When Does This Fish Spawn? Fishermen's Local Knowledge Of

Environ Biol Fish (2006) 76:371–386 DOI 10.1007/s10641-006-9043-2

ORIGINAL PAPER

When does this ﬁsh spawn? Fishermen’s local knowledge of migration and reproduction of Brazilian coastal ﬁshes

R. A. M. Silvano Æ P. F. L. MacCord Æ R. V. Lima Æ A. Begossi

Received: 4 November 2005 / Accepted: 21 March 2006 / Published online: 5 June 2006 Ó Springer Science+Business Media B.V. 2006

Abstract Fishermen’s local knowledge of fish- duction according to fishermen knowledge: fish ing resources may be an important source of species spawn in open ocean, near reefs or in information to improve artisanal tropical fish- coastal rivers (estuaries); some fishes reproduce eries management, such as those found in Bra- during the summer and others in winter, while zil, where most data on fish biology is lacking. some have more defined spawning months. The We aim to study the local ecological knowledge main fish migratory patterns mentioned by in- that Brazilian coastal fishers have about repro- terviewees were: long migrations along the ductive aspects (season, places and migration) coast, usually in the South to North direction, of 13 coastal fish species of commercial impor- short migrations among reefs, fishes that do not tance. We selected fishermen with more than migrate, migrations between the shore and open 30 years of fishing practice and we interviewed ocean and migrations between the sea and a total of 67 fishermen: 29 from the southeast- coastal rivers. Fishermen’s knowledge differed ern coast, from the communities of Puruba, among fish species: most fishermen did not Almada, Picinguaba and Bertioga, and 38 from know spawning places or seasons of large pe- the northeastern coast, from the communities of lagic fishes, which raised concerns of their pos- Valença, Arembepe and Porto Sauı´pe. In the sible depletion. We compared such interviews, we used standardized questionnaires ethnoichthyological information with available and showed photos of fish species. Our results scientific data, indicating promising insights indicate some general patterns in fishes’ repro- about reproduction and migration of Brazilian coastal fishes. Data gathered from local fishermen may provide inexpensive and prompt R. A. M. Silvano (&) information, potentially applicable to fisheries Depto. Ecologia, UFRGS, CP 15007, 91501-970 Porto management. Our approach might be useful to Alegre, RS, Brazil several other small-scale fisheries, especially the e-mail: [email protected] tropical ones, where there is a high diversity of P. F. L. MacCord Æ R. V. Lima Æ A. Begossi target species and a low biological and ecolog- Museu de Histo´ ria Natural, I.B. UNICAMP, ical knowledge about these species. C.P. 6109, 13084-971 Campinas, SP, Brazil Keywords Fishery management Æ Marine fishes Æ P. F. L. MacCord Programa de Po´ s Graduaçaõ em Ecologia/ IB Ethnoichthyology Æ Fish reproduction Æ Fish UNICAMP, Campinas, SP, Brazil migration

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Introduction context of complex, poorly known tropical fisheries (Johannes et al. 2000; Mathooko 2005). On Human communities that depend upon natural the other hand, the researcher should not to take resources show often a detailed knowledge of the fishermen’s LEK without checking (Huntington biology and ecology of plants and animals (Gadgil 2000). Ethnobiological surveys bring the meth- et al. 1993; Berkes 1999; Diamond 2005). Ethno- odological and conceptual tools to ‘‘translate’’ biology is the scientific discipline dedicated to the fishermen’s knowledge, providing useful, and of- study of such local ecological knowledge (Berlin ten new, information and guidelines to the related 1992; Berkes 1999). Albeit one of the first surveys disciplines of fish biology and fisheries, and aiding analyzing local ecological knowledge of Carib- to devise a multidisciplinary approach to marine bean fishers about marine fishes was published conservation (Drew 2005). about 40 years ago (Morril 1967), it was only Notwithstanding its high relevance to fisheries recently that the potential contribution of this management, reproductive patterns of tropical kind of survey to advance fisheries science has marine fishes are still poorly known, mainly due been widely recognized (Huntington 2000; to the difficulties involved in studying marine fish Johannes et al. 2000; Haggan et al. 2003; Drew reproduction (Sadovy 1996). Fishermen usually 2005; Mathooko 2005). Indeed, studies recording exploit fish schools (Parrish 1999) and fish and analyzing fishers’ LEK have been useful to spawning aggregations (Coleman et al. 1996), and better understand local fishing practices and cus- fishermen’s knowledge about spawning patterns tomary or common management rules (Johannes and migratory behavior of marine fishes has ad- 1978, 2002; Berkes 1999), to gather new biological vanced biological research (Johannes 1981; information about fish ecological aspects, such as Johannes et al. 2000). Nevertheless, fishermen’s migration, feeding habits and reproduction knowledge usually deals with local processes (Johannes 1981, 1988; Johannes et al. 2000; Val- (Degnbol 2005), which may reduce its efficacy to bo-Jorgensen and Poulsen 2001; Silvano and reveal general biological patterns of fishes, espe- Begossi 2002, 2005), to improve impact assess- cially for those with a wide geographical distri- ments (Johannes 1993) and to aid in marine bution. However, such general patterns can be conservation (Roberts et al. 2003; Drew 2005), addressed by a comparative approach, which among other applications. Furthermore, fishers’ integrates results of standardized surveys con- LEK may compliment conventional biological ducted in two or more fishing communities, surveys, by improving research design (Poizat and located along the distributional range of a fish Baran 1997), by providing more refined and lo- species or a fish community (Valbo-Jorgensen cally based data (Aswani and Hamilton 2004; and Poulsen 2001; Silvano and Begossi 2005). Huntington et al. 2004), and by revealing past Brazilian artisanal coastal fishermen are low- abundance trends in the population of exploited income people and usually they show a weak fishes, from before conventional fish landing data political organization (Diegues 1999), often being started to be recorded (Pauly 1995; Saénz-Arroyo neglected in official management measures et al. 2005). Fishermen’s knowledge of fishing (Begossi and Brown 2003). Although these fish- resources is especially important to support fishermen have developed a detailed knowledge of eries management in developing tropical coun- the classification and the ecology of marine and tries, due to the lack of research and biological estuarine fishes (Cordell 1974; Marques 1991; data on exploited fishing resources in a local or Begossi and Figueiredo 1995; Paz and Begossi regional scale (Johannes 1998; Johannes et al. 1996; Silvano and Begossi 2005), most of the 2000; Drew 2005; Mathooko 2005). Unfortu- published surveys do not address in detail po- nately, fishermen’s LEK has been often over- tential applications of local knowledge in fisheries looked or dismissed by biologists (Johannes 1993; management. Coastal fisheries management in Huntington 2000). Such resistance to consider Brazil is restricted to few and relatively inflexible fishermen’s LEK and points of view might be broad rules, due in part to a lack of data on the harmful to fisheries management, especially in the biology and ecology of target fishes. Indeed, most

