habitat choice in adult longfin damselfish: territory characteristics and relocation times

Habitat choice in adult longfin damselfish:

territory characteristics and relocation times

Karen L. Cheney, Isabelle M. Cote*

Centre for Ecology, Evolution and Conservation, School of Biological Sciences,

University of East Anglia, Norwich NR4 7TJ, UK

Received 5 March 2001; received in revised form 29 November 2001; accepted 5 January 2002

Abstract

Habitat selection by coral reef fish during initial settlement has been shown to depend on various

biotic and abiotic characteristics. However, relatively little is known of the factors influencing habitat

choice by adults during post-settlement processes such as relocation or migration. In this study, we

first characterised the habitat of longfin damselfish (Stegastes diencaeus Jordan and Rutter)

territories to quantify territory variability. Characteristics such as percentage cover of rock, sand, live

coral and distance from sand were highly variable, while territory area, turf and macro algae cover

were relatively uniform across territories.

We then assessed the importance of specific habitat characteristics by experimentally removing

damselfish and measuring recolonisation times in relation to these characteristics. The presence of

nest sites markedly increased the speed of territory recolonisation after experimental removals. Other

variable territory characteristics such as substrate type, rugosity and the presence of cleaning stations

did not affect recolonisation speed. In general, males recolonised territories faster than females, and

males were more likely to recolonise territories previously owned by males with an active nest site.

Thus, intraspecific competition for high-quality nest sites may generate sex differences in territory

relocation and highly stable sex-specific patterns of adult distribution.

D 2002 Elsevier Science B.V. All rights reserved.

Keywords: Eupomacentrus diencaeus; Habitat selection; Reef fish distribution; Territoriality

1. Introduction

Local patterns of adult distribution in marine organisms with high potential for larval

dispersal can be determined at the time of recruitment through habitat selection for

0022-0981/02/$ - see front matter D 2002 Elsevier Science B.V. All rights reserved.

PII: S0022 -0981 (02 )00500 -2

* Corresponding author. Tel.: +44-1-603-593172; fax: +44-1-603-592250.

E-mail address: [email protected] (I.M. Cote).

www.elsevier.com/locate/jembe

Journal of Experimental Marine Biology and Ecology

287 (2003) 1–12

characteristics such as substratum type (Sale et al., 1984; Shulman, 1984; Wellington,

1992; Tolimieri, 1995, 1998; Danilowicz, 1997; Gutierrez, 1998), depth (Jones, 1986;

Gutierrez, 1998) and conspecific density (Sweatman, 1983; Jones, 1987; Booth, 1992;

Booth and Beretta, 1994). Such preferences have been shown to have direct effects on

juvenile growth and survival (Jones, 1988, 1990; Wellington, 1992). However, density-

dependent post-settlement processes such as predation (Carr and Hixon, 1995), competi-

tion (Forrester, 1995) and relocation (Bartels, 1984; Robertson, 1988) can also markedly

affect the distribution of subadults and adults. Relocation and migration have been

particularly overlooked despite the fact that, even in territorial species, adults and

subadults have been reported to move, sometimes in substantial numbers, both within

contiguous habitat (Robertson and Foster, 1982; Itzkowitz, 1985) and between discon-

nected habitat patches (Jones, 1987; Robertson, 1988; Forrester, 1990).

The factors motivating adult reef fish decisions to switch territories are not clear, but

probably involve a search for higher quality territories. In high-density populations,

individuals may initially have to settle in suboptimal habitats (Itzkowitz, 1977) and then

move to better territories when they become available. Itzkowitz (1991), for example,

showed that male beaugregory damselfish (Stegastes leucostictus Muller and Troschel)

were likely to move to new, artificial nest sites only if the latter were of similar or superior

quality to their original nest site. These switches were accompanied by increased

reproductive performance.

