THE NAUTILUS

Volume 128, Number 2

June 20, 2014

ISSN 0028-1344

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THEe^NAUTILUS

Volume 128, Number 2

June 20, 2014

ISSN 0028-1344

Erratum

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THE NAUTILUS 128(2):31-39, 2014

Page 31

New Scutellastra (Gastropoda: Patellidae) species from the

lower Miocene Navidad Formation of central Chile and the

lower-middle Miocene Cantaure Formation of Venezuela

Sven N. Nielsen

Uiiiversidad Austral de Chile

Institiito de Ciencias Ainbieiitales y Evolutivas

Casilla 567, ValdiUa, Chile

[email protected]

Bernard Landau

Naturalis Biodiversity Center

Leiden, The Netherlands

and

Centro de Geologia da Universidade de Lisboa

Lisboa, Portugal

and

International Health Centres

Av. Infante de Henrique 7

Areias Sao Joao, P-8200-261

Albufeira, Portugal

l)[email protected]

ABSTRACT

New species of the patellid gastropod genus Saitellastra are

described from the lower Miocene Navidad Formation of cen-

tral Chile and the lower-middle Miocene Cantaure Formation of

Venezuela. Saitellastra arayae new species is the first fossil

record of a patellid from Chile and S. venezuelana new species

is the first from tropical America. This genus is today restricted

to soutliem Africa and the western Pacific, with the exception of

S. mexicana from Central America. Together with contempora-

neous fossil occurrences in New Zealand, these new records

show that Saitellastra had a much wider distribution when

water temperatures were higher during tlie mid-Cenozoic.

Additional Keywords: Paleontology, marine, Patellogastropoda

INTRODUCTION

The systematic position of the genus Scutellastra H. and

A. Adams, 1854 was considered problematic in the

Southern Synthesis (Lindberg, 1998), whereas it is

clearly included in Patellidae in the phylogenetic analysis

of Ridgway et al. (1998). Further investigations by

Koufopanou et al. (1999), Lindberg (2007), and Nakano

and Ozawa (2007) suggested that the genus Scutellastra

may be polyphyletic with respect to the genera Helcion

Montfort, 1810 and Cijmhiila H. and A. Adams, 1854.

Lindberg (2007) also mentioned a “scutellastrid clade”

being the “sister taxon of [. . .] the Patellidae” but did

not formally name it. It would contain the genera

Scutellastra, Helcion, and Cijmbula. We here follow the

traditional view that recognizes tire sole family Patellidae

within Patelloidea (Bouchet and Rocroi, 2005).

Extant species of Scutellastra are present almost

exclusively in the Atlantic, in southern Africa, and in the

Indian Ocean, in southern Australia (Ridgway et al.,

1998), with few species extending throughout the west-

ern Pacific, and only S. mexicana (Broderip and

Sowerby, 1829) is present in the eastern Pacific, in the

Americas, ranging from western Mexico to Peru. Species

of Saitellastra are usually intertidal, but some live just

subtidally (Lindberg, 1998). They are generally found in

tropical to subtropical regions, with the species of south-

ern Australia being the temperate end-members. The

fossil record of Scutellastra goes back to the upper

Cretaceous of Japan (Kase and Shigeta, 1996).

Patellogastropod limpets are a dominant group today

in intertidal environments of Pacific southern South

America, represented by the nacellid genus Nacella

(Valdovinos and Riith, 2005; Gonzalez-Wevar et al.,

2011) and the lottiid genus Scurria (Espoz et al., 2004).

The Chilean fossil record of limpets in beds older than

Pleistwene is scarce. Only a single specimen of Nacella

{Patinigera) intifo'nna DeVries, 2008 from the Huenteguapi

Sandstone (Le Roux et al, 2008) on Arauco Peninsula and

the liolotype of Nacella (Patinigera) nielseni DeVries,

2008 from Cliiloe Island (Watters and Fleming, 1972) are

known from the Neogene of Chile. The only other large

patellogastropod limpet is the smooth-shelled nacellid

Cellana fiienzalidai (Hemi, 1969) from the Pliocene of

northern Chile (Herm, 1969; Lindberg and Hickjiian,

1986). All these belong in the family Nacellidae.

The tropical American Neogene patellogastropod lim-

pet record is even poorer, with no fossil species at al!

described from any of the ricli Caribbean or Panamic

Pacific assemblages, except for the shell illustrated by

Lindberg (2007, text-figure 8B) as Scutellastra sp. from

Page 32

THE NAUTILUS, Vol. 128, No. 2

the Pliocene of Venezuela. Lindberg (2007) did not provide

the localih' for thi.s specimen, but, according to its localit)'

record, it was collected at approximately 300 m soutli of

Casa Cantanre (D.R. Lindberg personal communication),

which is the only place where we have found representa-

tives of Saiiellmtra in Venezuela. It is undoul)tedly con-

specific witli the new Venezuelan species described herein

and dates therefore from the Miocene, not Pliocene.

GEOLOGY OF FOSSIL-BEARING LOCALITIES

Chile: The Chilean Scufella.stra comes from reddish

sandstones of the Navidad Formation (Encinas et ah,

2006) at Punta Perro, central Chile (Figure 1). These

deposits were dated as upper Miocene by Finger et al.

(2007, locality PPN) l)ased on misidentified foraminifera

(Finger, 2013). The mollusk assemblage was interpreted

as being reworked from lower Miocene betls based c)n

comparison with southern Peni (DeVries and Frassinetti,

2003; Finger et ah, 2(M)7), an inteqiretation confirmed

with strontium isotope dating by Nhelsen and Glodny

(2009). Recently revised identification of stratigraphic

index foraminifera (Finger, 2013), however, now agrees

with a lower Miocene age for the Navidad Formation

and also explains earlier assignments to younger ages

(e.g., Ibaraki, 1992). These new data provide a consensus

for the lower Miocene age of the Navidad Formation.

The displacement scenario of Finger et al. (2007) is still

valid, although it has now been demonstrated that con-

temporaneous shallow-water deposits were transported

into bathyal depths, as indicated by the strontium isotope

data of Nielsen and Glodnv (2009). Many shallow-water

taxa occur in sediments intercalated with deep-water

deposits containing a completely different fauna (see

Finger et ah, 2007).

The assemblage from the same concretionaiy block that

contained the Saitellastra specimen includes Pinna

semicostata, Ghjcijineris sp., Astele chilemis, Echimyphoria

monilifer, Distorsio ringens, Glossaulax pachystomi,

Magnatica suhsohda, Sinnm mbglohosiim, Lampnxkmiina

(Hinidkifa, Testallium cepa, Austrotxmm echimdata. Inquis-

itor lingidacaninns, Dentalinm sp. and represents the ty{3-

ical shallow-water Navidad assemblage (see Finger et al.,

2007; Griffin and Nielsen, 2008; Kiel and Nielsen, 2010).

Venezuela: The Venezuelan Saitellastra material

lierein described and discussed comes from the San Jose

de Cocodite region in the Paraguana Peninsula of north-

ern Venezuela (Falcon State). The collection site where

S.N. Nielsen and B. Lanciau, 2014

Page 33

it originates from is located in the Caiitaure area, 3.4 km

West of the cliurch of tlie village of San Jose de Cocodite

(as tile crow flies), at an altitude of approximately 140 m

al)()ve sea-level on a acacia and cactus covered area

approximately 400 m south of Casa Cantaure with

tlie approximate geographic coordinates: 11°56'24.]" N,

70° 01' 04.5" W (Figure 2; location of Casa Cantaure

after Griffiths et al., 2013: 11°56'35.9" N, 70°01'10.8" W).

The specimens were collected from a thick, friable,

yellow, fine sandstone !)ed containing an abundant and

diversified molluscan assemblage (mostly gastropods and

bi\-alves with rare iiautiloid cephalopods), as well as

other elements such as barnacles and corals. This bed is

part of the Cantaure Formation (Jung, 1965; Hunter and

Bartok, 1974), which, as a whole, according to Diaz de

Gcunero (1974), is correlated with the planktonic forami-

niferaJ biozones Glohigerinatella insueta and Praeorbulina

glomerosa of Bolli (1966), biozones N7 and N8 of Blow

(1969), which in turn, according to tlie latest geologic time

scale of Gradstein et al. (2012), correspond to the Lower

to Middle Miocene transition, upper Burdigalian to lower

Langhian. Key (1996) corroborates this biostratigraphic

correlation stating that the Cantaure calcareous nanno-

fossil assemblage contains the Helicosphaera amj)lmpert.a

and Sphenolithus heteromorj)hus markers corresponding

to the biozones NN4 and NN5 of Martini (1971), which

broadly correlate with the above inentioiied foraminif-

eral zones.

In several recent papers, however, tli(> (iantaure For-

mation continues to be assigned to the Lower Miocene,

Burdigalian, after the traditional correlation of Diaz de

Gamero (1974) and Key (1996). Aguilera and Rodrigues

de Aguilera (1999), !)ased on planktonic foraminifera

data from a personal communication by Collins, place

the Cantaure Formation in the Lower Miocene,

Burdigalian. Griffiths et al. (2013), based on ^'Sr/^^^Sr

isotope data obtained from corals, assign an age of

between 16.3 and 16.6 Ma to the fossils of Cantaure,

placing them in the Burdigalian. Tliese antliors further

comment that the isotopic results obtained are in good

agreement with the traditional biostratigraphic age esti-

mates for the Cantaure Formation based on the identifi-

cation of the N7-N8 planktonic foraminiferal zones by

Diaz de Gamero (1974) and the nannofossil biozones

NN4-NN5 by Rey (1996). Anderson and Roopnarine

(2005), on the other hand, in their Table 2, place tlie

Cantaure Formation in the Burdigalian-Langhian, strad-

dling tlie Lower-Middle Miocene boundary.

The Cantaure Formation consists of a sedimentary

sequence approximately 75 in thickness and mainly

Figure 2. Geographic location of the study site in tlie Cantaure region, Paragiiana Peninsula, in Venezuela.

ir 56’ 24.1

Page 34

THE NAUTILUS, Vol. 128, No. 2

composed of fossiliferous silts, siltv sandstones, and fine to

medium sandstones interbedded with thin algal lime-

stones (Hunter and Bartok 1974; Lexico Estratigrilfico de

Venezuela, 1997; Aguilera et ah, 2013). A diverse fossil

assemblage, particiilarlv rich in mollusks, !)ut also featur-

ing corals, decapods and cirripedian cnistaceans, and fish

renuiins, has been identified in the sediments of the

Cantaure section, especiiilly in its lower part (e.g. Jung

1965; Nolf and Aguilera 1998; Aguilera and Rodrigues de

Aguilera, 1999; Griffiths et al., 2013). Locally, decimetric

boulders of limestone with in .sifn -attached valves of the

shallow marine bivalve Spomhjhis sp. may be observed

within the friable fine sandstone beds. This fossil assem-

blage is indicative of a shallow to coasted tropical marine

environment, with clear water and marine eiihaline salin-

ity (Jung, 1965; Di'az de Gamero, 1974; Nolf and Aguilera,

1998; Aguilera et al., 2013; Griffiths et ah, 2013).

MATERIALS AND METHODS

The Cliilean mateiicil described herein comes from

the CovaceUcli and Fnissinetti collection, housed in the

Museo Nacional de Historia Natural, Santiago, Chile. The

Venezuelan material described liere is from the Gibson-

Smith collection, housed in the Natiirhistorisches Museum

Basel (NMB), Switzerhuid, and the Bernard Landau

collection, lioused in the Natiirhistorisches Museum

Wien, Austria.

In the .systematic descriptions of the new species, we

have followed the moqihometric model suggested by

Jerardino and Navarro (2008) and MacClintock (1967)

(Figures 3-6). Following the convention established by

MacClintock (1967), the layers are numbered by refer-

ence to tlie myostracurn (m); starting at the outside of

the .shell, they are designated m+3, m+2, m+1, m and

AIL

u

Figures 3-6. Mor|ihoinetric nieasuremeiits of patellogastropod limpet; Scutellastra veneztielana new species. Figures 3-5 adapted

from Jerardino and Navarro (2008, p. 1025, fig. 1 ) TL = Total length; AL = anterior length; PL = posterior length; TW = total width;

LAV = lateral width; AIL = anterior inner length; IL = inner length; PIL = posterior inner length; IW = inner width; LIAV = lateral

inner width. Figure 6 adapted from MacClintock (1967) in = myostracurn.

ventral

dorsal

myostracurn

posterior

anterior

— — — —

TW

PL

S.N. Nielseii and B. Landau, 2014

Page 35

m - 1. Although MacClintock (1967) distingiiislied ni+2

and iri+3 layers in the shells of patellids, Ridgeway et al.

(1998) found that these were not clearly separable. In

both new species the width of the in+1 layer is greater

than half of the width of the combined outer (m + 1,

m+2, in+3) layers (see Figures 3, 5, 7, 10, 12). Tliis

character is seen in tlie genera Scutellastra, Lottia, and

Acmaea (Ridgeway et al, 1998). The relatively large and

solid shells with strong radial dorsal sculpture suggest

placement in the genus Scufellastra.

