studies on the systematics of the berothidae, part i: a. redescription

STUDIES ON THE SYSTEMATICS OFTHE BEROTHIDAE,

PART I: A. REDESCRIPTION OFTHE ’GENUS SPHAEROBEROTHA NVAS,

WITH A CRITIQUE OFTHE TAXONOMIC CHARACTERS

USED IN THE BEROTHINAE (NEUROPTERA)

BY ELLIS G. MAcLEoD

INTRODUCTION

AS a by-product o.f the recent review by P. A. Adams and myself(NiacLeod and Adams, I967) of the subfamily classification of theBerothidae, new information has been uncovered concerning theidentity and taxonomic status of some of the more poorly knownmembers of this family. Because of the general difficulty in securingtype specimens, for study and because of the great paucity ot theseinsects in most collections, I do not consider it advisable to. undertake,at this time, the full, species-level revision which this family so badlyneeds. Nevertheless, sufficient data, are now at hand to encouragethe view that a long-term, piecemeal study of the more disorderlysections of the, family may ultimately produce a better taxonomicunderstanding o.f these insects.

A.ccordingly, the present series of publications has been initiatedfor the purpose of providing a vehicle for the redescription of keygenera and species, for the dis.cussion of the taxonomic utility ofvarious systematic characters, for the reevalua.tion of the validityand systematic position of the numerous genera of the Berothinae,and for the publication of such ecological and morphological informa.-tion as provides insight into the adaptive basis of certain evolutionaryspecializations. If successful, this approa.ch should eventually pro-duce at least a partial, generic-level revision of the family and shouldlay a better to.undation for a future., thoroughgoing revision of thespecies than exists at the present time.

The first paper of this series will be devoted to. a redescription ofthe poorly known genus Sphaeroberotha Naves and to a preliminarydiscussion of the relationships of certain genera of the Berothinae.In the course of this discussion I have set forth my views concerningthe usefulness of certa.in of the. generic-level taxonomic characters

1Department of Entomology, University of Illinois, Urbana, Illinois.Manuscript received by the editor December 2, 1967

342

1967] MacLeod Berothidae 343

which have been used in the past and have noted which additionalcharacters 1 feel should be carefully evaluated in future descriptiveand taxonomic work with this family.

The Genus. S1)haeroberothaNav{ts based his description of the type species of Sphaer’oberotha,

S. dumonti, on a single, female. Although he included the speciesCostachella yeyri (Esben-Petersen) in his new genu.s as a secondspecies, his characterization of Sl)haeroberotha was, in fact, basedalmost solely on S. dumonti since he had only a drawing, whose ac-curacy he questioned, o,f the wings of Esben-Petersen’s species forstudy. Through the. great kindness of Dr. S. Kelner-Pillault o.f theMus4um National d’Histoire Naturelle, Paris, I have been permittedto borrow and study the ho.lotype o.f S. dumonti from which thefollowing redescriptions and discussions are drawn.

8lhaeroberotha NavesSI)haeroberotha Naves, 1930, Broteria 26: 133-134. Type species (by

.original designation)" S1)haeroberotha dumonti Naves, 1930. Tjeder,1958, South African Animal Life 6: 281-282. MaeLeod and Adams,1967, Psyche 74: 254, 256, 258.

Description. A genus belonging to, the Berothinae. Face short,extending only slightly ventrad of eyes; vertex a slightly raised tri-angular area., with two weak lateral tubercles at its posterolateralcorners; postgenae moderately inflated. Scape only slightly elongate;antennomeres distal to scape subspherical, each with two. rings ofdistally directed setae which extend as far as the next antennomere;distal antennomere more elongate, acuminate. Mouthparts well de-veloped, mandibles wit’h tips crossing when jaws are closed. Pronotummoderately robust, slightly wider than long, with two transverse

furrows, strongly deflexed laterally. Fore legs lacking raptorialmodifications.Wings (Fig. ) elongate, with rounded tips, lacking any indica-

tion of falcation; Sc connected distally to R1 by an oblique crossvein a.t edge of pterostigma; two radial cross veins; free basal pieceof MA (Fig:. , b) oblique, slightly proximal to basal fork of MP;a weak series of outer gradate crossveins present. Fore wing" prob-able vestige of recurrent humeral vein present basally in costal cellas a steeply inclined .crossvein with a dis:a.1 fork; stems of M and Rfusing just proximal to basal subcostal crossvein; outer gradatecrossvein between the branches o.f MP generally in line with gradatesabove and below this vein, not far proximal to these; jugal vein and

