Fusceulima and Halielloides ( Gastropoda : Eulimidae ) in the southwestern Atlantic , with descriptions of two new species of Fusceulima

Fusceulima Laseron, 1955 and Halielloides Bouchet & Waren, 1986 are reported from the southwestern Atlantic for the first time. Halielloides ingolfiana Bouchet & Waren, 1986 is considered a valid name, and the taxon is distinct from its previous senior synonym Eulima verrilliana Bush, 1909 in shell shape and absence of an umbilicus. Fusceulima saturata sp. nov. is characterized by a conical shell, domed at the apex, a short ovoid aperture, and a distinct dark-brown spiral band near the suture. Fusceulima toffee sp. nov. also has a conical shell and a dome-shaped apex, but has a taller high aperture and the entire shell is dark brown. Halielloides cf. ingolfiana, Fusceulima cf. boscheineni, and Fusceulima cf. minuta, are here recorded from the continental shelf off southeastern Brazil. The nominal species of these taxa were originally described from the northeastern Atlantic. Their status remains dubious because of lack of information on their biology, anatomy and molecular data. Regarding shell morphology, we cannot distinguish the Brazilian specimens from species in the northeastern Atlantic with the material available. A series of short grooves close to the suture of the protoconch in all species of Fusceulima, including the type species Fusceulima jacksonensis Laseron, 1955, is a possible diagnostic character for this genus.

Eulimidae Philippi, 1853 consists of a large and diverse group of marine microgastropods, with approximately 1000 valid species (BOUCHET 2014), most of which are parasites of echinoderms (WARÉN 1984).The Eulimidae is one of the five most species-rich families of marine gastropods in the Indo-Pacific (BOUCHET et. al. 2002, ALBANO et al. 2011).This diversity is not related exclusively to the number of extant species, but also to the wide range of parasitic strategies (TAKANO & KANO 2014).Despite these large numbers and differences in life habits, the morphology of the shell differs mainly in the general shape: some genera are very distinct in the latter and are easily recognizable, whereas others have very similar morphology and their limits are difficult to identify.The scarcity of morphological characters on the shell (e.g., axial ribs and spiral cords) poses a challenge for taxonomists (DGEBUADZE et al. 2012).
Recent oceanographic campaigns off southeastern Brazil and collection-based studies have revealed large numbers of unknown eulimids.A taxonomic review of the Eulimidae (in progress) using the material from the Campos Basin, southeastern Brazil, has led to the discovery of several species, including species of Fusceulima Laseron, 1955 andHalielloides Bouchet &Warén, 1986, which are the topic of this paper.
Members of Fusceulima occur in all oceans (THIELE 1925, LASERON 1955, BOUCHET & WARÉN 1986), but in the Atlantic, records of this genus are limited to the north -northeastern region.Fusceulima currently comprises 18 species (BOUCHET & GOFAS 2014).The distribution of Halielloides is limited to the North Atlantic (BOUCHET & WARÉN 1986), with only two valid species.Halielloides nitida (Verrill, 1884) is the only member of the genus known from the western Atlantic, on the USA coast, based on a synonymy (WARÉN 1991).No records from Fusceulima and Halielloides have been reported from the southwestern Atlantic.
In this study, we report for the first time the presence of species of Fusceulima and Halielloides in the southwestern Atlantic, based on empty shells collected in the Campos Basin,[40][41].

MATERIAL AND METHODS
The taxonomic identifications were based on conchological comparisons with type material and/or original descriptions and illustrations.Almost all species treated here were illustrated using scanning electron microscope (SEM) images.
The terminology used for conchological features follows BOUCHET & WARÉN (1986).The method used to count the number of whorls of the protoconch is in accordance with LEAL (1991).
Most of the material examined is from two projects: (HAB) Projeto Habitats -"Heterogeneidade Ambiental da Bacia
Diagnosis.Short, cylindrical shell, convex whorls, a strongly arched but weakly opisthocline outer lip.An ovate aperture and reflected inner lip with a distinct umbilical fissure (based on BOUCHET & WARÉN 1986).
