How Does a Whale Fall Community Affect Ocean Sediment – How Does a Whale Fall Community Affect Ocean Sediment

The early on Pleistocene whale-autumn community of Bargiano (Umbria, Central Italy): Paleoecological insights from benthic foraminifera and brachyuran crabs

Angela Baldanza, Roberto Bizzarri, Federico Famiani, Alessandro Garassino, Giovanni Pasini, Marco Cherin, and Francesco Rosatini

Article number:
Copyright Paleontological Lodge, April 2018

Author biographies
Plain-language and multi-lingual abstracts
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Submission:  2 May 2017 . Acceptance:  24 March 2018

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New insights into communities of benthic foraminifera and decapods, associated with whale-fall events (WFE) in a relatively shallow sea environment, are reported hither for the first fourth dimension from the early Pleistocene of Bargiano (southwestern Umbria, Italy). The inferred paleodepth of these WFEs is non greater than 100−150 m and, on the basis of more full general stratigraphic data, took identify over an estimated flow of about l,000 years. The foraminifera assemblages associated with these WFEs are depression in number of planktonic and benthic taxa, and six benthic species boss: the shallow infaunal species
Bigenerina nodosaria, Bannerella gibbosa, Marginulinopsis costata,


Five. striatissima, along with the epifaunal species

Lenticulina calcar

Siphotextularia concava. Considering these opportunistic species respond to curt-term favorable atmospheric condition by increasing in number and maintaining stable populations, the presence of high numbers of individuals of these species in association with three recognized WFEs provides evidence that a nutrient-rich environment favored their proliferation. The occurrence of previously unreported benthic foraminifera taxa (across the 3 WFEs), forth with the presence of the crab species
Albaidaplax ispalensis
(Goneplacidae) and
Chlinocephalus demissifrons
(Euryplacidae) (in at least one WFE), offer new insights into shallow sea whale-fall fossil communities.

Angela Baldanza (corresponding author), Department of Physics and Geology, Academy of Perugia, Via A. Pascoli ‒ I-06123 Perugia, Italy,

Roberto Bizzarri, Department of Physics and Geology, University of Perugia, Via A. Pascoli ‒ I-06123 Perugia, Italian republic,

Federico Famiani, Paleontological Museum – Mountain Subasio Regional Park, Loc. Cà Piombino – I-06081Assisi, Perugia, Italy.

Alessandro Garassino, Natural History Museum, Paleontology Department, Corso Venezia 55 – I-20121 Milan, Italy.

Giovanni Pasini, Via Alessandro Volta 16 – I-22070 Appiano Gentile (Como), Italian republic.

Marco Cherin, Department of Physics and Geology, University of Perugia, Via A. Pascoli ‒ I-06123 Perugia, Italy,

Francesco Rosatini, Via Strada di San Giacomo 1/D – I-06132 Fontignano (Perugia), Italy.

Keywords: Benthic communities; Decapoda; Whale-fall events; Paleoecology; Calabrian; Paleo-Mediterranean Bounding main

Final citation: Baldanza, Angela, Bizzarri, Roberto, Famiani, Federico, Garassino, Alessandro, Pasini, Giovanni, Cherin, Marco, and Rosatini, Francesco. 2018. The early Pleistocene whale-fall community of Bargiano (Umbria, Central Italy): Paleoecological insights from benthic foraminifera and brachyuran crabs.
Palaeontologia Electronica
21.1.11A 1-27.

 Copyright: April 2018 Paleontology Gild. This is an open up admission article distributed under the terms of Attribution-NonCommercial-ShareAlike 4.0 International (CC By-NC-SA 4.0), which permits users to copy and redistribute the material in any medium or format, provided it is not used for commercial purposes and the original author and source are credited, with indications if any changes are made.


The carcasses of great whales (including both baleen and sperm whales) are the largest class of detritus to collect on the sea flooring. Expressionless whales, whose bodies consist largely of proteins and lipids, typically sink to the sea floor where they serve equally an enormous, temporary source of organic enrichment: a forty-ton gray whale, for example, supplies approximately two 1000000 g C, equivalent to >2000 years of background C flux in the expanse beneath and around the carcass (Smith, 2006).

Over the last ii decades, research into modernistic and fossil whale-fall events (WFE) has increased considerably, providing new insights on whale-autumn ecosystems (Smith et al., 2002, 2015; Smith and Baco, 2003; Goffredi et al., 2004, 2008; Smith, 2006; Dominici et al., 2009; Danise et al., 2010, 2014; Higgs et al., 2010, 2012; Lundsten et al., 2010a, 2010b; Danise and Dominici, 2014; Roman et al., 2014; Taboada et al., 2016).

A whale carcass on a seabed provides a massive banquet for other organisms, and sea floor feeding and dismantling of a carcass past macro- and micro-scavengers and carnivores has been observed and studied in the Pacific and Atlantic Oceans, deep-body of water Antarctic, and other internal seas (Fujioka et al., 1993; Bennett et al., 1994; Wada et al., 1994; Smith et al., 2002, 2015; Smith and Baco, 2003; Goffredi et al., 2004; Fujiwara et al., 2007; Lundsten et al., 2010a, 2010b; Amon et al., 2013; Linse et al. 2014; Smith et al., 2014; Sumida et al., 2016). The surface area around a decomposable carcass becomes a specialized, if temporary ecosystem rich in organic matter and sulfides, a kind of “habitat island” on the seafloor (Smith et al., 2015).

