MARINE AND FISHERY SCIENCES 33 (2): 227-246 (2020)

https://doi.org/10.47193/mafis.3322020301106

ABSTRACT. We compiled georeferenced records of deep demersal fishes from the Colombian

Caribbean Sea in order to assess the level of survey coverage and geographic completeness of

species richness inventories at a scale of 15 min by 15 min cells, in view of threats from fishing and

oil and natural gas exploration. We identified a rich fauna with a minimum of 362 species registered.

Areas with high observed and predicted species richness were identified. Survey coverage and geo-

graphic richness completeness resulted in being deficient with no cell reaching the status of well-

sampled spatial unit, being 83% of the Colombian Caribbean Exclusive Economic Zone bottoms

unexplored, particularly depths beyond 1,000 m. A plea is made for renewed survey efforts with a

focus on the protection of the Colombian Caribbean deep-sea biota.

Key words: Colombian Caribbean, deep fishes, records, soft-bottoms, species richness.

Evaluación de la diversidad de la fauna de peces demersales profundos del Mar Caribe colom-

biano

RESUMEN. Se recopilaron registros georreferenciados de peces demersales profundos del Mar

Caribe colombiano con el fin de evaluar el nivel de cobertura de la prospección y la integridad

geográfica de los inventarios de riqueza específica a una escala de celdas de 15 min por 15 min, en

vista de las amenazas de la pesca y la explotación de petróleo y gas. Identificamos una rica fauna

con un mínimo de 362 especies registradas. Se identificaron áreas con alta riqueza específica obser-

vada y predicha. La cobertura de los registros y la completitud geográfica de la riqueza resultaron

ser deficientes sin que ninguna celda alcanzara el estado de unidad espacial bien muestreada,

estando el 83% de los fondos de la Zona Económica Exclusiva del Caribe colombiano sin explorar,

en particular aquellas profundidades superiores a los 1.000 m. Se hace un llamamiento para renovar

los esfuerzos de prospección con un enfoque en la protección de la biota de aguas profundas del

Caribe colombiano.

Palabras clave: Caribe colombiano, peces de fondo, registros, fondos blandos, riqueza específica.

INTRODUCTION

With the migration of the fishing frontier into deeper waters and the offs-

hore search for oil and natural gas it is now more important than ever to

improve the knowledge on the deep sea biota, particularly with regard to

MARINE IMPACTS IN THE ANTHROPOCENE

Assessment of deep demersal fish fauna diversity of the Colombian

Caribbean Sea

CAMILO B. GARCÍA*and JORGE M. GAMBOA

Departamento de Biología, Universidad Nacional de Colombia, Carrera 45 # 26-85, Bogotá, Colombia

227

Marine and

Fishery Sciences

MAFIS

*Correspondence:

cbgarciar@unal.edu.co

Received: 5 July 2020

Accepted: 14 September 2020

ISSN 2683-7595 (print)

ISSN 2683-7951 (online)

https://ojs.inidep.edu.ar

Journal of the Instituto Nacional de

Investigación y Desarrollo Pesquero

(INIDEP)

This work is licensed under a

Creative Commons Attribution-

NonCommercial-ShareAlike 4.0

International License

fishes, since they are candidates for exploitation

(e.g., Páramo et al. 2017; Grijalba-Bendeck et al.

2019) or might be affected as bycatch (e.g., Pára-

mo et al. 2012). In Colombian Caribbean waters

deep sea research has been sporadic, but three

main periods can be distinguished. In the late six-

ties and early seventies during the past century,

research vessels like the Oregon, Oregon II (U.S.

Fish and Wild Life Service) and Pillsbury (Uni-

versity of Miami, U.S.A.), as well as several

others, conducted trawling both on the continen-

tal shelf and notably on the continental slope, plus

some trawls occurred at depths beyond 1,000 m.

Later on, from the early part of the 20th century,

interest in continental slope fauna gained momen-

tum with the Macrofauna I and II campaigns that

trawled down to a depth of 500 m (see for instan-

ce Saavedra-Díaz et al. 2000; Roa-Varón et al.

2003; Saavedra-Díaz et al. 2004; Roa-Varón et al.

2007). A decade later in the context of exploration

for oil and natural gas, demersal fish samples

were taken in deeper waters around 800 m (ANH

I and II campaigns, Polanco et al. 2010). In sear-

ching for new fishery resources, Páramo et al.

(2011) conducted trawling at depths between 200

and 550 m during years 2009 and 2010. Since

then, to our knowledge, no mayor deep sea traw-

ling surveys have been undertaken in Colombian

Caribbean waters.

Paramount in diversity studies (and with a

focus on conservation) is the characterization of

the geographic distribution of diversity, which

goes hand in hand with the assessment of the

completeness of sampling of surveys, i.e., whet-

her an asymptote in the species accumulation

curve has been reached (Soberón et al. 2007; Hor-

tal et al. 2015). The composition of species lists is

necessary but not sufficient for effective diversity

management and conservation. In this study we

assess the level of survey coverage and geograp-

hical completeness of species richness inventori-

es by constructing a spatialized estimate of deep

soft-bottom demersal fish fauna diversity in

Colombian Caribbean waters, including the San

Andrés and Providencia archipelago and in the

process we highlight areas that have been well-

and poorly surveyed and those that have never

been visited.

MATERIALS AND METHODS

Georeferenced records of demersal deep fishes

were obtained from Polanco (2015), García and

Armenteras (2015, see sources cited), García

(2017a), Acero et al. (2018), Polanco et al. (2019)

and the Global Biodiversity Information Facility

(GBIF 2020). Only records with associated depth

registers deeper than 200 m were included. Valid-

ity of taxonomic names was checked against the

Catalog of Fishes (Fricke et al. 2019). A database

was constructed with the software ModestR (Gar-

cía-Roselló et al. 2013) which allowed further fil-

tering in order to lessen redundancy in the geo-

graphic records. Because our interests focused on

Colombian Caribbean waters, records in our

sources outside the current Colombian Exclusive

Economic Zone (EEZ, obtained from Flanders

Marine Institute 2019) were excluded.

ModestR produces files directly usable for the

module KnowBR (Lobo et al. 2018; Guisande

and Lobo 2019) of the application RWizard

(Guisande et al. 2014) designed to conduct a

search of both well and poorly surveyed spatial

units. The study area, i.e. the Colombian

Caribbean EEZ, was divided into spatial cells of

size 15 min (circa 28 km) by 15 min. We chose

this size as being consistent with previous simi-

lar analyses (García 2017b, 2018). Further set-

tings were as follow: curve =‘Rational’

(Ratkowski 1990) one of the options for adjust-

ing a function to the accumulation of species

with records that function as surrogates for effort

(Lobo 2008; Lobo et al. 2018); estimator =1,

meaning that we constructed the species accumu-

lation curve using the formula from Ugland et al.

(2003); cutoff =1, meaning that if the quotient

228 MARINE AND FISHERY SCIENCES 33 (2): 227-246 (2020)

between number of records and number of

species is 1 in a given spatial unit it is considered

non-informative and completeness is not calcu-

lated; cutoffCompleteness =0, meaning that if

the value of completeness is below this value of

completeness is not calculated and cutoffSlope =

1, meaning that if the final slope of the accumu-

lation curve is higher than this value, complete-

ness is not calculated.

We used the function SurveyQ (survey quality)

of KnowBR to depict well- and poorly-surveyed

equal area cells geographically. Default values

were used. For well sampled spatial units a slope

<0.02, completeness >90% and R/S (records

over species) >15. For poorly sampled cells a

slope >0.3, completeness <50% and R/S < 3.

RESULTS

After filtering, we included a total of 362

species and 6,211 records in the database, these

being the basis of the analysis (Appendix 1). A

map of the study area, records and the shape of the

Parque Nacional Natural Corales de Profundidad

(http://www.parquesnacionales.gov.co) is shown

in Figure 1. The bulk of the records were located

between 200 m and 1,000 m representing 98% of

all records. In turn, records between 200 m and

300 m represented 52% of all records. Apart from

a couple of records in the San Andrés and Provi-

dencia archipelago, sampling was concentrated on

229

GARCÍA AND GAMBOA: DEEP DEMERSAL FISH DIVERSITY IN THE COLOMBIAN CARIBBEAN

Figure 1. Deep demersal fishes record locations inside the Colombian Caribbean Exclusive Economic Zone and Parque Nacional

Natural Corales de Profundidad.

82° W 80° 78° 76° 74° 72° 70°

16° N

14°

12°

10°

8°

Colombia

Records

PNN Corales de Profundidad

Colombian Exclusive Economic Zone

the continental slope with a few records from the

abyssal plains (Figure 1). Table 1 shows those

species with more than 100 records in the database.

The division of the study area in equal area

cells (15 min by 15 min) resulted in 85 cells.

Records from each cell revealed some areas

where sampling was concentrated. The Gulf of

Salamanca, Guajira, Palomino and mixed

coralline bottoms showed cells with an elevated

number of records while most of the cells

received fewer records (Figure 2). Taking into

account that cells depicted were the ones with at

least one record, it was clear that most of the

study area had never been visited (Figure 2). The

Colombian Caribbean EEZ below 200 m amounts

to circa 385,000 km2while the added cell area is

65,072 km2, so around 83% of the deep sea bot-

tom has never been visited. The scarcity of

records resulted in 31 out of the 85 cells for which

completeness could not be calculated as the R/S

quotient was one. Not surprisingly, the observed

richness (Figure 3) closely followed the distribu-

tion pattern of records in cells with a correlation

of 0.9 (Pearson index, p <0.001). The Darien area

is added as harboring a high number of demersal

fish species (Figure 3).

