A Preliminary Study of Bryophytes and Invertebrates of Soil Crusts in the Little Desert National Park and Surrounds

2006·2.4 MB·English

BryophyteSpecialIssue
A
preliminary study ofbryophytes and invertebrates ofsoil
crusts in the Little Desert National Park and surrounds
Josephine Milne1 Megan Short2 and Karen Beckmann'
,
'NationalHerbariumofVictoria,
RoyalBotanicGardensMelbourne,BirdwoodAvenue,SouthYarraVictoria3141
2LifeandEnvironmentalSciences,DcakinUniversity,221 BurwoodIIwy,Burwood,Victoria3125
Abstract
This study is preliminary to ongoing investigations ofsoil crusts and associated invertebrates in
north-west Victoria, locusing on the Little Desert National Park. Ninety quadrats from nine sites
were sampled. Eighteen bryophytespecies (nine mosses, nine liverworts) were identifiedwithin the
quadrats. All invertebrates were from the Phylum Arthropoda. Overall abundance and diversity of
invertebrateswas low. Whilesamplinginthedriermonths isvaluable forobservingthedynamicsof
soil crusts in this region, a more comprehensive assessment ofspecies diversity is gained by sam-
plingduringwetterperiods. (TheVictorianNaturalist123(4),2006, 195-203)
Introduction
Soil generally is considered a precious particular landuse (e.g. grazing, cultivation)
resource, but what is the value placed on on the dynamics ofsoil crusts (Eldridge et
the organisms that comprise soil crusts? a!. 2000) and the effect of management
Bryophytes, together with lichens, fungi activities (e.g. burning oft) on these crusts
and cyanobacteria (blue-green algae) make (Eldridge and Tozer 1997a; Hodgins and
up the biological or cryptogamic crusts Rogers 1997). Thesestudiesconcentratedon
that play a very important role in protect- areas ofwestern New South Wales, south-
ing soils ofthe arid and semi-arid zones of westernSouthAustraliaandQueenslandand
Australia, including sensitive rangelands highlighted the diversity ofcryptogamic
(Eldridge and Tozer 1996; 1997a; 1997b; organisms in soil crusts andthe abiotic con-
Hodginsand Rogers 1997, Rosentreterand ditions conducive to development ofthese
Eldridge 2002). Biological crusts protect crusts. In Victoria, short lists ofbryophytes
soils from erosion, regulate infiltration of have been included in vegetation studies of
rainfall, provide a suitable microhabitatfor Hattah Lakes (Willis 1970) and Wyperfeld
germination ofseed, photosynthesisewhen (Scott 1982)National Parks, but thereareno
moist, therefore acting as a carbon sink formal systematic studies of soil crust
(Moore 1998; Eldridge 2000), and provide bryophyte species, the invertebrates that
food and shelter for invertebrates. In turn, inhabit them, or studies focusing on the
invertebrates play an important role in the dynamicsofsoilcrusts.
regulation ofdecomposition and nutrient The objectives ofthis preliminary study
cycling within the crust and soil beneath were to record the composition and abun-
(Belnap 2001). Recent research (Eldridge dance of soil crust bryophytes and docu-
2005) has highlighted the importance of ment the invertebrate fauna inhabiting
biological crusts as indicators ofthe effec- these crusts in the semi-arid zones of
tiveness of landscape management. north-western Victoria, in particular the
Conservation of soil crusts requires not Little Desert National Park (LDNP), Little
only an understanding ofthe organisms Desert Lodge, North Goroke State Forest
that comprise them, butalso ofthe interac- andJane DuffReserve.
tions that occur within them, and how
Methods
speciescomposition variesgeographically.
