First records of snailfishes (Pisces: Liparidae) from the Galapagos Islands, with descriptions of two new species, Paraliparis darwini and Paraliparis galapagosensis

10 Pages·2002·4.6 MB·English

by  David L Stein

;
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume53,No. 11,pp. 151-160, 2 figs. August22, 2002
First records of snailfishes (Pisces: Liparidae)
from the Galapagos Islands, with descriptions oftwo new species,
Paraliparis darwini and Paraliparis galapagosensis
by
David L. Stein
NOAA/OARR/0SS3
1315East WestHighway, Rm. 11805
SilverSpring, Maryland20910
email: [email protected]
and
NataliaV. Chernova
ZoologicalInstitute
RussianAcademyofSciences
199034St. Petersburg, Russia
email: antarctdOzin.ru
Snailfishes (Family Liparidae, Order Scorpaeniformes) areverywidelydistributed both
in depth and geographically; morethan 130speciesare known from theSouthern Hemi-
sphere,andfivespeciesfromdeep,easternPacific,equatorialwaters.Wereporthereinthe
first record of the family from the Galapagos Islands, and describe two new species,
ParaliparisdarwiniandP.galapagosensis.
Liparidfishes aredistributedworldwide in cold andcool waters atdepths fromthe intertidalto
over7000 m.Theyarehighlyspecioseandmorphologicallydiverse,occupyingawidevarietyofhab-
itats includingbenthic, epibenthic, andpelagic zones. Inthe Southern Hemisphere, thereare at least
130 described species (Andriashev 1986; Stein and Tompkins 1989; Stein and Andriashev 1990;
Stein,Melendez,andKong 1991;AndriashevandStein 1998;Stein,Chernova,andAndriashev2001
ChernovaandSteininpress;andothers)fromtheAntarctic,Australia,Chile,Argentina,andotherlo-
cations. In the tropical and subtropical Pacific, the five known species are from great depths
{CareproctuslongifillsGarman 1892, SWofPanama,3343 m;ParaliparisfimbriatusGannan 1892,
SW ofPanama, 3241 m; P. latifrons Garman 1899, Panama Bay, 3279 m; P. angustifrons Garman
1899,PanamaBay,935 m;P. attenuatusGarman 1899,PanamaBay, 1650 m).Nospecieswerepre-
viously known fromthe Galapagos Islands (Grove and Lavenberg 1997).
In this paper, we report the first records of the family from the Galapagos, and describe
ParaliparisdarwiniandParaliparisgalapagosensis,twonewspeciesfromupperslopedepths. Both
specimens were collected by the Johnson Sea-Link, a three-person research submersible especially
well suited forcollecting specimens in topographically complex environments where trawl nets are
unusable.
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Methods
WefollowpreviouslypublishedmethodsofAndriashevandStein(1998)andSteinetal. (2001)
forcounts, measurements, and abbreviations fordescribing liparids. Proportions are given as % SL
(%HLinparentheses). TerminologyofmouthandlowerjawfollowSteinetal. (2001). Countswere
obtainedusingradiographs.PectoralgirdleswereremovedandthenclearedandstainedwithAlizarin
Red S bythe method ofAndriashev, Neyelov, and Prirodina (1977).
The following abbreviations forcounts and measurements have been used:
Forcounts
V: vertebrae (abdominal + caudal)
D: dorsal-fmrays
A: anal-fmrays
C: caudal-fmrays
P: pectoral-fm rays (upper lobe rays +notchrays + lower loberays)
gr: gill rakers
pc: pyloric caeca
Rad: radials ofpectoral girdle
Forsensoiypores:
io: infraorbital
n: nasal
pm: preoperculo-mandibular
temporal
t:
Formeasurements:
aAf: distance from centerofanus to anal-fm origin
bd: maximumbody depth
bdA: body depth at anal fin-origin
E: horizontal diameterofeye
go: length ofgill opening
HD: maximum headdepth
HL: head length
HW: headwidth
io: interorbital width (between uppermargins ofeyes)
Ij: lowerjaw length
LPL: greatest length oflower lobe ofpectoral fin
ma: length from mandibular symphysis to centerofanus
NL: length ofshortestnotchray ofpectoral fin
po: postocularhead length
preA: preanal-fm length
preD: predorsal-fin length
sn: snout length; tip ofsnout to anteriormargin ofeye
SL: standard length
uj: upperjaw length
UPL: greatest length ofupper lobe ofpectoral fin
Collection abbreviations follow Leviton etal. (1985). Figures arebythejuniorauthor. Pectoral
girdle preparations are depositedwith the specimens.
