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Psyche 7:235-240, 1894.

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PSYCHE.
ON A RATIONAL NOMENCLATURE OF THE VEINS OF INSECTS, ESPECIALLY THOSE OF LEPIDOPTERA.
BY A. S. PACKARD, PROVIDENCE, E. I.
Hitherto there has been an unfortu-
nate lack of uniformity in the nomencla- ture of the veins of the wings, different names having been applied to the veins
of different orders.
In his paper on the phylogeny and
ontogeny of the veins of the wings of
Lepidoptera Spuler has, however, given
us a simple scheme and a nuonbering of
the veins which will, we think, apply
in general to the wings of insects of all orders.
Redtenbacher had previously pointed
out that the geologically older
Orthoptera and Neuroptera have a
much I icher and more complicated
venation than the Coleoptera, Lepi-
doptera, Hyn~enoptera and Diptera ;
thus among the Rhynchota, the oldest
forms, the Cicadidae and Fulgoridae
have a much greater number of veins
than the Hcrniptera. There is no
doubt but that the oldest insects were
provided with an excess of veins, that
on the other hand in the course of
development this superfluity has dis-
appeared by a process of reduction,
and in this way a simpler system of
venation has resulted. It is also to be
observed that the size of the wings has
had a considerable influence on the
number of the veins, since small forms
almost without exception have fewer
veins than insects with large wings."
Redtcnbacher also believes bb that the
normal type of a differentiated wing
may be found in those insects whose
fore and hind wings are most similar in
sim and shape," and states that the
venation is not useful as an ordinal
character, but is of more seivicc in
separating suborders and families.
We agree with Spuler in rejecting
Reclicnbacher's system, which is partly
based on Adolph's untenable theory of
convex and concave veins, but more
especially for the reason that Redten-
bacher assumes that the primitive
form of venation is that of the Ephe-
meridae. He remarks : "There is
scarcely another group of insects whose
wings show the primitive type, the
fan-shaped form, as the May-flies." It
may bc objected to this that the
Ephemeridae, though in most respects
generalized and primitive insects, yet
are, as regards the wings, highly
modified or specialized. That this is
the case is also suggested by the reduc- tion or atrophy of the month-parts.
Pucht 7 235.241 tprc.1903). lu~fdlpsychmilclub org/W-2JS html



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236 ps ycffl?. [May 1895.
On the other hand the retention of
sexual organs paired throughout, the
ducts remaining separate, with open,
paired outlets, shows that the May-flies ale, in this respect, more primitive
than any other winged insects. But as
regards the thorax and the 'wings, we
observe that in them a high degree of
modification has taken place. Thus
the two pairs of wings are very unlike
in size and shape, and this feature is
a secondary one. Hence the large
number of main longitudinal veins in
the wings of Ephemera is a case of
irrelative repetition of parts mostly
situated in the fan-like field, due to a process of specialization, a process
which is manifested in quite another
way in the wings of the Dermaptera,
also a primitive type. Redtenbacher
regards the eleven longitudinal veins
(I-XI) of Ephemerids as the normal
number, and considers that the
Trichoptesa, Lepicloptera, ctc., have
lost certain of the veins by a
PI-ocess of reduction. This view has
been adopted by Comstock in his mg-
gestive paper, å´ Evolution and Tax-
onomy," but it seems to us to be
untenable, the anal field (" faltentheil'' of Spuler) not being of primary impor-
tance. On the other hand Redten-
hacher's use of Roman numerals for
the main veins, and of a combination
of Roman and Arabic numerals for
their branches, is very convenient.
Spuler divides the wings of each pair
into an outspread poslion (Sfreiten-
theil), and a folded part (Palten-
theil).
The veins of the former area
he numbers in thc same manner as
Redtenbacher, beginning on the costal
edge of the wing, while those of the
folded area (the submedian and internal
or first and second anal veins of other
authors) he does not name, but simply
numbers with Greek letters P. He
considers that Hagen was right in
believing the Phryganidae, Tipulariae
aiid some Microlepidoptera to be Forms
with a schematic, /. e., primitive vena- tion (Stettin. Ent. Zeit., p. 316, 1870). Spuler shares the opinion of Fritz
Muller (Tennitidac), Braner and Red-
tenbacher (Libellnlidae) , and Haase
(Papilionidae), that the costa is only a hypodermal structure, a thickening of
the edge, which does not have a
trachea as its origin (adage), and
which therefore has nothing to do
with the veins.
Spuler also shows that the venation
of the Orthoptera, especially their most generalized form Blatta, is fnnda-
mentally nearly identical with that of
the Lepidoptera, veins I-V being
readily homologized with those of the
latter group ; so also with the most
generalized Hemiptera (Fulgora. Fig.
I). We may also draw attention to
Fig. I.
the remarkable resemblance in the
venation of the generalized Psocid
genus Amphientomum, which at first




