Deep-Sea Benthic Foraminiferal Recolonisation Following A

Marine Micropaleontology 44 (2002) 57^76 www.elsevier.com/locate/marmicro

Deep-sea benthic foraminiferal recolonisation following a volcaniclastic event in the lower Campanian of the Scaglia Rossa Formation (Umbria^Marche Basin, central Italy)

S. Galeotti a;b;Ã, M. Bellagamba a, M.A. Kaminski c;d, A. Montanari e

a Istituto di Geologia dell’Universita', Campus Scienti¢co, Localita' Crocicchia, 61029 Urbino, Italy b Centro di Palinologia dell’Universita', Campus Scienti¢co, Localita' Crocicchia, 61029 Urbino, Italy c Research School of Geological and Geophysical Sciences, Birbeck College and University College London, Gower Street, London WC1E 6BT, UK d KLFR, 3 Boyne Avenue, Hendon NW4 2JL, UK e Osservatorio Geologico di Coldigioco, 62020 Frontale di Apiro, Italy

Received 1 February 2000; received in revised form 23 July 2001; accepted 23 July 2001

Abstract

We studied the distribution of deep water agglutinated foraminiferal (DWAF) assemblages across a 15-cm-thick volcaniclastic layer in the lower Campanian Scaglia Rossa limestones of the Umbria^Marche Basin. Above the volcaniclastic layer, which is devoid of foraminifera, a remarkable pattern of recovery among DWAF has been observed. The complete recovery of DWAF in terms of trophic groups and complexity of assemblages is observed in the first 5 cm above the volcaniclastic layer, representing 4.8 kyr based upon the mean sedimentation rate of the Campanian Scaglia Rossa Formation. In its initial stage, the recovery pattern is remarkably similar to that observed following the 15 June 1991 Mount Pinatubo ashfall in the abyssal South China Sea where various species of Reophax, a small organically cemented species of Textularia, and the calcareous species Quinqueloculina seminula and Bolivina difformis are the earliest recolonisers on top of the tephra layer. Such similarities between modern and fossil analogues strengthens the reliability of environmental reconstructions based on DWAF. ß 2002 Elsevier Science B.V. All rights reserved.

Keywords: Benthic foraminifera; Volcanic ash; Substrate disturbance; Biotic recovery

1. Introduction prerequisite for understanding the processes re- lated to disequilibrium in deep-sea benthic com- Information on life history and community munities, which constitute a key component of the structure of deep-sea benthic foraminifera is a Earth’s largest biotope. Recolonisation of deep- sea substrates, in particular, is a topic that has received growing interest in recent years with sev- * Corresponding author. eral studies on both modern and fossil benthic E-mail address: [email protected] (S. Galeotti). foraminiferal communities following major envi-

MARMIC 851 10-4-02 58 S. Galeotti et al. / Marine Micropaleontology 44 (2002) 57^76 ronmental disturbances (see Alve, 1999 for a re- (2001), who documented the initial stage of review of the topic). colonisation on top of the tephra layer deposited In the fossil record, attention has recently fo- in the South China Sea as a result of the 1991 cused on recolonisation patterns by benthic fora- eruption of Mount Pinatubo in the Philippines. minifera following global catastrophic events such The initial observations suggest that faunal re- as the Cretaceous Oceanic Anoxic Events (Kuhnt, covery (largely by species of Reophax, Subreo- 1992; Coccioni et al., 1995; Peryt and Lamolda, phax, and a minute organically cemented species 1996), the Cretaceous^Tertiary Boundary event of Textularia) had begun as early as 3 years fol- (Kuhnt and Kaminski, 1993, 1996; Coccioni and lowing the eruption. Galeotti, 1994, 1998; Speijer and van der Zwaan, The process of faunal decimation and recovery 1994, 1996; Galeotti, 1998), and the Palaeocene^ following tephra falls must undoubtedly be a Eocene Boundary event (Kaminski et al., 1996; common occurrence in the geological record, con- Speijer et al., 1995, 1997). Successive recolonisa- tributing to local di¡erences in the patch structure tion of the sea £oor has also been suggested to and successional patterns observed in deep-sea explain the patterns observed in the vertical suc- benthic foraminifera. However, a major unknown cession of ‘£ysch-type’ foraminiferal assemblages question is whether or not this process leaves an within turbidite sequences in alpine and boreal observable fossil record, and if so what the time- regions (Gru«n et al., 1964; Butt, 1981; Verdenius scale of faunal recovery in the deep sea is. In this and Van Hinte, 1983; Kuhnt and Kaminski, paper we address these questions by documenting 1989). the pattern of the recolonisation through a high In the modern ocean, recolonisation by benthic resolution study of deep water agglutinated fora- foraminifera has been observed in shallow water miniferal (DWAF) assemblages from below, with- environments following arti¢cial disturbance (El- in, and above a 15-cm-thick volcaniclastic layer in lison and Peck, 1983; Schafer, 1983), in the deep a deep-sea sequence of the Scaglia Rossa Forma- sea following bottom current disturbance (Kamin- tion exposed in the Furlo Gorge (Umbria^Marche ski, 1985), and after local volcanic ashfalls (Finger Apennines, central Italy). and Lipps, 1981). In an experiment using recolonisation trays at an abyssal site in the Panama Basin, Kaminski et al. (1988a,b) identi¢ed several 2. Geological and stratigraphical setting species of benthic foraminifera as opportunistic, including Psammosphaera, Reophax excentricus The upper Turonian^middle Eocene Scaglia and Reophax dentaliniformis. Rossa Formation in the Umbria^Marche Basin Deep-sea basins that experience seasonal dys- consists of regularly bedded pink and reddish oxia also support communities of opportunistic limestones interbedded with reddish marly layers benthic foraminifera. In a study of the deep deposited under well oxygenated conditions in a San Pedro Basin o¡ southern California, Ka- lower bathyal depositional environment (Arthur minski et al. (1995) reported a faunal assem- and Fischer, 1977; Kuhnt, 1990). The faunal and blage consisting of Psammosphaera, Reophax, £oral associations of the Scaglia Rossa mostly and a minute organically cemented species of consist of calcareous nannofossils and subordi- Textularia that was interpreted as opportunistic. nate planktonic foraminifera along with rare Many of the species were the same as those benthic foraminifera (mainly DWAF). The found in the recolonisation trays in the Panama DWAF assemblages in these limestones are Basin, suggesting that di¡erent types of distur- highly diversi¢ed and are unique within the bance may result in similar benthic communities. Upper Cretaceous. They include elements of The sole observations of in situ recolonisation by mixed calcareous and organically cemented bathy- modern deep-sea benthic foraminifera in a vast al assemblages, purely agglutinated ‘£ysch-type’ disturbed habitat were carried out by Hess and assemblages, and a number of species known Kuhnt (1996), Hess (1998), and Hess et al. from Upper Cretaceous sequences deposited be-

MARMIC 851 10-4-02 S. Galeotti et al. / Marine Micropaleontology 44 (2002) 57^76 59

