INTRODUCTION
Ecuador possesses a great variety of coastal environments allowing for a high diversity of marine species. However, systematic studies on marine invertebrates are scarce, especially in polychaetes. In continental Ecuador, only 29 families, 53 genera and approximately 75 species of polychaetes are recorded so far. Hartman (Reference Hartman1939) pioneered polychaete studies by describing four new species and ten new records from Puna and Santa Clara Islands (Guayas Province). Later, Cruz et al. (Reference Cruz, De Gonzáles, Gualancañay and Villamar1980) reported four new records from benthic samples collected on the Estero Salado of Guayaquil Gulf. In the same Gulf, near that area, 29 species of polychaetes were identified by Villamar (Reference Villamar1983), one of which was a new species of Grubeulepis. The same author in 1989 reported nine new records at Canal del Morro and Jambeli in the Guayaquil Gulf. In the intertidal zone of Monteverde (Guayas Province), Villamar & Cruz (Reference Villamar and Cruz2007) later reported three new records for Ecuador. A new species of Australonuphis used as fishing bait, was described by de León-González et al. (Reference León-González, Cornejo-Rodriguez and Degraer2008) in Santa Elena Bay (Guayas Province). In northern Ecuador, very little is known about this group, and only one ecological study has been carried out by Villamar (Reference Villamar2006) in the intertidal zones of Manabi and Esmeraldas Provinces. In that paper, the author reported 12 genera and 27 species, of which 14 constituted new records for this country.
In order to increase the knowledge about this group of invertebrates on the Ecuadorian coast, we herein provide a checklist of the polychaete fauna from Bunche and Cabo San Francisco Beaches (Esmeraldas Province), northern Ecuador.
MATERIALS AND METHODS
Samples were collected in March 2009 (dry season) in the intertidal zone of two sandy beaches: Bunche (0°39′01.98″N 80°03′55.01″W) and Cabo San Francisco (0°38′16.35″N 80°3′14.07″W) located in the Esmeraldas Province, of northern Ecuador (Figure 1). The beach dynamics are different. Bunche is characterized as a low energy beach, a calm and protected area, with soft sloping banks and very fine particle sand, while Cabo San Francisco is characterized as a high energy environment (subjected to frequent and severe storms), with high slopes and larger than average grain size. Fresh water discharges affect both beaches. Sediment samples were sieved through a 1-mm mesh. All the polychaetes species were separated from the sediment, except one: Phyllodoce multiseriata Rioja, 1941, which was found in tidal pools at low tide. Specimens were fixed in 10% formalin and later preserved in 70% ethanol. The identifications were made using the taxonomic keys of Fauchald (Reference Fauchald1977) and de León-González et al. (Reference León-González, Bastida-Zavala, Carrera-Parra, García-Garza, Peña-Rivera, Salazar-Vallejo and Solís-Weiss2009). Samples were deposited at the Polychaetological Collection of the Universidad Autónoma de Nuevo León (Phyllodoce multiseriata, UANL 7803; Opisthosyllis arboricola, UANL 7804; Hemipodia pustatula, UANL 7805; Nereis riisei, UANL 7806; Pseudonereis gallapagensis, UANL 7807).
Fig. 1. Map of Ecuador indicating the sampling sites, Bunche and Cabo San Francisco Beaches.
RESULTS
Twelve polychaete species belonging to twelve genera and nine families were recorded from Bunche and Cabo San Francisco Beaches. This list includes eleven new records for Ecuador and one, Nereis riisei, for northern Ecuador. Short descriptions of the polychaete species sampled for this study are given below.
Fig. 2. Polychaetes from Bunche and Cabo San Francisco Beaches, northern Ecuador: (A) Hemipodia pustatula; (B) Microphthalmus arenarius, anterior end, dorsal view; (C) Scoletoma zonata, anterior end, ventral view; (D) Nereis riisei, anterior end, dorsal view; (E) Pseudonereis gallapagensis; (F) Armandia cf. salvadoriana; (G) Euzonus mucronata, whole specimen; (H) Phyllodoce multiseriata; (I) Pisione longispinulata, anterior end, dorsal view; (J) Malacoceros indicus; (K) Scolelepis (Scolelepis) acuta, anterior end, lateral view; and (L) Opisthosyllis arboricola, anterior end, dorsal view. Scale bars: A, C, E, F, G, H, J, K, L, 1 mm; I, 0.5 mm; B, D, 0.2 mm.
