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    This article was downloaded by: [118.92.18.216]On: 30 January 2015, At: 18:29Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number:1072954 Registered office: Mortimer House, 37-41 Mortimer Street,London W1T 3JH, UK

    New Zealand Journal of

    Marine and Freshwater

    ResearchPublication details, including instructions for

    authors and subscription information:

    http://www.tandfonline.com/loi/tnzm20

    Summer upstream

    migration of juvenile

    freshwater eels in New

    ZealandD. J. Jellyman abaMarine Laboratory, Department of Zoology ,

    Victoria University of Wellington , Private

    Bag, Wellington, New ZealandbFisheries Research Division , Ministry of

    Agriculture and Fisheries , P.O. Box 19062,

    Wellington, New Zealand

    Published online: 30 Mar 2010.

    To cite this article:D. J. Jellyman (1977) Summer upstream migration of

    juvenile freshwater eels in New Zealand, New Zealand Journal of Marine and

    Freshwater Research, 11:1, 61-71, DOI: 10.1080/00288330.1977.9515661

    To link to this article: http://dx.doi.org/10.1080/00288330.1977.9515661

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    N.Z. Journalof Marine andFreshwater Research11 1) :6 1 - 7 1 . March1977

    SUMMER UPSTREAM MIGRATION OF JUVENILE

    FRESHWATER EELS

    IN NEW

    ZEA LA N D

    D. J.

    JELLYMAN*

    Marine Laboratory, Department of Zoology, Victoria University of

    Wellington, Private Bag, Wellington, New Zealand

    ABSTRACT

    During summer, upstream migrations

    of

    juvenile freshwater eels (elvers)

    of

    both species (Anguflla australis schm idtii Phillipps,

    the

    short-finned

    eel, and

    A. dieffenbachii

    Gray,

    the

    long-finned

    eel) are

    observed

    at

    hydro-electric dams

    and waterfalls. The migration was sampled at Karapiro Dam on the Waikato

    River,

    1970, 1971, and 1974, and

    samples from

    a

    further

    11

    areas throughout

    the country were analysed.

    Elvers, active

    at

    night

    or

    during overcast

    and

    damp days, surmount steep

    obstacles

    by

    climbing; they adhere

    to

    damp surfaces

    by

    friction

    and

    surface

    tension, obtaining maximum resistance

    by

    undulating their bodies

    and

    keeping

    them closely adpressed

    to the

    substrate.

    The

    maximum size

    for

    vertical climbing

    is

    12

    cm .

    As

    elvers were sampled

    at

    sites where their migration

    was

    interrupted,

    no distinct schools were observed,

    but the

    elvers were strongly social

    and

    thigmotactic.

    Migrations are made up of elvers of similar size of one or both species.In

    large river systems, elvers apparently migrate successively further upstream each

    year for several, years. North Island samples contained elvers up to 3 y of age,

    but

    in the

    South Island elvers

    up to 7 y old

    were recorded; aging

    was by

    burnt

    otoliths. Migration

    is

    perhaps initiated

    by

    increases

    in

    water temperature,

    day

    length, and sociability; increased water flow may also coincide w ith 'runs '.

    Electric fishing of streams shows that the two species hav e different ha bita t

    preferences,

    and

    these

    are

    reflected

    in the

    proportion

    of

    elvers caught

    at

    different

    sites: longfins prefer swift stony rivers, shortfins silty backwaters and lakes,but

    both

    are

    often found together. Longfin elvers have thick dermal skin layers,

    and experiments in air of constant temperature and hum idity showed that they

    cannot survive prolonged exposure requiring cutaneous respiration

    as

    well

    as

    shortfins; longfins

    may

    also have h igher oxygen requirem ents.

    INTRODUCTION

    With the onset of summer each year, mass upstream migrations of

    juvenile freshwater eels of both Anguilla australis schmidtii Phillips,

    the short-finned eel, and A. dieffenbachii Gray, the long-finned eel, take

    place in rivers throughout the country. Such eels are referred to as

    elvers , as distinct from glass-eels , which is the stage at which eels

    arrive in fresh water.

