Environmental biology of fishes, tập 92, số 4, 2011

One of the earliest papers he wrote on the origins of the New Zealand freshwater fish fauna McDowall 1964 was probably instrumental in his gaining theopportunity to study at the Museum o

Robert M McDowall —taxonomist and biogeographer

Don J Jellyman

Received: 19 April 2011 / Accepted: 23 June 2011 / Published online: 14 July 2011

# Springer Science+Business Media B.V 2011

Keywords Biogeography Taxonomy Systematics

New Zealand Galxaias Native fish Introduced fish

Robert (Bob) McDowall died in Christchurch, New

Zealand on 20 February 2011 after a short illness Bob

was a widely published author and an acknowledged

world authority on the taxonomy and biogeography of

freshwater fishes In a preface to Bob’s recent

magnum opus on the osteology of the galaxiids and

allied genera (McDowall and Burridge 2011), the

series editor, Peter Bartsch called him “a complete

zoologist” and listed his knowledge and experience

ranged from“taxonomy, biogeography and systematics

to reproductive biology, behaviour, ecology,

evolu-tion, fisheries and conservation biology” So, how

did this talented and productive fisheries scientist

come to acquire a profound knowledge of not only

the freshwater fish fauna of New Zealand but of the

southern hemisphere?

Bob was born on 15 September 1939 in Palmerston

North in 1939, the second youngest in a family of

five Bob attended Victoria University of Wellington

(1958–62), completing an M Sc in zoology As a

young scientist, Bob recognised he had available to

him a virtually unstudied fish fauna, the nativefreshwater fishes of New Zealand The fauna issparse (~ 40 species) and characterised by a highdegree of diadromy Bob soon realised that thisfauna offered an opportunity to“explore pattern andprocess, cause and effect, evolution and biogeogra-phy, in a way that would have been much moredifficult in areas with more speciose faunas”(McDowall2010)

One of the earliest papers he wrote on the origins

of the New Zealand freshwater fish fauna (McDowall

1964) was probably instrumental in his gaining theopportunity to study at the Museum of ComparativeZoology, Harvard University USA (1965–68) where

he was exposed to the teachings of the likes of ErnstMayr, P J Darlington, Giles Mead, Alfred Romer andother luminaries of that era His Ph D was on thesystematics and phylogeny of the New Zealandgalaxiids The family Galaxiidae comprises a group

of southern hemisphere fishes whose wide geographicrange and the diversity of habitats they have colonisedare somewhat akin to the northern hemispheresalmonids The galaxiids were destined to become amajor research focus for Bob

Upon completion of his doctorate and return to NewZealand, Bob was instructed to commence research onthe diet of brown trout, Salmo trutta, something heregarded of much lesser importance than understand-ing the ecology of native species With a dogmatismthat often characterised his dealings with authority, heignored this directive and established an extensive field

programme studying the ecology of the migrations of

juvenile galaxiids (known locally as “whitebait”, and

considered a delicacy) He also commenced a

compre-hensive review of the taxonomy of the New Zealand

native fish fauna, an exercise that culminated in the

publication of his first book (McDowall1978a; revised

and enlarged as McDowall 1990) Both books were

illustrated with Bob’s own drawings and photographs

In the 1970’s and 80’s as a science administrator

responsible for over 60 staff, he became “lost in the

wilderness of bureaucracy” (McDowall 2010) A

timely invitation to be a keynote speaker at the first

international conference on diadromous fishes

(Boston, 1986) reinvigorated his research interests

and he commenced a study of diadromy that involved

broad scale geographic, taxonomic and behavioural

perspectives A further book followed (McDowall

1988), but more importantly, preparation for the book

provided the motivation and direction for his many

papers on diadromy and its evolutionary and

biogeo-graphic significance Bob’s contribution to New

Zealand fisheries science was recognised when he

was made a Fellow of the Royal Society of New

Zealand in 1984 In 1991 he gladly relinquished his

administrative role and returned to science He

officially retired from the National Institute of Water

and Atmosphere (NIWA) in 1999 (Jellyman2002) to

take up a James Cook Research Fellowship by the

Royal Society of New Zealand—this gave him 2 years

of scientific freedom to pursue his interests in

taxonomy and biogeography Upon completion of this

fellowship, he remained working as an emeritus

scientist at NIWA until his death (Fig.1)

As well as research on the taxonomy of the New

Zealand freshwater fish fauna, Bob also worked on

the faunas of South Australia, New Caledonia,

Patagonia, South America, the Falkland Islands, and

at the time of his death was engaged in revising the

taxonomy of South African galaxiids He never lost

the wonder of what he called the“Aha” moments, the

sudden dawning of an insight into a problematic

issue, or the chance observation of a rare or unusual

biological event When he was an undergraduate he

was fortunate enough to observe the spawning of

redfin bullies (Gobiomorphus huttoni)—this was one

such defining moment, and an experience he spoke of

even 50 years later

Although he worked extensively on the systematics

and taxonomy of southern hemisphere freshwater

fishes, Bob regarded his most significant scientificachievements as the papers he wrote on thebiogeography and benefits of diadromy As a fellowichthyologist (Gene Helfman, Professor Emeritus,University of Georgia) wrote in his recent condolences,

“while Bob worked chiefly in New Zealand, hisimpact was global” Another significant contribu-tion of Bob’s was the establishment of NewZealand Freshwater Fish Database in 1979—thisdatabase currently contains over 30 000 records ofspecies and their habitats, and has formed the basis

of many papers on distributions and habitat use ofnative and introduced fish by Bob and others (e.g.,McDowall and Taylor 2000; Jowett and Richardson

2003; Leathwick et al.2005)

Being a man of forthright opinion and logical andpersuasive argument, Bob relished spirited debate,especially with other researchers who did not sharehis views on dispersal biogeography (e.g McDowall

1978b; Craw 1979) He thrived on challenge to theextent that he once entered a national television quizshow purely for the sake of the experience Through-out his career, Bob displayed an unswerving and oftendogmatic pursuit of the truth and scientific excellence

As a young graduate, he successfully opposed theproposed introduction of largemouth bass, Micro-pterus salmoides, into New Zealand (McDowall

1968), a bold move as it was in direct opposition tohis then boss He had a particular aptitude forexpansive thinking, being able to see the “wood” aswell as the “trees”, a talent that meant he frequentlycame into conflict with bureaucrats who thought interms of the immediate and affordable He refereedmany papers (over 50 in his last year alone) and hiscomments were sometimes blunt but invariably fair—

he had a wonderful command of written English andconsequently could not abide poor grammar, some-thing many authors discovered to their detriment

He served on editorial boards of The Journal ofRoyal Society of New Zealand Royal, Reviews inFish Biology and Fisheries, Journal of Biogeography,Zoosystematics and Evolution, and Smithiana (Journal

of the South African Institute of Biodiversity)

Although he was very supportive of the concept of

a non-government management of fish and gameresources (as in the New Zealand Fish and GameCouncils), he was also a strong advocate of profes-sionally trained managers and the role of science inmaking informed management decisions He main-

tained strong conservation ethics—in the mid-1970s,

when there was continuing activity to drain wetlands

and fell native forests in New Zealand, Bob was astaunch advocate of conservation of these resources—

Fig 1 Bob McDowall through the ages Top left: The McDowall

family, 1948 Bob is on the far left Top right: The young angler;

one of Bob ’s first trout caught by fly, Kaituna River, Lake

Rotoiti, New Zealand, 1950 Center left: Bob holding his son

Stephen, and his wife Ainslie with daughter Susan Wellington,

New Zealand, 1972 Center right: Bob working at his desk, NIWA, Christchurch, New Zealand, 1999 Bottom left: Bob catching Galaxias, Chatham Islands, New Zealand, October,

2010 Bottom right: Bob McDowall, December 2010

he maintained a consistent message to any who would

listen, that the way to conserve freshwater fish in New

Zealand was through conservation of their habitat

From his research on galaxiids he was very aware of a

parallel threat, that of the deleterious impact of exotic

organisms on locally adapted and vulnerable floras

and faunas like the New Zealand forests and

freshwater fishes Partly for these reasons he gained

election to the New Zealand Conservation Authority,

the agency charged with oversight and policy of the

Crown estate in New Zealand While he was only

able to serve 2 years on this council before ill health

forced his resignation, his contribution was significant

and widely appreciated

Bob had a passion for writing, as evidenced by his

230 papers (in 66 different journals), 14 books and

300+ reports and popular articles A remarkable

feature of these papers is that Bob was the sole author

of 3/4 of them, and of the 57 joint-authored ones,

there were only 14 where he wasn’t the first author—

such was his dominance in his field At the time of his

death, he had just completed a comprehensive review

of the osteology and taxonomic relationships of the

galaxiids, retropinnids, and their allies (McDowall

and Burridge2011), and an extensive (872 p) book on

historic and contemporary Maori freshwater fisheries

(McDowall 2011)—this book gave him the

opportu-nity to bring together his twin loves of biology and

art As a side interest, he had also almost completed a

book on the diet of early New Zealand explorers His

diversity of interests is also evident from the topics he

wrote on which included papers on taxonomy and

distribution of fish, ontogenetic changes in body size

and shape, impacts of introduced and invasive species,

fish and art, the advantages of diadromy, and many

more He took pleasure in having distinctive titles,

believing he needed to capture a readers interest at the

earliest opportunity—among his titles are captivating

phrases like“When galaxiid and salmonid fishes meet

—a family reunion in New Zealand”, ‘The ii’s have

it ”(a taxonomic paper), “In defence of the caring

male”, “Shoot first and then ask questions”, and

“Falklands: fact, fiction or fiddlesticks…”

He had a particular aptitude for making science

palatable to lay people—among his writings are a

series of 50 articles on topics ranging from fish

biology and taxonomy, conservation, the role of

predators, biographies, and angling memoirs,

pub-lished in New Zealand’s Fish and Game magazine He

also gave many lectures and keynote addresses inNew Zealand and overseas Among the many awardsand accolades he received throughout his career, hewas recently awarded the Le Cren medal from theBritish Isles Fisheries Institute, a medal awarded to anindividual who has “made a lifelong contribution toall aspects of the study of fish biology and/or fisheriesscience, with a focus on conservation, training orpublic understanding of the discipline”—this medalwill be awarded posthumously

Bob leaves a huge written legacy in the worlds offisheries taxonomy, biology and biogeography Hischallenge to his contemporaries has always been topush the boundaries of understanding by asking the

“why” questions—why has this distinctive behaviourevolved? What are the ecological benefits of thisadaptation? This refreshing approach to biology, aswell as his extensive publications will be enduringmemory to this gifted, productive, and often provoc-ative man who gave so much to fisheries science over

a 50 year research record Maori have a saying aboutthe death of a respected person, that a mighty tree hasfallen in the forest, but in so doing it creates theopportunity for new growth; Bob McDowall was such

a tree but his legacy lives on in his writings and themany lives he influenced

References

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Jellyman DJ (2002) Bob McDowall —his contribution to New Zealand ’s freshwater fish N.Z J Mar Freshw Res 36:1 –12

Jowett IG, Richardson J (2003) Fish communities in New Zealand rivers and their relationship to environmental variables NZ J Mar Freshw Res 37:347 –366

Leathwick JR, Rowe D, Richardson J, Elith J, Hastie T (2005) Using multivariate adaptive regression splines to predict the distributions of New Zealand ’s freshwater diadromous fish Freshwat Biol 50:2034 –2052

McDowall RM (1964a) Affinities and derivation of the New Zealand freshwater fish fauna Tuatara 12(2):59–67 McDowall RM (1968a) The proposed introduction of the large mouth black bass Micropterus salmoides (Lacepede) into New Zealand NZ J Mar Freshw Res 2:149–161 McDowall RM (1978a) New Zealand freshwater fishes—a guide and natural history Heinemann, Auckland, 230 pp McDowall RM (1978b) Generalized tracks and dispersal in biogeography Syst Zool 27:88 –104

McDowall RM (1988a) Diadromy in fishes Migrations

between freshwater and marine environments Croon

Helm, London, 308 pp

McDowall RM (1990a) New Zealand Freshwater Fishes: A Natural

History And Guide Heinemann-Reed, Auckland, 553 pp

McDowall RM (2010) New Zealand Freshwater Fishes In

Noakes DLG (ed) An Historical and Ecological

Biogeog-raphy, Vol 32 Fish and Fisheries Series Springer 449 pp

McDowall RM (2011) ‘Ikawai Freshwater fishes in Maori Culture

and Economy ’ Canterbury University Press 872 p

McDowall RM, Burridge CP (2011a) Osteology and

relation-ships of the southern freshwater lower eutelostean fishes.

Zoosyst Evol 87:5 –183

McDowall RM, Taylor MJ (2000a) Environmental indicators of

habitat quality in a migratory freshwater fish fauna.

Environ Manage 25:357 –374

The papers and books of Bob McDowall

Allibone RM, Crowl TA, Holmes JM, King TM, McDowall

RM, Townsend CR, Wallis GP (1996) Isozyme analysis of

Galaxias species (Teleostei: Galaxiidae) from the Taieri

River, South Island, New Zealand: a species complex

revealed Biol J Linn Soc 57:107–127

Crow SK, McDowall RM (in press) Ontogenetic changes in

morphology of flathead galaxiid fishes (Osmeriformes:

Galaxiidae) in South Island, New Zealand NZ J Mar

Freshw Res 45

Gross MR, Coleman RM, McDowall RM (1988) Aquatic

productivity and the evolution of diadromous fish

migra-tion Sci 239:1291 –1293

Jowett IG, Richardson J, McDowall RM (1996) Relative effects

of in-stream habitat and land use on fish distribution and

abundance in tributaries of the Grey River, New Zealand.