Hemipodus pustatulus: Friedrich, Reference Friedrich1956: 61, figure 4A–C.
Hemipodus ciliatus: Hartmann-Schröder, Reference Hartmann-Schröder1959: 150, figures 128–131.
Hemipodus longipapillatus: Hartmann-Schröder, Reference Hartmann-Schröder1965: 150, figures 118–121.
Hemipodia pustatula: Böggemann & Eibye-Jacobsen, Reference Böggemann and Eibye-Jacobsen2002: 83, figures 136–138.
MATERIAL EXAMINED
Bunche Beach, two complete specimens, six anterior and three posterior fragments, 21 March 2009, coll. Berenice Trovant and Santiago Tineo.
DIAGNOSIS
Up to 40 mm long and 1 mm wide, 118 chaetigers. Live specimens bright red; preserved specimens orange to brown. Mandibles with simple rod-like ailerons. Pharynx covered by two types of papillae, with ‘U’-shaped ridges: digitiform ones with 10–15 ridges and conical ones with 8–10 ridges. Branchiae absent. All parapodia uniramous, with one long subtriangular prechaetal lobe and one rounded to triangular postchaetal lobe; prechaetal lobe with digitiform distal process from median parapodia to the posterior end of the body. Parapodia with one aciculum and compound spinigers. Globular dorsal cirri. Triangular ventral cirri on anterior parapodia, on median parapodia subulate and on posterior parapodia rounded to oval. Pygidium unknown.
DISTRIBUTION
Pacific Ocean: México (Bahía Kino and Mazatlán), El Salvador (Metalio, Acajutla, Las Salinas, La Libertad, San Diego, Toluca, Amate Campo, Las Hojas, Los Blancos, El Zapote, La Chepona and El Cuco), Costa Rica, Panama and Chile (Bahía San Vicente); Atlantic Ocean: Caribbean Sea, Costa Rica, Panama and Brazil (Paranagua).
Microphthalmus arenarius: Westheide, Reference Westheide1973: 243–248, figures 1–3; Salazar-Vallejo & Rizzo, Reference Salazar-Vallejo, Rizzo, de León-González, Bastida-Zavala, Carrera-Parra, García-Garza, Peña-Rivera, Salazar-Vallejo and Solís-Weiss2009: 227–244.
MATERIAL EXAMINED
Cabo San Francisco Beach, six complete specimens, two anterior and two posterior fragments, 20 March 2009, coll. Berenice Trovant and Santiago Tineo.
DIAGNOSIS
Up to 1.5 mm long and 0.3 mm wide, 22 chaetigers. Preserved specimens brown. Cylindrical body, with three antennae, median one inserted at posterior margin of prostomium. Six pairs of long tentacular cirri inserted on following three segments. A lateral pair of red eyes, located posteriorly in the prostomium. Simple palps with palpophore reduced. Notopodia reduced with compound spinigers, neuropodia with falcigers. An expanded anal plate on the posterior end. The anal plate is smooth and at least as wide as the last chaetiger. Two anal cirri, twice as long as anal plate.
DISTRIBUTION
Atlantic Ocean: Bermudas.
Lumbriconereis zonata: Johnson, Reference Johnson1901, 408–409, pl. 9, figures 93–100.
Lumbrineris zonata: Hartman, Reference Hartman1944, 146–147; Fauchald, Reference Fauchald1970, 112, pl. 18, figure E–I; Fauchald, Reference Fauchald1973, 24.
Scoletoma zonata: Carrera-Parra, Reference Carrera-Parra, de León-González, Bastida-Zavala, Carrera-Parra, García-Garza, Peña-Rivera, Salazar-Vallejo and Solís-Weiss2009: 263–276.
MATERIAL EXAMINED
Bunche Beach, one complete specimen, three anterior fragments and one posterior fragment, 21 March 2009, coll. Berenice Trovant and Santiago Tineo.
DIAGNOSIS
Up to 65 mm long and 0.5 mm wide, for 303 chaetigers. Preserved specimens pink to brown, highly iridescent, with a little lateral black intersegmental dot in the middle of each segment throughout body. Subtriangular prostomium. Maxillary formula as follows: with five pairs of maxillae. Carrier jaws attached to the maxilla I throughout the base. Plate connecting the maxilla I with II, broad and slightly sclerotized. Maxilla II as long as maxilla I. Maxilla I with a single tooth, maxilla II with three teeth and accessory lamella. Accessory lamella wider than maxilla II and occupies three-quarters of its lateral border. Maxilla III with two teeth, maxilla IV with a single tooth, both fully pigmented. Notopodia reduced with small dorsal cirri. Postchaetal lobe slightly longer than the prechaetal one. Without branchiae. Simple hooded hooks with short cap and simple limbate chaetae from chaetiger 1. Pygidium with two pairs of short anal cirri.
