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Home , Blattodea, Blattoidea, Dictyoptera, Dictyoptera (beetle), Mastotermes, Mastotermes darwiniensis

Systematic (1992) 17,253-268

Phylogeny of the

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; darwiniensis, and [Isoptera]) are analysed based on seventy morphological, developmental and behavioural characters. The record and the 'living fossil' genera , 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 + Cryptocercidae as a sister group to Mantodea, with the Isoptera as a sister group to that complex. Arrangements within the are equivocal, with Termopsidae and 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 or early Car- , termites) remains unresolved.· There is a boniferous, and mantids radiated from roaches in the general consensus that these three taxa represent a mono- mid-. 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 . 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 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 of 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 (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 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 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 with modern roaches that Ruiz & Martinez-Delclos, 1986): all produce oothecae (excepting a few, derived groups; Jarzembowski (1981) described the Valditermes Roth & Willis, 1958). brennae from an early 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 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 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 . 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 Car- 'Living fossils': Cryptocercus and Mastotermes boniferous and , all of the external ovipositors were long (Carpenter & Burnham, 1985). In the , Among extant forms, the three of the a variety of lengths were found (Vishniakova, 1968, 1971). family Cryptocercidae and the termite Mastotermes dar- By the Cretaceous, all ovipositors were quite short or winiensis Froggatt are considered to be relicts or 'living absent (F. M. Carpenter, pers. comm.). No modern roach, fossils'. Each of these groups is enshrined in the concept that or termite, has an external ovipositor, but mantids do as modern insects they have large numbers of uniquely pri- have external valves. Modern roaches have three pairs of mitive characters. Here we examine those interpretations, internal valves that are frequently called internal ovipositors and re-evaluate selected characters. Phylogeny of the Dictyoptera 255

Cryptocercus (also questioned by Emerson, 1935; Boudreaux, 1979). One inconsistency is that if there was a common ancestor Much of the debate on the phylogeny of the Dictyoptera with the symbionts, then that suggests that there was a involves Cryptocercus and its affinity to the termites. A secondary loss of the gut fauna, and coincident change in common notion is that advocated by Cleveland (1934: diet and habits, in all other roaches (Boudreaux, 1979). 332), ' ... the evidence is overwhelming that Oyptocercus Alternatively, one could hypothesize that is either the ancestor of termites or is closely is more closely related to the termites than to the rest of related to the ancestor, which is extinct.' He based that the roaches, but that is difficult to support based on any conclusion on similarities of morphology, habitat and character except the gut fauna. Suggestions are occasionally gut symbionts, and it has been adopted in general dis- made that either Cryptocercus or termites could have cussions of termite origins ever since (e.g. McKittrick, acquired the gut fauna from the other (Emerson, 1935; 1964; Krishna, 1970; Wilson, 1971; Burnham, 1978; Boudreaux, 1979; Hennig, 1981), but that hypothesis has Seelinger & Seelinger, 1983; Nalepa, 1984). never been accepted because ofdifferences in sexuality and The key question is whether the Cryptocercidae are physiology between Protozoa found in Cryptocercus and in a sister group to the termites (reviewed in Boudreaux, termites (Honigberg, 1970; Bobyleva, 1975). Differences 1979; Hennig, 1981), or whether they actually share more among Protozoa within the two hosts are unsurprising: derived characters with the modern roaches (discussed in there has probably been at least 100 million years of vir- Hennig, 1981; Seelinger & Seelinger, 1983). tually independent evolution since both of these lineages Many investigators have studied Cryptocercus anat- first acquired gut symbionts, and numerous specializations omy and morphology (e.g. Judd, 1948; Nutting, 1951; or physiological adaptations could have occurred since Marks & Lawson, 1962; McKittrick, 1964, 1965; Brossut, divergence. 1973; Roth & Alsop, 1978; Deleporte et al., 1988), gut Successful Protozoa transfaunation between Cryptocercus symbionts (e.g. Cleveland, 1949; Cleveland et aI., 1934; and (Termopsidae) has been repeatedly Nutting, 1956; Bobyleva, 1975), and social behaviour demonstrated under experimental conditions in which (Cleveland et aI., 1934; Seelinger & Seelinger, 1983; insects have been defaunated and received Protozoa from Nalepa, 1984; 1988a, b), often comparing those characters the alternate host either by feeding on recently removed to structures, Protozoa or behavioural patterns found hindguts (Cleveland, 1934) or by innoculation per rectum in termites. There is general agreement that the family of hindgut material (Nutting, 1956). Recent work (Thorne, Cryptocercidae is primitive relative to other living roaches 1990) has demonstrated that aggression and rapid con- (Princis, 1960; McKittrick, 1964, 1965; Roth, 1968; tIuber, sumption behaviours exist in these two insect groups. 1974; Boudreaux, 1979; Hennig, 1981). However, the Modern Cryptocercus and Zootermopsis live sympatrically, phylogenetic relationship between the Cryptocercidae and within the same decomposing logs in parts of California, the Blattodea remains uncertain because the character Oregon and Washington in the United States. It is likely systems have not been evaluated in light of modern sys- that their ranges, or those of their ancestors, overlapped tematic principles and techniques, and our analysis revealed more extensively in the past. It is thus probable that op- no synapomorphies for Blattodea minus Cryptocercidae. portunities for transfer of hindgut protozoa via aggression and consumption have occurred over evolutionary time. This hypothesis for the presence of similar cellulolytic Protozoan gut symbionts: a homologous character shared protozoa in both modern lineages (Thorne, 1990, 1991) by Isoptera and cryptocercids? must be considered as a valid alternative to the established theory that the symbionts were passed to Cryptocercus and Cryptocercidae is one of two groups of roaches that to termites through a common ancestor. It seems plausible feeds on wood: the other is the Panesthiinae. The Panesthi- to interpret the shared habitat and nutritional physiology inae have bacterial gut symbionts to aid in digestion of of Cryptocercus and the Isoptera as a result of convergence cellulose. The Cryptocercidae have protozoan symbionts and transfaunation (but see Nalepa, 1991). which are quite similar to those found in some termites. Presence of such similar gut fauna has frequently been cited as a character linking Cryptocercus'with the termites. Cryptocercus characters which are derived in comparison This contention must be reconsidered. to the Isoptera It appears certain that some of the intestinal Protozoa found in the lower termites and in Cryptocercus are derived While Cryptocercus is clearly primitive with respect to from a common ancestral flagellate. The protozoa are many of the other living roaches, it has many character morphologically very similar (reviewed in Bobyleva, 1975), states which are derived in comparison to the Isoptera and immunological characterizations of the Protozoa (Table 1). suggest that the gut fauna of the two hosts are related (Ablin, 1965; Ablin & Ritter, 1967). It is not clear, however, that these flagellates were Mastotennes darwiniensis transferred to Cryptocercus and to modern termites by a common ancestral host, which is the general conception Egg deposition: the 'ootheca'. Unlike all other termites, 256 Barbara L. Thorne and lames M. Carpenter

