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FOR THE -BEOFLE
FOR EDVCATION
FOR SCIENCE
LIBRARY
OF
THE AMERICAN MUSEUM
OF
NATURAL HISTORY
Sound at
A.M Nh |
190986
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—
— a
Journal of
Entomology and Zoology
VOLUME XV, 1923
PUBLISHED QUARTERLY BY
POMONA COLLEGE DEPARTMENT of ZOOLOGY
CLAREMONT, CALIFORNIA, U. S. A.
45-401 47 |» ~ Pye
CONTENTS OF WOE aa
Volume XV, Number 1
Merrill, Ida; Schoonover, E.
A Model of the Nasal Chambers
of a White Mouse at Birth, 1.
Chamberlin, R. V.
North American
Mimetus, 3.
Hilton, W. A.
The Neryous System and Sense
Organs, XII, 11.
Species. of
Volume XV, Number 2
Hilton, W. A.
Nervous System and Sense Or-
fangs; KITT. 17.
Cole, F. R.
Notes on the Early Stages of
the Syrphid Genus Microdin
(Diptera), 19.
Cole, F. R.
Notes on California Bombyliidae
with Descriptions of New
Species, 21.
Marimon, Sarah
‘Notes on the Color Changes of
Frogs, 27.
Volume XV, Number 3
Meadows, Donald C,
Notes on the Lepidoptera of
Southern California No. 1. 33.
Dodds, Clifford T.
A List of Coleoptera Collected
on the Beach During the
Summer of 1921 at Laguna
Beach, 35.
Campbell, Arthur S.
Some Common Chinese Mol-
lusca, 37.
Hilton, W. A.
The Nervous System and Sense
Organs, XIV, 43.
Volume XV, Number 4
Dodds, Clifford T.
A New Salt Marsh Mealy Bug,
57%.
Campbell, Roy E.
Notes on the Life History of
Dinaparte Wrightii Horn, 61.
Hilton, W. A.
Nervous System and Sense Or-
gans XALV Cont.,” 67.
INDEX TO VOL. XV
Bombyliidae, 21
Campbell, A. S., 37
Campbell, R. E., 61
Chamberlin, R. V., 3
Coles kt Rs, 19; 21
Coleoptera, 37, 61
Color Change, 27
Dinaparte, 61
Diptera, 19, 21
Dodds, C. T., 35, 57
Frogs, 27
Eiton We =Aw ls 175 43.) 67
Lepidoptera, 33
Marimon, S., 27
Meadows, D., 33
Mealy Bug, 57
Merrill. Tent
Microdon, 19
Mimetus, 3
Model, 1
Mollusca, 37
Mouse, 1
Nasal Chamber, 1
Nervous System, 11, 17, 43, 67
Schoonover, E, 1
Sense Organs, 11, 17, 43, 67
Spiders, 3
Syrphid, 19
A Model of the Nasal Chamber of a
White Mouse at Birth
Ida Merrill and Eugenia Schoonover
Two models were made by the blotting paper method. In one
of these the outer portion of the epithelium was taken as the outer
limit and the lining of the cavity as the other. The other model
was built from the plates which were cut from the interior of the
Upper figure model of the cavity, from outside.
Lower figure model of epithelium from outside. Both figures X40.
2 Journal of Entomology and Zoology
model. The larger model gives a picture of the mucous membrane
and the nasal chambers, the smaller shows the shape of the nasal
chamber. The knob on the inner side of the larger model is Jacob-
son’s organ. The drawings are by Elizabeth Keyes.
(Contribution from Zoological Laboratory of Pomona College.)
Upper figure section of the model of epithelium.
Lower figure model of epithelium from the outside. Jacobson’s organ, the
little elevation in the lower center. X40.
eee
The North American Species of Mimetus
By Ralph V. Chamberlin
In his “Araneides of the United States,’ Hentz describes three
species under the genus Mimetus; namely, interfector, tuberosus
and syllepsicus. Of these three syllepsicus has not since been defi-
nitely identified, while twberosus is generally regarded as a synonym
of interfector, a disposition with which no fault can be found. In
‘1882 Emerton described a male from Connecticut under the name
M. epeiroides; but the practice in recent years has been to refer all
individuals of the genus found in the United States to one species,
interfector, and, accordingly, in current catalogues epeiroides has
been placed in synonymy with that species.
However, a careful study of ample material of Mimetus from
various parts of the country reveals that there are at least five
clearly distinct species that have been confused under the name
interfector. One species occurs on the Pacific Coast apparently
from Washington to southern California and eastward to Texas.
Two species occur in the Northeastern States, the commoner of
these ranging southward as far as northern Georgia. The other
two species are common in the Southern States; and one of them is
found as far northward as Long Island, N. Y. It seems reasonably
certain that it was one of these two southern forms that was de-
scribed by Hentz as interfector, the particular one being fixed, it is
believed, by the figure of the palpus as indicated later in the notes
on the species. Tuberosus is left as a synonym of interfector; but
syllepsicus cannot be placed at present and is apparently different
from any of the five species here listed.
The males of these five species are easily recognized by the
characters presented in the palpus, the armature of the ectal mar-
gin of the cymbium providing a convenient index. Another readily
observed character of diagnostic importance occurs in the ter-
minal portion of the bulb which in the retracted organ lies adjacent
to the base of the embolus and presents typically two flat or lamellar
lobes projecting proximad. In one species (notius) one of these
lamellar lobes is aborted and in another (puritanus) the second lobe
is itself partly divided or bilobed.
The four species of which females are known may be sepa-
rated in that sex by the characters of the epigynum, which is in the
form of a strongly chitinized, transversely furrowed, caudally pro-
jecting lobe. At the caudal end of the epigynum, or near it on
its dorsal side, is an opening or pit and cephalad or proximad of
this on the dorsal side is a separately chitinized median longitudinal
piece or strip. The position and form of this epigynal opening or
pit, the size and position of caudal end of the median dorsal strip,
4
Journal of Entomology and Zoology
and the form and position of the spermathecae as revealed in
cleared specimens furnish good diagnostic characters.
Adults of the species here listed may be identified by means of
the following keys in conjunction with the accompanying figures of
palpi and epigyna.
a.
b.
a.
a.
/
Key to Males
Ectal margin of cymbium of palpus with no chitinous, spiniform
process proximad of the curved or bent apical one.
Ectal border of cymbium with an elevated and sharply limited
lobe at caudal end of scabrous portion of margin, the surface
of the lobe covered with minute prickles; bulb with two apical
lamellar lobes Cie Syn rs ead ees M. interfector Hentz
b.’ Scabrous portion of ectal margin of cymbium not ending
caudally in any such sharply defined lobe; apical portion of
bulb bearing only one developed lamellar lobe, the ectal one
being aborted and at most represented by a minute tooth
(Pe A ss Ce. nee ee ae ate M. notius sp. nov.
Ectal margin of cymbium with one or two chitinous processes
or spines proximad of the apical one.
b. With only one spine on margin of cymbium proximad of the
apical one, this toward the base; border scabrous from
apical to basal spine (Fig. 3)..... M. epeiroides Emerton
~
b.’ With two spines on ectal border of cymbium proximad of
the apical one of which the more distal one is sometimes
weak; margin scabrous only from apical spine to the more
distal marginal one.
c. Proximal marginal spine contiguous, or nearly so, with
basal lobe or auricle of cymbium; apical portion of bulb
with neither lamellar lobe at all subdivided or present-
ing processes (Big. 2)... oe M. hesperus sp. nov.
Proximal marginal spine well removed from basal lobe
of cymbium; apical portion of bulb with the larger,
more mesal, lamellar lobe partly subdivided, being
extended at its mesodistal corner (Fig. 1)..........
PP ONT oR seg Py M. puritanus sp. nov.
~
Key to Females.
The opening or pit located at extreme caudal end of epigynum
and visible in ventral view, the end in this view appearing
notched at the middle; median dorsal strip extending nearly to
caudal end of epigynum...... Sta h.o) Yorke M. puritanus sp. nov.
The pit is on the dorsal surface just proximad of caudal end of
Pomona College, Claremont, California 5
epigynum and thus not visible from below, the end not appear-
ing notched at middle; dorsal strip ending considerably proxi-
mad of end of epigynum.
b. Opening with no tooth or process from each lateral margin,
not thus partially subdivided; spermathecae essentially
longitudinal ;dorsal strip broader (Fie. 10) ......6. 66...
Re eee erg tet eee nee ANNs ets er M. notius sp. nov.
b.’ Opening partly divided into a distal and proximal portion
by lateral processes; caudal portion of spermathecae bent
at right angles, a distinct enlarged anterior and porterior
portion being connected by a narrower isthmus; dorsal
. strip narrower.
c. Lateral margins of epigynum not indented; isthmus of
of spermathecae narrower, curved, concave ectally.
A Nee Uns ened 1 Oe Ea te nee Tei M. interfector Hentz
c.’ Lateral margins of epigynum indented near level of
caudal ends of spermathecae; isthmus of spermathecae
thick, straight (Figs 7 and 8)....M. hesperus sp. nov.
Mimetus hesperus sp. nov.
In the male of this species the ectal margin of the cymbium of
the palpus bears two conspicuous black spines proximad of the
“apical curved one as in puritanus; but in the present species the
more proximal of these spines is in the re-entrant angle above
basal lobe or auricle, whereas it is distinctly distad of this position
in puritanus. A readily noted difference in the bulb is that the
larger lobe at apex of bulb is entire in hesperus, with no separate
process from inner distal corner as in the eastern form; and be-
tween this lobe and the conductor there are two folds of conical out-
line not present in the latter species (Cf. fig 2). The female
differs conspicuously in not having the epigynal opening ter-
minal and thus producing a median notch when viewed from below.
The epigynum in its structure most resembles that of interfector,
but differs in outline and in the form of the spermathecae (Cf.
figs. 7 and 8).
Type Locality —California: Claremont. Type, a male, M. C. Z.
No. 530.
Other Localities—California: Stanford; Washington: Camp
«matilla; Utah; Texas: San Antonio, Austin.
Mimetus puritanus sp. nov.
Mimetus interfector Emerton (nec. Hentz), Trans. Conn. Acad.
Sen, 1882; 6, p. 16, ple 35, fie 3:
—_
6 Journal of Entomology and Zoology
Mimetus interfector Keyserling (in part, including those fig-
ured), Spinnen Amerikas, Theridiidae 2, 1886, p. 6, pl, 11, fig. 137.
This species is in the female sex at once distinguishable from
all the others in having the epigynal pit at the caudal apex and
visible as a median notch from below (Fig. 6). The male may be
separated from the other species occurring in the eastern and
southern States by the presence of two subapical spines on the ectal
margin of the cymbium; and from the western hesperus, as indi-
cated above, by the position of the more proximal of these spines
and the form of the larger lamellar lobe of the bulb, which is unlike
that of any other species (Fig. 1).
Type Locality—New York: Ithaca. Type, M.C. Z. No. 535,
a male.
Other Localities—New York: Long Island, Sea Cliff; Maine:
Ogunquit; Mass.: Ipswich, Plymouth; Conn.: New Haven; Vir-
ginia: Great Falls, Falls Church; Georgia: Thompson’s Mills.
Mimetus epeiroides Emerton
Trans. Conn. Acad. Sci., 1882, 6, p. 17, pl. 3, fig. 4.
Known only from the male which is clearly distinct from the
other species in characters of the palpus. In this the ectal margin
of the cymbium possesses a single spine toward the basal lobe, in
distinction from the two preceding species in which there are two
spines on the margin, and from the two following ones in which
there is no marginal spine proximad of the distal one. The ectal
border is scabrous over its entire length from apex to basal spine.
The terminal portion of bulb bears two lamellar lobes, both of which
are simple.
Type Locality.—Mass.: Essex.
Immature specimens referred to this species have also been
taken by Mr. Emerton at other places in eastern Massachusetts and
at Providence, Rhode Island.
Mimetus interfector Hentz
Journ. Boston Soc. Nat. Hist., 1850, y, p. 3, pl. 4, fig. 12, 138.
Mimetus tuberosus Hentz, ibid., p. 3, pl. 4, fig. 14.
Of each of the two species of Mimetus occurring commonly in
the southern States, individuals may be found which match the fig-
ures of interfector given by Hentz reasonably well. I believe the
species to be fixed, however, by the figure of the palpus of the male
which, in spite of its general inadequacy, shows two prominent
lobes projecting proximad from the bulb that are apparently the
two lamellar lobes present in the one species, whereas in the other
species, listed below as M. notius, sp. nov., there is but a single
Pomona College, Claremont, California 7
lamellar lobe. In the species thus considered to be fixed as the
true interfector of Hentz the ectal margin of the cymbium lacks
spines; the scabrous border ends proximally abruptly in a lobe
elevated above the general surface and on which the area of
prickles is broader, a very characteristic feature enabling one to
detect the species at a glance (Fig. 5). The form of the opening
of the epigynal pit is similar to that of hesperus, being partly sub-
divided by a projection from each lateral margin and thus differing
from that of notius. The spermathecae also present a caudal and
an anterior larger lobe connected by a narrower, weakly curved,
isthmus.
Type Locality —Alabama.
Other Localities —Alabama: Morgan, Birmingham; Georgia:
Atlanta; Louisiana: Shreveport, Covington, Shrewsbury; North
Carolina; New York: Sea Cliff.
Mimetus notius, sp. nov.
In this species the opening of the epigynum lacks projections
from its lateral margins, and the median dorsal strip is broader and
more conspicuous than, e. g., in M. interfector or M. hesperus; the
spermathecae are essentially longitudinal as shown in fig. 10. The
male differs from all the others here considered in having on the
distal portion of bulb of palpus only a single lamellar lobe, the ectal
one being absent or represented only by a slight tooth at base of the
developed lobe. The ectal margin of the cymbium lacks spines
proximad of the apex and its scabrous border runs out gradually,
not ending in any such abruptly elevated lobe as occurs in inter-
fector.
Type Locality—Runnymede. Type, a male, M.C. Z. No. 551.
