Fig. 16. The hokkaidoconchid gastropod Ascheria salina sp. nov. from early Oligocene seep deposits at Cerro La Salina (block 1, B, C; block 2, A) Talara Basin, northern Peru. A. Paratype (NRM Mo187059) showing fine spiral sculpture on early whorls (A2, close-up of the apical part). B. Paratype (NRM Mo187060) large fragmentary specimen. C. Holotype (NRM Mo187061) showing opisthocline axial ornament and faint spirals on last whorl (C1–C3, different lateral views). All in lateral view.

Remarks.—Ascheria salina sp. nov. differs from A. elenen­sis by the shape of the axial ornament, which is strongly opisthocline just below the suture in A. elenensis, whereas the ribs become opisthocline only in the lower half of the whorl in A. salina. Furthermore, A. elenensis grows to a much larger size and appears to be higher spired (having a narrower apical angle) and A. salina has a higher whorl profile than A. elenensis. The Late Jurassic type species A. gigantea differs from A. salina by having spiral ornament also on large whorls, whereas in A. salina such spirals are only present on very early whorls (Kaim et al. 2014). Ascheria? sp. from Barbados (including the specimens reported as “cerithid sp.” by Kugler et al. (1984: pl. 7: 6–8), and as Abyssochrysos sp. by Gill et al. (2005: fig. 5D) has more oblique or sloping axial ribs and spiral ornament is also present on large specimens, in contrast to A. salina. As Ascheria? sp. from Barbados is most likely of Eocene age (Kiel and Hansen 2015), the early Oligocene Peruvian Ascheria salina is currently the youngest member of the genus; it is also the smallest species of Ascheria.

Stratigraphic and geographic range.—Type locality and horizon only.

Family Buccinidae Rafinesque, 1815

Genus Colus Röding, 1798

Type species: Murex islandicus Mohr, 1786, by subsequent designation; Recent, North Atlantic Ocean.

Colus sekiuensis Kiel and Goedert, 2007

Fig. 17A, B.

2007 Colus sekiuensis sp. nov.; Kiel and Goedert 2007: 43, figs. 3D, E.

Material.—One specimen (NRM Mo187062; H = 21 mm) from Cerro La Salina block 2, one specimen (NRM Mo187063; H = 7 mm) from Cerro La Salina block 4. Both upper Oligocene, Talara Basin, Peru.

Remarks.—The specimens are rather imperfectly preserved; with their evenly convex whorl profile and spiral sculpture consisting of 11 low, nearly flat-topped, beaded spiral cords, the specimens are identical to those of Colus sekiuensis known from whale and wood falls from early Oligocene strata in Washington state, USA (Kiel and Goedert 2007). Cyrtochetus? chiraensis Olsson, 1931 from the Chira Forma­tion in the Talara Basin, northern Peru, which is stratigraphically well below the Heath shale and now considered to be late Eocene in age (Higley 2004), is remarkably similar. That species differs from Colus sekiuensis by having finer and more numerous spiral cords with a more convex profile compared to the nearly flat-topped spiral of Colus sekiuensis. In general, the buccinid genus Cyrtochetus differs from Colus by having a varix at the aperture and denticles on the inner lip (Garvie 2013); however, those features cannot be recognized in the available material.

Olsson (1931) reported two species of Siphonalia? from Belén and Pajarabobo, S.? belenensis and S.? tessaria. Both differ from Colus sekiuensis by their more angular whorl profile. At least judging from the published drawing, Siphonalia noachina (Sowerby, 1846) as illustrated by Ortmann (1902: pl. 34: 5) from the late Oligocene to early Miocene of Pata­gonia, Argentina, has a similar overall sculpture, but a more convex whorl profile and fewer spirals per whorl.

In his faunal list for the limestone rocks near Belén and Pajarabobo, Olsson (1931: 22) lists “Austrofusus? belenensis n. sp.”, but a species of this name is neither described nor figured in his report. He also did not figure any neogastropods other than S.? belenensis and S.? tessaria from the Heath shale. Hence, Austrofusus? belenensis is a nomen dubium.

