DIMITRI FORERO¹

¹American Museum of Natural History, New York, NY 10024-5192, USA, and Department of Entomology, Comstock Hall, Cornell University, Ithaca, NY 14853–2601, USA. idf2@cornell.edu.

Hemiptera is the fifth largest group of insects after Coleoptera, Diptera, Hymenoptera, and Lepidoptera (Schuh and Slater 1995; Grimaldi and Engel 2005; Cameron et al. 2006), and the most diversified group of non-endopterygote insects (Kristensen 1991), with approximately 82.000 described species (Arnett 2000). Their feeding habits range from phytophagy to predation, including ectoparasitism and hematophagy. Many of them are important pest species to cultivated crops and some are important vectors of human diseases.

This review deals with the phylogenetic systematic hypotheses for Hemiptera. Throughout the text, common phylogenetic terminology is used in cladistics that the nonfamiliarized reader may find confusing. The reader shall refer to general references for an explanation of those terms and the logic behind it (e.g., Hennig 1966; Kitching et al. 1998; Schuh 2000; Albert 2005).

The number of described species within Hemiptera is not exactly known. It is considered to have roughly 82.000 described species (Slater 1982; Hodkinson and Casson 1991; Arnett 2000), a number that must be seen as conservative. For instance, an estimate of the total world fauna using Asian tropical rain forest samples lies between 184.000 and 193.000 species (Hodkinson and Casson 1991). In some diversified groups, as Cicadellidae, the proportion of undescribed tropical taxa may be as high as 90% (Dietrich and Wallner 2002). Of the total described species, merely three families account for most of the diversity: Cicadellidae, with approximately 22.000 species (Auchenorrhyncha) (Metcalf 1962a, 1962b, 1962c, 1962d, 1963a, 1963b, 1963c, 1963d, 1964a, 1965a, 1965b, 1966a, 1966b, 1966c, 1966d, 1967, 1968; Oman et al. 1990; McKamey 2001), and Miridae and Reduviidae (Heteroptera), with more than 10.000 and 6.000, respectively (Schuh 1995; Maldonado 1990).

Characters defining Hemiptera

Hemiptera has long been recognized as a monophyletic group (Hennig 1969; Carver et al. 1991). It can be recognized by the particular structure of the mouthparts: the mandibles and maxillary laciniae are modified into concentric stylets, the mandibular enclosing the maxillary ones, both forming the food and salivary channels; the multisegmented sheetlike labium is covering the mandibular and maxillary stylets; and the maxillary and labial palpi are always absent (Weber 1930; Hennig 1969, 1981; Cobben 1978; Kristensen 1991).

Paraphyly of Homoptera

Morphological evidence, nonetheless, pointed out that Homoptera was probably paraphyletic (e.g., Goodchild 1966; Schlee 1969d; Bourgoin 1986a, 1986b, 1993; Sweet 1996), or at least that evidence of its monophyly was not documented (Schlee 1969d). Some of the alleged characters supporting "Homoptera" mentioned were: enlarged foramen in the head, large sutures defining the mandibular plate, forewing larger than hindwing, reduced tarsomeres, and simple sperm (Bou- dreaux 1979; Hamilton 1981). Nonetheless, some of these characters are not synapomorphies (e.g., forewing –hindwing character), or may be based on reductions, which are difficult to homologize (e.g., number of tarsomeres).

Hennig (1969, 1981) doubted the monophyly of Homoptera, stating that the characters used for distinguish it from Heteroptera were symplesiomorphies. He recognized three groups within Hemiptera: Sternorrhyncha, Auchenorrhyncha, and Heteropterodea (as "Heteropteroidea"), the latter clade formed by Coleorrhyncha + Heteroptera (Schlee 1969d). The paraphyly of Homoptera was further corroborated with 18S rDNA sequence analyses (Wheeler et al. 1993; Campbell et al. 1994, 1995; Sorensen et al. 1995; von Dohlen and Moran 1995).

Even though evidence is compelling towards a paraphyletic "Homoptera", and that this has been adequately communicated to a more general audience (e.g., Carver et al. 1991; Kristensen 1991; Gullan 2001; Fagua 2005), it is still frequent to see references to "Homoptera" in areas such as Integrated Pest Management (e.g., Pedigo 1996), or in general entomological textbooks (e.g., Arnett 2000), practice that should be avoided.

