Research Article |
Corresponding author: Antonella Petrocelli ( antonella.petrocelli@iamc.cnr.it ) Academic editor: Antonella Lugliè
© 2019 Antonella Petrocelli, Ester Cecere, Fernando Rubino.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Petrocelli A, Cecere E, Rubino F (2019) Successions of phytobenthos species in a Mediterranean transitional water system: the importance of long term observations. In: Mazzocchi MG, Capotondi L, Freppaz M, Lugliè A, Campanaro A (Eds) Italian Long-Term Ecological Research for understanding ecosystem diversity and functioning. Case studies from aquatic, terrestrial and transitional domains. Nature Conservation 34: 217-246. https://doi.org/10.3897/natureconservation.34.30055
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The availability of quantitative long term datasets on the phytobenthic assemblages of the Mar Piccolo of Taranto (southern Italy, Mediterranean Sea), a lagoon like semi-enclosed coastal basin included in the Italian LTER network, enabled careful analysis of changes occurring in the structure of the community over about thirty years. The total number of taxa differed over the years. Thirteen non-indigenous species in total were found, their number varied over the years, reaching its highest value in 2017. The dominant taxa differed over the years. The number of species in each taxonomic division also varied. In addition to the centuries-old exploitation of its biotic resources, mainly molluscs, the basin has been subject for a long time to a range of anthropogenic driving forces linked to urbanisation, shipbuilding, agriculture and military activities, which have caused chemical and biological pollution, eutrophication and habitat destruction. It may therefore be assumed that these changes were closely related to human activities. Indeed, it was observed that the reduction of only one of these forces, i.e. urban sewage discharge, triggered the recovery of phytobenthos. Therefore, it may be assumed that if the anthropogenic pressure on the Mar Piccolo was eased, it could once again become the paradisiacal place it was held to be in ancient times.
LTER, Mar Piccolo, Mediterranean Sea, phytobenthos, transitional water systems
Human beings play a fundamental role in the ongoing degradation of coastal marine ecosystems (
The availability of historic data on benthic populations could allow better measurement of the changes (
The Mar Piccolo of Taranto (southern Italy) has belonged to the Italian LTER network since 2011 (LTER_EU_IT_095; https://deims.org/ac3f674d-2922-47f6-b1d8-2c91daa81ce1) (
The aim of the present study was to increase our knowledge of the structure of phytobenthic assemblages in the Mar Piccolo of Taranto and to reconstruct the history of macrophytic communities in terms of both species composition and dominance changes during the last 30 years. For this purpose, seasonal quantitative data, collected at various sites in the Mar Piccolo in 2008 (Project VECTOR http://vector.conismamibi.it) and from spring 2011 to winter 2018 (I-LTER network), were carefully analysed and compared with the floristic data collected in 1989. At the same time, again on the basis of already published data, speculative parallels were drawn with the demographic and socio-economic development of the town of Taranto (https://tinyurl.com/yacl8t6t;
The Mar Piccolo of Taranto is a lagoon-like basin located north of Taranto. It has a surface area of 20.72 km2 and is divided into two sub-basins known as the First Inlet, to the west and the Second Inlet, to the east (Fig.
In 2008, sampling exercises were carried out at monthly intervals at three sites, two in the First Inlet (Station A and Station B) and one in the Second Inlet (Station C).
Station A (40°29'35"N, 17°14'17"E) was characterised by soft bottoms and artificial hard substrata, mainly concrete blocks. Station B (40°30'01"N, 17°15'10"E) was characterised by a soft muddy bottom. Station C (40°29'40"N, 17°19'18"E) had a soft muddy bottom with scattered concrete blocks.
In the period spring 2011 - winter 2018, four sites were seasonally sampled, two in the First Inlet (Station 1 and Station 2) and two in the Second Inlet (Station 3 and Station 4) (Fig.
Station 1 (40°28'46"N, 17°13'41"E) was the only urban station, located in the old town. Artificial hard substrata prevailed, composed mainly of discarded plastic used in mussel farming activities but also concrete quays. Station 2 (40°30'03"N, 17°15'30"E) was located next to the mouth of a small river, with a soft muddy bottom and a few artificial substrates, mainly unlawfully built concrete quays used by mussel farmers. Station 3 (40°29'39"N, 17°19'22"E) was the only station characterised by natural hard substrata, rare in the Mar Piccolo, i.e. small rocks and the remains of a stone dock. It was close to the mouth of a small river. Station 4 (40°28'20"N, 17°18'25"E) was located in a zone where mussels and other seafood are manually cleaned, bagged and sold. A soft muddy bottom and concrete platforms were the main substrata.
