3785The ability to detect and locate the source of a distant chemical stimulus is an essential process in the lives of benthic crustaceans. Decapod crustaceans, including Achelata (spiny lobsters), Homarida (clawed lobsters), Astacida (crayfish) and Brachyura (crabs), rely on chemical signals to drive a diversity of behaviors ranging from conspecific interactions (Atema, 1995;Karavanich and Atema, 1998; Dunham, 1999, 2000;Gleeson, 1982Gleeson, , 1991 and predator avoidance (Berger and Butler, 2001) to den selection (Berger and Butler, 2001; Eggleston, 1998, 2000;Nevitt et al., 2000), grooming behaviors (Barbato and Daniel, 1997;Daniel et al., 2001) and food detection and localization (Kanciruk, 1980;Reeder and Ache, 1980;Devine and Atema, 1982;Giri and Dunham, 1999;Dunham et al., 1997;Keller et al., 2003).Chemical stimuli are detected by a multitude of chemoreceptive structures on crustaceans. Although chemoreceptive sensilla can be found on virtually all body surfaces, they are most concentrated on the appendages, particularly the antennules, antennae, dactyls and mouthparts (Ache and Macmillan, 1980;Derby, 1982;Schmidt, 1989;Schmidt and Gnatzy, 1984; Derby, 2001, 2002a;Garm et al., 2003). The antennules in particular have long been considered to be the primary chemoreceptive organ of the spiny lobster (Fig.·1). Each antennule is composed of four segments, the most distal of which bifurcates into a lateral flagellum and a medial flagellum. Each flagellum is composed of annuli that bear a complement of chemo-and mechanosensory sensilla that vary in morphology, distribution and pattern of innervation. Many studies have shown that the antennules are important for distance chemoreception in lobsters (Reeder and Ache, 1980;Devine and Atema, 1982) and other decapod crustaceans (Hazlett, 1971a;Kraus-Epley and Moore, 2002); however, it is not clear which populations of antennular sensilla are involved in this behavior.Chemosensory information from the antennular sensilla is transmitted to the central nervous system in two parallel pathways: the aesthetasc/olfactory lobe pathway and the nonaesthetasc/lateral antennular neuropil pathway (Schmidt and Ache, 1992, 1996a,b;. The aesthetasc/ olfactory lobe pathway originates in clusters of olfactory Benthic crustaceans rely on chemical stimuli to mediate a diversity of behaviors ranging from food localization and predator avoidance to den selection, conspecific interactions and grooming. To accomplish these tasks, Caribbean spiny lobsters (Panulirus argus) rely on a complex chemosensory system that is organized into two parallel chemosensory pathways originating in diverse populations of antennular sensilla and projecting to distinct neuropils within the brain. Chemosensory neurons associated with aesthetasc sensilla project to the glomerular olfactory lobes (the aesthetasc pathway), whereas those associated with non-aesthetasc sensilla project to the stratified lateral antennular neuropils and the unstructured median antennular neuropil (the nonaesthetasc pathway). Althoug...