    Present address:

    Fisheries Research Division, Ministry

    of

    Agriculture

    and

    Fisheries,

    P.O. Box

    19-062, Wellington,

    New

    Zealand.

    Received8July1975;revision received21July1976.

    Fisheries Research Division Publication

    262

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    62 N.Z.

    J O U R N A L

    OF

    M A R I N E

    &

    F R E S H W A T E R R E S E A R C H [ M A R C H

    TABLE 1Species

    proportions

    (%) and

    mean lengths L,

    cm) of

    migratory

    elvers

    (Anguilla

    spp.) sampled 1970, 1971, 1974.

    ARE A

    Te Kauwhata

    Karapi ro

    DAT E

    Jan.

    1971

    J a n - Fe b

    1970,

    1971, 1974

    Matahina

    Mokau River

    Shannon

    Makara si te

    a

    Makara si te

    b

    D obs on

    Lake Coleridge

    Aviemore

    Waitaki

    Roxburgh

    Jan

    1974

    Jan

    1971

    Feb.

    1971

    Feb.

    1972

    Jan. -Feb.

    1972

    Jan.

    1971

    Jan.

    1971

    Feb.

    1971

    Feb.

    1971

    Jan-Feb.

    1971

    n

    7 4 3

    3646

    1 6 8

    2 3 2

    2 3 3

    169

    4 4

    2 1 6

    8 0

    2 6

    6 5

    2 0 7

    %

    100

    6 2

    7 9

    18

    18

    9 6

    6 1

    12

    0

    0

    9

    1

    A. austral.

    L

    8. 1

    9.0

    9.0

    9.4

    10.2

    7.5

    7.9

    10.1

    13'.

    5

    12 . 0

    (1

    S.D.)

    (0.87)

    (0.98)

    (0.88)

    (0.81)

    (0 .66)

    (0.75)

    (1.22)

    (0.88)

    ( L 8 6 )

    (0.30)

    is

    Range

    5 . 9 - 1 1 . 4

    6 . 6 - 1 2 . 9

    7 . 2 - 1 1 . 5

    7 . 6 - 1 1 . 1

    8 . 3 - 1 1 . 6

    6 . 2 - 1 1 . 3

    6 . 3 - 1 1 . 6

    8 . 1 - 1 1 .

    8

    10 8 16 1

    11.7-12.3

    %

    0

    38

    2 1

    8 2

    8 2

    4

    39

    88

    1 00

    1 00

    9 1

    9 9

    A. di effenbac hli

    L(l S.D.)

    9'.

    9 .

    1 0 .

    11.

    8 .

    8 .

    1 1 .

    1 6 .

    1 6 .

    1 6 .

    1 4 .

    9 (L36)

    2 (0.86)

    2 (0.99)

    1 (0.80)

    1 (0.91)

    4 (0.92)

    9 (1.30)

    2 (1.81)

    9 (2.79)

    7 (2.41)

    9 (4.00)

    Range

    6 .

    8 .

    8 .

    9 .

    7 .

    7 .

    9 .

    1 2 .

    1 1 .

    1 2 .

    1 0 .

    8-15.1

    1-12.5

    3-13.0

    4-13.6

    4-10.2

    2-10.4

    7-15.8

    9-21.9

    6-23.7

    6-22-3

    8-31.5

    As glass-eels

    are

    generally sedentary during their first year

    in

    fresh

    water, elvers which have been resident

    in

    streams

    and

    rivers

    for a

    year

    or more comprise most

    of the

    m igration. B ecause man y eels spend their

    first year

    in

    fresh water

    in the

    upper estuarine

    or

    tidal area,

    the

    greatest

    concentrations

    of

    elvers

    are

    recorded from

    the

    lower reaches

    of

    water-

    ways.As

    elvers make their

    way

    steadily upstream , their num bers become

    reduced

    by

    mortality

    and by

    diversion into tributaries

    or

    other suitable

    habitat.

