Ecology and Behaviour of the Ladybird Beetles (Coccinellidae)

Ivo Hodek, A. Honek, Helmut F. van Emden, Ivo Hodek, A. Honek, Helmut F. van Emden

• Publisher: Wiley-Blackwell
• Year: 2012
• Print ISBN: 9781405184229
• eBook ISBN: 9781118223222
• Edition: 1
• Topic: Biological Sciences
• Subtopic: Entomology
• Index: Biological Sciences

Ladybirds are probably the best known predators of aphids and coccids in the world, though this greatly underestimates the diversity of their biology. Maximising their impact on their prey is an important element in modern conservation biological control of indigenous natural enemies in contrast to the classical approach of releasing alien species.

Ivo Hodek is one of the most internationally respected experts on coccinellids who has researched these insects for his entire career. He has now brought together 14 scientists of international standing to author 12 chapters, making this book the definitive treatment of coccinellid biology and ecology.

This volume covers the rapid scientific developments of recent years in the understanding of coccinellid phylogeny, the semiochemicals influencing their behaviour and of molecular genetics. Recent insights in relation to intraguild predation and the assessment of the predatory impact of coccinellids are also covered.

Other special features of the volume are the extensive references covering the literature from both East and West and a taxonomic glossary of the up-to-date nomenclature for species of coccinellids as well as of other organisms mentioned in the text.

While aimed at researchers, university teachers and agricultural entomologists, the book is readable and appropriate for others who just have a liking for these interesting and attractive insects.

Chapter 1

Phylogeny and Classification

Oldrich Nedv

d¹ and Ivo Ková

²

¹ Faculty of Science, University of South Bohemia & Institute of Entomology, Academy of Sciences, CZ 37005

eské Bud

jovice, Czech Republic

² Emer. Scientist of the National Museum, Prague; Current address: Zichovec, Czech Republic

1.1 POSITION OF THE FAMILY

1.1.1 The Cerylonid Complex

The family Coccinellidae includes approximately 6000 described species in some 360 genera and 42 tribes. Coccinellids belong to the superfamily Cucujoidea of the Coleoptera suborder Polyphaga, and the family is a member of the phylogenetic branch frequently referred to as the Cerylonid complex or series of families, which is composed of Alexiidae, Cerylonidae, Coccinellidae, Corylophidae, Discolomatidae, Endomychidae (s. lat. including Mychotheninae, Eidoreinae and Merophysiinae) and Latridiidae (Crowson 1955, Lawrence & Newton 1995). Bothrideridae were added later (Pal & Lawrence 1986). Monophyly of the Cerylonid series was based on morphological characters (Slipinski & Pakaluk 1992) and confirmed by parsimony analysis of molecular data (Hunt et al. 2007, Robertson et al. 2008).

Phylogenetic relations between the families and subfamilies included in the Cerylonid complex are rather complicated and not fully resolved (Slipinski & Pakaluk 1992). Early morphological studies supported a clade Endomychidae plus Corylophidae as the sister group of Coccinellidae (Crowson 1955, Sasaji 1971a). Affinities have been proposed between Endomychidae and Coccinellidae (Pakaluk & Slipinski 1990; Burakowski & Slipinski 2000) due to the presence of a characteristic basal (= median) lobe of male genitalia, pseudotrimerous tarsi and absence of coronal suture on the head of larvae (except in Epilachninae); Eupsylobiinae (Endomychidae) and Coccinellidae (Pakaluk & Slipinski 1990) due to the long penis and coccinellid-like tegmen; Mycetaeinae (Endomychidae) and Coccinellidae (number of abdominal spiracles, open middle coxal cavity, Ková

1996; distinct pronotum with sublateral carina, hidden mesotrochantin, Tomaszewska 2000); Alexiidae (= Sphaerosomatidae) and Coccinellidae (Slipinski & Pakaluk 1992); Endomychidae plus Alexiidae and Coccinellidae (Pakaluk and Slipinski 1990); or between Corylophidae and Coccinellidae (Sasaji 1971a; anterior tentoria separated, frontoclypeal suture absent, common type of antenna and tarsus, Tomaszewska 2005).

