Table Of Content©SociedadEspañoladeMalacología Iberus,25 (2):73-143,2007
Species and diagnosis of the Families and Genera of
Solenogastres (Mollusca)
Especies y diagnosis de las Familias y Géneros de los Moluscos
Solenogastros
ÓscarGARCÍA-ÁLVAREZ*andLuitfriedv.SALVINI-PLAWEN**
Recibidoel23-V-2005.Aceptadoel15-X-2007
ABSTRACT
Although Solenogastres molluscs are notrare animáis, knowledge ofthisgroup ¡svery
¡ncompletesince ¡nformation on ¡ts biology, diversityand geographical distribution ¡s still
limitedandunequal.Overthelastfewdecades, however, researchontheirmorphological
and organizational diversity has increased considerably, and the systematics has been
modified. Forthis reason, thiscontribution summarizesthe diagnoses ofall families and
genera, and compilesa listofthe specieswith ¡nformation abouttheir holotype, distribu-
tion and themostrecentliterature. The holotypeofAnameniaheathiLeloup, 1947, isde-
signatedasneotypeofNeomeniagorgonophilaKowalevsky, 1880.
RESUMEN
Aunque los moluscos solenogastros no son animales raros, el conocimiento que se tiene
sobreestegrupoes muyincompleto,ya quelas informacionessobresu biología, diversi-
dadydistribucióngeográficasonaún limitadasydesiguales. Noobstante, en lasúltimas
décadaslas investigacionessobresudiversidad morfológicayorganizativa se han incre-
mentado notablementeysu sistemática ha sufrido diversas modificaciones. Por ello en
esteartículosehaceuncompendiodelasdiagnosisdetodaslasfamiliasylosgénerosy
serecopila un listadodelasespeciescon informaciónsobresu holotipo, distribuciónyla
bibliografía más reciente. Se designa el holotipo de Anamenia heathi Leloup, 1947,
comoneotipodeNeomeniagorgonophilaKowalevsky, 1880.
KEYWORDS:Mollusca,Solenogastres,diagnosis,families,genera,species.
PALABRASCLAVE:Mollusca,Solenogastres,diagnosis,familias,géneros,especies.
INTRODUCTION
TheSolenogastresareaclassofmol- extremelylimitedanduneven,anddata
luscsofwhichourknowledgeisfairly ontheirbiologycontinuétobescarce.
incomplete. Despite progress in Solenogastres arenot, however, arare
researchingthisclassoverthelastfew animalgroupsincetheyarepresenton
decades,informationontheirdiversity mostmarinebottoms,fromcoastaláreas
and geographical distribution is togreatdepths.
*DepartmentofAnimalBiology,FacultyofBiology,UniversityofSantiagodeCompostela,E-15782Santiago
deCompostela.Spain.E-mail:[email protected]
**Zoologie,UniversitátWien,Althanstrafie14,A-1090Wien,Austria.
73
Iberus,25 (2),2007
Study of the Solenogastres com- 1951; Hoffman 1947/1949; Leloup
mencedinthelastdecadesofthe 19th 1950;Thompson1960;Schwabl1963).
century when Tullberg, in 1875, At the end of the 1960's, the 20th
describedthefirstspeciesNeomeniacari- century passed into a third period.
nata,collectedontheSwedishWestcoast. Salvini-Plawen undertooknew studies
Intheinitialyearsofresearch,thenumber onSolenogastresand,afewyearslater,
ofspeciesdescribedincreasedconsider- Scheltemastartedherworkonthisclass
ably, thanks to the meticulous work of molluscs. These authors described
carried out and improvements to the numerousspecies,makingcontributions
methodsusedforcollectingsamplesat onanatomy,systematicsandbehaviour
greatdepths. Mostofthenew species ofthe group and carryingout modern
carnefromsamplingscarriedoutinthe synopsesontheirorganization(Salvini-
courseofscientificexpeditionsconducted Plawen 1971, 1972, 1985; Scheltema
atthattimeandfrommarineresearchin 1988,1993;Scheltema,Tsecherkassky
thevicinityofBiologicalStations. and Kuzirian, 1994). Salvini-Plawen
Thehistoryofresearchonthismol- (1967) compiled diagnoses of known
luscanclassmaybedividedintothree generaandconstructedkeysforidentify-
largeperiods.Intheinitialperiod,from ingfamiliesandgenera.Someyearslater,
theendofthe19thcenturytothethird he published the monograph on
decade ofthe 20th century, numerous "Antarktische und subantarktische
workswerepublishedonSolenogastres, Solenogastres" (Salvini-Plawen 1978),
byauthorssuchasDanielssen, Heath, on which the current classification is
Hubrecht,Koren,Kovalevsky,Nierstrasz, based.Inthecourseofthepastdecade,
Marión, Odhner, Pruvot, Thiele and new researchers have joined Salvini-
Wirén, inter alia. They described new Plawen and Scheltema in the study of
species,inmanycasesfromtheEuropean thisclassofmolluscs,conductingworks
coastline, and also studied numerous on morphological and organizational
materialfromthevariousscientificexpe- diversity,basedonthedescriptionofnew
ditionsconductedinseas, unexplored species,contributingmuchdataontheir
untilthattime.ThusNierstrasz(1902) biology,distribuciónanddevelopment.
