probably more closely related to 
each other than to other groups reviewed here. 

In spile of si?e differences, Biomphohpa and 
GyrocochLu show structural consistencies in 
shell and anatomy which suggest common an- 
cestry. C) roctK.hlea is an evolutionary experi- 
ment in whorl size increase which reaches its 
extreme in G Kinuinciu. 

Nautdiropa with pitted apical sculpture, 
depressed spire, and strong radial ribbing may be 
related to the GvroeiH hlea-Btotnphalopa com- 
plex even though radular morphology, penial 
structure, shell coiling pattern and microsculp- 
lure are against close relationship, 



The highly modified Letomola contortus is 
possibly a derivative of the Gyrocochlea-\z\z\ of 
specialisation which has undergone size and 
shell sculpture reduction associated with a shift 
in microhabitat to life on rock surfaces. En- 
larged, bulbous protoconch and elongate 
genitalia indicate descent from a large ancestor 
whose organ systems are consistent with the 
elongate Gyrocochlea pattern. Barrier forma- 
tion and sulcus development in L. contortus can 
be correlated with change to small size and al- 
tered lifestyle. The bilobed kidney (with acutely 
angled ureteric arms) is probably a secondary 
modification directly attributable to reduction in 
whorl count. 

Rhophodon also has apertural lamellae but 
these are independently derived from those of 
Letomola. The coiling pattern, whorl profile and 
strong radial sculpture suggest affinities with 
Gyrocochlea lineage. However the apical sculp- 
ture does not. 

Coenocharopa is diverse in subtropical rain- 
forests. The various species lead relatively 
specialised lifestyles among the dirt particles of 
the litter, on rock surfaces, or among moss on 
logs. Their shells have undergone reduction in 
primary calcified sculpture and developed fine 
deciduous periostracal ribs and extensions. 
Radial apical sculpture, enlarged penial com- 
plexes and expanded epiphallic pore suggest dis- 
tant relationship with the CralopalElsothera 

Problems associated with Discocharopa have 
been discussed in the systematic section. In 
essence the subtropical Discocharopa may be a 
a locally derived group. Apical sculpture indi- 
cates possible relationships to the Cralopa-El- 
sothera group. 

Egilomen is represented by the depressed, 
widely umbilicate and strongly ribbed E. 
cochlidium and the globose, almost imperforate, 
more heavily ribbed E. globosa. The bilobed 
kidney (without evidence of whorl number 
reduction) suggests a quite separate derivation to 
other radially-ribbed genera reviewed herein. 

Omphaloropa varicosa with spirally Urate 
apex, wide umbilicus and large tubular penial 
verge does not appear to have any close 
equivalent among other subtropical species. 
Another species with spirally lirate protoconch, 
Sinployea intensa, represents a remnant of a 
more widespread Pacific group. 

Epiphallic and penial modifications distin- 
guish Rotacharopa while the peculiar apical 

sculpture relates the genus to the Pacific Island 
Microcharopa and North American Radiodiscus. 
Knowledge of Australian Charopidae is still at 
a rudimentary level and reflected by the mosaic 
nature of morphological trends outlined above. 
Production of a formal generic phytogeny would 
be a futile exercise at this stage. Rather I have 
proposed a scheme of hypothesised relationships 
(Fig. 149) which summarises the preceding dis- 
cussion and information given elsewhere in this 
study. It identifies the probable multiple origins 
of the fauna and outlines major trends discussed 


A preliminary computer-based cladistic 
analysis using PAUP was carried out. Seventy- 
five characters (42 anatomical, 33 conchologi- 
cal) (Appendix) were chosen and ordered. The 
resultant cladogram (Fig. 150) shows that taxa 
with highly differentiated shells such as 
Coenocharopa and Omphaloropa are distin- 
guished from the more generalised groups such 
as Gyrocochlea and Rhophodon. However, 
there are also some unusual associations where 
a high degree of conchological convergence is 
involved, viz. Ngairea levicostata and Mus- 
sonula verax ; N. murphyi and Hedleyoconcha 
delta. Egilomen globosa is associated with the 
Rotadiscinae even though the latter has marked- 
ly different sculptural and anatomical features. 
Elsothera and Cralopa are associated with taxa 
lacking a complete secondary ureter and the 
conchologically specialised Setomedea is 
separated from other species lacking a complete 
secondary ureter. Gyrocochlea, Biomphalopa 
and Nautiliropa were grouped together, and the 
highly modified Letomola contortus was placed 
near these genera. 


The Australian charopid fauna possibly con- 
sists of 250-300 species but prior to the present 
study, which introduces 27 species, only 110 
species level taxa had been named. Hence, with 
probably less than half the available species 
described, and an even smaller number subjected 
to detailed analysis, it is premature to be confi- 
dent about understanding their biogeography. 
The treatment below provides a synoptic review 
of climatic, geologic and vegetational changes in 
the east coast region during the Tertiary; brief 
discussion of charopid distribution patterns; and 


synthesis placing the taxa of this study into 
broader biogeographic context. 


