INTRODUCTIONTop

A crucial goal in fish population ecology is to describe the temporality of key life-story traits, such as growth, reproduction (spawning) and subsequent recruitment in the benthic system. This information is essential for proper management of fish species subjected to human exploitation (Caldow and Wellington 2003Caldow C., Wellington G.M. 2003. Patterns of annual increment formation in otoliths of pomacentrids in the tropical western Atlantic: implications for population age-structure examination. Mar. Ecol. Prog. Ser. 265: 185-195., Morgan 2008Morgan M.J. 2008. Integrating reproductive biology into scientific advice for fisheries management. J. Northwest Atl. Fish. Sci. 41: 37-51., Smallwood et al. 2013Smallwood C.B., Hesp S.A., Beckley L.E. 2013. Biology, stock status and management summaries for selected fish species in south-western Australia. Fish. Res. Rep. No. 242. Department of Fisheries, Western Australia.).

The pomacentrids (damselfishes) are a diverse fish family, including ca. 29 genera and 396 species distributed throughout tropical to temperate oceans of the world (Robertson 1998Robertson D.R. 1998. Do coral reef fishes have a distinctive taxonomic structure? Coral Reefs 17: 179-186., Nelson 2006Nelson J.S. 2006. Fishes of the World, (4th. ed.), Wiley, New York., Eschmeyer 2015Eschmeyer W.N. (ed.) 2015. Catalog of Fishes. Electronic version accessed January, 2015. ). They are among the first fishes described by Linnaeus, back in the 18th century, and they have subsequently received the attention of other well-known fish taxonomists and naturalists (Bleeker 1877Bleeker P. 1877. Mémoire sur les Chromides marins ou Pomacentroides de l’Inde archipelagique. Nat. Verh. Holl. Maatsch. Haarlem. 3: 1-166., Cuvier and Valenciennes 1830Cuvier G., Valenciennes A. 1830. Histoire naturelle des poissons. Tome cinquiéme. Chez F.G. Levrault, Paris.). In terms of the number of species, this family is the third-largest fish group in coral-reef ecosystems, after Gobiidae (>1500 species) and Labridae (>600 species) (Wainwright and Bellwood 2002Wainwright P.C., Bellwood D.R. 2002. Ecomorphology of feeding in coral reef fishes. In: Sale P.F. (ed.), Coral reef fishes: dynamics and diversity in a complex ecosystem. Academic Press, San Diego, CA, pp 33-56.), although they often reach larger abundances on reefs (Frédérich et al. 2009Frédérich B., Fabri G., Lepoint G., et al. 2009. Trophic niches of thirteen damselfishes (Pomacentridae) at the Grand Récif of Toliara, Madagascar. Ichthyol. Res. 56: 10-17.).

Most damselfishes are territorial and show aggressive behaviour when defending their territories (Randall et al. 1997Randall J.E., Allen G.R., Steene R.C. 1997. Fishes of the Great Barrier Reef and Coral Sea. Crawford House Publishing, Bathurst, New South Wales., Randall 2005Randall J.E. 2005. Reef and shore fishes of the South Pacific: New Caledonia to Tahiti and the Pitcairn Islands. University if Hawai’i Press, Honolulu, HI, USA., Gordon et al. 2015Gordon T.A.C., Cowburn B., Sluka R.D. 2015. Defended territories of an aggressive damselfish contain lower juvenile coral density than adjacent non-defended areas on Kenyan lagoon patch reefs. Coral Reefs 34: 13-16.); this behaviour has led to a plethora of studies that have used them as model organisms to test a range of ecological and behavioural questions (Frédérich et al. 2009Frédérich B., Fabri G., Lepoint G., et al. 2009. Trophic niches of thirteen damselfishes (Pomacentridae) at the Grand Récif of Toliara, Madagascar. Ichthyol. Res. 56: 10-17.). Pomacentrids deposit elliptical eggs, which have a tuft of adhesive filaments, on the substratum (Hutchinson 2006Hutchinson D.S. 2006. The Encyclopedia of Fishes: A Complete Visual Guide, Sydney, Frog City Press.). During the incubation time, males, but in some cases females, guard eggs until they hatch, frequently attacking intruders (Allen et al. 2006Allen G.R., Cross N.J., Allen C.J. 2006. Labridae. Labfinae. In: Zoological Catalogue of Australia. Volum. 35. Fishes. pp. 1368-1418). This parental care may range from hiding eggs to guarding their offspring in elaborately prepared structures for up to several months (Balshine and Sloman 2011Balshine S., Sloman K. A. 2011. Parental Care in Fishes. In: Farrell A.P., (ed.), Encyclopedia of Fish Physiology: From Genome to Environment, volume 1, pp. 670-677.). Moreover, damselfishes change their colour patterns according to their reproductive patterns (Souza et al. 2011Souza A.T., Ilarri M.I., Medeiros P.R., et al. 2011. Unusual colour patterns of territorial damselfishes (Pomacentridae: Stegastes) in the Southwestern Atlantic. Mar. Biodivers. Rec. 4: e101.). The bridal colour (Bakker and Mundwiler 1994Bakker T.M., Mundwiler B. 1994. Female mate choice and male red coloration in a natural three-spined stickleback (Gasterosteus aculeatus) population. Behav. Ecol. 5: 74-80.), size (Schmale 1981Schmale M.C. 1981. Sexual selection and reproductive success in males of the Bicolor damselfish, Eupomacentrus partitus (Pisces: Pomacentridae). Anim. Behav. 29: 1179-1184., Côté and Hunte 1989Côté I.M., Hunte W. 1989. Male and female mate choice in the red lip blenny: why bigger is better. Anim. Behav. 38: 78-88.) and courtship behaviour are phenotypic characteristics of males to improve their reproductive success (Knapp and Kovach 1991Knapp R.A., Kovach J.T. 1991. Courtship as an honest indicator of male parental quality in the bicolor damselfish (Stegastes partitus). Behav. Ecol. 4: 295-300.). Because patterns of sexual development differ among pomacentrids, attempts to assess gonadal development require data of the different phases of the fish life cycle and at different times of the year (Sadovy de Mitcheson and Liu 2008Sadovy de Mitcheson Y., Liu M. 2008. Functional hermaphroditism in teleosts. Fish Fish. 9: 1-43.). Recruitment of pomacentrids is widespread on onshore reefs, but juveniles may prefer alternative microhabitats to adults; ontogenetic shifts in habitat preferences can influence the spatial distribution of adult and juvenile damselfishes (Lirman 1994Lirman D. 1994. Ontogenetic shifts in habitat preferences in the three-spot damselfish, Stegastes planifions (Cuvier), in Roatan Island. J. Exp. Mar. Biol. Ecol. 180(1): 71-81.).

