In most species of fish, when breeding occurs territories are not formed and eggs not guarded, but released and fertilized without any subsequent care. To be able to maintain a stable population, some species produce thousands or even tens of thousands of eggs with the hope that some will develop into adult fish. Characins for example spawn over plants or over course ground cover so that eggs obtain some sort of protection till the larvae become mobile. Eggs of many marine fish species float and can thus be transported over long distances in the oceans.
Cichlids, however, have parental care protecting eggs, wrigglers and fry, and many species develop sexual dimorphism and establish territories. This extreme investment in their progeny pays off, as they don’t have to produce so many eggs to maintain stable and resilient populations.
The breeding season
For optimum results, the breeding season for cichlids is limited to the very best time of the year in regards to food availability, higher temperature, slower water flow, water clearness which allows them to maintain visual contact with the fry, and, most importantly, enough food for individuals to get into breeding condition and for their fry to develop as quickly as possible.
For cichlids and other fish, it is also important to synchronize spawning so that the sheer number of fry overwhelms predators which then won’t be able to get them all. For example, Etroplus maculatus, the orange chromide, in Sry Lanka was shown that pairs which reproduced outside the time when most pairs did were not as successful in bringing up their fry (Ward et al., 1981). Some events like the phases of the moon help in achieving finer synchronization (Nakai et al., 1990). In this particular experiment the authors proposed that such synchronization “suggested that the nocturnal guarding efficiency of eggs, especially exposed ones, is improved by the maximal amount of moonlight during full moon and, in species whose young leave the breeding sites about two weeks after spawning, the survival of dispersing young is enhanced by the darkness of moonless nights”.
Another interpretation (Knowlton, 1979) is that breeding synchronization prevents males from looking for another breeding partner and hence abandon breeding efforts, since all females are occupied.
While in some cichlid species breeding takes place twice a year, most species would only attempt to breed once, but there will always be isolated pairs which breed out of synchrony with the rest of the population.
For some species of cichlids, the breeding period is limited to the dry season prior to rainfall; this is the case for most Central American cichlids. There is one exception though: stable environments with a year round warm temperature and availability of food may sustain breeding pairs all year long, even if they may peek at a certain period. This is the case for example for Herichthys minckleyi populations inhabiting thermal springs in the Cuatro Cienegas valley in northern Mexico, and for both Herichthys bartoni and H. labridens in the thermal springs in the Valle of Rioverde in San Luis Potosi [Mexico]. Interestingly enough, Herichthys carpintis, native to the Panuco drainage but exotic in a few thermal springs in the later valley, are able to imitate their relatives and even if most of their breeding efforts happen in April as in most of their range, a notable number of pairs are also seen during the rest of the year. One may be tempted to generalize this phenomenon and think that probably this is the case for all cichlids inhabiting springs and lakes, but it is not.
In the numerous clear water cenotes in the eastern part of the Yucatan peninsula, inhabited by several cichlid species (although not all species in each Cenote), breeding happens in an established breeding season, although interestingly it does not coincide with the season for their populations dwelling in rivers. For example, even if the breeding season of Parachromis multifasciatus and Rocio octofasciata normally peaks in April, I have witnessed that in Cenote Escondido they prefer to breed in September or October, while for Vieja melanurus and Cribroheros robertsoni in the same cenote, I have only found pairs during July or August. Why this is the case, is to my knowledge not yet researched, but given the small size of the habitat it may be due to the limited availability of suitable breeding places.
In the Nicaraguan crater lakes, something similar happens, with the bulk of the breeding efforts for the cichlid species living there occurring in December, which does not overlap with that of cichlid species in rivers in the area, which mostly breed during the spring. Cichlid breeding in lakes after the rainy season in Nicaragua, which extends from May to October, has logic, since rain runoffs bring into the lakes a significant amount of food and nutrients that are necessary for the growth of plankton and algae, from which both parents and fry benefit.
In the Amazon basin in South America, breeding comes when rains start in October or November, where the rivers expand significantly and floods big extensions of forest. This assures that fry are large enough when the dry season comes and the consequential decrease in food sources occur. An example of this is Symphysodon aequifasciatus, whose life history has been studied (Crampton, 2008).
In Sri Lanka, the orange chromide, Etroplus maculatus, and the green chromide, Etroplus suratensis, have breeding tactics which have been studied by Ward & Samarakoon (1981). They observed that these species reproduce twice during the year when water conditions are favorable for nest construction and maintaining visual contact with offspring, which happens during the drier pre-monsoonal and monsoonal seasons from May through September, when water turbidity decreased and salinity increased. Breeding activity of both species peaked in July.
In Lake Malawi, Lance Smith (2000) studied the reproductive biology of the open water Copadichromis chrysonotus and determined that breeding was mostly continuous (with a couple of peaks) from August to May, which corresponds roughly to the spring and summer in the southern hemisphere, with a rainy season from mid-November to April. This seems roughly the case for other Malawi cichlid species as well, which may benefit from nutrient run-offs into the lake.
How many mates?