In subtropical and tropicnl eastern Australia 
distribution of land snails and rainforest arc 

linked In I be Chatoptdae, which is acknow- 
ledged to be a Gondwanan family (Van Bruggen, 
\9HQ) ihis relationship probably had its origins 
in Ihe Late Cretaceous when mesic forests were 
more widely spread than at present. Since the 
Cretaceous few Charopidae have made the Iran 
sition todnersclemphyll forest, in spite of potent 
environmental change which has favoured the 
development of \erophilic communities. This 
indicates a strong ecological bond between these 
tiny snails and closed forests w hich is prohahK 
due to convergent habitat preference, The 
ecological factors which favour development 
and persistence of rainforests e.g. moisture and 
euOMphic soils, are also those which favour the 
SUCCCS5 Of iCrTCSlrial molluscs For SnailS, rain- 
loresi alsr provides sheltei and food- These 
conclusions are not very different from those of 
Solem.Clinoand Roscoe(UJKi I who found that 
C and spatial quality were significant fac- 
tors in the survival of snail species in the 
ManuVau Peninsula area of New Zealand, and 
Box-. '34), who considered that shelter. 

lime and moisture svere key (actors in determin- 
ing local occurrences of British land molluscs. 

ta separated from Gondwanaland in the 

early I crtiary reaching its present position at the 

d .. Of ihe Asian plate in the mid-Miocene 

(Kcwji, 1981; Kemp and Galloway, 1981). In 

die liar I) leriiarv, most of Austiali'a would have 

resided within p weak band oi wind circulation 

L| id the surrounding warm seas would have 
resulted in widespread precipitation. But with 
opening of the southern ocean and drill 
northward into lower latitudes. significant chan- 
ges in Climate occurred. Circum-polai circula- 
tion significantly reduced heattransport between 

i equator and pole resulting in latitudinal 
Stratification, of watei temperatures. Conse- 
quently around the middle of the Miocene, ihe 
sea surface cooled and precipitation on land 
declined. From that lime aridity has been a 
rlnn , dor m Australia's cl i male with wet- 
ter phases occurring only briefly Rainloresi 
communities, widespread at the beginning of the 
Ternary, retreated to refugia and were replaced 
by a \cne adapted biota. However, in WiStCttl 

Australia, two geological events contributed to 
the maintenance of mesic communities in the 
wake of the a rid phases w I itch elsewhere resulted 
in the removal ol the widespread rumforcst flora 
(Martin. 1*378). Firstly, episodic uplift of the 
eastern highlands began in the Paleocene with 
major elevation occurring in the late Miocene 
and Pliocene Secondly there was widespread 
igneous activity ;n eastern Australia along and 
adjacent to the eastern highlands (Wellman and 
MaeOougall. 1974). In northern NSW viil- 
canifim continued into the mid-Miocene while in 
SEQ peak activity occurred in the Oligoccnc and 
Plio-rlcistooenCi The eastern highlands ensured 
orographic rainfall and enabled species to radiate 
a 1 1 U u d i n a I ly , while i n d i I e c 1 1 y . \ u I ea n i sm 
pio\ided favourable soil-water Conditions in a 
basically Jrj infertile confincm. Ihe fact thai 
most land snails in eastern Australia now occur 
m 'rainforest' indicates lhat Ihe persistence 
moist refugia was critical to Iheir survival in this 
region 'he major refuges during these Urv 
periods were in the moist uplands, and in 
loWlandS along major drainage: lines — areas 
which were not large, but Wet. 

Climatic changes in ihe late Quaternary were 
similar in amplitude to those experienced" in the 
Miocene and Pliocene but occurred in more 
rapid succession. Galloway and Kemp (1981) 
concluded that these changes must have placed 
considerable stress on montane and coastal en- 
vironments and that modern communities in 
these situations are recent phenomena consisting 
of biota which have survived in isolated refuses. 

Recent studies by Kershaw (1980, I9&I) on 
pollen samples from crater lakes on the Athcrton 
I ablcland. confirmed lhat the climatic fluctua- 
tions of the late Tertiary had dramatic effects on 
the Australian vegetation. Kershaw's icsults 
showed that between 83,500 and ftOQG BP, 
complex rainforest covered the Athcrton 
Tableland underrainfall much bighei than rodav 
These conditions persisted until about 38,000 BP 
ii a majoi change occurred and wei adapled 
araucarian forests of the Athcrton Tableland 
acre replaced by drier communities charac- 
terised by ElicafyptUS and Casuaritia. These 
dominated Ihe are* until 8-10,000 BP when rain- 
-.' again emerged from retreats. Kershaw 
| 1 980, 198 I ) Was further able to demi mSft&HC that 
these local, environmental changes could be ap- 
plied to other areas along the east coast. 

The penological studies of Kershaw (1481) 

*y Hypothesised relationships o| charopid genera re- leased in lift* study 




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and Martin (!P81) demonsiruied the. drastic ef 

feCtS *if mid to late Tertiary climate on mesic 
communities and exposed some important prin- 
ciples penmen? lo east coast biogeography. 
Firstly, areas and their faunas have been isolated 
an •> number Of occasions thus promoting al- 
lopatric specialion> and subsequently unified lu 
allow for dispersal and the evolution of sym- 
patic congeneric pairs. Secondly, certain ele- 
ments of the biota found in the "modern" 
northern Australia have been present in Australia 
for a long time and many ol these taxa presently 
confined lu the north were present in south-east- 
ern Australia When it was adjacent to Antarctica. 
Thirdly, some species in northern Australia 
have becomu extinct in fairly recent times. 
These principles have particular application to a 
low vagility Gondwanan group of organisms 
such as the charopid land snails. 