Abbreviations: SCO. PL, Museo Nacional de Historia

Natural, Santiago, Chile; NHMW, Naturhistorisches

Museum Wien, Austria; NMB, Natiirhistori.sches

Museum Basel, Switzerland.

SYSTEMATIC PALEONTOLOGY

Superfamily Patelloidea Rafinesque, 1815

Family Patellidae Rafinesque, 1815

Genus Scutellastra H. and A. Adams, 1854

Type Species: Patella plicata Born, 1778 (= P.

harhara Linnaeus, 1758) by subsequent designation

of Wenz (1938, see Ridgway et al., 1998); Recent,

South Africa.

Scutellastra arayae new species

(Figures 7-9)

Description: Shell large, tliick, oval, lieaGly ornamented

with very coarse irregular radiiil ribs projecting notalrly at

margin. Seven primary ribs, one secondary rib in each

interspace, one tertiary rib between primaries and

secondaries, additional lesser ribs in all interspaces

and on major ribs. Apex situated anteriorly. Muscle scar

horseshoe-shaped, open anteriorly. Venter with deep, U-

shaped myostracum (m); m-1 about 1/3 totiil width of

shell at level of opening of myostracum; m+1 wide, widtii

about 4/5 in of m-1, in+2 and 3 narrow (numbering

following MacClintock, 1967).

Type Material: Holotype SCO. PI. 6650 from Punta

Perro, height 55 mm, diameter 93 mm (incomplete) x

92.6 mm, Covacevich and Frassinetti locality 241080.1.

Concretionary block with small Pinna seniicostata.

Type Locality: Punta Perro, lower Miocene Navidad

Formation, central Chile.

Other Material Exannined: Known only from holotype.

Distribution: Only known from the type locality.

Etymology: Named after Ivette Araya, paleontology

collections manager at Museo Nacional de Historia Nat-

ural, Santiago.

Measurements: Measurements follow Jerardino and

Navarro (2008). Total length (TL) > 93 mm; anterior

length (AL) ~ 51.3 mm; posterior length (PL) unknown;

total width (TW) 92.6 miii; lateral width (L\\') 50 mm;

anterior inner length (AIL) 19 mm; inner l('ngtli (IL)

60 mm; posterior inner length (PIL) unknown; inner width

(IW) 36.2 inm; lateral inner widtli (LIW') > 26.2 inm.

Discussion: Scutellasfra arayae new- species lias few

coarse projecting ribs and can be easily distingiiislied

from species with a smooth or crenulate margin, includ-

ing S, mexicana. The apex of S. arayae lies anteriorly

while that of S. flexuosa (Quoy and Gaimard, 1834) from

Australia is nearly central. Smtellasira cliaj)inaiii (Tenison-

Woods, 1876) from Australia and S. longicosia (Lamarck,

1819) from South Africa have better defined and more

projecting ribs. Scutellastra harhara (Linnaeus, 1758)

and S. exusta (Reeve, 1854), botli from South Africa liave

more and finer ribs.

According to Beu and Miixwell (1990) two fossil species

of Sadelkistra are knowm from New Zealand, tlie upper

Oligocene-early Miocene S. aurorae Fleming, 1973 and

the early Miocene S. coojjeri (Powell, 1938), wliich would

both be roughly contemporaneous vrith the Chilean spe-

cies. judging from the original figures, S. aurorae has a

rather smooth margin and more and weaker ribs while

S. coirperi has stronger projecting ribs than S. arayae.

Several species of Scutellastra are known to be gar-

deners. Some species garden coralline algal substrate in

their periphery, feeding either on those coralline algae or

on other red algae growing on this substrate while other

species maintain patches of algae on which they graze

(Lindberg, 2007). Patch and periphery gardeners can be

recognized moiphologically since peripheiy gardeners

exliibit an anteriorly extended shell while the shells of

patch gardeners have a rounded anterior end. Both

forms can thus be recognized in the fossil record and

their ecology may be inferred as was done by Lindi^erg

(2007) who figured the tyjiically extended shell form ol a

fossil specimen from Venezuela that was reported as of

Pliocene age but, as mentioned earlier, is the Miocene

species described below. Scutellastra arayae does not

show an anteriorly extended shell and is therefore con-

sidered to belong to the patch-gardening group.

Scutellastra venezuelana new species

(Figures 10-14)

Scutellastra sp. — Lindberg, 2007, p. 230, fig. SB.

Description: Shell moderately large, up to 90 mm in

length, solid, depressed, pear-shaped, witli anterior end

produced, but not constricted at neck. Sculpture of

coarse radial ribs on dorsum, most of which of primary

strength and deeply corrugate the margin. Venter with

deep, U-shaped myostracum (m); m-1 about total wndtli

of shell at level of opening of myostracum; m+1 wide,

about equal in wddth to in-1, in+2 and 3 narrow' (num-

bering following MacClintock, 1967).

Type Material: Holotvpe NHMW 2013/0566/0002,

height 9.7 mm, diameter 34.0 mm x 22.5 mm; paratvpe

1, NHMW 2013/0566/0001, height 16.9 mm, diameter

Page 36

THE NAUTILUS, Vol. 128, No. 2

Figures 7-14. Saitellastra specie.s. 7-9. Scutellastra araijae new species. Holotvpe SGO.PI.665{), height 55 mm, diameter >93 x

92.6 mm. Punta Perro, lower Miocene NaMdad Forination, central Chile. 10-14. Scutellastra venezuelana new species. From 1 km

southwest of Casa Cantaure, about 10 km west of Pueblo Nuevo, Falcon, Venezuela, Cantaure Formation, upper Biirdigalian-lower

Langhian, lower-middle Miocene boundary'. 10-11. Holoty|re NHMW 2013/0566/0002, height 9.7 mm, diameter 34.0 mm x 22.5 mm.

12-14. Paratvpe 1 NHMW 2013/0566/0001. Height 16.9 mm, diameter 60.5 mm x 43.0 mm.

S.N. Nielsen and B. Landan, 2014 l^ige 37

60.5 nnn x 43.0 mni; paratvpe 2, NHMW 201 3/0566/

0003, height 10.6 nnn, diameter 38.5 mm x 27.8 mm;

parahpe 3, KIIMW 2013/0566/0004, lieight 10.5 mm,

diameter 40.3 mm x 27.6 mm; paratvpe 4, NMB

1120223, height 15.9 nnn, diameter 62.0 inm x 42.2 mm,

NMB locality 17516; paratvpe 5, NMB II20224, height

14.7 mm, diameter 59.0 mm x 43.2 mm, NMB local-

ity 17516.

Type Locality': 1 km southwest of Casa Cantaure,

about 10 km west of Pueblo Nuevo, Falcon, X'eneznela,

Cantanre Formation, upper Bnrdigalian-lower Langhian,

lower-middle Miocene !)oundarv.

Other Material Examined: Miiximurn dimensions:

diameter 89.1 mm x 58.7 mm. NHMW 2013/0566/0005

(8); NMB locality 17516 NMB (31 unnumbered speci-

mens), same locality as type material.

Di.strib«tion: Only known from the type locality.

Ety'mology: Named after the country of origin,

\'enezuela. The gender of Scutellastra is feminine.

Measurements: Measurements follow Jerardino and

Navarro (2008) of paratvpe 1. Total length (TL) 60.5 mm;

anterior length (AL) 36.5 mm; posterior length (PL)

29.3 mm; total width (TW) 43.0 mm; later;il vvadth (LW)

26.9 mm; anterior inner length (AIL) 20.3 mni; inner

length (IL) 29.2 mm; posterior inner length (PIL)

12.7 mm; inner w'idth (IW) 16.1 mm; lateral inner width

(liW) 16.8 mm.

Discussion: The rostration developed in Scutellastra

venezuelana new species makes it superficially similar to

the extant Scutellastra cochlear (Born, 1778) from the

coasts of South Africa, but this species differs in having

an even more pronounced rostration, which is somewhat

pinched at tfie neck. Although this tendency to rostration is

most strongly developed in S. cochlear tuid S. venezuelana,

this character is probably convergent as it is developed to

some degree in several other Scutellastra species; i.e. the

eastern Pacific species S. mexicana (Broderip and Sowerby,

1829) and the Indo-Pacific species S. (rfjtima (Pilsbry,

1927). More importantly, S. venezuelana differs from both

S. cochlear and S. mexicana in having coarser dorsal ribs,

almost all of which are of primary strength, whereas both

S. cochlear and S. mexicana have finer ribbing, with sev-

eral orders of major and minor ribs.

Despite the superficial similaritv’ between the shells of

S. cochlear and S. venezuelana, there are no common

factors between the Venezuelan and South African mol-

luscan faunas, and tlie new Venezuelan species is much

more likely to be related to the eastern pacific S. mexicana.

The similarity between these species is likely to be due to

common ecological factors. According to Lindberg (2007),

the tendency to rostration and an angular rather than

rounded profile when viewed from the dorsal aspect are

associated with gardening limpets that maintain the gar-

den around the periphery of the shell. Lindberg (2007)

suggested that the rostration might allow these limpets to

graze their gardens without leaving the vicinity of their

home depressions.

Scutellastra venezuelana lu'w spt'cies is (jiiite unlike

the Chilean S', arai/ae new species, which is less flat-

tened, the primarv ribs are much broader and th(‘ shell

does not develop the pronounced rostration so typical of

S. venezuelana, S. cochlear and S. mexicana.

CONCLUSIONS

In the Recent American shallow marine faunas,

Scutellastra is known only from the extant sp(‘ci('s S.

mexicana occurring from \\'est Mexico to Pern. This

paper introduces two new species from the American

fossil record. Scutellastra araijae new species is the

oldest and the southernmost record from the Americas.

It is also the first patellid recognized from Cenozoic

deposits of Chile. Scutellastra venezuelana new specie's

is only slightly younger, straddling the earlv-middle Mio-

cene boundary', it is the first record for the genus in the

tropical American Neogene and the Caribbean/western

Atlantic indentified to species-level. This new re'cord

adds the genus Scutellastra to the list of Paciphile gen-

era, i.e., genera which, follow'ing the closure of the Cen-

tral American Seaway, disappeared from the Caribbean

and became restricted to the Pacific side of their original

wider distribution. For full list of Paciphile genera see

Landau et al. (2009).

The biogeographic pattern of dispersal of patellid gas-

tropods has been fairly controversial. This is not helped

by the very poor fossil record for the group. “Patella”

sotjaensis Kase and Shigeta, 1996 from the upper Creta-

ceous of northern Japan, assigned to Scutellastra bv

Ridgway et al. (1998) is the oldest record for the genus,

altliough the generic placement was (piestioned by

Koufopanou et al. (1999). Together vCth the records

from New Zealand, these new lower and lower-middle

Miocene American records are among the oldest undis-

puted records for the genus. Koufopanou et al. (1999)

suggested that S. mexicana was a relict from a formerly

widespread Tethyan distribution of early Scutellastra

species (Powell, 1973; Ridgway et al, 1998). They pre-

dicted that further Scutellastra should be discovered in

the Atlantic and Mediterranean regions. These findings

support their hypothesis.

Through comparison vUth the current biogeographic

distribution of Scmtellastra spp., as far as tbe Chilean

record is concerned, it becomes evident that this fossil

species is yet anotlier piece of evidence that ( 1 ) shallow-

water or, as in this case, even intertidal tiixa were

displaced into bathyal depth of the Na\idad Formation

(see Finger, 2013), (2) sea surface temperatures along the

Chilean coast were significantly higher during the early

M iocene than they are today (Nielsen and Glodny, 2009),

and (3) discovery of this new species confirms the results

of Kiel and Nielsen (2010) tliat, although the Navidad

assemblage is relatively well described, there remain

many more species to be discovered. The \enezuelan

Page 38

THE NAUTILUS, Vol. 128, No. 2

record ( I ) illiistrate.s again tlie iinportanee of the Cantaure

as.seinl)lage, as one of the vei-v few tropical Neogene

assenililages representing rocky hottoin liabitats (Yenneij,

2001; Landau et ah, 2009; Landau and \'enneij, 2010),

and (2) adds to the nuniher of species knowii to have been

distril)uted throughout the Neogene Gatunian province,

hut today restricted to tlie Pananiie Pacific.

AC K N OW'LE DG M ENTS

Ivette Arava and tlie late Daniel Erassinetti (Museo

Naeional de Historia Natural, Santiago, Chile) are

thanked for access to the collections under their care.

Thanks to Carlos Marcpies da SiKa of the Universih’

of Lisfxni, Portugal, and Toinovuki Nakano of Nagova

University’, Japan for their help and advice in preparing

this paper.