Stratigraphic and geographic range.—Oligocene to early Miocene of western Washington state, USA; early Oligo­cene of the Talara Basin, northern Peru.

Buccinidae indet.

Fig. 17C.

Material.—One specimen (NRM Mo187064, H = 7 mm) from the Belén seep site, upper Oligocene, Talara Basin, Peru.

Remarks.—The specimen differs from Colus sekiuensis by its distinctive opisthocline axial ribs and its seemingly more convex whorl profile. Among the buccinids reported by Olsson (1931) from northern Peru, Cyrtochetus? chiraensis Olsson, 1931, has an overall similar shape, but lacks distinctive axial ornament. Olsson’s Siphonalia species discussed above in the context of Colus sekiuensis differ from this species by their more angular whorl profile.

Subclass Heterobranchia Burmeister, 1837

Family Acteonidae d’Orbigny, 1843

Genus Acteon Montfort, 1810

Type species: Bulla tornatilis Linnaeus, 1758, by original designation; Recent, North Atlantic Ocean.

“Acteon” sp.

Fig. 17D.

Material.—One specimen (NRM Mo187065, H = 3.3 mm) from Cerro La Salina block 7, upper Oligocene, Talara Basin, Peru.

Remarks.—The single available specimen is 3.3 mm high, has a distinct shoulder, and is sculptured by numerous fine, beaded spiral cords. A similar species is Acteon annectens Meyer, 1885, from the late Eocene Moodys Branch Formation in Mississippi, USA (Dockery 1977). Acteonids have also been reported from Late Cretaceous seep deposits in Japan (Kaim et al. 2009) and Oligocene seep deposits in Washington, USA (Kiel 2006). The generic name is given in inverted commas here because genera among the Acteonidae are mainly distinguished based on anatomy and radula rather than on conchological characters (Rudman 1971).

Family Cylichnidae Adams and Adams, 1854

Genus Cylichna Lovén, 1846

Type species: Bulla cylindracea Pennant, 1777, by subsequent designation; Recent, North Atlantic Ocean.

Cylichna sp.

Fig. 17E.

Material.—One specimen (NRM Mo187066, H = 5 mm) from Cerro La Salina block 1, one unnumbered specimen from Cerro La Salina block 6. Both upper Oligocene, Talara Basin, Peru.

Remarks.—The two available specimens are 5 mm high, have an evenly convex whorl profile, and show numerous fine spiral incisions. Olsson (1931) rarely reported specimens smaller than a centimeter and, hence, such small opisthobranchs are virtually unknown from the Peruvian fossil record. A similar species is Cylichna atahualpa (Dall, 1908) from the Pacific coast of Costa Rica (Valdés and Camacho-García 2004). Cylichna-like shells have occasionally been reported from fossil seep deposits, including an Early Cretaceous site in California (Kaim et al. 2014), a Paleocene site in Spitsbergen (Hryniewicz et al. 2019), and an Oligocene site in Washington, USA (Kiel 2006).

Phylum Arthropoda Von Siebold, 1848

Class Malacostraca Latreille, 1802

Order Decapoda Latreille, 1802

Suborder Pleocyemata Burkenroad, 1963

Infraorder Axiidea Saint Laurent, 1979

Family Callianassidae Dana, 1852

Subfamily Eucalliacinae Manning and Felder, 1991

Genus Eucalliax Manning and Felder, 1991

Type species.—Callianassa quadracuta Biffar, 1970, by original designation; Recent, western Atlantic Ocean.

Eucalliax capsulasetaea sp. nov.

Fig. 18.

Zoobank LCID: urn:lsid:zoobank.org:act:3742EC6D-CCAA-443D-A113-16EA7C8C30CD

Etymology: In reference to capsulated setae, one of diagnostic features of the species.

Type material: Holotype: right propodus (NRM Ar69394) from block 2. Paratypes: right propodus (NRM Ar69376) from block 6, left carpus (NRM Ar69377) from block 7, right propodus (NRM Ar69383) from block 4, and four specimens from block 2: left propodus (NRM Ar69393), right propodus (NRM Ar69397), right propodus (NRM Ar69398), left propodus (NRM Ar69401).

