U.S. NATIONAL MUSEUM PROCEEDINGS, VOL. XXIII PL. XXI Sc Cu J 1st.A
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^nt.
Mid.
THE HERMIT CRABS OE THE PAGURUS BERNHARDUS
TYPE.
By James
E. Benedict,First Aifxistaid Curator,DirmonofMarineInvertebrates.
Iti.s thepurpose ofthis paperto bringto notice that section of the genus
Pagurus
of which P.henihardm
is the type.Pagurm
hernhnrdnH occurs in the shallow waters of northwestern Europe, and is representedby
P.acadianus in the waters ofnortheast- ern North America; hy P. alad:e/ms, P. aleuticm, and P. ochotemis in the waters of the northwest coast;by
P. patagoniensis and P.harljajci' inPatagonian waters.
Of
these species P. acadianuH and P. alaskemis aremore
nearly relatedthan any others.Next
is P. hernha/rdus,more
closely related to the formertwo
species than to P. aleuticus,which,inturn, ismuch more
closely related to the firstthree than to the spiny-handed forms, P. ochoteiisk^ P. patagoniemis. and P. Imrhiger.In the author's opinion the validityof thespecificdistinctions
made
can only becontradicted, if at all,after
more
collecting has })een donein intermediate localities,
when
it is possible that themore
closely relatedAmerican
formsmay
be reducedto subspecies.KKY TO THE SPECIES.
ff. Handsgranulated.
h. Widthofthelefthandatbase lessthanone-halfofitslength
bernhardufi(p.452)
.
//. Widthofthelefthandatl)aseone-halfofitslength.
c. Dactylesoftheambulatoryfeetnotgroovedonoutermargin.
d. Acicleslender, withoutserrateouteredge acadianus (p.454).
d. Aciclebroad,edgeserrate alaskensis (p.456).
'•. Dactylsoftheambulatoryfeetwithadeep groove onthe outermargin aleuticus(p.460).
a. Handsspiny.
h. Spinesofthe liaudswithout ))lack points ochoten.vx (p.463).
I>. Spinesofthehandswith black jjoints.
r. Antennulesniuclilongerthanthe eyes patagonienm(p.465).
c. Antennulesnotaslongasthe eyes barbif/cr(p.466) .
Proceedings U.S.National Museum,Vol.XXIII—No. 1216.
451
452 PROCEEDINGS OF THE NATIONAL MUSEUM.
vol.xxiii.PAGURUS BERNHARDUS
(Linnaeus.)Cancer heni]iardus Linn.eus, Syst. Nat., 1758, p. 631; Mus. Lud. Ulr., 1764,p.
454.
—
Herbst, Natur.derKrabbenundKrebse,II, 1796, p. 14, pi.xxii,fig. 6.
Astacus benihardus Pennant, Brit. Zool., IV, 1777, p. 23,pi. xvii.
—
De
Geer, Mem. pourservira 1'Hist, desInsects,VII,1778, p.405,pi.xxii,figs.3-12.—
Olivier,Enc. Meth. Insects,VIII, 1791,p. 641.
PagarmbernhardufiFabricius, Suppl. Ent. Syst., 1798, p.411.
—
Latreille, Hist.
des Crust.VI, 1805, p. 160; Gen. Crust, et Ins., I, 1807,p. 46; Consid.Genr.
surlesCrust,des Aradi.etdesinsect, 1810,p.422.^—Lamarck,Hist,desAnim.
sansVert.,V,1818,p.220.
—
Desmarest,Consid. surlesCrust., 1825,p.173,pi.
XXX,fig. 2.
—
Milne Edwards, Ann. des Sci. Nat., 2d ser., VI, 1836, p. 266;
Hist.Nat.desCrust., II,1837,p.215; Atlasdu RegneAnim., 3d ed.,Crust.,pi.
XLiv, fig.2; Ann. des Sci. Nat., 3d ser., X, 1848, p. 59.
—
Bell, Brit.Crust., 1853, p. 171.—White,Pop. Hist. Brit. Crust., 1857, p. 74.—Bate, Kept. Brit.
Assoc, 1865, p. 52.
—
Norman, Kept. Brit. Assoc, 1868, p. 264.
