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because it is hypothesized that liver mass reflects status-dependent differences in energy expenditure, but not energy storage or energy acquisition. Dominants perform more energetically costly behaviors (e.g. social policing and care) compared with subordinates, supporting the notion that energy expenditure drives liver investment. Dominants in large groups with many subordinates to monitor and dominants that own multiple territories with a large area to patrol tend to have smaller livers. Subordinates do not appear to use the liver as a strategic energy storage organ. Although subordinates feed more frequently than dominants, a negative correlation is found between feeding rates and liver size. This provides evidence that liver investment patterns are linked to status-driven differences in energy expenditure but not energy intake or storage.
586:, specifically the defense of their young. Dominant female breeders perform the most care and also display the highest levels of plasma testosterone compared to other individuals within the social group. Dominant male breeders provide a similar amount of care as the subordinate helpers, but breeding males also have the highest levels of 11-ketotestosterone, an important fish androgen. Breeders have higher levels of both androgens compared to helpers, but no matter what the sex or status of the fish, there is a weak but significant positive correlation between testosterone levels and the frequency of care. Androgens may promote the defense of young, which contradicts the commonly reported trade-off between androgen levels and degree of parental care (androgen levels and degree of parental care are usually negatively correlated).
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higher plasma levels of 11-ketotestosterone than newly subordinate males, but both types of males have similar testosterone levels. The ratio of 11-ketotestosterone to testosterone, which demonstrates the physiological importance of testosterone conversion to 11-ketotestosterone, has been shown to be positively correlated with submissive behavior in female winners and weakly positively correlated with aggressive behavior in male winners. Different androgens play equivalent roles in female versus male dominance establishment, and relative levels of 11-ketotestosterone and testosterone are implicated in female dominance behavior and perhaps the behavior of both sexes.
492:. In one study, 71% of male vacancies were filled by joiners. Joiners are incoming fish that are not originally part of the group. However, only 15% of female vacancies were filled. Helpers increase their frequency of cooperative behavior following the removal of a female breeder but not after the removal of a male breeder. This suggests that female breeder vacancies are typically filled by subordinate helpers who inherit from within the group, and male breeder vacancies are commonly filled by joining individuals, who are either existing breeders or helpers from other groups.
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genetic quality and resources. Males holding only one territory may provide their mates with significant assets but may not be able to outcompete neighbors for additional breeding positions because of their smaller body size and possibly higher parasite load. The lack of differences between monogamous and polygynous groups in terms of offspring survival, a measure of reproductive success, suggests that there may be few, if any, fitness consequences of polygynous pairing for females.
77:
524:, while pair females shared the investment in territory maintenance and direct brood care. There is high energy expenditure for submissive behavior in daffodil cichlids, which might be an indicator of submissive behavior being a reliable signal among cichlids. The considerable energy expenditure involved in territory defense suggests that these costs, in addition to risk, are probably taken into account when doing cost-benefit analyses.
549:
a year after the first check. The sizes of the breeder male, breeder female, and the largest helper within a group do not influence the parameters of the long-term field study, and also do not correlate with the sizes of these categories of fish after one year. This suggests that group size is the critical variable determining the success of the individuals in the group, rather than body size or fighting ability of group members.
52:
304:, Africa and are widespread in the southern part of the lake. They are found along the rocky coastlines of the countries of Burundi, the Democratic Republic of the Congo, Tanzania, and Zambia. There are a number of different geographical varieties. The variety known as the "daffodil" is very popular and is found along the steep rocky slopes of Kantalamba and Kambwinba.
574:
discriminate between size-matched familiar and unfamiliar male neighbors and whether they respond more aggressively toward unfamiliar males. The results showed that males spent significantly more time near the territorial boundary that they shared with unfamiliar neighbors and directed significantly more aggressive behavior toward these individuals.