In Australia, a number ofstudies have Studyarea
examined the composition ofsoil crusts in The Little DesertNational Park is located
arid and semi-arid areas and rangelands in the Wimmera 375 km north-west of
(Eldridge and Tozer 1996; 1997a; Eldridge Melbourne. The area is described as semi-
1998a; 1998b; Eldridge2001), the impactof arid with mean daily maximum summer
Vol. 123 (4) 2006 195
Bryophytespecialissue
temperaturesranging from 28 to30°C and Brown StringybarkEucalyptusbaxteri,with
mean maximum winter daily temperatures large patches ofheath and Mailee-broom-
ranging from 14 to 15 °C (Bureau of bush Melaleucauncinata, particularly in the
Meteorology August 2004). Mean annual eastern and central blocks. The western
rainfall is415 mm with mostofthe rainfall block is almost all brown stringybark with
occurring from May to October (Bureau of small scattered patches ofgum-box-Buloke
Meteorology August 2004). The Wimmera woodland (consisting of Yellow Gum
plains were originally covered by wood- woodland and Slender Cypress Pine wood-
lands ofYellow Gum, Buloke and Black land) and Mallee-broombush (Land
and Grey Box with largeexpanses ofgrass- Conservation Council 1985). The LDNP
land between the woodlands (Land occurs predominantly in what is now
Conservation Council 1085). Since referredtoastheWimmera Bioregion (DSE
European settlement most ofthe natural 2006). Part of the western block of the
vegetation has been cleared for agriculture LDNP is also within the Lowan Bioregion
and the LDNP is all that remains ofthe (DSE 2006). Ecological Vegetation Classes
original vegetation. Thenational parkbegan have been determined for the two biore-
as a small reserve for the protection ofthe gionswithintheLDNP(DSE2006).
Malleefowl (National Parks Service 1996). The first fieldtrip in November 2003 sur-
dInevtehleolpattehe19a6r0esatfhoerreagwreircueltpulraen.sTtohifsurptrhoe-r vLeDyNedPsaintedssiintetshewi‘tehaisntetrhnebLliotctlke' Doefsetrhte
posal met with strongpublic opposition and Lodge (Fig. I). The second fieldtrip con-
pinar1k.96T8hteheLaDreNaPwahsaspreoxcplaainmdeeddaonvaetriontahle d‘uwecstteedrninaJnudnecen2t0r0al4bsluorcvkesy'edofsittheesLinDNthPe
years, and by 1988 comprised 132 000 ha andintheNorth GorokeState Forest. Inthis
(National Parks Service 1996). The vegeta- study, a total ofnine sites was examined in
tion ofthe national park is predominantly detail (Fig. 1). It was originallyproposed to
Victoria•South
Australiaborder
Fig. 1. Location ofstudy sites and predominant vegetation type at each site. I. Whimpy’s Little
DesertLodgeNatureTrail ‘claypan 1’ (SlenderCypress Pine Woodland); 2. Whimpy’s LittleDesert
LodgeNature Trail ‘claypan 2’ (Slender Cypress Pine Woodland): 3. Kiala camp ground (Yellow
Gum Woodland); 4. Salt Lake Road (Hcathland); 5. Stringybark Walk (Slender Cypress Pine
Woodland); 6. Mt MoffatTrack(Yellow Gum Woodland); 7. MtMoffatTrack,just north ofEast-
WestRoad(YellowGum Woodland);8. SouthernendofSambell’sTrack(YellowGumWoodland);
9.NorthGorokeState Forest(YellowGumWoodland).
196 The Victorian Naturalist
Bryophytespecialissue
samplealongtransects in diversevegetation coxylon F.Muell. subsp. leucoxylon (Fig.
types within the national park, but once in 2b). At each ofthese sites there was some
the field it became evident that sites which clay component in the soil. The Jane Duff
soley consisted ofsandy soils supported lit- Reserve, 5 km westofMitre, was visited en
tleorno soil crusts.Anybryophytespresent routetothenational park.
were restricted to the base of shrubs.
aThnedreifnorweo,osdalmapnldinsgdtoomoiknpaltaecedibnyheSaltehnldaendr DaAtatrcaonlsleecctti(o1n00 m) was set out at each
Cypress Pine Callitris gracilis R.Baker site. Soil crust bryophyte and lichen
(Fig. 2a) or Yellow Gum Eucalyptus leu- species were sampled from 30 x 30 cm
quadrats, at 10 m intervals along the tran-
Fig. 2 a. SlenderCypress Pine Callitrisgracilis b. Yellow Gum WoodlandEucalyptus leucoxylon
,
subsp. leucoxylon.
