ANARTIA, 25 (2013 “2015”): 32 - 46

ISSN: 1315-642X

The Presence of Long-Beaked

Common Dolphins (Delphinus spp.)

off Central-Western Venezuela

Jaime Bolaños-Jiménez

1,2

*, Graciela Castro-Pérez

Olga Herrera-Trujillo

, Lenin Oviedo

5,6

, Daniel Palacios

Leonardo Sánchez-Criollo

8,9

, María F. Puerto

Leonardo Sifontes

, María G. Silva-Hernández

and Auristela Villarroel-Marín

Asociación Civil Sea Vida (Sea Vida), A.P. 162, Cagua, Estado Aragua,

Venezuela 2122.

Instituto de Ciencias Marinas y Pesquerías (ICIMAP), Universidad

Veracruzana, Veracruz, México.

Universidad de Carabobo, Facultad de Ciencias y Tecnología (FACYT),

Naguanagua, Estado Carabobo, Venezuela.

Mares Venezuela, Caracas, Venezuela.

Proyecto Delphinus, Isla de Margarita, Venezuela.

Laboratorio de Ecología de Pinnípedos, Instituto Politécnico Nacional-

Centro Interdisciplinario de Ciencias Marinas, La Paz, BCS, México.

Marine Mammal Institute, Oregon State University, Hatfield Marine Science

Center. 2030 SE Marine Science Drive, Newport, OR 97365, USA.

Laboratorio de Paleontología, Instituto Venezolano de Investigaciones Científicas

(IVIC), Altos de Pipe, Estado Miranda, Venezuela.

Centro de Investigación de Cetáceos, Capítulo Zulia, Maracaibo, Venezuela.

*Corresponding author: bolanos.jimenez@gmail.com

Abstract

The common dolphin (genus Delphinus) is regularly found in re-

gions with upwelling-modified waters. The “Venezuelan stock” of

the common dolphin is documented to occur in north-eastern Vene-

zuelan waters, particularly in the “Eastern Upwelling” eco-region.

Records of Delphinus for other eco-regions of Venezuela are extremely

sparse. Information was compiled from three independent cetacean

studies conducted during 2009-2011 in the states of Aragua (Central

Coast eco-region), Miranda (Unare-Píritu eco-region) and Zulia

(Gulf of Venezuela eco-region), respectively. Fourteen records of

Delphinus were compiled for this study: 10 sightings (71,5%) off the

coast of Aragua State; two (14,3%) off Miranda State; as well as one

sighting (7,1%) and one stranding (7,1%) in the Gulf of Venezuela.

Records in the states of Miranda and Zulia were related to

upwelling-modified waters. We found no evidence of upwelling in

waters off Aragua State (71,5% of the records).

Keywords: Common dolphin, Delphinus, distribution, Caribbean

Sea, Venezuela, coastal upwelling.

Presencia de delfín común de rostro largo

(Delphinus spp.) al norte y oeste de la costa

continental de Venezuela

Resumen

El delfín común (género Delphinus) es frecuentemente encon-

trado en zonas de surgencias. El “Stock Venezolano” del género Del-

phinus está ampliamente documentado para las aguas del nor-oriente

de Venezuela, particularmente en la eco-región “Surgencia Orien

tal”. Registros de esta especie en otras regiones venezolanas son ex

tremadamente escasos. En el presente trabajo se recolectó informa

ción proveniente de tres diferentes proyectos de campo efectuados en

los estados Aragua (Eco-región Costa Central), Miranda (Eco-re

gión Unare-Píritu) y Zulia (Eco-región Golfo de Venezuela). Se do

cumentaron 14 registros de Delphinus: 10 en Aragua (71,5%), dos en

Miranda (14,3%), así como un avistamiento (7,1%) y un varamiento

(7,1%) en Zulia. Los registros en Miranda y Zulia estuvieron relacio

nados con zonas de surgencia; sin embargo en el presente estudio no

encontramos evidencias de surgencia en la costa de Aragua, donde

ocurrió el 71,5% de los avistamientos.

Palabras clave: Delfín común, Delphinus, distribución, Mar Caribe

Sur, surgencia costera.

Long-beaked common dolphins in Venezuela 33

INTRODUCTION

The common dolphin (genus Delphinus) is known to occur in

three general areas in the western Atlantic: 1) off the east coast of

Canada and the United States (Western North Atlantic stock); 2) off

north-central Venezuela (Venezuelan stock); and 3) southwestern

Atlantic (SWA) (Jefferson et al. 2009, Tavares et al. 2010). For the

SWA, Tavares et al. (2010) identified two stocks off southern Brazil

(Para and Rio de Janeiro states) and one in Argentina (Patagonia re

gion).

