Population biology of the freshwater shrimp Paratya curvirostris (Heller, 1862) (Decapoda: Atyidae)

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Population biology of the freshwatershrimp Paratya curvirostris (Heller,1862) (Decapoda: Atyidae)Alan Carpenter a ba Department of Zoology , University of Canterbury , Christchurch, New Zealandb Agricultural Research Division , Ministry of Agriculture andFisheries , P. O. Box 1654, Palmerston North, New ZealandPublished online: 30 Mar 2010.

To cite this article: Alan Carpenter (1983) Population biology of the freshwater shrimp Paratyacurvirostris (Heller, 1862) (Decapoda: Atyidae), New Zealand Journal of Marine and FreshwaterResearch, 17:2, 147-158, DOI: 10.1080/00288330.1983.9515992

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New Zealand Journal of Marine and Freshwater Research, 1983, Vol. 17 : 147-1580028-8330/83/1702-0147S2.50/0 © Crown copyright 1983

147

Population biology of the freshwater shrimpParatya curvirostris (Heller, 1862) (Decapoda: Atyidae)

ALAN CARPENTERtDepartment of ZoologyUniversity of CanterburyChrist church, New Zealand

Abstract The life history of the endemic NewZealand freshwater shrimp Paratya curvirostris(Heller, 1862) (Decapoda: Atyidae) in 3 streams inNorth Canterbury is described. Each female carriedon average 2000(±634) eggs and incubation tookabout 28 days at 14-18°C. Mixohaline planktoniclarvae were found in the Ashley Estuary and insmall numbers in fresh water. Post-larvae had acarapace length of 0.5-1.5 mm, and most developedinto males which occurred in freshwater streams. Ata carapace length of 5.0-6.0 mm males developedinto females which could be ovigerous at any time ofyear and could bear several successive broods.Estimated growth rate was greatest from late winterto early summer with a maximum weekly rate of 1.5mm carapace length calculated in spring. Thebehaviour of a female during egg hatching isdescribed in detail.

Keywords Atyidae; Paratya curvirostris; lifehistory; protandry; freshwater crustaceans;metamorphosis; salinity tolerance

protandry in the species and discussed its ecologicalsignificance.

Published information on the biology of Atydiaeis generally scattered and fragmentary, althoughresearch in Australia has provided detailed informa-tion on populations and life histories of Paratyaaustraliensis (Williams 1977; Gemmell 1978; Wil-liams & Smith 1979; Smith & Williams 1980) andCaridina nilotica (Glaister 1976). Early research onParatya compressa in Japan by Yokoya (1931) andKamita (1958a, b) has recently been complementedby Shokita's (1973,1976,1979) work on the naturalhistory of the Atyidae of the Ryukuku Islands(Japan).

Tropical Atyidae received attention from Darnell(1956) and more recently Hunte (1975, 1978) andHart (1980, 1981), and the life history of theEuropean atyid Atyaephyra desmaresti was studiedby Vorstmann (1955).

The embryology and development of atyids havebeen studied intensively (Ishikawa 1885; Gauthier1924; Nair 1949; Babu 1963; Glaister 1976), butlittle information on the larval habitat is availablewith the exception of the work of Williams (1977).

The research reported in this paper complementsa study of protandry in P. curvirostris (Carpenter1978), and deals with the habitat and populationecology of the species.

INTRODUCTION

The endemic atyid shrimp, Paratya curvirostris(Heller, 1862), is a characteristic inhabitant oflowland streams in many parts of New Zealand. It israrely found at altitudes above 40 m and typically isassociated with macrophyte beds or fringingvegetation (Carpenter 1976).

The population biology of P. curvirostris has beenstudied by Nielson (1972) who published a briefsummary of 3 years' fieldwork in Canterbury, andCh'ng (1973) who made a short study of P.curvirostris in the Horokiwi Stream near Wellington.Carpenter (1978) described the occurrence of

Received 11 June 1982; accepted 3 February 1983

tPresent address: Agricultural Research Division, Minis-try of Agriculture and Fisheries, P. O. Box 1654,Palmerston North, New Zealand

STUDY STREAMS

Three North Canterbury streams (Fig. 1) weresampled monthly: the South Branch of theWaimakariri River and the Ohoka Stream fromMarch 1975 to February 1976 inclusive, andSaltwater Creek from April 1975 to March 1976inclusive.