steinandchernova: galapagos liparids 153
Taxonomic Descriptions
GenusParaliparis Collett, 1878
Paraliparis Collett, 1878:34 (type species Paraliparis bathyhii Collett, 1878 by monotypy).
Burke, 1930:154; Andriashev, 1954:464; Cohen, 1968:385; Stein, 1978a:5, 1978b:37; Andriashev,
1986:14; Stein, C—hemova, andAndriashev, 2001:360.
Diagnosis. Onepairofnostrils(nostrilssingle).Ventralsuckingdiskabsent.Asingletermi-
nal (sensu Andriashev, 1986)orsuprabranchial (sensu Burke, 1930)porepresent intemporal canal.
Pectoral fin divided into two lobes or not; if present, lower lobe not forming a single filament.
Pseudobranch absent. Coronal pore absent. Barbels orskin flaps on head absent.
Paraliparis darwinin. sp.
Figure
1
—
Diagnosis. AParalipariswithteeth inupperjawabsent, in lowerjaw uniserial exceptvery
closetosymphysis.Teethinlowerjawbluntcanines,ovalincross-section.Chingelatinous,chin-pore
paironanteriorratherthanventral surface. Headlength21.8%SL,itswidth65%,anddepth84%, its
length. Upperjaw44%) HL. Body depth at anal-fin origin 22.1 (102). Lowerpectoral-fin lobe 65%)
length ofupper l—obe. P 23 (15+2+6), V 66, C 8. Rad4 (3+1), round.
HOLOTYPE. CAS 86576. Female, 131 mm SL, 149 mmTL. "Johnson Sea-Link" Dive 3971,
NE Pacific, Isla Wolf, Galapagos Islands, Ecuador, 637 m, suction collector, 23 November 1995.
Coll. J. E. McCoskeret al. —
CountsANDMeasurements. V66(12+54),D58,A53, P23 (15+2+6),C 8(4/4),gr 12,pc
10.HL21.8,HW 14.1 (65),HD 18.3(84.2),sn8.4(38.6),E3.8(17.5),po 1 1.5(52.6),io8.5(39.3),go
3.8 (17.5), uj 9.5 (43.9), Ij 8.5 (39.3), preD 33.5. preA40, ma 15.6, aAf22.9, bd 26 (1 19), bdA 22.1
(102), UPL 15.3 (70—), NL 5.3 or 35%o UPL, LPL 10.7 or 70% UPL.
Description. Headdeepatocciput, notcompressed, itswidth 1.3 in itsdepth. Anterodorsal
contourofheadslopingatangleofabout45°,slightlyconcaveaboveeye. Snoutlarge,deep,rounded,
gelatinous, slightly projecting beyond upperjaw, its length 2.2 times eye diameter. Subrostral fold
deep, well developed, coveringupper lip endrely. Nostril with raisedrim, not large, horizontal with
lowermarginofeye. Eyelongitudinallyoval,moderatelylarge,nottouchingdorsalcontourofhead.
Interorbital2.2 timeseye. Mouth subtenninalandsmall,oral cleftreachingtobelowanteriormargin
ofpupil, posterior end ofmaxilla (underthick soft skin cover) below posteriorthird quarterofeye.