Great-whale carcasses (baleen and sperm whales) are able to support both a sequence of heterotrophic and chemosynthetic microbial assemblages and metazoan communities in the energy-poor deep body of water, all of which pass through a series of 3 principle and overlapping ecological stages (Smith et al., 2002, 2015; Smith and Baco, 2003; Smith, 2006). Each phase is characterized by a diverse nutrient chain and diverse beast, with differing species associated with bounding main or internal-bounding main WFEs. The whale-fall ecosystem supports the evolution of a characteristic fauna, which reaches and maintains high species abundance for decades. Thus, many new species and evolutionary novelties have been recorded, including bone-eating annelid worms, gastropods, a variety of grazers on sulphur leaner, in addition to typical mollusk and serpulid assemblages (Smith et al., 2015, and references therein).

Research on WFEs is commonly focused on assemblages of microbes or of mollusks and other metazoans (Smith et al., 2015, and references therein); less is known, conversely, about benthic foraminifera and decapods associated with whale falls (encounter Appendix 1). If the presence of decapods (amid other crustaceans such equally amphipods, isopods, and rare barnacles and ostracods) is already documented, notwithstanding, data about benthic foraminifera are rarely reported (Fujioka et al., 1993; Wada, 1993; Wada et al., 1994; Naganuma et al., 1996; Gooday and Rathburn, 1999; Rathburn et al., 2003; meet also Geist, 2010). The modern-day WFEs that have been analyzed to date unremarkably involve events at great depths; in those environments, benthic foraminifera assemblages were generally scarce while typical deep-bounding main species that tolerate low oxygen content were noted (Gooday and Rathburn, 1999).

Research on fossil whale falls from the Eocene to the Pliocene (meet Appendix ane) also contributed to descriptions of faunal assemblages (mainly invertebrates) in shelf to bathyal paleoenvironments (Lancaster, 1986; Amano and Little, 2005; Nesbit, 2005; Amano et al., 2007; Dominici et al., 2009; Danise et al., 2010, 2014). Among these findings, only Amano and Petty (2005) and Amano et al. (2007) reported data nearly benthic foraminifera assemblages associated with fossil WFEs.

The unusual assemblage of benthic foraminifera and the decapod fauna recovered at the Bargiano site (southwestern Umbria, primal Italy), both of which are related to the presence of organic affair in proximity to a whale carcass, stand for a novelty for fossil whale-fall events, peculiarly for the Quaternary.

The silty clayey marine sediments of the early Pleistocene Bargiano site (southwestern Umbria, central Italian republic) are already known in the paleontological record for their exceptional and unusual preservation of
Ambergrisichnus alleronae
fossil cololites. Baldanza et al. (2013) and Monaco et al. (2014) described several WFEs preserved at Bargiano (and more generally in the Allerona area) that appear to take occurred over a relatively curt span of time.

Paleontological excavations in the Allerona area during 2003, 2008, and 2016 yielded 3 partial skeletons of baleen whales. Although fossils of these marine mammals are abundant in other areas of the Italian peninsula (e.g., Tuscany, Piedmont, Emilia Romagna, and Salento; meet Bianucci, 2014), the Allerona finds represented the first reports of cetaceans in Umbria. The specimens are still under grooming and report, but preliminary observations let usa to refer them to at least two different mysticete families: Balaenidae and Balaeonopteridae. These data add together to the massive presence of sperm whales, documented by the occurrence of
Ambergrisichnus alleronae
(Baldanza et al., 2013; Monaco et al., 2014). Indeed, the entire Allerona area, not merely the well-known Bargiano site, represents a sea flooring paleoenvironment deeply conditioned by WFEs and past the dispersion of decayed cetacean remains. The Bargiano scenario reported here permits u.s.a. to speculate on the original ecology scenario and on the paleoecological relationships amidst and behavior of the marine invertebrates collected within and around the whale falls.

The benthic infaunal foraminifers collected at Bargiano (Appendix two and Appendix 3), preliminarily reported by Rosatini (2016), nonetheless announced dissimilar to those reported from fossil or mod-twenty-four hours WFEs. In addition, decapod specimens, located around or between the basic of a big cetacean, were recovered (for a detailed assay, see Appendix 4 and Appendix v), including a new species of brachyuran crab (Asthenognatus alleronensis), previously unknown from the Pleistocene of the Mediterranean area and recently described by Pasini et al. (2017).


s figure1The study site (Effigy 1.1) lies close to the town of Allerona and is office of the South Valdichiana Basin. During the early Pleistocene, the basin accommodated a relatively shallow marine environment (on the order of a maximum of 100−150 1000). The geological, sedimentological, and stratigraphic description tin be found in Baldanza et al. (2013) and Monaco et al. (2014).

In the Bargiano section, massive to thinly-laminated offshore marine deposits crop out, ranging from clay to clayey silt to silty clay (Figure 1.2). Calcareous nannofossil and foraminifer assemblages from this site betoken an early Pleistocene (Calabrian) age (CNPl8 Zone
Backman et al., 2012; MPLe1 Zone
Iaccarino and Premoli Silva, 2007), and a relatively shallow offshore environment (about 100−150 one thousand). The recent recovery of a baleen whale skeleton (Effigy 1.three) has allowed us to consummate a vertical stratigraphic contour of the site.