Fig. 1. Location map of the studied section. low the Carbonate Compensation Depth. Ac- 3. Material and methods cording to Kuhnt (1990), the agglutinated foraminiferal assemblage of the Scaglia Rossa is in- To document changes in benthic foraminiferal dicative of a water depth between 1500 m and assemblages and recolonisation patterns above 2000 m. the volcaniclastic layer, a quantitative analysis In outcrops of the lower Campanian Scaglia of DWAF was carried out on acid residues or Rossa sequence at Furlo (Marche region of Italy, washed residues obtained from 18 samples. Sam- Fig. 1), Mattias et al. (1988) described a discrete ples were collected across a 255-cm interval, from 15-cm-thick bentonitic layer. According to the 125 cm below the volcaniclastic layer (FB1 to litho-, magneto- and biostratigraphic investigation FB6), within it (FB7a, b), to 115 cm above it of the Furlo sequence by Alvarez and Lowrie (FB8 to FB17). The interval spanning the ¢rst (1984), this bentonitic layer falls within the Globo- 5 cm above the volcaniclastic layer was sampled truncana elevata planktonic foraminiferal zone, every centimetre (FB8 to FB12). and within the lower part of Chron 33r of early Considering the duration of Chron 33r (Grad- Campanian age. Based on detailed geochemical stein et al., 1995) and its thickness in the Furlo and mineralogical analyses, Mattias et al. (1988) section (Alvarez and Lowrie, 1984), the mean sed- suggested a volcanic origin for this bentonite. The imentation rate during the early Campanian at layer is predominantly comprised of calcium dioc- Furlo was about 8.6 m/Myr. Therefore, assuming tahedric montmorillonite derived from the trans- a constant sedimentation rate and geologically information, in a marine environment, of ¢ne vol- stantaneous deposition of the volcaniclastic layer, canic glass originating from sub-aerial volcanic the studied interval represents approximately 370 activity. The sudden deposition of a 15-cm-thick kyr. Although a hiatus is known to occur within (post-compaction thickness) volcanic ash layer the Scaglia Rossa within the basal Palaeocene would have created a sudden disturbance of the (e.g. Dingus, 1984), there is no visible evidence sea £oor ecosystem, including mass mortality of of hiatuses or turbidites within the studied inter- benthic organisms. This event, therefore, repre- val. sents an interesting case for a high-resolution Limestone samples were soaked in 5% diluted study of the pattern of recolonisation of the HCl, wet sieved with tap water and a 63-Wm benthic community, which in this bathyal envi- screen, and oven dried. The s 63-Wm residues ronment is normally characterised by a highly di- were split with a microsplitter into subsamples versi¢ed assemblage of DWAF. containing approximately 300^400 specimens

MARMIC 851 10-4-02 60 S. Galeotti et al. / Marine Micropaleontology 44 (2002) 57^76

Table 1 Morphology and inferred life position of DWAF genera identi¢ed across the volcaniclastic layer at Furlo

MARMIC 851 10-4-02 S. Galeotti et al. / Marine Micropaleontology 44 (2002) 57^76 61 each, which were all picked, counted, mounted on positionally similar samples were identi¢ed using micropalaeontological slides for quantitative anal- the Bray^Curtis dissimilarity index (Bray and Cur- ysis, and then stored in the collections of the Paly- tis, 1957) in association with the nearest neighbour, nology Center at the University of Urbino. The or single linkage, clustering technique. two samples representing the bentonite layer were washed with tap water through a 63-Wm sieve. We largely adopted species concepts of Kamin- 4. Results ski et al. (1988a,b), Kuhnt (1990), Kuhnt and Kaminski (1990, 1993), Wightman and Kuhnt Results are summarised in Table 2 together (1992), Kaminski and Geroch (1993), and Kuhnt with the mean distance of each sample from the et al. (1998). Specimens not identi¢ed to the ge- bentonitic layer. DWAF assemblages from the neric level are reported as indeterminate forms. surveyed interval are rich and highly diversi¢ed, For each sample, the Fisher-K index and the per- consisting of not fewer than 70 species represent- centages of epifaunal, shallow infaunal, and deep ing 33 genera. A minor degree of uncertainty con- infaunal forms were calculated. The identi¢cation cerning the number of occurring species is due to of epifaunal v. infaunal groups was carried out the presence of specimens not identi¢ed to the using the morphogroup classi¢cation of Jones generic level in some samples. Preservation is gen- and Charnock (1985), Tyszka (1994), Nagy et al. erally good, though one of the samples above the (1995), and Van Den Akker et al. (2000). The bentonitic layer (FB10) exhibits a relatively poor inferred mode of life for each taxon is reported state of preservation. The most common compo- in Table 1. nents of the DWAF assemblage are Paratrocham- Tubular forms occurring in the studied material minoides spp., Trochamminoides dubius, Trocham- (mainly Rhizammina indivisa, Rhizammina algae- mina spp., Subreophax spp., Saccammina spp., formis and Rhabdammina sp.) have been combined and tubular forms (mainly Rhizammina algaefor- into a single group. Since only small fragments of mis). Other important components are Haplo- these delicate forms occur in the studied material, phragmoides spp., Reophax spp., and Ammos- counting them as individual specimens would phaeroidina pseudopauciloculata. overestimate their relative abundance. At the The relative abundance of each species £uctu- same time, considering the numbers of fragments ates greatly in the surveyed interval, and the larg- as counts, as is often the case, a¡ects the relative est £uctuations are observed just above the volca- abundance of the non-tubular taxa. As a compro- niclastic layer (see Table 2 and Fig. 2). The mise, we subdivided the number of fragments by a bentonitic layer itself is virtually devoid of fora- constant value of 15. This number was derived by minifera (both benthic and planktonic). Our ex- considering the minimum number of unbranched amination of a large volume of material collected fragments obtainable from the holotype of R. al- from the lower half of the ash layer revealed just gaeformis (which comprises the largest proportion one specimen of Glomospira serpens, while the of the tubular fragments). Although arbitrary, this samples collected from the upper half of the layer method enables us to obtain an estimate of tubular were completely barren. specimens that is closer to reality than counting Major di¡erences in diversity, community single fragments as specimens. structure, feeding and habitat preferences are ob- Statistical tests, including R-mode principal com- served among benthic foraminiferal assemblages ponent analysis (PCA) and Q-mode cluster analy- below and above the volcaniclastic layer. Cluster sis, have been performed on the untransformed analysis clearly separates two major branches at a data set after exclusion of rare species (those that Bray^Curtis index value of 0.3 (Fig. 3). The ¢rst occur as one or two specimens in fewer than three group consists of four samples, representing the non-adjacent samples) and grouping species that ¢rst 4 cm above the volcaniclastic layer (the ‘re- have a discontinuous, scattered distribution to the colonisation interval’), whereas the second group generic level. For cluster analysis, groups of com- includes all the remaining samples. A further sub-