Type locality: Cerro La Salina seep deposits, Talara Basin, Peru.

Type horizon: Presumably the early Oligocene part of the Heath shale.

Material.—The type material and further fragmentary specimens from Cerro La Salina blocks 1, 2–4, 6–8. Complete left carpus (NRM Ar69377), block 7. Seven near-complete left propodi: NRM Ar69378, block 7; NRM Ar69380 (counterpart NRM Ar69379), block 7; NRM Ar69381, block 7; NRM Ar69389 (counterpart NRM Ar69390), block 8; NRM Ar69393, block 2; NRM Ar69401 (counterparts NRM Ar69395, NRM Ar69400) block 1; NRM Ar69402, block 2. Five near-complete right propodi: NRM Ar69376, block 6; NRM Ar69383 (counterparts Ar0069384, NRM Ar69385), block 4; NRM Ar69394, block 2; NRM Ar69397, block 2; NRM Ar69398, block 2. Three fragmentary propodi: NRM Ar69382, block 7; NRM Ar69387, block 1; NRM Ar69396, block 2. One broken right fixed finger: NRM Ar69399, block 2. One complete right dactylus: NRM Ar69388, block 1. One fragmentary dactylus: NRM Ar69386, block 1. All upper Oligocene, Talara Basin, Peru.

Dimensions (in mm).—Measurements are given only for sufficiently preserved propodi (maximum height and length of the manus). NRM Ar69376, H = 12.6, L = 11.5; NRM Ar69378, H = 11.9, L = 12.8; NRM Ar69383, H = 16.0, L = 16.2; NRM Ar69389, H = ~10.0, L = 11.4; NRM Ar69393, H = 18.3, L = 18.1; NRM Ar69394, H = 11.6, L = 10.5; NRM Ar69397, H = 8.6, L = 8.5; NRM Ar69398, H = 12.2, L = 11.4; NRM Ar69401, H = 15.2, L = 19.1.

Diagnosis.—Major manus subrectangular; outer and inner lateral surfaces of manus densely covered with round tubercles, often with individual or serial setal pits; fixed finger with longitudinal keel.

Description.—Chelipeds (pereiopod 1) robust. Major carpus taller than long, not armed; upper margin keeled; lower and proximal margins forming continuous rounded margin with an indent at articulation with merus; distal margin with flange. Major propodus rectangular, robust; upper and lower margins keeled, parallel to each other or slightly converging distally; distal margin with or without blunt tooth below articulation with dactylus; area around articulation with dactylus forming bulge; inner and outer lateral surfaces strongly convex, densely covered with tubercles, tubercles closer to margins larger than those closer to longitudinal axis; larger tubercles occur closer to margins; tubercles usually with individual or serial setal pits. Outer lateral surface of fixed finger with longitudinal keel; occlusal margin of fixed finger with serrated keel forming blunt tooth distally. Dactylus robust, unarmed, tip slightly hooked; outer lateral surface with row of round setae positioned close to occlusal margin.

Remarks.—The species is placed within Eucalliax based on the subrectangular manus, the relatively short fixed finger with a triangular outline, and the presence of a longitudinal keel on the outer lateral surface of the fixed finger. This combination of features is characteristic for the genus as discussed in detail previously (Hyžný 2012; Hyžný and Hudáčková 2012).

Eucalliax capsulasetaea sp. nov. was presumably hetero­­chelous, as are most representatives of Callianassidae (Dworschak et al. 2012; Hyžný and Klompmaker 2015). Consequently, all the studied, rather large and massively built cheliped elements are interpreted to represent remains of major claws of pereiopod 1. The remains are nearly equally distributed between right and left chelae (five versus six propodi, respectively). Callianassid ghost shrimps in general show random asymmetry (sensu Palmer 2009) in Recent (e.g., Sakai 1969; Labadie and Palmer 1996) and fossil examples (e.g., Hyžný 2012; Hyžný and Hudáčková 2012), and Eucalliax capsulasetaea sp. nov. supports this observation.