The diversity of damselfishes drops rapidly with increasing latitude (Kingsford 1999Kingsford M. 1999. Territorial Damselfishes. In: Andrew N. (ed.) Under Southern Seas: The ecology of Australia’s rocky reefs. Sydney, Univerisity of New South Wales Press Ltd.). This pattern can be observed in the oceanic archipelagos of the northeastern Atlantic, including the Azores, Madeira, Canaries and Cabo Verde. In the Azores, two species cohabit, Similiparma lurida (Cuvier, 1830) and Chromis limbata (Valenciennes, 1833) (Santos et al. 1997Santos R.S., Porteiro F.M., Barreiros J.P. 1997. Marine fishes of the Azores: anannotated check-list and bibliography. Arquipel. Life Mar. Sci. Suppl. 1: 1-244., Leite et al. 2009Leite J.R., Bertoncini A.A., Bueno L., et al. 2009. The occurrence of Azores Chromis, Chromis limbata in the south-western Atlantic. Mar. Biodivers. Rec. 2: e145., Afonso et al. 2013Afonso P, Porteiro F.M., Fontes J., et al. 2013. New and rare coastal fishes in the Azores islands: occasional events or tropicalization process? J. Fish Biol. 83: 272-294., Froese and Pauly 2015Froese R., Pauly D., (eds). 2015. FishBase. World Wide Web electronic publication. www.fishbase.org). Both species also occur in Madeira, in addition to Abudefduf saxatilis (Linnaeus, 1758) and Chromis chromis (Linnaeus, 1758) (Freitas and Araújo 2006Freitas M., Araújo R. 2006. First record of Sergeant Major Abudefduf saxatilis (Linnaeus, 1758) (Pisces: Pomacentridae), from the island of Madeira (NE Atlantic Ocean). Bocagiana 218: 1-6., Wirtz et al. 2008Wirtz P., Fricke R., Biscoito M.J. 2008. The coastal fishes of Madeira Island - new records and an annotated check-list. Zootaxa, 26: 1-26., Froese and Pauly 2015Froese R., Pauly D., (eds). 2015. FishBase. World Wide Web electronic publication. www.fishbase.org). In the Canary Islands, Stegastes imbricatus Jenyns, 1840 also occasionally appears in addition to the species mentioned before (Brito et al. 2002Brito A., Pascual P.J., Falcón J.M., et al. 2002. Peces de las Islas Canarias. Catálogo comentado e ilustrado. Francisco Lemus Editor, La Laguna. 419 pp., Froese and Pauly 2015Froese R., Pauly D., (eds). 2015. FishBase. World Wide Web electronic publication. www.fishbase.org). Finally, in Cabo Verde Islands, five species are abundant, A. saxatilis, Abudefduf taurus (Müller and Troschel, 1848), Chromis lubbocki Edwards, 1986, Similiparma hermani (Steindachner, 1887), and S. imbricatus; and seven species are occasional, Abudefduf hoefleri (Steindachner, 1881), C. chromis, Chromis cyanea (Poey, 1860), Chromis multilineata (Guichenot, 1853), Microspathodon chrysurus (Cuvier, 1830), S. lurida, and Stegastes leucostictus (Müller and Troschel, 1848) (Wirtz et al. 2013Wirtz P., Brito A., Falcón J.M., et al. 2013. The coastal fishes of the Cape Verde Islands – new records and an annotated check-list (Pisces). Spixiana 36: 113-142., Freitas 2014Freitas R. 2014. The coastal ichthyofauna of the Cape Verde Islands: a summary and remarks on endemism. Zoologia Caboverdiana 5(1): 1-13., Hanel and John 2014Hanel R. John D.H.C. 2014. A revised checklist of Cape Verde Islands sea fishes. J. Appl. Ichthyol. 31 (1): 135-169.).

Damselfishes are excellent species as biological indicators, since they are small-sized, abundant, non-migratory and easily recognizable in the field and they are not usually a fishing target (Linton and Warner 2003Linton D.M., Warner G.F. 2003. Biological indicators in the Caribbean coastal zone and their role in integrated coastal management. Ocean. Coast. Manag. 46: 261–276.). In the Canary Islands, S. lurida, before known as Abudefduf luridus (Cooper et al. 2014Cooper W.J., Craig R.A., Jacob R.E., et al. 2014. Re-description and Reassignment of the Damselfish Abudefduf luridus (Cuvier, 1830) Using Both Traditional and Geometric Morphometric Approaches. Copeia 14(3): 473-480.), is a common fish inhabiting shallow-water rocky bottoms, especially vegetated reefs (Brito et al. 2002Brito A., Pascual P.J., Falcón J.M., et al. 2002. Peces de las Islas Canarias. Catálogo comentado e ilustrado. Francisco Lemus Editor, La Laguna. 419 pp., Tuya et al. 2004Tuya F., Boyra A., Sanchez-Jerez P., et al. 2004. Relationships between rocky-reef fish assemblages, the sea urchin Diadema antillarum and macroalgae throughout the Canarian archipelago. Mar. Ecol. Prog. Ser. 278: 157-169.), being abundantly captured by the artisanal fisheries fleet through traps deployed at <50 m depth (García-Mederos et al. 2015García-Mederos A.M., Tuya F., Tuset V.M. 2015. The structure of a nearshore fish assemblage at an oceanic island: insight from small scale fisheries through bottom traps at Gran Canary Island (Canary Islands, eastern Atlantic). Aquat. Living Resour. 1: 1-10.). Knowledge on this species in the Macaronesian area is limited to its ethology and spawning seasonality in the Azores Islands (Mapstone and Wood 1975Mapstone G.M., Wood E.M. 1975. The ethology of Abudeduf luridus and Chromis chromis (Pisces: Pomacentridae) from the Azores. J. Zool. 175(2): 179-199., Afonso and Santos 2005Afonso P., Serrão Santos R. 2005. Within-nest spawning-site preferences of female bluefin damselfish: the effect of early-stage eggs. Acta Ethologica 8: 5-11.). In this study, the main goal was to provide biological and ecological information on the population structure of S. lurida on rocky reefs at Gran Canary Island (Canary Islands, northeastern Atlantic). First, we assessed the reproductive ecology and inferred growth patterns. This approach provides important information from a fisheries perspective, i.e. spawning seasons and first maturity sizes. Second, we analysed the spatial and temporal patterns of abundance, describing annual recruitment patterns and therefore connecting the timing between reproduction and subsequent recruitment. Additionally, we sought to determine whether variation in abundance patterns of adults, sub-adults and juveniles varied at small spatial scales in relation to habitat composition and complexity. Overall, integration of this information provided insight into the life history of this species.