    The

    most easily observed

    and

    well-known migrations

    are

    those

    whose progress

    is

    interrupted

    by

    hydro-electric dams.

    Karapiro

    Dam, 130 km

    upstream

    on the

    Waikato River,

    is the

    first

    of several power stations

    on

    this river.

    The

    ann ual elver m igration the re

    is sometimes great enough

    to

    clog

    the

    chambers

    of

    turbines shut down

    for maintenance. Smaller migrations

    are

    known

    at

    other hydro stations

    and waterfalls. Normally elvers

    can

    surmount such obstacles

    by

    leaving

    the water

    and

    climbing along splash zones

    of

    waterfalls

    or

    discharge

    chutes. Some falls

    can act as an

    effective barrier

    to the

    passage

    of

    elvers, particularly

    if

    there

    is a

    swift current rushing over bare smooth

    rock: Huka Falls

    on the

    upp er W aika to R iver prevent eels from enter-

    ing Lake Taupo (Cairns 1941).

    MA T E R I A L S

    AND

    ME T H O D S

    The elver migrations were sampled

    by the

    author

    at

    Karapiro

    Dam

    (February

    1.970,

    January

    1971,

    Feb ruary 1974),

    Te

    Kauwhata

    on the

    Waikato River (January 1971),

    and at two

    sites

    in the

    M akara Stream,

    Wellington, during January

    and

    February

    1972. The

    downstream site

    at

    the Makara Stream (Site

    a in

    Table

    1) was 1.2 km

    from

    the

    mouth ,

    while Site

    b was 2.5 km

    from

    the

    m outh . Elvers from seven ad ditional

    localities were obtained during January

    and

    February

    1971

    with

    the

    co-operation

    of

    hydro station staff

    (Fig. 1). The

    specimens from

    the

    Mokau River were collected

    by Mr J.

    Josephs

    of New

    Plymouth

    in

    January

    1971.

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    1977]

    J E L L Y M A N M I G R A T I N G E L V E R S

    63

    FIG. 1Locationsmen-

    tioned

    in the

    text

    at

    which elvers

    (Anguilla

    spp.) were sampled,

    1970-74.

    170 E

    Lake Cofetidge

    Aviemore*

    W a i t a k i *

    Roxburgh*

    Hydro station

    Further elvers from theW aikato R iver w ere collectedby the author

    at the E l b o w , 16 kmfrom theriver m ou th, w hile fishingfor glass-eels

    during August andSeptember 1970.Theseareincluded here,but, com-

    pared with the summer migration, thenumbersof elvers moving during

    the winter

    are

    small.

    The elversat the dams were densely concentrated, and samples could

    be scooped out with a hand-net or large container. At the E lbow ,

    Makara St ream, and M oka u Rive rs, set-nets were use d. Elvers were

    preserved with 7% formalin, or frozen, or sent to thelab orato ry alive

    in plastic bags containingalittle waterandinflated with oxyg en.

    MIGRATORY BEHAVIOUR

    PERIODICITY

    Most samples were collected during January andFeb ruary (Table1).

    Discussions with hydro station staff indicated

    a

    consistency

    in

    arrival

    times of elvers from year to year; C airns (1 941) recorded the first

    appearance

    of

    elvers

    at the

    lowest hy dro station

    on the

    W aikato River,

    then slightly above Karapiro, between 18 and 22Jan ua ry during four

    consecutive years, 1936-39. My data are similarly consistent: in 1970

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    64 N.Z. JOURNAL OF M AR INE & FRESHWATER RESEARCH [MARCH

    the run com men ced in m id -J an ua ry (precise date of first arrivals un-

    known), in 1971 on 18 January, in 1973 on 17 January, and in 1974 on

    15 Jan uary (no data are available for 1 972 ).

    Some elvers move upstream in the company of glass-eels during

    winter and. spring months. During August 1970 a sample of 206 elvers

    was caught am ong several thousan d glass-eels at the E lb ow . These

    were all shortfins (m ean length 8.6 cm, rang e 6.9-1 3.2 cm ). In ad dition ,

    during Septem ber-D ecem ber 1971, 94 elvers were caught at the mo uth

    of the Makara Stream among a total catch of 773 glass-eels: 93 were

    shortfins (m ean length 13.9 cm, range 8.2-27.4 c m ) , and some stom achs

    contained marine invertebrates. Probably, these elvers had been resident

    in the lower estuary or the adjacent sea.

