Santa Catalina (Lequeitio, Basque Country): An ecological and cultural insight into the nature of prehistoric fishing in Cantabrian Spain

Journal of Archaeological Science: Reports 6 (2016) 645–653

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Santa Catalina (Lequeitio, Basque Country): An ecological and cultural insight into the nature of prehistoric fishing in Cantabrian Spain Eufrasia Roselló-Izquierdo a, Eduardo Berganza-Gochi a, Carlos Nores-Quesada b, Arturo Morales-Muñiz a,⁎ a b

Laboratorio de Arqueozoología, Depto. Biología, Universidad Autónoma de Madrid, 28049 Madrid, Spain Departamento de Biología de Organismos y Sistemas, Universidad de Oviedo, 33071 Oviedo, Spain

a r t i c l e

i n f o

Article history: Received 31 March 2015 Received in revised form 19 May 2015 Accepted 1 June 2015 Available online 10 June 2015 Keywords: Fish Fishing Taphonomy Cantabrian region Spain Upper Paleolithic Epipaleolithic

a b s t r a c t The paper presents the fish collections retrieved in the site of Santa Catalina. The results from this analysis are explored in terms of the Richness, Diversity, Equitability and Trophic level values of the 61 archeological levels that span from the Upper Magdalenian (15 ky cal BC) to the Azilian (10 ky cal BC). These results are discussed within the frame of theoretical proposals on the origin and evolution of prehistoric fishing in the Cantabrian region, and evidence that some of these proposals appear to be in need of revision. © 2015 Elsevier Ltd. All rights reserved.

1. Introduction Considered long to be the cradle of prehistoric research in the Iberian Peninsula, the Upper Paleolithic (36–12 ky cal BP) of the Cantabrian region is still lagging behind the study of fish remains when compared with groups such as mammals, mollusks or even birds (Straus, 2005; Álvarez-Fernández, 2011). Such drawback has had a negative impact in all attempts to provide a holistic view of the economy and subsistence strategies of those past hunter and gatherer populations. Among the reasons that might explain such contingency, ÁlvarezFernández (2011) mentions: (1) burial of the intertidal zones due to alluviation, colluviation and wind-blown sand, (2) a lack of accurate information about the position of present day sites relative to the prehistoric coastline, (3) a proportional over-representation of deposits younger than 16 ky cal BP, (4) an impossibility to carry out reliable intra- and inter-site comparisons due to studies systematically lacking key data on the area and volume of the excavated deposits, (5) improper retrieval methods that include non-sieving and an inadequate sorting of organic remains by ill-prepared students resulting in huge losses of small remains, (6) a failure to study marine animals or to leave them identified at a non-informative level (i.e., “Pisces indet.”), and (7) little ⁎ Corresponding author. E-mail addresses: [email protected] (E. Roselló-Izquierdo), [email protected] (E. Berganza-Gochi), [email protected] (C. Nores-Quesada), [email protected] (A. Morales-Muñiz).

http://dx.doi.org/10.1016/j.jasrep.2015.06.002 2352-409X/© 2015 Elsevier Ltd. All rights reserved.

attention being paid to taphonomic issues (Álvarez-Fernández, 2011: 329–330). Two major contingencies that need to be added to this list include (1) sea-level rise that during the past millennia flooded the Cantabrian continental shelf, doing away with most of the sites carrying evidence of marine fishing, and (2) an absence of adequate fish reference collections, in both Cantabrian Spain and the Iberian Peninsula at large, that partly explain point (6) in Álvarez-Fernández's list of problems. Such lack of reference materials is particularly harmful in the case of the three salmonid “species” from this region (i.e. the brown trout, the sea trout and the salmon) whose bones are often impossible to be set apart from each other. Given the aforementioned state of affairs it should come as no surprise to learn that (1) most of the Cantabrian sites that have thus far provided fish remains are located at some distance from the coast (i.e. are inland sites in the strict sense of the term, and were more so during most of the Middle and Upper Paleolithic times when sea levels were lower than they are today) and the fish remains therein found not only (1) constitute ludicrously small samples, both in absolute and relative terms when compared to the remaining fauna, but also (2) are mostly confined to the genus Salmo that either lives permanently (brown trout) or else seasonally penetrates deeply (sea trout, salmon) into freshwater. Such limitations have not prevented theories being put forward on the issue of prehistoric fishing in the Cantabrian region (Clark, 1976, 1983; Cleyet-Merle, 1990; Hayden et al., 1987; Le Gall, 1998, 1999; Meléndez et al., 1986; Pokines and Krupa, 1997; Russ, 2010; Straus,

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E. Roselló-Izquierdo et al. / Journal of Archaeological Science: Reports 6 (2016) 645–653