The determinants of territory quality can include any territory feature that can influence

fitness-related parameters such as growth rates, reproductive success, parasite loads and

risk of predation. Habitat characteristics such as nest site quality can have an obvious

direct effect on reproductive success, but other attributes may also be important in defining

habitat quality. Solandt et al. (in press) found that areas of high macroalgal cover

influenced territorial and breeding behaviour of longfin damselfish (Stegastes diencaeus

Jordan and Rutter). Algal availability has also been related to reproductive success in the

beaugregory damselfish (Itzkowitz and Slocum, 1995) and to foraging success in territorial

triggerfish (Chen et al., 2001). Limited shelter abundance can generate strong intraspecific

competition in some species which leaves a significant proportion of the population

without access to a territory and associated mating opportunities (e.g. Kroon et al., 2000),

while shelter size on a territory can determine predation risk to the territory owner (Hixon

and Beets, 1993). The presence of a cleaning station has yet to be considered but it could

affect adult habitat selection since proximity to cleaners is correlated to lower ectoparasite

loads in longfin damselfish (Cheney and Cote, 2001).

On coral reefs, habitat quality is often inferred from a comparison between reef areas

with and without the species of interest (e.g. Ebersole, 1985; Tolimieri, 1998). Such

comparisons generate descriptions of optimal and suboptimal habitats at a scale which

may be too coarse to relate to individual adult relocation decisions, since most relocating

adults move within habitats (Robertson and Foster, 1982; Itzkowitz, 1985; Wellington and

Victor, 1988). Moreover, for abundant species such as many damselfish, there may be few

areas within a habitat which are not encompassed within territories. As an alternative

approach to quantifying habitat quality, we therefore used the time taken to recolonise

artificially vacated territories. In saturated habitats, individuals should only switch to better

quality habitats, leading to intense competition for these high-quality sites. Thus, the speed

K.L. Cheney, I.M. Cote / J. Exp. Mar. Biol. Ecol. 287 (2003) 1–122

of colonisation of recently vacated territories should be related to territory quality

(Wellington and Victor, 1985, 1988), with high-quality sites being recolonised faster than

those of poor-quality (Itzkowitz et al., 1995).

In this paper, we investigate the factors affecting relocation decisions in the territorial

damselfish S. diencaeus. First, we quantify existing variability in territory features of

longfin damselfish, and then relate recolonisation speed of experimentally vacated

territories to specific habitat characteristics. We also consider behavioural costs (in terms

of defense against intruders) and benefits (in terms of foraging frequency) of territory

ownership, which may be independent of measurable habitat characteristics but important

for territory quality, and also relate these to relocation speed. Finally, we compare territory

quality for male and female longfin damselfish and consider the impact of observed sex-

related differences in territory requirements on damselfish distribution.

2. Materials and methods

2.1. Study site and species

Our study was conducted between May and August 2000 on the fringing reefs off the

Bellairs Research Institute, in the Barbados Marine Reserve, Barbados, West Indies

(13j10VN, 59j30VW). The study site was located in the spur and groove zone of the

South Bellairs reef (100–200 m from shore) and in the patch reef of North Bellairs (80–

130 m from shore) at depths ranging from 3.5 to 6 m. These reefs are degraded and are

predominantly covered in turf algae, which are grazed by herbivores such as damselfish

(Pomacentridae), parrotfish (Scaridae) and surgeonfish (Acanthuridae). There is very little

macroalgal and live coral cover, and the principal large corals found on the spurs are

Siderastrea siderea Ellis and Solander and Montastrea annularis Ellis and Solander.

We focused on longfin damselfish (S. diencaeus), an abundant species in which both

sexes aggressively defend mutually exclusive territories (ca. 1 m2) against a variety of

intruders. Each territory provides the resident with food resources in the form of a tended

algal mat (Robertson, 1984), and males may establish a nesting site within their territory

by removing algae from a suitable vertical area of coralline rock. Following a lunar cycle,

males with nest sites entice females into their territory to lay eggs during ca. 20 days/

month. Males then guard eggs in their nest for 5 days until hatching.