Webb (1950, l%8) expressed the results ol 
past environmental changes through a present 
Clay classification ol structurally based rainforest 
types. In a biogcographic synthesis, Webb and 
fracey (1V81) demonstrated that the popular 
concept of tropical and subtropical rainforest 
encompasses a complex mosaic o( community 
types each reflecting a past scries of climatic 
change. This division of the "rainforest massif 
into regions and provinces associated with 
present climatic regimes, which t*l the same time 
reflected quite different histories of climatic 
change, provides a practical framework for inter- 
preting the impact of these changes in the context 
ol the fragmentation and isolation i_»f rncsic com- 
munities. At a regional level the results indicate 
that the cooler forests of the southeast (cool 
temperate to subtropical) have been separated 
from the warmer forests of the northeast (tropi- 
cal), probably since the late Miocene arid IT) 
phases. A dry corridor between Gladstone and 
St Lawrence, geographically distinguished by a 
lack of coastal and subcoastal highlands, ensured 
that the biotas of the two regions developed in 
[solAlioil while a dry corridor between 
Townsvillc and Bow en isolated the Eungella 
rainforests from the main massif of northern 
rainforest. At a provincial level, temperate 
sts of the cooler Tasmanian and Victorian 
regions arc segregated from the more northern 
warm \ernpemte, and CQDl to warm subtropical 
forest while a third drier subtropical forest type 
occurs subcoastally in southern Queensland. 

Considering the close ecological ties between 

Clmi-'fidac and closed ,,.k*--!s, Webb's rain- 
forest classification makes it possible to relate 
past climatic and geological events to present 
day charopid distribution. 

Charopid Distribution 

At the family level distribution of Charopidae 
in Australia is basically east coast. Species are 
found in moist cool temperate forest of the south- 
east region (Smith and Kershaw, 197°; Smith 
and Kershaw, 1985), warm temperate and sub- 
tropical forests of northern NSW and SEQ, in the 
structurally subtropical forests near Mackay. 
MliQ, and the tropical coastal forests between 
Ingham and L'ooktown. NEQ. A small but Sig- 
nificant radiation of eharopids is also present in 
souiluvesl Australia (Solum, 1983), several 
species exist across the north ol the comment; 
f Salem i (984), and a presumably relict species 
Piiionte.'ia aemula (Tale, 1 894), is found in the 
rernnanl rnesophylic fores! of Central Austral a 
Alt indications are that the distribution of the 
group in Australia was much more extensive in 
the past. 

Of the 50 species reviewed herein 32 are en- 
demic to the main feres ol subtropical reforest 
between Dorngo. NSW (30'30'S) and Rock- 
bamptnn MEQ (23*30'S). Three species \ iz. 
Hedlt\\ocottcha delta, Coenocharopa tnuUh 
radtatu and CraU>pa stroudensis extend further 
Lh, while 14 species occur outside this core 

Discpckaropa npcrta from the drier rain- 
forests of SEQ ranges to northern and central 
Australia, the Bismarcks, Philippines, Indonesia, 
New Hebrides, Kcrmadcc. Fiji. Samoa, Austral 
and Society Islands (Snlem, 1983, pp 74-81) 
The small, scmiarborcal Sinploveu iniensa has a 
long, narrow, coastal distribution which encom- 
passes a number of sublittoral rainforest patches. 
Wide ranging species also include the litter 
dwelling Omphaloropa nuicosa which is found 
from the McPherson Ranges on the NSW/Old 
border, to west of Rockhampton, MEQ: 
Setomedea st'ticostata and Ngairea dorrigoensis 
which range from Dorrigo, NSW. to just north of 
Brisbane, SEQ, and the litter dwelling 
Coewcharopa patricostata with a range from 
Dorrigo. NSW. to just north of Gympic. SEQ. 
The north-south linear ranges of these species 
arc between 350—700 km and are the upper ex- 
tremes where the majority of species have ranges 
of less than 200km. However in general ihe 

FIG 150. Consensus irec derived from PAUP analysis. 



















pro toscrobicu lata 

































- I 

MPMniRso-niFourrNsi and minfum 

raiiges da rtoi approach Sotetn s predicted vultw 

lor all land snails of less than SO km (Solem, 

With the exception nl Rhophodon roimatu, 
Whose distribution is considered to be poorly 
known, the smallest linear ranges of subtropical 
endemic species are those of Rhophodon 
aliz#h£th&fi afld Setomedea nudicostata which 
N ur in the relatively small and unusual moist 
subtropical refugium in the Bobby Range, south- 
west of Gladstone, SEO. These examples sug- 
gest that species ranges may be determined by 

the size of parrtcula ! Test patches and 

hence subject to considerable pressure from 
changing climatic conditions. Cocnochnrupn 
idbaftensis, Rhophodon kempseyensis and 
Offfold ro.itonus fnim the Vcssahah-Carrai 
limestone scries. we>i of Kempsey, NSW, are 
further examples of environmentally restricted 

Coenocharopu lias four endemic, often over- 
lapping species in subtropical forests with com- 
paratively large linear ranges (180M86 km) 
indicating that the genus has undergone a long 
period of environmental sifting. In 
(rvrocochleu, linear ranges are relati\elv short 
(54-108 km) and patterns of overlap arc minimal 
w ith only Gxrucoehleu \initutaa and G. 
noAirtla-m:ilata showing minor sympatic occur- 
rence. It ts probable thai the GyrWQcitlw radia- 
tion is a relatively recent phenomenon in the 
subtropical region. 