DISTRIBUTION
Pacific Ocean: United States (Dillon Beach, Tomales Bay, Bodega Lagoon and Puget Sound region), Costa Rica and Panama (Venado Beach).
Nereis riisei: Grube, Reference Grube1857: 162; Monro, Reference Monro1933: 43; Hartman, Reference Hartman1940: 221, lám. 33, figure 37; Fauchald, Reference Fauchald1977: 31, figure 8C–E; Taylor, Reference Taylor, Uebelacker and Johnson1984: 31–38, figures 31–36A–G; de León-González & Solís-Weiss, Reference León-González and Solís-Weiss2000: 560; Santos & Lana, Reference Santos and Lana2003: 12, figures 34–40; de León-González et al., Reference León-González, Bastida-Zavala, Carrera-Parra, García-Garza, Peña-Rivera, Salazar-Vallejo and Solís-Weiss2009: 325–354.
MATERIAL EXAMINED
Bunche Beach, one incomplete specimen, 21 March 2009, coll. Berenice Trovant and Santiago Tineo.
DIAGNOSIS
Specimen of 10 mm long, 1 mm wide, 55 chaetigers. Preserved specimen unpigmented. Prostomium pentagonal in shape, with two pairs of eyes in trapezoidal arrangement, the posterior pair larger. A pair of short antennae that do not exceed in length the distal part of the palps. Globose palps, subspherical palpostyles. Peristomium as long as the following two segments, but less wide, with four pairs of tentacular cirri, the pair of larger cirri reach chaetiger 3. Pharyngeal paragnaths not visible given the size of specimen. Subconical dorsal ligule, rounded median ligule in all notopodia. Neuropodia with subtriangular ligule on anterior parapodia, and subulate on median and posterior parapodia. Dorsal cirri inserted medially on dorsal ligule, ventral cirri inserted below the ventral ligule. Notochaetae with homogomph spinigers in all parapodia; in median and posterior parapodia one homogomph long bladed falciger appears. Neuropodia with homogomph and heterogomph spinigers and short bladed heterogomph falciger. Pygidium unknown.
DISTRIBUTION
Pacific Ocean: Mexico (Santa Maria Bay, Gulf of California, Guerrero, Socorro Island), Costa Rica (Nicoya Gulf, Papagayo Gulf), Panama (Gorgona Island, Coiba Island), Colombia (Gorgona Island) and Ecuador (La Libertad, Galapagos Island); Atlantic Ocean: Gulf of Mexico (Florida, Texas, Veracruz, Ciudad del Carmen, Alacranes Reef, Cayo Arcas, Triángulos Oeste, Cayo Nuevo) and Brazil (Maranhão, Piauí, Ceará, Rio Grande do Norte, Paraíba, Pernambuco, Bahia, Alagoas); Caribbean Sea: Bonaire, Anguilla, St Eustatius, Aruba, Curaçao, Barbados, St Vincent, Grenada, Barbuda, Antigua, Jamaica, Cuba (La Habana Bay), Venezuela (Cariaco Gulf), Colombia (Nenguangue Bay), Panama (Galeta Reef) and Mexico (Nicchehabin Reef, east of Allen Point, Ascencion Bay and Cozumel Island).
Pseudonereis gallapagensis: Fauvel, Reference Fauvel1939: 328; Reference Fauvel1953: 215, figure 110A–C; Hartman, Reference Hartman1940: 231; Rioja, Reference Rioja1960: 297; Hartmann-Schröder, Reference Harmann-Schröder1962: 432; Wesenberg-Lund, Reference Wesenberg-Lund1962: 84, figure 32; Imajima, Reference Imajima1972: 97, figure 28A–J; Fauchald, Reference Fauchald1977: 32, figure 4G–H; Liñero-Arana & Reyes-Vázquez, Reference Liñero-Arana and Reyes-Vázquez1979: 8, figures 1–10; Rozbaczylo & Bolados, Reference Rozbaczylo and Bolados1980; 219, figure A–D; Wu et al., Reference Wu, Ruiping and Yang1985: 220, figure 124A–F; Hylleberg et al., Reference Hylleberg, Nateewathana and Bussarawit1986: 14, figure 8A–P; Bakken & Wilson, Reference Bakken and Wilson2005: 534; de León-González et al., Reference León-González, Bastida-Zavala, Carrera-Parra, García-Garza, Peña-Rivera, Salazar-Vallejo and Solís-Weiss2009: 325–354.