The South Branch arises as a seepage frombeneath the stopbanks of the main WaimakaririRiver, 5 km north-west of Christchurch. It flows for10 km through farmland before joining the mainriver near the northern motorway bridge 6 km northof Christchurch. Along much of its course it is fastflowing (up to 0.9 m s"1) over a shallow (<0.5 m)coarse-gravel bed. The sampling site was a 30 mStretch along the southern stream bank adjacent toDickies Road bridge, 2 km from the confluence withthe main river (5 on Fig. 1). Immediately below thesampling area the stream was heavily polluted by

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148 New Zealand Journal of Marine and Freshwater Research, 1983, Vol. 17

Fig. 1 Map of New Zealand showing location of study streams. Closed circlesindicate sampling stations. 1, Ashley River; 2, Waimakariri River; 3, SaltwaterCreek; 4, Ohoka Stream; 5, South Branch.

effluents from 2 meatworks. Below the outfalls a'pollution biota' occurs (Winterbourn et al. 1971)and no shrimps were found.

The catchment of the Ohoka Stream (4 on Fig. 1)lies between the Main Drain and North Eyre Roadand is mostly high-quality mixed cropping land. Thestream's basal flow originates from land runoff andrising artesian undercurrents. It flows for about 5km before joining the Main Drain and KaiapoiRiver about 3 km from the confluence of the

Kaiapoi with the Waimakariri. The Ohoka has amoderate flow (0.6-0.8 m s~x) and a bed composedof fine gravels. The sampling site consisted of 25 mstrips on each side of the stream about 200 m abovethe confluence of the Ohoka with the Kaiapoi Riverand Main Drain.

Saltwater Creek (3 on Fig. 1) runs into the AshleyEstuary 20 km north of Christchurch. It is fed byseveral large springs about 1 km east of Ashleytownship and is supplemented by runoff from

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Carpenter—Population biology of Paratya 149

Fig. 2 Number of mature and primary female shrimps found at each sampling site each month. Solid bars, ovigerousfemale shrimps.

Ashley Downs, 5 km west of the headwaters, duringwet periods. It runs 7 km across the northern end ofthe Canterbury Plains to enter the northwest cornerof the Ashley Estuary. The substratum of the creekis mostly mud, although patches of gravel alsooccur. Saltwater Creek is up to 1.5 m deep above theregion of tidal influence and current speed is usuallyless than 0.8 m s"1. Three sampling stations wereestablished on the stream: Station 1, at the head ofthe estuary; Station 2, 3 km further upstream; andStation 3, 5 km from the estuary.

METHODS

The number of eggs carried was assessed for 40 adultfemales 5.1-8.9 mm long. Eggs were removed fromthe pleopods with 10% KOH, spread in a petri dish,and photographed by placing the dish on photo-graphic paper and exposing it for 5 seconds. Eggswere recorded with a hand counter on examining a10 x 10 on print. This method allowed the small andnumerous eggs to be counted without magnificationand with a high degree of repeatibility.

Adult shrimps were collected from the studystreams at approximately monthly intervals for ayear. Collections were made by sweeping a hand net(195 |j.m mesh, 20 cm diameter) through the fringevegetation (Carpenter in press) along a 2 m stretchof bank. This procedure was modified at 2 sites inSaltwater Creek. At Saltwater Creek 1, shrimpswere caught by sweeping beds of Ruppia polycarpaMason until about 50 amimals had been taken,whereas at Saltwater Creek 2, samples were takenfrom beds of Elodea canadensis Michaux. Fringesamples at this station were taken from the standard2 m stretch of bank.

In the laboratory, samples were sorted and allshrimps were preserved in ethylene glycol afterfixation for 24 h in 10% formalin. Carapace length(c.l.) (the distance from the base of the supraorbitalspine on the left side of the carapace to the postero-dorsal edge of the carapace) was measured using aneyepiece micrometer fitted to an Olympus model Xstereoscopic microscope. Shrimps under 3.0 mm c.l.were regarded as juveniles, as they possessed nosecondary sexual characters. Males above this sizewere identified by the distally broadened propodus

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' 60-

13 23 2i 44 77 35 59

of the third and fourth pereiopods and by thepresence of an appendix masculina on the secondpleopod. Females were identified by the absence ofboth the broadened propodus and the appendixmasculina. Mature females also had deeper andbroader abdominal sternites than males.