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sight, from the shape and size of the
wings, reminds one of a Micropteryx
or Eriocephala, while it also has a few
scales like those of these moths.
But that the system of venation of
Spuler is morphologically the correct
one is fully and satisfactorily proved
by the ontogenetic development of the
veins. Fritz Miiller (Kosmos i, p. 390)
was the first to examine the incipient
venation of two semi-pupal moths
( Castnia avdaltu). He observed that
in the immature pupa the cross veins
were wanting", and that different longi- tudinal veins, which afterwards more
or less completely disappeared, were
present, and hence he regarded the
pupal venation as the primitive one.
This view Spuler has adopted and
extended, and it plainly enough, sup-
ported by the researches of Brauer and
Redtenbacher on the venation of the
nymph of Odonata, solves the problem
of the venation of insects in general,
and especially for Neuroptcra, Trichop-
tera, Mecoptera (Panorpidae), Lcpi-
doptera and Diptera.
Spuler's method was to strip off the
loose skin of a caterpillar just beginning to pupate, and examine the incipient
venation of the wings of the young
pupa on the living insect. He placed
the living pupa in water and then,
since the process of thickening and
resulting concealment of the veins of
the wing is retarded, the tracheal
branches become slightly enlarged,
filled with air, and thus are more
easily seen. Hence small pupae from
which the larval skin has just been cast, and are transparent, are the fittest
objects for examination.
The primitive and generalizecl con-
dition of the semipupal wing is shown
a
in Spuler's figure of Cerwa vinula
(Fig. 21, to which we have added the
Fig 2
numbering of all Hie veins.
He shows
that the fundamental pupal venation of
Lepidoptera will also apply toorthop-
tera (Blatta), Hemiptera, Trichoptera,
etc. He proves that the cross veins are
of quite secondary and subordinate
importance. The rcsults of Spuler's
investigations, extended through dif-
fercnt groups from Tineina to I<ho-
palocera, and illustrated by many
figures, are both interesting and con-
vincing. The comparison of the
venation of the fore wing of the adult
Fig. 3.
of Gracilwia spingella (Fig. 3 A)
compared with that of its setnippa
(Fig. 3 B), shows that the generalized




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238 PSYCHE. [May 1895.
venation of the latter is similar to that of Micropteryx, veins IVi IV., not
being 'connected by a cross vein with
I11 and its branches, and veins 11 and
å
111, with their branches, being sepa-
rate. The veins and their numbering
Fig 4.
arc indicated by Spuler's figure of
TaZac$oria å´pseudobomhyceZl (Fig. 4)
and one we have drawn of flr.picilus
musteZinus (Fig. 5).
consist of two layers, an outer (IT) and inner (c), which takes a stain and lies
^r ^
fig. 6.
next to the hypodermis (hy). In the
cavity of the vein is the trachea (fr),
which shows more or less -distinctly
the so-called spiral thread ; within the cavity are also Semper's rib (r) and
blood corpuscles (hc), which proves that the blood circnlatcs in the veins of the completely formed wing, though this
does not appiy to all Lepidoptera with
- -
hard mature wings. I have
been able to observe the same
structure in scctions of the
wing of Zygaena.
A cross-section of a vein of
,
the immature pupa of Pieris
brassicae shows that the
large trachea is first formed,
and that it extends along the
track between the protoplas-
mic threads connecting the
two hypodermal layers.
The main tracheae throw
off on both sides a number of
secondary branches showing
at their end a cell with an
Fig. 5.
intracellular tracheal struct-
The structure of a complete vein is
ure ; these accessory tracheae afterwards described by Spuler. In a cross-section branch out. of a Noctuid (Triphaena å´promtb The accessory or cross-traclicae often Fig. 6) the chitinous walls are seen to disappear, though in some moths they



