N.Z. Journal of Marine and Freshwater Research 11 (2) : 357-73. June 1977

FRANCES B. MJCHAELIS*

Cawthron Institute, Nelson, and Zoology Department, University ofCanterbury, Christchurch, New Zealand

ABSTRACT

The flora and fauna of Pupu Springs (40°51'S, _172° 46'E) and five otherNew Zealand cold springs are described. In Pupu Springs there are 16 species ofalgae, 10 species of bryophytes (including three species of liverworts), and 5species of angiospcrms. The fauna includes a phrcatic form (an eyeless planarian,Dugesia sp.), a possible glacial relict (the caddis fly Rakiura vernale), and coldstcnothcrms (e.g., the caddis fly Psilochorema tautoru). The most abundantanimals in the New Zealand springs are Mollusca, Crustacea, and' larvae ofPlccoptera, Diptera, and Trichoptera.

Pupu Springs consist of five biotopes and associated biocoenoses.

INTRODUCTION

Although a limited amount of work has been done overseas, in NewZealand studies of the flora of cold springs are limited to a limnologicalsurvey of Western Springs, Auckland, by Johnstonc (1972); the onlypublished account of the fauna of cold springs in New Zealand is thatof Marshall (1973), who studied the Avonhead Springs, Christchurch.

Pupu Springs, at Takaka, NW Nelson, South Island, the largest coldsprings in New Zealand, and five other cold springs were included inthis study: Western Springs, Auckland; Hamurana Springs, Rotorua;Otangaroa Springs, Pularuru (all in the North Island); Three Springs,at Kimbell near Fairlie; and the spring at Lake Hayes, Queenstown(both in the South Island). The species of plants and animals presentin the springs were recorded, and, at Pupu Springs, their distributionswere investigated to determine whether different assemblages wereassociated with different substrates and water velocities. For physico-chemical details and locations of all these springs, see Michaelis (1976a).

METHODS

Preliminary studies were made at Pupu Springs on five dates betweenJanuary and October 1970, followed by regular sampling at least onceevery 4 weeks from October 1970 to February 1972. The five other coldsprings were visited during 1970-72.

"•Present address: c/o G. G. Wells, P.O. Box 147, Rotorua, New Zealand.Received 17 October 1975; revision received 31 May 1976.

358 N.Z. JOURNAL OF MARINE & FRESHWATER RESEARCH [JUNE

TABLE 1—Substrate, water depth, relative water velocity, and major species ofplants at sampling sites in Pupu Springs shown in Fig. 1.

Site Substrate Depth(m)

Velocity

Boulders 0.6

2 Bedrock 4.33 Vertical concrete 0.3

Boulders 0.6

Moderate

SlightModerate

Strong

Major Plants

5

6

7

89

10

Gravel

Gravel

Silt and water

GravelGravelBoulders

0.5

2.5

<0.8

2.56.50.5

Strong

Strong

Very slight

SlightModerateVery strong

Cratoneuropsis relaxaFissidens rigidulusSpirogyra sp.Cratoneuropsis relaxaCratoneuropsis rela xaFissidens rigidulusLophocolea austrigenaL. minorNeesioscyphus phoenicorhizMSCyathophorum bulbosumJuncus microcephalusSynedra ulna (summer only)Achnanthes sp. (summer only)Myriophyllum elatinoidesSynedra ulna (summer only)Achnanthes sp. (summer only)Nastiirt ium in icrophylliimLemna minorNasturtium microphyllumNasturtium microphyllumEntophysalis rivu/arisHUdenbrandia rivuiaris

COLLECTION OF SPECIMENS

At the six cold springs studied, collections for identification ofbryophytes, angiosperms, and associated fauna were made by hand eitherwhile wading or while scuba diving. Algae were preserved in eitherLugol's iodine or 3% formalin, higher plants were preserved in glycerol,by a technique modified from Johansen (1940), and invertebrates werepreserved in 70% ethanol. Insect and. mollusc eggs were hatched andadult insects were reared from larvae and pupae for identification. When-ever possible, adult insects were taken by sweep netting around the edgeof the springs.

VEGETATION MAP OF PUPU SPRINGS

The areas of all plant species except Nasturtium microphyllum wereconstant over the study period, although there were seasonal changesin growth form (Michaelis unpublished 1974). A vegetation map wascompiled from data gathered on 4 April 1970, 28 July 1970, 5 December1970, and 29 October 1971. Mapping was by direct underwater observa-tion, by measurement along transect ropes, and by photographs ofemergent watercress beds (Michaelis 1976b). The final vegetationmap was superimposed on a contour map of the springs, so that depthdistribution and maximum depth of each plant species could be seen.

1977] MICHAKLIS—PUPU SPRINGS BIOLOGY 359

FIG. 1—Pupu Springs, showing regular sampling sites (boxes 1-10), details ofwhich are given in Table 1. Depth contours at 2 m intervals indicated bydotted lines. Lines joining X—R. and X—S indicate representative transects(Fig. 3).

