Distribution patterns and diversity of invertebrates of temperate rainforests in Tasmania with a focus on Pauropoda

2008·5.8 MB·English

by  Penelope Greenslade

MemoirsofMuseumVictoria65: 153-164(2008)
ISSN 1447-2546(Print) 1447-2554(On-line)
http://www.museumvictoria.com.au/About/Books-and-Journals/Journals/Memoirs-of-Museum-Victoria
Distribution patterns and diversity ofinvertebrates oftemperate rainforests in
Tasmania with a focus on Pauropoda
PenelopeGreenslade
SchoolofBotanyandZoology,AustralianNationalUniversity,GPOBox,ACT,0200,Australia;CentreforEnvironmental
Management, School of Science and Engineering, University of Ballarat, PO Box 663, Ballarat, Victoria, Australia
([email protected])
Abstract Greenslade,P.2008.DistributionpatternsanddiversityofinvertebratesoftemperaterainforestsinTasmaniawithafocus
onPauropoda.MemoirsofMuseumVictoria65: 153-164.
Pauropodaarerecordedforthe firsttimefromTasmania. Nineteen species arelistedfrom alarge collectionof
specimensmadeduringanintensivesurveyoftemperaterainforestinTasmania.Akeyisprovidedfortheidentificationof
Tasmanianspecieswhicharerecordedbyvouchernumberandthebiogeographicalaffinitiesofthefaunaarediscussed.
Taxic richness ofPauropodainTasmanianrainforests is comparedwiththatofseveral othergroups oflitterandbark
invertebrates fromthe same collections.All groups show similarpatterns ofhightaxic richness inthe northwest and
southeast regions with lowerrichness at higher altitudes. The implications forthe conservation ofTasmania’s highly
endemicinvertebratefaunaisdiscussed.
Keywords Acarina,Amphipoda,Diplopoda,Isopoda,Opiliones,Symphyla,regionaldiversity,conservationpriorities.
Introduction foundinthecentralAmazonforestwhereover50specieswere
recordedfromasinglevegetationtype(Scheller, 1994, 1997).
The Pauropoda is a Class ofminuteArthropoda belonging to The Australian fauna is estimated to comprise at least 1000
the Subphylum, Myriapoda. It comprises around700 hundred species ofwhich only 18 species (2%) have been recorded so
speciesworldwidethataregroupedintofivefamilies.Thetotal far.PreviousrecordsarefromQueensland(Remy, 1959),New
worldfaunaislikelytobeintheregionof5000speciesasmany South Wales (Harrison, 1914), Victoria (Tiegs, 1943, 1947;
regions and ecosystems are under-collected (Scheller, 1990). Remy, 1949),NorthernTerritory(GreensladeandMott, 1979)
Adults have eight to eleven pairs of legs, lack eyes and are and Western Australia (Remy, 1957; Postle et al., 1991). A
either whimtemor lightly sclerotised with a maximum length of checklist forAustralia is available (ABRS, ABIF, 2002). No
about1.5 (Scheller, 1990).Pauropodatendtobesoil-living species of pauropod have been recorded previously for
butare alsofoundinrotting logs, underbarkandinmoss and Tasmania.
leaflitter.Theyaregenerallymostabundantinmoistsoilsthat Althoughpauropodsoccurunderbothnativevegetationand
have high levels oforganic matter. Theirfeeding habits vary; crops, densities are variable. In studies overseas, densities of
some groupsfeed onfungi and decaying organic matterwhile pauropods ofup to several thousand have been found in soils
othersmayconsumemoresolidfood(Scheller, 1990). (Adis et al., 1999; Hagvar and Scheller, 1998; Lagerlof and
AlthoughPauropodahavebeenfoundinhumidsoilsunder Scheller, 1989) and, on a site in Denmark, they were more
native vegetation as well as on agricultural land throughout abundant than Psocoptera, Coleoptera and Symphyla, and a
theAustraliancontinent,verylittleisknownofthecomposition quarter as abundant as Acarina (Hagvar and Scheller, 1998).
ofthefauna. They areparticularly species-rich and abundant Similar densities have been found at some sites inAustralia,
inforestsoilsbutasoilsamplefromunderanynativevegetation indicatingthattheycompriseasignificantpart(5%ofindividuals)
islikelytocontainsomespecimens.Suchdataasexistindicate ofthearthropodfauna(GreensladeandMott,1979).Abundances
thatthereisaveryhighlevelofspeciesendemismofprobably arereducedbyfire(BrozaandIzhaki, 1997).
over 90%. Moreover, there are likely to be 20 to 30 species The invertebrate biodiversity of cool temperate rainforest
belongingto five orso generawithin a single vegetationtype wassampledintensivelyandextensivelyinTasmaniaaspartof
(Postle et al., 1991). This is the same order ofmagnitude as theNationalRainforestConservationProgram(NRCP)(Coyet
.
