Review of western Atlantic species of cocculinid and pseudococculinid limpets, with descriptions of new species (Gastropoda: Cocculiniformia)

33 Pages·1995·28.4 MB·English

by  James H Mclean

Review of Western Atlantic Species of
COCCULINID AND PSEUDOCOCCULINID LlMPETS,
New
with Descriptions of Species
(Gastropoda: Cocculiniformia)
James H. McLean1 and M. G. Harasewych2
CONTENTS
ABSTRACT 3
INTRODUCTION 3
MATERIALS AND METHODS 3
KEY TO WESTERN ATLANTIC COCCULINIDAE AND PSEUDOCOCCULINIDAE 4
SYSTEMATICS 6
Family Cocculinidae 6
Genus Cocculina Dali, 1882 8
Cocculina rathbuni Dali, 1882 8
Cocculina messingi, new species 11
Cocculina emsoni, new species 13
Genus Coccopigya Marshall, 1986 13
Coccopigya spinigera (Jeffreys, 1883) 13
Coccopigya mikkelsenae, new species 14
Genus Coccocrater Haszprunar, 1987 16
Coccocraterpocillum (Dali, 1890), new combination 16
Coccocraterportoricensis (Dali & Simpson, 1901), new combination 17
Genus Fedikovella Moskalev, 1976 17
Fedikovella caymanensis Moskalev, 1976 17
Fedikovella beanii (Dali, 1882) 19
Family Pseudococculinidae 22
Genus Notocrater Finlay, 1926 24
Notocraterhoubricki, new species 24
Notocrater youngi new species 26
,
Genus Tentaoculus Moskalev, 1976 27
Tentaoculus eritmeta (Verrill, 1884), new combination 27
Tentaoculusgeorgiana (Dali, 1927), new combination 28
Genus Caymanabyssia Moskalev, 1986 28
Caymanabyssia spina Moskalev, 1976 28
Genus Kaiparapelta Marshall, 1986 28
Kaiparapelta askewi, new species 28
Genus Copulabyssia Haszprunar, 1988 29
Copulabyssia leptalea (Verrill, 1884) 29
Genus Amphiplica Haszprunar, 1988 29
Amphiplica venezuelensis McLean, 1988 29
REALLOCATED TAXA 30
“Cocculina” conica Verrill, 1884 30
“Cocculina” dalli Verrill, 1884 30
“Cocculina” reticulata Verrill, 1885 31
1. Natural History Museum ofLosAngeles County, 900 Exposition Boulevard, Los Angeles, California 90007.
2. DepartmentofInvertebrateZoology,NationalMuseumofNaturalHistory,SmithsonianInstitution,Washingt
D.C. 20560.
Contributionsin Science, Number453, pp. 1-33
Natural HistoryMuseumofLos AngelesCounty, 1995
“Cocculina” lissocona Dali, 1927 31
“Cocculina” rotunda Dali, 1927 32
“Cocculina” superba Clarke, 1960 32
ACKNOWLEDGMENTS
32
LITERATURE CITED 32
2 Contributions in Science, Number453 McLeanand Harasewych: Cocculiniform Limpets
ABSTRACT.WesternAtlanticmembersofthelimpetfamiliesCocculinidaeandPseudococculinidaeare
reviewed. Previouslydescribed species are allocated to currentgenera, and new speciesare described. A
keyto species isprovided.
FamilyCocculinidae: Thegenus CoccocraterHaszpmnar, 1987, is recognized in thewestern Atlantic.
NewspeciesareCocculinamessingi,Cocculinaemsoni,andCoccopigyamikkelsenae.Newcombinations
&
areCoccocraterpocillum(Dali, 1890)andCoccocraterportoricensis(Dali Simpson, 1901). Moskalev’s
(1976)assignmentofFedikovellabeanii(Dali,1882)isconfirmed,andanEnglishtranslationoftheoriginal
descriptionofFedikovellacaymanensisMoskalev,1976,isgiven.LectotypesaredesignatedforCocculina
rathbuniDali, 1882, Coccocraterpocillum (Dali, 1890), andFedikovella beanii(Dali, 1882).