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revealed mixed parentage in 80% of the examined groups. A case of shared maternity was detected when a subordinate female bred alongside the dominant female in a social group. This was not found to be true for the males, though. Extra-pair paternity was assigned to other dominant males who held their
358:
When parents and helpers care for offspring, the protection may reduce the predation risk to offspring, which may allow mothers to invest less per single offspring. Daffodil cichlid helpers protect the group offspring and reduce the offspring's mortality rate. Therefore, dominant females are expected
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These fish have a light-colored tan body washed with hints of yellow and bluish-purple spots. The yellow is stronger along the upper portion of the body and onto the dorsal fin, and around the base of the pectoral fin. There are two vertical crescent-shaped bars just behind the eye highlighted with a
315:
Daffodil cichlids inhabit rocky coastlines and swim in large schools that often consist of hundreds of fish. When breeding, however, they will form monogamous pairs and spawn in caves. They are found in waters at depths of 32 feet (10 m) or deeper. They feed on swarms of plankton drifting in the lake
548:
Breeders living in a large group also benefit because individuals feed more often, have lower workloads, and enjoy greater reproductive success. Helpers in larger groups also feed more frequently but do not have lower workloads. Large groups are also more likely to contain a breeding male and female
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Group size positively influences the survival of group members. Group size correlates positively between years; in one study, 34% of the groups became extinct within one year, but none of the large groups became extinct. Large groups are more likely to contain small-sized helpers the subsequent year
487:
In addition to dispersing to join a new group, subordinates may also gain dominant breeding status by inheriting the top position of their current group. Inheritance happens whenever the dominant individual dies or leaves the group. The pathway to breeder status varies between males and females as a
457:
The degree of multiple paternity in a population of daffodil cichlids is important because sneaking fertilization often results in multiple paternity. The dominant male usually sires all offspring, but not always. When twelve different broods were looked at, it was found that 44.2% of the young were
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among every fish in the group. The breeder males and females are dominant individuals who have offspring, while the helper cichlids are subordinate. Dominance is typically determined by size. The effect of social position was investigated on growth rates and other physical conditions. In the absence
417:
concentrations when paired with subordinates. These results suggest that dominant individuals experience higher cortisol levels as well as higher growth rates since higher cortisol levels in the blood indicate that an individual either has less stress or better internal control when stressed. These
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The daffodil cichlid inhabits permanent social groups composed of one breeding pair and helpers of both sexes. Absolute breeding success (assessed by a combined measure of clutch size and egg survival probability) has been shown to be higher for breeding pairs with helpers compared to those without
307:
Like other
Lamprologini cichlids, daffodil cichlids are highly variable and are found in all kinds of habitats. They are found both at the surface and in very deep waters, but all species are substrate spawners. They have a body that can be somewhat elongated to very elongated. Their colors tend to
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The daffodil cichlid is very active and will swim in all areas of the aquarium. A minimum of 15 gallons is recommended for daffodil cichlids living in a tank with no other species of fish, with 20 or 35 US gallons (76 or 132 L) being better. A larger tank of 50 US gallons (190 L) or more
602:
Paradoxically, monogamous males occupy higher-quality territories with more shelter and fewer predators. Monogamous males also provide more parental care than polygynous males, but the number and survival of young do not vary according to male mating behavior. Females have a trade-off between male
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is regarded as a beneficial mating strategy for males, whereas females often suffer a reduction in pair male contributions. Some males hold only one territory with one breeding female while other males hold multiple territories, each one with its own breeding female. Polygynous males are larger in
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benefits to helpers. When given a choice between associating with unfamiliar kin or unfamiliar non-kin, juvenile daffodil cichlids spend a significantly longer time associating with kin. Relatedness, rather than familiarity, is more important in the association preferences of the daffodil cichlid,
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Some territorial species have been shown to react more aggressively toward unfamiliar conspecifics (members of the same species) than established neighbors, a behavior referred to as the "dear enemy" phenomenon. Laboratory experiments were conducted to determine whether male daffodil cichlids can
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that influences aggression and dominance. Sex differences in androgens have been investigated and show that newly dominant females have higher plasma levels of testosterone than newly subordinate females, but both types of females have similar 11-ketotestosterone levels. Newly dominant males have
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can explain the evolution of cooperative breeding, and the distribution of relatives within a population may influence the benefits of cooperative behavior. Females are more likely to inherit the breeding position of their mother or sister in larger groups. Helper to breeder relatedness decreases
324:
Daffodil cichlids are graceful fish with bodies that are elongated with a continuous dorsal fin. The tail fin is lyre-shaped and they develop long flowing filaments on all unpaired fins. They usually reach up to about 4–5 inches (10–13 cm) in length, but can sometimes get a bit bigger in the
474:
Social status and sex also influence how much daffodil cichlids invest in their livers. Males and females (controlling for body size) have similar liver investment, and subordinates of both sexes have relatively larger livers compared with dominants. Social status results in liver size disparity
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levels, and decreased fitness for subordinates. However, in cooperatively breeding species like the daffodil cichlid, the dominant individuals exhibit higher levels of glucocorticoids, which may be a consequence of the demands of maintaining high social rank and suppressing the reproduction of
287:
has a black stripe running from its eye to its gill cover and a yellow spot just above it, both of which are absent in N. pulcher. Because
Neolamprologus pulcher is the older of the two scientific names, the rules of scientific nomenclature would make this the correct name for the species. The
483:
The social hierarchy is not rigid, as helpers can also become breeders. The most common way for the change is via dispersion into other groups. When helpers disperse by moving into other groups, they can increase their status by becoming a breeder or by moving up in the dominance hierarchy.
