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Phalacrocorax [Stictocarbo] Kenyoni, Which Was De- Scribed by Siegel-Causey (1991) from the Aleutian Islands Using Midden Remains and Exist- Ing Skeletal Specimens

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Phalacrocorax [Stictocarbo] Kenyoni, Which Was De- Scribed by Siegel-Causey (1991) from the Aleutian Islands Using Midden Remains and Exist- Ing Skeletal Specimens The Auk 117(2):308-320, 2000 A CRITICALEVALUATION OF KENYON'SSHAG (PHALACROCORAX[STICTOCARBO] KENYONI) SIEVERT ROHWER,1'3 CHRISTOPHER E. FILARDI,1 KIMBERLY S. BOSTWICK,2 AND A. TOWNSENDPETERSON2 'Burke Museum and Department of Zoology, Box 353010, University of Washington, Seattle, Washington 98195, USA; and 2Natural History Museum and Biodiversity Research Center, University of Kansas, Lawrence,Kansas 66045, USA ABSTRACT.-We examine the validity of Phalacrocorax [Stictocarbo] kenyoni, which was de- scribed by Siegel-Causey (1991) from the Aleutian Islands using midden remains and exist- ing skeletal specimens. We emphasize a morphometric evaluation of the taxon using 224 skel- etal specimens of North Pacific cormorants, but we also evaluate the qualitative characters originally used to characterize R kenyoni. Principal components and discriminant function analyses of 14 skeletal characters failed to support the validity of the species. Similarly, all seven of the character states that Seigel-Causey described as unique to R kenyoni also were found in R pelagicus and P urile. Thus, the three type specimens of R kenyoni appear to be R pelagicus. Although we could not confirm the validity of P kenyoni, our morphometric anal- yses revealed that R pelagicus individuals from the central Aleutians are smaller than those from surrounding populations. Received 11 December 1998, accepted 20 July 1999. SIEGEL-CAUSEY (1991) described a new spe- kenyoni, based on detailed examinations of a cies of cormorantfrom the Aleutian Islands;he more limited reference series of specimens. named it Stictocarbo (=Phalacrocorax) kenyoni, Rohwer and Filardi were responsible for the reflecting his prior studies of major clades in morphometric analyses, and Bostwick and Pe- the Phalacrocoracidae (Siegel-Causey 1988). terson performed the qualitative analyses. Siegel-Causey first discovered this bird in mid- den remains from Amchitka Island in the far METHODS western Aleutian Islands but later found three recent skeletal specimens that he ascribed to Hereafter,we refer to the three North Pacific cor- kenyoni and designated as types (Siegel-Causey morants as kenyoni, pelagicus, and urile for Kenyon's 1991, Siegel-Causey et al. 1991). No other spec- Shag (P kenyoni), Pelagic Cormorant (P pelagicus), imens of this species exist. The external ap- and Red-faced Cormorant(P urile),respectively. Ta- ble 1 provides general localities and dates of collec- pearance of R kenyoni remains undescribed, tion for the specimens used in our morphometric and the species was not accepted as valid by analyses. The three kenyonispecimens were from AOU (1998). AmchitkaIsland, Alaska;the 25 urilespecimens were To facilitate an evaluation of the validity of P from Prince William Sound, or from other Alaskan kenyoni, the University of Washington Burke localities to the west and north; localities for the 196 Museum (UWBM) salvaged every intact cor- pelagicusspecimens are plotted in Figure 1. About morant recovered from the beaches of Prince one-third of the 196 pelagicusspecimens were col- William Sound, Alaska, following the Exxon lected from May to August and thus are likely to be Valdezoil spill of 24 March 1989. All were pre- from breeding areas. Although many of the speci- served as skeletal specimens, and when car- mens were not yet of breeding age, young pelagicus casses were not too rotten, associated extended are known to summer near the colonies where they hatched (Palmer1962). wings and flat skins also were preserved. Our Qualitative characters.-Because Seigel-Causey's di- analyses focused primarily on a morphometric agnosis of kenyoniwas based on seven autapomorph- evaluation of P kenyoni using 224 skeletal spec- ic skeletal characters(Siegel-Causey 1991:appendix imens of cormorantsfrom the North Pacific.We 2) and its small size, we paid special attention to the also evaluated the qualitative characters that qualitative characters of skeletal morphology de- Siegel-Causey (1991) used to characterize R scribed in his Appendix 1. Our interpretationsof Sie- gel-Causey's characterswere based on standard avi- an anatomicalreferences (Howard 1929, George and 3E-mail: [email protected] Berger 1966, Baumel et al. 1979, Baumel and Witmer 308 This content downloaded from 129.237.46.100 on Thu, 18 Sep 2014 14:09:14 PM All use subject to JSTOR Terms and Conditions April 2000] Evaluation of Kenyon's Shag 309 TABLE 1. Specimens examined by species, locality, and season (breeding = 1 May to 31 August). Map sym- bols for pelagicus refer to Figure 1. Breeding Nonbreeding Locality Map symbol season season R kenyoni Aleutian Islands 0 3 R urile Alaska 15 10 R pelagicus Valdez and Southeast Alaska V 29 124 Aleutian Islands A 17 1 Pribilofs and northern Alaska p 0 3 British Columbia and Washington W 3 3 California C 2 6 Russia R 7 0 Japan J 1 0 Totals 74 150 1993) and on careful inspection of specimens, in- 48557, 48617) and six female (UWBM 14484, 38795, cluding one of the paratypes (UWBM 18613). For the 42012, 43071, 44195, 44402) pelagicus, and six male purpose of evaluating the validity of each character (UWBM 48619, 50610, 52084, 52090, 52091, 52094) in diagnosing kenyoni, we assembled a reference se- and six female (UWBM 50611, 50612, 52086, 52089, ries of six male (UWBM 22442, 44193, 44194, 44403, 52093, 52107) urile, the other two "Stictocarbo" shags 120? 