A section about 2 m thick (WS), measured from some 70 cm higher up the skeletal remains to about one chiliad below, was described (Effigy one.ii-three, Figure two). Within that section, three main decapod horizons were recognized, although we as well noted the irregular occurrence of decapods throughout the unabridged department (Figure ii). In addition, 4 shorter sections (S1, S2, S3, and S4) were described and sampled at increasing distances from the whale skeleton in social club to investigate lateral and vertical variations in microfossil distribution (Figure one.two, Figure 2). Because of the slight n-eastern inclination of the dirt beds, these 5 sections permitted the reconstruction of a 2.8 m-thick composite stratigraphic section (Effigy ii). Finally, a 30 cm-thick section was described in the master cololites field (CFT: Cololite Field Trench department, Figure 1.2, Effigy ii), about 50 m northeast of and 4 m stratigraphically above the cetacean skeleton. Such interpolation was allowed to recognize a total of 25 irregularly-spaced stratigraphic levels (Figure 2), 22 of which lie in the main section with 3 in the CFT section.


s figure2Forty samples were nerveless from the 25 measured levels across the six sections (Figure 1.2, Figure 2). About 200 g of sediment from each sample were processed for micropaleontological analyses. The sediment was done with hydrogen peroxide and water then sieved through a 63 μm mesh to obtain the rest. Residues more often than not did not exceed two one thousand. The unabridged washed residue was observed under a stereomicroscope (Nissho optical, TZ240) for semi-quantitative assay of full and relative abundances and species diversity of foraminifera. Selected benthic foraminifera specimens were analyzed using a scanning electron microscope (SEM – Philips, 515) for the identification of morphological characteristics.

Of the twenty decapod specimens recovered, but the seven best-preserved, partially in life position and with exposed dorsal and ventral views, were considered. The specimens, embedded in pocket-size blocks of blue-greyness clay and covered by a thin patina of yellow-brown iron oxide were advisedly prepared and, due to the delicate nature of the fossils and the loose matrix, stock-still with polyvinyl acetate for study and preservation. The specimens were labeled with the MUAL (Museo di Allerona) institution code, progressively numbered, catalogued, and stored in the museum.


Recent field surveys (2015 and 2016) enabled the completion of the preliminary stratigraphic contour of the Bargiano site reported by Monaco et al. (2014). Crude silty clay beds dip more-or-less uniformly north-due east across the study area, but local stratification can rarely be recognized considering of the presence of three main families of joints (late Pleistocene-Holocene tectonics).

s figure3With regard to the fossil record, the species of malacofauna at Bargiano are reported in Monaco et al. (2014, table 1), accompanied locally by echinoids, and showed no pregnant variations through the sections shown in Figure two, whether vertically or laterally. Decapod crustaceans, together with undescribed thalassinidean and goneplacid burrows (Pasini et al., 2017, p. 69), were highly abundant only within a curt radius from the whale bones (Figure 3), while their occurrence in the rest of the blended section was sporadic.

The foraminifera assemblages recovered from all the samples were, in general, depleted in both planktonic and benthic taxa, although relative-affluence values (RA) highlighted the predominance of six epifaunal or shallow infaunal taxa, with the latter type being more common (Figure 4).

Whale-autumn Events

s figure4Throughout the department, three primary WFE horizons were recognized (Figure 2). They were stratigraphically numbered and named WFEs i−iii. The lowermost event (WFE 1: Figure 1.2, Figure two) is characterized by Cololite Horizon #25 (shut to the excavation site; Figure i.2, 1.five, Figure 2) and #27 (about 22.5 m east; Figure ane.2, 1.4, Figure ii). WFE two corresponds to the baleen whale skeleton horizon (Figure ane.iii, Figure 2); fragments of a cololite (Cololite #24) were noted close to the whale remains, but the cololite’s original position was non establish. Finally, WFE 3 is the main area in which
Ambergrisichnus alleronae
fossils were establish (Figure 1.2, Figure two), as described past Baldanza et al. (2013) and Monaco et al. (2014). The CFT section was measured and sampled close to Cololite #four (Effigy 1.2). These WFEs are in addition to the three (or four) events already noted throughout the Montemoro department described past Monaco et al. (2014, figure ane.ii).

Planktonic Foraminifera

Across the six sections, planktonic foraminifera were largely comparable to those reported past Monaco et al. (2014). The assemblages in both the Bargiano and Montemoro sections included very few taxa
(Globigerinoides ruber, Globigerinoides sacculifer, Globigerina bulloides, Globorotalia inflata, Globigerina cariacoensis,

Orbulina universa),

and abundance was low.
Turborotalita quinqueloba
sporadically occurred in depression frequencies. Plankton reached mutual values in merely a few samples: at the top of S4, in samples 3-7 of S1, at the top of WS, and at the tops of S3 and S2 sections (see Figure two). In the CFT department (Effigy two), planktonic foraminifera were common everywhere, with
O. universa
arable at the base of operations.

Benthic Foraminifera

Preliminary data reported by Monaco et al. (2014) indicated that epifaunal and shallow infaunal foraminifera taxa were predominant at the Bargiano site, confirming the abundance of degraded organic affair derived from WFEs.

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General observations.
The assemblages were very depression in number of taxa, and relative abundances (RA) were likewise low, except in the case of

Bigenerina nodosaria, Bannerella gibbosa, Marginulinopsis costata, Lenticulina calcar, Siphotextularia concava,


V. striatissima. The benthic taxa showed low species-diversity values (SD), varying from 17 to 1 as a function of distance from the whale skeleton or cololites. Lower SD values were observed in the WS section. SD values were highest, particularly in the lowermost levels, in sections S1 and S4, too equally in S2 and S3, located at varying distances from the whale skeleton (Figure 1, Figure 2), while the WS section showed medium SD values (from 8 to 6). In the CFT section, the assemblages were always poor, whereas the SD, which included planktonic taxa, varied from 20 to 15.