1.1.2 Sister Families

Tomaszewska (2000) argued monophyly of the Endomychidae plus Coccinellidae from the common characters of procoxal cavity externally open, tarsal formula 4-4-4, abdomen with five pairs of functional spiracles and median lobe without additional struts. However, the polyphyletic nature of the family Endomychidae has been known for a long time (Slipinski & Pakaluk 1992) and was confirmed with molecular analyses by Robertson et al. (2008) that grouped the subfamily Anamorphinae with the Corylophidae, and the rest with the Coccinellidae. Later analysis by Tomaszewska (2005), based on adult and larval characters, failed to confirm a sister relationship of the Endomychidae and Coccinellidae.

The comprehensive molecular study of beetles by Hunt et al. (2007) placed the family Alexiidae or the pair of Alexiidae plus Anamorphinae (Endomychidae) as the sister group of the Coccinellidae. Molecular analysis of the Cerylonid complex by Robertson et al. (2008) proposed the subfamily Leiestinae (Endomychidae) or the complex of Endomychidae (part) plus (Corylophidae plus Anamorphinae (Endomychidae)) as sister groups of the Coccinellidae.

1.1.3 Feeding Habits

In either case, the sister group of Coccinellidae, and their common ancestor, were probably fungivorous (Giorgi et al. 2009). Feeding on hemipterans (mainly Sternorrhyncha) has evolved predominantly in coleopteran lineages that contain fungus feeders (Derodontidae, Silvanidae, Laemophloeidae, Nitidulidae, Endomychidae, Anthribidae) and whose ancestors were fungus feeders (Coccinellidae) or sap feeders (Scarabaeidae: Cetoniinae) (Leschen 2000).

1.1.4 Monophyly of Coccinellidae

The monophyly of the family Coccinellidae was based on morphological synapomorphies, and has repeatedly been supported by molecular phylogenetic analyses (Hunt et al. 2007, Robertson et al. 2008, Giorgi et al. 2009), even early ones (Howland & Hewitt 1995) that did not place the family in the Cucujiformia. The first to recognize the morphological synapomorphy (although the term did not exist at that time) of the family, the siphonal structure of penis, was Verhoeff (1895).

1.2 CHARACTERISTICS OF THE FAMILY

The family Coccinellidae may be distinguished from the rest of the cerylonid complex by the combination of the following adult characters: (i) five pairs of abdominal spiracles, (ii) tentorial bridge absent, (iii) anterior tentorial branches separated, (iv) frontoclypeal suture absent, (v) apical segment of maxillary palps never needle-like, (vi) galea and lacinia separated, (vii) mandible with reduced mola, (viii) procoxal cavities open posteriorly, (ix) middle coxal cavities open outwardly, (x) meta-epimeron parallel sided, (xi) femoral lines present on abdominal sternite 2, (xii) tarsal formula 4-4-4 or 3-3-3, tarsal segment 2 usually strongly dilated below, (xiii) male genitalia with tubular curved sipho, distally embraced by the tegmen. Larvae are armed with setae and setose processes (three pairs on each abdominal segment), antennae of one to three segments, not over three times as long as wide, frontoclypeal suture absent (except some Epilachnini). Pupae of coccinellids are of the type pupa adectica obtecta – all appendages are glued to the body by exuvial fluid. The pupa is attached to the substrate by the tip of the abdomen. Coccidophagy is a synapomorphy for the family, other feeding habits, including mycophagy, are secondary (Giorgi et al. 2009).

Liere and Perfecto (2008) suggested that the evolution of coccinellid morphology was influenced by relationships with ants. The result of a parallel evolution in several subfamilies is a mosaic of various grades of characters and/or apparent similarities, which mosaic may considerably obscure the true phylogenetic relationships within the family. One of these characters may be the enlarged eye canthus, formerly considered an apomorphy of the Chilocorinae, but which probably evolved independently in the Platynaspidini (Slipinski 2007). Mouthpart morphology was used in determining diet and host specificity in Coccinellidae (Klausnitzer 1993, Klausnitzer & Klausnitzer 1997), but it is also constrained phylogenetically (Samways et al. 1997). Major larval types were defined by LeSage (1991). Pupal morphology was used for phylogenetic purposes separately from other characters (Phuoc & Stehr 1974).