Family Pseudococculinidae: The genera Notocrater Finlay, 1926, Tentaoculus Moskalev, 1976, and
Kaiparapelta Marshall, 1986,are recognized in the western Atlantic. NewspeciesareNotocraterhoub-
ricki, N. youngi, and Kaiparapelta askewi. New combinations are Tentaoculus eritmeta (Verrill, 1884)
and T.georgiana (Dali, 1927). Waren’s (1991)assignmentofCopulabyssia leptalea is confirmed,andan
English translation of Caymanabyssia spina Moskalev, 1976, is given. A lectotype is designated for
Tentaoculusgeorgiana (Dali, 1927).
Six taxa originallyproposed in Cocculina Dali, 1882, are removed from Cocculinidae or Pseudococ-
culinidae andassigned to otherfamilies, although not confirmed as valid species.
INTRODUCTION cific. This number can be expected to increase as
studiesusingresearchsubmersiblescontinue.Allof
Limpets of the families Cocculinidae and Pseudo- the recently added new species and records have
cocculinidaeoccuron biogenicsubstratesatbathy- cometolightfromworkwithsubmersiblesin both
al and abyssal depths. The recent collection of a thewesternAtlanticandeasternPacific,fromeither
number ofnew species as well as fresh material of
sampling of biogenic substrates or experimental
some previously described species has enabled us
work with larval settling.
toreevaluateallthepreviouslydescribedtaxafrom
the tropicalwestern Atlantic and the northwestern MATERIALS AND METHODS
Atlantic.
Untilrecentlythesefamilieswerepoorlyknown. Species forwhich we have freshly collected material are
A worldwide renaissance of interest in the group treated here in detail; those known to us only from the
started with Moskalev (1976), followed by Hick- type material are briefly treated. Revised English trans-
man (1983), Marshall (1986), Haszprunar (1987, ldaetsicornisboefdtbhyeMoroisgkinaalleRvus(1s9i7a6n)daersecrailpstoioinnscloufdtedw.ospecies
1988a, 1988b), McLean (1987, 1988, 1991, 1992), Two species described in 1882 by Daliwere based on
andDantartandLuque(1994).Thenumberofgen- material from widely separated localities, without desig-
era has increased: currently there are 6 genera rec- nation of type localities. We have located only some of
ognized in the family Cocculinidae and 12 in the theoriginalsyntypesintheUSNMcollection,butwetake
family Pseudococculinidae. this opportunityto designatelectotypesand correspond-
During the 19th century there were nine taxa ingtypelocalitiesforCocculinarathbuniDali,1882,and
proposed inthegenus Cocculina from thewestern Fedikovellabeanii(Dali,1882),inordertofacilitatecom-
Atlantic. Original descriptions and illustrations of parisonwiththefreshlycollectedmaterialreportedhere-
the species described byVerrilland Daliwere cop- in.New material, collected using the research submersi-
iedbyPilsbry(1890)andThiele(1909).C.W.John- blesJohnson-Sea-LinkIandIIandClelia,hascomefrom
son (1934) and Abbott (1974) listed 10 and 11 spe- severalsources:threespecieswerecollectedinthecourse
cies,respectively,butnonewinformationwasadd- of experimental work on larval settling conducted off
ed because few species were reported subsequent New Providence Island, Bahamas, by Craig Young, Ro-
to their initial descriptions. landEmson,andPaulA.Tyler;twospeciesweresampled
Although many ofthe previously described spe- duringstudiesofstalkedcrinoidsoffGrandBahamaIsland
cieshavenotbeencollectedagain,thereissufficient by Charles Messing; two species from off St. Vincent,
information in their original descriptions to allow LesserAntilles,werefoundinthecollectionsoftheHar-
most ofthem to be morepreciselyallocatedto the borBranchOceanographicMuseum,andonespecieswas
genera that are now available. Of the 13 species colRlaedctueldaeoffweSroeuthexCaamrionleindawbiythHasrcaasnenwiyncgh.electron mi-
originally described in Cocculina, only 5 now re- croscopy(SEM)afterdissolutionoftissuesinNaOH,air-
main in the family Cocculinidae. Two are now drying,andcoatingwith carbonandgold-palladium. For
placed in the Pseudococculinidae. Six other taxa specimensinwhichthepreservationwasadequate,bodies
proposed in Cocculina are removed to genera in were critical-point dried and coated, and the external
other families but are not allocated at the specific anatomy examined with SEM.
