PacificScience(1974), Vol. 28, No.4, p. 467-504 Printedin GreatBritain Algal Flora of Some North Island, New Zealand, Lakes, Including Rotorua and Rotoiti I VIVIENNE CASSIE2 FROM AN EXAMINATION over a 3-yearperiod of isms were counted and identified according to more than 370 samples, an accountis given of the method described by Cassie (1969). In the the chief components of the phytoplankton, 3rd year, net samples were examined. Vertical together with brief mention of neuston, auf hauls were made at selected stations from wuchs, and benthos communities. The specific bottom to surface with a tow net having a composition of algal classes and pollution 160-#mesh. Net samples provided apicture of tolerance ofdifferent species are discussed. biomass,whereas jarsamples gaveinformation From 1966 to 1969 lakes Rotorua and on abundance of smaller nannoplankton. For Rotoitipossessedarichandvariedalgalflorain counts from concentrated jar samples, cells which the same three species of diatoms pre rather than colonies were estimated, in an dominatedinfluctuatingamountsthroughallor attempt to give a precise picture of total pre part of each year. Chlorophyceae, particularly served phytoplankton. For net estimates, Chlorococcalesand desmids,werenumerousin colonies instead of single cells were taken as species but less prolific than diatoms in cell units for assessment of abundance. Samples numbers, except for planktonic Mougeotia. from Lake Ohakuri and other hydroelectric Blue-greenalgaeare knownto form noxious lakes were collected with a Mikrops tow net blooms at times, though no such blooms (50 meshes per cm), and those from Lake occurred within the sampling period. Dino Horowhenua were concentrated by settling phyceae and Cryptophyceae were relatively from jars containing 300 ml oflake water. unimportant. The present study was undertaken to deter mine the basic composition of the algal flora MORPHOMETRY comprising the phytoplankton and other com Lake Rotorua occupies a basin produced by munitiesinseveralNorthIsland,New Zealand, subsidence, whereas Lake Rotoiti fills a lakes. Ofthe370samples, 260camefrom lakes drowned v2.11ey. Both are affected by aerial top Rotorua and Rotoiti. In addition, 91 samples dressing and sewage effluent, though the effect were examined from the hydroelectric lakes ofeffluentin the latteris muchless thanin the constructed on the Waikato River (lakes former (Fish 1969b). Both are tectonic lakes, Ohakuri,Atiamuri, Whakamaru, Maraetai, and i.e., owe their origin to volcanic action. Hot Karapiro), and 22 from Lake Horowhenua springs occur at various places along their near Levin(Fig. 1 and Table 1). shores (but algae from these habitats have not beenstudiedinthissurvey).Bothlakesare86m above sea level. The drainage area into Lake METHODS Rotoiti (67 km2) is much less than that into During the first 2 years; 200-ml samples of Lake Rdtorua. According to the Ministry lakewaterfromlakesRotoruaandRotoitiwere of Works Kaituna Investigation Report, preserved in Lugol's Solution and the organ- 1965, the catchment area of Lake Rotorua is 526 km2• (Maximum recorded depth is 1 This work was supported by grants from the 44.5 m [Fish and Chapman 19691-) Lake Fisheries Research Division, Marine Department, and Rotoiti descends more abruptly to depths from the Botany Department, The University of rangingfrom40to70m andhasamaximumof Auckland. Manuscriptreceived4May 1973. 