NZ J Mar Freshw Res 30:463 –475

Kelly DJ, McDowall RM (2004) Littoral invertebrate and fish

communities pp 25.1 –25.14 In: Harding JS, Mosley MP,

Pearson CP, Sorrell BK (eds) Freshwaters of New Zealand

New Zealand Hydrological Society: New Zealand

Limno-logical Society Christchurch, N.Z

King CM, Roberts CD, Bell BD, Fordyce RE, Nicoll RS,

Worthy TH, Paulin CD, Hitchmough RA, Keyes IW, Baker

AN, Stewart AL, Hiller N, McDowall RM, Holdaway RN,

McPhee RP, Schwarzhans W, Tennyson AJD, Rust S,

McAdie I (2009) Phylum Chordata: lancelets, fishes,

amphibians, reptiles, birds and mammals In: Gordon DP

(ed) New Zealand inventory of biodiversity Canterbury

University Press, Christchurch, pp 432 –551

Koehn JD, McDowall RM (2004) Invasive species: fish and

fisheries: workshop overview, then and now NZ J Mar

Freshw Res 38:383 –389

Lee DE, McDowall RM, Lindqvist JK (2007) Galaxias fossils

from Miocene lake deposits, Otago, New Zealand: the

earliest records of the Southern Hemisphere family

Galaxiidae (Teleostei) J Roy Soc NZ 37:109 –130

Li J, McDowall RM (2010) Phylogenetic position of the

enigmatic Lepidogalaxias salamandroides with comment

on the orders of lower euteleostean fishes Mol Phylogenet

Evol 57:932–936

McDowall RM (1962) A redescription of the fresh-water finned bully Gobiomorphus huttoni (Ogilby) Trans Roy Soc NZ Zool 3:3–15

red-McDowall RM (1964b) Affinities and derivation of the New Zealand freshwater fish fauna Tuatara 12:59 –67 McDowall RM (1964c) A bibliography of the indigenous freshwater fishes of New Zealand Trans Roy Soc NZ Zool 5:175 –182

McDowall RM (1964d) A consideration of the question “What are whitebait? ” Tuatara 12:134–146

McDowall RM (1964e) Studies on the biology of the red-finned bully Gobiomorphus huttoni (Ogilby) I—Habitat and species interrelationships Trans Roy Soc NZ Zool 4:175 –182 McDowall RM (1965a) The composition of the New Zealand whitebait catch, 1964 New Zeal J Sci 8:285–300 McDowall RM (1965b) Descriptive and taxonomic notes on Grahamichthys radiatus (Valenciennes), Eleotridae Trans Roy Soc NZ Zool 7:51–56

McDowall RM (1965c) A new species of Retropinna from Lake Omapere, North Auckland Rec Dominion Mus 5:89–91 McDowall RM (1965d) Studies on the biology of the red- finned bully Gobiomorphus huttoni (Ogilby) II—Breeding and life history Trans Roy Soc NZ Zool 5:177–196 McDowall RM (1965e) Studies on the biology of the red- finned bully Gobiomorphus huttoni (Ogilby) III—Food studies Trans Roy Soc NZ Zool 5:233–254

McDowall RM (1966a) Further observations on Galaxias whitebait and their relation to the distribution of the Galaxiidae Tuatara 14:12–18

McDowall RM (1966b) A guide to the identification of New Zealand freshwater fishes Tuatara 14:89–104

McDowall RM (1967a) New land-locked fish species of the genus Galaxias from North Auckland, New Zealand Breviora 265:1–11

McDowall RM (1967b) Some points of confusion in galaxiid nomenclature Copeia 1967:841–843

McDowall RM (1968b) The application of the terms mous and catadromous to the southern hemisphere salmonid fishes Copeia 1968:176–178

anadro-McDowall RM (1968c) Interactions of the nature and alien faunas of New Zealand and the problems of fish introductions Trans Am Fish Soc 97:1–11

McDowall RM (1968d) Oceanic islands and endemism Sys Zool 17:346–350

McDowall RM (1968e) The proposed introduction of the mouth Black Bass Micropterus salmoides (Lacepede) into New Zealand NZ J Mar Freshw Res 2:149–161

Large-McDowall RM (1968f) The status of Nesogalaxias nicus (Weber and de Beaufort) (Pisces: Galaxiidae) Breviora 286:1–8

neocaledo-McDowall RM (1968g) Galaxias maculatus (Jenyns), the New Zealand whitebait NZ Fish Res Bull 2, 84 pp

McDowall RM (1969a) Extinction and endemism in New Zealand land birds Tuatara 17:1–12

McDowall RM (1969b) A juvenile of Aplochiton Jenyns Copeia 1969:631–632

McDowall RM (1969c) Lunar rhythms in aquatic animals—a general review Tuatara 17:133–144

McDowall RM (1970a) Comments on a new taxonomy of Retropinna (Galaxioidei: Retropinnidae) NZ J Mar Freshw Res 4:312–324

McDowall RM (1970b) The galaxiid fishes of New Zealand.

Bull Mus Comp Zool 139:341–431

McDowall RM (1970c) A second species of Galaxias common

to Tasmania and New Zealand (Pisces: Galaxiidae) Rec

Dominion Mus 7:13 –19

McDowall RM (1971a) Fishes of the family Aplochitonidae J

Roy Soc NZ 1:31 –52

McDowall RM (1971b) The galaxiid fishes of South America.

Zool J Linn Soc 50:33 –73

McDowall RM (1971c) The identity of Eleotris radiata

Valenciennes (Pisces: Eleotridae) Copeia 1971:731 –732

McDowall RM (1972a) Freshwater fishes and rivers of Australia

(by J.S Lake) —a book review Tuatara 19:128–129

McDowall RM (1972b) The species problem in freshwater fishes,

and the taxonomy of diadromous and lacustrine populations

of Galaxias maculatus (Jenyns) J Roy Soc NZ 2:325 –367

McDowall RM (1972c) The taxonomy of estuarine and

brackish-lake Retropinna from New Zealand (Galaxioidei:

Retropinnidae) J Roy Soc NZ 2:501–531

McDowall RM (1973a) Designation of a lectotype for Eleotris

gobioides Valenciennes (Pisces, Eleotridae) Copeia

1973:169–170

McDowall RM (1973b) Galaxias indicus Day, 1888 —a nomen

dubium J Roy Soc NZ 3:191 –192

McDowall RM (1973c) Limitation of the genus Brachygalaxias

Eigenmann, 1928 (Pisces: Galaxiidae) J Roy Soc NZ

3:193 –197

McDowall RM (1973d) Proposed suppression of the name

Galaxias delfini Philippi 1895 (Pisces: Galaxiidae) Z.N.

(S.) 1877 Bull Zool Nom 30:88 –89

McDowall RM (1973e) Relationships and taxonomy of the

New Zealand torrentfish, Cheimarrichthys fosteri Haast

(Pisces: Mugiloididae) J Roy Soc NZ 3:199 –217

McDowall RM (1973f) The status of the South African galaxiid

(Pisces: Galaxiidae) Ann Cape Province Mus 9:91 –101

McDowall RM (1973g) Zoogeography and taxonomy Tuatara

20:88–96

McDowall RM (1974) Specialization in the dentition of the

Southern graylings-genus Prototroctes (Galaxioidei:

Pro-totroctidae) J Fish Biol 6:209–213

McDowall RM (1975a) Gerald Stokell’s Freshwater and

diadromous fishes of New Zealand in the context of his

other published works, with a bibliography of his

publications J Roy Soc NZ 5:219 –223

McDowall RM (1975b) A revision of the New Zealand species of

Gobiomorphus (Pisces: Eleotridae) Rec Nat Mus NZ 1:1 –32

McDowall RM (1976a) Fishes of the family Prototroctidae

(Salmoniformes) Aust J Mar Freshw Res 27:641 –659

McDowall RM (1976b) Notes on some Galaxias fossils from

the Pliocene of New Zealand J Roy Soc N Z 6:17 –22

McDowall RM (1976c) The role of estuaries in the life cycles

of fishes in New Zealand Proc NZ Ecol Soc 23:27 –32

McDowall RM (1976d) The status of Stromateus maculatus

Forster—a fish from New South Wales J Roy Soc NZ

6:489–491

McDowall RM (1976e) The taxonomic status of the Galaxias

populations in the Rio Calle Calle, Chile (Pisces:

Galaxiidae) Stud Neotr Fauna 11:173–177

McDowall RM (1977) The possible effects on fishes and

eucalypt enrichment of New Zealand beech forests NZ J

Forest 22:45 –52

McDowall RM (1978c) New Zealand freshwater fishes: a guide and natural history Heinemann Educational, Auckland,

230 pp McDowall RM (1978d) Generalised tracks and dispersal in biogeography Sys Zool 27:88 –104

McDowall RM (1978e) A new genus and species of galaxiid fish from Australia (Salmoniformes: Galaxiidae) J Roy Soc NZ 8:115 –124

McDowall RM (1978) Patterns in the derivation of a New Zealand fish fauna pp 203 –218 Proceedings of the international symposium on marine biogeography and evolution in the Southern Hemisphere, vol 1, DSIR, Auckland, N.Z.

McDowall RM (1978g) Sexual dimorphism in an Australian galaxiid Aust Zool 19:308 –314

McDowall RM (1979a) The centrolophid genus Tubbia (Pisces: Stromateoidei) Copeia 1979:733 –738

McDowall RM (1979b) Fishes of the family Retropinnidae (Pisces: Salmoniformes)—a taxonomic revision and syn- opsis J Roy Soc NZ 9:85–121

McDowall RM (ed) (1980a) Freshwater fishes of south-eastern Australia (New South Wales, Victoria and Tasmania) Reed, Sydney, 208 pp

McDowall RM (1980b) Mobil New Zealand nature series — freshwater fish Reed, Wellington, 80 pp

McDowall RM (1980c) Charles Douglas, explorer: his notes on freshwater fishes J Roy Soc NZ 19:311 –324

McDowall RM (1980d) First adults of Schedophilus maculatus Guenther, 1860 (Stromateoidei: Centrolophidae) J Roy Soc NZ 10:141 –151

McDowall RM (1980e) Freshwater fishes and plate tectonics in the south western Pacific Palaeog Palaeoc Palaeoe 31:337 –351

McDowall RM (1980f) Seriolella caerulea Guichenot, 1848 in New Zealand waters (Stromateoidei: Centrolophidae) J Roy Soc NZ 10:65 –74

McDowall RM (1981a) Foreword pp 197 In: Berra TM (ed) An atlas of distribution of the freshwater fish families of the world University of Nebraska Press, Lincoln, Nebraska McDowall RM (1981b) Nomina nuda and the availability of lizard names; Comment NZ J Zool 8:127–128

McDowall RM (1981c) The relationships of Australian freshwater fishes pp 1253 –1273 In: R.A Keast (ed.) Ecological Biogeography of Australia Mono Biol 41:1253 –1273 McDowall RM (1981d) A sub-dorsal fin pore-canal system in Schedophilus maculatus Günther (Pisces: Centrolophidae) Copeia 1981:492 –494

McDowall RM (1982) The centrolophid fishes of New Zealand (Pisces: Stromateoidei) J Roy Soc NZ 12:103 –142 McDowall RM (1983) La faune exotique des poissons d ’eau douce de Nouvelle Zelande son histoire et son impact Comp Rend Soc Biogeo 59:223 –230

McDowall RM (1984a) Trout in New Zealand waters: the biology and management of trout in New Zealand’s lakes and rivers Wetland, Wellington, 120 pp

McDowall RM (1984b) The New Zealand whitebait book Reed, Wellington, 210 pp

McDowall RM (1984c) Designing reserves for freshwater fish

in New Zealand J Roy Soc NZ 14:17 –27 McDowall RM (1984d) Exotic fishes: the New Zealand experience In: Courtenay WR, Stauffer JR (eds) Distri-

bution, biology and management of exotic fishes John

Hopkins University Press, Baltimore, USA, pp 200–214

McDowall RM (1984e) Galaxiidae pp 126–127 In: Daget J,

Gosse JP, Thys van den Audenaerde DFE (eds) Checklist

of the Freshwater Fishes of Africa Orstom, Paris

McDowall RM (1984) Southern hemisphere freshwater

salmo-niforms: development and relationships pp 150 –153 In:

Moser GH (ed) Ontogeny and systematics of fishes,

American Society of Ichthyologists and Herpetologists

McDowall RM (1985) River estuaries in the life cycles of New

Zealand fish species pp 557 –570 In: Yanez-Arancibia A

(ed) Fish community ecology in estuaries and coastal

lagoons Ciudad Universitaria, Mexico, UNAM Press

Mexico

McDowall RM (1987a) Book review: The biology of perch and

related fish By J.F Craig, 1987 333 pp Croom Helm Ltd.

NZ J Mar Freshw Res 21:647 –8

McDowall RM (1987b) Impacts of exotic fishes on the native

fauna In: Viner AB (ed) Inland Waters of New Zealand.

