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146 F. H. Edgeworth<br />

however, had stated (1892) that <strong>in</strong> an embryo Amphiuma (size not stated)<br />

<strong>the</strong>re were two pits beh<strong>in</strong>d <strong>the</strong> fourth cartilag<strong>in</strong>ous gill-arch, i.e. five br<strong>on</strong>chial<br />

segments. Marcus (1908) stated that <strong>in</strong> larvae <strong>of</strong> Hypogeophis seven gillclefts<br />

are developed videe <strong>in</strong>fra, p. 149), i.e. five br<strong>on</strong>chial segments.<br />

In Menopoma <strong>and</strong> Ellipsoglossa seven gill-clefts are developed, <strong>the</strong> last<br />

two beh<strong>in</strong>d <strong>the</strong> fourth br<strong>on</strong>chial bar <strong>and</strong> its <strong>muscles</strong>. No muscle-plate or<br />

cartilag<strong>in</strong>ous bar is developed <strong>in</strong> this fifth br<strong>on</strong>chial segment.<br />

The sum <strong>of</strong> this evidence suggests that five br<strong>on</strong>chial bars with related<br />

<strong>muscles</strong> existed <strong>in</strong> Amphibian ancestors, but that <strong>the</strong> fifth disappeared l<strong>on</strong>g<br />

ago. Druner videe supra, pp. 129, 130) came to <strong>the</strong> c<strong>on</strong>clusi<strong>on</strong> that Urodeles<br />

orig<strong>in</strong>ally possessed a greater number <strong>of</strong> br<strong>on</strong>chial arches than four. The<br />

evidence merits discussi<strong>on</strong>. As regards <strong>the</strong> nerves found by him, it is possible<br />

that <strong>the</strong>y have relati<strong>on</strong> solely to <strong>the</strong> sixth <strong>and</strong> seventh gill-clefts. As regards<br />

<strong>the</strong> <strong>muscles</strong>, <strong>the</strong> phenomena <strong>of</strong> <strong>development</strong> do not bear out <strong>the</strong> <strong>the</strong>ory that<br />

<strong>the</strong> stump <strong>of</strong> <strong>the</strong> sixth gill-cleft separates a Transversus ventralis iv from a<br />

Transversus ventralis v. Thus <strong>in</strong> Menopoma <strong>the</strong>re is <strong>on</strong>ly <strong>on</strong>e primordium<br />

developed-that <strong>of</strong> Transversus iv. This, at first, lies solely <strong>in</strong> <strong>the</strong> fourth<br />

br<strong>on</strong>chial segment, <strong>the</strong>n under <strong>the</strong> stump <strong>of</strong> <strong>the</strong> sixth gill-cleft, <strong>the</strong>n posterior<br />

to it. The variati<strong>on</strong> is due to <strong>the</strong> fact that, <strong>on</strong> <strong>the</strong> partial atrophy <strong>of</strong> <strong>the</strong><br />

sixth gill-cleft, <strong>the</strong> Transversus iv, <strong>the</strong> h<strong>in</strong>d end <strong>of</strong> Subarcualis rectus iv, <strong>and</strong><br />

<strong>the</strong> fourth br<strong>on</strong>chial bar, migrate <strong>in</strong>to <strong>the</strong> fifth br<strong>on</strong>chial segment; <strong>and</strong>, <strong>on</strong><br />

<strong>the</strong> atrophy <strong>of</strong> <strong>the</strong> seventh gill-cleft, still fur<strong>the</strong>r back. There is a similar<br />

progressive variati<strong>on</strong> <strong>of</strong> <strong>the</strong> Transversus iv <strong>in</strong> relati<strong>on</strong> to <strong>the</strong> sixth gill-cleft<br />

<strong>in</strong> Ellipsoglossa, <strong>and</strong> <strong>of</strong> Transversus iii <strong>in</strong> relati<strong>on</strong> to <strong>the</strong> fifth gill-cleft <strong>in</strong><br />

Necturus. The evidence, as regards muscle elements, is thus limited to <strong>the</strong><br />

discovery <strong>of</strong> a Levator arcus v <strong>on</strong> <strong>on</strong>e side <strong>in</strong> <strong>on</strong>e larva <strong>of</strong> Trit<strong>on</strong>. I have not<br />

seen this, however, <strong>in</strong> <strong>the</strong> many larvae <strong>of</strong> Menopoma, Ellipsoglossa, <strong>and</strong><br />

Trit<strong>on</strong> crist. I have exam<strong>in</strong>ed.<br />

It is probable, <strong>the</strong>refore, that <strong>the</strong> case described by Druner is an <strong>in</strong>stance<br />

<strong>of</strong> fluctuati<strong>on</strong>, possibly <strong>of</strong> atavistic fluctuati<strong>on</strong>, from <strong>the</strong> usual number <strong>of</strong><br />

four br<strong>on</strong>chial bars <strong>and</strong> related <strong>muscles</strong> <strong>in</strong> Urodela.<br />

In support <strong>of</strong> <strong>the</strong>se c<strong>on</strong>clusi<strong>on</strong>s, it may be added that <strong>in</strong> Lysorophusa<br />

member <strong>of</strong> <strong>the</strong> ancestral Urodeles, from <strong>the</strong> Pennsylvanian deposits-<br />

Sollas found <strong>on</strong>ly four br<strong>on</strong>chial bars, <strong>the</strong> first three c<strong>on</strong>sist<strong>in</strong>g <strong>of</strong> ceratobranchial<br />

<strong>and</strong> epibranchial elements, <strong>and</strong> <strong>the</strong> fourth <strong>of</strong> a tra<strong>in</strong> <strong>of</strong> fragments.<br />

The case <strong>of</strong> Necturus, with <strong>on</strong>ly three branchial bars <strong>and</strong> related <strong>muscles</strong>,<br />

probably comes under a different category. All Amphibia, with excepti<strong>on</strong> <strong>of</strong><br />

Megalobatrachus max. (adult), Necturus <strong>and</strong> Proteus, have four br<strong>on</strong>chial bars<br />

<strong>and</strong> related <strong>muscles</strong>. Megalobatrachus max. has two branchial bars <strong>in</strong> <strong>the</strong><br />

adult state (Druner), but <strong>on</strong>ly <strong>the</strong> early stages <strong>of</strong> larval <strong>development</strong> have<br />

been published hi<strong>the</strong>rto-by de Lange. InNecturus <strong>the</strong> <strong>development</strong>, described<br />

above, permits <strong>of</strong> comparis<strong>on</strong> with Menoporna.<br />

In Menopoma seven gill-clefts are formed, all <strong>of</strong> which reach <strong>the</strong> ectoderm.

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