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homolateral; axons extend forwards only on the side of the brain from which they originate. The higher Malacostraca, including the stomatopods, have projections that are heterolateral; axons from each olfactory lobe branch to both sides of the protocerebrum. Thus, whereas in higher malacostracans, and indeed in dicondylic insects, pathways from the olfactory lobes show many elaborations, when comparing basal taxa the ground plan of the insect olfactory system is almost identical to that of basal malacostracans. The third center to be considered is called the central complex. This is a system of interconnected neuropils present in the brains of all arthropods that reaches great elaboration and complexity in the neopteran insects. Common to all arthropods, the central complex is associated with, and in orthopterous insects is known to develop from, a specific set of pioneer axons. These fasciculate to cross the midline via a rostral commissure, and then shift their locations to a more posterior commissure and in so doing set up a system of chiasmata that lay down the basic framework for a neuropil that in the Neoptera consists of sixteen modules or subunits (Boyan et al., 2008). This switch of an axon trajectory from one commissure to another behind it is unique to the protocerebrum and has been termed by Boyan et al. (2008) as “fascicle switching”. Modules that result from this axonal rerouting can develop in two neuropils: the one that is associated with the rostral commissure, called the protocerebral bridge, and one that is associated with the more posterior commissure to which fascicle-switching axons project, called the central body. In the Neoptera, the central body comprises two major divisions, the fan-shaped body and the ellipsoid body, connected to a pair of small, layered neuropils called the noduli (Loesel et al., 2002). This mid-line central complex, which is now known to be associated with the control of limb movements (see Poeck et al., 2008), has reached its greatest elaboration in those insects and crustaceans that show high levels of dexterity, but is much simpler in organization in species that have simpler modes of locomotion. In the Machilidae, the central complex consists of two layers of neuropil supplied by rostral axons that decussate into the central body neuropil through a system of chiasmata. There is no protocerebral bridge neuropil, nor noduli. Instead, the neuropil of the central body has the appearance of an elongated spindle, with two distinctive strata. The same organization typifies non-reptantian decapod crustaceans, such as caridids (shrimps). Their central complexes and those of machilids appear identical (Strausfeld, 2009). Examination of entomostracan brains also reveals a midline neuropil, but this is both extremely small, has a single chiasmatal element, and is entirely unstratified. Critics will certainly claim that the three examples reviewed above are insufficient to show that malacostracans and insects might derive from a common ancestor. And indeed there is no way of demonstrating conclusively that they have or have not. However, would the insects have arisen from an entomostracan-like ancestor (see Glenner et al., 2006), and would another entomostracan-like antecedent have also given rise to the malacostracans, then each of the features discussed above would have had to evolve independently in the Malacostraca and Insecta. The organization of the olfactory lobes and their central projections would then have to be interpreted as examples of convergence, as would the evolution of four nested optic neuropils and their chiasmata. Likewise, central complexes that appear to have identical architectures in machilids and malacostracans would have had to have evolved convergently. On the other hand, the most parsimonious explanation for all these commonalities is that these features common to the brains of insects and malacostracans were also present on a common malacostracan-like ancestor. There are many other features of insect brains, which have not been considered here, that speak to a malacostracan ancestry. One is the connection between the labral neuropils and the dorsal protocerebrum. Another is the organization 25
