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PELVIC and THIGH MUSCLES of ORNITHORHYNCHUS by HELGA S

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PELVIC and THIGH MUSCLES of ORNITHORHYNCHUS by HELGA S PELVIC AND THIGH MUSCLES OF ORNITHORHYNCHUS By HELGA S. PEARSON University College, London OF recent years attempts to establish homologies between the muscles found in one class of Vertebrates and those in another have been placed on a much sounder basis by the study of the fossil bones of common ancestral forms. These bones often show muscle insertions as well marked as those on the bones of animals living to-day. Moreover, being often intermediate in shape, they indicate how the skeletal differences between living groups have arisen, differences intimately connected with the different position and action of the muscles moving that skeleton. Watson in England, Gregory and Camp in America, were pioneers in this line of work. More recently Romer', using similar methods and also considering the nerve supply, has made a very careful comparative study of the locomotor apparatus in Amphibia, Reptiles and Mammals with especial reference to the mammalian line of descent. Of these three classes he himself dissected three living types, Cryptobranchus, Iguana and Didelphys, but at that time no Monotreme, relying on Westling's descrip- tion of conditions in this group. Recently I have had the opportunity ofdissecting the hind limb musculature of two specimens of Ornithorhynchus (kindly lent me for this purpose by Prof. D. M. S. Watson). I find that, in the light of Romer's results from other groups of vertebrates, many of Westling's homologies can no longer be ac- cepted; moreover they are unaccompanied by any figures. Although since the days of Cuvier and Meckel there have been other, partially illustrated, descrip- tions by Mivart, Alix, Coues, Manners-Smith and Frets, the homologies of these investigators are for the most part equally unacceptable, so that a new and fully illustrated description of these muscles seems to be needed. I have myself dissected specimens of Romer's three types, and also of other lizards and of the marsupial Sarcophilus, and I have had the opportunity of discussing some of the difficult points with Dr Romer himself, convincing myself of the validity of almost all his homologies. Most of the pelvic and thigh muscles of Ornithorhynchus can readily be brought into line with these. As might be expected, they are essentially mammalian in their arrangement but sometimes still suggest a reptilian origin. A few appear to be unique. 1 Romer (1922), "The Locomotor Apparatus of Certain Primitive and Mammal-like Reptiles. Bull. Am. Mu8. Nat. Hist. XLVI. Pelvic and Thigh Mucltes of Ormithorhynchus 153 I. MUSCLES SEEN WITH THE ANIMAL ON ITS SIDE "Cruro-coccygeus." Origin. In front by a few fibres arising from the anterior end of the iliac blade at its internal corner. Then along a short tendinous raphe stretching between this and the spine of the first sacral vertebra superficial to spinalis and semispinalis lumborum. Then along the spines of the first sacral to the fifth caudal vertebrae. Then from the fascia of the tail external to all the tail muscles, and a little way on to the ventral surface of the tail beneath the transverse process of the fifth caudal vertebra (or thereabouts). Insertion. (a) By a wide, flat tendon which passes round the back of the ankle, onto the under surface ofthe foot internal to the origin of gastrocnemius and (in the male) external to the spine. Here it is superficial to all the tendons of the leg muscles. (b) By a partly fleshy, partly tendinous insertion on to the preaxial border of the tibia, internal to gracilis and just distal to the insertion of semimem- branosus and semitendinosus. This enormous muscle is the hardest of all the monotreme muscles to inter- pret. I should judge it to be used by the animal in kicking back the earth scraped outfrom the burrow, and know of nothing that really corresponds to it in any reptile or higher mammal. Against its being part of glutaeus maximus, as it has usually been named (e.g. Manners-Smith', Westling2), is its insertion; this is on the lower leg, to which it passes superficially to biceps, instead of on the femur internal to biceps. Its mode of insertion would seem rather to relate it to a part of the reptilian flexor mass, the origin of which had not travelled down the ilio-ischiadic (mammalian sacro-ischiadic) ligament to the tuber ischii, as that of the semimembranosus and semitendinosus of most mammals has done. It may be related to the cruro-coccygeus of Didelphys, a slender muscle passing from the side of the tail to the lower leg, and represented in various other mammals by slips joining or accompanying semitendinosus or biceps (Romer, p. 573). This, Romer suggests, may further represent the long tendon of the reptilian caudi-femoralis (caudi-femoralis longus). The primitive caudi-femoralis is, as Romer points out, closely associated with the long flexor muscles, inserting into the flexor mass in Urodeles and primitive frogs. Frets3 calls the anterior part of the disputed muscle in Ornithorhynchus glutaeus maximus, the hinder part caudo-femoralis. Biceps [Reptilian ilio-fibularis; Romer, p. 564]. Origin. From the ischial tuberosity. Passes deep to "cruro-coccygeus" and superficial to all the femoral muscles. 