Other Localities—Florida: Altoona, Daytona; Louisiana:
Shreveport, Mansura; North Carolina: Raleigh.
Fig. 1. Mimetus puritanus sp. nov. Right palpus of male,
subectal view. 2. M. hesperus sp. nov. Right palpus of male,
similar view. 3. M. epeiroides Emerton. Right palpus of male
(type) from a more dorsal aspect, the hematodocha distended. 4.
M. notius sp. nov. Right palpus of male, subectal aspect. 5.
M. interfector Hentz. Right palpus of male, subectal aspect.
Fig. 6. Mimetus puritanus sp. nov. Epigynum, ventral
view. 7. M. hesperus sp. nov. Epigynum, ventral view. 8. M.
hesperus sp. nov. Epigynum viewed from above by transmitted
light, showing opening, dorsal strip, and the spermathecae in sil-
houette. 9. M. interfector Hentz. Epigynum in ventral view.
10. M. notius sp. nov. Epigynum viewed from above by trans-
mitted light to show form of opening and of dorsal strip and the
spermathecae in silhouette.
XII. Enteropneusta
For our general knowledge of the central nervous system of
this group we have the papers of Spengel, 1884-1894, Bateson, 1886.
Of the development of the nervous system and the larvae the
work began in 1870 with the study of the so-called Tornaria larvae.
Bateson, 1884-5, worked out the life history of a Balanoglossus
form and later Spengel, 794 and Morgan, 791 and ’94 gave an ex-
tended account of the Tornarian forms, including a good account of
the nervous system. Ritter, 94 and Davis, ’08, gave stages in the
development of Tornaria and Dolichoglossus, and Herder, 1909, also
gave an account of development in which the nervous system was
included.
In various accounts of the position and structure of the nervous
system especially as summarized in text books and other places,
there seems at times to be some difference in the descriptions but
I think for the most part the differences are in the way of express-
ing much the same idea so that no real difference is introduced.
In all cases the nervous system is as a whole epidermal much
as in Phoronis and in starfish. The epithelium everywhere is more
or less made up of columnar cells at the surface with a deeper
nervous layer of fibers, in part branches from the surface cells, and
a few deeper cells. In places the epidermic nervous system is
more marked. The whole body then might be described as covered
with a plexus of nerve cells and fibers; the thicker parts of the
plexus in places form the so-called nerves. The chief nerves of this
sort are a dorsal and ventral tract in the body region below the
collar with a circular band connecting these at the lower edge of
the collar, and a concentration of fibres about the base of the pro-
boscis, but the greater concentrations are in the collar itself. In
the dorsal and ventral surfaces of the collar just under the epi-
dermis is a concentration of nerve cells and fibers but the chief and
central concentration of nervous tissues is in the form of a thicker
cord running through the cavity of the collar on the dorsal side,
although connected with the epidermis at each end. This central
nervous system is continuous with the proboscis thickening in front
and as just described, with the dorsal and circular nerve tracts
behind.
To summarize, the nervous system may be described as fol-
lows: |
~ 1. General epidermal plexus continuous with other parts.
2. Basal proboscis ring continued into the proboscis by a
more diffuse band.
3. Ventral body nerve continued into ventral collar as a thin
layer.
pire A EATICLC Et De
Fig. 25. Diagrams to show the position of the nervous system in Dolicho-
( Nj
glossus. Nervous system shown by heavy lines below the surface.
1. Longitudinal section. 2. Cross section through the proboscis.
3. Central portion much enlarged. 4. Another part of the surface.
5. Neural epithelium much enlarged.
Pomona College, Claremont, California 13
4. Dorsal collar nerve somewhat cut off from the two follow-
ing.
5. Dorsal proboscis nerve continued above.
6. Central nervous system running through the central region
of the collar on the dorsal side and continuous above with the pro-
boscis nerves and below the collar with the dorsal body nerve.
The dorsal nerve of the collar and the thick central nervous
system of the collar are more or less joined by the strands and
they together make a sort of nerve tube thin on the dorsal side but
thick below.
The histological structure of the nervous system reveals be-
sides the usual epithelial cells of the surface, bipolar supportive
Fig. 24. Nervous System and Sense Organs of Enteropneusta. A. Diagram
of Balanoglossus showing position of the nervous system. B. Anoth-
er diagram of Balanoglossus in sectional view. Spengel. C, D.
Sections of developing nervous system. Morgan. E. Larva after
Herder. F. Apical eye of tornaria larva. Spengel. G. Eyes of
Tornaria after Morgan. H. Eye of Tornaria after Spengel.
14 Journal of Entomology and Zoology
cells reaching from the surface to the depths of the nervous system
and also probably bipolar sense cells as well as more or less deeply
placed multipolar nerve cells giving off fibers to the nerve areas.
The epidermis is a general organ of sense, the exact nature of
which has not been very clearly determined. Spengel considers
that about the proboscis in its ventral face there are points espe-
cially sensitive. In fact at this place he describes a deep depression
which he regarded as a special sense organ.
In the larval stage the first suggestion of a nervous system we
find in the development of the apical plate which in later stages
develops eye spots as simple caps of ectodermal cells surrounded by
pigment. The eye spots become anterior in position with a pocket
of the clear cells each ending in a point. Between the two eye-
cups a mass of pigment develops. At the base of the apical plate
nerve fibers begin to be seen.
At metamorphosis in a region where the collar will develop a
transverse groove forms near the mid-dorsal line. In the mid-
dorsal region a strip of ectoderm not crossed by grooves makes the
beginning of the neural plate. It sinks beneath the surface and
folds of the adjacent ectoderm or neural folds meet over it, and in
this way the neural tube is formed.
BIBLIOGRAPHY
Bateson, ‘W.
18294. The Early Stages of the Development of Balanoglossus. Q. Jour.
mic. se., vol. 24, pp. 207-235, pl. 18-21.
1885. Later Stages in the Development of Balanoglossus kovalevskyi.
With a suggestion of the Affinities of Enteropneusta. Q. Jour.
mic. sc. vol. 25, supp. pp. 81-122, pl. 4-9.
1886. Continued account of the Later Stages in the Development of
Balanoglossus kovalevskyi, and the Morphology of the Enterop-
neusta. Q. Jour. mic. sc. vol. 26, pp. 512-533, pl. 28-33.
1886. On the Morphology of the Enteropneusta. Stud. M. Z. vol. 3, pp.
37-65, pl. 7-12.
Davis, B. M.
1908. The early Life History of Dolichoglossus pusillus. Univ. Calif.
Pub. zool. vol. 4, no. 3.
Dawydoff, C.
1909. Beobachtungen uber den Regenerationsprozess bei den Enterop-
neusten. Zeit. f. wiss. zool. Bd. 93, pp. 287-305, pl. 13-16, 23 text
fig. P. 293 N. Syst.
Haldeman, G. B.
1886. Notes on Tornaria and Balanoglossus. J. Hopkins Univ. Cire. vol.
6, pp. 44-5.
Herder, K.
1909. Zur Entwickelung von Balanoglossus clavigerus. Zool. anz. vol.
34,
Pomona College, Claremont, California 5
Hilton, W. A.
1919. The Central Nervous System of Dolichoglossus. Jour. Ent. Zool.
vol. 9, pp. 59-61, figs. 1-5.
Kovalevsky, A.
1866. Anatomie des Balanoglossus (Delle Chiaje). Mem. Acad. inp. se. °
St. Petersburg. 7e ser. vol. 10, no. 3, 3 pl.
Koehler, R.
1886. Contribution a l’etude des Enteropneustes. Recherch anat. sur le
Balanoglossus sarniensis nov. sp. Internat. Monats. Anat. Hist.
vol. 3, pp. 139-190, pl. 4-6.
Metchnikov, E.
1870. Untersuchungen ueber die Metamorphose einiger Seenthiere. 1.
Ueber Tornaria. Zeit. f. wiss. zool. vol. 20, pp. 131-144, pl. 13.
Morgan, T. H.
1891. The Growth and Development of Tornaria. Jour. Morph. vol. 5
pp. 407-458, pl. 24-28.
1894. The Development of Balanoglossus. Jour. Morph. vol. 9, pp. 1-86,
pl. 1-6.
Ritter, W. E.
1894. On a New Balanoglossus and its Possession of an Endostyle.
Zool. anz. vol. 17, pp. 24-380.
1900. The Movements of Enteropneusta and the mechanism by which
these are accomplished. Biol. Lect. Woods Hole. vol. 3.
Spengel, J. W.
1884. Zur Anatomie des Balanoglossus. Mitt. zool. St. Neap. vol. 5, pp.
494-508, pl. 30.
1893. Die Enteropneusten des Golfes von Neapel und der angrenzenden
Meeres-Abschnitte. Fauna Flora Golfes Neap. 18 Monog. 758 pp.
37 pl.
Spengel, J. W.
1877. Ueber den Bau und die Entwickelung des Balanoglossus. Amtl.
Ber. 50. vers. d. Naturf. u. artze. Munchen.
1903. Neue Beitrage zur Kenntniss der Enteropneusten. 1. Ptychodera
fluva Esch. zool. Jahrb. Bd. 18, pp. 271-326, pl. 24-29, 5 text figs.
1904. 3. Eine neue Enteropneusten art aus den Golf. von Neapel. Zool.
Jahrb. vol. 21, pp. 15-362, pl. 20-22, 10 text figs.
Stiasny, G.
1914. Studien ueber die Entwickelung des Balanoglossus clavigerus
Delle Cheaje. II Darstellung der weiteren Entwickelung bis zur
metamorphose. Mitt. zool. St. Neap. vol. 22, pp. 255-290, pl. 59.
13 text figs.
1914. Studien ueber die Entwickelung des Balanoglossus clavigerus Delle
Cheaie. I Die Eentwickelung der Tornaria. Zeit. f. wiss. zool.
vol. 110, pp. 36-75, pl. 4-6, 24 text figs.
XIII. Cephalodiscus and Rhabdopleura
CEPHALODISCUS. The first report on this animal including a
sketch of its anatomy was by McIntosh, 1887, and later by Lang,
1890, and Harmer. Delage and Herouard, 1897, summarize the
knowledge of the nervous system about as follows:
The nervous system is a thickening of the epidermis on the
dorsal surfaces of the tentacles. The histological nature of the
nervous system was a little studied, but cells and fibers under the
epithelium as in echinoderms were described.
Mastermann, 1903, describes the central ganglion over the sub-
neural blood sinus; its position is as in Actinotrocha. This gives
off below a pair of large nerves each of which divides into six
branches for the six pairs of tentacle arms. Above it is prolonged
Fig. 26. A. Cephalodiscus showing location of the nervous system after
Hammer. B. Cephalodiscus after Masterman. C. Rhapdopleura
showing position of nervous system after Delage and Herouard.
18 Journal of Entomology and Zoology
into two large branches which follow the dorsal line of the epistome.
Laterally from the ganglion two other nerve branches go to the
epistomal disc. On the ventral surface of the trunk is a medial
longitudinal band which is continued into the peduncle. According
to Mastermann the mid-dorsal and two lateral epistomal branches
have homlogues in Balanglossus and Actinotrocha.
RHABDOPLEURA. The account of the structure of Rhabdopleura
which is usually given is that of Fowler, 1892. Other accounts
which however give little of the nervous system are those of Allman,
1869, and Lankester, 1874.
The central nervous system is represented by a thickening of
the ectoderm in the median region of the neck below the nucal pore
between the branches of the tentacles. There is a differentiation
of nervous tissue much as in Rhabdopleura or Balanoglossus. A
black pigment spot is located at the tip of the preoral lobe and may
be an eye-spot.
~
Fowler, G. H.
1892. The Morphology of Rhabdopleura normani Allm. Festachrift.
zur 70 Geburtst. R. Leuckart’s Lepzig. pp. 293-297. pl. 30.
Harmer, S. F.
1887. Cephalodiscus. Zool. of H. M. S. Challenger, vo. 22, part 63, pp.
39-47, 2 figs, 7 pl.
BIBLIOGRAPHY
Lang, A.
1890. Zum Verstandniss der Organisation von Cephalodiscus dodecalo-
phus McIntosh. Jen. Zeit. f. Nat. vol. 25, pp. 1-12.
Lankester, E. R.
1874. Remarks on the Affinities of Rhabdopleura. Q. Jour. mic. se. vol.
14, pp. 77-81.
184. Contribution to the Knowledge of Rhabdopleura. Q. Jour. mic. se.
vol. 24, pp. 622-647, 5 pl.
Mastermann, A. T.
1897. On the Diplochorda. 1. Actinotrocha. 2. Cephalodiscus. Q. Jour.
mic. sc. vol. 40, pp. 281-866, pl. 18-26.
1903. On the Diplochorda. 4. On the central complex of Cephalodiscus.
Med. Q. Jour. mic. se. Ns. vol. 46, pp. 32-33.
Schepotief, A.
1908. Cephalodiscus dodecaloptuis. Zool. Jahrb. anat. vol. 25, pp. 405-
494.
Notes on the Early Stages of the Syrphid
Genus Microdon (Diptera)
By Frank R. Cole, Stanford University
The peculiar larvae of the Syrphid flies of the genus Microdon
have been described by several entomologists, but they are known
in only a few species. Enthusiasts in past years placed these
bizarre forms among the molluscs in two or three instances, and
one entomologist stated that they were the early stages of a Coccid
on oaks. Wheeler has given a very interesting account of some
of these early stages and the habits of the flies. In America the
larvae are recorded only from ants nests, but Wasmann states that
they may be found in the nests of certain wasps and termites.
They live in nests in the soil, under rocks or under the bark of
old logs.
The larvae creep very slowly, with a wave-like motion of the
flat ventral sole, which is fringed and applied closely to the sur-
face over which they are travelling. Their food is probably, as
Wheeler believes, the minute pellets of food ejected from the
hypopharyngeal pockets of worker ants after the moisture has been
extracted. There is evidently one bua in a year, the flies emerg-
ing in May and June.
