Pleurobranchaea inconspicua Bergh, 1897 (Opisthobranchia: Pleurobranchidae): Redescription and distribution from Argentina and Colombia

2004·5.4 MB·English

by  Claudia Muniain

Bonnerzoologische Beiträge Band 55 (2006) Heft 3/4 Seiten 291-300 Bonn, November2007
Pleiirobranchaea iuconspicua Bergh, 1897
(Opisthobranchia: Pleurobranchidae):
Redescription and distribution from Argentina and Colombia
&
Claudia Muniain'>, Néstor E. Ardila^) Juan Lucas Cervera^'
1'Museo Argentino de Ciencias Naturales "Bernardino Rivadavia", Buenos Aires, Argentina
2)Universidad de los Andes, Laboratorio de Biología Molecular Marina, Laboratorio de Botánica y Siste-
mática, Bogotá, Colombia
3'Universidad de Cádiz, Departamento de Biología, Cádiz, Spain
*Paperpresentedtothe 2nd International Workshopon Opisthobranchia, ZFMK, Bonn, Germany, September20thto 22nd, 2006
Abstract. Ataxonomic slT.idy based on fresh specimens fromArgentina and Colombia, and vouchermaterial from the
collectionsoftheMuseoArgentinodeCienciasNaturales"BernardinoRivadavia"(MACN)andMuseodeHistoriaNa-
turalMarinadeColombia(MHNMC-INVEMAR)wasconducted.Thefirststudybyscanningelectronmicroscopy(SEM)
oftheradular,jawelementsandreproductive anatomy, paying special attention tothe internal cuticularstyletofthepe-
nis, is presented. Previous descriptions ofP. incoiispicua and itsjunior synonyms, P. hamva. P. Iiedgpelhi, andP. gela,
arecomparedwiththepresentresults;P. boiuiieaeis synonymizedandachecklistofthedistinguishingfeaturesofrele-
vant species within the genus Pleurohraiichaea is provided. This work aims at discussing the proposed amphiatlantic
distributionofP. incoiispicuaandatsolvingtheproblemsofsynonymies. Itprovidesastartingpoint foracomparative
studyofspecimensofthe genusPleurobninchaea fromboth sidesoftheAtlantic Ocean andthe Mediterranean Seathat
willbecomplementedwith molecularstudies in future times.
Keywords.Taxonomic redescription, Colombian Caribbean,Argentina, amphiatlantic distribution.
L INTRODUCTION
ThegenusPleiirobranchaea Meckel inLeue, 1813 isrep- et al. (2006), although both works refer to the original
resented in temperate-wann waters, by several species description only.
with wide geographical distribution ranges. Pleiirobran-
chaeainconspiciia Bergh, 1897wasdescribedfromnorth- In Marcus & Gosliner's revision (1984), the names P.
emBrazil (type locality: Sergipe, 10"54'S, 37°7'W),and hedgpethi Abbott, 1952, P. hamva Marcus and Marcus,
its distribution was later extended under the same name 1955 (mispelledhannva)andP.gela MarcusandMarcus,
orunderjuniorsynonymsofspeciesandsubspeciestosev- 1966wereconsideredJuniorsynonymsofP. inconspiciia.
eral localities in the Western Atlantic Ocean (USA, Abbott(1952) describedP. hedgpethifrom PortAransas
Caribbean, Brazil, Argentina) and to localities more dis- (GulfofMexico); this speciesbecame themost frequent-
tantfrointhetype locality, such asthe Mediten^anean Sea ly mentioned junior synonym ofP. inconspiciia for the
(Israel) and WestAfrica (Ivory Coast to Nigeria). Western Atlantic, even more frequently than the senior
synonym. However, in the same revision, these authors
Marcus & Gosliner (1984) gave the most recent revi- erected the species P. bonnieae for one specimen from
sion ofthe subfamily Pleurobranchaeinae, which includ- Florida region. According to Valdes et al. (2006) this
ed 14 species ofthe genus Pleiirobranchaea. However, namecouldbeanotherjuniorsynonym ofP. inconspiciia.
theseauthorsdidnotincludethe speciesP. morosa Bergh,
1892,P morula Bergh, 1905, P melcmopiis Bergh, 1907, Several authors considered the anatomy ofthe reproduc-
P. algoensis Thiele, 1925, P.japónica Thiele, 1925, and tive system and the penial moiphology as the most use-
P. dorsali Allan, 1933, because they were poorly de- ful characters to discriminate species ofthe genus Pleii-
scribed. Nevertheless, P. japónica was redescribcd by robranchaea coiupared to the classical hard structures of
TsuBOKAWA et al. (1992), confirming the validity ofthe radulae andjaws (Marcus 1961; Marcus & Gosliner
species and P. morosa has been recently mentioned froiu 1984; Willan 1987).
