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PHYLUM ARTHROPODA METAZOA I UNIVERSIDAD INDUSTRIAL DE SANTANDER

Introduction Multicellular life arose in the Precambrian Period. While many animal phyla appear to have originated near the end of the Precambrian (e.g., putative annelids), the first diverse assemblages of animals is not known until the Cambrian, the so-called Cambrian Explosion (520 million years) (Conway Morris, 2003). Among the diversity of groundplans for building an animal, the phylum Arthropoda, which has achieved a level of evolutionary success unrivaled in evolutionary history, is clearly more dominant on Earth relative to all others (Grimaldi & Engel, 2005). The arthropodan groundplan is the most commonly encountered form of life, having radiated into more species and into more habitats than any other lineage (Grimaldi & Engel, 2005).

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Fig. 1. Arthropod tagmosis. 1. Myriapoda (Millipede) with two tagmata (head and body); 2. Crustacea (lobster) with two tagmata (cephalothorax and abdomen); Chelicerata (scorpion and spider) with two tagmata (cephalothorax and abdomen); Hexapoda (insect) with three tagmata (head, thorax and abdomen).

Main morphological characters defining Arthropoda (Triplehorn & Johnson, 2005):

The body segmented, the segments usually grouped in two or three rather distinct regions (fig. 1) Paired, segmented appendages (from which the phylum gets its name). Bilateral symmetry. A chitinous exoskeleton, which is periodically shed and renewed as the animal grows (Ecdysis). A tubular alimentary canal, with mouth and anus. An open circulatory system; the only blood vessel ussually being a tubular structure, dorsal to the alimentary canal, with lateral openings in the abdominal region.

Nervous system consisting of an anterior ganglion or brain located above the alimentary canal; a pair of connectives extending from the brain around the alimentary canal and paied ganglionated nerve cords located below the alimentary canal. Striated skeletal muscles. Respiration by means gills, or trachea and spiracles.

Two competing theories are currently relevant to the arthropod sister group: either onychophorans or tardigrades are the closest relatives of arthropods.

Fig. 2. Phylogeny of panarthropod phyla. Left: Onychophora, Tardigrada and Euarthropoda from Nielsen (2001); right: Tardigrada, Onychophora and Euarthropoda from Edgecombe (2009).

Euarthropods are typically classified into four subphyla: Subphylum Chelicerata Most of them are terrestrial and many are aquatic or semiaquatic (Triplehorn & Johnson, 2005). Animals belonging to this subphylum lack antennae and tipically have six pairs of appendages. The first pair are the chelicerae, and the rest are leglike. Endite lobes of the pedipalps (second pair of appendages in the arachnids), or the leglike appendages in the horseshoe crabs,function as jaws. The body usually has two distinct divisions, an anterior region called prosoma (or cephalothorax) and a posterior region called the opisthosoma (or abdomen) (Fig. 3). The prosoma bears the chelicerae and the leglike appendages. The genital ducts open to the outside near the anterior end of the opisthosoma. Most chelicerates have and extra leg segment, the patella, between the femur and the tibia. The legs are generally uniramous; that is there is no exite or exopodite (Triplehorn & Johnson, 2005). Chelicerates has diversified to use all the major feeding strategies: predation, parasitism, herbivory, scavenging and eating decaying organic matter (De la Fuente, 1994).

Fig.3.Chelicerata: groundplan. Left, spider; right, scorpion.

Subphylum Myriapoda Myriapods are terrestrial; they have a single pair of antennae and, in most cases, simple eyes. The body is divided into numerous segments and possess numerous pairs of uniramous articulated legs. The mouthparts lie on the underside of the head, with an "epistome" and labrum forming the upper lip, and a pair of maxillae forming the lower lip; palpus are usually present in first and second maxillae. A pair of mandibles lie inside the mouth. Myriapods breathe through spiracles that connect to a tracheal system similar to insects. There is a long tubular heart that extends through much of the body, but usually few, if any, blood vessels . Centipedes have a single pair of legs on each body segment and they have a pair of venomous claws (called maxillipedes) that are used for both defense and for capturing and paralyzing prey (Triplehorn & Johnson, 2005).

Fig. 4. Myriapods ground plan. A, Millipede; B. Centipede.

Subphylum Crustacea Most of them are marine, but many occur in fresh water, and a few are terrestial. The appendages and body regions vary greatly in this group, but tipically are two pairs of antennae, the functional jaws consist of endite lobes of the gnathal appendages and many of the appendages are biramous (De la Fuente, 1994).

Fig. 5. Crustacea: ground plan of a shrimp.

Subphylum Hexapoda Five groups of Hexapoda are traditionally recognized: the wingless Collembola, Protura, Diplura and thysanura, and the winged Pterygota (Insecta) (Richard and Davies, 1957). Their body plan consists of a head, thorax and abdomen. The hexapoda head has a pair of mandibules, a pair of maxillae, a hypopharynx and a labium. There are three segments on the thorax, each segment supporting a pair of legs a few insects are legless, and some larvae have additional leglike appendages, such as prolegs on the abdominal segments). The first two thoracic segments also bear a pair of wings, but these are often reduced or lost. On the abdomen there are ten or 11 segments, though it may seem like fewer in some species. No locomotor appendages on the abdomen of the adult (except in some primitive hexapods); abdominal appendages, if present, located at the apex of the abdomen and consisting of a pair of cerci, an epiproct, and a pair of paraprocts (Triplehorn & Johnson, 2005).

