Translittoral Talitridae (Crustacea: Amphipoda) and the need to reserve transitional habitats: examples from Tasmanian saltmarshes and other coastal sites

9 Pages·1997·2.8 MB·English

by  A M M Richardson

Memoirs ofthe Museum ofVictoria 56(2):52l-529 (1997)
TRANSLITTORAL TALITRIDAE (CRUSTACEA: AMPHIPODA) AND THE NEED
TO RESERVE TRANSITIONAL HABITATS: EXAMPLES FROM TASMANIAN
SALTMARSHES AND OTHER COASTAL SITES
Alastair M.M. Richardson, Roy Swain and Vera Wong1
Department ofZoology, University ofTasmania, GPO Box 252C, Hobart, Tas. 7001, Australia.
'Present address: 8 Cedex Ct, Glen Waverley, Vic. 3150, Australia.
Abstract
Richardson, A.M.M., Swain, R. and Wong, V., 1997. Translittoral Talitridae (Crustacea:
Amphipoda) and the need toreservetransitional habitats: examplesfromTasmanian salt-
marshes and other coastal sites. Memoirs ofthe Museum ofVictoria 56(2): 521-529.
Transitionalhabitats,e.g.,thosethatliebetweenseaandland,areoftenignoredbecauseof
thedifferent backgroundsofmarineandterrestrialbiologists,buttheyareimportantforthe
seriesofspecieswhichspantheboundary.Theamphipod familyTalitridaehasperhapsthe
widesttranslittoral rangeand soprovidesevidencefrom whichtheevolutionaryprocessof
land colonisation can be deduced. The talitrids found on two sea-land transitions in Tas-
mania,asandy-beachtorainforestandasaltmarshtodrywoodland,aredescribed,andthe
significance oftheir distributions is discussed. The need to reserve the full range ofsuch
translittoral series is noted. This will not necessarily be achieved by establishing reserve
boundaries on the basis ofvegetation associations.
Introduction marinetofullyterrestrial.Thebestexamplesare
the amphipod crustacean family Talitridae and
Transitionalhabitats,thatisthoselyingbetween thepulmonatemolluscfamilyEllobiidae.Trans-
major habitats (land and sea, freshwater and littoralgroupsofthistypeprovideveryvaluable
land) tend to fall through the net ofbiological opportunities to reconstruct the evolutionary
investigations (Little, 1990). Marine biologists events leading to life on land (Little, 1989,
afenrdentterbreasctkrigarlobuinodlsogitshtastdtehveeylohpavferolmittsluecthodidfo- t1h9e90)s,paecnidestheiynbveolcvoemdemaorererveallautaibvelleysticlllowsheleyn-
with each other and so the zones ofinteraction related, as in the two families mentioned
between their spheres of operation tend to be above.
ignored. While marine biologists often work in In this contribution we will present evidence
the intertidal zone, their studiestend to be con- of translittoral distributions in the Talitridae
centrated well belowthe high watermark, while and demonstratethat such distributionsrequire
fewterrestrialbiologistsapproachthehighwater a broader approach to reserve design than has
markfrom theotherdirection.Thustheamount been adopted hitherto.
ofstudy on the intermediate zone is small, and
thisisparticularlyobviouswhenthetransitional
zone is extensive, as it is in the most sheltered Talitridae
sea-land interfaces, in saltmarshes and man- The Talitridae include the only fully terrestrial
groves. members of the Amphipoda, one of only two
And yet the interface between the two major crustacean groups (the other is the Isopoda)
habitats is in many ways more important than which havemembersthatarefullyterrestrial, ie.
the sea or land alone. The colonisation ofland independent of free water for all of their life
was one of the greatest evolutionary events in stages. The Talitridae have been loosely classi-
the history of those seven phyla that have fied by Bousfield (1984) into four groups, the
achievedterrestriality.Althoughmostofthecol- palustral talitrids (marsh-hoppers), the beach-
onisation events happened a long time ago, the fleas,thesandhoppersandthelandhoppers.The
habitats in which they occurred are still there, marsh-hoppers include the most plesiomorphic
and in some cases species can be found filling forms and they are found in the lower levels of
many ofthe intermediate stages. In a few taxa, saltmarshesand mangroves,wheretheyhavean
which might be called translittoral (Chester, amphibious lifestyle, entering and swimming
b1u9t9i2)o,nssersipeasnonfinspgectiheesceantnirbee froaunnged wfirtohmdifsutlrli-y epflfaicceisen(tlTyasimnasneiaaw:atCehresetveern, a1t9l92o,wStioduet.hInAfsroicmae:
521
522 A.M.M. RICHARDSON
Griffiths, 1976) their distributions extend into nents,contemporarywithanextensiveradiation
fully freshwaterstreams. In south east Australia ofthe angiosperm flora in Gondwana.