MATERIALS AND METHODSTop

Study area

This study was performed in Gran Canaria Island (Fig. 1), which is located at the centre of the Canarian Archipelago, with ca. 45 km diameter and a maximum elevation of 1950 m above sea level (Carracedo et al. 2002Carracedo J.C., Pérez F.J., Ancochea E., et al. 2002. Cenozoic volcanism II: In: Gibbons W. and Moreno T., (eds) The Canary Islands. The Geology of Spain. The Geol. Soc. Lond, Total alkali versus silica diagrams (TAS) with analyses of Canaries volcanic rocks, pp. 439-472.). Sediments and rocky reefs mainly compose the nearshore bottoms, with a high variability in the distribution and complexity of these habitats, which can be colonized by a range of canopy-forming species (Tuya and Haroun 2006Tuya F., Haroun R.J. 2006. Spatial patterns and response to wave exposure of photophilic algal assemblages across the Canarian Archipelago: a multiscaled approach. Mar. Ecol. Prog. Ser. 311: 15-28., Tuya et al. 2014Tuya F., Ribeiro-Leite L., Arto-Cuesta N., et al. 2014. Decadal changes in the structure of Cymodocea nodosa seagrass meadows: Natural vs. human influences. Estuar. Coast. Shelf. S. 137: 41-49.).

Biological sampling

A total of 629 specimens were collected, from January to December 2012 at Telde (east of the island, Fig. 1), through bottom trapping carried out by the local artisanal fleet. Traps were deployed between 18 and 30 m depth, either individually or in strings of 2-3 traps along a fishing rope. The number of days on which traps were placed on the seafloor varied between 3 and 10. For each individual, the total length (TL, mm), total weight (TW, g), and gutted weight (GW, g) were recorded. The sex and stage of sexual maturation (EMS) were recorded by a macroscopic examination of the gonads. Gonads were first removed and weighed to the nearest 0.01 g (GNW). Maturity stages were classified as immature (I), developing (II), spawning (III), regressing (IV) and regenerating (V) (Brown-Peterson et al. 2011Brown-Peterson N.J., Wyanski D.M., Saborido-Rey F., et al. 2011. A standardized terminology for describing reproductive development in fishes. Mar. Coast. Fish. Dynam. Manag. Ecosys. Sci. 3: 52-70.). The sagittae otoliths were used to determine fish age. They were extracted from all individuals, cleaned and stored dry in plastic vials.

Age and growth

The right otolith was selected to determine fish age. As otolith weight is considered as an indicator of fish growth rate (e.g. Pawson 1990Pawson M.G. 1990. Using otolith weight to age fish. J. Fish. Biol. 36: 521-531., Fletcher 1991Fletcher W.J. 1991. A test of the relationship between otolith weight and age for the pilchard Sardinops neopilchardus. Can. J. Fish. Aquat. Sci. 48: 35-38., Cardinale et al. 2000Cardinale B.J., Nelson K., Palmer M.A. 2000. Linking species diversity to the functioning of ecosystems: on the importance of environmental context. Oikos 91: 175-183.), the criterion was based on the absence of significant differences in mean weight between left and right otoliths (Student t-test, t=0.0098, p=0.992, n=200).

First, whole otoliths were immersed in a 1:1 glycerin–alcohol solution and observed under a stereomicroscope (NIKON SMZ 1000) using reflected light and a dark background. This method, however, did not provide reliable age estimates, so a random subsample of 200 otoliths was selected to perform the study through sectioning. For that, whole otoliths were embedded in epoxy resin and two or three transverse sections (1 mm thick) were cut through the central region with a slow-speed circular saw (Buehler, ISOMET-TM) to obtain a section that included the otolith core. Otolith sections were then mounted on a glass slide using Crystalbond as a mounting media and polished using decreasing grit abrasive paper (3M Lapping Film). Sections that included the core were examined under a compound microscope with transmitted light (Axioplan, Zeiss; Carl Zeiss Inc., Oberkochen, Germany) connected to a digital camera (ProgRresTM C10 plus; Jenoptik, Jena, Germany). Under transmitted light, the core and opaque bands appear as dark rings, and the wider translucent bands as clear or hyaline rings. The count path of the annuli was from the nucleus towards the tip of the inner face next to the sulcus, where the deposition of seasonal rings appeared clearly defined (Fig. 2a).

