Two New Genera of Sough American Cockroaches Superficially Resembling Loboptera, with Notes on Bionomics (Dictyoptera, Blattaria, Blatterllidae).
Psyche 73:196-207, 1966.
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TWO NEW GENERA OF SOUTH AMERICAN
COCKROACHES SUPERFICIALLY RESEMBLING
LOBOPTERA, WITH NOTES ON BIONOMICS
(DICTYOPTERA, BLATTARIA, BLATTELLIDAE) . All cockroaches initially extrude the ootheca with the keel or the micorpylar ends of the eggs facing dorsally, but some species rotate the egg case before depositing it (Roth and Willis, 1954, 1958). According to McKittrick ( I 964), one of the important characters for separating the Plectopterinae from the Blattellinae (both are in the Blattellidae) is the position in which the ootheca is carried just before it is deposited by the female. In the Plectopterinae, the keel of the egg case remains upright until deposition; in the Blattellinae, the o~theca is rotated so the keel and micropylar ends of the eggs face laterally, behavior characteristic of the Ectobiinae and Ny~tibo~inae (Blattellidae)
as well as of all the Blaberidae (ovoviviparous and viviparous species). The new genera here described were recognized as a result of studies stimulated by observing the lack of rotation by a species which formerly had been assumed to be one of the Blattellinae. Therefore, in addition to presenting descriptions and biological notes, this paper provides an application of the higher cate- gories in the classification of McKittrick ( 1964). On June 11, 1965, one of us (L.M.R.) received some living specimens of Loboptera thaxteri Hebard from Buenos Aires, Argen- tina.I When we found that the females of thaxteri do not rotate the o6theca before its deposition, we investigated the taxonomic position of Loboptera Bsunner. McKittrick (1964) had reported the rota- tion of the ootheca by Lobopterella dimidiatipes (Boli~ar),~ and as a result of this and other observations she placed it in the Blattelli- nae.
[Loboptera decipiens (Germar) also rotates its otitheca before deposition (Lefeuvre, 1959; Roth, unpublished observations).] We *Entomology Research Division, Agr. Res. Serv., U. S Department of Agriculture, Washington, D. C.
**Pioneering Research Division, U. S. Army Natick Laboratories, Natick, Massachusetts.
Manuscript received by the editor September 21, 1966. 'The species has been cultured easily on Purina laboratory chow. How- ever, individuals of thaxteri were heavily parasitized by the fungus Herpo- myces lohopterae Thaxter.
'For many years authors referred dimidiatipes to Loboptera, and Mc- Kittrick did so. Princis (1957) based the genus Lobopterella upon it.
3 9661 Gurney and Roth - Cockroaches I97 therefore sent specimens of thaxteri to Dr. McKittrick without iden- tifying the material, and she placed it in the Plectoptei-inae. She wrote, "When I looked at it, I thought 'Loboptera' and was thor- oughly surprised when I checked the genitalia.') In the past years, various species seeming to belong to the Old World genus Loboptera Brunner [type species, L. decipiens (Ger- mar),3 an Old World species] have been described from the New World. Such generic assignment of New World species has been based primarily on superficial appearance. Two South American species, in particular thaxteri Hebard and insularis R. S. Albuquer- que and Gurney, have a general appearance that is so similar to decipiens that Hebard (1932) assumed thaxteri to be an introduc- tion from the Old World. In fact, no previously described genera appear adequate for the inclusion of thaxteri and insularis. There- fore in order to assign these species in a manner consistent with current generic concepts in the Blattaria, we are describing a sepa- rate genus for each. Although the two seem congeneric in general appearance, the differences noted in the generic key following to distinguish Agmoblatta, new genus, from Isoldaia, new genus, are fundamental.
Agmoblatta, new genus
This genus closely resembles Loboptera in general external ap- pearance; it differs as noted in the following key. Its closest rela- tive is Isoldaia, also known from eastern South America. Generic description: Size medium for Blattellidae. Body with sparse slender setae slightly shorter than 1st antennal segment; moderately glossy.
