McGill University, Department of Earth & Planetary Sciences
Invertebrate Paleontology EPSC 334

Laboratory #4: CNIDARIANS & BRYOZOANS

Although we spontaneously associate reef-building with corals, the term reef is used by geologists for any structure constructed by organic activity and that rose above the seafloor.  Reefs are recognized in the fossil record by the greater compaction of sedimentary layers on either side of it. Recent corals belong to an order of cnidarians which did not arise until the Mesozoic era, about 230 million years ago. They were preceded by other organisms that seem to have had similar environmental preferences and erected reefs around which diverse marine communities thrived.

During the Precambrian, bacterial communities (probably dominated by cyanobacteria) trapped sediments and precipitated minerals from seawater which accumulated in reef-like stromatolites. Extensive algal mats became gradually restricted in the last part of the Proterozoic eon, possibly because herbivores had evolved and started grazing on them.

Archaeocyathids, the first major reef-building metazoans, were widespread but declined before the end of the Cambrian. During the rest of the Paleozoic, stromatoporoids (poriferans) were the main reef builders, with lesser contributions from corals, calcareous algae and bryozoans. This exercise introduces you to the general features of two metazoan phyla among reef builders, CNIDARIA and BRYOZOA.

Like the poriferans, cnidarians are diploblastic i.e. the embryo has only two differentiated cellular layers from which 9-2 cell types differentiate and develop into distinct tissues that fulfill different functions. The cnidarian polyp has a jar-shaped body, lined inside with digestive tissue, opened at the top in a mouth surrounded by a ring of tentacles. In most classes, the inner digestive surface is increased by radial folding to produce incomplete partitions called mesenteries.

PHYLUM CNIDARIA, CLASS HYDROZOA

Millepora is a hydrozoan known as the “fire coral”. Examine any one of the specimens under a stereomicroscope. Are all the pores in which individual polyps were living of the same size or do they show more than one size (1a)? What does this indicate about the polyps (1b)?Are there septa projecting into each pore, as in scleractinian corallites (1c)?

Millepora It is found in a range of growth forms that are adapted to different degrees of ambient water turbulence.  How would you label the growth form of each specimen (2a)? Which one of these growth forms is most likely to be an adaptation to relatively low water turbulence (2b)?

PHYLUM CNIDARIA, CLASS SCYZOPHOA

CONULARIDS are an enigmatic group (stratigraphic range), now extinct, which is suspected of belonging to the cnidarians. Examine and sketch the specimen provided (3a). Was it anchored to the bottom, swimming or floating (3b)?
PHYLUM CNIDARIA - Class ANTHOZOA - Order SCLERACTINIA

The stony CORALS (as opposed to the “soft corals” of the subclass OCTOCORALLIA) are members of the subclass ZOANTHARIA within the class ANTHOZOA. As a coral grows, it secretes a CaCO3 tube or corallite whose diameter expands until the polyp reaches adult size and then becomes cylindrical. As the polyp rises within the corallite, the successive positions of its base are marked by tabulae (large plates across the tube) or dissepiments (small, curved plates near the rim of the calyx). In many corals, radial plates called septa are secreted in (and support) the mesenteries on the floor of the calyx.

Recent corals, from the order SCLERACTINIA, appear in the fossil record during the Triassic (early Mesozoic), i.e. after the spectacular Permian extinction that marked the end of the Paleozoic era. Scleractinians are sometimes referred to as the "hexacorals" because the pattern of septa in each corallite displays 6-fold symmetry.  After the first 6 septa appear additional septa are added in multiples of 6, giving a symmetrical structure. The septa form ridges on the outside of the corallites (unlike those of the rugose corals, which form grooves) and the wall of the calyx (epitheca) is formed by the fusion of the outer edges of the septa.

Examine the scleractinian corals set aside for this question. Name one solitary genus (4a). Colony form, which describes the relations of the corallites, is described separately from colony habit. Find a colonial genus whose form can be describe as plocoid (4b). Find a colonial genus whose habit can be described as branching (syn. ramose) (4c), platy (4d), massive (4e).
 
