J. G. Myers.
On the Systematic Position of the Family Termitaphidiae (Hemiptera, Heteroptera) with a Description of a New Genus and Species from Panama.
Psyche 31(6):259-278, 1924.

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19241 Systematic Position of the Family Termitaphididce 259 ON THE SYSTEMATIC POSITION OF THE FAMILY TERMITAPHIDIDB (HEMIPTERA, HETEROPTERA), WITH A DESCRIPTION OF A NEW GENUS AND
SPECIES FROM PANAMA.
In
1902 Wasmann erected the genus, Termituphis, on a peculiar termitophile which he called Termitaphis circumvallata, and which he considered an aberrant aphid. Silvestri, des- cribing two additional species in 1911, recognized that the genus was not even homopterous and established for it the new family Termitocoridse, which he placed in the sub-order, Heteroptera. In 1914 a further species was described in a preliminary manner by Mjoberg, while in 1921 Silvestri recorded from India a fifth species of the genus. These references were all listed in the Zoological Record under the family, Aphididse, and apparently received no attention from heteropterists. The list of species was brought up to eight by Morrison in 1923. Such in brief is the history of the genus.
The writer is indebted to Dr. W. M. Wheeler for the op- portunity to study and describe a ninth species collected at Panama and to offer some suggestions on the relationships of the family to other heteroptera. Mr. Harold Morrison had also received specimens of the same insect from Panama, and was about to describe it but has very generously turned over his material to me.
The Rev. E. Wasmann most kindly sent for comparison the unique specimen of the type of the genus. Thanks are due also to Dr. W. M. Mann for bringing this valuable type from Europe. Previous workers have invariably referred their mate- rial to the type-genus, but Wasmann's type shows that it is decidedly not congeneric. A new genus, Termitaradus, is there- fore erected here for the Panama species and its allies, which Contributions from the Entomological Laboratory of the Bussey Ins- titution, Harvhrd University, no. 243.




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260 Psyche [~ecem ber
undoubtedly include all species described subsequently to T. circumvallata.
None of the previous workers on these insects has made any suggestion as to the position of the family in the Heteroptera. Nor have the heteropterists themselves given the matter any attention although the very detailed and entirely adequate des- criptions and figures of Silvestri and of Morrison were all that could be desired in the absence of actual specimens. Silvestri's family name must be changed in accordance with the International Rules, which state that the family name must be derived from that of the type-genus. Termitaphis Wasm. was the original genus, and was used by Silvestri as the type-genus. The genus, Termitoccris apparently does not exist. The family name must therefore be Termitaphididse. Dr. Wheeler drew my attention to this point. Superficially the insects of this family are remarkably dis- tinct from all other Heteroptera. This unique appearance is in keeping with a habitat shared, so far as known, by no other members of the sub-order. All the species collected have been found in the nests of termites, and such characters as are entirely peculiar to the family may be tentatively explained as results of adaptation to the termitophilous habit. Reuter's (1912) Bemerkungen uber mein neues Heterop- terensystern was taken as the latest authoritative and compre- hensive review of heteropterous taxonomy. In following the key to families and also in comparing the separate diagnoses of Reuter's series and superfamilies, it was found that the Termitaphididce were best placed in or near the series Phloeobiotica, a group established to contain the two families of bark-bugs, the Aradidse and the Dysodiidse, of which the latter is now by most authorities, e. g. Parshley, 1921, con- sidered a sub-family of the former.
Reuter's diagnosis of this series is as follows (1912, p. 32) .- Unguiculi semper aroliis destituti.
Caput horizontal, inter
antennas longe prolongaturn, utrinque tuberculo antennifero plerumque acuto instructum, bucculis sulcum rostralem for- mantibus. OcdU desunt. Rostrum quadrz-articulatum, sed ar- ticulo primo minutissirno, aegre distinguendo. Antenna capite



