Amphipacifica : journal of systematic biology

152 Pages·1994·14.9 MB·English

AMPfflPACIFICA is an international jour- Editorial Board.
nal ofaquatic systematicbiologyaimedprimarily Dr. E. L. Bousfield, Managing Editor, Ottawa,
at publication of monographic studies that are Dr. D. G. Cook, Technical Editor, Greely, ON.
overlylarge(25-50-t-printedpages)fortraditional
taxonomic journals. The journal has featured Advisory Board.
studiesoncrustaceansofthefaunisticallyrichand D. R. Calder, Royal Ontario Museum,Toronto, ON,
palaeobiologically ancient North American Pa- Canada.
cific coastal marine region. However, the geo- F. R. Cook, NorthAugusta, ON, Canada.
graphicscopeextendstoothermarineregions,and C. G. Gruchy, North Gower, ON, Canada.
to adjacentfresh waters. Faunistically, it encom- P. H. LeBlond, GalianoI., BC, Canada.
Z. Kabata, Nanaimo, BC, Canada.
passes otherfree-living and parasitic invertebrate
G. G. E. Scudder, University ofBritish Columbia
taxa,andaspectsofaquaticvertebrateanimalsthat
Vancouver, BC, Canada.
may involve systematics, ecology and behaviour.
C.-t. Shih, Taiwan Fisheries Research Instiute,
AMPHIPACIFICA
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AMPHIPACIFICA
JOURNAL OF AQ.UATIC SYSTEMATIC BIOLOGY
Volume Number
3, 2
Nov. 15, 2001
AMPHIPACIFICA,
Journal ofAquatic Systematic Biology (ISSN No. 1189-9905) is published
quarterly by Amphipacifica Research Publications, 1710-1275 Richmond Rd.,Ottawa, Ontario.,
Canada, K2B 8E3, Annual subscription rates are $40. US or $50. Canadian funds.
DEDICATION
The Journal AMPfflPACIFICA is dedicated to the promotion ofsystematic biology and
to the conservation ofEarth’s natural resources.
This issue is dedicated to the memory ofPonald Evan McAllister, 1934 - 2001.
Coverdesign: Adapted from the title page ofS. J. Holmes (1904). “Amphipod
Crustaceans ofthe Expedition.”
Harriman Alaska Expedition, pages 233-244.
AMPHIPACIFICA VOL. 3 NO. 2. Nov. 15, 2001 1 ^
Donald Evan McAllister, 1934-2001:
A
Tribute
During his career at the National Museum (now
Canadian Museum of Nature) as Curator of Fishes
(1958-1986),SeniorResearchScientist(1986-1994),
Researcher Emeritus and Research Associate (1994 -
2001),DonaldEvanMcAllister madesignificantcon-
tributions not only in his formal ichthyological and
curatorial studies, butalsoin his thoughtful advocacy
of responsible environmental management, and for
protectingglobal biodiversity. Hefoundedandedited
twoscientificjournals,taughtandinspiredmanyindi-
viduals, and impressed all those he met as a warm,
friendly, humble, andpatient mandevoted tobothhis
familyandhis science. Butaboveall elseDonwould
havebeenmostpleasedbyacolleagues'scharacteriza-
tionofhimas atruefriendofthe Earth.
Donwasbom23August, 1934,inVictoria,B.C. He
tookhis B.A., M.A. (onfreshwatersculpinsofBritish
Columbia),andPh.D.attheUniversityofBritishCol-
umbia. His thesis for the latter, the Evolution of
Branchiostegals and Classification of Teleostome
Fishes, was published as a museum bulletin in 1968.
DuringhisPh.D. studies,hespentayearasanNSERC
Visiting Student at the Museum of Zoology, Univer- Donald Evan McAllister, 1934-2001
sityofMichigan. Hewas thefirstichthyologistonthe papers, books, popular articles, and book reviews,
staffoftheNational Museum since itscreationaspart ranging from his Ph.D. thesis and a Guide to the
oftheGeologicalSurveyin 1842,andwas,unquestion- FreshwaterSportfishofCanada, taxonomicrevisions,
ably, one ofthe most energetic, innovative, issue-ori- papers oncomputers inmuseums, methodsofcollect-
entedandproductive members ofthe staff. Although ing and preserving fishes, and the use of x-rays in
Donlackedadministrativeambition,orthepatiencefor taxonomic studies. He was alsothe originatoroftwo
it,herantheichthyologysectionwithverveandinfec- environmentaljournalstowhichhe contributedexten-
tiousenergy. He attracted, trained andencouraged a sive overviews and informed opinion.