Completeness ranged from 6.9% to 76.9% in

cells with 75.9% of them (41 out of 54 informa-

tive cells) showing completeness above 50%; but

notice that no cell reached 100% completeness

(Figure 4). The ten cells showing completeness

above 70% were well- distributed along the coast

line (Figure 4), suggesting that the survey effort

was not spatially biased in terms of completeness.

No cell attains the status of ‘high quality sur-

vey’ (Figure 5); but most cells were labeled as

‘fair quality survey’ cells with a number of inter-

dispersed ‘poor quality survey’ cells that is in line

with the image in Figure 4.

DISCUSSION

Our focus on the current Colombian EEZ

caused 32 species and associated records to be

dropped from the database (Appendix 2). These

records are located to the west of the San Andrés

and Providencia archipelago and offshore Pana-

ma outside of Colombian EEZ. Thus, it is doubt-

ful whether the species concerned should be

included in deep sea fishes national biodiversity

230 MARINE AND FISHERY SCIENCES 33 (2): 227-246 (2020)

Table 1. Species with more than 100 georeferenced records in the database. Minimum, maximum and mean depths.

Species Records Min depth (m) Max depth (m) Mean depth (m)

Dibranchus atlanticus 207 198 1,440 412

Nezumia aequalis 150 223 1,143 392

Coelorinchus caelorhincus 134 200 810 327

Laemonema goodebeanorum 126 223 777 348

Chauliodus sloani 125 191 4,151 433

Synagrops bellus 122 192 810 352

Poecilopsetta inermis 113 229 750 292

Chlorophthalmus agassizi 109 200 776 301

Malacocephalus occidentalis 106 200 801 314

Neoscopelus macrolepidotus 104 276 900 475

Chaunax suttkusi 103 223 801 333

lists, although their presence in waters of the

archipelago that belong to Colombia is likely.

Notice that Bolaños-Cubillos et al. (2015) and

Acero et al. (2019) mention several of the species

excluded here in their species lists for the San

Andrés and Providencia archipelago.

The scarceness of surveys and samples east of

the archipelago and in depths beyond 1,000 m

231

GARCÍA AND GAMBOA: DEEP DEMERSAL FISH DIVERSITY IN THE COLOMBIAN CARIBBEAN

Figure 3 Observed richness of deep demersal fishes in cells of 15 min by 15 min in the Colombian Caribbean Exclusive

Economic Zone.

Figure 2. Spatial distribution of deep demersal fishes records in cells of 15 min by 15 min in the Colombian Caribbean Exclusive

Economic Zone.

626

548

469

391

313

235

157

79

1

15° N

14°

13°

12°

11°

10°

9°

8°

82° W 80° 78° 76° 74° 72°

Guajira

Golfo de Salamanca

Mixed coralline bottoms

139

122

105

87

70

53

36

18

1

15° N

14°

13°

12°

11°

10°

9°

8°

82° W 80° 78° 76° 74° 72°

Guajira

Golfo de Salamanca

Mixed coralline bottoms

Palomino

Darien

shown here clearly points to the need of increased

survey efforts, including for those unexplored

areas as well as renewed survey efforts in areas

visited in the past, as even the upper slope that

has received most records is at best ‘fairly-sam-

pled’, according to the criteria here and at our

232 MARINE AND FISHERY SCIENCES 33 (2): 227-246 (2020)

Figure 4 Percentage completeness of deep demersal fishes inventories in cells of 15 min by 15 min in the Colombian Caribbean

Exclusive Economic Zone.

Figure 5 Survey quality status of deep demersal fishes in cells of 15 min by 15 min in the Colombian Caribbean Exclusive

Economic Zone.

77

68

59

51

42

33

24

16

7

15° N

14°

13°

12°

11°

10°

9°

8°

82° W 80° 78° 76° 74° 72°

Guajira

Golfo de Salamanca

Mixed coralline bottoms

Palomino

Darien

15° N

14°

13°

12°

11°

10°

9°

82° W 80° 78° 76° 74° 72° 70°

High quality survey

Fair quality survey

Poor quality survey

spatial scale. Nevertheless, this study reveals a

rich deep sea fish fauna taking into account that

the 362 species in our database are a lower limit

of species richness. Considering the values of

percentage completeness and the current geo-

graphic coverage of surveys, more records for

species already in the database, i.e., extension of

their presently observed distribution, and more

species currently not in the database, are to be

expected in future campaigns.

The distribution of records with species fol-

lows the usual patterns for large diversity data-

bases: few species with many records and many

species with few records. Thus, 49.2% of the

species (178 out of 362) show five or fewer

records while just 3.0% of the species (11 out of

362) show more than one hundred records

(Appendix 1). Interestingly Epigonus occidental-

is (Goode and Bean 1896) and E. pandionis

(Goode and Bean 1881), postulated as possible

candidates for fishing due to their frequency in

trawls (Páramo et al. 2017) do not belong to the

most common species in the database but rank

low to intermediary in terms of records (Appen-

dix 1). Our time window spans decades while

Páramo et al. (2017) are just snap shots of deep

demersal fish presence and abundance in a limit-

ed depth range. This contrast highlights the use-

fulness and, indeed, a need for monitoring over

long periods and extended areas if we are to

understand the dynamics of ecosystems and the

biology of the species.

In view of the results it is probably risky to

postulate areas for conservation purposes.

Observed richness suggests continental slope

areas adjacent to the Gulf of Salamanca, Rosario

Island archipelago (mixed coralline bottoms),

slope areas to the north of Guajira Peninsula and

Darien as locations of accumulation of species.

Interestingly, the Gulf of Salamanca shelf (<200

m), since here the slope (>200 m) is known to

harbor significant numbers of elasmobranchs and

bony fish species (García 2017b, 2018; but

notice that García 2018 includes some slope

records). The ecological and biological reasons

for these findings are an open question worth

investigating.

The only national natural park dedicated to

deep sea biota in the Colombian Caribbean is the

Parque Nacional Natural Corales de Profundidad

established to protect deep sea coral species. The

park includes soft and mixed bottoms from 34 m

to 1,240 m depth. Using the module MRFinder of

ModestR a species list was composed for the

park resulting in 106 species with records inside

the park area (Appendix 3). Thus, although not

intentionally, Corales de Profundidad might also

be protecting a significant fraction of deep dem-

ersal fishes in the Colombian Caribbean. The

attractive effect of structures on fishes is well-

known, thus the presence of coral formations in

the general area of the park is probably conducive

of high species richness. Notice that cells in the

park area and in their vicinity show high ob-

served richness.

This research is one of the first steps in endeav-

ors of studying the diversity of deep sea demersal

fish species beyond the compilation of species’

names. Much work remains to be done both for

scientific and practical purposes with a focus on

protecting Colombian Caribbean deep sea biodi-

versity as derived from this assessment.

ACKNOWLEDGMENTS

Comments by two anonimous reviewers helped

to improve the manuscript.

