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Hypotheses of phylogenetic relationship among skuas. Heavy black branches are those on which a Catharactalike plumage morphology can be inferred. Open branches are those on which a Stercorariuslike plumage morphology can be inferred. Branch lengths are not proportional to evolutionary change. (a) Major features of the topology proposed by Cohen et al. (1997) based on mtDNA sequences. Either the Stercorarius or the Catharacta morphology must have evolved twice on this tree. (b) Alternative topology proposed herein. Each morphology need only evolve once on this tree.
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Last year, Cohen et al. presented molecular data suggesting the surprising result that both currently recognized genera of skuas, Stercorarius and Catharacta (Aves: Stercorariidae), are not monophyletic. However, the most enigmatic conclusion from their analysis, that S. pomarinus is sister to C. skua, rests solely on mtDNA sequence data. When the...
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... Birds used overwintering habitats September-May (Table 2) Our results of disparate migratory routes of sympatrically breeding congeners also raise additional questions about the evolutionary origins (Braun & Brumfield, 1998;Chu et al., 2009;Cohen et al., 1997) and biogeographic spread of North American jaegers. For We grouped the movements off of Japan as a staging period because they proceeded a 5-month migratory loop over Micronesia and the bird also stopped in the Oyashio Current in the spring before returning to the Arctic. ...
Polar systems of avian migration remain unpredictable. For seabirds nesting in the Nearctic, it is often difficult to predict which of the world's oceans birds will migrate to after breeding. Here, we report on three related seabird species that migrated across four oceans following sympatric breeding at a central Canadian high Arctic nesting location. Using telemetry, we tracked pomarine jaeger (Stercorarius pomarinus, n = 1) across the Arctic Ocean to the western Pacific Ocean; parasitic jaeger (S. parasiticus, n = 4) to the western Atlantic Ocean, and long-tailed jaeger (S. longicaudus, n = 2) to the eastern Atlantic Ocean and western Indian Ocean. We also report on extensive nomadic movements over ocean during the postbreeding period (19,002 km) and over land and ocean during the prebreeding period (5578 km) by pomarine jaeger, an irruptive species whose full migrations and nomadic behavior have been a mystery. While the small sample sizes in our study limit the ability to make generalizable inferences, our results provide a key input to the knowledge of jaeger migrations. Understanding the routes and migratory divides of birds nesting in the Arctic region has implications for understanding both the glacial refugia of the past and the Anthropocene-driven changes in the future.
... It has previously been shown that in 2 polymorphic species, the arctic (S. parasiticus) and pomarine skua (S. pomarinus), color variation has evolved independently by mutations in the melanocortin-1 receptor (MC1R), and some of the mutations implicated in color variation have occurred independently in the 2 lineages (Mundy et al. 2004;Mundy 2013, 2017). The third species is the south polar skua (S. maccormicki), one of the great skua species (previously comprising the genus Catharacta), which is more closely related to pomarine than arctic skuas (Braun and Brumfield 1998). Three color morphs have been defined in the south polar skua, with ventral plumage coloration varying from pale buff in the pale morph to dark brown in the dark morph ( Figure 1; Ainley et al. 1985;Olsen and Larsson 1997). ...
... An obvious next step to identify genetic variation contributing to the plumage polymorphism in south polar skuas would be to conduct a Genome-wide association study (GWAS), an approach that has succeeded in uncovering candidate loci affecting melanin coloration in other birds (crows, Poelstra et al. 2014;wood warblers, Toews et al. 2016; Swainson's thrushes, Delmore et al. 2016;and ruff, Küpper et al. 2016). The low level of genetic variation in south polar skuas, which is presumably related to a low effective population size resulting from a founder event (Ritz et al. 2008), would increase the power of this approach (Braun and Brumfield 1998;Janssen and Mundy 2017). ...