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THE NAUTILUS 128(2):40-50, 2014

Page 40

The family Caecidae (Gastropoda: Caenogastropoda)

in Argentine waters

Guido Pastorino

Ignacio Luis Chiesa

Museo Argentiiio de Ciencias Naturales “Bernardino Rivada\ia”

A\'. Ange! Gallardo 470

C14()5DJR Cindad Autdnoina de Buenos Aires, ARGENTINA

[email protected]

ABSTRACT

This is the first formal report of ineinhers of the family

Caecidae in Argentine waters. Caecum striattim de Folin,

1868, C. strij^osuiii de F'olin, 1868, and C. achiroua (tie Folin,

1867) are re-described from shallow waters off Piedras

Coloradas (~40°53.()81' S, 65°07.592' W), Rio Negro Proxince,

Argentina. 'Phis is the farthest south record of these species

wliich were previously recorded from USA, Bahamas, Panama,

Brazil, and Uruguay. The authors also make obsenations about

the different ontogenetic stages of the studietl species. Scan-

ning electron microscope illustrations of radula and operculum

are provitled for the first time.

Additional Kf’i/word.s: Argentina, Caecuim Patagonia, taxonomy

INTRODUCTION

The family Caecidae comprises marine caenogastropods

with simple cylindrical (Caecinae) or almost planispiral

(Ctiloceratinae) very small shells, usually around 2-3 mm

whicli in rare cases are larger than 5 mm. The Caecinae

inhabit tropical and temperate environments, mostly in

shallow waters. The early works of Carpenter (1858) and

de Folin (1877) established that at least three different

growth stages are present in representatives of the group.

However, Bandel (1996) reported more complicated

arrangements, which may be imi(|ue for each species.

Probably because of small size, particular ontogeny, and

somewhat conservative shell mor]ihology’, the taxonomy of

this interesting group is far from complete. In addition,

most of the species have been described based solely

on shell characters. However, some earlier workers (e.g.,

Gotze, 1938; Marcus and Marcus, 1963; Draper, 1979;

Bandel, 1984; etc.) described the radiilar moqihology of

some species. Marcus and Marcus (1963) presented draw-

ings of tlie anatomy, operculum, and radulae of what they

identified as C. cometim and C. pulchellum from the

littoral of Sao Paulo, Brazil. The actual identities of these

species are need of revision.

The first descriptions of species of Caeaim from the

southwestern Atlantic are those of de Folin (1868; 1874) as

reported by Klappenbach (1964). Inter, Lange de Morretes

(1954) de.scrihed a new species from Sao Paulo State,

wliich, together with his previous list (1949) increa,sed

the number of species of Caecinae knowm from Brazil.

These former workers are pioneers in the study of this

complex family; however, only in more recent years the

revision of type specimens led to a better understanding

of the identities of those nominal species. Absaliio (1994;

1995; 1997), Gomes and Absalao (1996), and Absalao and

Gomes (2001) made the first attempts, using modem

criteria, to review the family in the southwestern Atlantic.

More recently, Lima et al. (2013) improved on the

traditional format of species descriptions with an ontoge-

netic approach that we attempted to follow here. Lima

et al. (op. cit.) reported more than 30 species living along

Brazilian coast.

In the odier countries of southem South America otlier

than Bi'cizil, recent species of Caecidae have been

described from Cliile (Stiiardo, 1962; 1970; Di Geronimo

et al., 1995) and Uruguay (Klappenbach, 1964; Scarabino,

2004). Faiinati (1994) reported the presence of Caecum

antillanim Carpenter, 1858 from Holocene deposits from

Bahia Blanca, Buenos Aires Province, Argentina. In adtli-

tion, Penchaszadeh (1973) cited the presence of Caecum

sp. as part of the diet of the sea star A.sf rojjecten hrasiliensis

collected off Buenos Aires Province. The latter, as far as

we know, constitutes the only published report of recent

members of the family Caecidae from Argentina.

In this paper we describe, for the first time, three

recent representatives of this intriguing family from

Argentine waters. The study includes SEM illustrations

of the radulae, opercula, and remarks on the ontogeny of

some of these species.

MATERIALS AND METHODS

The material described herein was collected during a

sampling project focused essentially on small peracarid

crustaceans from shallow waters in San Matias Gulf, Ri'o

Negro, Argentina, during January of 2005. The samples

were obtained using a van Veen grab and a Raiischert

sledge, deployed from a small boat in several stations

off Piedras Coloradas (40°53.08r S, 65°07.592' W). The

grab area was 0.05 m“. The sledge opening measured

55 X 15 cm and was ecjuipped with nylon net of 1x1 mm

mesh size. The samples were manually sieved 10 times,

and then the sorted material was fixed with formalin 4%

on sea water, and later preserved in 70% ethanol. Table 1

lists the stations where Caecidae were present, including

die fisliing gear, geo-referenced locality, depth, and sedi-

ment grain size.

Due to small size, radulae were taken dissolving the

whole animal on a hanging drop slide with sodium hypo-

chlorite. Once clean, the radula was moved to another

slide filled with distilled water in which a piece of photo-

graphic film was glued to the bottom of the cavity with

the emulsion side up. Once the water evaporated, the

film was removed and attached to a SEM stub, and

coated with gold-palladium. Shells were cleansed in an

ultrasonic cleaner and observed and photographed

under SEM at the Museo Argentino de Ciencias

Naturales (MACN).

The genus Caecum sensu lato usually develops a

deciduous and spiral protoconch. The protoconch is gen-

erally lost and a septum closes off the first stage of the

teleoconch. This latter could be ornamented with a

structure more or less developed (finger-like, flat,

subquadrate, etc.) called mucro. Sometimes the mucro

pierces the septum and is clearly distinguishable as in

Figures 4-8, or could be less differentiated, as in

Figures 12-15. The teleoconch could develop several

ontogenetic stages, herein referred to, if the protoconch

is present, as stages I, II, III, and so on, or, if the

protoconch is lacking, as stages X, Y, Z, etc. The ontoge-

netic stages of the teleoconch could be still attached, in

which case a fracture line is visible.

The material is housed at the invertebrate collection of

die MACN.

RESULTS

Six of the 21 samples contained several specimens of

three different species of Caecidae in different ontoge-

netic stages. Caecum striatum de Folin, 1868 was the

commonest and the other species, C. strigosum de Folin,

1868 and C. achirona (de Folin, 1967) appear to be rare.

The sediment where this fauna live is mainly sand of

medium and fine grain. They were found between 12-

18 in depth, most of tliein alive and associated witli dif-

ferent species of ainphipods, mainly belonging to species

in the family Phoxocephalidae.

SYSTEMATICS

Family Caecidae Gray, 1850

Subfamily Caednae Gray, 1850

Genus Caecum Fleming, 1813

Caecum striatum de Folin, 1868

Figures 1-25

Caecum striatum (de Fol.) — de Folin, 1868: 49, pi. 5,

fig. 3; Rios, 1994: 56, pi. 18, fig. 207; Gomes and

AbsaJao, 1996: 519, fig. 7; Absalao and Gomes, 2001:

12, figs. 8-9 (lectotype designated).

Caecum striatum, van ohsoleta de Folin, 1874: 212.

Caecum antillanim Car|3enter, 1857. — Rios, 1994: 56,

pi. 18, fig. 203.

Description: Shell: Protoconch unknown. Teleoconch

X (first stage) very small, less tlian 1.5 mm; tubular,

slightly and regularly curved; apical caliber somewhat

larger than apertural; periostracum transliicent-broumish,

tliick, brittle when dry, covered with longitudinal micro-

scopic (but visible under stereoscopic microscope), close-

spaced, continuous, weakly sinuous striae; shallow thin

grooves among striae, faint circular lines (groxMli lines?)

crossing striae and producing wavy ends to those striae.

Septum fiat to slightly convex; mucro thin, finger-shaped,

weakly projected, flat; rising from the interior covered by

septum, sometimes partially broken (Figure 8), positioned

on dorsal margin. Teleoconch Y (second stage) (Figures 1

and 3 show the starting point indicated by a sudden increase

in diameter tliickening) smiiU, about 1.5 mm, moderately

curved; apertural diameter slightly larger than apical; api-

cal region circular, with slight constriction; rounded hemi-

spheric septum, with flat, polygonal dorstil mucro, slightly

twisted to left, sometimes very weak; oblicpie rim always

present between septum and end of striae; septum and

mucro whitish. This was the most abundant stage found.

R,<\i3ula (Figures 17-18): Raehidian tooth somewhat

semicirculai' in outline, with 12-13 short cusps, tlie cen-

tra! larger that lateral cusps; lateral teeth with 12-13 short

Page 42

THE NAUTILUS, Vol. 128, No. 2

Figures 1-9. Teleoconch oi' Caecum striatum de Folin, 1868. 1. MACN-in 39530-1. Teleoconch X and Y, showing the periostraciiin

broken, arrow heads probable facture line between two ontogenetic stages, X and Y. 2. MACN-In 39530-2. Scale bar = 500 pm.

3. Detail of Figure 1 showing the probable fracture line between two ontogenetic stages. Scale bar = 100 pm. 4-8. Five lateral views

of septum and mucro. 4. Detail of specimen in Figure 1. 5-6. MACN-In 39530-3. 7. MACN-In 39530-4. 8. MACN-In 39530-5.

Scale bar = 100 pm. 9. Detail of the ornamentation of the shell of the specimen in Figure 2. Scale bar = 50 pm.

G. Pastorino am! I.L. Chiesa, 2014

Page 43

Figures 10-16. Teleoconch of Caecum striatum de Folin, 1868. 10. MACN-In 39531-1, teleoconch. 11. MACN-In 39531-2,

teleococh, showing the periostracuin broken. Scale bar = 500 iJirt. 12-15. Four different wews of septum and imicro ol different

specimens. 12. Detail of apical extreme of Figure 10. 13. MACN-In 39531-3, Detail of septum and mucro, arrow heads imicro.

14. Detail of the apical extreme of specimen in Figure 11. 15. Lateral view of the apical extreme of specimen in Figure 13. Scale

bar = 100 pm. 16. Detail of the ornamentation of the shell in Figure 15. Scale bar = 20 pm.

cusps, larger than those of rachidian and smaller than

those of inner marginal teeth; inner marginal tooth long,

larger than all otliers, with 12-15 large, shar|3 cusps; at

end of cusps, a deep furrow (f) shows the starting point of

the long tooth stalk; outer marginal long, slender, with 12-

14 cusps smaller than those of inner marginal tooth.

Radulae show similar features at all the growth stages.

OPERCULU.vi (Figures 19-25); Similar in all growth

stages, circular, thick, corneous, external surface slightly

and mainly in the center concave, multispiral, sculptured

with a thick subquadrate cord of 4-5 whorls, separated by

a deep furrow, sometimes partially covered; internal sur-

face convex, attachment area spanning half of totid sur-

face, small central hole present; internal and external

surface closely attached; margin of inner surface reflected

over outer surface and covering its margin.

Material Examined: MACN-In 39535, St. 5; MACN-

In 39533,St. 6; MACN-In 39532, St. 15; MACN-In 39534

St. 19; MACN-In 39536, St. 18; MACN-In 39530/1-5

(illustrated specimens); St. 18; MACN-In 39531/1-3

(illustrated specimens), St. 18; all off Piedras Coloradas,

San Matias Gulf, Ri'o Negro Province, Argentina.

Page 44

THE NAUTILUS, Vol. 128, No. 2

Figures 17-25. Radula and operculum oi' Caecum striatum de Folin, 1868. 17. Dorsal xaew of the radiila, scale bar = 5 pm.

18. Detail ol the rachidian tooth, scale l)ar = 2 pm. 19-25. Operculum. 19. Twisted external \iew. 20. Side \iew. 21. Twisted internal

\iew. 22. Operculum attached, critical point dried. 23. External \iew. 24. Inteniiil \iew. 25. E.xternal \iew with furrows uncovered.

Scale l)ar = 100 pm. A!)l)reviation: f, furrow in the inner inargin;il tooth.

Distribution: Florida, USA; Bahamas; West Indies

(according to Lightfoot, 1992); Panama; Pernambuco

state, Fernando de Noronha Is., Rio de Janeiro, Brazil

(according to Leal, 1991; Absalao and Gomes, 2001 (as

C. stri^osinn)) and Rio Negro, Argentina.

Eemarks: Two ontogenetic stages (X and Y) are attrib-

uted to this species. As no complete or united specimen

was found, the ontogenetic order was arranged accord-

ing to tlie diameter of the aperture and septum area of

each stage and the general mor|3hology of the shell. The

G. Pastorino and I.L. Chiesa, 2014

Page 45

stage Y is tlie usually described form; however, the most

alrundant stage was stage X.

Absalao and Gomes (2001) designated lectotypes of

C. striatum and C. stri^osum and opened the discussion

about the possibility of these two names being synonyms.

We found enough distinction to maintciin the two species

separate until more information is available.

There is a series of errors on the publication dates of

the two species. Previous authors (i.e., Rios, 1985; 1994;

Leal, 1991; Ligthfoot, 1992; Absalao and Gomes, 2001)

considered 1867 as the publication date of C. strigosum.

Rehder (1946) completed the collation of de Folins "Les

Fondes de la Mer” previously published by Winkworth

(1941). According to them, both descriptions, from the

first volume of this work, were published in 1868.

Caecum strigosum de Folin, 1868

Figures 26-34

Caecum strigosum (de Fob) — de Folin, 1868: 53, pi. 5,

fig. 51869; : 261;

Caecum strigosum de Folin, 1867. — Rios, 1985: 44,

fig. 194; 1994: 57, pL 18, fig. 208; Leal, 1991: 86, pi. 13,

figs. H-I; Ligthfoot, 1992: 28, fig. 31; Absalao and

Gomes, 2001: 11, figs. 7, 8.