REFERENCES

ACERO A, TAVERA JJ, POLANCO A, BOLAÑOS-

CUBILLOS N. 2019. Fish biodiversity in three

northern islands of the seaflower biosphere

reserve (Colombian Caribbean). Front Mar

233

GARCÍA AND GAMBOA: DEEP DEMERSAL FISH DIVERSITY IN THE COLOMBIAN CARIBBEAN

Sci. 6: 113. doi:10.3389/fmars.2019.00113
ACERO A, POLO-SILVA CJ, LEÓN G, PUENTES V.
2018. First report of a sleeper shark (Somnio-
sus sp.) in the southern Colombian Caribbean.
J Appl Ichthyol. 34 (4): 981-983. doi:10.1111/
jai.13712
BOLAÑOS-CUBILLOS N, ABRIL-HOWARD A, BENT-
HOOKER H, CALDAS JP, ACERO A. 2015. Lista
de peces conocidos del archipiélago de San
Andrés, Providencia y Santa Catalina, reserva
de biosfera Seaflower, Caribe occidental co-
lombiano. Bol Investig Mar Cost. 44 (1): 127-
162. doi:10.25268/bimc.invemar.2015.44.1.24
FLANDERS MARINE INSTITUTE. 2019. Maritime
boundaries geodatabase: maritime boundaries
and Exclusive Economic Zones (200NM),
version 11. [accessed 2020 July]. https://www.
marineregions.org/. doi:10.14284/386
FRICKE R, ESCHMEYER WN, VAN DER LAAN R. 2019,
editors. Catalog of fishes: genera, species,
references. [accessed 2020 June]. http://
researcharchive.calacademy.org/research/
ichthyology/catalog/fishcatmain.asp.
GARCÍA CB. 2017a. Presence and distribution of
Chimaeriformes in the Colombian Caribbean
Sea. Pan Am J Aquat Sci. 12 (1): 85-88
GARCÍA CB. 2017b. What do we know about soft-
bottom elasmobranch species richness in the
Colombian Caribbean and of its spatial distri-
bution? Reg Stud Mar Sci. 9: 62-68.
doi:10.1016/j.rsma.2016.11.006
GARCÍA CB. 2018. Spatial richness patterns of
soft-bottom fish in the Colombian Caribbean
continental shelf and slope. Acta Biol Colomb.
23 (1): 59-65. doi:10.15446/abc.v23n1.66530
GARCÍA CB, ARMENTERAS D. 2015. Atlas de la
ictiofauna demersal de fondos blandos del
Caribe continental colombiano: aproximación
a su biodiversidad. Bogotá: Universidad
Nacional de Colombia, Publicaciones Facul-
tad de Ciencias. 765 p.
GARCÍA-ROSELLÓ E, GUISANDE C, GONZÁLES-
DACOSTA J, HEINE J, PELAYO-VILLAMIL P,
MANJARRÉS-HERNÁNDEZ A, VAAMONDE A,
GRANADO-LORENCIO C. 2013. ModestR: a
software tool for managing and analyzing
species distribution map databases. Ecogra-
phy. 36: 1202-1207. doi:10.1111/j.1600-
0587.2013.00374.x
[GBIF] GLOBAL BIODIVERSITY INFORMATION
FACILITY. 2020. GBIF Home Page. [accessed
2020 June]. https://www.gbif.org.
GRIJALBA-BENDECK M, PARAMO J, WOLFF M.
2019. Catch composition of deep-sea resources
of commercial importance in the Colombian
Caribbean. Rev Biol Mar Oceanogr. 54 (2):
188-197. doi:10.22370/rbmo.2019.54.2.1891
GUISANDE C, HEINE J, GONZÁLEZ-DACOSTA J,
GARCÍA-ROSELLÓ E. 2014. RWizard Software.
http://www.ipez.es/RWizard. University of
Vigo, Vigo.
GUISANDE C, LOBO JM. 2019. Discriminating
well surveyed spatial units from exhaustive
biodiversity databases. R package version.
2.0. https://cran.r-project.org/web/packages/
KnowBR.
HORTAL J, DE BELLO F, DINIZ-FILHO FAA, LEWIN-
SHON TM, LOBO JM, LADLE RJ. 2015. Seven
shortfalls that beset large-scale knowledge of
biodiversity. Annu Rev Ecol Evol Syst. 46:
523-549. doi:10.1146/annurev-ecolsys-112414-
054400
LOBO JM. 2008. Database records as a surrogate
for sampling effort provide higher species
richness estimations. Biodivers Conserv. 17:
873-881. doi:10.1007/s10531-008-9333-4
LOBO JM, HORTAL J, YELAB JL, MILLÁN A, SÁN-
CHEZ-FERNÁNDEZ D, GARCÍA-ROSELLÓ E, GON-
ZÁLEZ-DACOSTA J, HEINEE J, GONZÁLEZ-VILAS
L, GUISANDE C. 2018. KnowBR: an applica-
tion to map the geographical variation of sur-
vey effort and identify well-surveyed areas
from biodiversity databases. Ecol Indic. 91:
241-248. doi:10.1016/j.ecolind.2018.03.077
PÁRAMO J, FUENTES D, WIFF R. 2017. Population
structure and distribution of deep-water Cardi-
nal Fish Epigonus occidentalis (Epigonidae)
and  Epigonus pandionis (Epigonidae) in the
234 MARINE AND FISHERY SCIENCES 33 (2): 227-246 (2020)

Colombian Caribbean Sea. J Ichthyol. 57 (3):
424-433. doi:10.1134/S0032945217030109
PÁRAMO J, SAINT-PAUL U. 2012. Deep-sea
shrimps Aristaeomorpha foliacea and Pleoti-
cus robustus (Crustacea: Penaeoidea) in the
Colombian Caribbean Sea as a new potential
fishing resource. J Mar Biol Assoc UK. 92 (4):
811-818. doi:10.1017/S0025315411001202
PÁRAMO J, SAINT-PAUL U, MORENO F, PACHECO
M, ALMANZA M, RODRÍGUEZ E, ARDILA G,
BORDA C, BARRETO-GONZÁLEZ H. 2011. Crus-
táceos de profundidad en el Caribe colombia-
no como nuevo recurso pesquero. Informe
Final Santa Marta: Universidad del Magdale-
na, Santa Marta.
POLANCO A. 2015. Dynamics of the continental
slope demersal fish community in the Colom-
bian Caribbean-Deep-sea research in the
Caribbean [PhD thesis]. Giessen: Justus-Lie-
big University Giessen, Bogotá: Universidad
Nacional de Colombia. 192 p.
POLANCO A, ACERO A, GARRIDO M. 2010. Aportes
a la biodiversidad íctica del Caribe colombia-
no. In: INVEMAR, editors. Biodiversidad del
margen continental del Caribe colombiano.
Serie de Publicaciones Especiales, Invemar.
20. p. 318-353.
POLANCO A, DUEÑAS LF, LEÓN J, PUENTES V.
2019. New records and update on the geo-
graphic distribution of the Bony-eared Ass-
fish,  Acanthonus armatus Günther, 1878
(Ophidiidae, Neobythitinae), in the Caribbean
region. Check List. 15 (5): 767-772. doi:10.
15560/15.5.767
RATKOWSKI DA. 1990 Handbook of nonlinear
regression models. New York: Marcel Dekker.
241 p.
ROA-VARÓN A, SAAVEDRA LM, ACERO A, MEJÍA
LS. 2007. Nuevos registros de peces para el
Caribe colombiano de los órdenes Myctophi-
formes, Polymiixiformes, Gadiformes, Ophi-
diiformes y Lophiiformes. Bol Investig Mar
Cost. 36: 181-207. doi:10.25268/bimc.inve-
mar.2007.36.0.206 
ROA-VARÓN A, SAAVEDRA LM, ACERO A, MEJÍA
LS, NAVAS G. 2003. Nuevos registros de peces
óseos para el Caribe colombiano de los órde-
nes Beryciformes, Zeiformes, Perciformes y
Tetraodontiformes. Bol Investig Mar Cost. 32:
3-24. doi:10.25268/bimc.invemar.2003.32.0.257
SAAVEDRA-DÍAZ LM, ACERO A, NAVAS GR. 2000.
Lenguados de la familia Paralichthyidae (Pis-
ces: Pleuronectiformes) conocidos del inclu-
yendo un nuevo registro para el área. Rev
Acad Colomb Cienc Exactas Fis Nat. 24 (91):
295-310.
SAAVEDRA-DÍAZ LM, ROA-VARÓN A, ACERO A,
MEJÍA LS. 2004. Primeros registros ícticos en el
talud superior del Caribe colombiano (órdenes
Albuliformes, Anguilliformes, Stomiiformes,
Ateleopodiformes, Aulopiformes y Pleuronec-
tiformes). Bol Investig Mar Cost. 33: 159-183.
doi:10.25268/bimc.invemar.2004.33.0.254
SOBERÓN J, JIMENEZ R, GOLUBOV J, KOLEFF P.
2007. Assessing completeness of biodiversity
databases at different spatial scales. Ecogra-
phy. 30: 152-160. doi:10.1111/j.0906-7590.
2007.04627.x
UGLAND KI, GRAY JS, ELLINGSEN KE. 2003. The
species-accumulation curve and estimation of
species richness. J Anim Ecol. 72: 888-897.
doi:10.1046/j.1365-2656.2003.00748.x
235
GARCÍA AND GAMBOA: DEEP DEMERSAL FISH DIVERSITY IN THE COLOMBIAN CARIBBEAN

APPENDIX 1

Deep demersal fish species list (> 200 m depth)

of the Colombian Caribbean Sea with records and

depth ranges.

236 MARINE AND FISHERY SCIENCES 33 (2): 227-246 (2020)

Class Order Family Species Records Min depth Max depth Mean depth

Actinopterygii Anguilliformes Chlopsidae Robinsia catherinae 1 300 300 300

Actinopterygii Anguilliformes Colocongridae Coloconger meadi 60 218 777 361

Actinopterygii Anguilliformes Congridae Ariosoma balearicum 3 532 857 706

Actinopterygii Anguilliformes Congridae Bathycongrus bullisi 26 229 366 286

Actinopterygii Anguilliformes Congridae Bathycongrus vicinalis 1 366 366 366

Actinopterygii Anguilliformes Congridae Bathyuroconger vicinus 9 515 732 627

Actinopterygii Anguilliformes Congridae Japonoconger caribbeus 22 269 549 358

Actinopterygii Anguilliformes Congridae Parabathymyrus oregoni 2 314 315 315

Actinopterygii Anguilliformes Congridae Pseudophichthys splendens 57 204 803 392

Actinopterygii Anguilliformes Congridae Rhynchoconger flavus 3 732 732 732

Actinopterygii Anguilliformes Congridae Xenomystax austrinus 23 458 732 509

Actinopterygii Anguilliformes Congridae Xenomystax bidentatus 16 296 698 405

Actinopterygii Anguilliformes Congridae Xenomystax congroides 18 210 352 237

Actinopterygii Anguilliformes Moringuidae Neoconger mucronatus 3 200 265 243

Actinopterygii Anguilliformes Muraenesocidae Cynoponticus savanna 2 746 746 746

Actinopterygii Anguilliformes Muraenidae Gymnothorax conspersus 9 256 307 262

Actinopterygii Anguilliformes Muraenidae Gymnothorax polygonius 2 200 203 202

Actinopterygii Anguilliformes Nemichthyidae Avocettina infans 3 350 445 360

Actinopterygii Anguilliformes Nemichthyidae Labichthys carinatus 1 940 940 940

Actinopterygii Anguilliformes Nemichthyidae Nemichthys scolopaceus 6 300 3,978 654

Actinopterygii Anguilliformes Nettastomatidae Hoplunnis diomediana 1 453 453 453

Actinopterygii Anguilliformes Nettastomatidae Hoplunnis megista 2 366 366 366

Actinopterygii Anguilliformes Nettastomatidae Hoplunnis tenuis 5 201 366 215

Actinopterygii Anguilliformes Nettastomatidae Nettastoma melanura 5 503 732 621

Actinopterygii Anguilliformes Nettastomatidae Venefica procera 3 613 860 834

Actinopterygii Anguilliformes Ophichthidae Ophichthus cruentifer 4 496 699 548

Actinopterygii Anguilliformes Ophichthidae Ophichthus puncticeps 4 295 900 448

Actinopterygii Anguilliformes Synaphobranchidae Atractodenchelys phrix 2 393 500 447

