Spatial and seasonal patterns in species composition of fish larvae in the Canary Islands by Agencia Canaria de Investigación, Innovación y Sociedad de la Información (ACIISI)

INFORME TÉCNICO DEL INSTITUTO CANARIO DE CIENCIAS MARINAS

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

Mirjam Bachler

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Instituto Canario de Ciencias Marinas

Agencia Canaria de Investigación, Innovación y Sociedad de la Información

Gobierno de Canarias Autora: Mirjam Bachler. OE Clinical Trial Center Medical University Innsbruck

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Mirjam Bachler.

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SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

RESUMEN Se estudió la composición y presencia de especies de larvas de peces alrededor de las Islas Canarias en relación a la influencia de las características hidrográficas sobre el patrón de distribución espacial y estacional. Se proporciona una lista de taxa de larvas con los caracteres taxonómicos más importantes. Las larvas, la mayoría en fase larvaria tardía, fueron capturadas en diferentes estaciones durante seis campañas pelágicas realizadas entre 1997 y 2002, en distintas estaciones del año, tanto en el reino nerítico como en el oceánico de Lanzarote, Fuerteventura, Gran Canaria y Tenerife. Se recolectaron un total de 5088 larvas, pertenecientes a 16 órdenes, 54 familias y 70 especies, donde Engraulidae y Gobiidae dominan en la comunidad larvaria. Se documentan cinco nuevas citas para el área del Archipiélago Canario. En el área de las Islas Canarias se distinguen dos periodos estacionales con diferentes condiciones hidrográficas, en invierno con un afloramiento de baja intensidad y alta en primavera y verano. La floración fitoplanctónica invernal tardía (LWB) junto con el aumento del afloramiento de la costa Oriental Africana coincide con un aumento de la densidad y del número de especies de larvas. La diversidad local y el número de especies fueron más altas en un área influenciada por el afloramiento del Noroeste Africano. Especialmente en las estelas y remolinos de las islas, las larvas se acumulaban solapándose las especies neríticas y oceánicas. Los resultados corroboran las hipótesis de que las Islas Canarias son una importante área migratoria facilitando la colonización por especies de peces de zonas tropicales y templadas, el afloramiento conduce a un aumento de la producción local de larvas de peces, y las estelas y remolinos favorecen la abundancia y la diversidad de larvas de peces y proporcionan adecuadas zonas de cría.

ABSTRACT The species composition and occurrence of fish larvae in the area of the Canary Islands was studied regarding the influence of hydrographic features on spatial and seasonal distribution patterns. A comprehensive annotated larvae taxa list with the most important taxonomic characters is provided. Mostly late-stage larvae were caught by six pelagic cruises from 1997 to 2002 during different seasons in both the neritic and the oceanic realm off Lanzarote, Fuerteventura, Gran Canaria, and Tenerife. A total of 5088 larvae, allocated to 16 orders, 54 families and 70 species, were collected with Engraulidae and Gobiidae dominating the larval assemblage. Five new records could be documented for the Canarian Archipelago. In the area of the Canary Islands two main seasonal periods with different hydrographic conditions can be recognized, low intensity of the upwelling in winter and intense upwelling in spring and summer.The late winter bloom (LWB) and an associated stronger West African coastal upwelling coincided with an increased diversity and species number of fish larvae.The local diversity and species richness were highest in an area mostly influenced by the NW African upwelling. Especially in the islands’ wakes and eddies the larvae accumulated with spatial overlap of oceanic and neritic species. These results corroborate the assumptions that the Canary Islands are an important migratory area facilitating the colonization by tropical and temperate fish species, upwelling enhances local productivity, and the islands’ wakes and eddies positively affect fish larvae abundance and diversity and provide good nursery grounds.

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ZUSAMMENFASSUNG Die Artenzusammensetzung und das Vorkommen von Fischlarven im Bereich der Kanarischen Inseln wurden hinsichtlich des Einflusses der hydrographischen Bedingungen auf die räumlichen und saisonalen Verteilungsmuster untersucht. Außerdem wurde eine kommentierte Liste der Larventaxa mit den wichtigsten taxonomischen Merkmalen erstellt. Die Fischlarven, welche sich überwiegend in einem späten Entwicklungsstadium befinden, wurden bei sechs pelagischen Forschungsfahrten im Zeitraum von 1997 bis 2002 während unterschiedlicher Jahreszeiten im neritischen und ozeanischen Bereich rund um Lanzarote, Fuerteventura, Gran Canaria und Tenerife, gefangen. Insgesamt konnten 5088 Larven gesammelt werden, wobei sie 54 Familien, 77 Genera und 70 Arten zugeordnet werden konnten. Die Gemeinschaft der Larven wurde von den Engraulidae und den Gobiidae dominiert. Auch konnten fünf neue Arten für die kanarische Inselgruppe dokumentiert werden. Im Bereich der Kanarischen Inseln sind zwei Perioden mit unterschiedlichen hydrographischen Bedingungen vorherrschend. Zum einen, mit einem schwachen Auftrieb (Upwelling) im Winter und zum anderen, ein starkes Upwelling im Frühjahr und Sommer. Die Algenblüte gegen Ende des Winters (LWB) und ein verstärktes Upwelling an der Westafrikanischen Küste fällt mit einem einhergehenden Anstieg der Diversität und Artenzahl der Fischlarven zusammen. Räumlich gesehen waren die lokale Diversität und Artenzahl in den vom Westafrikanischen Auftriebsgebiet beeinflussten Bereichen am höchsten. Speziell in den bewegungsstillen Bereichen (Wakes) der Inseln und den von den Inseln induzierten Wirbeln (Eddies) akkumulierten die Fischlarven mit einer räumlichen Überlappung von neritischen und ozeanischen Arten. Diese Resultate bekräftigen die Annahmen dass die Kanarischen Inseln ein Gebiet für erhöhte Besiedlungraten von tropischen als auch temperaten Fischarten sind, das Upwelling zu einer höheren Produktivität beträgt, und die Wakes und Eddies der Inseln die Abundanz und Diversität von Fischlarven positiv beeinflussen und gute Aufwachsgründe bieten.

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INDEX 1 Introduction 11 1.1. Background and goals of this study

1.2. Oceanographic characterization of the area

2 Material and methods 19 2.1. Surveys and sampling method

2.2. Material treatment and taxonomic approach

2.3. Multivariate community analysis

2.4. Anova analysis

2.5. Geographical information system (gis) distribution maps

3 Results 27 3.1. Fish larvae taxonomic composition

3.2. Annotated larvae taxa list

¬ Albuliformes

¬ Anguilliformes

¬ Saccopharyngiformes

¬ Clupeiformes

Engraulidae 33 Clupeidae 34

¬ Argentiniformes

Argentinidae 35 Microstomatidae 37 Platytroctidae 38

¬ Stomiiformes

Gonostomatidae 39 Phosichthyidae 44 Stomiidae 47 Stomiinae 48 Astronesthinae 49 Melanostomiinae 49 Idiacanthinae 50

¬ Aulopiformes

Aulopidae 51 Synodontidae 52 Chlorophthalmidae 54 Notosudidae 55 Scopelarchidae 56 Alepisaurinae 58 Omosudinae 59 Paralepididae 60

¬ Myctophiformes

Myctophidae 63 Myctophinae 64 Lampanyctinae 69

¬ Lampriformes

Lampridae 75 Regalecidae 76

¬ Gadiformes

Macrouridae 78 Macrourinae 78 Melanonidae 78

¬ Stephanoberyciformes

Melamphaidae 79

INDEX ¬ Beryciformes

Diretmidae 80 Trachichthyidae 81

¬ Gasterosteiformes

Syngnathidae 83

¬ Scorpaeniformes

Scorpaenidae 84 Sebastinae 84 Scorpaeninae 85

¬ Perciformes

Moronidae 86 Percichthyidae 86 Serranidae 87 Serraninae 88 Anthiniinae 88 Callanthiidae 89 Apogonidae 90 Carangidae 91 Sparidae 92 Pomacentridae 94 Labridae 95 Chiasmodontidae 96 Blenniidae 97 Gobiidae

Gempylidae 99 Trichiuridae 100 Nomeidae 103 Caproidae

¬ Pleuronectiformes

104

105

Bothidae 105

¬ Tetraodontiformes

108

Balistidae 108 Tetraodontidae 109

3.3. Community analysis

111

3.4. Diversity of fish larvae

114

3.5. Biogeography

120

4 Discussion

125

4.1. Sampling method

125

4.2. Species composition

126

4.3. Occurrence and distribution patterns

130

4.4. Conclusion

133

5 Acknowledgements

135

6 References

137

7 Appendix

153

7.1. Overview table of all species encountered

153

7.2. Selected pictures of larvae

156

Engraulidae 156 Microstomatidae 156 Platytroctidae 157 Gonostomatidae 157 Sternoptychidae 158 Stomiidae 158 Aulopidae 160 Synodontidae 160 Chlorophthalmidae 161 Notosudidae 161 Scopelarchidae 161 Alepisauridae 162 Paralepididae 162

INDEX Myctophidae 162 Regalecidae 163 Macrouridae 164 Melanonidae 164 Diretmidae 164 Trachichthyidae 165 Syngnathidae 165 Diretmidae 166 Moronidae 166 Howellidae 167 Serranidae 167 Callanthiidae 167 Epigonidae 168 Carangidae 168 Chiasmodontidae 169 Chiasmodontidae 169 Gobiidae 169 Gempylidae 171 Scombridae 171 Caproidae 172 Bothidae 172 Tetraodontidae 173

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

1 INTRODUCTION

1.1. BACKGROUND AND GOALS OF THIS STUDY In recent years, the oceanography in the area of the Canarian archipelago, Eastern Central Atlantic, has been studied in particular regarding complex mesoscale hydrographic features (BARTON et al. 1998; BASTERRETXEA et al. 2002; HERNÁNDEZ-GUERRA et al. 2002 and 2003; JOHNSON and STEVENS 2000; KNOLL et al. 2002; LLINÁS et al. 2002; NAVARRO-PÉREZ and BARTON 2001; NEUER et al. 2002) and associated plankton and nekton distribution patterns (ARÍSTEGUI et al. 1997 and 2001; BASTERRETXEA et al. 2002; BÉCOGNÉE et al. 2006; BORDES et al. 1999. DAVENPORT et al. 2002; JOHN et al. 2001, 2004a and 2004b; MAYR 2004, NEUER et al. 2002; RODRÍGUEZ et al. 1999, 2001, 2004 and 2006; SANGRA et al. 2001; UIBLEIN et al. 1996, 1998, UIBLEIN and BORDES 1999; WIENERROITHER et al. 2009). The Canary Islands are a region of high productivity and high diversity close to the North West African Coast at about 28 to 29°N (Figure 1). Fuerteventura is the closest island to the African continent at about 90 km distance from the Moroccan coast, while the entire archipelago extends up to 400 km further west (BARTON et al. 1998). Typical for the Canary Islands is the narrow shelf adjacent to steep slopes which leads to near-shore conditions similar to those in the open ocean (NAVARRO-PÉREZ and BARTON 2001) and may induce coastal trapping of open-ocean dwellers and intense trophic interactions between oceanic and neritic fauna (UIBLEIN and BORDES 1999). BRITO et al. (2002) provide a comprehensive list of marine fish species from the area of the Canary Islands. Recent findings from pelagic cruises indicate that the total number of species may be significantly higher and further investigations should be carried out, in particular of deep pelagic fishes (WIENERROITHER 2005). Fish larvae occurring in this area have been studied only to a limited extent (BÉCOGNÉE et al. 2006; JOHN et al. 2001 and 2004a; RODRÍGUEZ et al. 1999, 2001, 2004 and 2006). Based on the growing knowledge of fish diversity in the Canary Islands, various attempts have been made to characterize the fish species composition of this are zoogeographically. According to earlier studies of mainly mesopelagic fishes, the archipelago lies within the North Atlantic Subtropical Faunal region with the eastern islands (Fuerteventura and Lanzarote) belonging to the Temperate-Semisubtropical region and the western islands (Gran Canaria, Tenerife, La Gomera, La Palma and El Hierro) being part of the Tropical-Semisubtropical region (BACKUS et al. 1977).This separation follows a hydrographic gradient from east to west. BORDES et al. (1999) and following workers have emphasized the need of a micro-biogeographical approach that takes local oceanographic conditions more into consideration.

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1 INTRODUCTION

November 1997

April 1999

May 1999

January 2000

November 2000

March 2002

Figure 1. Chlorophyll distribution around the Canary Islands during the months of investigation (www.oceancolor.gsfc.nasa.gov)

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SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

Regarding deep-water demersal and coastal fishes, the close affinity of the Canary Islands with the adjacent NW African coast, the Mediterranean outflow, the next-closest island Madeira, and seamounts in the immediate surroundings has been outlined (e.g. BRITO et al. 2006; LLORIS et al. 1991; UIBLEIN et al. 1999). Further studies that carefully integrate findings on fish distribution patterns and oceanographic conditions will certainly contribute to more detailed insights into both the small- and the large-scale zoogeography of the Canary Islands. Studies of fish larvae have particular relevance in this context, as they provide information on migration, colonization, and local recruitment patterns. Examples for the dependence of fish larval distribution on hydrographic features and how such features influence their community structures or distribution and affect the survival of single specimens are numerous (ACEVES-MEDINA et al. 2004; CHIH-HAO and TAI-SHEN 2002; FOCK and JOHN 2006; JOHN et al. 2001 and 2004b; LEIS 1986; LOBEL and ROBINSON 1988; LOGERWELL and SMITH 2001; SABATÉS and MASÓ 1992; SASSA et al. 2002 and 2004; SPONAUGLE et al. 2005). Since the Canarian Archipelago is a region of high hydrographic variability, it is highly interesting for studying fish larvae (BÉCOGNÉE et al. 2006; JOHN et al. 2004a; RODRÍGUEZ et al. 1999, 2001, 2004 and 2006) and to link the hydrographic features around the Canarian Archipelago to their distribution, especially regarding larvae with different habitats of origin. This study contributes to further knowledge of the diversity, abundance, and distribution of fish larvae in the Canary Island area based on examination of material collected during six pelagic cruises in the area.The main objectives are (1) to provide an annotated list of fish larvae taxa caught during these surveys, (2) to relate their distribution and abundance to oceanographic processes around the Canary Islands, and (3) to identify spatial and seasonal patterns in community structure and diversity. In order to provide sufficient background information for the discussion and conclusions to be drawn, the currently available information on the hydrographic situation around the Canarian Archipelago and the possible consequences for fish larvae abundance and diversity is reviewed in the section below.

1.2. OCEANOGRAPHIC CHARACTERIZATION OF THE AREA In general, the waters around the Canary Islands are oligotrophic and the autotrophic community is dominated by picoplankton, while the mesozooplankton is composed mainly of copedods (ARÍSTEGUI et al. 2001). Microheterotrophic organisms such as ciliates transfer the primary production to the higher trophic level of the mesozooplankton. Only a small proportion of the total primary production from larger cells (microplankton) is directly transferred to mesozooplankton herbivores (ARÍSTEGUI et al. 2001). A phytoplankton bloom and peak of primary production occurs in late winter in the investigated area, the so-called late winter bloom (LWB) seen in Figure 1a, d and f (ARÍSTEGUI et al. 2001; DAVENPORT et al. 2002). In addition to this LWB a local permanent increase in production around the islands can be observed then. This is a result of the perturbation of oceanic and atmospheric flows created by the islands themselves (SANGRA et al. 2001), as depicted in Figure 1a-f. Noteworthy, the chlorophyll distribution is not related to the mesozooplankton biomass distribution (ARÍSTEGUI et al. 2001). Particularly copepods, the main food source of fish larvae, are spatially negatively related to chlorophyll minima, with the highest abundance occurring at the margins of areas with high primary productivity (BARTON et al. 1998; CHIH-HAO and TAI-SHEN 2002). The peak in mesozooplankton abundance is temporally delayed from the phytoplankton peak, which as already mentioned before occurs from March to June (ARÍSTEGUI et al. 2001). This is caused by the prolonged duration of energy transfer into the higher trophic level of mesozooplankton via microheterotrophic organisms, as described above.

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1 INTRODUCTION

Further reasons for local higher production in the area of the Canary Islands are the characteristic oceanic events, such as coastal upwelling, upwelling filaments, Islands’ wakes and eddies (ARÍSTEGUI et al. 1997; BARTON et al. 1998). These mesoscale hydrographic features induce an enhanced zooplankton biomass, creating the nutritional regime necessary to sustain fish larvae and other higher trophic levels (YEBRA et al. 2004). The eggs of most fishes are pelagic and the majority of larvae undergo a pelagic phase in which they inhabit mainly the upper layers of the water column. This is related to the distribution of their food sources, therefore fish larval distribution differs from adult occurrence. Additionally, larvae often drift far away from their spawning areas (ACEVES-MEDINA et al. 2003), as most are not able to overcome current speeds and the flow velocities of mesoscale hydrographic features. The mean swimming velocity of larvae is around 20.5 cm per second, though there are remarkable differences between families (LEIS and CARSON-EWART 1997; LEIS and FISHER 2006). For example apogonids only have an average swimming speed of 6.3 cm per second (LEIS and CARSON-EWART 1997), which is clearly below the flow velocities of on- and offshore currents (MITTELSTAEDT 1983). One possibility for larvae to influence their distribution is the vertical migration. By changing layers with different directional flows they might be able to maintain their location (RODRÍGUEZ et al. 2001). Nevertheless, patterns of ichthyoplankton distribution can be interpreted in terms of the physical characteristics of an area (RODRÍGUEZ et al. 2001). One such physical characteristic of the area of investigation is created by the Canarian archipelago itself. It presents a barrier within the southward flowing Canary Current and acts as a transitional zone from the shelf upwelling region of northwest Africa to the open ocean waters of the eastern North Atlantic (BARTON et al. 1998). As a result of this hydrographic position BARTON et al. (1998) divided the area into four hydrographic subdivisions, which can be seen in Figure 1a-f. The main separation is into a northern and a southern part of the archipelago. In the north the flow of the Canary Current is undisturbed, with an eastern area affected by the African upwelling and an unaffected western area. The southern area is characterized by perturbations caused at the island-ocean interface. Such disturbances are island wakes in connection with winds, island upwelling and island induced eddies (NAVARRO-PEREZ and BARTON 2001). This area is also differentiated between eastern areas which are affected by coastal upwelling and western areas which are not. Due to the perturbation of the Canary Current in its southward flow the current is mixed vertically at the island’s margin (BARTON et al. 1998), resulting in a meandering current (NAVARRO-PÉREZ and BARTON 2001). The Canary Current itself originates from the southward flowing Azores Current which splits into several branches. The easternmost branch, which turns into the Canary Current, flows southward parallel to the African coast and through the Canarian Archipelago (HERNÁNDEZ-GUERRA et al. 2002; JOHNSON and STEVENS 2000; KNOLL et al. 2002). The current consists of different water masses with different properties. The Surface Waters (SW), as the uppermost mixed layer, is created under the influence of local atmospheric conditions (KNOLL et al. 2002). Around the islands the Surface Waters reach a depth of 150 m (KNOLL et al. 2002), though the extension of this layer varies seasonally with a peak during spring (ARÍSTEGUI et al. 2001; BARTON et al. 1998). An increased insolation during summer leads to a strong heating of the SW, producing a temporal thermocline (ARÍSTEGUI et al. 2001; BARTON et al. 1998), which prevents nutrients from permeating the euphotic zone. This limits phytoplankton growth during summer (ARÍSTEGUI et al. 2001). Cooling and wind stress obliterate this thermocline in autumn, resulting in homogenously distributed chlorophyll in the surface mixed layer once again (ARÍSTEGUI et al. 1997). Below the SW, the North Atlantic Central Waters (NACW) flow unaffected by atmospheric conditions, extending to about 700 m depth (KNOLL et al. 2002). These are followed by the Antarctic Intermediate Water (AAIW), which forms an intermediate layer in the depth range of 600 – 1100 m (HERNÁNDEZ-GUERRA et al. 2003; KNOLL et al. 2002; LLINÁS

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et al. 2002). This layer mixes with surrounding waters, moves northwards and extends westwards to the Island of Tenerife (HERNÁNDEZ-GUERRA et al. 2003). The deepest layer has its origin in the Mediterranean Sea. This very saline European Mediterranean Water (EMW) lies deepest directly above the bottom (HERNÁNDEZ-GUERRA et al. 2003; KNOLL et al. 2002) and entrains layers of the same density. The flow of the different water masses varies seasonally. In spring and summer the Canary Current is strongest, with SW and NACW yielding the highest intensity of southward transport in spring, turning to flow northwards in summer and autumn (HERNÁNDEZ-GUERRA et al. 2003; JOHNSON and STEVENS 2000; KNOLL et al. 2002). The current strength of the AAIW is strongest during autumn (KNOLL et al. 2002), when northward transport is at a minimum (HERNÁNDEZGUERRA et al. 2003). Furthermore, a permanent counter current exists within the Canary Current at a depth of about 200 – 400 m (KNOLL et al. 2002; MITTELSTAEDT 1983).This counter current is mainly due to a compensating force in the horizontal water distribution. The south-eastern region of the North East Atlantic is characterised by an excess of water due to the equatorial counterand undercurrent transporting water to the east. Simultaneously, a deficit of water along the northern African west coast occurs as the Canary Current flows westwards away from the coast. This forces the water from the equator to the north (MITTELSTAEDT 1983). With few exceptions, the counter current usually does not reach the inshore region, but if it reaches the coast it can become subsurface or even surface (MITTELSTAEDT 1983; vAN CAMP et al. 1991). The counter current is strongest during summer since the equatorial counter current is most intense then and therefore provides an excess of water. Adding to this effect, the Canary Current itself is strengthened by powerful trade winds, so the counter current does not reach to the surface (MITTELSTAEDT 1983). But during autumn the trade winds are weakest, causing the northward current to often reach the surface at the slope (KNOLL et al. 2002). Nevertheless, southward winds exist and the velocity of the surface counter current does not exceed a few cm per second, while the deeper layer reaches a velocity to 5 – 10 cm per second (MITTELSTAEDT 1983). In the water street west of the Canary Islands between the islands and the West African coast, which is called La Bocaina, the northward counter current down to 500 m is strongest with up to 9 cm per second in autumn (KNOLL et al. 2002). Counter currents may also interrupt the coastal upwellings and may be influence the circulation of the upwelling waters along the West African coast (MITTELSTAEDT 1983). These upwellings are caused by the strong trade winds blowing from the northeast parallel to the coast line (ARÍSTEGUI et al. 1997; JOHNSON and STEVENS 2000; vAN CAMP et al. 1991). At the level of the Canarian Archipelago, the winds blow parallel to the coast favouring a strong upwelling, while in the north, the winds blow onshore, keeping the upwellings weak (NAVARRO-PÉREZ and BARTON 2001; vAN CAMP et al. 1991). The winds advect the surface waters from the coast due to shear force, but the entire wind force cannot be transmitted to the deeper layers, causing the overall water flow to be directed about 45° from the wind direction (Ekman transport).Therefore, the water flows directly to the west. Due to the movement of water towards the open ocean and the decreasing depth towards the shelf, water flowing onshore is forced upwards (MITTELSTAEDT 1983), thus creating upwelling.The upwelling water reaches the surface on the shelf break or over the mid shelf and frequently occurs inshore as well (MITTELSTAEDT 1983). The upwelled waters originate at a depth of 100 – 200 m (KNOLL et al. 2002), have temperatures of 15 – 17°C and can be traced up to 200 – 300 km from the shelf break in upwelling regions north of Cape Blanc (MITTELSTAEDT 1983). Consequently, the upwelled water causes a gradient in salinity and temperature with the surrounding surface waters from the east to the west. At the level of the Canary Islands, the temperature increases due to insolation, while the salinity decreases

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1 INTRODUCTION

from east to west. The strength of these gradients rises with stronger upwelling (KNOLL et al. 2002). The area of influence can be clearly seen in Figure 1a-f with an enhanced chlorophyll production. Simultaneously with the trade winds, the upwellings show a seasonal pattern, (HERNÁNDEZ-GUERRA et al. 2002; NAVARRO-PÉREZ and BARTON 2001;VAN CAMP et al. 1991).The main determinant of the seasonal variation of the trade winds and subsequently for the intensity of the upwelling is the meridional shift of the Azores High (HERNÁNDEZ-GUERRA et al. 2002; MITTELSTADT 1983). The Azores High reaches its most northern position in summer and in autumn, resulting in the strongest winds and for the upwelling most favourable wind direction in summer. Consequently, the upwelling is most intense in summer north of 25°N (ARÍSTEGUI et al. 1997; KNOLL et al. 2002; NAVARRO-PÉREZ and BARTON 2001; vAN CAMP et al. 1991) at the Canary Islands. During the winter the upwelling is low or non existent due to weak winds and their variability in direction (BARTON et al. 1998), see Figure 1a, d and e. Even a warm onshore flow due to onshore winds may occur (VAN CAMP et al. 1991). However, despite the low intensity of the winds in winter, upwelling generally occurs throughout the year (NAVARRO-PÉREZ and BARTON 2001). Furthermore, upwelling can be observed west of the islands, depicted in Figure 1 b, c and f, with an enhanced chlorophyll production west of Gran Canaria and Fuerteventura. Upwelling varies with wind and coastal currents due to irregularities of coastline, bathymetry and influence of ocean currents (MITTELSTAEDT 1983). In general, variations in the offshore flow of up to 20 cm per second and the onshore flow from 5 – 15 cm per second can occur (MITTELSTAEDT 1983).Vertical velocities of up- and downwellings amount to approximately 50 m per day (BASTERRESTXEA et al. 2002; MITTELSTAEDT 1983). By meandering the Canary Current develops so-called upwelling filaments, which are mostly formed at the cups of the African coast and can reach more than 100 km offshore and so influence the Canary Islands directly (BARTON et al. 1998; JOHNSON and STEVENS 2000; KNOLL et al. 2002; NAVARRO-PÉREZ and BARTON 2001). The filament generated between Cap Yubi and Cape Bojador can extend to the east and southeast or towards the south of the island Gran Canaria (ARÍSTEGUI et al. 1997; BARTON et al. 1998; BASTERRESTXEA et al. 2002; vAN CAMP et al. 1991), see e.g. Figure 1d. A strong filament of semi-permanent nature is located at Cape Ghir near 31°N. It has its highest intensity in summer and autumn and reaches beyond 13°W (JOHNSON and STEVENS 2000; VAN CAMP et al. 1991). From the interaction of the meandering upwelling filaments and perturbations of the islands in the flow of the Canary Current, so-called eddies develop. These are vortices of waters and can be orientated in a clockwise (anticyclonic) or in a counterclockwise (cyclonic) direction. In the area of the Canary Islands the island induced eddies are common, especially at the island Gran Canaria (ARÍSTEGUI et al. 1997; BASTERRESTXEA et al. 2002). The island perturbs the flow of the Canary Current resulting in anticyclonic eddies to the south and cyclonic eddies to the southwest of the island (ARÍSTEGUI et al. 1997). One feature of a cyclonic eddy is its enhanced phytoplankton productivity at the periphery while at the core the amount is lower (ARÍSTEGUI et al. 1997). Cyclonic eddies bring deep nutrient rich waters to the surface in their core and move it away from the centre to the periphery, providing nourishment for phytoplankton. In older eddies the pump get weaker and the amount of chlorophyll increases at the core (ARÍSTEGUI et al. 1997). Cyclonic eddies are associated with low values of zooplankton biomass, causing further phytoplankton accumulation (ARÍSTEGUI et al. 1997). Contrarily, anticyclonic eddies submerge water at the core accumulating chlorophyll at a depth of about 150 m (ARÍSTEGUI et al. 1997). Zooplankton biomass accrues around the phytoplankton with high feeding and growth rates (HERNÁNDEZLÉON 1991). Anticyclonic eddies are therefore associated with sinking phytoplankton and picoplankton, concentrating biomass at their core (YEBRA et al. 2004)

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

The eddy generated by Gran Canaria is very common. It often spins off and can live up to several weeks, but gets weaker since no motor exists and it is influenced by wind stress (ARÍSTEGUI et al. 1997). The filament southeast of Gran Canaria sometimes ends in an eddy (BARTON et al. 1998) or interacts with the eddy of Gran Canaria, leading to an entrainment of chlorophyll rich waters into the peripheries (ARÍSTEGUI et al. 1997). Additional eddies are occur frequently, such as those generated by the outflow of Mediterranean Water. This warm and saline water spreads out as a tongue into the North Atlantic and results in a Mediterranean eddy, a so-called meddy, due to the density differences between the two water masses (JOHNSON and STEVENS 2000). Beside eddies, Gran Canaria generates a wind- and current-deprived area in the lee of the island. It is characterized by a diurnal heating cycle and a phytoplankton accumulation (BASTERRESTXEA et al. 2002). If the wind-sheltered areas heat up, a warm wake develops which can extend leeward some tens of kilometres (BASTERRESTXEA et al. 2002). In summer, a general warming of the lee occurs (VAN CAMP et al. 1991). In general, more or less motionless areas and warm wakes develop in the lee of every island, especially at islands with higher altitude (BASTERRESTXEA et al. 2002). The wake zone is a retention area for fish eggs and larvae (HERNÁNDEZ-LÉON 1991; RODRÍGUEZ et al. 2001). Not only the wakes, but all the described hydrographic features affect the distribution patterns of fish eggs and larvae. The community structure is related to the hydrography, vertically as well as horizontally (JOHN et al. 2004b; RODRÍGUEZ et al. 2001; 2004 and 2006; SASSA et al. 2002 and 2004). In general, the retention of neritic fish larvae within the neritic zone can be a problem, since larvae cannot overcome the currents and are in danger of being washed away (RODRÍGUEZ et al. 2001). Some features lead to a drift of larvae away from the favoured area, such as currents in general and along the flanks of the islands, local upwelling and migrating eddies. Other features prevent larvae from being advected out of an area. The wakes in the southwest or the dividing area of the currents north of the islands function as a retention area (RODRÍGUEZ et al. 2001). Further upwelling, and particularly upwelling filaments, influence the neritic community by averting the larvae and displace the oceanic community (RODRÍGUEZ et al. 2001). Other features, particularly eddies, even enhance larval survival (LOGERWELL and SMITH 2001). Larvae are re-circulated within eddies which provide a nursery ground (RODRÍGUEZ et al. 2001). Since eddies have a lifespan of several weeks, this would be enough for fish larvae to complete their development after which they can overcome the flow speed and return to their original habitat (RODRÍGUEZ et al. 2001). Thus, eddies prevent larvae from being averted by the Canary Current and provide better feeding conditions increasing larval survival (LOGERWELL and SMITH 2001).

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

2 MATERIAL AND METHODS

2.1. SURVEYS AND SAMPLING METHOD In the period from 1997 – 2002 six cruises were undertaken around the Canarian Archipelago at different seasons during the day as well as during the night and at different habitats and depths, respectively. The cruises were carried out by the Canarian governmental nautic school vessel B/E “La Bocaina”, with a length of 29.7 m, a width of 8 m and a draught of 3.1 m, were its brutto weight is registered as 205 tons (BORDES et al. 1999). All six cruises resulted in catching fish larvae, “La Bocaina 11/97”, “ECOS 04/99”, “Mesopelagic 05/99”, “Pelagic 01/00”, “Pelagic 11/00” and “La Bocaina 03/02”. All cruises with its stations are shown in Figure 2 –Figure 6. Commercial fisheries trawls were used with a mesh size of 2 mm and the opening of the net was stabilized by a steel ring of 1.5 m in diameter. The trawls were hauled in a horizontally orientation in different oceanic habitats, where the hauls were proceeded above bottom depths from 40 m – 3400 m. In waters with a bottom depth exceeding 200 m, trawls were categorized according to the common oceanic habitat classification into epipelagic hauls, which ranges from 0 – 200 m depth, and mesopelagic hauls, which ranges from 200 – 1000 m depth. Trawls undertaken at bottom depths shallower than 200 m were grouped as neritic. It has to be taken into consideration that trawls having the start within a certain habitat but during process reach into another area were accounted according to their main trawling time within a certain realm. Details to the single stations as date, initial and final position trawling and bottom depths are listed in Table 1 –Table 6. Since the net had no opening-closing devices, the avoidance of catching fish and fish larvae of an undesirable depth while descending and ascending the net to and from the designated trawling depth, had to be done by manipulating the speed of the research vessel. When descending the net, the speed of the ship was lowered to a level below the speed of the releasing cable. So the net was standing vertically in the water column and the opening was closed. In the intended depth the ship fastened the velocity, the net was brought into a horizontal position and the gear opened. While processing the opening was kept unclosed due to weights at the lower and floating devises at the upper part of the ring. During the trawl a net sounder positioned at the anterior end of the net transmitted data of depth, net opening diameter and speed every five minutes to the bridge, so that the depth and diameter of the net could be controlled by changing the velocity of the vessel. For ascending the trawl was closed due to a sudden stop of the ship.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

2 MATERIAL AND METHODS

Table 1 Characteristics of the trawl tows from the cruise “La Bocaina 11/97”. Station numbers (St. No.) are given. The habitat, trawls were taken in, is labelled in the column with “H”. n refers to the neritic, e to the epipelagic and m to the mesopelagic habitat. St.No.

Date [1997]

Time

Initial Latitude

Final Longitude

Latitude

Longitude

Trawl depth

Bottom depth

[range, in m]

09.11

20:35-21:35

28°07,46’ N

14°02,48’ W

28°05,96’ N

14°05,92’ W

71-78

1360-1413

10.11

17:20-18:18

28°06,38’ N

13°57,83’ W

28°05,17’ N

14°00,42’ W

500-523

1067-1390

10.11

20:12-21:12

28°10,04’ N

14°09,46’ W

28°09,72’ N

14°04,70’ W

34-69

617-1457

10.11

11:50-12:24

28°10,23’ N

14°05,19’ W

28°10,02’ N

14°07,18’ W

22-35

49-85

12.11

06:40-07:10

28°02,50’ N

14°26,28’ W

28°02,21’ N

14°24,23’ W

40-44

130-530

13.11

06:25-06:52

28°51,55’ N

13°54,29’ W

28°50,16’ N

13°54,51’ W

28-43

656-860

14.11

14:15-15:15

28°55,23’ N

13°34,57’ W

28°54,25’ N

13°38,19’ W

13-42

128-217

15.11

10:20-11:00

28°58,67’ N

13°28,74’ W

28°56,98’ N

13°30,11’ W

53-83

101-120

15.11

19:25-20:25

29°23,12’ N

13°12,23’ W

29°20,37’ N

13°11,96’ W

527-557

1280-1350

16.11

00:10-01:23

29°20,42’ N

13°21,88’ W

29°16,58’ N

13°23,92’ W

20-64

94-144

16.11

03:05-04:05

29°22,50’ N

13°33,36’ W

29°19,72’ N

13°33,59’ W

34-46

586-692

16.11

23:25-00:35

28°50,06’ N

13°53,01’ W

28°47,27’ N

13°52,11’ W

31-43

70-115

18.11

06:05-07:05

28°07,95’ N

14°01,63’ W

28°06,34’ N

14°04,23’ W

90-107

1183-1442

20.11

19:45-20:45

27°38,30’ N

15°37,64’ W

27°36,98’ N

15°39,98’ W

592-603

1638-1667

20.11

23:50-00:50

27°40,99’ N

15°44,21’ W

27°43,36’ N

15°46,66’ W

28-63

116-344

21.11

21:20-22:20

28°07,23’ N

15°47,59’ W

28°05,07’ N

15°50,17’ W

42-66

104-112

23.11

22:08-23:08

27°50,20’ N

15°19,27’ W

27°52,80’ N

15°18,27’ W

30-52

136-231

24.11

00:25-01:00

27°51,58’ N

15°18,71’ W

27°49,87’ N

15°18,71’ W

116-146

181-260

Table 2 Characteristics of the trawl tows from the cruise “ECOS 04/99”. Station numbers (St. No.) are given. The habitat, trawls were taken in, is labelled in the column with “H”. n refers to the neritic, e to the epipelagic and m to the mesopelagic habitat. Sat.

Date

No.