QUANTITATIVE SAMPLING AT PUPU SPRINGS

Sites for sampling plants and animals were chosen away from theboundaries of plant species, and were sampled regularly betweenSeptember 1970 and December 1972 by scuba diving, snorkelling, orwading (Fig. 1, Table 1). A 0.42-mm-mesh, 0.09 m2 Wisconsin trap(Welch 1948) was used to contain bare and bryophyte-covered bouldersand to sample submerged and emergent rooted, angiosperms. A Besssampler (Hess 1941) was used to sample floating angiosperms.

None of the quantitative data for plants is presented here, but theplant samples were used to provide quantitative animal samples. Within24 h of collection, animals were washed from, the living plant sampleswith tap water, and passed through a series of three sieves (smallest meshsize 0.42 mm).

Animals were sorted, into taxonomic groups and preserved in 70%ethanol for later sorting to species. Because Potamopyrgus antipodarumwere often present in large numbers, their dry weight was determinedand. converted via a factor to numbers per sample; these gastropodswere also counted underwater on bare, unstable sand and gravel on11 March. 1971. For each species of invertebrate, results were expressedas mean number of individuals per square metre, and 95% confidenceintervals were calculated.

BlOTOPES

Species of plants which covered a significant area and were restrictedto a given substrate and water velocity type were termed characteristicspecies (the 'exclusive' species of Poore 1955). Other species which were

360 N.Z. JOURNAL OF MARINE & FRESHWATER RESEARCH [JUNE

not restricted were termed 'indifferent' or 'casual' species (Tansley1946). Animals were grouped according to Thorup (1966). Character-istic species of animals were present in the favoured area with a numericalabundance of at least 1000 per square metre but of less than 10 persquare metre in other areas. Numerous species were present in thefavoured area at an abundance of at least 1000 per square metre, lessnumerous species at least 100 per square metre, and rare species at lessthan 100. Where the plant or animal species present on a substrate typein a particular water velocity included characteristic species, the substratetype and water velocity was termed a biotope (Dahl 1908), and theplant and animal species constitute a biocoenosis (Mobius 1877, citedby Hutchinson 1967).

RESULTS

TERRESTRIAL VEGETATION

Remnant stands in the Waikoropupu Valley indicate an originalcover of podocarps and hardwoods, including Nothofagus (beech). Thiscover was removed before the earliest photographic record of thesprings (about 1904). By then, Leptospermum had colonised the areaaround both springs. There was an extensive growth of the fern Blechnumcapense between the Main Spring and the Dancing Sands. Flax andcabbage trees were present, and a native rush, June us section Genuini(?/ . gregiflorus - det. E. Edgar) was just downstream of the DancingSands, but is no longer present. Gold mining in the area between thesprings and Fish Creek about 1900 resulted in mine tailings still visibletoday.

The main species of terrestrial plants now found at the springs andalongside the outflow are listed in Appendix 1; the native species arefound throughout New Zealand in lowland habitats. Native podocarp-hardwood forest now forms a dense cover on the island between thesprings, and to the cast of both the Dancing Sands and the springsoutflow. Shading of the springs by trees is negligible except near thisisland, and very little leaf litter enters the springs. Manuka and kanuka,gorse, and scotch broom form scrub to a height of 8 m around parts ofboth springs and along the west side of the outflow. Some of the shore-line is in grass for access and recreation, and flax and cabbage treesare no longer common. Dairy cattle are grazed on pasture to the westof the Main Spring and outflow and, at times, have access to the entiresprings area.

AQUATIC VEGETATION

ALGAE: Sixteen species of probably cosmopolitan algae were recordedfrom Pupu Springs (Table 2). The most common of these algae haveall been recorded in hard freshwater springs overseas, e.g., Batracho-sperrnum (by Round 1965), Hildenbrandia (by Botosancanu & Negrea1961), cyanophytes (by Prescott 1969), Achnanthes and Vaucheria sp.(by Whitford 1956). Chlorophytes and xanthophyceans are less com-monly recorded elsewhere. No planktonic algae were seen in the springs.

1977] M I C H A E L I S — P U P U SPRINGS BIOLOGY 361

TABLE 2—Species of algae recorded from Pupu Springs, January 1970-January 1972(*det. F. Drouct)

CYANGPHYTA DIATOMACEAE*Entophysalis rivularis Kutz. Achnanthes sp.*Nostoc parmeiiodes Kutz. Cocconeis placentula Ehr.Nostoc verrucosum (L.) Vauch. Cymbella sp.Microcoleus'! sp. Gornphonema sp.Oscillaton'al sp. Naviciila sp.

Synedra ulna (Nitzch.) Ehr.CHLOROPHYTA

Chaetophora elegans (Roth.) AgardhSpirogyra sp. (sterile)

CHRYSOPHYTA RHODOPHYTAXANTHOPRYCEAE Batrachospermum sp.

Vaucheria sp. (sterile) Hildenbrandia rivularis IJebm.