The taxa are epifaunal and shallow infaunal. Detritivores are near common amongst the epifauna
(Lenticulina, Cancris, Cassidulina, Siphotextularia, Gyroidina,


Bannerella, Fursenkoina, Bolivina, Nonionella, Melonis, Bulimina,
predominant among shallow infauna. Generally subordinate genera include the herbivorous epifaunal
Martinottiella, Quinqueloculina, Lobatula,
Asterigerinata; the shallow infaunal
Ammonia; the epifaunal suspension feeders
Lobatula; and the shallow infaunal omnivore


s figure5Ascendant taxa.
Representatives of Families Textulariidae, Vaginulinidae, and Eggerellidae (come across Appendix 2) reached considerable statistical frequencies in sediments close to skeletal remains (Effigy 2, Figure iv) and predominated the highly depleted assemblages (Effigy four, Effigy five, Figure half-dozen). Every bit reported past Murray (2006), the preponderance of foraminifera living in fine-grained sediment below the photic zone are infaunal, feed on detritus or leaner, and are abundant in the uppermost one−2 cm of sediment. Textulariidae, in particular, are passive detritivores (detrital and bacterial scavengers), while Vaginulinidae are stationary infaunal deposit feeders (Murray, 1991). The highest RA values (varying from mutual to abundant to very abundant; run across Figure 4) in all the sections were reached by the shallow infaunal taxa
Bigenerina nodosaria, Siphotextularia concava
(Textulariidae), and
Bannerella gibbosa
(Eggerellidae) and the Family Vaginulinidae species,
Marginulinopsis costata, Vaginulina
V. striatissima
(both shallow infaunal species) and

Lenticulina calcar
(epifaunal). Moreover, when the SD values were very low (in the WS department, for instance), only these six taxa occur in assemblages. Equally Figure four makes clear, the RA of the six taxa showed varying trends across the 25 levels and the three WFEs.

In all sections, unexpectedly,
Bigenerina nodosaria
was the dominant taxon: its RA varied from mutual to abundant, with the exception of the S2 department where information technology was common at the base of operations and decreased upwards to rare (Figure 4, Effigy five.1-3). Much is known about the lifestyle and environmental of
B. nodosaria. Mojtahid et al. (2010), for case, in enquiry at a 320-thou deep station in Biscay Bay (NE Atlantic) under eutrophic weather condition, identified it as a shallow infaunal species with a penetration depth of from 0.vii to ii.0 mm; the authors also characterized
B. nodosaria
as a mutual taxon all around the Mediterranean in circalittoral and upper bathyal muds (Sgarella and Moncharmont-Zei, 1993). de Stigter et al. (1998) establish
B. nodosaria
from the shelf/upper slope to the eye gradient in the Southern Adriatic Sea, and Kaminski et al. (2002) and Chendes et al. (2004) reported
B. nodosaria
in large numbers beneath 140 m in the Marmara Sea, with a critical signal at virtually 220 thou. Fontanier et al. (2002) identified
B. nodosaria
as a eutrophic species, well developed in environments where at that place was high marine productivity. Finally, Gross (2000) observed an elevated hateful migration speed for this taxon during food-enrichment experiments.

s figure6The presence of
Bigenerina nodosaria
in high numbers across Sections WS, S1, and S4 (the latter two are located virtually 6.v m from the whale skeleton, denoted WFE 2) suggests that the habitat was ideal for its omnivorous lifestyle, and that the increase in organic matter in sediments below the carcass was non a barrier to development. It is worth noting that
B. nodosaria
became rare in Section S2, probably because, at 8.5 yard from the skeleton, the content of organic matter in the sediments was considerably reduced. In Section S3,
B. nodosaria
was again abundant and mutual only, in this scenario, the most likely source of nutrients was WFE 1 (Figure 1, Figure two), as documented by the presence of Cololite #27 almost 6 m west of Section S3. The occurrence of
B. nodosaria
in highly depleted assemblages, such as those found in this study, is additional disarming evidence for the opportunistic character of this species and its chapters to drift to food-rich areas ideal for evolution and reproduction.

A like behavior seems to characterize
Bannerella gibbosa
(Figure 4, Figure 5.viii-nine), a shallow infauna and detritivorous species that occurs abundantly in Section S4 and which is common and arable in Sections WS and S2 (in levels close to the whale skeleton). In Section S3,
B. gibbosa
is rare, becoming common only in the uppermost samples. The species appears to take an affinity for sediments rich in abundant but likely depression quality organic matter, though its behavior is not known in particular. The other representative of the Textulariidae, the epifaunal detritivore

Siphotextularia concava
(Figure 4, Figure five.10-15), was common in Sections S4 and S1, except in i basal sample of S1 in which information technology was abundant. In Section WS, it was generally rare to common, constantly common in Section S2, and very abundant in Department S3. This species too shows a high affinity for sediments rich in organic affair but does not seem to tolerate high organic flux, as suggested past its rarity in sediments proximate to skeletal remains. An epifaunal lifestyle probably affected the development of
Due south. concava, and it was probable unable to develop if nutrients at the sediment-water interface were overloaded.

The stationary infaunal deposit feeder family Vaginulinidae (Murray, 2006) is represented at the Bargiano site by
V. striatissima

Marginulinopsis costata

(probably shallow infaunal taxa, considering the test morphologies and position of aperture) and
Lenticulina calcar
(a detritivore epifaunal species that prefers common cold water; see Murray, 2006). In particular,
V. striatissima
(Figure 4, Figure vi), until now reported but from Pleistocene clays, shows high frequencies. The Bargiano specimens are covered by well-developed costae and brandish a prominent basal spine (Effigy six) and are, moreover, very large, ranging 4−5 mm in length.