M
Azoreanarchipelagoby alaquias(2001 and Cervera
)
Fig. 1. Pleiirohranchaeainconspiciia Bcrgh 1897.A. Dorsalviewoflivingspecimen fromColombia(Photo: N.Ardila). B. Dor-
sal viewofhead. C. Detail ofthegill andthe pairsofpinnules. D. Footshowingaconspicuous spur. E. Livingspecimen from Pa-
mm
tagonia (80 length) (Photo: C. Muniain). F. Detail ofrhinophores and sensory papillae on the oral veil (Photo. C. Muniain).
Bonnerzoologische Beiträge 55 (2006) 293
UnlikemostofthepreviousdescriptionsofP. iiiconspicini Twopenisesandtheirrespectivecuticularstyletswerepre-
and of other species of the genus conducted from pre- viously critical-point-dried for SEM (MACN-in: 36985,
served material, often in bad conditions, this work re- 37064).
describes the characteristics ofrecently collected speci-
mens ofP. inconspiciia from two distant localities.
RESULTS
3.
This isthe first study using SEM forhard structures ofP.
inconspiciia, especially the examination of the penial 3.L Systematic description
stylet. Detaileddescriptionsoftheexternalappearanceand
thehard structures, aswell asthereproductive system, of Nudipleura Wägele & Willan, 2000
specimens from distant localities help to understand the Pleurobranchoidea Gray, 1827
intraspecific variability in this species, and allow to dis- Pleurobranchidae Gray, 1827
cuss the possible amphiatlantic distribution ofP. incon- Pleurobranchaeinae Pilsbiy, 1896
spiciia by comparison with material from other Atlantic Pleurohranchaea Meckel in Leue, 1813 (type Pleuro-
localities.Here, synonymizationofotherPleurohranchaea brancliidiiim niecl<elii Blainville, 1826)
species with P. inconspiciia is outlined.
Pleurohranchaea inconspicua Bergh, 1897 (Figs 1-5)
2. MATERIALSAND METHODS Pleurohranchaea inconspicua Bergh, 1897 (Bergh 1897:
49-51,pi. 8, figs2-10. Marcus& Gosliner 1984: 24-27,
Specimens were collected over a wide range of depths figs IB, 10-ll.Rlos 1984:206-207,pl.69, fig. 987. Mu-
from shallow intertidal areas down to 150 m in depth. A NiAiN 1997: 4, 17, 34. Ardila & Rachello 2004: 62,
totalof16specimenswerepreservedin70%ethanol. Pho- fig.3).
tographs and videos were obtained with a digital camera
from living specimens. Part ofthis material is deposited Pleurohranchaea hedgpethiAhhott, 1952 (Abbott 1952:
in the Museo de Historia Natural Marina de Colombia 1-2, pi. 1, figs 1-8. Marcus & Marcus 1959: 253 fig.
(MHNMC-INVEMAR) as well as in the MuseoArgenti- 6. Marcus & Marcus 1960: 253-254, fig. 6. Nussen-
no de Ciencias Naturales "Bernardino Rivadavia" Meyer 1965: 143-145, figs 1-2. Marcus & Marcus
(MACN). 1967a: fig. 55C. Marcus & Marcus 1967b: 200. Mar-
cus & Marcus 1969: 18. Abott 1974: 348, fig. 4201.
Colombia: Fourspecimenscollectedwith abottomtrawl
(9 X 1 m opening, 16 m in length) during the research Pleurohranchaea hamva Marcus & Marcus, 1957 (Mar-
cruisesINVEMAR-MACROFAUNA11, 2001 aboardthe cus & Marcus 1957: 21-27, figs 40-52).