The samples were handpicked from within a 50×50 cm square, randomly placed on the bottom. Three replicates were collected at each station. Seaweed thalli were stored in plastic bags and transported to the laboratory within a few hours. There, the macrophytes were sorted and each species was identified and weighed on a triple-beam balance. Data were expressed as kg wet weight m-2 (hereafter kg m-2). A list of species was compiled for each dataset, including the unpublished floristic data from 1989, with nomenclature as in
The quantitative data recorded in 2008 were not processed, since the sampling sites were not the same as those of 2011–2018. Therefore, only the total number of species, taxonomic divisions, biogeographical elements and dominant species were compared.
Since seasonal samplings started in spring 2011, to allow statistical tests to be performed on all four seasons of the year, the analysis was carried out during a period embracing the spring, summer and autumn of one year together with the winter of the following year (spring 2011 - winter 2012).
All univariate and multivariate analyses were performed using PRIMER v.6 (Primer-E Ltd., Plymouth, UK).
Two different matrices of the means were constructed from the three replicates collected at each station. The first matrix considered all the identified taxa and the second matrix only the NIS taxa. From the absolute abundance matrices (taxa vs. samples), the Bray-Curtis similarity index was calculated after log(x+1) transformation, in order to remove the effects of orders-of-magnitude differences between samples, to normalise the data and to increase the importance of smaller values, such as the mid-range species (
The PRIMER ‘DIVERSE’ routine was used to calculate the taxonomic richness (S), taxon abundance (n°taxa), Pielou’s evenness index (J’) and Shannon-Weaver diversity index (H’) for each sample.
The statistical significance of spatial and temporal variations in the community structure across the defined factors, “year”, “season”, “basin” and “station”, were tested by one-way analysis of similarities (ANOSIM). A two-way crossed analysis (year x station) was performed to highlight possible differences from year to year. In addition, bi-dimensional representations of the statistical comparisons amongst the samples collected during the seven years at the four stations were performed by means of non-parametric multidimensional scaling (nMDS) with superimposed hierarchical clustering and a cut-off at 80–90% similarity.
The SIMPER routine was used to identify the species that contributed most to dissimilarities amongst sampling sites (one-way procedure) and to explain the changes in terms of biomass or species composition (two-way procedure, years vs. sites).
All the multivariate tests were performed on the Bray-Curtis similarity matrix including all the identified taxa.
The floristic lists of species collected in 1989, 2008 and 2011–2018 are reported in Table
Trend in the total number of species. 2018 data for winter only. In addition to the analysed years 2008 and 2011–2018, unpublished floristic data from 1989 have been included.
Floristic list (presence/absence data) of seaweeds and phanerogams recorded in the Mar Piccolo in the different years. B=biogeographic element, A=Atlantic, C=Cosmopolitan, CT=CircumTropical, IP=IndoPacific, M=Mediterranean.
B | 1989 | 2008 | 2011 | 2012 | 2013 | 2014 | 2015 | 2016 | 2017 | 2018 | |
---|---|---|---|---|---|---|---|---|---|---|---|
Unidentified Bacillariophyta | + | + | + | + | |||||||
Unidentified Cyanophyta | + | + | |||||||||