DSIR, Wellington, N.Z, pp 333–347

McDowall RM (1987c) The occurrence and distribution of

diadromy in fishes Am Fish Soc Symp 1:1–13

McDowall RM (1987d) Zoogeography —the native fishes In:

Viner AB (ed) Inland Waters of New Zealand DSIR,

Wellington, N.Z, pp 291 –306

McDowall RM (1988b) Diadromy in fishes: migrations

between freshwater and marine environments Croom

Helm, London, 308 pp

McDowall RM (1990b) New Zealand freshwater fishes: a natural

history and guide Heinemann Reed, Auckland, 553 pp

McDowall RM (1990c) Filling in the gaps —the introduction of

exotic fishes into New Zealand In: Pollard DA (ed)

Introduced and translocated fishes and their ecological

effects Australian Society for Fish Biology, Magnetic

Island, Townsville, Queensland, pp 69 –82

McDowall RM (1990d) Freshwater fishes and fisheries of New

Zealand: the angler’s Eldorado Rev Aquatic Sci 2:

281–341

McDowall RM (1990e) When galaxiid and salmonid fishes

meet- a family reunion in New Zealand J Fish Biol

37:35–43

McDowall RM (1991a) Freshwater fisheries research in New

Zealand: processes, projects, and people NZ J Mar

Freshw Res 25:393 –413

McDowall RM (1991b) The ii ’s may have it at the end—

patronyms should be amended only if demonstrably

incorrect NZ Natur Sci 18:25 –29

McDowall RM (1992a) Diadromy: Origins and definitions of

terminology Copeia 1992:248 –251

McDowall RM (1992b) Global climate change and fish and

fisheries: what might happen in a temperate oceanic

archipelago like New Zealand Geo-J 28:29 –37

McDowall RM (1992c) Particular problems for the conservation

of diadromous fish: viewpoint Aquat Cons 2:351 –355

McDowall RM (1993a) Implications of diadromy for the

structuring and modelling of riverine fish communities in

New Zealand NZ J Mar Freshw Res 27:453–462

McDowall RM (1993b) Native trout of western North America:

[book review] NZ J Mar Freshw Res 27:382–384

McDowall RM (1993c) New Zealand ’s quest for the rainbow’s

end: [letter to the editor] NZ J Mar Freshw Res 27:379 –380

McDowall RM (1993d) A recent marine ancestry for mous fishes? Sometimes yes, but mostly no! Environ Biol Fish 37:329–335

diadro-McDowall RM (1994a) Gamekeepers for the nation: the story

of New Zealand ’s acclimatisation societies 1861–1990 Canterbury University Press, Christchurch, 508 pp McDowall RM (1994b) Correction of the type locality of Stokell ’s smelt, Stokellia anisodon (Stokell) 1941 (Pisces: Retropinnidae) J Roy Soc NZ 24:261 –265

McDowall RM (1994c) Distinctive form and colouration of juvenile torrentfish, Cheimarrichthys fosteri (Pisces: Pinguipedidae) NZ J Mar Freshw Res 28:385 –390 McDowall RM (1994) Family Centrolophidae Fishes of Australia ’s south coast 832–838

McDowall RM (1994) Family Galaxiidae Fishes of Australia ’s south coast: 234 –238

McDowall RM (1994) Family Nomeidae Fishes of Australia ’s south coast: 838 –843

McDowall RM (1994) Family Prototroctidae Fishes of Australia’s south coast: 233

McDowall RM (1994) Family Retropinnidae Fishes of Australia’s south coast: 232–233

McDowall RM (1994i) Fish imagery in art 70: Polynesian Maori carving of the Maui myth Environ Biol Fish 41:330

McDowall RM (1994j) On size and growth in freshwater fish Ecol Freshw Fish 3:67 –79

McDowall RM (1994k) The origins of New Zealand ’s chinook salmon, Oncorhynchus tshawytscha Mar Fish Rev 56:1 –7 McDowall RM (1994l) Salmons and their allies In: Paxton JR, Eschmeyer WN (eds) Encyclopedia of fishes NSW University Press, Kensington, NSW, pp 113 –118 McDowall RM (1994m) The Tarndale Bully, Gobiomorphus alpinus Stokell (Pisces, Eleotridae) revisited and rede- scribed J Roy Soc NZ 24:117 –124

McDowall RM (1995) Seasonal pulses in migrations of New Zealand diadromous fish and the potential impacts of river mouth closure NZ J Mar Freshw Res 29:517–526 McDowall RM (ed) (1996a) Freshwater fishes of south-eastern Australia Reed, Sydney, 247 pp

McDowall RM (1996) Ayson, Lake Falconer 1855–1927: Farm labourer, rabbit inspector, acclimatisation officer, fisheries inspector pp 24 –25 The Dictionary of New Zealand biography Volume three, 1901 –1920, Auckland University Press, Auckland

McDowall RM (1996c) Diadromy and the assembly and restoration of riverine fish communities: a downstream view Can J Fish Aquat Sci 53:219 –236

McDowall RM (1996d) Etymology of some common names for New Zealand freshwater fishes Pacific Sci 50:117 –121 McDowall RM (1996e) Threatened fishes of the world: Prototroctes oxyrhynchus Gunther 1870 (Prototroctidae) Environ Biol Fish 46:60

McDowall RM (1996f) Volcanism and freshwater fish ography in the North Island of New Zealand J Biogeol 23:139–148

bioge-McDowall RM (1997a) An accessory lateral line in some New Zealand and Australian galaxiids (Teleostei: Galaxiidae) Ecol Freshw Fish 6:217 –224

McDowall RM (1997b) Affinities, generic classification and biogeography of the Australian and New Zealand mud-

fishes (Salmoniformes: Galaxiidae) Rec Aust Mus

49:121–137

McDowall RM (1997c) The evolution of diadromy in fishes

(revisited) and its place in phylogenetic analysis Rev Fish

Biol Fish 7:443 –462

McDowall RM (1997d) Indigenous vegetation type and the

distribution of shortjawed kokopu, Galaxias postvectis

(Teleostei: Galaxiidae), in New Zealand NZ J Zool

24:243 –255

McDowall RM (1997e) Note of the conservation status of the

giant bully, Gobiomorphus gobioides (Teleostei: Eleotridae).

J Roy Soc NZ 27:163 –172

McDowall RM (1997f) Two further new species of Galaxias

(Teleostei: Galaxiidae) from the Taieri River, southern New

Zealand J Roy Soc NZ 27:199 –217

McDowall RM (1998a) Driven by diadromy: its role in the

historical and ecological biogeography of the New

Zealand freshwater fish fauna It J Zool 65:73 –85

McDowall RM (1998b) Fighting the flow:

downstream-upstream linkages in the ecology of diadromous fish

faunas in West Coast New Zealand rivers Freshwat Biol

40:111–122

McDowall RM (1998c) Phylogenetic relationships and

ecomorphological divergence in sympatric and allopatric

species of Paragalaxias (Teleostei: Galaxiidae) in high

elevation Tasmanian lakes Environ Biol Fish 53:

235 –257

McDowall RM (1999) Book review: Fish migration and fish

bypasses Edited by M Jungwirth, S Schmutz and S.

Weiss Fishing News Books, Oxford, 1998 J Fish Biol

54:1137 –1138

McDowall RM (1999b) The caudal skeleton in Galaxias and

allied genera (Teleostei: Galaxiidae) Copeia 1999:932 –939

McDowall RM (1999c) Diadromy and genetic diversity in

Nearctic and Palearctic fishes Mol Ecol 8:527 –528

McDowall RM (1999d) Different kinds of diadromy: different

kinds of conservation problems ICES J Mar Sci 56:410–413

McDowall RM (1999e) Freshwater fishes as indicators of

habitat health in New Zealand: the implications of being

migratory In: Holt A, Dickinson K, Kearsley GW (eds)

Environmental indicators: Proceedings of the symposium.

University of Otago, Dunedin, Environmental Policy and

Management Research Centre, pp 83 –84

McDowall RM (1999f) Further feral poeciliid fish in New

Zealand fresh waters, with a key to species NZ J Mar

Freshw Res 33:673 –682

McDowall RM (2000a) The Reed field guide to New Zealand

freshwater fishes Reed, Auckland, 225 pp

McDowall RM (2000b) Biogeography of southern cool

temperate freshwater fishes: evidence from macroparasitic

faunas J Biogeol 27:1221 –1229

McDowall RM (2000c) Biogeography of the New Zealand

Torrentfish, Cheimarrichthys fosteri (Teleostei: Pinguipedidae):

a distribution driven mostly by ecology and behaviour.

Environ Biol Fish 58:119–131

McDowall RM (2000d) Gerald Stokell 1890–1972:

horticultur-alist, ichthyologist pp 499–500 In: Dictionary of New

Zealand biography Volume five, 1941–1960, vol 5 Allen

and Unwin: Department of Internal Affairs, Wellington

McDowall RM (2000e) Identity of the Smith of Galaxias

smithii Falk Is J 7:3 –7

McDowall RM (2001a) The Reed New Zealand nature series— freshwater fishes of New Zealand Reed, Auckland, 95 pp McDowall RM (2001b) Anadromy and homing: two life- history traits with adaptive synergies in salmonid fishes? Fish Fisheries 2:78 –85

McDowall RM (2001c) Diadromy, diversity and divergence: implications for speciation processes in fishes Fish Fisheries 2:278 –285

McDowall RM (2001d) Getting the measure of freshwater fish habitat in New Zealand Aquat Ecosys Health Manag 4:343 –355

McDowall RM (2001e) New Zealand stream invertebrates: ecology and implications for management: stream inver- tebrate ecology comes of age (book review) NZ Sci Rev 58:29 –30

McDowall RM (2001f) The origin of the salmonid fishes: marine, freshwater or neither? Rev Fish Biol Fish 11:171 –179

McDowall RM (2001g) Parioglossus (Teleostei: Gobioidei: Microdesmidae) in New Zealand NZ J Mar Freshw Res 35:165–172

McDowall RM (2001h) The principal caudal fin ray count: a fundamental character in the galaxioid fishes NZ J Zool 28:395 –405

McDowall RM (2002a) Accumulating evidence for a dispersal biogeography of southern cool temperate freshwater fishes J Biogeol 29:207 –220

McDowall RM (2002b) Book reviews: Migration of freshwater fishes, by M C Lucas and E Baras 420 pp Blackwell Science, Oxford, 2001 J Fish Biol 60:1613 –1614 McDowall RM (2002c) Centrolophidae - medusafishes (ruffs, barrelfishes) pp p 3767 –3770 The living marine resources

of the western central Pacific Fao, Rome McDowall RM (2002d) How many species of Galaxias are there in South Africa? Ichthos 67:10 –11

McDowall RM (2002e) Provenance and status of Galaxias smithii Regan (1905) (Teleostei: Galaxiidae) J Nat Hist 36:1129–1134

McDowall RM (2003a) Book review: New Zealand fish: a complete guide, by C Paulin, C Roberts, A Stewart, and

P McMillan NZ J Mar Freshw Res 37:219–220 McDowall RM (2003b) Hawaiian biogeography and the islands ’ freshwater fish fauna J Biogeol 30:207–220 McDowall RM (2003c) Impacts of introduced salmonids on native galaxiids in New Zealand upland streams: a new look at an old problem Trans Am Fish Soc 132:229 – 238

McDowall RM (2003d) In defence of the caring male Trends Ecol Evol 18:610 –611

McDowall RM (2003e) Protocols for enumerating the principal caudal rays of fishes NZ J Zool 30:259 –291

McDowall RM (2003f) Salmons and their allies In: Cogger

HC, Eschmeyer WN, Paxton JR, Zweifel RG (eds) Ecology of animals: reptiles, amphibians and fishes Fog City Press, San Francisco, pp 335–341

McDowall RM (2003g) Shads and diadromy: implications for ecology, evolution and biogeography Am Fish Soc Symp 35:11–23

McDowall RM (2003h) Variation in vertebral number in galaxiid fishes (Teleostei: Galaxiidae): a legacy of life history, latitude and length Environ Biol Fish 66:361 –381

McDowall RM (2003i) Variation in vertebral number in

galaxiid fishes how fishes swim, and a possible

explanation for pleomerism Rev Fish Biol Fish

13:247–263

McDowall RM (2004a) Ancestry and amphidromy in island

freshwater fish faunas Fish Fisheries 5:78 –85

McDowall RM (2004b) Book review: Gunn, J.M., Steedman,

R.J and R.A Ryder (ed.) Boreal shield watersheds —lake

trout ecosystems in a changing environment Lewis, Boca

Raton 501 pp Prog Phys Geog 28:608 –609

McDowall RM (2004c) The Chatham Islands endemic galaxiid:

a Neochanna mudfish (Teleostei: Galaxiidae) J Roy Soc

NZ 34:315 –331

McDowall RM (2004d) Letter to the Editor: Protocols for citing

the authority for zoological names: comment NZ J Mar

Freshw Res 38:95 –196

McDowall RM (2004e) Shoot first, and then ask questions: a

look at aquarium fish imports and invasiveness in New

Zealand NZ J Mar Freshw Res 38:503–510

McDowall RM (2004f) What biogeography is: a place for

process J Biogeol 31:345–351

McDowall RM (2005a) [Book Review]: Freshwater fishes of

north-eastern Australia By B Pusey, M Kennard, and A.

Arthington NZ J Mar Freshw Res 39:1196 –1197

McDowall RM (2005b) Diadromy, history and ecology: a

question of scale In: Dufour SP, Prévost E, Rochard E,

Williot P (eds) Fish and Diadromy in Europe (ecology,

management, conservation) Springer, Bordeaux, France,

pp 5 –14

McDowall RM (2005) Ernst Mayr: [obituary] pp 138 –141

Ann Rep Roy Soc NZ 2005

McDowall RM (2005d) Falkland Islands biogeography:

con-verging trajectories in the South Atlantic Ocean J Biogeol

32:49 –62

McDowall RM (2005e) Falklands: fact, fiction or fiddlesticks: a

place for process J Biogeol 32:2187 –2187

McDowall RM (2005f) Historical biogeography of the New

Zealand freshwater crayfishes (Parastacidae, Paranephrops

spp.): restoration of a refugial survivor? NZ J Zool 32:55–

77

McDowall RM (2005g) Letter to the Editor Comment:

American Fisheries Society makes changes in the end.

NZ J Mar Freshw Res 39:1199 –1200

McDowall RM (2005h) Making a living in Red Pond: a

snapshot of the diet of a populaion of Aplochiton zebra

(Teleostei: Galaxiidae) at the Falkland Islands NZ J Zool

32:23 –27

McDowall RM (2006a) Crying wolf, crying foul, or crying

shame: alien salmonids and a biodiversity crisis in the

southern cool-temperate galaxioid fishes? Rev Fish Biol

Fish 16:233 –422

McDowall RM (2006b) Fish, fish habitats and fisheries in New

Zealand Aquat Ecosys Health Manag 9:391 –405

McDowall RM (2007) Hawaiian freshwater fishes: the role of

amphidromy in history, ecology and conservation biology.

pp 3–9 In: Evenhuis NL and Fitzsimons JM (ed) Biology

of Hawaiian streams and estuaries

McDowall RM (2007b) On amphidromy, a distinct form of

diadromy in aquatic organisms Fish Fisheries 8:1–13

McDowall RM (2008a) Jordan ’s and other ecogeographic rules,

and the vertebral number in fishes J Biogeol 35:501 –508

McDowall RM (2008b) Process and pattern in the biogeography

of New Zealand—a global microcosm? J Biogeol 35: 197–212

McDowall RM (2008c) Why are so many boreal freshwater fishes anadromous? Confronting `conventional wisdom ’ Fish Fisheries 9:208 –213

McDowall RM (2009a) Concluding remarks on the Halifax symposium Am Fish Soc Symp 69:813 –821

McDowall RM (2009b) Early hatch: a strategy for safe downstream larval transport in amphidromous gobies Rev Fish Biol Fish 19:1 –8

McDowall RM (2009c) An ichthyological lucky dip, or “What freshwater fish lives where? ”: Review of: T M Berra, Freshwater fish distribution Environ Biol Fish 4:147 – 151

McDowall RM (2009) Kenneth (Radway) Allen MSc DSc Cambridge FRSNZ Roy Soc N.Z Yrbk 2009

McDowall RM (2009e) Making the best of two worlds: diadromy in the evolution, ecology and conservation of aquatic organisms Am Fish Soc Symp 69:1–22