Phylogenetic hypotheses

1. Paraneoptera

Paraneoptera is the group of Neoptera that includes Thysanoptera, Psocodea ("Phthiraptera" + "Psocoptera" [Lyal 1985; Yoshizawa and Johnson 2006]), and Hemiptera (Kristensen 1991; Wheeler et al. 2001). Kristensen (1991) listed for this clade the following putative synapomorphies: maxillary lacinia elongate and slender, detached from stipes; postclypeus enlarged; reduced number of tarsomeres (three or less); cerci absent; at most six Malpighian tubules; abdominal ganglia fused in one mass; gonangulum in the females fused with tergum nine (Scudder 1961); and sperm acrosome without perforatorium (Jamieson et al. 1999). Yoshizawa and Saigusa (2001) proposed nine additional synapomorphies from the front wing axillary sclerites for the group.

Some authors have included Zoraptera in Paraneoptera based on morphological characters (Hennig 1969, 1981; Kristensen 1981; Wheeler et al. 2001; Beutel and Weide 2005), in which the inclusive group of Psocodea + Thysanoptera + Hemiptera is denominated Acercaria, due to the absence of cerci (Börner 1904). The inclusion of Zoraptera in Paraneoptera, nonetheless, is highly controversial (Hennig 1969, 1981; Yoshizawa 2007), and has not always been followed (e.g., Kristensen 1991; Grimaldi and Engel 2005). The putative synapomorphies of Zoraptera + Acercaria are all reductions (number of Malpighian tubules, number of tarsomeres, and fusion of abdominal ganglia) (Hennig 1969, 1981; Kristensen 1981; but see Hünefeld 2007), and thus, difficult to homologize.

Potential synapomorphies have been proposed for Zoraptera + Acercaria based on head capsule characters (Beutel and Weide 2005), and on genitalic characters both for Zoraptera + Psocoptera, and for Zoraptera + Hemiptera (Hünefeld 2007), but have not been tested in a phylogenetic analysis. Nonetheless, some authors had rejected the inclusion of Zoraptera in Paraneoptera, based on phylogenetic analyses of 18S rDNA sequences, which place Zoraptera closely related to Dictyoptera (Wheeler et al. 2001; Yoshizawa and Johnson 2005), and on wing base morphology (Yoshizawa 2007). These evidences support the hypothesis that Zoraptera is not a member of the Paraneoptera.

Relationships among members of Paraneoptera (in the restricted sense of Kristensen 1991) are still unclear. Some phylogenetic hypotheses show a polytomy among Psocodea, Thysanoptera, and Hemiptera (e.g., Kristensen 1991; Beutel and Gorb 2001). In other instances Hemiptera is the sister group of Thysanoptera + Psocodea (e.g., Whiting et al. 1997; Gorb and Beutel 2001; Wheeler et al. 2001; Willmann 2004), or the sister group of Thysanoptera (e.g., Hennig 1969, 1981; Kristensen 1981; Whiting et al. 1997; Wheeler et al. 2001; Yoshizawa and Saigusa 2001; Cranston and Gullan 2003). The latter clade, Hemiptera + Thysanoptera, is named Condylognatha (Börner 1904), and is supported by a few synapomorphies (Kristensen 1981), namely the modification of the mandibles and maxillary laciniae into stylets (Hennig 1969, 1981; Heming 1980), specialized sclerotized rings between antennal flagellomeres (Seeger 1975), and the particular conformation of the axillary sclerites of the forewing (Yoshizawa and Saigusa 2001). The character of the flagellomeres rings should be further evaluated since it was found only in Thysanoptera and in Pentatomidae (Heteroptera) (Seeger 1975).

In those phylogenetic analyses of Hexapoda including paraneopteran representatives, usually the taxon sample is small (e.g., Whiting et al. 1997) or some groups are missing completely (e.g., Thysanoptera: Kjer 2004), rendering the conclusions of paraneopteran relationships inadequate. When representatives of all the three major clades are included, the results are either an unresolved polytomy (Kjer et al. 2006) or unlikely poly- or paraphyletic groups (e.g., Hemiptera paraphyletic respect to Thysanoptera: Whiting et al. 1997; Paraneoptera and Hemiptera polyphyletic: Wheeler et al. 2001; Paraneoptera paraphyletic with Psocodea as basal-most Neoptera and Polyneoptera as sister group of Heteropterodea + Auchenorrhyncha: Whiting 2002; Psocodea paraphyletic respect to Thysanoptera: Kjer et al. 2006). The relationships among members of Paraneoptera are an open field of research.

2. Hemiptera Hemiptera is an ancient lineage with fossils known since the Early Permian (Kukalová-Peck 1991; Shcherbakov and Popov 2002). Hemiptera is a monophyletic group, based on the unique structure of the mouthparts (see above), which consist of a labium enclosing the maxillary and mandibulary stylets (Kristensen 1991). Yoshizawa and Saigusa (2001) proposed another potential synapomorphy, the fork of the anterior axillary fold-line of the forewing, a character that is independent from that one of the mouthparts, which reinforces the monophyly of Hemiptera.