344 Psyche [December

lobe absent; wi.h weakly developed lines o flexion present betweenMA and MP and between CuP and A; crossveins and torkingslacking dense tults o. erect setae; similar dense tufts trom trichosorsot wing margin also lacking. Hind wing: with C and Sc runningvery closely together or most o distance rom wing base to ptero-stigma; no basal crossvein in subcostal space; stem o NI passing be-neath R, but ree trom this vein to base ot wing; CuA parallelingposterior margin and reaching into distal /3 o.t wing, with numer-ous pectinate marginal torks; no trace ot CuP present; jugal veinabsent; a weak line ot flexion developed between bases o. NI andR, continuing a short distance distal to the. basal tree piece o.t MA..Male unknown. Female (Figs. 2, 3, 4): 7th sternite (Figs.. 2,

3, 7S) widely unsclerotized posteriorly and in midline, representedas two separated bilateral pieces, without the development o.t thin,rod-like lateral sclerites; 8th sternite (Figs. 2, 3, 8S) thin and strap-like laterally, with an anteriorly directed median lobe; 8th gono-coxites (Figs. 2, 3, 8gcx) represented as a small concave sclerite justanterior to, gonopore; 9th gonocoxites (Figs. 2, 3, 9gcx) large, withwell-developed, elongate hypocaudae; 9th gonapophyses lacking; othsternite (Fig. 3, oS) represented as a weakly sclerotized area dorsado 9th gonocoxites; genital chamber (Fig. 2. gen. ch.) large, withduct trom collaterial gland emptying into. it posteriorly, widely con-fluent with copulatory bursa (Figs. 2, 3, 4, co,p. b.), closed by an-teria.d appression ot membranous area between 9th gonocoxites againstlarge convex lobe ot weakly sclerotized cuticle just po.steriad o 8thsternite; copulatory bursa, somewhat small, consisting ot membranousdistal portion and a well-sclerotized, coiled proximal area rom whichthe spermathecal duct originates; spermathecal duct. (Fig. 4, sp. d.)elongate and looped; spermatheca (Figs. 2, 4, sp.) intricately coiled,sclerotized, with a prominent spherical diverticulum and with ap-parent cuticular attachments or glandular epithelium along one sidejust proximal to origin ot ertiliza.tion canal; tertilization canal lack-ing usual ringe o cuticular attachments tor glandular epithelium(Fig. 4, fert. c.).Type species" Sphaeroberotha dumonti Naves.

Sphaeroberotha dumonti NavesSphaeroberotha dumonti Navfis, 1930. Broteria 26: 134-135, fig. 44. De-

scribed from a single female now located in the collections of the Mus6umNational d’Histoire Naturelle, Paris, MaeLeod and Adams, 1967, Psyche74 254.


Description. Face and vertex dark yellow, with stout whitesetae of moderate length and with a suggestion of dark spotting aboutthe bases of the vertex setae; labrum with a transverse row of thin-ner, more-elongate setae. Scape short, twice as long as broad, setaelike those of vertex but darker, right antenna with fifty-six anten-nomeres distal to scape, each with a thin brown ring, setae of a darkamber color. Thorax tannish brown (probably much discolored),with no indications of any original mottling of color pattern, Pro-thoracic legs pale yellow, femora and tibiae with a few scattereddarker spots.Wings (Fig. x) with anterior margin of fore wing slightly con-

cave; no maculations on membrane, but with longitudinal veins offore wing with dark dots at setal bases; outer gradate series with 4crossveins in the anterior wing, 3 crossveins in the posterior wing.;fore wing length 6.7 mm.Thorax and wing margin with sparse elongate white setae (prob-

ably mostly rubbed off), posterior wing margins with more-elongate,silky pale yellow seta.e; setae on wing veins white, shorter than thoseon body, directed basally at about a. 45 angle to longitudinal axisof vein, length about equal to, average distance between veins; noevidence of squamae on bo.dy, legs, or wings.Abdominal structures as in Figs. 2-4: 8th sternite with short setae

and with a.n apodeme reenforcing the strap-like lateral arms, an-teriorly directed median lobe slightly pedunculate in lateral view,somewhat spatulate in shape when viewed ventrally; cuticle between8th sternite and 8th gonocoxites a weakly sclerotized convex lobe;9th gonocoxites somewhat triangula.r in lateral aspect, with a. distinctposterior lobe. Sclerotized, proximal portion of bursa wound in asnail-like coil of decreasing diameter, from the tip of which thespermathe.cal duct originates; spermatheca with coiling pattern as inFig. 4.The pinned type is missing its left antenna distal to, the scape,

right fore leg distal to the femur, mesothoracic legs distal to, theircoxae, metathoracic legs distal to the coxa (right) and femur (left)and mo,st of the left hind wing. The abdomen has been removed,cleared, and placed in glycerine in a microvial on the pin with thespecimen. The holotype was collected at a light in May, x9:7 byC. Dumo.nt at Nefta, Tunisia.