RA^Ancon, intheCaribbeancoastofColombiaat70and
150 m depth; two specimens offDibulla (11° 25' 34"N, Pleurohranchaea hedgpethi hamva Marcus, 1961
73° 27' 40"W), E-lOO, 150 m depth (MHNMC-INVE- (Marcus Er. 1961: 141. Marcus& Marcus 1967a: 48).
MAR: 3872, 3873), two specimens offBuritaca (11° 18'
28"N, 73°46'50"W) E-108. 109, 70mdepth (MHNMC- Pleurohranchaeagela Marcus& Marcus, 1966(Marcus
INVEMAR: 3874, 3875). One specimenwascollectedby & Marcus 1966:"i74, figs 35-37.
SCUBA from Neguanje Bay, 5-10 m depth in a culture
ofbivalves (Ardila priv. coll.). Pleurohranchaea honnieae Marcus & Gosliner, 1984
(Marcus & Gosliner 1984: 29-32, figs ID, 13-14.).
Argentina: One specimen from Puerto Quequén (38°
34'S, 58° 38"W) Buenos Aires, 12/1928, MACN-in: 3.LI. External features
18234; onespecimenfromPuertoQuequén, BuenosAires, Length of the living animals ranges between 13 and 80
2/1929, MACN-in: 18312; two specimens from Puerto mm.The body is oval and elongate. The ground colour is
Quequén, BuenosAires, I/I934, MACN-in: 412; sixspec- pale cream, with a dense dark brown reticulate along the
imens from Creek Bay (41° 6'S, 63° 56'W), San Matías mantledorsal region,beinglighterand lessobvious in the
Gulf, Rio Negro, MACN-in: 36985; one specimen from foot dorsal region and absent in the well developed
Larralde (42° 25'S, 64° 19'0), San José Gulf, Chubut, rhinophores (Fig. lA, B, E). Bright white spots are scat-
10/1/2006, MACN-in: 37064. teredall overthemantleandmoreconspicuousonthedor-
sal footsurface. Theoral veil isbroadwith sensorypapil-
Two specimens were dissected, and their jaws and radu- lae along the anterior edge. Rinophores are held erect in
laewereremovedandmountedforScanningElectronMi- the living animals (Fig. IB, F), whereas in the preserved
croscopy(SEM)examination (MACN-in: 36985, 37064). ones they are in a lateral position.
294 Claudia Muniain et al.: Pleurohranchaea iuconspicua fromArgentina and Colombia
mm
Themantle isreducedand fusedwith theoral veil; a large geal bulb. The radularfomiulaofone specimenof40
gill is visible on the right side ofthe body. The gill is at- in life is 30 X 64.0.64 (MACN-in: 36985). The radula
tachedalongmostofthebodywith about26pairs ofpin- lacks a rachidian tooth, but a difference among the first
nules per gill (Fig. 1 C, E). The anal papilla lies over the fourlaterals isnoticeable in someradularrows (Fig. 2A).
seventh pinnule. The foot is rounded in front. The soft Lateral teetharebicuspid, largestin themiddleofthehalf-
mantle, notcovering the foot, is visibleto mostofitspart row and smaller towards the outerborder (Fig. 2B). The
(Fig. 1 A, E). The coloration in the ventral foot region is jaw elements (rodlets) bear4 to 8 denticles from the out-
translucent, and the internal organs can be observed by emiost and innermost region (Figs 2C. D).
transparency.Attheposteriorend,thefootends inaround-
ed shape, with a large metapodial gland (yellowish) ex- Thereproductivesystemarrangementissimilartothatof
tending longitudinally from theposteriormantle towards otherPleurohranchaea species (Fig. 3). The tubular am-
the central foot sole (Fig. 1 E). Dorsally to the metapodi- pulla forks near its outlet into two branches, the shorter
al gland, the foothasabrown orblacktiny spur(Fig. 1A, branch istheoviductthatentersthecapsulegland, andthe
D). other branch is the efferent duct that splits in two. One
branch enters the prostate and then continues within the
3.1.2. Anatomy penial sac (Fig. 3, number4). The secondbranch leadsto
Ashell is lacking. Thepharyngealbulb frequentlyprotmd- the bilobed seminal receptacle, which is connected to the
edinpreservedspecimensthroughtheeversión ofthevery bursa copulatrix by a long and thin convoluted duct. The
extensible oral tube. Jaws consistoftwo paired and elon- bursa copulatrix is large and round.
gate chitinous plates on the lateral walls ofthe pharyn-
Fig. 2. A-D. Scanning electron photographs ofradula ofP. inconspiciiu MACN-in: 36985. A. Radula lacking a rachidian cen-
tral tooth. B. Outennost teeth on the same radula side (right). C, D. Jaw showing platelets with4-8 denticles from outennostre-
gion. D, Jaw showing platelets from innennost region.
