G. S. Tulloch.
The Proper Use of the Terms Parapsides and Parapsidal Furrows.
Psyche 36(4):376-382, 1929.

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Psyche
[December
THE PROPER USE OF THE TERMS PARAPSIDES
AND PARAPSIDAL FURR0WS.l
The terms parapsides and parapsidal furrows are very frequently misapplied by morphologists and systematists working with Hymenoptera. An examination of the litera- ture reveals the fact that the term parapsidal furrows has been used to designate either of two pairs of longitudinal furrows that may be present on the mesonoturn of certain Hymenoptera, viz., certain Xiphydriids, Ampulicids, For- micids (pf and no of Figs. 1, 2, 4, and 5). Likewise, the term parapsides has been used to designate the lateral re- gions delimited by either pair of the so-called parapsidal furrows (pa and sc of Figs. 1, 2, 4, and 5, I1 of Fig. 3). Since it is quite obvious that two distinct pairs of furrows or two distinct sets of regions cannot have the same terms applied to them, it may be of interest to persons working with Hymenoptera to have the correct application of the terms re-established as intended by their author. Moreover, as these characters are diagnostic and widely used in the classification of Hymenoptera, it is quite essential that the correct terms should be brought to the attention of present workers so that their usage may be stabilized and unneces- sary confusion eliminated in future literature. The term parapsides was first used by Macleay, 1830, (page 148, footnote 1) to designate certain lateral regions occurring on the mesonoturn of Polistes billardieri Fabr.2 Naturally the furrows delimiting these regions are the parapsidal furrows although Macleay does not specifically designate these furrows as such. Figure 3 of this paper is 1 Contribution from the Entomological Laboratory of the Bussey Institu- tion, Harvard University, No. 323.
2 Dalla Torre, 1904, considers P. billardieri Fabr. a variety of P. crinitus Felton.




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19291 Use of Terms Parapsides and Parapsidal Furrows 377
a portion of the mesonotum of P. billardieri Fabr. bearing the parapsides and parapsidal furrows taken from Mac- leay's original figure and is enlarged to twice the size of his figure. In his explanation of the plate he designated the parapsides of the rnesonotum with the Roman numeral 11. Since it is perfectly clear from his figure as well as from his description (page 148, footnote 1) that he intended the term parapsides to be applied to the lateral regions of the mesonotum delimited by the longitudinal (parapsidal) fur- rows, it is logical that all homologous regions and furrows in other Hymenoptera should be similarly designated as it 3
0 n sc try) P^ 5& Pa sc
Fig. 1. Portion of the mesonotum of Xiphydria. Fig. 2. Portion of the mesonotum of Ampulex. Fig. 3. Portion of the mesonotum of Polistes billardieri Fabr. Fig. 4. Portion of the mesonotum of Myopopone castanea muculata Roger. Fig. 5. Portion of the mesonotum of Paraponera cluvata F. Sm. is one of the important principles of comparative morphol- ogy to apply the same term to the homologous anatomical parts in different groups.
If all Hyrnenoptera had a mesonotal structure similar to that of P. billardieri Fabr. the matter would have been much simplified and no confusion would have resulted in the litera- ture. However, in certain Hymenoptera (Xiphydria, Am- pule~, Paraponera, Figs. 1, 2, and 5) another pair of longi- tudinal furrows appears extending backward from the ante-