—
Benedict, Ann.
&
Mag. Nat. Hist. (6), XVIII, 1896, p. 99.Pagurus sfrebloiiyx Leach, Malac. Brit., 1815, pi. xxvi, figs. 1-4.
—
Latreille,
Encyc Meth.,1825, pi. cccix, figs. 3-6.
Pagurus uUdianu^W. Thompson, Rept. Brit. Assoc, 1843, p. 267.
—
Bell, Brit.
Crust., 1853, p. 180.—White, Pop.Hist.Brit.Crust., 1857, p.76.—Bate, Rept.
Brit.Assoc,1856,p. 52.
Eupagurusbernhardusvar.A. granulutaandvar.B.granalala-denticuluta Brandt, Middendorff'sSibir.Reise, Zool., 1851,p.107.
Benihardusstreblony.rDana, ProcAcad.Nat.Sci.,Phila.,VI,1852,p.6.
EupagurusuUdiami.sStimpson, Proc.Acad.Nat.Sci.,Phila.,X,1858,p.236.
Eupagurus bernhardus Stimpson, Proc.Acad. Nat. Sci., Phila.,X, 1858,p.236.
—
Heller,Crust. Sudl.Eur.1863,p.160.
—
Henderson,Proc.RoyalPhys.Soc, IX, 1886,1).68.—PococK,Ann.Nat.Hist.,6thser.,1889,p.427.
The
anterior portion of the carapace i.s usually aljout a.s broad as long, varying- in individualsto l)roader or longer.The
three projec- tions of the frontarepointed; the median isa little in advanceof the lateral,whichare eacharmed
with a single tubercular spine.The
eye scalesare suljoval.armed
justbelow the point with a spinewhich pro- jects and gives the scales a sharp appearance.The
eyestalks are stout, constricted in the middle, dilated atthe cornea.The
terminal joints of the antennula are a little in advance of the corresponding- parts of the antenna.The
acicle oftheantennaisthree-sided, slender, and smooth, except on the inner edge,where
it is rough and hairy.The
right cheliped is stout and strong; in thesmaller specimens it does not reach the tips of the ambulatory legs, while in older and larger specimens itoften exceedsthem
in length.The
upper surface of the carpus isarmed
with anumber
of short spines, the larger of which are in the inner marginal row.The hand
is about one-third longer than the carpus; the dactyl is usuallylonger than thepalm; its average length in 21 specimens was found to equal the breadth ofthe palm.The
left cheliped is nuich smaller than the right and variesmuch
inrelative length, sometimes reaching the base of the dactylof^P. licrnhardusisheremadethe typeofthe genus.
HERMIT
('11.1IIS—BENEDK
T.453
thelatter,sometimes notto the middleofthepalm.
The
widthof thepalm
averages alittle less than one-halfof itsleno-th.The
carpal and propodal joints of the ambulatoiy legs are spiny above; the dactyls are curved andtwisted.A
good description of the color can not ])emade
from the speci-mens
at hand. Stripes of red areshown
on the three distal joints of the aml>ulatory legs.The
upp(M- surfaceof
the hands, near the prehensile edges of the lingers and along the middle of the palm,is tinged with red.
The
measurements of the 2i specimens gi\en below will probablyshow
the average relative measurements of thePAGURUSBERN'HAROrS.
species.
The
carapace^ is measuredfrom
the point of the medianpro-jection of the front to the margin of the hardened portion; the hand
from
thetip of theimmovable
linger to the middle ofthe marginwhen
the
hand
is bentdownward from
the carpus; the dactylfrom
thetip tothe condyle.Many
of the specimensare worn. This will in partaccount for the
more
slender dactyls of the ambulatorylegs, as the thin edgesbecome worn
in themovements
of the animal overa rough bottom?The
fact that thelingers of the large chelipeds areworn
otf at the tip also reducestheir length in proportion to the width ofthe hand.454 PROCEEDINGS OF THE NATIONAL MUSEUM.
VOL. XXIII.RECORD OF
SPECl.AIKNSEXAMINED.
Norway; (1. O. i-^ars(YaleUniv. Mus.)• Shetland; A. M. Xorinan (YaleUniv.