Halielloides cf.ingolfiana Bouchet & Warén, 1986 Figs 2-10 Characterization.Vitreous shell, cylindrical with a broad, obtuse apex, reaching 2.8 mm in length.Larval shell about 1.7 whorls, diameter about 400 µm; smooth.Teleoconch with up to six whorls and a rounded outline, strongly convex; suture deep and distinct, sinuous; subsutural zone distinct, narrow, occupying about 0.2 height of each whorl; surface sculptured by indistinct growth lines; incremental scars strong, well demarcated, appearing irregularly in number (4-6) and intervals (0.4-1.2 whorls).Last whorl comprising 0.4 total shell length; base elongated with rounded outline.Aperture high, elongated, ovoid, slightly pointed posteriorly with a rounded anterior margin; outer lip thin, with a sinuous profile, deeply sinuate below suture, evenly curved downwards, projecting most in the lower half; inner lip slightly projected into parietal region; columella straight.Small umbilical fissure.
Type locality of Halielloides ingolfiana.Off Iceland, Verrill, 1884 = Eulima verrilliana Bush, 1909) as a very similar species that differs in size (larger) and in having a more oblique aperture and weaker incremental scars.BOUCHET & WARÉN (1986) also mentioned the possibility of these two species becoming synonyms, and later, WARÉN (1991), based on additional material from other localities in the northeastern Atlantic, concluded that there is more variation in specimens of H. ingolfiana than in the material observed by BOUCHET & WARÉN (1986), which led him to confirm H. ingolfiana as a synonym of H. nitida.However, examination of the holotype of Eulima verrilliana (Fig. 11) revealed a different shell, with a more-conical shape and less-convex whorl profile.In addition, E. verrilliana lacks the umbilical fissure, as stated by VERRILL (1884) in the original description.The umbilical fissure is a diagnos- verrilliana is broken and does not permit a comparison.Thus, we consider H. ingolfiana as a valid name, and also consider that the generic assignment of Eulima verriliana requires further investigation.
Shells from southeastern Brazil (Figs 2-10) are very similar in shape and sculpturing (growth lines) to the holotype of H. ingolfiana, with strongly convex whorls and an easily recognizable umbilical fissure (Fig. 8).However, the mean diameter of the protoconch is smaller (narrower) in shells from Brazil, approximately 400 µm, and the protoconch reaches about 1.7 whorls.Shells of H. ingolfiana have a protoconch with approximately 1.2 whorls and a diameter of 530 µm.Concerning the color, the Brazilian specimens are also vitreous or whitish, but some have a slightly yellow tint (Fig. 5).
Considering H. ingolfiana as a valid name, there are no records of Halielloides from the western Atlantic.Halielloides cf.ingolfiana is quite distinct from any other named western Atlantic eulimid; the extremely close similarity in the shell morphology of Brazilian specimens with the shells of H. ingolfiana from the northeastern Atlantic prevents us from discriminating them.However, since we have no data to support the amphi-Atlantic distributions of this species, and given the limited bathymetric distribution (restricted to the continental shelf) of the material from southeastern Brazil, we hesitate to identify the material examined here as H. ingolfiana.A more reliable identification will depend on examination of soft parts and material from the continental slope off Brazil.
Fusceulima cf.boscheineni Engl, 1998 Figs 22-28 Characterization.Shell conical with a wide spire angle, and a broad, obtuse apex, reaching 1.5 mm in length.Larval shell approximately 1.5 whorls, diameter about 240 µm; smooth, with short, fine striations below suture, from the nucleus to the strongly recurved terminal scar at the transition to teleoconch.Teleoconch with up to three whorls of slightly sinuous outline, with a very slight convexity on the lower region; suture shallow, rectilinear; subsutural zone distinct, occupying about 0.2 height of each whorl; surface glossy and smooth; incremental scars weak, approximately 4, with    Remarks.Although the holotype of F. boscheineni is a larger shell, reaching six whorls, with a somewhat elongate shape, while all shells from Brazil are shorter with the maximum of five whorls, some paratypes of F. boscheineni have shorter shells, similar to the shells from Brazil.Fusceulima projectilabrum Bouchet & Warén, 1986 from the northeastern Atlantic, including also records from Greenland (WARÉN 1989), has a similar laterally expanded aperture and an umbilical chink, but differs in the almost cylindrical shape and the colorless shell.
Since we have no data to evaluate the possibility of a wide range of distribution in the Atlantic Ocean for species of Eulimidae, we refrain from describing a new species.Fusceulima boscheineni is limited to the continental shelf of the Canary Islands while Fusceulima cf.boscheineni is restricted to the continental shelf of the Campos Basin.