154 PenelopeGreenslade
al., 1993).Thisforesttypeoccupiesjustunder600,000hectares CollectingsitesasgiveninCoyetal. (1993).
iinnTnaosrtmhawneisatearnndTraesprmeasneinat,sw1i0t%hootfhtehrelsetsastee.xItteinssmiovsetarperaosmiinnetnhte Loc. 1 - NRCP 1 NW Tasmania, Savage River (41°19.1'S,
145°16.2'E and 41°18.5'S, 145°16.3'E), callidendrous
northeast and on the east coast and occasional patches in the m
southwest. The survey sampled 19 localities covering all the Nothofaguscunninghamiirainforest,alt.500 (GridReference
CQ558,247 and CQ560,255). Conservation status: Forest
nature conservation regions of the state accepted at the time
Reserve,partofthe’’SavageRiverPipelineCorridor”.
t(h1e99i0n)vearntdebmraajteorfraauinnaforceosmtptryepheesnswiivtehlyt,heiadiemntoiffydiongcuamreenatsinogf Loc. 2 - NRCP 2, NW Tasmania, Bradshaws Road
greatest species richness and phyletic diversity and assessing (41°49.9'S, 145°37.0'E),callidenmdrousandthamnicNothofagus
idmopcaucmtesntaotniotnheoffautnhae (iCnovyerteetbraalt.e, f19a9u3n)a. wItithwians athemafjiorsrt Ccounnsneirnvgahtaimoinirstaaitnufso:reMstt,Malut.r8c4h0ison(GRriivdeRrefReesreernvcee.CP854,680)
vegetationtypeinTasmaniaandthefirsttoassesstheconservation Loc. 3 - NRCP 3, central Tasmania, Projection Bluff
status ofthis highly endemic and species-rich fauna. The data (41°43.1'S, 146°43.5'E), high altitude callidendrous raminforest
also provided a basis for comparison with otherforest faunas dominated by Nothofagus cunninghamii, alt. 1100 (Grid
withinandoutsideTasmaniaforgroupsthatweretargeted. Reference DP770,812). Conservation status: World Heritage
survLeiys.tAednaersetimtahetedPa1u4r2o,p0o00daspsepceicmieensscooflliencvteertdebdruartiensgwetrhee AreaL.oc. 4 - NRCP 4, NW Tasmania, Cradle Mountain
collected, among them 1087 (0.8%) Pauropoda. No earlier campground(41°35.4'S,145°55.9'E),highaltitudecallidendroums
publications record specimens from Australian Nothofagus rainforestdominatedbyNothofaguscunninghamii,alt. 880
forests.Nineteenspecieswerefoundinthesurvey,ofwhich 17 (Grid Reference DP109,955). Conservation status: part
are new. Keys to families, genera, subgenera and species are conservationarea.
presented. Pauropodbiodiversity ateachlocalityis compared Loc. 5 - NRCP 5, NE Tasmania, Mt Victoria (41°20.4'S,
with that of other invertebrate groups from the same survey 147°49.9'E), callidendrous Nothofagus cunninghamii forest,
andthedatausedtomakeinferencesregardingtheconservation alt. 900 m (Grid Reference EQ693,233). Conservation status:
valueofeachregionofTasmania. forestreserve.
Loc. 6 - NRCP 6, NE Tasmania, Mt Michael (41°10.9'S,
MaterialsandMethods. 148°00.4'S), heavily logged callidendrous Nothofagus
m
cunninghamii rainforest, alt. 740 (Grid Reference
Twelvesitescoveringeachoftheconservationregionsaccepted EQ845,406). Conservationstatus: bluetierforestreserve.
at that time (1990) in which rainforest occurred, and Loc. 7 - NRCP 7, SE Tasmania, Big Sassy Creek,
representing the four main rainforest communities: (42°08.5'S, 147°54.3'E), small strip of Atherosperma
Callidendrous,Thamnic,ImplicateandOpenMontaneforest, moschatum dominated callidendrous rainforest along the
were selected for intensive sampling. The sampling was creek, alt. 400 m (Grid Reference EP749,332). Conservation
stratified sothatthere weretwo replicates in each region and status: forestreserve.
twotothreereplicatesofeachofthefourrainforesttypes.The Loc. 8-NRCP8,SETasmania,SandspitRiver(42°42.1'S,
sites arelistedbelow (Locs 1-12). Besidesthe 12 majorsites, 147°51.5'E), mixed Eucalyptus regnans and Atherosperma
some collecting was carried outin supplementary sites (Locs moschatum callidendrous forest, alt. 180 m (Grid Reference
13-19). On each of the 12 main sites the same sampling EN703,713). Conservationstatus: forestreserve.
methods were used; Tullgren funnel extraction of soil, moss Loc. 9 - NRCP 9, SW Tasmania, Frodshams Pass
andleaflitter,pitfalltrapping,yellowpantrapping,sweeping, (42°49.7'S, 146°22.9'E), callidendrous/thamnic and implicate
hand collections and pyrethrin knockdown of tree trunks rainforestdominatedbyNothofaguscunninghamii,alt. 620m
(PKD) (Coy et al., 1993). Only Tullgren funnel extraction of (Grid Reference DN497,580). Conservation status: World
leaf litter and humus was done on the supplementary sites. HeritageArea.
Pauropods were found most abundantly infunnel extractions Loc. 10-NRCP 10, SWTasmania, MtField, belowLake
ofsoil cores, moss andlitter, although afew specimens were Fenton (42°40.9'S,146°37.5'E), open montane low canopy
collected by other methods. Sampling was carried out in a m
single season-autumnforthemostpart.All specimenswere Nothofagus gunnii forest, alt. 980 (Grid Reference
DN695,746). Conservationstatus: nationalpark.
pgrievsenerivnedthienNetRhCanPolr.eDpeotrati(lCeodydeestcarli.,pt1i9o9n3s).ofthelocalitiesare (43°L0o8.c2.'S1,1 1-47N°5R4.C5P'E),11,smSaEllTarsemmannaian,t ToafsmcaanlliPdeennidnrsouuls-a
Holotypes of the species to be described in a later
thamnic intermediate cloud forest dominated by Nothofagus
publication(Scheller,inprep),willbelodgedintheAustralian
cunninghamii remains within an eucalypt dominated forest,
tNhaetiQounaelenInVsieccttoCroilalMecutsieounma.ndparatypesandothermaterialin mixed forest, alt. 400 m (Grid Reference EN742, 346).