To assess the precision of readings, one experienced reader counted opaque bands without knowledge of fish size at least twice. To minimize reading bias, the two readings were separated 2-3 months after randomization in the process of readings. When readings differed, a third reading was taken. An otolith was only considered unreadable, and therefore excluded from the analysis, when the differences between readings did not improve after this procedure. The coefficient of variation (CV=SD⁄mean) was used to measure precision of annuli counts together with a paired t-test to statistically compare differences between readings (Chang 1982Chang W.B. 1982. A statistical method for evaluating the reproducibility of age determinations. Can. J. Fish. Aquat. Sci. 39: 1208-1210., Campana 2001Campana S.E. 2001. Accuracy, precision and quality control in age determination, including a review of the use and abuse of age validation methods. J. Fish Biol. 59: 197-242.). To assess the yearly pattern of deposition of otolith annuli, the appearance of each otolith margin was recorded as opaque or translucent. The timing of annulus formation was examined by plotting the percentage occurrence of otoliths with a peripheral opaque band as a function of the sampling month (Morales-Nin 1992Morales-Nin B. 1992. Determination of growth in bony fishes from otolith microstructure. FAO Fish. Tech. Paper 322: 1-51.). January 1 (peak spawning; see results) was considered the birthdate, hence their annuli count was assigned to equal age. Finally, the von Bertalanffy growth equation was used to describe the growth of the species; it was fitted to the observed individual length-at-age data, rather than the frequently used mean length-at-age, to show individual growth variability:

L = L_∞ [1 – е^{–k (t – t0)}]

where L is the TL (cm), L_∞ is the asymptotic length, k is the growth coefficient, t is the age (years) and t₀ is the hypothetical age at which length is zero. To better estimate growth parameters, ages of smallest individuals (undifferentiated) were used for both sexes (García-Mederos et al. 2010García-Mederos A.M., Tuset V.M., Santana J.I., et al. 2010. Reproduction, growth and feeding habits of stout beardfish Polymixia nobilis (Polymixiidae) off the Canary Islands (NE Atlantic). J. Appl. Ichthyol. 26: 872-880.). Hotelling’s T2 test was used to compare growth parameters between males and females (Cerrato 1990Cerrato R.M. 1990. Interpretable statistical test for growth comparisons using parameters in the von Bertalanffy equation. Can. J. Fish. Aquat. Sci. 47: 1416-1426., Gordo 1996Gordo L.S. 1996. On the age and growth of bogue, Boops boops (L.) from the Portuguese coast. Fish. Manag. Ecol. 3: 157-164.).

A multiple linear regression model was used to evaluate whether age determination was predicted by otolith weight, otolith diameter and fish size (Boehlert 1985Boehlert G.W. 1985. Using objective criteria and multiple regression models for age determination in fishes. Fish. Bull. 83: 103-117., Schwamborn and Ferreira 2002Schwamborn S.H.L., Ferreira S.P. 2002. Age structure and growth of the dusky damselfish, Stegastes fuscus, from Tamandaré reefs, Pemambuco, Brazil. Environ. Biol. Fishes. 63(1): 79-88.). All variables used in the multiple linear regression were log-transformed to conform the assumptions of linearity, i.e. normality and homogeneity of variances (Zar 1996Zar J.H. 1996. Biostatistical Analysis, 3rd ed. Prentice Hall International, New Jersey, 662 pp.).

Reproductive biology

The spawning pattern was assessed from monthly changes in the frequency of the maturity stages and the gonadosomatic index (West 1990West G. 1990. Methods of assessing ovarian development in fishes: a review. Aust. J. Marine. Freshwater Res. 41: 199-222.):

GSI = 100 (GNW ⁄ GW)

Size at first maturity was based on the examination of males and females in mature phases (phase III, phase IV, or phase V) and immature individuals collected during the spawning period. The TL of all individuals was used to estimate the size at first maturity (L₅₀), defined as the size at which 50% of all fish are at sexually mature phases. Maturity curves were adjusted using the logistic curve (Pope et al. 1975Pope J., Margetts A., Hamley J., et al. 1975. Manual of methods for fish stock assessment. Part 3-Selectivity of fishing gear. FAO Fish. Tech. Pap. 41(1): 1-73.):

P = 100 / (1 + exp (a + bTL))

where P is the percentage of mature individuals as a function of size class (TL), and a and b are specific parameters which can change during the life cycle. A logarithmic transformation was initially applied to calculate the parameters a and b by means of linear regression. An Analysis of Covariance (ANCOVA) was used to compare the curves of maturity between sexes.

The sex ratio of the population (males: females), and the sex ratio by size intervals (10 mm) were determined. Sex ratios were tested statistically for significant deviations from the expected 1:1 ratio via chi-square tests (α=0.05).

Abundance patterns

Spatial and temporal variation in the abundance of S. lurida was studied from visual censuses carried out on a monthly basis between January and December 2012, at two randomly selected localities (Fig. 1). One locality is in the northwest (NW, Agaete, 28°5’57.58”N, 15°42’33.48”W) and the other in the east of the island (E, Telde, 27°59’21.68”N, 15°22’12.96”W); visual counts were performed between 0.2 and 5.8 m depth. Only individuals >2 cm (TL) were counted to optimize the in situ identification. On each sampling occasion, n=5 replicated 25 m transects were haphazardly surveyed during daylight hours. The abundance of fish within 2 m of either side of each transect (100 m²) was recorded on waterproof paper by the same diver, according to standard procedures for the study region (Boyra et al. 2004Boyra A., Sanchez-Jerez P., Tuya F., et al. 2004. Attraction of wild coastal fishes to Atlantic subtropical cage fish farms, Gran Canaria, Canary Islands. Environ. Biol. Fishes. 70(4): 393-401., Tuya et al. 2004Tuya F., Boyra A., Sanchez-Jerez P., et al. 2004. Relationships between rocky-reef fish assemblages, the sea urchin Diadema antillarum and macroalgae throughout the Canarian archipelago. Mar. Ecol. Prog. Ser. 278: 157-169.). Individuals were categorized as juveniles (TL<4 cm), sub-adults (TL 6-8 cm) and adults (TL>10 cm) based on morphological characteristics, fundamentally related to their colouration and body size (Mapstone and Wood 1975Mapstone G.M., Wood E.M. 1975. The ethology of Abudeduf luridus and Chromis chromis (Pisces: Pomacentridae) from the Azores. J. Zool. 175(2): 179-199.). This approach was subsequently endorsed by our data (see results). Concurrently, the diver counted on his way back the number of large (>1 m) and small (<1 m) topographic elements of the rocky substrate (i.e. cracks, crevices, caves, holes per 100 m²), and visually estimated the percentage cover of algae, following standardized procedures via the Linear Point Intercept sampling technique (Ohlhorst et al. 1988Ohlhorst S.L., Liddell W.D., Taylor R.J., et al. 1988. Evaluation of reef census techniques. Proc. 6th Intl. Coral Reef. Symp. 2: 319-324. Townsville, Australia.). The type of substrate was recorded every metre along the 25 m long transect (i.e. 25 points per transect). Water temperature on the bottom was registered using an underwater thermometer.