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378 Psyche [December
rior margin of the mesonoturn and converging more or less in their course. It is these furrows that have been erro- neously identified by some workers as parapsidal furrows for years; likewise, the lateral regions delimited by them and the true parapsidal furrows have been erroneously identified as parapsides. Since the correct term which should be applied to these furrows is little known, it may be of interest to review briefly the terms that have been applied to them, and, also to indicate the term which should have priority over the others as determined by its commonly accepted and correct usage in the literature. Morphologically these furrows which extend backward from the anterior margin of the mesonoturn a varying dis- tance and at a varying angle of convergence have been con- sidered to form the lateral limits of the prescutum when they converged sufficiently to fuse into a single median line. They have been correctly referred to as "prescutal sutures" by Crampton, 1926, In such cases as Paraponera (Fig. 5) they clearly limit the prescutum, but in many forms like Ampulex (Fig. 2) they open out and approximate parallel lines. It is obvious in such cases that the preseutum is not ciearly delimited and that it fuses with the scutum to form the general region prescutum plus scuturn.1 Although, as just stated, the failure of these furrows to converge and fuse makes it impossible to determine the posterior lateral limits of the prescutum; their presence, if even of short length extending backward from the anterior margin of the mesonofcum, indicates at once the lateral limits of the prescutum in the anterior region. In some forms such as Polistes these furrows are absent (Fig. 5) and in a similar case it has been assumed by a recent worker that the pre- scuturn extends laterally to the parapsidal furrows. An examination of the immature pupal stages of Polistes re- veals the presence of these prescutal sutures which gradu- ally disappear as the chitin hardens before attaining the adult condition. In at least one species of Polistes 1 The writer is cognizant of the fact that when the furrows con- verge and fuse, Snodgrass, 1926, does not consider the region delim- ited by them as the prescutum but as a part of the muturn. The writer does not on the basis of the evidence presented accept this view.



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19291 Use of Terms Parapsides and Parapsidal Furrows 379 the prescutal furrows are present in the adult. In Pepsis, a somewhat similar form, the mesonotum is, at first glance, composed of a large central area bounded laterally by the parapsidal furrows. A closer examination of the mesonotal surface reveals the presence of a pair of short indistinct furrows extending backward from the anterior margin. These are the remains of the prescutal furrows and delimit the anterior lateral borders of the prescutum. From the foregoing it is quite evident that the huge, apparently un- differentiated area present in Polistes and Pepsis is a com- bination of prescutum plus scutum and in no case does the prescutum extend laterally to the parapsidal furrows. Mayr, 1861, was the first to mention these furrows which morphologically limit the prescutum in some forms and in his early writings applied to them the indifferent term "con- vergirende Fwchen." Later, 1878, he identified them erro- neously with the furrows of Macleay calling them Purap- sidenfurchen. Kokouyew, 1898, designated these furrows as notauli. Emery, 1900, noted the misinterpretation of Mayr and designated the central furrows as Mayrian furrows. Morley, 1903, used the term notauli in the same sense as Kokouyew. Schmeideknecht, 1907, uses the term notauli for the central furrows but gives it as a synonym of Parap- sidenfurchen which is obviously incorrect. It may be seen from this brief sketch of the terms applied to these furrows that convergirende Furchen, notauli, and Mayrian furrows are synonymous. Since the term notauli has been used more widely than the other two terms and is perhaps the most appropriate of the three because, by derivation, it means "hollows on the back," present workers who have not already adopted this term to indicate the central furrows should do so and thus aid in avoiding any further confusion in the literature.
Some very clear distinctions between parapsidal furrows and notauli are evident upon an examination of different Hymenoptera. Both pairs of furrows are extremely vari- able in character, yet the following generalizations may be made. The parapsidal furrows may be present and the notauli absent, or vice versa, and occasionally in some forms both parapsidal furrows and notauli may be absent. In



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380 Psyche [December
forms such as Xiphidria (Fig. 1) the parapsidal furrows extend from the transcutal suture of the mesoscufcum to the anterior margin of the mesonotum. In forms like Para- ponera (Fig. 5) they extend only halfway to the anterior margin of the mesonoturn. Other gradations may be noted by simply examining other forms which always exhibit the tendency of the furrows to be lost in the anterior region and to be retained in the posterior region of the mesonotum. This would seem to indicate that the parapsidal furrows ex- tend forward from the posterior region of the mesonoturn or from the transcutal suture of the mesoscutum a varying distance in different forms. Another feature of the parap- sidal furrows is that they usually extend forward parallel to each other (Figs. 4 and 5) or they diverge (Figs. 1 and 2) and rarely, if ever, have their terminal points at a posi- tion which is convergent from the assumed starting point at the transcatal suture. In the form of Polistes (Fig. 3) figured in this paper the.parapsida1 furrows appear to con- verge though their terminal points are at a position which is divergent from the points of intersection of the parap- sidal furrows and the transcutal suture. The distinctive characteristics of notauli are to some degree the converse of the characteristics of parapsidal furrows. In forms like Paraponera (Fig. 6) and Xiphydria (Fig. 1) they converge backward from the anterior margin of the mesonotum, and fuse to form a median line. In forms like Ampules they extend backward in a course closely approximating parallel lines but do not.reach to the transcutal suture of the meso- scutum. In forms such as Myopopone (Fig. 4) they extend but a short distance backward and in Polistes (Fig. 3) they are absent. From the above facts the following generaliza- tions may be made: notauli apparently extend backward from the anterior margin of the mesonotum a varying dis- tance and at a varying angle of convergence, occasionally approximating parallel lines in their course but rarely, if ever, diverging from their assumed starting points at the anterior margin of the mesonotum.
Occasionally in some forms such as Xiphydria. (Fig, 1) a median line (me of Fig. 1) is present extending longitudi- nally along the central region of the mesoscutum. The pres-