[1999]

Time

Initial Latitude

Final Longitude

Latitude

Trawl depth

Bottom depth

Longitude

[range, in m]

08.04

19:23-20:50

28°51,75’ N

13°44,02’ W

28°49,01’ N

13°46,02’ W

28-39

117-220

09.04

21:17-22:00

28°50,30’ N

13°51,52’ W

28°47,30’ N

13°52,67’ W

8-39

79-99

10.04

05:13-06:00

28°49,69’ N

13°52,47’ W

28°47,29’ N

13°52,89’ W

25-39

98-112

10.04

21:28-23:30

28°10,07’ N

14°00,31’ W

28°09,57’ N

14°07,11’ W

19-50

102-695

11.04

09:45-11:40

28°07,29’ N

14°03,65’ W

28°05,62’ N

14°09,88’ W

382-511

1264-1351

12.04

22:13-23:41

28°05,76’ N

14°03,85’ W

28°04,43’ N

14°08,34’ W

23-52

1400-1544

13.04

21:51-22:55

28°08,43’ N

14°12,47’ W

28°05,15’ N

14°15,71’ W

19-40

67-314

14.04

05:38-06:45

28°07,85’ N

14°02,32’ W

28°06,33’ N

14°06,65’ W

47-67

1254-1359

14.04

21:20-22:15

28°49,61’ N

13°45,97’ W

28°46,41’ N

13°47,25’ W

18-33

87-165

n m

15.04

10:35-12:25

28°49,82’ N

13°37,89’ W

28°45,50’ N

13°41,62’ W

258-600

1031-1077

15.04

15:02-15:55

28°51,30’ N

13°41,63’ W

28°48,16’ N

13°43,95’ W

12-40

756-770

22.04

22:35-23:25

27°42,78’ N

15°49,74’ W

27°39,77’ N

15°48,58’ W

38-49

1145-1315

23.04

01:05-02:05

27°46,05’ N

15°48,70’ W

27°44,15’ N

15°46,49’ W

33-53

82-90

23.04

16:55-18:40

27°38,99’ N

15°39,43’ W

27°36,21’ N

15°43,42’ W

480-694

1069-1384

23.04

21:50-22:45

27°43,08’ N

15°43,70’ W

27°41,13’ N

15°42,58’ W

24-40

57-115

24.04

00:00-01:20

27°43,78’ N

15°47,00’ W

27°41,37’ N

15°43,46’ W

18-61

100-123

25.04

22:40-23:45

28°11,38’ N

16°50,51’ W

28°07,53’ N

16°47,98’ W

25-47

74-132

26.04

01:01-02:09

28°07,60’ N

16°46,16’ W

28°10,95’ N

16°50,31’ W

45-71

75-176

26.04

12:10-13:50

28°10,71’ N

16°53,77’ W

28°05,89’ N

16°51,04’ W

553-716

1213-1409

30.04

07:06-08:00

28°04,60’ N

17°21,20’ W

28°02,37’ N

17°19,43’ W

34-52

90-107

30.04

17:13-18:20

28°18,98’ N

17°06,61’ W

28°18,09’ N

17°03,89’ W

255-535

944-1283

30.04

21:52-23:50

28°06,84’ N

17°23,31’ W

28°02,36’ N

17°19,96’ W

25-70

110-118

01.05

21:15-23:10

28°41,90’ N

17°59,46’ W

28°34,61’ N

17°56,36’ W

18-42

96-784

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

Table 3. Characteristics of the trawl tows from the cruise “Mesopelagic 05/00”. Station numbers (St. No.) are given. The habitat, trawls were taken in, is labelled in the column with “H”. n refers to the neritic, e to the epipelagic and m to the mesopelagic habitat. Sat.

Date

No.

[1999]

1 2 3 4 5 6

11.05 11.05 11.05 12.05 12.05 12.05

Time 05:55-07:05 08:35-09:40 22:15-23:15 09:05-10:05 21:35-22:35 23:45-00:45

Initial

Final

Trawl depth

Bottom depth

Latitude

Longitude

Latitude

Longitude

[range, in m]

28°39,50’ N 28°34,57’ N 28°49,29’ N 28°48,30’ N 29°04,05’ N 29°02,45’ N

13°26,49’ W 13°31,57’ W 15°22,68’ W 15°23,67’ W 16°56,10’ W 16°58,18’ W

28°36,90’ N 28°31,85’ N 28°46,86’ N 28°46,40’ N 29°02,46’ N 29°00,60’ N

13°29,24’ W 13°34,12’ W 15°24,52’ W 15°25,80’ W 16°58,30’ W 17°00,20’ W

38-40 495-561 42-47 517-591 448-526 48-66

1327-1349 1287-1306 1646-1782 1940-2525 1660-2120 1463-2119

H e m e m m e

Table 4 Characteristics of the trawl tows from the cruise “Pelagic 01/00”. Station numbers (St. No.) are given. The habitat, trawls were taken in, is labelled in the column with “H”. n refers to the neritic, e to the epipelagic and m to the mesopelagic habitat. Sat.

Date

No.

[2000]

1 2 3 4

19.01 19.01 22.01 22.01

Time 18:30-19:55 22:10-23:10 14:35-15:50 23:15-23:45

Initial

Final

Trawl depth

Bottom depth

Latitude

Longitude

Latitude

Longitude

[range, in m]

27°56,97’ N 28°00,05’ N 28°12,13’ N 28°11,49’ N

15°16,52’ W 15°20,45’ W 15°31,58’ W 15°38,13’ W

28°00,37’ N 28°03,48’ N 28°12,62’ N -

15°18,07’ W 15°21,56’ W 15°35,50’ W -

467-566 17-50 356-402 37-66

765-845 105-294 949-1455 60-122

m n m n

25.01

16:25-18:00

27°41,51’ N

15°31,39’ W

27°40,27’ N

15°34,77’ W

490-510

950-1260

6 7 8 9 10

25.01 26.01 26.01 27.01 27.01

20:21-21:15 15:20-16:40 20:15-21:35 15:25-16:55 20:15-20:45

27°44,58’ N 28°04,16’ N 28°05,24’ N 27°49,08’ N 27°41,65’ N

15°24,40’ W 15°55,10’ W 15°49,56’ W 15°54,01’ W 15°42,91’ W

27°43,82’ N 28°02,39’ N 28°02,71’ N 27°51,64’ N 27°42,36’ N

15°28,42’ W 15°56,49’ W 15°53,57’ W 15°56,73’ W 15°44,61’ W

10-70 578-610 40-45 493-503 25-34

76-186 1239-1644 101-280 767-1096 75-86

n m n m n

27.01

21:45-22:45

27°41,82’ N

15°43,27’ W

27°43,23’ N

15°46,05’ W

41-64

75-92

Table 5 Characteristics of the trawl tows from the cruise “Pelagic 11/00”. Station numbers (St. No.) are given. The habitat, trawls were taken in, is labelled in the column with “H”. n refers to the neritic, e to the epipelagic and m to the mesopelagic habitat. Sat.

Date

No.

[2000]

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18

10.11 12.11 12.11 14.11 14.11 15.11 15.11 17.11 19.11 19.11 19.11 20.11 20.11 21.11 21.11 21.11 21.11 22.11

22.11

Time

Initial

Final

Trawl depth

Bottom depth

Latitude

Longitude

Latitude

Longitude

[range, in m]

20:38-21:28 18:28-19:17 21:59-22:46 11:00-11:52 22:33-23:13 21:20-22:03 23:04-23:53 17:21-18:39 11:00-12:14 13:28-14:38 18:24-20:04 18:28-19:35 21:16-22:18 12:08-13:23 15:01-16:01 18:36-19:10 21:30-22:34 11:53-12:53

28°10,69’ N 28°03,04’ N 28°03,38’ N 27°59,53’ N 27°44,96’ N 27°41,04’ N 27°42,21’ N 27°38,12’ N 27°37,49’ N 27°40,32’ N 27°41,39’ N 28°02,70’ N 28°03,32’ N 27°37,21’ N 27°36,26’ N 27°33,81’ N 27°37,91’ N 27°36,13’ N

15°31,73’ W 15°21,37’ W 15°20,70’ W 15°20,53’ W 15°26,21’ W 15°42,19’ W 15°44,79’ W 15°47,02’ W 15°45,05’ W 15°48,57’ W 15°43,84’ W 15°56,76’ W 15°51,38’ W 15°39,71’ W 15°40,36’ W 15°21,66’ W 15°38,55’ W 15°48,77’ W

28°10,60’ N 28°00,52’ N 28°00,94’ N 27°57,86’ N 27°44,22’ N 27°41,85’ N 27°43,75’ N 27°39,52’ N 27°39,54’ N 27°41,92’ N 27°44,76’ N 28°01,24’ N 27°59,90’ N 27°35,82’ N 27°37,30’ N 27°32,73’ N 27°36,44’ N 27°35,26’ N

15°34,48’ W 15°20,28’ W 15°20,73’ W 15°21,14’ W 15°28,23’ W 15°44,83’ W 15°46,99’ W 15°50,05’ W 15°47,82’ W 15°51,22’ W 15°48,54’ W 15°58,53’ W 15°53,30’ W 15°42,30’ W 15°38,11’ W 15°19,18’ W 15°41,52’ W 15°46,87’ W

55-90 17-45 43-90 37-64 20-48 37-64 47-71 434-606 622-813 310-530 24-57 444-900 37-46 495-1009 592-637 29-31 31-37 464-924

132-853 103-263 98-414 102-240 59-75 98-202 92-112 927-1653 928-1287 1174-1650 98-228 1634-1765 88-117 1344-1620 1316-1688 2189-2438 1305-1603 1440-1690

e n e n n n n m m m n m n m m e e m

14:50-15:55

27°36,01’ N

15°46,38’ W

27°37,05’ N

15°48,94’ W

496-630

1358-1564

Inf. Téc. Inst. Canario Cienc. Mar. n°13

2 MATERIAL AND METHODS

Table 6 Characteristics of the trawl tows from the cruise “La Bocaina 03/02”. Station numbers (St. No.) are given. The habitat, trawls were taken in, is labelled in the column with “H”. n refers to the neritic, e to the epipelagic and m to the mesopelagic habitat. Sat.

Date

No.

[2002]

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27

07.03 08.03 09.03 09.03 10.03 10.03 11.03 12.03 13.03 13.03 14.03 14.03 14.03 14.03 15.03 15.03 15.03 15.03 15.03 16.03 16.03 16.03 17.03 17.03 17.03 17.03 18.03

Time 22:34-23:52 19:54-20:59 17:35-19:00 21:55-23:03 00:10-01:12 10:39-11:05 21:20-21:54 20:35-21:35 13:18-14:00 16:18-17:20 10:05-10:40 14:01-15:05 16:56-18:20 21:30-22:00 10:55-11:30 12:52-13:30 15:06-15:45 16:50-17:45 21:09-21:45 13:18-14:35 20:10-20:52 22:14-22:50 00:24-01:05 02:47-03:40 19:56-20:48 22:03-23:35 00:46-02:00

Initial

Final

Latitude

Longitude

28°06,91’ N 28°09,49’ N 28°04,15’ N 28°08,22’ N 28°04,46’ N 28°08,50’ N 28°09,89’ N 28°08,54’ N 28°09,46’ N 28°07,78’ N 28°07,30’ N 28°04,74’ N 28°04,17’ N 28°08,76’ N 28°04,08’ N 28°03,70’ N 28°02,72’ N 28°03,37’ N 28°03,69’ N 28°03,42’ N 28°11,42’ N 28°03,20’ N 28°03,08’ N 28°02,86’ N 28°10,48’ N 28°04,36’ N 28°04,56’ N

14°13,87’ W 14°00,21’ W 14°14,83’ W 14°03,44’ W 14°14,80’ W 14°02,93’ W 14°01,28’ W 14°06,45’ W 14°10,29’ W 14°06,21’ W 14°01,00’ W 14°01,90’ W 14°00,61’ W 14°12,22’ W 14°09,47’ W 14°09,75’ W 14°10,50’ W 14°09,79’ W 14°09,38’ W 14°15,01’ W 13°59,82’ W 14°10,10’ W 14°10,24’ W 14°10,33’ W 14°02,56’ W 13°58,91’ W 13°59,73’ W

Latitude 28°03,95’ N 28°10,33’ N 27°59,83’ N 28°07,31’ N 28°02,18’ N 28°07,89’ N 28°09,79’ N 28°07,36’ N 28°09,23’ N 28°08,34’ N 28°07,29’ N 28°03,75’ N 28°02,22’ N 28°07,43’ N 28°02,02’ N 28°01,63’ N 28°00,98’ N 28°05,29’ N 28°01,49’ N 28°00,26’ N 28°09,35’ N 28°01,27’ N 28°00,83’ N 28°00,79’ N 28°09,90’ N 28°03,31’ N 28°04,00’ N

Longitude 14°16,75’ W 14°03,85’ W 14°17,44’ W 14°07,31’ W 14°17,21’ W 14°05,88’ W 14°03,55’ W 14°08,74’ W 14°07,76’ W 14°03,69’ W 14°03,49’ W 13°59,02’ W 13°57,74’ W 14°13,26’ W 14°11,03’ W 14°11,07’ W 14°12,46’ W 14°08,10’ W 14°11,04’ W 14°16,79’ W 14°01,68’ W 14°11,37’ W 14°11,73’ W 14°11,80’ W 13°59,80’ W 14°03,46’ W 14°03,56’ W

Trawl depth

Bottom depth

[range, in m]

24-60 13-54 440-665 22-130 34-123 617-915 98-219 269-529 19-31 429-572 31-140 323-570 632-1035 15-43 30-99 185-211 311-378 470-695 30-139 308-498 15-33 215-261 310-427 565-980 20-33 25-62 310-525

140-284 217-606 1107-1327 1000-1211 747-1006 980-1162 137-464 656-1026 40-77 959-1198 1246-1347 1406-1447 1447-1529 96-211 1439-1677 1461-1680 1000-1690 1364-1491 1462-1672 1262-1592 43-570 1499-1701 1505-1750 1523-1750 51-529 1374-1577 1364-1548

H n e m e e m e m n m e m m n e e m m e m e m m m e e m

Figure 2. Map of all trawls of the cruise „La Bocaina 11/97”. Station numbers refer to Table 1 (map from WIENERROITHER).

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

Figure 3. Map of all trawls of the cruises „ECOS 04/99” and “Mesopelagic 05/99”. Station numbers refer to Table 2 and Table 3 (map from WIENERROITHER).

Figure 4. Map of all trawls of the cruise „Pelagic 01/00”. Station numbers refer to Table 4 (map from WIENERROITHER).

Inf. Téc. Inst. Canario Cienc. Mar. n°13

2 MATERIAL AND METHODS

Figure 5. Map of all trawls of the cruise „Pelagic 11/00”. Station numbers refer to Table 5 (map from WIENERROITHER).

Figure 6. Map of all trawl of the cruise „La Bocaina 03/02”. Station numbers refer to Table 6 (map from WIENERROITHER).

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

2.2. MATERIAL TREATMENT AND TAXONOMIC APPROACH The material caught at the stations of the six successful cruises was first fixated in 7 % formalin and then transferred to 70 % ethanol. The fish larvae were sorted out of the material and were incorporated and stored, respectively, at the fish larvae collection of the Zoological Museum and University Copenhagen (ZMUC), Denmark. The specimens were identified to the lowest taxonomic level possible with the support from Dr. Hans-Chrisitan John at the Zoological Institute and Museum (ZIM), Hamburg and with identification literature and guides including BERTOLINI et al. (1956), Dana Report Nr. 40 (1953), FAHAY (1983), Moser et al. (1984), MOSER (1996), MOSER and WATSON (2001), LEIS and RENNIS (1983), OLIVAR and FORTUÑO (1991), RICHARDS (2006), and WHITEHEAD et al. (1984-86). From the latter two references only single chapters were used and cited throughout the entire annotated larvae taxa list. Due to the relatively low sample size, meristic counts did often match only with fractions of the entire range mentioned in the literature and hence the latter was provided thus allowing also a wider use of the diagnostic characters examined. The reference numbers assigned by the Zoological Museum, University of Copenhagen (ZMUC) are provided for each taxon and the respective collection reference is given at the end of the annotated larvae taxa list. In the taxonomic list (Table 7) all collected specimens of all developing stages are listed with habitat of origin and total number of specimens per taxon. The larvae taxa list was ordered according to NELSON (2006) and the validity of scientific genus and species nomenclature were cross-checked with ESCHMEYER (1998), the Eschmeyer’s online checklist (http:// www.calacademy.org/research/ichthyology/) and the online catalogue of fishes (http://www.fishbase.com). The habitat of the adult fish (habitat of origin) is specified after WHITEHEAD et al. (1986) and simplified to the classification of neritic, bathydemersal, epipelagic, deep pelagic and benthopelagic habitats of origin. Within the neritic species, coastal pelagic as well as littoral-demersal species are summarized (bottom depth is not deeper than 200 m). The epipelagic species inhabit the upper 200 m of the water column in the oceanic realm and within the deep pelagic species, meso- as well as bathypelagic species are combined (all pelagic species living deeper than 200 m). Bathydemersal species imply slope and deep bottom dwelling species as well as deep bottom associated species (with a bottom depth of more than 200 m). The classification benthopelagic is used, if species live bathydemersal as well as epi- or deep pelagic. In the annotated larval list, specimens in late transforming, juvenile or even adult stages were excluded. In this list the larval development and their important diagnostic characters of the orders, families and the lowest identified taxonomic level for each stage are described. Orders and families are well described since the pre-sorting of larvae should follow this order. For a further identification the genera and species description is very detailed and of crucial importance within each taxonomic level is the combination of all available diagnostic characters. All these characters are listed in following order in species description after diagnostic importance, starting with meristic (numbers of vertebrae, myomeres, fin rays etc.) and development information, like the size of hatching, flexion (the upwards bending of the notochord tip forming the caudal fin base) and transformation (loosing larval characters, complete the development of all adult characters and migrating to the adult habitat). Many meristic characters and development information are missing due to a lack of knowledge during the larval development in the literature. More descriptive information is given for the morphological as well as for the pigmentation development, since both are essential for larval identification. Within the certain families e.g. Myctophidae the photophore development is described, if required for identification. All literature references are listed per taxon description given. Within the reference material the number of individuals per taxon, the station number (S) and the individual collection numbers are listed. Also the seasonal and spatial distribution is described. Specimens, which reached a late juvenile or even adult stage are not mentioned and described in the annotated larvae taxa list. The leptocephalis are only described to the family level, since no further identification was possible due to their bad condition. E.g. myomeres were not countable since they are smashed. Due to the impossibility of identification down to a lower level all leptocephali are excluded from all further analyses.

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2 MATERIAL AND METHODS

2.3. MULTIVARIATE COMMUNITY ANALYSIS Multivariate analyses were performed with the software package PRIMER (Plymouth Routines Multivariate Ecological Research). For analysing the community structure the numbers of larvae were standardized to a trawling time of one hour. A Bray – Curtis rank similarity matrix was calculated using root-root transformed data. The root-root transformation has the advantage of scaling the abundant species down and that the similarity coefficient is invariant to a scale change (FIELD et al.1982). Additionally, the Bray Curtis similarity weighs rare species lower in comparing samples and avoids the effect of rare species (FIELD et al. 1982). The similarities between species composition were performed by a hierarchical agglomerative clustering with group average linking and multi – dimensional scaling (MDS), where the dissimilarities are directly proportional to the distance between the interpoints (FIELD et al. 1982). MDS was performed based on different factors, to investigate the effect of seasonal, regional and spatial conditions on the species assemblage. A one-way analysis of similarity (ANOSIM) was applied to determine the significance of trends within the different factors, used already in the preceding analysis. During all calculations only species with a relative abundance of more than 0.2 % of the total catch were included to avoid the effect of rare species (RAMOS et al. 2006). Additionally, specimens in a juvenile or even adolescence development stage were excluded to evade the ontogenetically different behaviour. To deal with the effect of different taxonomical levels all specimens of higher taxonomic level than species level were treated as species, as long as they definitively represent another species. Consequently, all damaged larvae were excluded, since it could not be assured being another species as already identified.

2.4. ANOVA ANALYSIS For investigating the differences in the abundance and diversity of fishes a one factorial ANOVA with the software STATISTICA was performed. The Shannon Index H’, the Pielou’s Index of Evenness J’, the Simpson’s diversity Index D, the total number of species S and the total number of individuals N were grouped to spatial and seasonal factors. Whisker plots with a confidence interval of +/- 95% (Standard error only used at the total number of individuals) indicate significant differences. Before running the ANOVA the normal distribution was tested by using the Kolmogorov-Smirnov & Lillefors (Prof. WAHL M. and Dr. LENZ M., pers. comment). The Levene’s test was performed to check for homogeneity and in case of heterogeneity the data were transformed with ArcSin to buffer the variances (Prof. WAHL M. and Dr. LENZ M., pers. comment). Does the ANOVA show significant differences, a post-hoc test (Bonferroni test) was used to check between which factors the differences are significant (Prof. WAHL M. and Dr. LENZ M., pers. comment).

2.5. GEOGRAPHICAL INFORMATION SYSTEM (GIS) DISTRIBUTION MAPS For illustrating the distribution of the species inhabiting the oceanic and neritic areas during their larval life, maps were created with the GIS program MapInfo Professional 7.5. Information about the habitat of origin of different species was taken from the previous already mentioned identification literature, especially from WHITEHEAD (1986). The habitat types of origin are listed in Table 7. Abundances are indicated by different coloured and sized circles described in the particular maps.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

3 RESULTS

3.1. FISH LARVAE TAXONOMIC COMPOSITION From the six successful cruises a number of 57 trawls resulted in capturing fish larvae, where 19 were epi- and 25 mesopelagic and 13 neritic trawls. In total, 4622 specimens of fish larvae and 466 leptocephali were collected, belonging to 54 families. 47 families and 77 genera could have been distinguished within the fish larvae, but only 70 species. Some specimens could not be identified to lower taxonomic levels than to family level due to their bad condition or larvae were not allocated to adult fish yet. In total, the minimum number of species and lower taxa, which represent one species within the fish larvae, although they could not be identified to species level, were 92. Within the leptocephali 12 specimens belong to the order of Albuliformes, where the larvae could be identified only to the level of the suborder Notacanthoidei. All other leptocephali belong to 7 families within the order of Anguilliformes, whereas two specimens could not be allocated to a family and none of the other leptocephalis could be identified to genus or species level. The most common family was Engraulidae with its only representative Engraulis encrasicolus comprising 63.3 % of all sampled larvae.The second most common family was the Gobiidae with 19.8 %. Except for the Gonostomatidae (3.8 %), Myctophidae (2.9 %), Synodontidae (1.8 %), Phosichthyidae (1.3 %), Paralepididae (1.1 %) and Scombridae (1.1 %) all other families were represented with a percentage lower than 1 %.The proportion of neritic larvae was highest with a percentage of 87.0 % and bathydemersal larvae were represented only with 1.3 %. Epipelagic larvae made up 1.6 % and deep pelagic larvae 10.1 % of the total catch. Beside the dominating Engraulidae and Gobiidae the community was very diverse with the most diverse family being the Myctophidae.

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3 RESULTS

Table 7 Taxonomic list of all specimens caught during the cruises “La Bocaina 11/97”, “ECOS 04/99”, “Mesopelagic 05/99”, “Pelagic 01/00”, “Pelagic 11/00” and “La Bocaina 03/02”; their habitat of origin according to WHITEHEAD et al. (1986) and total number are listed; systematic order according to NELSON (2006). The new records for the Canary Islands are written in bold letters. Taxon Order Albuliformes (Suborder Notacanthoidei) Unidentified Order Anguilliformes Unidentified Family MURAENIDAE Unidentified Family OPHICHTHIDAE Unidentified Family NEMICHTHYIDAE Unidentified Family CONGRIDAE Unidentified Family NETTASTOMATIDAE Unidentified Family SERRIVOMERIDAE Unidentified Family EURYPHARYNGIDAE Unidentified Family ENGRAULIDAE Engraulis encrasicolus (Linnaeus, 1758) Family CLUPEIDAE Sardinella spp. Valenciennes, 1847 Family ARGENTINIDAE Argentina sphyraena Linnaeus, 1758 Glossanodon leioglossus (Valenciennes, 1848) Family Microstomatidae Bathylagus sp. Günther, 1878 Dolicholagus longirostris (Maul, 1948) Family PLATYTROCTIDAE Sagamichthys schnakenbecki (Krefft, 1953) Family GONOSTOMATIDAE Cyclothone sp. Goode & Bean, 1883 Cyclothone acclinidens Garman, 1899 Cyclothone alba Brauer, 1906 Gonostoma denudatum Rafinesque, 1810 Family STERNOPTYCHIDAE Maurolicus muelleri (Gmelin, 1789) Family PHOSICHTHYIDAE Polymetme corythaeola (Alcock, 1898) Vinciguerria sp. Jordan & Evermann, 1896 Vinciguerria attenuata (Cocco, 1838) Vinciguerria nimbaria (Jordan & Williams, 1895) Vinciguerria poweriae (Cocco, 1838) Family STOMIIDAE Astronesthinae Chauliodus sp. Bloch & Schneider, 1801 Idiacanthus fasciola Peters, 1877 Melanostomiinae Stomias spp. Cuvier, 1816 Stomias boa (Risso, 1810)

Original habitat

Total Number

neritic to bathydemersal

neritic

deep pelagic

neritic to bathydemersal

415

neritic to bathydemersal

deep pelagic

neritic

2925

neritic

neritic to bathydemersal bathydemersal

2 9

deep pelagic deep pelagic

1 1

deep pelagic

deep pelagic deep pelagic deep pelagic deep pelagic

1 1 1 172

deep pelagic

benthopelagic deep pelagic deep pelagic deep pelagic deep pelagic

1 1 32 23 3

deep pelagic deep pelagic deep pelagic deep pelagic deep pelagic deep pelagic

3 4 1 1 2 1

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SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

Family AULOPIDAE Aulopus filamentosus (Bloch, 1792)

neritic to bathydemersal

Synodus spp. Scopoli, 1777

neritic

Synodus saurus (Linnaeus, 1758)

neritic

Synodus synodus (Linnaeus, 1758)

neritic

neritic to bathydemersal

Ahliesaurus berryi Bertelsen, Krefft & Marshall, 1976

deep pelagic

Scopelosaurus spp. Bleeker, 1860

benthopelagic

Family SYNODONTIDAE

Family CHLOROPHTHALMIDAE Chlorophthalmus agassizii Bonaparte, 1840 Family NOTOSUDIDAE

Family SCOPELARCHIDAE Benthalbella infans Zugmayer, 1911

deep pelagic

Scopelarchus analis (Brauer, 1902)

deep pelagic

Unidentified

deep pelagic

Alepisaurus ferox Lowe, 1833

epi- to deep pelagic

Omosudis lowii Günther, 1887

deep pelagic

1 49

Family ALEPISAURIDAE

Family PARALEPIDIDAE Lestidiops spp. Hubbs, 1916

deep pelagic

Lestidiops sphyrenoides (Risso, 1820)

deep pelagic

Macroparalepis spp. Ege, 1933

deep pelagic

Sudis hyalina Rafinesque, 1810

deep pelagic

Family MYCTOPHIDAE Benthosema suborbitale (Gilbert, 1913)

deep pelagic

Centrobranchus nigroocellatus (Günther, 1873)

deep pelagic

Ceratoscopelus maderensis (Lowe, 1839)

deep pelagic

Ceratoscopelus warmingii (Lütken, 1892)

deep pelagic

Diaphus spp. Eigenmann & Eigenmann, 1890

deep pelagic

Diogenichthys atlanticus (Tåning, 1928)

deep pelagic

Hygophum hygomii (Lütken, 1892)

deep pelagic

Hygophum reinhardtii (Lütken, 1892)

deep pelagic

Lampadena sp. Goode & Bean, 1893

deep pelagic

Lampanyctus sp. Bonaparte, 1840

deep pelagic

Lampanyctus crocodilus (Risso, 1810)

deep pelagic

Myctophum nitidulum Garman, 1899

deep pelagic

Myctophum punctatum Rafinesque, 1810

deep pelagic

Nannobrachium atrum (Tåning, 1928)

deep pelagic

Notoscopelus spp. Günther, 1864

deep pelagic

Notoscopelus resplendens (Richardson, 1845)

deep pelagic

57 12

Symbolophorus veranyi (Moreau, 1888)

deep pelagic

Taaningichthys minimus (Tåning, 1928)

deep pelagic

Unidentified

deep pelagic

epi- to deep pelagic

deep pelagic

benthopelagic

deep pelagic

Family LAMPRIDAE Lampris guttatus (Brünnich, 1788) Family REGALECIDAE Regalecus glesne Ascanius, 1772 Family MACROURIDAE Macrourinae Family MELANONIDAE Melanonus zugmayeri Norman, 1930 Family MELAMPHAIDAE Melamphaes spp. Günther, 1864

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3 RESULTS

Family DIRETMIDAE Diretmus argenteus Johnson, 1864 Family TRACHICHTHYIDAE

deep pelagic

Hoplostethus spp. Cuvier, 1829

benthopelagic

Hoplostethus mediterraneus Cuvier, 1829

benthopelagic

neritic to epipelagic

neritic to bathydemersal neritic

2 5

neritic

deep pelagic

neritic neritic to bathydemersal

1 4

neritic to bathydemersal

neritic

Microichthys coccoi Rüppell, 1852 Family CARANGIDAE

deep pelagic

Campogramma glaycos (Lacepède, 1801) Trachurus spp. Rafinesque, 1810 Family SPARIDAE

benthopelagic benthopelagic

6 12

Dentex spp. Cuvier, 1814 Pagrus pagrus (Linnaeus, 1758) Family POMACENTRIDAE

neritic neritic

4 1

Chromis sp. Cuvier, 1814 Abudefduf luridus (Cuvier, 1830) Family LABRIDAE

neritic neritic

1 2

Thalassoma pavo (Linnaeus, 1758) Family CHIASMODONTIDAE

neritic

deep pelagic

neritic

neritic to bathydemersal neritic to bathydemersal neritic neritic neritic

30 2 288 590 4

Diplospinus multistriatus Maul, 1948 Family TRICHIURIDAE

deep pelagic

Benthodesmus elongatus (Clarke, 1879) Lepidopus caudatus (Euphrasen, 1788) Family SCOMBRIDAE

benthopelagic benthopelagic

1 6

Scomber colias Gmelin, 1789 Family NOMEIDAE

epi- to deep pelagic

Cubiceps gracilis (Lowe, 1843) Family CAPROIDAE Capros aper (Linnaeus, 1758)

epi- to deep pelagic

neritic to bathydemersal

Family SYNGNATHIDAE Entelurus aequoreus (Linnaeus, 1758) Family SCORPAENIDAE Helicolenus dactylopterus (Delaroche, 1809) Scorpaena spp. Linnaeus, 1758 Family MORONIDAE Dicentrarchus spp. Gill, 1860 Family PERCICHTHYIDAE Howella brodiei Ogilby, 1899 Family SERRANIDAE Serranus sp. Cuvier, 1816 Anthias anthias (Linnaeus, 1758) Family Callanthiidae Callanthias ruber (Rafinesque, 1810) Family APOGONIDAE Apogon imberbis (Linnaeus, 1758) Family EPIGONIDAE

Unidentified Family BLENNIIDAE Parablennius spp. Ribeiro, 1915 Family GOBIIDAE Crystallogobius linearis (Düben, 1845) Lesueurigobius heterofasciatus Maul, 1971 Unidentified Type I Unidentified Type II Unidentified Family GEMPYLIDAE

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SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

Family BOTHIDAE Arnoglossus spp. Bleeker, 1862 Arnoglossus imperialis (Rafinesque, 1810) Arnoglossus rueppelii (Cocco, 1844) Bothus podas (Delaroche, 1809) Family BALISTIDAE Balistes capriscus Gmelin, 1789 Family TETRAODONTIDAE Canthigaster capistrata (Lowe, 1839) Sphoeroides spp. Anonymous, 1798

neritic to bathydemersal neritic to bathydemersal neritic to bathydemersal neritic to bathydemersal

9 4 1 16

neritic

neritic neritic

1 8

3.2. ANNOTATED LARVAE TAXA LIST

¬ Albuliformes NOTACANTHOIDEI In the leptocephali of the Notacanthoidei the most important character are the V-shaped myomeres and instead of a normal caudal fin a post-caudal filament is present. These two diagnostic characters in combination is leading to this suborder, but it should be verified by the further characters. The dorsal and pelvic fins are present and the pectoral fin is small. The gut is strait and subterminal. The pigmentation consists of a ventral series of melanophores and sometimes a row below the midlateral level.

-- Reference: CASTLE (1984);

¬ Anguilliformes In most families is the body slender.The number of myomeres can reach up to 290 and are W-shaped.The pectoral fins are reduced or lost.The leptocephalis have a leaf-like body shape and are almost transparent except for the eyes and the pigmentation. The head is small and the teeth are directed forward.Two developmental larval stages are known (Figure 7), which are easy identifiable. The engyodontic stage is defined by the needle-like teeth, an unformed nasal capsule, an undifferentiated medial finfold and hypurals. Leptocephalis larger than 20 mm TL are

Figure 7. Engyodontic and euryodontic stage of a congrid leptocephalus (CASTLE 1984)

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3 RESULTS

in the euryodontic stage. This stage is characterized by the embedding of the engyodontic teeth and the replacement by shorter, broad-based teeth. Furthermore, the lower jaw shortens relative to the upper jaw, the head decreases in relative length and the fins and hypurals get differentiated. The family identification follows the Morphology and pigment patterns. A short identification guide of the leptocephalis of the present study is given in Figure 8, where the diagnostic characters are clearly visible in leptocephali.

-- Reference: CASTLE (1984);

Ophichthidae Congridae Nettastomatidae Serrivomeridae Muraenidae Nemichthyidae Eurypharyngidae

Figure 8. Identification guide of the anguilliform and saccopharyngiform families of the present study. + = all or most species; (+) = some species. Modified after CASTLE (1984).

¬ SACCOPHARYNGIFORMES The caudal fin is absent or reduced. The gill openings are ventrally situated. The dorsal and anal fins are long based. The body of the leptocephali is deep. The myomeres are V-shaped. An identification guide for the family of Eurypharyngidae is given in Figure 8. Within the area of investigation the family Eurypharyngidae is known to be monogeneric and monospecific, therefore the specimens are most likely Eurypharynx pelecanoides VAILLANT, 1882.

-- Reference: NELSON (2006);

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SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

¬ CLUPEIFORMES Depending on their yolk sac size, clupeoid larvae hatch at 2 – 5 mm notochord length. Larvae of demersal adhesive eggs hatch at a more developed stage than those of pelagic eggs, but have pectoral fin buds and a continuous finfold present at hatching in common. The sequence of the appearance and ossification of fin rays is headed by the caudal, followed by the dorsal, anal, and pelvic, and ends with the pectoral fin. The fin formation is not fully completed until transformation, which occurs at approximately 20 mm standard length (SL). For identification, counting the number of myomeres or vertebrae is the best tool, however, pigment patterns can be useful, as well. Clupeids can be differentiated from engraulids by the larger gut length and the position of the dorsal to the anal fin. Furthermore, the number of myomeres between those two fins is important for identifying to lower levels within the families. But during metamorphosis the vent, the dorsal and the anal fin migrate forwards relative to the number of myomeres.

-- Reference: OLIVAR and FORTUÑO (1991);

Clupeoidei > Engraulidae In most engraulids the posterior end of the dorsal fin is located over the anterior beginning of the anal fin, which is the most important character leading to this family and easily visible (Figure 27). Engraulids also have a shorter gut than clupeids.

-- References: MCGOWAN and BERRY (1984);

• Engraulis encrasicolus Meristics: Dorsal rays:

14 – 17

Anal rays:

14 – 22

-- Morphology The larvae are elongated with a body depth of only 8 – 9 % of the SL at the level of the pectoral fins. The gut extends to a length of 78 % of SL in early larvae and decreases to 70 % in larvae more than 20 mm SL, but generally terminates between the middle and the end of the dorsal fin. The swim bladder is clearly visible and produces an indentation of the gut. The position of the terminate gut and the fin ray count were confirmed to be of high diagnostic value in this study.

-- Pigmentation In very early developmental stages pigmentation is apparent ventrally between the pectoral and caudal region. A more important marker for identifying is the number of 5 – 8 melanophores on the foregut and 3 – 4 on the swim bladder, which fade away during growth. In larvae of more than 10 mm in length one large spot is found at the anus and up to four more are embedded in the body behind it (Figure 26). Fin pigmentation is only found on the anal fin with a number of 8 spots and a band located posteriorly, but was not visible in the fish larvae examind in this study.

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3 RESULTS

-- References: MCGOWAN and BERRY (1984); OLIVAR and FORTUÑO (1991);

-- Reference material: 2925 specimens; 11/97: S4 (P18858-999; P181000-3692); S6 (P183693-3704); S7 (P183705-3719); S10 (P18818-848); S13 (P18849-857); 04/99: S14 (P183720); 05/99: S1 (P183721-3725); 03/02: S4 (P18814-815); S5 (P18816); S7 (P18817); S10 (P18801-805); S11 (P18806-810); S14 (P18811-813);

-- Distribution: During winter, larvae were mainly found at the epipelagic stations southeast of Fuerteventura at a depth range from 22 – 219 m during the day as well as during the night. Some specimens were caught southeast of Fuerteventura at a neritic station at a depth range of 15 – 43 m during the night as well as at a epipelagic station in 31 – 140 m depth during daytime. The highest numbers of E. encrasicolus larvae of the entire period of investigation were caught in autumn. During this period, the largest amounts of larvae were caught at neritic stations southeast of Fuerteventura at a depth of 22 – 35 m during the daytime, southeast of Lanzarote at 13 – 41 m depth during daytime and northeast of Lanzarote at a depth of 20 – 64 m during the night. Smaller numbers of larvae were found at epipelagic stations west of the channel between Lanzarote and Fuerteventura at 28 – 43 m and southeast of Fuerteventura at 90 – 107 m depth each around dawn. In spring, fewer specimens were caught. One larva was found at a mesopelagic station southwest of Gran Canaria at a depth of 480 – 694 m during the day. This single larva may derive from an earlier, shallower tow, most probably it remained in the net from the previous neritic haul and contaminated the mesopelagic trawl. Some more larvae occurred at a station far east off Fuerteventura at a depth of 38 – 40 m at dawn.