Differences in the total abundance of juveniles, sub-adults and adults among months (fixed factor) and localities (random factor) were tested through ANOVAs from square root transformed data; this was necessary to avoid heterogeneous variances. Multiple linear regressions tested whether the number of small and large topographic elements (100 m⁻²), the type of substrate, and the algal cover (per transect) affected abundances of the damselfish over time. To retain variables with good explanatory power, the Akaike information criterion (AIC) routine was used as a selection criterion for each model (the smaller the value the better the model, Anderson and Legendre 1999Anderson M.J., Legendre P. 1999. An empirical comparison of permutational methods for tests of partial regression coefficients in a linear model. J. Stat. Comp. Simul. 62: 271-303.), and the contribution of each independent variable to each model was described with partial r² values. Collinearity diagnoses among independent (predictive) variables were carried out through Spearman-rank-correlations. All analyses were based on a ‘forward’ selection procedure.

RESULTSTop

Population structure

Fish ranged in size from 56 to 157 mm TL, and weighed between 3.9 and 76.6 g TW. A total of 419 males, 113 females and 110 immature specimens (gonads were characterized by small, thin and translucent filaments) were identified. Significant differences (Student t-test, t=11.22, p=0.002) were found between the mean sizes of sexes; males were larger than females (Table 1). The size and weight distributions differed significantly between males and females (Kolmogorov-Smirnov test, length: d=6.00, p<0.0001; weight: d=5.67, p<0.0001).

The overall ratio of males to females was 1:0.26; the hypothesis of uniformity between sexes was therefore rejected (χ² =180.5, p<0.05). Females were found in most size intervals, but males were more abundant in the larger size interval (130-140 mm) (Table 2).

Growth

The readings coincided for 171 otoliths (85%) and were dissimilar for the remaining 29 (15%). There was no significant difference in age estimation between readings (CV=2.7%; t=–0.094, p>0.05). The growth rings, opaque and translucent, were usually well visible (Fig. 2). Marginal zone analysis showed a pattern of alternating narrow translucent zones and wide opaque zones, forming one annulus per year. Otoliths with an opaque edge (faster growth) were more abundant (52.9-87.5%) from November to March (spawning season, see below), while otoliths with a translucent edge (slower growth) were more common during the remaining months (Fig. 3).

Fish ranged in age from 0 to 18 years; the age of most fishes was between 8 and 10 years. Females reached higher maximum ages (18 years) than males (14 years). Significant differences in mean sizes between sexes were obtained from V and X age classes (Supplementary material, Table S1). A considerable variability in the length-age relationship indicated considerable differences in individual growth. The growth curve obtained by age-at-length data was well described through a von Bertalanffy growth fit, attaining a determination coefficient (r²) of 0.439 for males and 0.707 for females (Fig. 4, Table 3). Significant differences were found between the von Bertalanffy growth curves of both sexes (Hotelling’s T²-test, = 68.654>T₀² _0.05,3.181 = 8.515).

Partial regression coefficients indicated that radius diameter (r²-partial=0.775, t=13.728, p<0.001) and otolith weight (partial r²=0.5575, t=6.367, p<0.001) were significant predictors of fish age (Fig. 5). By contrast, fish length did not contribute to explaining variation (partial r²=0.338, t=–0.728, p=0.338).

Reproduction

Males in phase III were observed from November to May, with a maximum peak in February (Fig. 6a). The maximum occurrence of spawning females (phase III) also occurred from November to March, including a peak in February (Fig. 6b). The presence of regressing females (phase IV) was observed from February to August. Females at immature, regressing and developing/regenerating phases (V, I and II, respectively) were found throughout the entire year. A slight increase in phase III females was observed in May, which would likely result in spawning activity during June, suggesting the possibility of a secondary breeding season (Fig. 6b). For males, the GSI increased from November to May, including a peak in January (0.258) (Fig. 6c). The GSI of females also increased from November to May, with a peak of maximum activity in February (3.897) (Fig. 6d). Overall, integration of EMS and GSI results with maturity phases throughout the year indicated a spawning season from November to May. The maturity curves were clearly different between males and females (ANCOVA, F=34.811, p<0.001); the size at which 50% of individuals are mature was 103.44 mm TL for males and 84.71 mm TL for females (Fig. 7).

Abundance and recruitment patterns

The abundance of juveniles, sub-adults and adults varied inconsistently over time between localities (ANOVA: Locality × Month, p<0.05, Table 4). However, juvenile abundances were significantly higher in April at both localities (9.0±2.6 ind/100 m² at Agaete; 11.0±2.7 ind/100 m² at Telde, Fig. 8a, b). The abundance of sub-adults attained higher mean values in July at Agaete (19.6±2.8 ind/100 m²), while the larger values were observed between November and January (14.2±0.7 ind/100 m², 13.2±2.8 ind/100 m², respectively) at Telde (Fig. 8c,d). Adults peaked in abundance at similar times at both localities; November (22.4±1.8 ind/100 m²) at Agaete and November-December (28.2±2.1 ind/100 m², 23.8±1.7 ind/100 m², respectively) at Telde (Fig. 8e, f).

The percentage of sandy cover in the case of juveniles and adults, and the percentage cover of algae for sub-adults, accounted for the largest contribution to variability in fish abundances at Agaete. However, the number of small topographic elements was the largest contributor to variation in the abundance of sub-adults and adults at Telde (Supplementary material, Table S2). For the majority of specimens, the best models provided by the AIC routine included both the number of small topographic elements with algae and with sandy cover, i.e. patterns of abundance were, in most cases, affected by these two descriptors of the habitat (Table S2). Juvenile abundance was significantly predicted by the presence of large topographic elements at Telde (Table S2), i.e. juveniles were particularly abundant on high relief areas.