Rhodophyta | |||||||||||
Acanthophora nayadiformis (Delile) Papenfuss | IP | + | + | ||||||||
Agardhiella subulata (C. Agardh) Kraft et M.J. Wynne | A | + | + | + | |||||||
Aglaothamnion tenuissimum (Bonnemaison) Feldmann-Mazoyer | A | + | + | ||||||||
Aglaothamnion tripinnatum (C. Agardh) Feldmann-Mazoyer | A | + | |||||||||
Alsidium corallinum C. Agardh | A | + | + | + | + | + | + | + | + | + | + |
Alsidium helminthochorton (Schwendimann) Kützing | M | + | |||||||||
Amphiroa beauvoisii J.V. Lamouroux | A | + | + | + | + | + | + | + | + | ||
Antithamnion cruciatum (C. Agardh) Nägeli | A | + | + | + | + | + | + | ||||
Antithamnion tenuissimum (Hauck) Schiffner | M | + | |||||||||
Callithamnion corymbosum (J.E. Smith) Lyngbye | A | + | + | + | + | + | + | + | |||
Caulacanthus cf. ustulatus (Mertens ex Turner) Kützing | C | + | + | + | + | + | + | + | |||
Ceramium cimbricum f. flaccidum (H.E. Petersen) G. Furnari et Serio | C | + | + | + | + | ||||||
Ceramium comptum Børgesen | A | + | + | ||||||||
Ceramium diaphanum (Lightfoot) Roth | C | + | |||||||||
Ceramium incospicuum Zanardini | M | + | + | + | |||||||
Ceramium siliquosum (Kützing) Maggs et Hommersand | C | + | + | + | + | + | + | ||||
Chondracanthus acicularis (Roth) Fredericq | C | + | + | + | + | + | + | + | + | + | |
Chondracanthus teedei (Mertens ex Roth) Kützing | C | + | + | + | + | + | + | + | |||
Chondria capillaris (Hudson) M.J. Wynne | C | + | + | ||||||||
Chondria dasyphylla (Woodward) C. Agardh | C | + | |||||||||
Chylocladia verticillata (Lightfoot) Bliding | A | + | + | + | + | + | + | ||||
Corallina officinalis Linnaeus | C | + | + | + | + | + | |||||
Dasya baillouviana (S.G. Gmelin) Montagne | CT | + | |||||||||
Dasya ocellata (Grateloup) Harvey | A | + | + | + | |||||||
Dasya punicea (Zanardini) Meneghini | A | + | |||||||||
Dasya rigidula (Kützing) Ardissone | A | + | + | ||||||||
Dasysiphonia sp | + | + | + | ||||||||
Ellisolandia elongata (J. Ellis et Solander) K.R. Hind et G.W. Saunders | A | + | + | + | + | + | + | + | + | ||
Erythrocladia irregularis Rosenvinge | C | + | + | ||||||||
Gayliella flaccida (Harvey ex Kützing) T.O. Cho et L.J. McIvor | C | + | + | + | |||||||
Gelidiella lubrica (Kützing) Feldmann et Hamel | IP | + | + | + | |||||||
Gelidium crinale (Hare ex Turner) Gaillon | C | + | + | + | + | + | + | + | |||
Gelidium pusillum (Stackhouse) Le Jolis | C | + | + | + | + | ||||||
Gigartina cf. pistillata (S.G. Gmelin) Stackhouse | A | + | + | ||||||||
Gracilaria bursa-pastoris (S.G. Gmelin) P.C. Silva | C | + | + | + | + | + | + | + | + | + | + |
Gracilaria dura (C. Agardh) J. Agardh | A | + | + | ||||||||
Gracilaria gracilis (Stackhouse) Steentoft, L.M. Irvine et Farnham | A | + | + | + | + | + | + | + | + | + | |
Gracilaria longa Gargiulo, De Masi et Tripodi | M | + | + | ||||||||
Gracilariopsis longissima (S.G. Gmelin) Steentoft, L.M. Irvine et Farnham | C | + | + | + | + | + | + | + | + | + | |
Grateloupia cf. filicina (J.V. Lamouroux) C. Agardh | C | + | + | + | + | + | + | + | + | ||
Grateloupia minima P.L. Crouan et H.M. Crouan | A | + | + | ||||||||
Grateloupia turuturu Yamada | IP | + | + | + | + | + | + | + | + | ||
Griffithsia schousboei Montagne | A | + | |||||||||
Herposiphonia secunda (C. Agardh) Ambronn | CT | + | + | ||||||||
Herposiphonia tenella (C. Agardh) Ambronn | CT | + | + | + | + | + | + | ||||
Heterosiphonia crispella (C. Agardh) M.J. Wynne | A | + | |||||||||
Huismaniella nigrescens (Feldmann) G. Furnari, Cormaci, Alongi et Perrone | M | + | |||||||||
Huismaniella ramellosa (Kützing) G.H. Boo et S.M. Boo | CT | + | |||||||||