RESULTS

EggsIn general, the numbers of eggs per femaleincreased with carapace length and ranged from 900to 4000 per individual. The average number of eggscarried by the 40 females examined was 2010(±643).j ^ e relationship between maternal body size andnumber of eggs is given by the equation

Fig. 3 Percentage of ovigerous females with eggs soon to number of eggs = -764 + 407 x carapace lengthhatch taken each month. (f = 0.377; P<0.01).

F M A M J J A S O N DMonths

40-

20-

0

April 26 69% August 155 December

n

17

40-

20-

May

"LrTh-r-.

234 September 26

rTHT -rTrn

January

rf

95

Iit 40-

20-

0

40a

20-

92%

n

r =

June 97 October

Ik-

276 February 86

68%

aJu l y 107 N o v e m b e r 33 M a r c h 115

0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9Carapace length (mm)

Fig. 4 Size distribution of the shrimp population at Saltwater Creek 1, April 1975-March 1976.

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There was no significant difference between thenumber of newly laid eggs per female and thenumber of eggs per female close to hatching,indicating no losses during development.

The pattern of occurrence of ovigerous femalesvaried between sites and no general pattern could bedetermined (Fig. 2). Ovigerous females werepresent throughout the year in all 3 streams. No eggswere found to be close to hatching (as indicated bywell developed eyes) from March to July inclusive.However, from August to November and inJanuary, the percentage of females with eggs aboutto hatch was 30-40%. The percentage was evenhigher in February of both years and in December,reaching a maximum of 70% in February 1976 (Fig.3).

In the laboratory, egg development took about 28days at temperatures ranging between 14 and 18°C.Of 30 females whose eggs hatched in the laboratory,

all but 2 moulted within 2 days of the completion ofegg hatching.

Hatching of eggsOne female Paratya curvirostris was kept alivesuccessfully from egg extrusion to hatching. It took1.5 days between the first and last egg hatching andfor infertile eggs and debris to be cleaned from thepleopods.

As hatching proceeded, the female's behaviourconsisted of 3 cyclic phases: periods of restalternating with periods of strong pleopod beating,and less frequent periods of egg mass cleaning.Cleaning of the egg mass was carried out by thefourth and fifth pereiopods, which were scrapedacross the egg mass surface. Material removed bythis activity, presumably empty egg cases and deadlarvae, was passed immediately to the mouth via thethird maxillipeds and ingested.

40-

0Carapace length (mm)


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40-

20-

0

April 51 August 30 December 80

J]

40-

^ 2 0 -

ig 0I"-40H

May 23 September 33

MTh

January 41

1iTk

20-

0

40-

20-

June 21

T l

October 29 February

K

0

July 28

• i i p

November 33 March 85

0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5Carapace length (mm)

I 9


Periods of pleopod beating consisted of shortbursts of rapid activity alternating with longerperiods of slower movement. The bursts of rapidactivity usually involved 3-5 non-synchronised beatsof all the pleopods. This cycle was repeated anumber of times, sometimes with cleaning activityincluded as well. Rest periods became longer as thehatching period progressed. The space between thelast 2 pairs of egg-carrying pleopods was clearedcompletely of eggs before hatching from moreanterior spaces began.

LarvaeThe eggs of Paratya curvirostris hatch into typicalprimitive caridean zoeae (Gurney 1942). The newlyhatched larva of this species has been fully describedby Ch'ng (1973).

During sampling for adult shrimps some larvaewere found in all 3 freshwater study streams, but

never more than 10 per month. However, in theAshley Estuary at the mouth of Saltwater Creekgreater concentrations of larvae were found in abrief survey, suggesting that the larvae may occurlargely in brackish water (Carpenter 1982).