DISCUSSIONTop

The Canary damsel, Similiparma lurida, is quite similar in size (157 mm TL) to other pomacentrids from the Macaronesian archipelagos, such as C. limbata (120 mm SL) and S. hermani (160 mm SL), but smaller than A. saxatilis (229 mm TL), C. chromis (250 mm TL) and A. taurus (250 mm TL) (Froese and Pauly 2015Froese R., Pauly D., (eds). 2015. FishBase. World Wide Web electronic publication. www.fishbase.org). Our findings clearly indicated that this species is of early rapid growth, because it reached half of its asymptotic length within the first year. In pomacentrids, small-sized species seem to have a faster growth (Dulčić and Kraljević 1995Dulčić J., Kraljević M. 1995. Age, growth and mortality of damselfish (Chromis chromis L.) in the eastern middle Adriatic. Fish. Res. 22: 255-265., Tzioumis and Kingsford 1999Tzioumis V., Kingsford M.J. 1999. Reproductive biology and growth of the temperate damselfish Parma microlepis. Copeia 1999(2): 384-361., Wilson and Meekan 2002Wilson D.T., Meekan M.G. 2002. Growth-related advantages for survival to the point of replenishment in the coral reef fish Stegastes partitus (Pomacentridae). Mar. Ecol. Prog. Ser. 231: 247-260.), although it may be a response to unpredictable recruitment success (Longhurst 2006Longhurst A. 2006. The sustainability myth. Fish. Res. 81: 107-112.). Moreover, differences in growth were obtained between sexes: males grow faster (0.28 years^–1) than females (0.23 years^–1), favouring the presence of males in larger sizes. Although this is not uncommon for many fishes, it is particularly important for fish species with nuptial behaviour, in which males have to defend their territories and care for their offspring (Breder and Rosen 1966Breder C.M., Rosen D.E. 1966. Modes of reproduction in fishes. T.F.H. Publications, Neptune City, New Jersey. 941 pp., Dulčić and Kraljević 1995Dulčić J., Kraljević M. 1995. Age, growth and mortality of damselfish (Chromis chromis L.) in the eastern middle Adriatic. Fish. Res. 22: 255-265., Bracciali et al. 2014Bracciali C., Piovano S., Sarà G., et al. 2014. Seasonal changes in size, sex-ratio and body condition of the damselfish Chromis chromis in the central Mediterranean Sea. J. Mar. Biol. Assoc. U. K. 94(5): 1053-1061., Allen et al. 2006Allen G.R., Cross N.J., Allen C.J. 2006. Labridae. Labfinae. In: Zoological Catalogue of Australia. Volum. 35. Fishes. pp. 1368-1418). Some studies have also indicated that fish size is correlated with social dominance, which additionally favours the acquisition of food, mate choice, retention and reproductive success (e.g. Hoffman 1985Hoffman S.G. 1985. Effects of size and sex on the social organization of reef-associated hogfishes, Bodianus sp. Environ. Biol. Fishes. 14: 185-197., DeMartini 1988DeMartini E.E. 1988. Size-assortative courtship and competition in two embiotocid fishes. Copeia 1988: 336-344., Folkvord 1991Folkvord A. 1991. Growth, survival and cannibalism of cod juveniles (Gadus morhua L.): effects of feed type, starvation and fish size. Aquaculture 97: 41-59., Webster and Hixon 2000Webster M.S., Hixon M.A. 2000. Mechanisms and individual consequences of intraspecific competition in a coral-reef fish. Mar. Ecol. Prog. Ser. 196: 187-194., Hobbs and Munday 2004Hobbs J-P.A., Munday P.L. 2004. Intraspecific competition controls spatial distribution and social organization of the coral-dwelling goby Gobiodon histrio. Mar. Ecol. Prog. Ser. 278: 253-259., Fero and Moore 2008Fero K., Moore P. 2008. Social spacing of crayfish in natural habitats: what role does dominance play? Behav. Ecol. Sociobiol. 62: 1119-1125.). From an ecological perspective, a sexual variability of certain life-history traits may represent a trade-off, the costs paid in the currency of fitness when a beneficial change in one trait is linked to a detrimental change in another (Charnov and Krebs 1973Charnov E.L., Krebs J.R. 1973. On clutch size and fitness. Ibis 116: 217-219., Stearns 1989Stearns S.C. 1989. The evolutionary significance of phenotypic plasticity. Bioscience 39: 436-445., Lester et al. 2004Lester N.P., Shuter B.J., Abrams P.A. 2004. Interpreting the von Bertalanffy model of somatic growth in fishes: the cost of reproduction. Proc. R. Soc. Lond. Ser. B. Biol. Sci. 271: 1625-1631.). Fast growth may involve a cost in terms of reproduction, as early maturing occurs at a larger size; when growth is delayed, however, maturing occurs at a smaller size (Stearns and Koella 1986Stearns S.C., Koella J.C. 1986. The evolution ofphenotypic plasticity in life-history traits: Predictions of reaction norms for age and size at maturity. Evolution 40: 893-913., Charnov 2008Charnov E.L. 2008. Fish growth: Bertalanffy k is proportional to reproductive effort. Environ. Biol. Fishes. 83: 185-187., this study).

