G. W. Yeates N.Z. Soil Bureau, Lower Hutt

N.Z. Soil Bureau Scientific Report 77 Department of Scientific and Industrial Research Lower Hutt, 1986 Bibliographic Reference: YEATES, G. W. 1986: Enchytraeidae - some population estimates for grasslands and a New Zealand bibliography N.Z. Soil Bureau Scientific Report 77 14 p.

ISSN 0304-1735

Editing: David Isaacs Typing: Tessa Roach Draughting: Carolyn Powell

V. R. WARD, GOVERNMENT PRINTER, WELLINGTON, NEW ZEALAND-1986 CONTENTS

ABSTRACT 5 INTRODUCTION 5 METHODS 5 RESULTS AND DISCUSSION 6 1. Tussock grasslands 6 2. Pasture sui"veys 6 3. Vertical distribution in horotiu soil; grass species and insecticide effects 7 4. Wairarapa fertiliser : stocking rate trial 8 5. insecticide X ryegrass cultivar trial 8 6. Effects of repeated use of oxamyl 8 7. Phenology in three pastures 8 Conclusions 12 Acknowledgments 12 REFERENCES 12 ANNOTATED BIBLIOGRAPHY 13

Figures Temporal variation in total enchytraeid populations in Horotiu sandy loam, under 4 ryegrass cul­ tivars 7 2 Phenology of enchytraeid populations in Horotiu sandy loam, under mature pasture 9 3 Phenology of enchytraeid populations in Otorohanga silt loam, under mature pasture 10 4 Phenology of enchytraeid populations in Te Kauwhata clay loam, under mature pasture II

Tables Vertical distribution of enchytraeid populations in Horotiu soil under cocksfoot and ryegrass with various treatments 6 2 Estimated enchytraeid populations in Kokotau silt loam under various management regimes 8 3 Enchytraeid populations in various treatments at the Wairakei trial .. 8 4 Effects of insecticidal rates of oxamyl on enchytraeid populations below 4 ryegrass cultivars 8 5 Correlations between 3-weekly estimates of enchytraeid populations and soil moisture and soil tem­ perature II s

ABSTRACT

The occurrence of enchytraeids in survey samples from tussock grasslands and pastures is recorded. In pasture trials, populations are affected by fertiliser rate, sward productivity and stocking rate. Effects of insecticides and nematicides on populations under pasture are variable. Sampling each three weeks gave annual mean populations of 19 500-52 100/m2, and their seasonal variation correlates significantly with soil moisture and temperature. Thirty-seven references to work on enchytraeids in the New Zealand region are annotated. Keywords: Enchytraeidae; pastures; grasslands; phenology; pesticides; population numbers; bibliography

INTRODUCTION

Although enchytraeids (Annelida : Oligochaeta : European workers have shown the role of enchy­ Enchytraeidae) were first recorded from New Zealand traeids in soil decomposition processes (e.g., Abra­ soils last century (Beddard 1893), for many years only hamsen 1983, Anderson et al. 1983, Phillipson et al. passing reference was made to them (e.g., Stout 1954). 1979). Kasprzak (1982) suggested that in cultivated Luxton (1976) provided no New Zealand references soils enchytraeids may play a greater role in minerali­ to enchytraeids in his bibliography of soil animals in sation of organic matter than do lumbricid earth­ New Zealand. However, during the last 20 years, worms. Standen (1984) compared enchytraeid and population estimates of enchytraeids have been made earthworm populations in 14 long-established hay in a range of soils. These estimates have generally meadows. been incidental to more general soil zoological or pes­ This paper presents a series of population esti­ ticide studies. mates for enchytraeid populations extracted during European work on Enchytraeidae was stimulated soil nematode studies and provides an annotated by the basic taxonomic revision of Nielsen and Chris­ bibliography for Enchytraeidae in the New Zealand tensen (1959), and general accounts of the biology of region. As no attempt has been made to distinguish, enchytraeids, their sampling, abundance and distri­ enchytraeid species, detailed analysis of population bution are given by O'Connor (1967) and Dash (1983). responses to treatments is not made.