2.2. Territory characteristics

Thirty-five damselfish were chosen haphazardly within the study site. The territory of

each damselfish was mapped by observing each fish for a minimum of 15 min to locate

territorial boundaries. To assess variability in territory characteristics of longfin damsel-

fish, we quantified the following habitat parameters. The area of each territory was

measured using coloured nuts to divide the area into triangles, the sides of which were

calculated using straight-line measurements. The areas of each triangle were then

calculated and summed to obtain the total planar territory area. Substrate cover was

estimated using a 1-m2 quadrat with a 10� 10 point grid, which was systematically moved

K.L. Cheney, I.M. Cote / J. Exp. Mar. Biol. Ecol. 287 (2003) 1–12 3

to cover the entire territory. Under each point, substratum type (rock, rubble, sand, dead

coral) and live cover type (coral, sponge, turf, macro algae) were recorded. Substratum

cover and live cover were then expressed as a proportion of total points in a territory. The

planar distance of each territory to the nearest sand patch and height above the sand were

measured. Rugosity was estimated using SCI (substrate complexity index), i.e. the ratio of

a 1-m length of chain as it follows the contours of the substratum compared to the straight-

line distance between the ends of the chain. An SCI of 1.0 indicates a flat surface while

increasing SCI indicates substrata of increasing complexity. Conspecific density surround-

ing each focal fish was measured by counting the number of longfin damselfish within a 3-

m radius. Conspecific neighbours usually shared territorial boundaries. Conspecific

density is essentially identical to total damselfish density because dusky damselfish (S.

dorsopunicans Poey) and threespot damselfish (S. planifrons Cuvier), the only two species

to interact with longfin damselfish on our study site, very rarely occurred within 3 m of a

focal longfin damselfish. Bicolor S. partitus Poey and yellowtail damselfish Micro-

spathodon chrysurus Cuvier were present but were not considered because they interact

rarely with longfin damselfish. In fact, the territory boundaries of both species often

overlap with those of longfin damselfish, suggesting little competition for space or

resources (Robertson, 1984). Finally, we also noted the presence of active nest sites in

each territory, which was ascertained through observations during the reproductive period

prior to the removal experiment, and whether there was a cleaning station occupied by

cleaning gobies, Elacatinus spp., in the territory.

2.3. Behavioural observations

Behavioural observations were carried out on a subsample of 18 focal damselfish (all

males) to generate information on some of the costs and benefits of territory ownership.

Each male was observed for seven 15-min periods between 0900 and 1100 h over the 2

weeks before removal. The number of intruders onto the territories, the number of

chases and the number of bites taken on the substratum by the territory owners were

recorded.

2.4. Damselfish removals

The 35 focal damselfish were removed from their territory between 1400 and 1600 h

during non-spawning periods over 4 months (May–August 2000). Individual damselfish

were herded into a barrier net surrounding the territory, captured with a hand net and

removed from the reef. The standard length of each damselfish was measured to the

nearest mm and each fish was sexed by external examination of the urogenital pore.

Each vacated territory was monitored after 15, 30, 60, 120 and 180 min on the day of

removal. After this period, each territory was monitored 12–15 and 24 h after removal,

and then daily until recolonisation occurred. After damselfish were removed, a

neighbouring conspecific often began patrolling the vacant territory but still remained

within its own adjacent territory. The time of onset of such patrols was recorded, as was

the time when complete recolonisation occurred. Although it was obvious when a

neighbour was patrolling both territories, the identity of the final recoloniser (i.e.


neighbouring or non-neighbouring fish) could not usually be firmly established, and

hence the characteristics of the territory abandoned by each recoloniser could not be

measured. The recoloniser was later sexed by observing spawning behaviour or by

external examination of urogenital pore. Territory boundaries rarely altered during the

study period.