Vol. 123 (4) 2006 197
.
Bryophytespecialissue
tsehcety.aSroeilinsduircfaatcievefeoaftutrheeslairkeeliimhpooodrtoafntsoaisl TinabtlheeL1i.ttBleryDoepsheyrtteNsatrieocnoarldePdarwki,thLiitntlqeuaDdersaetrst
crust formation. Biotic and abiotic aspects Lodge and North Goroke State Forest Victoria,
were recorded for each quadrat along a Australia.
transect, including characteristics of soil Taxa
surface morphology e.g. slope within a
Mosses
quadrat, surface microtopography and Biyaceae
crust coherence (see Eldridge and Tozer Rosulabryumbillardierei(Schwagr.)
1997). Within each quadrat the vascular J.R.Spence*
plant cover and leaflitter cover were esti- Rosulabryumcampylothecium(Taylor)
mated. The percentage total soil crust J.R.Spence
Ditrichaceae
cover within the quadratwas then estimat- Eccremidiumsp.
ed together with the proportion ofthe Gigaspermaceae
algal, bryophyte and lichen components. Gigaspermumrepens(Hook.)Lindb.
Small samples ofmosses and liverworts Leucobryaceae
were taken to confirm identification. Some Campytopusintrqflexus(Hedw.)Brid.
collections ofthe liverwort Ricein were PoPloyttyrtirchiaceheuamcjuniperinitmHedw
fertile. Their spores were examined with a Pottiaceae
scanning electron microscope (SEM) BarbulacalycinaSchwagr.
because the microscopic structure of BarbulacrinitaSchultz
spores assists in the identification ofthese Didymodontorquatus(Taylor)Catches.
plants. The relationship between the suite TTorirqtuueltarealnltaarpcatpiiclala(tHaa(mHpoeok).fW.ilasnodnWilson)
of soil crust species and vegetation type, Broth.
leaflitter cover, topography, soil type and Splachnaccae
associated water retention will be reported Tayloriaoctoblepharum(Hook.)Mitt.*
elsewhere. Liverworts
Soil crust samples (10 x 10x2 cm deep) AcErnoibgomleblalcaeatehallinaG.A.M.Scottand
were collected from quadrats along each K.G.Beckm.
transect and the macro-invertebrate fauna Lethocoleapansa(Taylor)G.A.M.Scottand
extracted in the laboratory using Tullgren K.G. Beckm.
funnels (Gullan and Cranston 2000). In the Amelliaceae
first fieldtrip, soil crust samples were taken Gongylanthusscariosus(Lehm.)Steph.
Aytoniaceac
from four quadrats per transect, but as this Asterelladrummondii(Hook.f.andTaylor)
yielded a low number ofinvertebrates, five R.M.Schust. exD.G.Long
quadrats were sampled along each transect Asterellasp.
duringtheJune2004samplingperiod. After Fossombroniaceae
the invertebrates were extracted, any moss- FossombroniaintestinalisTaylor
esand liverwortspresentwereidentified. Fossombroniasp.
Ricciaceae
Soil from the November2003 crust sam- Ricciapapulosa(Steph.)Steph.
ples was potted out in small sterile pots Ricciasp.
filled with sterile coarse sandto determine
whether spores or asexual propagules of * Recordedatstudysites,butnotinquadrats.
mosses and liverworts were resting in the were identified (Table 1) from 90 quadrats
soil. The pots were placed inambient light sampled from ninesites. A furthertwomoss
and temperature, watered regularly with species Rosulabryum billardierei and
distilled water and covered with a plastic Tayloria octoblepharum were recorded in
sheettoavoidcontamination. the vicinity ofsome ofthe quadrats. Ofthe
Taxonomic nomenclature follows
12 moss taxa, five were from the family
Streimann and Klazenga (2002) for moss- Pottiaceae and two from the family
es, and McCarthy (2003)for liverworts. Bryaceae. The predominant liverworts
Results recorded were the thallose genera Asterella
Crustfloristics andRicciaand the leafy species,Lethocolea
Eighteen bryophyte species (nine mosses pansa and Fossombronia (Table 1 ). The
andnineliverworts)representing 11 families Jane Duff Reserve proved rich in Riccia
198 The Victorian Naturalist
Bryophytespecialissue
with three species being recorded, R. Table 2. Invertebrates recorded in soil crust
cavenosa,R. cristallinaandR. multifida. within the Little Desert National Park, Little
Two mosses, Fissidens sp. and Funaria Desert Nature Lodge and North Goroke State
sp., that were not recorded in the quadrats forest,Victoria,Australia.