successionofassistantcurators, technicians,contract- From the time of his appointment as curator, Don
ees, visiting scientists, post-doctoral students, assist- researchedArctic marinefishes with a view toacom-
ants and volunteers. His example of work ethic, panionbooktothefishesoftheotherCanadiancoasts.
compassion, and concernforthose aroundhim left its Heproducedapreliminarychecklistin 1960,followed
markon everyone whopassed though his section. by keys anda series ofpapers. A distributional atlas,
Donwas,firstandforemost,acollectionbuilder. On morepapersandabibliographyfollowed in the 1970s
hisarrival atthemuseum,therewerefewerthan4,5(X) and 1980s. Hecompiledanextensivedatabase which
specimens offishes in the collection. Through every hissuccessor,BrianCoadhascontinued. Withthehelp
means available to him, staff expeditions, donations, ofanumberofmuseum staffandthefinancial support
purchases and trades, he built the collection into a ofthe Department of Fisheries and Oceans, the long
world-class resource for taxonomic and life-history envisioned bookis close tocompletion.
studies. By the year 2000, there were more than Hewasapioneerandadvocateoftheuseofcomput-
585,000 specimens from all parts of the world, a ers in museums which led tohis devising a standard-
remarkable achievementon its own. But, at the same izeddatasheetforrecordingdetailedinformationinthe
time, Don was also a compulsive author who left a field. Healsocompiledanasyetunpublishedcompre-
comprehensive published record ofhis interests. His hensive list of ichthyological terms and their defini-
bibliography exceeds 620 items* including scientific tions. Thecomputerizeddatabaseofthefishcollection
AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 2
formedthebasisforalistofthefamiliesandspeciesof reefsandfindingnon-destructivemethodsforthehar-
fishesoftheworldwhichlistsmorethan 20,000valid vest ofaquariumfishes by Philippinefishermen. He
taxa. participatedin Canada'sBiodiversityConventionAd-
Don held adjunct professorships at Carleton and visoryGroupandadvisedonCanada’sstrategyonthe
Ottawa Universities, taught ichthyology at the latter, Riotreaty signedin 1991.
and participated on graduate student committees at Duringaperiodofmuseumdownsizing in 1993, in
both. In 1974, he organizedthe American Societyof a characteristic selfless gesture, he volunteered early
Ichthyologists and Herpetologists annual meeting in retirement to free a position so that a younger staff
Ottawa, onlythe thirdtime the Societymetin Canada membercould be retained. Following retirement he
andthefirstmeetingthatwasbilingual Donwasvery didnot "easeup" oneditingtheconservation-oriented
.
activeontheFishandMarineMammalssubcommittee journals he had initiated, nor did he slacken his own
oftheCommitteeontheStatusofEndangeredWildlife perceptive contributions tothem.
inCanadaandwrote thefirstlistofendangeredfishes EarlyinhiscareerDonhadmadepersonalcontribu-
in Canadain 1970. tionstostudiesofinvertebrateanimals,largelythrough
In the late 1980s and throughout the 1990s, Don his enthusiastic participation in two Museum expedi-
increasingly applied his wealth ofexperience and his tions in the 1960s with Drs. Edward L. Bousfield
social conscience to concerns for the planet. His (coastal southeastern Alaska) and Arthur H. Clarke
writingandactivitiesoninternationalcommitteeswere (eastcoastofHudsonBay), DuringtheAlaskaexped-
directed at a wider audience than his ichthyological ition,Don’skeeneyes spottedseveral unusual amphi-
colleagues, and he moved into the Museum’s newly- pods thatwereectoparasiticonrockfishandgreenling
created Biodiversity group. He had by this time cod that he had collected. Back at the museum lab,
foundedandeditedthequarterly"SeaWind",nowinits mostofthecrustacean species provednew toscience,
15thyearofpublication,andhemortgagedhishouseto oneofwhich,ParalafystiusmcallisterUwas namedin
fundconservationandtrainingprogramsorganizedby his honour. His passing has left a void that no single
itspublisher, "OceanVoiceInternational". In 1991 he individual is everlikely tofill.
originated and edited eight volumes of the quarterly
Canadian Biodiversity (later retitled "Global Biodi- FrancisR. Cook,CharlesG. Gruchy,and
versity")publishedbyCMN. In2000, itwasreplaced BrianW. Coad
A
with "Biodiversity: Journal ofLife on Earth" and *AcompletebibliographyofDonMcAllister'spublications
publishedbytheTropical Conservancy. He was esp- has been compiled by Brian Coad and will appear in the
ecially active in efforts to conserve the world’s coral CanadianField-Naturalist.