BRYOPHYTES: There were seven species of moss and three speciesof liverwort in Pupu Springs (Table 3). In these native bryophytes,Sainsbury (1955) discerned an endemic element, e.g., Hypopterigiumfiliculaeforme; a subantarctic clement, e.g., Fissidens rigidulus; a cosmo-politan element, e.g., Drepanodadus aduncus; and an Australianelement, e.g., Cyathophorum bulbosum. Sainsbury states that some ofthese mosses, e.g., Cratoneuropsis relaxa, are often found growing insprings and. streams, but others found submerged in Pupu Springs areusually found only on damp rocks, e.g., Cyathophorum bulbosum. Theliverworts Lophocolea spp. and Neesioscyphus phoenicorhizus werewell developed over large areas of the springs, although it is rare forliverworts to be important members of submerged communities inrunning waters (Hynes 1970).

Aquatic mosses and liverworts were not common in the other NewZealand cold springs, but the mosses Cratoneuropsis relaxa, Drepano-dadus spp., and Fissidens rigidulus were each found in two of the springsstudied (Table 3).

ANGIOSPERMS: Three of the five species of angiosperm recorded inPupu Springs were introduced to New Zealand (Table 3). Native speciesof Myriophyllum and Potamogeton were found in several of the springs,and Lemna minor occurred in all the springs that had areas of calmwater. Of the introduced species, Callitriche stagnalis grew in most ofthe springs listed, but Salvinia and Lagarosiphon major were restrictedto the North Island springs. The most widespread plants were theintroduced watercresses, Nasturtium microphyllum in the South Island,and N. offwinale in the North Island. Most emergent or submergedangiosperms were firmly attached to the substrate in moderate to verystrong water velocities, but floating-leafed plants were restricted to calmwater. No submerged angiosperms grew in the springs at Lake Hayes orin Three Springs, as the water was too shallow.

362 N.Z. JOURNAL OF MARINE & FRESHWATER RESEARCH [JUNE

TABLE 3—Species of aquatic plants, excluding algae, recorded from six cold springs inNew Zealand on the dates indicated and grouped by growth forms (some data forWestern Springs from Johnstone (1972); * — introduced species, X = present, - =not found)

Species ^a\<^

ni m m m m'P «* « c5 < " w -b o w /S p *t dc*^A c« H O ^

SUBMERGED Mossi-:sAcrocladium cuspidatum (Hedw.) Lindb.Cratoneuropsis re/axa (H.f. & W.) Fieisch e BrothCyathophonan butbosum (Hedw.) C M .Drepanoclodus aduncus (Hedw.) Warnst.D. fontinatiopsis (CM.) Dix.iuiu'nodiurn hispidum (\\S. & W.) Jaeg.Fissidens rigidulus C M .Hypopterygium ftlie ulacforme (Hedw.) Brid.Thamnium pandum (H.f. & W.) Jaeg.Thiudiopsis furfurosa (H.f. & W.) Fieisch

SUBMERGED LIVERWORTS

Lophocolca ausivigena (H.f. & T.) G.L. & N.L. minor NeesNeesioscyphus phocnicorhizus R. GrolieRiccai'dia sp.Riccioearpus natans (L.) Corda ap. Opiz

SUBMERGED ANGIOSPERMS

*Lagarosiphon major (Ridley) MossMyriophylium elatinoides Gaud.M. propinquum A. Cunn.Potamogeton sp.

EMERGENT ANGIOSPERMS

*Junctts microcepltalus H. B. K.* Nasturtium niicrophyllunt (Boenn.) Rchb.^Nasturtium offieinale R. Br.

FLOATING-LHAFI-;D ANGIOSPERMS

*Ca!!itriche xtagiialis Scop.Poiamogeion cheesemanii A. Bennett

FREE-FLOATING ANGIOSPERMS

Lemna minor L.*Safvinia sp.

DISTRIBUTION IN PUPU VSPRINGS: The distribution of bryophytesas a group and of each species of angiosperm was clearly defined, andthere were no transition zones between them. The map of aquaticvegetation (Fig. 2) shows that Pupu Springs is a mosaic of plant species.This distribution of plants reflects the varied substrate and water velocitytypes within the springs, and is possibly modified in places by lightintensity.

In Pupu Springs, angiosperms cover a greater area than bryophytes.Lemna minor, Juncus microcephalus, emergent Nasturtium micro-phyllum, and Myriophylium elatinoides all cover significant areas (11%,

1977] MICHAELIS—PUPU SPRINGS BIOLOGY 363

Wi mosses/liverworts

B Cailitriche stagnaiis

1 Juncus microcephakis

<m Lemna minor

m Myriophylhim elatinoides

•:•:• Nasturtium niicrophylhim (emergent}[!: Nasturtium micrpphyik/m (submerged)

No symbol denotes absence of higher plants

FIG. 2—Pupu Springs, showing aquatic vegetation. Depth contours at 2 m intervalsarc indicated by dotted lines. The maximum area covered by emergentNasturtium microphylum during the study period is shown, as mapped on23 July 1970.