Actinopterygii Anguilliformes Synaphobranchidae Dysommina rugosa 2 365 450 429

Actinopterygii Anguilliformes Synaphobranchidae Ilyophis brunneus 8 515 1143 692

Actinopterygii Anguilliformes Synaphobranchidae Synaphobranchus affinis 1 732 732 732

Actinopterygii Anguilliformes Synaphobranchidae Synaphobranchus oregoni 8 265 515 301

Actinopterygii Ateleopodiformes Ateleopodidae Ijimaia antillarum 30 329 698 423

Actinopterygii Ateleopodiformes Ateleopodidae Ijimaia loppei 1 503 503 503

Actinopterygii Aulopiformes Bathysauridae Bathysaurus mollis 2 1,800 4,151 2,976

Actinopterygii Aulopiformes Chlorophthalmidae Chlorophthalmus agassizi 109 200 776 301

Actinopterygii Aulopiformes Chlorophthalmidae Parasudis truculenta 34 223 561 311

Actinopterygii Aulopiformes Ipnopidae Bathypterois bigelowi 38 223 940 456

Actinopterygii Aulopiformes Ipnopidae Bathypterois grallator 5 1627 4,151 2,435

Actinopterygii Aulopiformes Ipnopidae Bathypterois phenax 6 821 1,800 1,230

Actinopterygii Aulopiformes Ipnopidae Bathypterois quadrifilis 5 515 900 683

Actinopterygii Aulopiformes Ipnopidae Bathypterois viridensis 17 276 900 475

Actinopterygii Aulopiformes Ipnopidae Bathytyphlops marionae 1 1,251 1,251 1,251

Actinopterygii Aulopiformes Ipnopidae Ipnops murrayi 3 821 1,800 1,416

Actinopterygii Aulopiformes Notosudidae Scopelosaurus smithii 2 622 622 622

Actinopterygii Aulopiformes Synodontidae Saurida brasiliensis 5 270 613 427

Actinopterygii Aulopiformes Synodontidae Saurida caribbaea 9 200 303 219

Actinopterygii Aulopiformes Synodontidae Saurida normani 3 198 298 251

Appendix 1. Continued.

237

GARCÍA AND GAMBOA: DEEP DEMERSAL FISH DIVERSITY IN THE COLOMBIAN CARIBBEAN

Actinopterygii Aulopiformes Synodontidae Synodus poeyi 1 198 198 198

Actinopterygii Batrachoidiformes Batrachoididae Porichthys plectrodon 5 200 370 241

Actinopterygii Beloniformes Hemiramphidae Hemiramphus balao 2 276 803 406

Actinopterygii Beryciformes Anoplogastridae Anoplogaster cornuta 1 300 300 300

Actinopterygii Beryciformes Berycidae Beryx decadactylus 1 378 378 378

Actinopterygii Beryciformes Berycidae Beryx splendens 2 366 369 367

Actinopterygii Beryciformes Diretmidae Diretmoides pauciradiatus 9 200 732 268

Actinopterygii Beryciformes Diretmidae Diretmus argenteus 6 205 792 454

Actinopterygii Beryciformes Holocentridae Ostichthys trachypoma 6 200 274 236

Actinopterygii Beryciformes Trachichthyidae Gephyroberyx darwinii 8 300 375 322

Actinopterygii Beryciformes Trachichthyidae Hoplostethus mediterraneus 2 288 435 362

Actinopterygii Beryciformes Trachichthyidae Hoplostethus occidentalis 64 329 558 350

Actinopterygii Cetomimiformes Barbourisiidae Barbourisia rufa 1 1,080 1,080 1,080

Actinopterygii Gadiformes Bregmacerotidae Bregmaceros atlanticus 53 200 801 319

Actinopterygii Gadiformes Macrouridae Bathygadus favosus 21 515 1,440 758

Actinopterygii Gadiformes Macrouridae Bathygadus macrops 96 269 810 422

Actinopterygii Gadiformes Macrouridae Bathygadus melanobranchus 28 365 900 634

Actinopterygii Gadiformes Macrouridae Cetonurus globiceps 5 613 1,097 906

Actinopterygii Gadiformes Macrouridae Coelorinchus caelorhincus 134 200 810 327

Actinopterygii Gadiformes Macrouridae Coelorinchus caribbaeus 96 200 503 276

Actinopterygii Gadiformes Macrouridae Coryphaenoides mexicanus 12 198 1,296 884

Actinopterygii Gadiformes Macrouridae Coryphaenoides zaniophorus 25 518 940 636

Actinopterygii Gadiformes Macrouridae Gadomus arcuatus 17 477 1,080 665

Actinopterygii Gadiformes Macrouridae Gadomus dispar 4 457 622 490

Actinopterygii Gadiformes Macrouridae Gadomus longifilis 23 365 1,251 644

Actinopterygii Gadiformes Macrouridae Hymenocephalus aterrimus 1 576 576 576

Actinopterygii Gadiformes Macrouridae Hymenocephalus billsam 2 223 373 361

Actinopterygii Gadiformes Macrouridae Hymenocephalus italicus 91 269 940 368

Actinopterygii Gadiformes Macrouridae Kuronezumia bubonis 3 515 732 586

Actinopterygii Gadiformes Macrouridae Malacocephalus laevis 13 329 612 387

Actinopterygii Gadiformes Macrouridae Malacocephalus occidentalis 106 200 801 314

Actinopterygii Gadiformes Macrouridae Nezumia aequalis 150 223 1,143 392

Actinopterygii Gadiformes Macrouridae Nezumia cyrano 16 453 960 630

Actinopterygii Gadiformes Macrouridae Nezumia suilla 5 365 640 572

Actinopterygii Gadiformes Macrouridae Sphagemacrurus grenadae 6 684 1,143 779

Actinopterygii Gadiformes Macrouridae Squalogadus modificatus 4 1,080 1,251 1,139

Actinopterygii Gadiformes Macrouridae Trachonurus sulcatus 10 612 1,251 814

Actinopterygii Gadiformes Macrouridae Trachonurus villosus 3 515 814 714

Actinopterygii Gadiformes Macrouridae Ventrifossa macropogon 7 223 640 411

Actinopterygii Gadiformes Macrouridae Ventrifossa mucocephalus 5 450 732 481

Actinopterygii Gadiformes Merlucciidae Merluccius albidus 47 219 662 297

Actinopterygii Gadiformes Merlucciidae Steindachneria argentea 66 200 770 298

Actinopterygii Gadiformes Moridae Gadella imberbis 32 223 801 374

Actinopterygii Gadiformes Moridae Laemonema goodebeanorum 126 223 777 348

Actinopterygii Gadiformes Moridae Physiculus fulvus 6 275 503 341

Actinopterygii Gadiformes Phycidae Urophycis cirrata 11 270 470 331

Actinopterygii Lophiiformes Antennariidae Fowlerichthys radiosus 1 192 192 192

Actinopterygii Lophiiformes Chaunacidae Chaunax pictus 57 270 810 417

Actinopterygii Lophiiformes Chaunacidae Chaunax suttkusi 103 223 801 333

Actinopterygii Lophiiformes Diceratiidae Bufoceratias wedli 4 405 493 478

Actinopterygii Lophiiformes Lophiidae Lophiodes beroe 2 512 512 512

Class Order Family Species Records Min depth Max depth Mean depth

Appendix 1. Continued.

238 MARINE AND FISHERY SCIENCES 33 (2): 227-246 (2020)

Actinopterygii Lophiiformes Lophiidae Lophiodes monodi 7 313 520 346

Actinopterygii Lophiiformes Lophiidae Lophiodes reticulatus 11 200 500 240

Actinopterygii Lophiiformes Lophiidae Lophius gastrophysus 17 286 561 347

Actinopterygii Lophiiformes Lophiidae Lophius piscatorius 4 200 500 240

Actinopterygii Lophiiformes Melanocetidae Melanocetus murrayi 7 523 857 562

Actinopterygii Lophiiformes Ogcocephalidae Dibranchus atlanticus 207 198 1,440 412

Actinopterygii Lophiiformes Ogcocephalidae Dibranchus tremendus 6 1,006 1,463 1,107

Actinopterygii Lophiiformes Ogcocephalidae Halieutichthys aculeatus 20 286 295 291

Actinopterygii Lophiiformes Ogcocephalidae Malthopsis gnoma 12 280 491 301

Actinopterygii Lophiiformes Ogcocephalidae Ogcocephalus declivirostris 3 365 223 221

Actinopterygii Lophiiformes Ogcocephalidae Ogcocephalus parvus 6 270 360 308

Actinopterygii Lophiiformes Ogcocephalidae Ogcocephalus pumilus 1 290 290 290

Actinopterygii Lophiiformes Ogcocephalidae Zalieutes mcgintyi 29 201 498 246

Actinopterygii Lophiiformes Oneirodidae Dolopichthys pullatus 1 4,029 4,029 4,029

Actinopterygii Lophiiformes Thaumatichthyidae Thaumatichthys binghami 1 1,251 1,251 1,251

Actinopterygii Myctophiformes Myctophidae Bolinichthys supralateralis 26 195 792 361