grew in thepots from soi! samples collect- Morphospeciescollected
ed inNovember2003. Nov Jun
The impact ofseason on the percentage 2003 2004
ofsoil crustcoverandthecontributionpar- Order
ticular cryptogams made to crust cover is Arancae _
1
depicted in Fig. 3. Sampling in June 2004, Acari 6 9
after substantial rainfall, showed that algae Hymenoplera 1 2
and liverworts, particularly Asterella sp., Coleoptera(larvae) - 2
Ffoosrmseodmbtrheonpiraedsopm.inaanndtLceotmhpoocnoelnetaspoafntshae HDLieeppmtiiedprotapetr(elaraarv(alea)rvae) 1-- 11-
soil crusts (Figs. 4d and 5b). In contrast, Blattodea 1 _
crusts sampled during the dry period of Collembola 1 5
November 2003 consisted mainly of Total 10 21
mosses(Fig. 3).
worts Gongylcmthus scariosus, Lethocolea
Invertebrates pansa and Riccia multifida have not been
All invertebrates collected in this study documented in previous soil crust studies.
were from the Phylum Arthropoda. Differences between the suite of species
Increasedabundanceandactivity ofinverte- recorded can be attributed to vegetation
brates was noted in the June 2004 sampling communities, soil types, level of distur-
period. They were observed crawling over bance (Eldridge and Tozer 1996; 1997a;
the soil crusts, whereas none was observed Hodgins and Rogers 1997) and sampling
in thedrierconditions in June2003. No dif- season. Also, because ofthe small size and
ference in either abundance or diversity of ephemeral nature of many soil crust
extracted macro-invertebrates was found bryophytes, taxacanbe overlooked. In this
between the two sampling periods, with the study, the season in which surveys were
exception ofthe insect order Collembola conducted influenced the taxa recorded
(springtails). The majority of arthropods and, in particular,theirrelativeabundance.
extracted were mites and springtails (Table Substantial rainfall in early winter (June
2). Only a small number of ants were 2004) influenced the dynamics ofthe soil
extracted from the soil crusts, but our field crust cover at the study sites, and the
observations suggest that ants are present at ephemeral nature ofliverworts became
mostsites but appeared to be movingacross quite apparent. There had been heavy rain-
soil crusts, between patchesofshrub and lit- fall intheweekspriortothistripand liver-
tercover, rather than inhabiting the areas of worts formed oneofthe predominant com-
soil crusts. Eight ant species were recorded ponents ofthe soil crusts. In the November
moving across transects: Anonychomyrma 2003 fleldtrip, much ofthe liverwort bio-
sp., Iridomyrmex sp. (meat ant), mass wras not evident, being in a dormant
Camponotus sp., Doleromyrma sp., summer phase and nearly impossible to
Pheidole sp. I, Pheidole sp. 2, detect, orresting in thesoilaseitherspores
Rhytidoponera sp., and Tapinoma sp. In the or asexual propagules, which produced
June 2004 sampling period, there were newplants with the onsetofrain (Fig. 5b).
