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Occurrence of benthic foraminifers, decapods, and other crustaceans on whale-falls, as reported in literature.

Age Region Depth/Environment Cetacean Taxon Crustaceans Benthic foraminifers References
Other Decapods
Eocene USA (LA) Shelf Archaeocete Arcoscalpellum jacksonensis Calappa sp.   Lancaster, 1986
late Oligocene USA (WA) Bathyal Mysticete   Callianopsis clallamensis   Nesbit, 2005
Miocene Japan (northwestern Hokkaido) Bathyal primitive Mysticete     Martinotiella communis, Spirosigmoilinella compressa Amano and Little, 2005
Miocene Japan (eastern Hokkaido) Upper bathyal (200-500 m) to Lower sublittoral (50-200 m) Indeterminate     large (∼300 μm diameter) benthic biloculine miliolid foraminifera, benthic foraminifera Amano et al., 2007
Pliocene Italy - Castellarano (Reggio Emilia) Shelf Mysticete   Decapods   Danise and Dominici, 2014
Pliocene Italy - Ponte a Elsa (Pisa) Shelf Mysticete   Decapods (?Goneplax sp.)   Danise and Dominici, 2014
Pliocene Italy - Orciano Pisano (Pisa) Shelf Mysticete   Decapods ( Eriphia sp.)   Danise and Dominici, 2014, Dominici, pers. comm. 2016
early Pleistocene Italy - Bargiano (Terni) - WFE2 100-150 m Mysticete   Albaidaplax ispalensis, Chlinocephalus demissifrons, Asthenognatus alleronensis*, ?Goneplax sp*., Jaxea sp.*, (?)thalassinidean and (?)goneplacid burrows Dominant: Bannerella gibbosa, Bigenerina nodosaria, Lenticulina calcar, Marginulinopsis costata, Siphotextularia concava, Vaginulina striatissima (see Appendix 2 for the complete list) this work; *Pasini et al., 2017
early Pleistocene Italy - Bargiano (Terni) - WFE1,3 100-150 m Odontocete (sperm whales)   (?)thalassinidean and (?)goneplacid burrows Dominant: Bannerella gibbosa, Bigenerina nodosaria, Lenticulina calcar, Marginulinopsis costata, Siphotextularia concava, Vaginulina striatissima (see Appendix 2 for the complete list) this work
Present day NW Pacific (Torishima Seamount) 4037 m Mysticete   Munidopsis spp. Cassidulinoides parkerianus (dominant), ‘Rhabdammina’ (?=Saccorhiza) ramosa, Reophax scorpiurus, Recuroides parkerae, Cystammina pauciloculata, Textularia kattegatensis, Textularia sp., Gyroidina quinqueloba, Tosaia hanzawai, Fursenkoina sp., Melonis pompilioides, Epistominella exigua, Spiroloculina sp. Fujioka et al., 1993;Wada, 1993; Wada et al., 1994; Naganuma et al., 1996
Present day NE Pacific (off southern California, Santa Catalina Basin, San Clemente Basin, San Nicolas, San Diego Trough, Santa Cruz Basin) 960-1960 m Mysticete Small lysianassid amphipods, Illyarachna profunda Lithodid crabs (Paralomis multispina?), Galatheids (Munidopsis quadrata*), Indeterminate crustaceans   *Bennet et al., 1994, Smith and Baco, 2003
Present day           Uvigerina peregrina, Globobulimina spp. Rathburn in Gooday and Rathburn, 1999
Present day Oceanic North Atlantic, South Atlantic, western Pacific, South Pacific (several sites) unknown unknown   Galatheidae indet.   Smith and Baco, 2003
Present day NE Pacific (Monterey Canyon) 382-289 m Mysticete Amphipods Indeterminate decapods, Chionoecetes tanneri, Munidopsis bracteosa, M. albatrossae, M. cascadia, Colossendeis gigas, C. japonica, Paralomis multispina, Paralomis verrilli, Galatheidae indet. Epistominella pacifica* Goffredi et al., 2004; Lundsten et al., 2010b; *Geist, 2010
Present day North Sea (Kosterfjord, Sweden) 30-125 m Mysticete Orchomene obtusa, Ostracods* Hyas araneus* Benthic foraminifers* Dahlgren et al., 2006;