2.5. Statistical analysis

There was no difference among months in recolonisation times or any other measured

variables, hence the results of all removals were pooled together. Recolonisation times

were log-transformed to meet the assumptions of parametric testing. Other data could not

be transformed to achieve normal distribution, therefore, were analysed using nonpara-

metric statistics. The significance of multiple comparisons of habitat differences between

males and females was adjusted using sequential Bonferroni corrections (Rice, 1989).

Only results that remained significant after correction are reported.

3. Results

3.1. Variability in territory characteristics

The percentage cover of rubble, sand, dead coral, live coral and sponge was highly

variable (coefficient of variation, V >100%) but territories appeared to be relatively

uniform in rock, macroalgae and turf algae cover (Table 1). In terms of physical

Table 1

Variation in characteristics of longfin damselfish territories

Component Range MeanF S.D. Coefficient of

variation, V

Pearson’s

correlation

r P

Percent abiotic Rock (%) 43.2–100 78.3F 12.1 15 � 0.06 0.74

cover Rubble (%) 0–11.1 1.0F 2.3 232* 0.12 0.50

Sand (%) 0–30.2 3.6F 6.0 167* 0.16 0.38

Dead coral (%) 0–90.0 5.1F 5.6 110* � 0.24 0.17

Percent live Live coral (%) 0–28.4 9.01F10.2 113* � 0.20 0.26

cover Turf (%) 22.0–100 65.7F 17.9 27 0.20 0.24

Sponge (%) 0–5.0 0.7F 1.9 253* 0.07 0.68

Macroalgae (%) 0–16.4 4.1F 2.3 43 0.23 0.19

Other physical Area (m2) 0.5–2.6 1.2F 0.4 37 � 0.20 0.26

characteristics Damselfish density

(fish per m2)

0.8–1.2 1.0F 0.1 12

Height from sand (m) 0–3.2 1.2F 0.6 49 � 0.98 0.57

Distance from sand (m) 0–2.4 0.3F 0.5 212* 0.13 0.46

Rugosity 1.0–2.5 1.25F 0.2 14

Pearson’s correlation relating habitat characteristics to recolonisation times.

* Indicates values of over 50%.


characteristics, territory area, surrounding density of longfin damselfish and rugosity

did not vary greatly whereas distance from sand was more variable (Table 1). Fifteen

of the 25 territories occupied by males had an active nest site; 10 territories were

occupied by females. Seventeen territories had a cleaning station within their boun-

daries.

There was a significant difference between male and female territories in their distance

from sand (meanF S.D.: males: 0.15F 0.29 m, females: 0.82F 0.71 m; t-test for unequal

variances: t=� 2.89, df = 10.3, P= 0.016), with males being closer to a sand patch than

females. There were no other significant differences in habitat characteristics between

males and females (t-test, all P>0.05). In addition, females were as likely as males to have

a cleaning stations within their territories (v2 = 0.41, df = 1, P= 0.52).

3.2. Costs and benefits of territory ownership

The commonest intruders onto longfin damselfish territories were bluehead wrasse

(Thalassoma bifasciatum Bloch) (46% of all intrusions) and brown chromis (Chromis

multilineata Guicherot) (24% of all intrusions). Overall intrusion rates onto damselfish

territories varied from 0 to 223 fish per 15 min. Chase rates by resident damselfish ranged

from 0 to 35 chases per 15 min. There was a significant correlation between intrusion and

chase rates (r = 0.72, df = 16, P= 0.001), suggesting that intrusion rate reflects the cost of

territory ownership.

The number of intrusions onto territories was not correlated with any of the habitat

characteristics measured (Spearman’s rank correlations, P>0.05 in all cases). Intrusion

rates were similar on territories with and without nest sites (Mann–Whitney U-test:

U = 39, N1 = 10, N2 = 10, P= 0.44). However, there were significantly more intrusions onto

territories that contained cleaning stations (Mann–Whitney U-test: U = 17, N1 = 9, N2 = 9,

P= 0.04).