Diptera, Coleoptera and Lepidoptera larvae Thegrowth ofliverworts from soil collect-
inthecrustsamples. ed in November 2003 is evidence that the
soil does acts as a diaspore bank. In June
Discussion 2004, gemmae were detected amongst the
The majority of bryophytes recorded in leaves ofthe liverwort Lethocolea pansa
this study also have been documented in indicatinga strategy in this species ofpro-
othersoil crust studies (Eldridge and Tozer ducing many asexual propagules at the
1996; 1997a; 1997b; Eldridge et a/. 2000; beginning ofthe growing season, prior to
Thompson and Eldridge 2005). The liver- the production ofgametangia (male and
Vol. 123 (4) 2006 199
Bryophytespecialissue
Fig.3. Mean percentagecoverofmoss,liverwort, lichen andalgaein soilcrustsatsitessurveyed in
theLittleDesertNational Park,LittleDesertLodgeandNorthGorokeStateForest,Victoria.
female sex organs) (Beckmann 1993). It new growth is initiated once favourable
became apparent that once pots were conditionsreturn (Beckmann 1993). In this
allowed to dry out, some liverwortspecies state, the presence ofthese plants is diffi-
e.g. Fossombronia sp. and Lethocolea cult to detect and would explain the lower
pansa, shrivelled and dried very quickly percentageofliverwortcrustcomponent in
and were difficult to detect on the soil sur- the November 2003 sampling (Fig. 5a). In
face. The stems ofthese perennial species contrast, the liverworts Riccia and
often act as tubers that persist after the Asterella were recorded during the
extremitieshavedried and deterioratedand November 2003 sampling. These species
Fig.5a. Patchofdrycryptogamiecrust,November2003, b. magnifiedsectionofsoilcrustaftersig-
nificantrain, showing growth ofephemeral liverwortsFossombroniasp.,Lethocoleapansaand the
mossEccremidiumsp.,June2004.
200 The Victorian Naturalist
Bryophytespecialissue
Fig. 4 a. Dry moss cushion, partially inundated with sand, b. moss cushion after rain, c. mosses
betweenpatchesoi toliose lichen,d.algalcrustin Callitrisgraciliswoodland.
Vol. 123 (4) 2006 201
Bryophytespecial issue
demonstrated a strategy ofdesiccation tol- From these observations it is recom-
erance where plants employed various mended that future work on the study of
mechanisms, in this case scales, to facili- soil crusts involve sampling during the
tate survival ofmatureplants which rapid- wetter months to attain a more accurate
ly recover after rain. The inrolled thalli picture ofthe contribution ofthe various
(flattened plant body) ofthese species had groups that make up soil crusts. However,
protective scales and were visible in some surveying overall crust cover in the drier
quadrats during the November 2003 sam- months is valuable in determining which
pling. Gongylantku$ scariosus, Lethocolea species are more tolerant to desiccation
pansa andEnigmella thallina all produce and, to observe the dynamic nature ofthe
spores in capsules that develop under the soil crusts.
soil in elaborated stem tissue (marsupia).
Acknowledgements
These marsupia persist in the soil after the This research was supported by funding
parent plant has shrivelled or decayed received from the Norman Wettenhall
(Beckmannand Scott 1980; 1992). Foundation. We thank Alain Braithwaite,
Mosses and lichens also took advantage Ranger, Little DesertNational Parkforhisassis-
of the availability of moisture (Fig. 4c). tance, Little Desert Lodge for allowing us to
Moss cushions ofRosulabryumcamylothe- work on their property and Anneke
Veenstra-Quah for herassistance with the map.
cCiaummpy,/Boaprubsuliantcraolfylceixnuas,alBl.schroiwnietda eavnid- I1n0v0e2r6t5e8b.rates were collected under permit No.
dence ofnew growth. These species pos-
sess morphological characteristics (e.g. References
Beckmann K.G (1993) The biology and taxonomy of
hyaline (colourless) leaf tips, leaf cell some Australian marsupial liverworts. (Unpublished
papillae(thickeningson cell wall), twisting PhDthesis.MonashUniversity).
aannddrToolzleinrg1o9f9l6e)avtehsa)t(eSncaobtlte1t9h82e,mEtlodtroildegre- BesAcucskatmrriaaolnsinau.sKL(iGLnedahbnemdr.g)SicaSott1te5,pGh7aA9n-iM84.-(1a98l9i)veGrownogrytlnanetwhutso
ate arid and semi-arid environmental con- Beckmann KGand ScottGAM (1992) A newthallose
ditions. During dry conditions, moss cush- genusofleafy liverwort from Australia. Journalof
ions often are partially inundated by sand BeBlrnyaoploJgy(21070.12)97M-i3c0r5o.besand microfauna associated
and brown in colour(Fig. 4a). Aftersignif- with biologicalsoilcrusts. InBiologicalSoilCrusts:
icant rain, cushions rehydrate and growth SEtdrsucturBee,lnFaunpctainodn,OanLdLMaanngaeg.em(eSnptr,ingpepr-1V6e7r-l1a7g4:.