studiedtheSolenogastresfromtheSiboga Inthepastthreedecades sincethe
expeditiontothearchipelagosofSouth- publicationbySalvini-Plawen (1967)
westAsia;Thiele(1902,1906)described ofthekeyforidentifyingthegeneraand
speciescollectedbyValdiviaintheIndian the compilation of their diagnoses,
Ocean; Pelseneer (1903), Nierstrasz knowledgeonthediversityoftheclass
(1908),Thiele(1913a)describedsamples has increased substantially. For this
takenduringvariousAntarcticexpedi- reason,thiscontributionprovidesasy-
tions;andThiele(1911,1932)compiled nopsisofthediagnosesofthefamilies
theSolenogastresfromArcticexpeditions. and genera, and compiles the species
Otherstudieswerealsoconductedon with information about the holotype,
anatomy(Thiele1894;Nierstrasz,1905; distributionandmostrecentliterature.
Heath1905,1914),physiology(Pruvot
1890b; Heath 1904a, 1908), behaviour
(Heath1904b)andlarvaldevelopment SYSTEMATICS
(Pruvot1890a,1892).Compilationswere
publishedbySemroth(1893),Nierstrasz To date, the class Solenogastres
(1909),Thiele(1913b)andHoffmann includes about 250 described species.
(1929-1930). Thecurrentsupragenericclassification
Over the following three decades, (Salvini-Plawen, 1978; Handl and
researchonSolenogastreswaslesspro- Todt,2005)isbasedonthecharactersof
ductive, but several results were of themantlecoverwithcuticleandscle-
special interest, such as those dealing rites, as well as on the types of the
with the anatomy, physiology and radula(monoserial,biserial,distichous,
development(Baba1938, 1940a, 1940b, tetraserial,polyserial/polystichous)and
74
GarcÍA-ÁlvarezANDSALVINI-PLAWEN:SpeciesanddiagnosisofSolenogastres
onthepairedventralforegutglandular of radula may be differentiated
organsassociatedwithit. (Salvini-Plawen,1978;Handl,2002):
-Monoserialtype: each rowformed
Charactersofthesclerites byonetoothoronemoreorlesswide
Scleritesarefoundinsidethecuticle píatewithavariabledenticulation,char-
orprojectingoutofthesame.Thesizeof acteristic of the genera Dondersia,
the sclerites varíes from micrometric Nematomenia,Alexandromeniaandothers.
sizesto3or4mminlength, andthere -Biserialtype: eachrowformedby
aretwobasictypesthatarerelatedtothe twoteethordenticulatedplates,charac-
thicknessofthecuticle.Thethincúneles, teristic ofthe family Simrothielliedae;
particular to the more primitive thepectinatedradula, verycharacteris-
Solenogastres, correlatewithsmoothor tic of the genus Anamenia may be
striated scales arranged in one layer, includedwithinthisgroup.
whereasthickcuticles,characteristicsof -Distichoustype: eachrowformed
themoreevolvedSolenogastres,corre- byapairofhook-shapedteeth,charac-
late with acicular sclerites, which are teristicofWirenia,Pruvotina,Epimenia.
arrangedintwoormorelayers.Inthe -Tetraserialtype: eachrowformed
orderCavibelonia, scleritesarehollow byfourteeth,characteristicofthegenus
aciculars with thick or thin walls, Imeroherpia.