440:
than helpers, as well as faster- and longer-swimming sperm and a higher percentage of motile sperm compared to helpers. The sperm of large helpers are characteristically similar to those of breeders, but helpers have smaller testes. The small size of the helpers' testes, coupled with the
540:
The group size of daffodil cichlid helpers is highly variable and typically ranges from one to fourteen individuals, with larger groups living in larger territories. Additionally, group size is more strongly correlated with territory quality than is breeder size.
395:
Since daffodil cichlids are a species that uses cooperative breeding, this means that each individual is ranked in terms of social status. Breeders have a higher social rank than non-breeders. Social status has significant effects on daffodil cichlid behavior.
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not fathered by the dominant male, and multiple paternity was found in 5 of 12 broods (41.7%), with 8 of 35 young (22.9%) being sired by males other than the territory owning breeder males. This is an exceptionally high rate of extra-pair paternity among
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is a very oxygen-rich lake so bubblers need to be going day and night, even if there are plants, to simulate their natural environment to full effect. Nitrates levels should be no more than 25 ppm; daffodil cichlids also can't tolerate a pH less than 7.
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will select for increased reproductive investment by subordinate sneaking males, relative to those of dominant males. This reproductive investment exists in daffodil cichlids in the form of reproductive suppression. Breeders have considerably larger
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opportunities with females due to the existence of a dominance hierarchy in competitive breeders. Studies have suggested that socially subordinate helper males sneak fertilizations from dominant breeding males. If sneaking does happen, then
351:
helpers. Egg size tends to decrease as the number of helpers within groups increases, which suggests that the presence of helpers enables breeder females to strategically reduce their investment per egg in a manner that maximizes breeder
1187:
Stiver, K.A.; P. Dierkes; M. Taborsky; S. Balshine (2004). "Dispersal patterns and status change in a cooperatively breeding cichlid
Neolamprologus pulcher: evidence from microsatellite analyses and behavioural observations".
1341:
Grantner, A.; M. Taborsky (August 1998). "The metabolic rates associated with resting, and with the performance of agonistic, submissive and digging behaviours in the cichlid fish
Neolamprologus pulcher (Pisces: Cichlidae)".
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being very deep and the water tends to stay stable. There should be normal water changes of only 10% to 20% a week, or more frequent small changes depending on the nitrite/ammonia levels and stocking numbers.
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to reduce their investment per egg when more helpers are present. Experiments show that females do indeed reduce egg size with increasing number of helpers but not when perceived neighbor density is high.
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Dierkes, Petra; Dik Heg; Michael
Taborsky; Eva Skubic; Roland Achmann (July 2005). "Genetic relatedness in groups is sex-specific and declines with age of helpers in a cooperatively breeding cichlid".
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size, body-scraped less (they suffered less from ectoparasites), have larger testes (when controlled for body mass), and have higher circulating levels of 11-ketotestosterone than monogamous males.
781:
Balshine, Sigal; Brenda Leach; Francis Neat; Hannah Reid; Michael
Taborsky; Noam Werner (July 2001). "Correlates of group size in a cooperatively breeding cichlid fish (Neolamprologus pulcher)".
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which is advantageous because not all familiar individuals within a cooperatively breeding group are relatives. Having the ability to recognize kin from non-kin brings fitness advantages through
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Daffodil cichlids cannot handle large water changes very well unless the new water chemistry closely matches the water they are in. This inability to tolerate large water changes is due to
1260:
Taves, Matthew D.; Julie K. Desjardins; Sandeep Mishra; Sigal
Balshine (April 2009). "Androgens and dominance: Sex-specific patterns in a highly social fish (Neolamprologus pulcher)".
1218:
Stiver, K.A.; J. Fitzpatrick; J.K. Desjardins; S. Balshine (February 2006). "Sex differences in rates of territory joining and inheritance in a cooperatively breeding cichlid fish".
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bit of blue. The dorsal fin is lyre-shaped and they develop long flowing filaments on all unpaired fins. The fins are tipped with an icy blue. They have brilliant blue eyes.