The foraging rates of resident damselfish varied from 76 to 144 bites per 15 min

(meanF S.D.: 110.3F 19.1 bites per 15 min) but were not correlated with any of the

habitat characteristics measured (N = 18, all P < 0.35). Also, foraging rates were not related

to the presence of a cleaning station (t=� 0.44, df = 16, P= 0.66) or the presence of a nest

site (t =� 0.84, df = 16, P= 0.41).

3.3. Recolonisation of vacant territories

All experimentally vacated territories were eventually recolonised by a conspecific. The

time taken for neighbours to begin patrolling the vacated territory ranged from 3 min to 24

h, while the time taken for complete recolonisation of territories ranged from 30 min to

960 h.

Recolonisation times for all territories combined were not influenced by any of the

measured characteristic of territories (Table 1). In addition, there was no difference in

recolonisation times between territories with cleaning stations and those without

(t=� 0.75, df = 33, P= 0.46). These results held when males and females were considered

separately. However, the presence of a nest site had a significant effect on recolonisation

speed. Territories originally owned by males with a nest site were patrolled by neighbour-


ing fish significantly sooner (ANOVA: F = 7.14, df = 33, P < 0.005) and eventually

recolonised faster (ANOVA: F = 31.32, df = 32, P < 0.001; Fig. 1) than territories of males

without nests or territories of females.

Fig. 1. Time taken to recolonise territories originally owned by males with or without nests, or by females. Means

are shownF S.E. Sample sizes are shown in parentheses.

Fig. 2. Time taken by males, females and subadults to recolonise vacant territories. Means are shownF S.E.

Sample sizes are shown in parentheses.


The measured costs and benefits of territory ownership did not influence recolonisation

speed. Recolonisation time was not correlated with either the rate of intrusions on each

territory (rs =� 0.15, N = 18, P= 0.62) or the foraging rate of the original owner

(rs =� 0.03, N = 18, P= 0.71).

Males recolonised territories significantly faster than females (t =� 2.58, df = 31,

P= 0.02; Fig. 2). Males recolonised 13 of the 15 territories initially occupied by males

with nests, while female territories were mainly recolonised by females (Table 2). The

territories of males without nests were as likely to be colonised by males as by females

(Table 2).

There was a significant size difference between the sexes (males: 88.9F 5.7 mm;

females: 83.8F 6.9 mm; t-test: t= 3.32, df = 33, P= 0.032), and between males with and

without nests (males with nest: 91.0F 4.9 mm; males without nest: 85.5F 5.5 mm; t-test:

t= 2.62, df = 23, P= 0.015). Recolonisation speed increased with the size of the original

owner (rs =� 0.53, N = 35, P= 0.001). This relationship, however, disappeared when we

considered males with nests (rs =� 0.24, N = 15, P = 0.40), males without nests

(rs =� 0.41, N = 10, P= 0.23), and females (rs =� 0.29, N = 10, P= 0.42) separately.

4. Discussion

Territory relocation by adult longfin damselfish occurred readily when space became

available on the reef. We found that the main determinant of territory quality for male

damselfish was the presence of a nest site, since experimentally vacated territories were

recolonised significantly faster, and predominantly by males, when they contained an

established nesting area. No other territory characteristic influenced recolonisation speed.

For females, however, no clear pattern emerged.

Caution should be used in the interpretation of recolonisation speed as an index of

habitat quality. Although it is generally true that better-quality territories will be

recolonised more quickly (Itzkowitz, 1977, 1991), the speed of recolonisation will also

depend on the difference in territory quality between territories which are available and

those currently occupied. In habitats of uniformly high quality, recolonisation speed may

therefore bear little relation to territory quality. This may not have been a problem in our

study site since habitat within our study site appeared extremely variable. In addition,

relocation speed will be influenced by the costs incurred by individuals during

relocation. Male damselfish, for example, should not relocate during the nest guarding

period owing to the cost of losing their brood. In addition, the costs of relocation may

Table 2

Numbers of males, females and subadult longfin damselfish recolonising territories originally owned by various

damselfish types

Original territory owner Recoloniser

Male Female Subadult

Males with nest (N= 15) 13 2 0

Males without nest (N= 10) 4 4 2

Females (N = 10) 1 9 0


be prohibitive when there are many non-territorial individuals (floaters) vying for

unoccupied space. In our study, the latter two costs were not applicable since the

removals were carried out during the non-breeding period and there were no floaters in

the population.