arrangedbothradiallyandtangentially. -Polystichous/Polyserialtype:each
The latter may be located in a single rowwithmanydenticles/teeth,typical
layer,andthusarearrangedobliquely,or of the genera Proneomenia and Dory-
mayformtwoormorelayersofsclerites menia.
interlacing among themselves almost
perpendicularly.Theseareembeddedin Characters of the ventral foregut
thecuticlearrangedsoastoformvery glandularorgans
smallangleswiththemantleepithelium Theventralforegutglandularorgans
(termedasskeletalbyScheltema, 1999; may be absent (Hemineniidae,
Scheltema and Schander, 2000). In NeomeniidaeandBirasoherpia).
addition, SterrofustiaandCavibelonia When these organs are present,
mayhavescleriteswithahookeddistal various types of arrangement may be
endand/orwithanasymmetricallyflat- recognized (Salvini-Plawen, 1978;
teneddistalportion,andinCavibelonia HandlandTodt,2005):
scleritesmayoccurwithadistallyasym- - Types with the main cell bodies
metricaxeorharpoonshape(termedas subepithelially arranged and opening
"cáptate"byArnofsky,2000). intercellularly. This canbe a clustered
type:maincellbodiesofglandssubep-
Charactersoftheradula itheliallyarrangedandopeningintercel-
Solenogastres normally possess a lularlyciósetogetherasapairedgroup
radula,althoughinmanyspecies,thisis directlyintothepharynx;andatypeA:
regressive.Theradularapparatuscom- with subepithelially arranged cell
prisesaradulawithavariablenumber bodiesintoapairofoutleadingducts.
oftransversalrowsofdifferentshaped - TypeB: with apaired outleading
teeth, a cellular reinforcement and an duct,theintercellularlyopeningsubep-
operativemusculature.Groupsofodon- ithelialglandcellsbeingsurroundedby
toblasts in the dorsal radular sheath anoutermusculature.
secrete radula teeth; there is no true - Type C: with a paired duct (sur-
radularmembrane,ratherakindofpre- roundedbymusculature)ofepithelially
ribbonofamaterialidenticaltothatof arrangedglandcells.
theteeth(Wolter,1992).Usedteethare -TypeofSimrothiella: withbulbous
lostor retained in the anterolateral or organs enclosing elongate epithelial
ventralradularsac. glandcellswithmaincellbodiescurved
Depending on the shape and tobelongitudinallyarranged(modified
arrangementoftheteeth,varioustypes typeC).
75
Iberus,25 (2),2007
-TypeDofAmphimeniidae: witha ordersSterrofustia(3families)andCavi-
paired,ramifiedduct,eachbranchbeing belonia(12families).ThePholidoskepia
terminallyprovidedwithapackageof representthebasalgroupandshowdif-
glandcells. ferentlinesofdevelopmentwithinthe
TheClassisdividedintofourorders, orderand,ontheonehand,gaveriseto
grouped within two supraorders. The theNeomeniamorpha,ontheothertothe
supraorderAplotegmentariaincludesthe Sterrofustiaamongwhich,inturn,might
orders Pholidoskepia (6 families) and havebeentheoriginofthepresumably
Neomeniamorpha(2families);thesupra- monophyletic Cavibelonia (Salvini-
order Pachytegmentaria includes the Plawen,1978,1985,2003).