123 Environ Biol Fish (2006) 76:371–386 373 of the available data about reproduction of Bra- Materials and methods zilian marine fishes are in scattered reports, dis- sertations and surveys (some with data collected Study sites as much as 40 years ago), usually published in local or regional journals with narrow circulation We studied seven artisanal fishing communities, (Almeida 1965; Sadowski and Dias 1986). four in the North of Saõ Paulo State, southeastern The main goal of this paper is to analyze local Brazilian coast, and three in the North of ecological knowledge on reproduction and Bahia State, northeastern Brazilian coast (Fig. 1, migration of 13 marine and estuarine fishes, Table 1). These study sites and fishing commu- through interviews with experienced fishermen nities were selected because of the lack of de- in seven artisanal fishing communities along the tailed ethnoichthyological surveys in these areas, southeastern and northeastern Brazilian coasts. compared to other coastal areas in southeastern We also compare fishermen’s knowledge with and northeastern Brazil (Marques 1991; Begossi available biological data from the ichthyological and Figueiredo 1995). We also selected the stud- literature, to propose insights that could improve ied regions according to logistic and opportunistic our current biological and ecological knowledge advantages: these sites have been the focus of and contribute to fisheries management. The other research projects and the fishing commu- results of this study may also be applicable to nities of Arembepe and Valença were already similar sites where these fishes also occur, espe- addressed in previous surveys (Kottak 1967; cially when broad patterns in migration and Cordell 1974). reproduction of the studied coastal fishes are On the northeastern coast of Bahia, the arti- observed. sanal fishermen exploit lobster, shrimp and fishes

Fig. 1 Map of Brazilian coast showing the studied sites (a) migration between the shore and open ocean, (C) long Arembepe, (b) Valença, (c) Ubatuba region, where are migration from South to North direction, (D) migration located the fishing communities of Almada, Puruba and between the sea and coastal rivers and (E) short migration Picinguaba and (d) Bertioga. Main migratory patterns around nearby reefs. Source of maps: E.E. De Miranda & mentioned by fishermen (Table 4) are illustrated: (A) A.C. Coutinho. Brasil Visto do Espaço. Campinas: migration along the coast (not long distances), (B) Embrapa Monitoramento por Sate´lite, 2004

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Table 1 Fish species addressed in this survey, with the most mentioned common names and total number of interviewed fishermen, for each fish species and fishing community Fishes Fishing communities, Bahia coast Fishing communities, Saõ Paulo coast Total of interviewees Arembepe Valença Porto Sauı´pe Almada Picinguaba Puruba Bertioga BA SP All

Caranx spp.a Guaricema Xumberga Guaricema Olhudo Olhudo Carapau Xareú382967 Centropomus parallelus Robalo Robalo Robalo Robalo Robalo Robalo peva Robalo 38 29 67 Cynoscion jamaicensis Pescada Pescada 23 23 Epinephelus marginatus Garoupa Garoupa d Garoupa Garoupa Garoupa Garoupa 23 29 52 Euthynnus alleteratus Bonito Bonito Serrinhae Bonito 29 29 Lutjanus synagris Arioco´ Cioba Arioco´ 38 38 Micropogonias furnieri Corvina Corvina Corvina Corvina 29 29 Mugil platanus Tainha Tainha Tainha Tainha Tainha Tainha 23 29 52 Mycteroperca acutirostris Badejo Badejo Badejo Badejo Badejo Badejo mira Miracelo 38 29 67 Pomatomus saltatrix Enchova Enchova Enchova Enchova Enchova Enchova 23 29 52 Scomberomorus brasiliensis Cavalab Cavala Cavala 38 38 Seriola lalandi Enchovac Olhete Enchova 38 38 Trichiurus lepturus Espada Espada Espada Espada 29 29 Number of interviewees 13 10 15 10 7 4 8 38 29 67 When more than one name to a fish species was frequently cited in a given fishing community, alternative names are mentioned in footnotes. BA = Bahia, northeastern Brazilian coast, SP = Saõ Paulo coast, southeastern Brazilian coast a