As a genus, Delphinus appear to prefer upwelling-modified wa

ters in tropical-temperate areas (Jefferson et al. 2009). Along the

Venezuelan coast, common dolphin distribution seems to mirror the

west–east gradient of coastal upwelling, with major concentrations

along the “Eastern Upwelling” eco-region in the north-eastern coast,

specifically off Araya and Paria Peninsula (Oviedo et al. 2010). The

Eastern Upwelling eco-region is defined by an extended shelf (over

100 km north) and high level of primary productivity due to seasonal

coastal upwelling (Lorenzoni 2000, Miloslavich and Klein 2008)

(Figure 1). Rueda-Roa and Muller-Karger (2013, Fig. 3) identified 13

upwelling foci distributed all along the Venezuelan coast.

The Venezuelan stock has previously been identified as belong-

ing to the long-beaked common dolphin, Delphinus capensis

(Bermúdez-Villapol and Boher-Bentti 2003, Ramírez-Carroz and

González-Fernández 2004, Bolaños-Jiménez et al. 2006) on the basis

of external features as described by Heyning and Perrin (1994). More

recently, Esteves and Oviedo (2007) reviewed the taxonomy of com

mon dolphins in Venezuela on the basis of morphometric features.

These authors found some morphological differences of Venezuelan

specimens as compared to other regions of the world and proposed

that common dolphins of the Venezuelan stock could be a dwarf

morphotype. On the basis of taxonomic uncertainty, Esteves and

Oviedo (2007) cautiously treated these populations as a single taxon,

Delphinus spp.

Up to now, the only confirmed records of common dolphins in

areas outside the Eastern Upwelling and Cariaco Trench eco-regions

were based in a single stranding in the Gulf of Venezuela (Ramírez-

Carroz and González-Fernández 2004) and two sightings off Aragua

Bolaños-Jiménez et al.

ANARTIA, 25 (2013 “2015”): 32 - 46

State (Cobarrubia and Bolaños-Jiménez 2008). Recent records on

the species occurrence in nearby waters of the southern Caribbean:

off Aruba (Luksenburg 2013), off the Colombian Guajira (Palacios

et al. 2012) and Venezuelan waters off southwestern Grenada (Smul

tea et al. 2004, 2013) offer evidence of the presence of common dol

phin associated with upwelling modified waters at regional scale.

The distribution, movements and abundance of highly mobile

marine species are best studied at large scales but most of the research

effort focuses on small areas (Cheney et al. 2013). Development of

collaborative efforts among research teams working in adjacent areas

and integration of multiple data sources are being actively used to

better build up a more real picture of the contemporary ranging pat

terns and abundances of several potential populations of dolphins

inhabiting these areas (Cheney et al. 2013). In this paper, we inte-

grated data sources from three independent studies to present evi-

dences of the presence of free-ranging common dolphins in central-

western Venezuela which confirm the occurrence of the species in re-

gions others than the Eastern Upwelling and Cariaco Trench eco-

region.

METHODS

Information was gathered as part of three field projects con-

ducted separately by independent teams in the States of Aragua

(Central Coast eco-region), Miranda (Unare-Píritu) and Zulia (Gulf

of Venezuela), respectively (see Fig. 1).

Description of eco-regions

Miloslavich and Klein (2008) described 13 Venezuelan marine

eco-regions: 1) Orinoco delta and Atlantic Front, 2) Eastern Upwel

ling, 3) Unare-Píritu, 4) Central coast, 5) Golfo Triste, 6) Tocuyo,

7) Paraguaná, 8) Golfete de Coro, 9) Maracaibo Estuary, 10) Gulf of

Venezuela, 11) Insular, 12) Oceanic eco-regions and 13) Cariaco

Trench. For the interest of this study, we considered five eco-regions

(see Fig. 1):

Eastern Upwelling. From the east of Peninsula de Paria to Mo

chima Bay. Characterized by shallow continental shelf with coarse

sandy bottoms and rocky shores. High primary and secondary pro

Long-beaked common dolphins in Venezuela 35

duction due to seasonal upwelling fronts. This eco-region the islands

of Margarita, Coche, Cubagua, Los Frailes, Los Hermanos and Los

Testigos (Miloslavich and Klein 2008). This eco-region includes the

following upwelling foci: Don Pedro, Las Galdonas, Carúpano,

Manzanillo, Macanao, El Guamache, Golfo de Cariaco and Mo-

chima (Roa-Rueda and Muller-Karger 2013).