Population structure and growthSaltwater CreekLength-frequency histograms of shrimps collectedfrom 3 stations on Saltwater Creek from April 1975to March 1976 are shown in Fig. 4-6.

The smallest size class of shrimp caught was 0.5-1.0 mm c.l. However, the next size class (1.0-1.5mm c.l.) was much more common. Such smallshrimps were found only at Saltwater Creek 1, theonly estuarine station sampled. The only individualssmaller than these were the zoeae, indicating thatthe size at which the animal becomes a post-larva isin the range 0.5-1.5 mm c.l. Most recruitment of

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40-

20-

April 255

ITL

August 363

September 333

December 314

January 314

20-

0

June 284 October 505 February 400

40-

20-

July 370 November 390 March 523

0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 8 9Carapace length (mm)

Fig. 7 Size distribution of the Saltwater Creek shrimp population, all 3 stations summed, April 1975-March 1976.

young occurred from May to October and in March1976. However the overall recruitment patternobserved may not be reliable, as sample sizes inApril, September, and December were low (Fig. 4).

At all 3 Saltwater Creek stations, shrimps in thesize range 3.0-6.0 mm c.l. were most common inmost months. The monthly size distribution ofshrimps collected at Saltwater Creek 2 did not varymuch during the study and no post-larvae occurredat this station (Fig. 5). A relatively constantsize/frequency distribution was also found at theupstream Station 3, although the population at thisstation was fairly small (Fig. 6).

In Fig. 7 the data for all 3 Saltwater Creek stationsare combined. This provides the best perspective ofthe population dynamics which cannot be inter-preted from data from a single site.

Females were categorized as (a) ovigerous, (b)mature (above 5.0 mm c.l.), and (c) primary (below5.0 mm c.l.), the latter being individuals that had

not undergone protandry (Berkeley 1930; Butler1964; Carpenter 1978). Very few females werefound in Saltwater Creek except at Station 2 (Fig. 2)where the majority of the breeding populationoccurred. The number of primary females was lowthroughout the year, and April, November, andDecember were the only months in which noovigerous females were found. The ratio of males tofemales was 7:1.

One of the most useful methods of estimatinggrowth rates from polymodal field data is theprobability paper technique outlined by Cassie(1950). Because of the short life history andrelatively long breeding season of Paratya curviros-tris, this method was not successful in this study.Instead, growth was estimated by plotting modalvalues shown in monthly population structurehistograms as suggested by Simpson et al. (1963).

In Saltwater Creek, recruitment occurred fromMarch to July and young shrimps grew rapidly from

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40-

20-

0

40H

March 74 July 292 November 170

20-

0

40-

20"


309 September 146 January 292

l i l t - .June 98 October 122 February 134

° 0 1 2 3 4 f T ] 8 ^ 0 1 5 6 7 8 9 0 1 2Carapace length (mm)

Fig. 8 Size distribution of the shrimp population in the Ohoka Stream, March 1975-February 1976.

late winter to reach their full size in mid summer(Fig. 10). An older generation appeared as aconstant-sized component and it is likely that theseanimals formed the main breeding population, i.e.,they bred in their second summer.

Ohoka Stream and South BranchIn both the Ohoka Stream (Fig. 8) and the SouthBranch (Fig. 9) the smallest size classes of shrimpscaught were 1.5-2.5 mm c.l., suggesting that the firstpost-larval stage(s) occurred elsewhere, probablydownstream of the sampling sites. Similarly fewlarge shrimps (over 6.0 mm c.l.) occurred at eithersite. The smallest shrimps were found in April in theSouth Branch and March-August in the OhokaStream, but in both streams these small shrimpswere at least 0.5 mm c.l. longer than the expectedsize of first stage post-larvae.

Females were very scarce in the South Branchexcept for primary females in August. No ovigerous

individuals were found in May, June, October, orMarch (Fig. 2). Similarly, no ovigerous femaleswere found in the Ohoka in early winter and veryfew were taken in autumn or spring. The massiveincrease in numbers of females in February mayhave been related to the placement of culvertsacross the stream in December just below the studysite. This effectively stopped further upstreammovement of young shrimps, and older individualsmay also have been washed down through theculverts. Nielson (unpublished notes) found adultshrimps in drift samples in the Avon River in 1941-1942.

