Female ornamentation and courtship behavior in Cryptoheros: Implications for classification?

No group of Central American cichlids have received as much attention the past 15 years that the species of the genus Archocentrus (sensu Kullander, 2003). Allgayer (2001) suggested an alternative classification by describing the genus Cryptoheros, but it has been questioned repeatedly, if this classification really reflects the phylogenetic relationship (see Archcentrus vs. Cryptoheros).

Accoding to Allgayer (2001), Cryptoheros differs from Archocentrus in the habit of breeding in crevices rather than on open surfaces. A closer look at the reproductive behavior of the Cryptoheros species (sensu Allgayer), however, reveals differences within the genus: A further evidence that this group is not monophyletic?

Female ornamentation and courtship behavior

The Cryptoheros species can be devided in two groups depending on the presence of presence of female ornamentation and courtship behavior (I have no observations on A. altoflavus, but it probably belongs to the latter group):

Ornamentation and courtship absent	Ornamentation and courtship present
A. spilurus*	A. myrnae
A. sajica	A. nanoluteus
	A. nigrofasciatus
	A. septemfasciatus
*observations made on animals coming from Lago Yojoa

Female ornamentationa and courtship behavior is briefly documented in the following using A. myrnae as an example. The presence of ornamentation (changed and intensified color especially in the region of the abdomen) and courtship displays (female presents its belly bending itsself and struting the fins) could be observed in all species listed above. Intensity and details of courtship behavior may vary across species.

	A. myrnae in normal coloration: As described by Loiselle (1997), this species is very colorful. All observations were made on fish collected in the Rio Uatsi, Costa Rica (2002) and their descendants. A male is shown in the upper picture, a female in the lower one.
	When a female gets ready to spawn, the behavior and coloration changes. Females occupy crevices and defend them aggressively. In the picture, the courtship coloration of A. myrnae is shown. Note the dark area ranging from the beginning of the ventrals to the pectorals to the black dots in the center of the body to the beginning of the anal fin. The blue coloration is intensified along the abdomen of the fish. This color pattern is only present during courtship, i.e. usually from the time the females occupy a territory until they spawn.
	While aggressively chasing other females and small males, females ready to spawn actively court dominant males. They swim back and forth from the crevice to the male, bend themself while they cant over right in front of the male (picture) and try to attract the male into the crevice. In A. myrnae it could also be observed that females quiver during courtship (similar to the quivering of many Central American cichlids during brood care).
	After spawing, females show a broodcare color pattern, which is very similar in all Cryptoheros (also the non-courters): fading of colors and an intensifying of melanin based color patters. The males show a similar coloration, however, in most cases the color switches with the emergence of the fry.

These observations have their limitations (only two pairs from aquarium strains observed in A. nanoluteus and A. septemfasciatus; small founder populations in A. myrnae and A. nigrofaciatus). Nonetheless, an important questions must be raised: Is the presence of female ornamentation and courtship behavior an autapomorphy characterizing a monophyletic group? If so, what does this mean for the classification of the Archocentrus species?