93.5 m in the center basin (Irwin 1969). The 2 DepartmentofBotany,TheUniversityofAuckland, PrivateBag,Auckland, New Zealand. waters of Lake Rotorua that flow into Lake 467 30 HPS 28 468 PACIFIC SCIENCE, Volume 28, October 1974 N o , 25 5,0 Miles FIG. 1. Southernpartofthe North Island ofNew Zealand, showing location oflakes studied and sequence of hydroelectriclakeson Waikato River, withitsorigininLakeTaupo. KEY: " L. Horowhenua; 2, L. Aratiatia; 3, L. Ohakuri; 4, L. Whakamaru; 5, L. Maraitai; 6, L. Waipapa; 7,L. Arapuni; 8,L. Karapiro; 9,L. Rotorua; 10,L. Rotoiti. Rotoiti via the Ohau Channel mix with those pasture farming; whereas only part of Lake of the latter only in the shallow basin at the Rotoiti is affected by farming, much of the western end, after which they flow down the surrounding drainage area being clothed in main outlet, the Kaituna River (Fish 1969b). bush. Jolly found the former to be usually Lake Rotorua has been classified by Jolly homothermous, whereas the latter, with some (1968) as being among those lakes directly stratification, was warm monomictic (cE. affected by their proximity to improved Hutchinson 1957). Algal Flora ofSome North Island Lakes-CAssIE 469 TABLE 1 SAMPLINGSTATIONSINLAKES ROTORUAAND ROTOITI DATE STATION DESCRIPTION LAKE ROTORUA May 1966-May1969 A offshore(OpenLakeStation; surface,9m, 17 m) B inshore(Kawaha Point; pooledsample) C inshore(MokoiaIsland; pooledsample) June 1967-May1968 A offshore(OpenLakeStation;surface, bottom) D offshore(Hamurana-MokoiaIsland; surface, bottom) June 1968-May1969 A offshore(OpenLake Station; bottomto surface) LAKE ROTOITI January-May 1967 Wrights Bay offshore(surface, 4m, 8m) (Te PohoeBay) June 1967-May1968 1 inshore, westend(surface,in 11 m) 2 offshore, oppositeWrights Bay(surface, in42m) 3 offshore, oppositeHauparuBay(surface,in59m) June 1968-May1969 3 offshore, oppositeHauparu Bay(bottomto surface) NOTE: ForlocationofRotoruastationsseeCassie(1969,fig. 1);forlocationofRotoitistations1-3seeFishand Chapman(1969,fig. 1). ENVIRONMENTAL FACTORS in mid-February 1966 (Irwin 1968). Maximum andminimumtemperaturesinLakeRotorua at Climate andTemperature a depth of1 m were found to exceed those at During the 3-year period, monthly rainfall the surface in November 1966 and May 1967. varied from 0.83 inches in March 1969 to Below 1 m there was also a decrease of up to 20.87 inches in August 1967. The warmest 1.0°C in temperature with depth; but the month also was March 1969 with 116 hours of decreaseneverexceeded1.0°Candatthemany sunshine. Lowest insolation (70.5 hours) bathythermograph stations there was no occurred in June 1968. There was no pro change. However, a decrease of 1.6° C was nounced rainy season, but hours of sunshine recorded by officers ofthe Marine Department were more numerous from January to March between surface and bottom at the O.L.S. on than in other months. October 1967 was 27 February 1967(Cassie 1969, table 3). notable for its warm air temperatures (4°C InLakeRotoitiIrwindetectedathermocline above normal) and low rainfall (1.4 inches between 10 and 20 m where temperatures below normal [N. G. Robertson, personal ranged from 20° to 16°C (a gradient 0.4° C communication]). per m). Hypolimnion temperatures ranged In Lake Rotorua surface water temperatures from 11.9° to 12.8°C. Jolly (1968) recorded at the open lake station (O.L.S.) were also surfaceandbottomwintertemperaturesinLake higher in November 1967 than ~t any time in Rotoiti as being 11.0°C and 10.5°C, respec the preceding and following years. Figures tively. In Lake Rotorua Irwin found that from thefirst sampling yeararegiveninCassie thermal stratificationwas ill-defined or absent; 1969, fig. 2. Subsequent data show that the whereasinLakeRotoitistratificationdidoccur. lowest temperaturewas 8.6° Con22 July 1968 and the highestwas 21.4°C on 12March 1968 ChemicalConditions (G. R. Fish, personal communication). Surface temperatures in Lake Rotorua The1955-1956valuesquotedbyJolly(1968) (measured by bathythermograph in daylight) for ammonia-nitrogen and phosphate were were found to vary from 13° to 14°C (2 to fairlylow; whereas nitratenitrogenwas hardly 4 May 1967) and varied from 22,50 to 24.5°C detectable in either lake. Silicate ranged from 30-2 470 PACIFIC SCIENCE, Volume 28, October 1974 TABLE 2 NUMBERS OF SPECIESIDENTIFIED FROMDIFFERENTALGAL CLASSES ANDHABITATS TOTALNUMBER PLANKTON AUFWUCHS BENTHOS IDENTIFIED Chlorophyceae 44 7 51 Xanthophyceae 2 2 Chrysophyceae 7 1 8 BaciIIariophyceae 8 11 5 24 Euglenophyceae 1 1 4 6 Dinophyceae 6 6 