MATERIAL EXAMINED
Cabo San Francisco Beach, one complete specimen, 20 March 2009, coll. Berenice Trovant and Santiago Tineo.
DIAGNOSIS
Specimen of 11 mm long, 0.5 mm wide, 61 chaetigers. Preserved specimen light brown. Sub-pentagonal prostomium. A pair of antennae and a pair of globose palps. Two pairs of black eyes. Four pairs of tentacular cirri, dorsal one longest. Birramous parapodia. Cirriform dorsal cirri in all parapodia. All notopodia highly developed, with ligule, lobes and chaetae. Pharynx armed in both rings, maxillary ring, Group I, two small cones; Group II, three pectins; Group III, four pectins in oval arrangement; Group IV, eight small cones and 20 wide basally, in the distal region, four in accordance pectins' growing arrangement. Oral ring, Group V, one big cone; Group VI, a short transverse bar into a cone shape; Groups VII–VIII, 22 cones in two lines. Dorsal cirrus inserted distal to the dorsal ligula in posterior parapodia. Notopodia with homogomph spinigers and neuropodia with homogomph spinigers and heterogomph falcigers. Pygidium with two ventrolateral anal cirri.
DISTRIBUTION
Pacific Ocean: Ecuador (Galapagos Islands), Peru and Chile; Atlantic Ocean: Gulf of Mexico, Panama, Brazil, South Africa; Indian Ocean.
Armandia salvadoriana: Hartmann-Schröder Reference Hartmann-Schröder1956: 95, figures 6–8.
MATERIAL EXAMINED
Bunche Beach, eight complete specimens, 21 March 2009, coll. Berenice Trovant and Santiago Tineo.
DIAGNOSIS
Up to 13 mm long and 1 mm wide, 35 chaetigers. Preserved specimens unpigmented. Elongate body with a lateral groove. Prostomium long, conical and acuminate; large nuchal organs, without ocelli. Cirriform branchiae, longer than notochaetae, recurved over the body, present from chaetiger two to the end of body, although in some individuals they may be absent in the last chaetigers. Anal funnel long with 17–19 rings, marginal festoon with 6–7 lanceolate papillae and with a long medial cirri. With 23–25 pairs of lateral eyes starting from chaetigers 6–7.
DISTRIBUTION
Pacific Ocean: El Salvador (San Juan, Pajarita and Madre-Sal) and Costa Rica (Golfo de Nicoya).
REMARKS
In the original description of A. salvadoriana Hartmann-Schröder, Reference Hartmann-Schröder1956, the number of pairs of eyes is 15, while our specimens have 23–25 pairs of eyes. Type material of A. salvadoriana was not observed; however, the Ecuadorian form is similar to the El Salvadorian one in all the remaining characters.
Euzonus mucronata: Blake, Reference Blake, Blake, Hilbig and Scott2000: 152–153, figure 7.2A–C, F.
MATERIAL EXAMINED
Bunche Beach, six complete specimens, 21 March 2009, coll. Berenice Trovant and Santiago Tineo.
DIAGNOSIS
Up to 14 mm long and 0.5 mm wide, 38 chaetigers. Live specimens unpigmented except for the pink gut; preserved specimens unpigmented. Body divided into three regions: cephalic region that includes the prostomium and the first two chaetigers; a thoracic region that includes the chaetigers 3–11 and the abdominal region that comprises two chaetigers without branchiae, 18 chaetigers with branchiae and eight post-branchial segments. Cylindrical body on the anterior segments, with ventral groove on branchial region that continues to the posterior end. Prostomium small and conical, buccal organs present. Parapodia birramous; all chaetae are capillaries; notochaetae longer than neurochaetae. Smooth branchiae with two branches. Pygidium with a long ventral cirrus and seven pairs of digitiform lateral lobes.
DISTRIBUTION
Pacific Ocean: Canada, United States (Dillon Beach) and Mexico.
Phyllodoce (Anaitides) multiseriata: Salazar-Vallejo, Reference Salazar-Vallejo1989, 45.