the genus Eorchestia is widespread in salt- Tasmaniahasaremarkably rich talitrid fauna
marshes and the lower reaches of associated (Richardson, 1993, 1996) which includes rep-
streams(Richardson, 993). Thebeachfleasand resentatives of all Bousfield's groupings. The
1
sandhoppersarethetypicaltalitridsofthe inter- taxonomy of the landhoppers is well-known
tidal zone, distinguished by the adaptations of (Friend, 1987)andthediverseintertidalfaunais
sandhoppersfortheirparticularhabitat, ieloose currently being described by the senior author.
sand. Thus the sandhoppers are large, robust Friend (1987) divided the Tasmanian landhop-
and spiny talitrids, sometimes with unusual perfauna intothreeecological groupings, oneof
morphologicaladaptationsforlifeinsand. They which he called the coastal group. Five or six
are efficient burrowers. The beachfleas, in con- species in three genera make up this group and
trast, arc not well-developed forburrowing, pre- they have distributions which are strongly
m
ferring to seek refuge under cast seaweed or restricted to a narrow zone, less than 100
stones (non substrate modifiers: Macintyre, wide, immediately behind the high tide mark.
1963, or nestlers: Griffiths, 1976); they are gen- Tasmania also retains some places, especially
erally more lightly built and weakly spined. in the west, where transitions from land to sea
Beachfleasareusuallyfoundon rockyormuddy remain undisturbed, and clothed with native
shores, but there is at least one undescribed vegetation. Theseprovideimportantopportuni-
species in Tasmania which has become conver- ties to study the natural distributions oftalitrid
gent with the sandhoppers and forages on mid- amphipods, isopodsand othergroupsacrossthe
tidal wetsand duringlowtide, takingrefugeina transition. We will consider two cases, a sandy
shallow burrow during high tide. beach to wet sclerophyll forest at South Cape
The landhoppers are found in a range ofter- Bay and a saltmarsh to dry woodland transition
restrial habitats in the Southern Hemisphere, at Lutregala Marsh on Bruny Island. Details of
Japan, the Phillipines, Central America, the these studies are described elsewhere (South
Caribbean and many oceanic islands. In Tas- Cape Bay: Richardson et al., 1991; Lutregala
maniatheyarcfound throughoutthe island, but Marsh: Richardson and Mulcahy, in press).
wheretheyhavecolonisedcontinents(Australia,
southern Africa) they are confined to the higher
Study areas and methods
rainfall zones around the continental margins.
Bousficld (1984) recognises two subgroups, the South Cape Bay
plesiomorphic simplidactylate genera, which SouthCapeBayisintheextremesouthofTas-
seem tobe descended from a marsh-hopperlike mania (Fig 1). It faces almost due south, but is
ancestor, and the cuspidactylate genera which sheltered fromtheprevailingsouthwesterlyand
apparently arose from the beachfleas. The sim- westerly winds by promontories to the west.
plidactylate formsareconfined toAustralia and Because ofstrong wave action, the sand above
southern Africa. thehighwatermark iscolonisedbyfewforedune
Thus the Talitridae show a truly translittoral plants, so that low coastal scrub and forest
range from the semi-aquatic marsh-hoppers to appears within a few metres of high water.
thelandhoppers. ThesisterfamilytotheTalitri- Closest to the shorethe shrubs Correa backhou-
dae is the Hyalidae, and it is widely- accepted siana, Cyathodesabietina, Westringia brevifolia
thattalitridsdescendedfromahyalid-likeances- and Olearia phlogopappa form a low dense
tor (Bousficld, 1984). Aquatic hyalids may be cover, five to ten metres deep, which gives way
found adjacent to intertidal talitrids in fully to low forest with emergent trees ofEucalyptus
marine and brackish waters, providing a valid nitida, sassafras,Athewsperma moschatum, cel-
extension to the translittoral series. ery top pine, Phyllocladus aspleniijblius, and
Itseemslikelythatthetalitridamphipodscol- southern beech, Nothofaguscunninghamii. This
onised land relatively recently compared to coastal forest has an understorey of native
other terrestrial arthropods. Little (1990) noted laurel, Anopterusglandulosus, and often cutting
that the fossil record ofthe amphipods is only grass, Gahnia grandis. The ground layer is
from the Quaternary, but Bousfield (1984) and mostly absent, apart from a few mosses and
Bousficld and Poinar (1995) suggests a mid- herbs. The vegetation becomes lower and more
Cretaceous origin for the simplidactylate land- wind-prunedfromthewesterntotheeasternend
hoppers of the Southern Hemisphere conti- ofthe bay.