Alterations in somatic growth are not always reflected in otolith morphometry, due to continuous growth and metabolic non-dependence between the two processes (Reznick et al. 1989Reznick D., Lindbeck E., Bryga H. 1989. Slower growth results in larger otoliths: An experimental test with guppies (Poecilia reticulata). Can. J. Fish. Aquat. Sci. 46: 108-112., Fowler and Doherty 1992Fowler A.J., Doherty P.J. 1992. Validation of annual growth increments in the otoliths of two species of damelfish from the southern Great Barrier Reef. Aust. J. Mar. Freshwater Res. 43: 1057-1068., Secor and Dean 1992Secor D.H., Dean J.M. 1992. Comparison of otolith-based back-calculation methods to determine individual growth histories of larval striped bass, Morone saxatilis. Can. J. Fish. Aquat. Sci. 49(7): 1439-1454., Xiao 1996Xiao Y. 1996. A general model for estimating tag-specifi c shedding rates and tag interactions from exact or pooled times at liberty for a double tagging experiment. Can. J. Fish. Aquat. Sci. 53: 1852-1861.). Many studies have indicated that otolith thickness, or weight, can explain 80-95% variation in fish age (Fletcher 1991Fletcher W.J. 1991. A test of the relationship between otolith weight and age for the pilchard Sardinops neopilchardus. Can. J. Fish. Aquat. Sci. 48: 35-38., Newman et al. 1996Newman S.J., Williams D.McB., Russ G.R. 1996. Age validation, growth and mortality rates of the tropical snappers (Pisces: Lutjanidae), Lutjanus adetii (Castelnau, 1873) and L. quinquelineatus (Bloch, 1790) from the central Great Barrier Reef, Aust. Mar. Freshwater Res. 47(4): 575–584., Labropoulou and Papaconstantinou 2000Labropoulou M., Papaconstantinou C. 2000. Community structure of deep-sea demersal fish in the North Aegean Sea (northeastern Mediterranean). Hydrobiologia 440: 281-296.). However, Pawson (1990) Pawson M.G. 1990. Using otolith weight to age fish. J. Fish. Biol. 36: 521-531.concluded that this technique has a limited application in ageing fish from wild populations with highly variable growth rates. Schwamborn and Ferreira (2002)Schwamborn S.H.L., Ferreira S.P. 2002. Age structure and growth of the dusky damselfish, Stegastes fuscus, from Tamandaré reefs, Pemambuco, Brazil. Environ. Biol. Fishes. 63(1): 79-88. found a linear correlation (partial r²=0.739) between otolith weight and age of the pomacentrid Stegastes fuscus, but they recommended that the high variability of otolith weight-at-age hampered the use of this variable for an accurate prediction of age. In our study, otolith thickness provided a better relationship with age than weight. This is most likely because the otolith weight may underestimate the age of older fish (Beckman et al. 1991Beckman D.W., Stanley A.L., Render J.H., et al. 1991. Age and growth-rate estimating of sheepshead Archosargus probatocephalus in Louisiana waters using otoliths. Fish. Bull. 89: 1-8., Wilson et al. 1991Wilson C.A., Dean J.M., Prince E.D., et al. 1991. An examination of sexual dimorphism in Atlantic and Pacific blue marlin using body weight, sagittae weight, and age estimates. J. Exp. Mar. Biol. Ecol. 151: 209-225., Ferreira and Russ 1994Ferreira B.P., Russ G.R. 1994. Age and growth of the coral trout, Plectropomus leopardus Pisces: Serranidae from the Northern Great Barrier Reef, Australia. Fish. Bull. 92: 46-57., Worthington et al. 1995Worthington D.G., Fowler A.J., Doherty P.J. 1995. Variation in the relationship between otolith weight and age: implications for the estimation of age of two tropical damsel-fish (Pomacentrus moluccensis and P. wardi). Can. J. Fish. Aquat. Sci. 52: 233-242., Newman et al. 1996Newman S.J., Williams D.McB., Russ G.R. 1996. Age validation, growth and mortality rates of the tropical snappers (Pisces: Lutjanidae), Lutjanus adetii (Castelnau, 1873) and L. quinquelineatus (Bloch, 1790) from the central Great Barrier Reef, Aust. Mar. Freshwater Res. 47(4): 575–584., Tuset et al. 2004Tuset V.M., González J.A., Lozano I.J., et al. 2004. Age and growth of the blacktail comber, Serranus atricauda (Serranidae), off the Canary Islands (centraleastern Atlantic). Bull. Mar. Sci. 74: 53-68.).

There are many spawning strategies by marine fishes, including variation at daily, lunar and seasonal scales (Robertson 1991Robertson D.R. 1991. The role of adult biology in the timing of spawning of tropical reef fishes. In: Sale P.F. (ed.), The ecology of fishes on coral reefs, pp. 356-386. San Diego: Academic Press.). Our results demonstrated that S. lurida has a long spawning period (ca. 7 months) that coincides with an increase in fish growth and the appearance of denser otolith rings. For pomacentrids, seasonal cycles of spawning and recruitment can be synchronized, particularly when one main spawning peak dominates the spawning period (Robertson 1990Robertson D.R. 1990. Differences in the seasonalities of spawning and recruitment of some small neotropical reef fishes. J. Exp. Mar. Biol. Ecol. 144(1): 49-62.). This occurred for the population of S. lurida at Gran Canaria Island; recruitment of juveniles reached a maximum in April, immediately after the main peak of spawning (February). This outcome suggests that the duration between gamete release and the appearance of recruits >2 cm is about 2 months. This result sounds plausible, given the fact that larval dispersion in most pomacentrids is reduced (10-24 days, Thorrold and Milicich 1990Thorrold S.R., Milicich M.J. 1990. Comparison of larval duration and pre- and post-settlement growth in two species of damselfish, Chromis atripectoralis and Pomacentrus coelestis (Pisces: Pomacentridae), from the Great Barrier Reef. Mar. Biol. 105: 375-384., Nemeth 2005Nemeth R.S. 2005. Population characteristics of a recovering US Virgin Islands red hind spawning aggregation following protection. Mar. Ecol. Prog. Ser. 286: 81-97.), and that males guard fertilized eggs for only a few weeks (Thresher 1984Thresher R.E. 1984. Reproduction in reef fishes. T.F.H. Publications, Neptune City., Asoh and Yoshikawa 2002Asoh K., Yoshikawa T. 2002. The role of temperature and embryo development time in the diel timing of spawning in a coral-reef damselfish with high-frequency spawning synchrony. Environ. Biol. Fishes 64: 379-392., Bessa and Sabino 2012Bessa E., Sabino J. 2012. Territorial hypothesis predicts the trade off between reproductive opportunities and parental care in three species of damselfishes (Pomacentridae: Actinopterygii). Lat. Am. J. Aquat. Res. 40(1): 134-141.). Some authors have suggested that some variability in the time of recruitment may arise as a result of several spawning acts, or that larval duration varies due to environmental uncertainty (Robertson 1990Robertson D.R. 1990. Differences in the seasonalities of spawning and recruitment of some small neotropical reef fishes. J. Exp. Mar. Biol. Ecol. 144(1): 49-62., Thorrold and Milicich 1990Thorrold S.R., Milicich M.J. 1990. Comparison of larval duration and pre- and post-settlement growth in two species of damselfish, Chromis atripectoralis and Pomacentrus coelestis (Pisces: Pomacentridae), from the Great Barrier Reef. Mar. Biol. 105: 375-384.). The fact that recruitment patterns were studied in shallower waters (0.2-6 m) relative to collections of specimens via fishing traps (18-30 m) does not seem to disturb interpretations, as the habitat is the same: rocky reefs on infralittoral bottoms.