The morphology of the test (encounter Appendix three for detailed description and taxonomy notes), which deviates from the characteristics of
Vaginulina striatissima
(original drawn by Schrodt, 1890, plate 21, figure 9a-b: Ellis and Messina, 1940-2006), leads usa 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 institute in the CFT section, compared to those constitute in sections S1−S4, prove differences in length of the examination and in number of costae (radiating from basal spine to concluding bedroom), which become thinner and more anastomized (Figure 6).

Lenticulina calcar
Marginulinopsis costata
(Figure 4, Effigy  5.6-5.vii) showed similar trends forth the 22 lowermost levels, with irregular, very arable to abundant occurrences between WFE 1 and WFE 2, common abundances (WFE 2, from Levels 12 to 15), and very abundant occurrences higher up WFE 2 (between Levels 15 and 22).

The RA of
Marginulinopsis costata
was abundant and common in Department S2; in the WS section, the species was common beneath and above the skeletal remains, becoming abundant in samples closest to the bones (WFE two). In Section S1, information technology was abundant overall and dominated the assemblages whereas, in Section S4, its RA decreased from common to rare in the uppermost sample. The behavior of
Grand. costata
is non articulate: with a shallow infaunal lifestyle, it prefers soft sediments and reaches a big size (over 6 mm), testify that food-rich sediments are an ideal habitat for this opportunistic species.

Lenticulina calcar, an epifaunal detritivore that prefers cold water, showed 4 peaks of abundances (Figure 4): at the base and height of Section S3 (coinciding with WFE 1), in the WS samples (close to the skeleton remains), and in the terminal three samples from Department S1.
Lenticulina calcar
has been reported in several other marine shoreface to offshore deposits in southwestern Umbria, while
Chiliad. costata
is a typical inhabitant of offshore deposits (Bizzarri et al., 2015).

Analogous distributions were observed in the CFT section in which benthic communities were dominated by abundant to very abundant
Bigenerina nodosaria, Siphotextularia concava, Bannerella gibbosa, Vaginulina striatissima, Marginulinopsis costata,
Lenticulina calcar
(Figure 4).

Subsidiary species.
Other taxa were also noted, although their abundance was low and their occurrence was irregular across the 6 sections. Among them,
Quinqueloculina seminula
was mutual below and to a higher place the bones in the WS section. In Sample 3 of Section S1, species of the
(Ammonia parkinsoniana, A. beccarii, A. tepida, A. perlucida)
were mutual, as was
Bolivina spathulata
Bulimina spinata. In all samples from S1,
Uvigerina mediterranea, a shallow infaunal cold-h2o detritivore, was mutual. Conversely,

Bulimina marginata
was mutual from Sample 4 to the elevation of the section.

Few differences were noted in the CFT section, where the epifaunal herbivorous species
Asterigerinata mamilla
and the suspension feeders
Heterolepa floridana
Lobatula lobatula
occur commonly. The species
B. spathulata, B. marginata, B. spinata, Cancris auriculus, Cassidulina laevigata, Melonis barleanum, Nonionella turgida, and
U. mediterranea
were rare and sporadic.

This state of affairs is noteworthy because the lifestyles of
(Murray, 1991), equally a shallow infaunal herbivore (Ammonia) and a shallow infaunal detritivore (Bulimina), are linked by like behavior that leads both species to colonize burrows where they feed upon bacteria on the couch walls. Jorissen et al. (1992) reported
B. marginata
as an opportunist species found as many equally iv cm below the surface of the sediment and able to respond to loftier food availability.


s figure7All decapod crustaceans institute at the Bargiano site came from the WS section and mainly belonged to two species:
Albaidaplax ispalensis
Garassino, Pasini and Castro, 2013 and
Chlinocephalus demissifrons
Ristori, 1886 (Figure seven; see Appendix 4 and Appendix 5 for taxonomic descriptions; run across also Karasawa and Kato, 2003, and Schweitzer et al., 2010). They were accompanied by the recently described crab
Astenognathus alleronensis
Pasini, Garassino and De Angeli, 2017 (family Varunidae), the outset written report of this genus in the Pleistocene worldwide, by fragments of

sp. and
sp., and by undescribed crustacean burrows (Pasini et al., 2017, p. 69). Nosotros notation that the Bargiano specimens of
A. ispalensis
were larger both than the type series and other specimens reported from Italia and as well bear a poorly distinct and obsolete tooth on the anterolateral margins.

In comparing to Ristori’due south specimen and the specimen reported by Garassino et al. (2004, p. 275: figures. 15, sixteen),
Chlinocephalus demissifrons
from Bargiano showed some variations in the outline of the carapace (which was more than elliptic in shape), fewer raised transverse carinae, and decoration that was more pitted dorsally and on the P1. This carapace variability is likely due to growth stages, as has been noted among several extant representatives of the Euryplacidae (Castro and Ng, 2010, p. 6). These differences could likewise propose, however, the presence of a local “blazon” not withal recognized in fossil species or they may reflect small normal variations. In whatsoever case, the differences in our stance are too inconsistent to allow u.s.a. to describe a new taxon for the Bargiano specimens.


The entire Allerona area represents a unique paleoenvironment in which several WFE took place over the bridge of about one hundred thousand years (Monaco et al., 2014), with a likely increase in WFE frequency in the uppermost sequence. These values, too as the sedimentation rates reported below, are derived from the general stratigraphic reconstruction at the calibration of the basin (Baldanza et al., 2011; Bizzarri et al., 2015). Because the Bargiano site alone, in a stratigraphic section of about five m in thickness and across the 22 levels analyzed, a total of three WFEs were recognized. If the estimated sedimentation rate of 1m/10ky is right, stratigraphic and sedimentological constraints imply that they probably occurred in an interval of virtually 50,000 years, with a frequency of about 18−20 k.y.

