Checklist of marine demersal fishes captured by the pair trawl fisheries in Southern (RJ-SC) Brazil

Lista de peixes demersais marinhos capturados pela pesca de parelha no Sul (RJ-SC) do Brasil

Abstract:

Demersal fishery resources are abundant on continental shelves, on the tropical and subtropical coasts, making up a significant part of the marine environment. Marine demersal fishery resources are captured by various fishing methods, often unsustainably, which has led to the depletion of their stocks. In order to inventory the marine demersal ichthyofauna on the Southern Brazilian coast, as well as their conservation status and distribution, this study analyzed the composition and frequency of occurrence of fish captured by pair trawling in 117 fishery fleet landings based in the State of São Paulo between 2005 and 2012. The ichthyofauna consisted of 245 species (81 families, 32 orders and 2 classes). Among the species, 50 species were classified as constant, 38 accessory and 157 accidental. Still, 13.47% of the species were listed as endangered; 35.29% of the Chondrichthyes and 11.85% of the Actinopterygii (15.1% of the total species) belong to the endemic fauna of the Biogeographic Province of Argentina. The richness was directly related to the oceanographic features of the study area, which determined the southern boundary of occurrence of several tropical species and the northern boundary of occurrence for temperate species. This is the region with the highest abundance of fishery resources in Brazil.

Keywords:
ichthyofauna; inventory; fishery resources; conservation

Resumo:

Nas costas tropicais e subtropicais, sobre as plataformas continentais, os recursos pesqueiros demersais são abundantes e considerados uma parcela importante do ambiente marinho. Os recursos pesqueiros demersais marinhos são capturados por diversas modalidades de pesca, muitas vezes de forma insustentável, o que vem provocando o esgotamento de seus estoques. Com o objetivo de inventariar a ictiofauna demersal marinha da costa sudeste-sul do Brasil, assim como seu status de conservação e distribuição, foram analisadas a composição e frequência de peixes capturados em 117 cruzeiros de pesca entre 2005 e 2012 pela frota de arrasto de parelha sediada no Estado de São Paulo. A ictiofauna foi composta por 245 espécies (81 famílias, 32 ordens e 02 classes), sendo 50 consideradas como constantes, 38 acessórias e 157 acidentais; 13.47% apresentam algum grau de ameaça; 35.29% dos Chondrichthyes e 11.85% dos Actinopterygii (15.1% do total) pertencem à fauna endêmica da Província Zoogeográfica da Argentina. A riqueza observada está diretamente vinculada às características oceanográficas da área de estudo, sendo estas responsáveis pelo limite meridional de ocorrência de várias espécies tropicais e o limite setentrional de espécies de regiões temperadas. Esta é a região com maior abundância de recursos pesqueiros do Brasil.

Palavras-chave:
ictiofauna; inventário; recursos pesqueiros; conservação

Introduction

Demersal fishery resources are abundant on continental shelves, on the tropical and subtropical coasts, making up a significant part of the marine environment. Oceanographic conditions influence the quality and potential of such resources, as well as regionally limit the use of certain types of vessels and fishing gear (Yáñez-Arancibia & Sánches-Gil 1988, Dias-Neto & Ximenes 1998).

The capture of demersal species has been carried out by artisanal and industrial fishing for several decades and it is one of the most important activities for the fishing industry in Brazil (Castro et al. 2003). Small-scale fishing consists of beach seining, gillnets, long-lines, stern trawlers or double rig trawlers along estuarine and coastal and regions. Industrial fishing is carried out by medium and large-sized vessels using pair trawls and otter trawlnets, stern trawlers or double rig trawlers, gillnets and long-lines, as well as fishing with traps, coves and purse seines (Sudepe 1985, Valentini et al. 1991, Ibama 1993, 1995, Haimovici 1997).

The industrial pair trawling stands out among various modalities and categories of fishing aimed to demersal fish. Species belonging to the family Sciaenidae (Weakfishes), such as Micropogonias furnieri (Whitemouth croaker), Macrodon atricauda (Southern King Weakfish) and Cynoscion jamaicensis (Jamaica weakfish); in addition to species of the family Balistidae, such as Balistes capriscus (Grey triggerfish) and family Ariidae, such as Genidens barbus (White sea catfish) are the most targeted along with elasmobranchs (Valentini et al. 1991, Ibama 1993, 1995, Haimovici 1997, Castro et al. 2003, 2007, Castro & Tutui 2007).

According to Caddy & Sharp (1986), it is necessary to bear in mind that exploited species are linked to others that may be essential for the economic and biological productivity of the ecosystem. This is the only way to minimize the adverse effects of fishing and not to compromise the system’s ability to sustain productive human activity. Therefore, the proper management of exploited resources and aquaculture requires knowledge about the dynamics of communities/populations, their ecological interactions, the integration with physicochemical and biological processes at spatial and temporal scales, as well as changes in fishery dynamics (Yáñes-Arancibia et al. 1985, Davis & Anderson 1989, Aryuthaka & Thumbthimsang 1992, Castelo et al. 2007, Castro & Tutui 2007).

This study provides a checklist of marine demersal ichthyofauna of the southern coasts of Brazil captured by the fishery fleet pair trawling based in the state of São Paulo and comments upon the composition, frequency of occurrence, conservation status and geographical distribution of the recorded species.

Material and Methods

The pair trawling fleet operation area had its limits under the coordinates 23º04’S / 42º20’W near Cabo Frio - RJ and 29º19’S / 49º41’W in Torres - RS. Prior to SMA-SP Resolution No. 69 of 2009, activities were concentrated on the south coast of São Paulo under the coordinates 24º31’S / 46º39’W near Peruíbe - SP and 28º37’S / 48º37’W near Cabo de Santa Marta - SC ( Figure 1).

Figure 1
Areas of legal practice of pair trawling fishing in the State of São Paulo (Espírito Santo to Rio Grande do Sul). The study area is highlighted in black.

The fleet operated between a depth of 14 and 55 meters (m ± SD = 29.2 ± 9.7m). For each year, the following means ± standard deviation and amplitudes were observed: 30.4 ± 8.6m (22 and 50m) in 2005; 27.9 ± 7.5m (20 and 42m) in 2006; 33.2 ± 13.9m (14 and 52m) in 2007; 25.2 ± 11.9 (15 and 55m) in 2008; 28.5 ± 10.9m (14 and 50m) in 2009; 30.1 ± 9.7m (26 and 52m) in 2010; 30.4 ± 6.3m (26 and 50m) in 2011; and 30.9 ± 6.6m (26 and 44m) in 2012.