McDowall RM (2010b) New Zealand freshwater fishes: an historical and ecological biogeography Springer, Dordrecht,

449 pp McDowall RM (2010c) Biogeography in the life and literature

of John Muir: a ceaseless search for pattern J Biogeol 37:1629 –1636

McDowall RM (2010d) Why be amphidromous: expatrial dispersal and the place of source and sink population dynamics? Rev Fish Biol Fish 20:87 –100

McDowall RM (2011b) Ikawai: Freshwater fishes in Mäori culture and economy Canterbury University Press, Christchurch,

872 pp McDowall RM (2011c) Dispersal ability: animal In: Simberloff

D, Rejmánek M (eds) Encyclopedia of biological sions University of California Press, Berkeley, pp 54 –159 McDowall RM, Allibone RM (1994) Possible competitive exclusion of common river Galaxias (Galaxias vulgaris)

inva-by koaro (G brevipinnis) following impoundment of the Waipori River, Otago, New Zealand J Roy Soc NZ 24:161–168

McDowall RM, Allibone RM (2004) Threatened fishes of the world: Galaxias cobitinis McDowall and Waters, 2002 (Galaxiidae) Env Biol Fish 70:42

McDowall RM, Banister KE (1985) Pikes, salmon, argentines.

pp 44 –59 In: Banister KE, Campbell A (eds) The encyclopedia of underwater life Allen Unwin, London,

287 pp McDowall RM, Banister KE (1986) Pikes, salmon, argentines.

pp 52 –67 In: Banister KE, Campbell A (eds) All the worlds ’ animals—fishes Torster Books, New York, 158 pp McDowall RM, Burridge CP (2011b) Osteology and relation- ships of the southern freshwater lower euteleostean fishes Zoosyst Evol 87:7 –185

McDowall RM, Chadderton WL (1999) Galaxias gollumoides (Teleostei: Galaxiidae), a new fish species from Stewart Island, with notes on other non-migratory freshwater fishes present on the island J Roy Soc NZ 29:77–88 McDowall RM, Charteris SC (2006) The possible adaptive advantages of terrestrial egg deposition in some fluvial diadromous galaxiid fishes (Teleostei: Galaxidiae) Fish Fisheries 7:153 –164

McDowall RM, David BO (2008) Gobiopterus in New Zealand

(Teleostei: Gobiidae), with observations on sexual

dimor-phism NZ J Mar Freshw Res 42:325–331

McDowall RM, Eldon GA (1980) The ecology of whitebait

migrations: (Galaxiidae: Galaxias spp.) NZ Fish Res Bull

20, 170 pp

McDowall RM, Eldon GA (1996) Threatened fishes of the world:

Neochanna burrowsius (Phillipps, 1926) (Galaxiidae).

Environ Biol Fish 47:190

McDowall RM, Eldon GA (1997) Reproductive cycling and

fecundity estimation in the upland bully J Fish Biol

51:164 –179

McDowall RM, Frankenberg RS (1981) The Galaxiid fishes of

Australia (Pisces: Galaxiidae) Rec Aust Mus 33:443 –605

McDowall RM, Fulton W (1978a) A further new species of

Paragalaxias from Tasmania (Pisces: Galaxiidae) with a

revised key to species Aust J Mar Freshw Res 29:659 –

665

McDowall RM, Fulton W (1978b) A revision of the genus

Paragalaxias Scott (Salmoniformes: Galaxiidae) Aust J

Mar Freshw Res 29:93–108

McDowall RM, Gosztonyi AE (1974) Zoogeography of Galaxias

maculatus in South America Copeia 1974:978 –979

McDowall RM, Kelly GR (1999) Date and age at migration in

juvenile giant kokopu, Galaxias argenteus (Gmelin)

(Tele-ostei: Galaxiidae) and estimation of spawning season NZ

J Mar Freshw Res 33:263 –270

McDowall RM, Lee DE (2005) Probable perciform fish scales

from a Miocene freshwater lake deposit, Central Otago,

New Zealand J Roy Soc NZ 35:339 –344

McDowall RM, McIntosh A (2008) Freshwater fishes In:

Winterbourn MJ, Knox GA, Burrows C, Marsden ID (eds)

The Natural history of Canterbury Canterbury University

Press, Christchurch, pp 616 –650

McDowall RM, Nakaya K (1987) Identity of the galaxioid

fishes of the genus Aplochiton Jenyns from southern

Chile Jpn J Ichthyol 34:377–383

McDowall RM, Nakaya K (1988) Morphological divergence in

the two species of Aplochiton Jenyns (Salmoniformes:

Aplochitonidae): a generalist and specialist Copeia

1988:233–236

McDowall RM, Pankhurst NW (2005) Loss of negative

eye-size allometry in a population of Aplochiton zebrai

(Teleostei: Galaxiidae) from the Falkland Islands NZ J

Zool 32:17 –22

McDowall RM, Pole M (1997) A large galaxiid fossil

(Tele-ostei) from the Miocene of Central Otago, New Zealand J

Roy Soc NZ 27:193 –198

McDowall RM, Pusey BJ (1983) Lepidogalaxias

salaman-droides Mees —A redescription, with natural history notes.

Rec West Aust Mus 11:11 –23

McDowall RM, Rowe DK (1996) Threatened fishes of the

world: Galaxias gracilis McDowall,1967 (Galaxiidae).

Environ Biol Fish 46:280

McDowall RM, Stevens MI (2007) Taxonomic status of the

Tarndale bully Gobiomorphus alpinus (Teleostei: Eleotridae),

revisited-again J Roy Soc NZ 37:15–29

McDowall RM, Stewart AL (1997) Further specimens of

Agrostichthys parkeri (Teleostei: Regalecidae) with natural

history notes In: Séret B, Sire JY (eds) 5th Indo-Pacific Fish

Conference, Société Française d ’Ichtyologie: Institut de

recherche pour le dévelopment Nouméa, New Caledonia,

pp 165–176 McDowall RM, Suren AM (1995) Emigrating larvae of koaro, Galaxias brevipinnis Gunther (Teleostei: Galaxiidae), from the Otira River, New Zealand NZ J Mar Freshw Res 29:271 –275

McDowall RM, Taylor MJ (2000b) Environmental indicators of habitat quality in a migratory freshwater fish fauna Environ Manag 25:357 –374

McDowall RM, Wallis GP (1996) Description and redescription

of Galaxias species (Teleostei: Galaxiidae) from Otago and Southland J Roy Soc NZ 26:401 –427

McDowall RM, Waters JM (2002) A new longjaw Galaxias species (Teleostei: Galaxiidae) from the Kauru River, North Otago, New Zealand NZ J Zool 29:41 –52 McDowall RM, Waters JM (2003) A new species of Galaxias (Teleostei: Galaxiidae) from the Mackenzie Basin, New Zealand J Roy Soc NZ 33:675 –691

McDowall RM, Waters JM (2004) Phylogenetic relationships in

a small group of diminutive galaxiid fishes and the evolution of sexual dimorphism J Roy Soc NZ 34:23–57 McDowall RM, Whitaker AM (1975) The freshwater fishes In: Kuschel G (ed) Biogeography and ecology in New Zealand Junk, The Hague, pp 277 –299

McDowall RM, Allibone RM, Chadderton WL (2005) Falkland Islands freshwater fishes: a natural history Falklands Conservation, London, 102 pp

McDowall RM, Robertson DA, Saito R (1975) Occurrence of galaxiid larvae and juveniles in the sea N.Z J Mar Freshw Res 9:1 –9

McDowall RM, Hopkins CL, Flain M (1976) Fishes In: Jolly

VH, Brown JMA (eds) New Zealand lakes Auckland University Press, Auckland, pp 292 –307

McDowall RM, Graynoth E, Eldon GA (1977) The occurrence and distribution of fishes in streams draining the beech forests of the West Coast and Southland, South Island, New Zealand J Roy Soc NZ 7:405–424

McDowall RM, Clark BM, Wright GJ, Northcote TG (1993) Trans-2-cis-6-nonadienal: the cause of cucumber odor [sic]

in Osmerid and Retropinnid smelts Trans Am Fish Soc 122:144–147

McDowall RM, Mitchell CP, Brothers EB (1994) Age at migration from the sea of juvenile Galaxias in New Zealand (Pisces: Galaxiidae) Bull Mar Sci 54:385 –402 McDowall RM, Main MR, West DW, Lyon GL (1996) Terrestrial and benthic foods in the diet of the shortjawed kokopu, Galaxias postvectis Clarke (Teleostei: Galaxii- dae) NZ J Mar Freshw Res 30:257 –269

McDowall RM, Jellyman DJ, Dijkstra LH (1998a) Arrival of an Australian anguillid eel in New Zealand: an example of transoceanic dispersal Environ Biol Fish 51:1 –6 McDowall RM, Paxton JR, Eschmeyer WN (1998b) Salmons and their allies In: Paxton JR, Eschmeyer WN (eds) Encyclope- dia of fishes Academic, San Diego, CA, pp 113 –117 McDowall RM, Allibone RM, Chadderton WL (2001) Issues for the conservation and management of Falkland Islands freshwater fishes Aquat Conservat 11:473–486

McDowall RM, Kennedy EM, Alloway BV (2006a) A fossil southern grayling, genus Prototroctes, from the Pleisto- cene of north-eastern New Zealand (Teleostei: Retropinni- dae) J Roy Soc NZ 36:27 –36

McDowall RM, Kennedy EM, Lindqvist JK, Lee DE, Alloway

BV, Gregory MR (2006b) Probable Gobiomorphus fossils

from the Miocene and Pleistocene of New Zealand

(Teleostei: Eleotridae) J Roy Soc NZ 36:97–109

McIntosh AR, McDowall RM (2004) Fish communities in

rivers and streams pp 17.1 –17.9 In: Harding JS, Mosley

MP, Pearson CP, Sorrell BK (eds) Freshwaters of New

Zealand New Zealand Hydrological Society: New Zealand

Limnological Society, Christchurch

Metcalfe JD, Arnold G, McDowall RM (2002) Migration In:

Hart PJB, Reynolds JD (eds) Handbook of fish biology

and fisheries Blackwell Publishing England, Oxford, pp

175 –199

Riehl R, McDowall RM, Greven H (2002) Zur biologie von drei Galaxiiden-Arten Neuseelands Verh Aquarienfische 2:123–129

Saruwatari T, Oohara I, Orr JW, Kobayashi T, McDowall RM (2000) Phylogeny of lower Euteleosts reconstructed from mtDNA analysis DNA Polym 8:96 –101

Waters JM, McDowall RM (2002) Phylogenetic placement of retropinnid fishes: Data set incongruence can be reduced

by using asymmetric character state transformation costs Sys Biol 51:432 –449

Waters JM, McDowall RM (2005) Phylogenetics of the australasian mudfishes: Evolution of an eel-like body plan Mol Phylogenet Evol 37:417 –425

Movements of hatchery-reared lingcod released on rocky

reefs in Puget Sound

Jonathan S F Lee&Barry A Berejikian&Michael B Rust&Ken Massee&

Terry Wright&Kyle Brakensiek&Scott Steltzner&H Lee Blankenship

Received: 21 January 2010 / Accepted: 18 May 2011 / Published online: 5 August 2011

# Springer 2011

Abstract Fourteen sub-adult hatchery-reared lingcod

(Ophiodon elongatus) were released onto reefs in

South Puget Sound, Washington, USA to evaluate

their movement behavior Acoustic telemetry revealed

variation in movement among individuals that was

related to body size Larger lingcod tended to leave

the release reef sooner than smaller lingcod Four

lingcod left the reefs less than 10 days after release,

while three lingcod left between one and 4 months

after release Seven lingcod remained at the release

reefs for the entire 5-month study, though they didmake apparent short-term (< 24 h duration) excur-sions away from the reefs Data suggest that thefrequency and duration of excursions increase withage and size in both wild and hatchery lingcod.Movement data from these hatchery lingcod andpreviously published studies on wild lingcod arecompared

Keywords Ophiodon elongatus Acoustic telemetry Stock enhancement Hatchery Behavior

Environ Biol Fish (2011) 92:437 –445

DOI 10.1007/s10641-011-9859-2

J S F Lee ( *):B A Berejikian:M B Rust:K Massee

National Marine Fisheries Service, Northwest Fisheries

Science Center, Manchester Research Station,

P.O Box 130, Manchester, WA 98353, USA

e-mail: jon.lee@noaa.gov

T Wright

Northwest Indian Fisheries Commission,

6730 Martin Way East,

Squaxin Island Tribe,

S.E 3100 Old Olympic Highway, Box 3, Shelton,

necessary for fish to display natural movement

patterns as adults (Pacific salmon: Dittman and

Quinn1996) Further, some species of wild fish that

are translocated have shown homing behavior back

to the original site of capture (rockfish and lingcod:

Matthews1990, 1992; Yamanaka and Richards 1993;

Reynolds et al 2010), raising the question of

whether hatchery-reared fish will display fidelity to

the release site after they are “displaced” from the

hatchery

Stock enhancement may be a potential tool for the

management of lingcod (Ophiodon elongatus Girard),

which are economically important to commercial and

recreational fisheries along the west coast of North

America, from Alaska to California Due in part to

overfishing, average catch declined 36% from a

coastwide average annual catch of 4501 mt between

1980 and 1989 to an average annual catch of 2876 mt

between 1990 and 1997 (Jagielo and Wallace 2005)

Beginning in 1998, fishery restrictions were enacted

and coastal lingcod were declared rebuilt in 2005

(Jagielo and Wallace2005) Similarly, lingcod

histor-ically made important contributions to fisheries in

Puget Sound, a large glacially formed estuary in

Washington State (Bargmann1982) However, while

lingcod stocks have been declared rebuilt on the

coast, strong fishing restrictions remain on Puget

Sound lingcod

Stock enhancement programs can take many years

to carefully plan, develop, evaluate, and refine

(Blankenship and Leber 1995) As a first step,

small-scale releases may improve assessments of the

potential for released hatchery fish to survive, behave

naturally, and bolster fish stocks In this study,

telemetry-tagged hatchery lingcod were released into

South Puget Sound and tracked with acoustic

receivers to determine how long hatchery-reared fish

would survive and remain near the release sites

Materials and methods

Egg collection and rearing are detailed in Rust et al

(2005) Briefly, wild-spawned eggs were collected in

winter 2000 from the marine area south of the Tacoma

Narrows Bridge (South Puget Sound, hereafter SPS)

and reared at the NOAA, Northwest Fisheries Science

Center, Manchester Research Station near Port

Or-chard, Washington, USA (Fig.1) Eggs were brought

into 20 L incubators, and transferred to mesocosmbags at hatch Lingcod were transferred to shore-based tanks at metamorphosis, and then to net pens atapproximately 150–250 mm