Sketch the shape of a single corallite from a colonial coral, indicating its genus and specimen number (5a). Focus on a single quadrant of the corallite. Label on your sketch the different orders of visible septa (i.e. septa of different sizes that are repeated either 6, 12 or 24 times) as “1st”, “2nd”, “3rd”,  etc.  Are the larger septa more or less numerous than the shorter septa (5b)?

Note that the colonies show different degrees of corallite integration. Choose any one of the colonial corals and describe in your own words the relations among corallites (6). For example, does each corallite has its own wall, are these walls separated by empty space or by skeletal structures that are neither walls nor septa; if there are no walls, do the septa of adjacent corallite touch or merge?

The origin of scleractinian corals is unclear. They secrete their hard parts in a pattern that is clearly different from their Paleozoic predecessors. Their soft parts suggest that they are more closely related to sea anemones (order ACTINARIA within the subclass ZOANTHARIA) than to the extinct Paleozoic orders of corals.

PHYLUM CNIDARIA - Class ANTHOZOA - Order TABULATA

These are a diverse group of middle Paleozoic corals in which the septa are missing or very short, essentially rows of spines, commonly in multiple of 6, extending inwards from the walls of the corallites.  Tabulae are prominent, crossing the corallites perpendicular to their length.

Examine Lichenaria, one of the first corals in the fossil record. Note that It is of very simple structure, with small corallites.

The family Tetradiidae is a primitive family in which the corallites are even smaller, polygonal or square, and without pores or true septa. In Tetradium the corallites are square and have 4 partitions that resemble septa.  When the partitions grow to meet towards the middle, the corallite is divided into 4 new corallites. Examine it under a stereomicroscope and use the polished specimen to sketch four adjacent corallites containing partial partitions (7).

The family Favositidae includes the so-called "honey-comb" corals. Their colonies are composed of contiguous corallites in hexagonal patterns, with spiny septa and pores in the corallite walls joining the polyps. Their resemblance to chaetetids (seen last week) who are now known to be sclerosponges generated vigorous debate about their true nature. What evidence of their affinity did Paul Copper (now at Laurentian University) report from a Canadian specimen in 1985 (Nature, vol. 316, p. 142)?

Examine an example of the common Silurian and Devonian genus Favosites.  Make a sketch of Favosites Draw a longitudinal section (8a) and a transverse section (8b), labelling the tabulae, septa and corallites.

The family Halysitidae is familiarly known as the "chain" corals.  The corallites are attached to each other only along two edges and form loops that are chain-like in transverse section. Halysites and Catenipora are representative. Manipora is a late Ordovician genus with more than one row of corallites in each rank.

In the family Auloporidae, Aulopora is a creeping, encrusting genus composed of trumpet-shaped corallites. Look for specimens where it covers other organisms.

Members of the family Syringoporidae are often referred to as the “organ pipe” corals. Examine the specimens of Syringopora, a very common genus (Silurian to Carboniferous) in which cylindrical corallites are filled with nested funnel-shaped tabulae.” Is there evidence of connections (phaceloid or dendroid) among the soft tissues of corallites? (9a). Describe the mode of preservation of the specimens: are the corallites filled by the same material as the space between them (9b).

Family Heliolitidae - The subcircular corallites are surrounded by cystose or tubulate tissue secreted by the connective tissue between the polyps. Examine the commonest genus Heliolites. Why are heliolitids sometimes classified apart from other tabulates (10)?

PHYLUM CNIDARIA - Class ANTHOZOA - Order RUGOSA

This order consists of Paleozoic solitary and colonial corals with septa arranged in a bilateral pattern. Solitary rugose corals are often called "horn corals".

The sequence in which the septa appear can be determined by cutting a corallite transversely at the tip of the horn (secreted at the earliest stage of its life).  After the 6 first septa, called primary septa, major septa are added in 4 sectors only. For this reason, rugose corals are sometimes called "tetracorals". 

The horn-like shape of the skeleton indicates that, as the coral grew, it secreted a calyx of progressively larger size and moved itself up the tube.  Changes in growth rate produce the grooves (called rugae) after which rugose corals are named. At each stage, the polyp was supported by a flat platform called tabula and/or the smaller curved peripheral plates called dissepiments.  These structures are easily seen in longitudinal sections.

