life-history traits, geographical range and conservation
TRANSCRIPT
1
Life-history traits, geographical range and conservation aspects of reef fishes from the
Atlantic and Eastern Pacific
Short title: Atlantic and Eastern Pacific reef fishes
Juan Pablo Quimbayo1*, Fernanda Carolina da Silva2, Thiago Costa Mendes3, Débora Silva
Ferrari2, Samara Leopoldino Danielski2, Mariana Gomes Bender4 Valeriano Parravicini5, Michel
Kulbicki6, Sergio Ricardo Floeter2
Affiliations:
1 Center for Marine Biology, Universidade de São Paulo, Brazil
2 Marine Macroecology and Biogeography Lab, Universidade Federal de Santa Catarina, Brazil
3 Instituto do Mar, Universidade Federal de São Paulo, Brazil
4 Marine Macroecology and Conservation Lab, Universidade Federal de Santa Maria, Brazil
5 École Pratique des Hautes Études, Université de Perpignan, France
6 Institut de Recherche pour le Développement IRD, UMR Entropie, Université de Perpignan,
France
*Corresponding author: Juan Pablo Quimbayo, email: [email protected]
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Abstract
Reef fish represent one of the most diverse vertebrate groups on Earth, with over 7000 species
distributed around the globe. This richness is not evenly distributed geographically. The Atlantic
(AT) and the Eastern Pacific (EP) encompass 30% of the global fish fauna. These areas have
been considered the most isolated from the marine biodiversity hotspot in the Indo-Pacific due to
distinct physical barriers, such as the Tethyan closure and the distance between the EP and the
western Pacific. Despite their comparatively lower species richness, these realms host unique
fish assemblages characterized by a remarkable proportion of regional endemics and species with
large body size. Here, we present the largest database of life-history traits, biogeographical and
conservation aspects presently available for the reef fish fauna of the AT and the EP realms. The
database includes 21 traits distributed into behavioral (home range, diel activity, group size, level
in the water column, three measures of preferred temperature), morphological (maximum body
size, size class, body shape, aspect ratio, caudal fin, mouth position) and ecological (trophic
level, diet, spawning strategy, depth of occurrence, two allometric constants, pelagic larval
duration and life span), as well as biogeographical (geographic range index, range extension,
species distribution in 20 marine provinces, Latitude North and South of occurrence, Total
number of provinces where species occur, occurrence in the AT and EP), and conservation
aspects (IUCN status, vulnerability and global market price). We compiled these data through a
careful review of 104 local checklists published between 1982 and 2020, online repositories,
local reports, books, and monographs on specific families or genera. We limited our database to
localities situated between latitudes 51ºN and 45ºS that including shallow and upper mesophotic
biogenic and/or rocky reefs habitats. Our database covers 2,198 species belonging to 146
families and 655 reef fish genera distributed in two marine realms (1,458 in the AT, 829 in the
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EP, and 89 in both realms) and 20 marine provinces. This database of reef fish offers the
opportunity to explore novel ecological and evolutionary questions at different scales and
provides tools for species conservation based on these traits.
Keywords: Taxonomic richness, behavioral traits, morphological traits, ecological traits,
biogeographical information, threatened species, fish price, species distribution, checklist,
Atlantic, Eastern Pacific.
METADATA
CLASS I. DATA SET DESCRIPTORS
A. Data set identity: Life-history traits, geographical range and conservation aspects of reef
fishes from the Atlantic and Eastern Pacific
B. Data set identification code:
• Fish_aspects_EasternPacific_Atlantic_Realms.csv;
• Fish_aspects_EasternPacific_Atlantic_Realms.txt
C. Data set description
1. Originators: Juan Pablo Quimbayo1*, Fernanda Carolina da Silva2, Thiago Costa Mendes3,
Débora Silva Ferrari2, Samara Leopoldino Danielski2, Mariana Gomes Bender4 Valeriano
Parravicini5, Michel Kulbicki6, Sergio Ricardo Floeter2
1 Center for Marine Biology, Universidade de São Paulo, Brazil
4
2 Marine Macroecology and Biogeography Lab, Universidade Federal de Santa Catarina, Brazil
3 Instituto do Mar, Universidade Federal de São Paulo, Brazil
4 Marine Macroecology and Conservation Lab, Universidade Federal de Santa Maria, Brazil
5 École Pratique des Hautes Études, Université de Perpignan, France
6 Institut de Recherche pour le Développment IRD, UMR Entropie, Université de Perpignan,
France
2. Abstract: Reef fish represent one of the most diverse vertebrate groups on Earth, with over
7000 species distributed around the globe. This richness is not evenly distributed geographically.
