Fig. 1. Pelvis of Tympanuchus pallidicinctusFig. 1.Pelvis ofTympanuchus pallidicinctus. A. Lateral view. × 1. B. Ventral view. × 1⅛.
The terminology applied to the synsacral vertebrae by different authors varies. The terminology proposed by DuToit (1913) is employed in the present account. See myFig. 1B. This terminology differs considerably from that used by Howard (1929). DuToit divides the fused synsacral vertebrae into the following five groups, listed in anteroposterior sequence: (1)synsacro-thoracic, which bear movable ribs; (2)synsacro-thoraco-lumbar, which lack movable ribs but possess well developed laterally directed parapophyses, in addition to the more dorsally directed diapophyses; (3)synsacro-lumbar, which lack parapophyses, although possessing inconspicuous diapophyses; these vertebrae are shortened anteroposteriorly and are so firmly fused together that often the number present can be determined only by counting the intervertebral foramina; (4)synsacro-sacral, which have much more pronounced transverse processes than do the synsacro-lumbar vertebrae; these transverse processes are expanded distally where they fuse with the ilium and represent both parapophyses and diapophyses partly or completely fused together plus sacral ribs (detectable only in the embryo); there are considered to be two of these vertebrae; they are situated at approximately the level of the acetabulum; (5)synsacro-caudal, which include the remainder of the fused vertebrae; no marked gross morphological features differentiate the synsacro-sacral and the synsacro-caudal groups of vertebrae. The boundaries between all but the last two groups of vertebrae are usually, but not always, easily determined. It may be difficult to determine whether a vertebra with rudimentary parapophyses belongs to the synsacro-thoraco-lumbar or the synsacro-lumbar group. Sometimes a parapophysis will be better developed on one side of a vertebra than on the other.
Femur
Thetrochanteris a large squarish tuberosity on the lateral surface of the proximal end of the femur. Thetrochanteric ridgeis a sharp, longitudinal (relative to the femur) ridge forming the anterior edge of the trochanter. Theobturator ridgeis a short, blunt, longitudinal ridge forming the posterior edge of the trochanter. Theanterior intermuscular lineis a slight ridge extending distally from the trochanteric ridge. Theposterolateral intermuscular lineis a slight ridge extending distally from the obturator ridge. Theposterior intermuscular lineis a slight, longitudinal ridge on the mid-posterior surface of the femur. Theinternal condyleis a large rounded articular prominence on the medial side of the distal end of the femur. On the lateral side of the distal end of the femur are two articular prominences—the lateralmost, smaller one is thefibular condyle, separated by thefibular groove(visible from posterior aspect only) from the larger and more medialexternal condyle. Thepopliteal areais a depression on the posterior surface of the distal part of the femur immediately proximal to the condyles.
Tibiotarsus and Fibula
Theinner cnemial crestis pronounced and directed anteriorly on the anterior surface of the proximal end of the tibiotarsus. Theouter cnemial crestis pronounced and directed anterolaterally on the anterolateral surface of the proximal end of the tibiotarsus. Therotular crestis transverse and forms the anterior border of the proximal end of the tibiotarsus; the crest extends between the dorsal ends of the two cnemial crests and also extends medialto the inner cnemial crest. Thefibular crestis longitudinal on the lateral surface of the tibiotarsus and fuses with the middle part of the fibula. Thefibular tubercleis small and on the lateral surface of the fibula near the level of the middle of the fibular crest. Theanteromedial intermuscular lineis a slight ridge extending from the inner cnemial crest down the anteromedial surface of the tibiotarsus. Theanterolateral intermuscular lineis a slight ridge extending from the fibular crest down the anterolateral surface of the tibiotarsus. Thesupratendinal bridgeis a transverse bony arch over a longitudinal groove near the distal end of the anterior surface of the tibiotarsus.
Tarsometatarsus
Thehypotarsusis a large, pronounced, squarish protuberance on the posterior surface of the proximal end of the tarsometatarsus and contains grooves and canals for the passage of the flexor tendons. The longitudinal ridges forming the lateral and medial edges of the posterior surface of the hypotarsus are termedcalcaneal ridges. Theposterior metatarsal crestis long and sharp; it is continuous with the medial calcaneal ridge that extends most of the way down the posterior surface of the tarsometatarsus medial to the midline; there is an opening between this crest and the tarsometatarsus immediately distal to the hypotarsus. Themedial metatarsal depressionis large; it is on the medial surface of the proximal end of the tarsometatarsus. Theanterior metatarsal grooveis a longitudinal groove in the midline of the proximal part of the anterior surface of the tarsometatarsus. The threetrochleaeare large rounded articular prominences at the distal end of the tarsometatarsus; there is one at the base of each of the digits II, III, and IV. The termdistal foramen(as used by Howard) refers to a short, anteroposteriorly directed canal that perforates the tarsometatarsus a short distance proximal to the intertrochlear notch between the trochleae for digits III and IV. Beginning at the middle of this canal and extending distally at a right angle to it is theintertrochlear canal, which opens via the terminal foramen into the intertrochlear notch between the trochleae for digits III and IV.
Nerves
For ease of description I have coined terms for the major divisions of the femoral and sciatic nerves.
Muscles
My terminology follows that of Fisher (1946) and Fisher and Goodman (1955) except for Mm. femoritibialis externus, flexor cruris lateralis (accessory head), and obturator internus et externus. Fisher (1946:547) states that most of his names for the hip and thigh muscles are those of Howell (1938) and the names for the shank and foot muscles are those of Hudson (1937). Fisher deviates, without explanation, from Howell's terminology in respect to Mm. vastus medialis and femoritibialis internus, M. caudofemoralis, M. flexor cruris lateralis, and Mm. obturator internus and obturator externus. Fisher's synonymy of these muscles (1946: table 42)is in error. Fisher understandably deviates from Hudson in respect to Mm. extensor brevis digiti III and extensor proprius digiti III (see Holmes, 1962), although Fisher's synonymy is in error here. See my table 1.
