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TRANSLITTORALTALITRIDAE (CRUSTACEA: AMPHIPODA) AND THE NEED TO RESERVE TRANSITIONALHABITATS: EXAMPLES FROM TASMANIAN

SALTMARSHESAND OTHERCOASTALSITES

AlastairM.M. Richardson, Roy SwainandVeraWong1

Departmentof Zoology, University of Tasmania,GPOBox 252C, Hobart,Tas. 7001, Australia.

'Present address: 8 CedexCt, Glen Waverley,Vic. 3150, Australia.

Abstract

Richardson,A.M.M.,Swain, R.andWong, V., 1997. Translittoral Talitridae (Crustacea:

Amphipoda)andtheneedtoreserve transitional habitats:examplesfrom Tasmaniansalt-

marshes andother coastal sites. MemoirsoftheMuseumofVictoria 56(2): 521-529.

Transitionalhabitats,e.g.,those thatliebetweenseaandland, are often ignoredbecause of thedifferentbackgroundsofmarine andterrestrial biologists,but they areimportantforthe seriesofspecieswhichspantheboundary.The amphipodfamily Talitridae hasperhapsthe widesttranslittoralrangeandsoprovides evidence fromwhichthe evolutionary processof landcolonisationcan be deduced.Thetalitridsfound ontwosea-land transitionsin Tas- mania,asandy-beachto rainforestandasaltmarshtodry woodland,are described,andthe significance oftheirdistributions isdiscussed.The need to reserve thefull rangeof such translittoral seriesis noted. Thiswill notnecessarily be achieved byestablishing reserve boundarieson the basisofvegetation associations.

Introduction

Transitionalhabitats, thatisthoselyingbetween major habitats (land and sea, freshwater and land) tend to fall through the net ofbiological investigations (Little, 1990). Marine biologists andterrestrialbiologistsdevelopfromsuchdif- ferent backgrounds that they have little to do witheach otherand so the zones ofinteraction between their spheres of operation tend to be ignored. While marine biologists oftenwork in the intertidalzone,their studiestendtobe con- centratedwell belowthe high watermark,while fewterrestrialbiologistsapproachthehigh water markfromthe otherdirection.Thustheamount ofstudyon the intermediate zoneissmall, and

thisisparticularlyobviouswhenthe transitional zone is extensive, as it is in the most sheltered sea-land interfaces, in saltmarshes and man-

groves.

Andyet the interface between thetwo major

habitats is in many waysmore important than the sea or land alone. The colonisation of land was one of the greatest evolutionary events in the history of those seven phyla that have achievedterrestriality.Although mostofthecol- onisation eventshappened a long timeago, the habitats in which they occurred are still there, and in some cases species can be found filling many ofthe intermediate stages. In a fewtaxa, which might be called translittoral (Chester,

1992),seriesofspeciescan befound withdistri- butions spanning the entire range from fully

marinetofullyterrestrial.Thebestexamplesare theamphipod crustacean family Talitridaeand thepulmonatemollusc familyEllobiidae.Trans-

littoralgroups ofthistype provide very valuable opportunities to reconstruct the evolutionary events leading to life on land (Little, 1989,

1990),andtheybecome morevaluablestillwhen

the species involved are relatively closely- related, as in the two families mentioned above.

In thiscontribution wewill presentevidence of translittoral distributions in the Talitridae anddemonstratethatsuchdistributions require a broader approach to reserve design than has been adopted hitherto.

Talitridae

TheTalitridae include theonly fully terrestrial

members ofthe Amphipoda, one of only two crustacean groups (the other is the Isopoda) which havemembersthatarefully terrestrial,ie.

independent of free water for all oftheir life stages. The Talitridae have been looselyclassi-

fied by Bousfield (1984) into four groups, the palustral talitrids (marsh-hoppers), the beach-

fleas,thesandhoppers andthelandhoppers.The marsh-hoppersinclude the mostplesiomorphic forms and they arefoundin thelower levelsof saltmarshesandmangroves,wheretheyhavean amphibious lifestyle, entering and swimming

efficiently inseawater even atlowtide. In some

places (Tasmania:Chester, 1992,South Africa:

