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Using Egg Density and Egg Mass Techniques for Incubation Stage Assessment to Predict Hatch Dates of Greater Flamingo Phoenicopterus Ruber Roseus Eggs

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Using Egg Density and Egg Mass Techniques for Incubation Stage Assessment to Predict Hatch Dates of Greater Flamingo Phoenicopterus Ruber Roseus Eggs 131 Using egg density and egg mass techniques for incubation stage assessment to predict hatch dates of Greater Flamingo Phoenicopterus ruber roseus eggs N. Jarrett1, V. Mason1, L. Wright2 & V. Levassor1 'The Wildfowl and Wetlands Trust, Slimbridge, Gloucestershire GL2 7BT, UK. Email: nigel. jarrett0w w t. org. uk Centre for Ecology, Evolution and Conservation, School of Environmental Sciences, University of East Anglia, Norwich NR4 7TJ, UK. Egg density and the egg mass techniques for incubation stage assess­ ment were developed to predict the hatch dates of Greater Flamingo Phoenicopterus ruber roseus eggs laid in captivity at WWT, Slimbridge, UK. The accuracy of each technique was tested on 20 parentally incubat­ ed eggs by comparing actual hatch date with predicted hatch date. For the egg mass technique a strong positive correlation existed between actual and predicted fresh mass, suggesting that model accuracy was high. Both techniques predicted hatch dates within two days 80% of the time. These techniques were found to be useful for accurate incubation stage assessment of Greater Flamingo eggs and the authors encourage aviculturalists managing captive colonies to use them. Key Words: egg mass, egg density, incubation stage assessment, Greater Flamingo, Phoenicopterus ruber roseus Each year at The Wildfowl and shallowly flooded islands. Single eggs Wetlands Trust (WWT), Slimbridge, UK, are laid on these nest mounds, and up to 60 pairs of colonially breeding cap­ nest-defence duties are shared by both tive Greater Flamingos Phoenicopterus sexes until a chick is hatched after 26- ruber roseus compete for nest sites. 32 days' incubation. Fighting between Breeding pairs usually construct nest nest mound occupants and their neigh­ mounds, often up to 0. 3m high and just bours or pairs that intrude into the 0. 2m apart, on and from the substrate of colony often results in eggs being dis- © Wildfowl & Wetlands Trust Wildfowl (2003) 54: 131-142 132 Predicting hatch dates of Greater Flamingo eggs placed from nests. As flamingos do not estimate incubation stage of Waterbird attempt to retrieve displaced eggs, up eggs of unknown lay-date, but neither to 25% of annual egg production is lost are particularly suitable for flamingos. in this way (Pickering 1992). The candling technique involves Displaced eggs are usually assessing egg air space development replaced, sometimes within minutes, by shining a light source into the egg so by eggs laid by intruding pairs. In 2000 that the size of the air space can be there were five cases of eggs being laid visualised and compared to known pat­ on nest mounds by intruding females terns of enlargement. However, the within two hours of the loss of the pre­ candling technique is difficult to per­ vious nest occupants' egg. In all cases, form on flamingo eggs, which are nor­ the 'new' eggs were adopted by the mally thick-shelled and, sometimes, original nest occupants (N. Jarrett, marked on the surface with scratches pers. obs.). In order to free nest sites or nest debris. Moreover, candling is a for other pairs to use, aviculturalists technique best practised in a low light sometimes discard these 'new' eggs environment, often not attainable in a after 32 days' incubation, believing field situation. them to be beyond their hatch-by date, The egg flotation technique involves when, in fact, they are viable, although immersing the egg in a container of at an earlier stage of development (N. water so that the angle and height of Jarrett, pers. obs. ). Until now there has the egg in the water column can be not been a reliable or safe field tech­ measured against patterns that are nique for aviculturalists to use when calibrated for different incubation attempting to estimate the incubation stages. Although Alberico (1995) has stage of flamingo eggs. demonstrated that this technique does Avian eggs lose approximately 16% not affect egg hatchability, immersion of their initial mass at an almost linear in water is known to carry a risk of rate during incubation, almost entirely damage to the developing embryo due to diffusive water loss through the (Gabel & Mahan 1996). This risk factor, eggshell pores (Drent 1970; Rahn & Ar coupled with the impracticality of car­ 1974). This has enabled field workers to rying an immersion vessel and uncont­ develop several techniques, based on aminated water for flotation into the egg mass change over the incubation field, leads the authors to believe that period, to determine the incubation the flotation technique is inappropriate stage of eggs of species with known for flamingo eggs. incubation periods. Egg candling (eg The egg density (eg Furness & Hanson 1954; Weller 1956) and egg Furness 1981) and egg mass (eg flotation (eg Carroll 1988; Walter & Westerkov 1950; Lind 1991) techniques Rusch 1997; Brua & Machin 2000) are for incubation stage assessment have the most commonly used techniques to been adopted by relatively few workers Predicting hatch dates of Greater Flamingo eggs 133 but represent reliable and safe alterna­ balance, and size was recorded to the tives to egg candling and egg flotation nearest 0. 