1983, 1992, 2005; Straus et al., 2002). Recently there has been a revival of these theoretically-bent papers with many hypotheses being proposed that concentrate on the role fishes played on the diet of the Cantabrian hunter–gatherer populations as well as in the character of what have been labeled as the fisheries of the Cantabrian region (Adán et al., 2009; Álvarez-Fernández, 2011; Turrero et al., 2008, 2012, 2013, 2014a,b). In this paper, we will present the fish assemblages retrieved in the Upper Magdalenian–Azilian site of Santa Catalina, explore their structure from an ecological standpoint, and discuss the data provided by these samples in light of some of the proposals put forward on the nature of prehistoric fishing along the former Cantabrian coast. 2. Material and methods The cave of Santa Catalina (geographical coordinates: 02°30′33″ longitude W; 43°22′35″ latitude N; UTM coordinates: X = 539.746, Y = 4.802.749, Z = 35 masl) is one in a karstic system of four cavities located at the base of the lighthouse of the city of Lekeitio (Vizcaya, Basque country, Spain) on a cliff presently overlying the Cantabrian sea (Fig. 1). It is a small cave (86 m2) with two entrances, whose strategic location when the sea level was lower allowed it to have control of the narrow coastal plain that extended to the former shoreline, as well as access to a wide spectrum of biotopes ranging from rivers and estuaries to the low mountain peaks (i.e. below 1000 m) of the Bermeo-Arno anticlinal range. From 1982 until 2000 the site witnessed fifteen one-month excavation campaigns at one of the entrances where a non-cemented archeological deposit had been discovered during prospection operations. The excavation was laid out into seven 1 m2 squares (A4, A6, A8, B4, B6, B8, D4) and the deposit excavated following the sedimentary features of the statigraphy in each of the squares. These features, varying in depth from 1–5 cm, were recognized as independent levels. On the basis of their archeological and sedimentological features a total of 61 levels have been grouped into three stages that represent a typical shell midden deposit in the upper two stages and a midden deposit with scarce shells on the lowermost stage (Berganza et al., 2012). In relation to the Greenland GISP2 ice core record in the 38.0–5.2 ky cal BC window, Stage III (31 levels; Table 3) represents an Upper Magdalenian that started to form under the cold and humid conditions that correspond to the latest moments of the GS-2 stadial and that, during the initial stages of the GI-1 interstadial, witnessed an improvement of the climate

followed by a short series of cold pulses. Stage II (15 levels; Table 4), represents a Final Magdalenian that coincided with the latest stages of the GS-1 interstadial that were characterized by milder temperatures and increased rainfall but that also featured harsher environmental conditions at the end of the sequence that may indicate cold pulses associated with the Younger Dryas period. Lastly, Stage I (15 levels; Table 5), represents a moment from the Azilian (i.e. local Epipaleolithic) culture featuring temperate climatic conditions, slightly colder than those existing in the region today. AMS radiocarbon dates on short-lived materials, mostly bone, though exhibiting some degree of overlap, seem fully coincidental with the archeological, sedimentological and climatic data sets (i.e., Upper Magdalenian: 15.007–12.887 cal BP; Final Magdalenian: 14.540– 11.602 cal BP; Azilian: 12.664–10.158 cal BP) (Table 1) and indicate that the occupation of Santa Catalina lasted from the latest moments of the Pleistocene to the earliest moments of the Holocene (i.e. the “Tardiglacial” in a loose sense of the word). Although in terms of excavated volume all stages exhibited comparable values (i.e. Stage III: 2.98 m3; Stage II: 3.49 m3; Stage I: 2.56 m3), not all fish remains were retrieved in the same manner. In this way, most of the sediment extracted from Stage I was dry-sieved through a 2.5 mm-wide mesh. But starting already from Level 10 (Stage I), the sediments from the B8 sector were wet-sieved in a tower of which the widest screen was 1 mm and the narrowest was 0.5 mm. From Level 18 (Stage II) onwards, this procedure was carried out on all samples. After drying, all materials were subjected to a process of sorting with the help of stereomicroscopes. The identification was carried out with the help of the reference collection of one of the authors (AMM) housed at the Laboratorio de Arqueozoología of the Universidad Autónoma de Madrid. To facilitate comparisons, both the NISP (i.e. number of identified remains) and MNI (minimum number of individuals) as defined by Reitz and Wing (1999) have been used as the estimators of abundance. MNI estimations took into account those elements that could be identified to taxon. In the case of paired bones, the convention of selecting the highest number of elements, whether right or left, as indicative of the MNI was followed. In the case of vertebrae these were first allocated to a size group through comparison with specimens of known size, and each group was taken to represent an MNI = 1. NISP and MNI were calculated for each of the 61 levels and their values later pooled for each of the stages. For evaluating the biological nature of the samples, a series of measures and indexes have been utilized. For each level, richness,

Fig. 1. Location of the site of Santa Catalina in the Cantabrian region and layout of the ground plan of the cave with an indication of the area at one of the entrances where the excavations took place.

E. Roselló-Izquierdo et al. / Journal of Archaeological Science: Reports 6 (2016) 645–653 Table 1 Radiocarbon dates from the Santa Catalina levels. All AMS dates except those from Teledyne Isotopes (I) (calibration through Calib611). Stage

Code

Date BP

Calibrated date BP (2σ)

I I I I I II II II II II III III III III III

Ua-24651 Ua-2358 I-16247 Ua-4280 Ua-2360 Ua-43257 I-15061 Ua-43258 Ua-43259 Ua-24652 Ua-42320 Ua-24654 Ua-24655 Ua-13876 Ua-13877

9760 ± 65 10530 ± 110 9510 ± 270 10100 ± 85 9180 ± 110 10392 ± 179 11460 ± 420 10911 ± 244 11961 ± 61 11155 ± 80 12146 ± 98 11225 ± 80 12345 ± 85 12405 ± 95 12425 ± 90

11,071–11,311 12,089–12,644 10,158–11,718 11,328–12,020 10,171–10,609 11,602–12,640 12,519–14,540 12,374–13,306 13,657–13,990 12,775–13,245 13,751–14,257 12,887–13,300 14,007–14,945 14,059–15,007 14,093–15,024

diversity and equitability have been estimated (Reitz and Wing, 1999). In the case of richness we have taken the S index in a slightly altered manner to refer not only to the number of species but also of any other taxon, be it genus, family or order in any given level. This was done because in many cases, identifying fish bones to the level of species is next to impossible for a variety of circumstances (e.g. lack of diagnostic features, bone damage). Given that many remains can only be allocated within higher taxonomic categories yet represent species present in a sample that would have been otherwise overlooked, incorporating higher taxonomic categories into the S value counts prevents one from further deflating the actual diversity of a sample. A more practical reason to proceed in this way was to make our estimates comparable with those from other studies. Diversity within each of the levels has been estimated by calculating the Shannon–Weaver function (H′) as defined by the formula: H0 ¼

 XS   i i logp p i¼1

where: pi s

the relative abundance of the ith taxon within the sample using the MNI as the estimator of abundance the number of taxonomic categories (species, genera, families, etc.) within the level.