Actinopterygii Myctophiformes Myctophidae Dasyscopelus selenops 2 275 520 396

Actinopterygii Myctophiformes Myctophidae Diaphus adenomus 1 567 567 567

Actinopterygii Myctophiformes Myctophidae Diaphus bertelseni 2 200 200 200

Actinopterygii Myctophiformes Myctophidae Diaphus dumerilii 26 200 727 299

Actinopterygii Myctophiformes Myctophidae Diaphus effulgens 2 223 365 330

Actinopterygii Myctophiformes Myctophidae Diaphus garmani 24 200 1,829 409

Actinopterygii Myctophiformes Myctophidae Diaphus lucidus 23 191 755 327

Actinopterygii Myctophiformes Myctophidae Diaphus minax 1 270 270 270

Actinopterygii Myctophiformes Myctophidae Diaphus rafinesquii 5 320 1,800 633

Actinopterygii Myctophiformes Myctophidae Diaphus splendidus 13 200 857 262

Actinopterygii Myctophiformes Myctophidae Diaphus taaningi 1 270 270 270

Actinopterygii Myctophiformes Myctophidae Lampadena luminosa 9 200 730 422

Actinopterygii Myctophiformes Myctophidae Lepidophanes guentheri 21 200 857 302

Actinopterygii Myctophiformes Myctophidae Myctophum nitidulum 16 200 727 340

Actinopterygii Myctophiformes Neoscopelidae Neoscopelus macrolepidotus 104 276 900 475

Actinopterygii Myctophiformes Neoscopelidae Neoscopelus microchir 23 223 803 372

Actinopterygii Notacanthiformes Halosauridae Aldrovandia affinis 5 515 1,097 559

Actinopterygii Notacanthiformes Halosauridae Aldrovandia gracilis 8 395 1,710 1,039

Actinopterygii Notacanthiformes Halosauridae Halosaurus guentheri 32 276 1,143 433

Actinopterygii Notacanthiformes Halosauridae Halosaurus ovenii 88 276 803 406

Actinopterygii Notacanthiformes Notacanthidae Notacanthus chemnitzii 1 724 724 724

Actinopterygii Notacanthiformes Notacanthidae Polyacanthonotus merretti 2 679 857 738

Actinopterygii Ophidiiformes Aphyonidae Barathronus bicolor 22 365 1,251 625

Actinopterygii Ophidiiformes Bythitidae Calamopteryx robinsorum 2 200 201 201

Actinopterygii Ophidiiformes Bythitidae Cataetyx laticeps 7 732 1,800 1,119

Actinopterygii Ophidiiformes Bythitidae Diplacanthopoma brachysoma 62 274 776 401

Actinopterygii Ophidiiformes Bythitidae Saccogaster staigeri 1 356 356 356

Actinopterygii Ophidiiformes Ophidiidae Acanthonus armatus 13 2,215 2,564 2,366

Actinopterygii Ophidiiformes Ophidiidae Bassozetus robustus 1 1,240 1,240 1,240

Actinopterygii Ophidiiformes Ophidiidae Bathyonus laticeps 2 1,627 2,983 2,531

Actinopterygii Ophidiiformes Ophidiidae Benthocometes robustus 4 223 303 287

Actinopterygii Ophidiiformes Ophidiidae Dicrolene introniger 37 395 1,296 715

Actinopterygii Ophidiiformes Ophidiidae Lamprogrammus brunswigi 2 1,317 1,317 1,317

Actinopterygii Ophidiiformes Ophidiidae Lamprogrammus niger 4 515 3978 1,105

Actinopterygii Ophidiiformes Ophidiidae Lepophidium brevibarbe 18 210 505 277

Class Order Family Species Records Min depth Max depth Mean depth

Appendix 1. Continued.

239

GARCÍA AND GAMBOA: DEEP DEMERSAL FISH DIVERSITY IN THE COLOMBIAN CARIBBEAN

Actinopterygii Ophidiiformes Ophidiidae Lepophidium cultratum 2 210 210 210

Actinopterygii Ophidiiformes Ophidiidae Lepophidium kallion 1 219 219 219

Actinopterygii Ophidiiformes Ophidiidae Lepophidium profundorum 8 207 404 288

Actinopterygii Ophidiiformes Ophidiidae Lepophidium robustum 1 200 200 200

Actinopterygii Ophidiiformes Ophidiidae Lepophidium zophochir 3 210 210 210

Actinopterygii Ophidiiformes Ophidiidae Luciobrotula corethromycter 16 540 821 612

Actinopterygii Ophidiiformes Ophidiidae Monomitopus agassizii 47 365 1,251 528

Actinopterygii Ophidiiformes Ophidiidae Neobythites gilli 78 200 500 320

Actinopterygii Ophidiiformes Ophidiidae Neobythites marginatus 68 205 670 305

Actinopterygii Ophidiiformes Ophidiidae Neobythites monocellatus 4 229 334 257

Actinopterygii Ophidiiformes Ophidiidae Neobythites ocellatus 19 192 366 234

Actinopterygii Ophidiiformes Ophidiidae Penopus microphthalmus 1 1,006 1,006 1,006

Actinopterygii Ophidiiformes Ophidiidae Xyelacyba myersi 5 1,251 1,440 1,301

Actinopterygii Osmeriformes Alepocephalidae Alepocephalus australis 8 1,097 1,317 1,146

Actinopterygii Osmeriformes Alepocephalidae Bathytroctes microlepis 2 1,097 1,097 1,097

Actinopterygii Osmeriformes Alepocephalidae Conocara macropterum 10 821 1,463 1,125

Actinopterygii Osmeriformes Alepocephalidae Leptoderma macrops 1 690 690 690

Actinopterygii Osmeriformes Alepocephalidae Narcetes stomias 7 558 1,829 1,509

Actinopterygii Osmeriformes Alepocephalidae Rouleina attrita 4 686 1,271 828

Actinopterygii Osmeriformes Alepocephalidae Talismania antillarum 2 457 457 457

Actinopterygii Osmeriformes Alepocephalidae Talismania homoptera 5 515 640 593

Actinopterygii Osmeriformes Alepocephalidae Xenodermichthys copei 23 360 640 433

Actinopterygii Osmeriformes Argentinidae Argentina brucei 33 198 439 266

Actinopterygii Osmeriformes Argentinidae Argentina striata 72 200 500 266

Actinopterygii Osmeriformes Bathylagidae Dolicholagus longirostris 6 445 857 699

Actinopterygii Osmeriformes Microstomatidae Xenophthalmichthys danae 7 205 505 323

Actinopterygii Osmeriformes Opisthoproctidae Opisthoproctus soleatus 1 374 374 374

Actinopterygii Perciformes Acropomatidae Caraibops trispinosus 15 192 523 220

Actinopterygii Perciformes Acropomatidae Synagrops bellus 122 192 810 352

Actinopterygii Perciformes Acropomatidae Verilus atlanticus 10 192 496 250

Actinopterygii Perciformes Acropomatidae Verilus pseudomicrolepis 12 247 308 262

Actinopterygii Perciformes Acropomatidae Verilus sordidus 2 201 201 201

Actinopterygii Perciformes Ariommatidae Ariomma bondi 3 351 428 383

Actinopterygii Perciformes Ariommatidae Ariomma melanum 2 298 366 355

Actinopterygii Perciformes Bathyclupeidae Bathyclupea argentea 24 402 732 448

Actinopterygii Perciformes Bathyclupeidae Bathyclupea schroederi 16 205 810 387

Actinopterygii Perciformes Callionymidae Synchiropus agassizii 8 274 352 306

Actinopterygii Perciformes Callionymidae Synchiropus dagmarae 1 229 229 229

Actinopterygii Perciformes Caproidae Antigonia capros 37 198 505 240

Actinopterygii Perciformes Caproidae Antigonia combatia 51 192 520 243

Actinopterygii Perciformes Carangidae Decapterus macarellus 3 198 235 226

Actinopterygii Perciformes Carangidae Decapterus tabl 21 200 507 229

Actinopterygii Perciformes Carangidae Selar crumenophthalmus 7 246 2,195 575

Actinopterygii Perciformes Carangidae Selene brownii 3 200 504 403

Actinopterygii Perciformes Carangidae Trachurus lathami 2 207 207 207

Actinopterygii Perciformes Emmelichthyidae Erythrocles monodi 1 201 201 201

Actinopterygii Perciformes Epigonidae Epigonus macrops 13 200 914 613

Actinopterygii Perciformes Epigonidae Epigonus occidentalis 28 366 823 434

Actinopterygii Perciformes Epigonidae Epigonus pandionis 55 223 720 343

Actinopterygii Perciformes Gempylidae Diplospinus multistriatus 8 200 445 298

Actinopterygii Perciformes Gempylidae Lepidocybium flavobrunneum 2 1,251 1,271 1,261

Class Order Family Species Records Min depth Max depth Mean depth

Appendix 1. Continued.