I am not using Fisher and Goodman's term femoritibialis externus; this muscle is here considered as a part of M. vastus lateralis. A great deal of confusion surrounds the terminology of the muscle complex here termed Mm. vastus lateralis and vastus medialis. Hudson (1937), Hudson,et al.(1959), Fisher (1946), and Fisher and Goodman (1955) have used different terminology for this complex. Most of the confusion stems from Gadow's (1891) unclear description of this complex, which he subdivided into two units termed Mm. femori-tibialis externus and femori-tibialis medius. Many birds have three parts to this complex. It is difficult to determine how to apply Gadow's two terms to these three parts. As nearly as I can determine, the correct method is that of Hudson,et al.(1959); but because Gadow's terms have been used in different ways (even by the same worker), it seems best to abandon these terms. Berger (1956:272) believes that the muscle unit that Fisher and Goodman term M. femoritibialis externus represents a head of M. vastus lateralis; I am accepting his opinion. For the three parts of the complex under discussion, I am using the terms M. vastus medialis and M. vastus lateralis pars lateralis and pars postica.Fisher (Fisher, 1946; Fisher and Goodman, 1955) considers the muscle here termed M. femorocruralis as an accessory head of M. flexor cruris lateralis. The two muscle units in question are closely associated; they insert broadly on opposite sides of a common tendinous raphe. Howell (1938:73) considers this to be a secondary fusion of unrelated muscles. Romer (1927:366) states that in the chick embryo M. femorocruralis is in reality a shank muscle that migrates into the thigh during development. Therefore, Fisher's usage of a single name for these two unrelated muscles is unsatisfactory. I am using Howell's terminology in which the name flexor cruris lateralis represents the main head only of Fisher's M. flexor cruris lateralis and the name femorocruralis represents Fisher's accessory head.Gadow (1891) divides the obturator complex into two muscles (or muscle groups), which he terms M. obturator and Mm. accessorii M. obturatoris. He states that the former is homologous with the mammalian obturator internus and the latter with the obturator externus. Hudson (1937), accepting Gadow's homologies, renamed these muscles M. obturator internus and M. obturator externus. Nearly all subsequent workers have followed Hudson's terminology, with its implication that these muscles are homologous with the mammalian muscles of the same name. Howell (1938) is an exception. He points out (pp. 78, 79) that the obturator internus of Hudson is homologous with the obturator externus of mammals. His evidence is convincing: "In origin the obturator is somewhat suggestive of the mammalian obturator internus, for which it has uniformly been mistaken. That the latter interpretation is incorrect, however, is attested by the facts that it receives twigs of n. obturatorius within the pelvis, passesthroughthe obturator foramen rather than dorsal to the border of the ischium, and it is segregated from any muscle with tibial innervation. Insertion has shifted only to a slight and unimportant degree as compared with that of themammalian obturator externus, and beyond question it is the equivalent of that muscle. The stimulus for a longer muscle, has been the same, resulting in the extension of origin to within the pelvis of the externus in birds and the internus in mammals, but the obturator internus is an extension of a part of the gemellus mass and this does not occur in any vertebrate class but Mammalia." Howell applies the term M. obturator to the entire obturator complex.Romer (1927), studying the development of the thigh musculature in chick embryos, concluded that the entire obturator complex is homologous with the mammalian obturator externus plus quadratus femoris. He considered the avian M. flexor ischiofemoralis to be the homologue of the mammalian obturator internus.Gadow, in his work on the ratites (1880:34), states that M. obturator (obturator internus of Hudson) cannot be homologous to the mammalian obturator internus, but must represent the obturator externus. His reasoning is as follows: "Als M. pectineus kann man diesen Muskel nicht auffassen, da er auf der Aussenfläche des Trochanter major inserirt, ferner auch nicht als M. obturator internus der menschlichen Anatomie, da er nicht vom Plexus ischiadicus, sondern vom Plexus cruralis aus innervirt wird. Seiner Innervation und Insertion nach wäre er nur mit dem M. obturator externus zu vergleichen, wobei er seinen Ursprung im Verhältniss zum Menschen nur bedeutend weiter auf das Os ischii und Os pubis distalwärts ausgedehnt hätte und so allerdings der Lage nach mit Ausnahme seines Insertionsdrittels ein 'internus' geworden wäre."Since Gadow gives different homologues for M. obturator in two of his works (1880 and 1891), one would suspect that he had changed his opinion in the interim; however, there is no evidence that he did so. In 1880 he gives supporting evidence (quoted above) for his view; in 1891 he does not. After describing (1891:173) how the origin of M. obturator in bird ancestors presumably migrated from a location outside the pelvis to a position inside the pelvis prior to the meeting of the pubis and ischium external to the muscle, he states: "Eine ähnliche Entwicklung ist für denObturator internusder Säugethiere anzunehmen, welchem derM. obturatorder Vögel entspricht." A similar development in mammals is impossible, owing to the different relationship of the muscle to the pelvic bones in this class. Gadow says nothing more about the mammalian homologue of M. obturator. In view of this discrepancy, Gadow can hardly be considered as a supporter of the idea that the avian M. obturator is homologous with the mammalian obturator internus.The evidence is conclusive, it seems to me, that the obturator internus of Hudson is not homologous with the mammalian obturator internus. Therefore, the term obturator internus is inappropriate for the avian muscle and must be abandoned. I shall follow Howell (1938) in naming the entire obturator complex M. obturator. This term, of course, is not used in the sense in which it is used by Gadow. The use of the term obturator externus for the entire complex is avoided because it may not correspond exactly to the mammalian obturator externus. As mentioned previously, Romer considers the avian muscle to be homologous not only with the mammalian obturator externus but also with the quadratus femoris.