TRANSLITTORAL TALITRIDAE AMPHIPODA IN TASMANIA 523
Lutregala Marsh
Lutregala Marsh is situated on the west coast
ofBruny Island, which isjust offthe south east
coast of Tasmania (Fig. 1). The marsh has a
diverse talitrid fauna, and, importantly, a rela-
tively undisturbed transition from saltbush salt-
marshthroughsalinetussockgrassland tonative
terrestrial vegetation, despite some grazing by
cattle on the higher levels. The marsh occupies
approximately 10 ha.
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, and extended into ter-
restrial vegetation up to 50 m beyond the
extreme high water mark. Pitfalls were con-
structed in the same way as at South Cape Bay.
Transect A was set and retrieved three times
between March and October 1990 during over-
night low spring tides, but Transect B was only
LutregalaMarsh used once because ofpersistent flooding.
Kilometers
S.CapeBay In the second approach to sampling, collec-
tions were made extensively over the marsh.
From a recent, colouraerial photograph, collec-
Figure 1. Tasmania, showingthe locations ofthe two tion siteswerechosen thatcouldeasilybeident-
study sites mentioned in the text. ified on the ground, and which represented the
range ofvegetation types present. At each sam-
pling site, at least 20 amphipods were collected
The topography immediately inland from the with an aspirator, and the same environmental
beach consists of a series of two or three low variables that were collected at each pitfall site
ridges,apparentlystabiliseddunes,separatedby were measured. These collections were also
damper swales. The most landward ridge made between March and October 1990. This
becomeshigherfromwesttoeast,butattheeast- period covers both the non-breeding (winter)
ern end of the bay erosion has cut into these and early breeding season of the species
structures, eliminatingthe lowerseaward ridges present.
and forminga sand cliffapproximately 8 metres Results
high. The soils are sandy, but at theeastern end
ofthebaywaterloggedclayeysoilsappearwithin South Cape Bay
30m inland ofthe top ofthe sand cliff. Twelve species, three ofwhich were supralit-
Two transect lines were set up in February toral sandhoppers, are listed in Table 1. Two
1989, at right angles to the high water mark, undescribed species ofTasmanorchestia Friend
from below the extreme high water mark of were identified, and none of the sandhoppers
spring tides (EHWS) to between 40 and 70 m hasbeenplaced inaspecifictaxonbecauseofthe
inland. Ateach metermarkalongthesetransects poor taxonomic knowledge ofthis group.
a pitfall trap was set. The traps consisted of a The catches from the pitfall traps (Figure 2a-
plastic container of 5 cm diameter and 10 cm d)show distinct zonations in the distribution of
wdeapsthfl,uslehtwiintthotthheessuoriflascoetohfatthethseanlidp,osfoitlhoertlreaapf ssepreiceise,s.wiAtmho'Tnaglotrhcehessatnida'hospppeecriess,1thfeoruenidscalcolseeastr
litter. Saturated picric acid solution (5-10 ml) to the sea, followed by 'Talorchestia' species 2,
wasplaced in thetrapsasa non-volatilepreserv- which in turn is replaced by 'Talorchestia'
ativeand Petri dishlidsweresupportedontwigs species 3. The replacement of sandhoppers
over the traps to prevent rainwater from enter- ('Talorchestia' species 3) by landhoppers (Aus-
innigghttsheimn.FeTbrraupasrywaernedasnetotfhoerroonfe14sedraiyessionfJtuelny aotrvnoedtrrlooanipdseewsvamesraarnlietnviiegmrhutsmsoFtrrheeireentdhw)aansis2nvoetroryavpessrhla(ar=pp.a2tTmah)lle,.
1989.
