Mode of Growth and Functional Morphology of Autozooids in Some Recent and Paleozoic Tubular Bryozoa

Abstract

Membranous structures reflecting functional organs are recognizable in a relatively few tubular Bryozoa of Paleozoic age belonging largely to the order Trepostomata. Some skeletal structures also seem to reflect functional organs in a generalized way. Thin sections, including both hard and soft parts, of several genera of Recent tubular Bryozoa of the order Cyclostomata provide a first approximation to the shape, size, and position of cuticular or membranous structures in autozooids that might be preserved under exceptional conditions in fossils. Potentially preservable cuticular or membranous structures include: (1) outward opening funnel-shaped terminal-vestibular membranes and sphincter muscle regions; (2) flask- or sac-shaped membranous sacs; and (3) the spherical-to-formless sex organs and brown bodies. Most of the diaphragms common to trepostome autozooecia presumably formed floors for living chambers of successive functioning bodies in the degeneration-regeneration cycle. The position of some skeletal intrazooecial structures within living chambers must have been lateral to functioning organs. Mural spines that have a definite distributional pattern might represent calcified attachment points for ligaments or muscles. Skeletal cystiphragms, hemiphragms, ring septa, and autozooecial wall thickenings all seem to be lateral features which provided significant modifications to the shape and size of the autozooidal living chamber. These and other skeletal structures appear to have been developed by zooids growing with colony-wide cyclic coordination so that skeletal structures commonly display a constant relative spacing or size correlation in the growth sequence of a colony. Hemiphragms, cystoidal diaphragms, ring septa, and skeletal cystiphragms and funnel-cystiphragms in some species are perhaps more comparable in cycle with basal diaphragms of autozooecia, suggesting that their distribution might have been controlled largely by degeneration-regeneration cycles. Closely tabulated mesopores seem to provide an expression of the most frequent colony-wide cycles in many species and can be correlated one-to-one with some mural spines and skeletal cystiphragms. Perhaps these most closely spaced structures reflect an increase in length of soft parts during a single functional stage of the degeneration-regeneration cycle. Some monticuliporid and diaphragmed trepostomes contain a second type of cystiphragm that forms small flask-shaped chambers filled with brown deposits that suggest a concentration of organic material during the life of the colony. These chambers do not preclude retractable lophophores but almost certainly the inflexible necks restrict significantly the room for passage of membranous structures. Because of this restriction and the scattered or thinly cyclic distribution of flask-shaped chambers known from only a few species, a primary food-gathering function does not seem feasible for them. Possibly, these restricted chambers had a reproductive function, conceivably comparable to the male zooids with reduced numbers of tentacles reported in a few species of cheilostome Bryozoa. Regardless of function, if the flask-shaped chambers and their inferred organs were zooids, they represent <i>intrazooecial</i> polymorphism, contrasting morphologically with the alternating and consistently present living chambers that presumably contained food-gathering organs. The shape, size, and position of food-gathering organs seem more likely then to be reflected by intrazooecial structures that are repeated regularly in autozooecia, such as basal diaphragms, cystiphragms, hemisepta, ring septa, and annular thickenings of zooecial walls.