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18341 Systematic Position of the Family Termitaphidich % 1 plerumqw longwres, QuadnarticidatcB, mpe crosses. Hemielytra e clavo, corio et membrana composita. Clams apicem versus sensim angustatus, apicem scutelli nunquam auperans. Mem- brana venis nomulus irregularibus et anastomosamtibus vel raro his tot& destituta. Meso-et metaphum simplå´iCT(~ Coxgs poså£ico rotatorice. Tarsi biarticulut-'. Corpus superne et interne deplana- turn.
The characters italicized are those which are clearly ex- hibited also by the TermitaphidiSos. The widest divergence lies in the wing characters, both pairs of wings being completelyabsent in the latter genus. But presence or absence of wings was never even a family character and there are Aradida with both pairs missing. There is therefore a strong presumption that the Ter- mitaphididae are related to the Aradidz. The presumption is
rendered almost a certainty by three other consideration's now to be examined in some detail.
Reuter (1912) laid considerable emphasis on the presence or absence of arolia as a taxonomic character. The Aradidae are said to possess no arolia and it was largely on this account that Reuter was unable to agree with Kirkaldy and with Bergroth that the Aradidse exhibit marked affinity with the Fentatomoids. The Termitaphididae on the other hand are furnished with very well-developed arolia ahown clearly in Silvestri's excellent figures (1911, 1921). Whether this deficiency should be taken to in- dicate lack of affinity between the Termitaphids and the Aradids ia questionable, since it is doubtful whether these organs afford such good taxonomic characters as has been supposed. In fact Renter, who used their presence or absence so largely, has him- self shown (1912) that they are probably of directly adaptive origin, varying apparently with the habitat even in genera of the same family. In the present case however no decision as to the importance of the arolia is essential to the argument since the Aradid genus, Ctemmewus Bergroth, (Dysodiknse, Mezirin~) possesses arolia as welt-developed as those of Termituphis, or as those of any of Reuter's aroliate families-Miridffi, Pentatomidae etc. The arolia of Ctenoneurus hochstetteri (Mayr) are shown in figure 9. This constitutes the first supplementary proof of the relationship of Ternitaphis to the Aradidse. Similar structures



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262 Psyche [~ecember
.
occur in certain species of the genera Aradus, Dysodius and Isoderrnus.
Incidentally the term "arolium" is used in general insect morphology and in hemipteran taxonomy with several different meanings which urgently need elucidation. Crampton (1923) applies the name primarily to the undivided pad-like structure between the claws of Orthoptera, e. g. Periplaneta. Further he mentions that the arolium in certain Hymenoptera and Homo- tera may be partially divided or faintly marked off into two lateral portions. There is no reference in Crampton's paper to the fact that in Heteroptera the arolium is always divided and in fact is referred to by taxonomists only in the plural. As illus- trative of the most exact use of the term in Hemipterology, figure 11 shows the arolia of a Mirid after Knight (1923). The same drawing shows also the pseudarolia which in many Mirids are greatly developed and perhaps take the place of the true arolia which are reduced to mere bristles. Knight's arolia arise as shown in the figure truly between the claws and are probably homologous with the undivided arolium described by Crampton. But in Pentatomids, Coreids, some Aradids and in Termitaphididce, the present writer finds that the arolia do not arise between the claws, but each from the base of the corres- ponding claw as shown in figures 7, 9 and 12. In these families it would seem that the so-called arolia are really homologous with the pseudarolia of the Miridse, while the true Mirid arolia are represented by bristles between the claws as shown in the Pen- tatomid, Euschistus (fig. 12) and in a Termitaphid in Silvestri's drawings. Organs evidently exactly homologous with the so- called arolia of Euschistus, Ctenoneurus and Termitaphis are des- cribed and figured in the Coreid, Anasa, by Tower (1913) as pulvilli.
Whether the appendages figured in Termitataradus and in Ctenoneurus constitute true or pseudarolia or pulvilli does not affect the question of relationship since they are obviously homo- logous structures in the two genera.
In 1920 Spooner for the first time recorded a peculiar con- dition in the Aradid head in which the rostra1 sets, instead of proceeding more or less directly cephalad and then caudad to



