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Population Differentiation and Rapid Evolution of Egg Color in Accordance with Solar Radiation

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Population Differentiation and Rapid Evolution of Egg Color in Accordance with Solar Radiation The Auk 125�4):796–802, 2008 Th American Ornithologists’ Union, �����2008. Printed in USA. POPULATION DIFFERENTIATION AND RAPID EVOLUTION OF EGG COLOR IN AccORDANCE WITH SOLAR RADIATION DAVID C. LAHTI1 Department of Biology, Morrill Science Center, University of Massachusetts, Amherst, Massachusetts 01003, USA Abstract.—A�����frican populations ��������� ���� ��the ������Village �Weaver���� �Ploceus cucullatus) are subject to brood parasitism by the egg-m�m��king Diederik Cuckoo �Chrysococcyx caprius), which has resulted in high variation in egg color between individuals. Considering the nesting environment of the Village Weaver and the putative pre-parasitism ancestral egg color, I �ypothesiz that blue-green pigment m�y protect Village Weaver mb�yos from solar radiation. O this basis, more intense �more chromatic) blue-green egg colors m�y have evolved by natural selection in Village Weavers in the �bsence �� D�derik Cuckoos. Moreover, even in African populations that are parasitizd, Village Weaver eggs m�y have diverged in accordance with nesting light environments. I accordance with the proposed �ypothesis, over the past �w� centuries �w� introduced populations have evolved more intense blue-green egg colors compared with their source populations. Tw� African source populations also d�ff� from each other in the predicted direction, the population with the sunnier nesting climate having more intensely blue-green eggs. These results suggest that upon relaxation of brood parasitism by an egg m�m��, Village Weaver eggs m�y �dapt to solar radiation. Received 13 February 2007, accepted 23 February 2008. Ky word: bir��d��� eggs������, egg�� color�����, intro����duce������d species������, Ploceus cucullatus, rapid evolution, relaxd selection, solar radiation, Village Weaver. Diferenciación entre Poblaciones y Evolución Rápida del Color de los Huevos de Acuerdo a la Radiación Solar Resumen.—��Las ��pob�����laciones ������africanas� d Ploceus cucullatus está sujetas � parasitism� d crí� por parte d la especie q� �mita huevos Chrysococcyx caprius, lo que ha conducid� � una alta variació entre individuos en el color d los huevos. Considerand� el �mbiente d anidació d P. cucullatus y el color ancestral putativo previo al origen del parasitism�, planteé la hipótesis d que la pigmentació �zul-verd pod�í� proteger � los mbriones d P. cucullatus d la radiació solar. S�bre esta base, los huevos d color �zul-verd má intenso �má cromático) pod�ían haber evolucionad� por selecció natural en ausencia d C. caprius. Admá, incluso en poblaciones africanas q� son parasitadas, los huevos d P. cucullatus pod�ían haberse diferenciad� d acuerd� � la luminosid�d d los �mbientes d anidació. D acuerd� � �� esperad� segú la hipótesis propuesta, � �� largo d los últimos dos siglos, los huevos d dos poblaciones introducidas han evolucionad� hacia colores �zul-verd má intensos en comparació con sus poblaciones originales. Dos poblaciones fuente �bicadas en África tamb�é d�fieren entre í en la direcció predicha: la població con el clim� d anidació má solead� presenta huevos con color �zul-verd má intenso. Estos resultados sugieren q� ante una disminució d la presió d parasitism� por parte d una especie �mitadora d huevos, los huevos d P. cucullatus pod�ían �daptarse � la radiació solar. Bird egg color is � promising trait for evolutionary analysis b- within � clutch and distinctive within the population. These traits cause it appears to b subject to � m�dest array �� selection pres- permit � female to recogniz her �w eggs and thus avoid brood sures �Uderwood and Sealy 2002, Kilner 2006) and because parasitism by the egg-m�m��king D�derik Cuckoo �Chrysococcyx changes in egg color tend to b evolutionary �Punnett 1933, Gosler caprius, hereafter “cuckoo”; Victoria 1972, Lahti and Lahti 2002). et al. 2000). “Ground color” is the relatively uniform background Distinctiveness and consistency of egg colors have no know color of an egg, as opposed to spots or other markings that can also function besides egg recognition, and these traits decayd in 75– b present; in many cases, ground color and markings are subject to 150 generations following �w� independent introductions to is- distinct sources of selection �Gosler et al. 2005, Westmoreland and land without egg-m�m��king brood parasites (Lahti 2005). These Kiltie 2007). I the African Village Weaver �Ploceus cucullatus), changes, in turn, have compromised egg recognition in the intro- both ground color and spotting �� � female’ eggs are consistent duced populations �Cruz and Wiley 1989, Lahti 2006). 1E-mail: lahti@bio.�mass.d� The Auk, Vol. 125, N�mb� 4, pages 796–802. ISSN 0004-8038, electronic ���������������ISSN����������� 1938-4254. 2008 by Th American Ornithologists’ Union. All rights reserved. Please direct all requests for permission to photocopy or reproduce article content through the University of California Press’ Rights and P�missions wbsite, http://www.ucpressjournals. com/reprintInfo.as�.