Article beginning on page 311.
Psyche 7:311-314, 1894.

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PSYCHE.
THE EPHEMERIDAE AND VENATION NOMENCLATURE. BY VERNON L. KELLOGG, STANFORD UNIVERSITY, CALIF. That consummation devoutly to be
wished, the agreement among cntotnol-
. ogists on a rational nomenclature for
the veins of the wings of insects seems
to be a probability of the near future.
Just at present one of the moot points
is presented by the wings of the Eplie-
meridae. * Redtenbacher in his elab-
orate study of wing-venation chose,
rather unhappily it now seems, the
Ephemerid wing as type of the existing
generalized wing. In this wing, in
addition to the usually distinct and
easily recognized five principal longi-
tudinal veins (exclusive of the anal
veins), viz. : costa (marginal vein) sub- costa, radius, media and cubitus, there
are two other apparently equally iin-
portant and fundamentally distinct
longitudinal veins, one lying between
radius and media, the other between
media and cubitus. These two veins
are called by Redlenbachcr TV and VI,
all the orders of insects.
* Com~to~li,
adopting in the main Redtenbacher's
nomenclature, explains the presence of
these two veins (IV and VI), naming
them pertinently premedia and post-
media, differently from Redtenbacher.
Professor Cornstock pointed out that
the veins in question were wanling in
the wings of paleozoic insects, and
stated his belief that the veins have
been secondarily acquired among the
Ma:-flies by a sttaightening ont of the
zigzag lines between two series of cells. That is, they are essentially venae
sfwiae. f Spuler, studying the on-
togeny or the lepidopterous wing found
no indication of the two veins, and
suggested a system of no~nenclatui e
which omits any reference to veins IV
and VI of Redtenbacher as independent
longitudinal veins. Spulcr's concln-
sions and system are accepted by $ Dr.
111, V, VII. Redtenbacher, believing 1-111, these veins to be incident to the facial t sp~ier, A., ZIW PIiylugeuie d Ontogtniie dee Flligel- insect Wing SOUgl,t to find them in
genders der Sclimetterlinge,
Zettsch. f. wiss. Zoologie, Bd.
LIII, s. ~~~-646, t. XXV, XXVI.
--
* Redtenbacher, Josef., Vergleichende Studien iiber das i. Packard, A. S., On a.Rationa1 Nomenclature of the Flugelgesder der Tnsecten, Ann. d, k. It, naturliistorischen Veins of Insects, especially those of Lepidoptera, Psyche, Hofuiuseams, Wien, Ed. I. s. 153-232, t. IX-XX, 1886. Ma?, 1895.




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312 ps 2TlYE. [December 1895.
Packard.
In his most recent *writing
on the matter, Professor Cornstock
states that in his present opinion it
would lie well not to recognize pre-
media and postmedia as belonging to
the group of principal longitudinal
veins.
As a note, perhaps of interest, I offer' a biief account of certain observations
recently made on Ephemcrid wings.
A comparison of the figures of Red-
tenbaeher and Comstocl< show that
although both refer to a premedial and
a postmedial vein they differ in their
interpretations of what constitutes these veins and their respective branches (see figures I and z, after Redtenbacher and
Fig. T
Coinstock, respectively ). What Pro-
fessor Coinstock defines to be vein IV
and its branches includes part of Red-
tenha&erYs rein 111 and all of his veins IV, V, VI, and their branches. Vein
V of Cums~ock is Redtenliacher's vein
VII, and Coinstock's veins VI and VII
are included by Recltenbacher among
the anal veins. + Scudder's description
* Comstodt, J. H., The Venation of the Wings of Insects, pp. 75-91,inThe Elements of Insect Anatomy,by Cornstock and Kellogg, 1895,
tScudder, S. H., The Pretertiary Insects of North America, pp. 5-6, 18p
of the venation of the Ephemeridae
agrees with Comstock's in that both
make radius (scapuhiris of Scuclder.
after Heer) a simple (unbranched)
vein, but Scndder does not recognize
any independent longitudinal veins
Fig. 2
between radius and media or between
media md cubilus. Comstock's veins
IV and V are Scudcler's externo-median,
and his veins VI and VII are Scudder's
interno-median. That is, Scudder
makes premedia. a branch of media, and
postn~cdia a branch of cubitus.
None of these interpretations of the
Ephemerid venation seems to me the
correct one ! The presumptuousness of
this statement should be less offensive
when we recall the fact that no two of
the three already offered interpretations agree. A characteristic of radius
noticeable in generalized wings and
strenuously preserved in the specialized wings, is its branched condition. Just
as sub-costa is characterized by its uni- formly unbranched condition (exccpt-
ing in the wings of a few very
generalized insects, as the Blattidae) , and media is characterized by its
tendency to lose its basal half, so radius and cubitus are characterized by the
persistence of their branches. Radius
in its mode of branching also shows