METHODS

Unless otherwise specified, populations reported Sample means and standard errors are given. here were recovered by the use of a Baermann tray, The bibliography was checked against the 'Biblio­ as described by Yeates (1978). No special care was graphy of Non-Marine Invertebrate.s of the New taken with the handling of the samples and thus Zealand Region', maintained at Entomology Divi­ physical damage may have reduced enchytrae!d sion, D.S.l.R., Auckland, for publications containing recoveries. In each case reference is made to the significant data; anecdotal or general references have nematological paper in which further location, treat­ been omitted. ment, sampling and agronomic data are contained. 6

RESULTS AND DISCUSSION

1. TUSSOCK GRASSLANDS (1985). Enchytraeid population estimates (thou­ All the samples listed by Yeates (1974) were /m2 in 0-10 cm soil, n~5) were: extracted by Baermann funnel and examined for enchytraeids. They were detected only in the bottom Site Soil · Enchytraezd 5 cm of leaf base of Chionochloa rubra from the Hari­ population 2 hari sites. Dr J.D. Stout (unpublished 1973) found (thousands/m ) enchytraeids in standing Chionochloa litter from the I. Horotiu sandy loam 56.5± 8.8 McKerrow site, and from standing litter of Chionoch­ 2. Otorohanga silt loam 27.0± 6.3 loa rigida in South Canterbuty (Williams 1977). 3. Te Kauwhata clay loam 12.8± 2.9 2. PASTURE SURVEYS 4. Oruanui hill soil 55.4± 13.1 Enchytraeids were found in 8 of the 77 pasture 5. Ohinepanea sandy loam 39.2± 14.8 samples extracted by Baermann funnel and examined 6. Horotiu sandy loam 17.2± 3.6 by Yeates (1975). The sites, which were all sampled 7. silt loam 23.8± 7.6 in October or November, were: 8. Hamilton clay loam 20.0 (n~2) Site no. Soil 9 Ohaupo silt loam 24.3± 2.2 11 Claremont silt loam 10. Puniu silt loam 22.9± 4.9 18 Pawson silt loam 11. Dunmore silt loam 31.0±11.3 23 Wairepa silt loam 12. Te Kauwhata clay loam 20.1± 1.6 36 Taupe sandy silt 13. Taupe ash soil 29.6± 18.4 37 Kaingaroa silt 14. Oruanui soil 11.7 ± 2.6 44 Egmont brown loam I 5. Ohinepanea sandy loam 61.1±10.6 46 Egmont black loam 16. Ohinepanea sandy loam 71.4± 7.0 51 Levin silt loam It is apparent from these samples that enchytraeids were present at moderate to high population levels at Sixteen mature pasture sites in the Taupe-Wai­ all sites. Generally, high populations would be kato-Bay of Plenty were quantitatively sampled in expected at this time of year. Details of seasonal vari­ August 1982 for the work reported by Watson et al. ation at three of the sites is given in section 7.

Table I Vertical distributiori of enchytraeid populations (thousands/ml) in Horotiu soil under 'Grasslands Apanui' cocksfoot (Dactylis glomerata L.) and 'Grasslands Nui' ryfgrass (Lolium perenne L.) with various treatments.

May 1982 May 1983 Depth mown mown grazed untreated oxamyl-treated interval cocksfoot rye grass ryegrass ryegrass ryegrass (cm) (n=2) (n=2) (n=3) (n=S) (n=2)

0-10 2.5 56.8 15.8 10 11 10-20 5.1 0.8 2.4 2.4 3.6 20-30 0.3 1.3 0 1.2 0 30-40 0.3 0 0 1.4 1.2 40-50 0.5 0 0.2 0.4 0.6 50-60 0.3 0.3 0 1.0 0.2 60-70 0 0.5 0 0.1 0 7

H all cultivars ±LSD X Grasslands N ui 40,000

x ~ E ~c 0 :"§ 20,000 ::l "- 0 ------0..

x

0'---'~~~-'-~~--'--~~~~~~'--~~-'-~~-'-~~~~~~~~~~~~~~- 30 30 5 6 26 19 31 10 4 2 Sep Nov Feb Apr May Jui Aug Nov Jan Mar May 1981 1982 1983

Figure I Temporal variation in !Ota! enchytraeid populations in Horotiu sandy loam, under 4 ryegrass cultivars (untreated and oxamyl treated).

