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Egg Hatching and Life History of A EGG HATCHING AND LIFE HISTORY OF A FAIRY SHRIMP BRANCHINECTA MACKINI DEXTER (CRUSTACEA: ANOSTRACA) IN A MOHAVE DOCKET PLAYA DRY LAKE)' DESERT (RABBIT 11-AFC-2 LARS CARPELAN LON R. BROWN AND H. DATE Universityof California, Riverside, California 92502 RECD. MAY 01 2012 Abstract. Observations on populations of Branchinecta mackini in temporaryponds, which last for as little as 3 days to as long as 4 months, on a desert playa showed that hatching of dehydratedcysts (eggs) follows initial entry of water into the basin if salinity remains low. Salinity ranges from an initial 0.5%o to as much as 34%o as the pond evaporates. Hatching is continuous at constant low salinity, but since salinity generally increases rapidly, initial hatch is usually of short duration. Additional periods of hatching follow further inflows of water or after melting of ice, that is, after reductions in salinity. The duration of hatching is in- versely proportional to rate of increase in salinity. When salinity of small-volume summer ponds increases at rates above 500 ppm (1,000 Vmhos) per day, there is virtual inhibitionof hatching. Laboratory studies showed that egg hatching was controlled by both salinity and oxygen operating in various combinations to inhibit or stimulate hatch. The hatching characteristics of desiccated eggs collected from the dried mud of the lake basin differedfrom non-dried eggs obtained from laboratory cultures after ejection from ovisacs of living females. The findingthat a salinity-oxygencomplex regulates hatching in a desert pond permitstenta- tive explanation of a differencebetween branchiopods of humid and arid regions. In both cases the branchiopods are characteristicof astatic waters, and stimulation of the egg to hatch must be by some factor that changes at the time of origin of the temporary pond. In humid regions the factors of concern are temperatureand oxygen. If the water undergoes signifi- cant changes in salinity,as it does in arid regions, control of egg hatch may be by both salinity and oxygen, with temperaturelimited to control of rate of development. INTRODUCTION salinityaffects the rate of excystmentof Artemia for The Anostraca (fairyshrimps) are branchiopod salinc, a specialized Anostracan,exceptional waters. The con- crustaceans whose usual habitats according to its adaptationsto hypersaline fromits dormantstage by ex- Dexter (1959) are "rain pools and temporary trol of emergence has been clarifiedby Clegg pondsthat form from melting snow and ice." The ternalosmotic pressure there are reportsof osmotic dormantstage, which is usually an embryonated (1964). Although ionic adaptations of active stages of other egg or cyst,permits survival during periods when and Cole and Brown the ponds are dry. Because mere wettingoften branchiopods(e.g., Broch 1969, seems to have writtenthe is not sufficientto bringabout resumptionof de- 1967), Horne (1967) only paper reportingthat salinity inhibits the velopment,various factorsor combinationsof fac- offreshwater Anostraca in naturalwaters. tors, reviewed by Hutchinson (1967, p. 562), hatching of freshwaterAnostraca have been have been suggestedas stimulifor hatching. Fail- Most studies made not in arid but in humid regions, where ure to develop has sometimesbeen attributedto changes in salinityare negligibleand where reg- an obligatediapause, but it is morelikely an adap- ulationof hatch is predominantlyby oxygencon- to stimuliother than wetting,thereby in- tation centration. Thus Broch (1965) foundthat stim- under certainconditions un- hibitingdevelopment ulation of the dormantstage of Chirocephalopsis completionof life cycle. favorableto bundyi(Forbes) is by low oxygenconcentration. below was foundthat In the study reported it Low oxygentension can also inhibitexcystment as eggs of Brachinectamackini (Dexter) hatchingof reportedby Moore (1963), who foundthat eggs desertpool was regulatedmainly by in a Mohave of two speciesof Anostraca-Streptocephalusseali of oxygen changes in salinity,with involvement Ryder and Eubranchipus holmani (Ryder) influenceof temperature. tensions, and without would hatch only in well-aeratedwater. Bishop well withenviron- The lifecycle is so synchronized (1967) foundthat low oxygen concentrationsin- mentthat thereis littlewastage of eggs. hibit hatching of the conchostracan,Limnadic that Jenningsand Whitaker (1941) reported stanleyanaKing. Brewer (1964) reportedthat 1 Supportedby National Science FoundationGrants lowering of oxygen tension is the stimulusfor on the Ph.D. GB 3500and GB 6668. This paperis based hatchingof a calanoid copepod, Diaptomus stag- thesisof L. R. Brown. Manuscriptreceived December 18, 1969; acceptedAugust 8, 1970. nalis Forbes. Sussman and Halvorson (1966, 42 LON R. BROWN AND LARS H. CARPELAN Ecology,Volume 