Fig. 6. Hexapoda: apterygota. A, Thysanura; B, Protura; C, Collembola; D, Diplura.

Class Insecta The major advance made by thier descendants was the evolution of wings, facilitating dispersal and therefore, colonization of new habitats. Their body is divided into head, thorax and abdomen.The insect head consists of six segments: the first and third have no appendages and the second bears a pair of antennae (homologous to the antennules of the Crustacea) (Owain & Davies, 1957); segments four through six support modified appendages used in feeding (a pair of mandibules, a pair of maxillae, a hypopharynx and a labium). The thorax carries three pairs of legs and usually one or two pairs of wings. The abdomen is devoid of ambulatory appendages, and the genital opening is situated near anal extremity of the body. Postembryonic development is rarely direct and metamorphosis is usually undergone .

Fig. 7. Insect morphology: ground plan of a grasshopper.

Purpose of the laboratory Recognized phylum Arthropoda, based on morphological characters and general descriptions. Differentiate phylum Arthropoda from others.

Materials Macroscopic samples of Chelicerata, Myriapoda, Hexapoda y Crustacea. Dissection equipment Stereoscope Microscope

Activity 1. Draw the main cephalic, thoracic and abdominal structures (mouth parts, legs, wings, spiracles, parameres and cerci.) of each arthropod subphylum.

Activity 2. Based on the table exemplified below, using the 31 characters from the list, write 1, if the character is present and 2, if it is absent. On the required cells, specify a number (e.g. number of legs) or a type (e.g. type of larva).

List 1: 1-> Body divided into three tagmas. 1= presence, 2= absense. 2-> Articulated legs. 1= presence, 2= absense. 3-> Bilateral symmetry. 1= presence, 2= absense. 4-> Mouth originated from the blastopore. 1= presence, 2= absense. 5-> Chitinous exoskeleton. 1= presence, 2= absense.

6-> Number of tagmas. 7-> Mesosome. 1= presence, 2= absense. 8-> Mesothorax. 1= presence, 2= absense. 9-> Tentorium. 1= presence, 2= absense. 10-> Maxillae. 1= presence, 2= absense. 11-> Quelicerae. 1= presence, 2= absense. 12-> Mandibules. 1= presence, 2= absense. 13-> Ommatidia. 1= presence, 2= absense. 14-> Oceli. 1= presence, 2= absense. 15-> Compound eyes.1= presence, 2= absense. 16-> Pedunculate eyes. 1= presence, 2= absense. 17-> Antennae. 1= presence, 2= absense. 18-> Palps. 1= presence, 2= absense. 19-> Labrum. 1= presence, 2= absense. 20-> Number of legs. 1= presence, 2= absense. 21-> Patela. 1= presence, 2= absense. 22-> Number of segments in the proleg. 23-> Wings. 1= presence, 2= absense. 24-> Gills. 1= presence, 2= absense. 25-> Telson. 1= presence, 2= absense. 26-> Spiracles. 1= presence, 2= absense. 27-> Larva type. 28-> Nauplius larva. 1= presence, 2= absense. 29-> Number of abdominal segments. 30-> Tritocerebrum. 1= presence, 2= absense. 31-> Deuterocerebrum. 1= presence, 2= absense.

REFERENCES Boudreaux, H.B. 1979. Arthropod phylogeny with special reference to insects. Wiley, New York. Conway Morris, 2003. he Cambrian explosion of metazoans. Pp. 1332. In Mller, G. B., and S. A. Newman (eds.),Origination of Organismal Form: Beyond the Gene in Developmental and Evolutionary Biology. MIT Press; Cambridge, Massachusetts, 332 pp. Grimaldi, D. & Engel, M. 2005. Evolution of the Insects. Cambridge University Press. De la Fuente, J. 1993. Zoologia de Artropodos, Mcgraw-Hill, Interamericana de Espaa, S.A. 950 pp. Edgecombe, G. 2010. Arthropoda phylogeny: An overview from the perspectives of morphology, molecular data and the fossil record. Arthropod Structure & Development 39, 74-87. Mayer, G., 2006. Origin and differentiation of nephridia in the Onychophora provide no support for the Articulata. Zoomorphology 125, 112. Mallatt, J.M., Giribet, G., 2006. Further use of nearly complete 28S and 18S Rrna genes to classify Ecdysozoa: 37 more arthropods and a kinorhynch. Molecular Phylogenetics and Evolution 40, 772794. Nielsen, C. 1995. Animal evolution, interrelationships of the living phyla. Oxford University Press, Oxford. Nielsen, C. 2001. Animal Evolution: Interrelationships of the Living Phyla [2nd Edition]. Oxford University Press; Oxford, UK. Owain, W & Davies, R.G. 1957. A General Textbook of Entomology. Methuen & Co Ltd. Ninth Edition, 727 pp. Triplehorn, C.A. & N.F. Johnson. 2005. Borror and DeLong's Introduction to the Study of Insects. Thomson Brooks/Cole, Seventh Edition, USA, 864 pp.

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