Fusceulima cf.minuta (Jeffreys, 1884) Figs [31][32][33][34][35][36] Characterization.Shell conical to subcylindrical, with obtuse apex, reaching 1.8 mm in length.Larval shell with 1.8 whorls, diameter about 330 µm; smooth, with short, fine striations below suture, from the nucleus to the strongly recurved terminal scar at the transition to teleoconch.Teleoconch with up to three whorls with sinuous outline, and a convexity on the lower region; suture deep, rectilinear; subsutural zone distinct, occupying about 0.35 height of each whorl; surface glossy and smooth; incremental scars strongly demarcated, about five in irregular intervals of 0.3-1.1 whorls.Last whorl about 0.5 total shell length; base elongated with slightly rounded outline.Aperture ovoid, pointed posteriorly with a rounded anterior margin; outer lip thin, of sinuous profile, deeply sinuate below suture, evenly curved downward, with the most projecting part at the middle; inner lip slightly projected into pari-etal region; columella straight.Imperforate.Shell vitreous, occasionally with brownish spots.
Type material of Fusceulima minuta.Lectotype NHMUK 1885.11.5.2064;Paralectotypes NHMUK 1885.11.5.2064, NHMUK 19790227 Remarks.The lectotype of F. minuta (Fig. 29) is slightly more conical, but the paralectotypes (Fig. 30) have a more cylindrical shell shape, similar to the shells from Brazil.In addition, the irregular light-brown spots of the paralectotypes are also found on shells from Brazil.The material available to us was composed exclusively of empty shells, and for this reason we cannot distinguish the specimens from Brazil from the eastern Atlantic shells of F. minuta.Fusceulima cf.minuta is limited to the continental shelf of the Campos Basin, and we have no records of it in greater depths such as in the northeastern Atlantic, where F. minuta reaches 662 m.However, the uncertainty about the possibility of amphi-Atlantism and the limited bathymetric distribution of the material examined here lead us to refrain from identifying Fusceulima minuta in Brazil.BOUCHET & WARÉN (1986) discriminated Fusceulima minuta  from two other species of Fusceulima from the northeastern Atlantic (F.thalassae Bouchet &Warén, 1986 andF. lineata (Monterosato, 1869); F. minuta is also different from F. projectilabrum in the expanded outer lip and the presence of an umbilical chink in the latter species.Fusceulima ingolfiana Bouchet & Warén, 1986 differs in the strong convexity of the whorls.Fusceulima sordida (Watson, 1897) is quite similar to the paralectotypes of F. minuta in its cylindrical shape, but the whorls are almost perfectly flat and according to the description of WATSON (1897), F. sordida possesses a minute chink behind the inner lip that is not present in F. minuta.Color of fresh shells brownish, with whitish or transparent apex; dark-brown spiral band at subsutural zone; beach-worn shells usually white.Dimensions.Holotype with six whorls; length 2.6 mm; width 0.9 mm; height of aperture 0.8; width of aperture 0.5 mm.Remarks.Fusceulima saturata sp.nov.resembles Fusceulima minuta (Figs 29-30) in the general shape of the shell, but F. saturata is taller, more conical, while F. minuta has a conical to subcylindrical shell.Furthermore, F. saturata has a shallowly demarcated suture.Fusceulima saturata differs from F. projectilabrum and F. boscheineni  principally in the absence of an umbilicus.Fusceulima saturata differs markedly from Fusceulima ingolfiana in the strong convexity of the whorls in the latter species.In addition, the color pattern of Fusceulima saturata differs from all the other Atlantic species.The general shape and color pattern of this species is more similar to Fusceulima goodingi Warén, 1981, described from New Zealand waters (WARÉN 1981), but the shells studied by us are even taller and F. goodingi possesses whorls that are almost cylindrical in contrast to the slightly sinuous outline of the whorls in F. saturata.
Description.Shell conical with a broad, obtuse apex, reaching 2.9 mm in length.Larval shell about 1.8 whorls, diameter about 350 µm; smooth, with short, fine striations below suture, from the nucleus to the strongly recurved terminal scar at the transition to teleoconch.Teleoconch with up to 3.2 whorls of sinuous outline, with a convexity on the lower region; suture more impressed, rectilinear; subsutural zone distinct, occupying about 0.15 height of each whorl; surface glossy and smooth, weak incremental scars, about six with irregular intervals of 0.3-0.9whorls.Last whorl with about 0.6 total shell length; base elongated with slightly rounded outline.Aperture high, ovoid, acute posteriorly with a rounded anterior margin; outer lip thin, of sinuous profile, sinuate below suture, evenly curved downward, with the most projecting part at the upper half; inner lip slightly projected into parietal region; columella straight.Imperforate.Color of fresh shells brownish, with whitish or transparent apex; false suture distinguished by a dark brown color; beach-worn shells usually cream or yellowish.