As discussed above, Hemiptera was traditionally divided in two groups: "Homoptera" and Heteroptera. Compelling evidence, nonetheless, suggest that Hemiptera is composed of three main clades: Sternorrhyncha, Auchenorrhyncha, and Heteropterodea, in which Sternorrhyncha is the sister group of Auchenorrhyncha + Heteropterodea (e.g., Hennig 1969, 1981; Kristensen 1975; Wootton and Betts 1986; Carver et al. 1991; Wheeler et al. 1993). Of these, only the monophyly of Auchenorrhyncha is still debatable (see below). Zrzavý (1990) termed the clade Auchenorrhyncha + Heteropterodea as Euhemiptera.

Shcherbakov and Popov (2002) proposed an alternative classification for Hemiptera, mostly following Börner (1904), in which they recognized five suborders and an "ancestral" group, the Archescytinoidea. In their view, the Auchenorrhyncha (as Cicadina) is paraphyletic, and the extant hemipterans that represent the sister group of the Heteropterodea (as Peloridiina + Cimicina) are the Cicadomorpha. They presented a phylogram depicting relationships among these groups (Shcherbakov and Popov 2002: see their fig. 179), which despite a list of characters supposedly supporting the nodes, can hardly be viewed as a phylogenetic hypothesis. However, Bourgoin and Campbell (2002), based on unpublished molecular data sets (18S rDNA), also argue that Auchenorrhyncha is paraphyletic, and that the sister group of Heteropterodea is Cicadomorpha, similar to the proposal of Shcherbakov and Popov (2002).

Yang (2004) proposed a scheme of relationships within Hemiptera based on morphological characters that differ from that presented by Wheeler et al. (1993), mainly in that Psylloidea is the sister group of Heteropterodea, Sternorrhyncha and Cicadomorpha are paraphyletic, and in the internal relationships of the Heteroptera infraorders (Leptopodomorpha, Cimicomorpha, and Pentatomomorpha are in a polytomy, Enicocephalomorpha is the sister group of this clade, and the remaining infraorders forms a basal group in an unresolved polytomy). Nonetheless, several flaws in the analysis, such as choosing a priori a "functional" outgroup (within Hemiptera) and not stating how the analysis was done, prevent his discussion of relationships being useful.

Sorensen et al. (1995) proposed unnecessary new names for clades already named (e.g., in Wheeler et al. 1993), in part, because Auchenorrhyncha was found to be paraphyletic respect to Heteroptera, and also because they wanted to recognize what they considered suborders of Hemiptera. They proposed: Clypeorrhyncha (for extant Cicadomorpha), Archaeorrhyncha (for Fulgoromorpha), Peloridiomorpha (for Coleorrhyncha [Peloridiidae]), Neohemiptera (for Fulgoromorpha + Heteropterodea), and Prosorrhyncha (for Heteropterodea) (Table 1). Nonetheless, these names may lead to confusion, in particular if Auchenorrhyncha is not paraphyletic (see below for discussion). Even more, the names Cicadomorpha, Fulgoromorpha, and Heteropterodea, independently of the outcome of the phylogenetic relationships within Hemiptera, convey a clear meaning already used in Hemiptera literature. Bourgoin and Campbell (2002) argue in favor of dropping the names proposed by Sorensen et al. (1995). Because the ICZN (1999) do not regulate the application of names above family rank, it depends on the researchers to favor the use of certain names. Vibrational communication is diverse in Hemiptera, with species producing sound in terrestrial as well as aquatic ecosystems, being substrate communication common (Crocroft and Rodríguez 2005). There may be stridulation, tymbal, and percussional vibration (Claridge 1985; Hoy and Robert 1996; Gogala 2006; Hoch et al. 2006; Soulier-Perkins et al. 2007). It has been suggested that a tymbal-like communication is synapomorphic for Auchenorrhyncha + Coleorrhyncha + Heteroptera (Hoch et al. 2006).

3. Sternorrhyncha

Sternorrhyncha is a monophyletic group (Schlee 1969c; von Dohlen and Moran 1995), which includes four main groups: Psylloidea, Aleyrodoidea, Aphidoidea, and Coccoidea (e.g., Schlee 1969b; Carver et al. 1991; von Dohlen and Moran 1995; Bourgoin and Campbell 2002).