Description; Protoconch unknown; teleoconch

medium sized, tubular, slightly curved, about 2 mm with

a clear, somewhat oblique, swelling, right at the end of

the aperture; sculptured with longitudinal striae, some-

times obsolete, similar to those described for C. striatum

but shallower, thinner and with more wavy pattern; sep-

tum evenly curved, hemispherical, without rim, protruded;

mucro small, sometimes very weak or obsolete, twisted to

left (Figure 32).

Radula similar to that of C. striatum. Operculum sim-

ilar to C. striatum but the attachment area at the internal

surface is smaller (Figure 34).

Material Examined; MACN-In 39537, St. 4; MACN-

In 39538/1^, St. 18, all off Piedras Coloradas, San Matias

Gulf, Rio Negro Province, Argentina.

Distribution; According to Rios (2009), from Maranhao

to Sao Paulo, Brazil; however, this author considers

C. striatum as a synonym. The distribution of both spe-

cies may overlap.

Remarks; According to Absalao and Gomes (2001)

C. striatum and C. strigosum should be treated as syno-

nyms. No doubts both species are really closer. However,

the presence of the apertural swelling in C. strigosum

together with the hemispherical septum and the almost

obsolete mucro clearly separates die latter species. In

addition, the smaller attachment area of the operculum

of C. striatum adds to the separation of the two species.

However, it still remains to be investigated whether

these differences represent just steps in the ontogeny of

a single species.

Caecum achirona (de Folin, 1867)

Figures 35-49

Brochina achirona de Folin, 1867: 57, pi. 3, fig. L

Caecum achironum de Folin, 1867. — .Ybsalao and

Gomes, 2001: 13, figs. 20, 21 (lectotvpe designation).

Description: Shell (Figures 35-44): Protoconch plani-

spiral with one whorl, transluscent, vitreous, witli several

very weak, ftiint cords on a crinkly surface; transition to

teleoconch I well defined. Teleoconch I and II of similar,

short length, witli a weak increase in diameter; transition

to teleoconch II appears as slight constriction; two other

constrictions are also apparent. Teleoconcli X short, ~l/3

length of teleoconch Y; transition to teleoconcli Y showai

as an increase in diameter; teleoconch Y large, strong.

Septum large, dome- or finger-shaped, tliick, flattened

above, lower part somewhat obli(|ue; mucro not visilde.

Complete shell (X-l-Y) moderately large, about 2.5 mrn in

length, curved, tapering toward tlie end, strong; anterior

diameter hviee as large as posterior one; shell translucent;

aperture drcukir, with sharp lip. Shell surface smooth

covered with fine growth lines only visible under SEM;

periostraciim whitish, transluscent, very thin.

Radula (Figures 45^6): Rachidian tooth small, flat-

tened, outline semicircular, with ~10 very small cusps;

lateral tooth small, visible behind inner marginal, with

about 12 small cusps larger than the rachidian tooth

cusps; inner marginal tooth thick, strong, witli about 6

thick, strong, rectangular cusps, larger than the cusps of

rachidian and marginal teeth, a deep furrow present at

the end of the cusps lateral tooth (f in Figure 43); outer

marginal tooth long, slender, thin, with 10-12 small and

sharp cusps.

Operculum (Figures 47-49): Circular, thick, with the

external surface slightly concave, with a thick spiral cord,

covered; internal surface convex, attachment area

appears to cover the whole surface, a central hole at the

center of the spiral formed by the margin of the spring;

margin of the inner surface is reflexed covering the mar-

gin of the outer surface.

Material examined; MACN-In 39529/1-4, St. 5, off

Piedras Coloradas, San Matias Gulf, Rio Negro Prov-

ince, Argentina.

Distribution: The actual distribution of this species is

hard to know as it is difficult to ascertain the taxonomic

circumscription of the nominal species treated by differ-

ent authors. The species has been apportioned to nortli-

eastern Brazil (Pernambuco and Briiia states) by de Folin

(1867). Lightfoot (1992) reported it from Tobago and

Uruguay; it was however not mentioned liy Scarabino

(2004; Umguay).

Remarks; There are some moiphological differences

between the material of Caeaim achirona described here

and the lectotypes illustrated by Absalao and Gomes

(2001), particularly the “longitudinal microstriation,”

Page 46

THE NAUTILUS, Vol. 128, No. 2

Figures 26-34. Caecum strigosum de Folin, 1868. 26. MACN-In 39538-1, scale bar = 500 |im. 27. Detiiil of the swelling around

the aperture in Figure 26. Scale bar = 100 pm. 28, MACN-In 39538-2. Scale bar = 500 pm. 29. Detail of the swelling around the

aperture in Figure 28. Scale bar = 100 pm. 30. Apical Mew of the septum and mucro of specimen in Figure 31. Scale bar = 100 pm.

31. MACN-ln 39538-3, scale l)ar same in Figure 26. 32. MACN-In 39538-4, apical Mew of septum and mucro. 33. Detail of

the ornamentation of the teleoconch of specimen in Figure 28. Scale bar = 50 pm. 34. Internal and external Mew of the operculum.

Scale bar= 100 pm.

(^. Pastoriiu) and I.L. Cliiesa, 2014

Page 47

Figures 35-44. Caecum achirona (de Folin, 1867). 35. .MACN-In 39529-1, protocoiich, still attached to teleoconch 1 and II.

arrows head probable lacture point. Scale bar = 200 pm. 36. Det;iil of the protoconch of Figure 35, arrow heads the boundarv edge

with teleoconch. Scale bar= 100 pm. 37. MACN-In 39529-2, protoconch. Scale bar same as for F’igure 36. 38-39. .V1ACN-Iti 39529-

3, two Mews of teleoconch X and Y, arrows head the probable fracture point, scale bar = 500 pm. 40-41. Details of the septum

of Figures 38 and 39. Scale bars: 40= 200 pm, 41 = 100 pm. 42. MACN-ln 39529-4, detail of the septum from Figure 43. Scale bar =

50 pm. 43. MACN-In 39529-4, teleoconch x and y, arrow heads probable fracture point. Scale bar = 200 pm. 44. Detail of the

surface of the shell. Scale bar = 100 pm.

Page 48

THE NAUTILUS, Vol. 128, No. 2

Figure.s 45-49. Caecum acliinma (de P’oliii, 1867). Radula and operculuin. 45. Dorsal \iew of tlie radula, scale bar = 20 fiin.

46. Detail of the lateral teeth, scale bar = 5 pm. Abbreviations: f, furrow; im, inner marginal tooth; 1, lateral tooth; om, outer

marginal tooth; r, racbidiaTi tooth. 47-49. Three views of the operculum. 47. External view. 48. Twisted view, still attached.

49. Internal view. Scale bar = 100 pm.

vvliicli, according to these authors, characterizes the spe-

cies. All tlie specimens stndied here are smooth. Tliis

oniamentation appears to l)e a variable cluiracter (F. B.

Lima, in lift.), all other leatnres allocated the material

into C. achiroiia. In addition, the differences with C. someii

de Folin, 1867 are also not clear. Absiiliio and Gomes

(2001) designated lectotvpes of the latter and considered

both as different species. According to the illustrations in

Absalao and Pizzini (2002, pi. 4, figs. 30-32) the shell in

C. sonicri presents an apertural constriction that is absent

in C. achiroiia.

"Farluhiin" maaclianicum Di Ceronimo, Privitera,

and Valdovinos, 1995 from the Pacific entrance of the

Strait of Magellan in about 100 m depth, is vaguely

similar. This latter species is smaller in size, reaching

not more tlian 2 nun of shell length, the septnm is

blunter and the aperture margin is somewhat reflected.

Also, the protoconch appears to be the same diameter

all along the entire whorl, wdiile the Atlantic species is

smaller in the first half. Ganging from the number of

individuals found. Caecum achirona is a locally uncom-

mon species.

DISCUSSION

The study of the family Caecidae from the southvv'estern

Atlantic is far from complete. The particular shell mor-

pholog)' with several ontogenetic stages and variable

ornamentation (Absalao and Pizzini, 2002), small size,

and the stereotyped original illustrations are probably

altogether responsible for this scenario. In addition, most

of tlie papers written so far described only the shell, with

more or less details. Radular characters are usually not

G. Pastorino and I.L. Chiesa, 2014

Page 49

included (!)ut see Marcus ant! Marcus, 1963; Draper,

1979; Bandel, 1984). Even when radular characters are

included tlie rare it is difficult to determine taxonomic

relationships. In the material studied liere, tlie morphol-

og)-’ of the radula of C. striatum and C. sirigosum clearly

differs from that of C. achirona. The presence of a par-

ticular inner marginal witli few, flat, and blunt cusps in

the latter could well justify a separate generic allocation.

However, as the characterization of most of the species is

still based on shell features, the use of radular characters

for generic allocations is still difficult. Absalao and Pizzini

(2002) discussed the artificial siibgeneric arrangement in

the subfamily Caecinae used by other authors. We agree

that the knowledge of the relationships within the fam-

ily is still very incomplete to warrant accurate alloca-

tions of species in subgeiiera or even in genera other

than Caecum.

Judging by their recorded distributions, all three spe-

cies reported here appear to be common in the shallow-

water meiofauna along the Atlantic coast. The area of

San Matias Gulf is part of the southern limits of the

Argentine malacological province, according to different

authors who agree considering the Peninsula Valdes area

as its southernmost boundary.

Members of the family Caecidae have been recorded

from Argentine waters. Some observations, as associated

fauna or as prey, reported in ecological or marine biology

papers, recognized caecids as part of food v/ebs. How-

ever, no formal descriptions had been published so far. A

possible reason could be the larger size of the traditional

mesh used in marine surveys that render this ty|3e of gear

ineffective to collect members of the family.

Arnaud and Poizat (1979) published some remarks on

the ecology of three species of Caecum from the Medi-

terranean Sea. They showed that each species have dif-

ferent requirements of depth and habitat. In that sense,

the hydrodynamic and the size of the sand grain play a

crucial role in the distribution of species. They also

mentioned the vertical migration of these species dur-

ing two seasons: spring and summer with two different

purposes, feeding and reproduction. Both species here

described were collected during the Southern Hemi-

sphere summer (in January). No egg capsules were found

together with the adults; however, new collections in

process particularly designed for tliis group could show

their presence.

ACKNOWLEDGMENTS

Silvio F. B. Lima (Brazil) showed his new approach to

the Caecidae and through the friendly discussion helped

to clarify the identity of the material. Renata dos Santos

Gomes (Brazil), Mauro Pizzini (Italy) and Robert

Moolenbeek (The Netherlands) kindly sent publications

and advice that really helped to finish this manuscript.

Brenda Doti and Daniel Roccatagliata (Argentina)

helped to collect and sort the samples. We acknowledge

funding by the Consejo Nacional de Investigaciones

Cientificas y Tecnicas (CONICET) of Argentina, to

wliich GP belongs as members of the “Carrera (!(‘I

Investigador Cientifico y Tecnico” and IG as a fellow.

LITERATURE CITED

Absalao, R.S. 1994. A new species of the genus Caecum

(Prosobranchia: Me.soga.stropo(la) from Soutlieni Brazil.

Journal of Concholog)’ 35: 137-140.

Absalao, R.S. 1995. Ocorrencia de Caecuui hutoti Jong &

Cooinans no Brasil (Rissoidea; Gastropoda; Mollusea).

Biociencias 3 (1): 207-21 1.

Absalao, R.S. 1997. Caecum eliezeri sp. nov. (Prosobranchia:

Mesogastropoda): A new species from Brazil. The \'eliger

40: 271-273.

Absalao, R.S. and R.S. Gomes. 2001. The species usually

reported in the siibgenus Brochina (Caecum, Caecidae,

Caenogastropoda) from Brazil and some relevant Upe

specimens from western Atlantic. Bolletino Malacologico

37: 9-22.

Absalao, R.S. and M. Pizzini. 2002. Critical amilysis of su!)-

generic taxa of the Subfamily Caecinae (Caenogastropoda:

Caecidae). Archiv fiir Molluskenkiinde 131:167-183.

Arnaud, P. M. and C. Poizat. 1979. Donnees ecologiques snr

des Caecidae (gasteropodes prosoliranches) dii Golfe de

Marseille. Malacologia 18: 319-326.

Bandel, K. 1984. The radulae of Carilibean and other

Mesogastropoda and Neogastropoda. Zoologische

Verhandelingen 214: 1-188.

Bandel, K. 1996. Pliylogeny of the Caecidae (Caenogastropoda).

Mitteilungen aus dem Geologisch-Palaontologischen

Institut der Universitiit Ham!)i!rg 79: 53-1 15.

Carjienter, P.P. 1858. First steps towards a monograph of the

Caecidae, a family of Rostriferous Gasteropoda. Proceed-

ings of the Zoological Society of London 26: 413-444.