Lowerjawsubtenninal.Chingelatinous,anteriormarginprominentbeyondlowerjaw,chinporeson
its anterior rather than ventral surface. Teeth on upperjaw absent. Teeth on lowerjaw primarily
uniserial;in3shortobliquerows(about3,4and6teeth)atsymphysis,butposteriorlyformingasingle
longrowofno fewerthan25 teeth onright sideofjaw. Teethsmall canines, not sharp, nottouching
each other; in the uniserial row posterior teeth slightly larger than anterior; in dorsal view (e.g.,
cross-section) teeth oval. Sensorypores less than nostril in diameter; pores on snout with thickened
margins(contoured),incomparativelydeeppitsofsofttissueandthuslooksimilartonostrilindiame-
ter. Pores 2-6-7-1,postorbital porepresent. Chinporesnotcloselyset, interspaceaboutequal todis-
tancepmi-pm2,eachinashallowpitofsofttissue.Gillopeningsmall,equaltohorizontaldiameterof
eye, entirelyabovepectoral-finbase. Gill opening locatedrelativelyhigh on side ofbody, itsdorsal
end horizontal with upper margin ofeye. Opercular flap small, triangular, its rounded tip closer to
lowerend ofgill opening and level with uppermargin ofeye.
Pectoral finnotched,notchrays2. Uppermostpectoralrayhorizontalwithuppermarginofeye.
Upperlobenotreachinganal-finorigin. Dorsalmost(6th)lowerloberaylongest,lowermostpectoral
rayinsertedbelowanteriorthirdofpostocularspace.Coracoidnotprojectingfromventralcontourof
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STEIN ANDCHERNOVA: GALAPAGOS LIPARIDS 155
body. Lowerpectoral-finlobelength65%thatofupperlobe. Rightand leftlowerpectoral-fin lobes
nottouching. Atransverse,thick,pocket-shapedskinfoldpresentonisthmusanteriortolowerlobes.
Radials 3+1, round or rounded, unnotched, R3 distinctly smallerthan others. Scapula with slender
helve ofmoderate length, coracoid with very slender, long helve, base notchedventrally.
Bodyhump-backed,deep(depthatanal-finoriginaboutequaltoheadlength). Anusbelowthird
quarterofpostocularspace. Anusandsmallgenitalpapillainashallowpit,surroundedbysoftradial
skin folds. Dorsal contour ofbody widely rounded. Maximum depth at dorsal-fin origin. Two free
intemeuralspresentbetweenneural spines4, 5,and6. Firstdorsalraybetweenneuralspines6and7.
Dorsalandanal finsnotdeep: atmid-lengthofcaudalpartofbody, findepthlessthanbodydepth. In
posteriorthirdofcaudalpartofbody,dorsalfinslightlydeeperthananalfin.Dorsalandanalfinsover-
lapabouthalfofcaudal fin. Gelatinoustissuemoderatelydeveloped; thickeronsnoutandchin. Skin
thin, semitransparent, covered with thick mucous layer, opaque after fixation. Free neuromasts not
mm
found. Pyloriccaecadigitate,tipssharp, longestca. 5.3%SL. Ovarianeggsunripe, lessthan0.5
in diameter.
Inalcohol,bodycolorpale,yellowishbrown.Marginsofdorsalandanalfins,caudalandpectoral
fins blackish. Ventrally, snout and head, inner subrostral fold, lips and gill opening rosy brown.
Mouth andgill cavities duskygray. Gill archespale. Peritoneum ink-black. Stomach, pyloriccaeca,
and intestine pale. —
Distribution. Known only from the holotype, collected offIsla Wolf, Galapagos Islands.
Thespecimenwascollectedfromagentlysloping, sandbottomusingrotenoneandasuctioncollec-
tor. Also collected during the same dive were Chauliodus sp., Stylephorus chordatus, Symphurus
diabolicus, and som—e invertebrates.
Etymology. Named after Charles Darwin, describer of the biological diversity of the
Galapagos Island fau—naand its significance.