Regarding the presumed depth of these events, some information is provided by foraminifers.
Orbulina universa, Globigerinoides ruber,
Globigerinoides sacculifer
are all symbiont-bearing species,
Globigerina bulloides

Turborotalita quinqueloba

are surface dwellers and are consistently full-bodied in the surface layer at an average depth of <50 k, and
Globorotalia inflata
is a surface-subsurface dweller at an average depth of betwixt l m-100 grand (Kucera, 2007; Rebotin et al., 2017).

These fluctuations in abundance appear related to body of water-surface temperature and nearly likely, for symbiont-begetting taxa, to the depth of light penetration. Mud particles dispersed in seawater heavily influence light penetration and, for the Bargiano-Allerona surface area where clay and silt were regularly supplied past river-mouth discharge, the waters were surely less transparent and less able to exist penetrated by light. This datum is aligned with what would be expected at intermediate depths, and our approximate of a depth of 100 m−150 g remains viable. In accordance with these values, the Bargiano site—and the entire Allerona surface area more generally—represent a meaning point of reference for the written report of relatively shallow fossil WFEs.

In society to suggest a comparison, available data on benthic foraminifera and crustacean associations from both fossil and present day WFEs are synthesized in Appendix one. It is clear that data on benthic foraminifers are fewer (with regard to both fossil and present-day events) than those related to crustacean communities, which are oftentimes documented in present-twenty-four hours events. Furthermore, the fossil tape of benthic foraminifera and decapod crustacean communities associated with WFEs is deficient and poorly reported (Appendix 1).

Concerning the benthic foraminifera associations reported from fossil WFEs (Appendix 1), few data exist. Amano and Little (2005) identified
Martinottiella communis

Spirosigmoilinella compressa
as biostratigraphic markers of the Miocene Chikubetsu Fm. in which remains of a archaic mysticete were found, only they were not directly connected with the skeleton. Amano et al. (2007), working with undetermined cetacean remains from Miocene deposits in Japan, reported large (~300 μm diameter) benthic biloculine miliolid foraminifera.

More than information come from present-day WFEs. Wada et al. (1994) described how sediments rich in fatty acids, collected below a whale skeleton on the Torishima seamount (S of Japan), hosted a foraminifera aggregation characterized by
Cassidulinoides parkerianus
‘Rhabdammina’ (?=Saccorhiza) ramosa, Reophax scorpiurus, Recurvoides parkerae, Cystammina pauciloculata, Textularia kattegatensis,
sp. (all agglutinated species);
Gyroidina quinqueloba, Tosaia hanzawai, Fursenkoina
Melonis pompilioides, and
Epistominella exigua
(all rotalids); and
sp. (a miliolid). The assemblage included both agglutinated and rotalid species, and epifaunal and infaunal genera (following the ecological description reported past Murray, 2006) seem to be represented as. Unfortunately, data regarding the relative abundance of these specimens were not reported, and just
C. parkeriana
was characterized as ascendant.

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Rathburn (unpublished information in Gooday and Rathburn, 1999, p. 199) reported that “whale falls favor common species (e.g.,
Uvigerina peregrina

spp.), which are typically plant in eutrophic atmospheric condition”; the authors as well believed “it is possible that a succession of foraminiferal species may respond to the unlike phases of the carcass decomposition procedure.”

Although the Bargiano foraminifera assemblages differ from those reported from nowadays-24-hour interval WFEs, some similarities can be identified. These largely involve families Textularidae and Eggerellidae and, more by and large, the occurrence of agglutinated species. At Bargiano, detritivore epifaunal species
(Lenticulina, Cancris, Cassidulina, Siphotextularia, Gyroidina,
and shallow infaunal species
(Bannerella, Fursenkoina, Bolivina, Nonionella, Melonis, Bulimina, Elphidium)
dominate, while epifaunal herbivore and break feeders and the shallow infaunal omnivore species are subordinate. Compared to information from Wada et al. (1994) the Bargiano benthic foraminifera assemblage shows comparable organization, although ecology conditions (4,037 m in the Wada group’s work) and sea bottom sediments are quite unlike. At Bargiano, moreover, the influx of organic matter was non linked to a single whale fall but was repeated and more than-or-less continuous over time.

Opposite to decapods, distributed mainly at and around WFE 2, benthic foraminifers were influenced past all the Bargiano WFEs. First, the overall reduced number of taxa could point stressful ecology weather, whereas the variability in SD demonstrates that the highest concentration of organic matter in sediments close to the whale skeleton became a limiting factor for the development of benthic taxa and for epifaunal taxa in particular. The shallow infaunal species
Bigenerina nodosaria, Bannerella gibbosa, Marginulinopsis costata,
5. striatissima; the epifaunal species
Lenticulina calcar, and

Siphotextularia concava

dominated the poor assemblages, suggesting that food enrichment was, conversely, favorable for them. As opportunistic species, they responded rapidly to short-term favorable circumstances past increasing their numbers; given their consistent presence in assemblages (across all 25 levels), it is possible to hypothesize that they were sedentary and carried out the life bike through time, increasing in number and maintaining a stable population on the seabed in response to an increase in organic affair.

As reported by Smith et al. (2015), the dispersion of whale biomass leads to a “bull’s eye” of elevated microbial enrichment and degradation activity in the sediment, which declines to background atmospheric condition across about ten m. Goffredi and Orphan (2010) also indicated that the sphere of influence of whale-fall-derived nutrients expanded laterally upwardly to 20 one thousand away, with changes in bacterial diversity. Co-ordinate to this evidence, our analyses of samples from the half dozen short sections (Effigy 1, Figure 2) located at varying distances from the whale carcass and close to fossil ambergris (cololites), highlight changes in benthic foraminifera assemblages and place ascendant taxa continued both to bacterial growth and to nutrient accumulation in the sediment. The increase of nutrients, relative to whale biomass dispersion and evidenced by frequencies of opportunistic benthic foraminifera, seems to confirm dispersion laterally around carcasses for at to the lowest degree x 1000. Nosotros would add together that the repeated overlap of WFE over time facilitated the persistence of opportunistic taxa on the sea bottom, some of which causeless specialized forms, and favored their reproduction (as in the CFT section, for case).