Fleming, ]. 1813. Mollinsca. In: Brewster, D. (ed.) The

Eclinbiirgh encyclopaedia: conducted by 13avid Brewster,

with assistance of gentlemen eminent in science and liter-

ature. Vol. 7. Edinburg.

Farinati, E.A. 1994. Micromoluscos (Gastropoda y Bivalvia)

del Holoceno del area de Bahia Blanca, Argentina.

Ameghiniana 31: 30.3-315.

Folin, L. de 1868. Les Ponds de la Mer 1, Baie de Bahia: 48-

51; Rade de Rio de Janeiro: 51-54, pi. 5. Saxy, Paris.

Folin, L. de. 1868-9. Obserx'ations on the septum of the

Caecidae; and some remarks on the sulrject of tlie suppres-

sion of tlie genera Brochina and Strehloceras or Phlehoceras.

Journal of the Linnean Society, Zoologx’ 10: 254—264.

[According to tlie JLS, pages 197-260 were pulilished on

November 26, 1868 and 261-330 on JaniuiA- 8, 1869].

Folin, L. de 1874. Sur les cotes du Bresil. Les Foods de la Mer

2: 210-214, pis. 9-10. Saxy, Paris.

Folin, L. de 1877. Note relatix'e au genre Parastrophia . journal

de Conchyliologie 25: 203-207.

Di Geronimo, I., S. Prixatera, and C. Valdoxinos. 1995

Fartulum magellanicum (Prosobranchia, Caecidae): .A

new species from the Magellanic Proxince. Boletin de la

Sociedad de Biologia de Concepcion 66: 1 1.3-1 IS.

Gomes, R.S. and R.S. Absalao. 1996. Usta conientada e ilustrada

dos Caecidae (MoUusca, Prosobranchia, Mesogastropoda)

da operayao Oceanografica Geomar XII. Rexista Brasileira

de Zoologia 13: 513-.531.

THE NAUTILUS, Vol. 128, No. 2

Page 50

Gotze, E. 1938. Ban und Leben von Caecum glahrum

(Montagu). Zoologi.sclie.s Jahrbucli (Abteiliing Systeinatik

und Okologie) 71: 1-190.

Gray, J. E. 1850. Catalogue of the mollusca in the collection

of the British Museum. Part II, Pteropoda. Newman,

Lomlon, 45 pp.

Klappenbach, M. 1964. La familia Caecidae (Moll. Gastr.) en

aguas uruguayas. Comunicaciones de la Sociedad

Malacologica del Uruguay 1 (6): 145-149.

Lange de .Morretes, F. 1949. Ensaio de Catalogo dos moluscos

do Brasil. Ar(juivos do Miiseu Paranaense 8: 1-216.

l^ange de Morretes, FI 1954. Dois novos moluscos do Brasil.

Ar(|uivos do Museu Paranaense 10: 331-336.

Leal, J.H. 1991. Marine Prosolrranch Gastropods from Oce-

anic Islands off Brazil. W. Backhuys, Oegstgeest, 418 pp.

Lightfoot, J. 1992. Caecidae of the Western Atlantic part 2,

conclusion. Of Sea and Shore 15: 23-31.

Lima, S.F.B., F.N. Santos, and R.S. Absalao. 2013. New species

oi' Caecum (Caenogastropoda: Rissooidea: Caecidae) from

the Atlantic coast of South America (Brazil) with a

description of the protoconch and growrih stages. Zoologi-

cal Science 30: 767-778.

Marcus, E. and Marcus, Ev. D. B. -R. 1963. Mesogastropoden

\'on der Kiiste Sao Paulos. Abhandlungen der

Mathematisch-Natunvissenschaftlichen Klasse. Akademie

der Wissenschaften und der Literatur Mainz 1963,

1: 5-105.

Penchaszadeh, P. 1973. Comportamiento trofico de la estrella

de tmir Astropecten hrasiliensis. Ecologia 1: 45-54.

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tion ol Les Fonds de la Mer. Proceedings of the Malaco-

logical Society of London 27: 74-75.

Rios, E.C. 1985. Seashells of Brazil. Editora da Fundayao

Universidade do Rio Grande, Rio Grande, 328 pp.

Rios, E.C. 1994. Seashells of Brazil. Editora da Fundayao

Universidade do Rio Grande, Rio Grande, 368 pp.

Rios, E. 2009. Compendium of Brazilian Sea Shells. Evangraf,

Rio Grande, RS, 676 pp.

Scarabino, F. 2004. Lista sistematica de los gastropoda marinos

y estuarinos \4\qentes de Uruguay. Comunicaciones de la

Sociedad Malacologica del Uruguay 8 (84-85/86-87):

305-346.

Stuardo, J. 1962. Caecum chilense, nuevo molusco para Chile.

Gayana, Zoologia 5: 3-9.

Stuardo, J. 1970. Sobre los representantes chilenos de la familia

Caecidae (Mollusca: Gastropoda). Boletin de la Sociedad

de Biologia de Concepcion 42: 183-190.

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tion. Proceedings of the Malacological Society of London

24(4): 149-151.

THE NAUTILUS 128(2):51-54, 2014

Page 5 1

A new melanopsid (Gastropoda) species from the middle Miocene

Kiipres Basin (Bosnia and Herzegovina)

Thomas A. Neubauer^

Oleg Mandic

Mathias Harzhaiiser

Geological-Paleontoiogical Department

Natural History Museum Vienna

1010 Vienna, Austria

[email protected]

mathias.liarzliauser@nlini-\\'ien.ac.at

[email protected]

ABSTRACT

Melanopsis fateljensis (Caenogastropoda: Cerithiinior|3lia:

Melanopsidae) is described as a new species frosn the early

Middle Miocene lacustrine deposits of the Kupres Basin. Sim-

ilarities to other co-occurring inelanopsids are discussed. Its

unique inor|)hology, in particular the elongate, stepped spire

with prominent spiral I)ijlges, clearly distinguishes the new

species from all other Melanopsidae known from the Neogene

of Europe.

Additional Keywords: Melanopsis, freshwater gastropod, new

species, Dinaride Lake System, Langhian

INTRODUCTION

Recently, Neubauer et al. (2013a) provided a taxonomic

revision of the molluscan fauna of the Kupres Basin

in Bosnia and Herzegovina, which was only poorly

known by then (Brusina, 1902; Kochansky-Devide and

Sliskovic, 1981; Jurisic-Polsak and Sliskovic, 1988). The

long-lived freshwater lake present in this basin during

the early middle Miocene is part of tlie Dinaride Lake

System, a collective of early to middle Miocene fresh-

water lakes in the Dinaride Mountain Chain (Figure 1;

Krstic et ah, 2003; Harzhauser and Mandic, 2008; De

Leeuw et al, 2012; Mandic et al., 2012). The well-

preserved fauna proved to be highly endemic, with

two genera and five species newly described and 30%

of the fauna endemic to Lake Kupres. Even well-

studied coeval, nearby lakes, like Lake Sinj (Neubauer

et al., 2011) and Lake Gacko (Neubauer et al., 2013b),

showed only a low faunistic affinity, with a miudmuin of

38.9% of shared taxa. Another striking feature was the

high percentage of sculptured morphologies, including

' Corresponding author

teleoconch microsculpture in one species, interpreted

as a reaction to the oversatiiratioii of ciilcium carbonate

in the water under the existing alkaline, hard-water con-

ditions (e.g., West et al, 1991).

Shortly after, another taxonomic work on the ostracod

and gastropod fauna of the region around Kupres was

published by Krstic et al. (2013). This study dealing with

an outcrop located al)oiit 2 km ENE of Fatelj Hill

revealed a different assemblage with species known from

the Sinj, Drnis, and Gacko basins (Figure 1) and none of

those described by Neubauer et al. (2013a). The fauna is

characterized by few species of the genera Gyranhis,

Fossandus, Bania, and “ Pseud amnicold’ (both occurring

species were recombined with Bania by Neubauer et al.,

2013a). Such a faunal composition with pulmonate and

typical pioneer species corresponds to those found in the

Gacko and Sinj basins and points to rather stressed,

ephemeral conditions at the basin margin in this partic-

ular stratigraphic level (Mandic et al., 2009; 2011;

Neubauer et al., 2011; 2013b). These deposits were

classified as “Ottnangian” (middle Burdigalian) by Krstic

et al (2013), but should rather be placed in the early

Langhian as discussed by Neubauer et al. (2013a).

The aim of the present study is to fix a misidentifica-

tion of a melanopsid species by Neubauer et al. (2013a).

Our taxonomic reinvestigation revealed clear differences

to the species with which the taxon was previously iden-

tified in Neubauer et al (2013a) and to all otlier

inelanopsids known from the European Neogene, urging

the description of a new species.

MATERIALS AND METHODS

The material derives from an outcrop at the nortliwestem

slope of a small hill, termed Fatelj, about 4 km S\V of the

town Kupres in Bosnia and Herzego\ina (43°58'17.2" N,

17°14'06.9" E, 1140 m). Tlie section is approximately 3 m

diick and covers three lithological units briefly discussed

Page 52

THE NAUTILUS, Vol. 128, No. 2

Figure 1. Geographical over\iew over the study area with

indication of the main sediinentarv' Irasins harboring paleo-

lakes mentioned in the te.xt (modified after Neuhauer et al.,

2013a).

in Neuhauer et al. (2013a). An age of tlie deposits of 15.5 ±

0.2 Ma (= early Langliian or early Badenian in terms ol

regional Paratethys stages) is suggested by the appearance

of the dreissenid i)iv;ilve Mijtilojysis aletici (Brusina, 1907),

which is a good hiostratigrapliic marker due to the rapid

evolution of these bivalves in the Dinaride L;ike System

(Kochansky-Devide and Sifskovic, 1978, 1981; De Leeuw

et al., 2010; Harzliauser and Mandic, 2010). Eleven sam-

ples were tiiken from units 1 and 3. Samples 090709/4 and

090709/5, contiiining the herein investigated species, were

treated with diluted hydrogen peroxide and washed

through two sieves with 2 mm and 0.5 mm mesh size.

When necessary', specimens were cleaned from sedi-

ments with an ultrasonic device. The material is stored

in the collection of tlie Natural History' Museum of

Vienna, Austria (NHMW 201 1/0138).

SYSTEMATIC PALEONTOLOGY

Class Gastropoda Cuvier, 1795

Subclass Caenogastropoda Cox, 1960

Order Cerithiimoiyiha Golikov and Starobogatov, 1975

Superfamily Cerithioidea Fleming, 1822

Family Melanopsidae H. Adams and A. Adams, 1854

Subfamily Melanopsinae H. Adams and A. Adams, 1854

Genus Meianopsis Ferussac, 1807

Melanopsis fateljemis new species

(Figures 2-11)

Meianopsis sp. — Brusina, 1902: pi. 29, figs 23-26.

Meianopsis mojsisovicsi (Neumayr, 1880) comb. nov. —

Neuhauer et al., 2013a: 137, figs 5E-F, I-K (non

Melanopttjchia Mojsisovicsi Neumayr, 1880).

Diagnosis: Shell conical, with elongate and distinctly

stepped spire, witli strong bulges below the sutures and a

marked shoulder between whorl flank and base; aperture

small, ovoid, with very' small anterior canal.

Description: Shell liigh-conical, slender, elongate,

witli 7-10 whorls; proportions variable, with broader

shells sometimes present. Protoconch bulbous, dome-

sliaped, highly convex; number of whorls unknown;

initial part elevated, not covered by successive whorls;

surface smooth. First few shell whorls form regularly

conical outline; beginning about with fifth whorl, weak

shoulder emerges near upper suture; shoulder increases

in strength incrementally, e\'entually forming a broadly

convex bulge. At transition of whorl flank to base,

marked angle occurs, forming second, weaker l)iilge;

weak and broad concavity' is formed between both

bulges, only visible on last 1-2 whorls. Upper bulge may

have irregular course at upper suture in some specimens.

Last whorl reaching 60-70 % of total height; base

straiglit. Aperture small, slender-ovoid, with anterior

and posterior tip forming acute, almost rectangular

angles; callus weakly expressed, glossy; outer lip shar]3ly

terminated, not reflected; siphonal canal very' short and

narrow, not extended or reflected; fasciole narrow, very

weak. Growth lines prosocyrt to slightly sigmoidal

(because of bulges), usually indistinct; occasionallv and/

or temporarily more prominent (Figures 9-10). Coloring

verv rarely preserved, consisting of thin, vertical to

slightly sigmoidal, moderately-spaced, dark yellow to

orange lines; occasionally they fonn widely-spaced zigzag-

lines on earlier whorls (penultimate w'horl upwards;

see Figure 6).

Type Material; Holotype (Figures 2G3), NHMW

2()ll/0138/()107a, 13.97 mm height x 5.80 mm width;

Paratype 1 (Figures 4-5), NHMW 2011/0138/01071),

11.49 mm height x 5.45 mm width; Paratvpe 2

(Figures 9-10),“ NHMW 2011/0138/0184, 12.41 mm

length X 5.61 mm width.