Geographic distribution within a genus is 
generally north-south. Rhophodon* with a single 
speci.-s in !he southeast region arid sh species 
from Kempsey. NSW, lo the Bobby Range 
SEO. <s illustrative. Setontedea and Hed- 
ItyocoflCfia have more striking disjunct rtortb- 
Kuuth distributions with relict species in central 
and north Queensland. Ngairea has its main 
radiation in the subtropical forests but also has 
an outlier species in the I Ha warm region ol 
southern NSW. The southern limitsof Llsoihent 
arc not known but its occurrence in the subtropi- 
cal forest is also considered to be relict. As a 
rule, genera lend to have relicts in the north 
rather than vice versa indicating that some 
genera which are present in the south were also 

I ttflW ^ the north during cooler times, 

A major exception to the typical north-south 
pattern IsCraiopa which has a single widespread 
species. C simtnlcnsis, in the coastal and sub- 
Coastal SUblfOpfcal rainforests; a closely related 
SpCClCs;, C. kapituucttsts. on Mt Kaputar, west of 
NarrabrL NSW; and a further widespread, but 

geographically disjunct species. C cariessi, on 
the limestone outcrops of the Great Dividing 
Range. Unlike other genera now confined to the 
Cast, Cralnpa has been able to survive in largely 
snail depauperate areas further west, 

Although patterns of geographic distribution 
are usually north-south, large wet areas such as 
the MePherson Ranges provide Sufficient width 
in the rainforest massif to allow for localised 
speciation in an east-west direction, e.g. Ngairea 
and Gyracitchlca. 

There is limited evidence for altiludinai dif- 
ferentiation. E#ilomen each lid i urn and 
CoenDcharopQ alata show preference for mon- 
tane habitats in contrast to the foothill/lowland 
distrihulion of their congeners. Ngairea cot- 
uricoJti is microsympatrie wilh A', tlorngornsts 
in the Great Dividing Range, and extends to the 
foothills whereas A 1 , ddrrigdensls occurs only In 
the highlands. Other examples are seen in 
G\rocochlea and Rhophcdon. 

Coherent distribution patterns among closely 
related genera are less obvious but this is probab- 
ly due io gaps in phvletie knowledge. Where 
patterns emerge they appear to be similar to 
those discussed above GyroOichha which is an 
endemic subtropical group has its nearest rela- 
tive, Hiomphutopa, in the rainforests of central 
and north Queensland, In conimsi die closely 
related MtiSSOftufat lenwvbbia and Ngairea 
have overlapping distributions in SEQ. 

Pathways of DiFrFRFNTiATtoN 

I consider that the environmental changes as- 
sociated ft ith mid It) late Tertiary aridity were the 
major factors affecting the present makeup and 
distribution of the Australian Charopidae. Two 
levels of changes arc seen as particularly sag* 
RtfiGltU- those which produced generic differen- 
tiation and majur mtrageneric disjunctions, and 
those which produced fine grain distribution ad- 
justments within larger areas of rainforest. The 
first level probably dates from the onset of aridity 
in the Miocene, while the second is most likely 
rotated tO drastic short term climatic oscillations 
of the Quaternary. 

Id the north the formation ol dry corridors, 
between St Lawrence and Roekhampton, and 
Howe n and low nsville(Fig 1 j correlate wnh the 
current disjunct distributions o\' Hcdleyocondta 
and Setomcdca. Persumablv these groups once 
ranged along the entire moist coastal region 
from north Queensland to northern NSW. As 
mesie habitats retreated, the ancestral range be- 
came severely fragmemc-d, and mid-east 


Queensland and northeast Queensland popula- 
tions would have been isolated from each other 
iiiu.l from mwe B©UihCffl subtropical raniloresis 

vrhnreal Hedlt'voconcfw responded more 
dramatically disappearing from the notophyll 
vine forests of mid-east Queensland. In the 
north. //. aitaketoae survived only in the al- 

(udinil relu.eja of Ml. Bellenden Ker, while in 

I »Otttb, persistence of suitable habitat enabled 
H. delta to flourish in a wide area of moist 
subtropical rainforest in northern New South 
Wales and southern Queensland. 

Sctumcdea would have endured similar en- 
vironmental pressures to fledleyoconcha but. 
prObAbly because of greatei adaptability Or 
peculiar microhabitat requirements, managed to 
maintain S. }atiac in mid-east Queensland, and 
at least two other spc< ,! 1 Ifl north (Jurrnsland. 
In the north S&OtHeaea was most likely restricted 
to montane refugia at some lime in the drier past, 
hut subsequently lias been able to disperse into 
the foothills and lowlands (5. montcithi). In 
&0Uth6rn Queensland and northern New South 
Wales, S. scticpstuta (Medley, 1924) occupies 
upland rcgufia as well as lowland notophyll vine 
forests also indicating dispersal from high to 
altitudes in wetter times. Further examples 
of large-scale tntragenenc disjunctions which 
can be related to mesic habitat regression include 
the separation of BiompltalopQ concinnQ and/1 
n;ctJ\a, and ihc isolation ol Ngtlir&l murpftyt in 
Ihe lllawarra region, southern New South Wales, 
from the main mass ottfgaircn specks fUlthei 