*Danise et al., 2014

Present day NE Atlantic (Porcupine Seahight) 2555-2710 m Odontocete (Phocoena phocoena) Eurythenes gryllus, Tmetonyx sp., Abyssorchomene sp. Munidopsis crassa   Kemp et al., 2006
Present day Off southern Japan (Kyushu Island) 219-254 m Odontocete (sperm whales) Ostracoda, Heteralepas sp., Nebaliidae indet., Gammaridae indet., Cumacea indet., Euphausiacea indet., Alpheus sp. (macrochirus group?), Eualus sp. cf. kikuchii, Processa philippinensis, Plesionika crosnieri, Plesionika grandis, Callianassa s.l. sp., Cestopagurus sp. nov., Paguristes albimaculatus, Nematopagurus lepidochirus, Nematopagurus spinulosensoris, Eumunida sp., Galathea spp., Munida spp., Galatheidae indet., Homola orientalis, Homolidae indet., Ethusa sp., Cryptocnemus obolus, Leucosiidae indet., Merocryptus lambriformis, Macrocheira kaempferi, Pugettia minor, Oxypleurodon stimpsoni, Trachycarcinus sagamiensis, Cancer gibbosulus, Cancer japonicus, Carcinoplax surugensis, Medaeus serratus, Xanthidae ind.   Fujiwara et al., 2007
Present day NW Pacific (Peter the Great Bay, East Sea, Russia) 30 m Mysticete Amphipods, Isopods     Pavlyuk et al., 2009
Present day NE Pacific Ocean (off Vancouver Island, British Columbia) 1288 m Mysticete Amphipoda sp., Munnopsidae sp., Mysida sp. Paralomis multispina, Pandalopsis ampla   Lundsten et al., 2010a
Present day Antarctic (Kemp Caldera, East Scotia Sea) 1444-1447 m Mysticete Jaera tyleri, Lysianassidae indet., Ilyarachna sp., Sericosura sp. Nematocarcinus sp.   Amon et al., 2013; Linse et al. 2014
Present day Antarctic (continental slope off Anvers Island) 1430 m Mysticete Amphipoda (unidentified) Paralomis birsteini   Smith et al., 2014
Present day SW Atlantic (São Paulo Ridge, off Brasil) 4204 m Mysticete Stephonyx sp., Epicaridea sp., Copepoda Munidopsis spp.   Sumida et al., 2016




Alphabetical list of Foraminifera species found in WFE 1-3 at Bargiano.

Planktonic species

Globigerina bulloides d’Orbigny, 1826

Globigerina cariacoensis Rögl & Bolli, 1973

Globigerinoides ruber d’Orbigny, 1839

Globigerinoides sacculifer Brady, 1877

Globorotalia inflata (d’Orbigny, 1839)

Orbulina universa d’Orbigny, 1839

Turborotalita quinqueloba (Natland, 1938)

Benthic species

Ammonia beccarii (Linnaeus, 1758)

Ammonia parkinsoniana (d'Orbigny, 1839)

Ammonia perlucida (Heron-Allen & Earland, 1913)

Ammonia tepida (Cushman, 1926)

Asterigerinata mamilla (Williamson, 1858)

Bannerella gibbosa (d’Orbigny, 1826)

Bigenerina nodosaria d’Orbigny 1826

Bolivina spathulata (Williamson, 1858)

Bulimina marginata d’Orbigny, 1826

Bulimina spinata Cushman & Campbell, 1935

Cancris auriculus (Fichtel & Moll, 1798)

Cassidulina laevigata d’Orbigny, 1826

Heterolepa floridana (Cushman, 1918)

Lenticulina calcar (Linnaeus, 1758)

Lobatula lobatula (Walker & Jacob, 1798)

Marginulinopsis costata (Batsch, 1791)

Melonis barleeanus (Williamson, 1858)

Nonionella turgida (Williamson, 1858)

Quinqueloculina seminula (Linnaeus, 1758)

Siphotextularia concava (Karrer, 1868)

Uvigerina mediterranea Hofker, 1932

Vaginulina striatissima Schrodt, 1890




Description of morphological test character of Vaginulina cf. V. striatissima. (Figure 6, in the text).