Cryoptophyceae 1 1 Myxophyceae 10 3 5 18 Total 79 22 15 116 2-5 mg/liter in Lake Rotoiti and from 1 DissolvedOxygen 6mg/liter in Lake Rotorua-Iow values for As with the surface waters of other New thermal waters (Jolly 1968: 240). Fish (1969b) Zealand lakes, those of Rotorua and Rotoiti reported that in Lake Rotorua during the last showed a high percentage saturation at most 2decades therehas beenatremendous incre2se seasons, especially in spring when there was a in the forces promoting eutrophication, due high rate ofphotosynthesis. Jolly (1968) ga:e particularly to land development, c~using the following figures to indicate the range 111 erosion and water runoff. The populatIon of percentagesaturationofdissolvedoxygenfrom Rotorua City has increased greatly, and surface to bottom in the two lakes: Lake phosphate-phosphorus and. nitrate-nitr~~en Rotoiti-surface 122-90, bottom 130-29; Lake havebeenretainedinthelakeInvastqu?ntltIes. Rotorua-surface 138-95, bottom 130-94. As a result the standing crop of organisms is During March and April in Lake Rot~iti, now much greater. minimal values for oxygen were recorded Just Data on the phosphate-phosphorus, nitrate below the thermocline. In the hypolimnion a nitrogen, and ammonia-nitrogen content of mean annual minimum of dissolved oxygen Lake Rotorua from June 1967 to May 1968 was estimatedbyJollyas being3.1 ppm. More (Fish1969b: table 1) showthatthe totalinflow recently, in 1967-1968, Fish(1969a) found the of these nutrients greatly exceeds the surface mean annual minimum in the hypolimnion to outflow.Themaximumamountofphosphate-P be no greater than 0.08 ppm. A comparison recordedbyFish(7,692 kg or22percent) came between the figures of Jolly (1968) and those from the Hamurana Stream. Nitrate-N was ofFish(1969a) showsthatminimalvalueswere most abundant from the Awahou Stream lower in 1967-1968 than in 1956-1957. In (30,583kgor28percent).Thebulkofa~monia 1956-1957 the dissolved oxygen in ppm in N (95,500 kg or 66.5 percent) flowed Into the Lake Rotorua ranged from 12.4 to 9.0 (Jolly lake through the Waiohewa Stream. The 1968);in1967-1968thosevalues were11.2-3.5 amount of phosphate-P and ammonia-N (Fish 1969a). entering the lake from sewage outflow during the same period was smaller (14.2 percent and 19.9 percent, respectively). No nitrate-N was COMMUNITIES P!?Jtoplankton recorded from this source. It seems likely, ac cording to Fish, that agricultural development Theappendixatthe end ofthiswork consti provides more nutrients that promote eutro tutes a list of all algae identified during the phication than does sew~ge inflow. F~rther, survey, together with their abundance on a during periods of flood111g the quantIty of 1-to-5 scale (Cassie 1961). The symbols p, a, nutrients in the lake can increase within a few and b indicate their habitat: whether phyto days by two or more orders ofmagnitude. plankton, aufwuchs, or benthos. Records for Algal Flora ofSome North Island Lakes-CAssIE 471 FIG. 2. Melosira grantllata Ralfs. Auxospore. Film FIG.3.PeridinicumstriolattlmPlayf.FilmS/12A. x1500. 14/24A. x2135. a and b are fragmentary. Table 2 indicates the phyceae, Dinophyceae, and Chrysophyceae distributionofspeciesamongthedifferentalgal seldom achieving ecological importance, at classes. The phytoplankton component is by leastinpreservedsamples.Ingeneral, no single far the largest, mainly because it was the phytoplankter dominated to the exclusion of principal object of study. Numbers of species all others. Over the 3-year sampling period in each category are therefore merely an thepatternofabundanceofthedifferentdomi interim guide to those actually present. nants and subdominants (5 and 4 on the Seventy-nineofthetotallistof116specieswere abundance scale in the appendix) did not vary found to lead a planktonic existence. These greatly. were not necessarily obligate phytoplankters, Fishand Chapman(1969) notedthat greatest but were the ones that most characteristically phytoplankton