240 MARINE AND FISHERY SCIENCES 33 (2): 227-246 (2020)

Actinopterygii Perciformes Gempylidae Nealotus tripes 5 205 360 262

Actinopterygii Perciformes Gempylidae Neoepinnula americana 13 229 333 249

Actinopterygii Perciformes Gempylidae Promethichthys prometheus 17 298 807 386

Actinopterygii Perciformes Gempylidae Ruvettus pretiosus 3 396 558 461

Actinopterygii Perciformes Haemulidae Haemulon aurolineatum 2 286 286 286

Actinopterygii Perciformes Haemulidae Haemulon boschmae 2 286 295 291

Actinopterygii Perciformes Lutjanidae Lutjanus vivanus 1 324 324 324

Actinopterygii Perciformes Lutjanidae Pristipomoides aquilonaris 1 198 198 198

Actinopterygii Perciformes Lutjanidae Pristipomoides macrophthalmus 3 201 280 220

Actinopterygii Perciformes Mullidae Upeneus parvus 7 219 792 417

Actinopterygii Perciformes Opistognathidae Lonchopisthus lemur 32 200 300 242

Actinopterygii Perciformes Opistognathidae Lonchopisthus micrognathus 2 265 265 265

Actinopterygii Perciformes Percophidae Bembrops anatirostris 41 198 500 314

Actinopterygii Perciformes Percophidae Bembrops gobioides 6 360 540 439

Actinopterygii Perciformes Percophidae Bembrops magnisquamis 1 540 540 540

Actinopterygii Perciformes Percophidae Bembrops ocellatus 22 223 670 410

Actinopterygii Perciformes Percophidae Bembrops quadrisella 3 290 384 380

Actinopterygii Perciformes Priacanthidae Heteropriacanthus cruentatus 4 333 432 406

Actinopterygii Perciformes Sciaenidae Protosciaena bathytatos 12 240 512 340

Actinopterygii Perciformes Sciaenidae Protosciaena trewavasae 8 191 201 199

Actinopterygii Perciformes Scombrolabracidae Scombrolabrax heterolepis 1 396 396 396

Actinopterygii Perciformes Serranidae Baldwinella aureorubens 25 198 351 249

Actinopterygii Perciformes Serranidae Baldwinella eos 1 316 316 316

Actinopterygii Perciformes Serranidae Bathyanthias cubensis 7 192 280 209

Actinopterygii Perciformes Serranidae Bathyanthias mexicanus 1 300 300 300

Actinopterygii Perciformes Serranidae Bullisichthys caribbaeus 1 219 219 219

Actinopterygii Perciformes Serranidae Hyporthodus flavolimbatus 1 270 270 270

Actinopterygii Perciformes Serranidae Hyporthodus nigritus 2 200 333 267

Actinopterygii Perciformes Serranidae Hyporthodus niveatus 1 316 316 316

Actinopterygii Perciformes Serranidae Plectranthias garrupellus 2 200 200 200

Actinopterygii Perciformes Serranidae Pronotogrammus martinicensis 1 219 219 219

Actinopterygii Perciformes Serranidae Serranus atrobranchus 3 300 370 323

Actinopterygii Perciformes Sparidae Pagrus pagrus 1 207 207 207

Actinopterygii Perciformes Symphysanodontidae Symphysanodon berryi 3 200 280 216

Actinopterygii Perciformes Synagropidae Parascombrops spinosus 28 192 750 243

Actinopterygii Perciformes Trichiuridae Benthodesmus simonyi 15 223 659 302

Actinopterygii Perciformes Trichiuridae Benthodesmus tenuis 45 223 732 371

Actinopterygii Perciformes Trichiuridae Lepidopus altifrons 8 320 366 334

Actinopterygii Perciformes Trichiuridae Lepidopus caudatus 6 240 404 293

Actinopterygii Perciformes Trichiuridae Trichiurus lepturus 3 220 250 239

Actinopterygii Pleuronectiformes Bothidae Chascanopsetta lugubris 7 366 576 417

Actinopterygii Pleuronectiformes Bothidae Monolene atrimana 2 223 373 298

Actinopterygii Pleuronectiformes Bothidae Monolene megalepis 10 206 505 235

Actinopterygii Pleuronectiformes Bothidae Trichopsetta caribbaea 7 192 300 209

Actinopterygii Pleuronectiformes Bothidae Trichopsetta ventralis 2 198 252 225

Actinopterygii Pleuronectiformes Cynoglossidae Symphurus diomedeanus 2 290 316 303

Actinopterygii Pleuronectiformes Cynoglossidae Symphurus ginsburgi 11 296 491 348

Actinopterygii Pleuronectiformes Cynoglossidae Symphurus hernandezi 12 204 300 228

Actinopterygii Pleuronectiformes Cynoglossidae Symphurus marginatus 62 265 698 355

Actinopterygii Pleuronectiformes Cynoglossidae Symphurus piger 17 203 750 276

Actinopterygii Pleuronectiformes Cynoglossidae Symphurus stigmosus 1 274 274 274

Class Order Family Species Records Min depth Max depth Mean depth

Appendix 1. Continued.

241

GARCÍA AND GAMBOA: DEEP DEMERSAL FISH DIVERSITY IN THE COLOMBIAN CARIBBEAN

Actinopterygii Pleuronectiformes Paralichthyidae Ancylopsetta cycloidea 9 192 269 207

Actinopterygii Pleuronectiformes Paralichthyidae Citharichthys cornutus 5 192 300 207

Actinopterygii Pleuronectiformes Pleuronectidae Poecilopsetta beanii 4 223 333 283

Actinopterygii Pleuronectiformes Pleuronectidae Poecilopsetta inermis 113 229 750 292

Actinopterygii Polymixiiformes Polymixiidae Polymixia lowei 72 200 940 282

Actinopterygii Polymixiiformes Polymixiidae Polymixia nobilis 7 366 512 412

Actinopterygii Scorpaeniformes Peristediidae Peristedion ecuadorense 34 223 810 374

Actinopterygii Scorpaeniformes Peristediidae Peristedion gracile 27 191 940 270

Actinopterygii Scorpaeniformes Peristediidae Peristedion greyae 53 265 803 334

Actinopterygii Scorpaeniformes Peristediidae Peristedion imberbe 1 219 219 219

Actinopterygii Scorpaeniformes Peristediidae Peristedion longispatha 38 223 766 414

Actinopterygii Scorpaeniformes Peristediidae Peristedion miniatum 42 229 720 291

Actinopterygii Scorpaeniformes Peristediidae Peristedion truncatum 18 223 731 348

Actinopterygii Scorpaeniformes Scorpaenidae Neomerinthe beanorum 3 200 300 260

Actinopterygii Scorpaeniformes Scorpaenidae Phenacoscorpius nebris 2 300 300 300

Actinopterygii Scorpaeniformes Scorpaenidae Pontinus longispinis 65 200 491 254

Actinopterygii Scorpaeniformes Scorpaenidae Pontinus nematophthalmus 9 200 698 288

Actinopterygii Scorpaeniformes Scorpaenidae Pontinus rathbuni 6 200 324 227

Actinopterygii Scorpaeniformes Sebastidae Helicolenus dactylopterus 3 293 662 366

Actinopterygii Scorpaeniformes Sebastidae Trachyscorpia cristulata 1 540 540 540

Actinopterygii Scorpaeniformes Setarchidae Ectreposebastes imus 5 400 732 550

Actinopterygii Scorpaeniformes Setarchidae Setarches guentheri 53 200 510 273

Actinopterygii Scorpaeniformes Triglidae Bellator brachychir 3 200 300 209

Actinopterygii Scorpaeniformes Triglidae Bellator egretta 2 200 203 201

Actinopterygii Scorpaeniformes Triglidae Prionotus beanii 1 219 219 219

Actinopterygii Scorpaeniformes Triglidae Prionotus stearnsi 10 213 351 273

Actinopterygii Stephanoberyciformes Gibberichthyidae Gibberichthys pumilus 3 720 732 728

Actinopterygii Stephanoberyciformes Stephanoberycidae Stephanoberyx monae 10 395 1,143 786

Actinopterygii Stomiiformes Gonostomatidae Gonostoma atlanticum 16 200 792 255

Actinopterygii Stomiiformes Gonostomatidae Sigmops elongatus 72 200 2341 452

Actinopterygii Stomiiformes Gonostomatidae Triplophos hemingi 28 300 732 424

Actinopterygii Stomiiformes Phosichthyidae Ichthyococcus ovatus 5 210 755 288

Actinopterygii Stomiiformes Phosichthyidae Pollichthys mauli 38 191 857 341

Actinopterygii Stomiiformes Phosichthyidae Polymetme corythaeola 19 275 857 487

Actinopterygii Stomiiformes Phosichthyidae Polymetme thaeocoryla 13 223 274 272

Actinopterygii Stomiiformes Phosichthyidae Yarrella blackfordi 26 283 1,143 625

Actinopterygii Stomiiformes Sternoptychidae Argyripnus atlanticus 1 260 260 260

Actinopterygii Stomiiformes Sternoptychidae Argyropelecus aculeatus 48 205 857 479

Actinopterygii Stomiiformes Sternoptychidae Argyropelecus gigas 1 540 540 540

Actinopterygii Stomiiformes Sternoptychidae Maurolicus muelleri 2 234 493 364

Actinopterygii Stomiiformes Sternoptychidae Polyipnus asteroides 50 205 567 338

Actinopterygii Stomiiformes Sternoptychidae Polyipnus clarus 4 205 265 259

Actinopterygii Stomiiformes Sternoptychidae Polyipnus laternatus 2 366 549 412

Actinopterygii Stomiiformes Sternoptychidae Sternoptyx diaphana 43 197 4,151 585

Actinopterygii Stomiiformes Sternoptychidae Sternoptyx pseudobscura 3 445 857 585

Actinopterygii Stomiiformes Stomiidae Aristostomias grimaldii 2 640 640 640

Actinopterygii Stomiiformes Stomiidae Aristostomias xenostoma 3 600 792 708

Actinopterygii Stomiiformes Stomiidae Astronesthes macropogon 9 225 1,710 412

Actinopterygii Stomiiformes Stomiidae Chauliodus sloani 125 191 4,151 433

Actinopterygii Stomiiformes Stomiidae Eustomias schmidti 2 450 857 586

Actinopterygii Stomiiformes Stomiidae Heterophotus ophistoma 3 640 732 671

Class Order Family Species Records Min depth Max depth Mean depth

Appendix 1. Continued.