Mus."). Firth ofClyde; John Murray(16988). Firth of Forth; A. M. Nor- man(1(5979). Channel Islands; Edward Lovett(6526). Jersey;
A.M.
Nor- man(6792). Holland (Yale Univ. Mus.). Europe(16980).Measurements ofPagurnsbernhardus.
HERMIT
('/,'.IBS—BEXEDTCT. 455
I
icaii I'onn the granules iiuike a .sharpand rather thinodj^e.
The worn
condition of the European specimens
may
aceount foi- a part of this diii'erenee.The
eyestalks in theAmerican form
arerehxtivel}^ laro-er, asshown
in tig. 1.The
measurements of acadianun weremade from
specimensaveraging-largerthan those with which they are com- pared.Much
of theutility of the studyof geographical distributionPagurttsacadiantjs.
of a genus
must
belost ifforms closely relatea aregroupea unaer onespecific name.
The name
hernhardiis as readilydesignates our speciesfrom
its associates,^?/56.s<?ew»s, hroyeri^ etc., as will acadianus, but the lattername
willmuch
better emphasize the difference separating itfrom
theEuropean
species orfrom
themore
closely related North Pacific form.RECORD OF SPECIMENS EXAMINED.
From the Grand Bank of Newfoundland to themouthof Chesapeake Bay, 7 to
I'tio fathoms, U.S.FishCommission.
Station.
456 PROCEEDIXGS OF THE XATIOXAL MUSECV.
vor..xxiTi.Off Cape Sable, NovaScotia, 16 fathoms(12604); (lloucester Harbor, Massachu-
setts, 7 to 10 fathoms (2564, 2610, 2S49); off Cape Ann, 19 to 29 fatlioms (25S0, 2597); off Cape Cod, 10 to 34 fathoms (XVSH, 4582, 458;',, 5034); Mncyurd Sound, 17 fathoms (3882, 3884,4542); off Gay Head (14397); Block Island Sound, 13 to ISifathoms (4543, 12853).
(IJoiieester<l(t)i((fl(nis.
Southwestern edge of Grand Bank, 200 fathoms(3721); off St. PetersBank, 265 fathoms (3762); Georges Bank, 32 to 46 fathoms(3723, 3758-3760); offPlymouth, Massachusetts (3534); Grand Manan,
New
Brunswick, S. F. Cheney (12332).MeasurementofPa<juri<.^<H-<t<li((iiiix.
ITKi:
MIT
rUAIlS—J]EXED
T('T.457
thr('(>-sidod; sidos flat, iridescent.
The
inner maro-iii isarmed
with from \'l to 1<> shoi't conical teeth, and isset with bunches ofhair.Tile
merus
of tlic right cheliped is very stout and strong-, and<>xtt'iids l)eyond the eye
by
about one-half itslength.The
carpus isaimed
on itsinner margin with a lino of stout spiues; there are alsoPAnURUS ALASKENSIS.
two
longitudinal rows of spineson its upper surface; it is elsewhere thickly set withspiny granules.The
hand is setwith spiny granules forming aborder on the outer margin.On
the lingers the surfaceisvery coarsely granular.
The
left cheliped is nnicli smaller than the right: its carpus has arow
of strong spiues on the iniuM- margin, anda parallelrow
farther458 PROCEEDINGS OF THE NATIONAL MUSEUM.
vol.xxiii.down
on tlu> outersurfjife; otherwise it is set with spiny oriiiudes us in the hirge hand.The
(hietyl does notshow from
alcove anv tiat-tened surfaee, but
from
the prehensile edge to the outermargin it isevenly rounded.
The
ambulatory legs of the rightside overreaeh the right oheliped but little; in very large specimens not at all.The
uppersurfaee of themerus
joints are a littleflattened.The upper
margin of the car- pus isarmed
witha singlerow
ofspines.Upper
surface of propodal joints flattened,armed
with arow
of short spines on the sunnnit and elsewhere with spin}' granules.The
dactyls are very wide, com- pressed, and twisted.The
upper surface is convex, itssummit
and margins each set with arow
of granules.Between
these rows are long%smooth
surfaces.The
inner surface of the dactjd isflat.The
outer surface is very convex near the proximal end, butbecomes much more
flattenednear the tip.In alcoholic specimens the general color a})ove is a light purple with iridescent reflections; below, light, tinged with reddish.