Comparisons. The new species is most similar in counts to P. galapagosensis (below) but
differsmostsignificantly innumberofgillrakers(12 vs. 8). Itsgeneralaspectisquitedifferent,hav-
ingamuchlessroundedandlongerheadandlongerabdominalcavity. Specifically,itdifferspropor-
tionallyfromP.galapagosensisinheadshape(anterodorsalcontourat45°,snoutprojectingvs.head
evenlyrounded,snoutnotprojecting),headdepth(84.2vs. 100%HL),snoutlength(39vs.34%HL),
upperjawlength(43.9vs.36.8%HL),andbodydepthatanus(102vs.75%oHL).ItisalsosimilartoP.
merodontiis Stein, Melendezand Kong 1991 fromChile,butdiffersmostdistinctlyinthenumberof
vertebrae(66vs.62-64),radials(3+1 vs.3+0),andpyloriccaeca(1 vs.6-8).Proportionallyitdiffers
mostinlongerhead(21.8vs. 15.1-17.2)andsnout(8.4vs.4.4-5.7),greaterpredorsal-finlength(33.5
vs. 19.3-26.9), preanal-fin length (40 vs. 34.5-37.6), and body depth (26 vs. 17.3-20.2).
Paraliparisgalapagosensis n. sp.
Figure 2
—
Diagnosis. AParalipariswithupperjawdentitionlimitedto3^isolatedteethoneachside
ofsymphysis;lowerjawteethuniserialexceptveryclosetosymphysis,recurvedanteriorly,tipsover-
lappingtoformasharpcuttingedge.Chinnotgelatinous,chinporepaironventralsurfaceofchin.HL
20.7% SL, its depth equal to its length. Upperjaw 37% HL. Bodydepthatanal-finorigin 15.4 (75).
Lowerpectoral-fi—nlobeequaltolengthofupperlobe.P22(15+1+6),V67,C8.Rad4(3+1),round.
Holotype. CAS 86737. Female, 94 mm SL, 104 mm TL. "Johnson Sea-Link" Dive 3949,
Cabo Rosa, S ofIsla Isabela, Galapagos Islands, Ecuador, 710 m, suction collector, 11 November
1995. Coll. J. E. McCoskeret al. Specimen compressed, head strongly flexedventrally.
CountsandMeasurements. V67(12+55),D60,A55,P22(15+1+6),C8(4/4),gr8,pc9.
HL20.7%SL,HW-(-), HD20.7(100),sn6.7(33.7), E4.1 (20.0),po 1 1.1 (58),io6.7(32.6),go3.3
(15.8), uj 7.6 (36.8), Ij 8.7 (42.1), preD 30.6 (136.1), preA 33.0 (147.2), ma 13.6 (60.4), aAf 16.3
1
156 PROCEEDINGS OFTHECALIFORNIA ACADEMYOF SCIENCES
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STEIN ANDCHERNOVA: GALAPAGOS LIPARIDS 157
(72.6),bdca.27(108),bdA 15.4(74.7),UPL 13.0(63.2),NL-(-),LPL 13.0(63.2), LPL 100%UPL,
pc 6.5 (31.2). —
Description. Head deep at occiput, anterodorsal contour evenly curved from occiput to
above eye; from there, sloping almost vertically to tip ofsnout; in life, probably quite compressed.
Snoutshallow, rounded,notgelatinous,onlyslightlyprojectingbeyondupperjaw, short, 1.7eyedi-
ameter. Subrostralfolddeveloped.Nostrilwithraisedrim,notlarge,aboutonahorizontalwithlower
marginofeye.Eyeround,notlarge,farbelowdorsalcontourofhead. Interorbital 1.6eye.Mouthinfe-
rior and small, oral cleft reaching to below anterior margin ofeye, posterior end ofmaxilla (under
thicksoftskincover)belowposteriorVaofeye. Lowerjawincluded. Chinnotgelatinous,roundedin
lateralview,slopinggraduallyposteriorly,chinporesonitsventralsurface. Teethonupperjawpres-
ent, but only 3^ isolated teeth on each side ofthe symphysis. Teeth on lowerjaw uniserial except
closetosymphysis, fonningonelongrowofnofewerthan40teethonrightsideofjaw. Teethsmall,
sharp,recurvedanteriorlysothattipsoverlaptoformasharpcuttingedge.Teethgraduatedinsize,be-
comingincreasinglylargeposteriorly;ovalincross-section. Sensoryporessmallerthannostrilindi-
ameter; pores on snout without thickened rims (not contoured), not in pits ofsoft tissue; similar to
nostrilindiameter. Pores2-61-1-1, postorbitalporenotfoundowingtoskindamage,probablypres-
ent. Chinporesnotcloselyset(interspaceaboutequaltodistancepmi-pm^), levelwithchinsurface,
not in pits. Posteriorto postorbital pore, arow ofabout 14 freeneuromasts presentalongmidline of
body.Gillopeningshort,0.8eyediameter,entirelyabovepectoral-finbase.Gillopeninglocatedrela-
tivelyhighonsideofbody,levelofventralendwellaboveuppermarginofeye.Opercularflapsmall,
triangular, its tip above middle ofgill opening, well above horizontal through uppermargin ofeye.