Clupeidae The body is elongate and has an approximate number of 50 myomeres.The relation of gut to body length is longer in clupeids than in engraulids and there is a gap between the posterior margin of the dorsal fin and the anterior margin of the anal fin.

• Sardinella sp. The number of vertebrae is less than 50 in the east Atlantic species in contrast to the genus of Sardina with more than 50 vertebrae. The larvae of Sardinella (exception S. tawilis (HERRE, 1927)) have 16 – 20 dorsal and 14 – 23 anal fin rays. The number of melanophores on the hindgut is smaller in the genus Sardinella than in Sardina. Using the fin ray count and the number of vertebrae/myomeres made it easy to distinguish Sardinella and Sardina. Both were found to be easily countable in well preserved larvae.

-- References: BERTOLINI et al. (1956); MCGOWAN and BERRY (1984); OLIVAR and FORTUÑO (1991);

-- Reference material: 2 specimens; 11/97: S7 (P183726); 03/02: S4 (P183727);

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SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Distribution: The larvae were found during the night at a neritic station southeast of Lanzarote at 3 – 41 m depth in autumn and at an epipelagic station southeast of Fuerteventura in a depth of 22 – 130 m during winter.

¬ ARGENTINIFORMES Families of this order are grouped into two suborders of deep-sea fishes. Characteristic for the suborder of Argentinoidei is the voluminous medial finfold in which the dorsal and anal fins form unattached to the body margin. First the pectoral fins form which remain very small in argentinids and in some microstomatids (Bathylaginae), suborder Alepocephaloidei, and subsequently the caudal fin develop. In the suborder of Alepocephaloidei the adipose fin is usually lacking. The preanal length is shorter (50 – 70 % SL) than in the Argentinoidei (70 – 95 % SL). The gut is straight and extends about to the midpoint of the body length and in all Argentinoidei the terminal section of the gut is deflected. In most Argentinoidei the dorsal and anal fin formation takes place during flexion. The anal fin develops just behind the deflection point of the gut and is located at the almost same spot as in the adults. The pelvic fin forms last at about mid-body, at the dorsal fin level in Argentinidae and Microstomatidae. During transformation the body deepens, the snout elongates and the eye increases in size. But the transformation to demersal juveniles may be protracted in some species by remaining in their pelagic habitat and keeping the larval pigmentation.

-- Reference: AHLSTROM et al. (1984 a); OLIVAR and FORTUÑO (1991); RICHARDS (2006a);

Argentinoidei > Argentinidae Characteristic for Argentinids is the series of ventral blotches from the pectoral fins to the caudal region. The number of these blotches is constant for each species, as well as the sequence of their appearance. In some species these blotches form lateral bars. The head is relatively small with a rounded profile. The eye is rounded to slightly ovoid. Transformation is gradual with an increase in body depth, head length, eye size and pigmentation ending in the pelagic juvenile stage. The prolonged transformation terminates when scales and a silvery integument are formed, and can be completed up to a size of 100 mm.

-- Reference: AHLSTROM et al. (1984 a);

• Argentina sphyraena Meristics:

Development:

Vertebrae:

46 – 55

Dorsal rays:

10 – 12

Anal rays:

11 – 15

Pelvic rays:

10 – 12

Pectoral rays:

15 – 18

Inf. Téc. Inst. Canario Cienc. Mar. n°13

Hatching: 7 mm Flexion: ~ 17 mm

3 RESULTS

-- Morphology: The anus is situated at about 2/3 down the total body length. The caudal fin is rounded at first, but becomes biloped during development. The dorsal and anal fin appear disattached from the body margin within the finfold. First outlines of dorsal and anal fins become visible with 14 – 15 mm and flexion starts at 17 mm. At a length of 27.5 mm the full number of fin rays is developed. The adipose fin forms from the finfold and the pelvic fins forms last at about 27.5 mm.

-- Pigmentation: There is no head pigmentation present in this family. A. sphyraena has only 7 ventral pigment patches, of which 5 are preanal and 2 are postanal. These patches do not extend to the dorsal end, but some stellate chromatophores are present on the gut. This pigmentation pattern is quite unique, but to do not confuse the larvae with other argentinids counting the pectoral fin rays verifies the identification to this species.

-- Reference: BERTOLINI et al. (1956); COHEN (1984); SCHMIDT (1918);

-- Reference material: 2 specimens; 11/97: 04/99: S19 (P191592); 03/02: S6 (P191593);

-- Distribution: A. sphyraena is a new record for the area of the Canary Islands. The previous closest Distribution was along the West African coast. The larvae were found at a depth range of 980 – 1162 m southeast of Fuerteventura and at 553 – 716 m depth west of Tenerife each during daytime in winter and spring, respectively.

• Glossanodon leioglossus Meristics:

Development:

Vertebrae (Myomeres):

49 – 51 (51)

Dorsal rays:

12 – 14

Anal rays:

10 – 13

Pelvic rays:

11 – 12

Pectoral rays:

19 – 20

Flexion: 15 mm

-- Morphology: The anus is located at 3/4 of total body length. At a length of 14 mm the disattached development of dorsal and anal fin within the finfold starts, connecting to the body as larval development progresses. The dorsal fin is located more posterior than in Argentina sphyraena. The high pectoral fin ray number is the best distinguishing character from A. sphyraena and was the only possibility to identify the larvae, since the pigmentation pattern was not that clear. At 30 mm length the full fin ray numbers are developed in the larvae, followed by the formation of the adipose fin at about 33 – 35 mm.

-- Pigmentation: Characteristic is the pigment stripe from the snout to the operculum, interrupted by the eye. The total number of ventral stripes is 9, where 7 are positioned preanally and 2 postanally.The stripes increase towards the dorsal margin with development, but become restricted ventrally during later development.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Reference: BERTOLINI et al. (1956); SCHMIDT (1918);

-- Reference material: 2925 specimens; 03/02: S3 (P191596-1598); S6 (P191602); S16 (P191599-1601); S18 (P191594-1595);

-- Distribution: All specimens were found during winter on the cruise sampling southeast of Fuerteventura and were mainly caught at mesopelagic depths within a range of 470 – 915 m during the daytime and within 440 – 665 m in the afternoon. Some larvae were also caught at an epipelagic station at a depth of 185 – 211 m during the day.

Microstomatidae This family consists of two subfamilies, the Microstomatinae and the Bathylaginae, in which big eyes are characteristic. In Bathylaginae the head size is moderate while the shape is variable. The snout is more elongated than in argentinids. The eye shape varies from almost round to elliptical and may be stalked. The pigmentation patterns are variable and can either have large isolated melanophores laterally on the last section of the gut and a single large blotch in the middle of the pectoral fin and the anus, or they can have a series of small dots along their body, as seen in bathylagids. The head is pigmented, especially on both jaws and on the opercle. The flexion is delayed and may be not completed by the end of the larval stage. Bathylaginae have a direct transformation of morphological changes such as an increase in body depth, eye size, head size and pigmentation. Also, the gut becomes coiled and covered by a black peritoneal sheath.

-- Reference: AHLSTROM et al. (1984 a); FAHAY (1983); NELSON (2006);

• Bathylagus sp. The body is elongated. Most species of this genus generally have stalked eyes, but the proportions vary among species. The pelvic fin is positioned below the dorsal fin. The adipose fin is present, but can be reduced to be nearly indiscernible in some species.

-- Reference: FAHAY (1983);

-- Reference material: 1 specimen; 05/99: S6 (P191604);

-- Distribution: The larva was caught at an epipelagic station in a depth range of 48 – 66 m far north off La Gomera and Tenerife at night during late spring.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

• Dolicholagus longirostris Meristics: Vertebrae:

48 – 51

Dorsal rays:

10 – 12

Anal rays:

19 – 21

Pelvic rays:

9 – 10

Pectoral rays:

9 – 12

-- Morphology: The larvae have a slender body with a slightly elongated snout. The eyes are stalked and the length of the stalks in relation to the head length change during development.The mean values of the percentage of stalk to head length are 54% in preflexion larvae, 48% during flexion and 27 % in postflexion larvae (Figure 28).

-- Pigmentation: D. longirostris develops a pattern of heavy pigmentation. At the posterior section of the gut a lateral series of small melanophores and a series of rectangular melanophores along the hypaxial myomeres form in preflexion larvae. In late larvae, the lateral gut melanophores span the entire gut. The anterior series has a wider spacing than the one found on the posterior gut section. Also, along the epaxial myomeres rectangular melanophores develop from the posterior to the anterior part.The head is fully pigmented from the opercle to the jaws. Although the single specimen examined was not in the best condition, the unique pigmentation pattern made the species identification possible.

-- Reference: AHLSTROM et al. (1984 a); FAHAY (1983);

-- Reference material: 1 specimen; 05/99: S6 (P191603);

-- Distribution: The only specimen was found at an epipelagic station at a depth of 48 – 66 m far north off La Gomera and Tenerife at night during late spring season.

ALEPOCEPHALOIDEI > Platytroctidae The body shape is slender and some members of this family develop photophores during the yolk sac stage. In some species a white tissue develops at the eyes and a spine is present at the cleithral symphysis. The distinctive character for this family is the external very black coloured tube and sac above the cleithrum, which contains a luminous fluid. The preanal length exceeds the midpoint of the body with an interspace between anus and anal fin origin.

-- Reference: RICHARDS and HARTEL (2006);

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

• Sagamichthys schnakenbecki Meristics: Vertebrae:

48 – 51

Dorsal rays:

16 – 18

Anal rays:

15 – 17

Pelvic rays:

Pectoral rays:

15 – 19

-- Morphology: Photophores are well developed and a white tissue surrounds the eyes, which does not occur on the infraorbitals. The anal fin originates well behind the dorsal origin.There is no spine at the cleithral symphysis. I identified the specimen based on the presence of the external tube (Figure 29) and the fin ray count.

-- Reference: RICHARDS and HARTEL (2006);

-- Reference material: 1 specimen; 04/99: S14 (P 17875);

-- Distribution: The only specimen of S. schnakenbecki was caught at a mesopelagic station south of Gran Canaria in a depth range from 480 – 694 m during the day in spring.

¬ STOMIIFORMES The Morphology of the larvae is very diverse. The body shape varies from slender, elongate with a long, trailing gut and long jaws and small elliptical eyes to a less slender shape with the gut reaching farther anterior or even close to the midpoint of the body and round or almost round eyes. Already during the larval period most larvae develop photophores.

-- Reference: OLIVAR and FORTUÑO (1991);

GONOSTOMATOIDEI > Gonostomatidae The body is slender to elongate. The base of the anal fin is almost as long as the postanal length. Usually fins develop at the same position as in adult fish (exception for Pollichthys). The adipose fin can be absent or present. Pectoral fins are often situated on peduncles. For identification the dorsal and anal fin, ray counts are an important character.There are two ways of photophore development. Either they have a simultaneous formation without associated pigmentation (Cyclothone, Diplophos and Manducus), or a successive appearance with associated pigmentation formation.

-- Reference: AHLSTROM et al. (1984 b); FAHAY (1983); RICHARDS (2006b);

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

• Cyclothone sp. The body is elongated with the gut extending to the midpoint of the body or just behind it. A conspicuous swim bladder is located posteriorly on the gut. The dorsal and anal fins are situated at opposite positions far back on the body. There is a gap between the pelvic and the anal fins. The pigmentation on the ventral side from the anus to the tail is typical for this genus. The overall pigmentation pattern is the best distinguishing character. Most or all ventral photophores form at once during the late post-larval stage. Photophores are lacking on the isthmus and have a low number in the ventral series. Meristic counts are in the range of 12 – 15 rays in the dorsal fin, 29 – 31 rays in the anal fin and 29 – 33 vertebrae. Beside the typical body shape and the fin positions, the fin ray counts were used to diagnose Cyclothone sp. Species identification was not possible due to the bad condition and the destroyed pigmentation pattern of the larva.

-- Reference: AHLSTROM et al. (1984 b); FAHAY (1983); OLIVAR and FORTUÑO (1991);

-- Reference material: 1 specimen; 05/99: S2 (P2014861);

-- Distribution: One specimen was caught at a mesopelagic station east of Fuerteventura in a depth range from 495 – 561 m during the day in late spring.

• Cyclothone acclinidens Meristics: Vertebrae:

30 – 32

Dorsal rays:

13 – 14

Anal rays:

18 – 20

Pelvic rays:

5–6

Pectoral rays:

8 – 10

-- Pigmentation: Melanophores are located on the fore- and midbrain and on the symphysis of the lower jaw. 12 external and 10 internal spots develop at the base of the anal fin. About 11 melanophores are situated ventrolateral at the anterior section of the body. Additional melanophores form along the dorsal side of the body. One spot forms at the dorsal side of the tail. In species identification, the fin ray count is not useful since the ray counts are overlapping between species. Here the pattern of pigmentation is diagnostically important and was clearly visible in the larva studied; the external and internal spots were all clearly visible.

-- Reference: BADCOCK (1986a); OLIVAR and FORTUÑO (1991);

-- Reference material: 1 specimen; 11/97: S4 (P2014859);

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Distribution: A single specimen was caught at a neritic station southeast of Fuerteventura at a depth of 22 – 35 m during the daytime in autumn.

• Cyclothone alba Meristics: Vertebrae:

30 – 31

Dorsal rays:

12 – 14

Anal rays:

17 – 20

Pelvic rays:

Pectoral rays:

9 – 10

-- Pigmentation: The very small larvae develop 5 and older larvae 7 – 8 melanophores on the ventral side of the body. Later, a series of melanophores form on the ventrolateral side of the anterior portion of the body. A row of melanophores is located along the dorsal margin of the body, with four spots at the caudal region and another five spots beneath the origin of the dorsal fin. There is no pigmentation on the base of the caudal fin. This pigmentation pattern was clearly visible in the studied specimen and, again, the fin ray count was not useful, since the count is overlapping with other Cyclothone species.

-- Reference: BADCOCK (1986a); OLIVAR and FORTUÑO (1991);

-- Reference material: 1 specimen; 11/97: S10 (P2014859);

-- Distribution: One specimen was found at a neritic station northeast of Lanzarote at 20 – 64 m depth during the night in autumn.

• Gonostoma denudatum Meristics:

Development:

Vertebrae:

Dorsal rays:

14 – 15

Anal rays:

28 – 30

Pelvic rays:

Pectoral rays:

11 – 12

Transformation: ~ 34 mm

-- Morphology: The number of myomeres is 38. The body is slender with a pointed snout. The eyes are slightly ovoid with the longer axis orientated vertically. The swim bladder is conspicuous. The origins of the dorsal and anal fin are situated opposite of each other, with the anal fin displaying a longer base. The pelvic fin bulbs are located under the posterior end of the swim bladder. At 14 – 15 mm the first photophores appear and temporal rudiments of pelvic fins develop. With further development an adipose fin appears with pigmentation at its base. The studied specimens showed the fin positions typical for the family of

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

Gonostomatidae and the fin ray count was indicative for the genus. The species identification was easiest done by using the pigmentation pattern at the caudal peduncle described below. This pattern is unique and was clearly visible in my specimens (Figure 30).

-- Pigmentation: Intestinal blotches are situated from above the gut to the pigmented swim bladder. One spot is located at the end of the gut. A series of melanophores extends from the posterior end of anal fin to the caudal fin. Scattered pigments develop on the base of the caudal fin. A prominent stripe of pigments is formed posteriorly diagonally over the caudal fin base from the dorsal margin of the caudal peduncle to base of the lower caudal fin rays. Further pigments are found on the head, above the eye and two spots on the hindbrain. Pigmentation is also present at the tip of the lower jaw.

-- Reference: AHLSTROM et al. (1984 b); BADCOCK (1986a); BERTOLINI et al. (1956);

-- Reference material: 172 specimens; 11/97: S4 (P2015008); S6 (P2015009-10); S13 (P2014862-2015005); 04/99: S13 (P2015011); S16 (P2015012); 05/99: S6 (P2015013-2015033); 01/00: S1 (P2015006-07);

-- Distribution: G. denudatum was caught in spring, autumn and winter. In spring, some specimens were found at neritic stations southwest of Gran Canaria in a depth range from 18 – 61 m during the night and in late spring at epipelagic stations of La Gomera and Tenerife in a depth of 48 – 66 m at night. The highest number of larvae during the entire investigation period was caught in autumn at an epipelagic station southeast of Fuerteventura at 90 – 107 m depth in the morning. Further larvae were found epipelagic west of the channel between Fuerteventura and Lanzarote in a depth of 28 – 43 m in morning and only one specimen was caught at the neritic station southeast of Fuerteventura at 22 – 35 m depth during the daytime. During winter, only two specimens were found at a mesopelagic depth of 765 – 845 m in the late daytime.

• Sternoptychidae Most larvae forms are still not known. Distinctive characters for this family are the presence of a pseudobranch and photophores, which coalesce in clusters or groups and develop prolongated and sequentially.The succession of development starts in most larvae either with the OP3 or Br photophore. The body can be very deep and laterally strongly compressed with a long or elongated and only slightly compressed tail. The pigmentation is very rare in all species and patterns are very useful tools in species identification together with meristic characters and morphometrics. During transformation the larvae descend to their adult habitats.

-- Reference: AHLSTROM et al. (1984 b); BADCOCK (1986b); RICHARDS (2006c);

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

• Maurolicus muelleri Meristics:

Development:

Vertebrae:

33 – 35

Dorsal rays:

10 – 11

Anal rays:

22 – 28

Pelvic rays:

Pectoral rays:

17 – 19

Flexion: 5.5 – 6.5 mm

-- Morphology: The body is very slender with a body depth of 11 % of NL in very early larvae which deepens to about 25 % of SL in specimens of about 20 mm. The preanal length in early stages extends slightly posteriorly of the midpoint of the SL and increases up to 2/3 of SL with development. The swim bladder is conspicuous and situated at the middle of the gut length. Eyes are ovoid, where the longer axis is orientated vertically, but the eyes become round with further development. The adipose fin is present.The origin of the anal fin is positioned behind the origin of the dorsal fin.The sequence of fin formation is from caudal to anal, dorsal, pelvic and pectoral fins.

-- Photophores: Photophores develop gradually and clumped. The formation of the photophores varies between individuals of the Atlantic, Mediterranean and Pacific. But the clumped photophores together with the slender body shape are a good indicator for the subfamily of the Maurolinae. In larvae at the very late postflexion stage the adult photophore patterns and meristic counts are very useful tools in species identification.

-- Pigmentation: FAHAY (1983) described 7 – 10 spots which develop along the anal fin base in larvae with a size of 6 – 10 mm. These spots disappear with further development. OLIVAR and FORTUÑO (1991) noted a lack of pigmentation for larvae up to a size of 7 mm, after which some melanophores develop above the swim bladder. In later larvae melanophores form dorsally and ventrally on the caudal peduncle according to both guides. In larvae larger than 10 mm pigmentation spreads from the head to the peduncle over the dorsum and at 15 mm SL the pigmentation covers the entire dorsal region. In the late postflexion larvae studied the pigmentation on the caudal peduncle consisting of a band of scattered melanophores reached from the ventral to the dorsal margin of the caudal peduncle at the level of the posterior group of clumped photophores (Figure 31).

-- Reference: BADCOCK (1986b); FAHAY (1983); OLIVAR and FORTUÑO (1991);

-- Reference material: 3 specimens; 05/99: S2 (P2015034-36);

-- Distribution: The specimens were caught at an epipelagic station far north of Gran Canaria in a depth range of 42 – 47 m in the night during late spring.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

PHOSICHTHYOIDEI > Phosichthyidae Larvae are slender with a long preanal length. The eyes are usually ovoid but become round in latest larval stages. Trunk pigmentation varies.The main feature of this family is the development of photophores without any associated pigmentation, but later the pigments develop simultaneously. For the identification of larvae meristic counts, morphometrics and pigmentation patterns are of essential importance.

-- Reference: RICHARDS (2006d);

• Polymetme corythaeola Meristics: Vertebrae:

43 – 45

Dorsal rays:

12 – 14

Anal rays:

30 – 34

Pelvic rays:

Pectoral rays:

9 – 10

-- Morphology: The body is slender, with a moderately sized head and a large mouth. The anal fin origin is directly below the end of the dorsal fin and reaches to the caudal peduncle. An adipose fin is present at the midlevel of the posterior half of the anal fin. The specimen was identified based on adult characters.

-- Reference: AHLSTROM et al. (1984 b); BADCOCK (1986c);

-- Reference material: 1 specimen; 11/97: S13 (P2015037);

-- Distribution: A transforming specimen was caught at an epipelagic station southeast of Fuerteventura at 90 – 107 m in early morning during autumn.

• Vinciguerria sp. The body is slender and deepens during transformation.The straight gut extends to about 75 % of SL, but decreases in length during transformation.The snout is pointed and the head length is about 20 % of SL and increases during transformation.The eyes are ovoid and slightly stalked.The sparse pigmentation pattern is the best character for species identification. Prominent hereby is the spot on the caudal peduncle. During transformation the photophores develop simultaneously. The sequence of fin formation starts with the caudal, continuous with the dorsal and anal fin, the pelvic buds and the pectoral rays form at transformation. The adipose fin is present, but inconspicuous and becomes more obvious with development. This specimen was in a bad condition and could not be identified to the species level.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Reference: Fahay (1983);

-- Reference material: 1 specimen; 01/00: S3 (P2015096);

-- Distribution: In winter the specimen was caught at a mesopelagic station north of Gran Canaria in a depth of 356 – 402 m during the day.

• Vinciguerria poweriae Meristics:

Development:

Vertebrae:

38 – 39

Dorsal rays:

13 – 15

Anal rays:

12 – 14

Pelvic rays:

Pectoral rays:

9 – 11

Flexion: 4 – 6 mm SL Transformation: 13 – 14 mm, gradually

-- Morphology: The body is slender and the eyes are round. The origin of the anal fin base is positioned posterior of the dorsal fin. Further description of Morphology can be found at the genus description of Vinciguerria.

-- Pigmentation: The prominent caudal spot is positioned at the middle of the peduncle and is still visible at transformation.There is no further pigmentation on swim bladder, anal or caudal fin base or on isthmus. The caudal spot and the absence of other pigmentation were the best character combination leading to this species.

-- Reference: BADCOCK (1986c); FAHAY (1983);

-- Reference material: 3 specimens; 11/97: S13 (P2015093); 05/99: S3 (P2015094); S6 (P2015095);

-- Distribution: In autumn only one transforming specimen was caught at the epipelagic station at 90 – 107 m depth in the morning. In late spring two specimens were caught at epipelagic stations at night, one far north off Gran Canaria in a depth of 42 – 47 m and the other one off of La Gomera and Tenerife in a depth range of 48 – 66 m.

• Vinciguerria nimbaria Meristics:

Development:

Vertebrae:

40 – 42

Dorsal rays:

14 – 15

Anal rays:

13 – 16

Pelvic rays:

Pectoral rays:

9 – 11

Inf. Téc. Inst. Canario Cienc. Mar. n°13

Flexion: 4 – 6 mm SL Transformation: 13 - 16 mm, gradually

3 RESULTS

-- Morphology: The origin of the anal fin is positioned at the midpoint of the dorsal fin.The eyes are round. Further description of Morphology can be found at the genus description of Vinciguerria.

-- Pigmentation: The caudal spot is located ventrally on the caudal peduncle and is lost during transformation. There is no pigmentation of the gas bladder, but above the anal fin two to three spots develop. Further two or three spots are placed in a vertical line at the caudal fin base and a narrow line is present along the isthmus. In the studied specimens all pigmentation patterns were clearly visible.

-- Reference: BADCOCK (1986c); FAHAY (1983); JESPERSEN and TÅNING (1926); OLIVAR and FORTUÑO (1991);

-- Reference material: 23 specimens; 11/97: S4 (P2015084-85); S6 (P2015076); S9 (P2015077); S10 (P2015079-2015083); S13 (P20150702015075); S18 (P2015091-92); 04/99: S12 (P2015078); 05/99: S1 (P2015086-88); S3 (P2015090); S4 (P2015089);

-- Distribution: V. nimbaria occurred at the neritic stations, southeast of Fuerteventura at a depth of 22 – 35 m during the day, and northeast of Lanzarote at a depth of 20 – 64 m during the night and also east of Gran Canaria at 116 – 146 m during the night. Larvae were also found at epipelagic stations west of the channel of Lanzarote and Fuerteventura at depths of 28 – 43 m in the morning and southeast of Fuerteventura at 90 – 107 m also during the morning, where four specimens were in the transforming stage. One specimen was caught at the mesopelagic station northeast of Lanzarote at a depth of 527 – 557 m in the evening. In spring one specimen was caught at the epipelagic station southwest of Gran Canaria at 38 – 49 m in night. Further larvae occurred during late spring mainly at epipelagic stations far east of Fuerteventura at depths of 38 and 40 m in morning and far north of Gran Canaria at a depth of 42 – 47 m during the night. One further specimen was caught at a mesopelagic station far north of Gran Canaria at 517 – 591 m during the day.

• Vinciguerria attenuata Meristics:

Development:

Vertebrae:

40 – 41

Dorsal rays:

13 – 15

Anal rays:

13 – 16

Pelvic rays:

6–7

Pectoral rays:

9 – 10

Flexion: 4 – 6 mm SL Transformation: 13 - 17 mm, gradually

-- Morphology: The origin of the anal fin is at the middle of the dorsal fin. The eyes are tubular. Further description of Morphology can be found at the genus description of Vinciguerria.

-- Pigmentation: The caudal spot is positioned median on the caudal peduncle, but is sometimes lost during transformation. The difference to V. poweriae is the pigmented swim bladder, but no additional pigments exist.The pigmentation on the swim bladder was always clearly visible in the studied larvae and was the easiest character for distinguishing V. attenuata from the other Vinciguerria species.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Reference: BADCOCK (1986c); FAHAY (1983); JESPERSEN and Tåning (1926); OLIVAR and FORTUÑO (1991);

-- Reference material: 32 specimens; 04/99: S12 (P2015042-46); S14 (P2015047); S16 (P2015038-41); 05/99: S1 (P2015048-49); S2 (P2015051); S3 (P2015052-2015065); S4 (P2015050); S6 (P2015066-2015069);

-- Distribution: During spring larvae were present southwest of Gran Canaria at an epipelagic station at 38 – 49 m and at a neritic station at 18 – 61 m both during night. One specimen was also caught at a mesopelagic station south of Gran Canaria at 480 – 694 m during the day. In late spring larvae were mainly found at epipelagic stations during the night. Most were found far north off Gran Canaria at a depth of 42 – 47 m and far east off Fuerteventura at 38 – 40 m and some more occurred at an epipelagic station north of La Gomera and Tenerife at a depth of 48 – 66 m. Two additional specimens were caught, one at a mesopelagic station during the day east of Fuerteventura at 495 – 561 m and one far north of Gran Canaria at a depth of 517 – 591 m.

STOMIOIDEA > Stomiidae > Chauliodontinae Yolk sac larvae have several melanophores on the caudal peduncle, but these are lost soon. No other pigments develop during larval stage. The body is slender with round cross section and do not deepen with development. The head is small with ovoid eyes and a short snout. The gut is thinner and longer than in larvae of Stomias and extends beyond the anal fin origin. The finfold is small but develops further in the posterior part of the body. Dorsal and anal fins form during the late postflexion stage. Some larvae shrink during transformation.

-- Reference: GIBBS (1986a); KAWAGUCHI and MOSER (1984);

• Chauliodus sp. The body is elongate.The dorsal fin bears only 5 – 7 rays and is positioned very anteriorly of the body with an elongated first ray. The anal fin has 10 – 13 rays and develops, analogous to the adipose fin, very posterior on the body close to the caudal peduncle. Two ventral series of photophores develop on each side of the ventral midline. The number of vertebrae ranges from 51 – 62. Some of the specimens studied had already developed photophores and an elongated first ray of the dorsal fin. Other larvae had no pigmentation at all and had no developed dorsal or anal fin (Figure 34).

-- Reference: GIBBS (1986a); KAWAGUCHI and MOSER (1984);

-- Reference material: 4 specimens; 11/97: S4 (P2015103); 05/99: S2 (P2015100); S3 (P2015101); S6 (P2015102);

-- Distribution: Larvae were caught at a neritic station southeast of Fuerteventura at a depth of 22 – 35 m during the day in autumn. In late spring three larvae occurred at a mesopelagic station east of Fuerteventura, one at 495 – 561 m depth during the day, one at an epipelagic station far north off Gran Canaria at a depth of 42 – 47 m during the night and one an epipelagic station north of La Gomera and Tenerife in a depth range from 48 – 66 m during the night.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

Stomiinae The slender larvae hatch with 3 – 4 mm NL and possess an elongate yolk sac. The body deepens during the late postflexion stage.The head is small with a flat snout and ovoid eyes.The very long gut is nearly as long as the larvae itself with a deflected terminal gut section. Larvae have a small finfold which is further developed at the posterior part of the body. The opposing anal and dorsal fin are situated far back on the body. The pelvic buds do not develop until or just prior to transformation.

-- Reference: KAWAGUCHI and MOSER (1984);

• Stomias sp. A series of melanophores forms between the body and gut, which extends to the tip of the notochord in early larvae and is already lost before flexion. Most species form a new series along the ventral midline of the gut from the isthmus to the vent. The genus Stomias has 64 – 83 vertebrae and is therefore easy to distinguish from the genus Macrostomias, which has a higher number of vertebrae of 164. This myomere/vertebrae count and the fin position were the most important and easy visible characters used in the diagnosis.

-- Reference: KAWAGUCHI and MOSER (1984);

-- Reference material: 2 specimens; 11/97: S13 (P2015108); 05/99: S3 (P2015107);

-- Distribution: Larvae were found at an epipelagic station during the night, far north of Gran Canaria at a depth of 42 – 47 m in late spring and southeast of Fuerteventura at a depth of 90 – 107 m in autumn.

• Stomias boa Meristics:

Development:

Vertebrae:

64 – 83

Dorsal rays:

16 – 22

Anal rays:

18 - 25

Pelvic rays:

4–5

Pectoral rays:

6–9

Ventral Photophores:

82 – 91

Start of transformation: 30 mm

-- Morphology: The gut length is close to 90 % of SL and has a deflecting terminal gut section, which was clearly visible the studied specimen (Figure 36). The larvae have an elongated head with very small ovoid eyes. Fin ray counts listed above are valid for all species of the genus Stomias.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Pigmentation: The ventral spots disappear at about 20 mm. The larvae have a series of midlaterally melanophores along the body and a pigmentation on head as well as on dorsal, anal and caudal fins, whereas in the studied specimen the dots along the anal fin base were the most distinctive ones (Figure 36).

-- Reference: FAHAY (1983); KAWAGUCHI and MOSER (1984); OLIVAR and FORTUÑO (1991);

-- Reference material: 1 specimen; 11/97: S9 (P2015106);

-- Distribution: The specimen was caught at a mesopelagic station northeast of Lanzarote in a depth range from 527 – 557 m in early night time during autumn.

Astronesthinae Larvae of this family show a great variability in pigmentation and Morphology. The body form can be from slender and round in cross section to laterally compressed. The gut can be from being trailed to having a deflected terminal gut section. Distinctive characters are the well developed medial finfold and the advanced position of the dorsal fin relative to the anal fin. All three specimens probably represent three different species. At least two similar specimens (Figure 32 - Figure 33) had no pigmentation at all, but seem to be two different species, since the gut of one specimen is deflected directly anteriorly of the anal fin origin, while in the other specimen the gut is deflected far in advance.The latter specimen might belong according to KAWAGUCHI and MOSER (1984) to the genus Astronesthes.

-- Reference: KAWAGUCHI and MOSER (1984);

-- Reference material: 3 specimens; 11/97: S10 (P2015099); S13 (P2015097); 03/02: S11 (P2015098);

-- Distribution: In autumn one larva of the Type II was caught at an epipelagic station southeast of Fuerteventura at a depth of 90 – 107 m in morning and another one was found at a neritic station northeast of Lanzarote at a depth of 20 – 64 m at night. In late winter the third specimen occurred at an epipelagic station in a depth range from 31 – 140 m in daytime.

Melanostomiinae In general the Morphology and pigmentation are very variable among this family. But with the exception of the genus Eustomias the larvae are elongated having an ovoid cross section of the body and a laterally compressed large head with small ovoid eyes. The gut is deflected and does not extend the finfold, but may extend beyond the anal fin origin. The anal and the dorsal fin form at the posterior part of the body opposite in position. One or more melanophores form at each epaxial myomere and each hypaxial myosepta.

-- Reference: KAWAGUCHI and MOSER (1984); RICHARDS (2006e);

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

-- Reference material: 1 specimen; 11/97: S13 (P2015105);

-- Distribution: The specimen occurred at the epipelagic station southeast of Fuerteventura in a depth range from 90 – 107 m during morning in autumn.

Idiacanthinae The family Idiancanthidae consists of the only genus Idiacanthus. Larvae of this genus are extremely elongated and flattened with a very small finfold. Very prominent are the elliptical eyes on long stalks with their cartilaginous supporting rods. The deflected and trailing gut extends beyond the origin of the anal fin.The dorsal fin already starts to form during the preflexion stage anteriorly to the anal fin and is longer than the latter. During transformation rays add sequentially anteriorly to the dorsal an anal fins, so that in adult fish the dorsal fin extend to about 2/3 and the anal fin to about 1/2 of the total body length. During larval stages the pectoral fin is well developed, but gets lost during transformation. If pelvic fins form, than in transforming larvae, whereas the pelvic fins develop always in females, but not in all males. The pigmentation pattern consists of melanophores on each hypaxial myomere, which are spreading into the myosepta with development. Further melanophores are found internally on the isthmus and along the midline of the gut.

-- Reference: KAWAGUCHI and MOSER (1984); RICHARDS (2006f)

• Idiacanthus fasciola Meristics:

Development:

Vertebrae (Myomeres):

~ 78 (~ 78)

Dorsal rays:

54 –80

Anal rays:

29 – 55

Pelvic rays:

6 (females)

Transformation of male: 35 mm

Pectoral rays:

Transformation of female: 45 mm

-- Morphology: The body is very elongated with a long head and flat snout. Eyes are stalked, but shorten gradually with further development. The gut is straight and deflected, but gets incorporated during transformation. Dorsal finfold is stronger developed than the ventral is. During transformation the body shrinks and becoming deeper. Also photophores begin to form and the eyestalks disappear totally. The fin rays complete, while the pectoral fins disappear during transformation. The female and male develop a gender dimorphism. While males form no barbel and do mostly have no pelvic fins, large postorbital photophores develop. Females form out a barbel on the lower jaw and have always a pelvic fin, but they develop only small postorbital photophores. Males transform at about 35 mm, while females do so with 45 mm. The female specimen studied was in a very late postflexion stage or already in the transformation since some photophores had already developed.

-- Pigmentation: A number of six spots form on isthmus and a series of melanophores develop ventrally along the gut.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Reference: GIBBS (1986b); KAWAGUCHI and MOSER (1984); RICHARDS (2006f);

-- Reference material: 1 specimen; 03/02: S6 (P2015104);

-- Distribution: The only specimen was caught at a mesopelagic station southeast of Fuerteventura at a depth of 617 – 915 m in daytime during late winter.

¬ AULOPIFORMES Larvae are elongated and slightly to strongly laterally compressed. After hatching the eyes are unpigmented, the mouth is not fully developed and the yolk sac has a small to moderate size. The preanal length is very variable and can extend to only less than one third to more than two thirds of the body length. Generally no spines are developed and the pigmentation is very rare, but latter can reach a full body coverage. Prominent for most families is the strong peritoneal pigmentation. The larvae of the suborder Synodontoidei are elongate with a round to ovoid cross section. The adipose fin is present. The pelvic fins form very anteriorly at the abdomen. Also small teeth on both jaws are very prominent in that suborder. The suborder Chlorophthalmoidei has in common that they have no swim bladder or fine spines and the pelvic fins are positioned at the abdomen. In most Chlorophthalmoidei the pigmentation is rare until transformation. The number of peritoneal patches is very variable.

-- Reference: DITTY, FAROOQI and SHAW (2006); WATSON (1996a);

SYNODONTOIDEI > Aulopidae Larvae are elongate with a large head and a curved head profile. Between anus and anal fin origin a large interspace is present. The peritoneal pigmentation is very distinctly developed and expands with larval development. Additionally a laterally patch on the caudal peduncle is very common. The larval development is direct and gradually.

-- Reference: AMBROSE (1996a); OKIYAMA (1984);

• Aulopus filamentosus Meristics: Dorsal rays:

Anal rays:

11 – 12

Pelvic rays:

Pectoral rays:

12 – 13

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

-- Remarks: The larvae were identified by family description and the very distinctive peritoneal patch and the pigmentation on the caudal peduncle (Figure 37). Only two Atlantic Aulopus species exist, whereas Aulopus cadenati POLL, 1953 is known to be restricted to the Southeast Atlantic.

-- Reference: SULAK (1986a);

-- Reference material: 2 specimens; 11/97: S4 (P2393621); S10 (P2393620);

-- Distribution: Both specimens were caught at neritic stations in autumn. At the station southeast of Fuerteventura the larva was present within a depth of 22 – 23 m during daytime and northeast of Lanzarote the larvae was found in a depth range of 20 – 64 m during the night.