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December ,895 1 1's ITHi??. 313
a rccogiiiz;ible consistency. Broadly
stated, tlic manner of the branching is
this: the stem forks rather near its base, the upper branch, which either does
not fork again (in more speciiilizecl
wings) or gives off a few branches (a
more gcneriilized condition) appearing
to be move ilirectly a continuation of
the basit1 trunk thsin the lower ILn'i'mcl~, which usually displays a "branching
away" cliamcter, and which is almost
always repeatedly forked and briinclicd. This repentedly lorking lower branch
is (lie radial sector of authors. In the more genen-ilizecl venation the sector
l~r~iticlies from' the radial stem 11c;ir its base, and is many-forked. The modi-
ticittions which 111e sector and its
branches exhibit, due to 111e specializing tendency ofthe wing toward narrowness
with iiccoiiipanyit~g coalescence and
disappearance of vein branches are the
reduction in the nunilxr (coalescence)
of the branches and the movement of
the point of origin of the sector I'artlier and farther away from the base of thc
wing.
Now although the lCpliemeric1 wings
are in point of specialization in advance (shown by the reduction of thc liincl
wing, and the specialization of the
thorax) of the genenil rank of the
f:imily among insects (piired genital
openings, etc.), the wings have by no
means reached that degree of special-
ization where radius has become an
iinbranched vein. In fact, radius in
the Epliemerid wing is, to my mind, in
very generalized condition. The many-
branched radial sector departs from the
stem very near its base. so near indeed, that by a slight modification it has
become apparently entirely distinct from raclius, and. in some Mayflies even
apparently joined at its base with media. Such an apparent or even real clissocia- tion of a branch from its original stem
and re-association with another vein is
lot ;in iiiicomtnon plienoinetion in the nodification of venation ; note among
the Lepidoptera tile association of the
branches of media, after the base of
media has disappeared. with mdins and
e~il)ittis.
Fig. 3,
No is this condition of radi~is and
its sector unique with tlie May-flies.
Among Neiiropteroid insects in general
tlie sector usually arises near the base of the radial stem (Odonata, Sialiclae
ct al.) , and nut unfrequently is appar- ently dissociated from the radial stem,
and re-associated with media, as in
certain Odonata and Perlidae. and,
;~niong unrelated forms, in Enii~ia,
Fulyora et at. In some cases the base
of media is intimately united (coalesces) with the base of radius, as in Neinura
(see figtire 3). In fact the crowding
together of the vein stems at the base of the wing brings about much distortion
and motlification of tliese one-time
mutually independent and co-important
trunlts.




