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The Adaptive Significance of Local Variations in the Bill and Jaw Anatomy of North European Red-Necked Grebes Podiceps Grisegena

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The Adaptive Significance of Local Variations in the Bill and Jaw Anatomy of North European Red-Necked Grebes Podiceps Grisegena The adaptive significance of local variations in Gr the bill and jaw anatomy of North European Red-necked Grebes Podiceps grisegena be A to Tb JON FJELDSÅ Re ad FJELDSÅ, J. 1982 : The adaptive significance of local variations in the bill and jaw anatomy of North European Red-necked Grebes Podiceps grise- gr~ gena. - Ornis Fennica 59:84-98. gel Red-necked Grebes from the northernmost breeding populations in Fin- tio land and Russia develop a long and slender bill. This trend, and also SM the large size of East Siberian and Nearctic Red-necked Grebes, might thz be associated with a change in the diet from invertebrates to fish in Fu areas where the piscivorous Great Crested Grebe is missing. A sample of 87 Red-necked Grebes from a Danish wintering area comprised both inl South Scandinavian and slender-billed northern types. These were studi- tu( ed in order to discover to what extent the morphological and anatomical dif variation affects the diet. tio The slender-billed Red-necked Grebes converge with Great Crested Grebes in the ability to open the bill rapidly and keep a strong grip on Fo the prey without sliding of the mandible . Both adaptations may be valu- ha able for catching fish. Study of the stomach contents showed that the ho, Red-necked Grebes of South Scandinavian type fed mainly on annelid, Sul gleaned, from plants and picked up from the bottom, while individuals of the northern type specialized on squids and, fish. th( Although this does not prove anything, it provides support for the Ca assumption that the morphological variation was caused by selection due Tl to the presence or absence of Great Crested Grebes . wl Jon Fjeldsd, Zoological Museum, University of Copenhagen, Universi- no tetsparken 15, DK-2100 Copenhagen 0, Denmark . zol The Red-necked Grebe Podiceps gri- Finnish birds, was later synonymized. th( segena has two distinct subspecies However, although there is consider- Ar (Storer 1979), the West Palaearctic able overlap with .the other European Cr grisegena and the Nearctic and East birds, my data (Fjeldså 1982) con- cri Siberian holboellii, the latter being firm that there is a clinical change in 19 rather pale and similar in size to the this part of the range . As we move et Great Crested Grebe Podiceps crista- from southwestern Scandinavia to Sio tus. Birds from Balchash ("balcha- near the northern distribution limit of N( schensis") are described as rather the species (inland areas from Kuopio th( long-winged. All the specimens I have to Finnish Lapland, Kola, Archan- ta( seen from this area are, however, gelsk, Dvina) the mean wing length mi within the grisegena range, and hardly increases by 1 .5 0/o and the tarsus by va qualify for taxonomic recognition. 2 .5 0/o, while the bill increases in Gr Similarly, the name schioleri, proposed length by as much as 11 0/o (Fig. 1) ve by Hortling (1929) for the rather big and becomes more slender, its shape Gr J. Fjeldsd : Functional anatomy of North European Red-necked Grebes 85 approaching that of the Great Crested Grebe. The interpretation of this trend will be the objective of this study. A possible interpretation and how to test it The morphological change in northern Red-necked Grebes may reflect adaptations to climate or to habitat gradients. However, according to general ecogeographical rule, adapta- tions to a colder climate should give smaller relative change in bill length than in wing length as we move north. Further, the summer temperatures of inland areas change little with lati- tude, and the wintering areas of the different Red-necked Grebe popula- tions (Danish waters and according to Folkestad 1978 northwest Norway) FiG. 1. Bill lengths of Red-necked (dots) have similar temperatures . Finally, and Great Crested Grebes (open circles) from different populations, measured on specimens holboellii shows no variation from the in breeding plumage in the Zoological Mu- subarctic zone, with its bleak lakes, to seums of the Universities of Copenhagen, Oslo the warm climate and rich lakes of the and Helsinki ; the Swedish Museum of Natural Canadian forest/prairie ecotones. History, Stockholm ; the American Museum of Natural History, New York ; the Australian Thus, considering the species as a Museum, Sidney, and the museum of CSIRO's whole, the geographical variation does Wildlife Research, Canberra. Bottom : bill not follow the climate or lake type lengths of the grebes collected in Danish zones. winter quarters and used in this study. It might be that the large size of the Red-necked Grebes in North America is due to the lack of Great ern coastal zone. Population fluctua- Crested Grebes (compare the diet of tions (Thomasson 1953), gene flow cristatus, e.g. in Cramp & Simmons contact