Editing Cichlid (section)

===Brood care===
{{Further|List of fish species that protect their young}}

====Pit spawning in cichlids====
Pit spawning, also referred to as substrate breeding, is a behavior in cichlid fish in which a fish builds a pit in the sand or ground, where a pair court and consequently spawn.<ref name=Alonso-2011>{{cite journal |last1=Alonso |first1=Felipe |last2=Cánepa |first2=Maximiliano |last3=Moreira |first3=Renata Guimarães |last4=Pandolfi |first4=Matías |date=22 July 2011 |title=Social and reproductive physiology and behavior of the Neotropical cichlid fish Cichlasoma dimerus under laboratory conditions |journal=Neotropical Ichthyology |volume=9 |issue=3 |pages=559–570 |doi=10.1590/S1679-62252011005000025 |doi-access=free |hdl=20.500.12110/paper_16796225_v9_n3_p559_Alonso |hdl-access=free }}</ref> Many different factors go into this behavior of pit spawning, including female choice of the male and pit size, as well as the male defense of the pits once they are dug in the sand.<ref name=Nelson-1995>{{cite journal |last=Nelson |first=C. Mindy |date=1 January 1995 |title=Male size, spawning pit size and female mate choice in a lekking cichlid fish |journal=Animal Behaviour |volume=50 |issue=6 |pages=1587–1599 |doi=10.1016/0003-3472(95)80013-1 |s2cid=54249065 |doi-access=free }}</ref>

Cichlids are often divided into two main groups: mouthbrooders and substrate brooders. Different parenting investment levels and behaviors are associated with each type of reproduction.<ref name=Duponchelle-2008>{{cite journal |last1=Duponchelle |first1=Fabrice |last2=Paradis |first2=Emmanuel |last3=Ribbink |first3=Anthony J. |last4=Turner |first4=George F. |title=Parallel life history evolution in mouthbrooding cichlids from the African Great Lakes |journal=Proceedings of the National Academy of Sciences |date=7 October 2008 |volume=105 |issue=40 |pages=15475–15480 |doi=10.1073/pnas.0802343105 |pmid=18824688 |pmc=2563094 |bibcode=2008PNAS..10515475D |doi-access=free }}</ref> As pit spawning is a reproductive behavior, many different physiological changes occur in the cichlid while this process is occurring that interfere with social interaction.<ref name=Brown-1978>{{cite journal |last1=Brown |first1=Dean H. |last2=Marshall |first2=Joseph A. |title=Reproductive Behaviour of the Rainbow Cichlid, Herotilapia Multispinosa (Pisces, Cichlidae) |journal=Behaviour |date=1978 |volume=67 |issue=3–4 |pages=299–321 |doi=10.1163/156853978X00378 }}</ref> Different kinds of species that pit spawn, and many different morphological changes  occur because of this behavioral experience.<ref name=Alonso-2011/>

Pit spawning is an evolved behavior across the cichlid group. Phylogenetic evidence from cichlids in Lake Tanganyika could be helpful in uncovering the evolution of their reproductive behaviors.<ref name=Muschik-2012>{{cite journal |last1=Muschick |first1=Moritz |last2=Indermaur |first2=Adrian |last3=Salzburger |first3=Walter |title=Convergent Evolution within an Adaptive Radiation of Cichlid Fishes |journal=Current Biology |date=December 2012 |volume=22 |issue=24 |pages=2362–2368 |doi=10.1016/j.cub.2012.10.048 |pmid=23159601 |s2cid=18363916 |doi-access=free |bibcode=2012CBio...22.2362M }}</ref> Several important behaviors are associated with pit spawning, including parental care, food provisioning,<ref name=Wisenden-1995>{{cite journal |last1=Wisenden |first1=Brian D. |last2=Lanfranconi-Izawa |first2=Tanya L. |last3=Keenleyside |first3=Miles H.A. |title=Fin digging and leaf lifting by the convict cichlid, Cichlasoma nigrofasciatum: examples of parental food provisioning |journal=Animal Behaviour |date=March 1995 |volume=49 |issue=3 |pages=623–631 |doi=10.1016/0003-3472(95)80195-2 |s2cid=41529998 }}</ref> and brood guarding.<ref name=Ota-2014>{{cite journal |last1=Ota |first1=Kazutaka |last2=Kohda |first2=Masanori |title=Maternal Food Provisioning in a Substrate-Brooding African Cichlid |journal=PLOS ONE |date=9 June 2014 |volume=9 |issue=6 |pages=e99094 |doi=10.1371/journal.pone.0099094 |pmid=24911060 |pmc=4049616 |bibcode=2014PLoSO...999094O |doi-access=free }}</ref>