I am not using Fisher and Goodman's term femoritibialis externus; this muscle is here considered as a part of M. vastus lateralis. A great deal of confusion surrounds the terminology of the muscle complex here termed Mm. vastus lateralis and vastus medialis. Hudson (1937), Hudson,et al.(1959), Fisher (1946), and Fisher and Goodman (1955) have used different terminology for this complex. Most of the confusion stems from Gadow's (1891) unclear description of this complex, which he subdivided into two units termed Mm. femori-tibialis externus and femori-tibialis medius. Many birds have three parts to this complex. It is difficult to determine how to apply Gadow's two terms to these three parts. As nearly as I can determine, the correct method is that of Hudson,et al.(1959); but because Gadow's terms have been used in different ways (even by the same worker), it seems best to abandon these terms. Berger (1956:272) believes that the muscle unit that Fisher and Goodman term M. femoritibialis externus represents a head of M. vastus lateralis; I am accepting his opinion. For the three parts of the complex under discussion, I am using the terms M. vastus medialis and M. vastus lateralis pars lateralis and pars postica.
Fisher (Fisher, 1946; Fisher and Goodman, 1955) considers the muscle here termed M. femorocruralis as an accessory head of M. flexor cruris lateralis. The two muscle units in question are closely associated; they insert broadly on opposite sides of a common tendinous raphe. Howell (1938:73) considers this to be a secondary fusion of unrelated muscles. Romer (1927:366) states that in the chick embryo M. femorocruralis is in reality a shank muscle that migrates into the thigh during development. Therefore, Fisher's usage of a single name for these two unrelated muscles is unsatisfactory. I am using Howell's terminology in which the name flexor cruris lateralis represents the main head only of Fisher's M. flexor cruris lateralis and the name femorocruralis represents Fisher's accessory head.
Gadow (1891) divides the obturator complex into two muscles (or muscle groups), which he terms M. obturator and Mm. accessorii M. obturatoris. He states that the former is homologous with the mammalian obturator internus and the latter with the obturator externus. Hudson (1937), accepting Gadow's homologies, renamed these muscles M. obturator internus and M. obturator externus. Nearly all subsequent workers have followed Hudson's terminology, with its implication that these muscles are homologous with the mammalian muscles of the same name. Howell (1938) is an exception. He points out (pp. 78, 79) that the obturator internus of Hudson is homologous with the obturator externus of mammals. His evidence is convincing: "In origin the obturator is somewhat suggestive of the mammalian obturator internus, for which it has uniformly been mistaken. That the latter interpretation is incorrect, however, is attested by the facts that it receives twigs of n. obturatorius within the pelvis, passesthroughthe obturator foramen rather than dorsal to the border of the ischium, and it is segregated from any muscle with tibial innervation. Insertion has shifted only to a slight and unimportant degree as compared with that of themammalian obturator externus, and beyond question it is the equivalent of that muscle. The stimulus for a longer muscle, has been the same, resulting in the extension of origin to within the pelvis of the externus in birds and the internus in mammals, but the obturator internus is an extension of a part of the gemellus mass and this does not occur in any vertebrate class but Mammalia." Howell applies the term M. obturator to the entire obturator complex.
Romer (1927), studying the development of the thigh musculature in chick embryos, concluded that the entire obturator complex is homologous with the mammalian obturator externus plus quadratus femoris. He considered the avian M. flexor ischiofemoralis to be the homologue of the mammalian obturator internus.
Gadow, in his work on the ratites (1880:34), states that M. obturator (obturator internus of Hudson) cannot be homologous to the mammalian obturator internus, but must represent the obturator externus. His reasoning is as follows: "Als M. pectineus kann man diesen Muskel nicht auffassen, da er auf der Aussenfläche des Trochanter major inserirt, ferner auch nicht als M. obturator internus der menschlichen Anatomie, da er nicht vom Plexus ischiadicus, sondern vom Plexus cruralis aus innervirt wird. Seiner Innervation und Insertion nach wäre er nur mit dem M. obturator externus zu vergleichen, wobei er seinen Ursprung im Verhältniss zum Menschen nur bedeutend weiter auf das Os ischii und Os pubis distalwärts ausgedehnt hätte und so allerdings der Lage nach mit Ausnahme seines Insertionsdrittels ein 'internus' geworden wäre."
Since Gadow gives different homologues for M. obturator in two of his works (1880 and 1891), one would suspect that he had changed his opinion in the interim; however, there is no evidence that he did so. In 1880 he gives supporting evidence (quoted above) for his view; in 1891 he does not. After describing (1891:173) how the origin of M. obturator in bird ancestors presumably migrated from a location outside the pelvis to a position inside the pelvis prior to the meeting of the pubis and ischium external to the muscle, he states: "Eine ähnliche Entwicklung ist für denObturator internusder Säugethiere anzunehmen, welchem derM. obturatorder Vögel entspricht." A similar development in mammals is impossible, owing to the different relationship of the muscle to the pelvic bones in this class. Gadow says nothing more about the mammalian homologue of M. obturator. In view of this discrepancy, Gadow can hardly be considered as a supporter of the idea that the avian M. obturator is homologous with the mammalian obturator internus.
The evidence is conclusive, it seems to me, that the obturator internus of Hudson is not homologous with the mammalian obturator internus. Therefore, the term obturator internus is inappropriate for the avian muscle and must be abandoned. I shall follow Howell (1938) in naming the entire obturator complex M. obturator. This term, of course, is not used in the sense in which it is used by Gadow. The use of the term obturator externus for the entire complex is avoided because it may not correspond exactly to the mammalian obturator externus. As mentioned previously, Romer considers the avian muscle to be homologous not only with the mammalian obturator externus but also with the quadratus femoris.
