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Community Versus Biocoenosis in Multivariate Analysis of Benthic Molluscan Thanatocoenoses

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Community Versus Biocoenosis in Multivariate Analysis of Benthic Molluscan Thanatocoenoses Rivista Italiana di Paleontologra e Stratigrafia COMMUNITY VERSUS BIOCOENOSIS IN MULTIVARIATE ANALYSIS OF BENTHIC MOLLUSCAN THANATOCOENOSES DANIELA BASSO & CESARE CORSELLI Received AprìL 11, 20A1; accepted January 7,2002 Key-words: Multivariate statistical analysis, Thanatocoenosis, variate analysis (classification, ordination, similarity and dissimilarity Molluscs, Recent, soft bottoms, Tyrrhenian Sea analysis). This procedure appears to be the most suitable for the iden- tification of "natural" grouping of biotopes, since the results are not Riassunto. Nell'ecologia del benthos, le comunità e le biocenosi obscured by the diffuse occurrence of the most common and ubiqui- sono unità descrìttive non perfettamente corrispondenti. Benché lo tous specres. schema concettuale basato sulle comunità, originarìamente definite con un approccio statistico quantitatiyo, appaia più adeguato ad inquadrare i risultati di un trattamenro sratistìco di tanato- dati di Introduction cenosi, per il benthos mediterraneo questo schema appare un'eccessiva semplificazione delle realì unità ecologiche. Al contrario, la bionomia The use of more or less conservative associations bentonica con le biocenosi, identificate da un gruppo di specie fedeii al biotopo (indipendentemente dalla loro abbondanza) deriva da un among several organisms as descriptive units for a con- approccio qu.rlitat ìvo che ha ri\contr.ìto maggior \llccesso rrr i ricerca- sidered area is widely accepted as a major tool in benth- rori dell'area mediterrrnea. Un gruppo di dodici ranaroceno.i rìr- ic ecology and palaeoecology. However, different con- reniche è stato elaborato con entrambi gli approcci per mertere in luce ceptual approaches produced several frameworks of una strategia pratica di analisi statistica multivariata dei modelli di dis- tribuzione in paleoecologia del benthos, che combini i vantaggi degli classification, which are only partially comparible. approcci qualitativo e quantitativo. Nel caso in cui si debba trart:ìre sra- Among them, those based on the concepts of biocoeno- tisticamente una matrice di grandi dimensioni di dati di tanatocenosi sis and community have been wideiy applied in the bio- bentoniche, si raccomanda di usare un approccio qualitatìvo per un;r logical literature. drastica riduzione della matrice di partenza, trattenendo solo le specie con un preciso significato bionomico, prima di procedere all'analisi At the end of XIX century Karl Mobius (1877) multìvariata quantitativa (classificazione, ordinamento, analisi della introduced the new term " biocoenosis " to include all similarità e dissimilarità). La procedura appare la più adeguata all'iden- individuals (plants or animals) living in rhe same area, tìfìcazione di gruppì "naturali" di biotopi, in quanto il risultato non è being closely connected to the mean environmental con- confuso dalla dominanza delle specìe più comuni ed ubiquiste. ditions of their habitat and by reciprocal interspecific Abstract. Community and biocoenosis as descriptive units for functional relations. The concept of biocoenosi.s is srrict- benthic ecology are not perfectly interchangeable. Although the con- Ìy associated to that of biotope, defined as the physical ceptual framework based on communities, originally defined by a sta- space (surface or volume) where the dominant condi- tistical quantitative approach, appears to be the most suitable in the tions (biological and abiological environmenral vari- statistìcal treatment of thanatocoenoses data, this frarnework appears to oversimplify the picture of the most important ecological units in ables) are homogeneous. The biocoenoses have been the Mediterranean benthos. On the contrary, the benthic bionomy widely used to describe the distribution of benthic with the biocoenoses, identified by a group of characteristic species organisms, in relation ro the environmenral variables, (disregarding their abundance) derives from a qualirative approach within the framework of the benthic bionomy, especially which has been more successfully adopted for the research in the Mediterranean area. A group of twelve thanatocoenoses from the developed in the Mediterranean by the French scienrists Tyrrhenian Sea has been treated with both approaches with the aim to of the Station Marine d'Endoume (among others, Pérès identìfy a practical strategy for analysing multispecies distribution pat- & Picard 1964). terns in benthic paleoecology, trying to combine the advantages of The delimitation of the biocoenoses is carried out both quantitative and qualitative approaches. \When dealing with large- sized data matrices of benthic thanatocoenoses, it is recommended to