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INSECTS Arthropoda INSECTS 579 [See also Disease, article on Infectious Disease; Im- species of animals on earth. Insecta alone accounts for mune System, articles on Microbial Countermeasures to about two-thirds of the animal species (Hammond, Evade the Immune System and Structure and Function 1992). Researchers have diagnosed the phylum Arthro- of the Vertebrate Immune System; Plagues and Epidem- poda in different ways. The most generally accepted fea- ics; Vaccination; Viruses.] tures are a chitinous exoskeleton and jointed append- ages. Other commonly cited synapomorphies (shared, BIBLIOGRAPHY derived homologous features) include an open circula- Bush, R. M., C. A. Bender, K. Subbarao, N. J. Cox, and W. M. Fitch. tory system, Malpighian tubules used for nitrogenous "Predicting the Evolution of Human Influenza A." Science 286 waste concentration and excretion, the hemocoel body (1999): 1921-1925. Use of molecular techniques to study anti- cavity, the lack of cilia on any body cells, the lack of genie drift. Cox, N. J., and C. A. Bender. "The Molecular Epidemiology of In- nephridia (the excretion organ of segmented worms), fluenza Viruses." Seminars in Virology 6 (1995): 359-370. and separate sexes (as opposed to a near sister group, Cox, N. J., and K. Subbarao. "Global Epidemiology of Influenza: segmented worms, which can be hermaphroditic). Some Past and Present" Annual Review of Medicine 51 (2000): 407- experts have included the lobopods, Peripatus, for ex- 421. ample, as arthropods but most recognize those interest- Dowdle, W. R. "Influenza A Virus Recycling Revisited." Bulletin of ing creatures as a separate phylum, the Onychophora. the World Health Organisation 77 (1999): 820-828. The exact topology of relationships among major groups Glezen, W. P., and R. B. Couch. "Influenza Viruses." In Viral Infec- tions of Humans, edited by A. S. Evans and R. A. Kaslow, 4th of the phylum Arthropoda is a contentious issue. The ed., 473-505. New York, 1997. General overview of influenza. phylogenetic tree in Figure 1 summarizes the relation- Murphy, B. R., and R. G. Webster. "Orthomyxoviruses." In Fields ships among major groups of arthropods. Note that the Virology, edited by B. N. Fields, D. M. Knipe. and P. M. Howley, sister group of insects is myriapods. Other recent mo- 3d ed., 1397-1445. Philadelphia, 1996. General overview of in- lecular considerations list Crustacea as the sister group. fluenza. Support for this arrangement with Crustacea is sparse Nicholson, K. G., R. G. Webster, and A. K. Ray (eds). Textbook of and does not overwhelm the purported synapomorphies Influenza. Oxford, 1998. In-depth chapters covering many areas of influenza biology. of Insecta and Myriapoda when morphological and Reid, A. II., J. K. Taubenberger, and T. G. Fanning. "The 1918 Span- molecular lines of evidence are evaluated together ish Influenza: Integrating History and Biology." Microbes and (Wheeler et al., 2001). Whether myriapods form a mono- Infection 3 (2001): 81-87. phyletic group (a group consisting of a common ances- — ROBIN M. BUSH tor and all of its descendants) is unclear. Myriapods could be merely a paraphyletic group (an evolutionary grade consisting of some, but not all, descendants of a INSECTS common ancestor). Both points of view have propo- Insects are the most diverse group of organisms on earth and seem to have been diverse since at least the Permian period, about 250 million years ago. This qualifies them FIGURE 1. Relationships Among Classes of Arthropoda. as the most successful animals ever to have lived on C. Ritey Nelson. earth. Scientists recognize at least 750,000 species of in- Annelida sects and place them in the order Insecta. They estimate, however, that the total number of living insect species is ten million or more. The number of beetle species Onychophora alone (order Coleoptera) at about 500,000, is roughly twice that of the nearest other major group (green plants). This amazing richness is evidence that evolution has taken varied paths to fill or subdivide niches. Larger (and smaller) organisms seem not to have exploited niches to this extent. Many researchers have posed the Arthropoda question Why are there so many insect species? This question may be rephrased as What key adaptations have allowed insects to be so species rich? Insects as Arthropods. Insects are the most spe- cies-rich class in the phylum Arthropoda. Arthropods Myriapoda include major groups such as the extinct trilobites, cheli- cerates, crustaceans, myriapods, and insects. The phy- lum Arthropoda contains roughly three-quarters of the Insecta 580 INSECTS nente. In any event, the ancestral insect (hexapod) is six legs; tagmatization, having the body divided into thought to have evolved from some member of this myr- three regions; mandibles with two condyles; wings; the iapod group. The controversy, in part, relates to exactly ability to compactly fold these wings; and complete met- which myriapod, whether millipede, centipede, symphy- amorphosis. lan, or pauropod, shares a common ancestor with in- The oldest insect