level. Parametersforshellmeasurementsareshowninfigure
ThefourspeciesofCocculinidaeandfivespecies 1. The abbreviations for the measurements are used in
the key to species that follows.
ofPseudococculinidae described from thewestern AbbreviationsofmuseumrepositoriesareLACM,Los
Atlantic over the last two decades bring the total Angeles County Museum of Natural History; HBOM,
to nine species of Cocculinidae and eight species Harbor Branch Oceanographic Museum, Fort Pierce,
ofPseudococculinidae,a total thatgreatlyexceeds Florida; and USNM, National Museum of Natural His-
the numberofspecies known from the eastern Pa- tory,Washington, D.C.
Contributionsin Science, Number453 McLeanand Harasewych: CocculiniformLimpets 3
) . .
Figure 1. Shell measurements. P = apex position, measured from anteriormargin ofthe shell to anterior margin of
the protoconch; SH = shell height, maximum dorsoventral dimension measured perpendicular to the plane of the
aperture; SL = shell length, measured from the anterior margin to the posterior margin of the aperture; SW — shell
width, maximum lateral dimension ofaperture measuredperpendicularto the SL axis.
KEY TO - Shell low (SH/SL > 0.30); apex nearly central
WESTERN ATLANTIC COCCULINIDAE (P/SL < 0.6)
AND PSEUDOCOCCULINIDAE Coccopigya mikkelsenae, new species
6) Copulatory organ at tip of right oral lappet
1) Apical fold of protoconch short and broad; (Cocculina) 7
protoconch sculpture of reticulate or concen- - Copulatoryorganbranchedfromrightcephalic
tric net; radula slightly asymmetrical, 3 inner tentacle Coccocrater 9
- lAaptiecraalltfeoeltdho(fCopcrcuoltionciodnaec)h long and narrow;2 7) Shellhig(h(SH/SL > 0).40),with 1 anteriorand
2 posterior keels; apex posterior of center (P/
pamerntoartsitococalmo,onsc4ihinngsnceutrlhprltaeutaerdresa;lotfreapedtuuhslta(uPlssotesrueodnocgcrlyoysctcaaulslsyimno--r 10)- SSnheLealr>lcle0o.n6wt)e(r.S(.HP.//CSSoLLcc<<ul00i..n368a)),emlsaocnkii,ngnkeeewlss;peacpieex8s
idae) 10
.
2) Protoconch with concentric sculpture Fedi- 8) Shell with radial sculpture moreprominent than
(
kovella 3 concentric sculpture; pigmented eyes absent .
- Protoconch with reticulate (honeycomb) Cocculina rathbuni Dali, 1882
sculpture 4 - Shell with concentric sculpture more promi-
3) Apex oFveedrihkaonvgeilnlgapcoasytemrainoernmiasrgMionskoaflesvhe,ll19.7.6 nenetntthanaxiaCloscccuullptiunrae;mepsisgimnegnit,endeewyesspepcriese-s
- Apexnotoverhangingposteriormarginofshell 9) Shellhigh(SH/SL > 0.45);apexnearposterior
Fedikovella beanii (Dali, 1882) margin (P/SL > 0.7)
4) Shellwithpitsinradialrows;periostracumwith Coccocraterpocillum (Dali, 1890)
raised hairs or spines Coccopygia 5 - Shell low (SH/SL) < 0.45); apex anterior of
- Shell without pits; per(iostracum sm)ooth . . 6 center (P/SL < 0.5)
5) Shellhigh(SH/SL > 0.35);apexnearposterior Coccocraterportoricensis
margin of shell (P/SL > 0.8) (Dali &: Simpson, 1901)
. Coccopigya spinigera (Jeffreys, 1883) Shell with interior septum; protoconch finely
4 Contributions in Science, Number453 McLean and Harasewych: Cocculiniform Limpets
Figures2-5. Cocculina rathbuniDali, 1882. 2. Dorsal, lateral,andventralviews ofshell (USNM 860363). Scale bar
= 1.0 mm. 3, 4. Dorsal and lateral views of protoconch of specimen in figure 1. Scale bars = 50 jam. 5. Detail of
protoconch sculpture in figure 3. Scale bar = 5 jam.