The Atlantic (AT) and the Eastern Pacific (EP) encompass 30% of the global fish fauna. These
areas have been isolated considered the most isolated from the marine biodiversity hotspot in the
Indo-Pacific due to distinct physical barriers, such as the Tethyan closure and the distance
between the EP and the western Pacific. Despite their comparatively lower species richness,
these realms host unique fish assemblages characterized by a remarkable proportion of regional
endemics and species with large body size. Here, we present the largest database of life-history
traits, biogeographical and conservation aspects presently available for the reef fish fauna of the
AT and the EP realms. The database includes 21 traits distributed into behavioral (home range,
diel activity, group size, level in the water column, three measures of preferred temperature),
morphological (maximum body size, size class, body shape, aspect ratio, caudal fin, mouth
position) and ecological (trophic level, diet, spawning strategy, depth of occurrence, two
allometric constants, pelagic larval duration and life span), as well as biogeographical
(geographic range index, range extension, species distribution in 20 marine provinces, Latitude
North and South of occurrence, Total number of provinces where species occur, occurrence in
the AT and EP), and conservation aspects (IUCN status, vulnerability and global market price).
5
We compiled these data through a careful review of 104 local checklists published between 1982
and 2020, online repositories, local reports, books, and monographs on specific families or
genera. We limited our database to localities situated between latitudes 51ºN and 45ºS that
including shallow and upper mesophotic biogenic and/or rocky reefs habitats. Our database
covers 2,198 species belonging to 146 families and 655 reef fish genera distributed in two marine
realms (1,458 in the AT, 829 in the EP, and 89 in both realms) and 20 marine provinces. This
database of reef offers the opportunity to explore novel ecological and evolutionary questions at
different scales and provides tools for species conservation based on these traits.
3. Keywords: Taxonomic richness, behavioral traits, morphological traits, ecological traits,
biogeographical information, threatened species, fish price, species distribution, checklist,
Atlantic, Eastern Pacific.
CLASS II. RESEARCH ORIGIN DESCRIPTORS
A. Overall project description
1. Identity: Life-history traits, geographical range and conservation aspects of reef fishes from
the Atlantic and Eastern Pacific
2. Originators: Juan Pablo Quimbayo1*, Thiago Costa Mendes2, Mariana Gomes Bender3
Valeriano Parravicini4, Michel Kulbicki5, Sergio Ricardo Floeter6
1 Center for Marine Biology, Universidade de São Paulo, Brazil
2 Instituto do Mar, Universidade Federal de São Paulo, Brazil
3 Marine Macroecology and Conservation Lab, Universidade Federal de Santa Maria, Brazil
6
4 École Pratique des Hautes Études, Université de Perpignan, France
5 Institut de Recherche pour le Développment IRD, UMR Entropie, Université de Perpignan,
France
6 Marine Macroecology and Biogeography Lab, Universidade Federal de Santa Catarina, Brazil
3. Period of study: Data of source publications range from 1982 to 2020.
4. Objectives: (1) Compile the occurrences of all extant reef fish species in the tropical and
subtropical reefs of the Atlantic and Eastern Pacific realms (b) identify behavioral,
morphological and ecological traits, as well as biogeographical and conservation aspects of reef
fish from the Atlantic and Eastern Pacific.
5. Abstract: Same as above
6. Source: Foundation for Research on Biodiversity – France | São Paulo Research Foundation
(FAPESP 2018/21380-0) and Conselho Nacional de Desenvolvimento Científico e Tecnológico
(CNPq 563276/2010-0)
B. Specific subproject description
1. Sites description
Our research comprises 104 localities from the Atlantic and Eastern Pacific marine realms
(Kulbicki et al. 2013) located between latitudes 51ºN and 45ºS (Table 1; Figure 1a). Both realms
are characterized by have a high level of isolation from the past and current biodiversity center,
with low similarity and high endesmim (Kulbicki et al. 2013, Pellissier et al. 2014). The Atlantic
is considered the second most isolated marine realm due to three main biogeographic barriers:
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the emerged Rea Sea land bridge c. 12-18 Myr which separated its fish fauna from the Indian
realm (Floeter et al. 2008, Belmaker et al. 2013); the closure of the Isthmus of Panama c. 3Myr
which disconnected the Atlantic from the Eastern Pacific; and the cold Benguela Current which
has largely constrain the colonization of Atlantic reefs by species arriving from the Indian realm
for at least 2 Myr (Floeter et al. 2008). The Eastern Pacific (EP) is recognized as the most
isolated marine realm due to an approximately 5000-km-wide expanse of deep open ocean that
separates Clipperton Island, the westernmost locality in that region from the nearest Central
Pacific islands. This stretch of deep ocean is known as the Eastern Pacific Barrier “EPB”
(Robertson and Cramer 2009). Moreover, the closure of the Isthmus of Panama c. 3 Myr,
isolated the EP from the Caribbean provinces (Coates and Stallard 2013). Finally, the EP is
strongly influenced by two cold currents (the California current in the north and Humboldt
current in the south) which divide the tropical and subtropical fish fauna in two subset species
with different temperature affinity (Robertson and Cramer 2009). The localities distributed in
these two realms encompass 20 marine provinces (six in the Eastern Pacific and fourteen in the
Atlantic; Figure 1a), which were defined by Spalding et al. (2007) as larger areas that have
distinct biotas consistent across evolutionary time periods.