7%, 16%, and 16% respectively of the total area of the springs). Themoss Cratoneuropsis relaxa (9%) and "liverwort" grouping of Lopho-colea spp., Neesioscyphus phoenicorhizus, and Cyathophorum bulbosum(about 19%) cover a total of 28% of the area of the springs. A total of501 m2 (28% of the area of the springs), generally where the substrateis unstable, is devoid of bryophytes or angiosperms.

Figure 3 shows representative transects across Pupu Springs.

ANIMALS PRESENT

PUPU SPRINGS: The aquatic animals found in the springs during thestudy are listed in Table 4. No attempt was made to identify Protozoa.Of about 46 aquatic species recorded, nearly half were insects. Threespecies have been introduced to New Zealand: Lumbriculus variegatus,Lymnaea columella, and Salmo trutta. The plecopteran Zelandopeiiaagnelis was recorded by McLellan (1967) from Pupu Springs, but hisspecimens came from nearby Fish Creek (D. A. Craig, pers. comm.),where it was taken in the present study also.

Several of the invertebrate species are undescribed, including a newgenus in the Orthocladiinae (Diptera) and probable new species ofDugesia (Tricladida), Tryssalurus and Zelandobates (Acarina), and

364 N.Z. JOURNAL, OF MARINE & FRESHWATER RESEARCH [JUNE

"3

-o g3 c

Cmo I

o

(HI) L|]d0Q

1977] MICHAELIS—PUPU SPRINGS BIOLOGY 365

Hydrobiosis and Oxyethira (Trichoptera). The trichopteran Rakiuravernale McFarlane, 1973 was described as a result of the collection ofadults during this study (Michaelis 1973).

OTHER SPRINGS: The faunas of the other five cold springs were similarto that of Pupu Springs (Table 4). The only species found in every coldspring was the widely distributed Potamopyrgus antipodarum (Gastro-poda), but other species found in at least three of the six springs wereParanephrops planifrons, Xanthocnemis zealandica, and Hydrobiosisparumbripennis. The vertebrate fauna in Hamurana Springs included theintroduced rainbow trout Salmo gairdneri.

DISTRIBUTION IN PUPU SPRINGS: The total numbers and speciescomposition of invertebrates varied between sampling sites, and many ofthe differences between sites could be related to substrate type and watervelocity.

The presence of higher plants was correlated with an increase in thetotal number of invertebrates (Tables 1 & 5). A substrate of boulderswith bryophytes (Sites 1 and 4) had more invertebrates than a similarsubstrate at a similar depth without bryophytes (Site 10). The fauna ona substrate of gravel with angiosperms in shallow water (Site 5) wasmore abundant than the fauna on a similar substrate that was toounstable for the growth of angiosperms (Michaelis unpublished 1974).The fauna was slightly more abundant on bryophytes at Site 4 than ondifferent species of bryophytes at Siie 1. Differences in water velocityrather than the presence, or absence, or the specific composition of higherplants may have accounted for some of the differences in invertebratesbetween these sites.

Although there were differences in the total abundance of invertebratesat different sites in water less than 1 m deep (Table 5), the proportionsof animals from each taxonomic group were relatively constant. At allsites in shallow water, Potamopyrgus antipodarum comprised 88-96%of total invertebrates, except at Site 10 (56%). At all sites in shallowwater, except Site 10, the order of numerical abundance was

Potamopyrgus > Other invertebrates > Insects(minus insects)

whereas at Site 10 it was

Potamopyrgus — Insects > Other invertebratesReduced numbers of P. antipodarum and other invertebrates exceptinsects at Site 10 may be related to the strong water velocities and lackof higher plants at that site.

Total invertebrates decreased with an increase in depth from 0.6 mto 6.5 m (Table 5). (Site 10 is disregarded as it is the only samplingsite without higher plants.) On the same species of plant, abundancedecreased with increasing depth of water, e.g., animals on Nasturtium,microphyllum were 10 times more abundant at the water surface (Site 7)

366 N.Z. JOURNAL OF MARINE & FRESHWATER RESEARCH [JUNE

TABLE 4—Species of aquatic animals collected in Pupu Springs and five other New Zealandcold springs (excluding spring outflows) on the sampling dates indicated (some datafor Western Springs taken from an unpublished 2-page set of class notes "The macrofaunaof Western Springs" issued by Zoology Department, Auckland University about 1962,and from Johnstone (1972) and Pullan et al. (1972); * == introduced species; t = singlespecimen only; X = present; - = not recorded)

Species g a s a& p s . * s * ~p s ;

P - «j w 5 ^ w •= ^ _ Z ^ ^o dot^ h 3 co

(£ w H O K ?