150i 180 150? 120? ARCTIC OCEAN LOCALITIES V Valdez & Southeast Alaska A Aleutian Islands P Pribilofs & Northern Alaska W British Columbia & Washington C California R RussiaRUSALSK sK J JapanRUSAA 60 60 P(3) V(153) CANADA (7) ~~~A(18) W(6) CHINA 1 40'- ~~~~~~~~~~~~~~~~~~~~~~~~U.S.A.40 C(8) JM1 PACIFIC OCEAN km 0 500 1000 20 -, , , , _ , - 20O 120? 150? 180c 150c 120O FIG. 1. North Pacific localities for the 196 pelagicus specimens used in the morphometric analyses. Symbols for the locality names are followed by sample sizes in parentheses. This content downloaded from 129.237.46.100 on Thu, 18 Sep 2014 14:09:14 PM All use subject to JSTOR Terms and Conditions 310 ROHWER ET AL. [Auk, Vol. 117 TABLE 2. Description of the 14 skeletal charactersused in the morphometricanalyses. Terminologyused in the characterdescriptions follows Baumel and Witmer (1993). Character Description Cranium depth Distance from center of external base of cranium to highest point of calvaria (skull cap) Cranium length From craniofacialflexion zone (hinge) to dorsal rim of foramen mag- num Maxilla length From craniofacialflexion zone (hinge) to rostrum of maxilla (tip of premaxilla) Coracoid length From acrocoracoidto lateral process of sternal end of coracoid Humerus length From head of humerus to ventral condyle at its distal end Ulna length From olecranon to ventral condyle at its dorsal end Carpometacarpuslength From carpal trochlea to distal end of major metacarpal Sternum length From dorsal spine of sternal rostrum to caudal border of sternal body at midline Sternum width Transversedistance between the first costal processes of each side Synsacrum length From corpus (body) of cranial-mostsynsacral vertebrato that of the caudal-most synsacral vertebra Pelvis width Distance between left and right antitrochantersthat form the dorsal rims of the acetabula Femurlength From tip of femoral trochanterto lateral condyle at its distal end Tibiotarsus length From patellar articularsurface of the cranial cnemial crest to medial condyle at its distal end Tarsometatarsuslength From the intercondylareminence at its proximal end to the most distal metatarsaltrochlea in the region. All urile reference specimens were USNM. After using these specimens to learn to re- from Alaska. It was important that the pelagicus ref- peat our measurements, Brian Schmidt measured the erence series included large birds taken outside the kenyoni type for us. Aleutians because kenyoni is small and has been re- Resolvingproblems with the morphometricdata.-Al- ported only from the Aleutians. For pelagicus males, though most of the Valdez birds were in good con- these specimens came from Prince William Sound, dition, some were too rotten to be sexed by gonads, Alaska (n = 2), Russia (n = 3), and Washington (n = and two were initially misidentified to species by 1); for pelagicus females, these specimens came from their preparators. Additionally, a few specimens Russia (n = 2), Washington (n = 3), and British Co- from the Valdez collection and from other museums lumbia (n = 1). The birds from Russia and Prince had broken elements that could not be measured. We William Sound were large. Each character was scored have dealt with these problems as follows. on each specimen by Bostwick and Peterson to eval- Specimens with missing characters were included uate the hypothesis that these qualitative characters in the morphometric analyses only if two or fewer diagnose the species. characters were missing. Twenty-seven of the 225 Quantitative characters.-For our morphometric specimens we measured had missing characters; 1 analyses, we used the 14 skeletal measurements de- with many missing characters was excluded, 23 had fined in Table 2. All were large enough to be taken a single missing character, and 3 had two missing accurately with dial calipers. Some of these mea- characters. We estimated values for missing charac- surements are difficult to replicate unless other ters in univariate regressions using the character workers first learn to match the measurements we most strongly correlated with the missing character made for individual specimens (available from Roh- as an estimator. All species and sexes were combined wer or Filardi). The problem lies not in the repeat- for this regression analysis because sex and species ability of our measurements, but in interpreting our determinations had not yet been evaluated. We later definitions for them (Table 2). Unless the same mea- reassessed these estimates using correlations be- surements are taken, errors between observers seri- tween characters for within-sex and within-species ously affect results. With one exception (see below), analyses; however, changes in estimates were so Filardi made all measurements. small (typically <1 mm) that reestimates were not The holotype specimen for kenyoni is housed at the used.
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