DiscussionSuch special,ized features as the large., anteriorly directed 8th

sternum, well-developed hypocaudae, manner of closure of the genital

PSYCHE, 1967 VOL. 74, PLATE 37

pt

cop. b. fert.

:c/’0.5

MACLEOD- BEROTH1DS


chamber, and elongate wings show that SI)haeroberotha quite properlybelongs to the Be.rothinae. Within this group, however, its systematicposition is far from. clear. Indeed, because of the poor state o.f ourknowledge concerning many other genera of the Berothinae, it is noteven certain that Sphaeroberotha represents a valid genus.

Along with the berothines Protobiella Tillyard, SpermophorellaTillyard, Nodalla Naves, Costachillea Nawls, and CycloberothaNavfs, Sphaeroberotha shares the feature of a rounded outer wingmargin. However, as the members of the other three subfamiliesthe Berothidae share this same characteristic, by itself such a. similar-ity shows nothing about the relationships of these genera of theBerothinae. These six genera do, however, possess certain othersimilarities and, as they have, to varying degree.s, been treated asa unit in previous taxonomic works, they will be. considered togetherhere.

Prior to his description of S/)haeroberotha, Navels himself (1929)separated the first three of these genera from the other two by thepresence of a fu.sion between Sc and R1 in the region of the pteros-tigma which was presumed to be lacking in Costachillea and Cy-cloberotha (and in the subsequently described St)haeroberotha) where

EXPLANATION OF PLATE 37Drawn from the female holotype of Shaeroberotha dumonti Navels in

the Museum National d’Histoire Naturelle, Paris. All from camera lueidatracings. Fig. 1. Right fore- and hindwing. Figs. 2 and 3. Lateral andventral aspects of the tip of the abdomen. (Setae indicated only on 8th-ster-nite, 9th gonocoxites, and hypocaudae. Right hypo.cauda omitted from Fig. 3.)Fig. 4. Copulatory bursa and spermatheca, dorsal aspect with posterior enddirected toward top of page.

Abbreviations: a--anus; 1A, 2A, 3Am 1st, 2nd, 3rd anal veins;b--basal free piece of anterior median vein; coll. gl. mcollaterial glandduct and opening; cop. b. copulatory bursa; Cu--base of eubital vein;CuA-- anterior cubitus; CUP-- posterior eubitus; left. c. fertilizationcanal; 8 gcx, 9 gcxSth, 9th gonoeoxites; gen. ch.--genital chamber;gl? possible cuticular canals from spermathecal gland cells; gpr--gono-pore hye hypocauda M base of median vein MA anterior medianvein; MP-- posterior median vein; ptpterostigma; Rl--first branchof radial vein; Rs--radial sector; rhvmvestige of recurrent humeralvein; 6S, 7S, 8S, 10Sm6th, 7th, 8th, 10th sternites; Sc--subcostal vein;sp--spermatheca; sp.d. mspermathecal duct; 6T, 7T, 8T6th, 7th, 8thtergites; 9T -t- ectmfused 9th tergite and eetoproct.

’Navfis further separated these three genera by placing Protobiella andSpermoI)horella in one tribe, the Spermophorellini, and Nodalla in a secondtribe, the No.dallini. No diagnostic features by which these two tribes mightdiffer are apparent from his descriptions; indeed the Spermophorellini arenot even included in his key to tribes (pp. 17-18) in his revision of 1929!


these veins are united by a short crossvein. This oppo.sing set ofcharacters has been much used in the supraspecific taxonomy of theBerorhidae, although there are several complicating features of thisarea of the wing which undermine the usefulness of this. set as ataxonomic character.

For example, there are’ many cases in which the approach of Scand R1 is so close that the decision as to. whether these, veins areactually fused or not becomes arbitrary. A more subtle source ofco,nfusion arises from varia.tions in the degree of obliquity of the cross-vein, so that in some cases an apparent fusion of Sc and R1 occurswhen in reality these veins are quite separate. The ease with whichthis condition can mimic a fusion o.f these veins is further compoundedby the fact that the true termination of Sc is often masked in thethickening and pigmentation of the adjacent pterostigma. In addi-tion to these complexities, this set of taxonomic chara.cters is alsooften subjected to a high degree of individual variability. I feel,therefore, that the degree of fusion of these veins, is of a. limitedutility and that the use of this feature at the tribal level can prob-ably not be justified. On this basis, then, the genera Protobiella,Spermophorella, and Nodalla cannot be. excluded from considerationin ascertaining the relationships o,f Sphaeroberotha.

