THE TASM ANIAN M UDFISH, GALAXIAS CLEAVERI · galaxias, Galaxias brevipinnis G unther, and the m...

TH E T A SM A N IA N M U D F ISH , GALAXIAS CLEAVERI SCOTT, 1934, IN VICTORIA

J. D . K o e h n a n d T. A. R a a d i k

Freshw ater E cology Branch, D epartm en t o f C onservation and E nv ironm en t,123 Brown Street, H eidelberg, V ictor ia 3084

K o e h n , J. D . & R a a d i k , T. A ., 1991:12:31. T he T asm an ian m udfish, G a la x ia s c leaver iScott, 1934, in V ictor ia . P ro ceed in gs o f the R o y a l S o c ie ty o f V icto r ia 103 (2): 7 7 -8 6 .ISSN 0035 -9211 .G a la x ia s c leaver i Scott, 1934 is recorded for the first t im e from the A ire R iver basin in

w estern V ictoria, ex tend ing the know n w esterly d istr ibu tion o f the species. T he habitat, general b io logy and behav iou r o f the species are described and com parisons m ade w ith three species o f N ew Zealand m udfish (N eoch an na spp.). G. c leaveri is nocturnal and able to surv ive per iods w ithou t free surface water. Large areas o f poten tia l G. c leaveri habitat have been destroyed in V ictor ia , and the preservation o f freshwater w etland hab itats is essen tia l to the surviva l o f the species in th is State, w here a conservation status o f vu lnerable rem ains justif ied .

T H E TA SM A N IA N M U D F IS H , Galaxias cleaveri Scott, a m em ber o f the sou thern hem i­sphere fam ily G alax iidae, is a scaleless salm oni- form fish first described from T asm an ia in 1934 although specim ens had been dug up a t S trahan in w estern T asm an ia in 1900 (H all 1901, Scott 1934). T he species w as considered endem ic to T asm an ia un til 1980 w hen specim ens were d iscovered on m ain land A ustra lia (Jackson & D av ies 1982). In the 56 years since its d iscovery G. cleaveri has been recorded only spasm od i­cally, an ind ica tion th a t it is rare in term s o f d is tr ibu tion and abundance. C onsequently little is know n o f the general biology and ecology o f th is species; indeed, no t un til 1986 d id anyone repo rt th a t it possesses a m arine larval stage (F u lton 1986). In the p resen t paper we h ighlight the p resence o f G. cleaveri in V ic to ria and p ro ­v ide updated in fo rm ation to help fu rthe r w ork and the deve lopm ent o f m anagem ent s tra t­egies.

D eta iled descrip tions o f G. cleaveri w ere given by A ndrew s (1976), M cD ow all & F rankenberg(1981) and C adw allader & B ackhouse (1983), and the last au tho rs p rov ided a co lour p h o to ­graph. Even so, w orkers less skilled in the tax ­onom y o f fish m ay experience difficulty in d istingu ish ing G. cleaveri from the broad-finned galaxias, Galaxias brevipinnis G un ther, and the m ou n ta in galaxias, Galaxias olidus G un ther, w hich m ay occur w ith in the sam e river system .

T he follow ing m orpho log ical features m ay be used as a sim plified guide to d istingu ish ing adu lt G. cleaveri from G. brevipinnis and G. olidus (Fig. 1).

1. A nal fin slightly b eh ind orig in o f dorsal fin in G. cleaveri.

2. Shape o f dorsal and anal fins: low, rounded to ovoid, elongated posterio rly in G. cleaveri.

3. Shape o f caudal fin: rounded to tru n ca ted in G. cleaveri.

4. Shape o f flanges on caudal peduncle: large, long and ra ised in G. cleaveri.

5. Shape o f pecto ra l fins: large and rounded in G. cleaveri.

6 . Sm all head in G. cleaveri.1. Size o f eyes: sm all in G. cleaveri.8. Large long tub u la r nostrils: m ore p ronounced

in G. cleaveri.M orphological s im ilarities are exh ib ited w ith

th ree species o f N ew Z ealand m udfish (M cD ow all & W h itaker 1975, M cD ow all 1990): the C an terbury m udfish, Neochanna burrowsius (Ph illipps) (Skrzynski 1968, C adw allader 1975); the brow n m udfish, N. apoda G u n th e r (E ldon 1968, 1971); and the b lack m udfish, N. diversus Stokell (T hom pson 1987, M cD ow all 1990). In h ab ita t and hab its G. cleaveri shows sim ilarities to these species and to the dw arf galaxias, Gal- axiella pusilla (M ack) (B ackhouse & V anner 1978, Beck 1985, H um phries 1986).