Hydrolithon cruciatum (Bressan) Y.M. Chamberlain | A | + | + | ||||||||
Hydrolithon farinosum (J.V. Lamouroux) Penrose et Y.M. Chamberlain | C | + | + | + | + | ||||||
Hypnea cornuta (Kützing) J. Agardh | IP | + | + | + | + | + | + | + | + | ||
Hypnea musciformis (Wulfen) J.V. Lamouroux | CT | + | + | + | + | + | |||||
Hypnea spinella (C. Agardh) Kützing | CT | + | + | + | |||||||
Jania rubens (Linnaeus) J.V. Lamouroux | C | + | + | + | + | ||||||
Jania virgata (Zanardini) Montagne | A | + | |||||||||
Laurencia intricata J.V. Lamouroux | A | + | |||||||||
Lomentaria clavellosa (Lightfoot ex Turner) Gaillon | A | + | |||||||||
Lomentaria compressa (Kützing) Kylin | M | + | |||||||||
Lophosiphonia obscura (C. Agardh) Falkenberg | C | + | |||||||||
Nitophyllum albidum Ardissone | M | + | |||||||||
Osmundea oederi (Gunnerus) G. Furnari | A | + | + | + | |||||||
Osmundea pelagosae (Schiffner) K.W. Nam | M | + | |||||||||
Peyssonnelia bornetii Boudouresque et Denizot | M | + | |||||||||
Phymatolithon calcareum (Pallas) W.H. Adey et D.L. McKibbin ex Woelkerling et L.M. Irvine | A | + | |||||||||
Polysiphonia denudata (Dillwyn) Greville ex Harvey | C | + | + | ||||||||
Polysiphonia elongata (Hudson) Sprengel | A | + | + | ||||||||
Polysiphonia morrowii Harvey | IP | + | + | + | + | + | + | ||||
Polysiphonia subulata (Ducluzeau) Kützing | A | + | + | + | + | ||||||
Porphyra linearis Greville | A | + | + | ||||||||
Porphyra umbilicalis Kützing | C | + | + | + | |||||||
Pterocladiella capillacea (S.G. Gmelin) Santelices et Hommersand | C | + | + | ||||||||
Pterocladiella melanoidea (Schousboe ex Bornet) Santelices et Hommersand | A | + | + | + | |||||||
Pyropia leucosticta (Thuret) Neefus et J. Brodie | A | + | |||||||||
Radicilingua reptans (Kylin) Papenfuss | A | + | |||||||||
Radicilingua thysanorhizans (Holmes) Papenfuss | A | + | + | + | + | + | + | + | + | ||
Rhodymenia ardissonei (Kuntze) Feldmann | A | + | + | ||||||||
Rhodymenia pseudopalmata (J.V. Lamouroux) P.C. Silva | A | + | + | + | |||||||
Rytiphlaea tinctoria (Clemente) C. Agardh | A | + | |||||||||
Solieria filiformis (Kützing) P.W. Gabrielson | A | + | + | ||||||||
Spyridia filamentosa (Wulfen) Harvey | C | + | + | + | + | + | + | + | + | + | + |
Stylonema alsidii (Zanardini) K.M. Drew | C | + | |||||||||
Stylonema cornu-cervi Reinsch | A | + | |||||||||
Wrangelia penicillata (C. Agardh) C. Agardh | CT | + | |||||||||
Unidentified non geniculate Corallinaceae | + | + | |||||||||
83 | 24 | 34 | 22 | 29 | 25 | 32 | 33 | 34 | 37 | 18 | |
Ochrophyta | |||||||||||
Colpomenia peregrina Sauvageau | C | + | + | + | + | + | + | + | |||
Colpomenia sinuosa (Mertens ex Roth) Derbès et Solier | C | + | + | + | |||||||
Cutlera chilosa (Falkenberg) P.C. Silva | M | + | |||||||||
Cutleria multifida (Turner) Greville | C | + | + | ||||||||
Cystoseira barbata (Stackhouse) C. Agardh | IP | + | + | + | + | + | |||||
Cystoseira compressa (Esper) Gerloff et Nizamuddin | A | + | + | + | + | + | |||||
Dictyota dichotoma (Hudson) J.V. Lamouroux v. dichotoma | C | + | + | + | + | + | + | + | + | + | |
Dictyota dichotoma (Hudson) J.V. Lamouroux var. intricata (C. Agardh) Greville | C | + | + | + | + | + | + | + | + | + | + |
Ectocarpus siliculosus (Dillwyn) Lyngbye | C | + | + | + | + | + | + | ||||
Feldmannia mitchelliae (Harvey) H.-S. Kim | C | + | |||||||||
Halopteris filicina (Grateloup) Kützing | C | + | + | ||||||||
Hincksia dalmatica (Ercegović) Cormaci et G. Furnari | M | + | + | + | |||||||
Nemacystus flexuosus var. giraudyi (J. Agardh) De Jong | M | + | |||||||||
Padina pavonica (Linnaeus) Thivy | CT | + | + | + | + | + | + | + | |||