    C L I M B I N G

    During visits to Ka rap iro Da m in 1970 and 197 1, little daytime activity

    of elvers was observed. However, power station operators report that

    during the peak of the season, elvers actively ascend the spillway and.

    face of the dam during daylight if rain is falling or the weather overcast.

    Large numbers of elvers were present at night. Larger eels of both

    species were also present. These would immediately move out of a

    strong torch beam, but elvers were less disturbed but would eventually

    leave an area of strong illumination.

    Concentrations of elvers were greatest where running water entered

    the main pool, and here the concrete was covered for a width of up

    to 3 m by a mass of elve rs. Progress against the spillway cu rre nt w as

    either in a series of vigorous darts forward or by crawling along the

    damp margin at the edge of the main flow.

    The method of vertical climbing consisted of a series of jerky move-

    ments, comm encing with the body forming a com pacted W shape.

    While the tail remained stationary, the head was extended forward and

    swung to one side. The hind part of the body was then pulled upward.

    The head alternately swung to the right and left, and so the body

    formed a W or an M shape. A fuller acco unt of similar vertical

    climbing by South African elvers is given by Skead (1959). Climbing

    elvers appeared to cling by both friction and surface tension, as elvers

    climbing vertically could not pass over any object which caused a

    substantial break in contact between themselves and the substrate.

    The lengths of elvers found, climbing a vertical wall were as follows

    with the figures in brackets being measurements of a random sample

    collected from the base of the spillway:

    Shortfins: m ean length 8.8 cm (9. 5), range 7.4-11.8 (7 .4- 1 2.9 ),

    n = 174 (227)

    Longfins: m ean length 9.6 cm (10 .8), range 7.9-12.0 (7 .7 -1 4.8 ),

    n = 50 (257)

    T he p ercen tage of eels less tha n 10 cm long in the clim bing sam ples

    was 9 1 % and 7 2 % for shortfins an d longfins respectively; equivalent

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    1977] JE L L Y M A N M I G RA T I N G E L V E RS 65

    figures for the general sample were shortfins 73% and longfins 28%.

    Small size is an advantage in climbing, as the surface area to weight

    ratio decreases with increasing size. The maximum length for vertical

    climbing is prob ably 12 cm, which corresp ond s to a w eight of 2.5 g.

    Elvers also showed distinct .thigmotaxis and crammed together in all

    crevices and pools on the spillway and elsewhere. These aggregations

    were partly enforced by the limited cover and resting areas available,

    but laboratory observations confirmed this social behaviour. When

    placed in an aquarium containing several sections of pipe as cover, the

    elvers did not disperse but congregated under one pipe. Thigmotaxis

    was also demonstrated in escape reactions. If the leading members of a

    column of climbing elvers were disturbed, their vigorous attempts to

    escape were transmitted through the whole column, and most elvers

    would rapidly re-enter deeper water.

    H A B I T A T PRE FE RE N C E S

    A comparison between glass-eel and elver catches for the Waikato

    River shows gross differences in the species composition for the two

    stages. Th us, the gran d me an for the E lb o w glass-eel samples (N =

    3004) recorded to early 1974 is 98% shortfin in contrast to 62% short-

    fin, elvers at K arap iro

    (N

    = 36 46 ). Elvers attempting to enter La ke

    Waikare (Te Kauwhata) were all shortfins (Table 1), and local fisher-

    men confirmed that this lake is inhabited by shortfins only.

    During electric fishing in Wellington streams and rivers, small eels of

    both species were frequently observed to show definite habitat prefer-

    ences.