3. VERTICAL DISTRIBUTION IN HOROTIU SOIL; in 1982 and 1983; the greatest contrast in the top GRASS SPECIES AND INSECTICIDE EFFECTS 10 cm was between mown cocksfoot and mown rye­ Cores to 1 m were taken on two occasions, once grass. The total nematode populations in these two to compare enchytraeids under various grasses (Yeates treatments also differed markedly, but in the opposite et al. 1983) and once to assess the effects of insecti­ direction (cocksfoot, 4.2 million/m2; ryegrass 1.6, mil­ cidal rates (0.5 kg a.i./ha) of oxamyl (section 6). lion/m2) (Yeates et al. 1983). The bacterial-feeding Enchytraeids were recorded up to 70 cm below the Rhabditida averaged 808 000/m2 in 0-10 cm soil below surface (Table 1). Oxamyl treatment had no marked cocksfoot but only 268 000/m2 below ryegrass. It is effect on population levels, and populations in the top possible that Rhabditida and Enchytraeidae use a 10 cm of soil below grazed 'Nui' ryegrass were similar common food resource. 8

4. WAIRARAPA FERTILISER: STOCKING RATE 6. EFFECTS OF REPEATED USE OF OXAMYL TRIAL Forty-eight large plots of Horotiu sandy to gravelly loam were resown with ryegrass in autumn 1.981; each Samples taken in July 1971 (wet season) from the plot received one of the 4 ryegrass cultivars 'NZ. per­ trial on Kokotau silt loam, described by Yeates (1976), ennial ryegrass', 'Grasslands Nui', 'Grasslands Man­ were extracted by Baermann funnel. Their enchy­ awa', 'Ellett'. The trial was grazed and subplots treated traeid populations are listed in Table 2. The average with 0.5 kg a.i./ha oxamyl granules each 10-15 days population over 16 paddocks was 18 800/m', and the during the growing season. About each 2 months all increases due to api)lication of lime or superphos­ plots were sampled, duplicate sets of cores being taken from each subplot (96 sets in total); during winter, phate, while not significant, indicate greater primary production due to these applications. The depression only the 'Nui' plots were sampled (Yeates and Barker 1986). of enchytraeid population at the heavier grazing pres­ sure (P c. 0.05) presumably reflects less litter due to Each summer, enchytraeid populations declined to greater herbage removal by grazing animals at this almost zero in all plots (Fig. 1). Analysis of variance showed populations under each cultivar were not sig­ level; treading effects are considered unlikely. nificantly different. On seven sampling occasions populations in the oxamyl-treated subplots were sig­ nificantly higher or lower than in the untreated sub­ plots (Table 4); the same comparisons for total nematodes all show significant reductions in the oxa­ Table 2 Estimated enchytraeid populations (/m 2, 0- myl-treated subplots. The mean population in 10 cm soil depth) in Kokotau silt loam under 2 various management regimes (n = 8 in each untreated plots was 16 OOO/m (0-10 cm soil depth). case).

Treatment Population/mi Table 4 Effects of insecticidal rates of oxamyl on enchytraeid populations below 4 ryegrass Sheep/ha 14.8 31 ODD± 9 250 cultivars; probability of the treatment effect is 22.2 6 600± 2 250 given for each sampling Lime (t/ha) 0 10200± 4 020 5.6 27 400± 9 810 Untreated Ox amyl p Superphosphate (kg/ha) 125 14200± 3640 500 23 400± 10 690 All cultivars February 1982 I 600 900 0.004 August 1982 16 700 21 OOO 0.107 January 1983 10 100 ID 300 0.081 5. WAIRAKEI INSECTICIDE X RYEGRASS March 1983 850 2 OOO 0.037 CUL TIVAR TRIAL 'Nui' ryegrass A grazing trial sown on Oruanui sandy loam in February 1982 2 100 I 680 0.041 1982 was sampled in April-May 1984 (Yeates and May 1982 28 700 19 500 0.010 Prestidge 1986). A total of 64 samples was examined November 1982 17 200 24 100 0.089 from 'Grasslands Ruanui' and 'Ellett' ryegrass plots, some of which had received spring applications of 7. PHENOLOGY IN THREE WAIKATO PASTURES oxamyl at insecticidal rates. These were significantly more enchytraeids below At three sites mowing trials were carried out in 'Ruanui' ryegrass and significantly fewer enchytraeids old pasture to assess response to control of white clo­ in the oxamyl-treated plots than in the untreated ones ver pests (Watson et al. 1985). Each three weeks sam­ (Table 3). ples were taken from four replicates of the following treatments: (a) untreated control (b) nematicide treated (oxamyl or fenamiphos) Table 3 Enchytraeid populations (/m2 in 0-10 cm soil) (c) nematicide treated plus urea application. in various treatments at the Wairakei trial; Over the 12-month sampling period mean popu- probability of the cultivar and treatment lations in untreated plots were: effects is given. Horotiu sandy loam 52 100/m2 Otorohanga silt loam 38 500/m2 Variable Population/ml p Te Kauwhata clay loam 19 500/m'. For the 18 sampling occasions analysis of variance Ellett 32 000 gave: 0.019 Ruanui 51 OOO 16 significant (P < 0.05) site effects 9 significant (P < 0.05) site X nematicide X Untreated 65 OOO nematicide brand interactions 0.001 Oxamyl 18 OOO 8 significant (P < 0.05) nematicide X nematicide brand interactions. 9