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In cooperatively breeding groups that have a mix of related and unrelated individuals, being able to identify and differentially cooperate with relatives can bring indirect
1378:
1438:
Bachar, Zina; Lyanne
Brouwer; Dik Heg; Michael Taborsky (2005). "Large group size yields group stability in the cooperatively breeding cichlid Neolamprologus pulcher".
1717:
978:
Buchner, A.S.; K.A. Sloman; S. Balshine (October 2004). "The physiological effects of social status in the cooperatively breeding cichlid
Neolamprologus pulcher".
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Usually, dominant individuals within group-living species are frequently aggressive towards subordinates, and that aggression can lead to chronic stress, higher
1756:
1468:
Frostman, Polly; Peter T. Sherman (August 2004). "Behavioral response to familiar and unfamiliar neighbors in a territorial cichlid, Neolamprologus pulcher".
1810:
901:
Le Vin, A.L.; B.K. Mable; K.E. Arnold (May 2010). "Kin recognition via phenotype matching in a cooperatively breeding cichlid, Neolamprologus pulcher".
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be brown, yellow, blue, black or a combination of all four. Black is usually a striping, either vertical or horizontal. Like other genera in the tribe,
557:
Daffodil cichlids are a very territorial species. They will try to drive out any perceived intruders that wander into their territory, whether it be a
462:
vertebrates, which suggests that male helpers may have produced these young before being expelled from the territory in response to their reproductive
1825:
1691:
1295:
Taborsky, M.; A. Grantner (December 1998). "Behavioural time-energy budgets of cooperatively breeding
Neolamprologus pulcher (Pisces: Cichlidae)".
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after breeding, which is a cumulative measure of the previous months' reproductive success and may explain why large groups do not become extinct.
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steeply with increasing helper age, particularly for breeding males. Helper to helper relatedness is age-associative and also declines with age.
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620:
would be required if mixing with other species. These fish need good water movement along with very strong and efficient filtration.
342:
Daffodil cichlids are omnivorous and feed on swarms of plankton in the water column as well as small crustaceans and invertebrates.
819:
Wong, Marian; Sigal Balshine (2011). "The evolution of cooperative breeding in the African cichlid fish, Neolamprologus pulcher".
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Mileva, Victoria R.; John L. Fitzpatrick; Susan Marsh-Rollo; Kathleen M. Gilmour; Chris M. Wood; Sigal Balshine (November 2009).
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1820:
1735:
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physiological equivalence of their sperm compared to breeder sperm, imply that the helpers are reproductively suppressed.
76:
449:
Mixed parentage is not too uncommon in the daffodil cichlid. Genetic data collected in groups of daffodil cichlids from
292:, a Belgian who set up a collection station, for the export of Tanganyikan cichlids in 1971, named "Fishes of Burundi."
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Desjardins, Julie; John Fitzpatrick; Sigal Balshine; Kelly Stiver; Nicole Mulligan; Glen Van Der Kraak (March 2008).
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1643:
1562:
1513:
1100:
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own social groups, but no subordinate males were found to have fathered any young in any of the examined groups.
284:
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Desjardins, Julie K.; John L. Fitzpatrick; Kelly A. Stiver; Glen J. Van der Kraak; Sigal Balshine (May 2008).
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increased levels of cortisol possibly correlate with size determining rank within the dominance hierarchy.
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1146:
Sopinka, N.M.; J.L. Fitzpatrick; J.K. Desjardins; K.A. Stiver; S.E. Marsh-Rollo; S. Balshine (July 2009).
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levels and grew the most. Dominant fish in groups of breeders and helpers had significantly higher plasma
32:
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756:
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were two distinct species. Now they are considered the same species, the only difference being that
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520:, it was found that pair males invested their energy expenditure almost exclusively in intrafamily
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Fitzpatrick, J.L.; J.K. Desjardins; K.A. Stiver; R. Montgomerie; S. Balshine (October 2005).
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aquarium reaching up to 6 inches (15 cm). They can live 8 – 10 years with proper care.
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1065:"Male reproductive suppression in the cooperatively breeding fish Neolamprologus pulcher"
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1379:"The Stress Response of the Highly Social African Cichlid Neolamprologus pulcher"
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1148:"Liver size reveals social status in the African cichlid Neolamprologus pulcher"
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1016:"Multiple paternity in the cooperatively breeding fish Neolamprologus pulcher"
867:"Mothers adjust egg size to helper number in a cooperatively breeding cichlid"
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where it prefers locations with plenty of sedimentation. The common names for
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water along with microorganisms such as small crustaceans and invertebrates.
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Stiver, K.A.; J.L. Fitzpatrick; J.K. Desjardins; S. Balshine (April 2009).