In May, 1917, the writer found a number of pupae of Micro-
don cothurnatus Bigot, while collecting in the Hood River Valley
of Oregon. The type of this species came from “Mt. Hood,” prob-
ably somewhere in the valley north of the mountain. While pull-
ing off the bark from an old pine log an ant’s nest was uncovered,
and among the frenzied inhabitants of the nest a number of
Microdon pupae were noticed. The ant was later determined as a
subspecies of Camponotus maculatus. At this date, May 19, there
were no larvae of the fly present and the pupae were all fully
developed. Eighteen pupae were taken, most of them rather closely
crowded near the entrance to the nest; all around them were empty
puparia, bearing evidence that the nest had been used for several!
seasons by the fiies. There were several adult flies around the log,
some of them freshly emerged, but the ants were so aroused at the
disturbing of their domestic tranquility that they quickly drove out
any strange insect that came near. The puparia taken were allowed
to become too dry and only two adults emerged out of the lot.
In April, 1921, some observations were made on another
species of Microdon. A student at Stanford University, Mr. Her-
bert Mason, found a single larva in a nest of Camponotus maculatus
vicinus Mayr. This specimen was reared by Mr. Carl Duncan and
the specimen and notes regarding it kindly turned over to the
writer. The species proved to be Microdon piperi Knab, a beautiful
dark blue species which ranges north along the Pacific coast region.
The larva was not closely examined by the writer, but in the
notes made on the specimen the color was given as largely pale
20 Journal of Entomology and Zoslapy
bluish green, with median ridge and the margins of the body
brown. The median ridge was quite prominent in the larva. The
coarse reticulum on the body has a pattern somewhat like that
figured for M. tristis, as can be seen from the figure. The length
was 11 mm.
The puparium shortens to about 9mm. The reticulum is much
more distinct than in the larva and two prothoracic tubercles push
out (in the specimen described one of the tubercles did not push
through the body wall). The reticulation is arranged in a more or
less symmetrical design and when examined under a high magni-
fication is seen to be made up of two types of processes; those on
the dorsal ridge and along the sides just above the fringe are of a
shape which might be likened to an inverted wine glass and the
other processes are quite short and composed almost entirely of the
white stalked portion (see figs. 6 and 7). The base in both cases is
dark brown and the apical portion white. From above the body
appears to be covered with white discs arranged in a reticulated
pattern, the center of each disc with a depression and a minute
cavity which appears to penetrate almost or quite through the
body wall. These minute structures may function as pores. The
anterior margin of the ventral fringe of the body is deeply notched
in the middle as shown in figure 4. The structure of the marginal
fringe is shown enlarged in figure 8. The fly emerges from the
puparium by breaking off the cover in three rather symmetrical
pieces, illustrated in figure 2.
The specimen described was taken the last of March and soon
commenced to pupate, the puparium being fully colored by April 8.
The adult emerged just a month later.
Wheeler notes that the most typical and frequent hosts of th-
Microdon larvae are ants of the genus Formica but Wasmann has
recorded a species of Camponotus in Madagascar as a host.
pew,
2 ae a Sy Nee
Ln
Fara daeda a custananes SS
Pre OTM SS
Fig. 1. Puparium of Microdon piperi Knab; 2, anterior portion of puparium,
showing symmetrical breaking; 3, posterior respiratory tubercle;
4, marginal fringe of puparium, showing split in anterior region;
5, reticulations of two kinds, those with a short, and those with a
high base; 6, and 7, portions of the reticulations more highly mag-
nified; 8, marginal fringe, greatly magnified,
Notes on California Bombyliidae with
Descriptions of New Species
Frank R. Cole, Stanford University, Cal.
The sun-loving Bombyliidae have always been a favorite group
with the writer, as the rather abraded specimens in his earliest
collections will bear evidence. California is rich in species of these
flies and notes on a few of the interesting forms are given below.
During the past two summers the writer has spent some time
in Mill Creek Canyon in San Bernardino County. Paracosmus
morrisont O. S. is a very common form in this locality and is
usually taken along roads and paths in the bright sunlight. Aphoe-
bantus vittatus Coq., a trim, beautifully marked little species,
occurs along with the above, but is not so common and is often
harder to catch. Villa squamigera Coq. and Villa mira Coq. are
not uncommon in the Mill Creek region, the latter species more
abundant in August, when it is found out in the sandy river
washes. Villa miscella Coq. is seldom seen and is quite wary,
flying up and down sandy roads for long distances when disturbed.
In Glen Martin, in this same general region but at a higher alti-
tude, one occasionally finds Rhabdoselaphus setosa Cresson, a little
species with a very long proboscis; it is usually taken on the wing
in the middle of the day, hovering near the ground. With the first
days of autumn specimens of Villa autumnalis Cole begin to ap-
pear, frequenting the yellow flowers of Hricameria and Chry-
sothamnus, and now and then a specimen of the beautiful golden
Lordotus diversus Coq.
Villa chromolepida new species.
Female. Length 7mm. Black, clothed with bright iridescent
scales; front tibiae without bristles; wings hyaline.
Head rather large in proportion to the body; proboscis pointed
and scarcely projecting beyond the oral margin; palpi small, black.
cylindrical and black pilose. Antennae black, first joint about
twice as long as second and with black pile; third joint twice as
long as first two combined and gradually tapering toward apex
(see fig. 9), the apical bristle minute. Frons shining black, with
erect black pile and sparse golden scales which are purple in
certain lights. Face projecting (see fig. 10), shining black, with
scales like frons but denser, pile short, black, reclinate. Cheeks
shining black, bare of pile or scales. Occiput black, densely
clothed with scales like those on face and frons; next the hollowed
out back of the head a line of short, fine, yellowish pile.
Mesonotum and scutellum shining black, with golden green
scales, purple by reflection; the median portion of dorsum with
22 Journal of Entomology and Zoology
erect blackish pile, the front and margins with white pile, stiff a>!
erect just back of the head. Pile of scutellum sparse and white.
Pleura shining black, with rather long, dense white pile on the
upper mesopleura, the lower part of mesopleura and other pleura!
plates with sparse black pile, not obscuring the ground color; sti7,
blackish bristle-like pile above front coxae. The coxae and pleura
with a few seattered iridescent scales. Stem of halteres yellow, the
knob white, with a black mark on anterior margin; tuft of pile
before halteres largely yellow.
Abdomen black, with erect white pile on sides of first and on
anterior corners of second segment; beyond this the pile is very
sparse, black, reclinate and scarcely noticeable. On each side of
posterior margin of first visible tergite some scales like those on
thorax; on the other abdominal tergites and sternites there is a
dense covering of tomentum or scales, largely colored like those of
thorax and in a definite design on dorsum; in the center of each
tergite beyond the first visible one a round spot with sparse bla
scales, on each side a larger oval spot covered with black scales
which have a purplish color in some lights; these lateral spots miss-
ing on seventh segment, which is almost wholly covered with irides-
cent scales. The venter black, with a wide median portion cloth? !
with black tomentum, the sides with iridescent scales as on th?
more or less telescoped, the last two segments scarcely visible;
color of pollen and pile as in male. Apices of femora an ochre
dorsum. Femora and bases of tibiae brownish yellow, the rest of
legs black; all the spines and pile of legs black, front tibiae without
bristles, the anterior tarsi with claws almost as large as on the
other tarsi; femora with a few yellowish, iridescent scales and
some black ones; tibiae and tarsi with black scales. Wings hya-
line, iridescent; the costa and veins at base yellowish, toward
posterior margin black; fork of radius rather angular at base.
The epaulets with purplish iridescent scales.
Holotype, a female, collected in Mill Creek Canyon, Cal., July
20, 1920 (F. R. Cole), in the collection of the California Academy
of Sciences.
The type female is the only specimen known‘and is not closely
related to any species seen by the writer. In Coquillett’s table of
species it would run to mercedis. It is distinct from any described
Mexican species.
Amphicosmus vanduzeei new species.
Female. Length 6 mm. A slender species, the body largely
shining black, the legs yellow.
Upper two-thirds of frons black, including the large ocellar
tubercle, the lower third yellow; pile sparse, fine, white, the narrow
orbits silvery pollinose. Face short, projecting, the central portion
shining black, sides yellow and with silvery pollen; antennal fovae
Pomona College, Claremont, California 25
deep and connected; first antennal joint slightly longer than
second, yellow; second and third joints black, the third joint about
as long as the first two combined, narrower (see fig. 6), with a
short sub-apical style. Vertex and upper occiput rather full (see
fig. 7), black, the lower occiput and cheeks yellow, occiput largely
silvery pollinose, the pile minute and whitish.
Mesonotum and scutellum shining black, the pile on median
portion of mesonotum and on scutellum short, blackish, on margins
of mesonotum white. Humeral callosities yellow, silvery pollinose;
a silvery pollinose, white pilose spot just back of humeri. Prescu-
tellar callosities partly yellow. Pleura shining black, the upper
mesopleura, the metapleura and hypopleura silvery pollinose and
white pilose. Halteres white.
Abdomen largely shining black, rather broad posterior mar-
gins of all segments yellowish; apical half of seventh visible seg-
ment lemon yellow; yellow on first segment reaches lateral margins,
on the second to sixth segments it does not do so. Pile of abdomen
very fine, sparse, white, longer on sides of first and second. Venter
largely brownish yellow, blackish at base, lemon yellow on genitalia.
Femora, tibiae, first tarsal joint, apex of fifth and base of claws
honey yellow; third and fourth tarsal joints, apex of second and
base of fifth blackish. Coxae and trochanters colored like femora,
a black spot below on base of trochanters. Wings hyaline, all
veins yellow at base, toward apex and posterior margin blackish.
All cells on posterior margin of wing wide open (see fig. 8).
Holotype, a female, collected at Palm Springs, Cal., May 20,
1917 (E. P. Van Duzee), in collection of California Academy of
Sciences. The type a unique.
This species differs from elegans Coquillett in having the first
antennal joint yellow and in the greater extent of black on the
abdomen. Coquillett gives no structural characters to distinguish
his species. The above described species differs from cincturus
Williston, from Mexico, in the smaller size and in the color of the
antennae and legs, cincturus having entirely black legs.
Metacosmus nitidus new species.
Female. Length 5.5 mm. Head black, a small amount of yel-
low on sides of oral margin. Ocellar tubercle slightly above
middle of frons but the lower ocellus nearly in the center; upper
half of frons with white pile, the lower part with black; frons
shining black, the narrow orbits silvery pollinose. Antennae black,
rather short and thick, the second joint larger than first (see fig.
4). Upper face and lower frons near base of antennae silvery pol-
linose; face short, shining black, distinctly projecting. Occiput
thinly gray pollinose, short, sparse white pilose; on the under side,
back from mouth opening, an oval yellow spot on each side of
middle. Proboscis not projecting beyond oral margin.
24 Journal of Entomology and Zoology
Thorax shining black, the dorsum with short, sparse white
pile; scutellum shining black, with short white pile. Humeral
callosities and lower half of pleura gray pollinose. Stem and under
part of knob of halteres blackish, the most of knob white.
Abdomen shining black, finely punctate and with short, sparse
whitish pile; hind margins of visible segments one to four narrowly
yellowish white, broader on the first. Abdominal pile appears
white in certain lights but is largely dark colored. Sternite of
seventh segment projects downward in a noticeable triangle as seen
in profile. Pile around genitalia rather dense and whitish. Ven-
ter black, the hind margins of first five segments yellowish white.
Legs wholly black, the pile fine and.short. Wings hyaline, rather
broad and rounded, the veins black and strong; R2+3 curved
slightly forward at tip (see fig. 3).
Holotype, a female, collected at Huntington Lake, Fresno
County, California, 7000 feet, July 15, 1919 (E. P. Van Duzee), in
the collection of the California Academy of Sciences.
Paratypes.—Two females, taken in the type locality, July 8,
1919, by Mr. E. P. Van Duzee.
This species is evidently near M. exilis Coquillett, but differs
in the color of the legs and in the wing venation. The only other
species in the genus is mancipennis Coquillett an eastern form,
which has the face and the stems of the halteres white.
Acreotrichus maculipennis new species.
A velvety brown species with thickly spotted wings; the pro-
boscis slightly longer than the head.
Male. Length 4.25 mm. Head black, brownish pollinose, the
face and vertex with rather long and erect black pile. Occiput
rather flat; occiput and cheeks with black pile. Oral opening
large, the antennae placed on the upper edge (see fig. 2) ; first and
second anetnnal joints rather slender, the first slightly longer than
the second, the third slightly longer than the first two combined
and considerably widened near the middle, the style short and
subapical (see fig. 2); pile on upper side of all antennal joints
black. Proboscis black, projecting twice the length of the antennae
beyond the oral margin. Palpi black, very slender, with black
pile, projecting beyond oral margin about one-third as far as
proboscis.
Thorax black; mesonotum velvety black, shading to a sepia
brown on the margins; the pile of dorsum erect and yellowish,
appearing brown in certain lights. Scutellum velvety black, with
comparatively long, coarse yellowish pile. There are indications of
two median black vittae on the anterior part of the mesonotum,
separated by a fine brown line. Pleura brown pollinose, the sparse
Pomona College, Claremont, California Zs)
pile on mesa- and sterno-pleura brown. Stem of halteres yellowish,
the knob yellow above and blackish brown below.
Abdomen black, sepia brown pollinose, with rather long, erect
yellowish pile, nowhere dense enough to obscure the ground color.
Venter like the dorsum, the pile shorter and more reclinate. Sev-
enth visible segment projecting over the small eighth, the genitalia
quite small, colored like the abdomen, the upper and lower forceps
about equal in size and closing over the internal organs. Knees
reddish, the rest of legs black; coxae and femora with long black
pile. Wings whitish hyaline, densely maculated with dark gray
and with remarkable thickenings of the membrane, some of which
appear to form supernumerary cross-veins (see fig. 1). The veins
near the posterior margin of the wing are wavy.