Costachillea hs been considered to differ from Sphaeroberotha andCycloberotha by the presence of an "internal radial vein" (i.e. thevein labeled b in Fig. of the present paper) in t’hese latter twogenera, which is presumed to, be lacking in Costachillea. Here again

3Because of the rather far distal position of the basal piece of MA inmany berothids, it might be considered that this vein is a true crossvein,the real basal piece having been lost with the close approximation of thestems of R and M. This possibility seems to be strengthened by the fact thatin the fore wings o.f several genera (Trichoberotha and some species ofLornamyia show this especially well) there does seem to be an additional,more proximal vein between the nearly adjacnt bases of R and M at thelevel of the m-cu crossvein, which is the more usual location of the basalpiece of MA in most other Neuroptera.

feel, however, that this more proximal crossvein is a secondary develop-ment in those genera where it is present and that the true basal piece ofMA is really the more distal vein. My principal reason for this conclusion isthat I am unable to find a second, more-proximal vein in the fore wings ofsuch generalized berothids as Cyrenoberotha and Rh.achiberotha, where thebases of R and M are rather widely separated and where such a vein, ifpresent, should be easily seen. Again, in the even more generalized ber.othidrelatives of the Neurorthidae and Dilaridae, where the relationships of theveins at the base of the fore wing are equally well made out, only a singlevein is present between these longitudinal veins, which in the nallachiine


one can, I feel, maintain strong reservations about the importance o.fthis set of a.lternatives as a taxonomic character. The degree of de-velopment of the basal piece of MA is subject to a good deal ofvariation, largely because of the weakening of its sclerotization whichresults from the line of flexion passing through it. This difficulty indetecting the vein is compounded when the vein is locat:ed in arelatively more pro.ximal position between the approaching bases o.fR and M.4 As a result of these considerations, I believe that it wouldbe premature to. assume that Costachillea cannot be closely related(or perhaps congeneric) with Sphaeroberotha until, at a minimum,specimens of Costachillea have been examined to see if perhaps apoorly sclerotized basal piece of MA is not present after all whichwas missed by Nawis.The final genus of bero.thine’s with rounded wing apices which

should be considered is Cyclobe’rotha. The characters which haveseemed to separate this genus from Sphaeroberotha are the number o.fradial crossveins, the number of branches of the radial sector, andthe presence or absence of squamae on the females. With respect tothe number of radial cro,ssveins, Na.vs (929, I93O) has stated thatCycloberotha is characterized by the possession of three. Althoughthis is true for the type species, C. rnjobergi (E,sben-Petersen), Nav,shas. erred in evaluating this feature in the seco.nd species of this genus,C. neuropunctata (Esben-Petersen), where only two radial cross-veins are present. Esben-Pe.tersen (I917) states this quite clearly inhis original description of this species and I have verified this factfrom the examination o.f a series of specimens of this species in mycollection. The. number of radial cro.ssveins is frequently rather con-stant within groups of closely allied species and it may be that C.neuropunctata has been incorrectly assigned to Cycloberoth.a. Prece-dent, however, exists in both Berotha a.nd in the. rather well-studiedLorn.amyia for including in the same genus species with differing num-bers of radial crossvein.s so that C. neuropunctata should not beexcluded fro,m Cycloberotha on the basis o.f this feature alone. In any

Dilaridae is relatively distal in position and is slightly sinuate as it is inthe fore wings of many berothids. Unfortunately, comparisons with theclosest relatives of the Berothidae, the Mantispidae, cannot be drawn, sinceeven in the most generalized of the Platymantispinae, the bases of R and Mof the fore wing are too closely adiaeent to permit study.

4In evaluating Naves’ descriptions an additional, essentially semanticproblem is added to these sources of confusion: a vein is only an "internalradial vein" if it is located proximal to the fork of MP. Where the veinis located distal to this fork it no longer qualifies as an "internal r,adial vein"and the taxon is described as lacking the vein (el. Navels, 1929, p. 23).

3 5o Psyche [December

case, attempts to separate Sphaeroberotha and Cycloberotha on thestrength of this single character is made impossible by the presentheterogeneous composition of Cycloberotha.