Petalonia fascia (O.F. Müller) Kuntze | C | + | + | ||||||||
Scytosiphon lomentaria (Lyngbye) Link | C | + | + | + | + | + | |||||
Sphacelaria cirrosa (Roth) C. Agardh | C | + | + | + | |||||||
Sphacelaria fusca (Hudson) S.F. Gray | C | + | + | + | |||||||
Sphacelaria rigidula Kützing | C | + | + | ||||||||
Undaria pinnatifida (Harvey) Suringar | IP | + | |||||||||
20 | 4 | 3 | 3 | 9 | 8 | 9 | 10 | 11 | 14 | 7 | |
Chlorophyta | |||||||||||
Bryopsis corymbosa J. Agardh | A | + | |||||||||
Bryopsis cupressina J.V. Lamouroux | M | + | + | ||||||||
Bryopsis pennata J.V. Lamouroux | A | + | + | + | |||||||
Bryopsis plumosa (Hudson) C. Agardh | C | + | + | + | |||||||
Caulerpa cylindracea Sonder | CT | + | + | ||||||||
Chaetomorpha linum (O.F. Müller) Kützing | C | + | + | + | + | + | + | + | + | + | + |
Cladophora dalmatica Kützing | A | + | + | ||||||||
Cladophora glomerata (Linnaeus) Kützing | C | + | + | + | + | ||||||
Cladophora hutchinsiae (Dillwyn) Kützing | C | + | + | ||||||||
Cladophora laetevirens (Dillwyn) Kützing | C | + | + | + | + | + | + | + | |||
Cladophora lehmanniana (Lindenberg) Kützing | A | + | + | ||||||||
Cladophora prolifera (Roth) Kützing | A | + | |||||||||
Cladophora ruchingeri (C. Agardh) Kützing | A | + | |||||||||
Cladophora rupestris (Linnaeus) Kützing | A | + | + | + | |||||||
Cladophora sericea (Hudson) Kützing | C | + | + | ||||||||
Codium fragile subsp. fragile | IP | + | + | + | + | + | |||||
Ulva compressa Linnaeus | C | + | + | + | + | + | + | ||||
Ulva curvata (Kützing) De Toni | A | + | + | + | + | + | |||||
Ulva flexuosa Wulfen | C | + | + | + | + | ||||||
Ulva intestinalis Linnaeus | C | + | + | + | + | + | + | + | |||
Ulva laetevirens Areschoug | C | + | + | + | + | + | + | + | + | + | + |
Ulva prolifera O.F. Müller | C | + | + | + | + | + | + | + | |||
Ulva pseudorotundata Cormaci, G. Furnari et Alongi | A | + | + | + | + | + | + | + | + | + | |
Ulva rigida C. Agardh | A | + | + | + | + | + | |||||
Umbraulva dangeardii M.J. Wynne et G. Furnari | A | + | + | + | + | + | + | + | |||
Valonia macrophysa Kützing | CT | + | |||||||||
26 | 9 | 10 | 9 | 12 | 16 | 11 | 10 | 15 | 11 | 8 | |
Spermatophyta | |||||||||||
Cymodocea nodosa (Ucria) Ascherson | A | + | + | + | + | + | + | + | + | ||
Ruppia cirrhosa (Petagna) Grande | C | + | |||||||||
Zostera noltei Hornemann | A | + | |||||||||
3 | 0 | 0 | 1 | 2 | 1 | 1 | 1 | 2 | 1 | 1 | |
TOTAL | |||||||||||
132 | 37 | 47 | 35 | 52 | 50 | 53 | 54 | 62 | 63 | 34 |
Species | 1989 | 2008 | 2011 | 2012 | 2013 | 2014 | 2015 | 2016 | 2017 | 2018 |
---|---|---|---|---|---|---|---|---|---|---|
Agardhiella subulata (C. Agardh) Kraft et M.J. Wynne | + | + | + | |||||||
Caulerpa cylindracea Sonder | + | + | ||||||||
Codium fragile subsp. fragile | + | + | + | + | + | + | ||||
Colpomenia peregrina Sauvageau | + | + | + | + | + | + | + | |||
Dasysiphonia sp | + | + | + | |||||||
Grateloupia minima P.L. Crouan et H.M. Crouan | + | + | ||||||||
Grateloupia turuturu Yamada | + | + | + | + | + | + | + | + | ||
Hypnea cornuta (Kützing) J. Agardh | + | + | + | + | + | + | + | + | ||
Hypnea spinella (C. Agardh) Kützing | + | + | + | |||||||
Osmundea oederi (Gunnerus) G. Furnari | + | + | + | |||||||
Polysiphonia morrowii Harvey | + | + | + | + | + | + | ||||
Solieria filiformis (Kützing) P.W. Gabrielson | + | + | ||||||||
Undaria pinnatifida (Harvey) Suringar | + | |||||||||
TOTAL 13 | 3 | 5 | 3 | 4 | 5 | 7 | 7 | 7 | 9 | 4 |
The number of species in each taxonomic division (i.e. Rhodophyta, Ochrophyta, Chlorophyta and Spermatophyta) also varied (Fig.