SEA WIND is published quarterlyand may beobtained through membership orsubscription from
OCEAN VOICE lOTERNATIONAL, P.O. Box 37026, 3332 McCarthy Road Ottawa, ON KIV OWO,
Canada, www.ovkca . PH; [phone], Fax [phone]. Regularmemberships: $25. peryear.
.
AMPHIPACIFICA VOL.3 NO. 2. Nov. 15, 2001 3
THE AMPHIPOD GENUSALLORCHESTES IN THE NORTHPACIFIC REGION:
SYSTEMATICS AND DISTRIBUTIONAL ECOLOGY.
By E. A. Hendrycks^ and E. L. BonsHeld ^
Abstract
ThetalitroideanamphipodfamilyHyalellidaeBulycheva1957,emendBousfield, 1996,andherewithrede-
fined,encompassesthemarinegeneraAllorchestesDana,ParhyalellaKunkel,ExhyalellaStebbing,MarinohyaleHa
Lazo-Wasem& Gable,andtheneotropicalfreshwatergenusHyalellaSaussure. FromthePacificcoastofNorth
Americaare newly described and figuredAllorchestes rickeri n. sp. (S.E. Alaska to Central California) andA.
priceaen, sp. (S.E,AlaskatosouthernVancouverIsland). Characterstatesofreproductivemorphology arenewly
utilized in redescribing and refiguring Allorchestes angusta, Dana (Alaska to California), A. bellabella Barnard
(KamchatkaandBeringseatoOregon),A.carimtaIwasa(NorthernSeaofJapan,BeringSeaandS.EAlaska), A.
malleolaStebbing(northern SeaofJapanandOkhotskSea),A.hirsutaIshimaru (southernSeaofJapan),andun-
describedspeciesofParhyalellaandHyalella. Numericalanalysisofselectedcharactersandcharacter states,partly
developed recently by Lazo-Wasem & Gable, supports taxonomic recognition of family Hyalellidae and its
distinctionfromthe closelyrelatedmarinefamilyHyalidae,andindicatesdegreesofmorphologicalsimilarityand
probableclosenessofrelationshipbetweenandamongitsfiverecognizedgenera. SpeciesofAllorchestesdominate
littoralmarinehabitatsofborealandcool-temperateopenandsemi-protectedsandbottomshavingsubmergedalgal
andphytalmats. SpeciesofHyalellaoccurintidal freshwaterportionsofestuariesandincoastalfreshwaterlakes
andponds ofwestern NorthAmerica, fromAlaskatoSouthernCalifornia.
INTRODUCTION
The talitroidean genusAllorchestes wasfirstpro- described by Stimpson (1857) from the Central Cali-
posedbyDana(1849), basedon species shortlyafter- forniacoast,hasnotbeenrecognizedsubsequentlyand
wards described as A. compressa Dana 1853, and A. maybesynonymouswithA.angustaDana. Stebbing
novizealandiae Dana, 1853,fromtheAustralian-New (1899,1906) described in detail A. malleolus from
Zealandregionofthe SouthPacific. Memberspecies seaweedin theYellow Seaand SeaofJapan. During
do share with member species of talitroidean family the twentieth century, three further western Pacific
Orchestiidae(nowTalitridae) someapomorphicchar- species weredescribed:A.vladimiri Derzhavin, 1937
acter states such as maxilla 1 palp minute or lacking, fromtheSeaofOkhotsk,A.carinataIwasa, 1939from
andtelsonentireorapicallycleft. However,thename OkhotskSeaandSeaofJapan,andA.hirsutaIshimaru,
''Allorchestes^^ (“other orchestes”) is somewhat of a 1995, from the Sea ofJapan. Tzvetkova (1990) ex-
misnomer since member species are mainly aquatic tendedtherangesofA.carinataandA.bellabellatothe
and apparently non-saltatory, not terrestrial or semi- Kamchatkapeninsularregion. Previous recordsfrom
terrestrial orcapable ofjumpingactively inthe air the Japanese region were summarized by Ishimaru
.
IntheNorthPacificregion, Allorchestesangusta (1994).
was described from the coast of California by Dana Along the North American Pacific coast. Stout
(1856), reaffirmed by Stimpson (1857), and subse- (1913) described, but did not illustrate, Allorchestes
quently found to be the common species ofthe entire oculatusfromtheLagunaBeachregionofS.Califomn-
region (below). In 1854-55, Stimpson described ia,a specieslatersynonymizedwith A.angustaDana.