Synodontidae Larvae of Synodontinae hatch with 2.5 – 3.8 mm and flexion occurs at a size of 7.0 – 10.2 mm. Body and gut are very elongate and round in cross section and latter has a length from 60 % – 90 % of BL. The head is round with round to slightly ovoid eyes and the snout is round too in early larvae, but the snout flattens dorsoventrally at midflexion stage. The mouth is large extending to the midlevel of the eye and is full with small teeth in both jaws.There is no apparent gas bladder. Prominent is the series of paired lateral gut blotches develops, which remain throughout the larval phase, but gets less distinct as they become internal. Additional pigments are positioned at the base of the anal fin and the tip of the notochord or at the caudal fin base. Shortly after flexion first fin rays develop at the anal fin and after their completion at a size of about 14 mm, the pelvic and pectoral fin rays formation starts. The anlagen of the short based dorsal fin are not visible until 13 mm and dorsal fin formation is completed with 20 mm.The adipose fin is present and forms at a size of 10 mm and is located directly above the anal fin. Larvae of most species reach a large size up to 40 mm.

-- Reference: LEIS and RENNIS (1983); STEVENS and MOSER (1996);

• Synodus sp. Most species of the genus Synodus have an extremely elongated body and the prominent ventral pigment blotches. The best tool for species identification is the use of the meristic characters and the number of the peritoneal dots in late larvae.

-- Reference: SULAK (1986b);

-- Reference material: 4 specimens; 11/97: S16 (P2393626-29);

-- Distribution: Synodus sp. was caught at a neritic station west of Gran Canaria at 42 – 66 m depth during the night in autumn.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

• Synodus saurus Meristics: Myomeres:

55 – 58

Dorsal rays:

11 – 13

Anal rays:

9 – 12

Pelvic rays:

Pectoral rays:

12 – 14

-- Morphology: The body is elongate with round to dorsolaterally flattened head and large mouth with teeth in both jaws. The eyes are round. The adipose fin is present. The sequence of fin formation starts with caudal and continuous with anal, dorsal, pectoral and ends with the pelvic fin.

-- Pigmentation: There is one blotch under the operculum and five more peritoneal dots dorsolaterally along the gut, which were clearly visible in the studied specimens (Figure 38). With the number of the spots it is easy to distinguish between S. saurus and S. synodus. Additionally a series of small spots develop along the anal fin base.

-- Reference: DITTY, FAROOQI and SHAW (2006); SULAK (1986b);

-- Reference material: 4 specimens; 11/97: S16 (P2393623-25); 04/99: S13 (P2393622);

-- Distribution: The larvae were caught in autumn during night at neritic stations, so west of Gran Canaria at 42 – 66 m depth and southwest of Gran Canaria at 33 – 53 m depth.

• Synodus synodus Meristics: Myomeres:

56 – 58

Dorsal rays:

11 – 14

Anal rays:

8 – 10

Pelvic rays:

Pectoral rays:

11 – 13

-- Morphology: The elongate body is round to dorsolaterally flattened in cross section. The head has a large mouth with teeth in both jaws. Eyes are round. Adipose fin is present. The sequence of fin formation starts with caudal and continuous with anal, dorsal, pectoral and ends with the pelvic fin.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

-- Pigmentation: One blotch is located under the operculum and 12 – 13 more peritoneal dotes dorsolaterally along the gut are present and were easy countable in the studied larvae (Figure 39). Additionally, a spot is positioned at the end of the anal fin and caudal fin base. Spots develop at the cleithrum and on the snout.

-- Reference: DITTY, FAROOQI and SHAW (2006); SULAK (1986b);

-- Reference material: 74 specimens; 11/97: S4 (P2393675-679); S6 (P2393649-55); S7 (P2393646-48); S10 (P2393656-74); S13 (P2393630-45); S16 (P2393681-703); 11/00: S13 (P2393680);

-- Distribution: The larvae were only caught in autumn mainly at neritic stations, so west and northwest of Gran Canaria in a depth from 37 – 66 m at night, southeast of Fuerteventura at a depth of 22 – 35 m in day and south- and northeast of Lanzarote at 3 – 41 m depth in day and 20 – 64 m at night. Some larvae were also found at an epipelagic station southeast of Fuerteventura at a depth of 90 – 107 m in the morning.

CHLOROPHTHALMOIDEI > Chlorophthalmidae Larvae have an extremely short gut and very large interspace from anus to the anal fin origin. A single peritoneal pigment section is located behind the pectoral fin base overlying the gut and an additionally patch lateral at the caudal fin base. The anal and dorsal fins have a short base, but the pectoral fins are well developed.The dorsal and pelvic fins are placed anteriorly of the body. Most larvae are caught close to the surface.

-- Reference: DITTY (2006a); OKIYAMA (1984); SULAK (1986c);

• Chlorophthalmus agassizii Meristics:

Development:

Vertebrae (Myomeres):

47 (46 – 48)

Dorsal rays:

10 – 11

Anal rays:

7–9

Flexion: 9.0 mm

Pelvic rays:

8–9

Transformation: 25 mm, gradually

-- Morphology: The body is slender and round in cross section. The preanal length is 45 % of SL during all larval stages. The large interspace between anus and anal fin origin also remains throughout larval development. The adipose fin is present and the pelvic fin bases are fused. The development is gradual and the eyes are migrating dorsally.

-- Pigmentation: The prominent single spot laterally at the midline of the caudal fin base is very distinctive and clearly visible in the studied larvae (Figure 40). With development further spots add. At early stages two additional spots dorsally and laterally of the caudal peduncle occur temporally. Also the internal peritoneal pigmentation is a very distinctive character and quite visible still in postflexion larvae (Figure 40) and remains until the transformation stage.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Reference: DITTY (2006a); FAHAY (1983); OKIYAMA (1984);

-- Reference material: 7 specimens; 11/97: S6 (P2393708-710); S13 (P2393704-705); S16; 11/00: S17 (P239370-707);

-- Distribution: All specimens were found at epipelagic stations in night or in the morning during autumn. Larvae were caught west of the channel between Lanzarote and Fuerteventura at 28 – 43 m in the morning, southeast of Fuerteventura at a depth of 90 – 107 m in the morning and southeast of Gran Canaria at a depth of 31 – 37 m during the night.

Notosudidae Larval features are similar throughout the whole family. The body is elongate and only slightly compressed, but with development is getting more lateral compressed towards the tail. Also the head is increasingly becoming depressed with growth. The wedge shaped snout is initially short and increases in length during development. The eye is ovoid with a longer horizontal axis but getting more roundish during transformation. On the slightly stalked eyes a conical mass of choroid tissue is positioned at posterior part. Except for Ahliesaurus the preanal length is about 40 -50 % of BL in early larvae and increases with growth up to 60 % of BL. A large interspace is present between anus and anal fin origin. Notosudids have no spines. The pigmentation is used as distinctive character for species identification. There is no peritoneal pigmentation developed and other pigmentation is rare. Melanophores are mostly reduced to the area of the caudal peduncle, on the finfold or fins before transformation. The only exception is the genus Ahliesaurus, which has a midlateral melanophore series. The length of flexion is at 10 – 12 mm and length of transformation takes place at 25 – 45 mm.

-- Reference: OKIYAMA (1984); WATSON and SANDKNOP (1996b);

• Ahliesaurus berryi Meristics:

Development:

Vertebrae (Myomeres):

47 – 50 (47 – 50)

Dorsal rays:

10 – 11

Anal rays:

19 – 21

Pelvic rays:

Pectoral rays:

10 – 12

Flexion: 10 - 12 mm

-- Morphology: The body is very elongated. The preanal length is at 57 – 60 % of SL. In the interspace of the origins of pelvic and dorsal fins lie one or one and a half myomeres, when the larva is more than 20 mm in length.

-- Pigmentation: The area of the caudal peduncle and caudal fin rays is full of small dots, which increase in number and extension during development. Additional pigmentation develops on anal and adipose fins. A distinctive feature for this species is the presence of six internal spots below the lateral midline, where the fourth spot is located above the anus. These small internal spots

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

were still visible in the studied postflexion larvae (Figure 41) and were an important diagnostic character together with the number of the myomeres for this species.

-- Reference: FAHAY (1983); KREFFT (1986a); RICHARDS (2006g);

-- Reference material: 2 specimens; 11/97: S10 (P2393711-712);

-- Distribution: Two specimens were caught at a neritic station northeast of Lanzarote at depths of 20 – 64 m during the night in autumn.

• Scopelosaurus sp. Most species, which occur in the North Atlantic, have 53 – 61 myomeres. The pelvic fins are positioned in front of the dorsal fin and 4 – 7.5 myomeres lie in between. The preanal length is shorter than that of Ahliesaurus berryi with about 40 % of SL. The number of vertebrae is more than 52. All species of that genus have the caudal peduncle pigment patterns.

-- Reference: FAHAY (1983); KREFFT (1986a);

-- Reference material: 3 specimens; 04/99: S5 (P2393713-714); 01/00: S6 (P2393715);

-- Distribution: One specimen was caught at a neritic station southeast to Gran Canaria at a depth of 10 – 70 m in the night during winter and other specimens were found southeast of Fuerteventura in a depth range from 382 – 511 m in day during spring.

ALEPISAUROIDEI > Scopelarchidae Except for Benthalbella the development is direct and transformation is completed at a length at 30 – >80 mm SL. Benthalbella retain its larval form in its prolonged period of growth till a very rapid transformation takes place. Newly hatched larvae are totally unpigmented (with some exceptions) and translucent. The mouth is partially formed, when recently hatched, and the preanal length is with 30 % of the body length short.The shape of the body is quite various within this family and can be deep bodied with a large head to moderately slender and elongate with a moderately small head, but they have a concave dorsal profile of the anterior body section in common. Also typical for this family is the low positioned large mouth with teeth in very early larvae and a long and wedge shaped to pointed snout. Another family feature is the peritoneal pigment except in Benthalbella and the ovoid eyes with the longer axis orientated vertically, but tends to incline forward. Species can only be distinguished with meristics, especially the myomere and the pectoral fin ray counts.

-- Reference: JOHNSON (1984a); WATSON and SANDKNOP (1996a);

-- Reference material: 1 specimen; 03/02: S20 (P2393719);

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Distribution: The scopelarchid was caught at a mesopelagic station southeast of Fuerteventura in a depth range from 308 – 498 m during the day in winter.

• Benthalbella infans Meristics:

Development:

Vertebrae:

55 – 58

Dorsal rays:

8–9

Anal rays:

20 – 25

Pelvic rays:

Pectoral rays:

25 – 28

Flexion: 15 mm Transformation: 50– 76 mm; rapidly

-- Morphology: The pelvic fins are anteriorly positioned to the pectoral fins and are also longer than the latter ones. Adipose fin is elongated throughout the larval development. The head length is about 30 % of SL.

-- Pigmentation: No pigmentation develops until transformation, except for the choroid tissue. The missing pigmentation and the myomere and pectoral fin ray count leaded to this species.

-- Reference: DITTY (2006b); DITTY (2006d); FAHAY (1983); JOHNSON (1986a);

-- Reference material: 2 specimens; 01/00: S1 (P2393716); S3 (P2393717);

-- Distribution: During winter two specimens were found at mesopelagic stations in daytime, so one east of Gran Canaria at a depth of 467 – 566 m and another north of Gran Canaria at a depth of 356 – 402 m.

• Scopelarchus analis Meristics:

Development:

Vertebrae:

44 – 49

Dorsal rays:

7–9

Anal rays:

21 – 26

Pelvic rays:

Pectoral rays:

18 – 22

Hatching: < 4.8 mm Flexion: 10 – 13 mm Transformation: 25 – 55 mm; gradually

-- Morphology: The shape of the body is moderately elongate, but anteriorly deeper. The snout is long. The teeth are well developed and very outstanding (Figure 42).The short based dorsal fin is located far anterior of the body. Pelvic fins have a long base, where the origin lies under the dorsal fin. The origin of the anal fin is at about 50 % of the SL.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

-- Fin formation sequence: Caudal fin, dorsal fin, anal fin, upper pectoral fin rays, pelvic fin and lower pectoral fin rays;

-- Pigmentation: Until the postflexion stage there is no pigmentation except for the choroid tissue. The peritoneal patch develops with a size of 10 mm and is situated above anterior middle of gut and shifts forward to the level of the pectoral fins at about 15 mm. Additionally a pair of dorsolateral peritoneal patches above the hindgut between pelvic fins and anus develops at 20 mm. During transformation stripes form above and below the lateral midline of the posterior body at a size of 25 mm. Further pigments develop on pelvic fin base after 25 mm and on snout and jaws with 30 mm. The peritoneal pigment sections fuse at a size of 33 – 55 mm. In the studied specimen these peritoneal pigment sections were already fused, though not clearly visible. The species identification resulted from the meristic counts alone.

-- Reference: DITTY (2006b); DITTY (2006d); JOHNSON (1986a); WATSON and SANDKNOP (1996a);

-- Reference material: 1 specimen; 04/99: S5 (P2393718);

-- Distribution: The only specimen was caught southeast of Fuerteventura at a mesopelagic station at a depth of 382 – 511 m during the day in spring.

Alepisauridae > Alepisaurinae The larvae are moderately elongate with a large curved head. The preanal length increases with development from 37 % of BL in preflexion to 81 % in transforming larvae. Prominent for that family are the big canine teeth on the dentary. All fins are small, only the pectoral fins are moderate in size and pigmented. The peritoneal pigmentation is present, but indistinct.

-- Reference: AMBROSE (1996b); JOHNSON (1984);

• Alepisaurus ferox Meristics:

Development: Hatching: < 5 mm

Dorsal rays:

26 – 45

Anal rays:

14 – 19

Pelvic rays:

8 – 10

Pectoral rays:

12 – 15

Flexion: 6 – 8.5 mm Transformation: 16 – 30 mm; gradually

-- Morphology: The body is short with a large, broad head and canine teeth on lower jaw. The latter character is the most important one leading to the genus Alepisaurus. Four preopercular spines develop at flexion stage and distinct bony ridges dorsally on the head, what was not well developed in my specimen. The pectoral fin tips are close to the anus. The dorsal fin is long based. The adipose fin is present.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Pigmentation: During preflexion stage small dots are present on pectoral fins. In postflexion stage the pigmentation of anal fin origin and dots dorsally on head are prominent. A large blotch is present at adipose fin origin and below. Melanophores are located dorsolaterally on the trunk. The entire pigmentation pattern is well developed in the studied specimen. During the transformation the spots spread dorsally and laterally along dorsum and tail.

-- Reference: AMBROSE (1996b); DITTY (2006b); FAHAY (1983); JOHNSON (1984); POST (1986a);

-- Reference material: 1 specimen; 11/00: S12 (P2393720);

-- Distribution: The only specimen was caught west of Gran Canaria within a depth range from 444 – 900 m at late daytime in autumn.

Omosudinae Larvae have a very large head with a straight head profile and a pointed snout.The mouth is very large too with large canine teeth on dentary and palatine.The body is short and stout, but becomes moderately elongate and laterally strongly compressed in adult stage. The fins are small, especially the pectoral fins. The peritoneal pigment sections have small interspaces. Atlantic species differ from Pacific ones in the smoothed or armed edges of head. A different intense pigmentation at the same sizes occurs and a pigmentation band above posterior part of anal fin are absent in Atlantic species and present in Pacific species. The transformation is gradual.

-- Reference: OKIYAMA (1984); POST (1986b);

• Omosudis lowii Meristics: Vertebrae (Myomeres):

39 – 41 (39 – 41)

Dorsal rays:

9 – 11

Anal rays:

13 – 14

Pelvic rays:

Pectoral rays:

12 – 13

-- Morphology: The body is short and the head is very large with an almost straight head profile. The very large mouth is full with canine teeth on the palatines and dentaries (Figure 43), which made it easy to distinguish from other Alepisauridae as Alepisaurus. Tiny teeth develop on the premaxillaries. Larvae have small pectoral fins with the anus far behind the tips. The dorsal fin is short and its origin is located after midbody level. The pelvic fins are situated at the same level as the dorsal fin is. The rays of the dorsal, anal and caudal are formed by 11.8 mm SL. The transformation is gradual.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

-- Pigmentation: Three to four peritoneal gut patches form temporary at a size of 6 mm and vanish at 30 mm. Rare pigmentation occurs in the area of the caudal peduncle. Prominent are the four to five small dots along the ventral side posteriorly to the anus in smaller larvae.

-- Reference: FAHAY (1983); POST (1986b);

-- Reference material: 1 specimen; 11/00: S12 (P2393721);

-- Distribution: A single specimen was caught west of Gran Canaria in depths from 444 – 900 m in daytime during autumn.

Paralepididae The subfamilies of Paralepidinae and of Sudinae form this family. The Sudinae differ by having a short body with long pectoral fins and a large head. The peritoneal patches forms earlier and serrated ridges develop along the ventral margin of the preopercle, snout and above the eyes. Paralepidinae are in general more elongated with a short trunk throughout the whole development. The myomere counts are higher than in Sudinae and the pigmentation is rarer. Prominent throughout the whole family are the peritoneal pigment patches, which increase in number with the lengthening of the gut. Other pigments are located on top of head, at the dorsal and anal fin base, in the area of caudal peduncle region and during transformation dorsal pigments add. The head is large with a length up to 40 % of SL having an elongated pointed snout. The eyes are slightly to strongly ovoid, but becoming round during transformation. Transformation takes place, when the black peritoneum is fully developed.

-- Reference: DITTY (2006c); OKIYAMA (1984); POST (1986c);

• Lestidiops sp. The body is elongated, but the head with the large and round eyes is deep in early larvae and increases in length with development. The short gut has no peritoneal pigmentation when the larva hatches, but melanophores appear when the gut grows in length during development. The anal fin is far back on the body and develops 25 – 35 rays. The pelvic fins are situated far posteriorly close to the anal fin origin.

-- Reference: OLIVAR and FORTUÑO (1991); POST (1986c);

-- Reference material: 49 specimens; 11/97: S6 (P2393745-759); S10 (P2393725-729); S13 (P2393730-742); S18 (P2393743); 04/99: S6 (P2393723); S12 (P2393744); S16 (P2393724); 05/99: S1 (P2393760-763); S3 (P2393770); 11/00: S17 (P2393764-768); 03/02: S18 (P2393722); S24 (P2393769);

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Distribution: The specimens were caught in autumn, winter and spring. In autumn larvae were mainly present in the morning time at epipelagic stations west of the channel between Fuerteventura and Lanzarote at 28 – 43 m, southeast of Fuerteventura at 90 – 107 m and south of Gran Canaria at 31 – 37 m. Some were also found at neritic stations, so northeast of Lanzarote at 20 – 64 m depth at night and east of Gran Canaria in a depth of 116 – 146 m also at night. In winter specimens were caught southeast of Fuerteventura at 470 – 695 m during daytime and at 565 – 980 m during the night. In spring late larvae occurred during the night at epipelagic stations southeast of Fuerteventura and Gran Canaria in depths of 23 – 52 m and 38 – 49 m respectively. Further specimens were found southwest of Gran Canaria at a depth of 18 – 61 m in night. In late spring larvae were present only at epipelagic stations, so far east of Fuerteventura at a depth of 38 – 40 m in the dawning and far north of Gran Canaria at a depth of 42 – 47 m in night.

• Lestidiops sphyrenoides Meristics: Vertebrae:

89 – 92

Dorsal rays:

9 – 11

Anal rays:

28 – 31

Pelvic rays:

Pectoral rays:

-- Morphology: The shape of the body is elongate. The anal fin is short and positioned slightly behind the midpoint of the body. This species is easy to distinguish from other Lestidiops species by the counts of the fin rays.

-- Pigmentation: Prominent are the peritoneal patches. For further information, look at the family description.

-- Reference: POST (1986c);

-- Reference material: 1 specimen; 05/99: S6 (P2393771);

-- Distribution: The only specimen was present in late spring at an epipelagic station far north of La Gomera and Tenerife at a depth of 48 – 66 m during the night.

• Macroparalepis sp. The larvae are very elongate with a small head and large round eyes. When larvae are more than 10 mm in length the gut is curved under the head. The peritoneal pigment sections can reach a maximum number of 10, with one exception in M. affine EGE, 1933 with a maximum of 12. Additional pigmentation is distributed on the caudal fin. Anal fin ray counts range from 22 – 28 and the vertebrae from 80 – 102 in species of the eastern north Atlantic.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

-- Reference: FAHAY (1983); POST (1986c);

-- Reference material: 2 specimens; 03/02: S11 (P2393772-773);

-- Distribution: The specimens were only found in winter at an epipelagic station southeast of Fuerteventura in a depth range of 31 – 140 m during the day.

• Sudis hyalina Meristics:

Development: Hatching: < 3.8 mm

Dorsal rays:

12 – 16

Anal rays:

21 – 24

Pelvic rays:

Pectoral rays:

13 – 15

Flexion: 7 mm Transformation: 20 mm

-- Morphology: The body is short with a large head and an elongate snout. The long pectoral fins are elongated and can reach up to the level of the dorsal fin origin, which was clearly visible in the studied specimen (Figure 44). The gut is long and the peritoneal pigment sections are early developed. Along the ventral edge of the preoperculum spines develop. On the angle of the preoperculum a big spine develop, which is smooth with one retrorse hook, while S. atrox ROFEN, 1963 has serrated edges with one or more antrorse hooks. The myomere count is low.

-- Pigmentation: In preflexion stages 3 – 4 patches develop and increase in number up to 7 – 8 patches in late larval stage. The pigment pattern on the trunk is patchy. Additional pigmentation is on the pectoral fin present already in early stages and at the bases of dorsal, anal and caudal fins. In transforming larvae 6 patches are at the dorsal line and one patch above and below the midline of the caudal peduncle, which is already clearly visible in the studied specimen (Figure 44).

-- Reference: DITTY (2006c); FAHAY (1983); OKIYAMA (1984);

-- Reference material: 1 specimen; 11/97: S6 (P2393774);

-- Distribution: The only specimen was caught at an epipelagic station west of the channel between Fuerteventura and Lanzarote at a depth of 28 – 43 m at late night in autumn.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

¬ MYCTOPHIFORMES This order consists of two families Neoscopelidae and Myctophidae. The species inhabit pelagic or benthopelagic deep sea habitats. Only six species in three Genera exist in the family of Neoscopelidae in the Atlantic, Pacific and Indian Ocean. Myctophidae are highly diverse with more than 235 in 32 genera, but all are luminous and occur primarily midwater of all oceans.

-- Reference: MOSER and WATSON (2001); MOSER and WATSON (2006a);

Myctophidae The yolk sac larvae hatch with a length of 2 mm and the notochord flexion occurs within a size range from 0.5 – 2 mm. The size at midflexion is about the half of the maximal larval size. Also transformation takes place within a short length interval of 2 mm and in most myctophid larvae it occurs at 12 – 19 mm, but ranges from 10 – 30 mm. The shapes of the head, body and gut are characteristic for most species and within most genera similarities in the overall habit exist. Most larvae are moderately slender, with the exceptions of Hygophum reinhardtii, which is very elongate and some species of Myctophum and Lampanyctus, which are robust.The shape of the eyes is very diverse in development and has therefore a diagnostic character, at least for sorting the larvae into subfamilies. Most of Lampanyctinae possess round and mainly sessile eyes, with the exception of the genera Notolychnus, whereas larvae of Myctophinae have elliptical, stalked eyes with a choroid tissue on the ventral margin. In most myctophid larvae the s-shaped gut extends to about the midpoint of body, where its form and length can be used as further diagnostic feature. With the exception of two genera of Notolychnus and Taaningichthys, myctophids develop first the Br2 photophore during the larval stage (see Figure 9). It is located posteroventral to the orbit and it seems to migrate to the branchiostegal membrane beneath the orbit during transformation. The sequences of the development of the further photophores are useful diagnostic characters. For the identification to generic and species level the melanophore pattern can be a very useful character. Because myctophids species possess distinct pigmentation, it is useful for identification to species level, with the exception of the genus Diaphus, where only some patterns are identified.

Figure 9. Schematic illustration of an adult Myctophidae with photophores labelled (after HULLEY 1986)

-- Reference: HULLEY (1986); MOSER and WATSON (2001); MOSER et al. (1984);

-- Reference material: 2 specimens; 11/97: S6 (P2393907); 01/00: S3 (P2393908);

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

-- Distribution: The specimens were caught north of Gran Canaria at a depth of 356 – 402 m during the day in winter and at an epipelagic station west of the channel between Fuerteventura and Lanzarote in depths of 28 – 43 m during late day in autumn.

Myctophinae • Benthosema suborbitale Meristics:

Development:

Vertebrae:

33 – 35

Dorsal rays:

11 – 14

Anal rays:

16 – 19

Pelvic rays:

Pectoral rays:

12 – 15

Hatching: ~2.0 mm Flexion: 5 – 7 mm Transformation: ~10.0 mm

-- Morphology: The body is short and deep.The gut is short with a deflected terminal section, which is disappearing in late postflexion stage. The eyes are narrow with a lunate choroid tissue at the ventral margin.

-- Pigmentation: A pair of melanophores is embedded just anterior to cleithral symphysis at 4 mm and coalesces in midline of lower jaw symphysis in further development. Two embedded blotches are located anterior to pectoral fin base under the opercle.

-- Photophore development: The Br2 forms at 5 mm, PO1, and PO2 at 9 mm.

-- Reference: FAHAY (1983); HULLEY (1986); MOSER and WATSON (2001); MOSER and WATSON (2006b); MOSER et al. (1984);

-- Reference material: 3 specimens; 05/99: S3 (P2393777); 01/00: S5 (P2393775-776);

-- Distribution: The larvae were caught in winter south of Gran Canaria at 49 – 510 m during the day and in late spring at an epipelagic station north of Gran Canaria at a depth of 42 – 47 m during the night.

• Diogenichthys atlanticus Meristics:

Development:

Vertebrae:

31 – 35

Dorsal rays:

10 – 12

Anal rays:

14 – 18

Pelvic rays:

Pectoral rays:

12 – 15

Hatching: < 2.9 mm Flexion: 6.0 – 6.9 mm Transformation: 13.5 – 14.5 mm

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Morphology: The body is moderately slender in preflexion stage and increases in depth with growth. The snout is acute becoming relatively shorter with further development.The eyes are initially elliptical, but get rounder in later stages.The ventral choroid tissue is absent. The prominent symphyseal barbel forms at 5 mm and is present till transformation. This barbel together with the pigmentation pattern led me to this species.

-- Pigmentation: Three dots form on the symphyseal barbel. One pair of melanophores is embedded above hindbrain. A ventrolateral pair of spots is located just posterior to the cleithrum. Further a dorsolaterally pair on terminal section and two pairs are placed along the midgut. These spots can increase in number up to six in late postflexion larvae. A series of melanophores forms out just posterior to the anus, which increases in number up to 12 in postflexion stage. Further single spots form out just anterior to the pectoral fin base as well as at the base of caudal fin in flexion larvae and posterior to the dorsal and adipose fin in large larvae. Additionally pairs of melanophores form at the bases at the anal fin rays in early postflexion larvae, which increase in number with further development.

-- Photophore development: Br2 forms at 6 mm, PO2 at 7 mm, PO5 at 8.5 mm, and AOa1 at 11 mm.

-- Reference: FAHAY (1983); HULLEY (1986); MOSER and WATSON (2001); MOSER and WATSON (2006b); MOSER et al. (1984);

-- Reference material: 2 specimens; 04/99: S14 (P2393802); 01/00: S1 (P2393801);

-- Distribution: Larvae were caught only during the day east of Gran Canaria at 467 – 566 m in winter and south of Gran Canaria in a depth of 480 – 694 m in spring.

• Hygophum hygomii Meristics:

Development:

Vertebrae:

36 – 38

Dorsal rays:

13 – 15

Anal rays:

20 – 22

Pelvic rays:

Pectoral rays:

14 – 17

Flexion: 6.0 – 7.0 mm Transformation: 13 – 14.5 mm

-- Morphology: Eyes are slightly elliptical and unstalked with a brownish choroid tissue at the ventral margin.The body and gut are moderately long with a distance from snout to anal fin origin (Sn - A) less than 60 % of SL. The Br2 photophore forms lately in larval period.

-- Pigmentation: The prominent series of pairs ventrally on isthmus und posterior to cleithrum is present. One large melanophore is positioned dorsolateral on the terminal gut section. In some larvae a spot occurs on lower jaw.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

-- Remarks: The larvae were easy to confuse with H. benoiti (COCCO, 1838), but Sn - A is more than 60 % of SL in postflexion larvae. With this character it was easy to differentiate in the studied larvae. The eyes are narrower and the snout is more pointed than in H. hygomii.

-- Reference: FAHAY (1983); HULLEY (1986); MOSER and WATSON (2001); MOSER and WATSON (2006b); MOSER et al. (1984);

-- Reference material: 18 specimens; 11/97: S4 (P2393808); S10 (P2393803-807); 04/99: S5 (P2393809); 01/00: S1 (P2393810-818); S5 (P2393819); 11/00: S19 (P2393820);

-- Distribution: In spring a transforming specimen was caught at a mesopelagic station southwest of Fuerteventura at 382 – 511 m in daytime. In autumn larvae occurred southeast at the neritic area of Fuerteventura at a depth of 22 – 35 m at day, northeast at the neritic area of Lanzarote at 20 – 64 m at night and one transforming larva in a mesopelagic depth range of 496 – 630 m during the day. In winter time specimens were found east of Gran Canaria at 467 – 566 m at late daytime and one transforming specimen in a depth of 490 – 510 at daytime.

• Hygophum reinhardtii Meristics:

Development:

Vertebrae:

38 – 40

Dorsal rays:

13 – 15

Anal rays:

21 – 25

Pelvic rays:

Pectoral rays:

13 – 16

Hatching: < 3.4 mm Flexion: ~8.8 – 10.3 mm Transformation: 15.0 – 16.4 mm

-- Morphology: The head is long and flat.The eyes are on short stalks, which were not distinct in the studied specimens, and very ovoid with a choroid tissue on the ventral margin.The body is very slender with an elongate gut, which was the most important character together with the pigmentation pattern for diagnosis. Only H. reinhardtii has this slender body shape with a straight gut and this kind of pigmentation pattern, which cannot be confused with another pattern of a myctophid (Figure 45).

-- Pigmentation: The pigmentation pattern consists of a median spot on the isthmus and three pairs of melanophores posterior to the cleithrum. One spot is located on the opercle. Series of paired melanophores form along the gut with a number of pairs up to 8 and 15 along the anal fin base. As further pigmentation 5 – 10 myospetum dashes are situated from the posterior part of the anal fin base to the caudal fin base, where one melanophore is located on the midline of the caudal peduncle. All pigmentation was clearly visible in the studied specimens and essential for identification.

-- Reference: FAHAY (1983); HULLEY (1986); MOSER and WATSON (2001); MOSER and WATSON (2006b); MOSER et al. (1984);

-- Reference material: 5 specimens; 11/97: S10 (P2393823-824); S13 (P2393821-822); 11/00: S19 (P2393825);

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Distribution: All specimens were caught in autumn, so at a neritic station northeast of Lanzarote at 20 – 64 m in night and at an epipelagic southeast of Fuerteventura at 90 – 107 m depth at late night, almost morning. One transforming specimen was found southwest of Gran Canaria in a depth of 496 – 630 m during daytime.

• Symbolophorus veranyi Meristics:

Development:

Vertebrae:

39 - 40

Dorsal rays:

12 – 14

Anal rays:

21 – 23

Pectoral rays:

12 – 14

Transformation: ~20 mm

-- Morphology: The body is moderately elongate. The head is broad and flat with a pointed snout. The narrow eyes are placed on short stalks and have a small choroid tissue, which is characteristic for this species and clearly visible (Figure 46). The pectoral fins are enlarged with a very high number of fin rays and a wing shaped base.The rays extend till beyond the anus.The latter was not always present in the studied specimens, since the rays are very fragile. Only one specimen still had elongated pectoral fin rays reaching beyond the anus (Figure 47). Also the pelvic fins are large and form early.

-- Pigmentation: Spots are situated on the tip of the upper and lower jaw. A large melanophore is located on the posterior edge of the opercle and a few spots along the gut with one spot placed on the terminal gut. Further pigments are at the base and rays of the pectoral fin. In general the number of pigments is decreasing towards the end of larval development, so the pigmentation was quite reduced in the studied specimens (Figure 46 –Figure 47).

-- Photophore development: Only Br2 forms at 12 mm during larval stage.

-- Reference: FAHAY (1983); HULLEY (1986); MOSER and WATSON (2001); MOSER et al. (1984); OLIVAR and FORTUÑO (1991);

-- Reference material: 2 specimens; 11/97: S6 (P2393905); 04/99: S12 (P2393895); S14 (P2393894); S16 (P2393896-897); 05/99: S5 (P2393898); S6 (P2393899-904);

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

• Myctophum punctatum Meristics:

Development:

Vertebrae:

Dorsal rays:

13 - 14

Anal rays:

20 – 22

Pectoral rays:

14 – 15

Flexion: 7 mm Transformation: ~ 21 – 22 mm

-- Morphology: The body is elongate with a pointed, flat snout. The stalked eyes are narrow and have a conical choroid tissue. The pectoral fins are often enlarged and broad with wing shaped bases, but this characteristic was not as distinct as in Symbolophorus veranyi in the specimens studied.

-- Pigmentation: Each a series of dots is present along the edges of upper and lower jaw. Several spots are placed on the upper edge of the opercle and a series of spots extends ventrally from the head to the anus. A melanophore is located at the base of the pectoral fins and a few along its rays. Further pigments often form on the posterior rays of the dorsal, anal and adipose fin. One single spot is situated middorsally anterior to the adipose fin base and midventrally an embedded melanophore at the end of the anal base as well as at the caudal fin base. Characteristic for this species is the spot at the midline of the caudal fin base, which was clearly visible in the studied larva.

-- Reference: FAHAY (1983); HULLEY (1986); MOSER et al. (1984); OLIVAR and FORTUÑO (1991);

-- Reference material: 1 specimen; 04/99: S12 (P2393830);

-- Distribution: A single specimen was caught at an epipelagic station southwest of Gran Canaria at a depth of 38 – 49 m during night in spring.

• Centrobranchus nigroocellatus Meristics:

Development:

Vertebrae:

35 – 40

Dorsal rays:

9 – 11

Anal rays:

16 – 19

Pelvic rays:

Pectoral rays:

13 – 17

Hatching: < 2.8 mm Flexion: 5.4 – 6.3 mm Transformation: ~ 12.0 mm

-- Morphology: The body is deep and laterally compressed. The head is large with a bulbous snout, which was very distinct in the studied specimens as well as the shape of the eyes, which is narrow with an extremely elongate and unpigmented conical choroid tissue. The terminal gut is slightly deflected. The dorsal and ventral finfold are enlarged.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Pigmentation: There is midlaterally pigmentation on the isthmus present and one pair of spots posteriorly of the cleithrum. Melanophores are located on the branchiostegal membrane. Further spots form at the posterior margin of the orbit at 12 mm.

-- Photophore development: Only Br2 forms at 5 mm during larval stage.

-- Reference: HULLEY (1986); MOSER and WATSON (2001); MOSER and WATSON (2006b); MOSER et al. (1984);

-- Reference material: 2 specimens; 11/97: S13 (P2393778-779);

-- Distribution: Two specimens were caught at the epipelagic station southeast of Fuerteventura at 90 – 107 m at the dawning during autumn.

Lampanyctinae • Diaphus sp. Two types of Diaphus are described, where one type has a slender body, a small head and develop melanophores postanally along the midventral line. The second one has a deeper body, a bulbous head and a single spot on caudal fin base. Both have in common, that they are not or rarely pigmented on head, but possess melanophores at the midgut region, at the anteroventral surface on liver and on gas bladder. Further embedded spots at the base of caudal fin rays are common. The genus of Diaphus develops the most photophores during larval stages.The photophore Br2 and/or PO5 forms first and PO2 secondly. Afterwards the rest of PO series, VO1, VO5, OP2, VLO and PVO form out.

-- Reference: MOSER and WATSON (2001); MOSER et al. PAXTON (1984);

-- Reference material: 8 specimens; 11/97: S4 (P2393800); S10 (P2393793-799);

-- Distribution: The larvae were only caught at neritic stations in autumn, so southeast of Fuerteventura at 22 – 35 m depth at day and northeast of Lanzarote at a depth at 20 – 64 m at night.

• Notoscopelus sp. The shape of the body is moderately deep with a relatively large head and large eyes. The gut is short in preflexion stage, but increases in length during flexion. Pigmentation occurs at the tips of both jaws, above the brain, on the gas bladder, and laterally at the cleithral region in early postflexion larvae. Additional melanophores form out along the lower jaw, on the hindbrain and on the nape. A series of spots are located along each side of the dorsal midline starting from the midbody and expend along entire dorsum. Some melanophores form out on pelvic and anal fin rays in some species in late postlarvae. Embedded myoseptal dashes develop on trunk and also at the caudal peduncle in postflexion stages in the specimens

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

studied. Blotches are situated along the septum and along the anal and caudal fin base with further pigmentation on the base of caudal, pelvic and anal fin rays in late postflexion stage of some species.