SystematicclassificationofClassSolenogastrestothelevelofgenus
ClassSOLENOGASTRESGegenbaur,1878
SupraorderAPLOTEGMENTARIASalvini-Plawen,1978
OrderPHOLIDOSKEPIASalvini-Plawen,1978
FamilyDondersiidaeSimroth,1893
DondersiaHubrecht,1888
NematomeniaSimroth,1893
IchthyomeniaPilsbry,1898
StylomeniaPruvot,1899
HeathiaThiele,1913
MicromeniaLeloup,1948
LyratoherpiaSalvini-Plawen,1978
HelluoherpiaHandlandBüchinger,1996
SquamatoherpiaBüchingerandHandl,1996
FamilySandalomeniidaeSalvini-Plawen,1978
SandalomeniaThiele,1913
FamilyLepidomeniidaePruvot,1902
LepidomeniaKowalevsky,1883
NierstrasziaHeath,1918
TegulaherpiaSalvini-Plawen,1983
FamilyGymnomeniidaeOdhner,1921
GymnomeniaOdhner,1921
WireniaOdhner,1921
GenitoconiaSalvini-Plawen,1967
FamilyMacellomeniidaeSalvini-Plawen,1978
MacellomeniaSimroth,1893
FamilyMeiomeniidaeSalvini-Plawen1985
MeiomeniaMorse,1979
MeioherpiaSalvini-Plawen,1985
Incertasedis1
PholidoherpiaSalvini-Plawen,1978
OrderNEOMENIAMORPHASalvini-Plawen,1978
FamilyNeomeniidaeIhering,1876
NeomeniaTullberg,1875
FamilyHemimeniidaeSalvini-Plawen,1978
HemimeniaNierstrasz,1902
ArchaeomeniaThiele,1906
SupraorderPACHYTEGMENTARIASalvini-Plawen,1978
OrderSTERROFUSTIASalvini-Plawen,1978
FamilyPhyllomeniidaeSalvini-Plawen,1978
PhyllomeniaThiele,1913
76
GARCLA-ÁLVAREZANDSALVINI-PLAWEN:SpeciesanddiagnosisofSolenogastres
HarpagoherpiaSalvini-Plawen,1978
LituiherpiaSalvini-Plawen,1978
OcheyoherpiaSalvini-Plawen,1978
FamilyImeroherpiidaeSalvini-Plawen,1978
ImeroherpiaSalvini-Plawen,1978
FamilyHeteroherpiidaeSalvini-Plawen,1978
HeteroherpiaSalvini-Plawen,1978
Incertasedis2
RhabdoherpiaSalvini-Plawen,1978
OrderCAVIBELONIASalvini-Plawen,1978
FamilyPruvotinidaeHeath,1911
SubfamilyPararrhopaliinaeSalvini-Plawen,1978
PararrhopaliaSimroth,1893
PruvotinaCockerell,1903
LabidoherpiaSalvini-Plawen,1978
SubfamilyEleutheromeniinaeSalvini-Plawen,1978
EleutheromeniaSalvini-Plawen,1967
GephyroherpiaSalvini-Plawen,1978
LuitfriediaGarcía-ÁlvarezandUrgorri,2001
SubfamilyLophomeniinaeSalvini-Plawen,1978
LophomeniaHeath,1911
MetameniaThiele,1913
HypomeniaVanLummel,1930
ForcepimeniaSalvini-Plawen,1969
SubfamilyHalomeniinaeSalvini-Plawen,1978
HalomeniaHeath,1911
SubfamilyUnciherpenaeGarcía-Álvarez,UrgorriandSalvini-Plawen,2001
UncimeniaNierstrasz,1903
SialoherpiaSalvini-Plawen,1978
UnciherpiaGarcía-Álvarez,UrgorriandSalvini-Plawen,2001
Subfamilyuncertain
ScheltemaiaSalvini-Plawen,2003
FamilyRhopalomeniidaeSalvini-Plawen,1978
RhopalomeniaSimroth,1893
PruvotiaThiele,1894
DinomeniaNierstrasz,1902
DriomeniaHeath,1911
EntonomeniaLeloup,1948
UrgorriaGarcía-ÁlvarezandSalvini-Plawen,2001
FamilyAcanthomeniidaeSalvini-Plawen,1978
AcanthomeniaThiele,1913
AmboherpiaHandlandSalvini-Plawen,2002
FamilyAmphimeniidaeSalvini-Plawen,1972
AmphimeniaThiele,1894
ProparameniaNierstrasz,1902
AlexandromeniaHeath,1911
PachymeniaHeath,1911
SpengelomeniaHeath,1912
ParagymnomeniaLeloup,1947
MeromeniaLeloup,1949
PlathymeniaSchwabl,1961
SputoherpiaSalvini-Plawen,1978
UtralvoherpiaSalvini-Plawen,1978
77
Iberus,25 (2),2007
FamilySimrothiellidaeSalvini-Plawen,1978
SimrothiellaPilsbry,1898
CyclomeniaNierstrasz,1902
KruppomeniaNierstrasz,1903
BiserrameniaSalvini-Plawen,1967
BirasoherpiaSalvini-Plawen,1978
HelicoradomeniaScheltemaandKuzirian,1991
PlaweniaScheltemaandSchander,2000
SpiomeniaArnofsky,2000
AploradoherpiaSalvini-Plawen,2004
FamilyDrepanomeniidaeSalvini-Plawen,1978