MARMIC 851 10-4-02 62 Table 2 Distribution of DWAF across the volcaniclastic layer at Furlo Sample FB1 FB2 FB3 FB4 FB5 FB6 FB7a FB7b FB8 FB9 FB10 FB11 FB12 FB13 FB14 FB15 FB16 FB17 Mean distance from the 3125 385 340 315 35 30.5 0 0 +0.5 +1.5 +2.5 +3.5 +4.5 +10 +19,5 +27 +36 +100 bentonite layer (cm) Ammobaculites sp. ^ 0.5 ^ ^ ^ 0.5 ^ ^ ^ ^ 1.5 0.5 ^ ^ ^ 0.9 2.0 0.5 Ammodiscus cretaceus ^ ^ 0.5 ^ ^ 0.5 ^ ^ ^ ^ ^ ^ ^ ^ 1.0 ^ ^ ^ Ammodiscus glabratus 1.7 ^ ^ 1.9 0.9 1.5 ^ ^ 1.4 0.4 0.5 0.5 2.8 1.5 1.0 ^ 0.5 ^ Ammodiscus peruvianus ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ 0.5 Ammodiscus planus 2.6 ^ 0.5 4.9 1.8 2.9 ^ ^ 1.4 1.8 ^ 1.5 1.4 5.8 2.4 0.4 0.5 ^ Ammodiscus sp. ^ 1.0 ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ .Glot ta./Mrn irplotlg 4(02 57^76 (2002) 44 Micropaleontology Marine / al. et Galeotti S. Ammosphaeroidina 7.7 3.0 10.0 8.7 8.5 7.8 ^ ^ 7.1 8.3 0.5 5.8 6.2 3.4 2.9 6.7 12.7 10.2 pseudopauciloculata Aschemocella sp. ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ 0.5 ^ ^ ^ Bulbobaculites 2.1 1.0 1 1.0 1.8 ^ ^ ^ 2.8 1.3 1.0 1.5 1.9 3.4 1.4 4.5 1.0 0.9 problematicus Dorothia sp. 0.4 ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^

AMC8110-4-02 851 MARMIC Glomospira charoides ^ ^ ^ 0.5 ^ ^ ^ ^ 0.5 ^ ^ ^ 0.9 ^ ^ ^ ^ 0.5 Glomospira glomerata ^ 0.5 2.0 1.0 0.4 0.5 ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ 0.5 ^ Glomospira gordialis ^ 2.0 2.5 1.0 6.3 1.0 ^ ^ 1.4 ^ 1.0 2.9 3.8 3.4 2.4 ^ 2.4 1.9 Glomospira serpens 0.4 0.5 ^ 0.5 0.9 ^ ^ ^ ^ 0.4 ^ 0.5 ^ 0.5 ^ 0.4 ^ 0.5 Glomospira spp. 3.0 0.5 ^ 1.0 0.9 2.0 ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ 0.5 0.4 Glomospirella gaultina 2.6 1.0 ^ 2.9 1.3 2.0 ^ ^ 1.4 2.2 0.5 1 ^ 4.9 1.4 0.9 1.0 1.9 Haplophragmoides ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ 0.4 ^ ^ suborbicularis Haplophragmoides walteri 5.1 11.4 4.5 2.9 3.6 2.4 ^ ^ 2.4 7.0 7.0 0.5 ^ 12.6 6.3 4.5 6.3 10.2 Haplophragmoides sp. ^ ^ ^ ^ 0.4 ^ ^ ^ ^ 0.4 1.5 ^ 1.4 ^ ^ 0.4 0.5 0.5 Hippocrepina depressa ^ ^ 0.5 ^ ^ ^ ^ ^ ^ ^ 0.5 ^ ^ ^ ^ 0.4 ^ ^ Hormosina excelsa ^ ^ 1.5 1.0 0.4 0.5 ^ ^ ^ ^ ^ ^ ^ ^ ^ 0.9 ^ 0.5 Hormosina sp. 1.3 0.5 ^ ^ ^ 0.5 ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ Hormosinella cf. distans 3.0 2.5 5.5 1.0 4.0 2.4 ^ ^ 5.7 5.3 7.0 2.4 6.2 4.9 4.3 4.5 1.5 2.3 Hyperammina dilatata ^ ^ ^ 0.5 ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ Hyperammina sp. 0.4 1.5 ^ ^ ^ 0.5 ^ ^ ^ 0.4 ^ ^ 0.5 ^ ^ 0.4 1.5 1.4 Kalamopsis grzybowskii ^ ^ 1.5 2.4 0.4 2.0 ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ 1.0 ^ Karreriella conversa 0.4 ^ ^ ^ ^ ^ ^ ^ ^ 0.4 ^ ^ ^ ^ ^ ^ 0.5 ^ Karrerulina sp. 0.9 4.5 4.5 1.0 5.4 2.0 ^ ^ 2.4 0.9 4.5 10.2 1.4 4.4 5.3 4.5 4.4 6.9 Lituotuba lituiformis 0.4 ^ ^ 1.9 ^ 1.0 ^ ^ 0.5 ^ ^ ^ 2.8 ^ 0.5 ^ ^ 0.5 Lituotuba sp. ^ ^ ^ ^ ^ 0.5 ^ ^ ^ ^ ^ ^ ^ ^ ^ 0.4 ^ ^ Paratrochamminoides ^ ^ 0.5 ^ ^ ^ ^ ^ ^ ^ ^ 1 ^ ^ ^ ^ 0.5 ^ heteromorphus Paratrochamminoides 7.2 9.4 3.0 6.3 10.7 5.4 ^ ^ 3.3 5.7 3.0 4.4 10.4 6.3 13.0 9.4 4.9 6.0 sp. 1 (Kuhnt, 1990) Paratrochamminoides 6.0 5.9 2.5 3.9 6.3 2.0 ^ ^ 2.4 4.4 3.5 3.4 5.2 7.2 12.5 5.8 8.8 6.0 sp. 2 (Kuhnt, 1990) Table 2 (continued) Sample FB1 FB2 FB3 FB4 FB5 FB6 FB7a FB7b FB8 FB9 FB10 FB11 FB12 FB13 FB14 FB15 FB16 FB17 Mean distance from the 3125 385 340 315 35 30.5 0 0 +0.5 +1.5 +2.5 +3.5 +4.5 +10 +19,5 +27 +36 +100 bentonite layer (cm) Paratrochamminoides/ 0.9 ^ 0.5 ^ ^ 1.0 ^ ^ 4.7 4.8 7.5 4.9 ^ ^ ^ ^ 1.0 ^ Trochamminoides juvenile Pseudobolivina lagenaria ^ ^^^^^^^^^^ 0.5^^^^ ^^ Pseudobolivina cf. munda 0.4 0.5 ^ 1.0 0.4 0.5 ^ ^ 6.1 6.1 3.0 3.9 4.3 1.9 1.0 0.4 1.5 0.5 Pseudobolivina variabilis 0.9 ^ ^ ^ ^ ^ ^ ^ 0.5 ^ ^ ^ ^ ^ ^ 0.9 ^ ^ Recurvoides nucleosus ^ ^ 1.0 0.5 ^ 0.5 ^ ^ ^ 0.9 ^ ^ ^ ^ 2.4 ^ 0.5 ^ Recurvoides sp. 0.4 ^ 2.0 0.5 0.4 1.0 ^ ^ 0.9 0.4 1.0 1 ^ 1.0 ^ ^ 1.0 0.5 .Glot ta./Mrn irplotlg 4(02 57^76 (2002) 44 Micropaleontology Marine / al. et Galeotti S. Reophax minutus 3.8 5.4 3.5 3.4 3.6 2.9 ^ ^ 14.6 7.9 8.5 8.7 6.6 2.9 10.1 4.9 6.3 5.6 Reophax subnodulosus ^ 2.5 2.0 ^ ^ 1.0 ^ ^ 0.5 ^ 0.5 ^ 0.9 ^ 1.0 ^ 0.5 0.5 Reophax sp. 5 ^ ^ ^ 1.0 ^ 1.0 ^ ^ 2.4 ^ 1.0 ^ 1.4 ^ ^ ^ 1.0 0.9 (Kuhnt, 1990) Reophax spp. ^ ^ ^ ^ ^ ^ ^ ^ 0.5 ^ ^ ^ ^ ^ ^ ^ 0.5 ^ Rzehakina epigona ^^^ ^^^ ^ ^^0.5^^^^^ ^^