PLATYHELMINTHESTemnocepliala novaezelandiae HaswellCur a pwguis (Weiss)Dugesia n.sp.l ?Dugesia n.sp.2

ROTIFER ALecane''! sp.Flosculanidae (1 genus)

MEMATODADorylaiminae (2 genera)Chromadoridac (1 genus)

ANNELIDAHaplotaxidae

*Lumbi'iettlus variegattis (Miiller)Oligochaeta

TARDIGRADA (I genus)

CRUSTACEAOSTRACODA

Herpetocypris pascheri BrehmUnidentified Ostracoda

COPEPODA

Tropocyclops pras'uuis (Fischer)

AMP HI POD A

PavacaUiope kariiane BarnardParaleptamphopus subterraneus (Chi I ton)Paraleptamphopus n.sp. 7

DECAPOD A

Pavaiya curvirosti'is (Heiier)Paranephrops planifrons White

ACARINATryssaturus n.sp.?Zelandobates n.sp.

INSECT AODONATA

Xanthocnemls zealandica McLachlan

EPHEMEROPTKRA

Deleatidii m myzobvanchia PhillipsDfleatidium sp.Zephlebia deniaia (Eaton)Zephk-bia scifa (Walker)Zephlebia verslcolor (Eaton)

1977] MICHAELIS—PUPU SPRINGS BIOLOGY 367

82-

Ills

Mefuik'p toper la diminuta KimminsSienopei'lci prasina (Newman)Zeiandobius furcillatusi TillyardUnidentified Plecoptera

Hydora sp.

PTKRA

Unidentified Chironomidaetihivononnts Zealandicus HudsonMaoridiamesa liarrisi PagastOrHiocladiinae(tphydridae (i species)IAiiiiiophora sp.•StraUomyidae (1 species)Unionia sp.

ICMOPTItRA

licophidaetelotessica vhelva McFarlanelicopsychidae'lelicopsyche poutini Mci'arlane'iakiura vernale McFarianedropiilidae).xycfhira n.sp.)xyeihira. spp.in identified HydroptilidaeitoceridaeSudsonenia amabilis (McLachlan)''ripleciides obsoleta (McLachlan)yce nt ro po d id aeJolyplectroptis puerilis (McLachlan)yacophih'daelydrobiosis parumbripeunis McFarlailydrobiosis n.sp.?ysilochorema lautoru McFarlane^silochorema Ibidens McFarlaneICOSTOMATIDAi:• ointxia gtmni (McFarlane)teconesus sp.Hinga feredayi (McLachlan)'yvnocentria e.vecta McLachlan'yenocentrodes sp.Jnidentified Sericostomatidae

[ympluda nilens Butler

H.LUSCAyinnaca columella Say'hysa and/or Physastraotamopyrgus antipodai'iim (Gray)phaerium novaezelandlae Deshayes

iiguilla ausfralis schmidtii Phillipsntfiiilla dieffenbachii Gray'nidentified eelsahuo trulla L.ahno gairdfieri Richardsonhdypnodon breviceps Stokeil

46 10

368 N.Z. JOURNAL OF MARINE & FRESHWATER RESEARCH [JUNE

TABLE 5—Mean numbers of the snail Potamopyrgus antipodarum, insects, and other invertebratesper square metre at nine sites in Pupu Springs (Fig. 1), arranged in order of increasing depth(for raw data see Michaelis (unpublished 1974); 95% confidence intervals given in parentheses;- = not calculable; e — estimated)

Site

1051476829

Depth(m)

0.50.50.60.60.82.52.54.36.5

Potamopyrgus

225

e 4 0e 180

e 3 02224

900000000000000500500000000

(±1 700)(-13 000)

_

(+•500)( + 360)( + 3 200)(±390)

Insects

1 500 ( + 650)370 ( + 470)

4 700 ( + 3 900)1 800 ( + 1 000)

200 ( + 74)150(490)16 ( + 23)25 (-1-23)9 (±6)

215I1

Others

900 ( + 290)300 (1-810)900 ( + 990)500 (+2 700)800 ( + 750)800 ( + 317)680 ( + 190)220 (+350)530 (-1-90)

52846

190324324

Totals

200 (+7000 ( + 7000 -000000 -400 (+2

700)400)

200)200 ( + 520)300 ( I K500 ( + 1

) 000)200)

than at depths 2.5 m and 6.5 m (Sites 8 and 9) and, on Cratoneuropsisrelaxa, were 20 times more abundant at depth 0.6 m (Site 1) than atdepth 4.3 m (Site 2). All the major invertebrates decreased in abund-ance with an increase in depth, e.g., numbers of Potamopyrgus anti-podarum decreased at least 10-fold (Table 5) as did numbers of otherinvertebrates excluding insects (i.e., mainly crustaceans). The decreasein numbers of insects was even more marked.

Some species of invertebrates were widespread within the springs.Four main substrate and water velocity types had an abundant faunaand accounted for about 75% of the springs area; the remaining areawas unstable sand and gravel (Table 6). Common invertebrates on atleast three of these four types were four crustacean species, five insectspecies (including three trichoptcrans, and the mollusc Potamopyrgusantipodarum).

Some species had restricted, distributions within the springs, and thuscharacterised three of the biotopes discerned.