D IS T R IB U T IO N

Previous records

G. cleaveri has been found to be patch ily d is tr i­bu ted in coastal areas in the no rth , sou th and west o f T asm an ia (A ndrew s 1976, M cD ow all & F rankenberg 1981, Fu lton 1990) bu t was re­po rted to be absent from F linders and K ing

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F igu re 1. M orphological features d istinguish ing G a la x ia s c leaver i (B) from G. o lid u s (A) and G b rev in in n h (C \ (A fter M cD ow all & Frankenberg 1981.) e v ip in m s yC).

Islands in the 1960s (F rankenberg 1967). A ndrew s (1976) was surprised at the absence o f G. cleaveri from the Bass S trait islands and m ain land A ustralia, even though at th a t tim e he d id no t know th a t the species possessed a m arine juven ile stage.

G. cleaveri was first recorded on m ain land A ustra lia in 1980 from the south-east side o f W ilsons P rom onto ry , V icto ria (Jackson & D av ies 1982). In 1983 ano ther ind iv idua l was recorded from the low er reaches o f the W ye R iver, O tw ay Ranges (K oehn & O ’C onnor 1990a; specim en N M Y A7594, D epartm en t o f Ichthyology, M useum o f V ictoria, M elbourne), ex tend ing the know n range o f the species in to w estern V ictoria. These tw o sites are referred to herein as sites 1 and 2 (Fig. 2, Tab le 1).

Subsequently , G reen (1984) repo rted G cleav- eri from a d ra in on F linders Island, Bass S tra it (specim ens Q V M 1984/5/6, Q ueen V ic to ria

A rt G aller>’’ L aunceston , and N M V A3391). T h is record com p leted a d istr i- bu tion pa tte rn co incid ing w ith th a t o f o th e r galaxnd species hav ing a trans-B assian d is tr i­bu tion nam ely G. brevipinnis, G. maculatus (Jenyns), G. truttaceus (V alenciennes) an d Gal- axiella pusilla (F rankenberg 1967).

It is no t surprising th a t G. cleaveri has been recorded in only tw o o f the num erous o th e r su r­veys prev iously conducted to determ ine the d is­tr ib u tio n o f freshw ater fish in coasta l V ic to ria (see K oehn 1990, K oehn & O ’C onno r 1990a K oehn et al. 1991). Sites sam pled during those surveys were m ostly in stream s ra th e r th an in

F igu re 2. L oca lities from w h ich G a la x ia s c leaveri has been recorded in V ictor ia (□ ); • = add itiona l sites surveyed in th is study, but w here G. c leaver i w as not found . Inset: areas w here G. c leaveri w as p rev iously recorded by Jackson & D av ies (1982) (1) and K oehn & O ’C onnor (1990a) (2).

sw am p and d ra in hab ita ts favoured by th is spe­cies. In add ition , specim ens o f G. cleaveri m ay have been m isidentified , particu larly before the species was recognised as occurring in V ic­to ria.

New records

In 1990 we surveyed likely G, cleaveri hab ita ts (sm all creeks, d ra ins and sw am ps) in the low land coastal p la ins o f the O tw ay R anges betw een Skenes C reek (143°43'E, 38°43'S) and the Jo h an n a R iver (143°23'E , 38°45'S) (Fig. 2). T w enty-four sites w ere sam pled using a Sm ith R oo t M odel 12 backpack electrofisher and d ip nets. D eta ils o f all sites sam pled and fish species collected are inc luded in K oehn et al. (1991). E leven specim ens o f G. cleaveri w ere collected from th ree sites (sites 3 ,4 and 5; Fig. 2 and Table 1) in the A ire and C alder R iver valleys, 35 km w est o f the locality on the W ye R iver w here the species was recorded by K oehn & O ’C onnor

(1990a). Specim ens have been deposited w ith the M useum o f V ictoria: N M V A 9512 (C alder R iver); N M V A 9513, A 9 5 10 (A ire R iver, drain); N M V A 9 5 1 1 (A ire R iver, b illabong).

P opu la tion density o f G. cleaveri a t the th ree sites could no t be estim ated because o f the dense vegetation and because o f the cryp tic and noc­tu rna l hab its o f G. cleaveri. T he op tion o f c lear­ing aquatic vegetation to sam ple m ore effec­tively was re jected as all sites w ere sm all in area.

Associations

All o ther species associated w ith G. cleaveri (Table 1) are com m on in the area (K oehn & O ’C onnor 1990a), are d iad rom ous (w ith the ex­cep tion o f Pseudogobius olorum), an d m ost have been recorded from low-lying sw am py hab ita ts . P. olorum is usually residen t in low er freshw ater o r estuarine areas as well as in coasta l lagoons (A llen 1989), and the specim ens o f G. brevipin-

SiteN o. W aterway

M ap N o . & Grid Ref.