Longfin damselfish territories were highly variable in some habitat features, such as

coral cover and distance from sand, but less variable in other characteristics such as algal

cover, depth and area. While high variability could have offered scope for observing

habitat preferences during relocation, it may also suggest that these characteristics were

unimportant to damselfish during initial territory settlement. In fact, most habitat

characteristics, whether highly variable or not, did not influence territory recolonisation

speed. These results are surprising since we had expected, for example, that territories

with cleaning stations would be recolonised more quickly. Although benefits of

proximity to cleaning stations have been measured for longfin damselfish (Cheney and

Cote, 2001), these limited benefits appear to be outweighed by the increased rates of

intrusions caused by the presence of a cleaning station within a territory. Similarly, we

had expected more intense competition for sites where algal cover is greater or of higher

quality (Itzkowitz and Slocum, 1995). However, although spatial variation in algal

quality which can affect damselfish distribution may exist (e.g. Solandt et al., in press),

turf alga was the predominant substratum cover on our degraded study site, and foraging

resources appeared abundant. It remains possible that the lack of preference for specific

features of the physical habitat by adults simply mirrors a similar absence of habitat

associations of juvenile S. diencaeus (Booth and Beretta, 1994), although young longfin

damselfish preferentially settle near conspecifics while relocating adults did not show this

tendency.

The only territory feature that influenced the speed of vacant territory colonisation was

the presence of a nest site. Selecting a territory with an established nest site is clearly

advantageous for males moving from territories without a suitable nest, since the new

nest is already cleaned and females will be familiar with this oviposition site. Males

moving from territories already containing a nest site may also benefit if the new nest is

of higher quality than the former. Although reproductive success in damselfish depends

largely on male courtship intensity and the presence of eggs in the nest (reviewed in

Petersen, 1995), evidence suggests that oviposition site quality can also be an important

factor to enhance reproductive success (e.g. Itzkowitz and Makie, 1986; Itzkowitz, 1990;

Sikkel, 1995). If high-quality sites are limited on the reef, then competition for territories

with the best nest sites should be intense (Wellington and Victor, 1985, 1988). Several

lines of evidence suggest that this is the case for longfin damselfish. First, a large

proportion of males in our sample (< 40%) did not have nests within their territories.

Second, males recolonised territories faster than females, suggesting that they are under

greater pressure to improve territory quality. Third, territories with nests were recolonised

fastest of all vacant territories, underlining the importance of nest sites. Finally, the largest

males had territories with nests, which would be expected if nest acquisition was a

competitive process.

The importance of nest sites for males generated sex differences in at least one territory

characteristic. Males were found on territories that were situated closer to sand. Such

territories tended to be situated on the periphery of reef patches and spurs and were


characterised by the presence of large vertical surfaces (e.g. side of coral head, or boulders)

which were suitable as nest sites. This apparent segregation of male and female territories

should be stable over long periods since the territories of a given sex were predominantly

recolonised by individuals of that sex.

Elucidating the dynamics of adult territoriality is essential to understand fully the

patterns of reef fish distribution. Robertson (1995, 1996) has shown that interspecific

competition among adult damselfish was important in determining patterns of abun-

dance. Intraspecific competition may have similarly strong effects when high-quality

resources are limited. We have shown that this is the case for territories with nest sites.

Given that only male damselfish provide parental care, intraspecific competition for

high-quality nest sites should generate sex differences in frequency of territorial

relocation and sex-specific patterns of distribution since nests are non-randomly

dispersed through the habitat. Such sex/habitat associations have only recently been

described (Sikkel et al., 2000), but they may be a common feature of reef fish

distribution.