524 A.M.M. RICHARDSON
Table 1. Talitrid amphipods collected at South Cape Bay and Lutregala Marsh, with their
phylogenetic affinities, ecological groupings and abundance. Bousfield's (1984) groupings are as
follows. I:i: palustral talitrids. II:iii: cuspidactylate beachfleas having a 4-dentate lacinia mobilis
on the left mandible. Ilia: sandhoppers having a 4-dentate lacinia mobilis on the left mandible;
III:ii: sandhoppers having a 5-dentate lacinia mobilis on the left mandible. IV:i: simplidactylate
landhoppers. IV:ii: cuspidactylate landhoppers.
Taxon Bousfield Landhopper Abundance Abundance
:
Group Ecological at South at
(Bousfield Group (Frienc Cape Bay Lutregala
1
1984) 1987)
Eorchestiapalustris Richardson I:i — Common
Beachflea ll:iii —Common
Talorchestia sp. 3 III:i Common —
Talorchestia sp.1 Ill:ii Common —
Talorchestia sp. 2 IIF.ii Common
Austwtwides maritimns Friend IV:i Coastal Common —Common
A. longicomis Friend IV:i Western forest Rare
Keratroides rex Friend IV:i Coastal Scarce Scarce
Mysticotalitrus Iasmaniae (Ruftb) IV:i Eastern forest Common Common
M. ayptus Friend IV:i Eastern forest Scarce Scarce
K. vulgaris (Friend) IV:i Eastern forest Common —Scarce
K. angulosus (Friend) IV:i Eastern forest Common —
Neorchestiaplicibrancha Friend IV:i Western forest Common
Tasmanorchestia Friend sp.1 IV:ii Coastal Rare
Tasmanorchestia Friend sp.2 IV:ii Coastal Rare Rare
On transectAtherewasacompletereplacement drier site about 1 km from the beach. Further
ofsandhoppers by A. maritimus from trap 9 to inland still, N. plicibranchawas the only species
trap 10, with no sandhoppers in trap 10 and no collected in rainforest several kilometers from
A. maritimus in trap 9 after 14 nights ofcatch- the coast.
ing. Once beyond the landward range of the
sandhoppers, the catch at S. Cape was domi- Lutregala Marsh
nated by A. maritimus for the next 20-30 m Eight species of talitrid amphipods were
inland. Tasmanorchestiasp.andKeratroidesrex identified at Lutregala Marsh. These included
also appeared, but in very low numbers, and onespeciesfromeachofthepalustralandbeach-
only on transect B in the summer series. fleagroups,threecoastalspeciesandthreeforest
Samples collected by hand, during the day, species (Table 3). Oneofthe coastal species was
(Table 2) from the four swales on S. Cape Bay anapparentlyundescribedspeciesofTasmanor-
transect B did not contain the sameproportions chestia Friend.
ofspecies as the pitfall traps at all sites. At the Profiles of the transects, the distributions of
seaward end, hand samples failed tocatch sand- amphipods and the levels ofthe environmental
hoppers but caught a much highernumberofA. parameters are shown in Figs 3a and 3b. Both
maritimus and K. vulgaris. Further landward, transectscrossaslightriseattheirseawardends,
most species weremuch lesslikelytobetrapped but Transect B isclearly much lowerthan Tran-
thancaught by hand, and onlyA. maritimusand sectAand liabletotidalfloodingovermostofits
M. tasmaniae were over-represented in the length. At their landward ends both transects
traps. rise slightly and pass into terrestrial vegetation.
Furtherinland,beyondthetraplines,theland- The most striking contrast in the amphipod
hoppercommunity in forest litterchanges, with fauna is the absence of Eorchestia palustris on
notrace ofthecoastal group speciesorMystico- the drierTransect A, where it is replaced in the
talitrustasmaniae.K. angulosusandNeorchestia lower sites by the beachflea. On Transect B the
plicibrancha become the dominants, with beachflea is mostly confined to the upper levels
Austwtwides longicomis Friend appearing at a ofthetransect, beyond the range ofE. palustris.
^
—
TRANSLITTORAL TALITRIDAE AMPHIPODA IN TASMANIA 525
t Talorchestiasp1 3.0 Talorchestiasp1
20 30 40
Talorchestiasp2
h: Talorchestiasp2
fe
3.0 10 20 30 20 30
Talorchestiasp3 3.0 Talorchestiasp3
10 20 30 40 50 80
30 40 50 60
3.0 Austrotroidesmaritimus
Austrotroidesmaritimus
o i
.