I am following the policy of Wilcox (1948) and Berger (1952) in latinizing the term anterior, changing it to anticus. When preceded by the feminine word pars, the feminine ending is used (antica).In table 1 my terminology is compared with that of Fisher and Goodman (1955), Howell (1938), Hudson (1937), and Gadow (1891). The terminology of Fisher (1946) is identical with that of Fisher and Goodman (1955) except that in his earlier work Fisher did not describe or name M. femoritibialis externus, and M. lumbricales of his earlier work is not mentioned in his later work. The terminology of Hudson,et al.(1959) is identical with that of Hudson (1937) except that the manner in which the femoritibialis complex is subdivided is identical with that of Gadow (1891) and different from that in Hudson's earlier work; also the abbreviations p. ext. and p. int. are substituted in his later paper for pars anterior and pars posterior, respectively, of M. adductor longus et brevis.
I am following the policy of Wilcox (1948) and Berger (1952) in latinizing the term anterior, changing it to anticus. When preceded by the feminine word pars, the feminine ending is used (antica).
In table 1 my terminology is compared with that of Fisher and Goodman (1955), Howell (1938), Hudson (1937), and Gadow (1891). The terminology of Fisher (1946) is identical with that of Fisher and Goodman (1955) except that in his earlier work Fisher did not describe or name M. femoritibialis externus, and M. lumbricales of his earlier work is not mentioned in his later work. The terminology of Hudson,et al.(1959) is identical with that of Hudson (1937) except that the manner in which the femoritibialis complex is subdivided is identical with that of Gadow (1891) and different from that in Hudson's earlier work; also the abbreviations p. ext. and p. int. are substituted in his later paper for pars anterior and pars posterior, respectively, of M. adductor longus et brevis.
I gratefully acknowledge the generous help of Professor A. Byron Leonard, under whose guidance this study was conducted and thank Professor E. Raymond Hall, Professor Howard A. Matzke, and Dr. Irwin Baird for numerous helpful suggestions and criticisms.
For help in collecting specimens I thank J. R. Alcorn, W. C. Glazener (through the courtesy of the Texas Game and Fish Commission), Dr. Harrison B. Tordoff, Jerry Tash, William Brecheisen, and Louis Brecheisen. I thank also Edwin Gebhard of the Kansas Forestry, Fish and Game Commission for help in locating the Lesser Prairie Chickens.
I am grateful for the assistance of Mrs. Chester Alexander and Dr. L. C. Dahl in translating a Russian and a Dutch reference, and thank George Young and James Bee for making equipment used in my study.
All of the original drawings exceptFig. 1were made by me, although the final inking ofFigs. 12through19was done by Bret Waller.Fig. 1was drawn by Kay Swearingen.
I was aided in this study during the summer of 1960 by a research grant from the University of Kansas.
Although no special study was made of the skeleton, certain conspicuous variations are discussed here.
There are a few pronounced differences between the pelvis ofTympanuchusand that ofPedioecetes. Whereas in the former the thick lateral iliac process has a pronounced overhang with the ventral edge lateral to the ischium (Fig. 1), inPedioecetesthere is no overhang at all and the edge of this process is much thinner. The ischium inPedioecetesis wider (in dorsoventral extent), especially posteriorly, than inTympanuchus. InTympanuchusthe posteroventral margin of the ischium is rounded and is free from the pubis, whereas inPedioecetesit is pointed and fused with the pubis.
InTympanuchus cupido(both subspecies) the lateral iliac process extends farther ventrally than inT. pallidicinctus, approaching or extending ventral to the level of the pubis in the former species; also the edge of this process is thicker inT. cupido.
All specimens studied have a single synsacro-thoracic vertebra. The number of combined synsacro-thoraco-lumbar and synsacro-lumbar vertebrae is eight in each specimen ofTympanuchusand in one specimen ofPedioecetes phasianellus jamesiand is seven in three specimens of the latter. In most specimens ofTympanuchusthere are three synsacro-thoraco-lumbar and five synsacro-lumbar vertebrae, although in two specimens (T. pallidicinctus) there are four of each group; in one of these latter two specimens the parapophysis on one side of the fourth synsacro-thoraco-lumbar vertebra is small. The first (of five) synsacro-lumbar vertebra has a rudimentary parapophysis on one side in one specimen ofTympanuchusand on both sides in another specimen. One specimen ofPedioecetes phasianellus jamesihas five synsacro-lumbar vertebrae and the others have four; all have three synsacro-thoraco-lumbar vertebrae.
For each nerve (or plexus) the condition found in most specimens of the Lesser Prairie Chicken (T. pallidicinctus) is described first. Following this, variations from the typicalT. pallidicinctuscondition are given forT. pallidicinctus, then forT. cupido(both subspecies considered together), and finally forP. p. jamesi.
Lumbosacral Plexus,Figs. 2,3
T. pallidicinctus
Description.—Eight spinal nerves contribute to the lumbosacral plexus. These are the second through the ninth synsacral spinal nerves (S2 to S9). The entire ventral ramus of each of these nerves, excepting S2 and S9, contributes to this plexus. The ventral ramus of S2 divides into two branches, only the posterior of which contributes to the plexus; the anterior branch directly innervates muscles of the abdominal wall (as does the entire ventral ramus of S1). The ventral ramus of S9 divides into two branches, only the anterior of which contributes to this plexus; the posterior branch contributes to the more posteriorly situated pudendal plexus.
Each root of the plexus corresponds to a single spinal nerve except one spinal nerve (S5—the furcal) that contributes a root to both the femoral nerve and the sciatic nerve; thus typically the plexus has nine roots (but see below). The four anteriormost roots (S2 to S5) contribute to the femoral nerve, although the contribution from S2 is small. S3 and S4 contribute to the obturator nerve. The five posteriormost roots (S5 to S9) contribute to the sciatic nerve, although the contribution from S9 is relatively small.
Individual Variation.—In all specimens (of all species) examined, the right and left sides of the plexus in any one individual were practically identical. In T.p. 2 (Fig. 2B), there appear to be two furcal nerves; S5 is typical, but a small branch of S4 apparently also contributes to the sciatic nerve. In T.p. 5, S9 is unique in dividing into three branches; the anterior two join the sciatic nerve separately; the posterior one joins the pudendal plexus as usual.
