Molecular Evidence for Extrapair Paternity and Female-Female Pairs in Antarctic Petrels

Considerable interspecific variation exists in the frequency of extrapair fertilizations (EPFs) in birds. In general, EPFs are more common and occur at higher frequencies in passerines than in nonpasserines (Westneat and Sherman 1997). Lower rates of EPFs are typical for territorial nonpasserines as well as those that breed colonially (Westneat and Sherman 1997). This seems to contradict Birkhead and Møller's (Birkhead and Møller 1992, Møller and Birkhead 1993) hypothesis of intense sperm competition in colonial birds. Their arguments were based on the assumption that the need for nest defense in dense aggregations restricts the ability of males to guard their mates, and that the high number of potential extrapair mates available in colonies selects for a high rate of extrapair copulations (EPCs). In contrast, Westneat and Sherman (1997) found no correlation across species between the frequency of EPFs and nesting dispersion, local breeding density, or breeding synchrony, although EPFs were related to nesting density within species. This suggests that EPC rates are not informative regarding EPF rates in colonial birds (Westneat and Sherman 1997), or that the pattern reported by Møller and Birkhead (1993) does not hold true when more species are included. The conflicting evidence regarding the relationship between extrapair activities and breeding density calls for more empirical studies, especially among colonial nonpasserines. Social monogamy is the predominant mating system in the Procellariiformes (Warham 1990). Several aspects of their breeding biology may, however, provide favorable opportunities for extrapair sexual activity. First, colonial breeding provides ample opportunities for EPCs because many potential partners are available at close range (Birkhead and Møller 1992, Møller and Birkhead 1993). Second, when the sexes are spatially and/or temporally separated, as may be the case in procellariiforms where adults seek food far from the colony, males have few cues to assess whether their mates have been unfaithful. Hence, few reasons exist to expect a facultative decrease in male parental investment if cuckolded, in contrast to the case for many territorial species where a male may have more reliable cues to his mate's unfaithfulness (e.g. female disappearance, high intrusion rate, etc.). Accordingly, colonial breeding may facilitate EPCs for both sexes, and colonial species may be expected to display high rates of EPC. Paternity studies require error-free sex determination of adults. This is straightforward for clearly dimorphic species, or if sex determination can be done from genitalia during the fertile period. But if fieldwork can be performed only during the nestling period, sex determination may be problematic for largely monomorphic species, and indirect methods must be used. In the Antarctic Petrel (Thalassoica antarctica), the two sexes differ slightly in mean body size. Lorentsen and Røv (1994) used this difference to determine the sex of Antarctic Petrels by discriminant function analysis (DFA). The procedure correctly determined the sex of 92% of the birds in a sample from the same year. This does not necessarily imply a similar resolution if the discriminant function is adopted for samples from other years, or if data are collected by other observers. Moreover, although useful for many purposes, 92% resolution in sex determination is insufficient for paternity studies. Therefore, we performed molecular sexing of all breeding adults according to the PCR-based method of Griffiths et al. (1998). The main aim of our study was to analyze whether extrapair paternity occurs in a colonial procellariiform, the Antarctic Petrel. We also tested the robustness of morphological sex determination (from DFA) across seasons relative to that obtained from molecular techniques.