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19241 Systematic Positionof the Family Terrnitaphididce 263 enter the labial trough or rostrum, are coiled several times, like a watch-spring, in a semi-circular sheath formed by the tylus. The setae are thus extremely long. That such an' extra- ordinary condition should previously have escaped the notice of hemipterists is probably the result of t,he heavy chitin- isation and black coloration of the head, which renders this structure entirely invisible in the untreated insect. The present writer noticed the setal coil independently in 1920 in the newly hatched nymph of Ctenoneurus, in which the coil shows as a dark mass against the soft white nymphal tissues. This ar- rangement of the trophi is present in an almost identical condi- tion in the Termitaphididce and constitutes the second supple- mentary proof of the relationship of these interesting termitophiles with the Aradids. To these two families alone of the Heteroptera are the coiled set'= apparently confined. Here we meet the difficulty that the feeding-habits of Aradidse and even more so of the Termitaphididse are very little known. It seems likely that the insects of both families suck the sap of trees or the moisture of dead wood and of fungi. Ob- viously only liquid nutriment could be taken up by such mouth- parts.
The first important character in which the Termitaphididce ap- pear to differ from the Aradidee lies in the extraordinary develop- ment of laminae on the margin of the body, round every portion of the periphery.
These laminee are furnished with stout outwardly directed bristles and with peculiar flabella, so named by Morrison. In some Aradids there is a lobulate expansion of the flattened lateral margin of the body. Such lobes are conspicuous i4n the imago of Dysodius lunatus (Fabr.) of which Dr. Nathan Banks has shown me specimens from Panama. In addition, Dr. Wheeler collected at Barro Colorado Island, Canal Zone, Panama, , a single Dysodius nymph, probably referable to D. lunatus. This nymph, which is apparently in the third stadium, shows the marginal lobes very well-developed and offering striking points of resemblance to those of Termitaphis. There are twelve rounded lobes on each side of the body, not including projections of the head. The first is pro-, the second meso- and the third and fourth together metathoracic, while the rest pertain to the



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264 Psyche [December
abdomen.
Every lobe (fig. 10) is furnished with an irregular series of long conical processes, evidently hollow and provided with a rather thick but elongate distal flagellum usually more or less curved. The flagella are very liable to be broken off, particularly from processes near the apices of the lobes; and many are missing in the nymph under study. In the pinned imago of Dysodius lunatus no trace of the flagella is discernible, but in alcohol specimens examined later they are as well-marked as in the nymph. In the nymph there is thus a striking simi- lanty to Termitaphis in the essential features of the marginal lobes. The number and distribution of the lobes themselves, their division into lobules or processes, the presence on every lobule of an easily detachable solid appendage arising apparently at the base of the lobule and running through or beneath it,s axis to protrude beyond its apex-in all these particulars there is practical agreement between the two genera. These constitute a third group of facts which may reasonably be considered to support the hypothesis of relationship between the Termitaphi- didae and the Aradidie. The most striking superficial difference lies in the fact that the lobes in Dysodius are widely separated and thus fail to form such a continuous peripheral margin as in the Termitaphididce. In the Dysodius nymph the conical processes with flagella are present also on the margin and pro- jections of the head, and on the segments of the antennae.
The metanotum is provided with two lateral lobes instead of one as in Termitaphis and allies.
It seems probable that marginal laminae in Terrnitaradus constitute a defensive apparatus enabling the insect to withdraw all its appendages under cover. For such withdrawal the form and articulation of the peculiar antennae are especially adapted. Were the laminae closely appressed to the substratum there would remain no unprotected part of the whole periphery. A similar developn~ent of lateral laminae is frequent in myrme- cophiles and termitophiles, not,ably in the larva of Microdon and in certain beetles and Myriapoda. In' the termitophilous milli- pedes of the genera Leuritus Chamberlin and Gasatomus Cham- berlin the general form of the body segments with their lateral lobes is strikingly reminiscent of the condition in Termitaradus



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19241 Systematic Position of the Family Termitaphididc~ 265 Wasmann (e. g. 1911, pp. 228-230) recognises this type of lateral laminat'ion of body segments coupled with flattening of the ventral surface, as a direct adaptation to termitophily or myr- mecophily-as a protection against the owners of the nests in which these arthropods live. It is a modified form of the adap- tive type which he designates "der Trutztypus." All the spe- cimens of Termitaphididae so far known have been collected in company with termites or in their nests. The total absence of eyes and ocelli in Termitaphids is prob- ably correlated with life in the gloomy recesses of the termite nest. The Aradids, t,hemselves living in a cryptozoic habitat, have advanced a stage in this direction in that ocelli are lacking. The absence of wings in Terrnitaphids is similarly explicable. The peculiar structure of the antennae, by which a superficially cryptocerate condition has been achieved, has been explained as a provision for tucking these organs under the cephalic laminae. The antennae are inserted very near the lateral margin and are folded in towards the rostrum.
The chief remaining morphological distinction between the Termitaphids and the Aradids lies in the structure of the ros- trum and related parts. The head itself differs considerably. In the former it is more flattened and exhibits on side margins and fore-border a remarkable lamination with division into two main lobes on each side. This condition could perhaps be derived from that of a typical Aradid by an antero-lateral extension and lamination on each side of the tylus, so that the latter instead of forming the anterior projection of the head as in most Aradids, came to lie at the posterior end of a deep incision extending caudad from the anterior margin of the head. So far as the rostrum is concerned the Aradids show a condition which has been described as apparantly three-segment- ed but really four-segmented. As a matter of fact, in Cteno- neurus at least, (fig. 8) the second segment is peculiarly cons- tricted where it lies between the bucculse, but four distinct segments are easily discernible. In Terrnitaphids the bucculse form no appreciable sulcus for the rostrum. Wasmann des- cribed and figured the rostrum of Termitaphis as three-segmented and such it decidedly appears to be to all but the most searching



