Non-metric variation of the middle phalanges of the human toes (II-V): long/short types and their evolutionary significance.

The human lateral toes are characterised by extreme reduction compared with other primates, and in particular other hominoids. Some phalangeal non-metric variants have been well identified in humans, in particular: triphalangeal/biphalangeal patterns, and the presence/absence of phalangeal secondary centres of ossification. The purpose of the present study was to describe and analyse an original non-metric variation of the middle phalanges of the lateral toes. The material consisted of 2541 foot radiographs that came from 2541 different European adult individuals. Two morphological types of the middle phalanx were defined as a simple binary trait: long type (L) and short type (S). In feet with a triphalangeal pattern in all lateral toes (1413 cases), a mediolateral increasing gradient was observed in the occurrence of type S: 8.1% in II; 30.7% in III; 68.4% in IV; and 99.1% in V. In feet with a biphalangeal pattern in one or more lateral toes (III-V; 1128 cases), type S occurred more frequently than in triphalangeal feet. Of the 30 theoretical arrangements of the L/S types in the lateral toes (II-V) in a complete foot, only 13 patterns were observed. Seven patterns represented 95.6% of the population: LLSS (20.9%), LLLS (17.1%), LSS (15.9%), SSS (14.5%), LSSS (12.7%), LLS (10.1%) and SSSS (4.4%). Type L can be interpreted as the primitive pattern (plesiomorphy), and type S as a derived pattern (apomorphy) that seems specific to the human species (i.e. autapomorphy). Within the specific evolution of the human foot in relation to the acquisition of constant erect posture and bipedalism, the short type of the middle phalanges can reasonably be considered as directly linked to the reduction of the lateral toes.