C rcation o| these major dry corridors probably 
also led to generic differentiation between north 
and south I his is best illustrated by the disjunct 
distribution of the closely related Gyr0C0$hlca 
JtKeaSt Queensland) and Biotnphalopa (mid- 
! and north Queensland). 
In southeast Queensland, regression of com- 
plex mesie habitats to upland refugia and their 
replacemenl by less complex miciophyll vine 
toresls in Ihe lowlands between dvmpic and 
Rockhnmpton, correlate with disjunctions in 
several groups. Rhophodon clizabethae, Gyro- 

Mtffl curlisKtnu. St'tomeden nudtcostata and 
lirea caHahcufata occur within this region in 
wetter moniane refugia and are geographically 
isolated from congeners further south. This 
isolation has been severe in/?, elizuhediac and5. 
umltcosttitti which arc restricted to relugial 
notophyll viae forests in the Bobby Range. SW 

"I Miriam Vulc and S cmplicuhuit which io- 
halnts ofll) the moial forests of the mountain 

summits between ( 'oiosstum Creek and Mt Lar- 
com. In conlrast U. curtisitma is (ound in both 
moist and dry toTCSW II probably survived 
climatic attrition in scattered refugia and sub- 
sequently dispersed into the drier microphvlt 
vine tort-sts and (ringing open lores! In com- 
parison with R. elizahetiiae. S. Hudteosiata and 
N. canahrulala. ihe larger G curn\:ana has less 
specific microhabitat requirements. It has more 
generalised pallia! structures indicating that it 
may have had less difficulty in adapting to drier 
environments. The fragmented distribution of 
Rotacharopu is also probably due fo these loeuil 
environmental chang 

Although ;Yl;./i <: u- has not colonised these di ie| 
microphvll vine forests, Ixnwchhia protoscro- 
bictdtiia, which is considered to be a derivative 
Of the \y.inr,'Lt group, has done so successfully 

Within the main humid subtropical rainforest 
block, which includes the Border Ranges. Great 
Dividing Range (from Dorrigo, NSW to the Mis- 
take Mountains, SEQ) and their foothills and 
lowlands, ehaiopiu 1 evolution has been much 
more complex The long-lrtm stability of ihcse 
subtropical forests was ensured by the 
favourable volcanic soils ami above average 
rainfall, even at lower altitudes. Hence while 
ihe.e was fragmentation and retreat of moist 
forest into favourable refugia such as gully 

tveflds, mvnne alluvii ;i "d mountain sutnroii 

dry periods, alternate periods of higher rainfall 
would have led to many episodes o? cxpan 
and amalgamation, hi some areas such as t>ic 
drier western edges of the Border Ranges and 
Great Di\ idfr, and |ripIaCO$Whefe$QJ] typCS 
poor, periods ol lower rainfall did cause a - 
in vegetation type to drier miciophyll vine forest. 
In higher reaches e.vv l.ammglon PlaU 

'temper it ra in fore si pi r&fatcu in seit cN d 

refugia. These events would have proyidedcon- 
siderable opportunity for local speciation. dis- 
persal and isolation. The results have been 
'fine-grain' speciation patterns and evolution or 
several endemic genera. 

The ancestral siock o( Gymcnchlea presumab- 
ly hail a distribution which covered the upl. 
and lowlands of the eastern Border Ranges. I D - 
lowing climate-induced liabila! fragmentation, 
the heavily ribbed Gyrocochlea vinitinchi wouJd 
have been isolated in higher pans of the 
Lamington Plateau from ihe sparsely ribbed 
forms of the foothills and with ihe shift CO drier 
forest type* in western areas, 6 convohua be- 
came 51 r Mated from more castcrh 
paw S in the. palm- 



dominated forests around the Darlington Range. 
In wetter times, G. vinitincta dispersed into the 
range of G. paucilamellaia. Similarly evolution 
of local Rhophodon probably involved frag- 
mentation of a wide ranging ancestral population 
into foothill (R. consohrinus) and upland (R. 
peregrinus) species. Subsequently R. pcregrinus 
expanded its range to become sympatric with R. 

Aridity episodes probably isolated the weakly 
ribbed Ngairea corticicola in the subtropical 
notophyll vine forests of the uplands and 
foothills from the heavily ribbed stock in more 
restricted temperate montane refugia. In the 
western highlands of the Border Ranges the 
development of drier vine forests effectively iso- 
lated N. levicostata from the widespread mon- 
tane species, N. dorrigoensis. As aridity phases 
were replaced by periods of higher rainfall, N. 
dorrigoensis was able to disperse from montane 
refugia and gradually occupy the ranges of its 
two congeners. 

It is more than coincidental that the expansion 
of rainforest during wetter periods favoured those 
species restricted to moist montane refugia. 
These communities were able to expand into 
lower areas where the soil type was suitable, and 
into areas with less suitable soils where rainfall 
was adequate. On the other hand the lowland and 
foothill rainforest communities isolated in gully 
heads would have expanded to produce a more 
contiguous block, but would not have expanded 
to higher altitudes. In contrast drier rainforests 
were environmentally cornered, and consequent- 
ly, the charopids in them, limited in their dispersal 
e.g. N. levicostata and (7. convoluta. 