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remain permanently. Fig. 7 ;'T~ rep- venations. There is no sharply defined resents these secondary veins in the
difference between reticulated and a
non-reticulated venation ; no genetic
difference exists between the two
t, kinds of venation, since there occur
true Blattidae with and without a
reticulated venation (Spuler).
It may be remarked that Spnler
agrees with Brauer and Redten-
bacher, as well as Haase, that
Adolph's system of convex and con-
Fig. ,.
cave veins is entirely erroneous.
edge of the fore wing of Lavernu van- We adopt, then. Spuler's system of ella, arising from a main trachea (tr)
venation, and earnestly trust that it
passing
through
vein I, two
of the
twigs ex-
tending- to
the cen-
tre, sliow-
ing that
the latter
has no
homology
with a
vein. Only rarely and in
strongly developed thick
folds are the cross-tracheae
provided with a chi tinons
thickening, ;is for cxam-
pie in Cossus Zigni-perda,
Since front we11 accessory
tracheae the cross-veins in
lepiclopterous wings are de-
veloped, we can recognize
in them the hornolo~ies of
the net-veins in reticulateil
Fig. 8.




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240 ps 2Txf~T. [May 1895.
may be generally accepted, as simple,
intelligible, and applicable to all orders of insects, based as it is on ontogcnetic, as well as anatomical, grounds.
The following system applies to the
Lepidoptera as well as all other orders. Fig. 8 represents the venation of a
Notodontian (Heterocampa oliliqua),
We merely deviate, from motives of
convenience, from Spuler's numeration
of the two anal veins, by numbering
them VI and VII, instead of designat-
ing them by the Greek letters a P.
The following table will show the
numbers and names of the five veins (if
the outspiead portion of the wing and
two (rarely three) of the fan-like or
inner portion. Instead of denoting the
veins by the noun and adjective as,
foi example the median vein, we may
call it in descriptions or diagnoses,
media.
I. Costa.
11. Subcosta (~adius) .
111. Media.
IV. Cubitus (median icin of some
authors).
V. First and (s~~bmedian).
VI. Second anal (internal).
VII. Third anal.
Istferufure relating to the nenation of
the wings of insects.
Yurine, L. Nouvelle m&hode cie clafeser les Hymdnopthres et les Dipleres.
14 PI.
Genfeve, 1807.
- - . Observations sur les ailes des
Hym&nopl&.es. (M6ni. Acad. Turin.,
xxiv, pp. 177-214, 1820.)
Lefebzwe, A. Communication verbale sur
a pterologie dea L6pidoplbres. 3 Pi.
(Ann. Soc. Ent. France, xi, pp. 5-33.
Rev. Zool., v, pp. 52-55. PI. I. 1892).
Burmeistrr, H. Untersuchungen ueber die
Flugeltypen der Coieopteren. (Abh.
naturf. Ges. Halle, ii, pp. 125-I+>. I PI. 1854.)
Ramand, B. E. de. Tablcmi de I'nile
;i,p&ieure des Hyrnhopt&res. I PI.
1839. (Revue Zool., ii, p. 339, 1839.)
Westwood, J. 0. Notes on the wing veins
of insects. (Trans. Ent. Soc. London.
Ser. 2, iv, pp. 60-64.
1857.)
Srhiner, J. R. Ueber das FlUgelgeader
der Dipteren. (Verh. K. K. Zool.-bot.
Ges. Wie!~., 1864, pp. 193-200. I PI.)
Newman, B. Memorandum on the wing-
rays of insects. (Trans. Ent. Soc.
London. Ser. 2, iii,pp. 225-231. 1855.)
Sjeyer, A. Oken's Isis, p. 94, 1839.
-- .
Stettin. Ent, Zeit., 1870, p. 202.
Figs.
Hagen, H. Ucher rationelie Benennung
des Geiiders in den Flugeln der Insekten. Stettin. Ent. Zeit., p. 316. Figs. 1870. ~ o g b , Ofto, Das FlLigelgeZder der Kafcr. Erlangen. 1875.
Muller, Fritz.
Beitrage zur Kenntniss der
Termiten. Jenaisch. Zeitschr. f. Ndturw. 1875-
Sciussure, H. de. Etudes sur l'aile der
Orthoplbres.
(Ann. sc. nat. 6 sir., x.)
Koite, H. y. Das Fltigeigeader der Psociden und seiner systematische Bedeutung.
(Stettin, Ent. Zeit., 1880, pp. 179-186. I PI.)
-- . Beitrag zur systematik der
Lepidoptera. (Berlin. Enl. Zeitschr.
Bd. xxvii, pp. 217-224. 1883.) "
-- . Die zwischenraume zwischen
den Punctstreifen der punktiert gestrei- fen Fliigeldecken der Coleoptera als
rudimentire Rippen auf gefasst.
(Jahresb. Westfsl. Prov. Ver. Wiseen.
u. Kunst. Munster., 1886, p. 57-59.
I PI.)