BIOTOPES

The major substrate types and water velocities in the study area maybe divided into biotopes and associated biocoenoses as listed in Table 6.

Biatope 1 had no characteristic plant species; the algae Hildenbrandiarivularis and Entophysalis rivularis, dominant in this biotope, were alsofound in Biotope 2. The biotope was characterised by the absence ofbryophytes and angiosperms and by a moderately abundant fauna.

Biotope 2 had seven characteristic species of plants, of which six werebryophytes and one was an epiphytic green alga. There were also twoindifferent species of algae (see Biotope 1) and another four species ofbryophytes which were casual species in this biotope. The fauna onboulders in shallow water was very abundant, and there were twocharacteristic species. No characteristic species of animals were associ-ated with bedrock and slight water velocity (Table 6), perhaps because

1977] MICHAELIS—PUPU SPRINGS BIOLOGY 369

such bedrock was found only at a depth of 4.3 m: the number of speciesdecreased with depth throughout the springs.

Blotope 3 had three characteristic angiosperm species, two of whichwere introduced, to New Zealand, but had no characteristic species ofanimals, although it supported a moderately abundant fauna of specieswidespread in the springs.

Biofope 4 had no characteristic plant and animal species and the faunawas not abundant.

Blotope § had two characteristic species of plants of which Nasturtiummicrophyllum was dominant. Associated with this was an abundantfauna, including four characteristic species of invertebrates of which twowere introduced.

DISCUSSION

Many of the invertebrates found in these New Zealand cold springsbelong to genera endemic to New Zealand, e.g., Paranephrops; Zephlebia,Megaleptoperla, Conuxia, Psilochorema, Hydrobiosis, Rakiura, andZelolessica, which belongs to a family, Helicophidae, endemic to Aus-tralia and New Zealand.

.Most of the aquatic species in the cold springs are widely distributedin running waters in New Zealand, e.g., Lumbriculus variegalus, Xantho-cnemis zealandica, Conuxia gunni, Hydrobiosis parumbripennis, Zelo-lessica cheira, and Potamopyrgus antipodarum. Other species found inPupu Springs may be confined to Nelson, Marlborough, the west coastof the South Island, and. to the North Island, e.g., Paranephrops plani-frons (see Hopkins 1970); or to Nelson and the west coast of the SouthIsland only, e.g., Helicopsyche poutini and Oxyethira n.sp. (A. G.McFarlane, pers. comm.) or may occur only rarely east of the SouthernAlps, e.g., Psilochorema taut or u (A. G. McFarlane, pers. comm.).

Some species are found in the cold springs during part of their lifehistory but spend the remainder downstream in estuaries or in the sea,e.g., Paratya curvirostris and Anguilla spp. Paracalliope karitane(Amphipoda: Eusiridae), known only from Pupu Springs and from thelower reaches of the Hutt River, Wellington, may be such a species(Barnard 1972).

Some species occur in cold springs far outside their normal geographicrange and may be classed as relicts. Rakiura vernale (Trichoptera:Helicopsychidae), found in Pupu Springs, appears to be a glacial relict(Michaelis 1973).

Some of the animals in the cold springs studied originated fromgroundwater. Two such phreatic species were collected: a white, eyelessplanarian, Dugesia n.sp., known only from Pupu Springs, and anamphipod, Paraleptamphopus subterraneus, from Otangaroa Springs.Many overseas studies of cold springs have reported phreatic fauna,

o

TABLE 6—Biotopes and biocoenoses in Pupu Springs (* = species introduced into New Zealand)

fo.

a

b

Substrate

Bedrock andboulders

Bedrock

Boulders

Gravel

Unstable sandand gravel

Silt and water

BlOTOPE

WaterVelocity

Very strong

Slight

Moderate-strong

Slight-strong

Slight-strong

Very slight

Area(%of

springs)

26

23

16

Characteristic Plants

None

1 Spirogyra sp.1 Cratoneuropsis rclaxa| Cyathophorum hulbnsumYFissidens rigidulusLophocolea austtigena

I L. minorj Neesioscyphus phoenicorhiztis*Juncus nv'crocephahisMyriophyllum eJaiinoides

* Nasturtium ni.icrophyll.um(submerged)

None

* Nasturtium microphyllum(emergent and floating)Lerrma minor

BIOCOENOSIS

Characteristic Animals

Dugesia sp.Deleatidium myzohranchia

/None

f Conuxia gunni\Zelolessica cheira

None

None

*Lumhricidua variegatusHerpetocypris pascheriTropocyclops prasmus

*Lymna?a columello.

Numerous Animals

Potamopyrgus antipodarum

/Paro.call.iope karitane\ Potamopyrgus antipodarum

{ Paracalliope karitanej Polypiectropus pncriiis| Psilochorema ttiutoru\_Potamopyrgus antipodarum.

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Potamopyrgus antipodarum

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1977] MICHAELIS—PUPU SPRINGS BIOLOGY 371

including species of Planaria (e.g., Carpenter 1928), Amphipoda andSyncarida (e.g., Efford 1959), and Isopoda (e.g., Minckley 1961).