Cariaco Trench. It is a depression in the continental shelf, located

in front of Barcelona and the Bay of Mochima, states of Anzoátegui

and Sucre. This eco-region is anoxic below 250m and is character

ized by the existence of chemosynthetic ecosystems. It is a unique

system in the world by permanent anoxic characteristics and high

surface productivity due to the intense seasonal upwelling (Milo

slavich and Klein 2008).

Central Venezuela. As part of the “Central Venezuela” eco-

region, the coast of the State of Aragua is characterized by narrow

continental margin, rocky shores, coarse sandy beaches and high wa

ter transparency (Miloslavich and Klein, 2008). This eco-region does

not include an upwelling focus (Rueda-Roa and Muller-Karger

2013).

Bolaños-Jiménez et al.

ANARTIA, 25 (2013 “2015”): 32 - 46

Figure 1. Relative location of the eco-regions mentioned in this study: Gulf

of Venezuela, Central Coast, Unare-Píritu, Cariaco Trench and Eastern

Upwelling.

Unare-Píritu. Shallow coastal lagoons and sandy beaches, high

sediment loads from the Unare and Tuy rivers. It is characterized by

high secondary production, mostly of crustaceans and bivalves. Up

wellingin Cabo Codera is considered unusual in that it can be present

when there are no other upwelling fronts (Miloslavich and Klein

2008). This eco-region includes the Unare and Cabo Codera upwel

ling foci (Rueda-Roa and Muller-Karger 2013).

Gulf of Venezuela. Characterized by a wide continental shelf

and shallow bottom (50 m). High primary and secondary production

of shrimp due to large upwelling fronts. Bottom communities domi

nated by detritivorous organisms (Miloslavich and Klein 2008). This

eco-region includes the Punta Cardón and Puerto Cumarebo upwel

ling foci (Rueda-Roa and Muller-Karger 2013).

Field effort

The surveys off the coastal regions of Aragua and Miranda

States were designed to obtain baseline information utilizing encoun-

ter information and photo-identification data of coastal dolphin popu-

lations. However, it is important to consider the following aspects:

1) The surveys for Aragua State are a continuation of the effort re-

ported by Bolaños-Jiménez et al. (2007, 2012); and 2) surveys off Mi-

randa State were intended to document oceanic cetaceans in the tran-

sition area between the coastal environment off Miranda State

(Unare-Píritu eco-region) and the western portion of the Cariaco

Trench eco-region. Surveys in the State of Aragua were conducted

from January 22-March 3, May 2-July 1, July 23-August 4, and August

25-September 12, 2009 and from August 19 through September 4

2011 (Bolaños-Jiménez et al. 2012, Castro-Pérez 2009, Sifontes 2013).

Surveys in the State of Miranda wereconducted betweenJanuary-July

2011. Based on the latter, surveys differed in routes and frequency;

however standard sighting information as well as standard photo-ID

data were equally collected and applied in both locations.

Dolphins were identified according to descriptions in Jefferson

et al. (1994) and Heyning and Perrin (1994). For each sighting, the

dolphins were determined to be the common dolphin on the basis of

the long snout and the crisscross color pattern that are diagnostic of

the genus (Jefferson et al. 1994, Heyning and Perrin 1994). The

number of dolphins around the boat was estimated every 5-10 min

Long-beaked common dolphins in Venezuela 37

and the maximum number was recorded at the end of the encounter.

Photographs of the dolphins were taken with SLR cameras (Nikon

D60 and Canon Rebel Xti EOS400, equipped with 18-55 mm and

70-200 mm lenses). According to standard procedures for photo-ID

projects, we attempted to photograph the dorsal fin of every dolphin

in each sighting (Bolaños-Jiménez et al. 2012). The encounter ended

when all of the dolphins had been photographed or the dolphins left

the area. The bottom depth at the sighting location was derived from

the map of the southern Caribbean from BlueChart Americas 2008

using Mapsource™ (Garmin) software.

For the stranding in the State of Zulia, one of us (MFP) had the

opportunity to photograph skeletal remains found on a beach in Cai

mare Chico, Guajira Peninsula (Fig. 4). Craniological features and

tooth counts were used for species identification. The skull could not

be collected.