Allorchestes rubricorniSy A. penicillatuSy and A.jap- Significantcontributionstothesystematicsofregional
onicus from coastal waters ofJapan. Despite the in- speciesofAllorchestesweremadebyBarnard,includ-
adequatedescriptionsandlackoffigures,thesespecies ing redescriptions of A. angusta (1952, 1954, 1970,
continued to be listed in Bate (1862) and even in the 1979), A. carinata (1979), and aformal new descrip-
recent catalogue ofIshimaru(1994). Character states tionofA.Z>e//a^e//a(1974, 1979,firstfiguredin 1954).
oftheeyesofA.japonicussuggestedaspeciesofHyale North Pacific species, including those of Hawaii
similar to H.pontica (see Della Valle 1893), but was (Barnard1970),werecompiledbyBarnard&Karaman
latertransferredtothesynonymyofA.compressaDana (1991), and those of North America by Bousfield
byBamard&Karaman(1991). Allorchestesseminuda (2001)
^ Research Division, Canadian MuseumofNature, Ottawa, Canada KIP6P4.
^1710-1275RichmondRd., Ottawa, ON, Canada K2B 8E3.
7
AMPHIPACIHCA VOL.3 NO. 2. Nov. 15, 2001 4
Inapreliminary reporton species offamilyHyal- MATERIALS AND METHODS.
idaeintheNorthAmericanPacific region, Bousfield Stationlistspertinenttofieldmaterialutilizedin
(1981) includedeight species ofA//orc/ie5re5, These this study are provided in Bousfield (1958, 1963,
andearlier recordsofA.angustaandA.bellab&ellaBar- 1968);Bousfield&McAllister(1962); andBousfield
nard 1974 were summarized by Ricketts Calvin &
Jarrett 1981),
(
(1968), Barnard (1975), Austin (1985) and Staude Thefollowingabbreviationsareusedinthetextand
(1987).
figures:
Basedmainlyontheresults of regionalfieldexpe-
ditions of the National Museum of Natural Sciences
(now the Canadian Museum of Nature), the present ABlR-2 - abnrtoeondnlaam1,el2la
studyextendsknowledgeofthesystematicsanddistri- CLSP - clothespin spine
butional ecology ofthe genus Allorchestes and other CX
- coxal plate
members offamilyHyalellidaeonthe Pacificcoastof DACT
- dactyl
NorthAmerica.
EP - abdominal side plate
ACKNOWLEDGMENTS GNl-2 - gnathopod 1, 2
HD
- head
Forassistancewithf^eldworktheauthorsaregreatly
LFT - left
indebted to regional marine laboratories, notably the
LL - lowerlip(labium)
PacificBiological Station,Nanaimo,B.C.,thePacific MD
- mandible
Environmental Institute, West Vancouver, B. C., the MX
BamfieldMarineStation,B.C.,andtheFridayHarbor MXP1-D2 - maxilla 1. 2
Laboratories, WA. Much of the detail has been ac- - maxilliped
knowledged in previous station lists (below). Dr. PP3L-7 -- ppleeroaepoopdods3,4, 5, 6,
Craig P. Staude, Dr. Colin B. Levings, the late Dr. PLP - palp
JosephineF.L.Hart,andthelateDr.DanielB.Quayle RET
- retinacula
were especially helpful with thefield collecting. RT
- right
The authors are grateful to Drs. Peter Slattery and SP - spine
Charles E. O'Clairforproviding valuable studymate- T ' - telson
rialfromtheBeringSeaandAleutianIslandsregions. U
- uropod
Dr. Nina Tzvetkova, Zoological Museum, St. Peters- UL
- upperlip(labrum)
burg, provided helpful commentary and loan ofvalu- UROS - urosome
able westernNorthPacificmaterial. Wemuch value X
- enlarged
the careful research of Dr. Shin-ishi Ishimaru on
im - immature
AllorchesteshirsutainJapanesecoastalwaters,andfor
juv - juvenile
hiskindpermissiontoreproducehispublishedfigures
ov - ovigerous
ofthat species(1995)asdeemedessential tothecohe- subad. - subadult
sionofthis study.