120,000 b a .. . a • I b :/ .. ·.b :1 b :1 b; :/ :1 ·. :, 90,000 a a. a I ab I a ,, I a . ,1 I ab a I I I

~ E I a ; a/ c a/ . I -0 I .... 60,000 ab . I ~ ., ::J a Q. a· I 0 \ ab;. a\ o_ a': I I I I I I I I I a a a:/ a 1:/ I . I :a 30,000

14 14 Sep Mar Sep 1982 1983

Figure 2 Phenology of enchytraeid populations in Horotiu sandy loam, under mature pasture. Untreated (n = 4), oxamyl treated (n = 2) and fenami"phos treated (n = 2) plots were sampled; treatments which do not share the same leuer on a sampling occasion differ si"gnificantly IP < 0.05). 10

120,000

c a

90,000

:: a

a

~ E c -0 a ~ 60,000 a "'Q_ 0 I\ (L I I \ a a I a \ a I \ a a I ab I \ :··· 1 \ a .. /\ / a ~ I·. a a ; ,1 ·... I I I al 30,000 \ I \ I : a \ I I I I al I ?. . ·a· ••• ''a .. a I I a I I a... a: I .: ...

b. : .... b .·a·. a a a.· ·.a b 0 7 22 7 Sep Feb Sep 1982 1983

Figure 3 Phenology of enchytraeid pOpulations in Ocorohanga silt loam, under mature pasture. (See Fig. 2 for key). II

60,000

untreated a/ -- oxamyl I a\ ...... fenam iphos l •... ·.. \ a.. . a . . ~. L- .a a a: ·I ·. ·.a··\ ~ E . • · ·•• / a a\I' a a c \ .. -0 a ] 30,000 a \ ·.a "Q_ \ 0 a I a_ c .. la \ b ... .-a I la abr>. a I . a; / a I I a I b

OL---'--.._----'---'----'-....L..-L--1...-~----L---'---'--'--L--'---'-----'-~ 21 8 21 Sep Mar &p 1982 1983

Figure 4 Phenology of enchytraeid populations in Te Kauwhata clay loam, under mature pasture. (See Fi"g. 2 for key).

These results are summarised graphically (Figs 2- 4). Soil moisture was measured and temperature was Table 5 Correlations between 3-weekly estimates of assessed for each sampling occasion. The enchytra:eid enchytraeid populations and soil moisture and populations at each site were significantly (P < 0.05- soil temperature p < 0.001) correlated with soil moisture and temper­ ature for the sampling occasion or for three weeks Variable Horotiu Otorohanga Te Kauwhata earlier (Table 5). sandy-loam silt loam clay loam Thus, over 12 months: (a) populations at the 3 sites differed significantly Soil moisture 0.843*** 0.497* 0.837*** (b) populations varied markedly with time, being Soil moisture 3 weeks 0.727*** 0.707** 0.665** lowest at the driest time of year earlier (c) soil moisture and temperature correlate with Soil temperature 0.696** 0.547* 0.723** population changes Soil temperature 3 0.760*** 0.565* 0.635** (d) nematicide effects vary with location, time and weeks earlier nematicide. 12

CONCLUSIONS

Enchytraeids are widespread in New Zealand and their contribution to organic matter dynamics in grasslands and the population densities are high by New Zealand pastures appears warranted. European standards. Effects of the pesticides applied appear to vary with time and soil. Stocking rate, sward Acknowledgments composition and productivity all influence enchy­ Messrs G. M. Barker, G. C. Crouchley, R. N. Wat­ traeid populations. Populations appear to be lowest at son and Dr R. A. Prestidge, all of the Ministry of the driest time of the year. In view of the diversity Agriculture and Fisheries, have provided access to of enchytraeid faunas recorded overseas and their trials under their control. Dr T. K. Crosby searched apparent complementary interaction with other ele­ the computer data-base, 'Bibliography of Non-Marine ments of the soil fauna, specific investigation of the Invertebrates of the New Zealand Region', at Ento­ abundance, diversity and phenology of enchytraeids, mology Division, D.S.l.R.