AMC8110-4-02 851 MARMIC Saccammina grzybowskii 3.0 2.5 2.0 2.9 0.9 2.0 ^ ^ 0.9 3.9 1.5 7.3 3.3 2.4 2.4 8.5 0.5 2.8 Saccammina placenta 5.5 6.9 5.0 4.9 5.8 4.4 ^ ^ 4.7 4.4 3.0 5.3 5.2 10.2 6.3 6.7 4.9 3.2 Saccammina sphaerica ^ ^^^ ^^^ ^ ^^^ ^^^^0.4^^ Saccorhiza ramosa ^ ^ 1.0 1.0 ^ 1.0 ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ 0.9 Spiroplectammina dentata 0.4 ^ 1.0 0.5 ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ 0.4 ^ ^ Spiroplectammina laevis 0.9 ^ ^ 1.0 ^ 1.5 ^ ^ ^ 3.1 1.0 ^ ^ ^ ^ 0.4 1.5 ^ Spiroplectammina spp. ^ 0.5 ^ ^ 0.9 ^ ^ ^ ^ ^ 1.0 2.4 0.5 ^ ^ ^ ^ ^ Subreophax guttifer 0.9 2.5 2.5 2.9 3.1 3.4 ^ ^ 2.4 6.1 3.0 1.9 0.9 1.5 0.5 0.9 3.4 2.3 Subreophax pseudoscalaris 0.4 ^ ^ ^ ^ ^ ^ ^ 0.5 1.3 ^ ^ ^ 1.0 ^ ^ 0.5 0.5 Subreophax scalaris 0.4 2.0 1.0 2.4 3.6 0.5 ^ ^ 4.7 2.2 6.5 2.9 1.4 2.4 3.8 1.3 1.5 1.4 Subreophax splendidus 2.1 8.4 4.0 6.3 4.0 3.4 ^ ^ 6.1 2.2 7.0 9.2 5.2 3.4 6.7 6.3 3.9 7.4 Subreophax spp. ^ ^ 1.0 0.5 0.4 2.4 ^ ^ ^ ^ 1.5 0.5 ^ ^ ^ ^ ^ ^ Trochammina deformis 7.2 5.9 8.5 3.9 6.3 5.4 ^ ^ 2.4 1.3 3.0 2.4 4.3 2.4 1.4 3.6 6.3 4.6 Trochammina ^ ^ 0.5 0.5 ^ ^ ^ ^ ^ 0.4 ^ ^ ^ ^ ^ ^ 1.0 ^ cf. gyroidiniformis Trochammina spp. 2.1 4.0 4.0 6.8 4.5 5.9 ^ ^ 1.4 0.4 3.0 ^ ^ ^ 0.5 ^ 0.5 0.5 Trochamminoides dubius 9.4 3.5 5.5 4.4 3.6 8.3 ^ ^ 3.8 8.8 4.5 2.4 10.4 2.9 1.4 9.4 1.0 7.9 Trochamminoides septatus 0.4 ^ ^ ^ 0.4 ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ Trochamminoides ^ ^^^ ^^^ ^ ^^^ ^^0.5^^ ^^ subcoronatus Trochamminoides spp. 3.8 2.5 3.5 3.9 ^ 2.4 ^ ^ 4.2 ^ 2.0 0.5 ^ 0.5 1.4 0.9 2.0 0.5 Tubular forms 6.0 5.9 7.0 4.9 4.0 6.3 ^ ^ 4.7 5.7 6.5 6.3 8.5 5.3 4.3 6.3 7.3 6.0 Turritellella sp. ^ ^ ^ ^ ^ 0.5 ^ ^ ^ ^ ^ 0.5 ^ ^ ^ ^ ^ ^ Uvigerinammina jankoi 4.3 1.0 4.0 1.0 1.3 6.3 ^ ^ 0.5 ^ 1.0 1 0.9 0.5 1.4 0.4 ^ 0.5 Indeterminate forms 1.7 0.5 0.5 1.9 2.7 1.0 ^ ^ 0.9 0.9 1.5 2.4 0.9 2.9 0.5 2.2 2.4 1.4 Number of specimens 235 202 201 206 224 205 0 0 212 228 200 206 211 206 208 223 205 216 63 64 S. Galeotti et al. / Marine Micropaleontology 44 (2002) 57^76

Fig. 2. Vertical distribution of selected DWAF taxa across the volcaniclastic layer at Furlo. Notice the scale break between 0 and 5 cm above the volcaniclastic layer. division of the second cluster enables us to distin- slightly higher proportions of the Subreophax guish samples FB12^FB16. and Reophax minutus assemblage, and lower rela- The ¢rst and second axes of the PCA accounted tive abundance of the Paratrochamminoides^Tro- for 78% and 6% of total variance, respectively. chamminoides^Trochammina assemblage. Reophax Loading scores for the ¢rst two axes are reported minutus, which forms a separate assemblage, is a in Table 3. Based on PCA scores, we identi¢ed relatively common taxon throughout the studied ¢ve assemblages denominated by the dominant interval, but shows a peak in relative abundance taxa (Fig. 4). These display a distinctive distribu- just above the volcaniclastic layer. The Pseudo- tion across the volcaniclastic layer. A comparison bolivina cf. munda assemblage, which consists of of the relative abundance of faunal groups iden- P. munda and small specimens of Paratrocham- ti¢ed by the PCA in the pre-ashfall, ‘recolonisa- minoides and Trochamminoides, is found almost tion’, and post-recovery intervals is given in Fig. 5. exclusively in the ‘recolonisation interval’. A Pre-ashfall and post-ashfall assemblages are third PCA assemblage comprised of Subreophax quite similar, despite displaying some minor dif- scalaris, Subreophax splendidus, Karrerulina spp., ferences in the species composition of the DWAF. and Hormosinella cf. distans shows highest rela- In particular, the post-ashfall assemblages show tive abundance in the ‘recolonisation interval’,