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daffodil cichlid, when it was still known as N. brichardi, was named after
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The defense behavior of daffodil cichlids has been shown to be linked to
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1514:"Sex and status in a cooperative breeding fish: behavior and androgens"
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1563:"Costs and benefits of polygyny in the cichlid Neolamprologus pulcher"
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In most vertebrates, aggression and dominance are tightly linked to
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259:. This species can reach a length of 10 centimetres (3.9 in)
516:
When metabolic rates for daffodil cichlids were measured using a
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Within a group of breeding and helper cichlids, there exists a
865:
Taborsky, Barbara; Eva Skubic; Rick Bruintjes (March 2007).
504:. Fish, however, do not only have testosterone but also
488:
result of sex differences in the costs of dispersal and
312:
will readily mate with females of other Lamprologini.
594:
Male daffodil cichlids are facultatively polygynous.
1014:
Achmann, R.; P. Dierkes; M. Taborsky (August 2008).
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409:of breeders, dominant individuals had higher liver
1101:"Mixed parentage in Neolamprologus pulcher groups"
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496:Physiological consequences of behavior
1386:Physiological and Biochemical Zoology
1262:General and Comparative Endocrinology
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1788:EED94365-D713-2D87-DAE0-8EACE028FF3C
1811:IUCN Red List least concern species
1521:Behavioral Ecology and Sociobiology
1344:Journal of Comparative Physiology B
1023:Behavioral Ecology and Sociobiology
783:Behavioral Ecology and Sociobiology
653:IUCN Red List of Threatened Species
479:Flexibility in dominance hierarchy
14:
300:Daffodil cichlids are endemic to
275:Previously, it was believed that
1826:Taxa named by Ethelwynn Trewavas
1202:10.1111/j.0022-1112.2004.00427.x
1165:10.1111/j.1095-8649.2009.02234.x
1120:10.1111/j.1095-8649.2009.02173.x
992:10.1111/j.0022-1112.2004.00517.x
950:10.1111/j.1461-0248.2005.00801.x
833:10.1111/j.1469-185x.2010.00158.x
75:
721:Lee, David (27 November 2010).
263:. It can also be found in the
1:
1582:10.1016/j.anbehav.2007.09.037
1232:10.1016/j.anbehav.2005.06.011
915:10.1016/j.anbehav.2010.02.006
426:Mature males may compete for
1274:10.1016/j.ygcen.2008.12.018
532:subordinate group members.
1847:
1452:10.1163/156853905774831891
727:African Rift Lake Cichlids
1533:10.1007/s00265-007-0504-1
1482:10.1007/s10228-004-0223-9
1035:10.1007/s00265-008-0587-3
400:Physiological differences
214:Trewavas & Poll, 1952
207:
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177:
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72:Scientific classification
70:
48:
39:
30:
23:
705:. February 2013 version.
691:; Pauly, Daniel (eds.).
296:Distribution and habitat
285:Neolamprologus brichardi
1470:Ichthyological Research
1190:Journal of Fish Biology
1152:Journal of Fish Biology
1108:Journal of Fish Biology
980:Journal of Fish Biology
642:Bigirimana, C. (2006).
1831:Fish described in 1952
1821:Taxa named by Max Poll
1644:Neolamprologus pulcher
1614:Neolamprologus pulcher
1309:10.1006/anbe.1998.0918
695:Neolamprologus pulcher
646:Neolamprologus pulcher
569:Individual recognition
271:Taxonomy and phylogeny
225:Neolamprologus pulcher
181:Neolamprologus pulcher
25:Neolamprologus pulcher
1356:10.1007/s003600050162
1082:10.1093/beheco/ari090
884:10.1093/beheco/arm026
795:10.1007/s002650100343
660:: e.T60604A12382292.
422:Competition for mates
1446:(11–12): 1615–1641.
723:"Brichardi Cichlids"
460:cooperative breeding
346:Cooperative breeding
506:11-ketotestosterone
406:dominance hierarchy
211:Lamprologus pulcher
42:Conservation status
1069:Behavioral Ecology
871:Behavioral Ecology
821:Biological Reviews
757:"Daffodil Cichlid"
578:Effect of hormones
522:agonistic behavior
445:Multiple paternity
253:princess of Zambia
1798:
1797:
1770:Open Tree of Life
1606:Taxon identifiers
1029:(10): 1581–1589.
755:Brough, Clarice.
553:Territory defense
433:sperm competition
369:fitness (biology)
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1576:(5): 1771–1779.
1570:Animal Behaviour
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1303:(6): 1375–1382.