In Eucalliax capsulasetaea sp. nov., the manus is approximately as long as tall (H/L ratio = 0.93–1.10), but one relatively large specimen is distinctly longer (H/L ratio = 0.80; see Dimensions above). NRM Ar69401 is also less tuberculate than other specimens. Nevertheless, all specimens are considered conspecific for several reasons. In extant representatives of Eucalliacinae (i.e., the species of Calliax Saint Laurent, 1973, and Eucalliax Manning and Felder, 1991), considerable variation in the manus H/L ratio is observed and distinctly elongated forms are not uncommon, especially in larger individuals (Dworschak 2006; Hyžný and Gašparič 2014). Also, the presence of setal pores on top of the tubercles covering the lateral surfaces of the elongated manus supports the conspecificity of all callianassid specimens from Cerro La Salina.

The setal pits positioned at the top of tubercles represent capsulated setae. They are not uncommon in hermit crabs (Paguroidea) and have been documented both in Recent (Komai 2003; Osawa 2012; Komai and Rahayu 2014) and fossil forms (Fraaije et al. 2011, 2015; Hyžný et al. 2016). Capsulated setae are rare in ghost shrimps. They have not been mentioned in taxonomic descriptions, but a quick survey of the published figures indicates their presence at least in fossil and extant species of Callianopsis Saint Laurent, 1973 (Karasawa 1997; Schweitzer Hopkins and Feldmann 1997; Lin et al. 2007), although they are limited to a more distal region (i.e., close to the articulation with the dactylus) and their number usually is not exceeding ten. In extant Neocallichirus karumba (Poore and Griffin, 1979) and its supposed fossil relatives, there are several (1–3) such tubercles present at the area of articulation with the dactylus (Dworschak 2008; Hyžný et al. 2016). Thus, Eucalliax capsulasetaea sp. nov. bearing numerous tubercles with capsulated setae is unique among both extant and fossil ghost shrimp species described to date, to our knowledge. The new species is morphologically closest to Eucalliax burckhardti (Böhm, 1911) from the Maastrichtian and Danian of Argentina and Mexico (Hyžný et al. 2013). Both species share densely tuberculated propodi; E. burckhardti, however, does not possess capsulated setae. Besides the presence of capsulated setae, the new species further differs from the type species E. quadracuta in the absence of diagnostic acute projections on the distodorsal and distoventral corners of the carpus and on the distodorsal corner of the propodus (Biffar 1970). Cenozoic representatives, such as E. vicetina Beschin, Busulini, De Angeli, and Tessier, 2002, from the middle Eocene of Italy; E. pseudorakosensis (Lőrenthey in Lőrenthey and Beurlen 1929) from the middle Miocene of Europe; E. yatsuoensis (Karasawa 1993) from the middle Miocene of Japan; and E. miyazakiensis Karasawa, 1993, possess smooth lateral surfaces of the major cheliped propodi. In contrast to the new species, E. yoshihiroi Karasawa, 1992, from the middle Eocene of Japan has sparse tubercles on the lateral surfaces of the major cheliped propodus, but it is not clear from the published figures alone whether setae are present at the top of the tubercles; no further details on this feature are mentioned in the description (Karasawa 1992). Newly presented material of Eucalliax reported from the late Eocene of Italy (Beschin et al. 2019) is too fragmentary and, based on the published figures, these specimens may not be congeneric with Eucalliax.

McLaughlin and Lane (1975) hypothesized that capsulated setae might act both to detect and repel predators, but more studies are needed to assess their function. The function of a limited number of capsulated setae in the ghost shrimps Neocallichirus and Callianopsis may be different than in hermit crabs and the here described new species of Eucalliax with numerous such setae. It is also worth noting that, whereas all Callianopsis species seem to possess capsulated setae, in Neocallichirus they are present only in the group of species morphologically close to N. karumba, a species differing profoundly from N. horneri Sakai, 1988, the type species of Neocallichirus. Consequently, the presence or absence of capsulated setae on ghost shrimps are not always a feature of generic importance, as currently understood.