====Mouth brooding vs. pit spawning====
One of the differences studied in African cichlids is reproductive behavior. Some species pit spawn and some are known as mouth brooders. Mouthbrooding is a reproductive technique where the fish scoop up eggs and fry for protection.<ref name=Duponchelle-2008/> While this behavior differs from species to species in the details, the general basis of the behavior is the same. Mouthbrooding also affects how they choose their mates and breeding grounds. In a 1995 study, Nelson found that in pit-spawning females choose males for mating based on the size of the pit that they dig, as well as some of the physical characteristics seen in the males.<ref name=Nelson-1995/> Pit spawning also differs from mouth brooding in the size and postnatal care  exhibited. Eggs that have been hatched from pit-spawning cichlids are usually smaller than those of mouthbrooders. Pit-spawners' eggs are usually around 2&nbsp;mm, while mouthbrooders are typically around 7&nbsp;mm. While different behaviors take place postnatally between mouthbrooders and pit spawners,  some similarities exist. Females in  both mouthbrooders and pit-spawning cichlids take care of their young after they are hatched. In some cases, both parents exhibit care, but the female always cares for the eggs and newly hatched fry.<ref>{{cite journal |last1=Sefc |first1=Kristina M. |date=21 July 2011 |title=Mating and parental care in Lake Tanganyika's cichlids |journal=International Journal of Evolutionary Biology |volume=2011 |page=470875 |doi=10.4061/2011/470875 |pmid=21822482 |pmc=3142683 |doi-access=free }}</ref>

====Pit spawning process====

Many species of cichlids use pit spawning, but one of the less commonly studied species that exhibits this behavior is the Neotropical ''Cichlasoma dimerus''. This fish is a substrate breeder that displays biparental care after the fry have hatched from their eggs. One study<ref name=Alonso-2011/> examined reproductive and social behaviors  of this species to see how they accomplished their pit spawning, including different physiological factors such as hormone levels, color changes, and plasma cortisol levels. The entire spawning process could take about 90&nbsp;minutes and 400~800&nbsp;eggs could be laid. The female deposits about 10&nbsp;eggs at a time, attaching them to the spawning surface, which may be a pit constructed on the substrate or another surface. The number of eggs laid was correlated to the space available on the substrate. Once the eggs were attached, the male swam over the eggs and fertilized them. The parents would then dig pits in the sand, 10–20&nbsp;cm wide and 5–10&nbsp;cm deep, where larvae were transferred after hatching. Larvae began swimming 8 days after fertilization and parenting behaviors and some of the physiological factors measured changed.

====Color changes====
In the same study, color changes were present before and after the pit spawning occurred. For example, after the larvae were transferred and the pits were beginning to be protected, their fins turned a dark grey color.<ref name=Alonso-2011/> In another study, of the rainbow cichlid, ''Herotilapia multispinosa'',<ref name=Brown-1978/> color changes occurred throughout the spawning process. Before spawning, the rainbow cichlid was an olive color with grey bands. Once spawning behaviors started, the body and fins of the fish became a more golden color. When the eggs were finished being laid, the pelvic fin all the way back to the caudal fin turned to a darker color and blackened in both the males and the females.<ref name=Brown-1978/>