Coloration is evolutionarily labile and so provides an excellent trait for examining the repeatability of evolution. Here, we investigate the repeatability of the evolution of polymorphic variation in ventral plumage coloration in skuas (Stercorarius: Stercorariidae). In 2 species, arctic (S. parasiticus) and pomarine skuas (S. pomarinus), plumage polymorphism was previously shown to be associated with coding changes at the melanocortin-1 receptor (MC1R) locus. Here, we show that polymorphism in a third species, the south polar skua (S. maccormicki), is not associated with coding variation at MC1R or with variation at a Z-linked second candidate locus, tyrosinase-related protein 1 (TYRP1). Hence, convergent evolution of plumage polymorphisms in skuas is only partly repeatable at the level of the genetic locus involved. Interestingly, the pattern of repeatability in skuas is aligned not with phylogeny but with the nature of the phenotypic variation. In particular, south polar skuas show a strong sex bias to coloration that is absent in the other species, and it may be that this has a unique genetic architecture.
... In contrast, while some species of great skuas do show plumage colour variation [7,8], this variation is more continuous and a discrete polymorphism as found in Arctic and pomarine skuas is absent. The phylogeny of skuas has previously been a source of contention, but there is considerable evidence that pomarine skuas are the sister group to the great skuas, with the other smaller species-Arctic skuas and long-tailed skuas (S. longicaudus)-forming a separate clade [9][10][11]. ...
A key outstanding issue in adaptive evolution is the relationship between the genetics of intraspecific polymorphism and interspecific evolution. Here, we show that the pale/dark ventral plumage polymorphism that occurs in both the pomarine skua (Stercorarius pomarinus) and Arctic skua (S. parasiticus) is the result of convergent evolution at the same locus (MC1R), involving some of the same amino acid sites. The dark melanic MC1R allele in the pomarine skua is strongly divergent from the pale MC1R alleles. Whereas the dark allele is closely related to MC1R alleles in three species of great skua (S. skua, S. maccormicki, S. lonnbergi), the pale pomarine skua MC1R alleles present a star-like pattern in an intermediate position on the haplotype network, closer to alleles of the long-tailed skua (S. longicaudus). Variation at other nuclear loci confirms a close relationship between the pomarine skua and the great skuas. The plumage polymorphism in pomarine skuas might have arisen in the common ancestor of pomarine and great skuas, only being retained in pomarine skuas. Alternatively, the pale and melanic MC1R alleles may have evolved independently in different lineages and been brought together in pomarine skuas by hybridization. In this case, introgression of a pale MC1R allele into the pomarine skua from another skua lineage is most likely. Our current data do not permit us to distinguish between these hypotheses, and assaying genome-wide variation holds much promise in this regard. Nevertheless, we have uncovered an intriguing example of a functionally important allele within one species that is shared across species.
... A year later, Braun & Brumfield (1998) reanalysed the Cohen et al.'s (1997) data using a maximumlikelihood approach and cladogram searches constrained for the monophyly of Catharacta. They recovered a cladogram similar to that of Cohen et al. (1997), the difference being the position of S. pomarinus as sistertaxon to a clade comprising the Catharacta species ( Figure 1B). ...
... Figures 1A and 1C). Braun & Brumfield (1998) defended their findings by arguing that they were more consistent with pre-cladistic, plumage-, body mass-, and behaviour-based hypotheses of a "natural" Catharacta. They furthermore suggested that, if their cladogram ( Figure 1B) proves "correct", it would be reasonable to place S. pomarinus in its own genus, for which Coprotheres would be available. ...
... Nevertheless, his main recommendation was to merge all species into a single genus, the older Stercorarius. Andersson (1999b: 212) further commented that "if Catharacta is retained ... the Braun & Brumfield (1998;both constrained (B) and unconstrained (C) for the monophyly of Catharacta). Numbers refer to percentage bootstrap (100 replicates) and Goodman-Bremer support (Grant & Kluge 2008b). ...
In this contribution I comment on the generic taxonomy of skuas, Stercorariidae, based on the currently available hypotheses of phylogenetic relationships for the group – i.e., the cladograms. Specifically, the different cladograms were examined following Hennig's principle of reciprocal illumination, in which a given hypothesis is evaluated by the extent to which it agrees with competing hypotheses. Currently, all species are often assigned to genus Stercorarius. However, chewing lice- (Insecta, Phthiraptera), behaviour- (territorial display and calls) and molecular-based (mitochondrial DNA) hypotheses of phylogenetic relationships all indicate that Pomarine Skua Stercorarius pomarinus and the species formerly placed in genus Catharacta are monophyletic and sister to a clade comprising Long-tailed S. longicaudus and Parasitic S. parasiticus Skuas. Therefore, contrary to the prevailing view that all species within the family should be placed in a single genus, I argue herein that in a cladistic-based classification by sequencing, both S. parasiticus and S. longicaudus should retain their generic name, whereas S. pomarinus should be transferred to Catharacta, as C. pomarina. © 2016, Sociedade Brasileira de Ornitologia. All rights reserved.