�������������� DOI: 10.1525/��au������������k.2008.07033 — 796 — 04_Lahti_07-033.indd 796 10/21/08 2:38:02 PM OCTOBER 2008 — EVolUTION OF BLUE-GREEN EGG ColoR — 797 A unanswered question is what sources �� selection remain much solar radiation for the resulting heat to b dissipated, heat to act on ground color �� Village Weaver eggs when brood parasit- will build up in the shell and b conducted into the egg, ulti- ism by cuckoos is removed as � factor. Th ground color �� Village mately killing the mb�y�. Th only direct dmonstrations of Weaver eggs in Africa varies from white to mdium greenish-blue this possibility are with artificial pigments �Montevecchi 1976, �Collias 1984); in the introduced populations on the island of His- Bertram and Burger 1981). Short of this xtrem point, �bsor- paniola and Mauritius, the egg colors are � subset �� the respec- bance of light by eggshell pigment would b beneficial, insofar as tive ancestral ranges in western Africa and South Africa (Lahti it forestalls �bsorbance by the mb�y�. 2005). Egg recognition m�y still function, to som xtent, in the Transmittance in the ultraviolet �U�) wavelength range can introduced populations in the contex� of conspecifi� parasitism b � danger to vertebrate mb�yos irrespective �� heating, causing �Freeman 1988), but there is no reason to believe that the xtent developmental �bnormalities even at common temperate levels �� of such parasitism would d�ffer between source and introduced xposure �Blaustein and Belden 2003, Perotti and Diegeuz 2006). populations. Blue-green egg color does not appear to function in A poultry study �Shafey et al. 2002) showd that eggshells are par- crypsis �Gö�mark 1992, Westmoreland and Kiltie 2007); regard- ticularly fficient at blocking U� light by �bsorbing it and that the less, crypsis would not aid Village Weaver eggs, because nests are brow eggs of at least one breed are more ffective than white eggs enclosed and the eggs are not visib� through the weave �Moreau in this regard. Biliverdin, the d�minant pigment in blue-green 1960). A recently proposed signaling �ypothesis �Moreno and eggs, has � high �bsorbance peak in the U� �Kennedy and Vevers Osorno 2003) relies on the inspection �� eggs by males and sub- 1973, Inomata et al. 2005). Consequently, biliverdin, including sequent m�dulation �� paternal feeding behavior. This �ypothesis that occurring in eggs, has been �ypothesizd to function in pho- is not relevant to the Village Weaver. Males d� not generally have toprotection in invertebrates �M�Donagh 2001), fish �Y�maguchi opportunities to �bserve egg colors, because males are xcludd et al. 1976), and lizard �Austin and Jessing 1994). by females from the enclosed nests throughout the egg stage and I general, in m�derate light environments where sunlight have not been �bserved entering them; moreover, in none �� the is filtered through vegetation, mb�yonic health and survival study populations d� male Village Weavers regularly provision m�y b aidd by blue-green eggshell pigmentation that can �b- young �Collias and Collias 1971; D. C. Lahti pers. �b). O possi- sorb and reflect incident solar radiation rather than allowing its bility is that egg ground color serves no function in the introduced transmission into the egg interior. I particular, because Village populations. I� so, ground color under relaxd selection might b Weavers tend to nest in isolated trees and denud the vegetation xpected to becom more variab� or—especially if there are costs around their nests (prob�b�y for predator-avoidance and mate- to pigment production or deposition—to decrease in intensity. attraction functions; Collias and Collias 1959, 1969, 1970), their A alternative �ypothesis is that solar radiation has an in- globular nests are commonly xposed to direct light through- fluence on egg color that is detectab� in the �bsence �� functions out the d�y. Th females generally lay eggs when the nests are for crypsis or egg recognition in this species. I particular, bird still green �Jacob et al. 1978); given the translucence of the nest- eggs m�y b subject to mb�yonic d�mage from solar radiation, ing materials, the light environment of the nest’ interior is blue- and pigment m�y b photoprotective �M�A�d�wie 1886, R�manoff green, similar to � forest shad as describd by Edler �1993). and R�manoff 1949). Theoretically, solar d�mage to bird mb�yos Blue-green eggs in such an environment would reflect light can tak �w� form: conducted heat from light �bsorbd by the across the visible spectrum in proportion to the incident light in- shell surface, and light transmitted through the eggshell. Pigment tensity and would block U� transmittance as well. This �ypoth- increases �bsorbance and so is unlikely to �meliorate conductance esis is consistent with the �bservation that the three ploceine from the shell surface. Pigment decreases transmittance, however, weavers in Africa that build nests similar to that of the Village and in so doing would decrease any �dverse ffects �� light passing Weaver and appear not to have varying egg colors in response through the shell.
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