production in Lake Rotorua occupiedthathabitat.Withinthephytoplankton occurs in the northeastern quadrant where communitythe chlorophycean componentwas waters move toward the main outlet, Ohau the strongestonewhen numbers ofspecies are Channel. These workers have also published considered. Only rarely did green algae domi maps showing beds of submerged Lagaro nate, however. The most abundant were siphonandElodeathatoccuratintervalsnearthe Bacillariophyceae (Cassie 1969), with Myxo- western shore ofthelake. Lowercell counts in 472 PACIFIC SCIENCE, Volume 28, October 1974 samples from inshore western waters are con phytoplankton bloom. Very frequent were sistentwiththefactthatFitzgerald(1969)found rapidly moving uniflagellateChromulina sp., no an antagonism between aquatic macrophytes more than 6 to 12p,in diameter. Itis not con and competing phytoplankton. (Cf. Cassie fined to theneuston, however, since specimens 1969, who recorded lower cell numbers at havebeenobservedinfresh subsurfacesamples Kawaha Point.) It is interesting to note in from both lakes. Small flagellated chryso this connection that Mitchell (1971) found monads are known to become attached to the in Tomahawk Lagoon (no. 2) on the south surface film, surround themselves with an side of Otago Peninsula that large crops envelope, and orient their chromatophores at of either phytoplankton or macrophytes right angles to the incident light. When light were always present, though never both rays are concentrated on them, the cell acts as together. alensfromwhichthelightis reflected(Ruttner R. M. Cassie (personal communication) has 1963: 136). It could be that the phenomenon shownbystatisticalanalysis ofa detailedsetof observedonLakeRotoitiwasasimilarone,but samples taken over 24 hours that the phyto the exact species responsible for the present plankton population of Lake Rotorua is rela phenomenon was not identified. Although 12 tively homogeneous despite differences due to otherspecieswereobservedinthesample,none depths and stations. Melosira granulata (Fig. 2) was as abundant as the minute Chromulina. in particular shows a fairly consistent vertical Other species, including Anabaena flos-aquae, homogeneity in distribution. By comparison Melosiradistans,Mougeotiasp.(narrowfilaments), Jolly (1959) found in her net samples that Rhizosoleniaeriensis,Cystodiniumcornifax,Asterio Melosira was by far the most abundant phyto nella formosa, and Bot,:yococcus braunii are all plankter and Chlorophyceae were subordinate known to occur in subsurface phytoplankton. though several genera, including Mougeotia, (See Cassie 1969: figs. 4, 6, 8, 9.) Cosmarium, Dicryosphaerium, Spon4Ylosium(prob Other species common or occasional in the ably including Sphaerozosma), and Closterium, phytoplankton at various times of the year were conspicuous. All but the first two were included Microrystis aeruginosa, Anabaena circi representedbysmallnumbers. Jollyalso found nalis, Pandorina morum, Trachelomonas volvocina, Staurastrum, Dinobryon, and Peridinium (prob Dinobryon y€ lindricum, and Nephrorytium agardhi ably P. striolatum, Fig. 3) to be moreabundant anum(Figs. 4-9). in Lake Rotoiti. Asterionella was insignificant. Staurastrum and Staurodesmus were nearly ItappearsfromacomparisonofJolly'Srecords always present, though seldom in large with those of the present survey th3t, apart numbers; e.g., Staurastrum floriferum (Fig. 4), fromthescarcityofAsterionella,theproportion S. leptocladum var. insigne (Fig. 10), S. sexangu of most of the planktonic algae has remained lare (Fig. 11), and Staurodesmus unicornis V2!. approximately the same. Two chlorophycean ceylonensis (Fig. 12). Staurodesmus convergens var. genera not recorded by Jolly but which laportei (Fig. 13) has been found to date only achieveddominanceattimes between1967and in the "hydroelectric" lakes on the Waikato 1969 were Actinotaenium pyramidatum and RiverandinLakeRotoma(nearLakeRotoiti). Actinastrumhantschii. Cyclotella stelligera (Fig. 14), though dominant A single fresh neuston (surface film) sample once in Lake Rotorua, was usually more collected in September 1969 from the surface abundant in Lake Rotoiti. of Lake Rotoiti between Wrights (Te Pohoe) Bay and Cherry Bay was examined. Here the Aufwuchs surface water displayed parallel rows of golden brown organisms which, on micro The aufwuchs community, i.e., organisms scopic examination, consisted of a number of attached to ormoving upon aliving substrate, normally planktonic species intermingled with occursinlakes Rotoruaand Rotoitimostcom masses of winged pine pollen embedded in monly on the macrophytes Lagarosiphon mqjor clumps of amorphous golden brown organic and Nitella hookeri. A few samples have been matter, which could have been a disintegrated examinedfromthelittoralzone(shallowinshore Algal Flora ofSome North Island Lakes-CAssIE 473 FIG. 4. Microrystisaeruginosa Klitz. and Staurastruntfloriferttnt (West & West) Brook. Film 14/19A. x3400. FIG. 5. AnabaenacircinalisRabenh. Film 9/31. x3600. 474 PACIFIC SCIENCE, Volume 28, October 1974 areas up to 5m deep). Twenty-two species of algae have been identified from the aufwuchs (Table 2). Diatoms predominate (11 species) and ChloropfDiceae are abundant (7 species). There was no opportunity to study seasonal behavior of the principal species. Many more will doubtless be revealed by some intensive investigations. Among the diatoms, pennate forms are the most conspicuous. Epithemia sorex, rather like a set offalse teeth when two adjacent fructules are seen in girdle view (Fig. 15), is one of the commonest (cf. Flint 1938, who noted a winter dominance and a year-roundoccurrenceofE.sorexinLakeSarah, Cass District, S. Island, New Zealand; and Castenholz 1960, who recorded this species in Alkali Lake, Lower Grand Coulee, Washing ton, as a prominent member of a pennate diatom community during autumn). Epithemia zebra (Fig. 16) is less frequently encountered. Epithemia and Eunotia sp. attach themselves firmly to leafand stem apices of Lagarosiphon and Nitella. Ofthe unattachedspecies, Melosira lineata(3 to 10cellsperchain, Figs. 17, 18) and Fragilariacrotonensis(Fig. 19)arelessfrequently encountered. Fragilaria virescens (Fig. 20) may FIG. 6.Pandorinamorttm(Mull.)Bory.Film9/28. x4000. FIG. 7. Trachclomonasvolvocina(Ehr.)andAsterioncllaformosaHass. Film9/17A. x4000. Algal Flora ofSome North Island Lakes-CAssIE 475 FIG. 8. Dinob~yon cylindriCtimva!. alpinum(1mb.) Bacbm. Film12/16. x1125. also occur. Many different small green algae, including Closteriumparvulum and C. aciculare (Figs. 21, 22), Scenedesmus, Kirchneriella, Cruci genia(Fig. 23), Selenastrum, and Mougeotia(wide cells), are intermingled with normally plank tonic species such as Melosira granulata, M. distans, Mougeotia(narrow cells), and numerous species of Staurastrum. Filaments of LyngbJla frequently enmesh fronds of the macrophyte Lagarosiphonnearthesurface. Othersworthyof mention among the aufwuchs community are Rhopalodia gibba, Ankistrodesmus falcatus, and the desmid Sphaerozosma aubertianum vaL com pressum, and Tabellariaflocculosa(Figs. 24-26). Equallycommonamongaufwuchsorphyto planktonareEudorinaelegans,Pediastrumduplex, and P. boryanum (Figs. 27-29), as well as Ankistrodesmusand Sphaerozosma. The position of Peroniella planktonica (Fig. 30) is hard to categorize, since It is planktonic merely by growing as an epiphyte and being carried up into thephytoplanktonbyits filamentous host. A specimen ofNitella hookeri collectedfrom Cherry Bay by B. Coffey showed a growth at branch apices ofanalgacloselyresembling the FIG. 9. Nepbroeytitlm agardbianllm Nag. Film 6j20A. x4000. genus Thamniochaete (cf. Bourrelly 1966). The 476 PACIFIC SCIENCE, Volume 28, October 1974 FIG. 10.Staurastrumleptoc/adiumvar. insigneWest & West. Film15/10. x1800. FIG. 11.Sta!lrastrumsexangulare(Bulnh.) Lund. Film12/11A. x1275. l2\LIO;U;.~u:::z:zEmlU,l1l!.nu::::'SiS
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