A ltitude(m )

D ateSam pled

N o.C ollected

T ota l Length

R ange (m m ) A ssoc. Species

1 Freshw ater CreekA 8119 493746 20 29 .1 0 .8 0 5 37 -75* Sfeel,C gal

(tributary) 16.12 .80 102 W ye R iverB 7620 514196 10 ' 14.09.83 1 93* A m m ,C gal,A grayl,

T up ,B tr3 ■ C alder R iver (drain) 7620 178056 20 17.10.90 1 90 Sfeel,C gal4 A ire R iver (drain) 7520 147063 10 31 .05 .90 2 5 6 -5 7 Sfeel, Cgal

24 .08 .90 5 7 4 -9 6 Sgal,Bgal,Bsg -

17 .10 .90 2 8 5 -9 0 —

5 A ire R (b illabong) 7520 151068 15 16.10 .90 1 80 —

am m = lam prey am m ocoetes, G eo tr ia a u stra lis or M o rd a c ia m o rd a xCgal = com m on galaxias, G a la x ia s m a cu la tu sSfeel = short-finned eel, A n g u il la au stra lisSgal = spotted galaxias, G a la x ia s tru tta ceu sB g a l = broa d-fin ned g a la x ia s, G a la x ia s brev ip in n isT up = tupong, P seu d a p h r itis u rv illi iBtr = brow n trout, S a lm o tru ttaAgrayl = A ustralian grayling, P ro to tro c tes m a ra en aBsg = b lue-spot goby, P seu dogob iu s o lo rum

A from Jackson & D av ies (1982)B from K oehn & O ’C onnor (1990a)* standard lengths

T a b le 1. Sum m ary o f site and co llection details for G a la x ia s c leaveri in V ictoria.

nis co llected w ere juven iles m igrating upstream to adu lt hab ita t.

T he association o f eels and o ther species o f galax iids w ith N ew Z ealand m udfish (Neo- channa spp.) has been observed by E ldon (1968), and G. cleaveri has been associated w ith o ther galax iids and w ith sou thern pygm y perch, Nan- noperca australis G u n th er (Scott 1936, 1971).

In G. cleaveri hab ita ts surveyed on m ain land A ustra lia the o th er residen t galax iid species are essentially free-sw im m ing, w hereas G. cleaveri is ben th ic. T hus G. cleaveri m ay face na tu ra l in terspecific com petition o r p red a tio n only from eels w hich are also ben th ic and m ay occupy sim ilar hab ita ts.

Biogeography

T he d is tr ibu tion o f freshw ater na tive fishes in the O tw ay region appears to be p rim arily re lated to geom orphological cond itions th a t ex isted during and after the last g laciation 5 ,000-20 ,000 years ago (K oehn & O ’C onnor 1990a). Such con­d itions restr ic ted non -d iad rom ous freshw ater species to the larger B arw on R iver system to the no rth and to the A ire and G ellib rand R iver system s to the south-w est, w hereas only d iadro- m ous species inhab it the short coastal stream s.

T he d iad rom ous lifecycle o f G. cleaveri accoun ts for its occurrence in the W ye R iver.

F rankenberg (1974) suggested th a t G. trutta­ceus and G. brevipinnis, bo th species w ith life­cycles sim ilar to th a t o f G. cleaveri, m ay have m igrated to m a in land A ustra lia from T asm an ia w hen a land bridge ex isted du ring th e P le is to ­cene glaciation. A sim ilar m ig ra tion m ay be sug­gested for G. cleaveri. F u lton (1986) described a re tu rn to fresh w ater by ju v en ile G. cleaveri and suggested a m arine phase in the species’ lifecycle (F u lton 1990). T he d is tr ib u tio n and residency o f the larval phase o f G. cleaveri is unknow n, as is the possib ility o f land-locked po p u la tio n s o f the species no t possessing a m arine life phase. Such popu la tions are know n in o th er no rm ally d iad rom ous galax iid species (P o llard 1972 H um phries 1989, F u lton 1990, M cD ow all 1990).