T. cupido
Individual Variation.—S2 or S5, or both, may contribute to a limited extent to the obturator nerve. In T.c.p. 3 (Fig. 3A) and T.c.a. 1 and 2, much of the plexus has shifted one segment anteriorly, relative to the synsacral vertebrae (the so-called prefixed condition); the roots of the femoral nerve are S2, S3, and S4 (all large); the furcal nerve is S4 (in T.c.a. 1, S5 gives an extremely small root to the femoral nerve, thus making two furcal nerves); six roots (S4 to S9) contribute to the sciatic nerve; S3 and S4 remain as the main contributors to the obturator nerve except in T.c.a. 2 in which only S2 and S3 contribute to it.
P. p. jamesi
Individual Variation.—In P.p. 1, the plexus resembles the typical condition inT. pallidicinctus. In P.p. 2, 3, and 4, the plexus is prefixed. P.p. 2 resembles T.c.p. 3. In P.p. 3 and 4 (Fig. 3B) there are two furcal nerves (S4 and S5); S2 to S4 are the main contributors to the femoral nerve; only S2 and S3 contribute to the obturator nerve; S4 to S9 contribute to the sciatic nerve (the anteriormost and posteriormost roots are small).
Femoral Nerve,Figs. 4,5
T. pallidicinctus
Description.—The femoral nerve is short, dividing inside the pelvis into six major divisions—anterior, middle, posterior, anterodorsal, dorsal, and posterodorsal. The anterodorsal and posterodorsal divisions are short, failing to extend so far laterally as the inguinal ligament; the posterodorsal division is also small and is usually covered by other divisions and is not visible when viewed from the ventral side.
The anterior division passes ventral to Mm. iliotrochantericus medius and iliacus and dorsal to the anterior end of the inguinal ligament. The division branches into two parts. The anterior part extends around the posterior border of M. extensor iliotibialis anticus and sends several twigs to the lateral surface of this muscle. The posterior part passes between the proximal parts of Mm. extensor iliotibialis anticus and extensor iliotibialis lateralis and supplies the skin.
The middle division passes ventral to Mm. iliotrochantericus medius and iliacus and dorsal to the inguinal ligament. The division branches into a large but variable number of parts. A variable number of branches (usually two) pass posterior to M. extensor iliotibialis anticus and penetrate the medial surface of M. extensor iliotibialis lateralis. Several branches supply the fused Mm. vastus lateralis and vastus medialis. The posteriormost branch of this division passes between Mm. ambiens and vastus medialis,giving twigs to the lateral surface of M. ambiens, and sometimes also to the medial surface of M. vastus medialis, and terminates in M. femoritibialis internus.
The posterior division, which does not subdivide, spirals completely around M. psoas (passing in turn anterior, dorsal, posterior, and ventral to it) and gives twigs into this muscle. This nerve then extends distally into the proximal part of the shank and there has a nonmuscular termination.
The short, thick anterodorsal division, partly covered by the anterior division, turns dorsally and passes through the femoral notch of the ilium and penetrates the deep surface of M. gluteus profundus.
The slender dorsal division passes ventral to M. iliotrochantericus medius and dorsal to the inguinal ligament and penetrates the ventral surface of M. iliacus.
The small, short posterodorsal division penetrates the ventral surface of M. iliotrochantericus medius.
Individual Variation.—In two legs the anterior division gives a twig or two twigs to M. extensor iliotibialis lateralis. The dorsal division may fuse proximally with either the anterior or middle division, thus appearing to be a branch of one of these divisions. In one leg (Fig. 5A), there are two separate branches (both fused with the middle division) to M. iliacus. On both sides of one specimen (Fig. 5A), the anteriormost branch of the middle division, which supplies M. extensor iliotibialis lateralis, gives off a twig that anastomoses with the branch of the anterior division that supplies M. extensor iliotibialis anticus. On both sides of another specimen, the anterodorsal division passes lateral to the anterior end of M. iliotrochantericus medius instead of through the femoral notch, which is lacking.
T. cupido
Individual Variation.—In three legs, the anterior division gives twigs into M. extensor iliotibialis lateralis. The dorsal division is fused proximally with the middle division in one instance. In three cases, a twig from the middle division anastomoses with the branch of the anterior division supplying M. extensor iliotibialis anticus. In the example shown inFig. 5B, a twig comes off the cutaneous branch of the anterior division, perforates the ventral part of M. iliacus, and rejoins the cutaneous branch. In both legs of one specimen, the cutaneous branch of the anterior division perforates the anterior edge of M. extensor iliotibialis lateralis instead of passing between the latter and M. extensor iliotibialis anticus. The posteriormost branch of the middle division, which terminates in M. femoritibialis internus, perforates the medial part of M. vastus medialis in one leg. In another leg, one of the branches to the fused Mm. vastus lateralis and vastus medialis sends a twig into M. extensor iliotibialis lateralis.
P. p. jamesi
Individual Variation.—In three legs, the anterior branch of the anterior division is cutaneous and the posterior branch supplies M. extensor iliotibialis anticus. The dorsal division may fuse proximally with either the anterior or middle division. In one leg (Fig. 4B), there are two branches to M. iliacus, one associated with the anterior division and one with the middle division.
Obturator Nerve
T. pallidicinctus
Description.—The long slender obturator nerve passes along the oblique iliac crest and divides into several branches immediately before reaching the obturator foramen. One or two branches, which do not pass through the foramen, penetrate the superficial surface of M. obturator pars postica. Several small branches (variable in number and arrangement) pass through the obturator foramen and supply pars ventralis, pars dorsalis, and pars antica of M. obturator. When pars ventralis and pars dorsalis are fused, one branch perforates the proximal end of this mass and reaches pars antica. One large branch passes through the obturator foramen dorsal to the tendon of M. obturator pars postica, then turns ventrally, passing lateral to the latter; the branch passes between Mm. adductor superficialis and adductor profundus and gives twigs to each of these two muscles.
Individual Variation.—None of significance in any of the three species.