521

https://doi.org/10.24199/j.mmv.1997.56.46

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522 A.M.M. RICHARDSON

Griffiths, 1976) their distributions extend into fullyfreshwaterstreams. InsoutheastAustralia the genus Eorchestia is widespread in salt-

marshes and the lower reaches of associated streams (Richardson, 1993).Thebeachfleasand sandhoppersarethe typicaltalitridsoftheinter- tidal zone, distinguished by the adaptations of sandhoppersfortheirparticularhabitat,ieloose sand. Thus the sandhoppers are large, robust and spiny talitrids, sometimes with unusual morphological adaptationsforlifeinsand.They

are efficient burrowers. The beachfleas, in con-

trast,arcnotwell-developedforburrowing,pre- ferring to seek refuge under cast seaweed or stones (non substrate modifiers: Macintyre, 1963,ornestlers: Griffiths, 1976); they are gen- erally more lightly built and weakly spined.

Beachfleas are usuallyfoundonrocky ormuddy

shores, but there is at least one undescribed species inTasmania which hasbecome conver- gent with thesandhoppers and forages on mid-

tidalwetsand duringlowtide,taking refugeina shallow burrow during high tide.

The landhoppersare found ina range ofter- restrial habitats in the Southern Hemisphere, Japan, the Phillipines, Central America, the Caribbean and many oceanic islands. In Tas- maniathey arcfound throughouttheisland,but wheretheyhavecolonised continents(Australia, southernAfrica)theyareconfinedtothehigher rainfall zones around the continental margins.

Bousficld (1984) recognises two subgroups, the plesiomorphic simplidactylate genera, which seemtobedescendedfromamarsh-hopperlike ancestor, and the cuspidactylate genera which apparently arose from the beachfleas. The sim- plidactylate formsareconfinedtoAustraliaand southern Africa.

Thusthe Talitridae showa truly translittoral range from the semi-aquatic marsh-hoppers to thelandhoppers.Thesisterfamilyto theTalitri- dae is the Hyalidae, and it is widely- accepted that talitridsdescendedfroma hyalid-like ances- tor (Bousficld, 1984). Aquatic hyalids may be found adjacent to intertidal talitrids in fully

marine and brackish waters, providing a valid extension to the translittoral series.

Itseemslikelythat thetalitridamphipodscol- onised land relatively recently compared to otherterrestrialarthropods. Little (1990) noted that the fossil record ofthe amphipods is only from theQuaternary, but Bousfield (1984)and Bousficld and Poinar (1995) suggests a mid- Cretaceous origin for the simplidactylate land- hoppers of the Southern Hemisphere conti-

nents,contemporarywithanextensive radiation ofthe angiosperm flora in Gondwana.

Tasmaniahasaremarkablyrich talitridfauna (Richardson, 1993, 1996) which includes rep- resentatives of all Bousfield's groupings. The taxonomy of the landhoppers is well-known (Friend, 1987)andthe diverseintertidalfaunais currently being described by the senior author.

Friend (1987) divided theTasmanian landhop- perfaunaintothree ecological groupings,oneof which he called the coastal group. Five or six species inthreegenera make upthisgroupand they have distributions which are strongly restricted to a narrow zone, less than 100 m

wide, immediately behind the hightide mark.

Tasmaniaalso retainssomeplaces,especially in the west, where transitions from land to sea remain undisturbed, and clothed with native vegetation.Theseprovideimportant opportuni-

tiesto studythe natural distributionsoftalitrid

amphipods, isopodsandothergroupsacross the transition. Wewill considertwo cases, a sandy beach to wet sclerophyll forest at South Cape Bay and asaltmarshtodry woodlandtransition at Lutregala Marshon Bruny Island. Details of these studies are described elsewhere (South Cape Bay: Richardson et al., 1991; Lutregala Marsh: Richardson and Mulcahy, in press).

Studyareasand methods South Cape Bay

SouthCapeBayisintheextremesouth of Tas- mania(Fig 1). It facesalmost duesouth, but is shelteredfromthe prevailingsouthwesterlyand westerly winds by promontories to the west.

Because ofstrongwave action, the sand above thehighwatermarkiscolonisedbyfew foredune plants, so that low coastal scrub and forest appears within a few metres of high water.