====Pit sizes====
Females prefer a bigger pit size when choosing where to lay eggs.<ref name=Nelson-1995/> Differences are seen in the sizes of pits that created, as well as a change in the morphology of the pits.<ref name=York-2015>{{cite journal |last1=York |first1=Ryan A. |last2=Patil |first2=Chinar |last3=Hulsey |first3=C. Darrin |last4=Streelman |first4=J. Todd |last5=Fernald |first5=Russell D. |title=Evolution of bower building in Lake Malawi cichlid fish: phylogeny, morphology, and behavior |journal=Frontiers in Ecology and Evolution |date=27 February 2015 |volume=3 |doi=10.3389/fevo.2015.00018 |s2cid=7458974 |doi-access=free }}</ref> Evolutionary differences between species of fish may cause them to either create pits or castles when spawning. The differences were changes in the way that each species fed, their macrohabitats, and the abilities of their sensory systems.<ref name=York-2015/>

====Evolution====
Cichlids are renowned for their recent, rapid evolutionary radiation, both across the entire clade and within different communities across separate habitats.<ref name=Duponchelle-2008/><ref name=Muschik-2012/><ref name=York-2015/><ref name=Zardoya-1996/><ref name=Goodwin-1998>{{cite journal |last1=Goodwin |first1=Nicholas B. |last2=Balshine-Earn |first2=Sigal |last3=Reynolds |first3=John D. |title=Evolutionary transitions in parental care in cichlid fish |journal=Proceedings of the Royal Society of London. Series B: Biological Sciences |date=7 December 1998 |volume=265 |issue=1412 |pages=2265–2272 |doi=10.1098/rspb.1998.0569 |pmc=1689529 }}</ref><ref>{{cite journal |last1=Hulsey |first1=C Darrin |last2=Hollingsworth |first2=Phillip R |last3=Fordyce |first3=James A |title=Temporal diversification of Central American cichlids |journal=BMC Evolutionary Biology |date=December 2010 |volume=10 |issue=1 |pages=279 |doi=10.1186/1471-2148-10-279 |pmid=20840768 |pmc=2944184 |doi-access=free |bibcode=2010BMCEE..10..279H }}</ref> Within their phylogeny, many parallel instances are seen of lineages evolving to the same trait and multiple cases of reversion to an ancestral trait.

The family Cichlidae arose between 80 and 100 million years ago within the order Perciformes (perch-like fishes).<ref name=Zardoya-1996>{{cite journal |last1=Zardoya |first1=Rafael |last2=Vollmer |first2=Dana M. |last3=Craddock |first3=Clark |last4=Streelman |first4=Jeffrey T. |last5=Karl |first5=Steve |last6=Meyer |first6=Axel |date=22 November 1996 |title=Evolutionary conservation of microsatellite flanking regions and their use in resolving the phylogeny of cichlid fishes (Pisces: Perciformes) |journal=Proceedings of the Royal Society of London |series=Series B: Biological Sciences |volume=263 |issue=1376 |pages=1589–1598 |doi=10.1098/rspb.1996.0233 |pmid=8952095 |bibcode=1996RSPSB.263.1589Z |s2cid=45834758 |url=http://nbn-resolving.de/urn:nbn:de:bsz:352-opus-36277 }}</ref> Cichlidae can be split into a few groups based on their geographic location: Madagascar, Indian, African, and Neotropical (or South American). The most famous and diverse group, the African cichlids, can be further split either into Eastern and Western varieties, or into groups depending on which lake the species is from: [[Lake Malawi]], [[Lake Victoria]], or [[Lake Tanganyika]].<ref name=Zardoya-1996/><ref name=Goodwin-1998/> Of these subgroups, the Madagascar and Indian cichlids are the most basal and least diverse.{{citation needed|date=November 2020}}