Table 1.

Derived characters unique to Cryptocercus Apterous cl': seventh sternum expanded cl' + ｾＺ seventh tergum expanded Loss of mandibular and hypopharyngeal cephalic glands First pair of ovipositor valves with small moveable spines Derived characters found in Crytocercus and other primitive cockroaches Styli absent in ｾ ｾ Presence of inferior hypostomal glands Spermatheca opens through a pore midline of eighth sternum Proventriculus: primary + secondary denticles conspicuously dissimilar Proventriculus: highly modified interdental areas (intercalary plate) Proventriculus bilateral symmetry Paraprocts V-shaped in crossections (fiat in Isoptera) Sex determination XXIXO

Mastotermes darwiniensis does not lay its eggs singly. Methods Instead, eggs are deposited in an organized mass or pod (Hill, 1925). This egg pod is often called an 'ootheca', but Largely based on an extensive literature review we com- we concur with others (Gay, 1970; Watson et al., 1977; piled data on six taxa key in the phylogeny of the termites Grasse, 1986) in thinking that that is an inappropriate and the larger dictyopteran complex. Groups included in term. It is certainly not an ootheca of the type produced the analysis were: by the mantids and roaches. The Mastotermes egg pod, (1) Mastotermes darwiniensis - the only living member which consists of two neat rows, is covered by a very thin of the Mastotermitidae, which is generally regarded as film (Hill, 1925; McKittrick, 1964). The roach-mantid a primitive family. This family is composed of at least ootheca has a discrete outer shell which results from the thirteen fossil species from four genera which had a broad tanning of protein and organic calcium salts secreted by geographic distribution during the Tertiary (Emerson, the colleterial glands after deposition of the eggs (reviewed 1965). in McKittrick, 1964; Boudreaux, 1979). In roaches there (2) Termopsidae and (3) Kalotermitidae - characters is a distinct keel on the hard ootheca shell, and in mantids from both fossil and living members of these termite oothecae come in a variety of shapes. The form of the families were evaluated. Both of these families are thought Mastotermes egg pod is really more similar to that of to have originated reasonably near the stem of the Isoptera acridids, and may well be primitive relative to the ootheca (reviewed in Emerson & Knshna, 1975). of roaches and mantids (Boudreaux, 1979). (4) Cryptocercidae - the roach family whose sole Mastotermes and Archotermopsis. The Mastotermitidae is Cryptocercus. There are three living species of is defined by ancestral characters rather than by uniquely Cryptocercus. derived characters. This has led to a lot of confusion and (5) Blattodea (excluding Cryptocercidae) and (6) Man- potential misinterpretation. While the sole surviving species, todea - dictyopteran orders. We tried to determine likely Mastotermes darwiniensis, has retained a number of con- character states of the ground plan in each order, basing spicuous characters which are probably ancestral (egg pod, judgements on the fossil record and present understandings enlarged pronotum, expanded hindwing jugal lobe), it of phylogenies (for roaches: Princis, 1960; McKittrick, also has a large number of derived features (Table 2). 1964; Roth, 1968; Huber, 1974), which are basically non- Another living termite with a very primitive morphology existent for the mantids (but see Beier, 1968). We could and social system is the Himalayan termite Archotermopsis not determine whether Blattodea in this sense is mono- wroughtoni, a member of the Termopsidae. phyletic; that question is beyond the scope of this analysis. Space limitations prohibit a complete discussion of this The characters adduced in this analysis are listed below, topic, but conspicuous derived character states for these along with the codes assigned to the character states. two taxa are listed in Table 2. (Characters omitted from the analysis due to ambiguity Mastotermes darwiniensis thus fits a common evolution- of state, non-independence or lack of variance within the ary pattern. It is primitive in some respects but it also has considered taxa are listed in Appendix 1.) The character a large number of derived characters. Thus this species state matrix for the taxa is shown in Table 3. Character state cannot be viewed as a clear window into the ancestry of definitions and polarities largely followed prevailing the- the Isoptera. ories of evolutionary change within the complex variously termed (Martynov, 1925), Paurometabola Phylogeny ofthe Dictyoptera 257

Table 2.