DrepanomeniaHeath,1911
FamilyStrophomeniidaeSalvini-Plawen,1978
StrophomeniaPruvot,1899
AnameniaNierstrasz,1908
FamilyProneomeniidaeSimroth,1893
ProneomeniaHubrecht,1880
DorymeniaHeath,1911
FamilyEpimeniidaeSalvini-Plawen,1978
EpimeniaNierstrasz,1908
EpiherpiaSalvini-Plawen,1997
FamilySyngenoherpiidaeSalvini-Plawen,1978
SyngenoherpiaSalvini-Plawen,1978
FamilyRhipidoherpiidaeSalvini-Plawen,1978
RhipidoherpiaSalvini-Plawen,1978
ThieleherpiaSalvini-Plawen,2004
FamilyNotomeniidaeSalvini-Plawen,2004
NotomeniaThiele,1897
KeytoFamilies
1.Scleritesasscalesorasscalesandsolidacicularspicules.Cuticlethinormoderately
thickrelativetothebody 2
1. Sclerites as acicular spicules only. Cuticle generally thick, with epidermal
papillae 3
2.Withscales,ocassionallyacicularsclerites 4
2.Withlancelolatedsclerites(withdistalwings)andsmoothorgroove-likescales,
sometimeswithsolidacides.Withoutventralforegutglandularorgans.Withtwo
pairsofcopulatorystyletswithglands.Withrespiratoryorgans 12
3.Onlywithsolidacicularsclerites 13
3.Withhollowacicularsclerites 17
4. With claviform scales. With monoserial radula of serrate plates. Subepithelial
ventralforegutglandcells(typeA) FamilyMacellomeniidae
4.Withoutclaviformscales 5
5.Withoutradula.Subepithelialventralforegutglandcells(typeA)
FamilyDondersiidae(part)
5.Withradula 6
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GARCÍA-ÁLVAREZANDSALVINI-PLAWEN:SpeciesanddiagnosisofSolenogastres
6.Monoserialradulawithpaireddenticles. Subepithelialventral foregutglandcells
(typeA) FamilyDondersiidae(part)
6.Othertypeofradula 7
7.Radulaformedofsimplyserrateplates(withdenticlessubequalinsize) 8
7.Othertypeofradula(heterodenticulateplatesordistichous) 9
8.Biserialradula(probably)ofserrateplates. Seminalreceptaclesinbundles ...
FamilySandalomeniidae
8.Monoserialradula.Subepithelialventralforegutglandularorgans(typeA) ...
GenusPholidoherpia
9.Biserialradula(withpairedplatesorbarsbearingdenticles).Ventralforegutglan-
dularorgansnotofso-calledtypeA(pairedductwithsubepitheliallyarranged
glandcells) FamilySimrothiellidae(part)
9.Raduladistichous 10
10.Ventralforegutglandularorgansofaccumulatedsubepithelialpharyngealglands
(clusteredtype) FamilyGymnomeniidae
10.Ventralforegutglandularorgansofductswithsubepitheliallyarrangedglandcells
(typeA) 11
11.Withonlyonetypeofscales FamilyLepidomeniidae
11.Withthreeormoretypesofscales,oneprojectingradiallyfromthecuticle
FamilyMeiomeniidae
12.Thincuticlewithoutepidermalpapillae.Withscalesandsolidspicules.Withshort
harpoon-shapedbodiesscleriteswithpedestalrestrictedtocertainpartsofthe
body.Radula(ifpresent)polyserial FamilyHemimeniidae
12.Thickcuticle,ingeneralwithepidermalpapillae.Scleritesassolidaciculesand/or
elongate groove-shaped elements, often distally lanceolate. Radula unknown
/absentinknownspecies FamilyNeomeniidae
13.Withoutradula GenusRhabdoherpia
13.Withradula 14
14.Radulatetraserial.Ventralforegutglandularorganswithsubepithelialglandcells
surroundedbymusculature(typeB) FamilyImeroherpiidae
14.Othertypeofradulaandofventralforegutglandularorgans 15
15.Biserialradula(rowsofpaireddenticulateradulaplatesorbars).Ventralforegut
glandularorgansofaccumulatedsubepitheliallyopeningglandcells (clustered
type) FamilySimrothiellidae(part)
15.Raduladistichous 16
16.Subepithelialglandcellbodiesofventralforegutglandularorgansfree(typeA) .