... They became efficient hunters, sometimes taking even relatively large vertebrate prey (Andersson, 1999) in spite of the constraints imposed by their evolution from gull-like ancestors (Mayr, 2011; Weir and Mursleen, 2012). There is a consensus that the ancestral members of Stercorariidae resembled jaegers rather than skuas (Cohen et al., 1997; Braun and Brumfield, 1998; Andersson, 1999; Ritz et al., 2008; Chu et al., 2009), i.e. they were relatively small and agile and had clearly different adult and juvenile plumages. De Korte and Wattel (1988) regard it as plausible that these basal skuas/jaegers were kleptoparasites and resembled modern parasitic jaegers (S. parasiticus) in morphology and lifestyle. ...
... Despite the current consensus that Pomarine jaegers are more closely related to skuas than to other jaegers, there are two contrasting interpretations of this evolutionary process; both views are supported by different kinds of empirical evidence. On the basis of nuclear DNA and skeletal morphology, Braun and Brumfield (1998) and Chu et al. (2009) proposed that skuas evolved in the northern hemisphere and that Pomarine jaegers diverged from that lineage at an early stage, retaining the ancestral jaeger traits (i.e. a clear difference between a juvenile gull plumage and a black-and-white adult plumage; elongated tail feathers; and small body size). According to the other interpretation, advocated by Cohen et al. (1997) on the basis of mitochondrial DNA and ectoparasites (see also Ritz et al., 2008), and proposed earlier by Andersson (1973) on the basis of behavioural traits, Pomarine jaegers are closely related to great skuas (Catharacta skua) and evolved relatively recently. ...
Background: The Fennoscandian tundra differs from other arctic regions hosting brown lemmings (Lemmus spp.) in that it supports exceptionally low numbers of avian predators. Jaegers (Stercorarius spp.) play a central role in the guild of predators exploiting lemming outbreaks, as they are long-lived and have a vast supply of marine survival resources. Collared lemmings (Dicrostonyx spp.) evolved much earlier than brown lemmings (Lemmus spp.).
Hypothesis: Lemming–vegetation dynamics and summer predation are causally interconnected. Avian predators that are adapted to exploit lemming outbreaks evolved in tundra areas where lemming–vegetation interactions generate density oscillations with a mean period short enough to provide dependable breeding resources for predators.
Predictions: Long-tailed jaegers (Stercorarius longicaudus), primarily exploiting collared lemmings, diverged from parasitic jaegers (S. parasiticus) more than 2 million years ago. Pomarine jaegers (S. pomarinus) evolved more recently in response to the emergence of regular oscillations of brown lemming populations. The current breeding distribution of jaegers should reflect the distribution of the two lemming genera, except for areas where the oscillations generated by lemming–plant interactions are too erratic to provide dependable breeding resources.
Empirical evidence: Our hypothesis accounts for the current distributions, behaviours, and ecologies of the three jaeger species. In addition, long-tailed jaegers did diverge early from other Stercorariidae. But we still do not know whether the timing of the divergence matches the evolution of collared lemmings. Evidence suggests that either: (1) Pomarine jaegers diverged from the skua (Catharacta spp.) lineage early, before skuas had developed their distinctive morphological traits; or (2) Pomarine jaegers diverged recently from great skuas (Catharacta skua) and then reverted to the ancestral jaeger morphology. Only the latter interpretation is consistent with our hypothesis, so we need to have more reliable timing of the evolution of
long-tailed and Pomarine jaegers.
... The Stercorariidae constitutes a small, distinctive family of seven kleptoparasitic seabird species all now regarded as within the genus Stercorarius (Remsen et al. 2013), although much debate exists about their systematics; sometimes, more species and a second genus are recognized within the family (Cohen et al. 1997;Braun and Brufield 1998;Ritz et al. 2008;Chu et al. 2009). The family comprises the larger and mostly Brown skuas and the smaller, more agile jaegers. ...