On 2 August 2004, 14 age-4 lingcod (seven males,522–575 mm; seven females, 602–719 mm) wereimplanted with acoustic tags (VEMCO, V13P-1H-

69 k coded transmitter, dimensions: 44 mm length by

13 mm diameter) Acoustic tags emit long-rangeacoustic pulses that allow continual monitoring of ananimal’s movement The tags allowed for the uniqueidentification of each tagged individual, and alsoemitted depth data The acoustic pulses were emittedevery 40–120 s throughout the life of the tag (326 dfor acoustic emissions and 244 d for depth sensor dataemissions)

Methods for surgical intraperitoneal implantation

of acoustic tags in adult lingcod followed thosesuccessfully used by Griffin (2000) Tags wereinserted through an approximately 1.5 cm incisionmidway between the pectoral fins and anus, andgently moved anteriorly before suturing Sex, length,and weight were recorded at the time of tagging.Males were identified by the presence of narrow andpointed genital papillae After surgery, fish were heldfor several days of pre-release monitoring One fishdied within 48 h post-surgery Its acoustic tag wasrecovered and implanted in another hatchery lingcod

of the same sex

On 12 August 2004, the 14 tagged lingcod weretransported with a boat to two reefs comprised ofrocky relief outcroppings (Fig 1) Zee’s Reef wasdesignated a marine conservation area in 2002 (W.Palsson, pers comm.) by the Washington Department

of Fish and Wildlife (WDFW); approximate depths inthe area ranged from 3 to 25 m The second releasereef was Itsami Reef, an artificial reef constructed byWDFW near Johnson Point (Hueckel and Buckley

1987; Fig 1) Depths in the area generally rangedfrom 9 to 40 m At each reef, the fish were placed inmesh bags and brought by SCUBA divers to thebottom for release Four females and three males werereleased at Zee’s Reef; four males and three femaleswere released at Itsami Reef

Pulses from the acoustic tags were recorded bysubmerged VEMCO VR2 acoustic receivers Similarresearch using this acoustic equipment has indicatedhigh detection probabilities up to a range of 500 m(VEMCO Inc., pers comm.) Two receivers were

deployed at each release site (Fig.1) in an attempt to

cover the entire reef and to provide redundancy in

case of receiver failure or loss Each receiver was

attached midway on a 7.6 m line that was anchored to

a 1 m auger and suspended in the water column by a

28 cm diameter trawl float Receivers were deployed

by SCUBA divers at Zee’s reef on 24 June 2004, and

at Itsami Reef on 7 August 2004 Receivers were

removed for data retrieval and replaced by SCUBA

divers at Zee’s reef on 21 September 2004, and at

Itsami Reef on 25 September 2004 Final recovery of

all acoustic receivers occurred from 22 to 23 January

2005 All receivers were functioning at the time of

recovery Data were also accessed from VR2 receivers

deployed by other researchers for separate projects,

allowing for the detection of movement to areas off

the release sites (Fig.1)

Detection data can be erroneously interpreted as fish

survival and site residency when an active acoustic tag

is expelled or if a fish dies and remains within the range

of receiver detection A fish was considered to be alive

at least until their last detection on a new receiver(indicating fish movement) or to the last time its tagtransmitted a depth change that could not be explained

by the predicted tidal height plus tag resolution (0.44 mresolution) A predator that consumes a tagged fish andleaves the range of receivers could be erroneouslyinterpreted as a tagged fish that leaves the site However,since predators would excrete the tag within severaldays, any additional tag movement after several dayswould confirm that the fish was alive when it left thesite Survival rates cannot be determined once a fishleaves the range of the receivers; therefore estimatedsurvival durations for each fish should be consideredminimums Equipment and acoustic properties canperiodically result in false tag detections Typically, afalse detection may be indicated by a single detectionthat is not accompanied by another that same day(Dagorn et al.2006) A fish was therefore considered to

be present at the release reef on a particular day only if

Fig 1 Map of South Puget

Sound, Washington, USA,

with approximate locations

of submerged acoustic

receivers and release sites.

The pairs of receivers

flanking each release site

were closer to the release

sites than depicted but were

spaced further apart on the

figure to prevent symbols

from overlapping Plus

symbols represent release

sites

there were two or more detections that day (see also

Starr et al 2004, 2005) Since body size, sex, and

location differences have been suggested to affect

movement in lingcod (Hart 1943; Mathews and

LaRiviere 1987; Jagielo 1990), general linear models

(binomial distribution, logit link) were used to test

whether the proportion of days spent at the release reef

(days spent at reef divided by duration of study) was

predicted by body length at release, sex, site, or the

interaction between body length and sex Male body

length did not overlap with female body length, so

body length and sex were perfectly confounded

Consequently, we centered body length by sex and

site for analyses

The method of assigning presence at the reef based

on a minimum of two acoustic detections per day is

sensitive to excursions away from the reef that lasted

more than one day (Starr et al 2004; Starr et al

2005) Lags between detections occurring on the

same day were interpreted as fish movement away

from the reef (“apparent short excursions”) However,

lags can also occur when a fish is inside the

monitored area if an acoustic pulse is temporarily

obscured by an acoustic shadow (in structurally

complex habitat) or ambient acoustic noise such as

that from turbulent water conditions, boat traffic, or

other acoustic tags (Heupel et al.2006; Simpfendorfer

et al 2008) To minimize these false absences, lags

that lasted less than one hour were excluded from

analyses While the possibility of false absences

cannot be ruled out, the results from this study show

that lags between detections correlate with biological

variables, which would not be expected if the lags

were due to abiotic interference This lends support to

the interpretation that lags in this study represent real

fish movement away from receivers Only lingcod

that were detected at least twice per day, every day, for

the duration of the study, were analyzed because only

this group contained enough individuals (n=7) and a

wide-enough range of data (several months) to allow

statistical analyses A generalized linear mixed model

(Poisson distribution, log link) was used to test

whether the number of days since release and body

length at release predicted the frequency (number per

day) of apparent short excursions A linear mixed

model was used to test whether apparent short

excursion duration was significantly predicted by

body length at release or the number of days since

release Data were log-transformed The residuals

were slightly skewed on a normal probability plot,but this was accepted given the robustness of linearmodels and the strength of the resulting p-values(Neter et al 1996)

Results

Seven of the 14 lingcod were present at the reefs everysingle day throughout the duration of the study(hereafter termed‘residents,’ Table1) All of these fishhad depth changes greater than would be expectedfrom an expelled tag or dead fish, within one day ofthe end of the study, indicating that the fish were alive(Table 1) The number of days spent at the release reefdecreased with body length at release (Fig 2; generallinear model, chi-square=1049.56, df=1, p<0.001),and was significantly affected by sex (general linearmodel, chi-square=58.47, df=1, p<0.001), site (generallinear model, chi-square=203.94, df=1, p<0.001), andthe interaction between sex and length (general linearmodel, chi-square=91.86, df=1, p<0.001) There was

no fishery monitoring component to this study, but arecreational angler incidentally caught one of thelingcod almost 5 years later at the release reef (Table1).The angler noticed the acoustic tag and notified theWashington Department of Fish and Wildlife

The seven resident lingcod were also used inanalyses that provided information on apparent shortexcursions from the reef that were shorter than 1 day

in duration The average resident lingcod made 0.5apparent short excursions per day, each lasting anaverage of 157 min The frequency of apparent shortexcursions was not significantly related to fish length

at release (generalized linear mixed model, F=2.07,df=1, 1152, p=0.15), but increased with the number

of days since release (generalized linear mixed model,F=189.22, df=1, 1152, p<0.001, Fig.3)

Apparent short excursion duration increased nificantly with fish length at release (linear mixedmodel, F=11.31, df=1, 5.17, p=0.019, Fig 4), andthe number of days since release (linear mixed model,F=33.78, df=1, 600.38, p<0.001, Fig.5)

sig-Discussion

Both hatchery-reared lingcod released in the currentstudy and wild lingcod released in previous studies

have exhibited substantial within-population

variabil-ity in range of movement In this study, half of the

hatchery-reared lingcod were at the release reefs every

day for the entire study The 2009 capture of one ofthe fish on the release reef shows that hatchery-rearedlingcod can stay at or return to release reefs several

Fig 3 Apparent short excursion frequency increased with the number of days since release Acoustic detection lags greater than one hour in duration were interpreted as instances where lingcod left the reef (apparent short excursions) These excursion analyses were only conducted for the seven lingcod that were detected at the release reef at least twice every day Each data point represents the month ’s excursion frequency averaged over all lingcod (±SE), but data analyses were conducted on the original daily data (not averaged over each month)

Fig 2 Larger lingcod tended to leave the release reefs sooner than

smaller lingcod Itsami Reef is represented by solid symbols and

lines Zee ’s Reef is represented by open symbols and dashed lines.

Triangles and circles represent females and males, respectively

Table 1 Fourteen lingcod were released on Zee ’s Reef and

Itsami Reef in Puget Sound, Washington State Table shows

release reef, lingcod length at release, sex, and the date each

lingcod left the reef for more than 1 day Five of the seven

lingcod that left for more than 1 day were never again detected

at the reef The seven lingcod that “did not leave” the reef were still present at the reef when the study ended on 22 January

2005 (Zee ’s Reef) or 23 January 2005 (Itsami Reef) Release reef Length (mm) Sex Fish ID# Release date Date left reef Most recent evidence fish was alive Zee’s 719 F 93 8/12/2004 8/12/2004 Periodically in Tacoma Narrows (8/17/04 to

10/12/004) Itsami 686 F 85 8/12/2004 10/10/2004 At Dana Passage (10/11/04); Back at Itsami

(12/5/04 to 12/18/04) Itsami 647 F 83 8/12/2004 11/1/2004 At Pt Richmond (11/4/04 to 11/10/04); At

Vashon Island (11/10/04) Itsami 602 F 82 8/12/2004 Did not leave Depth change of 10 m in 7 min on 1/22/05 Zee ’s 665 F 86 8/12/2004 Did not leave Depth change of 5 m in 4 min on 1/22/05 Zee’s 612 F 84 8/12/2004 Did not leave Depth change of 2 m in 21 min on 1/22/05 Zee ’s 676 F 81 8/12/2004 Did not leave Depth change of 21 m in 1 min on 1/21/05 Zee ’s 575 M 89 8/12/2004 8/12/2004 Not detected after leaving release reef Itsami 526 M 87 8/12/2004 8/19/2004 Back at Itsami (8/29/04 to 9/4/04); At Dana

Passage (9/4/04) Itsami 536 M 88 8/12/2004 8/20/2004 Not detected after leaving release reef Itsami 534 M 91 8/12/2004 11/25/2004 Not detected after leaving release reef Itsami 526 M 92 8/12/2004 Did not leave Caught by recreational angler at release

reef in 2009 Zee ’s 522 M 94 8/12/2004 Did not leave Depth change of 11 m in 2 min on 1/21/05 Zee ’s 550 M 90 8/12/2004 Did not leave Depth change of 10 m in 4 min on 1/21/05

years after release In contrast, the other seven lingcod

apparently left the release reefs between a few hours

after release and 4 months after release The

possibil-ity that predators removed these tagged fish from the

reef can be excluded for four of the seven lingcod,

which were detected at other locations (hence

moving) between 9 and 69 d after leaving the reef(average: 39 d)

Movement variation among individuals is consistentwith a recent study that tracked acoustically-tagged wildlingcod to describe space use within the home area(Tolimieri et al.2009) Space use varied as a function ofdiel and tidal cycles, but in different ways depending

on the particular individual (Tolimieri et al 2009).Movement variation among individuals has alsobeen observed in mark-recapture studies rangingfrom the Seymour Narrows to Neah Bay, where81% to 91% of recovered wild lingcod moved lessthan 8 km (5 miles) from the release site and 9% to19% moved a greater distance (Hart 1943; Jagielo

1990) In contrast, Mathews and LaRiviere (1987)found that half of the wild lingcod in northern PugetSound were recaptured less than 8 km from therelease site, while the other half were recapturedfrom a greater distance, leading them to propose thatmovement variation may also exist among popula-tions In a recent acoustic tracking study in PrinceWilliam Sound, 45% of wild lingcod dispersed fromthe release site (Bishop et al 2010)

Variation in lingcod movement among hatcheryindividuals (this study) and among wild individuals(previous studies) is paralleled in salmonids, wherethe tendency to stray varies among wild individualswithin a population, among wild populations, andamong hatcheries (Quinn 1993; Waples 1999) Sim-ilarly in wild tilefish, site fidelity varied amongindividuals within a season, but also between seasons(Yokota et al 2006) Controlling for release season,hatchery tilefish appeared to show greater site fidelitythan wild fish, though the authors suggest thathatchery rearing temperature may have caused thisdifference As we learn more about how hatcherypractices may affect variation in fidelity to releasesites, it may be possible and preferable for hatcheryrelease group behavior to match the variation in sitefidelity of receiving populations

This study and previous studies can improve ourunderstanding of some of the causes of variation inmovement Some of the among-individual variation inlingcod movement in this study was explained bybody size and sex Similarly, variation in wild lingcodmovement has been attributed to differences in bodysize (Bishop et al.2010), sex (Jagielo1990; Smith et

al 1990), and sexual maturity (Hart 1943; Jagielo

1990) Male and female lingcod off the Pacific coast

Fig 5 Apparent short excursion duration increased with the

number of days since release Acoustic detection lags greater

than one hour in duration were interpreted as instances where

lingcod left the reef (apparent short excursions) These

excursion analyses were only conducted for the seven lingcod

that were detected at the release reef at least twice every day.