The ichthyofauna from 117 fishery fleet landings of the industrial pair trawling fleet based in the State of São Paulo were sampled from March 2005 to May 2012. The sampling order was randomly. Only landings at the ports located in the municipalities of Santos and Guarujá-SP, Brazil were recorded.

The following sampling routine was performed for qualitative purposes: (a) the whole screening operation of the marketable product was followed up by collecting all rejected material among commercial products and also part of this products; (b) samples of the rejected and discarded products that fishermen sampled during the whole fishing operation were collected for further evaluation in the laboratory; c) co-ordinates of the most distant bids, as well as of the main fishing grounds and their respective depths, were annotated with the masters. Samples of rejected and discarded fauna were also obtained through 15 shipments to the studied fleet.

The collected specimens were referred to the laboratory in iceboxes. Samples were identified, counted and had their total length measured (in mm). Subsequently, the specimens were fixed in 10% formaldehyde and preserved in 70% ethyl alcohol. Voucher specimens of the some species collected during the study were deposited in the regional collection of fish from the Atlantic Forest coast of the “Acervo Zoológico da Universidade Santa Cecília” (AZUSC), in the city of Santos-SP, Brazil.

Fish species in the samples were identified according to Figueiredo (1977), Figueiredo & Menezes (1978, 1980, 2000), Fischer (1978), Menezes & Figueiredo (1980, 1985), Compagno (1984, 1988), Cervigón et al. (1992), Carvalho-Filho (1999), Moura et al. (2001), Carpenter (2002), Fischer et al. (2004), Marceniuk (2005), Leis (2006), Moura & Lindeman (2007), Sampaio & Nottingham (2008), Carvalho-Filho et al. (2010), Gomes et al. (2010), McBride et al. (2010), Rosa & Gadig (2010), Tavera et al. (2011, 2012), Ruocco et al. (2012), Frable et al. (2013), Menezes et al. (2015), Knudsen & Clements (2016), Last et al. (2016a, b, c); Marceniuk et al. (2016). The systematics and nomenclature followed Nelson et al. (2016) and Eschmeyer et al. (2017), respectively.

Species were classified from their frequencies of occurrence and according to the Dajoz’s (1983) scale as “constant” when present in more than 50% of the samples; “accessory” when frequencies ranged from 25 to 50% and “accidental” when frequencies were below 25%.

The conservation status of each species were based on the International Red List of threatened species (IUCN - International Union for Conservation of Nature 2017) and the Brazilian Federal list of threatened species, Ordinance number 445 of the MMA - Brazilian Environment Ministry, December 17th 2014 (MMA 2014).

Species were classified in the biogeographical categories proposed by Floeter et al. (2008) and Luiz Jr. et al. (2008).

Results

Of the 117 fishery fleet landings, the majority occurred in 2011 (20.52%) and the minority in 2010 (5.98%) (Table 1).

The richness consisted of 245 species (Table 2) distributed over 2 classes, 32 orders, 81 families and 170 genera. Out of the total species analyzed, 86.12% of the species belong to Class Actinopterygii and 13.88% to Class Chondrichthyes (Table 3). Of the 81 families comprising the captured ichthyofauna, six families had the highest species richness (33.5% of the total), namely Carangidae (6), Sciaenidae (19), Paralichthyidae (11), Haemulidae and Serranidae (10) and Engraulidae (8).

Table 1
Distribution of the number of fishing operations (N) and percentage (%) in relation to the study period.

	2005	2006	2007	2008	2009	2010	2011	2012	Total
N	10	17	13	18	20	7	24	8	117
%	8.5	14.5	11.1	15.3	17.0	5.9	20.5	6.8	100

Table 2
Ichthyofauna caught by the pair trawling fleet of the state of São Paulo, Dajoz scale (DS): Co = constant, Ac = accessory, Ad = accidental; geographic distribution (GD): CT = circumtropical, TA = Trans-Atlantic (both sides of the Atlantic), WA = Western Atlantic (Northern and Southwestern Atlantic), SWA = Southern West Atlantic (from northern Brazil to Argentina), SSWA = Southern South West Atlantic (species with affinity to temperate areas occurring from Argentina and Uruguay to southern Brazil), Ca = Caribbean (from Florida to Venezuela), Br = Brazilian Province (the area between the Orinoco Delta in Venezuela and Santa Catarina in Brazil) and EP = Eastern Pacific; conservation status in the IUCN Red List (IUCN 2017), in the Brazilian list (MMA 2014): X = Not Evaluated; LC = Least Concern; DD = Data Deficient; VU = Vulnerable; NT = Near-Threatened; END = Endangered, CR = Critically Endangered; vouchers.