-- Reference: MOSER and WATSON (2001); MOSER et al. (1984);

-- Reference material: 5 specimens; 04/99: S14 (P2393892-893); 05/99: S4 (P2393891); 01/00: S1 (P2393889); S3 (P2393890);

-- Distribution: All studied specimens of that genus were caught exclusively at mesopelagic depths around Gran Canaria in winter and spring. At winter time larvae were caught east and north of the island at 467 – 566 m depth in late daytime and at 356 – 402 m during the day. In spring specimens occurred south and far north off the island at 480 – 694 m and at 517 – 591 m during the day, respectively.

• Notoscopelus resplendens Meristics:

Development:

Vertebrae:

35 – 38

Dorsal rays:

21 – 24

Anal rays:

17 – 20

Pelvic rays:

Pectoral rays:

11 – 13

Hatching: < 2.4 mm Flexion: ~ 5.0 – 6.5 mm Transformation: ~ 20.0 mm

-- Morphology: The body is initially slender and the head relatively small, but both becoming deeper and laterally compressed with further development. The snout is acute in preflexion stage, but forms blunt after flexion. The teeth are well developed and hooked posteriorly on lower jaw. The eyes are large and almost round. The gut is short and strongly sigmoid. The longer dorsal fin has a higher ray count than the anal fin.

-- Pigmentation: Melanophores are positioned at the tips of jaws and at the cleithrum. A transverse pair above the midbrain develops in some larvae and a series of melanophores along the hypaxial region above the anal fin base. A paired series form on the dorsum extending to the adipose fin was found in most of the studied specimens.The sequence of melanophore development starts with a pair above cerebellum at 4 mm, at 5 mm one spot on midline of the nape and one further anteriorly to the midbrain. Several dashes form along the lateral midline of midbody, where it starts at the level of the anterior margin of the dorsal fin. Further a series of melanophores develop along the anal fin base at 5mm. With 8.5 mm pigmentation occurs at the angle of the jaws in some larvae and at 9.5 mm one or more melanophores on the throat or in some specimens also anterior to the forebrain. At a size of 12.5 mm pigmentation on the pectoral and anal fin rays and at the hypural margin occurs. At 14.5 mm melanophores on the edge of branchiostegal membrane appear. Most of the studied specimens were larger than 14.5 mm and had developed the entire pigmentation pattern.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Photophore development: The Br2 photophore forms at 4.2 mm, followed by PO5 at 6.2 mm, Vn at 9.2 mm, and PLO at 16.2 mm.

-- Remark: To distinguish late postflexion larvae of N. resplendens from other species of that genus, counting the gillrakers is necessary as it is in adult identification (WIENERROITHER, pers. comment). The numbers of gillrakers of N. resplendens of the upper and lower first arch are 5 - 7 + 1 + 13 - 16 and total 19 – 23.

-- Reference: FAHAY (1983); HULLEY (1986); MOSER and WATSON (2001); MOSER and WATSON (2006b);

-- Reference material: 57 specimens; 11/97: S13 (P2393832-839); 04/99: S12 (P2393840-842); S14 (P2393843-846); S16 (P2393847-855); 05/99: S4 (P2393856); S6 (P2393857-878); 01/00: S1 (P2393879-881); S7 (P2393882); 03/02: S3 (P2393883); S14 (P2393884); S24 (P2393885-888);

-- Distribution: In spring time specimens were caught around Gran Canaria, so southwest of the island at a neritic station at 18 – 61 m at night and at an epipelagic station at 38 – 49 m at night. At the day specimens were caught south of the island at a depth of 480 – 694 m. During late spring larvae occurred far north of Gran Canaria at 517 – 591 m during the day and at an epipelagic station north off La Gomera and Tenerife in a depth range of 48 – 66 m depth at night. In autumn some larvae were caught at the epipelagic station southeast of Fuerteventura at 90 – 107 m at late night. In winter specimens were mainly caught at mesopelagic stations, so east and west of Gran Canaria in depths of 467 – 566 m at late night and 578 – 610 m, respectively, where at the latter station one transforming specimen occurred. Further larvae were found south and southeast of Fuerteventura in depths of 440 – 665 in late daytime and 565 – 980 m in the night, respectively. One specimen was caught at a neritic station southeast of Fuerteventura in a depth range of 15 – 43 m during the night.

• Lampadena sp. Pigments develop above the brain, gut and gas bladder. Most species have large melanophores along the dorsal midline and postanally on the ventral midline. The other species form smaller, but more numerous melanophores in the dorsal and ventral series. Further pigments are embedded above the spinal column in some species. The sequence of the early forming photophores is Br2, PO5 and PLO.

-- Reference: MOSER and WATSON (2001); MOSER and WATSON (2006b); MOSER et al. (1984);

-- Reference material: 1 specimen; 03/02: S11 (P2393826);

-- Distribution: The only specimen was caught at an epipelagic station southeast of Fuerteventura at 31 – 140 m depth during the day in winter.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

• Ceratoscopelus warmingii Meristics:

Development:

Vertebrae:

35 – 36

Dorsal rays:

13 – 15

Anal rays:

13 – 15

Pelvic rays:

Pectoral rays:

12 – 15

Hatching: < 3.0 mm Flexion: ~ 5.0 – 6.0 mm Transformation: > 15.0 mm

-- Morphology: The body is slender with a small head. The eyes are moderately large and round to slightly ovoid. The snout becomes blunt in late larvae. The gut is slightly sigmoid and reaches to midbody level.

-- Pigmentation: A pair of spots forms on the terminal gut, which is very distinct (Figure 45). One to four spots above the spinal column of the caudal peduncle as indicated in the literature were not visible in the studied specimens. Also, a single dot located just posterior to the anal fin base was not clearly visible in the studied material. Embedded melanophores were weak but present these specimens in the optic region and also above the hindbrain in some late larvae.

-- Photophore development: Br2 and Vn form at 5 mm, while PO2 and PLO form first during transformation.

-- Reference: FAHAY (1983); HULLEY (1986); MOSER and WATSON (2001); MOSER and WATSON (2006b); MOSER et al. PAXTON (1984);

-- Reference material: 8 specimens; 11/97: S4 (P2393785); S9 (P2393792); S10 (P2393786-789); S13 (P2393790-791);

-- Distribution: The larvae were exclusively caught during autumn and mainly at neritic stations southeast of Fuerteventura and northeast of Lanzarote at a depth of 20 – 64 m during night and day. Further larvae were found at an epipelagic station southeast of Fuerteventura at 90 – 107 m depth and at a mesopelagic station northeast of Lanzarote at a depth of 527 – 557 m in early night.

• Ceratoscopelus maderensis Meristics:

Development:

Vertebrae:

35 – 38

Dorsal rays:

13 – 15

Anal rays:

13 – 15

Pelvic rays:

Pectoral rays:

13 – 14

Hatching: < 5.0 mm Flexion: ~ 6.0 mm Transformation: ~16.0 mm

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Morphology: The body is moderately slender with a relatively small head. The eyes are round and possess a choroid tissue. The gut is slender and slightly s-shaped.

-- Pigmentation Dots are present in the occipital region, laterally on the foregut and at the terminal section. Three or four larger spots are positioned on the dorsal midline of the caudal peduncle, which is the main character to distinguish from C. warmingii, were clearly visible in the studied specimens. Smaller melanophores develop on the dorsal midline of caudal peduncle too, but becoming, except for a few, embedded later.

-- Photophore development: Br2, Vn, PLO and PO5 form at 7 – 11 mm.

-- Remarks: Easy to confuse with Lepidophanes guentheri (GOODE & BEAN, 1986), but the melanophores laterally on foregut are lacking in that species. Further is C. maderensis deeper bodied than L. guentheri.

-- Reference: FAHAY (1983); HULLEY (1986); MOSER and WATSON (2001); MOSER and WATSON (2006b); MOSER et al. PAXTON (1984);

-- Reference material: 5 specimens; 05/99: S1 (P2393780-781); S2 (P2393782); 11/00: S12 (P2393783-784);

-- Distribution: Larvae were caught in late spring far east of Fuerteventura during the late night from 38 – 40 m and during the day at 495 – 561 m depth. In autumn juveniles were found in depths from 444 – 900 m west of Gran Canaria during the late day.

• Lampanyctus sp. The body is deep with a large head.The jaws are in some species elongate with distinctive teeth.The transformation is abrupt. Only Br2 forms during larval stage. Pigmentation patterns are very divers. Melanophores can develop on tip of the lower jaw, between the eyes, on the back and side of the head, at the adipose and the pectoral fin, along the myosepta and in the cleithral region.

-- Reference: MOSER and WATSON (2001); MOSER and WATSON (2006b); MOSER et al. (1984);

-- Reference material: 1 specimen; 05/99: S6 (P2393828);

-- Distribution: One specimen was found at an epipelagic station nor th of La Gomera and Tenerife at 48 – 66 m depth in night during late spring.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

• Lampanyctus crocodilus Meristics:

Development:

Vertebrae:

35 – 37

Dorsal rays:

13 – 15

Anal rays:

16 – 18

Pelvic rays:

Pectoral rays:

13 – 16

Flexion: ~ 6 – 7 mm Transformation: ~20.0 mm

-- Morphology: The body is relative deep with a short trunk. The eyes are round and have no choroid tissue.

-- Pigmentation: Some melanophores form dorsally on the head and blotches in the occipital region. Dashes are embedded at the peritoneum and at the anteriorly myosepta and increase in their numbers with further development. A large melanophore is present in the occipital region as well as on the terminal gut. Some additional pigments are located on the pectoral fin base and blotches along the dorsal midline between the dorsal and the adipose fin.

-- Reference: FAHAY (1983); HULLEY (1986); MOSER and WATSON (2001); MOSER and WATSON (2006b); MOSER et al. (1984);

-- Reference material: 1 specimen; 04/99: S12 (P2393827);

-- Distribution: A single specimen was caught at an epipelagic station southwest of Gran Canaria at a depth of 38 – 49 m at night in spring.

• Taaningichthys minimus Meristics:

Development:

Vertebrae:

39 – 41

Dorsal rays:

11 – 13

Anal rays:

11 – 14

Pelvic rays:

Pectoral rays:

15 – 17

Flexion: ~ 7.0 – 8.5 mm Transformation: ~ 21.0 mm

-- Morphology: The body is slender with a relative long and elongate gut. The head is small with round eyes and the lower jaw projects beyond the upper, which was clearly visible in the studied specimen.The larvae have a high count of vertebrae for myctophids and are therefore easily distinguishable. No photophores form at all during larval development, even Br2 would be lacking, but photophores had already developed in the studied larva since it was already in the transforming or even juvenile stage.

-- Pigmentation: Melanophores are present at the terminal gut. One to several pigments are embedded above the spinal column. Dots are located at the midline above the brain (cerebellum and medulla) and at the caudal peduncle region. At a size of 10 mm a

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

paired series of melanophores develop at the base of the dorsal and anal fin and becomes extended up to the caudal fin base in late postlarvae. Further dots are found on the margin of the hypural region.

-- Reference: FAHAY (1983); HULLEY (1986); MOSER and WATSON (2001); MOSER and WATSON (2006b); MOSER et al. (1984);

-- Reference material: 1 specimen; 11/00: S12 (P2393906);

-- Distribution: One transforming, almost juvenile T. minimus was caught west of Gran Canaria in a depth of 444 – 900 m in late daytime during autumn.

¬ LAMPRIFORMES The larvae hatch at a large size with a functional gut and large pigmented eyes. The head is laterally strongly compressed. Lampriformes possess highly protractible jaws with the upper jaw elongated, which develops precocious, but are variable in shape among species. The antiorly rays of the dorsal and pelvic fins are elongated and in some species various swellings and ornamentation develop. The dorsal fin base is elongated with a high number of fin rays. All larvae are strongly pigmented throughout the whole development. Melanophores are concentrated laterally and along the ventral as well as the dorsal margin of the body. Further pigmentation is found on the swellings of the dorsal and pectoral appendixes. Some larvae possess ink glands, which develop early and may be functional in young fish. The transformation is mostly gradual.

-- Reference: OLNEY (1984); OLNEY (2006a);

Lampridae Larvae are well developed when hatching with little or no yolk material. The jaws are protractible and the gut is already differentiated with an open lumen. Larvae are relatively slender in early stages but increase in depth with further development. The preanal length is 40 – 50 % of SL in most larvae. The anteriorly rays of the dorsal and pelvic fins are elongated.

-- Reference: CHARTER and MOSER (1996); OLNEY (1984); OLNEY (2006a);

• Lampris guttatus Meristics: Vertebrae (Myomeres):

46 – 50 (46 – 50)

Dorsal rays:

48 – 55

Anal rays:

33 – 42

Pelvic rays:

13 – 17

Pectoral rays:

21 – 24

Caudal rays:

30 – 32

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

-- Morphology: Lampris is slender in early stages, but increases fast in body depth with development. In later stages they are deep bodied with a symmetrical caudal fin. Larvae of about 10.6 mm SL have already the body shape of the adults.The caudal fin is almost complete at 8.6 mm SL. The base of the pectoral fin is initially vertically orientated, but moves into the adult horizontal position by about 10 mm SL. It was easy to identify the larvae, since they already had the adult body shape and developed fin rays to verify the identification with meristic counts.

-- Pigmentation In early larvae small dots are restricted to the region dorsally on the head and under the eyes. Internal pigmentation is concentrated above the gut. Pigmentation expands all over the body in form of stellate melanophores, except for the caudal peduncle.

-- Reference: OLNEY (1984); OLNEY (2006b);

-- Reference material: 1 specimen; 11/97: S13 (P38128);

-- Distribution: The only specimen was caught at an epipelagic station southeast of Fuerteventura at a depth of 90 – 107 m at late night, almost morning in autumn.

Regalecidae The body is extremely elongate and anal fin is absent. Spinules form on caudal and pelvic fin rays. Only one stout pelvic fin ray develops, with a second much reduced one, while other Lampriformes have three or more pelvic fin rays. The number of vertebrae is more than 60.

-- Reference: OLNEY (1984); OLNEY (2006c);

• Regalecus glesne Meristics:

Development:

Vertebrae (Myomeres):

143 – 153 (124 – 153)

Dorsal rays:

260 – 412

Anal rays:

Pelvic rays:

1 (second reduced)

Pectoral rays:

12 – 13

Caudal rays:

3–4

Flexion: 45.8 mm

-- Morphology: The newly hatched larvae are laterally compressed with a large head. The anteriorly part of body is deeper. The mouth is protractile and upward orientated as clearly visible in Figure 48.The large, round eyes are initially slightly stalked, a characteristic that had been already lost in the studied specimens. The dorsal fin moves forward until reaching the median level of the eye

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

and form out up to 300 rays, where the length of the rays is longer in the first third and is descending towards the tail. In the younger specimen under study (Figure 48) the dorsal fin did not reach even the level of the eye, whereas in the older one, the dorsal fin was already at the median level of the eye. The first three rays are connected with a membrane, while the fourth, very elongated ray extending the whole body length, is free to its base from that membrane. The pectoral fin has three rays, which are free from a membrane. The myomeres are increasing in number during postflexion development. The caudal fin has four rays, the anal fin is absent and the pelvic rays are initially four but decrease in number with development till only one ray remains.

-- Pigmentation: Scattered pigments are along the elongate fourth fin ray. Other pigmentation is present below the origin of the elongate dorsal rays and posteriorly along the trunk. Further pigmentation occurs along the ventral midline, at the end of the caudal peduncle, on the head and behind the thoracic belt. Along the trunk nine small melanophore groups develop dorsally and eleven along the ventrally profile, where each group grow up to the lateral line till they connect each other. Along the dorsal and the ventral profile of the caudal peduncle little spots and scattered pigmentation, respectively form out and get more intensive with further development.

-- Reference: BERTOLINI et al. (1956); OLNEY (1984); OLNEY (2006c);

-- Reference material: 2 specimens; 04/99: S13 (P38129); S14 (P38130);

-- Distribution: The specimens were caught only in spring south and southwest of Gran Canaria, whereas a juvenile specimen was caught at a mesopelagic station at a trawling depth of 480 – 694 m during the day and the larva was found at a neritic station at a depth of 33 – 53 m also during the night.

¬ GADIFORMES More than in a half of the Gadiformes fish indications of a caudal fin is lacking. If present, then the elements of the caudal fin are highly divers. Most fish possess five pectoral radials and the fin itself is positioned high on the body. The pelvic fins are located far anteriorly of the body (thoracic) and are usually elongate in larvae. In common are very high vertebral counts, and high counts of dorsal and anal fins with long fin bases. There can be 1 – 3 dorsal and 1 – 2 anal fins. The posteriorly dorsal and anal fin rays are separated from caudal fin rays, except in Muraenilepis and Macrourines. The mouth position is very variable within the order and barbels are present in many genera, which form on the symphysis of lower jaw.The larvae have already a well developed pigmentation pattern when hatching. The gut is short and coiled and the preanal length varies form one third to about the half of SL. Larvae do not have spines and the transformation is gradual. The juvenile stage is pelagic.

-- Reference: COHEN (1984); FAHAY (1983); FAHAY and MARKLE (1984);

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

Macrouridae No specialized larval developments exist and adult organ morphologies develop directly at a small size. As only temporary larval developments the stalked pectoral fins and an undeveloped snout form out. Knowledge is limited to alevins, where adult identification characters can be used. The macrourids are extremely elongate and the head length can make up to about 10 % of TL mostly having a protrusible mouth. The tail is moderate in length to very elongate and lack a differentiated caudal fin. They have a filiform caudal end instead. The dorsal spine serration form out late in larval development. Further, within the family the number of branchiostegal rays is 6 – 7. The gillrakers of the first arch is in Macrouroidinae lath-like shaped and 25 – 30 in number, while in Macrourinae the range varies from only 5 – 28 and having a tubercular shape. But more easily distinguishable characters are the one or two based dorsal fins and their relative length of dorsal to anal fin rays. Species of macrouroidines have a one based dorsal fin, while the macrourines have a two based dorsal fin, with the first one short based and having the first dorsal spine shortened and the second elongated. In both subfamilies the rays of the second dorsal and anal fins are confluent, but the anal fin rays are longer in relation to the dorsal fin rays in Macrourinae (Figure 49) and the other way around in the Macrouroidinae. This character is very easy to see, therefore it is possible to differentiate late stage larvae at least to subfamily level. But other important features, which would be crucial for larval identification to a low taxonomic level are not described for Macrouridae yet.

Macrourinae The anal fin rays are longer than the dorsal fin rays (Figure 49).The second spinous ray of the dorsal fin can be smoothed or serrated. Light organ can be present or absent, bulbous or tubular. Mouth position is terminal, sub-terminal or inferior. Chin barbel is present or absent.

-- Reference: FAHAY and MARKLE (1984); MERRET (2006); OLIVAR and FORTUÑO (1991);

-- Reference material: 5 specimens; 05/99: S2 (P375467); 01/00: S5 (P375466); 03/02: S16 (P375464-465); S22 (P375463);

-- Distribution The alevins were caught from winter to late spring. One specimen was found at a mesopelagic station at 950 – 1260 m depth during the day in winter. During very late winter alevins were caught southeast of Fuerteventura in a depth range of 185 – 211 m during the day and 215 – 261 m during the night. In late spring one alevin was caught at a mesopelagic station east of Fuerteventura at 495 – 561 m depth during the day.

Melanonidae The family consists only of one genus with two species. The body is slender and decreases towards the posterior part of the body ending in a small and narrow caudal fin.The head is blunt with a large mouth. A chin barbel is absent.The dorsal and anal fins are long based, almost connected to the procurrent caudal fin rays, which extend far anteriorly on the caudal peduncle. The pectoral fins are short based and located at the midheight of the body depth at the level of the dorsal fin origin. The pelvic fins are positioned at the level just posteriorly to the pectoral fins having 7 fin rays. On the smallest caught specimen all fins were formed possessing the adult body shape with 15 mm.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Reference: FAHAY and MARKLE (1984); LINDQUIST, SHAW and FAROOQI (2006);

• Melanonus zugmayeri Meristics: Vertebrae:

58 – 62

Dorsal rays:

67 – 80

Anal rays:

52 – 61

Pelvic rays:

5–7

Pectoral rays:

14 – 16

Caudal rays:

55 – 60

-- Morphology: Between the first and the second dorsal fin a small interspace is present. The anal fin originates far posteriorly on the body and anteriorly fin rays are elongated. The pectoral and the pelvic fins are small, with their bases at the same level of the body.

-- Pigmentation: Distinctive is the dark pigmentation on the body and on the peritoneum, which is prominent for that species and very distinct in the studied specimen (Figure 49).

-- Reference: COHEN (1986); FAHAY and MARKLE (1984); LINDQUIST, SHAW and FAROOQI (2006);

-- Reference material: 1 specimen; 05/99: S5 (P375468);

-- Distribution: This species is a new record around the Canary Islands, but were also found at the Azores and Madeira before. The single specimen was caught north of La Gomera and Tenerife at a depth of 448 – 526 m during the night in late spring.

¬ STEPHANOBERYCIFORMES Stephanoberyciformes have a high number of pelvic and caudal fin rays. Spines are present in dorsal, anal and pelvic fins. Also procurrent spinous caudal fin rays are developed in that order.

-- Reference: FRIAS-TORRES (2006a); RICHARDS (2006h);

Melamphaidae The early larvae are slender, but deepen with further development. The dorsal fin is more or less positioned oppositely to the anal fin, but the origin of the dorsal fin is more anteriorly. The dorsal fin forms out 1 – 3 weak spines with 9 – 18 rays,

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

while the anal fin has one single spine with 7 – 10 rays. The pelvic fins form already in preflexion stage and have one weak spine with 6 – 9 rays, which are long, fragile and pigmented.The origin of the pelvic fin is close anteriorly or posteriorly to the pectoral fin level, but migrates more posteriorly with development. The number of pectoral fin rays varies from 14 – 17. The principal caudal ray number is 10 + 9 and the procurrent spines have a number of 3 – 4. The number of vertebrae ranges from 24 – 31. The pattern of pigmentation varies strongly among species. Prominent are the spots at the posterior end of the dorsal and anal fin, which spread out with development. If a more intense pigmentation appears then it does so on the cranium, peritoneum and on the caudal peduncle.

-- Reference: FAHAY (1983); FRIAS-TORRES (2006a);

• Melamphaes sp. The myomere counts ranges from 25 – 31 and the dorsal fin ray counts from 10 – 14. At the dorsal margin of the head and the eyes a bulge is formed out. In very early larvae two spots are present at the anlagen of the dorsal and anal fin, which grow with further development to a line along the dorsal and ventral margin, some larvae form a band between these pigmentation areas at the level of dorsal and anal fin in late stages. Additionally pigments appear on the cranium, peritoneum and at the posterior region of the caudal peduncle.

-- Reference: FAHAY (1983); KEENE and TIGHE (1984);

-- Reference material: 2 specimens; 04/99: S14 (P412418); 05/99: S6 (P412419);

-- Distribution: Larvae were only caught in spring southwest of Gran Canaria at a depth of 480 – 694 m in day and epipelagic north of La Gomera and Tenerife at 48 – 66 m depth during the night.

¬ BERYCIFORMES The high number of pelvic and caudal fin rays is prominent for this order. Dorsal, anal and pelvic fin spines develop as well as spinous procurrent caudal fin rays. The body is slender in early larvae, but deepens with development. The pelvic fins with its long pigmented rays form rapidly anterior on body, but move more posteriorly with development. In many families the second or third ray of the dorsal fin is elongate. The development is gradual and direct.

-- Reference: KEENE and TIGHE (1984); RICHARDS, LYCZKOWSKI-SHULTZ and KONIECZENA (2006a); WATSON (1996b);

TRACHICHTHYOIDEI > Diretmidae The early larvae are slender, but getting deeper with development. A short spine develops on the frontal bone above each eye and a longer parietal spine on each side, which is directed backwards. This spination is typical for that family (Figure 51). Further on each side a preopercular spine develops, which is depending on the species postero- or anteroventrally orientated. The ridge of the opercle is not serrated.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Reference: KEENE and TIGHE (1984); RICHARDS, LYCZKOWSKI-SHULTZ and KONIECZENA (2006b);

• Diretmus argenteus Meristics: Vertebrae:

28 – 30

Dorsal rays:

25 – 29

Anal rays:

20 – 24

Pelvic spine + rays:

I+6

Pectoral rays:

16 – 20

Principal caudal rays:

8–9

The body is relatively slender in early stages smaller than 4 – 5 mm, but deepens with development. The preopercular spine is directed posteroventrally (Figure 51).

-- Pigmentation: Scattered pigment pattern all over the body, but some specimen are described with vertical bands on the body profile.

-- Reference: RICHARDS, LYCZKOWSKI-SHULTZ and KONIECZENA (2006a); RICHARDS, LYCZKOWSKI-SHULTZ and KONIECZENA (2006b);

-- Reference material: 2 specimens; 01/00: S3 (P40339); 03/02: S3 (P40338);

-- Distribution: Larvae were only caught at mesopelagic depths at day time in winter southeast of Fuerteventura at 440 – 665 m depth and north of Gran Canaria at a depth of 356 – 402 m.

Trachichthyidae The body is very deep and laterally strongly compressed. The heavy strong head spination forms at the margin of the preopercle and opercle. Further spines appear at the interopercle and subopercle and posttemporal. Spines are present on scales and lateral scales when developing and on bases of the dorsal fin rays. The pelvic fins are well developed. The number of vertebrae ranges from 26 – 29.

-- Reference: KEENE and TIGHE (1984); RICHARDS, LYCZKOWSKI-SHULTZ and KONIECZENA (2006c);

• Hoplostethus sp. The body and the head are very deep and laterally strongly compressed (Figure 52).The head is larger than or about the size of 1/3 of SL in adults. The posttemporal and preopercle spines might be still present in adults.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

-- Reference: KEENE and TIGHE (1984); MAUL (1986); RICHARDS, LYCZKOWSKI-SHULTZ and KONIECZENA (2006a); RICHARDS, LYCZKOWSKI-SHULTZ and KONIECZENA (2006c);

-- Reference material: 9 specimens; 11/97: S16 (P40349); 01/00: S5 (P40343); S7 (P40346-347); S9 (P40344); 03/02: S3 (P40342); S6 (P40341; P40348); S12 (P40345);

-- Remark: All specimens are probably Hoplostethus atlanticus COLLETT, 1889. The suggestion is based on the observation, that larval Hoplostethus develop little stings collaterally along the future dorsal fin spines (Figure 53). Consequently the fin formula could be determined.

• Hoplostethus mediterraneus Meristics: Vertebrae:

Dorsal spines + rays:

IV – VII + 12 – 14

Anal spines + rays:

III + 9 – 11

Pelvic spine + rays:

I+6

Pectoral rays:

14 – 16

Caudal rays:

-- Morphology: The body and the head are very deep and laterally strongly compressed. The head is larger than 1/3 of SL in adults. The posttemporal and preopercle spines are still present in adults. Although the specimen had already developed fin spines, the little stings along the fin spines were still present and clearly visible (Figure 53).

-- Reference: KEENE and TIGHE (1984); MAUL (1986); RICHARDS, LYCZKOWSKI-SHULTZ and KONIECZENA (2006a); RICHARDS, LYCZKOWSKI-SHULTZ and KONIECZENA (2006c);

-- Reference material: 1 specimen; 04/99: S10 (P40340);

-- Distribution: The only specimen was caught in spring at a mesopelagic station east of the channel between Fuerteventura and Lanzarote at a depth of 258 – 600 m in the day.

¬ GASTEROSTEIFORMES This order consists of very specialized fishes. The shape of the body is highly variable and specialized. But they have in common that the branchiostegal rays are reduced to a number of only 1 – 5. The mouth is small and has often a tubular shape. The postcleithrum is consistent of only one bone or even absent. The pelvic girdle is never directly attached to the

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

cleithrum.The pelvic fins, if present are very small and located at about the midpoint of the body.The dorsal and anal fins are positioned far back of body in an opposing position. The body is covered by spinules or rows of lateral bony scutes. Males may possess a specialized putch for receiving and incubating the eggs. Species with broadcast spawn or nests have larvae, which troughrun the entire development of pelagic larvae, but species which are breeding, set already young juveniles, having adult characters, free. Meristic characters and pigmentation are quite variable in this order.

-- Reference: DITTY et al. (2006); FRITZSCHE (1984);

SYNGNATHOIDEI > Syngnathidae This family consist of the following two subfamilies, first the Syngnathinae and second the Hippocampinae. Both are elongated fish, but with different posture of the body. But prominent for this family is the presence of a cutaneous skeleton, where bony scutes unite and form rings along the entire body. These rings may be very helpful in identification. After DAWSON et al. (1984) these rings are classified into the trunk rings, starting at the pectoral fin base and end with the ring at the anus, and the tail rings, which start with the first ring after the anus and the last ring on tail, but excluding the one bearing the caudal fin. Further, subdorsal rings are trunk and tail rings, which are bearing the dorsal fin. The mouth is strongly tubular and toothless. Pelvic fins are absent. The anal fin is small with few rays and no spines are developed in the dorsal fin. The dorsal fin is located above the anus, around the midpoint of body.The males incubate the fertilized eggs in pockets or skin folds.The young Syngnathidae form dark bars along their body.

-- Reference: BERTOLINI et al. (1956); DAWSON (1986); DITTY et al. (2006); FRIAS-TORRES (2006b); FRITZSCHE (1984);

• Entelurus aequoreus Meristics: Dorsal rays:

37 – 47

Anal rays:

4–9

Rings:

28 – 31 + 60 – 69

Subdorsal rings:

11 – 7 + 2 – 4

-- Morphology: The body is very elongate and the anus is placed at the anteriorly level of the dorsal fin. When larvae are newly hatched a medial finfold and pectoral fins are developed, but both disappears with late development. The subdorsal rings are clearly visible and easy to count (Figure 54).

-- Pigmentation: Stellate melanophores are present along the dorsal and little spots along the ventral margin of the body. Further stellate melanophores develop also along the dorsal medial finfold in young larvae and later along the gut.

-- Reference: DAWSON (1986); MUNK and NIELSEN (2005);

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

-- Reference material: 1 specimen; 11/97: S4 (P39922);

-- Distribution: This species is a new record from the area of the Canary Islands, but was already recorded from the Azores before. The only specimen was caught at a neritic station southeast of Fuerteventura at a depth of 22 – 35 m during the day in autumn.

¬ SCORPAENIFORMES All fish have the suborbital stay in common. Larvae possess large pectoral fins, which are pigmented and have an extensive head spination, which varies among the Scorpaeniformes. Many species have an extended pelagic juvenile stage.

-- Reference: LEIS and RENNIS (1983); RICHARDS (2006i); WASHINGTON et al. (1984);

SCORPAENOIDEI > Scorpaenidae The myomere count is 24 – 27. Preflexion larvae are elongate to moderately deep bodied but becoming more robust in the postflexion stage. The gut is initially straight, but coils fast, what results in a preanal length of 1/3 to 2/3 of BL. The gas bladder is small. Pigmentation is in general very light, but prominent on the pectoral fins (Figure 55). Further pigments are located on the head, along the ventral and lateral midlines on the caudal peduncle and on the gut.The shape of the head is very variable and the eyes are large and round. The mouth is moderate to large and full with teeth, which form after flexion at a size of 5.5 – 5.8 mm. The ridges and spination on head are well developed, with the largest spines found at the preoperculum. The large pectoral fins form early and are fan shaped with rays extending to the anus and even reaching the caudal fin in some species. Between the anus and the origin of the anal fin an interspace is present. The posterior anal fin spines form first as soft rays and transform to spines before settlement. There are no scales before the settlement developed.

-- Reference: HARDY (2006a); LEIS and RENNIS (1983);

Sebastinae • Helicolenus dactylopterus Meristics:

Development:

Vertebrae (Myomeres):

23 – 25 (9 – 12 + 16 – 18)

Hatching: 1.9 – 2.6

Dorsal spines + rays:

XI – XIII + 10 – 14

Anal spines + rays:

III + 3 – 5

Flexion: 4.5 – 6.6 mm

Pectoral rays:

17 – 20

Transformation: > 19 mm

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Morphology: The larvae hatch at a size of 1.9 – 2.6 mm and are moderately deep bodied, but deepen more with development. Prominent is the spongy tissue, which develops anteriorly in the dorsal finfold during preflexion stage and persists almost throughout the whole larval period. Spines are lacking on the second infraorbital bone and on the cleithrum.

-- Pigmentation: Pigments are present on the dorsolateral surface of the gut, above the brain, on the lower jaw, along the ventral margin of the caudal peduncle and on the pectoral fin.

-- Reference: HARDY (2006a); WASHINGTON, ESCHMEYER and HOWE (1984); WASHINGTON et al. (1984);

-- Reference material: 2 specimens; 04/99: S5 (P791441); S14 (P791442);

-- Distribution: In spring a juvenile was caught southeast of Fuerteventura at a mesopelagic station at a depth of 382 – 511 m in daytime and a larval specimen was caught south of Gran Canaria in a depth range from 480 – 694 m during the day.

Scorpaeninae • Scorpaena sp. The suborbital stay is present and may be spinated in some species, but not so in the studied specimens (Figure 55). Also, preopercle spines are present, the first being the longest. The second and third are developed in the entire genus, but the fourth and fifth spine can be absent. Spines are usually present at the posttemporal, preoperculum and at supra- and cleithral too. Further spines are present on the head.The dorsal fin consists of 12 spines and 7 – 10 rays.The anal fin has 3 spines and 5 rays and the pectoral fin ray number ranges from 16 – 21, where some rays may be branched.

-- Reference: HUREAU and LITVINENKO (1986);

-- Reference material: 5 specimens; 11/97: S10 (P791444-445); S16 (P791447); 04/99: S22 (P791443); 11/00: S1 (P791446);

-- Distribution: In autumn some specimens were caught at neritic stations during the night, so northeast of Lanzarote at a depth of 20 – 64 m and west of Gran Canaria at 42 – 66 m depth. Also at one epipelagic station north of Gran Canaria at a depth of 55 – 90 m during the night some larvae were found. Only one specimen caught at a neritic station west of La Gomera in a depth range from 25 – 70 m also during night, but in spring.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

¬ PERCIFORMES Most species have spines in the dorsal and the anal fins and lack the adipose fin. The pelvic fins have one spine and five or fewer soft rays. The number of the branchiostegal rays is seven or lower. Only four gill arches are present.

-- Reference: NELSON (1994);

PERCOIDEI > Moronidae The lateral line, which is not developed in larvae, is very long and can reach the posterior end of the caudal fin and an additional row scales above and below the lateral line on the caudal peduncle develops. Moronids hatch at 1.7 – 3.7 mm and flexion takes place at 7 – 9 mm. The dorsal and anal fin rays are completed shortly after flexion and first scales appear at a size of 16 – 25 mm.

-- Reference: JOHNSON (1984);

• Dicentrarchus sp. The preoperculum is serrated, where the spines of the ventral edge are orientated forward. Two unconscious spines on the opercle are still present in adults. Two dorsal fins develop and the caudal fin is moderately forked. Two species occur around the Canary Islands, Dicentrarchus labrax (LINNAEUS, 1758) and D. punctatus (BLOCH, 1792). But since larvae information is only available for D. labrax and the meristic counts are the same in both species, it is impossible to definitely distinguish one from the other (Figure 56).

-- Reference: BERTOLINI et al. (1956); TORTONESE (1986b);

-- Reference material: 5 specimens; 11/97: S4 (P43721); S5 (P43718); S10 (P43719); 03/02: S4 (P43720); S19 (P43722);

-- Distribution: Larvae occurred in autumn and in winter. In autumn specimens were caught mainly at neritic stations, so southeast of Fuerteventura at 22 – 35 m depth during day and northeast of Lanzarote at a depth of 20 – 64 during the night. Another larva was found at an epipelagic station south of Fuerteventura at about 40 m depth during the early morning. In winter larvae were caught at epipelagic stations southeast of Fuerteventura in a depth range of 22 – 139 m during the night.

Percichthyidae According to NELSON (2006) Howella is only provisionally placed under the Percichthyidae, whereas RICHARDS and LARA (2006) treat Howella as an own family. Although I follow NELSON (2006) in the taxonomic order, the following description is only valid for species of Howella. The body is moderately elongate and laterally compressed, but deepens with larval growth. The head is large as well as the mouth and the eyes are. The gut is initially straight, but coils during preflexion stage to a

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

preanal length of about 60 % of BL. The flexion takes place at around 3.5 mm and the dorsal and anal fin rays are completed at a size of 4.5 mm. Sub-, pre-, and opercle spination is developed. All fins have a low count and between the dorsal fins a large interspace is present. The pectoral fins are elongated and may reach to the origin of the anal fin. The caudal peduncle is long and the caudal fin is forked.

-- Reference: JOHNSON (1984b); NELSON (2006); RICHARDS and LARA (2006);

• Howella brodiei Meristics: Vertebrae:

24 – 26

Dorsal spines + rays:

VII – VIII + I + 8 – 10

Anal spines + rays:

III + 7 – 8

Pelvic spine + rays:

I+5

Pectoral rays:

14 – 16

-- Morphology: Four opercular spines are present. Sub- and preopercle end in each one large spine. The second dorsal fin and the anal fin are positioned in the posterior part of the body in an opposite position. Originally the pectoral fin is elongate and reaches beyond the anal fin origin, but all elongate pectoral fin rays broke off (Figure 57).