T he p resent d is tr ibu tion o f G. cleaveri closely conform s to the region encom passed by the land bridge (W ilsons P rom on to ry to C ape O tw ay). L arvae develop ing in m arine w aters w ou ld be d ispersed m ore w idely and the species w ou ld be expected to be m ore w idely d is tr ibu ted . T he oc­currence o f larval galax iids as far as 700 km from the coast o f N ew Z ea land supports the theo ry o f M cD ow all (1978) th a t long-range d ispersa l o f

d iad rom ous species m ay occur, as is exh ib ited by G. truttaceus and G. brevipinnis w h ich are w idespread in V ic to rian coasta l s tream s (K oehn & O ’C onno r 1990a, 1990c). T he m ore restr ic ted d is tr ib u tio n o f G. cleaveri suggests th a t the larvae m ay be confined to estuaries, although

- fu rthe r surveys are needed to e luc idate d ispersal m echan ism s.

H A B ITA T

A com prehensive descrip tion o f the und istu rbed h ab ita t a t site 1 was g iven by Jackson & D av ies(1982). S ite 2 w hich is also u n d is tu rb ed is a sm all steep stream w ith a poo l-riff le sequence d ra in ­ing m oun ta inous fo rest country . T he substrate consisted o f cobbles and gravel, the flow was high, fast and tu rb id , the conductiv ity was 150 EC and w ater tem p era tu re w as 10°C. I t is poss­ib le th a t the specim en found a t th is site had been w ashed from areas o f low-lying pastu re during recent ra ins.

All new sites w ere characterised as being m odified w ith all r ip a r ian vegetation rem oved, and sites 3 and 4 had also been channelized.

Site 3 is a shallow channel (1 m w ide, 0.2 m deep) w ith a 0.8 m m ud substra te and w ith w ater 2 0 -3 0 m m deep d ra in ing from a spring in a pas­tu red paddock in to the C alder R iver. A t the tim e o f co llection (17 O ctober 1990) the channel had recently been excavated and little vegetation was presen t. D isso lved oxygen concen tra tions were 5.7 m g/L , pH was 7.0 and conductiv ity was 680 EC at 8.0°C. O n 27 F ebruary 1991 the d ra in was heav ily vegetated w ith a varie ty o f native and in troduced species o f aqua tic and pastu re veg­e ta tion , the w ater was 2 0 -3 0 m m deep, and the m ud substra te was 2 0 0 -4 0 0 m m th ick.

S ite 4 is a shallow d ra in (0 .2 -0 .4 m deep) w ith a silt substra te lead ing from a spring in a cleared paddock in to the A ire R iver. O n 24 A ugust 1990 m ost o f the site consisted o f a 2 m w ide channel together w ith a larger 15 m x 20 m area, and was densely vegetated w ith aqua tic species. C onduc­tiv ity was 185 EC a t 10°C. O n 17 O ctober 1990 d isso lved oxygen concen tra tion was 5.6 mg/L, pH was 6.7, and conductiv ity was 850 EC at 17°C. O n 27 F eb ruary 1991 there w as 100 m m o f m ud and a little w ater up to 20 m m deep in cattle h o o f p r in ts in the channel. T he larger area had sh runk to 3 m x 20 m w ith up to 100 m m o f w ater and 300 m m o f m ud. Sections o f the d ra in o ften becom e dry during sum m er bu t o ther sections always rem a in m o ist due to an underg round spr­ing (D . D enney pers. com m .).

S ite 5 is a sm all b illabong (60 m x 5 m x 0.8 m deep) abou t 30 m from the A ire R iver. N o flow was apparen t and the substra te was silt w ith dense aqua tic vegetation. O n 16 O ctober 1990 the w ater was dark tan n in in co lou r and h ad a d isso lved oxygen concen tra tion o f 4 .0 m g/L , pH o f 6.4 and conductiv ity o f 190 EC at 16.5°C. O ne specim en o f G. cleaveri was co llected from ju s t inside a large log ly ing in the w ater. O n 27 F e­bruary 1991 the site was com pletely dry, a con ­d ition no t unusual fo r th is season (D . D enney pers. com m .).

All sites are at low a ltitudes (a m ax im um o f 20 m above sea level) and close to the sea (a m ax i­m um o f 8.5 km from the sea bu t only 3 km from brack ish w ater). Except fo r site 2, all sites h ad no d iscern ib le flow and had m ud o r silt substrates and dense aqua tic vegetation. A lthough A ndrew s (1976) considered th a t G. cleaveri to l­era ted brack ish w ater, all ou r specim ens w ere collected from fresh w ater. T he d ra in lead ing from site 3, how ever, flowed in to reaches o f the A ire R iver w hich are know n regularly to con ta in an estuarine salt-wedge u n d er low flow con­d itions (J. K oehn pers. obs.).