10
Z
EXPLANATION OF PLATE
Fig. 1. Wing of Acreotrichus maculipennis n. sp.; fig. 2, head of A. macu-
lipennis; fig. 3, wing of Metacosmus nitidus n. sp.; fig. 4, head of
M. nitidus ; fig. 5, wing of Paracosmus morrisoni O. S.; fig. 6, antennae
and front of head of Amphicosmus vanduzeei n. sp.; fig. 7, head of
A. vanduzeei; fig. 8, wing of A. vanduzeei; 9, antenna of Villa
chremolepida n. sp.; fig. 10, head of V. chromolepida; fig. 11, head of
Rhabdoselaphus setosus Cresson.
26 Journal of Entomology and Zoology
Female. In general very much like the male but lighter in
coloration. Pile of cheeks and lower occiput yellowish, on the rest
of the head and on the antennae reddish brown. Eyes widely
separated, the pollen of frons more buff colored than in male, the
pile shorter. Pollen of mesonotum much lighter in color than in
male, the pile shorter and paler. Ground color of coxae and pleura
yellowish brown in some specimens, the pile yellow. Knob of
halteres scarcely darkened below. Abdomen in dried specimens
yellow, also the tibiae except apices and bases of the four front
tarsi. Pile and fine setulae of femora and tibiae yellowish.
Holotype, a male, and allotype, a female, collected on the sand
dunes near Golden Gate Park, San Francisco, Cal., September 10.
1920 (F. R. Cole), in the collection of the California Academy of
Sciences.
Paratypes.—Two specimens in the Cal. Acad. of Sci., taken in
the type locality, and five specimens in the writer’s collection, taken
with the typés.
In 1895 Coquillett described Acreotrichus americanus from a
single male specimen taken in the state of Washington. This litt'e
species has hyaline wings, the antennae are quite different and the
proboscis comparatively longer. In May, 1917, the writer too
a single male specimen of A. americanus near Hood River, Ore-
gon; it appears to be a rare species. A. atratus Coquillett, from
Mexico, has a slender third antennal joint, three times as long as
the first two combined and of nearly an equal width; the wing;
are grayish hyaline. The three other known species in the genus
are described from Australia.
Notes on the Color Changes of Frogs
Sarah Marimon
In all these experiments I chose two identical frogs, and kept
one in normal conditions while the other was being subjected to
change.
Tree frogs, Hyla regilla.
I. Junel16. Hot water (about 30° to 35° C.). Left the frogs
for one hour.
The spots of the frog faded out, giving a lighter appearance.
However the background seemed much the same as the control.
Control. Tap water (about 15°-17° C.). Spots distinct.
Grayish green frog.
II. June17. Hot water. The frogs for this experiment had
peculiar red and green markings.
The whole tone was lighter at the end of an hour and one-half.
Spots somewhat more indistinct than at first.
Control. Color unchanged.
III. June 16. Cold water. (Cooled with ice—2° C.) The
frog was somewhat darker in color; the spots stood out more dis-
tinctly than previously.
Control. Tap water (15-17° C.) Color unchanged, spots
showing distinctly—not so distinctly as those of the frog in cold
water.
IV. June17. Cold water. a. The frogs were rather light in
color. Darker spots more distinct.
b. Two frogs grayish green in color. The color became
darker, spots more distinct.
V. Junel17. 5:00 P. M. Two frogs with red streaks down
the backs.
One jar covered with green tissue paper, the other left as a
control.
June 18. 10 A. M. Lighter in tone than the control. The
red streak changed to light sandy color. Spots lighter,—greenish
along the sides.
Control. Same as the day before, apparently. Spots dark
grey, grey sides, broad reddish streak down the back.
11 A.M. The frogs reversed.
June 19. Red streak narrower, sandy colored. The whole
cast of the frog was lighter and more greenish.
28 Journal of Entomology and Zoology
“Streak dark reddish. Frog much darker than the one in green
jar.
VI. June 21. Green and cold. To see which has the greater
effect, the background or the temperature.
a. Two frogs rather light in color.
The spots are more distinct but the whole color is lighter
than the control.
b. Two frogs rather dark in color.
Slightly lighter. The dark colored frogs do not seem to change
as readily as the lighter ones.
These experiments would seem to indicate the greater effects
of the background. However there was some chance for error
here, because (1) the experiment was only over a period of two
hours, and (2) because the frogs objected to the cold water, and
when they were not watched, they would climb up out of the water
and cling to the side of the jar.
VII. June 17. I put two frogs in a jar lined with leaves.
One frog very reddish, the other grayish green.
June 21. The grayish green frog much greener, lighter in
tone.
The reddish frog much lighter in tone but still decidedly red-
dish in color.
June 23. The red frog still reddish.
The green background lightened it but did not change its color.
VIII. June17. 5 P.M. Red cover to the bottle. Placed the
frogs in the jars.
June 18. 10 A. M. Slightly darker. Control. Color un-
changed.
June 20. a. About the same shade as the other frog (i. e. the
control) only with a more reddish tinge. b. Distinctly lighter, and
more reddish in color.
June 21. Frogs had each a sandy streak down the back.
The streak brighter reddish. The whole tone of the frog
slightly darker than the control. The frogs reversed. Streak
sandy colored.
June 22. Streak brightly reddish. Whole tone of frog much
lighter. Streak sandy colored. Darker than the one in red.
VIII. (a) June 17. 5 P.M. Took two greyish frogs with
no particular color showing. Placed one in a jar covered with
yellow tissue paper. The other frog used for a control.
June 18. Much lighter than the control. Seemed to have a
yellowish tinge. Spots faded somewhat.
Reversed the frogs.
Pomona College, Claremont, California 29
June 19. Lighter, the spots more faded than when in the con-
trol. The difference between the two not so marked as on June 18.
June 20. Slightly lighter, more yellowish in tone. Results
not so distinct.
VIII. b. June 21. Two frogs with reddish streak (June 20).
The streak more yellowish, now has a distinctly yellowish tone.
Srots lighter. Whole tone more yellow than control.
Reversed the frogs.
June 22. Yellowish in tone. Red streak now very yellowish.
Spots lighter.
{X... June 17.. 5, P. M... Two frogs, dark in color,;. with-red
streaks down the back. One in blue covered jar, one control.
June 18. 10A.M. Frog much lighter than the control.
The red streak along the back now sandy colored, however,
still with the reddish tint. Control color unchanged. Noticeably
darker than the one in blue.
The frogs reversed.
June 19. Lighter, the red streak sandy colored, same width as
before. Sides light grayish green. Whole tone lighter than the
one in the control.
The frogs reversed.
June 20. Lighter in tone, more greenish tinge. The red
streak now sandy, slightly greenish also.
I observed some pigment cells under the microscopes. The
melanophores (black) were the most noticeable, although on close
observation yellow and bluish grey pigment cells could be seen.
I stimulated the piece of skin with ice; in some cases the black
cells seemed to expand and in others this could not be seen. Some
such action, however, would be necessary to cause the darkening in
color brought about by cold.
The stimulations with heat were somewhat less definite than
with cold; however twice the contraction of the melanophores, due
to a heat stimulus, was observed.
Left the two dead frogs for six hours. When I observed them
again they were both remarkably lighter than they had been when
they were killed.
I took a bit of their skin and observed it again. One portion
was much lighter and had several isolated melanophores. I cooled
this piece of skin with ice, then stimulated it with hot water. The
pigment cells seemed to expand.
Conclusions:
1. The tree frog changes its color in response to heat, cold
and changes in the color of its environment.
30 Journal of Entomology and Zoology
2. The frog does not actually change color so much as it gets
lighter or darker in response to stimuli. There seems to be, how-
ever, some actual change in color.
3. The colors, blue, green and yellow cause the frog to get
lighter in color. The results with red were so irregular as to sug-
gest that the change might be due to some other agent than the color
environment.
4. When there is a reddish color present, i. e., red streak, the
red environment intensifies this coloration. When, however, there
is no red color present the red environment does not develop it.
5. This same phenomena is true of green coloration. Thus a
red frog does not seem to be able to change into a green one, nor a
green frog into a red one.
6. The changes in coloration or intensity seem to be due to
the expansion of the pigment cells.
.
Experiments with Rana sp.
I. May 26. Light. I left the frog in the light (sunlight,
although not direct) for one hour. At the end of this time it was
remarkably lighter than the one in the dark room.
II. May 26. Dark. Frog like the one in light. I left it in
the dark room for one hour. At the end of this time it was much
darker than the one in the light.
Reversed frogs.
Left two hours. At the end of this time the two frogs were
the same color again.
May 28. Repeated the first step of the light and dark experi-
ment with the same results.
May 29. Placed one frog on a white reflecting surface but not
in the sunlight. In one hour very little change.
At the end of the hour, placed the frog in the sunlight, still
on a white, reflecting surface. Remained there for one hour. At
the end of this time it was very much lighter than the one in the
semi-darkness.
Sunlight then has more effect than diffused light, or perhaps
the difference is caused by the difference in temperature.
III. May 29. Placed one frog in a rather dark but not abso-
lutely dark place, used rather as a control than as an experiment.
Apparently it did not change color.
Left it for another hour. The supposition was that it did not
change color in the second hour, since the first hour had no effect.
However at the end of the hour it was much darker than the
one in the sunlight,
Pomona College, Claremont, California a
IV. May 27. Heat and cold. Placed a frog in water of 30°
C., left it for an hour and one-half.
At the end of this time it was decidedly lighter.
Placed a frog in water of 3° C., left it for an hour and one-half.
At the end of this time it was decidedly darker.
There was a great deal of difference in the color of the two
frogs at the end of the exveriment.
V. May 28. Frog in water 30° C. Left one hour. Much
lighter than one in cold.
Frog in water 3° C. Left one hour. Much darker.
Reversed the frogs at 2:24 o’clock.
At 2:45, the two frogs had reached the same color.
VI. June l. Frog in water of 30° C., left one hour. Much
lighter.
Frog in water 3° C., left one hour. Much darker.
VII. May 26. Acid. Placed one frog in a weak acid (HCL)
solution. Left for several hours. There seemed to be no change
in color—possibly a little lighter than the control.
Control. Placed one frog in water, otherwise its environment
was the same as the one in acid.
No change in color.
VIII. Alkali. Placed one frog in a weak alkali (NaOH).
Left it for several hours. There seemed to be no change in color—
possibly slightly darker than the control.
EXPERIMENTS WITH A LOCAL FROG
IX. Rana draytonii
May 27.
X.- Cold 3° C. Found a frog among the others identical in
color.
Left in cold for one hour. Darker at the end of this time in
comparison with the control.
XI. May 29. Light. Placed Rana draytonii in sunlight for
an hour and one-half. At the end of this time it was very much
lighter.
Control. Kept the control in semi-darkness. Did not change
color.
Conclusions:
1. These frogs change color under certain conditions of heat,
cold, light, dark, or excitement. Acids and alkalis have little if
any effect.
2. a. Heat and light cause the frog to lighten in color.
There is evidence that heat is the true agent, and light only as it is
associated with heat.
b. Cold and dark cause the frog to darken in color,
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Notes on the Lepidoptera of Southern
California. No. 1
DONALD C. MEADOWS
Two days during the second week of April 1922 were spent
collecting Lepidoptera at Corn Spring, Chuckawalla mountains,
Riverside county, California. The Chuckawallas are typical Colo-
rado desert mountains, being low and rough, and having the vege-
tation for the most part confined to sandy washes. Corn Spring
lies on the north side of the range in a canyon of the same name.
It is a small palm covered oasis having many introduced plants
as it is the home of an old prospector, who has a house and garden
at the spring. The elevation is approximately 1500 feet.
Fourteen species of butterflies were collected and three ob-
served. The nomenclature used is that of Barnes and McDun-
nough’s Check List.
1. Pieris protodice, form vernalis—Edw. Three males and
two females taken. Fairly common around spring.
2. Nathalis iole—Bdv. Five males collected. Found spar-
ingly flying over bare, windswept desert mosaic. One specimen
taken near mouth of Corn Spring canyon far from any vegetation.
3. Eurymus eurytheme, form keewaydin—Edw. Two males
and two females taken. Common near spring.
4. Danais archippus—Fabr. One specimen seen at spring.
5. Danais berenice—var. strigosa—Bates. One specimen
seen with the above flying among the palms at Corn Spring.
6. Melitaea Neumoegeni—Skin—Wright. Fourteen males
and five females of this interesting species were taken. Probably
the most common butterfly of that locality.
7. Chlosyne californica—Wright. Nine males and five fe-
males taken in a small canyon about two miles above the spring.
These butterflies seemed to be very local in their distribution, one
small canyon being the only place that they were found. Types
figured by Wright from specimens taken in Colorado Desert, South-
eastern California. The Chuckawallas are at the northern edge
of the type locality.
34 Journal of Entomology and Zoology
8. Vanessa cardui—Linn. A few specimens seen flying in
Corn Spring canyon.
9. Apodemia mormo—Feld. One female taken.
10. Apodemia virgulti—Behr. One male taken flying with
the above. Contrary to expectations these two species were not as
common as in other parts of the desert.
11. ‘Calephilis nemisis—Edw. One male and two females
taken in canyon about two miles above spring.
12. Atlides halesus—Cramer. One female taken. Only one
other seen flying around a species of mistletoe.
13. Brephidium exilis—Bdv. Few Lycaenidae were found,
Two B. exilis were taken flying over grass growing near spring.
14. Hemiargus hanno—Stoll. Two males taken near spring.
This is a Mexican butterfly and only occasionally reported from
California.
15. Hemiargus isola—Reak. A male taken in canyon above
spring.
16. Pyrgus tessellata—Scud. A common butterfly through-
out desert. Very common around Corn Spring.
17. Thanaos clitus—Edw. Another common species in vicin-
ity of spring. A very fast flyer and difficult to catch. Six speci-
mens taken.
In all sixty seven specimens were taken near the spring.
A List of Coleoptera Collected on the
Beach During the Summer of 1921 at
Laguna Beach, California
CLIFFORD. TS DODDS
Determined by Dr. E. C. Van Dyke of the University of California.
CICINDELIDAE
Cicindela trifasciata Fab. var. sigmoidea Lec.
CARABIDAE
Dyschirius marinus (Lec.)