The number of bra.nches of the radial sector is likewise unsuitableas a criterion for distinguishing these two genera, since C. neuro-punctata and S. dumonti both have. an identical pattern of branchingof the radial sector. It is true that the type specimen of C. mjobergihas one fewer branch in the radial sector of the fore wing, but ashas been recently noted (MacLeod and Adams, 1967) slight dif-ferences in the number of branches of the radial sector are probablytoo variable a feature to use as a taxo,nomi.c character of genericdistinction. The. presence of squamae on the females of Cycloberothais likewise an insufficient reason for separating these two. genera, sinceotherwise closely related species are known in both Berotha andLomamyia which differ in the presence or absence of these squamaeon their females.

With the present state of our knowledge, concerning the generapresumed to be somewhat related to Sl)haeroberotha, it is not possibleto more than guess at what the. diagnostic features of the genus mightbe, assuming that it is actually a valid entity. Certainly none of thetaxo.nomic characters discussed above is sufficiently free from sus-picion to serve as a guide, in this direction. As a tentative guess, sinceI have not o.bserved them in the form seen in Shaeroberotha else-where in the family, I might suggest that the sclero.tized, "snail-shell"specialization of the proximal portion of the copulatory bursa and thepresumptive glandular elements asso.ciated with the distal portion ofthe spermatheca (rather than in their more usual position along thefertilization canal) may perhaps prove of diagnostic importance.There is a similar sclerotization of this po.rtion of the copulatorybursa in Cyrenoberotha o.f the. distantly related Cyrenoberothinae, al-though here the sclerotized portion is not coiled and of course thereare many other differences. Again, in the somewhat more closely re-lated Naizema (but not in Trichoberotha), there is a tendency for aweak sclerotization of this portion of the copulatory bursa (MacLeodand Adams, 1967). Here again, however, there is no tendency for acoiling of this sclerotized portio.n. Finally, Trichoberotha (to be re-described in a subsequent paper of fhis series) has a superficiallysimilar, snail-like coil in the sperm-conducting pathway; however,here it is the spermatheca itself, rather than a p.ortion of the copulatorybursa, which shows the apparent similarity. T’here are, in addition,

1967] 34acLeod Berothidae 35

ma.ny other important differences between Sphaeroberotha and Trich-oberotha.The foregoing analysis, which I regret has been unable to settle

any question as to the position or validity of Slhaeroberotha, em-phasizes the great need for a better understanding of the detailedstructure of a large segment of the species and genera of theBerothinae. Future descriptive work must carefully re-evaluate theeffectiveness of the taxonomic characters which have been used in thepast and must make available the condition of characters whose im-portance has usually not been properly appreciated up until now. Inparticular, I feel that among the more important features tor thecharacterization of valid genera are the structure of the male andtemale genitalia and o.f the internal, sperm-conducting and storagesystem of the female. Such details of the male as the degree of de-velopment o.f the gonarcus and gonocoxites and the basic form andlength of the mediuncus and its terminal setae will probably be ofgreat help in this attempt. In the female, the size and shape of the7th and 8th sternites, the degree of development of hypocaudae, thepresence or absence of 9th gonapophyses, and, especially, the structuraldetails of the copulatory bursa and spermatheca should also be care-fully evaluated.

LTERAT’VE Cn’EDKRb’c, L.

1922. Berothidae. Beitrige zu einer Monographie der Neuropteren-Familie der Berothiden. Stett. Ent. Zeit. 83: 49-88.

1923. Neuroptera suecinica baltiea. Die im baltisehen Bernsteineingeschlossenen Neuropteren des westpreussischen Provinzial-Museums in Danzig. Stett. Ent. Zeit. 84: 62-92.

ESBEN-PETERSEN, P.1917. Australian Neuroptera. Part III. Proc. Linn. Soc. New South

Wales 42: 203-219.MAcLoD, E. ANn P. ADAMS.

1967. A review of the taxonomy and morphology of the Berothidae,with the description of a new subfamily from Chile (Neuroptera).Psyche 74: 237-265.

Through the courtesy of Dr. O. S. Flint, the berothids of the Neuropteracollection of A. Alfieri, recently acquired by the U. S. National Museum,have been loaned to me for study. The collection contains specimens ofNodalla, including the holotype of N. aegyptiaca Navels, the type speciesof this genus. Preliminary study of these specimens reveals the same basicpattern of wing venation and female genitalia as Sphaeroberotha andindicates a probable synonymy of these two taxa. This material will bedescribed in a future paper of this series.


NAVES, L.1927. N6vroptres d’Egypt et de Palestine. me partie. Bull. Soc. Roy.

Entom. d’Egytpe 1926: 192-216.1929. Monografia de la familia de los Berotidos. Mere. Acad. Ci.

Zaragoza 2: 1-106.1930. Insectos del Museo de Paris. 6" Serie. Broteria (Set. Zool.) 26:

120-144.

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