Order/ Family/ Species	DS	GD	IUCN	MMA	Voucher
CARCHARHINIFORMES
Triakidae
Mustelus schmitti Springer, 1939	Ad	SSWA	END	CR	AZUSC3405
Carcharhinidae
Carcharhinus brachyurus (Günther, 1870)	Ad	CT	NT		AZUSC3791
Carcharhinus falciformis (Bibron, 1839)	Ad	CT	VU		AZUSC3354
Carcharhinus limbatus (Valenciennes, 1839)	Ad	CT	NT		AZUSC3350
Galeocerdo cuvier (Péron & Lesueur, 1822)	Ad	CT	NT		AZUSC3383
Rhizoprionodon lalandii (Valenciennes, 1839)	Co	WA	DD		AZUSC3293
Rhizoprionodon porosus (Poey, 1861)	Ad	Ca+SWA	LC		AZUSC4811
Sphyrnidae
Sphyrna lewini (Griffith & Smith, 1834)	Ad	CT	END	CR	AZUSC0893
Sphyrna zygaena (Linnaeus, 1758)	Ad	CT	VU	CR	AZUSC3780
SQUATINIFORMES
Squatinidae
Squatina guggenheim Marini, 1936	Ac	SSWA	END	CR	AZUSC3203
TORPEDINIFORMES
Torpedinidae
Tetronarce puelcha (Lahille, 1926)	Ad	SSWA	DD	VU	AZUSC3543
Narcinidae
Narcine brasiliensis (Olfers, 1831)	Co	WA	DD		AZUSC3424
RAJIFORMES
Rajidae
Atlantoraja castelnaui (Miranda Ribeiro, 1907)	Ac	SSWA	END	END	AZUSC3147
Atlantoraja cyclophora (Regan, 1903)	Ad	SSWA	VU		AZUSC3346
Atlantoraja platana (Günther, 1880)	Ad	SSWA	VU		AZUSC3349
Psammobatis extenta (Garman, 1913)	Ad	SSWA	LC		AZUSC3446
Psammobatis lentiginosa McEachran, 1983	Ad	SSWA	DD		AZUSC3442
Rioraja agassizii (Müller & Henle, 1841)	Co	SSWA	VU	END	AZUSC3432
Sympterygia bonapartii Müller & Henle, 1841	Ad	SSWA	DD	END	AZUSC3782
PRISTIFORMES
Rhinobatidae
Pseudobatos horkelii (Müller & Henle, 1841)	Ac	SSWA	CR	CR	AZUSC3454
Pseudobatos percellens (Walbaum, 1792)	Co	TA	NT		AZUSC3456
Zapteryx brevirostris (Müller & Henle, 1841)	Co	WA	VU	VU	AZUSC3591
MYLIOBATIFORMES
Dasyatidae
Hypanus americanus (Hildebrand & Schroeder, 1928)	Ac	WA	DD		AZUSC3355
Dasyatis hypostigma Santos & Carvalho, 2004	Co	SSWA	DD		AZUSC3359
Hypanus guttatus (Bloch & Schneider,1801)	Ad	Ca+Br	DD		AZUSC3358
Pteroplatytrygon violacea (Bonaparte, 1832)	Ad	CT	LC		AZUSC3786
Gymnuridae
Gymnura altavela (Linnaeus, 1758)	Co	TA	VU	CR	AZUSC3387
Myliobatidae
Aetobatus narinari (Euphrasen, 1790)	Ad	CT	NT		AZUSC3291
Mobula hypostoma (Bancroft, 1831)	Ad	CT	DD	VU	AZUSC3400
Mobula thurstoni (Lloyd, 1908)	Ad	CT	NT	VU	AZUSC3403
Myliobatis freminvillei Lesueur, 1824	Ad	WA	DD	END	AZUSC3412
Myliobatis goodei Garman, 1885	Ad	WA	DD	CR	AZUSC3418
Rhinoptera bonasus (Mitchill, 1815)	Ad	WA	NT		AZUSC3783
Rhinoptera brasiliensis Müller, 1836	Ad	WA	END	CR	AZUSC0877
ELOPIFORMES
Elopidae
Elops smithi McBride, Rocha, Ruiz-Carus & Bowen, 2010	Ad	WA	DD		AZUSC1360
ALBULIFORMES
Albulidae
Albula vulpes (Linnaeus, 1758)	Ac	WA	NT		AZUSC1742
ANGUILLIFORMES
Muraenidae
Gymnothorax conspersus Poey, 1867	Ad	WA	X		AZUSC3719
Gymnothorax funebris Ranzani, 1839	Ad	WA	LC		AZUSC2108
Gymnothorax moringa (Cuvier, 1829)	Ad	WA	LC		AZUSC2710
Gymnothorax ocellatus Agassiz, 1831	Ac	Ca+SWA	LC		AZUSC2010
Muraena retifera Goode & Bean, 1882	Ad	WA	LC		AZUSC2299
Ophichthidae
Echiophis intertinctus (Richardson, 1848)	Ad	WA	LC		AZUSC2109
Ophichthus ophis (Linnaeus, 1758)	Ad	TA	LC		AZUSC2256
Ophichthus gomesii (Castelnau, 1855)	Ac	WA	LC		AZUSC3673
Muraenosocidae
Cynoponticus savanna (Bancroft, 1831)	Ad	Ca+Br	LC		AZUSC3761
Congridae
Conger orbignianus Valenciennes, 1837	Ad	SSWA	X		AZUSC1120
CLUPEIFORMES
Pristigasteridae
Chirocentrodon bleekerianus (Poey, 1867)	Ac	Ca+Br	LC		AZUSC1488
Pellona harroweri (Fowler, 1917)	Co	Ca+Br	LC		AZUSC1477
Engraulidae
Anchoa filifera (Fowler, 1915)	Ad	Ca+Br	LC		AZUSC1490
Anchoa marinii Hildebrand, 1943	Ad	SSWA	LC		AZUSC4620
Anchoa spinifer (Valenciennes, 1848)	Ad	Ca+Br+EP	LC		AZUSC4621
Anchoa tricolor (Spix & Agassiz, 1829)	Ad	SWA	LC		AZUSC4625
Anchovia clupeoides (Swainson, 1839)	Ad	Ca+Br	LC		AZUSC1476
Anchoviella lepidentostole (Fowler, 1911)	Ad	Br	LC		AZUSC1487
Engraulis anchoita Hubbs & Marini, 1935	Ad	SSWA	X		AZUSC2160
Lycengraulis grossidens (Spix & Agassiz, 1829)	Ad	Br+SSWA	LC		AZUSC4663
Clupeidae
Brevoortia pectinata (Jenyns, 1842)	Ad	SSWA	LC		AZUSC3897
Harengula clupeola (Cuvier, 1829)	Co	WA	LC		AZUSC1328
Opisthonema oglinum (Lesueur, 1818)	Ad	WA	LC		AZUSC1327
Sardinella brasiliensis (Steindachner, 1879)	Ad	SSWA	X		AZUSC2164
SILURIFORMES
Ariidae
Aspistor luniscutis (Valenciennes, 1840)	Co	Br	X		AZUSC1976
Bagre bagre (Linnaeus, 1766)	Ac	Ca+Br	LC		AZUSC1367
Bagre marinus (Mitchill, 1815)	Ad	WA	LC		AZUSC4602
Cathorops spixii (Agassiz, 1829)	Ad	Ca+Br	X		AZUSC1518
Genidens barbus (Lacepède, 1803)	Co	Br+SSWA	X	END	AZUSC1371
Genidens genidens (Cuvier, 1829)	Co	Br+SSWA	LC		AZUSC1756
AULOPIFORMES
Synodontidae
Synodus bondi Fowler, 1939	Co	WA	LC		AZUSC2090
Trachinocephalus myops (Forster, 1801)	Ac	CT	LC		AZUSC1470
GADIFORMES
Merlucciidae
Merluccius hubbsi Marini, 1933	Ad	SSWA	X		AZUSC1288
Gadidae
Urophycis brasiliensis (Kaup, 1858)	Ad	SSWA	X		AZUSC2122
HOLOCENTRIFORMES
Holocentridae
Holocentrus adscensionis (Osbeck, 1765)	Ad	TA	LC		AZUSC1699
OPHIDIIFORMES
Ophidiidae
Genypterus brasiliensis Regan, 1903	Ad	SSWA	X		AZUSC0905
Ophidion holbrookii Putnam, 1874	Ac	WA	LC	CR	AZUSC1352
Raneya brasiliensis (Kaup, 1856)	Ac	SSWA	X		AZUSC1811
BATRACHOIDIFORMES
Batrachoididae
Opsanus beta (Goode & Bean, 1880)	Ad	Ca	LC		AZUSC3144
Porichthys porosissimus (Cuvier, 1829)	Co	SSWA	X		AZUSC1146
Thalassophryne montevidensis (Berg, 1893)	Ad	SWA	X		AZUSC3088
MUGILIFORMES
Mugilidae
Mugil curema Valenciennes, 1836	Ad	TA+EP	LC		AZUSC1814