Mastotermes darwiniensis Archotermopsis wroughtoni

Imago characters Fused first and second teeth, left mandible Ocelli absent Forewing basal suture well defined Pronotum not expanded Fore- and hindwing scales long No expanded hindwing jugal lobe Pimpule wing micro-sculpturing Eggs laid singly Ninth sternum divided en Bacteroids absent Three sterna! glands Fewer ! segments, imago Sperm multiflagellate Fewer ma!pighian tubules, imago Styles on 00 only Reduced development of ovipositor valves Reduced cerci Eighth sternum divided (9) Ninth paratergites fused with tergum 9 ?Opening of spermatheca

.Non-imago characters Soldiers reduced no. antenna! articles Soldier cerci 5 articles Bladelike protruberance front coxa, soldier No reproductive soldiers Chemical (quinone) soldier defence Workers (as opposed to pseudergates) Hilly integrated termitophile Nest construction!covered foraging galleries Forage away from the nest Large colony population sizes (not included in character matrix)

(Hennig, 1981) or Lower (Kristensen, 1981). 2. NUMBER OF ANTENNAL ARTICLES (IMAGO), Polarity was conferred by the addition of a hypothetical [0] >35; [1] 30-31; [2] 23-27; [3] 11-21. ancestral taxon to the matrix, which was used to root the cladogram. This taxon had all-plesiomorph states, except 3. POSITION OF EYES. [0] no stereoscopic VlSlon; for characters 41 and 49, where we could not determine [1] eyes broadly separated and rounded, enabling stereo- the plesiomorphic condition. For these two characters the scopic vision. ancestor Was scored as ambiguous. Multistate characters were linearly ordered, except for characters 25, 28, 34, 4. OCELLI (IMAGO). [0] 3 (median ocellus present); 41, 45, 51, and 64, which were treated as nonadditive. [J.] 2; [2] none. Hennig (J.981: J.95) calls absence of the For these characters we regard the polarity as ambigu- median ocellus the only undoubted derived character ous, based on the information at hand. Cladistic analysis common to cockroaches and termites. Boudreaux (1979) (Hennig, 1966) was implemented with the Hennig86 says that this loss is common among insects and there- program (Farris, 1988). fore is suspect as a synapomorphy linking termites and cockroaches.

Character state definitions 5. LEFT MANDIBLE. [0] with 3 distinct marginal teeth; [J.] with fused first and second marginal teeth (i.e. with 2 1. ANTENNAE, [0] long and filiform; [1] short and marginal teeth now visible). (Emerson, 1962: 22; Grasse, moniliform. J.986: 506).

Table 3..Data matrix for Dictyoptera. A question mark indicates missing data.

Mantodea 001O? O??OO 01010 01?11 10011 10000 1?120 00000 1?002 ?OO?O 0?10? 00000 0?0?1 ?OOOO B1attodea 00010 00010 01000 01?10 10000 10001 1?11O 10100 11002 100?0 00100 00000 00011 00000 Cryptocercidae 00010 01110 01000 I???? ????? ????1 10111 10110 11002 100?1 0?100 00000 00021 00000 Mastotermitidae 11011 00011 10100 00100 10100 01210 01000 11101 00110 01100 21110 21222 20200 11222 Kalotermitidae 13011 00001 10101 00210 11102 02210 O?OOO 11103 2?10? 00110 11121 22132 11110 01111 Termopsidae 12020 10001 10100 00000 00100 02100 00000 11102 2?001 00110 01001 12111 10110 01111 258 Barbara L. Thorne and lames M. Carpenter

6. RIGHT MANDIBLE. [0] with no subsidiary tooth on Concurring with Hennig, Wootton (pers. comm.) doubts front base of first marginal tooth; [1] with subsidiary tooth if the wing homonomy of termites is plesiomorphic. He on front base of first marginal tooth. (Emerson, 1961: 123; reasons that membranous wings in termites are adaptations 1962: 24; Emerson & Krishna, 1975: 26). to slow, hovering, passive wind-dispersal modes of flight. Considering the variation of thickened forewings within 7. MANDIBULAR GLANDS. [OJ present; [IJ absent. the terminal taxa, possession of tegmina may not even be (Brossut, 1973; also discussed in Deleporte, 1988: 150-151). a groundplan trait of these groups (Kristensen, 1975: 12).