Sciatic Nerve,Figs. 6,7,8,9
T. pallidicinctus
Description.—The sciatic nerve passes through the anterior part of the ilio-ischiatic fenestra. Several branches diverge from the nerve immediately after it emerges from the fenestra. The main trunk of the nerve then extends distally through the thigh deep to M. extensor iliofibularis and superficial (lateral) to Mm. flexor ischiofemoralis, caudofemoralis, adductor superficialis, and femorocruralis. The main trunk subdivides into two large nerves—peroneal and tibial—that are adjacent and bound to each other throughout the thigh; the peroneal nerve lies anterior to the tibial. At the distal end of the thigh the main trunk splits grossly into two large branches that diverge and enter the shank. This division does not represent the separation between peroneal and tibial nerves, as is sometimes assumed; the anterior branch includes a part of the tibial nerve as well as the entire peroneal nerve.
A longitudinal groove is visible grossly on the lateral surface of the main trunk, except at the proximal end; distally a second groove is visible posterior to the first one (Fig. 6). The long anterior groove indicates the boundary between the peroneal and tibial nerves; this groove may disappear distally, although the posterior groove is always visible distally. The posterior groove, which is continuous with the division of the sciatic nerve into anterior and posterior branches, represents the boundary between two divisions of the tibial nerve. (This is discussed in detail below.) In the middle of the thigh the peroneal and tibial nerves are enclosed in separate connective tissue sheaths, although the two sheaths are fused together; the point of fusion is marked by the anterior groove. If the two sheaths are slit open, the two nerves can be removed and can be seen to be entirely separate. In the proximal part of the main trunk the peroneal and tibial components are enclosed in a single sheath and appear as an undivided trunk; but if the sheath is removed, the two components can be pulled apart rather easily, although there may be some intermingling of a few fibers. This separation can be extended to a point proximal to the origin of all the branches of the sciatic nerve; thus it can be determined which branches arise from theperoneal component and which from the tibial. (These branches arise from the sciatic nerve as, or immediately before, the nerve passes through the ilio-ischiatic fenestra; since this level of the intact nerve could not be adequately observed, it was necessary to cut the nerve inside the pelvis and pull the intrapelvic part of the nerve out through the ilio-ischiatic fenestra. In doing this, care had to be taken to avoid damaging the most proximal branches.)
Three main branches arise from the peroneal component (apart from the main trunk) and two from the tibial. Including the peroneal and tibial components of the main trunk, the sciatic nerve can be divided into seven major divisions—anterior peroneal, middle peroneal, dorsal peroneal, posterior or main peroneal (contributes to main trunk), anterior or main tibial (contributes to main trunk), middle tibial, and posterior tibial. Farther distally, the posterior peroneal division becomes the peroneal nerve and the anterior tibial division becomes the tibial nerve. For descriptive purposes, the term peroneal (or tibial)nervewill be applied only where the nerve is enclosed in its own sheath, but regardless of whether or not the sheath is fused with another; proximal to this, where the separation may not be precise, the terms peroneal (or tibial)divisionorcomponentwill be used.
The small anterior peroneal division arises from the anterior edge of the sciatic nerve. Immediately after emerging from the ilio-ischiatic fenestra, the division turns anteriorly and passes deep to M. piriformis, to which the division gives a twig (in some cases more than one twig), then continues forward to supply the posterior part of M. gluteus profundus.
The middle peroneal division branches into two parts. One part penetrates the deep surface of the anteroproximal part of M. extensor iliofibularis. The other part emerges between the proximal ends of Mm. extensor iliofibularis and vastus lateralis and penetrates the deep surface of M. extensor iliotibialis lateralis.
The dorsal peroneal division arises from the posterodorsal part of the peroneal component, then angles posteriorly, crossing the dorsal surface of the anterior tibial division and superficially appears to arise from the tibial component. The dorsal peroneal division usually subdivides into two unequal branches, both of which penetrate the deep surface of the proximal end of M. extensor iliofibularis.
The large middle tibial division soon subdivides into two branches that pass posterodistally lateral to M. flexor ischiofemoralis. One branch (usually the anterior one) passes lateral to M. caudofemoralis (both heads) and emerges between Mm. extensor iliofibularis and flexor cruris lateralis and enters the skin. The other branch passes deep to M. caudofemoralis pars iliofemoralis, and divides into several branches. Several tiny branches penetrate the deep surface of M. caudofemoralis pars iliofemoralis. Another branch also enters the substance of the latter and emerges from the ventral edge of it, giving a twig to pars caudifemoralis, then passes lateral to pars caudifemoralis and enters M. flexor cruris lateralis. Still another branch passes deep to both heads of M. caudofemoralis and enters the anterior part of M. flexor cruris medialis.
The small posterior tibial division arises from the posterior edge of the sciatic nerve. The division diverges from the remainder of the nerve, as thelatter passes through the ilio-ischiatic fenestra, and penetrates the dorsal surface of M. flexor ischiofemoralis.
Below the middle of the main trunk a bundle of fibers of moderate size separates from the anterior edge of the tibial nerve, leaves the tibial sheath, and enters its own sheath, lying superficially between the tibial and peroneal sheaths (Fig. 6). At the distal end of the thigh the sheath enclosing this bundle of fibers remains fused with the posterior edge of the peroneal nerve and passes with the latter (diverging from the remainder of the tibial nerve) through the tendinous guide loop for M. extensor iliofibularis, and then diverges from the peroneal nerve. Since this bundle of fibers is distributed with the peroneal nerve, and since the origin of the bundle may be easily overlooked, it has sometimes been misinterpreted as a branch of the peroneal nerve, whereas it almost certainly is a branch of the tibial nerve; this bundle will here be termed the paraperoneal branch of the tibial nerve.
A small but long branch separates from the posterior edge of the proximal end of the tibial nerve or from the tibial component proximal to this and extends distally for some distance adjacent to the tibial nerve, then passes posterodistally between Mm. extensor iliofibularis and flexor cruris lateralis and supplies the skin.