T he presence o f G. cleaveri a t these sites is consisten t w ith its occurrence elsew here in sw am ps, d ra ins and sem i-perm anen t w aters. G. cleaveri was collected from stagnan t poo ls in T asm an ia (A ndrew s 1976) and from a d ra inage system usually dry in sum m er on F linders Island (G reen 1984). F u lton (1986) also recorded G. cleaveri from a dry section o f the E sperance R iver in T asm an ia. In N ew Z ealand th ree species o f m udfish, Neochanna burrowsius, N. apoda and N. diversus, have also been described as specialised to life in sw am ps, creeks and d ra ins th a t tend to dry up in sum m er (E ldon 1968, 1978, 1979a, M cD ow all 1990).

Scott (1934) described G. cleaveri as one o f the m ost specialised galaxiids in hav ing adop ted a m ode o f life su ited to such hab ita ts. Such ad ap ­ta tio n and an association w ith low a ltitude, sw am py hab ita ts is likely to ind icate a high de­gree o f dependance on the p resence o f ap p ro p ri­ate hab ita ts. C ollection o f the species in V ic to ria from d isjunct and highly m odified areas con­ta in ing in troduced vegetation suggests th a t these popu la tions m ay be rem nan ts o f a larger p o p u ­la tion th a t once existed w hen su itab le hab ita ts were m ore w idespread.

B EH A V IO U R

W e kep t fou r G. cleaveri from site 3 in a glass aquarium (0.36 x 0.45 x 0 .10 m ) con ta in ing a silt

substra te and vegetation from the site. T he be­h av iou r and position o f the fish were observed at in terva ls th roughou t each day for th ree weeks, and a t 5 m inu te in tervals fo r 1 hou r during one night.

G enerally the fish w ere inactive during the day, resting e ither on the substrate o r am ongst vegetation, and were difficult to locate because o f th e ir co lour and cryptic hab it. Ind iv iduals spent tim e resting e ither on the substrate w herever cover was available, o r am ongst dense w eed ju s t below the w ater surface. In bo th s itu ­ations, several ind iv idua ls shared the sam e cover and were in physical con tact w ith each other. T h is behav iou r is s im ilar to th a t described by E ldon (1969) in Neochanna apoda w hich is also te rr ito r ia l and aggressive to o ther species in aquaria (E ldon 1968). Ind iv iduals o f G. cleaveri exh ib ited no such aggressive behav iou r to each o ther o r to ind iv idua ls o f Galaxias maculatus o r G. truttaceus w hich were p laced in to the aquaria a t d ifferent tim es. A t n ight G. cleaveri were m ore active, continually m ov ing around open areas “ brow sing” . A t least tw o o f the fish were in open areas a t each observation . They im m ediate ly re­trea ted in to the vegetation w hen exposed to e ither w hite o r red light bu t had always re­appeared in the unvegetated areas by the next observation .

O ur observations ind icate th a t G. cleaveri is a noctu rna l, cryptic species w hich often inhab its the aqua tic vegetation ra th e r th an the substrate. O ther species o f galaxiids repo rted to be noctu r­nal are Neochanna apoda (E ldon 1968), Gal­axias brevipinnis (G lova & Sagar 1989a), and Galaxias vulgaris Stokell (G lova & Sagar 1989b).

A ESTIV A TIO N

Scott (1934) gave details o f the ability o f G. cleaveri to aestivate, though u nder unnatu ra l cond itions. T h is ab ility has been no ted by sev­eral o ther au thors (F letcher 1907, H all 1901, F u lton 1986) though the ir descrip tions m ain ly concern recovery o f aestivating ind iv iduals. Fu l­ton (1986) p rov ided pho tograph ic ev idence o f G. cleaveri aestivating during m id -sum m er u n d er a rock at least 10 m from free w ater.

M cD ow all & Pusey (1983) repo rted aestiv ­a tion in Lepidogalaxias salamandroides, and aestivation o f Galaxiella pusilla was suggested by M cD ow all & F rankenberg (1981). H u m p h ­ries (1985) tested th is suggestion by m ain ta in ing specim ens o f G. pusilla in an aquarium for 36 days w hilst low ering w ater levels and m a in ta in ­

ing oxygen concen tra tions a t less th an 5 ppm . F ish surv ived on the surface o f the m ud an d in a sm all hole for several days in the absence o f surface w ater. M cD ow all (1990) p resen ted ev i­dence o f the ab ility o f the N ew Z ea land m u d ­fishes Neochanna burrowsius an d N. apoda to surv ive dry periods, though bo th E ldon (1978) and M ered ith (1985) conc luded th a t these spe­cies do no t exh ib it true aestiva tion du ring w hich the ind iv idua l becom es to rp id and the no rm al ra te o f m etabo lism decreases.