8. HYPOPHARYNGEAL GLANDS. [OJ present; [lJ 18. WING MICRO-SCULPTURING. [0] only papillae; absent. (Brossut, 1973, also discussed in Deleporte, 1988: [lJ papillae and pimpules; [2J papillae, pimpules and 150-151). tubercles. (Roonwal & Rathore, 1978; Roonwal et aI., 1979; Roonwal, 1981, 1985). 9. PRONOTUM. [OJ not expanded; [1] expanded to cover part or all of head. Based on studies of Problattaria and 19. FOREWING PRERADIUS. [0] present as two Archimylacridae, Tillyard (1937) reports that fossils in branches; [lJ absent. (Martynov, 1937, 1938; Grasse, which the pronotum has begun to expand forward to 1986: 449). overlap the head capsule were unknown before the Lower Permian. Thus the typical termite pronotum (unexpanded 20. FOREWING SUBCOSTA. [OJ short; [IJ long. (Mar- except in Mastotermes) may represent the ancestral state. tynov, 1937: 146). Boudreaux (1979: 217-218) states that The pronotum groundplan condition in mantids is this may be the only possible credible synapomorphy unclear. The expanded pronotum condition is apparently between roaches and termites, and that this too may be restricted to subordinate taxa in the higher mantid grades. a convergence. Similarly, prothorax elongation is characteristic only of higher-grade mantids (Kristensen, pers. comm.). 21. POSITION OF MEDIA. [OJ media closer to Cu; [lJ position of M ends close to tip of wing and tends to be 10. VENTROPLEURITE (pleurosternal region of the halfway between Rs and Cu, or closer to the Rs than to pterothorax). [0] present; [1] absent. (Matsuda, 1960: Cu in middle of wing. (Weesner, 1969: 38; Hamilton, 718,720). 1972: fig. 12; Emerson & Krishna, 1975: 25-26; Hennig, 1981: fig. 48). 11. AORTA/INCURRENT OSTIA/PERICARDIAL SEPTUM = ALARY MUSCLES (all related characters). 22. MEDIA BRANCHING. [0] highly branched; [IJ [OJ aorta begins at prothoracic incurrent ostia, 12 pairs simple. (Rehn, 1951: 20; Krishna, 1961; Boudreaux, incurrent ostia (3 thoracic, 9 abdominal), 12 pairs alary 1979: 220). muscles; [1] aorta arises from mesothoracic enlargements (prothoracic lost; meso- and meta- retained), 11 pairs 23. CLAVUS (ANAL AREA) OF FOREWING. [0] incurrent ostia (2 thoracic, 9 abdominal), 11 pairs alary large, with veins; [lJ reduced or suppressed. muscles. (Nutting, 1951). 24. FOREWING JUGAL LOBE. [OJ reduced; [1] slightly 12. SEGMENTAL ARTERIES = excurrent lateral enlarged, folded under wings when at rest. structures. [0] absent (no excurrent circulatory plan); [lJ present. Segmental arteries are absent in other hexapods. 25. HINDWING SUBCOSTA. [0] short; [1] long; [2J (Nutting, 1951; Boudreaux, 1979: 221; Kristensen, 1981). absent (Krishna, 1961: 315). Ordered nonadditively be- cause polarity is unclear. 13. GANGLIA. [0] first two abdominal ganglia not fused with metathoracic ganglion; [1] fused. (Boudreaux, 1979: 26. POSTCUBITUS OF HINDWING (PCu = lA). [0] 220-221). numerous branches; [IJ simple, short vein. (Laurentiaux, 1960; Hennig, 1981: 202; fig. 48). 14. RAPTORIAL FORELEGS. [OJ no; [lJ yes. 27. ANOJUGALAREA (= 'ANAL LOBE'; 'VANNAL 15. NUMBER OF TARSAL ARTICLES. [0] 5; [1] 4. FAN'; 'VANNUS'; 'JUGUM'; 'NEALA') - HINDWING. [OJ enlarged; [lJ reduced; [2J essentially or totally absent. 16. WINGS. [OJ present; [1] absent. The enlarged lobe found in the hindwing of roaches, mantids and termites of the family Mastotermitidae is 17. TEGMINA (forewings thickened). [0] absent; [IJ generally called an 'anal lobe'. This term is inaccurate present. Tegmina are known from fossil orthopteroids in or at least confusing. In 1937 Martynov published a de- the Carboniferous. Holmgren (1911) and Martynov (1937; tailed study of the basal portions of orthopteroid wings. 1938) argue that tegmina are a derived condition; Hennig He demonstrated that in the forewings of Mastotermes (1981: 198) considers membranous forewings the derived darwiniensis and cockroaches there is a broad thickened state and hypothesizes that tegmina are probably ancestral. 'basal stripe' just behind the third axillary sclerite. This Phylogeny ofthe Dictyoptera 259 stripe nearly reaches the small triangular projection of the Whatever the term, it should be realized that the anal lobe base of the anal portion of the wings (Fig. 1). The forewing or region (clavus) of the forewing and the enlarged field of anal lobe (or clavus, Wootton, 1979) of Mastotermes is not the hindwing are not homologous structures. All primitive well developed and shows only faint signs of tracheation. termites have an anal lobe (clavus) in the forewing, but it In the hindwings the same thickened stripe is seen posterior is not usually observed or illustrated because it is anterior to the third axillary. Martynov recognized the chitinous to the basal suture. Among termites the hindwing anojugal jugal stripes of the fore- and hindwings as homologous. expansion is found only in Mastotermes darwiniensis and Similarly, he suggested that the broad membranous ex- in fossil impressions of the Mastotermitidae. pansion behind the basal jugal stripe in the hindwing is homologous with the reduced jugal lobe in the forewing. 28. BASAL SUTURE. [0] absent; [1] suture weak and In the hindwing the region homologous with the anal lobe not visible for its entire length (Emerson, 1933: 175); or clavus of the forewing is in fact elongated and strongly [2] suture well pronounced in forewing; distinct line of narrowed. The first anal vein (AI = PCu in Fig. 1), defining weakness in hindwing. Ordered nonadditively because the anal region of the hindwing, is present in all primitive polarity is unclear. termites, not just Mastotermes. Wootton (1979) addressed homologies and termino- 29. CURVATURE OF FOREWING SUTURE. [0] logies of wing areas, and recommends reference to the primitively absent; [1] suture only slightly curved; [2] dictyopteran hindwing expansion as an anojugal area. sutnre more distinctively curved. (Emerson, 1933: 168).

AX3 3rd axillary --/---,;;"

BASAL PART OF FOREWING base of anal region_

1CuA.CU,

CuP =cU2

JUGAL REGION

BASAL PART OF HINDWING

Fig. 1. Basal portions of the wings of Mastotermes darwiniensis (redrawn from Martynov, 1937). 260 Barbara L. Thorne and lames M. Carpenter