Bemidion ephippigerum (Lec.)
Bembidion indistinctum De}.
*Bembidion cautum (Lec.)
Platynus californicus (Dej.)
HYDROPHILIDAE
Ochthebius interruptus Lec.
tCercyon fimbriatus Mann.
STAPHYLINIDAE
Bledius fenyesi Bnhr.
Cafius canescens Makl.
t{Cafius lithocharinus Lec.
tCafius lutetpennis Horn.
Thinopinus pictus Lec.
tHadrates crassus (Mann.)
Baryodma sulcicollis Mann.
HISTERIDAE
tAcritus maritimus Lec.
Saprinus scissus Lec.
Saprinus bigemmeus Lec.
tSaprinus sulcifrons Mann.
*This species is not recorded as being as far south as California
in Leng’s Catalogue of The Coleoptera.
36 Journal of Entomology and Zoology
MELYRIDAE
Endeodes basalis (Lec.)
ANTHICIDAE
Anthicus californicus Laf.
Anthicus maritimus Lec.
DERMESTIDAE
tDermestes marmoratus Say.
TENEBRIONIDAE
Eulabis obscura (Lec.)
Phaleria limbata (Horn)
-
CHRYSOMELIDAE
Diachus auratus (Fab.)
CURCULIONIDAE
Phycocoetes testaceus Lec.
+The names thus checked are recorded by Lea Myers, Coleop-
tera From The Claremont-Laguna Region. Jour. Ent. and Zool.
1918. Vol. X. No. 3. pp. 48-53.
Some Common Chinese Mollusca
ARTHUR S. CAMPBELL
During the last year I had the opportunity to collect and ex-
amine a number of the commoner littoral and freshwater shell-bear-
ing Mollusca occuring near Canton and at Chung Chow, Hongkong
territory. The shells enumerated include only a fair sample of
what might. be obtained after longer search under more favorable
conditions.
It is interesting to note the alliance of this fauna with that
of the islands of the Pacific and with that of the California coast.
A number of species occur here that are found on the opposite
shore but there is a very complex admixture of the more definitely
warm-water forms and with some species of endemic origin. The
observations of Ralph Arnold (Palae. San Pedro, Calif., Acad. Sc.
03) concerning the tertiary shells of San Pedro and Japan shows
us the affinities at once of the living shell-bearing mollusca of
these two regions and likewise adds to our observations concerning
the relationship between the whole Pacific molluscan complex.
The molluscan fauna of South China appears to be paleotropical
considered in its broadest aspect.
All shells were determined by Dr. H. A. Pilsbury of the Phila-
delphia Academy. In all there are one hundred and thirty-seven
Species represented in this collection.
(Contribution from the Zoological laboratory and Museum of
the Biological Survey of South China, of Canton Christian Col-
lege).
GASTROPODA
Bullidae
Bulla ampulla L.
Acmaeidae
Helcioniscus eucosmia Pils.
H. toreuma Rve.
Haliotidae
Haliotus diversicolor Rve.
Turbinidae
Turbo coronatus var. granulatus Gmel.
T. intercostalis Pils.
T. japonicus Rve.
Neritidae
Nerita lineata Gmel.
N. undata L.
N. crepidularia Lam.
N. albicilla L.
38 Journal of Entomology and Zoology
Solariidae
Architectonica perspectiva L.
Littorinidae
Littorina irrorata Say.
L. palliata Say.
Viviparidae
Viviparus rossgeri V. Mlldff.
V. ciliata Rve.
V. orientalis Lea.
V. chinensis Gray.
V. aeruginosus Rve.
Cerithiidae
Cerithium morus Brug.
Clava sinensis Gmel.
Melaniidae
Melania ebenina Brot.
Stombidae
Strombus pugilis var. alatus Gmel. .
S. canarium L.
S. succinctus L.
S. bittatus L.
Turritidae
Turris desbayesii Doumet.
Cassididae
Cassis japonica Rve.
C. inflata Shaw.
C. strigata Gmel.
Doliidae
Tonnia allium (Soub.) Dillon.
Pyrula dussumieri Val.
P. ficus L.
Cypraeidae
Cypraea arabica L.
C. carneola L.
C. errones L.
C. moneta L.
C. erosa L.
C. helvola L.
Muricidae
Murex torrefactus Sowb.
M. adustus Lam.
M. fulvescens Sby.
M. tribulus L.
Rapana bulbosa Sol.
Cymatium (Turrotriton) pfeifferiana Rve.
Gyrineum tuberculata Br.
Pomona College, Claremont, California
Thaisidae
Thais luteostoma Dillon.
T. lapillus L.
Nyctilochidae
Bursa rana L.
Distortrix reticulata Link.
Columbellidae
Columbella versicolor, Sby.
Buccinidae
Buccinum undatum L.
Eburna lutosa Lamb.
E. areolata Lamb.
Alectrion obsoleta Soby.
Buscyon perversa L.
B. (Sycotypus) canaliculata Say.
Trochidae
Monodonta labrio L.
Tegula rusticum Gmel.
T. nigerrima Gmel.
T. argyrostoma Gmel.
Astraea undosa Wood.
Volutidae
Mitra aurnita Desh.
Olividae
Olivella sayana Rav.
O. (Callianax) biplicata Sby.
O. scripta Lam.
Conidae
Conus suturatus Rve.
Conus carinalis Hw.
Conus sulcatus Hw.
Turritellidae
Turritella bacillum Kiener
Helicidae
Eulota similaris Fer.
Polygyra albolabris Say.
Camaena cicatricosa Mull.
Cyclophoridae
Cyclphorus elegans Mldff.
Pyramidellidae
Pyramidula alternata Say.
Naticidae
Natica (Polinices) mamilla L.
N. P. melanostoma Gmel.
N. P. didyma Bolton.
Sinum neritoideus L.
39
+0 Journal of Entomology and Zoology
Auriculidae
Melanpus luteus Guoy.
Scealidae
Epitonium lamellosa Lam.
Siphonaridae
Siphonaria japonica Don.
S. cornuta Gld.
S. sirius Pils.
PELECYPODA
Arcidae
Arca (Scapharca) campechiensis Gmel.
A. decussata Sby.
A. obtusa Rve.
A. granosa L.
A. (Brabatia) fusca Brug.
Parallelepipedum torta St. March.
Mytilidae
Mytilus smaragdinus Ch.
M. californicus Conrad.
M. edulis L.
Modiolus fortunei Dkr.
Septifer virgatus Wiegen.
Pinnidae
Pinna incurva Gmel.
Atrina tuberculosa Sby.
Pernidae
Malleus albus Lam.
Ostreidae
Ostrea lakerousi Lamb.
O. cristata Born.
Pectinidae
Pecten pyxidalus Boru.
P. circularis Sby.
P. circularis var. aequisulcatus Cpr.
P. gibbus var. irradians Lam.
Amusium pleuronectes L.
Spondylidae
Spondylus cruentata Lisch.
S. imperialis Chemi.
S. sinensis Sby.
Unionidae
Anodonta woodiana Lea.
Pomona College, Claremont, California
Veneridae
Tapes variegata Handley.
LT. tristis: Lam.
T. phillippinarum A and R.
T. phenax Pils.
Tivela stultorum Maue.
Gafarium divaricatum Gmel.
Venus (Chione) cancellata L.
V. C. thiara Dillw.
Mactridae
Mactra (Spirula) solidissima Dillw
Cardiidae
Cardium robustum Sol.
C. rugasum Sby.
C. sinensis Sby.
Chamidae
Chama rubea Rve.
Myidae
Corbula erythrodon Lamb.
Solenidae
Solen grandis Dkr.
Tellinidae
Tellina alternata Say.
Metis balaustina L.
Paphia striata Lam.
Caecella cumingi Desh.
Cyrenidae
Corbicula fuscata Lam.
C. fluminea Mull.
Ptericolidae
Ptericola pholadiformis Lamb.
_ Anomiidae
Anomia sir iplex D’Orb.
4]
XIV. Echinodermata
ASTEROIDEA
The nervous system of the starfish is about the same in all
forms which have been studied. Only minor unimportant differ-
ences can be recognized and some of these may be due to the differ-
ent conditions under which the observations were made or the
different methods employed.
Along the radial and circumoral ambulacral vessels on the
oral side is a median thickening of the surface epithelium. This
is the chief part of the nervous system, that is the superficial
radial and circumoral system. Separated from these portions by
connective substance there are in each arm on each side of the
middle line the deep radial bands while within the nerve ring
about the mouth there are two deep circumoral bands continuous
with the two in each arm.
From the superficial nervous system fibers may be traced
directly to the surface layers of the tube-feet. From the inside
nerve rings, fibers follow the ambulacral system. The superficial
system is merely a thickening of the epidermis in certain regions
while the deep system is a thickening of the surface of the ambu-
lacral system. Nerve strands from the circumoral rings, proba-
bly from the deep rings, run to the stomach and other viscera.
In addition to the parts of the nervous system just described
there is a rather diffuse network of fibers and probably cells, found
in the body-wall outside of the muscles. This last has been called
the coelomic.
Sense cells and perhaps something of a nerve plexus seem to
be present below the epidermis all over the aboral and lateral parts
of the starfish. Just what relationship all these parts of the ner-
vous system bear to each other or how they may be distinguished
from each other, has never been made entirely clear.
Almost any portion of the body seems to be sensitive to touch
and there may be other sensations without special organs for their
perception. At the tip of each arm a little tentacle or papilla
marks the end of the radial canal and the superficial nerve cord.
This little organ has a special epithelium and may be a special
organ of touch but Eimer, 1880, considers it as an organ of taste.
The eye-spot is the most marked sense organ of the starfish.
Each arm has, very near the termination of the radial nerve at the
tip of the arm, a bright red spot of pigment. A little closer exam-
ination of one of these spots shows it to be composed of a number
of distinct regions of color. In section these little areas are seen
to be little follicles lined with epithelial cells. The cells which line
the follicles are spoken of as the visual cells. These are clear at
their inner margins but pigmented farther down. Their inner
processes come into relation with the nerve strands at the bases of
a+ Journal of Entomology and Zoology
the eye-spots. Between and surrounding the visual cells are
numerous bipolar, elongate supportive cells which stain strongly
with connective tissue stains. In some cases the eye areas are
not in the form of follicles as Pfeffer, 1901, has shown in a species
of Astropecten. In those eye areas which appear as follicles a
lens has been described and figured by Pfeffer and others but I am
inclined to the interpretation of Cuenot, 1887, who believed that
no lens is present. In fact, in some sections which I have seen
there was no sign of even a membrane over the mouth of the
follicles.
In the superficial system many long supportive cells help to
make up the bulk of the nerve cord. These stain deeply with
usual stains and at their inner ends are more or less intertwined.
it)
A My
Fig. 27. A. Diagram of a Starfish cut so as to expose internal as well as
external parts of the nervous system. In the center the deep nerve
ring is shown by a dark curved line, the surface nerve ring by
a thicker line. These parts are continued into the arm cut longi-
tudinally on the right. Nerves to the tube-feet are shown. The
superficial nerve plexus and internal nerves are indicated. B.
Cross section of the radial nerve of starfish, superficial and deep
parts shown. C. Nerve cells and supportive cells from the central
nervous system. D. Section through one of the pedicellariae of sea-
urchin showing distribution of nerves, after Hamann. E. Section
through “taste knob” of sea-urchin. Hamann.
Pomona College, Claremont, California 45
In the past I have been inclined to consider these as in part at
least with conductive function, but I am sure the true nerve cells
are sometimes bipolar, possibly in some cases multipolar with
fibers running longitudinally and laterally in the nerve strands.
The true nervous elements are more delicate, their fibers or fibrils
cross each other at various angles but bear no other obvious
relations to each other.
Among the earliest works on the nervous system and sense
organs of starfish is that of Haeckel in 1859. In 1860, Wilson has
a remarkably clear and accurate paper on the nervous system of
the starfish. Another early paper was by Owsiannikow in 1871.
Teuscher in 1856, figures the nervous system but not in much detail.
Ludwig, 1878, has his figure of the nervous system in section often
copied. Hamann, 1883-5, shows more of the structure of the
nervous system and gives a good idea of the structure of the eye.
Cuenot, 1887, gives a clearer idea of eye structure but not much
more about the detail of the nervous system. Jickeli, 1888, recog-
nizes four chief parts of the nervous system of starfish: (1) The
ambulacral, (2) the sub-epidermal body plexus, (8) the deep
nerves, (4) the intestinal nervous system. Pfeffer, 1901, studies
the eyes particularly and distinguishes clearly between support-
ing cells and nerve cells. More recent papers of Pietschmann,
1906, and especially of Meyer, 1906, show details in the nervous
system. The last author distinguishes clearly between supportive
cells and nerve cells in the nervous system. He finds the suportive
cells uni- or bipolar and usually running from the ventral to the
dorsal side of the nerve bands. The nerve cells are bipolar or
multipolar with fine branches.
Romanes, 1885, found besides strong negative reactions against
injurious stimuli, positive reactions of a chemical nature which
he considered due to the sense of smell. This sense depended
somewhat on the physiological condition of the animal, chiefly
upon its degree of hunger. A starfish, kept several days without
food, immediately crawled near some presented. If a small bit of
food be withdrawn as the animal approaches, the starfifish could
be led about in any direction. By severing various parts of the
rays, Romanes found that this so-called olfactory sense was equally
distributed throughout the length of the body and by varnishing
the upper surface he found that the reactions were unaffected.
Also by placing a bit of food on the aboral surface no reaction
occurred. Preyer, 1886, found great differences in individuals
when stimulated with food.
Starfish are positively phototropic but largely lose this ten-
dency if the eye-spots are removed. Romanes found the sensi-
tiveness so great that starfish discriminated between ordinary pine
boards covering the tank and the same boards painted black.