Of the African cichlids, the West African or Lake Tanganyika cichlids are the most basal.<ref name=Muschik-2012/><ref name=Zardoya-1996/>
Cichlids' common ancestor is believed to have been a spit-spawning species.<ref name=Goodwin-1998/> Both Madagascar and Indian cichlids retain this feature. However, of the African cichlids, all extant substrate brooding species originate solely from Lake Tanganyika.<ref name=Duponchelle-2008/><ref name=Goodwin-1998/> The ancestor of the Lake Malawi and Lake Victoria cichlids were mouthbrooders. Similarly, only around 30% of South American cichlids are thought to retain the ancestral substrate-brooding trait. Mouthbrooding is thought to have evolved individually up to 14 times, and a return to substrate brooding as many as three separate times between both African and Neotropical species.<ref name=Goodwin-1998/>

====Associated behaviors====
Cichlids have a great variety of behaviors associated with substrate brooding, including courtship and parental care alongside the brooding and nest-building behaviors needed for pit spawning. Cichlids' behavior typically revolves around establishing and defending territories when not courting, brooding, or raising young. Encounters between males and males or females and females  are agonistic, while an encounter between a male and female leads to courtship.<ref name=Burchard-1964>{{cite journal |last1=Burchard |first1=John E. |title=Family Structure in the Dwarf Cichlid Apistogramma trifasciatum Eigenmann and Kennedy) |journal=Zeitschrift für Tierpsychologie |date=26 April 2010 |volume=22 |issue=2 |pages=150–162 |doi=10.1111/j.1439-0310.1965.tb01428.x |pmid=5890861 }}</ref> Courtship in male cichlids follows the establishment of some form of territory, sometimes coupled with building a bower to attract mates.<ref name=Nelson-1995/><ref name=York-2015/><ref name=Burchard-1964/> After this, males may attempt to attract female cichlids to their territories by a variety of lekking display strategies or otherwise seek out females of their species.<ref name=Nelson-1995/> However, cichlids, at the time of spawning, undergo a behavioral change such that they become less receptive to outside interactions.<ref name=Burchard-1964/> This is often coupled with some physiological change in appearance.<ref name=Alonso-2011/><ref name=Brown-1978/><ref name=Burchard-1964/>

====Brood care====
Cichlids can have maternal, paternal, or biparental care. Maternal care is most common among mouthbrooders, but cichlids' common ancestor is thought to exhibit paternal-only care.<ref name=Goodwin-1998/> Other individuals outside of the parents may also play a role in raising young; in the biparental daffodil cichlid (''Neolamprologus pulcher''), closely related satellite males, those males that surround other males' territories and attempt to mate with female cichlids in the area, help rear the primary males' offspring and their own.<ref>{{cite journal |last1=Dierkes |first1=P. |title=Reproductive parasitism of broodcare helpers in a cooperatively breeding fish |journal=Behavioral Ecology |date=1 September 1999 |volume=10 |issue=5 |pages=510–515 |doi=10.1093/beheco/10.5.510 |doi-access=free }}</ref>

A common form of brood care involves food provisioning. For example, females of lyretail cichlids (''Neolamprologus modabu'') dig at sandy substrate more to push nutritional detritus and zooplankton into the surrounding water. Adult of '' N. modabu'' perform this strategy to collect food for themselves, but dig more when offspring are present, likely to feed their fry.<ref name=Ota-2014/><ref name=Zworykin-2001>{{cite book |last1=Zworykin |first1=Dmitry |chapter=Parental brood provisioning in cichlid fishes by means of stirring up of the loose material of bottom substrate: a brief review |date=2001 |doi=10.13140/2.1.1937.5048 |title=Cichlid Research: State of the Art |pages=269–286 }}</ref> This substrate-disruption strategy is rather common and can also be seen in convict cichlids (''Cichlasoma nigrofasciatum'').<ref name=Wisenden-1995/><ref name=Zworykin-2001/> Other cichlids have an ectothermal mucus that they grow and feed to their young, while still others chew and distribute caught food to offspring. These strategies, however, are less common in pit-spawning cichlids.<ref name=Zworykin-2001/>