Heterogeneity within each of the levels has been estimated by calculating Equitability (Lloyd and Ghelardi, 1964) by way of the V′ estimator as defined by the formula: V0 ¼ H″ = log S where: H′ S

the Shannon–Weaver function the number of taxonomic categories within the level.

For both H′ and V′ calculations decimal logarithms have been used. H′ and V′ values for each stage were calculated by taking the pooled counts of taxa from all the levels from one stage as a single sample (Table 2). One final measure to describe the structure of the assemblages was the Trophic level (TL). This was calculated for each one of the levels, taking the pooled counts of taxa from all of those belonging to one stage as a single sample (Table 2). The theoretical aspects of the TL have been explained by Pauly et al. (1998, 2000). In this case, TL values have been taken from Froese and Pauly (2015). The single extrapolation we

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had to make was that of the sandeel genus Hyperoplus whose TL was not provided by this resource. We assigned this taxon the TL value given in Fishbase for the species Hyperoplus inmaculatus (i.e. 4.4). TL values are specified for each taxon in Table 2. Previous to subjecting the samples to statistical analyses, the pooled data from each of these three chronocultural stages were tested for normality by means of a Shapiro Wilk test. This was done online through the website http://sdittami.altervista.org/shapirotest/ShapiroTest/html. The analyses for significance in the observed differences among stages included a one-way ANOVA test for independent variables, followed by an a posteriori Tukey test. Both tests were carried out online at the website http://vassarstats.net/. 3. Results Table 2 provides an overview of the fish assemblages from Santa Catalina in terms of MNIs and, in a summarized manner, of NISPs. The taxonomic diversity included 32 species and 11 additional genera (of which Salmo, Alosa and Phycis do not incorporate any remains identified to the level of species), representing a minimum of 25 families grouped into a minimum of 11 orders. Indeed, a substantial number of the nonidentified remains, that together represent ca. 30% of the studied samples, appear to be, in our opinion, potentially identifiable to the level of the species/genus. Failure to identify them reflects an absence of the pertinent specimens in our reference collection. Given that fully 15% of the species recorded at Santa Catalina (i.e. pike, Esox lucius, cod Gadus morhua, Bull Rout Myoxocephalus scorpius, Eelpout Zoarces viviparus and Butterfish Pholis gunnellus) are presently alien or very rare finds in Iberian waters, makes us suspect that additional non-Iberian taxa may have been present in this potentially identifiable sector of the assemblages. One remarkable feature of the three major assemblages is that sheer size does not apparently correlate with diversity. As can be seen in Table 2, despite far smaller samples, both the faunas from Stage I (representing 6–7% of the identified and total count of fishes, respectively) and Stage II (representing ca. 20% of both the identified and total counts) featured an identical number of taxa (i.e. 32) that was larger than the 28 taxa recorded for Stage III which accounted for ca. 74% of the total number of remains and 75% of those identified to taxon. Retrieval methods contributed to dictate the size of these assemblages. Both wet-sieving (systematically applied in Stage III but partly in Stage II and only marginally in Stage I) and mesh size [2.5 mm (Stage I) vs. 1–0.5 mm (Stages II and III)] explain why the average number of remains per level is ca. 12 in Stage I, rises to 42 in Stage II and reaches 97 in Stage III. Such differences (Tables 3–5) are independent of the fact that the number of levels from Stage III doubles those from Stages II and I (i.e. 31 vs. 15) and make one suspect that substantial numbers of remains must have been lost in the upper two levels due to inadequate retrieval methods. If this were so, both the pooled collections from Stages II and I and those from their constituent levels would represent gross under-representations of the original samples that would render them, strictly speaking, not comparable to those from Stage III. Stages II and I must have been originally far more diverse than what is reported here. This phenomenon would still fit neatly within an overall scenario of temperatures on the increase from the Upper Magdalenian to the Final Magdalenian/Azilian, that would have fostered, among other things, an increase in the number of temperate Atlantic water species in the Gulf of Biscay at the end of the Pleistocene. A more consistent view of general diversity emerges when the ecological indexes are estimated for each level. Tables 3, 4 and 5 incorporate the individual values of Richness (S), Diversity (H′), Equitability (V′) and Trophic level (TL) for these 61 levels. Tables 6, 7, 8 and 9 summarize Richness, Diversity, Equitability and Trophic level data for the three stages into which samples have been grouped. Diversities have been plotted in Fig. 2 for a more clear appreciation of trends. In terms of Richness, the Final Magdalenian (Stage II) exhibited the highest values both in terms of the range of number of taxa per level

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Table 2 Overview, in terms of the MNI, of the fish assemblages from Santa Catalina grouped into stage (III: Upper Magdalenian; II: Late Magdalenian; I: Azilian). Numbers within brackets after the scientific names of each taxon indicate the Trophic level (TL) values. Taken from fishbase.org). Stage