Two fish species from the same genus were used in the survey: Caranx latus in Saõ Paulo fishing communities and Caranx crysos in Bahia fishing communities. We 76:371–386 (2006) Fish Biol Environ grouped the two for analysis, as they belong to same genus and are similar in appearance bAlso called ‘‘Sororoca’’ cAlso called ‘‘Olho de Boi’’ dAlso called ‘‘Badejo’’ eAlso called ‘‘Bonito serrinha’’ Environ Biol Fish (2006) 76:371–386 375 in reefs, open sea and estuaries, usually with small about fish reproduction and migration (Silvano boats and using several types of fishing gear, some and Begossi 2002). Indeed, local knowledge may of which are typical, such as the ‘calaõ’, a kind of be unevenly distributed among fishermen and seine net used to catch shrimp and small fishes selecting appropriate informants is an important (Cordell 1974). There are also large-scale fisheries step in gathering useful information (Johannes equipped with large boats and a crew of several et al. 2000). fishermen (up to 15), which go fishing over large We interviewed fishermen using standardized distances for several days at sea. In Bahia, such questionnaires with the questions: Where and fishing is usually done with hook and line, tar- when does this fish spawn? Does this fish migrate geting pelagic fishes (mainly Scombridae and (‘run’)? When, and to where (migratory routes)? Carangidae) and reef fishes (mainly Lutjanidae We showed fish species to fishermen as color and Serranidae), which are also exploited by photographs, in the same order for each inter- artisanal small-scale fishermen (Ivo and Hanson viewee. Details of this interviewing methodology 1982). Artisanal fishermen often work as a crew are in other surveys (Silvano and Begossi 2002, on large fishing vessels. 2005). We selected 13 coastal fish species for this On the southeastern Brazilian coast, artisanal survey (Table 1), in order to include representa- fishermen are the caiçaras, inhabitants of areas tives of the most important fish groups often with Atlantic Forest remnants, who rely mostly caught by these artisanal fishermen in several on coastal fisheries as a source of food and cash aquatic habitats: reef fishes from the Lutjanidae and who descend from indigenous Brazilians and (Lutjanus synagris) and Serranidae (Epinephelus Portuguese colonizers (Begossi 1998; Diegues marginatus, Mycteroperca acutirostris), estuarine 1999). Fish is the main protein source for caiçara fishes from the Mugilidae (Mugil platanus) and fishermen, who use mainly gillnet and hook and Centropomidae (Centropomus parallelus), line to catch fishes (mainly Pomatomidae, Serra- demersal fishes from Sciaenidae (Cynoscion nidae, Mugilidae, Sciaenidae, Centropomidae and jamaicensis, Micropogonias furnieri) and pelagic Carangidae), shrimps and squids (Begossi 1996). fishes from the Carangidae (Caranx crysos, Caiçara fishing communities have been facing C. latus, Seriola lalandi), Pomatomidae (Po- difficulties and conflicts, such as an increase in matomus saltatrix), Trichiuridae (Trichiurus tourism and competition with large-scale com- lepturus) and Scombridae (Euthynnus alleteratus, mercial fishing vessels. Brazilian governmental Scomberomorus brasiliensis). We made the sur- environmental agencies, besides being ineffective veys during July 2003 (Arembepe and Valença, in dealing with these problems, have been con- Bahia), August 2004 (all Saõ Paulo fishing com- straining the caiçaras’ subsistence and small-scale munities) and January 2005 (Porto Sauı´pe, Bahi- fishing activities (Begossi 1995; Begossi et al. a). The number of interviewed fishermen varied 2001). for each fish species, as some fish species were addressed in some fishing communities but not in Data collecting and analysis others (Table 1), due to differences in abundance and commercial importance of fishes among In each studied fishing community, we inter- studied fishing communities. We also reduced the viewed selected fishermen according to the fol- number of fish species in some communities in lowing criteria: those who fish as their main order to optimize our sampling effort and time economic activity (active or retired), and who spent during interviews. However, we included have at least 30 years of fishing practice in the some fish species in all surveys (Table 1). We studied region. We thus analyzed a restricted interviewed fishermen about Caranx latus in the sample of more experienced fishermen, whom we Southeast and about C. crysos in the Northeast. found based on data from previous surveys and This was due to possible differences in abundance interviews. We used such an approach because and commercial importance of these fishes be- previous ethnoichthyological surveys show that tween the two regions, and because we intended older fishermen are usually more knowledgeable to compare our results with previous surveys

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Table 2 Spatial and temporal patterns of spawning of coastal ﬁshes, numbers are % of interviewees who mentioned each data Time and place of spawningb Fish speciesa Cspp Cpar Cjam Emar Eall Lsyn Mfur Mpla Macu Psal Sbra Slal Tlep

Don’ t know where 48 21 65 40 55 39 24 15 39 52 42 66 41 Reefs 10 3 31 18 2 37 6 3 3 Reef’s crevices 1 8 14 5 10 Rivers (estuary) 43* 38 2 3 Rivers’ mouth 4 3 6 2 Submerged logs in the river 7 8 1 Open sea, offshore 15 1 28 * 38** 10** 1 8* 3 3 41* Mud bottoms 3 8 3 3 Patos’ Lagoon (South Brazil)c 1 3 19 2 Brazilian southern coast 3 3 23 15 Inshore, coastal waters 1 3 2 3 14 7 Offshore rocky outcrops (‘‘parce´is’’) 4 3 7 3 4 Coastal sandy beaches 3 14 2 2 3 Islands 3 2 7 3** 4 10 Do not know when 6 28 65 54 41 39 10 13 54 54 34 63 31 Summer 9** 1 9 12* 34** 3** 21 6 7 2** 2** 5* 14* Winter 4 6* 4 4 13 10** 8** 6* 8 8 5 10 Spring ** ** * ** * Autumn * Year round 3 3* 4 4 7 14 4 7 3 24 Total number of interviewees 67 67 23 52 29 38 29 52 67 52 38 38 29