Head very broad; interocular distance a little greater than distance between antennal sockets ; vertex smooth ; antennae unspecialized; maxillary palpus of moderate length, apical segment narrowly triangular. Pi-onotum hemispherical, moderately vaulted, posterior margin straight except near simply rounded lateral angles. Tegmen a narrow, subtriangular lateral pad ; wings absent. Front femur with anteroventi-a1 margin (Fig. 8) bearing a few strong spines, I or 2 of which toward knee of decreasing length, followed by about 5 piliform spines and 3 strong apical spines of increasing length; posteroventral margin with 3 or 4 spaced, strong, short spines in apical half, a slightly longer apical spine. Middle and hind femora with strong spines on both ventral margins. Middle and hind tibiae spined in 3 series on dorsal surface. Basal tarsomere of hind tarsus slightly more than one-half length of tarsus, ventral 'Hebard (1922, p. 332) indicated the type species.
198 Psyche [September
Figs. 1-9. Male genitalia of Loboptera, Lobopterella, and Agmoblatta (All figures except no. 2 drawn from KOH preparations) 1-3. Loboptera decipiens (Germar), male from culture at MacDonald College, originating in the Azores.
1. Ventral view of paraprocts and
2. Ventral view of subgenital plate. 3. Dorsal view of phallomeres.
4. Loboptera maroccana Bolivar, male from Tangiers. Dorsal view of phallomeres.
5-6. Lobopterella dimidiatipes (Bolivar), male from Fiji. 5. Dorsal view of phallomeres.
6. Ventral view of subgenital plate.
19661 Gurney and Roth - Cockroaches 199
surface with double row of closely set, strong, short setae, pulvill~~s occupying apical loth of ventral surface: small pulvilli on tarsomeres 1-4; claws simple, equal ; asolium of moderate size. Abdomen smooth (Figs. 15, 16) ; lateroposterior angles of terga simple; tesgum 7 of male with a wide median depression in which a tuft of hairlike setae arises
(Figs. 15, 20) ; cercus with about 9
segments. Male genitalia: Supra-anal plate broadly triangular, un- specialized; subgenital plate (Fig. 9) with specialized posterior margin, with styli ; paraprocts without conspicuous specialization ; phallomeres (Fig. 10) of plectopterine type, 1st sclei-ite of left phallo- mere ( LI ) a conspicuous framework, median sclerite (L2VM ) slender and elongate ; 2nd sclei-ite of right phallomere (R2) a con- spicuous hook. Female supra-anal plate triangular, simple ; subgenital plate broad, posterior margin weakly cleft medially and bent dorsad (Fig. 19).
Type-species : Lo boptera thaxteri Hebard. The name Aginoblatta is derived in part from the Greek word "Agn~os)" meaning a break or fracture and has reference to the cleft of the female subgenital plate.
Hebai-d (1932) described only the female of A. thaxteri and erred in suggesting that it is parthenogenetic; he based this suggestion on the fact that all his specimens were females. The male markings strongly resemble those of the female (Figs. 15, 16). The female genitalia of A. thaxteri and Loboptera decipiens are shown in Figs. 22 and 24. The female genitalia of Lobopterclla are illustrated by McKittrick (1964, p. 161). Agmoblatta thaxteri has one pair of spermathecae, but each spermatheca has a double terminal bulb (Fig. 25). Loboptera decipiens also has only one pair of spermathecae (Fig. 23), but each has many amber-colored tubules or branches. Sixteen of these branches terminate in rounded colorless bulbs in which the sperm are stored. Among the species studied by Mc- Kittrick ( 1964)) only members of the Blattidae and Cryptocercidae have "forked" (i.e., branched) spermathecae; none of the Blaberoi- 7. Loboptera decipiens (Germar), male from Madeira. Spines on antero- ventral margin of front femur.
8-9. Agmobiatta thaxferi (Hebard). 8. Female paratype, spines on anteroventral margin of front femur. 9. Male from Natick culture originat- ing in Buenos Aires, Argentina. Ventral view of subgenital plate. Abbreviations: L2VM=median sclerite (second or ventromedial sclerite of left phallomere) ; L3 3rd sclerite of left phallomere ; LPA :== left paraproct ; PS = piliform spines ; R2 = 2nd sclerite of right phallomere : R3 = 3rd sclerite of right phallomere; RPA= right paraproct; RS = right stylus; WPA = weakly pigmented area of subgenital plate.