30. PROVENTRICULUS SYMMETRY. [0] radial; [1] (female segment 8). [0] extensions lacking; [1] postlateral bilateral (Chopard, 1965; Deleporte, 1988: 145-146; extensions of shelf. (McKittrick, 1964: 100). Deleporte et aI., 1988). 43. EIGHTH FEMALE STERNUM. [0] entire; [1] 31. PROVENTRICULUS TEETH. [0] little differenti- divided. (Imms, 1919; Crampton, 1923; Fuller, 1924: ation in size and profile of primary and secondary plicae fig. 17; Grasse, 1986: 61, fig. 53, redrawn from Crampton, (= teeth); [1] primary and secondary denticles are con- 1923). spicuously dissimilar. (Judd, 1948: 101-103, 123). 44. NINTH FEMALE STERNUM. [0] entire; [1] divided 32. PROVENTRICULUS PULVILLI. [0] pulvilli of (2 plates). primary plicae (teeth) expand anteriorly and partially overlap corresponding teeth; [1] do not. (McKittrick, 45. OPENING OF FEMALE SPERMATHECA. [0] be- 1964: 75). tween eighth and ninth sterna; [1] on the ninth sternum; [2] through a pore midline of eighth sternum. (Imms, 33. SECONDARY ｐｕｌｖｉｾｌｉＮ [0] rounded, bulbous 1919: 121-123; Browman, 1935; Roonwal, 1956; Marks (termites); [1] flat (roaches, including Cryptocercus). & Lawson, 1962; McKittrick, 1965; Matsuda, 1976: 186; (Deleporte, 1988: 142, fig. 27). Boudreaux, 1979: 216). Note that Matsuda (1976: 191) stated, 'In 34. PROVENTRICULUS INTERCALARIES. [0] (Fernard, 1896; Ito, 1924) and Stagmomantis (Fig. 54B) longitudinal folds between teeth; [1] intercalary plate = the spermatheca consists of the pyriform reservoir and the highly modified new feature - sclerotizations ofphragmata canal that opens on the 8th sternum above the gonopore. between denticles and interdentiaries; [2] anastomosing The opening lies anterior to the 9th sternum that supports ridges. (Judd, 1948: 102, 105, 123, 126; McKittrick, 1964: the lateral valvulae.' We have not been able to determine 75; Deleporte, 1988). Ordered nonadditively because the generality of this state in mantids. polarity is unclear. Ordered nonadditively because polarity is unclear.

35. TERGA. [0] all visible; [1] seventh tergum (as well 46. FORM OF PARAPROCTS. [0] flat in cross-section; as sternum) posteriorly expanded to conceal completely [1] V-shaped in cross-section. (McKittrick, 1965: 19). segments 8, 9 and 10. 47. FUSED PARATERGITES. [0] absent; [1] present 36. TENTH TERGUM. [0] is true tenth tergum - mantids (ninth paratergites broadly confluent with tergum 9). (Walker, 1919, 1922; Matsuda, 1976: 187); [1] a composite (Fuller, 1924: 72; Roonwal, 1956; Marks & Lawson, 1962: segment in which the supraanal lobe and the true tenth 142-143; McKittrick, 1965: 19; Matsuda, 1976: 181). tergum are fused or undifferentiated (Matsuda, 1976: 181). 48. MALE GENITALIA. [0] well developed with three 37. NUMBER OF ABDOMINAL STERNA (IMAGO). asymmetric phallomeres; [1] highly reduced; no sclerotized [0] 11; [1] 10. (Snodgrass, 1931; Browman, 1935; Roonwal, copulatory structures present (Beier, 1956; Weesner, 1969: 1956; Waterhouse, 1970). 155; Matsuda, 1976: 171, 187; Boudreaux, 1979: 220).

38. OVIPOSITOR BLADES. [0] protruding from body 49. VASA DEFERENTIA. [0] open into ejaculatory duct cavity; [1] internal. from the ventral side; [1] from the dorsal side. (Weesner, 1969: 151; Matsuda, 1976: 184, fig. 53A and B). 39. SHAPE OF FIRST AND THIRD PAIRS OF OVI- POSITOR VALVES. [0] no odd shape or moveable spines 50. MALE STERNA. [0] seventh sternum not enlarged on first valves; [1] unique and distinctive, and with small to form genital plate, and does not cover sterna 8 and 9; moveable spines on first valves. (Marks & Lawson, 1962: [1] male sterna resemble female with enlarged seventh 155-156). sternum. (Roonwal, 1956).

40. DEVELOPMENT OF OVIPOSITOR VALVES. [0] 51. SPERM FLAGELLA. [0] single flagella; [1] afIagel- well developed; [1] reduced; [2] highly reduced; [3] only late; [2] multiflagellate. (Imms, 1948; Baccetti & Dallai, ventral valves present plus membranous elevations. (Imms, 1977, 1978; Baccetti et aI., 1981; Baccetti, 1987). Ordered 1919; Browman, 1935; Marks & Lawson, 1962: 142-143). nonadditively because polarity is unclear.

41. EGG DEPOSITION. [0] eggs laid in mass, 2 rows, 52. SPERM ACROSOME. [0] present; [1] absent. with secretory envelope; [1] eggs laid in ootheca with dis- (Baccetti et aI., 1971, 1974, 1981; Baccetti, 1979, 1987). crete outer case; [2] eggs laid singly. Ordered nonadditively because polarity is unclear. 53. STYLI ON MATURE FEMALE. [0] present; [1] absent (lost sometime during instars from immature to 42. EXTENSIONS OF LATEROSTERNAL SHELF imago). (Fuller, 1924: 74; Browman, 1935: 120, 121; Phylogeny of the Dictyoptera 261

Grasse, 1982: fig. 54). that ... termites and cockroaches were both derived from a common ancestral stock with XX/XY sex-determination, 54. CERCI (IMAGO). [0] 6 or more segments; [1] 3-5 with more rapid evolutionary loss of the Y chromosome segments; [2] 2 segments. (Emerson, 1933: 175; Krishna, in cockroaches.' Note: Luykx (1990) discovered one case 1961: 315). of XX/XO sex determination in termites: Stolotermes victoriensis (Termopsidae). Luykx considers this case 55. BACTEROIDS IN ADIPOSE TISSUE. [0] present; derived in convergence with cockroaches; not a [1] absent. (Koch, 1938; Jucci, 1952; Gharagozlou, 1966; synapomorphy. Grasse, 1986: 496). 66. CHROMOSOME NUMBER. [0] 2n ::;; 52; [1] 2n = 98. 56. SOLDIER SIZE AND HEAD SHAPE. [D] soldiers primitively absent; [1] large with flat elongated head; [2] 67. EUSOCIAL LIFE. [0] absent; [1] highly developed not so. with caste system.