Mugil liza Valenciennes, 1836	Ad	WA	DD		AZUSC2698
Polynemidae
Polydactylus oligodon (Günther, 1860)	Ad	WA	LC		AZUSC2089
Polydactylus virginicus (Linnaeus, 1758)	Ac	WA	LC		AZUSC2011
BELONIFORMES
Exocoetidae
Cheilopogon melanurus (Valenciennes, 1847)	Ad	TA	LC		AZUSC2709
Hemiramphidae
Hemiramphus balao Lesueur, 1821	Ad	TA	LC		AZUSC4008
Hemiramphus brasiliensis (Linnaeus, 1758)	Ad	TA	LC		AZUSC4604
Hyporhamphus unifasciatus (Ranzani, 1841)	Ad	WA+EP	LC		AZUSC4618
Belonidae
Ablennes hians (Valenciennes, 1846)	Ad	CT	LC		AZUSC2020
Strongylura marina (Walbaum, 1792)	Ad	WA	LC		AZUSC4672
Strongylura timucu (Walbaum, 1792)	Ad	WA	LC		AZUSC4603
Tylosurus acus (Lacepède, 1803)	Ad	TA	LC		AZUSC4619
CARANGIFORMES
Coryphaenidae
Coryphaena hippurus Linnaeus, 1758	Ad	CT	LC		AZUSC4601
Rachycentridae
Rachycentron canadum (Linnaeus, 1766)	Ad	CT	LC		AZUSC2244
Echeneidae
Echeneis naucrates Linnaeus, 1758	Ad	CT	LC		AZUSC3232
Carangidae
Alectis ciliaris (Bloch, 1787)	Ad	CT	LC		AZUSC1389
Caranx crysos (Mitchill, 1815)	Ac	TA	LC		AZUSC1319
Caranx hippos (Linnaeus, 1766)	Ac	TA	LC		AZUSC1658
Caranx latus Agassiz, 1831	Ad	TA	LC		AZUSC1322
Chloroscombrus chrysurus (Linnaeus, 1766)	Co	TA	LC		AZUSC1344
Decapterus punctatus (Cuvier, 1829)	Ad	TA	LC		AZUSC1792
Hemicaranx amblyrhynchus (Cuvier, 1833)	Ac	WA	LC		AZUSC1928
Oligoplites palometa (Cuvier, 1832)	Ad	Ca+Br	LC		AZUSC3037
Oligoplites saliens (Bloch, 1793)	Co	Ca+SWA	LC		AZUSC1317
Oligoplites saurus (Bloch & Schneider, 1801)	Ad	WA	LC		AZUSC3771
Parona signata (Jenyns, 1841)	Ad	SSWA	X		AZUSC3604
Pseudocaranx dentex (Bloch & Schneider, 1801)	Ac	CT	LC		AZUSC1329
Selar crumenophthalmus (Bloch, 1793)	Ad	Ca+Br	LC		AZUSC2447
Selene setapinnis (Mitchill, 1815)	Co	WA	LC		AZUSC1323
Selene vomer (Linnaeus, 1758)	Co	WA	LC		AZUSC1388
Seriola dumerili (Risso, 1810)	Ad	CT	LC		AZUSC3668
Seriola fasciata (Bloch, 1793)	Ad	TA	LC		AZUSC3677
Seriola lalandi Valenciennes, 1833	Ad	CT	LC		AZUSC2634
Trachinotus carolinus (Linnaeus, 1766)	Co	WA	LC		AZUSC1336
Trachinotus falcatus (Linnaeus, 1758)	Ad	WA	LC		AZUSC3149
Trachinotus goodei Jordan & Evermann, 1896	Ac	WA	LC		AZUSC2016
Trachinotus marginatus Cuvier, 1832	Ad	SSWA	X		AZUSC2678
Trachurus lathami Nichols, 1920	Ad	WA	LC		AZUSC1122
Uraspis secunda (Poey, 1860)	Ad	CT	LC		AZUSC1963
ISTIOPHORIFORMES
Sphyraenidae
Sphyraena guachancho Cuvier, 1829	Co	TA	LC		AZUSC1341
Sphyraena tome Fowler, 1903	Ac	SSWA	X		AZUSC1602
Xiphiidae
Xiphias gladius Linnaeus, 1758	Ad	CT	LC		AZUSC4598
PLEURONECTIFORMES
Paralichthyidae
Citharichthys arenaceus Evermann & Marsh, 1900	Ac	WA	LC		AZUSC4623
Citharichthys macrops Dresel, 1885	Ad	WA	LC		AZUSC1678
Cyclopsetta chittendeni Bean, 1895	Ad	Ca+Br	LC		AZUSC1651
Etropus crossotus Jordan & Gilbert, 1882	Ad	WA+EP	LC		AZUSC2159
Etropus longimanus Norman, 1933	Ad	SSWA	X		AZUSC4622
Paralichthys brasiliensis (Ranzani, 1842)	Ac	SWA	X		AZUSC1521
Paralichthys orbignyanus (Valenciennes, 1839)	Ad	SSWA	X		AZUSC1926
Paralichthys patagonicus Jordan, 1889	Ac	SSWA+EP	X		AZUSC1392
Syacium papillosum (Linnaeus, 1758)	Ad	WA	LC		AZUSC1549
Syacium micrurum Ranzani, 1842	Ad	WA	LC		AZUSC3036
Xystreurys rasile (Jordan, 1891)	Ad	SSWA	X		AZUSC4600
Bothidae
Bothus ocellatus (Agassiz, 1831)	Ad	WA	LC		AZUSC3086
Bothus robinsi Topp & Hoff, 1972	Ad	WA	LC		AZUSC2088
Achiridae
Achirus declivis Chabanaud, 1940	Ad	WA	LC		AZUSC2245
Gymnachirus nudus Kaup, 1858	Ac	WA	LC		AZUSC1949
Trinectes paulistanus (Miranda Ribeiro, 1915)	Ad	Ca+Br	LC		AZUSC4664
SYNGNATHIFORMES
Fistulariidae
Fistularia petimba Lacepède, 1803	Ad	TA	LC		AZUSC1330
Fistularia tabacaria Linnaeus, 1758	Co	TA	LC		AZUSC2018
Dactylopteridae
Dactylopterus volitans (Linnaeus, 1758)	Co	TA	LC		AZUSC1147
SCOMBRIFORMES
Gempylidae
Thyrsitops lepidopoides (Cuvier, 1832)	Ad	SSWA	X		AZUSC1148
Trichiuridae
Evoxymetopon taeniatus Gill, 1863	Ad	Ca+SWA+EP	LC		AZUSC2939
Trichiurus lepturus Linnaeus, 1758	Co	CT	LC		AZUSC1391
Scombridae
Scomber japonicus Houttuyn, 1782	Ad	CT	LC		AZUSC2286
Scomberomorus brasiliensis Collette, Russo & Zavala-Camin, 1978	Co	Ca+Br	LC		AZUSC1369
Scomberomorus cavalla (Cuvier, 1829)	Ad	WA	LC		AZUSC4599
Thunnus alalunga (Bonnaterre, 1788)	Ad	CT	NT		AZUSC1933
Centrolophidae
Seriolella porosa Guichenot, 1848	Ad	SWA+EP	X		AZUSC2636
Stromateidae
Peprilus xanthurus (Quoy & Gaimard, 1825)	Co	SWA	X		AZUSC1333
TRACHINIFORMES
Pinguipedidae
Pseudopercis semifasciata (Cuvier, 1829)	Ad	SSWA	X		AZUSC2119
Percophidae
Percophis brasiliensis Quoy & Gaimard, 1825	Ad	SSWA	X		AZUSC1145
Uranoscopidae
Astroscopus sexspinosus (Steindachner, 1876)	Ad	SSWA	X		AZUSC2113
Astroscopus y-graecum (Cuvier, 1829)	Ad	WA	LC		AZUSC2014
LABRIFORMES
Labridae
Xyrichtys novacula (Linnaeus, 1758)	Ad	WA	LC		AZUSC1157
Scaridae
Nicholsina usta (Valenciennes, 1840)	Ad	WA	LC		AZUSC3644
Sparisoma axillare (Steindachner, 1878)	Ad	Br	DD	VU	AZUSC1828
Sparisoma frondosum (Agassiz, 1831)	Ad	Ca+Br	DD	VU	AZUSC1799
PERCIFORMES
Centropomidae
Centropomus parallelus Poey, 1860	Ad	WA	LC		AZUSC1372
Centropomus undecimalis (Bloch, 1792)	Ac	WA	LC		AZUSC1363
Gerreidae
Diapterus auratus Ranzani, 1842	Ad	WA	LC		AZUSC3898
Diapterus rhombeus (Cuvier, 1829)	Co	Ca+Br	LC		AZUSC1325
Eucinostomus argenteus Baird & Girard, 1855	Ac	WA+EP	LC		AZUSC1326
Eucinostomus gula (Quoy & Gaimard, 1824)	Ad	WA	LC		AZUSC1359
Eucinostomus melanopterus (Bleeker, 1863)	Ad	TA	LC		AZUSC1625