Additional Measurements; 10.47 mm height x

4.74 mm width (Figure 6); 10.39 mm height x 4.17 mm

width (Figure 7); 14.10 mm height x 5.47 mm width

(Figure 11); 13.72 mm height x 5.41 mm width;

15.01 mm height x 5.67 mm width; 12.75 mm height x

6.17 mm width.

Type Locality: NW slope of Fatelj hill near Kupres,

-Bosnia and Herzegovina.

Stratum Typicum: Lower middle Miocene (= lower

Langliian, lower Badenian).

Material Examined: Several hundred specimens from

debris collection of Unit 3, 20 from siunple 090709/4 and

T.A. Keul)auer et al., 2014

Page 53

Figures 2-11. Melanopsis fateljeiisis Neuhauer new species, from the early middle MioceTie of the Fatelj hill, Kupres Basin,

Bosnia arul Herzego\ina. 2-3. HoloUpe (NHMW 201 l/()13S/()107a). 4-5. Paratvpe 1 (NHMW 201 l/()138/01()7h). 6. Specimen

showing zigzag pattern on penultimate whorl (NHMW 201 1/0138/0185). 7. Specimen with preserved coloration (NHMW 2011/

0138/0186). 8. Protoeonch vdew (NHMW 201 1/0 138/0 107c), 9-10. Paratvpe 2 (NHMW 2011/0138/0184). 11. Elongate specimen

(NHMW 2011/0138/0187). All illustrated .specimens are from sample 090709/7 (debris collection of Unit 3).

a single specimen from sample 090709/5, all from the

type locality.

Ety'mology: The new' species is named after the tvpe

localitv'.

Geographic Distribution: So far onlv known from

the Kupres Basin.

Taxonomic Remarks: This species is based on a mis-

identification fry Neubaiier et a!. (2013a), whom errone-

ously identified the present material as Melan()])sis

inojsisovicsi (Neumayr, 1880) described from the roughly

coeval localitv Dzepi (Bosnia and Herzegovina). A direct

comparist)!! is unfortunately impossible as tlie type mate-

ricil oi M. mx)jsisovicsi has been lost. Other material aviiil-

able from Dzepi and the descriptions and illustrations

of Neumayr (1880) still show the differences quite clear.

M. mojsisovicsi has a much higher last whorl and lacks

the subsutural bulges so distinct for M. fateljensis. The

columellar fold typical for M. mojsmwicsi is absent as

well. The two specimens illustrated in Brusina (1902: pi.

29, figs 23-26) as “Mekitiopsis sp.” correspond fully to the

present species. Melanopsis filifera NeumavT, 1880 from

the early middle Miocene deposits of Divar (Bosnia and

Herzegovina) also differs in a larger last w'horl and a w'eakly

to non-stepped spire. Aside from tliese we are not aware

of any otlier melanopsid species similar to M. fateljensis.

ACKNOWLEDGMENTS

We thank Medina Mandic (Vienna) for assistance in the

field and Alice Schumacher (Natural Histoi-y Museum

Vienna) for support with the photographs. We are

grateful to Vitaliy V. Anistratenko (Schmalhausen

Institute of Zoology of NAS, Ukraine) for his construc-

tive review. The work contributes to the project

“Ereshwater Systems in the Neogene and Qiiaternaty’

of Europe: Gastropod Biodiversity, Provinciality, and

Eaunal Gradients” financed by tlie Austrian Science

Eund (EWE project no. P25365-B25).

Page 54

THE NAUTILUS, Vol. 128, No. 2

IJTERATURE CITED

Baiidel, K. 2000. Speciation among the Melanopsidae

(Caenogastropoda). Special emphasis to the Melanopsidae

of the Fannoiiian Lake at Pontian time (Late Miocene)

and tlie Pleistocene and Recent of Jordan. Mitteilungen

aiis dem Geologisch-Paliiontologischen Institut der

Universitiit Hamburg 84: 1.31-208.

Bnisina, S. 1902. Iconographia Molluscomm Fossilium in

tellure tertiaria Hungariae, Croatiae, Slavoniae, Dalinatiae,

Bosniae, HerzegoUnae, Serbiae and Bulgariae inventoriiin.

Officina Soc. Ty|Wgraphicae, Agram, 30 plates.

De Leeuw, A., O. Mandic, A. VranjkoMc, D. Pavelic, M.

Harzhauser, W. Krijgsman, and K.F. Kuiper. 2010. Chro-

nolog)' and integrated stratigraphy of the Miocene Sinj

Basin (Dinaride Lake System, Croatia). Palaeogeography,

Palaeoclimatology, Palaeoecology 292: 155-167.

De Leeuw, A., O. Mandic, W. Krijgsman, K. Kuiper, and

H. Hr\’ato\ic. 2012. Paleoinagnetic and geochronologic

constriiints on the geodyiiamic evolution of the Central

Dinarides. Tectonophysics 530-531: 286-298.

Harzhauser, M., T. Kowalke, and O. Maiulic. 2002. Late Mio-

cene (Pannonian) Gastropods of Lake Pannon with Special

Emphasis on Early Ontogenetic Development. Annalen

des Naturhistorischen Museums in Wien 103A: 7.5-141.

Harzhauser, M. and O. Mandic, O. 2008. Neogene lake .systems

of Central and South-Eastern Europe: Faunal diversity’,

gradients and interrelations. Palaeogeography, Palaeocli-

matology, Palaeoecology 260: 417-434.

Harzhauser, M. and O. Mandic. 2010. Neogene dreissenids in

Central Europe: evolutionary shifts and diversity changes.

In: Van der Velde, G., S. Rajagopal, and A. Bij de Vaate

(eds.) The Zebra Mussel in Europe. Backhuvs Publishers,

Leiden/Margraf Publishers, Weikersheim, pp. 11-29.

]uri.sic-Polsak, Z. arid T. Sli.skoMc. 1988. Slatkovotlni gastropod!

neogenskih naslaga jugozapadne Bosne. Zlrornik referata

naucnog skupa "Minerali, stijene, izumrli i zi\i sMjet

BIH”, Zemaljski Muzej Bosne i HercegoMne, Sarajevo,

7.-8. Oktobar 1988: 167-174.

Kochanskw-Dexide, V. and T. Sli.skoUc, T. 1978. Miocenske

kongerije Hn’atske, Bosne i Hercegovine. Palaeontologia

jugoslavica 19: 1-98.

Kochansky-Dexide, V. and T. Sliskoxic. 1981. Mlade miocenske

kongerije 1 ix'anjskog, Dux'anjskog i Kupreskog polja u

jugozapadnoj Bosni i Hodova u Hercegovini. Palaeontologia

jugoslavica 25: 1-25.

Krstic, N., L. Saxac, G. Jox'anoxdc, and E. Bodor. 2003. Lower

Miocene lakes of the Balkan Land. Acta Geologica

Hungarica 46: 291-299.

Krstic, N., G. Jovanoxic and L. Saxac. 2013. Jezerski ostrakodi i

prateci mekiisci iz kupre.skog polja, donji deo dinaridskog

sistema jezera (Otnang) na xdsini od 1,150 m. Zapisnici

Sqrskog Geoloskog Drustx'a 2011: 1-25.

Mandic, O., D. Pax'elic, M. Harzhauser, ]. Zupanic, D.

Reischenbaciier, R.F. Sachsenhofer, N. Tadej, and A.

Vranjkoxic. 2009. Depositional history of the Miocene

Lake Sinj (Dinaride l.ake System, Groatia): a long-lived

liard-water lake in a pull-apait tectonic setting, journal of

Paleolimnology 41: 431-452.

■Mandic, O., A. De Leeuxv, B. Vukox-ic, W. Krijgsman, M.

Harzhauser, and K.F. Kuiper. 2011. Palaeoenvironmental

ex'olution of Dike Gacko (Soiitliem Bosnia and Herzegoxina):

Impact of the Middle Miocene Climatic Optimum on tlie

Dinaride Lake System. Palaeogeography, Palaeoclimatology,

Palaeoecologx’ 299: 475—492.

Mandic, O., A. De Leeuw, j. Bulic, K.F. Kuiper, W. Krijgsman,

aTid Z. Jurisic-Polsak. 2012. Paleogeographic evolution

of the Southern Pannonian Basin: 4()Ar/39Ar age con-

straints on the Miocene continental series of northern

Croatia. International journal of Earth Sciences 101:

1033-1046.

Neubauer, T.A., O. Mandic, and M. Harzhauser. 2011.

Middle Miocene Freshwater Mollusks from Lake Sinj

(Dinaride Lake System, SE Croatia; Langhian). Archiv

f’iir Molluskenkunde 140: 201-237.

Neubauer, T.A., O. Mandic, M. Harzhauser, and H. Hrvatox’ic.

2013a. A new Miocene lacustrine mollusc fauna of the

Dinaride Lake System and its p;ilaeobiogeographic, palaeo-

ecologic, and taxonomic implications. Palaeontology 56:

129-156.

Neubauer, T.A., O. Mandic, and M. Harzhauser. 2013b.

The Middle Miocene freshwater mollusk fauna of Lake

Gacko (SE Bosnia and Herzegoxina): taxonomic rexisioii

ami paleoenxironmental analysis. Fossil Record 16(1):

77-96.

Neumayr, M. 1880. V. Tertiiire Binnenmollusken aus Bosnien

und der Hercegoxina. Jalirbucli der kaiserlichen und

kdniglichen geologischen Reichsanstalt 30(2): 463-486.

West, K., A. Cohen and M. Baron. 1991. Moqrhology and

behaxior of cral)s and gastropods from Lake Tanganyika,

Africa: implications for lacustrine predator-prey coevolu-

tion. Evolution 45: 589-607.

THE NAUTILUS 128(2):55-58, 2014

Page 55

Attenuiconus marileeae, a new species of cone

(Gastropoda: Conidae: Puncticulinae) from Curasao

M.G. Harasewych

Department of Invertebrate Zoology

National Museum of Natural History

Smithsonian Institution

P.O. Box 37012

Washington, DC 20013-7012 USA

H a ras e wy ch @ s i . e d u

ABSTRACT

Attenuiconus marileeae new species is described from deep

reefs off southeastern Curasao. It resembles A. attenuatus,

A. honkeri, and A. aureonimbosns in size and general proportion

of the shell, but is readily distinguished on die basis of its distinc-

tive color pattern, which consists of a \a\ad orange-red base color

with three bands of irregular, white flainmules. Attenuiconus

marileeae was collected at substantially greater depths than any

of its Caribbean congeners. Only A. atireonirnljosus, from the

nortlieastem Gulf of Mexico, inhabits comparable deptlis. Like

all species of Attenuiconus, nearly all specimens A. marileeae

have one or more major repaired breiiks indicative of unsuccess-

ful attacks by crustaceans.

Additional Keywords: Deep Reef, Ciirasub, bottles, predation

INTRODUCTION

Over the past several years, sampling off soutliem Curasao

and adjacent islands using the manned submersible

CuRASUB as part of tlie Deep Reef Observation Project

(DROP), a collaboration between the Smithsonian and

Substation Curasao, has greatly enriched our knowledge

of tlie deep-reef faunas of tlie region and led to the discov-

ery of range extensions and new species in multiple phyla.

Among the many mollusks collected is a new species

of Attenuiconus , a genus of conids endemic to the tropi-

cal western Atlantic (Petuch, 2013; Tucker and Tenorio,

2013). Several specimens were among the hundreds of

molluscan shells found in multiple glass bottles recov-

ered from the ocean floor at depths ranging from 130-

168 m. The age of the bottles ranged from inid-19*''

century to modern. Although nearly all tliese specimens

were dead collected, they nevertheless provide insights

into the molluscan biodiversity of the region, and include

multiple range extensions and several new taxa. Many of

the shells had at least one drill hole, and were likely

brought into the bottles as food by small octopuses.

This new species is described and compared to

Attenuiconus attenuatus (Reeve, 1844), the ty|ie species

of Attenuiconus , a wide ranging species that occurs in

soutlieastem Florida and tliroiighout tlie Caribbean,

including Curasao, as well as to specimens of Sandeiiconus

sanderi (Wils and Moolenbeek, 1979), the type spe-

cies of Sandeiiconus, which was also present in the bot-

tle samples.

SYSTEMATICS

Family Conidae Fleming, 1822

Subfamily Puncticulinae Tucker and Tenorio, 2009

Genus Attenuiconus Petuch, 2013

Attenuiconus Petuch, 2013: 212-213. Type species: Conus

attenuatus Reeve, 1844, by original designation.

Diagnosis: “Shell small to average size for sulifamily,

very elongated, with straight sides and narrow, straight

apertures; spires low or flattened, with projecting,

mammiiate protoconchs of 2 or 3 whorls; spire whorls

may be flattened, or slightly canaliculated; shells gener-

ally smooth and polished, but some species have coarse

sculpture of fine spiral threads; shells generally colored

in yellows or oranges arranged in wide bands, but may be

colored pink, salmon, reddish-orange witli brown or

white longitudinal flainmules.” (Petuch, 2013: 212-213).