Calcareous hyaline test, elongated, slender, and strongly compressed with a progressive enlargement from the basal chamber towards the apertural chamber. Prominent basal spine, generally well preserved, sometimes broken in the apical portion. Test covered by numerous well developed costae. The shape of the test is almost straight on one side and slightly curved on the other. The maximum size of the test is close to the opening and can reach as much as 1 mm. The chambers, generally flattened, vary in number from eight to twelve (in larger specimens). Generally, the last two chambers are slightly inflated. The sutures between the chambers are usually difficult to visualize; a thin groove is evident in some specimens, usually between the last and third last chamber. The sutures are oblique, scarcely visible in the basal chambers, and become progressively more visible approaching the last chamber. The aperture is aligned with the straight edge and is eccentric and circular with a radiate edge.

The test is crossed for its entire length by rectilinear costae, often bifurcated and/or anastomosed, which are not interrupted by the sutures between chambers. The costae branch off from the upper part of the basal spine and appear to diverge into a herringbone pattern from the margin of the straight side, generating a thin flange-shaped projection (Figure 6.2-3). The costae crossing the last chamber converge in the radiate opening (Figure 6.8, 6.11). A smooth or thinly costate inclined surface connects the top of the last chamber with the aperture.

Dimensions. Maximum lengths are between 4.0-4.5 mm with a minimum of 2mm in broken specimens missing the last chamber. The width of the apertural chamber is generally less than one millimeter; the width of the aperture is approximately 0.1 mm.

Type locality. Whale-fall sites at Bargiano and Montemoro (see Monaco et al. 2014), southwestern Umbria, Allerona, Central Italy.

Type levels. Early Pleistocene, Calabrian, clay marine deposits of Chiani-Tevere Unit (Baldanza et al. 2013, 2014).

Observations. The morphology of the test, which deviates from the characteristics of Vaginulina striatissima (originally drawn by Schrodt, 1890, plate 21, figure 9a-b, Ellis and Messina, 1940-2006) leads us to hypothesize that modifications in the costae and spine of these foraminifera were induced by the soft clay substrate and a high availability of nutrients that stimulated reproduction. The specimens found in the CFT section, compared to those found in Sections S1-S4, show differences in length of the test and in number of costae (radiating from basal spine to last chamber), which become thinner and more anastomized (Figure 6.6-6.7, 6.9, 6.12).

The development in our specimens of costae and basal spine may fall within the specific variability of the taxon, though we do not exclude that the taxon we identified as Vaginulina cf. V. striatissima may be included into a new species, the origin of which may be closely linked to the peculiarities of the living environment.

Notes. The specimens figured in Ellis and Messina (1940-2006) as Vaginulina striata d’Orbigny, 1826 (figs. 36, 58) and Vaginulina striatissima Schrodt, 1890 (figs. 9a-9b) are drawings: d’Orbigny and Schrodt both reported all the morphological characters described in our specimens (test shape, aperture shape and position, costae, and basal spine), but not all together.



Morphology of specimens attributed to Albaidaplax ispalensis Garassino, Pasini and Castro, 2013 (Figure 7.1, in the text).


Superfamily GONOPLACOIDEA MacLeay, 1838
Family GONEPLACIDAE MacLeay, 1838
Subfamily GONEPLACINAE MacLeay, 1838
Genus ALBAIDAPLAX Garassino, Pasini and Castro, 2013

Type species. Albaidaplax ispalensis Garassino, Pasini and Castro, 2013, by monotypy.

Included species. Albaidaplax ispalensis Garassino, Pasini and Castro, 2013.

Albaidaplax ispalensis Garassino, Pasini and Castro, 2013
Figure 7.1

v. 1998 Goneplax gulderi Bachmayer; Mayoral et al., p. 508, 509, fig. 2 (5).

v. 1948 Goneplax cf. saccoi [sic]; Vía, p. 146, fig. 6.; Solé and Vía, 1989: 34.

v. 1989 Goneplax cf. saccoi [sic]; Solé and Vía, p. 34.

V* 2013 Albaidaplax ispalensis Garassino, Pasini and Castro, p. 362, 363, fig. 4.

v. 2014 Albaidaplax ispalensis Garassino, Pasini and Castro; Pasini et al., p. 252, fig. 8A.

