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Egg Size Determines Offspring Size in Neotropical Cichlid Fishes (Teleostei: Cichlidae) Author(S): Ronald M

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Egg Size Determines Offspring Size in Neotropical Cichlid Fishes (Teleostei: Cichlidae) Author(S): Ronald M Egg Size Determines Offspring Size in Neotropical Cichlid Fishes (Teleostei: Cichlidae) Author(s): Ronald M. Coleman and Alison P. Galvani Source: Copeia, Vol. 1998, No. 1 (Feb. 3, 1998), pp. 209-213 Published by: American Society of Ichthyologists and Herpetologists Stable URL: http://www.jstor.org/stable/1447721 Accessed: 01/05/2009 00:45 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=asih. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit organization founded in 1995 to build trusted digital archives for scholarship. We work with the scholarly community to preserve their work and the materials they rely upon, and to build a common research platform that promotes the discovery and use of these resources. For more information about JSTOR, please contact [email protected]. American Society of Ichthyologists and Herpetologists is collaborating with JSTOR to digitize, preserve and extend access to Copeia. http://www.jstor.org Copeia,1998(1), pp. 209-213 Egg Size Determines Offspring Size in Neotropical Cichlid Fishes (Teleostei: Cichlidae) RONALD M. COLEMAN AND ALISON P. GALVANI Within the family Cichlidae, the size of eggs We investigated the relationship between egg varies tremendously across species (Breder and size and hatchling size in 26 species of Neotrop- Rosen, 1966; Fryer and Iles, 1972; Coleman, ical cichlids. We also examined eggs and hatch- 1991). Sizes range from the small eggs (0.9 mm lings produced by 17 females within one species in diameter) of rams (Microgeophagusramirezi) to [convict cichlids, "Cichlasoma(Archocentrus)" ni- the massive eggs (4.5 mm in diameter) of some grofasciatum], and eggs and hatchlings from of the African mouthbrooding cichlids such as within single clutches of two species (convict Cyphotilapiafrontosa (unpubl. data). The hatch- cichlids and rainbow cichlids, Herotilapia multis- lings of the large-egged mouthbrooders are pinosa) to look at the effects of intraspecific and clearly much larger than the hatchlings of the intraclutch variation in egg size. rams and other small-egged species (pers. obs.). the between size and However, relationship egg MATERIALSAND METHODS hatchling size for smaller differences in egg size has not been For within the explored. example, Cichlid eggs are not spherical but generally the of sizes is Neotropical cichlids, range egg are better described as prolate spheroids (elon- much smaller: such as Cich- small-egged species gated spheres). To compare nonspherical eggs, lasoma dimerushave an diameter of average egg Coleman (1991) proposed using the "effective 1.3 whereas the mm, largest-egged species, diameter," which is the diameter the egg would "Cichlasoma tuba has an (Theraps)" average egg have if its contents were reshaped into a perfect diameter of 2.4 mm Does (RMC, unpubl. data). sphere. In the case of a prolate spheroid, the this small difference in size actu- relatively egg effective diameter (de), as used throughout this translate into a consistent difference in the ally paper, is the cube root of the major axis multi- size of the does the hatchlings? Furthermore, plied by the square of the minor axis (i.e., cube variation in size within a ac- egg single laying root of length X width X width). count for differences in size? Under- hatchling The systematics of Neotropical cichlids are in the of differential allo- standing consequences a state of flux, creating confusion in the generic cation to size between in a egg eggs single status of many of the species used in this study and will clutch and within among species help (Kullander, 1983; Stiassny, 1991). We follow the evolution us understand the factors shaping Stiassny (1991) in referring to certain of these of egg size. fishes by the genus " Cichlasoma,"including the not translate into A larger egg may necessarily section names provided by Regan (1906, 1906- a larger hatchling for several reasons. For ex- 1908) as though they designate subgenera; how- ample, a given measure of egg size may not be ever, the exact phylogenetic meaning of these a true representation of the energetic and ma- sections is uncertain. The 26 species examined terial resources of an egg. This could be the were Aequidens pulcher, Cichlasoma bimaculatum, case if larger eggs are simply filled with more C. dimerus, C. festae, C. pusillum, "Cichlasoma water than smaller eggs, and size differences (Amphilophus)" alfari, "C. (Am.)" citrinellum,"C. may solely be accounted for by water