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Table 1. List of localities (n=104) considered to compile local species chechklist of reef fish in
the Atlantic and Eastern Pacific. These checklists published between 1982 and 2020, online
repositories, local reports, books, and monographs on specific families or genera. The number
refer to ecoregions proposed by Spalding et al. (2007). Biogenic reefs are made up of hard matter
created by living organisms, whereas rocky reefs are formed mainly by inert substract.
Location Spalding
province
Latitude Longitude Reef type References
Abrolhos - Brazil 14 -17.78 -38.89 Biogenic/Rocky (Pinheiro et al. 2018)
Adriatic Sea 4 42.73 14.98 Rocky (Lipej and Dulčić 2010)
Antilles NE 12 21.3 -74.6 Biogenic (Robertson and Van-Tassell
2019)
Antilles SE 12 14.6 -61.1 Biogenic (Robertson and Van-Tassell
2019)
Antilles SW 12 11.8 -71.2 Biogenic (Robertson and Van-Tassell
2019)
Argentina 48 -42 -62 Rocky (Galván et al. 2009, Bovcon et
al. 2011, López et al. 2013)
Arraial do Cabo -
Brazil
47 -22.96 -41.58 Rocky (Pinheiro et al. 2018)
Ascension Island 15 -7.94 -14.37 Rocky (Wirtz et al. 2017)
Atol Rocas - Brazil 14 -3.87 -33.80 Biogenic (Pinheiro et al. 2018)
Azores 3 38.2 -26.8 Rocky (Serrão-Santos et al. 1997)
Bahamas 12 24.59 -77.23 Biogenic (Robertson and Van-Tassell
2019)
Bahia - Brazil 14 -12.91 -38.30 Biogenic/Rocky (Pinheiro et al. 2018)
Baja California 11 25.5 -112.5 Rocky (Pondella et al. 2005)
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Bermudas 12 32.37 -64.81 Biogenic (Robertson and Van-Tassell
2019)
Black Sea 7 42.85 33.47 Rocky (Yankova et al. 2010)
Bonaire Island 12 12.19 -68.65 Biogenic (Robertson and Van-Tassell
2019)
California 11 30.3 -120.3 Rocky (Robertson and Allen 2015)
Canaries 3 28.39 -15.31 Rocky (Dooley et al. 1985)
Cape Verde 16 15.63 -23.26 Rocky (Hanel and John 2015)
Caribbean NW 12 27 -94.4 Biogenic (Robertson and Van-Tassell
2019)
Caribbean SE 12 14.2 -62.3 Biogenic (Robertson and Van-Tassell
2019)
Caribbean SW 12 11.8 -71.2 Biogenic (Robertson and Van-Tassell
2019)
Cayman Island 12 19.4 -81.3 Biogenic (Robertson and Van-Tassell
2019)
Chile Central 45 -30 -72 Rocky (Ojeda et al. 2000)
Chile North 45 -23 -70.7 Rocky (Ojeda et al.. 2000; Pérez-
matus et al.. 2007)
Chile South 48 -43 -74 Rocky (Ojeda et al. 2000)
Clipperton Atoll 43 10.29 -109.22 Biogenic (Allen and Robertson. 1997;
Fourriére et al.. 2014)
Cocos Island 43 5.53 -87.05 Biogenic/Rocky (Fourriére et al. 2017)
Colombia
Caribbean
12 10.41 -75.56 Biogenic (Robertson and Van-Tassell
2019)
Colombia TEP 43 3.78 -77.85 Biogenic/Rocky (Robertson and Allen 2015)
Costa Rica
Caribbean
12 10.02 -83.1 Biogenic (Robertson and Van-Tassell
2019)
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Costa Rica TEP 43 9.60 -84.76 Biogenic/Rocky (Robertson and Allen 2015)
Cuba 12 21.74 -80.34 Biogenic (Robertson and Van-Tassell
2019)
East USA 6 32.7 -79.3 Rocky (Smith-Vaniz et al. 1999)
England South 2 51 -5.5 Rocky (Kulbicki et al. 2013)
Ecuador 43 -1.29 -80.45 Rocky (Robertson and Allen 2015)
Espírito Santo -
Brazil
14 -20.45 -40.05 Rocky (Pinheiro et al. 2018)
F. Noronha - Brazil 14 -3.85 -32.40 Rocky (Pinheiro et al. 2018)
Florida Keys 12 24.71 -81.67 Biogenic (Robertson and Van-Tassell
2019)
France - Atlantic 3 47.3 -3.9 Rocky (Bearez et al. 2017)
Gulf of Mexico NE 6 30 -88.6 Rocky (Robertson and Van-Tassell