Trend in taxonomic divisions. 2018 data for winter only. In addition to the analysed years 2008 and 2011–2018, unpublished floristic data from 1989 have been included. R=Rhodophyta, O=Ochrophyta, C=Chlorophyta, S=Spermatophyta.
In biogeographical terms, Cosmopolitan taxa ranked first each year, followed by Atlantic. Other interesting features were the low number of Mediterranean taxa and the increase in Indo-Pacific taxa from 2011 onwards (Fig.
Chorological spectrum of taxa collected in the Mar Piccolo over the years. 2018 data for winter only. In addition to the analysed years 2008 and 2011–2018, unpublished floristic data from 1989 have been included. A=Atlantic, C=Cosmopolitan, CT=CircumTropical, IP=Indo-Pacific, M=Mediterranean.
The dominant taxa, reaching a maximum yield of at least 5 kg m-2, differed over the years. Table
Maximum yearly yelds of dominant taxa (kg m-2) in the Mar Piccolo of Taranto. In brackets values lower than 5 kg m-2.
2008 | 2011 | 2012 | 2013 | 2014 | 2015 | 2016 | 2017 | 2018 (winter) | |
---|---|---|---|---|---|---|---|---|---|
Amphiroa beauvoisii | – | 8.7 | 17.2 | 10.9 | 10.8 | 8.5 | 7.3 | 6.5 | (0.93) |
Chaetomorpha linum | 29.9 | 10.8 | 11.3 | (3.8) | 5.4 | (0.4) | 6.1 | (3.6) | (3.1) |
Chondracanthus acicularis | (2.0) | (0.1) | (4.8) | 6.8 | (4.6) | (3.4) | (2.0) | 11.6 | (2.4) |
Dictyota dichotoma var. dichotoma | (0.1) | (0.2) | (0.8) | (0.1) | (2.2) | 6.5 | 10.3 | 11.4 | – |
Dictyota dichotoma var. intricata | (0.4) | 6.3 | (1.9) | 9.5 | 6.9 | (1.6) | (2.5) | 9.3 | 10.0 |
Ellisolandia elongata | – | 7.5 | (1.4) | 5.6 | (2.6) | 9.7 | (1.4) | 5.2 | (0,1) |
Hypnea cornuta | (2.3) | 7.7 | 7.3 | 10.1 | 16.1 | 8.4 | 5.0 | (4.1) | – |
Spyridia filamentosa | (0.9) | (0.1) | (0.2) | (2.6) | (0.5) | (0.2) | (1.8) | 7.1 | (0.1) |
Ulva intestinalis | – | – | (0.9) | – | (0.1) | – | 5.1 | (0.1) | (0.1) |
Ulva laetevirens | (1.2) | 9.7 | (3.4) | (3.8) | (1.1) | (2.8) | (4.6) | (2.9) | (1.1) |
Considering the study period as a whole, the yearly biomass values (mean ± SD) for the whole basin ranged from 1.4±0.9 kg m-2 in 2014–2015 to 1.8±1.4 kg m-2 in 2017–2018 (Fig.
Mean values (±SD) calculated for the years of the study, sampling sites and seasons. a, b, c biomass (g m-2) d, e, f species richness g, h, i ecological diversity.
Species richness (S) and ecological diversity (H’) increased over the years (Fig.
Only small, marginally significant differences in the macrophyte community structure were apparent over the years (RANOSIM = 0.038; p = 0.03). The two-way crossed ANOSIM showed that there were no differences in community structure from one year to the next, while there was variation between years distant from each other. For example, the pairwise test for 2011–2012 and 2017–2018 revealed significant variation in community structure (R = 0.363; p = 0.001), while the difference between 2016–2017 and 2017–2018 was not significant (R = 0.363; p = 0.3). Indeed, for immediately successive years, the R statistic was almost always negative.
The nMDS representation of the total biomass means per Station shows that the two stations in the Second Inlet were very close to each other and different from those in the First Inlet (Fig.
nMDS representation of the mean total biomass values reported for each sampling site in the period 2011–2018.
Considering the analysis of each site, Station 2 was found to differ sharply from all the other stations (Fig.