through the European popula- 1977, and holboellii, e.g. in Palmer tions and some interspecific contact in et al. 1962). The smaller bill dimen- all European winter quarters might sions in the East Siberian than in the explain the moderate amplitude of the Nearctic holboellii (Fig. 1) agrees with geographical variation of the Europe- the fact that the former has some con- an Red-necked Grebes. Altogether, the tact with Great Crested Grebes on variation in the species coincides well migration and in winter. The size with what we could expect, if the variation of Finnish Red-necked variation was due to interspecific Grebes could also be explained by the competition. More distribution and very sparse breeding of Great Crested biometric data from North Russia Grebes outside the southern and west- would be desirable to reinforce the 86 ORNIs FENNICA Vol. 59, 1982 documentation. Nevertheless, it ap- study of the birds from these important pears to be a reasonable working winter quarters for Red-necked Grebes hypothesis that Red-necked Grebes particularly interesting is the fact that evolve morphological adaptations they have very variable bills (Fig. 1), which permit them to live like Great which suggests that birds of both Crested Grebes in areas where this North Finnish/North Russian and species is missing. Alternatively, we South Scandinavian provenience are may suppose that the two were origin- represented. Altogether 87 Red-neck- ally allopatric ecological counterparts, ed and six Great Crested Grebes were and that the Red-necked Grebe collected after the oil-spill by the managed to invade the range of the Danish Game Research Station, Kale, Great Crested Grebe by means of and sent to the Zoological Museum, "character displacement", evolving a University of Copenhagen. smaller size and smaller bill suited for taking invertebrate prey (for diet in the breeding areas, see Madsen Anatomical studies 1957, Onno 1958, Markuze 1965) . Methods. In methods and terminology, the It is pertinent to test whether the anatomical study follows that of Fjeldså (1981) on Peruvian Grebes . morphological changes affect the birds' The jaws and jaw muscles of one Great ecology. This could be established by Crested and six Red-necked Grebes (including studying whether Red'-necked Grebes two of the most slender-billed) from Hatter change their ecology as we pass north Rev were dissected under a 10 .5-67.5 X zoom Kyowa dissecting microscope . In addi- through Finland. Unfortunately, tion an examination was made on the sculls though, a documented ecotope change of 26 Great Crested and, 19 Red-necked would be insufficient as evidence, as Grebes (including holboellii) . Distances from it could have three or more causes : articulations to insertion points of muscles were measured with caliper rulers, and the It could be due to the change in "power arms" calculated by the appropriate morphology, but it could as well re- trigonometric functions. The potential muscle flect changes in the habitat and food force is determined- from the number of fibres resources, or changes in resource ac- rather than the muscle mass (Bock 1974) . To obtain an index of fibre number, I calculated cessibility due to the lack, in the north, the physiological cross section of the muscles of interference competition from Great as insertion area times the sine of the insertion Crested Grebes (cf. Berndt & Drenck- angle. Since various parts of the muscle may hahn 1974). The data might con- have different insertion angles, the total cross section was calculated, section by section. This sequently be hard to interpret. was done at either end of each muscle, and It may be safer to make the study the mean of the two vallues was usedl. For in a common wintering area, where pennate fibre arrangements, the cross section the interference competition is slight, value. The potential speed, of contraction pinnation angle to give a functionally relevant and where all the birds use the same value. The potential speed of contraction food supply. is judged from the fibre lengths. Some card,- The opportunity to make such a board models, in 4 X scale, were used to determine the effects of coupling ligaments study arose in January 1979, when and changes in action angles of muscles as the 400-500 Red-necked Grebes were bones change position. Lever calculation was killed by an oil-spill on Hatter Rev used to determine muscle torques, while the (55°55'N, 08°50'E) in Danish coastal interaction of muscles was studied by con- structing freebody diagrams (Bock 1974), waters (Larsen 1979) . These birds can elaborated for non-gravity and, static situations, all be assumed to have fed in the and ~disregardling the load against which the same macrohabitat . What makes the muscles act. J. Fjeldsd : Functional anatomy of North European Red-necked Grebes 87 Due to the complexity of the superficial TABLE 1 . Some important distances (mm) and deep muscles and space considerations, and physiological cross section values (0, few aspects can be properly depicted .
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