A small branch separates from the anterior edge of the peroneal nerve a short distance above the distal end of the main trunk and passes distolaterally between Mm. extensor iliotibialis lateralis and extensor iliofibularis and supplies the skin.
A twig comes off the medial surface of the tibial nerve near the distal end of the main trunk, passes anteriorly deep to the peroneal nerve, and penetrates the lateral surface of M. femorocruralis; in some cases two twigs enter this muscle.
Individual Variation.—In one leg (Fig. 7), the twig to M. caudofemoralis pars caudifemoralis arises more proximally than usual and perforates pars iliofemoralis independently of the branch to M. flexor cruris lateralis. The nerve supplying M. flexor cruris lateralis does not perforate M. caudofemoralis pars iliofemoralis, but passes deep to it in three legs. In half the legs, the paraperoneal branch of the tibial nerve, after extending a short distance in its own sheath, enters the sheath of the peroneal nerve and appears grossly to unite with it; if, however, the sheath is slit open, the paraperoneal branch can be easily pulled apart from the posterior edge of the peroneal nerve; the paraperoneal branch is again enclosed in its own sheath at the distal end of the thigh. In one leg, the cutaneous branch of the peroneal nerve perforates the posteroproximal part of M. gastrocnemius pars externa; in three others, this branch is absent. In one of these last three legs (Fig. 7), the distal cutaneous branch of the tibial nerve is also absent. In three legs (of different specimens), a minute twig from the middle tibial division passes posteriorly deep to M. caudofemoralis pars caudifemoralis toward the tail (Fig. 7); this twig joins the pudendal plexus in one leg; in the other two the twig could not be traced to its termination. Minute twigs come off the peroneal nerve near the middle of the thigh and enter M. extensor iliofibularis in some legs. In a few cases, a minute nonmuscular twig arises from the peroneal nerve near the distal end of the main trunk and passes anteriorly deep to M. vastus lateralis pars postica (Fig. 7).
T. cupido
Individual Variation.—In several legs, the nerve supplying M. flexor cruris lateralis does not perforate M. caudofemoralis pars iliofemoralis, but passes deep to it. The branch to M. flexor cruris medialis arises from the posterior (rather than the middle) tibial division in one instance (Fig. 8). In one leg, a minute twig from the middle tibial division passes posteriorly and joins the pudendal plexus; in another, a similar twig is present but could not be traced to its termination. In some specimens, minute twigs come off the peroneal nerve near the middle of the thigh and enter M. extensor iliofibularis. In one leg, a nonmuscular twig arises from the base of the cutaneous branch of the peroneal nerve and passes anteriorly deep to M. vastus lateralis pars postica. In another leg (Fig. 8), a tiny additional twig arises from the posterior edge of the tibial nerve and subdivides, one branch joining the cutaneous branch of the middle tibial division and the other joining the distal cutaneous branch of the tibial nerve.
P. p. jamesi
Individual Variation.—In both legs of one specimen, the branch to M. flexor cruris medialis arises from the posterior (rather than the middle) tibial division; in three legs, this branch arises as an independent division of the tibial nerve (Fig. 6). (Only in one leg does this branch arise as inT. pallidicinctus.) The branch to M. flexor cruris medialis perforates the lateral part of M. flexor ischiofemoralis in one instance. In all legs except one (nerve possibly destroyed), a second twig to M. flexor ischiofemoralis arises from the branch to M. flexor cruris medialis (Fig. 6). In one leg (Fig. 9), an additional branch, arising as an independent division of the sciatic nerve, enters M. extensor iliofibularis distal to the point of entrance of the dorsal peroneal division; this extra branch arises posterior (adjacent) to the dorsal peroneal division, but it could not be determined with certainty whether it arises from the peroneal or tibial component. A minute twig from the branch to M. flexor cruris medialis passes posteriorly and joins the pudendal plexus in one leg (Fig. 6); in another, a similar twig is present but could not be traced to its termination. In nearly all the legs, minute twigs come off the peroneal nerve near the middle of the thigh and enter M. extensor iliofibularis (Fig. 6). In both legs of one specimen, the paraperoneal branch enters the peroneal sheath (although separable from the peroneal nerve). The distal branch to M. femorocruralis gives off a long twig to M. gastrocnemius pars media in one instance (Fig. 6).
Peroneal Nerve,Fig. 10
T. pallidicinctus
Description.—The branch that is given off in the thigh has been discussed above. The peroneal nerve passes, with the paraperoneal branch of the tibial nerve, through the guide loop for M. extensor iliofibularis. The peroneal nerve diverges from the paraperoneal branch and passes along the anterior (proximal) edge of the tendon of M. extensor iliofibularis medial to the common tendon of the lateral heads of Mm. flexor perforatus digiti IVand flexor perforatus digiti II and lateral to the common tendon of the anterolateral heads of Mm. flexor perforatus digiti IV, flexor perforatus digiti II, and flexor perforatus digiti III.
The peroneal nerve soon gives off a spray of branches that supplies the following: femoral head of M. tibialis anticus, tibial head of M. tibialis anticus (branch passes deep to femoral head), M. extensor digitorum longus (branch passes deep to tibial head of M. tibialis anticus), and M. peroneus longus. A part of the nerve may or may not pass through a notch in the proximal end of the lateral head of M. flexor digitorum longus. The nerve then extends distally along the anterolateral edge of the latter muscle and subdivides into two long branches. Gadow (1891) termed these branches the superficial peroneal and the deep peroneal; his terminology will be used here.
The superficial peroneal branch, after giving off, near its proximal end, one or two twigs into M. peroneus brevis, passes lateral to the retinaculum for the tendon of M. tibialis anticus, then across the intratarsal joint lateral to the latter, then lateral to the insertion of M. tibialis anticus, where the branch subdivides. One of the two resulting branches gives one or two twigs into M. extensor brevis digiti IV, then terminates nonmuscularly in the digits. The other branch passes between the main and accessory insertions of M. tibialis anticus and joins the branch of the deep peroneal which supplies M. abductor digiti II. (See next paragraph.)