Figs. 10-14. Male genitalia of Agmoblatta and Isoldaia (All drawn from KOH preparations)
10. Agmoblatta thaxteri (Hebard). Male from Natick culture originating in Buenos Aires, Argentina.
Dorsal view of phallomeres.
11-14. Isoldaia insularis (R. S. Albuquerque and Gurney), male holotype. 11. Dorsal view of phallomeres.
12. Ventrolateral view of apical portion of subgenital plate. 13. Ventral view of paraprocts and associated struc- tures.
14. Ventral view of subgenital plate.
Abbreviations : LI = 1st sclerite of left phallomere ; L2 = second dorsal sclerite of left phallomere; other abbreviations as in figs. 1-9. dea have this type of spermatheca. She believes that the branched spermatheca is a primitive condition. The many branched spermatheca in Loboptera dicipiens is unique for the Blattellidae and may repre- sent a relic character. Lobopterella dimidiatipes has 2 pairs of spermathecae ( McKittrick, 1964, p. I 53). The ootheca (Fig. I 7) of A. thaxteri has a deep groove ventrally
19661 Gm-ney and Roth - Cockroaches 20 I along the midline (Fig. 18) made by the upturned medial cleft (Fig. 19) of the subgenital plate; the cleft serves as a mold as the ootheca is formed and extruded posteriorly. The ootheca remains upright until deposition, and its keel rests in the medial marginal indentation of the supragenital plate. The eggs require water for development, and the females usually deposit their oothecae on the moist cotton of the water vials.
Observations were made on the courtship behavior of AgnzobZatta. A newly emerged female was placed with 2 old males. Initially, the males showed only mild interest in the female, but within an hour they were actively pursuing and courting her. The males antennated and palpated the female and turned with their backs toward her, stretching and arching their abdomens and fully exposing the gland an the 7th terga. However, the female did not respond by mounting and palpating the gland, and the males did not extend their phallo- meres and attempt to grasp her genitalia. This behavior is similar to that of BZattelZa germanica (L.) (Roth and Willis, 1952)~ Nauphoeta cinerea (Olivier), and other genera in which the male will not attempt to seize the female until she mounts or palpates his tergum (Roth and Barth, 1964). The male af h'qnzoblatta relies on antenna1 contact in pursuing the female; if he loses this contact, he seems to wander about haphazardly until he again makes contact, turns his back, and courts. Also, the male can discriminate between contact with a female and a male; males on touching each other did not court. This behavior is very similar to that shown by B. german- ica (Roth and JVilIis, 1952).
The female we used had not mated 2 hours after the start of ob- servations. However, females do mate soon after emergence since a pair was seen in copula in the typical opposed position (heads fac- ing in opposite directions) though the female was still very light in color and less than a day old. Subsequently, on days 13, 20, 25, 31, and 39, she deposited egg cases. Thereafter she deposited no more oothecae though she was kept until 62 days after emerging. Isoldaia, new genus
This genus is similar superficially to AgnzobZatta, from which it differs primarily in the lack of tergal specialization in the male and in the uncleft posterior margin of the female subgenital plate. Generic description : Agrees with AgmobZatta except as follows : About 10 to 12 piliform spines in apical half of anteroventral margin of front femur, in contrast to about 5 to 6 in AgmobZatta; maxillary
GURSEY AND ROTH - COCKROACHES
19661 Gurne~j and Roth -- Cockroaches 203 palpus longer, exemplified by antepenultimate segment clearly more than half as long as width of interocular space (palpus shorter in AgnzobZatta, antepenultimate segment decidedly less than half as long as width of interocular space) ; supra-anal plate of female more acutely triangular than in Agnzoblatta; male abdomen without tergal specialization; subgenital plate of female with entire, uncleft pos- terior margin. Male genitalia and subgenital plate are illustrated in Figs, 11-14.