2019)
Gulf of Mexico NW 6 26.9 -96.4 Rocky (Robertson and Van-Tassell
2019)
Gulf of Mexico SW 12 20.6 -96 Biogenic (Robertson and Van-Tassell
2019)
Galápagos
Archipelago
44 -0.62 -90.58 Rocky (McCosker and Rosenblatt
2010, Robertson and Allen
2015)
Georgia 6 31.03 -80.57 Rocky (Yankova et al. 2010, 2014)
Gorgona Island 43 2.97 -78.17 Biogenic/Rocky (Zapata and Morales 1997,
Robertson and Allen 2015)
Greece 4 38.2 24.9 Rocky (Papaconstantinou 1988, 2014)
Guadalupe Island 11 24.7 -108.4 Rocky (Reyes-Bonilla et al. 2010)
Guadeloupe -
Caribbean
12 16.19 -61.45 Biogenic (Robertson and Van-Tassell
2019)
11
Guatemala -
Caribbean
12 15.89 -88.43 Biogenic (Robertson and Van-Tassell
2019)
Guatemala TEP 43 13.82 -91.37 Rocky (Robertson and Allen 2015)
Guinea-Sierra
Leone
17 9.16 -14.46 Rocky (Floeter et al. 2008)
Gulf of California 11 28.75 -112.38 Rocky (Ayala-Bocos et al. 2018)
Hispaniola 12 18.3 -69.6 Biogenic (Robertson and Van-Tassell
2019)
Hump - Brazil 14 -5.94 -36.75 Biogenic/Rocky (Pinheiro et al. 2018)
Ilha Grande - Brazil 47 -23.47 -43.60 Rocky (Pinheiro et al. 2018)
Isla la Plata 43 -1.27 -81.06 Rocky (Robertson and Allen 2015)
Israel-Lebanon-
Syria
4 32.9 34.25 Rocky (Golani 2005, Ali 2018,
Bariche and Fricke 2020)
Jamaica 12 17.1 -77.4 Biogenic (Robertson and Van-Tassell
2019)
Juan Fernandez and
Desventuradas
46 -30.03 -79.8 Rocky (Dyer and Westneat. 2010;
Pérez-Matus et al.. 2014)
Lybia-Egypte 4 32.5 25.6 Rocky (Hassan and El-Silini 1999,
Elbaraasi et al. 2019)
Madeira 3 32.8 -16.6 Rocky (Wirtz et al. 2008)
Maghreb 4 37.15 6.4 Rocky (Golani 2005)
Malpelo Island 43 3.99 -81.61 Rocky (McCosker and Rosenblatt
1975, Rubio et al. 1992,
Robertson and Allen 2015)
Manuel Luiz -
Brazil
13 -0.86 -44.27 Biogenic/Rocky (Rocha and Rosa 2001)
Martinique 12 14.74 -60.92 Biogenic (Robertson and Van-Tassell
2019)
12
Mauritania 17 18.85 -16.59 Rocky (Froese and Pauly 2020)
Mediterranean E 4 33.28 30.1 Rocky (Golani 2005)
Mediterranean NW 4 42.25 8.25 Rocky (Psomadakis et al. 2012)
Mediterranean W 4 39.7 7.9 Rocky (Psomadakis et al. 2012)
Mexican Caribbean 12 21.24 -87.23 Biogenic (Robertson and Van-Tassell
2019)
Monterrey Bay 10 36.62 -121.9 Rocky (Burton and Lea 2019)
Namibia 50 -22.9 13.9 Rocky (Froese and Pauly 2020)
Navassa 12 18.39 -75.1 Rocky (Collette et al. 2003)
Nicaragua
Caribbean
12 17 -86 Biogenic (Robertson and Van-Tassell
2019)
Nicaragua TEP 43 11.68 -86.83 Rocky (Robertson and Allen 2015)
North Carolina 6 34.7 -75.6 Rocky (Smith-Vaniz et al. 1999)
Panama Caribbean 12 9.29 -78.17 Biogenic (Robertson and Van-Tassell
2019)
Panama TEP 43 8.51 -79.16 Biogenic/Rocky (Dominici-Arosemena and
Wolff 2006, Robertson and
Allen 2015)
Pelican Cays Belize 12 16.65 -88.2 Biogenic (Smith et al. 2003)
Peru 45 -12 -78 Rocky (Robertson and Allen 2015)
Portugal 3 39.6 -9.5 Rocky (Carneiro et al. 2014)
Puerto Rico 12 18.2 -66.2 Biogenic (Robertson and Van-Tassell
2019)
Puerto Vallarta -
Mexico
23 20.76 -105.56 Biogenic/Rocky (Robertson and Allen 2015)
Revillagigedos 43 18.77 -111.61 Rocky (Fourriére et al. 2016)
Saba Netherlands
Antilles
12 17.63 -63.23 Biogenic (Robertson and Van-Tassell
2019)
13
Salvador - Brazil 43 13.07 -89.16 Biogenic/Rocky (Pinheiro et al. 2018)
Santa Catarina -
Brazil
47 -27.59 -47.96 Rocky (Anderson et al. 2015)
São Paulo - Brazil 47 -24.98 -46.33 Rocky (Feitoza et al.. 2003; Menezes.