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1220:Animal Behaviour
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1114:(5): 1129–1135.
1105:
1096:
1087:
1086:
1084:
1060:
1047:
1046:
1020:
1011:
996:
995:
986:(4): 1080–1095.
975:
962:
961:
933:
927:
926:
909:(5): 1109–1114.
903:Animal Behaviour
898:
889:
888:
886:
862:
853:
852:
816:
807:
806:
778:
765:
764:
752:
731:
730:
718:
707:
706:
685:
679:
678:
676:
674:
669:
639:
470:Liver investment
257:lyretail cichlid
245:daffodil cichlid
228:is a species of
213:
183:
80:
79:
59:
54:
53:
35:
21:
1846:
1845:
1841:
1840:
1839:
1837:
1836:
1835:
1801:
1800:
1799:
1794:
1786:
1781:
1773:
1768:
1760:
1755:
1747:
1742:
1734:
1729:
1721:
1716:
1708:
1703:
1695:
1690:
1682:
1677:
1669:
1664:
1656:
1651:
1642:
1641:
1636:
1627:
1626:
1621:
1608:
1598:
1597:
1565:
1560:
1559:
1548:
1516:
1511:
1510:
1497:
1467:
1466:
1459:
1437:
1436:
1421:
1381:
1376:
1375:
1371:
1340:
1339:
1332:
1294:
1293:
1289:
1259:
1258:
1247:
1217:
1216:
1209:
1186:
1185:
1181:
1145:
1144:
1135:
1103:
1098:
1097:
1090:
1062:
1061:
1050:
1018:
1013:
1012:
999:
977:
976:
965:
938:Ecology Letters
935:
934:
930:
900:
899:
892:
864:
863:
856:
818:
817:
810:
780:
779:
768:
754:
753:
734:
720:
719:
710:
687:
686:
682:
672:
670:
641:
640:
636:
631:
622:Lake Tanganyika
613:Lake Tanganyika
609:
607:In the aquarium
592:
580:
571:
555:
538:
498:
481:
472:
451:Lake Tanganyika
447:
424:
402:
393:
365:
363:Kin recognition
348:
340:
335:
322:
302:Lake Tanganyika
298:
290:Pierre Brichard
273:
237:Lake Tanganyika
196:
185:
179:
166:
163:N. pulcher
74:
66:
55:
51:
44:
17:
16:Species of fish
12:
11:
5:
1844:
1842:
1834:
1833:
1828:
1823:
1818:
1816:Neolamprologus
1813:
1803:
1802:
1796:
1795:
1793:
1792:
1779:
1766:
1753:
1740:
1727:
1714:
1701:
1688:
1675:
1662:
1649:
1634:
1618:
1616:
1610:
1609:
1604:
1596:
1595:
1546:
1527:(5): 785–794.
1495:
1476:(3): 283–285.
1457:
1419:
1398:10.1086/605937
1392:(6): 720–729.
1369:
1350:(6): 427–433.
1330:
1287:
1268:(2): 202–207.
1245:
1226:(2): 449–456.
1207:
1179:
1133:
1088:
1048:
997:
963:
944:(9): 968–975.
928:
890:
877:(4): 652–657.
854:
827:(2): 511–530.
808:
789:(2): 134–140.
766:
732:
708:
689:Froese, Rainer
680:
633:
632:
630:
627:
608:
605:
591:
588:
579:
576:
570:
567:
563:heterospecific
554:
551:
537:
534:
529:glucocorticoid
497:
494:
480:
477:
471:
468:
446:
443:
423:
420:
401:
398:
392:
389:
364:
361:
347:
344:
339:
336:
334:
331:
321:
318:
297:
294:
272:
269:
219:
218:
217:
216:
205:
204:
198:
197:
186:
175:
174:
168:
167:
160:
158:
154:
153:
150:Neolamprologus
146:
142:
141:
136:
132:
131:
126:
122:
121:
119:Actinopterygii
116:
112:
111:
106:
102:
101:
96:
92:
91:
86:
82:
81:
68:
67:
49:
46:
45:
40:
37:
36:
28:
27:
15:
13:
10:
9:
6:
4:
3:
2:
1843:
1832:
1829:
1827:
1824:
1822:
1819:
1817:
1814:
1812:
1809:
1808:
1806:
1789:
1784:
1780:
1776:
1771:
1767:
1763:
1758:
1754:
1750:
1745:
1741:
1737:
1732:
1728:
1724:
1719:
1715:
1711:
1706:
1702:
1698:
1693:
1689:
1685:
1680:
1676:
1672:
1667:
1663:
1659:
1654:
1650:
1645:
1639:
1635:
1630:
1624:
1620:
1619:
1617:
1615:
1611:
1607:
1602:
1591:
1587:
1583:
1579:
1575:
1571:
1564:
1557:
1555:
1553:
1551:
1547:
1542:
1538:
1534:
1530:
1526:
1522:
1515:
1508:
1506:
1504:
1502:
1500:
1496:
1491:
1487:
1483:
1479:
1475:
1471:
1464:
1462:
1458:
1453:
1449:
1445:
1441:
1434:
1432:
1430:
1428:
1426:
1424:
1420:
1415:
1411:
1407:
1403:
1399:
1395:
1391:
1387:
1380:
1373:
1370:
1365:
1361:
1357:
1353:
1349:
1345:
1337:
1335:
1331:
1326:
1322:
1318:
1314:
1310:
1306:
1302:
1298:
1291:
1288:
1283:
1279:
1275:
1271:
1267:
1263:
1256:
1254:
1252:
1250:
1246:
1241:
1237:
1233:
1229:
1225:
1221:
1214:
1212:
1208:
1203:
1199:
1196:(1): 91–105.
