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Phylogeny of the Dictyoptera

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Phylogeny of the Dictyoptera Systematic Entomology (1992) 17,253-268 Phylogeny of the Dictyoptera 1 2 BARBARA L. THORNE ,2 and lAMES M. CARPENTER ,* IDepartment of Biology, Northeastern University, Boston, MA 02115, and 2Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138, U.S.A. Abstract. Relationships among six key dictyopteran taxa (Mantodea; Blat- todea (excluding Cryptocercidae); Cryptocercidae; Mastotermes darwiniensis, Termopsidae and Kalotermitidae [Isoptera]) are analysed based on seventy morphological, developmental and behavioural characters. The fossil record and the 'living fossil' genera Cryptocercus, Mastotermes and Archotermopsis are discussed in detail. Exact analysis of the character state matrix by implicit enu- meration (Hennig86) resulted in one cladogram, strongly supporting Blattodea + Cryptocercidae as a sister group to Mantodea, with the Isoptera as a sister group to that complex. Arrangements within the termites are equivocal, with Termopsidae and Mastotermes darwiniensis possible as the relatively most pri- mitive element of Isoptera. Introduction In 1937 (and further in 1938), another Russian paleoen- tomologist, A. V. Martynov proposed a phylogeny and Historical background evolutionary time frame (1937: fig. 21) in which roaches The phylogeny of the dictyopteran complex (mantids, and termites diverged in the late Devonian or early Car- cockroaches, termites) remains unresolved.· There is a boniferous, and mantids radiated from roaches in the general consensus that these three taxa represent a mono- mid-Carboniferous. Martynov suggested that the solitary phyletic grouping, but relationships among these 'orders', ancestor of the Isoptera (order 'Archisoptera') of the and how they fit in with other orthopteroids, are disputed. Devonian was the stem of all modern and extinct dictyop- Derived characters linking these include, for example, teroids (and also of all holometabolous orders). Martynov a perforated corporo tentorium (Kristensen, 1975; see obviously did not structure his arguments in the context also list in Boudreaux, 1979: 277). Boudreaux (1979) of modern systematic hypotheses, but some of his inter- considered the Blattaria and Mantodea to be sister groups, pretations are intriguingly compelling. For example, and the Isoptera to be the sister group of that complex. Martynov argued that homonomous wings were very Hennig (1981) combined the roaches and tennites into a likely the ancestral condition in winged insects. The order single order ('Blattodea') which he considered as the sister Isoptera was named as such based upon similarity in size, group to the mantids. Kristensen (1975) endorsed the shape and venation of the fore- and hindwings. Martynov same relationships as Hennig, but upon reevaluation emphasized the fact that termites have 'retained' extra Kristensen (1981) grouped all three taxa into the same basal branches of the radius (which he tenned 'preradius', order (Dictyoptera) and encouraged further work to re- synonymous with first and second radius (Comstock, solve the phylogeny. In a volume edited by Rohdendorf 1918; Tillyard, 1926; Emerson, 1933, 1965), radius (Smart, & Rasnitsyn (1980), paleoentomologists at the Soviet 1951, 1953), first radius and radial sector (Hennig, 1981» Paleontological Institute published their consensus opinion in both forewings and hindwings. The 'preradius' veins regarding the evolution of insect orders. They felt certain are absent in the forewings (present in the hindwings) of enough to propose relationships among virtually all modern roaches and mantids. Martynov suggested that if tennites and extinct groups, but the Dictyoptera were left dangling diverged from roaches, it is difficult to envision a reversal amidst the orthopteroids: unrooted and with question of the wing characters back to such striking homonomy. marks. Hennig (1981: 197) felt that Isoptera wing homonomy is secondary. ,. Present address: Department of Entomology, American Similarly, Martynov interpreted tegmina (found in Museum of Natural History, Central Park West at 79th Street, roaches and mantids) as a derived character state. He New York NY, 10024, U.S.A. argued that since tennites have 'retained' membranous Correspondence: Dr Barbara L. Thome, Department of En- forewings, one must again hypothesize that tennites, tomology, University of Maryland, College Park, MD 20742- or solitary proto-tennites, diverged from the basal stem 5575, U.S.A. of the modern 'roach' taxa rather than radiating from 253 254 Barbara L. Thorne and lames M. Carpenter evolved blattids with tegmina and non-homonomous (Walker, 1919, 1922; Marks & Lawson, 1962; McKittrick, wings. Tegmina have been considered an ancestral state 1964, 1965). These structures