Background: Skuas and jaegers (Charadriiformes: Stercorariidae) are seabirds breeding at moderate to high latitudes and some perform extensive post-breeding transequatorial migrations. Most species overwinter and perform significant portions of their migratory flyways along the Pacific coast of South America, but scant information is available on their at-sea ecology in this waters. Our aims in this study were to determine: 1) the timing of occurrence and fluctuations in abundance of skua and jaeger species, 2) their spatial distribution within the coastal zone and 3) at-sea behavior of birds, including flock size and interactions with other seabird species.
Results: Between July 2006 and October 2013, we conducted at-sea bird counts at Valparaiso Bay (33°S) in central Chile and confirmed the occurrence of Chilean skuas (Stercorarius chilensis), Brown skuas (S. antarcticus), and Parasitic jaegers (S. parasiticus). Parasitic jaegers are regular austral summer visitors (November to March), whereas Brown skuas occur in the area only in winter (July to October). Chilean skuas were regularly recorded year-round in the area with higher abundances between late winter and early spring (August to October). Brown and Chilean skuas where observed comparatively offshore, whereas Jaegers presented a more coastal distribution, probably associated to host presence. Chilean skuas kleptoparasitized similar-sized (shearwaters and fulmars) and larger seabird species (boobies), whereas jaegers chased only smaller coastal birds (gulls and terns). Brown skuas engaged in no kleptoparasitic behaviors. All three species were observed mostly as solitary individuals.
Conclusions: Skuas and jaegers showed in general a marked seasonality in their occurrence and abundance (only Chilean skua occurs year-round) and use this area as a commuting and stopover zone within their extensive migratory flyway along the southeastern Pacific.
Keywords:
... Relationships of Stercorarius skua are uncertain. Mitochondrial DNA analyses have suggested that S. skua is more closely related to Stercorarius pomarinus than to taxa breeding in the southern hemisphere (Blechschmidt et al. 1993, Cohen et al. 1997, Ritz et al. 2008, but see Braun & Brumfield 1998), whereas analyses of morphology and ectoparasites support monophyly of the 'great skuas' (Andersson 1999a, 1999b, Chu et al. 2009). As well as genetic differences from southern hemisphere skuas, Great Skua differs from all other skuas in plumage at most ages (Furness 1987, Malling-Olsen & Larsson 1997). ...
This paper is the seventh report of the TaxonomicSub-Committee (TSC) of the BOU RecordsCommittee (BOURC) relating to the British List.Species-level decisions are based on criteria out-lined by Helbig et al. (2002). The sixth report ofthe Sub-Committee was published by Sangsteret al. (2010a).The Sub-Committee has been working with theAssociation of European Rarities Committees’Taxonomic Advisory Group, which has recentlypublished recommendations online (Crochet et al.2010). The reports of the BOURC and its TSC,which are published in Ibis, remain the officialpublications of the BOU.Capped Petrel [Black-capped Petrel]Pterodroma hasitataCapped Petrel and Bermuda Petrel differ diagnos-ably in pattern and coloration of crown, hindneckand rump, and uppertail-coverts (Brooke 2004,Howell & Patteson 2008). Molecular phylogeneticanalysis further suggests that these are not closelyrelated taxa (Jesus et al. 2009). Capped Petrel andBermuda Petrel are therefore best treated as sepa-rate species:• Capped Petrel Pterodroma hasitata (polytypic)• Bermuda Petrel Pterodroma cahow (monotypic)Capped Petrel is in Category A of the BritishList.Water Rail Rallus aquaticusRecent vocal and molecular studies indicate thatWater Rail consists of two distinct lineages (deKroon et al. 2008, Tavares et al. 2010). Two maincall types have been recorded for Rallus aquaticusaquaticus: a long and complex call (the ‘pigsqueal’), and a short and simple call, both repeatedin series, whereas for Rallus aquaticus indicus, onlya complex call type has been recorded, which isdistinct from that of R. a. aquaticus (de Kroonet al. 2008). Preliminary playback experimentsshowed a lack of response of R. a. aquaticus to thecomplex calls of R. a. indicus, suggesting a biologi-cal significance for these differences (de Kroon et al.2008). Phylogeographical analysis of mitochondrialand nuclear DNA sequences places R. a. indicusin a separate clade from the clade containingR. a. aquaticus and Rallus aquaticus korejewi, andfurther showed that these clades are strongly differ-entiated (Tavares et al. 2010). The two groups alsodiffer in plumage (Cramp & Simmons 1980, deKroon 1991, Taylor & van Perlo 1998). Water Railis therefore best treated as two species:• Water Rail Rallus aquaticus (polytypic, withsubspecies aquaticus and korejewi)
... At lower taxonomic levels, problems remain unresolved for diverse reasons, e.g. complex patterns of molt for the Laridae (Chu, 1994Chu, , 1998 Pons, Hassanin & Crochet, 2005), and geographically variable polymorphism of definitive plumages for the Stercorariidae (Braun & Brumfield, 1998; Andersson, 1999; Ritz et al., 2008). The remaining 'wading' families, typified by terrestrial foraging for sessile, fossorial invertebrates or less commonly mobile (terrestrial, aerial, or natatorial ) insects, approach synonymy for the Charadriiformes for many (e.g. ...