REFERENCES

Note: Publications containing substantive information on New WATSON, R. N.; YEATES, G. W.; LITTLER, R. A.; STEELE, Zealand enchytraeids are contained solely in the annotated bibliography K. W. 1985: Responses in nitrogen fixation and herbage pro­ duction following pesticide applications on temperate pas­ tures. Proc. 4th Australasian Conference on Grassland Invertebrate Ecology (R. B. Chapman, Ed): 103-113. ABRAHAMSEN, G. 1983: Effects of lime and artificial acid rain on the enchytraeid (Oligochaeta) fauna in coniferous forest. WILLIAMS, P.A. 1977: Growth, biomass, and net productivity of Holarctic ecology 6: 247-254. tall-tussock (Chionochloa) grasslands, Canterbury, New Zealand. New Zealand Journal of Botany 15: 399-442. ANDERSON,]. M.; INESON, P.; HUISH, S. A. 1983: The effects of animal feeding activities on element release from deci~ YEATES, G. W. 1974: Studies on a climosequence of soils in tus­ duous forest litter and soil organic matter. Pp. 87-100 in sock grasslands. 2. Nematodes. New Zealand Journal of Zool­ Lebrun, P., Andre, H.M., de Medts, A.; Gregoire-Wibo, C. ogy J; 171-177. and G. Wauthy (Eds) 'New Trends in Soil Biology'. Pro­ ceedings of the eighth International Colloquium of Soil YEATES, G. W. 1975: Nematode genera from some New Zealand Zoology, Louvain. pastures. New Zealand Soil Bureau Scientific Report 21; 22 p. DASH, M. C. 1983: 'Biology of Enchytraeidae'. International Book Distribution, Dehra-Dun. YEATES, G. W. 1976: Effect of fertiliser treatment and stocking rate on pasture nematode populations on a yellow-grey earth. KASPRZAK, K. I 982: Review of enchytraeid (Oligochaeta, New Zealand Journal of Agricultur~l Re_search 19: 405-408. Enchytraeidae) community structure _and function in agri­ cultural ecosystems. Pedobiologia 23: 217-232. YEATES, G. W. 1978: Populations of nematode genera in soils under pasture. I. Seasonal dynamics in .dryland and irrigated LUXTON, M. 1976. Bibliography of soil animals in New Zealand, pasture on a southern yellow-grey earth. New Zealand Jour­ to 1967. New Zealand Soil Bureau Bibliographic Repon 18. nal of Agricultural Research-21: 321-330. 76 p. YEATES, G. W.; BARKER, G. M. 1986: Influence of repeated O'CONNOR, F. B. 1967. The Enchytraeidae. Pp. 213-257 in A. oxamyl application on nematodes beneath four ryegrass cul­ Burges and F. Raw (Eds) 'Soil Biology'. Academic Press, tivars on a yellow-brown loam. New Zealand Journal ofAgri­ London. cultural Research 29 (in press). PHILLIPSON, ).; ABEL, R.; STEEL, ).; WOODELL, S. R. j. YEATES, G. W.; PRESTIDGE, R. A. 1986: Nematode popula­ 1979: Enchytraeid numbers, biomass, and respiratory tions and their effects on herbage production in a·-volcanic metabolism in a beech woodland - Wytham Woods, Oxford. plateau pasture. New Zealand Journal ofAgricultural Research Oecologia 43: 173-193. 29 (in press). STANDEN, V. 1984: Production and diversity of enchytraeid, YEATES, G. W.; STANNARD, R. E.; BARKER, G. M. 1983; earthworms and plants in fertilised hay meadow plots. Jour­ Vertical distribution of nematode populations in Horotiu nal of Applied Ecology 21: 293-312. soils. New Zealand Soil Bureau Scientific Report 60. 14 p. 13