Acknowledgements

Thank you to the staff at the Bellairs Research Institute, especially its director, the late

Professor Joan Marsden, for logistical support. We thank Elizabeth Whiteman, John

Reynolds, Mike Haley and one anonymous reviewer who made suggestions which greatly

enhanced the paper. [AU]

References

Bartels, P.J., 1984. Extra-territorial movements of a perennially territorial damselfish, Eupomacentrus dorsopu-

nicans Poey. Behaviour 91, 312–322.

Booth, D.J., 1992. Larval settlement and preferences by domino damselfish Dascyllus albisella Gill. J. Exp. Mar.

Biol. Ecol. 155, 85–104.

Booth, D.J., Beretta, G.A., 1994. Seasonal recruitment, habitat associations and survival of pomacentrid reef fish

in the US Virgin Islands. Coral Reefs 13, 81–89.

Carr, M.H., Hixon, M.A., 1995. Predation effects on early post-settlement survivorship of coral reef fishes. Mar.

Ecol., Prog. Ser. 124, 31–42.

Chen, T.C., Ormond, R.F.G., Mok, H.K., 2001. Feeding and territorial behaviour in juveniles of three co-existing

triggerfishes. J. Fish Biol. 59, 524–532.

Cheney, K.L., Cote, I.M., 2001. Are Caribbean cleaning symbioses mutualistic? Costs and benefits of visiting

cleaning stations to longfin damselfish. Anim. Behav. 62, 927–933.

Danilowicz, B.S., 1997. The effects of age and size on habitat selection during settlement of a damselfish.

Environ. Biol. Fishes 50, 257–265.

Ebersole, J.P., 1985. Niche separation of two damselfish species by aggression and differential microhabitat

utilization. Ecology 66, 14–20.

Forrester, G.E., 1990. Factors influencing the juvenile demography of a coral reef fish. Ecology 71, 1666–1681.

Forrester, G.E., 1995. Strong density-dependent survival and recruitment regulate the abundance and species of a

coral reef fish. Oecologia 111, 137–143.

Gutierrez, L., 1998. Habitat selection by recruits establishes local patterns of adult distribution in two species of

damselfishes: Stegastes dorsopunicans and S. planifrons. Oecologia 115, 268–277.


Hixon, M.A., Beets, J.P., 1993. Predation, prey refuges and the structure of coral reef fish assemblages. Ecol.

Monogr. 63, 77–101.

Itzkowitz, M., 1977. Spatial organization of the Jamaican damselfish community. J. Exp. Mar. Biol. Ecol. 28,

217–241.

Itzkowitz, M., 1985. Aspects of the population dynamics and reproductive success in the permanently territorial

beaugregory damselfish. Mar. Behav. Physiol. 185, 57–69.

Itzkowitz, M., 1990. Heterospecific intruders, territorial defense and reproductive success in the beaugregory

damselfish. J. Exp. Mar. Biol. Ecol. 140, 49–59.

Itzkowitz, M., 1991. Habitat selection and subsequent reproductive success in the beaugregory damselfish.

Environ. Biol. Fishes 30, 287–293.

Itzkowitz, M., Makie, D., 1986. Habitat structure and reproductive success in the beaugregory damselfish. J. Exp.

Mar. Biol. Ecol. 97, 305–312.

Itzkowitz, M., Slocum, C.J., 1995. Is the amount of algae related to territorial defense and reproductive success in

the beaugregory damselfish? Mar. Behav. Physiol. 24 (4), 243–250.

Itzkowitz, M., Itzkowitz, D.E., Shelly, D., 1995. Territory use and disuse in the beaugregory damselfish. Bull.

Mar. Sci. 57, 653–662.

Jones, G.P., 1986. Food availability affects growth in a coral reef fish. Oecologia 70, 136–139.