III

Taxa

MNI

Chondrichthyes indet.(3.6) Lamna nasus (3.8) Rajiformes indet. (3.5) Anguilla anguilla (3.7) Conger conger (3.4) Alosa sp. (3) Sardina pilchardus (2.8) Engraulis encrasicolus (2.8) Esox lucius (4.1) Salmo sp. (3.6) Ciliata mustela (3.8) Gadus morhua (3.8) Gaidropsarus cf. vulgaris (3.5) Melanogrammus aeglefinus (3.5) Merlangius merlangus (3.5) Molva molva (3.8) Phycis sp. (3.8) Pollachius pollachius (3.8) Trisopterus luscus (3.8) Gadidae indet. (3.8) Belone belone (3.8) Atherina presbyter (2.8) Myoxocephalus scorpius Dicentrarchus labrax (3.5) Serranus cabrilla (3.5) Trachurus trachurus (3.3) Diplodus cf. vulgaris (3.4) Diplodus sp. (3.4) Pagellus erythrinus (3.4) Sparidae indet. (3.4) Chelon labrosus (2.1) Liza aurata (2.4) Mugil cephalus/Liza sp. (2) Coris julis Labrus bergylta (3.6) Labrus bergylta/Labrus bimaculatus (3.6) Labrus sp. (3.6) Symphodus cf. melops (3.5) Symphodus sp. (3.5) Zoarces viviparus Pholis gunnellus cf. Trachinus draco cf. Blennius ocellaris (3) Lipophrys sp./Parablennius sp. (3) Lipophrys sp. (3) Parablennius sp. (3) Blennidae indet. (3) Gobius sp. (3.2) Ammodytes cf. tobianus Hyperoplus sp. Scomber japonicus (3.3) P. flesus/P. platessa/L. limanda (3.5) Pleuronectidae indet. (3.5) Total MNI Total NISP identified Total NISP non-identified Total NISP

II %

MNI

1 2 6

0.13 0.27 0.83

4

0.55

1 439

0.13 61.4

70 1 1 2 1 1

9.8 0.13 0.13 0.27 0.13 0.13

5

0.7

3

0.41

1 1 1

1

1

I

0.13 0.13 0.13

%

Total

MNI

3 28 4

0.8 8.1 1.1

32 12

9.3 3.5

140 2 11 6

40.7 0.6 3.1 1.7

5

1.4

15

4.3

3 1 1 2 4 1 2 4

0.8 0.3 0.3 0.6 1.1 0.3 0.6 1.1

%

1

0.8

1 5 2 2 22 14

0.8 4.2 1.7 1.7 18.6 11.8

12 2 2 4 1

10.1 1.7 1.7 3.3 0.8

1 1 5 1 6

0.8 0.8 4.2 0.8 5

1 1 1

0.8 0.8 0.8

1 2

0.8 1.7

1

0.8

2

1.7

0.13

0.13

2 3 1

0.27 0.41 0.13

1 1

0.13 0.13

1 2

0.13 0.27

161 1 715 2459 927 3386

22.5 0.13 100 27.3 100

(3–17) and the mean number of taxa per level (9.46; Table 6). The Upper Magdalenian (Stage III), although featuring an identical range of taxa per level as the Azilian (i.e. 1–11), despite its far larger number of levels (31 vs. 15), exhibited a lower mean number of taxa per level (3.83) than the Azilian (5.9). Differences in this parameter seem also significant between the Final Magdalenian and Azilian so that taxonomic richness did not evidence any clear trend within the temporal interval marked by these three stages in terms of absolute number of taxa but fluctuated instead among them. Prior to testing the significance of the differences in the Diversity, Equitability and Trophic level measurements among stages, the levels

2 2 10 1 3

0.6 0.6 2.9 0.3 0.8

7

5.9

2

1.7

1 5 9 3 3 4

0.3 1.4 2.6 0.8 0.8 1.1

1 3 10 1 2 1

0.8 2.5 8.5 0.8 1.7 0.8

1 24

0.3 7

344 631 271 902

100 42.9 100

118 184 142 326

100 43.5 100

MNI

%

1 1 6 39 6 2 58 26 1 591 4 83 11 2 2 1 1 1 1 15 1 21 3 4 2 3 3 5 2 4 4 1 1 2 2 3 17 1 5 2 3 1 2 9 20 4 5 5 1 2 1 185 1 1177 3274 1340 4614

0.08 0.08 0.5 3.3 0.5 0.16 4.9 2.2 0.08 50 0.33 7 0.9 0.16 0.16 0.08 0.08 0.08 0.08 1.3 0.08 1.8 0.25 0.33 0.16 0.25 0.25 0.42 0.16 0.33 0.33 0.08 0.08 0.16 0.16 0.25 1.44 0.08 0.42 0.16 0.25 0.08 0.16 0.76 1.7 0.33 0.42 0.42 0.08 0.16 0.08 15.8 0.08 100 29 100

within each stage were subjected to a Shapiro–Wilk test for normality (W statistic). In the case of Diversity (H′), normality (N) was found for all three stages [Stage I (N = 15; Mean = 1,455; SD = 0,645; W = 0,922; p N 0,1; N); Stage II (N = 15; Mean = 1,653; SD = 0,429; W = 0,908; p N 0,1; N); Stage III (N = 31; Mean = 0,584; SD = 0,361; W = 0,929; p N 0,05; N)]. In the case of Equitability (V′) normality could not be assumed for Stage I levels (N = 15; Mean = 0,455; SD = 0,249; W = 0,555; p b 0,01; NN), but levels from both Stage II (N = 15; Mean = 0,788; SD = 0,125; W = 0,968; p N 0,1; N); and Stage III (N = 31; Mean = 0,452; SD = 0,245; W = 0,936; p N 0,05; N) conformed with normality. Finally, in the case of Trophic level (TL)

E. Roselló-Izquierdo et al. / Journal of Archaeological Science: Reports 6 (2016) 645–653 Table 3 Stage III (Upper Magdalenian) levels with their corresponding values for the different parameters taken to describe the structure of the fish samples (see text for further explanations).