Phyllodoce multiseriata: Glockner-Fagetti, Reference Glockner-Fagetti, de León-González, Bastida-Zavala, Carrera-Parra, García-Garza, Peña-Rivera, Salazar-Vallejo and Solís-Weiss2009: 409–424.
MATERIAL EXAMINED
Bunche Beach, eight complete specimens, two anterior and four median fragments, 21 March 2009, coll. Berenice Trovant and Santiago Tineo.
DIAGNOSIS
Up to 45 mm long and 1.5 mm wide, 155 chaetigers. Live specimens greenish with a brown intersegmental transverse band; preserved specimens with the same pattern colour. Prostomium chordate. With four antennae and nuchal papilla. A pair of red eyes. Four pairs of tentacular cirri. Pharynx with conical papillae arranged regularly in the basal ring. First two segments fused dorsally. Uniramous parapodia, rounded dorsal cirri, pyriform ventral cirri, smaller than the dorsal one. Neuropodia with spinigers. Pygidium with two anal cirri.
DISTRIBUTION
Pacific Ocean: Canada (British Columbia) and Mexico (La Aguada, Acapulco and Guerrero).
Pisione longispinulata: Aguado & San Martín, Reference Aguado and San Martín2004: 75–78, figures 2–3.
MATERIAL EXAMINED
Cabo San Francisco Beach, 82 complete specimens, nine anterior, two median and four posterior fragments, of which 29 were males (26 complete specimens and three anterior fragments); 20 March 2009, coll. Berenice Trovant and Santiago Tineo.
DIAGNOSIS
Up to 3.5 mm long, 0.5 mm wide, 36 chaetigers. Preserved specimens unpigmented. Body elongate and filiform. Small prostomium, buccal segment directed forward and partially fused to prostomium; without chaetae. A long pair of smooth palps. Buccal segment with dorsal tentacular cirri directed forward, elongated (approximately one-third of the length of the palps); short globular ventral cirri. This segment carries a pair of buccal aciculae that arise internally at the base of the dorsal cirri, firm, slightly emerging and distally expanded. Globular dorsal cirri with a distal papilla small and spherical. A pair of eyes located between chaetigers 2–3. Reversible pharynx, with two pairs of chitinous jaws seen by transparency. Brown mandibles occur in the third chaetiger. Parapodia with four chaetae, one simple chaetae and three falcigers (the first longest) with long secondary spines. Notoacicula embedded throughout the entire body, not protruding. The 29 males had a pair of copulatory organs located between chaetigers 12 and 27, and three of them also had a second pair located between chaetigers 25 and 28. Rounded pygidium with a pair of long anal cirri.
DISTRIBUTION
Pacific Ocean: Panama (Coiba Island).
REMARKS
Aguado & San Martín (Reference Aguado and San Martín2004) described Pisione longispinulata based on two males and found that they had one copulatory organ between segments 18 and 20, while in our specimens we found the first pair between segments 13 and 28. Furthermore, three specimens presented a second pair of copulatory organs. Given the number of specimens we have been able to observe, we believe there might be a wider variation in the number and in the distribution of the copulatory organs in this species.
Malacoceros indicus: Pettibone, Reference Pettibone1963: 99; Day, Reference Day1967: 477, figure 18.5p–u; Blake, Reference Blake1983: 219; Reference Blake, Blake, Hilbig and Scott1996: 105–107, figure 4.4; Delgado-Blas, Reference Delgado-Blas, de León-González, Bastida-Zavala, Carrera-Parra, García-Garza, Peña-Rivera, Salazar-Vallejo and Solís-Weiss2009: 589–615.
MATERIAL EXAMINED
Bunche Beach, one anterior fragment, 21 March 2009, coll. Berenice Trovant and Santiago Tineo.
DIAGNOSIS
Specimen of 25 mm long, 1 mm wide. Live specimen unpigmented except for the green digestive tube, preserved specimen unpigmented. Body wide anteriorly. Prostomium bell-shaped with frontal horns. A pair of red eyes. With cirriform branchiae from first chaetiger. Branchiae basally fused to postchaetal lamellae and short on posterior region of the body. Capillary chaetae simple. With neuropodial hooks. Hooded hooks bidentate from chaetigers 30 to 50. Pigidium unknown.
DISTRIBUTION
Pacific Ocean: Chile; Atlantic Ocean: Gulf of Mexico, South Africa; Indian Ocean: Gulf of Manaar, Mozambique and New Caledonia.