For the purpose of obtaining enough descendants to maintain a stable population, cichlids have evolved several breeding strategies that cover a wide range of behavior schemes known to vertebrates, from monogamy to the extended family. Here is list and definition for each one of them.
Monogamy in cichlids is widely used, particularly when the environmental conditions require a close and strong guard of eggs and fry, but it can relax if conditions are favorable. When I write monogamy I refer to a breeding effort or season, not paralleling the life-long monogamous life of many birds. Take for example Amatitlania nigrofaciata, the convict cichlid, whose males rarely desert females in the care of their babies, especially in a highly stressful situation, when fry guarded by just the female would be less likely to reach adulthood. When the environmental conditions are favorable, males are known, however, to desert brooding females. Those conditions are associated with low fry predation, reduced number of babies to take care of and proximity of babies to independence.
Monogamous cichlids rarely exhibit a strong sexual dimorphism, since males and females rely on other cues to mate.
Polygyny refers to the case when one male fertilizes the eggs of more than one female, which happens when the care of the eggs (and sometimes the fry) can be successfully carried out by the female alone.
Having as sister species Herichthys cyanoguttatus, the Texas cichlid, a monogamous North American cichlid species, the Cuatro Cienegas cichlid Herichthys minckleyi, which most likely inherited that behavior has mostly departed from monogamy in the Cuatro Cienegas valley springs. Males protect territories in which several females spawn and from where females and fry are subsequently expelled by the male, with the lone female raising the fry among vegetated areas (Artigas Azas, 2015). Clearly, the cooperation of the male is not necessary in raising the young as likely it inhabits in less competition and stressful conditions than those faced by H. cyanoguttatus in the rivers. A polygynous breeding strategy is then a more productive reproductive alternative producing more fry.
All Apistogramma species in South America utilize a similar strategy. Polygynous cichlids tend to exhibit a strong sexual dimorphism since the single male has to attract females to his territory in competition with many other males who simultaneously try to do the same. The beautiful colors and showy unpaired fins of Apistogramma males, which contrast with the modest coloration and short fins in females are an example of this.
Many mouth-brooding cichlids use a similar reproductive strategy. Think of mbuna cichlids in Lake Malawi where males establish and defend a territory where females come to spawn and have their eggs fertilized, and then leave to, sometimes, spending their brooding time in big groups that protect them. When females release their fry almost simultaneously the big number of them overwhelms predators.
Some mouth-brooding females, like those in the Malawi cichlid Fossorochromis rostratus extend parental care as long as or longer than what is provided by monogamous cichlids. Fry are cared for till they become so large that at one point they can’t all fit anymore in the mother’s mouth. Other mouth-brooding females, such as those of many rock-dwelling mbuna from Lake Malawi, for example Metriaclima lombardoi, immediately abandon their fry once incubation time is over, which happens about three and a half weeks after spawning.
In this breeding technique, a male fertilizes the eggs of two or more females, and a female gets her eggs fertilized by two or more males.
The school-forming Lake Tanganyika cichlid species Cyprichromis coloratus and C. leptosoma, which form three dimensional territories and spawn in the open water, are examples of this breeding strategy. In a study carried out by Anderson, Werdenig, Koblmüller & Sefc (2016) using DNA microsatellite markers, it was demonstrated that the eggs of 4 out of 12 examined brooding females of C. coloratus and 18 of 22 broods of C. leptosoma had multiple paternity, establishing these species as polygynandrous.
Another example is the case of the feather fins cichlid Ophthalmotilapia ventralis from Lake Tanganyika. Breeding males of O. ventralis maintain territories and most of them defend a rather small patch on a large rock in the upper reaches of the rocky habitat, where they build a small sand bower on top of the rock. Each male then vigorously courts females when they visit his territory, and those females seduced try to pick up the yellow lappets located at the tip of the male’s large pectoral fins, confusing them with her own eggs. The lappets are placed by the male at the center of the bower where he previously released some sperm, as this happens she gets her mouth-brood eggs fertilized. She then takes one of two courses of action, either she leaves the male’s territory looking for another male, or she stays and deposits one or more eggs which she immediately picks up (Konings, 2014), probably to be fertilized by another male.
In at least some Lake Malawi mbuna species, such as Metriaclima zebra, polygyny is extended to polygynandry by having females getting their eggs fertilized by several males (Parker et al., 1996).
Immler & Taborsky (2009) propose as the most likely explanation for the benefits of this behavior what is known as “sperm shopping” where sperm actually competes to fertilize the available eggs, so all eggs are properly fertilized by the best sperm.
In polyandry, one male fertilizes only one female’s eggs, and one female gets her eggs fertilized by more than one male. This may be the rarest of the mating systems.
Polyandry in cichlids has just been documented in some Julidochromis and Chalinochromis species in Lake Tanganyika (Konings, 2019:180). In Julidochromis marlieri and J. regani for example, males are almost always smaller than females, usually considerably. In J. marlieri, it is a common place situation that a larger female presides over the territory of two smaller males, which carry out the parental care of the eggs and fry. Females then spawn with two males and males just fertilize the eggs of a single female. With some variants this may be the case for all species in both mentioned genera.