FamilyPhyllomeniidae
16.Subepithelialglandcellbodiesofventralforegutglandularorganssurroundedby
musculature(typeB) FamilyHeteroherpiidae
17.Withhollow,hookshapedsclerites 18
17.Withouthollow,hookshapedsclerites 19
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Iberas,25 (2),2007
18.Ventralforegutglandularorganswithepithelialglandcells(typeC)
GenusScheltemaia
18.Ventralforegutglandularorgansofductswithsubepitheliallyarrangedglandcells
(typeA)orcircumpharyngealfollicularglands ... FamilyPararrhopaliidae(part)
19.Withoutradula 20
19.Withradula 28
20.Withoutventralforegutglandularorgans FamilyRhopalomeniidae(part)
20.Withventralforegutglandularorgans 21
21.Circumpharyngealfollicularglands FamilyPararrhopaliidae(part)
21.Ventralforegutglandularorgansofanothertype 22
22.Foregutglandularorgansoframifiedductswithterminalclustersofglandcells
(typeD),generallyopeningpre-radularly FamilyAmphimeniidae(part)
22.Foregutglandularorgansofanothertype 23
23.Twotypesofforegutglandularorgans: Gland cellssubepithelial (typeA)and
glandcellsepithelial(typeC) FamilyRhopalomeniidae(part)
23.Foregutglandularorgansofanothertype 24
24. Ventralforegutglandularorganswithsubepithelialglandcellssurroundedby
musculature(typeB) FamilyStrophomeniidae(part)
24.Ventralforegutglandularorgansofanothertype 25
25.Ventralforegutglandularorgansofductswithsubepitheliallyarrangedglandcells
(typeA) 26
25.Ventralforegutglandularorganswithepithelialglandcells(typeC) 27
26.Mantlescleritesacicularandinseverallayers ... FamilyRhopalomeniidae(part)
26.Mantlebodiesstoutwithcavityfilledbyamatrix,arrangedinonelayer
FamilyNotomeniidae
27.Mantlescleritesarrangedtangentiallyinseveralintercrossedlayers.Withoutrespi-
ratoryfolds FamilyRhopalomeniidae(part)
27.Mantlescleritesarrangedradiallyinonelayer.Withrespiratoryorgans
FamilyDrepanomeniidae
28. Withbiserialradula(seriesofpairedplatesorbars).Ventralforegutglandular
organsofaccumulatedsubepitheliallyopeningglandcells(clusteredtype)
FamilySimrothiellidae(part)
28.Othertypeofradulaandofventralforegutglandularorgans 29
29.Radulapectinate(monoserialordividedwithdenseelongatedenticles),ventral
foregutglandularorgansoftubeswithsubepithelialglandcellssurroundedby
musculature(typeB) FamilyStrophomeniidae(part)
29.Othertypeofradula(monoserial,distichous,polyserial/polystichous) 30
30.Withmonoserialradula 31
30.Othertypeofradula 32
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GarcÍA-ÁlvarezANDSad/INI-PlaweN:SpeciesanddiagnosisofSolenogastres
31.Ventralforegutglandularorgansofductswithsubepitheliallyarrangedglandcells
(typeA) FamilyAcanthomeniidae
31.Foregutglandularorgansoframifiedductswithterminalclustersofgland cells
(typeD) FamilyAmphimeniidae(part)
32.Withpolystichous/polyserialradula 33
32.Withdistichousradula 34
33.Ventralforegutglandularorgansofductswithsubepitheliallyarrangedglandcells
(typeA) FamilyRhipidoherpiidae
33.Ventralforegutglandularorganswithepithelialglandcells(typeC)
FamilyProneominiidae
34.Ventralforegutglandularorgansofductswithsubepitheliallyarrangedglandcells
(typeA) 35
34.Ventralforegutglandularorgansofanothertype 36
35.Withoutrespiratoryorgans FamilyRhopalomeniidae(part)
35.Withrespiratoryorgans FamilyPararrhopaliidae(part)
36.Ventralforegutglandularorganswithsubepithelialglandcellssurroundedby
musculature(typeB) FamilySyngenoherpiidae
36.Ventralforegutglandularorganswithepithelialglandcells(typeC)
FamilyEpimeniidae
ClassSolenogastresGegenbaur,1878
SolenogastresGegenbaur,1878,sensuSimroth,1893.Amphineurain:H.G.Bronn'sKlassenand
OrdnungenderTierreichs,3(1):131.