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266 Psyche [December
examination. Silvestri, however, in all his work characterizes it as four-segmented and shows four very distinct segments in his figures. Such distinctness is certainly in error. The second segment, reckoning on this basis, is very indistinctly articulated and the present writer is by no means sure that it constitutes a true segment. (Figs. 2, 3).
The dorsal pores described and figured by Morrison are unlike anything known in other Heteroptera. Possibly, how- ever, this worker's technique would reveal similar structures in other families.
To sum up it would appear that the Termitaphididae may be regarded as Aradoids specialized, in some respects degeneratively (absence of wings, eyes, ocelli and rostral sulcus), in others ad- ditionally (lateral lamination and armature and folded antennae in Terrnitaradus; physogastry in Termitaphid for a life of ter- mitophily.
The diagnosis of the series Phlceobiotica (==superfamily Aradoidea) as set out by Reuter in 1912 and quoted above, may be modified as follows to include the Termitaphidida--- Arolia present or absent; head horizontal, much prolonged between the antennae or else furnished with an acute antenni- ferous tubercle; a rostral sulcus formed by the bucculae present or absent ; ocelli absent ; rostrum 4-segmented, often thickened. Hemielytra when present formed of clavus, corium and mem- brane; clavus narrowed towards the apex and never reaching beyond apex of scutellum. Membrane with some irregular and anastomosing veins or rarely completely destitute of venation. Meso-and meta-pleura always simple. Posterior coxae rotatory. Tarsi 2-segmented. Bcdy except in Termitaphis flat)tened above and below.
This series and superfamily comprises two families dis- tinguished as follows.-
Tylus forming anterior projection of head ; bucculse forming a rostra1 &lcus; margin of body more or less simple or furnished with well separated irregular lobes. . . . . . . . . . . Aradidce (Spin.) Tylus at end of a deep incision extending caudally from anterior margin of head; bucculse forming no appreciable rostral sulcus; margin of body furnished with lobes, separate or fused,



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19241 Systematic Position of the Family Termitaphididce 267 which form a practically continuous lamina encircling the whole. Termitaphididce (n. n.)
As regards the position of the series Phlceobiotica, the dis- covery of arolia or similar structures in the Aradid, Ctenoneurus, is a further indication that Bergroth is correct in considering it nearest related to the Pentatomoids. Reuter was impressed by the fact that the eggs of Pentatomoids and of Coreoids are operculate, the embryo being furnished with a peculiar egg- burster for forcing up this lid; while the ova of Aradids, accord- ing to Heidemann, lack lids entirely and resemble more those of Lygaeids. The operculum and correlated egg-burster are, how- ever, by no means universal in the Pentatomoids, since they are totally lacking in the New Zealand Acanthosomatine genera, Oncacontias Breddin and Rhopalimorpha Mayr. The writer's notes on these insects are now in the press. In addition, obser- vations now being carried out on certain North American Coreoids indicate a lack of these structures in this superfamily also. Since the above was written I have seen Barber's (Psyche, 1923) des~ript~ion of the egg of aradus ,&lineatus, which has a distinct cap and chorial processes.
Very little is known under this heading. All the recorded specimens have been collected in association with termites, of which the following species have been identified. The hosts of Dr. Wheeler's Panama examples were kindly determined by Mr. Banks, those of the other Panama material by Dr. Snyder.- Termitaphis circumvallata Wasm.,
A mitermes foreli Wasm., Colombia.
Termitaradus mexicana (Silvestri), ,
Leucotermes tennis (Hag.), Mexico.
T. subafra Silv., . . Rhinotermes putorius Sjost. Africa.
T. australiensis (Mjob.), -
Coptotermes sp., Australia.
T. annandalei (Silv.), . . Coptotermes heimi Wasm., India. T. guiana (Morr.),
Leucotermes crinitus (Emerson), British Guiana. T. trinidadensidMorr.), . . . L. tennis (Hag.), Trinidad.