The data reported here demonstrate that at to the lowest degree six benthic foraminifera taxa were endemic to fossil WFEs at Bargiano and that these taxa multiplied (through asexual multiple fission: Murray, 2006) in response to increased availability of food associated with a surge in organic matter. In the Mediterranean Sea, De Rijk et al. (1999, 2000) reported that the bathymetrical distribution of the chief benthic foraminifera taxa was controlled by the flux of labile organic affair to the sea flooring and that the upper and lower depth limits of several taxa corresponded closely to estimated flux isolines. In our example, the isolines corresponded to the WFEs (WFE ane−three).

The scarcity of information on foraminifera at shallow WFEs, however, contrasts with detailed research regarding benthic foraminifera communities in similar contexts, such as those associated with cold seeps (Levin, 2005, and references therein) at which the interactions of microfauna with flow, chemistry, and microbe development is more conspicuously shown.

As indicated by Levin (2005), the composition of foraminifera communities at seeps and their relationship to environmental factors have largely been analyzed in diverse geographic settings (the Gulf of United mexican states, Central and Northern California, and off the coast of Nippon). Differences noted in densities for reference genera and species bespeak the biogeographic variation of communities, just well-nigh seep genera are also characteristic of other depression-oxygen and organic-rich settings (Rathburn et al., 2000, 2003; Bernhard et al., 2001; Robinson et al., 2004). There is little information about seep effects on diversity, merely studies have shown that diversity is reduced in seep sediments with respect to non-seep sediments (Robinson et al., 2004). Likewise, the vertical distribution of foraminifera varies with seepage, although the majority of seep species are considered infaunal. Infaunal foraminifera species tolerate low-oxygen and organic-rich weather (Rathburn et al., 2000) and thrive at varying maximum depths according to habitat (bacterial mats vs. mollusk beds, for example), with subsurface peaks between 2 cm and iv cm. Species characteristic of shallow (120 g) methane seeps in the Santa Barbara Basin off California include
Bolivina tumida, Epistominella pacifica, Oridorsalis umbonatus,
Uvigerina peregrina. Central and northern California seeps also support loftier densities of
Chilostomella, Globobulimina, Nonionella, Cassidulina,
(Bernhard et al., 2001; Rathburn et al., 2003).

It is noteworthy that five genera found by Bernhard et al. (2001) and Rathburn et al. (2003),
Bolivina, Uvigerina, Nonionella, Cassidulina,
Textularia, are among those nowadays as subsidiary taxa in Bargiano shallow whale-fall sediments; among these,
Uvigerina, Siphotextularia
(comparable to
Textularia), and

, highlight an affinity for the kind of paleoenvironment institute at Bargiano. The strong similarity between the two contexts (seeps and whale falls) is evident: both environments induce changes in associations (reducing diversity), and taxa are all infaunal opportunists, characteristics of oxygen-poor or organic-rich environments.

Regarding the decapods found at Bargiano,
Albaidaplax ispalensis
has previously been reported from the Pliocene and early on Pleistocene of Andalusia (Spain) and Tuscany (Italy) (Garassino et al., 2013, p. 363). This finding represents the showtime record of
in the Umbria Region and the second in the early Pleistocene from Italy.
Chlinocephalus demissifrons
has been reported from three Pliocene localities in northern Italy: Fornaci (Savona, Liguria) (Ristori, 1886); Biella (Piedmont) (Garassino et al., 2004); and Castellarano (Emilia Romagna) (Pasini and Garassino, 2013). Garassino et al. (2004), moreover, pointed out that the specimen assigned by Ristori (1891) to
Titanocarcinus sculptus
Ristori, 1891, from the Pliocene of Mucigliani, Siena (Tuscany, central Italia) could be a juvenile phase of

C. demissifrons
. Based upon Italian reports known to date, the Bargiano specimens stand for the starting time report of this species from Umbria, and the get-go in the early Pleistocene, extending the distribution and stratigraphic range of the species into the paleo-Mediterranean Basin. These data add to knowledge of decapod communities in or near fossil WFEs (Appendix ane).

The nearly complete listing of brachyuran (Crustacea, Decapoda) venereal was provided by Fujiwara et al. (2007: effigy 2b), who used high-definition Television receiver video over three years to written report and report on crustacean creature associated with sperm-whale falls in the waters of Cape Nomamisaki (SE Nippon) at depths of 219−250 m. They listed, amid other crustaceans, the following brachyuran crabs, assigning them to families, equally follows:
Homola orientalis
Henderson, 1888 (Homolidae);
sp. (Dorippidae);

Cryptocnemus obolus
Ortmann, 1893, and
Merocryptus lambriformis
A. Milne-Edwards, 1873 (Leucosiidae);
Macrocheira kaempferi
(Temminck, 1836),

Pugettia minor
Ortmann, 1893,
Oxypleurodon stimpsoni
Miers, 1886 (Majidae);
Trachycarcinus sagamiensis
Rathbun, 1932 (Atelecyclidae);
Cancer gibbosulus
(De Haan, 1833-1849) and
C. japonicus
Ortmann, 1893 (Cancridae);

Carcinoplax surugensis
Rathbun, 1932 (Goneplacidae); “
Medaeus serratus”

Sakai, 1965 (Xanthidae); and
sp. (Pinnotheridae); see Fujiwara et al. (2007: table iv).