-- Pigmentation: The little stellate melanophores distribute all over the body, but concentrate at the origin of the caudal peduncle. This is a prominent character for that genus and very distinct (Figure 57). Together with the meristic counts, species identification is possible. Further, the dorsal surface of the gut is strongly pigmented. Spots form at the tip of the snout.

-- Reference: RICHARDS and LARA (2006);

-- Reference material: 4 specimens; 11/97: S4 (P43725-726); S10 (P43723-724);

-- Distribution: Specimens were only present at neritic stations in autumn, so southeast of Fuerteventura at a depth of 22 – 35 m at daytime and northeast of Lanzarote at 20 – 64 m depth during the night.

Serranidae The family consists of four distinguishable morphological subfamilies. But all larvae hatch at 1.2 – 2.3 mm with a large yolk sac, unpigmented eyes and underdeveloped mouth. The gut is straight and reaches to or beyond the midbody level, but coils and shortens during preflexion stage. The shape of the body and head are variable from slender to deep and from broad to laterally compressed.The anal, pectoral and pelvic fins form early. If there are any elongate spines, then mostly the third dorsal or the pelvic spines are modified, but more spines can be elongate and modified as well. The pectoral fins may be enlarged and pigmented. The pigmentation is very variable, but in most larvae light to moderate. But melanophores are present and

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

accumulated in the area of the head, at the gas bladder, at the gut, dorsally on the trunk, dorsally and ventrally along the caudal peduncle and on the spinous dorsal, pectoral and pelvic fins. In some an armature on the head develops.

-- Reference: BERTOLINI et al. (1956); KENDALL Jr. (1984); RICHARDS, BALDWIN and RÖPKE (2006); WATSON (1996c);

Serraninae • Serranus sp. The body is deep. The pelvic fins and the dorsal spines form early, where latter are elongate. Pigmentation is found at the angular, cleithral symphysis, on the anus, on the trunk and the anal fin, along the ventral margin of the caudal peduncle and below the dorsal fin base. Further pigmentation is located at the membranes of the first dorsal, pelvic and anal fins.

-- Reference: JOHNSON (1984b); RICHARDS, BALDWIN and RÖPKE (2006);

-- Reference material: 1 specimen; 05/99: S1 (P43731);

-- Distribution: In late spring one specimen was caught at an epipelagic station far east of Fuerteventura at a depth of 38 – 40 m in the morning.

Anthiniinae • Anthias anthias Meristics:

Development:

Dorsal spines + rays:

X + 15

Anal spines + rays:

III + 7

Pelvic spine + rays:

I+5

Pectoral rays:

Flexion: 5.5 mm

-- Morphology: The body is moderately deep. At the operculum three series of short and stout spines are present, where the two more external series have each one spine strongly developed. These two spines are more or less overlapping each other and are directed towards the anus, which is prominent for that species and was clearly visible in the studied specimens (Figure 58). All spines at the operculum get stronger developed with larval growth, but are reduced towards the juvenile stage. Further, a supraorbital crest develops in later larval stages. All fins start to form at a size of approximately 5 mm. The third spine of the dorsal fin is elongated. The pelvic and the pectoral fins are well developed, where the latter is elongated.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Pigmentation: Three black dots are located ventrally posteriorly to the anus, where the first of these is the largest, and one further spot is present at the caudal fin base. The tips of dorsal and pectoral fin rays are pigmented as well as the anteriorly membrane of the first dorsal fin. Further pigmentation is situated at the peritoneum and above the forebrain.

-- Reference: BERTOLINI et al. (1956); OLIVAR and FORTUÑO (1991); TORTONESE (1986a);

-- Reference material: 4 specimens; 11/97: S10 (P43729); S13 (P43728); S18 (P43727); 04/99: S13 (P43730);

-- Distribution: In autumn specimens were caught at neritic stations in night, so northeast of Lanzarote at a depth of 20 – 64 m depth and east of Gran Canaria at 116 – 146 m. One more specimen was caught at an epipelagic station southeast of Fuerteventura in a depth range of 90 – 107 m in morning time. In spring one larva was found at a neritic station at a depth of 33 – 53 m during the night.

Callanthiidae Within this family the number of dorsal fin spines is 11 and the dorsal fin rays vary from 9 – 11. The lateral line runs from the level of the dorsal fin ignition to its end or even to the caudal peduncle. The number of vertebrae is 24.

-- Reference: NELSON (2006);

• Callanthias ruber Meristics: Dorsal spines + rays:

XI + 10 – 11

Anal spines + rays:

III + 9 – 10

Pelvic spine + rays:

I+5

Pectoral rays:

-- Morphology: The body depth is high and increases more with development. The preanal length is about the half of the BL, but shortens with further larval growth. Above the large and round eyes a serrated crest develops which smoothens later on and was almost not anymore visible in the studied specimens. The operculum carries two rows of spination and one scapular spine is present at the pectoral girdle.

-- Pigmentation: No pigmentation is present except on the peritoneum and little spots above the brain. The peritoneal spots were not visible in the studied specimens, but the spots above the brain were very distinct (Figure 59).

-- Reference: BERTOLINI et al. (1956); TORTONESE (1986a);

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

-- Reference material: 39 specimens; 04/99: S5 (P43760); 05/99: S2 (P43759); 03/02: S3 (P43765-767); S6 (P43768-770); S11 (P43754-758); S12 (P43737); S13 (P43738-745); S15 (P43751-753); S16 (P43746-750); S17 (P43732-736); S20 (P43761-764);

-- Distribution: Larvae were caught in late winter during the day southeast of Fuerteventura mainly at mesopelagic stations in a depth range of 311 – 1035 m. Some specimens occurred also at epipelagic stations at a depth of 30 – 211 m in day. In the spring larvae were found also southeast of Fuerteventura at a mesopelagic station at 382 – 511 m depth during the day and in late spring larvae were caught at a mesopelagic station east of Fuerteventura at a depth of 495 – 561 m in daytime.

Apogonidae Larvae hatch at 2.5 – 3 mm well developed, with pigmented eyes, functional large mouth and a small yolk sac. Flexion takes place at 3 – 4 mm. Most larvae are moderately elongate in early stages, but becoming deeper with development.The head is large. The preanal length reaches to about midbody level and the larvae have throughout all stages a distinctive gas bladder. Small spines form along the posterior margin of the preoperculum shortly before the flexion. Pigmentation may be present above the brain or on the dorsal profile of the head, on the branchiostegal membranes, the gas bladder, the gut, along the dorsal margin of the body, along the lateral midline and ventrally of the caudal peduncle.

-- Reference: SANDKNOP and WATSON (1996); JOHNSON (1984b);

• Apogon imberbis Meristics:

Development:

Myomeres:

Dorsal spines + rays:

VI + I + 9 – 10

Anal spines + rays:

II + 8 – 9

Pelvic spine + rays:

I+5

Pectoral rays:

10 – 12

Hatching: 2.5 mm

-- Morphology: The preanal length is to about midbody length. The body is moderately deep. The bluish colored eyes are not very large and the mouth is not open, when hatching. At the operculum spines form and at the preoperculum three short thin spines develop in early larvae, but change to a serrated margin with further larval growth. The supraorbital crest is distinguishable and develops to an eye surrounding protruding crest. All these spination and serrated margins were not visible anymore and the larval pigmentation was already developed in the late stage larva studied.

-- Pigmentation: Initially no pigmentation is present, but melanophores develop in the occipital and cephalic region and at the brain in postflexion stage.

-- Reference: BERTOLINI et al. (1956); TORTONESE (1986a);

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Reference material: 1 specimen; 11/97: S5 (P44296);

-- Distribution: Only one specimen was found at an epipelagic station south of Fuerteventura at a depth of about 40 m at the break of dawn in autumn.

Carangidae The family consists of four subfamilies. But larvae hatches at a length of 1.3 – 4.3 mm having a large yolk sac but underdeveloped mouth, no pectoral fin buds, and no eye pigmentation. The gut is initially straight and reaches up to 50 – 70 % of BL, but coils with development without changing the preanal length. Most early larvae are slender with a large head and increase their body depth with growth. On the head ridges and spines form out, where the two rows of spines at the preoperculum are prominent, with the larger spines developing along the posteriorly margin of the preoperculum. The largest spine hereby is situated at the angle of the preopercle, which is also the first to form in early larvae. The number of the spines is variable and increase during early development and gets reduced in early juvenile stage. Just shortly towards the juvenile stage, the remaining preopercular spines get completely overgrown by bones and tissue. Other spination occurs at the supracleithrum and the posttemporal bones until flexion stage, but no spines form at interoperculum and suboperculum.The often serrated supraoccipital ridge forms early in preflexion larvae and remains to late transformation stage, where it is also overgrown by bones and tissue. The frontal supraocular ridge may develop spines. The fin formation starts with the caudal fin rays in late preflexion stage, followed by the pectoral fins and soft rays of dorsal and anal fin. The spinous dorsal fin forms next, while the pelvic fins develop last. At the anteriorly margin of the first dorsal fin an antrorse spine is developed, but covered by skin. Pigmentation is very variable and ranges from light to almost complete, but melanophores concentrate at dorsum, the gas bladder, the gut, along the lateral midline, along the ventrally margin of the tail and laterally on the caudal fin base. The pigmentation gets more intense during postflexion stage, with the countershading initiate and vertical bars begin to form.The development is gradually and direct. No sudden metamorphose happens.The number of myomeres ranges at 23 – 26 or 27. The first two spines of the anal fin are separated from the rest and a distinctive gap in most species is present.

-- Reference: LAROCHE, SMITH-VANIZ and RICHARDSON (1984); WATSON et al. (1996);

• Campogramma glaycos Meristics: Dorsal spines + rays:

VI – VII + I + 26 – 28

Anal spines + rays:

II + I + 23 – 25

-- Morphology: The mouth is large, extending the level of the eye. The dorsal fin base is shorter than the anal fin base. The length of pectoral fin rays is extending the pelvic fins rays.The separated first two spines were clearly visible in the studied specimens (Figure 61).

-- Reference: SMITH-VANIZ (1986);

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

-- Reference material: 6 specimens; 11/97: S16 (P461539-44); S18 (P461545);

-- Distribution: All larvae were exclusively found at neritic stations around Gran Canaria in night during autumn. The depth ranges from 42 – 66 m western and from 116 – 146 m eastern of the island.

• Trachurus sp. The body is slender and its depth does not exceed 35 % of the SL. No dorsal fin spines or other rays are elongated. No finlets develop at all. The supraoccipital ridge is present in flexion and postflexion larvae and at the supraocular ridge, tiny spines develop. Both were not that clearly visible, but present in the studied specimens (Figure 62). At the posterior angle of the preoperculum a single spine forms and the posttemporal and supracleithral spines are developed very weak. Stronger pigmentation form dorsolaterally and along the lateral midline. Whereas light scattered pigmentation develops ventrolaterally on the myomeres and not in the hypaxial myosepta, like in other genera.

-- Reference: LAROCHE, SMITH-VANIZ and RICHARDSON (1984); LAROCHE et al. (2006);

-- Reference material: 12 specimens; 04/99: S6 (P461547-550); S16 (P461546); 05/99: S1 (P461551); 03/02: S11 (P461556-557); S13 (P461552554); S19 (P461555);

-- Distribution: All specimens were caught at cruises in late winter to late spring. Larvae were mainly found at epipelagic stations southeast of Fuerteventura within a depth range of 23 – 140 m at day as well as night and also at an epipelagic station far east off Fuerteventura at 38 – 40 m depth. Further larvae were found at a neritic station southwest of Gran Canaria at a depth of 18 – 61 m during the night and at a mesopelagic station at 632 – 1035 m depth during daytime.

Sparidae Hatching occurs at a size of about 2 mm and larvae have a large yolk sac, with a posteriorly oil globule, but unpigmented eyes, an undeveloped mouth and pectoral fins. Usually the myomere count is 24. Larvae are moderately compressed and the coiled gut reaches to 40 – 60% of BL. Spination develops on the head and at the pectoral girdle and is very variable within that family. The count of the second dorsal and anal fin is of about the same number. The pattern of pigmentation is very variable during yolk sac stage and getting lightly pigmented during preflexion stage, where melanophores concentrate at the ventrum, dorsally on the gas bladder, on the gut and in some species also dorsally on the head. During flexion and postflexion some larvae remain lightly pigmented, while others develop pigmentation along the lateral midline.

-- Reference: FAHAY (1983); POWELL and GREENE (2006); WATSON and SANDKNOP (1996c);

• Dentex sp. Larvae are slender, but adults are rather deep.The preanal length is to about midbody length, but decreases with growth.The count of the spinous and the soft rays of the dorsal fin have equal numbers.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Reference: BAUCHOT and HUREAU (1986); BERTOLINI et al. (1956);

-- Reference material: 4 specimens; 11/97: S4 (P50679-82);

-- Distribution: Larvae were only caught at a neritic station southeast of Fuerteventura at a depth of 22 – 35 m in daytime during autumn.

• Pagrus pagrus Meristics:

Development:

Vertebrae:

24 (10 + 14)

Dorsal spines + rays:

XI – XIII + 9 – 11

Anal spines + rays:

III + 7 – 9

Pelvic spines + rays:

I+5

Pectoral rays:

15 – 16

Principal caudal rays:

9+8

Flexion: 4.1 mm Transformation: 13 mm

-- Morphology: In early larvae the body is rather slender, but deepens with further development. The preanal length is a little less than midbody length and decreases during later stages. Only the pectoral fins are developed in early larvae. At the operculum a protruding crest is present, from which the spines form two rows, where the anterior series develop 7 and the posterior one develop 8 spines, where the 5th spine at the angle is the longest. This number of spines at the operculum are an important character of identification. No occipital crest is visible in early stages, but develops in larvae more then 5 – 6 mm. A supraocular crest is seen above the eye with 4 short stout spines. A supraoccipital crest develops, which is typically for the Pagrus up to a length of 15 mm. Further interopercular, suprachleithral, pterotic, tabular and postemporal spines develop. At 13 mm transformation takes place and crests start to disappear, but are still visible with larger sizes and were also encountered the studied specimen. At a size of 30 mm all spines have reduced totally. The anus finally migrates to the midpoint of body. At a size of 20 mm the settlement already takes place.

-- Pigmentation: In early larvae pigmentation occurs at the upper and lower jaw. Posteriorly to the anus along the ventral profile 6 – 7 or 3 – 4 spots form. Further dots occur on the ventral profile from anterior to the cleithral symphysis, at the base of the pelvic fins and anterior to the anus. The gut is pigmented on the postdorsal surface. In flexion stage the dots on the ventral midline posteriorly to the anus can be reduced to two dots and do not exceed the number of three in postflexion stage.

-- Reference: BAUCHOT and HUREAU (1986); BERTOLINI et al. (1956); POWELL and GREENE (2006);

-- Reference material: 1 specimen; 03/02: S1 (P50683);

-- Distribution: The only specimen was caught at the neritic station at a depth of 24 – 60 m at night during late winter.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

LABROIDEI > Pomacentridae Larvae hatch with a small yolk sac and pigmented eyes and a functional mouth. Larvae are slender when hatching, but increase their body height to more than 40 % of BL. The head is laterally compressed and a weak head spination develops in most. The mouth is small without teeth. Small teeth develop at a size 6 – 7 mm SL in both jaws and the mouth gets larger and may also get protrusible. The gut is short and coiled with a preanal length of being usually below 40 % of the BL, but increase in length with development up to about 50 % of BL.The sequence of fin formation is variable, but all may be already present at a size of 3 mm. Members of that family develop 8 – 17 dorsal spines and 10 – 18 dorsal soft rays, while the anal fin consists of 2 spines and of 10 – 18 soft rays.The count of myomeres is 25 – 27, but usual is 26.The pigmentation is restricted to the ventral margin of the tail, on the gut, on the head and on the inconspicuous gas bladder. One or more series of small opercular spines develop. Settlement may not occur until a size of 18 mm.

-- Reference: PARIS-LIMOUZY et al. (2006); RICHARDS and LEIS (1984); WATSON (1996d);

• Chromis sp. Meristics: Vertebrae:

Dorsal spines + rays:

XIII – XIV + 10 – 11

Anal spines + rays:

II + 10 – 12

-- Morphology: The body is moderately deep and laterally compressed. The head is short with a small, but strong protractible mouth and large eyes. One dorsal fin is developed, where the soft rays and the anteriorly spines are elongated.

-- Remark: The larva had already all adult meristic characters. The formation of stripes had already begun, which is characteristic for juveniles.

-- Reference: QUIGNARD and PRAS (1986a);

-- Reference material: 1 specimen; 11/97: S5 (P56344);

-- Distribution: In autumn the larva was caught at an epipelagic station south of Fuerteventura at a trawling depth of about 40 m during early morning.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

• Abudefduf luridus Meristics: Myomeres:

25 – 27

Dorsal spines + rays:

XIII + 15 – 16

Anal spines + rays:

II + 12 – 14

-- Morphology: The body is deep and laterally compressed. The head is short with a small protractible mouth. One dorsal fin is developed, where the soft rays are longer than the spinous part.

-- Reference: QUIGNARD and PRAS (1986a); WATSON (1996d);

-- Reference material: 2 specimens; 11/97: S10 (P56342-43);

-- Distribution: In autumn Abudefduf luridus was caught at an epipelagic station south of Fuerteventura at a trawling depth of about 40 m in early morning and at a neritic station northeast of Lanzarote at a depth of 20 – 64 m during the night.

Labridae Larvae hatch at a size of 1.5 – 2.7 mm having a large yolk sac, which is anteriorly protruding with the oil globule, if present, at the furthermost position. Most larvae are elongate and laterally compressed with a deep caudal peduncle, but some species are quite deep bodied. The number of myomeres ranges from 23 – 28, but is most times 25. The gut is straight, some kind of wrinkly and can reach up to 81 % of BL. When the gut coils is variable, sometimes also after flexion. The shape of head and snout is very variable. The mouth is small and tiny conical teeth form in both jaws at a size of t about 6.3 mm. No head spines are present. The dorsal fin is long based and its spines form first as soft filaments, but still can be distinguished by their triangular bases. Some spines may be elongated during postflexion stage.The origin of the anal fin is directly posteriorly of the vent. The last fin, which is formed, is the pelvic fin and can be delayed up to a size of 12 mm, but is usually ossified at about 7 mm.The count of the dorsal fin ranges within 8 – 13 for the spines and 7 – 15 for the dorsal rays.There are 14 – 15 principal caudal rays. The pigmentation is very variable, but most postflexion larvae are unpigmented. If present, melanophores form out above the vent, along the dorsal and ventral margins of the caudal peduncle, along the myosepta of the caudal region, dorsally on the gut, the brain, on the lower jaw and on the dorsal and anal fin elements. Already during preflexion stage melanophores occur at 2 or 3 of these locations. The development is direct and settlement takes place at a size of about 15 mm or some about 25 mm. No scales are formed before settlement. Some species bury themselves in the substrate during transformation. The best identification characters are the meristics like fin ray and spine counts.

-- Reference: LEIS and RENNIS (1983); QUIGNARD and PRAS (1986b); RICHARDS and LEIS (1984); JONES, LARA and RICHARDS (2006);

Inf. Téc. Inst. Canario Cienc. Mar. n°13

3 RESULTS

• Thalassoma pavo Meristics: Vertebrae:

Dorsal spines + rays:

VIII + 12 – 13

Anal spines + rays:

III + 10 – 12

-- Morphology: The body is slender and laterally compressed with a deep caudal peduncle. There are no head spines and the mouth is small. The single dorsal fin is long based, while the anal fin is about half the length.

-- Reference: QUIGNARD and PRAS (1986b);

-- Reference material: 1 specimen; 11/97: S6 (P5793);

-- Distribution: Only a single specimen was caught at an epipelagic station west of the channel between Lanzarote and Fuerteventura at 28 – 43 m depth during late night, almost in morning in autumn.

TRACHINOIDEI > Chiasmodontidae The larvae are slender and elongate with a short preanal length having a maximum of 50 % of the SL, which stays more or less stable throughout the entire larval development. Spination is present at the preoperculum, except for the genus Kali. Near or during the flexion tiny spines develop all over the body, except for the genus Kali. The pigmentation is strong, but restricted to the dorsally surface of the gut and gas bladder and spreads out during development. The caudal fin is the first to develop followed by the formation of the pecotral fins. The dorsal fin counts ranges from 6 – 13 spines and 18 – 28 rays. The anal fin may possess one spine and the rays range from 17 – 28.The pectoral fins form 10 – 15 soft rays, while the pelvic fins have one spine with 5 soft rays. The number of vertebrae varies from 33 – 48.

-- Remark: Spots are developed on the ventral and dorsal margin of the body, but staggered to each other. One spot is present at the caudal fin base and further dots at the forebrain.The body is full with tiny spines and a preopercular spination is present (the specimen is at least no Kali). There is one tooth on the tip of the lower jaw (Figure 63).

-- Reference: HARDY (2006b); WATSON, MATARESE and STEVENS (1984);

-- Reference material: 1 specimen; 11/97: S4 (P65181);

-- Distribution: The only chiasmodontid larva was found at a neritic station southeast of Fuerteventura in autumn at a depth of 22 – 35 m during daytime.

Inf. Téc. Inst. Canario Cienc. Mar. n°13

SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

BLENNIOIDEI > Blenniidae In general within this family great morphological variations occur. But in common is the elongate the body. The head is short, rounded and broad, but becoming elongate during further development. Small teeth develop. The preanal length is moderate to short, but usually to midbody length.The gas bladder is present.The number of myomeres ranges from 30 – 40. Preopercular spination is present and may be complex. The pectoral fin rays are very long. The pigmentation is initially very light, but increase in intensity with development. It is restricted to along the ventral midline of the body, in the area of the gut and at the dorsal surface of the gas bladder. For the identification the counts of the ventral fins are.

-- Reference: DITTY, CAVALLUZZI and OLNEY (2006); LEIS and RENNIS (1983); MATARESE, WATSON and STEVENS (1984);

• Parablennius sp. Body is elongate. In adults, tentacles form on nasal openings, above eyes in all Parablennius. Also canine teeth are present in lower jaws and in most in upper jaws. Usually no interspace is present between the two dorsal fins. Identification of the studied specimens to genus level was done using meristic counts (Figure 64).

-- Reference: ZANDER C. D. (1986);

-- Reference material: 2 specimens; 11/97: S16 (P75876-877);

-- Distribution: The specimens of that genus were found at a neritic station west of Gran Canaria at 42 – 66 m depth during the night in autumn.

GOBIOIDEI > Gobiidae Larvae hatch at a size of 1.7 – 4.4 mm with a functional mouth and a small yolk sac. The size at flexion varies from 2.7 – 3.8 mm. The number of myomeres ranges from 24 – 27. The shape of the body is variable from stout to moderately elongate. The gut is initially straight or curved and reaches to 50 – 65 % of SL. After flexion the gut is strongly s-shaped, but is never fully coiled. The gas bladder in larvae is large and is situated slightly anteriorly of the midbody level. The head is moderately in size and rounded with a gently profile, but changes in shape with development. The eyes are large. Teeth form in both jaws at a size of 7 mm. A medial finfold and the pectoral fins may be present already when hatching, and if not, they develop early. The bases of the fins are short with low fin ray counts, except for maybe the dorsal and the anal fins. Often the dorsal fins are separated, while the origin of the second dorsal fin is at about midbody level. The pelvic fin is the last to form at a size of about 7 mm with one spine and 2 – 5 rays. The pelvic fins can be united forming a sucking pelvic disc. The pigmentation is variable, but restricted to the dorsal surface of the gas bladder and the gut, especially on the hindgut anteriorly to the anus. Further areas of pigmentation are along the ventral margin of the body, at the anal fin base and often in the region of the isthmus. In many species melanophores are present in the area of the caudal peduncle, and on several fins. Less common, but in some species present, is the pigmentation at the dorsal surfaces of the tail, trunk and head. The pattern of pigmentation is important in the identification to genus or species level. The development is direct and no changes occur during settlement, except in some species the eyes move dorsally and the head gets laterally more compressed.

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3 RESULTS

-- Remark: I could distinguish two different types of larvae due to their different pigmentation patterns. But neither via the pigmentation pattern or fin ray counts the larvae could have been identified to a lower taxonomic level. Some further larvae could not be allocated to one of the distinguished types because of their bad condition. Type I gobies had a double row of stellate melanophores along the dorsal fin base (Figure 67) and Type II gobies had a blotch behind the dorsal fin base (Figure 68).

-- Reference: LEIS and RENNIS (1983); RUPLE (1984); YEUNG and RUPLE (2006);

-- Reference material: 882 specimens; 11/97: S4 (P782233-246; P782260-782847); S7 (P782849); S10 (P781972-782232); S13 (P781968-971; P782247-255); S16 (P782256-259); 04/99: S16 (P782848);

-- Distribution: Type I gobies was only found in autumn and mainly at neritic stations within a depth range of 20 – 66 m during the day as well as during the night. The highest number was caught northeast of Lanzarote and a lower number was each found southeast of Fuerteventura and west of Gran Canaria. Some larvae occurred also at an epipelagic station southeast of Fuerteventura in a depth of 90 – 107 m during late night, almost dawning. The type II gobies larvae were found only at neritic stations in autumn, so southeast of Fuerteventura and Lanzarote in a depth range of 13 – 42 m during the day and west of Gran Canaria at a depth of 42 – 66 m during the night. In the spring some specimens were caught at a neritic station southwest of Gran Canaria at 18 – 61 m depth in night. Some specimens were damaged or the pigments vanished, that identification was not possible. Larvae were caught at an epipelagic station southeast of Fuerteventura at a depth of 90 –107 m at late nighttimes, almost morning in autumn.

• Crystallogobius linearis Meristics: Vertebrae:

29 – 31

Dorsal spines + rays:

Males: II – III + I + 18 – 20 Females: 0 or rudimentary

Anal spine + rays:

I + 20 – 21

Pectoral rays:

15 – 19

-- Morphology: The body is elongate and laterally compressed. The entire body is transparent, so that internal melanophores and the gas bladder are clearly visible. Although the preserved specimens studied were not transparent anymore the melanophores on the gas bladder were still clearly visible (Figure 65). Sexual dimorphism is developed. Males form two front canine teeth in lower jaw when adult. The pelvic disc is deep in males but reduced or lacking in females. The first dorsal fin is differently developed in males and females (Figure 65).

-- Reference: MILLER (1986);

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SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Reference material: 30 specimens; 11/97: S10 (P781960-965); S13 (P781936); S18 (P781937-959);

-- Distribution: This species is a new record from the Canary Islands, but specimens were already recorded from Madeira (WIRTZ et al. 1995).The specimens were caught at neritic stations in autumn, so northeast of Lanzarote at 20 – 64 m during night and east of Gran Canaria in a depth of 116 – 146 m. A single specimen was found at an epipelagic station southeast of Fuerteventura at a depth of 90 – 107 m also in autumn.

• Lesueurigobius heterofasciatus Meristics: Vertebrae:

Dorsal spines + rays:

VI + I + 14 – 15

Anal spine + rays:

I + 14 – 15

Pectoral rays:

22 – 24

-- Morphology: Body is elongate. The head profile is rounded. An interspace between the two dorsal fins is present.

-- Pigmentation: The pattern of different intense brownish vertical bands along the body is very prominent and a distinctive character of that species and was also clearly visible in the studied specimens (Figure 66).

-- Reference: MILLER (1986);

-- Reference material: 2 specimens; 11/97: S16 (P781966-967);

-- Distribution: Two specimens were found at a neritic station west of Gran Canaria at a depth of 42 – 66 m during the night in autumn.

SCOMBROIDEI > Gempylidae The shape of the body is triangular, rectangular or elongate and laterally compressed. The number of vertebrae ranges from 32 – 58.The maxillae are exposed and anteriorly strong canine teeth are present.The base of the spinous dorsal fin is longer than the base of the soft fin rays. Usually 3 spines are present at the anal fin. The spines of the dorsal, anal and pelvic fins are serrated. All larvae of this family are pelagic.

-- Reference: COLLETTE et al. (1984): RICHARDS (2006j);

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• Diplospinus multistriatus Meristics: Myomeres:

58 – 61

Dorsal spines + rays:

XXX – XXXVI + 36 – 42

Anal spines + rays:

II + 28 – 33

Pelvic spines + rays:

I+0

Pectoral rays:

11 – 13

-- Morphology: In comparison to other species of that family the body is relatively deep. The ventral gut is distinctly flattened. At the lower jaw tip two forward protruding spines are present.The flexion takes place within a length of 8 – 10 mm SL.The dorsal spines are almost completely developed in larvae more than 8 mm SL. The very long spines of the dorsal fin and the spine of the pelvic fin are strongly serrated (Figure 69). The pelvic spine is also elongate, reaching beyond the vent while the rays are absent. The preopercular spine is only moderately serrated. No dorsal or anal finlets develop.

-- Pigmentation: The gular membrane is pigmented. Spots form along the base of the dorsal fin and on the membrane of the first dorsal fin. Further pigmentation is on fore- and midbrain, on the gut, on the tip of the lower jaw and posteriorly to the eyes.

-- Reference: COLLETE et al. (1984); FAHAY (1983); PARIN (1986a); RICHARDS (2006j);

-- Reference material: 3 specimens; 11/97: S10 (P73360); S16 (P73361-62);

-- Distribution: All specimens were only caught at neritic stations during the night in autumn, so northeast of Lanzarote at a depth of 20 – 64 m and west of Gran Canaria at a depth of 42 – 66 m.

Trichiuridae The body is elongate and laterally strongly compressed with an initially short gut, which elongates during flexion and postflexion stage. The maxilla is enclosed be the preorbital and the anterior are strongly developed. The fin formation starts with the development of the dorsal fin. Both, the dorsal as well as the anal spines are serrated, while the dorsal spines are not longer than the dorsal rays. And the two anal spines are located directly behind the vent. The pelvic fins are reduced to one spine and one ray, where the spine is serrated too. But the pelvic fins may be even absent. Additionally the caudal fin is reduced or absent too. Prominent for this family is the very high vertebrae count, which ranges from 100 to almost 200. For identification the meristic counts are crucial.

-- Reference: COLLETTE et al. (1984); RICHARDS (2006k);

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• Benthodesmus elongates Meristics: Dorsal spines + rays:

XLIV – XLVI + 104 – 109

Anal spines + rays:

II + 93 – 102

Pelvic spine + ray

I+1

Pectoral rays:

-- Morphology: The body is very elongate and laterally compressed. The pelvic fins are reduced and they are positioned below the origin of the pectoral fin. The fin spines are serrated.

-- Pigmentation: The pigmentation is the same as in Lepidopus caudatus with melanophores occurring along the gut, on the snout and on the top of the head. Especially on the dorsal region of the trunk pigmentation develops. When the dorsal rays form out, the pigments start to spread backwards to the caudal region.

-- Remark: Identification was done by family features and by meristics characters. The head was small with a protruding tooth on the tip of lower jaw. In the studied specimen the first dorsal spine was slightly longer and serrated only on the posterior edge.

-- Reference: OLIVAR and FORTUÑO (1991); PARIN (1986b);

-- Reference material: 1 specimen; 03/02: S16 (P73363);

-- Distribution: The only specimen was found in late winter during the day at an epipelagic station southeast of Fuerteventura at a depth of 185 – 211 m.

• Lepidopus caudatus Meristics: Dorsal spines + rays:

IX + 90 – 107

Anal spines + rays:

II + 60 – 65

Pelvic spine + rays:

Adults: I + 1 Juveniles: I + 2

Pectoral rays:

-- Morphology: The number of myomeres is 113 in specimens of 10 mm in SL and the flexion takes place at a SL of 12 mm at least in Mediterranean species. The body is very elongate with a body depth of about 10 % of NL in preflexion larvae and increases during flexion to about 14 % of SL, but decreases again to about 8 % of SL in larvae longer than 30 mm. The snout is small in early larvae becoming more acute with further development. The gut is very short throughout the whole larvae stages with a length of 1/4 – 1/2 of the SL. The first spine of the dorsal fin is elongated with an appendix, which is variable in length

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during development. The head spination consists of two or three preopercular spines, which are present in larvae from 10 – 30 mm in SL.

-- Pigmentation: Melanophores develop along the gut, on the snout and on the top of the head. A higher concentration of the pigments occurs along the dorsal surface of the trunk, which start to spread backwards to the caudal region, when the dorsal rays form out.

-- Remarks: Some sparse dots were present on the finfold and along the ventral midline of the caudal peduncle in the studied specimens. A lateral pigmentation on the head at the level of the eye was also found.

-- Reference: OLIVAR and FORTUÑO (1991); PARIN (1986b);

-- Reference material: 6 specimens; 03/02: S11 (P73369); S18 (P73364-68);

-- Distribution: Specimens were only caught southeast of Fuerteventura in late winter during the day at an epipelagic station in a depth range of 31 – 140 m and at an epipelagic station at a depth of 470 – 695 m.

• Scombridae The number of vertebrae varies from 31 – 66, but within the species the number is quite stable with a very narrow range of variability. The body is deep with a large head and a distinct snout, which may be elongated. The jaws are well developed with no canine teeth. Head spination is present at the opercle, supraocular and posttemporal and pterotic region.The preanal length is shorter than the midlength of the body in early larvae, but during development the anus migrates posteriorly. The caudal fin forms first. Usually the first dorsal fin forms before the second, except for the genus Scomber. Simultaneously to the first dorsal fin the pelvic fins form out in most genera, with one spine and 5 rays. The pectoral fins are positioned high on the body with a high ray count ranging from 19 – 36. Prominent for this family are the 5 – 12 finlets following the second dorsal and the anal fin. The full fin complements are fully developed with a size of about 15 mm. For species identification meristic characters are useful.The pigmentation is very variable within this family, but pigments occur mainly on head, ventrally on tail in early larvae increasing with development and dorsal pigmentation adds.

-- Reference: COLLETTE et al. (1984); FAHAY (1983);

• Scomber colias Meristics:

102

Development:

Vertebrae (Myomeres):

14 + 17 (31)

Dorsal spines + rays:

VIII – XI + 12 + 4 – 5

Anal spines + rays:

II + 11 + 5

Hatching: ~3 mm

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SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Morphology: The larvae hatch with a size of 3 mm and the tiny premaxillary teeth form already at 4 mm and persist throughout the larval development till the juvenile stage. The head is small with a length less than 1/3 of the SL and the snout is rounded, so the head profile curves gently. The jaws are equally in length and tiny premaxillary teeth develop. The number of myomeres is 31 and all vertebrae are ossified at a size of 7.6 mm. There are no larval specializations, so no spination forms out. Unique for this genus is, that the first dorsal fin develops after the second has formed and is completed with a size of 13 mm. In the studied specimens all fins as well as the finlets had already formed out (Figure 70).

-- Pigmentation: Strong pigmentation develops ventrally on the trunk especially at the level of the midbody and posterior to the anus along the tail. Further pigmentation is located at the fore- and hindbrain, at the tip of the lower jaw.The same pigmentation pattern is found in S. scombrus LINNAEUS, 1758, but it develops later in larval development and a pigmentation at the cleithral symphysis is present, which is absent in S. colias (Figure 70).

-- Reference: COLLETTE (1986); COLLETTE et al. (1984); FAHAY (1983);

-- Reference material: 50 specimens; 03/02: S1 (P74172); S4 (P74173-174); S5 (P74177); S6 (P74179); S11 (P74180-218); S14 (P74178); S17 (P74175-176); S19 (P74169-71);

-- Distribution: Larvae only were caught in late winter southeast of Fuerteventura, whereas most larvae occurred at epipelagic stations in a depth range of 22 – 140 m in day as well as in night time. Further each one specimen was caught at a neritic station in depths of 15 – 60 during the night and some larvae were caught also at mesopelagic stations in a depth range of 311 – 378 m and 617 – 915 m each during the day.

STROMATEIODEI > Nomeidae The body is deep during whole larval period and even increases with development to more than 1/3 of SL in postflexion larvae.The length of the gut is less than half of NL initially and increases to more than 50 % of SL in late larvae. Also the head length increases to about 1/3 of the SL. Flexion takes place at a size range from 4.5 – 5 mm SL. Along the preoperculum very small, almost not visible spines start to develop at 5.2 mm and disappear again at a size of approximately 11 mm of SL. Head pigmentation is present dorsally on the head, at the tips of the jaws, and ventrally from the snout to the cleithrum. Further melanophores develop at the dorsal and lateral surface of the gut and ventrally along the midline of the caudal peduncle in preflexion stage, but latter vanish during postflexion stage. All other melanophores increase in number and size in larvae exceeding 7 mm SL and stellate melanophores starts to cover the entire body.

-- Reference: OLIVAR and FURTAÑO (1991);

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• Cubiceps gracilis Meristics:

Development:

Vertebrae (Myomeres):

33 – 34 (31)

Dorsal spines + rays:

IX – XI + 22 – 24

Anal spines + rays:

II – III + 20 – 23

Pectoral rays:

20 – 24

Hatching: ~3 mm Flexion: ~6 – 9 mm

-- Morphology: In preflexion larvae the body is deep with 29.5 % of the NL and deepens more with development to 37 % of SL in postflexion stage, but decreases again in juveniles of more than 20 mm to 32 % of SL. The gut is short with 36 % of NL and increases with development to 55 % of SL. Flexion takes place at 6 – 9 mm. All fin rays of caudal, anal, pectoral and pelvic fins are developed at a size of 13.2 mm. The second dorsal fin forms after the development of the first dorsal fin is completed. Since the studied specimen was in the late postflexion stage, all rays had already developed and species identification was done by meristic counts. There are no preopercular spines present.