W e investigated w hether bu rrow ing an d aesti­vation could be induced in G. cleaveri by p lacing tw o ind iv idua ls (101 m m and 78 m m TL ) in to a glass aquarium (0.3 x 0.62 x 0.3 m ) con ta in ing a substrate o f soil and m ud, a large flat rock a t one end, dense aquatic vegetation in the m idd le , and a p iece o f log at the o ther end. T he w ater level was low ered artificially, and heating an d il lum i­nation w ere p rov ided during the day by an incandescent globe. O n day 10 som e tunnelling in the m idd le section o f the tan k was observed, and one fish was seen lying near the w ate r su r­face w here it spasm odically gu lped w ate r an d air. O n day 14, w hen the w ater level h ad fa llen to10 m m above the substrate, one fish was resting in a vertical shaft in the m ud w ith its head ju s t p ro trud ing . O n day 22 n e ith e r fish cou ld be seen and no surface w ater rem ained , though w ater was p resen t in the open ing o f the shaft. L a te r on the sam e day the heads o f bo th fish w ere pos­itioned in the shaft open ing and th e ir bod ies were u n d er the m ud in h o rizon ta l tunnels. O n day 25 a series o f sm aller open ings w ere ob ­served in a line d irec ted aw ay from th e large shaft, p resum ab ly along the h o rizo n ta l tunne ls. O n day 31 there was no w ater in the p it o f the large shaft, the m ud substra te h ad begun to dry, and one G. cleaveri h ad its head p ro tru d in g from the tunne l in to the shaft below the surface o f the substrate. O n day 32, a fte r 5 days w ithou t any free w ater, the fish in the tunne ls stopped m ov ­ing. O n day 42 w hen the substra te h ad d ried to only 30 m m th ick the tw o G. cleaveri w ere seen th rough cracks in the dry m ud ly ing in th e tu n ­nels. T he tank was then slowly rehyd ra ted , and the fish recovered m ovem en t and em erged w hen the m ud becam e soft. T hey bo th im m ed ia te ly fed on earthw orm s and show ed no ill effects from surv iv ing in stagnan t w ater fo r 14 days and w ithou t surface w ater fo r an o th e r 14 days.

W hilst no t physio logically confirm ing the ab ility o f G. cleaveri to aestivate, o u r study shows th a t the species can su rv ive perio d s w ithou t free surface w ater by burrow ing in to the substrate.

L IFE C Y C L E

Two m ale G. cleaveri co llected a t site 4 on 31 M ay 1990 w ere in a r ipe spaw ning stage (Po llard 1972), w hereas ind iv idua ls co llected on 24 A u­gust 1990 w ere all spen t o r undeve loped , in d i­cating a w in ter spaw ning. T hese observations are consisten t w ith those o f A ndrew s (1976) w ho reported fully developed eggs in a specim en o f G. cleaveri exam ined in T asm an ia du ring M ay. A fter ageing w h iteba it re tu rn in g to freshw ater as approx im ate ly 2 m on ths o ld, F u lto n (1986) sug­gested th a t G. cleaveri spaw n du ring m id-w in ter, and he believes (W. F u lto n pers. com m .) th a t juven ile G. cleaveri re tu rn to fresh w ater during spring along w ith o th er galaxiids.

C adw allader (1975) and E ldon (1979a) con ­cluded th a t Neochanna burrowsius in N ew Z ea­land spaw ns during la te w in ter and early spring in hab ita ts frequen ted by adu lts. In con trast, E ldon (1979a) suggested th a t N. apoda spaw ns during m ost m on ths o f the year, especially w hen a drought breaks. B ecause E ldon (1971) found th a t in an aquarium N. apoda deposited eggs above the w aterline, he specu la ted th a t in the w ild the species deposits eggs ou t o f the w ater am ongst dam p vegetation and detritus. T he spaw ning location o f G. cleaveri has no t yet been found.

P resen t ev idence ind icates th a t G. cleaveri is un ique am ongst galax iids in possessing the tw o characteristics o f aestiva tion and d iadrom y.

TH R EA TS A N D C O N SE R V A T IO N STA TU S

G eneral th rea ts to freshw ater native fish in V ic to ria have been described by K oehn & O ’C onno r (1990b) w ho considered h ab ita t re­m oval and a lte ra tion a p rim e reason fo r the decline o f m any species. T he re liance o f G. cleav­eri on specific h ab ita t w ould app ear to m ake it susceptib le to h ab ita t changes, particu larly the loss o f w etland hab ita t.