ANNOTATED BIBLIOGRAPHY

BEDDARD, F. E. 1893. Some new or little-known Oligochaeta. LUXTON, M. 1968: The fauna of New Zealand soils. In: Proceedings of the Royal Physical Society of Edinburgh 12: 31- Anzaas 40th congress, Christchurch, Abstracts section D. 45. Preliminary report of 5 following papers. Microscolex nova zelandiae n.sp., Fridericia antarctica n.sp, Hen/ea ventriculosa. LUXTON, M. 1982: Studies on the invertebrate fauna of New Zealand peat soils. I. General survey of the sites. Revue BENHAM, W. B. 1903: On some new species of aquatic Oligo­ d'Ecologie et de Biologie du Sol 19: 535-552. chaeta from New Zealand. Proceedings of the Zoological Society Methods;. comparison of sites and populations. of London 1903, ii: 202-232. Enchytraeus simulans n.sp, Achaeta maorica n.sp. LUXTON, M. 1982: Studies on the invertebrate fauna of New Zealand peat soils. II. Restiad . Revue d'Ecologie et de BENHAM, W. B. 1904: Phylum Annulata. Pp.276-285 in F. W. Biologie du Sol 19: 553-578. Hutton 'Index Faunae Novae Zealandiae'. i\1ean populations of 2462, 548, 5570/m2 • Fridericia galba, Hen/ea ventriculosa listed on p. 284; Appen­ dix lists Enchy!raeus simulans, Achaeta maorica on p. 346. LUXTON, M. 1982: Studies on the invertebrate fauna of New Zealand peat soils. IV. Pasture soils on Rukuhia peat. BENHAM, W. B. 1904: A note on the Oligochaeta of the New Pedobiologia 24: 297-308. Zealand lakes. Transactions and Proceedings of the New !vlean population of 1674/m2• Zealand Institute 36: 192-198. Enchytraeus simulans n.sp. (Taupo), Achaera maorica n.sp. LUXTON, M. 1983: Studies on the invertebrate fauna of New (Manapouri). Zealand peat soils. III. Fern peats. Revue d'Ecologie et de Biologie du Sol 20: 87-109. BENHAM, W. B. 1905: On the Oligochaeta from the southern islands of the New Zealand region. Transactions and Pro­ Mean populations of 17 946, 47 527/m2 • ceedings of the New Zealand Institute 37: 285-297. LUXTON, M. 1983: Studies on the invertebrate fauna of New Marionina antipodum n.sp., Enchytraeus albidus, Lumbricil­ Zealand peat soils. V. Pasture soils on Kaipaki peat. Pedo­ lus tnacquariensis n.sp. biologia 25: 135-148. BENHAM, W. B. 1905: Earthworms from the Kermadecs. Trans­ Mean population of 1297/m2• actions and Proceedings of the New Zealand Institute 37: 298- MARTIN, N. A. 1972: The effect of insecticides on the soil and 299. pasture fauna. Abstracts of the 14th International Congress of Six unidentified enchytraeids found in sample. Entomology, Canberra: 261. BENHAM, W. B. 1909: Report on Oligochaeta of the Subantarctic Preliminary report of the following paper. Islands of New Zealand. Pp. 251-294 in C. Chilton (Ed) MARTIN, N. A. 1975. Effect of four insecticides on the pasture The Subantarctic Islands of New Zealand. ecosystem. IV. Enchytraeidae and Diptera larvae heat­ Lutnbricus intermedius n.sp. extracted in water-filled funnels. New Zealand Journal of BENHAM, W. B. 1915: Oligochaeta from the Kermadec Islands. Agricultural Research 18: 313-315. Transactions and Proceedings of the New Zealand Institule 47: Populations not significantly reduced by insecticides. 174-185. Fredericia bollonsi n.sp., F.B. variety oliveri. MARTIN, N. A. 1983: Miscellaneous observations on a pasture fauna: an annotated species list. D.S.!.R. Entomology Divi­ BENHAM, W. B. 1915: On Lumbricillus macquariensis Benham. sion Report No. 3. 98 p. Transactions and Proceedings of the New Zealand Institute 47: Maximum population 47 OOO/m 2 • 189-191. L. 1nacquariensis junior synonym of L. intermedius. McCOLL, H.P. 1984: Nematicides and field population of enchy~ traeids and earthworms. Soil biology and biochemistry 16: 139- BENHAM, W. B. 1950: Oligochaeta from the Auckland and 143. Campbell Islands. N.Z. Departmenl of Scientific and Indus­ Populations correlated with soil moisture, not affected by trial Research: Cape Expedition Series Bulletin 10: 27-35. nematicides; maximum population c.47 000/m2• Three unidentifable specimens from 'leaf mould', Ewing Is., Auckland Islands. MILLER, R. B.; STOUT,.J. D.; LEE, K. E. 1955. Biological and CLARK, W. C. 1964: A new species of Sectone1na Thorne, 1930 chemical changes following scrub burning on a New Zealand (Family Nygolaimidae, Nematoda). New Zealand Journal of hill soil. N.Z. Journal of Science and Technology B37: 290- Science 7: 174-176. 313. Enchytraeidae as prey. Enchytraeids present 2 months after burning.