Conservationstatus: notreserved.
Loc. 12 -NRCP 12, SETasmania, MtMangana, Bruny I
Abbreviations
(43°22.1'S, 147°17.0'E), low-canopy callidendrous-thamnic
Collectors:ATW,A.Trumbull-Ward;DR,D.Rounsevell;HM, intermediaterainforestdominatedbyNothofaguscunninghamii
H.Mitchell;JD,J.Diggle;MN,M.Neyland;PG,P.Greenslade; on Mt Mangana, alt. 540 m (Grid Reference EM229, 980).
RC,R. Coy; SS, S. Smith. Conservationstatus: forestreserve.
DistributionpatternsanddiversityofinvertebratesoftemperaterainforestsinTasmaniawithafocusonPauropoda 155
Figure1.Locationofcollectingsites,numberedfrom1to12asinthetext.Numbersinbracketsindicatetotalnumberofpauropodspeciesfound
oneachsite.InsertshowsdistributionofrainforestinTasmaniatakenfromCoyetal.(1991).
Loc. 13 - NW Tasmania, Hibbs Lagoon (42°34'S, 205). Conservationstatus: WorldHeritageArea.
145°19.5'E), rainforest (Grid Reference CN623, 853). Loc. 17 - NE Tasmania, Simons Road (41°21.5'S,
ConsLeorcv.at1i4on-sNtatTusa:smSaonuitah,WMesttSCtornosnearcvhat(i4o1°n1A0'rSe,a.147°34'E), s1p4e7c°i3a1l.3f'oEr)e,s(tGryrimdaRneafgeeremnecnetEzQo4n3e5,fo2r1s2p).riCngotnasielrs.vationstatus:
rainforest (Grid Reference EQ476, 422). Conservation status: Loc. 18 - NTasmania, Saxons Creek (41°15'S, 146°40'E),
forestreserve.
Loc. 15 - N Tasmania, Asbestos Ranges (41°09'S, rainforest (Grid Reference DQ751, 316). Conservation status:
146°40'E),NorthLeg,rainforest(GridReferenceDQ727,444). notreserved.
Conservationstatus: nationalpark. Loc. 19 - SWTasmania, OldFarmRoad, MtWellington,
Loc. 16 - SW Tasmania, Riveaux Creek (43°10'S, (42°54'S, 147°16'E), Eucalyptus forest. Conservation status:
146°38.6'E), Huon pine rainforest (Grid Reference DN704, nationalpark.
))))
156 PenelopeGreenslade
ChecklistanddistributionsofTasmanianPauropoda Brachypauropodidae
Regions: NW - northwest, NE - northeast, N - north, C - GenusBrachypauropoidesRemy, 1952
central, SE-southeast, SW-southwest Typespecies:BrachypauropoidespistilliferRemy 1952
FamilyPauropodidae Brachypauropoides sp. nov. 19 NW,NE, SE
GenusAllopauropuss.str. KeytoTasmanianPauropoda
SubgenusAllopauropusSilvestri, 1902 1. Body fusiform; tergites entire, generally not distinctly ;
Typespecies:AllopauropusbrevisetusSilvestri, 1902 legslongerthanlengthofsegment Pauropodidae2
Allopauropus(Allopauropus sp. nov. 1 NW,NE, SE, SW Body oval, somewhat flattened; tergites most often
divided, generally distinctly sclerotized; legs equal to or
Allopauropus. (Allopauropus sp. nov. 2 SE shorterthanlengthofsegment Brachypauropodidae
SubgenusDecapauropusRemy, 1931
2. Alllegs5-segmented Cauvetauropus
Type species:Allopauropus(D.) cuenoti(Remy)
First and last pair of legs 5-segmented, all other legs
DecapauropuscuenotiRemy, 1931
6-segmented 3
Allopauropus(.Decapauropus sp. nov.3 SE,NW, SW,N 3. Anteriormarginofsternal antennalbranchnearlyalways
Allopauropus(.Decapauropus sp. nov.4 NW, SE distinctly shorterthanposteriormargin....Allopauropus5
Allopauropus{Decapauropus) sp. nov. 5 SW Anteriorandposteriormarginsofsternalantennalbranch
NW nearlyalwaysofsubequallength 4
Allopauropus{Decapauropus) sp. nov. 6
4. Pygidial sternumwithsetaebl +b2 Pauropus 19
Allopauropus{Decapauropus) sp. nov.7 NW, SE
Allopauropus{Decapauropus) sp. nov. 8 NW,NE, SW Pygidial sternumwithsetaebl only Stylopauropoides 13
GenusCauvetauropusRemy, 1952 5. Pygidial sternumwithsetaebl andb2only
A. {Decapauropus) 8
Typespecies: Cauvetauropusmicrochaetus(Remy, 1952)
AllopauropusmicrochaetusRemy, 1948 Pygidial sternumwithsetaebl andb2andb3
SubgenusNesopauropusScheller, 1997 A. {Allopauropus) s. str6
6. Temporal organs without anterior appendage; distal part
Type species: Cauvetauropus {Nesopauropus) biglobulosus ofT3cylindrical;analplatewithstraightposteriormargin
Scheller, 1997 A. (A.) sp. 1
Cauvetauropus{Nesopauropus) sp. nov. 9 NE Temporal organs with distinct anterior appendage; T3
with ovoid distal swelling; anal plate with long
GenusStylopauropoidesRemy, 1956 posteromedianprocess A. (A.) sp. 2
Typespecies: Stylopauropoidestiegsi(Remy, 1949) 8. Antennalglobulusgnotaswideasmaximumdiameterof
StylopauropoidesringueletiRemy, 1962 NW,N, SW tergalbrancht 9
Stylopauropoides sp. nov. 11 NW,NE, SE,N Antennal globulus g wider than maximum diameter of
tergalbrancht A. (D.) sp. 8
Stylopauropoides sp. nov. 12 NW,NE, SE, SW
9. Distal swellingonT3 absent 10
Stylopauropoides sp. nov. 13 NW, SE,C,NE,SW