1195:
1191:
1183:
1180:
1175:
1171:
1166:
1161:
1157:
1153:
1149:
1142:
1140:
1138:
1134:
1129:
1125:
1121:
1117:
1113:
1109:
1102:
1095:
1093:
1089:
1083:
1078:
1074:
1070:
1066:
1059:
1057:
1055:
1053:
1049:
1044:
1040:
1036:
1032:
1028:
1024:
1017:
1010:
1008:
1006:
1004:
1002:
998:
993:
989:
985:
981:
974:
972:
970:
968:
964:
959:
955:
951:
947:
943:
939:
932:
929:
924:
920:
916:
912:
908:
904:
897:
895:
891:
885:
880:
876:
872:
868:
861:
859:
855:
850:
846:
842:
838:
834:
830:
826:
822:
815:
813:
809:
804:
800:
796:
792:
788:
784:
777:
775:
773:
771:
767:
762:
758:
751:
749:
747:
745:
743:
741:
739:
737:
733:
728:
724:
717:
715:
713:
709:
704:
703:
698:
696:
690:
684:
681:
668:
663:
659:
655:
654:
649:
647:
638:
635:
628:
626:
623:
617:
614:
606:
604:
600:
597:
590:Mating system
589:
587:
585:
577:
575:
568:
566:
564:
560:
552:
550:
546:
542:
535:
533:
530:
525:
523:
519:
514:
511:
507:
503:
495:
493:
491:
485:
478:
476:
469:
467:
465:
461:
455:
452:
444:
442:
439:
434:
429:
428:fertilization
421:
419:
416:
412:
407:
399:
397:
391:Social status
390:
388:
385:
384:Kin selection
381:
379:
375:
374:kin selection
370:
362:
360:
356:
354:
345:
343:
337:
332:
330:
326:
319:
317:
313:
311:
305:
303:
295:
293:
291:
286:
282:
278:
270:
268:
266:
262:
258:
254:
250:
249:fairy cichlid
246:
242:
238:
234:
231:
227:
226:
212:
209:
208:
206:
203:
199:
194:
190:
184:
182:
176:
173:
172:Binomial name
169:
165:
164:
159:
156:
155:
152:
151:
147:
144:
143:
140:
137:
134:
133:
130:
127:
124:
123:
120:
117:
114:
113:
110:
107:
104:
103:
100:
97:
94:
93:
90:
87:
84:
83:
78:
73:
69:
63:
58:
57:Least Concern
47:
43:
38:
34:
29:
26:
22:
19:
1613:
1573:
1569:
1524:
1520:
1473:
1469:
1443:
1439:
1389:
1385:
1372:
1347:
1343:
1300:
1296:
1290:
1265:
1261:
1223:
1219:
1193:
1189:
1182:
1155:
1151:
1111:
1107:
1075:(1): 25–33.
1072:
1068:
1026:
1022:
983:
979:
941:
937:
931:
906:
902:
874:
870:
824:
820:
786:
782:
761:Animal-World
760:
726:
700:
694:
683:
671:. Retrieved
657:
651:
645:
637:
618:
610:
601:
593:
581:
572:
556:
547:
543:
539:
526:
518:respirometer
515:
502:testosterone
499:
486:
482:
473:
456:
448:
425:
403:
394:
382:
366:
357:
349:
341:
327:
323:
314:
309:
306:
299:
281:N. brichardi
280:
276:
274:
256:
252:
248:
244:
240:
224:
223:
222:
210:
180:
178:
162:
161:
149:
129:Cichliformes
24:
18:
1705:iNaturalist
1638:Wikispecies
1158:(1): 1–16.