Examine specimens of the solitary rugosan corals Streptelasma (Ordovician) or Caninia (Carboniferous).  Note that the septa are of several “orders” (i.e. different lengths) as in modern scleractinian corals. Some reach the center of the horn and others are confined to the margins.  In Streptelasma the septa reach the center and form an axial column (called columella) that sticks up from the base of the calyx. 

Draw a cross section of either Streptelasma or Caninia (only one!) and show the relationship of the tabulae, dissepiments and septa (11). You need to use both transverse and longitudinal sections to show every element in a block diagram.

Examine the horn corals and Calceola. Note that they are not straight cones nor perfectly cylindrical. What does this indicate about the life position (12)?

[See item posted on WebCT for this question.] What information about the Earth’s rotation was confirmed by Scrutton and Hipkin (1973) in their study of the growth bands of Devonian rugose corals (13a)?  In what climatic conditions must these rugose corals have grown to provide sufficient evidence to support  their conclusion (13b) ?

Not all rugosans were solitary. Colonial rugosans appeared in the middle Ordovician and  became abundant after the tabulates declined in the late Paleozoic. Some of the colonies consist of separate cylindrical corallites.  In others genera the corallites are in contact forming a honey-comb pattern. Find an example of a colonial rugose coral in the collection and describe its colony as phaceloid, cerioid, astreoid, aphroid, coenostoid or meandroid (14)?

PHYLUM BRYOZOA

This phylum is the only skeletonized invertebrate phylum to appear after the Cambrian period.  Bryozoans (also called “moss animals”) are small colonial metazoans that built a common skeleton of CaCO3 composed of minute tubes or boxes.  The colonies show the same diversity of shapes as those of corals and stromatoporoids. Bryozoans lack the septa of rugose corals. They can often be distinguished from tabulate corals by the smaller size of their tubes but there is a zone of overlap at a diameter of about 0.5 mm.

The phylum BRYOZOA is divided into several classes and orders. Only 3 orders have an extensive fossil record and only a few of the most common forms are presented in this exercise. 

PHYLUM BRYOZOA - CLASS GYMNOLAEMATA - Order CHEILOSTOMATA

Most of the Recent genera belong to the order CHEILOSTOMATE which arose during the Mesozoic.They secrete a skeleton composed of small boxes.  Each box has an aperture for the extrusion of the tentacles. In some cases the aperture is a double hole, and the second opening leads into a sack which is squeezed by a contractile muscle to expel water and push the feeding tentacles out.  The front surface of the boxes is commonly lacy, pierced by many pores.

Examine the form of the colony in the following Recent bryozoans Bugula, Retepora, Cellepora, Membranipora, Escara.  Find examples of the following forms: encrusting (15a), ramose (15b), tubular (15c).

Examine some of the common bryozoans of the early Paleozoic era, some of which  were important reef builders.  Examine the chart provided: did bryozoans build reefs in associations with other groups, or were they prominent during periods of decline of stromatoporoids and corals (16)?

CLASS GYMNOLAEMATA – ORDER CTENOSTOMATA  [Answer posted on WebCT.]  Why do these bryozoans leave trace fossils, not body fossils in sedimentary rocks (17)?

CLASS STENOLAEMATA - ORDER TREPOSTOMATA

Trepostomes have partitions within and between their tubular housings that resemble those of certain corals. To what order of coral (TABULATA, RUGOSA, SCLERACTINIA) are they most similar (18)?

Compare the following pairs of organism. Indicate which genus in each pair is a bryozoan, and explain how you can identify their “look alike” as either a tabulate coral, a scleractinian coral or a stromatoporoid (poriferan). If distinctive features are lacking on the specimens, explain what you might look for in either a transverse or a longitudinal section.
- Hallopora and Porites (19a).
- Constellaria and an unnamed specimen (19b).
- Prasopora and Favosites (19c).

CLASS STENOLAEMATA – ORDER FENESTRATA

Fenestrate bryozoans built lacy sheets pierced with large openings, and on which all the zooecia were facing the same way.  Examine the specimens provided.

Archimedes is particularly distinctive because its sheet is coiled into a helical crew. Examine the specimens and find one where the opening of the zoecia are visible. On which side of the sheet  are the zoecia openings: are they facing upward or downward  (20)?