to four and these coalesce by their dorsal walls along the line of the nema, and the sicula becomes embedded in the base of the polypary. In the family of the Diplograptidae the branches are reduced to two; these also coalesce similarly by their dorsal walls, and the polypary thus becomes biserial (diprionidian), and the line of the nema is taken by a long axial tube-like structure, the nemacaulus or virgular tube. Finally, in the latest family, the Monograptidae, the branches are theoretically reduced to one, the polypary is uniserial throughout, and all the thecae are directed outwards and upwards.
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1, Diptograptus, young sicula. |
20, Climacograptus Scharenbergi. |
The thecae in the earliest family—Dichograptidae—are so similar in form to the sicula itself that the polypary has been compared to a colony of siculae; there is the greatest variation in shape in those of the latest family—Monograptidae—in some species of which the terminal portion of each theca becomes isolated (Rastrites) and in some coiled into a rounded lobe. The thecae in several of the families are occasionally provided with spines or lateral processes: the spines are especially conspicuous at the base in some biserial forms: in the Lasiograptidae the lateral processes originate a marginal meshwork surrounding the polypary.
Histologically, the perisarc or test in the Graptoloidea appears to be composed of three layers, a middle layer of variable structure, and an overlying and an underlying layer of remarkable tenuity. The central layer is usually thick and marked by lines of growth; but in Glossograptus and Lasiograptus it is thinned down to a fine membrane stretched upon a skeleton framework of lists and fibres, and in Retiolites this membrane is reduced to a delicate network. The groups typified by these three genera are sometimes referred to, collectively, as the Retioloidea, and the structure as retioloid.
It is the general practice of palaeontologists to regard each graptolite polypary (rhabdosome) developed from a single sicula as an individual of the highest order. Certain American forms, however, which are preserved as stellate groups, have been interpreted as complex umbrella-shaped colonial stocks, individuals of a still higher order (synrhabdosomes), composed of a number of biserial polyparies (each having a sicula at its outer extremity) attached by their nemacauli to a common centre of origin, which is provided with two disks, a swimming bladder and a ring of capsules.
In the Dendroidea, as a rule, the polypary is non-symmetrical in shape and tree-like or shrub-like in habit, with numerous branches irregularly disposed, and with a distinct stem-like or short basal portion ending below in root-like fibres or in a membranous disk or sheet of attachment. An exception, however, is constituted by the comprehensive genus Dictyonema, which embraces species composed of a large number of divergent and sub-parallel branches, united by transverse dissepiments into a symmetrical cone-like or funnel-shaped polypary, and includes some forms (Dictyograptus) which originate from a nema-bearing sicula and have been claimed as belonging to the Graptoloidea.
Of the early development of the polypary in the Dendroidea little is known, but the more mature stages have been fully worked out. In Dictyonema the branches show thecae of two kinds: (1) the ordinary tubular thecae answering to those of the Graptoloidea and occupied by the nourishing zooids; and (2) the so-called bithecae, birdnest-like cups (regarded by their discoverers as gonothecae) opening alternately right and left of the ordinary thecae. Internally, there existed a third set of thecae, held to have been inhabited by the budding individuals. In the genus Dendrograptus the gonothecae open within the walls of the ordinary thecae, and the branches present an outward resemblance to those of the uniserial Graptoloidea. But in striking contrast to what obtains among the Graptoloidea in general, the budding orifices in the Dendroidea become closed, and all the various cells shut off from each other.
The classification of the Dendroidea is as yet unsatisfactory: the families most conspicuous are those typified by the genera Dendrograptus, Dictyonema, Inocaulis and Thamnograptus.
As regards the modes of reproduction among the Graptolites little is known. In the Dendroidea, as already pointed out, the bithecae were possibly gonothecae, but they have been interpreted by some as nematophores. In the Graptoloidea certain lateral and vesicular appendages of the polypary in the Lasiograptidae have been looked upon as connected with the reproductive system; and in the umbrella-shaped synrhabdosomes already referred to, the common centre is surrounded by a ring of what have been regarded as ovarian capsules. The theory of the gonangial nature of the vesicular bodies in the Graptoloidea is, however, disputed by some authorities, and it has been suggested that the zooid of the sicula itself is not the