Distal swellingonT3 present A. {D.) sp. 6
Stylopauropoides sp. nov. 14 NW,C,NE, SE,SW
10. Posteriormarginofanalplateincised 11
Stylopauropoides sp. nov. 15 NW,NE, SE
Posteriormarginofanalplaterounded A. {D.) sp.3
NW
Stylopauropoidessp. nov. 16
11. Seta on coxa oflast pair oflegs furcate; T3 with simple
GenusPauropusLubbock, 1867 pubescenthairs 12
Typespecies: PauropushuxleyiLubbock, 1867 Setaoncoxaoflastpairoflegs simple;T3 withbranched
pubescenthairs A. (D.) sp. 5
PauropusdolosusRemy, 1956 SE
12. Appendageofcollumsegmentincisedanteriorly;proximal
Pauropussp. nov. 18 NW,NE,SE, SW 2/3 ofT3 thickened A. (D.) sp.4
DistributionpatternsanddiversityofinvertebratesoftemperaterainforestsinTasmaniawithafocuson Pauropoda 157
Appendageofcollumsegmentpointedanteriorly;T3with Australiabutoveralongerperiod.Ofthese31 (80%)belonged
thinaxes A. (D.) sp.7 to eitherStylopauropoides orAllopauropus. As here, alarge
proportion ofthe faunawas undescribed as only two species
13. Branches ofanalplateseparatedbyV-shapedincision were already named. Allopauropus is near cosmopolitan and
14
the most widespread and most diverse genus of pauropods
Branches ofanalplateseparatedbyU-shapedincision withnearly200speciesknownworldwide,sothehighnumber
^
S. sp. 16. ofAllopauropusspeciesinAustraliaisnotsurprising.
The genusStylopauropoideshasitsmaindistributioninthe
14. Lateral appendages onanalplateabsent 15 southernhemisphere andis notoften encounteredinthenorth.
_Lat_eraltriangularappendagesonanalplatedistinct ItssoutherlyrangeindicatesaGondwananoriginanditsabsence
S. 13 from the southern part ofAfrica may be a result of lack of
collecting. Thirteen species have been described but only one
15. Distalappendagesofanalplatewithflatposteriorsurface from north ofthe equator (Ivory Coast and Guinea). A list of
16 speciesinthe genuswiththeirdistributionsis giveninTable 1.
Distal appendages of anal plate with convex posterior ThesixnewspeciesfromTasmaniasuggestsahighconcentration
surface S. sp. 14 ofspeciesinthisgenusinTasmanianrainforests.
Pauropusalso occursworldwidehenceis considerednear
16. Anteriodistal and posteriodistal corners of sternal cosmopolitan as well. The other genera, Cauvetauropus and
antennal branch equally truncate; pygidial bl evenly Brachypauropoides, have wide but more southerly
curved 17 distributions. Only ten species have been described in
Posteriodistalcornerofsternalantennalbranchdistinctly Cauvetauropus, buttheyaredistributedoveraverylargearea
moretruncatethananteriodistalone,pygidialbl undulate fromBrazil,France,northandtropicalAfrica,the Seychelles
S. sp. 11 and Sri Lanka. The same is true ofBrachypauropoides, now
knownfromMadagascar,BorneoandNewZealand.
17. T3withlongsimplebranchesorwithbranchedpubescence; AffinitiesarestrongbetweenTasmaniaandNewZealand.
setaeontergitesfine 18 As noted above for Tasmanian rainforest, the genus
T3 with very simple pubescence hairs; setae on tergites Stylopauropoides-withnearly40%oftotal species and57%
andheadclavate S. sp. 15 oftotal specimens collected-is well represented. The genus
isalsodiverseandabundantinNewZealandwheresixspecies
18. Distal partofall trichobothriaexceptforT5 with simple havebeendescribed(Remy,1952, 1956a, 1956b),oneofwhich
curvedpubescentbranches; stcylindricalblunt...S. sp. 10 is also known from mainland Australia (Remy, 1949). In
Distal part of all trichobothria without branches except anotherNewZealandcollection, 106pauropods were studied
T5 but with branched pubescent hairs; st thin, tapering, inwhichStylopauropoidesrepresented35%ofthespeciesand
pointed S. sp.12 74% ofthe specimens (Remy, 1952, 1956a, 1956b). Although
collectingmethodsandvegetationinthetwosurveysdiffered,
19. Pygidial st subcylindrical tapering; posteriorappendages the results indicate that the genus is particularly diverse in
ofanalplate shortandblunt P. dolosus thesetworegions. OthersimilaritieswithNewZealandarein
thegeneraBrachypauropoidesandPauropus.Theformerwas
Pygidial stsimilarto aknifeblade; posteriorappendages knownpreviouslyfromMadagascar,BorneoandNewZealand,
ofanalplatelong,thinandpointed P. sp. 18 and the Tasmanian species is close to one of the two New
Discussion Zealand species, B. pistillifer Remy. Pauropus dolosus is
commontoNewZealandandTasmaniaandA. sp. 1 iscloseto
Composition and affinities. In Tasmania, as elsewhere, the A.maoriorumRemyfromNewZealand.