fossils are pieces of Collembola sects. from the Rhvnie Chert of Scotland. Several specimens Insects as Hexapods. A well-supported tree show- including the described Rhyniella precursor have been ing phylogenetic relationships among living six-legged found in these lower Devonian deposits dated to ap- arthropods is shown in Figure 2. To produce this ar- proximately 400 million years ago. These specimens are rangement, Wheeler and colleagues (2001) reviewed the the first in the fossil record to show the six-legged con- vast morphological literature and added characters dition and having the body segmentation coalesced into given by sequences from two genes. It has become com- three main body regions: head, thorax, and abdomen. mon practice in recent years to list, the class to which Thus, the minimum age of the insect clade is 400 million insects belong as Hexapoda. But a rationale for recog- years. nition of all six-legged artliropods as class Insecta is sup- Numerous authors have speculated on the advantage portable as well. A primary reason for recognizing class that six legs and three body regions might have con- Hexapoda (rather than class Insecta) is generally given ferred on these species. The dual tripod gait, with six legs as the lack of resolved phylogenetic relationships among allows significant stability with few contact points on Collembola, Diplura, and Protura. Advocates maintain the substrate. Functional morphologists have studied that a named basal node will clarify the relationships this phenomenon, as have mechanical robot designers. with taxa further up the tree. Also, a need is articulated A metachronal gait is employed by most terrestrial ar- to recognize individual names for nodes from which Ar- thropods. It is typified by lifting one leg at a time from chaeognatha, Zygentoma, and winged insects emanate. the substrate while keeping the leg behind it down to The five groups of wingless animals in question here have bear the animal's weight. This allows good stability been included in the "Apterygota" in the past. Apterygote, while decreasing net energy use. By decreasing the num- in this sense, is clearly a paraphyletic group, and its use ber of legs to six (from the myriapod condition of many), as a formal clade name should be discouraged. Most mechanical simplification is achieved. Although it is in- (but not all) recent reviews of the classification status tuitive that simplification with no loss of stability or of these five orders list Collembola, Diplura, and Protura speed could give an organism a selective advantage, we as a monophyletic group, Entognatha This status is must note that other gait systems work. Is the six-legged based on the presence of mouthparts that can retract condition a breakthrough adaptation regarding diver- into a facial pouch, as well as several other character- sity? Not necessarily, but it could have allowed chan- istics. Archaeognatha and Zygentoma each form mono- neling of energy and integration resources elsewhere. phyletic groups (Figure 2). The rationale for use of the Resources needed for multiple limbs could be diverted taxon name Hexapoda does not hinge on attempts to to other important features or functions necessary for avoid paraphyletic groups. The second rationale, nam- survival and reproduction. Additionally, limbs freed ing of every node in a phylogenetic sequence, has been from use during locomotion could take on new roles as criticized regularly in the past as redundant and unnec- mouthparts for enhanced food acquisition and manipu- essary. On a more positive note, several characters sup- lation. Ancestral legs could also be modified into com- porting monophyly of Hexapoda (major body regions of plex genitalia and enhanced reproduction. head, thorax, and abdomen; six legs) have a long tradi- Tagmatization, the specialization of different regions tional use as defining characteristics for Insecta in the of the body for different functions, is not unique to in- general scientific community and for the public as well. sects, arthropods, or protostomes for that matter. What Although "tradition" alone is not a valid argument for a is unique for the head, thorax, and abdomen arrange- particular naming scheme, stability of use is desirable. ment seen in insects, however, is a significant simplifi- Here, class Insecta is used for the node connecting the cation of the annelidlike or myriapod model. The head Entognatha with the remainder of six-legged arthro- in insects is largely responsible for environmental sen- pods. sory perception and food acquisition. The thorax bears Insect Phylogeny and Key Innovations. We can legs and wings in insects. It can be considered the lo- begin to answer questions such as "Why are there so comotion center of the animal. In contrast, the abdo- many kinds of insects?" by considering a few key ad- men s particular functions of food processing and waste aptations. Six breakthroughs in morphological adapta- removal can be considered more as retained ancestral tion are largely responsible for the success of insects.
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