The ratio of males to females in the South Branchwas 15:1, contrasting with the 5:1 ratio in the OhokaStream and the 7:1 ratio in Saltwater Creek.

The modal size of the South Branch populationincreased steadily throughout the winter and in earlysummer was similar to that of shrimps fromSaltwater Creek. There was no net growth in the

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40-

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0

March 211

j

July 450 November 313

40-

20-

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sI

20-

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40-

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May 398 September 184 January 201

June 314 October 201 February 175

o4-P i P i i I i i P0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9

Carapace length (mm)

Fig. 9 Size distribution of the shrimp population in the South Branch, March 1975-February 1976.

Ohoka population until early spring (Fig. 10), butsubsequent rapid growth brought the mean size ofanimals in the population to a level similar to thatfound in the other 2 streams in early summer.

DISCUSSION

The average number of eggs carried by femaleParatya curvirostris is very high compared with mostother species of Atyidae. Advanced atyids of thegenera Caridina, Syncaris, Jonga, Halocaridina,and Troglocaris all have small numbers (5-100) oflarge eggs (Babu 1963; Hedgepeth 1968; Juberthie-Jupeau 1969; Shokita 1973, 1976, 1979; Chinnayya1974; Glaister 1976; Couret & Wong 1978) whereasthe more primitive genera Paratya, Atyaephyra, andAtya have large numbers (100-10,000) of small eggs(Vorstmann 1955; Chace & Hobbs 1969; Shokita1973, 1976, 1979; this study).

Shokita (1973; 1976; 1979) showed that specieswith small eggs usually have estuarine or marinelarvae whereas those with large eggs live exclusivelyin freshwater. Exceptions to this general rule areParatya compressa, which has numerous small eggsbut is confined to freshwater (Yokoya 1931; Kamita1958a, b), and Jonga serrei, which has a few largeeggs and estuarine larvae (Chace & Hobbs 1969).Thus Paratya curvirostris, with numerous small eggsand an apparently estuarine larva, conforms to thegeneral atyid pattern.

In Paratya curvirostris, as in all other speciesinvestigated, a linear relationship between the sizeof the female and the number of eggs carried wasfound (Shokita 1973,1976,1979; Hart 1981). The 28days that female P. curvirostris took to incubate theireggs at 14-18°C is similar to that of anothertemperature atyid, Atyaephyra desmaresti (Vor-stmann 1955), but contrasts with the incubation

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South Branch Stream

Fig. 10 Growth rates of shrimp populations from the 3 study streams, estimated from population size modes shown inFig. 7, 8, and 9. Lines fitted by eye.

periods found in tropical atyids, e.g., Caridinanilotica, in which incubation time varies cur-vilinearly with temperature from 11 days at 30°C to40 days at 18°C (Hart 1980).

Ovigerous females were found throughout theyear although not at all sites in all months. As noovigerous females had eggs about to hatch duringthe period March-July, it is possible that the eggs'overwinter', particularly as the water temperaturesin the study streams in March were higher than inAugust (the coldest month) when late stage eggswere found again (Carpenter 1976).

The growth rate of Paratya curvirostris estimatedfrom population histogram modes was similar tothat of Pandalus danae, a marine protandrousshrimp (Berkeley 1930) and for normallygonochoristic species such as Crangon septemspinosa(Wilcox & Jeffries 1973). However, the recordedgrowth rate of Paratya australiensis in Tasmania

(Walker 1972) is only about half that of P.curvirostris, whereas the tropical Caridina nilotica isabout twice as fast (Hart 1980).

Although the data obtained in this study areincomplete it is possible to reconstruct a probablelife history of the species in small coastal streams.During spring and summer, females living in thestream above the influence of salt water carry 900-4000 eggs for about 28 days, after which they hatch.The zoeae are planktonic and are carried down-stream by the current to the estuarine reaches.Development to metamorphosis takes an undeter-mined time, but as the first eggs probably hatch inAugust and recruitment was first recorded in May,larval life may be about 8 months.