Each data point represents the month ’s excursion duration

averaged over all lingcod (±SE), but data analyses were

conducted on the original daily data (not averaged over each

month)

Fig 4 Lingcod that were larger at release spent more time

away from reefs during apparent short excursions Acoustic

detection lags greater than one hour in duration were interpreted

as instances where lingcod left the reef (apparent short

excursions) These excursion analyses were only conducted

for the seven lingcod that were detected at the release reef at

least twice every day Each data point represents a single

lingcod ’s average excursion duration (±SE) over the entire

study, but data analyses were conducted on the original data

(not averages)

of Canada reach sexual maturity between 500 and

620 mm and 600–670 mm respectively (Richards et

al.1990) Some of the hatchery lingcod (males: 522–

575 mm; females: 602–719 mm) in the present study

were probably reaching sexual maturity, which may

have contributed to the variation in movement

behavior

Lingcod movement might also be influenced by

habitat characteristics of the release site (Ritter2008)

More lingcod remained at Zee’s Reef for the full

duration of the study than at Itsami Reef Zee’s Reef is

a naturally-formed reef composed of bedrock and

hardpan outcroppings, boulders, caves, and crevices

Itsami Reef is an artificial reef created in 1982 from

pieces concrete sidewalk, curb, and slabs piled

upon each other up to almost two meters high,

resulting in a matrix with crevices, overhangs, and

a complex geometry (Hueckel and Buckley 1987;

Mark LaRiviere, pers comm.) Larger sample sizes

and a different experimental design are necessary to

determine the influence of site characteristics on

movement variation

Stationary acoustic receivers in this and previous

studies have provided the ability to quantify the

duration of excursions from a home reef Starr et al

(2005) used stationary acoustic receivers to track 83

wild lingcod for up to 14 months in Alaska They

observed periods of site fidelity that were frequently

interrupted by excursions lasting a median of about

2 d (range=1−299 d) In contrast, mobile acoustic

receivers used to track acoustic-tagged wild lingcod

off Vancouver Island, British Columbia for 12 to 50 d

did not detect excursions from the release site

(Matthews 1992; Yamanaka and Richards 1993)

Stationary receivers used to track acoustic-tagged

wild lingcod in Prince William Sound detected few

excursions over the course of the study (Bishop et al

2010) Most of the lingcod in the present study also

made few long-term (> 1 d) excursions Starr et al

(2004,2005) hypothesized that lingcod make

excur-sions away from the home area to feed If so,

differences in the frequencies of long-term excursions

among studies may relate to differences in the local

availability of lingcod prey items

Excursion differences among studies may also reflect

natural increases in excursion frequency and duration

that may occur as lingcod age and grow This could

reflect increased food needs or decreased vulnerability

to predators (e.g., Dahlgren and Eggleston2000) This

ontogenetic increase in excursions is supported byStarr et al (2004, 2005), who tagged fairly largelingcod (female average: 99 cm, male average: 87 cm)and observed a greater number of long-term excursionsthan Matthews (1992), Yamanaka and Richards (1993)and the present study All three of the latter studiestagged smaller lingcod (female averages: 72 cm,

51 cm, and 66 cm, respectively; male averages:

64 cm, 49 cm, and 54 cm, respectively) A recentstudy that also detected few long-term excursionstagged lingcod that averaged 63 cm (Bishop et al

2010) Further, in the present study, apparent shortexcursion frequencies increased with days since release(age), and apparent short excursion durations increasedwith days since release (age) and with body size atrelease Thus, a positive relationship between excur-sions and body size or age is supported by trendsamong studies, comparisons of different-sized lingcod

in this study (effect of body size at release), andlongitudinal observations (effect of days since release)

of those individual lingcod in this study

In addition to effects of age and growth on shortand long excursions, age and growth may also affectdispersal (leaving without return) In this study, thenumber of days at the release reef significantlydecreased with lingcod length at release This rela-tionship was largely driven by the tendency for thesmallest individuals to remain at the release site forthe entire study (controlling for sex and site).Similarly in wild lingcod, size at tagging wasnegatively correlated with tendency to remain at therelease site (non-dispersal) (Bishop et al 2010).Previous studies have documented how hatcheryrearing can impede the expression of natural, adaptivebehavior (Olla et al 1998; Brown and Day 2002;Salvanes and Braithwaite2006) While this study wasnot designed to provide a direct comparison betweentelemetry-tagged wild and hatchery lingcod, thehatchery-reared lingcod displayed behavioral patternsthat have been reported for wild conspecifics Addi-tional important questions include how exact propor-tions of moving and staying individuals comparebetween wild and hatchery-origin fish when seasonand site are standardized, whether hatchery fish willstay at release sites for longer periods of time, showsimilarities or differences with wild fish in otherbehavioral traits, ultimately reproduce, or have nega-tive impacts on wild fishes; and how these parametersvary as a function of release age and study location

(Leber1995; Fairchild et al.2005) Future studies that

combine acoustic tagging and traditional tagging

methodologies with ecological, genetic, and health

monitoring of wild and hatchery individuals should

enable an evaluation of the costs and benefits of stock

enhancement (Liao et al.2003; Bell et al.2008)

Acknowledgements The authors thank Kelly Andrews for

providing acoustic telemetry data on lingcod #83, Megan

Moore for making Fig 1 , the Tacoma dive club, including

Mark LaRiviere and Dave DeGroot, for diving, and Karen

Grace-Martin for statistical advice Tom Flagg, Mark LaRiviere,

Nick Tolimieri, Culum Brown, and two anonymous reviewers

improved the manuscript with constructive comments JSFL

was supported by funds from the Science Consortium for

Ocean Replenishment (SCORE) during part of the data analysis

and manuscript preparation The views expressed herein are

those of the authors and do not necessarily reflect those of

funding agencies.

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Fluctuation and variation in stream-fish assemblages

after a catastrophic flood in the Miyagawa River, Japan

Yuichi Kano&Kaori Ohnishi&Yasuo Tomida&Naoyo Ikeda&Naomi Iwawaki&

Masahiko Miyagawa&Yasushi Harada&Hidetaka Ichiyanagi&

Katsutoshi Watanabe

Received: 11 July 2010 / Accepted: 18 May 2011 / Published online: 2 June 2011

# Springer Science+Business Media B.V 2011

Abstract In the autumn of 2004, a typhoon caused a

catastrophic flood of the Miyagawa River in Japan

Based upon snorkeling surveys conducted every

autumn from 2005 to 2009, we monitored the post

flood fluctuation of the local fish assemblages at nine

sites of both the main stream and subsidiary streams

of the river Results revealed that species richness

significantly increased from 2005 to 2009 In

addi-tion, the fish densities of eight species significantlyincreased over the same period, whereas the density

of one species decreased, and that of eight othersremained unchanged Categorization based on Eu-clidean distance revealed five main clusters from thenine sites Among these sites, fish assemblages withinsubsidiary streams were stable as they remainedwithin the same clusters while those in the mainstream were dynamically variable through time asthey changed cluster membership In addition, theEuclidean distance between two arbitrary fishassemblages was positively correlated with environ-mental distance (the Euclidean distance calculatedbased on river width, depth, velocity and pebble size),time distance, and spatial distance along the river Inconclusion, the fish assemblages were dynamicallyand regularly altered and varied in the five years afterthe flood, except for those in the subsidiary streams,and such variation was related to environmental,temporal and spatial variation

Keywords Cottus pollux Disturbance Liobagrusreini Niwaella delicata Pseudobagrus ichikawai Pseudobagrus nudiceps Typhoon Meari

Introduction

Flooding is a potentially important influence onstream-fish assemblages as represented by the floodpulse concept (Junk et al 1989) However, flood

DOI 10.1007/s10641-011-9861-8

Y Kano (*):H Ichiyanagi

Department of Urban and Environmental Engineering,

Graduate School of Engineering, Kyushu University,

744 Motooka, Nishi-ku,

Fukuoka 819 –0395, Japan

e-mail: kano@species.jp

K Ohnishi:N Ikeda:N Iwawaki:M Miyagawa

Ohsugidani Nature School,

199 Kuzu, Ohdai-cho,

Mie 519 –2633, Japan

Y Tomida

National Salmon Resources Center,

2-2 Nakanoshima, Toyohira-ku, Sapporo,

Department of Zoology, Division of Biological Science,

Graduate School of Science, Kyoto University,

Kitashirakawa-Oiwakecho, Sakyo-ku,

Kyoto 606 –8502, Japan

effects vary with flood severity (Resh et al 1988).

Thus, how catastrophic floods, i.e., those far outside

the range of usual seasonal water level fluctuations

and ordinary disturbances, influence stream fishes is a

topic of considerable interest in fish research For

example, Hoopes (1975) reported an extreme flood

event in the Appalachian Mountains that damaged

populations of the charr Salvelinus fontinalis, with a

particular impact upon small-sized individuals

Like-wise, fish fauna were drastically impacted over a short

period after another severe flood event in Arkansas

(Matthews 1986) Similarly, in Japan, the density of

the population of the sculpin Cottus pollux

consider-ably decreased due to a heavy flood event and

resulting environmental changes and refuge

limita-tions (Natsumeda 2003) On the other hand, some

surveys of flood events have reported few or no

notable impacts on fish species or populations (Tew et

al 2002; Lojkásek et al 2005; Pires et al 2008)

These reports, however, focused only on short-term

effects or on specific species

The Miyagawa River, flowing from the Odaigahara

Mountains to Ise Bay through Mie Prefecture, is one

of the most famous rivers in Japan for its clear water,

according to BOD (biochemical oxygen demand)

evaluation [MLIT (Ministry of Land, Infrastructure,

Transport and Tourism, Japan) 2010] The fish

biodiversity in the river is considerably high

com-pared with that in other river systems and includes

rare species such as the bagrid catfish Pseudobagrus

ichikawai, which has been designated as a national

natural monument species of Japan For example, 48

freshwater fish species were found in the Miyagawa

River, whreas 3–32 species were found in 14 other

river systems in Mie Prefecture (Miyamoto et al

2001) In the autumn of 2004, a heavy typhoon

(named “Typhoon Meari”) hit the region, with

maximum precipitation of 139 mm/h and 1000 mm

of continuous rainfall (Hayashi et al 2004) As a

result, debris flow and slope failure occurred at many

locations along the Miyagawa River, bringing disaster

to the region This represented the worst disaster for

the region for the last several decades, and it is said to

have been, in meteorological terms at least, an even

stronger event than the famous “Ise Bay Typhoon

Disaster” that killed more than 4500 people in 1959

(Hayashi et al.2004)

In this study, we monitored the fish assemblages at

nine sites in the main stream and subsidiary streams

of the Miyagawa River after the severe flood Giventhe fish assemblages endured a catastrophic flood wetested the hypothesis that the assemblage wouldchange through time along a recovery trajectory andthat such change would differ between the main andsubsidiary channels To assess long-term fluctuation

of overall fish assemblages, the monitoring wasimplemented for five years from 2005 to 2009 forall fish species present, with the exception ofartificially stocked species Analyses were conductedboth on each species and on integrated parameters offish assemblages, and here we report the results

Materials and methods

Study river

The Miyagawa River (Fig 1), covering a 920 km2watershed, is located on the Kii Peninsula of centralJapan The geological formation of the basin is steep,and human settlement has been restricted to onlyalong the main stream The climate is humidsubtropical with 2000–5000 mm precipitation peryear Precipitation is especially high in the typhoonseason (August to October; 200–600 mm per month).The mean monthly temperature ranges from 3.8°C(January) to 25.4°C (August)

-In the river, Anguilla japonica, Plecoglossusaltivelis altivelis, Oncorhynchus masou ishikawaeand Carassius spp have been artificially stockedevery year for game fishing [Miyagawa RiverFisheries Cooperative Association (MRFCA) pers.comm.], and the carp Cyprinus carpio (domesticated/colored type) has also been stocked by local residents

or has escaped into the river from domestic ponds(Table1), although native individuals of these speciesalso inhabit the river In addition, Opsariichthysuncirostris and Pseudobagrus nudiceps were uninten-tionally introduced from other regions of Japan, mostprobably from Lake Biwa (Tomita 1994), and theyhave became established in the river Of all the fishes,Pseudobagrus ichikawai is the most noteworthyspecies because it has been designated as a nationalnatural monument species of Japan and has beencategorized as “EN” (Endangered) in the MOE(Ministry of the Environment, Japan) Red List(MOE 2010) The population of P ichikawai in theMiyagawa River is regarded as particularly important,

as the river is one of the largest source areas for the

species (Watanabe1997)

Summary of the flood

Typhoon Meari brought a catastrophic flood to the

Miyagawa River from 29 to 30 September 2004

(Fig 2) The total amount of estimated precipitation

reached 600–1000 mm, with a maximum rate of

139 mm/h (Hayashi et al 2004), becoming the 11th

heaviest Japanese rainfall on record since such

weather observation records formally began in 1872

In addition, the precipitation data may have actually

been underestimated due to a certain failure in the

observation equipment Due to such heavy rain, the

Miyagawa River overtopped its banks, causing many

associated mudslides and debris flows in various

locations (Hayashi et al.2004)

Study sites and snorkeling survey

There were two large dams in the Miyagawa River,

the Miyagawa Dam and the Misedani Dam (Fig 1)

The Miyagawa Dam is a storage dam, and the

Misedani Dam is a power-generation dam without

any functions for water storage Thus, the Misedani

Dam overflowed violently during the flood Nine

study sites (A–I) were established on the main stream

and subsidiary streams of the river (Fig.1and Table2)

between the two dams Sites D–F and H were studysites previously established in 2000 (for P ichikawaiand P nudiceps in 2000), whereas sites A–C, G, and Iwere established in 2006–2009 Sites A, C, and Gwere situated in the subsidiary streams, and the streamclassification was “Aa” or “AaBb” (Kani 1944,

1984) Sites B, D, E and F were situated in themiddle reaches of the main stream and were classified

as “Bb” due to rippling flow Sites H and I, situatedbelow the Miyagawa Dam, were rather deep and slowand belonged to the habitat“BbBc” category

To obtain the relative density for each fish species, asnorkeling survey (as in Watanabe and Ito1999; Kano

et al.2010) was conducted for each site every autumnfrom 2005 to 2009 (Table 2) Surveys occurred atnight, between 20:00 and 23:00 The detection rates(detected individuals per all present individuals) by thesnorkeling survey were 21.5–28.7% (SE: 3.3–8.8%)and 34.7% (95% confidence limit: 27.0–43.1%) for P.ichikawai (Watanabe and Ito 1999) and O masouishikawae (Kano et al.2010), respectively, demonstrat-ing the validity of the snorkeling survey The surveywas conducted with three surveyors swimming slowlyupstream (100–200 m/h) with no overlapping or vacantareas The surveyors tallied the observed fish of eachspecies on a waterproof notebook No counts wereincluded for the five stocked species We consideredboth Rhinogobius flumineus and Rhinogobius sp CB

as Rhinogobius spp due to the difficulty of visual

Fig 1 Map of the

Miya-gawa River and study sites

(A–I) Detail is not shown

for the conservation of

Pseudobagrus ichikawai.