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I would also designate the two veins
either as original principal vcins (Red- called by Redmhacher VI (postmedia
tenbaclier) or as venae spziriae (Com-
of Comstock) and VII, simply as vein
stock).
VII. Cubitus as well as radius is More convincing than the argument characterized by the persistence of its from analogy for the correctness of this branches. As defined by Redtenbacher
cubitus in the Ephemerid wing has lost
all of its branches. This is extreme
specialization. In my opinion cubitus
in the Ephemerid wing forks at the very
base producing the effect of two inde-
.. ..: ,
IX
pendent longitudinal veins. The same Fir. 4 effect is sllo~n in Neln~ra (see figure ~lltcrl,ret~lt~or~ is o~ls ed fact t~lat 3, especially hind wing) and is inter- ^ ^^ trll& (original ^ of tile preted hy * Rcdtenbacher exactly as 1
of tile ,.aclial be dis-
would interpret the similar condition in tinctly traced, at least in observed
the Ephemeridae. Nemura, indeed, in
instances, as a primary branch of the
the condition of both radius find cubitus radial tracheal trunk, altliough the
is very like the Epllelnerid wing and it chitillous envelop of the sector2s t,.aclleal is interesting to note Redtenbacher's
branch, which gives the vein its visi-
interpretation (fia 3) of the venation of bility to the naked eye, is not present at this wing.
No longer constrained by
tlie base of the sector.
In studying the
the rigid limits which the application
vcllalio,l of certaill Blattid willgs I
of the theory of origin:~l convex and
foull~l t,,at Llle kving could be so
concave vcins entailed we can now
mounted that the trachea! trunks (or,
hardly justify tlie acceptance of two
more accurately, probably, the *å´ Rip-
such variant interpretations of two such penstriinge,'' relicts of the original
essentially similar wings.
tracheal trunks and identical in course
In my view, tlierefore, the veins of
with them), the foundations of the
the Ephemerid wing sho~ild lie liomol-
present visible veins, could frequently
ogized as indicated in f figure 4.
The
be seen and traced.
At the base of any
interpretation does away with any
wing the thick chitin envelop of a vein
recogllitio~l of veins IV bemedia) and
is often
althoug~l t,le t,.aclleal
VI (~osl~c~i~) as independent veins,
trunlc ,lcrsists,
In a nloullted wing of
-
- ~ Hexagenia sp. I have plainly ob-
* I recognize, of course, that Redlenbciclier's interprets- tiom are influenced always by the convex and concave vein served the branching trachea of the
theory.
sector arising from the radial trunk at
t Evidently, if this interpretation is correct and veins IV and VI are not elsewhere found as original principal veins, *Korscheit, E., and Heider, K.
Lehrbuch der verglei-
the veins should be numbered as fo1lov.s: I = costa, I1 = chenden Entwickiuiiesgescl~ichte der wirbdloscn Thiers, p, subcosta. I11 =radius, IV = media, V = cubitus. 864, 1890.



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December i8qs.J PLs' TCHt??. 315
an appreciable distance from the base
of radius. This wing also shows the
unity at the base of the wing of the two main branches of cubitus.
In the examination of a considerable
number of wings from various orders of
insects I have noted that anal veins
arise from certain tracheal trunks or
often apparently from one main trunk
which divides at the base of the wing
into several or many branches or rays ;
and that this main anal trunk is distinct from the main trunk or trunks which
run into the pre-anal arca of the wing,
and which are the fo~indations of the
principal longitudinal veins of the pre- anal arca. In the wing of Hexagenia
the veins corresponding to those veins
marked " anal " in figure 4 are supplied with tracheae from one main trunk,
the anal area trunk, while the first vein in front of these anal veins (called by
me part of cubitus) does not receive a
branch from this main anal trunk.
ON THE NEST AND PARASITES OF PROSOPIS VARZPRONS CRESSON.
BY A. DAVIDSON. M. D., LOS ANGELES, CAL. This bee and Ceratina dwpla are the
most common ones that tunnel in the
shoots of the elder and mustard in this
locality. The cells are built in stems
which the parent herself has hollowed
out, and measure on an average 4 lines
long, by 2 or 3 wide. They are lined
wit11 a thin, transparent layer of silk
which is spun by the parent bee, and
are filled three-fourths full with the
light yellow semi-fluid bee-food. The
egg is laid on top of this mass, and the cell is closed by the same silken tissue which, in its turn, forms the base of the succeeding cell. The last of the series, when finished, is further protected by a layer of pith of variable depth. The
cells are probably normally built con-
tig~~ously, but a few sometimes have
prtitions of pith, which may be the
work of more than onc bee.
Two broods at least are produced
annually.
By splitting' the twigs con-
taining the cells I was enabled to watch the larvae pass through their various
stages, and in a series gathered on
May j at Manzana, in the Antelope
Valley, all of which were apparently
newly constructed, the food was con-
sumed in 8 clays after the hatching of
the egg. The larvae at this stage are
quite active, and in their restlessness a few of them burst through the lining of
the cell on the exposed side and made
their escape. During this period of
activity they void a small quantity of
excrement which, being limited to one
end of the cell, simulates an artificial partition between the cells.
In from four to six days after the
food was consumed they passed into
the pupa stage, and on June 9th with
two exceptions all had taken their
fliglit, t.he time occupied in passing




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