Eugerres brasilianus (Cuvier, 1830)	Ad	WA	LC		AZUSC1875
Polyprionidae
Polyprion americanus (Bloch & Schneider, 1801)	Ad	CT	DD	CR	AZUSC3656
Mullidae
Pseudupeneus maculatus (Bloch, 1793)	Ad	WA	LC		AZUSC1661
Upeneus parvus Poey, 1852	Ac	WA	LC		AZUSC1351
Kyphosidae
Kyphosus vaigiensis (Quoy & Gaimard, 1825)	Ad	TA	LC		AZUSC3867
Kyphosus sectatrix (Linnaeus, 1758)	Ad	TA	LC		AZUSC0372
Serranidae
Diplectrum formosum (Linnaeus, 1766)	Co	WA	LC		AZUSC1331
Diplectrum radiale (Quoy & Gaimard, 1824)	Ac	WA	LC		AZUSC1355
Dules auriga Cuvier, 1829	Ad	SSWA	X		AZUSC1140
Epinephelus itajara (Lichtenstein, 1822)	Ad	WA	CR	CR	AZUSC1043
Epinephelus marginatus (Lowe, 1834)	Ad	WA+EP	END	VU	AZUSC2377
Epinephelus morio (Valenciennes, 1828)	Ad	WA	NT	VU	AZUSC3646
Hyporthodus niveatus (Valenciennes, 1828)	Ac	WA	VU	VU	AZUSC1600
Mycteroperca acutirostris (Valenciennes, 1828)	Ad	Ca+Br	LC		AZUSC1646
Rypticus randalli Courtenay, 1967	Ad	Ca+Br	LC		AZUSC3721
Serranus atrobranchus (Cuvier, 1829)	Ad	WA	LC		AZUSC3676
Pomatomidae
Pomatomus saltatrix (Linnaeus, 1766)	Co	CT	VU		AZUSC1630
Priacanthidae
Priacanthus arenatus Cuvier, 1829	Co	TA	LC		AZUSC1339
Pomacanthidae
Pomacanthus paru (Bloch, 1787)	Ad	WA	LC		AZUSC1342
Malacanthidae
Caulolatilus chrysops (Valenciennes, 1833)	Ad	WA	LC		AZUSC1605
Haemulidae
Anisotremus surinamensis (Bloch, 1791)	Ad	WA	DD		AZUSC1688
Anisotremus virginicus (Linnaeus, 1758)	Ac	WA	LC		AZUSC1343
Boridia grossidens Cuvier, 1830	Ad	SSWA	X		AZUSC1912
Conodon nobilis (Linnaeus, 1758)	Co	WA	LC		AZUSC1320
Genyatremus cavifrons (Cuvier, 1830)	Ad	Ca+Br	DD		AZUSC1968
Haemulon aurolineatum Cuvier, 1830	Ac	WA	LC		AZUSC1398
Haemulon steindachneri (Jordan & Gilbert, 1882)	Ac	Ca+SWA	LC		AZUSC1913
Orthopristis ruber (Cuvier, 1830)	Co	Ca+SWA	LC		AZUSC1138
Haemulopsis corvinaeformis (Steindachner, 1868)	Co	Ca+SWA	LC		AZUSC1345
Pomadasys crocro (Cuvier, 1830)	Ad	WA	DD		AZUSC3099
Lutjanidae
Lutjanus analis (Cuvier, 1828)	Ad	WA	NT		AZUSC2789
Lutjanus griseus (Linnaeus, 1758)	Ad	WA	LC		AZUSC3100
Lutjanus jocu (Bloch & Schneider, 1801)	Ad	WA	DD		AZUSC4542
Lutjanus synagris (Linnaeus, 1758)	Ad	WA	NT		AZUSC4321
Rhomboplites aurorubens (Cuvier, 1829)	Ad	WA	VU		AZUSC1423
SCORPAENIFORMES
Scorpaenidae
Scorpaena brasiliensis Cuvier, 1829	Ac	WA	LC		AZUSC1599
Scorpaena isthmensis Meek & Hildebrand, 1928	Ad	WA	LC		AZUSC3711
Scorpaena plumieri Bloch, 1789	Ac	WA	LC		AZUSC1611
Triglidae
Prionotus nudigula Ginsburg, 1950	Ad	SSWA	X		AZUSC1609
Prionotus punctatus (Bloch, 1793)	Co	Ca+SWA	LC		AZUSC1141
MORONIFORMES
Ephippidae
Chaetodipterus faber (Broussonet, 1782)	Co	WA	LC		AZUSC1334
ACANTHURIFORMES
Sciaenidae
Ctenosciaena gracilicirrhus (Metzelaar, 1919)	Co	Ca+Br	LC		AZUSC1142
Cynoscion acoupa (Lacepède, 1801)	Ad	Ca+SWA	LC		AZUSC3202
Cynoscion guatucupa (Cuvier, 1830)	Ac	SSWA	X		AZUSC1143
Cynoscion jamaicensis (Vaillant & Bocourt, 1883)	Co	Ca+SWA	LC		AZUSC1386
Cynoscion leiarchus (Cuvier, 1830)	Co	Ca+Br	LC		AZUSC1966
Cynoscion virescens (Cuvier, 1830)	Co	Ca+Br	LC		AZUSC1358
Isopisthus parvipinnis (Cuvier, 1830)	Co	Ca+Br	LC		AZUSC1390
Larimus breviceps Cuvier, 1830	Co	Ca+Br	LC		AZUSC1318
Macrodon atricauda (Günther, 1880)	Co	Br+SSWA	X		AZUSC1498
Menticirrhus americanus (Linnaeus, 1758)	Co	WA	LC		AZUSC1430
Menticirrhus littoralis (Holbrook, 1847)	Ad	WA	LC		AZUSC4730
Micropogonias furnieri (Desmarest, 1823)	Co	Ca+SWA	LC		AZUSC1144
Nebris microps Cuvier, 1830	Ac	Br	LC		AZUSC1467
Paralonchurus brasiliensis (Steindachner, 1875)	Co	Ca+SWA	LC		AZUSC1425
Pogonias cromis (Linnaeus, 1766)	Ad	WA	LC	END	AZUSC3206
Stellifer brasiliensis (Schultz, 1945)	Ad	Br	X		AZUSC1486
Stellifer rastrifer (Jordan, 1889)	Ad	Br+SSWA	LC		AZUSC1520
Umbrina canosai Berg, 1895	Ad	SSWA	X		AZUSC1123
Umbrina coroides Cuvier, 1830	Ad	WA	LC		AZUSC1365
Acanthuridae
Acanthurus chirurgus (Bloch, 1787)	Ad	TA	LC		AZUSC2120
SPARIFORMES
Lobotidae
Lobotes surinamensis (Bloch, 1790)	Ad	CT	LC		AZUSC2635
Sparidae
Archosargus aries (Valenciennes, 1830)	Ad	WA	X		AZUSC3239
Archosargus rhomboidalis (Linnaeus, 1758)	Ac	WA	LC		AZUSC1361
Calamus penna (Valenciennes, 1830)	Ad	WA	LC		AZUSC3035
Calamus pennatula Guichenot, 1868	Ad	WA	LC		AZUSC3089
Diplodus argenteus (Valenciennes, 1830)	Co	SWA	LC		AZUSC1366
Pagrus pagrus (Linnaeus, 1758)	Co	TA	LC		AZUSC1426
LOPHIIFORMES
Lophiidae
Lophius gastrophysus Miranda Ribeiro, 1915	Ad	WA	LC		AZUSC1608
Antennariidae
Antennarius striatus (Shaw, 1794)	Ad	WA	LC		AZUSC3143
Ogcocephalidae
Ogcocephalus vespertilio (Linnaeus, 1758)	Ac	Ca+SWA	X		AZUSC2707
TETRAODONTIFORMES
Ostraciidae
Acanthostracion quadricornis (Linnaeus, 1758)	Ad	WA	LC		AZUSC2117
Balistidae
Balistes capriscus Gmelin, 1789	Co	CT	VU		AZUSC1347
Monacanthidae
Aluterus monoceros (Linnaeus, 1758)	Ac	CT	LC		AZUSC1362
Aluterus heudelotii Hollard, 1855	Ad	CT	LC		AZUSC4517
Stephanolepis hispidus (Linnaeus, 1766)	Co	TA	LC		AZUSC1332
Molidae
Mola mola (Linnaeus, 1758)	Ad	CT	VU		AZUSC3950
Tetraodontidae
Lagocephalus laevigatus (Linnaeus, 1766)	Co	TA	LC		AZUSC1373
Lagocephalus lagocephalus (Linnaeus, 1758)	Ad	CT	LC		AZUSC3234
Sphoeroides greeleyi Gilbert, 1900	Ad	Ca+Br	LC		AZUSC2522
Sphoeroides spengleri (Bloch, 1785)	Ad	TA	LC		AZUSC3330
Sphoeroides testudineus (Linnaeus, 1758)	Ad	WA	LC		AZUSC2163
Diodontidae
Chilomycterus spinosus (Linnaeus, 1758)	Co	SWA	LC		AZUSC1139