Pectoralfinnotched,onenotchray.Uppermostpectoralrayabovehorizontalthroughuppermar-
ginofeye. Upperlobereachinganal-finorigin. Longestlowerloberayfifthfrombottom; lowermost
pectoral ray insertedbelowposteriorthirdofpostocularspace. Lowerpectoral-fin lobe length about
equal to that ofupper. The two lowerpectoral-fin lobes not touching. Radials 3+1, R3 smallest; all
round, unnotched. Scapula with slender short helve, coracoid with long, very slenderhelve.
Bodystronglyhump-backed,deepestjustposteriortogillopening.Anteriorofvertebralcolumn
stronglycurvedventrallytoparallel dorsal outlineofbody, probablynatural. Anusbelowgill open-
ing, almost between insertion of lowest pectoral-fin lobe rays. Ventral contour of body almost
straight.Onefreedorsalintemeuralpresent,insertionoffirstdorsalraybetweenvertebrae6/7.Dorsal
andanal finsnotdeep: atmid-lengthofcaudalpartofbody, findepthslessthanbodydepth. Inposte-
riorthirdofcaudalpartofbody, dorsal finnotdeeperthananal fin. Dorsal andanal finsoverlap first
third ofcaudal fin. Gelatinous tissue absent. Skin thin, semitransparent. Pyloric caeca digitate, tips
sharply pointed, longest ca. 6.5% SL.
Inalcohol,bodycolorpale,yellowishbrown.Marginsofdorsalandanalfins,caudalandpectoral
fins blackish. Ventrally, snout and head, lips and gill opening rosy brown. Mouth and gill cavities
dusky gray. Gill arch—es pale. Peritoneum inkblack. Stomach, pyloric caeca and intestine pale.
Distribution. Knownonlyfromtheholotype,collectedoffIslaIsabela, GalapagosIslands.
The specimenwas capturedusinga suction samplerabove a sandbottom withno current. Alsocol-
lected during the dive were Symphwus diabolicus, Bathypterois sp., Dibmnchus erinaceus, and
Rajella eisenhardti,—andnumerous shrimps and holothurians.
Etymology. ThenewspeciesisnamedaftertheGalapagosIslandsofEcuador,thelocation
ofits collection. —
Comparisons. In its counts and tooth arrangement, the new species is most similar to P.
dai-w'ini(see above forcomparison) although its appearance isquitedifferent. It isalso similartoP.
merodontus from Chile, but differs in number ofvertebrae (67 vs. 62-64), anal-fin rays (55 vs.
50-52),radials(3+1 vs. 3+0), andpyloriccaeca(9vs. 6-8). Proportionallyitdiffersmostnotablyin
itsgreaterbodydepth(ca. 27% SLvs. 17.3-20.2% SL),but italsodiffers significantlyin longerHL
1.
158 PROCEEDINGS OFTHECALIFORNIAACADEMY OF SCIENCES
Volume 53,No. 1
(20.7vs. 15.1-17.2)greaterpredorsal-finlength(30.6vs. 19.3-26.9),andshorterdistancefromanus
to anal-fin origin (16.3 vs. 19.4-24.4).