Jones, G.P., 1987. Some interactions between residents and recruits in two coral reef fishes. J. Exp. Mar. Biol.

Ecol. 114, 169–182.

Jones, G.P., 1988. Experimental evaluation of the effect of habitat structure and competitive interactions on the

juveniles of two coral reef fishes. J. Exp. Mar. Biol. Ecol. 123, 115–126.

Jones, G.P., 1990. The importance of recruitment to the dynamics of a coral reef fish population. Ecology 71,

1691–1698.

Kroon, F.J., de Graaf, M., Liley, N.R., 2000. Social organisation and competition for refuges and nest sites

in Coryphopterus nicholsii (Gobiidae), a temperate protogynous reef fish. Environ. Biol. Fishes 57 (4),

401–411.

Petersen, C.W., 1995. Male mating success and female choice in permanently territorial damselfishes. Bull. Mar.

Sci. 57, 690–704.

Rice, W.R., 1989. Analyzing tables of statistical tests. Evolution 43, 223–225.

Robertson, D.R., 1984. Cohabitation of competing territorial damselfishes on a Caribbean coral reef. Ecology 65,

1121–1135.

Robertson, D.R., 1988. Abundances of surgeonfishes on patch-reefs in Caribbean Panama: due to settlement, or

post-settlement events? Mar. Biol. 97, 495–501.

Robertson, D.R., 1995. Competitive ability and the potential for lotteries among territorial reef fishes. Oecologia

103, 180–190.

Robertson, D.R., 1996. Interspecific competition controls abundance and habitat use of territorial Caribbean

damselfishes. Ecology 77, 885–899.

Robertson, D.R., Foster, S.A., 1982. Off-reef emigration of young adults of the labrid fish Epibulus insidiator.

Copeia 1982, 227–229.

Sale, P.F., Douglas, W.A., Doherty, P.J., 1984. Choice of microhabitats by coral reef fishes at settlement. Coral

Reefs 3, 91–99.

Shulman, M.J., 1984. Resource limitation and recruitment patterns in coral reef fish assemblages. J. Exp. Mar.

Biol. Ecol. 74, 85–109.

Sikkel, P.C., 1995. Effects of nest quality on male courtship and female spawning-site choice in an algal-nesting

damselfish. Bull. Mar. Sci. 57, 682–689.

Sikkel, P.C., Fuller, C.A., Hunte, W., 2000. Habitat/sex differences in time at cleaning stations and ectoparasite

loads in a Caribbean reef fish. Mar. Ecol., Prog. Ser. 193, 191–199.

Solandt, J.L., Campbell, A.C., Haley, M.P., 2001. Incidence and effects of patchy Diadema antillarum Philippi

distribution on longfin damselfish population dynamics at Discovery Bay, Jamaica. Bull. Mar. Sci. (in press).

Sweatman, H.P.A., 1983. Influence of conspecifics on choice of settlement sites by larvae of two pomacentrid

fishes (Dascyllus aruanus and D. reticulatus) on coral reefs. Mar. Biol. 75, 225–229.

Tolimieri, N., 1995. Effects of microhabitat characteristics on the settlement and recruitment of a coral reef fish at

two spatial scales. Oecologia 102, 52–63.


Tolimieri, N., 1998. Contrasting effects of microhabitat use of large-scale adult abundance in two families of

Caribbean reef fishes. Mar. Ecol., Prog. Ser. 167, 227–239.

Wellington, G.M., 1992. Habitat selection and juvenile persistence control the distribution of two closely related

Caribbean damselfishes. Oecologia 90, 500–508.

Wellington, G.M., Victor, B.C., 1985. El-Nino mass coral mortality—a test of resource limitation in a coral reef

damselfish population. Oecologia 68, 15–19.

Wellington, G.M., Victor, B.C., 1988. Variation in components of reproductive success in an undersaturated

population of coral-reef fish: a field perspective. Am. Nat. 131, 588–601.


habitat choice in adult longfin damselfish: territory characteristics and relocation times

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