The ability to relate past events of climatic, 
geological and vegetations! changes to present- 
day distribution patterns in the Charopidae 
varies from group to group. In Coenocharopa. 
which contains six species with widely differing 
microhabitat preferences, these events have 
produced a very complex pattern of species over- 
lap. Indications are that the group has undergone 
a long period of environmental sifting that has 
blurred historical pathways. There is evidence 
for segregation o[ a mesic-adapled highland 
species (C. alata) and a species which became 
adapted to the more easterly warm moist 
notophyll vine forests of the foothills of the 
McPhcrSon Ranges (C. parvicostaia). In the 
north. C. macromphala and C. sordidits probably 
developed from a common ancestor in response 
to environmental change which isolated mesic 
communities in the D'Aguilar Range from the 

araucarian forests of the Blackbutt Range and 
Mt. Mudlo region. Most likely more than one 
instance of dispersal has brought about the com- 
plex sympatric patterns evident today. 

Nauiiliropa and Mussonula are endemic to this 
region and are probably recent local derivations 
of more generalised stock, viz. Gyrocochlea and 
Ngairea respectively. 

Restriction of charopids to wetter coastal 
habitats during the latter half of the Tertiary did 
not necessarily mean complete extinction of the 
family from drier western, sclerophyll- 
dominated areas. Where conditions were 
favourable i.e. where moist refugia could exist, 
charopids persisted e.g. Cralopa. As mesic 
habitats retreated eastward in the face of increas- 
ing aridity. C. kapuiarensis was isolated in the 
moist montane refugia of Mt Kaputar, near Nar- 
rabri, NSW, and a little further east, C. carlessi 
survived in scattered limestone refugia on the 
western edge of the Great Divide, albeit reduced 
to a series of cornered disjunct populations. 

Limestone habitats are extremely important 
refuge zones for land snails. Along the Great 
Divide, they typically support remnants of once 
more prevalent wet-adapted vegetation. The 
rocks trap available moisture in crevices and also 
provide protection from wildfires thus enabling 
small island communities, quite different from 
those in the surrounding open forest, to be main- 
tained. Land snails which inhabit these excep- 
tional refugia benefit from an ample supply of 
calcium. Because of long term isolation and 
need to adapt to a lifestyle among and on rocks, 
endemicity and specialization among snails is 
usually high. Limestones of the Yessabah-Car- 
rai block, west of Kempsey, are a case in point. 
Their isolation from the main subtropical rain- 
forest block, presumably due to mesic habitat 
regression, has resulted in separation of several 
charopid species from northern congeners e.g. 
Rhophodon kern psc\ easts and Coenocharopa 
yessahahensis. The monotypic Letomola^ which 
may be a derivative oi' the Gyrocochlea lineage 
of subtropical charopids, is also endemic to these 
limestones. All these species have highly dif- 
ferentiated shells and habitat-related radular 


Origins of the eastern Australian subtropical 
charopid fauna appear to be diverse. Groups 
with primitive pallial configurations such as 
Ngairea, Hedlevoconcha and Setomedea are 


probably remnants oi ''older stock with origins in 
the widespread Gonchvanan biota. This group 
seems to have suffered extensively from mesic 
habitat regression alihough Ngairea has 
managed to secondarily diversify in the subtropi- 
cal forests, \fu.\snnula and Fenwcbbia are rnosl 
likely recent, local derivatives. Cralopa and El- 
sothera have shell and anatomical features relat- 
ing them to New Zealand Charopidac and arc 
probable descendants of southern based groups 
thai have attenuated distributions in subtropical 
regions. Coenocharopa can be considered an 
early oIKhoot from this stem that radiated into 
diverse habitats in the subtropical region 
Rhophodon, with widely separated species along 
the east coast is probablv of southern origin 
representing an early attempt at habitat 
specialisation and like Coenocharopa and 
Nguirea has secondarily diversified in the sub- 
tropical forests. Gyrocorhlco, Btomphalopa, 
Nautiliropa and Letomola arc possibly part of 
the same lineage nsKhop/mdon, but more recent 
in origin. In the case of Letomola and 
Sautiliropa generic differentiation has been ae- 
L-nmpanicd by dramatic changes in conchologi- 
Cal and anatomical features. 

Of other genera, Sinployea is a remnant of a 
larger Pacific Island radiation, Egiiomen is of 
uncertain affinity but has anatomical features 
indicating only distant relationship with other 
subtropical groups; Omphaloropa has a conflict- 
ing combination of characters and unusual dis- 
tribution pattern which suggest that its 
relationships may be with species well to the 
north or south; Discocharopa remains 
problematic; and Rotachampa is related |o 
Pacific Basin and north American species. 


I owe special thanks to Mr Darryl Potter, 
Queensland Museum, foi his assistance in many 
aspects of this project. I am most grateful to the 
late Dr Alan Solem, Field Museum of Natural 
History, Chicago, USA, for his critical com- 
ments and constant encouragement. 

For the loan of material I thank staff of the 
Australian Museum, Sydney; Museum of Vic- 
toria, Melbourne, South Australian Museum, 
Adelaide; and British Museum of Natural His- 
tory, London. Other material was generously 
provided by Graeme Annabel!, Vince Kessner, 
and Terry Carless. 

Numerous specimens used in this study were 
collected during field work funded by the 

Australian Biological Resources Survey (1981- 


I wish to thank the following staff of the 
Queensland Museum • Dr Robert Raven (An 
nulogy) frti his help with the SEM and computer 
analysis; Dr Geoff Monteith (Entomology) for 
providing material and important information on 
collecting sites; Ms Gudrun Sai nes (Technician, 
Queensland Museum) foi help with the SEM: Mr 
Bruce Cowell and Mr Gary Cranitch (Photog- 
raphy) fot their assistance with all aspects of 
photography. In the early Stages of the project 
Miss Alison Green, formerly Technician 
Queensland Museum assisted \\\\\\ SEM arsd 
field work. 