Information gathered from scientific literature are marked ** for the same species in the study regions and * for other related species or the same species in other 76:371–386 (2006) Fish Biol Environ regions from Brazil or elsewhere aCodes for fish species (Table 1): Cspp (Caranx spp.), Cpar (C. parallelus), Cjam,( C. jamaicensis), Emar (E. marginatus), Eall (E. alleteratus), Lsyn (L. synagris), Mfur (M. furnieri), Mpla (M. platanus), Macu (M. acutirostris), Psal (P. saltatrix), Sbra (S. brasiliensis), Slal (S. lalandi), Tlep (T. lepturus). bLiterature sources to reproduction and migration of studied fishes: Almeida (1965), Ivo and Hanson (1982), Sadowski and Dias (1986), Fonteles-Filho and Ferreira (1987), Vazzoler (1991), Vieira and Scalabrin (1991), Ibagy and Sinque (1995), Haimovici and Krug (1992, 1996), Juanes et al. (1996), Muelbert and Sinque (1996), Martins and Haimovici (1997), Gillanders et al. (1999), Aliaume et al. (2000), Romagosa et al. (2000), Andrade et al. (2003), Barreiros et al. (2004), Teixeira et al. (2004) cThis spawning ground was only mentioned by southeastern fishermen Environ Biol Fish (2006) 76:371–386 377 made in the Southeast. We aggregated the an- and spawning sites, due to the low number of swers about these two fish species for analysis, citations (Table 2). We also used chi-square tests considering their overall similarity and taxonomic to compare frequency of doubts (number of relatedness (same genus) (Table 1). Furthermore, fishermen who answered ‘I do not know ‘) relative answers given by northeastern and southeastern to the total number of interviewed fishermen for fishermen about these two fish species did not each fish species. Such comparison may indicate differ regarding their main characteristics, such as differences in fishermen’s knowledge of distinct whether they migrate or not (v2 = 1.56, d.f. = 1, fish species (Silvano and Begossi 2002). P = 0.26), main patterns (v2 = 7.87, d.f. = 3, We show data as a percentage of all inter- P = 0.05) and seasons (v2 = 5.11, d.f. = 3, viewed fishermen for each fish species, in order to P = 0.16) of migration, or main sites (v2 = 5.85, standardize distinct sample sizes among fish spe- d.f. = 2, P = 0.05) and seasons (v2 = 5.65, cies (Table 1). The sum of percentages shown in d.f. = 3, P = 0.13) of spawning. rows of Tables 2–5 may sometimes be more than Whenever sample sizes (number of fishermen’s 100%, as some interviewees mentioned more citations) allowed, we compared fishermen’s an- than one aspect for the same fish species (Ta- swers about some general migratory and repro- ble 2). Among all data obtained, we show the ductive characteristics of fish species through chi- data mentioned by 10 fishermen or more (the square tests, using the number of interviewees’ most cited). To properly quantify, tabulate and citations (Fowler and Cohen 1990). Although compare the answer of each interviewed fisher- sometimes the same fisherman mentioned more man, we selected information of broader scope, than one feature for the same fish species (e.g. referring to general spatial or temporal features, spawning both in winter and summer), we con- counting how many fishermen mentioned that sidered samples as independent, as we believe information. For example, if an interviewee said that one answer would not be influencing the ‘‘this fish spawns in the open ocean, during the other, even when both answers were given by the hot season, mainly during January’’, we marked same person. Three fish species (C. jamaicensis, S. one fisherman referring to ‘‘open ocean’’ (habi- lalandi and S. brasiliensis) were excluded from the tat), ‘‘hot season’’ (spawning period), and ‘‘Jan- chi-square analyses comparing migratory patterns uary’’ (spawning month). Therefore, the sum of

Table 3 Spawning calendar of coastal ﬁshes according to ﬁshermen, numbers are % of interviewees who mentioned each data Fishes Autumn Winter Spring Summer All seasons Mar Apr May Jun Jul Aug Sep Oct Nov Dec Jan Feb Total number of interviewees

Caranx spp. 4**13 316741797**67 C. parallelus 10 4 3* 10* 7* 15* 7* 12* 10 16 13 10 67 C. jamaicensis 4 9 13 9 4 23 E. marginatus 4 10 10 4 6 10 12 12* 8 4 52 E. alleteratus 3 3 3 3 7 3 10 17** 21** 14 10 29 L. synagris 3 3 5 5 5 8 5 16 8 2** 3** 3 38 M. furnieri 3 14 21** 21** 28** 28** 10** 28** 28 21 10 29 M. platanus 2 2 15 44 42 29** 12** 10 4 4 4 2 52 M. acutirostris 4* * 1* 3* 3* 7* 12* 13 12 9 6 4 67 P. saltatrix * 4 6 6 4 12 13 15 13* 6* 4** 4** 52 S. brasiliensis 13 5 3 3 3 3 8 11 8** 11** 11 16 38 S. lalandi 3 3 11 3 11 5 11 38 T. lepturus 10 7 10 14* 21* 14* 14* 3* 3* 29 Information gathered from scientiﬁc literature are marked ** for the same species in the study regions and * for other related species or the same species in other regions from Brazil or elsewhere. Literature sources of scientiﬁc data are in Table 2

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Table 4 Spatial and temporal migratory patterns of coastal ﬁshes, according to ﬁshermen, numbers are % of interviewees who mentioned each data Migratory routes and seasons Fish species Cspp Cpar Cjam Emar Eall Lsyn Mfur Mpla Macu Psal Sbra Slal Tlep

Running fish (migrates) 96* 70 61 60* 100 50* 66** 100** 45* 83** 95 74* 86 Do not migrate (sedentary) 4 22 13 31* 32 34 49 8 3 11 14 Around reefs 1 25* 13 19 2 3 8 River (estuary)—sea 1 43 9 * 38** 2 ‘‘Arribaçaõ’’a 196169 3 Offshore-coast 9* 1 4 17 * 48* 12 1 6 3 * 14 North–South 4 3 2 3 10 1* 3 3 3 South–North 3 4 10 48 13 3 62** 6 44** 11 13 34 Deep waters–shallow waters 13 13 8 5* 2 18 11* Searching for food 16 6 4 2 10 5 7* 8 11 5 17 Everywhere along the coast 9 4 10 3 10* 2 1 2 18 11 10* Do not know when 3 6 26 10 3 12 15 3 22 3 Summer 33 9 9 10 45 17 8 7 4 26 13 3* Winter 4 16 8 3 22 10** 17** 6 15** 5 11 14 Autumn ** ** Spring ** * Year round 22 7 4 10 7 13 17 2 9 18 18 3 Along the tides 1 7 9 4 2 3