[[File:Managuense with eggs.jpg|thumb|210px|A substrate brooding female managuense cichlid, ''[[Parachromis managuense]]'', guards a clutch of eggs in the aquarium.|alt=]]
Cichlids have highly organized breeding activities.<ref name=Nelson/> All species show some form of parental care for both [[Egg (biology)|egg]]s and [[larva]]e, often nurturing free-swimming young until they are weeks or months old.
Communal parental care, where multiple monogamous pairs care for a mixed school of young have also been observed in multiple cichlid species, including ''[[Amphilophus citrinellus]]'', ''[[Etroplus suratensis]]'', and ''[[Tilapia rendalli]]''.<ref name=McKaye77>{{cite journal |author= McKaye, K.R.|author2 = N.M. McKaye | year = 1977 | title = Communal Care and Kidnapping of Young by Parental Cichlids| journal = Evolution | volume = 31 | pages = 674–681 | doi = 10.2307/2407533 |pmid = 28563477 |issue= 3 |jstor= 2407533}}</ref><ref name="Ward77">{{cite journal | author = Ward, J.A. |author2 = Wyman, R.L. |year = 1977| title= Ethology and ecology of cichlid fishes of the genus ''Etroplus'' in Sri Lanka: Preliminary findings | journal = Environmental Biology of Fishes | volume = 2 | issue = 2 | pages = 137–145 | doi = 10.1007/BF00005369 |bibcode = 1977EnvBF...2..137W |s2cid = 8307811 }}</ref><ref name="Ribbink81">{{cite journal | author = Ribbink, A.J. |author2 = Marsh, A.C. |author3 = Marsh, B.A. | year = 1981 | title = Nest-building and communal care of young by ''Tilapia rendalli dumeril'' (pisces, cichlidae) in Lake Malawi | journal = Environmental Biology of Fishes | volume = 6 | pages = 219–222 | doi = 10.1007/BF00002787 | issue = 2|bibcode = 1981EnvBF...6..219R | s2cid = 9546191 }}</ref> Comparably, the fry of ''[[Neolamprologus brichardi]]'', a species that commonly lives in large groups, are protected not only by the adults, but also by older [[Juvenile (organism)|juveniles]] from previous spawns.<ref>Steeves, Greg. [http://www.africancichlids.net/articles/neolamprologus_brichardi/ Neolamprologus brichardi.] {{Webarchive|url=https://web.archive.org/web/20080505172935/http://www.africancichlids.net/articles/neolamprologus_brichardi/ |date=5 May 2008 }} africancichlids.net. Retrieved 8 April 2008</ref> Several cichlids, including discus (''[[Symphysodon]]'' spp.), some ''[[Amphilophus]]'' species, ''[[Etroplus]]'', and ''[[Uaru]]'' species, feed their young with a skin [[secretion]] from mucous glands.<ref name=Loiselle/><ref name=baensch/>

The species ''[[Neolamprologus pulcher]]'' uses a cooperative breeding system, in which one breeding pair has many helpers that are subordinate to the dominant breeders.

Parental care falls into one of four categories:<ref name=baensch/> substrate or open brooders, secretive cave brooders (also known as guarding speleophils<ref name=Balon>{{cite journal |author=Balon, E.K. |year=1975 |title=Reproductive guilds of fishes: A proposal and definition |journal=Journal of the Fisheries Research Board of Canada |volume=32 |issue=6 |pages=821–864 |doi=10.1139/f75-110 }}</ref>), and at least two types of [[mouthbrooder]]s, ovophile mouthbrooders and larvophile mouthbrooders.<ref name="Keenleyside">{{cite book |last=Keenleyside |first=M.H.A. |year=1991 |chapter=Parental care |title=Cichlid Fishes: Behaviour, ecology and evolution |publisher=Chapman and Hall |place=London, UK |pages=191–208 |isbn=978-0-412-32200-6}}</ref>