Modern shorebirds (Neornithes: Charadriiformes) are among the most diverse, phylogenetically challenging, and evolutionarily critical of avian orders. Despite several morphological analyses and diverse molecular studies of the order, a consensus regarding relationships within the order, as well as clarity of membership of Turnix, has heretofore remained elusive. This paper describes a cladistic analysis of 1024 phenotypic characters (427 multistate, 209 of which were ordered), which represents an outgrowth of a prior higher-order study. The analysis was performed at species level for all families, exclusive of the highly derived, monophyletic, and intensively studied Alcidae and Lari (the analysis included three out-groups and an in-group, comprising 242 taxa). Characters analysed (excluding 83 poorly known, primarily myological characters) comprised 446 of the skeleton, 558 of the definitive integument, and 20 of natal patterns. Eighty parsimony-uniformative autapomorphies – neutral with respect to summary statistics – were included for descriptive and diagnostic purposes. The analysis found a large set of shortest trees: the majority-rule consensus of these was fully resolved, and most included nodes were unanimously conserved in the solution set. Support was significant for a majority of nodes, both by bootstrap percentages and (for higher-order nodes) support (decay) indices. All families were inferred to be monophyletic in this analysis, although the stability of relationships below intergeneric groups was generally found to be low. Some findings challenge previous morphological results, as well as many groupings variably inferred from molecular data. Close relationships were confirmed between: (1) the Jacanidae and Rostratulidae; (2) the Dromadidae and Haematopodidae; (3) the Ibidorhynchidae and Recurvirostridae; (4) among the Chionididae, Alcidae, Stercorariidae, Rynchopidae, and Laridae; and (5) between the Charadriidae and a clade comprising the Thinocoridae, Phalaropodidae, and Scolopacidae. Pedionomus was inferred to be the sister group of other Charadriiformes, with the Turnicidae arguably considered the sister group of the Charadriiformes or its basalmost member. The Jacanidae and Rostratulidae were strongly supported as sister groups, in agreement with most modern studies. Two points of controversy inferred herein were: Pluvianellus as the sister group of the clade comprising the Alcidae and Lari, and a closer relationship between Thinocoridae and Scolopacidae than between the former and Pedionomus. The topological neighborhood of weakest support involved the position of the Lari-Alcidae (especially relative to the Burhinidae) and the inclusion of Pluvianellus. A lesser point of contention was the rejection of the Glareolinae as closely related to the Laridae and allies, with the Laridae marginally favoured in some molecular studies. The Charadriidae were inferred to comprise four primary subgroups: lapwings (Hoploxypterus and Vanellus), dotterels (nine restricted genera), greater plovers (Pluvialis), and lesser plovers (Anarhynchus and Charadrius). The Scolopacidae comprised four subfamilies, the most speciose of which were the Calidridinae (sandpipers and stints), Tringinae (shanks, curlews, and godwits), and Scolopacinae (snipe and woodcocks). Lesser scolopacid clades included the ‘surfpipers’ (Arenaria and Aphriza) and the ‘meadowpipers’ (Philomachus and Tryngites). Biogeographic and evolutionary implications of the phylogeny are discussed, including the potential of fossils for stratocalibrations, and a revised classification of the order is proposed.© 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 567–618.