Remarks: In addition to the tv'pe species, which

ranges from southern Florida throughout the Caribbean,

Petuch (2013: 213) included within Attenuiconus: A.

eversoni (Petuch, 1987) from Honduras, A. honkeri

(Petuch, 1988) from Venezuela, A. ianotus (Cargile,

1998), from Honduras, Nicaragua and Colombia, as well

as A. paulosi (Petuch, 1988) from Venezuela and Colom-

bia. He noted that these species had previously been

included in Dauciconus Cotton, 1945 by Tucker and

Tenorio (2009), but that Attenuiconus may be distin-

guished from Dauciconus on the basis of its much

Page 56

THE NAUTILUS, Vo!. 128, No, 2

narrower and more elongate sliell, and by its projecting

protoconcli. Tucker and Tenorio (2013) included A.

(nireonimbosus (Petuch, 1987), a species from the west

coast of Florida, in Aiteiiuicoruts, but transferred A.

i^uofus to Kclh/coiius Petuch, 2013.

AttenuiconuH marileeae new species

(Figures 12-18)

De.seription: Shell (Figures 12-16) of moderate size

for genus (to 23 mm), with solid, narrow (lAV ^ 2.1),

straight-sided, conical, low-conical spire, projecting

protoconch, and narrow aperture. Protoconch (Fig-

ures 17-18) tall, conical, increasing in diameter from 291

pm to 850 pm in IVA evenly rounded, pitted glassv whorls.

Protoconch forms a broad, smootli varix prior to transition

to teleoconch (Figures 17, 18, an'ows), marked by devel-

opment of strongly tuherculate shoulder (17 ttihercles on

first teleoconch whorl, tubercles becoming weaker in suh-

se(|uent whorls, absent by 5*'' whorl). Teleoconch with up

to 8 shaiplv shouldered, stniight-sided whorls. Suture

adpressed in early whorls, mav become shallowlv

impressed in later whorls. Sutural ramp narrow, weakly

concave to flat, with 4-6 rounded cords between suture

and shoulder. Last whorl smooth except for 5-6 broad,

rounded spiral cords near anterior margin of shell. Aper-

ture long, narrow (17\V~1 1) with parallel sides, tleflected

from shell iLxis hv 11-14°. Shell base color golden orange

to orange red, with tliree hands of irregular wiiite mark-

ings: one at and below' the shoulder, one at mid-w'horl,

and one near the anterior margin of the shell. Band below'

shoulder broadest, consisting of ven' irregular, \’aguelv

sigmoidal w'hite flammules, w'hich mav he di\ided. White

flammules e.xtend over shoulder onto sutural ramp, hut

rarely reach suture. Wliite blotches in relatively narrow'

hand at mid-w'hoii range from small and compact (Fig-

ure 8) to large and amoqdious (Figure 10), while

flammules neiU' anterior margin tend to form diffuse,

ohli(|ue lines. Aperture color w'liite. Radula, operculum,

and periostracum unknow'n.

Tjpe Material: Holotvpe, USNM 1195478. Paratvpes

1—4, USNM 1240622, all from the ty]oe locality. Paratvpe

5, Petuch collection, also from the type localih'.

Type Locality: Off the Sea Arjuarium, Bapor Kihra,

Willemstad, Curayao, 12°04.48' N, 68°53.75' W, in

glass bottles collected at 130-168 m, using the Cuiusub

submersible.

Distribution and Habitat: This new species is pres-

ently know'll only from off the southeastern coast of

Curayao, at depths of 130-168 m. Nearly all specimens

have broken lips as well as one or more major repaired

breaks, the latter indicative of prior, severe hut unsuc-

cessful attacks by crustaceans.

Etymology: This new species is named in honor of

Marilee McNeilus in recognition of her longstanding

interest in mollusks and her support of research. She

participated in the submersible dives and assisted w'ith

the specimen sorting that led to the discovery of this

new' species.

Comparative Remarks: Attermiconus marileeae

resemliles A. atlenaatus, a wide-ranging species that

occurs in southem Curayao at shallower depths (Fig-

ures 1-3), in the size and proportions of the shell, hut

differs in having a more concave spire profile and more

prominent spiral sculpture between the shoulder and

suture. Atiemnconus marileeae is most readily distin-

guished from all its congeners by its bright orange-red

color and the large and distinctive patterns of w'hite

flammules that occur in three hands. The color patterns

in A. alieutiahts (Figures 1-5) and A. honkeri (Figures 6-

7) usually take the form of fairly w'ell-defined, parallel-

sided hands of color. Both A. eversoni (Figures 10-11)

and A. pouhm are easily distinguished from A. marileeae

by their flatter spires and more pointed early whorls, as

w'ell as hv their more uniformlv salmon-colored shells.

Attenuicomis aareonimhosus (Figures 8-9) is similar to

A. marileeae in shell shape and proportion. It shares a

similar pattern of irregular, nebulous white flammules,

hut is much paler in color. However, this species and A.

eversoni tend to have a series of very fine spiral brow'll

hands of spots, most evident between the shoulder and

mid-w'horl that are not present in A. marileeae.

Attemncxmus marileeae is easily separated from

Sandericomis sanderi, a species of similar size and base

color that also occurred among the specimens collected

from the bottles. Sandericomis sanderi (Figures 14-18)

has a broader shell with a flatter, more concave spire, a

sharper shoulder, and a broad whitish band with nebu-

lous margins at mid-w'horl.

DISCUSSION

Conus, one of tlie original Linnean genera (Linnaeus,

1758), had, until recently, been considered to he the

most species-rich modern marine genus, with more than

500 extant and several hundred extinct species (e.g.,

Bcickel, Korn and Kohn, 1995; Duda, Kohn and

Palumhi, 2001). Subsequent studies, which included

information on radular moqihologv and molecular data,

have partitioned the 743 cone species know'll at the time

among tliree families, fix’e subfamilies, and 1 14 living

genera, an arrangement that is more in line with those

of several other to.xoglossan families (Tucker and

Tenorio, 2013: 3). These authors note that the numbers

are expected to increase, and the relationships among

the taxa at all levels will continue to be refined as more

data become available.

Within this new paradigm of conoidean svstematics,

Attenuicomis represents a small, pooiiv know'ii genus

limited to the western Atlantic, with greatest diversity’ in

the southwestern Caribbean. The majority' of species

burrow' in sandy bottoms at depths ranging from 10 to

50 m, although some of the SW Caribbean species and

liave been reported to live on sponge reefs (Petuch,

M. II ;irase\wc!i, 2014

Page 57

Figures 1-23. Species of Attemdconus and Sandericonu.s. 1-5. Attemdconus attenuatus (Reeve, 1844), tvpe species of Attenuicomis

Petuch, 2013, 1. Apertural, 2. Dorsal, and 3. Apical vaews of USNM 876325, off southeast shore of Curasao, in 30-37 ni, on sandy

bottom. 4. Apertural and 5. Dorstd views of USNM 806476, off Dania, Florida, in 20 in, on sandy Irottoin, 6-7. Atleniiicorius Jtonketi

(Petuch, 1988). 6. Apertural, and 7. Dorsal views of the holotvpe, USNM 859946, off the Los Monges Islands, N'enezuela, in .35 in.

8-9. Attenuicomis aureonunlxmis (Petuch, 1987). 8. Apertural, and 9. Dorsal views of the holotvpe, USNM 859812, 50 km south of

Apalachicola, FL, in 150 m. 10-11. Attenuicomis eversoni (Petuch, 1987). 10. Apertural and 11. Dorsal views of the holotvpe, USNM

859878, off south coast of Utila Island, Biiliia Islands, Honduras, among live Agr/n'cw corals in 20 m. 12-18. Attenuicomis marileeae new

species. 12. Apertural, 13. Dorsal and 14. Apiciil views of the holotvpe, USNM 1195478. 15. Dorsal view of Paratvpe 1. 16. Dorsalvievv

of Paratvpe 2. 17. Lateral and 18. Dorsal views of protoconch of Paratvpe 3. All tvpe specimens from bottles collected in 130-168 m,

using the Curasub submersible, off the Sea Acjuarium, Bapor Kibra, Willemstad, Curasao. 19-23. Sandericomis sandeti (Wils and

Moolenbeek, 1979). 19. Apertural, 20. Lateriil, 21. Dorsal, and 22. Apical views of USNM 1240614, Off Marie Pampoen, Willemstad,

Curasao, in 297 m, on sandy bottom. 23. Apertural view of voucher specimen from the same localitv that was the source of COI barcode

sequence deposited in GenBank KJ751548. 2 cm scvile bar applies to all shells, 1 mm scale bar applies to scanning electron micrographs

of the protoconch. Arrows indicate transition from protocouch to teleoconch.

Page 58

THE NAUTILUS, Vol, 128, No. 2

2013: 213). Only A. mireonimhosus from the northea.st-

eni Gulf of Mexico wa.s reported from depths as great as

70-150 m. The depth at which A. marileeae has been

collected (130-168 in) is significantly greater than the

batlivmetric range for most Attenuiconus , but similar to

tliat of A. aureoniinbostis. As all available specimens of

A. nmrileeae were dead collected, it is possible that this

species inhabits somewhat shallower waters, and that the

shells may have rolled downslope and become occupied

l)y hermit crabs prior to being brought into the bottles.

However, other species of cones collected from tlie same

iiottles [(i.e., Sandericonus sanderi (Wils and Moolenbeek,

1979), Conasprelloides villepinii (P. Fischer and Bemardi,

1857), and Dalliamus inazei (Deshayes, 1874)] are all

known to inhabit the depths at which the bottles were

collected (Rosenberg, 2009).

Living specimens of Sandericonus sanderi (Fig-

ures 19-23) were collected nearby, but at substantially

greater depths (297 m).

ACKNOWLEDGMENTS

Tliis research was conducted while participating in the

Smithsonian Institution's Deep Reef Obserx-ation Pro-

ject (DROP) operating at the Curasao Seacjuarium.

Support from this program is gratefully acknowledged.

Thanks also to Adriaan Schrier and the staff and crew of

Substation Curasao and the Sea<|uarium for their gra-

cious hospitality and support. Special thanks to Marilee

McNeilus and Cristina Castillo for assistance in the field

and with sorting specimens, and to Yolanda Villacampa

and Yesha Shrestha for their contributions in the labora-

tory. Drs. Edward |. Petuch and |ohn Tucker pro-

vided helpful comments on drafts of this manuscript.

This is Occiin Heritage Foundation/Curacao Sea AcjiuiriunV

Substation Curacao (OHF/CSA/SC) Contribution Num-

ber 5.

LITERATURE CITED

Cargile, W.P. 1998. Description of Comi.s- ignotus, a new spe-

cies from Nicaragua. Siratiis 2(14): 9-14.

Deshayes, G.P. 1874. Description d’lin cone nouveau des Antil-

les. Journal de Conchyliologie 22: 62-66.

Duda, T.F. ]r.. A.]. Kohn, and S.R. Paliimbi, 2001. Origins of

diverse feeding ecologies within Conus, a genus of venom-

ous marine gastropods. Biological journal of the Linnean

Society 73: 391-409.

Fischer, P. and A.C. Bernardi. 1857. Descriptions d’especies

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secundum classes, ordines, genera, species, cum characteribus,

tlifferentiis, synontyiiis, locis . . . Edicio decima reformata.

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[iv] -h 823 pp.

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Education and Research Foundation, Charlottesville,

[v] 4- 154 4- A1-A4, 29 pis.

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mollusks. Coastal Education and Research Foundation.

Charlottesville, [vi] -f 217, 39 pis.

Petuch, E.j. 2013. Biogeography and Bicxliversity of Western Atlan-

tic Mollusks. Boca Raton, Florida, CRC Press, xvii 4- 2.34 pp.

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Atlantic Marine Mollusca. [WWW database (version 4.1.1)]

URL http:/Av'vwv. inalacolog.org/.

Tucker, j.K. and M.j. Tenorio, 2009. Systematic classification

of Recent and fossil conoidean gastropods. Hackenheim,

Conchbooks, 296 pp.

Tucker, J. K. and M. J. Tenorio, 2013. Illustrated Catalog of

the Living Cone Shells. Wellington, FL, MdM Publishing,

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Bijdragen tot de Dierkunde 49(2):255-260.

THE NAUTILUS 128(2):59-63, 2014

Page 59

First American record of the exotic slog Tandonia kusceri

(Gastropoda: Milacidae)

Joclieii Gerber

Collections Center — Invertel)rates

Field Museum of Natural Histor>'

1400 South Lake Shore Drive

Chicago, IL 60605-2496 USA

jgerber@riekliniiseiiin.()rg

ABSTRACT

The terrestrial slug Tandonia kusceri (Pulmonata: Milacidae) is

native to the Balkan Peninsula. This article reports the discov-

ery of the species for the first time outside southeastern

Europe, in Brookfield, Illinois, USA, a suburb of Chicago.

Descriptions and photographs of live animals and reproductive

organs are proMded to facilitate the recognition of this poten-

tial pest species preciously unrecorded from the Americas.