10 20 30 30 40 50
A Mysticotalitrustasmaniae Mysticotalitrustasmaniae
3.0 1 10 20 —30 40 50 60
jKeratroidesoaonogul»osuOs Keratroidesrex
o
Keratroidesvulgaris
10 20 30
3.0
3 Neorchestiaplicibrancha I Tasmanorch^est»iasp.
o
Neorchestiaplicibrancha
6m0etres 10 20 30 60
metres
A:SCapeA: February B:wS.CapeB: February
J^^^l Talorchestiasp1
Talorchestiasp1
\t 1
10 20 30 10 20 30 40 50 60 70
3.0 3.0 "
Talorchestiasp2 w^^^*^ Talorchestiasp2
10 20 30 10 20 30 40 50 60 70
Talorchestiasp3 3.0 "w^^^*^ Talorchestiasp3
it
10 20 30 40 50 60 70
3.0 -\Austrotroidesmaritimus»• 3.0 ;Austrotroidesmaritimus
:
3.0jMystico1t0alitrust2a0smaniae30 3.0 -Mystic1o0talitrus20tasman3i0ae 40 50 60 70
3.0"IKeratroi10desvul•g2a-0ris— 30 3.0 'Keratr1o0idesvu2l0garis -340 40 50 60 70
Keratroidesangulosus
Keratroidesangulosus <m
3.0 10 20 30 10 20 30 40 50 60 70
Neorchestiaplicibrancha 3.0"
Neorchestiaplicibrancha
•'•* T
60
metres 10 20 30 40 50 60 70
C:S.CapeA:July D:S.CapeB:July
m
iFmMnielgtawurnrhedeesr2dfeaur-roXdimniCTgsartttahhcepehae1Fvseweobhrfraiugtcaaehlridwytaariissldeyrsaiecemtaspt,ahctbhituphotoefdwesesaercaiehwnapesripxtdetfcaeeilndledgsteeridanopftesotaahct7eh0SspotmiurtftaaihnlndlC.lta.ThnpeheeeJvBuetalgryyea,tnsasTeteraicisotenmss;.aentxihteae.nvedTrehtdiec4ah0loramixzi(osAni)tsaollorga,4x,,,5s(mXis(B+)
526 A.M.M. RICHARDSON
Table 2. Catches oftalitrid amphipods by hand (first figure) or pitfall (second figure) at or near
South Cape Bay (no pitfalls were set inland ofthe 4th swale). Species abbreviations are as
follows. Tal 2: Talorchestia sp. 2; Tal 3: Talorchestia sp. 3; Austr mar: Austrotroides maritimus;
Kera vulg: Keratroides vulgaris; Kera angu: Keratroides angulosus; Myst tas: Mysticotalitrus
tasmaniae; Myst cryp: Mysticotalitrus crytpus; Neor plic: Neorchestia plicibrancha; Austr long:
Austrotroides longicornis.
Tal Tal Austr Kera Kera Myst Myst Neor Austr
2 3 mar vulg angu tas cryp plic long
— —
Swale 0/1 3/96 125/1 20/0
Swale 0—/3 125/4 78/0 6/0 30/0 3/0
Swale — 2—/4 128/1 11/0 9/8 8/0
Swale — — 40/0 75/2 4—/4 35/6
m
70 1 14 6
km
1 14 14
km
5
On both transects there is considerable overlap case beachfleas) are replaced by landhoppers,
between the distributions of the beachflea, since there are substantial overlaps both
coastal landhoppers, and forest landhoppers. between the distributions of the beachflea and
Fig 4 shows the distribution of species from thecoastal landhoppers and between thebeach-
the four ecological groups of amphipods that fleaandthemarsh-hopperswhichoccupyazone
were present (marsh-hoppers, beachfleas, closer to the sea.
coastal and forest landhoppers) over the marsh Thenumbersoftalitridspeciespresentatboth
paanldusttrhiesismraejstorrictveedgteotatthieownettmypaerss.hEvoergcehteasttioina esiitgehstiasthiLguht,re1g1alsapeMcairessha,t aSnoudthphyClaopgeenBetaiycaalnldy
where it is regularly inundated, whilethebeach- diverse. Although the phylogeny ofthe Talitri-
aflneda etxertreensdtsriamlorveegedteaetpiloyn.inTthoethfeoreusptpelranmdahrosph- adcaceepitsedno(tBoculsefairellyd,un1d9e8r4s,toMoodo,reit eitsagl.e,ne1r9a9l3l)y
persarerestrictedtotheterrestrialscrub,butthe that sandhoppers form a distinct, specialised
coastal landhoppersarefound both thereand in group that is unlikely to have produced any
the upper marsh.