For typing the manuscript. I am particularly 
grateful to Mrs Peta Woodgale, Queens! 

This work is based on a thesis which was 
submitted in fulfilment of the requirements for 
the degree of Doctor of Philosophy at the 
University of Queensland (i&88) and I would 
like to thank Professor Barrie larnieson. Depart- 
ment of Zoology, for his guidance, and criticisms. 

Finally. I wish to thank a special friend, J mi 
Alio has been most supportive during the course 
of this project. 


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1 .Tail differentiation (absent, 0; present,!) 

2. Foot shape (short broad, 0; bluntly rounded, 1; 

slender tapered, 2) 
3. Kidney shape (bilobed,0; moderately bilobed,l; 

weakly bilobed,2; almost unilobed,3) 
4. Configuration of kidney apex(simple,0;reflexed,l) 
5. Primary ureter configuration (simple.0; reflexed,l; 

coiled, 2) 
6. Length of secondary ureter (incompleted; com- 
plete, 1) 
7. Number of lobes in ovotestis (one,0; two,l; 

8. Ovotestis orientation (parallel, 0; right angles,l) 
9. Ovotestis shape (palmately clavate,0; finger-like,l; 

bilobed,2; unilobed,3) 
lO.Length of talon stalk (long,0; medium, 1; very 

ll.Talon shape (circular,0; finger-like, 1) 
12. Hermaphroditic duct kinking (unikinked,0; 

kinked, 1) 
13.Vas deferens shape (thin,0; weakly expanded,!; 

strongly expanded, 2) 
14. Nature of epiphallus - vas deferens junction 

(simple,0; complex,!) 
15. Differentiation of epiphallus (strong,0; weak,l) 
16. Presence of epiphallus (present,0; absent,l) 
17. Presence of epiphallic sheath (absent,0;present,l) 
18. Epiphallus length (short,0; elongate, 1) 
19. Epiphallic pilaster pattern (longitudinal, 0; 

20. Epiphallus configuration (ascending arm longer,0; 

ascending/descending arms equal, 1) 
21. Nature of epiphallus - penis junction (simple,0; 

complex, 1) 
22. Position of epiphallus - penis junction (apical,0; 

23. Nature of epiphallic entry into penis (through 

pore,0; through verge, 1) 
24. Position of penial retractor muscle insertion 
(penis,0; penis/epiphallus junction,!; epiphallus 
near penis,2; epiphallus far from penis,3) 
25. Presence of epiphallus - penis binding (absent,0; 

present, 1) 
26. Presence of penis sheath (absent,0; present, 1) 
27. Penis shape (pear shaped, 1; tubular,l) 
28. Penis expansion (slender, 0; moderately ex- 
panded,!; strongly expanded, 2) 
29. Presence of penial verge (absent,0; present, 1) 
30. Penial pilaster pattern (longitudinal,0; modified,!) 



31. Presence of internal penial collar (absent,0, 

32. Shape of genitalia (expanded.O; elongate, 1) 
33. Length of atrium (short,0; moderately long.l; very 

34. Length of vagina (short,0; moderately long,l; very 

35.Presence of preputial tube (absent,!); present,!) 
36. Presence of penial stimulatory pad (absent.O; 

present, 1) 

'.Presence of flared extension on radular teeth (ab- 
sent^; present,l) 
38. Configuration of radular row (transverse,0; v- 

39.Nature of central tooth (strongly tricuspid,0; weak- 
ly tricuspid, 1) 
40. Nature of lateral teeth (tricuspid^, bicuspid,!) 
41.Presence of lateromarginals (present.O; absent, 1) 
42. Nature of marginal teeth (multicuspid with 

variable-sized cusps.O; other, 1) 
43. Shell coiling pattern (loose,0; normal,!; tight,2) 
44. Nature of sutures (impressed,!); strongly im- 
pressed, 1 ; channelled,2) 
45. Presence of apertural barriers (absent.O; present, 1) 
46. Presence oi supraperipheral sulcus (absent.O; 

present,! ) 
47.Shape of umbilicus (closed,0;narrow-u.!;narrow- 

v,2; wide-v,3; cup-shaped,4; saucer shaped,5) 
48. Whorl profile (keeled.O; angulate.l; flattened 

above and below.2; rounded, 2; rounded below.4; 

laterally compressed,5) 
4s\Number of whorls (3.0-3.5.0: 3.5-4.0,1; 4.0- 

4.5,2; 4.5-5.0.3: 5.0-5.5,4: 5.5-6.0,5: 6.0-6.5,6) 
SO.Height of shell (0-1 mm,0, 1-2 mm,l; 2-3mm.2: 

3-4 mm.3; 4-5 mm,4; 5-6 mm, 5; 6-7 mm T 6) 
51. Shell diameter (1-2 mm,0; 2-3 mm, 1; 3-A mm,2; 

4-5 mm.3; 5-6 mtn.4; 6-7 mm,5: 7-8 mm,6) 
52.H/D ratio (0.40-0.50,0; 0.5 1-0.60,1, 0.61-0.70,2; 

0.71-0.80,3; 0.81-0.90,4) 
53. Spire protrusion (depressed,0; 0.01-0.20 mm,l; 