Romanes Preyer, Jennings and others have studied the righting
46 Journal of Entomology and Zoology
reactions of starfish in considerable detail. In general the star-
fish rights. itself by twisting about two or three of its rays until
the suckers on the ventral side have a firm hold of the supporting
surface and by controlling the twisting movement the body is
turned over. In this it is necessary that all five arms do not make
the attempt at once to bring the animal into a ventral position. If
five or four arms should work at once the animal could not turn
over. There must be some codrdination between the arms as is
seen when the circum-oral nerve is cut. In this case the codpera-
tion of the arms ceases. A single arm removed from the rest can
right itself. These experiments seem to show that the central
nerve ring acts merely as a conductor of impulses. The ventral
side of the starfish seems to be positively stereotropic.
If one arm of a starfish is stimulated the animal moves away
in a direction opposite to the stimulated arm. This looks like
intelligence, but when one arm is stimulated the tube-feet on this
arm draw in and according to the parallelogram of forces, a move-
ment away from the point of stimulation will take place. When
the starfish is stimulated as a whole the spines and pedicellariae
wave about and the jaws snap time and again. A separate exter-
nal stimulus is not necessary for each opening or closing of a pedi-
cellaria. Mechanical stimuli that are strong enough always cause
them to attack. Very light mechanical shock often produces no
effect even if repeated. There are some responses to food rather
than defensive movements, a nutrient juice causes the pedicellariae
to advance and open. Pedicellariae are often opened for attack.
If another starfish brushes against it, even when one of the indi-
vidual’s own rays cross, the pedicellariae may be advanced.
If closed pedicellariae are stimulated they must first be stimu-
lated to open before they will attack. Any stimulus which causes
the pedicellariae to rise will when repeated cause them to open.
Most stimuli which cause the pedicellariae to withdraw also cause
them to close. The larger pedicellariae are usually less inclined
to hold objects for a long time. Starfish seem to hold objects for
a longer time than sea urchins.
In starfish the pedicellariae seize and hold each other as well
as other objects. If a small bit of the body of a starfish, bearing a
single spine be cut from the rest, the pedicellariae seize any small
object which touches them. If the ventral nerve is cut or the
whole ventral side of the ray cut the pedicellariae continue to act,
but the cutting of the nerve acts as a stimulus. The transmission
of impulses seems to be by the nerve-nets over the body-wall.
Jennings has shown that the elevation of the groups of pedi-
cellariae or the rosettes to attack, is dependent upon the following:
1. Previous mechanical stimuli; 2. Preliminary chemical stimuli;
3. Foregoing chemical stimuli; 4. Cutting the radial nerve leaves
the rosettes in such a state that they attack more readily than
usual. 5. The rising of the rosettes in a central region leaves them
Fig. 28.
SENSE ORGANS OF STARFISH. From Campbell.
i
SSL ON Ee Cor
Ventral and lateral views of eye-pad Pisaster capitatus, showing
general relationship to terminal tentacle. X9.
Ventral view of eye-pad of Orthaster gonolena. X9.
Ventral view of eye pad of Pisaster ochraceus. X9.
Ventral view of eye-pad of Asterina miniata. X9.
Ventral view of eye-pad of Linckia colombiae. X9.
Ventral view of eye-pad of Asteropectin erinaceus. X9.
Ocellus from Orthaster gonolena to show general form. X350.
Drawn by camera lucida.
48 Journal of Entomology and Zoology
8. Ocellus from Linckia colombiae to show general features. X350.
Camera lucida.
9. Ocellus from Asterina miniata. X350. Camera lucida. General
view, note the clear central margin of pit.
10. Tactile organ from terminal tentacle of Linckia colombiae.
General view showing papillae and details. Camera lucida.
X350.
11. Single sensory cell from Linckia colombiae. Very greatly
magnified.
12. Sensory cells from Asterias rubens showing pigment. Re-
produced from Cuenot. Osmic acid. Greatly magnified.
13. General view of eye-pad of Asteropectin erinaceus. X3850.
Camera lucida.
14. Simple ocellus in an Asterias. Supportive cells dark. Sensory
cells lighter. Reproduced from Pfeffer. Diagramatic.
15. A more complex ocellus from Asteropectin mulleri. Note the
lens, other features as above. From Pfeffer. Diagramatic.
after subsidiance in such a state that they react more readily to
stimuli in a distant part of the body than the rosettes near the new
stimulus; 6. There are differences in the characteristics of indi-
viduals.
The opening of the pedicellariae depends upon:
1. Homogeneous preparatory stimuli
(a) Sometimes there is no response to the first stimulus.
(b) Sometimes the first stimulus causes retraction and
closing while later ones cause extension and opening.
(c) Sometimes with large pedicellariae the first stimulus
causes momentary opening, the next two or three have no visible
effect, the next pronounced opening.
2. Chemical stimuli of a certain character cause the pedi-
cellariae to open later and more readily under mechanical stimuli.
3. Chemical stimuli of a certain character cause later refusal
to open under usual mechanical stimulation.
4. Holding some object causes the pedicellariae after release
to refuse to open under other stimuli.
5. After closing the pedicellariae often open and close again
spontaneously, “snapping.” The foregoing action furnishes the
condition for the succeeding one.
In many cases the tube-feet are compelled to do much feeling
about before they find the object seized by the pedicellariae. In
exploring movements two or three rays are raised from the others
and swung about in the water; the other rays creep about. The
tip of the arm as well as the other parts of the arm are employed
in these feeling motions.
The relative intensity of illumination on different parts of the
body of the starfish may and at times does determine the direction
of movement without regard to the direction of the rays of light.
The ventral portion of the surface of the starfish is protected by
Pomona College, Claremont, California 49
movements more than the tips of the arms. After it has been at
rest for a time however the eye-spots are usually so placed as to
be protected from the light. The starfish in each case (Jennings)
moves towards that part of the body that is least illuminated.
There are a number of ways in which starfish right them-
selves according to Jennings:
1. The simplest method. Two adjacent arms twist their tips
with ventral faces inwards.
2. Two arms, the ventral faces not inwards but facing in the
same direction.
3. Three adjacent rays attack and usually turn by twisting
the outward rays.
4. Four rays take hold, two to right, two to left. Fifth ray
helped up, and swings over.
5. All rays attack release later of certain rays.
6. One ray twists and rights the whole.
7. Righting without attaching tube feet of any of the rays.
Raises disc strands on tips of arms then topples over.
If a starfish begins a reaction in a certain way it usually con-
tinues in the same way even in spite of opposing conditions.
When the starfish gets started it continues in the same way. The
variability of form in starfish that are righting themselves is
very great. No species rights itself in one way alone. When cer-
tain tube-feet are prevented from acting in righting movements
the others carry on the movements. In righting if one method
does not help another is used.
HABIT FORMATION
Preyer, 1886, Jennings, 1907, have brought further information
as the results of experiments to test habit formations in starfish.
By perventing certain rays to act in the righting reactions in star-
fish Jennings showed that he could establish temporary habits
and the slower formation of more lasting habits. The many
factors which determine the righting reactions have not a constant
tendency to make starfish turn on one given pair of rays. On the
contrary, they must sometimes act in one way, sometimes in an-
other. Therefore no very fixed habits are formed under normal
conditions.
in the righting reactions the impulse tends towards the ac-
complishment of the general turning of the starfish as a whole and
given parts sacrifice their own direction or even prevent it in the
general result.
We cannot assume single specific external stimuli as the deter-
mining factors for each separate movement, yet in some way the
relation of the organism to its environment has set in operation a
uniform action of which separate movements are parts.
50 Journal of Entomology and Zoology
ECHINOIDEA
The nervous system of sea-urchins may be compared with that
of starfish more easily than with that of other forms.
The nerves corresponding to the superficial radial and circum-
oral nerves are more deeply placed than in starfish and as in star-
fish are the most obvious parts of the nervous system. An epi-
neural space or tube on the outer side of the nervous band forms
the so-called ‘‘epineural cavity” or nerve tube, as interpreted by
Phouho, ’87, and others. The radial and circum-oral sinus follows
the nervous system on the inside.
The superficial radial system follows down the inside of the
shell in the center of the ambulacral area and these five strands
join with the circum-oral ring about the mouth opening.
From the nerve ring between the junctions of the five radial
nerves are branches to the intestine which go to make up the intes-
tinal plexus.- Nerves run out laterally from the radial nerves to
the tube-feet and also to the surface, to the bases of the spines and
to the ganglia at the bases of the spines. The radial nerves end
in the terminal tentacles through holes in the shell about the anal
region. It is by way of these openings, according to Phouho, that
the radial nerves contribute to the superficial nerve plexus just
outside the test of the sea-urchin. The deep radial nervous system
is but poorly represented, so little of it is present closely applied
to the superficial radial and circum-oral that it can hardly be recog-
nized apart from it.
According to some, a pentagonal area of aboral nerves sur-
rounds the anus and communicates with the genital organs and
with the external superficial system by means of fine fibers from
the radial nerves near their termination in the terminal tentacle.
It is quite probable that the superficial system communicates with
that of the shell at the aboral end not only through the so-called
ocular openings but also through the genital openings in the shell.
The surface of the body, the spines and the tube-feet, are all
organs of the tactile sense at least.
The so-called eye-spots at the terminal tentacle in the five ocu-
lar plates contains pigment and may have some sensitiveness to
light, but it is not like the eye-spots of starfish and may indeed not
be in any sense an eye-spot.
The chief parts of the system such as the radial and circum-
oral nerve bands are composed of about the same parts as in the
starfish. In smaller and perhaps younger specimens the outer
nuclear layer is thicker in proportion. Nerve cells are bi- and
multipolar. In some cases at least multipolar cells are found well
within the fibrous area of the strand. Many of the outer cells are
probably as in other echinoideans supportive in function. The
radial bands are thicker at the oral region and become somewhat
Pomona College, Claremont, California or
smaller at the region of the terminal tentacle in the ocular plate.
This might suggest something as to the nerve tracts or bundles of
fibers and gives an indication at least that fibers may convey im-
pulses at different distances such as in the central nervous system
of vertebrates.
The deep radial and circum-oral strands of sea-urchins are
poorly shown in section. Only a few cells scattered along the
inner margin of the fibrous region give an indication of this poorly
developed system.
In the sand-dollar, Dendraster excentricus some variation in
form is suggestive of value in comparison with other forms.
The righting reactions in sea-urchins are carried out with
greater difficulty than in starfish and only the fresher or more vig-
orous individuals are capable of the reaction.
Fig. 29. NERVOUS SYSTEM OF SEA-URCHIN. A. Diagram of nervous system of
sea-urchin showing in various ways the superficial and deep nerv-
ous system by having the superficial system cut away on part of
the two radial nerves at the left. Branches to the tube-feet shown
in the central of the three ambulacral areas. Nerves to the bases
of the spines show on the right. Superficial nerve plexus show in
the center. B. Diagram of the nervous system from the aboral
pole, showing the nerve connections at the genital openings and the
ends of the radial nerves at the five ocular plates. C. Diagram of
cross section of nervous system having branches to a spine and
a tube-foot after Delage and Herouard.
Although the eye-spots of sea-urchins are not well developed
they seem to avoid light and seek darker corners and sheltered
places. One form which has no eye-spots seems to avoid the light.
A sudden shadow falling on it causes it to direct its spine to the
%, *, *
ve
<7
hy, ©
Fig. 30. EXPLANATION OF FIGURES OF SAND-DOLLAR.
1. Diagram of one fifth of Aristotle’s lantern of Dendraster show-
ing three loops of the circumoral nerve ring, and parts of three
radial nerves, the central one partly hidden at its origin by the
lantern. The nerves are in black. X9.
2. Drawing of part of the first part of an oral radial nerve. X9.
3. Drawing of the lower end of an oral radial nerve. X9.
Pomona College, Claremont, California 6 6:
4. Drawing of the upper part of an aboral radial nerve. The
eye-spot region is up in the figure. X9.
5. Camera lucida drawing of a part of an aboral nerve showing
position of cell areas. X70.
6. Drawing of a section of an oral radial nerve. X300.
7. Drawing of a section of circumoral nerve. X300.
8. Drawing of a section of aboral nerve. X300.
9. Nerve cells from central regions of a radial nerve. The ar-
rangement is as shown in the drawing, cells of various levels shown
as one layer. Some of the processes possibly relate nearby cells,
but most fibers run into the general fibrous mass. All fibres or
fibrils are small. There is some indication of tigroid substance in
some of the cells. X450.
10. Nerve cells from near a lateral branch from the radial band.
X450.
shaded area. Uexkull, 1897, was of the opinion that the sea-urchin
possessed a special set of nerve fibers concerned with photic
responses.
If a bit of the test with one or more spines be separated from
the rest of the animal, the spine or spines may be stimulated to
react much as before. In the sea-urchins there are several kinds
of motile organs. There are the jaw-like organs or pedicellariae,
borne on movable stalks; there are the tube-feet and the long mov-
able spines. All these sets of organs are controlled by nerves, and
a nerve network connects all these motile organs. One general
network of nerves is within the shell and one without, and these are
connected with the five radial nerves and the circumoral nerve ring.
Each of these motile organs has a definite number of reaction or
responses and in these each group may act independently and each
organ may react as an independent individual. Each sea-urchin
then seems according to Uexkull to be made up of a colony of
almost independent structures yet all these are connected by the
nerve network and when one carries out a reaction others may
receive a stimulus to carry out its special activity.
The independence of these systems of spines, pedicellariae and
. tube-feet, and the definite character of their reflexes has been
clearly expressed by Von Uexkull He considers the sea-urchins as
made up of a “republic of reflexes.’’ Each reflex is of the same
value and is independent of the others except for the- nerve-net
connections between the systems. This group of chiefly independ-
ent systems has nothing like a central unity controlling them as a
whole and it is only by the synchronous course of different reflexes
that a unified action is simulated. The action is not unified but
the movements are ordered. Separate reflexes are so constituted
and so combined that the simultaneous but independent course of
reflexes in response to outer stimulus produces a definite general
action similar to the condition in animals with a common center,
54 Journal of Entomology and Zoology
The pedicellariae of sea-urchins refuse to seize or hold each
other or parts of the bodies or others of the same species. Von
Uexkull believes this is due to a presence of a substance “auto-
dermin” which is in the skin. In sea-urchin pericellariae have the
power of independent reactions. Each when isolated on a piece of
shell may behave as when on the body of the animal. The stimulus
from the pedicellariae need not pass through the radial nerves for
if the nerve is cut the reactions are as before.