The fauna of the New Zealand cold springs includes at least twospecies of Trichoptera that are presumably cold stenotherms: Rakiuravernale (see Michaelis 1973), and Psilochorema tautoru, which isgenerally found over 300 m above sea level (McFarlane 1951) but isnow known from two cold springs near sea level: Pupu Springs and aspring at Hunt's Beach, Westland (A. G. McFarlane, pers. comm.).Many species of stenothermous invertebrates confined to springs andspringbrooks have been recorded in other parts of the world, e.g., thegastropod Bythinella dunkeri by Thienemann (1912), the trichopteranApatidea muliebris by Nielsen (1950), and the triclad planarian Creno-bia alpina by Arnold & Macan (1969).

The biotopes in Pupu Springs may be compared with those in threecold springs in Denmark (Lille Blaakilde, Ravnkilde, and Rold Kilde).There, Thorup (1966) recognised five biotopes, three of which aresimilar to those in Pupu Springs, namely, stony bottoms ( = Biotope 1),areas of moss ( = Biotope 2) and swampy border areas ( = Biotope 5).The remaining biotopes of Thorup's springs (hygropetric areas and areascovered in dead beech leaves) were not present in Pupu Springs, butcan be found in the nearby spring-fed Fish Creek. The latter biotopewould not be expected in the springs because the forest that oncesurrounded the springs has been cleared.

By contrast, some springs have a single plant and animal communityand can be considered a single biotope, e.g., Silver Springs, Florida, withits beds of eel-grass, Sagittaria lorata, and associated aufwuchs on asubstrate of gyttja (Odum 1957), and Cone Spring, Iowa, with sevenspecies of invertebrates very evenly distributed on a substrate of sandwith pebbles (Tilly 1968).

ACKNOWLEDGMENTS

This work was financed by the Cawthron Institute and forms part of a Ph.D.thesis in the Zoology Department, University of Canterbury.

I acknowledge the help of Miss H. M. Campbell, the owner of Pupu Springs;of Entomology Division, DSIR, Nelson, by providing constant-temperature rearingfacilities; and of Nelson Underwater Club Inc.

I thank the following for assistance with identification of plant and animalspecimens: K. W. Allison (bryophytes); Botany Division, DSIR; E. A. Flint(algae); E. A. Hodgson (Neesioscyphus phoenicorhizus); R. Mason (aquaticangiosperms) F. R. Allison (triclads); R. O. Brinkhurst (annelids); F. Climo(molluscs); T. K. Crosby, D. J. Forsyth, and P. M. Johns (diptcrans); J. S.Dugdale (dipterans, lepidopterans); A. C. Eyles (hemipterans); R. R. Forster(arachnids); D. E. Hurley and M. Lewis (crustaceans); A. G. McFarlane (tricho-pterans); I. D. McLellan (plecopterans); G. W. Ramsay (mites); J. C. Watt(coleopterans); M. J. Winterbourn (ephemeropterans); and W. M. Wouts(nematodes).

My husband, Geoff Wells, assisted me with great skill both on field trips and inthe preparation of the figures. Finally I thank Drs Vida M. Stout and M. J.Winterbourn for their help during the study and in preparing the manuscript.

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372 N.Z. JOURNAL OF MARINE & FRESHWATER RESEARCH [JUNE

LITERATURE CITED

ARNOLD, F. & MACAN, T. T. 1969: Studies on the fauna of a Shropshire hillstream. Field Studies 3: 159-84.

BARNARD, J. L. 1972: The marine fauna of New Zealand: Algae-living littoralGammaridea (Crustacea: Amphipoda). N.Z. Oceanographic InstituteMemoir 62. 216 pp.

BOTOSANEANU, L. & NEGREA, S. 1961: Une oasis aquatiquc a faune reliquc dansla plainc du Danube inferieur. Hydrobiologia 18: 199-218.

CARPENTER, K. E. 1928: "Life in Inland Waters". Sidgwick & Jackson, London.267 pp.

DAHL, F. 1908: Grundsatze und Grundbegriffe der biocSnotischen Forschung.Zoologischer Anzeiger 33: 349-53.

EFFORD, I. 1959: Rediscovery of Bathynella chappuisi Delachaux in Britain.Nature 184: 558-9.

HESS, A. D. 1941: New limnological sampling equipment. American Society ofLimnology and Oceanography Special Publication 6. 5 pp.

HOPKINS, C. L. 1970: Systcmatics of the New Zealand freshwater crayfishParanephrops (Crustacea: Decapoda: Parastacidae). N.Z. Journal ofMarine and Freshwater Research 4: 278-91.

HUTCHINSON, G. E. 1967: "A Treatise on Limnology. Vol. IT. Introduction toLake Biology and the Limnoplankton". Wiley, New York. 1115 pp.

HYNES, H. B. N. 1970: "The Ecology of Running Waters". Liverpool U.P. 555 pp.

