Biologists strive to understand the evolutionary history and relationships of members of the animal kingdom, and all of life, for that matter. The study of phylogeny aims to determine the evolutionary relationships between phyla. Currently, most biologists divide the animal kingdom into 35 to 40 phyla. Scientists develop phylogenetic trees, which serve as hypotheses about which species have evolved from which ancestors

Recall that until recently, only morphological characteristics and the fossil record were used to determine phylogenetic relationships among animals. Scientific understanding of the distinctions and hierarchies between anatomical characteristics provided much of this knowledge. Used alone, however, this information can be misleading. Morphological characteristics may evolve multiple times, and independently, through evolutionary history. Analogous characteristics may appear similar between animals, but their underlying evolution may be very different. With the advancement of molecular technologies, modern phylogenetics is now informed by genetic and molecular analyses, in addition to traditional morphological and fossil data. With a growing understanding of genetics, the animal evolutionary tree has changed substantially and continues to change as new DNA and RNA analyses are performed on additional animal species.

Constructing an Animal Phylogenetic Tree

The current understanding of evolutionary relationships between animal, or Metazoa, phyla begins with the distinction between “true” animals with true differentiated tissues, called Eumetazoa, and animal phyla that do not have true differentiated tissues (such as the sponges), called Parazoa. Both Parazoa and Eumetazoa evolved from a common ancestral organism that resembles the modern-day protists called choanoflagellates. These protist cells strongly resemble the sponge choanocyte cells today (Figure 1).

Eumetazoa are subdivided into radially symmetrical animals and bilaterally symmetrical animals, and are thus classified into clade Bilateria or Radiata, respectively. As mentioned earlier, the cnidarians and ctenophores are animal phyla with true radial symmetry. All other Eumetazoa are members of the Bilateria clade. The bilaterally symmetrical animals are further divided into deuterostomes (including chordates and echinoderms) and two distinct clades of protostomes (including ecdysozoans and lophotrochozoans) (Figure 2ab). Ecdysozoa includes nematodes and arthropods; they are so named for a commonly found characteristic among the group: exoskeletal molting (termed ecdysis). Lophotrochozoa is named for two structural features, each common to certain phyla within the clade. Some lophotrochozoan phyla are characterized by a larval stage called trochophore larvae, and other phyla are characterized by the presence of a feeding structure called a lophophore.

Modern Advances in Phylogenetic Understanding Come from Molecular Analyses

The phylogenetic groupings are continually being debated and refined by evolutionary biologists. Each year, new evidence emerges that further alters the relationships described by a phylogenetic tree diagram.

Nucleic acid and protein analyses have greatly informed the modern phylogenetic animal tree. These data come from a variety of molecular sources, such as mitochondrial DNA, nuclear DNA, ribosomal RNA (rRNA), and certain cellular proteins. Many evolutionary relationships in the modern tree have only recently been determined due to molecular evidence. For example, a previously classified group of animals called lophophorates, which included brachiopods and bryozoans, were long-thought to be primitive deuterostomes. Extensive molecular analysis using rRNA data found these animals to be protostomes, more closely related to annelids and mollusks. This discovery allowed for the distinction of the protostome clade, the lophotrochozoans. Molecular data have also shed light on some differences within the lophotrochozoan group, and some scientists believe that the phyla Platyhelminthes and Rotifera within this group should actually belong to their own group of protostomes termed Platyzoa.

Molecular research similar to the discoveries that brought about the distinction of the lophotrochozoan clade has also revealed a dramatic rearrangement of the relationships between mollusks, annelids, arthropods, and nematodes, and a new ecdysozoan clade was formed. Due to morphological similarities in their segmented body types, annelids and arthropods were once thought to be closely related. However, molecular evidence has revealed that arthropods are actually more closely related to nematodes, now comprising the ecdysozoan clade, and annelids are more closely related to mollusks, brachiopods, and other phyla in the lophotrochozoan clade. These two clades now make up the protostomes.

Another change to former phylogenetic groupings because of molecular analyses includes the emergence of an entirely new phylum of worm called Acoelomorpha. These acoel flatworms were long thought to belong to the phylum Platyhelminthes because of their similar “flatworm” morphology. However, molecular analyses revealed this to be a false relationship and originally suggested that acoels represented living species of some of the earliest divergent bilaterians. More recent research into the acoelomorphs has called this hypothesis into question and suggested a closer relationship with deuterostomes. The placement of this new phylum remains disputed, but scientists agree that with sufficient molecular data, their true phylogeny will be determined.

Section Summary

Scientists are interested in the evolutionary history of animals and the evolutionary relationships among them. There are three main sources of data that scientists use to create phylogenetic evolutionary tree diagrams that illustrate such relationships: morphological information (which includes developmental morphologies), fossil record data, and, most recently, molecular data. The details of the modern phylogenetic tree change frequently as new data are gathered, and molecular data has recently contributed to many substantial modifications of the understanding of relationships between animal phyla.

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Glossary

Ecdysozoa
clade of protostomes that exhibit exoskeletal molting (ecdysis)

Eumetazoa
group of animals with true differentiated tissues

Lophotrochozoa
clade of protostomes that exhibit a trochophore larvae stage or a lophophore feeding structure

Metazoa
group containing all animals

Parazoa
group of animals without true differentiated tissues