OPHIUROIDEA
The nervous system of serpent-stars is more complicated than
that of starfish. The radial and circum-oral nerves are shut off
from the surface of the body and inclosed in a small cavity. The
more superficial radial and circum-oral nerves are well developed
and from the radial nerves fibers run out to the spines of the legs
and the tube-feet. These last are provided with ganglia at their
bases and with nerve strands running their length. The nerves
which run to the spines also have ganglionic thickenings upon
them at the bases of the spines. From the ganglia at the bases of
the tube-feet delicate strands run out to one epineural ganglion
for each tube-foot. On the ventral side of the disc on each side of
radial nerves lateral nerves run out to near the margin of the disc
communicating with the radial nerves and also connected with the
superficial nerve plexus on the lower side of the disc. There are
then in this way two lateral nerves from each arm base, and each
one of these sends out an inter-radial nerve. Nerves from the
circum-oral ring run to the teeth, five pairs in all.
The deeper radial and circum-oral nerves are closely applied
to the more superficial nerves and appear much like parts of them,
being represented by groups of dorsally placed cells. These deeper
nerves are two for each arm. The circum-oral ring about the
mouth sends out pairs of nerves to the muscles of the arm.
A system of so-called genital nerves is found quite distinct
and independent from the other systems. More or less isolated
loops of fibers are found in each area of the disc between the arms.
Hamann, 1888, gives one of the best accounts of the nervous’
system. Delage and Herouard also figure and describe the nerv-
ous system in this group. The first author describes and figures
nerves coming out laterally from the chief radial nerve to be
applied to the skin. These may be the cutaneous nerves of Delage
and Herouard.
Hamann also shows strands from these to the tube-feet where
ganglia are located and from these ganglia are nerves with ganglia
running in to the center of the arm, and also nerves arching up
dorsally to end in small ganglia. These are very much in the posi-
tion as shown in the diagrams and figures,
Fig.
Pomona College, Claremont, California 55
The chief radial nerves, as is well known, are concentrated
into ganglion-like swellings at the intervals between the vertebrae
and here from the dorsal nerve cord strands are sent into the mus-
cles of the arms.
jl.
eS? pe <i)
ee a
NERVOUS SYSTEM OF SERPENT-STARS. A. Diagram of the nervous
system of a serpent-star, a part of the disc and the bases of three
rays shown. In the upper right end ray the superficial nerve ring .
and radial nerve are removed to show the deeper nerves. In the
others and on the dise other nerves are shown. On the disc the
superficial nerve-net is given. Out from the radial and circum-oral
nerve the chief branches to the tube feet, etc., are shown. B. Dia-
gram of the nervous system of one of the arms cut across showing
large cavity of an arm in deep shading and the lowest cavity
within radial nerve, with branches to spines and tube-feet. C.
Diagram of section of an arm after Hamann. D. Through the arm
at another level. E. Section through radial nerve. Hamann. F.
Section through sense papilla. Hamann.
56 Journal of Entomology and Zoology
The parts of the nervous system are:
1. The superficial radial nerves. The chief branches: (a)
muscular nerves, (b) cutaneous nerves to tube-feet and to skin and
to spines. On each nerve to the tube-feet a ganglion is formed at
base of each tube-foot and strands run dorsally and centrally to the
intervertebral ganglia and ventrally to the two ventral ganglia or
epineural ganglia. (c) branches communicating with the lateral
nerves of the disc on each side of each radial nerve which in turn
have altogether 10 interradial nerves near the center of the disc
and branches to the superficial nerve plexus.
2. The superficial oral nerve is pentagonal in shape and gives
off: (a) nerves to the stomach, (b) a pair of dental nerves.
3. The deep radial nerves give off nerves to the muscles of the
arms.
4. The deep oral ring gives off: (a) interradial superior
nerves, (b) interradial internal nerves.
5. The genital nerves, independent of the others. Five dif-
ferent nerves between each radial area in the disc.
There are no eyes. The skin has no cuticle except at certain
points and these are the only ones where sense cells are located.
The tube-feet and spines are sensitive to touch. The palps are
sensitive to touch as well as parts of the general surface. The
extremity of the palps have sensory functions. The terminal
tentacle, it is thought, may be olfactory in function. The oral
palps have sensitive papillae well supplied with nerve cells.
The structure of the nervous system is somewhat like star-
fish but the central cords are parallel with more evident nerve cells
and the strands seem to have a more complicated structure.
Hamann’s work is perhaps the most valuable in this group.
Delage and Herouard make chief use of this in their work. Teus-
cher 1876, Land, ’76, Smith, ’79, and Ludwig, ’80, are the other
chief contributors who have considered the nervous system.
A New Salt Marsh Mealy Bug
(Friococcus palustris n.sp.)
CLIFFORD T. Dopps, University of California
Introduction.—While making a study of the insects of the salt
marshes and brackish waters of the San Francisco Bay region, I
chanced to find in considerable numbers, a mealy-bug on the salt-
marsh cord-grass (Spartina foliosa Trin.). It occurred on the
upper surface of the leaves and generally out of reach of the
ordinary high tides. The probable reason why this mealy-bug, as
well as the scale, Chinaspis spartinae Comst., occurs almost en-
tirely on the upper surface of the leaves is because of the fact that
during transpiration, water is given off from the lower surface
of the leaf, leaving after evaporation a considerable deposit of salt.
Type host and locality—From Spartina foliosa Trin., at Al-
monte, Marin Co., California, November, 1921. Found wherever
the host is located about the shore of Richardson’s Bay, an arm of
San Francisco Bay. :
Fig. B. Leaf of host plant Spartina foliosa Trin.; w, Chionaspis spartinae
Comst., adult female; x, H#. palustris, female before secreting sac;
y, sac that has been wet by the tide; z, normal sac.
Sac.—The natural sac is composed of fluffy white waxy fila-
ments (Fig. B, z), which after they have become wet by the tide,
and this is usually the case in nature, become a light ashy gray (1),
and have a more or less feltlike texture (Fig. B, y), thus offering
great protection, especially for the overwintering young. At the
posterior end of the sac there is an obscure opening, plugged with
wax filaments, where the young escape. The average length of
eis sac is 4 mm. for the adult females and somewhat less for the
males.
Adult female.—type—(Fig. A) Body smooth; six cephalotho-
racic spines on the dorsum, the two median anterior ones being
larger than the other four, all straight, slightly expanded at the
base, tapering to a rather blunt apex; eight pairs of very small,
blunt, conical, dorso-lateral marginal spines on each side; the
58 Journal of Entomology and Zoology
posterior spine of the last pair of each marginal series slightly
larger than the others. On the ventral surface there are sparsely
scattered hairs, arranged segmentally; four spiracles located pos-
terior to the coxae of the front and middle legs. Anal lobes not
chitinized, each with three small ventral and one large terminal
setae (Figs. C, D) and two dorso-lateral spines on the inner sur-
face. These spines are slightly larger than the cephalothoracic
spines mentioned above, not expanded at the base and very blunt.
The last pair of the marginal spines are located dorso-laterally
near the basal end of the anal lobe (Fig. E). The terminal setae
of the anal lobes are about two and one half times as long as the
anal lobes themselves (Fig. C), while the eight setae of the anal
ring are less than the length of the anal lobes. Antennae medium
stout, six to eight segmented (Figs. G, H, I, J,), the normal long
Fig. A. Eriococeus palustris n. sp., adult female cleared in caustic potash.
third divided into the third and fourth, and the normal ultimate
segment divided into the seventh and eighth. Apparently this di-
vision is not closely related to the moults. Legs rather slender
(Fig. F), claws not toothed, digitules with flat apical enlargements.
Male.—Body 1 mm. long; folded wings Prolene ’/, mm. be-
yond end of abdomen.
Pomona College, Claremont, California 59
Eggs.—Average 60 to 70 eggs per female, 92 highest number
noted. Ellipsoidal, pale cadmium yellow (1) ; .5 mm. long, .25 mm.
wide.
Type and paratypes deposited in the California Academy of
Sciences, paratypes also deposited as follows: United States Na-
tional Museum, Washington, D. C.; G. F. Ferris, Stanford Uni-
versity, Palo Alto, California; E. O. Essig, University of Cali-
fornia, Berkeley, California, and in my own collection.
Comparison.—This is a very distinctive species, the small
number of spines, their form, size and distribution separating it
quite widely from the known species of this state. The only
species that I have seen which at all resembles it is Hriococcus
inermis Gr., which is found on grass at Camberley, Surrey, Eng-
land.
Life history—As a rule the females come to rest with the
cephalic end of the body uppermost on the erect leaves, where they
— es
j>—1 >
Fig. C. Anal lobe showing relative positions of dorsa-lateral spines to ven-
tral setae.
Fig. D. Venteral aspect of anal lobe.
Fig. E. Dorsal aspect of anal lobe.
Fig. F. eg.
Figs. G, H, I, J, showing variation of antennae.
Note: (1) Nomenclature of Windsor and Newton’s water colors as given
in the “Glossary of Entomology”, Smith. Brooklyn Ent. Soc., Brooklyn, N. Y
1906.
60 Journal of Entomology and Zoology
secrete the sac about themselves. The eggs are layed in the
bottom of the sac, being quite closely packed with wax filaments.
As the female deposits the eggs her body is crowded forward, the
dead remains being found in the upper end of the sac. In some
instances in the laboratory, it was noted that undersized females
would secrete a sac, deposit a small number of eggs and die. Upon
dissection, it was found that there were often mature eggs in the
oviducts or partly developed eggs in the ovaries. The first laid
eggs in the bottom of the sac hatched several days in advance of
the others and thus the nymphs escaped before the later eggs
hatched.
Under laboratory conditions the first moult took place 17 days
after hatching and the second moult a week later. After the sec-
ond moult the individuals had lost the pale cadium yellow color and
became the violet gray of the adult. The day following the second
moult they secreted sacs, although they were very much smaller
than those which first made sacs under natural conditions. Nor-
mally they probably have five moults. The overwintering indi-
viduals found in nature were first instar nymphs which had not
left the sac.
Parasites —One Hymenopterous insect was found, Pseudo-
coccobius claussent Timberlake (2) which parasitized a large per-
centage of the mealy-bugs. As many as six of these parasites
were found in an adult female. They usually kill the female after
she has made the sac and before oviposition. They overwinter in
the sac as adults, emerging in the spring through circular holes
which they make.
Resistance to drowning.—Several experiments were made to
see if this insect was specially protected from the water. It was
found that submergence for three or four days had no ill effect on
an adult and that they could float on the surface of fresh water for
three weeks without dying. These results are of little significance,
however, as Mr. Floyd Wymore, in his B. S. thesis work at the
University of California, 1922, found that Pseudococcus gahani
Green, a terrestrial mealy-bug, not only could live under water but
laid eggs and otherwise lead quite a normal life.
Acknowledgements.—I am deeply indebted to Prof. G. F. Fer-
ris of Stanford University for numerous courtesies, especially for
determining this mealy-bug as a new species and for the permis-
sion to examine his collection of Coccidae. I am also indebted to
Mr. W. C. Matthews for photographing figure B, and to Prof. E. O.
Essig for suggestions and aid.
Note: (2) Mr. A. B. Gahan, Entomological Assistant of the United States
National Museum, writes as follows: “The parasite appears to be Pseudo-
coccobius clausseni Timberlake. This species was described from a single
male specimen bred from Hrium sp. [Lichtensioides Ckle.] * * * at Riverside,
Calif. Your males differ very slightly in the extent of yellowish color on
the face but I believe there is very little reason to doubt that they represent
this species with the type of which they have been compared.”
Notes on the Life History of Dinapate
wrightii Horn. (Col.)*
By Roy E. CAMPBELL, U.S. Bureau of Entomology, Alhambra, Cal.
In May, 1916, Mr. J. O. Martin, of Pasadena, after consid-
erable tedious scouting, discovered a log of the Washington Palm
(Washingtonia filifera) in Palm Canyon, on the Northwestern
border of the Colorado Desert, which contained partially-grown
larvae of Dinapate wrightii. Mr. Martin could hear more larvae
at work in the log, and decided to mark it and wait until the follow-
ing spring for further action. In 1917 he returned to Palm Can-
yon and sawed out several pieces from the fallen log, carried them
down the canyon to his automobile (a feat which was discovered
a little later by the writer to be quite laborious) and transported
them to Pasadena."
At the suggestion of Dr. F. H. Chittenden, the writer got into
communication with Mr. Martin and received directions to locate
the remaining 10 feet of the infested log. On May 19, 1917, the
writer went to Palm Springs and duplicated Mr. Martin’s actions,
except that the scouting was unnecessary. The logs were placed
in a wire cage, in Alhambra, and closely watched. Mr. Martin’s
efforts were rewarded by the appearance of the first beetle on
August 3, and emergences continued until thirty-one had appeared
by September 17. In the writer’s cage 3 adults appeared in the
latter part of July, and 2 in August.
When the sections were sawed from the log, a few larvae were
disclosed, some practically full-grown, while others were quite
apparently immature, indicating the possibility of two broods.
Also after the emergence of the beetles in 1917, larvae could still
be heard at work within the log.
On April 15, 1918, one piece of the log which had been trans-
ported to Alhambra, was cut up and examined. Nine larvae were
found, four of which were full-grown, and the rest not over half-
grown. These larvae were sent to Dr. Chittenden. Also one dead
adult female, which had failed to make its way out of the log,
was uncovered. The emergences of adults for that year from the
remaining piece of log were as follows:
* Bull. Brooklyn Ent. Soc. Vol. XII No. 5, pp. 107-110, December, 1917.
*Since this paper was presented for publication, an article by Dr. J. A. Comstock on “‘A
Giant Palm-Boring Beetle’? appeared in the March, 1922, Bulletin of the Southern California
Academa of Sciences (Vol. XXI, Part I). Besides giving many of his observations, it reviews
the literature on this interesting beetle.
