K.V. Makarov: "Morphology of Termitophilous Larvae of Orthogonius and Helluodes..." (1998)

 
K.V. Makarov 
 
Morphology of Termitophilous Larvae of Orthogonius and Helluodes and 
Position of the Tribes Orthogoniini and Helluodini in the Family Carabidae (Coleoptera) 
 
Entomological Review, Vol. 78, No. 9, 1998, pp. 1115-1125. 
 
Translated from Zoologicheskii Zhurnal, Vol. 77, No. 11, 1998. 
 
Abstract. Termitophilous larvae of the ground beetles Orthogonius acutangulus Chaud. and Helluodes taprobanae 
Westwood (Ceylon) are described. Larvae of O. acutangulus have a strongly differentiated secondary 
chaetome, lacking sensilla PA1, 2, 3, while setae ME13, 14 and TE1, 10, 11 have specifically dilated bases. According to 
the general character set, Orthogoniini cannot be considered closely related to either Lebiitae or Masoreitae 
(Graphipterini). A number of significant larval characters, such as well-developed lacinia, biserial nasale, ligula 
with additional sensilla (LA7), differentiated group of maxillary setae, pattern of the secondary leg chaetome, and 
structure of tarsus, suggest that Orthogoniini are remotely related to Pterostichitae and Harpalitae. Larvae of H. 
taprobanae, strongly differing from Orthogonius in habitus, have a number of convergently similar characters: 
weak pigmentation, reduced ocelli, etc. The unique characters of Helluodes are the specific morphology of ligula 
and the presence of a stout seta on 2nd maxillary palpomere. Larvae of Helluodini share no significant characters 
with the preimaginal stages of Anthiini and Helluonini. At the same time, Helluodes and Zuphiini have many significant 
larval characters in common: medially-directed angles of paraclypeus; elongate apical part of 4th antennal 
segment; 3rd antennal segment bearing a peculiarly flattened sensorium; small retinaculum; and absence of penicillus. 
Some of these characters are not found in other carabid larvae and thus can be considered synapomorphic, 
suggesting that Helluodini is a specialized group, derived from Zuphiini.