57. SOLDIER OCELLI. [0] soldiers primitively absent; 68. INQUILINES. [0] primitively absent; [1] inquilines [1] ocelli present (although vestigial); [2] absent. (e.g. termitophiles) absent; [2] inquilines fully integrated. (Seevers, 1957: 58, 280; Emerson, 1965: 28). 58. SOLDIER BLADELIKE PROTUBERANCE ON FRONT COXAE. [0] soldiers primitively absent; [1] 69. FORAGING AWAY FROM NEST. [0] group for- protuberances absent; [2] present. aging primitively absent; [1] foraging within nest wood only; [2] foraging away from nest. 59. SOLDIER CERCI. [0] soldiers primitively absent; [1] 5-7 articles, twice length of stylus; [2] 5 articles, ap- 70. NEST ARCHITECTURE. [0] nest building primi- proximate length of stylus; [3] 2 articles, stylus slightly tively absent; [1] excavation ofwood; no building of exten- longer than cerci. sive gallery network; [2] nest construction away from or within host wood. 60. REPRODUCTIVE SOLDIERS. [0] soldiers pri- mitively absent; [1] yes (both sexes); [2] no - soldier gonads reduced or vestigial. (Imms, 1919; Gay & Calaby, Results and Discussion 1970; Myles, 1986). Exact analysis of the matrix in Table 3 by implicit enumer- 61. SOLDIER DEFENCE. [0] soldiers primitively absent; ation resulted in one cladogram. It is shown in Fig. 2: the [1] mandibles only; [2] mandibles (mechanical defence) length is 109, with a consistency index (Kluge & Farris, plus chemicals. (Moore, 1968; Prestwich, 1984). 1969) of 0.88 and a retention index (Farris, 1989) of 0.84. This result is stable to successive weighting (Farris, 1969; 62. (WORKER) ANTERIOR CAECA OF THE ME- Farris et al., in prep.). The cladistic diagnoses (Farris, SENTERON. [0] present; [1] absent. (Noirot & Noirot- 1979, 1980) for the cladograrn are given in Table 4. The Timothee, 1969: 73). diagnoses show the distribution of all characters over the entire tree. 63. WORKER/PSEUDERGATE DEVELOPMENTAL PLASTICITY. [0] caste differentiation primitively absent; [1] substantial flexibility (reversionary and conversionary malts; stationary and saltatorial molts); [2] fairly rigid - Termopsidae no reversionary molts. (Watson & Sewell, 1985). [See also Noirot, 1985; Watson & Abbey, 1985; Grasse, 1986, for .------Mastotermitidae dicussion of worker/pseudergate distinction.] '----Kalotermitidae 64. NUMBER OF STERNAL GLANDS (PSEUDER- GATE/WORKER). [0] 3; [1] 1; [2] D. (Noirot, 1969: 98; .------Mantodea Roth & Alsop, 1978: 472). Ordered nonadditively because polarity is unclear. Blattodea 65. SEX DETERMINATION. [0] XX/XY; [1] XX/XO. (White, 1976; Luykx, 1983, 1990; Bedo, 1987). Bull (1983) "---{:ryptocercidae and Luykx (1990) regard the XO male condition as de- Fig. 2. Phylogeny of the Dictyoptera based on analysis of the rived. Luykx (1990) comments, 'The difference in their matrix in Table 3. The closer relationship of Blattodea + basic mechanism of chromosomal sex determination ... Cryptocercidae to Mantodea is strongly supported, corroborating argues against a simple evolutionary derivation of termites the hypothesis of Boudreaux (1979). Relationships of families (XX/XY) from cockroaches (XX/XO). It is more likely within the Isoptera are not firmly established. 262 Barbara L. Thorne and lames M. Carpenter

Table 4. Cladistic diagnoses for the dictyopteran cladogram (Fig. 2). Apomorphies are listed for each component (branch point; Nelson, 1979) and terminal taxon; ambiguous optimizations, where the state could not be determined, are indicated by question marks. The numerical codes for each apomorphy in Table 4 are listed in parentheses. Blattodea is not listed separately, as it has no apomorphies.