T he m ain tenance o f fish hab ita ts has been rec­ognised as a key issue in m anagem ent o f the S tate ’s freshw ater fish fauna (K oehn & O ’C on­n o r 1990c). W hilst up to one-th ird o f the S ta te ’s w etlands have been destroyed (D C E 1988), m ost o f the assessm ents have re la ted only to w ater- b irds; fu rth e r assessm ents in re la tion to changes to fish hab ita ts are necessary. C orrick (1981, 1982) and C orrick & N o rm an (1980) assessed coastal w etlands in sou thern V ic to ria and as­signed them to the fo llow ing categories.1. F looded r iv e r flats: land in u n d a ted fo r very

sho rt periods fo llow ing ra in o r flooding.

2. F reshw ater m eadow s: land w ith w aterlogged soil fo r up to 3 m on ths each year b u t w here surface w ater is shallow an d transien t.

3. Shallow freshw ater m arshes: land w ith w ater­logged soil th roughou t the year, an d w here surface w ater m ay be p resen t fo r 6 to 9 m onths.

4. D eep freshw ater m arshes: land in u n d a ted to a dep th o f m ore th an 1 m during years o f average o r above average ra in fall.

5. P erm anen t open fresh w ater: w ater storages and na tu ra l lakes deeper th an 1 m .

6 . Sem i-perm anent saline w etlands.7. P erm anen t saline w etlands.

F rom ou r know ledge o f the h ab ita ts o f adu lt G. cleaveri, shallow freshw ater m arshes (cat­egory 3) and deep freshw ater m arshes (category 4) appear to p rov ide p erm anen t areas o f hab ita t. F looded r iver flats and freshw ater m eadow s m ay be used tem porarily by the species during m ig ration o f w h iteba it o r m ig ration o f adu lts to estuarine areas. O pen fresh w ater (category 5) is un likely to be used, particu larly i f lack ing veg­etation . T here is no ev idence to suggest th a t

. saline w etlands (categories 6 and 7) p rov ide su it­able hab itats.

In a study o f the Snowy R iver and G ippsland Lakes catchm ents, C orrick & N o rm an (1980) concluded th a t 25% o f shallow and 34% o f deep freshw ater m arshes have been lost. W ith in the P o rt P h illip Bay region, valuab le w etlands lost include the E d ith v a le -C arru m -S ea fo rd Sw am ps (C ham pion 1977, D onnelly et al. 1985) and the sw am ps o f the low er Y arra and M aribyr- nong R ivers (C astelnau 1872, excerpts from a d iary kept by J. F lem ings rep rin ted in Shilling- law 1879, K enyon 1934, and D ucker 1985). A fter study ing w etlands betw een P o rt P h ill ip Bay and M t E m u C reek in w estern V ictoria, C or­rick (1982) concluded th a t 79% o f shallow and 66% o f deep freshw ater m arshes had been lost since E uropean settlem ent, an overall loss o f 73% o f po ten tia l G. cleaveri hab ita t. T he m ost extensive hab ita t loss, how ever, is in South G ippsland w here 95% o f the na tu ra l freshw ater w etland once p resen t has been destroyed (C or­rick 1981). South G ippsland also inc ludes the largest areas o f po ten tia l G. cleaveri h ab ita t because it con ta ins coasta l-d ra in ing w etlands only w ith in 40 km o f the coast, un like the o ther areas w here w etlands ex tend as far as 150 km in land.

M ore than 23,000 ha o f w etlands have been lost in South G ippsland , inc lud ing K oo-W ee- R up, C ard in ia and Y allock Sw am ps, and sw am ps along the Pow lett and T arw in R ivers.

C orrick (1981) p red ic ted fu rthe r loss o f w etland areas th rough drainage, clearing, cu ltiva tion and flood m itiga tion and irrigation works. H is p re­d ic tion appears fulfilled because ca lcu lations from recent stud ies o f th is area (C orrick unpub l. data) show an overall loss o f 99% o f po ten tia l G. cleaveri h ab ita t (shallow freshw ater marshes* 94%, deep freshw ater m arshes 99%).

T he loss o f such large areas o f h ab ita t su itab le fo r G. cleaveri m ust be the greatest th rea t to th is species in V ic to ria in recent tim es and m ay ac­coun t fo r its fragm ented d istr ibu tion . S im ilarily, F rankenberg (1974) stated th a t the range o f G. cleaveri in T asm an ia had undoub ted ly been fragm ented due to the d ra in ing and clearing o f sw am ps.