1.3 CHANGES IN THE CLASSIFICATION OF SUBFAMILIES

1.3.1 Morphologically Based Classifications

Six subfamilies of Coccinellidae were recognized by Sasaji (1968, 1971b): Sticholotidinae, Coccidulinae, Scymninae + Chilocorinae, Coccinellinae + Epilachninae. However, his knowledge of non-Japanese taxa was limited. Korscheffsky (1931) classified the genus Lithophilus in a separate subfamily on account of the tetramerous structure of tarsus. Klausnitzer (1969, 1970, 1971) distinguished this subfamily (as Tetrabrachinae) from its sister group Coccidulinae based mainly on six sclerites on the pronotum of larvae as apomorphy, while he considered adult tetrameric tarsi as a plesiomorphy. Ková

(1973) added this seventh subfamily (as Lithophilinae) to Sasaji’s phylogenetic tree, but later reassigned it to tribal level within the Coccidulinae and proposed another subfamily, the Ortaliinae (Ková

1996). Chazeau et al. (1990) retained Sasaji’s six subfamilies with slightly rearranged tribes.

The monophyly of several of the six subfamilies proposed by Sasaji has been disputed. The higher classification of Coccinellidae appears to suffer from the presence of para- and polyphyletic taxa (Vandenberg 2002). This problem is most conspicuous in the formerly basal subfamilies Sticholotidinae and Coccidulinae, and in the fauna of poorly studied regions.

1.3.2 Split of Sticholotidinae

In morphology based studies, Duverger (2003), Slipinski and Tomaszewska (2005) and Vandenberg and Perez-Gelabert (2007) proposed dividing up the highly diverse subfamily Sticholotidinae. In Sticholotis and allies, the terminal segment of the maxillary palp has a long obliquely oriented distal sensory surface on one side of the tapered apex, similar to the large sensory surface of the securiform (axe-shaped) palps of derived ‘true’ ladybirds; while in Sukunahikona and allies the sensory surface is small, oval and positioned distally, suggesting a basal position for this group. The subfamily Sticholotidinae also formed an unresolved polytomy in cladistic analysis by Yu (1994).

The subfamily Sticholotidinae was established by Sasaji (1968) with four tribes, whilst Sticholotidinae sensu Ková

(1996) contained 10 tribes. However, the phylogenetic relations among the constituent tribes remained obscure for a long time, in spite of an attempt at clarification by Gordon (1977). These Sticholotidinae s. lat. were characterized (Ková

1996), among other traits, by the shape of the apical maxillary palp segment which differs from the rest of Coccinellidae (narrowed apically, conical or barrel shaped). However, this applies to the recently re-established Microweiseinae (sensu Slipinski 2007), while in the Sticholotidinae s. str. the last segment is more or less enlarged, approaching the typical securiform shape found in the other Coccinellidae.

These two lineages differ substantially, such as in the shape of the metendosternite, presence or absence of anterolateral carinae of the pronotum, and also in characters of both male a...

Table of contents

• Cover
• Title page
• Copyright page
• Dedication
• Contributors
• Preface
• Introduction
• Taxonomic Glossary
• Chapter 1 Phylogeny and Classification
• Chapter 2 Genetic Studies
• Chapter 3 Life History and Development
• Chapter 4 Distribution and Habitats
• Chapter 5 Food Relationships
• Chapter 6 Diapause/Dormancy
• Chapter 7 Intraguild Interactions
• Chapter 8 Natural Enemies of Ladybird Beetles
• Chapter 9 Coccinellids and Semiochemicals
• Chapter 10 Quantifying the Impact of Coccinellids on their Prey
• Chapter 11 Coccinellids in Biological Control
• Chapter 12 Recent Progress and Possible Future Trends in the Study of Coccinellidae
• Appendix: List of Genera in Tribes and Subfamilies
• SUBJECT INDEX
• Color Plates