Tandonia kusceri is compared with three other Milacidae spe-

cies that have been introduced by humans into regions out-

side their native range, two of which have been recorded

from North America. The known distribution and ecology of

T. kusceri are summarized.

Additional Keijivords: Introduced species, genital anatomy

INTRODUCTION

Terrestrial slugs have a long history of being introduced

to the Americas from other continents. At least 26 spe-

cies have been reported as having established popula-

tions in the USA and Canada (Turgeon et ah, 1998;

Reise et ah, 2000, 2006; Grimm et ah, 2009). Introduced

slug species can pose threats to the environment, e.g., by

competing with native species (RoIIo, 1983), and they

can become important agricultural pests (Me Donnell

et ah, 2009 and references therein). Consequently, con-

siderable amounts of manpower and money are spent by

governments (e.g., U.S. Department of Agriculture

[USDA]) to prevent additional species from entering

North America, to limit the spread, and, if possible,

eradicate limited occurrences of newly introduced spe-

cies. Nevertheless, ever-increasing international trade

has the unfortunate side effect that additional alien slug

species (along with other mollusks and other animals and

plants) continue to be introduced to America (Robinson,

1999; Robinson and Slapcinsky, 2005). Efforts to prevent

new introductions and to limit the spread of alien spe-

cies, as well as attempts to discover the avenues of their

introduction, are more likely to be successful if intro-

duced species are documented as early as possible (Reise

et ah, 2000; 2006; Robinson and Slapcinsky, 2005).

In this article, the first American record of tlie alien

slug Tandonia kusceri (H. Wagner, 1931) is reported.

Photographs and descriptions of the animal and its gen-

ital anatomy are provided as identification tools.

MATERIALS AND METHODS

Voucher material has been deposited in the Mollusk

Collection of the Field Museum of Natural History,

Chicago (FMNH) and the USDA National Mollusk

Collection, Philadelphia (USDA):

USA, Illinois, Cook County, Brookfield, near the inter-

section of Jefferson and Harrison Avenues, 41°49'42" N,

87°51'23" W, 190 m a.s.h, crawling on concrete porch

after rain, at about 23:00h, 21 June 2013, leg. M.K.

Thayer (FMNH 328572: 8 specimens preserved in 70%

ethanol -b tissue samples of 5 of these specimens in

95% ethanol; USDA 140056: 2 specimens preseiwed in

70% ethanol).

Same locality, but found in front yard, buried 10 cm

deep in garden soil; 5 October 2013, leg. M.K. Thayer

(FMNH 328573: 1 specimen in 70% etlianol; FMNH

328574: 1 dried specimen [mummified after escape from

container in which specimens were kept]).

IDENTIFICATION

The descriptions (Figures 1^) given here are based on

the Brookfield specimens of T. kusceri.

External Morphology; Fully extended, mature spec-

imens ca. 7 cm long, rather slender. Length of mantle

shield in the front half of the body about % total body

lengtli. Breathing pore at about of the mantle length

from the anterior mantle margin, on the right side of the

mantle shield. Mantle shield with a horseshoe-shaped

groove open posteriorly. Surface of mantle shield gran-

ular. A keel extends along the middle of the dorsum.

Page 60

THE NAUTILUS, Vol. 128, No. 2

Figures i-4. Tandotiia kusceri from Bolingbrook, Illinois, USA (F’MNH 328572). 1. Extended specimen, length about 6.5 cm.

2. Contracted specimen. 3. Animal with sole exposed. 4. Distal portion of the reproductive tract. Abbrexiations: ag, accessory gland;

at, genital atrium; be, bursa copulatrix; bd, duct of bursa copulatrix; dep, distal part of epipluillus; dp, distal part of penis;

er, epiphallus retractor; ov, oxiduct; pep, proximal part of epiphallus; pp, proximal part of penis; pr, penis retractor; vd, vas deferens.

J. Gerber, 2014

Page 61

from tlie tail end to the posterior edge of the mantle

shield. Back and sides with about 15 diagonal rows

of flattened tubercles on each side. When resting, spec-

imens can contract strongly antero-posteriorly, so

as to attain nearly a half-circle shape when seen from

tlie side.

Overall color light yellowish- to pinkish-brown, becom-

ing lighter toward the sole. Black pigment concentrated

in the furrows separating the skin tubercles, which

results in a reticulated pattern. Keel lighter-colored

than back, without black pigment. Mantle diffusely

speckled with dark pigment. A dark band present on

either side along the branches of the horseshoe groove.

Another, somewhat less distinct longitudinal pigment

band in the center of the mantle shield. Head and

tentacles brownish-grey. Sole tripartite, uniformly pale

yellowish, without dark pigment. Mucus of body and

sole colorless, transparent, slightly milky when ani-

mal irritated.

Genitalia: Ovotestis in the dissected specimens (n=2)

large, suggesting sexual maturity. Hermaphroditic duct

long and thin. Albumen gland large (again, suggesting

sexual maturity), elongated and bent. Spermoviduct wide

and twisted. Vas deferens thin and almost i.5 times as

long as penis and epiphallus combined, opening apically

into the conically attenuated proximal end of the

epiphallus. Epiphallus tubular, wider distally than proxi-

mally, very long, about five times as long as the penis,

intensely coiled and twisted. A broad retractor muscle

inserting on the epiphallus about 'A of its length from its

proximal end. Boundary between epiphallus and penis

marked by the insertion of a thin second retractor mus-

cle. Penis with two distinct sections: a tubular proximal

part about as wide as the distal epiphallus or hardly

wider, and a distal part that is short and globular, about

twice as wide as the proximal part.

Oviduct tubular, straight or but lightly bent. Bursa

copulatrix large, spherical. Bursa duct thick, its diameter

slightly more than '/r of the bursa diameter, and about

twice as long as the bursa.

Accessory glands are two crinkled lobes, one roundish,

the other elongated, that are attached to the vagina at the

transition to the oviduct and bursa duct. The glands are

of a beige color (as opposed to the white surrounding

reproductive organs). Vagina and atrium short.

Epiphallus and bursa copulatrix with its duct were

examined for the presence of a spermatophore but none

was observed.

Distributioo: The native distribution area of T. kusceri

lies in the Balkans. It stretches from Central Serbia

through FYR Macedonia, northeastern Greece, and

Bulgaria to European Turkey and Southeast Romania

(Dedov and Mitev, 2011; Reischiitz, 1988; Wiktor, 1987;

2001). Occurrences of the species in Croatia, coastal

Southwest Ukraine, and Crimea are presumed to be the

result of human introductions (Son, 2010; Sysoev and

Schileyko, 2009; Wiktor, 1987; 1996).

Ecology: According to Wiktor (1987), T. kusceri is a

“species of veiy liigli ecological tolerance*, occurring in

biotopes of various liumiditv, most often found in places

of large quantity of loo.se stones, under which it shel-

ters. (Jeeurring in shrubs, woods, stone debris, and

.synantbropically. Usually in large numbers, especially on

limestone, and in biotopes heavily destroyed by man

(wasteliUid, dumps, ruins).” Recorded elevations range

from sea level (e.g., Varna, Bulgaria and Odessa, Uknune;

Wiktor, 1983; 1987; Son, 2010) to 1450 in a.s.l. (Osogovo

Mountiiins, FYR Macedonia; Dedov and Mitev, 2011).

Welter-Schultes (2012) states that T. kusceri occurs

“in Bulgaria in up to 2000 m.” However, none of the

references he gives contains siicli an elevation record

and it is unclear on what this statement is based.

According to observations at the B()ling!)r()()k site,

T. kusceri is only active at night. On 5 October 2013 two

specimens were discovered during the day, buried 10 cm

deep in garden soil. One of the specimens collected in

October 2013 was held captive until late March 2014.

The specimen was strictly nocturnal in its habits. During

the day it stayed buried in the soil covering tlie bottom of

the container rather than using items like pieces of tree

bark that were offered for shelter.

DISCUSSION

Several species in the Milacidae, namely Milax gfigaies

(Draparnaud, 1801), Tandonia budapestensis (Hazay,

1881) and Tandonia soiverhiji (A. Ferussac, 1823), are

known as invasives in areas far beyond their native south-

ern European range. Milax gagates has been introtliiced

around the world including North and South America

(Wiktor, 1987; Turgeon et al., 1998; Grimm et al., 2009;

McDonnell et al., 2009) and T. budapestensis has lieen

recorded in the eastern United States (Reise et al.,

2006). Consequently, it seemed likely that the milacids

from Brookfield belonged to one of tliese tramp species.

However, the attempt to assign the specimens to one of

these species based on external characters failed. Dissec-

tions clearly showed that the Brookfield specimens

were not conspecific with any of these invasives, but

that they instead belonged to Tandonia kusceri, a species

that had not been found previously outside of south-

eastern Europe.

The combination of an extremely long vas deferens

and epiphallus, the latter being intensely coiled and

twisted, a short, bipartite penis, and a large, bulbous

bursa with a thick tubular duct is unicpie among the

Milacidae and allows for easy recognition of T. kusceri.

Figure 4 shows the distal parts of the reproductive tract

of a specimen from Bolingbrook (FMNH 328572). It

resembles Wiktors (1987: 258-259, fig. 155) description

and drawing of the genital anatomy of T. kusceri very

closely. The only differences are: W'iktor shows a con-

striction at the insertion of the penis retractor, i.e., at

the epiphallus-penis boundary'. No such constriction is

discernible in the Bolingbrook specimen. Secondly, the

Page 62

THE NAUTILUS, Vol. 128, No. 2

broad retractor inserting on the epiphallus about 'A of

its length from its proximal end is neither mentioned

nor figured by Wiktor. However this feature is depicted

in two of the drawings of the genitalia of T. kusceri (as

Milax [M. ] kusceri) by Grossii (1983: 223, fig. 139; 124,

fig. 140).

Externally, the Bolingl)rook specimens agree w4th the

description of T. kusceri given by Wiktor (1987). Wiktor

states that extended specimens are up to 10 cm long. The

specimen figured by him (1987: 258, fig. 154) measures

about 6.5 cm.

Otlier milacid species recorded from North America

differ as follows (Wiktor, 1987):

Milax ^agates - Vas deferens short; epiphallus club-

shaped, proximally truncated, short; penis irregularly

rounded, short, almost half the length of the epiphallus;

bursa copulatrix elongated, its duct very short and indis-

tinct; accessory glands open into the atrium (not the

\'agina) through numerous tubules (characteristic for

genus Milax); atrial stimulator present (cliaracteristic for

genus Milax). Body tends to be uniformly dark grey or

blackish, without pigment spots.

Tandonia hudapestensis - Vas deferens short;

epiphallus short, as long as or slightly longer than penis,

cylindrical or club-shaped; penis irregularly rounded;

bursa copulatrix oval to cylindrical, its duct thick and

short. Body appearing variably blackish-broum due to

dense black spotting on a dull cream or orange back-

ground; keel olive or orange, without black pigment.

Wlien resting, animals often curl into a c-shape, as

opposed to other milacids which contract into a “hump”

(Kerney and Cameron, 1979; Reise et ah, 2006).

Tandonia sowerhtji has been ntroduced into large

parts of southern and western Europe as well as South

America and New Zealand, but has not been recorded

from North America. It can be similar to T. kusceri in its

external appearance (Wiktor, 2001) but differs clearly in

its genital anatomy (Wiktor, 1987; 2001): Vas deferens

long, narrow, twisted, about three times as long as the

epiphallus (Wiktor, 1987: 301 states “twice as long” but

fig. 215 on p. 302 shows the vas deferens to be clearly at

least three times as long as the epiphallus); epiphallus a

thick cylinder tapering and slightly bending toward the

vas deferens; a strong retractor muscle inserting later-

ally on the epiphallus about % of its length from its

proximal end; two additional muscles, inteqireted as

retentors, inserting laterally (pointing away from each

other) near the distal end of the epiphallus, close to a

constriction marking tlie epiphallus-penis boundary;

penis more or less cylindrical, about as wide as epiphallus,

its length less than half to ca. of the epiphallus length;

bursa copulatrix strongly elongated, tubular in younger

specimens, distally wider and clearly delimited from

duct in older specimens; bursa duct narrower and shorter

than bursa.

At this point it is not clear if the T. kusceri population

in Brookfield is the result of a recent introduction and

restricted to the immediate area around the collection

site, or if the species has been there for a number of

years and occupies a wider area. We do not know how

this exotic species was introduced. The fact that the

discoverer, Dr. Margaret K. Thayer, who has lived at this

address for the last 25 years and who, as an experienced

zoologist, is trained to take notice of unusual animal

occurrences, does not remember seeing this slug in pre-

vious years seems to point to a recent arrival on the

Thayer property. However, it is currently impossible to

state wdth certainty whether the slugs have recently

migrated from nearby yards and gardens where they

may have dwelled for some time and how and when they

arrived there. I intend to monitor the Tandonia popula-

tion in Bolingbrook and learn more about the extent of

the infestation in the coming months.

The apparently verv wide ecological amplitude of

T. kusceri and its synanthropic tendencies (Wiktor,

1987) suggest that the species could be successful in

l)ecoming established in North America.

AC K N O WLE D G M E NTS

Thanks are due to Dr. Margaret K. Thayer (FMNH-

Insects) for bringing the slugs to my attention, for

collecting specimens and for sharing her slug-related

observations witli me.

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