In N ew Z ealand popu la tions o f the th ree species o f m udfish have declined drastically w ith the loss o f h ab ita t due to sw am p drainage, deve lopm ent and agricu ltura l practices (M cD ow all 1990). W hilst concern has been ex­pressed fo r all th ree species, Skrzynski (1968) and E ldon (1979b) have questioned w hether Neochanna burrowsius can surv ive, especially w ith con tinued agricu ltu ra l m odifications. C adw allader (1975) suggested th a t p reservation o f h ab ita t areas th rough the estab lishm ent o f re­serves should be instigated for th is species. A popu la tion o f N. burrowsius estab lished in an artific ial pond (E ldon 1988) surv ived for several years before dying ou t as a resu lt o f a pro longed drough t (N Z M A F 1990)

O ther th rea ts to adu lt G. cleaveri, such as in terspecific com petition and p redation by in ­troduced trou t, Salmo trutta, Oncorhynchus m ykiis o r redfin, Perea fluvialitis, are unlikely to be m ajor, especially in sw am py hab ita ts w ith poo r w ater quality unsu itab le to these species. G. cleaveri w h itebait m ay be subject to p redation , how ever, w hilst m igrating upstream . H ab ita t d istu rbance and com petition from o ther in tro ­duced species such as carp, Cyprinus carpio, goldfish, Carassius auratus, and tench, Tinea tinea, are possib le bu t difficult to assess. Sedi­m en ta tion is un likely to affect G. cleaveri unless the h ab ita t areas becom e com pletely filled.

Because G. cleaveri is restric ted to a special­ised aqua tic hab ita t, c lim atic changes m ay have serious im pacts, although the im pact o f the G reenhouse Effect is as difficult to p red ic t as it is fo r o ther native species (B urchm ore 1990). Low er w in ter ra in fa ll m ay affect spaw ning and particu larly access to the sea, and an increase in the tida l l im it m ay a lte r availab le w etland hab i­ta t fo r m atu re G. cleaveri.

In a recen t rev iew o f the co nserva tion sta tus o f na tive fish in V ic to ria (K oehn & M orison 1990) G. cleaveri was listed as vu lnerab le , a category includ ing “ taxa no t p resen tly endangered bu t w hich are a t risk by hav ing sm all p opu la tions an d /o r by occupying restr ic ted h ab ita ts suscep­tib le to rap id en v ironm en ta l change an d /o r popu la tions w hich are declin ing a t a ra te th a t w ould render them endangered in the n ea r fu tu re ” . A lthough we have docu m en ted ad ­d itiona l localities fo r G. cleaveri, o u r resu lts re in force the ra rity o f the species. T he red u c­tions in availab le freshw ater w etlands an d on ­going th rea ts to such h ab ita ts ju s tify th e re te n ­tion o f G. cleaveri in the vu lnerab le category.

C O N C L U SIO N S

G. cleaveri is m ore w idespread in V ic to ria th an prev iously believed, occupying n a tu ra l and m odified hab ita ts along low land coasta l areas, a t least from W ilsons P rom o n to ry to the w estern Otways. T here is a need fo r fu rth e r surveys to determ ine the range o f the species in sw am py hab ita ts w ith in and ou tside o f th is area, p a r ticu ­larly on F rench Island w hich con ta ins rem n an t tea-tree sw am p h ab ita t th a t once ex isted th roughou t the K oo-W ee-R up sw am p area and the en tire W estern P o rt ca tchm ent. T he d is­covery o f G. cleaveri in th is area w ould strengthen argum ents th a t p resen t p opu la tions are rem nan ts o f a once m uch larger, m o re u n i­form ly d istr ibu ted popu la tion . C o llection o f w h itebait as they ascend coasta l s tream s m ay also be a useful m ethod o f determ in ing G. cleav­eri d is tr ibu tion (K oehn & O ’C onno r 1990a).

T he behav iou r and h ab ita t needs o f th is species are s im ilar to those o f the N ew Z ea land m udfishes. T he specific h ab ita t needs o f G. cleaveri m ake it particu larly susceptib le to h ab i­ta t changes; therefo re the m assive reduc tions in su itab le freshw ater w etland h ab ita ts have u n ­doubted ly been the greatest th rea t to the species. T he m anagem ent and conservation o f such w et­land areas are v ita l fo r the p reserva tion o f G. cleaveri in V ictoria.

A C K N O W L E D G E M E N T S

W e th an k Bill O ’C onnor, D am ien O ’M ahony and M athew W estaw ay for field assistance, D av id D enney for local in fo rm ation , A ndrew C orrick fo r the use o f unpub lished d a ta and D arw in E vans for h is com m ents on th e m an u ­script. W ordprocessing was com pleted by K ae

W inch and Irene P ren tice. T h is w ork w as com ­p leted as p a r t o f the S ilv icu ltu re System s P ro ject funded by the D ep a rtm en t o f C onservation and E nv ironm en t, V icto ria.

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