MARMIC 851 10-4-02 S. Galeotti et al. / Marine Micropaleontology 44 (2002) 57^76 65 although not immediately above the volcaniclastic catastrophically by the deposition of over 15 cm layer. of volcanic ash, an event that would have oc- Below the volcaniclastic layer, DWAF assem- curred within a day or two at the lower bathyal blages are characterised by a rather stable epifau- water depths represented by the Scaglia Rossa nal/infaunal ratio, with an average relative abun- (Wiesner et al., 1995). It is possible that some dance of infaunal forms around 22% (Fig. 6). In sediment-dwelling foraminifera reacted to burial this stratigraphic interval, DWAF diversity (Fish- by attempting to exhume themselves, and the spe- er-K) ranges from a minimum value of 10.7 in cies found within the basal centimetre of the ben- sample FB2 to a maximum of 16.5 just below tonite, Glomospira serpens, does possess the typi- the ash layer (FB6), with an average value of cal morphology of an active burrower. However, 13.3. The changes recorded by the PCA assem- the ability of foraminifera to burrow upward blages across the volcaniclastic layer are clearly through volcanic ash is apparently quite limited re£ected in the record of faunal parameters (epi- (see also Hess and Kuhnt, 1996, ¢gs. 3 and 4). faunal vs. infaunal abundance and diversity). The A remarkable pattern that can be interpreted ‘bloom’ in the elongate morphotypes that domi- as evidence of faunal recovery characterised the nate the Reophax minutus, Pseudobolivina cf. mun- sea £oor following the deposition of the volcanic da, and Subreophax assemblages is re£ected in a ash. Species such as Reophax minutus and Pseu- peak of infaunal forms occurring within the ‘re- dobolivina cf. munda, which behaved as oppor- colonisation interval’, with a maximum value of tunists in the recolonisation process, are inter- 46.2% recorded in the ¢rst centimetre above the preted as infaunal deposit feeders. Accordingly, volcaniclastic layer. Just above the volcaniclastic a special feature of infaunal elongate/lanceolate layer, diversity shows average values slightly low- morphotypes and, in particular of the genus er than those observed in the pre-ashfall interval, Reophax and its relatives seems to be their high dropping to its lowest value in sample FB12, just capability for dispersal. Forms belonging to the above the ‘recolonisation interval’. Subreophax assemblage (Subreophax scalaris, Although based on few samples, a gradual re- Subreophax splendidus, Hormosinella cf. distans, covery to average values comparable to that of and Karrerulina sp.), although bene¢ting from the pre-ash interval can be observed upsection. conditions following the deposition of the volca- Unlike the rest of the surveyed interval, a clear niclastic layer, apparently represent ‘second covariance between diversity and the proportion wave’ colonisers. of infaunal forms is observed in the ‘recolonisa- The stratigraphic distribution of PCA assem- tion interval’. blages reveals that the highest abundances of ‘recolonisers’ is restricted to the ¢rst 4 cm above the volcaniclastic layer. This interval corresponds to 5. Discussion 4.8 kyr, assuming a constant sedimentation rate of 8.6 m/Myr. However, this value for the time The DWAF assemblages studied at Furlo are signi¢cance of the disturbed fauna is certainly an generally similar to those reported by Kuhnt overestimate, owing to ubiquitous presence of bi- (1990) from the Upper Cretaceous Scaglia series oturbational mixing in the Scaglia Rossa. The of the classic Bottaccione section, near Gubbio. group most adversely a¡ected by the deposition The taxonomic composition of these assemblages of the volcaniclastic layer appears to be the Para- is indicative of a lower bathyal depositional set- trochamminoides^Trochamminoides^Trochammina ting at a palaeodepth of 1500^2000 m. assemblage. Prior to the deposition of the volcaniclastic layer, the DWAF assemblage is indicative of a 5.1. Comparison with Recent analogues certain environmental stability as indicated by the relatively minor £uctuations in the measured The recovery pattern of the DWAF in the Sca- faunal parameters. This stability was disrupted glia Rossa at Furlo displays remarkable simi-

MARMIC 851 10-4-02 66 S. Galeotti et al. / Marine Micropaleontology 44 (2002) 57^76

Fig. 3. Results of the Q-mode cluster analysis. The dendrogram is produced using the Bray^Curtis similarity measure on untransformed data in association with the nearest neighbour clustering method. Numbers to the right of the dendrogram are samples. larities to those observed following disturbance pered morphology with a ¢nely agglutinated wall, (actual or simulated) in modern deep-sea en- and biserial chamber arrangement. Subsequent vironments. Several species of the genus Reophax monitoring studies of the 1991 Mt Pinatubo teph- have been reported among the early colonisers of ra layer carried out between 1996 and 1999 (Hess, current-controlled substrates at the HEBBLE site 1998; Hill, 1998; Fisher, 1999; Hess et al., 2002) in the northwestern Atlantic (Kaminski, 1985) as have documented an increase in foraminiferal di- well as in a recolonisation tray experiment carried versity and the establishment of additional species out in the Panama Basin (Kaminski et al., that continue to colonise the substrate. The ‘sec- 1988a,b). According to Kaminski et al. ond wave’ of colonisers in the deep South China (1988a,b), Psammosphaera fusca and Hormosina Sea includes species of Subreophax and tubular ovicula are among the species that display good forms such as Rhizammina, while the abundance dispersal capability. Three years after the deposi- of the initial colonists has decreased (Hess et al., tion of volcanic material from the 1991 Mt Pina- 2002). Seven to eight years after the ashfall, the tubo eruption, various species of Reophax (i.e. R. trochamminid group (Trochammina ex gr. globi- scorpiurus, R. dentaliniformis, R. bilocularis), a geriniformis, Trochammina sp., Adercotryma glom- small organically cemented species of Textularia, erata) has increased markedly in abundance, and and the calcareous species Quinqueloculina semi- coiled multichambered forms such as Recurvoides, nula and Bolivina di¡ormis were reported as the Eratidus, Ammobaculites, Haplophragmoides, and earliest recolonisers on top of the tephra layer at Karrerulina are beginning to appear in the sam- stations collected from lower bathyal depths in ples. the South China Sea (Hess and Kuhnt, 1996). It In spite of the large age di¡erence between our is noteworthy that this species of Textularia locality and the modern faunas, it is remarkable shows remarkable morphological similarity to that the recolonisation pattern observed above the our Pseudobolivina cf. munda, which recolonised volcaniclastic layer at Furlo is entirely compara- the ¢rst 2 cm of sediment above the bentonite in ble (at least on a generic level) with patterns ob- the Scaglia Rossa at Furlo. Both species are char- served on modern deep sea substrates. In line with acterised by their small size and a lanceolate/ta- observations on modern assemblages, the genera