'Type-species : Lobo$tera insularis R. S. Albuquerque and Gurney. The name Isoldaia is adapted from the given name of Mrs. Isolda Rocha e Silva Albuquerque in recognition of her sustained efforts to broaden the knowledge of South American Blattaria. A habitus figure of a female of AgnzobZatta thaxteri is in Hebard ( 19321, and one of IsoZdaia insularix is in R. S. Albuquerque and Gurney ( 1963). Readers are referred to page 178 of McKittrick (1964) for information about the subfamily assignment of the genera wherein the phallomeres of LophobZatta Hebard, EuthZasto- bZatta Hebard, and SupeZZa Shelford represent the Plectopterinae, and Pseudomops Serville and BZatteZZa Caudell represent the Blattel- hae.
The following is a key for distinguishing Loboptera and several similar genera :
I. Conspicuous hook-shaped sclerite of male genitalia occurs as part of left phallo~nei-e (Figs,. 3-5, L3) ; one or both parapi-octs of male armed with a hook 01- spines (Fig. I) ; armature of ventro- anterior margin of front fenmi- (Fig. 7) includes heavy though short spines basad of 3 terminal ones (Type A) (Old 1,Vol-1 d )
. . . . . . . . . . . . . . . , . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . , . . . . . . . . . . , . . . . . . . , . . . . . . . . . . . . . . 2 - Conspicuous hook-shaped sclerite of male genitalia occurs as part of right phallomere (Figs. 10, I I, R2 j ; paraprocts of male simple, not distinctly armed (Fig. 13) ; armature of ventro- Figs. 15-20. 44gmoblatta thaxteri (Hebard). 1.5. Dorsal view of male; arrow points to the tergal gland on the seventh segment (X 5.2). 16. Dorsal view of female ( X 6.0). 17. Lateral view of ootheca (X 17.2). 18. Ventral view of ootheca; note deep medial longitudinal groove ( X 17.2). 19. Ventral view of end of the abdomen of a female forming an ootheca; arrow points to medial cleft in subgenital
plate (X 8.8). 20. Seventh tergum of male showing the medial tergal gland (X28.7).
Fig. 21. Lobopterella dim.'diatipcs (Bolivar) Seventh tergum of male ; arrow points to one of the 2 pits (X 21.8).
204 Psj~che [September
anterior margin of front femur includes delicate piliform spines (Fig. 8, PS) basad of 3 terminal ones (Type B) (New ........................................................................................ World) 3 2. Tegmina present as subtriangular lateral pads ; male subgenital plate simple, without styli (Fig. 2) ; dorsum of male abdomen unspecialized. (Canary Islands, Azore.~, Europe, North Africa, western Asia; other Old JVorld records subject to change with revised definition of genus) ........................... Loboptera Brunner -
Tegmina short, subquadrate, attingent or slightly overlapping ; male subgenital plate compIex, with styli (Fig. 6) ; tergum 7 of male abdomen with 2 s~nall poses without associated ,setae (Fig. 21). (Africa to Hawaii, other Pacific islands) ............................ ........................................................................ LoboptereZla Princis 3. Dorsum of male abdomen specialized on tergum 7 (Figs. 15, 20) ; posterior margin of female subgenital plate weakly cleft (Fig. 19). (Known only from Buenos Aires, Argentina) ............ .................................................................. A97720 blatta, new genus - Dorsum of male abdomen unspecialized; posterior margin of female subgenital plate entire. (Known only from San Sebastian Island about 60 miles east of Sgo Paulo, Brazil) ........................ ........................................................................ IsoZdaia, new genus Princis ( 1957) stated that the male tergum is unspecialized in LoboptereZZa; perhaps the pores on segment 7 were hidden under the 6th tergum in his museum specimens.
Several genera of Old JVorld Blattaria have been described to in- clude species appearing by traditional generic characters to be related to Lobofitera. Specimens of most of these genera are not available to us at present to permit us to stud~7 the characters shown by McICit- trick (1964) to have special significance in higher classification. When such a study eventually becomes possible, the resulting realign- ment of genesa may reveal which are most closely related to Agmo- bZatta and IfoZdaia and may suggest what the lines of evolutionary development have been.