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May 1895.1 PSYCHE. 241
Brazi-er, F. Ansichten iiber die pilluozo- ischen Insekten nnd deren Deutung.
Annal. K. K. naturh. Hofmus. Wien. i,
pp. 86-126. 2 PI. 1886.
Redfenbacker^ Joseph. Vergleichende Stu- dien ueber das Fitigeige5derder Insecten. (Annalen des K. K. Naturh. Hofmus.
Bd. i, pp. 153-231.) 12 Pis. Wien,
1886. Abstr. by J. H. Comstock in Amer.
Nat., xxi, pp. 932-93!\.. 1887.
Braver, F. 11. Redfentacher, V. Ein Beitrag zur Entwicklung des Flugelgeaders der
Iusekten, Zool. Anz. 1888, pp. 443-447.
Bonsdwf, A. von. Ueber die Ableitung
der Sculpt~~rverh~ltnissc bci den Deck-
fliigeln der Coleopteren. Zool. Anz.
Jahrg., xiii, 1890, pp. 342-346.
Haase, EricIt. Zur entwicklung der Fltigel- rippen der Schmetterlinge. Zool. Anz.,
xi", 1891, pp. 116-117.
S$iil% A, Zur Phylogenie und Ontogenie
des FItigelgeXders der Schmetterlinge.
Zeits. wissens. Zoologie, liii, pp. 597- 646. 2 PI. 1892.
Cornstock, % H. Evolutionand Taxonomy,
etc. Ithaca, N. Y. 1893.
Also the woi-ks of Kirby and Spcnce,
Burmeister, Doubleday, Herrich-Schaeffer, Westwood, lleer, Osten-Sacken, Scudder,
Adolph, Graber, Dyar, Kolbe, Packard, etc. THE GENUS OXYPTILA.
BY NATHAN BANKS, SEA CLIFF, N. Y.
Oxyptila is a genus of Thomisoid
spiders, described by Simon in 1864.
I consider the characters of this genus
in our fauna are that the quadrangle of
the M. E. is higher than broad, that
the P. M. E. are closer to each other
than to the S. E., and that the body
bears clavate hairs. Thus I would not
place in the genus 0. cineyea Em.
(New Eng. Thomisidae), as it bears
no clavate hairs and as it has much the
appearance ofa true Xystictis. In 1877
Thorell described one species of this
genus, 0. conspurcata,^ from Colorado.
In .188o Keyserling described two
species, 0. geo~giana and 0. neva-
densis. In 1882 Kyserling described a
third species, 0. mowoensis from Ft.
Monroe, Va. Dr. Marx in 1890
recorded this species from D. C. In
1892 I recorded both 0. geo~giana and
0. consptt~cata from Ithaca, N. Y. I
have since decided that 0. ~eorgiana
is the same as 0. co~zs$wcata; at
least I can see no other than color
differences between forms which agree
with the descriptions ; and the form
which I recorded from York as
0. geoygiana is not that species, but is new. I have since received two other
new species and obtained 0. monro-
ens-is from Long Island. The six
species may be tabulated as follows :-
I !
No spines above on metatarsus I, quite black species . . floridaria.