    F or exam ple, the catch of eels less than 26 cm long o n coarse

    gravel and rock su bstrate in, the W aika nae River (Fig. 1) was 94 %

    longfin and 6% shortfin (n = 41 2), but samples from the W aimeha

    Stream , 2.5 km north, w here the b ottom is sil ty mu d, contained 9 7 %

    shortfins a nd 3 % longfins (n = 1016) .

    T o exam ine hab itat preferences am on g eels 26 cm and less long, a

    scries of five stations, chosen for consistency of bottom type was electric-

    fished in the Makara Stream. Although shortfins were predominant in

    all areas, longfins preferred stony areas with swift flows to sillier areas

    with slower flows. Thus, in two stations where the bottom type was silt

    an d the flow rates 0.12-0.18 m .s-

    1

    , 99% of all eels

    (N =

    136) were

    shortfins, whereas in two upstream stations where bottom types were

    line-coarse rubble and. flow rates were 0.41-0.48 m.s~\ only 76% of all

    eels

    (N

    178) we re shortfins. In bot h species (bu t mo re obviously in

    shortfins), the smaller size groups predominated in downstream areas;

    this observation supports the concept of successive summer migrations

    penetrating further up stream.

    Sig

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    66 N.Z .

    JOURNAL

    OF

    MARINE

    &

    FRESHWATER RESEARCH [MARCH

    TABLE 2Species proportions (%) and mean lengths (L, cm) of samples of migratory

    elvers

    (Anguilla

    spp.) from base of spillway of K ara pi ro Dam (samples arranged

    by mon ths over the migration season; * = data from W oods (1964); - = not given;

    .. = not calculated)

    DATE

    19 January 1971

    26 January 1971

    28 January 1971

    28 January 1971

    5 February 1970

    12 February 1970

    19 February 1974

    *27 February 1962

    *5 March 1963

    Total

    n

    276

    484

    657

    333

    376

    647

    254

    35 7

    314

    3698

    86

    47

    63

    65

    62

    48

    74

    74

    76

    63

    L

    8 .4

    9 . 5

    9 .3

    9 .3

    8 .9

    8 .9

    8 .9

    9 .3

    9 . 7

    9 . 1

    A.au stralis

    (1 S.D.)

    (0.83)

    (1.03)

    (0.97)

    (1.15)

    (0.73)

    (0.81)

    (2.34)

    (2.22)

    Range

    6 .7-10.6

    7 .4-12.9

    7 .1-12.5

    6 .9-12.4

    7 .1-11 .1

    6 .6 -11 .6

    6 .8-11 .1

    -

    -

    %

    14

    53

    37

    35

    38

    52

    26

    26

    24

    37

    A.

    L

    9 . 0

    10.8

    10.3

    10.3

    9 . 3

    9 . 4

    10.2

    9 . 8

    10.4

    10.0

    dieffenbachii

    (1 S.D.)

    (0.97)

    (1.29)

    (1.26)

    (1.13)

    (1.42)

    (1.14)

    (1.24)

    (2.24)

    -

    Range

    7 .3-12.2

    7 .7-14.8

    7 .8-15.1

    7 .3-13.3

    7 .0-14.4

    6 .8-13.9

    7 .7-15.3

    _

    COMPOSITION

    OF

    SAMPLES

    SPECIES PROPORTIONS

    T ab le 1 gives the percentag es of each species for th e various sam ples.

    M ost specimens (8 9% ) came from the N orth Island. H ere, 6 5 % of the

    catch were shortfins, whereas in the South Island shortfins comprised

    only 6%. However, these figures are not representative of the country

    as a whole, as habitat has a marked influence on species composition.

    The fluctuations which occur in the proportions of both species

    arriving at Karapiro Dam are shown in Table 2. No trend is obvious,

    but the overall predominance of shortfins is apparent.

    LENGTH

    T h e m ean lengths given in Table 1 indica te the variable size of

    migrating elvers at different localities. In all elver samples containing

    both species, the mean length of longfins exceeds that of the shortfins,

    as it does in glass-eels (Jellyman, in press). As expected, elvers recorded

    at upstream sites are larger than those from down stream on the same

    river. This is seen in both the Makara Stream and the Waikato River,

    but not from the Waitaki River (Waitaki and Aviemore samples),

    where numbers are small and cover a large size range (12cm). General-

    ly, the length-frequencies of elvers eorrespond well to a normal distribu-

    tion pattern; Fig. 2 shows four typical samples.