MARMIC 851 10-4-02 S. Galeotti et al. / Marine Micropaleontology 44 (2002) 57^76 67

Table 3 Loading scores of identi¢ed taxa on the ¢rst two axes of the PCA Taxon Loadings First axis Second axis 1 Ammobaculites sp. 0.018 0.021 2 Ammodiscus spp. 0.135 30.111 3 Ammosphaeroidina pseudopauciloculata 0.333 30.281 4 Bulbobaculites problematicus 0.083 0.037 5 Glomospira spp. 0.160 30.204 6 Glomospirella gaultina 0.076 30.042 7 Haplophragmoides walteri 0.263 30.005 8 Haplophragmoides sp. 0.016 0.019 9 Hormosina spp. 0.021 30.076 10 Hormosinella cf. distans 0.187 0.158 11 Hyperammina spp. 0.022 30.030 12 Kalamopsis grzybowskii 0.019 30.081 13 Karrerulina sp. 0.188 0.151 14 Lituotuba lituiformis 0.022 30.033 15 Paratrochamminoides sp. 1 (Kuhnt, 1990) 0.337 30.116 16 Paratrochamminoides sp. 2 (Kuhnt, 1990) 0.265 30.006 17 Paratrochamminoides/Trochamminoides juvenile 0.071 0.341 18 Pseudobolivina cf. munda 0.094 0.288 19 Recurvoides spp. 0.045 0.007 20 Reophax minutus 0.296 0.523 21 Reophax subnodulosus 0.026 30.015 22 Reophax sp. 5 (Kuhnt, 1990) 0.024 0.037 23 Saccammina placenta 0.262 30.037 24 Saccammina sphaerica 0.145 0.069 25 Saccorhiza ramosa 0.011 30.041 26 Spiroplectammina spp. 0.050 0.025 27 Subreophax guttifer 0.113 0.023 28 Subreophax pseudoscalaris 0.013 0.013 29 Subreophax scalaris 0.111 0.224 30 Subreophax splendidus 0.254 0.201 31 Subreophax spp. 0.016 30.020 32 Trochammina deformis 0.207 30.304 33 Trochammina spp. 0.101 30.220 34 Trochamminoides dubius 0.270 30.179 35 Trochamminoides spp. 0.087 30.064 36 Tubular forms 0.285 30.017 37 Uvigerinammina jankoi 0.070 30.188 The ¢rst axis explains 78% of the total variance and the second axis explains 6% of the total variance.

Reophax, Hormosinella, and Pseudobolivina are in the ¢rst few centimetres above the bentonitic indicative of recolonisation following a major layer (see Table 2 and Fig. 2). It is possible that substrate disturbance. these forms, which presumably had an epifaunal An additional indication of stressed environ- to shallow infaunal position in the substrate, were mental conditions following the deposition of not able to reach the adult stage owing to the lack the volcaniclastic layer comes from the high of food resources on a comparatively sterile vol- proportion of juvenile and/or dwarfed specimens caniclastic substrate or because of competition for of the Paratrochamminoides^Trochamminoides resources from the established opportunists living group, which forms part of the Pseudobolivina on the sediment surface. Infaunal foraminifera cf. munda PCA assemblage and which is observed would in fact have no reason to colonise the ash

MARMIC 851 10-4-02 68 S. Galeotti et al. / Marine Micropaleontology 44 (2002) 57^76

Fig. 4. Plot of taxa scores on the ¢rst two axes of the PCA. The ¢rst and second axes explain 78% and 6% of total variance, respectively. Numbers refer to taxa as shown in Table 3. Five assemblages (encircled) are identi¢ed and denominated by the dominant taxa. layer itself because of a lack of food (in the form fragmentation just above the volcaniclastic layer. of bacteria). Fragments of Rhizammina indivisa collected in the The unchanged proportion of tubular forms ¢rst few centimetres above the volcaniclastic layer across the volcaniclastic layer seems to be at are smaller than those observed in the rest of the odds with the ¢ndings of Kaminski (1985) and studied stratigraphic interval. A possible explan- Kaminski et al. (1988a,b), who found that tubular ation for the increased fragmentation of Rhizam- epifaunal forms such as Dendrophrya have the mina above the volcaniclastic layer is scavenging lowest dispersal capability. Likewise, these forms by bioturbators, which following the deposition of are slow at recolonising the 1991 Mt Pinatubo ash the sterile volcaniclastic layer must have been con- layer in the South China Sea (see data tables in centrated near the sediment^water interface. Tub- Hess, 1998). The proportion of tubular forms in ular foraminifera are also more likely to be scav- the ‘recolonisation interval’ (with the exception of enged by surface-grazing macrofaunal burrowers sample FB9) is only slightly lower than that ob- than are infaunal species. Burrows ¢lled with tub- served in the pre-ashfall and post-recovery fauna. ular foraminifera have been observed in boxcores However, the relative abundance of tubular fora- from the South China Sea (Kaminski and Wetzel, minifera is certainly overestimated in these sam- unpublished data). ples because of poorer preservation and increased Five centimetres above the volcaniclastic layer

MARMIC 851 10-4-02 S. Galeotti et al. / Marine Micropaleontology 44 (2002) 57^76 69