Brief characterizations of Loboptera and 3 European species are included in Princis (1965). An illustration
of the paraprocts of Lobofltcra decipiens similar to our Fig. I, is in Bei-Bienko ( 1950, p. 169, fig. 67). The description of LoboptereZZa is in Princis ( 1957 j.
The present investigation tests the applicability of ILlcKittrick's higher categories and particularly her separation by genitalia and other characters of 2 groups of genera which for many years have been regarded as a single group, the Pseudomopinae. Our studies indicate that there is a basis for a fundamental distinction between
Gurnq) and Roth - Cockroaches
Figs. 22-23. Loboptera dccipiens (Germar), female from Natick culture, originating in France. 22. Posterior view of genitalia, 23, Spermathecae. Figs. 24-25. d4gmoblatta thaxteri (Hebard), female from Natick culture, originating in Buenos Aires, Argentina. 24. Posterior view of genitalia. 25. Spermathecae.
Abbreviations: inst. f. = intersternal fold; p p. paraprocts; pt. = paratergites; v.1 = first valve; vlf. la. = first valvifer arm; S VII = sternum VII; TX I tergum X.
206 Psyche [September
her 2 groups, Plectopterinae and Blattellinae, though additional characters such as wing venation seem to require evaluation before all genera placed in Plectopterinae by McKittrick remain there perm- anently. Most genera of Pseudomopinae, in the sense used by Hebard for many years, were placed in the family Blattellidae by Princis ( I 960) without subfamily separation. The subfamily name Pseudo- mopinae dates from Burr ( 1910, p. 152), who also used the family name Pseudomopidae. [An earlier use of Pseudomopinae by Rehn (1903, p. 260) is invalid because the name was proposed as a substi- tute for Phyllodrominae, and this substitute is required by Article 39 of the International Code of Zoological Nomenclature to be based on the valid name of the original type-genus, i.e. Blattella Caudel1.j Karny (1908) introduced Blattellidae, and that name has prionty for any higher category including both Blattella and Pseudomops. [The chronology of these group names was given by Icevan and Princis ( I 96 I ) .]
The correlation of differences in bionomics with the higher cate- gories was also tested in our study.
We thank Dr. K. Princis for identifying "Loboptera" thax~crz, Dr. Frances A. McKittrick for examining the same, Dr. R. K. Benjamin for identifying Herpoitzyces lobopterae, Mr. William Rahn for taking the photographs, and Mr. Peter Graves lor camera Lucida outlines of Figs. 22-24. We are also grateful to Dr. Pi-in..is for reading the manuscript and offering some very helpful suggestions. BEI-BIENKO, G. IA.
1950. Fauna of the U.S.S.R. Insects. Blattodea. (In Russian) 2001. Inst. Akad. Nauk., S.S.S.R., Moscow, n. s. No. 40, 343 pp., 132 figs.
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A new species of Loboptera Brunner. Ent. News 43 : 60-62, 2 figs. KARNY, H.
Die zoologische Reise des naturwissenschaftlichen Vereines nach Dalmatien im April 1906. B. Spezieller Teil. etc. 6. Orthoptera und Blattaeformia. Mitt. Naturw. Ver. Univ. Wien, 6: 101-113. KEVAN, D. K. McE., AND K. PRINCIS
1961. Blatta transfuga Brunnich, 1763 (Insecta, Dictyoptera) ; pro- posed suppression under the plenary powers. 2. N. (S.) 680, Bull. 2001. Nomencl. 18 (pt. 5) : 330-331.
19661 Gurney and Roth - Cockroaches 207
LEFEUVRE, J. C.
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1964. Evolutionary studies of cockroaches. Cornell Univ. Agric. Expt. Stat. Mem. 389: 197 pp., 205 figs., 6 text-figs. PRINCIS, K.
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1903. Studies in American Blattidae. Trans. Amer. Ent. Soc. 29: 259- 290.
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