Mean values reported for Station 2 in the period 2011–2018. a biomass (g m-2) b species richness c ecological diversity.
Within the nMDS plot of the biomass means calculated over the years versus the sampling sites, in the first two years (2011–2012 and 2012–2013), Station 2 grouped with all the other sites, while in the other five years, sharp segregation was evident (Fig.
nMDS representation of the biomass means calculated over the period 2011–2018 at the four sampling sites. In the plot, the symbols indicate the station and the numbers refer to the sampling year (1 2011–2012 2 2012–2013 3 2013–2014 4 2014–2015 5 2015–2016 6 2016–2017 7 2017–2018).
Station 3 showed the highest abundance of NISs, both for the study period as a whole and year by year (Fig.
Mean numbers (±SD) of NISs in the period 2011–2018, sampling sites and seasons. a, b, c biomass (g m-2) d, e, f species richness g, h, i ecological diversity.
LTER observations make it possible to detect the natural variability of ecological systems and the interaction between abiotic and biotic variables, as well as the effects on the environment of human activities (
In the Mar Piccolo, comparison of the situation dating back about thirty years and the results of the recent seven-year study show that qualitative and quantitative changes have occurred in the phytobenthic community and are ongoing. The statistical analysis showed that for the period 2011–2018, the structure of the phytobenthic community in the two basins of the Mar Piccolo differed and that a significant difference was also apparent between the sites investigated in the First Inlet, mainly due to the peculiar features of Station 2. At the same time, the results from the two stations of the Second Inlet were more similar even though with a significant difference in their community structure. Temporal variation was also observed, which was more evident when comparing distant years, while no significant differences were observed between successive years. Considering that the basin has been subject to a significant human pressure for centuries (especially from urban pollution and mussel farming) (
Similar developments occurred over the 20 years from 1983 to 2003 in the Venice Lagoon, where a marked change in species composition was recorded (
In the Orbetello Lagoon, a clear and progressive switchover of species occurred in the period 1983–2011, reflecting the worsening of ecological conditions in the lagoon, with the rise of Chaetomorpha linum (
Considering the floristic aspect, most of the species detected in the Mar Piccolo have also been reported in similar Mediterranean environments such as Greek and Cypriot TWSs (
Likewise, the chorological spectrum of Mar Piccolo phytobenthos has seen considerable fluctuation. The present situation is quite different from that of Italian marine flora in general, where the Atlantic and Mediterranean elements are prevalent, while the Indo-Pacific and Circum-Tropical elements are practically negligible (
The total of 129 taxa recorded in the Mar Piccolo in the 2011–2018 period is probably an underestimate, since only four coastal stations were seasonally sampled from a total surface area of about 21 km2. However, it was still comparable with that of other Mediterranean TWSs, considering that some of these environments have larger surface areas and probably offer a higher range of environmental conditions (Table
Number of seaweed and phanerogams taxa in some Mediterranean TWS. S=number of seaweeds; P=number of phanerogams.
TWS | S | P | Surface | Reference |
---|---|---|---|---|
Mar Piccolo | 126 | 2 | 21 km2 | this study |
Venice Lagoon | 296 | 5 | 432 km2 |
|
Thau Lagoon (France) | 179 | 2 | 75 km2 | Boudouresque et al. 2010 |
Nadoor Lagoon (Morocco) | 110 | 2 | 114 km2 |
|
Stagnone of Marsala | 108 | 4 | 20 km2 |
|
Mar Menor (Spain) | 69 | 2 | 135 km2 |
|
Orbetello Lagoon | 68 | 3 | 25 km2 |
|
Marano and Grado Lagoon | 41 | 4 | 160 km2 |
|
Acquatina Lake | 38 | 2 | 0.45 km2 |
|
Ganzirri Lake | 32 | 3 | 0.34 km2 |
|
Lesina Lagoon | 36 | 2 | 51.36 km2 |
|
Caprolace Lagoon | 28 | 3 | 2.26 km2 |
|
Faro Lake | 28 | 3 | 0.26 km2 |
|
Fogliano Lagoon | 10 | 2 | 3.95 km2 |
|
Quantitative dominance showed the same fluctuations amongst both years and seasons. In 1989, Gracilariaceae and Solieriaceae had the highest standing crop throughout the basin (
Considering differences in spatial patterns, it is well known that coastal lagoons are “a mosaic of assemblages”, mainly depending on abiotic factors (
Considering a longer time period of about one century, complemented with information on changes in both the demography of Taranto, based on decade-long census data (https://tinyurl.com/yacl8t6t) and the forcing factors affecting the Mar Piccolo, changes in phytobenthos can speculatively be related to human pressure.