2011)
São Tomé 17 0.19 6.61 Rocky (Wirtz et al. 2007)
Senegal - Gambia 17 13.91 -17.11 Rocky (Floeter et al. 2008, Froese and
Pauly 2020)
Sinaloan - Mexico 11 23.79 -107.23 Rocky (Robertson and Allen 2015)
South Carolina 6 32.7 -79.3 Rocky (Smith-Vaniz et al. 1999)
South Florida 12 24.71 -81.67 Biogenic (Robertson and Van-Tassell
2019)
Spain - Atlantic 3 43.9 -5.5 Rocky (Bañon et al. 2011, Báez et al.
2019)
St. Helena 15 -15.97 -5.71 Rocky (Brown et al. 2019)
St. Paul Rocks 14 0.86 -29.38 Rocky (Vaske et al. 2005)
Tres Marias Island 43 21.58 -105.81 Rocky (Erisman et al. 2011)
Trindade Island 14 -20.54 -29.19 Rocky (Pereira-Filho et al.. 2011;
Pinheiro et al.. 2015)
Turkey -
Mediterranean
4 35.48 32.55 Rocky (Bilecenoglu et al. 2002, Fricke
et al. 2007)
Uruguay 47 -35.3 -53.8 Rocky (Nion et al. 2002, Ruarte et al.
2009)
Venezuela 12 10.53 -61.33 Biogenic (Robertson and Van-Tassell
2019)
Virgin Islands 12 18.43 -64.65 Biogenic (Smith-vaniz and Jelks 2014)
Zumbi - Brazil 14 -8.71 -34.97 Biogenic/Rocky (Floeter et al. 2008, Pinheiro et
al. 2018)
14
We found 2,198 reef fish belonging to 146 families and 655 genera distributed heterogeneously
in 20 marine provinces (Figure 1b). The total number of species observed in the different marine
provinces ranges from 42 to 853 species (mean ± SD: 276 ± 197), the Tropical Northwestern
Atlantic province being the richest, whereas the Juan Fernández and Desventuradas Islands, the
poorest (Figure 1b). More species occur in the Atlantic realm than in the Eastern Pacific (1,480
vs 829 exclusive species, respectively), whereas only 89 occur in both realms (Figure 1a).
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Figure 1. Geographical distribution of 104 localities considered in this study (pink diamonds) (a). Each color on the map represents a 1
marine province according to Spalding et al. (2007). (b) Total number of species found in each marine realm defined according to 2
Kulbicki et al. (2013), who defined these realms how the 1st level of the dendrogram builted from cluster analysis used the 3
dissimilarity index ßsim. (c) Comparison of total number of species observed distribution across 20 marine provinces. 4
5
16
2. Experimental or sampling design 6
Data collection period 7
We compiled local checklists for 104 localities from articles published between 1982 and 2020, 8
found online through search tools (Web of Science, Scopus and Google Scholar), online 9
repositories (Robertson and Allen 2016, Pinheiro et al. 2018, Robertson and Van-Tassell 2019, 10
Fricke et al. 2020, Froese and Pauly 2020), local reports, books, and monographs on specific 11
families or genera. We used combinations of the following search terms, in English, Portuguese, 12
French, and Spanish: fish assemblages, fish checklists, new record, reef fish community, reef fish 13
geographical distribution, biogeographic patterns of reef fish, Eastern Pacific and Atlantic. We 14
did not consider Elamosbranchii because species in this group have different biological traits and 15
evolutionary histories when compared to bony fish (see for more details Kulbicki et al. 2013). 16
We limited our search to localities situated between latitudes 51ºN and 45ºS with biogenic and/or 17
rocky reefs habitats. Additionally, we only considered species occurring up to 60 meters deep 18
since this range was the most frequently observed in the compiled checklists and cover both 19
shallow and upper mesophotic biogenic/rock reefs (Rocha et al. 2018). Life-history traits were 20
compiled from the GASPAR (General Approach to Species-Abundance Relationships) research 21
group databases (Halpern and Floeter 2008, Luiz et al. 2013, Kulbicki et al. 2013, Parravicini et 22
al. 2013, Mouillot et al. 2014, Alzate et al. 2019), books and online repositories (Robertson and 23
Allen 2016, Robertson and Van-Tassell 2019, Froese and Pauly 2020). 24
25
26
17
3. Research methods 27
Taxonomy and systematics 28
The taxonomic classification follows the hierarchical system proposed by Linnaeus. All fish 29
names compiled from checklists, online repositories, local reports and monographs were verified 30
according to Eschmeyer's Catalog of Fishes (Fricke et al. 2020). 31
CLASS III. DATA SET STATUS AND ACESSIBILITY 32
A. Status 33
1. Latest update: 2020 34
2. Latest archive data: 2020 35
3. Metadata status: Metadata are complete 36
4. Data verification: All data was reviewed according to CLASS V section B and checked for 37
any input errors. 38
B. Accessibility 39
1. Storage location and medium: All data and metadata are stored with this publication in 40
Ecology journal. 41