    In a large river system, several separate migrations may take place

    concurrently during summer, with waves of eels, grouped approximately

    by size, penetrating further upstream each year; migrations apparently

    cease at an app rox im ate length of 30 cm, wh en a m ore sedentary mo de

    of life is adopted. Table 2 gives the mean lengths of elver samples

    collected from the base of the spillway at Karapiro Dam. During 1971,

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    1977]

    JELLYMANMIGRATING ELVERS

    67

    A austral is

    A dieffenbachi i

    Karapiro

    Dam

    28-1-71

    n=415

    Dobson

    8-1-11

    n=27

    Roxburgh

    25-1-71

    6 8 10 12 14 16 18 20

    n = 147

    10

    12 14 16 18 20

    l e n g t h

    (cm)

    L e n g t h

    (cm)

    2Length-frequency distributions

    of

    samples

    of

    elvers

    (Anguilla spp.)

    from four

    localities.

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    68

    N.Z.

    JOURNAL

    OF

    M A R I N E

    &

    FRESHWA TER RESEARCH [MARCH

    TABLE

    3Age

    distributions and length ranges (cm) of elvers in Waikato and Mokau

    River samples (SF=short-finned eel

    Anguilla australis,

    LF=long-finned eel

    A. dieffenbachii;

    =number aged; - = nil)

    AREA

    Waikato

    'Elbow'

    DATE

    August

    1970

    Waikato January

    Te Kauwhata 1971

    Waikato

    Karapiro

    D am

    Mokau

    January

    1971

    January

    1971

    SPECIES

    SF n

    Range

    %

    SF

    n

    Range

    %

    SF n

    Range

    m

    LF n

    Range

    /o

    SFn

    Range

    /

    0

    11

    6.9-8.4

    57

    7

    6.5-7.3

    21

    8

    6.6-7.7

    8

    1

    7.8

    6

    1

    7.6

    2

    8

    7 .

    7

    8,

    8 .

    A G E

    1

    15

    .2-10.7

    39

    6

    5-9.6

    76

    10

    .9-9.8

    77

    15

    .2-10.4

    63

    4

    .5-9.3

    55

    GROUPS

    2

    4

    11.0-12.9

    4

    2

    10.4-11.0

    3

    7

    10.2-12.3

    15

    10

    9.9-12.4

    31

    4

    9.8-11..1

    43

    3

    -

    _

    -

    -

    -

    _

    L F

    n

    Range

    8.5-10.1 10.6-11.3 12.2-13.0

    57 30 13

    seasonal variations in length of 1.1 cm and 1.8 cm occurred for short-

    fins and longfins respectively. Also, an increase or decrease in mean

    length

    for one

    species

    is

    paralleled

    by an

    equal change

    in the

    other

    species: the two samples collectedon 28 Jan u ary 1971 (Table 2) were

    takenan hour apart,and gave almost identical analyses.

    A G E G R O U P S

    Age groups could not be distinguished from examination of the

    length-frequency data,and age determinations were m ade by examina-

    tion of burnt otoliths. If an overlap in body length occurred between

    successive

    age

    groups

    the

    mid-point

    of the

    overlap

    was

    taken

    as the

    limit between groups. Similarly, if successive ranges did not meet, the

    mid-point of the difference wasdefinedas thegro up limit. Resu lts from

    Waikato and M okau Rivers are presented in Table 3. Unfortunately,

    South Island specimens were all preserved in strong formalin, and the

    otoliths were generally unreadable. Roxburgh and Aviemore samples

    were examined, and both contained eels from age groups 3-7, but

    deteriorationof otoliths prevented reliable reading.