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268 Psyche [December
T. insularis (Morr.) ,... . . . . L. tenuis (Hag.), Trinidad.
T. panamensis n. sp., . . .. . L. tennis (Hag.), \
'Panama.
L. convexinotatus snYder,f
In view of the additional species which have been brought t,o light within recent years coincident probably with intensified study of termites and of termitophiles, it would be premature to say much about distribution of the family. At present Central America seems to be the centre of greatest abundance but this may be due to greater collecting in the region. The distribution is certainly however practically circumtropical. In many res- pects it resembles that of Peripatus (sens. lat.) and may, as in the case of that genus indicate considerable antiquity as Mjoberg has suggested.
Habitat notes of the previously d&-cribed species are scanty in the extreme. Of T. mexicana, Silvestri (1911) writes "in cuniculis nidi Leucotermes tenuis (Hag.)." When describing T. annandalei the same writer states "in nido Coptotermes Heimi Wasm., in trunco arboris (Ficus bengalensis) emortui et super solui sistentis exempla nonnulla Dr. N. Annandale legit." Mjoberg found a number of examples of his Queensland species "under bark of dead eucalyptus trunks in the colonies of a white ant (Coptotermes sp.) in the open forest country." Dr. Wheeler found the Panama specimens within a termite nest (Leucotermes convexinotatus Snyder) the Termitaphids themselves being close to the cambiunl of the tree trunk from which they might probably have extracted nourishment. They were running about fairly actively.
What little is known of its habitat therefore seems to suggest that Termitaphids may have the same feeding habits as the Aradidse.
What advantage they derive from living in the termite nest is uncertain. Termitaradus is probably protected from the termites themselves by its lateral laminae and their armature. Termitaphis exhibits a certain degree of physogastry, a well- known feature of termitophiles and one which might indicate this genus as the more specialized of the two, though in the



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19241 Systematic Position of the Family Termitaphididce 269 structure of the body margin it is intermediate between the Aradids and Termitaradus.
Dr. Wheeler suggests they may extract nourishment from either the nest material or the contained debris. Wasmann (1902) considers Termitaphis and Termitococcus Silvestri as affording the only known cases of trophobiosis among termites. Of Termitococcus nothing has been reported since the original description. As regards Term'itaradus the peculiar dorsal pores discovered by Morrison may possibly secrete some material attractive to the termites, but only field observations can decide this point.
Most of the species of the family have been described from females alone. Males are now known of Termitaradus amandalei, T. quianct (?), and 2'. panamensis sp. 11. Silvestri has given good figures of the male genitalia. Nymphs - have been found of T. annandalei and of T. panamensis only. Silvestri has figured the outline of the body of t'he ultimate and second (?) instars in T. amandalei. Both these instars have one lateral lobe on each side of the body more than in the imago (female) the numbers being 14 and 13 respectively. The same difference is observable in T. panamensis, between all female nymphs examined and the common 13-lobe type of female adult. Silvestri considers the additional lobe in the nymph to belong to the nietathorax but to the present writer it seems to correspond exactly to the extra lobe present in the adult females of some of the species and shown by Morrison to pertain to the mesothorax. The reduction of lobes has gone furthest in T. insularis (Morr.) in which the female possesses only twelve, the number present in the males of those species in which both sexes are known. TAXONOMY.
An examination of the unique and beautifully preserved type of Termitaphis circumvallata Tasm. shows it to be a female, of a different genus from all later described species. Wasmann's figures (1902) express these divergences quite clearly. It is a very different-looking insect with a swollen egg-shaped body surrounded by an incurved and upcurved dorso-lateral, seg- mentally-divided lamina almost meeting on the anterior half of