During rains and storms 7 4 3 2 3 76:371–386 (2006) Fish Biol Environ Total number of interviewees 67 67 23 52 29 38 29 52 67 52 38 38 29 Information gathered from scientific literature are marked ** for the same species in the study regions and * for other related species or the same species in other regions from Brazil or elsewhere. Literature sources of scientific data and codes for fish species are in Table 2 aTypical seasonal migration, related to the water temperature and to the fish feeding behavior. Mentioned only by northeastern fishermen Environ Biol Fish (2006) 76:371–386 379

Table 5 Migration calendar of coastal ﬁshes, according to ﬁshermen, numbers are % of interviewees who mentioned each data Fishes Autumn Winter Spring Summer All seasons Mar Apr May Jun Jul Aug Sep Oct Nov Dec Jan Feb Total number of interviewees

Caranx spp. 166331 491316181367 C. parallelus 44312979610129767 C. jamaicensis 423 E. marginatus 2 4 2 2 10 10 4 4 2 52 E. alleteratus 17 10 10 3 7 7 10 14 31 31 21 29 L. synagris * 3*5*13*11*8*5*3*3*** *38 M. furnieri 3 ** 7 ** 14 ** 14 ** 14 ** 14 ** 14 17 7 10 7 3 29 M. platanus 19 40 ** 44 ** 33 ** 8 2 2 2 2 52 M. acutirostris 34331131464367 P. saltatrix 4 4 12 15 ** 17 ** 8 ** 13 ** 12 ** 15 ** 6 4 4 52 S. brasiliensis 1133333338881138 S. lalandi 5353 335838 T. lepturus 7333310102124*17*7*729 Most mentioned months for each fish species are highlighted. Information gathered from scientific literature are marked ** for the same species in the study regions and * for other related species or the same species in other regions from Brazil or elsewhere. Literature sources of scientific data are in Table 2 all fishermen that mentioned such general fea- C. parallelus, M. platanus and demersal M. fur- tures was used to calculate the percentages shown nieri (Table 2). By contrast, most of the inter- in Tables 2–5, according to procedures from other viewed fishermen did not know spawning places ethnoichthyological surveys (Silvano and Begossi or season of large pelagic fishes, such as E. al- 2002, 2005). leteratus, P. saltatrix and S. lalandi, as well as the demersal fish C. jamaicensis (Table 2). The number of fishermen’s citations for the Results three most mentioned fish spawning sites differed among fish species (v2 = 223, d.f. = 18, P < 0.01). We interviewed a total of 67 fishermen (Table 1) Thus, fishermen’s answers distinguish fishes that from 34- to 83-year-old (average = 63 years), who spawn mainly in reefs (E. marginatus, M. acuti- have been fishing in their respective regions for rostris and L. synagris), in coastal rivers or estu- an average period of 45 years. Notwithstanding aries (M. platanus and C. parallelus) and in open the potential differences in culture and knowl- sea (Caranx spp., E. alleteratus, M. furnieri and edge among studied fishing communities, we T. lepturus) (Table 2). Interviewees mentioned grouped data for analysis, in order to verify broad both seasons (Table 2) and specific months (Ta- patterns. Nevertheless, we also briefly addressed ble 3) of fish spawning. When they mentioned a some of the more remarkable differences be- season, we asked them to specify months, but they tween the answers given by fishermen from Saõ were not always able to do so. The calendar of fish Paulo and Bahia States. reproduction based on fishermen’s knowledge Comparison of percentages of fishermen’s indicates that most of the pelagic and reef fishes doubts (answers ‘I do not know where and when’, spawn during the summer and spring months Table 2) showed that spawning sites (v2 = 547, (Table 3). The estuarine M. platanus differed d.f. = 12, P < 0.001) and spawning seasons from all other fishes, spawning mainly during the (v2 = 537, d.f. = 12, P < 0.001) of some fish spe- winter months, C. parallelus spawns during both cies are better known to fishermen. Comparing winter and summer, and E. marginatus spawns in expected and observed values, fishermen showed late spring and early summer, but also during the fewer doubts about reproduction of estuarine winter. This fish showed a distinct pattern from