The shades of the squares

surrounding around the site

names correspond to those

in Figs 3 and 4

discrimination For Hemibarbus barbus/labeo,

Phox-inus oxycephalus jouyi, Pseudogobio esocPhox-inus,

Tribo-lodon hakonensis, Zacco platypus, and Zaccotemminckii, we stopped counting when the count

Fig 2 Fluvial landscape of the Miyagawa River as usual (left) (2010-01-20) and under the flooded condition (right) (2004-09-29, 13:22) [Photographs taken near site B (Fig 1 ) by T Takamatsu]

Family Species/

subspecies

Stocking (kg) (total in 2005 –2007) a Alien Red data list

ishikawae

1857 Domestic†Cyprinidae Carassius spp 50 Domestic†

Cyprinus carpio Unknown Domestic†Hemibarbus barbus

and/or H labeo Opsariichthys uncirostris Domestic Phoxinus oxycephalus jouyi

Pseudogobio esocinus Tribolodon hakonensis Zacco platypus Zacco temminckii Cobitidae Cobitis biwae

Siluridae Silurus asotus

Pseudobagrus nudiceps Domestic Gobiidae Gymnogobius urotaenia

Rhinogobius flumineus Rhinogobius sp CB Tridentiger brevispinis

Table 1 Fish list, stocking

amount and conservation

status (MOE 2010 ) of the

(pers comm.)†Native

indi-viduals were also inhabitants

reached 30 individuals because their abundance was

too great to count all individuals

Four environmental factors (river width, water

depth, water velocity, and substrate pebble size) were

measured in October 2008 for each site The river

width was measured each at 20-m intervals The water

depth, water velocity, and pebble size were measured

every three steps by walking zigzag in the river

(Bevenger and King1995) The values of each factor

at each site were averaged and then used as the

representative value of the site

Data analysis for increase or decrease in each species

To assess whether species richness and population

densities of respective fish increased or decreased

after the flood (from 2005 to 2009), analysis by the

generalized linear mixed model (GLMM) was

per-formed using the statistical software “R” (version

2.92) For the implementation of GLMM, the function

“glmmML” (glmmML package) was applied, in

which the year (2005, 2006, 2007, 2008, and 2009)

was the explanatory variable (as a metric variable)

and the species richness or density of each fish

species was the dependent variable The density of

each fish species was based on the number counted

during the snorkeling surveys, and the Poisson

distribution was fitted For the abundant species

(those for which we counted the first 30), a binomial

distribution was fitted, such that if the counted

number was over 30, the density was defined as“1”,

otherwise it was defined as “0” Sites A–F wereassigned as a random effect in the GLMM Tocalibrate the survey area, an “offset” argument in

“glmmML” function was used by substituting thelogarithm of the product of river width and surveylength (Table2) In the analysis for each species, siteswithout any count data for all years were omittedfrom the data on the assumption that the sites wereoriginally mismatched habitat for the species

Data analysis for fish assemblages

For the analysis, the density of each species (countnumber divided by survey area) was standardized Asfor species that were counted up to 30 individuals, thedensities were evaluated according to three levels: 0individuals = 0, 1–30 individuals = 1, and over 30individuals = 2, and the calculated values were thenstandardized in each species The Euclidean distancesbetween all possible pairs of the fish assemblageswere calculated, for which cluster analysis wasconducted using Ward’s method (Ward 1963) Theobtained dendrogram was divided into five groups.The same analysis was also conducted among thenine sites from the standpoint of environmentalfactors The four environmental factors were stan-dardized and clustered by the Euclidean distancesusing Ward’s method The dendrogram was dividedinto three environmental types

A model to explain the Euclidean distance oftwo arbitrary fish assemblages was made using

Table 2 Survey dates and environmental conditions of the study sites in the Miyagawa River

Site Stream classificationa Survey length

(m)

Distance from Misedani Dam (km)

Survey year and date

2000† 2005 2006 2007 2008 2009

a

According to Kani ( 1944 , 1984 )

† Survey was conducted only for Pseudobagrus ichikawai and Pseudobagrus nudiceps

Table 3 Total number of observed individuals by the snorkeling method and results of the GLMM analysis assuming the Poisson/ binomial distribution of fishes in the Miyagawa River

by GLMM [P value] (distribution)

2000 2005 2006 2007 2008 2009 Species richness A 4 3 3 4 0.08±0.04 [<0.05] (Poisson)

three parameters: the Euclidean environmental

distance, temporal distance, seasonal distance, and

spatial distance along the river between the two

assemblages The seasonal distance was the

differ-ence between the cumulative dates from January 1

to the survey date of particular year The spatial

distances between all possible pairs of the nine sites

were calculated by measuring the distance along the

river on the map Finally, a multiple regression

model was conducted in which the stepwise method

was used for the parameter selection (Pin-outwas set

at 0.05)

As for the most notable fish P ichikawai, Mori(2000) and Tokuhara (2005) suggested that the fishpreferred larger substrate stones as its habitatbecause large stones created void spaces whichthey could inhabit (no detail data were shown) Toexamine this assumption, the relationship betweenpebble size and density of P ichikawai wasanalyzed in this study A single regression model

was applied to average pebble size and P ichikawai

density

Results

By the snorkeling survey, we confirmed 18 natural

(and five stocked) species, as listed in Table 1 The

result of the snorkeling survey is shown in Table 3

The species richness in D–F and H was 8–10 (mean ±

SD: 6.75±0.96) in 2005 (just after the typhoon) and

increased to 12–13 (10.7±1.37) in 2009 The GLMM

analysis showed the significant trend in that the

species richness increased from 2005 to 2009,

showing a positive coefficient for year (Table 3)

Among species, the relative densities of Phoxinus

oxycephalus jouyi, Niwaella delicata, Liobagrus reini,

Pseudobagrus ichikawai, Pseudobagrus nudiceps,

Rhinogobius spp., and Cottus pollux were also

significantly increased from 2005 to 2009 The

analysis for Tribolodon hakonensis could not be

implemented because of statistical “perfect

separa-tion”, although this in itself implied that the fish had

increased On the other hand, Silurus asotus

signifi-cantly decreased after 2005 No significant increase or

decrease was detected in Hemibarbus sp./spp.,

Opsar-iichthys uncirostris, Cobitis biwae, Tridentiger

bre-vispinis, and Gymnogobius urotaenia

The environmental condition of each site and the

cluster analysis results are presented in Table 4 and

Fig 3, respectively The results corresponded to the

preliminary evaluation by Kani (1944,1984) (Table2)

In the dendrogram, sites H and I belonged to typeBbBc, with deeper water depth, slower velocity, andsmaller pebble size than other sites Sites B, D, E, and

F were categorized as type Bb with shallower depth,faster velocity, and larger pebble size than the typeAaBb, which is a typical characteristic of the middlereaches of the Miyagawa River Sites A, C, and G onthe subsidiary streams were type AaBb, showing thenarrowest width and fastest water velocity

The result of the cluster analysis for the fishassemblages is shown in Fig 4 The three type-AaBbsites (A, C, and G) belonged to the clusters I, II andIII, respectively, except for G in 2006, indicating solidstability among these fish assemblies On the otherhand, the four type-Bb sites (B, D, E, and F) changeddynamically; they first belonged to cluster IV during2005–2007, then to cluster III during 2007–2008, andfinally to cluster V during 2008–2009, except for D in

2009 As for the two type-BbBc sites (H and I), theyfirst belonged to cluster IV and then shifted to clusterIII and V, respectively

Results of the model predicting the Euclideandistance among the fish assemblages are shown inTable 5 The factors of Euclidean environmental,temporal, and spatial distances were selected by thestepwise method (multiple correlation coefficient: 0.37)

In short, the difference between the two arbitrary fishassemblages in the Miyagawa River were positivelycorrelated with environmental difference, temporaldifference, and spatial distance According to therelative standard error (SE/coefficient) and P-value,the environmental distance was the most effectivefactor, followed by temporal distance

The result of the single regression model betweenpebble size and Pseudobagrus ichikawai is shown in

Table 4 Fluvial environment of survey sites in the Miyagawa

River

Site Average value of environmental factors ±SD

Width (m) Depth (cm) Velocity

(cm/s)

Pebble size (cm)

1984 )

Fig 5 Data for A and C were omitted because no P.

ichikawai individuals were found in these sites

throughout the survey, and the habitats were likely

to be unsuitable for the fish owing to other reasons In

spite of the limited sample data, the analysis showed a

sufficiently low P-value and a high correlation

coefficient

Discussion

The fish assemblages of the Miyagawa River

exhibited a dynamic response and recovery following

the catastrophic flood of 2004 Almost half of all fish

species showed a significant trend toward increasing

post-flood density (Table 3) The available pre-flooddata for P ichikawai and P nudiceps were, unfortu-nately, not statistically comparable to the post-flooddata due to insufficient replications However, as thesurvey lengths were relatively long (250–390 m),comparison of sites may be reasonable and ofsignificance For example, in the year 2000, 121 and

19 P ichikawai were counted in sites F and H,respectively After the flood, this number had de-creased, with zero records for this species noted forboth these sites in 2005 It seems reasonable to suggestthat such a decrease is due to the impact of the flood

In D and E, the number of individual P ichikawai hadnot changed in contrast to changes in F and H As P.ichikawai prefers a habitat with large stones (Fig 5)

Fig 4 A cluster dendrogram of fish assemblages for each year ’s site data set and five distinct main clusters The shades of the squares surrounding the site names correspond to those in Fig 3

Factor Coefficient SE Relative standard error

Table 5 Results of multiple

regression model to explain

the Euclidean distance of

two arbitrary fish

assemb-lages in which the factors

were selected by the

step-wise method (Pin–out=0.05)

with void spaces (Mori2000; Tokuhara2005), sites D

and E with larger pebble size than sites F and H might

have been preferable for the fish

On the other hand in 2000, before the flood, P

nudiceps was unrecorded in any of the sites surveyed;

its numbers were recorded only in 2005 and later

Tomita (1994) had likewise reported that P nudiceps

had been previously distributed from the Misedani

Dam to the Town of Ryonai (about 1 km downstream

of site B) In 2000, this report was consistent with our

result, which showed no P nudiceps was found in any

sites However, the P nudiceps distribution extended

upstream to sites G and I until 2009 One possible

explanation for this is that the fluvial environment was

altered by the flood event to the benefit of this

particular species and its ecology, and this may have

aided in its migration An opposite case was reported

by Puckridge et al (2000); native fishes had a

advantage over exotic fishes during serial floods in

an Australian river As P nudiceps is a domestic alien

species, it is probable that the presence of this fish has

some negative ecological impact against the congener

P ichikawai This is an issue suitable for further study

It is noteworthy that the recovery of several fishes

(e.g., N delicata, T brevispinis and C pollux), which

abruptly increased in 2008 and 2009 (Table3), would

not have been detected if the survey had been

abandoned in 2007 Many surveys studying flood

effect on fish populations have assessed the situation

for only a short period after the flood (from

immediately after the flood to 19 months later, as in

Hoopes1975; Harrell 1978; Matthews 1986; Tew et

al.2002; Natsumeda2003; Lojkásek et al.2005;Čech

et al.2007; Pires et al.2008) The density of C pollux

in the Inabe River, Japan, had likewise been reported

to have been reduced by a heavy flood in 2003, but its

subsequent recovery was not reported (Natsumeda

2003) In our case we found that the recovery of C.pollux required 4–5 years (Table 3), although theperiod might be dependent on the magnitude of theflood and other localized conditions

Of all the species, Silurus asotus was the only fishthat significantly decreased from 2005 to 2009,suggesting that the fish decreased until 2009 MRFCA(pers comm.) suggested that a large number of S.asotus individuals drifted from the Miyagawa Damdue to the flood This suggestion might be true, as thedensity of S asotus in 2005 was exceptionally high insite H (Table 3), located just downstream of theMiyagawa Dam (Fig.1)

Over 30 individuals of Pseudogobio esocinus,Zacco platypus and Zacco temminckii were counted

in all locations and survey periods These fishes had thusbeen maintained at a moderate or high biomass levelsdespite the heavy flood On the other hand, Opsariich-thys uncirostris, Cobitis biwae, and Gymnogobiusurotaenia were maintained at low levels (0–4 individ-uals per site) in all locations and survey periods Wehave no information about whether these fishes wouldrecover further after 2009, or whether this low levelrepresents their natural density Of these, O uncirostris

is remarkable as a domestic alien species withpiscivorous traits This invasive fish has drasticallyexpanded in the Kyushu Region of Japan and has beencited as providing ecological disturbance for otherfishes (Onikura et al.2008) However, in the MiyagawaRiver, at least from our results, there is no evidence for

a tendency toward increasing density for this species,and little evidence for its invasiveness Given that theoriginal habitat of the fish is Lake Biwa (Onikura et al

2008), it might be difficult for the fish to establishstably in the flowing environment

These species-specific fluctuations brought anoverall fluctuation of fish assemblages, in which thephases of the fish assemblages shifted among theclusters as time advanced, except for type-AaBb sites,which stably belonged to the respective clusters(Fig.4) In addition, the difference in fish assemblagewas positively correlated with temporal distance,more significantly than with spatial and seasonaldistances (Table5) Although the pre-flood data werenot available, we can at least conclude that the fishassemblages largely changed not randomly, but with aconstant direction in 5 years after the catastrophicflood We suspect that such a steady dynamism might

Fig 5 The relationship between average pebble size and

Pseudobagrus ichikawai density at the sites (B, D, E, F, G,

and H) in 2008

be a process of ecological persistence and recovery of

the fish assemblages from a severe disturbance

brought on by the catastrophic flood In type-AaBb

sites in the subsidiary streams, on the other hand, the

fish assemblages were stable for 4 years after 2006

This may be because the fish assemblages in these

sites were not affected by the flood and/or recovered

before 2006 (no data exist for 2005) (Fig 4) In any

case, this also indicates the stability and persistence of

these fish assemblies, which were likely determined

by environmental conditions

In recent years, the frequency of heavy

precipita-tion has increased in Japan and some other regions of

the world [Endo et al 2005; Groisman et al 2005;

JMA (Japan Meteorological Agency) 2010a] JMA

(2010b) also presented a warning of the tendency

toward increasing frequency of 100-year heavy rain

events Thus, there may be an increasingly higher

probability that catastrophic floods, such as those

covered in this paper, will occur with greater

frequency in the future Furthermore, the importance

of disturbance, such as the “flood pulse concept”

(Junk et al 1989) and “intermediate disturbance

hypothesis” (Connell 1978), has been generally

discussed in ecology To understand the significance

of disturbance for freshwater fish in greater depth, a

program of preliminary surveys and long-timescale

monitoring for river/stream-resident fishes must be

planned, with the likelihood of future occurrences of

heavy floods in mind

Acknowledgements We are deeply grateful to C Wood for

English proofreading Our thanks also goes to R Fukui, M.