In the 1920s, Taranto had fewer than 100,000 inhabitants (https://tinyurl.com/yacl8t6t) and the only anthropogenic factors affecting the Mar Piccolo were mussel breeding and the shipyard of the Italian Royal Navy, established in 1889 (
In the 1940s, a change was already observed, linked to the presence of the First Squadron of the Italian Royal Navy fleet during the Second World War (Pierpaoli 1959). Considerable quantities of both fuel and the residues of smoke bombs were observable in the areas where the ships were moored and where the anti-aircraft defences were placed. This probably adversely affected seawater quality and seaweed assemblages, with Ulvales growing on the docks and a noticeably lower number of Ochrophyta and Rhodophyta species (Pierpaoli 1959).
The period from the 1950s to the 1970s saw a considerable increase in the population of Taranto (https://tinyurl.com/yacl8t6t), linked to the development of new economic activities, chief amongst which was the ITALSIDER steelworks (now Arcelor-Mittal Italy). New districts were built and the town expanded (https://tinyurl.com/ycm2y68w,
At the beginning of the 1980s, the population of Taranto peaked along with the industrial activities (https://tinyurl.com/yacl8t6t). The demographic and economic boom resulted in the creation of 14 sewage outlets, both urban and military, which discharged untreated effluent into the Mar Piccolo until the late 1990s. The most immediate consequences of these factors were the eutrophication of the basin’s waters and an increase in turbidity due to suspended particulate matter (
The beginning of the new century saw the start of a new era; the population in Taranto began to decrease (https://tinyurl.com/yacl8t6t) and nine urban sewage outlets were closed (
The Italian LTER network combines a number of marine, terrestrial and freshwater ecosystems where ecological data have been collected for several decades. By means of interdisciplinary activities and cross-ecosystem research, it aims to make historic datasets, in series of up to a century long, available for ecological research and preserve them for future generations (
In the Mar Piccolo, LTER studies enabled the analysis of historic qualitative and quantitative data and made it possible to draw up a history of its phytobenthos over about one century. On the basis of multidisciplinary observations of the whole basin conducted over many years (Petrocelli et al. unpublished data), it may be assumed that human activities, directly or indirectly, were the main cause of the changes. However, it is important to highlight that the basin showed high resilience. Indeed, following the removal of most of the urban wastewater discharges, which were the main cause of organic pollution, the basin was able to return to better conditions in just a decade. Therefore, it is realistic to hope that the Mar Piccolo could once again become the paradisiacal place described in ancient times, if only individual users adopt good habits. Indeed, strong pressures on the basin also arise from the careless dropping of litter of all types, such as mollusc shells, plastics and engine oil from fishing boats. In any case, long-term studies are ongoing, so it will be possible to monitor the changes year by year.
This research was carried out within the framework of the LTER network. The systematic sampling activities performed in these years benefited from the invaluable help of Giuseppe Portacci and Manuela Belmonte. The authors wish to thank Angél Pérez Ruzafa, the anonymous reviewer and the Subject Editor for their valuable suggestions that helped to improve the manuscript. George Metcalf revised the English text.
Biomass data in 2011–2012
Data type: measurement
Explanation note: Mean biomass values (g m-2), standard deviation and total number of taxa measured in each station and in each season in 2011–2012.
Biomass data in 2012–2013
Data type: measurement
Explanation note: Mean biomass values (g m-2), standard deviation and total number of taxa measured in each station and in each season in 2012–2013.
Biomass data in 2013–2014
Data type: measurement
Explanation note: Mean biomass values (g m-2), standard deviation and total number of taxa measured in each station and in each season in 2013–2014.
Biomass data in 2014–2015
Data type: measurement
Explanation note: Mean biomass values (g m-2), standard deviation and total number of taxa measured in each station and in each season in 2014–2015.
Biomass data in 2015–2016
Data type: measurement
Explanation note: Mean biomass values (g m-2), standard deviation and total number of taxa measured in each station and in each season in 2015–2016.
Biomass data in 2016–2017
Data type: measurement
Explanation note: Mean biomass values (g m-2), standard deviation and total number of taxa measured in each station and in each season in 2016–2017.
Biomass data in 2017–2018
Data type: measurement
Explanation note: Mean biomass values (g m-2), standard deviation and total number of taxa measured in each station and in each season in 2017–2018.