Taxonomic work on CMN amphipod collections
wasperformedinitiallybyELBasastaffmemberofthe SYSTEMATICS
National Museum of Natural Sciences at the Holly
Lane Laboratory, Ottawa, during the period 1979- Family Hyalellidae Bulycheva
1984. Originallineillustrationswerepreparedwiththe
capable assistance ofartist Roy E. Zittin, Cupertino, Hyalellidae Bulycheva, 1957: 18. Bousfield 1982:
California. Marjorie Bousfield provided translations 270; 1996: 176.
ofthe Russian literature. Talitridae(part) Stebbing 1906: 523. Gurjanova 1951.
The work was carred out mainly in the research Hyalidae(part)(subfamily)Barnard 1970: 268. Bous-
laboratoriesoftheCanadianMuseumof Nature. The field 1981: 176.
authors are especially grateful toassistantcollections Talitroidea (part) Barnard & Barnard 1983: 161.
manager Judith C. Price for help in retrieving, cata-
CMN
loguing, and labeling of amphipod material. TypegenusHyalella Smith, 1874.
AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 5
Genera: Allorchestes Dana, 1849: 136; Exhyalella plateslarge,subrectangularorovateanddistallyacute;
Stebbing, 1917, revised Lazo-Wasem & Gable 2001: marginal hooked setae shorttomedium in length.
51; MarinohyalellaLazo-Wasem & Gable, 2001: 67;
Parhyalella Kunkel, 1910:74. Laso-Wasem&Gable Distributional Ecology: Within the North Pacific
(2001) have recommended suppression of the enig- marine region, species of the genus Allorchestes are
matic genus Insula Kunkel, 1910, and we herewith largelyendemictocool-temperatewatersofbotheast-
concur. ern and western coasts. The tropical and warm-
temperategenusParhyalellaisrepresentedinTaiwan
Diagnosis:Bodysmooth,occasionallycarinateorpro- byP.kunkeliLazo-Wasem&Gableandinthesouthern
cessiferous,mainlyabdominally. Antennaerelatively Japan Seabyanundescribedspecies; intheHawaiian
short. Antenna 2 longer than antenna 1, peduncle Islands by P. pietschmanni Schellenberg; and on the
(male) occasionally stout and proximal flagellar seg- PacificcoastofNorthAmerica(BajaCalifornia)byP.
ments conjoint (incrassate). barnardi Lazo-Wasem & Gable. Species ofHyalella
Mouthparts of generalized talitroideanform. Man- occur in coastal fresh waters ofNorth America north
dible, leftlacinia5-6dentate; bladesofspinerowfew approximatelytothe tree line inAlaska.
(2-4); molarsetatending to reductionorloss. Max-
illa 1, palp minute or lacking. Maxilla 2, inner plate Remarks: Notall systematists(e.g.,Barnard&Kara-
with 1-2 prominent inner marginal setae. Maxilliped, man 1991; Lazo-Wasem & Gable 2001) accept the
outerplate short; palpmoderate, dactyl unguiform. original concept of family Hyalellidae and generic
Coxal plates 1-4 deep, subrectangular, posterior inclusionsthatwere splitofffromfamilyHyalidaeby
marginal cusp(acclivityofBarnard 1979)vestigial or Bulycheva(1957). Barnard (1969) subsequentlycre-
indistinct. Gnathopods usually strongly sexually di- ated Chiltoniinae, within the Ceinidae, to encompass
morphic. Gnathopod 1 (male),oftenmodifiedforpre- the antipodean freshwater genera Chiltonia, Austro-
amplexus; dactyl and propodal spines modified for chiltonia,andAfrochiltonia. Theaberranthyalidgenus
fittingintoprecopulatorynotchofperaeon2offemale. HyacheliaBdmdxdi1967:120),tentativelyplacedwithin
Gnathopod 2 (male) powerfully subchelate; propodal the talitroideanfamilyNajnidae by Bousfield (1982),
palm smoothlyconvexorweaklytoothed;carpal lobe wasalsoplacedwithinCeinidaebyBarnard&Karaman
welldeveloped,interiormarginlinedwithcombsetae. (1991). However,itstruefamilystatusremainsenig-
Peraeopods3-4slender,distalsegmentsposteriorly matic.