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270 Psyche [~ecem ber
of the body and showing the structure described in the key below. Some of the Panama material showed an upward curling of the laminae in alcohol, but the condition thus artificially produced was not even superficially similar to that in Termi- taphis. The Panama species is strongly flattened above and below, both in alcohol specimens and in life (Professor Wheeler), whereas Termitaphis would be rotund even were the laminae removed entirely. A new genus is therefore erected for the Panama species and for all other species described since T. circum- vallata. The Panama material was used as the genotype as it is the best known to me.
Silvestri's family diagnosis (1911, p. 232) may be modified by deletion of the phrase "corpus valde depressum," and by changing his statement regarding stigmata to read as follows: stigmata 9, of which two are thoracic. and seven abdominal. The two genera may be separated as follows: Termitaphis, Wasm. (1902, p. 105); Body egg-shaped, surrounded by a strongly incurved and upcurved, dorso-lateral segmentally di- vided lamina, the edges of which are furt'her divided into distinct, often quite distantly separated lobules each with a long fine almost smooth flagellum.
Type, T. circumvaUata Wasm.
Termitaradus, gen nov., Entire body strongly flattened above and below and surrounded by a flat lateral segmentally divided lamina the margin of which is crenulate forming short non- separated lobules, each provided with a short, circular, clavate or lanceolate flabellum with serrate edges. Type, T. panamensis, sp. nov.
In addition, the tylus, covering the setal coil, is in Ter- mitaphis strongly protuberant, while in the other genus it shares the general flattening of the body. In the structure of the ros- trum, antennae, legs and last ventral segments of the female the two genera are similar.
To Wasmann's original description of T. circumvallata may be added the following: marginal lamina on each side divided into 13 lobes (Wasmann did not count the minute 8th abdominal), bearing lobules as follows: 6, 3 (head), 9 (prothorax), 7 (meso- thorax), 7-8 (metathorax), 8-10) 8-9, 9-10, 8-9, 7, 6, 5, 3 (the 8



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19241 Systematic Position of the Family Termitaphididce 271 abdominal segments).
There are thus fewer head lobules than
in any other species of the family and more than the average of abdominal lobules. The flagella seem to me in most cases longer than figured by Wasmann. Both in the distinct separation of the marginal lobules and in the length and flagellate appearance of their appendages Termitaphis is clearly intermediate between the Aradid nymph described above and Termitaradus. The swollen form of the body is however very un-Aradoid, and may be best explained as an instance of termitophilic physogastry. THE SPECIES OF TERMITARADUS.
Eight species, including the new one described below may be referred to this genus.
Of these, one, namely T. australiensis
(Mjob.) is quite inadequately described and its relationships at present obscure.
The ihportant characters in the genus appear to be the form and average number of the flabella. It is therefore un- fortunate that these peculiar ~t~ructures are so easily detached. In their absence however, their number can be ascertained by counting the lobules in which they arise. The number of the lobes, at least in those species which show 13 or 14 on each side seems less reliable. One would of course have been inclined to regard the presence or absence of an additional lobe as a charac- ter at least of specific importance, but the following considerations have led the writer to reject it as such.- It is a single meristic character, and such are known to vary intraspecifically.
Nymphs in species in which the adult female is 13-lobed (T. annandalei and the normal 13-lobed Panama form) show the extra lobe clearly developed.
Specimens taken together in the case of three separate lots include a mixture of 13-lobe and 14-lobe examples. The specimens in these lots agree exactly in all other char- acters.
The males taken with these lots are all identical and all show 12 lobes, as does also the male of the typically 13-lobed T. annandalei.
The 14-lobed specimens have the lobes distributed as fol- ' lows in the ,manner indicated by Morrison.-2 to the head, one to the prothorax, 2 to the mesothorax, one to the metathorax and



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272 Psyche [~ecember
eight to the abdomen. Silvestri differs from this interpretation in considering the extra lobe (which by the way may be easily . distinguished by its smaller size) to belong to the metathorax in annandalei nymphs, and to the prothorax in the 14-lobed adults, while Morrison assigns it to the mesothorax. The writer agreed with Silvestri but a rigorous combined examination by Mr. Morrison and himself of material treated in different ways has led to the conviction that Morrison's interpretation is correct. The males differ from the females in that none of the ab- dominal segments after the seventh are furnished with lobes or form part of the marginal lamina. In number of flabella on head, thorax and segments 1 to 7 of the abdomen they agree with the corresponding female except that there is a tendency towards an average of one more flabellum in the abdominal segments. The following table shows the number of lobules and their flabella on one half of the body in the seven adequately known species, in the females only. The thirteen-lobed form of the Panama species has been taken as typical and the description founded on it alone. Should future work show that the four- teen-lobed form is specifically distinct there need then be no Head
Prothorax
-
Mesothorax
Metathorax
1 st abdominal
2nd
3rd "
4th "
5th "
6th "
7th ''
8th "




