Differentiation of the Carabid Antenna Cleaner

Abstract

The antennae of most carabids are groomed by tightly packed setae on the protibia. The mouthparts are not employed in antenna cleaning in ground beetles, but are so used in many 'Coleoptera (Jander, 1966, table 2). A protibial antenna cleaner may be primitively absent only in Nototylus (Banninger 1927: 771) but is secondarily lost in advanced paussines (Darlington 1950: 65). There is great variation in the degree of development of the cleaning setae and in protibial structure. This paper describes and analyzes structural differentiation of the carabid antenna cleaner, and presents preliminary data on grooming behavior. Work on the fine structure and histology of the cleaning setae and associated glands is in progress and will be reported elsewhere. The protibiae of about 100 genera representing 60 tribes were studied; 50 species were measured. A description of methods employed and a formal list of carabids examined in this study will be presented in a paper on prothoracic morphology of the Coleoptera. Structure. The antenna grooming setae are located on the medial face of the tibia; there are two types of setal aggregations. The major cleaning element is the setal band (Figs. 1, 21 SB). It is composed of very tightly packed setae arranged in a single file that always begins near the anterior spur and may extend nearly horizontally across the width of the medial face (Figs. 2, 7, 9) or may extend vertically up the length of the tibia for a considerable distance (Figs. 1, 3, 15 SB). The second type of cleaning cluster consists of relatively short, less densely packed vertical rows of setae, usually in single file, originating above the tibial spurs (Figs. 1, 21 ASR, PSR). In a few carabids dense setal rows, which are probably used in grooming, extend above both the anterior and posterior J A preliminary version of this work was submitted as a portion of a thesis to the Biology Department, Harvard University, in partial fulfillment of the requirements for the Ph.D. degree.

Phylogeny of carabid beetles as inferred from 18S ribosomal DNA (Coleoptera: Carabidae)

“…Morphological evidence for the placement of paussines as the sister group of other carabids is not abundant. Ancestral paussines had an apparently primitive antenna cleaner (Jeannel 1941; Hlavac 1971), although modifications of the antennae through evolution with ants (Darlington 1950) resulted in reduction of the antenna cleaner in most paussines (Ball & McCleve 1990). Paussines lack some traits of the larval preoral filter (Beutel 1993) present in other carabids, and have female genitalia more primitive in some regards than other carabids (Ball & McCleve 1990; Liebherr & Will 1999).…”

“…Although they possess a number of striking apomorphies (including an explosive chemical defense mechanism in adults and myrmecophilous habits with associated modifications of larval structure), they also possess features considered present in the groundplan of Carabidae. Evidence for the monophyly of remaining carabids (‘Anisochaeta’) is limited, consisting of a tendency of a proximal shift of one spur of the protibial antenna cleaner (Jeannel 1941; Hlavac 1971), characteristics of the preoral filter in larvae (Beutel 1993) and female genitalia (Liebherr & Will 1999). Within Anisochaeta, a number of taxa, including the supertribes Nebriitae (Nebriini, Opisthiini, Notiophilini, Notiokasiini) and Carabitae (Carabini, Pamborini, Cychrini), form an old radiation of basal lineages of carabids having, for the most part, exoskeletal characteristics considered primitive within carabids.…”

“…The large body of literature on carabid phylogeny has failed to reach a consensus about many aspects of tribal‐level relationships (see Ball 1979; Erwin 1979; Kavanaugh & Erwin 1991; Bousquet & Larochelle 1993). Detailed examination of the antenna cleaner (Hlavac 1971; Regenfuss 1975) and thoracic characters (e.g. Kavanaugh & Erwin 1991) indicates that the structures are not distributed in a pattern of simple transformation from ‘basal’ to ‘higher’ carabids.…”

“…In the last few decades, new characters have been examined, ranging from traits of the exoskeleton (e.g. Bell 1964; Bell 1967; Hlavac 1971; Regenfuss 1975; Hammond 1979; Evans 1982; Erwin & Stork 1985; Nichols 1985; Deuve 1988; Kavanaugh & Erwin 1991; Beutel & Haas 1996), wing venation (Ward 1979), female genitalia (Deuve 1993; Liebherr & Will 1999), muscles (Bils 1976; Baehr 1979; Burmeister 1980), ventral nerve cord (Heath & Evans 1990), digestive system (Yahiro 1990; Yahiro 1996), defensive glands and secretions (Moore & Wallbank 1968; Forsyth 1972; Kanehisa & Murase 1977; Moore 1979; Kanehisa & Kawazu 1985), larvae (e.g. Goulet 1983; Bousquet & Smetana 1986; Liebherr & Ball 1990; Beutel 1991; Arndt 1993; Beutel 1993), as well as chromosomes (Serrano 1981a,b; Serrano & Yadav 1984; Serrano 1992).…”

The integripennis species group of Geocharidius Jeannel, 1963 (Carabidae, Bembidiini, Anillina) from Nuclear Central America: a taxonomic review with notes about biogeography and speciation

“…Protibiae (Fig. 6F–I) with antenna cleaner of type B (Hlavac 1971), with both anterior (asr) and posterior (psr) apical setal rows and concave apico-lateral notch (tbn). Profemora moderately swollen.…”