thickness prior to compaction was probably much higher) sealed o¡ the entire benthic fauna. Studies of the 1991 Mt Pinatubo ash layer reveals that an ash thickness in excess of about 4 cm results in total mortality of the benthic foraminifera within a few years, as oxygen becomes de- pleted beneath the ash layer. The bentonite layer at Furlo contains dispersed pyrite crystals, indi- cating a reducing environment (Mattias et al., 1988). At Furlo, the mechanism responsible for total mass mortality would have been burial and subsequent oxygen depletion of the pore waters, which would have killed o¡ any surviving deep Fig. 5. Distribution of PCA assemblages across the volcani- infaunal species. clastic layer. Pre-ashfall values are calculated as a simple In terms of the recolonisation mechanism, the average of data from sample below the volcaniclastic layer. lateral migration of initial recolonisers from areas Likewise, post-recovery values represents an average of rela- una¡ected by the ashfall seems most appropriate. tive abundances recorded from samples above the ‘recoloni- It is interesting to note that the species which sation interval’ (i.e. the ¢rst 5 cm above the volcaniclastic layer). P-T-T = Paratrochamminoides^Trochamminoides^Tro- recolonised the substrate after the ashfall all be- chammina assemblage. S-R-H = Saccammina^Rhizammina^ long to the deep infaunal morphogroup, implying Haplophragmoides assemblage. The relative abundance of that they might not be easily transported by bot- rare and indeterminate species (reported as R/I) is also re- tom currents or similar mechanisms. However, ported. the elongate/lanceolate morphology might be more an adaptation for mobility in the mixed layer of the sediment. In dysoxic environments, at Furlo, we observe a return to background val- species belonging to the deep infaunal mor- ues in the relative abundance of the initial recol- phogroup are found living at the sediment surface onisers (i.e. Reophax minutus, Hormosinella cf. (Kaminski et al., 1995). An obvious mechanism distans, and Pseudobolivina cf. munda), together for dispersal would be the drifting of gametes or with the disappearance of juvenile forms of the zygotes during/after reproduction, since numerous Paratrochamminoides^Trochamminoides group. species of benthic foraminifera possess £agellated This change is here interpreted as the ¢nal phase gametes (see table 1 in Lee et al., 1991). Addition- of the recolonisation process. Based on an estima- ally, resuspension and transport of the ¢rst recol- tion of the average sedimentation rate for Chron onisers by bottom currents is another mechanism 33r in the Scaglia Rossa at Furlo, this return to for dispersion. Living specimens of Reophax scotti pre-ashfall comparable assemblages occurred have been found in plankton tows collected from some 5 kyr after the deposition of the volcaniclas- the North Sea (John, 1987), although these organ- tic material. This estimate is at odds with the ¢nd- isms may have gone into suspension in shallow ings of Hess et al. (2002), who document the re- water areas during a major storm. turn of trochamminids to the 1991 Mt Pinatubo tephra layer some 5^8 years after the volcanic eruption. As mentioned above, this estimate of 6. Conclusions the recovery time at Furlo is certainly a¡ected by relatively intense bioturbation, which would Signi¢cant di¡erences in diversity, community have displaced part of the original recolonisation structure, feeding and habitat preferences are ob- assemblage upward. served among benthic foraminiferal assemblages It is most likely that the deposition of a c. 15- below and above the volcaniclastic layer, which cm-thick volcaniclastic layer (whose original is virtually barren of foraminifera.

MARMIC 851 10-4-02 70 S. Galeotti et al. / Marine Micropaleontology 44 (2002) 57^76

Fig. 6. Relative abundance of inferred infaunal and epifaunal taxa, and DWAF diversity (Fisher-K index) across the volcaniclastic layer at Furlo. The proportion of indeterminate species is also reported. Notice the scale break between 0 and 5 cm above the volcaniclastic layer.

Below the volcaniclastic layer, DWAF assem- above the bentonitic layer, in association with ju- blages are diverse with 28^34 species, and only mi- venile/dwarfed specimens of Paratrochamminoides nor £uctuations in the relative abundance of taxo- and Trochamminoides. Reophax minutus and P. nomic groups and faunal parameters are observed. munda probably represent the initial recolonisers Common taxa are Paratrochamminoides spp., Tro- that appeared following the deposition of the vol- chamminoides dubius, Trochammina spp., Subreo- caniclastic material. Between 2 and 4 cm above phax spp., Saccammina spp. and tubular forms. the volcaniclastic layer, the highest relative abun- The recolonising assemblage above the bento- dances of Subreophax scalaris, Subreophax splen- nitic layer is signi¢cantly di¡erent in terms of didus, Hormosinella cf. distans, and Karrerulina sp. both diversity and relative abundances of taxa are observed. These taxa represent the next stage and faunal parameters, showing reduced diversity in the recolonisation process, and although ex- and an increased proportion of infaunal forms. ploiting niches opened up by the mass mortality High relative abundances of Reophax minutus of the original fauna, are not as opportunistic as and Pseudobolivina cf. munda (the latter being al- R. minutus and P. munda. The recolonisation fau- most exclusively represented just above the volca- na is comprised mainly of morphologies belong- niclastic layer) are observed in the ¢rst centimetre ing to the infaunal morphogroup (Plate I), which

MARMIC 851 10-4-02 S. Galeotti et al. / Marine Micropaleontology 44 (2002) 57^76 71

Plate I. Typical specimens of the recolonisation fauna above the volcaniclastic layer. a. Hormosinella cf. distans Brady; sample FB8, U250 b. Subreophax splendidus Grzybowski; sample FB8, U250 c. Pseudobolivina cf. munda Krasheninnikov; sample FB10, U350 d. Reophax minutus Tappan; sample FB8, U226

apparently live at the sediment surface in dis- strengthens the reliability of environmental recon- turbed conditions, and are dispersed by deep-sea structions based on DWAF. currents. The succession of species above the volcaniclastic layer at Furlo displays interesting similarities to the succession of species found colonis- Acknowledgements ing the 1991 Mt Pinatubo tephra layer in the abyssal South China Sea. Profound di¡erences We thank J. Erbacher and F. Jorissen for their between the calculated duration of the recolonisa- critical review that greatly improved the manu- tion process at Furlo and that observed in the script. This is Contribution No. 66 of the Deep South China Sea are attributed to bioturbational Water Agglutinated Foraminiferal Project. mixing in the Scaglia Rossa, which resulted in an expanded record of the fossil ‘recolonisation assemblage’. Despite the passage of 80 million years Appendix A. Taxonomic reference list of Earth History, broad similarities at the generic level between modern and fossil analogues All DWAF species identi¢ed in the samples of