content. (Am.)" longimanus, "C. (Am.)" rostratum, "C. In such a case, the difference in egg size may (Nandopsis)" managuense, "C. (Archocentrus)" be insignificant to the resulting hatchling. This centrarchus,"C. (Ar)" nigrofasciatum,"C. (Ar.)" is particularly plausible because fish eggs are hy- septemfasciatum,"C. (Ar)" spilurum, "C. (Ther- drated and expand to some extent when they aps)" bifasciatum, "C. (Th.)" intermedium, "C. leave the female (Kamler, 1992). An ideal mea- (Th.)" nicaraguense, "C. (Th.)" sieboldii, "C. sure of egg size may involve a complete bio- (Th.)" tuba, Cleithracaramaronii, Crenicichlalepi- chemical analysis of the contents of an egg, re- dota, Herotilapiamultispinosa, Microgeophagus altis- porting protein, lipid, and carbohydrate con- pinosa, Nannacara anomala, Neetroplusnematopus, tent (e.g., Kamler, 1992); however, this is not and Pterophyllumscalare. practical for many studies. Indeed, for situations The fishes were maintained in the laboratory when the eggs are particularly small and yet using a typical freshwater aquarium setup. Each must be kept alive, even weighing individual setup consisted of a glass aquarium (38-280 li- eggs or measuring their volume can be difficult. ters), gravel, sponge filter, heater, and either ? 1998 by the American Society of Ichthyologists and Herpetologists 210 COPEIA, 1998, NO. 1 pieces of slate, flowerpots, or broken flowerpots. pect significant positive relationships in either All species examined were biparental substra- analysis. tum spawners (Loiselle, 1994), though differing To examine intraspecific variation, we sam- in the precise choice of spawning site (e.g., flow- pled eggs and hatchlings from single spawnings erpot, piece of slate, exposed glass tank bot- of 17 convict cichlid females. To assess the effect tom). Water temperatures were maintained be- of variation within a clutch in size of egg on size tween 24 and 28 C, and the light cycle was 12D/ of offspring, we obtained individual hatchlings 12L. Fishes were fed twice daily on Purina trout from eggs of known size using single clutches chow and Tetramin cichlid flakes with weekly of convict cichlids and rainbow cichlids. In both supplements of live brine shrimp. cases, eggs were scraped off the laying site as Aquaria were observed daily for the presence described above and their size measured in a of eggs or hatchlings. When a clutch of eggs was petri dish. For the within-clutch analyses, we se- sizes as as of more in- found, a sample of 30-50 eggs was scraped from lected extreme well eggs of the substratum with a blunt metallic spatula. termediate size to maximize the range egg sizes used. Each was measured under the The rest of the eggs were left to hatch. Eggs egg and then in a numbered were then either measured immediately or microscope placed in the As each stored in 70% isopropyl alcohol. Storage in iso- hatching cup hatching aquarium. hatched several the was propyl alcohol does not significantly alter egg egg days later, hatchling removed and in a numbered vial of 4% diameter (unpubl. data). placed To calculate effective diameter, we measured formalin. Eggs that did not hatch were dropped from The were the length and width of 20 eggs from each sam- subsequent analysis. hatchlings measured as above and similar using a Wild dissecting scope fitted with an regressions per- ple formed. ocular micrometer, calibrated to 0.01 mm. The (2.0 cm diameter, 1.5 cm 20 values were averaged to produce a mean egg Hatching cups size for the On the of high) were made from the bottom portion of a species. day hatching, vial secured with silicon to a hatchlings were collected with a turkey baster. plastic specimen small square of glass to make the cup sink. The Because live hatchlings are difficult to measure cups were placed in a half-filled 38-L aquarium, accurately, they were immediately preserved in with a heater and airstone. This hatching-cup 4% formalin and measured several days later. design proved successful in other experiments Three parameters were measured with each because it allows water to circulate over the eggs hatchling lying on its side: the total length of but keeps the eggs from blowing out of the the hatchling (TL), and the length and width cups. of the yolk sac. Typically, 20 hatchlings were measured to obtain a mean hatchling value for RESULTS the species, though in a few cases, particularly when were rare, fewer than 20 were hatchlings Mean hatchling length was positively corre- used [C. 13; "C. (Am.)" alfari 13; "C. festae lated with mean egg diameter in the 26 species (Am.)" rostratum 18; "C. (Ar.)" septemfasciatum examined (Fig. 1; r = 0.83; df = 24; P < 0.01). 16; "C. 19; "C. inter- (Th.)" bifasciatum (Th.)" Diameters of most of the eggs were 1.3-1.9 mm.
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