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19291 Use of Terms Parapsides and Parapsidal Furrows 381
ence of this median line is of minor significance since it is used in the classification of only one group of Hymenoptera. However, its presence along with the notauli and parapsidal furrows illustrates the complete number of longitudinal lines of furrows that have been observed on the mesonotum of Hymenoptera.
Much of the confusion existing in the literature has re- sulted from the mistaken impression that the terms parap- sidal furrows and notauli are synonymous. From the fore- going discussion it should be clear that this is not the caxe, since they are two distinct pairs of furrows, each pair so completely localized in position and possessing such distinct characteristics that misidentification should be impossible. The terms discussed may be defined as follows: 1. Parapsidal f urrows-longitudinal furrows extending anteriorly from the posterior region of the mesonotum or from the transcutal suture of the mesoscutum and varying in length and in their course from one approximating paral- lel lines to one which is strongly divergent but very rarely, if ever, converging from the original point in the posterior region of the mesonotum.
2. Parapsides- (sing. parapsis) lateral regions delim- ited by the parapsidal furrows and the tegulae. 3. Notauli-(sing. notaulus) longitudinal furrows ex- tending posteriorly from the anterior margin of the meso- notum, varying in length and in angle of convergence, and occasionally approximating parallel lines but rarely, if ever, diverging in their course from the anterior margin of the mesonotum.
Crampton, G. C. 1926. A phylogenetic study of the thoracic sclerites of the Psychodoid Diptera with remarks on the inter-relationships of the Nematocera. Ent. News, vol. 37, p. 33.
Dalla Torre. 1904. Vespidse, Gen. Inc., Fasc. 19. Emery, C. 1900. Intorno a1 Torace delle Formiche. Bull. Soc. Ent. Ital. 32: 1-17, 14 figs.




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382 Psyche [December
Enteman, W. 1904. Coloration in Polistes. Carnegie Insti- tution of Washington. Pub. No. 19.
Kokouyew, N. 1928. Fragments Braconologiques. Horae SOC. Entom. Rossicae, vol. 32, p.
Macleay. 1830. Explanation of the comparative anatomy of the Thorax of Winged Insects. Zool. Journal. Vol. 5, pp. 145-179.
Mayr, G. 1861. Die europaeischen Formiciden. Wien. 1887. Sudamerikanische Formiciden. Verh. zool-bot. Ges. Wien, vol. 37, pp. 511-632.
Morley, C. 1903.
The Ichneumons of Great Britain. Keys.
Plymouth.
Schmiedekneeht, 0. 1907. Die Hymenopteren Mitteleuropas. Jena, G. Fisher.
Snodgrass, R. E. 1910. The thorax of the Hymenoptera. Proc. U. S. Nat. Mus. No. 1774, vol. 39, pp. 37-91. Wheeler, W. M. 1910. Ants, their Structure, Development and Behavior. New York, Columbia Univ. Press. me. median line.
no. notauli.
pa. parapsides.
pf. parapsidal furrows.
pre. prescutum.
sc. scutum.
11. parapsides (Macleay's
original designation)
Figure 1. Portion of the mesonotum of Xiphydria. Figure 2. Portion of the mesonotum of Ampulex. Figure 3. Portion of the mesonotum of Polistes billardieri Fabr.
Figure 4. Portion of the mesonotum of Myopopone cas- tanea nzaculata Roger.
Figure 5. Portion of the mesonotum of Paraponera clavata F. Sm.




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