Mantodea + (Cryptocercidae + Blattodea) 27. Anojugal area of hindwing reduced (1) or absent (2)? 4. Ocelli (imago) 3 (0) or 2 (I)? 28. Basal suture absent (0), weak (1) or well pronounced (2)? 12. Segmental arterties present (1). 36. Tenth tergum composite structure (1). 17. Tegmina present (1). 37. Number of abdominal sterna (imago) 10 (1). 19. Forewing preradius absent (1). 38. Ovipositor blades internal (1). 21. Media closer to RS (1). 40. Ovipositor valves reduced (1) or highly reduced (2)? 26. Postcubitus of hindwing simple (1). 41. Eggs laid singly (2). 31. Proventriculus teeth with primary and secondary denticles 45. Opening of female spermatheca on the ninth sternum (1). dissimilar (1). 48. Male genitalia highly reduced (1). 33. Secondary pulvilli rounded (0) or flat (I)? 52. Sperm acrosome absent (1). 34. Proventriculus intercalaries longitudinal folds (0), 53. Styli on mature female present (0)' or absent (I)? intercalary plate (1) or anastomosing ridges (2)? 55. Bacteroids in adipose tissue present (0) or absent (I)? 36. Tenth tergum present (0) or composite structure (I)? 56. Soldiers large with flat elongated head (1). 38. Ovipositor blades protruding (0) or internal (I)? 57. Soldier ocelli present (1) or absent (2)? 41. Eggs laid in ootheca (1). 58. Soldier bladelike protuberance on front coxae absent (1). 42. Extensions of laterosternal shalf lacking (0) or present (I)? 59. Soldier cerci 5-7 articles, twice length of stylus (1). 45. Opening of female spermatheca through a pore midline of 60. Reproductive soldiers (1). eighth sternum (2). 61. Soldier defence mandibles only (1). 46. Form of paraprocts flat (0) or V-shaped in cross-section (I)? 63. Worker/pseudergate developmental plasticity present (1). 53. Styli on mature female absent (1). 64. Number of sternaI glands (pseudergate/worker) 1 (1). 64. Number of sternal glands (pseudergate/worker) 1 (1). 67. Eusocial (1) 65. Sex determination XX/XO (1). 68. Inquilines absent (1). Mantodea 69. Foraging within nest wood only (1). 3. Eyes broadly separated and rounded, enabling stereoscopic 70. Nest architecture excavation of wood (1). vision (1). Termopsidae 14. Forelegs raptorial (1). 2. Number of antennal articles (imago) 23-27 (2). 20. Forewing subcosta long (1). 4. Ocelli (imago) none (2). 24. Forewing jugal lobe enlarged (1). 6. Right mandible with subsidiary tooth on front base of first 25. Hindwing subcosta long (1). marginal tooth (1). 34. Proventriculus intercalaries anastomosing ridges (2). 27. Anojugal area of hindwing absent (2). Cryptocercidae + Blattodea 28. Basal suture weak (1). 4. Ocelli (imago) 2 (1). 40. Ovipositor valves highly reduced (2). 9. Pronotum expanded (1). 55. Bacteroids in adipose tissue absent (1). 30. Proventriculus symmetry bilateral (1). 57. Soldier ocelli absent (2). 33. Secondary pulvilli flat (1). Mastotennitidae + Kalotermitidae 34. Proventriculus intercalaries intercalary plate (1). 2. Number of antennal articles (imago) 30-31 (1) or 23-27 36. Tenth tergum composite structure (1). (2)? 38. Ovipositor blades internal (1). 5. Left mandible with fused first and second marginal teeth (1). 42. Extensions of laterosternal shelf present (1). 18. Wing micro-sculpturing papillae and pimpules (1). 46. Form of paraprocts V-shaped in cross-section (1). 21. Media closer to RS (1). Cryptocercidae 27. Anojugal area of hindwing reduced (1) or absent (2)? 7. Mandibular glands absent (1). 28. Basal suture well pronounced (2). 8. Hypopharyngeal glands absent (1). 29. Forewing suture distinctively curved (1). 16. Wings absent (1). 32. Proventriculus pulvilli expanded (0) or not (I)? 35. Seventh tergum posteriorIy expanded (1). 40. Ovipositor valves reduced (1) or highly reduced (2)? 39. Shape of first and third pairs of ovipositor valves distinctive, 43. Eighth female sternum divided (1). with moveable spines on first valves (1). 51. Sperm flagella single (0), aflagelIate (1) or multiflagellate 50. Male seventh sternum enlarged (1). (2)? 64. Number of sternal glands (pseudergate/worker) 0 (2). 53. Styli on mature female absent (1). 54. Cerci (imago) 3-5 segments (1). Isoptera (Termopsidae + (Mastotermitidae + Kalotermitidae» 55. Bacteroids in adipose tissue present (0) or absent (I)? 1. Antennae short and moniliform (1). 56. Soldiers not large with flat elongated head (2). 2. Number of antennal articles (imago) 30-31 (1) or 23-27 57. Soldier ocelli present (1) or absent (2)? (2)? 59. Soldier cerci 5 articles, approximate length of stylus (2) 4. Ocelli (imago) 2 (1). 60. Reproductive soldiers absent (2). 10. Ventropleurite absent (1). I!. Aorta arising from mesothoracic enlargements, etc. (1). Mastotermitidae 13. Ganglia fused (1). 2. Number of antennaI articles (imago) 30-31 (1). 21. Media closer to Cu (0) or RS (I)? 9. Pronotum expanded (1). 23. Anal area of forewing reduced (1). 27. AnojugaI area of hindwing reduced (1). Phylogeny ofthe Dictyoptera 263

Table 4. (continued)

32. Proventriculus pulvilli not expanded (1). 70. Nest architecture away from or within host wood (2). 40. Ovipositor valves reduced (1). 41. Eggs laid in mass, 2 rows, with secretory envelope (0). Kalotermitidae 44. Ninth female sternum divided (1). 2. Number of antenna] articles (imago) 11-21 (3). 45. Opening of female spermatheca between eighth and ninth 15. Number of tarsal articles 4 (1). sterna (0) 18. Wing micro-sculpturing papillae, pimpules and tubercles 47. Paratergites fused (1). (2). 49. Vasa deferentia open into ejaculatory duct from ventral 19. Forewing preradius absent (1). side (0) 22. Media simple (1). 51. Sperm flagella multiflagellate (2). 25. Hindwing subcosta absent (2). 57. Soldier ocelli present (1). 27. Anojugal area of hindwing absent (2). 58. Soldier bladelike protuberance on front coxae present (2). 40. Ovipositor valves present only as ventral valves (3). 61. Soldier defence mandibles plus chemicals (2). 51. Sperm aflagellate (1). 63. Worker/pseudergate developmental plasticity absent (2). 54. Cerci (imago) 2 segments (2). 64. Number of sternal glands (pseudergate/worker) 3 (0). 55. Bacteroids in adipose tissue absent (1). 66. Chromosome number 2n = 98 (1). 57. Soldier ocelli absent (2). 68. Inquilines present (2). 59. Soldier cerci 2 articles, approximate length of stylus (3) 69. Foraging away from nest (2). 62. (Worker) anterior caeca of the mesenteron absent (1).