pitted Tentaoculus 11 coarselypustulose;eyespresent;footandman-
( )
- Shelllackinginteriorseptum;protoconchwith- tle edge with fine brown spots
out pits 12 Notocrater houbricki, new species
11) Shell low (SH/SL < 0.40); apex near center - Protoconch coarsely pustulose; teleoconch
(P/SL < 0.6) finely pustulose; eyes absent; foot and mantle
Tentaoculus eritmeta (Verrill, 1884) uniformly beige in color
- Shellhigh(SH/SL > 0.40);apexnearposterior Notocrater youngi, new species
margin (P/SL > 0.7) 15) Shell without posterior ridges or indentations
Tentaoculusgeorgiana (Dali, 1927) in margin; teleoconchwith sharp conicalgran-
12) Teleoconch sculpture granulate ........ 13 ules forming diagonally reticulate pattern
. . .
- Teleoconch sculpture of raised concentric ..... Caymanabyssia spina Moskalev, 1976
ridges 16 - Shell with 2 posterior ridges producing inden-
13) Protoconchwith anastomosing threads; teleo- tations in margin; teleoconch lacking diago-
conch sculpture granulate Notocrater 14 nally oriented conical granules
( ) . . .
- Protoconchwithprismaticcrystals;teleoconch ..........Kaiparapelta askewi, new species
sculpture of anastomosing network ..... 15 16) Shell small, length < 4 mm
14) Protoconch finely pustulose; teleoconch ....... Copulabyssia leptalea (Verrill, 1884)
Contributionsin Science, Number453 McLeanand Harasewych: CocculiniformLimpets 5
Figures6-11. CocculinarathbuniDali, 1882.6.Ventralviewofcritical-point-driedanimal(shellinfigures2-5).Scale
bar = 200 yum. 7. Right lateral view of head. Scale bar = 200 yum. 8. Right epipodial tentacle. Scale bar = 100 yum. 9.
Dorsalviewofradularribbon.Scalebar=20yum. 10.Lateralviewoflongitudinallycutradula,revealingrelativeheights
oftooth fields. Scale bar = 20 yum. 11. Detail ofpluricuspid and marginal teeth. Scale bar = 10 yum.
co = copulatory organ; ct = cephalic tentacle; et = epipodial tentacle; m = marginal teeth; pc = pluricuspid tooth;
pg = pseudoplicatid gill; r = rachidian tooth; 1 = first lateral tooth; 2 = second lateral tooth; 3 = third lateral tooth.
- Shell large, length > 7 mm of cocculinid genera on shell and radular charac-
. . . .Amphiplica venezuelensis McLean, 1988 ters; Haszprunar(1987, 1988b) treated anatomy of
cocculinidgeneraandcomparedthegrouptoother
SYSTEMATICS cocculiniform families. These works should be
Family Cocculinidae Dali, 1882 consulted for further details.
Six genera of Cocculinidae are currently recog-
DIAGNOSIS. Apical fold of protoconch short nized:CocculinaDali, 1882,CoccopigyaMarshall,
and broad; protoconch sculpture of reticulate net. 1986, Coccocrater Haszprunar, 1987, Paracoccu-
Radula slightly asymmetrical, inner lateral teeth 3. linaHaszprunar,1987,FedikovellaMoskalev, 1976,
REMARKS. Marshall (1986) defined a number and Teuthirostria Moskalev, 1976. No species of
6 Contributions in Science, Number4S3 McLean and Harasewych: Cocculiniform Limpets
Figures 12-14. Cocculina messingi, new species. 12. Dorsal, lateral, and ventral views ofholotype. Scale bar — 1.0
mm. 13, 14. Dorsal and lateral views ofprotoconch ofholotype. Scale bars = 50 fim.