manipulate the eggs and in the Dictyoptera by Hennig (1981: 198), and interpreted position them for the ootheca (reviewed in Marks & as a derived character state by Holmgren (1911) and Lawson, 1962; McKittrick, 1964). Boudreaux (1979: 220). Many ofthe long blattid ovipositors are so well preserved Martynov's ideas have received little attention, particu- in fossils that Vishniakova (1968) was able to determine larly from American orthopteroid systematists. Grasse that they are derived from the eighth and ninth abdominal (1986: 449-459), however, discussed Martynov's hy- sterna. This concerned her because the internal valves or potheses and found them quite plausible. ovipositors of modern roaches are associated with the seventh sternum (Walker, 1919, 1922; Roonwal, 1956; Marks & Lawson, 1962; McKittrick, 1964, 1965). (A The fossil record feature common to all the Dictyoptera is that in females the seventh abdominal sternum is extended posteriorly There are four reports of fossil termites from the Cre- to form a bursa or vestibulum.) Vishniakova (1968) ques- taceous, all determined as members ofthe Hodotermitidae tioned the homology of the external ovipositors of fossil (Emerson, 1967; Jarzembowski, 1981; Lacasa-Ruiz & roaches with the reduced internal ovipositor valves of Martinez-Delclos, 1986; Krishna, 1990). The oldest, modern cockroaches. Hennig (1981) was also worried Meiatermes bertrani, was found in a limestone deposit about including the fossil 'blattid-like' insects with long in Spain dating approximately 130 million years (Lacasa- ovipositors in the same taxon with modern roaches that Ruiz & Martinez-Delclos, 1986): all produce oothecae (excepting a few, derived groups; Jarzembowski (1981) described the termite Valditermes Roth & Willis, 1958). brennae from an early Cretaceous deposit in England The issue can be resolved based on the work of Nel (120 million years). Based on the forewing venation, (1929). He investigated the ontogeny of the genitalia in--- Valditermes was originally described as a member of the Orthoptera by examining serial sections ofall developmen- Termopsidae (a family which was formerly considered tal stages from embryos to adults. In studies of successive a subfamily of the Hodotermitidae; see Grasse, 1949, instars of Blattella nymphs he showed how the eighth and but also see Emerson (1965: 16) for arguments to retain later the ninth sterna completely telescope within the the original classification). Examination of additional seventh sternum which, simultaneously, gradually enlarges specimens revealed a possible anal expansion in the hind- and becomes specialized as the subgenital plate. Further, wing (Jarzembowski, 1984), characteristic of the family Nel demonstrated in nymphs from the first through third Mastotermitidae. Jarzembowksi felt that wing venation instars that the lateral and inner ovipositor lobes differen- characters still warrant placement of Valditermes in the tiate from the ninth sternum, and that the anterior lobes family Termopsidae, but shape of the hindwing is likely to (or ventral valves) are derived from the eighth sternum. stimulate a careful re-evaluation by termite taxonomists Thus, even though the three pairs of internal ovipositor (also suggested by Krishna, 1990). Aside from these valves appear to be associated with the seventh abdominal ambiguous specimens, the Mastotermitidae, which are sternum in modern roaches (because that is the only one considered by many to be the most 'primitive' termites, do that is visible), their embryological origin is from the eighth not appear in the fossil record until the Eocene (60mya) and ninth sterna, just as was observed in fossil roaches (reviewed in Burnham, 1978). by Vishniakova. It is therefore reasonable to interpret Fossil roaches, or at least 'roachoid' insects, have re- the external ovipositors of fossil blattids as homologous ceived a reasonable amount of attention because of one with the internal valves or ovipositors of modern forms. rather dramatic character. Virtually every blattoid insect These internal valves are visible, albeit highly reduced, found in fossil strata from Paleozoic times had a long in several termite families (Browman, 1935; Geyer, 1951; external projection that looks like a functional ovipositor Weesner, 1969). Of all termites, they remain most devel- (Martynov, 1937, 1938; Laurentiaux, 1951; Sharov, 1966; oped in Mastotermes darwiniensis (Crampton, 1920, 1923; Marks & Lawson, 1962; Vishniakova, 1968, 1971). So Browman, 1935; Roonwal, 1956; Geyer, 1951). many fossils have been found with the ovipositor and so few without that it is generally interpreted that the few without are males (F. M. Carpenter, pers. comm.). Based on specimens examined to date, the following pattern in ovipositor size appears over geologic time. In the
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