Although the support for the cladogram as measured by termite families is performed. That analysis is currentiy in the fit statistics is quite good, the result may be construed progress therefore we do not make any recommendations as surprising. Therefore, we performed supplemental on classification at this time. analyses designed to assess how robust the result is. First, This data set and analysis represent a synthesis of cur- we determined the number of steps required to attain rent information on basal groups comprising modern results according with received wisdom, using the 'XX' Dictyoptera. We hope that the paper will function as a interactive routine. For Blattodea + Cryptocercidae provocative stimulus for future work on these taxa. Inter- to be placed as closer to Isoptera than to Mantodea, a pretations and conclusions regarding the phylogeny of length of 113 is required, four more steps than required this group may well be revised as new information is for Fig. 2. In order for Cryptocercidae to be placed in compiled. Further comparative anatomical and molecular turn as closer to Isoptera than to Blattodea, a length approaches towards resolution of dictyopteran systematics of 119 is required, ten more steps than the cladogram will provide an important complement to this study. of Fig. 2. Thus, the closer relationship of Blattodea + Cryptocercidae to Mantodea is strongly supported, cor- roborating the hypothesis of Boudreaux (1979). Although the position of Cryptocercidae within the roach is Acknowledgments uncertain, the notion of roach in terms of the termites should be discarded. This paper is an expanded version of a symposium talk pre- We also investigated alternative arrangements within sented by B.L.T. at the 1988 meeting of the Entomological Isoptera. For Mastotermitidae to be placed as the relatively Society of America (Louisville, Kentucky) by invitation most basal element of Isoptera (that is, for Termopsidae of George C. Eickwort. We thank him for encouraging us and Kalotermitidae to appear as sister-groups in this analy- to investigate this topic. Frank M. Carpenter, William D. sis), a length of 112 is required, three more steps than Hamilton, Michael I. Haverty, Lynn S. Kimsey, Kumar Fig. 2. The weight of evidence for the relatively apical Krishna, David R. Maddison, Christine A. Nalepa, William placement of Mastotermitidae is thus not great. The L. Nutting, Louis M. Roth and J. A. L. Watson were each arrangement shown in Fig. 2 also resulted under each of instrumental in helping us to assemble data and to develop, these conditions: (1) all characters were treated as nonad- interpret or discuss concepts presented in the paper. We ditive, discarding all hypothesized polarities; (2) all traits sincerely appreciate each of their contributions. Sub- pertinent to social behaviour were deleted (i.e. characters stantive and constructive reviews of earlier drafts of this 56-61, 63 and 66-70); and (3) the behavioural traits manuscript by Lynn Kimsey, Kumar Krishna, Niels P. were deleted and all remaining features were treated Kristensen, Leigh R. Miller, William L. Nutting, Lisa as nonadditive. The relative disparity between the al- Vawter, J. A. L. Watson and Robin J. Wootton assisted ternative placements of Mastotermitidae was maintained greatly in clarifications and revIsions. During the course through all these analyses (as, for that matter, was the ofthis work B.L.T. was supported, in part, by NSF grant relative number of steps of attain roach paraphyly). Thus BSP-8607407 and a Science Scholars Fellowship from the on present evidence mastotermitids are evidently not Mary Ingraham Bunting Institute of Radcliffe College. living fossils, but this should not be considered as firmly J.M.C. was supported by NSF grants BSR-8817608 and established until a comprehensive analysis including. all BSR-9006102. 264 Barbara L. Thorne and lames M. Carpenter

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Appendix 1. Characters omitted from analysis. The following characters were omitted due to ambiguity of state, non- independence or lack of variance within the considered taxa.

Imago Soldier Size Pilosity Head pilosity .Head capsule shape Head mobility Compound eye shape Wing pilosity Compound eye pigmentation Size third antennal article No. antennal segments Postclypeus shape Frontal gland/fontanelle opening Tibial spur number and position Distinction of epicranial suture Tarsal pulvilli Ratio of third:fourth antennal segments Front coxae shape Antennal article shape Degree of development of seminal vesicles/accessory gland Protuberances above antennal fossae complex Sex of soldiers Intersternal fold - sclerotized or membranous Postclypeus longitudinal groove Forked spermatheca Molar ridges No. of testicular lobes Mandible serrations No. of chromosome arms (nombre fundamental) Soldier pronotum shape Chromosome centricity (acro-, meta- or telocentric) Coxae Distinction of epicranial suture Spines on front tibiae Secondary sclerotization of sternum 7 Workers Archidictyon Crop symmetry Position of anal veins in forewing Position of sternal glands Proportions of wing scales Crop/gizzard separation Position of media in hindwing Workers with functional eyes Meta- and mesothorax morphology Length of first section of hindgut Shape of compound eye Femoral grooves, front femora Ecology/biology/habitat Independent versus joint insertion of vasa deferentia Nitrogen fixation (exceedingly high in Mastotermes) Proventriculus: rigidity of dental belt Colony population size Proventriculus: pulvillar belt Intervalvular ring =basal ring No. of malpighian tubules in imago No. of malpighian tubules present during first larval instar No. of insertion groups of malpighian tubules Inferior hypostomal gland (= maxillary/accessory mandibular gland)

, .\