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Table 5 Stage I (Azilian) levels with their corresponding values for the different parameters taken to describe the structure of the fish samples (see text for further explanations).

Level

NISP

S

H′

ν′

TL

Level

NISP

S

H′

ν′

TL

29B 29C 29D 29E 29 F 29G 29H 30 30A 30B 30C 30D 30E 30F 30G 30H 30J 31A 31B 31C 31D 31E 31F 31G 31H 31J 31K 31M 31N 31P 32

1 1 15 100 41 166 141 20 5 30 16 28 45 22 37 33 28 85 172 182 201 152 202 341 165 122 42 46 10 4 6

1 1 3 6 4 5 5 8 2 2 2 3 1 2 4 2 3 5 3 3 4 7 5 11 5 4 5 2 3 4 1

0 0 0.6261 0.6602 0.6334 0.6796 0.9034 1.5382 0.6729 0.244 0.3767 0.3086 0 0.1834 0.7172 0.3048 0.4902 0.6656 0.7132 0.608 0.5424 0.8408 0.6447 0.8484 0.8281 0.9522 0.6123 0.4939 0.6388 1.3863 0

0 0 0.5699 0.3684 0.4569 0.4222 0.5613 0.7397 0.9708 0.352 0.5435 0.2809 0 0.2646 0.5173 0.4397 0.4462 0.4135 0.6491 0.5534 0.3912 0.4342 0.4254 0.3538 0.5145 0.6869 0.3804 0.7126 0.5814 1 0

3.6 3.6 3.48 3.59 3.59 3.6 3.61 3.44 3.54 3.58 3.6 3.6 3.6 3.55 3.59 3.59 3.6 3.6 3.59 3.6 3.62 3.6 3.6 3.6 3.61 3.61 3.6 3.58 3.58 3.6 3.6

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

9 12 11 11 3 14 1 10 16 15 2 20 41 12 7

4 6 6 5 3 2 1 8 7 10 2 11 10 8 6

1.2145 1.5376 1.72 1.1592 1.0975 0.5985 0 2.0249 1.7493 2.0691 0.6931 2.2891 1.941 1.9754 1.7539

0.8761 0.8581 0.9599 0.7202 1 0.8635 0 0.9737 0.8989 0.8986 1 0.9941 0.8429 0.9499 0.9789

2.88 3 3.13 3.11 3.3 2.8 3.4 3.36 3.02 3.04 3.15 3.15 3.11 3.45 3.4

estimations, normality could be assumed for levels from Stages I (N = 15; Mean = 3,153; SD = 0,195; W = 0,950; p N 0,1; N) and II (N = 15; Mean = 3,295; SD = 0,28; W = 0,938; p N 0,1; N), though not for those from Stage III (N = 31; Mean = 3,586; SD = 0,037; W = 0,599; p b 0,01; NN). In the light of the slight divergences in these two instances, it has been tentatively assumed that all samples conformed to normality. In terms of diversity (Tables 3, 4, 5 and 7) it does seem that the Upper Magdalenian faunas exhibited the lowest H′ values, both in terms of range (0.1834–1.5382) and mean (Ῡ = 0.6476). Those from the Final Magdalenian (range: 0.8802–2.1767; Ῡ = 1.6536) and the Azilian (range: 0.5985–2.2891; Ῡ = 1.5587), on the other hand, appear to be very similar and their differences non-significant. Indeed, one-way ANOVA test for independent samples evidenced significant differences to exist among all three stages (k = 3; N = 61; SS = 14.6340; df = 2;

Table 4 Stage II (Late Magdalenian) levels with their corresponding values for the different parameters taken to describe the structure of the fish samples (see text for further explanations). Level

NISP

S

H′

ν′

TL

16 16bis 17 18 19 20 21 22 23 24 25 26 27 28 29

17 3 21 18 9 89 88 96 35 41 22 54 33 13 92

8 3 10 6 7 15 17 17 11 11 11 5 6 3 12

1.758 1.0975 2.1767 1.2413 1.8882 1.8814 2.0604 2.0539 1.9375 1.7914 2.1546 1.1357 1.3528 0.8802 1.395

0.8454 1 0.9453 0.6379 0.9703 0.6947 0.7272 0.7249 0.808 0.7471 0.8985 0.7056 0.755 0.8012 0.5613