Remarks.Fusceulima toffee sp.nov.has a slightly sinuous whorl outline, similar to F. saturata sp.nov., but the elongate aperture (Fig. 50) clearly discriminates this species from the others that have a shorter aperture.In addition, the evenly curved outer lip has a different tortuosity from the common shape, not curving backward immediately below the suture, differing from all other species of Fusceulima.The color of F. toffee is more similar with the color of Fusceulima fulva (Watson, 1897), but F. fulva does not show the above-mentioned features of F. toffee.

DISCUSSION
Fusceulima and Halielloides are somewhat similar in shell morphology, especially in the dome-shaped apex and the ovoid aperture.Species of Halielloides have more-convex whorls, but Fusceulima ingolfiana Bouchet & Warén, 1986 is an exception, with very convex whorls, resembling Halielloides Bouchet & Warén, 1986.According to BOUCHET & WARÉN (1986) the cylindrical outline of the shell and the presence of an umbilicus are some of the diagnostic characters of Halielloides.However, some species of Fusceulima also have these features; for example, F. projectilabrum has an almost cylindrical shape, and F. boscheineni (and also F. cf.boscheineni, as discussed above) possesses a developed umbilicus.A small fissure is also present in Fusceulima sordida (Watson, 1897), an eastern Atlantic species, as stated by WATSON (1897) in the original description, and in F. projectilabrum (BOUCHET & WARÉN 1986).However, in all of the above cases, such a fissure characterizes an umbilicus similar to that present in F. boscheineni.
The four species of Fusceulima treated here have small and short grooves, mainly on the subsutural region of the protoconch whorls (34)(35)(36)(43)(44)(45)(51)(52)(53), especially visible close to the base of the protoconch and to the terminal scar, where this ornamentation stops, and is absent from the teleoconch whorls.This feature was not examined in the past in other eulimids and perhaps renders the classification of these species in the same genus more robust, even with different shapes and with the presence of an umbilical fissure in F. boscheineni.Among the species studied by us, this characteristic is especially visible in F. saturata , while in F. jacksonensis, the type species of the genus, some very tiny grooves can be seen (indicated by white arrows in Fig. 19), in spite of the eroded state of the shells.
Few species in other eulimid genera possess this feature (pers.obs.), and possibly this is not an exclusive feature of the genus.The possibility that the grooves are merely signs of growth of the shell in the protoconch region or are associated with the type of larval development, in this case lecitotrophy for all the species of Fusceulima studied by us, requires further investigation, but Halielloides cf.ingolfiana does not have grooves (Figs 9-10), nor do many other eulimids.
Thus, we consider that the umbilicus is a variable characteristic in Fusceulima and the short axial grooves on the protoconch are a potential diagnostic character.Their occurrence in other Fusceulima species requires further examination.ABSALÃO (2010) reported "Halielloides (?) sp." from Campos Basin, expressing uncertainty as to the generic assignment.This record was based on broken specimens dredged from 800 m depth (IBUFRJ 15086, IBUFRJ 17209) and cannot be evaluated here, as the shells are damaged and does not allow a reliable determination.Nevertheless, the presence of Halielloides cf.ingolfiana in the material examined in this study confirms that Halielloides occurs in the South Atlantic.Fusceulima, on the other hand, has wider geographic range, with most species recorded from the Indo-Pacific, Mediterranean and North Atlantic (THIELE 1925, LASERON 1955, BOUCHET & WARÉN 1986).
Our choice to report Halielloides cf.ingolfiana, Fusceulima cf.boscheineni and Fusceulima cf.minuta without a definitive determination (expressed by the use of "cf.") is based on the absence of evidence to support that a population from the western Atlantic continental shelf is somehow connected to those on the northeastern Atlantic continental shelf.Additionally, the lack of a clear morphological feature that could distinguish the shells from Brazil from the shells from the northeastern Atlantic belonging to the above-mentioned species led us to refrain from describing this material as new species.PIMENTA et al. (2009PIMENTA et al. ( , 2011) ) advocated that in the absence of supporting data to justify the existence of amphi-Atlantism in species of Pyramidellidae, this possibility should be viewed with caution, avoiding the introduction of new names or proposal of wide distributional ranges without proper foundation.Pyramidellidae also includes cases of shells with identical morphology occurring in shallow localities (continental shelf) of the eastern and western Atlantic.