Carapace rectangular, slightly wider than long, straight front wide as the orbits; wide orbits expanded distally with a short, triangular forward-directed tooth on outer orbital angle; supraorbital margins gently convex; anterolateral margins slightly convex with a smooth, short, barely distinct tooth; posterolateral margins slightly narrowing posteriorly; straight posterior margin; smooth dorsal surface of carapace; weak transverse ridges moderately convex; dorsal carapace without clear indication of regions; stout chelipeds with moderately long and large merus; subtriangular carpus spineless; palm smooth dorsally; dactylus and index nearly as long as the propodus; unarmed occlusal margins; ambulatory legs elongate, flattened, and slender.

Material and measurements. Five specimens in dorsal view, housed in the Museo di Allerona (MUAL) (MUAL 3 - carapace length: 28 mm; carapace width: 31 mm. MUAL 6 - carapace length: 22 mm; carapace width: 25 mm. MUAL 7 - carapace length: 15 mm; carapace width: 18 mm. MUAL 8 - carapace length: 25 mm; carapace width: 28 mm. MUAL 9 - carapace length: 28 mm; carapace width: 31 mm).




Morphology of specimens attributed to Chlinocephalus demissifrons Ristori, 1886 (Figure 7.2-3, in the text).


Family EURYPLACIDAE Stimpson, 1871
Genus CHLINOCEPHALUS Ristori, 1886

Type species. Chlinocephalus demissifrons Ristori, 1886, by monotypy.

Included species. Chlinocephalus demissifrons Ristori, 1886.

Note. Schweitzer et al. (2010) erroneously reported two species as belonging to this genus: C. demissifrons Ristori, 1886, and C. subovalis (Ristori, 1886). Ristori (1886) described Chlinocephalus based on the type and sole species, C. demissifrons. No species named subovalis is reported in the original text. The genus is therefore monotypic, as correctly reported by Karasawa and Kato (2003).

Chlinocephalus demissifrons Ristori, 1886
Figure 7.2-3

V* 1886 Chlinocephalus demissifrons Ristori, p. 101-103, pl. 2, figs. 5, 6.

v. 2003 Chlinocephalus demissifrons Ristori; Karasawa and Kato, tab. 5.

v. 2004 Chlinocephalus demissifrons Ristori; Garassino et al., p. 275-278, figs. 15, 16.

v. 2013 Chlinocephalus demissifrons Ristori; Pasini and Garassino, p. 321, fig. 2.

Suboval carapace wider than long, convex dorsally and nearly smooth; wide fronto-orbital margin extends over the orbits; frontal straight margin with weak median frontal incision and two distinct oblique pre-orbital incisions laterally; concave deep wide orbits obliquely directed to the frontal margin, with a small short median incision; supraorbital margins concave with a short, triangular point at the extremities; short anterolateral margins with two sharp, pointed, triangular spines, the first close to the extraorbital tooth, on the hepatic margin, the second on the epibranchial margin at the level of a smooth dorsal transverse carina that is slightly curved and convex in the middle; posterolateral margins convex wide, crossed by a second transverse dorsal carina, nearly straight medially; wide posterior margin gently convex; carapace dorsal region not distinct, slightly depressed between the two transverse ridges, and posteriorly, behind the second ridge toward the posterior margin; postfrontal region depressed slightly below level of remainder of carapace; suboval wide cardiac region poorly marked; dorsal carapace ornamented with sparse pits or coarse granulations, notably along the transverse ridges; P1 well developed, with elongate merus; globular carpus with a strong spine on inner distal margin; robust chelae with flattened inner palm; elongate, pointed, and armed dactylus and index, curved, converging distally; granulated chelipeds, possibly unequal (longer in male); elongate, flattened, and slender ambulatory legs, ovoid in transverse section; male thoracic sternum subcircular, with finely punctulate surface; sternites 5-7 petaloid in shape, sternite 8 reduced, poorly preserved; male pleon narrow, slender, T-shaped, with six movable subtrapezoidal somites plus telson, narrowly triangular; somites and telson slightly inflated longitudinally forming a median smooth, rounded, tubercle-like protuberance; wide trapezoidal s3 covering space behind P5 coxae; s4-s6 shorter, slightly narrowing to the triangular telson, rounded distally.

Material and measurements. Ten complete and incomplete specimens; we selected three specimens in dorsal and ventral view, slightly compressed dorso-ventrally (MUAL2: ventral view; MUAL10 - carapace length: 25 mm; carapace width: 32 mm. MUAL11-12 - carapace length: 32 mm; carapace width: 39 mm).


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