Discussion
Given the recent discoveries of many new snailfishes from the Southern Hemisphere (e.g.,
Andriashev and Stein 1998; Stein et al. 2001), the discovery of new liparid species from the
GalapagosIslandsextendstheverybroadbathymetricandgeographicdistributionofthefamily.The
islandsareisolatedbothaboveandbelowthewaterlevel(McCoskerandRosenblatt 1984;Groveand
Lavenberg 1997)andthustheoccurrenceofspeciesendemictothe islands, ratherthanoccurringon
themainland,is"normal."Estimatedichthyofaunalendemismis9.4%ofthe437speciesreportedby
Groveand Lavenberg(1997), thevastmajorityofwhichoccuratlessthan 100 mdepth,butonly23
species are known from greater depths (ibid.); we suggest that this probably reflects the lack of
deep-water sampling around the islands ratherthan a depauperate fauna. McCoskerand Rosenblatt
(1984) explained the occurrence ofendemic species by pointing out that most ofthe endemics are
fromfamilieswith lowvagilityoflarvae, i.e. short larval stageswithout longpelagicphases. This is
alsotrueofdeep-waterliparids(Stein 1980).Theypredictedthat"thedeep,nearshoresteepslopes. .
are likelyto contain interesting surprises modem submersibles allow mankind to reach new
. . . . . .
depths. . ."ThesenewspeciesofafamilypreviouslyunknownfromtheGalapagosfitthisprediction
in location and collection method.
Thezoogeographic/systematicrelationshipbetweentheGalapagosspeciesandtheliparidfauna
ofthe SouthPacific isunknown, althoughthegeneral relationshipoftheislands' shallow-waterfish
fauna is closest to the eastern tropical Pacific mainland, with nearly 60% of species in common
(McCoskerand Rosenblatt 1984; Grove and Lavenberg 1997).
With respect to their tooth arrangement and general morphology, the new species seem most
closelyrelatedtoP. merodontusfromChile,whichhasanedentateupperjawanduniseriallowerjaw
teeth,butdiffersinhaving3+0radialsratherthan3+1. Tootharrangementinbothnewspeciesseems
toplacethem intheParalipariscopeigroup(seeAndriashev 1986, Stein andAndriashev 1990, and
Steinetal. 1991),whichisnominallybasedonallmembershavingteethuniserial inatleastonejaw,
veryshortorporelikegillopeningwithreducedopercularflap,and3(3+0)pectoralradials.However,
intheirpectoral structureand somewhat largergill openingbothdifferfromP. copeiand its subspe-
cies {P. copeigibbeiicepsAndriashev 1982a,P. copeikergiielensisAndriashev 1982b, andP. copei
w//50«/Richards 1966).
Therehasbeennophylogeneticanalysisoftherelationshipsofthisgroup{P. copeicopeiGoode
and Bean 1896, P. copei gibbericeps, P. copei kergiielensis, P. copei wilsoni, P. merodontus, P.
nassariim SteinandFitch 1984,P. i/ee/ov?Andriashev \9^2h,P.paucidens Stein 1978b, P. rosaceus
Gilbert 1890)orthoroughexaminationoftheircomparativemorphology,andindeed,theiridentityas
a group is somewhat questionable. Pectoral girdle structure is unknown for P. nassariim. P.
paucidens,andP.rosaceus,andasdescribedabove,P.darwiniandP.galapagosensisdifferatleastin
theirpectoralgirdlemorphology.Thesedifferencessuggestthateithertoothpatternorpectoralgirdle
structure is not a good indicatorofrelationship. Resolution ofthis question awaits a full analysis.
Acknowledgments
The seniorauthorwas partially supported bythe Office ofOceanic and Atmospheric Research,
National OceanicandAtmospheric Administration. Thejuniorauthorwassupportedby the Russian
ScienceFoundation,GrantsNo99-04-49774,96-15-97881 andFGP"WorldOcean"Project 16. We
thankJ. McCoskerforbringingthespecimenstoourattentionandforprovidinghisdetailedfieldcol-
lection notes.
STEINANDCHERNOVA: GALAPAGOS LIPARIDS 159
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