In this strategy, such as in J. marlieri, the benefit for the female is that the smaller males perform almost all the brood care, allowing her more time to feed. The benefit for the smaller beta male is that he can fertilize a spawn even if he is not large enough to be an alpha male (which at one point he will become). The benefit for the alpha male is less clear, but it can be related to the fact that he will have the help of the beta male in guarding the fry, which will have a higher survival rate (Konings, 2019:180).
The extended family concept refers to species where two or more group members of both sexes reproduce, with some offspring remaining in the family.
A well-known example of this behavior is seen in Neolamprologus pulcher, a Lake Tanganyika endemic about which hundreds of research papers have been published. N. pulcher inhabits the upper layer of the rocky habitat, where monogamous pairs establish territories and breed on a regular basis. Some of the fry produced remain in the parent’s territory becoming helpers which take over part of the territorial defense, while some of the fry establish their own breeding pairs within the territory, which eventually grow into a colony in which every member is involved in the territorial defense against intruders.
You now have had a glimpse into one of the more fascinating aspects of the family Cichlidae, and in this article I have just scratched the surface of the many peculiarities of these wonderful animals. As always, for a more profound approach to these wonderful topics, I recommend the fascinating book “The Cichlid Fishes – Nature’s Grand Experiment in Evolution” by George Barlow, chapter four: mating games.
- Anderson, Caleb & A. Werdenig, S. Koblmüller, K.M. Sefc. 2016. "Same school, different conduct: rates of multiple paternity vary within a mixed-species breeding school of semi-pelagic cichlid fish (Cyprichromis spp.)". Ecology and Evolution. v. 6 (no. 1); pp. 37-45. DOI: 10.1002/ece3.1856 (crc07015) (摘要)
- Artigas Azas, Juan Miguel. 2015. "The desert cichlid: Herichthys minckleyi, part 2". Cichlid News Magazine. v. 24(n. 3), pp. 30-35 (crc06777)
- Crampton, William G. R. . 2008. "Ecology and life history of an Amazon floodplain cichlid: the discus fish Symphysodon (Perciformes: Cichlidae)". Neotropical Ichthyology. 10.1590/S1679-62252008000400008 (crc02522) (摘要)
- Immler, Simone & M. Taborsky. 2009. "Sequential polyandry affords post-mating sexual selection in the mouths of cichlid females". Behavioral Ecology and Sociobiology. v. 63(n. 8), pp. 1219-1230. DOI: 10.1007/s00265-009-0744-3 (crc03703) (摘要)
- Knowlton, Nancy. 1979. "Reproductive synchrony, parental investment, and the evolutionary dynamics of sexual selection". Animal Behaviour. (n. 27), pp. 1022-1033. DOI: 10.1016/0003-3472(79)90049-6 (crc04562) (摘要)
- Konings, Ad. 2019. "Tanganyika cichlids in their natural habitat, 4th edition". Cichlid Press. pp. 1-432. ISBN: 978-932892-26-0 (crc09277) (摘要)
- Konings, Ad. 2014. "Species profile: Ophthalmotilapia ventralis (Boulenger, 1898)". The Cichlid Room Companion. 检索 17- 4月-2021, 从: https://cichlidae.com/species.php?id=688 (crc10538) (摘要)
- Nakai, K. & Y. Yanagisawa, T. Sato, Y. Niimura & M. Gashagaza. 1990. "Lunar synchronization of spawning in cichlid fishes of the tribe Lamprologini in Lake Tanganyika". Journal of Fish Biology. v. 37(n. 4), pp. 589-598. DOI: 10.1111/j.1095-8649.1990.tb05891.x (crc03786) (摘要)
- Parker, Alex & I. Kornfield. 1996. "Polygynandry in Pseudotropheus zebra, a cichlid fish from Lake Malawi". Environmental Biology of Fishes. v. 47(n. 4), pp. 345-352. DOI: 10.1007/BF00005049 (crc05422) (摘要)
- Smith, Lance W. 2000. "The reproductive biology of an open-water spawning Lake Malawi cichlid, Copadichromis chrysonotus". African Zoology. v. 35(n. 2), pp. 151-164 (crc05541) (摘要)
- Ward, Jack A & Jayampathy I. Samarakoon. 1981. "Reproductive tactics of the Asian cichlids of the genus Etroplus in Sri Lanka". Environmental Biology of Fishes. v. 6(1), pp. 95-103. DOI: 10.1007/BF00001803 (crc02617) (摘要)
- Wisenden, Brian D. 1994. "Factors Affecting Mate Desertion by Males in Free-Ranging Convict Cichlids (Cichlasoma nigrofasciatum)". International Society for Behavioral Ecology. v.5(4), pp. 439-447. DOI: 10.1093/beheco/5.4.439 (crc01820) (摘要)
© Copyright 2021 Juan Miguel Artigas Azas, all rights reserved
Artigas Azas, Juan Miguel. (4月 17, 2021). "Spawning strategies in cichlids part 1 – Mating seasons and patterns". Cichlid Room Companion. 检索 6月 21, 2021, 从: https://cichlidae.com/section.php?id=318&lang=cn.