123 380 Environ Biol Fish (2006) 76:371–386 other reef fishes (L. synagris and M. acutirostris), tions differed among fish species regarding which spawn mainly during the spring (Table 3). migratory seasons (Table 4) (v2 = 65, d.f. = 24, For eight of the 13 studied fish species, at least P < 0.01): pelagic fishes, such as Caranx spp., one of the spawning months most mentioned by E. alleteratus and S. brasiliensis migrate during fishermen coincided with spawning months for the summer, while M. platanus, P. saltatrix and the same (or related) fish species gathered from C. parallelus migrate during the winter and the biological literature (Table 3). other fishes migrate year round (Tables 4 and 5). Contrarily to what we observed for fish repro- Fishermen’s answers suggest that some migration duction, few fishermen had doubts about fish patterns, such as long South to North migrations migration (Table 4). Despite all fish species being of M. platanus and P. saltatrix, as well as the regarded as migratory by at least some of the offshore and inshore migrations of M. furnieri interviewed fishermen (Table 4), answers (Table 4) may be related to reproduction, as they regarding whether a fish migrates (‘running’ fish) coincide with the mentioned spawning sites or not (sedentary) differed among the studied (Table 2) and months (Table 3) for these fishes. fishes (v2 = 104, d.f. = 12, P < 0.01). Thus, the Conversely, offshore and inshore migrations of majority of fishermen considered nine fish species some pelagic fishes, such as Caranx spp. and as being mostly migratory (Table 4), while four T. lepturus, may be related to feeding (Table 4). fishes, L. synagris (v2 = 1.61, d.f. = 1, P = 0.28), As observed for fish reproduction, fishermen’s M. furnieri (v2 = 2.2, d.f. = 1, P = 0.14) and data about migration of E. marginatus, Caranx M. acutirostris (v2 = 0.06, d.f. = 1, P = 0.8) were spp., M. furnieri, M. platanus and P. saltatrix equally quoted as sedentary or migratory. agree with proposed migratory patterns in scien- Although the reef fish E. marginatus was consid- tific literature (Table 4). Literature data about ered mainly migratory (v2 = 4.17, d.f. = 1, migratory periods, which were only available for P < 0.05), several fishermen (31%) also men- five of the studied fishes, usually agree with fish- tioned it as being sedentary (Table 4). ermen’s data (Tables 4 and 5). The interviewees mentioned four main patterns of fish migrations (Fig. 1), which differed for the studied fish species (v2 = 301.8, d.f. = 27, Discussion P < 0.01), thus making it possible to distinguish fishes that make long migrations along the coast, in Notwithstanding the extension of Brazilian coast, the South to North direction (M. platanus, P. sal- which harbors several fishing communities (Die- tatrix, Caranx spp., T. lepturus and E. alleteratus), gues 1999), we believe that our results may apply fishes migrating between the shore and open ocean to southeastern and northeastern Brazil, as long (or between deep and shallow waters) (M. furni- as our sample includes several fishing communi- eri), fishes migrating between the sea and coastal ties, which exploit various habitats (reefs, coastal rivers, according to prevailing tides and climatic waters, beaches and estuaries). However, perhaps conditions (C. parallelus), and reef fishes making our results would have more limited application short movements around nearby reefs (E. mar- to other Brazilian coastal regions with peculiar ginatus, L. synagris and M. acutirostris) (Table 4, environmental conditions and fishing communi- Fig. 1). Some fish species showed two migration ties, such as large estuarine lagoons in the far patterns according to the fishermen, such as M. South and the Amazon River mouth in the far platanus and Caranx spp. Besides both of these North. Although we showed the same fish species’ fishes migrating from the South to the North, the photographs to all interviewed fishermen, it might former also migrates between estuaries and the be that some fishermen referred to fishes other sea, and the latter also migrates between the shore than those shown to them. For example, north- and open-ocean (Table 4). eastern fishermen might be talking about Mycte- The calendar for fish migration according to roperca bonaci and Mugil liza, which are more interviewees (Table 5) generally agrees with the common there (Teixeira et al. 2004), instead of spawning calendar (Table 3). Fishermen’s cita- the congeners M. acutirostris and M. platanus (the

123 Environ Biol Fish (2006) 76:371–386 381 species shown in photographs), which occur the main issue in the studied fishing communities, mainly in the southeast. Some differences to the extent that fishermen did know reproduc- between answers of northeastern and southeast- tion habits of estuarine fishes. Moreover, inter- ern fishermen could, thus, be due to this distinct viewed fishermen did show good knowledge of perception. For example, most of the answers fish migration (Tables 4 and 5) as expected, con- regarding M. platanus reproduction came from sidering the importance of such knowledge to find the latter fishermen, who mentioned that this fish and exploit marine fish schools (Parrish 1999). spawns in the Brazilian South, more precisely in Another (and more plausible) explanation to the Patos Lagoon (Table 2). Conversely, northeast- observed lack of fishermen’s knowledge on ern fishermen might be dealing with another reproduction of some fishes might be that such similar Mugil species, which spawns elsewhere. events are relatively rare and difficult to observe, Only northeastern fishermen mentioned the ‘arr- contrarily to other biological aspects, such as fish ibaçaõ’ made by L. synagris and M. acutirostris. habitat and diet (Silvano and Begossi 2002). In This sort of seasonal migration, related to the such cases, reproduction of large pelagic fishes water temperature and to feeding, has been re- may be rare to fishermen due to two main factors, ported for M. bonaci (Teixeira et al. 2004) and L. not mutually exclusive. First, spawning occurs on purpureus (Ivo and Hanson 1982; Fonteles-Filho short and limited spatial and temporal scales. and Ferreira 1987) on the northeastern Brazilian Second, these fish species themselves are now coast, but it may not occur in the southeast. scarce and fishermen have less contact with them. Although dealing with supposedly more Indeed, some interviewed fishermen claimed that knowledgeable and experienced fishermen, we they were not able to mention reproductive pat- observed an overall lack of knowledge about fish terns because they rarely (or never) catch these reproductive patterns, especially for some large fishes with eggs. This situation deserves to be pelagic fishes with high market value. Conversely, further investigated through biological research, Pacific (Johannes 1981; Johannes et al. 2000) and as it raises concerns of over-fishing and depletion Caribbean (Coleman et al. 1996) fishermen are of these large pelagic fishes. A common pattern of very knowledgeable about fish spawning patterns. over-fishing in artisanal fisheries is increased cat- According to Diamond (2005), older people in ches of less valued, medium-sized or small her- traditional or illiterate societies are the equiva- bivorous and detritivorous fishes, associated with lents of our libraries, accumulating and transmit- a decline in catches of large, high value, piscivo- ting environmental knowledge. In this sense, one rous fishes (de Boer et al. 2001). Our results, thus, possible explanation for our results would be that raise some relevant questions: ‘Does the observed studied Brazilian fishermen might be gradually lack of fishermen’s knowledge of reproduction of loosing their local ecological knowledge, to the large reef and pelagic fishes indicate decreases in extent that they are in closer contact with urban the abundance of these fishes?’ Are large pelagic centers and media-derived information, and fish- fishes being replaced in fisheries by the well- ing is being replaced by other economic activities, known estuarine and demersal fishes?’ such as tourism (Begossi et al. 2001). If this is the Fishermen’s local ecological knowledge indi- case, then measures could be taken in order to cates general patterns in reproduction and avoid complete disruption of coastal artisanal migration of studied Brazilian coastal fishes. fishing, a relevant economic activity, especially These patterns usually agree with available sci- for poor people. Albeit in risk of vanishing due to entific data: for example, researchers already modernization and acculturation pressures documented the South to North spawning migra- (Johannes 1978), traditional knowledge and tion of P. saltatrix and M. platanus, which migrate associated management systems held by South against prevailing oceanic currents that will carry Pacific fishing communities for centuries have eggs and larvae back to estuaries in the South been recovering more recently (Johannes 2002), (Sadowski and Dias 1986; Vieira and Scalabrin showing that disruption of fishers’ culture is not 1991; Haimovici and Krug 1996; Juanes et al. irreversible. However, knowledge loss may not be 1996). Indeed, seven of the interviewed fishermen