-- Pigmentation: The pigmentation on the head and on the trunk is strongly developed. Already the late preflexion larvae lack the pigmentation series ventrally on the caudal peduncle, but a series of four to five spots occur on the caudal fin. During postflexion stage pigmentation spreads from the head dorsally backwards over the trunk. Dispersal of pigments also occurs on the caudal peduncle, where melanophores start to develop.

-- Reference: HAEDRICH (1986); OLIVAR and FURTAÑO (1991);

-- Reference material: 1 specimen; 03/02: S13 (P691561);

-- Distribution: A single specimen was found southeast of Fuerteventura at a depth of 632 – 1035 m in day during winter.

CAPROIDEI > Caproidae The body is very deep bodied and laterally highly compressed. An occipital crest is present and the eyes are large.The mouth is large and protractile. The dorsal spines are elongated and pectoral fin spines are present. The number of vertebrae is low.

-- Reference: GREENE and POWELL (2006); QUÉRO (1986);

• Capros aper Meristics:

104

Dorsal spines + rays:

IX – X + 23 – 25

Anal spines + rays:

II + 22 – 24

Pelvic spine + rays:

I+5

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SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

-- Morphology: Already during preflexion the body deepens and the size of the head increases. A medial serrated ridge is present at the cranium and a pair of serrated ridges develops along the lower jaw and at the supraocular region. At the preoperculum many spines develop and the entire body surface is full with tiny spines, which were clearly visible in the studied specimens at a size of about 10 mm (Figure 71).

-- Pigmentation: Large stellate melanophores are present dorsally, laterally and ventrally on the body, as well as very few spots are positioned on the head in newly hatched larvae. During preflexion the entire body gets pigmented with the exception of the caudal region (Figure 71).

-- Reference: QUÉRO (1986); TIGHE and KEENE (1984);

-- Reference material: 9 specimens; 03/02: S3 (P42161); S6 (P42162); S11 (P42163); S13 (P42164-165); S17 (P42166-168); S19 (P42169);

-- Distribution: All specimens were only caught southeast of Fuerteventura mainly at mesopelagic depths at the day in late winter, where the depths ranged of 311 – 1035 m. Some larvae were also found at neritic stations within 31 – 140 m in daytime and 30 – 139 m during the night.

¬ PLEURONECTIFORMES The body of the flatfish is highly laterally compressed, where one side becomes the blind underside with development. The gut coils and therefore the preanal length is short, which decreases even more with development. Initially the larvae have each one eye at one side, but during metamorphosis one eye migrates to the other side of the head. Consequently, both eyes are positioned then either on the left (sinistral) side, or on the right (dextral) side of the body. In most flatfish groups the nasal organ migrates also with the eye, but only close to the dorsal midline. The single groups of faltfish consist of species having the eyes on the same side. The dorsal and anal fin bases are extremely long. The pelvic fin on the blind side can be reduced, have a low fin ray count and are differently positioned.

-- Reference: AHLSTROM et al. (1984 C); OLIVAR and FURTAÑO (1991);

PLEURONECTOIDEI > Bothidae Species of that family have the eyes usually on the left side. The size at hatching ranges from 1.8 – 2.6 mm within that family. The body is highly compressed resulting in being very thin to even transparent. The eyes are ovoid with a choroid tissue on the ventral margin. The eye migrates through an opening under the anteriorly part of the dorsal fin. The dorsal and anal fin are very long based and reach to the caudal fin, which is very small or even reduced. Prominent for this family is the elongation of the first or second dorsal fin ray, which disappears, when the eye starts to migrate. Spination occurs at the region of the urohyal, basipterygia, cleithra and epiotics.The pigmentation is very light or not existent. Larvae of bothids reach a large size, before transformation starts.

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-- Reference: AHLSTROM et al. (1984 C); LARA (2006);

• Arnoglossus sp. The body is quite elongate.The second dorsal ray can be very elongate and ornamented.The pigmentation is very prominent for this genus and consists of melanophores above the brain, ventrally on the gut, at the dorsal surface of the gas bladder, along the dorsal and ventral midline and between the eyes. Some species have complete or partial bars at the tail region.The pigmentation in the studied specimens was clearly developed on the gas bladder, along the dorsal and ventral midline of the body (at the bases of the medial fins) and they had a partial bar at the tail region (Figure 72). Number of vertebrae ranges from 37 – 48. Arnoglossus develop scale spines.

-- Reference: AHLSTROM et al. (1984 C); NIELSEN (1986);

-- Reference material: 9 specimens; 04/99: S6 (P855956); 05/99: S1 (P855955); 03/02: S2 (P855957); S4 (P855951-954); S19 (P855949-950);

-- Distribution: Larvae were found during winter and spring only at epipelagic stations southeast of Fuerteventura during the night in depth ranges of 13 – 139 m. Only one specimen was caught in late spring far east of Fuerteventura at a depth of 38 – 40 m also during the night.

• Arnoglossus imperialis Meristics: Dorsal rays:

95 – 106

Anal rays:

74 – 82

-- Morphology: The body is rather elongate.The lower eye is more anteriorly positioned than the upper, when migrated to the adult position and both eyes are separated by a bony bridge in adults. The species identification was done only by meristic counts.

-- Pigmentation: The pigmentation follows the prominent larval pattern for that genus, with a partial bar at the tail region.

-- Reference: NIELSEN (1986);

-- Reference material: 4 specimens; 04/99: S16 (P855947); 05/99: S1 (P855945); 03/02: S15 (P855946); S19 (P855944);

-- Distribution: In winter a specimen was caught at an epipelagic station southeast of Fuerteventura at a depth of 30 – 99 m at daytime. In spring larvae occurred at a neritic station southwest of Gran Canaria at 18 – 61 m depth during the night and at an epipelagic station far east of Fuerteventura at a depth of 38 – 40 m also during the night.

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• Arnoglossus rueppelii Meristics: Dorsal rays:

110 – 118

Anal rays:

86 – 94

-- Morphology: The body is very elongate for a flatfish, which was prominent in the studied specimen. The eyes become very large in adults.

-- Pigmentation: The pigmentation follows the prominent larval pattern for that genus, with a partial bar at the caudal region.

-- Reference: NIELSEN (1986);

-- Reference material: 1 specimen; 04/99: S12 (P855948);

-- Distribution: The only specimen of A. rueppelii was caught at an epipelagic station southwest of Gran Canaria at a depth of 38 – 49 m during the night in spring.

• Bothus podas Meristics: Dorsal rays:

85 – 95

Anal rays:

63 – 73

-- Morphology: The body is very deep and compressed. The width between the eyes is much larger in males than in females, what is not developed in larvae.

-- Pigmentation: There is no pigmentation in larvae, except for pigmentation at the tip of the notochord in preflexion stage. The non-existent pigmentation (Figure 73) led me to this species and was verified by the fin ray counts.

-- Reference: NIELSEN (1986);

-- Reference material: 16 specimens; 11/97: S6 (P855964-965); S7 (P855961); S13 (P855966-968); 04/99: S5 (P855959); S6 (P855958); S8 (P855969); S13 (P855971); S14 (P855973); 05/99: S3 (P855962-963); 03/02: S2 (P855972); S3 (P855960); S5 (P855970);

-- Distribution: In autumn larvae were caught at epipelagic stations west of the channel between Lanzarote and Fuerteventura in depth of 28 – 43 m in the day and southeast of Fuerteventura at 90 – 107 m depth in early morning. Further larvae were caught

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at a neritic station southeast of Lanzarote in a depth of 13 – 41 m at daytime. In winter larvae were found southeast from Fuerteventura at a depth of 30 – 123 m during the night. Bothus podas was caught at a mesopelagic station southeast of Fuerteventura at 440 – 665 m depth during the late daytime. In spring specimens were found at a neritic station southwest of Gran Canaria at a depth of 33 – 53 m during the night and in the day at mesopelagic stations southeast of Fuerteventura in a depth range of 382 – 511 m as well as southwest of Gran Canaria at a depth of 480 – 694 m. In late spring some specimens were caught at the epipelagic station far north of Gran Canaria at a depth of 42 – 47 m during the night.

¬ TETRAODONTIFORMES All larvae are pelagic and most have a direct development. Only few characteristics are uniform among the Tetraodontiformes and some other are also found in other orders, e.g. the late formation of the caudal fin and scale specializations, which may form very early, as well as the dermal sac is not only found within this order. The large head with the large eyes and the small mouth, the presence of the gas bladder, the coiled and massive gut and the low number of myomeres are characteristic for this order. The pelvic fins are reduced or lacking. In the suborder Tetraodontoidei larvae are sheltered in a vesicular sac. In comparison to the rest of the body is the tail small and laterally compressed. Larvae are strongly pigmented. Except for the molids no dermal spines develop and the tail does not form. The larvae of the suborder Balistoidei are mainly deep and wide bodied. If the tail seems to be reduced, it never had developed. Larvae are moderately to strongly pigmented. In both suborders the transformation takes place at a small larval size and species have a long, not specialised juvenile stage.

-- Reference: ABOUSSOUAN and LEIS (1984); LEIS (1984);

BALISTOIDEI > Balistidae Larvae hatch at 1.5 – 2 mm NL and no jaws or eye pigmentation is developed.The body is less pigmented and cylindrical with a slightly small lateral compression in shape. With development the body deepens fast and is almost round in cross section at the level of the trunk, but the tail remains compressed. Only in newly hatched larvae a kind of inflated sack shelters the trunk, but disappears soon and close to the flexion, the gill opening reduces till a pore remains. In Balistidae the spines of the dorsal fins are first to form, followed by a simultaneously formation of dorsal rays, the anal fin and the pectoral fins. The caudal fin is the last developing. The fin formation is completed at around 5 – 6 mm SL. The pelvic fins are absent, but a modified pelvic spine develops. Prominent in this family is the elongation of the first spine, which can be armed with hooks during preflexion stage. During preflexion spines develop on preoperculum and disappears shortly before dermal spines develop on the jowl, cleithrum, on the forehead and laterally on the gut, where they spread all over the entire body by midflexion.This spinulated remain till into the juvenile stage, but do not transform into the prominent specialised scales in adults. Pigmentation concentrates on the brain and gut. In preflexion larvae a series of melanophores develops along the ventral midline from posteriorly of the anus to the tail, where bands may develop. The spinous dorsal fin membrane is strongly pigmented and from there pigments spread out all over the trunk with further development.

-- Reference: ABOUSSOUAN and LEIS (1984); LYCZKOWSKI-SHULTZ and INGRAM (2006);

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• Balistes capriscus Meristics:

Development:

Vertebrae:

Dorsal spines + rays:

III + 2 + 22 – 27

Anal rays:

23 – 26

Pelvic rays:

Pectoral rays:

13 - 15

Caudal rays:

Hatching: 1.7 mm Flexion: ~ 3 mm Transformation: ~ 10 mm

-- Morphology: The body is deep and the eyes are near the dorsal edge. The mouth is small. The second dorsal fin and the anal fin are opposite in their position. Only one spine is present instead of pelvic fin. Juveniles have no elongated caudal rays.

-- Pigmentation: In preflexion stage melanophores occur on the brain, nape, jowl and in gill cavity, on gas bladder, on the dorsal surface on the gut, along the ventral margin of the caudal peduncle and the caudal finfold, along the anal fin base, if present than few spots on above und below the notochord tip. The overall pigmentation pattern gets more intense with development. But there are no pigments on the front of the head, on the snout or chin, till late postflexion stage. During flexion melanophores develop during flexion on the membrane of the first and second dorsal spine. A vertically bar develops on the causal peduncle and is present at 3.7 mm. Some spots develop on caudal fin rays. During postflexion stage, the pigmentation spreads all over the body.

-- Reference: TORTONESE (1986d); LYCZKOWSKI-SHULTZ and INGRAM (2006);

-- Reference material: 3 specimens; 11/97: S4 (P88324-325); S16 (P88326);

-- Distribution: This species was exclusively caught at neritic stations in autumn. Larvae were found southeast of Fuerteventura at a depth of 22 – 35 m during the day and west of Gran Canaria at 42 – 66 m depth during the night.

TETRAODONTOIDEI > Tetraodontidae The development stage at which larvae hatch is depending on species. Consequently some have partially formed jaws and fully pigmented eyes, while other larvae have no jaws and no eye pigmentation developed. A viscular sac moderate in size shelters the head and the trunk of the larvae. But a large yolk sac is in common, which disappears during preflexion or persists till after flexion. The body is initially cylindrical and increases in heights and width with further development, but become laterally compressed. After the absorption of the yolk sac, the gill opening is closed to a pore by a thin membrane. The preanal length is initially around 50 % BL but increases to 75 % BL with further development. The tail is well developed till flexion and the notochord tip remains till short after postflexion. The buds of the pectoral fins may be absent or present, but the formation of the pectoral fins is the first in the fin development sequence. After the pectoral fins, the dorsal and the anal fin develop simultaneously and the last fin to form is the caudal fin. No spines form in dorsal or anal fins and the pelvic

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fins are absent. The vertebrae count is low and range within 17 – 19. The first body spinules appear at the ventral side of the larvae during preflexion and usually spread fast over the entire body, sometimes this may be delayed or no spreading occurs at all. The pigmentation is initially restricted to the area above gut, brain and the yolk sac, but covering the head and trunk before flexion. There is no distinct pelagic juvenile stage.

-- Reference: LEIS (1984); LYCZKOWSKI-SHULTZ (2006);

• Canthigaster capistrata Meristics: Dorsal rays:

9 – 10

Anal rays:

8–9

-- Morphology: The body is moderately deep. No fin spines are present. The head profile is concave and the jaws are protuberant, which was a prominent characteristics in the studied specimen.

-- Reference: MOURA and CASTRO (2002); TORTONESE (1986e);

-- Reference material: 1 specimen; 04/99: S13 (P89248);

-- Distribution: The only specimen was caught at a neritic station southwest of Gran Canaria at a depth of 33 – 53 m during night in spring.

• Sphoeroides sp. The body is moderately slender. The dorsal and anal fins are opposite in position. No pelvic fins develop. No information of pigmentation development is given, so no verification of species identification can be done. Most probable the specimens are Sphoeroides pachygaster (MÜLLER and TROSCHEL, 1848). In the studied specimens the body spinules were well developed (Figure 74).

-- Reference: LYCZKOWSKI-SHULTZ (2006); TORTONESE (1986e);

-- Reference material: 8 specimens; 11/97: S7 (P89252); S13 (P89253); S16 (P89249-251); 11/00: S1 (P89256); S11 (P89254-255);

-- Distribution: Specimens were only caught in autumn mainly at neritic stations. So southwest of Gran Canaria at a depth of 24 – 57 m in the evening, southeast Lanzarote at 3 – 41 m depth during the day and west of Gran Canaria at 42 – 66 m depth at night. Larvae occurred also at epipelagic stations north of Gran Canaria at a depth of 55 – 90 m at night and southeast of Fuerteventura at a depth range of 90 – 107 m during the morning.

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3.3. Community analysis The similarity in larval species composition among single stations is shown in Figure 10. The cluster analysis revealed no clear community groupings. An assemblage cluster of 5 groups can be distinguished at a similarity level of about 7 %. The subsequent MDS plots indicate which factor might influence the community structure.

Figure 10. The dendogram shows species clustering analyses of the fish larvae assemblage with group average linking based on a Bray – Curtis similarity matrix

At a significance level of 0.1 % ANOSIM calculated a difference (Global R: 0.28) between seasons with the difference being highest between January and November (Pairwise test: r: 0.412; significance level: 0.1) and January and May (Pairwise test: r: 0.398; significance level: 0.2) and lowest between April and May (Pairwise test: r: 0.021; significance level: 0.031). The MDS plot in Figure 11 shows the community structure based on the factor season, more precisely the month of material collecting. Not that obvious is the greatest difference between January and November. A tendency of a grouping between a day and night community can be seen in Figure 12, even if it is overlapping. ANOSIM found differences (Global R: 0.123) at a 0.1 % significance level. Daytime of collecting is reflected in the community structure. The factor habitat seems to influence the community structure of fish larvae slightly (Figure 13). A transition zone, especially between mesopelagic stations and epipelagic stations can be seen. With the exception of one station the neritic and mesopelagic stations are clearly separated. The ANOSIM test shows differences in the community structure (Global R: 0.121) at a significance of 0.1 %, where the greatest separation (Pairwise test: r: 0.16; significance level: 0.1) of communities is found between the neritic and mesopelagic habitat and the lowest between epi- and mesopelagic habitat (Pairwise test: r: 0.081; significance level: 6.1).

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Figure 11. 2-d MDS based on a Bray – Curtis similarity matrix. Symbols and numbers represent the different seasons of January (1), March (3), April (4), May (5) and November (11) of investigations.

Figure 12. 2-d MDS based on a Bray – Curtis similarity matrix. Symbols represent the collecting time during day (1) or night (2).

A grouping can be seen between the upper 150 m of collecting depth to the collecting depth of 150 to 600 m (Figure 14). Larvae found deeper than 600 m seem not to differ from the species community in the mediate depths. A general difference of communities is also reflected by ANOSIM (Global R: 0.165; significance: 0.1 %), where the greatest separation (Pairwise test: r: 0.214; significance level: 3.4) is found between the community of the SF and the NACW, whereas between the water mass of NACW and AAIW are almost no differences found (Pairwise test: r: 0.041; significance level: 36.2).

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Figure 13. 2-d MDS based on a Bray – Curtis similarity matrix. Symbols represent the habitat of neritic zone (1), epi- (2) and mesopelagic (3) zone.

Figure 14. 2-d MDS based on a Bray – Curtis similarity matrix. Symbols represent the depth distribution from 0 – 150 m (1), 150 – 600 m- (2) and more than 600 m depth (3).

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Figure 15. 2-d MDS based on a Bray – Curtis similarity matrix. Symbols represent the different islands of Fuerteventura (1), Lanzarote (2), Gran Canaria (3), La

There are differences in the larval community between the different islands (Figure 15). Especially Fuerteventura and Gran Canaria seem to have different communities. The two oceanic stations from La Bocaina east from Fuerteventura are similar to the Fuerteventura community. The oceanic stations northern from the Islands are similar to the community of GC. The two stations western of Tenerife and close to La Gomera are not similar at all. The global ANOSIM test showed there were significant differences (Global R: 0.162, significance: 0.1 %) in assemblage composition among the Islands. The highest difference (Pairwise test: r: 0.679; significance level: 6.7) shows ANOSIM between the Islands of Fuerteventura and Tenerife/La Gomera and Fuerteventura and the oceanic station north of Tenerife. Even the communities between Tenerife/ La Gomera and oceanic north of Tenerife are significantly different (Pairwise test: r: 0.5; significance level: 33.3) though with a low significant level. In general the Tenerife community is different from all other, but still most similar to the community of Gran Canaria (Pairwise test: r: 0.182, significance level: 7). Not overlapping at all are the communities between the La Bocaina assemblage and the community north of Tenerife (Pairwise test: r: 1.0; significance level: 33.3). Most similar are the communities of La Bocaina and the assemblage around Lanzarote (Pairwise test: r: -0.008; significance level: 50.8).

3.4. Diversity of fish larvae For investigating the influence of the different spatial and seasonal factors of seasons (Months), daytime, type of habitat, and regions (single islands) on indices of diversity (Total Species S, Shannon Index H’ and Simpson’s D) and Evenness (Pielou’s Evenness Index J’) ANOVA analyses were performed. The indices of H’, J’, D and S for each station are listed in Table 8. A scatter plot in Figure 16 shows eventual trends among the indices in dependence on the habitat type. Stations with only one single specimen have a Simpson Index of 1 and a Shannon Index H’ of 0 and consequently no Evenness Index can be calculated. A low Evenness J’ indicates a dominance phenomenon. Especially within neritic stations, dominance of species occurs (Figure 16), while in the epi- and mesopelagic realms the species are distributed more evenly.

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Table 8. The diversity indices of Total Species S, Pielou’s Evenness Index J’, Shannon H’, and Simpson’s D are shown for each station (stations resulted in the catch of only one species are excluded). Additionally, the habitat type of each station is listed, 1 refers to neritic, 2 to epipelagic and 3 to mesopelagic tows. Pielou’s Evenness

Total Species [S]

11/97/4

0.187

0.541

0.699

11/97/5

0.946

1.040

0.375

11/97/6

0.799

1.841

0.200

11/97/7

0.611

1.095

0.492

11/97/9

1.000

1.099

0.333

11/97/10

0.393

1.158

0.556

11/97/13

0.502

1.477

0.443

11/97/16

0.750

1.798

0.273

11/97/18

0.439

0.707

0.684

04/99/5

0.970

1.561

0.222

Index [J’]

Shannon Index [H’]

Simpson’s Diversity

Trawl

Index [D]

Habitat

04/99/6

0.832

1.154

0.388

04/99/12

0.870

1.692

0.231

04/99/13

1.000

1.792

0.167

04/99/14

0.929

2.138

0.148

04/99/16

0.798

1.660

0.265

05/99/1

0.916

2.013

0.155

05/99/2

0.917

1.475

0.265

05/99/3

0.649

1.349

0.426

05/99/4

1.000

1.099

0.333

05/99/5

1.000

0.693

0.500

05/99/6

0.677

1.624

0.273

01/00/1

0.773

1.244

0.375

01/00/3

1.000

0.693

0.500

01/00/5

0.961

1.332

0.280

01/00/7

0.918

0.637

0.556

11/00/1

1.000

0.693

0.500

11/00/12

1.000

1.099

0.333

11/00/17

0.863

0.598

0.592

03/02/1

1.000

0.693

0.500

03/02/2

1.000

0.693

0.500

03/02/3

0.917

1.643

0.220

03/02/4

0.914

1.471

0.260

03/02/5

1.000

1.099

0.333

03/02/6

0.943

1.834

0.180

03/02/11

0.549

1.205

0.487

03/02/12

1.000

0.693

0.500

03/02/13

0.805

1.116

0.398

03/02/14

0.865

0.950

0.440

03/02/15

0.811

0.562

0.625

03/02/16

0.895

1.241

0.322

03/02/17

0.937

1.030

0.380

03/02/18

0.819

0.900

0.469

03/02/19

0.928

1.494

0.250

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Figure 16. Scatter plots showing the diversity and evenness indices depending on the different habitat types. 1 refers to neritic, 2 to epipelagic and 3 to mesopelagic tows.

Figure 17. Whisker plots of the Pielou’s Evenness Index J’, Shannon Index H’, Simpson’s Diversity Index, number of total species S and number of total individuals N. All are in dependence of the habitats (1: neritic; 2: epipelagic; 3: mesopelagic). Means are shown and confidence interval is given with +/- 95 (in Total Individuals N the Standard Error is given).

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To test the significance of the differences of the Shannon H’, Pielou’s J’, Simpson’s D, number of total Species S and the number of total individuals N among the grouped stations after the factors of season, diurnal changes, vertical Distribution, horizontal Distribution an ANOVA was performed. Results are illustrated in Whisker Plots, which are summarized in Figure 17 to Figure 21. In general the confidence intervals overlap indicating no differences between the three habitat types (Figure 17). Only between the Evenness in the neritic zone and the mesopelagic zone is a significant difference, confirmed by the ANOVA results (J’: p < 0.01). Also the Bonferroni test confirms a significant difference between the habitat types 1 and 3 (p = 0.01292). The trend of an increasing Evenness from neritic to the epipelagic to the mesopelagic habitat can be recognized. The low Pielou’s J’ in the neritic realm indicates a dominance of some species, especially of Engraulidae and Gobiidae. This is also reflected in the lowest diversity index (Shannon and Simpson’s Index), though the total species number is highest. In the epiand mesopelagic realm the individuals are evenly distributed among the species, though the diversity (H’, D and S) is higher in the epipelagic habitat than in the mesopelagic zone. But in general the different habitats are not affecting the diversity significantly (H’,D and S: p > 0.05).

Figure 18. Whisker plots of the Pielou’s Evenness Index J’, Shannon Index H’, Simpson’s Diversity Index, number of total species S and number of total individuals N. All are in dependence of the months (1: January; 3: February; 4: April; 5: May; 11: November). Means are shown and confidence interval is given with +/- 95 (in Total Individuals N the Standard Error is given).

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The number of larval individuals shows a very high range in the neritic realm (Figure 17).This is caused by some single stations with an extremely high amount of Engraulidae and Gobiidae. In the mesopelagic realm the number of individuals and species is lowest and evenly distributed, reflected in the relatively high evenness. ANOVA shows no significant difference (all p > 0.05) between the months. The season is not influencing the indices. Only some trends can be seen. Evenness stagnates between January and May and decreases in November (dominance effect). In Figure 18 the diversity (Shannon and Simpson’s Index) and species number increases towards May, though the diversity is low again in November, while the species number is comparable with May. The Simpson’s Index is the same in March and April. The number of individuals is exceptionally high in November. The depth is influencing the Evenness and the number of total species significantly (p < 0.05). The difference is significant between the depth ranges of 0 - 150 m and 150 – 600 m. All other Indices do not differ significantly (p> 0.05). In Figure 19 the diversity H’, D and the number of species are highest within the surface waters, although D shows smaller differences and the number of species shows almost no difference between the two deepest layers. All are decreasing with depth, while Evenness shows a more or less differing pattern. The number of individuals is extraordinary high in the surface waters, what is mainly caused by the high amount of Engraulidae and Gobiidae especially in the neritic realm.

Figure 19. Whisker plots of the Pielou’s Evenness Index J’, Shannon Index H’, Simpson’s Diversity Index, number of total species S and number of total individuals N. All are in dependence of the depths (1: 0 – 150 m/SF; 2: 150 – 600 m/NACW; 3: > 600 m/AAIW). Means are shown and confidence interval is given with +/- 95 (in Total Individuals N the Standard Error is given).

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Figure 20. Whisker plots of the Pielou’s Evenness Index J’, Shannon Index H’, Simpson’s Diversity Index, number of total species S and number of total individuals N. All are in dependence of the Islands (1: Lanzarote; 2: Fuerteventura; 3: Gran Canaria; 4: Tenerife and La Gomera; 5: high oceanic in La Bocaina; 6: high oceanic far north off Gran Canaria; 7: high oceanic far north off Tenerife). Means are shown and confidence interval is given with +/- 95 (in Total Individuals N the Standard Error is given).

The region has no significant influence on the indices (all p > 0.05). In Figure 20 the diversity (Shannon and Simpson’s Index) and the number of species is decreasing from the most eastern islands towards the west with open ocean condition. Around Fuerteventura the diversities are highest. Within the oceanic stations far off the islands the number of species do not show remarkable chances, while the diversity (Shannon and Simpson’s Index) is highest in La Bocaina. But all diversities far off the islands are higher or as high as stations around the islands. The individual number is highest around the eastern most islands. The diversity (Shannon and Simpson’s Index) and the total number of species are higher in trawls conducted during night, but almost invariant during the daytime (Figure 21). The number of individuals is clearly higher during day, what is caused by the single stations with a high number of Engraulidae and Gobiidae.

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Figure 21. Whisker plots of the Pielou’s Evenness Index J’, Shannon Index H’, Simpson’s Diversity Index, number of total species S and number of total individuals N. All are in dependence of the daytime (1: during day; 2: during night). Means are shown and confidence interval is given with +/- 95 (in Total Individuals N the Standard Error is given).

3.5. Biogeography In Figure 22 – Figure 25 horizontal Distribution and abundance of larvae is shown. The larvae are arranged due to their origin and spawning habitat as noted in Table 7. The habitats of origin are summarized as neritic, oceanic (epi – to deep pelagic), neritic to bathydemersal species and benthopelagic species. All cruises (La Bocaina 11/97, ECOS 04/99, Mesopelagic 05/99, Pelagic 01/00, Pelagic 11/00 and La Bocaina 03/02) are combined in the illustrations. The different colours and sizes of the circles indicate the numbers of larvae, which are listed in the legend within the maps. The isobaths are labelled, so that the horizontal Distribution is comparable with the bottom depths as well. The occurrence of fish larvae is clearly not restricted to the neritic habitat (Figure 22). A neritic – oceanic exchange takes place. A high abundance of neritic fish larvae occurs within the 100 m isobath or close to it. But many small groups of neritic larvae occur deeper than the 500 m isobath, especially southwest of Gran Canary and southeast of Fuerteventura. Some neritic larvae were found at oceanic stations with a bottom depth of about 2000 m, north of Gran Canaria and east of Fuerteventura.

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Figure 22. Larval distribution of neritic species. Numbers of larvae are given in the legend. Grey isobaths are marked with the bottom depth (map modified after WIENERROITHER 2005 with MapInfo Professional 7.5).

Figure 23. Larval distribution of oceanic (epi- and deep pelagic) species. Numbers of larvae are given in the legend. Grey isobaths are marked with the bottom depth (map modified after WIENERROITHER 2005 with MapInfo Professional 7.5).

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Also the oceanic fish larvae show an accumulation in the southeast of Gran Canaria and southwest of Fuerteventura (Figure 23). Especially in the latter area a neritic – oceanic exchange takes place. Along the flanks of this island, as well as north of Fuerteventura, oceanic species can be seen at neritic stations.

Figure 24. Larval distribution of species being neritic as well as bathydemersal. Numbers of larvae are given in the legend. Grey isobaths are marked with the bottom depth (map modified after WIENERROITHER 2005 with MapInfo Professional 7.5).

Figure 25. Larval distribution of benthopelagic species. Numbers of larvae are given in the legend. Grey isobaths are marked with the bottom depth (map modified after WIENERROITHER 2005 with MapInfo Professional 7.5).

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Similar to species that were ecologically characterized as neritic to bathydemersal (Table 7, Figure 24), the fish larvae from benthopelagic species were also encountered in larger concentrations southeast of Fuerteventura (Figure 25). Around Gran Canaria fish larvae from these ecological groups were also encountered close to coast. Benthopelagic species occur mainly above bottom depths of 1000 to 2000 m (Figure 25).

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4 DISCUSSION

4.1. SAMPLING METHOD Traditional vertical plankton sampling techniques allow to catch also considerable quantitites of fish larvae during hauling-in of the net.This method causes considerable bias regarding vertical distribution patterns (SASSA et al. 2002). LEIS (1986) used the same method with a collapsing net, however in more recent years other more elaborated methods have been adopted. For instance, JOHN et al. (2001, 2004a, 2004b) used a multiple opening-closing net (multinet) for their investigations. When trawling nets are used (JOHN et al. 2001; LOBEL and ROBINSON 1988; LOGERWELL and SMITH 2001; PALOMERA et al. 1988; RODRÍGUEZ et al. 1999; SABATÉS and MASÓ 1992) mainly horizontal and virtually no vertical resolution of the distribution is obtained. JOHN et al. (2001) found differences in the catch between neuston nets and multinets, especially with diurnal catches. While the multinet net does not reveal diurnal changes, the neuston net does, due to its higher visibility to the larvae. Another bias arises from the relative coarse mesh size. A mesh size of 2 mm in the cod end selects only larger larvae and is inappropriate for obtaining smaller ones. Also, larger slim larvae might escape through the wide meshes. Paralepidids and Cyclothone sp. can easily escape and are therefore underrepresented, although they are common in adult stages around the Islands (WIENERROITHER 2003 and 2005). Meticulous larval investigations use nets with a mesh size between 200 and 500 µm (ACEVES-MEDINA et al. 2004; BÉCOGNÉE 2006; JOHN et al. 2001, 2004a and 2004b; LEIS 1986; LOBEL and ROBINSON 1988; LOGERWELL and SMITH 2001; PALOMERA et al. 1988; RODRÍGUEZ et al. 1999; SABATÉS and MASÓ 1992; SASSA et al. 2002, 2004). A further problem lies in the non-identical realisation of the trawls. Having no identical replicates leads to statistical problems. In addition, overlapping sampling depths cause a bias in the statistical analyses. The variable ranges of depth and durations make it difficult to give exact details about the occurrence of different species (WIENERROITHER 2003). Moreover, in respect to the horizontal distribution of sampling stations a bias lies in the accumulation of stations in some areas, e.g., southeast of Fuerteventura. The non-randomly and non-evenly distribution of the stations within an area might also lead to a problem in the application and interpretation of statistical analyses. Further, the analyses of horizontal distribution might reflect the sampling pattern and not the real in-situ small scale distribution of fish larvae within the entire area of investigation.

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4.2. SPECIES COMPOSITION 4.2.1. Comparison with earlier work The taxonomic composition of fish larvae matches quite well with the data given by RODRÍGUEZ et al. (1999), where the community was also dominated by neritic larvae (mainly clupeiforms), though with a lower percentage. Contrary, observations by RODRÍGUEZ et al. (2001) show an inverse pattern. A proportion of about 85% of the total number of fish larvae were oceanic, while only 11% were neritic, although the area of investigations were relatively close to Grand Canaria, and not further offshore than in other studies (RODRÍGUEZ et al. 1999; JOHN et al. 2004a).The lack of neritic larvae can be explained by the simultaneously observed hydrographic situation.The filament and the island induced eddy off Gran Canaria, which both count as retention area of neritic species (RODRIGÍUEZ et al. 2001), are bent southward seeming to spin off and not reach the island. One mechanism related to the upwelling filament is the transportation of the European anchovy, Engraulis encrasicolus from the West African coast to the islands. This species has the highest individual number of larvae in the present study. Larvae of E. encrasicolus are known to be present between December and March at Gran Canaria (BÉCOGNÉE et al. 2006), though around the Canary Islands E. encrasicolus are mentioned and confirmed as summer spawners (JOHN et al. 2004a; RODRÍGUEZ et al. 1999). The results of the present study give a high number of E. encrasicolus larvae in November 1997, when upwelling was weak. RODRÍGUEZ et al. (1999) calculated the age of European anchovy larvae around the Canary Islands with 1.2 days and 38.3 days at 4 mm and 19.2 mm, respectively, and a spawning origin north of Cape Yubi. The larvae in the present study tend to be around the larger size, so their spawning might occur around end of September or beginning of October, where the upwelling and filaments might be higher developed. Admittedly, no single European anchovy larva was caught in January 2000 and November 2000. During the entire period of investigation the high proportion of engraulid larvae among cluipeiform fish is unusual, since larvae of Sardinella aurita VALENCIENNES, 1847 have been reported as being predominant (BÉCOGNÉE et al. 2006; RODRÍGUEZ et al. 2001). In addition, the absence of European pilchard, Sardina pilchardus (WALBAUM, 1792) is uncommon in that region (BÉCOGNÉE et al. 2006; RODRÍGUEZ et al. 2001), but also JOHN et al. (2004a) did not find S. pilchardus larvae neither in winter, spring (March) or summer, a fact he thought to be noteworthy. RODRÍGUEZ et al. (1999) found out, that the European anchovy larvae were only associated with the northern area of the filament, with a peak in abundance at about 27.5°N and14.0°W, while the European pilchard larvae were concentrated at 27.0 – 27.5°N and 15.5°W. During our investigation, only the northern area of the filament meeting the islands of Gran Canaria and Fuerteventura was investigated and provided a high number of E. encrasicolus. Even at more northern stations larvae were found, following the pattern observed by RODRÍGUEZ et al. (1999). He suggested E. encrasicolus spawn along the western African coast, the eggs are transported by a filament to the Canary Islands and larvae development occurs during the transportation. Besides the general selection towards late stage larvae due to the coarse mesh size, this explains the catch of exclusively postflexion larvae. Within the Engraulis encrasicolus larvae one thing could be observed. The irises of the European anchovy larvae were sometimes silvery and sometimes black coloured. 99.97 % of the larvae caught during the daytime had silvery irises, whereas only one specimen (0.03 %) had black irises. This single specimen with dark eyes occurred at a mesopelagic station, but most likely remained in the net from the previous tow (contamination). This is indicated by the dark eyes since the previously undertaken neritic haul was done during the night and the anchovies captured during the night had black irises to an extent

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of 92.11 % and only 7.89 % of the larvae had silvery irises. In hauls taken during the dawn, some larvae had silvery and some dark irises. Anyway, the change in the coloration of the iris to a dark iris might indicate an adaptation to avoid being seen by predators during the night.The moon light could be reflected by the silvery irises and hence the pelagic larvae could be easily detected by potential predators. Probably therefore, the coloration of the iris changes into black during the night. Not the highest individual numbers, but the highest diversity was found among the myctophids. The most common genera were Notoscopelus, Hygophum and Ceratoscopelus, listed in decreasing order. Also in the multi-seasonal study of JOHN et al. (2004a) Notoscopelus was the overall most common genus, followed by Ceratoscopelus. Incongruently the third and fourth frequent myctophid group were the genus Lobianchia and the species Diogenichthys atlanticus, whereas Lobianchia was not present in our studies at all and D. atlanticus only with a few specimens. Our third most common genus was therefore only the fifth frequent one in their study. The study of RODRÍGUEZ et al. (1999) differs quite much from these patterns. He found the genus Lampanyctus the most common one followed by Diaphus, Diogenichthys, Ceratoscopelus and Hygophum. Notoscopelus was only present with a small portion of specimens. This might depend on the sampling season which was in summer (August 1993). During this time of the year JOHN et al. (2004a) could not record a high number of this genus either. Therefore, a low number of Notoscopelus larvae during summer might be a common pattern in the Canary Islands area. Notoscopelus was only identified to genus level in JOHN et al. (2004a) and RODRÍGUEZ et al. (1999). In the present study Notoscopelus resplendens was found to be most common as it was also the case for the adult representatives of the genus (WIENERROITHER 2003) collected in the same cruises. JOHN et al. (2004a) note the absence of the genus Ceratoscopelus in winter 1997. However, in the present study specimens of Ceratoscopelus warmingii were found in November 1997, while Ceratoscopelus maderensis was absent. In contrast, C. maderensis was found in winter 2000 though in a low number, while C. warmingii was completely absent then. In May 1999 C. maderensis showed the highest individual numbers, but no specimen of C. warmingii occurred. JOHN et al. (2004a) found only one single specimen of C. warmingii, but a peak in C. maderensis abundance during summer 1998. RODRÍGUEZ et al. (1999) found a similar pattern in summer (August 1993) with C. maderensis being more abundant, while C. warmingii occurred at lower numbers. Hence, the two Ceratoscopelus species seem to have opposing seasonal abundance patterns and a more or less exclusionary occurrence. While C. maderensis occurs mainly in summer, C. warmingii seems to be most abundant during the winter season. Concerning the vertical distribution, JOHN et al. (2001) mentioned Ceratoscopelus. warmingii as being most abundant in the upper 25 m, while fewer specimens were caught down to 50 m in the Angola-Benguela frontal zone. The present study confirms this vertical distribution pattern also in the region of the Canary Islands. Only one specimen derives from a deep tow at around 500 m depth northeast of Lanzarote. The specimen may however have been collected near the surface during the final hauling-in phase. The highest number of Diogenichthys was found in March.This fits perfectly to the pattern found by JOHN et al. (2004a) who reported the highest catch of D. atlanticus also in March. Noteworthy is the absence of the sternoptychid hatchet-fish genera in the present study, which were found in depths below 400 m (JOHN et al. 2004a; RODRÍGUEZ et al. 1999) around the Canary Islands. Sternoptyx HERMAN, 1781 as well as Argyropelecus COCCO, 1829 seem to have their spawning season in summer. However, many sternoptychid larvae were not identified to lower than family level (JOHN et al. 2004a; RODRÍGUEZ et al. 1999) and occurred all year round with a peak in March.