The deep peroneal branch passes through the retinaculum for the tendon of M. tibialis anticus, lying lateral, then deep, then medial to the latter; it crosses the intratarsal joint medial to the latter. Immediately above the insertion of M. tibialis anticus, the deep peroneal branch divides, one branch passing on each side of the main insertion. The branch passing lateral to the main insertion passes between the latter and the accessory insertion (medial to the latter) and is joined by a branch of the superficial peroneal nerve. This fused branch extends distally between Mm. extensor hallucis longus and extensor brevis digiti IV and medial to M. extensor brevis digiti III, giving twigs into the latter and into M. abductor digiti II before terminating nonmuscularly in the digits. The branch of the deep peroneal nerve that passes medial to the main insertion of M. tibialis anticus gives one or two twigs into the proximal head of M. extensor hallucis longus, then terminates nonmuscularly in the digits.
Individual Variation.—In four legs, the branch of the superficial peroneal nerve that usually joins the lateral branch of the deep peroneal nerve is lacking (Fig. 10B). In these legs it can be seen that Mm. extensor brevis digiti III and abductor digiti II are supplied by the deep peroneal nerve.
T. cupido
Individual Variation.—In two legs, the same branch that gives twigs into the proximal head of M. extensor hallucis longus also sends a twig into the distal head of this muscle (Fig. 10C).
P. p. jamesi
Individual Variation.—None of significance.
Tibial Nerve,Fig. 11
T. pallidicinctus
Description.—The branches given off in the thigh have been discussed in the account of the sciatic nerve. At the distal end of the thigh the peroneal nerve and the paraperoneal branch of the tibial nerve diverge from the remainder of the tibial nerve and pass through the tendinous guide loop for M. extensor iliofibularis whereas the remainder of the tibial nerve does not. This main part of the tibial nerve immediately divides into three main divisions—lateral, posterior, and medial.
The lateral division passes between Mm. flexor perforatus digiti IV and gastrocnemius pars externa and subdivides into two branches, one of which penetrates the medial surface of M. gastrocnemius pars externa. The other branch passes deep to the latter and sends twigs into the posterior head of M. flexor perforans et perforatus digiti II, then passes deep to the latter and enters M. flexor perforans et perforatus digiti III.
The posterior division sends a branch into the medial head of M. flexor perforatus digiti IV, then passes between the latter and the medial head of M. flexor perforatus digiti III, and extends distally giving off twigs to each of the three heads of M. flexor perforatus digiti IV, to each of the two heads of M. flexor perforatus digiti III, and to each of the three heads of M. flexor perforatus digiti II. The number and arrangement of these twigs is variable.
The medial division passes medial to the medial head of M. flexor perforatus digiti III, sends a twig to the lateral surface of M. gastrocnemius pars media, then passes into the shank musculature between Mm. plantaris and flexor hallucis longus, and sends a branch along the medial edge of M. flexor hallucis longus that gives several twigs into this muscle before terminating nonmuscularly. A small branch extends to M. popliteus, another to M. plantaris, and another to the posterior head of M. flexor digitorum longus. A nonmuscular branch passes between the medial and posterior heads of M. flexor digitorum longus and extends distally deep to this muscle. A long branch gives off near its proximal end a variable number of twigs that pass deep to M. plantaris and enter M. gastrocnemius pars interna; the branch then extends distally along the lateral edge of M. plantaris and terminates nonmuscularly.
The paraperoneal branch diverges from the peroneal nerve, passing medial and then distal to the insertion of M. extensor iliofibularis, whereas the peroneal nerve passes proximal and then lateral to this insertion. The paraperoneal branch passes deep to the lateral heads of Mm. flexor perforatus digiti IV and flexor perforatus digiti II and superficial to the tendon of the anterolateral head of M. flexor perforatus digiti IV and then passes distally along the anterolateral borders of the latter and the lateral head of M. flexor perforatus digiti III and the posterolateral border of M. flexor digitorum longus. This branch is thus separated from the peroneal nerve by M. flexor digitorum longus and by the fibula; the branch passes along the lateral surface of the tibial cartilage, continues lateral to the hypotarsus, then turns medially before extending distally between Mm. abductor digiti IV and flexor hallucis brevis, sending twigs into each of these muscles and a long twig into M. lumbricalis before terminating nonmuscularly.
Individual Variation.—In T.p. 3L,R (Fig. 11B), an extra branch arises from the tibial nerve as a separate (fourth) division; it enters the medial head of M. flexor perforatus digiti IV and also gives off a twig that anastomoses with the posterior division (left leg) or with the first branch of the posterior division (right leg). In T.p. 3R (Fig. 11B), a large extra branch arises from the proximal part of the medial division and passes medial and then deep to the medial head of M. flexor perforatus digiti III, perforates the tendinous part of the medial head of M. flexor perforatus digiti II, and joins the posterior division (lateral to the medial head of M. flexor perforatus digiti III). A similar branch is found in T.p. 3L except that it arises from the proximal part of the posterior (rather than the medial) division. In T.p. 3R (Fig. 11B), the branch to M. gastrocnemius pars externa arises so far proximally that it appears as a separate (fifth) division of the tibial nerve. In two legs, the branch of the medial division that supplies M. gastrocnemius pars media sends a twig into the distal end of M. femorocruralis (Fig. 11A).
T. cupido
Individual Variation.—In one leg, an extra branch of the medial division arises immediately distal to the branch to M. gastrocnemius pars media and enters the proximal end of the medial head of M. flexor perforatus digiti III. In one instance, the branch to M. gastrocnemius pars interna passes through a gap in the origin of M. plantaris rather than distal to the origin of the latter.
P. p. jamesi
Individual Variation.—The branch to M. gastrocnemius pars interna gives a minute twig to the deep surface of the free belly of M. plantaris in one leg.