3.1 3.33 3.23 3.41 3.35 2.97 3.03 3.08 3.09 3.16 3.38 3.57 3.51 3.63 3.58

MS = 7,320; F = 34,366; p b 0.0001) yet the a posteriori Tukey test confirmed these differences to be significant only between Stages I and III (p b 0,01) and between Stages II and III (p b 0.01) but not between Stages I and II (p N 0,05 NS). Given that the samples from the Upper Magdalenian are by far the largest ones and richness was essentially comparable to that of the ensuing stages (see above) what this means is that diversities during these initial moments of the occupation were heavily influenced by the abundance of very few taxa. Indeed, the data from Table 2 reveal that the abundance of the Genus Salmo throughout all of the levels from Stage III seems to be the reason that explains such comparatively reduced diversity at these earliest moments of the occupation at Santa Catalina. When equitability values are considered (Tables 3,4,5 and 8), a gradual shift towards a more even representation of taxa from the earliest to the latest levels of the occupation seems evident. In this case the change among the stages is not evident in the maximum values of the range, all very close to a V′ value of 1 (Level III: 0.9708; Level II: 0.9703; Level I: 0.9941) (Table 8) but rather in the shifts revealed by both the lowest values of that range (Level III: 0.2646; Level II: 0.5613; Level I: 0.7202) and the mean values (Level III: 0.5565; Level II: 0.7881; Level I: 0.9153) towards greater evenness. One-way ANOVA test for independent samples evidenced significant differences to exist among all three stages (k = 3; N = 61; SS = 2.105; df = 2; MS = 1.053; F = 21.135; p b 0.0001). The a posteriori Tukey test confirmed these differences to be significant between Stages I and III (p b 0.01) and also between Stages II and III (p b 0.01) but not between Stages I and II (p N 0.05). In other words, not only were a greater number of taxa recorded in the late Magdalenian and Azilian of Santa Catalina but in those stages the fish assemblages were not skewed towards one single taxon as was the case of Salmo during the Upper Magdalenian. Trophic levels evidenced the clearest trend of all estimators and allow one to further probe into the nature of these temporal shifts through time (Tables 3, 4, 5 and 9). In this case, both the minimal overlapping of the ranges between the Upper and Final Magdalenian stages and that of the Azilian, as well as the mean TL values, evidence significant differences. The ANOVA test for independent samples evidenced significant differences to exist among all three stages (k = 3; N = 61; SS = 2,142; df = 2; MS = 1.071; F = 50.047; p b 0.0001). The a posteriori Tukey test confirmed these differences to be significant Table 6 Summary of Richness (S) values for each chronocultural stage. Stage

N

Range

Ῡ

III II I

31 15 15

1–11 3–17 1–11

3.8 9.5 5.9

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Table 7 Summary of Diversity (H′) values for each chronocultural stage.

Table 9 Summary of Trophic level (TL) values for each chronocultural stage.

Stage

N

Range

Ῡ

Stage

N

Range

Ῡ

III II I

31 15 15

0.1834–1.5382 0.8802–2.1767 0.5985–2.2891

0.6476 1.6536 1.5587

III II I

31 15 15

3.44–3.62 2.97–3.63 2.88–3.45

3.6017 3.2625 3.1175

between not only Stages I and III (p b 0.01) and Stages II and III (p b 0.01), but also between Stages I and II (p b 0.05). From a qualitative perspective what this shift revealed was a change from a community of essentially predatory fishes (Upper Magdalenian: 3.6017) down the trophic chain towards one of essentially planktonic or microphagous feeders (Azilian: 3.1175). When one confronts the list of taxa (Tables 2–5) this numerical change reflects a shift from samples dominated by salmonid fishes to samples where Clupeiformes (i.e. sardines and anchovies) and, secondarily, Blennidae (i.e. blennids) came to dominate the samples. 4. Discussion Contrary to what seemed to have been the rule for Cantabrian prehistory up until now, where salmonid fishes were often the sole taxon recorded in archeological sites (but see Roselló and Brinkhuizen, 1994), the fish assemblages from Santa Catalina reveal an unprecedented taxonomical diversity. Even when considering the reported diversity as definitive, highly unlikely, given the substantial number potentially identifiable remains awaiting identification, what seems beyond question is that very few fish assemblages from prehistoric or fully historical times, in Europe and elsewhere, bear witness to such a diversity at the taxonomic level. More important, the data emerging from the Santa Catalina fish assemblages call into question several propositions put forward during these past thirty years on the issue of marine adaptations and the nature of Cantabrian fisheries during prehistoric (i.e. Upper Paleolithic and Mesolithic) times. 1. Onset and timing of marine fishing. Concerning the earliest records of marine fishes in the Cantabrian region Álvarez-Fernández (2011) writes that these “… comes from Middle Magdalenian contexts” by which he is referring to the single Pleuronectid (i.e. flatfish) remain reported by Morales (1984) at Tito Bustillo. Turrero et al. (2014a) further specify that “… regional marine fisheries seem to start, at least to a significant extent, in the Azilian” (Turrero et al. (2014a, b)), by which it is not totally clear whether they are referring here to “northern Iberia” as mentioned in the title of their article, or exclusively to the province of Asturias that is the only zone of the Cantabrian coast considered in their survey. The data from Santa Catalina evidence that a fishing activity was taking place in a marine environment in the Upper Paleolithic some five millennia previous to the Azilian and that this activity was taking place at an obviously larger scale than that provided by the single flatfish bone found at Tito Bustillo. In fact, this should come as no surprise given that marine fishes had been mentioned in the Aurignacian (35–28 ky BP) levels of Cueto de la Mina and in the Aurignacian and Gravettian (27–21 ky BP) levels of Aitzbitarte III (Rasilla Vives, 1990; Roselló and Morales, 2011). A reluctance to consider amphidromous taxa such as eels and salmonids, as

Table 8 Summary of Equitability (V′) values for each chronocultural stage. Stage

N

Range

Ῡ

III II I

31 15 15

0.2646–1 (0.9708) 0.5613–0.9703 0.7202–1 (0.9941)