242 MARINE AND FISHERY SCIENCES 33 (2): 227-246 (2020)

Actinopterygii Stomiiformes Stomiidae Malacosteus niger 10 400 732 502

Actinopterygii Stomiiformes Stomiidae Melanostomias macrophotus 1 404 404 404

Actinopterygii Stomiiformes Stomiidae Stomias affinis 32 205 857 420

Actinopterygii Stomiiformes Stomiidae Stomias longibarbatus 3 225 755 358

Actinopterygii Tetraodontiformes Triacanthodidae Hollardia hollardi 6 329 558 350

Actinopterygii Tetraodontiformes Triacanthodidae Parahollardia lineata 1 369 369 369

Actinopterygii Tetraodontiformes Triacanthodidae Parahollardia schmidti 5 200 360 222

Actinopterygii Zeiformes Grammicolepididae Grammicolepis brachiusculus 8 324 662 442

Actinopterygii Zeiformes Grammicolepididae Xenolepidichthys dalgleishi 25 200 439 245

Actinopterygii Zeiformes Parazenidae Cyttopsis rosea 95 223 732 324

Actinopterygii Zeiformes Parazenidae Parazen pacificus 2 274 352 281

Actinopterygii Zeiformes Zeidae Zenopsis conchifer 2 270 298 284

Actinopterygii Zeiformes Zenionidae Zenion hololepis 72 200 540 385

Elasmobranchii Carcharhiniformes Carcharhinidae Galeocerdo cuvier 1 365 365 365

Elasmobranchii Carcharhiniformes Scyliorhinidae Apristurus canutus 2 530 777 592

Elasmobranchii Carcharhiniformes Scyliorhinidae Apristurus parvipinnis 2 719 719 719

Elasmobranchii Carcharhiniformes Scyliorhinidae Apristurus riveri 3 860 960 889

Elasmobranchii Carcharhiniformes Scyliorhinidae Galeus arae 4 256 512 374

Elasmobranchii Carcharhiniformes Scyliorhinidae Galeus cadenati 4 256 512 436

Elasmobranchii Carcharhiniformes Scyliorhinidae Schroederichthys maculatus 1 274 274 274

Elasmobranchii Carcharhiniformes Scyliorhinidae Scyliorhinus boa 19 223 540 298

Elasmobranchii Carcharhiniformes Scyliorhinidae Scyliorhinus hesperius 9 290 549 341

Elasmobranchii Carcharhiniformes Triakidae Mustelus canis 2 298 316 307

Elasmobranchii Rajiformes Anacanthobatidae Schroederobatis americana 51 307 803 475

Elasmobranchii Rajiformes Rajidae Breviraja nigriventralis 18 457 732 512

Elasmobranchii Rajiformes Rajidae Breviraja spinosa 4 540 612 575

Elasmobranchii Rajiformes Rajidae Cruriraja rugosa 7 365 732 449

Elasmobranchii Rajiformes Rajidae Dactylobatus clarkii 18 366 512 421

Elasmobranchii Rajiformes Rajidae Dipturus bullisi 9 201 334 302

Elasmobranchii Rajiformes Rajidae Dipturus garricki 3 283 307 301

Elasmobranchii Rajiformes Rajidae Dipturus oregoni 1 396 396 396

Elasmobranchii Rajiformes Rajidae Dipturus teevani 3 240 576 407

Elasmobranchii Rajiformes Rajidae Fenestraja plutonia 1 428 428 428

Elasmobranchii Rajiformes Rajidae Fenestraja sinusmexicanus 2 485 485 485

Elasmobranchii Rajiformes Rajidae Gurgesiella atlantica 34 223 698 498

Elasmobranchii Squaliformes Centrophoridae Centrophorus granulosus 4 200 732 333

Elasmobranchii Squaliformes Centrophoridae Centrophorus squamosus 1 670 670 670

Elasmobranchii Squaliformes Dalatiidae Isistius brasiliensis 1 621 621 621

Elasmobranchii Squaliformes Etmopteridae Etmopterus bullisi 1 274 274 274

Elasmobranchii Squaliformes Etmopteridae Etmopterus carteri 8 283 343 292

Elasmobranchii Squaliformes Etmopteridae Etmopterus hillianus 6 180 540 351

Elasmobranchii Squaliformes Etmopteridae Etmopterus perryi 22 283 375 297

Elasmobranchii Squaliformes Etmopteridae Etmopterus pusillus 1 288 288 288

Elasmobranchii Squaliformes Etmopteridae Etmopterus schultzi 43 269 823 422

Elasmobranchii Squaliformes Etmopteridae Etmopterus virens 25 288 503 331

Elasmobranchii Squaliformes Somniosidae Somniosus cf microcephalus 1 200 200 200

Elasmobranchii Squaliformes Squalidae Squalus cubensis 4 198 274 263

Elasmobranchii Squatiniformes Squatinidae Squatina david 2 198 305 252

Elasmobranchii Torpediniformes Torpedinidae Tetronarce nobiliana 6 292 369 329

Holocephali Chimaeriformes Chimaeridae Chimaera cubana 1 234 234 234

Holocephali Chimaeriformes Chimaeridae Hydrolagus alberti 27 223 1,143 477

Class Order Family Species Records Min depth Max depth Mean depth

Appendix 1. Continued.

APPENDIX 2

Species excluded from data base because records

fall outside the Colombian Caribbean Exclusive

Economic Zone.

243

GARCÍA AND GAMBOA: DEEP DEMERSAL FISH DIVERSITY IN THE COLOMBIAN CARIBBEAN

Class Order Family Species Records Min depth Max depth Mean depth

Class Order Family Species

Actinopterygii Anguilliformes Congridae Acromycter atlanticus

Actinopterygii Anguilliformes Congridae Bathycongrus thysanochilus

Actinopterygii Anguilliformes Congridae Rhynchoconger gracilior

Actinopterygii Anguilliformes Nettastomatidae Hoplunnis similis

Actinopterygii Aulopiformes Giganturidae Gigantura chuni

Actinopterygii Aulopiformes Ipnopidae Bathytyphlops sewelli

Actinopterygii Aulopiformes Paralepididae Lestrolepis intermedia

Actinopterygii Aulopiformes Paralepididae Stemonosudis rothschildi

Actinopterygii Batrachoidiformes Batrachoididae Porichthys bathoiketes

Actinopterygii Ophidiiformes Ophidiidae Lepophidium entomelan

Actinopterygii Ophidiiformes Ophidiidae Lepophidium marmoratum

Actinopterygii Ophidiiformes Ophidiidae Lepophidium staurophor

Actinopterygii Osmeriformes Argentinidae Argentina stewarti

Actinopterygii Osmeriformes Argentinidae Glossanodon pygmaeus

Actinopterygii Perciformes Apogonidae Paroncheilus affinis

Actinopterygii Perciformes Labridae Decodon puellaris

Actinopterygii Perciformes Lutjanidae Rhomboplites aurorubens

Actinopterygii Perciformes Percophidae Bembrops macromma

Actinopterygii Perciformes Serranidae Serranus phoebe

Actinopterygii Perciformes Uranoscopidae Kathetostoma cubana

Actinopterygii Pleuronectiformes Paralichthyidae Ancylopsetta microctenus

Actinopterygii Pleuronectiformes Paralichthyidae Citharichthys dinoceros

Actinopterygii Stomiiformes Sternoptychidae Sonoda megalophthalma

Actinopterygii Stomiiformes Stomiidae Echiostoma barbatum

Actinopterygii Stomiiformes Stomiidae Eustomias acinosus

Elasmobranchii Rajiformes Rajidae Breviraja mouldi

Elasmobranchii Rajiformes Rajidae Fenestraja ishiyamai

Elasmobranchii Rajiformes Rajidae Leucoraja garmani

Elasmobranchii Squaliformes Etmopteridae Etmopterus robinsi

Elasmobranchii Squaliformes Squalidae Squalus mitsukurii

Myxini Myxiniformes Myxinidae Eptatretus caribbeaus

Myxini Myxiniformes Myxinidae Eptatretus multidens

Holocephali Chimaeriformes Chimaeridae Hydrolagus mirabilis 3 720 1,296 995

Holocephali Chimaeriformes Rhinochimaeridae Neoharriotta carri 14 288 485 329

Holocephali Chimaeriformes Rhinochimaeridae Rhinochimaera atlantica 2 914 917 916

Myxini Myxiniformes Myxinidae Eptatretus aceroi 2 705 705 705

Myxini Myxiniformes Myxinidae Eptatretus ancon 8 476 670 513

Myxini Myxiniformes Myxinidae Eptatretus wayuu 5 300 303 301

Myxini Myxiniformes Myxinidae Myxine mccoskeri 72 269 801 446

Myxini Myxiniformes Myxinidae Myxine robinsorum 2 783 821 793

APPENDIX 3

Deep demersal species (> 200 m depth) recor-

ded inside the Parque Nacional Natural National

Natural Park Corales de Profundidad in the

Colombian Caribbean.