Bilaterally symmetrical molluscs strictions causing lateral pouches.
withthemantle-epitheliumcoveredby Paired ventral and lateralnerve cords
a chitinous cuticle and by calcareous separatedthroughout,inpartganglion-
scleritesofaragonite.Bodylaterallynar- ated. Hermaphrodites, with paired
rowed,footrestrictedtoamedioventral dorsalgonadgenerallyopeningbytwo
groove. Subterminal pallial cavity short ducts into pericardium; pericar-
withoutctenidia,inpartwithsecondary dioductsopeningintospawningducts
respiratoryformations.Mouthopening (internalisedtérmino-lateralportionsof
behindorfusedwithapapilloussensor- pallialcavity).Withoutaortaandexcre-
ialcavity(atrium,vestibulum).Midgut tory organs. Marine, freely moving
straightandspacious,withoutsepárate upon sediments or living epizoic on
digestive gland, due to dorsoventral Cnidaria; carnivorous, mostly feeding
musculaturegenerallywithserialcon- onCnidaria.
SuperorderAplotegmentariaSalvini-Plawen,1978
AplotegmentariaSalvini-Plawen,1978.Zoológica,44(128):23.
Mantle sclerites as scales and/or cavity; mantle cuticle relatively
solid acicular spicules in one layer thin, mostly without epidermal
only, no sclerites with internal papillae.
81
Iberus,25 (2),2007
OrderPholidoskepiaSalvini-Plawen,1978
PholidoskepiaSalvini-Plawen,1978.Zoológica,44(128):23.
Mantlealmostexclusivelywithscaly withsubepitheliallyarrangedglandcell
sclerites.Ventralforegutglandularorgans bodies(clusteredtypeortypeA).
FamilyDondersiidaeSimroth, 1893
DondersiidaeSimroth,1893.Amphineurain:H.G.Bronn'sKlassenandOrdnungenderTierreichs,3
(1):225.
Synonyms:MyzomeniidaeThiele,1894(part).
Sclerites of two or more types of denticles. Ventral foregut glandular
scales, occasionallywithsolid acicular organsofductswithsubepithelialgland
spicules. Without epidermal papillae. cells (type A). Without respiratory
Radula monoserial, with paired long organs.
GenusDondersiaHubrecht,1888
DondersiaHubrecht,1888.DondersFeestbrendel,Amsterdam:324.
Typespecies:DondersiafestivaHubrecht,1888.
At least with two types of scales. lateral constrictions. Unpaired sec-
Mouth separated from the atrium. ondarygenitalopening.Withdorsoter-
Radula-plateswithonepairoflateral, minalsenseorgan.Withoutcopulatory
mediallycurveddenticles.Midgutwith styletsandrespiratoryorgans.
DondersiafestivaHubrecht,1888,inFeestbundelDondersTijdschr.Geneesk.(Amsterdam)
Holotype:GulfofNaples(Italy);60m;Mus.Nat.Hist.Leiden,Netherlands.
Literature:NierstraszandStork,1940,inZoológica,36(99).
DondersiaannulataNierstrasz,1902,inSiboga-Exp.Monogr.,47
Holotype:Bima,Sumbawa,Sunda-Sea(Indonesia)(SibogaSt.47);55m;Zool.Mus.Univ.Ams-
terdam,Netherlands.
Literature:Nierstrasz,1902,inSiboga-Exp.Monogr.,47.
DondersiacalifornicaHeath,1911,inMem.Mus.Comp.ZoolHarvardColl,45(1)
Holotype:SanDiego,(California,USA)(AlbatrossSt.4303);38m;Calif.Acad.Sci.,SanFrancisco,
USA.
Literature:Heath,1911,inMem.Mus.Comp.Zool.HarvardColl,45(1).
DondersiacnidevoransSalvini-Plawen,1978,inZoológica,44(128)
Holotype:RossSea(Antárctica);659-714m;SmithsonianInstitution(Nat.Mus.Nat.Hist.),
WashingtonDC,USA.
Literature:Salvini-Plawen,1978,inZoológica,44(128).
DondersiaindicaStork,1941,inSiboga-Exp.Monogr.,47b
Holotype:Bima,Sumbawa,Sunda-Sea,(Indonesia);55m;(Typematerialmissing).
Literature:Stork,1941,inSiboga-Exp.Monogr.,47b.
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