Pauropodidae is the most diverse family as all but one of 19 In conclusion the Tasmanian rainforest fauna is rich in
speciesbelongtoit.Ofitsfivesubfamilies,onlythePauropodinae speciesbutoflowgenericdiversity,andaffinitiesarestrongest
is present, already knownto bethe mostwidespread andmost atthespeciesandgenericlevelwithNewZealandasmightbe
species rich subfamily worldwide. The Tasmanian rainforest expected(Kantvilasetal., 1985).
faunaisrelativelyrichinspecies,butunexpectedlynospeciesin Habitats. The Tasmanian collections recorded pauropods
tdhiestrfiabmuitleydfEaumriylpyauwriothpordeipdraeseenwtaastivceosllienctBeodr.nTehoi,sNieswaGwuiidneelya fromtreetrunks (5 species) and tree ferns (1 species) forthe
and New Caledonia and on mainland Australia (New South first time. Both habitats were sampled by pyrethrin knock-
down. Numerous specimensofseveral species (15 species,71
WBrailteasi)n.aTnhdeCefnatmriallyEuisroaples,onfoorutnhderinnUcSoAolearndloactaolivteires20in00Gmreaitn rneoctorcdosm)mwoenreincooltlheecrtehdabfitraotms.mIonssraionnforsietsets,wmhoesrsectohmeymownelrye
theHTihmeamlaajyoarsit(yScohfeltlheer,NR19C9P0asnpdeciinmcelnusdebderleofnegretnoctesw)o.genera, has a higher moisture content which may account for higher
speciesrichnesshere.
Allopauropus with eight species and Stylopauropoides with
seven species,togethercomprising 80% ofthe species. Postle Abundance. Species varied in numbers of specimens caught
et al. (1991) found 39 species in forests in south Western fromoneindividualtoover200.Sevenspecieswererepresented
158 PenelopeGreenslade
Table1.ListofknownspeciesinthegenusStylopauropoideswiththeirdistributions.InadditionsixnewspeciesfromTasmaniaarerecordedin
thispaper.
Species Localities References
1. S.tiegsi (Remy) Australia(eastern) Remy 1949
NewZealand Remy 1952c, 1956a, 1956b
2. S. bornemisszaiRemy Australia(western) Remy 1957,
Postleetal. 1991
3. S. ringueletiRemy Argentina Remy 1962
Chile Scheller 1968
Tasmania Newrecord
4. S. lambdaRemy NewZealand Remy 1956b
5. S. subantarcticus Scheller CrozetIslands Scheller 1974
6. S. infidus(Remy) NewZealand Remy 1956a
7. S. duplex(Remy) NewZealand Remy1956a
8. S. bilobatusScheller NewCaledonia Scheller 1993
9. S. hirtus(Remy) NewZealand Remy 1952c, 1956a
10. S.delamarei(Remy) IvoryCoast Remy 1948
Guinea Remy 1959a
11. S. vadoni(Remy) Madagascar Remyl956c,
Remy&Bello 1960
12. S. incisusRemy &Bello Madagascar Remy&Bello 1960
13. S furcillatus(Remy) NewZealand Remy 1952c
.
NewCaledonia Scheller 1993
by fewerthan 20 individuals (37%), eight species represented although itis also possible that some ofthese species atleast
bybetween21 and99individuals(42%)andfourspecies(21%) arehaveapreferenceforothervegetationtypessuchaseucalypt
representedbyover 100individuals.Therewasadistincttrend forest.
forthe most numerous species to be the most widespread, as
Regionaldiversity.Faith(1992)describedamethodofranking
expected.
sites based onphyletic diversity in whichbranch lengths on a
Regional distribution. Because the NRCP material comes phylogenetically derived tree of relationships were used as a
almostexclusivelyfromrainforests (fig. 1), species occurrence measure of divergence and hence conservation uniqueness.
inothervegetationtypesisnotknown.Withinrainforests,most FaithandGreenslade(1994)providedapreliminaryanalysisof
species werefoundto be distributed widely inthe state being three groups of invertebrates from the NRCP sites which
recordedfromall majorregions (table. 2). Seven species were indicated that the northwestern sites (Locs 1, 2 and 4) were
widely distributed: A. sp.l,A. sp. 3, S. sp. 11, S. sp. 12, andP. majorcontributorstophyleticdiversity.Althoughphylogenetic
sp. 18,werefoundinfourregionsandS. sp. 13 andS. sp. 14in trees are not available for any of the groups (except for
fiveregions.Twootherspecies, S. sp. 15 andS. sp. 19 seemto Triaenonychidae Opiliones (Hunt, 1996), new taxonomic
have more northerly and easterly distributions; two, A. sp. 8 informationis availableforthreemore groups. Inthe absence
andS. ringueleti, were most commonlyfound inthe northern ofrelationship trees, the dataare usedto calculatethe simple
and southwest regions, and two species recorded from both measure of ”taxic” diversity for six groups; Diplopoda (R.