Paracocculina and Teuthirostria are known from Species in most genera have a pair of posterior
the western Atlantic or eastern Pacific. epipodial tentacles (figures 5, 8) and a pseudopli-
Four of the genera (excepting Fedikovella and catid gill (figures 15, 19), as defined by Haszprunar
Teuthirostria haveprotoconchsculptureofaraised (1987), on the right side.
)
honeycomb network aligned in longitudinal rows. Marshall considered the ridges on the edge of
The most significant character for separation of the rachidiantooth to representvestigesofthefirst
genera is the position of the copulatory organ in laterals, numbering the further laterals as 2-4.
thesesimultaneoushermaphrodites:modifiedtipof However,toavoidambiguity,we numbertheslen-
right oral lappet in Cocculina, branched from the der lateral teeth that have cusps as 1-3 (figure 10,
base of right cephalic tentacle in Coccopigya and 1-3).Thelargemulticuspidtooththatseparatesthe
Coccocrater, from right side of foot in Paracoc laterals from the marginals is here called the plur-
culina, but yet unknown in Fedikovella and Teu- icuspid tooth (figures 10, 11,pc) ratherthan lateral
thirostria. The genus Coccopigya can be recog- tooth 4, as it has no features in common with the
nized by its hirsuteperiostracum, but no clearsep- other lateral teeth. The pluricuspid teeth are the
aration of Cocculina, Coccocrater, and Paracoc- largestteethandareclearlythemosteffectiveteeth
culinacanbedefinedonshellorradularcharacters. in the row.
Contributionsin Science, Number453 McLean and Harasewych: Coeculiniform Limpets 7
Figures 15-19, Cocculina messingi, new species. 15. Ventral view of critical-point-dried animal of holotype. Scale
bar= 200/im. 16. Anteriormargin oforal lappet. Scale bar= 100/mi. 17. Copulatoryorgan. Scale bar= 100nm. 18.
Right epipodial tentacle. Scale bar = 100/im. 19. Detail ofpseudoplicatidgill. Scale bar = 20 Atm.
ct = cephalic tentacle; et = epipodial tentacle; m = mouth; ol = oral lappet;pg = pseudoplicatidgill.
Genus Cocculina Dali, 1882 In theeastern Pacific,thegenusisrepresented by
C.baxteriMcLean,1987,C.cowaniMcLean,1987,
Type species (subsequent designation, Dali, 1908: and C. craigsmithi McLean, 1992, the last unique
340): Cocculina rathbuni Dali, 1882.
in the family for its occurrence on whale bone.
DIAGNOSIS. Protoconch with honeycomb
sculpture,periostracum smooth; teleoconch sculp- Cocculina rathbuni Dali, 1882
ture offine radial ribsandgrowth lines; copulatory Figures 2-11
organ at tip of right oral lappet.
wesRtEerMnARAtKlaSn.ticCbocyctuhleintaypeisspreecpireessaenntdedtwionntehwe Cocpic.ul15i,nafirgsa.t5h,bu7;niPiDlaslbir,y,1818829:04:0123;2,Daplii.,2158,89fi:g3s.475,,
6 [copy of Dali]; Dali, 1908:340 [type designa-
species here described: C. messingiand C. emsoni.