1 mm using vernier callipers. (Jarrett & Warren 1996). Both tech­ niques require egg size (maximum The egg density technique length and breadth measurements) Hoyt (1979) described egg volume and egg mass data to be collected so (V) (cm3) as: that fresh egg mass and egg density can be estimated. Furthermore, both V=K\/. LB: ' equation 1 techniques assume that all eggs lose the same proportion of their fresh where L is egg length (cm), B is egg mass, at a constant rate, over a defined breadth (cm) and Kv is a shape coeffi­ incubation period with the implication cient. Hoyt (1979) observed that the that egg density and egg mass can be value of Kv was 0. 509±0.008 (S. D. ) for predicted for each day of incubation. most species of birds except those lay­ For the egg density technique, incuba­ ing very asymmetrical eggs. Equation 1 tion stage is indicated when the was used to calculate the egg volume observed egg density is entered into a of 217 Greater Flamingo eggs. Egg species-specific density loss equation. density (D) (g/cm3) was then calculated For the egg mass technique, incubation for each egg using the equation: stage is indicated when observed egg mass is referenced to a model of mass D=M/V equation 2 loss for an egg of that particular egg's estimated fresh mass. where M is egg mass (g). To calculate Here the usefulness of the egg den­ pre-hatch egg density, data were col­ sity and the egg mass techniques for lected from parentally incubated eggs incubation stage assessment to predict to estimate the proportion of initial egg the hatching dates of eggs laid by cap­ mass lost over the course of the incu­ tive Greater Flamingo at WWT bation period. Incubation period was Slimbridge are described and dis­ defined as the number of days between cussed. the onset of incubation and the begin­ ning of hatch (when the chick breaks into the air space and begins to vocalise Methods and tap with its beak on the inner eggshell). Thus the mass of 18 eggs Fresh egg mass and size measure­ was recorded within two hours of ovipo­ ments (maximum length and breadth) sition and daily after day 24 until the were recorded for 217 Greater hatching chick was audible within the Flamingo eggs laid in 2001, 2002 and air space of the egg. The average pro­ 2003. Mass was recorded within four portion of fresh mass lost was hours of oviposition to the nearest 0. 1 g U . 5%±0.56 (S. E. ) (range 10. 4-19. 5%) using a portable, levelled electronic over an average incubation period of 134 Predicting hatch dates of Greater Flamingo eggs 28.0 days (range 26-30 days). Thus pre-hatch egg density (Dp) Kw=Mf/LB2 equation 5 (g/cm:! ) for the sample of eggs was cal­ General Linear Models were used to culated using the equation: investigate how Kw differed between years and first, second, third and fourth Dp=Df.0. 855 equation 3 clutches (Norusis 2000). In the model where Df is fresh egg density (g/cm3) chosen, Kw was considered as the and 0. 855 is the average proportion of dependent variable with year and initial egg mass retained at the time the clutch number incorporated as main chick enters the air space. So that the effects. Interactions between these hatch date of eggs of unknown incuba­ independent variables were also inves­ tion stage could be predicted, a tigated. species-specific equation describing The accuracy of the sample mean egg density over the incubation period Kw was assessed by using equation A to was developed using regression analy­ estimate fresh mass of each egg and sis. then using a correlation to determine General Linear Models were used to the similarity between observed and investigate how fresh egg density dif­ predicted fresh egg masses. On the fered between years and first, second, assumption that all Greater Flamingo third and fourth clutches (Norusis eggs lost 14. 5% of their initial mass at 2000). In the model chosen, fresh egg a linear rate over the 28-day incubation density was considered as the depend­ period, tables were compiled to show ent variable with year and clutch num­ the predicted daily mass of eggs with ber incorporated as main effects. fresh masses ranging those of the 217 Interactions between these independ­ eggs sampled. (Appendix 1 provides an ent variables were also investigated. example of a table for eggs of fresh mass ranging from 150 to 165g). The egg mass technique Evaluating the techniques Hoyt (1979) described the relation­ ship between the fresh mass (Mf) (g) of The egg density and egg mass tech­ a bird's egg and its linear dimensions, niques for incubation stage assess­ as: ment were used to predict the hatching date of 20 Greater Flamingo eggs of Mf=Kw/.
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