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    1977] JELLYM AN MIGR ATING ELVER S 69

    MORTALITY

    Numerous dead elvers and others in very poor condition were found

    at Karapiro Dam, and predation by larger longfins and catfish Ictalurus

    nebulosus

    (Le Sueur) was observed. Other predators include brown

    trout Salmo trutta L., shags (cormorants), ducks, kingfishers, and

    wa ter rats.

    In live samples, the proportion of longfins that died in transit far

    exceeded that of shortfins. A trial was run to compare survival rates

    of elvers of both species exposed to humid air in a constant temperature

    cabinet: three trays containing ten elvers of each species were used,

    and the deaths recorded over a week. At 15c, 70% of the longfins

    died, but no shortfins; at 20c, 90% of the longfins and 40% of the

    shortfins died; and at 25c, 100% of both species died, but the longfins

    died before the shortfins. Thus, longfins are less able to survive pro-

    longed periods of exposure which require cutaneous respiration,

    especially under the additional stresses of high temperature and crowd-

    ing.

    D I SC U SSI O N

    Summer migration of juvenile freshwater eels is not confined to New

    'Zealand.

    Skead (1959) reviewed several years' observations on the

    migration of elvers of

    A. mosscimbicci

    Peters in the B uffalo Rive r, So uth

    Africa. Summer migrations of Australian elvers are recorded by Kershaw

    (1911) and Whitley (1929). A similar phenomenon was witnessed in

    New Guinea by Herre (1930), and summer migrations of juvenile

    European eels have long been known.

    In New Zealand and apparently elsewhere, the stimuli which initiate

    migration are unknown. Although low temperatures (less than 10c)

    could inhibit mass movements of elvers, the fact that summer migrations

    occur within a few weeks of each other throughout the whole country

    indicates that it is not the reaching of any specific temperature which

    triggers migration. Temperatures recorded in the Makara Stream over

    the migratory period ranged from 14.2-21.5c, bu t the two ru n s

    recorded at the downstream site did not correspond with any common

    value or fluctuation. The lunar cycle appears to have no direct effect, as

    seen by the consistency of arrival times each year at Karapiro Dam.

    A small migration was recorded at M aka ra during a fresh in the

    stream, but larger movements took place when the stream level was

    normal. Hardy (1950) observed that increased water flow did not

    induce a larger number of elvers to climb the margins of a small water-

    fall in a North Canterbury stream.

    Migration may be initiated by the summation of several factors. A

    temperature greater than a threshold value, the increased day-length of

    summer, and a rapidly increasing tendency to school shown initially by

    Downlo

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    70 N.Z. JOURNAL OF M AR INE & FRESHWATER RESEARCH [MARCH

    a few elvers, may all contribute to the onset of migration. Increased

    water flow producing a strong rheotactic reaction among elvers may

    also be a contributing agent, but is not the causative one.

    The annual migration enables small eels to populate upstream areas.

    If suitable habitat is encountered, many eels probably choose to remain

    rather than progress further upstream. In this way, eels are able to

    disperse and occupy the available habitats along the complete river

    system, including territories vacated by migrant eels.

    The differences noted between species proportions of glass-eels from

    the E lbo w and elvers from K arap iro Dam , and also the age group

    distributions from the E lbo w samp le (see T ab le 3) , indicate that a

    considerable proportion of shortfins choose to spend at least the first

    2 y of freshw ater life in th e up pe r estua rine a rea s. Age class 0 w hich

    dom inates the Au gust E lb ow samples, is pro m ote d to age class 1

    in the January Te Kauwhata and Karapiro Dam samples, where i t is

    again dominant. In both these latter samples, age class 0 represents the

    previous year's glass-eels and is understandably more prevalent in the

    downstream (Te Kauwhata) sample than in the Karapiro Dam sample.

    Longfins, which are absent from the E lb o w and Te K auw hata

    samples, are also most strongly represented at Karapiro Dam by age

    class 1. Although longfins prefer swift and stony waters typical of

    upstrea m areas, substantial num bers of age gro ups 0 and 1 live down-

    stream of Karapiro Dam. Therefore, although a large percentage of

    shortfin glass-eels remain resident in the lower reaches of rivers, those

    that do migrate upstream do so more rapidly than longfins.