673:16 November
559:conspecific
380:avoidance.
320:Description
1805:Categories
629:References
508:, another
490:inbreeding
464:parasitism
378:inbreeding
310:N. pulcher
277:N. pulcher
241:N. pulcher
1440:Behaviour
584:androgens
157:Species:
139:Cichlidae
95:Kingdom:
89:Eukaryota
1723:10154109
1679:FishBase
1629:Q1756298
1623:Wikidata
1590:29101068
1490:25786675
1414:24045567
1406:19807269
1364:23177724
1325:17466822
1282:19174165
1240:33618947
1174:20738479
1128:20735623
958:34517681
923:53152164
849:39910620
841:20849492
803:29900603
702:FishBase
596:Polygyny
536:Grouping
510:androgen
415:cortisol
411:glycogen
333:Behavior
265:aquarium
243:include
202:Synonyms
189:Trewavas
135:Family:
109:Chordata
105:Phylum:
99:Animalia
85:Domain:
62:IUCN 3.1
1697:2371813
1541:7980302
1317:9933533
1043:5648274
353:fitness
338:Feeding
267:trade.
233:endemic
230:cichlid
195:, 1952)
145:Genus:
125:Order:
115:Class:
60: (
1775:280185
1762:184151
1736:648820
1710:106777
1671:205342
1588:
1539:
1488:
1412:
1404:
1362:
1323:
1315:
1280:
1238:
1172:
1126:
1041:
956:
921:
847:
839:
801:
438:testes
191:&
1783:Plazi
1749:60604
1718:IRMNG
1658:46K3T
1586:S2CID
1566:(PDF)
1537:S2CID
1517:(PDF)
1486:S2CID
1410:S2CID
1382:(PDF)
1360:S2CID
1321:S2CID
1236:S2CID
1104:(PDF)
1039:S2CID
1019:(PDF)
919:S2CID
845:S2CID
799:S2CID
561:or a
1757:NCBI
1744:IUCN
1731:ITIS
1692:GBIF
1684:8752
1402:PMID
1313:PMID
1278:PMID
1170:PMID
1124:PMID
954:PMID
837:PMID
675:2021
658:2006
376:and
279:and
255:and
193:Poll
1666:EoL
1653:CoL
1578:doi
1529:doi
1478:doi
1448:doi
1444:142
1394:doi
1352:doi
1348:168
1305:doi
1270:doi
1266:161
1228:doi
1198:doi
1160:doi
1116:doi
1077:doi
1031:doi
988:doi
946:doi
911:doi
879:doi
829:doi
791:doi
662:doi
235:to
1807::
1785::
1772::
1759::
1746::
1733::
1720::
1707::
1694::
1681::
1668::
1655::
1640::
1625::
1584:.
1574:75
1572:.
1568:.
1549:^
1535:.
1525:62
1523:.
1519:.
1498:^
1484:.
1474:51
1472:.
1460:^
1442:.
1422:^
1408:.
1400:.
1390:82
1388:.
1384:.
1358:.
1346:.
1333:^
1319:.
1311:.
1301:56
1299:.
1276:.
1264:.
1248:^
1234:.
1224:71
1222:.
1210:^
1194:65
1192:.
1168:.
1156:75
1154:.
1150:.
1136:^
1122:.
1112:74
1110:.
1106:.
1091:^
1073:17
1071:.
1067:.
1051:^
1037:.
1027:62
1025:.
1021:.
1000:^
984:65
982:.
966:^
952:.
940:.
917:.
907:79
905:.
893:^
875:18
873:.
869:.
857:^
843:.
835:.
825:86
823:.
811:^
797:.
787:50
785:.
769:^
759:.
735:^
725:.
711:^
699:.
656:.
650:.
565:.
466:.
355:.
261:TL
251:,
247:,
1592:.
1580::
1543:.
1531::
1492:.
1480::
1454:.
1450::
1416:.
1396::
1366:.
1354::
1327:.
1307::
1284:.
1272::
1242:.
1230::
1204:.
1200::
1176:.
1162::
1130:.
1118::
1085:.
1079::
1045:.
1033::
994:.
990::
960:.
948::
942:8
925:.
913::
887:.
881::
851:.
831::
805:.
793::
763:.
729:.
697:"
693:"
677:.
664::
648:"
644:"
187:(
64:)
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