We generated monthly maps of satellite-measured chloro-

phyll-a by NASA’s MODIS/Aqua sensor for our study area, ob-

tained through NOAA Coastwatch,West Coast Node (http:// www.

coastwatch.noaa.gov/). We used these maps to determine the inten-

sity and extent of upwelling plumes along the coast of central and

western Venezuela during the survey periods.

RESULTS

We recorded 10 sightings (71,4%) off Aragua State, two sight

ings (14,4%) off Miranda State and one sighting (7,1%) and one

stranding (7,1/%) in the State of Zulia. All of the sightings were re

corded in waters over the continental shelf (Table 1, Fig. 2). Depth

ranged between 12-160 m (average 74 ± 48 m, see Table 1).

Nine out of 10 sightings recorded off Aragua between July-

November 2009 were of a small group that ranged from 2-6 individu

als. Because of the presence of one identifiable dolphins -particularly

a presumed mother with her calf- we believe that most of the sight

ings recorded in Aragua during July-September 2009 were of the

same group (Bolaños-Jiménez, pers. obs., Sifontes, pers. obs., Ar

mando Díaz, pers. comm., see Fig. 3A). No good quality pictures are

available for photo-identification purposes and no attempt has been

Bolaños-Jiménez et al.

ANARTIA, 25 (2013 “2015”): 32 - 46

Long-beaked common dolphins in Venezuela 39

Table I. Recent records of the common dolphin in central-western Venezuela (States of Aragua, Miranda and Zulia) during

2009-2011.

# Date Latitude

(N)

Longitude

(W)

Depth

(m)

N Nº calves and

juveniles

State of: Record

1 2008 11,893180 -71,239346 12 3 0 Zulia Sighting

2 26 March 2009 11,12881700 -71,80742100 NA 1 NA Zulia Stranding

3 01 July 2009 10,50111111 -67,75805556 45 60 ? Aragua Sighting

4 27 July 2009 10,51277778 -67,72611111 71 4 1 Aragua Sighting

5 04 Aug 2009 10,50194444 -67,77361111 90 4 1 Aragua Sighting

6 25 Aug 2009 10,52138889 -67,73138889 160 2 1 Aragua Sighting(*)

7 29 Aug 2009 10,50666667 -67,79250000 145 6 1 Aragua Sighting

8 07 Sept 2009 10,52166667 -67,67527758 141 4 1 Aragua Sighting

9 09 Sept 2009 10,51525778 -67,71277778 72 5 1 Aragua Sighting

10 11 Sept 2009 10,50722222 -67,73777778 68 5 1 Aragua Sighting

11 13 Nov 2009 10,50277778 -67,74199444 28 5 1 Aragua Sighting

12 22 July 2011 10,49611111 -65,92083333 58 200 ? Miranda Sighting

13 23 July 2011 10,41333333 -65,85833333 56 300 ? Miranda Sighting

14 02 Sept 2011 10,50250000 -67,73277778 15 1 0 Aragua Sighting(**)

(

): Mixed-species aggregation with approx. 20 Atlantic spotted dolphins (Stenella frontalis).

(

): Mixed-species aggregation with approx. 180 Atlantic spotted dolphins (S. frontalis).

Bolaños-Jiménez et al.

40 ANARTIA, 25 (2013 “2015”): 32 - 46

Figure 2. Sighting (brown triangles) and stranding (red square) records of

Delphinus spp. in central-western Venezuela, 2009-2011. This map was pro-

duced by using Maptools, available at www.seaturtle.org/maptool).

Figure 3. Long-beaked common dolphins (Delphinus spp.) in the states of

Aragua (A and B) and Miranda (C), central Venezuela. Photo credits: Leo-

nardo Sifontes (A), Jaime Bolaños-Jiménez (B) and Lenín Oviedo (C).

Long-beaked common dolphins in Venezuela 41

Figure 4. Skull of a common dolphin stranded in Caimare Chico, State of

Zulia, found on 26 March 2006. Photo: María Fernanda Puerto.

made to start a catalog aimed to target this species in Aragua State,

because of scarcity of encounters.

The specimen collected in Caimare Chico, Guajira Península,

State of Zulia, corresponded to the body condition Code 5 (skeletal

remains) according to the classification of Geraci and Lounsbury

(2005). On the basis of the high tooth count (over 45 per series) and

the presence of deep palatal grooves (see Fig. 4) this specimen was

confirmed to be a common dolphin, genus Delphinus. This determi-

nation was confirmed by specialists (see Acknowledgements).