1 Manners-Smith (1885), "On some points in the Anatomy of Ornithorhynchlu paradoxwu," Proc. Zool. Soc. 2 Westling (1889), "Anatomische Untersuchungen uber Echidna," Bihang k. Svenska Vet.- Akad. Handlingar, xv. 3 Frets (1910), "tOber den plexus Jumbo-sacralis, U.s.W.," AMorphol. Jahrb. xL. Anatomy ix 11 154 Helga S. Pearsron --BiC. Fig. 1. The right pelvic and thigh muscles of Ornithorhynchu8, as seen with the animal on its side. BIC. =biceps; CLO. =cloacal pouch; COC. =ischio-coccygeus; GAS. =gastro- cnemius; GLU. 1, 2. = glutaeals; PYR. = pyriformis; REC. = rectus; VAS. = vastus; X. = "cruro-coccygeus." Pelvic and Thigh Mumcle8 of Ornithorhynchus 155 Insertion. Into the fascia covering the lower leg muscles on the anterior and external surface of the lower leg. Glutaeals [Reptilian ilio-femoralis; Romer, p. 570]. In Ornithorhynchus there are two glutaeal muscles. (a) ( = " Glutaeus medius " of Westling and Frets). Origin. By a thin tendinous margin immediately under the fleshy origin of " cruro-coceygeus." Passes to the femur superficial to the axial musculature and to the other glutaeal. Insertion. On the posterior (external) border of the femur at the distal extremity of the crest running down that border from the great trochanter. The area of insertion is small, and distal to the long one of the other glutaeal. (b) ( = " Glutaeus minimus " of Westling and Frets). Origin. From nearly the whole of the glutaeal fossa of the ilium. Passes backwards and outwards, covering the origins of rectus and vastus. Insertion. On to the dorsal surface of the great trochanter and the crest running from that trochanter down the posterior (external) border ofthefemur. The first of these two glutaeals, (a), corresponds most nearly to the Eu- therian glutaeus maximus, the second, (b), to a fused glutaeus medius and minimus. In neither of my specimens was there any sign of a division of this second glutaeal into two, as described by Manners-Smith. Pyriformis [Reptilian caudi-ilio-femoralis? Romer, p. 572]. Origin. From the transverse process of the second caudal vertebra just dorsal to the anterior fibres of ischio-coccygeus. Insertion. On the ventral surface of the femur beyond the obturator inser- tions, and close to the most distally inserting part of the glutaeus, on the posterior (external) border of the bone. For this, as for the other muscles from tail to limb, it is hard to find the reptilian equivalent. The American workers have suggested that it is the reptilian caudi-ilio-femoralis (coccygeo-femoralis brevis). This in reptiles "inserts by a tendon into the femur part way down the shaft, reaching the bone between ilio-femoralis and the adductors" (Romer, p. 572). In higher mammals pyriformis inserts on the ventral surface of the great trochanter. The monotreme muscle, although undoubtedly representing the pyrifornis of higher forms, has an insertion some way down the shaft of the femur, in the position of the reptilian caudi-ilio-femoralis. This makes the suggested homology a very probable one. Rectus [= Reptilian caput iliacus triceps or extensor ilio-tibialis; Romer, p. 562]. Origin. From a small area on the ilium immediately in front of the acetabu- lum. There is a distinct tubercle for it at the acetabular end ofthe ridge between the glutaeal and iliac fossae of the ilium. f 11-2 156 Helga S. Pearson Passes between the glutaeals and ilio-psoas and then superficial to vastus. Its insertion may be taken with that of vastus below. Vastus [= Caput femoralis triceps or femoro-tibialis; Romer, p. 564]. Origin. From the whole of the dorsal surface of the femur except for the insertion places of the glutaeals and ilio-psoas on the dorsal surfaces of the two trochanters. The muscle has a thick fleshy body lying between vastus and the femur. Insertion. Together with rectus on the patella and patella ligament. Sartorius [= Caput acetab. triceps or Ambiens; Romer, p. 563]. Origin. From the extremity of the ilio-pectineal spine (the long process projecting forwards from the anterior border of the pubis just below the ilio- pubic symphysis). Insertion. On the front of the knee joint, proximally to and in a line with the insertion of gracilis, and at some little distance beyond the insertion of the adductors. As pointed out by Romer the sartorius of monotremes, like the ambiens of reptiles, arises from the pubis in front of and a little below the acetabulum, instead of from the superior iliac spine as in most higher mammals. Mono- tremes are unique, however, in having the anterior border of the pubis in this region drawn out into what I have termed above the ilio-pectineal spine, and it is from the tip of this spine that sartorius takes origin.
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