OtherwesternAtlantictaxadescribed inthisgenus tion]; Thiele, 1909:6,pi. 2, figs. 1, 2; C.W.John-
son, 1934:66 [checklist only]; Abbott, 1974:34,
are herein reallocated to other genera or enumer-
ated under a heading of species for which the ge- fig. 192; McLean, 1987:325, figs. 1-4.
nericaffinityorsynonymyhasnotbeenestablished. DESCRIPTION.Shell(figure2)mediumtolarge-
8 Contributions in Science, Number453 McLean and Harasewych: Cocculiniform Limpets
Figures20-24. Cocculina messingi, new species. 20. Dorsal view of radularribbon. Scale bar = 20 /im. 21. Lateral
view oflongitudinallycut radula revealingrelative heights oftooth fields. Scale bar - 20 ^m. 22. Detail ofrachidian,
lateral, and pluricuspid teeth. Scale bar = 10 jim. 23. Detail of marginal teeth. Scale bar = 6 /xm. 24. Detail of distal
ends ofouter marginal teeth. Scale bar = 2 ixm.
sized for family (maximum length 11 mm), thin, tern, aligned longitudinally in rows (figures 3, 4).
not eroded, white, periostracum thin. Shell height Surfacewithinnettedareaofprotoconchfinelypit-
moderate, that of illustrated specimen 0.35 times ted (figure 5). Tip ofprotoconch immersed in pos-
length.Anteriorandposteriorslopesnearlystraight, teriorslopeofshell.Teleoconchsculptureofraised
lateralslopesslightlyconvex.Outlineindorsalview concentric growth lines and fine radial striae; con-
elongate-oval, anterior end slightly narrower than centricsculpturemoreprominentthanradialsculp-
posterior end; aperture not planar, ends raised rel- ture, not raised at intersections with radial striae.
ative to sides of shell. Apex slightly posterior to Shell edge thin and sharp.
center,protoconch slightly below highest point of Dimensions. Length 11, width 6.5, height 2.75
mm
shell,extendingposteriorly.Protoconchlength240 (original description); length 5.7, width 3.9,
/urn, protoconch sculpture of honeycomb net pat- height 2.0 mm (figure 2).
Contributionsin Science, Number453 McLean and Harasewych: Cocculiniform Limpets 9
Figures 25-28. Cocculina emsoni, new species. 25. Dorsal, lateral, and ventral views of holotype. Scale bar =1.0
mm.26,27.Dorsalandlateralviewsofprotoconchofholotype.Scalebars=50/um.28.Detailofprotoconchsculpture
in figure 26. Scale bar = 5 nm.
External Anatomy (figures 6-8). Eyes lacking, 860363, LACM 151187, HBOM 065:03884. On
copulatory organ at tip of right oral lappet, basal palmetto (Sabalpalmetto) fronds deployed earlier
portion enlarged, producing bilobed effect; pseu- for sampling of invertebrate settling. According to
doplicatid gill on right side dorsal to oral lappet; R. Emson (pers. comm.), this speciesalongwith C.
pair of posterior epipodial tentacles (figures 6, 8), emsoni is frequently recruited on palmetto sub-
with tufts of cilia under high magnification (figure strates at this locality. Further details will be pro-
8); areaat side ofheadwith tufts ofcilia like those vided in a forthcomingpaperbyC.M. Young, P.A.
of epipodial tentacles (figure 7). Tyler, and R.H. Emson.
Radula (figures 9-11). Rachidian broad, outer REMARKS. Dali mentioned material of Coc-
edges weakly defined, tip with single small over- culinarathbunifromthreestations[Massachusetts,
hanging cusp; first lateral with four cusps on outer Barbados, and Martinique] in the original descrip-
edge, second with three, third singly cusped; plur- tion, without designation of a holotype or type
mm
icuspidlongandbroad,with innerandoutercusps. locality.Althoughthe 11 specimen(fromMas-
Marginals similar in size. sachusetts?) has not been located, the other two
NEW RECORD. Off Southwest Reef, New syntypes are present in the USNM. The Barbados
Providence Island, Bahamas (24°54'04"N, specimen (Blake sta. 288, USNM 333750) is a par-
77°33'14"W), 518 m, Johnson-Sea-Link II, dive tially dissected, dried animal with no shell. The
2317,9May1992.Dispositionofspecimens:USNM Martinique specimen (Blake sta. 195, USNM
10 Contributionsin Science, Number 453 McLean and Harasewych: Cocculiniform Limpets