Table 3
Total number of ichthyofaunal taxa captured by the pair trawling fleet based in the State of São Paulo.

Class	Chondrichthyes	Actinopterygii
Order	6	26
Family	11	70
Genus	22	148
Species	34	211

From the frequency of occurrence (FO) and the classification proposed by Dajoz (1983) the ichthyofauna consisted of 50 species (20.41%) classified as constants; 38 (15.5%) accessory species and 157 (64.08%) accidental species. Seventeen species (6.34%) had 100% frequency throughout the sample period: Dactylopterus volitans (Flying gurnard), Prionotus punctatus (Bluewing searobin), Chloroscombrus chrysurus (Atlantic bumper), Oligoplites saliens (Castin leatherjacket), Selene setapinnis (Atlantic moonfish), S. vomer (Lookdown), Conodon nobilis (Barred grunt), Orthopristis ruber (Corocoro grunt), Diplodus argenteus (South American silver porgy), Cynoscion jamaicensis (Jamaica weakfish), Menticirrhus americanus (Southern kingcroaker), Micropogonias furnieri (Whitemouth croaker), Chaetodipterus faber (Atlantic spadefish), Trichiurus lepturus (Largehead hairtail), Balistes capriscus (Grey triggerfish), Stephanolepis hispidus (Planehead filefish) and Chilomycterus spinosus (Southern burrfish).