northwest and southeast regions, A. sp. 4 and A. sp. 7. Six Mesibov, unpublished data), Opiliones (G. Hunt, unpublished
species, A. sp. 2,A. sp. 5,A. sp. 6, , C. sp.9, S. sp. 16 and P. data), euptyctime oribatid mites (Niebala and Colloff, 1997),
dolosuswereonlyfoundatasinglesiteandmaybeshortrange Amphipoda (A. Friend, unpublished data) and Isopoda (A.
endemics. This is a high proportion (30%) of rare species Green,unpublisheddata)inadditiontothePauropoda(tables 1
Distribution patternsanddiversityofinvertebratesoftemperaterainforestsinTasmaniawithafocuson Pauropoda 159
Table2.RecordsofPauropodaspeciesfromNRCPsiteswithtotalspecies,subgenera,genera,familiesandtaxicscoreforeachsite
Numberof
Sites records
Main Other
Species 1 2 3 4 5 6 7 8 9 10 11 12 Othersites sites sites
Allopauropus sp. nov. 1 + + + + + + 13,16 6 2
Allopauropus sp. nov. 2 + + + 3 0
Allopauropus sp. nov.3 + + + + + 13,16,18,19 5 4
Allopauropus sp. nov.4 + + + + 4 0
Allopauropus sp. nov. 5 16 0 1
Allopauropus sp. nov. 6 + 1 0
Allopauropus sp. nov.7 + + 2 0
Allopauropus sp. nov. 8 + + + 3 0
Cauvetauropussp.nov. 9 + 1 0
StylopauropoidesringueletiRemy, 1962 + 14,15,19 1 3
Stylopauropoides sp. nov. 11 + + + + + + + 17, 18 7 2
Stylopauropoides sp. nov. 12 + + + + + + + 13.16,17 7 3
Stylopauropoides sp. nov. 13 + + + + + + + + + 13,17,19 9 3
Stylopauropoides sp. nov. 14 + + + + + + + + + + + 11 0
Stylopauropoides sp. nov. 15 + + + + + 17 5 1
Stylopauropoides sp. nov. 16 + 13 1 1
PauropusdolosusRemy, 1956 + + + 3 0
Pauropussp. nov. 18 + + + + + + + 13,17 7 2
Brachypauropoidessp. nov. 19 + + + + + 13,17 5 2
Total species 11 7 2 8 5 9 10 7 5 2 5 10
Total subgenera 5 5 1 3 2 5 4 5 2 2 3 4
Total genera 4 4 1 2 2 5 3 4 3 2 3 3
Totalfamilies 2 2 1 1 1 2 2 2 1 1 1 1
TAXICSCORE 22 18 5 14 10 21 19 18 11 7 12 18
and 2). These groups include avariety ofdecomposer groups Theappendixlists speciesidentifiedfromthecollections. The
and a predatory and scavenger group (Opiliones) and were largernumbersofeuptyctimeoribatidAcarina,Diplopodaand
selected partly on the basis that they were characteristic of Opilione species compared with smaller detritivores such as
rainforest faunas and also because they showed a negative Pauropoda,maybeacharacteristicofrainforestfaunas.
response to disturbance (Coy et ah, 1993; Greenslade, 1992); Table 2 summarisesthe site record dataforall 19 species
hence they could act as surrogates for the whole rainforest of pauropod. The number of species, subgenera, genera and
fauna. They were also the only groups for which all species familiesateachofthemaintwelvelocalitiesisgivenandthese
identifications were available. Therewere some differences in figures summed, withoutweighting,to give an simple overall
the total number of species found in each higher taxon with ’’taxic” richness”. This method gives more weight to the
Opilionesbeingthelargestgroupwith34species,followedby phylogenetic diversity found on each site than does a simple
euptyctime oribatid Acarina with 31, Diplopoda with 29, speciesrichnesscountandisusedherebecauseoftheabsence
Isopodawith 28, Pauropodawith 19 andAmphipodawith 17. of any formal phylogenetic analysis. The collecting intensity
160 PenelopeGreenslade
Table3.Numbersofspecies,genera,familiesandtotaltaxicscoreofPauropoda,euptyctimeoribatidmites(NiebalaandColloff,1997),Opiliones
(G. Hunt, pers. comm.), Diplopoda (R. Mesibov, pers. comm.), Amphipoda(A. Friend, pers. comm.) and Isopoda (A. Green, pers. comm.)
recordedoneachofthetwelvemainNRCPsitesduringthesurvey.