Kato, M Mori, K Muraoka, K Nakamura, Y Nishi, H.

Nishimura, Y Shimatani, T Suzuki and T Takamatsu for great

assistance We want also thank Miyagawa River Fisheries

Cooperative Association for the kind assistance and useful

information provided This work was supported in part by

Global COE Program (Center of excellence for Asian

conser-vation ecology as a basis of human-nature mutualism), MEXT,

Japan, and by the Global Environment Research Fund of the

Ministry of the Environment, Japan (Subject No D-1008)

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Sound production in Etheostoma oophylax (Percidae)

and call characteristics correlated to body size

Patricia Speares&Carol Johnston

Received: 2 September 2010 / Accepted: 18 May 2011 / Published online: 31 May 2011

# Springer Science+Business Media B.V 2011

Abstract Sound production in fishes is common in

marine and freshwater species, however there are still

many vocal species for which sound production has not

been documented This paper is the first account of

sound production in the Guardian Darter (Etheostoma

oophylax) Laboratory recordings revealed that males

produced several vocalizations, including single pulse

knocks, multi-pulsed purrs, and tonal drums All

vocalizations were documented during agonistic and

courtship encounters, including spawning We also

investigated possible correlations between call

charac-teristics and male size Male standard length was found

to be correlated to inter-pulse interval of purrs, as well

as the slope of the drum vocalizations Determining a

link between male size and acoustic characteristics

could be the first step in documenting mate or

male-male assessment by acoustic communication in darters

Keywords Mate choice Vocalizations

Standard length Darters

Introduction

Sound production is important for agonistic, courtship,

and spawning interactions in both freshwater and marine

species of fishes (for a review, see Ladich 2004;Amorim 2006; Myrberg and Lugli 2006) However,

to date there are many species for which characteristics

of vocal repertoires have not been formally described.Darters of the subgenus Catonotus (Genus: Etheos-toma) are a group of small benthic fishes foundthroughout the Southeastern United States Fishes inthis subgenus exhibit a breeding ecology known asegg-clustering (Page 1985) Males use flat rocks asshelters and spawning habitat which they defendagainst predators (Knouft and Page 2004) andconspecific males Males court females to lay clusters

of eggs on the surface of their shelter Courtship inCatonotus darters utilizes multiple signal modalities,including visual signals, such as changes in colorationand the development of egg mimics (Knapp andSargent 1989; Page and Bart1989; Page and Knouft

2000), as well as acoustic communication (Johnstonand Johnson2000) Assessment of physical character-istics of potential mates occurs in darters (Knapp andSargent 1989) However, to date, the hypothesis thatmate or conspecific male assessment occurs by means

of acoustic characteristics has not been tested Foracoustic communication to be feasible we would firsthypothesize that call characteristics should be indicative

of male condition or size

The objective of this study is to document soundproduction in Etheostoma oophylax, in both agonisticand courtship contexts, and determine which acousticcharacteristics of the vocal repertoire of this species, ifany, are correlated to male size Other studies haveshown that size (standard length, SL) may be

correlated to reproductive success (Magnhagen and

Kvarnemo 1989) and nest holding ability (Stammler

and Corkum2005) in fishes

Methods

Etheostoma oophylax were collected from three sites in

Calloway County, Kentucky, USA during the spring of

2007 and 2008 Collections were made in April of 2007

from Sugar Creek (36.65788 N, -88.15735 W) and

Panther Creek (36.58149 N,-88.10930 W) In April of

2008, collections were also made from Anderson Creek

(36.69446 N, -88.14353 W) Fish were caught by kick

seining into a 10 foot net Fish were placed into coolers

with creek water and an airstone Specimens were then

transported back to Auburn University by car

Laboratory accommodations were 76 L glass aquaria

with small cobble for substrate, and a filter (Millennium

2000 with activated carbon filtration, Marineland Inc.)

with an airstone Males were either housed alone or with

one other male Females were housed with up to 6 other

females per tank The water temperature was maintained

at 20-23°C and fish were kept under a 14:10 light/dark

photoperiod, but also had access to natural lights

through windows All recordings were taken within

2 weeks of collection from the field

Recording

Etheostoma oophylax readily produced vocalizations

and spawned in captivity, therefore all recordings

were done in the laboratory The recording tank was a

76 L glass aquarium with sand substrate and a single

tile shelter The tank was located in a sound

dampened room and placed on a foam pad on a table

separate from all other equipment During the

recording procedure all filters, air pumps, and

non-essential electronics were unplugged in order to

minimize unwanted noise Prior to recording, the

spectrogram and power spectrum were examined for

the presence of electrical noise If electrical noise was

found and could be identified then it was eliminated

by unplugging or shutting off electronics

Before recording a single male was placed into the

recording tank and given time to acclimate and set up

residency under the tile shelter overnight In the

morning, another male and multiple females were

added simultaneously to the tank for agonistic and

courtship trials Recording began as soon as the fishbegan interacting (usually < 5 min) Trials lasted

30 min or until the fish no longer interacted If fishdid not interact, the non-resident fish were removedand replaced by new fish The hydrophone wasplaced less than 1 cm from the shelter so that therewould be minimal distortion of the signal due to thetank environment (Akamatsu et al 2002)

All recordings were done with a Bruel and Kjaerhydrophone (Model number 8103: sensitivity−211 dB

re 1 V/μPa; frequency response 0.1 Hz to 180 kHz)connected to a Bruel and Kjaer charge amplifier (model2635) The amplifier was connected to a cassetterecorder (Sony: model TC-D5 Pro II) which allowedthe observer to acoustically monitor the trial Therecorder was connected to a PC computer (Dell OptiplexGX754), where sounds were collected using Raven PC(version: 1.2.1, Laboratory of Ornithology, CornellUniversity, Ithaca, NY) and stored for later analysis.During the recording trials, an observer was alwayspresent in the room and recorded the context of thesounds Sounds produced during male-male interactionswere classified as agonistic interactions All interactionsbetween males and gravid females were considered to

be courtship interactions

There is no distinct movement or behavior associatedwith sound production in darters Therefore, if morethan one male was present in the tank, the calls wereassumed to be made by the fish which was the aggressor

in the interaction The aggressor was determined to bethe individual which approached or chased the othermale, or the male which inhabited the shelter or nest.Often, we were able to easily determine which malemade the vocalization by identifying the male under thenest and therefore closest to the hydrophone Two maledarters rarely had overlapping vocalizations however, ifthis did occur then these signals were not used in theanalysis Females have never been shown to producevocalizations in any species of darter, therefore, allsound production in courtship interactions was attributed

to males

Signal analysis

Recordings were dissected into individual signals forfurther analysis Pulsed calls included knocks andpurrs, and tonal calls included drums, as previouslydocumented to occur in other species of Catonotusdarters (Johnston and Johnson 2000)

Purrs were identified as many (> 2) pulses in rapid

succession Total call duration and fundamental

frequencies, the number of pulses, the frequency of

each individual pulse, and the inter-pulse interval

were also measured (Fig 1) The fundamental

frequency of each individual pulse, and the inter-pulse

interval were averaged for each call Knocks were

identified as vocalizations which contained only one

pulse The fundamental frequency and the total call

duration were measured for all knocks (Fig.2)

Drum vocalizations contain multiple tonal bands,

which exhibit frequency modulation In order to

describe the variation in these calls, we looked at call

characteristics that have not previously been examined

These included the number of inflection points and the

first and second slope of the call (Fig.3) An inflection

point was defined as a point in the call when the

frequency modulation changed direction (i.e the call

changed from an‘up sweep’ to a ‘down sweep’) After

the number of inflection points was determined, the

slope of the call before and after each inflection point

was calculated For simplicity, only the first and second

slopes of each call were used in the analysis The slope

was determined by calculating the change in frequency(Hz), divided by the time (ms)

slope ¼ $y$x¼$ frequency$ timeFor each drum, the frequency of the second tonal band,the total number of tonal bands, the fundamentalfrequency, and the total call duration were alsocalculated (Fig.3)

All durations, and number of pulses were calculatedusing the waveforms in the time domain Slopes andinflection points of drums were calculated using thespectrogram (Hanning window, FFT size: 2466 sam-ples, bin resolution: 17.9 Hz) All frequency variableswere calculated using power spectra (Hanning window,FFT: 10000 samples, bin resolution: 4.4 Hz) All callswere averaged for each male prior to analysis Descrip-tive statistics (mean, standard deviation, range, mini-mum, maximum) were calculated for each male for allvariables To test for correlations between male size(SL) and call characteristics, a Pearson r correlation testwas performed for all of the above mentioned callcharacteristics A multivariate analysis of variance

Fig 1 Waveform,

spectro-gram and power spectrum

of a purr vocalization which

contains a total of 7 pulses

as determined by the

wave-form The total duration,

inter-pulse duration and

pulse duration was

deter-mined from the waveform

as indicated and the black

arrow on the power

spec-trum indicates fundamental

frequency of the total purr

(Hanning window; FFT=

10000; bin resolution=4.4).

The amplitude is shown in

‘kilounits’ (kU) which is an

uncalibrated arbitrary unit

used by the software Power

spectrum levels also

represent uncalibrated

values

Fig 2 Waveform,

spectro-gram and power spectrum

(Hanning window; FFT=

1000; bin resolution=4.4)

for a knock vocalization.

Black arrow on power

spectrum indicates the

fundamental frequency.

Total duration was calculated

from the waveform as

indicated The amplitude is

shown in ‘kilounits’ (kU)

which is an uncalibrated

arbitrary unit used by the

software Power spectrum

levels also represent

uncalibrated values

Fig 3 A representation of

the drum vocalization The

total duration of the call was

determined from the

waveform as indicated An

example of an inflection

point and multiple tonal bands

is indicated by the white

arrows The fundamental

frequency is also indicated in

the spectrogram The black

arrows on the power spectra

(Hanning window; FFT=

10000; bin resolution=4.4)

indicate the (from left to right)

the fundamental frequency

and the frequency of the

second tonal band The

amplitude is shown in

‘kilounits’ (kU) which is an

uncalibrated arbitrary unit

used by the software Power

spectrum levels also represent

uncalibrated values

(MANOVA) was used to determine if statistical

differ-ences existed between call characteristics in aggressive

and courtship contexts Standard length of the fish was

used as a covariate in this MANOVA since some call

characteristics were found to be significantly correlated

to male size All statistical calculations were performed

using SPSS statistics package (version 16.0, SPSS Inc

Chicago, IL)

Results

Call description

In total, 431 calls were analyzed from a total of 12 males

(Table1) which ranged in size from 590–750 mm (SL)

Purrs, drums and knocks were recorded during both

agonistic and courtship contexts Courtship calls

contained purrs (37%, n=117), knocks (34%, n=

109), and drums (28%, n = 91) Courtship purrs

contained an average of 6.05 pulses, with a mean total

duration of 0.377 s Individual pulses had a mean

duration of 0.042 s with a mean inter-pulse interval of

0.030 s The mean fundamental frequency of the entire

purr was 38.75 Hz, and the average fundamental

frequency of each individual pulse was 58.78 Hz.Courtship knocks had a mean duration of 0.062 s and afundamental frequency of 74.17 Hz Courtship drumshad a mean duration of 0.457 s The mean fundamentalfrequency of the drum vocalizations was 99.15 Hz, andthe mean frequency of the second band was 146.76 Hz.The courtship drums also had a mean of 1.51 inflectionpoints, 2.38 tonal bands, a slope of 1.127 Hz/ms for thefirst slope and−0.772 Hz/ms for the second slope.During agonistic interactions 21% of calls werepurrs (n=25) 39% were knocks (n=44) and 39% ofcalls were drums (n=45) Aggressive purrs had amean total duration of 0.312 s; each pulse had a meanduration of 0.023 s and a mean inter-pulse interval of0.03 s The mean fundamental frequency of the entireagonistic purr call was 96.84 Hz and the meanfundamental frequency of each individual pulse was101.12 Hz Aggressive knocks had an averageduration of 0.035 s and a mean fundamental frequency

of 103.66 Hz Aggressive drums had a mean totalduration of 0.392 s, a mean fundamental frequency of152.02 Hz and a mean second band at 170 Hz.Aggressive drums also had an average of 2.83 bands,1.67 inflection points, a mean first slope of 1.492 Hz/sand a mean second slope of−1.31 Hz/s

Courtship Agonistic

Purrs Purr duration (s) 0.377±0.244 0.312±0.171 24.013 0.001 Purr Fundamental (Hz) 38.75±20.807 96.84±26.34 24.014 0.001

Inter-pulse Interval (s) 0.030 ±0.021 0.030±0.021 0.143 0.870 Pulse Duration (s) 0.042±0.025 0.023±0.004 0.588 0.585 Pulse Fundamental (Hz) 58.78±16.604 101.12±16.674 0.875 0.464 Knocks

Knock Duration (s) 0.062±0.081 0.035±0.021 0.000 0.997 Knock Fundamental (Hz) 74.17±74.17 103.66±32.196 0.608 0.456 Drums

Drum Duration (s) 0.457±0.326 0.392±0.227 0.275 0.765 Drum Fundamental (Hz) 99.15±56.079 153.02±51.124 0.933 0.425 Second Tonal band (Hz) 146.76±43.826 170±33.194 1.039 0.389

# inflection points 1.51±1.634 1.67±1.87 0.897 0.438

Slope 2 -0.772±1.441 -1.31±6.897 0.631 0.552

Table 1 Mean ± standard

deviation of the call

characteristics for courtship

and agonistic interactions of

Etheostoma oophylax F and

p values indicate results of

MANOVA conducted with

standard length (SL) as a

covariate Significant

p values are indicated in

bold font