A
redstreak runs around theprehensile edgeof the
thumbs
and behind thedact3'^ls to the inner margins of the hands. Thereis an oblongpatch of I'edon the outer distalmargins and on the inner uppersurface of the
merus
jointsof the cheliped.The
lowerouter surface of the car- pal joints of the ambulatory legs are pointedwith red.The
propodal joints and dactyls are longitudinally streakedwith red.This species isvery close to hcrnhardus, but is easily distinguished
by
its broader and shorter left hand,by
the wide dactyls of the aml)ulat()ry legs, ])y the acicula, and by its pearh" iridescence.Brai)dt recognized but one
North
Pacific '*variet}'^ of hernhardus besides the verydistinctochotensls,' of this hehad buta single speci-men
obtained b}^Wosnesenski
at Unalaska and believedby
Brandt to be identical with the verycommon form
on the English coast,which he designates under the descriptivephrase as "var. B. granu- lata-denticuUvtay
As
alashenms has amuch
greater resemblance to the truehernhardus than has aleuticiis^ Brandt's descriptive phrase ismade synonymous
with it,though from
the localit}' aleuticuswould
benmch more
likely to be obtained. Stimpson says underEupagu-
rus hernJiardus,^
"Specimens
have been sentfrom Puget Sound by
Dr. Kennerly." Specimensfrom
the Straits ofFuca
are small but readily distinguishedfrom
acadianus.'Ann. Lye. Nat. Hist.,
New
York,VII, p. 89.III:i:MIT
crabs— BENEDr(
T.i;i;(oi;i)
ok
sf'kciimens IvXAminki*.459
Alaska, frmii liiiinak Island to l'>rist(jl Bay,5 to47I'atli
sion steamerA/hafross, 1890and 1891:
<; U. S. Fisli Cnimiii.-
Station.
4r>o
PROCEEDINGS OF THE NATIONAL MUSEUM.
VOI-.XXIII.Mea!<iirniiiiil.^ nfl^<i</iirii.tuliufki'ii
Locality.
i
J^o. i2ir>.
HERMIT CRA
l',S~BEXEDI<T.4()1 end. This character of the dactyls is sufficient to distinfuiyh this
from
any other species of the hfrnhardustype. Tlic coloi- is dark inmostspecimens.
The
dact}^ of the largehand
Juisa horny tipon the prehensileedijePagurusaleuticus.
inold and young. Inalashensis this character ispresentin the
young
only, as is also trueof acadiaiwx.
Owen's
remarks onthetwo
specimenstakenatKamchatka
willapply wellenough to this species.The
colorisusualh^ ''a dirtybrown
hue, and thelefthand
is quitevariable.''462 PROCEEDINGS OF THE NATIONAL MUSEUM
RECORD OF SPECIMENS EXAMINED.
From Unalaska to Oregon, 13 to 238 fathoms; U. S. Fish Commission steamer
Albatros.^.1888to 1890.
Station.
HERMIT CRABS— BEXKDK
T.468
464 PROCEEDINGS OF THE NATIONAL MUSEUM.
and alaskmMs,the inner surfacehighlyiridescent.
The
outerang-le of the basal article isverymuch
produced,and theinner edge is spiny.Chelipeds notlong, the nierus extending butyery little
beyond
the eyes.Where
thepreceding species of thehe/'n/iardustypearegranu-Pagurusochotensis.
lar or with spiny granules, this species is thickly set with slender spines.
The
hands are hairy, the hairs not reachingto theend of the spines.The
color of alcoholic specimensis a straw 3'ellow. Slenderstreaks of red run longitudinally on the carpal, propodal, anddactyl jointsof theambulatory legs.The merus
joints hayetwo
transversestreaksof thesame
color.RECORD OF SPECIMENS EXAMINED.
From VancouverIslandtoSanDiego,California,20to62 fathoms; U.S.FishCom- missionsteamerAlbatross, 1888and1890.
station.