    Specific habitat preferences reduce intraspecific interaction and imply

    different feeding habits. Unfortunately, few data are available to com-

    pare the feeding habits of small eels of each species. It is not certain

    whether selective feeding occurs in areas where both species coexist:

    Cairns (1942) combined both species for the smallest size group he

    examined, and Hopkins (1970) assumed diets of both species were

    similar. Burnet (1952) showed that apparent feeding preferences in

    longfins largely reflected the abundance of organisms in the fauna.

    Thus, the suitability of the habitat rather than the availability of

    specific food types may be the primary requirement in habitat prefer-

    ence.

    Relative mortalities of asphixiated glass-eels and elvers suggest that

    longfins have higher oxygen demands than shortfins, but this has not

    been tested experimentally. If correct, the habitat preferences of both

    species would be plausibly explained. Also, the thicker dermal layer of

    longfins may result in less efficient cutaneous respiration than in short-

    fins. Under critical conditions, this and their possibly higher respiratory

    demands would lead to the increased mortality of longfins relative to

    shortfins.

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    1977] JELLYMANMIGRATING ELVERS 71

    ACKNOWLEDGMENTS

    I wish to thank the various personnel of the NewZealand Electricity Depart-

    ment

    and Mr J.

    Josephs

    of New

    Plymouth

    for

    providing elver samples.

    I

    also

    wish to thank Mr A. M. R. Burnet and Dr R. M. McDowall for adviceand

    criticism

    of the

    manuscript.

    LITERATURE CITED

    BURNET,

    A. M. R. 1952:

    Studies

    on the

    ecology

    of the New

    Ze ala nd long-finned

    eel, Anguilla dieffenbachii Gray. Australian Journal of Marineand

    Freshwater Research

    3

    (

    1

    )

    : 32-63.

    CAIRNS,

    D. 1941 :

    Life-history

    of the two

    species

    of New

    Ze alan d freshwater

    eel. Par t 1 -Taxonom y,

    age and

    growth, migration,

    and

    distribution.

    N.Z. Journal of Scienceand Technology, SeriesB, 23

    (

    2

    ) :

    53-72.

    1942: Life historyof the twospeciesof freshwater eel in NewZealand.

    II. Food,

    and

    inter-relationships with trout. N.Z.

    Journal

    of

    Science

    and Technology, Series

    B, 23

    (

    4

    )

    :

    132-48.

    HARDY, C. J. 1950: Summary of a report on eel trapping and observations at

    Lakes Taylor

    and

    Shcppard.

    North Canterbury Acclimatisation

    Society, Eighty-Sixth Annual Report:24-6.

    HERRE,

    A. W. 1930: A New

    Cuinea eel-fair.

    Science (

    New

    York

    ) 71

    : 16.

    HOPKINS,

    C. L. 1970:

    Some aspects

    of the

    bionomics

    of

    fish

    in a

    brown trout

    nursery stream.

    N.Z.

    Marine Department Fisheries Research Bulletin

    4.38 pp.

    JELLYMAN,

    D. J. (in

    p re ss ): Freshwater invasion

    of a New

    Zealand stream

    by

    glass-eels

    of two Anguilla spp.

    N.Z.

    Journal

    of

    Marine

    and

    Fresh-

    water Research.

    KERSHAW, J. A. 1911: Migration of eels in Victoria. Victorian Naturalist 27:

    196-201.

    SKEAD,

    C. J. 1959: The

    climbing

    of

    juvenile eels.

    Piscator

    13

    (

    46

    )

    :74-86.

    WHITLEY,

    G. P. 1929: An

    eel-fare

    at

    Parramatta.

    Australian Museum Magazine

    3

    (

    10

    )

    :

    348.

    WOODS,

    C. S. 1964:

    Fisheries aspects

    of the

    Tongariro power development

    project. N.Z. Marine Department Fisheries Technical Report 10.

    214

    pp.

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