Bonner zoologische Beiträge 55 (2006) 295
Fig. 3. Reproductive system ofP. inconspiciia MACN-in: 36985. Scale bar: Imm. Abbreviations: AM ampulla, BC bursa copu-
latrix. CGcapsulegland, EDefferentduct. LPSTpenis loopcontainingthecuticularstylet,OVoviduct, PPpenial papilla, PR pro-
state, PS penial sac, PSTpenis (cuticular stylet), SR seminal receptacle, VA vagina. For numbers see text.
The penis is an elastic cutilar stylet, that coils in numer- cuticule (Fig. 4E). Apparently, as observed from the de-
ous helically loops (Fig. 3, number 3), that finally enters tail ofthe final portion ofthe efferent duct, in its portion
inthe atriumthrough apenial papilla (Fig. 3, numbers 2, connectingwith theprostate, a central cuticularstylet in-
1). It is difficult to find the point where the efferent duct side it would be lacking (Fig. 4F).
and the penis are connected. When stretched out, the pe-
nis reaches a length of 16 and 20 cm in specimens of45 3.2. Geographic distribution
mm
and 80 length respectively. Observations ofthe pe-
nisunderSEMreveal anexternal cuticuleandinternalcu- The geographical distribution cited for P. inconspiciia is
W
ficular stylet throughout its length (Fig. 4A-E). from Cape Halteras, North Carolina 35° N 80° (USA)
W
to San José Gulf42° S 64° (Argentina) (See Table 1.
Atransversal section ofthepenis showsthatthe chitinous Fig. 5). Regarding the Caribbean distribution of this
stylet is not rounded and has a central cylindrical orifice. species,thefirstrecord fromColombia wasgivenbyArdi-
The shape and size ofthese penial stylets from two spec- LA & Rachello (2004), with specitncns from 70-150 m
imens examined under SEM are similar (Figs A, B, C). and an individual collected from 8-10 m, present study.
The width ofthose stylets is about 80 |am. The penis has
morethan 15 roundish loops(Fig.3, LPST, number3),and Marcus & Marcus (1969) recorded specimens for the
contains the cuticularstyletall along its length (Fig. 4D). first time (as P. hedgpethi) forArgentina (40" 32' S, 60°
The penial stylet runs only along one fifth ofthe penial 19' W, 19 specimens at 57 m) and (40" ITS, 60° 27' W,
296 Claudia Muniain et al.: Plciunhrcinchaea inconspiciiu from Argentina and Colombia
Fig. 4. SEM (cntieal-point-dned)ofreproductivesystemoíP. inconspiciiu(MACN-in: 36985; 37064). A. Penis, showingtheex-
ternal euticule and internal ehitinous stylet (MACN-in: 36985); B and C. Detail ofthe tranverse seetion ofthe cuticular styletof
two different specimens (B, MACN-in: 36985; C, MACN-in: 37064). D. Portion ofpenial loop with a length of20 em from spe-
cimenof80 mm in length (MACN-in: 37064). E. Dorsal viewofthe external euticule, showingaportion ofthecuticularstyletin-
side it (MACN-in: 36985). F. Detail ofthe region ofthe efferent duct that leaves the prostata (MACN-in: 37064).