2. Contact person: Juan Pablo Quimbayo, Center for Marine Biology, University of São Paulo, 42
Brazil. E-mail: [email protected]; [email protected] 43
18
3. Copyright or proprietary restrictions: There are no copyright or proprietary restrictions for 44
research or teaching purposes. 45
4. Proprietary restrictions: None. 46
a. Citation: This paper 47
CLASS IV. DATA STRUCTURAL DESCRIPTORS 48
A. Data set file 49
1. Identity: Fish_aspects_EasternPacific_Atlantic_Realms.csv 50
2. Size: header + 2,198 rows and 63 columns (622 KB) 51
3. Format and storage: Digital data file in *.csv; *.txt. Columns separate by semicolon and 52
decimal per coma. 53
4. Header information: See column description in section B. 54
5. Alphanumeric information: Mixed 55
B. Variable information 56
Each row in the database represents one species and columns show information on taxonomy 57
classification, behavioral, morphological and ecological traits, biogeography and conservation 58
information (Table 2; Figure 2). 59
19
Table 2. Column names (n=63), variable definition, variable nature and variable units for the 60
database. These columns contain information on behavioral, morphological and ecological traits, 61
as well as biogeographical, and conservation aspects relative to 2,198 reef fish species from the 62
Eastern and Atlantic realms. The realms were defined according to Kulbicki et al. (2013). 63
Variable Name Variable definition Nature Units fb_code This code is an identifier assigned to
each species according to FishBase (Froese and Pauly 2020)
Class Taxonomic class Order Taxonomic order Family Taxonomic family Genus Taxonomic genus
Species Taxonomic specific epithet
Name Scientific Name (Genus and specific epithet)
Home_range Area of activity:
• Sedentary or Territorial species (sed): species that remain in a restricted area (less than a few 10
m²) for extended periods (at least a month).
• Mobile species (mob): species that remain in area of more than 100 m² or traveling among different reef
areas.
• Very mobile (vmob): species which frequently change reefs or
travel large distances on the same reef daily.
Categorical (ordinal)
Diel_activity Period of the day that species feed
and breed:
• day: diurnal species - species active essentially during day time
hours.
• both: species active during both night and day or species with
crepuscular behavior.
• night: nocturnal species – species mainly active during the night.
Categorical (nominal)
20
Size_group Gregariousness behavior:
• solitary (sol): species that usually live alone.
• pair: species usually seen in pair.
• small groups (smallg): species that form small groups (3 to 20 individuals on average).
• medium groups (medg): species that form groups of 21-50 individuals on average.
• large groups (largeg): species that form large groups or schools (> 50 individuals on average).
Categorical
(ordinal)
Level_water Position in water column:
• bottom: species staying on the
bottom at all the times.
• low: species that live slightly
above the bottom but occasionally
rest on the bottom.
• high: species spending most of
their active time high above the bottom (several meters).
Categorical (ordinal)
Occurrence_temperature These data represent the averages
temperature estimated to each species from temperature reported to
each locality where these species occur. The temperature data were estimated from satellite data
averaged between 2002 and 2009 from BIO-oracle (Tyberghein et al.
2012), [Mean=TempOccurrence_mean,
Minimum=TempOccurrence_min, and
Maximum=TempOccurrence_max]
Continuous Degrees Celsius
Body_size_max Maximum total length of species. Continuous Centimeters
Size_class Maximum total length of species
divided into ordered categories according to Mouillot et al. (2014).
• s1: 0 - 7 cm
• s2: 7.1 - 15 cm
• s3: 15.1 - 30 cm
• s4: 30.1 - 50 cm
• s5: 50.1 - 80 cm
Categorical
(ordinal)
Centimeters
21
• s6: > 80 cm
Body_shape Morphological specialization for
swimming.
• box_shaped: globular form, as in Diodontidae.
• compressed: body flattened laterally.
• depressed: body flattened dorsoventrally.
• eel_like: body elongated, but with snake-like shape and locomotion.
• elongated: long in relation to length.
• fusiform: body spindle-shaped, the most hydrodynamic form.
Categorical
(nominal)
Aspect_ratio An indicative value of species activity calculated from the squared
height of caudal fin (h) divided by its surface (s) area (h2/s).
Continuous Nondimensional
Nphoto Number of photos on which the aspect ratio metric was based.
Discrete Nondimensional
Caudal_fin Shape of caudal fin:
• forked: caudal fin with a deeply
concave or excavated hind margin.