forms ancestral to the landhoppers. This, com-
bined with the absence of beachfleas and the
Discussion
abrupt transition from beach fauna to forest
At South Cape Bay the interface between the fauna, suggeststhat sandybeacheswereunlikely
beach and the adjacent forest is quite sharp. to have been the site where the colonisation of
There was complete replacement of sandhop- landbytalitridsbegan. Onthesaltmarshes,both
persbylandhopperswithin4metres,atthepoint ofthelikelyancestralgroupsarepresent(marsh-
wherewoodyvegetationformedaclosedcanopy hoppersandbeachfleas),thetransition toland is
and substantial amounts of organic matter gradual, distributions overlap substantially and
began to appear in the sandy soil. The coastal the saltmarsh habitat foreshadows land in the
landhoppers extend inland for another 50 m, structure and nutritional nature of its
beyond which the landhopper community vegetation.
becomes much less diverse than in the coastal In both these examples there are reasons why
zone. Furtherwork at South Cape Bay and else- the phenomena ofinterest cover a wider range
where in the west of Tasmania (Richardson, than "shore" or "land", however they might be
1993, in prep.) shows that the distance that the defined. Certainly vegetation patterns do not
coastal species penetrate inland is related to giveanyindicationoftheinland limitsofcoastal
exposuretowind,perhapsbecausethisincreases landhoppers in either location. There has been
the amount of salt spray deposited behind the some interest in the reservation of saltmarsh
shore. vegetation in Tasmania (Kirkpatrick & Glasby
At Lutregala Marsh, in contrast, there is no 1981), but bydefinition thathasreferredtosalt-
obvious point where intertidal talitrids (in this marsh plants. Thereare many records ofbeach-
TRANSLITTORAL TALITRIDAE AMPHIPODA IN TASMANIA 527
Mysticotalitrustasmaniae
Mysticotalitruscryptus
Keratroidesvulgaris
10animalspertrapnight
Austrotroidesmaritimus
Keratroidesrex
Tasmanorchestia sp.
•: Mysticotalitrustasmaniae O :Austrotroidesmaritimus
:M.cryptus : Keratroidesrex
A: Keratroidesrex A : Tasmanorchestia sp.
250m
A2 A4 A6 A8 A10 A12 A14 A16 A18
Mysticotalitruscryptus <
Keratroidesrex
10animalspertrapnight Austrotroidesmaritimus —
Tasmanorchestia sp.
Eorchestiapalustris
O Eorchestiapalustris
:
• Beachflea
:
Figure 4a, b. Maps ofLutregala Marsh, showing the
distributions ofthe four ecological groups oftalitrid
amphipods. A: marsh-hoppers and beachfleas; B:
coastal and forest landhoppers. The dashed line rep-
resents the approximate boundary of woodland.
Samples were collected by hand, and from the pitfall
FMiagrusrhe s3ah,owbi.ngPrtohfeilleevoelfstowfosoitlrmaonsiescttusreo,nsoiLluotrrgeagnailca tFrilalnesdecstysm(bAo:lsstarretfeorfttorasnpseeccitesAo:fBt:hestfaorrteostfltraanndsheoctppBe)r.
tchoentreenltatainvde asobiulnsdalainnciteyoatfeaamcphhtirpaopdpsintgrpaopspietdioni,napintd- glraonudph,opopretrhgerboeuapchofrlteoa,thoeppeanlusstyrmabloglrsoutpo(tEhoercchoeassttiala
falls. MHW: mean high water level. palustris).
528 A.M.M. RICHARDSON
fleas extending their range into terrestrial veg- etal., 1993, Morritt, 1987, 1988, 1989; Spiceret
etation, usually in situationswhere there iscon- al.. 1987; Spicer and Taylor, 1987, 1994), and
siderablesaltspray(egReid 1947,Bagenal 1957, Tasmania provides perhaps the best oppor-
Kuhnelt 1976, Bousfield 1982), and the coastal tunity in the world to combine ecophysiology
landhoppers, which are part ofthe translittoral with studiesofdistributionsandotheraspectsof
series, extend beyond the landward limit ofthe thebiologyofatranslittoralseries,providedthat
recognised saltmarsh plants. places where that can be done remain intact.