The present study also highlighted the influence of habitat structure on the spatial and temporal variability in the abundance of juveniles, sub-adults and adults of S. lurida on shallow-water bottoms. The paramount role of habitat structure (e.g. substratum composition) as a driver of fish distribution and abundance has been widely reported (Luckhurst and Luckhurst 1978Luckhurst B.E., Luckhurst K. 1978. Analysis of the influence of substrate variables on coral reef fish communities. Mar. Biol. 49: 317-323., Friedlander and Parrish 1998Friedlander A.M., Parrish J.D. 1998. Habitat characteristics affecting fish assemblages on a Hawaiian coral reef. J. Exp. Mar. Biol. Ecol. 224(1): 1-30., García-Charton and Pérez-Ruzafa 2001García-Charton J.A., Pérez-Ruzafa A. 2001. Spatial pattern and the habitat structure of a Mediterranean rocky reef fish local assemblage. Mar. Biol. 138: 917-934., Tuya et al. 2009Tuya F., Wernberg T., Thomsen M.S. 2009. Habitat structure affect abundances of labrid fishes across temperate reefs in south-western Australia. Environ. Biol. Fishes. 86: 311-319., 2011Tuya F., Wernberg T., Thomsen M.S. 2011. The relative influence of local to regional drivers of variation in reef fishes. J. Fish. Biol. 79: 217-234.). In general, our results suggest that rocky bottoms covered with algae interspersed with sandy patches seem an ideal habitat for S. lurida. This pattern, however, varied between localities. At Agaete, different habitat elements contributed to explaining variation in the abundances of sub-adults and adults, in particular the presence of small topographic elements and algal cover. A positive influence of algal cover may be explained by the fact that damselfish territories need to contain algae where females deposit their eggs (Knapp et al. 1995Knapp R.A., Sikkel P.C., Vredenberg V.T. 1995. Age of clutches in nests and within-nest spawning site preference of three damselfish species (Pomacentridae). Copeia 1995: 78-88., Navarrete-Fernandez et al. 2014Navarrete-Fernandez T., Landaeta M.F., Bustos C.A., et al. 2014. Nest building and description of parental care behavior in a temperate reef fish, Chromis crusma (Pisces: Pomacentridae). Rev. Chil. Hist. Nat. 87: 1-9.), which are further protected by males. Small-sized topographic elements, on the other hand, provide protection against predators. The presence of large topographic elements contributed to explaining the presence of juveniles at this locality. The tendency of juveniles to seek refuge after recruitment in the benthos, as way to avoid predation, has been reported for a range of reef fish species (Scharf et al. 2006Scharf F.S., Manderson J.P., Fabrizio M.C. 2006. The effects of seafloor habitat complexity on survival of juvenile fishes: Species-specific interactions with structural refuge. J. Exp. Mar. Biol. Ecol. 335: 167-176., Leitão et al. 2008Leitão F., Santos M.N., Erzini K., et al. 2008. The effect of predation on artificial reef juvenile demersal fish species. Mar. Biol. 153: 1233-1244.), including damselfishes (Almany 2004Almany G.R. 2004. Differential effects of habitat complexity, predators and competitors on abundance of juvenile and adult coral reef fishes. Oecologia 141(1): 105-113.). Hence, large ledges and outcrops, which we have included here as large topographic elements, seem to provide an ideal habitat for juveniles, which tend to concentrate in these areas. The positive influence of sandy cover on juvenile abundances may be an artefact, as large ledges and outcrops typically reduce water flow and ease sedimentation in their surroundings. Taken together, these outcomes may be indicative of a change in habitat use by S. lurida with ontogeny (e.g. an ontogenetic niche shift, Wilbur 1980Wilbur H.M. 1980. Complex life cycles. Ann. Rev. Ecol. Evol. Syst. 11: 67-93.). Ontogenetic shifts in microhabitat preferences by juveniles and adults of another pomacentrid (Stegastes planifrons) have also been registered (Lirman 1994Lirman D. 1994. Ontogenetic shifts in habitat preferences in the three-spot damselfish, Stegastes planifions (Cuvier), in Roatan Island. J. Exp. Mar. Biol. Ecol. 180(1): 71-81.); in particular, adults exhibit a preference for foliose coral heads, whereas juveniles exhibit a preference for dead foliose coral heads. At Telde, however, the number of small topographic elements exclusively contributed to explaining the abundances of sub-adults and adults. This fact may be explained by the presence of more homogenous bottoms at this locality. Therefore, the discrepancy in results between the two localities indicates that between-location variation in habitat composition and structure may affect partitioning of habitat niches between juvenile and adult populations of reef damselfishes.

ACKNOWLEDGEMENTSTop

We thank Nito and the crew of the vessel “Alvaro Tercero” for the supply of individuals. Our gratitude also goes to Nuria Raventos for her help with otolith readings. Also, we express our gratitude to two anonymous referees, whose comments significantly improved a previous draft.

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Supplementary material

The following material is available through the online version of this article and at the following link:
http://www.icm.csic.es/scimar/supplm/sm04343esm.pdf

Table S1. – Age-length key for S. lurida at Gran Canaria Island. n, number of individuals; SD, standard deviation; TL, total length; *, significant differences.

Table S2. – Results of multiple linear regression analyses testing for the relationship between the total abundances of juvenile, sub-adult and adult individuals and structural elements of the habitat for each locality. The Akaike information criterion routine was used to retain variables with good explanatory power.