Taxa Sites
1 2 3 4 5 6 7 8 9 10 11 12
species 11 7 2 8 5 9 10 7 5 2 5 10
Pauropoda genera 4 4 1 2 2 5 3 4 3 2 3 3
families 2 2 1 1 1 2 2 2 1 1 1 1
Taxicdiversity 17 13 4 11 8 16 15 13 9 5 9 14
species 17 12 4 0 6 3 8 6 6 4 10 15
genera 3 5 3 0 3 2 3 3 2 2 3 4
Oribatidmites
families 2 3 2 0 1 1 2 2 1 1 2 2
Taxicdiversity 22 20 9 0 10 6 13 11 9 7 15 21
species 11 12 1 1 12 10 7 10 4 1 3 8
Opiliones genera 6 9 1 1 10 7 6 9 2 1 2 6
families 3 3 1 1 3 3 2 2 2 1 1 3
Taxicdiversity 20 24 3 3 25 20 15 21 8 3 6 17
species 9 6 0 2 6 2 6 4 2 2 3 5
Diplopoda genera 6 5 0 2 5 2 5 4 1 1 3 4
families 3 2 0 2 1 2 4 4 1 1 2 3
Taxicdiversity 18 13 0 6 12 6 15 12 4 4 8 12
species 5 4 0 0 0 0 0 1 2 0 2 6
Amphipoda genera 4 4 0 0 0 0 0 1 2 0 2 5
families 1 1 0 0 0 0 0 1 1 0 1 1
Taxicdiversity 10 9 0 0 0 0 0 3 5 0 5 12
species 12 9 1 7 6 4 5 7 5 3 6 7
Isopoda genera 5 5 1 3 3 2 4 4 2 1 3 3
families 4 4 1 2 2 2 3 3 2 1 3 3
Taxicdiversity 21 18 3 12 11 8 12 14 9 5 12 13
Totaltaxicdiversity 108 97 19 32 66 56 70 74 44 24 55 89
Siteranking 1 2 12 10 6 7 5 4 9 11 8 3
wasidenticalatallthe 12mainsites soprovidingasolidbasis scoresareallhighaltituderainforest(over880m). Overallthere
forcomparison. The dataindicates thatthe Savage River site is atrend ofdecreasingtaxic diversitywithaltitudeforall sites
(NWLoc.1)isthemosttaxicrichsitefollowedbyMtMichael with a significant negative correlation of taxic diversity with
(NE, Loc. 6) and Big Sassy Creek (SE, Loc. 7). Three other altitude ofr= -.61, p<0.05. This findingis not surprising given
sites(NWLoc.2,SELoc. 8andSELoc. 12)allhaveanequal the sparser tree cover resulting in more open vegetation and,
score so high taxic richness appears to be spread over all presumably, higher dessication rates at the higher altitudes.
regionsexceptforthesouthwestandcentralnorth. AlthoughmontanevegetationtypeswerenotpresentatFrodshams
Total taxic richness for the five other groups and a mean Pass,itwaslocatedatahighaltitudeandthreetypesofrainforest
rankingforall sitesis showntogetherwiththepauropoddatain weresampledratherthanonlythemorecommonCalledendrous
tables 3 and 4. There is atendency for all the groups to show type,whichmaybethereasonforitslowspeciesrichness.
similar patterns of species richness between localities as that The Symphyla which were also studied from these
shown by the pauropods. Localities from the northwest and collections (Clarke and Greenslade, 1996) also showed a
southeast(1,2,6,7and 12)haveconsistentlyhighrankingsfor higherspeciesrichnessfromlocalities1and2inthenorthwest
all six invertebrate groups while localities 3, 4, 9 and 10 are than 7to 9 and 11 and 12. Material from sites 3 to 6 and 10
consistently low. The same pattern was found when the taxic werenotstudiedsothistaxonhasnotbeenincludedhere.
diversitywas calculatedusing aweighting system(notshown). Factors affecting the taxic diversity, apart from altitude,
Apartfrom Frodshams Pass (9), the otherthree sites with low rainforest type or region could be annual rainfall, area of
DistributionpatternsanddiversityofinvertebratesoftemperaterainforestsinTasmaniawithafocuson Pauropoda 161
Table4.Individualandmeanrankingsoftaxicscoresofselectedinvertebratesfromthetwelvemainrainforestsites
Pauropoda Oribatid Opiliones Diplopoda Amphipoda Isopoda
Acari
Sites
based ranks
ranks
diversity diversity diversity diversity diversity diversity rank
Ranking Ranking Ranking Ranking Ranking Ranking rank
Summed Mean summed
Taxic Taxic Taxic Taxic Taxic Taxic
Final
on
1 17 1 22 1 20 4 18 1 10 2 21 1 10 2 1
2 13 5 20 3 24 2 13 3 9 3 18 2 18 3 2
3 4 12 9 8 3 10 0 12 0 7 3 12 61 10 12
4 11 7 0 12 3 10 6 9 0 7 12 5 50 8 10
5 16 2 13 5 15 7 15 2 0 7 12 5 29 5 5
6 16 2 6 11 20 4 6 8 0 7 8 10 42 7 8
7 15 3 13 5 15 7 15 2 0 7 12 5 29 5 5
8 13 5 11 6 21 3 12 4 3 6 14 3 27 5 4
9 8 10 9 8 6 8 4 10 5 4 9 9 49 8 9
10 5 11 7 10 3 10 4 10 0 7 5 11 59 10 11
11 9 8 15 4 6 8 8 7 5 4 12 5 36 6 7
12 14 4 21 2 17 6 12 4 12 1 13 4 21 4 3
rainforest or conditions during collection. Highest rainfall is species occurs on each andnowhere else. Riveaux Creek,just
recordedfromhighaltitudelocalitiesaswellasintheextreme withintheWorldHeritageArea, isthe only site sampledwith
northwest and east, while lowest rainfall occurs in the the rare rainforesttree, Lagerostrobosfranklinii (Huon pine)
southeast.Itdoesnotappearthataveragerainfallisinfluencing andnoNothofagus.Thereservationstatusofthesethreesitesis
taxic richness. It should be noted, however, that rain fell relatively secure and not subjectto logging butone important
heavily during sampling atLoc. 2 andfaunawas particularly siteforinvertebrates, SimonsRoad,is subjecttologging. This
activeatthis siteduringcollection. siteshouldbedesignatedasaprotectedforestryreserve.