52, No. 1 p. 201) give even greatergenerality to oxygen as FIELD STUDIES ab- a stimulusin theirdiscussion of presenceand Introduction sence of oxygenas an environmentalinfluence on germinationof spores. This reportis restrictedto ponds appearingin Need for combinationsof factorshas been re- one basin on one dry lake bed on the Mohave ported. It has been known since 1941 that low Desert. Rabbit Dry Lake is in the Lucerne Val- oxygen stimulateshatching of aedine mosquito ley at an altitudeof 2,950 ft (950 m) in the south- eggs (Gjullin, Hegarty,and Bollen 1941). But easterncorner of the Mohave Desert in San Ber- Horsfall (1956 & 1958) showed that additional nardinoCounty, California (34?27' N, 117? W). factors,such as time for developmentand proper Althoughother California desert ponds have been temperature,are neededto "condition"Aedes eggs studied,this pond is chosen for reportbecause it beforethey can be stimulatedto hatchby low ox- was the most intensivelystudied (data are avail- ygenconcentrations. Interactions of two or more able fromsix differentfillings, during three sea- factorsare thus possible, and Weissman-Strum sons over a two-yearperiod) and also because it and Kindler (1963) have postulateda two-stage is the simplest biologically. One species of process of hatchingof A edes eggs that requires branchiopod-B. mackini Dexter-predominates low oxygen to stimulatebreakdown of a water- almost to the exclusion of other macroscopican- permeablebarrier, which then permitsthe entry imals. B. mackiniwas firstreported from brack- of water,an osmoticphenomenon. ish lakes in the state of Washingtonby Dexter It thus seems thatapparently conflicting reports (1956) and has since been reportedin California of what stimulatesand what inhibitsarthropod and Nevada (Dexter 1959). excystmentare due to specificdifferences, due to In nearby Mohave waters there are other involvementof more than one factor,or due to branchiopods,including another Branchinecta sp. all eggs of one speciesnot beingin the same "con- (B. lindahli) as well as Thamnocephalusplatyurus dition." The differencesin stimuli required by and Triops longicaudatus(which are oftenasso- humidregion (freshwater)anostraca such as Chi- ciated), togetherwith complex communitiesof rocephalopsis,which respondsto low oxygenand Cladocera, Ostracoda, Conchostraca,and Insecta. temperature,and the arid region (hypersaline) In Rabbit Dry Lake, the only macroscopicin- Artemia, which responds largely to osmotic vertebratein additionto B. mackiniis Branchi- changes,suggested that studyof a species inhabit- necta gigas Lynch. Unlike mackiniwhich is pres- ing waters of intermediatesalinity might con- ent in all ponds formed-winter, spring and tribute to understandingthe requirementsfor summer-B. gigas is presentin winterand spring resumptionof developmentand the broaderprob- only. It apparentlyis intolerantof summerheat. lem of adaptationto environment. B. gigas was firstreported from Washington in To put the Rabbit Dry Lake waters into per- 1937 and since then fromMontana, Nevada, and spective,we can compare them with conditions Utah (Dexter 1959); it has not previouslybeen reported elsewhere. In the tropical desert of reportedfrom California. Despite its large size northernSudan, Rzoska (1961) reportedconduc- (50-90 mm) it may be missed in samplingbe- tance at the end of the existenceof a temporary cause of its relativelysmall numbers. It is a pred- it in a ratio of pond to be 1,130 ,mhos (about 550 ppm). This ator on mackini,which outnumber gigas to 40,000 mackiniin some collections. terminalconcentration is about equal to the min- one was observedto consume about 35 young imum initial salinityof the Mohave desert pond, Gigas mackiniper day. Prey smallerthan 1.5 cm are not much than salinities by and greater reported eaten whole, larger ones are nipped in two and Prophet (1963) in studies in Kansas and Okla- onlythe abdomeneaten. Both prey and predator homa, where no pond had a conductivitygreater hatch at the start of the pond, so predationon than 555 Vmhos (about 275 ppm). The waters mackinioccurs from the verybeginning. B. gigas of Rabbit Dry Lake increase in salinity from is mentionedhere only incidentally, but it obviously about 0.5%yoto terminalsalinities about equal to is not unimportantto the general ecology of the sea water (34%yo)during the finalstages of sum- pond and to the populationdynamics of B. mac- mer ponds. Horne (1967) foundthat the salinity kini. Gigas is ignoredin the followingaccount in in several Wyoming ponds increased to a max- orderto focusattention on the egg-hatchingof B. imumof from0.5 to 1o%, except for one pond in mackim. which Branchinectalindahli and Streptocephalus texanus occurredin which the salinityincreased Field methods to 16.5%o The latterconcentration
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