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19241 Systematic Position of the Family Termitaphididce 273 Key to the females of the seven adequately described species. ................
Only 12 lobes to body margin on each side. T. insularis Morr.
13-14 lobes to body margin on each side. ..
Lobules of 2nd to 6th abdominal lobes not more than four. T. trinidadensis Morr.
Lobules of 2nd to 6th abdominal lobes six or more. Flabella short and rounded, at most hardly more than twice as long as broad.
8th abdominal lobe with two lobules; anterior abdom. segments with normally 7 or more lobules on each margin. Lobules of 2nd to 6th abdom. lobes not more than 7; flabella rounded. ................ T. mexicana Silvestri. Lobules of 2nd to 6th abdom. lobes 8 or more; flabella short clavate. .................. T. annandalei Silvestri. 8th abdominal lobe with 3 lobules; anterior abdom. seg- ments with normally 6 or fewer lobules on each margin. T. guiance Morr.
Flabella elongate, much more than twice as long as broad Flabella lanceolate,
very acute at apex; 8th abdominal
lobe with 3 lobules .............. T. panamensis sp. n. Flabella subcylindrical, rounded at apex or at most very obtusely pointed; 8th abdom. lobe with 2 lobules. T. subafra Silvestri.
Termitaradus panamensis sp. nov.
Male, female: Colour of alcohol specimens pale yellowish. Very similar save in details, to the Indian T. annandalei Silv., from which it differs in the smaller number of lobules on the marginal lobes, and in the shape of the flabella, which are elon- gate and lanceolate, the broadest part being nearer base than apex, the apex itself being very sharply pointed. The dorsal
surface of the body is minutely papillated, marked into in- numerable polygonal areas (chiefly irregular hexagons) by lines of slightly larger papillae, and supplied with numerous pores. The tibia1 comb is similar to that described and figured by



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274 Psyche [~ecem ber
Morrison in T. guiance. The male genital segments differ from those of T. annandalei, especially in the caudal margin of the seventh abdominal segment, (mesad of the marginal lamina, see Silvestri, 1921, fig. 111, 4) which is far less sinuate in the present species.
Length: male, 2.35; female, 2.40 mm.
Holotype (a slide mount) Type Cat. No. 27855 U. S. National Museum.
Allotype (slide mount, 13-lobed form). U. S. National Museum. Paratypes in U. S. Nat. Mus. and in coils. Dr. W. M. Wheeler and Museum of Comparative Zoology.
Described from eight lots, with data as follows.- 5 males and 3 females, Barro Colorado Id., C. Z., Panama, 20th June, 1924, W. M. Wheeler. No. 510 (in nest of Leucotermes convexinotatus Snyder) ;
2 females, same locality, 21st Feb., 1924, T. E. Snyder. (in nest of Leucotermes tenuis Hag.)
3 females, same locality, 6th June, 1923, Zetek-Malino coll. (with Leucotermes tenuis on soft dry wood on ground) Z.2081 10 females, 2 males and 4 nyn~phs, near Fort San Lorenzo, C. Z., Panama, 14th June, 1923, J. Zetek, Z.2128A. (on soft wood of tree-stump) ;
3 females and two nymphs, same locality and date, J. Zetek, Z.2132A. (with Leucotermes tennis) ; 1 female (2.2171): In moist very soft rotting log on ground, Rio Aejeta, C. Z., Panama, Aug. 19th, 1923. With L. tenuis
and Cornitermes acignathus Silv.
J. Zetek coll.
1 female (2.2263 S): In branch on ground, hard wet wood, Sweetwater, Fort Sherman, C. Z., Sept. 7th) 1923. With L. tenuis. Zetek-Malino coll.
3 females and one nymph (z.2264 A) in pieces of branches on ground, hard wood.
Other data as 2.2263 S.
Termitaradus guianae (Morr .)
Among the material kindly lent by the U. S. Bureau of Entomology through the courtesy of Mr. Morrison, were six females and one male from Rio Frio, Colombia, collected by Dr. W. M. Mann in February, 1924. These have been referred