• lanceolated: spear shaped caudal fin.
• pointed: when the dorsal and anal fins are joined to the caudal.
• rounded: caudal fin with only one lobe and concave margin.
• truncated: caudal fin with only one lobe and straight margin.
• heterocercal: asymmetric fin.
Categorical (nominal)
Mouth_position Anatomic jaw position:
• superior: lower jaw is protruding upwards.
• terminal: the upper and lower jaws are equal in length.
• subterminal: upper jaw is protruding downwards.
• inferior: mouth located in ventral position.
• tubular: enlargement of mouth cavity.
Categorical
(nominal)
22
• elongated: long snout due to the increase in the bones premaxillary,
maxillary and dental.
Trophic_level Position in the food web based on
diet composition and mean trophic level of food items
Continuous Nondimensional
Diet Main items consumed by species:
• herbivore-detritivore (hd): feeds predominantly on the epilithic
algal matrix (EAM).
• macroalgal feeder (hm): feeds predominantly on macroalgae
(>1cm high) or seagrass.
• sessile invertebrates (is): feeds
predominantly on sessile benthic invertebrates including corals,
sponges and ascidians.
• mobile benthic invertebrates (im): feeds predominantly on mobile
benthic invertebrates including crustaceans, annelids, echinoderms
and mollusks.
• planktonic (pk): feeds predominantly on invertebrate
material in the water column including zooplankton, and,
detrital aggregates.
• omnivore (om): feeds on a range of organisms and includes a large
proportion of animal material and plant and/or detrital material.
• piscivore (fc): feeds predominantly on fishes but may also include cephalopods.
Categorical (nominal)
Spawning The way that species deposit eggs in water:demersal: deposit their eggs
directly on the substrate such as rocks.
• live: species without an external larval stage.pelagic: eggs free in the column water
• attach: species that attach their eggs to objects that float in the
water rather than the bottom substrate.
Categorical (nominal)
23
• oral: species with parental care in which female or male keep the
eggs in their mouth until complete development.
Depth_min Minimal depth reported for the species.
Continuous Meters
Depth_max Maximal depth reported for the species.
Continuous Meters
Depth_range Difference between the maximum and minimum depth reported for the
species
Continuous Meters
Allometric_a Constant allometric coefficient
estimated from the relationship between size and weight
Continuous Nondimensional
Allometric_b Constant allometric coefficient
estimated from the relationship between size and weight
Continuous Nondimensional
Level_length_weight Taxonomic level at which the information was compiled
• family
• genus
• species
Categorical
PLD Pelagic larval duration. Discrete Days
PLD_level Taxonomic level at which the
information was compiled
• family
• genus
• species
Categorical
(nominal)
Life_span Maximum age expected according to growth curves reported in the
FishBase
Discrete Years
Geographic_range_index This index was estimated from
distribution data used to build range maps for each species, defined as the
convex polygon shaping the area where each species is present
(Parravicini et al. 2014). From these maps was sum the number of grid
cell (5º x 5º; corresponding to approximately 550 x 550 km) where
the each species has been recorded.
Discrete Nondimensional
Range_extension Classification on the geographical
origin of species:
Categorical
(nominal)
24
• invasive: species are non-native to
the ecosystem which can causes several alterations in the ecosystem and influence negatively native
species.
• lessepsian: migratory species from
the Red Sea to the Mediterranean.
• native: species with evolutionary history in the Atlantic or Eastern
Pacific.
• transatlantic: species with restricted distribution in one part of the
Atlantic but were recently reported in other localities.
• transpacific: species native from Pacific but were recently reported in Atlantic.