123 382 Environ Biol Fish (2006) 76:371–386

(10%), all from southeastern Brazil, mentioned show considerably inter-population variability that the former fish species migrates soon after regarding spawning sites, such as M. furnieri, (‘follows’) the latter. Thus, it would be worth which spawns in the sea (pelagic coastal waters) in checking this and the ecological factors influenc- Brazil (Vazzoler 1991; Ibagy and Sinque 1995) and ing such an alleged association between these two in estuaries in Uruguay (Vizziano et al. 2002). fishes: are P. saltatrix preying on M. platanus? Are Indeed, in our survey, fishermen’s answers also migratory movements of these two fish species indicated several spawning sites for this fish triggered by similar environmental cues, such as (Table 2). Sadowski and Dias (1986) made a differences in water temperature (Sadowski and broad mark-recapture survey of Mugil cephalus Dias 1986; Vieira and Scalabrin 1991)? Under- (possibly M. platanus) from 1954 to 1962 on the standing this would aid in managing and protect- Brazilian south and southeastern coasts, identify- ing spawning runs of two important target fishes. ing a South to North spawning run, but not finding Furthermore, some of these migratory patterns spawning sites. Authors then use pieces of evi- were also recorded in previous ethnoecological dence from other places (such as Mexico) to argue surveys in other fishing communities: marine that this fish could be spawning in open-ocean, fishermen from Bu´ zios Island (southeastern Bra- near the coast. However, this fish also makes short zil) and in Moreton Bay (eastern Australia) also lateral movements between estuaries and the sea mentioned a South to North migration of P. sal- during its spawning runs along the Brazilian tatrix, and its association with migration of mugi- southeastern coast (Sadowski and Dias 1986). lids (Silvano and Begossi 2005). We could not find Although eggs and larvae have not yet been ob- data on migration or reproduction of some fishes served in coastal rivers and estuaries, our data and our data may, thus, be the only information raise the hypothesis that perhaps some popula- currently available. Most of the literature referred tions of M. platanus could be spawning near or to in this survey (Tables 2–5) addresses other re- inside estuaries. In Northern Australia, aboriginal lated fish species, or the same fish species in other fishermen claim that the barramundi (Lates regions of Brazil and elsewhere. Thus, even ob- calcarifer, Centropomidae) spawns in rivers, con- served correspondence between fishermen and trarily to the opinion of ichthyologists, who affirm literature data does not necessarily mean redun- that this fish spawns in the coastal sea. However, dant information (Silvano and Begossi 2005). biological research in that region confirms that Fishermen’s knowledge recorded here some- aborigines are right: the barramundi population times disagrees with the scientific literature. Such shows a distinct spawning pattern there, differing discrepancy may provide new insights to be from populations already studied in other Aus- investigated by biological research, improving the tralian regions (Johannes 1988). existing knowledge base (Marques 1991; Aswani According to Vazzoler (1991), there are two and Hamilton 2004). According to fishermen, distinct populations of M. furnieri on the Brazil- overall migratory patterns of M. platanus match ian coast: one in the south and the other in the literature data. However, according to fishermen, southeast (encompassing our study region, this species spawns in rivers or estuaries and, Fig. 1). However, a recent study challenges this according to scientists, in the ocean (pelagic assumption: genetic similarities indicate that coastal waters) (Table 2). Such discrepancy might there could be just one population of M. furnieri arise from fishermen confounding spawning and in Brazil (Levy et al. 1998). Our results support nursery sites of this fish, to which estuaries provide the ‘two populations’ hypothesis, as most of the an important growing and nursery site (Vieira and interviewed fishermen said that this fish species Scalabrin 1991; Romagosa et al. 2000). However, does not migrate over long distances or in the considering the lack of scientific knowledge on South to North direction (Table 4). There would, reproductive behavior of this fish in our study re- thus, be few contacts between individuals from gions, fishermen’s answers reported here might be southern and southeastern Brazil. Therefore, worth checking, even if they sound unreliable to considering fishermen’s assertions, we may raise ichthyologists at first glance. Some fishes may the hypothesis that the observed gene flow