====Open brooding====
Open- or substrate-brooding cichlids lay their eggs in the open, on rocks, leaves, or logs. Examples of open-brooding cichlids include ''[[Pterophyllum]]'' and ''Symphysodon'' species and ''[[Anomalochromis thomasi]]''. Male and female parents usually engage in differing brooding roles. Most commonly, the male patrols the pair's territory and repels intruders, while the female fans water over the eggs, removing the infertile ones, and leading the fry while foraging. Both sexes are able to perform the full range of parenting behaviours.<ref name="Keenleyside"/>

====Cave brooding====
[[File:Cyphotilapia frontosa mouthbrooding.jpg|thumb|right|180px|A female ''[[Cyphotilapia frontosa]]'' mouthbrooding fry, which can be seen looking out of her mouth]]
Secretive cave-spawning cichlids lay their eggs in caves, crevices, holes, or [[Shell dweller|discarded mollusc shells]], frequently attaching the eggs to the roof of the chamber. Examples include ''[[Pelvicachromis pulcher|Pelvicachromis]]'' spp., ''[[Convict cichlid|Archocentrus]]'' spp., and ''[[Apistogramma]]'' spp.<ref name=baensch/> Free-swimming fry and parents communicate in captivity and in the wild. Frequently, this communication is based on body movements, such as shaking and [[pelvic fin]] flicking. In addition, open- and cave-brooding parents assist in finding food resources for their fry. Multiple neotropical cichlid species perform leaf-turning and fin-digging behaviors.<ref name="Keenleyside"/>

====Ovophile mouthbrooding====
Ovophile mouthbrooders incubate their eggs in their mouths as soon as they are laid, and frequently mouthbrood free-swimming fry for several weeks. Examples include many [[East African Rift]] lakes ([[Lake Malawi National Park|Lake Malawi]], [[Lake Tanganyika]], and [[Lake Victoria]]) endemics, e.g.: ''[[Maylandia]]'', ''[[Pseudotropheus]]'', ''[[Tropheus]]'', and ''[[Astatotilapia burtoni]]'', along with some [[South America]]n cichlids such as ''[[Geophagus steindachneri]]''.

====Larvophile mouthbrooding====
Larvophile mouthbrooders lay eggs in the open or in a cave and take the hatched larvae into the mouth. Examples include some variants of ''[[Geophagus altifrons]]'', and some ''[[Aequidens]]'', ''[[Gymnogeophagus]]'', and ''[[Satanoperca]]'', as well as ''[[Oreochromis mossambicus]]'' and ''[[Oreochromis niloticus]]''.<ref name=Loiselle/><ref name=baensch/> Mouthbrooders, whether of eggs or larvae, are predominantly females. Exceptions that also involve the males include eretmodine cichlids (genera ''[[Spathodus]]'', ''[[Eretmodus]]'', and ''[[Tanganicodus]]''), some ''[[Sarotherodon]]'' species (such as ''[[Sarotherodon melanotheron]]''<ref name=Kishida00>{{cite journal |last1=Kishida |first1=Mitsuyo |last2=Specker |first2=Jennifer L. |date=February 2000 |title=Paternal mouthbrooding in the black-chinned tilapia, ''Sarotherodon melanotheron'' (Pisces: Cichlidae): Changes in gonadal steroids and potential for vitellogenin transfer to larvae |journal=Hormones and Behavior |volume=37 |issue=1 |pages=40–48 |doi=10.1006/hbeh.1999.1556 |pmid=10712857 |s2cid=8128737 }}</ref>), ''[[Chromidotilapia guentheri]]'', and some ''[[Aequidens]]'' species.<ref name="Loiselle"/><ref name=Keenleyside/><ref name=Coleman99>{{cite journal | author = Coleman, R. |date=January 1999 | title = Mysterious mouthbrooders | journal = Cichlid News  | pages = 32–33}}</ref> This method appears to have evolved independently in several groups of African cichlids.<ref name="Nelson"/>