Giles: Multivariate Analysis of Pleistocene and Recent Coyotes 



375 



and a small, weak skull. The San Pedro Martir Coyote is medium-sized, broad- 

 skulled, and has a predominantly dark or reddish coat. 



Jackson (Young and Jackson, 1951) lists certain cranial characters that he 

 considers indicative of the various subspecies (table 1). This table presents a good 

 case for the use of multivariate analysis : it is clear that assessing by eye the rela- 



TABLE 1 



Some Cranial Characteristics of Coyote Subspecies 



Character 



C. 1. ochropus 



C. 1. mearnsi 



C. 1. testes 



C. 1. clepticus 



C. 1. OTCuttX 



Frontal region 



Nearly flat 



Depressed 



Flat 







Zygomatic arches 



Narrow 





Very broad 



Rostrum 













Breadth 



Narrow 





Narrow 



Broad 



Broad 



Length 



Long 



Short 



Long 



Short 





Skull 













Size 





Small 



Large 



Broad 



Large 



Breadth 



Narrow 



Medium 



Dentition 



Weak 



Weak, small 



Heavy 



Moderate 



Heavy 



Source: Young and Jackson (1951). 



tive merit of all these characters would be difficult. Certainly no single character is 

 sufficient to distinguish any one group. 



Grinnell (1933:113) states the following in regard to subspecific differentiation: 



Indeed, the three races which are here recognized are not sharply set off ; many specimens are 

 so indeterminable by any character whatsoever that their assignment on any save geographic 

 grounds has proved impossible to me. In other words, not only is there intergradation in the 

 usual subspecific sense, but individual variation is so great (at least in the characters of skulls 

 and teeth) that frequent specimens even from the metropolis of one or another of the races do 

 not fit that race ! 



The fossil canids from Rancho La Brea, as described by Merriam (1910, 1912) 

 and Stock (1938) can be arranged by size, smallest to largest, as follows: Urocyon 

 calif ornicus Mearns, the gray fox; Canis andersoni J. C. Merriam, a small coyote; 

 C. latrans orcutti, the coyote; C. petrolei Stock, a more wolf like coyote; C. lupus 

 furlongi J. C. Merriam, the wolf; C. milleri J. C. Merriam, intermediate between 

 C. I. furlongi and C. (Aenocyon) dirus Leidy, the dire wolf. 



The coyotes from the Rancho La Brea pits were first given specific rank by 

 Merriam (1910) as Canis orcutti. Two years later Merriam (1912) reduced the 

 group to the status of a subspecies of the California Valley Coyote: C. ochropus 

 orcutti. Hay (1927) regarded the orcutti group as a distinct species, but later 

 workers have considered them to be only a subspecies (e.g., Schultz, 1938). As all 

 Recent coyotes have been placed in one species, C. ochropus is a synonym of C. 

 latrans ochropus. Thus the Rancho La Brea form will be considered here as C. I. 

 orcutti. 



The characters by which Merriam (1912) distinguished Canis latrans orcutti 

 from other coyotes, especially C. I. ochropus, are: somewhat larger skull, broader 

 across the palate and zygomatic arches, higher and thicker mandible, and thicker 

 carnassials. 