MARMIC 851 10-4-02 72 S. Galeotti et al. / Marine Micropaleontology 44 (2002) 57^76 this study are alphabetically arranged with refer- Trochammina squamata var. charoides Jones and ence to their original description. Parker, 1860, p. 304. Ammobaculites sp. Glomospira glomerata (Grzybowski) Test consisting of 3^4 chambers in the coiled Ammodiscus glomeratus Grzybowski, 1898, pl.11, stage later uncoiling to form a rectilinear stage ¢g. 4. composed of 2^3 chambers. The wall is com- Glomospira gordialis (Jones and Parker) posed of medium to ¢ne grain-sized particles Trochammina squamata var. gordialis Jones and resulting in a somewhat smooth aspect. Aper- Parker, 1860, p. 304. ture is a round, simple opening at the end of Glomospira serpens (Grzybowskii) the last chamber. Ammodiscus serpens Grzybowski, 1898, pl. 11, ¢gs. 2, Ammodiscus cretaceus (Reuss) 3. Operculina cretacea Reuss, 1845, pl. 13, ¢gs. 64, 65. Glomospirella gaultina (Berthelin) Ammodiscus glabratus Cushman and Jarvis, 1928 Ammodiscus gaultinus Berthelin, 1880, pl. 1, ¢gs. 3a,b. pl. 12, ¢gs. 6, 6a. Haplophragmoides suborbicularis (Grzybowski) Ammodiscus peruvianus Berry, 1928, pl. 27. Cyclammina suborbicularis Grzybowski, 1896, pl. 9, Ammodiscus planus Loeblich, 1946, pl. 22, ¢g. 2. ¢gs. 5^6. Loeblich, 1946. Foraminifera from the type Pep- Haplophragmoides walteri (Grzybowski) per shale of Texas. J. Paleontol., 20, 130^139. Trochammina walteri Grzybowski, 1898, pl. 11, ¢g. 31. Ammodiscus sp. Hippocrepina depressa Vasicek, 1947, pl. 11, ¢gs. We place in this species specimens showing 1, 2. a slightly irregular planispiral coiling. Our Hormosina excelsa (Dylazanka) specimens are, therefore, comparable to Hyperammina excelsa Dylazanka, 1923, pl. 1, ¢g. 3. those reported as Ammodiscus sp. 1 from Hormosina velascoensis (Cushman) the Campanian and Maastrichtian Scaglia Nodosinella velasconsis Cushman, 1926, pl. 20, ¢g. 9. Rossa of the Gubbio section by Kuhnt Hormosina sp. (1990), who regarded them as intermediate Test coarsely agglutinated, composed of large forms between the genera Ammodiscus and subrounded chambers linked by thick connec- Trochamminoides. tions. Ammosphaeroidina pseudopaciloculata (Mjatliuk) Hormosinella cf. distans (Brady) Cystamminella pseudopaciloculata, Mjatliuk, 1966, pl. cf. Reophax distans Brady, 1881, p. 50. 1, ¢gs. 5^8; pl. 2, ¢g. 6; pl. 3, ¢g. 3. Specimens in our material di¡er from typical H. Aschemocella sp. distans in having a smaller test and ¢ner ag- This species resembles A. carpathica (Neagu) in glutination. possessing large chambers with a relatively Hyperammina dilatata Grzybowski, 1896, pl. 8, coarsely agglutinated wall. However, in our ¢g. 17. material we found only large fragments, which Hyperammina sp. prevented us from classifying it at the species Large subrounded chambers composed of a level. ¢nely agglutinated, thick wall. Di¡ers from H. Bulbobaculites problematicus (Neagu) dilatata in possessing less elongate, more regu- Ammobaculites agglutinans sp. problematicus Neagu, larly subrounded, instead of sack-like cham- 1962. bers. Dorothia sp. Kalamopsis grzybowskii (Dylazanka) Test ¢nely agglutinated, consisting of sub- Hyperammina grzybowskii Dylazanka, 1923, p. 65^66. rounded, in£ated chambers arranged in troco- Karreriella conversa (Grzybowski) chospiral coil, later becoming biserial. The Gaudryina conversa Grzybowski, 1901, pl. 7, ¢gs. 15, aperture is a low interiomarginal arch, at the 16. base of the last chamber. Karrerulina sp. Glomospira charoides (Jones and Parker) A small-sized species with a variably elongate,

MARMIC 851 10-4-02 S. Galeotti et al. / Marine Micropaleontology 44 (2002) 57^76 73

tapered test. The aperture is slightly produced Saccammina sphaerica Sars, 1872, p. 250. on a short neck. Saccorhiza ramosa (Brady) Lituotuba lituiformis (Brady) Hyperammina ramosa Brady, 1879, pl. 3, ¢gs. 14^15. Trochammina lituiformis Brady, 1879, pl. 5, ¢g. 16. Spiroplectammina dentata (Alth) Lituotuba sp. Textularia dentata Alth, 1850, pl. 13, ¢g. 13. Test ¢nely agglutinated with an irregularly coiled Spiroplectammina laevis (Roemer) initial portion. This form resembles Glomo- Textularia laevis Roemer, 1841, pl. 15, ¢g. 17. spira irregularis (Grzybowski) from which it is Spiroplectammina spp. di¡ers in having the tendency to uncoil in the We place in this group all specimens that, due ¢nal part of the test. to relatively bad preservation of Spiroplectam- Paratrochamminoides heteromorphus (Grzybow- mina in the HCl residues, could not be classi- ski) ¢ed at species level. Trochamminoides heteromorphus Grzybowski, 1898, pl. Subreophax guttifer (Brady) 11, ¢g. 16. Reophax guttifer Brady, 1881, p. 49. Paratrochamminoides sp.1 Kuhnt, 1990, pl. 5, Subreophax pseudoscalaris (Samuel) ¢gs. 12^13. Reophax pseudoscalaria Samuel, 1977, pl. 3, ¢gs. 4a,b. Paratrochamminoides sp.2 Kuhnt, 1990, pl. 5, Subreophax scalaris (Grzybowski) ¢gs. 14^16. Reophax guttifer var. scalaria Grzybowski, 1896, pl. 8, Paratrochamminoides/Trochamminoides juvenile ¢g. 26. We place in this group small-sized specimens Subreophax splendidus (Grzybowski) 1898 showing an irregular coiling of a few rounded Reophax splendida Grzybowski, 1898, pl. 10, ¢g. 16. to subrounded chambers resembling the initial Trochammina deformis Grzybowski, 1898, pl. 11, portion of the test in Trochamminoides and ¢gs. 20^22. Paratrochamminoides. Trochammina gyroidinaeformis Krasheninnikov, Pseudobolivina lagenaria Krasheninnikov, 1974, 1974, pl. 5, ¢gs. 7a^c, 8a^c, 9c. pl. 5, ¢gs. 1a,b, 2c. Trochamminoides dubius (Grzybowski) Pseudobolivina munda Krasheninnikov, 1973, pl. Ammodiscus dubius Grzybowski, 1898, pl. 8, ¢gs. 12, 2, ¢gs. 10^11. 14. Pseudobolivina variabilis Vasicek, 1947, pl. 1, Trochamminoides septatus (Grzybowski) ¢gs. 10^12. Ammodiscus spetatus Grzybowski, 1898, pl. 11, ¢g. 1. Recurvoides nucleosus (Grzybowski) Trochamminoides subcoronatus (Grzybowski) Trochammina nucleosus Grzybowski, 1898, pl. 11, ¢gs. Trochammina subcoronata Grzybowski, 1896, pl. 28^29. 9, ¢gs. 3a^c. Recurvoides sp. Turritellella sp. Test small, streptospiral. The 5^6 visible cham- A tubular form characterised by a series of close bers are di⁄cult to distinguish. Specimens in convolutions whose diameter slightly increases our material are often deformed. in size giving the test a somewhat conical Reophax minutus Tappan, 1940, pl. 14, ¢gs. shape. Aperture terminal, rounded. 4a,b. Uvigerinammina jankoi Majzon, 1943, pl. 2, ¢gs. Reophax subnodulosus Grzybowski, 1898, pl. 10, 15a,b. ¢gs. 17^18. Reophax sp. 5 Kuhnt, 1990, pl. 3, ¢g. 11. Rzehakina epigona (Rzehak) References Silicina epigona Rzehak, 1895, pl. 6, ¢g. 1. Saccammina grzybowskii (Schubert) Alth, A., 1850. Geognostich pala«eontologische Beschreibung d. na«schsten Umgebung von Lembereg. Haidingers Nat.wiss. Reophax grzybowskii Schubert, 1902, pl. 1, ¢g. 13. Abh. 3, 171^284. Saccammina placenta (Grzybowski) Alvarez, W., Lowrie, W., 1984. Magnetic stratigraphy applied Reophax placenta Grzybowski, 1898, pl. 10, ¢gs. 9^10. to synsedimentary slumps, turbidites and basin analysis: The

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