armed with singly inserted complex spines; dactyls SubfamilyChiltoniinaeBarnard 1969:467(part),is
simple. Peraeopods 5-7, bases broad, hind margin herewith removed from the Hyalellidae. Its genera,
lacking ‘‘surge seta” and corresponding indentation; along with marine genera Allorchestes and Parhyal-
distal segments typically slenderandweakly spinose; ella, had been placed within family Hyalellidae by
dactyls small tomedium, lackingmediansetaorlock- Bousfield (1982, 1996). Chiltoniin generalack: (1) a
ingspine;basisofperaeopod7,posterodistallobelying carpal lobe in male gnathopod 2; (2) a precopulatory
mediad ofischium. notch in peraeon 2 (female); and (3) a corresponding
Epimeral side plates ordinary, plate 2 deepest. modificationofthe propodanddactyl ofgnathopod 1
Pleopodsnormal,raminotsexuallydimorphic;pedun- (male), all suggesting separate phyletic origin.
cle with 2-3 small retinacula; innerrsimus proximally The Chiltoniinaecontinuedtoberecognizedwithin
with claviform "clothespin" spines. Uropods 1 - 2 family Ceinidae by Barnard & Barnard (1983) and
short,pedunculardistolateraland/ordistomedialspines Barnard & Karaman (1991). However, its principal
ordinary; rami variously (or not) marginally spinose, subfamily character states, involving sexually dimor-
apical spinesoccasionallystriate. Uropod3uniram- phicandothercharacters,donotconformwiththoseof
ous;ramusdistinct,short,notfusedtopeduncle. Telson the type genus Ceina. Subfamily Chiltoniinae may
plate-like;lobesfusedbasally,separatedbydistalnotch therefore require separate family recognition within
only, orfused totally. the Talitroidea, a proposal beyond the scope of the
Coxalgillsmedium,sac-like,subequal. Sternalgills presentstudy. SubfamilyHyalellinaeisthereforehere
(when present) paired, sublinear. consideredmonotypic withinfamilyHyalellidae.
Female: Peraeon segment 2 with anterodistal pream- ClassificatorystatusoffamUyHyalellidae
plexing (precopulatory) notch (Figs IB, 6). Brood Barnard & Karaman (1991) have retained some
AMPHIPACIHCA VOL. 3 NO. 2. Nov. 15, 2001 6
generaoffamilyHyalellidaewithin
familyHyalidae (sens,lat.l. appar-
ently without formal numerical
&
analysis. Laso-Wasem Gable
(2001) also include the genera
Parhyalella, Exhyalella, and
Marinohyalella within Hyalidae
(sens,lat.). However,allgeneraof
family Hyalellidae Bulycheva are
reliablydistinguishedfromallmem-
bers offamilyHyalidae (sens, str.)
bythefollowing character states:
1. Maxilla 1,palpvestigialorlack-
ing (vs. distinct, 1-2 segmented);
2. Telson plate-like, apex entire or
narrowly cleft (vs. fully bilobate,
cleftto base).
AdditionallywithinfamilyHyal-
ellidae: coxae 1-4 are not poster-
iorly cuspate (vs. often distinctly
cuspate); gnathopod 2 (male), car-
pal lobe is always presentand dis-
tinct (vs. lackingorvestigial);per-
aeopods 3-7, distal segments are
slender, weakly spinose (vs. stout,
strongly spinose); peraeopods 5-7,
baseslackposteriormarginal“surge
seta”andpit(vs.present);uropod 1,
distomedial and distolateral ped-
uncular spines ordinary (vs. often
strongly developed);and uropod3
uniramous, or ramus fused to the
peduncle (vs. occ. unequally bi-
ramous, outer ramus not fused to
peduncle). Sternal gillsarepresent
in one supergenus (Hyalella) (vs.
lacking in all Hyalidae including
freshwater members). Addition-
ally, the preamplexing notch (fe-
male)ispresentinailgeneraandspeciesofHyalellidae names are herewith regarded as nomena nuda and
and is highly variable in form (Figs. 1, 6). The suitablyreplaced inthis study.
preamplex-ing notch is yet known only within a few
subgroupsofHyalidaeand, whenpresent,isrelatively
simple inform. CharactersandCharacter States.
AlthoughacceptedbyTzvetkova(1990), Barnard Inprevious studiesofhyalellidandhyalidamphi-
&
Karaman(1991)didnotrecognize thenew generic pods,charactersandstatesmostfrequentlyselectedfor
and species names proposed by Bousfield (19^1) for species distinction have been those of the antennae,
North American Pacific species within families mouthparts, gnathopods, uropods, and telson (e.g.,
Hyalidae and Hyalellidae (e.g., within Allorchestes). Barnard 1979;Ishimaru 1995). In this studyofNorth
Bousfield (2001) tentativelyrecognizedthem inalist Pacifichyalellids,thosecharactersarealsoutilized,but
of amphipod species of North America, but these the taxonomic significance ofreproductive morphol-