The similarity shownbythe differentinvertebrate groups Information on invertebrates from the NRCP survey has
gives the taxic pattern more validity than that from a single already provided the basis for special protection for several
group. Moreover, the large number of species (158), genera localities and all the twelve main localities, not formally
(67)andfamilies(20)aswellashighergroups(six)contributing protectedbyotherlegislation,wereacceptedforlistingonthe
tothepatterns,addtoitssignificance.Theseresultsdemonstrate National Estate Register. With the enactment of the
thevalueofsuchtargetedsurveys, suchasthis one supported Commonwealth Environment Protection and Biodiversity
by the NRCP, especially when sampling is carried out Conservation Act 1999, however, land managers were no
strategically. longer obliged to consult the register when assessing
Conservationvalues. Few sites,apartfromthoseintheWorld applicationsfordevelopmentsonsiteswithnativeecosystems.
HeritageArea,wereformallyreservedwithintheparkssystem TherecommendationfromtheNRCP 1990/1 surveywasthat
atthetimetheprojectwascarriedout(1990),althoughseveral Locs. 1, 2, and 6, not then protected, should be designated
sitesarenow speciallyprotectedforestreservesandsoarenot specialreservestoprovideforimprovedmanagementofthese
subject to logging. Loc. 1 was gazetted in part to protect sitesforprotectionofinvertebratebiodiversity. Sincethattime
invertebrates, althoughonly acorridoralongthe SavageRiver thishaslargelybeenachieved.
piplelinehasbeenreserved; othertractsofforestinthisregion The conservation of Tasmanian rainforest is of high
remainunprotected.Locs1(SavageRiver),6(BigSassyCreek) importancebecauseitis onlyherethatlargetracts ofitremain
and16(RiveauxCreek)areuniqueinthatadifferentpauropod in Australia, most notably in the northwest (Anon, 2008).
162 PenelopeGreenslade
Decisions on selection of reserves until recently has been Faith,D.P 1992.Conservationevaluationandphylogeneticdiversity.
largelybasedonvegetationand,intheRFAprocess,vegetational BiologicalConservation61: 1-10.
associations were given most weight (Anon, 1997). Because Faith, D.P, and Greenslade, P. 1994. Phyletic diversity and the
invertebrates integrate environmental variables atafinerscale assessment of invertebrate biodiversity ofTasmanian rainforest
thandoplants andaremore species rich,they exhibitahigher areas.P.90in:ProceedingsoftheInternationalForestBiodiversity
Conference, Conservingbiologicaldiversity intemperateforest
levelofsmallscaleendemismthandoplants(Harvey,2002).It ecosystems - towards sustainable management. Centre for
haspreviouslybeennotedthatNothofagusrainforestexhibitsa Resource and Environmental Studies, Australian National
higher level of generic diversity for Collembola than other University:Canberra.
vegetationtypesinthestate(Greenslade, 1987) andtheresults Greenslade,P.1987.GenericbiogeographyofTasmanianCollembola.
oftheNRCPsurveyemphasisedthispoint.Thespeciesrichness Pp 653-60 in Soil Faunaand Soil Fertility in: Striganova, B.R.
ofinvertebratesfoundinthis vegetationtypeinTasmaniawas (ed.).ProceedingsoftheNinthInternationalColloquiumonSoil
estimatedataround750(excludingtwospeciesrichgroups,the Zoology1985.Moscow.
HymenopteraandDiptera) (Coy etal., 1993).Althoughonlya Greenslade,P. 1992.ConservingInvertebratediversityinagricultural,
proportionoftheknownfaunaisreportedinthispaper(20%), EfocroessytsrytemasndandnaEtnuvrailronemceonstys4t0e:m2s97i-n312A.ustralia. Agriculture,
thedataillustratetheirrelevancetolandmanagementespecially Greenslade, P., and Mott, J.J. 1982. Collembola and other soil and
in decisions onformal reserves because invertebrates have an surface fauna of native and improved pastures at Katherine,
apparenthighdegreeofendemism,eveninasingleforesttype NorthernTerritory.Pp299-307in:Lee,K.E.(ed.)Proceedingsof
suchasNothofagusrainforest. the Third Australian Conference on Grassland Invertebrate
Ecology.CSIRO:Melbourne.
Acknowledgments Hagvar,S.,andScheller,U. 1998.Speciescomposition,developmental
stages andabundanceofPauropodain coniferousforestsoils of
Special thanks are directedtothe collectors and are also due southeastNorway.Pedobiologia42: 278-82.
to the Parks and Wildlife Service, Forestry Commission and Harrison, L. 1914. On some Pauropoda from New South Wales.
DepartmentoftheArts,Sport,theEnvironmentandTerritories Proceedings ofthe Linnean Society ofNew South Wales 39:
(now Department of Environment, Water, Heritage and the 615-34.
Arts) for financially supporting the project. Grateful thanks Harvey, M. 2002. Short-range endemism amongst the Australian
are also dueto G. Hunt,A. Friend,R. Mesibov andA. Green fauna: some examples from non-marine environments.
InvertebrateSystematics16(4):555-70.
foridentifying material andtothereferees for suggestions as Hunt, G.S. 1996. A preliminary phylogenetic analysis ofAustralian
to improving the paper. Finally many thanks are due to U. Triaenonychidae (Arachnidae: Opiliones). Revue Suisse de
SchellerforhispreliminaryidentificationsofPauropoda. Zoologievol.horsserie:295-308.
Lubbock,J. 1867.OnPauropus,anewtypeofcentipede.Transactions
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