Goodchild (1966) based on the alimentary tract morphology regarded Aphidoidea as the sister group of Coccoidea + (Psylloidea + Aleyrodoidea). Schlee (1969a, 1969b) based on external morphology and male genitalic characters considered two sister groups within Sternorrhyncha: Psylloidea + Aleyrodoidea (Psylliformes), and Aphidoidea + Coccoidea (Aphidiformes). In some analyses Psylloidea is considered the sister group to the rest of Sternorrhyncha (e.g. Campbell et al. 1994, 1995). Despite these competing hypotheses, no modern comprehensive phylogenetic analysis is available for the group (but see review of previous phylogenetic hypotheses in Schlee 1969b).

3a. Aleyrodoidea. The Aleyrodoidea, or whiteflies (Figs. 1A, B, C), includes 1.556 valid species in Aleyrodidae, the only included family, which is subdivided into three subfamilies: Aleurodicinae, Aleyrodinae, and the controversial Udamoselinae (Mound and Halsey 1978; Martin and Mound 2007). Manzari and Quicke (2006) analyzed the phylogenetic relationships of the large and widespread Aleyrodinae, which contains most of the economically important pest species (Byrne and Bellows 1991; Martin and Mould 2007). Their analyses based on pupal case characters (Manzari and Quicke 2006) found that only 45% of the genera with multiple representatives were monophyletic, and that most of the tribes and economically important genera (e.g., Bemisia and Trialeurodes) are not monophyletic. They did not propose any new taxonomic scheme at the tribal level arguing that the data set used is not enough to provide adequate resolution.

3b. Aphidoidea. Aphidoidea has three families, Phylloxeridae, Adelgidae, and Aphididae, commonly called aphids (Figs. 1D, E), with some 4.500 described species (Remaudière and Remaudière 1997; Blackman and Eastop 2006). Some authors prefer to give them rank of superfamilies (e.g., Shcherbakov and Popov 2002).

Heie (1987) proposed a scheme of relationships for the Phylloxeridae and Aphididae based on morphology of extant and extinct taxa. Von Dohlen and Moran (2000) proposed a phylogenetic scheme for Aphididae using 12S and 16S mtDNA. They found little support and structure at deeper nodes, only recovering recognized tribes but not subfamilies. Martínez- Torres et al. (2001), also using a mitochondrial gene, found similar results. Ortiz-Rivas et al. (2004) analyzed the relationships of Aphididae based on a single nuclear gene (long wavelength opsin) and the topologies of their analyses were better resolved than in previous analysis. They identified three main clades, with Aphidinae and Lachninae always monophyletic. The relationships within Aphidinae were examined by von Dohlen et al. (2006) with three genes: leucine tRNA, COII mtDNA, and EF1a.

3c. Coccoidea. The group has about 7300 described species (Miller and Ben-Dov 2006), and the 20 or more families (Figs. 1F, G) are usually divided into two groups: Archaeococcoidea and Neococcoidea (Koteja 1974; Miller and Kosztarab 1979; Gullan and Kosztarab 1997). Gullan and Cook (2007) review the phylogeny and classification of the coccoids in detail. Miller (1984), Miller and Hodgson (1997), and Foldi (1997) proposed phylogenetic hypothesis based in morphological characters for the group. Nonetheless, their analyses had either a small taxon sampling or poor nodal support that renders the results non-conclusive.

Cook et al. (2002) assessed the phylogeny of the Coccoidea using rDNA sequences of the small subunit. They found that the neococcoids are a monophyletic group and that Pseudococcidae is the sister group to the rest of them. Eriococcidae was retrieved as a paraphyletic group in their analyses, corroborating previous views based on morphology (Cox and Williams 1987). The monophyly and relationships of the archaeococcoids are in doubt (Gullan and Cook 2007).

Two out of three of the largest families of Coccoidea have been assessed for phylogenetic relationships. Downie and Gullan (2004) assessed the phylogeny of the Pseudococcidae using DNA sequences of EF1a, 28S and 18S. They found that the three major clades of the analysis correspond to the subfamilies Pseudococcinae, Phenacoccinae and Rhizoecinae, with Sphaerococcinae as polyphyletic. Recently, Hardy et al. (2008) reassessed the relationships of the Pseudococcidae, using an expanded taxon sampling and adding morphological data. They recognized two main clades to which they apply the names Phenacoccinae and Pseudococcinae, including in the former the Rhizoecini.