Scolelepis (Scolelepis) acuta: Delgado-Blas, Reference Delgado-Blas2006: 79–81, figure 2; Delgado-Blas, Reference Delgado-Blas, de León-González, Bastida-Zavala, Carrera-Parra, García-Garza, Peña-Rivera, Salazar-Vallejo and Solís-Weiss2009: 589–615.
MATERIAL EXAMINED
Bunche Beach, five anterior fragments and one median fragment, 21 March 2009, and Cabo San Francisco Beach, seven anterior fragments, 20 March 2009, coll. Berenice Trovant and Santiago Tineo.
DIAGNOSIS
Up to 11 mm long and 1 mm wide. Preserved specimens unpigmented. Body elongated, conical prostomium and short peristomium. Two pairs of red eyes. Two long tentacular palps, slightly rugose, with two longitudinal rows of cilia. The first segment with short neurochaetae and without notochaetae. Chaetiger 5 without modified chaetae. Postchaetal lamella associated with notopodia and neuropodia of anterior region. All lamellae decrease in size toward posterior region. Branchiae from second chaetiger, basally fused to the lamella in most parapodia. Simple chaetae and neuropodial hooded hooks with three small teeth. Pygidium unknown.
DISTRIBUTION
Pacific Ocean: United States (San Diego, California).
Opisthosyllis arboricola: Góngora-Garza, Reference Góngora-Garza, de León-González, Bastida-Zavala, Carrera-Parra, García-Garza, Peña-Rivera, Salazar-Vallejo and Solís-Weiss2009: 615–639.
MATERIAL EXAMINED
Bunche Beach, one anterior and posterior fragment, 21 March 2009 and Cabo San Francisco Beach, one complete specimen, 20 March 2009, coll. Berenice Trovant and Santiago Tineo.
DIAGNOSIS
Up to 3 mm long, 0.5 mm wide, 50 chaetigers. Preserved specimens unpigmented. Pentagonal prostomium. Two pairs of eyes in trapezoidal arrangement. With three articulated antennae in the anterior region of the prostomium. Median antennae with 14 articles. Lateral antennae with ten articles, twice the length of the prostomium. Two free subtriangular palps. Anterior-body with long dorsal cirri formed by 14–22 articles; mid-body with long dorsal cirri with 16–17 articles, and posterior dorsal cirri with 12–14 articles. Triangular ventral cirri. Ventral parapodial lobe short, subconical and truncated, with prechaetal and postchaetal extension smaller than the last one. Anterior-body with 10–12 falcigers per fascicle, unidentate with well-defined tertiary teeth. Median-body with 7–9 falcigers per fascicle, dorsal falcigers with tertiary teeth and ventral smooth, all unidentate. Porsterior-body with 5–7 falcigers per fascicle, unidentate, smooth, without tertiary teeth. Pharynx wide, occupies almost the entire width of the body, as long as 7–8 segments. With middle-dorsal tooth in a posterior position. Middle-dorsal tooth single, large and yellow, in a posterior position. Proventricle as long as 8–9 segments, no observation of muscle cell ring. Dorsal acicular chaetae with rounded end and ventral acicular chaetae unidentate and smooth. The anterior region of the body with three aciculae, the median region with two aciculae and only one in the posterior region very close to the pygidium. Pygidium with two long articulate anal cirri. The cirri with approximately 12 articles each.
DISTRIBUTION
Atlantic Ocean: El Salvador (Estero Jaltepeque and Golfo de Fonseca).
ACKNOWLEDGEMENTS
This research was conducted by Berenice Trovant as part of the requirements for an MSc degree in ‘Biodiversity in tropical areas and its conservation’ at the Universidad Internacional Menéndez Pelayo (UIMP, Spain), a Masters programme funded by the Spanish National Research Council (CSIC, Spain) and carried out at the Universidad Central del Ecuador. This work would not have been possible without the help of the following people: Trovant's family, S.A. Tineo, J.M. Orensanz, J. Muñoz, G. Góngora, L.F. Carrera-Parra, D. Salcedo, V. Radashevsky, G. San Martín, A. Rizzo, S. Salazar-Vallejo, J.M. Serrano, F. Villamar, M. Cornejo, V. Carvajal, J. Santiana, C. Crowe, S. Heaslip and the staff of the Biological Reserve ‘The Congal’: Andrés, Fredy, Karina and Vicente. Also, we are grateful to Vivianne Solis-Weiss and the two anonymous referees that contributed substantially to improve the manuscript.