7 specimens at 44 m), and their geographical range has 4. DISCUSSION
beenextended to northern Patagonia someyears laterfor
the intertidal to 8 m (Muniain,1997, present study). Asmentionedabove,thiswork intendstoserveasastart-
ingpoint for furthertaxonomic and molecularstudies in-
Marcus & Gosliner (1984) cited localities frotn the volvingalargernumberoffreshmaterialtoverifywhether
Mediterranean Sea (Israel) and WestAfrica (Ivory Coast this species is also present along the western coasts of
to Nigeria). Further investigations will allow testing the Africa and the eastern Mediterranean (Israel) (amphiat-
hypothesis ofan amphiatlantic distribution. lantic species), as Marcus & Gosliner (1984) stated, or
Bonnerzoologische Beiträge 55 (2006) 297
-15°N ^^Caribbean Sea\ J
ATLANTIC
)
/• OCEAN
-15°S
.
j
y
- 30° s
J
-45° S
^
1
75° W 45° W 15°W 15° E
Fig. 5. Known distribution oíPleurobrauchaeu incunspicua Bergh, 1897, including its synonyms.
to confirmthe validity ofother species ofthe genus (e.g. imal. Besides all the facts stated above, Marcus &
P. tarda from Western and South Eastern Atlantic, Table Gosliner (1984) provide unclear infonnation about the
l,Fig. 5). collecting localitiesoftheirmaterial off! inconspiciia,but
theygiveageographical range inthewesternAtlanticthat
Thus, Valdés etal. (2006) suggest thatP. bonnieae Mar- would include the type locality ofP. bonnieae. Another
cus and Gosliner, 1984 could be ajunior synonym ofP. species of Pleiirobrancliaea, P. agassizi Bergh, 1897,
inconspiciia.Weagreewiththisopinion,sincethedescrip- would fitthegeographical range iorP. inconspiciia. How-
tionofthisspeciesisbasedonasinglepreservedand, very ever, the coloiu"ofliving specimens ofP. agassizi, its ba-
probably,juvenilespecimenandthe features suggestedto thymetricrangeandsomeanatomicaldifferencesconcern-
differentiateitfromothernominal species(P. inconspiciia ingtheradularteethandthereproductive system makeus
orP. vayssierei) are weak (presence ofa penial cuticle or considerthis species as valid until additional fresh mate-
diameter ofthe base ofthe penis as they can be artifacts rial can be studied.
ofpreservation). There is no description ofthe living an-
298 Claudia Muniain et al.: Plewobranchaea inconspicua fromArgentina andColombia
ExternalandinternalcharacteristicsofP. iuconspiciiahave from the Mediten"anean Sea (Israel), will allow to eluci-
been described from fresh collected specimens from two date the existence ofmore than one species ofthis genus
distant localities ofits wide distribution, as well as from inthe MediteiTaneanorconfirmthatP. meckeliiistheon-
preserved material. The number ofpinnae ofthe gill and ly species present in this region.
the presence ofa conspicuous caudal spur are shared by
all dissected specimens. All Patagonian living specimens Acknowledgements. The authors are grateful to the staff of
showedthecaudal spur;but it ispossiblethatafterpreser- Museo de la Plata, Patricia Sarmiento and MuseoArgentino de
vation the spur is not evident, and forthis reason in some Ciencias Naturales "Bernardino Rivadavia", Fabián Tricárico,
descriptions it might have notbeenmentioned. The same fortheirvaluable assistanceduring SEM sessions.AndtoNés-
hypothesis can be drawn for the posterior region ofthe tor Cazzaniga (Universidad del Sur, Bahía Blanca) for kindly
foot sole, inwhich ayellow mctapodial glandfonnsalon- providingspecimensforthisstudyfromRioNegro.Twoanony-
mousreviewersprovidedhelpful comments onthemanuscript.
gitudinal fuiTow,oftenvisiblebytransparencythroughthe
skin ofthe dorsal side ofthe tail. The internal anatomy, This study was funded byArgentinian grants from GEF-BIRF
the radula, thejaw elements and the aiTangcmcnt ofthe 28385/AR(A-CB-51)andANPCyT(PICT34111)and Spanish
grantGCL2006-05182/BOS(DGloftheEducationandScience
reproductiveorgans fullyagreewith previousdescriptions. Ministry). CM. is Research Memberofthe National Research
The morphology ofthe internal cuticularcylindrical stylet Council ofArgentina (CONICET).
issimilartothat illustratedby Marcus&Gosliner(1984,
fig. IB) by optical transverse section.
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300 Claudia Muniain etal.: Pleurohranchaea inconspiciia fromArgentina and Colombia
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