IOHANSEN, D. A. 1940: "Plant Microtechnique". McGraw-Hill, New York.523 pp.

JOHNSTONE, I. M. 1972: Limnology of Western Springs, Auckland, New Zealand.N.Z. Journal of Marine and Freshwater Research 6: 298-328.

MCFARLANE, A. G. 1951: Caddis fly larvae (Trichoptera) of the family Rhya-cophilidae. Records of the Canterbury Museum 5: 267-89.

MCLELLAN, I. D. 1967: New gripoptcrygids (Plecoptera) of New Zealand.Transactions of the Royal Society of N.Z., Zoology 9: 1-15.

MARSHALL, J. W. 1973: A benthic study of the Avon Spring Stream, Christchurch.Mauri Ora 1: 79-90.

MICHAELIS, F. B. 1973: The distribution and life history of Rakiura vernale(Trichoptera: Helicopsychidae). Journal of the Royal Society of N.Z.3: 295-304.

unpublished 1974: "The Ecology of Waikoropupu Springs". Ph.D.thesis lodged in library of University of Canterbury, N.Z. 158 pp.

1976a: Physico-chemical features of Pupu Springs. New Zealand Journalof Marine and Freshwater Research 10: 613—28.

1976b: Watercress (Nasturtium microphyllum (Boenn.) Rchb. andN. officinale R.Br.) in New Zealand cold springs. Aquatic Botany 2-317-25.

MINCKLEY, W. L. 1961: Occurrence of subterranean isopods in the epigeanenvironment. American Midland Naturalist 63: 452-5.

1977] MICHAELIS—PUPU SPRINGS BIOLOGY 373

NIELSEN, A. 1950: Notes on the genus Anatidea McLachlan with descriptions oftwo new and possibly endemic species from the springs of Himmcr-land. Enlomologiske Meddelelser'25: 384-404.

ODUM, H. T. 1957: Trophic structure and productivity of Silver Springs, Florida.Ecological Monographs 27: 55—112.

POORE, M. D. 1955: The use of phytosociological methods in ecological investiga-tions. 1. The Braun-Blanquct system. Journal of Ecology 43: 226-44.

PRESCOTT, G. W. 1969: "The Algae: A Review". Nelson, London. 436pp.

PULLAN, N. B., CLIMO, F. M. & MANSFIELD, C. B. 1972: Studies on the distribu-tion and ecology of the family Lymnaeidac (Mollusca: Gastropoda)in New Zealand. Journal of the Royal Society of N.Z. 2: 393-405.

ROUND, F. E, 1965: "The Biology of the Algae." Arnold, London. 269 pp.

SAINSBURY, G. O. K. 1955. A handbook of the New Zealand mosses. RoyalSociety of N.Z. Bulletin 5. 490 pp.

TANSLEY, A. G. 1946: "introduction to Plant Ecology". Allen & Unwin, London.260 pp.

THIENEMANN, A. 1912: Der Bergbach des Sauerlandes. Internationale Revue derGesamten Hydrobiologie, Biologische Supplement 4. 125 pp.

THORUP, J. 1966: Substrate type and its value as a basis for the delimitation ofbottom fauna communities in running waters. Pp. 59-74 in Cummins,K. W., Tryon, C. A. and Hartman, R. T. (eds)~ "Organism-SubstrateRelationships in Streams." Pymatuning Symposium in Ecology 4.University of Pittsburgh.

TILLY, L. J. 1968: The structure and dynamics of Cone Soring. Ecological Mono-graphs 38: 169-97.

WELCH, P. S. 1948: "Limnological Methods". Blakiston, Philadelphia. 381pp.

WHITFORD, L, A. 1956: Communities of algae in springs and spring streams ofFlorida. Ecology 37: 433-42.

APPENDIX 1—Species of terrestrial plants (excluding bryophytes, lichens, and grasses)surrounding Pupu Springs, December 1971 (* = introduced species)

Species

FfLICOPSIDAPhymatodes diversi'foliumBh'chnum capen.seGleichenia cmwinghamii

GYMNOSPERMAEDacrydium cupressinumPodocat'piis tolara

ANGIOSPERMAEMONOCOTY LED ONES

Phormium tenctxCordyliiie oust red isCortaderia sp,

Position

ForestForestForest, scrub

ForestForest

ShoreShoreShore

Species

DlCOTY LED ONES

* Ranunculus acrisFuchsia excorticcilaPittosponim eugenioides ?Lepto.spernwm scopavlumLeptospermum ericoidesMetrosideros colensoiWeimnannia racemosa

* Utex eut'opaens*Cytisus scopariusNothofagus fuscaNothofciqus menziesiiPseudapanax crassifoHiimCvathodes fasciculataC yathodes /uniperinaCoprosma tenuicoulisCoprosma robustaCoprosma cunnirvzhamii

*Myosotis caespitosa

Position

ShoreForestI 'orestScrubScrubForestForestScrubScrubForestForestForestForestForestForestForest, scrubForestShore