Closesttotheshoretheshrubs Correa backhou- siana, Cyathodesabietina, Westringiabrevifolia

and Olearia phlogopappa form a low dense cover, fiveto ten metresdeep, whichgives way

to lowforest with emergent treesof Eucalyptus nitida, sassafras,Athewspermamoschatum,cel- ery top pine, Phyllocladus aspleniijblius, and southernbeech,Nothofagus cunninghamii. This coastal forest has an understorey of native laurel,Anopterusglandulosus, andoften cutting grass, Gahnia grandis. The ground layer is

mostly absent, apart from a few mosses and herbs.The vegetation becomeslowerand more wind-pruned fromthewesterntotheeasternend ofthe bay.

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Kilometers

LutregalaMarsh

S.CapeBay

Figure 1.Tasmania, showingthe locationsofthetwo studysites mentioned in thetext.

Thetopography immediatelyinlandfromthe beach consists of a series oftwo or three low

ridges,apparentlystabiliseddunes, separatedby damper swales. The most landward ridge

becomeshigherfromwesttoeast,butattheeast- ern end of the bay erosion has cut into these structures, eliminatingthelowerseawardridges and formingasandcliffapproximately8metres high.The soilsaresandy, butatthe eastern end ofthebaywaterloggedclayeysoilsappear within

30m inland ofthe top ofthesand cliff.

Two transect lines were set up in February

1989, at right angles to the high water mark, from below the extreme high water mark of

spring tides (EHWS) to between 40 and 70 m

inland.At eachmetermarkalongthese transects a pitfall trap was set. The traps consisted ofa plastic container of 5 cm diameter and 10 cm

depth, let into thesoilsothat thelipofthe trap wasflushwiththe surfaceofthe sand,soilorleaf litter. Saturated picric acid solution (5-10 ml) wasplacedinthe traps as a non-volatilepreserv- ativeand Petridishlidsweresupportedontwigs overthe traps to prevent rainwaterfrom enter- ing them. Traps were set for one series often nightsinFebruary andanother of14 daysinJuly 1989.

LutregalaMarsh

LutregalaMarsh issituated onthe west coast ofBruny Island, which isjust offthe southeast coast of Tasmania (Fig. 1). The marsh has a

diverse talitrid fauna, and, importantly,a rela- tivelyundisturbedtransitionfromsaltbushsalt-

marshthroughsalinetussockgrasslandtonative terrestrial vegetation, despite some grazing by cattleon the higher levels. Themarsh occupies approximately 10ha.

Two sampling methods were employed to determine the distribution of amphipods.

Firstly,twotransectsofpitfalltraps,moreorless at right angles to the high water mark, were placed across the marsh, andextendedintoter- restrial vegetation up to 50 m beyond the extreme high water mark. Pitfalls were con- structed inthesame way asat SouthCapeBay.

Transect A was set and retrieved three times between March and October 1990 duringover- nightlow springtides, but Transect B wasonly used once because ofpersistent flooding.

In the second approach to sampling, collec- tions were made extensively over the marsh.

Fromarecent, colouraerialphotograph,collec- tionsiteswere chosenthatcouldeasilybeident- ified onthe ground, and which representedthe range ofvegetation types present. Ateach sam- plingsite, at least 20 amphipods werecollected with anaspirator, and the sameenvironmental variables thatwere collected at each pitfall site

were measured. These collections were also

made between March and October 1990. This period covers both the non-breeding (winter) and early breeding season of the species present.

Results South Cape Bay

Twelvespecies, threeofwhich weresupralit- toral sandhoppers, are listed in Table 1. Two

undescribed speciesof Tasmanorchestia Friend were identified, and none ofthe sandhoppers has beenplacedinaspecifictaxon because ofthe poor taxonomic knowledge ofthisgroup.

The catchesfromthe pitfall traps (Figure 2a- d)showdistinctzonationsinthe distributionof species.Amongthesandhoppers,there isa clear series,with'Talorchestia' species1 foundclosest tothe sea, followed by 'Talorchestia' species2,

which in turn is replaced by 'Talorchestia' species 3. The replacement of sandhoppers ('Talorchestia' species 3) by landhoppers(Aus- trotroidesmaritimusFriend)is verysharp. The overlap was never more than 2 traps (= 2m), and onseveral nights therewasnooverlapatall.