Lundsten et al. (2010b: figure 5D) reported
Chionoecetes tanneri
Rathbun, 1893 (Oregoniidae) every bit feeding on a whale tissue (site reported every bit Whale-1018) from the deep floor of Monterey Coulee, California (U.s.). Based upon observations of extant brachyuran crabs, the written report of
(both Goneplacidae) could exist strictly correlated with feeding behavior, as attested by the extant

. Regarding decapods from fossil WFEs, Lancaster (1986) and Nesbit (2005) reported the occurrence of
sp. in Eocene sediments (associated with archaeocete bones), and of
Callianopsis clallamensis
(Withers, 1924) in late Oligocene deposits (with mysticete remains), respectively.

Only a few generic reports have considered fossil whale-fall localities in Italia. Danise and Dominici (2014, p. 236, table i) noted that “decapods are also reported” among the creature associated with the fossil basic of cetaceans from the Pliocene of Orciano Pisano and Ponte a Elsa (Tuscany), and Castellarano (Emilia Romagna), Italy. The decapods observed, just non collected, at Ponte a Elsa were associated with the skeleton of a big baleen whale and possibly referable to
Goneplax. The specimens observed at Orciano Pisano, conversely, were closely associated with a cetacean skeleton and were referable to
(Brachyura, Eriphiidae). Unfortunately, due to their frail nature, the specimens were cleaved during the field work that was mainly focused on recovery of the fossil skeleton (Dominici, personal commun., 2016).

As a consequence, the specimens from Bargiano stand out as the only confidently identified fossil brachyuran reported from a fossil whale-fall environment. The presence of several specimens nearby the carcass of the cetacean can be interpreted as opportunistic behavior past these decapod crustaceans due to the presence of a keen, temporary availability of food on the ocean lesser.

Unfortunately, the fossil evidence does not allow to determinate confidently the straight interaction between decapod crustaceans and the cetacean remains. Their presence around the bones might reveal a temporary assemblage of opportunistic scavenger venereal due to the presence of a copious source of food for a relatively long menstruum. The Bargiano specimens, however, were collected just in proximity to whale bones and were evidently absent-minded in the others layers of the outcrop, where but a few, poorly preserved decapods of other species have been nerveless (Garassino and Pasini, personal obs., 2016).

Unlike WFE2, the WFE1 and WFE3 lacked skeletal remains and
cololites are the only evidence of sperm-whale-carcass accumulation on the sea floor. In fact, rare, unidentifiable bone fragments were establish all around these masses (Baldanza et al., 2013; Monaco et al., 2014). A large area around the study site was modified past homo intervention about 30 years ago, when about three m of soil were removed to reveal the marine deposits, accelerating the formation of deep runoff furrows (Effigy one.two). Part of the basic may well have been lost, then, specially if they were reduced to fragments. Such fragmentation probably had a biological origin; the devastation of lipid-rich sperm-whale bones, for example, could have been activated by boring worms such equally the extant
As reported by Higgs et al. (2012), evidence of
activity on fossil whale basic was documented in Pliocene deposits of Orciano Pisano, Tuscany. Therefore, the presence and activity of
in the Allerona WFE surface area during the Pleistocene cannot exist excluded.

Time to come enquiry will include documenting the presence and activeness of marine worms and, more generally, of chemosymbionts, on fossil bones, which at the moment are in the preliminary phase of restoration.


The paleontological heritage of the Bargiano site, where a cetacean skeleton and more than than 27 fossil ambergris structures (linked to the presence of sperm-whale carcasses, Figure 1.2, ane.four-5) have been plant, represents a window onto a shallow (100 m−150 g) early Pleistocene marine paleoenvironment that has proven ideal for an investigation of the relationships among opportunistic micro- and macrofauna in WFEs. The information regarding foraminifera communities and decapods recovered from Bargiano make articulate that a locally high nutrient flux significantly influenced biota in the start few millimeters or centimeters of sea flooring sediment. This increase in nutrients, related to whale-carcass biomass and evidenced by frequencies of opportunistic benthic foraminifera, appears to confirm lateral dispersion effectually the carcass for a radius of at least 10 m.

The entire Allerona expanse represents an unusual paleoenvironment where several WFEs took place over a span of well-nigh 100,000 years (Monaco et al., 2014) with a likely increase in frequency of WFEs during the last fifty one thousand.y. The information presented here allow us to recognize, in a 5-chiliad-thick stratigraphic department analyzed at a total of 25 levels, a total of three WFEs. If these events occurred as multiple cetacean expiry events, the shallow sea floor environment was enriched in organic matter, allowing opportunists (micro- and macrofauna, scavengers, and bacteria) to proliferate steadily.


We wish to thank G. Teruzzi, Museo di Storia Naturale di Milano, for the photos of crab specimens; L. Bartolucci, Section of Physics and Geology (Perugia University), for SEM photos of foraminifers; and S. Dominici, Museo di Storia Naturale di Firenze, for useful discussion and suggestions concerning whale falls in Tuscany. We are also grateful to the students of the Department of Physics and Geology and the Section of Chemistry, Biology and Biotechnology (Perugia Academy) for their contributions to paleontological excavations (June-July 2016) and for their recovery of decapod specimens. Finally, we wish to thank J. Luque, Northward. Higgs, and Thou. Hyžný for their suggestions for improving the manuscript, and W. Ricketts for the accurate revision of the English language text.


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How Does a Whale Fall Community Affect Ocean Sediment