Latitude Coordinate at which the species occurs. These data were compiled
based on species checklists. [Lat_Max_North, Lat_Min_South]
Continuous Degrees
Provinces Presence or absence in provinces distributed on 20 columns named
province_number (binary). These provinces were defined by Spalding
et al. (2007)
Binary 0/1
Total_occupancy_province Total number of provinces where
one species occur
Discrete
Ocurrence_Atlantic Presence or absence of each species in the Atlantic Realm
Binary 0/1
Ocurrence_EasternPacific Presence or absence of each species in the Eastern Pacific Realm
Binary 0/1
IUCN_status Conservation status of species
• NE: Not evaluated
• DD: Data deficient
• LC: Least concerned
• NT: Near threatened
• VU: Vulnerable
• EN: Endangered
• CR: Critically endangered
Categorical (nominal)
Vulnerability Continuous values ranging from 0 to100, provided by FishBase and
based on fish life-history traits. Values closer to 100 represent more
Continuous Nondimensional
25
64
Missing value code: NA 65
Figure 2. Schematic representation of some the behavioral (blue rectangles), morphological 66
(dark yellow rectangles), and ecological traits (green rectangles), as well as biogeographical 67
(purple rectangles), and conservation aspects (red rectangles) included in the database. From left 68
to right schematic figures represent: (1) Home range (very mobile/mobile/sedentary); (2) Diel 69
activity (day/both/nigh); (3) Size group (solitary/par/small group: Smallg/medium group: 70
Medg/large group: Largeg); (4) Level in the water (bottom/low/high); (5) Occurrence 71
temperature (minimal /maximum); (6) Body size (cm); Size class (s1: 0-7 cm/s2: 8-15 cm; s3: 72
16-30 cm); (7) Body shape (box shape/compressed/depressed/eel like/elongated/fusiform); (8) 73
Aspect ratio (h2/s); (9) Caudal fin (forked: Fk/lanceolated: Lc/pointed: Pt/rounded: Rd/truncated: 74
Td/ heterocercal: Ht); (10) Mouth position (superior: Sup/terminal: Ter/subterminal: 75
Subter/inferior: Inf/tubular: Tub/elongated: Elon); (11) Trophic level; (12) Diet (herbivore-76
detritivore: HD/macroalgal feeder: HM/sessile invertebrate feeders: IS/mobile invertebrate 77
feeders: IM/planktivores: PK/omnivores: OM/piscivores: PS); (13) Spawning 78
(demersal/live/pelagic/ attach /oral); (14) Depth range (minimal: min /maximum: max/range: 79
vulnerable species whereas species
with values closer to 0 are less vulnerables
Price_category Price value based on ex-vessel price for the species. Ex-vessel prices are
the prices that fishers receive directly for their catch, or the price at which
the catch is sold when it first enters the supply chain.
• unknow
• low
• medium
• high
• very high
Categorical (ordinal)
26
rang); (15) Geographic range; (16) Range extension (invasive: point line/lessepsian: white 80
dashed line/native: uniform line/ transatlantic: dashed line); (17) Marine provinces; (18) IUCN 81
status (NE: not evaluated/DD: data deficient/LC: least concern/NT: near threatened/VU: 82
vulnerable/EN: endangered/ CR: critically endangered); (19) Price category according to 83
FishBase (unknow/low/medium/high). 84
28
CLASS V. SUPPLEMENTAL DESCRIPTORS 86
A. Data acquisition: See details in the section Class II, more specifically the Data collection. 87
B. Quality assurance/quality control procedures: The name of all records (i.e., 88
species) compiled from104 local checklists published between 1982 and 2020, were reviewed 89
and update according to Eschmeyer's Catalog of Fishes (Fricke et al. 2020). The 21 traits 90
distributed into behavioral (home range, diel activity, group size, level in the water column, three 91
measures of preferred temperature), morphological (maximum body size, size class, body shape, 92
aspect ratio, caudal fin, mouth position) and ecological (trophic level, diet, spawning strategy, 93
depth of occurrence, two allometric constants, pelagic larval duration and life span), were also 94
revised in two ways: 1) checking the several information available in the online repositories 95
(Robertson and Gerald 2015, Pinheiro et al. 2018, Robertson and Van-Tassell 2019, Froese and 96
Pauly 2020); and 2) reviewed criteria used by Mouillot et al (2014). The biogeographical 97
(geographic range index, range extension, species distribution in 20 marine provinces, latitude 98
north and south of occurrence, Total number of provinces where species occur, occurrence in the 99
AT and EP), and conservation aspects (IUCN status, vulnerability and global market price), were 100
reviewed according to external repositories (Froese and Pauly 2020, IUCN 2020). 101
C. Computer programs and data processing algorithms 102
This procedure was performed in R environment ver. 3.2.4 (R Core Team 2020) using plyr 103
(Wickham 2011), reshape (Wickham 2007), ggplot2 (Wickham 2016), rgeos (Bivand et al. 104
2019b), rgdal (Bivand et al. 2019a), rfishbase (Boettiger et al. 2012), sp (Pebesma and Bivand 105
2005) packages. 106
29
Acknowledgments 107
We thank the following projects for helping in the generation of this database: French 108
Foundation for Research on Biodiversity (FRB) and (CESAB) for funding the GASPAR working 109
group (PI: M.K), SISBIOTA‐Mar (PI: S.R.F.; CNPq 563276/2010‐0 and FAPESC 6308/2011‐8), 110
The National Center for Ecological Analysis and Synthesis (NCEAS) of UC Santa Barbara, USA 111
(S.R.F. post doc grant), CNPq (Brazil) for funding the Synthesis on Biodiversity and Ecosystem 112
Services (SinBiose; PI: M.G.B.; CNPq 442417/2019-5), FAPESP (Brazil; PI:J.P.Q. 2018/21380-113
0). This study was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível 114
Superior (CAPES, Brazil), finance Code 001 through a master’s scholarship awarded to S.L.D. 115
and D.S.F. and doctorate scholarship F.C.S. We thank Carlos E.L. Ferreira, Juan José Alvarado, 116
and Andrea Arriaga-Madrigal for continuing inputs and discussions over reef fish life-history 117
traits. 118
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