In the case of Eulimidae, a similar approach should be employed.There is little available knowledge about the duration of the larval phase in this family.However, the type of protoconch present in the species studied by us indicates that it is short.The species of Halielloides and Fusceulima treated here do not have more than two protoconch whorls, and the whorls are more than 200 µm in diameter, suggesting lecithotrophic development (WARÉN 1984).Rectilabrum lanceolatum Bouchet & Warén, 1986 is one of the few species with inferred planktotrophy and anatomical studies that confirm an amphi-Atlantic distribution, with records from the northeastern Atlantic and Virginia, USA (BOUCHET & WARÉN 1986, 1994, ROSENBERG 2009); planktotrophy is not a predominant characteristic in deep-sea eulimids from the North Atlantic (BOUCHET & WARÉN 1994).In addition, there is no evidence of rafting that would guarantee continuity of such distant populations.Although other intrinsic factors can affect dispersal ability (PAULAY & MEYER 2006), paucispiral protoconchs are not related to wide distributions, and thus the taxa treated here as "cf." possibly have a short planktonic period or lack of this period, which reduces their dispersal capacity.
Thus, the question of taxonomic treatment remains: such taxa are either considered conspecific under the application of the strict morphological species concept, in this case, based on the available shell morphology data, or they are deemed cryptic but distinct species when it comes to shell-morphology.An accurate taxonomic study including the morphology, biology and genetics of species provisionally identified here is necessary to elucidate the taxonomy of these specific cases, or even a possible complex of sibling species.
.7-1.0 whorls.Last whorl about 0.6 total shell length; base elongated with rounded outline.Aperture ovoid, pointed posteriorly, laterally expanded, with a rounded anterior margin; outer lip thin, with sinuous profile, deeply sinu-ate below suture, evenly curved downward, with the most projecting part in the lower half; inner lip slightly projected into parietal region; columella straight.Small, chink-like umbilicus.Fresh shells with brownish/gold spots.

Figs 38- 45
Figs 38-45Melanella sp.1:Absalão & Pimenta, 2005: 36, fig.98.    Diagnosis.Broad shell with dark brown spiral band at subsutural zone; strong grooves at subsutural region of protoconch.Description.Shell conical with a broad, obtuse apex, reaching 2.7 mm in length.Larval shell with 1.75 whorls, diameter about 220 µm; smooth, with short, fine striations below suture, from the nucleus to the strongly recurved terminal scar at the transition to teleoconch.Teleoconch with up to four whorls of slightly sinuous outline, with a slight convex-

Fusceulima and Halielloides (Gastropoda: Eulimidae) in the southwestern Atlantic, with descriptions of two new species of Fusceulima Leonardo Santos de Souza & Alexandre Dias Pimenta
Bouchet & Warén, 1986955 and Halielloides Bouchet &Warén, 1986are reported from the southwestern Atlantic for the first time.Halielloides ingolfianaBouchet & Warén, 1986is considered a valid name, and the taxon is distinct from its previous senior synonym Eulima verrilliana Bush, 1909 in shell shape and absence of an umbilicus.Fusceulima saturata sp.nov. is characterized by a conical shell, domed at the apex, a short ovoid aperture, and a distinct Laseron, 1955-brownspiralband near the suture.Fusceulima toffee sp.nov.also has a conical shell and a dome-shaped apex, but has a taller high aperture and the entire shell is dark brown.Halielloides cf.ingolfiana, Fusceulima cf.boscheineni, and Fusceulima cf.minuta, are here recorded from the continental shelf off southeastern Brazil.The nominal species of these taxa were originally described from the northeastern Atlantic.Their status remains dubious because of lack of information on their biology, anatomy and molecular data.Regarding shell morphology, we cannot distinguish the Brazilian specimens from species in the northeastern Atlantic with the material available.A series of short grooves close to the suture of the protoconch in all species of Fusceulima, including the type species Fusceulima jacksonensisLaseron, 1955, is a possible diagnostic character for this genus.KEY WORDS.Eulimids; Eulimoidea; Halielloides ingolfiana; taxonomy; West Atlantic.