It is possible that the larvae of Paratya curvirostrismaintain themselves in estuarine waters bybehavioural reactions to salinity change similar tothose described by Hughes (1969 a, b) for various

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Peneidae. As they grow, larvae probably movetowards the head of the estuary where theymetamorphose and continue to move upstream.Most develop into males, but about 1.5% becomeprimary females. After further growth the malesundergo a sex change to become secondary females(Carpenter 1978). After completing the transforma-tion, which takes a number of moults, they lay theirfirst clutch of eggs. They may produce a clutch amonth for 7 or 8 months, before dying at amaximum size of about 10 mm c.l. in the autumnand winter. The maximum length of post-larval lifeis likely to be about 2 years, as has been suggestedby Williams (1977) for Paratya australiensis.

The need for upstream movement is morecomplicated than it appears at first sight, for it wasfound that after a flood the modal size class of theshrimp population at Saltwater Creek 1 increased.Thus, upstream movement is necessary for bothreaching the area where breeding occurs, and toreturn animals to that reach of the stream aftereither floods or drift. Shaw (1981) mentioned large-scale upstream migration in the Waikato River. Allthe direct evidence that has been reported herepoints to a cyclic migratory life cycle, butcircumstantial evidence indicates much remains tobe clarified. Particularly, the life history of Paratyacurvirostris in streams such as the South Branch,which appear to prevent a migratory life historyoccurring, needs elucidating.

In New Zealand, retention of the migratory lifehistory may have been permitted by the absence ofcompetitively superior Palaemonidae from freshwa-ters and the upper reaches of estuaries. Availableevidence suggests that the primitive Atyidae cannotcoexist with palaemonids (Reik 1959; Orr 1971;Walker 1972), and it is only when they are the onlyfreshwater shrimps present that the primitiveAtyidae form a major component of freshwaterecosystems.

ACKNOWLEDGMENTS

I am grateful to Dr M. J. Winterboum for his enthusiasmfor this project. Dr W. C. Clark and Dr M. B. Jonescontributed to my understanding, particularly of protandryand the mixohaline life history. The figures were expertlydrafted by Raewyn Moar, and Monica Liddell typed themanuscript.

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propinqua De Man. Indian journal of fisheries 10A :107-117.

Berkeley, A. A. 1930: The post-embryonic developmentof the common pandalids of British Columbia.Contributions to Canadian biology and fisheries N.S.6: 79-214.

Butler, T. H. 1964: Growth, reproduction and distributionof the pandalid shrimps in British Columbia.Journal of the Fisheries Research Board, Canada21 : 1403 -1452.

Carpenter, A. 1976: Biology of the freshwater shrimpParatya curvirostris (Heller, 1862) (Decapoda:Atyidae). Unpublished M.Sc. thesis, Departmentof Zoology, University of Canterbury.

1978: Protandry in the freshwater shrimp Paratyacurvirostris (Heller, 1862), with a review of thephenomenon and its significance in the Decapoda.Journal of the Royal Society of New Zealand 8 :343-358.

1982: The habitat and distribution of thefreshwater shrimp, Paratya curvirostris (Heller,1862) (Decapoda: Atyidae) in North Canterbury.Mauri ora 10 : 85-98.

Cassie, R. M. 1950: The analysis of polymodal frequencydistributions by the probability paper method. NewZealand science review 8: 89-91.

Chace, F. A.; Hobbs, H. H. 1969: The freshwater andterrestrial decapod crustaceans of the West Indieswith special reference to Dominica. United StatesNational Museum, bulletin 291. 258 p.

Chinnayya, B. 1974: The embryonic and larval develop-ment of Caridina weberi De Man in the laboratory(Decapoda: Atyidae). Broteria serie trimesral-ciencias natrais 43 : 119-134.

Ch'ng, T. K. 1973: Aspects of the biology of thefreshwater shrimp Paratya curvirostris (Heller) inthe Horokiwi Stream. Unpublished B.Sc.(Hons.)project, Zoology Department, Victoria Universityof Wellington.

Couret, C. L.; Wong, D. C. L. 1978: Larval developmentof Halocaridina rubra Holthius (Decapoda: Aty-idae). Crustaceana 34 : 301-309.

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Documents

Population biology of the freshwater shrimp Paratya curvirostris (Heller, 1862) (Decapoda: Atyidae)