When considering only the Critical, Endangered and Vulnerable categories, both by the IUCN (2017) criteria and by Brazilian federal Legislation (MMA 2014) we observed a total of 33 threatened species (13.47%) in this study. Also, when considering only the threat categories and choosing the most conservative classification, three distinct and worrying scenarios are drawn by correlating the conservation status with the frequency of occurrence (constant, accessory and accidental) (Table 4).

Table 4
Checklist of the threatened species captured according to the International Union for Conservation of Nature (IUCN 2017) and the Brazilian list (MMA 2014) and their frequency of occurrence (Dajoz 1983).

	Accidental	Accessory	Constant
Critically endangered	Mustelus schmitti	Pseudobatos horkelii	Gymnura altavela
	Sphyrna lewini	Squatina guggenheim
	Sphyrna zygaena	Ophidion holbrooki
	Myliobatis goodei
	Rhinoptera brasiliensis
	Epinephelus itajara
	Polyprion americanus
Endangered	Sympterygia bonapartii	Atlantoraja castelnaui	Rioraja agassizii
	Myliobatis freminvillei		Genidens barbus
	Epinephelus marginatus
	Pogonias cromis
Vulnerable	Carcharhinus falciformis	Hyporthodus niveatus	Zapteryx brevirostris
	Atlantoraja cyclophora		Pomatomus saltatrix
	Atlantoraja platana		Balistes capriscus
	Tetronarce puelcha
	Mobula hypostoma
	Mobula thurstoni
	Sparisoma axillare
	Sparisoma frondosum
	Epinephelus morio
	Rhomboplites aurorubens
	Mola mola

Most of the captured species (35.51%) during this study were distributed in the Atlantic Ocean (WA), followed by the SSWA (15.10%), circumtropical (CT) species (13.06%), Trans-Atlantic (TA) species (11.02%), Ca+Br (9.8%), Ca+SWA (4.9%), SWA (2.45%), Br (2.04%), Br+SSWA (2.04%), and WA+EP (1.63%). One species (0.41% of the total) was found in each of the following areas: TA+EP, SWA+EP, SSWA+EP, Ca+Br+EP, Ca+SWA+EP, and Ca.

Discussion

The richness of demersal organisms is directly related to the continental shelf width, sediment types, oceanographic conditions, the geological events that shaped the continental shelf, as well as its positioning relative to the equator characterizing the environmental temperature (Lowe-McConnell 1987, Longhurst & Pauly 2007).

Although many results of this study can be explained by the oceanographic features of the pair trawling fleet operation area in the state of São Paulo, biotic factors interfere with the species distribution through inter- and intraspecific relations, such as the predator-prey effect and the competitive interactions for food (Sanders 1969, Moyle & Cech 1998, Menge & Olson 1990, Barry et al. 1996, Akin et al. 2003).

Overall, the widest and largest continental shelf is in southern Brazil, consisting of muddy sediment and under the influence of the oscillation of the subtropical convergence between the warm waters of the Brazilian current and the cold waters of the Malvinas current. The region has upwelling zones that during the summer and spring cause the South Atlantic Central Water (SACW) to penetrate the shelf reaching the coastal areas with a thermocline of 10 and 15 meters. During the winter, SACW retracts and Tropical Water (TW) fills the space. There is significant water input from continental drainage. Due to such characteristics, the region represents the Southern boundary of occurrence of several tropical species (e.g.: Hypanus guttatus, H. americanus, Anchoa tricolor, Rypticus randalli) and the Northern boundary of temperate species (e.g.: Atlantoraja castelnaui, A. cyclophora, Sardinella brasiliensis, Boridia grossidens). Moreover, the region has the highest abundance of fishery resources in Brazil (Pires-Vanin et al. 1993, Matsuura 1995, Castro & Menezes 1998, Menezes et al. 2003, Braga & Niencheski 2006, Castro et al. 2006, Castro et al. 2008, Amaral & Nallin 2011, Menezes 2011).