244 MARINE AND FISHERY SCIENCES 33 (2): 227-246 (2020)

Class Order Family Genus Species

Actinopterygii Anguilliformes Colocongridae Coloconger Coloconger meadi

Actinopterygii Anguilliformes Congridae Bathycongrus Bathycongrus bullisi

Actinopterygii Anguilliformes Congridae Pseudophichthys Pseudophichthys splendens

Actinopterygii Anguilliformes Congridae Xenomystax Xenomystax congroides

Actinopterygii Aulopiformes Chlorophthalmidae Chlorophthalmus Chlorophthalmus agassizi

Actinopterygii Aulopiformes Chlorophthalmidae Parasudis Parasudis truculenta

Actinopterygii Aulopiformes Ipnopidae Bathypterois Bathypterois bigelowi

Actinopterygii Aulopiformes Synodontidae Saurida Saurida brasiliensis

Actinopterygii Beryciformes Trachichthyidae Hoplostethus Hoplostethus occidentalis

Actinopterygii Gadiformes Bregmacerotidae Bregmaceros Bregmaceros atlanticus

Actinopterygii Gadiformes Macrouridae Bathygadus Bathygadus macrops

Actinopterygii Gadiformes Macrouridae Coelorinchus Coelorinchus caelorhincus

Actinopterygii Gadiformes Macrouridae Coelorinchus Coelorinchus caribbaeus

Actinopterygii Gadiformes Macrouridae Gadomus Gadomus arcuatus

Actinopterygii Gadiformes Macrouridae Hymenocephalus Hymenocephalus billsam

Actinopterygii Gadiformes Macrouridae Hymenocephalus Hymenocephalus italicus

Actinopterygii Gadiformes Macrouridae Malacocephalus Malacocephalus laevis

Actinopterygii Gadiformes Macrouridae Malacocephalus Malacocephalus occidentalis

Actinopterygii Gadiformes Macrouridae Nezumia Nezumia aequalis

Actinopterygii Gadiformes Macrouridae Ventrifossa Ventrifossa macropogon

Actinopterygii Gadiformes Merlucciidae Steindachneria Steindachneria argentea

Actinopterygii Gadiformes Moridae Gadella Gadella imberbis

Actinopterygii Gadiformes Moridae Laemonema Laemonema goodebeanorum

Actinopterygii Gadiformes Moridae Physiculus Physiculus fulvus

Actinopterygii Lophiiformes Chaunacidae Chaunax Chaunax pictus

Actinopterygii Lophiiformes Chaunacidae Chaunax Chaunax suttkusi

Actinopterygii Lophiiformes Lophiidae Lophiodes Lophiodes monodi

Actinopterygii Lophiiformes Ogcocephalidae Dibranchus Dibranchus atlanticus

Actinopterygii Lophiiformes Ogcocephalidae Halieutichthys Halieutichthys aculeatus

Actinopterygii Lophiiformes Ogcocephalidae Malthopsis Malthopsis gnoma

Actinopterygii Lophiiformes Ogcocephalidae Ogcocephalus Ogcocephalus declivirostris

Actinopterygii Myctophiformes Myctophidae Dasyscopelus Dasyscopelus selenops

Actinopterygii Myctophiformes Myctophidae Diaphus Diaphus effulgens

Actinopterygii Myctophiformes Myctophidae Diaphus Diaphus garmani

Actinopterygii Myctophiformes Myctophidae Diaphus Diaphus lucidus

Actinopterygii Myctophiformes Neoscopelidae Neoscopelus Neoscopelus macrolepidotus

Actinopterygii Myctophiformes Neoscopelidae Neoscopelus Neoscopelus microchir

Actinopterygii Notacanthiformes Halosauridae Halosaurus Halosaurus guentheri

Actinopterygii Notacanthiformes Halosauridae Halosaurus Halosaurus ovenii

Actinopterygii Notacanthiformes Notacanthidae Notacanthus Notacanthus chemnitzii

Actinopterygii Ophidiiformes Bythitidae Calamopteryx Calamopteryx robinsorum

Actinopterygii Ophidiiformes Bythitidae Diplacanthopoma Diplacanthopoma brachysoma

Actinopterygii Ophidiiformes Ophidiidae Benthocometes Benthocometes robustus

Actinopterygii Ophidiiformes Ophidiidae Monomitopus Monomitopus agassizii

Actinopterygii Ophidiiformes Ophidiidae Neobythites Neobythites gilli

Actinopterygii Ophidiiformes Ophidiidae Neobythites Neobythites marginatus

Actinopterygii Osmeriformes Argentinidae Argentina Argentina striata

Actinopterygii Perciformes Acropomatidae Caraibops Caraibops trispinosus

Appendix 3. Continued.

245

GARCÍA AND GAMBOA: DEEP DEMERSAL FISH DIVERSITY IN THE COLOMBIAN CARIBBEAN

Actinopterygii Perciformes Acropomatidae Synagrops Synagrops bellus

Actinopterygii Perciformes Acropomatidae Verilus Verilus atlanticus

Actinopterygii Perciformes Acropomatidae Verilus Verilus pseudomicrolepis

Actinopterygii Perciformes Bathyclupeidae Bathyclupea Bathyclupea argentea

Actinopterygii Perciformes Bathyclupeidae Bathyclupea Bathyclupea schroederi

Actinopterygii Perciformes Caproidae Antigonia Antigonia capros

Actinopterygii Perciformes Caproidae Antigonia Antigonia combatia

Actinopterygii Perciformes Carangidae Decapterus Decapterus tabl

Actinopterygii Perciformes Carangidae Selar Selar crumenophthalmus

Actinopterygii Perciformes Epigonidae Epigonus Epigonus macrops

Actinopterygii Perciformes Epigonidae Epigonus Epigonus occidentalis

Actinopterygii Perciformes Epigonidae Epigonus Epigonus pandionis

Actinopterygii Perciformes Lutjanidae Pristipomoides Pristipomoides macrophthalmus

Actinopterygii Perciformes Opistognathidae Lonchopisthus Lonchopisthus lemur

Actinopterygii Perciformes Percophidae Bembrops Bembrops anatirostris

Actinopterygii Perciformes Percophidae Bembrops Bembrops gobioides

Actinopterygii Perciformes Percophidae Bembrops Bembrops ocellatus

Actinopterygii Perciformes Percophidae Bembrops Bembrops quadrisella

Actinopterygii Perciformes Serranidae Baldwinella Baldwinella aureorubens

Actinopterygii Perciformes Serranidae Bathyanthias Bathyanthias cubensis

Actinopterygii Perciformes Serranidae Hyporthodus Hyporthodus nigritus

Actinopterygii Perciformes Symphysanodontidae Symphysanodon Symphysanodon berryi

Actinopterygii Perciformes Trichiuridae Benthodesmus Benthodesmus simonyi

Actinopterygii Perciformes Trichiuridae Benthodesmus Benthodesmus tenuis

Actinopterygii Pleuronectiformes Bothidae Monolene Monolene atrimana

Actinopterygii Pleuronectiformes Bothidae Monolene Monolene megalepis

Actinopterygii Pleuronectiformes Cynoglossidae Symphurus Symphurus marginatus

Actinopterygii Pleuronectiformes Paralichthyidae Ancylopsetta Ancylopsetta cycloidea

Actinopterygii Pleuronectiformes Pleuronectidae Poecilopsetta Poecilopsetta beanii

Actinopterygii Pleuronectiformes Pleuronectidae Poecilopsetta Poecilopsetta inermis

Actinopterygii Polymixiiformes Polymixiidae Polymixia Polymixia lowei

Actinopterygii Scorpaeniformes Peristediidae Peristedion Peristedion ecuadorense

Actinopterygii Scorpaeniformes Peristediidae Peristedion Peristedion gracile

Actinopterygii Scorpaeniformes Peristediidae Peristedion Peristedion greyae

Actinopterygii Scorpaeniformes Peristediidae Peristedion Peristedion longispatha

Actinopterygii Scorpaeniformes Peristediidae Peristedion Peristedion truncatum

Actinopterygii Scorpaeniformes Scorpaenidae Neomerinthe Neomerinthe beanorum

Actinopterygii Scorpaeniformes Scorpaenidae Pontinus Pontinus nematophthalmus

Actinopterygii Scorpaeniformes Setarchidae Setarches Setarches guentheri

Actinopterygii Stomiiformes Phosichthyidae Polymetme Polymetme corythaeola

Actinopterygii Stomiiformes Phosichthyidae Polymetme Polymetme thaeocoryla

Actinopterygii Stomiiformes Sternoptychidae Argyripnus Argyripnus atlanticus

Actinopterygii Stomiiformes Sternoptychidae Argyropelecus Argyropelecus aculeatus

Actinopterygii Stomiiformes Sternoptychidae Argyropelecus Argyropelecus gigas

Actinopterygii Stomiiformes Sternoptychidae Maurolicus Maurolicus muelleri

Actinopterygii Stomiiformes Sternoptychidae Polyipnus Polyipnus asteroides

Actinopterygii Stomiiformes Sternoptychidae Polyipnus Polyipnus laternatus

Actinopterygii Stomiiformes Stomiidae Chauliodus Chauliodus sloani

Actinopterygii Tetraodontiformes Triacanthodidae Parahollardia Parahollardia schmidti

Actinopterygii Zeiformes Grammicolepididae Xenolepidichthys Xenolepidichthys dalgleishi

Actinopterygii Zeiformes Parazenidae Cyttopsis Cyttopsis rosea

Class Order Family Genus Species

Appendix 3. Continued.

246 MARINE AND FISHERY SCIENCES 33 (2): 227-246 (2020)

Actinopterygii Zeiformes Zenionidae Zenion Zenion hololepis

Elasmobranchii Carcharhiniformes Scyliorhinidae Scyliorhinus Scyliorhinus boa

Elasmobranchii Rajiformes Rajidae Breviraja Breviraja spinosa

Elasmobranchii Rajiformes Rajidae Dipturus Dipturus bullisi

Elasmobranchii Rajiformes Rajidae Gurgesiella Gurgesiella atlantica

Holocephali Chimaeriformes Chimaeridae Hydrolagus Hydrolagus alberti

Myxini Myxiniformes Myxinidae Myxine Myxine mccoskeri

Class Order Family Genus Species