DISCUSSION

The common dolphin is regularly found in upwelling-modified

waters (Jefferson et al. 2009, Oviedo et al. 2010). In Venezuela, this

species has rarely been seen outside the Eastern Upwelling eco-

region (see Cobarrubia and Bolaños-Jiménez 2008) but, recently,

a sighting has been confirmed in Venezuelan waters southwestern

from Grenada (Smultea et al. 2004, 2013). We found that sightings of

common dolphins in the State of Aragua presented in this paper were

not related to upwelling-modified waters, but satellite imagery re

vealed a high level of pigments, probably because of both chlorophyll

and colored dissolved organic matter (CDOM) (Klein, pers. comm.

2013). The discontinuity of records presented here might be an arti

fact of the few sightings included in the study; nevertheless, it pro

vides grounds to speculate on the possibility of: 1) a more continuous

presence of Delphinus throughout the Venezuelan coast and 2) ex

ploratory movements of dolphins from one upwelling focus to an

other. Because of the opportunistic nature of the records and the

small number of sightings we cannot rule out or confirm any of the

alternatives.

The presence of Delphinus in the State of Miranda (Unare-

Píritu eco-region) was expected, because the area is adjacent to the

Eastern Upwelling eco-region and there exists a previously published

sighting around Cabo Codera (see Jefferson et al. 2009). Our results

are in agreement with Cobarrubia and Bolaños-Jiménez (2008), who

concluded that –based on the paucity of sightings– the common dol-

phin was not a regular inhabitant in the State of Aragua. As for the

nine sightings recorded in Aragua between July-November 2009.

Bolaños-Jiménez et al. (2012) speculated that they could be related to

the effects of Cabo Codera upwelling. Nevertheless, our review of

chlorophyll images did not reveal the occurrence of an upwelling

plume during the time of sightings.

Until recently, the occurrence of Delphinus in the Gulf of Vene

zuela was based on the finding of skeletal remains in year 2002

(Ramírez-Carroz and González-Fernández 2004). Our addition of

one stranding and one sighting in that region (see Table 1, Fig. 2) con

firms the occurrence of free-ranging Delphinus in waters of the Gulf

of Venezuela. Recently, some sightings have been recorded to the

east of the Gulf of Venezuela in Aruban waters (Luksenburg 2013,

Smultea et al. 2004, 2013) and to the west, off the Colombian side of

the Guajira Peninsula (Palacios et al. 2012). Palacios et al. (2012) ten

tatively proposed the existence of a “Guajiran stock” that would be

separated from the “Venezuelan stock” by outflow of warm water

from the Lake of Maracaibo and the Gulf of Venezuela (Palacios et

al. 2012; Fig. 1). According to authors, this stock would inhabit the

Bolaños-Jiménez et al.

ANARTIA, 25 (2013 “2015”): 32 - 46

waters surrounding both the Colombian and Venezuelan sides of the

Guajira Peninsula. Palacios et al. (2012) also acknowledged that a

more or less continuous distribution of common dolphins from

northeastern Venezuela to northern Colombia is a possibility that

cannot be ruled out until further survey work is conducted. Contin

ued research efforts in the southern Caribbean will further help eluci

date the occurrence of Delphinus as well as its association with envi

ronmental parameters.

ACKNOWLEDGEMENTS

The ICLAM-MINAMB (Instituto para la Conservación y Con

trol de la Cuenca del Lago de Maracaibo, Ministerio del Poder Popu

lar para el Ambiente) provided logistical support for the participation

of two of the authors (MFP and LSC) in field work in the State of Zu-

lia. The field work in Aragua and Miranda States was possible be-

cause or the support of Cetacean Society International (CSI), Hu-

mane Society International (HSI), Fondo IEA-Provita, Fundacite

Aragua/U.T. MPPCTI, Gobernación del Estado Miranda, Policía

del Estado Miranda, the Rufford Foundation, Armando José Díaz,

and Familia Macía Castillo. Ignacio Moreno and Charley Potter

confirmed the identification of the specimen stranded in the State of

Zulia. Julián Castañeda, Eduardo Klein, Mari Smultea, Meike Holst

and Cathy Bacon provided useful references and information.

Ninive Espinoza produced the map in Figure 1. This manuscript was

greatly improved by comments from Dagmar Fertl and Jolanda Luk

senburg.

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