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524 A.M.M. RICHARDSON

Table 1. Talitrid amphipodscollected at South Cape Bay and Lutregala Marsh, with their phylogenetic affinities, ecologicalgroupings and abundance. Bousfield's (1984) groupings are as follows. I:i: palustral talitrids. II:iii: cuspidactylate beachfleas havinga 4-dentate lacinia mobilis on the left mandible. Ilia: sandhoppers havinga 4-dentate lacinia mobilison the left mandible;

III:ii: sandhoppers havinga 5-dentate lacinia mobilison the left mandible. IV:i: simplidactylate landhoppers. IV:ii: cuspidactylatelandhoppers.

Taxon Bousfield Landhopper Abundance: Abundance

Group Ecological at South at (Bousfield Group (Frienc1 Cape Bay Lutregala

1984) 1987)

Eorchestiapalustris Richardson I:i Common

Beachflea ll:iii

Common

Talorchestiasp. 3 III:i Common

Talorchestiasp.1 Ill:ii Common

Talorchestiasp. 2 IIF.ii Common

Austwtwides maritimns Friend IV:i Coastal Common Common

A. longicomis Friend IV:i Westernforest Rare

Keratroides rex Friend IV:i Coastal Scarce Scarce

Mysticotalitrus Iasmaniae(Ruftb) IV:i Eastern forest Common Common

M. ayptusFriend IV:i Eastern forest Scarce Scarce

K. vulgaris(Friend) IV:i Eastern forest Common Scarce

K. angulosus(Friend) IV:i Eastern forest Common

NeorchestiaplicibranchaFriend IV:i Western forest Common

Tasmanorchestia Friend sp.1 IV:ii Coastal Rare

Tasmanorchestia Friend sp.2 IV:ii Coastal Rare Rare

OntransectAtherewasacompletereplacement ofsandhoppersbyA. maritimusfrom trap9to trap 10, with no sandhoppers intrap 10andno

A. maritimus in trap 9 after 14 nightsofcatch- ing. Once beyond the landward range of the sandhoppers, the catch at S. Cape was domi- nated by A. maritimus for the next 20-30 m

inland. Tasmanorchestiasp.andKeratroidesrex also appeared, but in very low numbers, and only on transect B in thesummer series.

Samples collected by hand, during the day, (Table 2) from the four swaleson S. Cape Bay transect Bdidnotcontainthesameproportions ofspecies as thepitfall trapsat all sites. Atthe seawardend, handsamplesfailedtocatchsand- hoppersbut caughtamuchhighernumberofA.

maritimus and K. vulgaris. Further landward, mostspeciesweremuchlesslikelytobetrapped than caught by hand,andonlyA.maritimusand M. tasmaniae were over-represented in the traps.

Furtherinland,beyondthetraplines,theland- hoppercommunity in forestlitterchanges,with notraceofthe coastalgroupspeciesor Mystico- talitrustasmaniae.K.angulosusandNeorchestia plicibrancha become the dominants, with Austwtwides longicomis Friend appearingata

drier site about 1 km from the beach. Further inlandstill,N.plicibranchawastheonlyspecies collected in rainforest several kilometers from the coast.

LutregalaMarsh

Eight species of talitrid amphipods were

identified at Lutregala Marsh. These included onespeciesfromeach ofthe palustralandbeach-

fleagroups, three coastal speciesandthreeforest species(Table3). Oneofthe coastal specieswas an apparently undescribedspeciesofTasmanor- chestia Friend.

Profiles ofthe transects, the distributions of amphipodsandthe levelsoftheenvironmental parameters are shown in Figs 3a and 3b. Both transectscross aslight rise attheirseawardends, butTransectB isclearly muchlower than Tran- sectAandliabletotidalfloodingovermostofits length. At their landward ends both transects riseslightly andpass into terrestrialvegetation.