62 Journal of Entomology and Zoology
August 1, 1918—1 male.
August 2, 1918—1 male.
August 2, 1918—1 male.
August 8, 1918—1 male.
Sept. 2, 1918—1 female, elytra deformed.
Since there evidently was still another brood, or some larvae
were slower in developing, the remainder of the log was kept, and
three beetles emerged in 1919 as follows:
July 24, 1919—1 male, large fine specimen.
July 25, 1919—1 male, small specimen.
Aug. 25, 1919—1 female, average specimen.
No further attention was paid to the log until April 1, 1920,
when out of mere curiosity, it was cut up. To the writer’s great
surprise one live larva was found.
It did not appear to be quite full-grown, or at least was a little
undersized, and was soft and flabby. Although it was not ex-
pected that it could mature, a hole was bored in the end of a piece
of the log, near and parallel to the surface, the larva put in, the
hole corked up and the piece of wood placed upside down. The
larva soon began to bore into the wood.
On May 24 it was examined again. The larva had continued
boring into the wood, parallel with the bark, filling up the hole
behind it, and had turned around in the hole and was headed up-
ward. It finally worked a little to one side, and started upward
parallel with the other gallery. It was then transferred to another
piece of log, and put in a hole bored about 2 inches deep. During
the transfer, the photograph of the larva in the gallery shown in
Plate I, A, was taken.
The cork plug was removed frequently and the progress noted.
Not much eating was done after the above date, and on July 12
the writer was delighted to find that the pupa had formed. It
was creamy white, with dark eyes. By August 4 the legs, mouth-
parts and head were turning brown, and on August 8 the adult
formed. It was put back in the hole and the latter plugged up.
The beetle proceeded to the top of the gallery and ate its way up-
ward and outward. It emerged from the log on August 23, a
medium-sized female. The gallery eaten by the larva between the
time it was put in on May 24, and pupation on July 12, in which
pupation took place, is shown in Plate I, B—C, and the exit hole
eaten by the beetle at C—D. The walls of the gallery made by the
larva are much smoother than those made by the beetle as the latter
ate its way out. An exterior view of the exit hole is shown in
Plate I, F.
A resume of the above indicates the following:
May, 1916, Palm log with immature larvae discovered in Palm
Canyon by Mr. Martin.
EXPLANATION OF PLATE I
A. Mature larva of Dinapate wrightii in gallery just preparatory to
pupation. B—C. Parallel section of gallery eaten by larva between May 24
and July 12, in which pupation took place. C—D. Hole eaten by adult in order
to escape from log. E. External view of exit hole. |
64 Journal of Entomology and Zoology
May, 1917, Log removed to Alhambra, California.
July and August, 1917, 5 adults emerged from log.
April, 1918, 4 full-grown and 5 partly-grown larvae observed
in one piece of log.
August and September, 1918, 4 adults emerged from re-
mainder of log.
July and August, 1919, 3 adults emerged.
April 1, 1920, one nearly full-grown larva found in log.
July 12, 1920, larva pupated.
August 8, 1920, adult formed.
August 23, 1920, adult emerged from log.
The partly-grown larvae observed by Mr. Martin in 1916 must
have been the ones to emerge in 1917 and 1918, indicating the life
cycle to be at least 2 or 3 years. However, they may have been
more than one year old in 1916. Mr. Martin believes that the
small larvae observed in 1917 were from a brood deposited after
the log was discovered in May, 1916. However, it is apparent that
there was no deposition after the logs were taken in May, 1917,
and it seems probable that the latest deposition possible was from
beetles which emerged in the summer of 1916. If this is true,
then the life cycle of the beetle emerging in 1920 was practically
4 years. It is possible that deposition occurred prior to 1916,
which would make the life cycle 5 years or more. Beetles emerg-
ing in the other years must have been from 1 to 3 years old at the
time the log was discovered. If this is so, it would make 4 sep-
arate broods, which seems improbable. It is the writer’s opinion
that there probably were two broods, and that the life cycle of
Dinapate wrightii may vary from 3 to 5 years. It is certain that
the period can be four years. The quantity and quality of food
accessible to each individual larva no doubt had much to do with
the rate of development, but probably other factors enter in also.
If the log contained only one brood, then the variation in the length
of life would be still greater.
It is interesting to note that when Mr. H. G. Hubbard visited
Palm Canyon in February, 1897, he observed that ‘all larvae were
thoroughly dormant and very flaccid. There are no young, and
evidently all are of the same age and nearly or quite adult. I
feel sure that they are more than one year old, and probably more
than 2 years old, but no doubt they would have issued in July or
August of this year.’””
Specimens sent to Washington by Mr. Hubbard did emerge
in August. His belief that the life cycle would be at least 3 years
is demonstrated by the writer’s experience.
? Ent. News, Vol. X, No. 4, pp. 228-230, 1899.
Pomona College, Claremont, California 65
Mr. Richard T. Garnett visited Palm Canyon on May 21 and
22, 1917, and after extended search, found an infested log, from
which he took 133 adults, 28 pupae and 17 larvae. One fresh exit
hole was observed. This and other observations indicate that the
period of emergence of the beetles extends from the latter part of
May to the early part of September, and it is probable that ovipo-
sition also takes place during this period, perhaps continuing a
little later. Mr. Garnett observed two sizes of larvae in the log.*
Only one pupal record was obtained, but judging from this,
and the condition of the insects on the various dates the log was cut
into or examined, it seems that the pupal period is about one
month, and the adult may remain in the log two weeks from the
time it forms until it eats its way out. Plate I, C—D, shows that
the beetle had to bore nearly an inch from the end of the gallery
in which pupation took place to the outside of the log.
In view of the relatively large numbers of such a rare beetle
collected by Mr. Garnett, Mr. Martin and the writer, Hubbard’s
fears that the insect was about to become extinct are quite un-
founded. The two infested logs were found in the same canyon
but more than a mile apart.
*Ent. News, Vol. XXIX, pp. 41-44, Feb. 1918.
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Journal of Entomology and Zoology 67
HOLOTHUROIDEA
In sea-cucumbers the chief parts of the nervous system are
much as in other groups but the superficial and deep radial and
circum-oral systems are quite distinct from each other.
The more superficial system is composed of five strands in
an epineural cavity under the longitudinal radial muscles but well
in from the surface of the body. The oral ring circles the peris-
tome; at the base of the tentacles between its radial branches
there are strands, one for each tentacle; other branches go to the
pharynx and intestinal tract. The epineural cavity seems not
present in some forms, possibly due to contraction of the animal.
The radial nerves end at the anal end of the body but there is no
special terminal tentacle. The radial nerves give off branches
to the tube-feet and also to the skin; two nerve plexuses have been
recognized, a superficial just under the epithelium and a deeper
one in the body-wall. Both of these networks receive some
branches from the radial nerve.
The deeper nerve ring or hyponeural divides into two strands
on the inside of each superficial radial nerve according to Hero-
uard, ’87. This deeper system is chiefly motor while the super-
ficial system is sensory, a generalization which he extends to other
echinoderms. Branches from the deep system are said to supply
muscles of the body-wall and lantern region.
Among the earlier works dealing with the nervous system of
holothurians was that of Krohn, 1841, where the radial nerves
were noticed but little detail given. Semon, 1883, and especially
Hamann, show the general form and histological structure of the
nervous system. Herouard, ’87-’89, brings out some points, espe-
cially emphasizing the motor and sensory divisions of the nervous
system, as already noted. |
Gerould, 796, shows the nervous system in Caudina but little
is said about it. Clark in Synapta, 1898, shows the nervous system
in section. Red spots at the bases of the tentacles, the so-called
eyes, are figured.
Five radial nerves are recognized and smaller branches to the
tentacles. Each radial nerve is divided longitudinally into an
outer and inner band as in other forms, but unlike others has no
vessel of any kind accompanying the nerves and no spaces or
lacunae. Each tentacle nerve sends off branches to the digits so
that almost the whole surface of the tentacle becomes sensory. On
the bases of the tentacles and in the ectoderm over the body are
sense buds or tactile papillae such as described by Hamann, ’83.
Under each of these is a small ganglion. From the lower side of
the circum-oral ring, between every two tentacles, a broad nerve
Re rove
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68 Journal of Entomology and Zoology
runs to the ectoderm of the oral disc and to the muscles of the
oesophagus.
Ackerman, 1902, gives figures of the nervous system in Cucu-
maria. Retzius, 1906, by means of the silver method gives a
mosaic picture of the epidermal cells. Between these cells are
small oval fields, the sense cells between the polygonal areas or
supportive cells. These are partly between two cells, partly be-
tween several supporting cells; they are not regularly arranged.
Reimers, 1912, discusses the development of Synapta and gives
something of the nervous system. MHaanen, 1914, in Mesothuria,
is not inclined to accept Herouard’s (1890) suggestion that the
inner nerve band is chiefly a motor nerve. Very fine intestinal
nerves from the circum-oral nerve ring are found in this form as
well as the thicker nerves found by other observers. Every ten-
tacle and every foot has its own nerve, the first from the circum-
oral nerve ring, the second from the radial nerves. The foot
nerves are .029 inches broad and smaller and more circular in out-
line than the tentacle nerves. There seem to be at least some
Ras, 4
AIM Be ane,
ach
Fig. 32. NERVOUS SYSTEM oF HoLoTHUROIDEA. A. Diagram of a sea-cu-
cumber showing superficial and deep central systems, branches to
tentacles and tube-feet and the inner and outer nerve plexus. B.
Section through body-wall of Holothuria showing central band
in dark with nerve to a tube-foot. C. Nerve supply to tube-foot,
Hamann. D. Sense papilla of Synapta supplied by a nerve. Ha-
mann. E. Oral end of Synapta showing location of sense pores.
Pomona College, Claremont, California 69
motor and probably some sensory fibers in these. Sense cells and
an epithelial plexus were not clearly seen in this form. Retzius
found sense cells in the skin chiefly about the mouth opening, in
the tentacles and the tube feet. In this form the peripheral nerve
fibers were not found. Crozier, 1915, discusses the sensory reac-
tions of Holothuria surinamensis Ludwig.
The nervous system does not have to be intact for the act of
autotomy but it is more successfully carried out when it is unin-
jured.
The animals are reactive to tactile, vibratile, photic, and chem-
ical stimuli, and practically indifferent to heat in the way of a
sensation.
The parts of the body are sensitive in the following order,
beginning with the most sensitive: (1) tentacles, (2) anterior end,
(3) posterior end, (4) papillae, (5) pedicels (Podia), (6) mid-
body surface.
The tube-feet discs are positively stereotropic. This shows in
the righting reaction. The arms are photokinetic, negatively pho-
totropic; they do not respond to increase in light intensity, but re-
spond negatively to decrease in light intensity. The whole surface
is sensitive in this way. The fluorescent-skin pigment is possibly
concerned.
Dissolved substances representing those homologous to human
taste qualities for sour, bitter eee salt and alkaline, are effective
as stimuli.
CRINOIDEA
There are three distinct parts of the nervous system:
1. The superficial epidermal.
2. The deep oral system, according to the suggestions of
Delage and Herouard.
3. The deep aboral system.
The superficial oral system is much like the radial and circum-
oral system of starfish, with the nerve ring and radial nerves run-
ning down the surfaces of the ambulacral grooves in each arm with
branches to the surface and to the little elevations covered with
sense hairs.
The deep oral system according to Delage and Herouard’s
interpretation is in the connective tissue under the epidermis and
consists of a central nerve ring and strands down each arm with
branches to the pinnacles.
The deep aboral system develops later than the oral in the
young form. It is in the center of the so-called chambered organ.
There is a central mass of nervous matter in the chamber; strands
run out from this towards the arms and fork but are united again,
70 Journal of Entomology and Zoology
to form a ring or pentagon of nervous tissue. From this ring
strands run out to each arm and branch and are distributed to the
arms, running embedded in the ossicles of the arms.
Carpenter, 66, and Marshall, ’84, found that the aboral nerv-
ous system controls the movements of the animals. If the cham-
bered organ is destroyed the animal is paralyzed, but it will swim
readily or make the necessary movements just as well when the
whole ambulacral nerve ring and alimentary canal are removed.
Fig. 33. NERVOUS SYSTEM OF CRINOIDS. A’. Diagram of a section through the
body of a crinoid showing nervous system by heavier lines. B.
Diagram of a section of the nervous system of a crinoid, nerves in
black, after Marshall. C, D, and E. Diagrams of the central nerv-
ous system of Crinoids, after Marshall and Carpenter. F. Dia-
gram of the plan of the nervous system of a crinoid.
The commissural connectives between the aboral nerves co-
ordinate movements and if these are cut the arms move independ-
ently.
The position of the radial cords within the bony plates comes
about gradually from larval conditions when they are open,
trough-like grooves. These grooves gradually close in.
The cirri each have nerves from the central aboral nerve
mass. The arms, the cirri and the palps are tactile organs.
Hamann has shown nerve endings in the surface epithelium
as well as by means of little projections with fine hairs at their
ends.
Among the important contributions to the nervous system of
this group are those of Carpenter, 1865-84, Teuscher, ’76, Ludwig,
"77, Hamann, ’87, Cuenot, 91. The papers of Hamann, Carpenter,
Marshall and Haanen are among the most valuable contributions
to our knowledge of the nervous system,
Pomona College, Claremont, California 71
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Pomona College, Claremont, California 73
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VOLUME FIFTEEN NUMBER ONE
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» MARCH, 1923
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A Mopet or THE Nasat CHAMBER OF A WuitE Mouse at BirtH—
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DECEMBER, 1923
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NOTES ON THE LIFE HISTORY OF DINAPARTE WRIGHTII HORN
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Bt 61
NERVOUS SYSTEM AND SENSE ORGANS XIV CoNntT.—W. A
Hilton
oer ee eeee ee eee eeee eee eeeeeeeee ee ee
°
oeoer ee eeeeee ee sees eeereee eee eee ee see eee ee eee eee
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