Muscle scars are preserved in a number of specimens of Eucalliax capsulasetaea sp. nov. These are observed in specimens in which the cuticle is not fully preserved on the inner and outer sides of the manus (Fig. 18F), or, sometimes, on the cuticle. These scars represent closer (adductor) muscles (see Klompmaker et al. 2019), whereas the much smaller (abductor) opener is not seen here.

Eucalliax spp. have been found in the Late Cretaceous, Paleocene, Eocene, Miocene, Pliocene, and Recent (Hyžný and Klompmaker 2015: supplementary file 1). Thus, the new species is the first record from the Oligocene.

Stratigraphic and geographic range.—Type locality and horizon only.

Discussion

With Olsson’s (1931) and our study combined, 39 species of mollusks have been reported from the seep deposits in the Talara Basin, but only five species are shared between Olsson’s (1931) and our study (Pseudophopsis peruviana, Conchocele tessaria, Pleurophopsis unioides, P. talarensis, and “Vesicomya” tschudi; Table 2). The main reason for this difference is that Olsson did not consider small specimens: many species reported here are smaller than ca. 10 mm. Another reason might be that the sediments from which Olsson (1931) extracted his most diverse fauna (“between Belén and Pajarabobo”) appear to have been mostly eroded. This might explain why only five out of 13 of Olsson’s (1931) larger bivalve species are shared with our study.

Notably among the taxa we found are Coccopigya sp., which represents the oldest record of this genus, and Ascheria salina, representing the youngest and smallest member of Ascheria. Fossils are rare in the deep-water deposits of the Heath shale (Olsson 1931; Palacios Moncayo 1994) and, hence, methane seeps with their rapid in situ carbonate precipitation are more likely to preserve specimens. Consequently, the deep-water genera Neilo, Propeamussium, Coccopygia, Cantrainea, and Colus, and also the seep-restricted taxa Pyropelta, Provanna, Ascheria, and perhaps Retiskenea are here reported for the first time from the fossil record of Peru.

Among decapods, callianassid ghost shrimps are common associates of ancient seep environments (Klompmaker et al. 2018: fig. 4). Klompmaker et al. (2018) argued that they may be more commonly reported from the fossil seeps compared to modern seeps because these burrowing shrimps are difficult to catch in today’s ocean bottoms. The earliest records of callianassids from seeps thus far are mid-Cretaceous in age (Karasawa 2011; Kiel et al. 2013; Agirrezabala et al. 2013). Eucalliax had not been reported from a fossil seep thus far, unlike Callianassa, Callianopsis, Calliax, Glypturus, and Neocallichirus (see Klompmaker et al. 2018).

Conclusions

Combined with Olsson’s (1931) earlier work, our study revealed a remarkably diverse fauna of mollusks at the early Oligocene seeps in the Talara Basin in northern Peru, encompassing close to 40 species. The majority of them belong to genera known from both coeval and Recent seeps. The biogeographically closest ties are with late Eocene to Oligocene seep faunas in the northeastern Pacific region, with which they share two species and twelve genera. Notable is the absence of larger Bathymodiolus species from the seeps in the Talara basin, as these mussels are quite common in Oligocene seep deposits in the North Pacific region. A newly described ghost shrimp species of Eucalliax is the first occurrence of this genus reported from a seep.

Acknowledgments

We thank Ali Altamirano (Museo de Historia Natural, Lima, Peru) for assistance in the field, Cris Little and Jordan Bestwick (both Leeds University, UK) for letting us use their images of Pseudophopsis peruviana, Andrzej Kaim (Institute of Paleobiology, Polish Academy of Sciences, Warsaw, Poland) for comments on the gastropods, Jim Goedert (Wauna, USA) for comments on the provannids, Judith Nagel-Myers (St. Lawrence University, Canton, USA) and Leslie Skibinski (PRI) for loans of specimens, and the reviewers Kazutaka Amano (Joetsu University of Education, Japan) and Krzysztof Hryniewicz (Institute of Paleobiology, Polish Academy of Sciences, Warsaw, Poland) for their critical comments that greatly improved the manuscript. Financial support was provided by Vetenskapsrådet through grant 2016-03920 to SK, and by the Slovak Research and Development Agency through contract APVV-17-0555 to MH.

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