In a survey of the crustacean and mollusc
fauna of Tasmanian saltmarshes, Wong et al. Acknowledgements
(1993) recorded the factors causingdisturbance
at 52saltmarshesand 1 1 otherbrackish marshes Weacknowledge theTasmanian Department of
aroundtheTasmanian coast. Aswell asgrazing, Environment and Land Managaement for per-
invasion by exotic species, landfill, catchment mission to work within the Western Tasmania
modification, fire and off-road vehicles, all World Heritage Area, and Julian Bush and the
noted by Kirkpatrick and Glasby (1981), they Tasmanian Conservation Trust for access to
also recorded the condition of the vegetation Lutregala Marsh. Several pepople helped in the
immediately behind the marshes. Very few of field; thanks to Andrew Cawthorne, Stefan
the marshes retained intact backing vegetation, Eberhard, Maria Moore, Mark Nelson, Caron
since land clearance for grazing behind the Summers and Jerry Tupacz.
marshesisverycommon (indeed, grazing in the
marshesisoneofthemostcommon disturbance
factors). Wong et al. (1993) recommended nine References
marshes for listing on the Register of the Bagenal, T. B., 1957. The vertical range ofsome lit-
National Estate and added another three toralanimalson St. Kilda. ScottishNaturalist69:
marshes of high conservation value, but even 50-51.
amongst this group, only five were recorded as Bousfield,E.L., 1982.TheamphipodsuperfamilyTal-
hmaavyinbgeirnetlaacttivbealcykeiansgyvteogestealteicotns.aTlthmuasrswhheislefoirt eiFctaormloioilgdyeya.TaNilnaitttirhiodenaaeNl:oMrsytushsteeeamsuatmtesircnsofaPnaNcdaitfduiircsatlrreiSgbciuiotenin.ocnea1,l.
reservation on the basis of values within the Ottawa, Publications in BiologicalOceanography
marsh as defined by the vegetation, there are II: 1-73.
very few opportunities to reserve the complete Bousfield, E.L., 1984. Recentadvancesinthesystem-
series of distributions of the translittoral atics and biogeography oflandhoppers (Amphi-
talitrids. poda: Talitridae) ofthe lndo-Pacific region. Pp.
Fortunately, the situation with sandybeaches 171-210 in: Radovsky F.J.. Raven, P.H. and
and their backing vegetation is not so bad. at Sohmer, S.H. (eds), Biogeography ofthe tropical
Pacific.AssociationofSystematicCollectionsand
mlmeauansctihaionWfottrhhleedwcHeoeasrstitetralnigeeshAawrlieftaho.ifnElttshheeewhiWeselrsaent,de,irnnwlhoTewarese-r BousfttirheiealldB,aermEn.pLih.cieapnPo.ddBPfiorisonhamorp,TeMGru.tOsi.aeJru.y,m:a1m9Hb9oe5n.roNldueelpwuo.stietrsreos-f
rainfallareasthedunesthatbackthebeachesare the Dominican Republic. Journal ofCrustacean
oftencolonised bytheintroducedmarramgrass, Biology 15: 746-755.
Ammophila arenaria, but in another survey Chester, E.T., 1992. The life history ofthe palustral
(Richardsonetal.,inprep.),afewbeachesonthe talitrid Eorchestia palustris (Crustacea: Amphi-
eastcoast havebeen identified wherethecoastal poda: Talitridae). Unpublished Honours thesis.
vegetation remains undisturbed. University ofTasmania. 165 pp.
vatItioins rdeegcriestitoanbsles,otohfotuegnhhapvraegtmoatbiecm,atdhaetornestehre- Frienzpdoo,odagJ.:eAo.gT,arl1ai9pt8hr7yi..daTRehe)ecootrfedrTrseaosstfmrtiahaneliAaaum:spthsriyaslptioeadmnsaMt(uiAcsmsepuahmni,d-
basis ofvegetation patterns, without any refer- Supplement 7: 1-85.
encetoanimal distributions. While it isnot sur- Griffiths, C.L., 1976. Guide to the henthic marine
prising that there is often no clear correlation amphipods of southern Africa. South African
between animal distributions and plant associ- Museum: Cape Town. 106 pp.
ations, in some cases, such as the translittoral Kirkpatrick,J.B. andGlasby,J., 1981.Saltmarshesin
groups discussed here, defining reserves in veg- Tasmania. Occasional Paper Number 8. Depart-
etational terms can lead tothe failuretoprotect ment ofGeography, University ofTasmania. 62
ssiigdneirfaibclaentinztoeorleosgticianltphheenecoompehnyas.ioTlhoegryeoifslcaonnd- Kuhneptlph.et.AnK.i,m1a9l76K.inSogidlobmi.olFoagybewritahnsdpeFcaibalerr:efLeroenndcoento
colonisation in talitrid amphipods (e.g., Moore 483 pp.