by identifying a list of the characterisric species, classi- use a qualitative approach for data reduction, on the basis of their sig- fied in function of their fidelity to rhe biotope. For nificance in benthic bionomy, prìor to perform the quantitative multi- example, the exclwsir,,e characteristic (excl) species are those Dipartimento dì Scienze Geologiche e Geotecnologie, Unìversità degli Studi di Milano Bicocca,Pzza della Scienza 4, 20126 Milano. E- mail cesare.corselli@unìmib.it; daniela.basso@runimìb.it. D. Basso & C. Corsellì Stotion STl2 STl0 STl3 BorA BorB E\TO Sco2E Ellg AmlSE BelTE MC22 EllS Abra alba 221911 2e Abra nitida 143 11 Abn ovata 1034 542 205 Abra prismatica 61112 24C Acanthocardia echinata a A c a nth oc a rd i a p a u c i c o st ata 3798 A c a nth oc a rd i a tu b e rc u I ata Aciniopsis hiia 12 Acrnopsls cancellata 139 131 6 xo Aclis minor Acmaea virginea 321 17 18 Acteon tomatilis 1 Actonia testae z Aequipecten commutatus A ln Aequipecten operculais 7 10 17 207 Alvania consociella 21 Alvania discors 32 117 228 10 342 4 Alvania lineata 24 30 Alvinia jetrreysi Anadara diluvii 12 Anodontia fragilis 4 Anomia ephippium 322457915 26 Apicularia similis ozz Aporrhais pespelecani 2123 1 28 Arca noae 312111 Arca tetragona 1 5 18 Arsenia punctura 11 Astarte fusca 14 1 2 Astafte sulcata 2 Astraea rugosa 3482 122 tJ c Atys brocchii 214 1 Axinulus croulinensis 75 Azorinus chamasolen 4 Barbatia barbata 26141 Barleeia rubra 14 43 Balhvaraa orenonhia Sîo'tion STl2 STl0 STl3 BorA BorB El70 Sco2E Ellg AmlSE BelTE MC22 Ells seta Dracnystoma 2236 5i 3ela nebula 21 3ela turgida 43 3ellaspira rigida Sellaspira septangul aris 21 3 itti u m gr. rettcul atu m 1839 4954 450 1143 2190 584 4367 4 3ittìum lacteum 10 3olinus brandaris 1 3ornia sebetia 96 114 42 Sulla striata 16 )alliostoma conulus 10 7 8 )alliostoma Iaugieri 23 20 ?alliostoma wiseri )alliostoma zizyphinum )allista chione )alyptraea chinensis 9881742 227 )antharus dorbignyi 3 11 ?apulus ungaricus 1 )ardiomya costellata )ardiomya stiolata Ò )ardita calyculata 14 15 jerastoderma glaucum 3801 5/.04 549 )e rith id i u m su bmam i I I atum Seithiopsis fayalensis n )erithiopsis scalaris 2 )erithiopsis tiara 1 )erithiopsis tubercul aris 123 )erithium vulgatum 2346 7780 2170 10 21 150 21 20 )hama gryphoides 17192 )hamelea gallina 8 '10 )hauvetia minima 32 51 )hauvetia vulpecula 1 )hlamys flexuosa 2hlamys glabra 32 )hlamvs multistriata 431 B enthic mo llus can tbanato c o enos e s 155 Stotion sTl2 sTl0 sTl3 BsrA BorB El70 sco2E Etl g AmtsE BetTE MC22 Eil S lChlamvs vaia 1J lE ,l lcn,y"útid" dotiotum 11 IChrysaltida excavata lClanculus conllinus 44 4 lClanculus cruciatus 47€r' lCl ath rom ang e I i a q u ad ril I u m 1 Clausinella brogniafti 5349 Columbella rustica 1 Columbellopsis minor 711 2 Comamondia gncilis 2411123 Conus venticosus 721644 C o ral I i op h i I a I ame I I o sa 3 Corbula gibba 2 19 49 22s 110 50 1 881 Crepidula unguiformis 651 Ctena decussata 2622916 Cultellus tenuis 53 11 Cuspidaria cuspidata 911 3 Cuspidaria rcstrata 1 Cyclope neritea 261 1824 57 2 Cylichna alba 10 Cylichna crossei 1 Cylichna cylindracea 41 Cylichnina subcylindrica Dentalium dentalis De ntal i u m i n aeq uicostat u m 44 Digitaia digitata 2 Diodora gibberula 4AE Diodon graeca 167 2 Diodora italica z Diplodonta apicalis rotundata Diplodonta 1 1 Divaicella divaricata 375729 Donax trunculus 1 Donax variegatus 3 10 Donax venustus 21 Stoîion STl2 STl0 STl3 BorA BorB E|TO Sco2E Ell9 Aml5E BelTE MC22 Ells Dosinia exoleta 8413 Emarginula adiatica Emarginula elongata 46 Emaryinula huzadii 11 6 9 Emarginula punctulum Emarginula rosea 1 Emarginula tenen Ensls ensr.s Ensis siliqua minor Epilepton clarkiae 4 Epitonium aculeatum Epitonium cl ath ratul u m 1 Epitonium èommune Epitonium tiberii J Epitonium tuftoni 1 Eulimella tunis Fossarus amblguus Fusinus gigliolii Fusinus pulchellus Fuslnus rosfrafus Fusinus rudis zu5 Fustiaria rubescens Galeodina carinata Galeomma tuúoni Gibberula caelata Gibberula miliaia 40 143 2.4 Gibberula philippii 12123 G i bbe ru I i na cl a nde sti n a 572 313 tl Gibberulina occulta 41 Gibbula adansoni 21 Gibbula ardens 46 Gibbula fanulum 1 42 Gibbula guttadaui 52 Gibbula magus 13 Gibbula rarilineata 13 17 23 4 1,56 D. Basso G C. Corsellt Stotion STl2 STl0 STl3 BorA BorB El70 Sco2E Ell9 AmlSE BelTE MC22 Ells Gibbula turbinones 't0 15 Gibbula vaia 21351 Glans aculeata Glans trapezia 29 40 52 Glycymerís glycimeris Glycymeis insubica Gonilia calliglypta 132 Goniostoma auiscalpium 34 62 Gouldia minima 17387016096 Hadiania craticuloides 1 Haliotis tuberculata 731 I Haminoea cymoelium Haminoea hydatis 44 1 Hiatella rugosa 6 10 3 4 1 4 42 Hinia incrassata 99 205 1 Hinia limata Hinia pygmaea tc Homalopoma sanguineum 22 42 20 17 Hyala vitrea 4 Hyalina mitrella Hyalopecten similis Q?2 108 Hydrobia stagnalis 66880 61952 17872 lrus irus 43 Jujubinus exasperatus 114 255 136 152 to Jujubinus
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