begins (Fig. 4b). Recruitment of new Berlin.1)
plants was particularly evident in DSE (2006) Department of Sustainability and
Einccornelymifdoiuurmqusap.d,rawthsicdhurhiangdtbheeenNorveceomrbdeerd ElEBdniroivldiogrgeoin1c)mael(n1st9o9iw8lea-b)csriLutsietc,hpevrniosetweaecntddor3msoJusolsfyets2h0ea0ns6doill.ivNearwtourrtasl.
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Desert was low. This tends to confirm the Eld1r8i,d9g-e24D.(2000)Ecologyandmanagementofbiolog-
observation that, as soil crusts are dry and ical soilcrusts:recentdevelopmentsand futurechal-
inhospitable for much ofthe year, there is lenges.TheBryologist103,742-747.
unlikely to be a suite of invertebrates EldInriBdigoeloDgi(c2a0l0S1o)ilBiCorluosgtisc.alStsrouicltucrreu,stsFuonfctAiuosnt,ralainad.
specifically inhabiting the soil crusts. Management, pp 119-131. EdsJ Belnap and O L
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the soil crust as habitat. The results from EltdorrisdgereDlJatainndgTtoozetrheMdEis(t1r9i9b7uat)ioEnnviorfontmeernrtiaclolloaucs-
this preliminary study support the conclu- bryophytesandlichensinsemi-arideasternAustralia.
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A
pictorial representation ofperistomal architecture
Chris Tyshing and Maria Gibson
PlantEcologyResearchUnit,SchoolofLifeandEnvironmentalSciences
DeakinUniversity,221BurwoodHighway,Burwood, Victoria3125
Abstract
The terminology associated with the use ofperistomes in the identification and classification of
mosses is cumbersome and difficult to understand. This paper provides a pictorial explanation of
peristomal architecture with its associated terminology, such as nematodontous and arthrodontous
peristomes, and the division ofthe latter into diplolepideous and haplolepideous peristomes. (The
VictorianNaturalist 123(4),2006,203-211
The moss plant normally seen and recog- hygroscopic movement in response to
nised is referred to as a gametophyte as it changes in humidityby bendingbackwards
produces the gametes, i.e. egg and sperm. and forwards (Proctor 1984). The move-
When the sperm fertilizes the egg a sporo- ment provides a gentle catapulting action
phytedevelops. Thesporophyteisephemer- for launching spores a short distance into
al and essentially remains dependent on its the air, where they may be caught by a
gametophyte parent (Fig. 1), i.e. nutrients gentle breeze and dispersed to an environ-
are obtained from the gametophyte parent ment suitable for germination. Subsequent
through the basal foot ofa stalk-like struc- to germination, spores will develop into
ture (the seta) that remains embedded with- another gametophyte generation.
in the parental gametophytetissue. A spore Hygroscopic movement of the exostome
capsuleterminatesthisseta(Fig. 1). may be particularly relevant in closed for-
Many mosses have one or more rings of est situations where opportunities for air-
teeth around themouthofthe capsule (Fig. transport ofspores needs to be maximized.
2). The teeth collectively are referred to as The inner ring ofteeth (endostome) (Fig.
the peristome (Fig. 1) and are protected by 2) may regulate spore dispersal bygradual-
an operculum or lid (Fig. I), which falls ly siftingthespores.
offwhen the spores are mature. However, As spore dispersal mechanisms in moss-
notall mosseshaveperistomes. es,peristomes are specialised, intricate and
The outer ring of teeth (exostome) in architecturally elaborate. Adaptive trends
double peristomes (Fig. 2) may exhibit ofmorphological characters have resulted
Vol. 123 (4) 2006 203