... This latter, more-specific hypothesis is not supported by our osteological data. Indeed, as Braun & Brumfield (1998) describe, a (S. pomarinus, C. skua) clade is not supported by the nuclear data of Cohen et al. (1997) either, and although Cohen et al.'s mitochondrial data do support it, the strength of that support can be debated. ...
... pomarinus, C. skua) clade is not supported by the nuclear data of Cohen et al. (1997) either, and although Cohen et al.'s mitochondrial data do support it, the strength of that support can be debated. Braun & Brumfield (1998) did not comment on Cohen et al.'s (1997) ectoparasite evidence, and that evidence does provide additional support for a (S. pomarinus, C. skua) clade; however, the support is inconsequential when viewed in a more global context: adding the ectoparasite characters of Andersson (1999a) to our skeletal data had no affect on the topology of the shortest tree. ...
... The simplest explanation for our skeletal data is not that S. pomarinus is the sister taxon to C. skua, with the clade thus formed being the sister to the remaining forms of Catharacta, but that all of the members of Catharacta form a monophyletic group, the sister to which is S. pomarinus. This is the hypothesis proposed by Braun & Brumfield (1998) and Andersson (1999b). ...
The skuas (Aves: Charadriiformes, Stercorariidae) consist of two assemblages. On the basis of size, plumage, and distributional similarities, each of the two assemblages has long been considered monophyletic, and this traditional hypothesis has commonly been manifested in the recognition of two genera, Stercorarius and Catharacta; conversely, more recently collected molecular and ectoparasite evidence yields an alternative hypothesis, in which one member of Stercorarius, Stercorarius pomarinus, is more closely related to the forms in Catharacta than to the other Stercorarius sp. In this study we used skeletal morphology to test the competing hypotheses of skua phylogeny. Cladistic analysis of 141 osteological characters provided strong support for the molecular/ectoparasite hypothesis. However, those skeletal data did not support a sister-taxon relationship between S. pomarinus and Catharacta skua, as inferred from mitochondrial DNA sequence data; instead, they resolved pomarinus as the sister of a monophyletic Catharacta. Additionally, our skeletal evidence did not support a sister-group relationship between skuas and auks, as constraining skua/auk monophyly increased the tree length by nearly 5%.
© 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 157, 612–621.
... Even with new species tree phylogenetic methods, disentangling the effects of hybridization and lineage sorting on a phylogeny of recently diverged species is and will continue to be difficult (Braun and Brumfield, 1998;Holder et al., 2001;Buckley et al., 2006). In this regard it is noteworthy that M. candei and M. vitellinus, despite hybridizing extensively where their distributions meet in western Panama, were not resolved as sister taxa in our phylogenetic analyses; in fact the trees that clustered them together had some of the highest deep coalescence scores (Table 2). ...
Although the power of multi-locus data in estimating species trees is apparent, it is also clear that the analytical methodologies for doing so are still maturing. For example, of the methods currently available for estimating species trees from multilocus data, the Bayesian method introduced by Liu and Pearl (2007; BEST) is the only one that provides nodal support values. Using gene sequences from five nuclear loci, we explored two analytical methods (deep coalescence and BEST) to reconstruct the species tree of the five primary Manacus OTUs: M. aurantiacus, M. candei, M. vitellinus, populations of M. manacus from west of the Andes (M. manacus (w)), and populations of M. manacus from east of the Andes (M. manacus (e)). Both BEST and deep coalescence supported a sister relationship between M. vitellinus and M. manacus (w). A lower probability tree from the BEST analysis and one of the most parsimonious deep coalescence trees also supported a sister relationship between M. candei and M. aurantiacus. Because hybrid zones connect the distributions of most Manacus species, we examined the potential influence of post-divergence gene flow on the sister relationship of parapatrically distributed M. vitellinus and M. manacus (w). An isolation-with-migration (IM) analysis found relatively high levels of gene flow between M. vitellinus and M. manacus (w). Whether the gene flow is obscuring a true sister relationship between M. manacus (w) and M. manacus (e) remained unclear, pointing to the need for more detailed models accommodating multispecies, multilocus DNA sequence data.
