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Phylogeny and Classification Chapter 1 Phylogeny and Classification Oldrich Nedv eˇ d 1 and Ivo Kov á rˇ 2 1 Faculty of Science, University of South Bohemia & Institute of Entomology, Academy of Sciences, CZ 37005 Cˇ esk é Bud eˇ jovice, Czech Republic 2 Emer. Scientist of the National Museum, Prague; Current address: Zichovec, Czech Republic Ecology and Behaviour of the Ladybird Beetles (Coccinellidae), First Edition. Edited by I. Hodek, H.F. van Emden, A. Honeˇk. © 2012 Blackwell Publishing Ltd. Published 2012 by Blackwell Publishing Ltd. 1 1.1 POSITION OF THE FAMILY 1.1.2 Sister families 1.1.1 The Cerylonid complex Tomaszewska (2000) argued monophyly of the Endomychidae plus Coccinellidae from the common The family Coccinellidae includes approximately 6000 characters of procoxal cavity externally open, tarsal described species in some 360 genera and 42 tribes. formula 4 - 4 - 4, abdomen with fi ve pairs of functional Coccinellids belong to the superfamily Cucujoidea of spiracles and median lobe without additional struts. the Coleoptera suborder Polyphaga, and the family is a However, the polyphyletic nature of the family Endomy- member of the phylogenetic branch frequently referred chidae has been known for a long time (Slipinski & to as the Cerylonid complex or series of families, Pakaluk 1992 ) and was confi rmed with molecular which is composed of Alexiidae, Cerylonidae, Coccinel- analyses by Robertson et al. (2008) that grouped the lidae, Corylophidae, Discolomatidae, Endomychidae ( s. subfamily Anamorphinae with the Corylophidae, and lat. including Mychotheninae, Eidoreinae and Mero- the rest with the Coccinellidae. Later analysis by physiinae) and Latridiidae (Crowson 1955 , Lawrence Tomaszewska (2005) , based on adult and larval char- & Newton 1995 ). Bothrideridae were added later (Pal acters, failed to confi rm a sister relationship of the & Lawrence 1986 ). Monophyly of the Cerylonid series Endomychidae and Coccinellidae. was based on morphological characters (Slipinski & The comprehensive molecular study of beetles by Pakaluk 1992 ) and confi rmed by parsimony analysis Hunt et al. (2007) placed the family Alexiidae or the of molecular data (Hunt et al. 2007 , Robertson et al. pair of Alexiidae plus Anamorphinae (Endomychidae) 2008 ). as the sister group of the Coccinellidae. Molecular Phylogenetic relations between the families and analysis of the Cerylonid complex by Robertson et al. subfamilies included in the Cerylonid complex are (2008) proposed the subfamily Leiestinae (Endomychi- rather complicated and not fully resolved (Slipinski dae) or the complex of Endomychidae (part) plus & Pakaluk 1992 ). Early morphological studies sup- (Corylophidae plus Anamorphinae (Endomychidae)) ported a clade Endomychidae plus Corylophidae as the as sister groups of the Coccinellidae. sister group of Coccinellidae (Crowson 1955 , Sasaji 1971a ). Affi nities have been proposed between Endomychidae and Coccinellidae (Pakaluk & Slipinski 1.1.3 Feeding habits 1990 ; Burakowski & Slipinski 2000 ) due to the presence of a characteristic basal ( = median) lobe of In either case, the sister group of Coccinellidae, male genitalia, pseudotrimerous tarsi and absence and their common ancestor, were probably fungivo- of coronal suture on the head of larvae (except in rous (Giorgi et al. 2009 ). Feeding on hemipterans Epilachninae); Eupsylobiinae (Endomychidae) and (mainly Sternorrhyncha) has evolved predominantly Coccinellidae (Pakaluk & Slipinski 1990 ) due to in coleopteran lineages that contain fungus feeders the long penis and coccinellid - like tegmen; Mycetaei- (Derodontidae, Silvanidae, Laemophloeidae, Nitiduli- nae (Endomychidae) and Coccinellidae (number of dae, Endomychidae, Anthribidae) and whose ancestors abdominal spiracles, open middle coxal cavity, Kov á rˇ were fungus feeders (Coccinellidae) or sap feeders 1996 ; distinct pronotum with sublateral carina, (Scarabaeidae: Cetoniinae) (Leschen 2000 ). hidden mesotrochantin, Tomaszewska 2000 ); Alexii- dae ( = Sphaerosomatidae) and Coccinellidae (Slipinski & Pakaluk 1992 ); Endomychidae plus Alexiidae and 1.1.4 Monophyly of Coccinellidae Coccinellidae (Pakaluk and Slipinski 1990 ); or between Corylophidae and Coccinellidae (Sasaji 1971a ; ante- The monophyly of the family Coccinellidae was based rior tentoria separated, frontoclypeal suture absent, on morphological synapomorphies, and has repeatedly common type of antenna and tarsus, Tomaszewska been supported by molecular phylogenetic analyses 2005 ). (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 fi rst to recognize the morphological synapomor- phy (although the term did not exist at that time) of Phylogeny and classifi cation 3 the family, the siphonal structure of penis , was 1.3 CHANGES IN THE Verhoeff (1895) . CLASSIFICATION OF SUBFAMILIES 1.3.1 Morphologically based classifi cations 1.2 CHARACTERISTICS OF THE FAMILY Six subfamilies of Coccinellidae were recognized by Sasaji (1968, 1971b) : Sticholotidinae, Coccidulinae, The family Coccinellidae may be distinguished from Scymninae + Chilocorinae, Coccinellinae + Epilachni- the rest of the cerylonid complex by the combination nae. However, his knowledge of non - Japanese taxa was of the following adult characters : (i) fi ve pairs of limited. Korscheffsky (1931) classifi ed the genus abdominal spiracles, (ii) tentorial bridge absent, (iii) Lithophilus in a separate subfamily on account of the anterior tentorial branches separated, (iv) frontoclypeal tetramerous structure of tarsus. Klausnitzer (1969, suture absent, (v) apical segment of maxillary palps 1970, 1971) distinguished this subfamily (as Tetra- never needle- like, (vi) galea and lacinia separated, brachinae) from its sister group Coccidulinae based (vii) mandible with reduced mola, (viii) procoxal cavi- mainly on six sclerites on the pronotum of larvae ties open posteriorly, (ix) middle coxal cavities open as apomorphy, while he considered adult tetrameric outwardly, (x) meta - epimeron parallel sided, (xi) tarsi as a plesiomorphy. Kov á rˇ (1973) added this femoral lines present on abdominal sternite 2, seventh subfamily (as Lithophilinae) to Sasaji ’ s phylo- (xii) tarsal formula 4 - 4 - 4 or 3 - 3 - 3, tarsal segment 2 genetic tree, but later reassigned it to tribal level within usually strongly dilated below, (xiii) male genitalia the Coccidulinae and proposed another subfamily, with tubular curved sipho, distally embraced by the the Ortaliinae (Kov á rˇ 1996 ). Chazeau et al. (1990) tegmen. Larvae are armed with setae and setose retained Sasaji ’ s six subfamilies with slightly rear- processes (three pairs on each abdominal segment), ranged tribes. antennae of one to three segments, not over three The monophyly of several of the six subfamilies times as long as wide, frontoclypeal suture absent proposed by Sasaji has been disputed. The higher (except some Epilachnini). Pupae of coccinellids are of classifi cation of Coccinellidae appears to suffer from the the type pupa adectica obtecta – all appendages are presence of para - and polyphyletic taxa (Vandenberg glued to the body by exuvial fl uid. The pupa is attached 2002 ). This problem is most conspicuous in the to the substrate by the tip of the abdomen. Coccido- formerly basal subfamilies Sticholotidinae and Coccid- phagy is a synapomorphy for the family, other feeding ulinae, and in the fauna of poorly studied regions. habits, including mycophagy, are secondary (Giorgi et al. 2009 ). Liere and Perfecto (2008) suggested that the ev - 1.3.2 Split of Sticholotidinae olution of coccinellid morphology was infl uenced by relationships with ants. The result of a parallel evolu- In morphology based studies, Duverger (2003) , Slipin- tion in several subfamilies is a mosaic of various grades ski and Tomaszewska (2005) and Vandenberg and of characters and/or apparent similarities, which Perez - Gelabert (2007) proposed dividing up the highly mosaic may considerably obscure the true phyloge- diverse subfamily Sticholotidinae . In Sticholotis netic relationships within the family. One of these and allies, the terminal segment of the maxillary palp characters may be the enlarged eye canthus, formerly has a long obliquely oriented distal sensory surface considered an apomorphy of the Chilocorinae, but on one side of the tapered apex, similar to the large which probably evolved independently in the Platy- sensory surface of the securiform (axe - shaped) palps naspidini (Slipinski 2007 ). Mouthpart morphology of derived ‘ true ’ ladybirds; while in Sukunahikona was used in determining diet and host specifi city and allies the sensory surface is small, oval and posi- in Coccinellidae (Klausnitzer 1993 , Klausnitzer & tioned distally, suggesting a basal position for this Klausnitzer 1997 ), but it is also constrained phylo- group. The subfamily Sticholotidinae also formed an genetically (Samways et al. 1997 ). Major larval types unresolved polytomy in cladistic analysis by Yu were defi ned by LeSage (1991) . Pupal morphology was (1994) . used for phylogenetic purposes separately from other The subfamily Sticholotidinae was established by characters (Phuoc & Stehr 1974 ). Sasaji (1968) with four tribes, whilst Sticholotidinae 4 O. Nedveˇd and I. Kovárˇ sensu Kov á rˇ (1996) contained 10 tribes. However, the independent, based mainly on morphology of larvae, phylogenetic relations among the constituent tribes while the tribe Coccinellini was considered polyphyletic remained obscure for a long time,
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