0.5565 0.7881 0.9153

marine rather than freshwater catches may have a lot to do with this hypothesis of marine fishes appearing late in the sequence. At Santa Catalina, although one cannot decide where the fishes had been harvested, the most plausible scenario on account of the overall composition of the fish faunas points to the marine environment. Given that salmon and occasionally sea trout concentrate at the mouth of rivers previous to their spawning run and that juvenile salmon do the same upon their return into the sea, a transitional water environment, such as an estuary, would more properly fit the taxonomic composition of the studied assemblages. 2. Composition of the catch: stasis vs. change. When it comes to diversity, the Santa Catalina assemblages reveal a situation that completely changes the paradigm of Cantabrian fish assemblages essentially dominated by salmonids at all times. In the case of families, for example, Turrero et al. (2014a) write “it is clear that more varied fish were caught in later periods … whereas Solutrean and Magdalenian catch were exclusively composed of salmonids” (Turrero et al. (2014a)). As can be seen in Table 2, the Late Magdalenian assemblages from Santa Catalina are as diverse as those from the Azilian, if not more, and incorporate fishes from no less than 25 families (i.e. Lamnidae, Rajidae, Anguillidae, Congridae, Clupeidae, Engraulidae, Esocidae, Salmonidae, Gadidae, Atherinidae, Cottidae, Moronidae, Serranidae, Carangidae, Sparidae, Mugilidae, Labridae, Zoarcidae, Pholidae, Trachinidae, Blennidae, Gobiidae, Ammodytidae, Scombridae and Pleuronectidae). Such number not only represents a four-fold increase in the number of families provided by Turrero et al. (2014a) for both the Azilian and ensuing Asturian (i.e. Mesolithic) periods in the Cantabrian area, but also a significantly higher number of families than that recorded in the Azilian of Santa Catalina (15). In other words, contrary to what has been postulated, the onset of the Holocene at Santa Catalina was accompanied by a 40% decrease in diversity at the level of the family that was used as the proxy for fish diversity by Turrero et al. (2014a). As was seen, both in terms of Richness (S) and diversity (H′) the Final Magdalenian (Level II) is the stage signaling the maximum fish diversity at Santa Catalina so the trend in family numbers seems consistent with these other ecological measures of diversity except for one caveat: the number of families decreases slightly through time (Table 2). The trend is far from significant at the quantitative level (i.e. Upper Magdalenian: 18 families; Final Magdalenian: 17 families; Azilian: 15 families) yet informative from a qualitative standpoint. Indeed, what such trend evidences is a gradual replacement during these five millennia that bridge the gap between the Pleistocene and the Holocene of cold-water taxa that belong to families with few species (e.g. Pholidae, Zoarcidae, Ammodytidae, Cottidae) by temperate water taxa belonging to richer families in terms of species as is the case of the sea breams (Sparidae) and blennies (Blennidae). Although in terms of sheer numbers the absolute number of taxa was not much altered through time, what one is witnessing here is a drastic faunal turnover. Indeed, the nature of the fish assemblages at Santa Catalina changed drastically in terms of families, genera and species throughout the five millennia of occupation not just in terms of presence and absence of specific taxa but also in terms of the abundances each one of these exhibited. Not a single taxon featured stasis throughout these five millennia, fluctuations being instead the rule. It is hard to imagine that humans remained as mere spectators of these complex dynamics (Kettle et al., 2008, 2010). Indeed, if fishing is the reason that explains these assemblages, then one can also postulate that humans qualitatively shaped the

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Fig. 2. Diversity values expressed by the Shannon–Weaver function (H′), for the 61 levels arranged into stages. Dotted lines correspond to an H′ = 1.

composition of the collections that were found. That, in spite of all these taphonomical, methodological and taxonomical contingencies that underlie the diversities of the Santa Catalina fish collections, the overall number of taxa remained essentially constant seems puzzling and could be another hint to prove the role played by humans in shaping the composition of the catch. As for the evolution of the Trophic level values, contrary to what has been postulated it seems clear that these changes meant a lowering of the mean TL values of the assemblages, not their increase (Turrero et al., 2014a). 3. Fish as a resource. Much has been written on the role that fishes at large and marine fishes in particular might have played in the subsistence of prehistoric hunter-gatherers of the Cantabrian region. Originally, in light of the scarcity of remains, certain authors had postulated a marginal role for fishes in those human communities (Clark, 1983; Straus, 1992). More recently, Turrero et al. (2014a) talk about “… the increasing importance of marine resources in the Upper Paleolithic” when discussing the results from their study on fishes and Adán et al. (2009) mention salmon as being both crucial in the human diet during the Upper Paleolithic in France, and also constituting a “quality resource” when referring to the Cantabrian region. One problem with most of these proposals is that one does not know the databases on which they are based or, when this is known, the databases are ludicrously small. A second, equally important problem is how does one assess the importance of fishes? Although a wealth of non-zooarcheological data, from cave art depictions to stable isotope analysis bear witness to the importance of fish within the symbolic realm and their use as food for Late Upper Paleolithic people, none of these data allow for straightforward interpretations or are devoid of problems (Adán et al., 2009; Richards and Hedges, 1999; Richards et al., 2005). At Santa Catalina we have a possibility to assess the contribution that fishes played within an animal assemblage that numbers close to 200,000 identified specimens (Table 10). Although for many reasons the vertebrate and invertebrate assemblages are non-comparable (e.g., most of the sea urchin samples are made up of spines of which a single specimen may bear hundreds) a comparison of the assemblages from the three vertebrate group appears feasible. Using NISP values, the data in Table 10 appear very eloquent. In this way, against a “static” background of mammals that represent 55–59% of all

vertebrates at all times, fish NISPs constitute a substantial third of the total fauna during the Upper Magdalenian but decrease to half that amount during the Late Magdalenian and a little bit further still during the Azilian (i.e., 13.5% vs. 15.7%). Although these figures cannot be taken at face value because systematic retrieval through wet-sieving with 1–0.5 m size meshes was only systematically applied in the Upper Magdalenian levels, fish losses must have not been too different to those experienced by birds and small mammals. One way or the other, if we restrict our comparisons to the earliest stage of the occupation one can see that fishes could by no means be considered a marginal resource at that time. Whether this has been also the case in the ensuing periods and to what extent can the figures from Santa Catalina be extrapolated to other sites of the Cantabrian region will remain open to question at this point. But even if the abundance trends seen in Table 10 could not be taken at face value, it seems clear that the data from the Upper Magdalenian stage would call into question the proposal that marine “fisheries” only became important in the Cantabrian region after the onset of the Holocene (Adán et al., 2009; Turrero et al., 2014a,b). 4. Seasonality. It has been recently proposed that, in the case of salmon, preferential winter harvest took place in the Cantabrian region from the Last Glacial Maximum onwards (Turrero et al., 2013). This hypothesis, based on a mere 25 vertebrae from mixed deposits covering a time span of more than ten millennia, seems a bit far-fetched. On the other hand, the data emerging from Santa Catalina are not always easy to interpret in terms of seasonality since many species (e.g. sea