.
TRANSLITTORALTALITRIDAE AMPH1PODA IN TASMANIA 529
Little, C, 1989. Comparative physiology asatool for manianTalitridae.JournalofCrustaceanBiologv
investigating the evolutionary routes ofanimals 16: 574-583.
onto land. Transactions ofthe Royal Society of Richardson,A.M.M.andMulcahy,M.E.,inpress.The
Edinburgh: Earth Sciences 80: 201-208. distributionoftalitridamphipods(Crustacea)on
Little, C, 1990. The terrestrial invasion. Cambridge a saltmarsh in southern Tasmania, in relation to
University Press: Cambridge. 304 pp. vegetation and substrate. Estuarine, Coastaland
Moore,P.G., Morritt,D.andStevenson,T.D.I., 1993. ShelfScience.
Morphology ofthe antennary gland exit duct in Richardson,A.M.M.,Swain.R.AndSmith,S.J., 1991
ecological and phylogenetic series of talitroid Local distributionsofsandhoppersand landhop-
Amphipoda(Crustacea).JournalofZoology 231: pers (Crustacea: Amphipoda: Talitridae) in the
187-201. coastalzoneofwesternTasmania. Hydrobiologia
Morritt,D., 1987. Evaporativewaterlossunderdesic- 223: 127-140.
cation stress in semiterrestrial and terrestrial Richardson, A.M.M., Swain, R. and Shepherd. C. (in
amphipods (Crustacea: Amphipoda: Talitridae). prep.)Asurveyofthefaunaoftheupperregionsof
Journal of Experimental Marine Biology and sandy beaches on the east coast of Tasmania.
Ecology 111: 145-157. Report to the National Estate Grants Program.
Morritt, D., 1988. Osmoregulation in littoral and ter- Spicer,J.I. and Taylor, A.C., 1987, Respiration in air
restrial talitroidean amphipods (Crustacea) from and water of some semi- and fully terrestrial
Britain.,JournalofExperimentalMarineBiology amphipods (Crustacea: Amphipoda). Journal of
andEcology 123: 77-94. Experimental Marine Biology and Ecology 106:
Morritt, D., 1989. Ionic regulation in littoral and ter- 265-277.
restrialamphipods(Crustacea:Amphipoda:Tali- Spicer, J.I. and Taylor, A.C., 1994, The origin and
tridae), JournalofExperimentalMarine Biology metabolicsignificanceofexosomaticwaterinthe
andEcology 132: 53-67. semi- terrestrial beachflea, Orchestia gammarel-
Reid, D.M., 1947.Talitridae(CrustaceaAmphipoda). lus (Crustacea: Amphipoda). Journal ofZoology
LinnaeanSocietySynopsesoftheBritishEauna7: 232, 617-632.
1-25. Spicer, J.I., Moore, P.O. and Taylor, A.C., 1987, The
Richardson, A.M.M., 1993. The distribution of physiologicalecologyofland invasion bytheTal-
coastal landhoppers (Crustacea: Amphipoda: itridae (Crustacea: Amphipoda). Proceedings of
Talitridae) related to sea spray at Cox Bight, theRoyalSociety ofLondon (Series B), 232: 95-
Tasmania. Papers and Proceedings ofthe Royal 124.
Society ofTasmania 127: 55-60. Wong, V., Richardson,A. M. M.andSwain,R., 1993.
Richardson, A.M.M., 1996. A new species ofProtor- ThecrustaceansandmolluscsofTasmanian salt-
chestiaBousfield(Crustacea:Amphipoda:Talitri- marshes.UniversityofTasmania: Hobart. Report
dae) from Maatsuyker Island, Tasmania, with a to the National Estate Grants Program. 95 pp.
keytothegenusandnotesonthediversityofTas-