Morse and Normark (2006) studied the phylogenetic relationships of Diaspididae using EF1a and 28S rDNA sequences. Most of the traditional groups recognized in classifications were broadly recovered, although none of them was strictly monophyletic. The remaining family, Coccidae, has never been subject of a comprehensive phylogenetic analysis. Qin and Gullan (1995) evaluated the relationships of one of the subfamilies, Ceroplastinae (Coccidae), finding that most of the genera were not monophyletic, and that most of the species should be grouped into the genus Ceroplastes.

3d. Psylloidea. Psylloidea (Fig. 1H) has more than 3000 described species (Hodkinson and Casson 1991; Hollis 2004; Burckhardt 2005). The classification of this group of sapsucking insects has been extremely artificial (Burckhardt and Lauterer 1989). White and Hodkinson (1985) first proposed a phylogenetic scheme of the Psylloidea using mostly nymphal characters. In their phylogenetic analysis they regard Psyllidae as the sister group of all other psylloids. Of the eight families recognized by them, two, Aphalaridae and Spodyliaspididae, were accepted in their classification as paraphyletic following an "evolutionary" approach (White and Hodkinson 1985: 264). Ouvrard (2002) reassessed the relationships of the Psylloidea using morphological characters of the adult thorax and 18S rRNA sequences. His results differ from those of White and Hodkinson (1985), although it is difficult to compare and interpret them because Ouvrard did not present a list of characters, matrix, or list of species used in his analysis. Nonetheless, he concluded that Aphalaridae and Psyllidae are paraphyletic.

Recently, Ouvrard et al. (2008) identified potential morphological synapomorphies for Psylloidea in relation to the wing base structure: absence of subalare, a median notal wing process, and of the anterior arm of the third axillary sclerite; no articulation with the second axillary sclerite; a weakly-sclerotized third axillary sclerite; and presence of a two-horned basalare. Hodkinson (1989) provided a review of the faunal elements present in the Neotropical region.

Much of the recent evidence for the paraphyly of Auchenorrhyncha come from molecular sets alone. Campbell et al. (1995) using 18S rDNA sequences found a polytomy among Fulgoromorpha, Cicadomorpha, and Heteropterodea when using the full dataset (their fig. 3). When using a restricted data set (homoplasious sites removed), either Fulgoromorpha or Cicadomorpha become sister groups of Heteropterodea (Campbell et al. 1995, their fig. 4), but when Peloridiidae was removed from this restricted data set, Fulgoromorpha become the sister group of Heteroptera. Sorensen et al. (1995) also using 18S rDNA found that Auchenorrhyncha is not monophyletic. Despite their small dataset, they found that Auchenorrhyncha is either paraphyletic or polyphyletic respect to Heteroptera. Bourgoin et al. (1997) in an analysis of the Fulgoromorpha found that this group is the sister group to Heteroptera, rendering Auchenorrhyncha paraphyletic. They also found that Cicadomorpha was not monophyletic if Psyllidae was included as part of the outgroup, showing that their analysis is sensitive to taxon sampling. Ouvrard et al. (2000) found a polytomy among Fulgoromorpha, Cicadomorpha, and Heteropterodea, similar to the results of Campbell et al. (1995). Urban and Cryan (2007) using an extended data set of four genes and 83 species of Fulgoromorpha found a monophyletic Auchenorrhyncha, which was the sister group of Heteroptera.

4a. Cicadomorpha. Within this group three superfamilies are recognized: Cercopoidea (spittlebugs or froghoppers) (Figs.1I, J), Cicadoidea (Fig. 1K), and Membracoidea (leafhoppers and treehoppers) 2005 229 2005 22 1 1 231-283 1982 3 1-455 1995 21 93-123 1998 50 1-152 1999 63 5-18 2000 142 185-194 1994 8 8 861-909 1999 13 13 309-350 2004 14 1-344 2004 18 18 467-522 2000 1.003 1987 47-53 1995 222 1996 361 1999 576 2001 346 2006 550 1993 536 1994 868 2005 400 2006 368 2003 1111 2001 39 4 4 177-207 2005 124 121-136 2006 two 1439 1904 27 511-533 1981 928 1970 404 1979 320 1986 15 4 4 237-252. 1986 22 4 4 413- 422 1993 29 3 3 225-244 2007 2002 4 67-82. 1990 . 26 4 4 555-564. 1997 13 207-224 1954 108 1-917. 2005 2 143-157817 1991 19 19 71-75 2002 75 1-2 1-2 57-59 1989 23 643-712. 1991 36 431-457 2006 64 1 1 27-34 1994 3 2 2 73-88 1995 20 3 3 175-194 1962 18 1 1 1-16 1971 7 4 4 737-770 2005 271 CSIRO^dDivision of Entomology 1991 2 2 429-5091137 2004 1995 27.3A 506 2002 27. 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