0.21-0.50 mm,2; 0.50-1.00 mm,3; 1.00-2.00 

mm. 4) 
54.Bodv whorl width (0.01-0.50 mm,0; 0.51-1.00 

mm.l; 1.01-1.50 mm.2: 1.51-2.00 mm,3; 2.01- 

2.50 mm,4; 2.51-3.00 mm,5; 3.01-3.50,6) 
55.SP/BWW ratio (0.00-0,10.0; 0.1 1-0.20,1; 0.21- 

0.30,2; 0.31-0.40,3; 0.41-0.50,4, 0.51-0.60.5; 

56. Number of ribs on body whorl (25-50.0: 51-75. 1 : 

76-100,2; 101-125,3; 126-150,4: 151-175.5; 

57.Ribs/mm (0.01-5.00,0; 5.01-10.00,1; 10.01- 

15.00,2; 15.01-20.00,3: 20.01-25.00,4; 25.01- 

30.00,5; 30.01-35.00,6) 
5S.UmbilicaI width (closed.0; 0.01-0.50,1: 0.51- 

1.00.2: 1.01-1.50,3: 1.51-2.00.4) 
5P.D/U ratio (0.01-2.50,0; 2.5 1-5.00, 1 ; 5.5 1-7.50,2; 

7.51-10.00,3; 10.00-closed,4) 
60. Nature of apical sculpture (spiral.O; predominantly 

radial,!; other,2) 

61 Presence of radialson terminal part of protoconch 
(absentO: present, 1) 

62. Presence of regular pits on protoconch (presenl,0; 
absent. 1) 

63. Presence of irregular pits on protoconch 
(present,0; absent, 1) 

64. Presence of apical spiral grooves (present.O; ab- 

65. Presence oT apical spiral segments (present.O; ab- 
sent, I) 

66. Presence of post nuclear spiral grooves (present.O: 

67. Presence of post nuclear spiral furrows (present.O: 
absent, 1) 

68. Presence of two-part sculpture on adult whorls 
(present.O; absent,!) 

69. Presence of post nuclear microradials (present.O; 
absent, 1) 

70.Presence of periostracal blades on major ribs 
(present,0; absent, 1 ) 

71. Presence of post nuclear periostracal setae 
(present.O; absent, 1 ) 

72. Shell colour (monochrome,0; flammulated,l) 

73. Presence of large major radial ribs (present.O; ab- 

74. Presence of elongate-bead' microsculptvirc 
(present.O; absent,!) 

75. Nature of beaded microsculplure (elongale,0; cir- 





NCHAR . . 

, 75 







SWAP . . . , , 







Weights applied 




Missing data code 

. - 9 



2.Statistics Derived from Consensus Tree 

Consensus fork index (component count) = 42 

CF (normalised) = 0.875 

Term information =400 

Mickevicrfs consensus information (CI) - 0.290 

Levels sum - 6473 

RohlfsCl(1) = 0.813 

Rohlfs - log CI(2) = 0.16566E-* ,03 

Weighted consensus fork = 0.361 



Character states for species in order of listing above. 


levicos : 

002000?????7777?????????????????????1001001000212443343??14201011011 101011? 
murphyi : 

00300021Q11Q0000010001000001001102001001002000204341334??3UG1 11 1111 101011? 

00200011010 10 00 000000002000100100000101100 10002 13351343? 7 2 3201 101011 1011 11? 

002 01 01 1070000000 100001 1000100 100000101 100 1300223351342 7 72320 110 10 11 101 111? 



1020002201 1100D000000001000100 110200100010 100020445344 6 7 7232 0101 11101 11011? 

0021202002000000000011020001000001010000001100343221221113 1001111101010101? 

00212020020000000000000300011000010000000021002311112122 121001111111011101? 

00212020020000000000100200010000010000000011003431211211 131001 1 11101010 101? 
monteit : 



002010?????????????????????????????? 100100 10002 12342334? 722211111111111011? 

02000 10002000000000000700001 01700 107000000 110033 11 10 111222 100 1111 1110110000 

vinitin : 

022001110101100001000110111000012200000000110043 2461060 3041101111111011001? 

02200110077010000 1000 100 11 10000122000000001100453341050104 11011111 110 L 1001? 

0220011000001000010001001110000122 0000000011004523420400041101111111011001? 
curtisi : 

2 200110000010000100010011100001220000000011004 3 22411403141200111111011101? 



contort i 

01000100120000000000000100100001 120100100121 1 15? 1010010??21201011 1110110 U? 

010001100200000000000002000100000101000000120034 311111101211011111110110001 



011101 10000020100000000200120000010000000012000320011111204101111111011001? 
earless . 

01110110011020100000000000120000210000000012002321212131122101111 1110 11001? 
peregri I 

012101100100000000000001010100012201000000211054 300001055201011111110111000 



kempsey : 

minutis : 

01210110010000000000000100010001020000000 0211054300011023201011111110110000 
colmani : 



01210110100000000000000100010001120000000 0211054301011001201011111110110000 

sericat : 

0221011001000100000000010001001002 0000000011000322312315204101111111011001? 

02210110011001001100000300010010020000000002 010322410304104101111111011001? 






annabel : 

kessner : 





Stanisic, J. 

Systematics and biogeography of eastern Australian Charopidae (Mollusca, Pulmonata) from 
the subtropical rainforests 1