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Another sternoptychid species, Maurolicus muelleri, is known to spawn over the shelf and slope, however the only specimen in our study was found far north from Gran Canaria where the bottom depth is more than 1300 m. This may indicate a drift by a northern filament from the West African coast into the open ocean. The seasonal occurrence of M. muelleri was incongruent with previous studies (JOHN et al. 2004a). These authors observed the highest catches during winter season, while no larva could be found during winter cruises in the present study, but only during May. The occurrence of Vinciguerria larvae is concentrated from the surface water down to 50 m, but when they start to transform they migrate vertically down to 300 – 400 m (BADCOCK 1994). In the present study most larvae were found in the upper meters, but since larvae were in post flexion stage and some already in the transformation stage, several specimens were found deeper, even down to 500 m already in the process of migrating down to their adult habitat. RODRÍGUEZ et al. (1999) found V. poweriae to be the most common species among the genus Vinciguerria during August 1993, while during our investigations V. attenuata was the most abundant species with a peak in May. V. attenuata was mainly found at oceanic stations far north of the islands, while V. nimbaria seem to be more associated to the islands and was intermediate in its occurrence with the highest abundance in autumn. V. poweriae was rare throughout the period of investigation. In JOHN et al. (2004a) V. nimbaria was most abundant throughout all seasons, except for autumn, which was not included in their investigations. This species had its peak during March. Intermediate in its occurrence was V. attenuata, but not found in winter and V. poweriae was lowest in abundance and not found in March. The overall tendency that can be seen is that V. poweriae is most abundant in summer (JOHN et al. 2004a; RODRÍGUEZ et al. 1999), which is congruent with BADCOCK (1984). In our study V. nimbaria has its peak in autumn, but no further information about this season can be found in other studies. In the study of JOHN et al. (2004a) V. nimbaria was most common in March. V. attenuata is most common in May according to the present study, and in summer according to the data from JOHN et al. (2004a).

4.2.2. Comparison with adult fish WIENERROITHER (2003 and 2005) investigated adult fish material caught in the same surveys the larval material of this study derives from. Concerning the adult fish fauna WIENERROITHER (2003) observed mesopelagic species from the cruise La Bocaina 11/97 and ECOS 04/99, where the families Myctophidae, Gonostomatidae, Phosichthyidae and Stomiidae made up about 95 % of the total catch around the Canary Islands, although trawls in neritic, epipelagic and mesopelagic realm were taken. Concerning only mesopelagic fish larvae, Gonostomatidae were the most abundant next to Myctophidae, Phosichthyidae and Paralepididae. The most diverse family within the larval community are the Myctophidae, which is similar to the findings of WIENERROITHER (2003 and 2005). Within the adult Myctophidae Hygophum hygomii was the most abundant species in all seasons (WIENERROITHER 2005), while only a few larval specimens were caught. Ceratoscopelus warmingii and Lobianchia dofleini (ZUGMAYER, 1911) were very abundant in the adult fish material (WIENERROITHER 2005). In the larval material some specimens of larval C. warmingii occurred, but no larval L. dofleini. Within the adult Gonostomatidae Cyclothone sp. was the most dominant genus (WIENERROITHER 2005), while only a few Cyclothone larvae were found in identical trawls. This result may at least in part reflect a general underprepresentation of small larvae due to the methodology as discussed before. Adult Gonostoma elongatum GÜNTHER, 1878 was more abundant than G. denudatum (WIENERROITHER 2005), while G. denudatum was abundant in larval material, no larval G. elongatum was found at all. Congruent with WIENERROITHER (2005), the paralepidid Lestidiops sp. was most abundant in autumn season in larval and adult study. Further, none of the demersal indicator species found by LORANCE et al. (2001) was represented in the larval material. Overall, the larval community sampled seems not to reflect the species composition of the adult fish assemblage. Only a small portion of the adult fish

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species lists of BRITO et al. (2002), Uiblein et al. (1996, 1998), LORANCE et al. (2001), and WIENERROITHER (2003 and 2005) is represented in the larval list. More sampling efforts using more appropriate techniques would certainly allow collecting a higher number of fish larvae taxa and also to possibly obtain more new records from this still underinvestigated area (WIENERROITHER 2005).

4.2.3. New records WIENERROITHER (2005) noted that the current number of fish species in the area of the Canary Islands still may further increase, if the pelagic sampling is continued. Also, previous surveys have not fully covered all habitats as well as the larval stages. One origin of enrichment for the area of investigation is most likely the passive drift of larvae. This can be supported with the new records of WIENERROITHER (2005), as individuals of several fish were near adolescence or even at the juvenile stage. This led him to propose larval transportation via water masses into that area, which is lso supported by the investigations of RODRÍGUEZ et al. (1999). Further, a northward transport of tropical species by the AAIW and also NACW reverse current are one source for new species around the Canary Islands (JOHN et al. 2004a). The Canaries have a subtropical littoral fish fauna, which is related to the one in the Atlantic-Mediterranean biogeographic province (BRITO et al. 2006). This fish fauna has a very low or even non-existent rate of endemism in each archipelago, because currents redistribute species between the islands (BRITO et al. 2006). Genetic studies showed that fish populations of the Azores are more separated from the other Islands (Madeira and Canary Islands), while Madeira and the Canaries are more related to each other and to the European mainland (Portugal and England), which leads to the suggestion of a southward colonization (DOMINGUES et al. 2007). Within the archipelagos of the Canaries, Madeira and Azores the migrations of neritic species show a northwestern trend (DOMINGUES et al. 2006). From the western coast of Portugal to the archipelagos of Madeira and Canaries a southward migration exists due to the transport by the Canary current (DOMINGUES et al. 2007). This southward transport is strongest in spring, summer and winter in the upper 200 m (JOHN et al. 2004a). On the other hand, 80 % of the new records of the littoral fish fauna during the nineties and the first years of the present decade were tropical (BRITO et al. 2005), a fact also observed by WIENERROITHER (2003). JOHN et al. (2004a) have concluded from own and historical data, that species intruded the Canary Islands from the northwards flow of NACW and SW in autumn. JOHN et al. (2004a) mentioned that northward currents add 4 to 8 tropical specimens per squaremeter to the native oceanic fauna. It is also thought that filaments can contribute neritic larvae from the African shelf to the local neritic populations of the Canary Islands (RODRÍGUEZ et al. 1999). The gadiform species Melanonus zugmayeri is distributed worldwide in tropical to temperate seas and inhabits the mesoand bathypelagic realm (COHEN 1984). Within the northeastern Atlantic M. zugmayeri is most common around the Azores, Madeira and west of the Canary Islands. So it is not astonishing to encounter this species around the islands, since interconnections due to migration between these islands (DOMINGUES et al. 2006 and 2007) and a southward transport within the season of spring (JOHN et al. 2004a) are common. Argentina sphyraena has been also recorded along the coast of Europe and Mediterranean Sea and along the coast of West Africa and inhabits the shelves and slopes (COHEN 1984). Since larvae were found around the western islands, which are most influenced by the African upwelling, it is most likely, that the larvae were entrained by an upwelling filament from the Marroccanean coast. The temperate Crystallogobius linearis inhabits the coastal habitat as well as the pelagic realm down to a depth of 400 m along the Atlantic European coast and the Mediterranean Sea (MILLER 1984). The Canary Islands are the southernmost record

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of C. linearis in the North Eastern Atlantic. Also the temperate species Entelurus aequoreus is common along the European coast and around the Azores. All specimens of both temperate species were caught during autumn. A southward transport must have occurred, though the autumn season is known as a time of northward transport of all water masses (JOHN et al. 2004a; KNOLL et al. 2002). This can be explained by JOHN et al. (2004a) since no AAIW northward flow was present in autumn 1997 and the SW and NACW had a clear southward geostrophic net transport during autumn 1997, when those specimens occurred. A southward transport is also most likely since C. linearis was recorded around Madeira for the first time already in 1995 (WIRTZ et al. 2007). TORTONESE (1984) noted that Microichthys coccoi is only known from the strait of Messina, but he pointed out, that some young Epigonus telescopus (RISSO, 1810). from the North Atlantic could have been confused with M. coccoi. So, it might be that M. coccoi is more common in the North Atlantic, than has been observed up to now. If M. coccoi really would occur only in the Mediterranean Sea, a passive drift out of the Mediterranean Sea by outflowing water is most likely.

4.3. Occurrence and distribution patterns 4.3.1.Vertical spatial occurrence and distribution RODRÍGUEZ et al. (2006) noted that the vertical distribution does not depend on the thermocline and the associated phytoplankton peak, but is more species-specific and related to other environmental conditions. Regarding the depth distribution of different communities the common separation of realms into neritic, epi- and mesopelagic is not sufficient. Nevertheless, there is a trend for the communities of the epipelagic zone to consist of neritic as well as mesopelagic larvae. At this stratum an exchange between the neritic and oceanic species takes place. For that reason, the diversity (Shannon and Simpson’s Index) and number of species are highest in the upper 150 m (although the neritic realm is included here) and decreases depending on the water masses in the present study, although a fine scale study on the vertical distribution of larvae by RODRÍGUEZ et al. (2006) showed, that larvae do not occur directly in the surface stratum but within 20 – 66 m depth, but the diversity was highest in 50 – 66 m depth. The species number is also highest in the most upper layer of the water collumn. This is because larvae of all oceanic species favour this stratum, especially the most divers family of myctophids. They mainly inhabit the upper 100 m and no specimens were found deeper than 150 m (SASSA et al. 2004). Allowedly, this relation is poor since no simultaneous measurements of the hydrographic situation were done and stratum information were taken from the literature. Anyway, the depth and hydrographic situation e.g. water masses should be considered and related to the fish larval distribution in future studies. Within the neritic realm, a dominance effect occurs because of the high number of Gobiidae and Engraulidae. Evenness differs significantly between the neritic and the mesopelagic realm. In the mesopelagic realm few species and the lowest numbers of individuals were found, but they were evenly distributed. Though most species inhabit the neritic and epipelagic realms, the evenness of the surface waters including the neritic realm is significantly lower compared to deeper layers, which is also caused by the engraulids and gobies. Anyway, the diversity (Shannon and Simpson’s Index) is highest in the epipelagic zone, since no dominance effect occurs and the community consists of neritic as well as oceanic species. The region, where this exchange takes place is in the Islands’ wakes and eddies as clearly seen in Figure 22 – Figure 23. Looking at the diversity (Shannon and Simpson’s Index) and especially at the species number around the single Islands, the main exchange happens southeast of Fuerteventura. Here, in the epipelagic realm close to the slope, productivity seems to be the highes among all areas of the archipelago which have been investigated so far.

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4.3.2. Horizontal spatial occurrence and distribution The African upwelling system seems to influence the larval community. A change in community is observed from the east to west congruently with the strength of the upwelling’s influence. This trend exists even at the highly oceanic stations. The more oceanic conditions prevail, the more distinct the communities become. The influence from the African coast becomes significant on the eastern edge of the Archipelago and stress was laid on the investigation of the eastern community, so only a few samplings were conducted around the western islands. Unfortunately no east-west comparisons are possible, but as the western islands are not influenced by the African upwelling, a significant difference in the community assemblage can be expected. The diversity (Shannon and Simpson’s Index) shows an decreasing trend from the East (Lanzarote and Fuerteventura) to the West. Within this trend Fuerteventura has the highest diversity, what is caused by the highly productive area southeast of the island as discussed in the section before. Even the highly oceanic stations show this trend. This reflects the strength of the influence by the Northwest African upwelling. Notable, the highly oceanic stations stations closest to the upwelling (N off Fuerteventura) show a particularly high diversity (Shannon and Simpson’s Index). This is an area only favored by oceanic species (Figure 23) and represents a highly productive area for the oceanic community. The most important area of neritic larvae within the oceanic realm seems to be the southwest and southeast of Gran Canaria and Fuerteventura, respectively in the present study. In this area most neritic larvae within the oceanic zone were found and Engraulis encrasicolus as well as Gobiidae make up the main portion of those neritic species. Within these areas two main mesoscale hydrographic features happens, the island induced wakes and eddies (RODRÍGUEZ et al. 2006). Especially downstream southwest of Gran Canaria the generation of warm wakes and eddies are known to be common (RODRÍGUEZ et al. 2001). In addition, this is also the collision zone between the filament originating from the West African coast and the island itself and its generated eddy, respectively. In this area an increased abundance of fish larvae in the oceanic zone can be recognized by BÉCOGNÉE et al. (2006), RODRÍGUEZ et al. (2001) and in the present study. Also LEIS (1986) found the highest accumulation and diversity of larvae downstream of oceanic islands. Especially eddies provide a retention and nursery area for fish larvae. The trajectory model showed that particles released north of the island were transported into the cyclonic eddy south of Gran Canaria, where they remain close to the island for several weeks (RODRÍGUEZ et al. 2001) and might be delivered to the island again when the larvae are old enough. For example eddies in the Florida current system are clearly associated with the delivery of neritic larvae to the coastal zone and larvae can even exceed their larval phase until they are transported back (SPONAUGLE et al. 2005). But usually eddies maintain a sufficient duration enabling larvae to complete their pelagic phase and return to parental habitats (LOBEL and ROBINSON 1988). Being trapped within an eddy can be beneficial for larvae, since eddies produce a high number of survivors due to better feeding conditions, while the neritic realm educe only a moderate number of larvae, and within an eddy the mortality rate is even lower than in surrounding waters (LOGERWELL and SMITH 2001). Many fish larvae are trapped or even seem to prefer being in an eddy. E.g. the pacific sardine, Sardinops sagax (JENYNS, 1842) were associated with mesoscale eddies within the oceanic realm (LOGERWELL and SMITH 2001). Also around Hawaii the densities larval fish were higher within an eddy than within the surrounding water (LOBEL and ROBINSON 1988). The number of larvae within an eddy was comparable with the amount found associated with a reef, but the proportion of mesopelagic, epipelagic and neritic larvae were approximately the same (LOBEL and ROBINSON 1988). However, eddies may also have an adverse affect on neritic recruitment, if they move too far away from the coastal habitat, although transformed fish can swim at least one to several kilometres to migrate back to coastal habitats (LOBEL and ROBINSON 1988).

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But not only neritic larvae accumulate in the southwest of Gran Canaria and especially southeast of Fuerteventura, also the number of oceanic species is high there and they occur even in the neritic realm. Also JOHN et al. (2004b) observed a mixing of oceanic larvae with neritic species onshore due to an onshore transport by an anitcyclonic eddy. Further, few oceanic specimens appear at neritic sites along the flanks of Gran Canaria and at the stagnation point of Fuerteventura and Lanzarote, where the current divides and flow is minimal. This can be explained by the trajectory model of RODRÍGUEZ et al. (2001). This study shows that particles released north of Gran Canaria within the oceanic realm may beach along the flanks (RODRÍGUEZ et al. 2001). Further onshore currents e.g. due to the local upwelling at the western sides of the islands (Figure 1b-c), might transport oceanic larvae into the neritic zone. The north of the islands is known to accumulate neritic larvae (RODRÍGUEZ et al. 2001), especially species with non-pelagic eggs like pomacentrids (LEIS 1986). The hydrographic feature, which is helping the larvae remain close to the islands, is the stagnation point. This is the area where the current divides and no flow is noticeable. A high concentration of neritic larvae and fish eggs north of Gran Canaria in the study of RODRÍGUEZ et al. (2001) shows the stagnation point being a favoured nursery ground for larvae. Also in the present study a high abundance of neritic larvae is found at the stagnation point of the island complex Fuerteventura and Lanzarote. Furthermore, on the eastern and western flanks of Gran Canaria the abundance of larvae was relatively high. But too few stations were taken to get an accurate picture of the distribution there, as well as in the north of Gran Canaria. Usually the flanks of an island are not areas with high larval fish abundance, as a previous trajectory model showed (RODRÍGUEZ et al. 2001). At the flanks of Gran Canaria, especially in the channel between Gran Canaria and Tenerife, a low abundance was found, which is also congruent with the low mesozooplanktonic biomass there (RODRÍGUEZ et al. 2001). The reason for the low abundance is the flank currents, which advect the plankton and prevent it from accumulating (RODRÍGUEZ et al. 2001). The trajectory model also showed particles, which were released along the flanks and transported out of the area, so flanks do not seem to be a preferred spawning area. Due to the very complex hydrographic situation around the Canary Islands with local upwellings and other hydrographic events (e.g. southeast of Fuerteventura), the overall larval distribution based on the hitherto available samples gives no clear picture or trend. A more fine-scale spatial investigation of planktonic communities and ichthyoplankton would be required for an enhanced interpretation in terms of the local physical characteristics (ARÍSTEGUI et al. 1997; HERNÁNDEZ-LÉON 1991; RODRÍGUEZ et al. 2001; SABATES and MASO 1992) and to tackle the question if anomalous local or mesoscale hydrographic events are reflected in changes in larval distribution (SABATES and MASO 1992).

4.3.3. Seasonal occurrence and distribution JOHN et al. (2004b) noted that the offshore spreading of neritic larvae is more an effect of hydrographic variabilities due to local upwellings than a seasonal effect (JOHN et al. 2004b). But the intensity of the upwellings and current strength is seasonally dependent. E.g. the upwelling is lowest in late autumn (Figure 1a and e). Consequently, the spreading of larvae is also lowest then. This is supported by the present study, since most neritic larvae caught in autumn were within the neritic realm. So the differences in spatial distribution are also seasonally caused. No significant seasonal differences in community structure were encountered in this study. Nevertheless, some trends in larval occurrence and distribution depending on the time of the year could be observed: it seems that the months of January and November represent two spawning events for different communities. Although there might be a further spawning event in May attention has to be paid to the area of investigation. During the May cruise only highly oceanic locations were chosen,

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which resulted in the catch of mainly oceanic species (Figure 23). Consequently the difference in community structure might originate in the spatial distribution of stations and not in the seasonal variation. The samples collected in May seem to be more similar to the community found in November, which also included more oceanic stations and oceanic conditions prevailed, since upwellings were low then (Figure 1a and e). The stations in March were within the sheltered area southeast of the island Fuerteventura, where the eddy spins of (RODRÍGUEZ et al. 2001). This may reflect more the hydrographic situation than a seasonal effect. The low evenness and high number of individuals in November is caused by the high proportion of Engraulidae and Gobiidae. Anyway, diversity (Shannon and Simpson’s Index) and the species number did not differ significantly between the seasons, but both tend to increase towards summer, when the upwelling along the African coast and the late winter bloom (LWB) become more intense, which can be followed in Figure 1. The upwelling and the late winter bloom increases the diversity and productivity in that area and this increased productivity might be preferred by many species for spawning to provide better conditions for the development of the larvae.

4.3.4. Diurnal occurrence and distribution Diurnal analyses showed small differences in community structure, attributable to differences in the appearance of Engraulidae and Gobiidae which where caught mainly during the day. LEIS (1986) observed that gobies are shallower during the day. In addition, more species are caught during the night, which indicates a net avoidance during the day. RODRÍGUEZ et al. (2006) found larvae to be denser in every stratum during the night, which might also reflect dial migration or net avoidance during day. They found no significant differences and suggested that if some species migrate, other species migrate opposing, so that there is no difference in the overall larval population. Also SASSA et al. (2004) found no diurnal vertical migration within the larval myctophids. And since the differences are not significant, it can be assumed that dial patterns do not affect the results of the present study.

4.4. Conclusion The fish larvae material investigated derives from rather unselective sampling methods which were primarily aimed at collecting adult pelagic fishes. However, the by-catch from the trawl tows resulted in a number and diversity of fish larvae sufficiently high for conducting a detailed taxonomic and ecological study of this important ecosystem component. Among the total 5088 individuals of mostly late-stage fish larvae from 16 orders, 54 families, and 70 species, five new records consisting of temperate as well as tropical species could be documented for the Canary Islands. Larval Engraulidae and Gobiidae dominated most of the samples. My investigations show that the overall diversity (Shannon and Simpson’s Index) increases towards summer, which is congruent with an increasing influence of the African upwelling. The densest fish larvae accumulations were encountered within the islands’ wakes and eddies. Especially southeast of Fuerteventura, in a hydrographically highly active area, oceanic and neritic larvae were concentrated and co-occurred at open-ocean as well as at coastal sampling stations. In an open-ocean area off northern Fuerteventura the highest diversity of oceanic species was encountered.These results strongly suggest that both largeand small- scale hydrographic processes and structures arising from surface and deep currents, upwelling, and the islands’ wakes and eddies clearly influence the local and seasonal distribution and abundance of fish larvae in the area of the Canary archipelago. In order to fully consider the significance of fish larvae as indicators of physical as well as biological conditions and changes in a wide array of habitats of a marine biodiversity hotspot like the Canary Islands, a methodologically more advanced, systematic monitoring program would need to be established.

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5 ACKNOWLEDGEMENTS

First of all I thank Franz Uiblein from IMR (Institute of Marine Research), Bergen, who gave supervision and supported me during my thesis. Further, I am grateful to Fernando Bordes from ICCM (Instituto Canario de Ciencias Marinas), Telde, for his great work as cruise leader and the translation of the abstract. I also thank Ulrike-Gabriele Berninger from the University of Salzburg, Department of Organismic Biology for her moral support and words of encouragement. I am grateful to Peter Rask Møller from the ZMUC (Zoological Museum and University Copenhagen) for providing the fish larval material. For introducing me into the skill of fish larval identification I am very grateful and thank Hans-Christian John from the ZIM (Zoological Institute and Museum), Hamburg. I also thank Martin Wahl and Mark Lenz both from the IFM-Geomar, Kiel, for their help in the statistical analyses. I am also grateful to Jørgen Nielsen (ZMUC) and Rupert Wienerroither (University of Salzburg, IMR) for valuable discussions.

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7 APPENDIX

7.1. OVERVIEW TABLE OF ALL SPECIES ENCOUNTERED Table 9 List of numbers of individuals of each species caught during all six cruises “Bocaina 11/97”, “ECOS 04/99”, “Mesopelagic 05/99”, “Pelagic 01/00”, “Pelagic 11/00” and “La Bocaina 03/02”. Individual numbers are given per cruise, systematic order according to NELSON (2006). Larvae illustrated below in a picture are written in bold letters. Order

Suborder

Family

11/97

04/99

05/99

Albuliformes

Notacanthoidei

Unidentified

Anguilliformes

Unidentified

Anguilloidei

Muraenidae

Congroidei

Ophichthidae

Nemichthyidae

Congridae

161

Nettastomatidae

Saccopharyngiformes

Saccopharyngoidei

Inf. Téc. Inst. Canario Cienc. Mar. n°13

01/00

11/00

03/02

1 1

2 125

Serrivomeridae

Eurypharyngidae

153

7 APPENDIX

Order

Family

Genus / Specie

11/97

04/99

05/99

Clupeiformes

Engraulidae

Engraulis encrasicolus

2902

Clupeidae

Sardinella spp.

Argentiniformes

Argentinidae

01/00

11/00

Argentina sphyraena

1 9

Glossanodon leioglossus Microstomatidae

Bathylagus sp.

Dolicholagus longirostris Platytroctidae Stomiiformes

Gonostomatidae

Sagamichthys schnakenbecki

Cyclothone spp. Cyclothone acclinidens

1 1

Cyclothone alba

Gonostoma denudatum

147

Sternoptychidae

Maurolicus muelleri

Phosichthyidae

Polymetme corythaeola

3 1

Vinciguerria spp.

Vinciguerria attenuata

Stomiidae

Vinciguerria nimbaria

Vinciguerria poweriae

Astronesthinae

Chauliodus spp.

5 2 1 3 1

Idiacanthus fasciola

Aulopiformes

Aulopidae Synodontidae

03/02

Melanostomiinae

Stomias spp.

Stomias boa

Aulopus filamentosus

Synodus spp.

Synodus saurus

1 1

Synodus synodus Chlorophthalmidae

Chlorophthalmus agassizii

Notosudidae

Ahliesaurus berryi

Scopelarchidae

Benthalbella infans

Scopelosaurus spp.

2 2

1 2

Scopelarchus analis

1 1

Unidentified Alepisauridae

Alepisaurus ferox Omosudis lowii

Paralepididae

Lestidiops spp.

1 34

Macroparalepis spp. Sudis hyalina Myctophiformes

Myctophidae

2 1 1

Benthosema suborbitale Centrobranchus nigroocellatus

Ceratoscopelus warmingii

Diaphus spp.

Diogenichthys atlanticus Hygophum hygomii

Hygophum reinhardtii

1 1

Lampadena sp.

Lampanyctus sp.

1 1

Lampanyctus crocodilus

154

Ceratoscopelus maderensis

Myctophum nitidulum

Lestidiops sphyrenoides

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Myctophum punctatum

Nannobrachium atrum Notoscopelus spp.

2 4

Notoscopelus resplendens

Symbolophorus veranyi

7 1

Taaningenichthys minimus Unidentified

1 1

Lampriformes

Lampridae

Lampris guttatus

Regalecidae

Regalecus glesne

Gadiformes

Macrouridae

Macrourinae

Melanonidae

Melanonus zugmayeri

Stephanoberyciformes

Melamphaidae

Melamphaes sp.

Beryciformes

Diretmidae

Diretmus argenteus

Trachichthyidae

Hoplostethus spp.

Syngnathidae

Entelurus aequoreus

Scorpaeniformes

Scorpaenidae

Helicolenus dactylopterus

Perciformes

Dicentrarchus spp.

Percichthyidae

Howella brodiei

Serranidae

Serranus sp. Callanthias ruber Apogon imberbis

Epigonidae

Microichthys coccoi

Carangidae

Campogramma glaycos Dentex spp.

1 2 1

1 1

1 1 6

Trachurus spp. Sparidae

Moronidae

Apogonidae

1 3

Callanthiidae

Scorpaena spp.

Anthias anthias

Hoplostethus mediterraneus Gasterosteiformes

4 1

Pagrus pagrus Pomacentridae

Pleuronectiformes

Chromis sp.

Abudefduf luridus

Labridae

Thalassoma pavo

Chiasmodontidae

Unidentified

Blenniidae

Parablennius spp.

Gobiidae

Crystallogobius linearis

Lesueurigobius heterofasciatus

Unidentified Type I

288

Unidentified Type II

589

Unidentified

Gempylidae

Diplospinus multistriatus

Trichiuridae

Benthodesmus elongatus

Lepidopus caudatus

Scombridae

Scomber colias

Nomeidae

Cubiceps gracilis

Caproidae

Capros aper

Bothidae

Arnoglossus spp.

Arnoglossus imperialis

Arnoglossus rueppelii Tetraodontiformes

Bothus podas

Balistidae

Balistes capriscus

Tetraodontidae

Canthigaster capistrata Sphoeroides spp.

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5 1

155

7 APPENDIX

7.2. SELECTED PICTURES OF LARVAE Engraulidae

Figure 26. Engraulis encrasicolus 21.1 mm [SL] with silvery eyes (left) and the overlapping dorsal and anal fin (right).

Figure 27. E. encrasicolus 24.0 mm [SL] with silvery eyes (left) and 23.7 mm [SL] with dark eyes (right).

Microstomatidae

Figure 28. Dolicholagus longirostris 20.1 mm [SL] with the prominent stalked eyes.

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Platytroctidae

Figure 29. Sagamichthys schnakenbecki 20.2 mm [SL]. The black tube above the cleithrum is clearly visible (right).

Gonostomatidae

Figure 30. Gonostoma denudatum 18.2 mm [SL] with photophores on the head, ventrally on the trunk (right) and three along the ventral margin of the caudal peduncle together with the characteristic pigmentation pattern at the caudal fin base (right).

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Sternoptychidae

Figure 31. Maurolicus muelleri 12.1 mm [SL] with the clustered groups of photophores (left) and the pigmentation pattern on the caudal peduncle (right).

Stomiidae

Figure 32. Astronesthinae 36.9 mm [SL] having an acute snout (left), a large finfold and a deflected terminal gut section (right).

Figure 33. Astronesthinae 23.6 mm [SL]. The advanced position of the dorsal fin relative to the anal fin is clearly visible (right).

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Figure 34. Chauliodus sp. 27.2 mm [SL] round in cross section and having no pigmentation.

Figure 35. Melanostomiinae 23.6 mm [SL]. The anal and the dorsal fin are situated far posteriorly at the body; opposite in position.

Figure 36. Stomias boa 32.6 mm [SL] with the elongated gut (deflected) and the opposite dorsal and anal fins (right).

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7 APPENDIX

Aulopidae

Figure 37. Aulopus filamentosus 12.2 mm [SL] with the characteristic pigmentation pattern of the peritoneal pigment and a dot at the caudal peduncle.

Synodontidae

Figure 38. Synodus saurus 20.5 mm [SL] with the round head and the series of paired lateral gut blotches.

Figure 39. Synodus synodus 21.5 mm [SL] and 25.7 mm [SL] with the round head (left) and the series of paired lateral gut blotches (right).

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Chlorophthalmidae

Figure 40. Chlorophthalmus agassizii 21.3 mm [SL] with the internal peritoneal pigmentation (left) and the prominent single spot laterally at the midline of the caudal peduncle (right).

Notosudidae

Figure 41. Ahliesaurus berryi 15.3 mm [SL] having the very elongated body (left) and the small dots on the caudal peduncle and caudal fin rays and the internal spots below the lateral midline (right).

Scopelarchidae

Figure 42. Scopelarchus analis 41.1 mm [SL] with the characteristic eyes (left) and the teeth on the tongue (right).

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Alepisauridae

Figure 43. Omosudis lowii 26.2 mm [SL] with the prominent large head (left) and the characteristic canine teeth on the palatines and dentaries (right).

Paralepididae

Figure 44. Sudis hyalina 17.0 mm [SL] having the elongated snout (left) and the long pectoral fins (right).

Myctophidae

Figure 45. Ceratoscopelus warmingii 11.7 mm [SL] with a pair of spots on the terminal gut (left) and Hygophum reinhardtii with the ovoid eyes having the choroid tissue and characteristic ventral pigmentation pattern (lright).

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SPATIAL AND SEASONAL PATTERNS IN SPECIES COMPOSITION AND OCCURRENCE OF FISH LARVAE IN THE AREA OF THE CANARY ISLANDS, EASTERN CENTRAL ATLANTIC

Figure 46. Symbolophorus veranyi 11.1 mm [SL] with the slightly stalked eyes (left) and the enlarged pectoral fins (right).

Figure 47. Symbolophorus veranyi 11.3 mm [SL]. The wing shaped pectoral fin base and its elongated rays reach beyond the anus.

Regalecidae

Figure 48. Regalecus glesne ca. 113 mm [SL] with the compressed large head.

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Macrouridae

Figure 49. Macrourinae 35.5 mm [SL] with the characteristic head (left) and the anal fin rays being longer than the dorsal fin rays (right).

Melanonidae

Figure 50. Melanonus zugmayeri 26.7 mm [SL] having the origin of the small pectoral and pelvic fins at the same level just behind the blunt head (left) and the small, narrow caudal fin (right).

Diretmidae

Figure 51. Diretmus argenteus 8.3 mm [SL] with the prominent head spination and the posteroventrally directed preopercular spine (left). The parietal spine in detail (right).

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Trachichthyidae

Figure 52. Hoplostethus sp. 12.0 mm [SL] and 12.7 mm [SL] having a deep (left) and laterally strongly compressed (right) body shape.

Figure 53. Hoplostethus mediterraneus 18.1 mm [SL] with tiny stings at the base of each fin ray (not along the first three spines starting from the right side).

Syngnathidae

Figure 54. Entelurus aequoreus 33.4 mm [SL] with the characteristic shape of the head (left) and the prominent subdorsal rings (right).

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7 APPENDIX

Diretmidae

Figure 55. Scorpaena sp. with 14.7 mm [SL] (above) and 9.8 mm [SL] (below). Characteristic is the head spination.

Moronidae

Figure 56. Dicentrarchus sp. 16.5 mm [SL]

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Howellidae

Figure 57. Howella brodiei ca. 8 mm [SL] (left) and 10.3 mm [SL] (right). Characteristic are the little stellate melanophores all over the body.

Serranidae

Figure 58. Anthias anthias 9.4 mm [SL]. Mark the elongated opercular spines directed towards the anus.

Callanthiidae

Figure 59. Callanthius ruber 12.7 mm [SL] with the prominent pigmentation above the brain.

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Epigonidae

Figure 60. Microichthys coccoi ca.17.2 mm [SL] with the prominent pigmentation pattern.

Carangidae

Figure 61. Campogramma glaycos 15.2 mm [SL] (left) and 12.4 mm [SL] (right) with the large spines at the preoperculum with its largest spine in the angle (left) and the prominent separated two first spines of the anal fin (right).

Figure 62. Trachurus sp. 14.1 mm [SL] with the prominent pigmentation pattern and dorsal and anal fins.

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Chiasmodontidae

Figure 63. Chiasmodontidae 14.5 mm [SL] with the staggered spots on the ventral and dorsal margin of the body.

Chiasmodontidae

Figure 64. Parablennius sp. 18.5 mm [SL]. Identification to genus level is done by meristic counts.

Gobiidae

Figure 65. Crystallogobius linearis 16.8 mm [SL] and 15.7 mm [SL] unfixed the body is transparent. Characteristic are the internal melanophores and sexual dimorphism. These specimens are males with their prominent dorsal spines and curved lower jaw (left).

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7 APPENDIX

Figure 66. Lesueurigobius heterofasciatus 19.1 mm [SL] with the prominent different intense brownish vertical bands along the body.

Figure 67. Gobiidae Type I 11.3 mm [SL] having a double row of stellate melanophores along the dorsal bin base (right).

Figure 68. Gobiidae Type II 10.3 mm [SL] having a blotch behind the dorsal fin base (right).

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Gempylidae

Figure 69. Diplospinus multistriatus 8.1 mm [NL] with the serrated pelvic spine is also elongated reaching beyond the vent while the rays are absent.

Scombridae

Figure 70. Scomber colias 18.9 mm [SL] (above) and 15.9 mm [SL] (below) with the characteristic pigmentation pattern (above) without the pigmentation on the cleithral symphysis (above left) and the anal and dorsal finlets (below).

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7 APPENDIX

Caproidae

Figure 71. Capros aper 10.6 mm [SL] having a deep body and the entire body surface is full with tiny spines.

Bothidae

Figure 72. Arnoglossus sp. 13.2 mm [SL] with the pigmented gas bladder and the prominent pigmentation pattern. The eyes have not migrated yet in that specimen (left).

Figure 73. Bothus podas 25.9 mm [SL] (left) and 12.1 mm [SL] (right) having no pigmentation and no migrated eye. The eyes have not migrated yet in that specimen.

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Tetraodontidae

Figure 74. Sphoeroides sp. 7.8 mm [SL] with the slender, but inflated, body shape (left) and the body spinules (right).

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173