of mechanosensory centers in the tritocerebrum, which are so distinctive in malacostracans and insects, yet appear to be lacking in the Entomostraca. A satisfactory resolution to discrepancies between neurophylogenetic reconstructions of arthropod relationships and reconstructions derived from molecular phylogenetics are not necessarily insoluble. Clarification of the position of the Remipedia may be crucial. Better resolution may also be obtained if molecular and neural characters can be combined in future analyses. References: Boyan GS, Williams JL, Herbert Z. 2008. Fascicle switching generates a chiasmal neuroarchitecture in the embryonic central body of the grasshopper Schistocerca gregaria. Arthropod Struct Dev. 37:539-544. Dohle, W. 2001. Are the insects terrestrial crustaceans? A discussion of some new facts and arguments and the proposal of the proper name ‘Tetraconata’ for the monophyletic unit Crustacea Hexapoda. Ann. Soc. Entomol. France 37, 85–103. Fanenbruck M, Harzsch S. 2005 A brain atlas of Godzilliognomus frondosus Yager, 1989 (Remipedia, Godzilliidae) and comparison with the brain of Speleonectes tulumensis Yager, 1987 (Remipedia, Speleonectidae): implications for arthropod relationships. Arthropod Struct. Dev. 34, 343–378. Glenner H, Thomsen PF, Hebsgaard MB, Sorensen MV, Willerslev E. 2006 The origin of insects. Science 314, 1883–1884. Hanström, B. 1926 Vergleichende Anatomie des Nervensystems der wirbellosen Tiere. (Facsimile reprint by A. Asher, Amsterdam 1968.) Holmgren, N. 1916 Zur vergleichenden Anatomie des Gehirns von Polychaeten, Onychophoren, Xiphosuren, Arachniden, Crustaceen, Myriapoden, und Insekten. K. Svenska Vetensk. Akad. Handl. 56, 1–200. Labandeira CC, Beal BS, Hueber FM. 1988. Early insect diversification: evidence from a Lower Devonian bristletail from Quebec. Science 242, 913–916. Lichtneckert R, Nobs L, Reichert H. 2008. Empty spiracles is required for the development of olfactory projection neuron circuitry in Drosophila. Development. 135:2415-2424. Loesel R, Nässel DR, Strausfeld NJ. 2002. Common design in a unique midline neuropil in the brains of arthropods. Arthropod Struct Dev. 31:77-91. Mallatt J, Giribet G. 2006 Further use of nearly complete, 28S and 18S rRNA genes to classify Ecdysozoa, 37 more arthropods and a kinorhynch. Mol. Phylogen Evol. 40, 772–794. Meinertzhagen, I. A. 1991 Evolution of the cellular organization of the arthropod compound eye and optic lobe. In Vision and visual dysfunction, vol. 2 (eds. J. R. Cronly-Dillon & R. L. Gregory). Evolution of the Eye and Visual System, pp. 341–362. London, UK: MacMillan Press. Meinertzhagen IA, Hanson TA. 1993 The development of the optic lobes. In The development of Drosophila melanogaster (eds M. Bate & A. M. Arias), pp. 1363–1491. New York, NY: Cold Spring Harbor Laboratory Press. Poeck B, Triphan T, Neuser K, Strauss R. 2008. Locomotor control by the central complex in Drosophila-An analysis of the tay bridge mutant. Dev Neurobiol. 68:1046- 1058. Regier JC, Shultz JW, Ganley AR, Hussey A, Shi D, Ball B, Zwick A, Stajich JE, Cummings MP, Martin JW, Cunningham CW. 2008. Resolving arthropod phylogeny: exploring phylogenetic signal within 41 kb of protein-coding nuclear gene sequence. Syst Biol. 57:920-938 26
Page 1 and 2: ISBN 978-88-96493-00-7 XXII Congres
Page 3 and 4: © 2009 Accademia Nazionale Italian
Page 6 and 7: SESSIONI E COMITATO SCIENTIFICO I.
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Sessione IV ENTOMOLOGIA FORESTALE
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Sessione IV - Entomologia forestale
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Sessione V - Ecologia e Etologia IL
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Sessione V - Ecologia e Etologia TA
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Sessione VI - Entomologia agraria I
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Sessione VI - Entomologia agraria S
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Sessione VI - Entomologia agraria R
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Sessione VI - Entomologia agraria A
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Sessione VI - Entomologia agraria V
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Sessione VI - Entomologia agraria B
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Sessione VIII BIOTECNOLOGIE ENTOMOL
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Sessione IX ENTOMOLOGIA MERCEOLOGIC
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Sessione X - Controllo biologico DA
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Sessione X - Controllo biologico MI
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Workshop Martedì, 16 giugno 2009 -
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INDICI
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Campolo · 90; 107; 121; 130; 132;
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I Iafigliola · 102; 320 Iannotta
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Pinzauti · 113 Piras · 76; 125; 3
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SOMMARIO PROGRAMMA XV LETTURE PLENA
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XXII CNIE - Accademia nazionale italiana di Entomologia

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