According to Menezes et al. (2003), 1,297 species of marine fish, belonging to 36 orders and 192 families, occur in Brazil. The results of this study showed the Southern demersal ichthyofauna accounted for 18.9% of species recorded in Brazil. The families Carangidae, Sciaenidae, Paralichthyidae, Haemulidae, Serranidae and Engraulidae showed high number of species, with most species being demersal, benthic or benthopelagic (Lowe-McConnell 1987).

According to Dajoz’s (1983) scale, we could hypothesize that the common richness of the ichthyofauna by landing is approximately 90 species (constant + accessory). However, this value differs from that observed in landings, which ranged from 49 to 104 species (mean and standard deviation = 68.65 ± 13.20), mainly due to the high number of accidental species.

According to criteria adopted by the International Union for Conservation of Nature (IUCN 2017), species classified as least concern (64.08%) were predominant, followed by not-evaluated species (14.69%), data deficient (8.16%), vulnerable and near-threatened (4.90%), endangered (2.45%) and critically endangered (0.82%). Based on Brazilian federal legislation (MMA 2014), only 10.61% of the species are threatened regarding the conservation status: 4.49% endangered, 3.67% vulnerable and 2.45% endangered. Proportionally, Class Chondrichthyes had higher number of highly endangered species (23.53% critically endangered, 11.76% endangered and 20.59% vulnerable). Based on the three scenarios observed regarding the conservation status of the captured species, the actions of fishery management are priority for the pair trawling fishing operating in the Zoogeographic Province of Argentina. We emphasize this study was carried out in the period before the prohibitions on the capture of several endangered species (MMA 2014); therefore, the continuous monitoring of the composition of this fishing modality is necessary. We also point out there were restrictions of fishing area for pair trawling fishing in the State of São Paulo (SMA 2009). Such restrictions occurred during the last 3 years of this study, which again emphasizes the need for continuous monitoring to verify changes in the composition of the species captured by this type of fishery.

According to Kotas (1991), the pair trawling fishing is characterized as active, multispecies and difficult to manage due to the large number of species captured. The most used management technique in these cases is the restriction of areas and/or periods (High et al. 1969, Caddy 1982), which often causes economic and social problems. There are other ways of ensuring greater selectivity and more sustainability in these fisheries without losing their economic viability, such as adequacy of fishing nets and the use of bycatch reduction devices (BRD). Several BRD models have been developed and tested in different types of environments worldwide, many of which are summarized in the study by Broadhurst (2000). Thus, studies on fishing technology aiming at minimizing the capture of endangered species are essential for biodiversity maintenance.

Caires (2014) listed 120 species (76.67% Actinopterygii and 23.33% Chondrichthyes) as endemic to the zoogeographic province of Argentina. Out of the 92 species of Actinopterygii listed, 24 species (26%) were observed in this study, namely: Conger orbignianus (Congridae), Sardinella brasiliensis (Clupeidae), Merluccius hubbsi (Merlucciidae), Urophycis brasileinsis (Gadidae), Genypterus brasiliensis and Raneya brasiliensis (Ophidiidae), Porichthys porosissimus and Thalassophryne montevidensis (Batrachoididae), Parona signata and Trachinotus marginatus (Carangidae), Sphyraena tome (Sphyraenidae), Etropus longimanus, Paralichthys orbignyanus, P. patagonicus and Xystreurys rasile (Paralichthyidae), Thyrsitops lepidopoides (Gempylidae), Pseudopercis semifasciata (Pinguipedidae), Percophis brasiliensis (Percophidae), Astroscopus sexspinosus (Uranoscopidae), Dules auriga (Serranidae), Boridia grossidens (Haemulidae), Prionotus nudigula (Triglidae), Cynoscion guatucupa and Umbrina canosai (Sciaenidae). Among the 28 species of Chondrichthyes, 11 species (39.3%) were captured in the present study, namely: Mustelus schmitti (Triakidae), Squatina guggenheim (Squatinidae), Atlantoraja castelnaui, A. cyclophora, A. platana, Psammobatis extenta, P. lentiginosa, Rioraja agassizii and Sympterygia bonapartii (Rajidae), Pseudobatos horkelii (Rhinobatidae) and Dasyatis hypostigma (Dasyatidae). Due to the distribution (Southeast region of Brazil to Argentina), we also consider the following species endemic to the zoogeographic province of Argentina: Tetronarce puelcha (Torpedinidae), Anchoa marinii and Engraulis anchoita (Engraulidae) and Brevoortia pectinata (Clupeidae). Thus, 35.29% of Chondrichthyes and 11.85% of Actinopterygii (15.1% of the total) belong to the endemic fauna of zoogeographic province of Argentina.

The importance of the zoogeographic province of Argentina in the diversity of marine and estuarine fish fauna of Southern Brazil was evidenced in two other studies carried out in the Laje de Santos Marine State Park (Luiz Jr et al. 2008) and in the Paranaguá estuarine complex (Passos et al. 2012). The similarity in species composition was low (<50%) due to the different types of environments (rocky reefs, continental shelf and estuary). Totally, 427 species were observed, out of which 46 species (10.77%) are endemic to zoogeographic province of Argentina, being the fourth geographic distribution with the highest number of species, only behind the Western Atlantic with 169 species (39.58%), circumtropical with 52 species (12.18%) and trans-Atlantic with 50 species (11.71%). There was higher rate of endemism in the zoogeographic province of Argentina among marine demersal species (14.29%), followed by estuarine (10.95%) and reef (4.69%) species.

Based on the results of this work, we highlight the need for studies addressing other aspects of exploited communities, such as diversity analysis, spatial and seasonal distribution, as well as on impacts on explored demersal fishes, such as types of fishing, pollution, degradation of coastal areas and the introduction of exotic species.

Acknowledgments

The authors thank all the fishermen from the “Pró-Pesca Project: fishing the knowledge” and the Zoological Collection team at Santa Cecília University.

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Publication Dates

• Publication in this collection
  21 Jan 2019
• Date of issue
  2019

History

• Received
  22 Aug 2018
• Reviewed
  23 Nov 2018
• Accepted
  12 Dec 2018