DUTCH, M. E.; STOUT,]. D. 1968: The carbon cycle in a beech NIELSON, C. O.; CHRISTENSEN, B. 1959: The Enchytraei-d.ae: forest ecosystem in relation to microbial and animal popu­ critical revision and taxonomy of European species. Natura lations. Transactions of the 9th International Congress of Soil Jutlandica 8-9: 1-160. Science 2: 37-48. Achaeta maorica, Hen/ea ventriculosa, Fridericia bollonsi, F. Enchytraeidae present but no separate counts given. b. v. oliveri, Marionina anri"podum listed from New Zealand. LEE, K. E. 1968: A preliminary study of soil animals and their STOUT, j. D. 1954: The ecology, life history and parasitism of relationships to some New Zealand soils. New Zealand Soil Tetrahymena (Paraglaucoma) rourata (Kahl) Corliss. Jour­ Bureau Bullelin 26(2): 168-183. nal of Protozoolagy 1: 211-215: Recorded from 1 of 45 sites surveyed. Infects Enchytraeidae. 14

STOUT,]. D. 1958: Biological studies of some tussock-grassland STOUT, J. D. 1968. Micro-norganisms and peat. Appendix 3, pp. soils. VII. Protozoa. New Zealand JOurnal of Agricultural 127-33 in 'Peat Classification by Pedological Methods, Research I: 974-984. Applied to Peats of Western Wellington, New Zealand', by Enchytraeidae found in 5 of 9 samples. W. F. Harris. N.Z. Department of Scientific and Industrial Research Bulletin 189. 138 p. Enchytraeids ill 2 Macquarie I. peat samples. STOUT, J. D. 1958: Aquatic oligochaetes oq::urring in forest litter. II. Transactions of the Royal Society of N.Z. 85: 289-99. YEATES, G. W. 1967. Studies on nematodes from dune sands. 9. Enchytraeids occurred in cultures from beech forest litter. Quantitative comparison of the nematode faunas of six localities. New Zealand Journal of Science JO: 927-948. STOUT,]. D. 1960. Biological studies of sOme tussock-grassland Present in all samples; 7-404/1000 g sand. soils. XVIII. Protozoa of two cultivated soils. New Zealand YEATES, G. W. 1968: Analysis of annual variation of the nema­ Journal of Agricultural Research 3: 237-244. tode fauna in dune sand, at Himatangi Beach, New Zealand. Enchytraeidae found in 5 of 9 protozoa cultures. Pedobiologia 8: 173-207. Present in all samples; 6-355/1000 g sand; correlation with STOUT,]. D. 1961: Biological and chemical changes following predatory nematode. scrub burning on a New Zealand hill soil. 3. Microbiological YEATES, G. W. 1969: Predation by Mononchoides potohikus changes. N.Z. Journal of Science 4: 740-52 .. {Nematoda:Diplogasteridae) in laboratory cU:lture. Nemato­ Enchytraeids in litter only; less common when pasture logica 15: 1-9. established. Enchytraeidae as prey. YEATES, G. W.; WATSON, R. N.; BARKER, G. M.; PRES­ STOUT, J. D. 1961. The protozoan fauna. P. 77 in 'Little Barrier TIDGE, R. A. 1985: Enchytraeid populations of some Wai­ Island, Hauturu'. (Comp. W. M. Hamilton). N.Z. Depart­ kato-Taupo pastures. Proceedings of the Fourth Australasian ment of Scientific and Industrial Research Bulletin 137. Conference on Grassland Invertebrate Ecology: 375. 198 p. Abstract of poster paper covering some aspects of work Enchytraeids in a loamy peat sample. reported in the present paper.