The most striking contrast in the amphipod fauna is the absence of Eorchestiapalustris on the drierTransect A,whereit isreplaced inthe lowersites bythe beachflea. OnTransect Bthe beachflea ismostly confinedto theupper levels ofthetransect,beyondthe range ofE.palustris.

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t

TRANSLITTORALTALITRIDAEAMPHIPODA INTASMANIA 525

Talorchestia sp1

fe

Talorchestiasp2

3.0 10 20 30

Talorchestiasp3

10 20 30

A Mysticotalitrustasmaniae

10 20 30

3Neorchestiaplicibrancha

A:S CapeA:February

\t

Talorchestia sp1

3.0

10 20 30

Talorchestia sp2

it

10 20 30

Talorchestiasp3

3.0 -\

Austrotroides maritimus

»•

3.0 10 20 30

j Mysticotalitrustasmaniae

3.0"I 10 20 30

Keratroidesvulgaris

•-

Keratroides angulosus

3.0 10 20 30

Neorchestiaplicibrancha

C:S.CapeA: July

3.0

Talorchestiasp1

h:

20 30 40

Talorchestiasp2

3.0

10 20 30 40 50 80

3.0 Austrotroides maritimus

o i

.

20 30

Talorchestiasp3

30 40 50 60

Austrotroidesmaritimus

30 40 50

Mysticotalitrustasmaniae

3.0 1

o

10 20 30

jKeratroides angulosus

ooo » O

40 50 60

Keratroides rex

Keratroidesvulgaris 3.0

o

60 metres

I Tasmanorchestiasp.

Neorchestiaplicibrancha

10 20 30

B: S.CapeB: February

J^^^l Talorchestiasp1

60 metres

60 metres

1

3.0 "

3.0 "

w

10 20 30

^^^^ Talorchestiasp2

40 50 60 70

w*

10 20 30

^^^^ Talorchestiasp3

40 50 60 70

w*

3.0

;

10 20 30

Austrotroidesmaritimus

40 50 60 70

:

3.0 -

10 20 30

Mysticotalitrustasmaniae

40 50 60 70

3.0 '

10 20 30

Keratroidesvulgaris-4

40 50 60 70

Keratroides angulosus <m

3.0"

10 20 30

Neorchestiaplicibrancha

40 50 60 70

•'•* T

10 20 30 40 50 60 70

D: S.CapeB: July

Figure2a-d Catches oftalitridamphipodsinpitfalltrapsatSouthCapeBay,Tasmania.Thehorizontalax, sism

metresfrom Trap1 which wasset attheseaward edge ofthe strandl.ne vegetation; theverticalaxisis log,,,(X + M whereXistheaveragedaily catchof eachspeciesineachpitfall.Thetransectsextended40m(A) or45m(B) inland duringthe February series,but were extendedto 70m inthe July series.

(6)

526 A.M.M. RICHARDSON

Table 2. Catches oftalitrid amphipodsbyhand (first figure) orpitfall (second figure) at or near South Cape Bay (nopitfallswere set inland ofthe 4th swale). Species abbreviations are as follows. Tal 2: Talorchestia sp. 2; Tal 3: Talorchestiasp. 3; Austr mar:Austrotroidesmaritimus;

Kera vulg: Keratroides vulgaris; Keraangu: Keratroidesangulosus; Mysttas: Mysticotalitrus tasmaniae; Myst cryp:Mysticotalitrus crytpus; Neorplic: Neorchestia plicibrancha;Austrlong:

Austrotroides longicornis.

Tal 2

Tal 3

Austr mar

Kera vulg

Kera angu

Myst

tas

Myst cryp

Neor

plic

Austr long

Swale Swale Swale Swale 70 m

1 km

5 km

0/1 3/96 125/1 20/0

— —

0/3 125/4 78/0 6/0 30/0

2/4 128/1 11/0 9/8

— —

40/0 75/2 4/4

— —

1 14

14

3/0 8/0 35/6

6 14

On both transects thereisconsiderableoverlap between the distributions of the beachflea, coastal landhoppers, andforest landhoppers.