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19241 Systematic Position of the Family Termitaphididce 275 provisionally to T. guiance. The male has twelve marginal lobes on each side, while five of the females have thirteen and the sixth shows fourteen. These seem all conspecific in spite of the divergence in the number of lobes. In this connection reasons have been adduced above for rejecting this character as specific among thirteen-and fourteen-lobed forms. Unfortunately the flabella are almost entirely lacking in the seven specimens, but the few that remain (fig. 13.) are identical in shape with those figures by Morrison in T. guiance. Moreover the number of lobules in the respective lobes corresponds very closely with that in 2'. guiance, the chief differences being as follows: Number of lobules
I
T. guiana
Colombian
material
1 --------
The variation in the considerable series of T. panamensis examined supports the suggestion that these differences come well within the range of intraspecific variability. In any case they seem an insufficient basis for specific rank. In conclusion the writer would express his deep indebtedness to Professor W. M. Wheeler, Professor C. T. Brues and Mr. Nathan Banks for references to and loan of literature and for much helpful advice; and to Mr. Harold Morrison not only for turning over the task of describing his new species but also for much time, and patient work in demonstrating the interpretation of the segmentation.
2nd lobe of mesothorax
2nd abdominal lobe
5th abdominal lobe
Literature Cited.
4
6
6 .
CRAMPTON, G. C. 1923, Preliminary note on the termi- nology applied to the parts of an insect's leg. Canad.
Entom. vol. 55, no. 6, pp. 126-132, PI. 3.



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27 6 Psyche [~ecem ber
KNIGHT, H. H., 1923.
Miridae: in, Hemiptera of Connecticut
Conn. State Geol. and Nat. Hist. Survey, Bull. 34. M.J~BERG, E., 1914.
Preliminary description of a new rep-
resentative of the family Termit'ocoridee. Ent. Tidskr., Uppsala, vol. 35, pp. 98-99, 2 figs. MORRISON, H., 1923. Three apparently new species of Terrnitaphis. Zoologica, vol. 3, no. 20, pp. 403-408, PI. 24. PARSHLEY, H. M., 1921.
Essay on the American species of
Aradus. Trans. Am. Ent. Soc., vol. 47, pp. 1-106, PI. 1-7. REUTER, 0. M., 1912.
Bemerkungen uber mein neues Het-
eropterensystem.
Ofv. Finsk. Vet.-Soc. Forh., 54, A, no. 6, pp. 1-62. SILVESTRI, F., 19 11.
Sulla posizione sistematica del genere
Termitaphis Wasm. (Hemiptera) con descrizione di due specie nuove.
Portici Boll. Lab. Zool., vol. 5, pp. 231-236. ibid., 1920. Contribuzione alia conoscenza dei Termitidi e Termitofili dell'hfrica occidentale.
Op. cit., vol. 14, pp.
265-319.
(includes just a copy of the description and figures of T. subafra from Silvestri, 1911').
ibid., 1921. A new species of Termitaphis (Hemiptera-Het- eroptera) from India. Rec. Ind. Mus., Calcutta, vol. 22, pp. 71-74, 3 text-figs.
SPOONER, C. S., 1920. A note on the mouth parts of Aradida? Ann. Ent. Soc. Am., vol. 13, pp. 121-122, 1 text-fig. TOWER, D. G., 1913. The external anatomy of the squash bug, Anasa tristis DeG. Ann. Ent. Soc. Am. vol. 6, pp. 427-441, PI. 55-58.
WASMANN, E., 1902.
Species novae insect orum termitophi-
larum ex America meridionali.
Tijdschr. v. Ent., vol. 45, pp. 75-107; Termitaphis, p.. 105, PI. 9, fig. 7, 7a-c.
ibid., 1904.
Res. Swed. 2001. Exped. to Egypt and White Nile, 1901; no. 13. Termitophilen aus dem Sudan. Pp. 1-21 tlaf, 1. Upsala.
ibid., 1911. Die Ameisen und ihre Gaste. 1st Congr. Int. d7Ent. Bruxelles, 19lO.P~. 209-234, PI. 12-17.



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Psyche
[December
Figures.
Termitaradus panamensis sp. nov.
Outline of body of female
showing lobes and sutures.
Do. Rostrum projecting from body and viewed from behind.
Do. Lateral view of head showing rostrum, setal coil and insertion of left antenna.
Do. Setal coil, semi-lateral view.
Do. Portion of marginal lobe ( pronotum, dorsal). Do. Flabellum much enlarged.
Do. Tip of tarsus showing claws and arolia. Ctenoneurus hochstetteri (Mayr) . Rostrum. Do. One claw and its associated arolium. Dysodius sp., nymph. Portion of marginal lobe (pronotum, dorsal).
Lygus vanduzeei Knt. Claws, arolia and pseudarolia (after Knight).
Euschistus variolarius (Pal. de Beauv.). Tip of tarsus from below. The small circles are insertions of long spines. The arolia are united to the claws only at the base. Termitaradus guiance (Morr.) (Colombian specimen). Fla- bellum,




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