NO.121(1. lIEliMIT
VRABS—BENEDHJT. 405
Alaska: W.H.Dall:
?Port Levasheff, 70 to80 fathoms, mud, stones (16778).
Sitka, 10 to25fathoms (14951).
Menzies Bay, Discovery Passage,BritishColmiibia,H fatlioms,soft hott ; Lieut.
CommanderH.E. Nichols, [T.S.N.(5929).
Victoria, BritishColumbia, 10fathoms; Dr. C. F. Newconihe (15801).
Straits ofFuca (3397).
PAGURUS PATAGONIENSIS
(Benedict).KiiIKKjui-m. patacjoniemi^Benedict,Proc. V.S.NationalMuseum,X\',1S92,jx 3.
Tlic iinterior portion of the carapaco is a little broader than long-.
The
front hawthreeroundedlobe«equally produced.The
laterallobesI
PaGURUSPATAGONIENSIS.
are
armed
with a .sinolc .small, sharpspine, which points directly for- ward.The
length of the eyestalks laid olf on the front equals the distancefrom
the angle of the front to the middle ofthe eyestalk on the opposite side; theyare stout,constricted in themiddle,anddilated andflattened at thecornea.The
eyescales are miu'h as in I\dld.sl'm-.s/.sy the subterminal spineis black tipped and shows
from
above; theacicles reach the base of the iiagella. In form they are very closely like that of hernhardus, and not so
much
flattened as in the related Alaskan species.Proc. N.
M.
vol. xxiii 30466 PROCEEDINGS OF THE XATIOXAL MUSEUM.
voi..xxni.The
chelipeds are a little shorter than the ambulatory legs of their respectivesides.The
carpus of the large cheliped is shapedlike that of P. hei'nhardus, but it is evenly set with short, sharp spines with black, horny tips.The
spines of the uppersurface of the hand are likethose of the carpus,andare arranged ins(n'enmore
orlessdistinct rows.The
hand ismore
than twice as longas broad; theoutermar- gin is arcuate; the inner margin is nearly straight.The
prehensile edges of thefingers arearmed
with large turbicles,andare hornynear the tips.The merus
joints of both chelipeds are very smooth on theirlarge surfaces, but bordered with spines below.The
leftcheliped reaches to the base ofthedactyl of the largehand.Itis similarin shape and armature, with the exception ofthe prehen-
sile edges of thefingers,whichare
much more
horny.The
carpus has an evenly convex outer and upper surfacearmed
with spines.The
dactyls of the ambulatory legs are curved and bent, as in hern- hardus:and alliedspecies, butare not compressed.Length
of the larger specimen 105 nun.; length of right cheliped 68mm.
; length of rightambulatory legs 85mm.
RECORD OF SPECIMENS EXAMINED.
Eastcoast ofPatagonia, 43fathoms,station 2768; U.S. FishCommission steamer
Alhatrotis, 1888 (16772), twospecimens.
PAGURUS BARBIGER
(A. Milne-Edwards).Bernliarduii harbigerA. Milne-Edwards,Crustacea, Mission Scientifiqiiedu Cap Horn, Paris, 1891,p.F'28,pi.iii,fig.1 a-e.
The
description of P.patag(mienf<i)<was
published before the U. S.National
Museum
received a copy ofthe above work, ora comparisonwould
have beenmade
andthe differencesin part given.As
the figures of P. harhiger weredrawn
])y A. Milne-Edwards and agree very well withhis descriptions, theymust,in the absence of the type specimens, beassumed
to be correct.The main
points of difference are as follows:The
front of P. harbigeris slightly three lobed, with the middle a little inadvance of thelateral lobes.The
length of the eyestalklaidofl'on the front equalsit in length.
The
carpus of the right hand isnearly rectangular.
The
outline of the left hand is evenly arcuate on both sides.The
figure of the secondfoot shows a broad dactyle, avery broad propodus arcuate beneath, the carpus is also broad, and both it and the propodus are
armed
with teeth or spines markedly differentfrom
those on the chelipeds. Notwithstanding these differ- ences, it is not impossible that the speciesmay
prove to be identical, as the typeandonlyspecimenof P. harhigerisverymuch
smallerthan either specimen in thismuseum. The
length of the rightcheliped isoiven as 23 nuu.