Fig4 shows the distribution ofspeciesfrom the four ecological groups ofamphipods that were present (marsh-hoppers, beachfleas, coastaland forestlandhoppers) overthe marsh and the major vegetation types. Eorchestia palustrisisrestrictedtothewetmarshvegetation whereitisregularlyinundated, whilethebeach-

flea extends more deeplyinto the upper marsh and terrestrial vegetation. The forest landhop- pers arerestricted totheterrestrialscrub,but the coastallandhoppersarefound boththereandin the uppermarsh.

Discussion

At South Cape Bay the interface between the beach and the adjacent forest is quite sharp.

There was complete replacement of sandhop- persby landhopperswithin4metres,atthe point wherewoodyvegetationformeda closedcanopy and substantial amounts of organic matter began to appear in the sandy soil. The coastal landhoppers extend inland for another 50 m, beyond which the landhopper community becomes much less diverse than in the coastal zone.Furtherworkat SouthCapeBay andelse-

where in the west of Tasmania (Richardson, 1993, in prep.)shows that the distance that the coastal species penetrate inland is related to exposuretowind, perhaps becausethisincreases theamount ofsalt spray deposited behind the shore.

At Lutregala Marsh, in contrast, there is no obviouspoint where intertidal talitrids (in this

case beachfleas) are replaced by landhoppers, since there are substantial overlaps both between the distributions ofthe beachflea and the coastallandhoppersand betweenthebeach-

fleaandthemarsh-hopperswhich occupyazone closerto the sea.

The numbersoftalitridspecies presentatboth

sitesis high, 11 speciesat South Cape Bay and eight at Lutregala Marsh, and phylogenetically diverse. Although the phylogeny ofthe Talitri- dae is not clearly understood, it is generally accepted (Bousfield, 1984, Moore et al., 1993) that sandhoppers form a distinct, specialised group that is unlikely to have produced any forms ancestraltothe landhoppers. This, com- bined with the absence of beachfleas and the abrupt transition from beach fauna to forest fauna, suggeststhatsandy beacheswereunlikely to have been the sitewhere the colonisation of landbytalitridsbegan.Onthe saltmarshes,both ofthelikelyancestralgroupsare present(marsh- hoppersandbeachfleas),the transitiontolandis

gradual, distributions overlapsubstantiallyand the saltmarsh habitat foreshadows land in the structure and nutritional nature of its

vegetation.

Inboth these examplesthere arereasonswhy

the phenomena ofinterest covera widerrange than "shore" or"land", howeverthey might be defined. Certainly vegetation patterns do not giveanyindicationoftheinlandlimitsofcoastal landhoppers in either location. There hasbeen some interest in the reservation of saltmarsh vegetation in Tasmania (Kirkpatrick &Glasby 1981),butbydefinition thathasreferred tosalt-

marshplants.Therearemanyrecords of beach-

(7)

Mysticotalitrustasmaniae

Mysticotalitruscryptus

Keratroidesvulgaris

10 animals per trapnight

Austrotroides maritimus

Keratroides rex Tasmanorchestia sp.

A2 A4 A6 A8 A10 A12 A14 A16 A18

10 animals per trapnight

Eorchestia palustris

Mysticotalitrus cryptus <

Keratroides rex

Austrotroides maritimus

Tasmanorchestiasp.

Figure 3a, b. Profile oftwo transects on Lutregala Marsh showingthelevelsofsoilmoisture,soilorganic contentandsoilsalinity ateach trappingposition,and therelative abundanceofamphipodstrappedin pit-

falls. MHW: meanhigh waterlevel.

:Mysticotalitrustasmaniae O :Austrotroidesmaritimus

:M.cryptus :Keratroides rex

A:Keratroides rex A : Tasmanorchestia sp.

250m

O:Eorchestiapalustris

:Beachflea

Figure 4a, b. Maps ofLutregala Marsh, showingthe distributions ofthe four ecologicalgroups oftalitrid amphipods. A: marsh-hoppers and beachfleas; B:

coastal and forestlandhoppers. Thedashedline rep- resents the approximate boundary of woodland.

Sampleswerecollectedby hand,and from thepitfall transects(A: startoftransect A:B: startoftransectB).

Filledsymbolsrefertospeciesoftheforestlandhopper group, or the beachflea, open symbolsto the coastal landhopper grouportothepalustralgroup(Eorchestia palustris).

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