Table 10 Overview of the faunal collections from Santa Catalina grouped into chronocultural stages. Values are expressed in terms of the NISP. The percentages of the mammal, bird and fish assemblages have been estimated from the combined values of the vertebrate samples. Total

Stage III (%)

Stage II (%)

Stage I (%)

Total (%)

Mammals Birds Fishes Ʃ vertebrates Mollusks Crustaceans Echinoderms Ʃ ID

4306 (58.8) 553 (7.5) 2459 (33.6) 7318 (92.4) 601 8 7827 15,754

2178 (54.4) 1194 (29.8) 631 (15.7) 4003 (41) 5767 96 73,893 83,759

753 (55.6) 417 (30.8) 184 (13.5) 1354 (27) 3694 163 86,654 91,865

7237 (57) 2164 (17) 3274 (25.8) 12,675 (56) 10,062 267 168,374 191,378

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breams, wrasses, blennies and other rock pool taxa) might have been year-round residents whereas others, such as codfishes and Clupeiformes, migratory to an extent that is yet to be determined. Yet, if one concedes that fishes behaved thousands of years ago much in the same way as their modern counterparts do today, it seems clear that the abundance of salmonids of all sizes at Santa Catalina (Roselló, unpublished data) contradicts the proposal of a winter harvesting as juveniles appear mostly during the spring and summer (Quero, 1984; Wheeler, 1969; Froese and Pauly, 2015). In fact, adult salmonids, meaning salmon and sea trout, appear in Cantabrian rivers essentially all throughout the year. Their value as seasonality indicators is thus questionable despite most salmons exhibiting a clear seasonal signal. At Santa Catalina one finds three clear instances of “summer bioindicators”, as are the horse mackerel (Trachurus trachurus; Stage III), the mackerel (Scomber japonicus; Stage II) and the needlefish (Belone belone; Stage I) (Fischer et al., 1987; Quero, 1984; Wheeler, 1969; Whitehead et al., 1986; Froese and Pauly, 2015) (Table 2). Summer fishing would also be suggested by the presence of sardines and anchovies, so that if salmonids were indeed taken during the winter, then the presence of summer fishing indicators would suggest an activity carried out throughout the year. As things presently stand, and although the issue must of necessity remain open, the data from Santa Catalina do not argue for a winter harvesting of fishes. 5. Conclusions Fishing is a phenomenon that, in terms of human activity, often combines issues concerning availability and choice. From a zooarcheological perspective, very rarely can the data be taken to reveal the structure of a former fish community or the decisions taken to fish a particular species at a given place and time and the fishing method and tackle selected to that end. From such standpoint, the fish remains from the site of Santa Catalina offer, for the first time in Cantabrian prehistory, the possibility to explore a variety of propositions, both theoretical and practical (i.e. also methodological), against a reliable dataset. Many issues remain open at this stage in relation with these fish assemblages. Two prominent ones concern the taphonomy and the size/weight inferences of the studied remains. In terms of taphonomy, a detailed analysis of traces and fracture patterns is currently under way to determine whether or not alternative accumulators might have played a role in the formation of the deposits. To this end, the inference of the sizes and estimation of the weights for each one of the thousands of fish bones will prove crucial as our preliminary assessments are revealing drastic size changes through time in individual species as well as on the assemblages taken at large. Equally crucial will be to determine the nature of the former shorelines, presently lying underwater, as these could inform about the nature of the biotopes where fishes could have been captured (Clark and Mix, 2002; Ehlers and Gibbard, 2004; Galparsoro et al., 2010). Finally, an isotopic and bio-molecular project is projected to start at the end of this year that will try to determine, among other things, the species represented in the unidentified fraction of the fish samples and the provenience of remains in terms of the biotopes that were harvested. For the moment, suffice it to say that the scenarios revealed by this collection of fishes seem to be far more diverse and far more labile that what had been hitherto postulated. As a result, one would do well to wait until Santa Catalina loses its singular character in the frame of prehistoric fishing in the Cantabrian region and Europe before putting forward more hypotheses about this fascinating phenomenon. Acknowledgments The authors wish to acknowledge the help received from all the research team members that worked at Santa Catalina along the years.

Special thanks are expressed to Igor Gutiérrez, Mikel Elorza, Pedro Mª Castaños, Victor Vásquez and Teresa Rosales for allowing use of the data from their respective faunal groups. José Luis Arribas is gratefully thanked for all the help received and Leif Jonsson for providing us with specimens and unpublished data from his databases. Laura Llorente is thanked for her collaboration in the preparation of the manuscript. The authors also would like to thank the two anonymous reviewers of this manuscript for their thoughtful and positive input. This research benefited from Grant HAR 2014-55722-P (“Ictioarqueologia de la Prehistoria cantábrica: Modelos para la caracterización de las primeras pesquerías europeas”) of the Spanish Ministerio de Economía y Competitividad. References Adán, G.E., Álvarez-Lao, D., Turrero, P., Arbizu, M., García-Vázquez, E., 2009. Fish as diet resource in North Spain during the Upper Paleolithic. J. Archaeol. Sci. 36 (3), 895–899. Álvarez-Fernández, E., 2011. 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