123 Environ Biol Fish (2006) 76:371–386 383 between southern and southeastern populations single, larger, and strategically placed marine re- of this fish could be related to passive transport of serves. Moreover, our results also show that eggs and larvae along the Brazilian coast, instead maintenance of exploited marine fish stocks may of migration and interbreeding of adults. Com- rely upon conservation measures from other bined evidence from our ethnoichthyological connected habitats and ecosystems, such as survey and literature thus suggest that more coastal rivers and estuaries. Most of the inter- detailed migratory surveys would be desirable to viewed fishermen mentioned the dependence of solve this important question. C. paralellus on estuaries and rivers for repro- Another discrepancy between local and scien- duction. Indeed, a biological survey also shows tific knowledge refers to spawning of reef fishes the importance of estuaries as nursery grounds for from the Serranidae family (E. marginatus and M. Centropomus undecimalis in Puerto Rico (Alia- acutirostris). According to interviewed fishermen, ume et al. 2000). Our results also aid in devising these fishes spawn near reefs and usually do not more flexible management rules: instead of clos- migrate, while according to the literature these ing all fishing activities during certain periods, fishes, especially Epinephelus spp., usually form managers might close fishing of some fish species large spawning aggregations and migrate over long (or fish groups) during their respective spawning distances to spawn away from their home reefs in seasons. the Caribbean (Coleman et al. 1996; Sadovy 1996). Interviewed fishermen in our survey did not mention such spawning aggregations, not even on the Conclusion northeastern coast, which is similar to the Carib- bean (tropical reefs, warm water). Furthermore, Data gathered from fishermen in this survey, such spawning aggregations have not yet been which agree with data acquired from other fish- documented in Brazilian coastal waters, albeit ermen elsewhere, and with literature data, might some researchers believe that they might occur indicate real biological patterns. We do not claim (Andrade et al. 2003). Our results suggest that that our ethnoichthyological data should replace serranid spawning aggregations may not occur in biological surveys, but our results could help to Brazil, or, if they occur, they are probably rare, elaborate such future surveys, to the extent that sporadic and usually not exploited by fishermen. we can devise testable hypotheses about fish The efficacy of marine reserves (and zoning migration and spawning, based on fishermen’s systems) to fishery management depends, among knowledge. Such an approach would save time other things, on whether reserves afford protec- and money, providing rapid results, readily tion of important habitats to critical and vulner- applicable to fisheries management, thus follow- able life-stages of exploited fishes, such as ing the ‘data-less’ approach proposed by Johan- spawning sites (Roberts et al. 2003). Our results nes (1998). Besides informing scientists and may thus be useful to set marine protected areas managers when devising management measures, and to devise ecologically sensitive zoning plans our data would help to achieve the necessary on the Brazilian coast, as we identify major involvement of the stakeholders (in our case, the spawning and migratory patterns of some Brazil- fishermen), in the design and implementation of ian coastal fishes, according to fishermen’s these measures, such as marine reserves (Roberts knowledge. For example, highly migratory fishes, et al. 2003). Applying fishers’ knowledge in such as M. platanus and P. saltatrix, might be management may not be an easy task, due to managed in a distinct fashion than less migratory differences between fishers’ and researchers’ or sedentary reef and demersal fishes, such as knowledge base and perceptions (Degnbol 2005). E. marginatus and M. furnieri. Following the The agreements observed here between fisher- reasoning of Roberts et al. (2003), the former men’s and scientists’ knowledge about fish would be best protected by several smaller mar- migration and reproduction may be a starting ine reserves along their entire migratory route, point to devise a ‘common ground’ knowledge, while the second may be best managed through which would be understandable to both

123 384 Environ Biol Fish (2006) 76:371–386 researchers and users, thus being applicable in Begossi A (1995) Fishing spots and sea tenure: incipient future co-management of fishing resources (De- forms of local management in Atlantic Forest coastal communities. Human Ecol 23:387–406 gnbol 2005). Our study shows that local fisher- Begossi A (1996) Fishing activities and strategies at Bu´ zios men’s knowledge has the potential to improve Island (Brasil). In: Meyer RM, Zhang C, Windsor ML, understanding of reproduction and migration of McCay BJ, Hushak LJ, Muth RM (eds) Proceedings of Brazilian coastal fishes, as already observed in the the World Fisheries Congress, Theme 2, Atenas, May 1992. Oxford & IBH Publishing, Calcutta, pp 125–141 tropical Pacific (Johannes 1981; Johannes et al. Begossi A (1998) Cultural and ecological resilience among 2000) and in the Mekong River (Valbo-Jorgensen caicaras of the Atlantic Forest coast and caboclos of and Poulsen 2001). Furthermore, the ethnobio- the Amazon. In: Berkes F, Folke C (eds) Linking logical approach and methods here reported social and ecological systems for resilience and sustainability. The Beijer International Institute of Eco- could also be useful to the study and management logical Economics, Stockholm, pp 129–157 of several other poorly known tropical fisheries, Begossi A, Figueiredo JL (1995) Ethnoichthyology of thus helping to improve dialogue and mutual southern coastal fishermen: cases from Bu´ zios Island understanding between fishers and researchers, and Sepetiba Bay (Brazil). Bull Mar Sci 56:710–717 Begossi A, Brown D (2003) Fisheries co-management which would ultimately benefit the development experiences in Latin America and the Caribbean. In: of sound marine conservation strategies (Drew Wilson DC, Nielsen JR, Degnbol P (eds) The fisheries 2005). co-management experience: accomplishments, challenges and prospects. Kluwer Academic Publishers Acknowledgements We thank to Brazilian fishermen of Fish and Fisheries Series 26, Dordrecht, pp 135–150 the studied communities for their kind cooperation, to Begossi A, Hanazaki N, Peroni N (2001) Knowledge and P. Zahorcsak for help in the fieldwork and data analyses; use of biodiversity in Brazilian hot spots. Environ, E. Camargo for help in the fieldwork, to A.S. Melo for Develop Sustain 2:177–193 advice in statistical analyses; to FAPESP for financial Berkes F (1999) Sacred ecology—traditional ecological support through research projects (01/05263-2, 01/14526-7 knowledge and resource management. Taylor & and 04/02309-1) and for grants to R.A.M. Silvano (01/ Francis, Philadelphia, 209 pp 07247-4) and to R.V. Lima (03/04154-0); to CNPq, for a Berlin B (1992) Ethnobiological classification. principles of productivity scholarships to A. 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